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                  <text>January,

- 1 -

1970

JOB PROGRESS REPORT

State

COLORADO

Project No.

W-4l-R-19

Work Plan No.

1

Job Title:

Sheep Distribution,

Bighorn

Period Covered:
Personnel:

Bighorn Sheep and Mountain
Goat Investigations
Job No.
Populations

12
and Herd Composition

June, 1968 to May, 1969

William

H. Rutherford

and Thomas N. Woodard

ABSTRACT

Bighorn sheep distribution and sex-age classification data collected during
the 1968 segment year are tabulated.
The Pikes Peak, Trickle Mountain,
TarryaI'l Mountains, Taylor River and Lake Fork of Gunnison River bighorn
herds show excellent production and high ewe:lamb ratios; the San Juan
Wilderness Area, Cow Creek-Wetterhorn,
San Luis Peak, and Battlement Mesa
herds show medium ewe: lamb ratios: and the Sangre de Cristo and Buffalo
Peaks herds show very low ewe: lamb ratios.
Distribution maps have been
worked on, but are not completed.
During the 1968 bighorn hunting season,
131 licenses were issued and 32 rams were killed, for a success ratio of
24-4 percent.
Successful hunters averaged 6.2 hunting days each, unsuccessful hunters averaged 8.5 days, and the average for all hunters was 7.9 days,
for a projected total of 1,015 man-days of sheep hunting.
Sixty-one percent
of permittees were hunting bighorns for the first time. First-time hunters
had a success ratio of 22 percent, while those who had hunted during previous years had a ratio of 39 percent.
Seventy-nine percent of all permittees
started their hunt on the first day of season.
Comments made by hunters
are tabulated.
Measurements of bighorn sheep heads taken during previous
years showed that only lout of 17 heads smaller than 3/4 curl had 5 growth
rings; all others had 2, 3, or 4 rings.
Of 53 heads larger than 3/4 curl,
one had 2 rings, two had 3 rings, twelve had 4 rings, four had 5 rings, and
the remaining 34 ranged in growth ring count from 6 to 13.

��- 3 -

BIGHORN

SHEEP DISTRIBUTION,

POPULATIONS

AND HERD COMPOSITION

William H. Rutherford

P. S. OBJECTIVE
Determine the statewide distribution,
sition of bighorn sheep in Colorado.

population

trends, and herd compo-

SEGMENT OBJECTIVES
1.
2.
3.
4.

Determine bighorn sheep distribution.
Determine population size or trend for each herd.
Determine sex and age composition of each herd.
Prepare detailed range or distribution maps for specific herds with
written descriptions for census.
Develop standardized forms which can.
be used by management personnel during routine census and for recording
long-term census information.

METHODS AND MATERIALS
No changes have been made since previous reports were written.
See:
Woodard, Thomas N. 1969. Bighorn sheep distribution, populations, and herd
composition.
In Federal Aid Game Research Report, January, 1969, pp. 33-38.

DESCRIPTION

OF AREA

This job is statewide in scope, but limited during each segment to those
areas which can be covered.
Activities during the 1968 segment year consisted of bighorn sheep herd observations in the areas listed in Table 1.

RESULTS AND DISCUSSION
Population

Status

Bighorn sheep distribution and sex-age classification data were obtained
during the summer by on-the-ground observation techniques using binoculars and spotting scope. Those data gathered during the winter were
obtained by aerial observation from either helicopter or fixed-wing craft,
in cooperation with Regional Game Biologists as a part of routine census
operations.
Results are presented in Table 1.

�-4 -

~ab1e 1.

Date

Bighorn sheep observations ~n Colorado; summer, 1968 and winter, 1968-69.

Area

Rams - Curl Class
!z
Full
3/4
~

Number of Sheel2
Yearlings
Ewes Male Female

Lambs

Unc1assHied

Total
Sheep

Sangre de
7-24-68 Cristo Range
Area No. 9

0

0

0

0

0

0

0

0

0

0

8-6-68

Lake Fork of
Gunnison R.
Area No. 33

0

0

0

0

0

0

0

0

0

0

8-8-68

Sheep CreekTrickle Mtn.
Area No. 10

1

0

0

0

24

2

6

23

0

56

Collegiate
8-10-68 Range
Area No. 11

0

0

0

0

1

0

0

0

0

1

3

1

1

0

21

0

0

8

0

34

Vallecito Cr.
2-12-69 Pine River
Area No. 28

1

2

2

0

5

0

0

1

0

11

Sheep Mtn.2-12-69 Wolf Cr. Pass
Area No. 15

3

3

3

0

18

0

0

10

0

37

1-8-69

Cimarron Peak
Area No. 16

Pikes Peak
2-18-69 Area No. 6

49*

118

0

0

46

8

221

Tarrya11 Mtns.
2-19-69 Area No. 23

8*

15

0

0

11

0

34

Buffalo Peaks
2-27-69 Area No. 12

6*

12

0

0

2

0

20

3-3-69
3-3-69

3-3-69

Black Canyon of
Gunnison R.
Lake Fork of
Gunnison R.
Area No. 33
San Luis Peak
Area No. 22

0

0

0

0

0

0

0

0

0

0

1

3

5

0

17

0

0

13

0

39

5

1

2

0

4

0

0

2

0

14

-------------------------------------------------------------------------------------------

�-

Table 1. Bighorn
continued.

sheep observations

5 -

in Colorado;

summer, 1968 and winter,

1968-69,

Number of Sheep
Yearlings
Ewes Male Female

Lambs

UnclassHied

'rotal
Sheep

Date

Area

Rams - Curl Class
Full
3/4
~
~

3-3-69

Cow CreekWetterhorn Pk.
Area No. 21

2

3

6

0

22

0

0

9

32

74

4

1

6

5

7

0

0

5

0

28

2

4

1

0

19

0

0

9

0

35

0

1

1

0

0

0

0

0

0

2

3-5-69

3-6-69

3-9-69

Taylor River
Area No. 26
Battlement
Mesa
Area No. 24
West Elk
Mountains

3-9-69

Sheep CreekTrickle Mtn.
Area No. 10

11

2

0

0

51

0

0

21

26

III

3-10-69

Sangre de
Cristo Range
Area No. 9

2

0

6

1

7

0

0

0

5

21

Totals

35

21

33

6

341

2

6

160

71

Add 63 rams unclassified

as to head size -

*

No classification

breakdown

given by Regional

738

Game Biologist.

Bighorn sheep hunters during the 1968 season were asked to report on observations of sheep during the hunt. Table 2 presents what is believed to be
the highest unduplicated counts of sheep seen by hunters in each of the
areas.
The term "unduplicated" means that attempts have been made to
eliminate obvious duplications whether noted by the hunter or not. Thus,
Table 2 may still contain a few undetected duplications, but should be
reasonably accurate.

�- 6 -

Table 2.

Hunter observations of bighorn sheep, 1968.

Area

Male

Adult
Female

Lamb

Yearling

Total

1.

Poudre River

9

25

9

5

48

3.

Geneva Creek

7

40

20

8

75

6.

Pikes Peak

8

33

17

3

61

9.

Sangre de Cristo Range

8

26

1

4

39

10.

Sheep Creek-Trickle Mtn.

8

24

23

8

63

*11.

Collegiate Range

11

2

o

2

15

14.

Glenwood Canyon

2

3

4

2

11

15.

Sheep Mountain

3

16

9

3

31

16.

Cimarron Peak

5

38

3

6

52

17.

Empire

2

o

o

o

2

6

14

7

2

29

.,b'(19. Bowen Pass-Clark's Peak
21.

Cow Creek-Wetterhorn Peak

9

20

11

5

45

22.

San Luis Peak

16

35

15

o

66

23.

MCCurdy Mountain

8

30

11

1

50

24.

Battlement Mesa

0

12

4

o

16

28.

Vallecito Creek

1

6

2

2

11

29.

Blanco River

0

7

5

1

13

31.

Mount Evans

5

35

15

9

64

32.

Georgetown

6

5

2

1

14

114

371

158

62

705

Totals

*

Includes sheep seen in upper Taylor Park, outside of open hunting area.
** Includes sheep seen just inside Rocky Mountain National Park boundary.

�- 7 -

rotal counts cannot be obtained in most areas because of the wide and
sparse distribution of the sheep. Herd classification counts are valuable
chiefly as trend data to aid in determining the status of each individual
herd and its reproductive success.
Following are comments on ewe:lamb
ratios for each of the areas on which sufficient ewes and lambs were
classified to provide a basis for comment.

Pikes Peak, Area No.6
This is the largest and healthiest herd in Colorado.
Mid-winter ewe:lamb
ratios have been consistently high for several years, indicating excellent
survival during the critical first year.
The 1968-69 winter ratio of
100:39 and the 1968 hunter-observed
ratio of 100:51, while good, are not
exceptionally high because non-productive yearling ewes were included in
the ewe classification.
True ratios have averaged about 100:80 for the past
several years.

Sangre de Cristo Range, Area No.9
Lamb survival has been poor in this herd for several years, and this trend
has continued through the 1968-69 season. Hunters observed only one lamb
during hunting season, and mid-winter aerial observations failed to find
any lambs. A specific study by a graduate student will be undertaken during
the next project segment, in an attempt to detect the causes of poor lamb
survival in this herd.

Sheep Creek-Trickle

Mountain,

Area No. 10

This herd is the second largest in Colorado, and is apparently just as
healthy as the Pikes Peak herd. Lamb survival is excellent, as evidenced
by a mid-summer observed ewe:lamb ratio of 100:96, and a ratio of 100:41
during mid-winter surveys.

Buffalo Peaks, Area No. 12
This herd, once one of the largest and most productive in the state, has
deteriorated to an alarmingly low level. No hunting season was held here
in 1968. Mid-winter surveys showed only 12 ewes and 2 lambs, for a ewe:
lamb ratio of 100:17.

Sheep Mountain-Wolf Creek Pass-Cimarron
Area Nos. 15, 16, and 28.

Peak-Pine

River-Vallecito

Creek,

These 3 areas cover the bighorn sheep range in the San Juan Wilderness Area.
The areas are treated separately for convenience in administering hunting
permits, but will be treated as one area for the purposes of this report.

�- 8 -

It is a large area with sheep scattered thinly and no concentrations at any
time of year.
It is also the most difficult area in the state in which to
obtain reasonably accurate and comparable population trend information.
The herd appears to be healthy and slightly increasing in numbers.
Hunter
observations indicated an ewe:lamb ratio of 100:23, while mid-winter surveys
(considered more accurate) showed a ratio of 100:43.

Cow Creek-Wetterhorn

Peak, Area No. 21

This is a healthy herd, although fairly small in numbers, in a rather inaccessible area. Again, as with the San Juan Wilderness Area herd, it is
difficult to census accurately.
Hunter observations showed a ewe:lamb ratio
of 100:55, and mid-winter surveys showed a ratio of 100:41.

San Luis Peak, Area No. 22
This is a rather small herd scattered over a very large area, difficult to
census, and difficult to establish population trends. Hunter observations
indicated a ewe:lamb ratio of 100:43, but mid-winter surveys found only
4 ewes and 2 lambs, too few to be used as an indication of herd status.

Tarryall

Mountains,

Area No. 23

After a long period of low numbers following the drastic die-off during the
winter of 1952-53, this herd, at one time the largest in Colorado, appears
to be finally starting an upswing.
Lamb survival is improving, with eleven
lambs being seen both by hunters and during mid-winter surveys.
The midwinter ewe:lamb ratio was 100:73.

Battlement

Mesa, Area No. 24

This is a difficult herd to census, because of the pinon-juniper and shrub
overs tory and sandstone cliffs which make up its habitat, certainly a
drastic departure from what is considered normal bighorn habitat in Colorado.
Ewe;lamb ratios appear good, but thus far the herd has shown no indication
of growth in numbers.
Hunter observations indicated a ratio of 100:33,
while the mid-winter ratio was 100:47.

Taylor River, Area No. 26
This area has been closed to hunting for several yea~s because of low
sheep numbers, and is now showing an increase in herd size. Productivity
and lamb survival appear good, with a mid-winter ratio of 100:71.

Lake Fork of Gunnison

River, above Lake City

This area has never been opened to bighorn hunting, and has no hunting area
number designation.
It will be recommended for an open season in 1969 by

�- 9 the Southwest Region.
The bighorn herd has been building for several years
and now contains Some good rams plus a very healthy ewe-lamb herd.
The
mid-winter ewe:lamb ratio was 100:76.

Distribution

Maps and Census Forms

Preparation of detailed range or distribution maps, census descriptions,
and census forms are still incomplete and not ready for submitting with
interim progress reports.
Work was accomplished on them during this segment, with the objective being to prepare them in finished form for the
final report.

Summary of 1968 Bighorn Hunting

Season

A total of 112 hunters, or 86 percent, of the 131 successful applicants for
1968 bighorn sheep licenses returned the questionnaire form which was sent
with the license (Appendix A). Data on the number of sheep observed and
reported by these hunters have already been presented (Table 2). The following section presents other data extracted from the 112 returned questionnaires •.
Table 3 presents the license and harvest
open to hunting during the 1968 season.

statistics

for each of the areas

The average hunter spent 7.9 days hunting sheep in 1968 as compared to 7.2
days in 1967, 6.6 days in 1966, and 6.4 days in 1965. The number of days
each individual hunted ranged from 1 to 22, for a projected total of 1,015
man-days for all sheep hunters participating.
Two permittees reported that
they did not hunt.
Successful hunters averaged 6.2 hunting days each, while
unsuccessful hunters averaged 8.5. Sixty-one percent of the permittees were
hunting bighorn sheep for the first time. Thus, of the 112 respondents, 08
were first-year hunters and 44 had hunted during at least one previous year.
First-year hunters reported taking a total of 15 rams, for a success ratio
of 22 percent.
Hunters who had previously hunted bighorn sheep reported
taking a total of 17 rams, for a success ratio of 39 percent.
Of the hunters who returned questionnaires,
88, or 79 percent, started
their hunt on the first day of season.
Six, or 5 percent, started on a
week day of the first week of season, and 7, or 6 percent, started on the
second weekend of the season.
Two, or 2 percent, did not indicate what
days they hunted.
The remaining 9 hunters, or 8 percent, started hunting
after the second weekend, with one hunter hunting only on the last day of
season.
Table 4 presents information on dead bighorn
obvious duplications eliminated.

sheep seen by hunters,

with

�- 10 -

Table 3.

License and harvest statistics, 1968 bighorn sheep hunting season.

Area

Applications
Received

Licenses
Issued

Reported
Kill

Reported
Wounded

I.

Poudre River

21

6

3

0

3.

Geneva Creek

55

6

1

0

6.

Pikes Peak

53

15

8

0

9.

Sangre de Cristo Range

47

10

1

0

10.

Sheep Creek-Trickle Mtn.

48

10

6

0

II.

Collegiate Range

46

4

0

0

14.

Glenwood Canyon

11

5

0

0

15.

Sheep Mountain

9

6

1

0

16.

Cimarron Peak

6

3

2

0

17.

Empire

7

6

1

0

19.

Bowen Pass-Clark's Peak

34

8

1

0

21.

Cow Creek-Wetterhorn Peak

13

8

2

1

22.

San Luis Peak

18

6

2

0

23.

McCurdy Mountain

26

8

1

0

24.

Battlement Mesa

9

6

0

0

28.

Vallecito Creek

10

6

1

0

29.

Blanco River

6

6

0

0

31.

Mount Evans

25

6

1

0

32.

Georgetown

17

6

1

0

462

131

32

1

Totals

�- 11 Table 4.

Area

Hunter observations

Ram

of dead bighorn

Dead this Year
Lamb
Ewe

3

1

6

1

Unknown

sheep, 1968.

Ram

Dead Previous Years
Lamb
Unknown
Ewe

1
2

2
1

9
15

l~~

3

16

2~~

1*

1·1\*

17
19

1

21
22

3

23

1

2

31

Totals

i'~

"k7~

1

1

1

1

Killed in snows lides.
Killed by poacher with

2

1

1

1

1

12

8

0

3

.22 rifle.

As expected, a variety of comments were made on bighorn sheep management
and hunting in general.
Approximately 60 respondents made such additional
remarks on their returned questionnaires.
Table 5 presents information on
the nature of these remarks and the number of hunters making each one.

Ram Head Measurements
Information was collected during the 1968 project year on correlation of
horn growth and age of bighorn rams, by locating successful sheep hunters
from previous years and obtaining measurements and age by growth rings of
the heads. While most of the data were from heads larger than 3/4 curl,
enough data from heads smaller than 3/4 curl were obtained to be worthwhile.
These data are presented in Table 6.

�- 12 -

Table S. Number of hunters expressing opinions on various subjects related
to bighorn sheep management and hunting, 1968.

Number of
Connnents

Item

Curl Regulations:
Favor half-curl
Opposed

Application

9

in certain areas

to harvest of half-curl

40

rams

Restrictions:

Leave restrictions

32

as they now are

3

applicants

Favor 3-year restriction

on successful

Favor 3-year restriction

on hunters who make a kill

Favor S-year restriction

on hunters who make a kill

Favor lifetime restriction

on hunters who make a kill

Other Connnents:
Favor exemption
requirement

3

Believe sheep and goat hunters should be allowed to take a
mule deer

3

Too many domestic sheep in the high country

8

Favor a November

*

of sheep and goat hunters from blaze orange

season in:

1

Area 10
Area 24

All hunters in these categories want application
is for unsuccessful hunters.

1

restriction

to remain as

�- 13 Table 6. Number of bighorn ram heads measured, 1968; by area where killed,
curl classification, and number of growth rings.

Area

Heads smaller
than 3/4 curl;
number of
growth rings
2
3
4
5

I.

Poudre River

2.

Gore Range

3.

Geneva Creek

4.

South Platte Canyon

1

5.

Rampart Range

1

6.

Pikes Peak

9.

Sangre de Cristo Range

10.

Sheep Creek-Trickle Mtn.1

1I.

Collegiate Range

12.

Buffalo Peaks

14.

Glenwood Canyon

15.

Sheep Mountain

1

19.

Bowen Pass-Clark's Peak

1

22.

San Luis Peak

23.

McCurdy Mountain

24.

Battlement Mesa

31.

Mount Evans

Totals

2

1

Heads 3/4 curl or larger;
number of growth rings
3 4 5 6 7 8 9 10 11
1

12

13

1
1
1

1

1

1

2

1

1
4

3

1

1

1

1

2
1

1

1

1

2

1
1

1
1
1

1

1

2

1

3

3

9

1

1

1

4
7

1

1

2
1

5

2

3

1
4

1

1

2 12

1
410

1
2

2

1

3

4

1

�- 14 -

Of 17 heads in the smaller than 3/4 curl class, only one had 5 growth rings;
all others had 2, 3, or 4 rings. Of 53 heads in the 3/4 curl and larger
class, one had 2 growth rings, two had 3 rings, twelve had 4 rings, and four
had 5 rings; the remaining 34 ranged in growth ring count from 6 to 13. Of
the 17 heads of less than 3/4 curl, 4 were taken from Western Slope areas
where rams of this class were legal during the 1968 season. The mean number
of growth rings of these 4 animals was 3.75. The remaining 13 heads were
taken from Eastern Slope areas during previous years when such rams were
legal, but which were restricted to 3/4 curl or larger for the 1968 season.
The mean number of growth rings of these 13 animals was 2.92.
The pattern was reversed for heads in the 3/4 curl or larger class. Of
these 53 heads, 9 were taken in the Western Slope areas described above and
had a mean growth ring count of 6.44. The remaining 44 were taken from the
Eastern Slope areas also described above, and had a mean growth ring count
of 6.89. Obviously, the samples are too small to place any confidence in
the figures. We need to obtain measurements on more heads to determine
possible incidence of older rams with less than 3/4 curl growth, and to
procure a larger sample from Western Slope areas.

Prepared by
William H. Rutherford
Wildlife Researcher

l

�- 15 APPENDIX
DIVISION

A

of GAME, FISH AND PARKS

6060 BROAm-JAY
DENVER, COLORADO
80216
825-1192
LICENSE NOo

_

BIGHORN SHEEP QUESTIONNAIRE

1968
1.

Were you successful in killing a ram? Yes
No
If so, what are the approximate horn measurements?
Length of horn around outside curve.
Basal circumference
R.H.
L.R.

_

spread

(mayor

-------- inches
L.R.
------ inches

may not be tip to tip)

incheso

Please circle the days on which you hunted (count part days as full days)
August:
17 18 19 20 21 22 23 24 25 26 27 28 29 30 31
September:
1
2
3
4
5
6
7
8
Area #1 (Poudre) only:

November

5 6 7 8
30; December 1 234
14
15
13
11 12

3.

Did you wound a ram and not get him?

4.

Rave you hunted bighorn

Yes

Row many bighorn
year's hunt?

------------

sheep in Colorado

If yes, what year(s) ?
_
Row many bighorn sheep have you killed

5.

at hair line.

R.R.

inches

------- inches

Widest
2.

------

_

sheep, including

No

_

other than this year? Yes

in Colorado?

the one killed,

Number

10

9

No

--------

did you see during this

Full curl Rams - Number
3/4 to full curl - Numb-e-r--------------1/2 to 3/4 curl - Number
Smaller than 1/2 curl - Number
Ewes - Numbe r
Lambs - Number

--------

--~-------------

--------

6.

Hould you please indicate on the enclosed map by an "X" the approximate location
where you saw bighorn sheep during your hunt, and include the numbers, age,
and sex of animals observed.
Also, circle location where you made your killo

7.

Did you observe any dead bighorn
Dead this year;
Dead previous

Number and sex

years:

sheep, other than ones killed by hunters?

----------

Number and sex

'-------

�- 16 1968-Sheep,

Page 2

8.

A plastic bag is enclosed for the collection of fecal samples.
If you kill
a ram, would you please strip out the pellets from the lower intestine (colon),
place them in the plastic bag, and give the bag to any Conservation Officer
or leave at :lny Game, Fish and Parks Office.

9.

Remarks and suggestions - other comments concerning your sheep hunt. We are
particularly interested in comments concerning "quality" sheep hunting,
comments on the half-curl versus three-quarter-curl
regulation, and your
opinion concerning restriction of application for bighorn sheep hunting to
once every three years, and kill to once every five years, or once in a
lifetime.

�January,

- 17 -

1970

JOB PROGRESS REPORT

COLORADO

State of
Project No.

W-4l-R-19

Work Plan No.

1

Job Title:

Job No.

13

Bi~horn Sheep Habitat Studies

Period Covered:
Personnel:

Bighorn Sheep and Mountain
Goat Investigations

June, 1968 to May, 1969

William H. Rutherford,

John P. Goettl, Thomas N. Woodard

ABSTRACT

Assignment of project pesonnel to other jobs left no time to accomplish
winter range surveys under this job. Laboratory personnel worked on analyses
of bighorn stomach sample and fecal collections.
Results of stomach sample
analyses are not conclusive, as only three samples, all from Pikes Peak,
were available for study. These three stomach contents samples averaged 51
percent grasses, 11 percent forbs, and 38 percent willow.
Lungworm larvae
were detected in all fecal samples examined, but quantitative data were not
taken.

��- 19 -

BIGHORN SHEEP HABITAT STUDIES
William

H. Rutherford

P. S. OBJECTIVE
Determine
graphical

the vegetative types, forage preferences,
features of areas now occupied by bighorn

competition and toposheep in Colorado.

SEGMENT OBJECTIVE
Determine the vegetative
winter ranges.

types and vegetative

composition

on bighorn

sheep

METHODS AND MATERIALS
No changes have been made since previous reports were written.
See:
Woodard, Thomas N. 1969. Bighorn sheep habitat studies.
In Federal Aid
Game Research Report, January, 1969, pp. 39-55.

DESCRIPTION

OF AREA

This job is statewide in scope, but limited during each segment
areas which can be covered.

to those

RESULTS AND DISCUSSION
This is essentially a negative report with respect to the objectives as
stated above.
Project personnel for Segment 19 consisted only of the project
leader and a temporary student assistant, both of whom concentrated on
collecting data for Work Plan 1, Job 12 and Work Plan 2, Job 4. No opportunity to work on habitat studies was available; therefore, the segment
objective was not met.
Some laboratory work was accomplished on analysis of stomach sample and
fecal collections.
Although these analyses are not covered in the segment
objective for this job, it appears that this is the most logical place to
report this activity.
Table 1, following, presents the analyses of three
bighorn stomach contents samples collected in 1967 and examined during this
project segment.

�- 20 Table 1. Percentage composition by point contact
sheep stomach contents samples, Colorado, 1967.

technique,

of three bighorn

Sample Number
Item

1

Date collected

2

3

9-67

9-67

9-67

Pikes Peak

Pikes Peak

Pikes Peak

Vegetative type of collection
location

alpine

subalpine

subalpine

Sex of sheep

male

male

male

Age of sheep

mature

mature

mature

Locality

where

Plant material

collected

encountered,

percentage:

Unidentified

grasses

29

58

56

Unidentified

forbs

5

4

10

Compositae

3

2

Salix

63

20

Agropyron

spp.

6

Bromus anomalus
Smilacina

spp ,

30

4
10

Fecal samples submitted by hunters during the 1968 season were examined by
laboratory personnel to determine whether lungworm larvae were present.
These samples were collected from Pikes Peak, Area No.6;
Georgetown, Area
No. 17; Bowen Pass, Area No. 19; Cow Creek, Area No. 21; and Empire, Area
No. 32. In addition, laboratory personnel made field collections of pellet
groups in the Trickle Mountain and Pikes Peak areas for the purpose of
obtaining lungworm larvae for laboratory research.
All collections showed
the presence of lungworm, but quantitative data were not collected, because
the number of larvae per gram weight of fecal material can vary so widely
that such data are not useful as an index of the level of infestation unless
very large and systematic collections are made.

Prepared

by _. ·.::.t~7;1.,....:. ~..:./..:.:Iz';"~_~_':-_&gt;1._lV_-:,-, _._.' +---";"--F~~
William H. Ruther
Wildlife Researcher

�January,

- 21 -

JOB PROGRESS

Work

REPORT

COLORADO

State of
No.

W-4l-R-19

Plan No.

2

Project

1970

Job Title:

Mountain

Period Covered:
Personnel:

June,

William

Bighorn Sheep and Mountain
Goat Investigations
Job No.

Goat Distribution,
1968 to May,

H. Rutherford

Populations

and Herd Composition

1969

and Thomas

N. Woodard

ABSTRACT

Mountain goat distribution and sex-age classification data collected during
the 1968 segment year are tabulated.
The two areas which have well-established goat herds, the Collegiate Range and Mount Evans, show very high
nanny:kid ratios and medium high kid:yearling ratios.
The trend in these
herds is definitely up.
Goats have recently been discovered in the Gore
Range.
Distribution maps have been worked on, but are not completed.
During the 1968 mountain goat hunting season, 19 licenses were issued and
15 goats were killed, for a success ratio of 79 percent.
Successful hunters averaged 2.4 hunting days each, unsuccessful hunters
averaged 7.5 days, and the average for all hunters was 3.5 days, for a
projected total of 67 man-days of goat hunting.
All but one of the permittees were hunting goats for the first time, and all permittees started
their hunt on the first day of season.
The harvest consisted of 6 male
and 3 female goats in the Collegiate Range and 6 female goats on Mount Evans.
The difficulty experienced by hunters in selecting male goats, particularly
on Mount Evans, means that the number of permits may have to be lowered.
Also, it appears that the need for goats as transplant stock precludes the
possibility of increasing the number of hunting permits in the near future.

��- 23 -

MOUNTAIN

GOAT DISTRIBUTION,
William

POPUIATIONS,

AND HERD COMPOSITION

H. Rutherford

p. S. OBJECTIVE
Determine the distribution,
goats in Colorado.

populations,

SEGMENT

and herd composition

of mountain

OBJECTIVES

1.
2.
3

Determine mountain goat distribution.
Determine population size or trend for each herd.
Determine sex and age composition of each herd.
4. Prepare detailed range or distribution maps for specific herds with
written descriptions
for census procedure.
Develop standardized form~
for use by management personnel during routine census and for recording
long-term census information.
c

METHODS

AND MATERIALS

No changes have been made since previous reports were written.
See:
Woodard, Thomas N. 1969. Mountain goat distribution,
populations, and
herd composition.
In Federal Aid Game Research Report, January, 1969,
pp. 57-59.

DESCRIPTION

OF AREA

This job is statewide in scope, but limited to those areas in which mountain
goat populations currently exist; i.~.,the Collegiate Range, Mount Evans,
The Gore Range, and the San Juan Mountains.

RESULTS AND DISCUSSION
Population

Status

Mountain goat distribution and sex-age classification
data were obtained
during the summer by on-the-ground observation techniques using binoculars
and spotting scope.
Those data gathered during the winter were obtained by
aerial observation from helicopter, in cooperation with Regional Game Biologists as a part of routine census operations.
Results are presented in
Table 1.

�- 24 Table 1.
1968-69.

Mountain

goat observations

Date

Area

7-17-68

Mt. Shavano

7-18-68

Male

5

in Colorado;

summer,

1968 and winter

Number of Goats
Adults
Female
Unc1ass.ified Yearlings

Kids

Total

I;'

22

2

11

-26

66

Sheep Mtn.

11

5

5

10

31

7-30-68

Mt. Evans

4

7

6

5

22

8-22-68

Mt. Shavauo=

18

10

28

8-22-68

Mt. Princeton*

5

8-22-68

Sheep Mtn. ~\-

22

2-12-69

Needles Mtns. ~\- 1

* Helicopter

2

5

1

4

26

2

6

counts.

Total counts cannot be obtained because of the generally rugged terrain
through which the goats are dispersed.
Thus, the data in Table 1 are valuable
primarily for determining the status of each individual herd, its sex-age
composition, and its reproductive success.
In all herds, nanny:kid ratios
are extremely high (close to 100:100), but kid:yearling ratios are only about
100:50, indicating that loss of kids through mortality before reaching the
yearling class is common.
Nevertheless, annual herd increment continues
to stay at a high level, the trend in goat numbers is uniformly up, and all
herds exhibit a healthy status.
The two areas into which immigration of goats has taken place, the Gore
Range and the Needles Mountains, deserve special mention.
A minimum of two
adult male goats are known to have been present in the Gore Range for at
least two years.
This area has been considered for a goat transplant, and
with the appearance of the immigrant animals, it was decided to go ahead
with transplant plans. Accordingly, a small shipment of 5 goats (1 adult
male, 1 yearling male, 2 adult females and 1 yearling female) was obtained
from South Dakota in July 1968 and released on Black Creek in the Gore Range.
These animals were still accounted for in October, 1968, but were not seen
during the winter of 1968-69.
In February, 1969, a helicopter flight was made by the Project Leader and
the Southwest Region Game Biologist to eva1uate.winter range conditions in
the West Needles and Needles Mountains north of Durango, as a prerequisite
to planning for goat transplants in this area. The six goats listed for the
Needles Mountains (Table 1) were observed on this trip; these were located
in Chicago Basin at the head of Needle Creek.
Prior to this, go~ts were not

�- 25 known to be present in the Needles Mountains.
These animals have almost
certainly come from the 1964 transplant of 9 goats released on Cottonwood
Creek southwest of Lake City which had since been given up as a failure,
since no trace of them could be found in the vicinity of the release site.

Distribution

Maps and Census Forms

Preparation of detailed range or distribution maps, census descriptions,
and census forms are still incomplete and not ready for submitting with
interim progress reports. Work was accomplished on them during this segment, with the objective being to prepare them in finished form for the
final report.

Summary of 1968 Mountain

Goat Hunting

Season

Of the total of 20 successful mountain goat license applicants, one was
determined to be a fraudulent purchase by a non-resident.
This hunter was
stopped before he began his hunt; therefore, the total stands at 19 instead
of 20. Of this total, 18 hunters returned questionnaires.
The average hunter spent 3.5 days hunting goats in 1968. Days hunted ranged
from 1 to 11, for a projected total of 67 man-days for all goat hunters.
Successful hunters averaged 2.4 hunting days, while unsuccessful hunters
averaged 7.5 days. All but one of the permittees were hunting goats for
the first time. In the Collegiate Range, 6 adult male and 3 adult female
goats were taken; on Mount Evans, 6 adult female goats were taken. All
hunters started their hunt on the first day of season.
Only one hunter felt that the present restrictions on applications for goat
licenses should be continued.
Two hunters felt that successful applicants
should be restricted to three years on re-applying.
Four felt that hunters
making a kill should be required to wait 5 years before re-applying, and two
felt that a once-in-a-lifetime
kill should be enough, with applications by
those who were successful in the drawing but unsuccessful in the hunt to be
limited to every 3 years.
The difficulty experienced by hunters in identifying and choosing male goats
on Mount Evans has some definite management implications.
This particular
herd, unlike the Collegiate Range herd, does not use the alpine habitat, but
instead, frequents old forest fire areas below timberline.
The cover is
heavy enough that positive identification of the sex of adult goats is difficult. It appears that reduction of the number of permits allowed will be
necessary, to avoid excess harvest of productive females.
With the need for surplus goats to furnish s tock for an intensified transplanting program, it appears that the number of hunting permits should not
be increased in any of the goat hunting areas, at least for the next few years.

v{ /i ,'.

r

Prepared

by

! /"--.r&gt;:.. .'

~/

j/

u/id!b:'i{ / ~ !fkitf;~;L(/(

William H. Rutherford
Wildlife Researcher

/
"

��January,

- 27 -

JOB PROGRESS

COLORADO

State of
No.

W-4l-R-19

Plan No.

2

Project
Work

Job Title:
Period

REPORT

Personnel:

Job No.
Mountain

Covered:

June,

William

Bighorn Sheep and Mountain
Goat Investigations

Goat Habitat

1968 to May,

H. Rutherford,

5

Studies

1969
John P. Goettl,

Thomas N. Woodard

ABSTRACT

Assignment of project personnel to other jobs left no time to accomplish
vegetation surveys on mountain goat range.
Aerial reconnaissance
during
late winter was done in the San Juan Mountains for the purpose of observing
winter range conditions in those areas which have been proposed as goat
transplant sites.
Only the Needles Mountains and the Wetterhorn Peak area
appear to have enough snowfree winter range to warrant further consideration
for goat releases.
Laboratory personnel worked on analyses of mountain
goat stomach sample and fecal collections.
Collections made in the alpine
area of the Collegiate Range averaged 80.5 percent grasses and grass-like
species, 17.2 percent forbs, and 2.3 percent woody species.
Analyses of
collections from sub-alpine range are not conclusive, as only one each from
Sheep Mountain and Mount Evans were collected.
These data are presented in
Table 1.
Lungworm larvae were detected in fecal samples collected from
two locations in the Collegiate Range, but quantitative data were not taken.

1970

��- 29 -

MOUNTAIN

GOAT HABITAT

William

STUDIES

H. Rutherford

p. S. OBJECTIVE
Determine the vegetative types, forage preferences, competition, and topographical features of present mountain goat-occupied areas in Colorado;
determine the vegetative types and topographical features of areas, particularly the Gore Range, which have been proposed as transplant sites for
mountain goats.

SEGMENT OBJECTIVE
Determine
ranges.

the vegetative

types and vegetative

composition

on mountain

goat

METHODS AND MATERIALS
No changes have been made since previous reports were written.
See:
Woodard, Thomas N. 1969. Mountain goat habitat studies.
In Federal Aid
Game Research Report, January, 1969. pp. 61-64.

DESCRIPTION

OF AREA

This job is limited in scope to those areas in which mountain goats presently occur (Collegiate Range and Mount Evans) and to those areas which have
been proposed as mountain goat transplant sites (Gore Range, Needles Mountains, Pagosa Peak, Rawah Peak, Mount Sneffe1s, Wetterhorn Peak, Sangre de
Cristo Range, Brown's Canyon, Spike Buck Creek, Graham Peak and Blanco
Basin) •

RESULTS AND DISCUSSION
Habitat Surveys
This is essentially a negative report with respect to the objective as
stated above.
Project personnel for Segment 19 consisted only of the project leader and a temporary student assistant, both of whom concentrated on
collecting data for Work Plan 1, Job 12, and Work Plan 2, Job 4. No
opportunity to work on vegetation surveys was available; therefore, the segment objective was not met.
Most of the areas currently on the State Game Manager's list of proposed
mountain goat transplant sites were observed during the winter of 1968-69.
These observations were conducted with fixed-wing aircraft during March,
1969, for the express purpose of determining whether winter range exists in

�- 30 -

sufficient quantity during the period of heaviest snow cover to warrant
further consideration as transplant sites. Comments on these areas follow:

Gore Range
Not surveyed during the winter of 1968-69, except by Northwest Region personnel who attempted unsuccessfully to locate the goats known to inhabit
this area. No data on extent of winter range have been recorded.

West Needles and Needles Mountains
This area was surveyed by helicopter in February 1969, at which time 6
goats were found in Needle Creek, and again by fixed-wing aircraft in
March 1969. All vegetation, both alpine and sub-alpine, in the West Needles
Mountains was deeply snow-covered, and no possibilities for winter range
could be found.
In the Needles Mountains the stream drainage pattern is to
the west, providing an abundance of south-facing slopes above timberline
which remain relatively snow free during winter.
This area appears to offer
excellent possibilities as mountain goat winter range.

Pagosa Peak
The only timber-free open area is confined to the peak itself, and this was
heavily snowed in. No snow free areas could be found. Even down in the
oak brush several thousand feet lower, the snow was deep and continuous.
On this basis, this is not a good transplant site.

Wetterhorn

Peak

This area covers a large expanse of high rugged mountains.
The ridge tops
seem to be well oriented to wind direction and exposure, as the area contained a number of snow free ridges, most of them with good vegetation.
This site warrants further survey work during the summer, as it appears to
offer adequate winter range for a goat transplant.

Mount Sneffels
Like Wetterhorn Peak, this is a very extensive area of high mountains, but
the ridge tops must be oriented in generally unfavorable directions since
very few of them were wind-blown free of snow. Those which were consisted
only of bare rock. Because the area is so large, goats might possibly
find a place to winter, but it would be very marginal and could not support
a herd of any great sizeo
This is not a good transplant site.

Graham Peak
This area was heavily snowed in, with much of it extensively timber covered.
Very little open area existed on ridge tops, and no vegetation showed. This
is not a good transplant site.

�- 31 Blanco Basin
Timber and vertical cliffs make up most of the area, which was heavily
snow covered.
No winter range potential could be detected.
This is not
a good transplant site.

Analysis

of Collections

Some laboratory work was accomplished on analysis of stomach sample and
fecal collections.
Although these analyses are not covered in the segment
objective for this job, it appears that this is the most logical place to
report this activity.
Table 1, following, presents the analyses of 12
mountain goat stomach contents samples collected in 1965, 1966 and 1967,
and examined during this project segment.
Pellet groups were collected on Mt. Shavano and Sheep Mountain in the
Collegiate Range during August and September, 1968, and examined by laboratory personnel to determine whether lungworm larvae were present.
All
collections showed the presence of lungworm larvae, which appeared to be
Protostrongylus stilesi. Quantitative data were not collected, because the
number of larvae per gram weight of fecal material can vary so widely that
such data are not useful as an index of the level of infestation unless
very large and systematic collections are made.

Prepared

�Table 1. Percentage composition,
1965, 1966 and 1967.

Item

Date Collected
Locality where
Collected
Vegetative Type
of Collection
Location
Sex of goat
Age of goat
Plant Material
Encountered
Percentage:
Unidentified
Grasses:
Poa spp.
Carex spp.
ASiroE:y:ronspp.
Kobresia spp.
Trisetum
sEicatum
Unidentified
forbs:
Arenaria
fendleri
Castilleja spp.
Compositae
Potentilla spp ,
Geum turbinatum
Trifolium spp.
Achillea
lanu10sa
Betula
Si1andulosa
Salix spp.
Vaccinium spp.

1

by point contact technique,

2

3

4

5

of 12 mountain

goat stomach content samples,

Sa!!!EleNumber
6
7

8

9

10

Colorado,

11

12

9-65
Mt.
Princeton

9-66
Mt.
Princeton

9-66
Mt.
Shavano

9-66
Mt.
Shavano

9-67
Sheep
Mtn.

9-67
Mt.
Evans

9-67
Mt.
Shavano

9-67
Mt.
Princeton

9-67
9-67
9-67
Mt.
Mt.
Mt.
Shavano Princeton Shavano

9-67
Mt.
Shavano

Alpine
Male
Mature

Alpine
Male
Mature

Alpine
Female
Kid

Alpine
Male
Mature

SubAlpine
Female
Mature

SubAlpine
Unknown
Mature

Alpine
Male
Mature

Alpine
Male
Mature

Alpine
Male
Mature

Alpine
Male
Mature

Alpine
Female
Mature

Alpine
Female
Mature

58
9
4
2
1

73
4
5
3

84

61
2
3
5

82

71
2
2

85
2

71

-

84

-

-

-

-

84

-

-

-

73
2
2
8

11

6

13

16

13

12

12

10

3

-

2

---1

3
1

1
4

-

2

-

-

76
3
2
7

9

14

8

12

--

3

3
2

--

---

-

-

6

1

1

7

-

2

-

-

4
2

-

1

-

-

2

-

3
-

6

1
1

-

--

4

-

3

-2

-

2

-

4

-

-

1
2

-

9

5

w

N

�- 33 -

January,

JOB FINAL REPORT

State of

COLORADO
------~~~~~-------

Project No.

W-40-R-lO

Antelope

Work Plan No.

1

Job No.

Job Title:

10

Review of Literature

Period Covered:
Personnel:

Investigations

April 1, 1963 to April 30, 1969

George D. Bear and Edgar J. Prenzlow

ABSTRACT·

An annotated bibliography (Prenzlow, E. J. 1965. A Literature Review
on Pronghorn Behavior.
Colo. Special Report No. 3, 28 pp.) has been
published and other references are being used in publications for jobs
within this project.

1970

�- 34 -

REVIEW OF LITERATURE
George D. Bear

Po So OBJECTIVE
To prepare an annotated bibliography to census methods
animals and review literature pertaining to population
methods.

for big game
determination

METHODS AND MATERIALS
Articles pertaining to population determination, census methods, animal
behavior, antelope food habits, and animal physiology were located and
abstracted.
The library at Colorado State University was the primary
source of material.

RESULTS AND DISCUSSION
A list of references was presented in the report for Segment 5 of this
project (January, 1965; pp 247-277).
Edgar J. Prenzlow published an
annotated bibliography on literature pertaining to antelope behavior
(1965.
A Literature Review on Pronghorn Behavior.
Colo. Dept. Game,
Fish, and Parks; Special Report No.3;
28 PP.). Other references are
being used in publications for jobs within this project.
This review
was also intended to support job planning, however, the review was
initiated at the same time as the field work on the jobs. Also, in view
of the fact that the Bureau of Land Management recently released a publication on antelope literature (Yoakum, Jo
1967. Literature of the
American Pronghorn Antelope.
U.S.D.lo; B.L.M; Reno, Nev. 82 PP.) it is
recommended this job be terminated at its present status.

Prepared

y

.'

bY~/
~
8·'cVL...-/
George' D. Bear
Wildlife Researcher

�- 35 -

January,

1970

JOB FINAL REPORT

State of
Project
Work

COLORADO
No.

W-40-R-10

Antelope

Plan No.

1

Job No.

Job Title:

Herd Structure

Period

April

Covered:

Personnel:

and Factors

Investigations

Affecting

11
Herd

Structure

1, 1963 to April 30, 1969

Georg,e Bear, Gary Myers,

Edgar Prenzlow

and Richard

Denney

ABSTRACT

Study areas were established in northwestern and northcentral Colorado.
The antelope herd on the northwestern Colorado (Moffat County) study area
declined rapidly under heavy harvest of 42-48 percent of the herd and increased with light harvests, approximately 20 percent.
It appears that
a harvest level somewhere between 20 and 40 percent would have maintained
a stable population.
Buck:doe ratios increased under the restricted harvest of the buck segment of the herd.
The mean age of the does in the
harvest and fawn:doe ratios showed a trend similar to the population
curve.
Very limited data were obtained from the northeastern Colorado
(Larimer County) study area due to closure of the land to hunting.
Hunters
tended to report fawns as adult antelope on the hunter report cards.
There
was a very high wounding loss during years of high antelope densitie~ this
loss decreased as the population densities declined.
Numbers of antelope
observed by the hunters and the time required for a hunter to bag an
antelope were generally related to antelope densities.
Mortality due to
predators and diseases appeared to be very light among the older fawns
and adult antelope.
There was a 12 percent return faom 167 antelope
fawns tagged on the study areas.
Due to the lack of known-age animals,
ages for antelope in this study were determined by the eruption and wear
technique.
Therefore, aging techniques tested in this study were subject
to the same errors.
Eye lens weights, horn length of bucks, head length,
head width, ramus length, and. diastema length increased rapidly in younger
animals.
Growth tended to level out from two years and older.

�- 36 -

RECOMMENDATIONS

This study did not produce adequate information concerning the effect
of hunting on population trends and herd structure.
A more intensive
study on a large herd would be useful in managing antelope herds in
the state.
Some information gathered under this job was presented in the following
report:
Bear, G. D.
1968. Hunter harvest and population trend of a
small herd of antelope located in Moffat County, Colorado.
Proc.
Antelope States Workshop 3:85-91.

�- 37

-

HERD STRUCTURE AND FACTORS AFFECTING
HERD STRUCTURE
George Bear

p. S. OBJECTIVE

Obtain information
herds.

on structure and mortality

factors affecting

antelope

METHODS AND MATERIALS

Mortality

Hunter Harvest
Check stations were established in the vicinity of each study area to
obtain information from animals killed during the hunting season. Age,
sex, and location of kill was recorded for each animal checked.
Age was
determined by the tooth eruption and wear technique.
Hunting information
(wounded and abandoned antelope observed, number of hours hunted, and
number of antelope observed) was recorded for each hunter checked.
Check stations and hunter report cards were used to determine the number
of antelope harvested on the study areas by hunters.
Wounding loss was
determined by randomly surveying the areas on foot, by vehicle, and fixedwing aircraft during the few days immediately following the hunting season.

Natural Mortality
Information pertaining to natural mortality was obtained whenever the
opportunity permitted.
Eagle nests on or near the study areas were periodically checked for antelope remains.
Dead antelope encountered on the
study area were examined to determine the cause of death. Animals found
on the "wounding loss" surveys and others collected for the food habits
and physiological studies were examined for parasites and diseased tissues.

Aging Techniques

Fawn Tagging
It is essential to have data from known-age animals if the aging techniques
are to be properly evaluated, thus fawns on the study areas were marked.
Due to the precociousness of young antelope, the fawns must be caught within
a few days after birth. Most antelope fawns in Colorado are born from May
25th to June 15th, hence, tagging operations were confined to this period.

�- 38 -

Fawns were located by watching a doe until she went to nurse her fawn;
then when the fawn hid itself, the worker used a long handled fish net
to capture the young animal. Metal tags were placed in the ears of the
animal, one in each ear. The sex of the antelope was noted prior to
release.
Tags were approximately one and one-half inches in diameter
and had an identification number and a notation (requesting the finder
to return the tag to the Department of Game, Fish and Parks at Fort
Collins) inscribed on it.

Eye Lens Weights
Antelope eyes collected at the check stations were tagged and preserved
in buffered formalin until the lenses could be removed, dried, weighed,
and compared with the age of the antelope.
Kolenosky and Miller's (1962)
technique for drying antelope eye lenses was followed.
A small sample of eye lenses were ashed and compared to the age of the
animal.
Small crucibles were weighed to the nearest one-ten thousandth
of a gram. The eye lenses were placed in the crucibles and heated in a
0
furnace at 300 C for four to six hours until the lens has started cooking
and assumed a black,foam-like consistency.
Oven heat was then increased
to 6000 C. When the lenses were cooked to a fine granular ash, crucibles
cont a Ln Lng vt.he lenses were removed from the oven and placed in a desiccator,
and allowed to cool for approximately one hour. The crucibles and lens
ashes were weighed to the nearest one-ten thousandth of a gram. Crucible
weight was subtracted from the total to determine the weight of the ashes.
The ashing technique was compared with the drying technique to determine
which was most reliable for aging antelope.

Mandible

and Head Measurements

Measurements were made on the lower jaws collected at the check stations.
The maximum length of the ramus was measured to the nearest millimeter,
not including any part of a tooth that extended forward from the anterior
end of the bone. Length of the mandibular diastema was measured to the
nearest one-tenth of a millimeter with a pair of calipers; measuring from
the posterior margin of the alveolus of the lower incisor to the anterior
margin of the first cheek tooth.
Head length and width were measured and recorded for the antelope brought
through the check stations.
Head length was measured from the occiput to
the incisors with calipers.
Measurements were recorded to the nea re st onehalf centimeter.
Head width was also measured with a caliper to the nearest
one-half centimeter, from a point posterior to the orbits and anterior to
the ears.

Horn Length
Horn length measurements were recorded for all buck antelope brought through
the check stations.
This measurement was made to the nearest one-half inch;
measured along the center of the outside surface following the curve of the

�- 39 -

of the horn from the base to the tipo Also, hunters who killed a buck
were requested to record the length of the buck's horns on a Hunter
Report Card. These data were compared with check station data to evaluate the possibility of using horn length, reported by hunters, as an
index to the age structure of the harvest.

DESCRIPTION

OF AREA

Study areas were established in Larimer and Moffat counties where major
antelope concentrations exist in two different vegetative types. The
Larimer County study area is located in the rolling prairie region
approximately 20 miles north of Fort Collins, Colorado, and is characterized by vegetation typical of shortgrass areas; blue grama (Boute1oua
gracilis), buffalo grass (Buch1oe dacty1oides), western wheatgrass
(Agropyron smithii), and others. This area covers approximately 88
square miles.
The Moffat County study area is located in the rolling
sandhi11s immediately south of Maybell, Colorado, and is characterized
by a moderately dense overs tory consisting primarily of bitterbrush
(Purshia tridentata) and big sagebrush (Artemisia tridentata).
Cheatgrass (Bromus tectorum) is the most abundant understory species occurring
on the area. The Moffat County area is approximately 30 square miles and
is enclosed by a sheep-tight fence. Both study areas are grazed by sheep
and cattle~

RESULTS AND DISCUSSION

Mortality

Hunter Harvest

Moffat County Study Area--Hunting permits for the area were of a "specified
permit" type (designating the sex of the animal that could be killed) from
1963 through 1966. In 1967, all permits were of the "either sex" type.
The numbers of permits issued varied throughout the study period
In 1963
and 1964 there were 250 permits (50 buck - 200 doe) issued (Table 1)0
This was reduced to 100 permits in 1965 and 1966; 25 buck - 75 doe, and
50 buck - 50 doe permits, respectively.
Only 60 permits were issued in
1967. Hunter success was very high (over 90 percent) for all years
except 1965, when the success was low due to a heavy snowfall in the area
during the hunting season.
o

�- 40 -

---

Total

Number

Fawns / 100 Does
Bucks/ 100 Does

400

350

300

0:: 250 ~
LLI
OJ

~

::&gt;

2

200

150 -

100 -

50 -

--Fig. 1.
surveys,

Tota 1 antelope
1963-1967.

-- --_ .•- _--_.--

numbers and ratios

..•...•

obtained

on aerial

antelope

�- 41 Table 1.

Hunter success on the Moffat County study area, 1963-1967.
Permits Issued
Doe
Total

Number
Harvested

Percent
Success

250

243

97

200

250

229

92

25

75

100

60

60

1966

50

50

100

92

92

1967

Either-Sex

60

57

95

Year

Buck

1963

50

200

1964

50

1965

Aerial counts indicated a rapid decline in the population from 1963 to the
period following the 1965 hunting season, then the population started increasing (Figure 1). The fawn:doe ratio increased from 75:100 in 1963 to
91:100 in 1964; decreased to 54:100 in 1965, then increased to 71:100 and
90:100 in 1966 and 1967 (significantly different at the 95% confidence level)o
Fawn:adult ratios showed a trend more closely resembling the population
trend, decreasing from 55:100 in 1963 to 43:100 in 1964, and 35:100 in 1965
These ratios then started upward in 1966 and 1967, 47:100 and 59:100 respectively.
The buck:doe ratios showed a slight upward trend under the
restrictive buck permits during the period from 1964 through 1967: 32:100,
40:100, 43:100, and 40:100 respectively.
However, these ratios were not
significantly different at the 95% confidence level
0

The mean age of the does harvested followed a pattern similar to the population trend. The mean age decreased from 203 years in 1963 to 102 years
in 1965, then increased to 204 years in 1966, and down to 2.0 years in the
1967 "either sex" season (Table 2). The fawn harvest was low in 1963, increased and remained high until 1967, when it decreased again (Figure 2)0
The low percentage of fawns in the harvest occurred when the fawn:adult
ratios were high, also when the population was high in 1963 and rapidly
increasing in 19670 Hunters were probably biased in selecting larger
animals when the population was high, and were less selective during the
population lowo Yearling and adult (two years and older) portions of the
harvest were alternately high and low: the adult class being high in the
harvest when the yearling class was low; and the reverse being true alsoo
It stands to reason the adult portion of the population would be low
following a year when yearling harvest was heavy, and the adult portion of
the population would increase following a year when yearling harvest was
light, because of the yearling recruitment into the adult class the following year.

�- 42 -

Table 2. Age of antelope checked at Moffat County hunter check stations,
1963-1967.
Age Class
(years)
1963
Fawn
1
2
3
4
5+
Total
Mean Age
1964
Fawn
1
2
3
4
5+
Total
Mean Age
1965
Fawn
1
2
3
4
5+
Total
Mean Age
1966
Fawn
1
2
3
4
5+
Total
Mean Age
1967
Fawn
1
2
3
4
5+
Total
Mean Age

Number

Bucks
Percent

Number

Does
Percent

Number

Total
Percent

5
16
12
8
6
2
49
2.5

10
33
25
16
12
4
100

26
54
13
18
20
4
135
2.3

19
40
10
13
15
3
100

31
70
25
26
26
6
184
2.3

17
38
14
14
14
3
100

23
14
21

32
19
29
15
6

41
27
20
15
7
8
ll8
2.0

35
23
17
13
6
7
101

64
41
41
26

34
22
22
14
6
4
102

8
10
4
1
0
1
24
1.2

33
42
17
4

II

4
0
73
1.9
4
3
5
1
4
0
17
2.4

101
24
18
29
6
24
101

4
100

II

8
191
2.0
12
13
9
2
4
1
41
1.6

29
32
22
5
10
2
100

6
7
13
8
9
7
50
3.1

12
14
26
16
18
14
100

13
5
4
4
3
5
34
2.4

38
15
12
12
9
15
101

19
12
17
12
12
12
84
2.8

23
14
20
14
14
14
99

6
13
5
3
3
3
33
2.3

18
39
15
9
9
9
99

5
6
4
2
1
1
19
2.0

26
32
21
11
5
5
100

11
19
9
5
4
4
52
2.2

21
37
17
10
8
8
101

�- 43 -

-------

Fawns
Yearlings
--

Ad u Its

50

,"

,
, \ V'
,
-. r: -/__/0
'\
" /,'
\
,
• I \

40

\

'\

IZ
ltJ

o

30

/ ,/

0.:::

\

-,

W
CL

\,

~

\

/

,,

\
\

20

"

"
/\

I

\
\

I

\ I

V

10

o =---19t63~~==""~~~1~9~6-4--",,=-~o'~I~"65~~~=~'=-'19-~G==-~-~'=--=r96J7~-'~c=
YEAR
Fig.2.
Percentage of fawn, yearling,
harvest, Moffat County, 1963-1967.

and adult antelope

occurring

in the

�-44

There did not appear to be any relationship between the percentage of
yearlings or adults in the harvest and the population trend. However,
graphs obtained by plotting the number of yearlings and adults in the
harvest, rather than using percentages, more closely resembled the
population curve.
Sample sizes for the doe segment in the various age
classes during the last three years were quite small. For example,
numbers in the yearling class were only 10, 5, and 6. Thus, the age
structure based on these small samples may be misleading.
Mean age of bucks decreased in 1964, then gradually increased until
1966. In 1967, when the either sex permits were issued, the mean age
dropped again.
Hunters were noticeably biased in selecting their bucks.
When large bucks were fairly numerous and easily found, hunters put
forth more effort in obtaining animals with large horns. Therefore,
the buck harvest is less apt to reflect the age structure of the population than the doe segment of the harvest.
An estimate of the pre-season population was derived from total counts
and fawn ratio counts.
For example, 428 animals were counted in the
spring of 1963. A ratio of 55 fawns per 100 adults was obtained on a
flight in August.
By multiplying 55 percent times 428 adults there was
estimated to be 235 fawns, added to 428 adults equals 663 animals (Table
3). Hunter harvest was combined with the estimated wounding loss for
1963 to calculate an overall herd drain of 282 animals.
A percent harvest figure was obtained by dividing the harvest by the estimated population. The percent of the total herd harvested in 1963 through 1967
was 42, 48, 17, 40, and 21, respectively.
In relating the percent harvest to the population trend, we see that the population decreased
slightly with a 42 percent harvest, then sharply declined following a
48 percent harvest the next year. There was a slight increase with a
17 percent and 40 percent harvest.
The population increased sharply
following the 21 percent harvest in 1967. It appears a harvest level
somewhere between 20 percent and 40 percent would have maintained a
stable population.
These data were not further presented in life tables because the mortality rate was not consistent from one year to the next, the recruitment
rate was not constant, and the age structure of the harvest was not consistent between years.
There was a tendency for hunters to record fawns as adult animals on
their report cards.
Check station data showed that fawns made up a
larger portion of the harvest than indicated by report cards (Table 4).
An average for the five year period indicated fawns comprised 15 percent
of the report card returns, and 24 percent of the antelope checked at
the check stations.
The sample size was very good; 88 percent of the
antelope harvested on the study area during the five-year period were
"aged" by check station personnel.

�- 45 Table 3.

Hunter harvest of antelope on the Moffat County study area, 1963-1967.

Hunting Season

1963

Harvest
Wounding Loss
Total

Bucks

Does

Fawns

Total

54
9
63

138
3
141

51
27
78

243
39
282

~'&lt;'Estimated
preseason population: 428 ad + 235 fawns
Percent of population harvested: 42%
1964

Harvest
Wounding Loss
Total

59
21
80

92
15
107

78
15
93

663

229
51
280

Estimated preseason population: 410 ad + 176 fawns = 586
Percent of population harvested: 48%
1965

Harvest
Wounding Loss
Total

19
1
20

24
2
26

17
0
17

60
3
63

Estimated preseason population: 280 ad + 98 fawns = 378
Percent of population harvested: 17%
1966

Harvest
Wounding Loss
Total

49
1
50

22
3
25

21
0
21

92
4
96

Estimated preseason population: 165 ad + 78 fawns = 243
Percent of population harvested: 40%
1967

Harvest
Wounding Loss
Total

24
0
24

13
1
14

20
1
21

57
2
59

Estimated preseason population: 178 ad + 105 fawns = 283
Percent of population harvested: 21%

-k

Based on aerial surveys.

�- 46 Table 40 Percentage of bucks, does and fawns harvested on the Moffat
County study area based on hunter report cards and check station data,
1963-1967.

Bucks

Year

Hunter ReEort Cards
Bucks
Does
Fawns
Total
(%)
(%)
(%)
Number

Check Station
Does
Fawns

(%)

(%)

(%)

Total
Number

1963

20

75

5

243

24

60

16

189

1964

21

53

26

229

26

40

34

191

1965

40

33

27

60

32

39

29

41

1966

45

38

17

92

50

27

23

82

1967

70

23

7

57

52

27

21

52

Mean

30

55

15

32

44

24

Some of the hunter observations followed a pattern similar to the population
trend (Table 5). The number of wounded antelope reported by hunters for
the five-year period was 68, 51, 3, 4, and 2. There was a very high wounding
loss during the years of high antelope densities, but this loss decreased
with a population decline.
The time required for a successful hunter to
bag an animal more than doubled as the population declined.
The number of
antelope observed per hunter per hour was very closely related to the time
each person spent hunting.
The ratio of bucks to does in the herd increased
under· the restricted "specified sex system", this same trend was genera lly
reflected in the number of bucks observed by the hunters.

Larimer County Study Area--Approximate1y
90 percent of the study area has
closed to antelope hunting since 1966, therefore, only three years (19631965) of reliable hunter harvest data were obtained.
This is not an
adequate sample to relate hunter harvest to population trends. Aerial
surveys indicated a general upward trend for this herd from 1963 to 1967,
during which 243, 209, 231, and 305 antelope were counted.
The buck:doe
ratios were 26:100, 21:100, 38:100, and 46:100, respectively.
The me.an age of antelope harvested ranged from 2.1 years to 3.0, which
was slightly higher than the mean age of animals cropped on the Moffat
County area (Table 6)0 The Larimer County herd was more static or stable,
thus more older animals appeared in the harvest.

�- 47 -

Table 5. Miscellaneous information collected from successful hunters at the Moffat
County antelope check stations, 1963-1967.

Category

1963

1964

47
2

70
8

135
5

Year
1965

1966

1967

17

43
5

29
5

113
19

24

37
1

18
2

182
7

183
27

41

80
6

47
7

49
92
1.3
71
25

51
52
3.3
16
31

17
32
2.9
11
36

48
40
4.1
10
26

35
28
5.9
5
24

140
104
2.0
52
31

148
47
4.5
10
75

23
50
3.8
13
45

38
37
4.3
9
47

19
18
6.1
3
23

189
101
1.6
63
29
68
54

199
48
4.1
12
60
51
26

40
42
3.4
12
42
6

86
39
4.3

54
24
6.0
4
24
4
4

Harvest
Bucks
First day
Second day
Does
First day
Second day
Both Sexes
First day
Second day
Hunter Observations
Buck Hunters
Number of hunters checked
Antelope observed/hunter
Hours hunted/hunter
Antelope observed/hunter/hour
Bucks observed/100 does
Doe Hunters
Number of hunters checked
Antelope observed/hunter
Hours hunted/hunter
Antelope observed/hunter/hour
Bucks. observed/IOO does
Buck and Doe Hunters
Number of hunters checked
Antelope observed/hunter
Hours hunted/hunter
Antelope observed/hunter/hour
Bucks observed/IOO does
Wounded antelope observed
Abandonded antelope observed

9

36
15
3

�- 48 Table 6. Age of antelope checked at the Larimer County hunter check stations,
1963-1965.
Age Class
(years)
1963
Fawn
1
2
3
4
5+
Total
Mean Age
1964
Fawn
1
2
3
4
5+
Total
Mean Age
1965
Fawn
1
2
3
4
5+
Total.
Mean Age
1966
Fawn
1
2
3
4
5+
Total
Mean Age

Bucks
Number
Percent

Number

6
11
9
1
6
3
36
2.5

17
30
25
3
17
8
100

3
4
3
1
2
1
14
2.4

21
29
21
7
14
7
99

9
15
12
2
8
4
50
2.5

18
30
24
4
16
8
100

5
6
8
3
4
1
27
2.4

18
22
30
11
15
4
100

4
1
1

67
17
17

6
0.8

101

9
7
9
3
4
1
33
2.1

27
21.
27
9
12
3
99

1
4
4

8
33
33

1
2
3

13
25
37

2
1
12
2.6

17
8
99

2
8
3.5

25
100

1
5
6
3
2
3
20
3.0

5
25
30
15
10
15
100

2

100

2

40

1
1
1

20
20
20

5
2.3

100

1
1
1

33
33
33

3
3.5

99

2
0.5

Does
Percent

100

Number

Total
Percent

�- 49 Hunters tended to report fawns as adult
Check station data indicated 18 percent
1963 to 1965, while hunter report cards
of eight percent fawns (Table 7). This
for the Moffat County study area.

antelope on the report cards.
of the harvest was fawns from
indicated the harvest consisted
is very similar to the results

Observations made by hunters were somewhat erratic and didn't follow the
trends established by the aerial surveys during the three-year period.
The average number of antelope observed per hunter for 1963, 1964 and
1965 hunting seasons was 14, 25, and 40, respectively (Table 8). Only
five hunters were checked in 1966 so that year's data is not included
in this discussion.
These averages are considerably lower than those
found for the Moffat County area, however, the density of antelope on
the Moffat County area was much greater.
Hunters on the Larimer County
area required 4.6, 8.1, and 3.2 hours to bag their animals during these
respective years.
There was considerable rain during the 1964 season,
thus poor weather likely accounted for the high number of hours (8.1)
required to make a kill. The average number of bucks observed per 100
does during the 1963-65 seasons was 22, 29, and 21, respectively.
Hunters on the Larimer County area observed two wounded antelope each
year. This was much lower than the wounding observations on the Moffat
County study area. Again, this is likely due to the lower antelope
densities on the Larimer County area.

Table 7. Percentage of bucks, does, and fawns harvested on the Larimer
County Study area based on hunter report cards and check station data,
1963-1965.

Bucks

Year

Hunter ReEort Cards
Bucks
Does
Fawns
Total
(%)
(%)
(%)
Number

(%)

Check Station
Does
Total
Fawns
(%)
(%)
Number

1963

71

21

8

87

58

26

16

55

1964

72

16

12

57

67

6

27

33

1965

59

35

6

105

55

40

5

20

Mean

66

26

8

60

22

18

�- 50Natural Mortality
Mortality due to predation and disease appeared to be very light among the
older fawns and adult antelope.
Cottontail rabbits and jackrabbits were
abundant on both areas and appeared to be one of the major food items for
predators.
Rabbit and squirrel remains were commonly found around the
eagle nests.
Sheep ranchers in the Moffat County study area maintained
very intense coyote control programs and the coyote population was very
low. Whereas, aerial antelope surveys on the Larimer County area indicated there was approximately one coyote per five square miles during
the winter months, but only about one coyote per seventy-five square miles
during the summer.
Coyotes apparently frequent the plains, and the areas
adjacent to them during the winter months, then move into the foothills
during the summer. Golden eagles were commonly seen on both study areas.
During the five-year period of this study only three antelope (two fawns
and one yearling doe) were suspected of being killed by coyotes.
One
fawn was killed by an eagle. Also, a doe fawn on the Larimer County area
was observed during February, 1964 to have deep wounds on her back, which
were thought to have been caused by an eagle. The fawn recovered and was
observed to be in good health at a later date.
During June, 1965, a doe on the Moffat County study area was observed
pursuing three badgers (an adult female and two partially grown young)o
The badgers were collected and their stomachs examined.
The stomachs of
the two young badgers were gorged with the remains of an antelope fawn.
Mortality due to diseases and parasites was negligible, as nearly as could
be ascertained in this study. A total of 56 dead antelope were examined
for parasites and diseases during the study periodo
Three of these
antelope had one to three wood ticks on them, one had necrotic stomatitis,
one a tapeworm cyst in the mesentery, and another a light infestation of
nematodes in the small intestine.
During March, 1966, two doe antelope were collected in Moffat County for
the physiological study. They both had abnormally long hooves and slightly
swollen leg joints.
One of these does chose to lie down and hide when she
was approached rather than flee with the other antelope associated with her.
Later she got up and attempted to move off, but had extreme difficulty in
walking.
The second doe was thought to be a healthy animal prior to collection. There were 79 other antelope observed in the area that morning and
five of these showed obvious signs of being lame. The lameness was suspected to be caused by a polyarthritis similar to a type occurring in
domestic sheep. Unsuccessful attempts were made to capture one of these
lame animals for intensive laboratory study. The lameness was evident only
during cold wet weather.
Carcasses of two very young fawns that died from some unknown cause were
found during the five-year study period. They had died while in a normal
resting posture, and obviously had not been molested or killed by predators.
Eartag returns and general observatiDns indicate there may be a high mortality among very young fawns, but carcasses were not found to substantiate
this.

�- 51 -

Table 8. Miscellaneous
Larimer County antelope

information collected from successful
check stations, 1963-1966.

hunters

at the

Year
Category

1963

1964

1965

1966

24
15

17
11

14

3
1

11
5

5
1

10

0
2

35
20

22
12

24

3
3

67
12
4.7
3
25

28
26
8.0
3
28

14
50
3.6
14
19

4
26
5.0
5
46

26
19
4.5
4
19

5
18
8.8
2
36

10
27
2.6
10
27

2
2
4.0
1
0

93
14
4.6
3
22
2
3

33
25
8.1
3
29
2

24
40
3.2
13
21
2
1

6
18
4.7
4
42
0
0

Kill
Bucks
First day
Second day
Does
First day
Second day
Both sexes
First day
Second day
Hunter Observations
Buck Hunters
Number ot' hunters checked
Antelope observed/hunter
Hours hunted/hunter
Antelope observed/hunter/hour
Bucks observed/100 does
Doe Hunters
Number of hunters checked
Antelope observed/hunter
Hours hunted/hunter
Ante lope observed/hunter/hour
Bucks observed/100 does
Buck and Doe Hunters
Number of hunters checked
Antelope observed/hunter
Hours hunted/hunter
Antelope observed/hunter/hour
Bucks observed/100 does
Wounded antelope observed
Abandoned antelope observed

�.• 52 -

Two adult antelope were known to have been struck and ki~led by automobiles on the highways associated with the study areas. The highway
"right-of-ways" were fenced with net-wire, and it was rare to see
antelope on the right-of-way.
Numerous antelope were found tangled
in fences in the vicinity of the study area. This was discussed in
the report on "antelope and net-wire fences" (Work Plan 2, Job 5) of
this project.

Aging Studies

Tagging

Fawns

A total of 167 fawns was tagged on the study areas. There were 97
females and 70 males in this sample. There was a total of 19 tagged
antelope (13 females and 6 males) known to have been harvested by hunters and one fawn was found dead a few weeks after tagging (Table 9).
Nine of the tagged antelope were shot when fawns, five as yearlings,
four ~s two-year olds, and two as three-year olds. This was a 12 percent return on the tags, and most of these were one year old or younger.
Only three marked animals are known to exist on the study areas at the
present date. It is interesting to note that similarity to findings of
Hoover, et, al. (1959) who tagged fawns on the same area in Larimer County
during the period 1947-1954.
They averaged 11.8 percent return on their
tags and all returns' came back within three years of the tagging date.
It appears t here is a very high mortality rate among tagged fawns.
Measurements were obtained from only a few marked animals following harvest. Therefore, data based on known-age antelope were very limited and
of little value in this study.

Table 9. Antelope
1964-l967~"•

tag return data, Larimer and Moffat

County study areas,

Age
(years)

Male

Sex
Female

Both

Fawn

3

6

9

1

1

4

5

2

2

2

4

3

0

2

2

Total

6

14

20

* Antelope were tagged as fawns less than three days old.

�- 53 Eye Lens Weights
A sample of the eye lenses were dried 192 hours to determine the rate
of decrease in weight when dried longer than 72 hours as used by Kolenosky
and Miller (1962). Dry weights obtained at 24 hour intervals showed an
average decrease in weight of 1.1, 0.9, 0.7, 0.5, and 0.4 percent (Table
10). This weight loss beyond the 72 hour drying time did not appear to
be significant, therefore, the 72 hour drying time was used in this study.
Ashed weights of 40 eye lenses are compared with their respective dry
weights in Table 11. The sample size is too small for a valid comparison,
however, there did not appear to be much difference between the trend
established by the dry lens weights as compared with the ash weights of
the same lenses. The ashing technique required less time, however, this
technique involved small weights (less than 20 mg) and appeared to be
more subject to errors. Therefore, the dry lens weights were used here.
Mean lens weights increased for each age class from fawns to the five year
old class. The increase was more rapid between the fawn, one year old,
and two year old age classes; increasing from 342.0 mg to 570.6 mg and to
662.0 mg, respectively (Table 12). The mean weights for the three year
old, four year old, and five year old (plus) classes were 743.6 mg, 749.3
mg, and 799.7 mg, respectively; therefore, lens growth was much slower in
the older 'age classes. Kolenosky and Miller (1962) found a very similar
growth pattern for antelope eye lenses collected in Saskatchewan.
There was considerable variation in the data and overlap in the confidence
intervals established for each age class older than "fawn".
Since the
age of each antelope was determined by the "tooth erruption and wear
technique", the eye lens technique is subject to the same errors. Ko1enosky
and Miller concluded "However, it has been demonstrated that lens growth
proceeds at a species-predictable
rate in antelope, and the growth curves
that have been established in this study can be verified when tagged
specimens of known age are available".
The results obtained in this study
further emphasize the need for known age antelope to fully evaluate eye
lens growth.

Mandible

and Head Measurements

Skull measurements increased from the fawn age class to yearlings, then
tapered off in the older age classes.
Fawns had a significantly (90 percent confidence level) shorter head than older animals; there was not a
significant difference in mean lengths for the older age classes (Tables
13 and 14). There was not a significant difference between bucks and does.
The average head length for fawns was approximately 22-23 centimeters;
yearlings, 28 centimeters, and adult antelope, 29-30 centimeters
Average
skull width measurements were not significant between any of the age
classes, however, there was a slight increase in width from the fawn age
class through the yearling age class.
Skull widths of buck and doe
antelope were very similar at the fawn age class (approximately 8.5 centimeters).
These widths for does increased to 9.6 - 10.0 centimeters for the
older age classes, while buck skull widths averaged 10.8 - 11.7 centimeters.
o

�Table 10. Weight of antelope eye lenses when dried at 60 degrees centigrade.

72
Hours

96
Hours·

Weight
Lost

120
Hours

Weight
Lost

Lens Weight {mgl
144
Weight
168
Hours
Lost
Hours

16
17
18
19
20

594.9
597.2
561.3
516.9
327.2
327.8
582.8
586.0
611.9
694.4
625.0
573.6
758.7
360.8
676.8
590.6
577.7
517.5
509.2
614.6

588.5
585.1
553.7
512.4
323.5
325.4
574.1
577 .2
610.2
683.5
614.3
571.2
750.6
357.1
666.5
588.9
567.7
515.8
507.4
610.5

6.4
12.1
7.6
4.5
3.7
2.4
8.7
8.8
1.7
10.9
10.7
2.4
8.1
3.7
10.3
1.7
10.0
1.7
1.8
4.1

579.3
576.9
547.3
510.5
320.3
321. 9
561.9
567.5
610.0
674.9
605.5
570.1
745.7
353.6
658.4
588.8
558.8
515.8
506.8
608.2

9.2
8.2
6.4
1.9
3.2
3.5
12.2
9.7
0.2
8.6
8.8
1.1
4.9
3.5
8.1
0.1
8.9
0.0
0.6
2.3

573.3
572.2
543.5
509.2
318.4
319.3
551.7
559.3
609.6
670.3
600.3
569.3
742.9
348.3
653.4
587.9
552.9
515.0
506.0
606.6

6.0
4.7
3.8
1.3
1.9
2.6
10.2
8.2
0.4
4.6
5.2
0.8
2.8
5.3
5.0
0.9
5.9
0.8
0.8
1.6

Total

11204.9

11083.6

121.3

10982.2

101.4

10909.4

Mean

560.2

554.2

6.1

549.1

5.1

545.5

Number

1
2
3
4
5
6
7
8
9
10
11
12
13
14

IS

Percent
Decrease

1.1

0.9

Weight
Lost

192
Hours

Weight
Lost

568.3
568.6
540.6
508.1
316.8
317.3
543.9
553.4
609.1
666.8
595.8
568.7
741.4
343.5
650.0
587.4
548.2
514.7
505.5
605.4

5.0
3.6
2.9
1.1
1.6
2.0
7.8
5.9
0.5
3.5
4.5
0.6
1.5
4.8
3.4
0.5
4.7
0.3
0.5
1.2

562.7
565.0
538.2
507.9
315.1
315.4
538.1
548.3
608.6
663.5
591. 9
568.5
741.3
340.0
646.5
586.8
543.2
514.7
505.5
604.7

5.6
3.6
4.4
0.2
1.7
1.9
5.8
5.1
0.5
3.3
3.9
0.2
0.1
3.5
3.5
0.6
5.0
0.0
0.0
0.7

72.8

10853.5

55.9

10805.9

47.6

3.6

542.7

208

540.3

2.4

0.7

0.5

0.4

VI

.p-

�- 55 -

Table 11. Comparing
related to age.

dry lens weight and the ashed weight of the lens as

Age Class
(years)

Number
in Sample

Fawn

9

344.8

12.6

7.3

0.5

1

15

549.6

25.5

1005

0.7

2

10

660.2

45.5

12.9

2.8

3

1

837.9

4

3

795.6

5+

2

801.0

Table

120

Dry eye lens weights

Dry Lens Weight {mg2
Mean
Std. Dev.

Ash Weight
Mean

of Lens {mg2
Std. Dev.

1602
14.2

34.8

0.4

14.1

as related to age for 270 ant e Lope iw

Age Class
(years)

Number
in Sample

Standard
Deviation

90% Confidence

Mean

Fawn

60

342.0

27.2

362.2 - 321.8

1

74

570.6

70.6

681.2 - 460.0

2

61

662.0

55.9

746.0 - 578.0

3

27

743.6

45.4

869.0 - 618.2

4

29

749.3

66.7

1000.6 - 498.0

5+

19

799.7

83.7

1197.6 - 401.8

~': Lens

weights

are given in milligrams

(mg) •

Intervals

�- 56 -

Mandible measurements increased in the younger age classes and was more
constant among the older age classes. Mean ramus and diastema lengths
were significantly different for fawns and yearlings.
There was not a
significant difference between the means for the older age classes.
There was no apparent difference due to sexo The mean ramus length
for fawns was 160 millimeters; yearlings, 209 millimeters, and older
animals, 217-220 millimeters (Tables 15 and 16). The mean diastema
length for fawns was 49.6 millimeters, yearlings 69.3 millimeters, and
older antelope 71.4-76.1 millimeters.

Horn Length
Horn length data collected at check stations during 1963 and 1964 showed
a rapid horn growth until the animal was approximately two years old.
Mean horn length for fawns and yearling bucks were 1 6 inches and 7.8
inches, respectively.
There wa.s a significant difference (90 percent
confidence level) between these age classes and older animals (Table 17).
Age classes two years and older had means greater than 12 inches, however,
there was not a significant difference between these classes.
These data
indicated buck antelope could be grouped into three distinct categories:
fawns, yearlings, and adults (two years and older), by using horn length
measurements.
0

During 1965 and 1966 report cards (145) were issued to successful buck
hunters when their antelope were examined at check stations.
Sixty-nine
percent of the report cards were returned.
Antelope ages were established by
using horn lengths recorded on the report cards. These data were related
to t~e ages established for the same antelope at the check stations.
It
was determined that the horn length data had a six percent error. Therefore, this technique appears to be adequate for establishing a general
age structure for buck antelope.
Ten years of data from Colorado indicates 60 to 72 percent of the annual harvest was bucks.
Based on a
sixty-nine percent report card return, ages could be determined for approximately forty percent of the antelope harvested in Colorado each yearo
This is a very easy technique to use, requiring only a measuring tape.
Since antelope harvest information in Colorado relies on hunter report
cards, this technique could be used to supplement harvest information.
It still needs to be tested on a larger scale and related to changes in
population structure.

�- 57 -

Table 13. Skull length and width measurements
by hunters.*

Age Class
(years)

Number in
Sample

for buck antelope

harvested

Mean
(± Standard Deviation)
Length
Width

90% Confidence
Length

Limits
Width

Fawn

10

23.0 + 1.9

8.6 + 0.7

19.9 - 26.1

7.5 - 9.7

1

21

27.8 + 0.6

10.8 + 0.8

26.8

28.8

9.5 - 12.1

2

19

28.8 + 1.7

11.3 + 0.8

26.0 - 31.6

10.0 - 12.6

3

2

29.7

11.4

4

12

29.8 + 0.2

11.7 + 0.8

29.4 - 30.2

10.4 - 13.0

5+

1

30.2

12.5

~~Measurements

are in centimeters.

Table 14. Skull length and width measurements
by hunters. *

Age Class
(years)

Number in
Sample

for doe antelope harvested

Mean
~+ Standard Deviation2
Length
Width

90% Confidence Limits
Length
Width

Fawn

5

21.7 + 0.8

8.4 + 0.7

20.4 - 23.0

7.3 - 9.5

1

5

28.4 + 1.6

9.6 + 0.5

25.8 - 31.0

8.8 - 10.4

2

5

29.3 + 0.9

9.7 + 3.1

27.8 - 30.8

4.7 - 14.7

3

5

29.2 + 1.5

10.0 + 1.4

26.7 - 31.7

7.7 - 12.3

4

2

28.5

9.6

5+

3

29.3

9.8

Measurements

are in centimeters.

oJ,

�- 58 -

Table 15. Relationship of ramus length and age for 153 antelope mandibles
collected at hunter check stations.
Number
In Sample

Mean
Length (rrnn)

90% Confidence
Interva 1 (mm)

Fawn

14

161

158 - 165

1~

23

207

204 - 210

2~

18

218

215 - 222

3~

6

224

220 - 228

4~

7

220

214 - 226

5~ - over

1

214

Fawn

23

160

159 - 161

1~

15

209

206 - 213

2~

20

215

212 - 218

3~

11

214

210 - 218

4~

8

219

214 - 224

5~ - over

7

221

216 _ 226

Fawn

37

160

158 - 162

1~

38

209

201 -211

2~

38

217

215 - 219

3~

17

218

211 - 225

4~

15

219

216 - 222

5~ - over

8

220

215 - 225

Sex and Age
(years)
Males

Females

Both Sexes

�- 59 -

Table 16. Relationship of diastema length and age for 153 antelope mandibles
collected at hunter check stations.
Sex and Age
(years)

Number
In Sample

Mean
Length (nnn)

90% Confidence
Interval (nnn)

Fawn

14

49.5

47.7 - 51.3

llz

23

69.2

67.6 - 70.8

2lz

18

72 .1

70.6 - 73.6

3lz

6

74.9

72 .4 - 77.4

4lz

7

72 .1

69.9 - 74.3

5lz - over

1

70.5

Fawn

23

49.7

48.8

llz

15

69.4

67.1 - 71.1

2lz

20

70.3

68.8 - 71.8

3lz

11

70.0

68.0 - 72.0

4lz

8

72.5

70.7 - 74.3

5lz - over

7

76.9

74.1 - 79.7

Fawn

37

49.6

48.8 - 50.4

llz

38

69.3

68.0 - 70.6

2lz

38

71.4

70.4 - 72.4

3lz

17

71.7

69.9 - 73.5

4lz

15

72 .3

71.1 - 73.5

5lz - over

8

76.1

72 .9 - 79.3

Males

Females
50.6

Both Sexes

�- 60 -

Table 17. Relationship between the horn length and age for 219 buck
antelope.*
Age Class
(years)

Number
in Sample

Mean Horn
Length

Standard
Deviation

90% Confidence
Interval

Fawn

26

1.6

0.4

0.9 -

1

71

7.8

1.4

5.5 - 10.1

2

62

12.1

1.2

10.2 - 14.0

3

31

13.5

1.1

11.7 - 15.3

4

22

14.0

1.3

11.9 - 16.1

5+

7

13.5

1.8

10.5 - 16.5

2.3

* Measurements are given in inches.

LITERATURE CITED
Hoover, R. L., C. E. Till, and S. Ogilvie. 1959.
Colo. Dept. Game and Fish, Tech. Bull. No.4.

The antelope of Colorado.
110 p.

Ko1enosky, B., and R. S. Miller. 1962. Growth of the lens of the pronghorn
antelope. J. Wildl. Mgmt. 26(1): 112-113.

Prepared by

,--.~.!J,
~.
George':Bear
Wildlife Researcher

�January,

- 61 -

1970

JOB FINAL REPORT

State of
No.

W:-40-R-10

Antelope

Investigations

Plan No.

1

Job No.

12

Project
Work

COLORADO

Job Title:

Antelope

Period Covered:
Personnel:

April

George

Census

1, 1963 to April 30, 1969

D. Bear

ABSTRACT

Aerial counts were generally highest in the early morning hours and lowest
during mid-day.
Antelope were most easily counted on clear days and when
vegetation was very green.
Air conditions generally had little effect on
the total number of antelope counted, but as the air became more turbulent
errors·increased
in the sex and age ratios.
Peak fawn counts were obtained
in late August and early September just before the hunting seasons.
Highest total counts were obtained from November through March when the herds
were large.
Direction of flight appeared to have little effect on counting
accuracy.
There was little difference between the total number of antelope
counted with the helicopter and total number counted with a fixed-wing
aircraft during the winter months.
The helicopter appeared to improve the
accuracy of classifying bucks, does and fawns.
Trends in population numbers
and fawn:doe ratios were reasonably accurate in the 60 and 40 percent sample;
but this sampling technique did not yield reliable information for buck:doe
ratios.
The permanent road transects were extremely variable and unpredictable.

�- 62 -

RECOMMENDATIONS

Recommendations

for aerial surveys are as follows:

10

Conduct counts at daylight,
free from snow.

2.

Make total annual counts with sex ratio classification
winter, or late winter and early spring.

3.

Make fawn counts in late August or September

4.

A rating sheet should be completed for each flight, evaluating
conditions and recording the time of flight.

5.

The feasibility
be explored.

on a clear day, and when the ground is

of establishing

in early

before the hunting

season.
census

trends from sample surveys needs to

Contents of this report were summarized and presented in the following
publication:
Evaluation of Aerial Antelope Census Techniques.
Colorado
Game, Fish and Parks Division, Outdoor Facts, Game Information Leaflet
Number 69 (in print).

�- 63 -

ANTELOPE

CENSUS

George D. Bear

p. S. OBJECTIVE
Develop

technique

for censusing antelope.

METHODS AND MATERIALS

Aerial

Variation

Census

Due to Time and Weather

Aerial counts were conducted with a fixed-wing airplane monthly on each
study area from June, 1963 to March, 1965 to determine daily and seasonal
variability in the aerial census technique.
Counts were conducted three
times a day, early morning, mid-day, and late afternoon, to determine the
most suitable time for aerial census. The Larimer County area was flown
in mile-wide strips.
Permanent land marks were used as boundary markers
to prevent overlap.
Due to shrub cover and concentration of antelope it
was most desirable to fly the Moffat County area in one-half mile-wide'
strips. A flying altitude of approximately 100-300 feet was maintained.
Other factors recorded on each count were: vegetative cover, relative
visibility due to ground cover, cloud cover, wind velocity, distribution
of antelope within the herd, approximate location on the study area
(strip number and distance along the strip), and activity of the antelope.
Ground cover was rated in one of three classes:
I - new snow, 100% snow
cover; II - 100% snow cover, but old snow with a lot of old tracks; and
III - (a) spotty snow cover, (b) bare ground. Light conditions were rated
in one of three classes:
I - clear skies; II - broken overcast 50 percent
of the time; and III - solid overcast.
Air conditions were also rated in
one of three classes:
I - good, solid air; II - mild to moderate downdraft and turbulence; and III - severe turbulence and downdrafts.

Variation

Due to Type of Aircraft

Comparisons were made between aerial census data collected using a helicopter and a fixed-wing airplane.
Helicopter and airplane counts were made
in the same areas and under similar conditions when possible.

Variation

Due to Flight Direction

Fixed-wing

flights were made from May, 1965 to August,

1967 to determine

�- 64 the effect of flight direction on census variability.
Flights were
limited to early morning hours and clear calm days when possible.
Two
flights were made on the same area on successive days. The first-day
flight was conducted in an east-west direction and the second-day flight
in a north-south direction.

Sample Trends
Trends established from random samples were compared with trends established from counts of the total land area. Aerial surveys were flown in
the fall and late winter from 1963 to 1967. The entire areas were flown
in strips, as described above. Trends were plotted for the total number
of antelope counted, buck-to-doe ratios, and fawn-to-doe ratios. Three
samples (60 percent, 40 percent, and 20 percent of the total land area
surveyed) were selected and similar trends plotted for each sample. The
strips used in each sample were selected using a "Table of Random Numbers".

Ground Census
Ground counts, using binoculars and a spotting scope, were conducted from
a vehicle while driving along a permanent transect in each study area.
The road transects were 23 miles and 11 miles long on the Larimer and Moffat
County study areas, respectively.
These counts were made once a month,
three times a day (morning, noon, and evening).
The sex, age, and location
were recorded for all antelope sighted.
Light and ground cover were recorded as for the aerial counts. Road conditions were recorded in one of
three classes:
I - good, eaSily traveled; II - fair, occasional places
of difficult travel; and III - poor, difficult to travel, requires driver's
attention most of the time.

DESCRIPTION

OF AREA

Study areas were established in Larimer and Moffat Counties where major
antelope concentrations exist in two different vegetative types. The
Larimer County study area is located north of Fort Collins, adjacent to
the Colorado-Wyoming
state line. The area is approximately 88 square miles.
The terrain is gently rolling and the vegetation primarily blue grama,
buffalo grass, and other typical shortgrass prairie flora.
Shrubs such as
four-wing saltbush and winter fat are common only in the bottom lands in
the southern portion of the study area.
The Moffat County study area is in the sandhills immediately south of
Maybell.
It consists of a single pasture, approximately 30 square miles,
which is enclosed by sheep-tight fencing. The area has rolling terrain.
Bitterbrush, sagebrush, and rabbitbrush are the dominant vegetative species
on the area, bitterbrush being the most important.
Cheatgrass is the most
abundant understory vegetation.
Antelope on this study area are confined by
sheep-fences; whereas on the Larimer County study area there appears to be
a limited amount of movement between the study area and adjacent lands.

�- 65 -

RESULTS AND DISCUSSION

Aerial

Variation

Census

Due to Time and Weather

Due to the large number of variables associated with each flight and the
small number of flights, the data testing daily, seasonal, and direction
of flights could not be given statistical parameters.
The cOmments
offered here are derived from visual evaluation of the data.
Aerial antelope counts were generally highest in the early morning hours
and lowest during mid-day (Tables 1 and 2). Evening observations were
between these counts.
Light conditions seemed to be one of the most
important factors affecting the counts. When skies were cloudy the
antelope were harder to see, thus counts were lower than expected under
good light conditions.
Ground cover was also important.
Bare ground
was the best background for spotting antelope; with an ideal situation
when the vegetation was growing and very green.
Spotty-snow cover was
the poorest because antelope blended into the background.
The Moffat
County area is a shrubby vegetative type, thus even when there was a
couple feet of snow on the ground the shrubs were sticking up through
the snow, therefore presenting a very mottled background.
Air conditions
generally had little effect on the total number of antelope counted.
However, as the air became more turbulent the sex and age ratio counts were
subject to additional error.
This was because the pilot could not fly
safely at low enough altitudes to distinguish the sex or age of antelope.
There wasn't a definite pattern in the age or sex ratios due to census
conditions.
Antelope herds were small, approximately 3-15 animals, throughout the
summer months, until October.
Then animals grouped into large winter
herds.
High total counts and fawn counts were obtained in late August and
early September just before hunting season. By October fawns were difficult to distinguish from adult antelope.
Large total counts were obtained
from November through March when the herds were large. Young bucks were
difficult to detect in the larger herds during this period. Variability
in the counts due to the time of day was not as critical since the large
groups were easily seen even during mid-day.
Variability in the counts
during the winter months was due to the inability of the observer to get
an accurate count on the larger herds of 75-200 animals.
It took less
time to fly the aerial counts on the study areas when antelope were in
large herds than during the summer months when they were scattered.
Data
from both study areas indicated similar trends.

Variation

Due to Type of Aircraft

Several types of aircraft were used for the census work. These included
a DeHavi1and Beaver, PA18 Piper Cub, Cessna 180, Cessna 182, Hughes Helicopter, and a super charged Bell 47G3B helicopter.
The fixed-wing aircraft were of the high-wing type, which are recommended for census work.

�Table 1. Aerial census with a fixed-wing airplane on the Larimer County antelope study area, 1963-1964.

Month

Time

Census Conditions
Ground Light Air

Antelope Counted
Bucks Does Fawns Total

Buck-Doe-Fawn
Ratio

Census Time
(hours)

Ave. Herd
Size

June - 1963 Morning
Noon
Evening

III
III
III

II
III
II

I
II
II

29
39
47

39
45
50

4
3
14

72
87
111

74-100-10
87-100-7
94-100-28

1.33
1.37
1.47

3.7
2.8
4.3

July

Morning
Noon
Evening

III
III
III

I
II
III

I
II
II

50
66
22

109
34
45

66
30
34

225
130
101

46-100-61
194-100-88
49-100-76

1.47
1.07
1.23

4.8
5.7
5.1

August

Morning
Noon
Evening

III
III
III

II
III
II

I
II
II

41
36
46

78
73
57

38
53
43

157
162
146

53-100-49
49-100-73
81-100-75

1.15
1.20
1.37

4.1
6.2
4.9

September

Morning
Noon
Evening

III
III

No flight
I
II
I
I

60
74

120
108

96
90

276
272

50-100-80
69-100-83

1.25
1.33

6.7
5.9

ANTELOPE HUNTING SEASON
October

Morning
Noon
Evening

III

November

Morning
Noon
Evening

III
III
III

December

Morning
Noon
Evening

January-1964 Morning
Noon
Evening

III

I
I
No flight
I
I

40

63

52

155

63-100-83

1.20

5.0

29

68

30

127

43-100-44

1.22

6.4

II
II
I

33
13
21

125
26
80

85
26
62

243
65
163

26-100-68
50-100-100
26-100-78

1.22
1.10
1.25

15.2
13.0
16.3

III
III

No flight
III
I
III
I

49
44

173
62

--

222
106

28-100
67-100

l.18
l.08

44.4
35.3

III
III
III

I
II
II

1.05
1.05
l.08

18.0
22.0
30.7

II
III
II

III
III
II

.
0\
0\

--

-~~234
38
154
25-100
~h'(154
-30
87
34-100
35
149
-184
25-100
(*42 unclassified, **37 unclassified)

---------------------------------------------------------------------------------------------------------------------

�Table 1. Aerial census with a fixed-wing airplane on the Larimer County antelope study area, 1963-1964 (continued).
Month

Time

Census Conditions
Ground Light Air

Ante10Ee Counted
Bucks Does Fawns Total

Buck-Doe-Fawn
Ratio

Census Time
(hours)

Ave. Herd
Size

February-1964 Morning
Noon
Evening

III

III
No flight
No flight

II

38

159

--

197

24-100

1.13

32.8

Morning
Noon
Evening

III
III
III

I
I
III

I
II
II

64
52
52

123
139
85

---

187
191
137

52-100
37-100
61-100

1.25
1.03
1.22

8.5
14.7
9.8

II
II
I

60
52
52

88
53
87

----

148
105
139

68-100
98-100
60-100

1.25
1.05
1.30

4.1
4.8
3.5

III
III
III

64
27
11

59
56
20

37
39
16

160
122
47

108-100-63
48-lO0-70
55-100-80

1.15
1.08
1.20

5.9
8.1
5.9

I
II
I

64
40
39

90
116
123

82
72
81

~'(248
228
243

71-100-91
34-100-62
32-100-66

1.32
1.20
1.22

6.4
7.1
7.8

1.03
1.05

10.6
14.5

March

No flight

April
May

Morning
Noon
Evening

III
III
III

Morning
Noon
Evening

III
III
III

August
September

II
II
I
No flight

June
July

--

III
III
III
No flight

Morning
Noon
Evening

III
III
III

I
I
I

-k12

ANTELOPE HUNTING SEASON
No flight

October
November

Morning
Noon
Evening
Note:

No flight due to weather
~'c209
25-100
106
-26
I III
~h'c203
21-100-11
118
23
43
II
I
(*77 unclassified, ** 19 unclassified)
No other flights were made, the pilot and airplane were not available.
III
III

0\

"-.I

�Table 2.

Aerial census with a fixed-wing airplane on the Moffat County antelope study area, 1963-1964.

Month

Time

Census Conditions
Ground Light Air

Ante10Ee Counted
Bucks Does Fawns Total

June - 1963

Morning
Noon
Evening

III
III
III

I
I
I

I
II
II

106
28
43

162
78
157

67
36
52

335
115
252

65-100-41
28-100-12
27-100-33

1.30
1.72
1.82

8.1
5.2
5.0

July

Morning
Noon
Evening

III
III
III

I
II
I

I
II
I

61
75
91

207
71
139

176
40
102

444
186
332

29-100-85
106-100-56
65-100-74

1.02
0.83
0.90

7.8
6.9
6.1

August

Morning
Noon
Evening

III
III
III

II
II
I

I
II
II

105
69
106

256
173
273

193
108
213

554
350
592

41-100-75
40-100-62
39-100-78

1.18
0.88
0.97

11.1
8.1
7.4

September

Morning
Noon
Evening

III
III
III

ANTELOPE HUNTING SEASON
68
145
104
317
29
82
53
166
62
118
96
276

47-100-72
35-100-65
53-100-81

1&amp;10
0.93

4.9
5.7
5.8

October

Morning
Noon
Evening

III
III

II
I
II
III
No flight

59
28

236
138

116
50

411
25-100-49
~\-231
20-100-36
*15 Unc l.a ss,

0.88
0.72

12.8
21.0

November

Morning
Noon
Evening

III

I
No flight
No flight

I

80

330

--

410

24-100

1.00

22.8

December

Morning
Noon
Evening

II
II
II

I
I
III

I
I
I

67
78
27

311
328
97

----

378
406
124

22-100
24-100
28-100

0.92
0.95
0.75

47.2
67.7
31.0

I
I
I

76
78
93

244
263
257

---

--

320
341
350

31-100
30-100
36-100

0.80
0.83
0.75

80.0
85.3
87.5

January-1964
February

I
I
1

I
II
II

Buck-Doe-Fawn
Ratio

Census Time
(hours)

--

Ave. Herd
Size

No flights
Morning
Noon
Evening

I
I
I

I
I
I

----------------------------------------------------------------------------------------------------------------------

0'\

00

�Table 2.

Aerial census with a fixed-wing airplane on the Moffat County antelope study area, 1963-1964 (continued).

Month

Time

March - 1964

Morning
Noon
Evening

Census Conditions
Ground Light Air
I
I

I
I
No flight

April

No flights

May

No flights

June

No flights

July

Morning
Noon
Evening

III
III

I
I

AnteloEe Counted
Bucks Does Fawns Total
90
85

240
295

104
147
I
I
69
66
II
I
No flight due to weather

Buck-Doe-Fawn
Ratio

240
380

37-100
29-100

0.73
--

110.0
190.0

137
50

388
185

71-100-93
96-100-72

0.88
0.68

7.2
6.6

--

.

Morning
Noon
Evening

September

Morning
Noon
Evening

112-100-91
414
137
124
I
I
153
~\-251
126-100-88
51
58
I III
73
~\-69Unc Lass,
No flight due to weather
ANTELOPE HUNTING SEASON
53-100-61
206
96
59
I
I
III
51
139-100-113
26
81
23
II
I
III
32
66-100-63
42
153
67
II
III
III
44

October

Morning
Noon
Evening

III
III
III

II
II
I

Morning
Noon
Evening

I
I

I
I
No flight

November

December

III
III

75
53
74

199
134
203

Ave. Herd
Size

--

August

I
I
II

Census Time
(hours)

---

--

274
187
277

38-100
40-100
36-100

0.93
0.68

6.6
8.7

1.20
0.77
0.80

4.5
2.9
4.6

0.83
0.70
0.73

14.4
15.6
21.3

98.0
0.67
33-100
-196
147
I
49
39.5
0.57
158
--I
-(Note: A large group of approximately 125 antelope moved out of the study area)

-

No flights

-----------------------------------------------------------------------------------------------------------------------

0'\
\.0

�Table 2.

Aerial census with a fixed-wing airplane on the Moffat County antelope study area, 1963-1964 (continued)._
Census Conditions
Ground Light Air

Month

Time

January-1965

Morning
Noon
Evening

I

February

Morning
Noon
Evening

March

Morning
Noon
Evening

Ante10Ee Counted
Bucks Does Fawns Total
53

107

--

160

Buck-Doe-Fawn
Ratio

Census Time
(hours)

Ave. Herd
Size

50-100

0.57

160.0

III
No flight
No flight

III

I

I
No flight
No flight

I

40

120

--

160

30-100

0.63

160.0

III

I
No flight
No flight

I

71

83

--

154

86-100

0.83

51.3

-..j
0

�- 71 -

The Beaver and Cessnas had aSide-by-side
seating arrangement for the
pilot and observer.
In the Piper Cub the observer was seated behind
the pilot, thus the observer had poor forward visibility, but better
side-to-side VLSLon.
The Piper Cub was better for census work because
it could fly slower and was more maneuverable, whenever situations required these flight characteristics, such as circling large herds.
Helicopters had great maneuverability with excellent observer visibility. They were slow in cruise-flight and only had approximately a
2-l/2-hour fuel supply. Fixed-wing aircraft were affected to a greater
extent by winds and rough air. They had a faster cruising speed and
approximately a 5-l/2-hour fuel supply. Rental costs of the airplanes
was $20-25 per hour, the Hughes was $65 per hour, and the Bell was $80
per hour. The Hughes proved to be unsatisfactory due to its flight
limitations.
Six paired (helicopter: fixed-wing airplane) flights were made in July,
October, December, January, and February (Tables 3 and 4). Due to problems in scheduling a helicopter, these flights were generally not on
consecutive days. A 24% higher count was obtained with the helicopter
in July than the fixed-wing plane. The total counts in December to
February were nearly equal, averaging 3% variation between the helicopter
and fixed-,wing counts. The fixed-wing flight in October yielded a 23%
higher count than the helicopter count. Helicopter counts in all pairs
showed less variation between morning, noon, and evening counts; also,
there was less variation in the buck:doe:fawn ratios for the helicopter
counts than for the fixed-wing counts.
The helicopter appeared to be better than the fixed-wing aircraft for
counts when the antelope are scattered, or when more accurate sex and
age ratio counts are desired.
However, the expense of renting a helicopter is several times that for a fixed-wing airplane.
The total
counts made with a fixed-wing plane on the wintering herds were nearly
equal to the helicopter counts.

Variation

Due to Flight Direction

Eight paired flights were made during the months April through August
to test the effect of flight direction on the total number of animals
counted (Table 5). Higher total counts were obtained on the east-west
flights in six of the eight pairs. However, poorer light conditions
prevailed on the north-south flights during three of the six pairs, which
likely contributed to the lower counts. There were differences in the
sex and age ratios, but there did not appear to be a distinct pattern.
It would appear, from this small sample, that the direction of flight has
little effect on counting accuracy.

�- 72 -

Table 3. Aerial census conducted on the Larimer County study area comparing helicopter
and fixed-wing airplane counts.

Census Conditions
Ground Light Air

Ante10Ee Counted
Bucks Does Fawns Total

Buck-Doe-Fawn
Ratio

Census
Time
(hours)

Airplane

Month

Time

Fixed-wing

July

Morning
Noon
Evening

III
III
III

I
II
III

I
II
II

50
66
22

109
34
45

66
30
34

225
130
101

46-100-61
194-100-88
49-100-76

1.47
L07
1.23

Helicopter

July

Morning
Noon
Evening

III
III
III

I
II
II

I
II
II

92
44
63

120
68
99

76
45
57

288
157
219

77-100-63
65-100-66
64-100-58

2.12
2.00
2.03

Fixed-wing

Feb.

Morning
Noon
Evening

III
III
No flight
No flight

II

38

159

197

24-100

L13

Helicopter

Feb.

Morning
Noon
Evening

III
III
III

I
I
III

I
I
I

39
59
48

97
134
123

136
193
171

40-100
37-100
39-100

1.32
1.33
1.33

Fixed-wing

Mar.

Morning
Noon
Evening

III
III
III

I
I
III

I
II
II

64
52
52

123
139
85

187
191
137

52-100
37-100
61-100

1.25
1.03
1.22

Helicopter

Mar.

Morning
Noon
Evening

III
III
III

I
I
I

I
I
I

39
63
66

96
150
174

135
213
240

41-100
42-100
40-100

1.78
2.30
1.25

Fixed-wing

Nov.

Morning
Noon
Evening

No flight
III
I
III
I

Morning
Noon
Evening

II
II
No flight
No flight

Helicopter

Dec.

"

,

26
III
106
*209
25-100
II
43
ll8 23 **203
21-100-ll
(*77 Unclassified
**19 Unclassified)
I

99

189

288

52-100

L03
L05

�- 73 -

Table 4. Aerial census on the Moffat
fixed-wing airplane counts.

County study area comparing

helicopter

Census Conditions
Ground Light Air

Ante10ee Counted
Bucks Does Fawns Total

and

Buck-Doe-Fawn
Ratio

Census
Time
(hours)

Aircraft

Month

Time

Fixed-wing

July

Morning
Noon
Evening

III
III
III

I
II
I

I
II
I

61
75
91

207
71
139

176
40
102

444
186
332

29-100-85
106-100-56
65-100-74

1.02
0.83
0.90

Helicopter

July

Morning
Noon
Evening

III
III
III

I
I
I

I
I
I

157
·104
123

245
172
274

184
133
191

586
409
588

64-100-75
60-100-77
45-100-70

1020
1.50
1.50

Fixed-wing

Oct.

Ho'rn Lng
Noon
Evening

III
III
III

II
II
I

I
I
II

75
53
74

199
134
203

274
187
277

38-100
40-100
36-100

0.83
0.70
0.73

Helicopter

Oct.

Morning
Noon
Evening

III
III
III

I
I
I

I
I
I

61
52
50

98
95
96

214
208
204

62-100-56
55-100-64
52-100-60

1058

Fixed-wing

Dec.

Morning
Noon
Evening

II
II
II

I
I
III

I
I
I

67
78
27

311
328
97

378
406
124

22-100
24-100
28-100

0.92
0 95
0.75

Morning
Noon
Evening

III

89

151

70-100

1.35

Helicopter

Dec.

No flight due to weather
III
I
62
No flight due to weather

55
61
58

0

Fixed-wing

Jan.

Morning
Noon
Evening

I
III
III
53
107
No flight due to weather
No flight due to weather

160

50-100

0 57

Helicopter

Jan.

Morning
Noon
Evening

I
I
I

416
332
373

28-100
31-100
32-100

0 63
0.62
0.70

I
II
III

I
I
I

92
78
90

324
254
283

0

0

�- 74 -

Table 5. Aerial census on the study areas evaluating variability due to direction
flown.
Location
and Date

Direction
of Flight

Census Conditions
Ground Light Air

Ante10Ee Counted
Buck-Doe-Fawn
Bucks Does Fawns Total
Ratio

Moffat County
Aug. , 1965

Aug. , 1966

June, 1967

July, 1967

E-W

III

I

I

77

136

74

287

57-100-54

+'l-S

III

I

I

40

129

68

237

31-100-52

E-W

III

I

I

56

54

42

152

104-100-78

N-S

III

II

I

40

76

54

170

53-100-71

E-W

III

I

I

22

71

41

134

31-100-58

N-S

III

II

I

27

68

15

llO

40-100-22

E-W

III

I

I

23

68

53

144

33-100-77

N-S

III

I

I

18

58

52

128

31-100-89

E-W

III

III

I

32

75

107

43-100

N-S

III

III

I

29

68

97

42-100

E-W

III

I

I

63

168

231

38-100

N-S

III

III

II

14

49

63

29-100

E-W

III

I

II

79

177

68

324

45-100-37

N-S

III

I

I

106

173

77

356

61-100-45

E-W

III

II

I

89

183

135

407

49-100-74

N-S

III

III

I

84

177

122

383

47-100-69

Larimer County
May, 1965

April, 1966

Aug. , 1966

Aug. , 1967

�- 75 Sample Trends
Trends plotted from the number of antelope counted on the fall surveys in
the 60 percent and 40 percent samples resembled the trends plotted for the
totaL (100 percent) survey (Fig. 1 and 4). The trends plotted for the 20
percent sample showed greater variance.
This was true for both study areas.
Trends established for the 60 percent and 40 percent samples from the late
winter surveys on the Larimer County study area also closely resembled the
trend for the total count; while the 20 percent sample showed only slight
variations (Fig. 5). However, all samples taken from the late winter surveys on the Moffat County study area showed extreme variations.
Differences between the two areas were due to the distribution of the antelope
on the areas. Antelope on the Moffat County area were grouped into one to
three very large herds during the winter months, so that most of the antelope
were included in the samples or entirely omitted. Antelope on the Larimer
County study area were more evenly distributed in smaller herds, thus the
sample trends did not show the extreme deviation from the basic trend as
found on the other area.
Trends plotted for fawn:doe ratios in the 60 percent and 40 percent samples
closely resembled the trend for the total survey (Figs. 1 and 3). The trends
for the 20 percent sample varied considerably from the other trends. This
same pattern occurred for the buck:doe samples in the fall surveys on the
Moffat County area (Fig. 2). Only the 40 percent sample in the fall surveys on the Larimer County area resembled the trend for the total area.
Trends for all samples in the spring surveys on the Larimer County area
varied from the basic trend.
Approximately one-third of Colorado is occupied by antelope and censused
in its entirety each year. It would be a great money- and time-saver if
population trends could be established from samples and flights covering
100 percent of the area made only once every five years. With the reduction of census time required, flights could be made only under ideal conditions, therefore yielding more accurate information.
The data presented
here indicated trends in population numbers and fawn:doe ratios were reasonably accurate in the 60 and 40 percent samples.
This sampling technique
did not yield reliable information for buck:doe ratios.
Other factors,
such as herd distribution, had definite effect on the trends established
from samples
The data presented here are inadequate and inconclusive.
There is a need for detailed research on sampling techniques.
Larger study
areas should be used for such an investigation.
o

Ground Surveys
The road counts were extremely variable and unpredictable (Tables 6 and 7).
In general antelope were most easily seen in the morning or evening and when
the skies were clear
However, antelope near the road would often run when
the vehicle passed by, thus were not recorded on other counts during the same
day. It would appear that an observer could gather much better information
seeking out known antelope concentrations rather than remaining on established
road transects.
o

Prepared

by

/4~.v6".&lt;~
George D. Bear
Wildlife Researcher

�- 76 -

600

--100percen t
.--~ 60
"
40
"
• _ •• 20

450

II

~

w

co

~

:::&gt;
Z 300

150

200

v»

150

w

o
Ca
o
o
0::: 100

w
Q..

rJ)

Z
~

-c

u,

50

o~----~~~----~~
~~
1%3

1%4

1%5

~

1%6

~~_
1%7

YEAR
Fig. 1. Trends in total number of antelope counted and fawn:doe ratios
established for various sample sizes using the fall aerial survey data
from the Moffat County study area.

�- 77 -

,,---..

--

_II

r., InII

100 perc en t
60

II

40

"

20

"

1000

800
en

,
,
,
,
,
,
,

••
;

w

o
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,
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o

o
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w
e,

en
~

v
:::&gt;

lID

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400

,

,

,

,

200

OL-

~~~
"h:
_
1963

__~~

III _I;~

1964

YEAR

~

~

1965

1966

~~1967

Fig. 20 Trends in buck:doe ratios established for various sample sizes
using the fall aerial survey data from the Moffat County study area.

�-

78 -

(400) ,
~

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100

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w

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o
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(200) t.
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~

«

u;

20

0L-------~19~G~3~----~1~9~6~4------~19~6~5~----~1*9~6~6------il~9~67~
YEAR

Fig. 3. Trends in buck:doe and fawn:doe ratios established for various
sample sizes using the fall aerial survey data from the Larimer County
study area.

"
II

"

�- 79 -

100 percen t

"'--'--'--_,1/1,

60

"

•

40

"

20

"

.1,

--

•;r 1- ~,

400

300

200

100

O~ __~

•

~~;~\;_'
1963

~~~------~~--------~~~-~~--~~~I~-~
1964

196

1966

1967

YEAR
Figo 40 Trends in total number of antelope counted established
sample 'sizes using the fall aerial survey data from the Larimer
areao

for various
County study

�- 80 -

100

~~, -1_ ..-,
,,---~

80
~~

V)

0
£:I

~

~~

w

60

0
0
•....

~

~,

.,

,~

0:::
W

e,

40
V)

~

U

:::&gt;

co

20

"

,
,,,

"

.•,
,.

'4_."-

.,

,

~

20 -

100 percent
60
"
40
"

..f

,

/

#

-¢

V"#

,

"*

~~.

0

250

50

YEAR
Fig. 5. Trends in total number of antelope counted and buck:doe ratios
established for various sample sizes using the spring aerial survey data
from the Larimer County study area.

�Table 6.

Road counts on the Larimer County antelope study area, 1963-1965.
Ante10Ee Counted
Does
Fawns
Unc1ass.

Month

Time

Census Conditions
Light
Road
Ground

June 1963

Morning
Noon
Evening

III
III
III

III
II
III

II
II
II

---

Morning
Noon
Evening

III
III
III

I
I
II

I
I
I

1
1
4
1
7
2
No antelope observed

--

Morning
Noon
Evening

III
III
III

II
I
III

I
I
I

11

---

--

Morning
Noon
Evening

III
III
III

I
I
II

I
I
I

4
10
1

July

August

September

Bucks

4
4
-4

2

19
.

1

--

--

--

--

--

8

2

--

--

7

4

--

20
25
13

12
9
5

--

4

7
--

Total

Buck-Doe-Fawn
Ratio
0-100-25

5
4
6

50-100-0

8
10

25-100-25
14-100-29

38
4
11

58-100-42

43
44
19

25-100-60
40-100-36
8-100-38

-

0-100-60

00

t-'

October

No counts

November

Morning
Noon
Evening

III
III
III

December

Morning
Noon
Evening

III
II
I

I
I
I

9
5
2

No counts
III
III

I
I

I
I

No antelope observed
No antelope observed

January 1964 Morning
Noon
Evening

III
III
III

II
I
I

I
I
I

No antelope observed
No antelope observed
No antelope observed

February

No counts
I
II
III

I
I
I

5
26
-No antelope observed
No antelope observed

March

April

Morning
Noon
Evening

III
III
III

34
26
4

11
8
2

17
23
14

71
62
22

24-100-32
19-100-31
50-100-50

26

57

19-100-0

No counts

-------------.-------------------------------------------------------------------------------------------------------------

�Table 6.

Road counts on the Larimer County antelope study area, 1963-1965 (continued).

Month

Time

Census Conditions
Ground
Light
Road

May 1964

Morning
Noon
Evening

III
III
III

II
I
I

.II
II
II

Morning
Noon
Evening

III
III
III

I
I
I

I
I
I

2
No antelope observed
1
5
--

Morning
Noon
Evening

III
III
III

III
II
II

I
I
I

--

Morning
Noon
Evening

III
III
III

I
II
III

I
I
I

-- _.-

June

July

August

September
October

November

Bucks
4
--

7

Antelope Counted
Does
Fawns
Unc1ass.
8
3
10

1

9
3
6

-4

Total

Buck-Doe-Fawn
Ratio

5

1.3
3
22

70-100

--

3

50-100

--

6

20-100

8
10
10

0-100-40
900-100-0
150-100-0

29
5
14

37-100-44
0-100-25
80-100-40

---

--

--

-

1

5
1
2

2

1

--

--

--

5

16
4
5

7
1
2

---

3

50-100

00
N

No counts
Morning
Noon
Evening

III
III
III

III
III
II

II
II
II

5
2
7

19
10
13

10
8
5

--

Morning
Noon
Evening

II
III
III

I
I
I

II
II
I

5
--

24
9
39

15
3
12

II
I
I

I
I
I

December

No counts

January 1965 Morning
Noon
Evening

III
III
III

17

2

-4
No antelope observed
No antelope observed

--

34
20
25

26-100-53
20-100-80
54-100-38

-2

49
12
70

21-100-63
0-100-33
44-100-31

--

6

50-100-0

--

--

--------------------------------------------------------------------------------------------------------------------------

�Table 6.

Road counts on the Larimer County antelope study area, 1963-1965 (continued).
Census Conditions
Ground
Light
Road

Bucks

February 1965 Morning
Noon
Evening

III
III
III

III
II
II

I
I
I

No antelope observed
No antelope observed
----

March

III
III

I
I

I
I

No antelope observed
No antelope observed

Month

April

Time

Morning
Noon
Evening

No count
No counts

Anteloj2e Counted
Does
Fawns
Unclass.

5

Total

Buck-Doe-Fawn
Ratio

5
00

w

�Table 7.

Road counts on the Moffat County antelope study area, 1963-1964.
Ante10Ee Counted
Fawns
Unc1ass.

Total

Buck-Doe-Fawn
Ratio

121
74
106

32-100-28
33-100-43
18-100-21

170
73
136

65-100-75
36-100-19
47-100-59

212
65
110

43-100-59
42-100-35
36-100-72

2

29
15
29

44-100-31
63-100-25
58-100-67

4

--

20

60-100-40

10

6

36

11-100-55

No antelope observed
1
6
6
No antelope observed

--

13

17-100-100

III

I
II
II

I
I

II
II

No antelope observed
58

58

Month

Time

Census Conditions
Road
Ground
Light

Bucks

Does

June 1963

Morning
Noon
Evening

III
III
III

I
I
I

II
II
II

24
14
14

76
42
76

21
18
16

--

Morning
Noon
Evening

III
III
III

I
II
I

II
II
II

46
17
31

71
47
66

53
9
39

--

Morning
Noon
Evening

III
III
III

III

40
13
19

93
31
53

55
11
38

24
10

III

I
I
I

Morning
Noon
Evening

III
III
III

I
I
I

I
I
I

7
5
7

16
8
12

5
2
8

--

Morning
Noon
Evening

III
III
No counts
III
I

I

6

10

I

2

18

Morning
Noon
Evening

III
III
III

III

Morning
Noon

II
II

July

August

September

October

November

December

I

I

January 1964

No counts, roads closed by snow

February

No counts, roads closed by snow

March

No counts, roads closed by snow

--

--

--

--

---

--

-1

-------------------------------------------------------------------------------------------------------------------------

00

~

�Table 7.

Month

Road counts on the Moffat County antelope study area, 1963-1964 (continued).

Time

Census Conditions
Light
Road
Ground

April 1964

No counts

May

No counts

Bucks

Ante10ee Counted
Does
Fawns
Unc1ass.

Total

Buck-Doe-Fawn
Ratio

71
33
81

54-100-22
24-100-33
56-100-:06

77
167

32-100-94
80-100-53

Morning
Noon
Evening

III
III
III

1I
I
II

II
II
II

25
5
28

46
21
50

10
7
3

Morning
Noon
Evening

III
III
No count

I
I

I
I

11
56

34
70

32
37

August

Morning
Noon
Evening

III
III
No count

I
I

I
I

31
2

56
14

37
11

2
6

126
33

55-100-66
14-100-79

September

Hor nLng

III
No count
III

I

I

3

1

--

--

4

300-100-0

Noon
Evening

I

I

6

8

2

4

20

75-100-25

Morning
Noon
Evening

III
III
III

I
I
I

I
I
I

38
10
24

51
19
32

43
13
34

--

132
42
90

75-100-84
53-100-68
75-100-106

June

July

October

Note:

--

---

--

4

00
\.Jl

---

Additional counts could not be made due to snowfall on the area, which closed the roads.

��- 87 -

January,

1970

JOB FINAL REPORT

State of

COLORADO

Project No.

W-40-R-10

Antelope

Work Plan No.

1

Job No.

Job Title:

Techniques

Period Covered:

April 1, 1963 to April 30, 1969

Personnel:

for Determining

Investigations
13

Population

Trends

George D. Bear

ABSTRACT

Present census methods can yield accurate information for antelope population trends. There appears to be little need for complex formulas as used
in deer and elk population studies. Basic trend data for the study areas
is presented in Jobs 11 and 12 of this project.

��- 89 -

TECHNIQUES

FOR DETERMINING

POPULATION

TRENDS

George D. Bear

P ..S. OBJECTIVE
Determine

the most effective

technique

for determining

population

trends.

during a five-year

period

METHODS AND MATERIALS
Establish population trends for data collected
under Jobs 11 and 12.

RESULTS AND DISCUSSION
The basic trend of the populations on the study areas as related to hunter
harvest is presented in Job 11 (Herd Structure and Factors Affecting Herd
Structure) of this project.
Trends as related to 20, 40, and 60 percent
sample sizes is discussed in the report for Job 12 (Antelope Census) of
this project.
Complex population formulas as used in deer and elk population studies are not needed for antelope.
Application of Kelker1s and
Eberhardt1s formulas resulted in very unrealistic population estimates
(Segment 5 report for this project).
Antelope occupy open rangelands and
aerial surveys conducted under the proper conditions and at the proper
times can yield accurate information on population trends without adjustments by using formulas.

Prepared

by --::-,,-..:...~_;_v·'~f-_-...:4:.!::...-·.~~£.Z...·_~~
G~orge ~Bear
~
Wildlife Researcher

_

��January,

- 91 -

1970

JOB PROGRESS REPORT

State of

~C~OL~ORA~D~O~

Project No.

W-40-R-10

Antelope

Work Plan No.

2

Job No.

Job Title:

Investigations
2

Food Habits of Antelope

Period Covered:
Personnel:

_

May 1, 1968 to April 30, 1969

George D. Bear and Jerry Robinson

ABSTRACT

Analysis of individual antelope rumen samples has been completed.
Plant
species in each sample have been identified and weights determined.
Percent composition has been determined for each plant species occurring on
the vegetation surveys. These data need to be grouped according to the
seasons of the year for final analysis, and the vegetation survey data
related to the rumen analysis data. Graphs have been prepared showing
daily and seasonal variations in antelope feeding patterns.

��- 93 -

FOOD HABITS OF ANTELOPE
George D. Bear

P. S. OBJECTIVE
To determine

antelope

forage preferences

and use.

SEGMENT OBJECTIVES
1.

Complete

analysis

of rumen samples and vegetation

2.

Prepare manuscripts.

surveys.

METHODS AND MATERIALS
A detailed description of the procedures for collecting data was presented
in segment report W-40-R-9 for this job (Colorado Game Research Report,
January 1969).

DESCRIPTION
A detailed
job.

description

was presented

OF AREA
in segment report W-40-R-9

for this

RESULTS AND DISCUSSION
Rumen samples were taken from antelope systematically collected on study
areas in Moffat and Saguache counties from 1965 to 1967. Additional samples
were collected from antelope harvested on study areas by hunters.
The
rumen samples have been sorted, the individual plant species identified,
and the weight of each species determined for each rumen sample. Vegetation
surveys were conducted on the study areas during the collecting periods to
detennine the plants available to the antelope.
Percent composition has
been computed for each plant species on each survey.
These data still need
to be compiled into seasonal trends based on plant growing periods, and the
rumen sample data related to the vegetation survey data. Observations on
antelope feeding behavior have been compiled and graphs constructed showing
daily and seasonal variations in feeding patterns.
A manuscript for the
Journal of Wildlife Management is pending final analysis of the data.

Prepared by
George -if.iear
Assistant Wildlife

Researcher

��January,

- 95 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-40-R-10

Antelope

Work Plan No.

2

Job No.

Job Title:

Physiological

Period Covered:
Personnel:

Investigations
3

Studies

May 1, 1968 to April 30, 1969

George D. Bear

ABSTRACT

Physiological and morphological data were obtained from 58 pronghorn
antelope collected on two study areas from June, 1965 to April, 1968.
A manuscript titled "Seasonal trends in fat levels for pronghorn antelope
in Colorado" has been prepared to submit to the Journal of Mammology for
publication.
The remainder of the data will be prepared for publication
during the next work segment.

1970

��- 97 -

PHYSIOLOGICAL

STUDIES

George D. Bear

P. S. OBJECTIVE
To establish
and season.

physiological

and morphological

norms for antelope

by sex, age,

SEGMENT OBJECTIVE
Analyze

data collected

in segments

METHODS

7, 8, and 9 and prepare manuscripts.

AND MATERIALS

A detailed description of the procedures for collecting data was presented
in segment report W-40-R-7 for this job (Colorado Game Research Report,
January, 1967, Part 1).

DESCRIPTION

OF AREA

Detailed descriptions were presented in two previous progress reports for
this job; W-40-R-7 and W-40-R-8 (Colorado Game Research Report, January,
1968, Part 1).

RESULTS AND DISCUSSION
Physiological and morphological data were collected from 58 antelope collected on the study areas during the period from June, 1965 to April, 1968.
Only the data on body fat levels have been analyzed.
A manuscript titled
"Seasonal trends in fat levels for pronghorn antelope in Colorado" has been
prepared and submitted to the Journal of Mammalogy for publication.
This
paper discusses the seasonal trends of visceral fat, kidney fat index,
thoracic fat, rump fat, and bone marrow fat for buck and doe antelope.
The
remainder of the data collected in this job will be prepared for publication
during the next work segment.

Prepared by _--"'=~_-f.~~_--...:&lt;fJ,:::........' _~=:
..:.....
_=_
G~~.
Bear
Assistant Wildlife Researcher

��Jan~ary,

- 99 -

1970

JOB FINAL REPORT

State

No.

W-40-R-10

Antelope

Plan No.

2

Job No.

Project
Work

COLORADO

of

Investigations

5

Job Title

Evaluation of Structures Placed in Net Wire
Fences to Facilitate Antelope Movements

Period

May 1, 1967 to April

Covered:

Personnel:

George

30, 1969

D. Bear

ABSTRACT

Several types of crossing devices were constructed in net-wire fences and
their use by antelope evaluated under natural conditions on rangelands in
northwestern Colorado.
The following types of structures were evaluated:
Paul's pass, Antelope pass, Powder River pass, and a section of fence
measuring 32 inches in total height.
Antelope passes evaluated in 1967
measured four feet by four feet.
Antelope use was nearly doubled by placing
this pass in an artificial corner or offset.
Nineteen percent of the
domestic sheep that approached this pass crossed it, while 58 percent of
the antelope crossed it.
In 1968 the dimensions of these passes were
increased to six feet by six feet; antelope usage dropped to three percent,
and no sheep crossed these larger structures.
Neither antelope nor livestock crossed the Powder River passes.
Usage of Paul's pass by all classes
of animals was very low, less than five percent.
Adult and yearling antelope
readily crossed the 32-inch fence; and many fawns started crossing it by
September.
Observations in this study indicated that this type of fence
is adequate to contain livestock under normal conditions on the open range.

��- 101 -

EVALUATION OF STRUCTURES PLACED IN NET WIRE
FENCES TO FACILITATE ANTELOPE MOVEMENTS
George D. Bear

New concepts in range management practices have greatly increased the
number of fences bisecting rangelands occupied by pronghorn antelope.
Antelope readily crawl under barbed wire fences constructed on cattle ranges.
However, most net wire fences constructed on sheep ranges are effective
barriers to antelope movements.
Construction of numerous sheep-tight fences
approximately 42-44 inches high in northwestern Colorado during 1964 caused
concern among local sportsmen.
Antelope were seen walking along these
fences seeking new travel routes to formerly used waterholes and wintering
grounds.
Many animals became entangled in fences when attempting to force
their way under or to jump across.
Intensive studies on antelope and various
fence types has been in progress in Wyoming since 1963. Therefore, Wyoming
personnel were consulted on the problem before the Colorado Division of
Game, Fish and Parks and the Bureau of Land Management in Colorado began a
study to evaluate various types of structures placed in sheep-tight fences
to facilitate antelope movements.
The study was conducted in two localities in northwestern Colorado; near
Sunbeam along the Yampa River and near Great Divide along the Little Snake
River.
Both areas are characterized by rolling terrain and a shrub overstory (sagebrush, Artemisia tridentata and bitterbrush, Purshia tridentata).
Annual precipitation is approximately eight inches.
Antelope in these areas
make short migrations, approximately 10 miles, between summering and wintering grounds, and also group-up along the river and permanent waterholes
during the latter part of the summer.

P. S. OBJECTIVE
Evaluate structures installed in sheep-tight fences as a means
free movement of antelope through net wire fences.

for permitting

The following report was presented at the Western Association of Game and
Fish Commissioners; June 28, 1969. These data were also published in the
following:
- 1968 Proceedings of the Third Biennial
Workshop, pp. 15-22.

Antelope

States

- Colorado Game, Fish and Parks Division, Outdoor Facts,
Game Information Leaflet Number 71 (in print).

�- 102 METlJODS AND MATERIALS

Construction
Several types of crossing devices were constructed in net-wire fences and
evaluated under natural conditions on open rangelands.
Paul's pass was
designed by Paul Applegate of the Bureau of Land Management.
It was a
rectangular pen 50 feet across and 100 feet long constructed in the fenceline (Fig. 1). Periphery of the pass was constructed with a three-strand
barbed wire fence. The fence bisecting the center of the enclosed area
(on the same plane as the original fence) was 26-inch woven wire. The
theory for constructing this type of structure is that the outer barbed wire
fence will prohibit movement of cattle, but will allow the antelope to crawl
under.
Then the inner fence (26-inch net wire) will prohibit the movement
of sheep, but is of a minimum height to permit antelope to cross over it.
The other type of crossing devices resembled minature catt1eguards.
The
antelope pass was constructed of 1-inch steel bar. Due to the light construction of these passes, they were limited to 4 feet widths to prevent
vehicles from crossing over them. The first passes evaluated in 1967
measured 4 feet by 4 feet. The pass was set on a pair of railroad ties
which elevated it approximately 6 inches off the ground.
One-half of the
passes were merely inserted in the fenceline, the others were placed in
offsets constructed in the fence1ine (Fig. 2). This was done by placing
a 90 degree turn in the fence and running this leg of the fence out for
25 feet, then angling it back toward the original fence1ine.
An antelope
pass was then placed in the center of the 25-foot leg of the offset.
In
1968, the dimensions of the passes were increased to 6 feet by 6 feet, and
they were elevated to 16 inches off the ground. Also, all passes in 1968
were placed in offsets.
In addition to these newly installed structures, two "Powder River" passes
and a.section of 32-inch fence, which had been installed about one year in
advance, were included in the study. The Powder River passes were made by
cutting a standard cattleguard in half, thus giving a minature cattleguard
measuring 4 feet wide and 6 feet long. This structure was placed in the
fenceline over a two-foot deep pit. These passes were installed in the
32-inch fence. The fence was constructed with a 26-inch net wire placed
2 inches above the ground and a strand of barbed wire placed 4 inches over
the woven wire.
It paralleled the Yampa River for approximately 4 miles,
blocking the routes normally used by antelope which watered at the river
during the summer months.

Evaluation

of Structures

Movement of antelope and livestock around the structures and the 32-inch
fence was determined by track counts and by direct observations during the
period between May 1 and November 1, in both 1967 and 1968. The structures

�- 103 -

were built on graded or cleared areas, which facilitated the track counts.
Four track counts, with two-day intervals between counts, were made during
each month.
The tracks were erased with a garden rake following each
count. During dry periods water was hauled to the areas and the soil moistened by using a portable pump. This technique maintained a better tracking
surface.

RESULTS AND DISCUSSION
Antelope

Pass

The smaller antelope passes constructed in 1967 were a better type of
structure for permitting antelope across the fencelin~.
The percentage of
animals crossing this type of structure was nearly doubled by placing it in
an artificial corner or offset.
Of the antelope approaching the pass placed
in the offset, 58 percent crossed, while only 30 percent crossed the pass
placed in the straight fenceline (Table 1). Nineteen percent of the domestic
sheep crossed the pass in the offset, and eight percent crossed the pass
set in the straight fenceline.
No cattle crossed the antelope passes in the
fence line , and three percent (a calf and a yearling) crossed the passes
placed in an offset.
Modifications made on the antelope pass in 1968 greatly reduced the percenta.ge of animals crossing over the pass. The dimensions were increased
and the passes raised to 16 inches off the ground in an effort to reduce
the number of livestock crossing the passes.
Only one percent of the antelope crossed these new passes, and no livestock crossed.
Paul's Pass and Powder River Pass
Use of the Paul's and Powder River passes was very low during both years of
the study. No livestock or antelope were known to use the Powder River
passes.
Cattle and horses did not cross the Paul's passes.
In 1967, two
percent of the antelope crossed the fenceline using this type of structure,
and three percent crossed in 1968. One percent of the sheep crossed through
the Paul's passes during both years.
The percentage of antelope entering the
outer barbed wire fence, but not crossing the center fence, increased from
three percent in 1967 to 10 percent in 1968; while SO percent of the sheep
entered the outer exclosure in 1967 and 18 percent in 1968.
The 32-inch Fence
Adult and yearling antelope readily crossed the 32-inch fence. A total of
76 adult and yearling animals were observed to approach the fence; 74 of
these jumped across.
During July and August, the fawns would follow the
adults to the fence, then run frantically along the fence after the older
animals crossed, but would not attempt to cross.
On several occasions does
were observed to jump back and forth across the fence several times as if
trying to coax their fawns across.
In October, 42 percent of the fawns were
observed jumping across the fence (Table 2). Cattle were observed moving
along this fence on many occasions; none of them attempted to jump across.

�\"""
\

\

\ \
,

'

- 104 -

(f)

-

IL

\

(l)

-0

Cd)

&lt;..'~
o (])
O,;~

-~

.

(\.I

�- 105 Table 1. Antelope and livestock movement around and across the various types
of structures.

Type of Structure,
Year and Species

Number of
Attempts

Number
Crossing

Percent
Crossing

AnteloEe
1967
Antelope Pass in offset
Antelope Pass
Paul's Pass
Powder River Pass

66
30
100
206

38
39
2
0

58
30
2
0

1968
Antelope Pass in offset
Paul's Pass
Powder River Pass

200
250
60

1
7
0

1
3
0

SheeE
1967
Antelope Pass in offset
Antelope Pass
Paul's Pass
Powder River Pass

370
285
360
0

70
24
5
0

19
8
1
0

1968
Antelope Pass in offset
Paul's Pass
Powder River Pass

339
981
35

0
1
0

0
1
0

Cattle
1967
Antelope Pass in offset
Antelope Pass
Paul's Pass
Powder River Pass

74
53
138
11

2
0
0
0

3
0
0
0

1968
Antelope Pass in offset
Paul's Pass
Powder River Pass

202
135
123

0
0
0

0

4
8

0
0

0
0

0
0

Horses
1967 - No Observations
1968
Antelope Pass in offset
Paul's Pass

�- 106 -

Table 2.
Colorado.

Observations

of anteldpe

Number Crossing
Bucks Does Fawns

Date

crossing

the 32-inch

fence near Sunbeam,

Number not Crossing
Bucks Does Fawns

Percent Crossing
Does
Fawns
Bucks

July 28-30

7

10

0

1

0

12

88

100

0

August 26

1

11

0

0

0

6

100

100

0

September

17

0

1

0

0

0

0

October

15-20

6

38

15

0

1

20

100
100

97

42

Spi11et, et a1. (1967) concluded from their work in Wyoming that the following fence types appeared to offer t~e best possibilities for least interference with. antelope movements· and yet provide for sheep management:
(a) 32'-inch net wire fence; (b) 26-inch net and barbed wire 4 inches above
it; (c) standard cattleguards. They stated, "A fence 32 inches high is
apparently the maximum that most antelope will readily cross. Standard
cattleguards will satisfactorily permit the movement of adult and yearling
antelope if they are placed so that they can be readily located. A corner
location for cattleguards is more readily found and crossed by antelope" ..
Here in Colorado, we found that the use of artificial corners (offsets)
increased the use of the antelope passes.
However, our findings differ from
those of the Wyoming study in that antelope did not readily cross a 6-foot
cattleguard or antelope pass. Approximately 60 percent of the antelope
crossed a 4-foot pass, but only one percent crossed the passes when they
were increased to 6 feet. If adult animals are hesitant to cross these
structures, the fawns will be less likely to use them. The 4-foot passes
were not satisfactory for preventing livestock movements across them.
Spillet, et al. (1967) recommended the use of a fence not exceeding 32
inches in height.
Data from our study indicated that this type of fence
was acceptable for yearling and adult animals.
Fawns started crossing this
fence by October.
It would appear that fawns may readily cross this fence
by December when the winter migration starts.
It is believed that a 32-inch
fence on antelope ranges would provide less interference to antelope movements than a higher fence with antelope passes installed in it. Observations
in this study indicated that this type of fence is adequate to contain livestock under normal conditions on the open range. The study in Wyoming found
that the 32-inch fence was adequate for sheep managment.
Sundstrom (1968),
in his study on water consumption by antelope, found that fawns had started
taking water by August.
Ther~fore, water developments should be made in an
area prior to, or at the time of, fence construction to provide the fawns
with water until they learn to cross the fence.

�- 107 -

Paul's pass received very little use; however, it should be further evaluated.
There was a noticeable increase of activity around these passes the
second year.
The percentage of antelope entering the outer barbed wire
exclosure increased from three to ten percent.
Antelope may more readily
accept a structure varying from this basic design.
It should be tested in
fenced areas where it is not essential for antelope to migrate semi-annually,
thus permitting them to become accustomed to the structure.
At the present time, there does not appear to be a clear-cut solution for
getting antelope across livestock fences.
Apparently there are regional
differences in antelope accepting various types of crossing structures and
fences.
Rouse (1962) found antelope in Carter County, Montana, had learned
to jump fences and experienced no difficulty in clearing fences up to 45
inches in height.
Reports by Spillet, et al. (1967) and Zobell (1968) indicate antelope in central Wyoming more readily cross horizontal barriers
than antelope on the Colorado study area did. It presently appears that a
32-inch maximum height on net wire fences with proper water developments may
partially alleviate the problem in northwestern Colorado.

LITERATURE

Rouse, C. H.
Antelope

CITED

1962. Antelope and sheep fences.
Conference.
pp. 45-47.

Trans.

1962 Interstate

Spillet, J. J., J. B. Low, and D. Sill.
1967. Livestock fences -- how
they influence pronghorn antelope movements.
Utah State Univ. Agri.
Exper. Sta. Bull. No. 470.
79 pp.
Sundstrom, C. 1968. Water consumption
tion related to water in Wyoming's
States Workshop. pp. 39-46.
Zobell, R.
Trans.

Prepared

by pronghorn
Red Desert.

antelope and distribuProc. 3rd Antelope

1968. Field studies of antelope movements on fence ranges.
33rd North American ,:,rildlife
Conference. pp , 211-217.

by
George D .,CSear
Wildlife Researcher

��- 109 -

January, 1970

JOB PROGRESS REPORT

State of

COLORADO
.----~~~~~~-----------

Project No.

W-105-R-9

------------------------

Work Plan No.,
Job Title:

4

_

An Ecological Investigation of the
Cache la Poudre Deer Herd, Colorado

Job No.,

l~(_E_i~g_ht_h
__Y_e_a_r~)

Population Density and Structure

Period Covered:

March 1, 1968 to February 28, 1969

Personnel:

Allen E. Anderson

_

�- 110 -

POPULATION

DENSITY AND STRUCTURE

Allen

E. Anderson

p. S. OBJECTIVES
I.

Estimate
population density on each of five selected study locations
\
be Li.eved to be representative of the lower-winter, middle-winter,
upper-winter,
transitional, and summer ranges of the herd to: (a) relate density to changing environmental conditions (Work Plan 3, Jobs
1 and 3), and (b) elucidate density-elevational
relationships, particularly between years.
"

2.

Estimate sex and age structure (composition)
vide basic information on herd dynamics.

SEGMENT

of the population

OBJECTIVE

Tabulate, summarize, analyze, and interpret collected
literature, and prepare manuscripts
for publication.

RESULTS

to pro-

data, review

recent

AND DISCUSSION

Progress
Estimation of herd structure and population densities on selected areas was
completed prior to June 1, 1965. The manuscript "Frequency distributions
of
mule deer fecal group counts" (D. C. Bowden, A. E. Anderson, and Do E. Medin)
was tentatively accepted by the Journal of Wildlife Management for publication
in late 1969. A computer program was written and preliminary analyses of 33
site factor variables related to cumulative counts of mule deer fecal groups,
1961-65, were initiated.
Analyses of herd structure samples - time - weather
relationships
were underway as Segment 9 ended.

Future
Complete

analyses

Prepared

by:

during

Segment

albA/C', ~~?1./1
Allen E. Anderson
Wildlife Researcher

Plans

10 and begin preparation

of final manuscripts.

�January, 1970

- III -

JOB PROGRESS REPORT

State of

COLORADO
.------~~~~---------An Ecological Investigation of the

Project Noo

W__-_10_5_-_R_-_9

Work Plan No.__~~5---- __---------Job Title:

Job Noo

1 (Eighth Year)

----------------------------

Physical Characteristics

Period Covered:
Personnel: .

Cache la Poudre Deer Herd, Colorado

_

March 1, 1968 to February 28, 1969

Allen E. Anderson

�- 112 -

PHYSICAL

CHARACTERISTICS

Allen E. Anderson

Pg Sg OBJECTIVE
Measure relevant physiological and morphological characteristics of deer
collected seasonally so that: (a) the response of the herd to its environment can be more adequately interpreted, (b) establish "physiological norms",
and (c) provide basic morphological data as related to sex and age classes.

SEGMENT OBJECTIVE
Tabulate, summarize, analyze, and interpret collected data, review recent
literature, and prepare final report and publication manuscripts.

RESULTS AND DISCUSSION
Progress
Physiological and morphological studies on four deer collected each month,
1961-1965, were completed April 27, 1965. The number totaled 192 regular
collections plus 33 from supplementary sources. Compilation and tabulation
of the many thousands of data entries has been completed.
Card punching,
selection of appropriate statistical applications, computer programming,
interpretation, and final literature review remain to be completed.
A
manuscript, "Erythrocytes and leukocytes in a Colorado mule deer population"
(A. E. Anderson, D. E. Medin, D. C. Bowden) was completed and submitted to
the Journal of Wildlife Management.
Two manuscripts, "Efficient sectioning
of incisors for estimating ages of mule deer" (J. A. Erickson and W. A.
Seliger) and "Antler morphometry in a Colorado mule deer population" (A. E.
Anderson and D. E. Medin) were accepted by the Journal of Wildlife Management and will appear in the April, 1969, and July, 1969 issues, respectively.
The manuscript "Estimating age in mule deer - a comparison of methods"
(J. A. Erickson, A. E. Anderson, and D. E. Medin) was being revised for resubmission to the Journal of Wildlife Management as Segment 9 ended. The
manuscript "Antler phenology in a Colorado mule deer population" (A. E.
Anderson and D. Eo Medin) was submitted to the Journal of Mammalogy. About
1,500 relevant references were added to the project bibliographical files.
Future Plans
Blood and tissue chemistry data will be related to several environmental
variables.
Planning of analyses, selection of variables, and tabulation of
these data for transfer to computer punch cards will begin during the first
half of Segment 10. Similar activity relative to external body measurements
and all organ and gland data will occupy the second half of Segment 10. The
extensive review of pertinent literature will continue with more effort being
expended on unpublished material.

Prepared

?:~&gt;~

by ~
Allen E. Anderson
Wildlife Researcher

�January, 1970

- 113 -

JOB PROGRESS REPORT

State of

COLORADO
----------------------------

Project Noo

__-_l_0_5-_R_-_9
W

Work Plan No.

5

----~-----------------

Job Title:

An Ecological Investigation of the
Cache la Poudre Deer Herd, Colorado
Job NOo

2~(~E_i~g~h_t_h_Y_e_a_r~)

Reproductive Studies

Period Covered:
Personnel:

_

March 1, 1968 to February· 28, 1969

Allen E. Anderson

_

�- 114 -

REPRODUCTIVE

STUDIES

Allen E. Anderson

P. S. OBJECTIVE
Determine the reproductive pattern of the deer herd to provide data on:
(a) morphology of the reproductive organs as related to age and season,
(b) gross and net productivity between years, and (c) tentatively, the
relationship of productivity to measured factors of the environment
(Work Plan 3).

SEGMENT OBJECTIVE
Tabulate, summarize, analyze, and interpret collected data, review recent
literature, and prepare final report and publication manuscripts.

RESULTS AND DISCUSSION

Progress
Reproductive data gathering from collected deer was completed April 27,
1965
Ovarian collections from hunter killed deer were completed with the
1964 hunting season. Most of the data have been compiled and tabulated.
Selection of appropriate statistical applications, interpretations, and
final manuscript writing remain to be completed.
General job status is
that of mid-analysis.
The following thesis was completed; Markwald, R. R.
1968. Histological and histochemical study of testicular periodicity in
mule deer. M.S. Thesis. Colorado State Univ., Fort Collins.
165 p.
0

Future Plans
Place data on computer

Prepared

by

cards and initiate analyses

e~

~
Allen E. Anderson
Wildlife Researcher

o

�- ll5 -

January, 1970

JOB PROGRESS REPORT

State of

COLORADO
----------------------------

Project Noo

W__-_10_5_-_R_-_9

Work Plan No.
Job Title:

~5

_

An Ecological Investigation of the
Cache 1a Poudre Deer Herd Colorado
Job Noo

3 (Eighth Year)
--------~~--------~--------

Harvest Analysis

Period Covered:
Personnel:

_

March 1, 1968 to February 28, 1969

Allen E. Anderson

�- 116 -

HARVEST ANALYSIS
Allen E. Anderson

p. So OBJECTIVES
1.

Determine the age structure of the deer herd kill to provide estimates
of (a) net productivity (percent female yearlings in the kill), and
(b) the effects of hunting regulations.

2.

Locate the distribution of the kill by sub-unit to relate age and sex
structure of the kill to elevational levels and harvest intensity.

3.

Measure relevant physical characteristics of the kill to provide a
possible index of herd response to winter food quality.

RESULTS AND DISCUSSION

Progress
Check station sampling of the fall harvest was completed with the 1965
hunting season.
Collected data have been tabulated and statistical treatment has been completed.
Most of the literature has been reviewed.
Objective 3 has been met in the Work Plan 5, Job 1 Manuscript to be published
in the Journal of Wildlife Management in 1969 entitled: "Antler morphometry
in a Colorado mule deer population" (A. E. Anderson, D. E. Medin).

Prepared

by

r2do?V

e

U?412.'VJ4(Tlr./

Allen E. Anderson
Wildlife Researcher

�- 117 -

JOB PROGRESS REPORT

State of

COLORADO
--------------~------------

Project No

W_-_l_0_5_-_R_-_l_0_

o

Work Plan No.

4
----~----------------

Job Title:
Period Covered:
Personnel:

An Ecological Investigation of the
Cache la Poudre Deer Herd, Colorado
Job NOo

l

Population bensity and Structure
March 1, 1969 to February 28, 1970
Allen E. Anderson

(~N_i_n_t_h
__Y_ea
__r~)

_

�- 118 POPULATION

DENSITY

AND STRUCTURE

Allen E. Anderson

P. S. OBJECTIVES
1.

Estimate population density on each of five selected study locations
believed to be representative of the lower-winter, middle-winter,
upper-winter,transitional,
and summer ranges of the herd to:
(a) relate density to changing environmental conditions (Work Plan 3, Jobs 1
and 3), and (b) elucidate density-elevational
relationships, particularly

2.

between

years.

Estimate sex and age structure (composition)
vide basic information on herd dynamics.

SEGMENT

of the population

OBJECTIVE

Tabulate, summarize, analyze, and interpret collected
literature, and prepare manuscripts for publication.

RESULTS

to pro-

data,

review

recent

AND DISCUSSION
Progress

Estimation of herd structure and population densities on selected areas was
completed prior to June 1, 1965.
Analyses of herd structure samples-timeweather relationships were essentially completed as Segment 10 ended.
Publications:
Bowden, D. C., A. E. Anderson, and D. E. Medin.
1969.
Frequency distributions of mule deer fecal group counts.
J. Wildl. Mgmt.
33(4):895-905.
Manuscripts

in preparation:

Anderson, A. E., D. E. Medin, and D. C. Bowden.
1970. Mule deer fecal
group counts related to site factors on winter range.
Manuscripts

submitted:

None

Future

Plans

Complete and submit final manuscripts for the Journal
ment and the Journal of Range Management.

Prepared

by

E" {:;~r&lt;£[(!jz.4-Cyt..../
Allen E. Anderson
Wildlife Researcher

u--i:?-i!P-:&gt;(../

of Wildlife

Manage-

�- 119 -

JOB PROGRESS REPORT

State of

COLORADO
.------~~~~---------An Ecological Investigation of the
W_-_l_0_5_-~R---l-0----~--Cache la Poudre Deer Herd, Colorado

Project NOo
Work Plan No
Job Title:

5

e

Job NOo

l~(~N_i_n_t_h_Y_e_a_r_)~-----------

Physical Characteristics

-----------------------------------------------------------

Period Covered:
Personnel:

_

March 1, 1969 to February 28, 1970

Allen E. Anderson

��- 121 -

PHYSICAL CHARACTERISTICS
Allen E. Anderson

P. S. OBJECTIVE
Measure relevant physiological and morphological characteristics of deer
collected seasonally so that: (a) the response of the herd to its environment can be more adequately interpreted, (b) establish "physiological norms",
and (c) provide basic morphological data as related to sex and age classes.

SEGMENT OBJECTIVE
Tabulate, summarize, analyze, and interpret collected data, review recent
literature, and prepare final report and publication manuscripts.

RESULTS AND DISCUSSION
Progress
Physiological and morphological studies on four deer collected each month,
1961-1965, were completed April 27, 1965. The number totaled 192 regular
collections plus 33 from supplementary sources.
All data were placed on
punch-cards and preliminary analyses of blood and tissue chemistry data were
underway as the Segment ended. The review of literature was continued and
about 500 references were obtained for project files during Segment 10.
Publications:
Anderson, A. E., D. E. Medin, and D. P. Ochs.
1969. Relationships of carcass
fat indices in 18 wintering mule deer.
Proc. Annu. Conf. Western Assoc:
State Game and Fish Commissioners.
Erickson, J. A., and W. G. Seliger.
1969. Efficient sectioning of incisors
for estimating ages of mule deer. J. Wildl. Mgmt. 33(2):384-388.
Manuscripts

in preparation:

Anderson, A. E., and D. E. Medin.
1970. Vitamin A in the liver and blood
of mule deer. A.A.A.S. and New Mexico Acad. Science, 46th Annual
Meeting, Las Vegas, New Mexico.
April 22-25.
Anderson, A. E., D. E. Medin, and D. C. Bowden.
1970. Temporal fluctuations
in blood serum electrolytes and protein in a mule deer population.
J.
Wildl. Mgmt.
Anderson, A. E., D. E. Medin, and D. C. Bowden.
in adrenal ascorbic acid and adrenal weight
J. Mammalogy.

1970. Temporal variation
in a mule deer population.

�- 122 -

Anderson, A. E., D. E. Medin, and D. C. Bowden.
1970. Temporal variation
of vitamin A in the blood and liver of a mule deer population.
J.
Marrnnalogy.
Nicolls, K. E. 1970. Cytometry and volumetry of acidophil cells in the
hypophysis cerebri pars distalis of Colorado mule deer (odocoileus
hemionus hemionus) relative to seasons of the photo period and antler
cycles.
Z. fur Zellforsch. und Mikros. Anat.
Roughton,

R. D. 1970.

Manuscripts

Age structure

in browse populations.

Ecology.

submitted:

Anderson, A. E., and D. E. Medin.
1970. Antler phenology of a Colorado
mule deer population.
Southwestern Nat. 15(2 or 3). (Accepted).
Anderson, A. E., and D. E. Medin.
1970. Erythrocytes and leukocytes in
a Colorado mule deer population.
J. Wildl. Mgmt. 34(2).
(Accepted).
Erickson, J. E., A. E. Anderson, D. E. Medin, and D. C. Bowden.
1970.
Estimating age in mule deer-comparison of techniques.
J. Wildl. Mgmt.
34(3).
(Accepted).
Future Plans
Complete and submit final manuscripts

Prepared by

(;;-;.i.£~/c/ E ~,:.,
Allen E. Anderson
Wildlife Researcher

to appropriate

~.Jc-1-/

Journals.

�- 123 -

JOB PROGRESS REPORT

State of

COLORADO
.-------~~~~---------An Ecological Investigation

Project No.

W-105-R-10

-----------------------

of the
Cache la Poudre Deer Herd, Colorado

Job No .,

2 &lt;.;..N_i_n_t_h
Y_e_a_r.:-.)

Work Plan No.,

~-5-- __---__--------

Job Title:

Reproductive Studies

Period Covered:

March 1, 1969 to February 28, 1970

Personnel:

Allen E. Anderson

_

�- 124 -

REPRODUCTIVE

STUDIES

Allen E. Anderson

P. S. OBJECTIVE
Determine the reproductive pattern of the deer herd to provide data on:
(a) morphology of the reproductive organs as related to age and season,
(b) gross and net productivity between years, and (c) tentatively, the
relationship of productivity to measured factors of the environment
(Work Plan 3).

SEGMENT OBJECTIVE
Tabulate, summarize, analyze, and interpret collected data, review recent
literature, and prepare final report and publication manuscripts.

RESULTS AND DISCUSSION
Progress
Reproductive data gathering from collected deer was completed April 27,
1965. Ovarian collections from hunter killed deer were completed with the
1964 hunting season. Most of the data have been compiled and tabulated.
Publications:

None

Manuscripts

in preparation:

Manuscripts

submitted:

None

None
Future Plans

Place data on computer

cards and initiate analyses.

Prepared by
Allen E. Anderson
Wildlife Researcher

�- 125 -

JOB PROGRESS REPORT

COLORADO
---------------------------

State of

Project Noo
Work Plan Noo,
Job Title:

W__-_lO_5_-_R_-_l_0
~5----__-----------

_

An Ecological Investigation of the
Cache la Poudre Deer Herd. Colorado
Job Noo,

3~(~N~i~n_t~h~Y~e~a~r~)

Harvest Analysis

Period Covered:

March 1, 1969 to February' 28, 1970

Personnel:

Allen E. Anderson

_

�- 126 -

HARVEST ANALYSIS
Allen

E. Anderson

p. S. OBJECTIVES
1.

Determine the age structure of the deer herd kill to provide estimates
of (a) net productivity
(percent female yearlings in the kill), and
(b) the effects of hunting regulations.

2.

Locate the distribution of the kill by sub-unit to relate age and sex
structure of the kill to elevational levels and harvest intensityo

3

Measure relevant physical characteristics
possible index of herd response to winter

0

of the kill to provide
food quality.

a

RESULTS AND DISCUSSION

Progress
Check station sampling of the fall harvest was completed with the 1965
hunting season.
Collected data have been tabulated and statistical treatment has been completed.
Most of the literature has been reviewedo

Pub lications:
Anderson, Ao Eo, and D. Eo Medin.
1969. Antler morphometry in a
Colorado mule deer population.
J. Wildlo Mgmt.
33(3):520-533

0

Manuscripts

in preparation:

Manuscripts

submitted:

None

None

Future

Plans

Incorporate findings relative to Objectives 1 and 2 in a Department
Reporto
The first draft will be completed during Segment 110

Prepared

by ( ~7(/

E ~0~f'/!&lt;f-2-r.J

Allen E. Anderson
Wildlife Researcher

Special

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                  <text>April

1970

- 1 -

JOB PROGRESS

State

COLORADO

of
No.

W-37-R-23

Game Bird

Plan No.

1

Job No.

Project
Work

REPORT

Job Title:
Period

Summarization

Covered:

April

and Publication

1, 1969 through

Survey
12

of Pheasant

March

Research

Findings

31, 1970

P. S. OBJECTIVE
To publish

pheasant

management

and research

SEGMENT
1.
2.
3.
4.

findings

in Colorado.

OBJECTIVES

To assemble and compile all pheasant data pertinent to the manuscript.
To make a list of all photographs
that will appear in the manuscript.
To extract and compute all necessary file data for completing
tables
and figures.
To secure photographs for the manuscript
(not currently available)
0

METHODS
Methods

and materials

were

AND MATERIALS

described

RESULTS

in previous

segment

reports.

AND DISCUSSION

Wayne W. Sa nd f or t and Harold M. Swope, formerly connected with the job,
were assigned to new duties during the segment.
Therefore, no work was
accomplished
on the job.
Other W-37-R personnel will be assigned to
complete the work in Segment 24.

.:\(
Prepared

by

(-, 7

/.

~\~"~,~.
_'~L(~.~'~.1~(_'_·~.:_7~~._·&lt;_
..=~_J_'~~~'
..
~.
Howard D. Funk
Section Chief, Small Game Research

_

��April

1970

- 3 -

JOB FINAL

State

COLORADO

of
No.

W-37-R-23

Game Bird

Plan No.

1

Job No.

Project
Work

REPORT

Job Title
Period

Covered:

Personnel:

Pheasant

Hen Harvest

Survey

14

Investigation

April

1, 1969 through March 31, 1970

Warren

D. Snyder

ABSTRACT

Ring-necked
pheasant populations were censused along with collection of
environmental
and harvest data on two large study areas in Northeast
Colorado from 1963 through 1968.
The purpose of the study, experimental
hen harvest evaluation, was not enacted due to public and political
pressures.
However, much supplemental
information was obtained.
Analysis
showed that the birds-per-mile
index obtained during late-summer production
counts was a better indicator of population level and trend than the
previously used fall population index.
A new fall index, CY, multiplying
the crowing index times the young-per-hen,
was found to be equal to the
birds-per-mile
index in quality (r = .97**).
This would eliminate the
need for winter or spring sex ratio counts when hens are not included in
legal harvest.
A close relationship was found between the young-per-hen
index and the percentage of hens with broods (r = .95**).
Birds per mile
observed during spring sex ratio counts were least accurate of the population indices.
Existing sample sizes were found inadequate for evaluation
of proposed hen harvest.
Measures of hunting pressure, harvest and land
closure to hunting were obtained.
Both study areas were mapped by cover
type.
Approximately
80 percent of the land in both areas was devoted to
winter wheat production on a production-fallow
rotation basis.
Precipitation, vegetative growth and height, and spring plowing of wheat stubble
were compared with pheasant densities.
June 1 wheat height was the only
factor showing significant correlation with pheasant density (r = .67*).
Recommendations
for reenactment of the hen harvest study were provided
along with recommendations
for pheasant management census under existing
harvest conditions.

�-4 -

RECOMMENDATIONS

1. Successful hen harvest is a substitute for density dependent natural
mortality.
Yet little knowledge exists as to the proportion of mortality
that results from either density dependent or density independent influences.
The type of mortality, time of mortality, degree of annual turnover and many other unknowns exist.
Therefore, a well planned and conducted mortality study would provide invaluable knowledge upon which to
base hen harvest.
2. Harvest of hen pheasants in the future should also be based on a
thorough evaluation as outlined in the "Conclusions
and Recormnendations"
section of this report.
3. Under current "cocks only" harvest in Colorado, sex ratio counts
should be discontinued.
More brood count replicates should be completed.
4.
The fall population index is not an accurate measure of population
trend.
A new CY index should be substituted for it and tested with other
indices before submitting season recommendations.

�-

PHEASANT

5 -

HEN HARVEST

INVESTIGATION

Final Report with Recommendations
Reenactment

Warren

for Future

of the Study

D. Snyder

Acknowledgements
The writer is deeply indebted to Harold M. Swope, who designed and initiated
this study.
His able instruction, guidance and assistance resulted in
quality census data facilitating this analysis.
It is truly regrettable
that he was unable to carry this study to a successful completion.
Wayne W.
Sandfort, who provided much assistance and direction during initial phases
of the study, is to be commended.
Sincere thanks are expressed to Dr. David C. Bowden, consulting statistician,
who provided or assisted with much of the analysis presented here.
Several personnel were responsible for collecting field data. William E.
Martin, Michael R. Sterling, William E. Jones, J. Steven Lange, William
Carpenter, and Wayne Russell were among the primary participants.
Their
fine efforts are to be commended.
Gratitude is expressed to Wildlife
Conservation Officer Trainees and Research Personnel who provided temporary assistance.
Thanks are also expressed

to Howard D. Funk for editing

this manuscript.

P. S. OBJECTIVE
To determine how the inclusion of one hen in the bag limit will affect a
dryland pheasant population, its harvest, and its hunter recreation time.

METHODS

AND MATERIALS

Review of Literature
Library research produced little information relative to the effects of
removing females from upland game bird populations.
Several Western
States, known to allow hunter harvest of hen pheasants, were requested to
send all pertinent data.

�- 6 A questionnaire was prepared requesting answers to specific questions.
This was sent to all State and Provincial Conservation Agencies.
A final
synopsis was compiled from the data contained in the completed questionnaires and accompanying materials (Swope 1964a).

Selecting

and Defining

the Study Areas

Study areas (Fig. 1) were selected by a process of elimination based primarily on previous surveys conducted under the pheasant work plan. Areas
were compared on the basis of: (1) size, (2) pheasant populations, (3)
vegetative cover and land use practices, an.d (4) homogeneity within each
area. A central township was selected in each area for more intensive study.

Population

Census

Spring Sex Ratio Counts
Random roadside surveys were conducted during early morning and evening
periods beginning when harem activity neared a mid-April peak. Counts
were discontinued in early or mid-May when the proportion of hens declined
indicating the onset of nesting.
An average of approximately 577 pheasants
were tallied in 256 miles of counts each spring.
Harems were flushed when possible.
Counts usually were conducted during
periods of calm, fair weather; however, counts were occasionally conducted
during wet, drizzly weather.
During such periods birds remained in the
open because of the wet vegetation.
Twenty miles-per-hour was used as a
standard speed.

Crowing

Counts

The Kimball crowing-count method, as revised in Colorado in 1953 (Sandfort
1960a), was used on three ten-stop routes in each study area (Fig. 1).
If possible, at least three replicates were made per route. Stations were
approximately two miles apart.
Two minute counts were begun at 50 minutes
before sunrise and completed about ten minutes after sunrise.
Counts were
used when wind conditions were less than five miles per hour. The high
count per station was used to obtain a route average.
Route averages were
then combined to obtain a study area average.

Rooster

Crowing Observations

Crowing roosters were frequently observed at a distance on mornings when
wind prevented completion of a crowing census.
The frequency of crowing
was timed, recorded and harem sizes were determined.
A spotting scope
was used in these observations.

�7

study ~I
Areas
Colorado

EXPERIMENTAL

,,
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SCALE
-- .

o 1 2 3 4 5

••••••••••Brood Route

-._.- Crowing and
Brood Route
Central Study
Township

AREA

~ig. 1. Census routes and the central study township within each hen harvest

study area.

lOmi.

�- 8 -

Production

Counts

These counts were usually employed during August and into September.
Hens,
young and cocks were tallied on seven roadside routes in each area (Fig. 1).
Observed broods were flushed when possible during these early morning or
evening counts.
Age estimates were obtained on clearly observed broods.
Most counts were restricted to periods of little or no wind and fair weather.
Three or more replicates were usually obtained per route (Sandfort 1960b).

Hunting
Hunting

Pressure

and Harvest

Information

Surveys

Aerial hunting pressure checks were made on the opening weekend of season
each year. A pre-season flight a few days before the season was conducted
in 1968 to index the number of vehicles resident along the census routes.
Vehicles parked at farmyards, parked elsewhere, or moving along roads were
recorded along the 174 square mile survey route in each study area. Some
additional hunting pressure information was obtained during varied periods
in 1965, 1966, and 1968, while contacting hunters.

Hunting

Season Recommendations

Recommendations concerning season length and bag and possession
were submitted each year based on census information.

limits

Land Posting Surveys
Surveys to determine the amount of land unposted, closed to hunting, or
posted "Hunting by Permission Only" were conducted each year in the experimental area. These were also extended to the control in 1967 and 1968
(Snyder 1969a).

Hunting

Success

Field contacts were used to measure hunting success on the two study areas
through several weekends of the 1968 season.
Check stations were set up
during evening hours at approaches to the highway between the two study
areas on the first and second weekends of season.
Hours hunted and birdsper-hunter information was obtained.

Sampling

Environmental

Conditions

Cover Mapping
Aerial photos of study sections were located in the appropriate Soil
Conservation Service Office.
Details of each section (field boundaries,
roads, farm lots, etc.) were transposed from the photo to graph paper.

�- 9 Vegetative cover types were placed on the graph paper copy during field
inspections of the section.
Graph squares were tallied for each field or
cover type and converted to acres to obtain the acreage per section
(Snyder 1966).

Precipitation

Measurement

Precipitation gauges were mounted on steel posts in occupied farm lots
nearest the corners of the two study townships.
The eight farmers receiving these agreed to keep precipitation records throughout the year.
Each
gauge station recorder was given a clipboard and a supply of precipitation
record forms. Cooperators were contacted periodically to pick up precipitation data and leave additional forms.

Vegetative

Growth Measurements

Transects for measuring vegetative vigor were established in two sma11grain fields at each corner of the experimental and control townships.
These extended into the fields at 45 degree angles from either field edge.
Fifty vegetative height measurements, one pace apart, were made along each
transect periodically during the period of spring growth each year.
Final
measurements were completed late in summer to record stubble heights, and
residual weed growth following wheat harvest.

Photo Hubs
Twenty-six photos were taken annually during November to record roadside
and field conditions at thirteen photo hub sites.
Seven of these sites
were in the experimental area and six were in the control.

Progress of Spring Tillage of Wheat Stubble
Two roadside surveys of stubble fields were conducted in late April and
early May each year to determine the progress of initial spring ploWing.
The length or width of each field adjacent to the road traveled was measured by speedometer readings.
Percentages plowed, unplowed and partially
plowed were ascertained.

Non-measured

Environmental

Influences

Hailed areas were located and mapped.
In some instances, areas were searched
for evidence of pheasant mortality.
The six year period of study was generally free of severe blizzard conditions.
As a consequence, little winter
mortality surveillance was conducted.

�- 10 DESCRIPTION

OF AREA

This study was initiated in the extreme northeast corner of Colorado in
parts of Phillips, Sedgwick, Logan, and Yuma counties.
As initially
established, the experimental area contained approximately 580 square miles.
The experimental area was defined as: That area north of U. S. Highway 6,
east of State Highway 59, south of the South Platte River and west of the
Nebraska State Line. The control area contained 436 square miles of land,
to the south of U. S. Highway 6 in Phillips County, from Holyoke west, and
included parts of eastern Logan County and Northern Yuma County.
Both
areas lay within Colorado's best pheasant range.
Both areas were essentially High Plains Tableland, very flat and uniform
in character.
Elevations ranged from about 3,400 feet in the eastern part
to about 4,000 feet in the west.
The areas were sparsely populated.
Average farm size exceeded two square miles.
A close examination of crop
types, precipitation and other characteristics of the two areas follows
in the section on environment.

RESULTS

AND DISCUSSION

The primary objective of this study was not attained.
Due to public and
political pressures, the first proposed experimental hen harvest in 1963
was called off before it could be implemented.
As a consequence, this
report provides findings concerning environment, census and harvest based
on "cocks-only" harvest.
But in spite of this initial failure, the study
has provided a vast quantity of excellent information.
This information
will be invaluable in conducting a future experimental hen harvest.
In
addition, evaluation of the census data has provided improved census procedures for use in bettering pheasant management.
Environmental influences governing pheasant abundance are more clearly understood.

Review of Literature
Swope (1964a) summarized literature and information obtained and used
during initial phases of this study. Swope (1968) also compiled and
edited an additional quantity of literature concerning the basis for hen
harvest and a summarization of initial study results for Division personnel
information.
In addition, a large volume of literature was reviewed and
used in evaluating the study results.
Many of the references are listed
at the end of this report.

Selecting

and Defining

the Study Areas

Criteria for establishing the two study areas were listed in the preceding
section on methods.
As reported by Swope (1964b), the two areas selected
were the only ones available in Colorado which could meet the basic requirements.
Area boundaries were retained as initially established throughout
the six-year study.

�- 11 -

Within each study area, townships for intensive study were selected.
These townships were centrally located with large buffer zones surrounding
them.

Population
Definition

Census

of Terms

The following presentation involves three separate censuses: (1) the sex
ratio count, (2) the crowing count, and (3) the production or brood count.
Several indices and index combinations are summarized, analyzed, and discussed on the following pages.
To help the reader distinguish the various
components from one another, abbreviated symbols have been substituted in
a transitional manner.
The following table provides a reference to these
index symbols (Table 1).

Table 1. A list of index abbreviations,
in pheasant population census.

Abbrev.

their deriviation

Representation

and representation

Source of Derivation

C

Number of cocks

(1)

Spring crowing count

H

No. of hens per cock

(2)

Spring sex ratio count

Y

No. young per hen

(3)

Production

CH

Total hens in spring

Censuses

1 and 2 above

CHY

Total young

(based on hen pop.)

Censuses

1, 2 and 3 above

CY

Total young

(based on cock pop.)

Censuses

1 and 3

bpm

Birds-per-mile

in spring

(2)

Spring sex ratio count

BPM

Birds-per-mi1e

in fall

(3)

Production

count

SH

Percent hens observed with broods
(successful hens)

(3)

Production

count

spi

Spring population
(p = C + CH)

Censuses

1 and 2

FPI

Fall population index
(P
C + CH + CHY)

Censuses

1, 2 and 3

Correlation
Correlation

coefficient
coefficient

index

significantly
significantly

count

greater than 0 at 5 percent
greater than 0 at 1 percent

level.
level.

�- 12 Sex Ratio Counts
Two indices, hens-per-cock (H) and birds-per-mile
(bpm) were obtained from
this spring census.
David C. Bowden, Ph. D., provided an analysis of these
data. Sample sizes, sex ratios, standard errors, confidence intervals and
estimated sample size needed to be within 10% and 20% of the "true" sex
ratio at 95% confidence level are shown in Table 2. Sex ratios differed
by 10 percent, but not by 20 percent between the two study areas.
Question
arises as to what percent harvest would we have to attain to be able to
detect a significant change in sex ratio between the two areas.
Some
examples are included to illustrate.
Example

1.

l20~
12

100 cf
60

Post season number
Post season sex ratio

108
2.7

40
1

Fall pre-season sex ratio
harvest + 5% mort.
20/0 ~
and 60%
25%~

l20~

1006'

30

60

Post season number
Post season sex ratio

90
2.25

40
1

Fall pre-season

sex ratio

cf only season but 10% ~

Example

2.

mortality

d'

2.25 X 20%
2.25 + .45
The difference

between

.45
2.70
the two ratios is 20% of the lower.

Based on this information, we should use the sample size estimated needed
to predict within 0.1 of the ratio. Or, on the average about 20 to 25
counts.
That number is attainable and within reason.
New procedures outlined in the Conclusions and Recommendations section would probably reduce
the required sample size.
Table 3 summarizes and illustrates the birds-per-mile
(bpm) data for the
six year period.
Note that the estimated number of counts needed to attain
95% confidence within 0.1 of the ratio is beyond the capacity of present
census procedures.
Those at the 0.2 width of ratio would have to be used.
Again, pairing procedures outlined in the Conclusions and Recommendations
Section should increase the quality of samples.
I~oth the sex ratio (H) and birds-per-mile
(bpm) indices were analyzed on
an individual census count basis.
From the results in Tables 2 and 3, the
sex ratio information appears to be the more reliable of the two.

Crowing

Count Census

In evaluating the crowing indices (C) the high count per station was obtained
from available replicates.
High counts for the ten stations were then
averaged to obtain a route index. This procedure has been used in Colorado
pheasant census for a number 6f years (Sandfort 1960a).
It is preferrable

�- 13 -

to the average-per-station for two major reasons. First, pheasant crowing
intensity usually peaks during a certain period each spring (Swope 1967).
Second, weather conditions exert high variance on crowing counts. If
several replicates are obtained, the high count usually represents the
morning when weather and crowing intensity were both near optimum. Use
of the high count helps eliminate variance due to counts completed when
crowing intensity and weather were at different levels. Table 4 provides
a summary of route averages and the number of replicates obtained per route.

Table 2. The ratio of hens per cock, confidence interval and sample size
estimates based on spring sex ratio data.

Year

No. Rtes.
Sampled

Total
Birds

Hens/Cock

Std. Error
of Ratio

95% Confidence
Interval

SamE1e Size Estimate
at 0.2
at 0.1
Ratio
Ratio

Experimental Area
1963

14

792

1.82857

.13555

-.I

.29279

35.89

8.97

1964

9

582

2.23333

.23246

+

.53605

51.85

12.96

1965

11

411

2.23622

.05834

+

.12998

3.72

.93

1966

6

473

2.37857

.10461

+

.26895

7.67

1.92

1967

13

1038

2.78832

.13634

+

.29708

14.76

3.69

1968

11

725

3.09605

.23635

+

.52659

31.82

7.96

Control Area

1963

5

251

1.64211

.11907

+

.33054

20.26

5.06

1964

9

677

2.11982

.14429

+

.33273

22.17

5.54

1965

9

238

1.90244

.10lO7

+

.23307

13 .51

3.38

1966

5

372

2.32143

.15706

+

.43600

17.64

4.41

1967

12

590

2.18919

.13464

+

.29634

21.99

5.50

1968

12

778

3.29834

.16639

+

.36623

14.79

3.70

�- 14 Table 3. Pheasants observed per mile, confidence interval and sample size
estimates based on spring sex ratio data.

Year

No. Rtes.
Sampled

Total
Miles

Total
Birds

Birds/
Mile

St. Error
of Ratio

Confidence
Interval

Number Routes Needed
at 0.1
at 0.2
Ratio
Ratio

Experimental Area
1963

14

427.7

792

1.85410

.22358

+

.48293

94.98

23.74

1964

9

207.0

592

2.85990

.29749

+

.68601

51.78

12.95

1965

11

284.1

411

1.44667

.12610

+ 28095

41.49

10.37

1966

6

193.3

473

2.44697

.35167

+

.90414

81.91

20.48

1967

13

343.2

1038

3.02739

.41124

+

.89609

113.90

28.47

1968

11

245.0

725

3.22449

.28798

+

.64162

43.55

10.89

Control Area

1963

5

180.0

251

1.39444

.19642

+

.54526

76.45

19.11

1964

9

184.0

677

3.67935

.24676

+

.56903

20.83

5.21

1965

9

282.8

238

.84158

.12398

+

.28590

103.86

25.96

1966

5

179.9

372

2.06782

.53980

+ 1.49848

262.57

65.64

1967

12

293.9

590

2.00749

.21033

+

.46294

63.81

15.95

1968

12

253.8

778

3.20725

.36820

+

.81041

76.62

19.15

Analysis of variance was completed on the crowing count samples by Dr.
Bowden (Table 5). He did not find a significant interaction between years
and areas. But, area crowing means were widely different on the two areas
among years. To illustrate, C was 67.56 on the experimental area and 41.90
on the control in 1966. The difference was 25.66. In 1968, respective
indices of 36.27 and 37.23 on the two areas provided a difference of only 1.0.
If future hen harvest was detrimental to the experimental area population,
it must be detected by the years x areas interaction. Based on this analysis,
Dr. Bowden recommended a minimum of five crowing routes per area. These
would be needed to detect differences between areas when year effect was
considered.

�- 15 -

Table 4. The average rooster crowing rate per route and area during the six
year census period. l./

Year

East Route
Aver.
Replicate

Central Route
Aver.
Replicate

West Route
Aver.
Replicate

Area
Average

Experimental Area
1963

62.9

5

43.3

5

38.0

4

48.1

1964

71.6

5

75.3

4

64.7

4

70.5

1965

49.0

3

45.6

5

48.1

3

47.5

1966

78.8

4

73.8

3

50.1

3

67.6

1967

62.8

3

64.9

4

68.2

4

65.3

1968

41.0

2

40.3

2

30.4

2

37.2

Route X

61.0

57.2

49.9

56.0

Control Area
1963

42.0

5

50.8

4

42.2

5

45.0

1964

49.2

5

58.7

4

62.1

4

56.7

1965

25.0

3

23.9

4

29.4

4

26.1

1966

50.0

3

40.4

4

35.3

4

41.9

1967

71.0

3

57.3

3

36.7

4

55.0

1968

42.5

3

40.5

3

25.8

2

36.3

Route X

46.6

45.3

38.6

1/ High count per station data were used to compute route averages.

43.5

�Table 5.

Analysis of variance summary for crowing count data.

Source of Variation

D.F.

Sum of Squares

Total

359

125,972 •40000

Areas

1
4

Routes/Areas
Years

Mean Square

F

14,187.77778

14,187.7778~

9.4006 *

6,036.95556

1,509.2381

4.8080 *
/

5

40,260.10000

8,052.020o-P11.5817

*

Areas X Years

5

7,236.85556

1,447.3711

Not sign.

Routes X Years/Areas

20

13,904.71111

695.2356

Stations/(Routes/Areas)

54

16,950.66667

313.9012~

270

27,395.33333

101.4642

2.0818
I

Years X Stations/
(Routes/Areas)

I-'

3.0937 ***

0'\

�- 17 -

The analysis did find a signficant difference between areas (when years were
not considered), among years and among routes within areas (Table 5). A
decline in pheasant density from east to west seemed evident in both areas.
A significant difference among stations within routes was also noted by
the analysis.
This was probably due to the typical early morning pattern
of crowing intensity illustrated by Kimball (1949), Swope (1967), and others.
A peak in crowing activity was reached about 40 to 45 minutes before sunrise. After the peak, crowing activity gradually declined.
This trend
appeared quite evident in averaging samples from the six routes for the six
years (Fig. 2). The undefined peak possibly resulted from different starting times, different peaks of crowing intensity, or different pheasant
densities associated with stations along the six routes.

Influence

of Sex Ratio on Crowing

Intensity

Swope (1964b) found evidence that the number of hens within a rooster's
harem influenced his crowing rate. The fewer hens he possessed, the more
frequently he crowed (Fig. 3). When comparing crowing rates on the two
areas, equal sex ratios would not influence the comparison.
But, if hen
harvest materially changed the sex ratio on one area,then
the influence of
sex ratio would have to be considered when comparing crowing indices.
Sample sizes listed in Fig. 3 represent crowing intervals rather than
roosters.
Although a trend is evident, I do not consider the data to be
highly reliable.
A much larger sample is needed before making adjustments
in the crowing rate to compensate for sex ratio differences.
Gates (1966) was unable to test sex ratio influence on crowing rate as an
independent variable in his evaluation.
However, his analysis indicated
pheasant density was more influential than sex ratio.
Cocks crowed more
frequently in areas of higher density.
This variable should also be considered in future hen harvest research and evaluation.
It should be noted
that Gates study was conducted on an area possessing much lower rooster
densities than those in the Colorado study areas.

Production

Census-Brood

Counts

Sampling and Ana1ysis--These
counts, conducted from midsummer through late
summer were designed to measure production.
In a species, such as pheasant,
with high annual mortality and production, this census was the most important.
Two main indices were derived from the brood census data.
These were (1)
birds-per-mi1e
(BPM), and (2) young-per-hen
(Y). Table 6 presents the mean
annual ratios of these on the two study areas.
Fig. 4 shows the respective
BPM trends on the two areas through the six year study.

�- 18 -

50

40
w
~
m
~
00

~
.~
30
~
0
~

u

~
w
~
m

20

1/
77.910 95.1% 100

100

100

90.8%

85.1%

79.8%

76.8%

r~~__
~~__
~~__
~~~~__
~
1/ Percentage

o

100

1

2

3

4

Fig. 2. The average crowing
on 6 routes through 6 years.

of the average of stations

5

6
Station

7

8

3 through 6

9

10

intensity per station based on 133 counts

�-

19 -

6

,
,,
~'56
,
,

5

,
,
,

,
,
,,

,076
,
,,

4

,
3

,,
,
,

,

,
,,

130 (Sample size)
2

o

1

2

3 or more

Hens Per Rooster
Fig. 3
The apparent relationship between sex ratio and the crowing
frequency of roosters.
The line was fitted visually.
Sample size
refers to number of crowing intervals.

�- 20 -

Table 6. The mean annual young per hen and birds per mile ratios obtained
on the two hen harvest study areas.

Year

No.
Counts

Yg. /Hen

Std. Error
of Ratio

Confidence Birds
Interval Per Mile
95%

Std. Error
of Ratio

Confidence
Interval

EXEerimental Area
1963

19

4.17

.5832

1964

34

2.53

1965

28

1966

±

±

1.225

2.220

.2982

.2015

.4111

2.11

.2119

.4323

3.78

.3055

.6269

1.72

.2454

.5036

28

4.85

.1798

.3689

4.30

.3567

.7319

1967

28

4.62

.2786

.5717

3.65

.5236

1.0744

1968

14

5.22

.6818

1.472

2.27

.4970

.7962

1.23

.2320

±

0.627

+

- 1.074

Control Area
±

±

1963

20

3.65

.3804

1964

30

2.45

.3173

.6489

.75

.1598

.3268

1965

28

3.40

.6649

1.3644

.80

.1083

.2222

1966

27

5.56

.2696

.5543

2.95

.2920

.6004

1967

29

5.49

.2100

.4301

3.71

.4301

.8808

1968

15

5.75

.5998

1.2866

2.64

.4573

.9809

.4856

�- 21 -

5

4

,,
,

/
/
/
/

\

/

\

/

3

(

,
I

.•.....

::E:
0..

I

I

Q)

I

....•
rl

::E:
Q)

,,
,,
,

I

III

\..l

\

/

I

I

2

0..

J
I

s&gt;
\..l

Q)

Vl
.a

0
Vl

+'
~
Vl
10

Q)
.c 1

0..

o

63

64

65

66

67

Year
Fig. 4
The number of pheasants per mile observed during brood counts &lt;SPM)
on the two areas during the six year study.

68

�- 22 -

Dr. Bowden conducted analysis of variance on these ratios.
The results
are presented in Table 7.
There was no significant difference in either
index between areas.
Looking only at BPM the interaction between areas and
years was significant at the 5% significance level. However, the Y interaction of areas and years was not significant.
He concluded that if the
effect of hen harvest reached the magnitude of year effect brought on by
environmental influences, then it could be detected.
We would expect little
influence of hen harvest on the ratio of young per hen. But, BPM would be
affected if hen harvest lowered production.
Based on the analysis, Dr.
Bowden stated that about thirteen routes instead of the present seven would
be needed to estimate Y and BPM for an area within a particular year. Two
to four replicates were conducted per route, and this number was considered
in the analysis.

Period and Peak of Hatch--Brood age estimates were obtained each year to
detect possible differences in the period and peak of hatch.
Little discernable difference between the two areas was noted.
Therefore, information
from the two areas was combined and converted to percentages for year to
year comparison.
Fig. 5 indicates the earliest hatch occurred in 1963 with
the latest production peak in 1965. Some bias was introduced into the comparisons by differences in the period of observation (Table 8) among the
different years.
Early sampling would count first hatches more often, thus
skeWing the projected peak. The influence this had on the projections is
uncertain.
The exceptionally late hatch in 1965 probably was not fully
measured by BPM in relation to other years.
Comparison of harvest survey
data and the BPM index supported this hypothesis.
When interpreting Fig. 5,
the reader should assume that late summer production was at a higher level
than shown.
Some broods hatched even as late as September, after the majority of the census had been completed.

Midsummer Versus Late Summer Census--An attempt to determine the best
period for production census is shown in Table 9. Production BPM indices
were computed for bi-weekly periods and contrasted with average brood ages.
In 1963 the highest BPM index occurred in the last half of July.
In 1964
the first half of August produced the best count.
But during the last four
years of study, mid-September counts produced the highest indices.
Young
averaged about five weeks of age in 1964. During the other five years the
best counts were obtained when most birds were seven to nine weeks of age.
Table 9 by itself, does not yield any best bi-weekly interval for production
census.
The period and duration of hatch is one factor of influence.
Weather conditions, which vary from year to year, must be considered.
Although BPM indices were highest in mid-September from 1965 through 1969,
late August and early September census data were considered better in other
respects.
Older broods, especially those beyond nine weeks, gradually lost
ties with their hen and joined with other broods.
Age estimates, young per
hen and broodless hen figures were less accurate in late September.
Variance within and among counts became greater.
Klonglan (1955) obtained
similar findings.

�- 23 -

Table 7. Analysis of variance of experimental and control area young-per-hen
(Y) and birds-per-mi1e (BPM) information from 1964 through 1968. l/
Source of
Variation

d.f.

Sum of Squares

Mean Square

F

)
)
)

3,1231

)
)
)

3.1045 *

Birds-per-Mile (BPM)
Total

69

156.42015

Area

1

6.34368

6.34368

Route/Area

12

24.37489

2.03124

Year

4

77.70242

Area X Year

4

9.86551

2.46638

48

38.13365

.79445

Route X Year/Area

F1 ,12 at 90% = 3.18

at 95% = 4.75

F4 ,48 at 95% = 2.6

Young-per-Hen (Y)
Total

69

l39.29797

Area

1

2.52732

2.52732

Route/Area

12

10.16620

.84718

Year

4

83.92165

Area X Year

4

2.63014

.65753

Route X Year/Area

48

40.05266

.83443

F1 ,12 at 90% = 3.18

)
)
)

2.9332

)
)
)

.78797

F4 ,48 at 50% = .85

1/ 1963 data were omitted in these analyses due to differences in sample size.

�- 24 -

•

,\.

,

25

1',
I

"

I

I

20

m
0

.u
t::

.....,

I
I
I

4-1

I

,

I

10

I

Q)

I

Q)
p..,

I

.
.""

I

"

.
.
.~

5

.;I

~

\

\

\
\
\

./

,
,.

,
\
\

/

I

o
,...

\,

~,
.
,
I \
-.;j-.

I

:::r::

I

\

I

~ I

o
.u

,. .
'(,

~ I

15

\

\ I

I
I
I
"1

...c:

~

. .\

"""

.'"

\

\

" ,

" ,,

.

\

-

0
2
May

June

-

-- .. . .
.... -----

\.-

\

3
July

4

1

2
3
August

4

25

./\
•

eo/!
\,Q

\

•

:!it

20

\

/

\
\
\

15

\

\

...c:
o

\•....•..

.u

m

:::r::
4-1

\..

\

"

.

""'",\\

10

,.,

0

.u
t::
Q)

o
,...
Q)
p..,

\'
.

5

\

"

.•.•...

--...•'-

"

, .•.....
,,~.,...

0
1

Fig. 5.

2

3
May

4

1

2
3
June

4

The period and peak of production

1

2

3
July

as derived

4

1

2
3
August

from brood age estimates.

4

1

�= 25 Table 8.

The period of production

census during the six year study.

Period of Census
Year

Beginning

Date

Ending Date

1963

July 25 J)

September

3

1964

July 5

September

6

1965

August

3

October

1966

August

2

September

14

1967

August

1

September

29

1968

August

5

September

12

1

Jj Earlier counts were conducted but not included for analysis.

Very young broods were difficult to census.
Broods under four weeks of age
were difficult to see and to flush. The best census seemed to occur when
the majority of the broods were six to nine weeks old. Fairly reliable
counts of broods in this age interval were made, since hens could be
readily distinguished from young.
During the study, hatching usually peaked in late June or early July.
Adding six to nine weeks for the birds to reach the optimum age puts the
best census period from the second week in August through early September.
Management census information used in season recommendations
in Colorado
have consistently been collected in late July or early August.
In 1965,
especially, and in the following three years, counts and season recommendations were completed when conditions were only marginal for production
census.
Much better counts could have been obtained in late August and
early September.

Morning Versus Evening Counts--Both morning and evening production counts
were made.
Previous studies have shown better and more consistent counts
during morning census (Bennett and Henrickson 1938) and others.
Analysis
was completed to determine if this was true during this study.
Evaluation
of production census is partially summarized in Table 10. From 1964
through 1966 morning counts produced a higher BPM index.
Evening counts
provided a higher index in the remaining three years.
To measure variance,
morning and evening count totals were plotted on a frequency polygon (Fig. 6).
Successive counts on anyone
route were rotated between mornings and evening
as much as possible.
Thus, it was assumed that morning and evening counts

�- 26 -

were distributed equally among the fourteen routes on the two areas.
Fig.
6 represents 161 morning counts and 157 evening counts.
The lack of single
defined peaks in the figure was due to different population levels obtained
during the six years of study.
The occurrences indicate that, in this
study, distribution of evening counts was generally equal in range and
quality with morning counts;

Table 9.
intervals

A comparison of birds per mile
of production census.

July 4-17

July 18-31

1j

(BPM) and brood ages during biweekly

Aug. 1-14

Aug. 15-28

Aug. 29Sept. 11

1.92

1.84

1.71

1963 BPM

2.41

X Brood Age

6.81

8.24

9.35

11.2

Sept. 12-15

1964 BPM

0.77

1.41

2.14

1.26

1.36

X Brood Age

4.14

5.51

4.91

7.22

9.10

1965 BPM

0.82

1.33

1.29

1.84

X Brood Age

5.40

5.70

5.85

7.44

1966 BPM

2.68

3.91

3.83

5.31

X Brood Age

6.14

7.65

8.18

7.1

1967 BPM

2.64

3.87

3.13'1:..1

4.85

X Brood Age

5.89

6.59

8.53

9.17

1968 BPM

1.90

2.48

2.61

3.90 '1:..1

X Brood Age

6.35

7.38

9.09

9.45

II Underline BPM represents

the highest

II Sample size relatively small.

for the year.

']j

�- 27 -

Table

10.

A comparison

of morning

and evening pheasant

A. M.

production

counts.

Year

Miles

Birds

BPM

Miles

P. M.
Birds

BPM

1963

1019

1725

1.69

963

2098

2 .180'c

1964

629

945

1.5l')'c

911

1353

1.48

1965

787

1061

1.350'(

774

1010

1.31

1966

770

2999

3. 89~'c

693

2216

3.20

1967

795

2355

2.86

690

3100

4.49')'c

1968

529

1247

2.36

192

530

2.76·'"

4529

10335

2.28

4223

10307

-,"

The highest

2.44

of the two indices per year.

Population

Indices

Need for a better understanding of the sex ratio, crowing and brood censuses, and their indices led to the following analyses.
Simple linear
correlations were used almost exclusively in the following section.
Since
both areas received the same treatment (cocks only harvest, season length,
etc.) and census throughout the study, the information from the two
areas was used in unison here.
Thus, sample sizes were increased from six
to twelve in all correlations except those with harvest.
Similar correlations were conducted on Small Game Management Unit No. 1
(Fig. 7) for comparison purposes.
This unit contained about 3,400 square
miles in Northeast Colorado, including the two study areas.
Information
presented here was based on fourteen years of sampling on nine census
zones and routes (Fig. 7) (Sandfort 1960a, b). Unit 1 sample size exceeded
that for either study area (Table 11), but fewer brood count replicates
were usually run.

�- 28 -

24
22
20
18
Q)

.

(J

s:: 16

,,\,
, • - Evening Counts
• •
•
•• ••

,..,..
Q)

:::)

(J
(J

14

0
4-1
0

12

:&gt;-.

g 10
Q)

:l
0'

,..
Q)

8

•

,,

••...

Ji..

6

- Morning

Counts

,,

,,

4
2
0
0
N
I

0

0

-.:t

0

0

0
0

0
N
r-l

0
-.:t

0

0

\0
r-l

r-l

co

0
0

0
N
N

0

0

N

\0
N

-.:t

\0

co

I

I

I

I

r-l
N

I

r-l

r-l
\0

r-l

I

I

r-l

I

r-l

I

r-l

I

r-l

0

N
r-l

&lt;r

r-l
\0
r-l

I

co

r-l

I

-.:t

co

0

r-l
N
N

&lt;r

r-l

r-l

Pheasants

r-l

r-l

Observed

N

r-l

N

r-l
N

per Count

Fig. 6. A comparison of the range and distribution of 161 morning
157 evening production counts obtained during the six year study.

and

�- 29 -

-Qt'i
o
9
i
o

o
o

e
8(Jl
rT

8~
~:.J-.....-1J--~
:1)

•

•

::&gt;

---Akron

- - - -,""'yuma

t.J" - •• _ tL.~_~

g

-

.-. •

ay

KEY

---

aa -

Management Uhit Boundary
Census Route
Highway

Fig. 7.

Small Game Management Unit 1 and census routes within the unito

�- 30 Table 11. A comparison of sampling
Small Game Management Unit 1.

on the hen harvest

Sample Unit

Study Areas

study areas and

Management

Unit

Sex Ratio
No. of Routes

Random

Average Miles

256

918

Birds Tallied

577

6,312

Spring

Winter

No. of Routes

3

9

Average No. Counts

11

21

No. of Routes

7

9

Average Miles

630

834

Birds Tallied

1,508

1,789

Period of Census

Crowing

9 Zones

Count

Brood Count

Correlation

of Indices

Several different indices were available from the sex ratio, crowing and
production censuses.
Kimball (1949) described two that were incorporated
into Colorado pheasant census procedure.
These, the spring population
index (spi) and the fall population index (FPI), were represented by respective formulas:
p = C + CR, and P = C + CR + CRY.
The spring population index (spi) represented breeder density, while the
FPI measured fall density. The bpm and BPM observations, respectively,
obtained during sex ratio and production counts provided second measures
of pheasant abundance during spring and fall periods.
In other words, spi
and bpm were independent, directly comparable indices of spring density on
a year by year basis.
The FPI and BPM were independent, directly comparable
indices of fall density.
In addition, harvest surveys provide a third
independent measure of fall density.

�- 31 -

I)reeding Population Indices--In initial analysis of spring data, spi and
bpm were plotted on a scatter diagram to observe their relationship.
Analysis revealed a significant.correlation
coefficient of .65* (Fig. 8).
This
fair relationship provided iittle confidence in the quality of either index.
One index or the other could approximate actual population trend or both
could be equally erroneous.

Fall Indices--The FPI and BI~ (Fig. 9) were compared next.
A coefficient
of .90** indicated a more significant relationship between these two indices.
Again, the coefficient does not compare these indices with unknown fall
population trends.
It only compares the two with each other.
But as the
variance between two or more independent samples decreases, our confidence
in their accuracy increases.
To illustrate, 9, 10, and 11, and 4, 9, and
15 might represent two respective samples of an actual population.
We
would place higher confidence in the low variance group, 9, 10, and 11,
than in the other when predicting the population mean.
In the same manner,
high correlation coefficients
indicate low variance between samples on a
year to year basis.

Fall Index-Harvest Relationships--Samples
of hunter success and harvest
provided additional measures of fall population level and trend.
They
were compared here with the fall indices by correlation analysis to indicate
whether B~ or FPI was the better predictor of fall population
level.
Harvest data were not obtained on a study area basis except in 1968.
Therefore, Colorado Small Game Hunter Harvest Survey data from Sedgwick, Phillips,
Yuma and Logan counties were combined and used (Grieb and Hunter 1964, 1965,
1966,1967),
(Grieb 1968), (Funk and Grieb 1969).
The two study areas lie
within these four counties.
BPM and FPI indices from the two study areas were respectively combined and
averaged.
As a result, sample sizes were reduced to six.
These were compared by linear correlation with two measures of harvest: (1) total harvest,
and (2) birds per hunter.
As shown in Table 12, B~ correlated much more
closely with both harvest indices than FPI.
This provided evidence that
B~ was the more accurate fall index on the study areas.
To supplement study area results, BPM and FPI from Management Unit 1 were
tested against the same four-county harvest figures.
Sample sizes from
this unit included 13 years of harvest data and fall indices from nine sex
ratio, crowing and brood routes (Fig. 7). Correlation
results are shown
in Table 12. Birds-per-hunter
samples more closely compared with B~
than
with FPI.
But little difference was detected when FPI was tested against
the two harvest indices.
In interpreting the data it should be noted that
the B~-FPI
correlation coefficient
for the Management Unit was only .68,''',
(Table 13): much lower than the .90,',,', coefficient
for the B~-FPI
relationship on the two study areas.
Brood samples usually were not as intensive
nor conducted during the optimum census period in the Management Unit.
Thus,
the BPM data obtained there were not considered as accurate as those representing the study areas.

�- 32 -

0

250
230

::c
u

0 0

210

+

u

190

H

0.

~

H

150

c

0
oM

.jJ
Aj

0

130

r-I

0

0

::l

0.
0

a.

0

0

'0

s:::

0

0

170

110

0'1

~

'0
(IJ
(IJ

1.1

90

rn

70

0

Fig. 8 The relationship between the spring population index and
pheasants per mile observed during spring sex ratio counts.

�- 33 -

o

o

o

o
o
o

G

o

r

.90

lIS

,
I

2

4

3

Pheasants Observed Per Mile

5

(BPM)

Fig. 9. The relationship between the fall population
per mile observed during brood counts.

index and pheasants

�- 34 -

Table 12. Correlation coefficients relating harvest figures to fall
indices of pheasant populations on the study areas and Small Game Management Unit 1. 1/

Study Area

1/

Index

BPM

FPI

Management
BPM

Unit
FPI

Total Harvest

0.89

0.66

0.70

0.70

Birds/Hunter

0.85

0.66

0.71

0.46

Harvest

1/ Phillips, Sedgwick, Yuma and Logan County Small Game Hunter Harvest
Survey data were used in all correlations.
1/ Indices from the two study areas were combined reducing the sample size
to six. Management Unit sample size was thirteen.

Additional data support the hypothesis that BPM was more accurate than FPI
as an indicator of population level and trend on the study areas.
EPM
involves only the relationship between birds and miles based on a rather
large sample obtained after production was essentially complete.
In contrast,
the FPI was based on three separate censuses.
Errors inherent in anyone
of the three would be multiplied in obtaining the end result.
Hen mortality
between spring and production census was not considered.
The FPI also
assumed that hens with and without broods were observed in direct relation
to their number.
It was concluded from these factors that the study area BPM was the better
estimator of population level and trend. This index was then used as a base
for testing the various components of the FPI.

Density and Non-density Factors--When reviewing this analysis, it should be
understood that H, the number of hens per cock, by itself does not measure
density.
It must first be combined with C. Thus, CH represents hen density.
The same is true for Y. It is an index of production, but must be used in
conjunction with C or CH to represent production density.
Of the three
basic components, C, H, and Y, only C represents density.

Correlation of FPI and its Components with BPM--Regression analysis by
Bowden (Table 13) provided comparisons of all basic study area FPI components
with BPM. Cock populations (C) were not significantly correlated with EPN.
However, hen densities (CH) were (r -= .65')',). Production measured by (Y),
independent of dens ity, was highly related to BPM (r = .70')'''',). This provided
initial evidence of the importance of production in determining fall density.

�- 35 -

Table 13. Relationships
per-mile (BPM) expressed

of several population indices with fall birdsas .corre Lat Lon coefficients.

Variable
CH

Correlated with BPM
CY
CRY
C+CH

Location

C

H

Y

Study Areas
n = 12

.48

.40

.70~b'~.65"'~

•97"k"k

Unit 1
n = 14

.57-k

-.12

.79,'d&lt;.42

•75"'~"'~•7l,'d&lt;

~'~ Correlation
~b'~ Correlation

significantly
significantly

greater
greater

. 91-k-k

than 0 at 5 percent
than 0 at 1 percent

C+CH+cHY

.62"~

.90.,'d~

.48

.68,'d~

level.
level.

Cock densities (C) and hen densities (CH) were individually combined with Y
to relate production to spring breeder densities.
The respective CY and
CHY formulas, representing production denSity, were compared with BPM
(Fig. 10 and Fig. 11). The CY index showed highly significant correlation
with BPM (r = .97.,'d~)
(Table 13). The CHY-BPM correlation coefficient was
.91.,''':',
only slightly above the original FPI-BPM correlation where r = .90.,'~"'~.
This analysis indicated cock densities were better than hen densities for
estimating fall populations.
Note, however, that adding breeders to young,
as in the FPI formula (C + CH + CRY) did not result in a higher correlation
with BPM. Apparently breeder populations did not add to the accuracy of
fall density estimates except through their role in determining production.
Management Unit 1 data (Table 13) supported study area findings.
Although
the coefficients were much lower, the same trend in fall density comparisons
was evident.
Respective correlation coefficients obtained, when BPM was
plotted against FPI, CRY, and CY, equalled .68, .71, and .75. Again the
simple CY index showed the best relationship with BPM.
Tests were not conducted to determine if CY-BPM correlation coefficients
were significantly higher than those obtained using CHY or FPI. Sample
sizes were not considered adequate to permit findings of significant differences, even if they did exist.
However, the hypothesis that CY exceeded
CHY and FPI as a more accurate measure of fall density can still be made.
The similar trend of the coefficients in both the study area and management
unit samples supports this hypothesis.
To briefly summarize, comparisons of study area BPM and FPI with harvest
survey data indicated BPM was the more accurate of the two. It, in turn,
was used to compare components of FPI. Of these, the CY index, basing
production on spring cock densities, was most closely related to BPM. The
CHY index, measuring production based on spring hen densities was equal,
if not slightly better than the original FPI formula.
Less confidence
could be placed in Management Unit 1 results due to greater variances and
lower coefficients among samples, but management results supported study
area findings.

�- 36 -

350

300

250
.•....

iJ
c
(!)

::c

200

~
Q.,
tJl

§
0

)04

x 150
(!)

+'
ro
P::

tJl

.;j
~
0

Sol

100

0

u

L

0

I

1
Pheasants

2
Observed

•4

3
Per Mile

5

(BPM)

Fig. 10 Correlation of the CY index with pheasants
(BPM) during brood counts.

observed

per mile

�- 37 -

900

o
800

o

700

o

600

••.....
1'C

o

500

u

c 400
&lt;Il

::c

L.!

&lt;Il
0.
0'1

c

;::I
0

0

300

&gt;t
&gt;&lt;
II)

c
&lt;Il

::c

200
T' - .91

100
~

o

1

Pheasants

2

Observed

4

3

Per Mile

5

&lt;BPM)

Fig. 11. Correlation of the CHY index with pheasants
mile &lt;BPM) during brood counts.

observed

per

�- 38 -

The Basis for the CY Index--One important question remains unanswered.
Why did the BPM index show such close correlation with the CY index when
the latter doesn't consider hen density?
Why was the relationship better
than with CRY or FPI? Differential spring to summer mortality between the
sexes may provide the answer.
Hen densities observed during production
counts appear slightly better related to the crowing index (r = .75**)
than to spring hen populations (r = •731d:).
Rooster populations were
reduced in number by hunting before the crowing count was taken. Hens,
which survived harvest and winter in greater numbers, were subject to the
major, but variable spring stress of egg laying, nesting, renesting, and
mortality due to stubble plowing and crop harvest.
Dahlgren (1963) found
evidence that highest annuai pheasant mortality occurred from spring to
summer.
Hens undoubtedly were more subject to this mortality than roosters.
Thus, the reduced hen population surviving after spring more closely approximated C than CH. But this may not be the correct explanation.
The basis
for the CY index may depend on other variables too, which are not yet
discovered.
While C represents the breeding population, Y in turn, represents the capacity of that population to reproduce.
Linder, et al. (1960) found that the
number of hens successful in bringing off a brood was the primary factor
determining the rate of chick production.
Clutch size, fertility and
hatchability were not important variables in production.
The same condition
was evident in the study area analysis.
Successful hens (SH) or the observed
proportion with young, were plotted against Y (Fig. 12). A close positive
correlation resulted (r = .95**). Nesting success (SH) was the primary
factor determining production.
When C and Y were individually compared with BPM, Y showed a much closer
relationship (r = .70**). The C-BPM correlation coefficient was only .48
(Table 13). When the C and Y correlation coefficients were individually
squared, C accounted for only 23 percent of BPM variablity.
In contrast,
49 percent of the variability was attributed to Y or production.
We can
conclude that nesting success, which for the most part determined production,
was the primary factor determining fall populations.
Environmental conditions which governed nesting success were of major importance.
The above explanation leads to a major consideration in pheasant census.
Where hens are not harvested and percentage rooster harvest is rather consistent from year to year, winter and/or spring sex ratio counts can be
eliminated.
This is in contrast to previously published statements by
Dale (1952), Wagner et al. (1965), and others who have stressed that sex
ratios must be considered in determining subsequent fall density levels.
However, in some states where a high percentage of cocks are harvested,
CH might serve as a better base than C. More testing is needed under a
wide range of conditions.

The Influence of Density on Production--Linder,
et al. (1960), Allen (1956),
and Wagner and Stokes (1968), all showed evidence of density dependent
influence on pheasant reproduction.
In other words, fewer young per hen

�- 39 were produced when breeding densities were high, and more young were produced per hen when densities were low. In this study, an inverse correlation
of -0.25 between C and Y produced little evidence of density dependent
influence on production.
No relationship between hen densities (CH) and
young per hen was found (r = .07). Percent successful hens also showed no
significant relationship with spi (r = -.22) or with C (r = -.27) However,
if we are to conclude that hens can be successfully harvested without detriment to reproduction, density dependent factors must be present at some
time of the year.

Winter Survival--Wagner
et al. (1965), Linder et al. (1960) and Wagner and
Stokes (1968) found that spring densities were directly proportional in
size to those present the previous fall. Or, density dependent environmental
factors were not influential or highly limiting to wintering populations.
Similar comparisons in this study were not as conclusive.
Young per mile
observed during production counts were plotted against the spring population
index of the following year. The resultant coefficient (r = .56) did not
show significant evidence of a direct relationship for the five year interval. A better relationship was found using management unit data for a
thirteen year period.
In this test, where CY was plotted against C of the
following spring, the correlation coefficient (r = .79**) was highly significant.
These tests hint that winter density dependent factors were not of
major importance in Northeast Colorado, but do not rule out the possibility
of their existence.

A Test of Spring bpm Quality--Earlier
in the text the spring population
indices (spi) were tested against spring birds-per-mile
(bpm) (Fig. 8).
In the test presented here, bpm were substituted for spi, combined with Y,
and plotted against BPM (Fig. 13). The result shows only a fair relationship (r = .58*). Previous analyses indicated both BPM and Y were highly
accurate indices.
If we accept this as true, we have further evicence that
bpm were not accurate spring measures of density.

Hunting

and Harvest

Information

Several measurements, important to management and reinactment of the study,
were obtained during the study. However, because the 1963 hen season was
cancelled, many important facets of this phase of study could not be completed.

Aerial Hunting

Pressure Surveys

In 1963, Swope established aerial survey routes to record changes in hunting
pressure.
Flights were conducted on the opening Saturday afternoon of
season each year except in 1968, when weather conditions delayed the flight
one day. Table 14 summarizes the findings.
The 1968 counts could not be
compared directly with those of previous years.
They were conducted under

�- 40 -

adverse weather conditions at a time when many hunters were starting their
return trip to Denver.
The higher proportion of vehicles in the control
may partially be a result of this movement toward home.

Table 14. Vehicles observed
pressure on the experimental

on aerial transects as indices of hunting
and control study areas.

Percent
Change

Non-local
Vehic1es11

Percent
Change

Observed
Control

Total

197

199

396

1964

221

184

405

+ 2.5

178

+ 5.3

1965

207

196

403

- 0.5

176

- 1.0

1966

265

291

556

+38.0

329

-1-

1967

327

264

591

+ 6.3

364

+10.6

1968'1:./165

247

412

Year

Vehicles
Exp.

1963

169

8.7

185

11 Based

on a 1968 preseason count of 121 and 106 resident vehicles respectively, in the experiment and control areas.
11 Census conducted 2nd day of season. Not believed directly comparable
with data from preceding years.

Aerial flights were based on a large sample, 174 square miles in each
area.
Data in Table 14 indicate hunting pressure changed considerably
between areas and from year to year.
General regional variations in hunting
pressure were somewhat in relation to pheasant abundance and season length.
A pre-season flight was conducted a few days prior to the 1968 season.
In
the experimental area, 121 vehicles were tallied as compared to 106 in the
control.
These local vehicles were subtracted from the totals of opening
weekend counts to provide an estimate of hunter influx (Table 14).
Only a small percent of the local vehicles tallied in the pre-season counts
were believed used in hunting during the opening weekend.
Observations on
study plots in Sedgwick County (Snyder 1969b) indicated less than twenty
percent of the hunters were driving local vehicles.
Some of these were
probably from local towns and not included in the pre-season census.
Others
may have been non-hunting farmers traveling to or from home.

�- 41 -

100
95
:r: 90
VI

II)

c:
Q)

:r: 85
.....•

z
II)
II)

Q)

0

80

u

s 75

VI

+'
c:
ClI

u
70
~
Q,

r =

65

.95

0
60

0

2.0

2.5

3.0

3.5

4.0

Young Per Hen

4.5

5.0

5.5

6.0

(y)

Fig. 12. A scatter diagram showing the relationship between the
observed percentage hens with broods and the young per hen ratio.

�- 42 -

20

0

18

0

c:
Ql

:I:
~
Ql
a.

16

0&gt;

14

§

0

0

)i

x
•...•.

12

0

Ql

M

0..

~
0&gt;

.~
~

0

(])

10

0

tIl

0

8

0..
tIl
Ql

6

0 0

M

•.-t

::i:

~
Ql
a.

4

III

.,. ... .58

0

III

+'
c:

2

III

III
Ql

..c

a.
0

1

2
Pheasants

4

3
Observed

Per Mile

Fig. 13 Birds per mile observed in spring multiplied
hen index in relation to the fall BPM index.

5

&lt;BPM)

by the young per

�- 43 Hunting

Pressure Following

the Opening Weekend

Systematic surveys were not conducted to measure hunting pressure following
the opening weekend of season.
However, partial inventories were obtained
while conducting other studies in 1965, 1966, and 1968. The results presented in Table 15 were contrasted with opening weekend estimates.
In
making this direct comparison, it was assumed that a square mile was observed for each linear mile travelled by vehicle.
Some biases and errors
were introduced, but the general comparison is believed valid.
In 1967, traffic counters were placed at a number of key locations to measure
traffic coming into the areas. Malfunctions of the equipment, along with
destruction by snow plows and vandals, resulted in a poor sample (Snyder 1968).

Table 15.
subsequent

Hunting pressure on the study areas during opening weekend
periods of the 1965, 1966, and 1968 hunting seasons.

Miles
Sampled

Period

Miles/
Party

1965

Miles
Sampled

Miles/
Party

1966

Miles
Sampled

and

Miles/
Party

1968

Opening weekend

348

1.57

348

.85

348

1.36 1/

Weekdays

541

42.23

651

23.25

269

26.90

Second weekend

80

8.00

140

6.36.

568

6.60

Third weekend

148

11.38

282

8.60

209

11.60 ?:.J

109

27.20

Fourth weekend
Fifth weekend

1/ Based

on the assumption 20 percent
local hunters were used in hunting.
1/ Second opening of split season.

Hunting

62

10.33

of the resident vehicles

plus non-

Season Recommendations

Sex ratio, crowing and production indices were used each year to provide
management recommendations
for setting hunting seasons.
Table 16 provides
season length information on the study areas during the six-year study.

�- 44 -

Land Posting Surveys
In the event of hen harvest, the percent of land open to hunting would
be an important consideration in harvest evaluation.
Because of threatened
mass closures, when the study was initiated, surveys were conducted to
determine the extent of land posting in the experimental area. In 1967
these were extended along survey routes established in the control.
Most
of the land remained unposted throughout the study period (Table 17). Little
evidence of year to year trend was evident.

Hunting

Success

In the event of hen harvest, hunter success would provide one of the best
measures of study success or failure.
Increased harvest as a result of
shooting hens on a sustained basis would provide one of the best methods of
study evaluation.
Over-harvest of hens would result in decreased production
and lower hunter success.
Hen harvest within the capacity of the population
would provide a sustained increase in harvest with subsequent increased
hunter opportunity.
Since hens were not allowed in the bag, hunter success was measured only
in 1968. Field contacts during the day, followed by check stations in the
evening, were employed during the first two weekends of season.
Due to manpower limitations, sampling was not uniform throughout the study areas.
Check stations were set up at approaches to the highway between the two
study areas.
As a result, most samples came from localities proximal to
these stations.
Adequate samples were not obtained in the northern part of
the experimental area, where pheasant densities were low; nor in the southern part of the control, where good populations existed.
Biases prevented
direct reliable comparison of indices on the two study areas.
This was in
direct contrast to pheasant density information presented in the previous
segment.
There, BPM, CY and FPI were higher in the control.
The data in Table 18 indicate hunting success dropped slightly on the
second weekend of season.
Confidence cannot be placed in data for the
remainder of the season due to inadequate sample size. But, the average
hunter wasn't very successful at bagging pheasants in Colorado's best pheasant range.
The hunter's inclination to walk the large fields declined as
the day and season progressed.
Need for habitat manipulation to facilitate
harvest as well as to improve production was evident.

Environmental

Influences

When the study was initiated, it was hoped that similar land type, land
use, cover types and climatic conditions would produce consistently similar
pheasant production and density on the two areas.
But, this situation did
not follow.
Pheasant populations on the control dropped dramatically below
those on the experimental area in 1964 and remained there in 1965 (Fig. 4).
Production brought the control population up to that on the experimental
area by 1967. In 1968, the experimental area density declined.

�- 45 Table 16. Hunting season information on the two study areas during the
six year study.
Year

Open Dates

Season Length

Bag and Poss.

1963

Nov. 9

Dec. 22

44

3 &amp; 6

1964

Nov. 8

Dec. 25

48

3 &amp; 6

1965

Nov. 14

Nov. 30

15

3 &amp; 6

1966

Nov. 12

Dec. 31

50

3 &amp; 6

1967

Nov. 11

Dec. 31

51

3 &amp; 6

1968

Nov. 16
Dec. 14

Dec. 1 &amp;
Dec. 31

33

3 &amp; 6

Table 17.

A comparison of land closure to hunting from 1963 through 1968.
Land Status
Posted "Hunting
by Permission"

Closed to
Hunting

Year

Area

Unposted

1963

Exper.

73%

23%

4%

1964

Exper.

72%

25%

3%

1965

Exper.

88%

10%

2%

1966

Exper.

84%

12%

4%

1967

Exper.

87%

8%

5%

Control

92%

3%

5%

Exper.

88%

5%

7%

Control

85%

6%

9%

1968

�Table 18.

Hunter success on the experimental and control study areas during the 1968 pheasant season.

Period

Location

Hunters

Hours
Hunted

Adult

First

Exper.

416

2,050

Weekend

Control

433

First

Both

Weekdays

Areas

Second

Exper.

Pheasants Harvested
Unclas sified
Young

Total

Birds
Per Hunter

Hours
per Bird

39

140

90

269

.65

7.6

2,020

26

104

117

247

.57

8.2

12

23

3

5

0

8

.67

2.9

104

410

8

32

l3

53

.51

7.7
+:-

Weekend

Control

90

489

6

20

15

41

.46

11.9

Third

Both

43

120

1

8

3

12

.28

10.0

Weekend

Areas

Second

Both

27

153

2

6

12

20

.74

7.6

Opening Dec.Areas
14 and 15

(j'\

�- 47 -

Fortunately, Swope (1964b) initiated a number of environmental measures at
the start of the study. These measurements disclosed some of the influences
responsible for population variances between the two study areas.
However,
some environmental factors could not be readily measured.
Herein lay one
of the major weaknesses in the study.

Long Term Environment
Approximately 580 square miles were mapped on the experimental area and 411
square miles were completed on the control from 1963 through 1966. Only
large continuous rangeland areas were excluded.
Cover type acreages and
percentages by area are presented in Table 19. Table 20 presents these
data on an "acres-per-section"
basis.
The various cover types were discussed by Snyder (1967). Most significant
among the figures was the high percent of land utilized for small grain
production.
Approximately 84 percent of the experimental area and 76 percent of the control were planted to small grains or were Summer fallowed.
Winter wheat was the predominate crop. Only small percentages of oats,
barley or rye were included in these figures.
During dry years with resultant crop failure, millets were substituted for small grains.
Since small
grains were produced only every other year, only half the acreage could be
considered productive pheasant habitat in anyone
year. Linder et a1.
(1960) and Trautman (1960) pOinted out the importance of small grain fields
for pheasant production.
The same is probably true in these study areas.
During the six year period of study, a pronounced trend toward deep well
irrigation was evident in both study areas.
Irrigated corn, sugar beets,
and beans was primarily located in the southeastern part of the experimental area and in the northeastern, eastern, and southern part of the
control.
About the same acreages were placed under irrigation on the two
areas.
The two areas contained similar cropland acreages.
Pastureland was the
one exception.
The average experimental section contained about 52 acres
of pasture, whereas about 87 acres of pasture were tallied per control
section.
Most were short-grass pastures of little value to pheasants.
The greater amount of pasture in the control was believed partially
sible for the increased proportion of fencerow found in that area.
fences were usually only marginal as pheasant habitat.

responPasture

The average control area section contained 9.4 acres of draw or low area
contrasted to only 3.4 acres per section in the experimental area.
In
many instances in the control, this cover was along intermittent drainages
and was of good to excellent quality.
In the experimental area, few draws
existed.
Low "slough type" areas were often of little value because they
were farmed when they did not contain water.
The experimental area contained more and better windbreaks than the control.
Most of these were located at occupied farm-lots, whereas more willows and
permanent cover were found along drainages in the control.

�- 48 -

Table 19. Cover types by acreage and percent on the experimental and control
study areas.

Cover Type

Experimental
Acreage
Percent

Control
Acreage

Percent

Crop Type
Green Wheat*
Fallow Wheat*
Corn
Sorghums, Millets
Pasture
Plowed
Miscellaneous**

156,668.4
158,042.3
4,313.0
17,205.2
30,940.5
1,556.4
2,011.0

41.29
41.65
1.14
4.53
8.15
.41
.54

97,741.8
101,406.9
3,827.4
11 ,611.1
35,599.2
1,432.4
1,418.0

37.23
38.62
1.46
4.42
13.56
.55
.54

Total Crop Types

370,736.8

97.71

253,036.8

96.38

Occupied Farm Lots
Vacant Farm Lots
Towns
Cemeteries
Highways
Railroads
Miscellaneous

1,580.7
342.6
656.0
10.4
161.7
50.9
33.4

.42
.09
.17
.003
.04
.01
.009

1,009.9
247.0
398.0
22.0
109.0
25.0
77.3

.38
.09
.15
.008
.04
.01
.03

Total Cultural Features

2,835.7

.74

1,888.2

.71

Tree Plantings
Draws &amp; Low Areas
Rights of Way
Fence Rows
Edge Cover
Miscellaneous

695.0
2,016.7
567.3
429.2
1,689.1
476.8

.18
.53
.15
.11
.45
.13

335.3
3,832.2
281.2
596.1
1,345.9
1,241.3

.13
1.46
.11
.23
.51
.47

Total Special Coverts

5,874.1

1.55

7,632.0

2.91

Cultural Features

Wild Areas and Special Coverts

GRAND TOTAL

379,446.6

* Or other cereal grain.
** Alfalfa was the major miscellaneous crop.

262,557.0

�- 49 -

Table 20. Cover type acreages
cover type percentages.

Cover Type

per 640 acre section based on study area

Experimental

Acres Per Section
Area

Control

Area

Cover Type
Green Wheat*
Summer Fallow
Corn
Sorghums, Millets
Pasture
Plowed
Miscellaneous**

264.26
266.56
7.30
28.99
52.16
2.62
3.46

238.27
247.17
9.34
28.29
86.78
3.52
3.46

Total Crop Types

625.35

616.83

Occupied Farm Lots
Vacant Farm Lots
Towns
Cemeteries
Highways
Railroads
Miscellaneous

2.69
.58
1.09
Trace
.26
.06
.06

2.43
.58
.96
.06
.26
.06
.20

Total Cultural

4.74

4.55

Tree Plantings
Draws and Low Areas
Right of Way
Fence Row
Edge Cover
Miscellaneous

1.15
3.39
.96
.70
2.88
.83

.83
9.35
.70
1.47
3.26
3.01

Total Special

Coverts

9.91

18.62

TOTAL ALL COVER TYPES

640.00

640.00

Cultural

Features

Features

Wild Areas and Special

Coverts

* Green wheat or other cereal grain.
** Alfalfa was the major crop.

�- 50 -

In sunnnary, it seems probable that differences between the two areas tended
to nullify each other and make the areas similar in their capacity to sustain pheasants.
More tree planting and larger acreages of small grain on
the experimental area were possibly compensated by more and better undisturbed permanent cover along drainages, and by slightly more roadside "edge
cover" in the control.
Most other land features were quite similar on the
two areas.

Annual Environmental

Measurements

Precipitation--Farmers
living in the vicinity of the central township corners, who expressed a willingness to cooperate, were asked to record precipitation.
Several became lax in their recording efforts as the study progressed.
Fortunately, others of the eight recorders, maintained good to
excellent records throughout the study.
Monthly and annual accumulations of precipitation, measured at the township corners, are provided in Table 21. Comparison with the long-term average for the area is provided.
Data considered too unreliable were excluded.
Better records were maintained during the growing season.
Sunnner precipitation was of more interest to farmers and was easier to record.
Fall and
winter moisture, usually in the form of snow, was light, infrequent, and
difficult to measure.

Vegetative Growth Measurements--Variability
of winter wheat growth was considered a potentially important factor in pheasant production.
In addition,
climatic influences on wheat might have a similar effect on pheasant production.
Therefore, wheat growth was measured each spring.
A sunnnary is
presented in Table 22. These data indicate variance in growth and height
among years was more noticeable than that between areas.
Total sunnnation
of average wheat growth measurements (Table 22) was 431 inches in experimental area contrasted to 409 inches on the control.

Stubble Height and Weed Overstory--In early fall, residual wheat stubble
and weed overstory were also measured.
Average stubble height for the six
years was 10.5 inches in the experimental area and 9.1 inches in the
control.
Weed overstory averaged 18.3 inches on both areas for the six
year period.

Photo Hubs--Vegetative
growth of the preceding growing season was pictorially illustrated by photos at designated photo hubs each fall. The purpose
of the photos was to detect differences between conditions on the two
areas.
However, a review of the photos indicated cover differences would
have to be quite dramatic to be detected.
Therefore, this phase of study
was discontinued.

�Table 21.

Inches of monthly and annual precipitation recorded on the experimental and control study areas.
1963

Month

Exp.

1964
Con.

Con.

Exp ,

1965
Exp. Con.

1966
Exp. Con.

1967
Exp.
Con.

1968
Exp. Con .

Long Term Mean ~/

January

.63 ))

.00 1/

.46

.58

1.51

.45

.17

.24

.04

.03

.31

February

.20

.38

.71

.19

.20

.22

T

T

.25

.25

.35

March

.58

.90

.47

.18

.79

.71

.32

.28

.10

.06

.99

April

.23

2.53

.51

.17

1.34

1.18

1.08

1.51

.94

.60

1.9l

May

4.08

1.33

1.69

4.51

3.73

.79

.71

5.12

6.10

2.42

2.46

3.19

June

.42

.80

4.26

7.51

5.39

6.24

4.68

6.38

6.02

2.37

1.64

3.37

July

3.05

2.62

.91

4.75

4.70

3.68

4.66

3.86

1.50

1.98

1.91

2.46

August

1.36

3.12

.32

1.60

1.19

4.34

5.09

2.03

.47

3.77

3.16

2.00

2.45

.45

.39

1.51

2.82

4.06

2.19

1.05

1.16

.03

.23

1.48

October

.95

.32

.14

2.15

1.48

.50

.75

.54

.60

.34

.43

.92

November

.45

.09

.04

.01

.01

.31

.16

.28

.35

.06

.12

.44

December

.15

.09

T

.90

.31

.17

.19

1.02

1.25

.42

.44

.39

11.56 25.09 20.75 23.93 20.99

21.85

19.48

12.72 11.33

17.81

14.55

13 .51 11.94

1/ Precipitation received at Holyoke, Colorado which lies between the two study areas was substituted where study
area information was not available.
Based on 68 years of U. S. Heather Bureau Records at Holyoke, Colorado.

'2:../

t-'
I

September

Total

V'I

�Table 22. Hheat growth, stubble height and weed overstory measurements collected on the experimental and
control area transects.

Date

1963
Exper. Cant.

April 10
12
14
16
18
20
22
24
26
28
30
May

2
4
6
8
10
12
14
16
18
20
22
24
26
28
30

2.28

1965
Exper. Cant.

1966
Exper. Cant.

5.39

11.37

1967
Exper. Cant.

4.00

3.50

6.75

6.25

8.53
8.60

7.90

9.74

9.39

5.32
9.96

11.71

14.28

16.61

11.81

11.05
12.39

21.27

17.19

22.31

19.68

1968
Exper. Cant.

2.43
7.90

10.53
9.75

8.50

13.75

13.75

19.95
17.82

18.95

20.94

22.85

21.38
19.00
23.52

10.83
17.81

7.35
26.40

26.23

24.43

33.87

27.52

tv

11.45

10.68

15.50

13 .40

19.46

8.10
20.64

7.21
13.80

_4.16
21.16

10.83
25.16

8.68
20.19

23.53

17.50

23.95

23.05

12.53
V1

21.84

June

1
3
5
7
9
11
13
Fall Measurements
of \fueat Stubble
Hei,
\leed over-story

1964
Exper. Cant.

12.30
26.34

25.36

21.36

9.61
10.18

7.47
10.02

.....U)

1

'·1")

11.85
13.13

�- 53 -

Progress of Spring Stubble P1owing--Cessation
of pheasant nesting usually
occurred before spring plowing of wheat stubble was completed.
Since wheat
stubble offered attractive nesting cover, delay in plowing could significantly affect pheasant production.
Considerable nest destruction and occasional hen mortality was sometimes reported during spring plowing by farmers.
Each spring, the progress of spring plowing was measured to detect possible
differences between areas.
Table 23 indicates plOWing in the control
usually lagged slightly behind that on the experimental area. Greater variation occurred among years than between areas.

Table 23. Relative progress
harvest study areas.

of spring stubble plOWing

on the two hen

Percent Stubble
Experimental Area

Plowed
Control

Date

Year

April 23

1968

45

35

24

1964

20

17

26

1963

85

84

1967

75

67

27

1966

18

23

28

1965

44

47

48

46

1963

98

96

1964

66

48

1966

80

81

1967

91

81

1968

89

82

85

78

85

79

Mean

May

10-12

Hean

May

18

1964

Area

�- 54 -

Unmeasured

Environmental

Influences

Winter Weather--Winter weather conditions were observed in the region during
the last four years of study. During the winter of 1965-66, one to two feet
of snow accumulated in late December and early January and persisted in part
through February.
Conditions were more severe in the experimental areas
due to greater snowfall.
Glazed snow conditions developed.
Pheasants concentrated in farmyards and she1terbe1ts and apparently survived with little
mortality through the period.
Crowing indices in spring were higher in the
experimental area despite the more severe snow conditions that prevailed.
Blizzard conditions occurred twice during the last four years of study. A
March, 1966 blizzard produced considerable snow. Four to six inches accumulated during the second occurrence in December, 1968. Neither lasted for an
extended period or appeared to significantly reduce pheasant populations.
Some mortality observed in road ditches after the 1968 storm may have been
due to hunters, who forced pheasants to leave cover areas during the storm.

Summer Climatic Factors-Hail--The
impact of summer storms, primarily hail
storms, on pheasant density was considered a second factor that could not
be readily measured.
Hail size, wind velocity, the amount of accompanying
rain, the time of occurrence in relation to period of hatch, and other
factors all determined the influence of such storms on pheasants.
Three hailstorms during June, 1964 blanketed almost the entire control area
(Swope 1965). He reported these storms may have played havoc with pheasant
reproduction.
The experimental area went unscathed by hail that year.
Hail
storms were not recorded in 1963 or 1965. Only one small area in the control
was severely hailed during August, 1966. Cloudbursts in the western part
of the control at that time sent flood waters east along several intermittent streams and considerable damage to vegetation along the drainages was
observed.
Again the impact on pheasants remained unknown.
In June, 1967
hail caused widespread crop damage in both study areas.
In the experimental
area, two severe storms hit parts of Sedgwick County.
A similar mid-June
storm hit much the same area in 1968.
The effect of these storms on pheasants was not determined.
Some searches
for mortality were made, but usually produced little evidence.
Finding
significant evidence during the summer would be difficult even if a large
crew was available to search, for the birds remain widely dispersed during
the nesting season.
Farmers in Some localities reported mortality following
the 1964 and 1967 storms.
But in other areas less severely hailed, pheasants apparently suffered little from the hail.
Drought--The hail storms in June 1964 were followed by July-August drought
that persisted late into the spring of 1965. From July, 1964 through April,
1965 moisture totaled 4.32 inches on the experimental area and 2.92 inches
on the control.
Normal expected precipitation for this period would be 11.25
inches. As a result, most of the winter wheat in the control area was
plowed under during the late spring of 1965. Surveys in the control central
township indicated only 12 percent of the land still remained in wheat.

�- 55 -

In contrast, soil moisture conditions were enough better in the experimental area so that a much smaller percentage was destroyed.
Again the
impact of drought and subsequent wheat destruction on pheasants could not
be directly measured.

The Relation
Pheasant Production
Influences

on Production

of Environment to
and Survival, and Harvest

and Survival

Martinson and Grondahl (1966) found a direct positive correlation between
pheasant production and May-June precipitation amounts in Southwestern North
Dakota.
That is, production was higher in wet years and poorer in dry
years.
Similar comparison of harvest in nine Northeast Colorado counties
with four station regional May-June precipitation averages for a 14 year
period produced a correlation coefficient of .14. Apparently other factors
exerted greater influence on pheasant production in Northeast Colorado.
Wheat growth, stubble height, spring plowing progress and other environmental
variables were plotted and tested against the study area BPM index. The
results are listed in Table 24. Of the several variables, only green wheat
height (Fig. 14) correlated significantly with BPM (r == .67",). The correlation of March-April moisture with BPM was quite poor (r == .07). However,
when data from 1964, a year of hail and summer drought, were removed, the
correlation rose to a significant level (r == .86) (Fig. 15). This supports
the contention that early spring moisture was important in its influence
on wheat growth and the onset of nesting, but adverse conditions following
in summer may discount influence of spring moisture.
During the study period, early spring moisture conditions were almost
routinely below normal.
If March through May moisture had been above normal,
the peak of hatch might have been moved up to the first part of June, or
even into May.
But again, excessive spring moisture might delay spring
plowing, resulting in high nest destruction and hen mortality.
More study
of the several variables is needed in the future.
Several of the factors in Table 24 are illustrated in conjunction with gPM
and the period and peak of hatch in Figures 16 through 19. Important variables that seem evident in these illustrations are summarized as follows:
1. Hatching peaked early in 1963 because June precipitation was abnormally
low. Poor production resulted.
2. Severe hail in the control in 1964 and in parts of the experimental
area in 1967 and 1968 occurred somewhat in unison with the main period of
hatch.
3. Drought conditions through the summer of 1964 curtailed reproduction.
These drought conditions persisted into the spring of 1965 retarding wheat
growth and delaying the peak of hatch.
Production in 1965 was sustained
through a long summer rainy period and possibly not completely measured by
brood counts.

�- 56 -

4. Below normal March-April precipitation, which slowed wheat growth in
spring, was a dominate characteristic throughout the period.
Wide population departures on the two areas in 1964 might be assessed to hail in
the control area, but were not as readily explained in the experimental
area.
5. Summer pheasant production was above average and sustained over long
periods in both 1966 and 1967. Above normal June-July moisture was received
both years.
6. A direct positive relationship was evident between June 1 wheat height
and fall pheasant densities.
The widest deviation from this relationship
occurred in 1967. However, during that year, considerable growth occurred
during the first part of June (Fig. 14).

Table 24. The relationship
density index.

Environmental

Spring

Plowing

May-June

of environmental

(Percent complete April 28)
(Study areas)

May-August

Precipitation

March-April

Moisture

(Study areas)

March-April

Moisture

(Excluding

Green Wheat Height
Wheat Stubble

to BPM, the fall

Correlation Coefficient
with BPM

Variable

Precipitation

variables

(Study areas)

.46

.30
.13

.07

1964)

(June 1)

Height

(Late summer)

Weed Overs tory (Late summer)

-.03
-.34

Heather, directly and through its impact on vegetation, seemed to govern
pheasant production and survival in Northeast Colorado.
Four main components were believed especially important during most years.
These were
(1) winter and early spring moisture conditions as they influenced wheat
growth in spring, (2) spring plowing of wheat stubble, (3) precipitation
amount and pattern received during the reproductive period, and (4) severity
of thunderstorms, especially hail, during the reproductive period.
Hinter
blizzards could be listed as an additional factor occasionally reducing
breeding stock. A hypothesis on the interaction of these and other influences is presented as follows:

�- 57 -

o

32

31

30

29

28

27

0

.-I
Q)

~

:I

26

0

'J

..c
eo
.0-1
Q)
..c

25

.u

co

Q)
..c

24

~

C-64
Severe
Hail

0

c

Q)
Q)

H

0

E-67

Considerable
growth in June

c&amp;

23

0

0

22

0

21

C-67

0

20

r

19

.65

0
!

1

2
Pheasants

Fig. 14.
density.

The relationship

3
observed per mile

4

5

(BPM)

between wheat height on June 1 and fall pheasant

�- 58 -

March April Moisture

3.5

- BPM

,-,

,'.,
.,
•...,C-64

,

• J

..•...•
E-64

3.0

2.5

o o
o

2.0

1.5

00
o 0

1.0

.5

00

o

r

o
1

o

= .07

with 1964 excluded
r

2

Pheasants

3
per mile

= .86

4

5

(BPM)

Fig. 15. March-April precipitation as an influence on fall pheasant
In 1964 hail and summer drought appeared to be overriding factors.

density.

�- 59 -

-1 aun.r

;0

se sa4~UI U1 ~461aH ~ea4M uaaJ~
00

1.0'&lt;:1'

f"lf"lf"l

NOOO

1.0'&lt;:1' N

0

00

1.0

f"lf"lNNNNNIl""lr-i

00
1.0
0\
r-i

-_ ....••----

~

- -... -- •.~ •...- ---

-- - - - - - ~ --

---_ •.. _--.

-

I/')

(WdS) anw

N

o

Jad s:tueslo""'td

0\00

t'-I.O

I/')

&lt;;j'f"lN

r-i

0

uOl~e~1d1~aJd ;0 sa4~uI

�Fig. 17
Normal precipitation, actual precipitation and green wheat height in comparison with the BPM
index and the peak period of production on the control study area. Letters at the base represent months
from March through August.

�26

,...
::f::

22

I

I

~~Jf..tj&gt;A';ClI;.~.Ii32):~"'g'U&lt;"""""'J.·'?~"~~~~~~;~~~:_}~:5:_~~:';.J

I

I

(:l~«'-'.itW.~'4.~~"').I~l1f.'~~;[f...(,~(5.~':;'R·(~O[?~:.t;.',)~&lt;l~~1
-1120

U

5

0-

co
4

18

(I)
.-I

.,-j
X

I

I-t

/'l/BPM

(l)

0.

•..-1

2

..c

a..

0.,

.,-j

.

+J

~1
10
(!J

+J
ro
+J

.:::

:.:.

III

s::
co

i::

..-10

3

-

1

.

- - -- - ---------------------------

0

14 hl
\...0

"-~-... ~~ '12 4-t
E~ .h~r--"·r';e.l·:;~··

Q.,

-----------------------

-~"."fl.if-NfJ'r.
,
{', ~~\?'(I

'~'~""

0

~ ~~~~':~r:!/~~[.~
~1

Vi

OJ

10

~

8
6
4
2
0

1963

1964

1965

1967

1966

1968

Fig. 18 March-April, May-June, July-August and annual accumulations of precipitation in relation
to the BPM index of fall density on the experimental study area
r

0'\

I-'

�,-. 5
:E

Il.

22

Q)

20

e4
..-I
oM3

:t:

18
~2
Vl

+I

16

§1
Vl

m

_I

fo

Ii

14

~~j~~~~~~~~~~~j~

'.,
....,.

////////////////
///////
/////////

.

- ,',' :.:&lt;....

/ / /' / / / / / / / / / / / / /

/////////////////
/////////////////
/////////////////
.J////////////////

d'

-::.. - ..

~----

----

~~~~~~~~-~~~~~~~~

~~~~j ~~~~~ ~~~~~~Ar:::::':-

////////////////
/(//////////////
////////////////
////////////~///

/////////////////

////////////////
)///////////////
////////////////
////////////////

/ / // / / // // / // // /

/ / / / / / / / / / / / / / / ~/

/ / / / / / / / / / / / / / / / 1:;:;

/////////////////
/////////////////
/////////////////
/////////////////
/////////////////

~

.

~~~~~~:;:; ~~:; j:;~~:; ~:;,..)

".".

•.

..-

--

,:....:...:

10...4-4

0
tIl

8
6
---::;:::::::::::::::,
..
-::/:;:&gt;;.;-::;::::;.;'

/) ,':;'l~:;::i~':):'~';:~

////////////////
////////////////
////////////////
////////////////
/;//////////////

4
2
0

1964

1965

1966

1967

u
Q)

..-::2:....

//////////////
//////////////
////////////////
////////////////

j ~~~~j~ ~~~~j~ j ~ ~~~:;~~~~:;~~~~:;~~~

1963

ro

+I

oM
0.
oM

~
Il.

~~~~~-~~~~~~~~~~~~~~~~~~~~~~~~~~/
-r
,,.,.,.".,
..'.,.,.,.
,.,.

///////////////
////////////////
////////////////
////////////////
////////////////
////////////////
////////////////
////////////////
///////////////
//////////////

+I

~~~~~~~~~~~~
12

.. ~.

////////////////

;::,:,.:. :&lt;:&lt;-'-, ..

s::

s

1968

Fig. 19 March-April, May-June, July-August and annual accumulations of precipitation
relation to the BPM index of fall density on the control study area.

in

Q)

~

0\
N

�- 63 1. Spring plowing of wheat stubble disrupts territories and initial nesting
efforts.
It reduces the available nesting cover in late April and early
May to near zero in most years.
2. Pheasant nesting is delayed.
Pheasants are then forced to nest in green
wheat.
If soil moisture is inadequate,as
in 1965, or winter and early
spring precipitation is below normal, wheat growth is retarded.
As a consequence, it does not provide suitable nesting cover when pheasants need it
most. Correlation of green wheat height and pheasant production supports
this hypothesis.
3. Delay of nesting subjects the eggs to stresses of summer heat and dryness.
If June and July are relatively wet and cool, nesting can be completed
successfully as in 1966 and 1967 and good production results.
But if summer
conditions are dry and hot, poor reproduction results.
Fortunately some
moderate weather conditions usually exist each summer.
Tenacious nesting
efforts by the hen payoff during such periods.
4. Impact of hail and severe thunderstorms is generally independent of
other environmental variables.
Fortunately, hail is often localized.
Storms
in the control during 1964 were an exception.
Unfortunately, we cannot assign numerical ratings to the impact of hail or
drought on pheasant reproduction.
Our measurement of environmental influences remains incomplete.
More refined analysis of the data available in
this study seems futile under these circumstances.

The Relationship

of Environmental

Influences

to Hen Harvest

Previous studies on pheasants, quail and other species indicate that respective influences exerted by density dependent and density independent environmental limitations determine the harvestable surplus.
The higher the
density dependent influences, the greater the allowable harvest. With that
in mind, we will briefly review the previously listed environmental influences.
Winter Survival--Management
Unit 1 data and that of previous studies indicate
density dependent factors were not of major importance in winter mortality.
On the study areas, winter cover conditions did not vary greatly in either
quality or quantity from year to year or between areas. Yet spring populations varied widely.
We have little evidence to indicate that winter acted
as a major bottleneck reducing populations to a consistent carrying capacity
level. However, we cannot exclude the possibility of subtle limitations due
to competition for winter cover, escape from predation, etc.

Nesting and Production Conditions--In early spring, wheat stubble offered
an abundance of attractive cover. Spring plowing destroyed all nests established in wheat stubble, changing the situation in a hurry. Only the meager
roadside cover and a few unfarmed areas remained to provide suitable nesting
cover.
If this limitation continued through the summer, available nesting
space would limit production and hens would be surplus.

�-

64 -

Farmers have reported finding nests in wheat fields during July harvest
and during late sununer tillage of stubble.
Green wheat apparently became
suitable as nesting cover in late May.
The competitive nesting bottleneck
was again removed.
Trautman (1960) in South Dakota and Linder et al.
(1960) in Nebraska, found that small grain fields provided valuable nesting
cover.
Nesting success was much higher in small grain than along roadsides
and other areas.
Northeast Colorado wheat fields provided vast nesting
areas in sununer in comparison to the number of birds available to use them.
It is difficult to believe hens must compete for nesting cover during the
sununer. Instead, density independent climatic factors seem predominate at
this time. Hail, high temperatures, drought or rain exert their influence
irrelevant to pheasant density.
A hypothesis was previously presented, stating that C rather than CH best
represented breeding population density because of sp~ing hen mortality.
If this is true, then the period of mortality became increasingly important.
If mortality occurred during spring plowing before green wheat was suitable
for nesting, we again had a harvestable surplus of hens. Farmers frequently
reported mortality of nesting hens while plowing wheat stubble.
But, if
little hen mortality occurred during this period, or if most of it occurred
after mid-May, then the harvestable surplus would be greatly reduced.
Production in wheat fields seemed to be the result of random nesting success in
a non-confining environment.
If so, production was a direct result of the
number of hens present.
Clearly, our limited knowledge of environmental
influence on pheasants does not permit us to draw conclusions at this time.

CONCLUSIONS

AND RECOMMENDATIONS

For Current Management
Conclusions
1. Of the population indices tested, birds per mile (BPM) and young per
hen (Y), obtained during production counts, and the crowing index (C),
obtained in spring, provided the most reliable study area trend data.
Birds
per mile (bpm) obtained during spring sex ratio counts were least accurate
of those tested.
2. A CY index (crowing index X young per hen) closely approximated BPM
as an index of population level and trend. Use of this index instead of
FPI eliminates need for spring sex ratio counts during "cocks only" hunting
seasons.
In other words, rooster densities more accurately measured breeding
capacity than hen densities in spring did. The reason for this was not clear.
4. The young per hen index (Y) was almost completely determined by the proportion of hens successful in nesting.
Therefore, clutch size, hatchability
and fertility were not important variables.
5. Spring breeding population levels partially determined the number of
young.
Therefore, they were, in part, responsible for fall density levels.
But productivity, determined by nesting success, was the primary factor
responsible for fall density levels.
6. Census information from Small Game Management Unit 1 involved larger
samples than those obtained on the study areas. However, study area data

�- 65 -

were far superior in accuracy.
More persons were involved in management
unit sampling, possibly accounting for some differences in sampling accuracy.
7.
Best production census occurred when the majority of the young reached
six to nine weeks of age. This peak period usually existed from mid-August
through early September in Northeast Colorado.
Brood counts obtained in
evening were equal in quality to those obtained in morning.
8. Northeast Colorado production was usually determined by four environmental variables: (1) winter and spring moisture and its influence on wheat
growth, (2) spring plowing of wheat stubble, (3) precipitation amount and
pattern during the reproductive period, and (4) severity of thunderstorms,
especially severe hail, during the reproductive period.
Some evidence was
found indicating that wheat growth in spring could be used as an indicator
of pheasant production success in dryland areas in Northeast Colorado.
9. There was little difference in the amount of land closure from one year
to the next on two study areas. No trend in closure could be detected.
10. The average hunter put in seven or more hours hunting time per bird
harvested.

Recommended

Applications

to Management

1. Sex ratio counts should be discontinued under existing "cocks only"
management.
2. The fall population index (FPI) should be replaced by the CY and BPM
indices for measuring
population trend.
3. Construct a regression line between CY and BPM using both study area and
management unit data for past years.
Use this graph as a reference in the
future. Each year, plot indices on this graph and compare them with the
regression line. If the two indices closely approach the line, then confidence in census results is increased.
In contrast, one or both indices are
suspect of error when they deviate widely from the regression line. In
this case, additional census is needed prior to submitting season recommendations.
4. Young per hen (Y) and percent successful hens (SH) should be compared
each year in the same manner that CY and BPM are tested.
Be sure all field
personnel flush and record hens with and without broods by standard, well
explained procedures.
5. More late summer brood count replicates are needed to improve the
accuracy of management census.
For example, in Management Unit 1, at least
one more replicate per route is needed.
Counts should be conducted from
mid-August through early September before submitting season recommendations.
Use evening as well as morning counts to increase sample size.

Reenacting

the Study

A complete program of education, coopL~ation and realignment with the farmers
of Northeast Colorado is needed before attempting to conduct this study a
second time. The reader is referred to Swope (1964b) for further information
concerning information and education procedures.

�- 66 -

Assuming public acceptance throughout the study, do we have adequate knowledge to successfully
evaluate hen removal and determine its impact on the
population?
Can we measure or account for all the variables affecting the
hen population and thus, derive an accurate conclusion?
I feel quite confident that the answer is yes.
Procedures for reenactment
of this evaluation,
based largely upon knowledge gained from past experience, are presented in
the following pages.

Personnel
Two full time research employees will be needed to conduct this study.
Both
should be resident in the region and have a trained hunting dog for use in
the census work.
Part-time employees might be effectively utilized during
the spring census period.
Additional Division personnel would be needed
during the hunting season.

Study

Area Size and Location

Increasing the size of the study areas does not seem possible.
Larger homologous sites are not available in Colorado.
However, they would be preferred to reduce the effect of local weather influences.
The experimental
area must retain defined, easily recognizable boundaries.
Lyon (1956) reviewed literature and summarized findings on pheasant movements.
Considerable movement was noted by stocked birds but not in resident
flocks.
When considering the home range of the species and average annual
movements, I do not think ingress or egress from these large study areas is
too important.
On this basis, I recommend that intensive study central townships be excluded in future tests.
Population

Inventory

Sex ratio, crowing, and brood count samples were all found to be inadequate
for obtaining statistically
valid evidence of the effects of hen harvest
within anyone
year.
The following recommendations
are presented to attempt
correction of this sampling deficiency.

Sex Ratio--First year pre-treatment
census should begin with the spring sex
ratio count in mid-April.
Thirteen routes, each 24 miles long, are needed
in each study area.
Counting procedures include:
1. Divide each route into two-mile segments with corresponding
tally forms
to record the birds observed per segment.
2. Select and census one route in each area Simultaneously.
Treat each
pair of routes in this manner.
Retain these pairs throughout the study.
3. Alternate the t,.-,observers between areas to remove bias due to their
individual ob se rva t Lo.i ab i i i t i.e s .
4.
Run each route in one direction only.
5. Complete two replicates per route.

�- 67 -

6. Record good harem flushes separate from those where conditions do not
facilitate a complete flush.
7. Maintain
a fifteen to twenty mile-per-hour
speed.
8. Terminate counts when altered sex ratios indicate the onset of nesting.
9. Run counts only under favorable weather conditions.
10. Check procedures used in Colorado (Sandfort 1960a) and in other states.

Crowing Counts--A minimum of five crowing routes are needed per area.
Each
should have ten stations spaced approximately
two miles apart.
Procedures
include:
1. Complete four replicates per station under favorable weather conditions,
if possible.
2. High-count-per-station
averages would be used in analysis.
3.
Pair routes and run simultaneously
in each area (as in 2 under sex ratio).
4. Follow standard Colorado census procedures
(Sandfort 1960a).
lwo variables that influence crowing counts should be examined (Swope 1967).
The influence that different sex ratios and densities exert should be determined, utilizing mornings when crowing trend counts cannot be effectively
conducted.
In addition, the distance that a crowing rooster can be heard
should be accurately determined under Northeast Colorado conditions.

Production Counts--Paired,
simultaneous counts would be run on the thirteen
routes set up for sex ratio counts.
1. Tally observations per two-mile segment.
2. Run routes in only one direction.
3. Initiate counts near August 1, and continue into mid-September.
4. Study brood ages and weather data to predict the optimum census period.
5. Conduct four replicates per route with complete or near complete flushes
recorded separate from poor observations.
Consider all birds observed in
BPM computations.
6. Be careful to distinguish hens with and without broods and to separate
broods if possible.
Assign a hen to broods (two or more young) flushed without a hen.
7. Conduct both morning and evening counts under favorable weather conditions
and follow procedures outlined by Sandfort (1960b).
8. Make supplemental brood counts in localities where severe hail has potentially reduced populations.

Pre-season and Post-season Sex Ratio Counts--Stokes
(1954) described procedures used to conduct fall and winter counts.
In general, two or more men
with dogs should attempt rapid and complete flushes of birds from winter
cover resting areas.
A minimum 1,000 bird sample per area is recommended
per census.
Take advantage of cold, windy days when birds are in cover.
Snow cover fac i Li t a t ee be t t c r counts, but cannot be relied upon in Northeast
Colorado.
Birds per mile indices should be considered secondary to the
primary objective.
Under heavy snow conditions obtain a total census of sample routes within
the two study areas by use of airplane or helicopter.
Sexes should be
tallied sepac~tely, if possible.

�- 68 -

Procedures

for Collecting

Environmental

Data

Spring Plowing of Wheat Stubble--This phase of study can be continued much
as previously conducted.
Standard routes on standard dates are preferrable.

Measurement of Wheat Growth--Select random fields throughout each area to
be measured on alternate years.
1. Recommended dates for these measurements include: April 1, May 1, May
15, June 1, and June 15.
2. Record dates when wheat begins to head out and when it reaches full
height.
Relate to measurement dates.
3. A faster, more effective technique for measuring cover is needed (see
Jones 1968).

Precipitation Measurements--Farmers
who express willingness to cooperate
would be asked to record precipitation.
Provide rain gauges and recording
forms to ten recorders distributed in each area. If possible, supplement
with automatic precipitation recorders, used at two or more locations.

Unmeasured Environmental Influences--There
is no easy or direct way of
assigning a numerical rating to a blizzard, hail or drought with respect
to its influence on pheasant reproduction and survival.
Normally, we expect
blizzards and droughts to affect populations in both areas on about the
same basis.
But hail storms can occur over a few square miles, or as in
1964, over an entire study area. The "backdoor approach", comparing populations after the storm with those before, in relation to other areas, seems
to be the best alternative.

Hail--Determination
of the size and area of hail occurrences is needed.
Residents in the hailed area should be contacted concerning hail size, intensity and duration.
Key locations would be searched if man-power and
time were available.
Relate information to period and peak of hatch.

Winter Storms--Key concentration areas in both study areas are to be searched
with dogs following severe winter storms.
Attempts would be made to relate
mortality to pheasant densities before and after the storm.

Harvest

Procedures

Past Procedures for Determining Hen Harvest--Swope (1964b) explained the
procedure used to compute the number of hen pheasants proposed for harvest
in 1963. This harvest was not implemented, so was not included in study
results.
A summary of this procedure follows.
1. The crowing index was used as a base.
It was estimated that roosters
could be heard over a four square mile area. Therefore, the approximate
number of cocks per squre-mile was obtained by dividing the index by four.
The 1963 index was 48. The number per square-mile was conservatively set
at ten.

�- 69 -

2. Spring and winter sex ratio data were compared.
The more conservative
spring ratio, where two hens were observed per cock, was used. Each section
therefore, contained twenty hens if it contained 10 roosters, or thirty
birds in total.
3. The young per hen (Y) was slightly over four in the experimental area.
Four was used as an index of prodcution.
4. Computations were made to determine the total fall population in the
SOO-plus square mile area.
They are listed as follows:
10 cocks x 500 sq. mi.
20 hens x 500 sq. mi.
10,000 hens x 4 young
one-half of 40,000
young hens + old hens

5,000
10,000
40,000
20,000
30,000

cocks
hens
young
young hens
total hens

5. Twenty percent of the total hens would equal 6,000 birds.
The requested
hen permits was dropped to 4,000 to compensate for spring and summer mortality and probably failure to observe brood less hens in direct proportion
to their number.
6. These figures were compared with the Colorado Small Game Hunter Harvest
Survey (Grieb and Hunter 1962) which showed that approximately 23,000 cocks
were shot, on the average, over the preceding seven years in Phillips and
Sedgwick counties.
The experimental area contained about half the pheasant
range in these two counties.

Future Procedures for Determining Hen Harvest--This same general procedure
should be u3ed in future study.
But, additional factors derived from this
six-year study and from literature can also be used.
In Table 25 I have attempted to compare (1) Small Game Hunter Harvest Survey
information (abbreviated to small game survey, (2) opening weekend aerial
hunting pressure surveys, and (3) the BPM index. Swope (1964b) stated that
the experimental area encompassed over half the pheasant range in Phillips
and Sedgwick counties.
It also included a majority of the better pheasant
range.
Therefore, two-thirds was used as an estimate of experimental area
portion of the hunting pressure and harvest for the two counties.
Grieb
(1968), and Funk and Grieb (1969) found that about 70 to 75 percent of those
hunting in the region were out on the opening weekends of 1967 and 1968.
As a result, 75 percent was used in projecting the opening weekend hunting
pressure for the first four years of study (Table 25). Grieb's (1968)
analysis indicated 38.7 percent of the pheasants were killed on the opening
weekend in 1967. In 1968, Funk and Grieb (1969) found that about 72 percent
of the birds were killed the first nine days of season.
Table 18 shows
that hunting pressure was about five times greater the opening weekend in
1968 than on the second weekend.
Based on these figures, about 55 percent
of the harvest occurred during the opcn Lng weekend in 1968. These harvest
estimates were used in both sets of opening weekend harvest estimates in
Table 25.
Small Game Survey figures were consistently above those projected from aerial
transect pressure survey data. The reasons for this wide divergence is
unclear.
Gross errors could exist in either or both sets of data.
Possibly

�- 70 -

two-thirds of the two-county total was too high a proportion of the experimental area harvest.
Errors also probably exist in the aerial survey
projections.
Season length, BPM and Small Game Survey information for 1963 and 1968 were
about the same. Therefore, we can project from the 1968 data, that 3,000
to 5,000 roosters were harvested on the experimental area during the opening
weekend of 1963. One or two hens in the bag would have increased hunting
pressure.
As a result, 4,000 hens might have been harvested the first weekend of the 1963 season.
This does not take into consideration harvest
reduction due to land closure.
Based on two-thirds of the two-county rooster harvest (Grieb and Hunter 1964),
the 1963 hen kill of 4,000 would have approximated about 40 percent of the
rooster harvest.
Accurate figures to determine the percent rooster harvest
are not available.
Sandfort (1954) found that about 45 percent of the roosters were taken in the Fleming area. Even if 60 percent were taken in 1963,
this would mean that less than one-fourth the hens would have been harvested.
I can only conclude from these meager projections that Swope's request for
4,000 hen permits was quite realistic.
Based on the preceding crude analysis, future hen harvest permit estimates can be obtained from fall indexSmall Game Survey comparisons.
The length of hen harvest seasons can also
be based on this information.
A third technique is also available as an aid in determining the number of
hen permits.
If pre- and post-season sex ratios and fall indices are available from the preceding year, a basic analysis can be completed.
The percent cocks harvested and total cock harvest can be related to the fall index
for the previous year (Petrides 1954). Assuming a constant percentage harvest, the fall indices for the subsequent treatment year can be used to
estimate the total fall hen population and the number to harvest.
Known hen harvest numbers, age ratio data, hunting pressure and other details
will be available after the study is underway.
Several methods are available which can be related to fall indices and subsequent harvest.
A partial
reference list to these includes Allen (1942), Dahlgren (1959), Kelker (1945),
Petrides (1949 and 1954), Selleck and Hart (1957), and Stokes (1954).
Wishart (1969) explained a new age technique, measuring the proximal primary,
that is applicable to both sexes. This measurement should be tested against
bursa and spur characteristics in Colorado to determine its degree of accuracyand
limitations.
Preliminary analysis in Colorado revealed that feather
measurements here were different from those in Alberta, but the technique
appeared to be a good one (Snyder unpublished).

Development of the Special Permit--Swope (1964b) explained procedures for
developing the permit.
Illustrations were included.
No recommendations for
improvement of this system are presented here.

�Table 25. Hunting pressure and harvest estimates in the experimental study area based on Small Game
Hunter Harvest Survey data and opening weekend aerial pressure surveys.
Small Game Survey Data
Opening
Weekend
Total
Harvest}.!
HuntersV

Opening
Weekend
Hunters}.!

Aerial Survel Estimate
Opening
Opening
Weekend
Weekend
Harve~t~.!
Hunters'll

BPM
Index

Year

Total Birds
Harvestedll

1963

9,71L'

3,572

--

2,679

--

1,250

2.22

1964

6,672

2,415

--

1,811

--

1,300

2.14

196.';

9,088

3,308

--

2,481

--

1,662

1.68

1960

12,243

3,871

--

2,903

--

2,354

4.27
I

1967

12,540

3,816

5,016

2,671

3,366

2,566

3.73

1968

8,739

3,274

4,806

2,423

2,920

2,000

2.17

II Based on 2/3 of total Phillips-Sedgwick County harvest.
~I Based on 2/3 of total Phillips-Sedgwick County

hunters.
1963 through 1966 hunter numbers were based on

11 Data based on Grieb (1968) and Funk and Grieb(1969).
1967 and 1968 data.

~I Based on birds per hunter (opening weekend) from Grieb (1968) and Funk and Grieb (1969).

51 From basic data in Table 15. Three hunters per vehicle was used as a base (Swope 1964a and b).
percent added to base to correct for additional hunters present second day.

Fifteen

-...J

I-'

�- 72 -

Controlling the Harvest--In 1963, four check stations were to be located at
key entrances into the experimental area (Swope 1964b).
Hunters were to
obtain their hen permit and also check out through these stations.
Similar
stations should be located on main access roads into the control to measure
hunter opportunity and harvest in that area.
Hen permits might be distributed through local license agents to area residents a few days prior to season.

Hunting Pressure--Aerial
surveys should be continued as in the past.
A preseason flight should also be made along each transect to determine the number of resident vehicles.
Ground surveys within the areas would determine
the percentage of local and non-local vehicles used in hunting.

Land Posting--Routes

and procedures

Evaluating

Procedures

Sampling

established

in the past should be resumed.

After the first year, all population, environmental and harvest data can be
analyzed and evaluated to determine where weaknesses exist, and if sample
sizes are adequate.
After this, sampling procedures will become routine.
Normally, winter and early spring conditions affecting survival should be
about the same on both areas.
Spring crowing counts are one of the best
indices of this. But fall, winter and spring sex ratio data will be invaluable to determine percentage hen harvest and the impact of this harvest
on breeding populations.

Evaluating

Hen Harvest

Results

Although sampling procedures become routine after the first year, analyzing
and interpreting the results will remain a constant challenge.
Two basic
models are presented in the following pages to aid in explanation of hen
harvest analysis.
Model l--Determining the percentage harvest of hens and cocks will be the
first procedure.
This information is based on:
(1) pre-season sex ratios,
(2) number of hens removed, and (3) post-season sex ratios.
An example is
illustrated, based on the assumption of equal populations on both areas.
Part 1:

Harvest

Data:

Area
Experimental
Hens
Cocks

Variable

Fall Indices

Control
Hens

Cocks

BPM

2.00

2.00

CY

190.00

190.00

�- 73 -

Part 1:

Harvest

Data,

(continued):

Area
Variable

Pre-season

Sex Ratio

EXEerimental
Cocks
Hens

Hens

Control
Cocks

100

110

100

10/

Unknown

110
4,000

Harvest

Unknown

0

Illegal
Post-season

Sex Ratio

207

100

25%

60%

248

100

10%

60%

Results
Percent Harvest

Illegal
Pre-season

Number

Harvest

16,000

14,500

16,000

14,500

4,000

8,700

1,600

8,700

Legal Harvest
Birds per Hour Hunted
Post-season

Number

12 ,000

12,700

8,700

.21

.15

5,800

14,400

5,800

The above data can also be determined using age ratios in place of sex
ratios (Selleck and Hart 1957). Young birds are more readily harves ted than
adults (Nelson 1948), so some bias in the results can be expected using the
age ratio method.
Part 1 of Modell
assumes equal harvest on both areas.
Hunting pressure
and harvest information from the two areas can be compared to determine if
this is the actual case. If not, hunting pressure and harvest information
can be used to predict post-season sex ratio and population differences
between the two areas.

Part 2: Winter to Spring Sex Ratios--The next procedure entails comparing winter to spring sex ratios.
If differences between winter and
spring sex ratios are noted they should be about the same for both areas.
Changes in sex ratios would only result from disproportionate mortality of
one sex during the period, or disproportionate censusing of the sexes in
winter and/or in spring.
We would expect to detect sex ratio differences
of ten percent or more as previously explained under "Results and Discussion".
Winter mortality of twenty percent for both sexes on both areas will be
assumed in continuing the model.

�- 74 -

EXEerimental Area
Hens
Cocks

Variable

Post-season

Number

Control Area
Hens
Cocks

12,000

5,800

14,400

5,800

9,600

4,640

11,520

4,640

Less 20 Percent Mortality
Survival to Spring

Part 3: Breeding POEulation Indices--Comparison
of breeding population
density on the two areas is the third step. If rooster harvest and winter
survival have been the same on the two areas, crowing indices should be
directly proportional to previous fall indices.
First year hen harvest
should not affect following spring rooster populations or crowing indices.
Hen harvest would have to reduce reproduction first. Potential hen harvest
reduction of roosters could not be detected until after the second hen
season.
The birds per mile (bpm) observed during past spring sex ratio counts was
unreliable.
Increasing the sample size would probably improve its quality,
so it should not be overlooked in comparing breeding population densities.
Comparisons can be made with the breeding population index as shown.

Variable

Crowing

Experimental

Index

Hen Index
Breeding

Population

Index (spi)

Area

Control Area

37.2

37.2

2.07

2.48

77.0

92.3

Part 4: Production--Production
data are presented in Model I using four
young-per-hen (Y) for both areas.
Note that a considerable difference in
fall population exists between the two areas. This is detected by the FPI
but not by the CY index. In previous study, spring sex ratios were consistently within ten percent of each other on the two areas.
In this hypothetical model, there were about twenty percent more hens-per-cock in the
control than in the test area. Thus, the CY index mayor may not be applicable in evaluating hen harvest.
As yet we do not understand the basis for
the CY index, so we cannot project its roll in hen harvest evaluation.

�- 75 -

Experimental

Variable

Successful

Hens

(75% nesting

(excluding mortality)

Less 30% summer and early fall mortality
by Sex

l6,590~

28,200

34,560

42,440

50,720

29,708

35,504

l3,118~

20,160~

(Y)

l5,344cf1

l31.4: 100

126.5:100

Fall Sex Ratio
Young-per-hen

4.0

4.0

Total Young

Fall Populations

8,640

7,200

success)

Young-per-successful-hen

Total Birds

Control Area

Area

4.0

4.0

422

498

FPI Total Young

38,400

46,080

FPI Total Birds

52,640

62,240

FPI

BPM (From Fig. 10) based on FPI
CY

1.15

1.15

148.8

148.8

Projection of this model into subsequent years of study would result in a
declining population on the test area and lowered harvest.
Hen harvest
would not be successful.
Model 2--The preceding hypothetical model assumed that pheasant survival and
production were not limited by density.
In other words, carrying capacity
had no influence on the population.
Model 2 projects the same basic data
but takes competition for survival and nesting cover into consideration.
It operates on the theory that higher density results in greater stress and
increased mortality.
At present we do not know to what degree density dependent variables operate, or when or how they affect Colorado pheasant populations.
Carefully measured and controlled hen harvest might provide some
of these answers.
Careful comparison of the CY, FPI, and BPM indices provides one method of doing this.
Part 1:

This repeats Part 1 of Modell.

Proceed directly

to Part 2.

�- 76 -

Part 2:

Winter

to Spring Survival

EXEerimental Area
Cocks
Hens

Variable

Total Birds
Mortality
Spring

mortality

Index

Hen Index
Population

Part 4:

Index (spi)

Production

4,640

hens

1/

Young per successful

hen

Total young
Total birds

(excluding mortality)

to fall 1/

less 30%

16,590

Fall population

1/ Disproportionate

4,350

Experimental

Control Area

37.2

34.8

2.07

2.48

77.0

86.3

Experimental

Successful

10,800

and Fall Measurements

Variable

Survival

25/0

Indices

Variable

Breeding

20%

due to higher density on control.

Part 3 : SEring POEulation

Crowing

5,800
20,200

9,600

Population

14,400

17,800

1:../

1/ Disproportionate

5,800

12,000

Post season sex ratio

Control Area
Hens
Cocks

mortality

Area

Control Area

75% = 7,200

67% = 7,236

4.0

4.0

28,200

28,944

42,440

44,094

29,708

less 32% = 29,984

l3,1l8

due to higher density

17,185

in the control.

12,799

�- 77 -

Part 4:

Production

and Fall Measurements,

Variable

Fall sex ratio
Young per total hens (Y)
FPI

Experimental

126.5:100
4.0
422

(continued)

Area

Control Area

l34.2 :100
4.0
466.3

FPI (total young)

38,400

43,200

FPI (total birds)

52,640

58,350

BPM (from Fig. 9) based on FPI
CY

1.15
148.8

1.35
139.2

Many variations of the two models presented here could occur.
For example,
hen harvest on the test area might increase rooster harvest on that area.
Or density-dependent
factors might operate in summer but not in winter or
vice versa.
By comparing sex ratios, age ratios, density indicators, and
production and harvest many of the variables can be checked by more than
one method.
This should provide increased confidence in the results.

Adjusting for Environmental Influence--As previously stated, the major weakness inherent in the study is our inability to measure the impact of differing environmental influences on reproduction.
For example, can we be certain
potential reproduction declines on the experimental area were due to local
spring storms or due to excessive hen harvest?
Our main hope in this case
would be to carry the study over a long enough period to eliminate the
short-term local influence of weather.
In past years following severe hail
or blizzards, pheasant populations have usually returned to pre-storm
levels within a couple years (Sandfort, Boeker and Swope 1954).
The probability that hail will completely b1anLet the experimental areas is
not too great.
If local populations within an area are reduced by a summer
storm these can be analyzed on a separate basis.
This is facilitated by
separating spring sex ratio and brood counts into two-mile segments.
Crowing
stations also are spaced at approximate two-mile intervals.
Additional
brood counts within the hailed zone can be conducted.
An estimate of production loss due to hail can then be obtained.
To illustrate, the follOWing
formula might be used to measure production loss due to hail in a localized
part of one area:

�- 78 -

First

Then

let

A

A

Study

area BPM or CY exclusive

B

Hailed

zone BPM or CY index

C

The study area bpi

D

The bpi

E

The projected

x D

E

(breeding

in the hailed

of the hailed

population

zone

index)

zone

BPM or CY index

and E - B represents

for the hailed

the production

zone

loss due to hail

C

Drawing Conclusions--The
main question concerning hen harvest is what percentage of hens can be taken as substitution
for density dependent natural mortality.
Two general procedures can be used to measure this.
First, by
plugging the variables of harvest and population indices into models we
can determine if hen harvest at a certain level reduces production.
We have
reviewed this general procedure on the preceding pages.
Previous analysis indicates that reproduction
is predominately
regulated by
density independent factors.
However, the possibility of hen mortality
occurring prior to the main nesting period exists.
Research into the degree
and period of potential hen mortality is needed.
The second evaluation technique would measure the end product, harvest.
If
hen harvest was successful,
the test area should produce sustained increased
harvest, or sustained increased recreational opportunity
for the hunter.
If
hen harvest curtailed reproduction
this would in turn reduce harvest on the
test area and reduce hunting opportunity.

LITERATURE

CITED

Allen, D. L.
1942.
A pheasant inventory method based upon
sex ratios.
Trans. N. Am. Wildl. Conf. 7:329-352.

kill records

and

1956.
The management outlook.
pp. 431-466.
In D. L. Allen (Editor),
Pheasants in North America.
Stackpole Co., Harrisburg, Pennsylvania,
and \"'ildl.Mgmt. Inst., Washington, D. C.
490 pp.
Bennett, L. J., and G. o. Hendrickson.
1938.
Censuring the ringneck
ants in Iowa.
Trans. N. Amer. Wildl. Conf. 3:719-723.
Dahlgren, R. B.
1963.
Rhythmic fluctuations in South
lations and associated adult mortality,
1947-62.
Conf. 28:284-296.

Dakota
Trans.

pheas-

pheasant popuN. Am. Wildl.

�- 79 -

Dale, F. H. 1952. Sex ratios in pheasant
Wildl. Mgmt. 16(2):156-163.

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J.

Funk, H. D., and J. R. Grieb.
1969. Colorado Small Game Hunter Harvest
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Grieb, J. R. 1968. Colorado Small Game Hunter Harvest Survey-1967.
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Colo-

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Survey-196l.
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, and
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, and
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, and
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, and
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Jones, R. E. 1968. A board to measure
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Kelker, G. H. 1945. Sex ratio equations and formulas for determining
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The crowing

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J. Wild1. Mgmt.

Klonglan, E. D. 1955. Factors influencing the fall roadside
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Linder, R. L., D. L. Lyon, and C. P. Agee.
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Trans. N. Am. Wi1dl. Conf. 25:214-230.
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on
Quart.

Martinson, R. K., and C. R. Grondahl.
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:;...tkota.
J. Wildl. M~mt. 12(1) :20-31.

NOl, "'"''

�- 80 -

Petrides, G. A.
age ratios.

1949.
Viewpoints on the analysis of open season
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1954.
Estimating the percentage kill in ringnecked
other species.
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sex and

pheasants

season.

and

Colorado

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1960b.
Pheasant brood survey.
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Quart. Res. Prog. Rpt.
April:49-55.

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, H. M. Boeker, and H. M. Swope.
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44 pp.

synopsisColo. Game,

�- 81 -

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___

, C. D. Besadny, and C. Kabat.
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Madison.
168 pp.

Wishart, W. 1969.
mal primaries.

Age determination
J. Wildl. Mgmt.

Prepared by
Warren D. Snyder
/
Wildlife Researcher

of pheasants by measurement
33(3):714-717.

of proxi-

��April

1970

- 83 -

JOB PROGRESS

State

of

_

No.

W-37-R-23

Game Bird

Plan No.

1

Job No.

Project
Work

~ __ ~C~O~L~O~RA~D~O

REPORT

Job Title
Period

Covered:

Personnel:

Pheasant
April

Nest

Site Selection

1, 1969 to March

Survey
16
Stud

31, 1970

Donald M. Hoffman, Warren D. Snyder, David de Calesta, Robert
L. Schmidt, Ronald B. Arant, Ronald G. Blumberg, Don Crane,
Courtney J. Crawford, Charles W. Reickert, Lawrence A.
Webster, and John F. Corey.

ABSTRACT

Three of the 4 quadrats (NE, ~~, and SW) were each stocked with 15 hens
and 2 cocks, and 1 quadrat (SE) was stocked with 16 hens and 2 cocks by
March 13, 1969 (Segment 22) in preparation for nesting studies in 1969.
Sixty-six pheasant nests were found in the 48 nesting plots representing
8 vegetative species or combination of species.
Thirteen nests were found
in 8 alfalfa plots, 12 nests were found in 8 intermediate wheatgrass
seeding plots, 11 nests in 4 volunteer forbs plots, 10 nests in 4 alfalfacrested wheatgrass
(mowed) plots, 7 nests in 4 winter wheat stubble plots,
and 6 nests in 4 tall wheatgrass
seeded plots.
Due to the variation in
numbers of plots by vegetative type available in 1969, number of nests per
cover type plot gives a better comparison of preference than total numbers
of nests by cover type.
On this comparison,
the first-year successional
weedy cover plots (volunteer forbs) ranked highest with 2.75 nests per
plot, followed by alfalfa-crested
wheatgrass
(mowed) plots with 2.50 nests
per plot, winter wheat stubble with 1.75 nests per plot, alfalfa with 1.63
nests per plot, intermediate wheatgrass
seeded "and tall wheatgrass
seeded,
each with 1.50 nests per plot.
Fewer numbers of nests were found in other
cover type represented.
Sixty-three percent of all nests in 1969 were
established
less than 15 feet from plot perimeters.
Average clutch size of
61 nests, where it was possible to determine this, was 8.1.
A minimum of
3 cocks and 13 hens, wild-trapped
in 1969 and early 1970, were restocked to
each of the 4 quadrats by March 11, 1970 for continuation
of the study in
1970.

�- 84 -

RECOMMENDATIONS

The change over of three cover types (winter wheat, hairy vetch, and white
sweet clover) to smooth bromegrass, intermediate wheatgrass,
and tall
wheatgrass.
respectively, was completed in 1969.
No new cover types are
recommended at this time, but field work on the study should be continued
for one more segment so stands of the new grasses can be adequately tested.

�- 85 -

PHEASANT

NEST SITE SELECTION

STUDY

Donald M. Hoffman

Field work on this study was conducted similar to the 2 previous years work
except an additional intensive nest search was completed in all types in
early July, 1969. The change over of vegetative types used very little or
not practical to establish along roadsides, started in the fall of 1968, was
completed in the fall of 1969. Intermediate wheatgrass was used to replace
all hairy vetch plots; smooth bromegrass was used to replace winter wheat
plots; and tall wheatgrass was used to replace all white sweet clover (and
volunteer forbs) plots.

P. S. OBJECTIVE
To compare pheasant nesting use of, and success in, (1) winter wheat, (2)
alfalfa, (3) crested wheatgrass,
(4) hairy vetch, (5) white sweet clover,
(6) alfalfa-crested wheatgrass mixture, (7) smooth bromegrass, (8) tall
wheatgrass, and (9) intermediate wheatgrass.

SEGMENT
1.
2.
3.
4.

OBJECTIVES

To maintain plots.
To measure nesting preference.
To determine nesting success by cover type.
To obtain wild pheasants and care for them following

METHODS

release

in plots.

AND MATERIALS

Wild pheasants were captured with a vehicular mounted cannon-net, field set
cannon-net, and hand netting at night in Phillips and Sedgwick counties in
Segment 22. Three of the 4 quadrats (NE, NW, and SW) were each stocked with
15 hens and 2 cocks and 1 quadrat (SE) was stocked with 16 hens and 2 cocks
by March 13, 1969 (Figure 1). An electric fence was placed around the quadrats and charged in March 1969 to reduce domestic cat problems.
A magpie
trap was built but caught no magpies.
Phenology measurements were again secured during 1969 to ascertain readiness
of the various vegetative species and cover types for use by nesting
pheasants.
An initial nest search in 1969 was made
with 8 men, including 5 W.C.O. Trainees
was systematically searched using ropes
segments.
Nests were marked, recorded,

during the period June 4 through 6
and others (Figure 2). Each plot
to divide the plots into narrow
and removed to encourage renesting.

�00
0'\

Fig. 1. Banding a wild-trapped pheasant cock prior to releasing at the Fort Collins Wildlife Research
Station. (D. Hoffman~ photo)

�- 87 -

o

1-1
4-l

..c
o
1-1
m
Q)
CI)

Q)

CI)

.u

~

�- 88 -

Figure 3 shows one of the nests found in an alfalfa vegetative plot. A
second nest search was made during the period July 1 through 12, 1969 by
2 men searching each plot systematically
in strips outlined by laths. No
ropes were used because of the limited number of men available for this
search.
Nests were again marked, recorded and removed.
A third and final
nest search was made during the period July 21-28, 1969, preceding and
following the mowing of the plots on July 24, 1969.
Distances were measured from each nest to the nearest edge and to the apex
or base of the triangular plot, whichever was closest, with a steel tape.
Each nest could later be located on a scaled diagram of the quadrats.
Numbers
of eggs, fate of the nest and estimated age of embryos were recorded.
In
addition, vegetative heights and vegetative species in the 4 cardinal directions from each nest were recorded.
Prior to the windrowing of the plots on July 24, 1969, 62 live pheasants
were recovered and placed Lri a covered holding pen. In September 1969, 14
additional wild-trapped pheasants (mostly juveniles) were added to the stock
held in this covered pen. Due to early, heavy, wet snows breaking down the
top of the large fenced pen, it became necessary to wing-clip and release
surviving birds into the southeast quadrat.
Several piles of brush were
added to the southeast quadrat to provide winter cover since vegetation
within the plots was flatteried from the early snows of October, 1969, and
never recovered.
Eleven wild pheasants were captured at the Fort Collins
Hildlife Research Station and added to the pen.
A great horned owl problem developed on the wing-clipped pheasants late in
the wintering period in the open holding pen so that it became necessary to
live-trap additional pheasants in the Holyoke area using a hand net and
spotlights from a vehicle to restock each quadrat with a minimum of 3 cocks
and 13 hens by March 11, 1970.
Piles of dead tree limbs were placed in the
remaining 3 quadrats to provide cover for the pheasants for the same reason
listed above.

Plot Composition
Plot Composition

for 1969 (Treatment

Following

1968 Nesting

Season)

In accordance with original plans, vegetative types used very little, or not
practical to establish along roadsides, were reworked following the 1968
nesting season.
These were then seeded to other vegetative cover types
thought to have potential for nesting cover.
Plot Number

1 - Hinter Hheat

Four of the 8 original plots planted to winter wheat were left in wheat
stubble for a residual cover check in 1969. The stubble was of excellent
quality for the first time during the study.
The other 4 were reworked and
seeded to smooth bromegrass.
Plot Number

2 - Alfalfa

These were all mowed and maintained following the 1968 nesting
that 1969 composition was comparable to 1968.

season

so

�ex&gt;
\0

Fig. 3.

A pheasant nest containing seven eggs, located in 1969.

(D. Hoffman, photo)

�- 90 -

Plot Number

3 - Crested

Wheatgrass

Four of these were mowed and 4 were unmowed following the 1968 nesting
season so that a residual cover check was secured in 1969 on half of the
plots and the other half were comparable to 1968.
Plot Number 4 - Hairy Vetch
All of these plots were reworked and seeded
following the 1968 nesting season.
Plot Number

to intermediate

wheatgrass

5 - White Sweet Clover

Four of these were reworked and seeded to tall wheatgrass and the other 4
were left unmowed following the 1968 nesting season.
The 4 unmowed white
sweet clover plots were used to check volunteer forbs residual cover in 1969.
Plot Number

6 - Alfalfa-Crested

Wheatgrass

Four of these were mowed and maintained as previously and 4 were left
unmowed following the 1968 nesting season.
The 4 unmowed plots were used
to test residual cover in 1969.
Plot Composition

for 1970 (Treatment

Following

1969 Nesting

Season)

The changing of vegetative types used very little, or not practical to
establish along roadsides, started in 1968, was continued in the fall of
1969. The 4 remaining winter wheat plots were reworked and seeded to ,smooth
bromegrass and the 4 remaining white sweet clover plots were reworked and
seeded to tall wheatgrass.
Plot Number

1 - Originally

Winter Wheat,

now Smooth Bromegrass

Four of the original winter wheat plots were reworked and seeded to smooth
bromegrass in the fall of 1968. These were all mowed in the fall of 1969.
The remaining 4 winter wheat plots were reworked and seeded to smooth
bromegrass in the fall of 1969.
Plot Number

2 - Alfalfa

These were all mowed and maintained following the 1969 nesting season so
1970 composition will be comparable with 1968 and 1969. Two of these plots
including Number 2, Interior, NW Quadrat, and Number 2, Exterior, NE Quadrat,
are approaching alfalfa-crested wheatgrass mixtures due to invasion by
crested wheatgrass.
Plot Number 3 - Crested Wheatgrass
Four of these were mowed and 4 were left unmowed following the 1969 nesting
season so that a residual cover check may be secured in 1970 on half of the
plots.
Composition for 1970 is therefore comparable to that of 1969.

�- 91 -

Plot Number 4 - Originally

Hairy Vetch, now Intermediate

Wheatgrass

All of these plots were reworked and seeded to intermediate wheatgrass
following the 1968 nesting season.
Four of these intermediate wheatgrass
plots were mowed and 4 were left unmowed following the 1969 nesting season,
so a residual cover check may be secured in 1970 on half of the plots.
Plot Number 5 - Originally

White Sweet Clover, now Tall Wheatgrass

Four of the original white sweet clover plots were reworked and seeded to
tall wheatgrass in the fall of 1968. These were all mowed in the fall of
1969. The remaining 4 white sweet clover plots (volunteer forbs in 1969)
were reworked and seeded to tall wheatgrass in the fall of 1969.
Plot Number

6 - Alfalfa-Crested

Wheatgrass

Four of these were mowed and 4 were left unmowed following the 1969 nesting
season, so that a residual cover check may be secured in 1970 on half of the
plots.
Composition for 1970 is therefore comparable to 1969.

RESULTS AND DISCUSSION
Location

of Nests

Sixty-six pheasant nests were found in the 48 triangular-shaped nesting
plots representing 8 vegetative species or combination of species.
Table
lists numbers of nests by various cover types and treatments.
Figures 4
through 7 show the location of each nest in the 4 quadrats.

1

By numbers of nests, alfalfa (13 nests) ranked highest for nest establishment, followed by intermediate wheatgrass seeded (12 nests), volunteer forbs
(11 nests), alfalfa-crested wheatgrass, mowed (10 nests), winter wheat
stubble (7 nests), and tall wheatgrass, seeded (6 nests).
The larger number
of nests found in 1969 (66 nests) compared with 1968 (33 nests) resulted
from the increased number of nest searches, intensity of searches, and less
predation of nesting hens by domestic cats due to use of an electric fence
around the quadrats.
Limited domestic cat predation occurred late in the
nesting season of 1969 when a cat found a crawl hole beneath the perimeter
fence. At least 4 incubating hens were destroyed in this manner.
Due to a variation in numbers of plots by vegetative type, total numbers of
nests by vegetative type does not give a true picture of preference.
For
this reason, numbers of nests per plot were calculated (Table 1).
The first year successional weedy cover plots (volunteer forbs plots) were
highest with 2.75 nests per plot, followed by alfalfa-crested wheatgrass
(mowed) plots with 2.50 nests per plot, winter wheat stubble with 1.75 nests
per plot, alfalfa with 1.63 nests per plot, intermediate wheatgrass, seeded,
and tall wheatgrass, seeded, each with 1.50 nests per plot. Fewer numbers
were found in the other cover types represented.

�Table 1.
1969.

Number of pheasant nests found in 48 plots representing various vegetative types and treatments,

Plot
Number

Quadrat
Type and Treatment

NW

NE

SW

SE

1

Winter wheat (stubble)

1

3

2

1

Smooth bromegrass (seeded)

0

0

2

Alfalfa (mowed)

5

3

Crested wheatgrass (mowed)

3
4

Total

Number
Plots

Nests/
Plot

1

7

4

1.75

2

0

2

4

0.50

4

3

1

13

8

1.63

0

0

1

0

1

4

0.25

Crested wheatgrass (unmowed)

0

0

0

1

1

4

0.25

Intermediate wheatgrass (seeded)

4

3

2

3

12

8

1.50
I

\0
N

Volunteer forbs
(old white sweet clover plots)

1

3

3

4

11

4

2.75

5

Tall wheatgrass (seeded)

2

2

0

2

6

4

1.50

6

Alfalfa-crested wheatgrass
(mowed)

3

2

3

2

10

4

2.50

Alfalfa-crested wheatgrass
(unmowed)

1

1

1

0

3

4

0.75

17

18

17

14

66

48

1.38

5

6

Total Average

�- 93 -

370'

•

Mesa (m)

Agin (rr)

2
12'

Ager~m)

~

••

Mesa-Agcr (m)

/.
1/

4

6

@'

•

~,

;esa

(m)

.&gt;

.. :::~~~!U~-: ~
::~:r. C
.~

~rr)

~

~O'

.:

~.5) ~.

46'

#71\~ ~
\

/I

Agel

~

~:l(urn)

~

\10'

/1

r-: 1

~esa

•

\\
Age~(:~

__

j

.....

..

.

"Il!('IIi;;

"f 1;:,1,\;,"';:1

&lt;.lil:tdr;lI, Pl""",:tp!

Sc],,'ct;IlII (:!\!,1y, r\or:·, PlaIt 1.

Location of plv·.qs::1rtnest s in northwest

r:

L~~e_~)~

..

1)e".ilrd\"'·;I,&lt;

-

~;

12

~

_. __ ._-----_._----_ _--_ __ __ _ .._._._---_._------.-.-.- 12'
---_._---------_._-------

Fir;. 4.

J~

II • ~ V

~ \

~

Brin

'7(----- ~

\'c,s! Site

.Ioh Hi
quadr at ,

1969.

_ ..__ ..-----_ ..._._._------

I

.-.- .. _. __ ._ ....•..

I

�- 94 -

370'

Agin (rr

Agin (rr)

4

4

12'

6

Mesa-Agcr (um)
12'
-----------------_._------_

D('[&lt;Iikd

Fig. 5.

:\n"":~'·II]{·i:!', of lndi\'idi'al
();':"\l':ll.
r'IH':':-anf
Scl c ct i on ~lud·;. \\',)1'1; 1'1&lt;111 1 .. Job 1(i

I'{c;-,!.

...

Sile

Location of pheasant nests in northeast quadrat, 1969

_-_.- ._ .. - _.__ _._
..

.....

�- 95 370'

\ \

Meal (um)

5

7

(..sa

Agin (rr-)

--

•

4

12'

/

•\ \

II /. /.

~.

4

2

/

•

Trae (c)

!oj • f

Agin (rr)

~VU/

I
~

~?
.&gt;

•
6

3

. --;. -: /~

Mesa-Agcr

7

(m)

Agcr (urn)

(m )

~o'~
~

!'.esa(m )

/

/

/

12'

/I
3

L~__~~_cr_(_m)
__ ~

\\
.

/

Kesa-

I

~r

5

j

~rin (rr)_

L..

Agel (rr )

12'

Detail.·"

;\I';';:il:~l';~ll'lIh of IIl&lt;!i"i&lt;!II,t! (lu'l&lt;brl.

Scl ccti.in

Stlldy. \\'",h

(um)

Plan]

I'h"''';'!llt

"Ie,;t Sit"

.. Job IG

Fig. 6. Location of J:heasant nests in southwest quadrat, 1969D

~

�- 96 ...

------.

370'

.

~-

Agcr (urn)

""'~

12'

3

•

6
Mesa-Agcr

(urn)

4
Agin (rr)

•5

J)e!;.1;1,-,1

\;; .. ,;;.,' ;;1("':1..; "f
Sell'L'[j,,;,

Fig. 7.

1::&lt;I;-;;d'I::1

"{lIdy.

\\"rk

011,,,1;.;.1. "!"''''',;!iI ~~(':,t Site
Plall 1. .Inh ]()

Location of pheasant nests in southeast quadrat,

1969.

�- 97 -

Sixty-three percent of all nests in 1969 were established less than 15 feet
from plot perimeters as shown in Table 2. During 1969, 41 nests were found
in exterior plots and 25 nests in interior plots.
The terms "interior and
exterior" refer to the location of the individual plots in relation to a
central point where all four quadrats join. These are indicated in the
margins of Figures 4 through 7.
Although wild pheasants were observed to fly in and out of the enclosures
from time to time, none of the nests could definitely be attributed to wild
hens, since none were seen leaving nests.
Disturbance of birds was held to
a minimum, but periodic measurements of vegetative heights was necessary so
this may have deterred wild hens from nesting in the experimental plots.

Table 2.

Distance

Distance
Nearest

From
Edge

of pheasant

nests from plot perimeters,

1969.

Number
Nests

Percent

0 - 5'

9

14

5 - 10'

13

20

10 - 15'

19

29

15 - 20'

9

13

20 - 25'

9

13

25 - 30'

4

6

30 - 35'

2

3

35 - 40'

1

2

Total

66

100

Clutch Sizes
Five nests contained an unknown number of eggs. These were nests where the
eggs had hatched or were destroyed by predators and scattered.
The average
clutch size of the remaining 61 nests was 8.1, ranging from 1 to 22 eggs.
The two largest clutches of 20 and 22 each, were undoubtedly the result of
more than one hen laying in each nest, based upon embryo ages.

�- 98 -

Considerable variation was found in clutch sizes between various cover
types as shown in Table 3. Largest average numbers of eggs per clutch were
found in crested wheatgrass, unrnowed (1 nest had 15.0 eggs), alfalfa-crested
wheatgrass, unrnowed (2 nests averaged 10.0 eggs), alfalfa, mowed (12 nests
averaged 9.6 eggs), winter wheat (7 nests average 8.4 eggs), tall wheatgrass,
seeded (6 nests averaged 8.3 eggs) and alfalfa-crested wheatgrass, mowed
(8 nests averaged 8.1 eggs).

Fate of Nests
Of the 66 nests found and recorded, 60 were determined to be in an active
state of incubation, 3 were hatched successfully, and 3 were unsuccessful
due to nest searching operations or Windrowing operations (Table 4).
All nests found to be in an active state of incubation were removed to encourage renesting and to increase the size of the sample.
The large number
of nests found to be in an active state of incubation in 1969 reflects the
greater intensity in nest search operations and an additional second nest
search not used previously.
One each, nest was hatched successfully in alfalfa, alfalfa-crested
grass (mowed), .and alfalfa-crested wheatgrass (unrnowed) plots.

wheat-

Period of Nest Establishment
Table 5 lists periods of nest establishment based upon aging embryos according to procedures outlined in Game Information Leaflet Number 15 (Sandfort,
1965).
An initial peak in nest establishment occurred during the period May 1-15,
1969. Following the first nest search of June 4-6, 1969 and removal of all
nests found, additional peaks in nest establishment occurred during the
periods June 1-15 and June 16-30, 1969. Some hens continued to recycle
until the pheasants were captured and removed to a covered pen just prior to
windrowing of the vegetation in 9 of the 12 plots in each quadrat on July
24, 1969.
Fourteen nests out of the 18 established prior to May 31, 1969 (78%) were
found in plots having good carryover
residual cover from 1968. This better
residual cover was found in half of the plots (unrnowed plots and plots which
contained alfalfa).
Eight nests out of the 13 established
plots having good carryover
residual

after July 1, 1969 (62%) were found in
cover from 1968.

Alfalfa normally regrows following mowing in late July each year, but the
alfalfa-crested wheatgrass (unmowed) plots provided excellent field mouse
habitat.
This resulted in mice clipping the young alfalfa and much less
residual alfalfa was found in the alfalfa-crested wheatgrass (unmowed plots)
than in similar plots which were mowed in 1968.

�Table 3.

Clutch sizes found in 66 pheasant nests in various vegetative types, 1969.

Clutch
Size

1
Brin
(rr)

Mesa
(m)

1
1

1

1
1

2

1

1

2

3
Agcr
(m)

Plot Number and Treatment
5
5
3
4
Agel
Meal
Agcr
Agin
(rr)
(rr)
(um)
(um)

1
Trae
(c)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(um)

Total

1'&gt;"
0

0

1

0

0

1

0

0

2

1

6
6
1
4
3
6
4
7
2
4
1
6
2
2
1
1
2
0
0
2
0
1
5

Total

7

2

13

1

1

12

11

6

10

3

66

Ave. Clutch

8.4

5.0

9.6

7.0

15.0

7.4

6.5

8.3

8.1

10.0

8.1

1
2

3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
Unknown

3

1
1

1
2

2

1

1

2

1
1
1

1
1
1
1

3
1

1
1
1

2

2

2

1
1
3

1

1

1
1
2

1
1
1

1
1~"

1

,',More than 1 hen 1ayed in nes t.

\0
\0

�Table 4.

Fate of 66 pheasant nests in various vegetative types, 1969.
Plot Number and Treatment
1
Brin
(rr)

2
Mesa
(m)

3

3

4

5

5

Type

1
Trae
(c)

Agcr
(m)

Agcr
(urn)

Agin
(rr)

Meal
(urn)

Agel
(rr)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(urn)

Total

Active

7

2

12

1

1

11

11

5

8

2

60

1

1

Hatched

1

Unsuccessful

Total

1
7

2

13

1

1

12

11

6

3

1

1

3

10

3

66

t-'

o
o

�Table 5.

Estimated period of nest establishment in various vegetative types, 1969. 1/

Period

1
Trae
(c)

1
Brin
(rr)

June 1-15

2

June 16-30

2

1

1

July 1-15

5
Agel
(rr)

2

1

2

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(urn)

3

5

3

2

1

2

3

6

1

3

1

2

13

Total

1

1

1

1

1

1

12

1

13
3

2

1

7

2

1

1

Unknown

Total

Plot Number and Treatment
5
3
4
Meal
Agin
Agcr
(urn)
(rr)
(urn)

2

3

2

May 16-31

July 16-31

3
Agcr
(m)

2

April 16-30
May 1-15

2
Mesa
(m)

11

6

1

18
17

.

2

7

I-'
0
I-'

1

2

1

1

4

10

3

66

1/ Period of nest establishment based upon aging of embryos according to Game Information Leaflet No. 15
(Sandfort, 1965).
July 21-28, 1969.

First nest search made June 4-6, 1969; second was made July 1-12, 1969; third was made

�- 102 -

Phenology
Heights of growth measurements were made on April 16 and 23; May 2, 13, and
23; June 2, 18, and 30; and July 17, 1969. Table 6 summarizes average
heights of vegetation on April 16, May 13, and June 18, and July 17, 1969.
Heights of vegetation was comparable to that in 1967, except for plots seeded
to grasses in the fall of 1968, with most plots having adequate cover for
pheasant nesting by May 13, 1969. The initial peak of nest establishment
found during the period May 1-15, 1969 indicates this was true. Heights of
vegetative growth appeared to reach a maximum for the season in mid-June 1969.
Plots were not irrigated
maintained with complete
new grass seedings.

in the spring of 1969 but good soil moisture was
plot irrigations in the fall of 1968 to establish

Height of Vegetation

Surrounding

Nests

Nests found prior to mowing and found in uncut plots provided an opportunity
for obtaining vegetative height measurements surrounding 60 nests.
Table 7
lists average heights and species of vegetation surrounding these nests.
The
average height of all measurements at the vicinity of the established nests
was 18.5 inches in 1969. This average vegetation height did not vary greatly
between periods of nest searches.

Stocking

Quadrats with Pheasants

for 1970

Prior to windrowing the plots on July 24, 1969 (Figure 8), 62 live pheasants
were recovered and placed in a covered holding pen. Six dead pheasants
were found in the nesting plots, including 4 eaten by a domestic cat which
had crawled under the perimeter fence at a wash-out, 1 killed by a bird dog
used in the recovery of the birds, and 1 killed by the windrowing machine.
Only 1 pheasant released prior to the 1969 nest searches could not be
accounted for.
In September 1969, 14 additional wild-trapped pheasants were added to the
stock held in a large covered pen. Four of these died shortly thereafter.
All surviving birds (58 hens and 10 cocks) were wing clipped and released
in the southeast quadrat on October 9, 1969, after early, heavy, wet snows
had broken down the covered pen top. Eleven wild-trapped birds were captured at the Fort Collins Wildlife Research Station and added to the open
pen.
A great horned owl problem developed late in the wintering period, resulting
in a loss of approximately one-third of the wing clipped pheasant population
prior to February 9, 1970 and an additional loss of one-third of the wing
clipped pheasant population from February 9, 1970 to March 11, 1970.
Thirty seven wild pheasants live-trapped in the Holyoke area with a hand
net and vehicle equipped with 2 spotlights, were added so that a minimum of
3 cocks and 13 hens were stocked in each of the 4 quadrats by March 11, 1970.

�- 103 -

Table 6.

Average height of vegetation in pheasant nesting plots, 1969.

Plot
Number

4-16-69

1

Vegetative Type
and Treatment

1

Smooth bromegrass (seeded)

2

Alfalfa (mowed)
New growth
Residual

3

Crested wheatgrass (mowed)

3

Crested wheatgrass (unmowed)
New growth
Residual

5

Intermediate wheatgrass
(seeded)

6.0
20.0

22.0

40.0

36.0

2.0

4.0

13.0

14.0

5.5
22.0

16.5

25.5

19.0

7.0

9.0

17.0

18.0

8.0
16.0

11.0

17.0

14.0

2.0

3.0

12.0

16.5

1.0
42.0

3.0

16.0

21.0

3.0

4.0

10.0

21.0

15.0
11.0

24.0
17.0

22.0
16.0

15.0
12.0

18.0
17.0

14.0
14.0

Volunteer forbs
New growth
Residual

5

Tall wheatgrass (seeded)

6

Alfalfa-crested wheatgrass (mowed)

6

7-17-69

Winter wheat (combined)
Volunteer
Stubble 1../

4

Average Height (Inches)
5-13-69
6-18-69

New growth
Alfalfa
Crested wheatgrass

4.5
7.0

Residual
Alfalfa

28.0

Alfalfa-crested wheatgrass (unmowed) 2/
New growth
Alfalfa
Crested wheatgrass

8.0

Residual
Alfalfa

15.0

1/ Winter wheat stubble was of excellent quality.
l/ New growth alfalfa was well clipped by mice in unmowed alfalfa-crested wheatgrass plots during 1969 growing season.

�- 104 Table 7.
1969.

Heights and composition of vegetation surrounding 60 pheasant nests,

Nest
Plot
Date
Number Quadrat Number Measured
1
2
3

SE
SE
SE

5-1
3-1
i -e

6-4-69
6-4-69
6-4-69

4
5
6

SE
SW
SW

2-E
1-E
6-E

6-4-69
6-4-69
6-4-69

7

SW

6-E

6-4-69

8
9
10
11
12
13
14
15
16

SW
SW
SW
NW
NW
NW
NW
NW
NE

5-1
I-I
1-1
2-1
6-1
2-E
2-E
4-E
2-E

6-4-69
6-4-69
6-4-69
6-5-69
6-5-69
6-5-69
6-5-69
6-5-69
6-5-69

17

NE

5-E

6-5-69

18
19
20

NE
NE
NE

5-E
4-E

6-5-69
6-5-69
6-5-69

21
22

NE
SE

2-1
4-1

6-5-69
6-6-69

23

SE

5-E

6-6-69

24

SE

5-E

6-6-69

25
26
27

NW
NW
NE

5-E
4-E
2-1

6-6-69
6-6-69
6-6-69

28

SE

6-E

7-1-69

29

SE

5-E

7-1-69

30

SE

4-E

7-2-69

31

SE

4-E

7-2-69

r-s

Composition

Ave.
Height
(Inches)

Mustard-cheatgrass
Crested wheatgrass
Winter wheat, stubble
Winter wheat, growing
Alfalfa
Tall wheatgrass, volunteer
Alfalfa
Crested Wheatgrass
Alfalfa
Crested wheatgrass
Bindweed
Winter wheat, growing
Winter wheat, growing
Alfalfa
Crested wheatgrass
Alfalfa
Alfalfa
Cheatgrass
Alfalfa
Crested wheatgrass-alfalfa
Cheatgrass
Crested wheatgrass
Cheatgrass
Winter wheat, growing
Bindweed
Hairy vetch-thistle
Alfalfa
Thistle
Cheatgrass-thistle
Cheatgrass
Crested wheatgrass
Kochia
Wild rye
Mustard
Cheatgrass
Alfalfa

20.0
16.0
20.0
22.5
23.5
15.5
19.0
14.0
20.7
23.0
9.0
37.3
37.0
23.8
13.8
15.5
16.5
16.5
22.3
23.0
14.5
17.5
9.3
37.8
12.7
13.0
22.5
15.5
13.0
9.0
20.5
7.7
28.0
30.3
11.3
23.3

Alfalfa
Crested wheatgrass
Cheatgrass
Crested wheatgrass
Dandelion
Cheatgrass
Hairy vetch
Intermediate wheatgrass
Intermediate wheatgrass
Thistle

17.7
14.0
18.0
25.0
6.0
10.0
13 .5
17.0
14.0
22.7

Ave. Height
for Period
(Inches)

19.3

-----------------------------------------------------------------------------------

�- 105 Table 7. Heights and compositon of vegetation surrounding 60 pheasant nests,
1969, continued.

Plot
Date
Nest
Number Quadrat Number Measured
32

SE

5-1

7-2-69

33
34

SW
SW

2-E
6-E

7-3-69
7-3-69

35

SW

4-1

7-1-69

36

SW

4-1

7-1-69

37
38
39

SW
SW
NW

2-1
6-1
6-E

7-7-69
7-7-69
7-9-69

40

NW

4-E

7-9-69

41
42
43
44
45
46

NW
NW
NW
NW
NW
NE

2-E
5-E
I-I
5-1
2-1
2-E

7-9-69
7-9-69
7-10-69
7-10-69
7-10-69
7-11-69

47

NE

5-E

7-11-69

48

NE

6-E

7-11-69

49

NE

6-E

7-11-69

50
51
52

NE
NE
NE

1-E
6-1
4-1

7-11-69
7-12-69
7-12-70

53

NE

5-1

7-12-70

54

SW

2-E

7-18-69

55

NW

4-E

7-21-69

Composition

Ave.
Height
(Inches)

Cheatgrass
Mustard
Tall wheatgrass
Alfalfa
Alfalfa
Crested wheatgrass
Hairy vetch
Mustard
Cheatgrass
Tall wheatgrass
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Cheatgrass
Mustard
Alfalfa
Mustard
Winter wheat (growing)
Bindweed
Alfalfa
Alfalfa
Crested wheatgrass
Cheatgrass
Squirrel-tail grass
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Winter wheat (growing)
Crested wheatgrass
Cheatgrass
Hairy vetch
Cheatgrass
Mustard
Winter wheat (growing)

18.0
19.0
22.0
18.5
23.5
19.0
10.0
12.0
12.3
14.0
18.0
17.5
23.0
l3.0
17 .5
21.5
17.8
24.3
25.5
14.8
20.3
18.4
22.0
12.7
15.0
25.0
17.0
16.3
12.0
29.8
l3.0
12.0
15.0
17.5
26.0
35.0

Alfalfa
Crested wheatgrass
Cheatgrass
Intermediate wheatgrass

27.3
14.0
17.0
19.5

Ave. Height
for Period
(Inches)

18.0

-----------------------------------------------------------------------------------

�- 106 -

Table 7. Heights and composition of vegetation surrounding 60 pheasant nests,
1969, continued.

Nest
Plot
Date
Number Quadrat Number Measured
56

NW

6-E

7-22-69

57
58
59

NE
NE
SW

5-1
5-1

7-22-69
7-22-69
7-23-69

60

SW

5-1

7-23-69

r-s

Composition
Alfalfa
Alfalfa-crested wheatgrass
Cheatgrass
Winter wheat (growing)
Bindweed
Kochia
Kochia

Ave.
Ave. Height
Height
for Period
(Inches)
(Inches)
18.0
13.5
17.5
26.5
7.3
10.0
30.5

18.3
Average height all measurements

18.5

�I--'
o
--.J

Fig. 8.

Vegetation on selected plots was windrowed on July 24, 1969.

(D. Hoffman, photo)

�- 108 -

Key to Abbreviations - Figures 4 through 7 and Tables 3 through 5.
Plants
ConnnonName

Symbol

Scientific Name

Agcr

Agropyron cristatum

Crested wheatgrass

Agel

Agropyron elongatum

Tall wheatgrass

Agin

Agropyron intermedium

Intermediate wheatgrass

As

Astragalus spp. (Madison strain) Hairy vetch

Brin

Bromus inermis

Smooth bromegrass

Meal

Melilotus alba

White sweet clover

Mesa

Medicago sativa

Alfalfa

Trae

Triticum aestivum

Winter wheat

Treatment - Following 1968 Nesting Season
(c)

Combined only - residual cover check in 1969

(cr)

Combined and reseeded

(m)

Mowed only

(mr)

Mowed and rototilled

(rr)

Reworked and reseeded

(um)

Unmowed - residual cover check in 1969

LITERATURE CITED
Sandfort, W. W. 1965. Aging pheasant embryos. Colo. Game, Fish and Parks
Dept., Outdoor Facts, Game Information Leafl. No. 15. 2 p.

Prepared by
Wildlife Researcher

�- 109 -

JOB PROGRESS

State

COLORADO

of
No.

W-37-R-23

Game Bird

Plan No.

1

Job No.

Project
Work

REPORT

Job Title

Pheasant

Period

April

1, 1969 to March

Robert

L. Schmidt

Covered:

Personnel:

Roadside

Cover

Survey

18

Evaluation

Study

31, 1970

and Warren

D. Snyder

ABSTRACT

Seeding of roadside study plots with grasses and grass-legume
cover was
continued in 1969.
Spring seeding efforts were generally unsuccessful.
Modification
of the grass drill to provide better seed placement proved
worthwhile.
Fair to good stands were established west of Holyoke after
August rains.
More than one-half the study plots will be available for
nest search in 1970.

��- 111 -

PHEASANT

ROADSIDE

COVER EVALUATION

Warren

STUDY

D. Snyder

P. S. OBJECTIVE
To compare pheasant production under natural roadside conditions with the
following cover types to be established along roadsides: (1) grass, (2) grasslegume mixtures.

SEGMENT
To establish grass and grass-legume
east Colorado.

METHODS

OBJECTIVE
cover on roadside

test plots in North-

AND MATERIALS

A listing of methods and materials used in this study was presented in the
previous Segment Report (Snyder, 1969). Changes and additions during this
work segment include:
replacement of shoe-type furrow openers on the grassland drill with double disk openers.
Depth bands on these flexible attachments permitted seed placement at 3/4 to 1 inch depth.
Also, herbicide treatment of a number of study plots was conducted in June after late spring rain
prompted rapid weed growth.
A 2-4-D Amine solution was applied at a rate of
two pints per acre using a trailer type boom sprayer.

RESULTS AND DISCUSSION
Study plots believed to have adequate stands for nest searching in 1970 are
listed in Table 1. Their respective control plots are also included.
The shoe-type furrow opener attachments used on the drill during 1968 and the
spring of 1969 did not permit proper depth control in seed placement.
In
addition, the ground was opened and disturbed permitting excessive moisture
loss. As a result, only poor to fair stands were achieved.
May and June rains produced heavy weed growth on the plots.
Spraying with
2-4-D Amine base herbicide in mid-June only partially controlled weed growth
and subsequently, several plots were mowed.
Where grass stands were considered inadequate, the plots were cultivated and reseeded.
The double disk
furrow openers with depth band attachments were much more effective in producing adequate stands.
Loose soil and weeds along plot boundaries both
were found detrimental to grass establishment.

�- 112 Most of the plots listed in Table 1 were attained through July-August seeding
after summer fallow.
August rains were adequate west of Holyoke.
Those
plots to the east did not receive sufficient rain to germinate seed. October
snows stimulated some seed germination, but cold temperatures were not conducive to stand establishment.
Several of the plots designated exclusively for grass contained fair stands
of alfalfa after the late summer seeding.
Apparently sufficient alfalfa seed
leaked from the legume drill box to produce these stands.
Observations were made of a pure dense stand of intermediate wheatgrass
planted several years ago by a local farmer.
In 1969 weeds were almost completely excluded from the stand. Grass density appeared to stunt growth,
resulting in only fair pheasant nesting cover.
October snows completely
flattened the grass, so that no winter cover was available.

LITERATURE

CITED

Snyder, W. D. 1969. Pheasant roadside evaluation study. Colorado
Game, Fish and Parks, Game Research Report. April, pp.37-45.

Prepared

by

U'~(/).¥J
Warren D. Snyder
Wildlife Researcher

Div. of

�- 113 -

Table l. Roadside study plots with fair to good grass stand and their
controls as of April 30, 1970.
Seeded

Group

Associated

Plot

Control

fj OJ
p.
a
e-,
fj-.-I
fj O.j...l
E-I
-.-10
0
,...
.j...IOJ
-.-I
.j...I
ctlCf.l
OJ
:&gt;
0
0
OJ ofj a
U)
HO
0

Z

0
E-I

p::

fj1/ OJ2/
p.afj
.
.-I :&gt;.
E-I
fj O.j...l
-.-10
0
,...
.j...IOJ
•.-1
.j...I
ctlCf.l
OJ
:&gt;
0
0
0
OJ Ofj
HO
0
sr:

1

7

43(2)

13

- 2 -L

P. Travis

18 - 5 - C1

2 - 7 - 43

24

- 2 -21

P. Travis

24 - 2 - C1

1 - 8 - 44

21 - 5 - G

w. Sprague

20 - 2 - C2

o,

,...

OJ

il
;:l

-.-I
..c:
UJ

~

OJ
fj

bO
ctI

Farmer

21 - 5 - L
2 - 8 - 44

28

- 6 -G

20

w. Sprague

28 - 6 - L
1 - 7 - 44

21

- 8 -G

21

- 8 - L

P. Lappart

- 3 - C1

28 - 5 - C1
29

- 1 - C2

21

- 1 - C2

16 - 3 - C1

1 - 6 - 44

11 - 7 - L

C. Beeber

11 - 8 - C2

2 - 6 - 44

11 - 6 - G

C. Beeber

11 - 4 - C2

11 - 6 - L
1 - 7 - 45

12

3 - 7 - 45

15 - 3 - G

- 7 - Gl/

10 - 1 - C1

v. Rufert

1 - 4 - C1

v. He1goth

15 - 4 - C2

v. He1goth

15

15 - 4 - L
4

- 7 - 45

15 - 7 - G
15

- 8 - C2

- 7 -L

5 - 7 - 45

9 - 5 - G

H. Hassler

9 - 5 - C2

- 7 - 45

8 - 3 - L

L. Miller

8 - 4 - C1

6

8 - 2 - C2
7 - 7 - 45

17

- 8 -G

L. Boller

17 - 8 - L
8 - 7 - 45

17

- 6 -G

17

- 6 -L

18 - 1 - C1
18

L. Boller

- 7 - C2

17 - 5 - C2

----------------------------------------------------------------------------

�- 114 -

Table 1. Roadside study plots with fair to good grass stand and their
controls as of April 30, 1970, continued.

GrouE

Seeded

Plot

Associated

rd/01:-1
0

fj

fj •
..-!
O-IJ

0

• ..-!

0-

•..-!

)..&lt;
&lt;!)

~;::)
Z

9

..c
en

~

0
H

&lt;!)

-IJ

m

&lt;!)
00

fj

p::;

- 7 - 45

10 - 7 - 45
12

•..-!

eo

- 7 - 45(6)

o

o

-IJ&lt;!)

moo
o

fj

&lt;!)
0-

fj

O-IJ

H

p..-!

H
)..&lt;
&lt;!)

:&gt;

Ol=l
0
....:10 U

o

0

• ..-!

•..-!

-IJ&lt;!)

-IJ

t\lCI)

o

Farmer

&lt;!)
00

o

e-,

)..&lt;
&lt;!)

:&gt;

Ol=l
0
....:10 U

7 - 2 - L
7 - 2 - G
7 - 3 - Gli

K. C. Young

8 - 5 - C1

K. C. Young

7 - 4 - Cl

18 - 5 - G

L. Miller

13 - 2 - C1

18 - 5 - L
13 - 7 - 45(6)

0

0-

&gt;.

Control

18 - 6 - G

18 - 4 - C2
K. C. Young

13

- 1 - C1

18 - 6 - L
14 - 7 - 45

18 - 7 - G

K. C. Young

7 - 5 - C1

- 7 - 45

32 - 8 - G

L. Roll

32

15

32 - 8 - L
1

- 7 - 46

13 - 8 - G

- 1 - C1

29 - 3 - C2
C. Einsphar

12 - 3 - C1

H. Gall

14 - 6 - C2

D. Neiman

35 - 8 - C1

- Gli

Unknown

9 - 2 - C1

- 1 - c21

Unknown

16 - 6 - C1

- c21

Unknown

11 - 1 - Cl

13 - 8 - L
2 - 7 - 46

14 - 7 - G
14 - 7 - L

1 - 6 - 46

35

- 1 -G

35 - 1 - L
1 - 7 - 47

9 - 3

1 - 6 - 48

17

1 - 10 - 44

12 - 6

on section divided into 1,2 mile segments running clock wise around
the section from the northeast corner.
Example: the east and west ~ mile
segments would be respectively numbered 3 and 4.
?:...I Cover types: C = grass, L = grass-legume, C1 = farmed roadside, and
C2 = unfarmed roadside.
11 Grass in plot seeded by farmer prior to study initiation.

1/ Location

�April

1970

- 115 -

JOB PROGRESS

Stdte

COLORADO

of
No.

W-37-R-23

Game Bird

Plan No.

3

Job No.

Project
Work

REPORT

Job Title

Effects
Distribution

Period

April

Covered:

Personnel:

Survey

8a

of Sagebrush Control on
and Abundance of Sage Grouse

1, 1969 to September

16, 1969

Clait Braun, Courtney Crawford, Howard Funk, Don Gore,
Don Hoffman, Scott King, Chet McCord, John Monarch, Ken
Morrison, Mike Robertson, Clayton Wetherill,
Terry A.
May (Principal Investigator).

ABSTRACT

Peak strutting ground counts of male sage grouse (centrocercus urophasianus)
totaled 357 birds, the highest count since the experimental
spraying was
completed in 1965. A total of 106 birds were captured and marked, including
28 females fitted with VHF radio transmitters.
These 28 females were monitored a total of 990 days (from April 7 to September 11), including 262
relocations.
Eight of the radio-frequency-marked
females were followed to
nests with a mean distance traveled to nests of 6.29 km (range: 2.19-13.70 km)
from the strutting ground where captured.
Mean clutch size of seven nests
was 7.4 (range: 7-8 eggs).
Poor production,
apparently resulting from cold,
wet weather during the peak period of hatching, caused reduced brood sizes.
Females with broods moved soon after hatching to areas with abundant forbs
which served as food for juveniles.
No definite "travel lanes, to summering
areas were observed, but adequate cover between nesting and meadow areas
appeared to be important.
Females tended to use wet meadows during summer
while males tended to remain in upland sites near moist draws.
Movements
determined from harvested banded birds indicated general movements northwest,
west, and southwest with few birds moving south and southeast from strutting
grounds where banded.
No movements east across Highway 125 could be documented.

�- 116 -

RECOMMENDATIONS

1. Data gathered

in North Park suggested that a habitat manipulation
program resulting in partial kill or partial eradication of sagebrush on upland sites in amounts up to approximately
70 percent did not seriously
affect sage grouse populations during spring and summer months.
In fact,
opening up dense sagebrush areas, permitting forbs and grasses to grow
may improve these areas for summer use.
Until land managers present evidence that partial kill programs can be conducted successfully over large
blocks, sagebrush control programs should be completed in strip patterns.

2. Data on nest locations and distances traveled to nest sites suggested
that sagebrush areas near known summer use areas and wet meadows were
utilized as nesting habitat.
This fact should be considered in developing guidelines for sagebrush control programs.
3. Broods joined into small flocks once juveniles reached approximately
four weeks of age.
All brood census data should be completed within
four weeks of the peak period of hatching to obtain accurate brood counts.
Peak period of hatching can be estimated by adding 47 days to the observed peak of female attendance on strutting grounds.
4. Analysis of harvest data indicated that the sagegrouse population in
North Park could withstand heavier hunting pressure.
If strutting ground
counts are normal and production as indicated by brood counts is good, a
nine day season beginning on the permanent opening date and continuing
through the following weekend with bag and possession limits of three and
six should be considered.
Detailed hunter check data should be accumulated on both weekends and correlated with breeding counts the following
year.
5. Further work with similar objectives by students at this time would probably yield little.
In the future when another student is assigned to
this project in North Park, a permanent employee with the Colorado Division of Game, Fish and Parks should be in close contact with this student,
including considerable direct supervision in the field.
6. Needs for future work include: (a) develop more accurate census techniques for breeding populations,
(b) devise methods to adequately estimate
sex and age structure of the population,
(c) examine fall behavior of
sage grouse to learn if there is a relationship with the high proportion
of females harvested,
(d) experiment with hunting regulations to establish a hunting program designed to give hunters more time in the field
without harm to the sage grouse resource, and (e) investigate the winter
ecology of sage grouse.
Emphasis during future studies should be on
examining the effects of sagebrush eradication on sagegrouse populations
when they are concentrated during the months of November through March.

7.

Similar radio-telemetry
studies, if ever conducted, on sage grouse should
be planned so a crew of two follow no more than 12 instrumented birds
during a given time period.
More realistically,
this number should be
eight if even higher quality observations are desired.

�- 117 -

EFFECTS OF SAGEBRUSH CONTROL ON
DISTRIBUTION AND ABUNDANCE OF SAGE GROUSE
Terry A. May

In 1963, a two year study was initiated to investigate sage grouse
(Centrocercus urophasianus) abundance and distribution in North Park near
Walden, Colorado and to recommend patterns of sagebrush (Artemisia spp.)
control.
In June, 1965, the Bureau of Land Management sprayed approximately
4,000 acres of sagebrush following the design and specifications
recommended
at the termination of the pre-treatment study.
Immediately after spraying,
a post-treatment study was initiated to examine the immediate effects of
sagebrush eradication.
A third study is planned to examine long-term effects
of sagebrush eradication.
Work reported here pertains to the April through September activites of
female sage grouse in and around experimental spray areas and is the second
year of a two year study.
The first year of study was conducted by Poley
(1969).

P. S. OBJECTIVE
To determine the effects of 2,4-D spraying of sagebrush on: (1) sage grouse
abundance: (2) sage grouse distribution, and (3) vegetative composition and
density.

SEGMENT
1.
2.

OBJECTIVES

Measure sage grouse abundance and harvest in North Park, Colorado.
Investigate sage grouse distribution throughout spring and summer.

METHODS

AND MATERIALS

Abundance

and Harvest

An index to spring population level was calculated using counts of strutting
males as described by Gill (1965).
Productivity was estimated by recording
all sightings of unsuccessful and successful females along with the number
of chicks per brood.
Harvest data were obtained at the Cowdrey and Walden
hunter check stations during the opening weekend of the hunting season.
Age
and sex of birds were determined using the methods described by Crunden (1963)
and Dalke et al. (1963).
Population estimates were calculated from the
recovery of banded birds at hunter check stations.

�- 118 -

Capture

and Marking

Birds were trapped on strutting grounds in spring using the "spotlighting"
technique described by Pyrah (1959) while most females were captured using
a mobile cannon-net similar to the one described by Lacher and Lacher (1964).
Females were cannon-netted on primary mating areas just prior to daylight.
A noose-pole described by Zwickel and Bendell (1967) was used wit.h Some
success on juveniles.
All adults were banded with serially numbered, metal leg bands and numbered
plastic bandettes color coded to trapping location as described by Poley
(1969). Juveniles were marked with poultry patagium tags or banded when
they were old enough to retain leg bands.
Selected females were fitted with very high frequency (VHF) radio transmitters obtained from S. L. Markusen, Cloquet, Minnesota.
Transmitters were
attached similar to the methods described by Brander (1968) and Poley (1969).
Batteries with metal "tabs" attached to aid in soldering to lead wires were
also obtained from Markusen.
Harnesses were strengthened by running plastic
coated metal leader through the tubes forming the radio harness as recommended by Poley (1969) (Fig. 1). Instrumented females were monitored using
equipment described by Poley (1969).

Distribution
Distribution and movements were determined from observations of marked birds
and by tracking radio-frequency-marked
females.
Data on banded birds taken
by hunters were recorded at check stations.
General distribution was also
determined from observations of unmarked birds.

DESCRIPTION

OF AREA

Detailed descriptions of the study area were presented by Gill (1965) and
Poley (1969). Carr (1967) estimated a 72 percent kill on sagebrush for all
sprayed areas.

RESULTS AND DISCUSSION
Abundance
Strutting

Ground Counts

Strutting grounds on the study area were located and named by Rogers (1964)
and given code names by Gill (1965) (Fig. 2). Three strutting grounds
(SG-l, SG-9, and SG-lO) were located in strip-spray areas, while two
(SG-2 and SG-5) were located in block-spray areas.

�- 119 -

ANTENNA

TRANSMITTER

TAPE

BATTERY

Arrows are plastic covered leader material
that is inserted into tubes of transmitter.

Fig. 1.

Strengthened

transmitter

harness

for female sage grouse.

�- 122 -

Table 2. Comparison
sage grouse.

of 1960 through

1969 strutting

Year
1964 1965

Strutting
Ground

1960

1961

1962

1963

SG 1 1/

10

5

0

0

0

SG 2 2/

43

4

7

20

SG 3

14

0

0

SG 4

130

96

SG 5 2/

19

SG 6

ground counts of male

1966

1967

1968

1969

0

0

0

0

0

26

9

9

10

2

8

0

0

0

0

0

0

0

219

216

120

53

47

69

58

82

2

52

17

65

52

54

59

52

82

5

1

8

9

3

0

2

12

36

36

SG 7

0

7

5

0

0

0

0

1

3

2

SG 8

5

0

0

0

0

0

6

5

2

6

1/

167

109

71

85

99

52

97

81

71

117

SG 10 1/

17

77

28

19

17

0

6

9

0

16

SG 11

12

50

5

11

7

0

13

15

6

5

SG 12

14

12

0

0

0

0

0

0

1

3

Total

446

363

395

377

337

166

234

265

231

357

SG 9

1/ Strutting
2/ Strutting

grounds
grounds

in strip-spray
in block-spray

areas.
areas.

Counts on major strutting grounds (SG-4, SG-5, and SG-9) increased substantially while counts on smaller strutting grounds (SG-2, SG-8, SG-10, and
SG-12) remained stable or increased only slightly.
Gill (1965) suggested
that higher counts from smaller strutting grounds were actually better indicators of a high population level than counts on large strutting grounds.
Schlatterer (1960), working in Idaho, suggested that subadult females
visited strutting grounds after the peak in hen numbers.
Evidence from ages
of females captured on strutting grounds strongly suggested that subadult
females in Colorado also visited strutting grounds in larger numbers late
in the breeding season.

�- 123 The experimental sagebrush spraying in 1965 apparently did little to cause
relocation or abandonment of strutting areas.
The same strutting areas were
used by similar or greater numbers of birds in 1969 than before the spraying
was accomplished.

Production

Estimates

Productivity was relatively poor in 1969. The first brood was observed on
May 28, while the estimated peak of hatching occurred between June 1-10.
June was unseasonably wet and cold, which apparently resulted in decreased
chick survival and smaller brood sizes.
Large numbers of unsuccessful females
were observed which may have indicated a lower than normal success ratio.
During May, two broods, each having at least three chicks, were observed.
Nine broods with a mean of 5.2 chicks per brood were observed during June,
while 45 females were classified as being unsuccessful.
The mean brood size
dropped to three in July when 18 broods were observed.
After July 7, "gang
broods" were observed which made it difficult to obtain accurate brood counts.
Exchange of chicks between broods probably occurred readily at this time.
Brood counts after July 7 were further complicated because unsuccessful
females also seemed to associate with "gang broods".

Distribution

and Movements

In order to re-observe sage grouse throughout the spring and summer, emphasis
during spring was placed on banding and marking all birds possible.
A total
of 106 birds were captured and marked during 1969 (Table 3). This included
ten recaptures of birds banded prior to 1969.

Table

3.

Age, sex and location

Location

Adult

of sage grouse banded

Male
Subadult

Adult

in 1969.

Female
Subadult

Juvenile

SG 4

20(4)1/

13

5

9

SG 5

0

1

0

0

SG 9

11 (6 )1./

7

8

11

Other areas

3

0

4

7

7

Total

34

21

17

27

7

in parentheses

indicates

1/ Figure

recaptures

of birds banded

prior to 1969.

�- 126 -

Evidence suggested that the same nest sites were used in preceding years.
One nest with all eggs intact had small pieces of egg shells scattered in
the nes t bowl.

Distances

Traveled

to Nests

Eight of the radio-frequency-marked
females
distance of 6.29 km (range 2.19 - 13.70 km)
captured (Table 5). Females did not always
ground on which they were captured as other
passed on the way to nests.

nests were located with a mean
from the strutting grounds where
nest closest to the strutting
major strutting grounds were

Table 5. Distances and directions
sites from strutting grounds. 1/

traveled

by female sage grouse to nest

Banding

Location

SG 4

SG 9

A e
Adult

Subadult

Adult

3.34-WNW

2.l9-SSW

2.50-SSW

4.72-SE

11. 20-NNW

7. IS-NNW

Subadult

13.70-NNW
5.52-WSW

Mean Distances
4.03

1/ Distances

8.15

Traveled

to Nest Sites
4.84

are in kilometers.

Four subadults traveled a mean distance of 8.15 km (range 2.19 - 13.70 km)
to nests, while four adults traveled a mean distance of 4.43 km (range
2.50 - 7.15 km) to nests.
This suggested that nesting experience was important as subadults traveled farther and more variable distances to suitable
nest sites.
Only one female moved southeast from the strutting grounds to nest, while
three moved northwest, two moved southwest, and two moved west.
These data
indicated a general trend of westerly movements from strutting areas to
nesting sites.
Summer concentration areas for females and broods nearest to

�- 127 major strutting grounds were west of these strutting areas.
The experimental spray areas were north and east of these strutting grounds.
Females
appeared to select nest sites near summering areas, thus explaining, in part,
the general trend of westerly movements.
Of the 16 radio-frequency-marked
females not located on nests, five were
unsuccessful nesters, one was a predator victim, and ten were not accounted
for after initial capture or reobservation. Of the ten not classified, it
was probable that most of the transmitters ceased functioning.
It was possible some moved completely out of the study area, but this was probably the
exception rather than the rule. Six transmitters were returned during the
hunting season, and one was retrieved during the summer.
Four of these transmitters had broken antennas, and signals from a transmitter with no antenna
could not be received over 100 m.

Nest Sites
Fourteen nests of the year (Fig. 3) were located during the summer with two
of these being in sprayed areas.
One was in the experimental strip-spray
area, while the other was on a hillside sprayed by a private landowner.
In
both instances live sagebrush occurred within 2 m of the nests.
Of the 14
nests, 12 were under big sagebrush (Artemisia tridentata), one was under a
greasewood (Sarcobatus vermiculatis) bush, and one was under a rabbitbrush
(Chrysothamnus spp.) bush.
The mean height of vegetation over nest cavities
was 350 rom (range 280-430 rom). Relatively flat areas (0-10 percent slope)
with a mean average slope of 7.38 percent (range 0-25 percent) were preferred
for nests.
No preference for aspect was noted, although the largest number
of nests (7) had a southwest aspect.
In all situations, live sagebrush was
within 5 m of nests.
Nesting occurred in medium to tall brush, but nests
were typically in rather open sites or on the edges of open areas.
Frequently
(7 of 12 instances), sagebrush plants over nests were partially dead, but in
all situations, they provided excellent cover.
Nesting was not observed in dense, thick sagebrush areas.
It is my hypothesis
that habitat manipulation programs designed to obtain partial kills on dense
sagebrush areas may Lmprove these areas for nesting by opening up the cover.
Klebenow (1969) reported similar suggestions.

Nesting

Success

Of eight active nests, seven had known clutch sizes (4 with 7 eggs each, and
3 with 8 eggs each).
Three of the nests hatched successfully (Ion June 1
and 2 on June 6) with 85 percent of the eggs (18 of 21) hatching.
Thre~
females deserted their nests which was probably caused by observer interference.
Two nests were destroyed, possibly by some small mammal.
Egg
shells in the nests had small holes in the sides and were scattered around
the nest.
No instances of renesting were documented,
a rare occurrence in 1969.

and this phenomenon

was probably

�- 128 -

Artemisia
Location of Nest to
Sprayed Areas

Sarcobatus

Veg. Over Nes t
Mean Ht. of Veg.
Over Nest = 350 mm

CIl
.IJ
CIl
Q)

Z

~

o

o

z

Ave. Ht. of Veg. in mm Over Nest
N

w

E

S

Aspect
4
CIl
.IJ
CIl

of Nest Sites

Mean Ave. Percent
Slope = 7.36 Percent

3

Q)

Z

~

2

0

.1

0

z

o

Fig. 3.

Fourteen

2

4

6

8 10 12 14 16 18 20 22
Percent Slope of Nest Sites

sage grouse nests

located during

1969.

24

26

Chrysothamnus

�- 129 Brood Movements
Brood movements could be plotted on only one of the three radio-frequencymarked females that were successful (Fig. 4). This female (No. 886) moved
her brood a short distance from the nest into deep sagebrush immediately
after hatching on june 1. This brood remained in the vicinity of the nest
for several days, staying in medium to tall sagebrush.
On the 4th day after
hatching, this brood moved to an irrigation ditch approximate1y.75
km south
of the nest. The brood remained in this area for two days when contact with
the female was lost.
Movements of two successful females captured after their broods had hatched
were plotted during the spring and summer (Fig. 5 and Fig. 6). Female #2016
was captured in a small "pothole" just prior to sunset on June 20. This
"pothole" was an area where open water was present during April and May.
In
late May this water disappeared, and grass and forbs grew in abundance.
"Pothole" areas remained green until late June or early July, and observations of hens with chicks feeding in these areas were quite common.
This
agreed with the findings of 'Klebe now and Gray (1968) and Peterson (1970) who
found that forbs made up a large portion of the diets of juvenile sage grouse.
After six days, female #2016 moved to the Lake Creek area where she roosted
during the day in shrub-dominated sites adjacent to wet areas.
After 2
weeks, this brood moved to the reseeded ryefie1d and spent the remainder of
the summer in that area.
Female #2018 was captured 75 m south of the reseeded ryefie1d on July 9.
This brood spent the summer in the vicinity of the reseeded ryefie1d.
It appeared that hens with broods moved soon after hatching to areas where
moisture was abundant and vegetation was dominated by grass and forbs.
Female sage grouse with broods roosted during mid-day in sites with abundant
shrubs adjacent to wet areas.
These wet areas were typically "potholes"
previously described or near stream bottoms and irrigation ditches.
In all
Situations, forbs were available as food for juvenile sage grouse. Definite
"travel lanes" to Summer areas were not observed; but the amount of protective cover may have been important in determining movement patterns.
Females with broods typically had reached summering areas by the time chicks
were 4 weeks of age. At this time distinct broods were not common, and hens,
both successful and unsuccessful, were observed with chicks in "gang broods".
Considerable exchange of chicks between broods probably occurred at this
time, but juveniles were not radio-frequency-marked
so their movements after
reaching summering areas are not known.

Summering

Areas

Females, both successful and unsuccessful moved into wet areas during summer
while general observations indicated that males remained in upland sagebrush areas near draws with thick sagebrush and some moisture.
Observations
on marked and unmarked females indicated that they remained in or near wet
areas throughout the summer.
One heavy summer concentration area for females
and juveniles was the reseeded wild rye field which had several irrigation

�- 130 ditches running through it. Birds used this area for both roosting and
feeding.
Some birds would roost in the ryefie1d at night while others
would fly about .75-1.25 km south to a greasewood-rabbitbrush
dominated area
to roost at night.
In early September,
sagebrush areas.

Movements

females and juveniles

from Harvested

started moving back into upland

Birds

Movements plotted from banded birds harvested (Fig. 7) were similar to those
determined by tracking radio-frequency-marked
females.
In general, movements were northwest, west, and southwest of strutting grounds.
One adult
female was harvested along the North Platte River east of the strutting
ground where she was banded.
Another adult female was taken approximately
1.6 km north of McFarlane Reservoir approximately 27 km southeast of where
she was banded on the study area. Seasonal movements appeared limited primarily to the vicinity of the study area with some birds moving out of the
area to the south and southwest.
Movements to the north and northwest were
probably limited to the Independence Mountain and Trappers Gulch areas.
There were probably no movements east across Colorado Highway 125.

Harvest

Analysis

A total of 569 sage grouse were checked at the Cowdrey and Walden check
stations on September 13-14 (Table 6). Seventy percent were adults which
reflected poor production in 1969. Only 31.4 percent of the total kill were
males which probably could be explained by availability of birds and by
different behavior patterns between sexes.
Indicated hunting pressure in 1969 increased to a new high with a total of
662 hunters being checked.
Hunters spent slightly more hours (5.16) to
obtain each bird than in past years which resulted in an average of .86
bird bagged per hunter checked.
Twenty-three banded or marked sage grouse were recovered
Of the 23, eleven were banded in 1969, eight were banded
banded in 1967, and one was banded in 1966.

in the bag (Table 7).
in 1968, three were

Using the Lincoln index method to estimate a fall population level before
hunting, a total population of 4,046 birds was obtained for the northwest
quarter of North Park. This included 1,543 males, 1,306 females, and 1,197
juveniles with each estimate being calculated separately.
These estimates
mayor may not be va.':, however, they are not totally unrealistic and have
merit.
Theoretically, populations
patterns such as exhibited

of game birds which have promiscuous breeding
by sage grouse also have high annual turnover.

�- 131 -

Brown (1967) found that male sharp-tailed grouse (Pedioecetes phasianellus)
had an annual turnover that averaged 65 percent.
With similar occurrences
in sage grouse likely, it would appear that the population in North Park
could withstand heavier hunter harvest without serious effects to the sage
grouse population.
Using the population estimate of 4,046 birds, only 14.1
percent of the population in the study area was harvested in 1969. Up to
35-40 percent of this population could probably be taken each year without
harm to breeding populations the following spring.
With an effort to obtain
better hunter dispersion throughout North Park, a management program designed
to increase bag limits and provide longer seasons would be desirable.

LITERATURE

Brander, R. B. 1968. A radio-package
Mgmt. 32(3):630-632.

CITED

harness

for game birds.

J. Wildl.

Brown, R. L. 1967. Sharp-tailed grouse population study. Montana Fish
and Game Dept. Proj. W-9l-R-8, Job No. II-E. Job Completion Rept. 16 p.
Carr, H. D. 1967. Effects of sagebrush control on abundance, distribution
and movements of sage grouse.
Colorado Game, Fish and Parks Dept.
Proj. W-37-R-17, Work Plan 3, Job 8a. Job Compl. Rept. 106 p.
Crunden, C. W. 1963. Age and sex of sage grouse from wings.
Mgmt. 27(4):846-849.

J. Wildl.

Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F.
Schlatterer.
1963. Ecology, productivity, and management of sage
grouse in Idaho. J. Wildl. Mgmt. 27(4):811-841.
Gill, R. B. 1965. Effects of sagebrush control on distribution and abundance of sage grouse.
Colorado Game, Fish and Parks Dept. Proj.
W-37-R-17, Work Plan 3, Job 8a. Job Completion Rept.
185 p.
Klebenow, D. A., and G. M. Gray.
1968. Food habits of juvenile
grouse.
J. Range Mgmt. 21(2):80-83.
Klebenow, D. A. 1969. Sage grouse nesting
J. Wildl. Mgmt.
33(3):649-662.
Lacher, J. R., and Dorothy D. Lacher.
J. Wildl. Mgmt. 28(3):595-597.

1964.

and brood habitat

sage

in Idaho.

A mobile cannon net trap.

Peterson, J. G. 1970. The food habits and summer distribution of juvenile sage grouse in central Montana.
J. Wildl. Mgmt. 34(1):147-155.
Poley, B. E. 1969. Effects of sagebrush control on distribution and abundance of sage grouse.
Colo. Game, Fish and Parks Div. Proj. W-37-R-22,
Work Plan 3, Job No. 8a. Job Compl. Rept. 31 p.

�- 132 -

pyrah, D. B. 1959. Sage grouse population trend study. Sage grouse
trapping study. Wyoming Game Bird Survey. Proj. W-50-R-8, Work Plan
2, Jobs 1 and 2. p. 3&amp;-64.
Rogers, G. E. 1964. Sage grouse investigations in Colorado.
Game, Fish and Parks Dept. Tech. Publ. 16. 132 p.

Colorado

Schlatterer, E. F. 1960. Productivity and movements of a population of
sage grouse in southeastern Idaho. M. S. Thesis. Univ. of Idaho,
Moscow. 87 p.
Zwickel, F. C., and J. F. Bendel1. 1967. A snare for capturing blue
grouse. J. Wildl. Mgmt. 31(1):202-204.

Prepared by
Terry Ad May
Student Assistant

J

�- 133 -

Fig. 4. Movements of female 886 from June 1 to June 6.

�134 -

�135-

�- 136 -

1554

Fig. 7. Harvest locations in relation
sage grouse, September 13-14, 1969.

to banding

1623

location of hunter harvested

�Table 6.

Comparison of North Park sage grouse hunter check information, 1963-1969. II

Total
Birds

Birds
Per
Hunter

Hours
Hunted
Per Bird

58

506

1.03

4.86

45

41

179

.82

3.49

26

26

51

116

.77

5.40

56

45

71

44

263

.86

4.67

127

67

42

46

33

267

.89

4.41

80

135

42

156

141

58

512

.94

5.09

121

277

70

57

114

30

569

.86

5.16

Year

Bag
Limit

Hunters
Checked

Hours
Hunted

Adult
Hales

Adult
Females

Percent
Adults

1963

3

492

2,460

62

150

42

113

181

2

217

624

25

81

59

28

2

150

626

27

30

49

1966

2

306

1,227

31

116

1967

2

300

1,177

50

2

546

2,604

2

662

2,936

1964
1965 ])

1968
1969

Juvenile Juvenile Percent
Males
Females Juveniles

II Based on Walden and Cowdrey check station data during opening weekends.
II A one day season; hence only a one day check.

t-'
w
-....J

�- 138 -

Table 7.

Banded birds checked at the Cowdrey and Walden check stations.

Age

Sex

Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Adult
Juvenile
Adult
Adult
Adult

Fema1eQ.I
Male
Fema1e§j
Female
Male
Male
FemaleS I
Fema1e~:;

Ma1e71
Female§j
Male

Fema1e61
Male
Fema1e£/
Female
Male
Female
Male
Male
?

Female~./
?
?

Banding
Location

Date
Banded

SG 9
SG 9
SG 9
SG 9
SG 9
SG 9
SG 9
SG 9
SG 9
SG 4
SG 4
SG 4
SG 4
SG 4
SG 4
SG 4
SG 4
SG 6
SG 6

April 8, 1969
April 8, 1969
April 8, 1969
April 21, 1969
May 8, 1968
May 29, 1968
April 25, 1968
April 25, 1968
May 1, 1966
April 19, 1969
May 15, 1969
April 17, 1969
April 17, 1969
April 16, 1969
April 10, 1967
May 3, 1967
April 11, 1967
May 2, 1968
April 22, 1968
June 2, 1969
July 8, 1969
August 21, 1968
August 7, 1968

II
21
31
~j

Band
Number

Bandette
Number

Red 83
889
Red 458
1635
Red 80
886
Red 94
2013
Red 441
1603
Red 452
1617
851
Red 302
Red 304
853
Red 451
1466
2012
Yellow 424
Yellow 162
1668
Yellow 416
2005
Yellow 150
1654
Yellow 415
2004
831
Yellow 421
Yellow 195
1554
Yellow 403
835
1593
1577
Patagia1 Tag :/fo 52
White 56
2017
White 20
1623
White 17
867

II North of Lake John 1.2 km.
21 West side of reseeded ryefield.
31 Northwest of Cowdrey-Pearl Road junction with the Hill Ranch Driveway 2.1 km.
41 West .8 km and 150 yards north of southeast corner of Sec. 21, T10N, R81W.
51 Females radio-frequency marked in 1968.
61 Females radio-frequency marked in 1969.

II Male radio-frequency marked in 1966.

�April 1970

- 139 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-37-R-23

Work Plan No.

10

Game Bird Survey
1

Job No.

Job Title

Study of Hungarian

Partridge

Adaptability

Period Covered:

April

Personnel:

Donald M. Hoffman, Charles E. Brown, Gary Brown, David de
Ca1esta, Mark Frasier, Aden M. Greer, James D. Houston,
Roger Lowry, Jon Moser, Dale F. Reed, William I. Roland,
Warren D. Snyder, Henry Wilson and Michael Zgainer.

1, 1969 to March 31, 1970

ABSTRACT

Six hundred and twenty nine more pen raised Hungarian partridge were released in Moffat County on Isles Mountain southwest of Craig, Colorado at
three sites on April 19, 1969.
Intensive field searches were made by W-37-R Project personnel during the
months of June through September, 1969. Only 11 Huns were observed, including a brood of at least four chicks.
This is the first record of a brood
being observed by Division personnel in Moffat County, even though landowners had reported broods earlier.
A winter search of the release sites and surrounding areas in Moffat County
by two Researchers, six W.C.O. Trainees, three W.C.O. IS, and one Area
Supervisor in late February, 1970 resulted in no Huns being observed, no
field sign found, and no new leads secured from landowners interviewed.

�- 140 -

RECOMMENDATIONS

Even though efforts to study
populations were intensified
required and are recommended

the success of field released
in 1969, additional follow-up
to determine this.

Huns to establish
checks will be

Locations in Colorado other than those receiving transplarits to date appear
to have possibilities
as Hungarian partridge range~
It is recommended
these
be checked as trial release sites.
These include areas near Del Norte~
Norwood, Montrose, and Collbran.

�- 141 -

HUNGARIAN

PARTRIDGE

ADAPTABILITY

Donald M. Hoffman

P. S. OBJECTIVE
To determine the ability of Hungarian partridge to establish reproducing
populations in mixed brush1and-wheat1and
habitat in Moffat County, Colorado.

SEGMENT
1.
2.

Introduce
Determine

Hungarian partridge.
presence and dispersal

METHODS

OBJECTIVES

of Huns and success

of previous

plants.

AND MATERIALS

Field releases of pen-raised stock were made with the use of a helicopter
to fly the crated birds from Round Bottom on the north side of the Yampa
River to Isles Mountain south of the river (Figure 1).
Intensive field searches were made by Project W-37-R personnel during the
periods June 11-13, July 14-17, August 5-8, 18-22, 26-29, and September 2-5,
1969. In addition, persons living and working in or near the release site
were interviewed and observations reported were recorded.
A winter search of the release sites and surrounding areas in Moffat County
by 12 Division employees including 2 Researchers, 6 w.c.o. Trainees, 3
w.c.o. IS, and 1 Area Supervisor was made during the last week of February,
1970.

RESULTS AND DISCUSSION
On April 19, 1969, 629 additional pen raised Huns from Fort Collins Wildlife
Research Station and Little Hills Experiment Station holding pens (Figure 2)
were released at 3 sites on Isles Mountain, southwest of Craig, Colorado in
Moffat County.
Fifteen of these were pen raised by the Avian Science Department of Colorado State University and the remainder were pen raised at the
Fort Collins Wildlife Research Station.
A male Hungarian partridge released
in 1969 is shown in Figure 3. Locations, and numbers, of Hungarian partridge
released in Moffat County from 1964 through 1969 are shown in Figure 4. Table
1 summarizes all field releases made in Larimer, Moffat and Routt counties
since 1964. A total of 1,249 Huns have been released with most (1,100) being
released in Moffat County, 135 in Larimer County, and 14 in Routt County.
Table 2 summarizes Hungarian partridge reports and observations for Moffat
County for 1969. Although only 11 Huns were observed during the intensive

�I-'

..p-

N

Fig. 1.

Releasing Hungarian partridge transported
1969.
(D. Domenick, photo).

by helicopter,

to release site on Isles Mountain

southwest

of Craig

�I-'

+:-

w

Fig. 2. Reading band numbers of Hungarian partridge in holding pens at Little Hills Experiment Station,
1969. (D. Domenick, photo).

�~

+=-+=--

Fig. 3.

A male Hungarian partridge released in Moffat County in 1969.

(D. Domenick, photo).

�- 145 R. 91W.

COLORADO
!fI

,.

c

,

~
~

SCALE OF MILES
t

0

"

"

,-

T.6N

.j

,.

,. "
\

'

--~f-·.'

"t-

L

'

/'

Fig. 4. Locations of Hungarian partridge releases, Moffat County,
1964 through 1969.

�- 146 -

Table 1.

Hungarian

Date

Jan.

partridge

Number
Released

&amp; Feb., 1964

field releases

since 1964.

Source of
Stock

Release Area

Oregon &amp; Idaho
(wild trapped)

Larimer County, Douglas
Sec. 1, T8N, R69W

Lake

Sub-total

135

March 28, 1964

40

Idaho
(wild trapped)

Moffat County, Round Bottom
Sec 34, T6N, R92W

March 23, 1965

124

Oregon
(pen raised)

Moffat County, Round Bottom
Sec. 33, T6N, R92W

April

18, 1967

71

FCWRS
(pen raised)

Moffat County, Round Bottom
Sec. 21, T6N, R92W

April 30 and
May 1, 1968

236

FCWRS
(pen raised)

Moffat County, Round Bottom
Sec. 29, T6N, R92W

April 30, 1968

14

FCWRS
(pen raised)

Routt County, Northeast of
Hayden, Sec. 35, T7N, R87W

April

19, 1969

389

FCWRS (374) &amp;
CSU (15)
(pen-raised)

Moffat County, Isles
Mountain, Sec. 24 &amp; 25,
T5N, R92W

April

19, 1969

240

FCWRS
(pen raised)

Moffat County, Isles
Mountain, Sec. 16, T5N, R92H

Sub-total

Total

1,114

1,249

An additional 22 birds (11 pairs) were taken to Rocky Ford Experimental Bird
farm from this shipment.
On October 6, 1965, 51 (pen-raised) birds were received
from Oregon and were taken to the Fort Collins Wildlife Research Station for
experimental breeding purposes.
On February 19, 1970, 13 1969-hatched Huns from
eggs secured from wild-trapped birds in England were received from Winchester
Farms, Alton, Illinois and will be used for experimental breeding purposes in
1970 and later years.

&gt;',

�- 147 -

Table 2.

Summary

Date

partridge

Number Huns

Late Winter,
Spring

of Hungarian

1969

1969

Sununer 1969

July 16, 1969

August

5, 1969

Area

Moffat

County,

Observed

1969.

By

Several

Round Bottom area

L. Ellgen

2 (1 pair)

Isles Mountain,

BLM catchment

G. Hilton

4 (2 pair)

Isles Mountain, saddle south
of BLM catchment

G. Hilton

Few

Round Bottom, Fuhr Gulch

J. Counts

2 (1 pair)

Isles Mountain, south of
Meade Ranch headquarters

Meade Brothers

Isles Mountain, north of
Meade Ranch headquarters

Meade Brothers

2 (1 pair)

Isles Mountain

area

Meade Brothers

4

Isles Mountain,

Bowers

2 (1 pair)

Isles Mountain, Meade
Brothers property

D. de Colesta

Isles Mountain, Meade
Brothers property

D. de Colesta

6 (3 pair)

July 15, 1969

reports and observation,

2 (1 pair)

fields

B. Bowers

4 chicks
(~ grown)

Isles Mountain,

Bowers

fields

D. de Colesta

3 broods

Isles Mountain,

Bowers

fields

B. Bowers

Augus t 7, 1969

2 (1 pair)

Isles Mountain,

Bowers

fields

D. de Colesta

August

1 adult, male

Isles Mountain, Meade
Brothers property

August

October

6, 1969

19, 1969

20, 1969

60 to 80

Cedar Mountain

D. de Colesta
Hunters report
to D. Reed.

�- 148 -

field searches conducted in the summer of 1969, a brood of at least 4 chicks
was located.
This is the first record of a brood being observed by Division
personnel in Moffat County.
Past reports of peaks of Hun numbers sighted by land owners on Isles Mountain have coincided closely with wheat harvests.
This occurs normally in
late August in Moffat County.
Some of these reports of Huns may have been
confused with sharp-tailed grouse based upon observations made in 1969.
From 27 to 29 sharp-tailed grouse in 4 different groups were observed by D.
de Co1esta on or near wheat fields during the period August 16-29, 1969.
Three of these locations were the same or nearly the same as where groups
of Huns had been reported in recent years, mostly during wheat harvests.
Only 1 Hun was observed by de Co1esta during this same period.
It appears
that sharp-tailed grouse congregate in the wheat fields at harvest time and
may be difficult to distinguish from Huns by untrained observers.
There
exists a strong possibility that some of the past Hun reports on Isles Mountain were actually sharp-tailed grouse.
One report of 60 to 80 Huns seen in the Cedar Mountain area northwest of
Craig about October 20, 1969 was secured from hunters by Dale F. Reed. A
winter search of the release sites and surrounding areas including Cedar
Mountain was made by 12 Division employees during the last week of February
1970. No Huns were observed, no field sign found and no new leads were
secured from land owners interviewed.

Prepared

by

_M~~~='
~CWJ~-~~/.
~-~l

Donald M. Hoffma~
Wildlife Researcher

+~...:...~::.-:==-=~

�Aprll

l~/U

- 149 -

JOB PROGRESS REPORT

State of

~CO~L~O~RA~D~O~

Project No.

W-37-R-23

Work Plan No.

10

_
Game Bird Survey

Breeding

Job No.

2

of Hungarian

Partridge

Job Title

Experimental

Period Covered:

April 1, 1969 to March 31, 1970

Personnel:

Lawrence A. Webster, Robert L. Schmidt, John F. Corey,
Dr. John V. Shutze, Dr. Harry D. Muller, and Donald M.
Hoffman

ABSTRACT

Twenty more eggs (664 compared with 644) were laid by the breeders using
the 22 percent protein feed in comparison with the 28 percent protein feed
but more of the 22 percent protein feed eggs were also infertile (178 compared with 118). A higher percent hatch resulted from the 28 percent protein feed (53.11% compared with 47.14%) considering total eggs, and a
slightly higher percent hatch resulted from the 28 percent protein feed
(65.02% compared with 64.40%) considering fertile eggs. A higher percent
survival for the first seven days was found using the 28 percent protein
feed (92.92%) in comparison with the standard 22 percent protein feed
(89.93%).
More fertile eggs were produced in the six ground floored pens using 22
percent protein feed than in the wire floored pens (223 compared with 167)
and more fertile eggs were produced in the six ground floored pens using
28 percent protein feed than in the wire floored pens (176 compared with
169).

�- 150 -

RECOMMENDATIONS

With two years data comparing 28 percent protein game bird breeder feed
with the standard 22 percent feed and no clear advantage indicated in
using one over the other, it is recommended that the 1970 test series
compare 16 percent feed with 22 percent feed.
The wire-floored
series of breeder pens are adaptable for controlled tests
to compare the effects of extra lighting on fertile egg production due to
their construction
and location.
It is recommended that the necessary
wiring be done and these tests started in 1970.
The Huns received from Winchester Farms, Alton, Illinois will be held
separately in breeding pens in 1970 so that characteristics
may be observed
and measured toward improving our present stock for field survival.
Production

cost records

for Hungarian

partridge

will

be maintained

in 1970.

�- 151 -

EXPERIMENTAL

BREEDING

OF HUNGARIAN

PARTRIDGE

Donald M. Hoffman

Good progress continued in 1969 in improving production techniques for Hungarian partridge at the Fort Collins Wildlife Research Station.
From the 52
pairs (104 birds) held as breeders in 1969, there were enough birds for use
as breeders in 1970 (43 pairs) plus 674 surplus Huns which will be released
at 3 sites in Moffat County in April, 1970. Thirteen (5 hens and 8 cocks)
1969-hatch Huns were received from Winchester Farms, Alton, Illinois, on
February 19, 1970. Ten (5 pairs) of these will also be used for breeding
pairs in 1970, bringing the total number of breeding pairs for 1970 to 53.
The breeders used in 1969 were selected from the second and third hatches
and all were 1968-hatch birds.
The number of birds pen-raised at the Fort
Collins Wildlife Research Station to be released in the field (674) in 1970,
compares favorably with the 614 from this source released in 1969 and 250
released in 1968. See Table 1, Work Plan 10, Job 1, Job Progress Report for
numbers of birds released in the field from 1964 through 1969.

P. S. OBJECTIVE
To develop

game farm production

techniques

SEGMENT
1.

for Hungarian

OBJECTIVES

To measure the effects of 2 levels of protein
(22% versus 28%) on:
(a)
(b)
(c)
(d)

partridge.

in the breeder

feed

Egg production
Egg fertility
Egg hatchability
Chick survival for the first 7 days

2.

To measure the relationship of holding breeders
wire to fertile egg production.

3.

To compare the number of fertile eggs produced by early mating
with those mating late in the breeding season.

METHODS

on the ground and on

pairs

AND MATERIALS

The detailed arrangement of the Hungarian partridge breeding pens for 1969
is shown in Figure 1. Mated pairs for all except the late mated series
were selected and caught in the large Hungarian partrige holding pens
(Figure 2). Figure 3 shows the arrangement of these holding pens. Table 1
lists the pen schedule for the various tests run in 1969. One mated pair

�- 152 -

N

O£ 62 S2 L2 92

75'

6'4"

"11,7[\
co
1
fC')
fI'l

Ground-floored

Sorie,

37 38 39 40 41 42 43 44 45 46 47 4£

'v _a._ 'v _

"'--"1/_'-

"v _ ,--,-'\,11' _ '-- "l/ -,~

19'

6'4"

Wiro-floorod

Sories

v
C\I

b

rt)

C\I
C\I
C\I

CD

0

~

49~

51

Pens 4'-::9:-'"-';;-:51;-¥

--~

~--

(not used

in 1969)
SCALE e

I

en

I

I

0' 5' 10'

I

25'

Fig. 1.
U&gt;
L

.~t- -

Detailed Arrangement

_

r~---

~'G'-~
.__._-_
..
-- 48'----

Co

Partridgo

Breeding

of. Hunoarlan

Pene,

�t-'

VI
W

Fig. 2. Hand netting Hungarian partridge breeder pair following selection.
collection of pairs. (D. Hoffman photo).

The small pen is used to facilitate

�- 154 -

Covered
Brooder

Shelter

1

Houses

.r &gt;:

L-..-...L.--J--_"--_-1 /'_-I-_--'-_---J4'~8'-4

J

f

-0
t\J

4'

N

-

10

o

(\J

C\I
N

~ 15.5
---35
-t

,~15.5-l
---

~--------------I05'-----------,
Fig. 3.
Hungarian

Partr idge

Holdin CJ Pe ns

�- 155 Table

1.

Pen schedule

Pen
Number

Protein

22%

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48

T

Number Pens on

18

Key:

for

Hungarian

Level

28%

8'X8' 8' X 16'

T

0

6' X33'

x
x
x
x
x
x
x
x
x
x
x
x

T
T
T

T

Pairs
Early

Mated
Late

T
T
T
T
T
T

x
x
x
x
x
x

T
T
T
T
T
T
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x

T
T
T
T
T
T
T
T
T
T
T
T
T
T
T
T
T
T

Pen Floor
Ground
Wire
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
T
T
T
T
T
T
T
T
T
T
T
T

x
x
x
x
x
x
x
x
x
x
x

T
T
T
T
T
T
T
T
T
T

T == Pen on Test;

1969

Pen Size

T

T
x
x
x
x
x
x
x
x
x
x
x
x

partridge,

T
T
T
T
T
T
T
T
T
T
T
T

18

0

0

x == Pen not

on Test

0

6

6

12

12

�- 156 -

of Hungarian partridge was used in each pen. Pens 1 through 36 are located
in a converted nursery shadehouse with ground floors, and pens 37 through
48 are a separate series of wire floored experimental game bird breeding
pens.
Alternating sections of snowfence slats were removed from the north portion
of the shadehouse prior to the 1969 tests. Based upon encouraging results,
as measured in numbers of fertile eggs, the remaining alternate sections
from the south portion were removed in March 1970, prior to the 1970 test
series.

Protein Levels

in Feed

Pens 1-6 and 19-48 were used to test the effect of 2 levels of protein in
the feed. Birds in even numbered pens were fed a standard game breeders feed
containing 22 percent protein and those in odd numbered pens were fed a
similar feed, but containing 28 percent protein.
Eggs were marked and placed
in separate incubator drawers to facilitate determination of (1) fertile
eggs produced; (2) number of fertile eggs that hatched, and (3) numbers of
surviving chicks.
Chicks were held separately in the brooder until they
were 7 days of age to detect differences in survival.

Type of Floors
Pens 25-36 have ground floors and pens 37-48 have wire floors.
Fertile egg
production from the mated pairs placed in even numbered pens (22% protein
feed) on the ground were compared with even numbered pens on wire and odd
numbered pens (28% protein feed) on the ground were compared with odd numbered pens on wire.

Period of Mating
In 1969, pairs of early mated Hungarian partridge were placed in pens 8, 10,
12, 14, 16, and 18, on March 17, 1969, approximately 8 weeks after pairing
had first been observed in the runs. An additional twelve cocks from 1
holding pen and 12 hens from another holding pen were held in separate pens
from March 17, 1969 to April 2, 1969. It is not known if these birds were
mated prior to catching.
These 24 birds (12 cocks and 12 hens) were placed
in a common pen on April 2, 1969, and almost immediately began pairing.
Six
pairs of these later mated Huns were selected from these 24 birds and placed
in pens 7, 9, 11, 13, 15, and 17. Fertile eggs produced by Huns in early
mated pens (numbers 8, 10, 12, 14, 16, and 18) were compared with those from
the late mated pens (numbers 7, 9, 11, 13, 15, and 17). All pens were fed
22% protein feed.
In preparation for the 1970 tests, 6 pairs used for earliest mating pairs
were selected as soon as the birds were observed to start pairing in the
large runs (February 6, 1970) and the 6 pairs used for latest mating pairs
were selected from 10 unpaired hens and 10 unpaired cocks held in separate
pens from February 6, 1970 to March 23, 1970. These began to pair almost
immediately when recombined on March 23, 1970 and pairs were selected later
on the same date.

�- 157 -

RESULTS AND DISCUSSION
Protein Levels

in Feed

Table 2 lists a comparison of numbers of eggs produced and percent hatch of
eggs from breeder pairs in 18 pens using the standard 22 percent protein
level feed with 18 breeder pairs using a similar feed but containing 28 percent protein.
Twenty more eggs (664 compared with 644) were laid by the
breeders using the 22 percent protein feed in comparison with the 28 percent
feed, but more of the 28 percent protein feed eggs were fertile (526 compared
with 486).
Percent fertile eggs was 81.67 for the 28 percent protein feed,
and only 73.19 for the 22 percent protein feed. A higher percent hatch, considering total eggs, resulted from the 28 percent protein feed (53.11% compared with 47.14%) and an also slightly higher percent hatch, considering
fertile eggs only, resulted from the 28 percent protein feed (65.02% compared with 64.40%).
Table 3 lists a comparison of chick survival during the first 7 days on
these 2 different protein levels in the feed. All of the chicks from pens
fed the 22 percent protein feed were combined in this test so that 556 chicks
on this ration were used in comparison with 325 chicks on 28 percent protein
feed. A higher percent survival for the first 7 days was found using the 28
percent protein feed (92.92%) in comparison with the standard 22 percent
protein feed (89.93%) indicating the 28 percent protein feed was better than
the 22 percent.
Results in 1969 were directly opposite of those found in
1968 when it appeared the 22 percent protein feed was slightly better for
Hungarian partridge chick survival.
Birds in several of the pens died during the 1969 test series including a
cock in pen number 2 on May 6, 1969, a hen in pen number 7 on May 21, 1969,
and a hen in pen number 26 on July 2, 1969.
Pairs from alleyways using similar protein level feeds were used to replace the first 2 of these and the
last was not replaced due to the lateness of the laying season.
Several pens
had poor layers but this was not unusual.

Type of Floors
Table 4 lists a comparison of numbers of fertile eggs produced and percent
hatch of fertile eggs from 6 pens with wire floors and 6 pens with ground
floors using either 22 or 28 percent protein feeds. More fertile eggs were
produced in the 6 ground floored pens using 22 percent protein feed than in
the wire floored pens (223 compared with 167) and more fertile eggs were
produced in 6 ground floored pens using 28 percent protein feed (176 compared
with 169).
The ground floored pens used in this test were again all located in the north
portion of the old shadehouse, but every other section of snow fence was
removed prior to the 1969 tests. The ground floored pens were therefore
still partially shaded, while the wire floored pens had shade only from the
sides and shelters located in the north ends.

�Table 20 Numbers of eggs produced and hatchability of eggs laid by Hungarian partridge on two levels of
protein in the feed, 1969
0

Protein
Level

Number
Pens

Number
Eggs
Produced

Number
Infertile
Eggs

Number
Fertile
Eggs

Percent
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

22%

18

664

178

486

73019

313

47014

28%

18

644

118

526

81067

342

53011

Table 3

0

Percent
Hatch
Fert ile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

64040

20

153

9

65002

13

171

12

Survival of Hungarian partridge on two levels of protein in the feed, 1969

0

Protein
Level

Number
Pens

Number Chicks
Taken to
Brooder House

Number
Chicks Lost
First Seven Days

22%

34

556

56

500

89 93

28%

18

325

23

302

92 92

Number Chicks
Survived
Seven Days

Percent
Survival
First Seven Days

0

0

t-'

VI

00

�- 159 A comparison of numbers of fertile eggs laid by the breeders in pens 25-36
in the north section of the shadehouse in 1969 with alternating sections of
slats removed (399) with 1968 when the section was covered with the slats
(255) indicates that removal of half of the slats had considerable value.

Period of Mating
Table 5 lists a comparison of numbers of fertile eggs produced and percent
hatch of fertile eggs from 6 pens with early mating pairs and 6 pens with
late mating pairs, all using 22 percent protein feed. More fertile eggs
were produced by the early mating pairs (200 compared with 178) but a higher
percent of the fertile eggs from the late mating pairs hatched (75.28 percent compared with 58.00 percent).
This resulted in a higher number (134)
chicks hatched from the late mating pairs compared with the number of chicks
hatched from the early mating pairs (116).
This test cannot be considered a good one because of selection procedures
used.
Early mating pairs were selected on March 17, 1969, approximately 8
weeks after pairing had first been observed in the runs. Twelve cocks from
one run and 12 hens from a different run, also selected on March 17, 1969,
were held in separate pens until April 2, 1969. These were then allowed to
pair off and were used for late mating pairs.
It is unknown if all or part
of the late mating pairs selected had been paired earlier to other mates.
Also it is not known if the 6 pairs of early mating Huns were actually the
earliest to pair since approximately 8 weeks elapsed from the time the birds
were first observed to start pairing and the time they were selected for
breeders.
In 1970, earliest mating pairs will be selected for comparing
numbers of fertile eggs produced with later mating pairs and a better test
should result.

/

/

Prepared

by

I

, : ,/

J;/

•

':

~/.,---:,~/:-: .«

~-- __--~--------~------Donald M. Hoffman
Wildlife Researcher

/

�Table 4.

A comparison of eggs produced by Hungarian partridge held on wire with those held on the ground.

Treatment

Percent
Protein
Level

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Wire floors

22

6

262

88

28

6

219

22

6

28

6

Ground
floors

Number
Dead Number
Germs
Culls

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

167

103

39.31

61.67

4

60

3

45

169

96

43.84

56.80

3

70

5

236

15

223

150

63.56

67.26

14

59

2

200

22

176

133

66.50

75.57

6

37

4

Number
Fertile
Eggs

t-'

0-

0

Table 5. A comparison of eggs produced by Hungarian partridge breeders of early mating versus late matingo

Treatment

Percent
Protein
Level

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Number
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

Early mating

22%

6

242

40

200

116

47.93

58000

11

72

2

Late mating

22%

6

191

11

178

134

70.15

75.28

8

41

2

�April,

1970

- 161 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

.W-37-R-23

Work Plan No.

12

Job Title

Game Bird Survey
10

Job No.

Relationships of the Productivity and Distribution of
Wild Turkeys on the Uncompahgre Plateau to the Environment

Period Covered:

April 1, 1969 to March 31, 1970

P. S. OBJECTIVES
1. To determine the size, distribution, and productivity of the wild
turkey
population on the Uncompahgre Plateau.
2. Record movements of wild turkeys and determine how environmental factors
affect these movements.
3. Determine the sex and age structure of the turkey kill on the Uncompahgre
Plateau to provide estimates of total harvest and size of the turkey
population.
4. Determine the vegetative types present on the wild turkey summer and
winter range and the species composition and abundance of food producing species within each type. Also determine the elevation, describe
topography, and record certain climatic factors on part of the better
turkey summer and winter range.

SEGMENT OBJECTIVE
To compile,

analyze,

summarize,

and publish

information.

Progress
Field work was completed on October 20, 1967. All data have been summarized,
and most data have been analyzed.
A manuscript containing these data and
results from Work Plan 12, Job 14 is being edited and revised.

Future Plans
The manuscript will be finished in Segment
be in the form of a technical bulletin.

Prepared

24.

Publication

will probably

/1

}

l
~;t&lt;~._·_L-_.~~~
_

by __~_._~_-~~\~'~~/_'~/~·
__
Gary T-~ Myers
.
Wildlife Researcher

��April

1970

- 163 -

JOB FINAL REPORT

State of

COLORADO

Project No.

W-37-R-23

Work Plan No.

12

Game Bird Survey
13

Job No.

Job Title

Experimental

Period Covered:

April

Habitat

Improvement

1, 1969 to March

for Wild Turkeys

31, 1970

p. S. OBJECTIVE

To compare the effectiveness of (1) food plots and (2) supplemental
feeding stations in increasing turkey harvests.

SEGMENT OBJECTIVES

To compile, analyze,

summarize,

and publish

information.

ABSTRACT

A IS-page paper was prepared and presented at the Second National Turkey
Symposium, Columbia, Missouri, February 11 and 12, using results presented
in the Job Final Report for this job, April, 1969, pp. 111-151.

Reference:
Hoffman, Donald M. 1970. Problems involved with present habitat
management for Merriam's turkeys in Colorado.
Second National
Turkey Symposium, Columbia, Missouri. February 11-12, 1970.

Prepared

by

f)~!fl~~

Donald ~ffman
Wildlife

Researcher

v

��April 1970

- 165 -

JOB PROGRESS REPORT

State of

COLORADO
----------------------------

Project No.

W-37-R-23

Work Plan No.

12

Game Bird Survey
14

Job No.

Job Title

Use of Food Plots to Concentrate

Period Covered:

April 1, 1969 through December

Wild Turkeys
2, 1970

p. S. OBJECTIVES
Determine the effects of small food plots on fall concentrations
harvest of wild turkeys.

and

SEGMENT OBJECTIVE
To compile,

analyze,

summarize,

and publish

information.

Progress
Field work was completed December 2, 1968. Data were analyzed, written up,
and incorporated in a manuscript containing information gathered under
W-37-R Work Plan 12, Job 10. This manuscript is being edited and revised.

Future Plans
The manuscript will be finished in Segment
be in the form of a technical bulletin.

24.

Prepared by

Publication

'-'- .---;

~-'--:\

will probably

J} . 1L."
.,
::.!--.( (.&lt;L-

,'1
...........•

'-'

Gary T. Myers
'J
Wildlife Researcher

��April 1970

- 167 -

JOB PROGRESS

State of

REPORT

COLORADO
----------~~~-----------

Project No.

W-37-R-23

Work Plan No.

15

Game Bird Survey
Job No.

Job Title

Study of Mountain

Period Covered:

April 1, 1969 to March 31, 1970

Personnel:

Donald M. Hoffman,

2

Quail Adaptability

Ronald B. Arant and Torn W. Barnes

ABSTRACT

No additional mountain quail releases were made in Colorado during this
segment.
Young mountain quail produced at the Fort Collins Wildlife
Research Station in 1969 are being held for breeding stock in 1970 toward
future field releases.
A total of 372 wild-trapped mountain quail from
California and Oregon have been re1eas~d on the west side of the
Uncompahgre Plateau from 1965 through 1968. Although no additional
mountain quail sightings or reports were secured in a one week field search
of the release sites and vicinities on the west side of the Uncompahgre
Plateau in the summer of 1969, a covey of 20 mountain quail were observed
on the South Fork of Mesa Creek early in 1970 by Conservation Aide, Torn
Barnes of Nucla, Colorado.

��- 169 -

STUDY OF MOUNTAIN

QUAIL ADAPTABILITY

Donald M. Hoffman

P. S. OBJECTIVE
To determine the ability of mountain quail to establish
lations in areas of mixed brush, pinon-juniper, conifer
Uncompahgre Plateau.

SEGMENT
1.
2.

Introduce mountain
Determine presence

quail.
and dispersal

METHODS

reproducing popuand aspen on the

OBJECTIVES

of mountain

quail.

AND MATERIALS

A 1 week search of release sites on the western side of the Uncompahgre
Plateau was made in August, 1969. Individuals living and working in or
near the release sites were contacted and asked to report any sightings.

RESULTS AND DISCUSSION
No additional wild trapped mountain quail were obtained from other states.
Good progress was made in rearing pen-raised mountain quail at the Fort
Collins Wildlife Research Station in 1969, but the lack of production during
1968 did not allow for additional field releases.
The adult breeders, the
1969-hatched birds, and 12 wild trapped birds secured from Oregon on
January 18, 1970, are being held to propagate the species toward future
field releases.
Table 1 lists all past field releases and sources of stock
being held at the Fort Collins Wildlife Research Station for experimental
propagation purposes.
No additional mountain quail sightings were secured in a 1 week search of
release sites and vicinities on the west side of the Uncompahgre Plateau.
A covey of 20 mountain quail were observed by Conservation Aide Tom Barnes
on the South Fork of Mesa Creek early in 1970. Follow-up checks on this
report are planned for Segment 24.

Prepared

by

j~m~~

Donald M. offman
Wildlife Researcher

�Table 1. Summary of mountain quail field releases.

Release
Area

Number
Birds

Source of
Birds

August 19 and 25, 1965

143~'~

California
(wild trapped)

Mesa County, Indian Creek,
Sec. 27, Rl7W, T51N

August 20, 1966

163

California
(wild trapped)

Mesa County, Indian Creek
approximately 4 miles west
of 1965 releases

January 25, 1968

66

Oregon
(wild trapped)

Mesa County, Indian Creek,
approximately I mile east
of west Uncompahgre National
Forest boundary

Date

t-'
-...J

Total

372

* Sixteen additional mountain quail from these shipments were taken to Fort Collins Wildlife Research
Station for experimental propagation studies. On January 18, 1970, 12 Wild-trapped mountain quail
from Oregon were added to the breeding stock at the Fort Collins Wildlife Research Station. These
constitute the source of our breeding stock.

o

�April 1970

- 171 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-37-R-23

Work Plan No.

15

Game Bird Survey

Breeding

Job No.

3

of Mountain

Quail

Job Title

Experimental

Period Covered:

April 1, 1969 to March 31, 1970

Personnel:

Lawrence A. Webster, Robert L. Schmidt,
and Donald M. Hoffman

John F. Corey

ABSTRACT

Good progress was made in the experimental game farm propagation of mountain
quail in 1969, with~OO
eggs being laid by 28 breeders of mixed sexes
(18 hens and 10 cocks) and 68 young hatched, compared with no egg production
in 1968. Of the 68 young hatched in 1969, there are 45 healthy birds to be
used in the 1970 tests, plus 23 older breeders at the Fort Collins Wildlife
Research Station.
In addition, twelve wild trapped mountain quail from
Oregon were added to this breeding stock on January 18, 1970. Ten of these
wild trapped birds have survived.
.
A test to compare fertile egg production by three selected groups of two
hens and one cock each, with a community pen of six hens and three cocks
cannot be considered a valid test due to stresses induced on the breeders
by holding them in wire floored pens because of inadequate ground floored
facilities.

�- 172 -

RECOMMENDATIONS
The 1969 tests indicate that mountain quail breeders should not be held
in wire floored pens with one-half inch hardware cloth bottoms because
they soon become sore footed.
Earlier tests showed that they should not
be over-wintered in wire floored pens for the same reason.
Because of
stresses induced on breeders following placing in wire floored pens, the
testing of fertile egg production of three selected groups of two hens
and one cock each, with a community pen of six hens and three cocks cannot
be considered valid.
This test will be rerun as soon as adequate ground
floored facilities are built. A series of ten experimental ground floored
breeding pens designed similar to the present tinamou breeding pens will
be built and used for tests in 1970.
Breeders used in the 1970 tests should all be selected from 1969 hatched
birds to eliminate the variable encountered in using different age breeders.

�- 173 -

EXPERIMENTAL

BREEDING

OF MOUNTAIN

QUAIL

Donald M. Hoffman

P. S. OBJECTIVE
To develop

game farm production

techniques

SEGMENT
To measure the relationship
fertile egg production.

for mountain

OBJECTIVE

of flock mating and selected

METHODS

quail.

groups mating

to

AND MATERIALS

The wire floored quail pens number Q 1 and Q 3 were divided into 2 approximately equal sized pens each with a wood framework and 1" chicken wire.
Selected groups of 1967-hatched birds or earlier hatched birds, consisting
of 2 hens and 1 cock were placed in each of these 4 pens of equal size and
a flock of 8 hens and 4 cocks were placed in the larger undivided pen Q 2
on March 24, 1969. On April 2, 1969, after only 10 days confinement on
wire, 10 of the 12 mountain quail in pen Q 2 had extremely sore and bleeding
feet. Several of these birds on wire had started to develop a white diarrhea,
apparently due to stress conditions.
Some of the birds in the selected
groups of 2 hens and 1 cock each (4 pens) had also started to exhibit these
symptoms of stress, although not as many as in the more crowded community
breeding pen. "All birds were then moved to the ground flQored quail runs
shown in Figure 1. Pens 1, 3, and 5 contained selected groups of 2 hens
and 1 cock each, and pen 6 contained a community of 6 hens and 3'cocks.
The surplus breeders (5 sore footed hens plus 1 cock) were held in pen 4
and 1 hen plus 3 cocks were held in pen 2. Table 1 lists composition of
breeding pens for the 1969 tests.

RESULTS AND DISCUSSION
Table 2 lists total numbers of mountain quail eggs produced and numbers of
fertile eggs produced by individual pens in 1969. Although good progress
was made in 1969 in propagating mountain quail with 100 eggs being laid by
28 breeders of mixed sexes (18 hens and 10 cocks) and 68 young mountain
quail hatched, the test to compare fertile egg production of 3 selected
groups of 2 hens and 1 cock each, with a community pen of 6 hens and 3 cocks
cannot be considered a valid one. This was due to (1) stresses induced on
the breeders by confinement on wire, causing sore footed symptoms and death
in some cases, (2) stresses induced by additional handling of breeders necessary in order to replace losses, and (3) stresses induced on the selected

�I-'
-...J

00

Figure 3 - A group of 1969-hatched mountain quail at the Fort Collins Wildlife Research Station.
(D. Hoffman photo).

�I'
I-'

"-I

\.0

Fig. 4.

A close-up of a mature mountain quail hen.

(D. Hoffman, photo)

��April

1970

- 181 -

JOB FINAL

State

Work

COLORADO

of

Project

No.

W-37-R

Plan No.

17

Job Title
Period

REPORT

Covered:

Personnel:

Game Bird Survey
Job No.

Inventory
April

of Ptarmigan

1, 1965 to March

Clait E.
Schmidt,
Division
Students

1

Populations

31, 1970

Braun, Terry A. May, Glenn E. Rogers, Raymond K.
Wildlife Conservation
Officers of the Colorado
of Game, Fish and Parks; Ronald A. Ryder and
of Colorado State University.

ABSTRACT

Distribution
and populations of white-tailed
ptarmigan (Lagopus leucurus)
in Colorado were investigated, with major emphasis on comparing population
levels of hunted and unhunted areas.
Study locations were selected in
alpine areas of Rocky Mountain National Park, at Mt. Evans and Crown Point
in north-central
Colorado, at Independence Pass in central Colorado, and
at Mesa Seco in the southern part of the State.
Ptarmigan were found to
be distributed throughout the approximately
3,750 square miles of alpine
habitat.
Techniques were developed to accurately census breeding pairs
and to locate hens with broods.
Criteria were established based on pigmentation of the ninth primary to accurately separate adult and subadult
ptarmigan.
Descriptions
from which the sex of ptarmigan could be reliably
determined from late April to mid-September
were prepared, but no method
was found using external characters to reliably determine sex when birds
were in winter plumage.
Breeding dens i t i e s ranged from a low of six birds
per square mile on one hunted area to about 30 birds per square mile on
the unhunted area.
Breeding densities appeared to be related to quality
of habitat, although they may have been suppressed by hunting on one area;
however, hunting did not significantly
reduce breeding densities during
the period studied.
Timing of nesting activities varied with area and year
and were directly related to climatic conditions in June and early July.
Nesting success and production varied tremendously being higher in years of
mild climatic conditions in June and early July.
Fall population densities
ranged from less than 12 birds per square mile on one hunted area to as
high as 60 birds per square mile on the unhunted study area.
Exploitation
of fall populations up to 40 percent had no apparent effect on the population of one hunted area, but the annual exploitation
of over 50 percent of
the fall population on another hunted area may have suppressed breeding
populations.
Two hunted populations were maintained by immigration from
surrounding areas.

�- 182 -

RECOMMENDA TIONS

1. Techniques for aging and sexing white-tailed
ferred in usable form to Game Planning.

ptarmigan

should be trans-

2. Techniques for census of ptarmigan populations during breeding
brood periods should be made available to Game Planning.

and

3. Continued research on population densities and effects of hunting should
be conducted at Mt. Evans, Independence Pass, Crown Point, with Rocky
Mountain National Park being retained as a control
4. Population

studies at Mesa Seco should be discontinued.

5. Research should be emphasized exploring the relationship between June
and early July climatic conditions, nesting success, and fall production.

�- 183 -

INVENTORY

OF WHITE-TAILED

Clait

PTARMIGAN

POPULATIONS

E. Braun and Glenn E. Rogers

P. S. OBJECTIVE
To inventory

white-tailed

ptarmigan

RESULTS

in Colorado.

AND DISCUSSION

The typed manuscript of the final report was prepared, reviewed and submitted to Dr. Lee Yeager for final editing.
This manuscript has been
approved for publication as a Technical Publication of the Colorado Division
of Game, Fish and Parks.
The title will be "The White-tailed
Ptarmigan in
Colorado".

Prepared

by

----=6~V::=........:-- -..::~:....:....'
-L.~-=---=:':'_.
Clait E. Braun
Assistant Wildlife

Researcher

_

��April

1970

- 185 -

JOB PROGRESS REPORT

State of

COLORADO
----------~~--~----------

Project No.

W-37-R-23

Work Plan No.

19

Job Title

Experimental

Game Bird Survey
Job No.
Breeding

1

of Tinamou

Period Covered:

April

1, 1969 to March 31, 1970

Personnel:

Lawrence A. Webster, Robert L. Schmidt,
and Donald M. Hoffman

John F. Corey

ABSTRACT

A total of 493 eggs was laid by the 36 tinamou breeders consisting of 24
hens and 12 cocks on hand in May, 1969. Of these, 59 were culls, 166 were
infertile, 101 were dead germs, 29 pipped but did not hatch, 1 was crippled,
and 137 were of a good hatch.
This compares favorably with the number of
good hatch in 1968 (108).
A 7.57 percent better hatch resulted in using a normal incubation temperature of 99.750 F (35.37%) compared with a test incubation temperature
of 99.250 F (27.80%).
A larger number of total eggs was laid by the tinamou held in four pens
with selected groups of two hens and one cock each (260 eggs) compared with
a community pen of eight hens and four cocks (65 eggs). A larger number of
fertile eggs was also produced in the pens with selected groups (148 compared to 33).
More fertile eggs were laid by selected groups of two hens and one cock in
four pens fed a standard 22 percent protein feed (148) compared with a
similar number of birds and pens fed a 28 percent protein feed (87). Both
feeds were similar in content except for protein levels.
In addition, more
fertile eggs hatched by birds fed the 22 percent protein feed (91) compared
with the 28 percent protein feed (31).

�- 186 -

RECOMMENDATIONS

Based upon results of tests run in 1969, recommendations for 1970 tests
are to (1) compare egg hatchability using two different levels of humidity
(50 percent versus 55 percent) in the test and control incubators.
(2)
Secure one year's additional data on comparing flock mating to selected
group mating on fertile egg production, and (3) compare the effects of two
protein levels in the feed (22 percent versus 16 percent) on fertile egg
production.

�- 187 -

EXPERIMENTAL

BREEDING

OF TINAMOU

Donald M. Hoffman

An initial field release of 42 pale spotted tinamou was made on April 8,
1969 in Baca County, southeast of Springfield.
Good progress continued in
improving production techni~ues for pale spotted tinamou at the Fort Collins
Wildlife Research Station.
From the 36 breeders (24 hens and 12 cocks)
on hand in May, 1969 there are enough 1969 hatched birds for use as breeders
in the 1970 tests (36) plus 10 1969 hatched and 30 adult breeder surplus
tinamou.
These 40 were available for release in 1970.

P. S. OBJECTIVE
To develop

game farm production

SEGMENT
1.
2.
3.

techniques

for pale spotted

tinamou.

OBJECTIVES

To compare the effects of two levels of temperature (99.750 F versus
99.250 F) during incubation on egg hatchability.
To measure the relationship of flock mating and selected group mating
to fertile egg production.
To measure the effects of two protein levels in the feed (28% versus
22%) on fertile egg production.

METHODS

AND MATERIALS

The detailed arrangement of tinamou breeding pens is shown in Figure
The pen schedule for tinamou for 1969 is shown in Table 1.

1.

With the assistance of Wayne H. Bohl, Research Biologist, U. S. Fish and
Wildlife Service, in accurately determining sexes, selected groups of 2 hens
and 1 cock were placed in pens 1, 2, 4, 5, 6, 7, 9, and 10 and a community
flock of 8 hens and 4 cocks was placed in the combined pen 3, 8, and alleyway on April 7, 1969. Thus, a total of 36 birds were used in breeding
experiments in 1969. Of these, 2 birds (1 cock and 1 hen) died while in
the breeding pens.
Numbers of eggs produced and numbers of fertile eggs
produced were compared for selected groups (pens 2, 4, 6, and 10) and the
community group (pens 3, 8, and alleyway, combined) all on similar feed
(22% protein).
Settings of tinamou eggs were made alternately in the test and control incubators every 7 days insofar as possible in 1969, similar to 1968. Temperature in the control incubator (Robbins, Model H-7) was held at the standard
99.750 F level and temperature in the test incubator (Humidaire, Model 50)
was lowered to 99.250 F. The relative humidity was maintained in both incubators (control and test) at a normal 55 percent.
The hatching success of
eggs in the two incubators was then compared.

�- 188 -

~-------------30'--~--------~
12'

,,12'

.-

"1"

-0)

to

0

¢

to

/

-,

ro

ex&gt;

C\J

~

. I

\I,

-an
V

/

-,

-

0)

V
SCALE 8
L...._ I

I

O'

10'

5'

Fig. 1.
Detoiled

Arrangement

of Tlnamou Breeding Pens.

�- 189 1.

Table

Pen schedule

Pen
Number

for tinamou,

Protein Level
22%
28%

2

T

3

x

4

T

5

x
T
x
T
x

T

Alleyway

x

Pens on Test

T

x

T

Number

T

T

10

4

Groups

T

T

9

Mating
Selected

x

T

7

8

El.ocks

T

1

6

1969.

T
T

4

3

Key: T = Pen on Test; x = Pen not on Test.
.,'~
Pens number 3, 8, and alleyway was used as 1 combined

4

pen for flock mating.

Birds in even numbered selective group pens (4 pens with 2 hens and 1 cock
each) were fed a standard game bird breeder feed containing 22 percent
protein.
Birds in odd numbered selective group pens (4 pens with 2 hens and
1 cock each) were fed a similar feed but containing 28 percent protein.
Eggs
were marked and placed in separate incubator trays to facilitate determination of (1) fertile eggs produced and (2) number of fertile eggs that
hatched.

RESULTS AND DISCUSSION
Incubation

Temperature

Test

Following culling procedures, eggs produced by breeders were alternately
placed in the control and test incubators every seven days because only

�- 190 -

two incubators were available for use in this test. Table 2 lists hatching
success of eggs in the test incubator (temperature of 99.250 F) in comparison with eggs in the control incubator (temperature 99.750 F). Results
showed a 7.57 percent better hatch in the control incubator when considering
total eggs hatched (35.37% compared to 27.80%) and 8.83 percent better hatch
when considering fertile eggs hatched (55.17% compared to 46.34%).
Percent of dead germs was 4.9 percent higher in the test incubator (25.9%)
compared with the control incubator (21.0%).
Percent of infertile eggs was
3.3 percent more (40.0%) in eggs incubated in the test incubator compared
with eggs in the control incubator (36.7%). Variation in percentages of
numbers of pipped eggs which did not hatch and number of crippled birds did
not vary more than 1 percent between the test and control incubator.
Results indicated the normal incubation temperature of 99.750 F was therefore better than an incubation temperature of 99.250 F.

Flock Mating Versus Selected

Group Mating

Test

Table 3 lists a comparison of numbers of eggs produced and numbers of fertile
eggs produced by 8 hens and 4 cocks in a community pen with the same number
of birds in four pens of 2 hens and 1 cock each. All birds were fed a
standard 22 percent protein feed. A much larger number total eggs was laid
by the tinamou. held in four pens with selected groups of 2 hens and 1 cock
each (260 eggs) compared with a community pen of 8 hens and 4 cocks (65 eggs).
In addition, a much larger number of fertile eggs was laid in the pens with
selected groups of 2 hens and 1 cock per pen (148 compared to 33).
There was evidence of egg eating in the community pen which may explain part
of this wide variation.
Eggs were gathered once per day through the laying
season.
Based upon the one season's data, it appears that selected groups
of 2 hens and 1 cock are better than community pens.
Plans are to run this
test again for one more year.

22 Percent

Protein Versus

28 Percent Protein Feed Test

Table 4 lists a comparison of numbers of fertile eggs produced and number
of fertile eggs that hatched by 22 percent and 28 percent protein levels
in the feed. Selected groups of 2 hens and 1 cock each in pen numbers 2,
4, 6, and 10 were fed 22 percent protein feed and like numbers of birds in
pen numbers 1, 5, 7, and 9 were fed 28 percent protein feed. More fertile
eggs were produced by birds fed the 22 percent protein feed (148) compared
with 28 percent protein feed (87) and more of the fertile eggs hatched by
birds fed the 22 percent protein feed (91) compared with the 28 percent
protein feed (31). By percentages, 61.5 percent of the fertile eggs hatched
using the 22 percent protein feed, compared with 35.6 percent of the fertile
eggs which hatched using the 28 percent protein feed.
From these comparisons it is evident that the standard 22 percent game bird
breeder feed was better than the 28 percent protein feed for pale spotted
tinamou egg production during 1969.

�Table 2. A comparison of hatchability of pale spotted tinamou eggs under two levels of temperature, 99.750 F
(normal) versus 99.250 F, 1969.

Item
On test 1/

Date
Set

Number
Set

5/6/69
5/20/69
6/3/69
6/17/69
7/1/69
7/15/69
7/29/69
8/13/69
8/26/69

9
13
22
35
33
34
24
22
13

Sub-Total
Control 1/

5/13/69
5/27/69
6/10/69
6/24/69
7/8/69
7/22/69
8/5/69
8/19/69
9/3/69

Number
of
·Infertile

Dead
Germs

2
1
5

1
7
7

12
14
19
12

10

8
9

Number
Pipped but
Not Hatched

o

Number
of
Good
Hatch

Percent
Total
Eggs
Hatched

Percent
Fertile
Eggs
Hatched

o
o

6
3
.7

o
o

8
11

85.71
25.00
41.18
34.78
57.89
66.66
50.00
28.57
50.00
46.34

Number
Crippled

10

1

o
o
o
o

2

66.66
23.08
31.82
22.86
33.33
29.41
25.00
18.18
15.38

13 (6.3%)

0 (0.0%)

57 (27.8%)

27.80

2
3
5

8
5

o

4
10

2

o

1

o

o

6
4

205

82 (40.0%) 53 (25.9%)

10
26
33
38
39
24
32
16
11

5

o

o

o

11

4

1

5
9

12

o

14
17
16
2

o

11

o

4
2

50.00
34.62
42.42
44.74
41.03
8.33
34.38
25.00
18.18

80 (34.9%)

35.37

55.17

137

31.10

51.49

11
9

5
9

1
2
1

10

4

15

7

o

9
6
6

7

5
1
2

o

5
1

o
o
o

Sub-Tota 1

229

84 (36.7%) 48 (21.0%)

16 (7.0%)

1 (0.4%)

Totals

434

166

29

1

101

100.00
60.00
66.66
62.96
53.33
22.22
47.83
40.00
40.00

1/ Eggs incubated in a small Humidaire Model 50 automatic incubator using temperature of 99.25 o F and relative
humidity of 55 percent.

1/ Eggs incubated in a large Robbins Model H-7 automatic incubator using temperature of 99.75° F and relative
humidity of 55 percent.

t-'
t-'

\.0

�Table 3.

A comparison of numbers of pale spotted tinamou eggs by month and by flock and selective group matings.

Al2ril
June
Max
Ju1X
August
Pen No. Culls Total Culls Total Culls Total Culls Total Culls Total Culls Laid Avail.

Egg Totals
Se1ective
21
InFlock Pen-11
GrouE PenSet fertile Fertile Hatched Total Fertile Total Fertile

1

0

0

2

6

2

13

2

20

0

9

6

48

42

41

19

22

13

2

0

2

3

19

3

28

3

24

1

13

10

86

76

74

3

71

46

86

71

4

0

0

3

13

2

22

2

13

1

5

8

53

45

44

6

38

28

53

38

5

1

2

3

14

0

15

1

10

0

7

5

48

43

43

12

31

13

Flock

0

0

1

5

6

23

5

21

3

16

15

65

50

52

19

33

15

6

0

0

0

0

1

16

0

14

1

14

2

44

42

43

22

21

9

44

21

7

0

0

0

8

0

15

0

12

0

0

0

39

39

40

18

22

5

9

0

0

6

7

3

14

1

9

1

3

11

33

22

23

11

12

0

10

0

0

1

12

0

18

1

26

0

21

2

77

75

74

56

18

8

Totals

1

4

19

84

17

164

15

149

7

92

59

493

434

434

166

268

137

11 A flock of 8 hens and 4 cocks in commUnity type pen (total 12 birds).
£1 Selected group of 2 hens and 1 cock per 4 pens (total 12 birds).

All birds used in flock vs. selected group mating tests were on 22 percent protein feed.

65

33
•....
\D
N

65

33

77

18

260

148

�Table 4. A comparison of number of pale spotted tinamou fertile eggs produced and number of fertile eggs
hatched using 22 percent and 28 percent protein levels in the feed.
Protein Test
22%
28%
Fertile Hatched Fertile
Hatched

Pen Number

Culls

Laid

Egg Totals
Avail. Set
Infertile

1

6

48

42

41

19

22

13

2

10

86

76

74

3

71

46

71

46

4

8

53

45

44

6

38

28 .

38

28

5

5

48

43

43

12

31

13

Flock

15

65

50

52

19

33

15

6

2

44

42

43

22

21

9

7

0

39

39

40

18

22

5

22

5

9

11

33

22

23

11

12

0

12

0

10

2

77

75

74

56

18

8

18

8

Totals

59

493

434

434

166

268

137

148

91

87

31

Percent ha-tch (fertile eggs)

Fertile

Hatched

21

22

13

31

l3

9

I-'

\0

61.5

35.6

w

�- 194 -

East Pens versus West Pens
Although not written up as a part of the 1969 tests, it was found that a
difference existed in the numbers of eggs produced and numbers of fertile
eggs produced by either east or west locations of the pens. An attempt
was made to plant clumps of crested wheatgrass and tall wheatgrass in the
east pens early in the spring of 1969 so cover would be comparable to that
found in the west pens. These grass transplants did not take, resulting in
more of the eggs being laid without benefit of shade and apparently many of
these eggs became over-heated before being gathered.
Table 4 lists eggs laid by individual pens. Total numbers of eggs produced
in the 4 west pens (numbers 1, 2, 4, and 5) was 235 compared to 193 for the
4 east pens (numbers 6, 7, 9, and 10). Numbers of fertile eggs produced in
these 4 west pens was 162 compared to 73 for the 4 east pens. This comparison indicated that suitable cover in the breeding pens is important and
clumps of crested wheatgrass and tall wheatgrass with protective wire cages
were replanted in the east tinamou pen in early April, 1970.
Test pens were alternated by east and west locations in the 22 percent
versus 28 percent protein feed and flock mating versus selected group mating
tests, so cover differences should not have affected test results adversely.

Prepared

by

�April

1970

- 195 -

JOB PROGRESS

COLORADO

Sta te of
No.

W-37-R-23

Plan No.

19

Project
Work

Job Title
Period

REPORT

Covered:

Personnel:

Job No.
Study

April

Game Bird Survey

of Tinamou

2

Adaptability

1, 1969 to March

31, 1970

Charles Gibson (U.S. Forest Service),
Bert Widhalm, and Warren D. Snyder

Donald

M. Hoffman,

ABSTRACT

Forty-two pale spotted tinamou, Nothura darwinii sa1vadorii, were released
during April, 1969 on U. S. Forest Service rangeland in southern Baca
County, Colorado.
Searches in 1969 failed to provide evidence that the
species remained, survived or reproduced in the vicinity of the release
site.
Two unconfirmed
sightings were reported by local residents from
locations ten to fifteen miles away from the release site.

��- 197 -

STUDY OF TINAMOU ADAPTABILITY
Warren D. Snyder

P. S. OBJECTIVE
To determine the ability of pale spotted tinamou to establish reproducing
populations in areas of mixed tall and mid-grasses, sandsage and yucca on
the Comanche National Grasslands, Baca County, Colorado.

SEGMENT
1.
2.

Introduce
Determine

OBJECTIVES

pale spotted tinamou.
presence and dispersal

METHODS
Reference is made to Snyder
in this study.

of pale spotted

tinamou.

AND MATERIALS

(1969) for review of methods

and materials

used

RESULTS AND DISCUSSION
A gallinaceous guzzler in Section 13, Township 32 South, Range 46 West,
southeast of Springfield in Baca County was the site for release of fortytwo pale spotted tinamou.
These birds had been raised at the Fort Collins
Wildlife Research Station under Work Plan 19, Job 1. Most were released
directly from the crate on April 8, 1969. Ten were fenced in and held under
the guzzler canopy for three days before release.
Scratch grain and water
were provided.
Most of the birds ran or flew only a short distance upon
release.
The release site was located within a large Forest Service pasture that had
received little or no livestock grazing for several years.
The dry stream
bed of the Sand Arroyo drainage traversed the site. Sand sagebrush, yucca
and a variety of grasses and forbs were primary components of the habitat.
Bert Widhalm, Wildlife Conservation Officer, reported finding feathers
indicating mortality of one bird during a search on April 20th. No other
evidence of mortality or survival was obtained at that time. Carrizo
District personnel of the U. S. Forest Service visited the release site
several times, but were unable to find tinamou or evidence of their presence.
Searches by the writer were completed during June 20-21, 1969 and on December 27th in the release site pasture and surrounding areas.
A dog was used
during these searches.
No birds or evidence of their presence was found.

�- 198 -

U. S. Forest Service personnel received two unconfirmed reports of tinamou
sightings during 1969. Leroy Sunday, of Springfield, reported sightings
of 1, 1, and 2 birds in Sections 11, 14, and 34, respectively, of T. 31 S.,
R. 45 W. The nearest of these sightings was approximately ten miles east
of the release site, down the Sand Arroyo drainage.
The second reported sighting was of about six birds observed one or two
months after the release.
Mr. Jesse Tetters reported the observation in or
near Section 21, T. 33 S., R. 48 W., or about fifteen miles southwest of
the release site. These locations will be searched during the forthcoming
segment.
In summary, we have no evidence that the tinamou remained at or near the
release site. Two unconfirmed sightings by local residents provide some
evidence that the birds moved a considerable distance from the release site.

LITERATURE

Snyder, W. D. 1969.
Fish and Parks.

Prepared

CITED

Study of tinamou adaptability.
Colorado
Game Res. Rpt. April, pp. 177-181.

by __ -1v.,--",,--_~:.-.........:...-:-....:;;...~()=-..;:~l",-- ~_I-I-I~",,---_IJ.
__/ __
Warren D. Snyder ~
Wildlife Researcher

Div. of Game,

�</text>
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                  <text>July,

- 1 -

JOB PROGRESS

State

Work

COLORADO

of
No.

W-10l-R-12

Plan No.

1

Project

Job Title:
Period

REPORT

Covered:

Personnel:

Game Range
Job No.

Investigations
1

Effects of Different Intensities of
Simulated Browsing on Key Browse Species
April

1, 1969 to March 31, 1970

Harold

R. Shepherd

ABSTRACT

A typed first-draft manuscript,
inclusive of figures and tables, was
prepared and copies were submitted to reviewers for comments and
suggestions.

1970

��- 3 -

EFFECTS
SIMULATED

OF DIFFERENT INTENSITIES OF
BROWSING ON KEY BROWSE SPECIES
Harold R. Shepherd

Po So OBJECTIVE
To publish

results

of completed

field work.

SEGMENT OBJECTIVES
1.

To make revisions

2.

To submit manuscript

in the manuscript
to publisher

as necessary.
and follow through

to final publication.

RESULTS AND DISCUSSION
A typed first-draft manuscript, inclusive of figures and tables, was prepared and copies were submitted to five reviewers for their comments and
suggestions for improvement in content and format.
Edited copies and
written statements of comment and criticism have been received from two
reviewers.
Upon receipt of similar materials from the others, a final
draft will be prepared and submitted for publication as a Technica 1 Publication of the Colorado Division of Game, Fish and Parks.

Prepared

~ __"kit:)(~&gt;,{)l~n:t
j)
..
by;;;
Harold R. S~epherd
Wildlife Researcher

��- 5 -

July, 1970

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~---------

Project No.

W-10l-R-12

Work Plan No.

1

Job Title:

Game Range Investigations
Job No.

Paddock Studies on Effect of Varying

4
Intensities

Period Covered:

April 1, 1969 through March 31, 1970

Personnel:

Richard M. Bartmann

of Deer-Use

ABSTRACT

A final report was submitted for this job during Segment 11. This report
appears in the July 1969 Game Research Report, Part 3, pp. 305-393.

��July, 1970

- 7 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-10l-R-12

Work Plan No.

2

Job Title:

Publication

Game Range Investigations
Job No.

I

of Results of the Little Hills Grazing

Study

Period Covered:

April 1, 1969 through March 31, 1970

Personnel:

William T. McKean, Richard M. Bartmann, Bertram D. Baker,
Jack R. Grieb, Harold M. Swope, Laurence E. Riordan,
and David C. Bowden.

ABSTRACT

A second draft of five chapters of the Technical Bulletin was completed.
A first draft of the introductory chapter was almost completed, as well
as a section on plant responses based upon Canfield line transects.
nvO
papers were prepared for other Division publications.

��- 9 -

PUBLICATION OF RESULTS OF LITTLE HILLS GRAZING STUDY
William T. McKean
P. S. OBJECTIVE
To publish the results of the Little Hills grazing study.
SEGMENT OBJECTIVE
To prepare a final report on the study and publish the results in the
Division's Technical Bulletin series.
RESULTS AND DISCUSSION
Progress
1.

A second draft has been completed of the following material, including
figures, for a Division Technical Bulletin:
a.

Key browse production and utilization based upon stem length measurements before and after grazing.

b.

Utilization estimates of all forage classes including the less
abundant species.

c.

Vigor of Indian ricegrass (Oryzopsis hymenoides) and low phlox
(Phlox caespitosa).

d.

Pasture range condition and trend based on standard Big Game Range
Analysis procedures.

e.

Weight responses of livestock.

2.

A first draft of the introduction is completed. This includes descriptive sections on: location, past land use, climate, topography, soil
and vegetation.

3.

A first draft of the chapter on pasture vegetative response as indicated
by Canfield line transects has been completed by Richard M. Bartmann.

4.

Citations for other written materials prepared during the segment are
as follows:
McKean, W. T. 1970. Fencing for mule deer.
Div., Game Inf. Leaf1. 3p. I11us.

Colo. Game, Fish and Parks

McKean, W. T. 1970. Article about Little Hills grazing study.
Research Review, 1969. 1p. I11us.

Game

�- 10 -

Future

Plans

There remain to be written:
a section on methodology; a summary and
discussion of results; recommendations
for management; numerous modifications
to the text to shorten and make more concise; lists of tables and figures;
the appendix, literature cited, and acknowledgements;
changes needed after
the copy has been reviewed and edited by others; proof reading and other
followup details prior to publication.

?jt'-/d

:"I'

Prepared

•

bY:d~'~z,?'

I/' ",.

"--n
~c, :yt''';

~../'

William T. McKean
Wildlife Researcher

__.

• - .

ZC-~-f.-i'r

�July, 1970

- 11 -

JOB PROGRESS REPORT

State of

~~C~O~L~O~RAD~~O~

.Project No.

W-101-R-12

Work Plan No.

_
Game Range Investigations

2

Job No.

Job Title

Rodent Effects on Deer Winter Range

Period Covered:

April 1, 1969 to Harch 31, 1970

Personnel:

Harold R. Shepherd

3

ABSTRACT

Time budgeted for this job was used to complete a first-draft manuscript
for a final report on the "Effects of Variable Clipping on Key Browse
Species in Southwestern Colorado".
Consequent;ly, segment objectives
were not accomplished.

��- 13 -

RODENT

EFFECTS ON DEER WINTER

RANGE

Harold R. Shepherd

Po So OBJECTIVE
To complete the laboratory work on the food-habits
and publish results of the entire studyo

portion

of the study

SEGMENT OBJECTIVES
1.

To prepare a final report presenting

2.

To publish results

results

of studyo

of the study.

RESULTS AND DISCUSSION
No work was accomplished toward attainment of objectives during this segment.
The extra time realized was used in completing a first-draft manuscript for a final report on the "Effects of Variable Clipping on Key Browse
Species in Southwestern Colorado".

of!

Prepared

by

,q::4N--K'~

,

(-j;
f
A,·I? 4U,'j!
...
llU'e-{

Harold R. Shepherd'
Wildlife Researcher

��July, 1970

- 15 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-10l-R-12

Work Plan No.

3

Job Title:

Survey, Inventory,

Game Range Investigations
Job No.
and Analysis

3
of Deer and Elk Winter Ranges

Period Covered:

April 1, 1969 through March 31, 1970.

Personnel:

Keith Giezentanner and Bert Baker.
Cooperating:
Bill
Brown, Jim Cruse, and Jerry Martinez, all of the U. S.
Forest Service; Bob Jacobsen and Neil McCleery, U. S.
Bureau of Land Management.

ABSTRACT

Maps and reports of big game winter range inventories for National Forest
portions of Big Game Management Units 55 (Taylor River) and 67 (Tomichi),
and all of Unit 22 (Piceance) were completed, bound, and distributed to
appropriate Division Field Services and Game Planning Services offices.
Cooperative inventories for Units 55 and 67 were accomplished with the
Gunnison National Forest and those for Unit 22 with Craig District, Bureau
of Land Management.

��- 17 -

SURVEY,

INVENTORY,

AND ANALYSIS

OF DEER AND ELK WINTER

RANGES

Bertram D. Baker

P. S. OBJECTIVE
To prepare inventories of big game winter range resources on the Grand MesaUncompahgre, Gunnison, Rio Grande, and San Juan National Forests and Game
Unit 22 (Piceance) in Colorado.

SEGMENT

OBJECTIVES

1.

To prepare 2-inch scale vegetation type maps of areas within zones of
average winter use and show type acreages by landownership, range condition transect locations, and other pertinent information from completed cooperative inventories with the Gunnison National Forest and
Craig District of the Bureau of Land Management.

2.

To prepare t-inch scale big game winter range maps of Management Units
22 and 67 on base maps furnished by the BLM for Unit 22 and Gunnison
National Forest for Unit 67.

3.

To prepare winter

4.

To bind 2-inch scale maps in holders and t-inch scale maps,
and reports in binder file folders for Units 22 and 67.

5.

To distribute to appropriate Division offices winter range inventory
maps and binder file folders for Units 22, 55, and 67.

range inventory

METHODS

reports

for Units 22 and 67.
summaries,

AND MATERIALS

Collections fo field data had already been accomplished in previous years
by interagency-approved
techniques that have been detailed before in these
reports (Denney 1962:51-96; Baker 1964:90-92; 1966:55-56).

RESULTS
Map Preparation,
Gunnison

National

AND DISCUSSION

Binding,

and Distribution

Forest

Final copy, color-coded, and map-holder bound 2-inch scale township plat
maps of the winter range inventory for Unit 55 (Taylor River) were transferred to appropriate Division offices.
The maps had been held pending
final approval of and simultaneous release with the unit winter range

�- 19 also went directiy to the Northwest Regional Office to be available for
immediate use. Mr. Giezentanner color-coded the four copies of each map
and brought legends up-to-date with addenda notations.
Since Unit 22 map parts are too large for convenient binding and use from
map holders, methods of filing and handling final copies are optional by
Division section.
Research Center W-lOl-R Project files will receive and
retain one complete set of 2-inch scale maps.
Winter range and key area boundary lines shown on 2-inch scale maps of Unit
22 were transferred in red pencil to five ~-inch scale BLM base maps.
These
~-inch maps,trimmed
to about 11- by IS-inch size sheets, also had unit
boundaries, numbers, and names inked on them in black.
A copy of each map
was placed as a fold-out in each of five binder file folders currently
awaiting addition of finished narrative unit reports for distribution.

Data Reproduction,

Gunnison

National

Compilation,

and Analysis

Forest

Vegetation and land category acreages of National Forest winter ranges were
compiled and summarized in table form for Unit 67. The table is included-in
the inventory final report that is presented as APPENDIX A of this report.
Copies of the Unit 67 final report accompanied mapS and allied inventory
information in each of five binder file folders.
Preceding work on Unit 67
inventory summarization, however, the previously mentioned Unit 55 report
was given final approval early in the project segment.
Copies of the latter
report were made and bound in similar number and manner as Unit 67 materials
were handled.
The Division's Southwest Regional Office got three, Denver
Game Planning Services Section one, and Fort Collins Research Center one
binder file folder of each unit. Map holders of 2-inch scale maps were
transferred at the same time binder folders were turned over to Division
offices.

Grand Mesa-Uncompahgre,

Rio Grande,

and San Juan National

Forests

Nothing more was accomplished on data reproduction, compilation, and analyses
of these forests' winter range inventories.
Pending delivery of 2-inch
scale township plat maps for the latter two forests, the officework phase of
the inventory program must be delayed.
Loss of original field annotated air
photos of the survey done on the Grand Mesa-Uncompahgre
National Forest is
complicating proofwork that was started on 2-inch scale township plats on
hand from an earlier distribution.

White River Resource

Area

- Bureau of Land Management

The compilation of vegetation and land category acreages followed the sequence
of 2-inch scale map proofwork because so much Unit 22 map territory needed
correcting.
A table was first composed of acreages west of Piceance Creek

�- 20 (area corresponding to BLM Yellow Creek Management Area) in order to compare
results with data already available. Data for area east of Piceance Creek
(Little Hills Triangle) were summarized secondly. Then, all information
were combined to present vegetation and land category tabulation by landownership for the entire unit in the final unit report that was prepared.
Copies of the Unit 22 final report, presented herewith as APPENDIX B, accompanied maps and allied inventory information in each of five, 2-volume sets
of binder file folders. The Division's Northwest Regional Office was given
three sets of folders, Denver Game Planning Services section received one
set, and one set was retained in the Fort Collins Research Center files.
Two-inch scale winter range inventory maps were transferred Simultaneously
with binder file folders.

LITERATURE CITED
Baker, B. D. 1964. Browse transect analysis and application, pp. 87-106.
In Game Res. Rep., January (Part One). Colo. Dep. Game, Fish and Parks.
121 pp.
1966. Browse transect analysis and application, pp. 53-67
In
Game Res. Rep., July, Part One. Colo. Dep. Game, Fish and Parks. 93 pp.
0

Denney, R. N. 1962. Browse transect analysis and application, pp.Sl-96.
In Game Res. Rep., April. Colo. Dep. Game and Fish. 96 pp.

Prepared by

&amp;i;;,&lt;fN'I; j). ,~
Bertram D. Baker
Assistant 'Wildlife Researcher

�- 21 -

APPENDIX

BIG GAME WINTER

A

RANGE ANALYSIS

GAME UNIT 67 - TOMICHI

INTRODUCTION
The following report presents a description of Big Game Management Unit 67
and a review of a survey of deer and elk winter ranges of the unit within
Gunnison National Forest boundaries.
Field data were collected in the summer
of 1962 by Gunnison National Forest Range Analyst Jack McCrain, and Game,
Fish and Parks Division seasonal employees Dave Stearns and Gary Brown.
Inventory data for Unit 67 are more fragmentary than for Units 53, 54, 55,
and 63, also covered in 1962, because: (1) very limited pre-inventory information was available on winter distribution of deer and elk, and (2) being
the last area examined in a crowded summer schedule, Unit 67 suffered from
accumulative fieldwork overages from preceding units.
Furthermore, winter ranges in the southern part of Unit 67 are mainly occupied by elk. Comparatively light snow accumulations there most years allow
the elk freedom of movement over a large expanse of territory.
This circumstance made it difficult to determine average extent of winter range occupation as well as to pinpoint placed that would qualify as being key areas.
Cebolla District Ranger Wally Chesbro delineated what he believed to be key
areas, but time limitations prevented adequate examination of them.
Consequently, some of the key areas noted on 2-inch scale township plat maps lack
completely, or in part, vegetation type data.
Map production was accomplished principally by U. S. Forest Service Region 2
engineers in Denver.
Considerable map proofing and type acreage recomputations
and compilations were done by Regional Wildlife Management Biologist Jim
Cruse and the author.
Assistance in supplying and updating maps by Fishery
Management Biologist Bill Brown, currently assigned to the Gunnison National
Forest Range and Wildlife Staff, is gratefully acknowledged.

DESCRIPTION

OF UNIT

Location.--The major portion of Unit 67 lies in and corresponds with the
northwestern one-third of Saguache County.
A small northeastern part and
larger northern and western bordering portions are situated in Gunnison County.
Near the southwest corner of Unit 67, in the vicinity of Cathedral, a few
sections form a block that is located in Hinsdale County.
Colorado Game, Fish and Parks Division Laws and Regulations Handbook, 1965
(p. 12, Chap. 3 - Big Game) gives the boundaries of Unit 67 as follows:
"Those portions of Gunnison, Hinsdale and Saguache Counties east of a line

�- 22 -

along the divide between Cebolla and Cochetopa Creeks to Ground Hog Peak,
thence down Cathedral Creek to Spring Creek, thence down Spring Creek to
Cebolla Creek and down Cebolla Creek to State Highway No. 149; east of
State Highway No. 149 to its intersection with U. S. Highway No. 50: south
of U. S. Highway No. 50 to the Continental Divide; and west and north of
the Continental Divide to the divide between Cebolla and Cochetopa Creeks".
The only question concerning the preceding description is the definition
of "Ground Hog Peak".
Examination of best maps available, including the
U.S.G.S. Stewart Peak Topographic Quad, shows a "Ground Hog Park"; but,
there is no peak evident by that name.
It is assumed that "Peak" was
exchanged for "Park" when the description was prepared.
Because Cathedral
Creek flows through Ground Hog Park, the mixup is not of great importance.
Nevertheless, should the description ever be rewritten in the future, it
is recommended that the first part be changed to read "Those portions of
Gunnison, Hinsdale, and Saguache Counties east of the Cebolla-Cochetopa
Creeks divide from the Continental Divide to the origin of Cathedral Creek,
thence down C thedral Creek to Spring Creek , and so forth, as it reads
presently.
Area. -- Computed by use of a planimeter on a t-inch scale map, the total
area of Unit 67 was found to be 1,005.6 square miles, or 643,584 acres.
By similar means, it was also determined that there are about 497 square
miles (318,080 acres), or 49.4%, of the unit inside of the Gunnison
National Forest boundary.
Physical features. -- Moderately rough mountainous terrain characterizes
most of the southeastern two-thirds of Unit 67. A foothills zone, consisting of comparatively gently sloping hills and mesas, corresponds with
the northwestern one-third of the unit.
Highest elevations are found in the extreme eastern and southern parts.
Approaching very nearly 14,000 feet in elevation are Mt. Ouray in the east,
and San Luis and Stewart Peaks in the south. That portion of the Continental Divide coincidental with the crest of the Cochetopa Hills and the
unit's southeastern boundary varies elevationally between about 10,000
and 11,900 feet, perhaps averaging somewhere between 10,700 and 10,900 feet.
With high ground on the unit's south side, it naturally follows that most
streams flow northward.
Main drainage systems fitting that category from
east to west, in order, are Marshall, Long Branch, Needle, Razor, Cochetopa,
South Beaver, Willow, and Cebolla Creeks.
The Gunnison River and Tomichi
Creek drain westward on the unit's north boundary.
Needle Creek Reservoir is the largest, wholly contained lake within the
unit; however, at maximum capacity, it probably will not cover much over
65 acres.
The narrow Gunnison River neck of Blue Mesa Reservoir east of
the bridge on State Highway No. 149 is estimated to have approximately
640 surface acres at high water.
Vouga Reservoir on Razor Creek, MCDonough
Reservoir on Los Pinos Creek, four other small reservoirs in the Cochetopa
Creek drainage, and a few small scattered alpine lakes account for most
of the remaining dead water storage in Unit 67.

�- 23 -

Cochetopa Park is the largest park in Unit 67. It completely encircles
the monolithic Cochetopa Dome and lies along and between West Pass and
Archuleta Creeks on the east and middle Cochetopa Creek on the west.
Industry -- Production and sale of livestock and livestock products are
the leading industries in Gunnison and Saguache Counties (Colorado State
Planning Division 1961:848,877).
Cattle ranches in. the valleys of the
Gunnison River, and Tomichi, Needle, Razor, Cochetopa, and Cebolla Creeks
provide most of the production.
In both counties in 1960, income from mineral extraction and milling
ranked second to that provided by livestock production.
Principal minerals
that year in Gunnison County were coal, zinc, sand and gravel, and lead.
Saguache County's main minerals were uranium ore and gem stones (Colorado
State Planning Division 1961:848,877).
Gunnison, Gunnison County's largest city and the county seat, is the only
incorporated town in Unit 67. Actually, since highways there act as
coundaries of Units 54, 55, and 67 and meet within and divide the city,
only about the southern one-third of Gunnison lies in Unit 67.
Gunnison's population of 3,477 in 1960 approximated 63% of that for the
entire county (Colorado State Planning Division 1961:783).
The 1970
census should show gains in both population and economy of Gunnison city
and county due to currently expanding state-supported Western State College
and recreation developments at Blue Mesa Reservoir for water sports.

LIMITS OF AVERAGE WINTER RANGE
Long Branch Creek drainage and vicinity. -- A short bulge of winter range
extends southward up Long Branch Creek from the latter's confluence with
Tomichi Creek to just above the Long Branch Fire Station (old F. S. Guard
Station).
The upper average winter range limit line at its lowest point
lies at approximately 8,400 feet elevation on Tomichi Creek and reaches
about 9,300 feet at the highest point on the divide east of Long Branch
Creek.
Heavily timbered north slopes between Long Branch Creek and Owen
Creek very severely restrict winter use of that area. Consequently, the
upper limit line is placed immediately adjacent to and along the south
side of Tomichi Creek from Long Branch Creek to about Deer Haven Ranch
which lies at the mouth of Hicks Creek.
Needle Creek-Owen Creek-Tie Creek drainages. -- From Deer Haven Ranch, the
upper limits line angles southwestwardly, intercepting Tie Creek at about
8,700 feet. The line continues on to intercept the 9,400-foot contour on
the divide between Tie and Owen Creeks, traverses west across Owen Creek
at about 8,900 feet, and then varies between 8,400 and 10,000 feet in making
a southward loop in the middle Needle Creek drainage that includes Needle
Creek Reservoir.

�- 24 -

Prosser Creek-Razor Creek-Barret Creek drainages.-:Virtually
all of the winter range zone between Barret Creek and the west side of Razor Dome lies
outside of the National Forest.
Nevertheless, the upper limits line was
approximated on maps outside of the Forest Boundary between the aforementioned
topographic features except where it was looped briefly inside near Snyder
Gulch and Prosser Rock on the Razor Creek drainage.
Archuleta Creek-West Pass Creek drainages.--The upper winter range limit
line approaches 11,200 feet elevation on the south slopes of Razor Dome.
The entire north side (generally southerly exposures) of West Pass Creek is,
in fact, characteristically
open grass and/or sagebrush parkland that permits
unrestricted use by deer and elk most years to elevations of well over 10,000
feet.
All of Cochetopa Dome south of West Pass Creek has been included within the
zone of average winter use, even though its maximum elevation is 11,120 feet,
and much of its upper slopes consist of boulder fields and heavy timber.
That situation precludes occupation of the Dome's top most years by big game.
The south side of Lujan-West Pass Creek, from the Pine Creek confluence west
to Salaya Creek, is excluded from the winter zone. The Salaya-West Pass
Creek junction lies at about 9,400 feet. From there, and southeastward to
the head of Archuleta Creek on Cochetopa Pass, the upper limit holds close
to the 10,000-foot contour.
The line crosses the Continental Divide a short
distance into Unit 68, then returns to the summit of Cochetopa Pass. From
the Pass, the line goes southwestwardly, varying around 10,000 feet elevation
near headwater areas of Los Creek, Monchego Creek, Texas Creek, and Van
Tassel Gulch.
Cochetopa Creek-Los Pinos Creek drainageso--The upper limit line crosses
Cochetopa Creek and its West Fork at about 9,300 feet elevation and then
varies to a maximum of 10,000 feet in that vicinity.
No fieldwork was done in nor information gathered on extent of winter range
for T. 45 N., R. 1 E., N.M.P.M.; so, the tentatively drawn upper limits line
on the 1/4-inch scale unit map is not authentic for that township.
The upper limit line west of Los Pinos and Cochetopa Creeks varies considerably
in elevation.
It is located at about 9,600 feet on Ralston Creek, 10,000 feet
on Trail Creek, goes up to 10,800 feet on Pole Road Gulch, back to around
10,000 feet on Willow Creek and Townsite and Homestead Gulches, and then down
to 9,300 feet where it exits the National Forest on Bead Creek.
The line
perhaps averages slightly under 10,000 feet elevation in this sector of the
Forest.
Cathedral Creek-Spring Creek drainages.--Elevation
of the upper winter range
line approximates 10,000 feet in the area southwest of Los Pinos Pass and
in the vicinity of lower Cathedral Creek. The line drops to about 9,500
feet where it crosses Spring Creek and leaves Unit 67.
Generalo--Lack
of time prevented field reconnaissance of potential winter
range on the National Forest in the area from Los Pinos Pass northeastward
to upper East Beaver Creek and Stubbs Gulch.
However, the assumption was

�- 25 -

made by local Division and Forest Service Administrators,
and accepted by
us, that generally north and northwest exposures and high elevations
accompanied by heavy snow accumulations normally prevent winter use by
big game on that particular portion of the Forest.
That explains why
the 1/4-inch scale map of Unit 67 shows a big gap in the upper winter
range line north and west of the divide between the Cochetopa Creek and
Tomichi Creek-Gunnison River-Cebolla
Creek drainages.
Applicable to those parts of Unit 67 covered by this survey, the upper
winter range line exhibits so much elevational variability that no attempt
is made to present an average elevation of the line except where already
stated by locale.
Perhaps worthy of repetition, though, are the lower
and upper extreme elevations of 8,400 feet and 11,200 feet for the upper
limit line, respectively, near Long Branch Creek and Razor Dome.
At no point does a lower limit line enter the Gunnison National Forest.
Lowest point in the unit is at the bridge on State Highway No. 149 and
Blue Mesa Reservoir which is about 7,600 feet in elevation.

KEY AREAS
Previously discussed time limitations (see INTRODUCTION) for doing fieldwork severely restricted the amount of quality of information that could
be obtained relative to key areas in Unit 67. Largely through knowledge
supplied by local Division and Forest Service employees, several comparatively small areas of Cochetopa Creek and its eastern tributaries were
delineated as being key; some areas were designated specifically for elk.
No names were given to any of these key areas, and most were not visited
or type mapped.
The only clues to authenticity of these key areas are results of two pellet group count transects.
Noticeably more deer than elk
pellet groups were recorded on the key area west of Cochetopa Dome, an
area supposedly used mainly by elk. Another transect that was read on the
large, non-specific West Pass Creek key area showed a fairly substantial
pellet group count for elk and less for deer.
These results indicate that
much more information is necessary before accurate assessments can be made
of territories that might be considered as being key.
No indictment is intended by the foregoing discussions,
since a similar
situation was sidestepped when encountered on surveys of the Rio Grande
River country.
Like the latter area, which is characterized by extensive
parkland vegetative types used mostly by elk, the middle and upper
Cochetopa Creek drainage is so open most winters that the elk are not
consistently dependent on certain portions for existence.
The problem
was circumvented on the Rio Grande National Forest by recognizing the
difficulty of delineating key areas and establishing arbitrarily chosen
geographical subdivisions with which range transecting data could be
associated.
In contrast to apparent lack of decision about key areas east of Cochetopa
Creek, thorough field checking of areas west of Middle Cochetopa Creek revealed that use was both light and inconsistent enough by deer and elk to
prevent classifying any of that territory as being key.

�- 26 Two key areas are named and delineated on maps.
primarily by deer. They were designated because
tation and current data on browse age-form class
group counts strongly indicated that they should
scriptions of the areas are given as follows:
Long Branch key area.--All
drainage.

winter

Needle-Owen Creeks key area.--All
Dutchman (Tie) Creeks drainages.

range within

winter

These two are used
prior history of habicomposition and pellet
be so classified.
De-

the Long Branch Creek

range within

the Needle-Owen-

LANDOWNERSHIP
Presented before in this report was the finding that slightly over 49% of
Unit 67's total area lies within boundaries of the Gunnison National Forest.
An unknown small fraction of that 49% is possessed under private ranching
and mining patents.
It is estimated that of the remaining 51% non-Forest
land, approximately two-thirds is controlled by the Bureau of Land Management in public lands and one-third is in private ownership.
BLM and private
landownership is depicted graphically on the "Public Lands Outdoor Recreation
Map, Powderhorn Area", published and distributed jointly by the Division
and BLM, U. S. Department of the Interior.
A summary of vegetation and land category acreages by landownership within
Forest boundaries is presented on Table 1. It is emphasized that those
data are fragmentary, because type mapping was accomplished for only about
one-third of the National Forest big game winter zone. The information is
included here, however, to give some idea about relative amounts of the
vegetation types and proportions of private and Federal lands in the
territory covered by this inventory.
Moreover, the footnote on Table 1
gives additional explanation of the basis for summary data.

VEGETATION

TYPES

Discussion of principal categories of vegetative cover found on National
Forest big game winter range in Unit 67 is presented in paragraphs to follow.
This is done in spite of incomplete vegetation type acreage data for reasons
already explained.
Perhaps the most important thing that can be derived from
treating vegetation categories is an enumeration of dominant and principal
associated plants found in the categories.
Acreages dominated by Various
species would be of added benefit but are not possible or relevant in this
inventory.
Conifer type.--Regardless
of obvious inventory shortcomings, the Conifer
Type (Type 6) is very important to wintering big game on the National Forest.
Douglas fir (Pseudotsuga menziesii) is the most frequently named coniferous
species in type designations.
Less frequently cited as type species, in their
approximate decreasing order of importance, are lodgepole pine (Pinus contorta),
Engelmann spruce (Picea engelmanni), ponderosa pine (Pinus ponderosa), and
limber pine (Pinus flexilis).

�- 27 -

Table 1. Vegetation type and land category acreage sunnnary, big game
winter range within Gunnison National Forest and Came Unit 67.*

No Fo

Acreage
Pri.

Total

1,648

0

1,648

239

0

239

6,018

347

6,365

5 - Browse

251

76

327

6 - Conifer

7,099

30

7,129

1,309

20

1,329

40,560

566

41,126

57,124

1,039

58,163

Type or Land Category

1 - Grassland
2 - Meadow
4 - Sagebrush

10 - Broadleaf

and Rabbitbrush

Tree

Unclassified

TOTALS
~'r

Sunnnary represents only those townships where type mappings and/or an upper
winter range limit line was established.
At least three townships that are
assumed to have winter range on the Forest had nothing recorded for them nor
were maps issued.
Two townships contain only an upper limits line. Most of
the remaining 2-inch scale township plats show only partial type mapping
within the winter range zone.

It is of interest to note that lodgepole pine reaches southernmost limits of
contiguous distribution in Colorado on the eastern and southern perimeter
of the Gunnison Basin along and near the Continental Divide.
Recollections
by the author places the southern limit near Los Pinos Pass which is substantiated by a report of Barrell (1969:332) that places an association of
lodgepole and Colorado spruce (Picea pungens) east of the same passo The
winter range inventory report for a survey of Unit 76 (Rio Grande) lying
south of Unit 67 listed no lodgepole pine in vegetation type sunnnaries (Baker
1967:58).
Thus, except for possible isolated, sporadic occurrences as presented by Preston (1940:24) and Fowe11s (1965:373), lodgepole pine is rarely
found south of Game Units 66 (Lake Fork Gunnison) and 67 of the Gunnison Basin.
Concurrent with the decline in abundance of lodgepole in southern Colorado
is the comparable increase in abundance of aspen (Populus tremuloides) and
such conifers as Engelmann and Colorado spruces, white fir (Abies conco1or),
bristlecone pine (Pinus aristata), and limber pine. Aspen, in particular,

�- 28 -

is an intolerant fire species like lodgepole and apparently replaces lodgepole in that capacity south of the Gunnison River drainage system.
In any
event, aspen is a frequently-listed
codominant with all of the coniferous
species and for sizable acreages in Unit 67.
Of subdominant browse species found with the conifers, big sagebrush
(Artemisia tridentata) is probably the most abundant by being listed in
four types. Common juniper (Juniperus communis) and antelope bitterbrush
(Purshia tridentata) are also listed as being present but only in one type
each.
Lowest-level plants are led in type designations by herbaceous grasses;
more specifically, Arizona and Thurber fescues (Festuca arizonica; K.
thurberi) occur, either singly or in combination with each other, in four
Conifer Types.
The plant symbol "FES" that is used in type deSignations
indicates the presence of both species.
Actinea
tions.

(Hymenoxys
It appears

sp.) is the only forb listed in coniferous
only once.

type designa-

Sagebrush~.
-- The Sagebrush or Rabbitbrush Type (Type 4) ranks a close
second acreage-wise to the Conifer Type in this partial Unit 67 winter
range inventory.
As would logically be expected, the Artemisia and
Chrysothamnus genera dominate Type 4 lands. Big sagebrush is the most
abundant species of the former genus.
Douglas rabbitbrush (Chrysothamnus
viscidiflorus) is the most abundant species of the latter genus, although
various rabbitbrush mixtures occur that also include rubber rabbitbrush
(~. nauseosus) and Parry rabbitbrush (~. parryi).
The mixtures are designated by the symbol "CHY" in type designations.
Antelope bitterbrush ranks fairly close in importance to the aforementioned
composite shrubs in Type 4 associations.
It is most abundant in the area
from Needle Creek east to Long Branch Creek and enhances the forage of deer
Wintering in that locale.
Elsewhere in Unit 67, bitterbrush seems to occur
sporadically and not in great abundance.
Principal Type 4 grasses include Arizona and Thurber fescues as well as
Idaho fescue (K. idahoensis).
The latter is very closely related to Arizona
fescue but is less abundant.
Arizona fescue tends to inhabit lower elevational dry sites whereas Idaho fescue exhibits preference for higher
elevations and better sites.
Similar to the Conifer
Type 4 mixtures.

Type, actinea

is cited once as a type species

in

This 1962 inventory reveals that a m~n~mum of 408 acres of sagebrush and
rabbitbrush cover had been treated by herbicidal spray in measures to convert Type 4 vegetation to grassland.
Up-to-date information received from
Kufeld (1969: pers. comm.) shows that, through January 1, 1970, approximately
15,000 acres of National Forest sagebrush lands in Unit 67 have been treated,
mainly by application of herbicides and some by plowing.
Impact of this
human activity upon forage supplies for wildlife, in general, and big game,
in particular, is nearly impossible to assess.
By any standards, however,
the total acreage receiving treatment must be regarded as being substantial
and of a magnitude to make the environment espeCially less favorable for
deer.

�- 29 Grassland type.--Sagebrush dominates the scene over most of the lower e1evationa1 winter range area below and outside of the National Forest in Unit
67. This situation generally conforms with the vegetational aspect of like
zones of game units north of the main Gunnison River and Tomichi Creek.
A notable exception to the above generality exists in the Cochetopa Park
vicinity.
There, poorer site conditions apparently have allowed grassland
stands to become established and intermingled with sagebrush.
Other
evidence of lessened sagebrush dominance is that sagebrush plants generally
fall short of maximum vigor by averaging lower height and exhibiting lighter
stand densities.
Thus, the Cochetopa Park area firmly hints of being transition territory, fitting somewhere between strongly dominating sagebrush
of the main Gunnison Basin and adjoining 1ike-e1evational areas east of the
Cochetopa Hills divide in the Rio Grande drainage where grassland types rule
non-forested lands.
Of 12 Grassland Types recorded by the survey, seven types feature Arizona
fescue as a principal dominant or codominant and four types each have mountain
muh1y (Muh1enbergia montana) and its close relative slimstem muh1y (~. filicu1mis)
cited in a similar capacity.
Mountain muh1y occurs more charactistica1ly
on
higher, better sites; in contrast, slimstem muhly is usually found on lower,
drier places.
Both species, nevertheless, are frequently mixed in close
association.
Dryland sedges (Carex sPP.) were recorded present with fescues
in one type.
Browse species occurring intermittently but in great enough abundance to rate
type species designations in Grassland Types are Douglas rabbitbrush and big
sagebrush.
Actinea, a forb, is listed as a co- or subdominant species in
three Grassland Types. Another once-listed forb is goldaster (Chrysopsis Spg)
which is symbolized by "CRR".
Broad-leaf tree type.--The Broad-leaf Tree Type (Type 10) in Unit 67 is represented in six types by aspen in pure stands or in associations with conifers.
Codominant coniferous species include Douglas fir, Engelmann spruce, and
limber and ponderosa pines.
One type lists thin1eaf alder (Alnus tenuifolia)
as the principal broad-leaf tree species. Willows (Salix spp.) are cited as
subdominants with alder in this latter type designation.
Thurber fescue, both by itself and mixed with Arizona fescue, is accorded
type species designation in three Broad-leaf Types.
Other herbaceous ground
cover species were most certainly present but not prominent enough in the
general type aspects to warrant recognition.

RANGE TRANSECTS
Browse range condition transects.--Fourteen
transects were established and read
in Unit 67. Of the 14, ten were located on what is classified as key range.
The four other transects were read in areas originally believed to have been
key; but, light use by big game as indicated from pellet group counting and
browse hedging categorizations did not justify establishment of more transects
and/or setting up the areas as key.
All except one transect were located in sagebrush types. The exception was a
transect that was placed in a coniferous "6-DF-PP-Artr" type. Logically, big

�- 30 sagebrush is awarded key species status on a 11 of the transects, since it
clearly dominates compositions throughout.
Bitterbrush is abundant enough
on the Tomichi Creek key area transected types that it was awarded key
species status five times.
Copies of transect records and a summary of pertinent
accompany this report in the binder file folder.

data from them

Pellet group count indexes generally are so low that intensity of occupation
by big game can not be considered a serious influence over most Unit 67
winter ranges within the scope and time of this inventory
Two transects
each in the Needle and Owen Creeks drainages showed that deer had concentrated
in sizable numbers at those specific locations but not over all of the key
area.
Despite obvious anoma lies in distribution, annua 1. utilization surveys
have been and will continue to provide vigilance regarding possible overs t ock Lng ,
0

REFERENCES

CITED

Baker, B. D. 1967. Big game winter range analysis, Game Unit 76 - Rio
Grande.
pp. 55-620
In Game Reso Rep., Colo. Game, Fish and Parks
Div., Denver.
3:1-71 pp. Processed.
Barrell, J
1969. Flora of the Gunnison
Rockford.
494 pp.

Basin.

Natural Land lnst.,

Colorado State Planning Division.
1961.
the State of Colorado.
916 pp.

Colorado

1959-61, Yearbook

o

Fowells, H. A. 1965
Silvics of forest trees of the United States.
Forest Service, U.S.D.A., Agric. Hdbk. No. 271. 762 pp.
0

Preston, R. J., Jr. 1940.
PYess, Ames.
286 pp.

Rocky Mountain

trees.

Iowa State ColI.

January, 1970
Fort Collins

of

�- 31 -

APPENDIX

B

BIG GAME WINTER RANGE ANALYSIS
GAME UNIT 22 - PICEANCE

INTRODUCTION
This report presents results of a big game winter range inventory in
Game Management Unit 22 (Piceance).
Inventory fieldwork and officework were done in cooperative efforts of the BLM Craig District and
Division Federal Aid Project W-101-R in the period from April 1965
through February 1970. Original commitments for the task were made
in 1964 by BLM State Wildlife Specialist Carl Lind and W-101-R Project
Leader Richard Denney, both of whom are presently assigned and/or
employed elsewhere.
Mr. Lind assisted with fie1dcrew orientation and
training and provided program coordination at the state level in 1965.
After 1965,Pau1 Applegate replaced Mr. Lind as BLM State Office coordinator.
Interagency-approved
standard techniques were employed to accomplish the inventory (Denney 1962:51-96).
A large share of the credit for making the program go is due BLM Craig
District Assistant Manager Neil McCleery.
Helped by strong support from'
District Manager Keith Norris, Mr. McCleery furnished all-important
supervision in field and office phases of operations.
Since 1966, District Wildlife Management Biologist Bob Jacobsen has
handled much of the workload previously taken care of by Mr. McCleery
and is commended for his efforts.
Not to be overlooked in these acknowledgments are the many courtesies,
physical facilities, and general assistance furnished by Division Little
Hills Experiment Station Supervisor Bill McKean and Station Foreman Jack
Corey and BLM White River Resource Area Manager Stan Colby.
Finally but
certainly not in the least, the following BLM and Division seasonal and
part-time employees performed both field and office activities related
to type mapping, reading transects, and preparing maps:
Chuck Bullock,
Leroy Carlson, Bob Chavez, Ken Dillinger, Mike Dole, Larry ForbiS, Keith
Giezentanner, Ernie Hafertepen, Craig Ludwig, Bob Moore, Dave Morris,
Charles Robbins, Pete Scorup, and Bill Wolvin.

DESCRIPTION

OF UNIT

Location and size.--Big Game Management Unit 22 is located in westcentral
Colorado within the larger White River Basin geographical province.
A
very narrow strip along the unit's southern boundary lies in Garfield County.
All remaining territory is wholly contained within central Rio Blanco County.

�- 32 In broad terms, State Highway No. 64 along the White River on the north,
State Highway Noo 13 on the east, the crest of the Roan Plateau on the
south, and Hammond Draw and the crest of the Cathedral Bluffs-Calamity
Ridge chain of hills on the west form the unit's boundarieso
Using the more specific description of Unit 22 given in the Colorado
Game, Fish and Parks Division Laws and Regulations Handbook, 1965 (p. 4,
Chapo 3 - Big Game), the boundaries are as follows:
"That portion of Rio
Blanco County south of State Highway No. 64, east of Hammond Draw and the
divide between the Yellow-Piceance
Creeks and Douglas Creek drainages east
to its junction with State Highway No. 13, and west of State Highway No.
13 from this junction south to the Garfield County line; that portion of
Garfield County within the Piceance Creek drainage".
The preceding description is fairly definite except for western portions
of the unit in the vicinity of Hammond Draw and Calamity Ridge southward
to the Roan Plateau.
There, description fuzziness involves the considerable
territory that drains into the White River both east and west of Hammond
Draw and not into either Yellow or Douglas Creeks, which the description
implies.
In addition, to compound confusion Hammond Draw splits into separate and nearly equal forks near the draw's origin on Calamity Ridgeo
By consulting the U.S.G.S. Calamity Ridge Topographic Quadrangle (7.5
minute series; 1962) and BLM 1/2"-scale maps, the author arrived at the
following proposed description for Unit 22 boundaries:
"That portion of
Rio Blanco County south of State Highway No. 64, east of Boise Creek from
State Highway No. 64 to the creek's orlgln on Calamity Ridge, east of a
direct line between the Boise Creek origin and the U.S.G.S. Greasewood
Triangulation Station atop Calamity Ridge, east of the crest of the
Calamity Ridge-Cathedral
Bluffs hills between Greasewood Triangulation
Station and the junction of the Piceance-Douglas
Creeks divide and the
crest of the Roan Plateau, north of the crest of the Roan Plateau between
the latter junction and the Roan Plateau Crest-State Highway Noo 13 junction,
and west of State Highway No. 13 between the Roan Plateau Crest-State Highway
No. 13 junction and the junction of State Highway Noso 13 and 64; that portion of Garfield County within the Piceance Creek drainage".
In moving a portion of the northwest boundary eastward from Hammond Draw
to Boise Creek, some territory would be transferred from Unit 22 to Unit
21; but, the approximately
11 square miles involved would be negligible in
relation to benefits derived from having better defined and more practical
geographical landmarks for boundarieso
Jacobsen (1969) used a dot grid on township plats to arrive at a figure of
998.9 square miles, or 639,319 acres, total area in Unit 220
PhYSical features.--Combined
drainages of Piceance and Yellow Creeks correspond essentially with the area encompassed by Unit 22 boundaries and are
also Virtually synonymous with territory geologists deSignate as the
Piceance Creek Basin.
This basin is a broad structural depression underlain
by rocks of the Wasatch and Green River Formations.
The Wasatch Formation
consists mainly of relatively erodible shales, clay, and some sandstone
layers that form broad valleys or badlands between cliffs or ridges of the

�- 33 -

Green River Formation and the Mesa Verde Group.
The Green River Formation
is composed mostly of shale and marlstone, with some sandstone and limestone also present.
Oil-shale beds occur throughout the Green River
Formation which has a maximum thickness of about 3,500 feet and rocks
underlying mesas that are relatively resistant to erosion (U. S. Department
of Agriculture 1966:9).
As indicated in the previous paragraph, Piceance Creek Basin is an area of
diverse topography.
Differential resistance to erosion of underlying rock
layers has created sharp ridges, low to moderately-high
rounded hills,
abrupt cliffs, and rugged badlands interspersed with open valleys, upland
parks, and smaller basins.
Northerly flowing Piceance and Yellow Creeks and their many tributaries
dissect the landscape.
East of Piceance Creek, principal tributary drainage
systems are from north to south, respectively, Dry Fork, Greasewood Gulch,
Collins Gulch, and Thirteenmile and Fourteenmile Creeks.
Ryan Gulch, Black
Sulphur Creek, and Stewart Gulch drainages are just a few of many entering
Piceance Creek from the west and south. Again, north to south, respectively,
Yellow Creek has the main tributary drainages of Greasewood, Barcus, and
Duck Creeks feeding it from the West.
Westward flowing White River, at Unit 22's northern boundary, has several
generally short drainageways joining it from the south. Among the latter
from east to west, in order, are Sheep Creek (along and near State Highway
No. 13), Kendall and Hay Gulches, and Little Spring and Monument Creeks,
as well as aforementioned Boise Creek and Hammond Draw.
The reservoir on Division-owned Rio Blanco State Recreation Area (Rio Blanco
Lake) contains a maximum of 115 surface acres of dead-water storage and is
the only large body of water in Unit 22. Several small ponds, variously
used for irrigation, flood water control, stock water storage, and trout
fishing, are scattered throughout Unit 22.
As might logically be expected, basin-rim high ground on western, southern,
and eastern perimeters has the greatest elevations.
On the west, maximum
elevations include 8,420 feet near the origin of Black Sulphur Creek and
8,685 feet at the head of Big Duck Creek. At the south edge, there are at
least five high points on the Roan Plateau exceeding 8,400 feet, two of which
best 8,700 feet. Greatest elevation in the eastern part is reached at the
eastern end of Joe Bush Mountain.
Elevation there is 8,371 feet. Several
other ridges and mountains north and south and in the vicinity of Joe Bush
Mountain reach heights just below or above 8,000 feet, Kendall and Segar
Mountains included.
Climate.--Unit 22's highly variable climate can generally be described as
semiarid in having comparatively warm summers and cold winters.
Extremes
are due to moderately wide variations in elevations and exposures.
Annual
frost-free periods range from about 120 days at the unit's northwest corner
near the White River to less than 50 days at high elevations of the basin
rim.
Mean annual precipitation varies from about 12 to 25 inches.
It is distributed fairly uniformly throughout the year, with a little less than one-half

�- 34 usually occurring with the December-April period.
Summer thundershower
activity at higher elevations is typically of the gentle variety.
In
contrast, rain showers at lower elevations are affected by convective
conditions conducive to production of cloudbursts that result in highpeak-flow floods of short duration.
Preceding generalizations about Unit 22 climate are based upon information contained in a U.S. Department of Agriculture Report on the White
River Basin (1966:6,12-27).
Descriptions of local Piceance Creek Basin
climates are also included in the U.S. Department of Agriculture (1966)
report's discussion of soils which are condensed, in part, in following
paragraphs.
Soils.--There are four broad soil mapping units represented in Piceance
Creek Basin.
The largest unit (Unit 3) is composed of what are termed
"shallow rocky and deep moderately dark colored soils of the uplands".
This unit corresponds with lowest portions of the basin at elevations of
between about 6,000 and 7,000 feet. Slope gradients usually range between
10 and 60 percent.
Here the annual precipitation averages 12 to 15 inches;
the mean annual temperature is about 470 F; and, the frost-free period is
around 100 days. Summer storm waters frequently cause much damage to
bridges and stream crossings, create deep gullying and head cutting, and
leave flood deposits that cover vegetation on alluvial fans and bottomlands (U. S. Department of Agriculture 1966:12,13,17-19).
Considerably smaller in extent than Soil Mapping Unit 3 in Piceance Creek
Basin is Unit 4. Unit 4 is called the "Deep moderately dark colored soils
of the mesas and va lleys". Powell Park and Josephine Basin, just west and
southwest of Meeker, are represented by Unit 4 soils. Elevationa1 range
of these locations is 6,000 to 6,600 feet. Meeker has an average annual
precipitation of about 17 inches, a frost-free period of 94 days, and a
mean annual temperature of 440 F. Slope gradients of Unit 4 bottom1ands,
upland benches, and mesas range from less than five percent to about 15
percent.
Despite lower slope gradients, water erosion is a problem around
Meeker (U. S. Department of Agriculture 1966:19,21).
Soil Mapping Unit 5 soils rank second in extent to Unit 3 and generally lie
above Unit 3 and below Unit 7 in Piceance Creek Basin.
Unit 5 is termed the
''Moderately deep and deep, dark colored soils of the uplands".
Steep slope
gradients within the range of 10 and 60 percent are most common.
Unit
elevationa1 range is between 7,000 and 8,000 feet with extremes of 6,400
feet and 8,800 feet. Greater precipitation and lower mean annual temperatures
are associated with the greater elevations.
Annual precipitation ranges from
15 to 20 inches, and the frost-free period averages from 45 to 100 days.
Erosion is slight to moderate and generally confined to washing along stock
trails and minor gullying along drainageways (U. S. Department of Agriculture
1966:21-23).
The "Dark colored soils of the cold, forested mountain slopes" categorize
Unit 7 soils. This unit, being restricted to a narrow strip along the crest
of the Roan Plateau on Piceance Creek Basin's southern edge, is not very
extensive.
The landscape consists of ridges and steep lower mountain slopes
and valleys.
Slope gradients usually range from 10 to 50 percent at elevations

�- 35 -

between 8,000 and 8,800 feet. Mean annual precipitation ranges from 20 to
o
25 inches; mean annual temperature is less than 40 F; and, frost-free
growing seasons average less than 50 days.
Like in Unit 5 soils, erosion
is comparatively slight (U. S. Department of Agriculture
1966:25-27).
Industry.--Production
of oil and natural gas is by far the greatest contributor to Rio Blanco County industrial wealth.
In fact, the county
ranked first in 1963 Colorado yield of those fuels (U. S. Department of
Agriculture 1966:10).
Total value of "petroleum" in 1960 is listed at just
over $57 million (Colo. State Planning Division 1961:874).
Principal oil fields in Rio Blanco County are located near Rangely and in the
Wilson Creek area, both outside of Unit 22. However, Piceance Creek drainage
in the unit is the major gas-producing area in Rio Blanco County.
Most wells
there yield gas from the Green River and Wasatch Formations (U. S. Department
of Agriculture 1966:10).
Production and sale of livestock and livestock products comprise the second
leading industry in the county.
Income from those sources in 1959 totaled
over $3 million (Colo. State Planning Division 1961:874).
Cattle, sheep,
and horse ranches in the valleys of the White River and Piceance, Yellow,
and Sheep Creeks provide most of the livestock production.
Income from secondary minerals such as sand and gravel, uranium ore, and
coal, respectively, follow livestock in importance (U. S. Department of
Agriculture 1966:11).
Unit 22 has no incorporated towns or cities.
Meeker, county seat of Rio
Blanco County, is located about 2 miles east of Unit 22's northeast corner;
near the unit's northwest corner is Rangely, home of Rangely Junior College.
According to the U.S. Census Bureau (pers. comm. 1970), Meeker dropped in
population from 1,655 in 1960 to 1,536 in 1970 which contrasts with a gain
by Rangely (1,464 in 1960 to 1,638 in 1970). These population changes are
partially compensating and indicate a possible slight increase in the total
county population.
Both cities serve as trade centers for ranching, oil
and gas areas, and tourist business (Colo. State Planning Division 1961:
798-799).
Projected future developments of oil shale, agriculture, water, and recreation
resources and associated increases in human population of the White River
Basin are summarized by the report of the U.S. Department of Agriculture
(1966:73-92).
Therefore, only one comment seems appropriate here relative
to those subjects.
Increased activities above those of the present population density of 1.5 people per square mile now in the White River Basin
will simply mean more adversity for wildlife, in general, and deer, in
particular.

LIMITS OF AVERAGE

WINTER RANGE

From the beginning of the field investigations phase of this inventory, it
was obvious that determining average upper limits of the big game winter zone
would be difficult.
Continuous, intensive air and ground observations of big

�- 36 game distribution were lacking.
In substitution, pellet group count
indexes and browse age and form class determinations were relied upon to
establish where deer and elk were found most years within the period of
November 10 to April 20.
The aforementioned
5-l/3-month span accounts for the time that the major
portion of wintering deer are present in Unit 22. There is usually some
migration after November 10 and April 20, but most movement ordinarily
Occurs prior to both dates.
No attempt was made in the inventory to delineate summer and transitional
ranges, although Jacobsen (1969) logically treats summer range as the
difference between total unit area and that classified as winter range.
The foregoing procedure is questionable for some locales in the vicinity
of the upper winter line where ranges might more properly be called yeararound ranges. Nevertheless, the latter instances probably are of minor
importance because summer population densities and resultant effects on
vegetation are insiginificant in the total ecology of the deer.
Already implied, but not specified, is the finding that winter zone upper
limits apply almost universally to deer populations.
Elk have not been
plentiful enough anywhere in Unit 22 to have influenced placement of a
winter range boundary line. It is conceivable, however, that future
increases of elk numbers are likely and will complicate management of both
big game species in the unit.
Some medium to high elevation, north and east facing slopes at the east
end of Unit 22 (on and/or near Kendall and Segar Mountains) were found to
have received light winter use. These areas were first excluded as winter
range.
Upon further deliberation, the areas were included for two reasons.
First, small area map exclusions were thought to be possible future sources
of confusion.
And secondly, to show these island exclusions might imply
unimportance which we wanted to avoid.
The slopes in question are mostly
dense stands of high-value browse that help effect better deer nutrition
and distribution in less severe winters.
Finally, within limitations imposed by information available and that
obtainable by procedures used in the field the summers of 1965-67, following
paragraphs will describe the location of the average upper winter range
limit line as it appears on accompanying 2-inch scale maps. Elevational data
were obtained from U. S. Geological Survey Topographical Quads upon which the
winter line was transposed.
Yellow Creek area.--In his report, Jacobsen (1969) gives the mean elevation
of the winter range line for the Yellow Creek portion of Unit 22 at about
6,850 feet and the mean low elevation within the winter range of about 6,000
feet. It should be made clear that these elevational means apply to all
of the area west of Piceance Creek and not just to the Yellow Creek drainage.
Starting at the northwest corner of Unit 22, the upper winter line is located
near the surprisingly high maximum elevation of 8,000 feet on the north end
of Calamity Ridge. Wind-swept, comparatively snow-free conditions atop the
ridge probably account for deer using this locale. Also in the same vicinity
on the leeward northeast side of Calamity Ridge, the winter range line drops

�- 37 down to and roughly follows the 7,400-foot contour at the heads of Hay
Canyon, Little Spring Creek, and Trail Gulch.
From Trail Gulch, the line
drops sharply to the bottom of North Fork Greasewood Creek at a pOint about
one mile east of Thompson Spring; then it goes northeastward along the
North Fork bottom, varying from about 7,000 feet to a low of 6,400 feet,
and excluding the uor t hwes t; slopes of the dividing ridge between the North
and Middle Forks of Greasewood Creek.
Southeast slopes into the Middle Fork-East Fork Greasewood Creeks drainage
are included as winter range, elevations there also varying between 6,400
and 7,000 feet. The winter range line traverses the divide between East
Fork Greasewood and North Barcus Creeks at about the 6,600-foot level,
swings southwest up North Barcus and southeast back down Middle Barcus
Creeks to include almost all of the area between the two creeks below an
elevation of about 7,000 feet. Exceptions are places near the head of
North Barcus Creek that have elevations near 7,800 feet.
From the confluence of Middle and Main Barcus Creeks, the winter line
locates south and southwestward, mostly across contours, to intercept Duck
Creek's main forks near their junction.
Elevations vary from about 7,400
feet at the Middle-Main Barcus Creeks' junction to 6,500 feet where the
principal forks of Duck Creek jOin.
The winter zone south of Duck Creek includes most of 84 and Bar D Mesas,
with necks that project southwestward up Corral Gulch and Stake Springs
Draw above the old 84 Ranch headquarters.
Excluded in this vicinity, however, is nearly all of Wolf Ridge between Corral Gulch and Stake Springs
Draw.
The winter line joins Ryan Gulch about one-half mile west of the
old Ryan School.
Elevations of the line between Duck Creek and Ryan Gulch
vary within the range of 6,500 to 7,000 feet.
Ryan Gulch-Black Sulphur Creek-Stewart Gulch area.--Drainageways
of these
gulches, and drainages paralleling them in between, flow northeastward and
northward into Piceance Creek. Jacobsen (1969) included this area within
his report from which mean winter range elevations were cited previously
(see first paragraph herein under the foregoing Yellow Creek area heading).
Continuing discussion of winter range line location, the line is placed to
include lower southeast slopes into the Ryan Gulch bottoms from near Ryan
School southwestward to the multi-confluence
of Ryan, Galloway, Dry, and
Wagonroad Gulches.
Upland terrain between Ryan Gulch and Black Sulphur
Creek is Wagonroad Ridge.
Ridge territory is winter zone that lies northeast of the winter line extending roughly between Reigle Ranch on Ryan
Gulch and Duckett Ranch on Black Sulphur Creek.
Elevations reach a maximum
of about 7,100 feet atop the ridge and minimums of 6,600 feet on both Ryan
Gulch and Black Sulphur Creek.
Eastward from Black Sulphur Creek, in order, the dividing ridge between
Black Sulphur Creek and Yankee Gulch is excluded as winter range; conversely,
most of the lower Yankee Gulch-Eureka Creek upland is classified as winter
zone; and again, all of the dividing ridge between Eureka and Fawn Creeks
and the Fawn Creek bottoms is excluded as winter range.
From Fawn Creek eastward, the winter line is situated to include as winter
zone both upland and bottomland lower territories of Little Dry and Dry

�- 38 Gulches, Hunter Creek, Enoch and Big Jimmy Gulches,
Gulch, and West and Middle Stewart Gulches.

Willow

Creek, Scandard

The winter line crosses East Stewart Gulch at about 6,400 feet elevation,
drops down East and main Stewart Gulches on their northeast bottom-margins,
and touches Piceance Creek just east of the Walter 01d1and Ranch.
As already indicated, elevations of the upper winter line between Ryan and
Stewart Gulches are quite variable.
This is because drainage bottoms, mesas,
and ridges are irregularly included within the winter zone. In general,
however, maximum elevations vary around 7,000 feet and minimums about 6,400
feet, conforming with Jacobsen's (1969) overall mean of 6,800 feet.
Upper Piceance Creek-Fourteenmile
Creek-Thirteenmile
Creek-Sheep Creek area.-The winter line coincides with Piceance Creek from near the Walter 01d1and
Ranch southeastward upstream to a point about two miles west of Rio Blanco
Post Office.
Thus, the entire southeastern corner of Unit 22 lying south
of Piceance Creek and east of main Stewart Gulch is outside of the winter
zone.
From west of Rio Blanco at about 7,200 feet elevation on Piceance Creek,
the winter line goes almost due north in crossing contours to include the
heads of Coyote, Deer, and Davis Gulches.
Maximum elevations in this vicinity
reach about 8,000 feet. Northeast slopes of upper portions of the dividing
ridge between Davis Gulch and Fourteenmile Creek are excluded by the line
which drops into the latter stream at its confluence with Dark Canyon (not
to be confused with Dry Fork tributary of same name).
Thence, the line is
located in Dark Canyon, across Fourteenmile Ridge, and down to Thirteenmi1e
Creek near the Cox Ranch.
Elevations in this area reach 8,100 feet on
ridgetops and between 6,840 and 7,260 feet in bottoms of Fourteenmile and
Thirteenmile Creeks, respectively.
On the north side of Thirteenmile Creek, the winter line extends westward
downstream from near the Cox Ranch to the confluence of Open Gulch and
Thirteenmile.
The line then follows the bottom of Open Gulch and over the
ridge into Timber Gulch about 2-1/2 miles above the confluence of Joe Bush
and Timber Gulches.
As a consequence, the Goat Trail Ridge, Rough Gulch,
Rough Ridge, Hyberger Gulch, and Bald Knob areas and territory east to
State Highway No. 13 are all excluded as winter range.
The winter line bisects Joe Bush Mountain in crossing northeastward from
Timber Gulch into Joe Bush Gulch at about the 7,000-foot level. From the
latter pOint, the line goes up the bottom of Joe Bush Gulch, loops around
the extreme eastern end of Segar Mountain, and traverses back into Segar
Gulch near its origin.
This is all high ground, having elevations ranging
between 7,700 and 8,100 feet along the winter line.
From near the origin of East Fork Hay Gulch, the line angles northeastward
to its unnamed opposing drainage on the Sheep Creek side of the ridge. There
it joins and generally coincides with the 7,200-foot contour southward-bound
until the contour intercepts State Highway No. 13 about nine miles north of
Rio Blanco.
This is also the line's exit point from Unit 22 and falls
slightly north of the Sheep-Thirteenmile
Creeks divide on State Highway No. 13.
It is apparent from foregoing paragraphs that most of the area in Unit 22
east of Piceance Creek (Little Hills Triangle) is winter range.
Classification

�- 39 -

of various high-ground portions as winter range in this eastern sector
of the unit might be questionab1eo
However, when doubts occurred in the
course of fieldwork, areas were usually credited as being winter range
rather than allow an omission to occuro
Genera1.--In contrast to a mean elevation of about 6,800 feet west of
Piceance Creek, mean elevation of the winter line east of Piceance Creek
is estimated to be 7,400 feet between extremes of 6,400 and 8,100 feet.
It is logical to assume that exposure is the primary reason most of the
Little Hills Triangle is usable winter rangeo
Drainages and intervening
ridges are oriented either north and south or east and west and have steep
slope gradientso
As discussed by Yeager (1960:4-6)~ these slope and
topographic factors provide for maximum insolation benefits and territory
for use by wintering animals.
At no point does a lower limit line enter Unit 22 because
is open-ended at the north on the White River.

the winter

zone

KEY AREAS
Winter range of Unit 22 was divided into seven key area subunits based upon
geographical features.
Primary purpose of this partition was to satisfy
the need for localized place references with which to tie transect recordso
The agreement had previously been reached that all usable winter range
conformed with the definition of and could justifiably be called keyo
Descriptions of the seven key areas are presented in following paragraphs
and shown graphically on accompanying 1/4-inch scale unit mapo
Rattlesnake key areao--A1l winter ranges lying south of State Highway No.
64, east of Piceance Creek, north of Dry Fork Piceance Creek, and west of
East Fork Hay Gulch and the VT Trail.
Greasewood key area.--A11 winter range bounded on the west and south by
Piceance Creek, east by Collins and Little Corral Gulches, and north by
Dry Fork Piceance Creek.
Thirteenmi1e key area.--A11 winter range that lies north of Piceance Creek,
east of Collins-Little
Corral Gulches, south of Dry Fork Piceance Creek
and Timber Gulch, and west of the winter range line already described between
Timber Gulch and upper Piceance Creek.
Segar Mountain key area.--A11 winter range bounded on the west and south by
East Fork Hay Gulch, VT Trail, Dry Fork Piceance Creek, Timber Gulch, and
the winter range line between Timber Gulch and State Highway No. 13, east
by State Highway No. 13, and north by State Highway No. 64.
Hammond-Barcus key area.--A11 winter range bounded on the north by State
Highway No. 64 between Hammond Draw and Yellow Creek, west by Hammond Draw
and the winter range line between Calamity Ridge and Barcus Creek, south by
Barcus Creek, and east by Yellow Creek between its confluence with Barcus
Creek and junction with State Highway No. 64
0

�- 40 Barcus-Ryan key area.--A11 winter range bounded on the west by Yellow Creek
between its junction with State Highway No. 64 and confluence with Barcus
Creek, and the winter range line between Barcus Creek and Ryan Gulch, south
by Ryan Gulch, east by Piceance Creek, and north by State Highway No. 64
between Piceance and Yellow Creeks.
Ryan-Story key area.--A11 winter range bounded by Ryan Gulch on the north,
the winter range line between Ryan and main Stewart Gulches on the west
and south, and Piceance Creek on the east and north.
Story Gulch was first believed to have been the eastern terminus of winter
range south of Piceance Creek; but, field reconnaissance proved that the
terminus was at main Stewart Gulch about four miles west of Story Gulch.
Story (Gulch) was retained in the name because transect records already
contained it through the field season and chances were slight for future
conflict by its retention.

LANDOWNERSHIP
Federal Public Lands under jurisdiction of the U. S. Bureau of Land Management compose the largest landholding of winter range in Unit 22 (Table 1).
Approximately
84.5 percent falls into that category.
Private landowners
control the second largest block (30,595 acres, or 9%), and the Colorado
Division of Game, Fish and Park's lands under patent equal about 6-1/2 percent (21,926 acres) of the total winter range.
Relative to the latter
acreage, there seems to be little doubt about the value of Division aims
for acquiring and developing land in Unit 22 for big game, just as there
should be no question about BLM responsibilities and evident concern in
maintaining equitable deer populations on the great expanse of land under
their jurisdiction.
Tables 2 and 3 present summaries of vegetation type acreages by landownership for the Yellow Creek (west Piceance) and Little Hills Triangle (east
Piceance) portions of Unit 22 winter range.
In discussions to follow, some
gross comparisons of the two parts will be made for which the tables will
serve as useful references.

VEGETATION

TYPES

Unit 22 is top-ranked in hunter harvest of mule deer in Colorado (Table 4).
Importance of the Piceance-White River deer herd has long been known.
Indeed,
some of the first efforts of the Division's Federal Aid Section were directed
towards evaluating deer numbers and forage interrelationships
of what Carhart
(1943:18) termed the "most important flock of 'wild livestock' in Colorado".
Therefore, in light of winter range that must be considered as close to
optimum as any in Colorado, vegetation types are discussed in following
paragraphs to the end that amounts, characteristics, and proportions be used
as standards for better deer management other places in western Colorado.

�Table 1. Vegetation and land category acreage summary, big game winter range, Game Unit 22 (Piceance).
Landownership
TOTAL

PRIVATE

GF&amp;P

BLM

%

Total

No.
Acres

Total

224

0.8

2,918

0.9

22.2

8,044

26.3

71,760

21.2

1,006

4.6

6,457

21.1

52,881

15.6

0.3

0

0.0

62

0.2

852

0.2

171,129

59.8

12,739

58.1

7,649

25.0

191,517

56.6

13 - Saltbush

3,699

1.3

46

0.2

129

0.4

3,874

1.2

14 - Greasewood

1,599

0.6

1,868

8.5

249

0.8

3,716

1.1

688

0.2

0

0.0

0

0.0

688

0.1

1,168

0.4

1,394

6.4

7,781

25.4

10,343

3.1

338,529*

100.0

Total

No.
Acres

0

0.0

20.6

4,873

45,418

15.9

790

Total

No.
Acres

2,694

0.9

4 - Sagebrush or Rabbitbrush

58,843

5 - Browse
6 - Conifer

No.
Acres

1 - Grassland

Type or

Land Category

%

%

%

....

.f:'-

9 - Pinyon-Juniper

18 - Annuals
Cultivated and Hay
Totals

286,008

100.0

--

-

21,926

100.0

30,595

100.0

* This does not jibe with total winter range acreage as reported in Jacobsen (1969). Discrepancies
to later map revisions which had not been updated in the BLM report as of this writing.

are due

�Table 2. Vegetation and land category acreage summary, big game winter range, Yellow Creek (or Piceance
Creek West) part, Game Unit 22 (Piceance).
"';'\

Landownership
BLM

GF&amp;P

PRIVATE
No.
%
Acres
Total

TOTAL
No.
%
Acres
Total

Type or Land Category

No.
Acres

Total

No.
Acres

Total

1 - Grassland

2,564

1.6

0

0.0

0

0.0

2,564

1.4

4 - Sagebrush or Rabbitbrush

36,423

22.1

3,209

29.0

2,465

22.7

42,097

22.6

5 - Browse

1,594

1.0

74

0.7

58

0.5

1,726

0.9

117,150

71.2

4,981

45.0

4,671

43.1

126,802

68.0

13 - Saltbush

3,673

2.2

0

0.0

129

1.2

3,802

2.0

14 - Greasewood

1,432

0.9

1,769

16.0

234

2.2

3,435

1.8

668

0.4

0

0.0

0

0.0

668

0.4

1,030

0.6

1,039

9.3

3,282

30.3

5,351

2.9

100.0

10,839

100.0

186,445

100.0

9 - Pinyon-Juniper

%

%

~

N

18 - Annuals
Cultivated and Hay

--

Totals

164,534

100.0

11,072

~'~These acreages do not jibe with those reported in Jacobsen (1969). Discrepancies are due to later map
revisions which have not been updated in the BLM report as of this writing.

�Table 3. Vegetation and land category acreage summary, big game winter range, Little Hills Triangle (or
Piceance Creek East) part, Game Unit 22 (Piceance).
Landownership
TOTAL

PRIVATE

GF&amp;P

BLM

%

Total

No.
Acres

Total

224

1.1

354

0.2

15.3

5,579

28.2

29,663

19.5

932

8.6

6,399

32.4

51,155

33.6

0.7

0

0.0

62

0.3

852

0.6

53,979

44.4

7,758

71.5

2,978

15.1

64,715

42.6

13 - Saltbush

26

0.0

46

0.4

0

0.0

72

0.0

14 - Greasewood

167

0.1

99

0.9

15

0.1

281

0.2

Cultivated and Hay

138

0.1

355

3.3

4,499

22.8

4,992

3.3

Total

0

0.0

18.5

1,664

43,824

36.1

790

Total

130

0.1

4 - Sagebrush or Rabbitbrush

22,420

5 - Browse
6 - Conifer

Type or Land Category

1 - Grassland

9 - Pinyon-Juniper

%

121,474

100.0

%

.j::-

w

10,854

100.0

--

--

-

Totals

%

No.
Acres

No.
Acres

No.
Acres

19,756

100.0

152,084

100.0

�- 44 -

Table 4. Fifteen game management units with the highest total deer harvest,
ten-year period, 1959-68.
Unit Number

Name

Harvest

Rank

22

Piceance

57,137

1

42

Grand Mesa

45,078

2

31

Roan Creek

39,061

3

21

Douglas Creek

35,783

4

62

East Uncompahgre

34,548

5

32

Parachute

32,677

6

11

Strawberry

28,913

7

44

Brush Creek

27,638

8

33

Rifle

27,087

9

23

Miller Creek

26,245

10

40

Glade Park

25,641

11

70

San Miguel

24,880

12

43

Roaring Fork

24,875

13

71

Dolores

21,907

14

78

San Juan

21,183

15

�- 45 Pinyon-juniper type.--Ru1ing the vegetative mantle of Unit 22 winter range
is the Pinyon-Juniper Type (Type 9). Pinyon-juniper
is a woodland or foothills zone climax (Oosting 1956:296,303-304)
and is here represented in
types by approximately equal portions of Utah juniper (Juniperus osteosperma)
and pinyon pine (Pinus edu1is).
Rocky Mountain juniper (1. scopu10rum) is
also found in Type 9 associations but is sparsely abundant.
Type 9 dominance is shown in Table 1, although it is actually underplayed
there and on Table 3. If the approximately 39,000 acres of the 4- and 5-PJ
transition types found east of Piceance Creek are added to Type 9 acreages
on Tables 1 and 3, resulting totals are equal to the same 68% component
shown in Table 2.
Thus, with about 68% of the winter range covered by Type 9, it is appropriate
that all kinds of slopes, exposures, and terrains support this vegetation.
Some Pinyon-Juniper Types contain as much as 5,000 acres, with other blocks
frequently having over 1,000 acres.
Fortunately for interrupters like sagebrush, greasewood, and saltbush in drainage bottoms, upland sagebrush parks,
and upland browse types (particularly east of Piceance Creek) the vast
expanses of pinyon-juniper are broken up and more valuable for big game
naturally.
Significance of pinyon pine and junipers as cover for mule deer is widely
recognized and accepted.
Less well understood, however, is the presence of
these conifers in deer diets.
In Piceance Creek Basin, Carhart (1941: 9-11)
reported that pinyon occurred in greater amounts than did junipers in 37
winter- and 30 spring-collected
stomach samples.
More specifically, pinyon
comprised 21.41% of winter (Dec.-Feb.) diets and ranked second to big sagebrush (Artemisia tridentata).
Pinyon pine dropped to 7.20% of spring
(Mar.-May) diets and a ranking of sixth.
The same study showed that junipers
ranked third and lOth, respectively, as components of winter and spring diets.
Elsewhere in the Rockies, central and southern studies demonstrated that
junipers are components of mule deer diets (Smith 1952:153, Hill 1956:395,400,
Smith 1959:8-13, Anderson et al. 1965:353-364), but nutritional functions
of conifers are unknown and unexplained by researchers.
Suffice to say that
more than enough pinyon pine and juniper plant material probably exists to
satisfy all deer dietary demands in Unit 220
Another codominant evergreen tree occurring with pinyon-juniper
is Douglas
fir (Pseudotsuga menziesii).
This association is not common, as evidenced
by only two types totalling 2,300 acres in the vicinity of Thirteenmi1e,
Fourteenmi1e, and Davis Gulch areas of upper Piceance Creek.
From frequency of appearance in type designations and acreage involved, the
most common browse species in approximate order of abundance in Unit 22
Pinyon-Juniper Type vegetation are as follows:
1.

True mountainmahogany
to mountainmahogany,

(Cercocarpus montanus), hereafter shortened
25 different types for 105,115 acres.

2.

Big sagebrush,

3.

Utah serviceberry (Amelanchier utahensis), hereafter
as serviceberry, 20 types for 44,071 acres.

15 types for 87,093 acres.
referred

to

�- 46 4.

Antelope bitterbrush (Purshia tridentata),
brush, 9 types for 27,232 acres.

also called bitter-

Other important but less abundant browse type-species include Douglas and
dwarf rabbitbrushes (Chrysothamnus viscidiflorus; ~. depressus), black
sagebrush ~. nova), snowberry (Symphoricarpos sp.), Gambel oak (Quercus
gambeli) , and shadscale saltbush (Atriplex confertifolia).
The rabbitbrushes, snowberry, and black sagebrush are fairly abundant and widely
distributed.
Gambel oak is found almost exclusively east of Piceance
Creek.
Shadscale saltbush is not connnon with pinyon-juniper as indicated
by one type of about 600 acres having the species.
There are several unconnnon shrubby species associated with pinyon pine and
juniper that did not rate type species designation.
They include rubber
rabbitbrush ~. nauseosus), connnon winterfat (Eurotia lanata), bush rockspirea (Holodiscus dumosus), currants (Ribes sPP.), connnon chokecherry
(Prunus virginiana), gray horsebrush (Tetradymia canescens), ephedra
(Ephedra sp.) bush eriogonum (Eriogonum sp.), creeping mahonia (Berberis
repens), black greasewood (Sarcobatus vermiculatus), and cottonthorn horsebush (!. spinosa).
Martin ceanothus (Ceanothus martini) and greenleaf
manzanita (Arctostaphylos patula) are very rare and only known to exist at
the north end and top of Calamity Ridge.
Grass and forb cover is characteristically
light under pinyon-juniper overstory. Needleandthread
(Stipa comata) and wheatgrasses (Agropyron spp.),
latter symbolized by "AGR" and including mostly bluestem and beardless
bluebunch wheatgrasses ~. smithi; !. inerme), are abundant and prominent
enough, nevertheless, that they were named in seven types totalling about
8,000 acres.
Other very connnon grasses found with pinyon-juniper but not
rating type species designation are prairie junegrass (Koeleria cristata),
Indian and littleseed ricegrasses (Oryzopsis hymenoides; Q. micrantha),
bottlebrush squirreltail (Sitanion hystrix), and bluegrasses (Poa spp.).
Despite fair abundance and variety in pinyon-juniper associations, forbs
are lacking as type species.
However, several broad-leaf herbs were recorded in browse range transecting work. They include broom snakeweed
(Gutierrezia sarothrae) (also called a half-shrub), goldenweeds, (Haplopappus)
sPP.), phloxes (Phlox sPP.), arrowleaf balsamroot (Balsamorhiza sagittata),
yarrow (Achillea sp.), scarlet globemallow (Sphaeralcea coccinea), prickly
pear (Opuntia sPP.), and lupines (Lupinus spp.).
Sagebrush or rabbitbrush type.--Sagebrush or Rabbitbrush (Type 4) is the
second most extensive vegetation type in Unit 22, ranking thus on the basis
of strong representation west of Piceance Creek (Tables 1, 2). The Browse
Type east of Piceance Creek is more extensive than sagebrush (Table 3), but
the acreage advantage there is not great enough to reverse the situation
for the entire unit (Table 1).
Big sagebrush is strongly the dominant plant of Type 4 acreages.
It lacks
being listed as a type species only once in 36 different type designations
(that one time a 4-CHY type of 71 acres).
Furthermore, big sagebrush is
probably the most abundant woody species in all of the winter range on the
strength of being a type species in 73 of 143 total types in Unit 22.

�- 47 Importance of big sagebrush in the nutrition of Unit 22 deer can not be
overemphasized.
Applicable to this same area, Carhart (1941:9-11), reported that big sagebrush comprised about 37% and 27%, respectively, of
winter and spring rumen samples to lead forage consumed those seasons.
Moreover, big sagebrush composed 15% of fall rumen to rank second for that
season behind serviceberry which was 18%. Sagebrush failed to qualify as
year-round food, since the same study found big sagebrush absent from deer
rumens obtained in summer.
Big sagebrush very often exhibits tall, rank growth on bottomland sites
that are characterized by deep well-drained soils.
Site factors rather
than varietal differences evidently cause greater size. Beetle (1960:52)
said that size of big sagebrush "is a simple function of density and
favorableness of site conditions".
He added that big sagebrush is widely
distributed over a great diversity of sites but does avoid shallow soils.
Except for heavy consumption of grasses in spring, deer apparently use the
robust bottomland sagebrush primarily for cover.
Very closely associated with sagebrush in Unit 22 are rabbitbrushes which
rank as the second most abundant woody plant complex of Type 4 acreages.
On accompanying maps, forms, and data summaries, the type symbol "CHY"
stands for mixtures of Chrysothamnus composed principally of Douglas and
rubber rabbitbrushes.
Recalling that Type 4 is also equivalent to the
Rabbitbrush Type, there are three of these latter types that total 398
acres.
"CHY" dominated acres all occur on Yellow Creek bottoms.
East of
Piceance Creek there is one type of 1,247 acres which lists "CHY" secondarily
or codominantly.
Overall, Douglas rabbitbrush is probably the second most abundant shrubby
species in Type 4 acreages.
Basis for this reasoning is that Douglas rabbitbrush is listed as a type species in types containing over 16,700 acres
which are in addition to the approximately 1,600 acres mentioned in the
previous paragraph also containing Douglas rabbitbrush.
Ranking browse species abundance after Douglas rabbitbrush in Type 4
acreages is difficult.
Both rubber and dwarf rabbit brushes contribute substantially to the total shrubby component, with the former probably furnishing
more total volume in Unit 22. Dwarf rabbitbrush is given type species designation in one sagebrush type of 450 acres.
On the basis of type acreage representation, pinyon pine and junipers probably
follow sagebrush and rabbitbrush in volume of Type 4 plant materials present.
In particular, there are eight different sagebrush/pinyon-juniper
types
totalling 6,400 acres.
Caution should be exercised here, however, for permitting much credence on 4-PJ (and 4-J) acreages as indexes to abundance.
Crown cover percentages of pinyon-juniper average 10% or less when 4-PJ, or
4-J, are used in type designations, thus causing possible error through
overestimation.
Following pinyon pine and junipers in order of probable abundance in Type 4
associations is serviceberry.
It was recorded as a codominant type species
in four type designations that total 4,084 acres.
Behind serviceberry in
decreasing approximate order of abundance are: Gambel oak, shadscale saltbush, black greasewood, common winterfat, and snowberry, each represented by
minor acreages.

�- 48 Grasses account for all herbaceous type species plants occurring with sagebrush.
Wheatgrasses,
particularly bluestem singly and mixed with beardless
bluebunch, predominate.
The exotic Fairway crested wheatgrass ~. cristatum),
widely used in revegetation projects, was abundant enough to be listed as
a type species in one type of 59 acres
Next to wheatgrasses in abundance,
the very common needleandthread
provides excellent livestock forage.
Other
notable grasses are blue grama (Bouteloua gracilis), Sandberg bluegrass
(R. secunda), giant wildrye (Elymus cinereus), and cheatgrass brome (Bromus
tectorum,) which are all listed as type species at least once; prairie
junegrass, Indian ricegrass, and bottlebrush squirreltail are commonly present but not given in type designations.
o

Browse type.--The Browse Type (Type 5) is the third most extensive vegetation
type on Unit 22 winter range.
There are about 53,000 acres, or 16% of total
winter range in the type. Most of the 53,000 acres are found east of Piceance
Creek (Table 3).
Serviceberry is almost certainly the most abundant browse species in Unit 22
Type 5 vegetation, because it is a type species in 35 different types that
total 51,878 acres.
The preceding acreage lacks about 1,000 acres of that
for all of Unit 22 Type 5 lands (Table 1).
Due to just cited excellent abundance, coupled with high forage preference
by deer, serviceberry is uniquely important to the Piceance Basin deer herd.
Again referring to a food habits study, Carhart (1941:9-11) reported that
serviceberry was a major year-round component of mule deer diets on the White
River.
This shrub composed 17.6% and 54.2%, respectively, of fall and summer
collected rumens to be the leading forage those two seasons.
Serviceberry
ranked fifth in amount of winter forage consumed (7.5%) and second in spring
(20.8%); so, there should be little question about its preferability.
The
plant is plentiful and used, and application of the term "key" to a species
could never be more valid for a broad plant type than here.
Deer forage quality of Unit 22 winter range excels not only because of
serviceberry.
Additional important staples such as big sagebrush, mountainmahogany, and Gambel oak, respectively, augment serviceberry in 15 types of
39,286 acres, 17 types of 20,980 acres, and 15 types of 18,276 acres.
Other minor shrubs abundant enough in Type 5 acreages to qualify as type
species include Douglas, dwarf, and rubber rabbitbrushes, snowberry, bitterbrush, and greasewood
Browse plants present but not abundant nor conspicuous
enough for type designation are ephedra and gray horsebrush.
o

Similar to sagebrush/pinyon-juniper
mixtures which have sagebrush aspects
(see section on Sagebrush type), there are extensive areas in Unit 22 where
P-J, in low abundance, also mixes with vegetation having browse aspects
(20 types - 32,676 Ao)o
Both 5-PJ and 4-PJ associations have been mentioned
previously herein as transition types
I think that use of the term
"transition" is particularly appropriate for 4-PJ areas which seem to be
intermediate stages to 9-PJ finally.
On the other hand, since Oosting
(1956:296,303-304)
says that both pinyon-juniper and oak-mountainmahogany
(browse) types are foothills zone climax vegetation, there is room for conjecture on the ecological situation represented by high-density pinyon-juniper
o

�- 49 in browse.
To further speculate, I believe that 5-PJ types are essentially
very stable communities and probably require catastrophic influences to
effect changes, either towards mature woodland or mature shrub1and vegetation.
A wide variety of forbs are present in browse associations.
However, none
was prominent enough to warrant type species designation.
Commonly recorded
broad-leaf herbs "hit" in transecting work include mountain thermopsis
(Thermopsis montana), astragalus (Astragalus sPp.), fringed sagebrush
(Artemisia frigida), clovers (Trifolium spp.), eriogonums (Eriogonum sPP.),
goldenweeds, lupines, and phloxes.
Last, but certainly worthy of note and very important, are grasses that were
given type species recognition under the browse.
Need1eandthread
and b1uestem wheatgrass are cited for very substantial acreages.
Many other grasses
are found with browse.
They, along with those just mentioned, rated low
priority for citation whenever shrubs could logically be recorded instead.
Sedges

(Carex spp.) are cited for one type of minor acreage.

Cultivated and hay 1and.--On the basis of containing about 10,000 acres, the
category Cultivated and Hay Land ranks as the fourth most extensive cover
type of Unit 22 winter range (Table 1).
Regardless of extent of acreages,
this land category has been left for last in previous like reports on other
units; by so doing, hay and cultivated ground were inferred to have little
direct bearing on welfare of wintering deer.
In Unit 22, just as in many
other western slope locales, hay meadows furnish considerable first green
forage for about a month each spring.
Thus, for helping alleviate nutritional deficiencies at a most critical time for deer, hay and cultivated
areas deserve and are herein given more recognition.
More detailed definition is due the two-part title of this cover category.
The "Cultivated" portion is equivalent to annual and perennial crops of small
grains, grass for seed, corn, and so forth; these crops make up a small
percentage of the total acreage of the category.
"Hay Land", on the other hand, comprises the great bulk of the category and
includes both improved and unimproved kinds of ground. A partial listing
of plants found on improved meadows would include. alfalfa (Medicago sativa),
smooth brome (~. inermis), timothy (Ph1eum pratense), and orchardgrass
(Dactylis glomerata).
Unimproved meadows presently support mostly such
plants as b1uestem wheatgrass, cordgrass (Spartina sp.), foxtail barley
(Hordeum jubatum), sedges, and rushes (Juncus spp.), whether or not the
land had ever been plowed and seeded.
Saltbush type.--Type 13, the Saltbush Type, provides about 3,900 acres, or
one percent, of the total winter range in Unit 22 (Table 1).
In area and
source of forage, this cover type is unimportant.
It is valuable, nevertheless, by serving to break up expanses of pinyon-juniper and providing variety
of forage.
Contrary to being on bottom1ands, where first thought might place
them, Saltbush Types in Unit 22 are mostly upland sites. The sites are
underlain by the Green River Formation.
Of the three separate types present,
one is near the mouth of Piceance Creek, one is just east of the mouth of Yellow
Creek, and one encompasses most of Boise Creek Basip. All types are on northerly

�- 50 exposures that are near and break directly
White River.

or almost directly

into the

It is consistent that the highly alkaline soils derived from calcareous
portions of the Green River Formation should favor the alkaline-tolerant
saltbushes.
Of browse compositions determined from three condition transects in the Saltbush Types, shadsca1e saltbush is the most abundant species
on two. On the third transect, shadscale is a close second to big sagebrush.
Douglas rabbitbrush and bluestem wheatgrass also are listed as type species
in Type 13 associations.
Shadsca1e saltbush is not preferred deer forage. That deficiency is perhaps
remedied by the already-mentioned
presence of big sagebrush and Douglas
rabbitbrush, and to a lesser extent by minor amounts of serviceberry, mountainmahogany, and rubber rabbitbrush.
Other less desirable shrubs that occur
in minor quantities in Type 13 acreages are Gardner saltbush ~. gardneri),
black greasewood, common winterfat, snowberry, and gray and cottonthorn horsebrushes.
Most common grasses and forbs include bottlebrush squirreltail,
Indian ricegrass, saltgrass (Distichlis spo)~ cheatgrass brome, clovers,
onion (Allium sp.), prick1ypear, and phloxes.
Greasewood typeo--S1ightly
less than the total acreage of the Saltbush Type
is the Greasewood Type (Type 14); its 3,716 acres are also about equal to
one percent of the winter range area.
Similarity between Types 13 and 14 essentially ends with like total acreages,
however, since Type 14 is found exclusively on bottomland areas compared with
the generally upland locations of saltbush cover. Also, differences are
evident in the functions that each vegetation type serves.
Black greasewood,
a very poor forage species for game, mainly provides cover.
In contrast,
shorter-statured saltbush communities function mainly to provide a variety
of forage.
Big sagebrush is associated with greasewood in mixtures and typically exhibits
rank growth form like it does composing its own types (see section on Sagebrush type).
Therefore, except where rabbitbrushes are fairly abundant in
type compositions and when grasses green up annually in spring, Type 14
acreages apparently do not supply much deer forage.
Pinyon pine and junipers appear as type species in one type of 121 acres,
indicating a possible tendency of these conifers to invade and/or replace
greasewood in a manner similar to that occurring with sagebrush and browse
types.
From information on one transect in one type, subdominant grasses occurring
with greasewood include saltgrass, cheatgrass brome, and bottlebrush squirretail; forbs listed from the same sources are phloxes, mustard (Cruciferae),
and summercypress (Kochia sp.). Cheatgrass brome, in its early stages of
growth, probably contributes the most forage for deer of all plants present.
Grassland type.--Grassland Type (Type 1) vegetation is perhaps worthy of
note principally from the standpoint of ecological interest; because, based
solely on numbers of acres, the type can not be considered as being critical
to the upkeep of very many deer. In extent, Type 1 comprises only about
2,900 acres of Unit 22 winter range.

�- 51 Except for possibly 125 acres in two types (l-Agsm-Artr, l-Kocr-Agsm) of
unknown ecological origin, grassland vegetation in Piceance Basin resulted
from man-caused sagebrush or pinyon-juniper rehabilitation.
Despite some
reseeding of exotics, native grasses are still the most abundant herbaceous
cover in Type 1. The natives are led by bluestem wheatgrass and Indian
ricegrass.
Fairway crested wheatgrass, prairie junegrass, and cheatgrass
brome are other herbaceous plants recognized by being listed as type species.
Trees and shrubs cited as type species in Grassland Types are pinyon pine,
junipers, mountainmahogany,
big sagebrush, and Douglas rabbitbrush.
Thus,
grassland areas can not automatically be disregarded in deer range evaluations,
although abundance of woody forage plants is invariably poor whenever Type 1
is used to deSignate an area of vegetation.
Other types.--The Conifer Type (Type 6) and Annual Type (Type 18) each comprise less than 1,000 acres, or one-half of one percent, of the total winter
range.
Douglas fir and junipers are dominant overs tory tree species in Type 6
acreages which occur on high-elevation north slope sites in the extreme
northeastern and southeastern corners of the unit (Kendall, Davis, and Coyote
Gulches).
Subordinate browse species include serviceberry, Gambel oak, and
snowberry.
Also present but not used as type species is common chokecherry.
A great variety of herbaceous plants are present, too, as ground cover.
The Annual Type is composed of cheatgrass brome areas that are found only
west of Piceance Creek. One type is noted as a burn; the other adjoins and
could also have had the same history.
Douglas rabbitbrush and serviceberry
are abundant enough to rate type species mention with cheatgrass.
Browse
range condition transects revealed the presence of additional following
species: common winterfat, snowberry, mountainmahogany,
big sagebrush, dwarf
rabbitbrush, broom snakeweed, Indian ricegrass, needleandthread,
bottlebrush
squirreltail, and prairie junegrass.

BROWSE RANGE CONDITION
When this inventory started, it was hoped that at least one browse range
condition transect would be established and read in every browse containing
type. It was soon apparent that such a seemingly minor goal could not be
met with means available.
Consequently, in order to fulfill the minimum
need of a type map of the winter range in Unit 22, tighter priorities and
reduced transecting were effected.
Almost completely ignored by sampling
procedures, then, were types having browse canopy coverages of about five
percent or less or rank bottomland shrub growth; sparse understory pinyonjuniper and sagebrush stringer bottoms fit those latter categories where
undersampling occurred.
A total of 309 transects were finally read, however,
and copies of field data forms and summaries are filed and accompany the
binder file folders which hold this report.
In addition to browse range condition transects, 197 "non-transected"
write-ups were prepared for most types not having condition transects

type
installed

�- 52 in them. These forms served two purposes:
First, through listing of
plant species present and cursory age-form descriptions of browse, a
record of vegetation was made for the type.
Secondly, a pellet group
count was done and recorded on each write-up to furnish an index to
game and/or livestock use of the type, This latter information became
particularly useful by helping determine placement of the winter range
line. Copies of completed "non-transected"
type forms are also retained
in the binder file folders.
Due to sampling inadequacies, it is not feasible to statistically evaluate
condition transect data nor will it be possible in future years to reliably
determine trend of vegetation of the whole or subunits such as key areas.
Changes in transected types might be detectable, but even that will remain
to be verified through actual rereading.
Despite foregoing uncertainties
of inventory techniques, the following paragraphs will present attempts
to draw general and broad-based conclusions about winter range vegetation.
Composition,--A
composite picture of Unit 22 browse range condition transect
ratings is presented in Table 5. From the table, an overall rating of
''Medium +" is given for browse composition which is strongly favorable for
deer.
Deer have been proven to do best when they obtain sustenance from
as many good plant species as possible.
It is conceded that the ''Medium +"
rating value of 2.10 is far from maximum attainable, but many "High" ratings
were necessary to provide that average.

Table 5. Summary of ratings for browse and soil, 309 browse range condition
transects, Game Unit 22 (Piceance) big game winter range, 1965-67.
Item

Low

Rating
Medium
High

Total

Browse Composition

Overall or
Average Ra t Lngw

2.10 (Medium+)

No. of Transects

79

III

119

309

% of Total

26

36

38

100

Browse Density

1.46 (Low+)

No. of Transects

178

120

11

309

% of Total

57

39

4

100

Browse Vigor

1.38 (Low+)

No. of Transects

220

61

28

309

% of Total

71

20

9

100

Soil Stability

1. 87 (Medium-)

No. of Transects

64

221

24

309

% of Total

21

71

8

100

* Computed on basis of Low=l, Medium=2, and High=3.

�- 53 -

It is difficult to determine an accurate ranking of browse species abundance for Unit 22 winter range in its entirety.
Nevertheless, some indications of comparative abundance can be deducted from type acreage
summaries, be~ause type species citations result and depend upon abundance
after the initial aspect and timber designation requirements have been met.
Data on Table 6 also provide gross indexes to abundance of shrubby species.

Table 6. Frequency of shrubs listed as key species from vigor ratings on
309 browse range condition transects, by key and total area, Game Unit 22
(Piceance), 1965-67.

Key Area

Frequency Occurrence of Shrub Species Listed as Key 17,17
Artr AME Cemo Putr Chvd Quga Chde At co Arno Chna GUT Eula PRU

Rattlesnake

10

20

19

2

3

Greasewood

23

15

9

2

4

Thirteenmile

45

39

16

1

2

8

Segar Mountain

24

45

26

4

6

6

Hammond-Barcus

24

18

31

2

1

Bracus-Ryan

29

2

27

12

Ryan-Story

16

3

6

3

Totals

171

142

134

26

1

1

3

1

1

1

1

1

1

1

2
16

14

3

3

2

1

1

1/
- See Chapter 80, Region 2 Range Environmental Analysis
Service, or Denney (1962), for plant symbol meanings.

Handbook,

U. S. Forest

2/
- Vigor computations are made and scored for all species designated as being
"key" on a transect.
More than one species are very often listed for each
transect, thus allowing for the discrepancy between total frequencies and
total transects.

Thus, by referring to vegetation type summaries as supportive evidences, big
sagebrush justifies being ranked the most abundant and top-volume forage producer in Unit 220 Serviceberry and mountainmahogany
apparently are tossups
for second ranking.
The former is cited more often (62 types) but in lower
total acreage (101,211 A.) than the latter (43 types - 127,067 A.). Also
complicating interpretation here is the fact that mountainmahogany
occurs in

�- 54 -

low densities in large type tracts of pinyon-juniper, and both species
frequently are listed together in type designations.
Suffice to say that
having three high-value shrubs abound on the winter range has to be a key
factor in the long standing success of the associated deer herd.
There are three additional shrubs that further enhance Unit 22 deer winter
forage.
Based on type acreages in approximate order, fourth ranked bitterbrush in 10 types totalling 27,682 acres is most common west of Piceance
Creek, but occurs sporadically in fair abundance elsewhere.
Douglas
rabbitbrush is fifth ranked from presence in 20 types of 24,526 acres.
It
actually could be more abundant than bitterbrush, since it is distributed
everywhere, frequently below abundances that warrant giving it type species
designations (e. g. sagebrush types).
Perhaps a solid sixth in browse
abundance is. oakbrush (20 types, - 23,044 A.).
It is found almost
exclusively east of Piceance Creek at higher elevations.
Abundance of oakbrush can be discounted on the basis of volume-availability
because crowns
frequently are out-of-reach and stands are impenetrably dense.
Except by selectively eradicating pinyon-juniper to promote browse, altering
vegetative composition to benefit deer, is, in reality, subject to problems.
Care should be exercised that more undesirable compositions do not result
through well-meaning action.
There are some indications that nearly pure
stands of sagebrush and rabbitbrush occur under severe disturbance, for
example.
Density.--As shown in Table 5, densities of browse tend to be deficient
overall.
Additional key area average densities are given in Table 7; they
indicate that the two easternmost high elevation key winter range areas of
Segar Mountain and Thirteenmile have the greatest browse densities.
Also,
it can be noted that oakbrush is involved in the upper values of density
ranges for those same key areas.
Big sagebrush types provide upper values
of density ranges for the other two key areas east of Piceance Creek,
namely Rattlesnake and Greasewood.

Table 7. Browse densities
Game Unit 22, 1965-67.
Key Area

by key area, 309 browse range condition

No. Trans.

Densities
Average

Rattlesnake

32

13.8

Greasewood

32

15 6

56

22 3

Thirteenmile

0

0

Segar Mountain

54

Hammond-Barcus

60

12.0

Barcus-Ryan

55

6.5

Ryan-Story

20

7.4

22.2

as Percent

transects,
Crown Cover
Range

2 - 27
(9-PJ)
(4-Artr)
4 - 43
(9·-PJ)
(4-Artr)
6 - 55
(9-PJ) . (5-Quga)
6 - 47.
(9-PJ)
(5-SYM-QUE)
2 - 39
(5-AME)
(9-PJ)
o - 19
(9-PJ)
(9-PJ)
1 - 16
(9-PJ) - (4-Artr)

�- 55 Lowest browse densities are consistently associated with pinyon-juniper
vegetation on all seven key areas (Table 7). See MANAGEMENT RECOMMENDATIONS
section at end of report for proposed specific remedies to improve shrub
cover.
Vigor.--Of 309 browse range condition transects, 220 were rated low for
vigor (Table 5). Observations apply to the 3-year span of 1965-67, coincidentally with deer populations at low numbers.
With fewer deer, higher
plant vigor might have been expected but was not immediately possible.
Hedging (form) characteristics depend upon recent past use (within three
years prior to observations exclusive of current growth), so form classification actually does not reflect current grazing intensity and apparently
did not in this case, as well (Table 5).
Besides plant hedging or form characteristics, vigor ratings also involve
age criteria.
An integral of age is amount of decadence.
The rate that
plants mask dead parts (decadence) in recovering vigor through production
of new growth is variable, being dependent upon factors affecting growth
and attrition (e.g. effective precipition and grazing).
Moreover, as
slowly as shrubs ordinarily grow, several years may be required to increase
"young" components enough to effect changes in ratings.
Thus, age characteristics of browse populations are usually not as quickly demonstrative
of plant recovery as are form or hedging characteristics.
Soil stability. --Overa 11, soil stability has a ''Medium minus" rating from
about a median numerical value at 1.87 (Table 5). Like the item browse
composition, this component of range condition is not easily improved upon
by man.
It is, however, susceptible to rapid deterioration by many means.
Lowest parts of the basin lack vegetation; they are thus particularly
vulnerable to forces of heavy cloudbursts which also characterize the lower
zone. Conservative stocking of livestock and game must be observed if soil
losses are to be minimized.

BIG GAME DISTRIBUTION
Pellet group counts were done in conjunction with browse range condition
transect data collection and non-transected type records preparation.
All
groups were classified and recorded by spring-summer or fall-winter category
regardless of length of time in place. That resulted in cumulative indexes
of use rather than current stocking rate estimates such as are determined
in annual kinds of surveys.
Transect plots were mostly .Ol-acre circular.
Some few transects had 100-square-ft. circular plots employed.
Table 8 summarizes results of pellet group counts by key area and season of
use by deer.
It shows that winter range of the northeastern corner of Unit
22, as represented by Rattlesnake and Segar Mountain Key Areas, receive the
heaviest use and support the greatest numbers of wintering deer. The
Greasewood and Thirteenmile Key Areas follow Rattlesnake and Segar Mountain
fairly close in degree of winter deer occupation.
Winter use of HammondBarcus, Barcus-Ryan, and Ryan-Story Key Areas drops down to light stocking.

�- 56 -

It is appropriate that the greatest concentration of winter deer use falls
on the Rattlesnake Key Area which also contains a substantial acreage of
Game, Fish and Parks patented lands. Therefore, range improvement planning
east of Piceance Creek should include Rattlesnake Key Area in top priority.

Table '8. Indexes of deer stocking
key area, Game Unit 22 (Piceance),

Key Area

Number of
Transects

*

from pellet group count transects,
1965-67.

by

Mean Deer Pellet Groups/Acre
Spring-Summer
Fall-Winter

Rattlesnake

60

7

1300

Greasewood

72

5

980

Thirteenmile

78

101

888

Segar Mountain

73

84

1022

Hammond-Barcus

72

63

740

Barcus-Ryan

84

40

675

Ryan-Story

32

365

402

* Pellet group counts made both on browse range condition transects and "nontransected" form write-ups.
Does not include transect data where sheep use
was evident or suspected, because of inaccuracies that would be produced by
their presence.

The Ryan-Story Key Area apparently receives almost as much use by deer in
spring and summer as it does in fall and winter, ranking it first in springsummer use of all key areas.
High year-round ranges of Thirteenmile and Segar
Mountain Key Areas receive what might be considered low-moderate spring-summer
occupation that very likely has little influence on browse production and
health.
Elk occupation of Unit 22 winter range was almost nil as shown by pellet group
counts.
Heaviest use was recorded on Hammond-Barcus and Segar Mountain Key
Areas; fall-winter indexes there averaged less than three pellet groups per
acre and on other key areas zero-plus.
Impact of present elk populations thus
has to be comparatively unimportant on habitat.

MANAGEMENT

RECOMMEN~TIONS

Recommendations resulting from this inventory fall into either one of the
broad categories of unit boundary changes or vegetation manipulation activities.
The former have been discussed previously under the heading DESCRIPTION OF UNIT;

�- 57 the latter are given as follows:

I. Best information available leads this writer to believe that
pinyon-juniper manipulation is the most efficient and economical
means for increasing shrubby vegetation in Unit 22. In fact,
some chaining has already been done near lower Yellow Creek to
improve that area. Thus, it is recommended that chaining, cabling,
and/or dozing be continued but under general restrictions described
in following paragraphs.
First, concurring with information presented by Minnich (1969:35-36),
it is suggested that continuous chained areas not be too large.
Blocks or areas of undisturbed woodland should be interspersed with
chained areas in order that any continuous chaining does not exceed
one-quarter mile in width.
Secondly, more reliable and quicker improvement in browse cover is
obtainable if pinyon-juniper is treated where residual browse exists.
Understory shrub cover percentages vary considerably, thus making
each pinyon-juniper type subject to individual evaluation.
However,
there is enough P-J containing lower (five to 10 percent) amounts
of browse in desirable composition and on suitable terrain to furnish
almost unlimited acreage for rehabilitation on Unit 22 winter range.
2.

Where sagebrush forms characteristically
rank growth in bottomland
stands, it is recommended that type changes be effected by chaining,
plowing, or pipe harrowing and reseeding.
Mixtures of adapted
perennial grasses and legumes should be broadcast before (or between)
treatment(s) in efforts to minimize opportunities for cheatgrass
brome to take over the sites.
Where other cover is too distant, sparse, or absent for effective
protection, treatment should be done in blocks with treated blocks
alternating with untreated areas.
Since standards are not available
on optimum sizes and spacings of treated vs. untreated areas, use of
good judgment by local administrators is the best advice that presently can be given.

3.

There are mature oakbrush stands in the extreme northeast corner of
Unit 22 that could be improved for forage production.
Early spring
dozing of the tall, out-of-reach plants is suggested to open the
stands and stimulate sprouting.
Browse range condition transect
records and the 2-inch scale vegetation type map can be consulted
to assist with initial planning activities.
It might be rather
superfluous to add that, whatever range improvement projects are
devised, careful and detailed plans should be made prior to
implementation.

�- 58 -

REFERENCES

CITED

Anderson, A. E., W. A. Snyder, and G. W. Brown.
1965. Stomach content
analyses related to condition in mule deer, Guadalupe Mountains,
New Mexico.
J. Wi1d1. Mgmt.
29(2):352-366.
Beetle, A. A. 1960. A study of sagebrush$ The Section Tridentatae
Artemisia.
Univ. Wyoming Agr. Exp. Stag Bull. 368. 83 pp.
Carhart, A. H. 1941.
food requirements
(processed).

of

Deer-elk survey, Colorado, Vol. 5., Part I Deer
in Colorado.
Colo. Game and Fish Comm.
28 pp.

1943. The Piceance-White River deer herd.
Supp1. Rep., Colo. Game and Fish Comm. 20 pp.
Colorado State Planning Division.
1961.
of the State of Colorado.
916 pp.

Colorado

Deer-elk survey,
(processed).
1959-61, Yearbook

Denney, R. N. 1962. Browse transect analysis and application, pp. 5196. In Game Res. Rep., April.
Colo. Dept. Game, Fish and Parks.
101 pp. (processed).
Hill, R. R. 1956. Forage, food habits, and range management of mule
deer, pp. 393-414.
In W. p. Taylor (ed.). The deer of North
America.
The Stackpole Co., Harrisburg, and the Wildlife Mgmt.
Inst., Washington, D. C. 668 pp.
Jacobsen, R. 1969. Intensive inventory and analysis, C1 WBA-T1 Yellow
Creek for mule deer (Odocoi1eus hemionus).
BLM Craig Dist. Off.,
July.
25 pp.
(processed).
Minnich, D. W. 1969. Vegetative response and pattern of deer use
following chaining of pinon and juniper forest. J. Range Mgmt.
Abstracts of Papers, 22nd Ann. Mt. Amer. Soc. Range Mgmt.
59 pp.
Oosting, H. J. 1956. The study of plant communities.
2nd ed. W. H.
Freeman and Co., San Francisco and London.
440 pp.
Smith, A. D. 1959. Adequacy
wintering of mule deer.
Smith, J. G. 1952.
16(2):148-155.

of some important browse species in overJ. Range Mgmt.
12(1):8-13.

Food habits of mule deer in Utah.

J. Wi1d1. Mgmt.

U. S. Department of Agriculture.
1966. Water and related land resources,
White River Basin in Colorado.
Coop. Study Rep. of Colo. Water Conserve
Board and D.S.D.A. Econ. Res. Service, Forest Service, and Soil Conserve
Service.
November.
December.
92 pp. (Loose leaf).
Yeager, L. E. 1960. Factors affecting the quality and management of big
game winter range in Colorado.
Colo. Game and Fish Dept., Current
Rep. No. 30. January.
29 pp. (processed).
March, 1970
Fort Collins

�July,

- 59 -

JOB PROGRESS

REPORT

S ta te of

...::c:,;:O:,;:L::,.::O:;,.RA::;:;::D:..::O:...------

Project No.

W-lOl-R-12

Work

Plan No.

Game Range

Investigations

Job No.

4

1970

la

Job Title __~I~n~v~e~n~t~o~r~y~o~f~R~a~n~g~e~M~a~n~i~p~u~l~a~t~i~o~n~P~r~o~J~·e~c~t~s~i~n~C~o~l~o~r
Period Covered:

April

1, 1969 through March

Personnel:

Roland

C. Kufeld

31, 1970

ABSTRACT

An inventory was made of all range type-conversion
projects completed
during 1966 through 1969, in Colorado on lands administered by the U.S.
Forest Service, Bureau of Land Management and Bureau of Indian Affairs.
Acreages treated were:
Forest Service29,283 acres, Bureau of Land
Management - 74,517 acres, Bureau of Indian Affairs - 28, 287 acres,
Total - 132,087 acres.
Of this total acreage treated 70,547 acres were
seeded.
This report contains analyses of range manipulation project inventory
data by kind of treatment within individual National Forests, B.L.M.
Districts, Indian Reservations,
and Game, Fish and Parks Regions.
Data
are included on kinds of equipment and chemicals used for treating rangelands, plant species used for reseeding, and environmental
characteristics
of areas where range type-conversions
have been located.
Data on livestock and wildlife use prior to treatment are also presented.

��- 61 -

INVENTORY

OF RANGE MANIPULATION
Roland

PROJECTS

IN COLORADO

C. Kufe1d

Po So OBJECTIVE
To collect information which describes the location, extent, environmental
conditions, land use practices, type of treatment and effects of treatment
on the range, livestock, and wildlife, for all completed range manipulation
projects in Colorado that are located within the ranges of deer, elk and
sage grouse, and develop a processing system for information which describes
proposed range manipulation projects.

SEGMENT OBJECTIVES
1.

To inventory range type conversion
during 1966, 1967, 1968 and 1969.

2.

To compile,

3.

To develop a processing and filing system for information
posed range type-conversion
projects in Colorado.

codify, process,

projects

and analyze

in Colorado

inventory

completed

data.
about pro-

METHODS AND MATERIALS
An inventory of all range type-conversion
projects completed through 1965,
in Colorado, on lands administered by the U.S. Forest Service, Bureau of
Land Management and Bureau of Indian Affairs has been completed, and the
results reported by Kufeld (1968). This report concerns type-conversion
projects completed between January 1, 1966, and December 31, 1969. The
tables contained herein are designed in the same manner as those in the
1968 report, so a reader, if he so desires, can easily combine the data.
These table formats will also be used in future reports.
Two data collection methods were used. Most of the data were collected by
contacting offices of the U.So Forest Service, Bureau of Land Management
and Bureau of Indian Affairs, and recording pertinent range type-conversion
project information from records on file in those offices.
Office contact
inventory procedures have been described by Kufe1d (1968).
Some of the
data were provided by Regional Game, Fish and Parks Division biologists
using procedures outlined in "A System for Inventory, Evaluation and
Exchange of Information on Range Type Conversion Projects" (Kufe1d, 1968).
Inventory data were transferred from original data sheets to Mark Page
Reader Forms, and then to I.BoMo cards.
Data were analyzed by the Colorado
State University Computer Center.
Unfortunately,
records pertaining to
individual range type-conversion
projects were sometimes sketchy, and for
some projects it was not possible to obtain all of the inventory data
needed to complete the inventory form (Fig. 1). This report summarizes

�- 62 -

all of the inventory data available, and it must be pOinted out that
differences which occur in acreage grand totals among several tables
contained in the report are due to lack of data in some categories for
some individual projects.
For example, the total of acres seeded by
each seeding method (Table 25) is less than the total acres seeded
by the three agencies (Table 4), because information on the seeding
method was lacking for some individual projects.
Data in Tables 1
through 23 and 31 through 36 represent all of the range type-conversion projects inventoried, because the information in the categories
listed in those tables was available for all projects.
Data in Tables
24 through 30 represent only a portion of the projects inventoried,
because the data could not be obtained for some projects.
Color coded maps showing the location and type of treatment for each
type-conversion
project inventoried, and xerox copies of each inventory form were made and sent to the Game, Fish and Parks Regional
Manager in whose region the projects were located.

RESULTS AND DISCUSSION

Location

and Extent

of Completed

Range Type-Conversion

Projects

Acreages treated during 1966 through 1969, inclusive, on lands administered by the U.S. Forest Service, Bureau of Land Management, and Bureau
of Indian Affairs, and on all of the lands of these three agencies combined are shown by vegetative type and kind of treatment in Tables 1
through 4. A total of 132,087 acres were treated of which 57 percent
(74,517 acres) were treated by the Bureau of Land Management, 22 percent
(29,283 acres) by the Forest Service and 21 percent (28,287 acres) has
been treated on Indian Reservations
Including those projects completed
prior to 1966 (Kufeld, 1968), a total of 650,195 acres have been treated,
of which the Bureau of Land Management completed 51 percent (333,263 acres),
Forest Service 35 percent (225,777 acres), and Bureau of Indian Affairs
14 percent (91,155 acres).
Fifty-three percent (70,547 acres) treated
during 1966 through 1969, were seeded.
o

The number of acres treated per year has declined somewhat since 1965.
An average of 33,022 acres were treated annually during 1966, through
1969, compared with 55,340 acres treated per year from 1961 through 1965.
From the mid 1930's through 1965, most range type-conversion
efforts
were directed toward sagebrush control.
Pinyon-juniper control ranked
second in importance from 1956 through 1965. Between 1961 and 1965, inclusive, 47 percent (131,755 acres) of the acreage treated was to control
sagebrush, and 35 percent (97,681 acres) was for control of pinyon and
juniper.
During the 1966 through 1969 period, however, 50 percent
(66,484 acres) of the acreage treated was to control pinyon and juniper,
and 43 percent (56,214 acres) was to control sagebrush.
Most of the increased pinyon-juniper
control during 1966 through 1969, was attributed to
an extensive pinyon-juniper
control program on the Ute Mountain Indian
Reservation (Table 17). Sagebrush control still received primary emphasis
on Forest Service and Bureau of Land Management lands (Tables 1 and 2).

�- 63 -

Acreages treated by vegetative type and kind of treatment are shown for
individual National Forests, Bureau of Land Management Districts and
Indian Reservations in Tables 5 through 17. Total acreages treated by
these agencies are compared directly in Table 18, and the information
is shown by location within Colorado Division of Game, Fish and Parks
Regions in Tables 19 through 21.
The Uncompahgre National Forest ranked first among the National Forests
in the number of acres treated during 1966 through 1969. They treated
8,517 acres.
The Gunnison was second with 8,233 acres, and the White
River third with 5,326 acres. Among the Bureau of Land Management
Districts the Montrose District was first with 28,676 acres, Craig
second with 22,134 acres, and Grand Junction third with 19,969.
No
range type-conversion work was done during 1966 through 1969, on the
Arapahoe, Pike, Roosevelt and San Isabel National Forests, or the
Glenwood Springs B.L.M. District.
During the 1966 through 1969 period
the Southern Ute and Ute Mountain Indian Reservations treated 5,301 and
22,986 acres respectively.
During 1966 through 1969, 65 percent (85,612 acres) of the total acres
treated were located in the Southwest Game, Fish and Parks Region, 35
percent (45,839 acres) were in the Northwest Region, and less than 1
percent (636 acres) was in the Southeast Region.
No type-conversions
were done in the Northeast Region.

Range Type-Conversion
Treatmel'lts,sEqllipment,
Chemical Herbicides and Plant Species Used for Reseeding
The various kinds of treatments such as spraying, chaining, plowing, etc.
have been listed in Table 4. Most of the sagebrush control during 1966
through 1969, (47,338 acres or 84 percent) was done by spraying.
Six
percent (3,525 acres) was done by chaining, 6 percent (3,503 acres) by
plowing, and 3 percent (1,728 acres) involved the use of a brush cutter.
In the pinyon-juniper type 91 percent (60,684 acres) of the control work
done during 1966 through 1969, was by chaining.
Seven percent (4,600
acres) of the pinyon-juniper area treated was burned (Table 4), however,
this involved burning pinyon-juniper slash that had been previously chained
rather than burning standing vegetation.
Ninety-nine percent of the spraying in Colorado during 1966 through
has been done aerially, and has involved the use of 2,4-D herbicide
22).

1969,
(Table

Acreages treated by anchor chaining, cabling, railing and bulldozing are
shown in Table 23. More than 99 percent (64,349 acres) of this work involved the use of an anchor chain.
Less than one percent involved bulldozing only, and there were no cabling or railing projects during the
1966 through 1969, period.

�- 64 -

Crested wheatgrass (Agropyron cristatum) continues to be the most important grass species used for reseeding.
It was used on 84 percent (58,304
acres) of the total acres seeded (Table 24). Other important grasses were
intermediate wheatgrass (Agropyron intermedium) used on 45 percent (31,865
acres) of the seeded acres, Russian wi1drye (E1ymus junceus) (42 percent
or 29,866 acres), smooth brome (Bromus inermis), (41 percent or 28,787
acres), and other wheatgrasses
(Agropyron spp.)(36 percent or 25,632 acres).
Two forb species were used for reseeding.
Sweet Ciover (Me1itotus sp.)
was seeded on 20 percent (13,872 acres) of the seeded acres, and alfalfa
was used to reseed 15 percent or 10,640 acres.
No alfalfa had been used
to reseed Federally administered range lands in Colorado prior to 1966
(Kufe1d, 1968).
A substantial increase has occurred in the number of acres seeded with
browse species.
Prior to 1966, browse species were seeded on only three
percent of the total seeded area (Kufe1d, 1968). During the 1966 through
1969, period, however, 20 percent (13,865 acres) of the total seeded area
was seeded with fourwing saltbush (Atrip1ex canescens); bitterbrush (Purshia
tridentata), was used on 15 percent (10,866 acres); and mountain mahogany
(Cercocarpus montanus) was used on 6 percent (4,574 acres).
C1iffrose
(Cowania mexicana), was used for the first time in Colorado and was seeded
on 990 acres.
Plants were seeded in various combinations of species so
numerous that it is impractical to list all of the combinations.
Seeding methods varied depending on the vegetative type and kind of seed.
Comparatively
little seeding was done in the sagebrush type, however, the
two primary methods of seeding in the sagebrush type were drilling and
aerial broadcasting
(Table 25). In the pinyon-juniper
type aerial broadcasting was used almost exclusively for seeding grasses and forbs. Browse
seed was usually applied with a browse seeder or seed dribbler, even on
projects where browse seeding was accompanied by aerial broadcasting of
grasses and forbs.
Most reseeding (88.5 percent)
acre (Table 26).

was done at a rate of 5 to 10 pounds per

Environmental Characteristics
of Areas
Where Range Type-Conversions
Have Been Located
Pre-treatment plant densities on sagebrush and pinyon-juniper projects for
which plant density data were available are shown in Table 27. Pre-treatment densities on sagebrush control areas ranged as high as 80 percent,
although most of the work was done where densities were less than 40 percent. All of the pinyon-juniper
control work during 1966 through 1969,
was done in areas where plant density was under 40 percent.
Shrub composition before treatment, on projects where composition data were
available, ranged from 21 to 70 percent on sagebrush control areas and 51
to 80 percent on pinyon-juniper control areas (Table 28).

�- 65 During the 1966 through 1969 period, range conditions for livestock prior
to treatment (for those projects where pre-treatment
livestock range condition data were available) on sagebrush and pinyon-juniper areas was most
commonly considered "fair to good" or "goodlt, and none was considered "poor".
This is in contrast to those projects completed before 1966, (Kufeld, 1968)
when most sagebrush and pinyon-juniper control areas were felt to be in
"poor" condition for livestock prior to treatment (Table 29). Data on condition of these ranges for wildlife were not available.
Erosion
slightlt
30).

condition before treatment
to "moderate" on sagebrush

most commonly ranged from "none or
and pinyon-juniper control areas (Table

Pre-Treatment Livestock and Wildlife
Use on Range Type-Conversion Project Areas
Ninety-eight p~rcent (129,986 acres) of the total acreage of range treated
in Colorado by the three agencies during 1966 through 1969, was located
within grazing allotments.
Eighty-nine percent (115,661 acres) was grazed
by cattle and horses, and 11 percent (14,325 acres) was used by sheep and
goats (Table 31).
Ninety-six percent (126,738 acres) of the acreage treated during 1966
through 1969, was within the range of mule deer. Most of the treated deer
range (111,119 acres or 88 percent) was used by deer during the fall-winterspring period or on a yearlong basis.
The remaining 12 percent (15,619
acres) was summer range (Table 32). Twenty-nine percent (37,003 acres of
the treated deer range was considered heavily used by deer before treatment
(Table 33)0
A total of 35,287 acres of elk range were treated during 1966 through 1969.
Seventy-nine percent (27,881 acres) was used by elk during the fall-winterspring period or on a yearlong basis, and the remaining 21 percent (7,406
acres) was summer range (Table 34). Only 2 percent (761 acres) of the
treated elk range was considered heavily used before treatment (Table 35).
A total of 29,178 acres of sage grouse range were treated of which 62 percent (17,192 acres) was heavily used sage grouse range (Table 36)0
Prior to 1966, most of the range type-conversions
were designed primarily
to benefit livestock.
Modification of range vegetation to suit the specific
needs of livestock sometimes has an adverse effect on species of wildlife
whose requirements are different from those of livestock.
During the 1966
through 1969, period, however, there has been a substantial increase on some
National Forests and Bureau of Land Management Districts in efforts to improve
the range for wildlife as well as livestock.
This is particularly evident
among type-conversion
projects completed by the Montrose Bureau of Land
Management District where serious consideration has been given to maintaining
or improving wildlife habitat on all of the 1966 through 1969, projects that
were located in important wildlife areas.
Their sagebrush spray projects
were designed to spray in 200 yard strips'with 100 yard leave strips, and
no spraying was done within 200 yards of streams or ponds.
Their pinyonjuniper chainings were designed to leave strips or patches of standing
pinyon and juniper of at least 100 yards in width at 1/5 to 1/4 mile intervals,

�- 66 -

and along canyon rimso Chained areas were reseeded with browse as well
as grass specieso
Colorado Division of Game, Fish and Parks has participated financially toward improving wildlife habitat on many of the
Montrose BLM type-conversion projects, as well as projects on other BLM
Districts and National Forests in Colorado.
Financial participation by
the Division was usually in the form of contributing browse seedo
Closer interagency coordination in planning range-type conversion projects has been a significant factor during the 1966 through 1969, period
in maintaining and improving wildlife habitat in relation to range typeconversion worko
Proposed range type-conversions are reviewed and discussed by land management agency and Game, Fish and Parks Division personnel before the projects are implemented.
Projects which might have
a detrimental effect on wildlife are usually modified or deleted.

LITERATURE
Harrington, H. D. 1954.
Denver.
666 po

Manual

CITED

of the plants of Coloradoo

Sage Books,

Kufeld, Roland C. 1968
Inventory of range manipulation projects in
Coloradoo
Coloo Divo Game, Fish and Parks.
P-R Project W-lOl-RlO,
WP4, Jla, Quarto Repo July, Part 1, po 1-121.
0

Weintraub, Frances C. 1953
Grasses introduced into the United States.
U.S. Dept. of Agri., Agri. Handbook No. 58.
0

Prepared

by

f(do.4! C / %~P-UY/
Roland C. Kufeld
Wildlife Researcher

�- 67 -

Table 1. Acreages of rangeland treated during 1966 through 1969, inclusive,
in Colorado on lands administered by the U.S. Forest Service.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded As
Part of Treatment

Grassland

Spray

2

846

0

Meadow

Spray

1

240

0

Spray

2

355

0

Sagebrush

Spray

39

19,940

455

Rabbitbrush

Spray

3

860

0

Browse

Spray
Chain

2
1

467
300

0
300

Subtotal

3

767

300

Pinyon-Juniper

Chain

6

5,975

5,975

Annual Weeds

Spray

1

300

0

All Veg. Types

GRAND TOTAL

57

29,283

6,730

Perennial

Forbs

�- 68 -

Table 2. Acreages of rangeland treated during 1966 through 1969, inclusive,
in Colorado on lands administered by the U.S. Bureau of Land Management.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Grassland

Seed only
Plow

2
1

129
600

129
600

Subtotal

3

729

729

Seed only
Plow
Spray
Chain
Brush Cutter

1
7
18
3
3

120
2,802
27,398
3,525
1,728

120
2,802
0
1,730
0

Subtotal

32

35,573

4,652

Rabbitbrush

Seed only

1

373

373

Pinyon-Juniper

Seed only
Chain

1
31

1,200
31,723

1,200
23,657

Subtotal

32

32,923

24,857

Seed only
Plow

5
1

2,137
1,275

2,137
1,275

Subtotal

6

3,412

3,412

Sagebrush

Desert Shrub

Acres
Treated

Acres Seeded As
Part of Treatment

Abandoned

Lands

Seed Only

2

592

592

Accidental

Burn

Seed only

2

915

915

GRAND TOTAL

78

74,517

35,530

All Veg. Types

�- 69 -

Table 3. Acreages of rangeland treated during 1966 through 1969, inclusive,
in Colorado on lands administered by the U.S. Bureau of Indian Affairs.
Vegetative
Type

Kind of
Treatment

Sagebrush
Pinyon-Juniper

All Veg. Types

No. of
Projects

Acres
Treated

Acres Seeded As
Part of Treatment

Plow

1

701

701

Chain
Burn

1
1

22,986
4,600

22,986
4,600

Subtotal

2

27,586

27,586

GRAND TOTAL

3

28,287

28,287

�- 70 -

Table 4. Acreages of rangeland treated during 1966 through 1969, inclusive,
in Colorado on lands administered by the U.S. Forest Service, Bureau of Land
Management and Bureau of Indian Affairs.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Grassland

Seed only
Plow
Spray

2
1
2

129
600
846

129
600
0

Subtotal

5

1,575

729

Spray

1

240

0

Spray

2

355

0

Seed only
Plow
Spray
Chain
Brush Cutter

1
8
57
3
3

120
3,503
47,338
3,525
1,728

120
3,503
455
1,730
0

Subtotal

72

56,214

5,808

Seed only
Spray

1
3

373
860

373
0

Subtotal

4

1,233

373

Spray
Chain

2
1

467
300

0
300

Subtotal

3

767

300

Seed only
Chain
Burn

1
38
1

1,200
60,684
4,600

1,200
52,618
4,600

Subtotal

40

66,484

58,418

Seed only
Plow

5
1

2,137
1,275

2,137
1,275

Subtotal

6

3,412

3,412

Annual Weeds

Spray

1

300

0

Abandoned

Lands

Seed only

2

592

592

Accidental

Burn

Seed only

2

915

915

GRAND TOTAL

138

132,087

70,547

Meadow
Perennial

Forbs

Sagebrush

Rabbitbrush

Browse

Pinyon-Juniper

Desert

Shrub

Acres
Treated

Acres Seeded As
Part of Treatment

---

All Veg. Types

�- 71 -

Table 5. Acreages of rangeland treated during 1966 through 1969, inclusive,
on the Grand Mesa National Forest.
Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Perennial Forbs

Spray

2

355

0

Sagebrush

Spray

1

1,450

0

All Veg. Types

GRAND TOTAL

3

1,805

0

Vegetative
Type

Table 6. Acreages of range land treated during 1966 through 1969, inclusive,
in Colorado on the Gunnison National Forest.

Vegetative
Type

lUnd of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Sagebrush

Spray

13

8,233

0

All Veg. Types

GRAND TOTAL

13

8,233

0

Table 7. Acreages of rangeland treated during 1966 through 1969 inclusive on
the Rio Grande National Forest.
Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Meadow

Spray

1

240

0

Rabbitbrush

Spray

3

860

0

All Veg. Types

GRAND TOTAL

4

1,100

0

Vegetative
Type

�- 72 -

Table 8. Acreages of rangeland treated during 1966 through 1969, inclusive,
on the Routt National Forest.

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Sagebrush

Spray

4

1,590

0

Browse

Spray

1

67

0

All Veg. Types

GRAND TOTAL

5

1,657

0

Table 9. Acreages of rangeland treated during 1966 through 1969, inclusive,
on the San Juan National Forest.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grassland

Spray

2

846

0

Browse

Chain

1

300

300

Pinyon-Juniper

Chain

1

1,499

1,499

All Veg. Types

GRAND TOTAL

4

2,645

1,799

Table 10. Acreages of rangeland treated during 1966 through 1969, inclusive,
on the Uncompahgre National Forest.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Sagebrush

Spray

8

3,341

455

Browse

Spray

1

400

0

Pinyon-Juniper

Chain

5

4,476

4,476

Annual Weeds

Spray

1

300

0

All Veg. Types

GRAND TOTAL

15

8,517

4,931

�- 73 Table 11. Acreages of rangeland treated during 1966 through
on the White River National Forest.

1969, inclusive

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded As
Part of Treatment

Sagebrush

Spray

13

5,326

o

All Veg. Types

GRAND TOTAL

13

5,326

o

Table 12. Acreages of rangeland
on the Craig BLM District.
Vegetative
Type

Kind of
Treatment

Sagebrush

Pinyon-Juniper

treated during

1966 through 1969, inclusive

No. of
Projects

Acres
Treated

Seed only
Plow
Spray

1
3
7

120
1,623
8,457

120
1,623
0

Subtotal

11

10,200

1,743

Seed only
Chain

1
5

1,200
9,277

1,200
4,477

Subtotal

6

10,477

5,677

Acres Seeded As
Part of Treatment

Abandoned

Lands

Seed only

2

592

592

Accidental

Burn

Seed only

1

865

865

GRAND TOTAL

20

22,134

8,877

A 11 Veg. Types

�- 74 Table 13. Acreages of rangeland treated during 1966 through 1969, inclusive,
on the Montrose BLM District.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grassland

Seed only

1

100

100

Plow

1

600

600

Subtotal

2

700

700

Plow

4

1,179

1,179

Spray

9

16,241

0

Subtotal

13

17,420

1,179

Pinyon-Juniper

Chain

11

9,281

9,281

Desert Shrub

Plow

1

1,275

1,275

All Veg. Types

GRAND TOTAL

27

28,676

12,435

Sagebrush

Table 14. Acreages of rangeland treated during 1966 through 1969, inclusive,
on the Canyon City BLM District.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grassland

Seed only

1

29

29

Sagebrush

Spray

2

2,700

0

Rabbitbrush

Seed only

1

373

373

Pinyon-Juniper

Chain

3

586

586

Accidental Burn

Seed only

1

50

50

All Veg. Types

GRAND TOTAL

8

3,738

1,038

�- 75 Table 15. Acreages of rangeland treated during 1966 through 1969, incluSive,
on the Grand Junction BLM District.
No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Chain

3

3,525

1,730

Brush cutter

3

1,728

0

Subtotal

6

5,253

1,730

Pinyon-Juniper

Chain

12

12,579

9,3l3

Desert Shrub

Seed only

5

2,l37

2,l37

All Veg. Types

GRAND TOTAL

23

19,969

l3,180

Vegetative
Type

Kind of
Treatment

Sagebrush

Table 16. Acreages of rangeland treated during 1966 through 1969, incluSive,
on the Southern Ute Indian Reservation.
No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Plow

1

701

701

Pinyon-Juniper

Burn

1

4,600

4,600

All Veg. Types

GRAND TOTAL

2

5,301

5,301

Vegetative
Type

Kind of
Treatment

Sagebrush

Table 17. Acreages of rangeland treated during 1966 through 1969, inclusive,
on the Ute Mountain Indian Reservation.
No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Chain

1

22,986

22,986

GRAND TOTAL

1

22,986

22,986

Vegetative
Type

Kind of
Treatment

Pinyon-Juniper
All Veg. Types

�- 76 Table 18. Comparison of rangeland acreages treated during 1966 through 1969
on each National Forest, B.L.M. District and Indian Reservation in Colorado.
Land Management
Agency

fI)

+.J
fI)
(J)

1-1

0
I'&lt;.

..-t
C\I
s::
0
•.-1

+.J
C\I

Z

fI)

+.J
·
;:E: •.-1

()

•

1-1

o-l+.J

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Uncompahgre
Gunnison
White River
San Juan
Grand Mesa
Routt
Rio Grande
Arapahoe
Pike
Roosevelt
San Isabel

15
13
13
4
3
5
4
0
0
0
0

8,517
8,233
5,326
2,645
1,805
1,657
1,100
0
0
0
0

4,931
0
0
1,799
0
0
0
0
0
0
0

Subtotal

57

29,283

6,730

Montrose
Craig
Grand Junction
Canyon City
Glenwood Springs

27
20
23
8
0

28,676
22,134
19,969
3,738
0

12,435
8,877
13,180
1,038

Subtotal

78

74,517

35,530

Southern Ute
Ute Mountain

2
1

5,301
22,986

5,301
22,986

Subtotal

3

28,287

28,287

138

132,087

70,547

0

• fI)

r:Q •.-1

0

s::
0
•.-1

S::+.J
C\I

C\I

•.-1 :&gt;
"C

1-1

s:: (J)

H

fI)
(J)

~

GRAND TOTAL

�- 77 -

Table 19. Acreages of rangeland treated during 1966 through 1969 in the
Southwest Region of Colorado Game, Fish and Parks Division.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grassland

Seed only
Plow
Spray

2
1
2

129
600
846

129
600
0

Subtotal

5

1,575

729

Meadow

Spray

1

240

0

Sagebrush

Plow
Spray

5
32

1,880
30,515

1,880
455

Subtotal

37

32,395

2,335

Seed only
Spray

1
3

373
860

373
0

Subtotal

4

1,233

0

Spray
Chain

1
1

400
300

0
300

Subtotal

2

700

300

Chain
Burn

22
1

43,294
4,600

42,297
4,600

Subtotal

23

47,894

46,897

Desert Shrub

Plow

1

1,275

1,275

Annual Weeds

Spray

1

300

0

All Veg. Types

GRAND TOTAL

74

85,612

51,909

Rabb itbrush

Browse

Pinyon-Juniper

�- 78 Table 20. Acreages of rangeland treated during 1966 through 1969 in the
Northwest Region of Colorado Game, Fish and Parks Division.
Vegetative
Type

Kind of
Treatment

Perennial Forbs
Sagebrush

No. of
Projects

Acres
Treated

Spray

2

355

0

Seed only
Plow
Spray
Chain
Brush cutter

1
3
25
3
3

120
1,623
16,823
3,525
1,728

120
1,623
0
1,730
0

Subtotal

35

23,819

3,473

Browse

Spray

1

67

0

Pinyon-Juniper

Seed only
Chain

1
13

1,200
16,804

1,200
9,735

Subtotal

14

18,004

10,935

Desert Shrub

Seed only

5

2,137

2,137

Abandoned Lands

Seed only

2

592

592

Accidental Burn

Seed only

1

865

865

All Veg. Types

GRAND TOTAL

60

45,839

18,002

Acres Seeded as
Part of Treatment

Table 21. Acreages of rangeland treated during 1966 through 1969 in the Southeast Region of Colorado Game, Fish and Parks Division.
Vegetative
Type

Kind of
Treatment

Pinyon-Juniper

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Chain

3

586

586

Accidental Burn

Seed only

1

50

50

All Veg. Types

GRAND TOTAL

4

636

636

�- 79 Table 22. Acreages treated with aerial and ground spraying equipment and
2, 4-D, and 2, 4, 5-T chemical herbicides.

Number
Aerial

49,766

99

Ground

640

1

TOTAL

50,406

100

2, 4-D

49,832

99

2, 4, 5-T

489

1

Unknown

85
100

50,406

TOTAL

Table 23.

Acres Treated
Percent

Acreages treated by anchor chaining, cabling, railing and bulldozing.

Vegetative
Type

Anchor Chain
and Bulldozers
No. of
Projects Acres

Kind of EguiEment Used
Rail
Cable
and
Bulldozers
and Bulldozers
No. of
No. of
Projects Acres
Projects Acres

Bulldoze Onl~
No. of
Projects Acres

Sagebrush

3

3,525

0

0

0

0

0

0

Browse

1

300

0

0

0

0

0

0

Pinyon-Juniper

38

60,524

0

0

0

0

1

160

TOTAL

42

64,349

0

0

0

0

1

160

�- 80 -

Table

24.

Plant
Categories

Grasses

Forbs

Plant species used to seed Colorado
Scientific
Name

u

Percent of Total
Acres Seeded lj

cristatum

58,304

83%

Agropyron

intermedium

31,865

45

Elymus

junceus

29,866

42

Bromus

inermis

28,787

41

Agropyron

spp.

25,632

36

Agropyron

elongatum

3,372

5

Agropyron

smithii

815

&lt;2

Agropyron

sibiricum

766,

&lt;2

Agropyron

inerme

701

&lt;2

Agropyron

trachycaulum

446

&lt;2

Dactylus

glomerata

300

&lt;2

Bromus sppo

100

&lt;2

Hilaria

jamesii

67

&lt;2

Oryzopsiis lry;men6ides

67

&lt;2

13,872

20

10,640

15

Melitotus

sppo

Atriplex

canescens

13,865

20

Purshia

tridentata

10,866

15

4,574

6

990

2

Cercocarpus

montanus

Cowania mexicana

l/Scientific

Acres
Seeded

Agropyron

Alfalfa
Browse

range lands.

names were taken from Harrington

(1954) and Weintraub

(1953).

llRepresents
the percent of the total number of acres seeded in Colorado during
1966 through 1969 (70,547 acres) which has been seeded with each plant species.
Only includes species used on more than 2 percent of the rangelands seeded.

�Table 25.

Methods used for reseeding rangelands.

II
Method of Seeding

Ground
Broadcast
Acres
%11
Seeded

Browse
Seeder or
Dribbler
Acres
%11
Seeded

Aerial Broadcast
&amp; Browse
Seeder or
Dribbler
Acres
%11
Seeded

Vegetation
Type

Drill
Acres
Seeded

%11

Aerial
Broadcast
Acres
%Jj
Seeded

Grassland

0

0

0

0

729

100

0

0

0

0

729

Meadow

0

0

0

0

0

0

0

0

0

0

0

Perennial Forbs

0

0

0

0

0

0

0

0

0

0

0

Total
Acres
Seeded

I

Sagebrush

2,887

50

2,431

42

35

&lt;1

455

8

0

0

5,808

Rabbitbrush

373

100

0

0

0

0

0

0

0

0

373

Browse

0

0

0

0

0

0

0

0

300

100

300

Pinyon-Juniper

0

o 42,338

76

393

&lt;1

2,140

4

11,110

20

55,981

Desert Shrubs

1,607

47

530

16

1,275

37

0

0

0

0

3,412

Annua 1 Weeds

0

0

0

0

0

0

0

0

0

0

0

A bandoned Lands

592

100

0

0

0

0

0

0

0

0

592

Accidental Burn

865

95

0

0

50

5

0

0

0

0

915

TOTAL

6,324

45,299

2,447

2,595

11,410

!/Tables includes seeding information for only a portion of the seeding projects inventoried.
method of seeding was not available for many seeding projects.
~/percent of total acres seeded in each vegetation type.

68,110

Infonnation on

00
to-'

�- 82 -

Table 26.

Seed application rates used in reseeding Colorado rangelands.

Rate Seeded
Lbs. IAcre II

Number of
Acres 11

Percent of Total
Acres Seeded

1-2

2,330

3.6

3-4

2,796

4.3

5-6

47,576

73.5

7-8

4,892

7.6

9-10

4,773

7.4

11-12

2,250

3.5

13-14

100

0.1

+14

0

0.0

64,717

100.0

TOTAL

11 Rate seeded is shown to the nearest pound per acre.
~I Table includes seeding rates for only a portion of the seeding projects
inventoried. Information on seeding rates was not available for many
seeding projects.

�- 83 Table 27. Average plant densities on sagebrush and pinyon-juniper range
type-conversion project areas prior to treatment. 1/
Sagebrush
Percent of
Total Acres

Pinyon-JuniEer
Percent of
Acres
Total Acres

Percent Plant
Density

Acres

0-10

132

0.9

2,455

25.8

11-20

7,257

50.2

6,227

65.5

21-30

2,638

18.2

274

2.9

31-40

160

1.1

554

5.8

41-50

610

4.2

0

0.0

51-60

2,200

15.2

0

0.0

61-70

1,176

8.1

0

0.0

71-80

300

2.1

0

0.0

81-90

0

0.0

0

0.0

91-100

0

0.0

0

0.0

TOTAL

14,473

100.0

9,510

100.0

1/ Table includes plant density data for only a portion of the sagebrush and
pinyon-juniper control projects inventoried. Information on plant density
was not available for many sagebrush and pinyon-juniper projects.

�- 84 -

Table 28. Average shrub composition on sagebrush and pinyon-juniper range
type-conversion project areas prior to treatment. 1/

Percent Shrub
Cover

Acres

Sagebrush
Percent of
Total Acres

Pinyon-JuniEer
Percent of
Acres
Total Acres

0-10

0

0.0

0

0.0

11-20

0

0.0

0

0.0

21-30

2,390

18.1

0

0.0

31-40

4,852

36.6

0

0.0

41-50

1,456

11.0

0

0.0

51-60

3,275

24.8

828

22.1

61-70

1,250

9.5

1,427

38.0

71-80

0

0.0

1,499

39.9

81-90

0

0.0

0

0.0

91-100

0

0.0

0

0.0

TOTAL

13 ,223

100.0

3,754

100.0

1/ Table includes shrub composition data for only a portion of the sagebrush
and pinyon-juniper control projects inventoried. Information on plant density
was not available for many sagebrush and pinyon-juniper projects.

�- 85 -

Table 29. Range condition for livestock on sagebrush and pinyon-juniper range
type-conversion project areas prior to treatment. 1/

Range Condition
for Livestock

Sagebrush
Percent of
Acres Total Acres

Pinyon-JuniEer
Percent of
Total Acres
Acres

Sagebrush and PinyonJuniEer Combined
Percent of
Total Acres
Acres

0.0

0

6.0

0

0.0

Good to Excellent 2,230

19.8

1,499

4.3

3,729

8.0

Good

6,922

61.4

7,000

19.9

13,922

30.0

Fair to Good

1,534

13.6

25,548

72 .6

27,082

58.3

587

5.2

1,110

3.2

1,697

3.7

Poor to Fair

0

0.0

0

0.0

0

0.0

Poor

0

0.0

0

0 0
0

0

0.0

Very Poor to Poor

0

0.0

0

0.0

0

0.0

Very Poor

0

0.0

0

0.0

0

0.0

11,273

100.0

35,157

100.0

46,430

100.0

Excellent

Fair

TOTAL

0

1/ Table includes livestock range condition data for only a portion of the sagebrush and pinyon-juniper control projects inventoried. Information on livestock
range condition was not available for many sagebrush and pinyon-juniper projects.

�- 86 -

Table 30. Erosion condition on sagebrush and pinyon-juniper range typeconversion project areas prior to treatment. 1/

Sagebrush
Percent of
Total Acres

Pinyon-JuniEer
Percent of
Acres
Total Acres

Erosion
Condition

Acres

None or Slight

12,447

48.0

274

0.7

None to Moderate

10,312

39.7

12,958

31.6

Moderate

1,000

3.9

26,297

64.0

Moderate to Severe 1,096

4.2

0

0.0

Severe

0

0.0

0

0.0

None to Severe

1,100

4.2

1,500

3.7

TOTAL

25,955

100.0

41,029

100.0

1/ Table includes erosion condition data for only a portion of the sagebrush and pinyon-juniper control projects inventoried. Information on erosion
condition was not available for many sagebrush and pinyon-juniper projects.

�- 87 -

Table 31. Acreages of rangeland treated to improve conditions for four classes
of livestock.

Vegetative
Types

TYEe of Livestock Range
SheeE and Goats
Horses
Cattle and
Acres Seeded as
Acres
of
as
No.
Acres
Seeded
Acres
No. of
Treated
Part
of Treatment
Projects
Projects Treated Part of Treatment

Grassland

5

1,575

729

0

Meadow

1

240

0

0

Perennial Forbs

2

355

0

0

Sagebrush

65

52,416

4,773

6

3,763

1,000

Rabbitbrush

3

860

0

1

373

373

Browse

2

700

0

0

Pinyon-Juniper

32

56,666

48,600

6

7,819

7,819

Desert Shrub

4

1,907

1,907

2

1,505

1,505

Annual Weeds

1

300

0

0

Abandoned Lands

2

592

592

0

Accidental Burn

1

50

50

1

865

865

118

115,661

56,651

16

14,325

11 ,562

TOTAL

�Table 320 Acres of mule deer range treated in Colorado during 1966 through 1969 by the U. S. Forest Service,
Bureau of Land Management, and Bureau of Indian Affairs.

Total Acres of Seasonal Deer Range Treated
Spring
Fall
Spring
Summer
Winter
Fall
Summer
Fall
Spring
Yearlong

Vegetative
Type

Kind of
Treatment

Winter

Grassland

Seed only
Plow
Spray

129
600
0

0
0
0

0
0
0

0
0
846

0
0
0

129
600
846

Subtotal

729

0

-0

0
0
0

846

--0

--0

1,575

Spray

0

240

0

0

0

0

240

Spray

0

0

355

0

0

0

355

0
861
0
0
0

120
3,503
44,636
3,525
1,728

Meadow
Perennial Forbs
Sagebrush

Rabbitbrush

Browse

,

Total

Seed only
Plow
Spray
Chain
Brushcutter

0
2,642
24,482
541
0

0
0
4,,1(i)1
0
0

120
0
9,524
1,254
1,728

0
0
5,020
0
0

0
0
1,509
1,730
0

Subtotal

27,665

4,101

12,626

5,020

3,239

-861

53,512

Seed only
Spray

373
0

0
0

0
0

0
0

373
860

373

860

-0

0
0

Subtotal

-0

0
860

0

1,233

Spray
Chain

0
300

400
0

0
0

0
0

0
0

67
0

467
300

Subtotal

300

400

0

0

0

67

767

0

--

-------------------------------------------------------------------------------------------------------------------

00
00

�Table 32. Acres of mule deer range treated in Colorado during 1966 through 1969 by the U. S. Forest Service,
Bureau of Land Management, and Bureau of Indian Affairs (continued).

Vegetative
Type
Pinyon-Juniper

Desert Shrub

Winter

Total Acres of Seasonal Deer Range Treated
Spring
Fall.
Spring
Summer
Winte:r
Fall
Summer
Fall
Spring
Yearlong

Seed only
Chain
Burn

lt200
48,521
0

0
4,931
0

0
1,338
0

0
2,393
0

0
2,671
0

0
830
4,600

1,200
60,684
4,600

Subtotal

49,721

4,931

1,338

2,393

2,671

5,430

66,484

Seed only
Plow

0
1,275

405
0

0
0

0
0

0
0

1,275

405

0

0

-0

405
1,275

Subtotal

-0

0
0

0

0

300

0

0

0

300

452

0

140

0

0

0

592

80,515

10,077

15,619

8,259

5,9l0

6,358

126,738

Kind of
Treatment

Annual Weeds

Spray

Abandoned Lands

Seed only

All Vegetative Types GRAND TOTAL

Total

-

1,680

ex&gt;

1.0

�Table 33. Acres of mule deer range treated in Colorado during 1966 through 1969, by the D. S. Forest
Service, Bureau of Land Management and Bureau of Indian Affairs where deer use was heavy at the time of
treatment.

Vegetative
Type
Grassland

Sagebrush

Browse

Pinyon-Juniper

Annual Weeds

Kind of
Treatment

Acres of Heav~-Dse Seasonal Deer Range Treated
Spring
Fall
Spring
Sunnner
Winter
Winter
Fall
Summer
Fall
Spring

Yearlong

Total

Seed only
Plow

29
600

0
0

0
0

0
0

0
0

0
0

29
600

Subtotal

629

0

0

0

0

0

629

Plow
Spray
Chain

335
2,655
0

0
0
0

0
494
0

0
0
0

0
0
0

2,990

0

494

1,730

-0

335
3,149
1,730

Subtotal

-0

0
0
1;730

5,214

--

Spray
Chain

0
300

0
0

0
0

0
0

0
0

67
0

67
300

Subtotal

300

0

0

0

0

67

367

Seed only
Chain

1,200
22,804

0
1,841

0
0

0
2,393

0
2,255

0
0

1,200
29,293

Subtotal

24,004

1,841

0

2,393

2,255

0

30,493

0

0

300

0

0

0

300

27,923

1,841

794

2,393

3,985

67

37,003

Spray

All Vegetative Types GRAND TOTAL

\0

0

�Table 340 Acres of elk range treated in Colorado during 1966 through 1969, by the U. So Forest Service,
Bureau of Land Management and Bureau of Indian Affairs where elk use was heavy at the time of treatment.

Vegetative
Type

Kind of
Treatment

Acres of Seasonal Elk Range Treated
Spring
Spring
Sunnner
Winter
Fall
Fall
Summer

Fall
Winter
Spring

Yearlong

Total

Seed only
Spray

29
0

0
0

0
0

0
846

0
0

0
0

29
846

Subtotal

29

0

0

846

--0

0

875

Meadow

Spray

0

0

0

0

240

0

240

Perennial Forbs

Spray

0

0

355

0

0

0

355

Grassland

\0
I-'

Sagebrush

Browse

Plow
Spray

35
14,528

0
3,006

0
6,751

0
2,406

Subtotal

14,563

3,006

6,751

2,406

Spray
Chain

400
300

0
0

0
0

0
0

Subtotal

700

0

-0

0
565

0
1,600

35
28,856

565

1,600

28,891

67
0

0
0

467
300

--

--

-67

0

0

--

767

Pinyon-Juniper

Chain

3,859

0

0

0

0

0

3,859

Annual Weeds

Spray

0

0

300

0

0

0

300

19,151

3,006

7,406

3,252

872

1,600

35,287

All Vegetative Types GRAND TOTAL

�- 92 Table 35. Acres of elk range treated in Colorado during 1966 through 1969, by
the U. S. Forest Service, Bureau of Land Management and Bureau of Indian Affairs
where elk use was heavy at the time of treatment.
Acres of Heav:y:Use Seasonal Elk Range Treated
Spring
Fall
Spring
Summer Winter
YearWinter
Fall
Summer
Fall
Spring
long

Vegetative
Type

Kind of
Treatment

Grassland

Seed only

29

0

0

0

0

0

29

Sagebrush

Spray

0

0

330

0

335

0

665

Browse

Spray

0

0

0

0

67

0

67

All Veg. Types

TOTAL

29

·0

330

:0

402

0

761

Total

Table 36. Acres of sage grouse range treated in Colorado during 1966 through 1969,
inclusive in Colorado on lands administered by the U. S. Forest Service and Bureau
of Land Management.

Vegetative
Type

Kind of
Treatment

Sagebrush

Seed only

Acres of Sage Grouse Range Treated
AlII
Heavily Used
Range
Range jj
120

120

Plow

2,307

1,307

Spray

26,751

16,525

Total

29,178

17,952

1/
- Acres where sage grouse use was heavy at the time of treatment.

�-

IMPORTANT:

':J.:J

-

When completing the blanks on this form foHow the instructions in the
Interagency Council on Wildlife Ecology booklet entitled "A System for
Inventory and Evaluation of Vegetation Modification Projects".

GFP #4A COMPLETED VEGETATION MODIFICATION PROJECT
Colorado Game, Fish and Parks Division
It

PROJECT DESIGNATION
Number
1.

----------------------------------

11*

111*

LAND STATUS
3.
Natl. Forest
4.
BLM District
5.
State Agency
6.
SCS Area
7.
Indian Agency

IV

County
Township

Name

District or Unit

Name

_
Allotment

_

LOCATION
GoFo&amp;P. Region
9.
12.

2.

10. WoCoO" District
13.

H. Game Mgmt.
Unit No.

--

Drainage
Sections

Range

P.M.

Dominant
SITE DESCRIPTION
15. Species
14.* Vegetative Type
17. Topography
16.* Elevation
20. Soil Type
_
Exposure
Percent
Slope
19.
18.*
21. Erosion Condition
2lA. Method Used to Determine Erosion Condition
For Elk
For Deer
22. Range Condition for Livestock
--22A. Method Used to Determine Range Condo Livestock
Deer
Elk
23. Forage Production for Livestock
For Deer
For Elk
23A. Method Used Determ.Forage Prod.Livestock
Deer
Elk
24. Vegetative Cover
25. Shrubs
26. Herbs
26A. Grass
27. Litter
28. Ground
28A. Method Used to Determine Vegetative Cover &amp; Composition
29. Agencies
30. Date
Remarks

--- _
------------

-----------------_

-----------------------------------------------------------------

V

CLIMATIC DATA
31. Avg. Annual Precipitation
33. Avg. Annual Temperature
33B. Avg. July Temperature
35. Avg. Snow Depth on Area

VI

RANGE USE (At Time of Treatment)
37.* Kind of Livestock
39. Carrying Capacity
41. Stocking Rate

Fig. 1.

Seasonal
Distribution
32.
33A. Avg. Jan. Tempo
Length Growing Season
34.
36.* Stag Reporting

38.* Season of Use
40. Grazing System
Remarks

Form used for recording inventory data.

�_________

"TJ.--

Wildlife
44A. How Use Determined
Remarks__ ~
VII

Ut:C1'&gt;VU,

~~.

of Use
.,.--__
45.Agencies______________

amount;

_

of Use
46.Date

_
_

_

TREATMENT
47.* No. Acres
Treated

48.* Purpose of
_
Treatment,
_
49.* Kind of Treatment
-"..,,.50. Equipment Used,
_
51.* Date Treated
52.* Equipment, Chemicals
_
53.* Seeded: Yes
No
54.* How Seeded
55.* Date Seeded--5-6-.*--S-p-e-c-1-·e-s--S-e-e-d-e-d---------57.* Rate Seeded
58.* Protection Given
Location from Other projects
_
Remarks
_

------------------

MAP OF PROJECT LOCATION

T

R

--

6

5

4

3

2

1

6

5

4

3

2

1

~

8

9

10

11

12

7

8

9

10

11

12

18

17

16

15

14

13

18

17

16

15

14

13

19

20

21

22

23

24

19

20

21

22

23

24

30

29

28

27

26

25

30

29

28

27

26

25

31

32

33

34

35

36

31

32

33

34

35

36

R

.

T

6

5

4

3

2

1

6

5

4

3

2

1

7

8

9

10

11

12

7

8

9

10

11

12

18

17

16

15

14

13

18

17

16

15

14

13

19

20

21

22

23

24

19

20

21

22

23

24

30

29

28

27

26

25

30

29

28

27

26

25

31 '

32

33

34

35

36

31

32

33

34

35

36

*

Items "flagged" are or primary importance, and special effort should be
made to obtain information.
Form Completed by
Title
Agency
Date,
Show title and agency of all persons nar t i.c Lna t Ln o in cnmnletion
of formq,

_

�- 95 -

July,

1970

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-10l-R-12

Work Plan No.

4

Game Range Investigations
Job No.

2

Experimental Improvement of Oakbrush
Job Title __~O~n~D~e~e~r~a~n~d~E~l~k~W~i~n~t~e~r~R~a~n~g~e~s
__-~B~e~a~v~e~r~~C~r~e~e~k~
Period Covered:

April 1, 1968 through March 31, 1970.

Personnel:

Roland C. Kufeld, Ronald Lambertson,
Gary Montgomery, George Bock.

Mark Fra sier,

ABSTRACT

One hundred and fifteen circular .001 acre pellet plots, and 225 point
quadrat vegetation measurement plots were established on the 135 acre
Beaver Creek study area. Pre-treatment vegetation measurements were
made and pellet plots cleared during August, 1968. Accumulated deer
and elk pellets were counted in May, 1969. Vegetative ground cover
prior to treatment was 127.66 percent.
Percent composition was 43.16
shrubs, 34.36 forbs, 22.03 grasses and 0.33 lichens and moss. During
the period August 19, 1968, to May 6, 1969, the area received 68 deer
days and 43 elk days use per acre.
On July 10, 1969, 67 acres encompassing the eastern half of the study
area were sprayed with 2 pounds of 2-4-5TP and 10-1/2 gallons of water
per acre applied by helicopter.
The western half of the area will be
used for control.

_

��- 97 -

OAKBRUSH

EXPERIMENTAL ll1PROVEMENT OF
ON DEER AND ELK WINTER RANGES - BEAVER CREEK
Roland

C. Kufeld

INTRODUCTION
Gambel oak (Quercus gambellii) is one of the five most important deer and
elk winter browse plants in Coloradoo
Extensive stands of Gambel oak are
found on big game winter ranges throughout the western slopeo Many of
these oakbrush stands have grown beyond the reach of deer and elk, however,
and provide little forage for wintering big game animalso
Such overage
stands need to be treated to increase their production of available big
game winter forageo
This study is designed to determine if oakbrush productivity and the amount
of deer and elk use can be increased on overage oakbrush ranges by spraying
with 2-4-5TP to induce sproutingo
It is a cooperative project between
Colorado Division of Game, Fish and Parks and the U.So Forest Service, Routt
National Forest.
The 135 acre study area is located on the Routt National
Forest approximately 28 miles southwest of Hayden, Colorado (T3N, R89W, S 9,
10, 15, 16; 6th PoMo)o
Dominant plant species on the area are Gambel oak (Quercus gambellii), snowberry (Symphoricarpos sp.), serviceberry (Amelanchier alnifolia), and chokecherry (Prunus virginiana).
Two small aspen pockets (Populus tremuloides)
also occur on the area.
Elevation ranges from 8,000 to 8,500 feet. Slopes
are predominantly southwest facing and are relatively steep; from 30 to 90
percent.
The area is used by elk during the fall, winter and spring, and
yearlong by deer. Most deer use, however, occurs during the spring, summer
and fall. The area has been set aside for wildlife by the U.S. Forest
Service and is closed to livestock grazing.

Po So OBJECTIVE
To determine if deer and elk carrying capacity can be increased on oakbrush
winter game ranges by spraying with 2-4-5TP to induce sprouting.

SEGMENT OBJECTIVES
This study has been underway for two years, however, this is the first
annual job completion report written concerning the studyo A report was not
prepared for the first year because that year's activities were charged to
FW-19-C-28 (See Amendment number 1, December 19, 1968). Although this is
the second year of the study it is the first segment of W-lOl-R, WP 4, J2.
Therefore, this report will include progress made during the first two years
of the study. Objectives for the first two years are as follows:

�- 98 -

1.

To establish permanent point quadrat vegetative measurement
plots, and measure percentage composition and cover of all
plant species on the study area prior to treatment.

20

To establish permanent pellet group plots, and measure
and elk use on the study area prior to treatment.

3.

To establish permanent
treatment photos.

4.

To spray one half of the 135 acre study area with 2-4-5TP
after all pre-treatment measurements have been madeo

METHODS
Point Quadrat

photo point stations

deer

and take pre-

AND MATERIALS

and Pellet Group Plot Establishment

One hundred and fifteen .001 acre circular pellet plots, and 225 pOint
quadrat plots were established in the study area during the summer of
1968. Plots were located at 100 ft. intervals along parallel lines,
which stretch the entire width of the study area.
Separate lines were
used for pellet and point quadrat plots. A baseline was established
along one side of the study area, and the first plot on each line was
located on the baseline.
A detailed map of the study area was made
which shows the location of each plot.
The center of each pellet group plot was marked with a 3/8" round steel
stake. Each point quadrat plot was marked with two angle iron stakes
placed 67 inches apart over which the point quadrat frame can be erected.
Both stakes were located on the line. Plot stakes were marked with
numbered metal tags bearing the letters "pp" if it was a pellet plot or
"PQ" if it was a pOint quadrat plot, the line letter (there are 12 point
quadrat lines, A through L, and 6 pellet group plot lines, A through F),
and plot number along the line. Strips of plastic surveyor's flagging
were tied to each stake and to an overhanging tree branch near the stake.
Blue flagging was used for pellet plots, and red for point quadrat plots.
Plot locations were laid out with a K &amp; E compass set to allow for 14
degrees east declination, and a steel tapeo
When measuring distances no
allowance was made for slope.
The tape was always pulled tight (Figo 1).

Photo Point Establishment
Sixty-seven photo points were established and pre-treatment photos were
taken between August 30, and September 2, 1968
Certain pOint quadrat
and pellet group plots were selected for photo pOint stations.
An llx16
inch blackboard with the plot number written in chalk was suspended from
a nearby branch.
This became the photo center and focusing point. The
compass bearing and distance from the plot stake (the easternmost stake
0

�- 99 -

was always used in the case of pOint quadrat plots) to the center of the
blackboard, and the date and time of day the photo was taken were recorded.
The camera was held directly over the stake at from waist to
chest level
Two photos were taken at each station.
A black and white
photo was taken with a Ciroflex camera using 120 plus X film, and a 35
rom color slide was made with a Praktica camera using Kodachrome II film.
A sample photo point picture is shown in Fig. 20
o

Pre-treatment

Vegetation

Measurements

The vertical point quadrat method, originated by Levy and Madden (1933),
and further described by Medin (1965), utilizing both crown and basal
contacts was used to describe the vegetation during August, 1968. The
equipment consisted of a tubular 7/8" aluminum conduit horizontal frame
mounted on 5/8" steel rod uprights
Thumb screws on the horizontal frame
allowed adjustment to a height of 505 feet above the ground surface.
Two
small 3-inch line levels, facing at right angles, were attached to the
frame for leveling the equipment in both the vertical and horizontal planes.
Two sleeves with thumbscrew adjustments were welded to the bottom of the
vertical uprights.
The frame was positioned for reading by driving one
18" X 3/4" angle-iron stake into the ground, placing one sleeve on this
stake and orientating the frame on the west side and at a right angle to
the transect line of travelo
The westerly stake was then driven into the
ground through the sleeve and the frame leveled with thumbscrews.
After
the data for a plot had been recorded the thumbscrews were loosened and
the entire frame removed leaving the stakes in permanent positiono
o

Ten guide holes 6 inches apart were drilled through the conduit frame for
quadrat pin projection.
Pins were made of 3/16" carbon-tested drill rod
in 6-foot lengths and sharpened to a needle point with a bevel approximately
1 inch long.. Clothes pins were used as a brake to hold the quadrat pins at
any desired height.
Point contacts were recorded in two ways. All hits from a height of 5.5
feet to ground surface were "crown" contacts and recorded by I-foot height
intervals.
The same plant or different ones may have been hit once or more
than once. After crown contacts were recorded the pin was lowered until it
struck a plant base, litter, rock, erosion pavement, or bare soil; this
was called a "basal" contact.
Definitions of the various categories of
contacts are as follows:
Ground cover.--The proportion of the ground surface occupied by
vertical projection of live plant parts, both basal and aerial,
plus that occupied by nonliving matter such as litter, rock, and
dead plant parts.
Crown cover.--The proportion of the ground surface occupied by vertical projection of live aerial plant parts, plus that occupied by
nonliving matter such as litter, rock, and dead plant parts
o

Basal cover.--The proportion
parts at the ground surface,

of the area occupied by live plant
or the area defined by the live root

�- 100 -

crown, plus that occupied by nonliving matter such as litter, rock,
and dead plant partso
The basal area of plants with basal rosettes
was understood to be the area defined by the live root crown only;
the rest of the live parts were considered crown covero
Littero--Dead organic material either lying on or projecting above
the soil surfaceo
All dead plant parts, with the exception of woody
plants which were recorded by species, were considered as litter.
Bare soil.--All exposed mineral soil and rock particles up to 1/8inch diameter, and well-dispersed
rock particles up to 3/4-inch
diameter which do not provide a continuous covero
Rock.--Stones
sur f'ace ,

larger than 3/4-inch

diameter

appearing

at the soil

Erosion pavemento--Partic1es
of rock from 1/8 to 3/4-inch diameter
forming a continuous cover on the soil surface.
Individual rock
particles 1/8 to 3/4-inch diameter that do not form a continuous
cover were classified as bare soi1
9

The pOint quadrat technique is illustrated in Figo 3, and the form used
to record pOint quadrat data is shown in Figo 40
Two specimens of each species of plant encountered on point quadrat plots
were collected, mounted and identifiedo
Facilities of the Colorado State
University Herbarium were used in making plant identifications,
and
identifications were checked and annotated by CoS.U. Botany Department
personnel.
One set of specimens was contributed to the CeSoU. Herbarium,
and one set was deposited in the Herbarium at the Game Research Center.

Pre-treatment

Deer and Elk Use Measurements

Pellet groups were cleared from pellet plots on August 19, 1968, and
accumulated deer and elk pellet groups counted on May 6, 1969
A pellet
deposit was considered a "group" if it contained more than five pellets
and was distinct from other groups in or near the p1oto A group was
counted if more than half of it fell inside the plot.
0

The .001 acre circular plot boundary was delineated by stretching a 31
8•.3/4" card from a jacob staff placed in the ground adjacent to the center
stake.

Treatment
A few days prior to treatment the spray area, which encompassed the eastern
one-half of the study area or a tota 1 of 67 acres, was flagged with
fluorescent orange plastic sheeting and surveyor1s flagging.
This area
was sprayed with 2 pounds of 2-4-5TP and 10-1/2 gallons of water per acre
applied by helicopter on July 10, 1969, (Figo 5). Spraying was done by
Skychoppers, Inc. of Durango, Co1oradoo

�- 101 -

Fig. 1.

Fig. 2.

Surveying plot locations.

A sample photo point picture prior to treatment.

�- 102 -

Fig. 3.

Measuring

vegetation

with the pOint quadrat.

�- 103 POINT QUADRAT VEGETATIVE FORM
Area
Line &amp; Plot No.

Date
--------------------------- Vegetative
Type

1 2 3 4 5 6 7 8 9 10

-----------------------------------------------------------

-----------

p.~n No.

Species

Observers
Comments

H e~g
. ht 0 f Sspec i.es C ontac t
Ground
Surface

Under
l'

I' -2'

2' -3'

3'-4'

4'-5'

5'-5.5'

Tota 1
Hits

Dead
Browse
Rock
Litter
BareGrd
TOTALS
Line &amp; Plot No.

-----------------

Vegetative Type

----------

p.~n N o.

Species

1 2 3 4 5 6 7 8 9 10

Comments

_

H·h
e~gl t 0f Sspec i.es C ontact
Ground
Surface

Under
i:

I' -2'

2' -3'

3'-4'

4'-5'

5'-5.5'

Dead
Browse
Rock
Litter
BareGrd
TOTALS
Note:

Plot frame oriented East-West.

Pin #1 always westernmost.

Fig. 4~ Form used for recording point quadrat data.

Use dot tally.

Tota 1
Hits

�- 104 -

Figure

5 - Half of the area was sprayed by helicopter on July 10, 1969,
with 2 pounds of 2-4-5TP and 10-1/2 gallons of water per acre.

�- 105 -

Weather was generally excellent during the application.
The sky was
clear, temperature was in the 70's, and wind velocity was less than 5
miles per hour, except for a short period when gusts of 7 to 8 miles
per hour occurred.
Spraying was stopped at that point and resumed 2
hours later after wind velocity dropped to about 2 miles per hour.
Wind direction was from the west.
Oil sensitive cards were placed in selected point quadrat plots the
day prior to spraying, and retrieved the day following in order to
measure the spray coverage, and to determine the amount of drift onto
the 68 acre control area, which lies west of and adjacent to the sprayed
area. Almost no drift was recorded on the control area.

RESULTS AND DISCUSSION
Pre-treatment

Vegetation

Measurements

The pre-treatment vegetation description discussed here is for the entire
study area.
Separate vegetation descriptions for the spray and control
areas will be made in a future report.
First, however, it will be
necessary to classify pOint quadrat plots along the boundary between the
two areas according to whether they are sprayed or unsprayed.
This will
be done during the next segment.
Vegetative cover on the study area prior to treatment was relatively heavy.
The layers of live crown vegetation up to a level of 5.5 feet covered the
ground 127.66 percent.
The addition of dead browse, litter and rock resulted in a total ground cover from crown level material of 221.56 percent
(Table 1). Had the shrub and tree crown vegetation above 5.5 feet been
measured the ground cover percentage would have been much higher.
It was
not measured, however, because it was above the reach of game animals,
and could not be measured with the point quadrat technique.
The ground was covered with a heavy layer of litter.
Litter made up 71.11
percent of the cover at ground surface level, and 84.83 percent of the
ground cover above ground surface.
Shrubs (woody and semi-woody vegetation) were the most abundant species
comprising 43.16 percent of the total vegetation (Table 2). Forbs made
up 34.36 percent, grasses 22.03 percent and lichen and moss 0.33 percent.
Snowberry was the most abundant shrub measured, and it comprised 23.13 percent of the vegetation on the study area below 5.5 feet. Although Gambel
oak is the dominant plant species on the area it comprised only 8.12 percent of the total vegetation measured below 5.5 feet. Most of the oak was
too tall to be measured, as was most of the other major species, serviceberry and chokecherry, which respectively made up 3.72 and 6.12 percent
of the total vegetation below 5.5 feet.
Pre-treatment

Deer and Elk Use Measurements

Deer and elk use on the study area during the fall,winter
and spring of
1968 and 1969, was relatively heavy.
During the period August 19, 1968 to

�- 106 -

May 6, 1969, inclusive, pellet group counts indicated that the study area
received 68 deer days and 43 elk days use per acreo Separate pre-treatment game use estimates for the spray and control areas will be made in
a future report after pe11~t group plots along the spray-control area
boundary have been classified according to whether they are sprayed or
unsprayedo
.

LITERATURE CITED
Harrington, H. D. 1954
Denver
666 p.

0

Manual of the plants of Colorado.

Sage Books,

0

Levy, E. B., and E. A. Madden. 1933
The pOint method of pasture
analysis. New Zealand Jour. Agr. 46:267-279.
0

Medin, D•.E. 1965. Vegetative analysis.. Colo. Depto Game, Fish and
Parks. P-R Project W-l05-R-5, WP3, J3, Quarto Repo January, Part
3, po 345-398.

Prepared

by~CI~f4
Roland C. Kufe1d
Wildlife Researcher

�Table I.

Pre-treatment ground cover percentages of combined Beaver Creek spray and control areas.

% Crown Cover
Plant Species 11

Ground
Surface

Under l'

1'-2'

2'-3'

3'-4'

4'-5'

1.47
.18
.04
.93
4.36
.13
.62
10.00

1.78
.22

.89

.40

.27

2.00
1.42
.04

1.02
.98

.13
.27

--

.71

.09

--

21.24

9.33

4.57

1.02
.18
.09

.13
.04

.18

.09

5'-5.5'

Total Crown
Cover %

Woody and Semi-Woody
Amelanchier alnifolia
Artemisia tridentata
Populus tremuloides
Prunus virginiana
Quercus gambellii
Ribes sp.
Rosa sp ,
Symphoricarpos sp.

.04
.13

Subtotal

.17

17.73

Achillea lanulosa
Agastache urticifolia
Aster sp.
Artemisia dracunculus
Artemisia ludoviciana
Chenopodium berlandieri
Circium canescens
Crepis runcinata
Descurainia tichatdsonii
Collomia linearis
Erigeron spp.
Eriogonum umbellatum
Fra garia ova lis
Ga lium borea Le
Geranium richardsonii
Heracleum lanatum

.09

7.56
2.49
.84
.04
.04
.13
1.33
.13
.13
.67
2.67
.44
.22
3082
.53
.13

2.09
2.22
.13
.84
13.96

.04
1.69
1.11
.44
5.16

2.36

.40

4.81
.40
.08
7.86
10.36
.30
1.90
29.92
55.63

Forbs

.04

.04

,04
.22
.09
.04
.04
.44

-----------------------------------------------------------------------------------------------------------------

t-'

0
-....J

7.56
3.64
1.06
.13
.04
,13
1.60
.13
,13
.71
2.89
053
022
3.86
.57
.57

�Table 1.

Pre-treatment ground cover percentages of combined Beaver Creek spray and control areas (continued).

% Crown Cover
Plant Species 11
Forbs (continued)
Helenium hoopesii
Lactuca pulchella
Lathyrus leucanthus
Ligusticum sPa
Linaria vulgaris
Linum Lewf.s Lf
Mertensia franciscana
Osmorhiza occidentalis
Pedicularis grayi
Pteridium aguilinum
Senecio serra
Solidago canadensis
Swertia radiata
Taraxicum sp ,
Thalictrum fendleri
Tragopogon pratensis
Viguiera multiflora
Urtica gracilenta
Lupinus argenteus
Unknown Forb

Ground
Surface

.04

.04

.04

-Subtotal
Grass and Grasslikes
Bromus anomalus
Carex SPa
Elymus cinereus
Elymus glaucus
Ph1eum pratense
Poa spp.
Sti£!!..
lettermani

-- ---'-..~--,.....-.•--- .. - - -- ----- - ------

-

.29

.04

Under l'

027
.53
3.16
.62
.27
.04
2.00
.53
.09
.04

.04
.27
1.02
.04
1.87
.18
1.24
.09
33.47

1'-2'

1.33

2'-3'

.27

3'-4'

4' -5'

5'-5.5'

.27
053
4.80
.62
.71
.04
4.40
062
.09
.32
.04
.04
004
.27
2.22
.04
2.14
.27
2.79
.13

.04

.44
2.18
.09

.22

.27
.04
.04

.01

1.16

.04

.27
.09
.93
.04

.58

.04

--

-1.38

--

9.22

.08

Total Crown
Cover %

-0.00

0.00

44.15

2.22
.18
.04
2.44
1.06
1.02
.04
.09
.04
.44
.53
2.71
.80
.09
3.60
.13
.09
.22
16.67
.31
.67
17.34
.09
.09
-- - -- -- -- - - - -- - --- -- - -- - --- -- - - - - -- - - -- - -- - -- - -- - - -- -- - - -- - - -- --- --- - - -- - - -- -- -- - -- --

to-'

0

co

�Table I.

Pre-treatment ground cover percentages of combined Beaver Creek spray and control areas (continued).

% Crown Cover
Plant Species

Ground
Surface

1/

Grass and Grass1ikes
Agropyron spp.
Unknown Grass
Subtotal
Lichen and Moss
Lichen
Moss
Subtotal

Under i '

1'-2'

2.31
.22

.27

.39

25.81

2.14

.04

.36
004

3'-4'

4' -5'

5'-5.5'

Total Crown
Cover %

(continued)

004

.40

0.00

018
2013
71.ll
25.69

4.44
018
83042

Subtotal

ss.ri

TOTALS

100.00

Other
Dead Browse
Rock
Litter
Bare Ground

2 '-3'

.13

--

0.00

0.00

0..00

2004
013
1.24

.75

053

.13

004

88.04

3.41

.88

165.45

36.01

1l.72

1/ Scientific names are from Harrington (1954).

--

0.00

--

2.58
.22
0.00

.36
.04

t-'
0
\0

0.00

0000

.40

.36

.04

8.16
.31
84.83
0.00

--

--

.36

.04

5.22

2.72

.44

.57

28.08

--

93.30

221.56

�Table 2.

Pre-treatment composition of vegetative cover on combined Beaver Creek spray and control areas.

Level of Vegstation
Plant Species

11

Ground
Surface

Under l'

1 t -2'

2'-3'

3'-4'

4'-5'

1.13
014
.03
072
3037
.10
.48
7.73

1.37
.17

.70

.31

.21

1062
1072
.10
.65
10079

1.55
1010
.03

.79
.76

.34
3099

0

55

.07

16.42

7.23

3054

1.83

.79
.14
.07

.10
003

5'-5.5'

Tota 1. ,
Cover %

Woody and Semi-Woody
Ame1anchier a1nifolia
Artemisia tridentata
Populus tremu10ides
Prunus virginiana
Quercus gambe11ii
Ribes sPa
Rosa SPa
SymRhoricarpos spo

.03
.10

Subtotal

.13

l3.70

Achillea 1anulosa
Agastache urticifo1ia
Aster spo
Artemisia dracuncu1us
Artemisia 1udoviciana
Chenopodium ber1andieri
Circium canescens
Crepis runcinata
Descurainia richardsonii
Co11omia 1inearis
Erigeron spp.
Eriogonum umbe11atum
Fragaria ova lis
Ga lium borea le
Geranium richardsonii
Herac1eum 1anatum

.07

5.84
1092
.65
.CI03
.03
.10
1.03
.10
.10
.52
2.06
.34
.17
2.96
.41
.10

.03
1.31
.86

.10
021

.31

3.72
.31
.06
6.12
8.12
.23
1.47
23.l3
43.16

Forbs

.03

.03

.14

003
.17
.07
.03
.03
.34

5.91
2.81
.82
.1(1)'

007

.03
.10
1.24
.10
.10
.55
2.26
.41
.17
2.99
.44
.47

------------------------------------------------------------------------------------------------------------------

•.....
•.....
0

�Table 2. Pre-treatment composition of vegetative cover on combined Beaver Creek spray and control areas
(continued).

Level of Vegetation
Plant Species

1/

Forbs (continued)
Helenium hoopesii
Lactuca pulche11a
Lathyrus leucanthus
Ligusticum sp.
Linaria vulgaris
Linum lewisii
Mertensia franciscana
Osmorhiza occidenta1is
Pedicularis ~
Pteridium aqui1inum
Senecio serra
-Solidago canadensis
Swertia radiata
Taraxic:um sp ,
Tha1ictrum fend1eri
Tragopogon pratensis
Viguiera multiflora
Urtica graci1enta
Lupinus argenteus
Unknown Forb
Subtotal
Grass and Grasslikes
Bromus anoma 1us
Carex sp ,
E1ymus cinereus
Elymus glaucus
Phleum pratense
Poa spp.
Stipa 1ettermani

Ground
Surface

.03

Under l'

.21
.41
2.44
.48
.21
.03
1.55
.41
.07
.03

1'-2'

1.03

1.68
.07

.17

.21
.03
.03

.03

.89

.03

.21
.07
.72
.03

.22

25.87

7.12

003

1072
.79
.34
2.10
.10
12.89
.07

.14
.03
.07
.62
.07
.52

003

--

.03

.24

.21

3'-4'

4'-5'

5'-5.5'

.03

.34

.03
.21
.79
.03
1.44
.14
.96
.07

.03

2'-3'

.45

--

1.09

.03

.06

.03

.07

------------------------------------------------------------------------------------------------------------------

Total
Cover %

.21
.41
3.71
.48
.55
.03
3.40
.51
.07
.27
,03
,03
.03
.24
1.71
.03
1065
.21
2.19
.10
34.36

1.92
.82
.44
2.79
.17
13.65
.07

""'"
""'"
""'"

�Table 20 Pre-treatment composition of vegetative cover on combined Beaver Creek spray and control areas
(continued).

Level of Vegetation
Plant Species 11

Ground
Surface

Under l'

1'-2'

1079
017

021

2'-3'

3'-4'

4'-5'

5'-505'

Total
Cover %

Grass and Grasslikes (continued)
Agropyron sppo
Unknown Grass
Subtotal

-.10

030

19.97

.03

.27
.03

.30
.03

Subtotal

.03

.30

.33

TOTALS

.68

59.84

Lichen and Moss
Lichen
Moss

11

2.00
.17

Scientific names are from Harrington (1954).

1.66

25.20

8.42

22.03

3.60

1.83

031

99.88

~
~

I'.)

�July, 1970

- 113 -

JOB PROGRESS REPORT

Sta te of

.::.C.::.OL=OR.(\=D~O::._._
_

Project No.

W-10l-R-12

Work Plan No.

4

Job Title

Game Range Investigations
Job No.

3

Experimental Improvement of Oakbrush on
Deer, Elk and Cattle Ranges - Hightower Mountain

Period Covered:

April 1, 1969 through March 31, 1970.

Personnel:

Roland C. Kufeld,

Richard

Fentzlaff,

Fred Brooke.

ABSTRACT

A study area was selected at the northeastern end of the Grand Mesa
National Forest near Hightower Ranger Station (T9S, R92W, S17, 19, 20,
21, 28, 29, 30). Eight 60 acre habitat improvement units were established.
A thirty acre unit was laid out in the center of 6 of the 60 acre units.
The inside 30 acre portions will be treated and the outside 30 acres will
be left for wildlife cover. No 30 acre areas were established in the 2
control units.
All boundaries were marked with orange steel fence posts.
Sixty .002 acre circular pellet plots were established in each of the 8
habitat improvement units.
These were located using a modified system
of multiple random starts.
Plots were marked with orange 3/8" steel
stakes 24" long.
A vegetative measurement method was tested and selected for use during
the course of the study. The technique involves clipping and weighing
green weights of forbs and grasses by species, and estimating green weights
of individual browse species.
The sampling unit is a meter square plot.
Eighty preliminary meter square plot samples were taken on which to base
sample size estimates.
Herbage samples of various browse, grass and forb
species were collected periodically during the summer of 1969, and percentage moisture content determined.
A method was devised for converting
green weights of all plant species on the area to air dry weights.
Specimens of each species of plant found on the study area were collected,
mounted and identified.
A review of literature

was begun.

��- 115 -

EXPERIMENTAL
IMPROVEMENT OF OAKBRUSH ON
DEER, ELK AND CATTLE RANGES - HIGHTOWER MOUNTAIN
Roland

C. Kufeld

INTRODUCTION
Game and land managers have recognized the need for increasing the supply
of big game animals to meet the constantly rising demand for big game hunting.
The primary factor which limits the size of big game populations
in Colorado
is availability and condition of winter game range.
The amount of available
winter range is shrinking every year due to construction of dams and reservoirs, highways, urban, industrial and agricultural developments,
and oil,
gas and mineral exploration.
Still another factor is the practice of converting brushlands to grasslands primarily to improve the range for livestock
grazing.
Due to the declining availability
of winter range, efforts to increase big
game populations must be concentrated on protection and proper management
of existing winter ranges, and on improvement of wildlife habitat wherever
possible on these ranges to achieve greater productivity.
The oakbrush vegetative type is one of the most important types of winter
deer and elk range in Colorado.
As oakbrush becomes older, however, it
grows beyond the reach of deer and elk and loses its value as a game forage
species.
Since cattle grazing is also an important use of oakbrush ranges in Colorado,
this study will be designed to determine how overage Gambel oak winter game
ranges can be improved to increase deer and elk use and production of forage
for deer, elk and cattle on the same range.
This report

describes

activities

during

the first year of the study.

P. S. OBJECTIVE
To determine the extent to which deer, elk and cattle forage production and
game use can be increased and maintained through implementation
of practical
deer and elk habitat improvement measures on overage Gambel oak winter
game ranges.
SEGMENT
1.

To establish

eight,

2.

To install

permanent

OBJECTIVES

60 acre habitat
pellet

improvement

group plots

units.

in habitat

improvement

units.

�- 116 -

3.

To test selected

vegetative

measurement

4.

To collect and identify specimens
vicinity of the study area.

5.

To make a literature

techniques.

of each plant species

found in the

review.

METIlODS AND MATERIALS
Study Area Selection
The study area is located at the northeastern end of the Grand Mesa National
Forest near Hightower Ranger Station (T9S~ R92W, S17, 19~ 20, 21, 28, 29,
30) 6th P. M. Dominant plant species are Gambel oak (Quercus gambellii),
snowberry (Symphoriocarpos),
serviceberry (Amelanchier alnifolia), and
chokecherry (Prunus virginiana), and big sagebrush (Artem~sia tridentata).
Elevation ranges from 8,000 to 8,500 feet.
A number of potential study areas all located on U. S. Forest Service and
Bureau of Land Management Lands were considered when selecting a location
for this study.
The Hightower Mountain location was selected because: (1) the
area is representative of most of the oakbrush ranges in Colorado, and findings
derived there should be applicable in other areas of the state; (2) the oakbrush type there is large enough to permit establishment of 8, 60 acre habitat
improvement units in which to test vegetation modification methods; (3) slopes
are not so steep as to preclude testing of one or more vegetation modification
methods, slope varies from 0 to 30 percent; (4) the area recieves relatively
heavy use by deer and elk. Deer use the area primarily in spring, summer and
fall, but some winter deer use occurs during less severe winters.
Elk use
the area on a yearlong basis, but most elk use occurs during the fall-winterspring period; and (5) the area is grazed by cattle during the summer, and
the stocking rate is about average for most of the oakbrush ranges in the
state.
This will be a cooperative project between Colorado Division of Game, Fish
and Parks and the U. S. Forest Service, Grand Mesa-Uncompahgre
National
Forest.

Habitat

Improvement

Unit Establishment

Locations for habitat improvement units were tentatively selected through
the use of aerial photos and ground reconnaissance.
A joint field inspection
of the tentative locations was made by members of the U. S. Forest Service
and Colorado Game, Fish and Parks Division on June 11, 1969. During the
inspection, approval was given by the Grand Mesa-Uncompahgre
National Forest
Supervisor to establish habitat improvement units on these locations with the
stipulation that Unit Number 3, which is located adjacent to Buzzard Campground, not be treated by burning for aesthetic reasons.

�- 117 A randomized block design was used for assigning treatments
habitat improvement unitso
Treatments will be as follows:

to specific

Randomized
Block Number

Habitat Improvement
Unit Number

Treatment

1

1

Burn

2

Spray

3

Chain

4

Control

5

Spray

6

Control

7

Burn

8

Chain

2

Habitat improvement units were spaced as far apart as possible
minimize the effects that post treatment increases in deer and
suIting from animal preference to a particular unit would have
units.
The maximum spacing distance between units was limited
of the continuous oak type in the vicinity of the study area.
terval is 1/8 mile between units.

in order to
elk use reon adjacent
by the extent
The spacing in-

Each habitat improvement unit encompasses approximately 60 acres.
A 30 acre
unit was established in the center of each of the units that are slated for
treatment.
It is this 30 acre center portion of each unit that will be treatedo
The outside 30 acres will be left for wildlife covero No 30 acre units were
established in the centers of the 2 control unit~.
The shape of habitat improvement units is irregular and not uniform due to the
necessity of following the edges of the oak type when laying out unit boundarieso
Interior and exterior boundaries of habitat improvement units were laid out with
a K &amp; E compass set to allow for 14 degrees east declination, Fig. 1., and steel
tape. Orange steel fence posts were placed at each boundary turning point, and at

�- 118 -

intervals of 200 to 400 feet between turning points.
Boundary posts were
marked with numbered metal tags. Each tag bears the number of the habitat
improvement unit, and "A" if it is an outside boundary post~ or a "B" if it
is an inside boundary post, and the post number.
Both the outside and
inside boundary posts are numbered consecutively counterclockwise
around the
boundary (Fig. 2). Strips of fluorescent orange plastic surveyor's flagging
were tied to tree branches Jat..
approximately 50 foot intervals between boundary
posts.
When measuring distances no allowance was made for slope. The tape
was always pulled tight.

Pellet

Plot Establishment

Eighty temporary .001 acre circular pellet plots were sampled in order to
obtain a preliminary sample upon which to base sample size estimates.
Relative pellet sampling precision with various numbers of plots per habitat
improvement unit was computed, and considered. along with manpower requirements
in determining the number of pellet plots to establish per unit.
Sixty .002 acre circular pellet plots were subsequently established in each
of the 8 habitat improvement units.
Plots were located using the same
surveying techniques described in the previous section on habitat improvement unit establishment.
Pellet plots were established along randomly located lines which stretch
the entire width of the 60 acre unit.
Plot lines were laid out parallel
to each other.
One side of the 60 acre unit was used as a base line, and
the starting point of each pellet plot line was established along the base
line and marked with a pellet plot stake.
The distance between the starting
point stake and the first pellet plot on the line was selected at random
and varies from 0 to 100 feet. The distance between each pellet plot on a
line is 100 feet. Detailed maps were made of each habitat improvement unit
showing the exact location of each pellet plot.
Pellet plots were marked with orange steel stakes labeled with numbered
metal tags. Each tag bears the designation lip" indicating it is a pellet
plot, the line number, and the plot number.
Strips of blue plastic surveyor's
flagging were tied to the stake and to an overhanging tree branch near the
stake (Fig. 3).

Vegetation

Measurement

Methods

Three vegetation
measurement methods were considered
study.
These were the weight estimate, point quadrat
methods.

for use during
and dry weight

the
rank

The vegetation measurement technique selected involves the use of a meter
square sampling unit in which green weights of all grasses and forbs rooted
inside the plot, and of portions of all browse species which occur or
protrude into the plot up to a height of five feet above ground level will
be recorded.
Each forb and grass species will be clipped and weighed by
species to the nearest gram. The green weights of browse species will be

�- 119 -

estimated using a variation
Shoop and McIlvain (1963).

of the weight

estimate

technique

described

by

Green weights of all plant species encountered on meter square vegetative
measurement plots will be converted to air dry weights for two reasons:
(1) a great deal of variation in moisture content exists among species,
(2) moisture contents of individual species change drastically during the
summer as plants dry out and mature.
Herbage samples of various browse,
grass and forb species were collected periodically during the summer of
1969, and percentage moisture content determined.
A method was devised
for converting green weights of all plant species on the area to air dry
weights.
Samples of certain plant species will be collected at 10 day
intervals throughout each vegetative measurement period, and percentage
moisture contents will be determined.
These data will be used to convert
green weights of plants encountered on meter square plots to air dry weights.
The vegetation measurement method calls for a ten day pre-measurement
orientation period in order to train field personnel in the use of the method,
and to permit them to become proficient in estimating browse weights.
Since
several observers will be required to complete vegetation measurements a
mathematical procedure was devised for adjusting weight estimate data to
correct for observer error.
A detailed field manual describing the technique and mathematical procedures
for correcting for observer weight estimating error, and for converting
green weights to air dry weights was prepared and will be distributed to
field personnel who will assist in the study.
Eighty temporary meter square vegetation measurement plots were sampled in
order to obtain a preliminary sample upon which to base sample size estimates.

Plant Collection

and Identification

Three specimens of each species of plant found on the study area were collected, mounted and identified.
Facilities of the Colorado State University
Herbarium were used in making plant identifications, and identifications
were checked and annotated by C.S.U. Botany Department personnel.
One set
of specimens was contributed to the C.S.U. Herbarium, one set deposited in
the Herbarium at the Game Research Center, and one set used for field
reference.
Color photographs were taken of each species when collected,
and these will also be used for field reference.

Literature

Review

A review of literature to assemble information on treatment methods and
effects of treatment on the range, wildlife and livestock, optimum treatment
dates, seeding methods and rates, and the most desirable species for seeding
was initiated, and will be continued during the next project segment.

�- 120 -

RESULTS

AND DISCUSSION

Since this was the first segment of this study all of the work was directed
toward establishing habitat improvement unit boundaries, prellininary sampling
to determine the number of study plots needed for statistical reliablity,
establishment of sampling plots, plant collection, and literature review.
This has been described under the "Methods and Materials" section.
Thus, the
study has not yet reached the stage where findings are being made or results
obtained.
The only results discussed here are those related to preliminary
sampling for sample size determination, and plant identification.

Pellet Plot Establishment
Game use on the study area during the winter of 1969 was estimated at 23
deer days and 25 elk days use per acre on the basis of projections from
preliminary pellet plots used for sample size determination.
These use
figures are believed to be substantially below average for the area, because
snowfall in the area was exceptionally heavy that winter, and most of the
deer and elk moved to lower elevations.
Sixty circular pellet plots were subsequently established in each habitat
improvement unit, and plot size was increased from .001 to .002 acres in
order to increase the amount of area covered without substantially increasing
the cost of measuring plots.
With 60 .002 acre plots, deer and elk use
increases to 33 deer days and 36 elk days use per acre resulting from treatment can be detected as significant at the 95 percent confidence level.

Vegetation

Measurement

Plots

Seventy five vegetation measurement plots will be established in each
habitat improvement unit during the summer of 1970. Sample size computations
based on 80 preliminary meter square plots read during 1969 indicate 75 plots
will permit detection of changes resulting from treatment of 10 percent in
all vegetation, 15 percent in browse, 25 percent in forbs and 25 percent in
grasses, as significant at the 95 percent confidence level.

Plant Collection
A list of plant species collected
1969 is shown in Table I.

and Identification
on the study area during

the summer of

�- 121 -

LITERATURE CITED

Harrington, H. D. 1954.
Denver. 666pp.

Manual of the plants of Colorado.

Sage Books,

Kelsey, H. P., and W. A. Dayton. 1942. Standardized plant names.
McFarland Co., Harrisburg, Pa. 675pp.

J. Horace

Shoop, M. C., and E. H. McIlvain. 1963. The micro.-unit forage i.nventory
method. J. Range Mgmt. 16(4):172-179.

Prepared

by

~{h~

C. J.&amp;fdd

Roland C. Kufeld
Wildlife Researcher

�- 122 -

Table 1. Plants collected on the Hightower
during the summer of 1969.

Scientific

Name

11

Mountain

oak control

Common Name

study area

l:..1

FORBS
Achillea lanulosa Nutt.
*Agastache urticifolia Kuntz
Allium acuminatum Hook.
Aguilegia coerulea James
Ara~is drummondi Gray
Artemisia biennis Willd.
Artemisia dracunculus L.
Artemisia ludoviciana Nutt.
Aster adscendens Lindi.
Aster rubrotinctus Blake
Astragalus haydenianus A. Gray
Balsamorhiza sagittata (Pursh) Nutt.
Berberis repens Lindl.
Calochortus gunnisonii Watson
Capsella bursa-pastoris
(L.) medic.
Castilleja linariaefolia Benth.
Chenopodium berlandieri Moq.
*Cirsium canescens (Nutt.)
Delphinium nelsoni Greene
Delphinium occidentale (Wats.) Wats.
*Descurainia richardsonii (Sweet) O.E.Schultz
Disporum trachycarpum (S. Wats.) Benth.&amp;Hook
Erigeron flagellaris A. Gray
Erigeron formosissirnum Greene
Erigeron speciosus (Lindl.) DC
Eriogonum umbellatum Torr.
Epilobium paniculatum Nutt.
Fragaria americana (Porter) Britt.
Galium boreale L.
Gentiana heterosepala Engelm.
*Geranium parryi (Engelmo) Heller
Gilia aggregata (Pursh) Spreng.
Grindelia aphanactis Rydb.
Hackelia .leptophylla (Rydb.) Johnston
Helenium hoopesii A. Gray
Hydrophyllum fendleri (Gray) Heller
Iris missouriensis Nutt.
Lactuca scariola L.
Lappula redowskii (Hornemo) Greene
Lathyrus leucanthus (Rydbo)

Western Yarrow
Nettleleaf Giant Hyssop
Tapertip Onion
Colorado Columbine
Drummond Rockcress
Biennial Wormwood
Tarragon
Louisiana Sagebrush
Common Aster
Aster
Hayden Poisonvetch
Arrowleaf Balsamroot
Creeping Mahonia
Gunnison Mariposa
Shepherds Purse
Wyoming Paintedcup
Pit seed Goosefoot
Thistle
Minzies Larkspur
Duncecap Larkspur
Richardson Tanseymustard
Wartberry Fairybells
Trailing Fleabane
Fleabane
Oregon Fleabane
Sulfur Eriogonum
Autumn Willowweed
American Strawberry
Northern Bedstraw
Annual Gentian
Parry Geranium
Skyrocket Gilia
Gumweed
Stickseed
Orange Sneezeweed
Fendler Waterleaf
Rocky Mountain Iris
Prickly Lettuce
Stickseed
Aspen Peavine

�- 123 -

Table 1. Plants collected on the Hightower
during the summer of 1969, continued.

Scientific

Name 1/

FORBS,

Mountain

oak control

study area

Common Name

?J

continued

Lepidium densiflorum Schrad.
Ligusticum filicinum S. Wats.
Ligusticum porteri Coult. &amp; Rose
Linum lewisii Pursh
Lupinus argentues Pursh
Mertensia franciscana Heller
Moldavica parviflora (Nutt.) Britt.
*Musineon divaricatum (Pursh) Nutt.
Oenothera caespitosa Nutt.
"cOrthocarpus luteus Nutt.
Osmorhiza chilensis Hook. &amp; Arn.
Osmorhiza occidentalis (Nutt.) Torr.
Pachistima myrsinites (Pursh) Raf.
Pedicularis grayi A. Nels.
Penstemon
crandallii A. Nels.
Penstemon strictus Benth.
Phacelia heterophylla Pursh
Phlox longifolia Nutt.
Polemonium foliosissimum A. Gray
Potentilla pulcherrima Lehm.
Rudbeckia montana Gray
Senecio ambrosioides Rydb.
Senecio integerrimus Nutt.
Senecio serra Hook.
Smilacina stellata (L.) Desf.
Solidago sparsiflora A. Gray
Stellaria jamesiana Torr.
",Stellaria longifolia Muhl.
Swertia radiata (Kellogg) Kuntze
Taraxacum officinale Wiggars
"cThalictrum fendleri Engelm.
Tragopogon dubium Scop.
Urtica dioica L. var. procera (Muhl.) Wedd.
Valeriana occidentalis Heller
Veronica americana (Schwein.)
Viguiera multiflora (Nutt.) Blake
Viola rugulosa Greene
Wyethia amplexicaulis
(Nutt.) Nutt.

Prairie Pepperwood
Privet
Porter Ligusticum
Lewis Flax
Silvery Lupine
Franciscan Bluebells
Dragonhead
Musineon
Tufted Eveningprimrose
Yellow Owlclover
Sweetroot
Sweetanise
Myrtle Pachistima
Grays Pedicularis
Crandall Penstemon
Rocky Mountain Penstemon
Varileaf Phacelia
Longleaf Phlox
LeafyPolemonium
Beauty Cinquefoil
Coneflower
Ragweed Groundsel
Lambstongue Groundsel
Butterweed Groundsel
Starry Solomonplume
Threenerve Goldenrod
Chickweed
Chickweed
Showy Frasera
Common Dandelion
Fendler Meadowrue
Salsify
Big Sting Nettle
Western Valerian
American Speedwell
Showy Goldeneye
Cheyenne Violet
Mulesears Wyethia

-----------------------------------------------------------------------------

�- 124 -

Table 1. Plants collected on the Hightower Mountain oak control study area
during the summer of 1969, continued.
Scientific Name !I

Connnon Name ?:...I

GRASSES
Agropyron Smithi Rydb.
Agropyron trachycaulum (Link) Malte.
Bromus anomalus Rupro
Bromus inermis Leyss.
Dactylus glomerata L.
Elymus glaucus Buckl.
Festuca thurberi Vasey
Hordeum brachyantherum Nevski
Juncus balticus Willd.
Koeleria cristata (L.) Pers.
Orzyopsis hymenoides (R. &amp; S.) Ricker
Phleum pratense L.
Poa ampla Merr.
Poa canbyi (Scribn.) Piper
Poa pratensis L.
Sitanion hystrix (Nutt.) J. G. Smith
Stipa columbiana Macoun.
Stipa lettermani Vasey

Bluestem Wheatgrass
Slender Wheatgrass
Nodding Brome
Smooth Brome
Orchardgrass
Blue Wildrye
Thurber Fescue
Meadow Barley
Baltic Rush
Prairie Junegrass
Indian Ricegrass
Timothy
Big Bluegrass
Canby Bluegrass
Kentucky Bluegrass
Bottlebrush Squirreltail
Subalpine Needlegrass
Letterman Needegrass

BROWSE
Amelanchier alnifolia Nutt.
*Artemisia cana Pursh
Artemisia tridentata Nutt.
Ceanothus fendleri Gray
Cercocarpus montanus Raf.
*Chrysothamnus depressus Nutt.
*Chrysothamnus nauseosus (Pallas) Britt
*Crataegus erythropoda Ashe
Lonicera involucrata (Rich.) Banks
~'(Populustremuloides Mich.
Potentilla fruiticosa L.
Prunus virginiana L.
Purshia tridentata (Pursh) DC.
Quercus gambellii Nutt.
Rhamnus smithi Greene
Ribes cere~un Dougl.
*Rosa nutkana Presl.
Symphoricarpos sp.

Saskatoon Serviceberry
Silver Sagebrush
Big Sagebrush
Fendler Ceanothus
True Mountain Mahogany
Dwarf Rabbitbrush
Rubber Rabbitbrush
Cerro I-lawthorn
Bearberry Honeysuckle
Aspen
Bush Cinquefoil
Connnon Chokecherry
Antelope Bitterbrush
Gambel Oak
Smith Buckthorn
Wax Gooseberry
Nootka Rose
Snowberry

1/ Scientific names are from Harrington (1954).

II Common names are from Kelsey and Dayton (1942).
* Specimens of all plants except those prefixed by * have been annotated
by Dr. H. D. Harrington, Colorado State University Herbarium.

�- 125 -

Fig. 1.

Surveying

unit boundaries

and plot locations.

Fig. 2. Orange steel fence posts were placed at each boundary
point and at 200 to 400 foot intervals between turning points.

turning

�- 126 -

Fig. 3.
numbered

Pellet plots were marked with orange steel stakes to which
metal tags and strips of blue surveyor's flagging were attached.

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                  <text>July, 1970

- 127

JOB FINAL REPORT

State of

COLORADO

Project No.

W-38-R-24

Deer-Elk

Work Plan No.

1

Job No.

Job Title

Experimental

Trapping

Period Covered:

None

Personnel:

Raymond J. Boyd

Investigations

and Marking

5

Techniques

ABSTRACT

A final report was submitted on this job in the July, 1969 (Part One)
Game Research Report.
It should not have been included in the W-38-R-24
Project Segment job descriptions.

��July, 1970

- 129 -

JOB PROGRESS REPORT

State of

~C~O~L~O~RA~D~O~

_

Project No.

W-38-R-24

Deer-Elk

Work Plan No.

2

Job No.

Investigations
7A

Job Title

White River Elk Study - Publication ·of Results

Period Covered:

April 1, 1969 through March 31, 1970

Personnel:

Raymond J. Boyd

ABSTRACT

A "final" draft of the manuscript was completed and bids were obtained
for printing as a Division Technical Bulletin.
The rpinting contract
had not been awarded at the time this report was prepared.

��- 131 -

WHITE RIVER ELK STUDY

- PUBLICATION

OF RESULTS

Raymond J. Boyd

P. S. OBJECTIVE
Formulate

a management

plan for the elk and their habitat

SeGMENT
1.

Publish all the research
study.

1.

Publish as a Colorado

OBJECTIVE

findings

METHODS

on the White River.

resulting

from the White

River elk

AND MATERIALS

Game, Fish and Parks Division

Technical

Bulletin.

RESULTS AND DISCUSSION
A final draft of the manuscript, complete with figures, photographs and
tables was completed and edited.
This manuscript was taken to several
printing firms and bids secured for printing.
At the time this report was
prepared, no contract for printing had been let. It is anticipated that the
printing will be completed by August 15, 1970, at which time distribution
of the publication can begin.

��July,

1970

- 133 -

JOB FINAL REPORT

S ta te of _~,.;.-_.::..CO.::..L:::.O.:.:RA=:.:D~O::..-._
Project No.

W-38-R-24

Deer-Elk

Investigations

Work Plan No.

2

Job No.

8

Job Title

Hunter Harvest

Surveys

Period Covered:

April 1, 1969 through March

Personnel:

Raymond J. Boyd, Velma Merkle, Velma Fredrickson,
Mary Lu Selch, Pat Ruqua, Sara Johnson, Robert Tully,
Howard Funk and Jack Grieb.

31, 1970

ABSTRACT

Six different categories of 1969 elk license holders (either sex,
specified, antlerless, archery, regular bull and Sportsman's) were
randomly surveyed in order to determine various harvest information.
A total of 22,790 questionnaires were sent out to a portion of the
estimated 78,202 persons who purchased an elk license in 1969 (59,683
resident and 18,519 nonresidents).
The return from the sample totaled
17,091 for an overall return of 74.99 percent.
An estimated harvest
of 17,581 elk (11,204 bulls, 5,370 cows and 1,007 calves) was determined
for the 1969 elk season.
This harvest was accomplished by an estimated
66,791 hunters (49,308 residents and 17,483 nonresidents).
Data
collected from Sportsman's License holders also indicated that out of
an estimated 15,691 purchasers (11,422 residents and 4,269 nonresidents),
7,979 hunted one or more species of small game and 11,533 fished.
As a result of this random survey, a manual was prepared showing sample
sizes required, methods of drawing the sample, recorrnnended questionnaires
and informational letters to be sent out and suggested IBM punch-card
layouts for computer analysis of the questionnaire data.

�- 134 -

RECO'MMENDATIONS

1.

Since hunter report card returns are getting smaller each year,
there is reason to suspect that harvest and pressure information
generated by these cards is in error.
Therefore it is recommended
that all future big game harvest data be determined by random
surveys of all big game license.holders.
From aptoperly
designed
sample, far better quality information can be obtained.

2.

Complete instructions,sample
sizes, calculations,
IBM card lay~
outs and projection methods are detailed in the manual prepared as
part of this job.
This manual should be followed in all future
random surveys of big game license holders.

�- 135 -

HUNTER HARVEST

SURVEYS

Ra ymond J. Boyd

The purpose of this investigation is to develop a reliable post-season
mail survey, suited to our facilities, for estimating the 1969 elk harvest, hunter success and other related information needed by game managers.
Results of this study will serve as background for random surveys of other
game species hunted in Colorado.

Po So OBJECTIVE
To estimate the annual State harvest of all big game species, sex, age,
game management unit, county, hunter residence and type of license, season,
period of season and to obtain information on man-days of recreation involved.

METHODS AND MATERIALS
Since the final result of this investigation will be a procedure manual for
conducting a random survey of all elk license holders, no details of the
methods used will be presented in this report.
The reader is referred to
the procedure manual for details on methods used in this survey.

RESULTS AND DISCUSSION
The procedure manual developed by this job will be used as a guide by the
Division in all future random surveys for big game harvest statistics.
The Colorado Division of Game, Fish and Parks has, within its big game
license framework, six different kinds of elk licenses available to the
hunting public.
These six licenses are further broken down into two additional categories as there is a different license for residents and nonresidents.
In effect, then, 12 license types were sampled in this survey.
Table 1 indicates the license types surveyed, the number of questionnaires
sent, number of questionnaires returned before and after a follow-up letter
and the percent return on each type of questionnaire.

�- l36 -

Table

1.

Numbers

of questionnaires

sent, returned

and percent return,

Number of
Questionnaires
Sent Out

Number Returned
Before Fo11owup Letter

Number Returned
After Followup Letter

Percent
Return

Type
of
Survey

19690

Either-sex

License

147

104

10

77 .6

Specified

License

2,816

2,030

217

7908

Antlerless

License

6,254

4,815

448

8402

Archery

Elk License

1,102

692

93

7102

Regular

Bull License

6,223

3,352

589

6303

Big Game

3,090

2,131

170

74.5

Small Game

3,158

2,118

322

77.3

Total

22,790

15,242

1,849

74.9

Sportsman's

License:

Certain basic data had to be developed for each license category before the
total elk harvest and other related management information could be developed.
For each of the six license types it was necessary to determine:
sent out.

1.

Number

of questionnaires

20

Number

of usable cards returned

3.

Number

of reported

4.

Number

of each license type sold to residents

5.

Number of elk kills reported

and percent return.

no hunts by residents

and nonresidentso

by residents

and nonresidents.

and nonresidents.

From the above information, hunter success, total number of participating
hunters and total harvest could be calculated.
This information is shown
for each of the six license types as follows:
Regular

Bull License

sent out

= 6,2230

I.

Questionnaires

2.

Usable cards returned

3.

Reported

no hunts

= 3,940

(63 3% return).
0

= 287 (270 residents,

17 nonresidents).

�- 137 4.

Licenses

sold

5.

Kills reported

= 45,867 (34,750 residents,

11,117 nonresidents).

= 727 (474 resident kills, 253 nonresident

kills).

Success
Nonresident

Resident

253 = nonresident kill
1,016 = nonresident hunters
253 + 1,016 = .24901% success

474 = resident kill
2,712 =res ident hunters
474 + 2,712 = .17477% success
Number of Participating

Hunters
Nonresident

Resident
2,712
2,982

x

1,016
1,033

34,750

x = 31,604 resident

hunters

Total participating

regular bull elk hunters

x
11,117

x = 10,934 nonresidents
=

42,538

Total Harvest
Resident

Nonresident

31,604 X .17477

= 5,523 bulls
Antlerless

10,934 X .24901

= 2,723 bulls

Elk License

In 1969, there were 53 areas in the State set up with limited numbers of
antlerless elk licenses.
The antlerless elk harvest, hunter success and
number of participating hunters was determined for each area.
Rather than
show calculations of this information for each area, one example was chosen
as representative of all antlerless areas.
Area Q (Game Management

Units 77 and 78; see map in procedure

1.

QUestionnaires

2.

Usable cards returned

3.

Reported

4.

Licenses

5.

sent out

no hunts

manual)

400.

= 341 (85.2% return).

= 15 (13 residents,

2 nonresidents).

= 1,599 (1,078 residents, 521 nonresidents).
Kills reported = 244 (reSidents - 116 cows and 24 calves; nonresidents
sold

85 cows and 19 calves).
Success
Resident
140
resident kill
200 = resident hunters
140 + 200 = .70000% success

Nonresident
104 = nonresident kill
126 = nonresident hunters
104 + 126 = .82539% success

-

�- 138 -

Number of Participating Hunters
Resident

Nonresident

200 =
X
213
1,078

126
X
128 - 521

X = 1,012 resident hunters

X = 513 nonresident hunters

Total participating ant1er1ess elk hunters in Area Q = 1,525

0

Tota 1 Harvest
Nonresident

Resident
1,012 X .70000 = 708 elk
(587 cows and 121 calves)

513 X .82539 = 423 elk
(346 cows and 77 calves)

Either-sex Elk License
There was only one area in the State in 1969 that was set up as an eithersex elk hunt area.
1.
2.
3
4.
5.
0

Questionnaires sent out = 148.
Usable cards returned = 114 (77.0% return).
Reported no hunts = 7 (6 residents and 1 nonresident).
Licenses sold = 148 (106 residents and 42 nonresidents).
Kills reported = 12 (residents - 4 bulls, 4 cows, no calves;
nonresidents - 3 bulls, 1 cow, no calves).

Success
Resident
8 = resident kill
82 = resident hunters
8 + 82 = .09757% success
Number of Participating Hunters
Resident
82 = ..1L
88
106
X = 99 resident hunters

Nonresident
4 = nonresident kill
25 = nonresident hunters
4 + 25 = .16000% success
Nonresident
25 = L
26
42
X = 40 nonresident hunters

Total participating either-sex elk hunters = 139
Total Harvest
Resident
99 X .09756 = 10 elk
(5 bulls, 5 cows, no calves)

Nonresident
40 X .16000 = 6 elk
(4 bulls, 2 co;s, no calves)

�- 139 -

Specified

Elk License

There were 10 areas in the State in 1969 that were set up under the
specified elk license regulation.
Area E, on the White River, harvest
data is shown below as an example of harvest calculations for this type
of license.
Questionnaires

2.

Usable cards returned

3.

Reported

4.

= 3,109 (2,417 residents and 692 nonresidents).
Kills reported = 444 (residents - 136 bulls, 141 cows and 30 calves;
Licenses

5.

sent out

= 1,517.

1.

= 1,190 (78.4% return).

no hunts = 161 (142 residents

and 19 nonresidents).

sold

nonresidents

- 86 bulls, 48 cows and 3 calves).

Success
Resident

Nonresident

307 = resident kill
759 = resident hunters
307 + 759 = .40447% success
Number of Participating
Resident

137 = nonresident kill
270 = nonresident hunters
137 + 270 = .50740% success

Hunters
Nonresident

759 = ----"X~
901
2,417

270
289

2,036 resident

hunters

Total participating

hunters

X

= X
692

X = 646 nonresident

hunters

= 2,682

Tota 1 Harvest
Resident

Nonresident

2,036 X .40447 = 824 elk
(366 bulls, 378 cows, 80 calves)

646 X .50740 = 328 elk
(206 bulls, 115 cows, 7 calves)

Archery

Elk License

sent out

= 1,102.

I.

Questionnaires

2.

Usable cards returned

3.

Reported

no hunts

4.

Licens~s

sold = 1,254 (873 residents

5.

Kills reported

= 793 (71.9% return).

= 43 (34 residents

= 37 (26 residents

and 9 nonresidents).
and 381 nonresidents).

and 11 nonresidents).

Success
Resident
26
resident kill
523
resident hunters
26 + 523 = .04971% success

Nonresident
11 = nonresident kill
227 = nonresident hunters
11 + 227 = .04845% success

�- 140 Number of Participating Hunters
Nonresident

Resident
X

523 =
566

227 = ~
270
381

873

x = 320 nonresident hunters
Total participating archery elk hunters = 1,127

x = 807 resident hunters
Total Harvest

Nonresident

Resident
807 X .04971 = 42 bull elk

320 X .04845 = 14 bull elk

Sportsman's License
The Sportsman's License in Colorado enables a hunter to legally take an elk,
a deer, a lion and a bear, plus small game and fish. While this job was
set up to determine only the elk harvest for 1969, it was not practical to
survey Sportsman's License holders only on their elk hunting success. Therefore, the Sportsman's License survey was designed to determine harvest levels
on all species of game and fish that could legally be taken on the Sportsman's
License. As an example, however, the calculations below will show only how
many Sportsman's License holders actually hunted elk in 1969.
1. Questionnaires sent out = 3,090.
2. Usable cards returned = 2,301 (74.5% return).
3. Reported no hunts = 291 (265 residents and 26 nonresidents).
4. Licenses sold = 15,691 (11,422 resident and 4,269 nonresident).
Number of Participating Hunters
Resident
1,828 = ---..,;X~_
2,093
11,422
X = 9,976 resident elk hunters

179
205

Nonresident
X
4,269

X = 3,728 nonresident elk hunters

Total participating Sportsman's License holders = 13,704 hunters
Harvest St~tistics - 1969 Big Game Season
The following tables list all harvest data for the various elk license
categories for 1969 in Colorado, plus additional harvest data on deer, bear,
lion, small game and fishing that was determined for the Sportsman's License.
All of these data were determined from the random survey detailed in this

Prepared by

�- 141 -

Table 2

. Total elk harvest by Game Management Unit, 1969. !./

Game Management
Name
Unit No.
2
4
5
6
7
8
11
12
13
14
15
16
17
18
19
20
22
23
24
25
26
27
28
29
31
32
33
34
35
36
37
38
39
40
41
411
42
43
44
45
46

Brown's Park
Elk Head
Snake River
Cowdrey
Laramie River
Red Feather
Strawberry Creek
Williams Fork
Dunkley
Elk River
Yampa
West Side
Owl Mountain
Troublesome
Poudre River
Estes Park
Piceance
Miller Creek
White River
Sweetwater
Toponas
Sheephorn
Hot Sulphur
Boulder
Roan Creek
Parachut;e Creek
'Ziflc
'.:oloradoRiver
Castle Peak
Piney
Summi,t
Central City
Nt. Evans
Glade Park
Kannah Cr.
Surface Cr.
Grand Mesa
Roaring Fork
Brush Creek
Holy Cross
Bailey

Bulls

Cows

Calves

Total

48
281
158
101
36
14
87
220
385
330
167
22
50
210
70
206
32
212
382
80
103
36
68

34

1

87
39
8
55
27
55
62
208
34
10
15
153
27
59
5
200
293
43
47
16
93
17

26
5
8
9
2
8
7
34
2
1
6
24
5
13

83
281
271
145
52
78
116
283
454
572
203
33

35
52
6
5
31
7

14
108
22
14
14
56
51
22
42
14
195
177
235
24
9

2
30
31

3
4

29
1
56
17

9
0
8
2

55
90
126
48
8

7
19
15
5
1

71

387
102
278
32
447
727
129
150
57
192
24
14
2
141
57
14
14
94
1
115
41
42
14
257
286
376
77
18

-------------------------------------------------------------------------------

�- 142 -

Table 2.

Total elk harvest by Game Management Unit, 1969 (continued). 1/

Game Management
Unit No.
Name
461
47
48
49
50
52
53
54
55
56
58
59
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
90
Totals

Deer Creek
Frying Pan
Mt. Elbert
Buffalo Peaks
South Park
N. F. Gunnison
Coal Cr.
Sapinero
Taylor ~iver
Buena Vista
Eleven Mile
Pikes Peak
W. S. Uncompahgre
E. S. Uncompahgre
Smith Fork
Olathe
Alpine
L. F. Gunnison
Tomichi
Saguache
Grape Cr.
San Miguel
Dolores
Yellow Jacket
Mancos
Hermosa
Animas River
Creede
Piedra
San Juan
La Garita
S. F. Rio Grande
Conejos
Sand Dunes
Trinchera
Greenhorn
Picket Wire
Sangre de Cristo

Bulls

Cows

Calves

16
175
92
51
43
276
66
387
478
110
13

115
53
7
52
77
13
138
293
4
10

19
17
2
14
10
30
33
1

5

82
191
8
'14
232
139
III

13
9
5

2

67
49
51
53

6
10
12

269
14
185
333
28
66
444
644
196
560
830
187
255
218
30
8
22
91
50

59
123

10
17

19
261
298
188
475
458
51
193
125

5
29
75
64
107
91
6
48
32

7
7
12

7

11,204

5,370

1,007

Total

16
309
162
60
109
363
79
555
804
114
24
5
97
200
13
14
299
194
172
334
14
254
473
28
90
734
1,017
448
1,142
1,379
244
496
375
30
8
36
98
62
17,581

1/
- Includes harvest on Regular, Sportsman's, Specified and Antler1ess licenses
only.

�- 143 -

Table
Unit,

3 • Elk harvest by resident
1969. ]j
Unit

and nonresident

Bulls
NR

No.

Name

R

2
4
5
6
7
8
11
12
13
14
15
16
17
18
19
20
22
23
24
25
26
27
28
29
31
32
33
34
35
36
37
38
39
40
41
411
42
43
44
45
46
461

Browns Park
Elk Head
Snake River
Cowdrey
Laramie River
Red Feather
Strawberry Cr.
Williams Fork
Dunkley
Elk River
Yampa
West Side
Owl Hountain
Troublesome
Poudre River
Estes Park
Piceance
Miller Cr.
White River
Sweetwater
Toponas
Sheephorn
Hot Sulphur
Boulder
Roan Cr.
Parachute Cr.
Rifle
Colorado River
Castle Peak
Piney
Summit
Central City
Mt. Evans
Glade Park
Kannah Cr.
Surface Cr.
Grand Mesa
Roaring Fork
Brush Creek
Holy Cross
Bailey
Deer Creek

44
192
70
50
36
14
14
129
251
263
131
22
50
210
58
206
22
132
256
44
28
36
56
0
14
0
59
22
14
14
56
0
48
18
42
14
146
101
148
19
9
16

4
89
88
51
0
0
73
91
134
67
36
0
0
0
12
0
5
80
126
36
75
0
12
0
0
0
49
0
0
0
0
0
3
4
0
0
49
76
87
5
0
0

hunters

Cows

by Game Management

R

NR

R

Calves
NR

R

33
0
69
27
8
50
11
37
51
158
27
6
14
144
25
58
3
159
219
29
32
16
88
17
0
2
22
28
0
0
24
1
50
17
0
0
43
65
120
46
8
0

1
0
18
12
0
5
16
18
11
50
7
4
1
9
2
1
2
41
74
14
15
0
5
0
0
0
8
3
0
0
5
0
6
0
0
0
12
25
6
2
0
0

1
0
24
0
8
6
2
6
7
27
2
1
5
24
5
13
0
33
47
6
0
5
28
7
0
0
3
4
0
0
6
0
8
2
0
0
5
16
15
4
1
0

78
192
163
77
52
70
27
172
309
448
160
29
69
378
88
277
15
324
522
79
60
57
172
24
14
2
84
54
14
14
86
1
106
37
42
14
194
182
283
69
18
16

0
0
2
5
0
3
0
2
0
7
0
0
1
0
0
0
0
2
5
0
0
0
3
0
0
0
0
0
0
0
3
0
0
0
0
0
3
3
0
1
0
0

Total
NR

5
89
108
68
0
8
89
111
145
124
43
4
2
9
14
1
7
123
205
50
90
0
20
0
0
0
57
3
0
0
8
0
9
4
0
0
63
104
93
8
0
0

--------------------------------------------------------------------------------------

Total
Harvest

83
281
271
145
52
78
116
283
454
572
203
33
71
387
102
278
32
447
727
129
150
57
192
24
14
2
141
57
14
14
94
1
115
41
42
14
257
286
376
77
18
16

�- 144 Table 3 • Elk harvest by resident and nonresident hunters by Game Management
Unit, 1969 (continued). 11
Unit
Name

No.
47
48
49
50
52
53
54
55
56
58
59
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
90

Frying Pan
Mt. Elbert
Buffalo Peaks
South Park
N. F. Gunnison
Coal Cr.
Sapinero
Taylor River
Buena Vista
Eleven Mile
Pikes Peak
W. S. Uncompahgre
E. S Unc ompa hgre
Smith Fork
Olathe
Alpine
L. F. Gunnison
Tomichi
Saguache
Grape Cr.
San Miguel
Dolores
Yellow Jacket
Mancos
Hermosa
Animas River
Creede
Piedra
San Juan
La Garita
S. F. Rio Grande
Conejos
Sand Dunes
Trinchera
Greenhorn
Picket Wire
Sangre de Cristo
0

Resident &amp; Nonresident
Sub-totals
Totals

11

Bulls
NR
R

Cows
NR
R

Calves
NR
R

163
78
51
37
185
16
308
374
72
13
0
70
100
8
14
72
51
73
243
14
98
170
28
28
225
359
121
164
272
149
205
153
30
8
22
63
36

104
53
7
44
60
10
96
244
4
10
5
10
8
2
0
50
41
38
50
0
46
99
0
14
203
219
164
326
261
49
183
107
0
0
7
5
12

13
17

12
14
0
6
91
50
78
103
38
0
0
12
91
··0

0
160
88
38
26
0
87
163
0
38
219
284
75
395
561
38
50
65
0
0
0
28
14

11
0
0
8
17
3
42
49
0
0
0
3
1
3
0
17
8
13
3
0
13
24
0
5
58
79
24
149
197
2
10
18
0
0
0
2
0

6
0
0
.0
1
0
3
14
0
0
0
0
0
0
0
0
2
1
2
0
4
0
0
0

2

14
9
0
27
19
0
1
0
2
0
0

..

o·

0
4
9
10
0
6
17
0
5
29
34
54
64
57
6
44
27
0
0
7
0

0

41
10
43
34
0
4
5
0
0
·0
0
0

0

R

Total
NR

280
148
60
9S
254
26
433
637
76
24
5
82
108
10
14
122
96
120
303
14
150
286
28
47
457
612
339
554
590
204
432
287
30
8
36
68
48

29
14
0
14
109
53
123
166
38
0
0
15
92
3
0
177
98
52
31
0
104
187
0
43
277
404
109
587
792
40
64
88
0
0
0
30
14

Total
Harvest

309
162
60
109
363
79
555
804
114
24
5
97
200
13
14
299
194
172
334
14
254
473
28
90
734
1,017
448
1,142
1,379
244
496
375
30
8
36
98
62

17,581
7,127

4,077

11,204

Harvest by archers not included.

4,238 1,132
5,370

798

209

1,007

12,163

5,418

17,581

17,581

17,581

�- 145 -

Table 4 •

1969 regular bull license harvest by Game Management Unit.

Unit
No.

Name

3
4
5
6
7
8
11
12
13
14
15
16
17
18
19
20
21
24
25
26
27
28
31
33
34
35
36
37
41
411
42
43
44
47
48
49
52
53
54

Craig
Elk Head
Snake River
Cowdrey
Laramie River
Red Feather
Strawberry Creek
Williams Fork
Dunkley
Elk River
Yampa
West Side
Owl Mountain
Troublesome
Poudre River
Estes Park
Douglas
White River
Sweetwater
Toponas
Sheephorn
Hot Sulphur
Roan Creek
Rifle
Colorado River
Castle Peak
Piney
Sununit
Kannah Creek
Surface Creek
Grand Mesa
Roaring Fork
Brush Creek
Frying Pan
Mt. Elbert
Buffalo Peaks
N. F. Gunnison
Coal Creek
Sapinero

Resident

169
70
42
28
14
14
98
212
183
84
14
42
154
42
212
14
14
28
28
28
56
14
28
14
14
14
56
42
14
98
70
140
140
70
28
169
254

Nonresident

Total

24
24
61
24

24
193
131
66
28
14
87
134
346
232
120
14
42
154
54
212
14
14
64
89
28
68
14
77
14
14
14
56
42
14
147
119
213
152
70
28
205
36
315

73
36
134
49
36

12

36
61
12
49

49
49
73
12

36
36
61

---------------------------------------------------------------------------------------

�- 146 -

Table 4 •

1969 regular bull license harvest by Game Management Unit (continued).

Unit
No.

Name

55
56
61
62
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
84
85
90

Taylor River
Buena Vista
Wo So Uncompahgre
Eo S. Uncompahgre
Olathe
Alpine
L. F. Gunnison
Tomichi
Saguache
Grape Creek
San Miguel
Dolores
Ye ll.ow Jacket
Mancos
Hermosa
Animas River
Creede
Piedra
San Juan
La Garita
So Fo Rio Grande
Conejos
Sand Dunes
Greenhorn
Picket Wire
Sangre de Cristo
Total

Resident

Nonresident

Total

310
56
70
84
14
56
28
42
212
14
98
154
28
28
183
310
198
140
254
126
197
112
14
14
42
28

73
24
12
36

383
80
82
120
14
129
89
66
224
14
171
276
28
52
329
481
259
518
632
150
233
136
14
14
66
28

5,523

73
61
24
12
73
122
24
146
171
61
378
378
24
36
24
24

2,723

8,246

�,

- 147-

Table

5.

1969 elk harvest on antlerless and specified licenses (by elk area) e

Area

Unit
Number

No. of
Licenses
Issued

Bull
Harvest
RegoLic.

A

14

400

B

12
13

B-1

Cows

Calves

Success
Ratio on
Permits

325

208

34

6005

567

199

220
385
605

55
62
117

8
_7_
15

66~3

283
454
737

11

50

87

27

2

58.0

116

C

15
1/2-27

100

167
18
185

32
14
46

2
1
3

4900

201
33
234

D

1/3- 6
1/2-16

96

34
11
45

9
8
17

3
0
3

20.8

46
19
65

E

23
24

200
293
493

35
52
87

3700

447
727
1,174

AO
Ant I

2,113
996

22
22

Elk Harvest on Ant10

&amp; SEecified Licenses
Bulls

212
360
572

Total
Harvest

F

1/2-27
36

52

18
14
32

4
0
4

4
0
4

1504

26
14
40

G

28
37

499

68
56
124

93
29
122

31
9
40

3205

192
94
286

H

42

121

195

55

7

51..2

257

I

44
1/2-47

401

235
88
323

126
56
182

1.5
13
28

5204

376
157
533

1-1

1/2-47

99

87

59

6

6506

152

J

54

300

382

138

30

56.0

550

K

55

498

473

293

33

65 5

799

L

65

100

232

67

0

67 0
0

299

M

67

100

111

51

10

61••
0

172

0

--------------------------------------------------------------------------------------

�- 148 -

Table 50 1969 elk harvest on antler1ess and specified licenses (by elk area)
(continued).

Area

Unit
Number

No. of
Licenses
Issued

Bull
Harvest
RegoLic.

M-1

68

131

N

70
71
73

0

Elk Harvest on Ant1.
Cows

Calves

Success
Ratio on
Permits

269

53

12

49.6

334

350

185
333
66
584

59
123
19
201

10
17
5
32

66.6

254
473
90
817

74

549

439

261

29

52.8

729

P

1/2-75

200

320

102

14

58.0

436

P-1

1/2-75

459

319

196

36

50.5

551

Q

77
78

1,599

555
825
1,380

475
458
933

107
91
198

70.7

1,l37
lz374
2,511

R

76

451

196

188

64

55.9

448

R-1

80

399

255

193

48

60.4

496

R-2

79

100

187

51

6

57.0

244

S

81

248

218

125

32

6303

375

1/3-57
69
84
90

80

2
0
6

33.8

2
14
35

20

0
0
7
_0_
7
5
13
18

46.4

75
278
353

30.5

123
0
123
75

S-l
&amp; SS

T

U

2/3-19
20

278

39 . AO
391 AntI

1/2-50

AO
Ant I

W

3/4-25
26

Bulls

14
22
50
86

227
150

113

31

21
59
80

8

51
_0_
51

56
0
56

8
0

34

33

8

41.2

30

6
0
6

41.5

60
103
163

-.!±L
77

Total
Harvest

--.E.L

8
...n....

102
80
200

..JL

41
183
224

8
V

&amp; S~ecified Licenses

8

--------------------------------------------------------------------------------------

96

150
246

�- 149 Table 50 1969 elk harvest on antler1ess and specified licenses (by elk area)
(continued).

Area

Unit
Number

Noo of
Licenses
Issued

Bull
Harvest

200

20
108
22
150

80
60

7

Reg Lf,c ,
i

Elk Harvest on AntI.

&amp; SEecified Licenses
Bulls

Success
Ratio on
Permits

Total
Harvest

Cows

Calves

13
30
31
74

0
3
4
7

40,,5

33
141
57
231

34

1

5403

83

X

1/4-25
33
34

Y

2

Z

8
1/3-19

320

14
21
35

55
6
61

9
0
9

21.9

78
27
105

AA

2/3- 6
7

275

67
36
103

30
8
38

2
8
10

17'04

99
52
151

BB

1/2-17

30

25

5

2

23.3

32

CC

43

198

177

90

19

5500

286

DD

46

8

1

12.7

18

EE

52

149

276

77

10

58.4

363

FF

61
62

50

82
191
273

13
9
22

2
0
2

48,,0

97
200
297

GG

85

148

82

9

7

0

10.8

98

HH

1/2-50

9

19

6

19,,6

34

JJ

48
1/3-56

53
4
57

17
0
17

5704

162
41
203

5
5
10

0
0
0

50 0

5
---5

87

26

45.2

271

153

24

33,,7

387

AO
AntI

AO
AntI

AO
AntI

9

99
43

92
81
129

41

92

JL
129

KK

1/4-58
1/2-59

20

LL

5

250

158

MM

18

599

185

25

0

10

---------------------------------------------------~----------------------------------

�- 150 -

Table 50 1969 elk harvest on antler1ess and specified licenses (by elk area)
(continued).

Area

Unit
Number

No. of
Licenses
Issued

Bull
Harvest
RegoLico

NN

66

100

00

53
63

pp

Elk Harvest on Ant10
Cows

Calves

Success
Ratio on
Permits

139

49

6

5500

194

49

66
8
74

11
7
18

0
0
0

3607

77
15
92

1/2-16
1/2-17

100

11
25
36

2
10
12

1
4
5

17.0

14
39
53

TT

49

50

51

7

2

18.0

60

UU

29
38
381

80

16
2
0
18

7
0
0
7

31.2

23
2
0
25

VV

3/4-58

AO
Ant1

20
10

8

5

1

46 7

14

WW

21
22
31
32

AO
Ant1

30
30

0
13
0
0
13

0
5
0
2
7

0
0
0
0
0

33 3

14
18
14
2
48

14
14
28

&amp; SEecified Licenses
Bulls

0

0

Total
Harvest

XX

40

\0
AO
Ant1 20

22

17

2

82 0

41

ZZ

45

AO
50
Ant1 100

24

48

5

51.3

77

0

�Table 6 . Resident elk hunting pressure showing county of residence and number of participating license holders
by Game Management Unit, 1969.
County of
Residence
Adams
Alamosa
Arapahoe
Archuleta
Baca
Bent
Boulder
Chaffee
Cheyenne
Clear Creek
Conejos
Costilla
Crowley
Custer
Delta
Denver
Dolores
Douglas
Eagle
Elbert
E1 Paso
Fremont
Garfield
Gilpin
Grand
Gunnison
Hinsdale
Huerfano
Jackson
Jefferson
Kiowa
Kit C.?XS'n'

2

4

5

6

7

8

1

66

58

45

20

49

5

26

22

24

4

6

40 103

95

26

46

Unit Number
16
14
15

12

13

2

87

139

88

1

42

112

119

30

13

112

40

66

11

40

17

18

19

20

2

40

134

1

41

2

27

114

15

16

15

56
13

2
59
5

94

267

13

13

22

13

23

24

25

26

88

153

75

36

53

93

18

33

3
62
1

4
109
1

13

45

2

2

4
13

2

1

~
~

VI

13
1
13

95 165

61

21

45

17

62

3
6

26

16

11

13
13

13

29

7

16

188

150 165

54

13

2

29

17
13
26

47

6

26

83

17

40

2

51

161

376

19

72

7

13

2

2

2

1

15
54

15

13
2

146

2

29

30

83
32

17

303

42
85

13
2
245

31

32

14
1
5

4
297
1

78

13
87

16

10

7

89

29

2

55

47

2

58
3
32

1
4
80
6
56

1

1

101

153

2

2

3

2

16

93
55

9

3
163
1
2
1

-------- ----- ------------------------------------------------------------------------------------------------------

�Table 6 • Resident elk hunting pressure showing county of residence and number of participating license holders
by Game Management Unit, 1969, continued.
County of
Residence
Lake
La Plata
Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Sunnnit
Teller
Washington
Weld
Yuma
Sportsman's

Total

27

28

29

31

32

13

2

1

33

1

26

14

15

34

Unit Number
35
36
37

26

40

38

39

39l

40

41

42

43

44

45

46

43
6

13

1

33

150

1

3

2
491

43

1

13

13
13

1
1

14

13
5

1

2

1

15

1
13
13

1

44

13

1

1

13

17
1

92
13

2
4
11

13

32

1

1

26

129

37

1

3

74

78

10

30

62

14

88

2

482 1284

335

18

60

503

379

102

231

524

98

417

13

59

.po

11
5
3

1

51

~

I.J1

18

19

14
43

148

203 1071

17
2
104

1
162

17

24

677 1160

197

216

-------------------------------------------------------------------------------------------------------------------~

�Table 6 . Resident elk hunting pressure showing county of residence and number of participating license holders
by Game Management Unit, 1969, continued.

County of
Residence
Adams
Alamosa
Arapahoe
Archuleta
Baca
Bent
Boulder
Chaffee
Cheyenne
Clear Creek
Conejos
Cos til1a
Crowley
Custer
Delta
Denver
Dolores
Douglas
Eagle
Elbert
El Paso
Fremont
Garfield
Gilpin
Grand
Gunnison
Hinsdale
Huerfano
Jackson
Jefferson
Kiowa
Kit Carson
Lake
La Plata

51

52

53

Unit Numbe r
56
55
54

47

48

49

50

17

15

18

31

27

19

77

7

28

18

56

47

53

22

13

27

15
1

3
126

13

46
26
4

2
20
61

57

58

59

61

62

63

64

13

13
84

13

1
13

13

1
2

13

65

66

67

27

16

1

2

56
13
57
1

1

2

14
60

4
1
1
78

33

84

4
62
47
1
76

57
75

77

7
54

13
53

13
7
162

7
26

2

69

15

14

13

41
21

16
59

13
62
2

14

36
13

194
47

66
18

98

5

30

2

13

l3

1

1

1

93
2

69
15

13

7
1
271

6

26

13
13

55
13
15

13

122
54

323
42
18

20

15

381

553

34
26

26
26

73
15

t05
13

l3

2

13
49

38

32

43

28

1

99

40

1

14

14
l3

37

82

54

15

-------------------------------------------------------------------------------------------------------------------~

1

t-'
V1
V1

�Table 6 • Resident elk hunting pressure showing county of residence and number of participating license holders
by Game Management Unit, 1969, continued.
County of
Residence
Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Surmnit
Teller
Washington
Weld
Yuma
Sportsman's

Total

47

48

49

50

13

2 .

13

2

14

2

1

1
184

6
2
15

1
5

51

52

53

42

54

Unit Number
55
56

57

61

62

15
13
13

13

13

15

41

142

26

89

124

17

14
3

58

59

63

64

65

66

67

13

13
1
1

13

13

1

46

3
1

306

104

1
13

27
74

13

t-'

\.n
0'1

234
3

48

13

2

1

1

8
59

148

28

13

1

13

45

16

177

1

13
2

13

13

27
13

71
1
13

15
13

13

9
1
6

1
98

48

677

96

67

51

74

728

556

595

397

26

l3
16

3

3

2
2

15

134

213

63

12

15

15

32

53

13

1

220 1038 1885

354

92

163

136

267

405

171

2
13

2

l3

72

76

101

2.7 719

479

993

--------------------------------------------------------------------------------------------------------------------

�Table 6 . Resident elk hunting pressure showing county of residence and number of participating license holders
by Game Management Unit, 1969, continued.

County of
Residence

Adams
Alamosa
Arapahoe
Archuleta
Baca
Bent
Boulder
Chaffee
Cheyenne
Clear Creek
Conejos
Costilla
Crowley
Custer
Delta
Denver
Dolores
Douglas
Eagle
Elbert
El Paso
Fremont
Garfield
Gilpin
Grand
Gunnison
Hinsdale
Huerfano
Jackson
Jefferson
Kiowa
Kit Ca rson
Lake
La Plata

68

69

1

70

71

13

6

72

73

74

13

65
2
9
13
13

41
13

3

Unit Number
77
76
75

78

79

80

81

82

25

43

27
69
15
13

37
298
17

2
189
17

13

7
7
26

2
2

27
4
6

4
2

34
27
49
26
29

30
122
6

19
281

37
28

1

22

21
13

15
1

38

84

85

2

7
1

6

57

1
15
172

83

5-

1
3

13

1
6
21
16

1
13
16

52

13
16
23

3
13

83

4
31
4

6
54
118

115

255
15

2
1

1
35

2
2
81
104

13
22

36

26

1

8

213
46

13
13

43

10

13

13

124

92
17

13

13

2

107
19

139
19

220
23
6

119
45

175
17

70
8

6
2

38
1

13
13

13

13

22

2

6

49

2
15

13
2
6

41

14

13

6

62

136

25

2
:2

3

13

29

538 1040

3

11
6

2

6

29

1

2
372

103

---------- .---------------------------------------------------------------------------------------------------------

•.....
-...J
VI

�Table 6 • Resident elk hunting pressure showing county of residence and number of participating
by Game Management Unit, 1969, continued.

County of
Residence
Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Sunnnit
Teller
Washington
Weld
Yuma
Sportsman's
Total

68

69

70

71

72

73

74
13

14

Unit Number
77
76
75
4
30

3

13

78

79

19
12

2

80

81

82

83

license holders

84

85

2

1
179

1

1
26

10
186

54
378
43
3

13

40
13

6

26

2
14
46

282
13

13
122

46

14
46

54

19
6
6

13
13

2

13

19

47

4

10

00

6
1

41

13

167

13

13
13
118

13
101

4
2

16

2
3

1
103
1020

41

3

83

26
66

246

6

17

169

292
4
470

14

21

47

9

20
93

2

132

6
15

2

15
4

6

1

2

2

19

14

195

187

173

184

207

75 . 117

12

75

698

117

480 1181 1626 1235 1296 1400

512

t-'

Ul

2

13

15
29

13
13

73

55

45
2

26

1

13
177

105

17

6

29

37

955 1911

849

69

45

146

359

106

-------------------------------------------------------------------------------------------------------------------~

�Table 6. Resident elk hunting pressure showing county of residence and number of participating license holders
by Game Management Unit, 1969, continued.
County of
Residence
Adams
Alamosa
Arapahoe
Archuleta
Baca
Bent
Boulder
Chaffee
Cheye.nne
Clear Creek
Conejos
Costilla
Crowley
Custer
De1ta
Denver
nO'lores
Douglas
Eagle
Elbert
El Paso
Fremont
Garfield
Gilpin
Grand
Gunnison
Hinsdale
Hue r f ano

Jackson
.Ie f f'e rs on

Kiowa
Kit C8.c.son
Lake
La Plata

87

90

13

62

Unit Number
Sport~man i s
Unknown

357
58
364
62
18
11
408
149
18
33
7
14
25
139
1138
25
22
69
3

26
28

3

753
110
110
3
65
76
7
54
18
448
3
62
14
187

Total

2641
·714
2024
522
75
121
2508
870
35
128
296
37
37
161
583
6100
64
140
352
29
4712
641
1003
52
670
1262
25
157
238
3625
33
101
332
2354

-----------_._---------------------------------------------------------------------------------------------------

~

V1
\0

�Table 6 • Resident elk hunting pressure showing county of residence and number of participating license holders
by Game Management Unit, 1969, cuntinued.

County of
Residence

87

90

Unit Number
Sportsman's
Unknown

Total

-Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Summit
Teller
Washington
Weld
Yuma
Sportsman's

Total

364
18
3
22
175
7
83
158
98
7
18
25

13

3
108

.

29
18
245
36
95
113
39
18
50
44

13

30

436

172
3
6670

273

436

6670

2799
268
49
95
1879
143
589
984
1103
120
221
102
47
27
322
118
2099
460
976
940
328
32
209
40
300
132
28
1404
45

4811 501

t-'
0'\

0

�Table 7. Nonresident elk hunting pressure showing state of residence and number of participating license
holders by Game Management Unit, 1969.

-State of
Residence
Alabama
Arizona
Arkansas
Ca lifornia
Connecticut
Delaware
District of Columbia
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Kentucky
Louisiana
Maine
Maryland
Massachusetts
Michigan
Minnesota
Mississippi
Missouri
Nebraska

Unit Number
2

4

5

6

7

8

11

12

13

14

--

15

16

17

18

19

20

4
13
2

52

31

13

3

5

13
5

42

43

26
9

13
13

17

21

22

13
1
3

5

13
13
26
26
13

13
4
15
28

2
20
1
3

5
11

3

2
13

26
30
2

39
13
13
15

46
59

13

2
13
2
1

59
13

3

13

2
13
13

2

13
2
13
39

18

14

13

13

1
1

8
13

13

20
15

56
2

13

13
17
40
24

13
5

2
10
2

13

13
5

2

13

13
2

1

-------------------------------------------------------------------------------~------------------------------------

23

24

1
.3

6

11

1

52

20
103

5
1
4
9
4
7
3
1

8
1
7
15
7
13
4
1

1

1

17
8
2
13
3

29
14
2
23
4

...
'"...

�Nonresident elk hunting pressure showing state of residence and number of participating license
Table 7
holders by Game Management Unit, 1969 (continued)
0

State of
Residence

---

2

4

5

6

17

2

7

8

11

12

Unit N~mber
15
13
14

16

17

18

19

20

21

22

23

24

4
1

7
1

1
3
1
7
18
1
7
1
2
7
27

14
4
1
13
30
1
13
1
2
11
46

1
1
1
13
1

1
1
1
23
1

242

430

..

Nevada
New Jersey
New Mexico
New York
North Carolina
North Dakota
Ohio
Oklahoma
Oregon
Pennsylvania
South Carolina
South Dakota
Tennessee
Texas
Utah
Virginia
Washington
West Virgir.ia
Wisconsin
Wyoming
Mexico
Sportsman's
Total

13

29

1

1
2
17
13

1
1
13

6

13

2

13

2

13

13

39

1

2
26

13

2
1

52

2
17

13
26

13

3
1

15
2

13
27
1

13
39
1

13
6

26

26

13
26

2

13

44

58

30

3

18

59

34 278 304 116

6

52

188

13

15
13

4

13

117

109

149

30

24

14

39

8

10

333

366

530

143

59

59

190

50

25

13

27

--------------------------------------------------------------------------------------------------------------------

to-'
0'
N

�Table 7 • Nonresident elk hunting pressure showing state of residence and number of participating license
holders by Game Management Unit, 1969 (continued).

State of
Residence

Alabama
Arizona
Arkansas
California
Connecticut
Delaware
District of Columbia
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Kentucky
Louisiana
Maine
Maryland
Massachusetts
Michigan
Minnesota
Mississippi
Missouri
Nebraska

25

26

27

28

29

31

32

Unit Number
36
34
33

37

40

42

43

44

45

47

48

13
26
21

1

43

1

13

30

3

13

2
13
17

39

52
1

15

13

15

2

117
13

1
13
2
13

2
4

6

13

1

3

15

13

2
13

2
26

26

~

13

13

13

13
13

14

1

13
13

18
18

1

2

13
13

57

1

13
13

0\

w

2

13

13

13

13
2

19
16
13

18
2

16

14
13

2

13

6

66

2

--------------------------------------------------------------------------------------------------------------------

30

�Table 7 • Nonresident elk hunting pressure showing state of residence and number of participating license
holders by Game Management Unit, 1969,(continued).
State of
Residence

25

26

27

28

29

Nevada
New Jersey
New Mexico
New York
North Carolina
North Dakota
Ohio
Oklahoma
Oregon
Pennsylvania
South Carolina
South Dakota
Tennessee
Texas
Utah
Virginia
Washington
West Virginia
Wisconsin
Wyoming
-.
Mexico

13

Sportsman's

55

36

16

37

1

Total

202

133

71

146

27

31

32

Unit Number
33
34
36

37

39

40

42

43

44

45

47

48

1

2

2
15

13
2

1
13

26

26

15
15

26
30

1
10

26

66

6

7

39
t-

O'

+='

13

13

2

13

125

13

26

5

2

13

13
73

39
13

66

3

13

17

1

1

13

14

61

34

15

11

353

106

41

17

13

5

125

56

68

519

186

418

17

--------------------------------------------------------------------------------------------------------------------

46

10

215

14

�Table 7 • Nonresident elk hunting pressure showing state of residence and number of participating license
holders by Game Management Unit, 1969, (continued).
State of
Residence

Alabama
Arizona
Arkansas
California
Connecticut
Delaware
District of Columbia
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Kentucky
Louisiana
Maine
Maryland
Massachusetts
Michigan
Minne,sota
Mississippi
Missouri
Nebraska

50

52

1

13
1
58

53

54

55

13
34

13
30
68

26

13

15

15

56

58

Unit Number
63
61
62

64

65

66

67

68

70

71

39

52

103

43

72

73

13

26

13

1

13
43

13

26
47

13
13
44

31

13

16
13

26

3

4
1
1

13
3
13
15

13
14

7

3
19

14
27

14
13

13

13
15
1
14

•....

13

'"

13

V1

13
14
13

8

13
13

26
16

26
13
15
1
3
4

14
13

42

13
13
47

13

26
13
13
1

13
13
13

17
1

3
1

26
5

13

--------------------------------------------------------------------------------------------------------------------

�Table 7 • Nonresident elk hunting pressure showing state of residence and number of participating license
holders by Game Management Unit, 1969. (continued).
State of
Residence

Nevada
New Jersey
New Mexico
New York
North Carolina
North Dakota
Ohio
Oklahoma
Oregon
Pennsylvania
South Carolina
South Dakota
Tennessee
Texas
Utah
Virginia
Washington
West Virginia
Wisconsin
Wyoming
Mexico

50

52

53

5

54

55

56

58

61

Unit Number
62
63

64

65

66

67

68

70

71

13

71

72

73

15
1
13

2

26
3

13
2

6
26
13

14

2
39

32
34

13

13

13
13

30
15

3
41

13
14

13

13
39

13

13
13

13

2
2

26
5

13
15

46

63

•.....

13
26
103

13

1

114

97

13

111

157

27

2

15

'"'"

76

13
175

13

115

3

Sportsman's

17

79

34

94

138

15

7

21

43

24

2

89

77

51

15

45

128

18

43

Total

37

295

188

373

622

67

8

203

288

37

28

406

389

214

129

308

586

57

229

--------------------------------------------------------------------------------------------------------------------

�Nonresident elk hunting pressure showing state of residence and number of participating license
Table 7
holders by Game Management Unit, 1969 (continued)o
0

Sta t e of
Residence

Alabama
Arizona
Arkansas
California
Connecticut
Delaware
District of Columbia
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Kentucky
Louisiana
Maine
Maryland
Massachusetts
Michigan
Minnesota
Mississippi
Missouri
Nebraska

74

75

20

41
92
93

30

76

~·7

78

79

Unit Number
35
80
81

31
44
85

13

3
1

3
42

44
44
99

4

2
34

18

26

Unknown

Sportsman is

Total

16
24
169
490

81
386
625
2142
13
2
3
211
48
512
440
192
700
170
220
2
36
15
440
246
66
1375
206

1

4

4

13
13

11

15

73

17

22
13
5
26
22
5

18
5
5
18
13
18

13
13

13

14

39

3

13

14
3

75
16
84
118
59
84
59
118
8

4
4

15
2

39

58
6

13

104
18

13
18

2

2
13

138

26

17

1
22
1

84
33
33
356
16

--------------------------------------------------------------------------------------------------------------------

t-'

'"

-...J

�Table 7 • Nonresident elk hunting pressure showing state of residence and number of participating license
holders by Game Management Unit, 1969 (continued).
State of
Residence

Nevada
New Jersey
New Mexico
New York
North Carolina
North Dakota
Ohio
Ok l a homa

Oregon
Pennsylvania
South Carolina
South Dakota
Tennessee
Texas
Utah
Virginia
Washington
West Virginia
Wisconsin
Wyoming
Mexico

74

75

76

77

78

13

18

103

88

94

96

39
13

4
20

13
32
2

3
82

169

210

16

13

13

4
7
118

255

220

13
462

22
39
614

79

Unit Number
80
81

26
74

Unknown

13

39

66

85

28

33
8
16

1

186
270

4
18

110
24
92
1260

2
41

13
94

45

Sportsman is

16
8
50

13

15

16

13

5

Sportsman's

131

153

143

273

387

38

79

75

36

394

3931

Total

513

914

674

1461

1802

230

430

206

122

394

3931

2

Total

136.
2
632
56
33
2
532
1550
17
274
127
91
266
4945
14
18
15
10
296
15
1

17163

•.....
(j'\

ex&gt;

�Table 8 • Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969.
County of
Residence

(,oc;.ejosCostilla

Custer

Delta

Dolores

1

9

7

27

11
196
13

30

9

2

1

7

2

Clear Creek

Alamosa

Archuleta

Boulder

Chaffee

9
38

30

5

9

27

33
364
13
23
4

1

32

319

33
447

-...,_ ..-,.,-_.

Adams
Alamosa
Arapahoe
Archuleta
Ba ca

Bent
Boulder
Chaffee
Cheyenne
Clear Creek
Conejos
Costilla
Crowley
Custer
Delta
Denver
Dolores
Douglas
Eagle
Elbert
E1 Paso
Fremont
Garfield
Gilpin
Grand
Gunnison
Hinsdale
Huerfano
Jackson
Jefferson
Kiowa
Kit Carsor
Lake
La Plata

9
9

,.

67
306
21

11

9

46
36

56

262
43
2

53

30

19

1

395
51

9
21
19

49
9
9

21

6
3
1

9
9

~
\.0

9

2
122
2

t-'

9

291
40

2

2
9

67
4

41
41

9

1

11
2

20

11

9

9
7

2
27
9

48

35

9

4
112

97

9

372

-------------------------------------------------------------------------------------------------------------------

�Table 8.
Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969 (continued).
County of
Residence

Alamosa

Archuleta

Boulder

Chaffee

Clear Creek

Conejos

Costilla

Custer

Delta

Dolores

,----

Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Summit
Teller
Washington
Weld
Yuma
Sportsman IS

31
13

19

3

3

11

2

2
192

42

53

522
49

1
21
63
9
2
2

9

11

10
t-'
-...J

1
11

9
67

2

0

1
102

9

7

11
116

59

11

2

37
9
2
11

9

18

256

1

29

3
4

47

119

80

1
9
107

21
5

34

123

75

Total
96
1,803
504
1,329
389
972
33
242
1,003
-----_. __ ._----------------------------------------------------------------------------------------------------------

844

�Table 8 • Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969. (continued).

County of
Residence

Adams
Alamosa
Arapahoe
Archuleta
Baca
Bent
Boulder
Chaffee
Cheyenne
Clear Creek
Conejos
Costilla
Crowley
Custer
Delta
Denver
Dolores
Douglas
Eagle
Elbert
E1 Paso
.Fr emont

Garfield
Gilpin
Grand
Gunnison
Hinsdale
Huerfano
Jackson
Jefferson
Kiowa
Kit Carson
Lake
La Plata

Eagle

E1 Paso

Fremont Garfield

Gilpin

Grand

Gunnison

Hinsdale

Huerfano

Jackson

Jefferson

2
2
16
13

15

113

2

11

67

12

191

9

96

30

339

111

81

9

55

1

228

189

12
14

31

1

2
182
5

2
64
74

61
15

413

199
1
91

280

128

2

646
21

11
361
19

43
19

4
29
110
621
2
9
2

9

124
11
19

4
3

11
127
13

228
9
9

21

2

2
277

7

9
9
308
299
2
9
2

9
43
9

597
87
29

53
19

933
2
9

16
28

196
9
9
21

71

9
21
9

21
1

t-'
-...J
t-'

154

22

11

1

67

2
25

521

72

2
392

85
2

270
235

67

9
1

20

-------------------------------------------------------------------------------------------------------------------

�Table 8 • Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969 (continued).
County of
Residence

Eagle

El Paso

Fremont Garfield

Gilpin

Grand

Cunn i s on

Hinsdale

Huerfano

Jackson

Jefferson

247

2

.-.~.--.

Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Sunnnit
Teller
Washington
Weld
Yuma
Sportsman's
Total

37

9
9
51

5
75

1
3

5
9
2

9

92
9
15
6

13
9
18
4
97

9
296
19
49

2
1
16
4

25
2

2
14
2

3

3

9

4

t-'

7
2
2
9

19

-...J

9
2
9
280

26
6

7
18

N

168
9

2
2

14
11

13

1
19

2
31

9
9

9
2

13
46
2

62
2

9
16
1
4

4

2
1
209
3

269

1

13

298

13

366

492

182

31

185

15

2,222

20

147

1,942

130

3,098

4,306

583

392

1,896

126

-------------------------------------------------------------------------------------------------------------------

�Table 8 • Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969 (continued).

County of
Residence

Adams
Alamosa
Arapahoe
Archuleta
Baca
Bent
Boulder
Chaffee
Cheyenne
Clear Creek
Conejos
Costilla
Crowley
Custer
Delta
Denver
Dolores
Douglas
Eagle
Elbert
El Paso
Fremont
Garfield
Gilpin
Grand
Gunnison
Hinsdale
Huerfano
Jackson
Jefferson
Kiowa
Kit Carson
Lake
La Plata

Lake

La Plata

Larimer

Las Animas

Mesa

Mineral

11

78
4
14
13
11

111

3

9

42
40
41
29
19

9

10

39
2

2
11

49
11
9

2
9
13

311
2
2

23
9

Mo:Ffat Montezuma
9

11

26

Montrose

Ouray

Park

11

21

49

9

2

78

15
3

71

t-'
....•

w

16
2
9
1
16

4
92
2

22
35

142

35

6

7

42

94
13

1
1
34

2
9
4

58
9

29
4

9

1

2
143

25

33

2

2

4

31

9

1
219
3

2

7

114

19
19

13

13

36

160

9

9

3
10

37

3
23

120

9
2

94

2
1,460

9
9

2

37

9

4

1

2
13

11

9

-------------------------------------------------------------------------------------------------------------------

9

�Table 8 .• Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969 (continued).
County of
Residence

Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Sunnnit
Teller
Washington
Weld
Yuma

Lake

La Plata

Larimer

2

13
28

1,499

Las Animas

Mesa

Mineral

Moffat

Montezuma

Montrose

Ouray

Park

19

9
9

21

29

49

19

574
11
2

414

185

2

19
87

20

1
134

33

2
778

7
2
58
58

1
19
175

55
289

21

2
2

9

2

9
59

.po.

9
11

44

20
65

3

9

28

19

57
81
2

9

2

19

4
21
2

16

9
9
40

2
11
4

9

4

1
12
1
8

127

93

49

136

793

783

501

920

4

9

4

437

Sportsman's

42

364

338

31

Total

266

2,549

3,082

207

11

22

143

165

121

1,092

1,019

842

to-'

'-I

1

4

----------~--------------------------------------------------------------------------------------------------------

�Table 8 • Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969 (continued).
County of
Residence

Adams
Alamosa
Arapahoe
Archuleta
Baca
Bent
Boulder
Chaffee
Cheyenne
Clear Creek
Conejos
Costilla
Crowley
Custer
Delta
Denver
Dolores
Douglas
Eagle
Elbert
El Paso
Fremont
Garfield
Gilpin
Grand
Gunnison
Hinsdale
Huerfano
Jackson
Jefferson
Kiowa
Kit Carson
Lake
La Plata

Pitkin

Pueblo

Rio Blanco

37

240

17

143

21
9

15
203

Rio Grande

30
292
7
4
16
9

Sa ?"'lcS . ~.~t..!

466

45
60
45
2

384

243

11

3

2
2
2
119

15
430

17
75

1
5
10
161
25
126

l39
1
215

Routt

San Miguel

9

Summit

Teller

45

9
9

40

4

36

t-'
-...J
VI

94

9
9
815

9

19
4

120
21
29

2
47
293
11
2

San Juan

234
9
2
2

21
11
189
2

9

9
2

92
9

420
103

19

31

21

38
42

9
16

9

86

224

38

502

3

24

2

21
9

36

4

47

2
18

-------------------------------------------------------------------------------------------------------------------

85
9
9

�Table 8 • Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969 (continued).
County of
Residence

Pitkin

337

Sportsman's

152

1,290

Rio Blanco
133
9
5
22
128

9

Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Summit
Teller
Washington
Weld
Yuma

Total

Pueblo

10
3

9
11

Rio Grande
2
2

Routt

Saguache

239

31
4
2

28
58

40
5

258

t~5

18
19

63

San Juan

San Miguel

Sunnnit Teller

2

2

2

9
4

177
11

2

2

2

4

I-'
-...J

4
20

5
19
213

337
4
512

19
394

2

9

9

33
738

26

(j\

47

76
9
270

9
2
4
133

2

9
9

220
29

4

71
15

9

227
4

13

2

9

19

1

541

202

665

236

13

73

58

15

21

2,899

1,880

5,057

2,391

112

458

656

147

11

---------------_._--------------------------------------------------------------------------------------------------

�Table 8. Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969 (continued).
County of
Residence
Adams
Alamosa
Arapahoe
Archuleta
Baca
Bent
Boulder
Chaffee
Cheyenne
Clear Creek
Conejos
Costilla
Crowley
Custer
Delta
Denver
Dolores
Douglas
Eagle
Elbert
El Paso
Fremont
Garfield
Gilpin
Grand
Gunnison
Hinsdale
Huerfano
Jackson
Jefferson
Kiowa
Kit Carson
Lake
La Plata

Unknown

Sportsman's

357
58
364
62
18
11
408
149
18
33
7
14
25
139
1,138
25
22
69
3
753
110
110
3
65
76
7
54
18
448
3
62
14
187

Total

2,600
690
2,051
492
63
77
2,114
1,378
9
135
350
37
50
147
1,015
5,671
74
150
544
55
4,620
560
1,301
32
667
1,108
37
178
293
3,417
25
108
517
2,124

--------------_._----------------------------------------------------------------------------------------------------

t-'

~
~

�Table 8.
Resident elk hunting pressure showing county of residence and number of participating license
holders by county, 1969 (continued).
County of
Residence

Unknown

Sportsman's

Total

-Larimer
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Morgan
Otero
Ouray
Park
Phillips
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
Routt
Saguache
San Juan
San Miguel
Sedgwick
Summit
Teller
Washington
Weld
Yuma
Sportsman's

Total

364
18
3
22
175
7
83
158
98
7
18
25
29
18
245
36
95
113
39
18
50
44
172
3
376

2,840
228
38
195
1,816
182
672
1,400
1,396
133
247
91
88

23
385
100
2,015
358
823
941
413
34
205
4
359
144
29
1,428
29

6,670

49,405

J-I

-...J

ex&gt;

�Table 9.
Nonresident elk hunting pressure showing state of residence and number of participating license
holders by county, 1969.

State

Alamosa

Archuleta

Boulder

Chaffee

Count~
Clear Creek

Conejos

Delta

Dolores

Eagle

43

46

12
15
46

1
37
21
2

Fremont

-Alabama
Arizona
Arkansas
California
Connecticut
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Kentucky
Louisiana
Maine
Maryland
Massachusetts
Michigan
Minnesota
Mississippi
Missouri
Nebraska
Nevada
New Jersey
New Mexico
New York

30
64
105

15

102

54

21

12

3
24

12

33
12
15

12

24

3

12
15

12

30

12
24
12
3

12
3

18

12

134
3
27

3
3

149

30

12
9

12
12
12

12
19
82

12
67

24
12

-------------------------------------------------------------------------------------------------------------------

•....
-...J

\0

�Table ..g., Nonresident elk hunting pressure showing state of residence and number of participating license
holders by county, 1969 (continued),

State

North Carolina
North Dakota
Ohio
Oklahoma
Oregon
Pennsylvania
South Carolina
South Dakota
Tennessee
Texas
Utah
Virginia
Washington
West Virginia
Wisconsin
Wyoming
Mexico

Alamosa

Archuleta

Boulder

Chaffee

Count:t:
Clear Creek Conejos

Delta

Dolores

Eagle

Fremont

2
12
228

12

27

24

24
15
24
12

24
49

12
55
18

12
12

15
37
818

20

46

12
15

250

68

1
I-'

3
15

00

0

3

6

Sportsman

2

432

1

15

6

69

74

101

115

Total

29

2,219

25

86

36

245

345

687

522

--------------------------------------------------------------------------------------.--------------------------

55

�Table 9 . Nonresident elk hunting pressure showing state of residence and number of participating license
holders by county, 1969 (continued).

State

Count
Hinsdale Huerfano

Garfield

Grand

Gunnison

42
12

12

76

14

24
43
201

24
40
27

64

12

64
18
3
87

3

Jackson

Jefferson

Lake

LaPlata

Larimer

3

61
148
132

15

-.Alabama
Arizona
Arkansas
California
Connecticut
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Kentucky
Louisiana
Maine
Maryland
Massachusetts
Michigan
Minnesota
Mississippi
Missouri
Nebraska
Nevada
New Jersey
New Mexico
New York

18
18
12
3
27

40
3
95
43
12

1
15
1
1
12

1
3
15
26
12

40
12
12
102

IS

12

24 .
12

18
12

39

10
2
5
30

12
3
12

15
27

4

3

6

27
12

1

43

15
24

I-'
00

I-'

4

12

4

13
5

4

12

3

12
3

12

77

15
42
13

15

12

171

------------------------------------------------------------------------------------------------------------------

�Table 9 • Nonresident elk hunting pressure showing state of residence and number of participating license
holders by county, 1969 (continued).

State

North Carolina
North Dakota
Ohio
Oklahoma
Oregon
Pennsylvania
South Carolina
South Dakota
Tennessee
Texas
Utah
Virginia
Washington
Wes t Virginia
Wisconsin
Wyoming
Mexico

Garfield

Grand

Gunnison

Hinsdale

39
21
12

12
61

24
123

43

Count:t
Huerfano Jackson

Jefferson

Lake

7

LaPlata

Larimer

12
55

4

37

1
198

37

3
49
305
12

1
256

28

1

284
t-'

00
N

12
24

3

[1-0

Sportsman

169

82

331

168

6

60

Total

852

330

1,614

666

54

265

18

9

239

37

15

1,328

124

----------------------------------------------------------------------------------------------------------------

�Table 9 • Nonresident elk hunting pressure showing state of residence and number of participating license
holders by county, 1969 (continued).

State

Las Animas

Mesa

Mineral

Moffat

Count
Montezuma

Montrose

Ouray

Park

Pitkin

Rio Blanco Rio Grande

Alabama
Arizona
Arkansas
California
Connecticut
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Kentucky
Louisiana
Maine
Maryland
Massachusetts
Michigan
Minnesota
Mississippi
Missouri
Nebraska
Nevada
New Jersey
New Mexico
New York

12
37
6
18

92

77

12
42

30

89

12
70

1

49
12

12
40
223

3
43

12
9
3
16

12
6

52

12
27
12
12

28

12
12

12

12

12
18
24
15

4
1
1
1

17

30
10
15
38
10

24

37

00

12
5

24
1

12

37

12

55

12

24
24
41

6
13
1

1
12

3
1
40

18
17

12

6
1

12

36
31
10
95
1
35
5

6

---------------------------------------------------------------------------------------------------------------------

I-'

w

j

12
18

37

�Table 9 . Nonresident elk hunting pressure showing state of residence and number of participating license
holders by county, 1969 (continued).

State

Las Animas

North Carolina
North Dakota
Ohio
Oklahoma
Oregon
Pennsylvania
South Carolina
South Dakota
Tennessee
Texas
Utah
Virginia
Washington
West Virginia
Wisconsin
Wyoming
Mexico

6

Sportsman

20

Total

72

Mesa

Mineral

Moffat

12

12
3
122

1
1

15
12

Countl
Montezuma Montrose

24

12
12

Ouray

Park

18
18

Pitkin

13
24

24

24

26
74
12
32

58

10
13
89

2
64

12

252

12
54
1

12
117

121

89

1

Rio Blanco Rio Grande

24
80

12
61
I-'

00
.p..

3

5
5

2

26
5

3

27

12
12

86

171

83

68

73

74

14

75

440

77

337

837

362

324

424

422

55

328

1,345

434

1

---------------------------------------------------------------------------------------------------------------------

�Table 9 . Nonresident elk hunting pressure showing state of residence and number of participating license
holders by county, 1969 (continued).

State

Alabama
Arizona
Arkansas
California
Connecticut
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Kentucky
Louisiana
Maine
Maryland
Massachusetts
Michigan
Minnesota
Mississippi
Missouri
Nebraska
Nevada
New Jersey
New Mexico
New York

Routt

37
224
3
12
152
159
27
105
37
12
3
12
98
18
161
31
18

Saguache

40

Countl
San Miguel
San Juan

12

37

24

43

Sunnnit

Teller

12
24
6
12

3

12

12
6
6
3

55
12

12

24

12

Unknown

Sportsman

Total

15
88
92
692
6
43
11
142
84
68
231
23
38
4
11
4.
115
90
19
372
95
38
11
179
23

69
405
603
2,606
20
173
24
610
411
209
931
130
162
7
40
5
433
289
89
1,472
326
159
22
811
65

----------------------------------------------------------------------------------------------------------------

t-'

00
Vt

�Table 9 , Nonresident elk hunting pressure showing state of residence and number of participating
holders by county, 1969 (continued),

State

North Carolina
North Dakota
Ohio
Oklahoma
Oregon
Pennsylvania
South Carolina
South Dakota
Tennessee
Texas
Utah
Virginia
Washington
West Virginia
Wisconsin
Wyoming
Mexico
Sportsman

Total

Routt

Saguache

Count;l
San Juan
San Miguel

15

Teller

Unknown

3

30
40

12
15

64
3

12

153

80

12
12
58
24

Sunnnit

3
12

12

27

61

6

12

361

44

14

38

10

5

1,881

360

113

230

22

8

348

license

Sportsman

Total

6
14
103
302
11
88
4
46
36
680
10
11
3
1
98
4
4

26
26
411
1,441
59
338
43
78
189
4,246
23
22
20
25
347
45
5

3,931

l3 ,490

I-'

00

~

�- 187 -

Table 10 •

1969 regular bull license harvest by day of season of kill.

Date

Number
Killed

Percent

October 18

1,969

23.9

19

1,197

14.5

20

997

12.1

21

573

6.9

22

586

7.1

23

324

3.9

24

349

4.2

25

299

3.6

26

286

3.5

27

124

1.5

28

212

2.5

29

174

2.1

30

137

1.7

31

50

0.6

1

249

3.0

2

174

2.1

3

124

1.5

4

174

2.1

5

87

1.0

6

124

1.5

7

37

0.4

November

Total

8,246

= 79.7

�- 188 -

Table 11.

1969 ant1er1ess elk harvest by day of season of kill.

Date

Number
Kil1ed

Percent

October 18

1,269

19.9

19

1,043

16.3

20

808

12.6

21

594

9.3

22

404

6.3

23

280

4.4

24

211

3.3

25

304

4.8

26

223

3.5

27

108

1.7

28

166

2.6

29

126

2.0

30

94

1.5

31

95

1.5

1

190

3.0

2

132

2.1

3

90

1.4

4

60

0 9

5

108

1.7

6

72

1.1

November

Total

6,377

0

==

80.4

�- 189 -

Table 12.

No.

Archery elk harvest by game management unit, 1969.

Unit
Name

4
6
8
11
12
14
16
18
33
34
42
55
70
71
72
73
74
76
78
82
83

Elk Head
Cowdrey
Red Feather
Strawberry Creek
Williams Fork
Elk River
West Side
Troublesome
Rifle
Colorado River
Grand Mesa
Taylor River
San Miguel
Dolores
Yellow Jacket
Mancos
Hermosa
Creede
San Juan
Sand Dunes
Trinchera

Total

1/ Antlered only season in 1969.

Harvest
Bulls 11

Total

2
2
8
2
2
2
3
3
2
2
3
3
2
3
3
3
3
2
2

2
2
2
2
8
2
2
2
3
3
2
2
3
3
2
3
3
3
3
2
2

S6

S6

2
2

�- 190 -

Table 13.

Archery elk harvest by county, 1969.
Harvest

County Name

Archuleta
Conejos
Costilla
Dolores
Garfield
Grand
Gunnison
Hinsdale
Jackson
La Plata
Larimer
Mineral
Moffat
Montezuma
Pitkin
Rio Blanco
Routt
San Miguel

Total

1/ Antlered only season in 1969.

Bulls 1/
2

Total

2
I

1
4

4

7
7

7
7

2
2
2

2
2
2

4
4
1
4

4
4

2
2
-2

2
2
2

3

3

I

4

5

5

2

2

56

56

�Table 14.

Archery elk harvest by day of season and game management unit. 1/
1969
Date of Season

No.

Units
Name

August
16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31

2
4 Elk Head
6 Cowdrey
8 Red Feather
11 Strawberry Cr.
12 Williams Fork
14 Elk River
16 West Side
18 Troublesome
33 Rifle
34 Colorado R.
42 Grand Mesa
5S Taylor River
70 San Migue 1
71 Dolores
1
72 Yellow Jacket
73 Mancos
74 Hermosa
76 Creede
78 San Juan
2
82 Sand Dunes
2
83 Trinchera

1

2

1

Totals

5

1

SeEtember
2 3 4 5 6 7 8 9 10 11 12 13 14

1

1
2
2
3

3

2
2
2
2
1

2
3

1
1

1

2

1

2
2
3

2

1
3

Total

2
2
2
2
8
2
2
2
3
3
2
2
3
3
2
3
3

3
3

3

2
2
3

!/Data from hunter report cards.

1

5

1

3

2

1

4

4 1

262

1

4

2

6

56

I-'

1.0

I-'

�Table 15.

County

Adams
Alamosa
Arapahoe
Archuleta
Baca
Boulder
Chaffee
Clear Creek
Conejos
Custer
Delta
Denver
Douglas
El Paso
Garfield
Grand
Gunnison
Hinsdale
Jackson
Jefferson
Lake
La Plata
Larimer

County of residence of archery elk hunters showing number of hunters by unit hunted (continued).

31

33

34

35

36

37

38

39

Unit Hunted
41
42
43

1
. 1

44

45

3

1

46

461

47

1·

48

49

50

51

52

2
2

1

3

2

2
1

1

12
2

2

2

3

3

.p.

2

1
4

2

2

6

9

4
6

4

9

1

2

1
1

1
2
1

2

1

1

1

1
4

1
2

2

-----------------------------------------------------------------~--------------------------------------------------

I-'

\.0

1

�Table 15.

County
Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Otero
Ouray
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
San Juan
San Miguel
Teller
Weld

Total

County of residence of archery elk hunters showing number of hunters by unit hunted (continued).

31

33

34

35

36

37

38

39

Unit Hunted
43
41
42

44

45

46

461

47

48

49

50

51

52

2
1
1

2

6

16

3
1

7

4

4

t-'

\0
VI

1
1

1

7

22

3

1

3

1

6

1

33

12

13

1

3

1

10

13

13

--------------------------------------------------------------------------------------------------------------------

4

1

19

�Table 15.

County of residence of archery elk hunters showing number of hunters by unit hunted (continued).

County

53

54

55

1

2

56

57

58

59

Unit Hunted
66
65
62

72

71

70

68

67

77

76

75

74

73

78

1

Adams
Alamosa
Arapahoe
Archuleta
Baca
Boulder
Chaffee
Clear Creek
Conejos
Custer
Delta
Denver
Douglas
El Paso
Garfield
Grand
Gunnison
Hinsdale
Jackson
Jefferson
Lake
La Plata
Larimer

3

1

1

3

4
2
2

2
2

2

7

2

3

6
1

2

7

2

6

3

4

3

2

3

4
2

2
1

1

1

3

4

3

3
1

1

2

3

2

1
1

1

2

1

1

9

6
1

2

-----------------------------------------------------------------------------------------------------------------------

t-'
\0
0'\

�Table 15.

County

Las Animas
Lincoln
Logan
Mesa
Mineral
Moffat
Montezuma
Montrose
Otero
Ouray
Pitkin
Prowers
Pueblo
Rio Blanco
Rio Grande
San Juan
San Miguel
Te ller
Weld

Total

County of residence of archery elk hunters showing number of hunters by unit hunted (continued).

53

54

55

56

57

58

59

62

Unit Hunted
65
66
67

68

70

71

72

73

74

75

76

77

78

2
1
1
4

1

3

7

2
18

2

1

6

3

3

1

2
3

I

16

1

17

4
1

I

1
3
2
t-'

1.0

I

6

3

3

2

7

-..J

3

I

15
I

1

2

7
2

1
2

9

16

36

7

1

3

4

20

14

4

7

4

13

17

1

20

24

14

65

13

7

�Table

17.

State of residence of·archery elk hunters showing number of hunters by county hunted, 1969.

State

Archuleta

Arizona
Arkansas
California
Delaware
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Michigan
Minnesota
Missouri
Nebraska
Nevada
New Jersey
New Mexico
New York
Ohio
Oklahoma
Pennsylvania
South Carolina
Tennessee
Texas
Utah
Washington
Wisconsin

2

Total

11

Chaffee Conejos Costilla Dolores Eagle Garfield
2

2

Z

Grand Gunnison Hinsdale Huerfano Jackson

2
2

2

2
2

2

2

2

2

2

2

12

4

3

2
2

6

4

N

2

4

0

N

2
2
1
1
4

1
2
2

2

3
2

16

4
2
2
5

2
2

3

2

2

7

6

2
2

2

11

2

14

6

23

8

32

33

6

-----------------------------------~------------------------------------~~--------------------------------------------

2

�Table

17 •

State of residence of archery elk hunters showing number of hunters by county hunted, 1969.(continued)

State
Arizona
Arkansas
California
Delaware
Florida
Georgia
Illinois
Indiana
Iowa
Kansas
Michigan
Minnesota
Missouri
Nebraska
Nevada
New Jersey
New Mexico
New York
Ohio
Oklahoma
Pennsylvania
South Carolina
Tennessee
Texas
Utah
Washington
Wisconsin

Total

LaPlata Larimer Mesa Mineral Moffat Montezuma Montrose Ouray Rio Blanco Rio Grande Routt San Juan Total
3
4

2

4

9
4
1

4

2

1

28

2
2
2
2
2
2
2

2

4
4
2
11
7
7
31
11
16
16
4
2
2
16
2
15
38
13

2
3
1
2

3

2

2
2

1
4

3

8

4

4

1
2
2

2
5

8

1

2

4
2

4

2
2
2

2
4
4
2

6
2

2
2

2

6
2

6

2

2

2

2

2

22
4
2

14

4

20

33

39

2

9

6

6

1
6

7

2
35

15

13

9
2

320

N

0

v.&gt;

�- 204 -

Table 18. Total hunting days spent by archery elk hunters by Game Management
Unit, 1969.
Unit
Hunted

4
5
6
7
8
11
12
13
14
15
16
17
18
19
20
23
24
25
26
27
28
29
31
33
34
35
36
37
38
39
41
42
43
44
45
46
461
47
48
49
50

Resident

Nonresident

24
28
36
4

87
28
147
87
136
19
23 .
29
17
31
54
8
42
8
9
58
46

13
61
23
30
3
10
14
11
44
3
1

2

18
99
6
8
5
5
29
17
88
55
38
6
9
20
60
42
21
9

42
22
6

73
16
II

Total

24
28
36
4
87
41
208
110
166
22
33
29
31
31
65
52
3
43
8
·9
58
46
2
60
121
12
8
5
5
29
17
161
71
49
6
9
20
60
42
21
9

------------------------------------------------------------------------------

�- 205 -

Table 18. Total hunting days spent by archery elk hunters by Game Management
Unit, 1969 (continued).
Unit
Hunted

51
52
53
54
55
56
57
58
59
61
62
63
65
66
67
68
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
90

Resident

3
74
25
56
98
23
5

Nonresident

Total

46
49
10

3
4

3
74
37
72
165
23
5
5
9
26
100
4
67
20
15
11
39
106
44
95
112
96
516
69
57
26
152
107
5
5
46
52
14

Total

2,992

1,026

4,018

Ave.

5.72 days

4.52 days

5.36 days

13
16
67

5

9
21
68
49
11
15
11
39
90
7
74
103
69
293
40
30
26
112
68
5

5
32
4
18
9

16
37
21
9
27
223
29
27
40
39
5

�- 208 -

Table 21. Pre-season deer harvest by sportsman's license holders, by
Game Management Unit, 1969. 1,/
Unit
Number

1/

-

Buck Harvest
Nonresident
Resident

Total Harvest

7

14

3

17

47

14

3

17

78

53

14

67

82

40

10

50

90

28

6

34

Total

149

36

185

.

Does not include archery or primitive weapons harvest.

�- 209 -

Table 22. Regular season deer harvest by sportsman's license holders, by
Game Management Unit, 1969. 11
Unit
No.
1
2
3
4
5
6
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
381
39
40
41
411
42
43
44

Name

Green River
Browns Park
Craig
Elk Head
Snake River
Cowdrey
Red Feather
Buckeye
Blue Mountain
Strawberry Creek
Williams Fork
Dunkley
Elk River
Yampa
West Side
Owl Mountain
Troublesome
Poudre River
Estes Park
Douglas
Piceance
Miller Creek
White River
Sweetwater
Toponas
Sheephorn
Hot Sulphur
Boulder
Salt Creek
Roan Creek
Parachute Creek
Rifle
Colorado River
Castle Peak
Piney
Summd t;
Central City
Ra 1ston Creek
Mt. Evans
Glade Park
Kannah Creek
Surface Creek
Grand Mesa
Roaring Fork
Brush Creek

Bucks
NR
R
34
34
17
34
42
26
60
8
8
24
34
59
153
34
9
119
128
102
60
94
59

9
17
8
84
76
68

NR

R

134

9

30
30

42
34
17
8

Fawns
NR
R

Total
NR

R
51
34
59
68
59
34
68

8
15

8

Total
Harvest

134
45
30

8

15
30
30
15
15
15

17
18
17
59
60
76
17

30

8

15

8

25
42
51
118
213
119
17
17
162

8

15

17

59
42
68
59
17
8
17
17
34
60
34
59
42
8
17

Does

60
15

34
34
8
8
8
25
8
43
17
34
25
17

9
8

45
9
15
26
9
17
8

15
15
17

8
15
15

8
17
42
9

9

17

8
8
8
119
60
45

34
43
60

8

110
68
102
93
25
102
59
128
93
51
25
17
51
68
42
76
93
17
34
8
17
25
8
118
119
136

15
30
60
15
15
30

170
15
45
60
30

15
15

15
15

119
60
45

----------------------------------------------------------------------------------

185
34
59
113
89
34
68
8
40
72
111
133
228
149
17
17
162
170
125
68
147
93
25
162
89
128
93
51
8
40
32
51
68
42
91
93
17
49
8
17
25
8
237
179
181

�- 210 -

Table 220 Regular season deer harvest by sportsman's license holders, by
Game Management Unit, 1969 (continued). 11
Unit
No.
45
46
461
47
48
49
50
51
52
53
54
55
56
57
58
59
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
89
90
931
937
Totals

Bucks
Name

Holy Cross
Bailey
Deer Creek
Frying Pan
Mt. Elbert
Buffa 10 Peaks
South Park
South Platte
N.F. Gunnison
Coal Creek
Sapinero
Taylor River
Buena Vista
Badger Creek
Eleven Mile
Pikes Peak
W.S. Uncompahgre
E.S. Uncompahgre
Smith Fork
Olathe
Alpine
L.F. Gunnison
Tomichi
Saguache
Grape Creek
San Miguel
Dolores
Yellow Jacket
Mancos
Hermosa
Animas River
Creede
Piedra
San Juan
La Garita
S.F. Rio Grande
Conejos
Sand Dunes
Trinchera
Greenhorn
Picket Wire
Branson
Sangre de Cristo
Crow Valley
Bijou

R
8
17
17
42
42
68
34
26
51
76
128
128
59
9
76

Does
NR

R

Fawns
NR

NR

R

8
34
25
42
76
85
42
34
68
85
230
239
59
17
76

17
8
15

25
17
8

45
30
60
45
15

8
9
42
43

9

45
45
15
15

8
9
8

8

15
-IS

17
94
50
17
8
25
68
76
25
51
42
8
34
59
110
42
34
59
51
128
59
51
8
25
51
68

30
208
15
119
60
15
30
60
60
15
30
75
119
15
60
282
30
45

8
42
9
8
9
34
34
9

30
45
30

15

15

15
8

25
17
15
17
51
9
17
17
17
42
17

15
30

9
8

45
8

8

15
8

17
4,016 2,211

1,497

450

210

liThe estimated regular season deer kill was accomplished
resident hunters and 4,114 nonresident hunters.

R

25
136
59
25
17
59
110
85
2S
76
59
8
34
76
170
51
59
76
76
170
76
51
8
25
51
8
68
8
17

Total
NR

15

90
75
75
60
15
15
60
268
45
119
90
15
30
60
60
30
30
90
149
15
60
327
30
45

15

45 5,723 2,706

Total
Harvest
8
34
25
42
91
85
42
34
158
160
305
299
74
17
76
15
85
404
104
25
126
149
125
115
25
136
119
38
64
166
319
66
119
403
106
170
121
51
8
25
51
8
83
8
17
8,429

by an estimated 10,189

�- 211 -

Table 23.

Resident Sportsman's license S~ll

Game Survey, 1969.

Species

Number of
Hunters

Total
Bag

Ave. Season
Bag
Per Hunter

Scaled Quail

247

2,371

9060

2056

GSD1beh Quail

55

232

4.22

2.46

Bobwhite Qua il

179

832

4.65

2..
05

Chukar Partridge

99

421

4.25

2.12

Mourning Dove

1,923

30,480

15.85

3.56

Sage Grouse

717

2,581

3.60

1.90

Sharp-tail Grouse

117

240

2.05

1.63

Blue Grouse

1,162

2,928

2.52

1.91

Ptannigan

192

526

2.74

1.60

Pheasant

3,234

11,093

3.43

2052

Cottontail Rabbit

3,166

23,301

7.36

4.32

Snowshoe Hare

482

1~190

2.47

3.17

Fox Squirrel

136

481

3.54

2.20

Ducks

1,799

14,014

7.79

5.44

Geese

408

1,081

2.65

4.89

Ave. Days
Hunted
Per Season

�- 212 -

Table 240

Nonresident Sportsman's license Small Game Survey, 1969.

Number of
Hunters

Total
Bag

Ave. Season
Bag
Per Hunter

Ave. Days
Hunted
Per Season

Bobwhite Quail

9

42

4.65

15.00

Mourning Dove

10

70

7.00

1.00

Blue Grouse

29

77

2.66

1033

Cottontail Rabbit

56

158

2.83

3.50

Snowshoe Hare

48

77

1 60
0

4.16

Fox Squirrel

35

131

3 75

2075

Ducks

115

814

7.08

3031

Geese

10

40

4.00

4.33

Pheasant

31

93

3 00

1.40

Species

0

0

�- 213 -

Table 25.

Sportsman's license fishing survey, 1969.
Non-resident

Total

10,213

1,320

11,533

8,377

1,320

9,697

100

o

100

1,736

o

1,736

1 - Streams

1,911

362

2,273

2 - Lakes

2,036

645

2,681

3 - Both

6,266

313

6,579

1 - Bait

1,154

381

1,535

2 - Lures

541

264

805

3 - Flies

920

147

1,067

4 - Bait and Lures

1,740

205

1,945

5 - Bait and Flies

1,064

117

1,181

6 - Lures and Flies

1,258

98

1,356

7 - All Three

3,536

108

3,644

Total Trout Bag

502,377

22,215

524,592

Average No. Trout/Season

49.19

16.83

46.72

Days Fished/Season for Trout

16.77

4.28

15.68

Hours Fished/Day for Trout

4.94

3.27

4.79

Item

Resident

Number of Participants
Fished for Trout
Fished for Warm Water
Fished for Both
Fished ill.:

Type of Equipment Used

��July, 1970

- 215 -

JOB FINAL REPORT

State of

COLORADO

Project No.

W-38-R-24

Work Plan No.

4

Job Title

Deer-Elk Investigations
Job No.

3C

Evaluation of Deer~Scare Devices

Period Covered:

None

Personnel:

David F. Gordon

ABSTRACT

A final report for this job was published in the July, 1969 (Part One)
Game Research Report. It should not have been included in the W-38-R-24
job descriptions.

��July, 1970

- 217 -

JOB PROGRESS REPORT

State of

~C~O~L~O~RAD==~O~

Project No.

W-38-R-24

Deer-Elk

Investigations

Work Plan No.

6

Job No.

11

Job Title

_

Evaluation of the Effects
of Spring-Summer Grazing by Deer on Alfalfa

Period Covered:

April 1, 1969 - October

31, 1969

Personnel:

David F. Gordon, Harold M. Swope, Marion C. Coghill,
Steven F. Steinert, Julius J. Klein, Donald E. Speers,
John F. Corey, Dale F. Reed, Kenneth C. Dilinger,
coolidge Woodard, Rifl~ F~lls Hatchery personnel and
Rifle Gap Conservation Unit inmates.

ABSTRACT

Two cuttings of hay were harvested from the Rifle Gap study field. Hay
from the study plots was weighed at both harvests.
Plots with weights
of hay within five pounds of each other were paired and one plot of each
pair selected at random to be fenced.
Deer were counted hourly through
the night once a week during April and May.
In June three counts were
made and in July and August, two each. Data indicated that there is a
resident herd. The winter herd peaked, on the average, at 8:00 PM MST
then declined.
Resident numbers peaked at 9:00 PM and remained level
until 3:00 AM when they suddenly left the field. By midnight 60-70 percent of the total of all hourly counts had been attained.
The Little Hills study field was planted to Ladak alfalfa, with a nurse
crop of Colorado 37 oats, the end of May and a crop established.

��- 219 EVALUATION OF THE EFFECTS
OF SPRING-SliMMER GRAZING BY DEER ON ALFALFA
DAVID F

0

GORDON

Preparatory work establishing paired plots at Rifle Gap and planting
60 acres of alfalfa at Little Hills was completed this segmento The
report will be in two sections, Rifle Gap and Little Hillso
Po So OBJECTIVE
To determine the effect of spring-summer grazing by deer on production
of alfalfa
0

SEGMENT OBJECTIVES
1.

20
3
0

40

To establish stands of alfalfa on which to conduct the study.
To determine the number of deer using the study fieldso
To establish and fence paired plots in the study fieldso
To determine if there is a difference in production between grazed
and ungrazed plots.
METHODS AND MATERIALS - RIFLE GAP
Study Field

Preparation of Field
Bare spots left in the field last season due to poor seeding and irrigation
were reseeded by hand on May 20. These areas were watered along with the
rest of the field during regular irrigation.
Irrigation System
Pumping Station--The upper pumping station was established by filling in
the settling pond and installing three, 20-foot sections of 8-inch culvert
where the pond had been. The upper end of the first section is anchored
in a concrete header. The lower section rests solidly on the bank at the
foot of the old pond. The pipe slopes about 1 inch in 10 feet. The upper
section is covered with dirt to allow passage of machinery.
The pump tank was set on a flooring of redwood boards. A concrete slab
was poured for the pump and engine. The engine was converted to L-P gas
carburet ion and a 300-gallon propane tank installed for fuel supply.

�- 220 -

The pumping station in the middle of the field was removed and the
settling basin filled in. Sufficient pipe was secured to complete
the mainline from the middle of the field to the lower end.
Ditch Maintenance--Regular ditch cleaning_was accomplished with weedburner and shovelo The thick grass and weeds were cut down and the
less dense vegetation burned. Loose tumbleweeds, leaves and other
debris in the ditch were removed.
The diversion from West Rifle Creek to the reservoir was checked daily
during the major runoff to keep the diversioh ditch open and water in
it. As the runoff subsided this chore became a weekly inspection progressively ciosing off the creek to divert a full head of water into
the reservoir. On May 19 the creek was low enough to divert it entirely
into the reservoir. From the middle of June until irrigation was terminated the ditches, and especially the diversion, had to be checked
after each rain to be sure they were open and running.
A new headgate was built at the first diversion from the creek. This
was constructed of 2x4 spruce framing bolted and nailed together. Redwood 2xS's, grooved for hardwood splines, were bolted to the frame. A
spill was cut in the lower side above the gate which was three slabs of
redwood individually removable to adjust water flow. Concrete was
poured in the floor of the box.
Irrigation--Pipes were changed each morning. While one man shut down
and serviced the engine, the rest of the crew began moving lines. Service
consisted of checking the oil and filling when necessary. As soon as
service was complete the engine was restarted.
To move lines all but one lateral were shut down, in order not to overload the pump, moved to the next setting, hooked up, tested and the water
turned in. The last line was then shut off and reset. A maximum of six
lines, equalling 30 sprinkler heads, was operated in two sections of three
lines each. As water supply dwindled the number of effective lines had
to be decreased.
Irrigation schedule was as follows:
Begun April 21,
Interrupted April 29,
Restarted May 16,
Shut down for harvest June 20,
Restarted July 19,
Terminated August 1.
Hay Harvest
Cutting--Hay was harvested twice under contract with F. W. Scarrow of
Rifle, Colorado. The hay was cut with a Hesston, 1969, model 500, selfpropelled, swather-conditioner with l2-foot sickle-bar as follows:

�- 221 -

1.

Cut all hay the length of the field beginning from the ends of
the plots and working out to the edge of the field.

2.

Cut the hay in the study plots by having a helper stand at one
end, midway between the corner posts, for the swather operator
to sight on.

3.

Cut the rest of the hay parallel to the plots, a helper pulling
each corner marker as the swather approached and replacing it
in the same hole when the machine had passed.

Baling--Baling was accomplished with a tractor-drawn, John Deere, 1962,
model 2l4T, string-tie baler drawn by a Massey-Ferguson, 1967, model 165
tractor as follows:
1.

Bale all windrows running the length of the field outside the
ends of the plots.

2.

Bale all windrows within plots as follows:

3.

a.

Mark with a sheet of newspaper and tie the last bale from
the opening windrows.

bo

Subsequently mark and tie the last bale from each ploto

Bale all windrows between plots in the following sequence:
a.

All middle windrows up the field.

b.

All lower windrows back down the field.

c.

All upper windrows back up the field.

Study Plots
Harvest Data
Selecting Plot Pairs--A crew followed inunediately behind the baler weighing
the hay on a Chatillon spring scale, type 160, recording the weight of
each bale to the nearest half pound. The resulting data were grouped and
ranked from heaviest to lightest and the standard deviation calculated for
each group. One tenth of the highest value of s was rounded to the nearest
five pounds and this figure used as the upper limit of difference for
selecting plots to be paired.
Plot pairs were selected as follows:
1.

Select pairs with the least difference in weight between plots
up to a five-pound maximum.

2.

In the case of a choice between three plots with equal difference
in weight, select the two closest together.

3.
4.

Do not pair 200-foot plots with l50-foot plots.
Discard all 150-foot pairs.

�- 222 -

Fencing Plots--The plot to be fenced was selected by flipping a coin.
Heads picked the higher numbered plot; tails, the lowero Treated,
wooden, corner posts were set in holds dug 36-40 inches deep at each
corner. These were braced with 6-foot, steel T-posts set on a rock
in the ground. Line posts consisted of 10-foot, steel T-posts driven
every 10 feet between corner postso Two courses of 39-inch-wide
woven-wire were stretched between the corner posts and clipped to each
line post (Figo 1).
Deer Numbers
Deer counts on the study field were made from April 2 through August 18
as outlined under segment 23. In addition, at each count the position
of each group of one or more deer was marked on a map (Appendix I). Due
to other commitments and harvest interference, counts were missed at the
following times:
1. The second and third weeks of June (June 16 and 23).
2. The first two weeks of July (July 7 and 14).
3.

The first week of August (August 4) •

Counting was discontinued after the third week of August (August 18).
RESULTS AND DISCUSSION - RIFLE GAP
Segment objectives 1 and 3 have been fulfilled. Objectives 2 and 4 will
not be completed until the study is finished. Work on objective 4 could
not be started this year because plots could not be selected until the
second cutting had been completed and the hay weights analyzed.
Study Field
Preparation of Field
Inspection of the areas reseeded this spring showed a good stand of young
alfalfa.
Irrigation System
Pumping Station--The settling ponds were both filled on March 24 before the
snow had completely melted. About 9 inches of snow on top of frozen ground
at the bottom of the ponds was buried. The lower pond site was leveled out
on May 28. At the permanent station the culvert had to be installed before
thaw in order to begin irrigation as soon as possible. As the ground thawed
it settled under the upper two-thirds of the tube. Dirt and snow were removed from under the lower end of the first length of culvert and the middle
of the second length and backfilled with dry dirt packed down. The two
threatened sections of the tube were held up by these rests, the solid
supports at each end and the two jOint couplings until dry dirt could be
packed under their full length.

�- 223 -

Fig. 1.

Photograph of a fenced plot showing corner and line posts,
corner braces and woven wire on Rifle Gap Alfalfa Damage
Study Field.

�- 224 -

Table 1 summarizes the relative costs of regular and L-P gas for the
two years the study has been in effect.

Table 1. Summary of comparative costs of regular and L-P gas to operate
the Rifle Gap irrigation pump engine.
Gas

Hours

Gallons

Cost/Gal.

Total

Regular

1368

600

$.239

L-P

1584

660 l/

$8175

Cost/Hr. Gal./Hr.

Year

$143.40

$.105

.44

1968

$115.50

$.073

.42

1969

l/ThiS figure is less an estimated 180 gallons left in the tank at the end
of the season.

L-P gas is cheaper than regular, recovering the cost of conversion to L-P
carburet ion in three years. Since all that is required with L-P gas for a
season of operation is one oil change and oil level kept up, this is also
cheaper. When using regular gas, oil must be changed every three days as
well as keeping the crankcase full between oil changes. No information on
relative cost of maintenance is available.

Ditch Maintenance--A man was hired April 2 whose primary responsibility was
control of the irrigation water supply for peak efficiency. Secondly, he
was to service the engine and supervise changing irrigation pipes. Thirdly,
he was to improve the existing ditch system when time permitted. This latter
involved a good bit of lonely shovel work. The man hired, seriously neglected
his responsibilities. His employment was terminated April 29.
The new headgate worked well even though water washed under the gate before
the concrete floor was poured. This leak did not occur until flow in the
ditch was low enough not to cause an oversupply to the pump tank.
Irrigation--About the middle of May, water pressure at the pump dropped
drastically even though the headgate was open wide. A subsequent check revealed five minor leaks in the feeder ditch between the headgate and the
first flume, due to a buildup of algae and silt; and two culverts plugged
with tumbleweeds. The leaks were repaired and the pump shut down while the
ditch was reopened. When water had cleared irrigation was resumed.
Through negligence the pump engine was permitted to run out of oil causing
the crankshaft to freeze and break the housing. This necessitated replacement of the engine at a cost of $236.51 and interrupted irrigation from
April 29 to May 16.

�- 225 -

After April, irrigation pipe was changed under supervision of Rifle Falls
Hatchery personnel with a crew from the Rifle Gap Conservation Unit, five
days a week. On weekends one of the crew members took upon himself the
responsibility of supervision and did a very commendable jobo
Interruption of irrigation the first two weeks of May necessitated running
both upper and lower sets of sprinklers together as long as water supply
lasted in order to give the field adequate coverageo This made the task
much more unpleasant because of mud and wet alfalfa.
Hay Harvest
First cutting proceeded without mishap. Procedures for cutting and baling
were perfected and the hay stacked before it rained. Second cutting was
rained on twice in the windrow and once in the bale before it was stacked.
Study Plots
Harvest Data
Selecting Plot Pairs--Weights of hay from study plots are summarized in
Tables 2 and 3.
Calculated standard deviations were as follows:
First Cutting

NW Section, 26 pounds;
E Section, 51 pounds;
SW Section, 65 pounds.

Second Cutting

NW Section, 42 pounds;
E 'Section, 22 pounds

SW Section, 33 pounds
One tenth of the highest value of s for all six calculations rounds to 5
pounds. This figure was used as the upper limit of difference between plots
of a pair.
Plots to be paired (Fig. 2 and 3) were selected on the basis of second cutting
data for the following reasons:
1. Values of s were lower.
2. There were fewer weeds to bias the data.
3. More pairs could be selected. (Ten using first cutting weights,
14 using second).

�- 226 -

Table 2. ~eights of hay from tentative study plots on Rifle Gap Alfalfa
first cutting, 1969.
Damage Study Field
NW Section
Plot No.
Lbs. of Hay

E Section
Plot No.
Lbs. of Hay

SW Section
Lbs , of Hay

Plot No.

346.0

12

321.0

28

294.5

40

326.0

13

312.0

33

292.0

45

318.0

10

292.5

16

288.0

38

312.5

14

286.5

19

280.0

39

294.0

6

280.0

17

278.0!/

41

293.5

9

269.5

15

276.0

37

290.5

11

264.5

22

·275.0

42

289.0

7

258.5

23

251.0

36

286.0

2

255.5

.18

237.5

44

283.0

8

242.0

21

233.5

51

277 .5

5

237.0

32

229.5

46

266.0

4

236.0

29

216.5

43

262.0

3

231.5

20

192.0 .

47

249.5

1

224.01/

34

148.0

50

220.0

31

143.0

35

194.0

27

142.0

49

173.0

30

133.5

48

169.0

26

96.5

52

157.5

24

152.5

25

1/ These weights are about 5 pounds short due to reba1ing of broken bales.

�- 227 -

Table 3. Weights of hay from tentative study plots on Rifle Gap Alfalfa
Damage Study Field -- second cutting, 1969.
NW Section
Plot No.
Lbs.of Hay

E Section
Plot No.
Lbs. of Hay

SW Section
Plot No.
Lbs. of Hay

176.5

8

142.0

15

179.0

36

172.5

12

141.5

16

167.5

41

158.5

10

133.5

24

164.5

38

153.5

13

ll8.01/

19

164.0

37

145.0

6

u s.o

29

164.0

40

140.5

14

ll4.5

28

162.5

35

128.0

7

108.011

32

159.5

42

127.011

II

107.0

27

141.0

39

102.5

9

104.5

17

139.0

44

86.0

5

102.0

34

133.0

45

69.5

2

98.5

23

127.5

46

67.5

1

98.5

31

118.5

43

65.0

3

96.5 .

18

ll5.011

48

40.01:./

4

85.0

25

99.5

51

84.0

22

98.5

50

83.5

33

84.5

52

76.5

26

71.5

49

76.5

30

-- 'l:l

47

71.0

21

-- 2,..1

20

II These weights are about 5 pounds short due to reba1ing broken bales.

II These weights were lost due to broken bales.

�- 228 -

Fig. 2.

Photograph of Rifle Gap Alfalfa Damage Study Field With
Study Plots Established.

�"

. ,: '.' ....

rl&amp;-Jl-Jl-"-C:-

..

..
__ -

••

0-

'

:i

\\

i,

•

I

IN"

-- --=---•...-.-

.--

..-

1-i"''''''

..-...'--.

00...

00

••

••

0

:i

•

II
I

00

•.....•.....

•• ••
••

I

I I

0

10

. If')

.'

..

.

.r::"

-

,'-'" -r : - ---,'. ,

--:

Legend

---

",---_.i- -- - ,I

Open Plot

"

Closed Plot
Boundary of New Seeding
Boundary of Alfalfa
Plot

Field

Pairs

NW

E

SW

Open Closed

Open Closed

Open Closed .

2
I I
12

I

15

33

8

30
29
23

16
22
26
28
3I

40

7

27

34

10
6

13
14

38
35

1

37
41
42

Fig. 3. Map showing the location of paired plots on Rifle Gap Alfalfa Damage Study Field.

N
N
\0

�- 230 -

Fencing Plots--Fencing was completed in six days with the help of Project
and Little Hills personnel assisted by a crew from the Rifle Gap Conservation Unit. When stretching the top course of wire we chained the top
of the post, where the wire was fastened, to a parked pickup before applying
tension. This prevented the corner post from being pulled loose in the
ground. Gates for the plots were built by Little Hills personnel.
Deer Numbers
Deer Count Data
A shift in numbers of deer from old growth areas in the study field to new
growth is illustrated by a graph (Figo 4) plotted from data in Table 4.
Last year it was observed that the mature alfalfa scattered through the new
planting was consistently grazed to the exclusion of the innnature. This
year's observations indicated that at a definite time the new seeding
reached a point when it.was as palatable as the older plants. The reason
for this preference has not been determined.
Fig. 5 graphs deer numbers throughout the growing season. The date by
which most of the migratory deer left the field is indicated on this graph.
Table 5 sunnnarizes numbers of deer that were on the field at each hour
of each night of counting for the entire season.
The data from Table 4 were graphed on time of night (Fig. 6) to see if the
pattern of hourly visits was different in old and new growth. No marked
difference existed.
Count data from the entire season were divided into winter herd (Table 6)
and summer (resident) herd (Table 7). Graphing these results indicated
the winter herd numbers peaked at 8:00 PM MST (Fig. 7) and steadily declined
thereafter. There were no peaks after midnight. This general pattern of
behavior has been observed consistently in other studies, relating to
alfalfa and pinto beans, in other locations on the western slope of Colorado
from Little Hills in the north to Egnar in the south.
A graph of the nightly pattern for the residents (Fig. 8) indicated that
the entire herd had entered the field by 10:00 PM and left again in a
similar manner after 3:00 AMo
Simple inspection of deer-count data from previous years and other locations
indicated that, on the average, 60-70 percent of the total of all hourly
counts was attained by midnight. This year's data concur.
Deer Distribution
In order to get an idea of deer distribution over the field for the entire
sununer, a composite of all maps collected through the sununerwas made (Fig.
9). This indicates fairly universal coverage of the field with definite
concentrations at the northwest corner, the south edge and the east end.
These are areas of access for deer as reported under segment 23
This map
also illustrates the observation that during nights of counting, few deer
frequent the vicinity of the counting station.
0

�- 231 40

PERCENT OF AVERAGE NUMBER OF DEER
~

ON DATE- OF COUNT

"
I I
,I

RIFLE GAP- 1969

I I
I I
I I
I
I
I
I

30

rfJ

,,

-

"0
Q)

C

::J

Q)

I
I
I

Legend

I

'

~

I
I

New

Seedlno

G- -

-(!] Old

61'3 6/30

7/21 "Y"28 8/11

Growth

I

I

0

u
~
Q)

,,
,
,,
,, •
,,,

~
I

20

c

--

I

0

c
o
Q)

~

ell
0..

G

I

I
I
I
I

I
I

I
I
I
I

10
8

~
6

~

't!{

4
2

•

,,
&amp;-El.

0
4/2

41'3 4/16 4/23 4/30 &amp;'7 5/12 5/19~6

Date

of

fIl

8/18

Count

Fig. 40 Graph illustrating the shift of deer concentration
from old growth
to new growth alfalfa.
(Rifle Gap Ranch, Garfield County, Colorado - 1969)0

�Table 4.

Deer-count data from new seeding and old growth on Rifle Gap Alfalfa Damage Study Field - April 2 through May 26, 1969.
Percent

Mountain Standard Time

Date

1900
New Old

4/2
4/9

16

24

4/16

2200
New Old

2300
New Old

2400
New Old

0100
New Old

0200
New Old

0300
New Old

0400
New Old

0500
New Old

of
Average
New Old

2000
New Old

2100
New Old

2

70

1

8

5

17

o

15

1

8

o

13

o

9

o

10

o

7

o

6

1.6

11.9

9

33

8

19

11

27

17

30

11

38

16

12

4

24

11

26

6

32

o

14

18.0

27.9

8

44

10

37

8

48

5

49

5

34

5

41

12

35

4

17

3

28

13

16

13.3

38.4

4/23

7

o

24

5

22

23

30

3

18

7

20

12

14

o

15

o

13

21

9

6

3

o

28.8

7.7

4/30

o

o

28

12

14

18

10

6

7

3

12

13

7

3

11

2

4

4

5

1

o

o

16.2

6.2

5/7

o

o

13

o

2

6

5

11

6

o

3

11

9

o

o

9

o

8

3

o

8

o

8.2

4.6

5/12

o

o

7

o

9

o

2

2

10

3

o

2

10

o

4

4

6

5

5

o

o

o

8.7

1.6

5/19

o

o

o

o

5

1

2

o

3

2

3

o

o

3

5

5

o

5

o

o

o

o

2.9

1.6

5/26

o

o

o

o

4

2

2

o

o

o

1

o

5

o

o

o

1

o

o

o

o

o

2.6

0.2

4.9

3.2

15.1 11.3

9.3 12.3

10.9

7.5

8.5

9.2

6.4 10.0

5.1

7.7

3.9

3.7

Percent
of
Average

12.4 11.9

12.4 11.9

10.9 11.3

N
W
N

�- 233 40

PERCENT

OF AVERAGE NUMBER OF DEER
ON

DATE OF COUNT

RIFLE GAP - 1969

30

-

"0
Q)

c:
:J
0

U

...

Q)
Q)

20

a

0

.•..
c:

Q)
(.)

...

Q)

0..

Winter

Herd

10
8
6

•

4
2

4/2

4/9 4/16 4/23 4/30 5/7 5/12 5119 5/26

Date

of

~

619 &amp;130 ~I

7/Z8 8/11 8/18

Count

Fig. 5. Graph of the seasonal peak of deer numbers expressed in percent of
average. (Rifle Gap Ranch, Garfield County, Colorado ~ 1969).

�- 234 -

Table 5. Deer-count data from Rifle Gap Alfalfa Damage Study Field - April
2 through August 18, 1969.

2000 2100

2200

2300 2400

0100

0200

0300

0400

0500

Percent
of
Average

17

9

22

15

9

13

9

10

7

7

6.3

42

27

38

47

49

28

28

37

38

14

19.0

52

47

56

54

39

46

47

21

31

29

22.7

Mountain Standard Time
Date

1900

4/2
4/9

40

4/16
4/23

7

29

45

33

25

32

14

15

34

15

2

12.3

4/30

0

40

32

16

10

25

10

13

8

6

0

7.8

5/7

0

13

8

16

6

14

9

9

8

3

8

406

5/12

0

7

9

4

13

2

10

8

11

5

0

3.4

5/19

0

0

6

2

5

3

3

10

5

0

0

1.7

5/26

0

0

6

2

0

1

5

0

1

0

0

0 8

6/2

0

2

4

0

6

0

0

2

6

0

0

1.0

6/9

0

5

4

4

5

5

4

9

7

0

0

201

6/30

0

0

6

6

6

4

6

6

4

1

0

1.9

7/21

0

2

6

6

3

8

2

1

1

2

0

105

7/28

0

0

7

14

5

10

6

4

4

2

0

2.5

8/11

1

13

11

16

10

11

12

13

12

17

2

5.8

8/18

4

6

14

14

13

21

10

16

19

6

6.6

0

�- 235 40

PERCENT OF AVERAGE NUMBER OF DEER
ON HOUR OF COUNT
NEW SEEDING VS. OLD GROWTH
RIFLE GAP-1969
30

Legend:

0----0 New Seedino
EI- - -{!] Old Growth

-

"0

Q)

c:

~

0

U

~

Q)
Q)

20

CI
0

+-

c:
Q)
o

~

Q)
a.

10

,,

8

~
6
4
2
0
1900 2000

2100

2200

2300

2400

0100

Mountain Standard

Time

0200

0300

0400

0500

Fig. 6. Comparison of the pattern of deer numbers per hour in old and new
growth alfalfa. (Rifle Gap Ranch, Garfield County, Colorado - 1969).

�- 236 -

Table 6
Deer-count data from Rifle Gap Alfalfa Damage Study Field April 2 through April 30, 1969
0

0

Date

1900

4/2
4/9

40

4/16

Mountain Standard Time
2200 2300 2400 0100 0200

0300

0400

0500

2000

2100

17

9

22

15

9

13

9

10

7

6

42

27

38

47

49

28

28

37

38

14

52

47

56

54

39

46

47

21

31

29

4/23

7

29

45

33

25

32

14

15

34

15

2

4/30

0

40

32

16

10

25

10

13

8

6

0

57

13 1

1107

12.0

11.0

11.2

81

82

80

7.1

37

Percent
of
Average

0

0

0

0

0

0

�- 237 -

Table 7. Deer-count data from Rifle Gap Alfalfa Damage Study Field May 7 through August 18, 19690
Mountain Standard Time
2200 2300 2400 0100 0200

0300

0400

0500

9

8

3

8

10

8

11

5

0

3

3

10

5

0

0

0

1

5

0

1

0

0

0

6

0

0

2

6

0

0

4

4

5

5

4

9

7

0

0

0

6

6

6

4

6

6

4

1

0

0

2

6

6

3

8

2

1

1

2

0

7/28

0

0

7

14

5

10

6

4

4

2

0

8/11

1

13

11

16

10

11

12

13

12

17

2

8/18

4

6

14

14

13

21

10

16

19

6

Percent
of
Average

0.8

11.3

13.0

11.3 1101

1202

11.1

11.6

7.7

206

Date

1900

2000

2100

5/2

0

13

8

16

6

14

9

5/12

0

7

9

4

13

2

5/19

0

0

6

2

5

5/26

0

0

6

2

6/2

0

2

4

6/9

0

5

6/30

0

7/21

7.2

�- 238 40

PERCENT OF AVERAGE NUMBER OF DEER
ON HOUR OF COUNT
WINTER HERD
RIFLE GAP-1969

30

~Q)

c:

::J
0
0

~

Q)

20

CJ)

-C

0

c:
o
CJ)

...

CJ)
a,

10
8
6

4
2

1900 2000

2100

2200

2300

Mountain

2400

0100

Standard

Time

0200

0300

0400

Fig. 7. Graph illustrating the pattern of deer numbers per hour for the
winter herd.
(Rifle Gap Ranch, Garfield County, Colorado - 1969).

0500

�- 239 40

PERCENT OF AVERAGE NUMBER OF DEER
ON HOUR OF COUNT
RESIDENT HERD
RIFLE GAP- 1969

30

.,

-

."

c:::
;:,
0

(.)

.,.,...

20

0

0

-.,.,
c:::

...

o

11.

1900

2000

2100

2200

2300

2400

0100

Mountain Standard

Time

0200

0300

0400

Fig. 8. Graph illustrating the pattern of deer numbers per hour for the
resident herd. (Rifle Gap Ranch, Garfield County, Colorado - 1969).

O~OO

�~

....•. "-. .
" ,,,--X-'
. . . ... .....
,.. .,
.
. .'. .,

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:.

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•
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.
.
.
.
..
: ...
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: .. ,
, . ~--\.
.•..
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,--,,'
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.
.
•
•••••
•
•. . ..
.
'
•
• .'
..... .
. ::... '.' '... . .. .:.--.-.~.,

..•

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•

•
•

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·

••

lC__

,

__

'.

••

•• ':
:: ••••
1
..•
..••••
: ••• I

Leaend:

Group of one or more deer
Boundary of new seeding
- - - Boundary of olfalfo field

:: ••__e.!:" '"

~

Fig. 9. Map compiled from the entire season's data indicating the distribution
groups of deer.
(Rifle Gap Ranch, Garfield County, Colorado - 1969).

I

on the study field of

N

+'o

�- 241 -

Breaking the distribution down irttoperiods of time, Fig. 10 illustrates
that the majority of the deer remained in the old growth and perimeters of
the new growth until plants reached a certain maturity.
Fig. 11 pictures the same thing for the winter herd. Fig. 12 indicates a
more even distribution by the residents. Individual deer of this herd also
showed less fear of the counting station.

Conclusions
I.

There is something about young alfalfa plants that is unpalatable to
deer.

2.

There is a resident herd in this area.

3.

Peak visits followed an overall pattern similar to that of last year,
though not as exaggerated.

METHODS AND MATERIALS - LITTLE HILLS
Study Field
Preparation of Field
Final preparation for seeding the study field was begun April 22. The
field was disked twice and harrowed with a spike-tooth harrow once to
break up the clods. Leveling was done with the local Soil Conservation
Service District's land-plane drawn by a D-4 Caterpillar crawler. A
mixture of Ladak alfalfa (15 lb./A.) and Colorado 37 oats (30 lb./A.)
was sowed using a John Deer tractor, model 620, and l2-foot drill between
May 26 and 29.
Irrigation
Additional pipe was purchased and the mainline permanently installed the
full length of the field along the south edge, between May 27 and June 3
(Fig. 13). One section of 3,600 feet (60 T-valves) extended to the east
above the well, another of 3,840 feet (64 T-valves) to the west below the
well. The feeder line from the well was fitted with a reversible elbow
permitting water to be turned into either section.
Five lateral lines were spaced along the mainline in one section at a time.
East of the well there were 12 T-valves between each lateral, west of the
well there were 13 except at the end. A pipe trailer was used to transfer
lateral lines from one section to the other.

�~

.

lI._,,_lt-;c,-X-X-;e
___-'.!L;-X-

0

!.:....

.'

xI 0'.
\...
'.

___ .

•

Station

':'

.'

xI

I

)(

I

)C

"

i

(/)1

I

,

i·

i.'

;-.. ....•... :..,.--•••

0

~

0

:.

.,

0:.

:,'f

:.

•

:
0

•

0

i

i
~.

~••. ~'"

,

..•.-

· · " - ... ~ ~.,'"
·/

I
I
I

I

I

I

- - -

Legend'

-

-

-'

~,

- - - -

or

r

:.

:0'
: G'

.,0'0,

: -.' - :..

J

:'

~-----....

of

:'

•• • •

,. I'·

~~~-

_

0

.,.

/~-

..".. ",

•
0

,.:

•

. .
'.
. . . ....

..
.

,

" .

••

•
\,..'
\.
\

0

,.,~,'

• •

••

, •

I ,.
x.'

:

0'.

\

P~mp'"!1

•• 1,':
1

Yo,'

•

~

o.

v •
~

lI •

:

::

•

)(-.0;;1(_
)(1"-)(

! .! :

I

,

~~

'"

:
•I 0 0
I 0

-IC_

-'
• Group of one or more deer
Boundary of new seeding
Boundary of olfa If a field

, - -"

Fig. 10. Map indicating the distribution on the study field of groups of deer until deer concentration
shifted from old growth to new growth. (Rifle Gap Ranch, Garfield County, Colorado _ 1969).

N

+-N

�~

~"'_ I

,,_ It_,,_x-x-It

~

---.

.

I••.

•
• ••••

'.
.\
,.

. . .

...

,

-"

-)C_

•

,.-.-

" ....•

.::
,. .,.,
.
. .. ..:..... :,
..: . .,
. . '... ....'.,

x_ x_

;.

.1

• :

'1

••
!..

I

.::.'

•f

:.:.

of

o

•

•
-:r'\---

•

Pumping
• Station

, .

•

•
I '.
\
•

\

• I
•I

... -

/

,
I

, .... .;.
0

0

\\

\

..
•

:
:

.

. .....

o·
'

,....,.:..
.. .':.::.:;.

:'"
--"

• •

'~'':-';

. . e.. '
.
....
~::
:. --'

-

..••
!..--"'~

"

_...

•..... ...

.

_ •••_

__

•••
__ .:
t
-- ••
..,,-'
...• _------

Legend:
Group of one or more deer
Boundary of new seeding
Boundary of alfalfa field

~

Fig. 11. Map indicating the distribution on the study field of groups of deer only for the winter herd.
(Rifle Gap Ranch, Garfield County, Colorado - 1969).

N

+=-

w

�~

___

-

:\..

••
..-•••
W'J :;;;0-)(

- ••••..z

_)(

-

x-rr:

)C._x

••

-

..)C_

)L-1&lt;_

J(_
•

J(_

:

•• •

:

• 0

o

•

o •

o •

o.

....

.. . ..

;

•

•••

:

••

..

••

0

•:"

t :.

00

•

•••
•

••••

•

... .

.

••

••

--.~
I

...~_

)(

__

_ .•
.

." -

.•.
_

_ o

•

~
'" _. ,. ••
'.'

..'

0

•
I.'.
.:
I.

o
o

I

I

..-:

.:

.

•••
:

• ••! •.

•

•

0

'

,

I

I

,

t

N

~

x

o·

o

k .•...•...•
Statia.
.
:.

I

i
I

•

-.-X-=:.:)(•..-.

X;-o.•••••

• I

••

,.

)(

X--k __

•

\
\

•

.:

\
\
\ •
\

0

\.

.." .'

..•. _i...--.; .•.._
•

0

.

••

•• •••• •
•

••

oo •

•

o

• • ·0
•

•
•

.

I

...~

0

..... ; .;:...,--:., ". '

.:-:

" .",,:,

,
'"

.

-- -

I

I

0

••• ••••• •

•

- ---'

I

I

\

- - - - - ."

• !:.!sI.!M:

- •'.

•

• '.

.'

.•••
•••
__•
-_

'; !

I

.'
I
•:
!
•
•I
-_ •••
0
JI
~.•••
•••- J- - - -

- --

• Group of one or more deer
Boundary of new seeding
Boundary of alfalfa
field

Fig. 12. Map indicating the distribution on the study field of groups of deer only for the resident
herd.
(Rifle Gap Ranch, Garfield County, Colorado - 1969).

~
~

�CORRAL
PASTURE

GULCH

8

PASTURE

PASTURE

7

6

---------- -----------------",
/\.
"

-.~
.

..,.j'---&gt;-

\

/

"'~

~ 'r'e'~.t
.•.::---"'"\.
/
...,.,

- - --------

I
I

----

~-",

Legend:
r:;;::1

~

___

====t~ I

Direction

of

line

moves

- - - -'

Boundary

/

lateral

of alfalfa
IV

-I'V&gt;

Fig. 13. Map of Little Hills Alfalfa Damage Study Field showing major features and the pattern used when
changing lateral lines. (Meeker, Rio Blanco County, Colorado).

�- 246 -

Irrigation was begun on June 4. Sprinkler lines were changed every 12
hours during June and July, every 24 hours during August. The pattern
of change is diagrammed in Figo 140 The lateral lines were carried 120
feet down the mainline, skipping one T-va1ve each time, until the first
setting of the next lateral was reached. This process was then reversed
on the way back using T-va1ves that had been skipped on the way down.

RESULTS AND DISCUSSION - LITTLE HILLS
Segment objective 1 was attained this year. Objectives 2 and 4 will not
be complete until the study is finished. Work on objective 3'cou1d not
be done due to adverse weather conditions and related complications.
Work on objective 4 can not begin until plots are pairedo All work was
performed by Little Hills personnel.
Study Field
Preparation of Field
Preparation and planting of the field would have been completed earlier
except for a delay in obtaining the land-plane.
Irrigation
The procedure, as outlined above, permitted one complete irrigation of
the field every 13 days when pipes were changed at 12-hour intervals.
This pattern obtained the most uniform germination of the seed. The
entire field was irrigated completely three times through the summer.
Hay Harvest
The field was thoroughly and evenly seededo Growth was not uniform all
over the field, however, due to differences in soil fertility, drainage
and irrigation coverageo
The original plan was to harvest the oats by combine. Stubble would have
been baled for easy removal. This plan was abandoned because heavy snow
matted the oats to the ground the week before harvest was scheduled. Subsequent moisture prohibited any machinery from entering the field.
As an emergency measure 600 head of cows and calves belonging to Mr. Ivo
E. Shults grazed the field between November 17 and December 10
This
appears to have accomplished its end by removing approximately 90 percent
of the oat cover. The true effect will be learned next spring.
0

Study Plots
Inclement weather which prohibited harvest also cancelled fencing of the

�N

Legend:

f

+

Sprinkler ...head
Pipe joint

-e- Tee-valve
-+- Pipe joint

Lateral

Line
N
.po.

--.J

----

Mainline

Order and direction of lateral line changes

Fig. 14. Diagram of the order followed when changing lateral lines on Little Hills Alfalfa Damage
Study Field, Meeker, Rio Blanco County, Colorado.

�-

248

-

control meadow and marking of the study plot cornerso
be done next spring.

These tasks will

Deer Numbers
Deer counts were not attempted because the study field remained fallow
until after the deer had left for the summer.

Prepared by

iJ~

y:? ~

David F. Gordon
Asst. Wildlife Researcher

�- 249 -

APPENDIX I

��July, 1970

- 251 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-24

Deer-Elk

Investigations

Work Plan No.

11 B &amp; C

Job No.

1

Job Title

Physical Characteristics
White River and Rio Grande Elk

Period Covered:

April 1, 1969 through March 31, 1970

Personnel:

Raymond

J. Boyd

ABSTRACT

Due to reassignment to other jobs within this project, this job was
not completed, a rough draft of the breeding dates paper was finished,
however.

��- 253

-

PHYSICAL CHARACTERISTICS - WHITE RIVER AND RIO GRANDE ELK
Raymond J

0

Boyd

Po So OBJECTIVE
To publish data on elk physical characteristics.
SEGMENT OBJECTIVES
1.

Publish in the Journal of Wildlife Management an article comparing
physical characteristic differences in two elk herds that winter on
two different vegetative typeso

2.

Publish in the Journal of Wildlife Management an article on the
breeding dates of Colorado elk based upon fetal measurements.
METHODS AND MATERIALS

1.

Gather all comparative physical measurement data from the White River
elk herd and the Rio Grande elk herd and show differences attributed
to nutrition and complete a paper on the differences.

2.

Get together all fetal measurement data and write a paper on the
breeding season of Colorado elk based upon crown/rump measurements
of fetuses collected in December.
RESULTS AND DISCUSSION

Due to reassignment to other jobs within this project, this job was not
completed during the past segment. However, a first draft of the breeding
dates paper was completed.

��July,

1970

- 255 -

JOB FINAL

S ta te

of

Project
Work
Job

.:::C,:::;O.:::L,:::;ORAD:..=.::::.O:::...-_-..:..:...-._
No.

Plan

REPORT

No.

W-38-R-24

Deer-Elk

.=I.=l_B=--

_

Job

No.

Investigations

7

Ti t Le _......:;l\..:i;.;::o~G:.:r..::a:.::n:.::d:.::e:....-::E:.:1:.::k::......::S:...:t:..:u:.::d:.!y_-__=P....::u:.::b:...:l::i:..:c:.::a::.:t:.:i:..:o:.:n::......::o::f~R
_

Period
Personnel:

Covered:

None
Raymond

J.

Boyd

ABSTRACT

This
job was completed
and a final
report
submitted.
advertently
included
in this
segment.
The final
report
in the July,
1969,
(Part
One) Game Research
Report.

It

was inis published

��JU.lY,

.l:1/U

- 257 -

JOB PROGRESS

REPORT

State of

COLORADO

Project No.

W-38-R-24

Deer-Elk

Investigations

Work Plan No.

llc

Job No.

1

Job Title

White River Elk population

Components

Period Covered:

April

Personnel:

Edgar J. Prenzlow.
Check Station Assistant's Richard
M. Bartmann, George D. Bear, RaymondJ.
Boyd,
David
F. Gordon, Donald M. Hoffman, Richard M. Hopper,
Roland C. Kufeld, Gary T. Myers, Robert L. Schmidt,
Donald G. Smith, Steven F. Steinert, Harold M. Swope,
and Michael R. Szymczak.

1, 1969 through March 31, 1970

ABSTRACT

This is the fourth annual progress report of a five-year investigation
to determine the effects of specified permit seasons (limited number of
both antler1ess and antlered permits available to hunters) on the White
River elk herd in northwestern Colorado.
Estimated herd size, determined
by formulas using pre- and post-season sex and age ratios in combination
with harvest levels, has decreased from about 6,800 to 5,800 elk from
1966 to 1969. Pre-season sex ratios (bulls per 100 cows) and age ratios
(calves per 100 cows) were increased from about 29 to 50 and 58 to 60,
respectively.
The change in sex ratios resulted from issuing more
antlerless and fewer antlered permits to hunters in an attempt to
stabilize the female segment of this elk herd. Methods used to determine
harvest rates, sex and age ratios, and age structures of this elk herd
are compared and evaluated.
Also, data from check stations, hunter
report cards and random surveys are included.

��- 259 -

WHITE RIVER ELK HERD - POPULATION

COMPONENTS

Edgar J. Prenzlow

P. S. OBJECTIVE
To develop

a harvest

formula

for the White River elk herd.

SEGMENT OBJECTIVES
1.
2.

3.

4.

Review literature necessary for population analysis of the White River
elk herd.
Summarize, evaluate and file all previous information collected on the
White River elk herd to better acquaint new project personnel on the
history of this job.
Test a candidate elk harvest formula for Area E elk herd.
a. Maintain wintering population at about 5,000 elk by stabilizing
female segment of herd.
b. Narrow post-season sex ratios to 20 bulls per 100 cows in an attempt
to increase productivity
(calves per 100 cows) by specified permits.
Determine accurate sex and age structures of the harvest, hunter success
ratios, and amounts of recreation and associated recreation days expended
on the White River elk herd.

METHODS

AND MATERIALS

Data collection (aerial and ground sex and age ratio counts, check stations
and mail questionnaires
to determine harvest, age composition, recreation
days, etc.) for this project has not varied appreciably since its instigation
in 1966 (Prenzlow 1967, 1968). Likewise, procedures for determining total
population estimates have not changed.

DESCRIPTION

OF AREA

Physiographies of the White River area are included in Harris (1963) and Boyd
(unpublished).
Their manuscripts are concerned with all eight Game Management
Units (12, 13, 23, 24, 25, 26, 33 and 34) whereas this present investigation
is limited primarily to Area E, or Game Management Units 23 and 24 (Fig. 1).
Prenzlow (1967, 1968) also described the study area.

�- 260 -

••"T9

.Jl~

B

Fig. 1. Elk Management Area E (Game Management Unit~
23 and 24) showing boundaries, drainage patterns and
highway systems.
Solid c.ir~lee are active check
station locations.

STUDY

AREA

LEGEND
GAME

MANAGEMENT

UNIT

OOUNDARY

IIOHWAY=="""
OECONDARY ROAO ---IlIVERCII[EK
------&lt;::::::

Check Stations.

-.

�- 261 -

RESULTS

AND DISCUSSION

Sex and Age Composition

Aerial

Surveys

Pre-Season Sex and Age Ratios--The
1969 pre-season sex and age ratio
counts were flown in a helicopter on 29, 30 September and 1, 2 and 5
October.
These classifications
were confined to elk observed within
Area E (Table 1). A total of 1,358 elk were classified resulting in
a ratio of 50.2 bulls (including spikes) and 60.3 calves per 100 cows.
Summaries of past sex and age ratios for the years 1961 through 1969
are shown in Table 2. while Figs. 2 and 3 illustrate 95 percent confidence intervals for these ratios.

Table

Pre-season

1.

sex and age ratio

Mature

Location

Ripple Cr., Picket Pin
and Trappers Lake
Oak Ridge,

Big Beaver

Cr.

counts

in Area E, 1969.

Spikes

Cows

Calves

Total

13

31

103

45

192

2

14

27

14

57

Bulls

Miller

Cr. and Elk Cr.

2

2

13

10

27

South

Fork Drainage

41

34

158

110

343

25

34

139

89

287

Lost Park and Snell Peak

30

96

205

121

452

Total

113

211

645

389

1,358

Percent

8.3

15.5

47.5

28.7

100.0

Marvine Crs.,
Big Ridge

Ute Cr. and

�Table 2.

Summaries of pre-season sex and age ratios from Area E, 1961-1969.
Bulls

Cows

Calves

Year

No.

%

No.

%

No.

%

Total

Bulls

:

Ratio
Cows

:

Calves

1961

128

19.6

309

47.3

216

33.1

653

41.4

:

100

:

69.9

1962

216

22.1

462

47.4

298

30.5

976

46.8

:

100

:

64.5

1963

125

20.0

298

47.5

204

32.5

627

41.9

:

100

:

68.5

1964

120

17.5

363

52.9

203

29.6

686

33.1

:

100

:

55.9

1965

233

17.3

699

51.8

416

30.9

1,348

33.3

:

100

:

59.5

1966

86

15.4

299

53.7

172

30.9

557

28.8

:

100

:

57.5

1967

134

16.8

415

52.2

247

31.0

796

32.3

:

100

:

59.5

1968

135

18.7

366

50.7

221

30.6

722

36.9

:

100

:

60.4

1969

324

23.9

645

47.5

389

28.6

1,358

50.2

:

100

:

60.3

Ave.

167

19.5

428

49.9

263

30.6

858

39.0

:

100

:

61.4

N
0'\
N

�-

263

-

60

••• 50
~
0
~I

0
0

.!!

40
30

:I

co

20
0
1961

6~

63

64

65

66

67

68

69

70)

Years

Fig. 2.

Confidence intervals on pre-season sex ratios, Area E, 1961-1969.

,..
80

••• 70
~
0

U
0
0

---

60

50
•••
G)
&gt;

0

U

40
0
J961

62

63

64

65

66

67

68

69

710

Years

Fig. 3. Confidence intervals on pre-season age ratios (ca1ves:100 cows),
Area E, 1961-196,.1/

so
•••40

~

o

U

30

o
o

..:::..
20
.!!

:J

co

10

o L----19-6~,-----6~2----~63~----6-4-----6~5-----676----~6~7-----6~8~---76~9----~70
Years

Fig. 4.

Confidence intervals on post-season sex ratios, Area E, 1961-1969.11

1/ Confidence intervals were calculated ate( = 005 using a formula from
Riney (1956) as f&amp;l1~
100 calves/lOO cows + (100 Y!(calves + cows) calves/cows3)

�- 264 -

Post-Season Sex and Age Ratios--Post season sex and age ratio counts
were made in Area E on 1-3 December.
A total of 1,839 elk were classified resulting in a ratio of 16.4 bulls (including spikes) and 65.3
calves per 100 cows (Table 3). Summaries of prior post-season
sex and
age ratios for the years 1961 through 1969 are shown in Table 4. Confidence intervals oh post-season
sex ratios appear in Fig. 4.

Table

3.

Post-season

sex and age ratio

counts

in Area E, 1969.

Location

Bulls

Spikes

Cows

Calves

Total

Nine Mile

4

16

96

53

169

Uranium

0

4

34

25

63

7

37

197

134

375

Oak Ridge

7

20

102

69

198

Miller

3

6

33

22

64

Sand Pk. and east of Big
Beaver Cr.

5

23

196

130

354

South

3

31

354

228

616

Total

29

137

1,012

661

1,839

Percent

1.6

7.4

55.0

36.0

100.0

Pk.

Sawmill and west
Beaver Cr.

Ground

of Big

Crks.

Fork

Surveys

In 1967, 1968 and 1969 elk were classified by ground surveys during July.
This was done to estimate yearling-adult
cow ratios which can not be obtained
during aerial classifications
and also served as independent estimates of sex
and age structures.
It is important to determine the proportion of yearling
females in the herd and their influence on overall herd productivity.
Usually
only a small portion of yearling females breed, but there is considerable
conflicting literature on this subject.
Ground survey data are being reported
in a Game Information Leaflet.
Just the 1969 data have been tabulated for
this report.

�Table 4.

Summaries of post-season sex and age ratios from Area E, 1961-1969.

Year

No.

%

No.

%

No.

%

Total

Bulls

:

Ratio
Cows

:

Calves

1961

106

10.6

465

46.5

430

43.0

1,001

22.8

:

100

:

92.5

1962

125

13.8

446

49.2

335

37.0

906

28.0

:

100

:

75.1

1963

45

6.1

413

55.7

283

38.2

741

10.9

:

100

:

68.5

1964

117

6.0

1,130

58.3

692

35.7

1,939

10.4

: .100

:

61. 2

1965

85

5.2

930

57.1

613

37.7

1,628

9.1

:

100

:

65.9

1966

128

6.1

1,245

58.9

739

35.0

2,112

10.3

:

100

:

59.4

1967

214

9.2

1,318

56.4

806

34.5

2,338

16.2

:

100

:

61.2

1968

148

8.2

951

52.9

699

38.9

1,798

15.6

:

100

:

73.5

1969

166

9.0

1,012

55.0

661

36.0

1,839

16.4

:

100

:

65.3

Ave.

126

7.9

879

55.3

584

36.8

1,589

14.3

:

100

:

66.4

Bulls

Calves

Cows

N
0\
V1

�- 266 -

Observers on horseback, riding during the early morning and evening hours,
classified elk in different sections of the Flat Tops Primitive Area.
In
four days, two teams of two or three men classified a total of 969 elk in
July, 1969 (Table 5). Many more elk were seen on which classification
was
not possible.
The average size of elk groups observed was 32.3 animals.

Table 5. Composition
in Area E,1969.

of 969 elk classified

Mature
Bulls

Yearling
Bulls

during

Mature
Cows

summer

ground

counts

Yearling
Cows

Calves

Total

Number

Observed

52

65

440

133

279

969

Percent

Observed

5.4

6.7

45.4

13.7

28.8

100.0

16

17

28

22

20

30

21 1.7

2.2

14.7

4.4

9.3

32.3

(e.g. mature

bulls

observed

in 16 of 30 total

Frequency

in Groups

Mean

Composition

Group

1/

1/ Frequency
groups).

in groups

21 Mean group

size determined

Check

Surveys

Station

by percent

were

of elk observed

times

32.3.

Number of Elk Checked--Four
White River elk check stations (Meeker, New
Castle, Deep Creek, Ripple Creek) were operated for the first nine days
during the 1969 big game season.
Research personnel at the permanent check
station at Rifle also checked elk from White River units.
A total of 983
elk were checked from Game Management Units 12, 13, 23, 24, 25, 33 and 34.
Comparable data from other White River units also were collected because of
their importance in relation to Area E elk.
Table 6 lists the average number
of elk checked at each station from 1964 through 1969.
Table 7 lists checkout by date of 673 elk from Area E.
cent were checked through stations on the third through
the season.

More than 85 perthe seventh day of

�- 267 -

Table
herd,

6. Number of elk checked through stations from.White River elk
(Game Management units 12, 13, 23, 24, 25, 33 and 34) 1964-1969.

station

1964

1965

1966

1967

1968

1969

Average

Meeker

553

506

591

518

725

637

588

Deep Creek

338

227

220

303

308

139

256

New Castle

321

196

166

227

290

78

213

Rifle

137

138

124

92

83

73

108

32

137

159

112

120

129

Idaho Springs
Ripple

Creek

227

56

1,381

1,204

1,384

133
4

4

Hamilton

Total

110

1,381

1,633

983

1,328

Sex and Age Composition of Elk Checked--Percent
composition
(bulls, cows
and calves) of elk checked is presented in Table 8. Sex and age ratios
of the 673 were 77.0:100:8.9.
This compares with 126.8:100:21.2
in 1966,
108.1:100:32.1
in 1967, and 95.6:100:26.1
in 1968.
During the first three
years a constant number of antlered permits (2,500) were issued, but in
1969, antlered permits were reduced to 2,200.
Ant1er1ess permits varied
from 1,000 in 1966, to 1,500 in both 1967 and 1968 and to 1,000 again in
1969.
These changes partially account for the differences
in ratios.

Age Structure of Harvest--Of
673 elk checked, 437 were aged from Area E
with the technique described by Quimby and Gaab (1957).
Table 9 lists
these ages by sex, while Table 10 shows similar information by game management unit.
Teeth from 491 of the 653 elk aged (all units) were collected and taken to
the wildlife Laboratory at the Fort Collins Game Research Center.
Here the
teeth were again aged according to the cementum technique reported by Keiss
(1969).
Assuming ages as determined by the cementum method to be correct,
this comparison revealed an error of 1.6 percent on yearlings and 64.0
percent on two-year old and older elk (Tables 11 and 12) between the two
methods.

�Table

10.

Sex and age determined at check stations of 437 elk by unit of kill, Area E, 1969.

Age (Years)
~

l~

2~

3~

~

5~

6~

7~

8~

9+

Total

Number

0

37

12

0

2

2

0

1

0

0

54

% of Total

0

68.5

22.2

.0

3.7

3.7

.0

1.9

.0

.0

100.0

14.1

4.6

.0

.8

.8

.0

.4

.0

.0

20.7

118

41

Unit and Sex

23 - Males

% of all Males

.0

24 - Males
Number

7

24

11

6

1

0

a

1

209

% of Total

3.3

56.4

19.6

11.5

5.3

2.9

.5

.0

.5

.0

100.0

% of All Males

2.7

44.8

15.6

9.1

4.2

2.3

.3

.0

.3

.0

79.3

Number

2

4

11

8

4

2

4

1

3

1

40

% of Total

5.0

10.0

27.5

20.0

10.0

5.0

10.0

2.5

7.5

2.5

100.0

% of all Females

1.1

2.3

6.3

4.6

2.4

1.1

2.4

.6

1.7

.6

23.1

Number

16

19

18

15

19

7

18

6

7

9

134

% of Total

12.0

14.2

13.4

11.2

14.2

5.2

13.4

4.5

5.2

6.7

100.0

% of all Females

9.2

10.9

10.3

8.7

10.9

4.0

10.3

3.4

4.0

5.2

76.9

23 - Females

24 - Females

N
-..J

0

�- 271 -

Table II. Percent error in aging 183 yearling elk at check stations, all
White River Units, 1969.

Station

Dental
Cementum 1/

Field
2:../
Aged

Percent
Correct

Percent
Error

Meeker

125

122

97.6

2.4

Ripple Creek

8

8

100.0

0.0

New Castle

22

22

100.0

0.0

Deep Creek

12

12

100.0

0.0

Rifle

16

16

100.0

0.0

Total

183

180

98.4

1.6

1/ Number of teeth checked in laboratory using cementum technique.
2:../ Number of correct ages determined at check station using tooth replacement
and wear technique.

Table 12. Percent error in aging 308 elk older than yearlings at check
stations, all White River Units, 1969.
Percent
Correct

Percent
Error

79

37.4

62.6

16

9

56.3

43.7

New Castle

24

8

33.3

66.7

Deep Creek

34

5

14.7

85.3

Rifle

23

10

43.5

56.5

Total

308

111

36.0

64.0

Station

Dental
Cementum 1/

Field
Aged 2:../

Meeker

211

Ripple Creek

1/ Number of teeth checked in laboratory using cementum technique.
2/ Number of correct ages determined at check station using tooth replacement
and wear technique.

�- 274 -

Table 15. Composition
Game Management Unit.

of the 1969 estimated elk harvest
Data are from report card survey.

Unit

Antlered
No.

Antlerless
No.

%

in Area E by

Total
No.

%

%

23

429

25.4

398

23.5

827

48.9

24

409

24.1

456

27.0

865

51.1

Total

838

49.5

854

50.5

1,692

100.0

Table 16. Number of resident and non-resident hunters, estimated elk kill
and success ratio in 1969, Area E. Data are from report card survey.
Number of
Hunters

Success
Ratio

Resident

3,554

31.0

547

556

1,103

Non-Resident

1,225

48.1

291

298

589

Total

4,779

35.4

838

854

1,692

Class

Animals Harvested
Antlered
Antlerless

Total

Random Survey--To better estimate the elk kill in Area E, 1,517 of 3,200
permittees (47.4% sample) were contacted by letter and requested to give
information regarding their hunt. Appendix A is a replica of the stamped,
pre-addressed postcard sent to each hunter in the sample.
Of 1,517 hunters sampled, 1,190 (79.4%) returned usable cards. The estimated total harvest was 1,183 (c.r.± 127 2 1.96) elk with 37.0 percent of
the hunters being successful.
Table 17 gives results of the survey, including harvest by type of hunter, while Table 18 lists active (those who
actually hunted elk) number of hunters and success ratios.
Random sampling resulted in a more accurate projection than the report card
survey because more than 36 percent of each type of hunter responded to the
survey; whereas, the report card method over-estimated kill by each class
of hunters.
For instance, antlerless kill by non-residents was projected
to be 298 animals, although there were only 193 antlerless permits issued.
Small sample sizes, method of projection (success ratios of hunters with
antlered and antlerless permits are assumed to be comparable), and the

�- 275 -

fact that successful hUnters return report cards at a greater rate than
unsuccessful hunters, caused the over-estimates.
Also, many hunters
with regular elk and sportsman licenses report kills in Area E.
It is
likely that kills are made in Area E by persons without the proper permit;
but it is doubtful these exceed the confidence interval of the kill estimated by the random survey.
Table 19 compares kill estimates and proportions from report cards and random surveys by residency and type of
permit.

Table

17.

Results

of 1969 random

survey

of 3,200 Area E permittees.

Antlered

Antlerless

Res.

N. Res.

Res.

N. Res.

Total

1,616

584

807

193

3,200

Permits

Issued

Permits

in Sample

762

266

394

95

1,517

Percent

in Sample

47.2

45.5

48.8

49.2

47.4

Sample

Returned

593

210

308

79

1,190

Percent

Returned

77 .8

78.9

78.2

83.2

78.4

Percent

of Total

36.7

36.0

38.2

40.9

37.2

Permits

Reported

"No Hunts"

103

13

39

6

161

Estimated

"No Hunts"

281

36

102

15

434

Reported

"Kills"

136

86

171

51

444

Estimated

Total Harvest

371

239

448

125

1,183

Table 18. Number of hunters participating
during
and success ratios per license and active hunter.
survey.

Antler less

Antlered
Resident

Active

Hunters

1969 Area E elk season
Data projected from random

Non-Resident

Resident

Non-Resident

Total

1,335

548

705

178

2,766

Success

(Kill)/Lic.

23.0

40.9

55.5

64.8

37.0

Success

(Kill)/Hunter

27.8

43.6

63.5

70.2

42.8

�- 276 -

Table 19. Total resident and non-resident projected elk kill by type of
animal based on report cards and random survey, Area E, 1969.

Survey

Report

Card

Random

1/ Figures

Resident
Antlered
Antlerless

Non-Resident
Antlered
Antlerless

No.

547

(33.8)Y

556

(68.9)

291

(49.8)

298

(154.4)

1,692

(52.9)

371

(23.0)

448

(55.5)

239

(40.9)

125

(64.8)

1,183

(37.0)

success

ratios

in parentheses

indicate

per license

Total
Percent

issued.

Projections from the random survey indicated that 434 of 3,200 (13.6%) licenses
were not used.
The number of "no hunts" in Area E has averaged 7.0 percent
since 1966.
Other surveys by Prenz10w (1969 a,b) on elk Areas MM and T, and
by Turner (1967, 1968) in New Mexico, reported rates of unused licenses at
2.5, 4.0, 3.4, and 2.9 percent, respectively.

Harvest

in Relation

to Area

and Time

Location of Harvest--Harvest
by Game Management Unit was projected from the
random survey and appears as Table 20. About two-thirds
(63.6%) of the
harvest was from Unit 24. Report card data (Table 15) indicated 51.1 percent of the harvest was from Unit 24.

Table 20.
Management

Composition of 1969 estimated
Unit, random survey.

elk harvest

in Area E by Game

Unit

Antlered
No.
%

Ant1er1ess
No.
%

No.

%

23

227

19.2

204

17.2

431

36.4

24

383

32.4

369

31.2

752

63.6

Total

610

51.6

573

48.4

1,183

100.0

Total

�- 277 -

Hunter Effort--The Area E elk season provided 12,749 man-days of hunting
recreation in 1969. This was a decrease of about 9,000 man-days compared
with 1967 and 6,000 man-days in 1968 (Table 21). Three major reasons
accounted for this loss of recreation days.
In 1969, 3,200 total permits were issued compared with 4,000 in both 1967 and 1968; an increase
in the number of "no hunts" (Table 17), and severe weather conditions
during the 1969 season.
Mean number of days hunter in 1969 (4.6) also decreased from 5.7 and 5.2
in 1967 and 1968, respectively.
Probably, this decrease also can be
attributed to severe weather conditions during the 20 day season.

Estimates

of Mortality

Other Than Legal Harvest

Wounding Loss--Estimates of net wounding loss (number of elk reported
wounded minus number of wounded animals harvested) are presented in Table
22; while in Table 23, a comparison is presented between wounding loss
reported by hunters with antlered and antlerless licenses.
It is assumed
these wounding loss figures represent a minimum loss because all hunters
would not admit wounding an elk. Hunters who did not wound an animal,
most likely would not respond to the question positively.
The 9.2 percent figure compares closely to 9.0 and 7.4 percent reported
by Hay et ale (1961) and Hunter (1967): and also to the 9.4 and 7.7 percent determined for Area E in 1967 and 1968. Early records by Smith
(1959) indicated wounding loss affected about 1.5 percent of the elk herd
on the White River Plateau.
Tileston (1962) determined that elk wounding
loss averaged 10.1 percent (range 6.1 to 11.7) in Colorado from 1946 to
1953.

Population

Total Population

Estimates

and Projections

Estimates

Formulas were used to estimate the total number of elk present in Area E prior
to the 1969 season.
Prenzlow (1968) described this in detail with examples of
how ratios of antlered to antler less elk (from harvest and pre- and postseason classification counts) were used in a formula devised by Dr. Bowden of
Colorado State University, to estimate pre-season populations.

Pre-Season Population Estimate, 1969--An estimated 3,393 (C.I. = 83,2 =1.96,
~ = .05) elk were present in Area E prior to the 1969 season (Table 24). This
compares to 6,751, 6,313 and 6,261 estimated in 1966, 1967 and 1968, respectively.

�- 278 -

Table 21.

Recreation day yield of specified permits in Area E, 1967-69. 1/

Year and
Type of Permit

Active
Hunters

Recreation Days
Total
Mean/Hunter

Antlered

1,761

9,662

5.5

Ant1er1ess

1,224

6,742

5.5

Antlered

616

3,745

6.1

Ant1er1ess

216

1,468

6.8

Totals and Mean

3,817

21,617

5.7

Antlered

1,668

8,720

5.2

Ant1er1ess

1,110

5,389

4.9

Antlered

671

3,583

5.3

Ant1er1ess

296

1,645

5.6

Tota Is and Mean

3,745

19,337

5.2

Antlered

1,335

6,256

4.7

Ant1er1ess

705

2,958

Antlered

548

2,717

5.0

Ant1er1ess

178

818

4.6

Totals and Mean

2,766

12,749

4.6

Resident

Non-Resident

Resident

Non-Resident

Resident

Non-Resident

1/ Data from postcard surveys and systematic random samples (approximate 50 percent)
of n = 2,072 (1967), and 1,971 (1968) and 1,517 (1969).

�Table 22. Wounded elk salvaged by hunters and net wounding loss Area E, 1969.
Woundinl:1j
Loss
Net Loss
Wounded Ki lled

Number of Elk
Reported Ki lled

No. Wounded

Antlered

222

28

9

19

8.6

Antleress

222

13

7

6

2.7

Totals

444

41

16

25

5.6

License Type

Table 23.

Percent Loss

Area E wounding loss - comparison of antlered and antlerless hunters, 1969, random survey.

of Kill

Projected Elk Loss
No.

222

12.6

77

3.4

222

5.9

34

3.4

444

9.2

111

No. of
Hunters Reporting

No. of Elk
Rept. Wounded

Wounding Elk

No. of Elk
Rept. Killed

Antlered

803

28

3.5

Antler1ess

387

13

1,190

41

License Type

% of Hunters

% Loss

IV

'"
\0
I

Totals

�- 280 -

Table 24. Pre-season population
and age in Area E, 1969.

estimate

and proportions

of elk by sex

Item

Estimated
Number

Expected
Percent

Bulls

811

19.2

23.9

Cows

1,612

50.4

47.5

Calves

970

30.4

28.6

Total

3,393

100.0

100.0

1/ Percentages

expected

from 1968 post-season

2/ Percentages
helicopter.

observed when making

population

1969 pre-season

Observed
Percent

y

y

data.

classifications

by

Until 1969, total population data were believed to be reasonable estimates
of the actual population levels in Area E. For instance, predictions for the
percentages of bulls, cows and calves were previously within five percent of
what was actually observed during pre-season aerial observations.
However,
for 1969, the expected and observed percentages of bulls differed almost 20
percent (Table 24). This was a critical difference, because the change in
ratios of bulls from pre- to post-season is the major component of the
formula used to estimate the total pre-season popUlation.
Several possible factors could have caused this discrepancy (e.g. observer
error, sampling error, etc.); but it is my best judgment, based upon the
data obtained from Area E and adjacent areas, that early and rather severe
weather conditions caused an abnormal elk migration pattern.
As a result,
the data that were obtained were from two different populations of elk and
therefore, not valid.
Aerial trend data, collected independently from this investigation, also
revealed the estimate of 3,393 was extremely low (White 1970). During midJanuary counts, a total of 3,668 elk were observed within Area E indicating
that at least 4,851 (3,668 plus estimated harvest of 1,183) could have been
counted before the season.
Therefore, to accomplish the objective of testing an elk harvest formula,
it became necessary to use the 1969 projected population figure of 5,839
(Prenzlow 1969c) rather than the 1969 estimate of 3,393. This was thought
justified because past projections, after being corrected for over-winter
loss, have been close approximations of elk total population estimates
derived by formula.

�- 281 -

Post-Season Population Estimate, 1969--An estimated 4,646 Area E elk survived the 1969 season (Table 25). This was calculated by subtracting the
elk harvest, as determined by the random survey, by class of animal from
the pre-season population.

Table 25. Post-season population
and age in Area E, 1969.

estimate

and proportions

Y

of elk by sex

Item

Pre-Season
Population

Harvest

Bulls

1,119

610

509

11.0

9.0

Cows

2,938

488

2,450

52.7

55.0

Calves

1,772

85

1,687

36.3

36.0

Totals

5,829

1,183

4,646

100.0

100.0

Post-Season POEulation
No.
Exp. % y
Obs. %

on 1969 random survey.
by class of the total estimated number of elk
remained following the 1969 season.
11 Proportions actually observed when making 1969 post-season
by helicopter.

11

1/ Based

y Proportions

(4,646) that
classification

Total post-season populations have declined from 5,336 to 4,646 elk since
1966.
This was largely the result of increased harvest of the antlerless
animals.
Allowing more antlerless permits was necessary, however, so that
the composition of the wintering herd could be changed to meet the objective
of narrowing the sex ratio (more bulls per 100 cows).
Proportions
of bulls
have increased in the post-season populations
from 6.1 to 9.0 percent.

POEulation

Projections

Each year, based on the last post-season population estimate, a projection
is made for the next year.
Prenzlow (1968) described this procedure in
detail using 1967 data as an example.

POEulation Projection for Area E, 1970--Prior to the 1970 season, a projected 6,632 elk will be available to hunters in Area E (Table 26).
However, this figure may change slightly if productivity
differs significantly
from 60.3 calves per 100 cows.

�- 284 -

LITERATURE CITED
Boyd, R. J. (Unpub.). The elk of the White River Plateau.
Fish and Parks Div.* Denver. Tech. Bull. No.
•

Colo. Game,

Colorado Game, Fish and Parks Division*. 1970. 1969 Colorado big game game
harvest. Unpublished report. Colo. Game, Fish and Parks Div.*. Denver. Mimeo. 160 p.
Harris, J. T. 1963. Population dynamics of the White River elk herd,
Colorado. Ph.D. Thesis. Univ. Mich. Ann Arbor, Mich. 190 p.
Hay, K. G., G. N. Hunter, and L. Robbins. 1961. Big game management in
Colorado, 1949-1958. Colorado Game, Fish and Parks Df.v Ls Lorrs ,
Denver. Tech. Bull. No.8.
112 p.
Hunter, G. N. 1967. Colorado big game harvest, 1959-1965. Colorado Game,
Fish and Parks Division*. Denver. Special Mgmt. Report. No.1.
41 p.
Keiss, R. E. 1969. Comparison of eruption-wear patterns and cementum
annuli as age criteria in elk. J. Wildl. Mgmt. 33(1):175-180.
Prenzlow, E. J. 1967. Population components - White River elk.
rado. Job Completion Report, W-38-R-2l. p. 251-276.
1968. White River elk population components.
gress Report. W-38-R-22. p. 383-421.

Colorado.

ColoJob Pro-

1969a. Area MM season summary for 1968. Unpublished report.
Colorado Game, Fish and Parks Division*. Denver. Mimeo. 2 p.
1969b. Area T season summary for 1968. Unpublished report.
rado Game, Fish and Parks Division*. Denver. Mimeo. 2 p.
1969c. White River elk population components, Coiorado.
gress Report, W-38-R-23. p. 179-234.
Quimby, D. C., and J. E. Gaab. 1957.
indicator in Rocky Mountain elk.

Colo-

Job Pro-

Mandibular dentition as an age
J. Wildl. Mgmt. 21(4):435-451.

Riney, T. 1956. Differences in proportion of fawns to herds in red deer
Cervus elaphus from several New Zealand environments. Nature.
177:488-489.
Smith, D. G. 1959b. Collection of data on the White River elk herd.
Colorado. Job Completion Report, W-38-R-12. p. 33-39.

*1964 and prior known as Game and Fish Department. 1964 Changed to Game,
Fish and Parks Department. 1968 changed to Game, Fish and Parks Division.

�- 285 -

Ti1eston, J. V. 1962. A resume of Colorado big game research projects.
1939-1957. Colorado Game, Fish and Parks Division*. Denver, Tech.'
Bull. No.9.
81 p.
Turner, F. L. 1967. Elk population trends distribution and harvest information. New Mexico. Job Completion Report. W-93-R-9. 11 p.
1968. Elk population trends, distribution and harvest information.
New Mexico. Job Progress Report. W-93-R-10. 12 p.
White, C. E. 1970. Letter to W. W. Sandfort regarding White River elk
census 1970. Colorado Game, Fish and Parks Division*. Denver.
January 20, 1970. 2 p.

*1964 and prior known as Game and Fish Department. 1964 changed to Game,
Fish and Parks Department. 1968 changed to Game, Fish and Parks Division.

�- 286 -

APPENDIX A
POST CARD SURVEY - AREA E HUNTERS

Please list results from your own hunting effort and not information from those with whom you hunted in 1969.
1.
2.

Did you hunt elk on your specified license? Yes
No _
What Game Management Unit within the AREA did you hunt?
Unit
County
Drainage
Number of Days Hunted
In Each Unit

3.
4.
5.
6.

Did you kill an elk on your specified license? Yes
No
Did you kill a: Bull
Cow
Calf
Number of points on antlers: Right
Left
_
In what Game Management Unit listed in #2 did you kill?
Unit Where Killed
Date of Kill

_

7·

How many elk did you hit that you weren't able to get?

_

8.

Was your elk wounded before you shot it? Yes

_

No

�July, 1970

- 287 -

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~--------

Project No.

W-38-R-24

Deer-Elk

Investigations

Work plan No.

14

Job No.

1

Job Title __~M~i~d~d~l~e~P~a~rk~~D~e~e~r~S~t~u~d~y~-~P~o~p~u~l~a~t~i~o_n~D~i~s~t~r_i~b~u~t~~~'o~
_
Period Covered:

April 1, 1969 - March 31, 1970

Personnel:

R. Bruce Gill, Donald W. Reichert, Olaf C. Wallmo,
Robert L. Schmidt, Julius J. Klein, Willard Travnicek,
Steven Horn, and Paul F. Gilbert.

ABSTRACT

Deer winter range boundaries are mapped for "mild, moderate, and severe"
winter conditions in Middle Park. Trapping and marking results are
summarized for the 1970 trapping program during which 205 deer were
marked with neckbands and ear tagged.
Bucks comprised 31.7 percent of
these deer, does 32.2 percent, buck fawns 14.1 percent, and doe fawns
22.0 percent.
An average of 2.6 deer were tagged per day of trapping
effort compared to 3.7 per day in 1969. Maps are included depicting
the 1969 observations of neckbanded deer by quarters of the year.

��- 289 •

MIDDLE

PARK DEER STUDY - POPULATION

DISTRIBUTION

R. Bruce Gill

p. S. OBJECTIVE
To delineate deer concentration areas in Middle Park and relate changes
in deer distribution in time and space to accumulation and physical
properties of snowo

SEGMENT OBJECTIVES
1.

20
3.
4.

Define winter range boundaries of the Middle Park deer population
mild, moderate, and severe winter conditions.
Define sub-unit boundaries.
Delineate major deer concentration areas.
Relate changes in deer distribution to accumulation and physical
properties of snow.

under

METHODS AND MATERIALS
For a complete

description

of methods

and materials

see Gill

(1969).

RESULTS AND DISCUSSION

Patterns

of Distribution

Deer distribution patterns are complex phenomena, influenced by a number
of factors among which are behavior and density of the deer themselves;
availability of food, water, and cover; snow depth, snow crust; topography;
land use patterns; etc. One purpose of this study was to map winter distribution patterns of mule deer and to investigate the relationship of changes
in these patterns to changes in snow depth and crusting.
Another was to
provide information to be used in stratifying sampling schemata for population
dynamics studies.
Still a third was to learn if deer populations using
particular, discrete winter ranges retain this discretion during other
seasons of the year.
The relationship between deer distribution and snow has been reported by
Gilbert et al. (1970).
It was concluded that deer distribution patterns
are governed by both depth and crusting of snow. Depths in excess of 18
inches of uncrusted snow essentially precluded deer use, and depths of
crusted snow exceeding 12 inches also negated deer use. Since deer winter
distribution is determined largely by snow depths, winter range boundaries
are dynamic entities from year to year as well as from period to period

�- 290 -

within a yearo Figures 1-3 represent the kinds of deer distribution
patterns that might be expected in late February and March during socalled "mild, moderate, and severe" winter conditions and hence portray
the extremes and the median limits of deer concentrations in Middle Parko
Trapping and Marking
Trapping Results
The Game Management Unit system in Colorado was initiated so that animal
species could be managed on a population basis. In Middle Park it was
suspected that the deer management units were not distinct biological
units.. It was believed there were at least four discrete populations of
deer inhabiting the Middle Park valley and these deer did not confine
their movements to game management unit boundaries. To test this hypothesis a trapping and marking program was begun in January 1969
The
objective of this program was to tag and mark deer from winter concen-.
tration areas within the boundaries of each postulated sub-unit area
(Figso 4-5). Two hundred-five deer were tagged and released during the
period January-March, 19700 Of these 31.7 percent were bucks older than
one year, 32.2 percent were does older than one year, 14 6 percent were
buck fawns, and 22 0 percent were doe fawnso Trapping and tagging results are presented for each sub-unit area in Tables 1-4.. The 1970
trapping results differed slightly in percentages by sex and age group
from the 1969 results (Table 5).. An average of 206 deer were tagged per
day in 1970 as compared with 3 7 per day in 1969
The average number of
deer trapped per day (including recaptures) in 1970 was 6 9 deer per day.
Recapture records were not kept in 1969 so no comparisons in total catches
per day are possibleo
0

0

0

0

0

0

Neckbanded Deer Observations
Neckbanded deer sightings and returns for 1969 are summarized by quarters
of the year in Figures 6-9
The data, after only one year, are too meager
to merit detailed discussion, but some of the observations are interesting.
The January-March summaries revealed the majority of neckbanded deer observations (97 percent) were in the sub-unit where they were originally trappedo
Very few deer during this quarter moved from one sub-unit to anothero
0

So few observations were recorded during the summer and fall quarters that
generalizations from these data are hazardouso
During the October-December quarter 87 percent of the red neckbanded deer
were reported from the Muddy Creek (red) sub-unit, 100 percent of the blue
neckbanded deer were reported from the Blue River (blue) sub-unit, 73 percent
of the white neckbanded deer reports came from the Troublesome Creek (white)
sub-unit, and 100 percent of the yellow neckbands were from the Williams
Fork River (yellow) sub-unit. These preliminary data support the initial
selection of sub-unit boundaries as approximations of deer population boundaries"

�- 291 -

N

Fig. 1. Winter distribution patterns of ~iddle Park deer under mild
winter conditions.

�- 292 -

N

Green

Legend;
Grey = Light deer use.
Dark = Moderate to heavy deer use.

Fig. 2. winter distribution
winter conditions.

patterns ~f Middle Park deer under moderate

�- 293 -

Figo 3
Winter distribution
winter conditionso
0

patterns

of Middle

Park deer under severe

�- 294 -

SUB-UNITS
Muddy
Blue

Creek
River

Williams
1111111\

Troublesome

Figo 40

fork

River
Creek

Hypothetical deer sub-unit boundaries in Middle Parko

�- 295 -

N

Wms.

Green

Mtn. Res.

R = red neckbanding

site

= blue neckbanding

site

B

= white neckbanding site
Y = yellow neckbanding
site

W

Figo 50

Trapsite locations for the 1970 deer neckbanding programo

�Table 1.

Deer trapped and tagged in the Muddy Creek sub-unit, 1970

Date

Sex

Age

1-17-70
1-20-70
1-31-70
2 -1-70
2 -2-70
2-6-70
2-15-70
2-22-70
2-26-70
3-4-70
3-7-70
3-9-70

Doe
Doe
Doe
Doe
Buck
Buck
Doe
Buck
Doe
Buck
Doe
Doe

Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn

SUMMARY:

12 deer tagged

Earteg Numbers
Left
Right

Neckband

MP-333
MP-342
MP-352
MP-353
MP-355
MP-368
MP-397
MP-418
MP-430
MP-441
MP-447
MP-453

Red 0 14
Red 0 2
Red 0 12
Red 0 6
Red 0 5
Red 0 7
Red 0 4
Red 0 11
Red 0 17
Red 0 1
Red 0 16
No neckband

MP-333
MP-342
MP-352
MP-353
MP-355
MP-368
MP-397
MP-418
MP-430
MP-441
MP-447

------

16.7% (2) Mature Bucks
16.7% (2) Buck Fawns

Remarks
Hwy. 40 Sec. 23
Hwy. 40 Sec. 23
Hwy. 40 Sec. 23
Hwy. 40 Sec. 23
Hwy. 40 Sec. 23
Hwy. 40 Sec. 23
Red Mountain
Red Mountain
Red Mountain
Martin Ranch E. of Hwy. 40
Red Mountain
Red Mountain

58.3% (7) Mature Does
8.3% (1) Doe Fawns

N

~

0'1

�Table 2.

Deer trapped and tagged in the Blue River sub-unit, 1970

Date

Sex

Age

1-8-70
1-8-70
1-9-70
1-9-70
1-10-70
1-10-70
1-10-70
1-10-70
1-10-70
1-10-70
1-11-70
1-11-70
1-11-70
1-11-70
1-11-70
1-12 -70
1-12-70
1-12-70
1-12-70
1-13-70
1-14-70
1-14-70
1-15-70
1-15-70
1-15-70
1-15-70
1-16-70
1-16-70
1-17 -70
1-17-70

Doe
Buck
Buck
Buck
Buck
Doe
Doe
Buck
Doe
Buck
Doe
Buck
Doe
Doe
Buck
Buck
Buck
Doe
Doe
Buck
Buck
Doe
Doe
Buck
Buck
Doe
Buck
Buck
Buck
Doe

Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature
Fawn·
Mature
Mature

Eartag Numbers
Left
Right

MP-291
MP-292
MP-293
MP-294
MP-295
MP-296
MP-297
MP-298
MP-299
MP-300
MP-301
MP-302
MP-303
MP-304
MP-305
MP-307
MP-308
MP-309
MP-310
MP-312
MP-313
MP-314
MP-317
MP-318
MP-319
MP-320
MP-324
MP-325
MP-331
MP-332

MP-291
MP-292
MP-293
MP-294
MP-295
MP-296
MP-297
MP-298
MP-299
MP-300
MP-301
MP-302
MP-303
MP-304
MP-305
MP-307
MP-308
MP-309
MP-310
MP-312
MP-313
MP-314
MP-317
MP-318
MP-319
MP-320
MP-324
MP-325
MP-331
MP-332

Neckband

Blue 0 14
Blue 0 9
Blue 0 3
Blue 0 1
Blue 0 5
Blue 0 10
Blue 0 11
Blue 0 37
Blue 0 2
Blue no if
Blue 0 6
Blue 0 4
Blue 0 7
Blue 0 15
Blue 0 12
Blue 0 18
Blue 0 30
Blue 0 24
Blue 0 27
No neckband
Blue 0 16
Blue 0 28
Blue 0 29
Blue 0 287
Blue 0 8
Blue 0 31
Blue 0 34
Blue 0 41
Blue 0 35
Blue 0 40

Remarks

Shane Gulch
Shane Gulch
Shane Gulch
Shane Gulch
Shane Gulch
Williams Peak Road
Williams Peak Road
Harsha Gulch
Harsha Gulch
Harsha Gulch
Shane Gulch
Shane Gulch
Williams Peak Road
Williams Peak Road
Williams Peak Road
Shane Gulch (MP-306 Void)
Harsha Gulch
Harsha Gulch
Harsha Gulch
Harsha Gulch (trap mortality)
Williams Peak Road
Harsha Gulch
Shane Gulch
Harsha Gulch
Harsha Gulch
Harsha Gulch
Shane Gulch
Harsha Gulch
Shane Gulch
Harsha Gulch

-----------------------------------------------------------------------------------------------------------

N
\0
"-l

�Table 2.

Deer trapped and tagged in the Blue River sub-unit, 1970 (continued).

Date

Sex

Age

1-18-70
1-19-70
1-19-70
1-21-70
1-25-70
1-26-70
2-5-70
2-7-70
2-14-70
2-17-70
2-22-70
2-23-70
2-25-70
2-25-70
2-25-70
2-26-70
2-26-70
2-28-70
3-4-70
3-4-70
3-13-70
3-19-70
3-21-70
3-19-70
3-22-70
3-23-70
3-26-70
3-26-70

Buck
Buck
Buck
Buck
Buck
Buck
Doe
Doe
Doe
Doe
Doe
Buck
Doe
Doe
Doe
Doe
Buck
Doe
Doe
Doe
Doe
Buck
Doe
Doe
Doe
Buck
Doe
Doe

Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Fawn

SUMMARY:

58 deer tagged

Eartag Numbers
Left
Right

MP-336
MP-337
MP-338
MP-343
MP-350
MP-35l
MP-362
MP-376
MP-396
MP-404
MP-4l7
MP-423
MP-427
MP-428
MP-429
MP-432
MP-433
MP-437
MP-439
MP-450
MP-468
MP-479
MP-489
MP-480
MP-492
MP-495
MP-500
MP-502

MP-336
MP-337
MP-338
MP-343
MP-350
MP-35l
MP-362
MP-376
MP-396
MP-404
MP-4l7
MP-423
MP-427
MP-428
MP-429
MP-432
MP-433
MP-437
MP-439
MP-440
MP-468
MP-479
MP-489
MP-480
MP-492
MP-495
MP-500
MP-502

32.7% (19) Mature Bucks
13.8% (8) Buck Fawns

Remarks

Neckband

Blue 0 20
Blue 0 23
Blue 0 39
Blue 0 52
Blue 0 50
Blue 0 51
Blue 0 53
Blue 0 58
Blue 0 46
Blue 0 49
Blue 0 59
Blue 0 19
No neckband
No neckband
No neckband
Blue 0 57
Blue 0 44
Blue 0 60
Blue 0 47
Blue 0 46
Blue 0 26
Blue 0 22
Blue 0 37
Blue 0 48
No neckband
Blue no 1/:
No neckband
No neckband

Shane Gulch
Shane Guclh
Willia~ Peak Road
(trap mortality)
Shane Gulch
Harsha Gulch
Shane Gulch
Williams Peak Road
Williams Peak Road
Williams Peak Road
Williams Peak Road
Williams Peak Road
Williams Peak Road
Williams Peak Road
Williams Peak Road
Spring Creek Road (trap mortality)
Spring Creek Road
Spring Creek Road
Spring Creek Road
Williams Peak Road (trap mortality)
Williams Peak Road
Williams Peak Road
Spring Creek Road
Williams Peak Road
Spring Creek Road
Spring Creek Road
Williams Peak Road
Spring Creek Road
Spring Creek Road

34.5% (20) Mature Does
19.0% (11) Doe Fawns

N

~

00

�Table 3.

Deer trapped and tagged in the Troublesome Creek, sub-unit, 1970.

Date

Sex

Age

1-22-70
1-23-70
2-4-70
2-4-70
2-5-70
2-5-70
2-6-70
2-6-70
2-6-70
2-6-70
2-6-70
2-7-70
2-7-70
2-7-70
2-7-70
2-9-70
2-9-70
2-9-70
2-10-70
2-10-70
2-11-70
2-11-70
2-11-70
2-11-70
2-12-70
2-12-70
2-13-70
2-13-70
2-13-70
2-14-70
2-14-70

Doe
Doe
Buck
Doe
Buck
Buck
Doe
Buck
Buck
Buck
Buck
Doe
Doe
Buck
Buck
Buck
Doe
Buck
Doe
Buck
Doe
Buck
Doe
Doe
Doe
Buck
Buck
Buck
Doe
Buck
Buck

Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn

Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn

Eartag Numbers
Right
Left
MP-345
MP-347
MP-356
MP-354
MP-359
MP-360
MP-363
MP-364
MP-365
MP-366
MP-367
MP-369
MP-370
MP-371
MP-372
MP-377
MP-378
MP-379
MP-38l
MP-382
MP-385
MP-383
MP-384
MP-386
MP·.,387
MP-388
MP-389
MP-390
MP-391
MP-394
MP-395

MP-345
MP-347
MP-356
MP-354
MP-359
MP-360
MP-363
MP-364
MP-365
MP-366
MP-367
MP-369
MP-370
MP-371
MP-372
MP-377
MP-378
MP-379
MP-38l
MP-382
MP-385
MP-383
MP-384
MP-386
MP-387
MP-388
MP-389
MP-390
MP-391
MP-394
MP-395

Neckband
White 0 3
White 0 4
White 0 2
White 0 5
White 0 1
No ne ckband
White 0 12
White 0 20
White 0 14
White 0 13
White 0 9
White 0 15
White 0 23
White 0 19
White 0 7
White 0 10
White 0 11
White 0 18
White 0 21
White 0 25
White 0 22
White 0 8
White 0 16
White 0 6
White 0 39
White 0 17
White 0 29
White 0 33
White 0 26
White 0 42
White 0 34

Remarks

Corral Cr. Rd. Sec. 2
Corral Cr. Rd. Sec. 2
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec 11
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 2
Rifle Range
Rifle Range
Sulphur Gulch
Corral Cr. Rdo Sec. 11
Corral Cr. Rd. Sec. 11
Rifle Range
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11
Rifle Range
Corral Cr. Rdo Sec. 11
Corral Cr. Rdo Sec. 2
Corral Cr. Rdo Sec. 11
Rifle Range
Rifle Range
Corral Cr. Rdo Sec. 2
Corral Cr. Rd. Sec. 11
Corral Cro Rd. Sec. 11
Corral Cr. Rd. Sec. 2
Rifle Range
Rifle Range
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11

--------------------------------------------------------------------------------------------------------------

o

N
\.D
\.D

�Table 3.

Date
2-15-70
2-15-70
2-16-70
2-17-70
2-17-70
2-19-70
2-19-70
2-19-70
2-19-70
2-20-70
2-21-70
2-21-70
2-21-70
2-22-70
2-22-70
2-23-70
2-23-70
2-26-70
2-28-70
2-28-70
3-2-70
3-4-70
3-4-70
3-5-70
3-6-70
3-6-70
3-7-70
3-7-70
3-8-70
3-10-70
3-11-70
3-12-70
3-12-70

Deer trapped and tagged in the Troublesome Creek sub-unit, 1970 (continued).

Sex

Age

Buck
Doe
Doe
Doe
Buck
Doe
Buck
Buck
Buck
Buck
Doe
Buck
Buck
Buck
Buck
Doe
Buck
Buck
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Doe

Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Fawn

Ha t ure

Eartag Numbers
Right
Left
MP-399
MP-400
MP-401
MP-402
MP-403
MP-407
MP-408
MP-409
MP-4l0
MP-412
MP-414
MP-4l5
MP-416
MP-42l
MP-422
MP-424
MP-425
MP-43l
MP-435
MP-436
MP-438
MP-440
MP-442
MP-443
MP-445
MP-446
MP-449
MP-451
MP-452
MP-457
MP-459
MP-46l
HP-460

MP-399
MP-400
MP-401
MP-402
MP-403
MP-407
MP-408
MP-409
MP-410
MP-412
MP-414
MP-4l5
°MP-4l6
MP-42l
MP-422
MP-424
MP-425
MP-431
MP-435
MP-436
MP-438
MP-450
MP-442
MP-443
MP-445
MP-446
MP-449
MP-451
MP-452
MP-457
MP-459
MP-461
HP-460

Neckband

Remarks

White 0 30
White 0 24
White 0 36
White 0 37
White 0 28
White 0 52
No neckband
White 0 31
White 0 40
White 0 42
White 0 49
No neckband
No neckband
White 0 45
No neckband
No neckband
No neckband
White 0 53
White 0 57
White 0 35
White 0 58
No neckband
White 0 56
No neckband
White 0 46
White 0 47
White 0 27
No neckband
White 0 51
White 0 44
White 0 60
No neckband
White 0 50

Corral Cr. Rd. Sec. 2
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Seco 11
Corral Cre Rd. Seco 11
Corral Cr. Rd. Sec. 2
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Seco 2
Rifle Range
Corral Cro Rdo Seco 11
Corral Cr. Rdo Sec. 11
Rifle Range
Corral Cr. Rdo Sec. 2
Rifle Range
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11
Corral Cro Rd. Seco 2
Corral Cro Rd. Seco 11
Corral Cr. Rd. Seco 11
Old Ridgeway Rd.
Corral Cr. Rd. Sec. 11
Old Ridgeway Rd.
Rifle Range
Rifle Range
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Seco 11
Corral Cr. Rd. Sec. 11
Corral Cr. Rd. Sec. 11
Rifle Range
Rifle Range
Corral Cr. Rd. Sec. 11
Rifle Range

--------------------------------------------------------------------------------------------------------------

w

0
0

�Table 3

Deer trapped and tagged in the Troublesome Creek, sub-unit, 1970 (continued).

0

Date

Sex

Age

3-13-70
3-13-70
3-13-70
3-13-70
3-14-70
3-16-70
3--16-70
3-16-70
3-17-70
3-17-70
3-17-70
3-17-70
3-18-70
3-19-70
3-19-70
3-19-70
3-19-70
3-19-70
3-19-70
3-20-70
3-21-70
3-21-70
3-22-70
3-22-70
3-23-70
3-23-70
3-24-70
3-26-70
3-27-70
3-27-70

Doe
Doe
Doe
Doe
Doe
Doe
Buck
Buck
Doe
Buck
Doe
Doe
Buck
Doe
Doe
Buck
Buck
Doe
Buck
Buck
Buck
Doe
Doe
Buck
Buck
Doe
Buck
Doe
Buck
Doe

Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn

SUMMARY:

94 deer tagged

Eartag Numbers
Left
Right
MP-462
MP-463
MP-464
MP-465
MP-466
MP-467
MP-470
MP-471
MP-473
MP-474
MP-475
MP-476
MP-47R
MP-481
MP-477
MP-483
MP-484
MP-485
MP-486
MP-482
MP-491
MP-488
MP-493
MP-494
MP-497
MP-498
MP-499
MP-503
MP-501
MP-504

MP-462
MP-463
MP-464
MP-465
MP-466
MP-467
MP-470
MP-471
MP-473
MP-474
MP-475
MP-476
MP-478
MP-481
MP-477
MP-483
MP-484
MP-485
MP-486
MP-482
MP-491
MP-488
MP-493
MP-494
MP-497
MP-498
MP-499
MP-503
MP-501
MP-504

31 9% (30) Mature Bucks
1308% (13) Buck Fawns
0

Remarks

Neckband

Rifle Range
Corral Cr. Rdo Sec. 11
Corral Cro Rdo Sec. 2
Old Ridgeway Rd.
Corral Cr. Rd. Seco 11
Corral Cro Rdo Sec. 11
Corral Cr. Rdo Sec. 11
Old Ridgeway Rd.
Rifle Range
Rifle Range
Corral Cr. Rd. Sec. 11
Old Ridgeway Rd.
Rifle Range
Rifle Range
Rifle Range
Corral Cro Rd. Seco 11
Corral CroRdo Seco 11 (Trap Mortality:
Corral Cr. Rd. Seco 11
Corral Cr. Rd. Sec. 2
Old Ridgeway Rd.
Old Ridgeway Rdo
w
Corral Cro Rd. Sec. 11
0
Rifle Range
~
Old Ridgeway Rdo
Rifle Range
Corral Cro Rd. Sec. 11
Old Ridgeway Rd.
Corral Cro Rdo Sec. 11
Old Ridgeway Rdo
Rifle Range

No neckband
White 0 67
No neckband
White 0 68
No neckband
White 0 59
No neckband
White 0 54
White 0 61
No neckband
White 0 70
No neckband
No neckband
No ne ckband
No neckband
No neckband
No neckband
No neckband
No neckband
No neckband
No neckband
No neckband
White 0 63
No neckband
No neckband
No neckband
No neckband
No neckband
No neckband
No neckband

31 9% (30) Mature Does
2204% (21) Doe Fawns
0

�Table 4.

Deer trapped and tagged in the Hi11iams For River sub-unit., 1970.

Date

Sex

Age

1-13-70
1-14-70
1-14-70
1-15-70
1-15-70
1-16-70
1-16-70
1-16-70
1-17-70
1-17-70
1-18-70
1-18-70
1-20-70
1-20-70
1-21-70
1-22-70
1-23-70
1-24-70
2-4-70
2-5-70
2-6-70
2-7 -70
2-7-70
2 -7-70
2 -9 -70
2-14-70
2-14-70
2-15-70
2-20-70
2-22-70

Buck
Buck
Buck
Doe
Buck
Doe
Buck
Buck
Doe
Buck
Doe
Doe
Buck
Buck
Buck
Doe
Buck
Buck
Doe
Buck
Doe
Doe
Doe
Doe
Doe
Doe
Doe
Buck
Buck
Buck

Hature
Hature
Hature
Fawn
Hature
Fawn
Hature
Hature
Hature
Fawn
Hature
Hature
Hature
Hature
Hature
Fawn
Hature
Fawn
Fawn
Hature
Fawn
Hature
Fawn
Mature
Fawn
Fawn
Mature
Fawn
Mature
Fawn

Eartag Numbers
Right
Left
HP-3ll
HP-315
HP-316
HP-321
HP-323
HP-326
HP-327
HP-328
HP-329
HP-330
HP-334
HP-335
HP-339
HP-340
HP-344
HP-346
HP-348
HP-349
HP-358
HP-36l
HP-357
HP-373
HP-374
HP-375
MP-380
HP-392
MP-393
MP-398
MP-4ll
MP-419

HP-311
HP-315
HP-316
HP-321
HP-323
HP-326
HP-327
HP-328
HP-329
HP-330
HP-334
HP-335
HP-339
HP-340
HP-344
HP-346
HP-348
HP-349
HP-358
HP-36l
HP-357
HP-373
HP-374
MP-375
MP-380
MP-392
MP-393
MP-398
HP-4ll
MP-419

Neckband

Yellow 0 5
Yellow 0 9
Yellow 0 11
Yellow 0 16
Yellow 0 13
Yellow 0 4
Yellow 0 1
Yellow 0 14
Yellow 0 2
Yellow 0 3
Yellow 0 6
Yellow 0 147
Yellow 0 17
Yellow 0 19
Yellow 0 7
Yellow 0 8
Yellow 0 18
Yellow 0 26
Yellow 0 30
Yellow 0 21
Yellow 0 24
Yellow 0 34
Yellow 0 12
Yellow 0 31
Yellow 0 28
Yellow 0 46
Yellow 0 44
Yellow 0 23
Yellow 0 29
Ye ILow 0 22

Remarks

East end Cedar Ridge
Williams Fork Boat Ramp Road
Hilliams Fork Boat Ramp Road
Williams Fork DarnAccess Road
Williams Fork Boat Ramp Road
Williams Fork DarnAccess Road
Williams Fork Boat Ramp Road
Williams Fork Boat Ramp Road
Williams Fork Dam Access Road
Williams Fork Boat Ramp Road
Williams Fork Boat Ramp Road
Williams Fork Boat Ramp Road
Williams Fork Boat Ramp Road
Williams Fork Boat Ramp Road
Williams Fork Boat Ramp Road
Williams Fork Boat Ramp Road
Williams Fork Boat Ramp Road
Williams Fork Dam Access Road
West end Cedar Ridge
West end Cedar Ridge
West end Cedar Ridge
West end Cedar Ridge
West end Cedar Ridge
West end Cedar Ridge
East end Cedar Ridge
East end Cedar Ridge
East end Cedar Ridge
West end Cedar Ridge
West end Cedar Ridge
West end Cedar Ridge

-----------------------------------------------------------------------------------------------------------

w
0
N

�Table 4.

Deer trapped and tagged in the Williams Fork River sub-unit, 1970 (continued).

Date

Sex

Age

2-22-70
2-24-70
2-27-70
3-6-70
3-7-70
3-9-70
3-10-70
3-14-70
3-17-70
3-20-70
3-21-70
3-23-70

Buck
Doe

Mature
Mature

SUMMARY:

SUMMARY
OVER ALL
SUB-UNITS:

?

?

Doe
Doe
Doe
Doe
Buck
Buck
Doe
Doe
Buck

Mature
Fawn
Mature
Fawn
Fawn
Fawn
Fawn
Fawn
Mature

41 deer tagged

205 deer tagged

Eartag Numbers
Left
Right

MP-420
MP-426
MP-434
MP-444
MP-448
MP-454
MP-458
MP-469
MP-472
MP-487
MP-490
MP-496

MP-420
MP-426
MP-434
MP-444
MP-448
MP-454
MP-458
MP-469
MP-472
MP-487
MP-490
MP-496

Neckband
Yellow 0 33
Yellow 0 39
Yellow 0 54
Yellow 0 10
Yellow 0 47
Yellow 0 53
No neckband
No neckband
No neckband
No neckband
No neckband
No neckband

Remarks

West end Cedar Ridge
West end Cedar Ridge
West end Cedar Ridge
East end Cedar Ridge
West end Cedar Ridge
East end Cedar Ridge
West end Cedar Ridge
East end Cedar Ridge
East end Cedar Ridge
East end Cedar Ridge
East end Cedar Ridge
West end Cedar Ridge

34.1% (14) Mature Bucks
14.6% (6) Buck Fawns

22.0% (9) Mature Does
29.3% (12) Doe Fawns

31.7% (65) Mature Bucks
14.1% (29) Buck Fawns

32.2% (66) Mature Does
22.0% (45) Doe Fawns

neck injury
w
0
w

�Table 5.

Comparisons of the 1969 and 1970 Middle Park deer trapping and tagging results.

Sub-Units
Troublesome Creek
Percent
MF
FF
AF
FF
AM

AM

Totals
Percent
MF
AF

FF

22.2

23.6

39.4

16.9

20.1

14.6

29.3

31. 7

32.2

14.1

22.0

17.7

26.5

27.0

36.5

15.7

20.8

Williams Fork River
Percent
MF
FF
AM
AF

Year

Muddy Creek
Percent
AF
MF
AM*

FF

AM

Blue River
Percent
MF
AF

1969

13 .9

51.4

13.9

20.8

35.1

35.1

16.3

l3 .5 23.1

35.6

19.2

22.1

18.5

37.1

22.2

1970

16.7

58.3

16.7

8.3

32.7

34.5

l3.8

19.0

31. 9

31.9

l3 .8

22.4

34.1

22.0

Totals 14.2

52.2

14.2

19.4

33.7

34.8

14.7

16.8

27.3

33.8

16.7

22.2

27.9

27.9

*AM = Adult males (older than one year)
AF = Adult females (older than one year)
MF = Male fawns
FF = Female fawns

w

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'-

..

nec kbon ds

Blv.e..J..--/
neck'hands.
.'i:;. . 'i

-

-.

.~.

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Fig. 6.

Observations of neckbanded deer, January-March, 1969.

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�- 309 -

Maximum distances individual deer traveled from trapsites were: 40 airline
miles for red, 18 airline miles for blue, 26 airline miles for white, and
22 airline miles for yellow neckbanded deer. Five sightings of Middle
Park neckbanded deer were reported from areas outside the Park, three red
neckbands from the Oak Creek-Phippsburg areas and two blue neckbands from
the Piney Ridge area (Figso 8 and 9)e

LITERATURE CITED
Gilbert, Po F
0 Co Wallmo, and R, B. Gill. 1970. Effect of snow depth
on mule deer in Middle Park, Colorado. J. Wild1. Mgmt. 34(1):15-23.
0,

0

Gill, Ro Bo 1969. Middle Park Deer Study. Population distributiono
Colo. Game, Fish and Parks Div., Game Research Sectiono Fed. Aid
Projo W-38-R-23 Job Progress Rept. July Part 10 ppo 79-104

Prepared by
R. Bruce Gill
Wildlife Researcher

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                  <text>- 311 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-24

Deer-Elk

Investigations

Work Plan No.

14

Job No.

2

Job Title

Middle Park Deer Study
population Density and Structure

Period Covered:

October

14, 1969 to February

1, 1970

Personnel:

R. B. Gill. P. F. Gilbert. o. C. Wallmo, J. Cooney,
V. R. Clark, H. Donaho 7 D. Benson, L. H. Carpenter,
H. R. Shepherd, C. E. Braun, J. F. Corey, W. T. McKean,
H. D. Funk, W. H. Rutherford, W. T. Burkhard, L. M.
Finnell, D. Horak, B. D. Baker, C. woodward, D. T.
Weber, R. Eo Keiss, W. J. Adrian, and L. J. Griess.

ABSTRACT

Estimates of the total Middle Park winter deer populations were 10,640
in 1968, 9,112 in 1969, and 7,205 in 1970. Variances of each strata
were found to be related to each strata mean in an apparent curvilinear
fashion.
Frequency distributions of the data show similarities to a
negative binomial distribution.
Pre-season sex and age classifications
yielded ratios of 43 bucks:lOO does and 93 fawns:lOO does.
Post-season
snowmobile classifications produced ratios of 48 bU9ks:100 does and 82
fawns:lOO does, while post-season helicopter counts resulted in ratios
of 45 bucks:100 does and 77 fawns:lOO does. The estimates of sex and
age structure for. 1969-70 show a downward trend in both buck:doe and
fawn:doe ratios from the two previous years.
Similar trends are
apparent from check station data.

��- 313 -

MIDDLE PARK DEER STUDY
POPULATION DENSITY AND STRUCTURE
R. Bruce Gill

P. S. OBJECTIVE
To estimate the density and sex and age structure of the Middle Park
deer population in order to harvest this population more efficiently.

SEGMENT

OBJECTIVES

1.

Estimate
Park.

the size of the total winter

2.

Estimate
lation.

the sex and age structure

METHODS
Methods and materials
will not be presented

of the Middle

of deer

in Middle

Park deer popu-

AND MATERIALS

have been discussed
in this discussion.

RESULTS

population

in detail

by Gill

(1969) and

AND DISCUSSION
Density

Population

Estimates

The 1970 helicopter quadrat census was conducted on January 5-8, 1970.
A total of 13.9 hours (including ferry time) was used to count 80 square
mile sections or 5.8 sections counted per hour of flight time.
The cost
of censusing was $16.41 per count section.
Results of three years of
counts are summarized in Table 1. Mean numbers of deer have varied from
a high of 20.73 deer per square mile in 1968 to 12.29 deer per square mile
in 1970.
Projected total populations by sub-unit indicate that the Muddy
Creek and the Williams Fork River sUb-units have contributed most to this
decline (Table 2). Possible rei'lSOl1S
for this population decline are lower
fawn survival and higher hunting mortality encountered in 1969.
Estimates
of fawn populations declined 15.5 percent from 1968-1969 and 23.4 percent
from 1969-1970.
In contrast doe estimates declined 11.2 percent from
1968-1969 and 14.9 percent from 1969-1970 (Fig. 1). This decline was from
88 fawns per 100 does in 1968 to 85 fawns per 100 does in 1969 to 77 fawns
per 100 does in 1970 (Fig. 2).

�'l'ab1e1.

Numbers

of deer counted

per quadrat

within

the eight sampling

strata,

Williams
Muddy Creek
Muddy Creek
Blue River
Blue River
High DensitJ::
Low DensitJ::
High DensitJ::
Low DensitJ::
1968 1969 1970
1968 1969 1970
1968 1969 1970
1968 1969 1970

2
0
0
0
0
8
64
32
124
58
30
32

0
0
0
0
0

I,

0
J

0

---

&lt;)

9

0

6
18
99
39
66
46
87
31
6
12
0
0
22
37

34
9
20
6
2
2
42
28
5
12
0
0
5
1

---

,.
=-~442 478

44
20
27
0
1

'1

0
0
0

56

6
0
0
0
0
0
0
0
0
0

-----

IS

0
0
0
0
0
11

0

0

14
25
0
76
31
4
9
134
49
71
124
28

175

56

21

42

565

8.8

9.3

1.8

3.5

---

0

--0

=

26.0 23.9

_yst
v(yst)

=
=

0
1
0
30
0

19
0
1
36
88
20
4
102
15
105
58
17

14
0
0
24
77
18
3
20
83
136
49
6

4
10
0
30
32
0
0
31
0

465

430

107

47.1 38.8 35.8

1968
20.73
11.35

8
10
0
0
35
0
0
36
29

0
17
0
37
34
0
0
26
0

40
0
45
42
4

Park,

Colorado,

1968-1970.

Williams

Fk. R.
Hi9:h DensitJ::
1968 1969 1970

0
2
26
73
4

Middle

0

0
25
21
0

Fk. R.
Low DensitJ::
1968 1969 1970

0
6
14
0
0

0
0
3
0
0

0
0
1
0
0

Troublesome Cr.
Granby
High Densitx
Low DensitJ::
1968 1969 1970
1968 1969 1970

43
25
0
35
50
65
0
9
0
26

5
41
0
39
76
44
15
36
0
6

11
60
0
51
42
90
7
8
0
29

-------------

7
2
2
0
0
0

---

---

6
23
0
0
0
0
0

w
I-'

.p-

118

114

11.9 13.1 12.7

1969
15.52
4.25

105

131

46

20

3

1

21.0 26.2

9.2

4.0

0.6

0.2

1970
12.29
2.55

298

---

II

29

25.3 26.2 29.8

---

1.8

4.1

253

262

�Table 2.

Estimates

Sub-Unit
Sampling Area

of Middle Park deer population

by sub-unit sampling areas, 1968-1970.

1968

1969

1970

Muddy Creek

4,101

2,994

1,429

Blue River

4,074

3,531

3,290
I

Williams

Fork River

843

774

271

w
•.....
VI

Troublesome

Creek

1,679

1,910

134

306

10,640 ± 2,879

9,112 ± 1,975

7,206 ± 1,881

7,746 - 13,534

7,137 - 11,087

5,235 - 9,087

1,622

Granby

TOTALS
90% Confidence

Limits

�II, 000 ~
TOTAL

10,000 ~

POPULATION

9 ,000 ~
8 ,000 ~
~

w
w

7 ,000 ~

••
0

6,000 •

0

V)

~

w

w
•....
0\

5,000 •

c:Q

~

::::&gt;

Z

4,000 •

e..

3,000.
2,000.

DOES

.

••••••••••••••••••••••••••••••••••••
'iIr••••••••••••••••••••••••••••••••••••
e. · · · · · ·
FAWNS

,, , , ,, , , , , , '

••...............................

.-t

A

A

A

1968

1969

1970

BUCKS

41

1,OOO~

o.
Fig. 1.

Estimates

of Middle Park deer population

from helicopter

census data.

�1l0~
100~
FAWNS· '100

90~

DOES

•••....•..................•...•....••••.............••.......•..•........

~

80~
IJ)

w

70~

0
0

60~
0
0

--

BUCKS:l00 DOES

t-'
-...J

50~

.....•

ex
w
Q..

W

40~

IJ)

0

-

30~

I-

«
ex

20~
10.

O~

Fig. 2.

Post-season

A
1968

sex and age ratios estimated

•

•

1970

1969
from helicopter

classification

counts,

1968-70.

�- 318 -

Confidence

Limits

Confidence limits about the total population estimate are quite large
and increase with increasing population size (Fig. 3).
In 1968 when
the population was estimated at 10,640 deer the 90 percent confidence
limits were ± 2,879 while in 1970 with an estimated population of 7,205
the 90 percent confidence limits were ± 1,881 (Table 2). This characteristic results because the mean and variance of each stratum are not
independent but appear to be related in a curvilinear fashion (Fig. 4).
Frequency distributions
of the data readily reveal that they are not
distributed
normally about the mean (Figs. 5 and 6), but are highly
skewed toward the zero end.
Similar frequency distributions
have been
reported for pellet group data by Bowden et al. (1969) and McConnell
and Smith (1970).
The ninety percent confidence limits overlap for
each of the three years in which deer were censused (Fig. 3). Because
of this, means were compared for significant differences by using paired
"t" tests.
Both the 1968 and 1969 populations were significantly different from the 1970 population with a probability of less than 0.10 that
the differences
could have occurred by chance, while the 1969 population
was not significantly
different from the 1968 population.

Sex and Age Composition

Pre-season

Classifications

Pre-season deer classifications
were obtained from helicopter surveys
conducted on October 14 and 15, 1969.
Sampling was conducted so that
nearly all areas of suspected deer concentrations
within each sub-unit
were included.
The major objective for the pre-season classification
was to obtain estimates of the fawn:doe ratios, although bucks were
recorded as well.
The buck:doe ratio is believed to be lower than the
true population ratio because exclusively buck groups were seldom observed
in aggregates larger than 4 individuals
(Table 3) and, consequently,
there
was a greater probability of missing the smaller, less obvious, buck
groups than the larger doe-fawn groups.
Sample size resulting from this
survey was 1,065 deer classified with the resulting ratios:
43 bucks:lOO
does and 93 fawns: 100 does.

Post-season

Classifications

Snowmobile Classifications--Three
separate classifications
were conducted
along established snowmobile routes (Gill, 1969) during this segment, one
in November and two in December.
Ratios for the combined data from all
three sampling periods were 48 bucks:lOO does and 82 fawns:lOO does (Table
4).
This compares with ratios of 37 bucks:lOO does and 93 fawns:lOO does
in 1967 and 55 bucks:lOO'does
and 92 fawns:lOO does in 1968.

�- 319 -

14,000

13,000

12,000

11,000

0:::

w
w

10,000·

0
u,

0
en
0:::

9,000

W

co
~

::&gt;

Z

8,000

7,000

6,000

5POO~------~--------------~--------------~
1968
Fig. 3. Estimates
percent confidence

of total Middle
limits.

1970

1969
Park deer population,

bracketed

by 90

�~

VARIANCE

oJ
0

0
0
I

0
0

e".I
0
0

I

I

N

~
0
0
I

(]I

0)

0
0

0
0

I

I

~

8

(X)

0
0

W·

0
0

I

-

0
0
0

0
0

-N
0
0

-

e".I
0
0

-~
0
0

0)

(11

0
0

0
0

~
0
0

(X)

-

-W

0
0

0
0

N

0
0
0

•

•

o

I\l

0

,

.-

•

•
,-

-

(,II

~

•

O.

w
I'-&gt;

.:

0

•

~I

•

0

•

s
Fig. 4.

Relationship

between

strata means and variances

for Middle Park deer census data.

�- 321 -

80
IJ)
IJ)

-c

70

-.I

u
I
U

1968
60

-c

w

50

z
'-"

&gt;.

u

z
w

::::&gt;

C!J
w
a::
u,

30
20

10

•
O+-~--~--~--~~~~
01.

19

201

6q

401.

39

59

10~ 120/
99
119 139

801.

79

CLASSES

IJ)
IJ)

-c

80

80

70

70

-.I

1969

U

1970

60

60

50

50

..- 40
~

40

&gt;
Z

30

30

C!J 20
w

20

10

10

o

o~- •.....
--.,.-~::::~:::=;~~

I

u

-c

w

Z
'-"

u
w

::::&gt;

a::
u,

07
19

20

120, 0
20
40 60
80, 100, 120
739 40Z59 60779 80,'99 100'119
7139 719 739
'59 '79 '99
'119 "39
CLASSES

Fig. 5.
Frequency distributions
data, 1968-70.
.

CLASSES
for Middle

Park

helicopter

quadrat

census

�- 322 -

80
70
60

.-

50

'-'"

40

~
0

&gt;-

u
Z

30

w

:::&gt;

0

20

w
~

u,

10
0

a7 2q
19
39

4q 601
59

801

79

99

1001 12O,
119

139

CLASSES
COMBINED

DATA 1968-70

Fig. 6. Frequency distribution
for the combined
quadrat census data for the years 1968-70.

Middle

Park helicopter

�- 323 -

Table 3.

Results of pre-season deer classifications, October 14-15,1969.

Area

Bucks

Does

Fawns

Total

4
2
6

1

6
2
3
2
7

5

10
4
9
3
12
3

1
1
1

2

3
1
1
6

4
2

2
1

7

1

2

1

2

2

3

2

1
1

2
1

1

3
3
2

4
4
2

2

1
4

2

8

1

1

2

9
4
5

5
2
5

15
6
12

3
5
11

2
5
14

5
l3
25

Muddy Creek Sub-Unit
Gore Canyon to Sheep Creek

3
Sheep Cr. to Gore Pass Hwy.

6
1
1
3
2
1
3
Gore Pass Hwy. to Red Dirt Cr.

1
4

Red Dirt Cr. to Tyler Mtn.

Wolford Mtn.

Little Wolford Mtn.
S. Notch Gunsight Pass

1
2

N. Notch Gunsight Pass
3

4
3
3
2
4
3
6
4
3
4
7
7

5
1

--------------------------------------------------------------------------------

�- 324 -

Table 3. Results
(continued).

of pre-season

deer classifications,

Area

Troublesome

Creek

October

14-15,

Bucks

Does

Fawns

Total

4
3
2

3
2
4

8
8
9
7

3
1
1
2
1
2
1
1
3
2
2

2
2
1
4
1
2

Sub-Unit

E. Side Troublesome

Creek

to East Fork

1
3
3
7

E. Fork Troublesome
Creek Basin

Creek

to Slide

3

2

1

2
2
1
2

1
1
1
1
1
1

1

1
4

2

4
4
3
2

2
3
5
4

4
2
2
4
1
2
1

3
2
1
2
2
2
3

3
2
1

3
4
2

3
3
2
3

1

2
1

1

2

Dry Gulch

to Rock Creek

1
2
1
1
2

Corral

Creek

1969

1
1

1
1

3
5
3
2
8
2
4
1
3
5
4
4
1
1
1
2
1
7
6
7
9
6
2
7
4
3
6
3
4
4
7
8
3
1
4
4
5
4
1
3
2

-------------------------------------------------------------------------------

�- 325 -

Table 3. Results
(continued) •

of pre-season

Area

deer

classifications,

14-15,

October

1969

Bucks

Does

Fawns

Total

Mt. Bross

4

6
2
1

4
4
2

10
6
3

Ute Bill Creek

1
1

2

1

1
4

4
1
2
2
1
3
4
1
2

4
1
1
1
1
3
3
2

Sheriff

Creek
2
1
4

1

4

Kinney

Creek

McQueary
Drowsy

Smith

1

Creek

Water

Creek

4
3
4

Creek

8
4
3
4
6
6
7
3
3
4

1
3
1
1

1
3
1

2
6
2
1

1

2

4

.2

1

3

4

3

1
1

1
2

4
10
4
2
3

2
4
4

5
2

2
9
6

4

4

9

E. of Hwy. 125 and N. of Willow
Creek

Res.

1

E. of Hwy. 125 and S. of Willow
Creek

Res.

1

1

---------------------------------------------------------------------------------

�- 326 -

Table 3. Results
(continued).

of pre-season

deer classifications,

Area

Blue

River

Bucks

Does

October

14-15,

Fawns

Total

3

2

5
4

1
3
1

1
5
1

2
8
2

Sub-Unit

San Toy Mtn.
4

Trough

Eagle

Road S. to Eagle

Pass

Ranch

Pass Ranch

to Spring

Cr.

3
5
3
1

3
5
1
1
2

2

6

3

1
1
5
2
4

2
4

Spring

Creek

to Green Mtn.

3
2

4
1

2
Acorn

Creek

Ute Creek

to Ute Creek

to Shane

6

Gulch

13

12

31

11
4
4

11
6
2

22
10

15

9
7
3
5
4
14
1

28

1

Gulch

to Haystack

Mtn.

7
7
15
1

4

1
1
2
1

2

9

2
6
6

2
4

7
3

2

1

3
2
4

Shane

1969

3
1
1
1
2
1
1
2
4
1
2

2
1
1

13
9

12
13
33
2
4
6
2
3
1
4

2
5

4
1
2

4
1
7
8
2
4
2

----------------------------------------------------------------------------------

�- 327 -

Table 3. Results
(continued).

of pre-season

deer

Area

Shane

Gulch

classifications,

Bucks

to Haystack

Mtn.(cont.)

October

14-15,

Does

Fawns

Total

1
3
1
2
2
2

3
1
1
1
2
2

4
4
2
3
4
4
1

5
3

5
1

1
3

2
4

10
5
1
2
3
8

2
1

3
1

5
2

1
1
1
3
6

3

5
1
2
8
12
2

1
!5

1
5

1
1
3
3
4
2

1
1
6
5

1

E. Side Lawson

Ridge

to Kremmling
1
1
2
1

Williams

Fork River Sub-Unit

cottonwood

Beaver

Pass

1

Creek

2
1
2
Little

1969

Muddy

1

Creek

3
W. Side Williams Fork River
Battle Mtn. to Kinney Creek

1
3
5

1
2
14

from
2

2

7
2

1
4
2
1

2

2
4
9
8
13
4
1
4
2
3

--------------------------------------------------------------------------------

�- 328 -

Table 3. Results
(continued) .

of pre-season

deer classifications,

Area

October

14-15,

1969

Bucks

Does

Fawns

Total

1
3

3

6

1

1
2

2
4

10
3
3
7

TOTALS

195

451

419

1,065

PERCENT

18.3

42.4

39.3

100.0

W. Side Williams Fork River
Battle Mtn. to Kinney Creek

RATIOS:

from
(cont. )

100 Does

43 Bucks:100

Does

93 Fawns:l00

Table 4. Comparisons of post-season
snowmobile and helicopter
of mule deer population sex and age composition.

Sample

Date

Sample

Size

Bucks:lOO
Snowmobile

classifications

Ratios
Does

Fawns:lOO

Does

155

Classifications
76:100

94:100

2-3, 1969

226

48:100

88:100

15-18,

287

35:100

80:100

668

48:100

82:100

Nov.

18-22,

Dec.
Dec.

1969

1969

Totals

Helicopter
Jan.

Does

5-8, 1970

1,045

Classifications
45:100

77: 100

�-

329 -

Helicopter Classifications--During
the 1970 deer winter quadrat census .
1,045 deer were classified.
Ratios were 45 bucks:lOO does and 77 fawns:
100 does.
Comparing these data to the two previous years, the trend in
both buck:doe ratios and doe:fawn ratios has been downward (Fig. 2).

Check

Station

Data

Two check stations were operated for the first nine days of the 1969
big game hunting season to obtain estimates of the sex and age structure
of the harvest.
One was at Idaho Springs and one at Dillon.
The combined
sample size from these two stations was 1,717 deer, of which 1,479 could
be classified into sex and age categories.
Summaries of these data are
presented in Tables 5-9. Placement of deer into specific year classes
was done by the cementum aging technique (Erickson and Seliger, 1969).
Comparisons with the 1967 and 1968 data revealed that in 1969, for the
first time, the three year old age class did not contain a higher percentage of the t.ot.a
L sample than the two year old class.
All of the
1969 teeth were sectioned, stained, and mounted on permanent microscopic
slides.
This permitted time for lengthy examinations of questionable
teeth before assigning them to an age group.
In previous years teeth
were aged as rapidly as possible by laboratory personnel at the Game
Research Center.
I believe there is considerable opportunity
for error
in distinguishing
between two year old and three year old animals.
The
magnitude of this error is unknown as are possible means of eliminating
errors.
Tests should be conducted by collecting a large sample of teeth
from known-age deer and having personnel responsible
for aging teeth
assign each tooth to an age class with the same procedure now employed.
This age structure array could then be compared to the known age information and assessments made of the error associated with each year class
estimate.
As mentioned earlier, field classifications
of live deer indicated a
downward trend in ratios of bucks:lOO does and fawns:lOO does since 1968.
Similar trends are evident from check station ratios.
Ratios of bucks:
100 does, fawns:lOO does, and yearling does:lOO adult does all show declines over the three year sampling period (Fig. 7).
Reasons for these
declines are unknown, but all the evidence indicates declining popUlations
and not just chance phenomena associated with sampling error.
One interesting sidelight associated with the 1969 check stction collections was. the occurrence of a white-tailed
deer in the sample.
This
animal was a three year old buck taken from Tyler Mountain on 10-18-69.
This is the first white-tailed
deer checked from Middle Park during the
three year duration of this study.

�Table

5.

Sex and age classes

of deer checked

Unit
No.

Adult
Males

Yearling
Males

Male
Fawns

15

21
(22.6%)

12
(12.9%)

6
(6.5% )

9
(9.7%)

18

110
(25.4%)

62
(14.3%)

46
(10.6%)

17
(3.9%)

Headless
Males

at Idaho Springs
Adult
Females

and Dillon

checking

stations,

October,

1969.

Yearling
Females

Female
Fawns

Headless
Females

Headless
Unclass.

Totals

23
(24.7%)

4
(4.3%)

11
(11.8%)

7
(7.5%)

0
(0.0%)

93
{l00.0%)

110
(25.4%)

28
(6.5%)

28
(6.5%)

31
(7.2% )

1
(0.2% )

433
(100.0%)
I

27

54
(19.4%)

46
(16.5%)

18
(6.5%)

19
(6.8%)

70
(25.2%)

25
(9.0%)

17
(6.1%)

26
(9.4%)

3
(1.1%)

278
(100.0%)

28

63
(15.6%)

50
(12.4%)

48
(11.9%)

20
(5.0%)

119
(29.5%)

32
(8.0%)

36
(8.9%)

35
(8.7%)

0
(O.O%)

403
(100.0%)

37

102
(20.0%)

85
(16.7%)

42
(8.2%)

21
(4.1%)

119
(23.3%)

53
(10.04%)

39
(7.7%)

35
(6.9%)

14
(2.7%)

510
(100.0%)

Totals

350
(20.4%)

255
(14.9%)

160
(9.3%)

86
(5.0%)

441
(25.7%)

142
(8.3% )

131
(7.6%)

134
(7.8% )

18
(1.0%)

1,717'
(100.0%)

w
w

0

�Table 6. Sex and age classes
headless deer.

Unit
No.

Adult
Males

of deer checked

at Idaho Springs

Yearling
Males

Male
Fawns

and Dillon

Adult
Females

checking

Yearling
Females

stations,

excluding

Female
Fawns

Totals

15

21
(27.2%)

12
(15.6%)

6
(7.8% )

23
(29.9%)

4
(5.2%)

11
(14.3%)

77
(100.0%)

18

110
(28.6%)

62
(16.2%)

46
(12.0%)

110
(28.6%)

28
(7.3% )

28
(7.3%)

384
(100.0%)

27

54
(23.5%)

46
(20.0%)

18
(7.8% )

70
(30.4%)

25
(10.9%)

17
(7.4%)

230
(100.0%)
I

28

63
(18.1%)

50
(14.4%)

48
(13.8%)

119
(34.2%)

32
(9.2%)

36
(10.3%)

348
(100.0%)

37

102
(23.2%)

85
(19.3%)

42
(9.5%)

119
(27.0%)

53
(12.1%)

39
(8.9%)

440
(100.0%)

Totals

350
(23.7%)

255
(17.2%)

160
(10.8%)

441
(29.8%)

142
(9.6%)

131
(8.9%)

1,479
(100.0%)

w
w

t-"

�Table 7.

Number and percent

of deer checked in each age category by sex classes,

Unit
No.

Adult
Males

Yearling
Males

15

21
(53.8%)

12
(30.8%)

6
(15.4%)

39
(100.0%)

18

110
(50.5%)

62
(28.4%)

46
(21.1%)

218
(100.0%)

Male
Fawns

Totals

Adult
Females

excluding

headless

animals.

Yearling
Females

Female
Fawns

Totals

23
(60.5%)

4
(10.5%)

11
(29.0%)

38
UOO.O%)

110
(66.2%)

28
(16.9%)

28
(16.9%)

166
(100.0%)
I

27

54
(45.8%)

46
(39.0%)

18
(15.2%)

118
(100.0%)

70
(62.5%)

25
(22.3%)

17
(15.2%)

112
(100.0%)

28

63
(39.1%)

50
(31.1%)

48
(29.8%)

161
(100.0%)

119
(63.6%)

32
(17.1%)

36
(19.3%)

187
(100.0%)

37

102
(44.6%)

85
(37.1%)

42
(18.3%)

229
(100.0%)

119
(56.4%)

53
(25.1%)

39
(18.5%)

211
(100.0%)

Totals

350
(45.8%)

255
(33.3%)

160
(20.9%)

765
(100.0%)

441
(61. 8%)

142
(19.9%)

131
(18.3%)

714
(100.0%)

w
w

N

�Table

unit
No.

8.

Number

and percent

Adult
Males

yearlings

in the Middle

Yearling
Males

Park deer check, October,

Totals

Adult
Females

1969.

Yearling
Females

Totals

15

21
(63.6%)

12
(36.4%)

33
(100.0%)

23
(85.2%)

4
(14.8% )

27
(100.0%)

18

110
(64.0%)

62
(36.0%)

172
(100.0%)

110
(79.7%)

28
(20.3%)

138
(100.0%)

27

54
(54.0%)

46
(46.0%)

100
(100.0%)

70
(73.7%)

25
(26.3%)

95
(100.0%)

28

63
(55.8%)

50
(44.2%)

113
(100.0%)

119
(78.8%)

32
(21.2%)

151
(100.0%)

37

102
(54.5%)

85
(45.5%)

187
(100.0%)

119
(69.2%)

53
(30.8%)

172
(100.0%)

Totals

350
(57.9%)

255
(42.1%)

605
(100.0%)

441
(75.6%)

142
(24.4%)

583
(100.0%)

w
w
w
I

�Table 9.
1967-69.

Age structure

data from counts

of dental

cementum

layers

in incisor

teeth collected

from Middle

Numbers and Percent in Each Age Class
7 Yrs. 8 Yrs. 9 Yrs. 10 Yrs.
5 Yrs. 6 Yrs.
4 Yrs.

11+ Yrs.

Unc l..

Total

1
(0.2%)

0
(0.0%)

463
(100.0%)

1
(0.2%)

2
(0.5%)

424
0
(0.0% ) (l00.0%)

0
(0.0%)

2
(0.3%)

17
(2.2%)

765
(100.0%)

4
5
8
(0.5% ) (0.3% ) (0.3%)

17
(1.0%)

1,625
(100.0%)

Year

Fawns

1 Yr.

2 Yrs.

3 Yrs.

1967

99
(21.4%)

187
(40.4%)

45
(9.7%)

45
(9.7%)

40
(8.6%)

Males
12
18
(3.9%) (2.6%)

7
(1.5%)

5
(1.1%)

1
3
(0.2% ) (0.7%)

1968

82
(19.3%)

181
(42.7%)

51
(12.0%)

52
(12.3%)

23
(5.4%)

14
(3.3%)

8
(1.9%)

3
(0.7%)

4
(1.0%)

3
(0.7%)

1969

160
(20.9%)

255
(33.3%)

159
(20.8%)

92
(12.0%)

38
(5.0%)

13
(1.7%)

11
(1.4%)

9
(1.2%)

5
(0.7%)

4
(0.5%)

Total

341
(20.7%)

623
(37.7%)

225
(15.4%)

189
(11.4%)

101
(6.1%)

45
(2.7%)

31
(1.9%)

19
(1.2%)

14
(0.8%)

Oldest

Individual

Park deer,

Females
1967

80
(22.9%)

84
(24.0%)

39
(11.1%)

45
(12.9%)

39
(11.1%)

22
(6.3%)

16
(4.6%)

7
2.0%)

4
3
(1.1%) (0.9%)

4
(1.1%)

7
(2.0%)

0
(0.0%)

350
(100.0%)

1968

59
(20.1%)

58
(19.7%)

40
(13.6%)

60
(20.4%)

20
(6.8%)

8
(2.7%)

8
(2.7%)

8
(2.7%)

5
(1.7%)

7
(2.4%)

3
(1.0%)

4
(1.4%)

14
(4.8%)

294
(100.0%)

1969

131
(18.3%)

142
(19.9%)

192
(26.9%)

93
(13.0%)

51
(7.1%)

20
(2.8%)

13
(1.8%)

14
(2.0%)

19
(2.7%)

8
(1.1%)

3
(0.4%)

12
(1.8%)

16
(2.2%)

714
(100.0%)

Total

270
(19.9%)

284
(20.9%)

271
(19.9%)

198
(14.6%)

110
(8.1%)

50
(3.7%)

37
(2.7%)

29
(2.1%)

28
18
(2.1%) 1.3%)

10
(0.8%)

23
(1.7%)

30
(2.2%)

(100.0%)

Oldest

Individual

= 17 Yrs.

w
w

~

= 13 Yrs.

1.359

�- 335 -

BUCKS: 100 DOES

150~
140

~

130

~

120~

1 10 ~

." '" '" "

100 ~

fn

I"

"""

90~

I&amp;J

0
0

0
0

80~

I"

FAWNSzlOO
I"

I"

I"

I"

·DOES

I""

.""

70~

ADULT

IIII
IIIIII

II
IIII

"""""""'~

60~

a:

I&amp;J

0..

fn

50~
YEARLING

0

40~

t&lt;t:
0:

30~

DOES:

100

ADULT

DOES

20~

IO~
O~

A
1967

1968

1969

Fig. 7.

Check station ratios of bucks:lOO does, fawns:lOO does, and yearling

does:100

adult does,

1967-1969.

�- 336 -

LITERATURE

CITED

Erickson, J. A., and W. G. Seliger.
1969.
Efficient sectioning
incisors for estimating ages of mule deer.
J. Wildl. Mgmt.
33(2) :384-388.
Gill,

of

R. B.
1969.
Middle Park deer study.
Population density and
structure.
Colo. Div. Game, Fish and Parks. Game Res. Div.
Fed. Aid Proj. W-38-R-23.
Game Res. Rept. July, Part 1:105-122.

n';&gt;
Prepared

by

'"
J,
,)~J
-\..W

,~ R. Bruce Gill
Wildlife Researcher

I,

�- 337 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-24

Deer-Elk

Work Plan No.

14

Job No.

Investigations
3
and Mortality

Job Title

Middle

Park Deer Study Productivity

Period Covered:

January

1 - July 31, 1970

Personnel:

C. Grand Pre, R. Davies, R. Glaze, L. H. Carpenter,
W. Travnicek, G. Brown, and R. B. Gill.

ABSTRACT

Estimates of pre-natal productivity were 1.68 fetuses: doe from deer
collection data, and 1.67 fetuses per doe from highway killed deer.
Fawn survival was estimated at 73.2 percent from birth to the October
hunting season and 60.7 percent to January.
Ratios of male fawns:
female fawns were greater than 1:1. Average litter size based upon
the 1970 deer collection data was 1.9 fetuses per litter. Winter
mortality was estimated at 1,135 deer in 1970 compared with 639 fir
1969 and 1,706 for 1968. Mortality appeared to be positively related
to winter weather severity.
Mortality rates of the buck and fawn
categories generally exceeded their occurrence in the live deer postseason population.
Hunter harvest for the 1969 Middle Park deer
season was estimated at 4,151 deer compared with 2,104 in 1968 and
3,510 in 1967. Highway and railroad losses were tallied at 67 and 38
deer, respectively.

��- 339 -

MIDDLE PARK DEER STUDY PRODUCTIVITY AND MORTAILITY
R. Bruce Gill
P. S. OBJECTIVE
To estimate increments and losses to the Middle Park deer population in
order to formulate more efficient harvest regulations.
SEGMENT OBJECTIVES
1.
2.
3.

Estimate productivity rates of Middle Park deer.
Estimate mortality rates of Middle Park deer over the entire winter
range.
Estimate the magnitude of the hunter harvest.
METHODS AND MATERIALS

Methods and materials employed in this segment have been detailed previously
by Gill (1969).
RESULTS AND DISCUSSION
Productivity
Pre-natal Productivity
Thirty-four doe deer were collected between January 15 and May 31, 1970.
Eight does were collected from each of the sub-unit areas described by
Gill (1970), and two extra does were collected, one from the Blue River subunit and one from the Williams Fork River sub-unit. Locations of each
collection are depicted in Fig. 1. In addition, nine highway killed does
were collected and examined for pregnancy. Fetus:doe ratios averaged 168:100
from the collected does and 167:100 from the highway killed does (Tables
1 and 2). Pregnancy rates were 88.2 percent for collected does and 100.0
percent for highway killed does. Considering only pregnant does, fetus:doe
ratios were 190:100 from collected does and 167:100 for highway killed does.

Post-natal Fawn Survival
Gill (1970) reported a ratio of 93 fawns:100 does from Middle Park in midOctober, 1969. The true ratio of fawns:lOO does of reproductive age was
masked by a certain percentage of does (so-called "long yearlings") which
had not yet produced their first fawn. The proportion of these "long
yearlings" in the sample was approximated by the percentage of yearling does

�- 340 -

in the doe harvest sample (24.4 percent for 1969).
These were then subtracted from the doe sample and the fawn:doe ratio was recalculated as a
fawn:mother ratio.
The new ratio was 123 fawns:lOO mothers.
If we aSSume
that the pre-natal fawn:doe ratio of 168:100 was a true reflection of the
productivity of the doe population then 73.2 percent of those fawns survived
until the hunting season.
Conversely, the early fawn mortality rate was
26.8 percent.
The post-season fawn:mother ratio was calculated as 102:100.
Again, using
the collection ratio of 168 fawns:lOO mothers as the birth rate, the survival rate of fawns through the 1969 hunting season was 60.7 percent.

Doe Conception

Dates

All undamaged fetuses from collected does were measured for crown-rump
lengths and aged according to the fetal growth curve presented by Hudson
and Browman (1959). Back-dating from the dates of collection, dates of conception were estimated for each fetus.
These data are depicted in Fig. 2.
Of the 53 fetuses which could be measured, 88.7 percent were estimated to
have been conceived during the period November IS-December 10, 1969. The
range of conceptions was from November 16, 1969-January 11, 1970, a span
of 58 days.

Litter Sizes and Fawn Sex Ratios
Comparisons of litter size frequencies for 1968 and 1969 are presented in
Fig. 3. The average litter size (excluding the 0 class) was 2.00 young
per litter in 1969 and 1.90 young per litter in 1970.
Fawn sex ratios have favored males during both 1968-69 and 1969-70.
All
samples, including doe collections, highway kills, and check station ratios,
reveal ratios in excess of 100:100 (Fig. 4). This has been particularly
evident in samples for the Blue River sub-unit.

Mortality
Winter

Mortality

Estimates of Total Losses - Estimates of winter mortality have been conducted in Middle Park since 1968. Mortality has ranged from a high of 1,706
deer in 1968 to a low of 639 in 1969 (Table 3). Gilbert et ale (1970)
reported that deer population changes on one specific study area in Middle
Park were significantly and negatively correlated to November-February
precipitation.
Similar relationships existed between weather factors and total
mortality estimates.
The winter of 1967-68 was the most severe of the three
year sample period, and winter mortality was highest both in total estimated
losses and in percent of the live population which died (Table 4). Similarly, 1968-69 was the mildest winter of the three, and only 639 deer died
or 7.0 percent of the post-season live population.
The results for this
past winter (1969-70) were confusing because even though the weather data

�- 341 -

closely approximated those for 1968-69, the mortality data were more similar
to 1967-68.
This confusion may have resulted in part from the occurrence of
a larger percent of the wounding loss on winter ranges in 1969 than the two
previous years.
The 1969 hunting season was characterized by abnormally
high snowfalls and deer responded by descending from normal fall ranges to
the upper and middle winter ranges.
This was where the hunters encountered
them, and presumably, this was where much of the wounding loss occurred.
By
the time thewintet
loss surveys were conducted (June and July) deer carcasses
were decayed so badly that cause of death could not be determined, and consequently, deer perishing from hunter inflicted wounds could have been
tallied as winter mortality.

Sex and Age Structure of Winter Lost Deer - A summary of the winter loss
estimates by sex and age groups is presented in Table 5. The averages over
the three year study period were:
24.0 percent bucks, 29.0 percent does,
and 47.0 percent fawns. These same three groups were represented in the
post-season live deer population as follows:
22.3 percent bucks, 42.2 percent does, and 35.5 percent fawns. Generally, mortality rates in the buck
and fawn categories exceeded the percent occurrence in the live population
while does died in lesser percentages than their occurrence in the population.

Confidence Limits - Confidence limits about total winter loss estimates
were similar to those discussed for live deer population estimates (Gill,
1970). When winter loss estimates were high, confidence limits were wide,
and when estimates were low, confidence limits were narrow (Fig. 5). Also,
as was the case with live deer estimates, frequency distributions were
highly skewed toward the zero end (Figs. 6 and 7), and means and variances
of each strata were related curvilinearly
(Fig. 8).

Hunting Mortality
Estimates of the total Middle Park deer harvest have been made in three
ways:
1) from hunter report card estimates for Grand and Summitt counties;
2) through the combination of hunter report card estimates for Game Management Units 18, 28, and 37, and check station data (Gill, 1969); and 3) from
a regression equation describing the relationship between the Idaho Springs
checks of deer from Units 18, 28, and 37, and the Grand and Summit
counties
report card estimates (Table 6). The report card estimates for Grand and
Summit counties indicated hunting mortalities of 33.0, 23.1, and 57.6 percent of the 1968, 1969, and 1970 post-season populations, respectively.

Highway

and Railroad

Mortality

Sixty-seven deer were known to be killed by automobiles in Middle Park during
the period July 1, 1969 - June 30, 1970, and 38 were killed along the railroad tracks during the same period (Table 7). This total is down 21.1 percent from the previous year, and is the lowest figure since 1966-67.

�Table 1.

Collection
Period

January

February

March
April

Number of fetuses per doe from thirty-four deer collected in Middle Park, Colorado, winter, 1970.

Mudd~ Creek
Collection
Fetuses
Number
M F Total

70-4
70-5

2 0
1 1

Williams Fork River
Fetuses
Collection
Number
M F Total

2
2

Troublesome Creek
Collection
Fetuses
Number
M F Total

70-1
70-2
70-3

o 0

1 1
1 0

2
0
1

o2

2
0

70-6
70-7
70-8
70-14

3 0
1 1
1 1
1 1

3
2
2
2

70-9
70-10
70-11
70-15
70-1-A

1 1
2 0
o0
1 1
1 1

2
2
0
2
2

70-16

? ?

2

70-17
70-19

o1

1
2

70-22
70-23

2 0
1 1

2
2

70-22
70-21

2 2
2 0

2
2

70-18
70-24

1 1
2 0

2
2

70-25
70-26

o 2

2
2

70-27
70-32

2 0
1 1

2
2

70-30
70-31

o 0
1 0

0
1

70-28
70-29

2 0
2 0

2
2

70-33

1 0

1

l3 5

18

10 3

13

9 2

l3

70-12
70-13

o 0

2 0
2 0

May
Totals

Blue River
Collection
Fetuses
Number
M F Total

76

13

Male Fetuses:100 Female Fetuses = 244:100

Fetuses:100 Does = 168:100

w

~
N
,

�- 343 -

Table 2. Number of fetuses per doe from nine highway killed deer, Middle
Park, Colorado, winter, 1970.
Collection
Area

Fetuses
F
Total

Collection
Date

Collection
Number

M

2 -3-70

70-1-RK

?

1

2

Haystack Gulch off Hwy. 9

2 -8-70

70-2-RK

1

0

1

1 mi. west of Parshall

2-25-70

70-3-RK

1

0

1

Williams Peak Road

3-15-70

70-4-RK

1

0

1

0.1 mi. east of Windy gap

4-18-70

70-5-RK

o

2

2

Junction of King Gulch &amp; Hwy. 9

5-1-70

70-6-RK

1

1

2

East edge of Parshall

5-1-70

70-7-RK

o

2

2

~ mi. east of Parshall

5-1-70

70-8-RK

1

1

2

3 mi. east of Hot Sulphur Springs

5-3-70

70-9-RK

1

1

2

1.2 mi. west of Parsha1

6

8

15

Totals

75:100

Male Fetuses:100 female fetuses

167:100

Fetuses:100 does

Table 3. Estimates of Middle Park deer winter mortality by sub-unit sampling
areas, 1968-70.
Sub-Unit
Sampling Area

1968

1969

1970

Muddy Creek

817

162

327

Blue River

394

186

603

Williams Fork River

439

192

106

Troublesome Creek

56

99

99

Totals
90 Percent confidence
limits

1,706 ± 1,028

639 + 284

678 - 2,734

355 - 923

1,135 + 588

547 - 1,682

�Table 4.

Relationships

between deer winter mortality and weather factors, 1968-70.

Date

Inches
Pcpt

Inches
Snow

Ave.
Max.
Temp.

Ave.
Min.
Temp.

Ave.
Temp.

Nov. 1967

0.71

15.0

43.5

13.7

28.5

Dec. 1967

1.17

22.5

26.5

3.0

14.8

Jan. 1968

0.33

9.0

25.2

-6.7

9.3

Feb. 1968

1.62

36.5

30.4

3.3

16.9

Total

3.83

83.0

Ave. 31.4

3.3

17.4

Measured
Winter
Mortality

Percent of
Live Deer
Winter Population

1,706

16.0%

I

Nov. 1968

1.43

13.5

35.4

7.7

21.6

Dec. 1968

0.68

12.5

31.1

2.1

16.6

Jan. 1969

1.75

32.2

36.7

8.7

22.7

Feb. 1969

0.28

5.0

30.1

0.7

15.4

Total

4.14

63.2

Ave. 33.3

4.8

19.1

Nov. 1969

0.35

5.0

37.5

8.2

22.9

Dec. 1969

1.35

26.5

31. 9

1.1

16.5

Jan. 1970

0.89

19.5

27.7

3.7

15.7

Feb. 1970

0.35

6.0

34.5

5.1

19.8

Total

2.94

57.0

Ave. 32.9

4.5

18.7

w
+:'+:'-

.

639

7.0%

1,135

15.8%

�- 345 -

Table 5. Sex and age structure of deer winter losses compared to structure of
the post-season live deer population.

Total
Percent

Post-Season Live Deer POEulation
Total
Does
Fawns
Bucks
Percent
Percent
Percent
Percent

51.0

100.0

23.3

40.4

36.3

100.0

26.3

42.1

100.0

22.3

41.9

35.8

100.0

13.4

43.3

43.3

100.0

20.2

45.1

34.7

100.0

24.0

29.0

47.0

100.0

22.3

42.2

35.5

100.0

Year

Bucks
Percent

Winter Lost Deer
Fawns
Does
Percent Percent

1968

27.4

21.6

1969

31.6

1970
3 Year
Average

Table 6.

Estimates of the Middle Park deer harvest, 1967-69.

Year

County Report
Card Estimates

Ratio-Deer
Check Estimates

Regression Equation

Regression Equation
Estimates

1967

3,510

3,494

X2 = 161.95 + 3.248 Xl

3,066

1968

2,104

2,237

X2

321.41 + 3.140 Xl

2,909

1969

4,151

4,321

X2

52.00 + 3.329 Xl

4,503

�- 346 -

,
•
,
N

w8:~

70-1

•

•• 20.-12

• 70-17

~0-26

Hot
Kremmling

ZU-~~70-2~

70-)2

•

70-27

"70-2)

Green

Figo 10 Locations
studieso

Mtn.

Res.

of deer collection

sites for Middle Park productivity

Sulphur
Sprgs.

�- 347 -

Nov
Nov Nov Dee
15- 20 21-25 26-30 1-5

Dee
6-10

Dee Dee
11-15 16-20

on

c

•..0 30
Q.

1970

Q)

u
C

0

u

..•..

20

0

•..c
Q)

U

L.
Q)

10

Q..

Nov Nov Nov
Dee
15-20 21-25 26-30 1-5

Dee Dee
6-10 "-15

Jon

" -15

Fig. 2. Distribution of conception dates of does collected in Middle Park
in the winters of 1969 and 1970.

�- 348 -

1969

70~

62%
en
Ck:
W
lI-

o

60~
50~
40~

I-

Z

w

U
Ck:
w
e,

24%

12%
IO~

0%
jf
O~~------A~------~A~----~A~------~~.-------~A

o

234
LITTER

1970

70~

en

SIZE

67%

60~

Ck:
W
lI-

u..

o

40~

I-

Z
w

30~

~

w
c,
16%
·12%

5%

O~~----+-

A

+-

~

~

I

~0%

I

2

~

4

o

A

A

A

A

LITTER SI ZE
Fig.. 3.. Litter size frequencies
1969 and 1970 winters.

of does collected

in Middle Park in the

�- 349 -

260 ~
240.
220.

DOE COl ucr IONS /

/

200 ~

/j~~

I 80 ~

fI" /

(J)

w

0

Q

160 ~

0
0

140 ~

0:::
w

e,

I

..•

COllECTIONS
&amp; HIGHWAY KI l lS

j

120 ~

(J)

w
~

-c

100 ~

~

80.
60.

40.
20~

O~
1966

•

1967

1968

•

1969

•

1970

Figo 40 Ratios of male fawns:lOO female fawns from doe collection samples,
highway kill samples, and check station sampleso

�CI::
W
W

~~,
~
~
~
~
~
~
~
~
~
~
~
~

c
L&amp;..

o
CJ')

CI::
c:c
~
W

::;:)

w

VI

o

"~

z

",~
",~
",,~

~
~
~
~
~~,
~
~
~

"",""

."~

~
~

"" ""

,"""""

40

o~------------~------------~----------~
1967-68

Fig. 5.

1968-69

1969-10

Middle Park deer mortality estimates, bracketed by the 90 percent confidence limits

g

�90,

90,

90

80 I

80'

80

&lt;t

70 ---.

70-'·

70

z

60-1\

60-1

60

50-11

50

:l\

40

fJ)
fJ)

&lt;t
~

o
:t:

o
i.LJ

fJ)

i.LJ

u
z
w

501

a::
a::

g 4°1
:::&gt;

U

30

u,
0

\

\

w
0:::
l.&amp;J

u

a.

20 ...•

\

20,

IZ

10-.

\

t-'

20

-,

,

",

,

ZT

O-r

i

I

,.

T

0

I

2

3

4

0

I

2

3

4

0

VI

30

o ,

Fig. 6

w

I\

10.

10
0
0

I

2

3

CLASSES

CLASSES

CLASSES

1967-68

1968-69

1969-70

Frequency distributions of deer winter mortality data.

4

�- 352 -

100

90
80
70
60

-

50
40

&gt;

u

30

z

w

:::&gt;

o
w

20

Q::

IL.

10

o

2

3

4

CLASSES
COMBINED

DATA 1968-70

Fig ••7.. Frequency distribution for the 1968-70 combined deer winter
mortality data ••

�- 353 -

35.0

30.0

25.0

20.0

LLI

o

z
c(

0::
c(

&gt;

15.0

10.0

5,0

o.ot.~~~;:~~~~------~~
0.00

0.50

1.50

2.00

MEAN
Relationships between strata means and variances of Middle Park
Figo 8
deer winter mortality data, 1968-70 combinedo
0

~
2.50

�- 354 -

Table 7.

Known highway and railroad losses, Middle Park, 1963-70.

Year

Highway
Losses

Railroad
Losses

1963-64

90

1964-65

385

1965-66

7

Totals

1966-67

56

132

188

1967-68

54

60

114

1968-69

77

56

133

1969-70

67

38

105

LITERATURE CITED
Gilbert, P. F., O. C. Wa11mo, and R. B. Gill. 1970. Effect of snow depth
on mule deer in Middle Park, Colorado. J. WildL Mgmt. 34(1) :15-23.
Gill, R. B. 1969. Middle Park deer study - population productivity and
mortality. Colo. Div. Game, Fish and Parks. Game Res. Div. Fed.
Aid Proj. W-38-R-23. Game Res. Rep. July Part 1, 105-122.
Gill, R. B. 1970. Middle Park deer study - population density and
structure. Colo. Div. Game, Fish and Parks. Game Res. Div. Fed.
Aid Proj. W-38-R-24. Game Res. Rep. (In press).
Hudson, P., and L. G. Browman. 1959. Embryonic and fetal development of
the mule deer. J. Wi1d1. Mgmt. 23(3):295-304.

Prepared by

,
.~~.~---'-',~~~~------.-.\-.-\-R. Bruce Gill
Wildlife Researcher

�July,

- 355 -

JOB PROGRESS

REPORT

S ta te of

..;:C..;:O..;:L;,;:O;,:;RA=D..;:O:...-----

Project No.

W-38-R-24

Deer-Elk

Work Plan No.

14

Job No.

Job Title

Investigations
4

Middle Park Deer Study
Food Habits and Physical Characteristics

Period Covered:

October

1967 through April 1970

Personnel:

R. Bruce Gill, Willard Travnicek, Robert E. Keiss,
Joe Griess, Paul Gilbert, and Laren A. Roper

ABSTRACT

Fat content of bone marrow in collected deer declined at an average
daily rate of .61 and .46 percent in 1969 and 1970 (30 and 32 deer
sampled, r=.2 and -.5).
Similarly kidney fat indices decreased .31
and .33 percent in 1969 and 1970 (r= -.2 for both years).
Antler
beam diameters for all age classes in 1967 had overlapping confidence
limits except in the 1 year age class. Mean weights of the left
antler of yearling bucks is 70.9 ::t- 6.6 and 77.5 ::t- 10.4 grams for 1968
and 1969 (P=.05).
The number of points per antler for yearlings in
1967-1969 are comparable, except in 1967 which had fewer I-point
antlers (14, 20, and 17 percent in each respective year).
Blood serum
collections need more refined techniques as total protein, calcium,
magnesium, and zinc components appeared erratic in the sample.
Supplemental data to determine if E1aephora schneideri is present
indicate no occurrence, however, the data are inconclusive due to
small sample size (30) and the techniques for collections.
Wehrdikmansia cervipedis did occur in 70 percent of these samples.

1970

��- 357 -

MIDDLE PARK DEER STUDY FOOD HABITS AND PHYSICAL CHARACTERISTICS
Laren A

0

Roper

Po So OBJECTIVE
To measure selected physical characteristics related to the assessment of
physical condition of deer and to determine forage preferences of mule
deer in Middle Parko

SEGMENT OBJECTIVES
10

20
3

0

Estimate forage preferences of deer from stomach content analyseso
Estimate forage preferences of deer from tame deer observationso
Measure selected physical characteristics
of deer believed indicative
of the state of physical conditiono

METHODS

Forage Preferences
Procedures

are reported

by Gill

Forage Preferences
Procedures

are reported

AND MATERIALS

- Stomach

Content Analyses

(1969)0

- Tame, Trained

Deer Observations

by Gill (1969).

Physical

Characteristics

Procedures are reported by Gill (1969)0
In addition to those procedures
listed by Gill, blood samples from each deer collected for reproductive
studies (WP 14, Job 3) are analyzed for total protein and the parts per
million of calcium, magnesium, and zinc. Also an incisor tooth is collected
for age estimations by counting the cementum annuli in the sectioned tooth
(Low and Cowan, 1963)0 To supplement the yearling antler collections described by Gill, the number of points for each antler on all yearling bucks
checked through hunter check stations is recordedo
In 1967 diameter measurements of the main beam of all bucks were taken as well as a count of
the number of pOintso
The ears of 30 deer were collected to check for the occurrence of Elaephora
schneideri and Wehrdikmansia
cervipedis by counting microfilaria with a
dissecting microscopeo

�- 358 -

RESULTS AND DISCUSSION

Forage Preferences - Stomach Content Analyses
Eighty-four deer stomachs have been collected and analysis is pending ••

Forage Preferences - Tame, Trained Deer Observations
Food habits studies with tame deer were continuedo
This year this aspect
of Job 4 was conducted in conjunction with Job 5.. A summary of major
diet components is provided in Table I.. Further information is given in
the Job 5 report.

Table 1. Percent of total diets (bites) of 4 tame mule deer grazed on 3 study
areas on Middle Park winter range, 1969-70.
Type

Scientific Name

Shrubs
55.3%

Forbs
2701%

Grasses
17 6%
0

Percent

Purshia tridentata
Amelanchier alnifolia
Symphoricarpos oreophilus
Artemisia tridentata
Chrysothamnus biscidiflorus
Chrysothamnus parryi
Rosa sp••
Other (2 species)

35.4
10••
9
7.3
.8

Eriogonum umbellatum
Penstemon caespitosus
Phlox bryoides
Gutierrezia sarothrae
Artennaria parvifolia
Phlox multiflora
Castilleja flora
Penstemon cyathophorus
Other (9 species)

15.4
5.6
1.7
1.5
.8

Agropyron spicatum
Stipa pinetorum
Poa fendleriana
Boutelou gracilis
Koeleria cristata
Unidentified
Other (2 species)

.4
.••
2

.2
.1

••
7

.5
••
2
••
9
6.5
4 ••
8

4.4
1••
2
••
1

.7
t

�- 359 -

Physical Characteristics
This report only includes data concerning the measurement of physical
characteristics that may be indicative of the state of physical condition.
In 1969 collections began on January 16 and continued through April 15.
The mean fat concent for the 91 day period in January, February, March,
and April was 8705, 8001, 8900, and 6406 percent respectivelyo In 1970,
January 2Z-April 15 the mean fat content of the bone marrow was 6902,
7-7.0, 6906, and 5200 percent , The average decrease in bone marrow fat
is illustrated by plotting a simple regression (Figso 1 and 2)0 The
average decrease per day was .61 percent for 1969 and 046 percent for
19700 Apparently the winters of 1969 and 1970 were mild enough that
only a few deer had exceptionally low bone marrow fat content, and most
deer in the sample had fat content above 70 percent throughout the collection periodo
Plans are to test these data for curvilinear relationshipso
On March 5, 1965, 104 deer were collected in Middle Park (Keiss, 1965)0
Of these 50 had apparently died of malnutrition and starvation and 28
were in a weakened condition so that personnel captured them by hand and
sacrificed themo The average bone marrow fat was 606 percent in the
femurs and 25.4 percent in the cannon bone sampleso Fawns, yearlings,
and nine year olds all had femur fat below 3 percento The range for
deer in the yearling to 9 year age classes varied from 1203 to 36.9
percento This also includes deer that were killed by autos and trainso
When only deer that died of malnutrition or were caught and sacrificed
are included in the sample, the mean bone marrow fat of the femurs for
deer found on open range, at haystacks, and those caught and sacrificed
was 203, 09, and 101 percent.
If we can assume these data as representative of a state of physical
condition at or near death, the bone marrow fat content for deer collected
in 1969 and 1970 is below 35 percent in only 2 and 7 deer in 1969 and
19700 These comparisons may indicate that deer in 1969 and 1970 were
generally not suffering from severe malnutritiono
Kidney Fat Indices
The percent of kidney fat in deer collected in 1969 declined as the period
progressedo The mean for both kidneys was 3205, 3102, 1808, and 909 for
January, February, March, and April respective1yo In 1970 it was 3303,
2305, 1104, and 803 percento
Calculated regressions gave a mean decrease
in kidney fat per day of 031 and 033 percent for 1969 and 1970 (Figso 3
and 4)0 The correlation coefficient is -02 for both 1969 and 1970 data.
Plans are to analyze these data for curvilinear relationshipso
In white-tailed deer, Servinghaus et alo (1950) found evidence suggesting
antler growth and size are influenced by the quality and quantity of forage
available during the previous winter, particularly in yearling bucks and
less so in 2 year olds.

�- 360 -

100

t-

Z

w

U
0:::
W

Q..

b=-.61

Y = 79.8+ (- .61(x -46.2»

o

10

20

30

40

50

COLLECTION

60

70

80

90 100

DAYS

Figo 1. Regression of bone marrow fat in deer collected from January 15April 15, 1969.

�- 361 -

100

75

..-

z
w

U

0::
W
c,

50

b=-.46
A

r= 66.4 +(- .46( X-44. 7))

o

100

50
C.OlECTION

DAY!)

Figo 20 Regression of bone marrow fat in deer collected from January 26April 15, 1970
0

�- 362 -

80
75
70

b=-.31
V)

&gt;-

65

'9= 23.1 + (b)(X-43.9)

w

560
~
Iu..

55

w
....J
0 50

•
•

Z

-c

•

l-

J: 45

•

C&gt;
~
u..

40

•

0
~
u..

•
•

35

&gt;-

w

Z 30
0
~
I-

z

25

w

U

~

20

w
c,

15
10
•

•

•
•

•
••

5

• ••

0
0

10

20

30

40

50

COLLECTION

JAN ...•

j.- FEB ~j.-

60

80

90

---"I+-

APR

70

100

DAYS

MAR

Figo 3
Regression of percent of kidney fat in deer collected from January 15April 15, 1969
0

0

�- 363 -

60

•
55 •
•

(/)

&gt;-

w 50

b=-.33

Z

~= 1Z69 + (b)(X-41.9)

o
~

45

lLL.

W

-' 40

•

•

•

•

o

Z

-c 35
I-

•

J:

o 30
~

LL.

o 25

&lt;
LL.

Z 20

w

U
0::

w
c,

15
10

5

o+---~~~--~--~--~--~--~~~--~o
10
20
30
40
50
60
70
80
90
100
DAYS FROM STARTOF COLLECTIONS JAN. 26, 1970

JAN.f.-

FEB

~I+-

MAR

~I+-

APR

~I

Figo 40 Regression of percent of kidney fat in deer collected from January
26-April 15, 19700

�- 364 -

Initially in this study, antlers on deer in all age classes examined at
check stations were measured 1 inch above the burro
It was hoped this
would serve as an indication of the state of condition the animals were
In these data the 95 percent confidence
in during the previous wintero
limits overlap in all age classes except I-year old bucks (Fig 5)0
g

Since the first year's collection of data in 1967 it was decided to collect the left antler from as many yearling bucks as possible to determine
the mean mass in grams for this age class of deer
In 1968 and 1967 the
mean weight was 70.9 t 6.6 and 77.5 t 10.42 grams (P=o05)0 Using the
pooled variances for each set of data, a t-test for independent pairs
of data indicates there is no significant difference between the means
(Snedecor and Cochran, 1968)0
g

Our decision to use only data from yearling bucks has since been supported
by Anderson and Medin (1969)0 They suggest that for Rocky Mountain mule
deer yearling antler mass is the most sensitive and reliable indicator of
the condition of animals in the previous wintero
In conjunction with gathering antler mass information, the number of
points on yearlings was also recordedo
The average in 1967, 1968, and
1969 is presented in Figure 60 These data suggest fewer spike bucks
occurred in the kill in 1967, however, the data haven't been tested for
significance to date. In 1967 the percent yearlings with 1, 2, 3, and
4 point antlers were 14, 75, 10, and 1 percent, respectivelyo
The 1968
sample has 20, 71, 9, and 0 percent, while in 1969 the sample shows 17,
71, 11, and 1 percent.

Blood Serum Analysis
Blood samples obtained in 1969 from deer collected during January-April
were analyzed for total protein and parts per million of calcium, magnesium,
and zinco The results are erratic and appear inconclusive due mainly to
the lapse in time from collection of the sample and its arrival at the
laboratory for analysiso
Better sampling techniques are planned to alleviate these problems.
Table 2 contains a summary of the datao
Supplemental

Data

During the 1969 hunting season 30 deer were sampled for the occurrence of
Elaephora schneideri and Wehrdikmansia cervipedis microfilaria by collecting
ear samples from deer examined in check stationso
No evidence of ~o schneideri
was found, however, the collections were made from hunter killed deer which
had been subjected to freezing and thawingo
This could have obscured the
presence of this parasiteo
In future collections the forehead region may
give more reliable data, especially if analyzed soon after the animal is
collectedo
Thirty percent of the 30 samples had ~o cervipedis present.
The largest sample (22) of deer were from the Blue River game management
unito They had an incidence of 82 percent Occurrence of this apparently
harmless parasite (Table 3)0

�- 365 -

n=3

45

40

30
E
E
Z
~

w

tW

30

~

-c
0

25

20
n==180

2

3

4

5

6

7

8

9

10

AGE IN YEARS

Figo 50 Antler beam diameters and 95 percent confidence limits indicated by
the vertical line at each point (1967 ~ata).

�1 fbint =~

2 Points=D
3 Points=(])

100

4 Points=1I
90

80
70
I-

Z

60

w

u 50
~
W

W
0\
0\

0-

40
30
20

.

10

~~:.:

0

;l{~~t

.:.;::;:

1967

1968
YEAR

Fige

6e

Average number of pOints for yearling bucks in 1967, 1968, and 1969.

1969

�- 367 -

Table 20 Blood serum protein, calcium, magnesium, and zinc content of
Middle Park deer collected in 1969, January-Aprilo

Parts per Million
Total Protein

CA

MG

ZN

January (Sample Size: 6)
Maxo
Mino
Mean
February (Sample Size: 6)
Maxo
Mino
Mean
March (Sample Size: 8)
Maxo
Min.
Mean
April (Sample Size: 9)
Maxo
Mino
Mean

Table 3
Summary of the occurrence of Wehrdikmansia cervipedis microfilaria
in mule deer in Middle Park.
0

Herd Unit
18
27
28
37

No. in Sample
2
5
1
22

Number Infected

1
1
1
18

Percent Infected

50
20
100
82

�- 368 -

LITERATURE CITED
Anderson, Ao E., and Do E. Medino 1969. Antler morphology in a Colorado
mule deer population. Jo Wi1d1 Mgmt. 33(3}:520-533
0

0

Gill, Ro Bo 1969. Middle Park deer study: productivity and mortality.
Game Research Report. July, part two. PPo 141-1440
Keiss, Ro Eo 1965
Notes on deer and elk nutrition: winter 1964-65,
Coloradoo Unpubl. report, Coloo Game, Fish and Parks Depto 24 p.
0

Low, W. Ao, and I. McT. Cowano 1963. Age determination of deer by
annular cementum structure of dental cementumo Jo Wildlo Mgmto
27(3) :466-471.
Snedecor, G. Wo, and W. G. Cochran. 1968. Statistical methods.
Iowa State Press. 6th ed., 2nd printingo 593 po

The

Severinghaus, Co W., H. F. Maguire, Ro A. Cookingham and J. E. Tanck.
1950. Variations by age class in the.antler beam diameters of
white-tailed deer related to range conditionso No Amero Wildl.
Conf., Trans. 15:551-5700

Prepared

41/1Kr-

Wildlife Researcher Candidate

�July, 1970

- 369 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-24

Deer-Elk

Investigations

Work Plan No.

14

Job No.

5

Job Title

Middle

Park Deer Study - Range Fertilization

Period Covered:

May 1, 1969 through June 15, 1970

Personnel:

R. Bruce Gill, Paul F. Gilbert, o. C. Wallmo, Don W.
Reichert, Delwin Benson, Wayne Paintner, Robert Davies,
and Len H. Carpenter.
(Photos by Paul F. Gilbert).

ABSTRACT

Three study areas on native sagebrush range in western Grand County,
Colorado were selected and fenced.
Pre-treatment herbage production
and botanical composition measurements were made.
Production estimates
were made with an electronic capacitance meter and estimates ,of total
production of 1,522 pounds per acre at study area 1; 2,170 pounds of
herbage at study area 2; and 1~987 pounds of herbage at scudy area 3
were obtained.
Density measurements to give a measure of botanical
composition were made on permanently established plots at each study
area. Nitrogen fertilizer in the form of pelleted ammonium nitrate
at the rates of 0, 30, 60, 90, and 120 pounds of actual nitrogen per
acre was applied in late October, 1969. Pre-treatment grazing data
were gathered at each study area using tame, trained mule deer.
2,4-D herbicide treatments at the rates of 0 and 2 pounds per acre
were applied in early June 1970.

��- 371 -

MIDDLE PARK DEER STUDY
RANGE FERTILIZATION
Len H. Carpenter
This report presents the design, methods, and pre-treatment data gathered
for the first year of this study.
P. S. OBJECTIVE
To test the effects of fertilizers on deer forage and deer feeding intensity
to fertilization of critical winter ranges.
SEGMENT OBJECTIVES
1.
2.

To test the effects of nitrogen and 2,4-D on forage production on selected deer winter range areas.
To evaluate deer foraging responses to treated areas.
METHODS AND MATERIALS

Three study areas on native sagebrush range near Kremmling, Colorado were
chosen for this study (Fig. 1). These study areas were chosen so that
representative aspects of the mule deer winter range would be tested. The
treatment design chosen for each study area consists of 5 nitrogen rates in
combination with 2 rates of 2,4-D herbicide. This results in a randomized
block design of ten treatments replicated three times. Each of the ten
treatments was re-randomized for each study block (Fig. 2). The treatments
are as follows:
Fertilizer
1. 0 pounds elemental nitrogen per acre.
2. 30 pounds elemental nitrogen per acre.
3. 60 pounds elemental nitrogen per acre.
4. 90 pounds elemental nitrogen per acre.
5. 120 pounds elemental nitrogen per acre.
Herbicide
1. 0 pounds 2,4-D per acre.
2. 2 pounds 2,4-D per acre.
The nitrogen treatments were applied by whirlybird hand fertilizers in the
form of pelleted ammonium nitrate fertilizer (33.5% N.). The fertilizer
was applied at the Junction Butte site on October 30, 1969, the Flight1ine

�- 372 -

site was fertilized on November 2, 1969, and the Corral Creek site was
fertilized on November 3, 1969. Fall fertilization Was done so that maximum soil moisture would be available.
Herbicide treatments were applied by
a back-pack sprayer modified with a five foot boom with four nozzles.
A
nearly constant air pressure was maintained in the sprayer to assure even
application.
The 2,4-D was applied with water as a carrier at the rate of
five gallons per acre. The herbicide was applied on June 6, 1970 at the
Junction Butte and the Flightline Site. The Corral Creek site was treated
on June 7. Phenology of the vegetation at the time of application was favorable to obtain a successful big sagebrush kill. Weather conditions were
excellent at the time of application.
The 2,4-D, in combination with nitrogen fertilizer, was incorporated into
this study so that a maximum vegetational change could be obtained which
would hopefully result in more herbage being produced per unit acre. Much
of the area under study supports old, decadent, unproductive sagebrush which
seems to stymie the younger more productive plants.
It was thought that by
reducing the competition of the sagebrush plants with the herbicide and
applying the nitrogen to stimulate the remaining plants the total production
of an area could be increased.
Each of the three study areas has been fenced with an eight foot high deer
proof fence to prevent wild deer and livestock use. Each study block consists of a fenced area 520' X 220'. The ten treatments are pOSitioned in
two tiers of five treatments.
Each treatment plot is 100' X 100' (Fig. 3),
resulting in a total treatment area of 500' X 200'. This allows a border
of 10' on all sides between the treated plots and the fence.

Vegetation

Measurements

Although all 10,000 feet of each plot was treated, only a 9,000 foot square
will be measured.
This was done to reduce any leaching or drift effect that
may have occurred from an adjacent treatment.
Herbage

Yield

Herbage yield measurements were made with an electronic capacitance instrument Model 10lX (Fig. 4). This instrument when placed in vegetation measures
the capacitance or ability of that vegetation to hold energy from the headproduced electromagnetic
field. As the mass of vegetation is increased, the
ability to hold energy increases in the linear relationship.
Therefore, the
capacitance is closely related to the mass or weight of that vegetation
(Currie, unpublished).
For a more detailed presentation of this instrument,
the reader is referred to Neal and Neal (1966).
The method used consisted of a double sampling system so that a certain
ratio of meter read plots will be clipped.
A ratio of five meter read plots
to one clipped plot was chosen for the pre-treatment data. A total of 50
subplots was read for each 100' X 100' treatment plot resulting in a total
of 10 clipped plots per treatment.
The location of these 50 subplots was
chosen in a restricted random fashion so that they would cover the entire

�- 373 -

treatment in a representative manner.
Basically, the 50 plots were divided
into six lines of seven plots and one line of eight plots evenly spaced
across the treatment plot.
The capacitance meter plot was l' X 2' and was in three dimensions since
the probes are 18 inches long. The procedure used for a clipped plot was
as follows:
first, the immediate vegetation around the edge of the meter
that was not part of the subplot was clipped and removed to avoid interference; next plastic golf tees were placed under the corner probes of the
instrument so that the exact subplot location could be relocated when the
instrument was removed to do the clipping.
After the golf tees were located,
a first meter reading ~as made and recorded of the total vegetation on the
subplot.
The meter was then removed and the leafy shrub components of the
subplot were clipped and sacked separately by shrub species.
The meter was
then replaced on the sUbplot and a second meter reading was made and recorded.
The meter was again removed and the grass and forb components of the plot
were clipped and sacked separately as grasses and forbs. Finally, a third,
or reSidual, meter reading was made and recorded.
The various meter readings
were taken to evaluate the ability of the meter to estimate the separate
components of the vegetation making up a subplot. All production data are
on an oven-dry weight basis and measurements are made as soon as possible
after peak vegetative production is reached.
The data were analyzed by a
double sample computer program.

Botanical

Composition

In order to assess the various vegetational changes that may occur due to
the treatments, some measure of the relative abundance Of each species was
needed.
This measure might include frequency, density, basal area charting
or vegetational mapping.
To reduce the sample size needed for this measure,
permanent square foot density plots were established.
This will permit
measurements of a known area each year.
The term density in this report
means number of plants of each species per unit area.
Only six of the ten treatments on each study area were used for the density
measurements.
The six treatments chosen were the 0, 60, and 120 pound nitrogen treatments both with and without 2,4-D.
These six treatments represented the whole range from none to maximum application.
On each of the
above mentioned plots, 50 permanently marked square foot sample plots were
established in a restricted random fashion similar to the production subplot location method.
These plots will be read once each year and the
number of plants of each species rooted on the plot will be recorded.
In
addition, shrub heights are measured in tenths of feet so that changes can
be detected.

Deer Grazing Measurements
In order to measure mule deer response or reaction on these various plots,
tame, trained mule deer, property of the Rocky Mountain Forest and Range
Experiment Station, United States Forest Service, were used.
These tame

�- 374 -

deer allow strict control of grazing times and intensities.
Two types of
measurements were made with the grazing deer.
In order to measure specific
plant species and plant part preferences, a method was used which follows
that of Wa11mo and Neff (1970) where the deer were observed at close range
and the plant species and plant parts chosen were recorded (Fig. 5). The
other measurement made with the deer was a measure of treatment preference
(if any). This measure was made by releasing the deer into the study area
and observing and recording from a lookout tower the particular treatments
on which the deer graze.
Each study area grazing period corresponded to
the time that wild deer were using that portion of the winter range. The
study areas at the higher elevations were grazed in November and December,
while the lower study area was grazed in January and February.
The length
of the grazing periods varied from three to four days.
To obtain maximum data, each of the above two methods was alternated throughout a day by two hour grazing intervals.
This allowed three such intervals
in the course of one day, two intervals of one method and one interval of
the other method.
The order of this system was randomly chosen for each
study area. The deer were maintained on a minimal concentrate ration throughout the grazing periods.
Each evening the deer were removed to the central
holding pen for the night.

Plant Preference

Measurements

The plant selection method was used to obtain specific information as to
what individual plants and plant parts the deer choose on each treatment.
This method involved following the individual deer and identifying the
plant species that were being eaten.
Four tame deer and two observers were
used for each grazing interval.
However, only one deer per observer was
used in gathering data at anyone
time. The information obtained was transmitted into portable tape recorders and later transcribed onto forms
(Appendix I). Pertinent data gathered were the treatment number, the clock
time that deer entered the treatment, direction of travel, plant species
eaten, plant part eaten, and the number of bites of that particular part
eaten.
Other supplemental information such as behavior and urination and
defecation rates were recorded.
As these data were transcribed from the
tape recorder a map was constructed showing the grazing travel of that particular deer throughout the study area (Fig. 6). At the start of each
grazing trial a randomly chosen point in the exc10sure was used for the
starting point.

Treatment

Preference

Measurements

The method used to obtain treatment preference involved observing the deer
from an observation tower (Fig. 7). Each study area has a tower 10 feet
in height located midway along one side of the exc1osure.
Two observers
were present in the tower for each grazing interval and again, four deer
were used.
Each observer was responsible for observing two deer. The time

�- 375 -

spent on each plot was recorded (Appendix II) for both deer by clock minutes
while the grazing seconds were obtained for only one deer by using a tally
counter punched manually as that particular deer grazed.
This was done to
help tabulate the time spent on each plot activity, such as grazing or bedding time.
In addition, a map was drawn of the route of each grazing deer
to better understand grazing patterns.
Each observer changed deer between
trials and each deer was alternated on the data gathering method as to feeding seconds data or simply position data.

Other Measurements
Weather

Information

Weather information will be obtained from existing weather stations at Hot
Sulphur Springs and Kremmling.
Total moisture, moisture type, periods of
precipitation, and temperature records will be used to help interpret the
findings of the study.

Soil Descriptions
Soil descriptions for each study area were made by L. A. Fletcher, Soil
Scientist for the Soil Conservation Service.
These descriptions included
the pH of the soil at various levels, percent coarse material, soil profile,
soil temperature, soil
textures, drainage and permeability characteristics
and general aspects of each study area.
In addition, a soil chemical analysis was made for each study area at the soil testing laboratory at Colorado
State University.

Photographs
A permanent photo hub has been established on each treatment of each study
area (Fig. 8). This photo hub consisted of a meter square frame plot positioned on each treatment to delineate a specific photo area.
Both black
and white and color photographs were taken and will be repeated annually.
Plant Collections
A complete plant collection for each study area was made and verification
was obtained on the identification of species from the U. S. Forest Service
Herbarium in Washington, D. C.

DESCRIPTION
Junction

OF AREA

Butte Study Area

The Junction Butte study area is located on Colorado Division of Game, Fish
and Parks land and is approximately
four miles southeast of Kremmling, Grand
County, Colorado (NE~, Sec. 28, TIN, R8OW).
The site is on dry, native

�- 376 -

sagebrush range and is located on a long slope of about 8 percent. The
aspect is southerly with steep slopes of shallow soils and rock outcrops
immediately above the study area. The soil is well drained with a permeability of between .8 and 1.25 inches per hour. Annual precipitation is
about 11 inches of which about one-half occurs during the growing season.
Flightline Study Area
The Flightline study area is located about 20 miles northeast of Kremmling,
Grand County, Colorado (SW~, Sec. 36, T2N, R79W). The site is on native
sagebrush range located on a gently sloping to sloping uplands. The soil
is well drained with a permeability of between .2 and .6 inches per hour.
Annual precipitation is about 18 inches with about one-half falling during
the growing season.

Corral Creek Study Area
The Corral Creek study area is located about 21 miles northeast of Kremmling,
Grand County, Colorado (NE~, Sec. 36, T2N, R79W). This study area is located
on public land administered by the Bureau of Land Management. The site is
on native sagebrush range located on a sloping mountainside with an average
slope of about.8 percent. The aspect of this area is westerly. The soil is
well drained with a permeability of between .08 and .2 inches per hour.
Annual precipitation is about 18 inches again with about one-half of this
amount falling during the growing season.

RESULTS AND DISCUSSION
Vegetation Measurements
Herbage Yield
Table 1 summarizes the correlation coefficients obtained for each componen.t
meter reading for each of the study areas. High correlation coefficients
(.75 - .92) were obtained for all study areas for total weight estimates and
shrub component estimates. However, the forb and grass component correlation coefficients were not good (.37 - .53). In certain cases, the correlation coefficients were improved by subtracting the residual meter readings,
but generally, the benefit from this does not warrant the extra meter reading.
For this first year of pre-treatment data all ten treatments were analyzed
as one large sample to obtain total production in pounds per acre. This
resulted in an N of 100 for the small sample and an N of 500 for the large
sample for each study area. The following estimates were obtained. The
data from study area 1, the Junction Butte site, gave a mean clipped weight
per plot of 31.74 grams of herbage. This resulted in a total production
estimate of 1,522 pounds of herbage per acre. The variance of the mean was
computed to be 3.29 and the standard deviation of the mean was estimated to
be 1.81. The 95 percent confidence limits for the estimated total production were 1,349-1,695 pounds per acre.

�- 377 -

,
•
,
N

Study

Area

Area

3
Hot

Sulphur
Springs
CStudy

A rea 1

Scale: 1 in. =5 mi.

Fig. 1.

Location of the three study areaS.

�- 378 -

Junction

Butte

120 # N.

90 # N.

o # N.

30 # N.

30 , N.

21~2,4-0

0# 2,4-0

2# 2,4-0

2# 2,4-0

o./F 2,4-0

120# N.

0# N.

90# N.

60# N.

60# N.

0# 2,4-0

0# 2,4-0

2# 2,4-0

2# 2,4-0

0# 2,4-0

90# N.

30# N.

601 N.

120# N.

90# N.

2# 2,4-0

0# 2,4-0

0# 2,4-D

2# 2,4-0

0# 2,4-0

0# N.

0# N.

301 N.

60# N.

120# N.

0# 2,4-0

2# 2,4-D

2# 2,4-0

2# 2,4-0

0# 2,4-0.

120# N.

30# N.

90# N.

90# N.

120# N.

0# 2,4-0

0# 2,4-0

01 2,4-0

2# 2,4-D

2# 2,4-D

301 N.

60# N.

0# N.

0# N.

60# N.

21 2,4-D

0# 2,4-D

0# 2,4-D

2# 2,4-0

2# 2,4-D

\•

,

Flightl ine

Corral

Fig. 2.

Creek

Randomized

location of the ten treatments

N

on each study area.

\

•
,

�- 379 -

lOl.
220'

1

11-----------

520'

1--100'--1

Fig. J. Plot size and physical description of a stuQy area.

I

�- 380 -

Fig. 4. Electronic
measurements.

capacitance instrument Model 101 X used in herbage yield

�- 381 -

Fig.

5. Use of tame, trained mule deer in obtaining

plant selection

data.

�- 382

-

,.

~
/

./

t;7

V

( ,

l

~

.=Start
e=End
==Gate
A-Observation Tower
Fig. 6. Typical grazing patterns traveled
mule deer at the Flightline study area.

during grazing

trials by tame

�- 383 -

Fig. 7.

Observation

tower used in deer grazing trials.

�- 384 -

Fig. 8.

Hub for permanent photo record.

�- 385 -

Table 1. Correlation coefficients
ments with the capacitance meter.

Type of Meter Reading

obtained

Junction

for herbage

production

measure-

Study Areas and Coefficients
Butte
Flightline
Corral Creek

Total herbage weight estimation
(Total weight vs. total meter
read)

.75

.88

.88

Total herbage weight corrected
(Total weight vs. total meter read
minus residual meter reading)

.85

.86

.89

Shrub weight estimation (Shrub
weight vs. total meter read, Minus
shrub meter reading)

.83

.92

.91

Forb and grass estimation (Forb
and grass weight vs. shrub removed
meter reading)

.15

.46

.31

Forb and grass estimation corrected
(Forb and grass weight vs. shrub
removed minus residual reading)

.37

.38

.53

The data from study area 2, or the Flightline site, gave a total mean clip
weight of 45.16 grams per plot. This resulted in a total production estimate of 2,170 pounds of herbage per acre. The variance of the mean was computed to be 3.91 and the standard deviation of the mean was computed to be
1.98. The 95 percent confidence limits for the estimated total production
were 1,978-2,359 pounds per acre. Data from the Corral Creek site resulted
in a total mean clip weight of 41.38 grams per plot. This projected to a
total production estimate of 1,987 pounds of herbage per acre. The variance
of the mean was computed to be 3.04 and the standard deviation of the mean
was calculated to be 1.74. The 95 percent confidence limits for the estimated total production were 1,819-2,154 pounds.
As a result of the poor correlation coefficients for the forb and grass
components, no production estimates for these have been made.
It appears
that these estimates will have to be made completely by analyzing the forb
and grass clipped weights.
The three replicates allow a total of 30 such
plots for each treatment to make this estimate and plans for the following
year are to increase this to 15 clipped plots per treatment resulting in a
total of 45 clipped plots with the three replicates.

�-

386 -

Botanical Compositon
No pre-treatment data are presented for botanical composition. Since the
value of the permanently established plots will be to compare changes from
one year to another, the data gathered for the previous year are important
mainly for comparative purposes.

Deer Grazing Measurements
Plant Preference Data
A summary of the diets of the four tame, trained mule deer for each study area is
given in Tables 2, 3, and 4. The percent of the diet that each species makes
up is tabulated. No analysis of plant parts or individual deer differences
was made for these data. The diet composition represents all four deer for
all the grazing trials conducted during a grazing period at each study area.
Treatment Preference Data
Because the data taken are pre-treatment data, no treatment preferences
could be measured. However, it was felt that to better assess any changes
that might occur in grazing preferences it would be necessary to obtain Some
idea as to pre-treatment preferences in grazing area and plot locations. It
became obvious that the treatment block by the gate was visited most often.
Also, it seemed that each of the corner treatments at each study area was
preferred over the other treatments. This occurred primarily because they
seemed to prefer to stand in a fence corner for considerable portions of
time. Also, certain plots containing higher densities of plants such as
bitterbrush, Purshia tridentata, and serviceberry, Amalanchier alnifolia,
were visited more often. Data obtained from each study area are presented
in Tables 5, 6, and 7.

LITERATURE CITED
Currie, P. O. (unpublished). Development of the heterodyne vegetation meter
and partial test results on some range and wildlife vegetation types
in the Western U.S.A. Resume. 7 p.
Neal, D. L., and L. R. Neal. 1966. A new electronic meter for measuring
herbage yield. U. S. Forest Service. Res. Note PSW- 56. 4 p. Illus.
Wallmo, O. C., and D. J. Neff. 1970. Direct observation of tamed deer to
measure their consumption of natural forage. Range and wildlife habitat evaluation. A research symposium. U.S.D.A. Forest Service Misc.
Publ. No. 417. p. 105-110.

Prepared by ~~~·~L~~~n~H~~~·-~-f!~r-~-e~~·~e~r-:---------------Student Assistant

�-

387

-

Table 2. Junction Butte study area grazing observation data and diet
composition, grazing trial February 19, 1970 to February 22, 1970.

Plant Species

Ama1anchier
Agropyron

a1nifo1ia
spicatum

Symphoricarpos

oreophilus

Percent of Diet

36.5
16.5
12.6

Stipa pinetorum

6.3

Gutierrezia

5.7

sarothrae

Bouteloua

gracilis

4.6

Eriogonum

umbel1atum

4.0

Artemisia

tridentata

2.8

Phlox bryoides

2.8

Purshia

2.7

tridentata

Poa fendleriana

2.0

Artemisia

1.2

frigida

Chrysothamnus

spp.

0.8

0.5

Lichens
Berberis

repens

0.3

Penstemon

cyathoporus

0.2

Unknown

grass

Agropyron

smithii

0.2
0.1

Phlox multiflora

0.1

Koleria

0.1

Total

cristata

100.0

�- 388 -

Table 3. Flightline
composition, grazing

study area grazing observation data and diet
trial December 11, 1969 to December 14, 1969.

Plant Species

Percent of Diet

tridentata

57.2

Purshia
Eriogonum

12.6

umbellatum

Symphoricarpos

oreophilus

8.0

Stipa pinetorum

4.9

Agropyron

3.3

spicatum

Poa fendleriana

2.8

Penstemon

2.6

caespitosus

Phlox bryoides

2.3

Unknown

1.5

grasses

Castilleja

spp.

1.2

Amalanchier

alnifolia

1.0

0.4

Rosa sp.
Chrysothamnus
Penstemon

viscidiflorous

spp.

0.4
0.3

Phlox multiflora

0.3

Carex spp.

0.2

Unknown

forbs

0.2

Sitanion

hystrix

0.1

Penstemon

cyathoporus

0.1

Ko1eria

cristata

Artemisia

tridentata

Artemisia

tridentata

Total

0.1
0.1
spp. rothrockii

0.01

100.0

�- 389 -

Table 4. Corral Creek study area grazing observation data and diet
composition, grazing trial December 16, 1969 to December 18, 1969.

Plant Species

Percent of Diet

Purshia tridentata

33.6

Eriogonum umbel1atum

28.S

Penstemon caespitosus

14.1

Poa fendleriana

8.7

Stipa pinetorum

2.7

Antennaria parviflora

2.6

Arnalanchier alnifolia

2.3

Agropyron spicatum

2.2

Symphoricarpos oreophilus

1.7

Phlox rnultiflora

1.7

Chrysotharnnus viscidiflorus

0.6

Gilia candida

0.2

Unknown forbs

0.2

Senecio spp.

0.2

Senecio multilobatus

0.2

Juiperus scopulorum

0.1

Senecio integerrimus

0.1

Astragulus convallarius

0.1

Carex spp.

0.1

Aster spp.

0.1

Total

100.0

�- 390 Table 5. Junction Butte study area grazing observation data.
of feeding and location time per treatment plot.

Allocation

Plot No.

Total Minutes
on Plot

Total Minutes
Feeding on Plot

Percent of Total
Time Spent per Plot

1

207

8.27

9.9

2

301

36.95

14.5

3

186

19.77

8.9

4

195

24.37

9.5

5

661

60.15

31.7

6

136

0.00

6.5

7

304

16.72

14.6

8

7

0.00

0.3

9

21

0.00

1.0

10

64

0.00

3.1

Table 6. Flight line study area grazing observation data.
feeding and location tlineper treatment plot.

Allocation of

Plot No.

Total Minutes
on Plot

Total Minutes
Feeding on Plot

Percent of Total
Time Spent per Plot

1

398

13.35

20.1

2

57

10.88

4.0

3

79

9.57

5.2

4

24

4.78

3.5

5

14

0.47

7.3

6

409

38.08

19.5

7

352

12.95

16.8

8

17

1.57

4.0

9

55

14.60

6.4

10

155

28.33

13.2

�- 391 -

Table 7. Corral Creek study area grazing observation data.
feeding and location time per treatment plot.

Allocation of

Plot No.

Total Minutes
on Plot

To ta1 Minutes
Feeding on Plot

Percent of Total
Time Spent per Plot

1

334

39.3

23.0

2

11

0.0

0.8

3

299

22.0

20.6

4

14

2.2

1.0

5

50

0.0

3.4

6

446

29.8

30.8

7

23

0.5

1.6

8

64

5.4

4.4

9

60

0.0

4.1

10

149

0.0

10.3

�APPENDIX
Frrtilizrr Study:

I
Foragr Spluction

- 392

-

P &lt;l &lt;, r-.•. _-_.
{')

Date..... -- --. .......•-•.._-- _ ..•. _ .... _----_ ...•.._Hf'r,fn

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-,"::,::.l.::.'~~I:r~:~
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Rcco rd cr

e- ~

--'.

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Tr-mp '.

•

zrz:

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~-

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rr-c-

.0'

~~

O~O.

~-"--'.'.

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--'

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.~-'

._.

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I~.IV"&gt;...

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..,:..:-~";.':.::r••:=.:r~-•••.••...•....
-=------f..c"=";.rI..:,;:;,;r ••.•.••.

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Fer t Ll.L» cr

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Dat o .

- .._

- 393 -

::tudy:

FrE&gt;(&gt;

Gra:dne

\-lea t IH"'r Cond ,

-_._-- -.-----------_._---._--- .. -_ ....._-- --.. _---.
~.-.--.--... _-- .. --.....- ..-,.._-- .•------------ ..--_ .. _._~.•...-.
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__ w··

{indic te gn P, to~or, &amp;

••

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umhrrs

��July, 1970

- 395 -

JOB PROGRESS REPORT

State of

COLORADO
-----------~~~~~-----------

Project No.

W~38-R-24

Deer-Elk

Work Plan No.

14

Job No.

Investigations
6 (First Year)

Job Title

Middle
Physiology

Park Cooperative Deer Study
and Prevention of Deer Starvation

Period Covered:

June 1969 - June, 1970

Personnel:

Julius G. Nagy, Principal Investigator, James A.
Bailey, Investigator, Gene G. schoonveld, Graduate
Student.

ABSTRACT

Activities during this initial period of the projec~ involved construction of deer holding and handling facilities, acquisition of experimental animals, developing and refining techniques used in weighing
and obtaining blood and rumen samples, and conducting both short and
long term feeding trials.
There were indications that very high levels
of crude protein and NFE in a deer's diet can cause dietary upsets which
may possibly result in death.
The addition of protein to a high fiber
diet resulted in increased daily food intake and rate of passage,
although dry matter digestibility remained about equal.
Two adult deer,
fed an extreme malnutrition diet for 13 weeks were apparently able to
compensate for the poor quality diet by consuming more of the small
particle size feed. They maintained weight and remained in satisfactory
physical condition.

��- 397 -

PHYSIOLOGY

AND PREVENTION

OF DEER STARVATION

Julius G. Nagy

P. S. OBJECTIVE
To ascertain what takes place in deer physiology when starvation occurs, and
investigate the economic and biologic feasibility of preventing starvation.

SEGMENT
1.

Maintain

2.

Refine techniques for rumen content
for feeding, handling and examining

3.

Conduct metabolic
a.
b.

4.

b.
c.

5.

herd of 15 to 20 mule deer.
sampling
deer.

starvation

and improve methodology

trials:

To measure basic metabolic rate.
To determine the critical ambient
increase their rate of metabolism

Conduct
a.

an experimental

OBJECTIVES

temperature where deer must
to combat cold.

trials:

To detennine changes in deer physiology during the process of
starvation.
To determine the "critical" point of no return in deer starvation,
beyond which they will not recover.
To investigate methods of aiding deer recovery before the "critical"
point is passed.

Prepare report.

METHODS

AND MATERIALS

All deer on experimental trials are periodically weighed and samples of
blood and rumen fluid taken. Weighing is done by running the animal over a
scale and holding it there by means of a holding chute.
After weighing,
the animal is moved into a restraining chute constructed in a manner that
the animal is restrained by its head but access to the head, neck and right
or left regions of the body is possible.
Venous blood samples are easily
obtained by drawing blood from either right or left jugular veins using a
syringe of desired volume.
Rumen samples have been obtained using stomach tube technique which has
presented many problems and required much refinement.
PhYSical form of
diet, position in which the deer's head is held, excitability and period

�- 398 -

of time a wild deer can be restrained are problems encountered.
Rumen
samples can now be taken with relative ease, however this technique has
limitations which must be realized.
Varying amounts of saliva contamination
are always experienced and diets of different physical form, especially with
long fibrous particles, result in plugging the tube. Because of these limitations it would be more advantageous if rumen samples were obtained directly
through a rumen fistula.
Rumen fistu1ated deer allow for more frequent
rumen sampling regardless of form of diet with less injury to experimental
animals and elimination of saliva contamination.
Experimentation with
different rumen fistu1ation techniques and cannulas that can be applied to
deer are currently in progress.
To date, our technique and cannula have been
successfully tested using domestic goats; with slight refinements we believe
deer can also be successfully fistu1ated.

RESULTS
Maintaining

AND DISCUSSION
Experimental

Animals

All facilities for maintaining and handling experimental deer have been
constructed at Foothills Research Campus, Colorado State University." Six
large holding or exercise pens which have incorporated
within a group
isolation pen, fifteen individual isolation pens, work area for weighing
and handling deer and a research building are completed.
Figure 1 shows
actual physical layout of all facilities.
Efforts to obtain a tame herd of experimental deer were delayed by an
epizootic outbreak of Escherichia coli during summer of 1969 in our fawn
crop.
However, we were able to obtain experimental wild deer through cooperation with Colorado Game, Fish and Parks and U.S.D.I. Fish and Wildlife
Service.
Our fawn crop during the current year appears healthy; we have
experienced minimal losses and thus far results are very encouraging.

Refine Techniques

and Methodology

The techniques discussed under the Methods and Materials
devised and refined during the past segment.

Conduct

Metabolic

section were

Trials

Due to the loss of the fawn crop basic metabolic trials had to be postponed
but we concentrated instead on feeding trials using wild, adult deer.

Feeding

Trials

Trials of both short-term (21 day) and long-term (90 day) periods have been
conducted.
Four short-term trials designed to study effects various levels
of crude protein, fiber and nitrogen free extract (NFE) have upon digestibility and rate of passage through deer's digestive tract are completed.

�a WEIGHING AND HANDLING
b WORKING

CHUTE-

SPACE

W

\0
\0

OLA8
DIGESTION
CAGE
SHELTER

O

II
o

50
SCALE

Figure 1.

HOUSE
TRAI LER

100

1N FEET

Physical layout of deer holding and handling facilities.

�- 400 -

Initially the study was designed using four diets with combinations of
high and low crude fiber and protein (Table 1). The study was later changed
using two diets; deer on a dietary plane high in crude protein and NFE,
diets 2 and 4, were cyclic in daily food intake or off feed altogether.
Tentative results from these trials indicate very high levels of crude protein and NFE in a deer's diet can cause dietary upsets which may possibly
result in death.
Secondly, addition of protein to a high fiber diet resulted
in increased daily food intake and rate of passage although dry matter digestibility remained about equal.
Rate of passage was determined using stained
feed technique and determining 5-95 percent time required for stained particles to be eliminated.
Dry matter digestibility was determined through
use of digestibility cages and total fecal collection.
Table 2 compares
results obtained from two short-term feeding trials using diets 1 ahd 3.

Table 1. Proximate chemical analysis
feeding trials, oven-dry basis.

Proximate

Principles

of four diets used in short-term

Diet 1

Percent Composition
Diet 3
Diet 2

Diet 4

12.5

17.6

5.5

8.8

32.2

68.3

44.0

72.5

Crude Fiber

46.7

6.1

52.8

12.3

Ether Extract

2.5

2.9

2.5

3.0

Ash

6.1

5.1

5.2

3.4

Crude Protein
Nitrogen

Free Extract

A long-term trial was conducted, designed to study effects of an extremely
poor quality diet upon deer over an extended period such as experienced in
wild conditions during winter months.
Feed consisted of wheat straw and
cotton seed hulls which were ground and pelleted.
Diet used in this trial
is diet 3 (Table 1), also used in short-term trials.
Parameters investigated
were changes in daily food intake, weight change, serum urea, protein and
rumen pH, volatile fatty acids, total nitrogen, ammonia and total microorganism counts.
Chemical analysis of blood and rumen samples have not yet
been completed.
Weight loss for all experimental animals was not significantly greater than considered normal during winter months as reported by
other researchers, however, mean daily food intake increased considerably
during the trial as shown in Figure 2. It is theorized the animals were
able to compensate for a poor quality diet by consuming more small particle

�- 401 -

size. Experimental deer were therefore capable of maintaining weight and
remained in satisfactory physical condition throughout the trial in spite
of being fed what is normally considered an extreme mal-nutrition diet.
Research plans for 1970-71 include studying what effects form of diet,
particularly larger particle size, has upon digestibility, rate of passage
and feed impaction within the rumen. These studies will be closely related
with improving our fistu1ation technique and utilizing fistu1ated deer.

Table 2. Comparison of consumption, rate of passage and dry matter
tibility using two diets of different crude protein levels.

Animal

12.5 Percent Protein

5.5 Percent

Mean Daily Intake

Protein

(gms)

Deer 1

891

632

Deer 2

1449

1133

Deer 3

1173

1002
Rate of Passage

(hours)

Trial 2

36.5

57.5

Trial 3

45.5

54.0

Dry Matter

Digestibility

(percent)

Trial 2

42

40

Trial 3

44

47

Prepared

by --~T-------~----~~~--~~~¥-~
Nagy
Professor Wildlife

diges-

�- 402 -

2.4
~

2.2

2.0

1.8

'"'
eo

1.6

~

'-'
Q)

~

Cd

.IJ

1.4

p
H

e-,
M

~

1.2

Q)

~

1.0

p
Cd
Q)

~

0.8
1

3

2

4

5

6

7

8

9

10

11

12

HEEK
Figure

2.

Nean wcck Ly consumption
trial.

of tHO adult

dcc r , long-term

fc cdl n g

13

�July,

- 403 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-24

Deer-Elk

Investigations

Work Plan No.

15

Job No.

1

Job Title

An Investigation

of Deer-Auto

Accidents

Period Covered:

April 1, 1969 through March 31, 1970

Personnel:

Dale F. Reed, Dennis L. Money, Claudia A. Doose,
Kenneth C. Dillinger, William B. Zimmerman, Gary T.
Myers

ABSTRACT

Factors which might affect the number of deer-auto accidents were
measured on 20 miles of Highway 13 south of Meeker and on 18.5 miles
of Highway 82 near Glenwood Springs.
Deer densities, based on weekly
spotlight counts, averaged 125 near Highway 13 and 123 adjacent to
Highway 82. Most deer were present in both areas during March.
During
1969, automatic traffic counters recorded about 240,000 vehicles on
Highway 13 and 1,289,000 vehicles on Highway 82. Traffic on both highways was greatest in August.
Over 70% of the terrain near Colorado 82,
classified according to the number of contour intervals, was flat to
gently rolling and rolling, 10% moderately steep, 15% steep, and 3%
rugged.
Vegetation, typed within one quarter mile of both sides of
Highway 82, was 23% pinyon-juniper, 22% hayland, and 20% hay pasture.
Track counts, although poor, indicated a peak of deer migration across
Highway 13 around October 22, 1969. Colorado traffic accident reports
for 1965-1967 indicated that most deer-auto accidents occurred on
Sundays, and 47% accurred during the first three hours after sunset.
In addition to factors possibly affecting deer-auto accidents, the
number and location of deer killed in such accidents was recorded by
quarter mile interval.
Vehicles killed 109 deer on Highway 13 and
155 on Highway 82 during 1969.

1970

��- 405 -

DEER-AUTO ACCIDENT INVESTIGATION
Gary To Myers

Po So OBJECTIVE
Determine factors which cause deer-auto accidents on Highway 13 south of
the White River bridge in Rio Blanco County and on Highway 82 between
Basalt and Glenwood Springso
SEGMENT OBJECTIVES
1. Record all deer found killed by vehicles in the study area.
2. Estimate deer population densities adjacent to each highway.
3. Measure roadside deer activity along a 19 mile portion of Highway 13
and a 3 mile section of Highway 82.
4. Measure vehicle traffic volume and determine the time of heaviest traffic
of each highway.
5. Determine which driving and road conditions are most frequently associated
with deer-auto accidents.
6. Measure roughness of terrain adjacent to Highway 82.
7. Determine the type and acreage of each vegetative type within 1/4 mile
of Highway 82.
8. Measure the average speed of traffic on Highway 82.
9. Determine the time most deer-auto accidents occur.
10. Determine the time of peak roadside deer activity.
11. Measure general weather conditions in the study areas.
12. Measure the condition of deer killed by vehicles in the study areas.
METHODS AND MATERIALS
Methods used to determine the number and location of vehicle-deer kills (Fig.
1) and traffic volume have not changed (Myers, 1969)0 All other techniques
have been modified.
Deer Density Estimates
Deer density estimates were obtained by counting the number of deer within
sight of each highway at night with two spotlights attached to a vehicle
driven slowly up one side of the highway and down the other side. One
estimate was made each week on both highways except during summer on Highway
82. The number of deer seen in each marked quarter mile of highway was recorded.

�- 406 -

Fig. 1. One of many deer-auto
(Photo by Paul Gilbert).

accidents

occurring

each year in Colorado

�- 407 Deer Highway

Crossing

Estimates

A track strip was established in the median of 2-1/2 miles of Highway 82~
The strip, six feet wide, was rototilled and harrowed to a depth of about
two inches.
Deer tracks crossing the strip were counted and recorded
weekly during April, 1969 to determine the need for fencing to keep deer
off of the highway.
The time interval between erasure by harrowing and
the count was one day and night or roughly 24 hours.
A track strip about 19 miles long adjacent to Highway 13 was established
by bulldozing brush from an area at least six feet wide, disk plowing the
strip to a depth of four inches where pOSSible, and rototilling~
The strip
was also harrowed several times, and certain areas were hand raked to remove
rocks.
Deer tracks crossing the strip were counted and recorded at roughly
24 hour intervals, 14 times between May 16 and September 4, 1969, and 8
times between October 17 and October 26, 1969
Records of the number of
tracks seen crossing the strip in each marked quarter mile section of highway
were kept for each count on both Highway 13 and 82
0

0

Driving and Road Conditions
Driving and road conditions were recorded on Investigators Traffic Accident
Reports by the State Patrol when deer-auto accidents were reportedo
Copies
of these reports for 1968 and 1969 were obtainedo
Information regarding
driving and road conditions was taken from over 600 accident reports relating to deer-auto accidents occurring in 1968. This information was
coded, for summarization by computer.

Terrain Adjacent

to Highway

82

The area within 1/4 mile on each side of Highway 82 was divided into about
40-50 acre blocks on contour mapso
These blocks corresponded with the
quarter mile marked intervals of highway used to record the location of deer
crossings, sightings, and killso
Diagonals were drawn across each block
for each quarter mile section and the total number of 40 foot contour
intervals cut by both diagonals provided an index to measure slope steepness.
This index was used to place areas into six general categorieso
Blocks
having indexes from 1 to 10 were categorized as flat, 11 to 20 - gentle
rolling, 21 to 30 - rolling, 31 to 40 - moderately steep, 41 to 50 - steep,
and 51 to 60 ruggedo

Vegetation

Adjacent

to Highway

82

The fields designated on contour maps were drawn on aerial photographso
The various vegetative types containing 0 5 acres or more and within the
boundaries of the fields were designated on aerial photographs and, when
necessary, on foot to determine vegetative typeo Types were based on
descriptions set forth in Big Game Range Analysis Instructions (Interagency
Committee on Big Game Range Analysis, Unpublished Report, June, 1962)~
Quantity of vegetation was determined by planimetering each designated typeo
0

�- 408 Speed of Traffic
Two model NH~l Vehicle Speed Recorders were obtained from Newmeyer
Electronics, Glendale, Arizona
The Division of Highways installed buried
loops and provided detectors for use with the speed recorders to measure
speed of traffic at three different locations on Colorado 820 Eight loops,
two in each of the four lanes of traffic, were buried at each location to
enable measurements of speed of traffic traveling in various lanes and in
both directionso
Speed Recorders were operated intermittently during
February and March, 1969. The recorders provided information required to
calculate the average speed of traffico
o

Time of Deer-Vehicle

Accidents

The hour (Mountain Standard Time), day, and month of each accident reported
to the State Patrol during 1965-1967 was recorded on a code sheeto
Time
of accident was coded in relation to sunrise and sunset by use of maps based
on daily sunrise and sunset tables for five Colorado citieso
Accident times
were classified in one of 15 sunrise and sunset categorieso
All coded information was placed on punch cards and processed by computero
In addition, the time of death for each deer found killed on Highways 82
and 13 were estimated by use of thigh temperature, head temperature, rigormortis, or other characteristics
successfully used to estimate the time of
death of white-tailed deer (Gill and O'Meara, 1965)0

Time of Peak Roadside

Deer Activity

Deer detection devices were obtained and installed in the median of 1/4 mile
of Highway 82 to measure deer activit Yo These devices are described by Myers
(1970).

Weather
Weather instruments consisting of a recording rain gauge, maximum-minimum
thermometer, anemometer, and hygrothermograph
placed along each highway
were used to obtain general weather information.

Deer Condition
The condition of deer killed by vehicles in both study areas was determined
by kidney fat measurements described by Anderson (1965), and by measurement
of percent fat in the bone marrow of femurso
The later work was done by
personnel at the Research Laboratory in Fort Collins under the direction of
Robert E. Keisso

�- 409 RESULTS AND DISCUSSION
Number and Location
Highway

of Vehicle-Deer

Kills

13

Results indicate that at least 109 deer were killed on the 20 mile portion
of Highway 13 south of Meeker during 1969, compared to 118 in 1968. Thirtyone percent, or 34 deer, were killed in deer-auto accidents during October
(Fig. 2). Ten deer or less were killed during any other month with fewest
hit by motorists during December and January.
The greatest kill per mile
was 14 deer (Fig. 3). The highest kill per quarter mile was 8 deer in
Section 59. About 70 percent were hit between Sections 45 and 70.
Highway

82

One-hundred-fifty-five
deer were killed by vehicles on Highway 82 between
Glenwood Springs and Basalt during 1969 compared to 147 in 1968. Sixty-eight
percent of the kill, 106 animals, occurred during the first three months of
1969 (Fig. 4). Few deer were hit during the vacation months of May through
September, and 35 animals were hit during the last three months of 1969.
Over 25 percent of the deer (42) were killed in the second and third mile
south of Glenwood Springs.
More specifically, high kill areas were Sections
9, 8, 11, and 38 with kills of 16, 13, 9, and 9 deer respectively (Fig. 5).
The most concentrated kill per mile of highway included Sections 7-10 where
motorists collided with 39 deer during 1969.

Deer Density Estimates
Highway

13

One of the factors affecting the number of deer killed by vehicles was the
number of deer present near the highway.
A total of 6,538 deer was recorded
during 53 counts adjacent to Highway 13 in 1969 for an average of 125 deer per
count compared to 156 deer per count in 1968. Most were seen during late
winter and early spring.
The highest average, 285 deer, occurred in March
(Fig. 6). During all other months except May, June, July, and August, counts
averaged at least 95 deer per month.
Most deer (375) were seen in quarter mile
Section 78 (Fig. 7). One deer was seen in each of Sections 5 and 19.
Highway

82

A total of 4,047 deer was recorded adjacent to Highway 82 during 33 counts in
1969 for an average of 123 deer per count. About 90 percent of the deer were
present near Highway 82 during the first four months of the year.
Peak numbers
(402) were counted in March while counts averaged 10 or fewer deer during May
and September (Fig. 8). No counts were made during summer months.
The greatest
concentration of deer near one quarter mile of highway was 496 (Fig. 9). Over
2,000 deer, or 53 percent, were recorded along just three miles of highway.
Generally fewest deer were seen in Sections 41-60.

�50

040
lIJ

..J
..J

34

~

a::

~
•....
o

lIJ
lIJ

o
u,

o
m:
lIJ

ell

2

:)

z

o
JAN

Fig. 2.

FEB

MAR

APR

MAY

JUN

JUL

AUG

SEP

OCT

Monthly vehicle-deer kill on Highway 13 study area during 1969.

NOV

DEC

�o

o

2

o

9

2

2

4

2

3

II

II

8

12

14

II

8

:

3

: 6

Fig. 3. Location and number of deer killed by vehicles on Highway 13 study area during 1969. Each bar represents the
kill on one quarter mile of highway. Every fourth quarter mile section is numbered below the horizontal axis. Numbers
at the bottom of the graph are the numbers of deer killed in the four quarter mile sections between dotted vertical lines.
Quarter mile section number 1 begins at the Rio Blanco county line. Section 80 ends at the junction of Highway 13 with
Highway 64.

�50

040
UJ
-oJ
-oJ

-

~

9 r
30

.j:-

II II

t-'
N

0

u,
0

« 2
UJ
CD

:i

':)

2

10

0,--...1
JAN

FEB

MAR

APR

MAY

JUN

JUL

AUG

SEP

OCT

Fig. 4. Monthly vehicle-deer kill on Highway 82 study area during 1969.

NOV

DEe

�15

a:
l&amp;J
l&amp;J
C

l&amp;..

-

lOt

0

a:
l&amp;J
CD

2
:::&gt;

Z

5 I

"

18

6

16

:

13

. .

:

W

- --

_4_

o
24

~
to-'

9

9

13

:

II

8

4

.
19

:

6

3

2

6

6

2

?ig. 5. Location and number of deer killed by vehicles on Highway 82 studv area during 1969. Each bar represents
the kill on one quarter mile of highway. Every fourth quarter mile section of hLghway is numbered jus t beLow
the horizontal axis. Numbers at the bottom of the gra~h are the number of deer killed in the four quarter mile
sections ~etween dotted verticle lines. Section 1 is near Glenwood Springs citv limit. Section 74 is near the
3asalt citv limits.

�500

0

'"

IZ

:::)

0
0

I

-

a: aoo·~

~
•....

285

L&amp;J
1&amp;1
Q

"""

~
0
O!
1&amp;1

m
:IE
:::::&gt;

z

o
JAN

FEB

MAR

APR

MAY

JUN

JUL

AUG

SEP

OCT

NOV

DEC

~ig. 6. Average number of deer counted adjacent to Highway 13 study area by month
during 1969.

�7

3

228

G:
LLI
laI

JO·tI

0

~

I

180 ~

a: 150
I.IJ
en

137

15

I~

37

2

::)

z

SECTION NO. 0
DEER/MILE

24

: 114 : 203:

30

l 72

: 448

: 116 : 109 : 56

: 83

:

~ig. 7. Location of deer counted adjacent to Highway 13 study area in 1969.

,lit _-

,
+:t-'

VI
I

�0
W

I2

::&gt;

0

0

97'

•

FEB

MAR

a:
~ 300t"
0
U.

0

+'
t-'
0-

a:

ill

CO

:E

::&gt;
z

o
JAN

APR

MAY

JUN

JUL

AUG

SEP

OCT

NOV

DEe

Fig. 8. Average number of deer counted adjacent to Highway 82 study area by month during 1969.
No counts were made during months where * is shown.

�- 417 -

500

450

400

350

300

250
G:
1&amp;1
1&amp;1

N

0

0

N

•••• 200

0
IX

'"~

ell

::&gt;

z

~

~

100

50

o

4:
I:
16:
17 1 306! 274; 721154;

Fig. 9.

1969.

Location

196 :1I72~ 38 1178 1416 :

of deer counted adjacent

to Highway

82 study area in

�- 418 -

Deer Highway Crossing

Highway

Estimates

13

Results of track counts on Highway 13 were affected considerably by weather,
livestock activity in the track strip, texture and rockiness of the soil,
and other factors.
During a large majority of the track counts, at least
one and often several of these factors affected track. count accuracy.
Putting it bluntly, almost all of the track counts are inaccurate for one
reason or another, especially during October when frozen soil, mud, livestock drives, and other types of interference made it impossible to detect
deer tracks in large sections of the track strips.
Consequently readers
should interpret results with caution.
Twenty-two full length track counts were made on Highway 13 during 1969
(Table I). A total of 3,124 tracks were noted in the track strip during
May - October.
An average of 7 deer tracks were counted per mile of highway. The sixth and sixteenth mile of highway contained most tracks, 345
and 360 respectively.
An average of 95.7 tracks was seen per count in
May, 63.0 in June, 106.2 in July, 102.5 in August, 81.0 in September, and
223 5 in October.
0

Deer migrate from east to west across Highway 13 to their winter range
during October.
Eight track counts were made during this month in an
attempt to document when and where the migration crossed Highway 13. Considering only deer tracks which zema Ined- after subtracting east bound tracks
from west bound tracks, a minimum of 660 deer migrated across Highway 13
between October 16 and October 27, 1969 (Fig. 10). The peak of documented
migration during this period occurred around October 22. Important crossings
were in the sixth, tenth, eleventh, and sixteenth mile of highway where
roughly 60 percent of the documented migration crossed (Fig. II).

Highway

82

Crested wheat occurring near cover and water on the east side of about a
mile of Highway 82 may prevent mass movement of deer across the highway to
alfalfa fields.
If this is not the case and large numbers of deer cross
the highway, fencing may be required.
Track counts were conducted in the
median ofa mile of highway adjacent to the crested wheat area to get an
index of deer crossings.
Results indicate that there were no crossings on
May 7, 1969, and as many as 20 crossings on April 17, 1969 (Table 2).

�- 419 -

Table 1. Average number of deer tracks noted crossing the track strip
on Highway 13 during 22 counts in 1969.
Mile of
Highway

May

June

July

Aug.

Sept.

Oct.

Average

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20

4.7
1.0
0.7
2.0
17.3
9.0
2.3
0.0
0.3
1.0
3.7
5.0
4.3
9.0
2.0
4.3
8.7
6.0
1.7
12.7

0.0
0.0
0.5
1.0
4.5
9.5
2.0
1.5
0.0
1.0
2.5
2.0
9.5
5.5
3.5
6.5
1.0
0.0
6.5
6.0

5.3
1.5
1.5
1.3
1.3
6.8
2.5
10.8
2.2
4.0
9.2
6.7
10.5
8.3
5.3
7.5
5.7
4.8
6.0
4.8

11.5
2.5
1.0
2.0
3.5
16.0
8.0
2.0
6.0
12.5
5.0
2.0
13.0
5.0
1.0
2.0
4.0
2.0
0.5
3.0

18.0
0.0
0.0
0.0
7.0
19.0
3.0
1.0
0.0
1.0
21.0
1.0
0.0
3.0
0.0
0.0
0.0
0.0
3.0
4.0

7.6
16.3
8.8
1.3
3.5
25.9
0.6
0.0
1.5
17.5
19.4
25.0
10.1
11.9
25.0
35.6
3.1
4.9
1.4
4.3

6.7
6.7
3.8
1.4
5.0
15.7
2.3
3.3
1.7
8.9
11.7
12.0
9.2
8.9
11.2
16.4
4.3
4.1
3.1
5.6

Total
Tracks

287.0

126.0

637.0

205.0

81.0

1,788.0

3,124.0

Average

95.7

63.0

106.2

102.5

81.0

223.5

142.0

No. of
Counts

3

2

6

2

1

8

22

�- 420 -

16

WEST- EAST DEER

TRACKS - HIGHWAY 13- OCT.17-26,'69

.

148

140

120

100

f/)

~
0

&lt;I 80
0:
llL

0
0: 60

w
CD
~

:::&gt;

z
40
29

.-

20

17

18

19

20

21

22

23

24

25

26

27

OCTOBER
Fig. 10. Tracks counted which were made by deer migrating across
the track strip. Migrating deer are those animals moving west, for
which there is corresponding eastward movement.

�30

DEER

TRACKS

ADJACENT

TO HIGHWAY 13 (OCT. 17-26)

•

. TOTAL TRACKS CROSSING
BOTH WAYS
TRACKS MOVING WEST
ONLY (WEST- EAST
TRACKS)

O

25

200

200

200

+'
N
t-'

150

95

100
81

70

2B
12

0'

, , , , , , , , , , , 'r=r
I
2
3
4
5
6
7

"

, , , , , , , , , , , , , , , , , ,

F2

'

,

Mile section of
8
9
1011
12 13 14 15 16
Fig.
11.
Tracks
made
by
both
resident
and
migrating
deer
during October 17-26, 1969 track counts
Highway
adjacent to Highway 13.

�- 422 -

Table 20 Deer tracks counted in the median of Highway 82 during April
and May, 19690
Date of Count
10

AEril
17

30

May
7

Total

Average

25

0

0

7

0

7

1.75

26

2

1

0

0

3

0075

27

5

8

4

0

17

4025

28

3

11

5

0

19

4.75

Total

10

20

16

0

46

2088

Section Noo

Traffic Volume
Highway 13 had an annual traffic volume of about 240,000 cars with most
traffic in the months of June through October (Table 3)0 In comparison
the annual traffic volume on Highway 82 was about 1,289,000 carso Most
traffic on Highway 82 occurred during the summer vacation months (Figo
12). Putting it in simple terms, there was a maximum of about 41 vehicles
per hour on Highway 13 compared to a maximum of over 197 per hour on
Highway 82.

Driving and Road Conditions
Driving and road conditions associated with 1968 and 1969 reported accidents
are being coded for analysiso Results are not available at this timeo

�Table 3.

Traffic volume on Highway 13 and 82 study area during 1969.

Total

Number of Vehicles
AverageLDay
Hwy. 13
Hwy. 82

AverageLHour
Hwy. 13
HY7y. 82

Month

Hwy. 13

Hwy. 82

Jan.

13,272

87,144

428.13

2,811.1O

17.84

117.13

Feb.

12,958

85,206

462.79

3,043.07

19.28

126.79

Mar.

15,712

105,202

506.84

3,393.61

21.12

141.40

Apr.

16,412

93,242

547.07

3,108.07

22.79

129.50

May

19,043

100,291

614.29

3,234.19

25.60

134.80

Jun.

23,917

113,637

797.23

3,787.90

33.22

157.83

Jul.

27,859

133,096

898.68

4,293.42

37.44

178.89

Aug.

30,515

146,538

984.35

4,727.03

41.01

196.96

Sep.

23,318

113,812

777.27

3,793.73

32.39

158.07

Oct.

26,525

103,483

855.65

3,338 16

35 65
0

139.09

16,486

99,905

549.53

3,330 17

22.90

138 76

Dec.

14,343

107,413

462.68

3,464 94

19.28

144.37

1969

240,360

1,288,969

658.52

3,531 42

27.44

147.14

.j:'-

Nov.

0

0

0

0

0

N

w

�II HIGHWAY

13

HIGHWAY

82

o

197
20'0'

0'

JAN
1&lt;"
r i.g ,

1"

~L.

FEB

MAR

APR

MAY

JUN

JUL

AUG

SEP

O'CT NOV

Comparison of monthly traffic volume on Highway 13 and 82 study areas during 1969.

�- 425 -

Terrain Adjacent

to Highway

82

Over 50 percent of the land area within 1/4 mile of Highway 82 was
The remainder consisted
classified as flat and gentle rolling terraino
of 19 percent rolling, 10 percent moderately steep, 15 percent steep,
and 3 percent rugged terrain (Table 4)0

Table 40 Types of terrain within
Highway 82 study areao

1/4 mile of either

Quarter Mile Sections
Noo of
Sections

side of the Colorado

of Highway

Type of
Terra in

Terrain
Index

Flat

1-10

22

30.14

11-20

17

23 29

21-30

14

19.18

31-40

7

9 59

41-50

11

15 07

51-60

2

2.74

Gentle

Rolling

Rolling
Moderately
Steep
Rugged

Steep

Percent

0

0

0

Total

100 01
0

Vegetation

Adjacent

to Highway

82

The wrong conversion factor was used during planimetering and the error
was not discovered until report timeo As a result acreages are approximations subject to change.
Over 5,800 acres of vegetation located within one quarter mile of both
sides of Highway 82 were typedo
Types were comprised of 23 percent pinyonjuniper, 22 percent hay1and, 20 percent hay pasture, 13 percent sagebrush,
7 percent broadleaf trees, 6 percent browse, and lesser percentages of nine
other types (Table 5).

�- 426 -

Table 5. Acreage and percent of various vegetative
1/4 mile of the Highway 82 study area.

types present within

Vegetative
Type

Acres

Percent

Pinyon-juniper

1,367

23.4

Hayland

1,291

22.1

Hay Pasture

1,168

20.0

Sagebrush

742

12.7

397

6.8

Browse

350

6.0

Barren

164

2.8

Cultivated

123

2.1

Grassland

111

1.9

Culture

53

0.9

Annuals

35

006

Winterfat

23

004

Conifer

6

0.1

Orchard

6

001

Unknown

6

0.1

5,842

10000

Broadleaf

Total

Trees

Speed of Traffic
Speed data from tapes were compiled by two people. A comparison of their
work indicates that average speeds may vary a mile or so and sample size
from 5 percent to 6 percent due to interpretation of tape data (Table 6).
Results of reliable speed checks indicate that the average speed of traffic
ranged from 53.3 miles per hour on March 6, 1970 at station one to 60.0
miles per hour on February 23 at station two. Northbound traffic ranged
from 55.8 miles per hour to 59.3 miles per hour (Table 7).

�- 427 -

Table 6. Variation in speed data due to human judgment. Speeds below
were obtained by two people from the same speed tapeo Neither knew their
work would be checked against the work of another.
Individua 1 A
Average
Speed

Individual B
Sample
Average
Size
Speed

Hour

Sample
Size

1600-1659
1700-1759
1800-1859
1900-1959
2000-2059
2100-2159
2200-2259
2300-2359
2400-2459
0100-0159
0200-0259
0300-0359
0400-0459
0500-0559
0600-0659
0700-0759
0800-0859
0900-0959
1000-1059
1100-1159
1200-1259
1300-1359
1400-1459
1500-1559
1600-1659

80
108
72
89
40
39
28
17
13
9
4
5
5
6
18
50
105
101
110
95
90
111
80
101
30

56
58
54
58
59
58
55
54
55
59
53
52
54
56
55
55
58
59
59
58
58
56
56
55
58

82
109
72
87
39
38
28
17
13
9
4
5
5
6
20
50
89
95
111
95
90
108
80
114
29

55
57
54
57
59
58
55
54
55
59
52
52
54
56
56
54
58
61
58
58
58
56
55
58
59

1,406

56 9

1,494

55.8

Total

0

�- 428 Table 7. Speed data collected at three locations on Colorado 82 during 1970.
in parenthesis may be inaccurate due to equipment malfunction.

Figures

Date

Time of Data
Collection

Sample
Size(Car)

Average
Speed

Location

Direction
of Traffic

Feb. 3-4

1600-1600

1,429

57.7

Sta. 3

South

Dry Highway

Feb. 4-5

1600-1559

1,444

55.4

Sta. 3

South

Dry Highway

Feb. 5-6

1625-1625

1,494

55.8

Sta. 3

North

Dry Highway

Feb. 6

1200-2459

1,116

(57.7)

Sta. 3 North&amp;South Fig. may be
inaccurate due to
Equip. malfunc.

Feb. 12

1300-2400

822

59.7

unknown

South

unknown

Feb. 12

1300-1800

466

58.8

Sta. 2

North

unknown

Feb. 19

1500-2459

775

58.1

unknown

South

Dry Highway

Feb. 20

1000-2359

1,372

58.1

unknown

South

Dry Highway

Feb. 21

0900-1359

474

58.7

unknown

South

Dry Highway

Feb. 21

1945-2345

117

57.0

Sta. 2

South

Dry Highway

Feb. 22

0300-0700
1700-1800

255

58.3

Sta. 2

South

Dry Highway

Feb. 23

2100-2200
0500-0600

49

60.0

Sta. 2

South

Dry Highway

Feb. 27

0500-0700
1500-1600

212

59.3

Sta. 2

North

unknown

Mar. 4-5

1045-0145
0900-1000

877

(61.5)

Sta. 2

North

Dry Highway

Mar. 5-6

1030-1130

1,654

58.4

Sta. 2

South

Dry Highway

Mar. 5-6

1030-1130

1,692

55.7

Sta. 3

South

Dry Highway

Mar. 6

1100-2100

987

53.3

Sta. 1

South

unknown

Mar. 6-7

1130-1100

1,514

60.3

Sta. 2

South

unknown

Mar. 7-9

1100-1230

3,339

54.5

Sta. 1

South

Wet Highway

Mar. 7-9

1120-0900

3,172

(63.4)

Sta. 2

South

Wet Highway

Remarks

-------------------------------------------------------------------------------------------

�- 429 -

Table 7. Speed data collected at three locations on Colorado 82 during
in parenthesis may be inaccurate due to equipment malfunction.

Time of Data
Collection

Date

1970.

Figures

Sample
Size (Car)

Average
Speed

Location

Direction
of Traffic

Remarks

Unknown

Mar. 10-11

1000-0200

1,236

(66 2)
0

Stao

2

South

Mar. 17

1200-1500

206

57 0
0

Stao

3

South Snowing,wet,slush

Mar. 17-18

1300-1030

1,095

(76 3)

Stao

2

North
Snowing when put
on - Dry Highway when
off

Time of Deer-Vehicle

0

Accidents

Statewide
Over 50 percent of the deer-vehicle accidents in 1965-1967 occurred on Friday,
Saturday, and Sunday (Table 8). Almost 20 percent were on Sunday, 17 percent
on Saturday, and 16 percent on Friday.
This general pattern seems to hold
for all years except 1967 when slightly more accidents were on Friday rather
than Saturday.
Excluding Friday, roughly 12 percent of the deer-vehicle
mishaps occurred each weekday.
Almost 40 percent of the 1,441 deer killed by vehicles were hit between 6:00
PM and 9:00 PM Mountain Standard Time (Table 9). Fewest accidents, 7 5 percent, occurred between 9:00 AM and 5:00 PM. Looking at similar information
in another way, 48 percent of 1,624 deer-auto accidents during 1965, 1966
and 1967 which occurred at known times were during the first three hours
after sunset (Table 10). Over half of the accidents involving deer were
during the first hours past sunseto
The time of accident in relation to
sunrise and sunset varied from year to yearo
0

Highway

13 and 82

Combining information from Highway 13 and Highway 82, about 51 percent of the
deer-vehicle kills in 1969 occurred on Friday, Saturday, and Sunday (Table 11).
Over 20 percent were on Friday, 15 percent on Saturday, and 15 percent on
Sunday.
Fewest deer (9 6%) were killed on Monday.
0

�- 430 -

Table 8. Number of vehicle accidents involving deer and reported to
the Colorado State Patrol by day.
Day

No.

1965
Percent

Sunday
Monday
Tuesday
Wednesday
Thursday
Friday
Saturday

81
41
48
49
46
60
66

20.7
10.5
12.3
12.5
11.8
15.3
16.9

107
62
59

Total

391

100.0

Table 9.

Time of reported deer-vehicle accidents in Colorado.

Hour by
Military Time

No.

1965
Percent

1966
Percent

No.

1967
Percent

Total

Percent

55
93
103

19.4
11.2
10.7
13.1
10.0
16.9
18.7

126
84
88
72
92
111
110

18.4
12.3
12.9
10.5
13.5
16.3
16.1

314
187
195
193
193
264
279

19.3
11.5
12.0
11.9
11.9
16.2
17.2

551

100.0

683

100.0

1625

100.0

No.

72

1966
No. Percent

1967
No.
Percent

Total

Percent

3.69
2.25
3.28
3.07
5.12
4.10
1.84
1.64
1.23
2.25
.82
.41
1.02
.41
1.02
1.23
9.43
15.57
14.34
8.61
5.33
6.35
3.48
3.48

26
14
6
28
25
29
14
12
7
3
4
4
3
3
6
7
44
77
99
81
46
30
26
9

4.31
2.32
1.00
4.64
4.15
4.81
2.32
1.99
1.16
.50
.66
.66
.50
.50
1.00
1.16
7.30
12.77
16.42
13.43
7.63
4.98
4.31
1.49

53
29
33
48
64
63
34
27
15
22
11
9
12
5
16
19
126
189
208
168
105
91
57
37

3.68
2.01
2.29
3.33
4.44
4.37
2.36
1.87
1.04
1.53
.76
.62
.83
.35
1.11
1.32
8.74
13.12
14.43
11.66
7.29
6.32
3.96
2.57

99.97

603

100.01

1441

100.00

0100
0200
0300
0400
0500
0600
0700
0800
0900
1000
1100
1200
1300
1400
1500
1600
1700
1800
1900
2000
2100
2200
2300
2400

9
4
11
5
14
14
11
7
2
8
3
3
4

2.57
1.14
3.14
1.43
4.00
4.00
3.14
2.00
.57
2.29
.86
.86
1.14

5
6
36
36
39
45
33
30
14
11

1.43
1.71
10.29
10.29
11.14
12.86
9.43
8.57
4.00
3.14

18
11
16
15
25
20
9
8
6
11
4
2
5
2
5
6
46
76
70
42
26
31
17
17

Total

350

100.00

488

�- 431 Table 10. Number and percent of reported deer-vehicle accidents in Colorado in relation
to sunrise and sunset.

Time Interval

n t s
Dee
r - v e h i c 1 e A c c ide
Total
1967
1966
1965
No. Percent
No. Percent
No. Percent

Percent

Within 3rd hour
before sunset

4

1.0

1

.2

9

1.3

14

.9

Within 2nd hour
before sunset

5

1.3

10

1.8

10

1.5

25

1.5

At sunset or
within 1st hour
before sunset

11

2.8

17

3.1

27

4.0

55

3.4

Within 1st hour
after sunset

58

14.8

120

21.8

168

24.6

346

21.3

Within 2nd hour
after sunset

74

18.9

88

16.0

120

17.6

282

17.4

Within 3rd hour
after sunset

51

13.0

42

7.6

51

7.5

144

8.9

Within 4th hour
after sunset

24

6.1

37

6.7

30

4.4

91

5.6

Other times of
night

75

19.2

96

17.5

115

16.8

286

17.6

Within 3rd hour
before sunrise

7

1.8

14

2.5

19

2.8

40

2.5

l~ithin 2nd hour
before sunrise

6

1.5

14

2.5

13

1.9

33

2.0

Sunrise or within
1st hour before
sunrise

14

3.6

37

6.7

33

4.8

84

5.2

Within 1st hour
after sunrise

17

4.3

17

3.1

28

4.1

62

3.8

Within 2nd hour
after sunrise

9

2.3

8

1.5

15

2.2

32

2.0

Within 3rd hour
after sunrise

7

1.8

6

1.1

12

1.8

25

1.5

All other times
of day

29

7.4

43

7.8

33

4.8

105

6.5

Total

391

99.8

550

99.9

683

100.1

1624

100.1

�- 432 -

Table i r. Estimated
during 1969.

number of deer killed on Highway

HiBhwa~

82 and 13 by day

Day

No.

82
Percent

Noo

13
Percent

Total

Percent

Sunday

23

1595

10

1401

33

1501

Monday

15

1001

6

804

21

906

Tuesday

17

1105

8

1103

25

1l.4

Wednesday

19

1208

12

1699

31

1402

Thursday

17

1105

13

1803

30

13.7

Friday

34

23.0

11

1505

45

2005

Saturday

23

1505

11

1505

34

15,,5

Total

148

9999

71

100,,0

219

10000

HiBhwa~

On an hourly basis, 47 percent of the 129 deer killed at known times were hit
between 6:00 PM and 9:00 PM (Table 12). Most (13%) were hit between 7:00 PM
and 8:00 PM"

Time of Peak Roadside

Deer Activity

Information collected to determine the peak of roadside deer activity is
meaningless.
The bases which secured the electric eyes used to measure
deer activity in the median of Highway 82 were unstab1eo
The electric
eyes shifted away from the light sources and counted cars rather than deer.
In addition to this problem, deer were scarce and seldom crossed the portion
of highway instrumented with "deer detectors" as evidenced by the absence
of deer tracks in the median, the low vehicle-deer kill, and the low number
of deer counted in the area. A third problem was caused by human acttvity
in the area.
Results of "deer detector" counts appear in Table 13" Of interest is the
fact that vandals broke but two light sources from the end of February
through part of April (less than $20 damage which was easily repaired)9
Except for the unstable bases, equipment functioned properly in rain and
snow making it possible to collect a great amount of continuous data.

�- 433 -

Table 12g Estimated time of deer-auto accidents on Highway 82 and 13
study area.

Hour

NOg

Highway 82
Percent

0100
0200
0300
0400
0500
0600
0700
0800
0900
1000
1100
1200
1300
1400
1500
1600
1700
1800
1900
2000
2100
2200
2300
2400

4
2
2
5
4
1
0
0
0
0
4
0
0
0
1
2
6
13
13
8
10
8
14
5

3g9
200
2.0
4.9
3.9
100
000
000
0.0
0.0
3.9
0.0
0.0
0.0
1.0
2.0
5.8
12.•
8
12.8
7g8
908
708
13,,7
409

1
0
0
2
0
0
0
0
0
1
0
0
0
0
0
0
3
3
4
6
4
2
1
0

Total

102

100,,0

27

No.

Highway 13
Percent

Total

Percent

307
000
000
704
0 0
0.0
0,,0
0.•
0
0,,0
3,,7
000
0,,0
0 .•
0
000
0.0
0.0
11.•
1
l1.•
l
14g8
22.•
2
14.8
7.•
4
307
0,,0

5
2
2
7
4
1
0
0
0
1
4
0
0
0
1
2
9
16
17
14
14
10
15
5

.04
.02
,,02
.05
.•
03
.01
.00
.00
000
.•
01
003
,,00
.•
00
,,00
,,01
,,02
.07
.•
12
.13
.•
12
,,11
,,08
.11
004

99,,9

129

1.02

0

�- 434 -

Table 13. Time of activity recorded by detectors placed in the median of
Highway 82, February 28, - April 8, 1970. Although a few deer crossed
the median and were counted, most counts were triggered by vehicles and
human activity in and near the median. Data for Station 3 were summarized
for the period from Feb , 28, - March 27, 1970
0

Hour by
Military Time

Station 1

Station 2

0100
0200
0300
0400
0500
0600
0700
0800
0900
1000
1100
1200
1300
1400
1500
1600
1700
1800
1900
2000
2100
2200
2300
2400

393
330
335
443
215
124
26
64
39
37
26
127
189
70
68
192
170
255
181
403
632
1,083
994
409

42
87
49
60
329
106
36
1,550
1,244
1,195
737
702
580
340
129
240
288
25
6
29
3
6
66
97

Station 3

Total

28
48
5
13
0
0
4
39
21
40
168
5
12
28
36
25
0
4
6
5
0
1
1
1

463
465
389
516
544
230
66
1,653
1,304
1,272
931
834
781
438
233
457
458
284
193
437
635
1,090
1,061
507

Weather
Highway 13
Climatological data collected at the Highway 13 weather station appear in
Table 140 The average temperature from November 1969 through March 1970 was
26.5 0 F, about 3 OF warmer than the previous winter, at the Highway 13 weather
station. The mean low temperature was 15 30F, compared to 12 8°F the previous
winter. Almost 4 inches of precipitation was recorded for the four month period
in 1968-69, while during the same period in 1969-70, 3 23 inches of moisture fe110
0

0

0

Highway 82
Near Highway 82, the winter of 1969-1970 was relatively mild compared to the
winter of 1968-1969
The mean temperature at the weather station on Highway
82 was 24096°F in 1968-69 and 300540F in 1969-19700 The average low was
14040 F in 1968-69, while in 1969-70 it was 17090Fo
Precipitation for the
four month winter period was 3 9 inches in 1968-69 compared to 206 inches in
1969-70 (Table 15)0
0

0

�Table 14.

Climatological

data collected at the Highwav 13 weather station between April 1969 and March 1970.

Data Collected

Apr

May

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Jan

Feb

Mar

Maximum Temperature

(OF)

84

76

82

85

91

86

73

63

48

65

47

49

Minimum Temperature

(OF)

-4

22

25

21

41

29

6

-13

-10

-11

-4

-5

Ave. Maximum Temperature

(OF)

55.4

68.4

67.7

80.2

79.9

70.0

45.8

42.1

35.2

35.3

39.4

36.5

Ave. Minimum Temperature

(OF)

27.5

36.2

39.1

48.0

47.9

40.7

24.0

15.7

12.5

17.3

15.4

15.7

Total Precipitation

(IN.)

1. 34

.41

2.76

.73

1. 32

2.48

3.57

1.20

.61

1.02

.40

1.29

Ave. Wind Velocity

(M.P.H.)

1.35

4.81

3.68

3.97

3.19

4.91

3.28

3.57

2.59

4.26

3.26

3.19
I

+:-

w

U1

Table 15.

Climatological

data collected at the Highway 82 weather station between April 1969 and March 1970.

Data Collected

Apr

May

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Jan

Feb

Mar

Maximum Temperature

(OF)

82

82

88

92

96

93

73

61

53

52

55

59

Minimum Temperature

(OF)

16

23

23

21

40

30

21

-2

-7

-12

2

9

Ave. Maximum Temperature

(OF)

62.7

74.8

72.8

84.9

85.1

71.6

51. 3

46.5

39.8

37.7

46.6

44.9

Ave. Minimum Temperature

(OF)

29.3

37.5

41.6

49.8

48.9

40.7

29.3

18.7

16.2

12.6

19.6

22.7

Total Precipitation

(IN.)

.49

.50

2.71

.63

.54

1. 38

2.97

.30

1.23

.51

.60

1.02

Ave. 1~ind Velocity

(M.P.H.)

3.84

3.51

3.29

2.60

2.45

2.40

2.10

1.37

1.5

1.9

2.4

3.3

�- 436 -

Deer Condition
The weight of kidney fat from deer hit by vehicles on both study areas
during the period from October through March was highest, 64.1 grams,
in November and lowest, 13.0 grams, near the end of winter in March
(Table 16).
The percent of fat present in the bone marrow from femurs of deer killed
by vehicles on both study areas was lowest in April (Table 17). The
average fat content of the medial portion of the femurs from nine deer
killed in April was 26 27 percento The fat content rose to 44.39 percent in May and remained high until Decembero
0

Table 16. Average kidney fat weight (gm) by month of winter season from
animals killed on Highway 82 and 13
0

Fawn

Octo

Nov.

Deco

Jan.

Febo

Mar.

14.9

19.1

21 2

16.9

6.0

1.9

61.7

40 0
0

27.7

24.4

16.6

111.5

68 5

57.4

37.5

20.4

Yearling
Mature

52.3

0

0

Table 17. Percent fat content of bone marrow from the medial right or the
medial left portion of femurs from deer killed by vehicles on Highway 82
and 13 from 1968-1970.

Month

Fawn
Sample Percent
Size
Fat

February
March
April
May
June
July
August
September
October
November
December

6
13
3
4
0
0
0
1
10
6
3

25 55
12.92
2 .•
13
42 89
0

0

58 09
53.60
53 40
35.62
0

0

Yearling
Sample Percent
Size
Fat

0
5
1
0
2
0
1
0
4
2
0

42.86
63 74
0

75.96
67.69
71.86
81 27
0

Mature
Sample Percent
Size
Fat

1
8
5
5
0
1
0
0
4
1
0

77 .43
54 34
33.26
45.58
0

68.39

86.66
83.25

Total
Sample Percent
Size
Fat

7
26
9
9
2
1
1
1
18
9
3

32.96
31.42
26.27
44.39
75 96
68.39
67.69
58.09
65.00
62.91
35.62
0

�- 437 -

LITERATURE

CITED

Anderson, A. E., and D. E. Medin.
1965.
Two condition indices of the
Cache la Poudre mule deer herd and their application to management.
Colo. Game, Fish and Parks Dept., Outdoor Facts No. 23.
3 pp.
Gill,

J. D., and D. C. O'Meara.
1965.
Estimating time of death
white-tailed deer.
J. wildl. Mgmt.
29(3) :471-486.

Myers, G. T.
Progress

Prepared

by:

1969.
Deer-auto accident investigation.
Report, W-38-R-23.
p. 147-178.

/--J
,?-:/Z'-

f)

~;:-I~7

/-V-'
/ (..'4--t:",--&lt;-

Gary T. Myers
Wildlife Researcher

Colorado.

in

Job

��July,

lY7U

- 439 -

JOB PROGRESS

COLORADO

State of
No.

W-38-R-24

Deer-Elk

Plan No.

15

Job No.

Project
Work

REPORT

Job Title

Investigation

Period

April

Covered:

Personnel:

of One-Way

3

Deer

1, 1969 through March

Steve Steinert,
Dale F. Reed.

Investigations

Jack Corey,

Structures

31, 1970
Gary T. Myers

and

ABSTRACT

One-way structures that permit deer to move through deer-proof fences
from only one direction have been tested at the Little Hills Experiment
station paddocks.
The tests, based on mechanical counter readings and
track counts, indicate that 3/4 of all deer placed in the experimental
paddocks have used and re-used the one-way gate structures without the
aid of drift fences or the effect of funneling.
For two of the gates
tested, the frequency of use (preference) was significantly
different
2
(X = 4.08, P,0.05).
Two of the positions in the deer-proof fence did not show a significant
difference in use (X2 = 1.32, P~ 0.25).
In addition, one structure was
completely rejected because it failed the one-way criteria.
Two oneway fences have received only preliminary tests.
Information on frequency of use is necessary in order to modify or reject certain one-way
structures before they are subjected to extensive testing under varying
field conditions.

��- 441 -

INVESTIGATION

OF ONE-WAY

DEER STRUCTURES

Dale F. Reed

Many deer are killed on Colorado highways annually.
Some of these have been
trapped in the right-of-way where sections of the highway were fenced to prevent or reduce accidents involving both animals and people.
Much more
fencing (deer proof) is presently in the planning stages for critical deer
areas adjacent to the Interstate system.
Such structures should include
escape devices or one-way gates that permit deer to move through deer-proof
fences.
Deer that move around the ends of the fence, or from the side of
the road not fenced, will be able to pass through rather than be trapped and
more likely killed.
The object of this study is to develop and test one-way
structures most readily used by mule deer.

P. S. OBJECTIVE
Evaluate the effectiveness of several types of one-way gates and two fence
structures to permit one way movement of mule deer through existing deer
proof fences at Little Hills Experiment Station and elsewhere.

SEGMENT
1.
2.
3.
4.

OBJECTIVES

Design and construct several different t)~es of untested one-way deer
gates and two different one-way deer fences.
Install working examples of each structure in paddocks at Little Hills.
Test the effectiveness of each structure under controlled conditions.
Test those structures deemed effective under controlled conditions for
effectiveness under field conditions.

METHODS

AND MATERIALS

Design and Construction
One-way

Gates

The basic design for a one-way gate resulted from watching deer hit fences
in deer enclosures.
It was reasoned that the hole or opening on one side
must be larger than the other, thus allowing easy passage from one side and
difficult or impossible passage from the other.
The one-way characteristic
is, therefore, based on a funneling or cone principle.
Structures which lend
themselves readily to this principle are bailer tines.
Figure 1 shows 19
such tines arranged to provide an adjustable opening on one end. Table 1
lists the differences between all four one-way gates, three of which have
tines as a major component of construction.
The remaining type utilizes 10'
panels of woven wire in order to obtain a relatively long funnel effect.

�- 442 Fig. 1.

.
deer gate (with either Swinging
One-way
. g action) •
or Rigid sp r.i.n

.....,.: ..
""-,

~.. -:
,

---}'--

r
I

I

I

i
!

'fCf

II

;

'''--creosote
deer
fence
post

i

i
. 2" angleiron 1/8"
thick

I

!

Y-. I
I

ii
1

j
I

---'7..---

,

approx.

·1 .bole

].2"

ground

�- 443 -

Table 1. Outcome of testing and characteristics
tested at the Little Hills paddocks.

of four one-way

gates

Gate

Outcome of
Pre limina ry &amp;
Preference Tests

Operational
Principle

Component of
Construction

Width of
Large End

Width of
Small End

I

To be field tested

Rigid spring

Tines

26"

4"

II

Rejected during
preference tests

Swinging spring
(opens freely)

Tines

28"

0"

Rejected during
preliminary tests

Long funnel

10' x 6 '
Woven wire
panels

36"

8"

Tines

28"

4"

III

IIIb

To be field tested

One-way

Fences

Rigid spring
at ground level

Two possible one-way fences, similar to the overhanging and slanting fences
of Longhurst and Jones (1962), were constructed.
Each was made with variable vertical sections and with either a variable overhanging or slanting
section.
Table 2 lists the characteristics
and adjustable features of both
fences.
The variable sections were accomplished by use of telescoping
square steel tubing (Unistrut Western, Inc., 601 S. Jason St., Denver,
Colorado).

Installation

at Little Hills

Openings were constructed for the one-way gates at three designated places
(A, B, and C) in an existing deer-proof fence separating two 2 acre paddocks.
Construction was such that each of the one-way gates would fit any opening
and could be mounted in a direction oriented toward either paddock.
Larger openings, approximately 14' wide, were made for one-way fences in
a deer-proof fence also separating two paddocks.
The two fence sections
were installed so that they could be mounted in a direction oriented toward
either paddock.

Testing
One-way

Under Controlled

Conditions

Gates

The one-way gates were tested with small groups or individual
in the Little Hills paddocks throughout the year.

deer placed

�- 444 -

Table 2.

Characteristics

and adjustable

features

of one-way

fences I and II.

Max. -Min.
Vertical (In. )

Max.-Min.
Depth (In. )

Variable
Angle Limits
(from vertical)

Operational
Principle

I

66"-42"

64"-34"

450-900

Overhanging

II

72"-48"

90"-48"

1150-1300

Slanting

One-Way

Fence

Preliminary Tests--Preliminary
testing in the paddocks involved placing
small groups of deer in one paddock and food in another with a one-way gate
located in the deer-proof fence separating the paddocks.
The direction of
each gate was changed frequently to determine if negative passages (passages
against the funneling effect) could occur.
This procedure was repeated with
each of three gates until it had been established that they had positive
passages only (Fig. 2), and hence were one-way.
If a gate had negative
passages, it was modified until none occurred or it was rejected.
Estimates
of deer use were obtained by track counts on both sides of the gates to determine the direction and the number of deer passing through the gates over a
predetermined period of time which varied from one to several days.
Several
mechanical gate-use counters were used to supplement track count information.
Tests of Preference--Tests
of preference (frequencies of use) were made on
two gates at a time in several positions between two paddocks with individual
deer.
The location of each gate was changed in order to eliminate the preference of gates because of position.
With two gates (I and II or I and IIIb)
being tested in three possible positions, two of three possible sequences
were used (if A, B, and C are the positions; and I, II, and IIIb are the gates):
BI CII, BII CI, and AI Clllb, Alllb CI. Thus, the position of the gate was
reduced or eliminated as a possible bias and deer preference was tested.
Three gates were not tested Simultaneously during this segment because of
the greater number of days required.
Before the deer were tested individually, they were oriented in small groups
(Table 3) to each of the two gates for a period of 4 days.
Each gate was
rotated systematically
from position to position so that the deer could
experience one gate one day, another the next, etc. It was felt that to
have a preference, an animal must have experienced both possibilities.
Deer
were oriented in small groups rather than individually to reduce the number
of days necessary.
Once the group was oriented, all except one deer were returned to a holding
pen to await individual testing.
The deer to be tested was left in a paddock
not containing food. Both gates were then placed in position between the
paddock not containing food and the paddock with hay and pellets, thus
beginning the sequence.
Each deer was tested for 4 days, allOWing the sequence
to be repeated.
This procedure was repeated until each deer of the group was
tested.

�- 445 -

Fig. 2.
tests.

Positive

passage

of deer through a one-way

gate during preliminary

�- 446 -

One-way Fences
The number of deer crossing the one-way fences was to be counted each day
in a manner similar to that of the one-way gates. Preliminary tests had
just begun by the end of this segment, consequently, the procedures have not
yet been fully developed.
Testing Under Field Conditions
Three duplicates of the one-way gate deemed most effective under controlled
conditions were installed in a deer-proof fence adjacent to Highway 82. In
addition, because of the immediate need for such structures, three installations were recently made in the Vail underpass drift fences.
Effectivness of each structure under field conditions will be determined by
recording the number of tracks in and out of the gates and by gate-use
counters during the next segment.

RESULTS AND DISCUSSION
Controlled Conditions
One-way Gates
The one-way gates were tested with 45 deer placed in the Little Hills
paddocks throughout the year. These deer of various sex and age classes
were used in both preliminary testing and testing· for gate preference.
Preliminary testing of 3 groups of deer (Table 3) on gates I, II, and III,
provided information on whether the gates were one-way or not. The unlabeled
group of 11 fawns (Table 3) was used in repetitive te~ts on gate I which
was modified until essentially no negative passages occurred. Gate II was
the same as gate I except that it was made larger. Deer groups A and B
were used for tests on gates I, II, and III. Data on the number and percent
of passages (Table 4) indicate that both gates I and II are almost equally
effective, 94 and 100 percent respectively, in terms of the percent of
positive passages of the total passages made. Conversely, the percent of
positive passages (53%) was not considered great enough to warrant further
testing on gate III. Observation of deer going through the gate indicated
that they were follOWing a trail to the small unobstructed opening at the
negative end. The principle of a long funnel by itself was considered to
have failed and no modifications were made. Gate IIIb was modeled after
gates I and II, and was not involved in the preliminary testing.
Testing for gate preference involved 24 deer in orientation groups C through
L (Table 5). Gates I and II were evaluated Simultaneously with each of 19
individual deer. Thirteen of the 19 deer used the one-way gates. The total
frequencies of use (preference) on these two gates were significantly different (X2 = 4.08, P&lt;0.05).

�- 447 -

Table 3. Number and classification of deer in groups tested, gates tested,
and type of tests completed.
Group

Yearlings

Fawns

Does

Bucks

Total

Gates Tested

11
6
4

I
I,It,IlI
I,Il,III
I,ll
I,ll
I,II
I,ll

11

6
4

A
B
C

I

D

2

1
2

E

2

1
1

5
I

1

F
G
H

2
2

I

1

J
K
L

I
2

Total

23

I

1

1
5
4
1
2
2
1

8

45

2
2
I

10

4

r.rr
I,II
I,II
I,IIIb
I,IIIb
I,IIIb

Tests

Preliminary
Preliminary
Preliminary
Preference
Preference
Preference
Preference
Preference
Preference
Preference
Preference
Preference
Preference

Table 4. Number of deer passages for each one-way gate tested and percent
of positive passages.

Passages during
Prelimina~ tests
Pos.
No.
No.
Pos.
Neg.
%

Passages during
Preference tests
No.
No.
Pos.
Neg.

Total Passages
No.
No.
Pos.
Neg.

I

52

-2

94

51

-2

103

-4

96

II

29

0

100

20

-1

49

-1

98

III

40

-35

53

0

40

-35

53

7

0

100

IlIb

7

0

%

All Tests
Positive

�Table 5.

Frequencies of use (preference) of one-way gates and of one-way gate positions.

pos.

DOE
No. Group

.50 .50

1

--

a

a

--

a

a

I

GATE
II

IlIb

A

B

C

.50

.50

---

--

2

D

a

a

---

3

D

a

a

---

FAWN
No. Group

1

C

pos.

BUCK
No. Group I

GATE
II

D

a

a

2

E

.50

3

G

.50

I

GATE
II

Illb A

B

C

.75

.25

---

--

.75 .25

1

2

D

a

a

---

--

a

a

3

D

a

a

---

--

a

a

C

ros.
Illb A

B

C

---

-- a

a

.50

---

--

LOa

.50

---

--

.50 .50
I

4

F

.67

.33

---

--

La

a

4

G

.67

.33

---

--

La

a

4

G

.50

.50

---

--

.50 .50

.p.p00

5

G

.50

.50

---

--

.50 .50

5

H

.75

0

---

--

.25 .50

6

G

.67

.33

---

--

.33 .67

6

H

a

0

---

--

a

7

H

1.0

a

---

--

.50 .50

7

J

1.0

---

a

.50

-- .50

8

H

.50

.50

---

--

.50 .50

8

L

.75

---

.25

.75

-- .25

9

I

.75

.25

---

--

.25 .75

10

J

.33

--

.67

.67

--

.33

11

K

.75

--

.25

.25

--

.75

12

K

.75

--

.25

.75

--

.25

0

�- 449 -

Positions Band C used in the deer-proof
difference in use (X2 = 1.34, P &lt;0.25).

fence did not show a significant

Gates I and IIIb were likewise evaluated simultaneously, but with only 5
individual deer. Frequencies of use from this small sample were not significantly different (X2 = 2.35, P&gt;O.lO).
It is evident that from the X2 value of gates I and II that some factor is
influencing a greater number of deer responses to gate I. Since a possible
bias of position appears to be insignificant, it may be concluded that this
factor is some physical characteristic of one of the gates.
A suspected
physical characteristic of gate II that may be involved, is the noise that
is produced when it closes (metal against metal).
Gate I tines do no t open
or close freely - hence no noise is produced.
One-way

Fences

Since preliminary tests had just begun by the end of this segment, no analysis
can be made at this time. So far deer have been reluctant to use the one-way
fences in their lowest positions.
Only two positive crossings and one negative crossing have occurred during a total of 11 days when deer were in the
paddocks with fences in the positive direction.
Field Conditions
Deer use of one-way gates installed in the field had not occurred by the end
of this segment, consequently, no analysis or results can be discussed.
The
preliminary evaluation of the one-way fences must be completed before such
structures will be considered for field testing.

LITERATURE

CITED

Longhurst, William M., Milton B. Jones, Ralph R. Parks, Loren W. Neubauer,
Maynard W. Cummings.
1962. Fences for controlling deer damage.
Circular 514. pp. 7-13.

Prepared

by

~?i?f.~
Re~

Assistant

/.

#4

Wildlife

Researcher

and

��July, 1970

- 451 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-24

Deer-Elk

Work Plan No.

15

Job No.

Job Title

Evaluation

of Devices

Investigations
4

to Prevent Deer-Auto

Accidents

Period Covered:

April

1, 1969 through March 31, 1970

Personnel:

Dale F. Reed, Dennis L. Money, Kenneth C. Dillinger,
William B. Zimmerman, Claudia A. Doose, Gary T. Myers

ABSTRACT

The height of an existing fence adjacent to 3.8 miles of Highway 82
was not extended to eight feet because only seven deer, less than two
per mile, were killed by cars in the area to be fenced during 1968,
the extent of deer movement from the west across the highway and into
the fence was unknown, and the $25,000 required for fence extension
was unabailable.
Data relating to the effect of brush removal on the number of deer
killed by vehicles will not be analyzed until sample sizes are sufficient
to determine if differences in pre- and post-treatment data exist.

�- 452 -

EVALUATION

OF DEVICES

TO PREVENT DEER-AUTO

ACCIDENTS

Gary T. Myers

P. S. OBJECTIVE
Determine if a fence built adjacent to a portion of Highway 82 and brush
removal in strategic locations adjacent to Highway 13 south of the White
River bridge in Garfield County affects the number of deer killed by
vehicles in the areas involved.

SEGMENT
1.

2.

Determine if a 3.8 mile deer-proof fence adjacent to one side of Highway 82 near Cattle Creek effectively reduces the number of deer killed
there by motorists.
Measure the effect of roadside brush removal at selected deer crossings
on Highway 13 or Highway 82 on the number of deer struck by vehicles in
these areas.

METHODS
1.
2.

OBJECTIVES

AND MATERIALS

None
The Highway Department removed brush from all but four of the 40 miles
of right-of-way adjacent to the Highway 13 study area. Brush removal
was completed by April 1, 1970. Post-treatment kill and count data
were gathered weekly since completion of brush removal.

RESULTS AND DISCUSSION
The height of an existing fence adjacent to 3.8 miles of Highway 82 was not
extended to eight feet because only seven deer, less than two per mile, were
killed by cars in the area to be fenced during 1968, the extent of deer
movement from the west across the highway and into the fence was unknown,
and the $25,000 required for fence extension was unavailable.
Data relating to the effect of brush removal on the number of deer killed
by vehicles will not be analyzed until sample sizes are sufficent to determine if differences in pre- and post-treatment data exist.

Prepared

by
Wildlife

Researcher

�- 453 JOB

State

of

Project

PROGRESS

REPORT

COLORADO
--------~~~~~-------No.

Job

15

Investigations
5

No.

Work

Plan

Job

Ti t Le _-=E:.:f:.:f:.,:e:..:c:..:t=-.,:o:..:f=-":S:::,,pl:...e::.e=d-.:::.o.:.;n;,.....:.:N:..=u:::m.::b:..,
_

Period

No.

Deer-Elk

W-38-R-24

Covered:

Personnel:

April

1, 1969

Gary

T. Myers

through

March

31,

1970

P. S. OBJECTIVE

Determine
if traffic
moving
at a high average
speed
Jameron
Curve than traffic
moving
at a slow average

SEGMENT

1.
2.
3.

4.

kills more
speed.

deer

on

OBJECTIVES

Manipulate
the speed of traffic.
Measure
the average
speed of traffic.
Record
the number of deer killed.
Compare
the number of deer killed
by traffic
moving
at a lower
average
speed to the number killed by traffic
moving
at a faster
average
speed.

RESULTS

The Colorado
State Patrol
manipulation
of the speed
study can not be made.

AND

DISCUSSION

and .Division
of Highways
would not allow
of traffic
on Colorado
82. Therefore,
this

Prepared

by

Am r:P 111:;;&lt;-'1.'1
Gari~yers:
Wildlife
Researcher

��July,

1970

- 455 -

JOB PROGRESS

State of

REPORT

COLORADO
----------------------------

Project No.

W-38-R-24

Deer-Elk

Work Plan No.

15

Job No.

Job Title

Deer Underpass

Period Covered:

April

Personnel:

Gary T. Myers,

Investigations
6

Evaluation

1, 1969 through March

31, 1970

Bill Zimmerman,

and Dale F. Reed

p. S. OBJECTIVE

Determine if deer migrating from winter range on one side of Interstate
70 to summer range on the opposite side utilize and underpass constructed
in the area.

SEGMENT OBJECTIVE

Measure

deer use of the underpass.

METHODS

AND MATERIALS

Construction of the underpass was completed
of Highways toward the end of this segment.

RESULTS

by the Colorado

Department

AND DISCUSSION

Plans were prepared for the installation of electric-optical
detection
and counting equipment (Streeter Amet) at both ends of the underpass.
Installation and evaluation is anticipated during the first part of
the next segment.

c'~

Prepared

by

- I /J

--7[0:12 &lt;Lt: t, /y;j.)'
bale F. Reed
Assistant Wildlife

:1

~
Researcher

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                  <text>October,

- 1 -

JOB PROGRESS

State

Work

REPORT

COLORADO

of
No.

W-88-R-15

Migratory

Plan No.

1

Job No.

Project

Job Title
Period

1970

Covered:

Personnel:

Waterfowl
April

Production

15, 1969 to June

Bird

Investigations

1

Survey

30, 1969

C. Bryant and staff, Monte Vista National Wildlife Refuge;
R. Pearson, Brown's Park National Wildlife Refuge; C.
Hayes and J. Randall, Bureau of Sport Fisheries and Wildlife, W. Ladd and G. Will, Colorado State University;
G. Crawford, H. Funk, C. Hector, J. Hobbs, R. Hopper,
W. Russell, W. Rutherford, R. Reynolds, H. Schultz, J.
Wheeler and M. Szymczak, Colorado Division of Game, Fish
and Parks.

ABSTRACT

Weather conditions for waterfowl nesting and brood survival were generally
good in Colorado in 1969.
The total estimate of 69,025 duck breeding
pairs in major Colorado nesting areas was about 3,000 fewer than in 1968.
A major decrease in pairs occurred in the South Platte Valley while total
pairs in other areas either increased slightly or remained stable.
The
mallard remained the major breeding species in Colorado with an estimated
28,744 breeding pairs recorded.
Goose production in northwest Colorado
declined slightly from record 1968 levels with 425 goslings produced in
1969 compared with 487 the previous year.
The resident northcentral
Colorado Canada goose breeding population produced a total of 1,125
goslings, increasing the total population size to 4,208 birds.

��- 3 -

WATERFOWL

PRODUCTION

Michael

SURVEYS

R. Szymczak

P. S. OBJECTIVE
To formulate waterfowl

harvest

regulations.

SEGMENT OBJECTIVES
To determine, through statistically reliable sampling techniques, the number
of duck and goose breeding pairs, by species, in each major Colorado waterfowl breeding ground.

METHODS

AND MATERIALS

Present duck breeding-pair and production surveys
inventory of only major production areas.

consist of a breeding

pair

The 1969 duck surveys were conducted during the period May 12 to May 31.
Ground counts were made in the Yampa Valley and Brown's Park. Aerial surveys were conducted in North Park and the South Platte and Cache la Poudre
Valleys.
Intensive aerial counts combined with air-ground comparison
studies were conducted in the San Luis Valley.
All survey methods and sample areas remained the same as in previous years.
Flying was accomplished with a Cessna 185 aircraft.
Areas sampled by section or block were flown with one observer, while two observers were used in
sampling by transect.
Ground surveys of Canada goose production were conducted within the period
of April 15 to June 30. Production estimates in the Yampa Valley-Brown's
Park area were made from as complete a count as possible of breeding pairs,
goslings and eggs.
Gosling production and population size in the DenverBoulder-Fort Collins area were determined by conducting a one day count of
all geese on moulting and brood rearing areas and classifying birds as to
whether they were adults or goslings.
On the basis of these studies, a report is submitted
Fisheries and Wildlife, which constitutes Colorado's
continental cooperative breeding ground survey.

to the Bureau of Sport
part in the annual

RESULTS AND DISCUSSION
Weather conditions for waterfowl nesting were generally good in most areas
of the state.
The winter snow pack was considered normal in most areas with

�-4 -

heavy snows on some west slope drainages.
High water created some goose
nest flooding problems on the Yampa River. Water conditions were considered better than average for duck nesting in North Park, Brown's Park,
and the Yampa and Cache la Poudre Valleys.
Water in the South Platte and
San Luis Valleys was not as favorable as in 1968.
Figures in Table 1 reveal that the total duck breeding population was down
about five percent from the 1968 level. Only North Park and the Cache la
Poudre Valley showed increases from 1968 levels.
The most significant decline occurred in the South Platte Valley where the number of pairs recorded
were less than one-half of the previous year's total. Decreased water areas
in the South Platte Valley was probably responsible for decline in duck
numbers.

Table 1. Summary of Colorado duck breeding ground population
1969, with 1968 and the l5-year average for comparison.
Total Estimated

Breeding Pairs
From l5-yr.
Ave. 1954-1968

estimates,

Percent Change
From
From l5-yr. 1/
1968
Average

Area

1969

1968

San Luis Valley

27,425

27,611

27,549

North Park 1/.

22,111

19,777

7,093

+ 11.8

Cache la Poudre Valley

9,057

7,403

2,525

+ 22.3

+258.7

South Platte Valley

6,769

14,000

5,721

- 51. 7

+ 18.3

Yampa Valley

2,146

2,985

2,920

- 28.1

- 26.5

Brown's Park

1,517

938

206

+ 61. 7

+636.4

Totals

69,025

72,714

46,014

5.1

+ 12.1

Jj

0.7

0.4

1/ Does not include North Park because of change of method of projection
beginning in 1968.
1/ San Luis Valley averages are based on results of 1964-1968 only. The much
less intensive coverage of previous years is not included in the calculations.
1/ Aerial counts corrected by species from visibility ratio obtained in the
San Luis Valley during 1968 and 1969 only.

Table 2 lists composition of the total estimated breeding duck populations
for 1969, 1968, and the long-term averages by numbers and percent.
A significant change in species composition was recorded in 1969. The percent of
mallards in the breeding population declined and the percent of teals, particularly green-winged teal, increased.
A change in observers may have been

�-

5 -

partially responsible for this change in composition.
Green-winged teal
showed significant increases in North Park whereas the mallard breeding
population in South Platte declined by about 5,000 breeding pairs.

Table 2. Species composition of the Colorado breeding duck population,
1969, with 1968 and the l5-year average for comparison. !/
Percent Species

Species

Number of Breeding Pairs
1954-1968 2/
1969
1968
Average-

1969

1968

Composition
1954-1968
Average

Mallard

28,744

36,644

28,669

41.6

50.4

60.9

Blue-winged and
Cinnamon Teal

9,302

6,463

4,237

13.5

8.9

9.0

Pintail

4,355

7,970

3,306

6.3

11.0

7.0

Gadwall

5,942

8,425

4,l32

8.6

11.6

8.8

American Widgeon

1,870

343

547

2.7

0.5

1.2

Shoveler

2,750

3,645

1,785

4.0

5.0

3.8

Teal 12,924

5,411

1,591

18.7

7.4

3.4

Redhead

2,071

2,063

1,676

3.0

2.8

3.6

Other Divers

1,067

1,750

1,139

1.6

2.4

2.3

Totals

69,025

72,714

47,082

100.0

100.0

100.0

Green-winged

1/ Change in observers may have had an effect on species composition.
included here, are for the years 1964-1968

2/ San Luis Valley averages,

only.

Table 3 categorizes all geese observed on the breeding inventory in northwestern Colorado by age and location.
This information is compared with past
years in Tables 4 and 5. Total flock size and production show a decrease
from the record high numbers of 1968. Fewer adult birds were present on all
sections except the Little Snake River. More nesting pairs were recorded in
1969, but brood sizes were relatively smaller resulting in a decline in total
estimated production.
Undoubtedly, high water on the Yampa River had a
detrimental effect on production.
Figures presented for 1969 for the Green
River may not be comparable to those of 1968 as a less intensive survey was
conducted this year.

�- 6 -

Table 3. Number of Canada geese observed
County, Colorado, 1969.

and estimated

production,

Moffat

Nesting
Pairs

Non-nesting
Birds

Total
Adults

Estimated No.
Goslings 1/

Total
Birds

Yampa River
Craig to Juniper Spgs.
Juniper to Cross Mtn.
Lily Park

16 Jj
9 Jj

113
27
66

145
55
84

80 1/
48
44 !!./

225
103
128

YAMPA TOTALS

39

206

284

172

456

Green River
(Brown's Park)

20

171

231

144

375

Little Snake River
(Lower bridge to
State line)

27

228

282

109

391

GRAND TOTALS

96

605

797

425

1,222

Area

1/ Calculated

14

using average brood size and number of successful

nests.

1/ Includes nesting pairs associated with nests from which eggs were removed

for transplant purposes.
3/ Includes 29 goslings hatched
4/ Includes 18 goslings hatched

Table 4.

from eggs taken from nests before final survey.
from eggs taken from nests before final survey.

Total Canada geese observed,

Moffat

No. Geese Counted

County,

Colorado,

1969.

Percent

Change
From 1956-68
Ave.

1956-68

Area

1969

1968

Ave.

From 1968

Yampa River

456

624

357

-26.9

+ 27-.7

Green River

375

363

91

+ 3.3

+312.1

Little Snake River Jj 391

333

214

+17.4

+82.7

1,320

662

- 7.4

+ 84.6

Totals

1,222

1/ Not included in survey until 1962.

�- 7 -

Table 5.
1969.

Estimated

number of Canada goose goslings,

Moffat

County,

Colorado,

Area

No. of Goslings·
1956-68
1969
1968
Ave

Yampa River

172

239

132

-28.0

+ 30.3

Green River

144

139

32

+ 3.6

+350.0

1./ 109

109

67

0

+ 62.7

425

487

231

-12.7

+ 84.0

Little Snake River

Totals
1/ Not included

Percent
From 1968

Change
From 1956-68
Ave.

in survey until 1962.

Results of the June 19, 1970 census of Canada geese in the Fort Collins-BoulderDenver or foothills area are presented in Table 6. Comparative figures for
these specific areas for previous year are not available.
However, breakdowns by county or area basis are presented in Tables 7 through 9. The new
method of data collection, a one day count of moulting adults and young, does
not produce reliable information concerning nests.

Table 6.
Colorado.

Canada goose production

Production

Area

wellington

Water Area
Terry Lake
Douglas Lake
N. Poudre III
Divide No. 8
Elder Res.
Annex No. 8
Cobb Lake
Watson Lake

census of the Foothills

No.
Broods

2
4
5
9
0
3

Total
Fort Collins

Herring Lake
College Lake 1./
Dean Acres
Claymore Lake

3
6
4

Area, northcentral,

Total No.
Total No.
Goslings
Adults &amp; Yearlings

Total No.
of Birds

98
6
18
27
26
0
14
68

286
4
12
190
43
13
38
102

384
10
30
217
69
13
52
170

257

688

945

13
64
28
14

30
175
28
38

43
239
56
52

------------------------------------------------------------------------------------

�- 8 -

Table 6. Canada goose production census of the Foothills Area, northcentral,
Colorado (continued).

Production Area

Water Area

No.
Broods

Fort Collins
(Continued)

Sterling Gravel
Pits
Lindenmeier Lake
N. &amp; S. Grey Res.
Anderson's Pond
Parkwood Lake
Timnath Res.
Romily Gravel Pit
Fossil Creek Res.
Schuelke Res.

7
1
3
9
1
8
I
3
I

Total
Loveland

Flatiron Res.
Boedecker Res.
McNeil Res.

1
6

Total
Denver

Ish Lake
Crystal Lake
Terry Lake
Faivre Ponds
Hayden Lake
Rest Home Pond
Valmont Res.
Boulder Valley
Farm
King Pond

Total No.
of Birds

27
4
15

59
16
17

86
20
32

32

19

51

92

123

6

527
31
729
12
4

9

18
13

256

493

749

2

2

4

25

48

73

10

53

63

37

103

140

1

4

1

3

2
2

6
5

14

32

14
4

27

46
41

2

6

18
77

20
384

38
461

10

10

20

5

2

7

149
14

481
38

630

36

53

89

175

250

7

9

425
16

25

62

87

18

138
34

5

120
34
425
107

20

142

o

27

Total
Boulder

Total No.
Total No.
Goslings Adults &amp; Yearlings

1

1

Standley Lake
3
Sloans Lake
9
Denver City Park 41
Rocky Mtn. Arsenal 2
Federal Center
Pond 1.
Cline &amp; Kountee
Res.
6
Kendrick Res.
o
Bowles Lake 1./
Marston Res.
1
Kings Pond
6
Upper Tule Lake
o

o
98

52

523

112
162
27

�- 9 -

Table 6. Canada goose production census of the Foothills Area, northcentral,
Colorado (continued).
No.
Broods

Production Area

Water Area

Denver
(Continued)

Lower Tu1e Lake
S. Colo. Blvd.
and Quincey
Mohn Estate and
Blackmer L.
Reservoir No. 3

Total No.
Total No.
Goslings Adults &amp; Yearlings

o

o

18

18

5

13

16

29

3

9
6

10
7

19
13

426

1,318

1,744

1,125

3,083

4,208

2

Total
GRAND TOTAL

Total No.
of Birds

!/ Includes birds removed from area prior to census.
Table 7.

Larimer County Canada goose flock production information

Year

No. Nests
Established

No. Nests
Hatch

No. Birds
Raised

No. Birds
Planted

1957

0

0

0

31

31

1958

0

0

0

23

54

1959

0

0

5

48

60

1960

4

4

14

68

120

1961

7

6

20

95

210

1962

23

21

79

101

400

1963

43

31

100

0

500

1964

68

59

154

0

600

1965

79

60

178

0

650

1966

106

75

213
No

Approximate
Size of Flock

0
750+
Birds Moved Out
177
1,078

1967

173

124

374

1968

267

185

564

218

1,555

550

351

1,834

1969

�- 10 -

Table 8.

Boulder

Year

No. Nests
Established

County Canada goose flock production
No. Nests
Hatch

information.

No. Birds
No. Birds
Raised on Area Planted

Approximate
Size of Flock

1963

86

1964

97

175

1965

6

6

11

131

300

1966

22

15

52

116

400+

1967

36

28

81

177

500+

1968

57

44

132

0

149

0

1969

Table 9.

Year

1969

Denver area Canada goose flock production

No. Birds Raised

information.

No. Birds Moved Out

426

630

300

Approximate
Size of Flock

1,744

The data indicates that the northcentral Colorado resident breeding flock is
continuing to increase in spite of the continued removal of goslings from the
area for transplants in to other parts of the State.
The total flock size has
now increased to approximately 4,200 birds.

Fall Flight Prediction
Expected fall flights of ducks from Colorado's 1969 production will be above
average but slightly less than in 1968. Water conditions appear very favorable
for brood survival.
Production of geese in northwestern Colorado was below the 1968 level indicating
a slight decline in the fall flight.
The resident Canada goose breeding
grow and expand and should provide

flock in northcentral Colorado
excellent hunting locally.

Prepared
Michael R. Szymczak
Asst. Wildlife Researcher

continues

to

�October,

- 11 -

1970

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-88-R-15

Migratory

Work Plan No.

1

Job No.

Bird Investigations
2

Job Title

Trapping

and Banding Ducks and Geese

Period Covered:

July 1, 1969 to March 31, 1970

Personnel:

Charles Hayes, Jack Randall and Dale Horne, Bureau of
Sport Fisheries and Wildlife; Ralph Baker, Clait Braun,
Gary Brown, Bill Carpenter, David Cramer, Gurney Crawford,
Mark Frasier, Howard Funk, Larry Green, Bob Kitzmiller,
Roger Lowery, Kris Moser, Gene Nugent, Ray Schmidt,
Mike Szymczak, Ken Wagner, Mike Watkins, Clayton Wetherill,
Bert Widhalm, Gary Will, Henry Wilson, Mike Zgainer, and
.Richard Hopper, Colorado Game, Fish and Parks Division.

ABSTRACT

Nearly 8,750 ducks of 10 species were banded and released during 1969-70
(Segment 15). Mallards made up about 6,900 of this total. Green-winged
teals and pintails were thenext most abundant species banded, comprising
824 and 720, respectively.
Post-season banding (winter) contributed about
4,000 birds to the banded sample.
Pre-season banding in the San Luis
Valley and South Park produced 3,602 and 1,306 ducks, respectively.
Only
385 ducks were banded during the molting period in North Park.
About 1,300 Canada geese were banded in Colorado in Segment 15. Nearly
800 of these resulted from winter trapping operations in the Arkansas
Valley and the Cache la Poudre Valley.
The remaining bands were placed
on goslings transplanted in the South Platte Valley, San Luis Valley,
North Park, and on the Colorado River near Grand Junction.

RECOMMENDATIONS

1. Trapping and banding of molting ducks in North Park be discontinued
in favor of pre-season banding of free-flying ducks.

��- 13 -

TRAPPING

AND BANDING

DUCKS AND GEESE

Richard M. Hopper

This report summarizes all waterfowl banding activities of Project W-88-R-lS
for the Segment year April 1, 1969 to March 31, 1970. The actual analysis
of band recovery data occurs as a separate job (Work Plan 1, Job 3), and
work of this nature is not included here.
This report simply presents a
tabulation of numbers of ducks and geese banded by species and location
during the Segment.

P. S. OBJECTIVE
To formulate waterfowl

harvest

regulations.

SEGMENT

OBJECTIVES

1.

To trap and band ducks and geese for the purpose
life history, and annual mortality data.

2.

To report species and numbers of ducks winter banded in the South Platte
Valley, Arkansas Valley, and Bonny Reservoir as part of ~vork Plan 3,
Job 6.

METHODS

of accumulating

migration,

AND MATERIALS

Methods employed and materials used during Segment IS remained the same as
in previous Segments (Hopper 1968). Banding locations and times were the
same as in Segment 14 (Hopper
1969), except that Bonny Reservoir and the
Cache la Poudre Valley were added to the list of areas for inclusion in
the winter goose banding program.

RESULTS AND DISCUSSION
Ducks
The number and species of ducks banded during Segment IS is listed in
Table 1 by location of banding.
Nearly 8,750 ducks, representing 10 species,
were banded and released.
As usual, the mallard was the most abundant
species, comprising 6,906, or about 80 percent, of the total number of ducks
banded.
Pre-season banding in the San Luis Valley and winter banding in
eastern Colorado contributed most to the mallard total. Green-winged teals
and pintails were the next most numerous species in the banded sample, with
824 and 720, respectively.

�Table 1.

Number of ducks banded by species and location, 1969-70.

Species

North
Park

Cache la Poudre
Valley

South Platte
Valley

Bonny
Reservoir

Arkansas
Valley

San Luis
Valley

South
Park

Total

Mallard

148

591

1,236

645

915

2,545

826

6,906

Gadwall

11

---

1

---

---

15

---

27

American Widgeon

33

8

3

---

---

2

6

52

Green-winged Teal

131

27

---

---

---

529

137

824

Blue-winged and
Cinnamon Teal

6

---

---

---

---

133

23

162

Shoveler

8

---

---

---

---

4

---

12

t-'

~

Pintail

47

17

2

---

---

341

313

720

Redhead

---

---

---

---

---

32

1

33

1

---

---

---

---

1

---

2

385

643

1,242

645

915

3,602

1,306

8,738

Lesser Scaup
Total

�- 15 -

Poor trapping and banding success was attained on molting areas in North
Park during the summer, largely because we missed the peak of molting.
Only 385 ducks were banded, with mallards and green-winged teals accounting
for most of the total. Duck banding in North Park during future years, if
conducted at all, will be limited to the pre-season period after most birds
have ceased to molt their flight feathers and have acquired the ability to
fly. Few immature birds have been banded on molting areas in the past.
Pre-season banding should remedy this situation by placing more immatures
in the banded sample.
Pre-season banding was again accomplished in the San Luis Valley as part of
the continuing cooperative investigation in that area of Colorado.
This
operation was conducted during the period August 15 - September 20 and
resulted in the banding of 3,602 ducks.
The mallard quota (2,000) was
reached in the northern one-half of the Valley.
Green-winged teals and pintails made up most of the remaining number.
Pre-season banding at two locations in South Park (Antero Reservoir and the
Hartsel Ranch) yielded a banded sample of 1,306 ducks, of which mallards
comprised 826. Again, pintails and green-winged teals were the next most
abundant species, contributing 313 and 137, respectively.
The 1969-70 post-season (winter) mallard banding program was less successful
than in previous years because of mild weather that distributed the birds
widely and prevented them from concentrating in desirable trapping areas.
Only 3,849 mallards were banded, or about 1,350 shy of the 5,200-bird
quota.
The San Luis Valley was added to the list of winter banding locations
in Segment 15. The Job Progress Report for Work Plan 3, Job 6 covers this
particular work in more detail.

Geese
Nearly 1,300 Canada geese were banded in Colorado during the 1969-70
Segment (Table 2). Winter banding in the Arkansas Valley contributed
one-half of this total (600).

about

An additional 195 geese were winter banded in the Cache la Poudre Valley
near Wellington.
The remaining 476 bands were placed on Canada goose
goslings as part of our continuing transplant program.
The Colorado River
transplants were hatched from eggs collected on the Yampa River west of
Craig.
All other transplants resulted from eggs collected in the Denver
area and from birds trapped in the Denver and Fort Collins areas.

�- 16 -

Table 2. Number of Canada geese banded by location and season of
year, 1969-70.

Number Banded
Sunnner
Winter

Total

Arkansas Valley

600

600

Cache La Poudre Valley

195

195

Location

South Platte Valley

69

69

San Luis Valley

160

160

North Park

194

194

Colorado River

53

53

Total

476

795

1,271

LITERATURE CITED
Hopper, R. M. 1968. Trapping and banding ducks and geese. Colo. Game,
Fish and Parks Div., Game Research Rept. Oct. , p. 7-12.
Hopper, R. M. 1969. Trapping and banding ducks and geese. Colo. Game,
Fish and Parks Div., Game Research Rept. Oct. , p. 9-14.

J

'J."

I.

,

I. 71"'} / _
Prepared by: fi!&lt;-cf:. {;..!.d; I' t. crq)]}':; i

Richard M. Hopper . v
Wildlife Researcher

�October,

- 17 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-88-R-IS

Migratory

Work Plan No.

1

Job No.

Job Title

Analysis

of Waterfowl

Bird Investigations
3

Banding Data

Period Covered:

April 1, 1969 through March 31, 1970

Personnel:

Michael R. Szymczak

ABSTRACT

A complete tape of all banding and recoveries related to North Park has
been ordered from the Migratory Bird Populations Stations Bird Banding
Laboratory.
When this tape is received the data will be analyzed and a
final report written.

1970

��October,

- 19 -

JOB PROGRESS

State

of

Project
Work

REPORT

COLORADO
--------~~~~~---------No.

Plan No.

Covered:

Personnel:

Migratory

W-88-R-15
1

Job Title
Period

1970

Job No.
Waterfowl

October

Bird

Investigations
5

Kill Survey

1, 1969 through July 1, 1970

Velma Merkle,
Grieb, Robert

Velma Fredrickson,
Raymond
Tully, and Howard Funk.

Boyd,

Jack

ABSTRACT

Duck stamps sold in Colorado in 1969 numbered 34,281, the most since 1958.
Estimated number of duck hunters on the east slope increased to 27,058.
Regular season length on the east slope remained at 33 days, but bag and
possession limits were increased to four and eight, respectively;
two and
four of which could be mallards.
Regular season harvest was estimated at
180,000 ducks, an increase of 40,000 over 1968-69.
Harvest and hunting
pressure on the west slope remained relatively stable, with approximately
24,000 ducks taken by about 3,400 hunters.
About 2,700 hunters took an
estimated 23,000 ducks during the special San Luis Valley duck season.
Harvest during the special early teal season in specified east slope areas
was estimated at 5,395 birds killed by 866 hunters.
Approximately
7,000
hunters bagged about 40,000 ducks during the experimental
late pointsystem season in Central Flyway, Colorado.
Canada goose harvest was indicated to have decreased for the third consecutive year.
Number of hunters remained quite stable at about 13,000
but harvest fell from just over 19,000 in 1968-69 to just under 18,000 in
1969-70.
Numbers of birds present in the Arkansas Valley seemed adequate
for better harvest, but the half-day shooting regulation might have had
an effect to some degree.
An estimated 4,172 geese were harvested in the
northcentral Colorado counties of Larimer, Weld, Boulder, Morgan and
Adams, for an increase of 59 percent over the 1968-69 season.

��- 21 -

WATERFOWL

KILL SURVEY

Howard

D. Funk

This survey of Colorado small game harvest is a cooperative effort between
personnel of Federal Aid Projects W-88-R and W-38-R, and the Game Planning
Services section of the Colorado Division of Game, Fish and Parks.
Data presented herein are results from two separate surveys; (1) the regular
annual questionnaire survey limited to purchasers of combination small game
hunting and fishing, resident small game hunting, and non-resident small game
hunting licenses, and (2) the separate questionnaire survey of holders of
the increasingly popular sportsman's license which also allows small game
hunting.

P. S. OBJECTIVE
To formulate waterfowl

harvest

regulations.

SEGMENT
To estimate
by species,

OBJECTIVES

the State harvest of waterfowl for the 1969-70 hunting
County, and interval of the season.

METHODS

season

AND MATERIALS

Techniques utilized were similar to those employed in the past with hunter's
names drawn in a mechanical random fashion from duplicate license sales
stubs.
The cutoff dates for license sales figures used in projections were
April 30, 1970 for the regular survey and February 1, 1970 for the sportsman's license survey.
Feasibility of utilizing April 30 as a cutoff date for
both surveys should be examined for use hereafter.
One follow-up letter
was sent to non-reporting hunters after an interval of approximately
three
weeks.
The regular survey questionnaire was revised this year in an attempt
to simplify reporting by hunters and ultimately obtain higher reporting
rates and more valid harvest information than in past years.
The sportsman's questionnaire was somewhat more simplified than the regular
questionnaire.
Harvest data collected on this survey under Federal Aid
Project W-38-R were added to results of the regular survey (Boyd 1970).

RESULTS

AND DISCUSSION

Regular survey questionnaires were sent to 13,283 randomly selected 1969
license buyers about January 15, 1970. A total of 8,106 responded for a
return of 61.0 percent.
This was one of the lowest response rates since

�- 22 -

the survey was initiated, even with the more simplified questionnaire.
Of
the 8,106 returns, 4,409 reported hunting, 3,672 reported they did not hunt,
and 40 were not usable.
Percentages of license buyers actually hunting were
calculated from the survey sample by license type (Table 1). The largest
percentage of license buyers not utilizing the opportunity to hunt was found
in the combination hunting and fishing license category; these people
using their fishing privilege only. Total sales of regular type licenses
permitting small game hunting during 1969, other than sportsman's licenses,
amounted to 166,941.
It is estimated that 97,876 of these actually hunted
one or more species of small game.

Table 1. License sales, percent
license type, 1969 season.

Sales

use and projected

estimates

of hunters by

Percent Used for
Hunting Small Game

Projected
Number Hunters

License

Type

Resident

Combination

127,914

51.5

65,876

Resident

Small Game

36,424

81.6

29,722

2,603

87.5

2,278

Non-resident

Small Game

166,941

Sub-total

Sportsman's

97,876

License

Resident

11,422

62.8

7,173

Non-resident

4,269

7.0

298

Sub-total

15,691

7,471

182,632

105,347

TOTAL

Survey questionnaires were sent to 3,158 sportsman's license buyers.
A
total of 2,287 responded for a return of 72.4 percent, considerably better
than the response rate for the regular survey.
Of the 2,287 returns, 1,191
reported hunting small game while 1,096 reported they did not.
Percentages
of respondents indicating they hunted were calculated separately for residents
and non-residents.
Thus, of the total sale of sportsman's licenses for
1969 (15,691), it is estimated that 7,471 of the buyers of this type of
license took part in hunting one or more species of small game.

�- 23 -

Total estimated number of regular and sportsman's license buyers hunting
one or more species of small game in 1969 was 105,347 which compares with
134,606 for 1968, 98,618 for 1967, and 114,492 in 1966.
Colorado duck stamp sales are listed in Table 2. Numbers sold in 1969
(34,281) indicate the continuing annual increase in waterfowl hunters from
the low of approximately 18,000 sold in 1962. However, sales still are not
of the magnitude experienced in 1957 and 1958 when almost 42,000 were sold
during the most recent peak period of waterfowl populations.

Table 2.

Duck stamp sales for Colorado, 1954-1969.

Year

Number of
Stamps Sold

Percent Change From
Previous Year

1954

32,450

+ 20.5

1955

39,107

+ 20.5

1956

36,303

7.2

1957

41,794

+ 15.1

1958

41,897

+

1959

31,431

- 24.9

1960

30,592

2.7

1961

24,854

- 18.8

1962

17,701

- 28.8

1963

22,940

+ 29.6

1964

25,282

+ 10.2

1965

20,537

- 18.8

1966

29,377

+ 43.0

1967

31,064

+ 5.7

1968

31,218

+

1969

34,281

+ 9.8

.02

.5

�- 24 -

Duck stamp buyers are classified in Table 3 by the type of waterfowl hunting
in which they participated in 1969 and the previous 15 years. There was a
significant increase in numbers of duck hunters, the highest since 1958,
while goose hunters remained about average.

Table 3. Estimated number of duck and goose hunters, average number of days
hunted, season length, and bag and possession limits, by year, 1954-1969.

Season Length
{Days 2
Ducks
Geese

Bag and
Possession
Limit
Ducks
Geese

7.64

60

60

5-10

2-2

17,634

8.87

75

60

5-10

2-2

34,793

12,477

7.37

75

60

5-10

2-2

1957

37,166

12,057

6.52

75

60

5-10

2-2

1958

38,773

14,705

5.78

90

60

4-8

2-2

1959

29,060

l3 ,647

5.70

50

75

4-8

2 -2

1960

29,480

14,107

6.05

60

75

3-6

2-2

1961

22,920

11 ,245

4.40

30

60

3-6

2-2

1962

13,918

9,159

5.27

25 11

75

2-4 fi.1

2-2

1963

17,989

10,841

5.66

6.64

35 II

75

4-8 'LI

2 -2

1964

19,189

13,678

6.30

7.37

40 II

75

4-8

JJ

2-2

1965

15,374

11 ,344

5.62

5.53

36 11

75

4-8 ]j

2-2

1966

23,635

15,807

5.96

6.25

60 11

75

3-6 'LI

2-2

1967

25,347

l3,748

6.49

6.43

60 1/

75

3-6 1/

2 -2

1968

26,528

13 ,467

6.18

6.48

33 1/

75

3-6 'LI

2-2

1969

30,436

13,282

5.94

6.45

33 II

75

4-8 21

2-4 '2.1

Year

Number
of Duck
Hunters

Number
of Goose
Hunters

Average Number
Days Hunted For
Ducks
Geese

1954

31,834

12,l36

1955

37,816

1956

11 Hunting regulations for East Slope.

West Slope had a general season of 75
days with 4 ducks in bag, and 8 in possession in 1962; and 90 days, 4 in bag
and 8 in possession in 1963, 1964, and 1965; and 90 days, 5 in bag and 10 in
possession in 1966 and 1967; and 86 days, 5 in bag and 10 in possession of which
3 and 6 respectively, could be mallards in 1968; and 86 days, 5 in bag and 10
in possession in 1969.
21 Two mallards allowed in bag, and four in possession on East Slope.
II One mallard and pintail allowed in bag and two in possession on East Slope,
three mallards or three pintai1s on West Slope.
21 One goose per day and
±I One mallard in bag and two in possession on East Slope.two in poss. on W. Slope.

�- 25 Duck Harvest

Duck hunting statistics for the 1969 season are tabulated and compared with
results of past years in Table 4. Total retrieved harvest of 203,629 ducks
was significantly greater than in 1968, an increase of approximately 21 percent; and the largest since 1958. Increased numbers of hunters, relaxed
hunting seasons, favorable publicity, and special seasons offering a variety
of hunting opportunity were probably all responsible for this increase.
Estimates on wounding losses were not obtained through the questionnaire
this
year, but they probably remained at between 10 and 20 percent of the retrieved
bag.
Estimates on species composition of the 1969 bag also were not obtained from
the survey this year.
Data from the previous 15 years are shown in Table
5. Mallards are calculated to have been responsible for an average of about
69 percent of the total bag over the years. However, with the special
seasons in effect again in 1969, especially the late point-system season,
mallards probably comprised from 75 to 80 percent of the total harvest as in
1968 when the late mallard drake-only season was a heavy contributor.
Duck harvest by intervals and season types is illustrated in Table 6 for
both the east and west slopes of the State, the Central and Pacific Waterfowl Flyways, respectively.
Harvest during the regular season on the east
slope was indicated to have been greatest during the early portion of the
season when probably the greatest variety of ducks was available.
Harvest
was then reduced somewhat during mid-season but increased considerably from
mid-November through the end of the season as more mallards came south.
Total harvest on the east slope during the 1969 regular season, about 111,000
birds, was estimated to have increased approximately 43 percent over the
previous year's harvest of about 78,000 ducks.
Harvest of ducks on the west slope also was indicated to have been greatest
during the early season period.
It then dropped somewhat, but remained
relatively stable throughout the season on a harvest per day basis.
Total
estimated west slope harvest decreased about 8 percent from the 1968 figure
of 25,954 to 23,781 birds (Table 6).
Special Teal Season.--An optional nine-day early teal season was granted
the states in the Mississippi and Central Flyways in 1969. Colorado chose
to take the season from September 1-9, but restricted the open area to North
Park (Jackson County) and the portion of the State east of Colorado Highway
71, U. S. Highway 350, and U. S. Interstate Highway 25. Estimated harvest
was 5,395 birds by 866 hunters or approximately 3 percent of the total east
slope duck harvest by 3 percent of the total east slope duck hunters (Table 6).
San Luis Valley Experimental Season.--This was the seventh year of experimental duck hunting seasons in the San Luis Valley.
As in 1968, bag limits
were based on a point-system of 70 points.
Various other species of ducks
able to withstand increased harvest were placed in the low point category
along with the mallard drake in 1969. Results of the season obtained through
special field observations and surveys will be reported in a Migratory Bird
Populations Station Administrative Report later in 1970 (Geis et ale 1970).
However, information on the season gathered from this survey are presented
here.

�- 26 -

Table 4.

Date

Duck harvest statistics, 1954-1969.

Slope

1954
1955
1956
1957
1958
1959
1960
1961

Number
of
Hunters

Average
Seasonal
Bag

Total
Estimated
Harvest

Wounding Loss
Percent
Number

Total
Estimated
Hunting
Mortality

31,834
37,816
34,793
37,166
38,773
29,060
29,480
22,920

5.6
6.7
5.9
6.8
6.1
4.2
5.0
3.8

179,856
253,367
185,737
254,587
236,515
122,924
147,400
86,408

14.5
13.1
16.3
14.1
12.3
15.5
13.1
21.4

30,396
38,182
36,195
41,679
33,088
22,417
22,257
23,608

210,252
291,549
221,932
296,266
269,603
145,341
169,667
110,016

1962

East
West
TOTAL

11,349
2 569
13,918

2.6
3.5
2.8

29,507
8,992
38,499

13.5
11.7
13.1

4,603
1,187
5,790

34,110
10,179
44,289

1963

East
West
TOTAL

15,627
2,362
17,989

5.1
3.6
4.9

80,167
8,503
88,670

10.7
9.7
10.6

9,636
916
10,552

89,803
9,419
99,222

1964

East
West
TOTAL

16,311
2,878
19,189

4.0
4.0
4.0

65,244
11 ,512
76,756

10.6
10.4
10.6

7,764
1,335
9,099

73,008
12,847
85,855

1965

East
West
TOTAL

12,747
2,627
15,374

5.0
4.3
4.9

64,245
11,322
75,567

18.7
15.4
18.2

12,007
1,744
13,751

76,252
13,066
89,318

1966

East
West
TOTAL

19,494
4,141
23,635

5.0
4.4
4.9

97,860
18,345
116,205

16.4
20.4
17.1

19,278
4,715
23,993

117,138
23,060
140,198

1967

East
West
TOTAL

22,432
2,915
25,347

6.5
3.7
6.2

146,032
10,698
156,730

12.0
22.9
12.9

20,006
3,178
23,184

166,038
13,876
179,914

1968

East
West
TOTAL

23,006
3,522
26,528

6.2
7.4
6.4

142,578
25,954
168,532

10.9
10.9
10.9

17,466
3,179
20,645

160,044
29,133
189,177

1969

East
West
TOTAL

27,058
3 378
30,436

6.7
7.0
6.7

179,848
23 781
203,629

- 1/

- 1/

1/ Data not requested in survey questionnaire.

1/

�- 27 -

Table 5.

Species

composition

of the bag, 1954-1968.

IS-Year Averagez 1954-1968
Number Harvested
Percent of Total

Species

Mallard

99,840

68.5

Pintail

4,941

3.4

Green-winged

Teal

12,622

8.7

Blue-winged

Teal

5,509

3.8

Baldpate

1,720

1.2

Gadwall

4,279

2.9

Shoveller

1,615

1.1

Scaup

1,139

0.8

Redhead

1,154

0.8

Canvasback

1,062

0.7

Others and Unknown

11,821

8.1

TOTAL

145,702

100.0

Survey data indicate 2,706 hunters bagged an estimated 23,021 ducks during
the October 1-18 season (Table 6). This compares with 1968 figures of 3,083
hunters harvesting 22,242 birds which indicates harvest remained quite
similar between years.
However, numbers of hunters listed above are rather
meaningless and are no doubt somewhat deflated since they refer to successful hunters rather than estimates of total hunters.
On the other hand,
estimated harvest is probably somewhat inflated because total harvest is
projected from data from the successful hunters.
The special survey,
because of its intensity, will produce more valid estimates of hunter
participation and success.
Experimental Point-System Season.--A late point-system duck season was
granted Colorado and five other Central Flyway states in 1969 in place of
the mallard drake-only season of 1968. Dates of the season were from
December 13, 1969 through January 4, 1970 with a 40-point daily bag limit.

�- 28 -

Table 6. Ducks bagged and hunting pressure by season type, area, and interval
of the 1969 season.

Dates

Estimated Harvest
No. of
% of
Ducks
Total

Estimated
Hunting Pressure
No. of
% of
Hunters
Total

EAST SLOPE
Teal Season (Special)
Sept. 1-9

5,395

3.0

866

3.2

23,021

12.8

2,706

10.0

Oct. 25-Nov. 3

49,458

27.5

9,957

36.8

Nov. 4-13

28,236

15.7

6,575

24.3

Nov. 14-26

33,452

18.6

7,116

26.3

40,286

22.4

7,035

26.0

San Luis Valley (Special)
Oct. 1-18
Regular

Exp. Point-System Season
Dec. 13-Jan. 4

WEST SLOPE
Regular
Oct. 11-0ct. 30

6,706

28.2

1,301

38.5

Oct. 31-Nov. 19

5,042

21.2

1,260

37.3

Nov. 20-Dec. 9

4,685

19.7

1,301

38.5

Dec. 10-Dec. 29

5,517

23.2

1,280

37.9

Dec. 30-Jan. 4

1,831

7.7

568

16.8

�- 29 -

Mallard drakes counted 10 points each, while mallard hens and all other
species counted 40 points each.
Thus, hunting pressure was again directed
toward mallard drakes which band recovery data indicate are undershot in
the western Central Flyway.
A special, covering results of this season has
already been prepared and distributed (Funk et al. 1970).
Results of this comprehensive survey suggest that 7,035 hunters bagged
40,286 ducks during this experimental season (Table 6). However, the special
point-system season survey indicated 11,285 hunters were active during the
season and bagged 35,886 birds.
Here again, as with the San Luis Valley
season and for the same reasons, estimated hunters from this survey are
deflated and estimated harvest is somewhat inflated.
Thus, the special survey results are again no doubt more accurate.
Results between years, the 1968 drake-only season as compared with the 1969
point-system season, suggest little change in hunter participation and total
bag between season or regulation types.
This survey produced estimates of
7,454 hunters bagging 42,631 birds in 1968 as opposed to 7,035 hunters taking
40,286 ducks this last season.
Special survey results from both years,
although different than those above, also suggested little change between
years; 11,081 hunters with a bag of 37,786 birds in 1968 as compared with
11,285 hunters bagging 35,886 ducks in 1969.
The point-system season was considered as successful or probably even more
successful than the drake-only season of 1969. It was certainly popular
with the hunters with approximately 37 percent of the total hunters in the
State participating in the season and taking about 18 percent of the total
State duck harvest.

Goose Harvest
Results of the 1969 goose season presented in Table 7 indicate 13,282 hunters
bagged an average of 1.4 birds each for a season total of 17,939.
This
suggests a decrease of about 8 percent in harvest and a 2 percent decrease
in hunters from the previous year.
However, the 1969 harvest was still
above the l6-year average of 17,703 geese.
As with ducks, no data were collected in the questionnaire survey to permit crippling loss estimates.
Species composition of geese in the 1969 bag, as with all years of the survey,
remained about 99 percent, or above, Canada geese.
The remainder is composed
of snow geese and an even smaller portion of white-fronted
geese.
Food and water conditions were excellent during the 1969 Arkansas Valley
goose season.
Census figures indicated goose populations in the Valley
numbered about 60,000 through fall and winter, the highest number ever to
remain into January.
The major concentrations were in the Eads Lakes and
Two Buttes Reservoir areas, but there were good populations scattered throughout the fall and early winter on other bodies of water.
Numbers of geese wintering in the Denver-Greeley-Fort
Collins area also
increased significantly in 1969 and contributed to an increased harvest.

�- 30 Table 7.

Year

Goose harvest

Slope

1954
1955
1956
1957
1958
1959
1960
1961

1954-1969.

statistics,

Number
of
Hunters

Average
Seasonal
Bag

Total
Estimated
Harvest

Wounding Loss.
Percent
Number

Total
Estimated
Hunting
Mortality

12,136
17,364
11,541
12,057
14,705
13,647
14,107
11,245

0.7
1.0
1.0
1.2
1.3
1.6
1.1
1.3

8,168
17,711
11,310
14,589
19,704
21,972
15,659
14,056

22.8
18.3
21.6
23.5
22.3
17.8
20.7
24.5

2,410
3,884
3,116
4,473
5,655
4,730
4,087
4,568

10,578
21,248
14,426
19,062
25,359
26,702
19,746
18,624

1962

East
West
TOTAL

8,828
331
9,159

1.5
1.3
1.5

13,241
430
13,671

19.5
0.0
19.1

3,218
0 1/
3,218

16,459
430
16,889

1963

East
West
TOTAL

10,462
379
10,841

1.7
0.3
1.6

17,785
114
17,899

15.2
0.0
17 .8

3,184
3,184

20,969
114
21,083

1964

East
West
TOTAL

13,295
383
13,678

1.8
0.5
1.7

23,931
192
24,123

19.0
0.0
18.9

5,624 1/
05,624

29,555
192
29,747

1965

East
West
TOTAL

10,941
403
11,344

1.2
1.0
1.2

13,239
419
13,658

24.6
27.0
24.7

3,257
113
3,370

16,496
532
17,028

1966

East
West
TOTAL

15,443
364
15,807

1.9
1.1
1.9

29,613
420
30,033

16.2
0.0
16.2

5,745
87
5,832

35,358
507
35,865

1967

East
West
TOTAL

13,598
150
13,748

1.7
1.7
1.7

23,117
255
23,372

19.8
19.8
19.8

5,710
63
5,773

28,827
318
29,145

1968

East
West
TOTAL

13,050
417
13,467

1.5
0.7
1.4

19,088
291
19,379

16.6
16.6
16.6

3,806
58
3,864

22,894
349
23,243

1969

East
West
TOTAL

12,990
292
13,282

1.4
0.9
1.4

17,691
248
17,939

- ]j

- ]j

- 2/

Sixteen-year

average

goose harvest

(1954-69) is 17,703.

1/ No cripples reported on the west slope.

2/ Date not requested

in survey questionnaire.

o 1/

�- 31 -

For the fifth year, goose permits with a six-bird season limit and other
special regulations were in effect in most of the Denver-Greeley-Fort
Collins
area.
In 1969, the permit area was enlarged but the limit was removed from
number of permits to be issued and 8,342 hunters obtained permits.
A special
random survey sample drawn from these permittees indicated that 3,535 hunters
actually hunted geese one or more times in the area.
They bagged 2,431
geese in Larimer County; 1,112 in Weld County; 383 in Boulder County; 146 in
Morgan County; and 100 in Adams County.
This is a total harvest of 4,172
geese or an increase of 59 percent over the 2,631 birds bagged in 1968.

Waterfowl

Harvest by County

The reader is cautioned that information presented in this section is
subject to a great deal more error in accuracy than estimates in previous
sections, since the original sample has been broken down to county, thus
decreasing the size of sample on which to base estimates.
This is probably
even more true of geese than ducks because there are many more duck hunters.
Consequently, it is realized that in some counties, both duck and goose
harvest has been over-estimated,
and in others, underestimated.
However,
despite this error, these data represent the most accurate information
possible at the present time.
Tables 8 and 9 compare the 1969 duck and goose harvests, respectively, with
averages of previous seasons by county within each waterfowl region.
These
regional divisions of the State were located on the basis of waterfowl migration, location, and topography; and permit a closer evaluation of annual
harvest, yearly changes in harvest patterns, and the effect of different
types of hunting seasons in various portions of Colorado.
Grossly, it appears that the greatest duck harvest again occurred in the
Central Region, although the percentage of total State bag for this area
decreased somewhat from the previous year.
The harvest increased about
10,000 birds from about 69,000 bagged in 1968 to 79,000 in 1969. Greatest
percentage increase took place in the Northeast Region with an estimated
40,000 birds taken, as compared to 26,000 in 1968. Yet many hunters still
do not seem willing to travel to the eastern portions of the State because
of the two mallard limit during regular season.
The east slope again contributed the bulk of duck harvest in 1969, about 88 percent of the State
total.
Greatest harvest again took place in Weld County.
Number of ducks
bagged increased significantly in the Northwest Region on the west slope,
but decreased somewhat in other regions.
The 1969 goose harvest compared quite closely with the fifteen-year
average, but was down somewhat from the 1968 season results (Table 9).
The Southeast Region again was the high area of harvest with 60 percent of
the total State goose bag. Kiowa, Baca, Bent, and Prowers were the top
harvest counties in that order in the Region.
The Central Region was next
high with approximately 36 percent of the total State harvest.

�- 32 -

Table 8.

Duck harvest by Region and County.

Waterfowl
Region
and County

1969 Duck Harvest
Number
Percent
Harvested
of Total

Estimated 1969
Hunting Pressure
Number
Percent
Hunters
of Total

Fifteen-year Average
1954-1968
Number
Percent
Harvested
of Total

EAST SLOPE
NORTHEAST
Cheyenne
Kit Carson
Lincoln
Logan
Morgan
Phillips
Sedgwick
Washington
Yuma
NORTHEAST TOTAL

T
2,878
1,079
12,409
12,230
899
3,417
2,338
52216
40,466

T
1.6
0.6
6.9
6.8
0.5
1.9
1.3
2.9
22.5

27
460
135
2,057
2,111
81
541
568
649
6,629

0.1
1.7
0.5
7.6
7.8
0.3
2.0
2.1
2.4
24.5

428
1,505
1,032
7,131
9,553
189
3,737
2,081
32158
28,814

0.3
1.2
0.8
5.6
7.6
0.1
3.0
1.6
2.5
22.7

SOUTHEAST
Baca
Bent
Crowley
Huerfano
Kiowa
Las Animas
Otero
Prowers
Pueblo
SOUTHEAST TOTAL

1,259
5,216
2,518
T
3,957
1,079
7,733
1,798
32237
26,797

0.7
2.9
1.4
T
2.2
0.6
4.3
1.0
1.8
14.9

162
758
298
54
487
162
920
325
568
3,734

0.6
2.8
1.1
0.2
1.8
0.6
3.4
1.2
2.1
13.8

1,189
3,179
2,776
597
1,535
1,100
1,908
3,882
32333
19,499

0.9
2.5
2.2
0.5
1.2
0.9
1.5
3.1
2.6
15.4

CENTRAL
Adams
Arapahoe
Boulder
Douglas
Elbert
E1 Paso
Jefferson
Larimer
Weld
CENTRAL TOTAL

8,992
1,619
6,834
1,079
540
2,158
1,798
15,467
402645
79,132

5.0
0.9
3.8
0.6
0.3
1.2
1.0
8.6
22.6
44.0

1,542
325
1,543
108
108
406
352
3,086
5,710
13,180

5.7
1.2
5.7
0.4
0.4
1.5
1.3
11.4
21.1
48.7

7,447
7,659
153
274
1,033
1,446
11,646
272642
59,025

5.9
1.4
6.1
0.1
0.2
0.8
1.1
9.2
21.7
46.7

8,813
3,597
2,338
9,892
5,755

4.9
2.0
1.3
5.5
3.2

1,109
270
216
866
595

4.1
1.0
0.8
3.2
2.2

4,394
2,434
785
5,481
3,806

3.6
1.9
0.6
4.3
3.0

30,395

16.9

3,056

11.3

16,900

l3.4

SAN LUIS VALLEY
Alamosa
Conejos
Costilla
Rio Grande
Saguache
SAN LUIS VALLEY
TOTAL

1,725

-----------------------------------------------------------------------------------

�- 33 -

Table 8.

Duck harvest by Region and County (continued) .

Waterfowl
Region
and County

1969 Duck Harvest
Number
Percent
Harvested of Total

HIGH COUNTRY (E)
Chaffee
1,259
Clear Creek
Custer
1,079
Fremont
180
Gilpin
Jackson
Lake
Park
540
Teller
HIGH COUNTRY TOTAL 3,058

Estimated 1969
Hunting Pressure
Number
Percent
Hunters
of Total

0.7
0.6
0.1

0.3
1.7

Fifteen-year Average
1954-1968
Number
Percent
Harvested
of Total

81
27
108
108

0.3
0.1
0.4
0.4

454
86
288
820

0.4
0.1
0.2
0.6

27

0.1

81
27
459

0.3
0.1
1.7

230
134
259
44
2,315

0.2
0.1
0.2
0.0
1.8

WEST SLOPE
NORTHWEST
Garfield
Moffat
Rio Blanco
Routt
NORTHWEST TOTAL

2,901
380
571
1~878
5,730

12.2
1.6
2.4
7.9
24.1

312
81
64
165
622

9.2
2.4
1.9
4.9
18.4

1,553
543
731
747
3,574

8.2
2.9
3.9
4.0
19.0

4,280
5,233
1,784

18.0
22.0
7.5

751
663
331

22.2
19.6
9.8

11,297

47.5

1,745

51.6

3,088
4,544
2,410
180
10,222

16.3
24.1
12.8
1.0
54.2

95

0.4

20
40

0.6
1.2

2,188

9.2

166

0.7

291
20
81

8.6
0.6
2.4

595
3,04l1

2.5
12.8

64
516

1.9
15.3

127
23
70
1,427
93
911
15
194
2,860

0.6
0.1
0.4
7.6
0.5
4.9
0.1
1.0
15.2

HIGH COUNTRY (W)
Eagle
1,569
Grand
285
Gunnison
1,570
Pitkin
143
Summit
143
HIGH COUNTRY TOTAL 3,710

6.6
1.2
6.6
0.6
0.6
15.6

145
20
226
64
40
495

4.3
0.6
6.7
1.9
1.2
14.7

750
451
612
282
90
2,185

4.0
2.4
3.2
1.5
0.5
11.6

WEST CENTRAL
Delta
Mesa
Montrose
Ouray
W. CENTRAL TOTAL
SOUTHWEST
Archuleta
Dolores
Hinsdale
La Plata
Mineral
Montezuma
San Juan
San Miguel
SOUTHWEST TOTAL

------------------------------------------------------------------------------------

�- 34 -

Table 8.

Duck harvest by Region and County (continued).

Waterfowl
Region
and County

1969 Duck Harvest
Number
Percent
Harvested of Total

Estimated 1969
Hunting Pressure
Number
Percent
Hunters
of Total

Fifteen-year Average
1954-1968
Number
Percent
Harvested
of Total

Summary by Region
NORTHEAST
40,466
SOUTHEAST
26,797
CENTRAL
79,132
SAN LUIS VALLEY
30,395
HIGH COUNTRY (E)
3,058
NORTHWEST
5,730
WEST CENTRAL
11,297
SOUTHWEST
3,044
HIGH COUNTRY (W)
3,710
TOTAL OF REGIONS 203,629

19.8
13.1
38.9
14.9
1.7
2.8
5.5
1.5
1.8
100.0

6,629
3,734
13,180
3,056
459
622
1,745
516
495
30,436

21.8
12.3
43.3
10.0
1.5
2.0
5.8
1.7
1.6
100.0

28,814
19,499
59,025
16,900
2,315
3,574
10,222
2,860
22185
145,394

19.8
13.4
40.7
11.6
1.6
2.4
7.0
1.9
1.5
100.0

EAST SLOPE
WEST SLOPE

88.3
11.7

27,058
3,378

88.9
11.1

126,553
18,841

87.0
13.0

179,848
23,781

Table 9.

Goose harvest by Region and County.

Waterfowl
Region
and County

1969 Goose Harvest
Number
Percent
Harvested of Total

Estimated 1969
Hunting Pressure
Number
Percent
Hunters
of Total

Fifteen-year Average
1954-1968
Number
Percent
Harvested
of Total

EAST SLOPE
NORTHEAST
Cheyenne
Kit Carson
Lincoln
Logan
Morgan
Phillips
Sedgwick
Washington
Yuma
NORTHEAST TOTAL

71
18
106
177

0.4
0.0
0.1
0.6
1.0

65
13
13
195
507

0.5
0.1
0.1
1.5
3.9

21
28
14
147
557

0.1
0.1
T
0.8
3.3

18
124

0.1
0.7

91
156

0.7
1.2

514

2.9

1,040

8.0

50
95
90
1,002

0.3
0.6
0.5
5.7

T

-----------------------------------------------------------------------------------

�- 35 Table 9.

Goose harvest by Region and County (continued).
Estimated 1969
Hunting Pressure
Number
Percent
Hunters of Total

Fifteen Year Average
1954-1968
Number
Percent
Harvested
of Total

Waterfowl
Region
and County

1969 Goose Harvest
Number
Percent
Harvested of Total

SOUTHEAST
Baca
Bent
Crowley
Huerfano
Kiowa
Las Animas
Otero
Prowers
Pueblo
SOUTHEAST TOTAL

2,070
1,946
973
71
3,786
88
531
884
248
10,597

11.7
11.0
5.5
0.4
21.4
0.5
3.0
5.0
1.4
59.9

987
909
494
39
1,676
104
351
585
182
5,327

7.6
7.0
3.8
0.3
12.9
0.8
2.7
4.5
1.4
41.0

5,284
1,911
699
116
3,051
122
347
2,208
179
l3,917

30.3
10.9
4.0
0.7
17.4
0.7
2.0
12.6
1.0
79.6

18
495
584
18

0.1
2.8
3.3
0.1

350
65
596
13

2.7
0.5
4.6
0.1

159
T
3,307
12787
6,368

0.9
T
18.7
10.1
36.0

65
13
3,131
22026
6,259

0.5
0.1
24.1
15.6
48.2

316
138
55
3
6
19
160
667
870
2,234

1.8
0.8
0.3
T
T
0.1
0.9
3.9
5.0
12.8

35
88
71

0.2
0.5
0.4

39
117
156

0.3
0.9
1.2

7
140
121
2
2

0.1
0.8
0.6
T
T

194

1.1

312

2.4

272

1.5

3
38

T
0.2

CENTRAL
Adams
Arapahoe
Boulder
Douglas
Elbert
E1 Paso
Jefferson
Larimer
Weld
CENTRAL TOTAL
SAN LUIS VALLEY
Alamosa
Conejos
Costilla
Rio Grande
Saguache
SAN LUIS VALLEY
TOTALS
HIGH COUNTRY (EAST)
Chaffee
Clear Creek
Custer
Fremont
Gilpin
Jackson
Lake
Park
Teller
HIGH COUNTRY TOTAL

T

0.0

l3

0.1

22

0.2

18

0.1

39

0.3

3

T

18

0.1

52

0.4

66

0.4

---------------------------------------------------------------------------------

�= 36 Table 9.

Goose harvest by Region and County (continued) •

Waterfowl
Region
and County

Estimated 1969
Hunting Pressure
Number
Percent
Hunters of Total

1969 Goose Harvest
Number
Percent
Harvested of Total

Fifteen Year Average
1954-1968
Number
Percent
Harvested of Total

WEST SLOPE
NORTHWEST
Garfield
Moffat
Rio Blanco
Routt
NORTHWEST TOTAL

203

81.8

157

53.3

1
109

0.8
74.6

45
248

18.2
100.0

97
254

33.3
86.6

110

75.4

4
1
2

3.4
0.1
1.3

7

4.8

3
3

2.0
2.0

26

17.8

26

17.8

1,002
13,917
2,234
272
66
110
7
3
26

5.7
78.8
12.7
1.5
0.4
0.6
0.1
0.0
0.2

WEST CENTRAL
Delta
Mesa
Montrose
Ouray
W. CENTRAL TOTAL
SOUTHWEST
Archuleta
Dolores
Hinsdale
La Plata
Mineral
Montezuma
San Juan
San Miguel
SOUTHWEST TOTAL

T

19

6.7

T

19

6.7

38

13.4

T

0.0

HIGH COUNTRY (WEST)
Eagle
Grand
Gunnison
Pitkin
Sunnnit
HIGH COUNTRY TOTAL (W) Sunnnary by Region
NORTHEAST
SOUTHEAST
CENTRAL
SAN LUIS VALLEY
HIGH COUNTRY (E)
NORTHWEST
WEST CENTRAL
SOUTHWEST
HIGH COUNTRY (W)

514
10,597
6,368
194
18
248

2.8
59.0
35.6
1.1
0.1
1.4

1,040
5,327
6,259
312
52
254

7.9
40.2
47.2
2.3
0.3
1.9

T

T

38

0.2

TOTAL OF REGIONS

17,939

100.0

13 ,282

100.0

17,637

100.0

EAST SLOPE
WEST SLOPE

17,691
248

98.6
1.4

12,990
292

97.8
2.2

17,491
146

99.2
0.8

�- 37 -

Waterfowl Management Units
Again this year, harvest information has been gathered on the basis of
waterfowl management units (Fig. 1). The purpose of this is a better alignment of data to provide information about specific flocks of ducks and
geese. For example, Unit 1 encompasses the area utilized by wintering ducks
at Jumbo Reservoir, Unit 5 is North Park, and Unit 15 is the San Luis Valley.
In many cases, duck and goose flock boundaries transcend county lines and
it has been difficult to put together county information so that we could
look at the influence of hunting pressure and harvest on separate flocks.
We had hoped that management units would help solve this. However, there
are still problems with interpreting results which have to be examined by
year. Results of this portion of the survey are displayed in Tables 10 and
11 for ducks and geese, respectively. Estimated numbers of hunters and
harvest for the 1969 season are shown by management unit and compared with
the five-year averages from 1964-68. No other comments will be made at this
time.

LITERATURE CITED
Boyd, R. J. 1970. Hunter harvest surveys. Colo. Div. of Game, Fish and
Parks, Game Res. Rept. July, Part Two. pp. 133-213.
Funk, H. D., and J. R. Grieb. 1969. Colorado small game hunter harvest
survey -- 1968. Colo. Div. of Game, Fish and Parks. Ft. Collins,
July 1. 38pp.
Funk, H.D., R. M. Hopper, J. R. Grieb, D. Witt, G. F. Wrakestraw, T. Kuck,
D. E. Timm, and G. W. Merrill. 1970. Preliminary evaluation of the
1969-70 experimental point-system duck season within the High Plains
Mallard Management Unit of the Central Flyway. Cent. Fly. Tech.
Comm. Rept. March. llpp., 12 Tables and Appendix.
Geis, A. D., et a1. 1970. "Progress report
1969 San Luis Valley experimental duck hunting season". Mig. Bird Pop. Sta. Admin. Rept. In press.

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_~~/~~.
__
_

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-c,
Howard D. Funk
Section Chief, Small Game Research
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�- 39 -

Table 10.

Estimated duck hunters and harvest by management unit, 1964-1969.

Management Number
Unit
Hunters

1969 Season
Number
Percent
of Total Bagged

Percent
of Total

Five-year Average
1964-1968
Number
Number
Percent
Hunters of Total Bagged

Percent
of Total

EAST SLOPE
1

2,219

7.3

15,107

7.4

1,265

5.9

6,448

5.4

2

2,760

9.0

15,827

7.8

1,118

5.3

5,800

4.9

3

4,492

14.7

28,056

13.8

3,234

15.2

17,730

14.9

4

3,030

10.0

13 ,668

6.7

1,993

9.4

11 ,065

9.3

5

108

0.4

540

0.3

101

0.5

567

0.5

6

5,466

18.0

33,452

16.4

3,946

18.6

23,742

20.0

7

271

0.9

899

0.4

273

1.3

1,160

1.0

8

595

2.0

3,237

1.6

341

1.6

1,569

1.3

9

974

3.2

5,935

2.9

413

1.9

2,048

1.7

10

758

2.5

5,755

2.8

582

2.7

2,221

1.9

11

81

0.3

180

0.1

252

1.2

783

0.6

12

1,840

6.0

12,769

6.3

1,072

5.0

5,087

4.3

13

1,028

3.4

7,913

3.9

729

3.4

3,276

2.8

14

379

1.2

4,137

2.0

233

1.1

1,180

1.0

15

3,057

10.0

32,373

15.9

2,784

13.1

20,301

17.1

Sub-total

27,058

88.9

179,848

88.3

18,336

86.2

102,975

86.7

WEST SLOPE
16

3,378

11.1

23,781

11.7

2,933

13.8

15,783

13 .3

GRAND
TOTAL

30,436

100.0

203,629

100.0

21,269

100.0

118,758

100.0

�- 40 -

Table 11.

Management
Unit

Estimated goose hunters and harvest by management unit, 1964-1969.

Number
Hunters

1969 Season
Number
Percent
of Total Bagged

Percent
of Total

Five-year Average
1964-1968
Number
Number
Percent
Hunters of Total Bagged

Percent
of Total

EAST SLOPE
1

299

2.3

89

0.5

238

1.7

l31

0.6

2

546

4.1

372

2.1

479

3.5

504

2.3

3

2,117

15.9

2,158

12.0

1,803

l3.2

1,917

8.6

4

3,117

23.4

2,848

15.9

1,581

11.6

1,758

7.9

5

26

0.2

0

0.0

43

0.3

l32

0.6

6

1,377

10.4

991

5.5

1,179

8.7

1,122

5.0

7

26

0.2

0

0.0

41

0.3

23

0.1

8

52

0.4

71

0.4

86

0.6

155

0.7

9

52

0.4

0

0.0

69

0.5

64

0.3

10

1,624

12.2

3,998

22.3

1,971

14.4

4,844

21.9

11

1,286

9.7

2,211

12.3

2,646

19.4

5,986

27.1

12

1,650

12.4

3,450

19.2

2,147

15.8

3,849

17.4

l3

584

4.4

1,362

7.6

627

4.6

964

4.4

14

65

0.5

35

0.2

l38

1.0

149

0.7

15

169

1.3

106

0.6

262

1.9

213

1.0

12,990

97.8

17,691

98.6

13,311

97.8

21,812

98.6

Sub-total

WEST SLOPE
16

292

2.2

248

1.4

298

2.2

300

1.4

GRAND
TOTAL

l3 ,282

100.0

17,939

100.0

13,609

100.0

22,112

100.0

�October,

- 41 -

JOB PROGRESS

State

Work

REPORT

COLORADO
--------~~~~~----------

of

No.

W-88-R-15

Migratory

plan No.

1

Job No.

Project

Job Title
Period

1970

San Luis Valley

Covered:

Personnel:

May

Cooperative

11, 1970 to October

Mallard

Bird

Investigations
12

Investigation

18, 1970

Charles Hayes, Bureau of Sport Fisheries and Wildlife;
Ron Blumberg, Harvey Bray, Gary Brown, Harold Burdick,
Earl Cochran, Harry Dobbs, Gordon East, John Goettl,
Art Gresh, Lloyd Hazzard, Glen Hinshaw, Wayne Knisely,
Wilber Ladd, Jr., Bob Mangus, Dick McDonald, Kris
Moser, Dick Norman, Bob Rosette, Wayne Russell, Gary
Will, Clait Braun, Howard Funk, Mike Szymczak, Richard
Hopper, Colorado Game, Fish and Parks Division.

ABSTRACT

The duck breeding population estimate in the San Luis Valley for 1969
totaled 27,425 pairs, including 11,816 pairs of mallards.
This was nearly
equal to the 1968 figure for all species combined, but mallards were down
3,000 pairs in 1969 from the previous year.
Sampling error (± 22.8 percent) remained consistent with the three previous years.
A point-system regulation was again tested in the San Luis Valley during
the October 1-18, 1969 experimental
season.
This season was similar to
that held in 1968 except for some changes in point allocations among the
various species and sexes.
In 1969, pintail drakes, shovelers, all teal
and coots were added to the 10-point category along with mallard drakes,
mallard hens, canvasbacks,
and redheads counted 60 points each, with all
others

35 points

each.

Colorado personnel assisted in conducting 56 hunter performance
surveys
during the 1969 season, including 26 on private land and 30 on public
land.
Survey cards were sen,t to the Bureau of Sport Fisheries and Wildlife
for analysis.

��- 43 -

SAN LUIS VALLEY

COOPERATIVE

MALLARD

INVESTIGATION

Richard M. Hopper

This is a cooperative investigation between the Bureau of Sport Fisheries
and Wildlife and the Colorado Game, Fish and Parks Division.
The objectives
stated below relate to the collection of data necessary in evaluating the
experimental season and include only those for which the Colorado Division
had primary reponsibility.
Only results pertaining to these objectives are
presented here.
Overall results of the 1969 experimental season and interpretation of the data appear in Administrative Report No. 195 (Hopper, et al.,
1970) prepared by members of both agencies and distributed by the Bureau
of Sport Fisheries and Wildlife, Branch of Wildlife Research, Migratory Bird
Populations Station.

P. S. OBJECTIVE
To develop a harvest

formula

for the San Luis Valley mallard

population.

SEGMENT OBJECTIVES
This study was designed to determine the influence of various types of
hunting regulations and bag limits on the local breeding population of
mallards in the San Luis Valley.
Colorado's part of the study is covered
under the following segment objectives:
1.
2.

Determine the size of the 1969 breeding population in the San Luis
Valley.
Assist in determining the ability and/or willingness of hunters to abide
by specific regulations and determine wounding loss during the 1969 experimental season.

METHODS

AND MATERIALS

Techniques for conducting the 1969 breeding population survey, air-ground
comparison study, and hunter performance survey remained the same as those
in 1967 (Segment 13). A discussion of these methods appears in a previous
report (Hopper and Rutherford, 1968) and will not be repeated here.

RESULTS AND DISCUSSION
Breeding
Air-Ground

Comparison

Population

Survey

Study

Comparisons of air and ground counts of ducks on 122.5 miles of selected
transect are shown by species in Table 1. The proportion identified from

�- 44 -

the air (visibility ratio) was again highest for the mallard (0.272),
excluding the small sample of "other divers" (0.333). The three species of
teal were the most difficult to identify from the air, as witnessed by their
visibility ratios ranging from 0.020 to 0.056.
The visibility ratio for
cinnamon teal for example, means that the air crew identified, as cinnamon
teal, only 5.6 percent of the cinnamon teal actually present on the airground comparison transects.
These ratios were applied to the regular air
transect counts as correction factors in calculating estimates of total
breeding pairs for Russell Lakes and the major portion of the Valley,
excluding Monte Vista National Wildlife Refuge and the total-count areas.

Table 1. Air-ground comparison
in the San Luis Valley, 1969.

of ducks counted

on 122.5 miles of transect

Species

Estimated Breeding Pairs
Ground
Air
Pairs
Percent
Pairs
Percent

Mallard

474

48.7

129

64.2

0.272

Gadwall

98

10.0

22

10.9

0.224

Pintail

88

9.0

22

10.9

0.250

Shoveler

58

5.9

13

6.5

0.224

Blue-winged

30

3.1

1

0.5

0.033

Cinnamon

143

14.6

8

4.0

0.056

49

5.0

1

0.5

0.020

7

0.7

0

0.0

0.000

Redhead

26

2.7

4

2.0

0.154

Other Divers

3

0.3

1

0.5

0.333

976

100.0

201

100.0

0.206

Teal

Green-winged
American

Teal

Widgeon

Totals

Duck Breeding

Population

Proportion
Identified
From the Air

Estimate

Square miles of habitat, sampling intensity, and estimated number of breeding
pairs of ducks are listed in Table 2 for the six areas included in the 1969
survey.
Breeding pair estimates are also included from previous years for
comparison.

�Table 2. Breeding pairs by density type as estimated from the San Luis Valley regular air transects,
and nesting transects, 1964-1969.
Square Miles
Habitat

Percent
Sample

1964

Estimated Breeding Pairsll
1967
1968
1966
1965

Monte Vista NWR

22

5.50

3,944

3,000

3,964

3,509

4,054

4,420

Russell Lakes

6

33.33

1,452

689

1,440

828

1,845

876

San Luis Lakes

7

100.00

--

60

105

86

277

813

Mishak Lakes

4

100.00

--

187

338

532

204

313

Adams Lake

1

100.00

--

--

6

17

26

103

Type

1969

I

Remainder of Valley

1,265

17.95

27,366

18,866

18,111

24,171

21,205

20,900

Totals

1,305

17.80

32,762

22,802

23,964

29,143

27,611

27,425

11 Determined from nesting transects on Monte Vista National Wildlife Refuge and from aerial transects
on all other areas.

-I='
\J1

�- 46 -

The 1969 inventory yielded an estimated 27,425 breeding pairs of ducks, or
essentially the same as the 27,611 pairs obtained in 1968. This figure is
still somewhat lower than the high years of 1964 and 1967, but above the
1965 and 1966 estimates.
Species composition of the 1969 San Luis Valley breeding population is shown
in Table 3. Composition figures for past years are included for comparison.
The estimated mallard breeding population for 1969 was down 3,000 pairs from
1968, which resulted in a reduction in their composition from 54 percent of
the total number of breeding pairs of ducks in 1968 to 43 percent in 1969.
Each of the three species of teal increased substantially over 1968, making
up for the mallard deficit.
All other species contributed about the same
numbers of pairs in 1969 as in 1968. The 1969 sampling error, calculated
at
22.8 percent, remained consistent with the three previous years.

±

Harvest

Information

The point-system regulation was tested during the 1969 experimental duck
season in the San Luis Valley for the second consecutive year.
Some changes
in point allocations among the various species and sexes were made in 1969,
with pintail drakes, shovelers, all teal, and coots being added to the 10point category along with the mallard drake. Mallard hens, redheads, and
canvasbacks counted 60 points each, while all others tallied 35 points each.
All other rules remained the same as in 1968.
The 1969 regulation, although seemingly more complex, was based upon a more
realistic allocation of points, such as might be applied throughout one or
more flyways.
Mallard females were increased from 40 to 60 points in an
attempt to further reduce shooting pressure on these birds by encouraging
hunters to avoid them. Additional species and sexes that could withstand
greater harvest were placed in the low-point category (10 points) in an
effort to direct more shooting pressure toward them and away from mallard
females.
ObViously, the goal of the point-system regulation is to emphasize
the harvest of lightly shot species and sexes and discourage the harvest of
heavily shot species and sexes.
Thus, hunters were tested in the San Luis
Valley to determine their reaction to a point-system regulation relatively
more complicated than the one they faced in 1968.
State personnel assisted in collecting hunter performance data during the
1969 experimental season.
This work was necessary for determining hunter
attitude and compliance with the point-system regulation.
Fifty-six hunter
performance surveys were conducted by State men, including 26 on private
land and 30 on public land. The survey cards were sent to the Bureau of
Sport Fisheries and Wildlife for tabulation and analysis along with surveys
submitted by Federal personnel.
Results of this work are presented in the
joint report mentioned earlier in this report (Hopper, et al., 1970).

�Table 3.

Species composition of the San Luis Valley duck breeding population, 1964-1969.
1964
Pairs

%

1965
Pairs

%

1966
Pairs

%

1967
Pairs

%

1968
Pairs

%

1969
Pairs

%

Mallard

14,319

43.7

11,177

49.1

13 ,064

54.5

15,502

53.2

14,816

53.7

11,816

43.1

Gadwall

1,660

5.1

2,779

12.2

2,340

9.8

3,115

10.6

2,570

9.3

2,735

10.0

Pintail

7,531

23.0

1,878

8.2

2,644

11.0

3,085

10.6

2,737

9.9

2,347

8.6

263

0.8

1,981

8.7

409

1.7

2,100

7.2

1,561

5.6

2,889

10.5

Blue-winged Cinnamon Teal

2,337

7.1

3,249

14.2

1,719

7.2

2,905

10.0

3,310

12.0

5,496

20.0

Shoveler

1,030

3.1

978

4.3

1,788

7.5

1,503

5.2

1,211

4.4

1,055

3.8

93

0.3

88

0.4

0

0.0

157

0.5

0

0.0

0

0.0

5,017

15.3

667

2.9

1,448

6.0

455

1.6

1,252

4.5

1,031

3.8

512

1.6

5

--

552

2.3

321

1.1

154

0.6

56

0.2

Totals

32,762

100.0

22,802

100.0

23,964

100.0

29,143

100.0

27,611

100.0

27,425

100.0

~ .05 Standard Error ~/

+31.0%

Species

Green-winged Teal

.j::-

-...J

American Widgeon
Redhead
Other Divers 1/

+28.0%

+23.0%

1/ Includes lesser scaup, canvasback, ruddy duck, and common merganser.
~/ Excluding Monte Vista NWR and Russell Lakes.

+20.9%

+23.8%

+22.8%

�- 48 -

LITERATURE CITED
Hopper, R., H. Funk, R. Buller, A. D. Geis, and E. M. Martin. 1970.
Progress report: 1969 experimental duck hunting season in the San Luis
Valley of Colorado - an evaluation of the "point system'! in regulating
harvest. U.S.D.I., Bur. Sport Fisheries and Wildl., Migratory Bird
Pop. Sta., Admin. Rpt. No. 195. 20 p.
Hopper, R. M., and W. H. Rutherford. 1968. San Luis Valley cooperative
mallard investigation. Colo. Game, Fish and Parks Div., Fed. Aid
Game Res. Rpt., Oct. p. 33-42.

Prepared by

..JUt,- ,. ~7_% .Jf(;,)J'~
Richard M. Hopper
Wildlife Researcher

-~

�October

- 49 -

JOB PROGRESS

Work

REPORT

COLORADO

State of
No.

W-88-R-ls

Migratory

Plan No.

1

Job No.

Project

Job Title
Period

1970

Investigations
13

Determination
of Methods for Developing and
Managing Waterfowl Habitat--Shallow
Impoundment Study

Covered:

Personnel:

Bird

April

1, 1969 to December

Ralph

Baker

and Richard

15, 1969

Hopper

ABSTRACT

Pre-impoundment
studies of impoundment No. 1 were initiated in Segment 15.
Permanent, ~-square-meter
quadrats were selected as one means of describing
vegetation associated with the impoundment sites before and after flooding.
Forty-eight quadrats were established,
32 on a portion of the impoundment
characterized
by native meadow vegetation,
and 16 on a portion previously
farmed.
Forty-seven
species of plants were encountered
in the 48 quadrats,
46 in the native meadow quadrats, and eight in the quadrats located on the
previously farmed area.
Three species of grasses, inland saltgrass (Distichlis stricta) bluestem
wheatgrass
(Agropyron smithii), and sand dropseed (Sporobolus cryptandrus)
exhibited the highest frequencies
(percent of quadrate of occurrence)
in
the native meadow with figures of 93.8, 84.4, and 78.1 percent, respectively.
Inland saltgrass, bluestem wheatgrass,
and cheatgrass brome (Bromus tectorum)
showed the greatest densities, with averages of 168.5, 62.2, and 50.6 sterns
per quadrat, respectively.
Summercypress
(Kochia scoparia) occurred in all 16 quadrate on the previously farmed area, while cornmon Russian thistle (Salsoli kali) was found
in 14 (87.5%) and lettuce (Lactuca ludoviciana)
in 13 (81.2%).
Surnmercypress and cornmon Russian thistle also had the highest densities with
1,070 and 269 sterns per quadrat.
Twelve soil samples were collected at random on the impoundment site.
These
varied from sandy loam to clay loam in texture.
Hydrogen-ion
concentrations
(pH) range from 7.5 to 8.5. Phosphorus, zinc, and iron were present in
smaller amounts on the previously farmed area than on the native meadow
portion.

�- 50 -

RECOMMENDATIONS

1.

Eliminate plans for construction of two drawdown impoundments because
of the absence of a sufficient quantity of water.
Drawdown studies
can be initiated to the three existing impoundments upon termination
of studies involving permanent water levels.

2.

Spread bentonite over extreme north end of borrow area in an attempt
to reduce any water loss from the impoundment through seepage.

�- 51 -

SHALLOW

IMPOUNDMENT

STUDY

Richard M. Hopper

The shallow impoundment study continued in Segment 15 for the third consecutive year as part of a long-term investigation designed to acquire knowledge in the development and management of waterfowl habitat.
The first
years progress report covered the selection, location, and description of
the proposed impoundment sites (Hopper 1968), while the second years report
described the design and construction of the dikes and ditching systems for
the three permanent impoundments (Hopper 1969). This report covers mainly
pre-impoundment soil and vegetative studies of the first shallow impoundment
to be flooded.

P. S. OBJECTIVE
To determine the effects of (a) shallow, permanent impoundments; and (b)
shallow, drawdown impoundments on waterfowl use and harvest at Bonny Reservoir.

SEGMENT
1.
2.
3.

OBJECTIVES

Design and construct dikes and ditching system for two drawdown impoundments.
Conduct pre-impoundment studies of one permanent impoundment and two
drawdown impoundments.
Flood above three impoundment sites in September and record waterfowl
use until freeze-up.

METHODS

AND MATERIALS

Pre-Impoundment

Studies

Vegetation
Permanent quadrats were selected as one means for describing vegetation
associated with the impoundment sites before and after flooding.
Fortyeight, ~-square-meter
(~ m. x 1 m.) quadrats were established in impoundment
No. 1 on the area that will constitute the shallow-water and moist soil portion of the impoundment.
A different method of vegetative sampling is
planned for the deep-water portion of the impoundment (borrow area) adjacent
to the upstream face of the dikes where the fill material for the dikes
was obtained.
Locations of the 48 quadrats were determined through stratified random
sampling.
The ditch crossing impoundment No. 1 (Fig. 1) separates two portions
of contrasting vegetation and it was necessary to stratify the impoundment

�- 52 area for sampling purposes.
The area to the north of the ditch covers about
two-thirds of impoundment No.1 and consists of native. meadow-type vegetation that has been hayed and grazed in the past but not broken by the plow.
Land south of the ditch was farmed prior to 1969, with small grains as the
predominant crop. As a result, 32 quadrats were placed in the north area
and 16 in the south area.
Twelve reference points were randomly selected and permanently marked with
steel posts along the main (east) dike of impoundment No.1
(X - Z, Fig. 1),
with eight points in section X - Y and four in Y - Z. Corresponding points
were permanently marked at the opposite (west) side of the impoundment by
running a compass-line perpendicular to the main dike and across the impoundment from each of the 12 points on the main dike.
These reference points
were designated A through L.
Four quadrats were established along each compass-line within a 300-ft. distance west of the west edge of the borrow area (deep water portion).
This
300 ft. distance was stratified in an east-west direction into four, 75 ft.
segments, with one quadrat being randomly placed in each segment along each
compass line. These segments were designated as zones 1 through 4. This
will insure sampling of vegetation in various depths of water and on moist
and dry soils, since conditions will become progressively drier in an eastwest direction upon flooding of the impoundment.
The northwest and southeast
corners of each quadrat were permanently marked with 18 inch wooden stakes.
An aluminum tag, identifying a quadrat by reference point and number, was
placed on the northwest stake.
Designation A-2 on an aluminum tag, for
example, refers to the quadrat in the second 75 ft. zone west of the borrow
area along compass line A.
Information collected at each quadrat included: (1) date, (2) quadrat number,
(3) list of species present, and (4) number of individual plants or stems
of each species.
It was difficult to distinguish individual plants of some
species, especially many grasses and grass-like plants, thus stemS originating
at the soil surface were counted.
Analysis of vegetative data involved the
determination of frequency (percentage of quadrats in which a species occurred) and density (average number of stems or individuals per quadrat).
Photographs were taken along compass-lines A through L in an east-west direction.
Twelve quadrats were also randomly selected for photographing.
Quadrat photos were taken 10 feet from the southeast stake in an east-west
direction, with a yard stick placed behind the northwest stake.

Soils
A core of the top six inches of soil was collected adjacent to eight randomly selected vegetative quadrats and along four compass lines in the deepwater portion of impoundment No.1, with the aid of a soil auger.
The 12
samples were sent to the Colorado State University Soil Testing Laboratory
for analyses of the following items: pH, conductivity, lime, organic matter,
phosphorus, potash, zinc, iron, and texture.

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--=-~~-~

�- 54 RESULTS

Construction

AND DISCUSSION

of Drawdown

Impoundments

Because of the absence of a sufficient quantity of water to flood three
permanent impoundments plus two drawdown impoundments, the latter will not
be constructed and drawdown studies will be temporarily abandoned as part
of the overall investigation.
The three impoundments already in existence
can be utilized for drawdown studies following termination of studies involving permanent water levels.

Pre-impoundment

Studies

Vegetation
Forty-seven species of plants were encountered in the 48 ~-square-meter quadrats studied in impoundment No.1 during the period August 13-16, 1969
(Tables 1 and 2). We were unable to identify four of these plants, mainly
because of the lack of sufficient vegetative and flowering parts to work
with in keying them out. Forty-six of the 47 species were found in the 32
quadrats on the native meadow north of the ditch that crosses impoundment
No.1
(Table 1). While only eight species occurred in the 16 quadrats
located on the previously farmed area south of the ditch (Table 2). All
eight species found on the farmed area, except green brist1egrass (Setaria
viridus), were also found on the native meadow portion.
Tables 1 and 2 show the frequency (number and percent of quadrats of occurrence) and density (average number of stems per quadrat) for each plant
species encountered on the native meadow and farmed areas, respectively.
Only three plant species, all grasses, occurred in over 75 percent of the
32 native meadow quadrats.
These were: inland sa1tgrass (Distich1is stricta)
(93.8%), b1uestem wheatgrass (Agropyron smithii) (84.4%), and sand dropseed
(Sporobo1us cryptandrus) (78.1%).
Fleabane (Erigeron be11idastrum
was found
in 18, or 56.2 percent of the quadrats.
Two forbs, western ragweed (Ambrosia
coronopifo1ia) and summercypress (Kochia scoparia), had the next highest frequencies, with each occurring in 50 percent of the quadrats.
Six additional
species were found in 25 to 50 percent of the quadrats.
Thus, only 12 of
the 46 species encountered on the native meadow occurred in 25 percent or
more of the 32 quadrats.
Inland sa1tgrass, b1uestem wheatgrass, and cheatgrass brome(Bromus tectorum)
showed the greatest densities in the native meadow quadrats, with average
figures of 168.5, 6.2, and 50.6 stems per quadrat, respectively.
All other
species exhibited average densities of less than 31 stems per quadrat.
Summercypress occurred in all 16 quadrats on the previously farmed area,
while common Russian thistle (Sa1s01i ka1i) was found in 14 (87.5%) and
lettuce (Lactuca 1udoviciana) in 13 (81.2%) (Table 2). Groundcherry
(Physalis 10ngifo1ia) was the only other species of the eight encountered
that occurred in more than one quadrat.
Summercypress also had by far the
greatest density, averaging 1,070 stems, or in this case individual
plants, per quadrat.
These plants were noticeably stunted, as might be
expected with the extremely high density.
The density of common Russian

�- 55 -

thistle (269 stems per quadrat) amounted to only about one-fourth that of
summercypress, but was greater than any species in the native meadow.
All
other species in the farmed area had relatively low densities.
Species:
area curves, as described by Oosting (1956) were applied to the
native hay and farmed samples to determine the minimum number of ~-squaremeter quadrats required for adequate sampling of vegetation on each area.
This procedure indicated that the number of quadrats used (32 and 16, respectively) was sufficient for the present plant compositon; however, changes
in composition brought about by flooding the impoundment could result in a
revision of the number of quadrats needed.

Soils
Results of the analyses of 12 soil samples collected on August 18, 1969 are
shown in Table 3. Sample numbers AO, EO, HO, and KO were taken from the
borrow area (deep water area) where a foot or more of the top soil had been
removed and used for construction of the dike. These four subsurface samples
were clay loam in texture.
All other samples were taken from the original
soil surface and these consisted of lighter textures of loams and sandy
loams. Generally, it seems that surface soils of the impoundment are light
to medium in texture, but that at least some of the area is underlain with
heavy soils.
The subsurface samples (clay loams) were similar to the surface samples in
all respects except for lime and organic matter content.
The clay loams
were all high in lime content, whereas the lighter surface samples ranged
from low to high in this regard.
Organic matter content was generally lower
in the clay loam samples (0.8 - 1.3%) than in the loam and sandy loam samples
(1.3 - 3.6%). This difference in organic matter content would be expected
when comparing surface and subsurface soils.
Hydrogen-ion concentrations
(pH) of all soil samples ranged from 7.5 to 8.5,
or slightly to moderately alkaline.
Samples taken on the previously farmed
portion of the impoundment (Jl, J4, and K2) showed smaller amounts of phosphorus, zinc, and iron than did the samples collected from the native meadow.
Many years of cropping on farmland can be responsible for declines in these
elements.

Flooding

of Impoundment

Site

An attempt was made in mid-August to fill impoundment No. 1 from the existing
well.
It was obvious after several days that the well did not produce
enough water to meet this demand.
A sufficient quantity of water was
yielded to fill the borrow area and to begin spreading over the flats, but
at least an additional six vertical inches were needed to obtain the desired
amount of flooded area. A second well will be drilled during the summer of
1970. This well should produce more water than the first, and hopefully, the
two together will handle all three of the impoundments.

�- 56 -

Table 1. Frequency and density data for plant species encountered
quadrats on the native meadow portion of impoundment No.1.

Frequency
No. Quadrats
of
Occurrence

Percent

in 32

Density
Total Ave. No.
No.
Stems/
Stems Quadrat

Scientific

Species
Namel/
Common Name~/

Agropyron

smithii

Bluestem Wheatgrass

27

84.4

1,991

62.2

Ambrosia

coronopifolia

Western

16

50.0

146

4.6

Ragweed

Andropogon

gerardi

Big Bluestem

1

3.1

18

0.6

Aristida

longiseta

Red Threeawn

3

9.4

418

13 .1

Artemisia

caudata

Sagebrush

8

25.0

222

6.9

Artemisia

filifolia

Sand Sagebrush

2

6.2

Artemisia

ludoviciana

Louisana Sagebrush

8

25.0

123

3.8

Asclepias

incarnata

Swamp Milkweed

1

3.1

1

T

Asclepias

pumila

Plains Milkweed

2

6.2

20

0.6

Asclepias

speciosa

Showy Milkweed

2

6.2

4

0.1

Astragalus

spp.

Milkvetch

1

3.1

1

T

Bouteloua

gracilis

Blue Grama

11

34.4

978

30.6

Bouteloua

spp.

Grama

3

9.4

277

8.7

11

34.4

1,620

50.6

Bromus tectorum

Cheatgrass

Carex praegracilis

Sedge

7

21.9

549

17.2

Cirsium spp.

Thistle

2

6.2

4

0.1

Conyza canadensis

Horseweed

4

12.5

23

0.7

Danthonia

spp.

Danthonia

1

3.1

8

0.2

Delphinium

spp.

Larkspur

6

18.8

16

0.5

Distichlis

stricta

Inland Saltgrass

30

93.8

5,392

168.5

Fleabane

18

56.2

186

5.8

Erigeron

bellidiastrum

Brome

Fleabane

Elymus canadensis

Canada Wildlrye

1

3.1

2

0.1

Haplopappus
cephalus

Goldenweed

1

3.1

1

T

Rush

4

12.5

237

Juncus

spp.

phyllo-

7.4

�- 57 -

Table 1. Frequency and density data for plant species encountered in 32
quadrats on the native meadow portion of impoundment No.1, continued.

Species
Scientific Namell
Common Name21

Frequency
No. Quadrats
of
Occurrence
Percent

Density
Total Ave. No.
No.
Stemsl
Stems Quadrat

Kochia scoparia

Summercypress

16

50.0

772

24.1

Lactuca 1udoviciana

Lettuce

3

9.4

18

0.6

Liatris punctata

Dotted Gayfeather

7

21. 9

56

1.8

Me1i1otus alba

White Sweetclover

7

21.9

145

4.5

Muhlenbergia asperifolia

Alkali Muhly

1

3.1

4

0.1

Opuntia spp.

Prickly pear

1

3.1

1

T

Physalis longifolia

Groundcherry

5

15.6

13

0.4

Plantago spp.

Plantain

7

21.9

34

1.1

Polygonum aviculare

Prostrate Knotweed

2

6.2

3

0.1

Ratibida columnifera

Prairie coneflower

10

31.2

76

2.4

Salsola kali

Common Russian Thistle

8

25.0

463

14.5

Solanum rostratum

Buffalobur Nightshade

3

9.4

7

0.2

2

6.2

7

0.2

Alkali Sacaton

5

15.6

287

9.0

Sporobolus cryptandrus Sand Dropseed

25

78.1

986

30.8

Solidago graminifolia Grassleaf Goldenrod
Sporobo1us airoides

Stellaria media

Chickweed

2

6.2

5

0.2

Stipa comata

Needle and Thread

7

21. 9

690

21.6

Tragopogon porrifolius Salsify

2

6.2

3

0.1

Unknown No. 1

7

21.9

56

1.8

Unknown No. 2

3

9.4

9

0.3

Unknown No. 3

1

3.1

15

0.5

Unknown No. 4

1

3.1

2

0.1

II After Harrington (1954).
II After Kelsey and Dayton (1942).

�- 58 -

Table 2. Frequency and density data for plant species encountered
quadrats on the previously farmed area of impoundment No.1.

Scientific

Species
Namell

Connnon Name~./

Frequency
No. Quadrats
of
Occurrence
Percent

in 16

Total
No.
Stems

Density
Ave. No.
Stemsl
Quadrat

Kochia scoparia

Sunnnercypress

16

100.0

17 ,123

1,070.2

Lactuca

ludoviciana

Lettuce

13

81.2

58

3.6

Physalis

long ifo lia Groundcherry

5

31.2

8

0.5

Salsola

ka l i,

Connnon Russian

14

87.5

4,310

269.4

Setaria

viridus

Green bristlegrass

1

6.2

1

0.1

Solanum

rostratum

Buffalobur

1

6.2

1

0.1

1

6.2

2

0.1

Stellaria

media

thistle

nightshade

Chickweed

II After Harrington (1954).
~I After Kelsey and Dayton (1942).

Excessive seepage of water from the impoundment may be responsible for the
inability of the first well to fill the impoundment.
Soils at the extreme
north end of the borrow area appeared to be lighter in texture than the clay
loam soils found by sampling in other portions of the borrow area.
Bentonite
could be used to help remedy any seepage problem that may exist.
Intensive waterfowl use observations will not begin until impoundment No. 1
is filled to the desired level. However, waterfowl began using this impoundment innnediately upon initial flooding.
Water was maintained in the impoundment throughout the fall and winter.
Varying numbers of ducks were seen
using the impoundment during the fall, with a peak number of 43 individual
ducks being observed on October 9, 1969.

�Table 3

0

Soil test results of 12 samples collected in impoundment noo 1 on August 18, 1969.

Organic
Matter

Conductivity
(salts)

Lime

(%)

(umhos / em)

Phosphorus
(ppm)

Potassium
(ppm)

Zinc
(ppm)

Iron
(ppm)

No r,

Texture

pH
(paste)

AO

Clay Loam

709

High

103

0.6

8.7

417+

0.59

7.2

EO

Clay Loam

8.1

High

0.9

05
0

9.2

281

0029

7.9

HO

Clay Loam

7.9

High

1.1

05
0

7.4

417+

0.41

8.5

KO

Clay Loam

8.5

High

008

0.5

2.2

417+

1.38

6.5

B1

Loam

709

High

2.7

0.5

8.1

385

1.40

12.5

C3

Sandy Loam

7.5

Low

2.1

0.4

11.1

277

0 76

6.8

Sample

0

F1

Loam

7.5

Low

2.9

0.5

5.2

417+

0092

6.0

F4

Loam

708

Medium

2.7

05
0

5.9

417+

1.37

3.4

G2

Loam

707

Low

306

0.5

8.5

417+

1.90

4.8

J1

Loam

7.8

Low

2.5

0.3

4.1

417+

0042

2.7

J4

Sandy Loam

709

High

1.3

0.2

3.3

417+

0030

2.7

K2

Sandy Loam

709

High

108

004

3.3

417+

0.50

2.5

.
\JI

1.0

�- 60 -

LITERATURE CITED
Harrington, H. D. 1954. Manual of the plants of Colorado.
Denver. 666pp.

Sage Books,

Hopper, R. M. 1968. Determination of methods for developing and managing
waterfowl habitat--sha11ow impoundment study. Colo. Game, Fish and
Parks Div., Fed. Aid Game Res. Rpt., Oct., pp. 43-49.
Hopper, R. M. 1969. Determination of methods for developing and managing
waterfowl habitat--sha11ow impoundment study. Colo. Game, Fish and
Parks Div., Fed. Aid Game Res. Rpt. Oct., pp. 65-71.
Kelsey, H. P., and W. A. Dayton. 1942. Standardized plant names.
Horace McFarland Co., Harrisburg, Pa. 675pp.
Oosting, H. J. 1956. The study of plant communities.
Co ,, San Francisco. 440pp.

Prepared by ...••
£q...... .AA"..::W~J~ia~L:&lt;::'-)-..I.n~"~1--,~"2.L-...·
..!:::..,th~J.L:.'.J-?i~_
Richard M. Hopper~
Wildlife Researcher

J.

W. H. Freeman and

�October

- 61 -

JOB PROGRESS

State

Work

REPORT

COLORADO

of
No.

W-88-R-15

Migratory

Plan No.

1

Job No.

Project

Job Title
Period

1970

Determination
Waterfowl Habitat

Covered:

Personnel:

of Methods
- Ammonium

April

1, 1969 to March

Ralph

Baker,

Wilber

Bird Investigations

14

for Developing and Managing
Nitrate Pothole Blasting StudX

27, 1970

Ladd,

Jr., and Richard

Hopper

ABSTRACT

The most common species of vegetation adjacent to the potholes were
(Scirpus americanus) and sedge (Carex spp.).
Twenty other species,
grass and grass-like plants, also surrounded the potholes.

bulrush
mostly

Submerged aquatic vegetation was found in 70, or 91 percent of the 77 potholes included in this portion of the study.
Seven species of plants were
encountered, with muskgrasses
(Chara spp.) and pondweed (Potamogeton foliosus)
dominating in both frequency of occurrence and density.
Vegetation was
encountered more frequently in the 75 and 150 lb. potholes than in the 25
and 50 lb. sizes.
The two larger sized potholes also contained about twice
the density of vegetation as occurred in the smaller sizes.
Average loss of pothole depth from 1968 to 1970 varied from a low of 3.5
inches for 25 lb. potholes in Block B to a high of 7.5 inches for 50 lb.
potholes in Block B. Overall, the loss averaged 11.0 percent for all 25
lb. potholes, 13.1 percent for both 75 and 150 lb. potholes, and 15.1
percent for 50 lb. potholes.
Spring, summer, and fall observations
in 1969 resulted in a total 197 duckvisits for the 82 potholes studied.
Nearly all of this use occurred during
the spring period.
Mallards contributed
50 percent of the duck-visits,
followed by blue-winged
teals and gadwalls with about 22 and 18 percent,
respectively.
The 150 lb. potholes received the most overall use with an
average of 3.23 duck-visits per hour, followed by the 75 lb. potholes with
2036.
As in 1968, the 75 lb. potholes were the most efficient in regard
to cost per duck-visit in 1969.

��- 63 -

AMMONIUM

NITRATE

POTHOLE BLASTING

STUDY

Richard M. Hopper

The third consecutive year of field work was completed for the pothole
blasting study during Segment 15. The first year involved mostly designing
of the study, selection of a study area, and the actual blasting of the 84
potholes.
Figure 1 shows the layout of the study area. The second year,
as well as the third year reported here, included the evaluation of waterfowl
use of the potholes during the spring, summer, and fall periods.
Information
related to vegetation associated with the potholes was also collected during
the latter two years.
Results of the first two years of study appear in the
previous reports (Hopper 1968, Hopper 1969).

P. S. OBJECTIVE
To evaluate various size potholes blasted with ammonium nitrate in terms of
(a) life expectancy, (b) plant succession, (c) soil and water characters,
(d) waterfowl use and hunting potential, and (e) cost.

SEGMENT
1.
2.
3.

OBJECTIVES

Collect data pertaining to the vegetative, soil, and water characteristics associated with the potholes.
Obtain pothole measurements.
Collect waterfowl use and harvest data on and adjacent to the potholes.

METHODS

AND MATERIALS

Methods and materials employed during Segment 15 remained the same as those
used in Segment 14 (Hopper 1969) and will not be repeated here. An additional method was employed to describe the vegetation adjacent to the potholes.
Vegetation within 30 feet of the edge of each pothole was mapped by species
on graph paper (8 squares per inch).
Pacing was utilized to obtain the
approximate area occupied by each plant species or group of species.
Reference to the maps then revealed the relative abundance of each species by
pothole.
A number was assigned to each species ranging from 5 (most abundant) to 1 (least abundant) to serve as a scale of estimated importance.

RESULTS AND DISCUSSION

Vegetative,
Vegetation

Adjacent

Soil and Water Characteristics

to Potholes

Mapping of vegetation surrounding the 84 potholes resulted in the identification of 22 species of plants of obvious abundance on the study area (Table
1). Additional species are known to be present, but were considered of minor

�- 64 importance by the investigator.
Grasses and grass-like plants constituted over one-half of the species listed. Most are commonly associated
with moist environments such as that provided by the presence of the high
water table on the study area.
The most common plant species associated with the potholes on the study area
were Scirpus americanus (bulrush), and Carex spp. (sedge), the latter represented by at least two species (Table 2). These were also by far the most
abundant species in the individual blocks and adjacent to each of the four
different sizes of potholes.
Eleocharis macrostachya (spikesedge) is the
only other species that occurred in abundance in all three blocks and adjacent to all four pothole sizes. Juncus balticus (rush) was important in
all of these categories except the 150 lb. pothole size. Three other species,
Melilotus alba (sweetclover), Sitanion hystrix (squirrel tail), and Typha
latifolia (cattail), were fairly consistent in occurrence by block and
charge size but varied in rank of importance.
Melilotus alba is the only
species of the nine listed in Table 2 not classified as a grass or grasslike plant.

Submerged

Vegetation

in Potholes

Submerged aquatic vegetation in the 84 potholes was sampled during the period August 13-15, 1968. Sampling consisted of dragging a garden rake through
each pothole the following number of times according to charge size:
150 lb.-8; 75 lb.-6; 50 lb.-4; 25 lb.-3. One drag of the rake through a
pothole constituted one sample; thus, 441 samples were taken in the 84 potholes.
Seven potholes were later eliminated from the vegetative analysis
because they contained submerged tumbleweeds that hindered the sampling procedure.
Seven species of submerged aquatic plants were encountered during sampling
of the 77 potholes included in the analysis.
The frequency of occurrence of
each of these species is expressed as "pothole" frequency (PF) and as
"sample" frequency (SF) in Tables 3-4, with comparisons by block and charge
size.
"Pothole" frequency refers to the percentage of potholes in which a
given plant species occurred, while "sample" frequency means the percentage
of samples (individual drags) that yielded a given plant species.
Submerged
aquatic vegetation was found in 70, or 90.9 percent of the 77 potholes.
About 84 percent of the 410 samples produced vegetation.
Chara spp. (muskgrass) was the most frequently encountered species, being present in 74 percent of the potholes and 47 percent of the samples.
Potamogeton foliosus
was next in occurrence, with 47 and 42 percent, respectively, followed by
two other species of pondweeds (K. nodosus and
pectinatus).

K.

Submerged vegetation was more commonly encountered in Block A than in Blocks
B or C (Table 3, Fig. 1). All potholes yielded vegetation in Block A, compared to 88 and 86 percent in Blocks Band C, respectively.
"Sample" frequencies were 88 percent in Block A, as opposed to 81 percent in Block Band
82 percent in Block C. In Block A, Chara spp. was by far the most frequently
encountered plant, but its importance in this regard was equaled by some
species of the genus Potamogeton in Blocks Band C.

�- 65 Table 1.

List of major plants found adjacent

Scientific Name
and Abbreviation

1/

Common Name

Agropyron

desertorum

Apocynum

medium

Asclepias

incarnata

~. speciosa

to potholes.-

(Agde)

Wheatgrass

(Apme)

Dogbane

(Asin)

Milkweed

(Assp)

Bromus tectorum

Milkweed

(Brte)

Bromegrass

Carex spp. (Casp)

Sedge

Cicuta douglasi

Water hemlock

Distichlis

(Cido)

stricta

Elymus canadensis

(Dist)

Saltgrass

(Elca)

Wildrye

Eleocharis

macrostachya

(Elma)

Equisetum

kansanum

(Eqka)

Horsetail

Helianthus

annuus

(Hean)

Sunflower

Spikesedge

Juncus balticus

(Juba)

Rush

Kochia scoparia

(Kosc)

Summer-Cypress

Melilotus

alba (Meal)

Populus

angustifolia

(Poan)

Cottonwood

Sagittaria

latifolia

(Sala)

Arrowhead

Scirpus

acutus

~. americanus
Sitanion

Typha

(Scac)

Bulrush

(Scam)

Bulrush

hystrix

Sporobuolus

(Sihy)

airoides

latifolia

Sweetclover

Squirrel

(Spai)

Dropseed

(Tyla)

1/ Names according

to Harrington

Cattail
(1954).

tail

�- 66 -

Table 2. Order of estimated importance of most abundant
pothole study area, by block and pothole size.

Plant Species
A

Block
B

C

plants associated

with the

in Order of Estimated Importance
Charge Size
50
25
75

150

Total

Scam

Scam

Scam

Scam

Scam

Scam

Scam

Scam

Casp

Casp

Casp

Casp

Casp

Casp

Casp

Casp

Meal

Sihy

Tyla

Elma

Elma

Juba

Meal

Elma

Elma

Elma

Elma

Tyla

Juba

Elma

Elma

Tyla

Sihy

Juba

Juba

Sihy

Sihy

Tyla

Tyla

Juba

Spai

Meal

Elca

Juba

Tyla

Meal

Sihy

Meal

Juba

Meal

Sihy

The 150 and 75 lb. charge sizes had higher "pothole" and "sample" frequencies
than the 50 or 25 lb. sizes (Table 4). "Pothole" frequencies were 95 and
100 percent, respectively, for the former sizes and about 84 percent for
both of the latter ones. Similarly, "sample" frequencies were 88 and 90 percent for the two largest charges and 72 percent for the smallest sizes.
All
four charge sizes were nearly alike in regard to species importance, with
Chara spp. dominating in frequencies, followed by the three species of Potamogeton.
However, frequencies were considerably higher for these species in
the 150 and 75 lb. potholes than in the 50 and 25 lb. potholes.
Densities of the various species of submerged vegetation are compared in
Tables 5-6 by block and charge size. Density, as used here, refers to the
average number of rake teeth covered (maximum of 14) by each species of vegetation per sample (drag). The number of teeth covered by vegetation for all
species combined averaged 8.5 per sample, with Chara spp. and Potamogeton
foliosus each contributing 3.1 to this total.
Potamogeton nodosus and f.
pectinatus each yielding an average coverage of 1 tooth per sample, were the
next most abundant species.
Chara spp. was the major plant in Block A (4.3 teeth/sample) and second in
importance in Blocks Band
C (2.1 and 2.9 teeth/sample, respectively).
Potamogeton foliosus was the most abundant species in Blocks Band C, averaging 4.4 and 4.0 teeth covered per sample, respectively.
Block C supported
the greatest vegetative density of the three blocks (9.7 teeth/sample),
while Blocks A and B were practically equal in density with about 7.8 teeth
per sample.

�Table 3. Comparison of frequency of occurrence of submerged aquatic vegetation among blocks, as
determined from 410 samples in 77 potholes.

.

Freguenc:l!:
of Occurrence ~Percent2
Block C
Block B
SF
SF
PF
PF

PF

SF

Total

Species

Block A
SF?:..!
PF1/

Ceratophyllum demersum

0.0

0.0

4.0

0.8

0.0

0.0

1.3

0.2

Chara spp.

91.7

64.9

68.0

39.4

64.3

38.1

74.0

47.1

Najas guadalupensis

0.0

0.0

0.0

0.0

3.6

0.7

1.3

0.2

Potamogeton foliosus

20.8

12.2

52.0

56.1

64.3

56.5

46.8

42.2

Potamogeton nodosus

20.8

13.0

32.0

19.7

60.7

40.1

39.0

24.9
I

Potamogeton pectinatus

41.7

27.5

36.0

15.9

25.0

15.0

33.8

19.3

Zannichellia palustris

20.8

13.7

0.0

0.0

0.0

0.0

6.5

4.4

TOTAL

100.0

87.8

88.0

81.1

85.7

82.3

90.9

83.6

1/ Percentage of the total number of potholes in a given block in which a species occurred.
1/ Percentage of the total number of samples in a given block in which a species occurred.

0'\

-...J

�Table 4. Comparison of frequency of occurrence of submerged aquatic vegetation among charge sizes, as
determined from 410 samples in 77 potholes.

Species

Freguenc~ of Occurrence
50 lb.
75 lb.
150 lb.
SF?:.../
SF
PF
SF
PF
PFli

~Percent2
25 lb.
SF
PF

PF

SF

Ceratoph~llum demersum

0.0

0.0

0.0

0.0

0.0

0.0

5.6

1.8

1.3

0.2

Chara spp.

90.0

50.0

90.0

55.0

47.4

30.3

66.7

44.4

74.0

47.1

Najas guadalupensis

0.0

0.0

5.0

0.8

0.0

0.0

0.0

0.0

1.3

0.2

Potamogeton foliosus

60.0

49.4

70.0

56.7

26.3

21.0

27.8

18.5

46.8

42.2

Potamogeton nodosus

50.0

30.0

50.0

25.0

21.0

14.5

33.3

24.1

39.0

24.9

Potamogeton pectinatus

50.0

23.1

40.0

16.7

36.8

27.6

5.6

1.8

33.8

19.3

Zannichellia palustris

5.0

1.9

10.0

8.3

0.0

0.0

11.1

9.2

6.5

4.4

TOTAL

95.0

88.1

100.0

90.0

84.2

72.4

83.3

72 .2

90.9

83.6

Total

1/ Percentage of the total number of potholes of a given charge size in which a species occurred.
?:.../
Percentage of the total number of samples of a given charge size in which a species occurred.

0'\

co

�= 69 -

~o

0

&lt;,

U

'-&gt; O
"- .

~O
00 0
(,)
~0

0

0

o

0

0

0

0

0
0

0

o o

000
CO
~0
0
000
~0 0 0
o 0
~0

0

0

0
0 &lt;3

0

0

0

0

o

0

octo

o
o o o o 0
o
o
o
~o
o o o
o
o
o
~O 0
o o o
'\J

~0

&lt;,

0

0

F"i.g . 1. Poth
position andole
sizeblasti
of potholes
ng study
by ar
b~~~k~hoWing the

0

0

�- 70 -

Table 5. Density comparisons of submerged aquatic vegetation among blocks, as
determined from 410 samples in 77 potholes.
Density II
Species

A

Block
B

Ceratophy11um demersum

0.0

0.1

0.0

T?:../

Chara spp.

4.3

2.1

2.9

3.1

Najas guadalupensis

0.0

0.0

T

T

Potamogeton foliosus

0.6

4.4

4.0

3.1

Potamogeton nodosus

0.5

0.6

1.9

1.0

Potamogeton pectinatus

1.3

0.7

0.8

0.9

Zanniche 11ia palustris

1.0

0.0

0.0

0.3

TOTAL

7.7

7.9

9.7

8.5

C

Total

1/ Average number of rake teeth covered per sample.

I..! Less than 0.1.

Table 6. Density comparisons of submerged aquatic vegetation among charge
sizes, as determined from 410 samples in 77 potholes.

Species

150

Density 1/
Charge Sizes {lbs·2
75
50
25

Ceratophy11um demersum

0.0

0.0

0.0

T?:../

T

Chara spp.

3.8

3.3

1.9

2.3

3.1

Najas guadalupensis

0.0

0.1

0.0

0.0

T

Potamogeton foliosus

3.3

4.6

1.5

1.4

3.1

Potamogeton nodosus

1.1

1.2

0.5

1.4

1.0

Potamogeton pectinatus

1.1

0.8

1.3

T

0.9

Zanniche11ia palustris

0.2

0.5

0.0

0.7

0.3

TOTAL

9.5

10.5

5.2

5.9

8.5

1/ Average number of rake teeth covered per sample.

"fj Less than 0.1.

Total

�- 71 -

Potholes blasted with the two largest charge sizes (75 and 150 lb.) contained nearly twice the density of vegetation as occurred in those created
with 2S and 50 lb. charges, about 10 teeth per sample as opposed to less
than 6 (Table 6). Chara spp. was the most abundant plant for three out of
four of the charge sizes, the 75 lb. size being the only exception.
Potamogeton foliosus was first in density in the 75 lb. potholes and second in the
other three sizes of potholes.

Soil and Water Characteristics
The collection and analysis of soil and water samples was not done during
Segment 15. This work is planned for Segment 16.

Pothole Depth Measurements
One means used in the attempt to assess the life expectancy of the various
size potholes was to determine changes in pothole depth over the period of
the study.
Pothole depth measurements collected in March 1968 and March
and April 1970 are compared in Table 7. The figures shown are average depths
for all potholes of a given charge size in each block, i. e., the average
depth of the seven 25 lb. potholes in Block A was 45.4 inches in 1968. Individually, all potholes, except A254 and B253 were of greater depth in 1968
than in 1970. Potholes C253 and C1504 were omitted from the measurement
analysis because muskrat activity had diverted a stream of silt-laden water
into them.
Average loss of depth from 1968 to 1970 varied from a low of 3.5 inches for
25 lb. potholes in Block B to a high of 7.5 inches for 50 lb. potholes in
Block B. Overall, the loss of depth in two years averaged 11.0 percent for
all 25 lb. potholes, 13.1 percent for both 75 and 150 lb. potholes, and
15.1 percent for 50 lb. potholes.
Unfortunately, the above differences in depth between years may not reflect
the natural filling-in process of the potholes.
Flood waters covered the
entire pothole study area twice during the two years that lapsed between
depth measurements.
The South Fork of the Republican River, which flows
adjacent to the study area, overflowed its banks on July 26, 1968 and again
on August 22, 1969, causing no apparent damage to the potholes, but no doubt
contributing some to their loss of depth.
The flood waters rose gradually
over the study area and dense vegetation slowed its movement and prevented
damage to the potholes from washing action.
However, the extent of silt
deposition in the potholes is unknown.

Waterfowl
Determination

of Waterfowl

Use and Harvest

Use by Watching

Use of the potholes by waterfowl was observed during seven periods in the
spring, summer, and fall of 1969, starting on April 1 and terminating on
November 13 when the potholes froze.
These seven observation periods ranged

�- 72 -

from two to five days in length and totaled 24 days.
Potholes in Block A
were observed 33.99 hours, and those in Blocks Band C were watched 32.91
and 26.42 hours, respectively.
Potholes A1507 and C1504 were again eliminated from the study because of biased use figures.
Ducks were the major group of waterfowl to use the potholes in 1969, but
two instances of Canada goose-use were recorded in the spring.
The 82 potholes in all three blocks combined, received a total of 197 duck-visits during
the above periods and hours of observation.
Separately, Blocks A, B, and
C accounted for 68, 50 and 79 duck-visits, respectively.
These figures are
not completely comparable because the three blocks were not observed for
exactly the same morning and evening hours or for the same number of hours.
The total number of duck-visits recorded in 1969 (197) was considerably less
than in 1968 (502) because fewer hours were spent observing in 1969, particularly during the spring period when most of the duck use occurs.
Thus,
total amount of use is not comparable between the two years.
The following species of ducks, listed in order of decreasing occurrence,
were observed to utilize the potholes during the observation periods in 1969:
mallard, blue-winged teal, gadwall, American Widgeon, shoveler, ring-necked
duck, and green-winged teal. Mallards contributed 50 percent of the duckvisits, followed by blue-winged teals and gadwalls with about 22 and 18 percent respectively.
Tables 8-10 present duck use per hour of observation by observation period
and season of the year for the three blocks separately, while Table 11 shows
these data for all blocks combined.
Potholes in Block C received the most
use in 1969, just as they did in 1968, averaging 3.15 duck-visits per hour
of observation.
Blocks A and B received 2.22 and 1.52 duck-visits per hour,
respectively, in 1969, with these two blocks being reversed in amount of use
in 1968. Total use for all three blocks combined equalled 6.89 duck-visits
per hour. Differences in magnitude of the above figures between years cannot
legitimately be compared, since the observation periods may have occurred at
a time of peak duck use during one year and not the other.
Observations conducted during the spring period accounted for by far the
greatest amount of duck use, with an average of 19.42 duck-visits per hour
for all three blocks combined (Table 11). This compares with figures of 1.37
and 0.59 duck-visits per hour during the Summer and fall periods, respectively.
The same relationship existed for each of the three blocks individually.
The 150 lb. potholes received the greatest overall use of the four different
charge sizes in 1969, with an average of 3.23 duck-visits per hour for all
three blocks combined (Table 11). Next were the 75 lb. potholes with 2.36
duck-visits per hour, while the 25 and 50 lb. potholes received only 0.28
and 1.02 duck-visits per hour, repectively.
Reference to use in individual
blocks show that:
(1) 150 lb. potholes exceeded all others in Block A; (2)
150 and 75 lb. potholes led with almost identical figures of 0.55 and 0.52
duck-visits per hour, respectively, in Block B; (3) 75 lb. potholes exceeded
all others in Block C; and (4) 50 lb. potholes exceeded 25 lb. potholes in
all three blocks.

�Table 7.

Chargel/
Size
(Lbs.)

Comparison of pothole depth measurements in March 1968 and March-April 1970 by charge size and block.

Average DeEth ~Inches2
Total
1968

Block A
1970

Diff.

25

45.4

40.3

50

45.7

75
150

1968

Block B
1970

5.1

40.3

39.3

6.4

43.0

37.8

42.6

38.8

Diff.

Block C
1968
1970

Diff.

1968

1970

Diff.

Percent Loss
of Depth

36.8

3.5

39.5

33.8

5.7

41.8

37.2

4.6

11.0

47.9

40.4

7.5

47.3

40.0

7.3

47.0

39.9

7.1

15.1

5.2

39.9

34.1

5.8

40.8

35.3

5.5

41.2

35.8

5.4

l3 .1

3.8

43.2

36.7

6.5

44.8

37.5

7.3

43.4

37.7

5.7

l3 .1

1/ Twenty-one potholes of each charge size except for the 25 lb. and 150 lb. sizes which had only 20 each.

-...J
w

�- 74 Table 8. Duck use of potholes in Block A by charge size, based upon watching
from observation points, 1969.

Observation
Period

No. Hours
Observed

25

Average No. Duck-Visits/Hour
Charge Size 17
50
75
150

Total

Spring
4/1-4/3

3.08

0.0'1:../

2.60

2.60

7.96

13.16

4/28-5/1

8.58

0.0

0.0

0.82

3.13

3.95

Sub-total

11.66

0.0

0.69

1.29

4.40

6.38

6/2 -6/4

4.50

0.0

0.0

0.0

0.26

0.26

8/31-9/3

5.08

0.0

0.0

0.0

0.0

0.0

Sub-total

9.58

0.0

0.0

0.0

0.12

0.12

10/8-10/10

2.25

0.0

0.0

0.0

0.0

0.0

10/24-10/28

7.50

0.0

0.0

0.0

0.0

0.0

11/12-11/13

3.00

0.0

0.0

0.0

0.0

0.0

Sub-Total

12.75

0.0

0.0

0.0

0.0

0.0

33.99

0.0

0.24

0.44

1.54

2.22

Summer

Fall

TOTAL

1/ Seven potholes of each charge size except for the 150 lb. size which had
only six potholes.

1/ Combined duck use for all seven potholes in a row.

�- 75 Table 9. Duck use of potholes in Block B by charge size, based upon watching
from observation points, 1969.

Average No. Duck-Visits/Hour
Charge Size 1.7
50
75
150

Total

0.652:./

1.95

4.22

1.95

8.77

8.58

0.12

0.35

0.23

1.16

1.86

11.66

0.26

0.77

1.29

1.37

3.69

6/2 -6/4

6.00

0.0

0.50

0.33

0.17

1.00

8/31-9/3

1.75

0.0

0.0

0.0

0.0

0.0

Sub-total

7.75

0.0

0.39

0.26

0.13

0.78

10/8-10/10

4.00

0.0

0.0

0.0

0.25

0.25

10/24-10/28

8.00

0.0

0.0

0.0

0.0

0.0

11/12-11/13

1.50

0.0

0.0

0.0

0.0

0.0

Sub-total

13.50

0.0

0.0

0.0

0.07

0.07

32.91

0.09

0.36

0.52

0.55

1.52

Observation
Period

No. Hours
Observed

25

4/1-4/3

3.08

4/28-5/1
Sub-total

Spring

Sununer

Fall

TOTAL

1/ Seven potholes of each charge size.
2:./Combined duck use for all seven potholes in the row.

�- 76 Table 10. Duck use of potholes in Block C by charge size, based upon watching
from observation points, 1969.

Observation
Period

No. Hours
Observed

25

Average No. Duck-Visits/Hour
Charge Size 1.1
SO
75
150

Total

3.00

2.67

7.00

13.67

Spring
4/1-4/3

3.00

1.00'2:..1

4/28-5/1

4.92

0.41

0.41

4.47

1.42

6.71

Sub-total

7.92

0.63

1.39

3.79

3.54

9.35

6/2-6/4

3.25

0.0

0.0

0.0

0.72

0.72

8/31-9/3

1. 75

0.0

0.0

0.0

0.0

0.0

Sub-total

5.00

0.0

0.0

0.0

0.47

0.47

10/8-10/10

4.00

0.0

0.0

0.0

0.0

0.0

10/24-10/28

8.00

0.0

0.0

0.88

0.0

0.88

11/12-11/13

1.50

0.0

0.0

0.0

0.0

0.0

Sub-total

13.50

0.0

0.0

0.52

0.0

0.52

26.42

0.19

0.42

1.40

1.14

3.15

Summer

Fall

TOTAL

l/Seven potholes of each charge size except for the 150 lb. size which had
only six potholes.
~/ Combined duck use for all seven potholes in the row.

�- 77 Table 11. Duck use of potholes in Blocks A, B, and C combined by charge size,
based upon watching from observation points, 1969.

Observation
Period

No. Hours
Observed

25

Average No. Duck-Visits/Hour
Charge Size 1/
50
75
150

Total

Spring
7.55

9.49

16.91

35.60

0.53

0.76

5.52

5.71

12.52

31.24

0.89

2.85

6.37

9.31

19.42

6/2-6/4

13.75

0.0

0.50

0.33

1.15

1.98

8/31-9/3

8.58

0.0

0.0

0.0

0.0

0.0

Sub-total

22.33

0.0

0.39

0.26

0.72

1.37

10/8-10/10

10.25

0.0

0.0

0.0

0.25

0.25

10/24-10/28

23.50

0.0

0.0

0.88

0.0

0.88

11/12-11/13

6.00

0.0

0.0

0.0

0.0

0.0

Sub-total

39.75

0.0

0.0

0.52

0.07

0.59

93.32

0.28

1.02

2.36

3.23

6.89

4/1-4/3

9.16

4/28-5/1

22.08

Sub-total
Summer

Fall

TOTAL

1/ Twenty-one potholes of each charge size except for the 150 lb. size which
had only 19 potholes.
~/ Combined duck use for all 21 potholes of the same charge size in the three
blocks.

�- 78 The relationship of duck use and pothole cost among the four charge sizes
are compared in Table 12. The cost figures shown are merely comparative
and not actual, since they represent duck use that occurred only during the
observation periods.
The 75 lb. potholes cost less per duck-visit than the
other charge sizes in Block C ($2.17) and for all three blocks combined
($3.49).
The 150 lb. size was the least expensive in Block A ($2.51) while
in Block B the 50 lb. potholes cost the least. The 25 lb. potholes were
the most expensive of the charge sizes in all three blocks in relation to
duck use.
For all three blocks combined, the 75 lb. potholes were the most efficient
size in regard to cost per duck-visit in 1969, just as they were in 1968.
However, this has not been replicated in each block individually, but the
application of statistical tests should assist in the interpretation.
Such
tests will be conducted and presented in the final report.

Nesting

Study

Three mallard nests were discovered on the nesting transects in 1969, two
during the May survey and one on the June search.
Two of the nests were
destroyed and the other appeared abandoned.
One abandoned nest (mallard)
was found off-transect.
The four nests were 5, 36, 54, and 279 feet from the nearest pothole.
They
were located in dead sedge (Carex spp.) or bulrush (Scirpus americanus), or
a mixture of the two. Height of vegetation at the nest sites varied from
16 to 23 inches.

Harvest

and Hunter-Use

No harvest or hunter-use was observed on the pothole study area during the
periods: September 1-3 (special teal season), October 25-28, and November
12-13, 1969. The potholes froze over the night of November 13, making further
observations unnecessary.
Duck-use observations conducted during these same
three periods yielded a total of only seven duck-visits in the three blocks
combined.
Large numbers of ducks are attracted to the habitat immediately
adjacent to the pothole study area. This habitat consists of cattail marsh,
beaver ponds, and flooded timber.
Because of this local availability of
ducks, perhaps some could be encouraged to visit the pothole study area
through the use of decoys and calls.
To my knowledge this has never been
attempted.

�Table 12. Relationship of pothole cost and duck use among the four charge sizes by block during the
observation periods, 1969.

Charge
Size
(Lbs.)

Ave. Cost/
Pothole
(Dollars)

No.
DuckVisit

25

4.87

0

Block A
Cost!
Duck-visit
(Dollars)
---

Block B
Cost/
No.
Duck-visit
DuckVisits (Dollars)

3

11.36

No.
DuckVisits

5

Block C
Cost/
Duck-visit
(Dollars)

6.82

Total
No.
DuckVisits

Cost/
Duck-visit
(Dollars)

8

12.78
I

50

7.11

8

6.22

12

4.15

11

4.52

31

4.82

75

11.47

15

5.35

17

4.72

37

2.17

69

3.49

150

18.81

45

2.511/

18

7.32

26

4.3~/

89

4.0211

TOTAL

10.36

68

4.07

50

5.92

79

3.51

197

4.31

1/ Adjusted to express use of only six potholes of the 150 lb. charge size in Blocks A and C instead of seven.

-..J
\0

�- 80 -

LITERATURE

CITED

Hopper, R. M. 1968. Determination of methods for developing and managing
waterfowl habitat--arnmonium nitrate pothole blasting study.
Colo.
Div. Game, Fish and Parks, Fed. Aid Game Res. Rpt. Oct. p. 51-64.
Hopper, R. M.
1969. Determination of methods for developing and manging
waterfowl habitat--arnmonium nitrate pothole blasting study.
Colo.
Div. Game, Fish and Parks, Fed. Aid Game Res. Rpt. Oct. 73-85.

Prepared

by _£"-.. +-L=~:'=::. ~}L-0~~/_·-\-,-n:..-....L!j_---=:.2io....I....:;.4· .. 9;p.? 7:;'::;:&amp;::...11
Richard M. Hopper
Wildlife Researcher

_

�October

- 81 -

JOB PROGRESS

State

Work

REPORT

COLORADO

of
No.

W-88-R-15

Migratory

Plan No.

2

Job No.

Project

Job Title
Period

1970

Covered:

Personnel:

Improving
April

Experimental
Studies
Status of Canada Goose

1, 1969 to March

Bird Investigations

2
on
Populations

31, 1970

C. Hayes and D. Horn, Bureau of Sport Fisheries and
Wildlife; W. Carpenter, C. Crawford, G. Crawford, G.
East, H. Funk, R. Hopper, J. Monarch, D. Owens, C.
Slonaker, G. Steele, C. Wetherill, G. Will and M.
Szymczak, Colorado Division Game, Fish and Parks.

ABSTRACT

A total of 475 goslings were released in transplant areas in June and July
of 1969.
Sixty-nine goslings were released at the K-4 Ranch near Masters;
160 at San Luis Lakes in the San Luis Valley; 193 at Lake John Annex in
North Park and; 53 on the Colorado River near Lorna. Band recoveries indicate that birds released at the K-4 Ranch tended to remain in Colorado.
Approximately
10 percent of the geese released at the San Luis Lakes were
recovered the first year with Chaves County, New Mexico and the state of
Sonora in Mexico being the major recovery areas.
Goslings released in
North Park moved throughout the west and central portions of the Park
throughout late summer and early fall but showed tendencies to return to
the release site.
It was estimated thatmajor movement of North Park
goslings along migration paths began sometime between November 1 and
November 20. Approximately
11 percent of North Park goslings were recovered by hunters.
Recoveries indicate that North Park goslings migrated
along a Pacific Flyway route with the major portion of the birds wintering
on the Lower Colorado River in Yuma County, Arizona.
At least six nests
were established
in the transplant area along the Colorado River in midApril of 1969, two years after the original plant was made.

��- 83 -

EXPERIMENTAL STUDIES ON IMPROVING STATUS OF
CANADA GOOSE POPULATIONS
Michael

R. Szymczak

P. S. OBJECTIVE
To develop a technique for establishing a breeding population of Canada
geese in suitable habitat where they do not currently exist in Colorado.

SEGMENT OBJECTIVES
1.

Expand the breeding flocks of Canada geese in the South Platte Valley,
the San Luis Valley, and North Park.

2.

Continue efforts toward establishing the breeding
Canada geese in the Colorado River Valley.

METHODS

flock of Great Basin

AND MATERIALS

Canada goose goslings for transplant purposes were obtained from four different sources:
(1) goslings and some eggs from the metropolitan Denver area;
(2) goslings and some eggs from the Fort Collins area; (3) goslings from Valmont
Reservoir in the Boulder area and; (4) eggs from nest in danger of destruction
from high water along the Yampa and Little Snake Rivers in Moffat County.
All eggs were transported
to the Fort Collins Wildlife Research Station for
incubation and subsequent ralslng.
All goslings live-trapped were transported
as soon as possible to the release site. Birds obtained from eggs collected
in Moffat County were released along the Colorado River near Lorna. Birds
collected at other locations were released either at San Luis Lakes, Lake John
Annex or the K-4 Ranch.
Migration and mortality information were obtained
from the Migratory Bird Population Station.

from recovery

cards received

RESULTS AND DISCUSSION
South Platte Valley - K-4 Ranch

Sixty-nine goslings were released along the South Platte River on the K-4
Ranch near Masters, Colorado in July of 1968. The previous summer, 217 goslings
has been released on the same area.
According to visual sightings, there have been geese in the general vicinity
since the first gosling release.
The area has been closed to hunting.
The
local birds have been joined by migrant geese, considered part of the Hi-Line
Population, during the fall and winter period.
The local transplants utilized

�- 84 -

both Empire and Riverside Reservoirs which are near the transplant site.
As many as 1,000 geese have been observed in the area during the fall
and winter period.
According to hunting season band recoveries there has been some movement
of birds from the transplant site, but limited dispersion of the birds
from Colorado (Table 1). Less than five percent of the released goslings
have been recovered by hunters.
Only two of fourteen recoveries were
taken out-of-state.
One bird each was taken in the states of Kansas and
Oklahoma.
In addition, four birds released in 1968 were captured in July,
1969 while flightless in northwestern South Dakota.
All of the afore-mentioned states are considered outside of the range of the Hi-Line Population.
The South Dakota recoveries may have been the result of a "moulting
migration" by the geese in question from Colorado.
Pole-type nesting structures have been erected at the release site by G. I.
Crawford under Federal Aid Development Project W-llO-D.
The first signs of
nesting attempts were noted in the area in early spring of 1970.

Table l. Hunting recovery locations of Canada geese released
Ranch near Masters, Colorado in Weld County.

Year
No. Released
Area Recovered

1968
217
No. Recovered
Direct
Indirect

on the K-4

1969
69
No. Recovered
Direct
Indirect

Colorado
Morgan

Co.

S

0

1

N.A.

Weld Co.

0

2

1

N.A.

Adams Co.

1

2

0

N.A.

Kansas

0

1

0

N.A.

Oklahoma

0

1

0

N.A.

6

6

2

N.A.

Total

San Luis Valley - San Luis Lakes
In July of 1969, 160 goslings were released
east side of the San Luis Valley in Alamosa

at San Luis Lakes, located on the
County.
This was the first release

�- 85 -

in the San Luis Lakes area. Other releases on the Monte Vista and Alamosa
National Wildlife Refuge in 1967 and 1968 served to increase the size of
the local flock in the San Luis Valley.
Although no specific numbers of
total breeding pairs and production in the Valley are available, there
are indications that this population can support a limited harvest at this
time.
Transplants in the San Luis Lakes area were initiated in order to stimulate
establishment of a breeding population in the vicinity of San Luis and Head
Lakes.
The majority of the birds remained on San Luis Lakes during late
summer and early fall. Mortality during that period was considered minimal.

Band Recoveries
The location of band recoveries is presented in Table 2. Approximately 10
percent of the goslings released at San Luis were recovered the first year.
There was a definite variation in the distribution of recoveries of the birds
released on different dates.
One bird from each group was recovered in the
San Luis Valley.
Eight of the birds released on July 8 were taken near the
Bitter Lake National Wildlife Refuge near Roswell, New Mexico between
December 18, 1969 and January 5, 1970. Six of the July 22 birds were taken
in Mexico.
Five of the birds were taken in the Empalme Valley near Guaymas
on the Gulf of California in the state of Sonora and one at an unknown location
in the inland state of Chihuahua.
In 1967, a San Luis Valley transplant of
that year was also taken near Guaymas.
Three of the five birds taken in the
Empalme Valley were reportedly caught by hand on December 17, 1969. The
other two birds were taken in January and March indicating the birds may have
terminated their migration in that area.
The migration and/or recovery patterns of the 1969 San Luis Lake birds was
decidedly different than goslings released in the Valley the previous two
years.
Recoveries indicate that many of the 1967 and 1968 birds remained in
the San Luis Valley during the fall and winter period.
Most hunting recoveries from outside the Valley came from the Bosque del Apache National
Wildlife Refuge along the Rio Grande River in central New Mexico.
At least
five of the goslings released in 1967 or 1968 were recovered in Denver, their
natal area.
Four of the five were recaptured in the summer of 1969.

Spring Returns
Canada geese began returning to San Luis Lakes in early spring of 1970. As
many as 80 birds have been observed there during the spring and early summer.
A good number of these geese were probably transplants of the previous year,
but since none of these birds were color-marked, it is impossible to determine returnees through visual observation.

�- 86 -

Table 2. First year hunting recovery locations of Canada geese released
in 1969 at San Luis Lake, Alamosa County, San Luis Valley, Colorado.
Date Released
No. Released

July 8
80

July 22
80

Area Recovered

No. Recovered

No. Recovered

United States
Colorado
Costilla

Co.

o

1

Conejos

Co.

1

o

Chaves .Co.

8

o

o

1

o

5

o

1

9

8

New Mexico

San Miguel

Co.

Mexico
Sonora
Southwest

Chihuahua
Unknown

Total

North Park - Lake John Annex
In June and July of 1969 a total of 194 Canada goose goslings were released at
Lake John Annex, approximately eight miles west and 2 miles north of Walden,
Colorado in North Park. This was the initial release in an extensive program
which hopefully will establish a breeding population of Canada geese in this
high mountain park.

�- 87 -

There has been an extremely small, remnant population of Canada's in
North Park.
Birds have been observed through the summer months at both
McFarlane Reservoir and Lake John Annex.
At least one nest has been
found and photographed at the "Annex".
The population has not grown or
expanded.
Lake John Annex was picked as the release site for three reasons: (1) an
adult pair of the remnant population was known to be using the "Annex";
(2) excellent meadows for grazing were adjacent and, (3) most of the
adjacent land and water is controlled by the Division of Game, Fish and
Parks.
It was hoped that the young goslings
would associate with the
adult pairs through the fall and winter period and if possible, return
to North Park with them in the spring.

Local Dispersion
The goslings were released in two groups: 60 on June 25 and 134 on July 16.
Fifty-eight of the original 60 released carried green/white/green
neckbands.
Ninety-seven of the birds released on July 16 carried green neckbands with
white numerals 754 through 850. One of the birds in the June 25 plant died
before it was released from the holding pen. A chronology of events and
subsequent sightings in the vicinity of North Park are presented below.
The locations of sightings are coded and plotted in Figure 1.

Sightings
June 27
July 16
July 17
July 28

Aug. 11
Aug. 17
Aug. 21
Aug. 25
Aug. 27
Sept. 1

CD

59 birds released from pen
134 birds released
30 birds from the original release observed;
all of July 16 release present
25 birds sighted on the Bighorn Ranch
two
without neckbands of which one had been removed by
rancher.
28 geese observed at Boettcher Lake (]) (all banded)
19 at N. Delaney Butte
(all banded)
Feeders at Lake John Annex
closed
11 at Walden Lake
64 at Big Creek Lake
(all bandgd )
Geese observed at McFarlane Res. ~,
Hebron Sloughs (j), Case Flats
and Peterson
Pond
but no numbers noted.
Unknown number at (I)
14 banded birds at Cowdrey Lake
andG) on the N.
Platte River @
11 at Walden Lake @
Numerous birds at Lake John
87 birds on Lake John and Lake John Annex
Some observed on Walden Lake
16 collared birds at Lake John
2 collared birds at Lake John
2 collared birds at Cowdrey Lake

CD

CD

G)

®

CD

G)

®

®

Sept. 7
Sept. 16
Oct.
Oct.
Oct.
Oct.
Nov.
Dec.
Dec.

3
11
23
27
22
9
10

QJ;

@

CD

CD

Q

G)

CD

Q~

�z

o
...•
-n
o

3~·
Delane y 1./
Butte Ls. ~

OE
.

S.

8
&lt;::1 OA

Case &lt;!It'v
Flats

Peterson
Pond

9
Lake John or Lake John
1
Annex
Boettcher Lake
2
N. Delaney Butte Lake
3
Walden Lake
4
Big Creek Lake
5
McFarlane Reservoir
6
Hebron Sloughs
7
Case Flats
8
Peterson Pond
9
Cowdrey Lake
10
N. Platte R.
11, 12, 13

7

Hebron 0".
SloughS:
Me Farlone
Res.
SCALE

0

Fig. 1. Locations of Canada goose sightings
and winter of 1969.

2

4 Mi.

in North Park, sununer, fall

�- 89 -

The aforementioned sightings indicate that the birds dispersed from the
"Annex" and moved throughout the west and central portions of North Park.
However, they showed a tendency to return to the release site throughout
the summer, but particularly later in the fall.
Five geese were observed on McFarlane Reservoir on July 15. None of the
birds were neckbanded.
Two of the five were captured and both were adults.
The two birds captured were found to be not banded when captured.

Band Recoveries

and Sightings

Dispersion and Migration Chronology--Hunting
season recoveries and neckbanded sightings give indications of dispersion and migration chronology.
The location of band recoveries by date are presented in Table 3. Band
recoveries by time interval in any particular area are entirely dependent
on both hunting season dates and hunting pressure.
In other words, if
the season is not open or there are no hunters in an area when banded
birds are available, no recoveries can be expected.
Mid-October recoveries of North Park birds indicate some degree of dispersion in early fall. On October 18, geese were taken in Moffat County,
Colorado and near El Paso, Texas.
Approximately 85 of the 193 geese
released were known to still be in North Park at that time. One bird
was recovered near Prescott, Arizona in late October.
The location and
recovery dates of the October Texas and Arizona recoveries indicate that
these birds were probably wanderers and were not accompanied by a significant number of their North Park cohorts.
No birds were recovered during the November 1 through November 20 period,
however, the pattern of recoveries after that date (Table 3) in addition
to visual sightings indicate that major movement from the Colorado-Wyoming
area including North Park took place during that period.
On November 18, 30 geese were sighted on Mountain Home Reservoir in the
San Luis Valley.
All the birds were wearing green collars, at least some
of which were described as being green-white-green
in design, indicating
North Park birds released on June 25, 1969. Then, during the last ten days
of November, North Park birds were reported taken in Baca County, Colorado,
Green River, Utah and at Martinez Lake, Arizona.
After December 1, all
recoveries came from Arizona or Mexico.

Migration Routes and/or Harvest Areas--Twenty-two
or approximately 11 percent of the goslings released in North Park were recovered by hunters.
The location of recoveries recorded by release date are presented in
Table 4.

�Table 3. 1969-70 hunting season recoveries of North Park birds plotted according to time and approximate
distance from release site.

Area Recovered

October
1-10
11-20

21-31

Period of Recovery
November
December
1-10
11-20
21-30
1-10 11-20

21-31

1-10

January
11-20

21-31

United States
Colorado
Jackson Co.
Moffat Co.

2

Wyoming
Carbon Co.

1

1

1

Utah
Emery Co.

1

Colorado
Baca Co.

1

\0

Arizona
Yavapai Co.
Cila Co.
Texas
E1 Paso Co.

o

1
1

1

Arizona
Yuma Co.

2

4

Mexico
Baja

1

.Y

Coahuita
Sinaloa
1/ Recovered during the month of December; date not known.

1
2

1 1/

1

�- 91 -

Table 4. First year hunting recovery locations of Canada geese released
in 1969 at Lake John Annex, Jackson County, Colorado.
Date Released
No. Released

June 25
59

Area Recovered

No. Recovered

United

July 16
134

No. Recovered

States

Wyoming
Carbon Co.
Colorado
Jackson Co.
Moffat Co.
Baca Co.

2

1
2

1

Utah
Emery Co.

1

Arizona
Yavapai Co.
Gila Co.
Yuma Co.

1
4

1
2

Texas
El Paso Co.

1

Mexico
Baja
Northwest
Unknown

1

1

Coahuila
West-central

2

Sinaloa
Northwest

1

1

Total

6

16

�- 92 -

The location of recoveries indicate that the majority of North Park geese
crossed over the Continental Divide and migrated down the Pacific Flyway.
Seven of the 22 birds were recovered in the Central Flyway but three of
those seven were recovered in North Park or adjacent areas in Wyoming.
The Pacific Flyway-oriented birds apparently followed the traditional
route of some populations of Great Basin Canada geese, terminating migration in Arizona along the lower Colorado River.
Band recoveries on the
Imperial National Wildlife Refuge near Yuma, Arizona indicate that North
Park birds were wintering with birds banded at Wheatland Reservoir, a
moulting area for Great Basin geese in eastern Wyoming.
The first recovery of a North Park bird at the Imperial Refuge area was taken on
November 28 and the last approximately one month later indicating that a
number of birds may have terminated migration in that area. The two
birds taken near Las Mochis in the coastal state of Sinaloa could have
taken either the Pacific or Central Flyway route to their recovery location.
Only four birds were recovered in the Central Flyway, away from the release
location:
One early recovery near El Paso, Texas which was previously
mentioned; one in Baca County in southeast Colorado; and two in the Mexican
state of Chihauhaua somewhere near Babicora.
In addition to the one recovery in Baca County, at least four other neck-banded birds were known to
have remained at Turk's Pond in that same County throughout the winter.

Colorado

River

On July 8, 1969, 53 Great Basin Canada goose goslings were released on the
lower Colorado River near Lorna. These birds were obtained from eggs which
were collected in April from nests in danger of destruction from high water
along the Yampa River in Moffat County, Colorado.
The eggs were artifically
incubated and the birds raised at the Fort Collins Wildlife Research Station.
The birds when released wore green neck collars with two solid white dots.
The 1969 release was the third annual transplant in that area and brings the
total number of birds released there to 127.

Spring Reconnaissance
In mid-April of 1969 a trip was made by boat from Fruita downstream on the
Colorado River to the Utah border to get some idea of the progress of our
transplant program.
Approximately 50 geese were observed on the trip. The
majority were encountered near the transplant site. Only three green neckbanded birds were observed.
Although white markings were identifiable on
the neckbands, the exact design was not discernable.
The original 25 goslings released in the area were only two years old in the
spring of 1969 and would not be expected to begin nesting.
However, at least
six nests were established along this portion of the river, with four of these
on the island which served as a release site. Two-year old birds were apparently
capable of reproduction but normally do not nest until their third year. These
nests were probably produced by older migrant birds which wintered in the
area and remained to nest. The four nests near the release site possibly indicate the geese were reluctant to leave the area in which they were fed
artifically.

�- 93 -

Band Recoveries

and Sightings

According to band recoveries only three of 127 geese released have been
harvested by hunters.
All three of these birds were released in July
1968 and recovered during the 1969 hunting season.
Two birds were taken
near Glasgow in eastern Montana and one near Fort Collins, Colorado.
Both
harvest locations are considered to be within the Hi-Line Population area,
and outside the normal range of Great Basin goose population, west of the
Continental Divide.
In addition, at least ten birds of the 1968 plant
were sighted at College Lake near Fort Collins in January of 1970. The
lack of other recoveries and sightings in addition to the previously mentioned observations indicate that the transplanted birds are not moving
south, but staying in the transplant area, which is closed to goose hunting.
A portion of these birds have apparently accompanied migrant geese that
winter in the transplant area back north in the spring and then drifted into
the Hi-Line area.

Winter Populations
The transplant program along with the associated goose hunting closure has
apparently stimulated a small build-up in wintering numbers.
In January
of 1967, 71 geese were counted along the Colorado River from Delta to the
Utah border.
In January 1970, 335 geese were counted in the same area.

Prepared

by

~~72

.1A

Michael R. Szymczak
~~
Asst. Wildlife Researcher

J

��October

- 95 -

JOB PROGRESS

State

Work

REPORT

COLORADO

of
No.

W-88-R-15

Migratory

Plan No.

2

Job No.

Project

1970

Job Title

Arkansas

Valley

Period

November

22, 1969 to March

Covered:

Personnel:

Goose

Bird

Investigations

Flock Management

5
Studies

31, 1970

Velma Fredrickson,
Charlene Osborne, Ann Leck1er, Don
Bogart, John Casper, Rusty Forbes, Mark Frasier, Bernie
Goetze, John Gonzales, Larry Green, Jack Grieb, Mike
Hickey, Al Heins, Ray Johnson, Bob Kitzmiller, Rober
Lowry, Don Minnich, Horace Moorhead, Kris Moser, Gene
Nugent, Fred Roth, Wayne Russell, Bert Widhalm, Hank
Wilson, and Mike Szymczak

ABSTRACT

Geese were subject only to sunrise to 12:00 o·c1ock noon hunting throughout
the 1969-70 season.
Water levels on all reservoirs in southeast Colorado
were very high throughout the fall and winter period.
Food conditions were
excellent with temperatures above normal and precipitation
below normal.
Geese began arriving on November 11 and by November 21, approximately
57,000
geese were in the area.
Goose numbers in southeast Colorado varied from
55,000 to 65,000 throughout the winter.
The mid-winter
inventory of the
entire Short Grass Prairie Population totaled 147,414 birds, about 35,000
more than the previous years total.
Observation of goose flight patterns
throughout the season resulted in these conclusions:
(1) Without hunting
pressure, geese would fly twice a day despite changes in moon phases; (2)
The cessation of morning flights was closely correlated to light-of-moon
phases during the hunting season in the months of November and December;
(3) With firing lines operating, geese showed a more definite response to
1ight-of-moon phases, by discontinuing
morning flights, than under nonfiring line situations;
(4) Inclement weather stimulated morning flights
even during 1ight-of-moon periods.
Cold weather seemed to delay morning
flights even during dark-of-moon periods; (5) There were some indications,
particularly
toward the end of the season, that geese responded to morning
hunting pressure by delaying feeding flights until afternoon.
The harvest
in southeast Colorado was composed of a higher percent of immatures than
the birds taken earlier in the year on the Canadian staging areas.
An
estimated total of 10,597 geese were bagged by 5,327 hunters during the
1969-70 season in southeast Colorado.
At the Two Buttes Management Area
659 hunters bagged a total of 248 geese with 703 hunters harvesting 512
geese at the Lamar-Eads Management Area.
Bands were placed on 599 geese
post-season
in southeast Colorado.
Approximately
50 percent of the 1968-69
hunting season band recoveries of birds banded in southeast Colorado were
reported taken in Canada.
The annual mortality rate of all age classes
combined was calculated to be about 26 percent.

�- 96 -

RECOMMENDATIONS

1.

Continue observations of flight patterns of Canada geese to determine
what effect half-day hunting regulations have on their behavior.

2.

Record approximate time of day of harvest of birds from which tail
fans are collected in an effort to determine whether Canada goose flights
are age-specific in reference to time.

�- 97 -

ARKANSAS

VALLEY CANADA GOOSE FLOCK MANAGEMENT
Michael

STUDIES

R. Szymczak

Current emphasis in the Arkansas Valley study has been on the evaluation
of new approaches to goose management in southeast Colorado.
In recent
years, southeast Colorado populations have been characterized by an initial
pre-season population build-up followed by dwindling numbers as the season
progressed.
During the 1969-70 hunting season Colorado officials elected
to initiate a morning only hunting season in southeast Colorado in an
attempt to encourage geese to remain in the state throughout the winter
period.
A portion of this report will deal with the evaluation of that
regulation.

P. S. OBJECTIVE
To collect management information on the wintering
Canada Goose Population in southeast Colorado.

Short Grass Prairie

SEGMENT OBJECTIVES
1.

To determine the time of movement of geese into the Arkansas
and the size and distribution of the wintering flock.

Valley

2.

To obtain an estimate
flock.

of production

of the

3.

To determine
loss.

pressure

4.

To determine the annual mortality
pattern of the flock.

5.

To develop and evaluate
Colorado.

hunting

and the age composition

on the flock and the associated

and the migration

new approaches

METHODS

wounding

and/or harvest

to goose management

in southeast

AND MATERIALS

Periodic aerial counts of Canada geese (Branta canadensis parvipes) were made
in southeast Colorado.
The December 9 and January 7 counts were coordinated
with both ground and aerial counts made by various State and Federal personnel
throughout the wintering grounds of the Short Grass Prairie Canada Goose
Population.
These counts were made for the purpose of determining the size
and distribution of the wintering goose population.
Check stations were
operated at the Two Buttes Management Area and the Lamar-Eads Management Area,
which is an area for decoy hunting only.
Information on hunting pressure,
harvest and wounding loss was collected. Tail fans from bagged geese were

�- 98 -

collected at all check stations in order to determine age ratios in the
kill.
The weights of birds bagged were also recorded.
Goose feeding flights were observed periodically in the Eads Lakes area
and daily in the Two Buttes area.
Information on time of flight, numbers
of birds and general behavior of birds were recorded.
Geese were trapped and banded after the close of the hunting season to
obtain information on age ratios, mortality rates and migration patterns.
The effects of past hunting season regulations were evaluated and recommendations were made for the 1970-71 season.

RESULTS AND DISCUSSION

Water,

Food and Weather

Conditions

The water levels on all reservoirs, which serve as goose resting areas,
were very high with many near capacity throughout the 1969-70 season (Table
1). The high water virtually eliminated the open mud flats and bars normally utilized by the geese.
Above normal precipitation throughout most
of the area helped fill many of the reservoirs in addition to eliminating
the necessity for heavy use of stored irrigation waters.
All selected
stations recorded below average precipitation for the months of January,
February, November and December of 1969. Near or above normal levels were
recorded for the remaining months of 1969.
Food conditions throughout southeast Colorado were judged excellent for the
maintenance of Canada geese.
Above average rains through the spring and
early summer months of 1969 helped produce excellent crops of dryland milo.
Fall moisture insured the establishment of winter wheat stands in the same
dryland areas.
Heavy October snows rendered some milo fields unharvestable,
making the entire yield available for goose use.
A summary of weather conditions throughout the hunting season is presented
in Table 2. In general, temperatures were above normal and precipitation
below normal.
These factors combined to produce generally poor hunting
throughout the course of the season.
Occasional ground fog and wind provided the best opportunity for goose harvest during the season.

Migrational

Movements

and Distribution

of the Wintering

Population

Large numbers of Canada geese began arrlvlng in southeast Colorado about
November 11 in 1969. Approximately 57,000 geese were counted during the
first aerial survey on November 21. The results of all aerial surveys presented in Table 3 indicate that southeast Colorado retained between 55,000
and 65,000 geese during the hunting season.

�- 99 Table 1. Yearly precipitation totals for selected southeast Colorado weather
stations for 1969 and the water status of associated reservoirs.
Station Location
and
Associated Reservoirs
Rocky Ford

1969 PreciEitation (In. )
Departure
Total
from Norm

12.13

Near
Near
Near
Near
Near
17.04

Horsecreek
Blue Lake

John Martin Reservoir

22.72

Reservoir

Nee Granda
Flamingo Lake
Nee Noshe
Upper Queens Reservoir

Thurston

19.49

Two Buttes Reservoir

23.09

open
open
open
open
open

bars
bars
bars
bars
bars

Some open, bars
available
No open bars

Higher than
1968-69
Water at
capacity
Very small pool
Higher than
1968-69
Near capacity

Open shoreline
available; no bars
No open bars

No open bars

Shoreline open
No open bars
No open bars; heavy
vegetation on south bar
No open bars
No open bars

+5.29

Reservoir

SEringfield

No
No
No
No
No

Higher than
1968-69
Higher than
1968-69
Much higher
than 1968-69

Higher than
1968-69
Near 1968-69

Lower Queens Reservoir
Lamar

capacity
capacity
capacity
capacity
capacity

+8.94

Swede Lake
Sweetwater

Areas

+4.79

Reservoir

Eads

Resting

-,

-0.18

Dye Reservoir
Holbrook Reservoir
Cheraw Reservoir
Meredith Reservoir
Henry Lake
Las Animas

Water Level

Near capacity

No open bars

Much higher
than 1968-69

Open bars available

+8.36

�- 100 Table 2. Summary from selected southeast Colorado weather stations, November
1969 through January 1970.

Location

Temperature
Average
Departure from
Maximum Minimum
Normal Daily Ave.

Precipitation
..Departure
Monthly TotaL~.:/fromNorm

November
Rocky Ford

59.5

21.1

0.6

0.30(3)

0.16

Las Animas

59.4

24.7

1.5

0.28(1)

-0.20

Eads

57.7

25.6

0.06(1)

-0.43

56.5

26.7

0.27(2)

Rocky Ford

49.2

16.4

0.3

0.26(3)

-0.06

Las Animas

49.3

19.7

1.0

0.29(2)

-0.06

Eads

48.5

20.1

0.12 (3)

-0.18

Lamar

47.4

20.5

0.16 (3)

-0.18

Springfield

48.1

21.5

Rocky Ford

49.0

9.7

-0.6

0.12(1)

-0.31

Las Animas

48.6

13.7

-0.7

T (0)

-0.48

Eads

45.2

15.0

T (0)

-0.39

Lamar

49.0

15.3

T (0)

-0.50

Springfield

47.5

15.3

Lamar
Springfield
December

1.2

0.18(6)

January

2.3

0.01(1)

1/ Number of days of measurable precipitation in parentheses.

�- 101 -

Table 3. Results of Canada goose aerial surveys, southeast Colorado, 1969-70.

Nov. 21

Dec. 9

Date
Dec. 19

Jan. 7

Jan. 16

o

o

5

379

o

o

o

o

15

25

o

o

30

o

200

Meredith Reservoir

7,600

3,900

1,075

2,650

845

Holbrook Reservoir

o

100

400

600

1,200

Cheraw Reservoir

o

o

20

o

o

Horsecreek Reservoir

5,000

1,700

o

o

4

Blue Lake

2,200

4,000

o

o

o

201

1,200

1,000

o

o

6,800

1,900

o

o

55

o

1,900

400

o

o

Nee Noshe Reservoir

7,545

19,000

6,500

4,000

11,000

Upper Queens Reservoir

3,500

4,600

12,000

9,000

16,000

o

o

50

o

o

Two Buttes Reservoir

12,000

9,000

4,500

6,000

17,000

Turk's Pond

7,000

12,000

13 ,500

27,800

14,000

John Martin Reservoir

4,000

400

16,000

12,000

4,430

Bonny Reservoir

1,000

1,000

500

1,000

o

Total

56,846

60,700

55,980

63,444

64,759

Area
King Barret Sloughs
Arkansas River
King Barret-Dye Reservoir
Dye Reservoir

Eads Group:
Swede Lake
Sweetwater Reservoir
Flamingo Lake

Thurston

�- 102 -

The distribution of the birds within southeast Colorado remained essentially
the same as during the 1968-69 season.
Additional numbers during the early
part of the season, particularly in the Horsecreek-Blue Lake area, provided
increased hunting opportunities.
By mid-December most of the birds were
located in the John Martin, Eads Lakes, or Two Buttes-Turks area. Two Buttes
Reservoir, which in recent years has lost most of its goose population to
Turk's Pond shortly after opening day, retained huntable numbers throughout
most of the season.
The 1969-70 southeast Colorado goose population represents the largest
number of geese ever present in that area through the hunting season.
Comparative yearly figures are represented by the results of January inventories
recorded in Table 4. In recent years Colorado populations have been characterized by an initial pre-season build-up followed by dwindling numbers as the
season progressed.
In the past the birds have apparently moved south into the
Texas Panhandle area producing a gradual yearly increase in birds in that area
until the number of birds recorded during the January Inventory in the Buffalo
Lakes, Texas area exceeded the Colorado inventory (Fig. 1). During the 196970 season Colorado did not lose appreciable numbers to Texas, even though the
panhandle population also reached an all time high (Fig. 1). The increase in
the panhandle area may be the result of a continuing shift of the "Eastern
Segment" (Grieb 1970) of the Short Grass Prairie Population to panhandle
wintering areas.

Table 4. January
1948-1970.

Inventory

of Canada geese, Arkansas

Valley,

Colorado,

Year

Goose Count

Year

Goose Count

Year

Goose Count

1948

4,798

1955

25,110

1962

40,250

1949

12,286

1956

24,212

1963

35,889

1950

l3,170

1957

24,617

1964

33,750

1951

19,320

1958

35,894

1965

37,693

1952

30,463

1959

44,660

1966

38,635

1953

20,236

1960

37,394

1967

29,835

1954

20,280

1961

31,360

1968

42,682

1969

29,201

1970

63,444

1/ Inventory

of February

7, 1962 substituted

2/ Inventory

of February

15, 1966 substituted

for January,

1/

]j

1962 inventory.
for January, 1966 inventory.

�JANUARY INVENTORY

150,000

125,000

100,000

~
o

Vol

75,000
I

/
I

50,000
,...

,...
&lt;,

,...
./

,...

.,.,

...••.

.•....•

..... .•....•

•.•.•..

~

- - - ---

»&lt;

Colo-cado segment- - ---...........

_--

,/
,/
,/

&lt;,

Lakes se~ent

&lt;,

.,/

........,.,

'-

1

" '-

I
"

,...

"

1

1

25,000

62

61
Fig. 1.

63

64

65

66

67

68

69

Comparison of the Colorado and Buffalo Lakes (Texas) Segments of the Short Grass Prairie Canada goose

_ ....._ ••.1 ...••..~ __

70

�- 104 -

The results of coordinated aerial surveys conducted on the Short Grass
Prairie Canada goose wintering range indicate the population size is continuing to increase (Table 5).

Table 5. Post-hunting season status of Short Grass Prairie
population, 1960-70, data generally from regular mid-winter

Canada goose
inventories.

Year

Year

Number of Birds

Number

of Birds

1960

77,709

1965

103,435

1961

103,355

1966

1l0,485

1962

80,l33

1967

111,452

1963

93,940

1968

127,903

1964

81,221

1969

112,399

1970

147,414

!/ Inventory

of February

15, 1966, substituted

for unsatisfactory

1/

January

Inventory.

The Colorado population accounted for the major portion of the increase
(Table 6). Again, as during the previous year, the December Inventory
totaled fewer birds than the January count.
Coordinated counts were conducted in the second week of December and during the January Inventory.

Table 6. Results of Short Grass Prairie
hunting season, 1968 and 1969.

Canada goose coordinated

inventories,

State

December
1968

1969

1969

1970

Nebraska

1l,740

10,512

7,000

8,140

Colorado

28,633

60,700

29,201

63,444

Oklahoma

3,000

1,300

3,500

1,500

New Mexico

9,195

7,531

7,848

10,640

Texas

20,100

26,387

64,850

63,690

Totals

72 ,668

106,930

112,399

147,414

January

�- 105 -

Analysis

of Flight Pattern

Data

Eads Lakes
Pre-season flight pattern data indicate that geese were making two definite
flights daily.
Flights were generally low and most feeding locations were
not more than one mile from the reservoirs.
Geese congregated in specific
fields despite variation in time of departure.
It was noted that with
moderate winds (15-25 mph) , there was frequent interchange between feeding
locations and reservoirs throughout the day. There were no indications of
a tendency toward late afternoon or evening flights even though the light
of the moon was probably great enough to permit night feeding.
After November 22, the opening of the hunting season, morning flights were
negligible as there was a full moon on November 23 which permitted night
feeding (Table 7). Many flocks were observed returning to reservoirs the
morning of the 23rd, either from night feeding or from other reservoirs.
The afternoon flight on the 23rd began at approximately 3:45 p.m., 1:15
later than the afternoon flight of November 21, and continued into dark.
Flights were high and feeding locations were estimated to be five to six
miles from the reservoir.
The pattern of late afternoon-evening
flights
continued until November 29 when two-a-day flights were again initiated.
During the December 9 through 17 period, two-a-day flights were common.
Morning flights were generally sporadic and variable in direction.
Direction
of flights seemed to be dependent on location of firing-line hunters.

Table 7. Chronology of moon phase during the 1969-70
season in southeast Colorado.

Canada goose hunting

Month

Day

Moon Phase

November

9
16
23

New Moon
First Quarter
Full Moon

December

2
9
16
23
31

Last Quarter
New Moon
First Quarter
Full Moon
Last Quarter

January

7
14
22

New Moon
First Quarter
Full Moon

�- 106 -

Flocks would "probe" different areas until they found unmolested routes
and then following flocks would use these corridors until hunters moved
in. Flights were high and most feeding locations were more than 10 miles
from reservoirs.
Beginning December 15, morning flights were not complete with many birds remaining on the reservoir.
Afternoon flights
during the December 9 through 17 period were more direct and continuous.
Once a flight direction was established, the direction was essentially
maintained throughout the flight.
Flights were low and feeding locations,
for the most part, were within one mile of the reservoir.
There was no
morning flight on December 18. Morning flights did not begin again until
December 27 and then continued only through January 2. After January 2,
and through most of the remainder of the season, the geese flew only in
the afternoon.
During this period there was some variation in activity
of the geese during the morning.
Occasionally small flocks came into
the reservoir.
A few flocks did attempt to leave, but most returned to
reservoirs after being fired upon. On January 7 and 8, during the extreme
dark phase of the moon, birds were noted to be extremely restless and
feeding flights began between 11:45 and 12:00 noon. Afternoon flights
between January 6 and 11 were direct and short as most feeding locations
were within one mile of the reservoir.
Most birds returned to the reservoir before dark.

Two Buttes
Regular two-a-day flights were common during the pre-season period.
Unlike
the Eads area, two-a-day flights continued after the opening of the regular
season on November 22. However, afternoon flights gradually began later
and night flying during the period of November 22 through November 27 was
common.
Regular morning and afternoon flights began again on November 28.
On December 3 geese began using the millet field below the Two Buttes cabin.
On December 4 and 5 the geese were active throughout the day as inclement
weather set in. Birds continued extensive use of the millet field below
the cabin through December 6, the first day of the Two Buttes season. Most
flocks that attempted to leave the reservoir in the morning of the 6th
turned back at the firing line. A definite flight occurred in the afternoon.
Feeding flights on the 7th began about 10:00 a.m. and became more intensive
about 11:30 a.m. Many flights skirted the firing line by going easterly.
Generally, through the first week of the season flights began late in the
morning and continued into the aftern06n creating one continuous flight
daily.
Flights on December 10 were an exception, beginning about 7:30 a.m.,
apparently stimulated by the foggy, overcast conditions.
Through the first
part of the second week of the Two Buttes season, morning flights continued,
but they were generally not as heavy as the afternoon flights.
Toward the
end of the week (December 18), morning flights became extremely light.
Afternoon flights remained heavy and there were indications that the birds
were feeding at night.
Late afternoon and night flights were common through
December 28. The morning flights were extremely light with some variation
which seemed to be caused by weather conditions.
Increased movement was
noted on the morning of December 22, which had 30 to 40 mph winds, and on
December 27 when it snowed.
On December 29 geese began moving off the
reservoir at 11:00 a.m., but most turned back at the firing line. From

�- 107 -

December 30 through January 10 there was essentially no morning flight
from the reservoir.
There were large afternoon flights normally beginning
around 3:00 p.m. On January 9 and 10 some large incoming flights were noted.
From January 11 through the end of the season there were large morning
flights generally occurring during the 7:00 a.m. to 9:00 a.m. time period.
Geese using Meredith and John Martin Reservoirs followed the same general
pattern as those in the Eads Lakes area with some variation caused by local
weather conditions.
The only significant variation between flight pattern of birds on northern
areas and those at Two Buttes occurred at the beginning of the regular southeast season on November 22. Birds at Two Buttes did not respond to the season opening by going to late afternoon-night flying as quickly as those in
the northern areas.
The variation was probably caused by firing line
pressure; heavy in the north, none at Two Buttes.
The flight pattern information collected seem to indicate the following.
1.

Without hunting pressure,
changes in moon phases.

geese would fly twice a day despite

2.

The cessation of morning flights was closely correlated to lightof-moon phases during the hunting season in the months of November
and December.

3.

With firing lines operating, geese showed a more definite response
to light-of-moon phases, by discontinuing morning flights, than
under non-firing line situations.

4.

Inclement weather stimulated morning flights even during light-ofmoon periods.
Cold weather seemed to delay morning flights even
during dark-of-the-moon periods.

5.

There were some indications, particularly toward the end of the
season, that geese responded to morning hunting pressure by delaying
feeding flights until afternoon.

Age Composition
To obtain an estimate of the age composition of the harvest, tail fans from
birds bagged were collected.
Collection of tail fans on the two state-controlled hunting areas resulted in 597 usable fans. The age data, categorized
by method of hunting, indicates the percent of the harvest composed of
immatures increased from 1968-69 levels on both management areas (Table 8).
As during the previous year, immatures in 1970-71 were taken at a greater rate
in the decoy area than on the firing line.

�- 108 -

Table 8. Age composition of the goose harvest,
controlled hunting areas in southeast Colorado.

Year

Two Buttes
Adult
No.
%

1951-52

356

1957-58

by year, at two state

Check Station
(Firing Line)
Lamar-Eads
Young
Adult
No.
%
No.
%

46.8

404

53.2

{DecoY2
Young
No.

%

No Data

1958-59

929

47.9

1,010

52.1

1959-60

377

36.5

655

63.5

1960-61

612

45.5

732

54.5

1961-62

527

55.7

419

44.3

1962-63

204

48.9

213

51.1

1963-64

377

52.1

346

47.9

1964-65

442

63.1

259

36.9

1965-66

571

62.7

340

37.3

1966-67

217

45.3

262

54.7

1967-68

245

59.9

164

40.1

1968-69

136

61.8

84

38.2

79

51.3

75

48.7

1969-70

110

55.3

89

44.7

143

35.9

255

64.1

The age ratio of birds bagged on the two state areas showed some interesting
trends as the season progressed (Table 9). The percent of immatures harvested
on the decoy area increased until mid-season, then began to decline.
On the
firing-line area, percent of immatures in the bag generally increased throughout the season.
According to Rutherford's (1968) findings, juveniles normally
become less vulnerable as the season progresses irregardless of method of
harvest, firing-line or field hunting.
But Chapman, et al. (1969), working
with the dusky Canada goose in the Williamette Valley of Oregon, found that
restricting hunting to mornings only, resulted in a greater proportionate
harvest of immatures, as most adults apparently delayed flights from refuge
areas until afternoon.
This age-specific flight pattern was indicated by a
drastic change in the age ratio of the kill when full day hunting was initiated

�- 109

toward the end of the 1964-65 season.
The increased vulnerability
of
young-of-the-year
to morning only hunting may help explain why immatures
were apparently more vulnerable to hunting in southeast Colorado (1:1.37)
than earlier in the year on the Canadian staging areas 1:0.82 (Dzubin,
pers. comm).
But the reversal in age ratios after mid-season at the
Lamar-Eads area is really not explainable at this time.

Table 9.
southeast

Weekly age ratios of Canada geese brought
Colorado.

through

check stations,

Date

Lamar-Eads
Ratio
Adu1t:Imm.

11/22-11/28

1: 1.16

41

11/29-12/5

1: 1.49

84

12/6-12/12

1:2.22

87

1:0.78

87

12/13-12/19

1:3.63

74

1:0.63

65

12/20-12/26

1:1. 33

7

1:1

8

12/27-1/2

1: 1. 75

44

1:1.43

17

1/3-1/9

1:1. 23

47

1/10-1/15

1:1

14

l/Season

opened

on December

(Decoy~
Number

Two Buttes (Firing Line)
Ratio
Adu1t:Imm.
Number

1/

o
1:1.20

22

6, 1969.

Approximately 39 percent of all birds trapped post-season in southeast Colorado
in 1970 were immatures.
This figure is a substantial increase from the 16.8
percent trapped the previous year.
The sex and age composition of each
cannon-net catch is presented in Table 10.
A summary of all age data is presented in Table 11. In total, immatures made
up approximately 58 percent of the check station sample and 38 percent of the
trapped sample.
The apparent effect of half-day hunting on the vulnerability of different age
classes makes it impossible to estimate what percent of the goose population
is composed of birds of the year, and thus evaluate 1970 production of the
flock.

�- 110 Table 10. Sex and age composition of Canada geese trapped post-season
in southeast Colorado, 1970.
Location
and Date

Males
Adult
Imm.

Females
Adult
Imm.

Total

Percent
Immatures

Turk's Pond

1/31/70

49

31

52

44

176

42.6

Two Buttes

2/4/70

50

28

56

22

156

32.1

Turk's Pond

2/6/70

13

12

19

14

58

44.8

Turk's Pond

2/19/70

34

20

35

19

108

36.1

Turk's Pond

2/26/70

29

27

42

24

122

42.5

175

ll8

204

123

620

38.9

Total

Table 11. Age comparison of southeast Colorado Canada geese, 1969-70, as
estimated by check station and trapping results.
No. of
Young

Percent
Young

No. of
Adults

Percent
Adults

Total
Birds

Lamar-Eads

255

64.1

143

35.9

398

Two Buttes

89

44.7

llO

55.3

199

Meredith Lake

27

60.0

18

40.0

45

Check Station Sample

Total
Adult/Immature

371

271

642
1:1.37

Trapped Sample
Turk's Pond

191

41.2

273

58.8

464

Two Buttes

50

32.1

106

67.9

156

Total

241

Adult/Immature

379

620
1:0.64

�- 111 -

Hunting

Pressure

and Harvest

Despite the increase in the size of the Canada goose population in southeast Colorado, it was estimated that hunter participation fell to its
lowest level in 16 years (Table 12) (Funk and Tully, 1970).
It was estimated there were 5,327 active goose hunters, down 20 percent from the
previous years figures.
All major goose harvest counties, with the exception
of Kiowa, registered declines of from 33-42 percent in the number of active
goose hunters when comparing hunter participation during the 1969-70 season
with the 1968-69 season.
Kiowa County recorded a 70 percent increase in
hunters, possibly because of increased use of the Lamar-Eads Management Area.
Because of the lack of hunter participation,
total estimated retrieved
harvest for southeast Colorado was the smallest since the 1965-66 season
(Table 12). However, hunting success increased over the 1.6 season bag
recorded during the 1968-69 season, with each goose hunter harvesting approximately 2 birds per season during the 1969-70 season.

Table 12.

Goose hunting

season statistics,

1954-69.
Arkansas Valley
Estimated
Average
Goose Hunters
Season Bag

Year

Dates of Season

Stamp
Sales

Estimated
Kill

1954

11/1 - 12/30

32,450

7,071

1.04

7,372

1955

11/1 - 12/30

39,107

9,054

1.54

13,904

1956

11/9 - 1/7

36,303

9,833

1.05

10,276

1957

11/2 - 11/31

41,794

9,113

1. 39

12,656

1958

11/17 - 1/15

41,897

10,082

1.51

15,205

1959

10/26 - 1/8

31,431

8,888

1.61

14,309

1960

10/26 - 1/8

30,592

9,838

1. 39

13,629

1961

11/10 - 1/8

24,854

7,577

1.68

11,724

1962

10/31 - 1/13

17,701

6,021

1.58

9,495

1963

11/2 - 1/15

22,940

6,668

2.17

14,444

1964

11/2 - 1/15

25,282

8,016

2.30

18,474

1965

11/2 - 1/15

20,537

6,313

1.52

9,613

1966

11/19 - 1/15

29,377

9,357

2.59

24,269

1967

11/18 - 1/14

31,064

6,975

2.23

15,558

1968

11/16 - 1/15

31,218

6,668

1.66

11 ,046

1969

11/22 - 1/15

34,281

5,327

1.99

10,597

�- 112 -

Hunters in Kiowa County registered the largest harvest during the 1969-70
season (Table 13). The bag in both Baca and Prowers Counties, where hunter
participation was down about 42 percent, declined more than 50 percent in
comparison with the 1968-69 harvest.
Table 13. Comparison of the 1969-70 southeast Colorado goose harvest, by
county, with the IS-year average, 1954-68, based on the results of the
random small game survey.

County

Lakes and Reservoirs

Number and Percent of Geese Bagged
1969-70
15 Yr. Average
No.
Percent
No.
Percent

Kiowa

Eads and Blue

3,786

35.7

3,051

21.9

Baca

Two Buttes and Turk's

2,070

19.5

5,284

38.0

Bent

John Martin,
Horsecreek

1,946

18.4

1,911

13.7

Crowley

Meredith

and Henry

973

9.2

699

5.0

Prowers

Two Buttes and Eads

884

8.3

2,208

15.9

Otero

Horse Creek, Cheraw,
Dye and Holbrook

531

5.0

347

2.5

Pueblo

248

2.3

179

1.3

Las Animas

88

0.8

122

0.9

Huerfano

71

0.7

116

0.8

Total

10,597

Two Buttes and Lamar-Eads

Blue and

Management

l3,9l7

Areas

The number of hunters utilizing the Two Buttes Management area declined for
the fourth year in succession (Table 14). Hunting success was slightly better
than during the 1968-69 season with 659 hunters bagging 248 geese for an
average bag per hunter of 0.38 birds.
Hunting pressure declined as the season progressed but hunting success was irregular (Table 15) and, as would be
expected, closely dependent on whether morning goose flights occurred.

�- 113 Table 14. Goose harvest, hunting pressure, and hunter success, Two Buttes
Management Area.
No. of
Individual
Hunters

No. of
Hunter
Days

No. of
Geese
Bagged

Ave. Bag
per
Hunter

Ave. Bag
per
Hunter Day

1961-62

2,392

4,758

945

0.40

0.20

1962-63

1,479

3,178

418

0.28

0.13

1963-64

1,750

3,659

728

0.42

0.20

1964-65

1,996

3,946

721

0.36

0.19

1965-66

2,596

5,264

941

0.36

0.18

1966-67

1,257

2,413

493

0.39

0.20

1967-68

840

1,554

433

0.52

0.28

1968-69

792

1,309

243

0.31

0.19

1969-70

659

1,182

248

0.38

0.21

Year

Table 15. Hunter activity and goose harvest by weekly intervals, Two Buttes
Reservoir, 1969-70.
Week
Interval

Number of
Hunter Days

12/6 - 12/12

Successful
Hunter Days

No. of
Geese Bagged

Ave. Bag Per
Hunter Day

516 (43.7)

86

114

0.22

12/13 - 12/19

209 (17.7)

62

82

0.39

12/20 - 12/26

118 (10.0)

8

9

0.08

12/27 - 1/2

187 (15.8)

17

20

0.11

1/3 - 1/9

81 ( 6.9)

0

0

0.00

1/10 - 1/15

71 ( 6.0)

14

23

0.32

187

248

0.21

Totals

}j

1,182

1/ Percent of total in parentheses.

�- 114 The Lamar-Eads Management Area, in its second year of existence, recorded
an increase in hunter use in comparison with the previous season.
Approximately 700 hunters bagged S12 for an average bag per hunter of 0.73 (Table
16). Hunting pressure at the Lamar-Eads Area was generally consistent
throughout the season but hunting success, as at Two Buttes, was low during
full moon periods (Table 17).
Table 16.
Management

Goose harvest,
Area.

hunting

pressure,

Year

No. of
Individual
Hunters

No. of
Hunter
Days

No. of
Geese
Bagged

1968-69

S98

1,062

160

1969-70

703

1,361

S12

Table 17.
Management

Hunter activity
Area, 1969-70.

Week
Interval

Number of
Hunter Days

and goose harvest

1./

Successful
Hunter Days

and hunter

success,

Ave. Bag
per
Hunter

Lamar-Eads

Ave. Bag
per
Hunter Day

O.lS
0.73

by weekly

0.38

intervals,

No. of
Geese Bagged

Lamar-Eads

Ave. Bag Per
Hunter Day

11/22 - 11/28
216 (lS.9)
(Full moon 11/23)

31

42

0.19

11/29 - 12/S

133 ( 9.8)

SO

81

0.61

12/6 - 12/12

262 (19.3)

130

188

0.72

12/13 - 12/19

170 (12.S)

Sl

80

0.47

12/20 - 12/26
107 ( 7.9)
(Full moon 12/23)

6

7

0.07

12/27 - 1/2

160 (11. 8)

32

48

0.30

1/3 - 1/9

190 (14.0)

30

SO

0.26

1/10 - l/lS

123 ( 9.0)

9

16

0.13

339

S12

0.38

Totals

1/ Percent

1,361
of total in parentheses.

�- 115 Tables 18 and 19 indicate the effect of repeated trips by hunters on
their hunting success.
As during past years, the success of hunters
increased as the number of days hunted increased.

Table 18. The relationship of individual
Two Buttes Reservoir, 1969-70.

hunter activity

to hunting

Number of
Days Hunted

Total No.
of Hunters

Percent of
Hunters Succ.

No. of
Geese Bagged

1

379

43

11. 3

55

0.15

2

160

45

28.1

62

0.19

3

56

25

44.6

42

0.25

4

31

15

48.4

25

0.20

5

16

12

75.0

20

0.25

6

8

7

87.5

15

0.31

7

3

3

100.0

8

0.38

8

2

2

100.0

5

0.31

9

2

2

100.0

9

0.50

10

1

1

100.0

3

0.30

14

1

1

100.0

4

0.29

Totals

659

156

23.7

248

0.24

No. of
Succ. Hunters

success,

Ave. Bag/
Hunter Day

�- 116 Table 19.
Lamar-Eads
Number of
Days Hunted

The relationship
Area, 1969-70.
Total No.
of Hunters

of individual

hunter activity

to hunting

success,

No. of
Succ. Hunters

Percent of
Hunters Succ.

No. of
Geese Bagged

Ave. Bag/
Hunter Day

1

411

49

11.9

73

0.18

2

171

47

27.5

85

0.25

3

45

28

62.2

57

0.42

4

27

12

44.4

31

0.29

5

22

17

77.3

51

0.46

6

7

7

100.0

18

0.43

7

4

2

50.0

10

0.36

8

4

3

75.0

10

0.31

9

2

2

100.0

5

0.28

10

2

0

0.0

0

0.00

11

2

2

100.0

9

0.41

13

2

2

100.0

7

0.27

15

1

1

100.0

12

0.80

17

1

1

100.0

19

1.12

18

1

1

100.0

19

1.06

19

1

1

100.0

6

0.32

703

175

24.9

412

0.30

Totals

Banding

Investigations

Trapping and banding was accomplished after the close of the 1969-70 seasons.
Objectives were to band 500 geese in the Two Buttes area and the same number
at another location in the Arkansas Valley.
Because of the high water levels
of reservoirs in the northern areas, all trapping efforts were concentrated
in the Two Buttes area. A total of 599 new bands were placed on Canada geese.
Each individual catch is categorized by sex and age in Table 20. Approximately
47 percent of the banded birds were males.

�- 117 -

Table 20. Age and sex composition
Colorado, 1970.

Date

Males
Adult
Immature

Location

January

31

Turk's Pond

February

4

Two Buttes

February

6

February
February

of Canada geese banded

Adult

in southeast

Females
Immature

44

25

49

44

45

28

48

21

Turk's Pond

15

14

19

15

19

Turk's Pond

28

20

31

19

26

Turk's Pond

34

29

45

26

166

116

192

125

Res.

Total

Weights
The mean weights of all geese harvested on the two state-controlled
ment areas and birds trapped are presented in Table 21.

Table 21. Comparison of weights
southeast Colorado, 1969-70.

Source

manage-

of geese from trap and check station

Adult Ave.
Weight (lbs.) Range

samples,

Imm. Ave.
Total Ave.
Weight (lbs.) Range Weight (lbs.) Range

Two Buttes
Check Station

5.56(110)1/3.75-7.75 5.25(85)

3.25-7.25

5.43 (195) 3.25-7.75

6.03(379) 4.00-10.50 5.74(241) 3.38-8.75

5.92(620) 3.38-10.50

Two Buttes-Turks
Trapping
Lamar Mgmt. Area
Check Station

5.73(142)

1/ Sample sizes in parentheses.

3.50-9.50 5.41(248) 3.25-8.75. 5.53(390) 3.25-9.50

�- 118 -

In past years, mean weights of birds bagged on the Two Buttes Management
Area have been consistently larger than birds trapped post-season in the
Two Buttes-Turk's area. This year the reverse was true with trapped birds
averaging nearly .5 (one-half) lb. more than harvested birds.
Past
explanations for harvested birds weighing more have been: (1) hunters
selecting larger birds, (2) weight loss during the season, and (2) greater
trap vulnerability of sub-populations containing smaller birds.
The 196970 weights of harvested geese presented in Table 22 indicate that the geese
may have lost a small amount of weight on the average as the season progressed.
However, field personnel indicated that the birds maintained
better body condition throughout the hunting season than they had in previous years, possibly because the half-day season allowed geese to feed
unmolested during the afternoon.
The maintenance of good body condition
may have enabled the geese to add additional weight quickly at the end
of the hunting season.

Table 22. Average weights of geese bagged by time interval on two state
management areas in southeastern Colorado during the 1969-70 hunting season.
Two Buttes !/
Adults
Immatures

Interval

Lamar-Eads
Adults
Immatures

Nov. 22 - Nov. 28

5.92(21)

Nov. 29 - Dec. 5

5.72(31)

5.38(50)

Dec. 6 - Dec. 12

5.80(28)

5.47(54)

5.61(49)

5.41(37)

Dec. 13-

Dec. 19

5.56(16)

5.40(57)

5.46(47)

5.04(25)

Dec. 20 - Dec. 26

6.25(1)

6.18(4)

6.08 (3)

5.13(4)

Dec. 27 - Jan. 2

5.73(14)

5.57(27)

5.46(7)

5.22(8)

Jan. 3 - Jan. 9

5.60(22)

4.61(24)

0

0

Jan 10 - Jan. 15

5.21(7)

5.00(7)

5.88(4)

Jj

5.51(22)

5.25(11)

1/ Season opened December 6, 1969 at Two Buttes.
size in parentheses.

2/ Sample

The data presented in Table 23 indicate that some unusually large birds were
present in three of the five cannon-net catches.
Yet the average of any of
the five catches was greater than the average weight of harvested birds.

�- 119 -

Table 23. Weights of Canada geese captured with cannon-nets in southeast
Colorado, 1970.

Date

Location

January 31

Turk's Pond

Grouping

Number

AM

49
31
50
44

6.41
6.10
5.57
5.28

4.75-8.00
5.13-8.75
4.00-8.25
3.38-7.25

176

5.82

3.38-8.75

50
28
56
22

5.85
5.66
5.31
5.31

4.13-7.38
4.25-6.75
4.13-6.38
4.13-6.38

156

5.55

4.13-7.38

13
12
19
14

6.29
6.06
5.48
5.69

4.88-6.88
5.00-6.88
4.13-6.38
4.88-6.38

58

5.84

4.13-6.88

34
20
35
19

6.99
6.15
6.14
5.72

5.13-9.38
4.13-7.63
5.00-10.50
4.88-6.63

108

6.33

4.13-10.50

29
27
42
24

7.05
6.45
5.99
5.32

4.50-8.63
5.50-8.00
4.38-7.25
3.75-6.50

122

6.21

3.75-8.63

175
118
204
123

6.46
6.08
5.67
5.41

4.13-9.38
4.13-8.75
4.00-10.50
3.38-7.25

620

5.92

3.38-10.50

1M
AF

IF
Sub-totals
February 4

Two Buttes Res.

AM
1M

AF
IF
Sub-totals
February 6

Turk's Pond

AM
1M

AF
IF
Sub-totals
February 19

Turk's Pond

AM
1M

AF
IF
Sub-totals
February 26

Turk's Pond

AM
1M

AF
IF
Sub-totals
All Catches

AM
1M

AF
IF
GRAND TOTAL

Ave. Weight

Range

�- 120 -

At this point it seems the interpretation of weight data can be only
speculative.
Certainly weight, along with the comparative size of any
goose, gives some indication of body condition.
But until specific
studies are designed to determine what weight and/or body condition
indicates in terms of sub-speciation and/or productivity, interpretation
of weight data will continue to be mainly academic.

Distribution

of Harvest

The distribution of band recoveries through the 1968-69 season is presented in Table 24. The distribution of the 1968-69 harvest showed some
changes from recent trends.
The most significant change occurred on the
Canadian prairies where more banded birds were taken in Alberta than
Saskatchewan, reversing a trend which began during the 1964-65 season.
The Nebraska take of southeast Colorado banded birds declined approximately 50 percent from recent previous levels.
An important distribution
change which is not indicated by the table is the movement of Short Grass
Prairie birds into the Hi-Line population area in northcentral Colorado.
During the 1967-68 season, two southeast Colorado banded birds and one
panhandle of Texas bird were recovered in northcentral Colorado.
In
1968-69, four southeast Colorado birds and three Texas birds were taken
in that same area.
The provinces of Alberta and Saskatchewan along with southeast Colorado
continue to be the major harvest area of southeast Colorado banded birds.

Hunting

Mortality

Estimates

Tables 25 through 30 present band recovery data from all southeast Colorado
post-season banding efforts from 1951 through the hunting season of 1968-69.
Tables 25, 26, and 27 present mortality rates calculated by the compositedynamic method.
Tables 28, 29, and 30 present these same estimates calculated by the relative recovery rate method.
According to the composite method, both juveniles and adults continue to
have a first year mortality rate of about 28 percent (Tables 25 and 26).
Annual mortality rate for all age classes combined remains at 26 percent,
consistent with estimates for the previous two years (Table 27).
Analysis by the relative recovery rate method gives estimates of annual
mortality for birds banded during anyone
year.
By averaging these estimates, an average mortality rate for an age class and/or population can
be obtained.
Rates calculated by the relative method continue to be
slightly higher than those determined by the composite dynamic method.

�Table 24.

Percentages of total band recoveries, Arkansas Valley post-season

bandings, by area and year of recovery, all banding years combined.

195152

195253

195354

195455

195556

195657

195758

1958- 195959
60

196061

1961- 196262
63

PercenTotal No. tage of
1963- 1964- 1965- 1966- 1967- 1968- of Recov- Total Recoveries
67
68
64
65
66
69 eries

Far North above 530 10.5

9.0

9.0

10.7

10.5

6.2

6.3

7.0

7.9

7.7

7.9

6.3

15.5

15.2

4.7

10.4

7.5

9.9

240

9.2

23.4
17.6

31.0
24.3

33.3
17.3

28.2
10.5

29.7
9.4

39.3
9.8

35.4
14.6

33.3
9.4

38.7
9.0

34.9
9.3

33.3
19.4

28.4
17.4

26.8
8.5

27.9
10.9

19.4
16.4

14.2
24.6

23.4
17.0

762
389

29.3
14.9

0.7

1.5

4

0.2

0.7

0.6
T
0.6
0.2
6.2
28.9
0.3
0.6
0.8

Recovery Year
Area

Provinces below 530
26.4
Alberta
18.5
Saskatchewan
B. C., Manitoba,
1.3
Ontario
Central Flyway
1.3
Montana
N. Dakota
1.3
Wyoming
1.3
S. Dakota
7.9
Nebraska
25.0
Colorado
Kansas
Oklahoma
1.3
New Mexico
Texas
Panhandle
Waggoner Ranch
Gulf Coast
5.2

Pacific Flyway
Mississippi

1.1

0.5

0.5

1.0

1.3
0.7
0.7

3.2
23.1
0.5
0.5
1.6

4.3
22.9
0.5
2.4
1.9

6.0
23 ..
8
0.7
0.7
2.0

7.4
7.4
2.1

0.5

2.1

1.4

0.5
2.8

8.1
36.3

5.5
39.0
0.8

8.9
24.0

1.6

1.8

1.6
0.8
0.8

2.3
2.3

4.5
0.9

2.4

3.2

2.7

0.6

0.7

0.7

5.5
21. 7
1.0
0.5
1.0

6.7
30.1

2.0
1.4
8.8
35.4
0.7
1.4

0.7
0.7
9.7
32.4

4.6
0.8
0.8

4.0
2.0
1.0

7.3
2.4
0.6

2.7
2.0

0.8

1.5

1.2

1.4

0.7
1.4

1.3

4.9
28.3
0.7
1.4
0.7

7.2
36.6

6.4
26.6

2.8
0.7
0.7

0.7
1.4

2.8

1.4

3.9
5.2
1.2

0.5
5.4
31. 5
0.8

5.4

4.8

3.1
30.9

0.7

0.6
7.5
0.7

0.8

Flyway

Mexico
Total Number
of Recoveries

1.8

76

0.5

0.5

188

210

150

124

128

112

144

138

155

129

129

201

164

147

134

1.4
0.7
0.7

9.0
29.1

4.3
31.9

1.5
0.7

1.4

15
2
15
4
161
752
9
15
22

2.2

5.0
0.7
1.4

100
44
16

3.8
1.7
0.6

0.7

1.4

50

1.9

0.7

0.7

3

0.1

1

T

7.5

134

141

2604

....
N
....

�Table

25 . Composite

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965 -66
1966 -67
1967-68

Total
Recoveries

dynamic analysis

of band recoveries

from Canada geese banded as juveniles,

by Year Following
10
9

Arkansas Valley,

Colorado,

1951-1968.

1

2

3

4

5

6

Recoveries
8
7

Banding
11

12

13

14

15

16

17

18

300
628
578
0
0
0
0
176
298
167
248
123
399
251
198
0
285
270

31
58
52

25
31
25

11
33
25

11
20
24

4
16
20

5
8
11

4
7
11

7
6
9

1
4
2

1
4
4

0
0
4

0
5
1

0
2
3

1
1
1

1
0
1

0
0
0

0

1

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X

X
X

X

X
X
X
X

X
X
X
X

X
X
X

X
X

X

X

X

21
25
16
15
10
40
16
15

16
16
10
5
12
24
11
7

6
16
3
4
10
9
11
8

4
8
14
7
0
14
13
1

4
7
8
8
1
6
5

1
3
0
7
1
2

4
0
0
3
1

0
1
1
2

3
1
0

1
2

X

X

X

18
16

10

3921

333

192

136

116

79

38

30

26

11

12

4

6

5

3

2

0

1

1

Banded
Birds
Eligib le

3921

3651

3366

3366

3168

2917

2518

2395

2147

1980

1682

1506

1506

1506

1506

1506

928

300

Recoveries pe r
1,000 Banded

84.9

52.6

40.4

34.5

24.9

13.0

11.9

10.9
213.5

5.1

6.1

2.4

4.0

3.3

2.0

1.3
0.0
~ - 301. 7

1.1/

3.3

Alive Going
into Period

301.7 ,,216.8_ 164.:2. 123.8

89.3

64.4

51.4

28.6

23.5

17.4

15.0

11.0

7.7

3.3

28.17.

5.7
4.4
~=1172.1
25.7%

44/

Mor ta lity Ra te

39.5
867.1
24.6%

0

•....
N
N

�Table 26 •

Composite dynamic analysis of band recoveries

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
Total
Recoveries

from Canada Geese banded as adults, Arkansas Valley, Colorado,

1951-1968.

1

2

3

4

5

6

Recoveries b~ Year Fo11owins Bandins
12
11
10
7
8
9

344
650
900
0
0
0
0
347
217
250
306
334
369
335
780
0
389
646

45
71
85

23
37
55

19
24
50

10
23
31

7
21
25

12
6
20

5
11
9

3
6
12

3
9
10

2
7
6

2
7
2

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

33
21
27
29
26
35
24
55

27
9
16
13
18
20
12
40

17
7
6
18
12
14
15
20

8
4
12
12
5
4
10
22

5
4
8
4
13
11
4

8
2
4
7
3
5

2
1
6
3
2

2
1
3
2

2
0
2

2
1

0

X

X

X

29
35

26

5867

515

296

202

141

102

67

39

29

26

18

11

8

5

1

2

2

0

0

Banded
Birds
Eligible

5867

5221

4832

4832

4052

3717

3348

3014

2708

2458

2241

1894

1894

1894

1894

1894

944

344

Recoveries Per
1,000 Banded

87.8

,56.7

41.8

29.2

25.2

18.0

11.6

9.6

9.6

7.3

4.9

4.2

2.6

0.5

1.1

1.1

0.0 I

0.0

--

13

14

15

16

17

18

0
4
4

1
2
2

1
0
0

0
1
1

1
0
1

0
0

0

X
X
X
X

X
X
X

X
X

X

•....
N

w

I: = 311.2

223.4
Alive Going
Into Period

311.2

223.4

166.7 124.9

95.7

70.5

52.5

40.9
862.8

Mortality

Rate

28.2%

25.9%

31.3

21.7

14.4

9.5

5.3

2.7

1.1

2.2
r-=1l74.0

26.5%

0.0

�Table 27.

Composite dynamic analysis

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
Total
Recoveries

of band recoveries

from Canada Geese banded in all age classes, Arkansas Valley, Colorado,
b~ Year Following Banding
9
11
10
12

1951-1968.

1

2

3

4

5

6

Recoveries
7
8

644
1278
1478
0
0
0
0
523
515
417
554
457
768
586
978
0
674
922

76
129
137

48
68
80

30
57
75

21
43
55

11
37
45

17
14
31

9
18
20

10
12
21

4
13
12

3
11
10

2
7
6

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

54
46
43
44
36
75
40
70

43
25
26
18
30
44
23
47

23
23
9
22
22
23
26
28

12
12
26
19
5
18
23
23

9
11
16
12
14
17
9

9
5
4
14
4
7

6
1
6
6
3

2
2
4
4

5
1
2

3
3

0

X

X

X

47
51

36

9794

848

488

338

257

181

105

69

55

37

30

15

14

10

4

4

2

Banded
Birds
Eligible

9794

8872

8198

8198

7220

6634

5866

5409

4858

4438

3923

3400

3400

3400

3400

3400

1922

Recoveries Per
1,000 Banded

86.6 \ 55.0

41.2

31,3

25.1

15.8

11.8

10.2

7.6

6.8

3.8

4.1

2.9

1.2

1.2

0,6

0.5

3.9
2.7
2.:"=1176.7

2.1

13

14

15

16

17

0
9
5

1
4
5

2
1
1

1
1
2

1
0
1

0
1

X
X
X
X

X
X
X

X
X

X

•....
N

~

307.3

220.7

165,7 124,5

93.2

68.1

52.3

40.5
867.8

Mortality Rate

28.2%

25,2%

644

I

1.6

r = 307.3

219.1
Alive Going
Into Period

18

30.3

22.7

15.9

12.1

8.0

5.1

1.6

26.1%

�Table 28. Relative recovery rate analysis of band recoveries from Canada geese banded as juveniles, Arkansas
Valley, Colorado, 1951-1968.
Winter
Banded

Number
Banded

1950-51

300

CU ])

72

---

1951-52

628

196

138

.3121

1952-53

578

193

CU

.3339

1957-58

176

8U

39

1958-59

298

79

1959-60

167

1960-61

248

Number Recoveries
1 - n
2 - n

Recovery Rates
1 - n
2 - n

Survival
Rate

Mortality
Rate

.2400

.769

.231

.2197

.658

.342

---

.2216

.836

.164

54

.2651

.1812

.582

.418

52

36

.3114

.2156

1.049

51

36

.2056

.1452

.510

.490

1961-62

123

35

25

.2846

.2033

.854

.146

1962-63

399

95

55

.2381

.1378

.618

.346

1963-64

251

56

40

.2231

.1594

1.018

1964-65

198

31

CU

.1566

1966-67

285

CU

10

---

.0351

.592

1967-68

270

16

--

.0593
1.7589

70486

.736
.264

0749
.251

3,921

2.3898
Average survival rates
Average mortality rates

.11 Cannot use.

t-'
N
V1

.408

�Table 29. Relative recovery rate analysis of band recoveries from Canada geese banded as adults, Arkansas
Valley, Colorado, 1951-1968.
Winter
Banded

Number
Banded

Number Recoveries
1 - n
2 - n

1950-51

344

cu 1:..1

1951-52

650

1952 53

Recovery Rates
1 - n
2 - n

Survival
Rate

Mortality
Rate

89

---

.2587

.734

.266

229

158

.3523

.2431

.699

.301

900

313

CU

.3478

1957-58

347

CU

73

---

.2104

.913

.087

1958-59

217

50

29

.2304

.1336

.398

.602

1959-60

250

84

57

.3360

.2280

.793

.207

1960-61

306

88

59

.2876

.1928

.815

.185

p

1961-62

334

79

53

.2365

.1587

.658

.342

1962-63

369

89

54

.2412

.1463

.754

.246

1963-64

335

65

41

.1940

.1224

.697

.303

1964-65

780

137

CU

.1756

1966-67

389

CU

26

---

.0668

1.232

646

35

--

2.4556

1.7608

7.693

Average survival rates

.717

.769

Average mortality rates .283

.231

1967-68

-5,867

1:..1

Cannot use.

.0542

-

t-'
N
0\

�Table 30. Relative recovery rate analysis of band recoveries from Canada geese banded in all age classes,
Arkansas Valley, Colorado, 1951-1968.
Survival
Rate

Mortality
Rate

.2500

.752

.248

.2316

.676

.324

---

.2141

.855

.145

83

.2505

.1612

.494

.506

136

93

.3261

.2230

.889

.111

139

95

.2509

.1715

.687

.313

Winter
Banded

Number
Banded

Number Recoveries
1 - n
2 - n

Recover~ Rates
2 - n
1 - n

1950-51

644

cuy

161

---

1951-52

1,278

425

296

.3326

1952-53

1,478

506

CU

.3424

1957-58

523

CU

112

1958-59

515

129

1959-60

417

1960-61

554

1961-62

457

114

78

.2495

.1707

.712

.288

1962-63

768

184

109

.2396

.1419

.687

.313

1963 -64

586

121

81

.2065

.1382

.805

.195

1964-65

978

168

CU

.1717

1966-67

674

CU

36

---

.0534

.966

.034

1967-68

922

51

---

.0553
1.7556

7.523

.724
.276

.752
.248

9,794

2.4251
Average survival rate
Average mortality rate

1./ Cannot use.

•....
N
-...J

�- 128 -

LITERATURE

CITED

Chapman, J. A., C. J. Henny, and H. M. Wight.
1969. The status,
population dynamics, and harvest of the dusky Canada goose.
Wi1d1. Monogr. No. 18. 48 p.
Funk, H. D., and R. Tully.
1970. Colorado small game hunter harvest
survey --- 1969. Dept. Natural Resources, Colo. Game, Fish and
Parks Div., Fed. Aid Project W-88-R.
Grieb, J. R. 1970. The shortgrass
Wi1d1. Monogr. No. 22. 49 p.

prairie

Canada goose population.

Rutherford, W. H. 1968. Differential mortality of juvenile Canada
geese in the Arkansas Valley of Colorado.
Proc. Western Assoc.
Game and Fish Comms. 48:288-293.

Prepared
Szymczak
Asst. Wildlife Researcher

�October

- 129 -

JOB PROGRESS

State

Work

No.

Plan No.

Job Title
Period

REPORT

COLORADO

of

Project

1970

Covered:

Personnel:

W-88-R-1S

3
Investigation
December

of Mallard

Migratory

Bird Investigations

Job No.

6

Management

1, 1969 through

March

Units

of Eastern

Colorado

31, 1970

Richard Hopper, ,Jack Grieb, Mike Szymczak, Clait Braun,
Robert Kitzmiller, Larry Green, Ray Schmidt, David Cramer,
Velma Fredrickson,
Howard Funk and other members of the
Colorado Division of Game, Fish and Parks.

ABSTRACT

Winter banding efforts in nine study units in eastern Colorado in 1969-70
yielded a total of 3,849 mallards banded in January and February.
Size
of the peak wintering population increased substantially
over 1968-69
with about 4333,000 birds censused in early January 1970.
Sex ratio counts
during the banding period again indicated 60 percent of the wintering
mallard population was composed of males.
Analysis of band recovery data
was postponed until a complete tape on results since initiation of the
study in 1963-64 is obtained from the Bureau of Sport Fisheries and Wildlife.
The experimental
late point-system
season granted in portions of six states
in The Central Flyway was judged a success.
Harvest in Colorado during
the 23-day season was estimated at 43,000 birds of which 86 percent were
mallards.
The point-system
regulation seemed to have been effective in
placing emphasis on drake mallard harvest as approximately
90 percent of
the mallards taken were males.
Reaction to the regulation by hunters
was favorable.

RECOMMENDATIONS

1.

2.

Continue banding 600 mallards in each of the nine study units for one
more year until results of analyses indicate either the need to continue
at the same rate or to drop back to banding for the purpose of monitoring.
All other procedures would remain similar to previous years for at
least one more year.

��- 131 -

INVESTIGATION

OF MALLARD

MANAGEMENT

UNITS OF EASTERN

COLORADO

Howard D. Funk

Efforts were continued on the intensive Colorado winter mallard banding
program initiated in 1963. Purpose of the banding was to continue gathering
data on terminal wintering mallards in the State by study unit and also to
take part in the cooperative study of a similar nature in the Central
Flyway as a whole.
Because complete recovery data tapes were expected shortly
from the Bird Banding Laboratory in Patuxent, it was decided to delay analysis of recoveries until all data from initiation of the study to the present
had arrived.
Thus, this report covers only the banding efforts, population
and sex ratio surveys, and harvest and hunting pressure surveys conducted
during the winter of 1969-70.
Some information on the late experimental
point-system season in 1969-70 is also presented.

P. S. OBJECTIVE
To develop a harvest

formula

for Colorado

SEGMENT
1.

2.

3.

4.

mallard

management

units.

OBJECTIVES

To trap and band mallards after hunting season in the following eight
study areas for the purpose of obtaining migration, life history and
annual mortality data:
(a) South Platte Valley Denver to Greeley, (b)
South Platte Valley Greeley to Fort Morgan, (c) South Platte Valley Fort
Morgan to Sterling, (d) South Platte Valley Sterling to Julesburg,
(e) the Fort Collins area, (f) Bonny Reservoir, (g) Arkansas Valley
Pueblo to Rocky Ford, and (h) Arkansas Valley Rocky Ford to the Kansas
line. Feasibility of winter banding in the San Luis Valley will also
be investigated.
To conduct surveys for the purpose of obtaining winter population estimates of mallards; age and sex ratios in the hunter's bag, wintering
populations, and trap samples, and hunter pressure and harvest data by
study area.
To gather and tabulate band recovery data from the Bird Banding Office
from mallards banded in Colorado since 1963-64 for use in making preliminary estimates on migration and harvest routes and annual mortality for
use in final analysis during a later segment.
To conduct further special mallard seasons designed to take advantage of
surplus birds where feasible.

METHODS

AND MATERIALS

Trapping methods and materials were similar to those described in other
segment reports since 1964. Most birds were captured in cage traps with a
few taken by use of cannon-nets.
Birds were captured where and when possible,

�- 132 some at newly found trapping sites.
The San Luis Valley was added to the
list of study units with birds being banded at the Monte Vista National
Wildlife Refuge.
Periodic aerial censuses were conducted to obtain population estimates and locations of birds.
Sex ratio ground counts were also
taken during the period covered.
Much effort was expended by many people
in the experimental late point-system season in the way of issuing special
hunting permits, hunter performance surveys, special wing collection survey,
conduct and evaluation of a special hunter survey, and writing a comprehensive cooperative report involving six Central Flyway states.

RESULTS AND DISCUSSION

Trapping

and Banding

Results of the 1969-70 winter trapping and banding program are illustrated
in Table 1. With the addition of the San Luis Valley, we now have nine
study units in which birds were trapped (Units 1, 2, 3, 4, 6, 9, 12, 13, and
15; Fig. 1). Due to unseasonably warm weather and lack of manpower, no
banding was accomplished in Unit 1 and quotas of 600 per unit were not reached
in Units 6, 13, and 15. This was unfortunate, as we will have gaps in data
which will affect various analyses, especially mortality and survival estimates and life tables.
As always, we had problems in capturing females,
especially adults, but often made up discrepancies somewhat by banding over
the quotas on immature females.
Number banded in all units totalled 3,849
birds with fairly even ratios banded by age and sex on an overall basis.
The trapping site utilized in the San Luis Valley was the Monte Vista
National Wildlife Refuge.
Efforts will be made to trap in other areas in
the Valley in the future because of seemingly biased conditions.
Numbers of
adults trapped far outnumbered immatures and many retraps were taken from
previous bandings.
It is believed that birds present on the Refuge are
not representative of the entire San Luis Valley wintering mallard population.
Validity of the mallard wing aging
technique was investigated in two past
segments by holding groups of immature birds over winter and periodically
examining the progression of molt patterns.
The investigation showed that
age of immatures could be determined by examination of wings through February and even later each year. Thus, banders were advised of the validity of
the method in an article in the Journal of Wildlife Management (Hopper and
Funk 1970).

Winter

Sex RAtio Counts

As in previous years, ground sex ratio counts were conducted in the various
study units during the period of banding.
Results are presented in Table 2
by unit, with sample sizes by sex and percent males listed by dates.
Data
were collected from only five of the nine units but probably are representative of ratios in all areas.
Percent males in the samples varied somewhat
between dates and areas but in most cases averaged approximately 60 percent
as in past years.
The Arkansas Valley figures averaged only 54 percent males
this segment.
Some counts in the Arkansas Valley were conducted in Units 10
and 11 as well as Unit 12, but for simplicity, all were combined under Unit
12 for purposes of reporting (Fig. 1).

�Table 1. Numbers and percentages of mallards in the banded sample by sex, age, and area, eastern Colorado,
1969-70.

Management
Units

Adult

Irmnature

Number
Banded

No.

(1) Sterling-Julesburg

0

0

0

0

0

0

0

0

0

(2) Fort Morgan-Sterling

600

141

23.5

71

11.8

150

25.0

238

39.7

(3) Greeley-Fort Morgan

600

151

25.2

104

17.3

159

26.5

186

31.0

(4) Fort Collins

59l

153

25.9

112

19.0

150

25.4

176

29.7

(6) Denver-Greeley

36

15

47.7

1

2.8

15

47.7

5

13 .8

(9) Bonny Reservoir

645

155

24.0

156

24.2

150

23.2

184

28.6

(12) Rocky Ford-Lamar

686

152

22.2

163

23.8

l37

20.0

234

34.0

(13) Pueblo-Rocky Ford

229

150

65.5

39

17.0

29

12.7

11

4.8

(15) San Luis Valley

462

182

39.4

155

33.5

62

13.4

63

13.7

3,849

1,099

28.6

801

20.8

852

22.1

1,097

28.5

Male

Female
No.
%

%

Male
No.

Female
%

No.

%

t-'

w
w

�u'
c-

u:

m

/

c,.,
--I'"
•••••••
~

es
••••••

:it:

"
~

.'I

o

x

u

~
c.::I

Po

~

~

~

-

�- 135 Table 2.

Date

Mallard sex ratio ground counts by study units, 1969-70.

Location

Male

Number Ducks Counted
Female
Total

Percent
Males

Sterling-Julesburg (Unit 1)
1-26-70

South Platte River

337

145

482

69.9

1-29-70

South Platte River

387

176

563

68.7

1-31-70

South Platte River

339

182

521

65.1

2-18-70

South Platte River

349

236

585

59.6

2-19-70

South Platte River

356

223

579

61.5

2-20-70

South Platte River

321

197

518

62.0

2-20-70

Jumbo Reservoir

324

196

520

62.3

2-21-70

South Platte River

337

208

545

61.8

2,750

1,563

4,313

63.8

Sub-Totals

Fort Morgan-Sterling (Unit 2)
1-26-70

South Platte River

311

192

503

61.8

1-31-70

South Platte River

309

242

551

56.1

2-3-70

South Platte River

231

163

394

58.6

2-4-70

South Platte River

366

234

600

61.0

2-20-70

Prewitt Reservoir

333

203

536

62.1

2-21-70

Prewitt Reservoir

317

197

514

61.7

1,867

1,231

3,098

60.3

Sub-Totals
Greeley-Fort Morgan (Unit 3)
1-24-70

South Platte River

295

241

536

55.0

1-31-70

South Platte River

298

211

509

58.5

2-21-70

South Platte River

313

211

524

59.7

906

663

1,569

57.7

Sub-Total

�- 136 -

sex ratio ground counts by study units, 1969-70

Mallard

Table 2.

Number Ducks Counted
Total
Female
Male

Location

Date

Fort Collins

Area

(Continued) .

Percent
Males

(Unit 4)

2 -1-70

New Windsor

Reservoir

326

221

547

59.6

2-2-70

New Windsor

Reservoir

363

222

585

62.0

2 -3 -70

New Windsor

Reservoir

339

250

589

57.6

1,028

693

1,721

59.7

250

242

492

50.8

Sub-Totals

Arkansas

Valley

(Unit 12)

1-19-70

Verhoeff

1-20-70

Two Buttes Reservoir

128

90

218

58.7

1-23-70

John Martin

Reservoir

243

219

462

52.6

2-2-70

John Martin

Reservoir

225

187

412

54.6

2 -2 -70

Two Buttes Reservoir

223

185

408

54.6

2-2-70

Upper Queen's

64

40

104

61.5

1,133

963

2,096

54.0

Sub-Totals

Pond

Reservoir

It is difficult to realize the significance of results of sex ratio counts
after hunting season.
Males have averaged about 60 percent of Colorado
wintering mallard populations since initiation of the study in 1963-64.
Increased harvest of approximately 40,000 drakes during the late drake
season of 1968-69 (Grieb et al. 1969) and 35,000 during the late pointsystem season of 1969-70 (Funk et a1. 1970) apparently failed to have much
effect on sex composition of mallard flocks in the State.
However, sampling
methods and other unknown factors could be responsible for our inability
to detect changes.
Yet there is no reason to doubt that actual post-season
sex composition
of wintering Colorado mallard flocks is not close to 60 percent males.

�- 137 -

Winter Aerial Surveys
As in 1968-69, four periodic aerial censuses were conducted during December
and January to estimate numbers present by area (Table 3). Good production
on the breeding grounds in 1969 was reflected by increases in wintering
birds present.
Peak counts in each unit usually occurred during the midwinter survey the first part of January.
This meant that peak populations
of mallards were probably present during most of the late experimental pointsystem season from December 13, 1969 through January 4, 1970. High count
was approximately 433,000 birds in early January with most unit totals
decreasing by mid-January to an estimated total of 363,000 birds.
Some of
the reason for decreased numbers by mid-January could have been due to birds
moving south, as indicated by Arkansas Valley figures, or to other states.
However, the winter of 1969-70 was very mild and birds were scattered, making
census difficult.

Table 3.
1969-70.

Aerial duck counts by interval and study unit, eastern Colorado,

Number Ducks Counted by Date
Dec. 19-23, Jan. 5-6, 9, Jan. 19-20, 23
1969
1970
1970

Area and Study Unit

Dec. 9-10,
1969

(1) Sterling-Julesburg

23,200

18,263

20,300

17,030

(2) Fort Morgan-Sterling

27,800

34,830

46,650

32,380

(3) Greeley-Fort

33,990

46,900

46,550

28,295

(4) Fort Collins

45,775

40,398

43,200

46,425

(6) Denver-Greeley

79,935

65,400

88,328

52,850

(9) Bonny Reservoir

37,300

32,000

35,000

37,200

(1/) Arkansas Valley

77,025

82,765

105,416

113,660

47,470

35,340

432,914

363,l30

Morgan

(15) San Luis Valley

Grand Totals

1/ Units

325,025

10, 11, 12, and 13.

320,556

�- 138 -

Mallards must have been continuing to enter Colorado from the north from
mid-December to early January because numbers of birds increased even though
an estimated 40,000 mallards were removed from the population during the
late point-system season.
The peak in numbers in the winter of 1968-69
was estimated to have occurred in mid-December with approximately 300,000
present in eastern Colorado (Funk 1969). The 433,000 observed in early
January 1970 was an increase of about 133,000 birds over the previous year.
As in previous years, mid-winter populations of ducks were estimated at
about 98 percent mallards.

Recovery

and Mortality

Rates and Migration

Routes

With changes in work assignments, updating of recovery cards has been delayed.
Changes in format on recovery cards obtained from the Bird Banding Lab also
contributed to delay in processing as IBM programs utilized previously were
rendered useless.
However, complete IBM tapes with all winter banding and
recovery information from 1963-64 through 1969-70 are expected from the
Bird Banding Lab very soon. Analysis of data could be completed more economicallyand
completely when the tapes arrive.
Thus, no updating of recovery,
mortality and harvest distribution data will be presented here. Hopefully,
the bulk of analysis can be completed in the next segment.

Wing Surveys
Information on age and sex ratios in the harvest was again obtained by both
Federal and State wing surveys.
The former survey covered the entire
hunting season while the latter covered only the point-system season.
These
data are covered in the point-system report (Funk et al. 1970) and will be
utilized for final reports.
No information on the surveys will be presented
in this report.

Hunting

Pressure

Surveys

Duck hunting pressure and harvest by unit was again estimated by study unit.
These figures are shown in Table 4 and are derived from 1969-70 waterfowl
harvest survey figures (Funk 1970). Data include harvest from both the
regular 1969 duck season and the 1969-70 late point-system season.
Although
species composition was not estimated through the questionnaire survey,
overall composition of the harvest was judged to be nearly 80 percent mallards.
As shown in Table 4, average bag per hunter in northern Colorado was similar
to 1968-69 estimates of between 7 and 8 birds with a lower estimate in the
Arkansas Valley.
Hunting pressure was up significantly from 1968-69 resulting
in a higher total kill than a year ago. High numbers of ducks present
coupled with the more lenient point-system regulation must have contributed
to enticing hunters to take part in the season.
Numbers of hunters per 100 birds censused was again utilized to compare
hunting pressure by study unit.
Pressure was indicated to have increased
significantly between 1968-69 and 1969-70 in the northeast section of Colorado.

�- 139 -

Table 4. Hunter pressure, harvest, and population survey data for comparison
between management units and areas, 1969-70 regular and late point-system
seasons.

Estimated
Duck
Huntersz'/

Estimated
Ducks
Bagged~J

Average
Bag per
Hunter

Number
Ducks
Censused~J

No.
Hunters Present
per 100 Ducks
Censused

1

2,219

15,107

6.8

20,588

10.8

2

2,760

15,827

5.7

36,427

7.6

9

974

5,935

6.3

34,767

2.8

Total

5,953

36,869

6.2

91,782

6.5

3

4,492

28,056

6.2

42,480

10.6

4

3,030

13,668

4.5

43,124

7.0

6

5,466

33,452

6.1

77,888

7.0

Total

12,988

75,176

5.8

163,492

7.9

10

758

5,755

7.6

12

1,840

12,769

6.9

13

1,028

7,913

7.7

Total

3,626

26,437

7.3

88,402

4.1

351

9,352

Area

]J

Northeast

North-Central

Southeast

San Luis Valley

15

1/ See Fig. 1 for management

47,470

unit locations.

2/ Funk 1970.

"jj Data from Table 3, average

of first three counts, except that only the
January 9 count was uS2d for the San Luis Valley.
~/ Inventory figures for Arkansas Valley usable only as total.

�- 140 -

The average of 6.5 hunters per 100 birds censused was double that of a year
ago. Bonny Reservoir was again indicated to have been low in pressure even
though wintering birds were abundant.
Here, lack of favorable harvest habitat is an important factor in keeping participation and total harvest low.
However, hunters in the area seemed to have as much success as those in
most other areas.
Pressure in the north-central area was indicated to have
been similar to that of 1968-69.

The Experimental

Point-system

Duck Season

Following the 1968-69 experimental late mallard drake season, which seemed
to be a success, a late experimental point-system season was granted for
parts of six Central Flyway states in 1969-70.
Results of the season were
presented in a cooperative report to the Central Flyway Technical Committee
(Funk et al. 1970). Areas included in the experiment were the Central Flyway portions of Montana, Wyoming, Colorado, and New Mexico, along with
western portions of South Dakota and Nebraska.
Length of the season was 23
days, covering four weekends, from December 13, 1969 through January 4, 1970.
Shooting hours were from sunrise to sunset.
Daily bag was on a point-system
of 40+ points with mallard drakes counting 10 points each toward the total.
All other species and mallard hens counted 40 points each.
Participants
were required to have a special free permit to hunt in their possession.
Purpose of the point-system regulation was to continue placing pressure on
mallard drakes while allowing "mistake" birds to be legal.
Number of permits issued in Colorado was 17,308, similar to the drake season
of 1968-69.
Number of permittees who hunted was estimated at 11,285 or 65%
of the total recipients of permits.
Total harvest was estimated at 43,000
birds.
Both of the latter figures are very similar to those from the previous year. Hunter performance survey results suggested that 86 percent of
the total harvest in Colorado during the late season was composed of mallards
(78 percent males plus 8 percent females).
This represents an estimated male
composition in the mallard bag of 91 percent.
This compares favorably with
the 89 percent estimate on mallard male-female composition in the bag from
wing surveys.
Thus, the late point-system season regulation seemed to be a
feasible method of directing harvest toward drakes.
Reaction of hunters to the point-system regulation was judged to be very
good, based both on results of performance checks and comments by hunters
on harvest questionnaires.
The challenge to the hunter was still present
in that he could maximize the size of his daily bag by selctively shooting
drake mallards.

�- 141 -

LITERATURE

CITED

Funk, H. D. 1969. Investigation of mallard management units of eastern
Colorado.
Colo. Div. of Game, Fish and Parks Game Res. Rept. Oct.
pp. 131-156.
Funk, H. D. 1970. Waterfowl kill survey.
Colo. Div. of Game, Fish and
Parks Game Res. Rept. Oct. In press.
Funk, H. D., R. M. Hopper, J. R. Grieb, D. Witt, G. F. Wrakestraw, T. Kuck,
D. E. Timm, and G. W. Merrill.
1970. Preliminary evaluation of the
1969-70 experimental point-system duck season within the High Plains
Mallard Management Unit of the Central Flyway.
Cent. Fly. Tech. Comm.
Rept. March.
11 pp.+ 12 tables and Appendix.
Grieb, J. R., H. D. Funk, R. M. Hopper, G. F. Wrakestraw, and D. Witt.
1969.
Preliminary evalution of the 1968-69 experimental mallard drake season
in Central Flyway portions of Montana, Wyoming and Colorado.
Cent.
Fly. Tech. Comm. Rept. March.
8 pp. + 16 Tables and Appendix.
Hopper, R. M., and H. D. Funk.
1970. Reliability of the mallard wing agedetermination technique for field use. J. Wildl. Mgmt. 34(2):333-339.

,

(

~I
Prepared by

f(,,_{&lt;

(
I

(:

'\
I

.

7
/
I

:(-~

Howard D. Funk
Section Chief, Small Game Research

��- 143 -

JOB PROGRESS

State

of

Project
Work

~C~O~L~O~RA~D~O~

October

REPORT

_

No.

W-88-R-ls

Migratory

Plan No.

4

Job No.

Job Title
Period

Covered:

Personnel:

Trapping
April

1, 1969

1970

and Banding

through

March

Bird

Investigations

3

Doves
31, 1970

Charles Hayes, Dale Horne, and Jack Randall, Bureau of
Sport Fisheries and Wildlife; Clait Braun, William
Carpenter, Howard Funk, Larry Green, Chet McCord, Steve
Steinert, Mike Watkins, and Gary Will, Colorado Division
of Game, Fish and Parks.

ABSTRACT

Efforts initiated in 1964 to trap and band samples of mourning doves
(Zenaidura macroura) in Colorado were continued in 1969.
Cooperative
federal and state efforts resulted in 3,294 birds being newly banded.
Of this total, 1,071 were immatures; 1,345 were adult males while 8v8
were adult females.
Assigned quotas were accomplished for the eastern
slope of the Rocky Mountains, but not for western Colorado or extreme
eastern Colorado.
Wing molt data from trapped immatures indicated that
hatching was bimodal with mid-May and mid-June being major hatching
periods.
A sample of 172 hunter killed birds in southeastern
Colorado
game a young-to-old
ratio of 2.4:1.
Wing molt data from harvested immatures in this sample indicated that most early-hatched
young migrated
from Colorado prior to September 1.

RECOMMENDATIONS

1.
2.

3.
4.

5.

6.

Collection of wings from hunter-killed
mourning dover should be
terminated.
Analyses of migration patterns and mortality rates of doves banded in
Colorado should not be conducted until at least 400 band returns are
available.
Efforts to trap and band birds in extreme eastern Colorado should be
expanded.
The p. S. Objective should be modified since sufficient band returns
upon which to base a harvest formula will not be available in the
foreseeable future.
Trapping and banding efforts in Colorado in conjunction with overall
objectives of theCentral Management
Unit should be continued.
Final analyses of banding data should be conducted by theBureau of
Sport Fisheries and Wildlife for the entire Central Management
Unit.

��- 145 -

MOURNING

DOVE TRAPPING

AND BANDING

Clait E. Braun

The mourning dove is the most important migratory game species in North
America in terms of hunting opportunity and total harvest.
In Colorado, well
over 300,000 doves are harvested each year, as they are avidly sought by
sportsmen
(Funk and Tully 1970).
In view of hunter interest and potential
harvest, Colorado initiated intensive studies on dove migration patterns in
1962 (Funk 1964).
In 1964 an active trapping and banding program on mourning
doves was started in cooperation with personnel of the U. S. Fish and Wildlife Service.
This program was intensified in 1967, and over 2,000 doves
each year have been banded since that time. Data presented in this report
are those collected in 1969 during the sixth year of this continuing
investigation.

P. S. OBJECTIVE
To develop

a harvest

formula for doves in Colorado.

SEGMENT
1.
2.

OBJECTIVES

To band mourning doves in Colorado for the purpose of obtaining migration, life history, and annual mortality data.
To obtain age and sex ratio data, wing molt information, and estimates
on harvest size from hunters through personal contact and mail surveys.

METHODS

AND MATERIALS

Doves were trapped in suitable locations (feedlots, fallow fields, and other
areas where waste grain was abundant) through use of cage-type wire funnel
traps (Funk 1965). Wheat (Triticum aestivum) was the primary bait used,
although millet (Panicum spp.), milo (Sorghum vulgare), and a commercial
hen scratch were used in Some locations.
All doves trapped were banded with
U. S. Fish and Wildlife Service bands (size 3A) and classified as to age
and sex (adults only) (Reeves et al. 1968). Observations were made as to
primary feather replacement on all immature birds in order to estimate
hatching dates (Allen 1963).
Efforts were made to trap and band 4,000 doves, with 1,000 to be banded in
West Slope areas; 2,000 along the East Slope of the Rocky Mountains, and
1,000 in extreme eastern Colorado, following guidelines established by the
U. S. Fish and Wildlife Service (Ruos 1968).

�- 146 DESCRIPTION

OF TRAPPING

AREAS

East Slope trapping areas were located primarily in farming areas where
cattle feedlots, trench silos, and farm homesteads were interspersed with
cultivated areas.
In the Fort Morgan area, the density of farms was lower,
with resultant larger fields and fewer trees. Much of the upland portion
of this locality was in native grass and water was not abundant.
A similar
situation existed at the Rocky Mountain Aresenal near Denver where much
of the area was in native grass.
Doves at this location fly substantial
distances to obtain grit along roadsides.
Trapping on the West Slope was concentrated near Craig and Meeker and was
conducted along roads and in livestock use areas. Vegetation was dominated
by sagebrush (Artemisia spp.) and Gambel's oak (Quercus gambelli), with
ranches being widely separated.

RESULTS AND DISCUSSION
Cooperative Federal and State trapping and banding efforts were intensified
in 1969, with 3,294 doves being banded.
Of this total, 2,663 were banded
by Division personnel.
Doves were banded at eight different locations along
the East Slope and at three different areas on the West Slope.
The banding
goal of 4,000 birds was not accomplished, as no doves were banded in
extreme eastern Colorado, and only 703 of the 1,000 quota were banded in
western Colorado.
However, the total of over 3,200 doves banded surpassed
numbers trapped in any other year of the study. Sex and age distribution
of birds banded by area is presented in Table 1.
It is apparent from Table 1 that adults were more readily trapped than
immatures, possibly due to higher nutritional needs associated with nesting,
and that adult males occurred more frequently in the trapped sample than
adult females.
This contrasts to the 1968 data, when numbers of adult males
and females trapped were similar, and immatures comprised 63.1 percent of
the total number trapped (Funk 1969).
In 1969, the largest banding effort
was made in June prior to the time when immatures comprised a large portion
of the population.
However, trapping effort in 1968 was more uniform, with
the major effort being expended in July and early August.
Examination of
the age ratio of doves trapped in 1969 revealed that in late July and
August (Denver and Fort Morgan), immatures substantially outnumbered adults
in the trap samples.
The reason why adult males were more readily trapped
than females are not precisely known, but this phenomenon may be related to
time of day and behavior of male and female doves. Males apparently nest
during mid-day and are more vulnerable to trapping in late afternoon (Funk
1965, 1969).

Hatching

Data

Wing molt data were recorded for 1,047 of the 1,071 immature doves banded
in 1969. Estimated hatching dates (Allen 1963) are presented in Table 2.

�- 147 -

Table 1Colorado,

Sex and age distribution

of mourning

doves banded by area in

1969.

Area

Adult Male
Number
Percent
Banded

Adult Female
Number
Percent
Banded

Irrnnature
Number
Percent
Banded

Total

Eastern Sl0Ee,
Rocky Mountains
Fort Collins

944

53.8

565

32.2

247

14.0

1,756

Denver

33

12.6

55

21.0

174

66.4

262

Longmont

40

58.0

16

23.2

13

18.8

69

Vineland

97

32.3

85

28.3

118

39.4

300

Rocky Ford

0

25

100.0

25

Fort Morgan

3

2.0

1

.6

148

97.4

152

Monte Vista

8

38.0

1

5.0

12

57.0

21

Salida

3

50.0

3

50.0

0

1,128

43.5

726

28.0

737

28.5

2,591

139

31.2

106

23.8

201

45.0

446

Durango

2

22.2

0

7

77 .8

9

Meeker

76

30.6

46

18.6

126

50.8

248

217

30.9

152

21.6

334

47.5

703

Total all Areas 1,345

40.8

878

26.7

1,071

32.5

3,294

Sub-total

0

6

Western Sl0Ee,
Rocky Mountains
Craig-Hayden

Sub-total

�- 148 -

Table 2.

Estimated

hatching

A ri1
17-23
24-30

Number
Hatching

Number
Hatching

dates for wild-trapped

mourning

doves,

1969.

1-7

8-14

Ma
15-21

22-28

29-June 4

166

106

127

5

8

12

69

5-11

June
12-18

19-25

26-2

3-9

10-16

17-23

24-30

87

126

92

91

67

66

25

0

Ju1

Examination of data in Table 2 indicates that doves hatched as early as late
April (April 23), with hatching continuing until late July.
It is probable
that few doves hatch earlier than late April in Colorado, but some hatch
later than indicated.
These birds were not represented in the trapped sample
as trapping was terminated on August 21. Peak hatching periods were midMay and mid-June in 1969. However, hatching peaks are not clearly delimited,
probably because of unequal trapping effort from June through mid-August.
Also, it is possible that doves hatched in Colorado molt at different rates
than those in Indiana where the technique for estimating hatching dates
from primary molt progression was developed.
Upon tabulating the number of immature doves molting primaries 1 through 10,
it was apparent that there was a constant and rapid turnover of young doves
available for trapping in the populations sampled.
Over 75 percent (77.0)
of the immatures trapped were molting primaries 1 or 2, or had not shed
primary 1. In contrast, less than 3 percent (2.7) had molted more than 6
or 7 primaries, while no birds were trapped which were molting primaries 8
through 10. This strongly suggests that young doves rapidly left nesting
areas and migrated away from trapping sites. Where these birds congregated
and migrated to after hatching is not known.

Harvest
No effort was made to collect wings from throughout Colorado during the
1969 hunting season.
Wings of 172 birds harvested September 2 and 3 in
southeastern Colorado were ava~lab1e for analysis.
Of this total, 51 were
from adult birds and 121 were from irnrr.atures,
giving a young-to-o1d ratio
of 2.4:1.
Approximately 75 percent (75.2) of the young harvested had
molted 5 or fewer primaries, indicating they were less than 63 days of age.
This supports data collected in earlier years of the study which indicated
that most early-hatched young migrated from Colorado prior to September 1.

�- 149 -

Harvest data obtained through use of a mail survey of hunters after the
hunting season indicated that 25,119 hunters harvested 323,773 doves in
Colorado in 1969 (Funk and Tully 1970). Less than 30 band returns were
received from doves shot in 1969 that were banded in the same year.
While
sample sizes are too small for meaningful conclusions, it is evident that
doves banded in Colorado have a low hunting mortality. However, a substantial
amount of the known hunting mortality occurs in Colorado.
Because of low
band recoveries, meaningful data on migration and annual mortality are not
available.

LITERATURE
Allen, J. M.
Indiana.
4 p.

CITED

1963. Primary feather molt rate of wild immature
Ind. Dep. Conserv., Game Res. Sect. Circ. No.4,

Funk, H. D. 1964. Dove migration investigation.
Parks, Game Res. Rep., Fed. Aid Proj. W-88-R.

doves in
Indianapolis.

Colo. Div. Game, Fish and
Oct. p 89-113.

1965. Trapping and banding doves.
Game Res. Rep., Fed. Aid Proj. W-88-R.

Colo. Div. Game, Fish and Parks,
Oct. p. 161-169.

1969. Trapping and banding doves.
Game Res. Rep., Fed. Aid Proj. W-88-R.

Colo. Div. Game, Fish and Parks,
Oct. p. 157-162.

______, and R. J. Tully.
1970. Colorado small game hunter harvest survey,
1969. Colo. Div. Game, Fish and Parks, Fort Collins.
Mimeo. Rep. 35 p.
Reeves, H. M., A. D. Geis, and F. C. Kniffin.
1968. Mourning dove capture
and banding.
U. S. Dept. Interior, Bureau Sport Fisheries and Wildl.,
Spec. Sci. Rept. Wildl. No. 117, 63 p.
Ruos, J. 1968. Mourning dove banding needs, 1969-71.
Bureau Sport Fisheries and Wildl., Unpubl. Rept.

U. S. Dept. Interior,
13 p.

%' -r~?It?rZttun}
£

Prepared by._. ,,/tze ?
Clait E. Braun, Assistant

Wildlife

Researcher

��October

- 151 -

JOB PROGRESS

State

of

Project
Work

~C~O~L~O~RAD~~O

Plan No.

Period

Covered:

Personnel:

Migratory

4
------~--------------

Job Title

Band-tailed
April

REPORT

_

W-88-R-15

No.

1970

Pigeon

Bird

Job No.

Investigations

4

Investigations

1, 1969 through March

31, 1970

Jack Randall, Bureau of Sport Fisheries and Wildlife;
personnel of the U. S. Forest Service; Clait E. Braun,
William Carpenter, Velma Fredrickson,
Howard Funk, Chet
McCord, Richard McDonald, Marie Vendeville,
Ken Wagner,
Mike Watkins, Wildlife Conservation
Officers, and other
personnel of the Colorado Division of Game, Fish and
Parks.

ABSTRACT

Investigations
of band-tailed
pigeons (Columba fasciata) were initiated
in Colorado in 1969 in conjunction with the Cooperative Four Corners
States (Arizona, New Mexico, Utah and Colorado).
Major emphasis was placed
on delimiting present distribution
in respect to historic distribution,
ascertaining relative abundance, trapping and banding samples of pigeons
throughout the State and documenting breeding and nesting phenology.
Band-tailed pigeons were found to be distributed
throughout the forested
mountainous areas of Colorado from early May into October.
Estimated
numbers of pigeons present were not calculated but relative abundance was
greater in southern and southwestern Colorado than in northern and northwestern Colorado.
Abundance and distribution
appeared to be related to
presence of native foods and cultivated grain crops.
No real difference
could be detected between the 1969 survey of distribution
and abundance
and surveys conducted from 1946 to 1957.
Pigeons were trapped at five
different locations in 1969 and 1,600 birds were newly banded.
Data from
collected birds indicated that breeding and nesting were initiated prior
to June 25 and continued until at least mid-August.
Some data were obtained that indicated band-tails in Colorado were capable of multiple
nesting attempts and of producing up to two eggs per clutch.

RECOMMENDATIONS
1.
2.
3.

Efforts should be increased to trap and band wild band-tailed
pigeons
(especially immatures) over wider geographical
areas of the State.
An experimental hunting season for 1970 should be requested of the
Bureau of Sport Fisheries and Wildlife.
Plans should be formulated for an aviary study to document molts and
plumages from the immature to adult age classes.

��- 153 -

BAND-TAILED

PIGEON INVESTIGATIONS

C1ait E. Braun

Band-tailed pigeons, wild, native Co1umbids, occur only in North and Central
America.
Two populations are presently recognized north of Mexico, with the
coastal group (C. f. monilis) occurring principally west of the Sierra and
Cascade Mountai~s of California, Oregon, Washington, into Central British
Columbia.
The interior population (f. f. fasciata) is seasonally resident
in the forested mountains of Utah, Colorado, New Mexico, Arizona, southward
to Guatemala (Amer. Ornithol. Union 1957).
Systematic studies of regional or State populations of band-tails are
lacking, with most published references referring to distribution, finding
of a nest, or trapping and census techniques.
Notable exceptions are the
nesting studies of Glover (1953) and MacGregor and Smith (1955); general
life history work of Neff (1947) and Smith (1968); mortality studies of
Wight et al. (1967) and Silovsky (1969); and the behavior work by Peeters
(1962). Of the important contributions to understanding the ecology of bandtailed pigeons, only the general life history information presented by Neff
(1947) referred to the interior population.
Consequently, efforts were
initiated by the Four Corners States (Arizona, Colorado, New Mexico and
Utah) in 1967 to conduct cooperative studies on this little known population.

P. S. OBJECTIVE
To investigate life history and ecological facts pertinent to band-tailed
pigeons in Colorado as follows: range; relative numbers; relation between
numbers of birds present by area and such physical factors as weather, food,
vegetative cover, elevation, and salt; breeding and nesting chronology;
migration patterns and chronology; harvest patterns; and mortality rates.

SEGMENT
1.
2.
3.
4.

OBJECTIVES

To investigate distribution and relative numbers of band-tailed pigeons
in Colorado by area.
To investigate the correlation between abundance and distribution of
bandtails and physical features of the environment.
To investigate the breeding and nesting chronology of band-tails in
Colorado.
To trap, band, age, and sex as many birds as possible in selected
areas of Colorado to investigate migration patterns and chronology of
Colorado band-tailed pigeons throughout their range.

REVIEW OF LITERATURE
Much of the published literature concerning band-tailed pigeons relates to
general observations on distribution and the finding of a nest.
Of over 150
references examined, less than 20 were the result of organized scientific
study.

�- 154 Census techniques for band-tailed pigeons have been discussed by Sisson
(1968), Keppie (1970), and Keppie et al. (1970); while Miller and Wagner
(1955), and Silovsky et al. (1968) reported on sex and age characters.
Trapping techniques for band-tails have been mentioned by a number of workers,
among them Wooten (1955), Mace and Batterson (1961), Drewien et al. (1966),
and mortality rates have been examined by Wight et al. (1967) and Silovsky
(1969).
Intensive studies on nuptial behavior (Peeters 1962) and nesting
(Glover 1953; MacGregor and Smith 1955) have been conducted only in California.
Houston (1963) presented preliminary results of a study on the
reproductive ecology of this species in California.
Work by Neff (1947)
and Smith (1968) have contributed to the overall knowledge of band-tailed
pigeons.
Important contributions to an understanding of parasitic load
in this species have been made by Stabler and co-workers (1950, 1951, 1963)
and Sileo and Fitzhugh (1969).
Of 27 references concerning band-tailed pigeons in Colorado, 21 refer to
status or distribution within the State and are summarized by Bailey and
Niedrach (1965). Five of the remaining articles refer to parasites in wild
Colorado Columbids including band-tailed pigeons (Stabler 1950, 1951;
Stabler et al. 1950; Stabler and Matteson 1950; Stabler and Holt 1963). The
remaining brief reference pertains to nesting (Neff and Niedrach 1946).

METHODS

AND MATERIALS

Mail surveys of Wildlife Conservation Officers, U. S. Game Management Agents,
and U. S. Forest Service personnel were conducted in order to obtain information on past and present distribution of band-tailed pigeons in Colorado
(Appendix).
All persons contacted through the mail survey were also asked
to report all sightings of band-tails on standardized monthly observation
forms (Appendix).
In addition, project personnel systematically reported
their observations of pigeons on standardized cards (Appendix).
All
sightings of 100 or more pigeons were to be reported directly by telephone
to the Game Research Center in Fort Collins.
Efforts were made to trap pigeons wherever they were located feeding in
flocks larger than 50 birds.
Pigeons were trapped primarily through use of
cannon nets and modified cage-type wire funnel traps. One 30- by 75-foot
cannon net with l~-inch mesh was used with three cannons as described by
Dill (1969). Wire cage-type funnel traps (Funk 1965) with enlarged funnels
were utilized in some instances where space was limited or the cannon net
was unavailable.
Two 7- by 30-foot, 2 3/8-inch mesh mist nets were used
for 3 days following instructions of Reeves et al. (1968). All pigeons
trapped were separated into adult or immature classes (Silovsky et al. 1968),
and the sex of most adults was determined by cloacal inspection (Miller and
Wagner 1955). Following age and sex determination, pigeons were banded
with U. S. Fish and Wildlife Service bands (size 5), and released.
Barley (Hordeum vulgare) was most successfully used as bait for pigeons,
although wheat (Triticum aestivum) and milo (Sorghum vulgare) were used in
certain situations.
A few pigeons were trapped in orchards by using mist
nets placed adjacent to cherry (Prunus avium) trees.

�- 155 -

Band-tailed pigeons lost in trapping operations, those systematically
collected, and those shot by orchard growers near Hermosa were immediately
frozen and then transported to Fort Collins.
Upon processing, measurements
were taken of certain external characteristics to the nearest millimeter,
while gonads were measured to the nearest tenth of a millimeter.
Weights
were recorded to the nearest tenth of one gram. Status of the crop glands
was recorded with those crops showing glandular development being excised
and preserved in buffered formalin.
Upon dissection, gonads were measured,
excised, and preserved in buffered formalin.
Crop contents were dried and
stored in manila envelopes with gizzard contents being preserved in buffered
formalin.
All helminths found were preserved in either a 95-percent alcohol
and 5-percent glycerin solution (nematodes) or AFA (cestodes).
All carcasses
were saved for future mercury and fat analysis.
Usable skins were donated
to Colorado State University.
Standardized data sheets were utilized for
each bird (Appendix).

DESCRIPTION

OF TRAPPING

AREAS

Pigeons were trapped at locations near Salida, Rye, Durango, Monte Vista,
and Morrison in 1969. With two exceptions, trapping sites were located
along the edges of either newly planted (Salida, Rye, and Durango) or
harvested (Monte Vista and Morrison) grainfields (barley and wheat) located
along the foothills and in intermountain valleys.
These locations were
characterized by having an abundance of immediately adjacent roosting or
perching sites in trees or shrubs.
Trees and shrubs most commonly used were
ponderosa pine (Pinus ponderosa), narrowleaf cottonwood (Populus augustifolia),
and Gambe1's oak (Quercus gambe1i).
One trapping site near Durango was
located in an alfalfa (Medicago sativa) field along a stream course, while
the other exception, near Hermosa, was in a commercial cherry orchard.

RESULTS

AND DISCUSSION

Distribution
The distribution of band-tailed pigeons in Colorado prior to the advent of
settlement is poorly known.
Only a few scattered references concerning
pigeon sightings in Colorado are available prior to 1946. In 1945, general
studies of pigeons in Colorado were initiated under the auspices of the Game
and Fish Department which continued on a sporadic basis until the early
1950's.
As a result of these studies, the first general map of band-tail
distribution in Colorado was prepared (Neff and Culbreath 1947). Status
and distributional surveys were conducted annually from 1946 until 1957,
after which time no formal reports were prepared.
Of the reports prepared,
only those by Neff and Culbreath (1947), Kinghorn (1947), and Matteson
(1948) included distribution maps.
Sufficient data were obtained from the mail survey to plot the recent distribution of pigeons in Colorado (Fig. 1). Upon comparing the distribution of
pigeons in 1969 with the map prepared by Neff and Culbreath (1947), it is
apparent that only minor changes have occurred.
The most notable change is

�- 156 ~

that pigeons now occur north of the Cache 1a Poudre River in Larimer County,
whereas the earlier work indicated no sightings in this area.
It is probable
that the slight differences in distribution between 1946 and 1969 are a
function of numbers of observers and access.
Essentially, no real changes
in distribution of band-tailed pigeons have occurred since 1946.
Pigeons in 1969 were found to be widely distributed throughout the mountainous
areas of Colorado from the foothills of the Eastern Slope of the Rocky
Mountains, along the periphery of most of the intermountain valleys and
parks, to the forested mesas, plateaus, and canyons of the western and southwestern portion of the State.
They are extremely uncommon above tree line
(11,600 ft), and apparently are absent in North Park, South Park, and portions
of the Gunnison Basin as well as the sagebrush (Artemisia spp.)-pinyon
(Pinus edulis)-juniper(Juniperus
spp.) country in northwestern Colorado.

Abundance
At present, no adequate census techniques are available for estimating
numbers of band-tailed pigeons.
Studies in Oregon (Sisson 1968; Keppie 1970)
have been conducted on the use of an audio-index for census of this species.
Techniques developed in these studies have been recently proposed for widespread use in censusing pigeons (Keppie et al. 1970).
From reports of pigeon sightings it was possible to obtain a general
impression as to relative numbers of pigeons.
Reported observations of
pigeons were most abundant from the Glenwood Springs, Uncompahgre Plateau,
San Juan Basin, South Fork, Salida, and Rye-LaVeta areas.
Lesser numbers
of sightings were reported from the Evergreen-Morrison,
Buford, and Hot
Sulphur Springs areas.
Outside of these areas, reported sightings were
sparse.
While it is difficult to relate number of sightings to actual
densities of birds because of number of observers, access, time of year,
etc., it is possible to make a general assessment of pigeon abundance.
Apparently, band-tailed pigeons are most abundant south of a line from Denver to Glenwood Springs and Grand Junction.
While distribution and abundance
south of this line are not even, north of this line pigeon abundance and
distribution is even more discontinuous.
Reasons for the uneven distribution and abundance of pigeons in Colorado
are not known.
It is possible that the presence or absence of native foods
(berries and acorns) is most responsible in determining where pigeons occur
and how many can be supported.
Forested areas of northern Colorado are
noted for their lack of substantial numbers of species of berry and mast
producing plants.
Intermountain areas of this region are relatively high
in elevation, they have short growing seasons, and consequently, few cereal
grains are cultivated.
Pigeon distribution in this region is sparse and
numbers are apparently low. In contrast, forested areas of central and
southern Colorado have a variety of important berry (Sambucus spp., Prunus
spp., Lonicera spp., Crataegus spp.) producing bushes in addition to an
abundance of Gambel's oak (acorns).
Scattered throughout the lower valleys
of this region are farms which usually produce some cultivated grains.
Pigeons in this area are more widespread and apparently more abundant.
Whether
there is a cause-and-effect
relationship between food, abundance, and distribution of pigeons is not known, although general observations support
this possibility.

�COLORADO
MOFFAT

ROVTT

SEOGWICIC

LOGAN

WELl)

PHILLIPS

MORGAN
I rVMA

WASHINGTON

RIO

BLANCO

ELBERT

I(tTCARSON

LINCOLN

MESA

I-'

VI

CHEYENNE

A'

\

I CROWLEY

"

I KIOWA

I

SAGVACHE
BENT

J.AS ANIMAS

Fig. 1.

Band-tailed pigeon distribution, 1969.

I!'ROWERS

I SAC ••••

�- 158 Upon comparing locations and numbers of pigeons reported in 1969 with those
mentioned in early reports (Neff and Culbreath 1947; Kinghorn and Neff 1948;
Matteson 1948, 1949, 1950; Neff, 1951, 1952; U. S. Fish and Wildl. Servo
1954, 1955, 1957; Buller 1957), it is apparent that no real change has
occurred in numbers or distribution of pigeons in Colorado.
Indications
are that band-tails use the same general areas every year in approximately
the same numbers, with exact location used within a given area varying each
year.
The observed variation is probably related to food availability and
activities of the observers.
From what little meaningful data that are
available, it is concluded that pigeon numbers in Colorado have not noticeably
changed since 1946.
Relating pigeon abundance to physical features of the environment is difficult, if not impOSSible, without detailed study of the various habitats used.
Apparent relationships between pigeon distribution and abundance with presence or absence of native foods and cultivated crops have been previously
mentioned.
Upon listing reported observations of pigeons in 1969 by habitat
type (cropland, oak brush, pinyon-juniper, river bottom, orchard, ponderosa
pine, spruce-fir (Picea-Pseudotsuga &amp; Abies) forest, and alpine) and time
period (May through November) several generalizations can be made.
Upon
initially arriving in Colorado (early May) pigeons are most frequently seen
in cropland areas.
This pattern continues until early July when pigeons
are more frequently observed in river bottoms, orchards, ponderosa pine and
spruce-fir forests.
In August, pigeons continue to be seen in ponderosa
pine and spruce-fir forests, but sightings in areas dominated by Gambel's
oak and cultivated crops are most frequently reported.
In September, continuing into November, pigeon sightings are most frequently reported from
oak brush and forested areas.
It is doubtful that reported sightings give
an accurate picture of the habitats used by band-tailed pigeons.
Obviously,
pigeons in cultivated areas and in other easily accessible locations have a
better chance to be observed and counted than those in heavily forested or
more remote sites.

Banding
Prior to 1969, banding of band-tailed pigeons in Colorado was sporadic, and
only 557 had been banded (1945 through 1968). Of this total, 490 were
banded between 1945 and 1949, while 67 were banded between 1951 and 1968.
Through 1968, only one recovery (Mexico) had been reported.
Much of the
banding in earlier years was near Perry Park north of Colorado Springs,
near Westcliffe and west of Salida.
Intensive efforts were conducted in 1969 to trap and band pigeons in as
many areas as possible.
As a result of these efforts, 1,600 pigeons were
newly banded in five localities.
No recaptures of birds previously banded
in Colorado or elsewhere were obtained.
Results of trapping operations are
given in Table 1.
Information concerning the sex of each adult banded was recorded, but due
to a combination of factors these data are considered unreliable.
Factors
involved were the inexperience of the banding crews and large numbers of
birds to process quickly (up to 200 at one time). Sex of many birds was

�- 159 determined by cloacal inspection (Miller and Wagner 1955), but reliability
of this technique under field conditions where several persons were involved
was questionable.
Observations appeared to indicate that percentage of
each sex trapped varied with time of day. Males
appeared to predominate
before 10:00 A. M., and after 4:00 P. M., MDT, with females being more common
from 10:00 A. M. to 4:00 P. M.

Table 1.

Number of band-tailed

pigeons banded by area in Colorado,

Totals

Es timated
No. of
Birds Using
Trapping
Sites

2.9

308

500+

4

.8

524

600+

75.0

85

25.0

340

400+

144

35.6

261

64.4

405

500+

5

21. 7

18

78.3

23

300+

1,223

76.4

377

23.6

1,600

2,300+

Adults
No.
Banded
Percent

Immatures
No.
Banded
Percent

Area

Dates

Pueblo

June 25-28

299

97.1

9

Salida

June 25-30

520

99.2

Durango

July 2-31

255

Monte
Vista

Aug. 5Sept. 8

Morrison

Aug. 30-31

TOTALS

1969.

Upon examination of data presented in Table 1, it is apparent that percent
immatures trapped increased from late June to late August.
The immatures
trapped in June are difficult to explain as it apparently takes approximately 50-60 days for incubation and rearing of the young to flight stage
(Neff and Culbreath 1947). This would indicate that immatures flying in
late June were produced by parents that initiated nesting in early May.
It is possible that these young were produced in Colorado, or somewhere
south of Colorado.
The capture of flying young in June may not be unusual
as Matteson (1948, 1949) reported trapping one innna.tureon June 20, 1948
(total of 66 birds handled in June) and another on May 28, 1949 (total of
101 birds handled in May).
These birds were trapped in the Perry Park and
Westcliffe areas.
The high percentage of immatures trapped near Monte Vista in August is of
interest and may indicate that this trapping site was a staging area for
young pigeons produced over a large area. While number of pigeons banded
at Morrison was inadequate, it is possible that this flock was also comprised

�- 160 -

mostly of immatures.
This location could be another staging area for
immature birds.
While numbers of birds (Table 1) present at each trapping
site are estimates (probably conservative), it is apparent that once flocks
of pigeons are located, a substantial percentage of the flock can be readily
captured.
With the exception of Morrison, initial estimates of flock size
before trapping was initiated were approximately one-half or less of the
numbers actually trapped.
Therefore, observed flock size may have no correlation with actual number of birds using a particular area.

Mortality
Insufficient data were obtained to estimate mortality rates of band-tailed
pigeons banded in Colorado.
While Arizona and New Mexico conducted experimental hunting seasons on pigeons for the second consecutive year in 1969,
the season remained closed in Colorado.
Seasons in these two States were
conducted in limited areas from October 11 through 19, with bag and possession limits of 5 and 10. No pigeons banded in Colorado were reported shot
in Arizona or New Mexico in 1969. Only one recovery was reported through
August 1970 from the 1969 banding effort.
This bird was taken in Mexico
in June 1970.

Breeding

Phenology

Data on breeding phenology were obtained through crop and gonadal inspection
of birds shot by cherry growers near Hermosa, those accidentally lost in
trapping operations, and those systematically collected.
In all, 144 birds
were available for examination.
Of this number, 126 were adults (80 males
and 46 females), while the remaining 18 were immatures.
The sample included
birds taken from June 25 through October 6, but most (109) were from
July 9-13.
Data concerning crop activity and gonadal condition were tabulated but analyses were not completed during the segment.
Cursory examination of the
gonadal data indicates males and females were in breeding condition (enlarged
testes and egg follicles) from late June through mid-August.
Preliminary
results also indicate that some females were capable of laying at least two
eggs in quick succession.
There is some evidence, as yet inadequate, to
suggest that early hatched young of the year may be capable of breeding late
in their first year. Unfortunately, no collections were available from May
and early June, while collections after mid-August were inadequate to
accurately assess breeding phenology.
Even without these data, it is
apparent that pigeons breed and nest in Colorado from early June until sometime in October as birds collected in June and September had "active crops"
indicating that young were being fed. In all, crop glands of 33 of 126
adults for which data are available were lactating at time of death.
If
50-60 days are necessary for incubation and rearing of the young, it is
possible for pigeons in Colorado to nest twice, provided they are in breeding
condition when they arrive.

�- 161Fragmentary data collected in earlier work in Colorado (Neff and Culbreath
1947; Kinghorn and Neff 1948; Matteson 1948, 1949; Neff 1951) substantiates
the 1969 data as females with developed eggs in their oviducts have been
collected in early June, reports of nests with two eggs are on record, and
reports of eggs and young in nests are available from late August and September. Similar fragmentary information is available for New Mexico and
Arizona as up to two young have been found in nests from late February
through late September (Neff 1952; U. S. Fish and Wildlife Service 1954).
It is reasonable to conclude that the interior group of band-tailed pigeons
are multiple nesters and may produce up to two eggs per clutch.

Wing Measurement

and Molt Data

Measurements of primaries, rectrices, total length, carpal length, and bill
length were taken of all pigeons available for necropsy.
The major reason
for taking these measurements, other than to describe the morphology of
pigeons living in Colorado, was to examine the possibility of sexing and
aging birds from one or two wing measurements.
These data were tabulated
but not analyzed as sufficient sample sizes were not available.
It is
apparent that many of the measurements (rectrices, bill length, and some
primary lengths) will be useless for age and sex determination.
However, it
appears that a combination of carpal length and length of primaries 10
through 7 will be useful in separating immatures and adult males and females.
Once data from over 300 birds of known sex and age are available, final
analyses of the measurement data will be initiated.
Prior to dissection of collected birds, each adult specimen was closely
examined as to appearance and coloration of the forehead, upper breast, and
distinctness of the white crescent, and classified as to male and female.
Upon dissection, gonads were examined and the actual sex of each bird was
compared with the classification based on plumage data. Comparative data
were available for 119 adults (76 males and 43 females).
Of the 76 adult males compared, two (2.6 percent) were sexed incorrectly
while five (11.6 percent) of 43 females were classified as males from
plumage examination.
Total error in classifying pigeons to sex from plumage
examination alone was 5.9 percent.
While sample sizes are small, it is
apparent that an experienced observer can accurately sex pigeons from
plumage examination.
Use of this technique on live birds would bias sex
data in favor of males.
Systematic observations of primary feather replacement, except on a few
live-trapped birds and all birds collected, were not recorded in 1969.
Date concerning primary molt were available from 142 adults and 166 immatures.
Although samples are small, preliminary indications are that most adults
start replacing primaries 1 and 2 in early August (August 10), a few may
molt primary feathers in early July, with some birds replacing several primaries out of the normal sequence (1~10)
in late August and September.
Apparently, molting of primaries by adults is not sequential.

�- 162 Of the 166 irnmatures for which data are available, 59 had not shed primary
1, 56 were molting PI, 40 were molting P2, 5 were molting P3, 4 were molting
P4, and 2 were molting P5. These birds were trapped or shot between July 9
and October 6, with most (134) being trapped between August 10 and 17.
Hatching dates for these birds were estimated using unpublished data collected by H. M. Wight and his students (Personal communication, H. M. Wight).
Numbers of days used to back date for each primary class were: 0 = 36,
1 = 43, 2 = 54, 3 = 72, 4 = 81, 5 = 96. Number of days used are based on
the premise that it takes 30 days from hatching to fledging plus 13 days
until P1 is shed. It cannot be assumed that all birds not yet molting P1
are only 30 or 43 days old, so the number of 36 days was arbitrarily chosen
for the 0 class.
Unpublished data from Oregon (Personal communication,
H. M. Wight) indicate that birds kept in an aviary shed Pl-43 days, P2-54
days, P3-72 days, P4-8l days, and P5-96 days after hatching.
One bird
banded in Colorado on August 14 molting no primaries was retrapped on
September 10 when it had molted P2. Thus, it took 27 days to molt from 0 to
2 in this instance.
Estimated hatching dates for Colorado trapped birds are
presented in Table 2.

Table 2.

Estimated

hatching

Number
Hatching

band-tailed

pigeons,

1-7

8-14

June
15-21

22-28

29-Ju1y

2

9

4

12

22

52

6-12

13-19

July
20-26

27-2

3-9

51

5

2

1

&lt;May

Number
Hatching

dates for wild-trapped

31

5

August
10-30

0

1969.

5

31

1

It is apparent from Table 2 that hatching in 1969 was spread over a wide
time period (May 28 to August 31). The majority of the trapped immatures
for which data were available hatched between June 22 and July 12. Unfortunately, all but 27 of the 166 birds were from the Monte Vista area. The
nine birds from Durango all hatched prior to June 7. Only two of the Monte
Vista birds and none of the 18 Morrison birds hatched prior to June 7. It
is probable that the sample does not represent hatching dates for all
pigeons hatched in various areas in 1969. It is also possible that early
hatched pigeons were not available for trapping as only 11 of the 166 birds
examined had molted P3+.

�- 163 -

Parasitic

Load

Information on helminths was available for 144 band-tailed pigeons (126
adults and 18 immatures) necropsied.
Nineteen (18 adults, 1 immature) of
the 144 had helminth infections.
The infection rate of 13.2 percent is low,
and only one bird (adult female) was considered to be heavily infested.
Eleven of 46 adult females contained helminths, while only 7 of 80 adult
males were infested.
If adequate samples were available, this difference
would be significant.
Nematodes were found in 5 birds (3 males, 2 females)
while 14 contained cestodes and one (adult female) had both types of worms.
No adverse effects of the observed parasitism were noted. All worms found
were preserved for later identification.

LITERATURE

CITED

American Ornithologist's Union.
1957. Check-list of North American
5th ed. Lord Baltimore Press, Inc., Baltimore.
691 pp.
Bailey, A. M., and R. J. Niedrach.
1965.
Museum Nat. Hist. Vol. II. 895 pp.

Birds of Colorado.

birds.

Denver

Buller, R. J. 1957. Survey of band-tailed pigeon populations in Arizona,
Colorado, New Mexico, and Utah.
1957. U. S. Fish and Wi1dl. Serv.,
Branch of Game Mgmt., Albuquerque, New Mexico.
15 pp.
Dill, H. H. 1969. A field guide to cannon net trapping.
Bureau Sport
Fisheries and Wi1d1., U. S. Fish and Wi1d1. Serv., Div. Wi1dl. Refuges,
Minneapolis, Minn. Mimeo Rept., 21 pp.
Drewien, R. C., R. J. Vernimen, S. W. Harris, and C. F. Yocum.
1966.
weights of band-tailed pigeons.
J. Wildl. Mgmt. 30(1):190-192.

Spring

Funk, H. D. 1965. Trapping and banding doves.
Colo. Dept. Game, Fish and
Parks, Game Res. Rept., Fed. Aid Proj. W-88-R.
Oct. p. 161-169.
Glover, F. A.
fasciata)

1953. A nesting study of the band-tailed pigeon (Columba f.
in northwestern California.
Calif. Fish and Game 39(3):397-407.

Houston, D. B. 1963. The reproductive ecology of the band-tailed
J. Colo.-Wyo. Acad. Sci. 5(4):52.
Columba fasciata.

pigeon,

Keppie, D. M. 1970. The development and evaluation of an audio-index
technique for the band-tailed pigeon. M.S. ThesiS, Oregon State Univ.,
Corvallis.
_____ , H. M. Wight, and W. S. Overton.
1970. A proposed band-tailed pigeon
census.
Trans. No. Am. Wi1d1. and Nat. Res. Conf.
35:In Press.

�- 164 Kinghorn, R. G. 1947. Band-tailed pigeons.
Progress Rept., Fed. Aid Proj. W-37-R.

Colo. Dept. Game and Fish.
Oct. pp 1-4.

_____ , and J. A. Neff.
1948. Status of the band-tailed pigeon in Colorado, season of 1947. Colorado Dept. Game and Fish.
13 pp.
Mace, R. U., and W. M. Batterson.
1961. Results of a band-tailed pigeon
banding study at Nehalem, Oregon.
Proc. Ann. Conf. Western Assoc.
State Game and Fish Coroms. 41:151-153.
MacGregor, W. G., and W. M. Smith.
1955. Nesting and production of the
band-tailed pigeon in California.
Calif. Game and Fish. 41(4):315-326.
Matteson, C. P. 1948. Band-tailed pigeons.
Colo. Dept. Game and Fish,
Progress Rept., Fed. Aid Proj. W-37-R. Oct. pp 18-23.
1949. Band-tailed pigeons.
Rept., Fed. Aid Proj. W-37-R.

Colo. Dept. Game and Fish, Progress
Oct. pp 33-37.

1950. Band-tailed pigeons.
Report, Fed. Aid Proj. W-37-R.

Colo. Dept. Game and Fish, Progress
Oct. pp 39-40.

Miller, W. J., and F. H. Wagner.
1955. Sexing mature Co1umbiformes
cloacal characters.
Auk 72(3):279-285.
Neff, J. A.
pigeon.

by

1947. Habits, food and economic status of the band-tailed
U. S. Fish and Wi1d1. Serv., No. Am. Fauna 58,76 pp.

1951. Inventory of band-tailed pigeon populations in Arizona, Colorado and New Mexico, 1951. U. S. Fish and Wi1d1. Serv., Denver, Mimeo
Rept.
37 pp.
1952. Inventory of band-tailed pigeon populations in Arizona,
Colorado, and New Mexico, 1952. U. S. Fish and Wi1d1. Serv., Denver,
Mimeo Rept.
26 pp.
_____ , and J. C. Culbreath.
1947. Status of the band-tailed pigeon in
Colorado, season of 1946. Colo. Dept. Game and Fish, Mgmt. Div., Fed.
Aid Sect. Rept.
24 pp.
_____ , and R. J. Niedrach.
1946. Nesting
Colorado.
Condor 48(2):72-74.

of the band-tailed

Peeters, H. J. 1962. Nuptial behavior of the band-tailed
San Francisco Bay area. Condor 64(6):445-470.

pigeon

in

pigeon in the

Reeves, H. M., A. D. Geis, and F. C. Kniffin.
1968. Mourning dove capture
and banding.
U. S. Dept. Interior, Bureau Sport Fisheries and Wi1d1.
Spec. Sci. Rept. Wi1d1. No. 117, 63 pp.
Sileo, L., Jr., and E. L. Fitzhugh.
1969. Incidence of trichomoniasis
the band-tailed pigeons of southern Arizona.
Bull. Wi1d1. Disease
Assoc. 5:146.

in

�- 165 Silovsky, G. D. 1969. Distribution and mortality of Pacific Coast bandtailed pigeons.
M. S. Thesis, Oregon State Univ., Corvallis.
_______
, H. M. Wight, L. H. Sisson, T. L. Fox, and S. W. Harris.
1968.
Methods for determining age in band-tailed pigeons.
J. Wildl. Mgmt.
32(2):421-424.
Sisson, L. H. 1968. Calling behavior of band-tailed pigeons in reference
to a census technique.
M. S. Thesis, Oregon State Univ., Corvallis.
Smith, W. A. 1968. The band-tailed
and Game 54(1):4-16.

pigeon in California.

Stabler, R. M. 1950. Trichomonas gallinae
J. Colo.-Wyo. Acad. Sci. 4(2):83.

in Columbid

Calif. Fish

birds in Colorado.

1951. A survey of Colorado band-tailed pigeons, mourning doves,
and wild common pigeons for Trichomonas gallinae.
J. Parasitol.
37 (5) :470-472.
_____ , and Portia A. Holt.
1963. Hematozoa from Colorado birds.
and doves. J. Parasitol. 49(2):320-322.
____

, and C. P. Matteson.
1950. Incidence
Colorado mourning doves and band-tailed
36(6):25-26.

I. Pigeons

of Trichomonas gallinae
pigeons.
J. Parasitol.

in

_____ , Phyllis S. Limberg, and C. P. Matteson.
1950. Blood parasites
Colorado band-tailed pigeons.
J. Parasitol. 36(6):25.

in

U. S. Fish and Wildlife Service.
1954. Survey of band-tailed pigeon
populations in Arizona, Colorado and New Mexico with notes on Utah,
1953. U. S. Fish and Wildl. Serv., Branch of Game Mgmt., Albuquerque,
New Mexico.
35 pp.
1955. Survey of band-tailed pigeon populations in Arizona, Colorado, New Mexico, and Utah, 1954. U. S. Fish and Wildl. Serv., Branch
of Game Mgmt. Albuquerque, New Mexico.
18 pp.
1957. Survey of band-tailed pigeon populations in Arizona, Colorado, New Mexico and Utah, 1956. U. S. Fish and Wildl. Serv., Branch
of Game Mgmt., Albuquerque, New Mexico.
14 pp.
Wight, H. M., R. U. Mace, and W. M. Batterson.
of an adult band-tailed pigeon population
31(3):519-525.
Wooten, W. A. 1955. A trapping technique
Wildl. Mgmt. 19(3):411-412.

~;/
I

Prepared by

_[!__bt=--,..:..~_.

1967. Mortality estimates
in Oregon.
J. Wildl. Mgmt.

for band-tailed

//)

C.
__._D__- '__ /l_4_.._,._t.._1.._-1_.z...-,/;-- __ ----.,.

C lai t E. B raun
Assistant Wildlife

/S~t a . ~X'-;:
Researcher

pigeons.

J.

�- 166 STATE

OF COLORADO

John

A. Love,

DEPARTMENT

Governor

OF

NA TlJRAI

RESOURGS

DEPARTMENT

OF

NATURAL

RESOURCES

DIVISION OF GAME, FISH AND PARKS
Harry

R. Woodward,

T. W. Ten Evc k
Executive
Director
GAME,

FISH

AND

PARKS

COMMISSION

Director

May 9, 1969

Floyd

Getz,

LeRoy

Robson,

President

Harrv Combs,
C.

M.

Furncaux,
Ccrbar.

John

E. Holden,

K. Niess,

WilliJf1l

Cooperators

in Band-Tailed

From:

Howard D. Funk

Subject:

Band-tailed

MembN

W. Robinson,

Foro

51(on9,

Melllb~r

Dc.m

Sullie,

Memb!,f

Pigeon Investigation

Gentlemen:
We are initiating an intensive research program on band-tailed pigeons this year
and need your help to obtain as much vital data as possible.
Pigeons have not
been hunted in Colorado since 1950 when it was deemed populations were too low
to permit further seasons.
We intend to investigate the feasibility of again
hunting pigeons in the State through increased effort in estimating population
levels, locating concentration areas, trapping and banding, and finally by conducting experimental seasons in certain areaso The latter will probably not be
possible until at least 1970
What we need from you at this time is information
on pigeon range and sightings from your past experience and also information on
current observations.
Thus, you have been supplied with two questionnaireso The
first relates to past sightingso
Please fill in the required data and utilize
the enclosed map in plotting locations.
If you can find a better map, please
use it. Pay special attention to Question 3 and be sure to plainly mark locations of sightings of larger flocks on the map. Please return the questionnaire
and map to me as soon as possible.
0

You have also received a supply of Monthly Band-tailed Pigeon Observation formso
Information you supply will allow us to add to our knowledge on pigeon range and
habits within the state and will also help us in locating trapping sites. Use
one form each month, record observations and related data as they occur, and mail
the form to me at the end of each month.
This year I want to start collecting
data in May and some of you may observe birds as late as Novembero
Any and all
information you record on the various sightings will be appreciated!
Please
phone in sightings of larger flocks of birds as soon as possible after observation.
To start, our summer crew will be headquartering out of Durango but may shift to
the San Luis Valley area during the fall trapping period. All of you are more
than welcome to join in any portion of the program, such as trapping, whenever
your schedules permito
Thank you for your helpo

HDF:jc
Enclo

Member

Men:hcr

Pigeon Census

Howard D. Funk
Section Chief
Small Game Research

Prc sidcn r

Member

Or est
O.

To:

Vice

Secretnrv

Member

�- 167 BAND-TAILED PIGEON QUESTIONNAIRE
(Historic Range)

1.

Have you seen or heard of band-tailed pigeons being present
district in past years?
Yes
No
_

2.

Where were the pigeons sighted?
(See enclosed map and circle areas of
sightings.
Please roughly outline the boundary of your district.
Feel
free to show other si.ghtings outside your district.
All information
will be appreciated).

3.

Please list in the space provided below the locations, as close as possible, and approximate sizes of larger flocks (about 100 birds or more)
you have observed or heard of from reliable sources in past years.
If
possible, list the approximate dates of these sightings.
(Plot locations
of these larger groups of the enclosed map and designate them as 1, 2,
3, etc.).

Observers

Name

----------------------------

Address

_

on your

Please return to: Howard Funk
P. O. Box 567
Fort Collins,
Colorado

80521

�170 -

BAND-TAILED

PIGEON

OBSERVATION

Ti!:lr'

Locoti

011

SCCTI.

W\',,!hC'r

Other

ubSf"rV("f

____
._A.M.

P _M.

_

Altitude___

No.

CARD

.Vc q ct o tion

__

.

._..

(behavior,
fceding o r- t i vi t v , n oi s c s m crd c ,
t cnu c n c s s , fliUht
di '.t{'nC(',
I'!(".)

�- 171 -

COLLECTION FORM
Species

Date

Time

Collection No.

Location

Altitude

----_._---

_
County

-----

_ R.T.

------

---------------

Weather ----------

.-----------

Vegetation
Age

_

--------

Bill-----_-0 nun .
Male:

Testes

Female:

Weight

Sex

Ovary

gms .

Total Length_____________

R.

X

(Date)

--0

_

\Hng Length

nun.

L.

mm ,

----------x

_

X

No. Follicles

nun.

Follic le__ .

Df.am , La r ge s t;

nun.

mm ,

._nnn.

Oviduct

----

Crop

Heart

gms.

Fat Appearance
Primaries:

------

Wing

Tail:

Outer 10
9
8

7

Left
_

_

6
5----4
3

l~

3----

6
5------_
4
3-----2

Right
(Outside)

6
5

2
1

_

2

-------

1-----

_

1

--------

Plumage (Describe)--------

--_ .._---

Specimcns Snved
Ec t oparas i. t.es
Crop

_

_

Skin

Wt.

____

Parasi.tes
Gizzard Contents-----Remarks:

Carcass

--------

(Wet-including sac).

Viscera

Gonads

----

----------------

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                  <text>.Tarruar y , 1971

- 1 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-40-R-ll

Antelope

Work Plan No.

2

Job No.

Job Title

Investigations
2

Food Habits of Antelope

Period Covered:

May 1, 1969

to April 30, 1970

Personnel:

George D. Bear

ABSTRACT

No work was accomplished on this job during this segment.
Data shall be
analyzed and a final manuscript prepared during the next segment.

��January,

1971

- 3 -

JOB PROGRESS

S tate of
Project
Work

;;:.CO;;:.L::;:.O:::;RA:..:::;,:;D:::,:O:::...._
No.

W-40-R-ll

Antelope

Investigations

Plan No.

2

Job No.

3

Job Title
Period

REPORT

Covered:

Personnel:

Physiological
May

1, 1969

George

to April

Studies
30, 1970

D. Bear

ABSTRACT

No work was accomplished
on this job during this segment.
Data
analyzed and final reports prepared during the next segment.

shall

be

��January,

1971

~ 5 -

JOB PROGRESS

State of

REPORT

COLORADO
--------~~~~~----------

Project No.

w-4l-R-20

Work Plan No.

1

Job Title

Bighorn Sheep and
Mountain Goat Investigations
Job No.

12

Location and Distribution of
Bighorn Sheep Herds in Colorado

Period Covered:

June 1, 1969 to May 31, 1970

Personnel:

George D. Bear, William

H. Rutherford

ABSTRACT

Aerial and ground surveys were conducted in fourteen bighorn sheep areas
during the last segment.
An effort was made to obtain information on
herd composition, total numbers, seasonal distribution of herds, historical
data concerning the herds, and information on other ungulates utilizing the
areas.
The following herds were found to have decreased in bighorn numbers
during the past ten years: Mesa Verde National Park, Glenwood Canyon,
Cline top Mesa, Black Canyon of the Gunnison, West Elk Wild Area, Pole
Mountain, and Weminuche Creek-Cimarron Peak. Bighorn herds on Battlement
Mesa, Sheep Mountain-Mt. Hope, and Taylor Canyon have remained static during
the past ten years; while Sheep Creek-Trickle Mountain, La Garita Wilderness
Area, and Never Summer Mountains herds have increased.
There is not enough
information available on the Emerald Lake herd to establish a trend. Most
of these herds tend to winter and summer in the same areas, although the
wintering areas are much smaller and restricted.
Most sheep ranges are
utilized by deer and elk during the summer months.
Competition with domestic sheep seems to be restricted to the alpine ranges.
145 licenses were issued and 37 rams
During the 1969 bighorn hunting season
were killed, for a success ratio of 25.5 percent.
Successful hunters
averaged 4.5 hunting days each, unsuccessful hunters averaged 7.9 days,
and the average for all hunters was 7.0 days, for a projected total of 975
man-days of sheep hunting.
Sixty-five percent of permittees were hunting
bighorns for the first time. First-time hunters had a success ratio of 25
percent, while those who had hunted during previous years were 32 percent
successful.
Eighty percent of all permittees started their hunt on the
first day of season. Comments made by hunters are tabulated.
Measurements
of bighorn sheep heads taken during previous years showed that 54 percent
of the sample of heads of 3/4 curl or larger were of rams younger than 7
years, and 29 percent were of rams younger than 5 years.
g

��- 7 -

LOCATION

AND DISTRIBUTION

OF BIGHORN

George D. Bear and William

SHEEP HERDS IN COLORADO
H. Rutherford

P. S. OBJECTIVE
To determine numbers, location,
sheep herds in Colorado.

seasonal

range and composition

of bighorn

SEGMENT OBJECTIVES
1.
2.
3.
4.
5.

Determine bighorn sheep distribution.
Determine population size and trend for each herd.
Determine sex and age composition for each herd.
Determine correlation of age and horn growth of rams in each herd.
Prepare detailed range and distribution maps for each herd with written
descriptions for census.
Develop forms which can be used by management
personnel during routine census and for recording long-term census information.
METHODS

Distribution

AND MATERIALS

and Herd Data

Fourteen bighorn sheep areas were surveyed by horseback, aircraft, or foot
during the last segment.
The entire area was usually surveyed in a random
manner.
Information was collected on the location, total number, sex, and
age of each herd observed.
Interviews with Game, Fish and Parks personnel
and local residents, plus early Colorado Game and Fish Department Federal
Aid Reports, supplied additional information for each area. The Questionnaire
presented in Appendix A was sent to Colorado Game, Fish and Parks field personnel.
Questionnaire data have not been tabulated or analyzed.

Hunter Harvest and Horn Growth Data
One change has been made since previous reports were written.
For previous
methods and materials, see: Woodard, Thomas N. 1969. Bighorn sheep distribution, populations, and herd composition.
In Federal Aid Game Research
Report, January, 1969, pp. 33-38.
For this segment, an age-horn growth
relationship study was added as an objective.
Previously, age-horn-growth
data were included as a sidelight, with no specific place for presentation.
The collection of these data entails interviewing successful bighorn hunters
from past years and, if the head is still in possession, counting annual
growth rings and measuring amount of horn curl.

�- 8 -

RESULTS AND DISCUSSION
Distribution and Herd Data
Battlement Mesa (Mesa County)
Bighorn sheep in this area summer and winter along the narrow hogback
northwest of Collbran, extending from Horsethief Mountain to Horse
Mountain. The area is about 7,500-8,300 feet elevation. Southern
exposures are very steep eroded hillsides with sparse vegetation.
Northern exposures have a heady shrub (mostly oakbrush) and conifer
cover interspersed with grassy parks. Access is limited to foot
travel and a few knowledgeable horsemen. The area is not used by domestic livestock and receives very light use by other wild ungulates.
This herd appears to have been static in recent years. In 1960 there
was an estimated 10 sheep; 15 head were counted with a fixed-wing aircraft in 1964; 34 sheep observed on a helicopter survey in 1968; and
29 on a helicopter survey in 1969. An effort was made to count the
sheep from the ground during June, 1969. Only six sheep (one adult
ewe, one lamb, three yearlings, and one small ram) were observed. The
sheep were scattered in the brush-dominated areas, thus were difficult
to locate and classify. This area is best surveyed by aircraft during
the winter months.

Mesa Verde National Park (Montezuma County)
This is a rough canyon land in the pinyon-juniper vegetative type, interspersed with serviceberry and mountain mahogany. Elevation of the area
is approximately 7,000 feet. Apparently the sheep winter and summer in
the same localities.
Bighorn sheep (14 head) were released at Soda Canyon in 1945. The estimated population in 1960 was 50 head. Ranger Allen Atchinson was interviewed in July, 1969 and estimated the population at 25-30 sheep. Seven
ewes and three lambs were sighted from a helicopter at Pine Canyon in
June, 1969.

Glenwood Canyon (Garfield County)
A small herd of bighorns live in the deep canyon along the Colorado River
approximately 5 miles east of Glenwood Springs. They generally occupy the
range on the north side of the river between No Name Creek and Grizzly
Creek. The sheep scatter over the area during the summer months, and tend
to winter in an apple orchard at the junction of Grizzly Creek and the
Colorado River.
Seventeen sheep were trapped in the Tarryall Mountains in December, 1947
and released at Grizzly Creek. The estimated population in 1960 was

�- 9 -

35 head. There were nine animals at the apple orchard following the
1969 hunting season.
One old ewe died with very heavy lungworm infection during October and an adult ram was killed by an automobile,
so the remaining known population was seven sheep (two lambs, three
adult ewes, one yearling ewe, and one ram). The local Wildlife Conservation Officer thought there were more sheep higher up on the rim
of the canyon, however, helicopter and ground surveys have not substantiated this.

Cline top Mesa

(Garfield

County)

This area is very similar to the Glenwood Canyon Area; juniper-oakbrush
type interspersed with rugged cliffs.
Available information is quite
limited for this herd. Apparently they range between Canyon Creek and
Main Elk Creek approximately seven miles north of New Castle.
An "old timer" at New Castle, Melvin Lykes (lumberjack) was interviewed
on December 10, 1969. He stated there were 38 animals in the herd
during the 1930's.
He generally saw the sheep on lower Deep Creek, and
occasionally up higher on Deep Creek and on East Elk Creek.
Colorado
Game and Fish Division personnel estimated the 1960 popUlation at 54
head in the Clinetops and West Elk Creek. The local Wildlife Conservation
Officer estimates the present popUlation at 12 sheep.
Three adult rams
were found on Hadley Gulch during a helicopter survey, October 6, 1969.
Ground surveys have not been successful in locating the sheep.
Additional efforts should be made during the winter months with a helicopter.

Black Canyon of the Gunnison

(Gunnison County)

This is an extremely rugged canyon area in a juniper-oakbrush
vegetative
type at approximately 7,700 feet elevation on the Gunnison River.
This
appears to be a very good sheep area. It receives moderate deer-use
during the winter months.
There is very little information on the present herd status.
No sheep
were found on a helicopter survey in March, 1969.
Construction workers
at the Marrow Point Dam reported seeing a half dozen sheep in 1968. The
estimated population in 1960 was 35 head. An elderly rancher at Gunnison
said large herds, 150-200 bighorns, wintered in the valley from Gunnison
to Sapinero during the early 1900's.
Additional aerial surveys will be
made in this area.

West Elk Wild Area (Gunnison

County)

This area is located apprOXimately 25 miles northwest of Gunnison.
It is
believed that the sheep summer in the alpine ranges and descend to lower
areas to winter, however, this is not known for certain.
The 1960 population was estimated to be 75 animals.
The local Wildlife Conservation
Officer said the present popUlation may be 50 head.
A helicopter survey

�- 10 -

was conducted in the area last March and no sheep were found. Twentythree sheep were seen on Mill Creek by a cowboy during April, 1966. Two
were seen in West Elk Basin during April, 1969. More information is
needed for this herd.

Taylor River (Gunnison

County)

Bighorn sheep in this area winter on the rocky areas along the northside of the Taylor River from Almont and up the river approximately ten
miles.
This is a sagebrush-bunchgrass
type interspersed with conifers
at about 8,000 feet elevation.
They migrate a few miles further up the
river to summer.
Some move up into the alpine on Matchless Mountain.
These areas are utilized extensively by elk and deer. Population estimate by Department personnel in 1940 was 35; and 35 head in 1960. Thirtyfour sheep (15 ewes, 7 lambs, and 12 rams) were counted during a helicopter
survey on March 9, 1970.

Sheep Creek-Trickle

Mountain

(Saguache County)

This herd of bighorns winters on the southern and southwest exposure of
Trickle Mountain, approximately 15 miles west of Saguache.
The area has
large open bunchgrass parks interspersed with some stands of junipers and
conifers.
During the summer months some of the sheep move westward to
North Park, North Cochetopa Pass, and South Pass; which is more of a
conifer type at 9,000-10,000 feet elevation.
Others apparently move to
the alpine area on Antero Mountain.
This herd has been slowly increasing.
An aerial count in February, 1966
indicated there were 98 animals in the herd. Present estimates are 170
sheep.
The following ground count was made in June, 1969: 39 adult ewes,
24 lambs, 17 yearlings, and one half-curl ram, for a total of 81 sheep.
The following ground count was made in the same area in August, 1969:
43 adult ewes, 25 lambs, 8 yearlings, and 5 rams (two half-curls and
three one-quarter-curls)
for a total of 80 sheep.
A ground count in
December, 1969 tallied 82 adult ewes and yearlings, 20 lambs, and 12 rams
(one one-quarter-curl,
9 half-curls, and two three-quarter-cur1s).
Since
the yearling segment of the herd was not readily distinguished from adult
ewes in the December count, the yearling counts were combined with the
adult ewes to obtain the following ewe:lamb ratios: June, 100:43; August,
100:49; and December, 100:24.
This area is used extensively by a variety
of ungulates and should be watched closely to avoid overgrazing.
Domestic
cattle, deer, elk, and antelope use the same ranges as used by the bighorn
sheep.

La Garita Wild Area - Spring Creek (Saguache County)
The La Garita Wild Area is located approximately 6 miles north of Creede.
Most of the sheep winter and summer on the alpine ranges of Organ Mt., San
Luis Peak, Baldy Alto, and Baldy Chato. A few rams move down to a bunchgrass area on Cebolla Creek to winter and some rams summer in the eastern

�- 11 -

part of the wild area near Machin Lake. The Coleman and Hazzard Ranches
grazed domestic sheep in the alpine until 1961. Since then there has
not been any livestock grazing in the alpine.
Coleman Ranches took a
ten-year non-use permit for the area starting last year. This area receives moderate use by elk during the summer, and very light deer browsing.
Game and Fish Department personnel counted 26 bighorns in Spring Creek
during 1954. Population estimates for this area was 35 head in 1960. The
present population is estimated at 60 animals.
An aerial flight (Cessna 185) was conducted on November 28, 1969. There
was 3-4 feet of snow on the north exposures in the alpine areas, but the
ridgetops and south exposures only had scattered patches of snow. There
were numerous tracks along the ridgetops, but the only sheep we found
were on Baldy Alto.
Five rams (four three-quarter-curls,
and one halfcurl), two ewes, and two lambs were sighted.
Ground surveys were attempted
in late December, however, frequent storms permitted only a brief survey
of the Baldy Chato area. Bighorns were using the alpine ridge above Spring
Creek where winds kept the area free of snow. Fifteen sheep (10 ewes, two
lambs, and three rams) were seen in this area.
Another aerial (Cessna 185) survey was conducted on May 19, 1970. The
north exposures were still covered with deep snow, but the ridgetops and
south slopes were relatively free of snow. The following numbers of sheep
were counted: on Baldy Alto, 14 ewes, and 11 rams: and on Baldy Chato,
6 ewes, 1 ram, and 2 unclassified sheep.
The local Wildlife Conservation
Officer said there were five rams on CebQlla Creek during early May.
Two young rams wintered near the town of Creede.
Ground surveys were conducted in East Bellows, West Bellows, Farmers, East Willow, and West
Willow Creeks for bighorn sheep.
There was not any sign of sheep, other
than the two young rams. It is likely the deep snow in October trapped
the rams on the southern side of the wild area before they could migrate
to Baldy Alto area; thus they descended to the Creede area to winter.
According to some local residents there was a herd of bighorns on West
Bellows Creek until the 1950's.
It looks like a good transplant site.

Pole Mountain

(Hinsdale

County)

This is an alpine area approximately 27 miles west of Creede, Colorado.
Game and Fish Department personnel in 1940 estimated 150 bighorns summering
on Pole Mountain.
The sheep migrated northward to the Lake City area to
winter.
In the spring of 1940 a total of 98 sheep were classified in this
area; 38 ewes, 24 rams, 12 yearlings, 9 two-year-olds, and 17 lambs.
The
estimated population in 1960 was 40 head.
In the summer of 1969 a hiker
reported seeing 14 sheep.
A ground survey was conducted in the area during
December, 1969. A small band composed of four ewes, three lambs and one
yearling was found on the southeast side of Pole Mountain, which was relatively free from snow.

�- 12 -

There is a limited amount of winter range in this area and it is utilized
heavily by domestic sheep and elk during the summer months.
The local
Wildlife Conservation Officer has been working with the U. S. Forest
Service personnel to remove the bighorn wintering area from the domestic
livestock grazing allotment.

Sheep Mountain

- Mount Hope (MineralCourtty)

This area is located about twenty-five miles southeast of Creede.
Bighorns summer and winter on the alpine ranges.
A few descend to the rocky
cliffs on the southwest side of Sheep Mountain above Borns Lake to winter.
The alpine areas are also grazed by deer, elk and domestic sheep during
the summer months.
There was an estimated 35 sheep in this area in 1960. A total of 63 sheep
were counted in the Sheep Mountain area by Department personnel during 1962.
Present population estimates are 40 head.
A ground survey was attempted in this area during June, 1969. Only two
adult ewes were found in the area from Sheep Mountain to the south slopes
of Mount Hope. An aerial count (Cessna 185) was conducted on November 28,
1969. There was deep snow cover in the alpine, but the south exposures
had extensive cleared areas.
The following bighorns were found on the
ridgetops and southern exposures in the alpine:
Mount Hope, 11 ewes, 2
lambs, and 1 ram (half-curl); Sheep Mountain, 6 ewes, 3 lambs, and 1 ram
(half-curl).
There were not any sheep or tracks on Sawtooth or Table
Mountains.

Weminuche

Creek - Cimarron

Peak (HirtsdaleCounty)

This is the alpine area approximately 25 miles northwest of Pagosa Springs.
The area is grazed by elk and deer during the summer months.
Domestic sheep
formerly grazed this area during the summer months, however, the U. S.
Forest Service cancelled the grazing permit.
Estimated 1960 population in this area was 70 bighorns.
Present
made by the local Wildlife Conservation Officer was 35-40 head.

estimate

A ground survey was conducted in the area during August, 1969. Nine bighorns (7 ewes, 1 yearling, and 1 ram) were found in the alpine on the south
side of Weminuche Creek.
On the alpine ridge just west of Hossick Lake
there were 19 head (10 ewes, 7 lambs, and 2 yearlings).
There were tracks
on Cimarron Peak, but no sheep were sighted.
The local W.C.O. said he only
saw one lamb during the summer of 1968, which is reflected in the low
number of yearlings seen in 1969.
An aerial survey (Cessna 185) was conducted on November 28, 1969. There
was heavy snow cover in the alpine, however the southern exposures were
relatively free of snow. No sheep or tracks were seen in the alpine.
Eight
bighorn sheep (4 unclassified, 3 ewes, and 1 lamb) were seen in the timber

�-

13 -

and cliffs about half way down the mountain on the north side of Milk
Creek.
The timber made it very difficult to see the sheep; a helicopter
would be more suitable for the aerial survey work there.

Emerald Lake

(Hinsdale

County)

Bighorns summer in the alpine areas near Emerald Lake on Lake Creek and
along the Los Pinos River to the headwaters of Flint Creek.
There is a
vast expanse of summer range~ but suitable winter range is very limited.
The sheep winter in the cliffs on the south exposure at the junction of
Lake Creek and Los Pinos River.
This area is dominated by conifers and
aspen.
There is very little information on the number of animals in this herd.
The area was surveyed on foot and horseback during August, 1969.
Only
one adult ewe was seen. An aerial survey (Cessna 185) was conducted
during November, 1969 when eight bighorns were found in the alpine just
east of Emerald Lake. More survey work is needed in this area.

Never Summer Mountains

(Jackson County - Larimer

County)

Bighorns in this area remain in the alpine throughout the year.
They
winter on the lower ridges immediately above timberline, on the east side
of the mountain range within Rocky Mountain National Park.
They disperse
throughout the alpine area in summer.
A few animals apparently migrate
eastward across the North Fork of the Colorado River to Specimen Mountain
during the summer months.
This sheep area receives heavy use by elk and deer during the summer
months.
Areas outside the national park boundaries are used extensively
by domestic sheep during the summer months.
This herd has slightly increased during recent years.
The population was
estimated to be 40 animals in 1960. A total of 51 bighorns (24 ewes, 6
lambs, and 21 rams) were counted on an aerial survey during 1969. A ground
survey was conducted during August, 1969. Two adult ewes and a threequarter-curl ram were seen on Bowen Mountain, five ewes and one lamb on Mt.
Baker, and four ewes on Lead Peak.
The area was flown with a fixed-wing aircraft on December 1, 1969.
The
snow was very deep in the area, however, many ridgetops and steep south
facing slopes were exposed.
Eight ewes and a small ram were seen on top
of Mt. Neota.
There were tracks on the ridges along the east side of Lead
Mountain, Mt. Baker, and Bowen Mountain; however conditions did not permit
a closer survey.
Hunter Harvest

and Horn Growth Data

Bighorn sheep hunters during the 1969 season were asked to report on observations of sheep during the hunt.
Table 1 presents what is believed to be
the highest unduplicated counts of sheep seen by hunters in each of the areas.
The term "unduplicated" means that attempts have been made to eliminate
obvious duplications whether noted by the hunter or not. Thus, Table 1 may
still contain a few undetected duplications, but should be reasonably accurate.

�- 16 -

Table 2.

License and harvest statistics, 1969 bighorn sheep hunting season.

Area

First-Choice
Applications
Received

Licenses
Issued

Reported
Kill

1.

Poudre River

17

6

4

3.

Geneva Creek

55

6

2

6.

Pikes Peak

96

15

9

9.

Sangre de Cristo

53

10

2

10.

Sheep Creek-Trickle Mtn.

51

9

3

11.

Collegiate Peaks

33

4

o

14.

Glenwood Springs

2

4

1

l4a. Glenwood Canyon

5

4

1

15.

Sheep Mountain

21

10

1

16.

Cimarron Peak

4

2

1

17.

Empire

7

6

o

19.

Bowen Pass-Clark's Peak

54

12

2

21.

Cow Creek-Wetterhorn Peak

23

8

3

22.

San Luis Peak

18

6

1

23.

McCurdy Mountain

l3

8

1

24.

Battlement Mesa

7

6

2

26.

Taylor River

28

5

2

28.

Vallecito Creek

2

4

o

29.

Blanco River

3

2

o

31.

Mount Evans

37

6

o

32.

Georgetown

15

6

1

33.

Lake City

8

6

1

552

145

37

Totals

�- 17 -

Table 3. Number of hunters
to bighorn sheep management

expressing opinions
and hunting, 1969.

on various

subjects

Item

related

Number of
Connnents

Curl Regulations:
Favor half-curl in certain areas
Opposed to harvest of half-curl rams
Application

2

53

Restrictions:

Favor 2-year restriction on successful applicants
Favor 3-year restriction on successful applicants
Favor no restriction on hunters who make a kill
Favor 5-year restriction on hunters who make a kill
Favor lifetime restriction on hunters who make a kill
Favor allowing application for sheep and goat permits in
alternate years

2

1
4
2

4
1

Other Connnents:
Favor exemption of sheep and goat hunters from blaze
orange requirement
Favor a later season in:
Area l4A
Area 21
Area 24
Favor being allowed to refuse permit if not for firstchoice area
Domestic sheep a problem in the hunting area
Helicopter spooked sheep:
Area 19
Area 21
Too many non-hunters running around in sheep hunting
areas:
Area 6
Area 19
Favor setting of sheep season when no other season is in
progress

9
1

1
2

1
5
2

3

2
1
1

�Table 4. Number of bighorn ram heads measured, by area where killed, curl classification, and number
of growth rings.

Area
1.
2.
3.
4.
5.
6.

9.
10.
11.
12.
14.
15.
19.
21.
22.
23.
24.
31.

Poudre River
Gore Range
Geneva Creek
South Platte Canyon
Rampart Range
Pikes Peak
Sangre de Cristo Range
Sheep Creek-Trickle Mtn.
Collegiate Range
Buffalo Peaks
Glenwood Canyon
Sheep Mountain
Bowen Pass-C1ark's Peak
Cow Creek-Wetterhorn Peak
San Luis Peak
McCurdy Mountain
Battlement Mesa
Mount Evans

Totals

Heads Smaller
Than 3/4 Curl,
Number of
Growth Rings
2
5
3
4

2

3

4

Heads 3/4 Curl or Larger,
Number of Growth Rings
11
8
9
10
6
7
5

13

1

2

1

12

1
1
1
1

1
4

1

1

3

1

1

1

t-'

00

2

1

1

1

2
1
1

1
4
8

4

2

3

1

6

1

1

2
1

1

1

2
1

1

2

1

1

1

1

3

2
1
1

12

1

1
1

1

2

1

3

4

10

4

10

1
1

1

1

1

2

2

3

4

1

�- 19 -

Of particular interest is the proportion of 3/4 curl heads in the
younger age classes. Of the total of 56 heads of 3/4 curl or larger,
30, or 54 percent, have fewer than 7 growth rings; and 16, or 29 percent, have fewer than 5 growth rings. If the objective in bighorn
management is to confine the harvest to mature and over-mature rams,
and if 7 years of age is considered to be a minimum mature ram (as
in the system used by Nevada), the data in Table 4 strongly indicate
that Colorado's 3/4 curl criterion for a legal ram produces an
excessive harvest of young rams.

Prepared by/~~.-«J.
VCU ~•...
~
George KBear
Wildlife Researcher

�- 20 -

APPENDIX A
BIGHORN SHEEP INVESTIGATIONS
A comprehensive report on bighorn sheep in the State is being prepared

and

the following information is needed concerning bighorns in your district.
If certain information is unknown or questionable, please state this so an
effort can be directed toward gathering information in these areas.
Distribution
A map is enclosed; draw a solid line around the summer range occupied by
sheep at the present date, draw a broken or dotted line around the wintering
areas.
Has there been a change in this distribution?

Yes

_

No

_

If so, how has it changed and when?

Herd Composition and Trend
How many bighorns do you believe are in this area?
Is this herd increasing (

), decreasing (

), static (

_
), or unknown (

)7

If you have any historical information on this herd that should be included
in the report please list it here:

�List any survey information you have in your files that can be used in plotting trends.

Date

Type of
Survey*

Location

Ewes

I

Lambs

Bighorns Counted
Rams
J Unclass.
I 1/2 I 3/4 I Full I Unknown I

I

Total

N

I-'

i:Type of Survey: Aerial or Ground

�- 22 -

Other uses in the area occupied
Domestic

by the bighorns:

Livestock

Cattle:

None
, Light
Period of Use

", Moderate

, Heavy

_
_

Sheep:

None
, Light
Period of Use

, Moderate

, Heavy

_

Elk:

None
, Light
Period of Use

, Moderate

Deer:

None
, Light
Period of Use

, Moderate

None
, Light
Period of Use

, Moderate

---------------------------------

Wildlife
, Heavy

_

--------------------------------, Heavy

_

---------------------------------

Mountain
Goats:

, Heavy

_

---------------------------------

Human

Do you believe
distrubition

Management

None
,Light
, Moderate
, Heavy
Primary type(s)~people
using the area
campers, hunters, etc.

(hikers,
_

that any of the above uses have a definite

effect on

or have a limiting

Please comment:

Recommendations

Based on your knowledge
increased

effect on the herd?

(

of this area, do you believe

), maintained

How many bighorns

at the present

do you estimate

level (

this herd should be

), or decreased

should be maintained

(

in this herd?

)?

------------

�- 23 -

Are there any management

problems

should be known by a new

weD coming

Research

that are unique

to this area or herd that

into that district?

Please

list:

Recommendations

Are there any specific
should consider

problems

that the Bighorn

Sheep Research

solving which would assist you in managing

herds in your district?

Please be specific:

Project

the bighorn

�- 24 APPENDIX B
DIVISION OF GAME, FISH AND PARKS
6060 BROADWAY
DENVER, COLORADO 80216
825-1192
LICENSE NO .,

_

BIGHORN SHEEP QUESTIONNAIRE
1969

1. Were you successful in killing a ram? Yes

No

_

If so, what are the approximate horn measurements?
Length of horn around outside curve.
Basal circumference at hair line.
R.H.

R.H.

'---------inches

L.H.

L .H.,

inches

Widest spread (mayor may not be tip to tip)

inches.

-------- inches
-------- inches

2.

Please circle the days on which you hunted (count part days as full days)
August: 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31
September:
1
2
3 4
5
6
7
Area iF1 (Poudre) only:

November 26

27

28

29

30; December

Yes'---------

No,

1 2
567
_

3.

Did you wound a ram and not get him?

4.

Have you hunted bighorn sheep in Colorado other than this year? Yes
If yes, what year(s)?
~-----How many bighorn sheep have you killed in Colorado?

5.

3 4

No

Number

_

How many bighorn sheep, including the one killed, did you see during this
year's hURt?
Full curl Rams - Number
3/4 to full curl - Number.
1/2 to 3/4 curl - Number.
Smaller than 1/2 curl - Number
Ewes - Number
-------------Lambs - Number

----------_
_
_

------------

6. Would you please indicate on the enclosed map by an "X" the approximate location where you saw bighorn sheep during your hunt, and include the numbers,
age, and sex of animals observed. Also, circle location where you made your
kill.
7.

Did you observe any dead bighorn sheep, other than ones killed by hunters?
Dead this year:

Number and sex

Dead previous years:

~-------------

Number and sex

---------------

�- 25 1969-Sheep, Page 2
8.

A plastic bag is enclosed for the collection of fecal samples. If you kill
a ram, would you please strip out the pellets from the lower intestine (colon),
place them in the plastic bag, and give the bag to any Conservation Officer
or leave at any Game, Fish and Parks Office.

9.

Remarks and suggestions - other comments concerning your sheep hunt. We are
particularly interested in comments concerning "quality" sheep hunting, and
comments on the half-curl versus three-quarter-curl regulation. Observations
and general comments are welcome.

��January~

1971

- 27 -

JOB PROGRESS

State

Work

COLORADO

of

No.

W-41-R-20

Plan No.

1

Project

Job Title
Period

REPORT

Job No.

14

~E~v~a~l~u=a~t~i~o~n~o~f~~P~r~oLP~o~s~e~d~B~igg~h=o~r=n~S~h=e~e~p&amp;-~T=r=a~n~s~p~l~a~n~t
_

Covered:

Personnel:

Bighorn Sheep and
Mountain Goat Investigations

June 1969 to May
William

1970

H. Rutherford

ABSTRACT

Potential bighorn transplant sites are proposed usually by Game, Fish
and Parks or U.S. Forest Service personnel.
These sites are then evaluated
for suitability, first by a winter aerial survey to determine whether
snow-free areas exist, and then by a su~ner on-the-ground
survey of only
those sites which showed snow-free areas.
During the current segment, the Mount Wilson area in the San Juan Mountains
was reco~ended
for placement in very low priority, and the Mill Creek
area northwest of Gunnison was reco~ended
for placement in a very high
priority position for bighorn transplants.

��- 29 -

EVALUATION OF PROPOSED BIGHORN SHEEP TRANSPLANT SITES
William H. Rutherford

P. S. OBJECTIVE
To situate bighorn sheep transplant opportunities.

SEGMENT OBJECTIVES
To determine habitat characteristics of proposed bighorn sheep transplant
sites, and rate them by comparison with known occupied bighorn range.

METHODS AND MATERIALS
Potential bighorn sheep transplant sites are suggested or proposed by a
variety of individuals - Wildlife Conservation Officers, Wildlife Researchers,
and U.S. Forest Service personnel. The first activity toward evaluating
their suitability is a winter survey by aircraft during late February or
March, when snow depth is near maximum. Those sites which show little or no
snow-free areas are eliminated from consideration; those showing potential
winter range are scheduled for on-the-ground surveys during the summer months.
During the aerial survey, photographs are made of those snow-free areas which
show vegetation; these photographs are then used as guides for the on-theground survey. Summer surveys are confined to the areas found to be snowfree during winter, and consist of measurements of plant species composition
and density plus notations on nearness of cliffs and/or other rough rocky
terrain which sheep require for escape cover.

DESCRIPTION OF AREA
This job is statewide in scope, but limited during each segment to those
specific locations proposed as transplant sites.

RESULTS AND DISCUSSION
During the current segment, areas on Mount Wilson near Lizard Head Pass in
the San Juan Range and on Mill Creek northwest of Gunnison were surveyed on
the ground. Previously, aerial survey had established the presence of snowfree slopes in these two locations.
In the Mount Wilson area, the complex includes a much larger area than just
Mount Wilson itself. Aerial reconnaissance in March of 1969 showed about 150

�- 30 acres of snow-free ridgetops. All potential winter range is above timberline.
A horseback trip was made in August of 1969 with Forest Service personnel, at
which time gross observations were considered sufficient basis for placing
this area in very low priority as a transplant site. Heavy domestic sheep
use, poor range condition, unstable soil, and poor interspersion of cliffs
and vegetated areas characterized the site. This is historic bighorn range
and should not be rejected entirely as a potential transplant site, but
should be held in abeyance until the Forest Service takes action on closing
out the domestic sheep grazing. Should this occur, the area should be reevaluated as a potential bighorn transplant site.
The Mill Creek proposed transplant site is located on the east side of the
West Elk Wilderness Area. This is historic sheep range which still contains
a small remnant herd. The snow-free winter range is located on a south facing
slope on the north side of Mill Creek, approximately two miles downstream
from the Wilderness Area boundary, at an elevation of about 8,500 feet.
Interspersion of rocky cliffs and vegetated areas is excellent, no domestic
livestock competition exists, and plant species composition and density are
judged to be very good. Vegetation is a bunchgrass-shrub-aspen type, with
Festuca thurberi, Agropyron smithii, Danthonia intermedia, Bromus inermis,
~. anomalus, and Hordeum jtibatum making up most of the grasses and Prunus,
Symphoricarpos, Amelanchier, Ribes, Rosa, and Artemisia representing the
shrubs. Plant density was at least 75 percent even on slopes as steep as
40 degrees. On the basis of this favorable survey, the area is recommended
for a high-priority position on the list of bighorn transplant sites. One
other survey remains to be done in this area; namely, observation on the
amount of time required following a major snowstorm for the slope to again
show bare spots. This will be accomplished during the winter of 1970-71.

Prepared
Wildlife Researcher

�January,

1971

- 31 -

JOB PROGRESS

Sta te of

REPORT

C.;;;.o.;;;.L;:;.o~R;.;;;AD=.;O~
_

Project No.

W-4l-R-20

Work Plan No.

1

Bighorn Sheep and
Mountain Goat Investigations
15

Job No.

Job Title

Mortality

Factors

in Declining

Bighorn

Period Covered:

June 1, 1969 to January

Personnel:

Thomas N. Woodard, Ralph J. Gutierrez,
and William H. Rutherford

Sheep Herds

31, 1970
George D. Bear,

ABSTRACT

Factors possibly contributing to the bighorn sheep herd decline in the
Sangre de Cristo Mountains of Colorado were investigated by systematic
observations during the spring, summer and fall of 1969. High late
summer lamb mortality was found to be the suspected cause of the herd
decline.
Predation was not found to be significant but possible mountain
lion predation cannot be ruled out at this point. Mortality due to
accidents was not significant.
The probable most important factor is
disease possibly enhanced by inclement weather and/or nutritional deficiencies or imbalances.
No specific disease was noted, but it probably
involves respiratory difficulties caused by lungworm in combination with
pathogenic bacteria.

��- 33 -

:MORTALITY FACTORS IN DECLINING BIGHORN SHEEP HERDS
Thomas N. Woodard
P.S. OBJECTIVE
To identify factors responsible for, or contributing to, decline in bighorn sheep (Ovis canadensis) herds in Colorado.
SEGMENT OBJECTIVES
To determine lamb survival and mortality and reproductive potential of
the Sangre de Cristo bighorn sheep herd.
METHODS AND MATERIALS
Herd composition, number of live births, survival of lambs, and the importance of predation, accidents, weather, and disease were investigated by
systematic observations of the bighorn sheep herd during the period covered.
The external appearance of observed sheep was noted and fecal samples were
collected and analyzed for lungworm by the Baerman technique (Cable 1960).
For an effective investigation of the importance of disease it is desirable
to collect and necropsy mortalities, but no recent ones were located during
this period of research.
Though the study was not equipped with sophisticated weather instruments
some temperature readings and visual observations were recorded.
DESCRIPTION OF AREA
Location
The segment of the Sangre de Cristo range in which this study is being
conducted is located in southcentral Colorado approximately 15 miles southwest of Westcliffe. This part of the Sangre de Cristos separates Wet
Mountain Valley to the east from San Luis valley to the west. The east
side lies in the San Isabel National Forest and the west side lies in the
Rio Grande National Forest. The extreme northern boundary of the study area
is the Crestone Needles and the extreme southern boundary is Medano Creek.
However, most of the study area is in or directly adjacent to the Sand Creek
drainage.
Easiest access into the area is across Music Pass from the Wet Mountain
Valley side. The top of the pass lies on the ridge which forms the eastern
boundary of Sand Creek drainage and is accessible only to 4-wheel drive
vehicles or trail machines.

�- 34 -

Topography
The Sangre de Cristo Mountain range in this area is narrow and extremely rugged. Many peaks (Marble Mountain, Milwaukee Peak, Music Mountain,
Tijeras Peak and Cleveland Peak) reach an elevation of about 13,500 feet
(Fig. 1). The bighorn sheep summer range is characterized by steep slopes
cliffs, rock outcroppings and talus slides and is mostly above timberline
which occurs at approximately 11,000 feet.
Most of the study area is situated on the west side of the range and the
streams eventually drain into the San Luis Valley. The major drainages
are Sand Creek and Deadman Creek. However, several small creeks including Little Sand Creek, McKenny Creek, Short Creek, Pole Creek, and
Alpine Creek drain the southern part of the study area.

RESULTS AND DISCUSSION
Many of the bighorn sheep herds in Colorado are presently in a static
condition or are decreasing in size. Some herds, such as the Tarryall
herd in 1953, have had sudden major die-offs affecting all age classes
in the population. Other herds, such as the Sangre de Cristo herd, do
not exhibit a sudden die-off, but instead show a gradual decline of the
population. This study was initiated in an effort to determine the
factors contributing to the decline of the population size in these herds.
Sex and Age Composition
The potential breeding population in the Sangre de Cristos is dominated
by old age class animals. Seven mature rams and only one immature ram
were located. Of the mature rams, 5 were full curl and 2 were 3/4 curl.
They were located in Deadman Basin and Little Medano Creek. Eighteen
mature ewes and only 2 yearling ewes were located. They spent the first
half of the summer near the headwaters of Sand Creek and the last half
of the summer further south in and near Cleveland Gulch (Table 1).
It is impossible to approximate the age of the ewes by field observations
and Geist (1966) found that even the use of horn segment counts in aging
old ewes was invalid. However, it is a safe assumption that the majority
of the 18 adult ewes were in an old age class.
A total of 15 lambs was located. Most were dropped the last week of May
through the first two weeks of June. The lambing grounds are located in
the cliffs above Upper Sand Lake and on the s:ope above Lower Sand Lake
(Fig's. 2 and 3).
Based on the initial reproductive potential this herd should be growing
and productive. The total of 18 adult ewes and 15 lambs gives a maximum
ewe:lamb ratio of 100:83 which is comparable to the best herds in the
state.
Table 1 lists all sheep observations made in the Sangre de Cristos during
1969.

�- 35 -

Figure

1 - Topographic
area.

map, with 40 foot contour

intervals,

of the study

�- 36 -

Fig. 2.
Lake.

Bighorn

sheep lambing grounds on slope above Lower Sand Creek

�- 37 -

Fig. 3.
Lake.

Bighorn

sheep lambing grounds

in cliffs above Upper Sand Creek

�- 38 -

Table 1. Observations of bighorn sheep in the Sangre de Cristo Mountains in
Colorado during 1969.
Date
May 12
May 14
May 28*

Rams

Ewes

Lambs

3

2 - 3/4 curl
5 - Full curl

Yearlings

Location

1

Medano Creek
Little Medano Creek
Rams - Deadman Basin
Ewes - Willow Creek and
Sand Creek
Headwaters of Sand Creek
Headwaters of Sand Creek
Headwaters of Sand Creek
Headwaters of Sand Creek
Headwaters of Sand Creek
Headwaters of Sand Creek
Headwaters of Sand Creek
Headwaters of Sand Creek,
all located on slope
above Lower Sand Lake.
Headwaters of Sand Creek,
all located on slope
above Lower Sand Creek.
Headwaters of Sand Creek,
all located on slope
above Lower Sand Creek.
Headwaters of Sand Creek,
all located on slope
above Lower Sand Creek.
Headwaters of Sand Creek,
all located in cliffs
above Upper Sand Lake.
Headwaters of Sand Creek,
above and north of Upper
Sand Lake.
Headwaters of Sand Creek,
above and north of Upper
Sand Lake.
Headwaters of Sand Creek,
above and north of Upper
Sand Lake.
Headwaters of Sand Creek,
above and north of Upper
Sand Lake.
Headwaters of Sand Creek,
on ridgetop above Upper
Sand Lake.
Headwaters of Sand Creek,
on ridgetop above Upper
Sand Lake.

10

4

7
6

2
2
7

1
1

10
11

2
2
2

June 2
June 3
June 4
June 5
June 6
June 7
June 8
June 9

6

5
6

12

12

2
1

June 10

15

12

2

June 11

15

12

2

June 12

15

12

2

June 14

18

14

2

June 18

10

7

1

June 19

18

14

2

June 20

18

14

2

June 21

18

14

2

June 22
(Incomplete count)

8

11

June 23
(Incomplete count)

12

9

11
13
13
6

1

-------------------------------------------------------------------------------------

* Helicopter flight.

�- 39 Table 1. Observations of bighorn sheep in the Sangre de Cristo Mountains in
Colorado during 1969 (continued).

Date

Rams

Ewes

Lambs

Yearlings

9 total sheep.
June 24
(Incomplete count)

No classification.

June 28

3 total sheep.

No classification.

June 29

3 total sheep.

No classification.

July 1

5

1

July 2

2

1

July 3

18

15

July 4

12

10

July 5

13

10

July 6

16

13

July 8

13

11

July 23

13

11

July 24

6

4

August 12*

22 total sheep. No classification.
A number of lambs were present

August 15
August 16

2

1

1

(3/4-fu11 curl)

1

Location

Headwaters of Sand Creek,
on ridgetop above Upper
Sand Lake.
Headwaters of Sand Creek,
on ridgetop above Upper
Sand Lake.
Headwaters of Sand Creek,
on ridgetop above Upper
Sand Lake.
Headwaters of Sand Creek,
on slope above Lower
Sand Lake.
Headwaters of Sand Creek,
on slope above Lower
Sand Lake.
Headwaters of Sand Creek,
on knoll between Lower
and upper Sand Lakes.
Headwaters of Sand Creek,
on knoll between Lower
and upper Sand Lakes.
Headwaters of Sand Creek,
on slope above Lower
Sand Lake.
Headwaters of Sand Creek,
in cliffs above Upper
Sand Lake and at salt
lick below Music Pass.
Headwaters of Sand Creek,
on slope above Lower
Sand Lake.
Headwaters of Sand Creek,
on slope above Lower
Sand Lake.
Headwaters of Sand Creek,
north of Upper Sand Lake.
On ridgetop between Sand
Creek and Pole Creek.
On ridgetop at headwaters
of Little Sand Creek.
Same ewe located on August
15. Ram located on north
ridge of Deadman Basin.

--------------------------------------------------------------------------------------

* Helicopter flight.

�- 40 Table 1. Observations of bighorn sheep in the Sangre de Cristo Mountains in
Colorado during 1969 (continued).
Rams

Date

August 29

Sept. 10

1
(approx. 2 yr.
old)

Ewes

Lambs

2

1

Headwaters of Sand Creek,
on top of Milwaukee
Peak.

12

2

Cleveland Gulch on Lower
Sand Creek.

Yearlings

Location

Lamb Survival
The theory that major sheep mortality occurs in the transition from lamb
to yearling age class has been suggested as the reason for the decline in
many herds. The resultant low annual increment to the herds is not large
enough to replace normal mortalities in the lamb population.
Mills (1937) and Beuchner (1960) found abnormally high lamb mortality in
summer in Yellowstone National Park. Woodgerd (1964) discovered a lamb
mortality of 40 percent in the first 6 months of life in the sheep herd
on Wildhorse Island. In Colorado, Streeter (1969) found a very low survival to the yearling class on Buffalo Peaks.
The first year's results of this study showed high lamb mortality in late
summer is the probable cause of the herd decline in the Sangre de Cristos.
The maximum ewe:lamb ratio of 100:83 with 18 adult ewes and 15 lambs was
noted on July 3. On July 23, 13 adult ewes and 11 lambs were located, and
then on September 10, 12 adult ewes and only 2 lambs were located. Though
6 adult ewes we're not accounted for on September 10, and considering the
possibility that all 6 still had surviving lambs, the result would be a
maximum of 8 surviving lambs. Then the minimum mortality rate for lambs
would approach 50 percent. The actual mortality rate is certainly higher,
with the result that the annual increment to the herd cannot replace natural
mortalities in the population and therefore the herd gradually declines in
size.
Also, the fact that only 2 yearling ewes and one immature ram were located
in 1969 is additional proof of high mortality in the first year of life.
After July 23 the majority of the sheep left Sand Creek Basin for the rest
of the summer. The large ewe herd divided into smaller groups and due
partially to a prolonged period of inclement weather (Table 2) and an unfamiliarity with surrounding areas a substantLaI number of sheep were not
relocated for an extended period of time (Table 1). It was during this
critical time period that the suspected mortalities occurred, offering no
opportunity to actually observe what took place.

�- 41 Table 2. Weather observations made in the Sangre de Cristo Mountains in
Colorado during the summer of 1969.

Date

Observation

June 2

Dry, warm

June 3

Wind, rain, sleet and snow in afternoon for 1 hour.

June 4

Dry, warm

June 5

Dry, warm

June 6

Wind and rain in afternoon for 2 hours.

June 7

Wind, hail, rain and lightning in afternoon.
extensive than previous days.

June 8

Rain and hail in afternoon.

June 9

Rain and fog in morning.
wind in evening.

June 10

Light drizzle in early morning.
and cold all day.

June 11

Cloudy, fog, snow and cold all day.

June 12

Cloudy, snow and cold in afternoon for 2 hours.

June 13

Six inch snow accumulation in morning. Additional
snow and hail in evening. Temperatures - Low 34°,
High 49°.

June 14

Intermittent snow, hail, rain and fog.
tures - Low 37°, High 45°.

June 15

Intermittent snow, rain and fog.
Low 36°, High 42°.

Temperatures-

June 16

Intermittent snow, hail and fog.
Low 37°, High 42°.

Temperatures-

June 17

Intermittent snow and fog.
High 44°.

June 18

Fair and partly cloudy most of day. Rain in
evening. Temperatures - Low 36°, High 52°.

More

Heavy fog by evening.

Heavy rain, hail, and
Cloudy, windy

Tempera-

Temperatures - Low 38°,

�- 42 Table 2. Weather observations made in the Sangre de Cristo Mountains in
Colorado during the summer of 1969 (continued).

Date

Observation

June 19

Fair, partly cloudy and windy all day. No precipitation. Temperatures - Low 36°, High 59°.

June 20

Fair, partly cloudy and windy. No precipitation.
Temperatures - Low 40°, High 63°.

June 21

Clear, warm, windy.
High 66°.

June 22

Partly cloudy most of day. Rain and hail in
evening. Temperatures - Low 43°, High 62°.

June 23

Partly cloudy to overcast all day. Rain and hSil
in afternoon. Temperatures - Low 42°, High 56

June 24

Overcast, high winds, rain, and 2-3 inch accumulation of snow in evening and night. TemperaturesLow 29°, High 49°.

June 25

Overcast, snow, wind and cold.
No temperature readings.

June 26

Blizzard conditions.

June 28

Clear, warm and windy.
accumulation.

Five inch new snow

June 29

Clear, warm and windy.
High 64°.

Temperatures - Low 38°,

June 30

Clear, warm and gusty winds.
Low 39°, High 71°.

July 1

Clear to partly cloudy most of day. Heavy hail
and rain for 30 minutes in afternoon. Temperatures
Low 42°, High 67°.

July 2

Clear and warm in morning. Overcast and rain in
afternoon. Temperatures - Low 48°, High 73°.

July 3

Clear and warm in morning. Overcast and light
rain in afternoon. Temperatures - Low 43°, High 74°.

July 4

Overcast with rain and fog most of day.
tures - Low 43°, High 62°.

Temperatures - Low 44°,

Blizzard conditions.

Temperatures-

Tempera-

----------------------------------------------------------------------------

�- 43 Table 2. Weather observations made in the Sangre de Cristo Mountains in
Colorado during the summer of 1969 (continued).
Observation

Date

July 5

Clear in morning. Overcast with rain and hail in
afternoon. Temperatures - Low 42°, High 61°.

July 6

Partly cloudy and windy. No precipitation.
Temperatures - Low 47°, High 63°.

July 7

Clear and warm.

Temperatures - Low 43°, High 67°.

July 8

Warm and clear.

Temperatures - Low 43°, High 69°.

July 9

Light rain most of day.
High 50°.

July 10

Clear in morning. Overcast with rain and hail in
afternoon. Temperatures - Low 42°, High 64°.

July 11

Clear in morning. Overcast with rain and hail in
afternoon. Temperatures - Low 45°, High 66°.

July 12

Partly cloudy in morning. Overcast with rain and
hail in afternoon. Temperatures - low 45°, High 64°.

July 13

Partly cloudy in morning. Overcast with rain in
afternoon and evening. Temperatures - Low 47°,
High 71°.

July 14

Clear in morning. Overcast with rain, hail and
lightning in afternoon. Temperatures - Low 45°,
High 69°.

July 15

Clear in morning.

July 16

Rain and hail with fog in afternoon and evening.

July 17

Partly cloudy with some fog in morning.
with rain in afternoon.

July 18

Partly cloudy in morning.

July 23

Clear in morning. Overcast with rain and hail in
afternoon.
Temperatures from July 15 to July 23 - Low 440,
High 69°.

Temperatures - Low 410,

Overcast with rain in afternoon.

Overcast

�- 44 Table 2. Weather observations made in the Sangre de Cristo Mountains in
Colorado during the summer of 1969 (continued).
Date

Observation

July 24

Clear in morning. Overcast with heavy rain, hail
and lightning in afternoon. Temperatures - Low 420,
High 56°.

July 25

Clear in morning. Overcast with light rain in
afternoon. Temperatures - Low 43°, High 70°.

July 26

Clear in morning. Overcast with light rain in
afternoon. Temperatures - Low 44°, High 68°.

July 27

Clear in morning.

July 28

Clear in morning. Partly cloudy to overcast in
afternoon. No precipitation.

July 29

Clear to partly cloudy and warm.

August 2

Clear in morning. Overcast with rain in afternoon,
evening and night.
Temperatures from July 27 to August 2 - Low 440,
High 72°.

August 3

Partly cloudy with rain at night.
Low 45°, High 70°

August 4

Clear in morning.

August 6

Clear in morning. Overcast with light rain and wind
in afternoon.
Temperatures from August 4 to August 6 - Low 460,
High 72°.

August 7

Clear to partly clody with light rain.

August 11

Clear to partly cloudy.
Temperatures from August 7 to August 11 - Low 450,
High 71°.

August 12

Partly cloudy in morning. Light rain in evening and
during night.

Partly cloudy in afternoon.

Temperatures-

Overcast with rain in afternoon.

--------------------------------------------------------------------------------

�- 45 -

Table 2. Weather observations made in the Sangre de Cristo Mountains in
Colorado during the summer of 1969 (continued).

Date

Observations

August 13

Heavy fog and light rain all day.

August 15

Partly cloudy and warm.

August 16

Partly cloudy and warm.

August 17

Clear to partly cloudy and warm.

August 18

Clear to partly cloudy and warm.

August 19

Partly cloudy with fog on ridgetops.

August 21

Clear to partly cloudy in morning.
afternoon.

August 22-29

Partly cloudy and cool with intermittent light
rain.

August 30

Fog and light rain all day.
Temperatures from August 11 to August 30 - Low 400,
High 680•

Rain in

�- 46 -

Factors Contributing to Mortalities
Predation
No coyote (Canis latrans) predation on bighorn sheep in the Sangre de Cristos
was observed during 1969. Only two coyotes were seen on potential bighorn
sheep range and there were no observed interactions between sheep and coyotes.
Other authors report no significant predation on sheep by coyotes. Sperry
(1941) analyzed 8,339 coyote stomachs and reported that only a trace of bighorn sheep remains, which was probably carrion, was present. Honess and
Frost (1942) removed 131 coyotes from the Gros Ventre range in Wyoming, reducing the population by 75 percent. No corresponding increase in the sheep
population was noted. Smith (1954) found only 2.8 percent of collected coyote
scat in Idaho contained bighorn sheep remains.
One authentic case of coyote predation was noted by Crump (1958) who saw a
coyote chase and kill a lamb in the Torrey Creek drainage in Wyoming.
All available evidence indicates that golden eagles (Aquila chrysaetos) also
are not significant predators of bighorn sheep. Most authors have not seen
any attempted predation by golden eagles. However, Crump (1958) reports a
Wyoming Game and Fish Department pilot saw a golden eagle knock a lamb from
a cliff and then return to feed on it. Rush (1940) and Kennedy (1948) saw
golden eagles feeding on lamb carcasses but there was no conclusive evidence
that they actually killed the lambs.
Many golden eagles and one bald eagle (Haliaeetus leucocephalus) were seen in
potential bighorn sheep range in the Sangre de Cristos in 1969. Some of the
eagles passed directly over sheep at low elevations. Neither the eagles nor
sheep seemed to show interest in or notice the other at any time.
In contrast to coyote and eagle predation, the possibility of significant
mountain lion (Felis concolor) predation cannot be dismissed. Hornocker (1970)
found predation on bighorn sheep by mountain lions in Idaho to be of no significance. He felt that bighorn sheep group behavior preVents a successful
stalk by a mountain lion. However, Moser (1962) reported a large lion was
seen on sheep range at Empire, Colorado and the sheep population was reduced
by 4 animals in a period of 5 days. He also reported that a lion killed a
3 year old ram near Pikes Peak.
Though there were no sightings of mountain lions on potential bighorn sheep
range in the Sangre de Cristo Mountains in 1969, it is a distinct possibility
that some of the late summer lamb mortality which occurs after the sheep leave
the protective escape cover of the lambing grounds is the result of mountain
lion predation. However, it seems logical to assume that a lion would prey
on all age classes and not take only the young of a population.
Accidents
Although it is inevitable that some bighorn sheep are lost due to accidents,
this has to be considered a part of normal herd drain. Literature contains a

�- 47 few reports of observed sheep mortalities due to accidents, but during
field observations in the Sangre de Cristo Mountains in 1969 no bighorn
sheep of any age class were seen to be in danger of falling in even the
most difficult terrain.
The skeletal remains of a 9-year-old ram, the
death of which could have been the result of an accident, was found at
the bottom of a rock slide.

Weather
The bighorn sheep
adverse reactions
inclement weather
25 was especially

in the Sangre de Cristo Mountains showed no immediate
to bad weather.
They survived prolonged periods of
in excellent condition.
The time period around June
cold and wet with blizzard conditions prevailing (Table

2).
It is possible that weather of this type may be a delayed factor in abnormally high mortality rates.
Cold and wet weather may enhance the invasion
of disease-causing
organisms, especially in new born lambs, the effects of
which may not appear for weeks.
The suspected high late summer lamb mortality in the Sangre de Cristos
dicates that this could indeed be a contributing factor.

in-

Disease
Any statement concerning the causative organism of a possible disease condition in the Sangre de Cristo Mountains would be pure conjecture since no
mortalities were collected during this segment.
The external appearance
of the observed sheep was noted but they always appeared to be in excellent
condition.
The sheep never exhibited any of the typical signs of disease;
hunching up, emaciation, coughing, sneezing, or a mucoid discharge from the
nose.
However, if predation and accidents are not significant causes of the high
late summer lamb mortality in the Sangre de Cristo Mountains, then disease
is at least the probable proximate factor.
It is possible a disease complex
of rapid onslaught occurs, appearing explosively and attacking a large number
of lambs within a short period of time.
It is well documented that the majority of the Rocky Mountain bighorn sheep
in the United States are infected with lungworm (Protostrongylus
stilesi and
P. rushi).
Couey (1950) reported a 75-100 percent infection rate among bighorn sheep herds in Montana.
Infection rates ranged from 35-93 percent in
Idaho (Smith 1954).
In western Montana Forrester and Senger (1964a) reported
that 91 percent of 900 fecal samples from 10 herds contained Protostrongylus
larvae.
Pi1lmore (1961) reported that sheep herds in every area sampled in
Colorado except the Arkansas River Canyon herd are infected with lungworm.
Some historical die-offs of bighorn sheep herds have been attributed to 1ungworms.
Pillmore and Moser (1954) reported that necropsy of several mortalities

�- 48 from the Tarryall die-off in 1953 showed death due to verminous pneumonia
with Protostrongylus the causative organism.
Lungworm could be the causative organism of the herd decline in the Sangre
de Cristos but the collection and analysis of fecal samples was given low
priority during this segment. Pillmore (1956) found that fecal analysis
is useful only in determining if a herd is infected, but not reliable for
determining infection intensity. Samples were collected on July 3 and
again on September 3 (Tables 3 and 4). All adult samples and 3 of 6 lamb
samples collected on July 3, and all adult samples and 3 of 4 lamb samples
collected on September 3 were positive for lungworm.

Table 3. Results of fecal analyses on samples collected above Upper Sand
Creek Lake on July 3, 1969.

Sample

Dry Weight

Total Larvae

Larvae
per gram

1 - Adult

10.lg

129

12.8

2 - Adult

9.2g

400-450

43.4-48.9

3 - Adult

9.4g

340

36.2

4 - Adult

l3.4g

122

9.1

5 - Adult

l3.7g

95

6.9

6 - Adult

8.1g

181

22.3

7 - Lamb

7.3g

0

0.0

8 - Lamb

7.0g

0

0.0

9 - Lamb

7.3g

0

0.0

10 - Lamb

8.8g

4

.5

11 - Lamb

10.0g

120

12.0

12 - Lamb

7.6g

55

7.1

�- 49 Table 4. Results of fecal analyses on samples collected above Deadman
Lake on September 3, 1969.
Sample

Dry Weight

Total Larvae

Larvae
per gram

1 - Adult

l4.0g

500-550

35.7-39.3

2 - Adult

l2.2g

900-1000

73.8-81. 9

3 - Adult

l2.7g

390

30.7

4 - Adult

8.6g

550-600

63.9-69.6

5 - Adult

9.7g

500-550

51.5-56.7

6 - Adult

10.5g

900-1000

85.7-95.2

7 - Lamb

7.lg

0

0.0

8 - Lamb

5.2g

271

52.1

9 - Lamb

7.9g

319

40.4

10 - Lamb

10.lg

500-550

49.5-54.5

This is in agreement with results of other researchers. Pillmore (1956)
found positive lamb samples as early as mid-June. Forrester and Senger
(1964b) found 45-47 day old lambs shedding first stage protostrongylid
larvae. They also found protostrongylid larvae in the lungs of 3 necropsied
lambs; one 10 days old, one 2-3 days old and one near-term foetus. These
results suggest infection prenatally or soon after birth.
Though lungworm infection is undoubtedly a contributing factor, many mortalities due to non-verminous pneumonia have been reported. Potts (1937)
thought that Pasturella sp. and Corynebacterium pyogenes isolated from the
lungs were the cause of death in 2 rams in Rocky Mountain National Park.
Marsh (1938) also determined that the cause of losses of bighorn sheep
lambs on the National Bison range in Montana was due to pneumonia caused
by these two bacterial species. Some of the lamb mortality discovered by
Woodgerd (1964) on Wildhorse Island was due to a massive infection of ~.
pyogenes. In most of 8 mortalities at Glen Eyrie, Colorado in NovemberDecember 1959, Pasturella and Diplococcus organisms were located (Pillmore
1961). Post (1962) attributed the death of 17 sheep at the Sybille Experimental Unit in Wyoming to Pastuerellosis. He feels the short term dieoffs of sheep indicate a virulent organism and not the slow debilitating
diseas~ resulting from parasitism. However, he believes lungworm infection
may cause tissue damage which facilitates invasion of the bacterium. This
viewpoint is held by many other researchers.
Other pathogenic bacteria tave been isolated from the lungs of some mortalities.
Honess and Frost (1942) discovered late summer lamb mortality in the Crystal
Creek herd in Wyoming. The necropsy of two lambs showed death due to pneumonia
caused by a Streptococcus organism.

�- 50 Besides verminous and non-verminous pneumonias, other diseases are most certainly contracted by bighorn sheep. Scabies caused by the common mite
(Psoroptes equi var. ovis) has been the cause of some historical die-offs.
This possibility is not considered significant in the Sangre de Cristos for
two reasons. First, the pelage of all observed sheep was in excellent condition; and second, bighorn sheep usually contact scabies as a result of
contact with domestic sheep, none of which are grazed in this area of the
range.
Honess and Winter (1956) list other known infectious, non-infectious and
parasitic diseases of bighorn sheep.
Nutrition
If disease is a proximate factor in bighorn sheep herd reductions then nutritional deficiencies may be the ultimate factor (Streeter 1969). Although
this study is not investigating this possibility, a brief review of the
literature is appropriate.
Streeter (1969) made limited collections of forage species (four different
grass or grass-like species were collected) on alpine ranges in 1968 and
proximate analyses were conducted. Crude protein percentages in the Buffalo
Peaks area and phosphorus percentages in all collection areas were below the
recommended minimum levels for domestic sheep. This could be significant
because phosphorus deficiencies cause weak lambs and decreased milk production
in ewes (NRC 1964).
Cook and Harris (1968) conducted feeding tests with domestic sheep over a 3year period on Utah desert ranges known to be deficient in phosphorus and
protein. They found that supplements of phosphorus and protein in combination
increased wool yield and lamb crop.
These results point out the distinct possibility that nutritional deficiencies
or imbalances may be the major factors of bighorn sheep herd declines. However,
much additional research is needed before concrete facts can be established.

LITERATURE CITED
Beuchner, H. K. 1960. The bighorn sheep in the United States, it's past,
present and future. Wildlife Soc., Wildlife Monogr. 4. 174 p.
Cable, R. M. 1960. An illustrated laboratory manual of parasitology.
Burgess Publ. Co. 153 p.
Cook, C. W., and L. E. Harris. 1968. Nutritive value of seasonal ranges.
Agr. Exp. Sta. Utah State Univ. Bull. 472.
Couey, F. M. 1950. Rocky Mountain bighorn sheep of Montana.
and Game Comm. Bull. 2. 90 p.

Montana Fish

�- 51 Crump. W. I. 1958. Bighorn sheep reproduction and lamb survival study.
Wyoming Game and Fish Comm., Fed. Aid in Wildlife Restoration, Big
Game Survey. April: 1-28.
Forrester, D. J., and C. M. Senger. 1964a. A survey of lungworm infection
in bighorn sheep of Montana. J. Wildl. Manage. 28(3):481-491.
1964b.
lungworms.

Prenatal infection of bighorn sheep with protostrongylid
Nature. 201(4923):1051.

Geist, V. 1966. Validity of horn segment counts in aging bighorn sheep.
J. Wild!. Manage. 30(3):634-635.
Honess, R. F., and N. M. Frost. 1942. A Wyoming bighorn sheep study.
Wyoming Game and Fish Dept. Bull. 1. 127 p.
Honess, R. F., and K. B. Winter. 1956. Diseases of wildlife in Wyoming.
Wyoming Game and Fish Dept. Bull. 9. 279 p.
Hornocker, M. G. 1970. An analysis of mountain lion predation upon mule
deer and elk in the Idaho Primitive Area. Wildlife Soc., Wildlife
Monogr. 21. 39 p.
Kennedy, C. A. 1948.
29(1):68-69.
Marsh, H. 1938.
19(2):214.

Golden eagle kills bighorn lamb.

J. Mammal.

Pneumonia in Rocky Mountain bighorn sheep.

Mills, H. B. 1937.
National Park.

J. Mammal.

A preliminary study of the bighorn of Yellowstone
J. Mammal. 18(2):205-212.

Moser, C. A. 1962. The bighorn sheep of Colorado.
Fish Dept. Tech. Pub. 10. 49 p.

Colorado Game and

National Research Council. 1964. Nutrient requirements of domestic animals.
Nutrient requirements of sheep. Nat. Res. Council Pub. 1193. 90 p.
Pillmore, R. E., and C. A. Moser. 1954. Study of mortality factors.
Colorado Game and Fish Dep. Fed. Aid Quart. Rept. January:53-66.
Pillmore, R. E. 1956. Investigations of the life history and ecology of
the lungworm, Protostrongylus stilesi. Colorado Game and Fish Dept.
Fed. Aid Quart. Rept. April:47-70.
1961. Study of the lung nematodes of bighorn sheep; comparative
studies of infection intensities in wild and laboratory populations.
Colorado Game and Fish Dept. Fed. Aid Quart. Rept. October:85-97.
Post, G. 1962. Pasteurellosis of Rocky Mountain bighorn sheep (Ovis
canadensis canadensis). Wildlife Disease. 23:1-14.
Potts, M. K. 1937.
Hemorrhagic septicemia in the bighorn of Rocky
Mountain National Park. J. Mammal. 18(1):105-106.

�- 52 Rush, W. M.

1940.

Bighorns need a break.

Outdoor Life.

85(1):38-40,85.

Smith, D. R. 1954. The bighorn sheep in Idaho, its status, life history
and management. Idaho Dept. Fish and Game, Wildlife Bull. 1. 154 p.
Sperry, C. G. 1941. Food habits of the coyote. U.S. Fish and Wildlife
Servo Wildlife Res. Bull. 4:34-41.
Streeter, R. G. 1969. Demography of two Rocky Mountain bighorn sheep
populations in Colorado. Ph.D. Thesis. Colorado State Univ., Fort
Collins. 96 p.
Woodgerd, W.
Island.

Prepared

1964. Population dynamics of bighorn sheep on Wildhorse
J. Wildl. Manage. 28(2):381-391.

bY~uJ~x1
T~mas N. Woodard
Student Assistant

�January~

1971

- 53 -

JDB PROGRESS REPORT

State of

COLORADO

Project No.

w-4l-R-20

Work Plan No.

2

Job Title

Bighorn Sheep and
Mountain Goat Investigations
Job No.

4

Location and Distribution of
Mountain Goat Herds in Colorado

Period Covered:

June 1969 to November

Personnel:

William

1969

H. Rutherford

ABSTRACT

Mountain goat distribution and sex-age classification data collected
during the 1969 segment year are tabulated.
These data are of a general
nature, but indicate that nanny:kid ratios continue to remain at a high
level, with kid:yearling ratios considerably lower. The population trend
in the two areas which have well-established
herds, Mount Evans and the
Collegiate Range, is definitely up. The status of the goats in the Gore
Range and the Needles Mountains is less well known, due to small numbers
and rugged terrain.
Difficulty in obtaining accurate population data in
these two areas will probably continue.
During the 1969 mountain goat hunting season, 21 licenses were issued and
20 goats were killed, for a success ratio of 95 percent.
The average
hunter spent 4.2 days hunting goats in 1969, for a total of 88 man-days
of goat hunting.
All but two of the permittees were hunting goats for the
first time, and all but two started their hunt on the first day of season.
The harvest consisted of 3 adult female and 13 adult male goats in the
Collegiate Range, and 1 adult female and 3 adult male goats on Mount Evans.
Two hunters reported wounding and not getting a goat.
The assignment of
Division employees to accompany goat hunters was for assistance in choosing
a male goat, but the final choice was up to the license holder.
Harvest
data indicate that this procedure may have contributed to a reduction in
the legal harvest of female goats.

��- 55 -

LOCATION

AND DISTRIBUTION

OF MOUNTAIN

William

GOAT HERDS

IN COLORADO

H. Rutherford

P.S. OBJECTIVE
To determine numbers, composition,
mountain goat herds in Colorado.

seasonal

SEGMENT

OBJECTIVES

1.

Determine

mountain

2.

Determine

population

3.

Determine

sex and age composition

range,

and location

of

goat distribution.
size and trend for each herd.
of each herd.

4. Prepare detailed range and distribution

maps for each herd, with
written descriptions for census.
Develop forms which can be used
by management personnel during routine census and for recording
long-term census information.

METHODS

AND MATERIALS

No changes have been made since previous reports were written.
See:
Woodard, Thomas N. 1969. Mountain goat distribution, populations and
herd composition.
In Federal Aid Game Research Report, January, 1969.
pp. 57-59.

DESCRIPTION

OF AREA

This job is statewide in scope, but limited to those areas in which
mountain goat populations currently exist; i.~., the Collegiate Range,
Mount Evans, the Gore Range, and the San Juan Mountains.

RESULTS

AND DISCUSSION

Population

Status

Mountain goat distribution and sex-age classification
data were obtained
by on-the-ground observation techniques using binoculars and spotting
scope, and by requesting hunters to record their observations of the
animals.
Results are presented in Table 1.

�- 56 -

Table 1.
1969-70.

Mountain

goat observations

summer 1969 and winter

in Colorado;

Numbet·of

Goats·

Adults
Male

Female

Date

Area

Aug. 1969

Mt. Aetna*

1

Aug. 1969

Mt. Shavano *

2

18

9

Aug. 1969

Mt. Princeton*

1

3

Aug. 1969

Sheep Mountain*

10

20

Aug. 1969

Mt. Evans *

4

4

8/9/69

Mt. Shavano

2

19

4

8/9/69

Sheep Mountain

3

14

9/10/69

Needles

2/12/70
* Hunter

Unclassified

Yearlings

Kids

4

Total

5
17

54

22

3

29

20

14

64

4

12

7

17

49

3

3

12

35

Mtns.

2

1

Sheep Mountain

19

observations;

duplication

eliminated

8

3
19

where possible.

Total counts cannot be obtained because of the generally rugged terrain through
which the goats are dispersed.
Thus, the data in Table 1 are valuable primarily
for determining the status of each individual herd, its general sex-age composition, and its reproductive success for the given year.
It has consistently
held true that nanny:kid ratios are high in all herds, but kid:yearling ratios
are much lower, indicating that loss of kids through mortality before reaching
the yearling class is common.
This is probably a characteristic to be expected
in goat populations, and is no cause for concern as long as kid:yearling ratios
stay high enough to insure that annual herd increment is at a satisfactory level.
In the two areas which have well-established
herds, Mount Evans and the Collegiate
Range, mountain goat populations continue to expand in numbers and herds continue to exhibit a healthy status.
The status of the goats in the Gore Range
and the Needles Mountains is less well known.
Goats from the 1968 transplant
in the Gore Range are known to be still present, by tracks and droppings, but
sightings were not made in 1969 and no information on reproductive success is
available.
Only three goats were seen in the Needles Mountains in 1969 by
project personnel, but reports from private pilots indicate that at least 15
goats may be present.
Difficulty in obtaining accurate population data will
continue in these areas until the herds have grown considerably in numbers.

�- 57 -

Distribution

Maps and Census Forms

Preparation of detailed range or distribution maps, census descriptions,
and census forms are still incomplete and not ready for submitting with
interim progress reports.
Work was accomplished on them during this segment, with the objective being to prepare them in finished form for the
final report •

.Summary

of 1969 Mountain

Goat Hunting

Season

As determined by a special mail questionnaire to each goat hunter, 20
mountain goats were harvested by 21 hunters, for a success ratio of 95
percent.
The average hunter spent 4.2 days hunting goats in 1969.
Days hunted
ranged from 1 to 16, for a total of 88 man-days for all goat hunters.
All but two of the permittees were hunting goats for the first time.
All but two started their hunt on the first day of season; one started
on the second day and another did not start until the ninth day.
In the
Collegiate Range, 3 adult female and 13 adult male goats were taken; on
Mount Evans, 1 adult female and 3 adult male goats were taken.
Two hunters,
including the one who was unsuccessful, reported wounding and not getting
a goat.
By Commission action, the 1969 mountain goat hunting regulations were
amended with the requirement that a qualified Division employee accompany
each party of hunters.
The disproportionate
harvest of female goats in
1968 was considered to be detrimental to herd status; and to avoid a repetition, the Division employees were assigned to aid the hunter in selecting
a male goat.
Female goats, however, continued to be legal game, as regulations stipulated only that a goat have horns at least six inches long and
that no mature female accompanied by young may be taken.
Thus, the Division
employees assignment was for assistance only, and the final decision on
selection of an animal for harvest was up to the license holder.
Harvest data indicate that the 1969 regulations as amended may have been
effective in reducing the legal harvest of female animals.
The procedure,
however, is expensive and inefficient from the standpoint of Division personnel operations.
In view of the small number of licenses issued, the
potential for harvesting female goats is not great enough to be detrimental
to well-established
herds nor to justify such intensive efforts at control.
With the need for surplus goats to furnish stock for an intensified transplanting program, it appears that the number of hunting permits should not
be increased in any of the goat hunting areas, at least for the next few
years.

Prepared
Wildlife

Researcher

�January,

1971

- 65 -

JOB PROGRESS REPORT

State of

COLORADO
----------~--------------

Project No.

W-41-R-20

Work Plan No.

2

Job Title

Evaluation

Bighorn Sheep and
Mountain Goat Investigations
Job No.
of Proposed Mountain

Period Covered:

June 1969 to May 1970

Personnel:

William

6
Goat Transplant

Sites

H. Rutherford

ABSTRACT

Potential mountain goat transplant sites are proposed usually by Game,
Fish and Parks or U.S. Forest Service personnel.
These sites are then
evaluated for suitability, first by a winter aerial survey to determine
whether snow-free areas exist, and then by a summer on-the-ground survey
of only those sites which showed snow-free areas.
During the current segment, the Wetterhorn Peak area was found to be
suitable as a goat transplant site on the basis of habitat quality, but
was recommended for rejection because of an indigenous bighorn sheep
population, and the uncertainty of knowledge concerning interspecific
relationships of the two animals.
Sites in the Marcellina Mountain was
found to have sufficient snow-free slopes to warrant an on-the-ground
inspection, while the Ragged Mountain area was found to range from marginal to unsuitable with ~espect to amount of snow-free potential winter
range.
The latter area was recommended for rejection as a possible
transplant site.

��- 67 -

EVALUATION OF PROPOSED MOUNTAIN GOAT TRANSPLANT SITES
William H. Rutherford
P. S. OBJECTIVE
To situate mountain goat transplant opportunities.
SEGMENT OBJECTIVES
To determine habitat characteristics of proposed mountain goat transplant
sites, and compare them with standards established under Work Plan 2, Job
5.

METHODS AND MATERIALS
Potential mountain goat transplant sites are suggested or proposed by a
variety of individuals - Wildlife Conservation Officers, Wildlife Researchers,
and U. S. Forest Service personnel. The first activity toward evaluating
their suitability is a winter survey by aircraft during late February or
March, when snow depth is near maximum. Those sites which show little or
no snow-free area are eliminated from consideration; those showing potential
winter range are scheduled for on-the-ground surveys during the summer months.
During the aerial survey, photographs are made of those snow-free areas which
show vegetation; these photographs are then used as guides for the on-theground survey. Summer surveys are confined to the areas found to be snowfree during winter, and consist of measurements of plant species composition
and density plus notations on nearness of cliffs and/or other rocky terrain
which goats require for escape cover. Also, potential competition with
other grazing animals is noted.

DESCRIPTION OF AREA
This job is statewide in scope, but limited during each segment to those
specific locations proposed as transplant sites.

RESULTS AND DISCUSSION
During the current segment, on-the-ground surveys were conducted in the
Wetterhorn Peak area, which had been reported in the previous segment progress report as showing good potential, based on the results of a winter
aerial survey. Also, winter aerial surveys were conducted in the Marcellina
Mountain and Ragged Mountain areas during the current segment.
The Wetterhorn area is characterized by alpine tundra of expectionally high
quality. Plant density is good, and species composition tends toward high
percentages of the desirable Kobresia, Poa and Festuca. Interspersion of

�- 68 -

rocky cliffs and vegetated areas is good, and there is a minimum of
domestic sheep and elk competition. Winter range is located in the
alpine zone, with no possibility for wintering at lower elevations,
but there appears to be sufficient snow-free areas to support wintering goats. Thus, all ingredients for a successful goat transplant
appear to exist.
The major drawback to transplanting goats in the Wetterhorn Peak area
is the fact that this is a part of the normal range for a herd of bighorn sheep. There is very little documentation on interspecific relationships of these two animals, but the scant evidence at hand indicates that
bighorns do not tolerate mountain goat competition. Thus, it is felt
that this is a rather serious consideration, and that the area should
be rejected as a goat transplant site solely on this basis. Furthermore,
all future transplant site proposals should consider this aspect early
in the evaluation process, and should eliminate from consideration any
area having current or potential use by bighorns.
The Marcellina Mountain proposed goat transplant site was surveyed
aerially on April 10, 1970. This location showed 100 acres or more of
snow-free winter range on the south and west sides of the mountain, plus
Anthracite Canyon on the north side which has a great deal of snow-free
area. Interspersion of cliffs and vegetation is good, the area is judged
to have low potential for bighorn sheep, and is isolated well enough so
that little opportunity for goats to migrate exists. There is no potential
for domestic livestock competition, and nothing more than minor elk competition is foreseen. An on-the-ground survey will be conducted during
the next segment.
The Ragged Mountain, Chair Mountain and Huntsman Mountain proposed transplant site was also aerially surveyed on April 10, 1970. These lie in a
long chain oriented generally north-south. Ragged Mountain at the south
end showed some snow-free areas, but not extensive enough to consider as
a good release site. The further north the survey progressed, the more
complete became the snow cover. In summary, the south end is marginal
and the north end is completely unsuitable as a goat transplant site.
It is recommended that this area not be considered because of the proximity
of the good potential transplant site on Marcellina Mountain.

Prepared

bY*~#

William H. Rutherfo
Wildlife Researcher

�January, 1971

- 69 JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No.
Work Plan No.
Job Title

An Ecological Investigation of the
Cache 1a Poudre Deer Herd, Colorado

W-105-R-ll

Job No.

5

3 (Tenth year)

Harvest Analysis

Period Covered: March 1, 1970 to February 28, 1971
Personnel:

Allen E. Anderson

ABSTRACT

Publication plans our outlined.

��- 71 -

HARVEST ANALYSIS
Allen E. Anderson

P. S. OBJECTIVES
1.

Determine the age structure of the deer herd kill to provide estimates
of (a) net productivity (percent female yearlings in the kill), and
(b) the effects of hunting regulations.

2.

Locate the distribution of the kill by sub-unit to relate age and sex
structure of the kill to elevational levels and harvest intensity.

3.

Measure relevant physical characteristics of the kill to provide a
possible index of herd response to winter food quality.
RESULTS AND DISCUSSION
Progress

Check station sampling of the fall harvest was completed with the 1965
hunting season. Collected data have been tabulated and statistical treatment has been completed. Most of the literature has been reviewed.
Publications

None.

Manuscripts in Preparation
Manuscripts Submitted

None.

None.
Future Plans

Incorporate findings relative to Objectives 1 and 2 in a Department
Special Report. The first draft will be completed during Segment 12.

Prepared by ---L2!/h..u &amp; t2r-ceP/t4(r1L/
Allen E. Anderson
Wildlife Researcher

��January ~ 1971

- 73 JOB PROGRESS REPORT

State of

C~O~L~O~R~AD~O~ _

Project No.

W-l05-R-ll

Work Plan Noo

An Ecological Investigation of the
Cache la Poudre Deer Herd, Colorado

4
--------~------------

Job Title

Job No.

1 (Tenth year)

Population Density and Structure

Period Covered: March 1, 1970 to February 28, 1971
Personnel:

Allen E. Anderson

ABSTRACT

No publications.

Manuscripts submitted and in preparation are listed.

��- 75 -

POPULATION DENSITY AND STRUCTURE
Allen E. Anderson

P. S. OBJECTIVES
1.

2.

Estimate population density on each of five selected study locations
believed to be representative of the lower-winter, middle-winter,
upper-winter, transitional, and summer ranges of the herd to:
(a) relate density to changing environmental conditions CWork Plan
3, Jobs 1 and 3), and (b) elucidate density-elevational relationships,
particularly between years.
Estimate sex and age structure (composition) of the poulation to provide
basic information on herd dynamics.
SEGMENT OBJECTIVE

Tabulate, summarize, analyze, and interpret collected data, review recent
literature, and prepare manuscripts for publication.
RESULTS AND DISCUSSION
Progress
Estimation of herd structure and population densities on selected areas
was completed prior to June 1, 1965. Analyses of herd structure samplestime-weather relationships were essentially completed as Segment 11 ended.
Publications

None.

Manuscripts in Preparation
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1971.
related to mule deer fecal groups on winter range.

Shrub yield and use
J. Wildl. Mgmt.

Manuscripts Submitted
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1971. Mule deer fecal
group counts related to site factors on winter range. J. Range Mgmt.
Complete and submit final manuscripts.

Prepared by
Allen E. Anderson
Wildlife Researcher

��January, 1971

- 77 JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~---------

Project No.
Work Plan No.
Job Title

An Ecological Investigation of the
Cache la Poudre Deer Herd, Colorado

W-IOS-R-ll

Job No.

5

1 (Tenth year)

Physical Characteristics

Period Covered:

March 1, 1970 to February 28, 1971

Personnel:

Allen E. Anderson

ABSTRACT

Publications, manuscripts submitted~ and manuscripts in preparation are
listed.

��- 79 -

PHYSICAL CHARACTERISTICS
Allen E. Anderson

P. S. OBJECTIVE
Measure relevant physiological and morphological characteristics of deer
collected seasonally so that: (a) the response of the herd to its environment can be more adequately interpreted, (b) establish "physiological
norms", and (c) provide basic morphological data as related to sex and age
classes.
SEGMENT OBJECTIVE
Tabulate, summarize, analyze, and interpret collected data, review recent
literature, and prepare final report and publication manuscripts.
RESULTS AND DISCUSSION
Progress
Physiological and morphological studies on four deer collected each month,
1961-1965, were completed April 27, 1965. The number totaled 192 regular
collections plus 33 from supplementary sources. All data were placed on
punch-cards and preliminary analyses of blood and tissue chemistry data
were completed during Segment 10. The review of literature was continued
and about 1,000 references were obtained for project files during Segment 11.
Publications
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1970. Erythrocytes and
leukocytes in a Colorado mule deer population. J. Wildl. Mgmt.
34(2):389-406.
Anderson, A. E., and D. E. Medin. 1970. Vitamin A in the liver and blood
of mule deer. A.A.A.S. and New Mexico Acad. Science, 46th Annual
Meeting, Las Vegas, New Mexico. April 22-25. Abst. p. 37.
Erickson, J. A., A. E. Anderson, D. E. Medin, and D. C. Bowden. 1970.
Estimating ages of mule deer - an evaluation of technique accuracy.
J. Wildl. Mgmt. 34(3):523-531.

Manuscripts in Preparation
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1971. Temporal variation
of vitamin A in the blood and liver of a mule deer population. J.
Mammalogy.

�- 80 -

Anderson, A. E., D. E. Medin, and D. C. Bowden. 1971. Indices of carcass
fat in a Colorado mule deer population. J. Wildl. Mgmt.
Nicolls, K. E. 1970. Cytometry and volumetry of acidophil cells in the
hypophysis cerebri pars distalis of Colorado mule deer (odocoileus
hemionus hemionus) relative to seasons of the photo period and
antler cycles. Z. fur Zellforsch. und Mikros. Anat.
Roughton, R. D.

1970.

Age structure in browse populations.

Ecology.

Manuscripts submitted
Anderson, A. E., and D. E. Medin. 1970. Antler phenology of a Colorado
mule deer population. Southwestern Nat. 15(2 or 3). (Accepted).
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1971. Adrenal ascorbic
acid and adrenal weight in a Colorado mule deer population. J. Wildl.
Mgmt.

Future Plans
Complete and submit final manuscripts to appropriate Journals.

Prepared by

E:

~
cC;.:.dP~
Allen E. Anderson
Wildlife Researcher

�January, 1971
- 81 -

JOB PROGRESS REPORT

State of

~CO~L~O~R~AD~O~

Project No.

W-105-R-ll

Work Plan No.

_
An Ecological Investigation of the
Cache la Poudre Deer Herd, Colorado

5

Job Title

Job No.

2 (Tenth year)

Reproductive Studies

Period Covered:

March 1, 1970 to February 28, 1971

Personnel:

Allen E. Anderson

ABSTRACT

A publication is listed.

No manuscripts were submitted or prepared.

��- 83 -

REPRODUCTIVE STUDIES
Allen E. Anderson
P. S. OBJECTIVE
Determine the reproductive pattern of the deer herd to provide data on:
(a) morphology of the reproductive organs as related to age and season,
(b) gross and net productivity between years, and (c) tentatively, the
relationship of productivity to measured factors of the environment
(Work Plan 3).
SEGMENT OBJECTIVE
Tabulate, summarize, analyze, and interpret collected data, review recent
literature, and prepare final report and publication manuscripts.
RESULTS AND DISCUSSION
Progress
Reproductive data gathering from collected deer was completed April 27,
1965. Ovarian collections from hunter killed deer were completed with
the 1964 hunting season. Most of the data have been compiled, tabulated,
and placed on computer cards.
Publications
Short, Catherine. 1970. Morphological development and aging of mule and
white-tailed deer fetuses. J. Wildl. Mgmt. 34(2):383-388.
Manuscripts in Preparation:
Manuscripts Submitted:

None

None
Future Plans

Initiate analyses.

Prepared by
Allen E. Anderson
Wildlife Researcher

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April, 1971

JOB PROGRESS REPORT

State of

~C~O~~L~O~RAD~~O _

Work Plan No.
Job Title

1

Job No.

16

Pheasant Nest Site Selection Study

Period Covered:
Personnel:

Game Bird Survey

W-37-R-24

Project No.

April 1, 1970 to March 31, 1971

Donald M. Hoffman, Warren D. Snyde.r, Bruce C. Sigler, Dale W.
Stahlecker, Robert L. Schmidt, Lawrence A. Webster, John F.
Corey, Gary Brown, John Ellenberger, Mark Frazier, Jon Hooper,
Tom Lines, Roger Lowery, Jon Moser, Henry Wilson and Michael
Zgainer.
ABSTRACT

Each of the 4 quadrats was stocked with a m~n~mum of 13 hens and 3 cocks
by March 11, 1970 (Segment 23) in preparation for nesting studies in 1970.
Eighty pheasant nests were found in the 48 nesting plots, representing 6
vegetative species or combination of species. Sixteen nests were found in
8 alfalfa mowed plots; 15 nests Were found in 4 alfalfa-crested wheatgrass
mowed plots; 10 nests in 4 tall wheatgrass mowed plots; 9 each nests in
4 intermediate wheatgrass mowed and 4 intermediate wheatgrass unmowed plots;
8 nests in 4 tall wheatgrass seeded plots; and 7 nests in 4 smooth bromegrass mowed plots. Fewer numbers of nests were found in other vegetative
types and treatments represented. In comparing nests per vegetative type
and treatment, alfalfa-crested wheatgrass mowed, ranked highest with 3.75
nests per plot; followed by tall wheatgrass mowed, with 2.50 nests per plot;
intermediate wheatgrass mowed, and intermediate wheatgrass unmowed, each
with 2.25 nests per plot; tall wheatgrass seeded, and alfalfa mowed, each
with 2.00 nests per plot; and smooth bromegrass mowed, with 1.75 nests per
plot. Sixty-six percent of all nests in 1970 were established 15 feet or
less from plot perimeters. Average clutch size of 46 nests, where it was
possible to determine this, was 11.2. A minimum of 13 hens and 3 cocks,
wild-trapped in Phillips, Sedgwick, and Larimer counties in early 1971,
were restocked in each of the 4 quadrats by January 28, 1971 for continuation of the study in 1971 (Segment 25).

��"

-.)-

PHEASANT NEST SITE SELECTION STUDY
Donald M. Hoffman

Field work on this study was conducted similar to the previous year's work
with three separate nest searches completed during the nesting season of
1970. The change-over of three vegetative types used very little or not
practical to establish along roadsides was completed in the fall of 1969 and
no additional p1antings were made in 1970.
P. S. OBJECTIVE
To compare pheasant nesting of, and success in (1) winter wheat; (2) alfalfa;
(3) crested wheatgrass; (4) hairy vetch; (5) white sweet clover; (6) alfalfacrested wheatgrass mixture; (7) smooth bromegrass; (8) tall wheatgrass; and
(9) intermediate wheatgrass.
SEGMENT OBJECTIVES
1.

To maintain plots.

2.

To measure nesting preference.

3.

To determine nesting success by cover type.

4.

To obtain wild pheasants and care for them following release in plots.
METHODS AND MATERIALS

Wild pheasants were captured with a vehicle, spotlights, and hand nets at
night in Phillips and Sedgwick counties in Segment 23. All 4 quadrats were
restocked with a minimum of 13 hens and 3 cocks by March 11, 1970. Limited
Cooper's hawk predation occurred on the wing-clipped pheasants held in the
quadrats after this date but this was not serious.
Phenology measurements were again secured in 1970 to ascertain readiness of
the various vegetative species and cover types for use by nesting pheasants.
An initial nest search in 1970 was made during the period June 4 and 5 with
10 men, including 5 W.C.O. trainees and others. Each plot was systematially
searched using ropes to divide the plots into narrow segments. Nests were
marked, recorded, and removed to encourage renesting. A second nest search
was made during the period June 22 through July 3, 1970 by 2 men searching
each plot systematically in strips outlined by lath markers. A third and
final search was made by 5 men using methods similar to the second search
during the period July 20 through 27, 1970, preceding and following the windrowing of half of the individual plots on July 27, 1970. A well trained
bird dog belonging to Warren Snyder was used to advantage in the first and
last nest searches.

�-4-

Distances were measured from each nest to the nearest edge and to the apex
or base of the triangular plot, whi.chever was closest, with a steel tape.
Each nest could later be located on a scaled diagram of the quadrats.
Number of eggs, fate of the nest and estimated age of embryos were recorded.
In addition, vegetative heights and vegetative species in the 4 cardinal
direc.tions from each nest were recorded.
Prior to the windrowing of vegetation in half of the plots on July 27, 1970,
all but 4 of the 64 wing-clipped pheasants were caught with hand nets. Nineteen of these were released on Means Lake property in Phillips County and
the remainder were released outside the quadrats at the Fort Collins Wildlife
Research Station.
Two incubating hens were killed by the windrowing machine
and the other 2 were not recovered.
Five wild pheasants, including 3 hens and 2 cocks were captured with cloverleaf traps at the Fort Collins Wildlife Research Station during early
January, 1971. Fifty-seven additional wild pheasants, including 49 hens
and 8 cocks were captured with 4-wheel drive vehicles, hand spotlights, and
hand nets during 6 trap-nights in Phillips and Sedgwick counties.
These
were used to restock the 4 quadrats with a minimum of 13 hens and 2 cocks by
January 28, 1971 for continuation of the study in 1971. All pheasants were
wing-clipped and banded with aluminum leg bands prior to release in the
quadrats.

Plot Composition
Plot Composition

for 1970 (Treatment

Following

1969 Nesting

Season)

The changing of vegetative types used very little, or not practical to
establish along roadsides, started in 1968, was continued in the fall of
1969. The remaining winter wheat plots were reworked and seeded to smooth
bromegrass and the 4 remaining white sweet clover plots were reworked and
seeded to tall wheatgrass.
Plot Number

1.

Originally

Winter Wheat,

Now Smooth Bromegrass

Four of the original winter wheat plots were reworked and seeded to smooth
bromegrass in the fall of 1968. These were all mowed in the summer of
1969. The remaining 4 winter wheat plots were reworked and seeded to
smooth bromegrass in the fall of 1969.
Plot Number

2.

Alfalfa

These were all mowed and maintained following the 1969 nesting season so
1970 composition was comparable with 1968 and 1969. Two of these plots
including Number 2, Interior, NW Quadrat,
and Number 2, Exterior, NE
Quadrat, are approaching alfalfa-crested wheatgrass mixtures due to invasion
by crested wheatgrass.

�Plot Number

3.

Cres ted Wheatgras s

Four of these were mowed and 4 were left unmowed following the 1969 nesting
season so that a residual cover check could be secured in 1970 on half of
the plots.
Composition for 1970 was therefore comparable to that of 1969.
Plot Number 4.

Originally

Hairy Vetch,

Now Intermediate

Wheatgrass

All of these plots were reworked and seeded to intermediate wheatgrass
following the 1968 nesting season.
Four of these intermediate wheatgrass
plots were mowed and 4 were left unmowed following the 1969 nesting season,
so a residual cover check could be secured in 1970 on half of the plots.
Plot Number

5.

Originally

White Sweet Clover, Now Tall Wheatgrass

Four of the original white sweet clover plots were reworked and seeded to
tall wheatgrass in the fall of 1968. These were all mowed in the summer of
1969. The remaining 4 white sweet clover plots (volunteer forbs in 1969)
were reworked and seeded to tall wheatgrass in the fall of 1969.
Plot Number 6.

Alfalfa-Crested

Wheatgrass

Four of these were mowed and 4 were left unmowed following the 1969 nesting
season, so that a residual cover check could be secured in 1970 on half of
the plots.
Composition for 1970 was therefore comparable to 1969.

Plot Composition

for 1971 (Treatment

Following

1970 Nesting Season)

The changing of vegetative types used very little, or not practical to
establi8h along roadsides, started in 1968, completed in the fall of 1969.
No additional changes were made during 1970.
Plot Number

1.

Originally

Winter Wheat, Now Smooth Bromegrass

The 4 original winter wheat plots seeded to smooth bromegrass in the fall
of 1968, were left unmowed in the summer of 1970. The 4 original winter
wheat plots, seeded to smooth bromegrass in the fall of 1969, were mowed in
the summer of 1970. A residual cover check will be secured in 1971.
Plot Number 2.

Alfalfa

Four of these were mowed and 4 were left unmowed in 1970 so that a residual
cover check may be secured in 1971. Two of the alfalfa plots including
Number 2, Interior, NW Quadrat, and Number 2, Exterior, NE Quadrat, are
approaching alfalfa-crested wheatgrass mixtures due to invasion by crested
wheatgrass.
Plot Number

3.

Crested Wheatgrass

Four of these were mowed and 4 were left unmowed following the 1970 nesting
season so that a residual cover check may be secured in 1971 on half of
the plots.
Composition for 1971 is therefore comparable to that of 1970.

�-6-

Plot Number 4.

Originally

Hairy Vetch,

Now Intermediate

Wheatgrass

All of these plots were reworked and seeded to intermediate wheatgrass following the 1968 nesting season.
Four of these intermediate wheatgrass plots
were mowed and 4 were left unmowed following the 1970 nesting season, so a
residual cover check may be secured in 1971 on half of the plots.
Plot Number

5.

Originally

White Sweet Clover, Now Tall Wheatgrass

The 4 original white sweet clover plots seeded to tall wheatgrass in the fall
of 1968, were left unmowed in the summer of 1970. The 4 original white
sweet clover plots seeded to tall wheatgrass in the fall of 1969, were mowed
in the summer of 1970. A residual cover check will be secured in 1971.
Plot Number

6.

Alfalfa-Crested

Wheatgrass

Four of these were mowed and 4 were left unmowed folloWing the 1970 nesting
season, so that a residual cover check may be secured in 1971 on half of
the plots.
Composition for 1971 is therefore comparable to 1970.

RESULTS AND DISCUSSION
Stocking

of Pheasants

for 1970

An attempt to live-trap pheasants in September, 1969 in the Holyoke area
was made but losses folloWing capture were heavy.
Of 18 captured (mostly
juveniles) 4 died in transit to Fort Collins and 4 died while held in the
covered pen before moving to the southeast quadrat on October 9, 1969. At
least 2 additional birds from this source are known to have died while held
in the southeast quadrat.
The wing-clipped pheasants used in the 1969 nesting studies were recaptured
with hand nets at the end of the 1969 nesting season and placed in a large
covered pen in an attempt to hold these over for use in 1970. An unusually
heavy, wet snow in early October, 1969 broke the top of the pen down and 67
surviving pheasants (57 hens and 10 cocks) were released in the southeast
quadrat on October 9, 1969. Overwinter losses, mostly from great-horned owl
predation, were heavy.
Approximately 34 percent were lost from this cause
from October 9, 1969 to February 9, 1970 and an additional 33 percent were
lost from the same cause from February 9, 1970 to March 6, 1970. Limited
Cooper's hawk predation on the wing-clipped pheasants occurred after the
final stocking of the quadrats on March II, 1970, but this was not serious.

Location

of Nests

Eighty pheasant nests were found in the 48 triangular-shaped
nesting plots
representing 6 vegetative species or combination of species.
Figures 1
through 4 show the approximate location of nests in the various quadrats.
Table 1 lists pheasant nests found by vegetative cover type and treatments
in 1970.

�-7-

370'

4

2

Agin (urn)
""~

Mesa (m)

o
3
12' Agcr (m)

o
5

0::

Agel (rr)

~

o

0::
W

~

Brin (m)

0::

o
H
0::

~

Z

H

12'

6
Mesa~gcr(urn
Brin (rr)~

12'

Detailed

Fig. 1.

Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan I, Job 16

Nest Site

Location of pheasant nests in northwest quadratp 1970.

~
0
W

-

�-8-

370'

\\

2

\\

5

/00 6
Mesa- Agcr

II

12'

~
~
(l:

. ~1-,.

.:

Agel (rr) /

!o/

Agel (m)

7

7/ o 5

3
o Agcr (m)

(m)

I:t:

o
Brin

o

(m)

Brin (rr) ~

X

£iI

o

12'

4
Agin (

Mesa-Agcr (urn)

2

/

Ager

6

Fig. 2

G

Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan I, Job 16

(um~

3

12'

Detailed

~

~ 8

Nest Site

Location of pheasant nests in northeast quadrat, 1970.

)

-

�-9-

370'

4

7/

7

2
Mesa

(m) /

f
Brin (rr)

~

0::
W

6

E 0
Mesa-Agcr

(m)

~

~o'~

------0
12'

6

12'

Fig. 3 •

Location

Detailed Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan 1, Job 16

Nest Site

of pheasant

1970.

nests in southwest

quadrat,

�-10-

370'

3

6
12'

Mesa-Agcr (urn)

12'

12'

Detailed

Fige4 _

Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan 1, Job 16

Nest Site

Location of pheasant nests in southeast quadrat, 1970

0

�Table 1.

Number of pheasant nests found in 48 plots representing various vegetative types and treatments, 1970.

Plot
Number

Type and Treatment

NW

guadrat
NE
SW

SE

1

Smooth bromegrass (mowed)

1

3

1

1

Smooth bromegrass (seeded)

1

0

2

Alfalfa (mowed)

7

3

Crested wheatgrass

(mowed)

3

Crested wheatgrass

(unmowed)

4

Intermediate wheatgrass

(mowed)

Total

Number
Plots

Nests/
Plot

2

7

4

1.75

0

0

1

4

0.25

2

1

6

16

8

2.00

0

2

0

1

3

4

0.75

0

0

0

1

1

4

0.25

4

1

2

2

9

4

2.25
I

2

1

2

4

9

4

2.25

(mowed)

1

4

4

1

10

4

2.50

(seeded)

4

1

0

3

8

4

2.00

Alfalfa-crested wheatgrass (mowed)

5

2

2

6

15

4

3.75

Alfalfa-crested wheatgrass

1

0

0

0

1

4

0.25

26

16

12

26

80

48

1.66

4

Intermediate wheatgrass

5

Tall wheatgrass

5

Tall wheatgrass

6
6

Totals Average

(unmowed)

(unmowed)

I-'
I-'
I

�-12-

By numbers of nests, alfalfa mowed (16 nests) ranked highest for nest
establishment in 1970, followed by alfalfa-crested wheatgrass mowed (15 nests);
tall wheatgrass mowed (10 nests); intermediate wheatgrass mowed and intermediate wheatgrass unmowed (each with 9 nests); tall wheatgrass seeded (8
nests); and smooth bromegrass mowed (7 nests).
Fewer numbers of nests were
found in other vegetative types and treatments represented.
Numbers of nests per cover plot were calculated because of a difference in
numbers of plots by cover type and treatment.
In this comparison, alfalfacrested wheatgrass mowad ranked highest with 3.75 nests per plot, followed
by tall wheatgrass mowed with 2.50 nests per plot, intermediate wheatgrass
mowed and intermediate wheatgrass unmowed each with 2.25 nests per plot,
tall wheatgrass seeded and alfalfa mowed each with 2.00 nests per plot, and
smooth bromegrass mowed with 1.75 nests per plot.
Sixty-six percent of all nests in 1970 were established 15 feet or less from
plot perimeters as shown in Table 2. During 1970, 48 nests were found in
exterior plots and 32 nests in interior plots.
The terms "interior and
exterior" refer to the location of the individual plots in relation to a
central point where all 4 quadrats join. These are indicated in the margins
of Figures 1 through 4. Figure 5 shows an experimental plot of smooth bromegrass seeded in the fall of 1968 and mowed in the summer of 1969. A search
for nests in a plot of intermediate wheatgrass seeded in the fall of 1968
is shown in Figure 6. Figure 7 shows an experimental plot of tall wheatgrass seeded in the fall of 1968 and mowed in the summer of 1969. This
cover type furnishes good winter cover as well as good nesting cover at
the Fort Collins Wildlife Research Station.
Figure 8 shows the method used
in hand netting Wing-clipped pheasants located with the help of a well
trained bird dog. Pheasants were removed from the quadrats prior to windrowing of the vegetation in half of the plots on July 27, 1970.

Clutch Sizes
Thirty-four nests contained unknown clutch sizes.
These were nests where
the hen was still laying or nests which were abandoned for one reason or
another without incubation having started.
The average clutch size of the
remaining 46 nests which included both incubating nests and hatched nests
was 11.2 eggs, ranging from 5 to 21 (Table 3).
Considerable variation was found in clutch sizes between the various
cover types and treatments as shown in Table 3. Largest average numbers of
eggs per clutch was found in intermediate wheatgrass unmowed (6 nests
averaged 14.5 eggs); intermediate wheatgrass mowed (7 nests averaged 12.4
eggs); alfalfa mowed (9 nests averaged 12.2 eggs); smooth bromegrass mowed
(4 nests averaged 11.0 eggs); and alfalfa-crested wheatgrass unmowed (1 nest
averaged 11.0 eggs).

�I

I-'

W
I

Fig. 5. A plot of smooth bromegrass seeded in the fall of 1968 and mowed in the summer of 1969.
nests were found in the four smoothbromegrass, mowed plots in 1970. (D. Hoffman, photo).

Seven pheasant

�,....I

+:I

Fig. 6. Searching for nests in a plot of intermediate wheatgrass seeded in the fall of 1968 and left unmowed in the
summer of 1969
Nine pheasant nests were found in the four intermediate wheatgrass, unmowed, and nine in the four
intermediate wheatgrass, mowed plots in 1970.
(Do Hoffman, photo).
0

�I
•.....

V1
I

Fig. 7. A plot of tall wheatgrass seeded in the fall of 1968 and mowed in the summer of 1969.
found in the four tall wheatgrass, mowed plots in 1970. (D. Hoffman, photo).

Ten pheasant nests were

�I

~

0'\
I

Fig. 8. Hand netting a wing-clipped pheasant, located with the assistance of a well trained dog.
removed from the quadrats prior to windrowing of the vegetation.
(D. Hoffman, photo).

Pheasants

were

�-17-

Table 2.

Distance of pheasant nests from plot perimeters, 1970.

Distance
From Nearest
Edge

Number
Nests

Percent

0 - 5'

13

16

5 - 10'

25

31

10 - 15'

15

19

15 - 20'

11

14

20 - 25'

6

8

25 - 30'

9

11

30 - 35'

1

1

35 - 40'

0

0

Total

80

100

Fate of Nests
Of the 80 nests found and recorded, 66 were determined to be in an active
state of laying or incubation, 7 were hatched successfully, and 7 were
apparently abandoned for one reason or another. All nests were recorded
and removed to encourage renesting. This also prevented the recording of
the same nest more than once.
Of the 7 nests which hatched successfully, 2 were in alfalfa mowed plots,
2 in intermediate wheatgrass unmowed, 2 in alfalfa-crested wheatgrass mowed,
and 1 in intermediate wheatgrass mowed. Of 86 total eggs in 7 nests which
had successfully hatched, 71 (82.55 percent) of the eggs hatched.
Of the 7 apparently abandoned nests, 1 was hailed on with eggs showing hail
cracks, 1 was ruined by a predator (probably magpie or crow) and 5 were
abandoned for unknown reasons.
Periods of Nest Establishment
Table 5 lists periods of nest establishment based upon aging egg embryos
according to procedures outlined in Game Information Leaflet Number 15
(Sandfort 1965).

�Table 3.

Clutch sizes found in 46 incubating or hatched pheasant nests in various vegetative types, 1970.

Clutch
Size

1
Brin
(m)

1
Brin
(rr)

2
Mesa
(m)

3
Agcr
(m)

1
1

1
1

Plot Number and Treatment
3
4
4
5
Agcr
Agin
Agin
Agel
(urn)
(m)
(urn)
(m)

5

6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22

5

Agel
(rr)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(urn)

1
3
2
1

2

2
2

1
1

1
1

1
1

1
1
1
1

1

1

1
1
1

1
1
2

2
1

1
1
1

2

1
1

Total

1
5
6
6
3
1
3
3
2
6
2
2
1
3
1
0
1

Totals

4

0

9

3

0

7

6

5

3

8

1

46

Average
Clutch

11.0

-

12.2

8.0

-

12.4

14.5

9.4

9.3

9.5

11.0

11.2

I

I-'

00
I

�Table 4.

Fate of 80 pheasant nests in various vegetative types, 1970.

Type

1
Brin
(m)

Ac tive (Laying or
Incubated) 7

1
Brin
(rr)

2
Mesa
(m)

5
Agel
(rr)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(urn)·

Total

I

t-"

1

13
2

Abandoned or
Unsuccessful

1

7

Plot Number and Treatment
5
3
4
4
Agel
Agcr
Agin
Agin
(urn)
(m)
(urn)
(m)

1.0
I

Hatched

Total

3
Agcr
(m)

1

16

3

3

1

1

8

6

1

2

9

9

5

12

1

66

2

7

7

1

1

3

1

9

10

8

15

1

80

�Table S.

Estimated period of nest establishment in various vegetative types, 1970. 11

Period

1
Brin
(m)

1
Brin
(rr)

April 16-30

2
Mesa
(m)

3
Agcr
(m)

Plot Number and Treatment
5
3
4
4
Agcr
Agin
Agel
Agin
(urn)
(m)
(um)
(m)

2

5
Agel
(rr)

1

1

May 16-31

2

3

June 1-15

2

3

1

June 16-30

1

6

2

2

Total

4
2

3

1
1

6
Mesa-Agcr
(um)

1

1

May 1-15

6
Mesa-Agcr
(m)

3

2

3

13

16

5

3

1

1

2

5

1

24

I

N

July 1-15

1

1

1

July 16-31

1

Unknown

3

Total

7

1

16

2

1

6

1

3

1

1

3

1

3

3

9

9

10

8

IS

14
1

80

1/ Period of nest establishment based upon aging of embryos according to Game- Information Leaflet No. IS (Sandfort
1965). First nest search was made June 4-5, 1970; second was made June 22-Ju1y 3, 1970; third was made July
20-27, 1970.

0
I

�-21-

An initial peak in nest establishment occurred during the period May 16-31,
1970 or approximately 2 weeks later than in 1969. Heights of vegetation
on May 15, 1970 were about half that found on May 16, 1969 due to a cold,
wet spring in 1970. Weather factors, therefore, were the apparent cause of
this delay in nesting activity.
Additional peaks in nest establishment
occurred during the periods June 1-15, and June 16-30, 1970; similar to
those found in 1969. Highest number of nests (24) was recorded for the
period June 16-30, 1970. Some hens continued to recycle until removed
from the quadrats prior to windrowing of the vegetation in half of the individual plots on July 27, 1970.
Residual cover from 1969 was left in half of the crested wheatgrass, intermediate wheatgrass, and alfalfa-crested wheatgrass plots (12 plots out of
48) in order to compare nesting use with mowed plots. This residual cover
was flattened by the early, wet snows in September and October, 1969 and
did not recover so, for all practical purposes, standing residual cover was
non-existent in 1970. In 1970, only 3 nests out of the 19 established prior
to May 31, 1970 (15.78 percent) were found in plots having this flattened
residual cover from 1969. A good test of residual cover was not possible in
1970 for this reason.

Phenology
Heights of vegetation measurements were made on April 15 and 27; May 11 and
22; June 2, 12, and 23; and July 3 and 16, 1970. Table 6 summarizes average
heights of vegetation on April 15, May 11, June 12, and July 16, 1970.
Heights of the new growth vegetation on May 11, 1970 were generally about
half that found on May 16, 1969, but had largely caught up by mid-June.
The
cold, wet spring of 1970 probably caused this lateness of vegetation growth.
Weather factors also apparently caused a delay in nesting activity in 1970
(see section on periods of nest establishment).
In contrast to the 1969 season when vegetative growth appeared to reach a
maximum for the season in mid-June, most vegetative species in 1970 continued
to grow in height throughout the nesting season.
This was probably due to
differences in spring and summer moisture and temperatures.
Good early soil
moisture has been maintained for the 1969, 1970, and 1971 nesting seasons
by complete plot irrigations during the previous fall periods.
This was
required to establish new grass seedings prior to 1970, but was continued in
1970 to provide ade::Juatesoil moisture for the early 1971 growing period.

Height
Nl'st!'l
t u n

l

t Y

Surrounding

Nests

found prior to windrowing

and found in uncut plots provided

1'0'-

lu- I ~ht

ob

I n

l n

l n~

'1'111"., I 11Mlr. IIV(~I"I'f!.1'
t Lvv ho l gh t 01
1').'14

Itrs t

of Vegetation

v(~~('lnl
IwllI,htH

I

VI'

Hlld

l nc he a Willi

l1ll'tlFlllrelll('nI:R

'1fltwlclH

Iounrl

of

s u r

r

ounrl I ng

VI'II,,,ltlllon.

au r round f ng

IB ne e ts

an oppor7l~ no s t s •

1\11 IIVI'I'I1II,O

Found

V(1J.1;(llll-

du r l ng

LII ••

search period of June 4 and 5, 1970, compared with uve rag« vege
Lutlon heights of over 20 inches for the second and third nest searches.
I\n
ovo ru Ll average height of 19.41 inches was recorded for vegetation surroundt ng till' 7/. I1I'Rt~~rncn s u r od .
nest:

&gt;

�-22-

Table 6.

Average height of vegetation in pheasant nesting plots, 1970.

Plot
Number

Vegetative Type
and Treatment

4-15-70

1

Smooth bromegrass (mowed)

4.0

10.0

23.0

37.0

1

Smooth bromegrass (seeded)

4.0

12.0

12.0

10.0

2

Alfalfa (mowed)
New growth
Residual

2.0
4.0

9.0

21.0

26.0

3

Crested wheatgrass (mowed)

4.0

9.0

11.0

17.0

3

Crested wheatgrass (unmowed)
New growth
Residual

4.0
4.0

8.0

9.0

14.0

Intermediate wheatgrass
(mowed)

5.0

9.0

14.0

38.0

Intermediate wheatgrass
(unmowed)
New growth
Residual

5.0
3.0

8.0

16.0

36.0

5

Tall wheatgrass (mowed)

6.0

12.0

14.0

42.0

5

Tall wheatgrass (seeded)

2.0

11.0

10.0

14.0

6

Alfalfa-crested wheatgrass
(mowed)
New growth alfalfa
Crested wheatgrass
Residual alfalfa

1.0
4.0
3.0

10.0
9.0

18.0
13.0

31.0
18.0

Alfalfa-crested wheatgrass
(unmowed) !l
New growth alfalfa
Crested wheatgrass
Residual alfalfa

1.0
3.0
3.0

3.0
8.0

9.0
9.0

20.0
16.0

4

4

6

Average Height (Inches)
5-11-70
6-12-70
7-16-70

1/ New growth alfalfa was well clipped by mice in unmowed alfalfa-crested wheatgrass plots during 1970 growing season.

�-23-

Table 7.

Heights and composition of vegetation surrounding 74 nests, 1970.

Nest
Number

Quadrat

Plot
No.

Date
Measured

1
2
3
4
5
6

SE
SE
SE
SE
SE
SE

1-1
1-1
4-E
2-E
2-E
6-E

6-4-70
6-4-70
6-4-79
6-4-70
6-4-70
6-4-70

7
8
9

SW
SW
SW

5-E
5-E
6-E

6-4-70
6-4-70
6-4-70

10

NE

5-1

6-5-70

11

NE

6-E

6-5-70

12
13

NE
NE

1-1
5-1

6-5-70
6-5-70

14
15

NW
NW

2-1
5-1

6-5-70
6-5-70

16

NW

4-1

6-5-70

17

NW

1-E

6-5-70

18

NW

6-E

6-5-70

19
20

SE
SE

5-1
4-1

6-22-70
6-23-70

21

SE

4-1

6-23-70

22
23
24
25
26
27

SE
SE
SE
SE
S8

3-1
2-1
2-1
4-E
4-8
5-E

6-23-70
6-24-70
6-24-70
6-25-70
6-25-70
6-25-70

28

SE

29
'\0

81':

5-E
6-E

6-25-70
6-25-70

~I':

(,-'1')-70

II

nit;

()-I':
(,-I':

Composition

Ave. Ht.
(In.)

Smooth bromegrass
Smooth bromegrass
Intermediate wheatgrass
Alfalfa
Alfalfa
Alfalfa
Crested wheatgrass
Tall wheatgrass
Tall wheatgrass
Alfalfa
Crested wheatgrass
Tall wheatgrass
Cheatgrass
Alfalfa
Crested wheatgrass
Smooth bromegrass
Tall wheatgrass
Cheatgrass
Alfalfa
Cheatgrass
Thistle
Intermediate wheatgrass
Hairy vetch
Smooth bromegrass
Cheatgrass
Alfalfa
Crested wheatgrass

13.0
14.3
15.0
23.0
22.5
16.3
12.0
16.3
15.3
16.5
12.0
23.0
27.0
15.3
11.0
10.3
13.0
11.3
19.8
14.8
9.5
15.0
12.0
15.0
14.0
15.0
12.0

Cheatgrass
Intermediate wheatgrass
Cheatgrass
Intermediate wheatgrass
Cheatgrass
Crested wheatgrass
Alfalfa
Alfalfa
Intermediate wheatgrass
Intermediate wheatgrass
Tall wheatgrass
Cheatgrass
Cheatgrass

13 .0
24.0
12.7
26.8
13 .3
14.3
24.0
25.5
24.3
23.5
29.5
21.5
20.0
21.5

Ave. Ht.
for
Period (In.)

15.34

SE

(,-,/'j-10

Alfalfa
Alfalfn
A 1 III 1 I'll
Cn'Hl(~d

77.3
,/11 • ')
wlwntKrllHH

/.1 .1

----------------------------------------------------------------------------------

�-24-

Table 7. Heights and composition of vegetation surrounding 74 nests, 1970.
(continued) •

Nest
Number

Quadrat

Plot
No.

Date
Measured

32

SW

5-E

6-25-70

33

SW

6-E

6-26-70

34
35

SW
NW

4-1
4-1

6-26-70
6-29-70

36

NW

4-1

6-29-70

37

NW

5-E

6-29-70

38
39

NW
NW

2-E
6-E

6-29-70
6-30-70

40

NW

2-1

6-30-70

41

NW

6-E

6-30-70

42
43
44
45

NW
NW
NE
NE

5-1
5-1
1-1
5-1

6-30-70
6-30-70
6-30-70
7-2-70

46
47
48

NE
NE
NE

3-E
3-E
4-E

7-2-70
7-2-70
7-3-70

49

SE

6-E

7-20-70

50

SE

6-E

7-20-70

51

SE

5-E

7-20-70

52

SW

5-E

7-21-70

53
54
55
56
57

SW
SW
SW
SW
SW

1-E
4-E
4-E
4-1
2-E

7-21-70
7-21-70
7-21-70
7-21-70
7-21-70

58

SE

3-E

7-21-70

Composition

Ave. Ht.
(In.)

Tall wheatgrass
Cheatgrass
Alfalfa
Crested wheatgrass
Intermediate wheatgrass
Intermediate wheatgrass
Hairy vetch
Intermediate wheatgrass
Hairy vetch
Cheatgrass
Tall wheatgrass
Cheatgrass
Alfalfa
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Cheatgrass
Cheatgrass
Smooth bromegrass
Tall wheatgrass
Cheatgrass
Crested wheatgrass
Crested wheatgrass
Intermediate wheatgrass

29.0
16.0
25.5
21.0
33.0
26.0
8.3
31.3
16.3
16.0
32.7
20.0
19.5
23.0
16.5
17 .5
21.5
23.3
18.0
17.0
15.8
17.5
21.5
14.0
11.3
14.5
25.5

Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Tall wheatgrass
Winter wheat (volunteer)
Tall wheatgrass
Cheatgrass
Smooth bromegrass
Intermediate wheatgrass
Intermediate wheatgrass
Intermediate wheatgrass
Alfalfa
Crested wheatgrass
Crested wheatgrass

18.7
15.0
19.3
14.3
32.7
36.0
20.5
15.0
37.0
34.5
39.0
36.0
19.0
12.0
15.0

Ave. Ht.
for
Period (In.)

20.98

----------------------------------------------------------------------------------

�Table 7. Heights and composition of vegetation surrounding 74 nests, 1970.
(continued) •

Quadrat

Plot
No.

Date
Measured

59

NW

4-E

7-22-70

60

NW

4-E

7-22-70

61
62

NW
NW

2-E
4-1

7-22-70
7-22-70

63

NW

6-1

7-22-70

64

NW

I-I

7-22-70

65
66
67

NW
NW
NW

2-1
2-1
6-E

7-22-70
7-22-70
7-22-70

68

NE

2-E

7-23-70

69

NE

6-E

7-23-70

70

NE

4-1

7-23-70

71

NE

I-I

7-23-70

72

NE

5-E

7-23-70

73
74

NE
SE

5-1
2-E

7-23-70
7-23-70

Nest
Number

Composition

Ave. Ht.
(In.)

Intermediate wheatgrass
Cheatgrass
Intermediate wheatgrass
Cheatgrass
Alfalfa
Hairy vetch
Cheatgrass
Alfalfa
Crested wheatgrass
Smooth bromegrass
Cheatgrass
Alfalfa
Alfalfa
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Intermediate wheatgrass
Cheatgrass
Smooth bromegrass
Cheatgrass
Tall wheatgrass
Cheatgrass
Tall wheatgrass
Alfalfa

26.0
lO.7
34.7
14.0
14.5
9.5
lO.O
19.0
14.0
6.0
14.0
23.8
13.0
17.5
14.0
18.5
17.3
27.5
18.0
33.0
14.0
14.5
14.7
14.0
19.0
45.0
17.0

Ave. Ht.
for
Period (In.)

20.40
Average Height All Measurements

19.41

Stocking of Pheasants in Quadrats for 1971
1\ ml n I.mumof

13 hens and 2 cocks, wild-trapped in Phillips, Sedgwick and

Lllrllller cotlntiel'!
1n January, 1971, were wing clipped, 1ep;banded, and

rc l onscd In each of the 4 quadrats by January 28, 1971 for con tt.nuat Lon of
the 8 tudy in 1971 (Segment 25).

�-26-

KEY TO ABBREVIATIONS
Figures 1 Through 4 and Tables 3 Through 5
Plants
Symbol

Scientific Name

Agcr

Agropyron cristatum

Crested wheatgrass

Agel

~gropyron elongatum

Tall wheatgrass

Agin

Agropyron intermedium

Intermediate wheatgrass

Brin

Bromus inermis

Smooth bromegrass

Mesa

Medicago sativa

Alfalfa

Connnon Name

Treatment
Following 1969 nesting season
(m)

Mowed only

(rr)

Reworked and reseeded

(urn)

Unmowed - residual cover check in 1970

LITERATURE CITED
Sandfort, W. W. 1965. Aging pheasant embryos. Colo. Game, Fish and Parks
Dept. Outdoor Facts, Game Information Leaf1. No. 15. 2 p.

Prepared by

Ll~WJ~~
Donald M. Hoffma~
Wildlife Researcher

�-27April, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RAD~~O~·_

Project No.

W-37-R-24

Game Bird Survey

Work Plan No.

1

Job Title

Pheasant Roadside Cover Evaluation Study

Period Covered:
Personnel:

Job No.

18

April 1, 1970 to March 31, 1971

Dale W. Stahlecker and Warren D. Snyder.

ABSTRACT

Roadside seeding efforts to establish grass and grass-legume plots were
continued in 1970. Satisfactory stands were attained on twenty additional
plots, which will be included with their controls in the 1971 evaluation.
Nest searches through the summer measured nesting attempts and successes
on 48 seeded plots, 28 farmed controls and 23 unfarmed natural controls.
TWenty-two of the 29 nest sites were found in seeded plots, two were along
farmed roadside shoulders and five were in unfarmed controls. However,
most nests in seeded plots were found along the unseeded roadside edges
where green and residual weed cover dominated. Mammalian predators
destroyed 16 of the 29 nests, four were hatched, four were abandoned because of human disturbance and five could not be accurately classified.
Accumulations of residual grass and legume cover will potentially enhance
the test sites for nesting pheasants during successive years. Partial evaluation of cover conditions and nesting in older roadsides, previously seeded
by farmers, indicate new species should be tested to provide improved nesting cover.

��-29-

PHEASANT

ROADSIDE
Warren

COVER EVALUATION

STUDY

D. Snyder

P. S. OBJECTIVE
To compare pheasant production under natural roadside conditions with the
following cover types to be established along roads.Ldes: (1) grass and
(2) grass-legume mi.xtures.

SEGMENT
1.

To establish

2.

To measure

OBJECTIVES

grass and grass-legume
pheasant

production

cover on roadside

in test and control

test areas.

areas.

METHODS "AND MATERIALS
The reader is referred to Snyder (1969 and 1970) for methods
used in selection of roadside test areas and in establishing
grass-legume cover.

Measurement

of Pheasant

Production

in Test and Control

and materials
grass and

Plots

Nesting by pheasants in the seeded and control roadside plots was measured
by nest searches.
Dale Stahlecker, student assistant, began searches in
mid-June and continued through the summer.
Linear strips, in which width
varied in relation to cover density, were searched on foot. A stick was
used to part the vegetation for close, methodical inspection.
Nests, where
the hen was laying or incubating eggs, were marked by placing a lath nearby along the road shoulder.
These were checked again at later dates to
determine the fate of the nest.
All plots initially searched before August
I were searched a second time during the summer.
Characteristics
of the
nest site and nest were recorded, and shell fragments and nest bowls of
terminated nests were removed.
Pertinent items recorded at each nest included: date; group and plot location; location within plot; plot cover
type; cover type, height and density immediately surround the nest; status
of nest (laying, incubating, number of eggs, etc.); and fate of nest after
rE'ci1pckA.

�-30-

RESULTS AND DISCUSSION

Establishment

of Grass and Grass-Legume

Cover

Past Seedings
Marginal to satisfactory stands of grass and alfalfa were established
following seeding efforts in 1969 on a majority of the roadside study
plots.
Most of the plots, seeded during the spring of 1969, where seed
competition was severe in summer, produced disappointing stands.
Several
of these were reseeded again in 1969, others were replanted in 1970, and
a few were included in the first year evaluation.
Reseeding will be renewed on a few in 1971 attempting to increase grass and legume densities.
Fair to good grass stands, utilized in the 1970 nest searches, were
obtained west of Holyoke following August, 1969 rains.
Table 1 summarizes
the groups of plots utilized in 1970 nest searches.
Each group contained
either one or two seeded plots and one farmed control.
An unfarmed control was included in most groups.

Seeding

Efforts

in 1970

Approximately
twenty additional study plots will be available for nest
searching in 1971 due to seeding efforts in 1970. These were established
despite poor planting conditions.
Early spring seeding efforts were stymied
by wet soil conditions which prevented working the plots.
Several plots
prepared and seeded in early May produced suitable grass and legume stands.
Insects, destroyed one half-mile stand of grass, and one farmer's summer
employee destroyed another plot which was successfully reseeded.
Weed
competition and hot, dry summer weather reduced stand densities on some
spring seeded plots.
Only one inch of rain was received during the three-month period of midJune to mid-September.
Needless to say, seedings attempted during this
period did not germinate.
But, satisfactory grass stands were obtained
on several plots following September rains. Many sprouting alfalfa seedlings
in these plots were killed by frost while in the two-leaf stage.

Problems

in Establishment

of Grass and Legume

Stands

The primary problem in obtaining seeded stands was to place the seed in the
ground prior to receiving moisture that would germinate and sustain the
stand.
Firm soil was needed but hard rains frequently crusted soil preventing
sprouts from getting through.
Adequate moisture, too often, was followed by
hot-winds which dried the soil desiccating sprouting plants.
Weeds along
steep road shoulders also sapped moisture needed by seeds and plants along
the road edges.
Competitive weeds or loose soil along the field bordering
edge of the strips hampered seed germination.
Spring seedings were conSistently hampered by summer weed growth.
Efforts to suppress weed growth
by mowing resulted in distrubance and possible mortality of newly-hatched

�-31-

pheasant broods which would not leave the dense cover.
In two instances
pheasants attempted to nest in spring-seeded roadsides.
One of these
was killed on the nest while mowing.
In general, the best stands were
obtained from plots sunnner-fallowed and seeded in late July or August.

Measurement

of Pheasant

Production

in Seeded and Control

Plots

Tables 1 and 2 sunnnarize the results of nest searching on 48 seeded plots
and 51 control plots in 1970. The total number of nesting attempts, 29,
was considered low. Twenty-two of these occurred within seeded plots but
only seven hens actually utilized seeded grass or alfalfa.
Most of the
remaining 14 nests in the seeded plots were associated with various combinations of weeds and/or cheat grass cover.
Frequently, nests were
located at the base of unseeded road shoulders where residual weed cover
provided a protective canopy.
Kochia, cheat grass, wild lettuce and sunflowers were the most connnon annuals along unseeded or unfarmed portions
of all roadside plots.
Five nests located in the unfarmed (C2) controls were all associated with
annual forbs as were two nests along road shoulders in farmed (Cl) controls.
None were observed in grain stubble or fallow portions of the farmed cont ro Ls ,
Preliminary nest location results indicate residual cover was an important
attraction factor in nest site selection during the early spring.
Increased
use of green vegetation by hens was observed in late spring - sunnner nesting
efforts.
Most seeded stands lacked residual cover in 1970, therefore, they
may become more attractive to pheasants in future years.
Unfortunately,
two half-mile strips of seeded roadside were burned by farmers during the
early spring of 1971, destroying all residual cover.
Skunks, badgers and other unknown predators destroyed 16 of the 29 nests
located in the study plots.
Crushed and scattered shell fragments were
the main characteristic of destroyed nests.
Nest bowls were occasionally
torn apart.
In one instance, not included in this sample, pheasant remains
and shell fragments indicated that a hen was captured on the nest.
Past
observations have shown that skunks spend considerable time working roadsides in spring.
Undoubtedly, they are the primary predator affecting
roadside nesting success.
Four nests hatched successfully.
Nest search efforts, which flushed several
hens from their neets, were partially responsible for nest abandonment.
Overzealous checks to determine If hens had returned to their nest:.•increased
nb andontuon t • More rca t rnLnt will
be placed on follow-up nest checks in the
future.
Eggs were collected from abandoned nests to monitor for mercury and
pesticide residues.
Treatment of roadsides in 1970 by county maintenance personnal included mowing
along the upper edge of the road shoulder in mid and late summer to keep weed
cover below the level of the road surface.
Seeded stands were not mowed.
Few
if any of the unfarmed controls were mowed beyond the road shoulder.
Most of
th(' shoulder cover along the farmed controls which usually was only two to slx
foo'
In wIdth
WIlH
e l f m l nn t e d by 1'1111.
A few apo t a cont n l nt ng bind
weed,
Cnn ad I an t hLn tl.e , or poverty weed were sprayed by t.he county.

�Table 1.

Roadside treatment and control study plots nest searcheddtiriiigthe

Group Code_
!=l.o

.c

Q)

(/)

§

s::

or!

1-4

~

Z

E-i

1
2
1
2
1
1
2
1
2
1
1
3
4
5
6
7
8
9
10
12
13
14
1
1
2
1
2
1

7
7
6
6
7
8
8
10
10
11

7
7
7
7
7
7
7
7
7
7
7

7
6
7
7
7
7
6

Seeded Plot 1,.f

Q)

eo

!
42 (43)
42 (43)if
44
44
44 J)
44 J)
44
44 !if
44 if
44 (43)if
45 if
45 2:/
45 Jj
45
45
45
45
45
45 !if

45 (46)
45 (46)
45 !if
46 J)

46
46
47 i/
47 !if

48 4/

s

or!

or!

.j.J

.j.J

1-4
Q)

CJ

III
CJ

Q)
C/)

~.

13
24
11
11
21
21
28
12
28
30
12
15
15
9
8
17
17
7
7
18
18
18
35
13
14
9
9
17

0

:&gt;
0

u

s::

s::

0

0

or!

(/)
.j.J

or!

.j.J

.j.J

1-4
Q)

(/)

CJ

Q)

Q)
C/)

III
CJ

Z

Control Type 1
(Farmed~

Seeded Plot
s::

0

0

~

:&gt;
0

u

Sl1nimerof 1970.

s::

(/)
.j.J

or!

0

.j.J

.j.J

(/)

CJ

III
CJ

Q)

Q)
C/)

~

Z

21/g1
---------2
g 1 P
---------_.
7
gl 1 P
---------6
g
11 6 gl
8
21 8 gl
8
5
g
21 5 g
1 Pa,l A
6
g
28 6 gl
5
g
---------_.
5
g
----------6
g
30 6 g
7
g
12 7 8
g
4
15 3 gl 1 P
7
g
15 7 gl
5
8 1 U
---------3
g
8 3 gl
8
81 1 P, 1 A 17 8 gl 1 P
6
17 6 g
8 1 U
2
7 2 gl
8 1 S
3
g
7 3 g
5
18 5 gl 1 A, 1 P
8
6
g
18 6 gl
7
g
18 7 g
1
g
35 1 gl
8
gl 1 P, 1 U 13 8 gl 2 P
7
81 1 S
14 7 gl 2 P, 1 S
3
g
9 3 g
1 S
2
g
---------_ ..
1
---------_ ....
s

if

18
19
11
10
16
20
28
11
21
30
1
21
14
4
8
8
17
8
8
13
13

s::

0

or!

0

5
5
8
1
3
3
5
3
3
5
4
8
6
4
4
4
5
5
5
2
1
5
8
7
3

Control Type 2
(Unfarmed)
s::
0

0

or!

(/)
.j.J

or!

.j.J

.j.J

(/)

CJ

III
CJ

Q)

Q)
C/)

z

0

~

30
24

7
2

10
21
20
29
12

1
1
2
1
6

-------

1 U

-------

-------

35
13
10
15
15

7

16

6

-------

7

(/)
Q)

z

-------

15
15
9
8
17
17
17
7
18
18
18
26
8
14
10
10

7

(/)
.j.J

1 U

1 P

4
7
5
2
1
7
7
4
4
4
8
3
4
6
4
4

I

w

I'.)
I

1 A

2 Pa
1 P

Each seeded plot represents 1f4 mile of roadside.
~/ Seeded stand marginal to poor.
1/ Each 1/2 mile of section periphery was numbered in consecutive order, starting at the northeast corner and
running clockwise around the section to locate the study plot. Letter abbreciations are presented as follows:
g = grass, 1 = legume, A = abandoned nest, P = predator destroyed nest, Pa = predation after search, U =
undetermined nest fate, and S = successful nest.
~/ Grass stands seeded in previous years by resident farmers.

�-33-

Table 2. Characteristics
of pheasant
plots during the summer of 1970.

Part 1.

Fate of 29 pheasant

Successful

nests

nesting

attempts

in roadside

found along roadsides

4

--------------------------------------

Predator

destroyed

prior to search--------------

13

Predator

destroyed

after search ----------------

3

Abandoned

(primarily

Unknown

Part 2.

due to human

of nest attempts

Plot Type

Seeded

roadside

Farmed roadside

plots
controls
controls

(Cl)

Unfarmed

roadside

Part 3.

Cover type locations

Seeded

4

disturbance)--

5

----------------------------------------

Plot location

(C2)

and successes

No. Plots

Nest Attempts

Nests/Plot

48

22

0.46

28

2

0.07

23

5

0.22

of nest attempts

and successes

Cover Type

Number

grass dominant

4

Seeded alfalfa

dominant

Mixture of green and residual
cheat grass

3
weeds

study

or

22

�-34-

Characteristics

of Old Grass Stands for Pheasant

Production

Several plots, seeded by farmers in previous years, were utilized in the
study (Table 1). Two of these contained a western-crested
wheatgrass
combination; two contained dense stands of smooth brome; ten were composed of intermediate wheatgrass, and one contained Russian wild rye.
Most possessed near pure stands of the species planted and were attractive
in general appearance.
Pheasant production on these plots was not encouraging.
One successfully
hatched nest was located in an open stand of intermediate wheatgrass.
One
predator-destroyed
nest was found under a canopy of Russian wild rye.
Empirical evaluation indicated the brome, Russian wild rye and the crestedwestern wheatgrass combination generally did not possess adequate heights
to attract pheasants although densities were suitable.
In contrast, intermediate wheatgrass did not provide adequately dense foliage after becoming
root-bound.
The above mentioned grasses, like most species tend to decrease in height as
they increase in density and competition for moisture over a period of years.
Therefore, new stands of crested or western wheatgrass and brome may provide
attractive nesting cover only for two or three years after they are planted.
Intermediate wheatgrass may consistently produce adequate heights, but leaf
foliage decreases as plant density increases.
These and other species also
lodge under winter snows, diminishing their value for winter cover and spring
nesting cover (Fig. 1). Tall wheatgrass, which possesses a stiff strong
stem, does not lodge as readily (Fig. 2). It may prove to be a desirable
species where it will not hold snow to block roads.
Alfalfa, by itself,
provides favorable spring nesting cover, but loses its foliage in late
summer and is flattened by winter snow (Fig. 3). Present observations indicate that alfalfa-tall wheat mixtures produce a good cover combination,
but more evaluation is needed.
Considerable variation in soil and moisture conditions exits along Northeast
Colorado roadways.
Therefore, the same species may provide poor to excellent
nesting cover depending on the site. Roadsides along newly graded roads
lack humus and other nutrients characteristic of productive soils.
Germination and growth of seeded plants are reduced, so it may be preferable
to let weeds grow for a year to two before seeding newly elevated roadways.
Early spring fertilization could also be used advantageously here.

Loss of Roadside
While this study progresses, many roadside nesting and survival areas continue to be eliminated by back sloping of grader ditches and farming to the
road shoulder.
This occurs on old roadways as well as newly elevated ones.
Figure 4 illustrates elimination of roadside on a newly elevated road where,
in the foreground, wheat has already been seeded.
The stubble field will
be worked to the road shoulder in spring.
Most county roads in Phillips and
Sedgwick counties do not contain any roadside cover.

�-35-

Fig. 1. An old naturally reseeded roadside. Snows have lodged grass in
the grader ditch, however, weeds along the fence provide limited winter
cover for pheasants.

�-36-

Fig. 2. Tall wheatgrass holding snow along a seeded roadside.
Little
winter cover for pheasants is available to pheasants under these conditions.

�-37-

Fig. 3. Alfalfa along a roadside provided cover during the fall of 1970
before snows flattened it.

�-38-

Fig. 4. A newly elevated road showing back-sloping of the grader ditch
in preparation for farming to the road shoulder.
Pheasants have no place
to live or nest where this condition exists.

�-39-

In Phillips County, phone lines are presently being put underground,
eliminating roadside poles and providing additional areas for farmers
to eliminate roadside cover. In some instances poles bordering seeded
areas already have been eliminated. Hopefully, these plots can be
retained for use during the remainder of the study.

LITERATURE

CITED

Snyder, W. D. 1969. Pheasant roadside cover evaluation study. Colo.
Div. of Game, Fish and Parks. Game Research Report. April.
pp , 37-45.
1970. Pheasant roadside cover evaluation study. Colo. Div.
of Game, Fish and Parks. Game Research Report. April. pp.
109-114.

Prepared

by

U.r1t.,J
Warren D. Snyder
~,.)

Wildlife Researcher

��-41April, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RA~D~O~
__~----

Project No.

W-37-R-24

Work Plan No.
Job Title

Period Covered:
Personnel:

Game Bird Survey

3

Job No.
8a
Effects of Sagebrush Control
on Distribution and Abundance of Sage Grouse

April 1, 1970 to September 30, 1970

Warren Snyder, Larry Roper, Terry May, Bruce Sigler, Jerry
Whittaker, George Bock, David Croonquist, Don Gore, Courtney
Crawford, John Monarch and Howard Funk.

ABSTRACT
Counts of strutting grounds on the North Park study area showed peak numbers
of male sage grouse present about mid-April, 1970, about average for strutting
ground activity for the area. Numbers present peaked at 274 males, significantly below 1969, but otherwise the highest total since 1964. Data from check
stations operated at Cowdrey and Walden indicated hunting pressure was again
high with 564 hunters checked. However, harvest dropped to about half of the
1969 total with only 266 birds brought through the stations. Immature birds
made up 45 percent of the bag, an increase over the 30 percent of 1969, and
about average for the period since 1955 when check station activity was initiated in North Park.

��-43-

EFFECTS OF SAGEBRUSH CONTROL
ON DISTRIBUTION AND ABUNDANCE OF SAGE GROUSE
Howard D. Funk

An investigation of sage grouse abundance and distribution near Lake John
in North Park has been in effect since initiation in 1963. The job relates
to various specific study topics in various degrees of intensity.
More
recent study has been with regard to the immediate effects of a sagebrush
spraying project, accomplished in 1965, on sage grouse populations with a
study on the long-term effects of this spray project scheduled for a twoyear period beginning the spring of 1973. In the interim period, efforts
are to be limited to collection of study area strutting ground data,
banding samples of birds on these grounds, and operating check stations
near Walden and Cowdrey to obtain harvest data.
This report covers these
items.

P

S. OBJECTIVE

To determine the effects of 2,4-D spraying of sagebrush en: (1) sage grouse
abundance, (2) sage grouse distribution, and (3) vegetative composition and
density.

SEGMENT
the seasonal

OBJECTIVES

1.

To investigate
is controlled.

distribution

2.

To investigate sage grouse numbers
to the area treated.

of sage grouse after sagebrush

by age and sex classes

METHODS

AND MATERIALS

Strutting

Ground Counts

on and adjacent

Wildlife Conservation Officers conduct counts on a number of strutting
grounds in North Park annually.
Usually, three early morning counts are
made on each ground.
Data from these counts are gathered from the Regional
Biologist, high counts for each ground being reported herein.

Check Stations
Two check stations near Walden and Cowdrey are operated the first two days
of the season to collect various data on harvest such as harvest success,

�-44-

age, and sex ratios, banded bird information, and hunter participation for
a check on the year in comparison with previous years.
Project personnel
are responsible for the station with Conservation Officers also contributing
data from field checks.
Main emphasis is on checking harvest within the
study area.
Age and sex of birds are determined by methods described in
previous reports.

RESULTS

AND DISCUSSION

Strutting

Ground Counts

Results of strutting ground counts are displayed in Table 1. Peak counts
occurred about mid-April on most areas, which is about average since strutting
ground activity is subject to snow conditions in the area. Data on peak
numbers of females and total birds were not gathered for this report, but
should be obtained from the field men for continuity of data when preparing
for the final report.

Table

l.

Strutting

Peak counts of male sage grouse on strutting
1/
Ground-

SG

1

0

SG

2

8

SG

3

0

SG

4

SG

1970.

High Count Da te

April

14

66

April

14

5

57

April 21

SG

6

19

April

l3

SG

7

0

SG

8

7

April

14

SG

9

97

April

15

SG

10

16

April

15

SG

11

4

April 8

SG

12

0

Total

1/

Number Males

grounds,

Locations

274

of grounds

shown in previous

reports.

�-45-

Comparative data on annual high counts by strutting ground are presented in
Table 2. Most grounds show a definite decline in 1970 in numbers of males
present with the total males on all grounds about 25 percent below the 1969
total. However, the 1970 total remains the highest count on record, with
the exception of 1969, since 1964 when 337 males were observed. The year
of the indicated "crash" in population numbers was 1965 when males totaled
only 166. This was the year of the spray project, but spraying did not
take place until after the spring counts.

Table 2. Comparison of 1960 through 1970 strutting ground counts of male
sage grouse.
Strutting
Ground

1960

1961

1962

1963

Year
1964 1965

1966

1967

1968

1969

1970

SG

1

10

5

0

0

0

0

0

0

0

0

0

SG

2

43

4

7

20

26

9

9

10

2

8

8

SG

3

14

0

0

0

0

0

0

0

0

0

0

SG

4

130

96

219

216

120

53

47

69

58

82

66

SG

5

19

2

52

17

65

52

54

59

52

82

57

SG

6

5

1

8

9

3

0

2

12

36

36

19

SG

7

0

7

5

0

0

0

0

1

3

2

0

SG

8

5

0

0

0

0

o·

6

5

2

6

7

SG

9

167

109

71

85

99

52

97

81

71

117

97

SG

10

17

77

28

19

17

0

6

9

0

16

16

SG

11

12

50

5

11

7

0

13

15

6

5

4

SG

12

14

12

0

0

0

0

0

0

1

3

0

446

363

395

377

337

166

234

265

231

357

274

Total

Banding
Due to lack of manpower and available time, banding of samples of birds on
strutting grounds was not accomplished during the segment.

�Table 3.

Comparison of North Park sage grouse hunter check information, 1963-1970. 11

Total
Birds

Birds
Per
Hunter

Hours
Hunted
Per Bird

58

506

1.03

4.86

45

41

179

.82

3.49

26

26

51

116

.77

5.40

56

45

71

44

263

.86

4.67

67

42

46

33

267

.89

4.41

Year

Bag
Limit

Hunters
Checked

Hours
Hunted

Adult
Males

1963

3

492

2,460

62

150

42

113

181

1964

2

217

624

25

81

59

28

2

150

626

27

30

49

1966

2

306

1,227

31

116

1967

2

300

1,177

50

127

1965

u

Adult Percent
Females Adults

Juvenile
Males

Juvenile Percent
Females Juveniles

1968

2

546

2,604

80

135

42

156

141

58

512

.94

5.09

1969

2

662

2,936

121

277

70

57

114

30

569

.86

5.16

1970

2

564

2,617

58

89

55

53

66

45

266

.47

9.84

11 Based on Walden and Cowdrey check station data during opening weekends.

II A one day season; hence only a one day check station.

I
.po

0\
I

�-47-

Check Station

Results

Project personnel checked 266 sage grouse through check stations on September
11-12, 1970, the first two days of the season.
This was slightly less than
half the number checked in 1969. Number of hunters also decreased but not
to the extent of the harvest, thus increasing greatly the number of hours
spent per bird in the bag.
Percent immatures in the bag increased to about
45 percent, which was about average over the years.
No valid reason can be given for the sharp decline in harvest in 1970 or
the sequence of events over the past three years.
In 1968, the high count
on grounds was 231 birds, indicated havest was the highest since 1963 at
512 birds, and the bag contained 58 percent young, again the highest since
1963. Count results on the grounds increased to a high of 357 males in
1969, harvest increased to 569 birds, but the harvest contained only 30
percent young.
Then, 1970 count data on grounds produced 274 males, lower
than 1969 but higher than 1968, while harvest dropped to 266 birds but
immatures made up 45 percent of the bag. Adult females were harvested very
heavily in 1969
(48.6 percent of the bag), quite heavily in 1970 with 33.5
percent, and rather lightly in 1968 with only 26.4 percent.
Hunting pressure
was fairly even through these years with 546, 662, and 564 hunters per year
since 1968. Probably a variety of factors have been responsible for this
confusing sequence of data, including poor sampling in one or more areas of
data collection, including check stations.
Only two bands were turned in at check stations (1662-R466 and 1562-R498) as
compared to 23 the previous year.
The former was banded as a subadu1t male
on April 25, 1969 and the latter as an adult male on April 10, 1968, both
on strutting ground 9 (SG9). Both birds were shot north of Lake John the
first weekend of season.

Prepared

by
Howard D. Funk
Section Chief, Small Game Research

��-49-

April, 1971

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~---------

Project No.

Game Bird Survey

W-37-R-24

Work Plan No.

10

Job Title

Study of Hungarian Partridge Adaptability

Period Covered:
Personnel:

Job No.

1

April 1, 1970 to March 31, 1971

Donald M. Hoffman, Bruce C. Sigler, Dale W. Stahlecker, John F. Corey
and Charles E. Brown.

ABSTRACT
Six hundred and seventy four more pen raised Hungarian partridge were released
in Moffat County at 3 sites on April 24, 1970. This brings the total number of
Huns released in Moffat and Routt counties to 1,788 during the period of 19641970
0

Intensive field searches were made by W-37-R Project personnel during the months
of June, August and September, 1970. Only 10 Huns were observed by Project
personnel, these near 2 of the 3 1970 release sites. Some of the released Huns
were seen in the vicinity of all 3 release sites from May through August, 1970
by local residents. Although previously planted areas were searched, no reports
or observations of Huns could be secured, indicating little or no carryover populations and little or no increases in populations from previous years.
One brood of a hen with 2 chicks was reported by a rancher in the Jubb Creek
area in late August, 1970 in the vicinity of one of the 1970 releases. Reports
of Huns being seen distances of 4 and 5 miles from 2 of the 1970 release sites
were secured.

��-51-

HUNGARIAN

PARTRIDGE

ADAPTABILITY

Donald M. Hoffman

P. S. OBJECTIVE
To determine the ability of Hungarian partridge to establish
populations in mixed brushland-wheatland
habitat in selected
Colorado.

SEGMENT

OBJECTIVES

1.

Introduce

Hungarian

partridge.

2.

Determine

presence

and dispersal

METHODS
Field releases of pen-raised
April 24, 1970.

reproducing
sites in

of Huns and success

of previous

plant.

AND Y~TERIALS

stock were made in 3 areas in Moffat

County

on

Intensive field searches were made by Project W-37-R personnel during the
periods June 15-19; August 25-28; and September 14-17, 1970.
In addition,
persons living and working in or near the release sites were interviewed
and observations were recorded.

RESULTS

AND DISCUSSION

On April 24, 1970, 674 additional pen-raised Huns from the Fort Collins
Wildlife Research Station and Little Hills Experiment Station holding pens
were released in 3 areas of Moffat County.
Figure 1 shows a release made on
Johnson Gulch south of Craig and Figure 2 shows general habitat types of
this area.
Table 1 summarizes all field releases made in Colorado since
1964. A total of 1,923 Huns have been released with most (1,774) being
released in Moffat County, 135 in Larimer County, and 14 in Routt County.
Figure 3 shows the map location of the Johnson Gulch release with numbers
of birds released indicated, and Figure 4 shows the map locations of the
Price Creek and Jubb Creek releases, with the numbers of birds released
indicated.
Table 2 summarizes all Hungarian partridge reports and observations for
Moffat County for 1970. All reports and observations during 1970 were from
the vicinities of the April 24, 1970 releases except for a report of a
single Hun approximately 5 miles north of one of these 1970 release sites
and another approximately 4 miles north of another 1970 release site.
Only
10 Huns were observed by Project personnel, all of which were in 2 of 3
release areas for 1970. Photographs of several of these Huns were secured.

�I

VI
N
I

Fig. 1. Hungarian partridge release, John Gulch, south of Craig in Moffat County, April 24, 1970.
(D. Domenick, photo)

�I
I.J1

W
I

Fig. 2. Habitat in the vicinity of the Johnson Gulch release site, Moffat County. Vegetative types are
cultivated wheatlands bordered by big sagebrush, with a few scattered junipers on hillsides. (D. Domenick,
nhnrn)_

�-54Table 1.

Hungarian partridge field releases since 1964.
Number
Released

Source of
Stock

Release Area

Jan. &amp; Feb., 1964

135*

Oregon &amp; Idaho
(wild trapped)

Larimer County, Douglas
Lake Sec.l, T8N, R69W

Sub-total

135

March 28, 1964

40

Idaho
(wild trapped)

Moffat County, Round
Bottom Sec. 34, T6N, R92W

March 23, 1965

124

Oregon
(pen raised)

Moffat County, Round
Bottom Sec. 33, T6N, R92W

April 18, 1967

71

FCWRS
(pen raised)

Moffat County, Round
Bottom Sec. 21, T6N, R92W

April 30 and
May 1, 1968

236

FCWRS
(pen raised)

Moffat County, Round
Bottom Sec. 29, T6N, R92W

April 30, 1968

14

FCWRS
(pen raised)

Routt County, Northeast of
Hayden Sec. 35, T7N, R87W

April 19, 1969

389

FCWRS (374) &amp;
CSU (15)
(pen raised)

Moffat County, Isles
Mountain Sec. 24 &amp; 25, T5N,
R92W

April 19, 1969

240

FCWRS
(pen raised)

Moffat County, Isles Mtn.
Sec. 16, T5N, R92W

April 24, 1970

170

FCWRS
(pen raised)

Moffat County, Johnson Gulch
Sec. 15, T6N, R91W

April 24, 1970

234

FCWRS
(pen raised)

Moffat County, Jubb Creek
Sec. 35, T5N, R93W

April 24, 1970

270

FCWRS
(pen raised)

Moffat County, Price Creek
Sec. 30 &amp; 31, T4N, R95W

Sub-total

1,788

Date

Total

1,923

* An additional 22 birds (11 pairs) were taken to Rocky Ford Experimental
Bird Farm from this shipment. On October 6, 1965, 51 (pen raised) birds were
received from Oregon and were taken to the Fort Collins Wildlife Research
Station for experimental breeding purposes. On February 19, 1970, 13 1969hatched Huns from eggs secured from wild-trapped birds in England were received
from Winchester Farms, Alton, Illinois and were used for experimental breeding
purposes in 1970 and later years.

�R 92W

-55-

R 91W

"

\

It

;.

••

Fig. 3. Location and numbers of Hungarian partridge released, Johnson Gulch,
Moffat County, 1970.

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�-57-

Table 2. Summary of Hungarian partridge reports and observation, Moffat
County, 1970.
Date(s)

Number Huns

Area

Observed By

April, 1970

3 groups of
approx. 8 each

Jubb Creek, vicinity
of release sites

L. Kendall

1

Price Creek Road,
approx. 5 miles N. of
upper Price Creek
release site

May 21, 1970

B. L. M.
Personnel

Late May, 1970

2

Approx. 3 miles S. of
Jubb Creek release
sites

G. Hanson

June 16, 1970

3 (pair &amp;

Jubb Creek, vicinity
of release sites

B. Sigler

Price Creek, vicinity
of release sites

B. Sigler

single)
June 17, 1970

2 (pair)

June 18, 1970

5 (2 pair &amp; 1
single)

Jubb Creek, vicinity
of release sites

B. Sigler

May to early
August, 1970

Few

Price Creek, vicinity
of release sites

Stoddard

Late August, 1970

1 hen with 2
chicks

Jubb Creek, vicinity
of release sites

C. Hebblthwaite

Late August,
1970

Few

Yampa River, approx. 4
miles N. of Upper Jubb
Creek release site

C. Hebblthwaite

May to late
August, 1970

Few

Vicinity of Johnson
Gulch release site

Barnes

�-58-

Although areas of releases made prior to 1970 in Moffat County were searched,
reports and observations for these areas were nil, indicating an absence of
carry-over populations and little if any increase in populations.
One Hungarian partridge brood of a hen with 2 very small checks was reported
by rancher C. Hebblthwaire in the vicinity of the Jubb Creek release site
in late August, 1970.
Period of Release
Specific experiments have not been run to determine the best time for field
releases of Hungarian partridge in Colorado.
Traditionally,
the early
spring period has been selected because ~f the advantages of ample food
conditions with growing green grass and forbs and most of the severe winter
weather periods are past.
A field release time during the last half of April is a compromise between
the near approach of the egg laying season (early May under penned conditions)
and late snows which may occur as late as early May in areas selected for
Hungarian partridge releases.
Birds held in the large holding pens begin
to pair as early as February each year, and most are believed to be paired
by release time. By releasing all birds from a single holding pen at one
site, they have the opportunity to select the same mate upon release.
Handling of the birds is known to cause stress as measured in weight loss
and transporting the birds undoubtedly adds to this stress.
This may delay
the nesting activities of the birds somewhat; however, there should still
be ample time for the birds to establish territories and nest following
release.
Upon release from the crates, most of the Huns stay in a rather loose
group and appear to start pairing almost instantaneously.
Whether they pair
with the same birds as in the large holding pens is not known.
Birds observed
later at the release site or at a distance from the release site are usually
observed in pairs, so there appears to be no problem in the birds finding
mates following release.

Prepared

by

iJ~?r7~~

Donald M. Hoffman
Wildlife Researcher

~

�-59April, 1971

JOB PROGRESS REPORT

State of

...;C:..;O;;,.:L::;:O;.:.RAD=.;:.O
_
Game Bird Survey

W-37-R-24

Project No.

10

Job Title

Experimental Breeding of Hungarian Partridge

Period Covered:
Personnel:

Job No.

2

Work Plan No.

April 1, 1970 to March 31, 1971

Lawrence A. Webster, Robert L. Schmidt: John F. Corey, Donald M.
Hoffman and Dr. Harry D. Muller.

ABSTRACT
A test in which 12 pairs of qungarian partridge breeders fed a standard 22 percent
protein level game bird breeder feed compared with an equal number fed a similar
feed but containing 16 percent protein, indicated the 16 percent protein feed was
significantly better at the 10 percent level in numbers of fertile eggs laid (293
compared with 158). Egg fertility was also higher with the 16 percent protein
feed (86.94 percent compared with 73.49) resulting in higher numbers of chicks
hatched (234 compared with 130). Both levels of protein tested (22 percent and
16 percent) showed excellent results in the 7-day survival tests.
In a test where 6 pairs of early mating pairs were compared with an equal number
of late mated pairs, the late mated pairs produced more fertile eggs (129 compared
with 55) and a slightly higher percent hatch of fertile eggs was found with the
late mated pairs (80.62 percent compared with 78.18).
In another test which compared fertile eggs produced by 3 pairs rece~v~ng 2 extra
hours of artificial light each A.M., 3 pairs receiving 2 extra hours of artificial
light each P.M., and 3 pairs receiving no artificial light, it was found the
artificial lighting of the pens resulted in bringing breeder pairs into production
earlier than those which received no artificial light. However, total egg production and total fertile egg production were not increased with the use of artificial lights.
Production cost records were maintained in 1970. These indicated it costs a
minimum of $9.45 per Hun produced for field release using present methods, facilities, and equipment, including labor costs. If labor costs are excluded, a cost
of $2.20 per bird was calculated.

�-60-

RECOMMENDATIONS
One year's data have been collected in comparing 22 percent protein level
game bird breeder feed with 16 percent. Although results showed that the
16 percent protein feed was better for Hungarian partridge in 1970,
comparisons should be made for at least one more year. Extra lighting
tests started in 1970 should also be continued for at least one more year.
Progeny from highest producing pairs in 1970, Winchester Farms stock,
and the highest producing 1969-hatched adult breeders have been marked and
it is recommended that selection tests be started in 1971.
Numbers of Huns placed in rearing and conditioning pens at the Fort Collins
Wildlife Research Station and at the Rocky Ford Game Farm were recorded in
Segment 24. It is recommended that an accurate count of birds removed by
pen be made and a comparison of overwinter survival by various densities be
made in Segment 25.

�-61-

EXPERIMENTAL BREEDING OF HUNGARIAN PARTRIDGE
Donald M. Hoffman
Even though 36.90 percent fewer eggs were laid by the 52 pairs of Hungarian
partridge breeders in 1970, compared with an equal number of breeders in
1969 (1,147 eggs compared with 1,818), several refinements were made in
incubation procedures and the rearing of young. The 655 pen-raised Huns
released in the field in 1971, plus 20 extra birds held for breeders,
compares favorably with the 674 released in the field in 1970. A total of
614 were released in 1969 and 250 in 1968.
Refinements in procedures included (1) replacing the electric heat lamp
brooders with modified electric Jamesway heatring brooders, (2) eggs were
incubated at a humidity of 85.50 F wet bulb temperature in 1970 in place of
84.50 F used previously, (3) eggs were hatched at a temperature of 99.250 F
in 1970 in place of 99.500 F used previously, and (4) chicks were fed 27%
protein turkey starter crumbles in 1970 for the first seven days, 30% turkey
starter crumbles for the following two to four weeks, and 26% turkey grower
crumbles from the fourth week, with improved results. In 1969, 28% protein
game bird starter mash was fed for the first four weeks, followed by 26%
game bird grower crumbles and pellets.
P. S. OBJECTIVE
To develop game farm production techniques for Hungarian partridge.
SEGMENT OBJECTIVES
1.

To measure the effects of 2 levels of protein in the breeder feed
(16% versus 22%) on :
(a) Egg production
(b) Egg fertility
(c) Egg hatchability
(d) Chick survival for the first 7 days.

2.

To compare the numbers of fertile eggs produced by early mating pairs
with those mating late in the breeding season.

3.

To compare the numbers of fertile eggs produced by breeders subjected
to two extra hours of light each 24 hour period during (1) the early
morning and (2) the evening, with breeders receiving no artificial light.

4.

To determine production costs.

�-62-

METHODS

AND MATERIALS

The detailed arrangement of the Hungarian partridge breeding pens for 1969
is shown in Figure 1. Mated pairs for all except the late mated series
were selected and caught in the large Hungarian partridge holding pens
(Fig. 2). Table 1 lists the pen schedule for the various tests run in 1970.
One mated pair of Hungarian partridge was used in each pen.
Pens 1 through
36 are located in a converted nursery shadehouse with ground floors, and pens
37 through 51 are a separate series of wire floored experimental game bird
breeding pens.
Pens A-I, A-2, and A-3 are alleyway pens in the ground
floored series.
Protein Levels

in Feed

Pens 1-6 and 19-36 (24 pens) were used to test the effect of 2 levels of
protein in the feed. Birds in even numbered pens were fed a standard game
breeders feed containing 22 percent protein and those in odd numbered pens
were fed a similar feed, but containing 16 percent protein.
Eggs were
marked and placed in separate incubator drawers to facilitate determination
of (1) fertile eggs produced; (2) number of fertile eggs that hatched, and
(3) numbers of surviving chicks.
Chicks were held separately in the brooder
until they were 7 days of age to detect differences in survival.
Period of Mating
Pens 7-18 (12 pens) were used to compare numbers of fertile eggs produced
by earliest mating pairs with those mated late in the breeding season.
Even numbered pens were used for early mated pairs and odd numbered pens
for late mated pairs.
Six pairs used for earliest mating pairs were
selected as soon as the birds were observed to start pairing in the large
runs (February 6, 1970) and the 6 pairs used for latest mating pairs were
selected from 10 unpaired hens and 10 unpaired cocks held in separate pens
from February 6, 1970 to March 23, 1970. These began to pair almost
immediately when recombined on March 23, 1970 and pairs were selected later
on the same date.
Extra Lighting

Tests

The wire-floored series of pens are well suited for extra lighting tests
and the necessary wiring and automatic timing devices were installed late
in Segment 23. U. S. Weather Bureau sunrise-sunset tables were used to
determine times when lights should go on and off.
Pens 46-48 (3 pens) were lighted with 2 extra hours in the early morning
and pens 49-51 (3 pens) were lighted with 2 extra hours during the late
evening.
Pens 40-42 (3 pens) were used for controls and were treated
identically except no artificial lights were used.
Originally, pens 37-39
were also listed for controls, but the pair in pen number 39 escaped on
June 2, 1970, so this series of 3 pens was used only in comparing earliness
of egg production.
Records were maintained of dates of first eggs, numbers
of eggs laid, numbers of fertile eggs laid, and hatchability of eggs produced.

�-63-

N

IJ ~
-

CD

75'

6'4"

,

If]

'-

If)
Jf\

Ground-f loored Series

3738 394041

42 43 4445 46 47 4E

~ , v_ '-- 'v _ '---'v _
.'

\ V _'---,

v_ '-- v _
\

..
Wire - floored
CD

v

I-

N

A-2 ~

",'

b

-

0

N

N
(\J

01

N-.!.

-,

/

/

49~

.r:.

N

0

~1
A-3

SCALE 8

L

I

/

en

CD

I

I

0' 5' 10'

I

25'

~
L
/0)

Fig. 1.

U)

Detailed Arrangement

=,

an

5_'~

(11

N,

-

Ser

,

Partridge

/N

J'--'6---1

48' ----·-----······1

I:!

CD

l'" '

Breeding

of

Pins.

Hungarian

i,.

_

....l

�-64Covered
Brooder

Shelter

1

Houses

/~
l.--_-1- __

I'-__

I

..l..._---' _-+----L----.J4't-8'~

f

J
-0
t\J

4'

t

N

-in

o

C\I

C\I
C\I

•.4''i
-L__
~ 15.5
,"'15.5-1
---35
---

~_~

~

---l

~------------I05'-----------~
Fig. 2.
Hungarian

Partridge

Holdin q Pens

�-65Table l.

Pen schedule for Hungarian partridge, 1970.

Protein Level
Pen
Nwnber' 22%
16%

T
T

3
4

8x8

T

1
2

Pen Size ~Ft·2

T
T

5

8x16

x

x

x

x

x

x

x

x
x

x

7

x

x

8

x

x

9

x

x

10

x

x

11

x

x

12

x

x

13

x

x

14

x

x

15

x

x

16

x

x

17

x

x

18

x

x

20

T
T

21
22

T
T

23
24

T

25
26
27

T

T
T

Artificial Light
None
AM
PM

x

T

T

Mated
Late

x

6

19

6x33

Pairs
Early

T

x

T
T

x
x

T
T

x
x

T
T

x
x

T
T

x
x

T
T
T

x

x
x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

-----------------------------------------------------------------------------

�Table 1.

-66Pen schedule for Hungarian partridge, 1970.

Pen
Number

Pen Size ~Ft.2
8x8
8x16
6x33

28

Protein Level
22%
16%
T

29
30

T

T

31
32

T
T

33
34

T
T

35

T

Pairs
Early

(continued) .

Mated
Late

Artificial Light
PM
None

AM

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

36

T

37

x

x

x

38

x

x

x

39

x

x

x

40

x

x

T

41

x

x

T

42

x

x

T

43*

x

x

x

44"k

X

X

x

45"~

x

x

x

46

x

x

T

47

x

x

T

48

x

x

T

49

x

x

T

50

x

x

T

51

x

x

T

A-1"~

x

A-3"/( x
No. Pens
on Test 12
12
0
0
0
6
Key-T - Pen on Test
x - Pen not on Test
,.~Ind icates pens with Winchester stock birds.

x
x
6

3

3

3

�-67-

Production

Costs

Costs were estimated, based upon necessary labor, equipment, and facilities
required to produce a number of Hungarian partridge similar to anticipated
production at the Fort Collins Wildlife Research Station for 1971, using
methods now in use. These costs do not reflect the costs related to research
studies which have. been of considerable value in improving methods and
techniques for producing numbers large enough for adequate field releases.
Winchester

Farms Stock

Although not written as a separate test in 1970, 13 birds (5 hens and 8 cocks)
which were survivors from the 18 Hungarian partridge received from Winchester
Farms in Illinois in February, 1970, afforded the opportunity to test fertile
egg production and hatchability in 1970. These were all July, 1969-hatched
from eggs received at the McGraw Wildlife Preserve from wild Huns in England.
This new blood line will be introduced into our resident flock st.arting in
1971.
The Winchester Farms Huns were held in a separate pen and allowed to pair
naturally.
Pairs were placed in pens 43-45, A-I, and A-3. Records were
maintained of numbers of eggs laid, numbers of fertile eggs, and hatchability
of eggs produced.
Pens 43-45 were compared with pens 40-42 (controls).
Progeny from the first 2 hatches of 1970 were Wing-tagged on the left wing
in preparation for crossing with resident stock.
Because of limited numbers
of these progeny,S
adult cocks were also held over for use in 1971 crosses.

RESULTS

AND DISCUSSION

Protein

Levels

in Feed

Table 2 lists a comparison of numbers of eggs produced and percent hatch of
eggs from breeder pairs in 12 pens using a standard 22 percent protein
level feed with 12 breeder pairs using a similar feed but containing 16
percent protein.
A total of 122 more eggs (337 compared with 215) was laid
by the breeders using the 16 percent protein feed in comparison with the 22
percent protein feed. A total of 135 more fertile eggs (293 compared with
158) was laid by the breeders using the 16 percent protein feed in comparison
with the 22 percent protein feed. This difference is significant at the 10
percent significance level. In addition, 86.94 percent of the eggs laid by
breeders on 16 percent protein feed were fertile compared with 73.49 percent
for the 22 percent protein feed. A total of 234 chicks was hatched from
breeders on the 16 percent protein feed compared with only 130 for the 22
percent protein feed.
Excellent survival for the first 7 days was found for both the 16 and 22
percent protein level feeds (98.26 percent survival for the 16 percent feed
and 97.60 percent survival for the 22 percent feed) as listed in Table 3.
A very high survival rate also continued following the 7 day test, reflecting
the strong, healthy condition of the chicks from both levels of protein in
the feed and improvements in brooding methods used in 1970. The electric
heat lamp brooders were replaced by modified electric Jamesway heatring
brooders in 1970 for better results in both survival and condition of the
young birds.

�Table 2. Numbers of eggs produced and hatchability of eggs laid by Hungarian partridge on two levels of protein in
the feed, 1970.

Protein
Level

Number
Pens

Number
Eggs
Produced

Number
Infertile
Eggs

Number
Fertile
Eggs

Percent
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germ

Number
Culls

16%

12

337

34

293

86.94

234

69.44

79.86

9

50

10

22%

12

215

52

158

73.49

l30

60.47

82.28

2

26

5
I

~
co
I

Table 3.

Survival of Hungarian partridge on two levels of protein in the feed, 1970.
Number Chicks Taken
to Brooder House or
Battery Brooder

Protein
Level

Number
Pens

16%

12

230

22%

40

501

Number Chicks
Lost First Seven
Days

Number Chicks
Survived
Seven Days

Percent Survival
First Seven Days

4

226

98.26

12

489

97.60

�-69-

Based upon this one year's results, the 16 percent protein level appeared to
be considerably better than the 22 percent protein level, for Hungarian
partridge.
It is planned to run similar tests in 1971, but with 6 pens on
16 percent protein level feed and 6 pens on 22 percent protein level feed.

Period of Mating
Table 4 lists a comparison of numbers of fertile eggs produced and percent
hatch of fertile eggs from 6 pens with early mating pairs with 6 pens with
late mating pairs, all using 22 percent protein level feed. Considerably
more fertile eggs were laid by the late mating pairs (129 compared with 55)
and a slightly higher percent hatch of fertile eggs was found with the late
mating pairs (80.62 percent compared with 78.18).
This resulted in 104
chicks hatched from the late mating pairs compared with only 43 for the early
mating pairs.
Three of the 6 pens of early mating Huns laid no eggs compared
with only 1 of the 6 pens of late mating Huns which laid no eggs.
It therefore appears that variability of individual birds played a large part in the
results obtained.
No clear advantage was found in selecting earliest mating
pairs and, in fact, this proved to be disadvantageous
in 1970.

Extra Lighting

Tests

A comparison of numbers of eggs laid, numbers of fertile eggs, hatchability
of eggs, and numbers of chicks hatched by 3 pairs of Hungarian partridge
under artificial lighting for 2 extra hours in the A.M., 3 pairs under 2
extra hours during the P.M., and 3 pairs receiving no artificial lighting
(controls) is listed in Table 5. All pens received 22 percent protein level
feed and all were treated as alike as possible in other respects.
Originally,
the experiment was set up using 6 controls and 6 treated pens but a pair of
Huns in pen number 39 escaped on June 2, 1970 so this series of 3 pens
(Numbers 37-39) was not used except to compare earliness of egg production.
More eggs were produced by 3 pairs receiving no extra lighting or controls
(102) compared with those receiving 2 extra hours in the A. M. (83) and
those receiving 2 extra hours in the P. M. (71). In comparing numbers of
fertile eggs, both the pairs in 3 pens receiving extra morning light and
the 3 pens of controls laid 77 fertile eggs each, while the 3 pens receiving
extra evening light laid only 48 fertile eggs.
Considering hatchability of
fertile eggs, the controls ranked highest with 90.91 percent, followed by
those receiving 2 extra hours during the P. M. (87.50 percent) and those
receiving 2 extra hours during the A. M. (70.13 percent).
Numbers of chicks
hatched were 70 for the controls, 54 for the pens receiving extra morning
light, and 42 for the pens receiving extra evening light. Of the 9 pens
used in this test, the minimum number of eggs laid was 14 from pen Number 46,
(lighted for 2 extra hours in the A. M.), and the maximum number was 51 from
pen Number 42 (a control with no extra lighting).

�Table 4.

A comparison of eggs produced by earliest mated pairs of Hungarian partridge with those mated late, 1970!/.

Treatment

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Early Mated

6

64

Late Mated

6

141

Number
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

6

55

43

67.19

78.18

1

11

3

9

129

104

73.76

80.62

0

25

3

1/ A standard 22% protein game bird breeder feed was used in all pens.

Table 5. A comparison of eggs produced by Hungarian partridge pairs receiving two extra hours of light with those
receiving no extra light. !/
I
....•

0
I

Treatment

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Number
Fertile
Eggs

Two Extra
Hours A. M.

3

83

3

Two Extra
Hours P. M.

3

71

Controls

3

102

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

77

54

65.06

70.l3

5

18

3

23

48

42

59.15

87.50

1

5

0

20

77

70

68.63

90.91

1

6

5

1/ A standard 22% protein game bird breeder feed was used in all pens.

�Table 6.

I.

An estimate of minimum costs involved in producing Hungarian partridge. 1/

Capitol Investment
Breeding Pens - Set of 50 size 8' x 16' each
Brooder Houses - Set of 3 with shelters size 26' x 10'
Holding Pens - Set of 4 size 36' x 222'
Incubator
Brooders, feeders, and waterers
Small tractor with mower
Work facilities with egg storage room

$

7,500.00
3,000.00
10,000.00
2,000.00
500.00
1,000.00
2,500.00
$26,500.00

Sub-total
Forty year life expectancy - Cost per year
II.

Feed and Utilities
Feed for breeders and young per year
Utilities (heat, light, and water) per year

$

600.00
500.00
1,100.00

Sub-total
III.

662.50

$

Labor
Conservation Aide level, 2nd Step, for 5 months;
1 part time for 7 months per year
Retirement, hospitalization, etc. (7%) per year

5,400.00
400.00

Sub-total

5,800.00

TOTAL

$7,562.50

Estimate can produce 800 Huns per year with these facilities and labor.
Estimate of cost per Hun produced

=

$7.562.50
800

=

=

$1,762.50

Estimate of cost per Hun produced, labor excluded

1/ Costs do ~

800

=

$

9.45

s

2.20

include the following:
a. Land investment since value can probably be returned at end of use period plus interest, based
upon present trends.
b. Cost of breeding stock
c. Interest required on capitol investment.
d. Expenses incurred due to unforeseen losses.

I

"-J

I-'
I

�Table 7.

Source

A comparison of eggs produced by Winchester Farms stock Hungarian partridge with resident stock, 1970.

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Number
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

I

-..,J
N
I

Winchester
Farm, Illinois

3

83

26

55

46

55.42

83.67

1

8

2

FCWRS

3

102

20

77

70

68.63

90.91

1

6

5

�-73-

Even though the use of artificial lighting did not increase the total egg
production in 1970, it was found that the lights did bring the breeding
pairs into production earlier than the breeding pairs without artificial
lighting.
On May 15, 1970, all 6 of the lighted pens were in production and
had produced 22 eggs, compared with only 4 of the 6 original controls being
in production and having produced only 13 eggs.
The bringing of breeders
into production early may be of value, particularly when the next year's
breeders are selected from the first hatch of the previous year as is the
case at the Fort Collins Wildlife Research Station.
Because of the small samples possible in 1970, it is planned to rerun this
test for at least one more year, using the same number of pens in each sample.

Production

Costs

Table 6 lists estunates of minimum costs involved in producing Hungarian
partridge based upon estimates of capitol investment, feed and utilities,
and labor required to produce approximately 800 Huns, using procedures now
in use at the Fort Collins Wildlife Research Station.
A cost of $9.45 for
each Hun released in the field was calculated from this cost analysis,
including labor costs.
Most cost estimates on game bird propagation do not
include labor costs, which are the largest single annual cost item.
If
these labor costs are excluded, a cost of $2.20 per bird released in the
field was calculated.
Methods to reduce labor costs of propagating Huns
through rearing of young under semi-natural conditions are being tested.

Winchester

Farms Stock

Table 7 compares numbers of eggs laid, numbers of fertile eggs, hatchability
of eggs, and numbers of chicks hatched from 3 pens of pure Winchester Farms
stock with 3 pens of resident Huns. All birds received 22 percent protein
level feed and were treated as much alike as possible.
The 3 pens of Winchester Farms stock produced fewer numbers of eggs (83
compared with 102), fewer numbers of fertile eggs (55 compared with 77), and
fewer numbers of chicks hatched (46 compared with 70) in comparison with 3
pens of controls (resident stock).
In addition, hatchability of fertile
eggs was lower for the Winchester Farms stock (83.64 percent compared to
90.91) in comparison with the resident stock. Since the Winchester Farms
stock is only one generation removed from the Wild, these results were not
unexpected.
The Winchester Farms stock should be of considerable value in
introducing new bloodlines and may help prevent excessive future inbreeding.

Prepared

by

£~~~

Wildlife

Researcher

��-75April, 1971

JOB PROGRESS REPORT

State of

...;C::;.:O::,:L:;;O;,,;.RAD=;,;::O
_

Project No.

W-37-R-24

Work Plan No.

Game Bird Survey

12
Job No.
--=1.,:.0-,_
Relationships of the Productivity and Distribution of
~W~i~l~d~Tu~r~k~e,y~s~o~n~t~h~e~U~n~c~o~m~p~a=h~g~r=e~P~l~a~t~e~a~u~t~o~t~h~e~
_

Job Title
Period Covered:

April 1, 1970 to March 31, 1971

P.S. OBJECTIVES
1.

To determine the size, distribution, and productivity of the wild turkey
population on the Uncompahgre Plateau.
20 Record movements of wild turkeys and determine how environmental factors
affect these movementso
3. Determine the sex and age structure of the turkey kill on the Uncompahgre
Plateau to provide estimates of total harvest and size of the turkey
population.
40 Determine the vegetative types present on the wild turkey summer and winter
range and the species composition and abundance of food producing species
within each type. Also determine the elevation, describe topography, and
record certain climatic factors on part of the better turkey summer and
winter range.
SEGMENT OBJECTIVE
To compile, analyze, summarize, and publish information.
Progress
Field work was completed on October 20, 1967. All data have been summarized,
and most data have been analyzed. A manuscript containing these data and results
from Work Plan 12, Job 14 is being edited and revised.
Future Plans
The manuscript should be completed in Segment 25.
in the form of a technical bulletin.

c" "

(1

rt1, ~_

Prepared by: &lt;~~
oJ II~
Gary T. Myers
Wildlife Researcher

.'

Publication will probably be

��-77April, 1971

JOB PROGRESS REPORT

State of

C.=..O.=..L;;;,O;;;.;RA=D;;..;O~__:..
_

Project No.

W-37-R-24

Game Bird Survey

12

Work Plan No.
Job Title

Job No.

14

Use of Food Plots to Concentrate Wild Turkeys

Period Covered:

April 1, 1970 to March 31, 1971

P.S. OBJECTIVES
Determine the effects of small food plots on fall concentrations and harvest
of wild turkeys.
SEGMENT OBJECTIVE
To compile, analyze, summarize, and publish information.
Progress
Field work was completed December 2, 1968. Data were analyzed, written up,
and incorporated in a manuscript containing information gathered under W-37-R
Work Plan 12, Job 10. This manuscript is being edited and revised.
Future Plans
The manuscript should be completed in Segment 25.
be in the form of a technical bulletin.

Prepared by:

&lt;.p2"'-~~)
/J",~-f...4..
Gary T .'-yers
Wildlife Researcher

Publication will probably

��-79-

April, 1971

JOB PROGRESS REPORT

State of

~CO~L~O~RA~D~O~

Project No.

W-37-R-24

Work Plan No.
Job Title

Game Bird Survey
Job No.

2

Study of Mountain Quail Adaptability

Period Covered:
Personnel:

15

_

April 1, 1970 to March 31, 1971

Ronald B. Arant, Tom W. Barnes and Donald M. Hoffman.

ABSTRACT
Twenty-five surplus breeders, including eight 1969-hatched and seventeen
1967-hatched or older, were released on the South Fork of Mesa Creek in
Mesa County on June 23, 1970. The release site was approximately .6 mile
below the first crossing of the South Fork of Mesa Creek and approximately
5.7 miles above the last mountain quail sighting location. Birds released
included 14 hens and 11 cocks, which had produced no eggs during 1970.
Field trips to search release sites and surrounding areas on the Uncompahgre
Plateau were made in June and September, 1970. No mountain quail were seen,
no field sign found, and no new reports of sightings of the species were
secured.
Although mountain quail have been most difficult to propagate in captivity,
several problems have been solved. The species holds definite potential
for establishing a new game bird in the extensive areas of mountain shrub
and ponderosa pine zones on both the western and eastern slopes of Colorado.
If experimental propagation efforts are successful in 1971 and following
years at the Fort Collins Wildlife Research Station, it is recommended that
trial introductions be made in the Spanish Peaks area of Las Animas County
starting in 1972.

��-81-

STUDY OF MOUNTAIN

QUAIL ADAPTABILITY

Donald M. Hoffman

P. S. OBJECTIVE
To determine the ability of mountain quail to establish reproducing
populations in areas of mixed shrub and pine in selected sites in Colorado.

SEGMENT

OBJECTIVES

1.

Introduce mountain

quail.

2.

Determine

and dispersal

presence

METHODS

of mountain

quail.

AND MATERIALS

Twenty-five surplus mountain quail breeders from holding pens at the Fort
Collins Wildlife Research Station were banded with aluminum butt end leg
bands, transported to the South Fork of Mesa County in Mesa County in a
State pickup, and released on June 23, 1970.
Two separate field
of the Uncompahgre
living and working
any mountain quail

searches of the release sites and surrounding areas
Plateau were made in June and September, 1970. Individuals
in and near the areas were contacted and asked whether
had been observed.

RESULTS AND DISCUSSION
An observation of 20 mountain quail made by Conservation Aide Tom Barnes of
Nucla, Colorado on the South Fork of Mesa Creek reported in the Job Progress
Report for Segment 23 of this job for early 1970, was actually made in late
December, 1969. These were observed approximately one-half mile above the
old Bogus Mine in a pinon-cedar vegetative community.
Within their native
ranges, mountain quail that summer at higher altitudes frequently migrate
to lower levels in winter.
These were seen only one time, so they may have
moved into this lower area of South Fork of Mesa Creek from some unknown
higher area.
Twenty-five surplus mountain quail breeders, including eight 1969-hatched
and seventeen 1967-hatched or older birds, were released on the South Fork
of Mesa Creek in Mesa County on June 23, 1970. The release site was
approximately
.6 mile below the first creek crossing and approximately 5.7
miles above the last mountain quail sighting.
Birds released included 14
hens and 11 cocks, none of which produced eggs in 1970. Table 1 lists all
field releases of mountain quail and sources of stock being held at the
Fort Collins Wildlife Research Station for experimental propagation purposes.

�Table 1.

Summary of mountain quail field releases.
Number
Birds

Source of
Birds

Release
Area

Aug. 19 and 25, 1965

143*

California
(wild -trapped)

Mesa County, Indian Creek
Sec. 27, TS1N, R17W

Aug. 20, 1966

163

California
(wild-trapped)

Mesa County, Indian Creek
approx. 4 miles west of 1965
releases

Jan. 25, 1968

66

Oregon
(wild -trapped)

Mesa County, Indian Creek
approx. 1 mile east of west
Uncompahgre National Forest
boundary

Date

I

00

June 23, 1970

25

Total

397

FCWRS
(surplus breeders)

Mesa County, South Fork Mesa
Creek, Sec. 17, T49N, R16W

* Sixteen additional mountain quail from these shipments were taken to Fort Collins Wildlife Research
Station for experimental propagation studies. On January 18, 1970, 12 Wild-trapped mountain quail from
Oregon were added to the breeding stock at the Fort Collins Wildlife Research Station. These constitute
the source of our breeding stock.

N
I

�-83-

Field trips to search for mountain quail in the release areas and surrounding
areas on the Uncompahgre Plateau were made in June and September, 1970. No
mountain quail were seen, no field sign found, and no new reports of
sightings were secured from individuals contacted.

Prepared by

J/}~P2~
Donald M. Hoffman
Wildlife Researcher

��-85April, 1971

JOB PROGRESS REPORT

State of

C~O~L~·O~RAD~~O~_

Project No.

W-37-R-24

Work Plan No.
Job Title

Job No.

3

Experimental Breeding of Mountain Quail

Period Covered:
Personnel:

15

Game Bird Survey

April 1, 1970 to March 31, 1971

Lawrence A. Webster, Robert L. Schmidt, John F. Corey and Donald M.
Hoffman.

ABSTRACT

A new set of 10 mountain quail breeding pens was completed on April 13, 1970
and used for the 1970 season, but only one egg, which was infertile, was laid
by the entire group of 32 breeders of mixed sexes placed in the breeding pens
and no eggs were laid by 38 surplus birds of mixed sexes placed in holding
pens. The catching and sorting of the mountain quail breeders on April 14,
1970 apparently stressed these tempermental birds and resulted in almost no
egg production in 1970. This lateness in moving the breeders was due to a
delay in completing the new set of breeding pens.

��-87-

EXPERrnENTAL BREEDING OF MOUNTAIN QUAIL
Donald M. Hoffman

P. S. OBJECTIVE
To develop game fann production techniques for mountain quail.
SEGMENT OBJECTIVE
To measure the relationship of pair mating, selected group mating and flock
mating to fertile egg production.
METHODS AND MATERIALS
A new set of 10 mountain quail breeding pens was completed on April 13,
1970 (Fig. 1). It was planned to have these completed by mid-March but
inclement weather plus a manpower shortage delayed the completion.
Pairs of mountain quail were placed in pen numbers 1, 5, 7, and 9; trios
(2 hens and 1 cock) in pen numbers 2, 4, 6, and 10; and a community pen of
8 hens and 4 cocks was placed in the combined pen numbers 3, 8, and alleyway, on April 14, 1970. All birds were fed a standard 22 percent protein
level game bird breeder feed and all were 1969-hatched birds. Numbers of
fertile eggs produced were to be recorded by individual pens.
Ten wild-trapped mountain quail of mixed sexes received from Oregon in
Segment 23 wereplaced in combined holding pen Numbers 1 and 2, eleven
1969-hatched birds including 2 hens and 9 cocks not needed to fill the
experimental breeding pens were left in combined holding pen numbers 3 and
4, and seventeen 1967-hatched or older breeders of mixed sexes were placed
in combined holding pen Numbers 5 and 6. Figure 2 shows the arrangement
of holding pens.
RESULTS AND DISCUSSION
Although procedures for the 1970 mountain quail tests were carried out as
planned, except for being approximately I month late in completing a set
of 10 experimental breeding pens, only 1 egg was laid. This was laid on
May 1, 1970 in the community pen (combined pen Numbers 3, 8, and alleyway)
and was infertile.
It was found in previous year's tests that mountain quail stress easily.
Catching and sorting the breeders at approximately the time when they should
have started to lay may have been the reason for almost no egg production
in 1970.

�-88-

N

~---------------30'--~
1F-----12'--~

~

~--

12' ---41

o

SCALE 8

.I
·0'

t

I.

5'

10'

Fig. 1.
Petallod

Arranoornont

of

Mou"toln

Quail BrlGdhiQ

Pont.

�-89-

I(

. 20'

~

lE-a'-)E-8'-)f~

f

10

N
~

5

I

3

1

-

\
4

6

2
.

Scole:

o
60' ----------------~
Fig. 2.
Mounialn

Quail

Holding

Pens.

I": 15 ft.

15

30

�-90-

No eggs were laid by ten wild-trapped mountain quail of mixed sexes received
from Oregon in Segment 23, eleven surplus 1969-hatched birds (2 hens and
9 cocks), or seventeen 1967-hatched or older breeders of mixed sexes held
in separate mountain quail holding pens in 1970. It was necessary to catch
and move these surplus breeders in order to select breeders used in the
experimental breeding pens and to utilize holding pens with the best cover.
After it became apparent that surplus breeders held in holding pens were not
producing eggs, 25 (seventeen 1967-hatched or older and eight 1969-hatched)
were banded and released on the west side of the Uncompahgre Plateau on
June 23, 1970. As of February 9, 1971, there were 26, 1969-hatched and
9 wild-trapped mountain quail from Oregon on hand for use as breeders in
the 1971 tests. A total of 7 birds, including 6 1969-hatched and one wildtrapped, died while in wintering holding pens, prior to February 9, 1971.

Prepared

by

!l~'M~~
onald M.offman
Wildlife Researcher

�-91April, 1971

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~----------

Project No.

W-37-R-24

Game Bird Survey

Work Plan No.

17

Job Title

Inventory of Selected Ptarmigan Populations

Period Covered:
Personnel:

Job No.

2

April 1, 1970 to October 15, 1971

Clait E. Braun and Terry A. May.

ABSTRACT
Investigations of white-tailed ptarmigan (Lagopus leucurus) populations in
Colorado initiated in 1965 were continued in 1970 with major emphasis upon
populations levels, and relationships of nesting success and production to
climatic conditions. Breeding density decreases ranged from .6 to 6.0 birds
per square mile on all study areas from levels,observed in 1969. The observed
decreases were primarily the result of poor nesting success and production in
1969 on all areas and excessive harvests in 1969 at Independence Pass and Mto
Evans. Observed densities in 1970 ranged from 5.2 at Mt. Evans to 24.8 in
RMNP. Weather during periods of egg deposition and incubation in 1970 was warm
and dry, and nesting success ranged from about 50 to 75 percent. Survival of
chicks to September 1 was similar to all previous years studied. Average brood
size to September 1 was 4.5 chicks. Present data continue to indicate that
nesting success and production are closely related to amount of precipitation
and number of frost-free nights in June.

��-93-

INVENTORY OF SELECTED PTARMIGAN POPULATIONS
C1ait E. Braun

Knowledge concerning population levels of white-tailed ptarmigan and factors
influencing observed densities is essential for proper management. Substantial data are available relating to this small grouse in Colorado (Braun
1971; Braun and Rogers 1971; and Braun and Schmidt 1971), but additional
information is necessary for management programs. It has been hypothesized
by many workers that all species of grouse (family Tetraonidae) are cyclic,
exhibiting either short-term (3-5 years) or long-term (10-12 years) fluctuations. Present data concerning grouse inhabiting northern areas support
this hypothesis, but few long-term studies have been conducted on species
of grouse at the southern edge of their range. It has not been demonstrated
that any species of grouse is cyclic in Colorado. Relationships between
weather conditions and population levels of grouse have been suggested by
Siivonen (1957), Neave and Wright (1969), and others, but have not been
statistically examined for white-tailed ptarmigan. Campbell (1968) and
Francis (1967; 1970) have demonstrated close relationships between certain
weather parameters and populations of scaled (Callipepla squamata) and
California quail (Lophortyx ca1ifornicus). This project was undertaken to
examine long term population trends and effects of climatic conditions on
white-tailed ptarmigan in Colorado. Following recommendations of Braun and
Rogers (1970) all studies were discontinued at Mesa Seco.
P. S. OBJECTIVE
To test the hypotheses that (1) populations of white-tailed ptarmigan in
Colorado are not cyclic and (2) production and fall population densities
of this species in Colorado are related to temperature and moisture conditions
during June of the same year.
SEGMENT OBJECTIVES
1.

To estimate breeding population levels of ptarmigan in five study areas
(Mt. Evans, Crown Point, Independence Pass, Mesa Seco, and Rocky Mountain
National Park).

2.

To estimate nesting success and production in the above areas.

3.

To obtain weather data from a representative alpine area for use in
determining correlation between spring weather conditions and fall
ptarmigan populations.

4.

To compile data and prepare progress report.

�-94-

METHODS

AND MATERIALS

Techniques used were essentially those developed under Work plan 17, Job 1
and reported in detail by Braun and Rogers (1971).
Ptarmigan were censused
by traversing study areas using tape-recorded calls to locate territorial
males, breeding pairs, and hens with chicks.
Efforts were made with telescoping noose poles to catch all ptarmigan not banded previously.
All birds
caught were banded with serially numbered gold anodized aluminum leg bands,
with all adults being additionally banded with individually numbered blue
plastiC bandettes.
Green bandettes with black numerals were used to replace
any bandettes from previous years that had become too worn for recognition
of individual birds.
Weather data were obtained from the Institute of Arctic and Alpine Research
(INSTAAR), University of Colorado, for their D-l, Niwot Ridge Site.
This
site is located at 12,300 feet between the Rocky Mountain National Park
(RMNP) and Mt. Evans study areas.
Data collected at this site and in RMNP
have been previously ahalyzed and compared (Braun 1969), and differences
were found to be insignificant.
Weather data at this site have been systematically collected using standard instruments since 1952 (Marr 1967a; Marr
et ale 1968a and b).

DESCRIPTION

OF STUDY AREAS

Areas intensively investigated, with one exception, are those studied since
1966 under Work Plan 17, Job 1, and described in detail by Braun and Rogers
(1971). As suggested by Braun and Rogers (1970) at the cessation of the
initial 5-year project on inventory of ptarmigan populations, all work was
terminated at the Mesa Seco site.
The four current study areas are presented
in Figures 1-5.

RESULTS AND DISCUSSION
Breeding

Densities

Censuses of breeding birds were initiated on May 2 with few males being
observed on known territories.
The first breeding pair was observed on
May 4 in RMNP but most pairing did not occur until mid-May.
Weather conditions during the 1970 breeding period were similar to those experienced in
1968 and were colder and drier than in 1969. Snow levels in early May were
slightly less than in previous years studied except 1966. Timing of breeding
activities was earlier than in 1969 but generally was similar to previous
years aild suffered no major disruptions froni adverse weather.
Densities
observed are presented in Table 1.
Breeding
observed

densities decreased on all areas studied
in earlier years (Table 2).

in 1970 from levels

�-95-

7

-

(:
---..,,,

//~- ..

/

Fig.

1.

\
Mt. Evans study area.

T EVANS

ARE

COLORA

0

Stydl1 Area ~()unC»OJrlf

�-96-

.'.:./

CROWN

POINT AREA

COLORADO
-

Study Area Boundary

/-.--~--.
"/0..

Fig. 2.

Crown Point study area.

"...
.".~/

�-97-

ROCKY MOUNTAIN

NATIONAL

Pj\RK

SUNDANCE UNIT
It!

J

-

COLORADO
Study Area Boundary
SCALE

1111MILES

bE=:r::::=::JE3E3:::=J:E3~'3/=2=EE3=:r::::=::JE3===::::j:

Fig. 3.

Rocky Mountain National Park, Sundance Unit study area.

~ILE

�.--------.

Fig. 4.

.,
.~
C

ra

,.

Rocky Mountain

National

-98-

U)

t-

Z
:,:)

«r ~

~
c(

0..

-' &lt;tZ

c(

••••
::l

:z

&gt;L..

c

0

"0

::)
0

en

l!

(It

&lt; )., '&gt;

~

tal

-

::!

:i

"0

&lt;II

u

w
..,j

..I

IU

~~

0

I

-' -

0

...
~
,..

Z
0

.,

~
c( 0 0
0
Z
...J ~
0:
Z
~ ..,J 0
::)
&lt;C

c(

en

0

0

I- ~
Z

:&gt;

::l 0::I

0
~

&gt;~

study area.

/0
1&gt;

Mountain

LL

~ 0:
0 ...J
0 ...J
0=

Park, Fall River-Toll

�-99-

INDEPENDENCE
N

ASS AREA

COLORADO

I T .•..•

Study Aroo Boundory
o••

0,,0

0.000

~==E:~~~1/~2E:~~e+3:3~1
o~
e-3 F3
E3
:+

Fig.

5.

Independence

Pass

s tudy area.

�-100-

Table 1. White-tailed ptarmigan breeding population densities, all areas,
1970.

Square
Miles

No. of
Breeding
Pairs

Unmated
Birds

Total
Breeding Season
Population

Birds per
Square Mile

Tombstone RidgeSundance Mountain

1.25

8

5 males

21

16.8

Toll Memorial

.19

4

3 males

11

57.9

Fall River Pass

.70

9

3 males

21

30.0

2.14

21

11 males

53

24.8

Crown Point

1.93

8

4 males

20

10.4

Mt. Evans

1.54

3

2 males

8

5.2

Independence Pass

1.12

4

4 males

12

10.7·

Study Area
RMNP

Total RMNP

Table 2.

White-tailed ptarmigan breeding population densities, 1966-70.

1966

Birds eer Sguare Mile
1967
1968
1969

1970

RMNP

29.0

25.2

29.4

30.8

24.8

Crown Point

14.5

21.2

18.1

14.5

10.4

Mt. Evans

7.8

7.1

7.1

5.8

5.2

Independence Pass

18.7

18.7

21.4

17.9

10.7

Study Area

Decreases in breeding population levels were greatest at Independence Pass
and lowest at Mt. Evans. Reasons for the observed decreases are related to
the generally poor nesting success and production of young in 1969 at all
aceas and the continued heavy exploitation of fall populations by hunters

�-101~

at Independence Pass and Mt. Evans in 1969 (Braun and Rogers 1971). The
decreases at RMNP and Crown Point can only be attributed to poor nesting
success and production as no hunting is allowed in RMNP and no birds were
reported harvested at Crown Point in 1969. All decreases were _the result of
poor recruitment of young birds into the breeding population in 1970.

Nesting Success and Production
Nesting activities were initiated on all areas in 1970 between June 10 and
June 20. Incubation of one nest under observation was initiated on June 22.
Weather conditions during June and early July 1970 were warm and dry and
were similar to those in 1966 and 1968 when nesting success and production
were good to excellent. Estimated peak of hatching in 1970 was between
July 15-20 with the one nest under observation hatching on July 15-16.
This nest in RMNP contained six eggs and all successfully hatched.
The primary molt of 15 successful hens was examined in order to calculate
hatching dates. The earliest calculated hatching date was July 12 and the
latest was July 28. Eleven of the 15 successful females hatched their
clutches between July 15-20 while only one hen was calculated to have
hatched her clutch after July 25.
In 1970, 32 different hens were observed during the brood period (July 12September 15) with 27 of these being successful. While successful females
are more easily seen than those without chicks, the number of unsuccessful
females observed was low in 1970. When the five unsuccessful hens seen in
1970 are compared with the 22 brood less females reported in 1969 (Braun and
Rogers 1971), it is apparent nesting success was substantially better in
1970 than in 1969. Estimated nesting success in 1970 was 75 percent in RMNP,
65 percent at Crown Point, and ~ 50 percent at Mt. Evans and Independence
Pass.
Initial size of broods in 1970 upon hatching was six chicks. Brood size to
August 1 averaged 5.3 chicks, while each brood averaged 4.5 chicks on
September 1 (Table 3). This was an increase of 1.6 chicks per brood over
the 2.9 chicks per successful hen recorded to September 1, 1969.
Fall Densities
Estimates of densities of white-tailed ptarmigan on September 1, while
useful in illustrating population gain through production, are difficult to
accurately derive due to a number of variables. Estimated fall densities
for each area studied are presented in Table 4. These densities were
calculated following the three basic assumptions discussed in detail by
Braun and Rogers (1971).
Data presented in Table 4 present a good picture of actual populations at
RMNP and Crown Point. This is not true of the data for Mt. Evans and
Independence Pass. All that can really be said of the fall densities at
these two locations is that they were low.

�-102-

Table 3.

Number of broods and average brood size, all areas, 1970.-1/
Number of
Broods Observed

Average Number of
Chicks per Brood

July 1-15

0

0.0

July 16-31

7

5.3

August 1-15

4

4.0

August 16-31

15

4.5

September 1-30

7

3.1

Total

33

Date

Average

4.3

1/ Only distinct broods are included.

Hunting further reduced fall densities at Crown Point as hunters in the fall
of 1970 harvested 15 of 29 banded birds in the population. At Independence
Pass, only 1 of 12 banded birds present in the study area was harvested,
while the Mt. Evans area was closed to all hunting of grouse in 1970.
Hunter check stations were not operated in 1970 so total harvest on the
three study areas open to hunting is not known.

Relationships of Weather:Nesting Success and Production
Weather data obtained from the 12,300 level of Niwot Ridge, through the
courtesy of INSTAAR for the 1966-70 period are presented in Table 5.
Detailed analysis and statistical correlations will be completed for the
final report in 1974-75. The close correlation between June weather
conditions and ptarmigan nesting success and production previously suggested
from work conducted under Work Plan 17, Job 1, continued in 1970. In
general, weather conditions were warm and dry in June and July, 1970, and
nesting success and production improved over 1969 when the early Summer
weather was colder and wetter.

�Table 4.

Estimated fall densities of white-tailed ptarmigan, 1970.

Area

Estimated
Percent
Nesting
Success

Average
Brood Size
on
September 1

Total
Production

Total
Breeding
Population

Total
Population
on
September 1

Birds
per
Square
Mile

Percent
Gain
I

I-'
0
w

RMNP

75

4.3

64

53

112

52.3

57.1

Crown Point

65

5.0

25

20

43

22.3

58.1

Mt. Evans

50

3.0

5

8

12

7.8

41. 7

Independence Pass

50

3.0

6

12

17

15.2

35.3

I

�Llb1e 5.

~.Jeatherdata, Niwot Ridge, 12,300 ft., May-July, 1966-1970.
TemEerature

Mean Daily
Max.
Min.

Number Days
Minimum
Temperature
Less Than 32° F

(Marr 1967b; Clark 1970).
PreciEitation
Number Days
Relative
Ppt/Month
Humidity
(Inches)
90 or Above

Ave. WindSpeed/
Month

Year

Month

Max.

Min.

1966

May

57

6

38.1

26.4

20

1.7

16 1.1

13.2

June

58

22

48.6

33.3

12

1.8

19

11.0

July

65

38

58.2

43.5

0

2.6

25

8.2

May

54

-7

32.5

21.5

25

3.1

25

15.1

June

58

24

44.6

30.4

18

3.7

29

12.3

1967

1968

1969

1970

(mph)

July

62

32

54.7

40.7

0

2.9

27

10.0

May

50

2

33.5

20.2

29

2.6

27 Jj

16.6

June

61

11

50.2

33.7

12

0.65

18

13.9

July

62

28

54.1

39.6

2

2.1

25

9.4

May

57

13

41.5

28.0

22

5.7

23

11.0

June

57

15

40.9

28.8

19

5.7

29

13.8

July

62

33

55.2

41.0

0

2.5

17

10.6

May

51

4

40.0

25.0

25

0.55

22

14.5

June

65

15

47.6

33.0

11

2.5

19

12.6

July

62

30

55.7

41.2

1

2.1

19

9.9

1/ Data not available for 4 days.
1/ Data not available for 1 day.

I
•....

0
.po.
I

�-105-

LITERATURE CITED

Braun, C. E. 1969. Population dynamics, habitat, and movements of
white-tailed ptarmigan in Colorado. Ph.D. Thesis. Colorado State
Univ., Fort Collins. 189 pp.
1971. Habitat requirements of Colorado white-tailed ptarmigan.
Proc. Western Assoc. State Game &amp; Fish Comma 51: In press.
______ , and G. E. Rogers. 1970. Inventory of Ptarmigan populations.
Colo. Div. Game, Fish and Parks, Game Res. Rep., Fed. Aid Proj.
W-37-R. April. pp. 181-183.
, and
1971. The white-tailed ptarmigan in Colorado.
------Div.
Game, Fish and Parks. Tech. Bull. 27: In press.

Colo.

------ , and R. K. Schmidt, Jr.

1971. Effects of snow and wind on wintering populations of white-tailed ptarmigan in Colorado. pp. 238-250.
In Haugen, A. O. (Ed.). Proc. Snow and Ice Symposium, Iowa State Univ.,
Ames. 280 pp.

Campbell, H.
Mexico.

1968. Seasonal precipitation and scaled quail in eastern New
J. Wildl. Mgmt. 32(3):641-644.

Clark, J. M. 1970. Unpublished data.
Colo., Boulder.

Inst. Arctic and Alpine Res., Univ.

Francis, W. J. 1967. Prediction of California quail populations from
weather data. Condor 69(4):405-410.
1970. The influence of weather on population fluctuations in
California quail. J. Wildl. Mgmt. 34(2):249-266.
Marr, J. W. 1967a. Data on mountain environments. I. Front Range, Colorado,
sixteen sites, 1952-53. Univ. Colo. Studies, Sere BioI. No. 27. 110 pp.
1967b. Unpublished data.
Colo., Boulder.

Inst. Arctic and Alpine Res., Univ.

, J. M. Clark, W. S. Osburn, and M. W. Paddock. 1968. Data on
------mountain
environments. III. Front Range, Colorado, four climax
regions, 1959-1964.

Univ. Colo. Studies, Sere BioI. No. 29.

179 pp.

______ , A. W. Johnson, W. S. Osburn, and O. A. Knorr. 1968. Data on mountain
environments. II. Front Range, Colorado, four climax regions, 19531958. Univ. Colo. Studies, Sere BioI. No. 28. 169 pp.

�-106-

Neave, D. J., and B. S. Wright. 1969. The effects of weather and DDT
spraying on a ruffed grouse population. J. Wild1. Mgmt. 33(4):
1015-1020.
Siivonen, L. 1957. The problem of the short-term fluctuations in
numbers of tetraonids in Europe. Finnish Papers Game Res. 19:
44 pp.

Prepared by

_~tU~!:::..::-_,_y=--.:....

--.:..)~~o&lt;:..::.-

Clait E. Braun
Asst. Wildlife Researcher

_

�-107April, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RAD~~O~ _

Project No.

W-37-R-24

Work Plan No.
Job Title

Job No.

1

Experimental Breeding of Tinamou

Period Covered:
Personnel:

19

Game Bird Survey

April 1, 1970 to March 31, 1971

Lawrence A. Webster, Robert L. Schmidt, John F. Corey and Donald M.
Hoffman.

ABSTRACT
A total of 478 eggs was laid by the 36 pale spotted tinamou (24 hens and 12 cocks)
on hand in May, 1970. Of these 50 (10.5 percent) were culls, 132 (27.6 percent)
were infertile, and 296 (61.9 percent) were fertile.
A normal incubation relative humidity of 55-56 percent (84~-85~Owet bulb) was
considerably better than a test incubation relative humidity of 50 percent
(82-3/40wet bulb) both in the hatching of total eggs set and in the hatching of
fertile eggs. In addition, fewer dead germs and pipped eggs which did not hatch
occurred with the normal incubation relative humidity of 55-56 percent in comparison with a test incubation relative humidity of 50 percent.
Higher numbers of total eggs were laid by breeders held in 4 separate pens with
selected groups of 2 hens and 1 cock each (248 eggs) compared with a community
of 8 hens and 4 cocks (62 eggs). In addition, 190 fertile eggs were laid by the
4 selected group pens of breeders compared with only 32 for the community pen.
More fertjle eggs were produced by breeders fed a standard 22 percent protein
level feed (190) compared with a similar feed containing 16 percent protein
level feed (74). A higher percent of the fertile eggs were also hatched from
the birds fed the 22 percent protein level feed (70.00 percent) compared with
the 16 percent protein level feed (51.35 percent).
This study was scheduled to run through 1972-73 but poor overwinter survival
under penned conditions has prevented the releasing of suitable numbers in the
field. A final field release of all remaining stock (82 birds) was made in
Baca County, south of Vilas, on April 30, 1971.

��-109-

EXPERIMENTAL BREEDING OF TINAMOU
Donald M. Hoffman
Good progress was attained in refining techniques for rearing pale
spotted tinamou in captivity at the Fort Collins Wildlife Research
Station in 1970, but overwinter losses were again heavy. From the
36 breeders (24 hens and 12 cocks) held for 1970 tests~ there were
187 chicks hatched but only 62 survived through the fall and winter
periods. A final field release consisting of 59 1970-hatched tinamou
plus 23 surviving 1969-hatched adults was made in Baca County, south
of Vilas, on April 30, 1971.
Although a few tinamou (exact numbers unknown) are known to have escaped
from the holding pens at Rocky Ford, it appears that overwinter losses
approximated 50 percent9 with heaviest losses occurring during and
immediately following the heavy snow and severe cold temperatures in
early January, 1971. In addition, many of the surviving tinamou showed
the effects of having toes frozen during the winter period.
P. S. OBJECTIVE
To develop game farm production techniques for pale spotted tinamou.
SEGMENT OBJECTIVES
1.

To compare two levels of relative humidity (50% versus 55%) during
incubation on egg hatchability.

2.

To measure the relationship of flock mating and selected group mating
to total egg production and fertile egg production.

3.

To measure the effects of two protein levels in the feed (22% versus
16%) on fertile egg production and egg hatchability.

METHODS AND MATERIALS
The detailed arrangement of tinamou breeding pens is shown in Figure 1.
The pen schedule for tinamou for 1970 is shown in Table 1.
Selected groups of 2 hens and 1 cock were placed in pens 1, 2, 4, 5, 6, 7,
9, and 10 and a community flock of 8 hens and 4 cocks was placed in the combined pen 3,8, and alleyway on April 10, 1970. Thus, a total of 36 birds
was used in breeding tests in 1970. All breeders were dusted with louse
powder and primaries on both wings were clipped prior to releasing in the
breeding pens. Two birds died while in the breeding pens and 34 were recovered on September 28, 1970. Numbers of eggs produced and numbers of
fert:ileeggs produced were compared for the selected groups in pens 2, 4, 6,
and 10 and the community group in pens 3,8, and alleyway combined, on all
22 percent protein level feed.

�-110-

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~------------30'
I'
i"

,

-en

~

12

~

12

-"-

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0

sr

to

7

.'"
s»

1

to

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r0

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.

(j)

,.,
SCALE

Fig. 1.

Detailed

Arrangement

a

L~ __ I

I

O'

10'

5'

of Tlnamou Breeding Pens.

�-111-

Table l.

Pen schedule for tinamou, 1970.

Pen
Number

Protein Level
16%
22%
T

1
2

T

3

x

4

T

x
T

T
T
T

5

x

T

6

T
T

7
x

8

x
T

T

9
10

T

Alleyway

x

Number Pens on
Test

4

Key:

Mating
Flock*
Selected Groups
(8 Hens and 4 Cocks) (2 Hens and 1 Cock)

T = Pen on Test;

x
T
T

4

1

4

x = Pen not on Test·

* Pens numbe r 3, 8, and alleyway was used as 1 combined pen for flock mating.

Following normal culling procedures, settings of tinamou eggs were made
alternately (except for setting number 1) in the test and control incubators
every 7 days insofar as possible in 1970. Relative humidity in the control
incubator was held at the standard 55 percent relative humidity (8~0 wet
bulb) for the first 2 settings and then increased to 56 percent relative
humidity (85~0 wet bulb) based upon recommended procedures for Robbins
incubators. In the test incubators, eggs were incubated at a relative
humidity of 50 percent (82-3/40 wet bulb) using a Humidaire Model 50 for
the.first 3 settings and a Robbins Model C-l for the remainder. Temperature
in all incubators was maintained at a normal 99.750 F. Setting number 7
was placed in the control incubator rather than the test incubator because
of problems encountered in maintaining 50 percent relative humidity in the
Humidaire incubator and in anticipation,of securing a new Robbins Model C-l
incubator.

�-112-

Birds in even numbered pens 2, 4, 6, and 10 were fed a standard game
bird breeder feed containing 22 percent protein level. Those in odd
numbered pens 1, 5, 7, and 9 were fed a similar feed but containing 16
percent protein.
Eggs produced were marked to facilitate determination
of fertile eggs produced and numbers of fertile eggs that hatched.

RESULTS AND DISCUSSION
Table 2 lists a comparison of pale spotted tinamou production at the Fort
Collins Wildlife Research Station for the period 1967 through 1970, and
field releases made from 1969 through 1971. A gradual increase in numbers
of fertile eggs and numbers of chicks hatched is evident during the period,
using approximately equal numbers of breeders from 1968 through 1970.

Incubation

Humidity

Test

Table 3 lists hatching success of eggs in the test incubators (50 percent
relative humidity) in comparison with eggs in the control incubator (55
to 56 percent relative humidity).
Results showed a 6.38 percent better
hatch in the control incubators when considering percent of total eggs
hatched (46.58 % compared with 40.20%) and a 9.08 percent better hatch in
the control incubators when considering fertile eggs hatched (67.28%
compared with 58.20%).
Higher percentages of dead germs (21.64% for the
test incubators, compa~ed with 19.23% for the control incubator) and of
pipped eggs (6.70% for the test incubators, compared with 1.28% for the
control incubator) showed the incubation humidity was too low for this
species.
Results indicated the normal incubation relative humidity of 55-56 percent
(84~-85~0 wet bulb) was therefore better than a test relative humidity of
50 percent (82-3/40 wet bulb).

Flock Mating

Versus

Selected

Group Mating

Test

Table 4 lists a comparison of numbers of eggs produced and numbers of
fertile eggs produced by 8 hens and 4 cocks in a community pen with the
same number of birds in 4 pens of 2 hens and 1 cock each. All birds were
fed a standard 22 percent protein feed and treated as alike as possible.
A much larger number of total eggs was laid by the tinamou held in 4
separate pens with selected groups of 2 hens and 1 cock each (248 eggs)
compared with a community pen of 8 hens and 4 cocks (62 eggs).
In addition,
190 of the eggs laid by birds in the 4 selected group pens were fertile as
compared with only 32 for the community pen. Results obtained in 1970 were
similar to those secured in 1969.

�Table 2.

A summary of pale spotted tinamou production,

Item
Number of Breeders
Sexes

Total eggs laid

1967-1970, and field releases made 1969-1971.

1967

1968

Year
1969

1970

1971

17

37

36

36

0

24H;12C

24H; 12C

493

478

mixed
(connnunity pen)
177

mixed
(conununity pen)
335

1/

I

Total eggs set

166

316

434

I-'
I-'
W

428

I

Number fertile eggs

74

211

268

296

Percent fertile eggs

44.6

66.8

61. 8

69.2

Number chicks hatched

44

108

137

187

Percent fertile eggs hatched

59.5

51. 2

51.1

63.2

Number released, Baca County

None

None

42

40

82

1/ Original source of breeding stock provided by Foreign Game Introduction Program of United States Department
of the Interior included 20 birds received on June 25, 1966 and 17 birds received on March 8, 1967, all from
Argentina, South America.

�Table 3. A comparison of hatchability of pale spotted tinamou eggs under approximately 55 percent relative
humidity (normal) versus approximately 50 percent relative humidity, 1970.

Item

On Test

1/

Hatch
Number

Date
Set

Number
Set

1

5-18
6-1
6-15
7-6
7-20
8-3
8-17
8-31

12
14
21
43

3
5
8

10
12
14
16

3
4
4

19
8
8

Dead
Germs

Number
Pipped

4
7
6
5
5

o

6

2

2
2

o
1

4
2

Number
of
Cripples

Number
of
Good Hatch

Percent Hatch
Total
Fertile
Eggs
Eggs

1

4
3

9
8

33.33
21.42
47.61
34.88
54.54
40.74
42.85
34.78

44.44
30.00
58.82
62.50
72.00
57.89
60.00
53.33

78 (40.20)

40.20

58.20

10
12
15
20
13

76.92
66.66
53.57
54.05
41.93
30.43
U.93
14.81
57.69

76.92
80.00
83.33
71.42
81.25
43.75
56.52
25.00
88.23

o
o
o
o
o
o
o

10
15
18

27
21
23

6

4

8

5

194

60 (30.92)

42 (21.64)13 (6.70) 1 (0.51)

13
18
28
37
31
23
31
27
26

o

o
o
o
o

7
8

3
3
3
8
2
8

10

o

11
9

8
2

o

Sub-Total

234

72 (30.76)

45 (19.23) 3 (1.28) 3 (1.28) 109 (46.58)

46.58

67.28

Totals

428

132 (30.84)

87 (20.32)16 (3.73) 4 (0.93) 187 (43.69)

43.69

63.17

Sub-Total
Control Jj

33

Number
of
Infertile

2

5-25

4

6- 8

6
7
9
11
13
15
17

6-22
6-29
7-13
7-28
8-11
8-24
9-8

3

10
9

15

1
1
1

o
o
o
o
o
o
o
3

o

11

7

13
4

15

1/ Eggs incubated at 82-3/40 F wet bulb reading (50% reI. hum.) and temperature of 99.750 F. Setting numbers
1, 3, and 5 incubated in Humidaire Model 50 incubator and the remaining settings on test were incubated in a
Robbins Model C-l incubator.
~/ Eggs in settings number 2 and 4 incubated at 84~0 F wet bulb reading (55% reI. hum.) and the remaining
controls were incubated at 85~0 F wet bulb reading (56% reI. hum.) and temperature of 99.750 F. All settings
were incubated in a Robbins Model H-7 incubator.

I

I-'
I-'
+'
I

�Table 4. A comparison of numbers of pale spotted tinamou eggs laid by a flock consisting of 8 hens and
4 cocks with four selected groups of 2 hens and 1 cock each. 1/
Number
of
Infertile

Number
Fertile
Eggs

Number
of
Good Hatch

Pen
Number

Total
Eggs

Number
of
Culls

Number
Set

Flock

62

3

59

27

16

32

Sub-Total

62

3

59

27 (45.76)

16 (27.11)

32 (54.23) 16 (27.11)

Dead
Germs

16

Percent Hatch
Fertile
Total.
Eggs
Eggs Set
27.11

50.00

•...
•...
I

2

45

3

42

10

13

32

19

45.23

59.37

4

73

7

66

10

17

56

39

59.09

69.64

6

40

6

34

13

9

21

12

35.29

57.14

10

90

6

84

3

18

81

63

75.00

77.77

Sub-Total

248

22

226

36 (15.92)

57 (25.22) 190 (84.07)133 (58.84) 58.84

70.00

Totals

310

25

285

63

73

1/ All pens were fed 22 percent protein feed.

222

149

V1
I

�-116-

22 Percent

Protein

Versus

16 Percent

Protein

Feed Test

Table 5 lists a comparison of numbers of fertile eggs produced and numbers
of fertile eggs that hatched by tinamou breeders fed 16 percent protein
level feed with those fed a 22 percent protein level feed.
Selected groups
of 2 hens and 1 cock each in pen numbers 1, 5, 7, and 9 were fed the 16
percent protein feed and birds in pen numbers 2, 4, 6, and 10 were fed a
standard 22 percent protein level feed.
More fertile eggs were produced by the breeders fed the standard 22 percent protein level feed (190) compared with a similar feed containing 16
percent protein level (74). A higher percent of the fertile eggs hatched
were also from the birds fed the 22 percent protein feed (70.00%) compared
with the 16 percent protein feed (51.35%).
From these comparisons, it is evident that the standard 22 percent protein
level game bird breeder feed was much better than the 16 percent protein
level feed for pale spotted tinamou egg production and hatchability during
1970.

East Pens Versus West Pens
Although not planned as a separate test, a comparison of egg production by
east versus west pen location has shown the importance of providing adequate
natural cover in the breeding pens.
In 1969, a comparison of egg production
by 4 pens on the west side, each containing 7 or 8 established tall wheatgrass clumps, with 4 pens on the east side with no grass clumps, showed
highest numbers of total eggs were produced in the west pens (235 compared
to 193) and 162 fertile eggs were produced in the west pens compared with
only 73 fertile eggs in the east pens.
Prior to the 1970 egg laying season, a minimum of 8 clumps of tall and/or
crested wheatgrass were planted in the east pens and wire cages installed
to protect most of these growing clumps.
The established cover was left
in the west pens as well, so all pens had growing natural cover for 1970.
Table 6 lists a comparison of numbers of eggs laid and fertile eggs laid
by tinamou in the west pens with the east pens in 1970. Total eggs produced was 209 for the 4 west pens and 207 for the 4 east pens.
Numbers
of fertile eggs produced was 143 for the 4 west pens and 121 for the 4
east pens.
Location of pens plus a lack of adequate cover in the east
pens apparently caused this variation in egg production in 1969 but this
variation was considerably reduced in 1970 with the replanting of adequate
natural cover in the form of growing grass clumps.
The variation in
numbers of fertile eggs produced would probably be further reduced in
the future as the newly planted grass clumps mature but may never be
entirely eliminated because of a tall juniper windbreak on the west side

�Table 5. A comparison of numbers of pale spotted tinamou eggs produced and number hatched using 16 percent and 22 percent protein levels in the feed, 1970.

Pen
Number

Total
Eggs

Number
of
Culls

Number
Set

1

28

0

28

8

7

5

63

12

51

16

7

26

5

21

9

51

8

Sub-Total

168

25

Item

On Test
(16% Protein
Feed)

Control
(22% Protein
Feed)

Number
of
Infertile

Number
of
Good Hatch

Percent Hatch
Total
Fertile
Eggs Set
Eggs

Percent
Fertile
Eggs

13

46.42

65.00

71.42

22

13

25.49

37.14

68.62

5

6

10

47.61

62.50

76.19

43

40

1

2

04.65

66.66

06.97

143

69 (48.25)

36 (25.17) 38 (26.57)

26.57

51. 35

51. 74

Dead
Germs

,...
,...
•....•
I

I

2

45

3

42

10

13

19

45.23

59.37

76.19

4

73

7

66

10

17

39

59.09

69.64

84.84

6

40

6

34

13

9

12

35.29

57.14

61. 76

10

90

6

84

3

18

63

75.00

77.77

96.42

Sub-Total

248

22

226

36 (15.92)

57 (25.22)133 (58.84)

58.84

70.00

84.07

Total

416

50

369

105 (25.24)

93 (22.35)171 (46.34)

46.34

64.77

63.93

�Table 6. A comparison of numbers of pale spotted tinamou fertile eggs produced and number of fertile
eggs hatched by 4 west and 4 east pens, 1970.

Item
West Pens

Pen Number

Protein Level

Culls

Laid

Egg Totals
Set
Infertile

Fertile

Hatched

1

16%

0

28

28

8

20

13

2

22%

3

45

42

10

32

19

4

22%

7

73

66

10

56

39

5

16%

12

63

51

16

35

13

Sub-Total

22

209

187

44

143

84
I

t-"
t-"

East Pens

6

22%

6

40

34

13

21

12

7

16%

5

26

21

5

16

10

9

16%

8

51

43

40

3

2

10

22%

6

90

84

3

81

63

Sub-Total

25

207

182

61

121

87

Totals

47

416

369

105

264

171

00
I

�-119-

of the breeding pens, which provides additional shelter and shade to
the west pens. Natural cover in the form of growing grass clumps provides both shade and shelter for the breeders as well as for eggs laid.
By alternating pens by east and west location in all tests run in 1969
and 1970, bias due to pen location should have been eliminated.

��April, 1971

-121-

JOB PROGRESS REPORT

State of

~C~O~L~O~RAD~~O _

Project No.

W-37-R-24

Work Plan No.
Job Title

Job No.

2

Study of Tinamou Adaptability

Period Covered:
Personnel:

19

Game Bird Survey

April 1, 1970 to November 30, 1970

Donald M. Hoffman, Dale W. Stahlecker and Warren D. Snyder.

ABSTRACT
Forty pale spotted tinamou, Nothura darwinii salvadorii, were released in
May, 1970. This was the second release on U. S. Forest Service rangeland in
southern Baca County, Colorado. As in 1969, follow-up searches and checks
on reported sightings failed to provide evidence that tinamou remained, survived or reproduced in the vicinity of the release site.

��-123-

STUDY OF TINAMOU ADAPTABILITY
Warren D. Snyder
P. S. OBJECTIVE
To determine the ability of pale spotted tinamou to establish reproducing
populations in areas of mixed tall and mid-grasses, sandsage and yucca
on the Comanche National Gr~sslands, Baca County, Colorado
SEGMENT OBJECTIVES
1.

Introduce pale spotted tinamou.

2.

Determine dispersal of pale spotted tinamou and success of plant.
METHODS AND MATERIALS

Reference is made to Snyder (1969) for review of methods and materials used
in this study.
RESULTS AND DISCUSSION
Pale spotted tinamou raised at the Fort Collins Wildlife Research Station
under Work Plan 19, Job 1, were released on May 1, 1970 by Donald M.
Hoffman. This release included 15 males and 25 females. The location of
the release, along the Sand Arroyo in southern Baca County, was the same as
that of the previous year (Snyder 1969).
The vicinity of the release and surrounding localities were searched on
June 11-12, August 17-19 and November 1 and 2, 1970. The birds were not
observed, nor were there any indication of their presence in the area.
Bert Widhalm, District Conservation Officer, and Carrizo District personnel
of the U. S. Forest Service were contacted concerning sightings or reported
sightings. One local resident reported seeing the birds frequently during
the summer in a location approximately ten miles east of the release. Contact of this person left doubt as to the accuracy of his observations. His
description of the bird and its habits did not fit the tinamou.
In summary, there was no evidence of survival or reproduction of any of
the tinamou released either in 1969 or 1970. One additional release is
planned in 1970 with follow-up surveys to be made.

�-124-

Fig. 1. Pale spotted tinamou at release site in Baca County.
Don Domenick.

Photo by

�-125-

LITERATURE CITED
Snyder, W. D. 1969. Study of tinamou adaptability. Colorado Div. of
Game, Fish and Parks. Game Res. Rpt. April, pp. 177 -181.

Prepared

by

!f!.~yd'it~
Wildlife Researcher

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                  <text>-1-

April, 1971

JOB PROGRESS REPORT

State of

~C~O~~L~O~RAD~~O _

Work Plan No.
Job Title

1

Job No.

16

Pheasant Nest Site Selection Study

Period Covered:
Personnel:

Game Bird Survey

W-37-R-24

Project No.

April 1, 1970 to March 31, 1971

Donald M. Hoffman, Warren D. Snyde.r, Bruce C. Sigler, Dale W.
Stahlecker, Robert L. Schmidt, Lawrence A. Webster, John F.
Corey, Gary Brown, John Ellenberger, Mark Frazier, Jon Hooper,
Tom Lines, Roger Lowery, Jon Moser, Henry Wilson and Michael
Zgainer.
ABSTRACT

Each of the 4 quadrats was stocked with a m~n~mum of 13 hens and 3 cocks
by March 11, 1970 (Segment 23) in preparation for nesting studies in 1970.
Eighty pheasant nests were found in the 48 nesting plots, representing 6
vegetative species or combination of species. Sixteen nests were found in
8 alfalfa mowed plots; 15 nests Were found in 4 alfalfa-crested wheatgrass
mowed plots; 10 nests in 4 tall wheatgrass mowed plots; 9 each nests in
4 intermediate wheatgrass mowed and 4 intermediate wheatgrass unmowed plots;
8 nests in 4 tall wheatgrass seeded plots; and 7 nests in 4 smooth bromegrass mowed plots. Fewer numbers of nests were found in other vegetative
types and treatments represented. In comparing nests per vegetative type
and treatment, alfalfa-crested wheatgrass mowed, ranked highest with 3.75
nests per plot; followed by tall wheatgrass mowed, with 2.50 nests per plot;
intermediate wheatgrass mowed, and intermediate wheatgrass unmowed, each
with 2.25 nests per plot; tall wheatgrass seeded, and alfalfa mowed, each
with 2.00 nests per plot; and smooth bromegrass mowed, with 1.75 nests per
plot. Sixty-six percent of all nests in 1970 were established 15 feet or
less from plot perimeters. Average clutch size of 46 nests, where it was
possible to determine this, was 11.2. A minimum of 13 hens and 3 cocks,
wild-trapped in Phillips, Sedgwick, and Larimer counties in early 1971,
were restocked in each of the 4 quadrats by January 28, 1971 for continuation of the study in 1971 (Segment 25).

��"

-.)-

PHEASANT NEST SITE SELECTION STUDY
Donald M. Hoffman

Field work on this study was conducted similar to the previous year's work
with three separate nest searches completed during the nesting season of
1970. The change-over of three vegetative types used very little or not
practical to establish along roadsides was completed in the fall of 1969 and
no additional p1antings were made in 1970.
P. S. OBJECTIVE
To compare pheasant nesting of, and success in (1) winter wheat; (2) alfalfa;
(3) crested wheatgrass; (4) hairy vetch; (5) white sweet clover; (6) alfalfacrested wheatgrass mixture; (7) smooth bromegrass; (8) tall wheatgrass; and
(9) intermediate wheatgrass.
SEGMENT OBJECTIVES
1.

To maintain plots.

2.

To measure nesting preference.

3.

To determine nesting success by cover type.

4.

To obtain wild pheasants and care for them following release in plots.
METHODS AND MATERIALS

Wild pheasants were captured with a vehicle, spotlights, and hand nets at
night in Phillips and Sedgwick counties in Segment 23. All 4 quadrats were
restocked with a minimum of 13 hens and 3 cocks by March 11, 1970. Limited
Cooper's hawk predation occurred on the wing-clipped pheasants held in the
quadrats after this date but this was not serious.
Phenology measurements were again secured in 1970 to ascertain readiness of
the various vegetative species and cover types for use by nesting pheasants.
An initial nest search in 1970 was made during the period June 4 and 5 with
10 men, including 5 W.C.O. trainees and others. Each plot was systematially
searched using ropes to divide the plots into narrow segments. Nests were
marked, recorded, and removed to encourage renesting. A second nest search
was made during the period June 22 through July 3, 1970 by 2 men searching
each plot systematically in strips outlined by lath markers. A third and
final search was made by 5 men using methods similar to the second search
during the period July 20 through 27, 1970, preceding and following the windrowing of half of the individual plots on July 27, 1970. A well trained
bird dog belonging to Warren Snyder was used to advantage in the first and
last nest searches.

�-4-

Distances were measured from each nest to the nearest edge and to the apex
or base of the triangular plot, whi.chever was closest, with a steel tape.
Each nest could later be located on a scaled diagram of the quadrats.
Number of eggs, fate of the nest and estimated age of embryos were recorded.
In addition, vegetative heights and vegetative species in the 4 cardinal
direc.tions from each nest were recorded.
Prior to the windrowing of vegetation in half of the plots on July 27, 1970,
all but 4 of the 64 wing-clipped pheasants were caught with hand nets. Nineteen of these were released on Means Lake property in Phillips County and
the remainder were released outside the quadrats at the Fort Collins Wildlife
Research Station.
Two incubating hens were killed by the windrowing machine
and the other 2 were not recovered.
Five wild pheasants, including 3 hens and 2 cocks were captured with cloverleaf traps at the Fort Collins Wildlife Research Station during early
January, 1971. Fifty-seven additional wild pheasants, including 49 hens
and 8 cocks were captured with 4-wheel drive vehicles, hand spotlights, and
hand nets during 6 trap-nights in Phillips and Sedgwick counties.
These
were used to restock the 4 quadrats with a minimum of 13 hens and 2 cocks by
January 28, 1971 for continuation of the study in 1971. All pheasants were
wing-clipped and banded with aluminum leg bands prior to release in the
quadrats.

Plot Composition
Plot Composition

for 1970 (Treatment

Following

1969 Nesting

Season)

The changing of vegetative types used very little, or not practical to
establish along roadsides, started in 1968, was continued in the fall of
1969. The remaining winter wheat plots were reworked and seeded to smooth
bromegrass and the 4 remaining white sweet clover plots were reworked and
seeded to tall wheatgrass.
Plot Number

1.

Originally

Winter Wheat,

Now Smooth Bromegrass

Four of the original winter wheat plots were reworked and seeded to smooth
bromegrass in the fall of 1968. These were all mowed in the summer of
1969. The remaining 4 winter wheat plots were reworked and seeded to
smooth bromegrass in the fall of 1969.
Plot Number

2.

Alfalfa

These were all mowed and maintained following the 1969 nesting season so
1970 composition was comparable with 1968 and 1969. Two of these plots
including Number 2, Interior, NW Quadrat,
and Number 2, Exterior, NE
Quadrat, are approaching alfalfa-crested wheatgrass mixtures due to invasion
by crested wheatgrass.

�Plot Number

3.

Cres ted Wheatgras s

Four of these were mowed and 4 were left unmowed following the 1969 nesting
season so that a residual cover check could be secured in 1970 on half of
the plots.
Composition for 1970 was therefore comparable to that of 1969.
Plot Number 4.

Originally

Hairy Vetch,

Now Intermediate

Wheatgrass

All of these plots were reworked and seeded to intermediate wheatgrass
following the 1968 nesting season.
Four of these intermediate wheatgrass
plots were mowed and 4 were left unmowed following the 1969 nesting season,
so a residual cover check could be secured in 1970 on half of the plots.
Plot Number

5.

Originally

White Sweet Clover, Now Tall Wheatgrass

Four of the original white sweet clover plots were reworked and seeded to
tall wheatgrass in the fall of 1968. These were all mowed in the summer of
1969. The remaining 4 white sweet clover plots (volunteer forbs in 1969)
were reworked and seeded to tall wheatgrass in the fall of 1969.
Plot Number 6.

Alfalfa-Crested

Wheatgrass

Four of these were mowed and 4 were left unmowed following the 1969 nesting
season, so that a residual cover check could be secured in 1970 on half of
the plots.
Composition for 1970 was therefore comparable to 1969.

Plot Composition

for 1971 (Treatment

Following

1970 Nesting Season)

The changing of vegetative types used very little, or not practical to
establi8h along roadsides, started in 1968, completed in the fall of 1969.
No additional changes were made during 1970.
Plot Number

1.

Originally

Winter Wheat, Now Smooth Bromegrass

The 4 original winter wheat plots seeded to smooth bromegrass in the fall
of 1968, were left unmowed in the summer of 1970. The 4 original winter
wheat plots, seeded to smooth bromegrass in the fall of 1969, were mowed in
the summer of 1970. A residual cover check will be secured in 1971.
Plot Number 2.

Alfalfa

Four of these were mowed and 4 were left unmowed in 1970 so that a residual
cover check may be secured in 1971. Two of the alfalfa plots including
Number 2, Interior, NW Quadrat, and Number 2, Exterior, NE Quadrat, are
approaching alfalfa-crested wheatgrass mixtures due to invasion by crested
wheatgrass.
Plot Number

3.

Crested Wheatgrass

Four of these were mowed and 4 were left unmowed following the 1970 nesting
season so that a residual cover check may be secured in 1971 on half of
the plots.
Composition for 1971 is therefore comparable to that of 1970.

�-6-

Plot Number 4.

Originally

Hairy Vetch,

Now Intermediate

Wheatgrass

All of these plots were reworked and seeded to intermediate wheatgrass following the 1968 nesting season.
Four of these intermediate wheatgrass plots
were mowed and 4 were left unmowed following the 1970 nesting season, so a
residual cover check may be secured in 1971 on half of the plots.
Plot Number

5.

Originally

White Sweet Clover, Now Tall Wheatgrass

The 4 original white sweet clover plots seeded to tall wheatgrass in the fall
of 1968, were left unmowed in the summer of 1970. The 4 original white
sweet clover plots seeded to tall wheatgrass in the fall of 1969, were mowed
in the summer of 1970. A residual cover check will be secured in 1971.
Plot Number

6.

Alfalfa-Crested

Wheatgrass

Four of these were mowed and 4 were left unmowed folloWing the 1970 nesting
season, so that a residual cover check may be secured in 1971 on half of
the plots.
Composition for 1971 is therefore comparable to 1970.

RESULTS AND DISCUSSION
Stocking

of Pheasants

for 1970

An attempt to live-trap pheasants in September, 1969 in the Holyoke area
was made but losses folloWing capture were heavy.
Of 18 captured (mostly
juveniles) 4 died in transit to Fort Collins and 4 died while held in the
covered pen before moving to the southeast quadrat on October 9, 1969. At
least 2 additional birds from this source are known to have died while held
in the southeast quadrat.
The wing-clipped pheasants used in the 1969 nesting studies were recaptured
with hand nets at the end of the 1969 nesting season and placed in a large
covered pen in an attempt to hold these over for use in 1970. An unusually
heavy, wet snow in early October, 1969 broke the top of the pen down and 67
surviving pheasants (57 hens and 10 cocks) were released in the southeast
quadrat on October 9, 1969. Overwinter losses, mostly from great-horned owl
predation, were heavy.
Approximately 34 percent were lost from this cause
from October 9, 1969 to February 9, 1970 and an additional 33 percent were
lost from the same cause from February 9, 1970 to March 6, 1970. Limited
Cooper's hawk predation on the wing-clipped pheasants occurred after the
final stocking of the quadrats on March II, 1970, but this was not serious.

Location

of Nests

Eighty pheasant nests were found in the 48 triangular-shaped
nesting plots
representing 6 vegetative species or combination of species.
Figures 1
through 4 show the approximate location of nests in the various quadrats.
Table 1 lists pheasant nests found by vegetative cover type and treatments
in 1970.

�-7-

370'

4

2

Agin (urn)
""~

Mesa (m)

o
3
12' Agcr (m)

o
5

0::

Agel (rr)

~

o

0::
W

~

Brin (m)

0::

o
H
0::

~

Z

H

12'

6
Mesa~gcr(urn
Brin (rr)~

12'

Detailed

Fig. 1.

Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan I, Job 16

Nest Site

Location of pheasant nests in northwest quadratp 1970.

~
0
W

-

�-8-

370'

\\

2

\\

5

/00 6
Mesa- Agcr

II

12'

~
~
(l:

. ~1-,.

.:

Agel (rr) /

!o/

Agel (m)

7

7/ o 5

3
o Agcr (m)

(m)

I:t:

o
Brin

o

(m)

Brin (rr) ~

X

£iI

o

12'

4
Agin (

Mesa-Agcr (urn)

2

/

Ager

6

Fig. 2

G

Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan I, Job 16

(um~

3

12'

Detailed

~

~ 8

Nest Site

Location of pheasant nests in northeast quadrat, 1970.

)

-

�-9-

370'

4

7/

7

2
Mesa

(m) /

f
Brin (rr)

~

0::
W

6

E 0
Mesa-Agcr

(m)

~

~o'~

------0
12'

6

12'

Fig. 3 •

Location

Detailed Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan 1, Job 16

Nest Site

of pheasant

1970.

nests in southwest

quadrat,

�-10-

370'

3

6
12'

Mesa-Agcr (urn)

12'

12'

Detailed

Fige4 _

Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan 1, Job 16

Nest Site

Location of pheasant nests in southeast quadrat, 1970

0

�Table 1.

Number of pheasant nests found in 48 plots representing various vegetative types and treatments, 1970.

Plot
Number

Type and Treatment

NW

guadrat
NE
SW

SE

1

Smooth bromegrass (mowed)

1

3

1

1

Smooth bromegrass (seeded)

1

0

2

Alfalfa (mowed)

7

3

Crested wheatgrass

(mowed)

3

Crested wheatgrass

(unmowed)

4

Intermediate wheatgrass

(mowed)

Total

Number
Plots

Nests/
Plot

2

7

4

1.75

0

0

1

4

0.25

2

1

6

16

8

2.00

0

2

0

1

3

4

0.75

0

0

0

1

1

4

0.25

4

1

2

2

9

4

2.25
I

2

1

2

4

9

4

2.25

(mowed)

1

4

4

1

10

4

2.50

(seeded)

4

1

0

3

8

4

2.00

Alfalfa-crested wheatgrass (mowed)

5

2

2

6

15

4

3.75

Alfalfa-crested wheatgrass

1

0

0

0

1

4

0.25

26

16

12

26

80

48

1.66

4

Intermediate wheatgrass

5

Tall wheatgrass

5

Tall wheatgrass

6
6

Totals Average

(unmowed)

(unmowed)

I-'
I-'
I

�-12-

By numbers of nests, alfalfa mowed (16 nests) ranked highest for nest
establishment in 1970, followed by alfalfa-crested wheatgrass mowed (15 nests);
tall wheatgrass mowed (10 nests); intermediate wheatgrass mowed and intermediate wheatgrass unmowed (each with 9 nests); tall wheatgrass seeded (8
nests); and smooth bromegrass mowed (7 nests).
Fewer numbers of nests were
found in other vegetative types and treatments represented.
Numbers of nests per cover plot were calculated because of a difference in
numbers of plots by cover type and treatment.
In this comparison, alfalfacrested wheatgrass mowad ranked highest with 3.75 nests per plot, followed
by tall wheatgrass mowed with 2.50 nests per plot, intermediate wheatgrass
mowed and intermediate wheatgrass unmowed each with 2.25 nests per plot,
tall wheatgrass seeded and alfalfa mowed each with 2.00 nests per plot, and
smooth bromegrass mowed with 1.75 nests per plot.
Sixty-six percent of all nests in 1970 were established 15 feet or less from
plot perimeters as shown in Table 2. During 1970, 48 nests were found in
exterior plots and 32 nests in interior plots.
The terms "interior and
exterior" refer to the location of the individual plots in relation to a
central point where all 4 quadrats join. These are indicated in the margins
of Figures 1 through 4. Figure 5 shows an experimental plot of smooth bromegrass seeded in the fall of 1968 and mowed in the summer of 1969. A search
for nests in a plot of intermediate wheatgrass seeded in the fall of 1968
is shown in Figure 6. Figure 7 shows an experimental plot of tall wheatgrass seeded in the fall of 1968 and mowed in the summer of 1969. This
cover type furnishes good winter cover as well as good nesting cover at
the Fort Collins Wildlife Research Station.
Figure 8 shows the method used
in hand netting Wing-clipped pheasants located with the help of a well
trained bird dog. Pheasants were removed from the quadrats prior to windrowing of the vegetation in half of the plots on July 27, 1970.

Clutch Sizes
Thirty-four nests contained unknown clutch sizes.
These were nests where
the hen was still laying or nests which were abandoned for one reason or
another without incubation having started.
The average clutch size of the
remaining 46 nests which included both incubating nests and hatched nests
was 11.2 eggs, ranging from 5 to 21 (Table 3).
Considerable variation was found in clutch sizes between the various
cover types and treatments as shown in Table 3. Largest average numbers of
eggs per clutch was found in intermediate wheatgrass unmowed (6 nests
averaged 14.5 eggs); intermediate wheatgrass mowed (7 nests averaged 12.4
eggs); alfalfa mowed (9 nests averaged 12.2 eggs); smooth bromegrass mowed
(4 nests averaged 11.0 eggs); and alfalfa-crested wheatgrass unmowed (1 nest
averaged 11.0 eggs).

�I

I-'

W
I

Fig. 5. A plot of smooth bromegrass seeded in the fall of 1968 and mowed in the summer of 1969.
nests were found in the four smoothbromegrass, mowed plots in 1970. (D. Hoffman, photo).

Seven pheasant

�,....I

+:I

Fig. 6. Searching for nests in a plot of intermediate wheatgrass seeded in the fall of 1968 and left unmowed in the
summer of 1969
Nine pheasant nests were found in the four intermediate wheatgrass, unmowed, and nine in the four
intermediate wheatgrass, mowed plots in 1970.
(Do Hoffman, photo).
0

�I
•.....

V1
I

Fig. 7. A plot of tall wheatgrass seeded in the fall of 1968 and mowed in the summer of 1969.
found in the four tall wheatgrass, mowed plots in 1970. (D. Hoffman, photo).

Ten pheasant nests were

�I

~

0'\
I

Fig. 8. Hand netting a wing-clipped pheasant, located with the assistance of a well trained dog.
removed from the quadrats prior to windrowing of the vegetation.
(D. Hoffman, photo).

Pheasants

were

�-17-

Table 2.

Distance of pheasant nests from plot perimeters, 1970.

Distance
From Nearest
Edge

Number
Nests

Percent

0 - 5'

13

16

5 - 10'

25

31

10 - 15'

15

19

15 - 20'

11

14

20 - 25'

6

8

25 - 30'

9

11

30 - 35'

1

1

35 - 40'

0

0

Total

80

100

Fate of Nests
Of the 80 nests found and recorded, 66 were determined to be in an active
state of laying or incubation, 7 were hatched successfully, and 7 were
apparently abandoned for one reason or another. All nests were recorded
and removed to encourage renesting. This also prevented the recording of
the same nest more than once.
Of the 7 nests which hatched successfully, 2 were in alfalfa mowed plots,
2 in intermediate wheatgrass unmowed, 2 in alfalfa-crested wheatgrass mowed,
and 1 in intermediate wheatgrass mowed. Of 86 total eggs in 7 nests which
had successfully hatched, 71 (82.55 percent) of the eggs hatched.
Of the 7 apparently abandoned nests, 1 was hailed on with eggs showing hail
cracks, 1 was ruined by a predator (probably magpie or crow) and 5 were
abandoned for unknown reasons.
Periods of Nest Establishment
Table 5 lists periods of nest establishment based upon aging egg embryos
according to procedures outlined in Game Information Leaflet Number 15
(Sandfort 1965).

�Table 3.

Clutch sizes found in 46 incubating or hatched pheasant nests in various vegetative types, 1970.

Clutch
Size

1
Brin
(m)

1
Brin
(rr)

2
Mesa
(m)

3
Agcr
(m)

1
1

1
1

Plot Number and Treatment
3
4
4
5
Agcr
Agin
Agin
Agel
(urn)
(m)
(urn)
(m)

5

6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22

5

Agel
(rr)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(urn)

1
3
2
1

2

2
2

1
1

1
1

1
1

1
1
1
1

1

1

1
1
1

1
1
2

2
1

1
1
1

2

1
1

Total

1
5
6
6
3
1
3
3
2
6
2
2
1
3
1
0
1

Totals

4

0

9

3

0

7

6

5

3

8

1

46

Average
Clutch

11.0

-

12.2

8.0

-

12.4

14.5

9.4

9.3

9.5

11.0

11.2

I

I-'

00
I

�Table 4.

Fate of 80 pheasant nests in various vegetative types, 1970.

Type

1
Brin
(m)

Ac tive (Laying or
Incubated) 7

1
Brin
(rr)

2
Mesa
(m)

5
Agel
(rr)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(urn)·

Total

I

t-"

1

13
2

Abandoned or
Unsuccessful

1

7

Plot Number and Treatment
5
3
4
4
Agel
Agcr
Agin
Agin
(urn)
(m)
(urn)
(m)

1.0
I

Hatched

Total

3
Agcr
(m)

1

16

3

3

1

1

8

6

1

2

9

9

5

12

1

66

2

7

7

1

1

3

1

9

10

8

15

1

80

�Table S.

Estimated period of nest establishment in various vegetative types, 1970. 11

Period

1
Brin
(m)

1
Brin
(rr)

April 16-30

2
Mesa
(m)

3
Agcr
(m)

Plot Number and Treatment
5
3
4
4
Agcr
Agin
Agel
Agin
(urn)
(m)
(um)
(m)

2

5
Agel
(rr)

1

1

May 16-31

2

3

June 1-15

2

3

1

June 16-30

1

6

2

2

Total

4
2

3

1
1

6
Mesa-Agcr
(um)

1

1

May 1-15

6
Mesa-Agcr
(m)

3

2

3

13

16

5

3

1

1

2

5

1

24

I

N

July 1-15

1

1

1

July 16-31

1

Unknown

3

Total

7

1

16

2

1

6

1

3

1

1

3

1

3

3

9

9

10

8

IS

14
1

80

1/ Period of nest establishment based upon aging of embryos according to Game- Information Leaflet No. IS (Sandfort
1965). First nest search was made June 4-5, 1970; second was made June 22-Ju1y 3, 1970; third was made July
20-27, 1970.

0
I

�-21-

An initial peak in nest establishment occurred during the period May 16-31,
1970 or approximately 2 weeks later than in 1969. Heights of vegetation
on May 15, 1970 were about half that found on May 16, 1969 due to a cold,
wet spring in 1970. Weather factors, therefore, were the apparent cause of
this delay in nesting activity.
Additional peaks in nest establishment
occurred during the periods June 1-15, and June 16-30, 1970; similar to
those found in 1969. Highest number of nests (24) was recorded for the
period June 16-30, 1970. Some hens continued to recycle until removed
from the quadrats prior to windrowing of the vegetation in half of the individual plots on July 27, 1970.
Residual cover from 1969 was left in half of the crested wheatgrass, intermediate wheatgrass, and alfalfa-crested wheatgrass plots (12 plots out of
48) in order to compare nesting use with mowed plots. This residual cover
was flattened by the early, wet snows in September and October, 1969 and
did not recover so, for all practical purposes, standing residual cover was
non-existent in 1970. In 1970, only 3 nests out of the 19 established prior
to May 31, 1970 (15.78 percent) were found in plots having this flattened
residual cover from 1969. A good test of residual cover was not possible in
1970 for this reason.

Phenology
Heights of vegetation measurements were made on April 15 and 27; May 11 and
22; June 2, 12, and 23; and July 3 and 16, 1970. Table 6 summarizes average
heights of vegetation on April 15, May 11, June 12, and July 16, 1970.
Heights of the new growth vegetation on May 11, 1970 were generally about
half that found on May 16, 1969, but had largely caught up by mid-June.
The
cold, wet spring of 1970 probably caused this lateness of vegetation growth.
Weather factors also apparently caused a delay in nesting activity in 1970
(see section on periods of nest establishment).
In contrast to the 1969 season when vegetative growth appeared to reach a
maximum for the season in mid-June, most vegetative species in 1970 continued
to grow in height throughout the nesting season.
This was probably due to
differences in spring and summer moisture and temperatures.
Good early soil
moisture has been maintained for the 1969, 1970, and 1971 nesting seasons
by complete plot irrigations during the previous fall periods.
This was
required to establish new grass seedings prior to 1970, but was continued in
1970 to provide ade::Juatesoil moisture for the early 1971 growing period.

Height
Nl'st!'l
t u n

l

t Y

Surrounding

Nests

found prior to windrowing

and found in uncut plots provided

1'0'-

lu- I ~ht

ob

I n

l n

l n~

'1'111"., I 11Mlr. IIV(~I"I'f!.1'
t Lvv ho l gh t 01
1').'14

Itrs t

of Vegetation

v(~~('lnl
IwllI,htH

I

VI'

Hlld

l nc he a Willi

l1ll'tlFlllrelll('nI:R

'1fltwlclH

Iounrl

of

s u r

r

ounrl I ng

VI'II,,,ltlllon.

au r round f ng

IB ne e ts

an oppor7l~ no s t s •

1\11 IIVI'I'I1II,O

Found

V(1J.1;(llll-

du r l ng

LII ••

search period of June 4 and 5, 1970, compared with uve rag« vege
Lutlon heights of over 20 inches for the second and third nest searches.
I\n
ovo ru Ll average height of 19.41 inches was recorded for vegetation surroundt ng till' 7/. I1I'Rt~~rncn s u r od .
nest:

&gt;

�-22-

Table 6.

Average height of vegetation in pheasant nesting plots, 1970.

Plot
Number

Vegetative Type
and Treatment

4-15-70

1

Smooth bromegrass (mowed)

4.0

10.0

23.0

37.0

1

Smooth bromegrass (seeded)

4.0

12.0

12.0

10.0

2

Alfalfa (mowed)
New growth
Residual

2.0
4.0

9.0

21.0

26.0

3

Crested wheatgrass (mowed)

4.0

9.0

11.0

17.0

3

Crested wheatgrass (unmowed)
New growth
Residual

4.0
4.0

8.0

9.0

14.0

Intermediate wheatgrass
(mowed)

5.0

9.0

14.0

38.0

Intermediate wheatgrass
(unmowed)
New growth
Residual

5.0
3.0

8.0

16.0

36.0

5

Tall wheatgrass (mowed)

6.0

12.0

14.0

42.0

5

Tall wheatgrass (seeded)

2.0

11.0

10.0

14.0

6

Alfalfa-crested wheatgrass
(mowed)
New growth alfalfa
Crested wheatgrass
Residual alfalfa

1.0
4.0
3.0

10.0
9.0

18.0
13.0

31.0
18.0

Alfalfa-crested wheatgrass
(unmowed) !l
New growth alfalfa
Crested wheatgrass
Residual alfalfa

1.0
3.0
3.0

3.0
8.0

9.0
9.0

20.0
16.0

4

4

6

Average Height (Inches)
5-11-70
6-12-70
7-16-70

1/ New growth alfalfa was well clipped by mice in unmowed alfalfa-crested wheatgrass plots during 1970 growing season.

�-23-

Table 7.

Heights and composition of vegetation surrounding 74 nests, 1970.

Nest
Number

Quadrat

Plot
No.

Date
Measured

1
2
3
4
5
6

SE
SE
SE
SE
SE
SE

1-1
1-1
4-E
2-E
2-E
6-E

6-4-70
6-4-70
6-4-79
6-4-70
6-4-70
6-4-70

7
8
9

SW
SW
SW

5-E
5-E
6-E

6-4-70
6-4-70
6-4-70

10

NE

5-1

6-5-70

11

NE

6-E

6-5-70

12
13

NE
NE

1-1
5-1

6-5-70
6-5-70

14
15

NW
NW

2-1
5-1

6-5-70
6-5-70

16

NW

4-1

6-5-70

17

NW

1-E

6-5-70

18

NW

6-E

6-5-70

19
20

SE
SE

5-1
4-1

6-22-70
6-23-70

21

SE

4-1

6-23-70

22
23
24
25
26
27

SE
SE
SE
SE
S8

3-1
2-1
2-1
4-E
4-8
5-E

6-23-70
6-24-70
6-24-70
6-25-70
6-25-70
6-25-70

28

SE

29
'\0

81':

5-E
6-E

6-25-70
6-25-70

~I':

(,-'1')-70

II

nit;

()-I':
(,-I':

Composition

Ave. Ht.
(In.)

Smooth bromegrass
Smooth bromegrass
Intermediate wheatgrass
Alfalfa
Alfalfa
Alfalfa
Crested wheatgrass
Tall wheatgrass
Tall wheatgrass
Alfalfa
Crested wheatgrass
Tall wheatgrass
Cheatgrass
Alfalfa
Crested wheatgrass
Smooth bromegrass
Tall wheatgrass
Cheatgrass
Alfalfa
Cheatgrass
Thistle
Intermediate wheatgrass
Hairy vetch
Smooth bromegrass
Cheatgrass
Alfalfa
Crested wheatgrass

13.0
14.3
15.0
23.0
22.5
16.3
12.0
16.3
15.3
16.5
12.0
23.0
27.0
15.3
11.0
10.3
13.0
11.3
19.8
14.8
9.5
15.0
12.0
15.0
14.0
15.0
12.0

Cheatgrass
Intermediate wheatgrass
Cheatgrass
Intermediate wheatgrass
Cheatgrass
Crested wheatgrass
Alfalfa
Alfalfa
Intermediate wheatgrass
Intermediate wheatgrass
Tall wheatgrass
Cheatgrass
Cheatgrass

13 .0
24.0
12.7
26.8
13 .3
14.3
24.0
25.5
24.3
23.5
29.5
21.5
20.0
21.5

Ave. Ht.
for
Period (In.)

15.34

SE

(,-,/'j-10

Alfalfa
Alfalfn
A 1 III 1 I'll
Cn'Hl(~d

77.3
,/11 • ')
wlwntKrllHH

/.1 .1

----------------------------------------------------------------------------------

�-24-

Table 7. Heights and composition of vegetation surrounding 74 nests, 1970.
(continued) •

Nest
Number

Quadrat

Plot
No.

Date
Measured

32

SW

5-E

6-25-70

33

SW

6-E

6-26-70

34
35

SW
NW

4-1
4-1

6-26-70
6-29-70

36

NW

4-1

6-29-70

37

NW

5-E

6-29-70

38
39

NW
NW

2-E
6-E

6-29-70
6-30-70

40

NW

2-1

6-30-70

41

NW

6-E

6-30-70

42
43
44
45

NW
NW
NE
NE

5-1
5-1
1-1
5-1

6-30-70
6-30-70
6-30-70
7-2-70

46
47
48

NE
NE
NE

3-E
3-E
4-E

7-2-70
7-2-70
7-3-70

49

SE

6-E

7-20-70

50

SE

6-E

7-20-70

51

SE

5-E

7-20-70

52

SW

5-E

7-21-70

53
54
55
56
57

SW
SW
SW
SW
SW

1-E
4-E
4-E
4-1
2-E

7-21-70
7-21-70
7-21-70
7-21-70
7-21-70

58

SE

3-E

7-21-70

Composition

Ave. Ht.
(In.)

Tall wheatgrass
Cheatgrass
Alfalfa
Crested wheatgrass
Intermediate wheatgrass
Intermediate wheatgrass
Hairy vetch
Intermediate wheatgrass
Hairy vetch
Cheatgrass
Tall wheatgrass
Cheatgrass
Alfalfa
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Cheatgrass
Cheatgrass
Smooth bromegrass
Tall wheatgrass
Cheatgrass
Crested wheatgrass
Crested wheatgrass
Intermediate wheatgrass

29.0
16.0
25.5
21.0
33.0
26.0
8.3
31.3
16.3
16.0
32.7
20.0
19.5
23.0
16.5
17 .5
21.5
23.3
18.0
17.0
15.8
17.5
21.5
14.0
11.3
14.5
25.5

Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Tall wheatgrass
Winter wheat (volunteer)
Tall wheatgrass
Cheatgrass
Smooth bromegrass
Intermediate wheatgrass
Intermediate wheatgrass
Intermediate wheatgrass
Alfalfa
Crested wheatgrass
Crested wheatgrass

18.7
15.0
19.3
14.3
32.7
36.0
20.5
15.0
37.0
34.5
39.0
36.0
19.0
12.0
15.0

Ave. Ht.
for
Period (In.)

20.98

----------------------------------------------------------------------------------

�Table 7. Heights and composition of vegetation surrounding 74 nests, 1970.
(continued) •

Quadrat

Plot
No.

Date
Measured

59

NW

4-E

7-22-70

60

NW

4-E

7-22-70

61
62

NW
NW

2-E
4-1

7-22-70
7-22-70

63

NW

6-1

7-22-70

64

NW

I-I

7-22-70

65
66
67

NW
NW
NW

2-1
2-1
6-E

7-22-70
7-22-70
7-22-70

68

NE

2-E

7-23-70

69

NE

6-E

7-23-70

70

NE

4-1

7-23-70

71

NE

I-I

7-23-70

72

NE

5-E

7-23-70

73
74

NE
SE

5-1
2-E

7-23-70
7-23-70

Nest
Number

Composition

Ave. Ht.
(In.)

Intermediate wheatgrass
Cheatgrass
Intermediate wheatgrass
Cheatgrass
Alfalfa
Hairy vetch
Cheatgrass
Alfalfa
Crested wheatgrass
Smooth bromegrass
Cheatgrass
Alfalfa
Alfalfa
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Alfalfa
Crested wheatgrass
Intermediate wheatgrass
Cheatgrass
Smooth bromegrass
Cheatgrass
Tall wheatgrass
Cheatgrass
Tall wheatgrass
Alfalfa

26.0
lO.7
34.7
14.0
14.5
9.5
lO.O
19.0
14.0
6.0
14.0
23.8
13.0
17.5
14.0
18.5
17.3
27.5
18.0
33.0
14.0
14.5
14.7
14.0
19.0
45.0
17.0

Ave. Ht.
for
Period (In.)

20.40
Average Height All Measurements

19.41

Stocking of Pheasants in Quadrats for 1971
1\ ml n I.mumof

13 hens and 2 cocks, wild-trapped in Phillips, Sedgwick and

Lllrllller cotlntiel'!
1n January, 1971, were wing clipped, 1ep;banded, and

rc l onscd In each of the 4 quadrats by January 28, 1971 for con tt.nuat Lon of
the 8 tudy in 1971 (Segment 25).

�-26-

KEY TO ABBREVIATIONS
Figures 1 Through 4 and Tables 3 Through 5
Plants
Symbol

Scientific Name

Agcr

Agropyron cristatum

Crested wheatgrass

Agel

~gropyron elongatum

Tall wheatgrass

Agin

Agropyron intermedium

Intermediate wheatgrass

Brin

Bromus inermis

Smooth bromegrass

Mesa

Medicago sativa

Alfalfa

Connnon Name

Treatment
Following 1969 nesting season
(m)

Mowed only

(rr)

Reworked and reseeded

(urn)

Unmowed - residual cover check in 1970

LITERATURE CITED
Sandfort, W. W. 1965. Aging pheasant embryos. Colo. Game, Fish and Parks
Dept. Outdoor Facts, Game Information Leaf1. No. 15. 2 p.

Prepared by

Ll~WJ~~
Donald M. Hoffma~
Wildlife Researcher

�-27April, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RAD~~O~·_

Project No.

W-37-R-24

Game Bird Survey

Work Plan No.

1

Job Title

Pheasant Roadside Cover Evaluation Study

Period Covered:
Personnel:

Job No.

18

April 1, 1970 to March 31, 1971

Dale W. Stahlecker and Warren D. Snyder.

ABSTRACT

Roadside seeding efforts to establish grass and grass-legume plots were
continued in 1970. Satisfactory stands were attained on twenty additional
plots, which will be included with their controls in the 1971 evaluation.
Nest searches through the summer measured nesting attempts and successes
on 48 seeded plots, 28 farmed controls and 23 unfarmed natural controls.
TWenty-two of the 29 nest sites were found in seeded plots, two were along
farmed roadside shoulders and five were in unfarmed controls. However,
most nests in seeded plots were found along the unseeded roadside edges
where green and residual weed cover dominated. Mammalian predators
destroyed 16 of the 29 nests, four were hatched, four were abandoned because of human disturbance and five could not be accurately classified.
Accumulations of residual grass and legume cover will potentially enhance
the test sites for nesting pheasants during successive years. Partial evaluation of cover conditions and nesting in older roadsides, previously seeded
by farmers, indicate new species should be tested to provide improved nesting cover.

��-29-

PHEASANT

ROADSIDE
Warren

COVER EVALUATION

STUDY

D. Snyder

P. S. OBJECTIVE
To compare pheasant production under natural roadside conditions with the
following cover types to be established along roads.Ldes: (1) grass and
(2) grass-legume mi.xtures.

SEGMENT
1.

To establish

2.

To measure

OBJECTIVES

grass and grass-legume
pheasant

production

cover on roadside

in test and control

test areas.

areas.

METHODS "AND MATERIALS
The reader is referred to Snyder (1969 and 1970) for methods
used in selection of roadside test areas and in establishing
grass-legume cover.

Measurement

of Pheasant

Production

in Test and Control

and materials
grass and

Plots

Nesting by pheasants in the seeded and control roadside plots was measured
by nest searches.
Dale Stahlecker, student assistant, began searches in
mid-June and continued through the summer.
Linear strips, in which width
varied in relation to cover density, were searched on foot. A stick was
used to part the vegetation for close, methodical inspection.
Nests, where
the hen was laying or incubating eggs, were marked by placing a lath nearby along the road shoulder.
These were checked again at later dates to
determine the fate of the nest.
All plots initially searched before August
I were searched a second time during the summer.
Characteristics
of the
nest site and nest were recorded, and shell fragments and nest bowls of
terminated nests were removed.
Pertinent items recorded at each nest included: date; group and plot location; location within plot; plot cover
type; cover type, height and density immediately surround the nest; status
of nest (laying, incubating, number of eggs, etc.); and fate of nest after
rE'ci1pckA.

�-30-

RESULTS AND DISCUSSION

Establishment

of Grass and Grass-Legume

Cover

Past Seedings
Marginal to satisfactory stands of grass and alfalfa were established
following seeding efforts in 1969 on a majority of the roadside study
plots.
Most of the plots, seeded during the spring of 1969, where seed
competition was severe in summer, produced disappointing stands.
Several
of these were reseeded again in 1969, others were replanted in 1970, and
a few were included in the first year evaluation.
Reseeding will be renewed on a few in 1971 attempting to increase grass and legume densities.
Fair to good grass stands, utilized in the 1970 nest searches, were
obtained west of Holyoke following August, 1969 rains.
Table 1 summarizes
the groups of plots utilized in 1970 nest searches.
Each group contained
either one or two seeded plots and one farmed control.
An unfarmed control was included in most groups.

Seeding

Efforts

in 1970

Approximately
twenty additional study plots will be available for nest
searching in 1971 due to seeding efforts in 1970. These were established
despite poor planting conditions.
Early spring seeding efforts were stymied
by wet soil conditions which prevented working the plots.
Several plots
prepared and seeded in early May produced suitable grass and legume stands.
Insects, destroyed one half-mile stand of grass, and one farmer's summer
employee destroyed another plot which was successfully reseeded.
Weed
competition and hot, dry summer weather reduced stand densities on some
spring seeded plots.
Only one inch of rain was received during the three-month period of midJune to mid-September.
Needless to say, seedings attempted during this
period did not germinate.
But, satisfactory grass stands were obtained
on several plots following September rains. Many sprouting alfalfa seedlings
in these plots were killed by frost while in the two-leaf stage.

Problems

in Establishment

of Grass and Legume

Stands

The primary problem in obtaining seeded stands was to place the seed in the
ground prior to receiving moisture that would germinate and sustain the
stand.
Firm soil was needed but hard rains frequently crusted soil preventing
sprouts from getting through.
Adequate moisture, too often, was followed by
hot-winds which dried the soil desiccating sprouting plants.
Weeds along
steep road shoulders also sapped moisture needed by seeds and plants along
the road edges.
Competitive weeds or loose soil along the field bordering
edge of the strips hampered seed germination.
Spring seedings were conSistently hampered by summer weed growth.
Efforts to suppress weed growth
by mowing resulted in distrubance and possible mortality of newly-hatched

�-31-

pheasant broods which would not leave the dense cover.
In two instances
pheasants attempted to nest in spring-seeded roadsides.
One of these
was killed on the nest while mowing.
In general, the best stands were
obtained from plots sunnner-fallowed and seeded in late July or August.

Measurement

of Pheasant

Production

in Seeded and Control

Plots

Tables 1 and 2 sunnnarize the results of nest searching on 48 seeded plots
and 51 control plots in 1970. The total number of nesting attempts, 29,
was considered low. Twenty-two of these occurred within seeded plots but
only seven hens actually utilized seeded grass or alfalfa.
Most of the
remaining 14 nests in the seeded plots were associated with various combinations of weeds and/or cheat grass cover.
Frequently, nests were
located at the base of unseeded road shoulders where residual weed cover
provided a protective canopy.
Kochia, cheat grass, wild lettuce and sunflowers were the most connnon annuals along unseeded or unfarmed portions
of all roadside plots.
Five nests located in the unfarmed (C2) controls were all associated with
annual forbs as were two nests along road shoulders in farmed (Cl) controls.
None were observed in grain stubble or fallow portions of the farmed cont ro Ls ,
Preliminary nest location results indicate residual cover was an important
attraction factor in nest site selection during the early spring.
Increased
use of green vegetation by hens was observed in late spring - sunnner nesting
efforts.
Most seeded stands lacked residual cover in 1970, therefore, they
may become more attractive to pheasants in future years.
Unfortunately,
two half-mile strips of seeded roadside were burned by farmers during the
early spring of 1971, destroying all residual cover.
Skunks, badgers and other unknown predators destroyed 16 of the 29 nests
located in the study plots.
Crushed and scattered shell fragments were
the main characteristic of destroyed nests.
Nest bowls were occasionally
torn apart.
In one instance, not included in this sample, pheasant remains
and shell fragments indicated that a hen was captured on the nest.
Past
observations have shown that skunks spend considerable time working roadsides in spring.
Undoubtedly, they are the primary predator affecting
roadside nesting success.
Four nests hatched successfully.
Nest search efforts, which flushed several
hens from their neets, were partially responsible for nest abandonment.
Overzealous checks to determine If hens had returned to their nest:.•increased
nb andontuon t • More rca t rnLnt will
be placed on follow-up nest checks in the
future.
Eggs were collected from abandoned nests to monitor for mercury and
pesticide residues.
Treatment of roadsides in 1970 by county maintenance personnal included mowing
along the upper edge of the road shoulder in mid and late summer to keep weed
cover below the level of the road surface.
Seeded stands were not mowed.
Few
if any of the unfarmed controls were mowed beyond the road shoulder.
Most of
th(' shoulder cover along the farmed controls which usually was only two to slx
foo'
In wIdth
WIlH
e l f m l nn t e d by 1'1111.
A few apo t a cont n l nt ng bind
weed,
Cnn ad I an t hLn tl.e , or poverty weed were sprayed by t.he county.

�Table 1.

Roadside treatment and control study plots nest searcheddtiriiigthe

Group Code_
!=l.o

.c

Q)

(/)

§

s::

or!

1-4

~

Z

E-i

1
2
1
2
1
1
2
1
2
1
1
3
4
5
6
7
8
9
10
12
13
14
1
1
2
1
2
1

7
7
6
6
7
8
8
10
10
11

7
7
7
7
7
7
7
7
7
7
7

7
6
7
7
7
7
6

Seeded Plot 1,.f

Q)

eo

!
42 (43)
42 (43)if
44
44
44 J)
44 J)
44
44 !if
44 if
44 (43)if
45 if
45 2:/
45 Jj
45
45
45
45
45
45 !if

45 (46)
45 (46)
45 !if
46 J)

46
46
47 i/
47 !if

48 4/

s

or!

or!

.j.J

.j.J

1-4
Q)

CJ

III
CJ

Q)
C/)

~.

13
24
11
11
21
21
28
12
28
30
12
15
15
9
8
17
17
7
7
18
18
18
35
13
14
9
9
17

0

:&gt;
0

u

s::

s::

0

0

or!

(/)
.j.J

or!

.j.J

.j.J

1-4
Q)

(/)

CJ

Q)

Q)
C/)

III
CJ

Z

Control Type 1
(Farmed~

Seeded Plot
s::

0

0

~

:&gt;
0

u

Sl1nimerof 1970.

s::

(/)
.j.J

or!

0

.j.J

.j.J

(/)

CJ

III
CJ

Q)

Q)
C/)

~

Z

21/g1
---------2
g 1 P
---------_.
7
gl 1 P
---------6
g
11 6 gl
8
21 8 gl
8
5
g
21 5 g
1 Pa,l A
6
g
28 6 gl
5
g
---------_.
5
g
----------6
g
30 6 g
7
g
12 7 8
g
4
15 3 gl 1 P
7
g
15 7 gl
5
8 1 U
---------3
g
8 3 gl
8
81 1 P, 1 A 17 8 gl 1 P
6
17 6 g
8 1 U
2
7 2 gl
8 1 S
3
g
7 3 g
5
18 5 gl 1 A, 1 P
8
6
g
18 6 gl
7
g
18 7 g
1
g
35 1 gl
8
gl 1 P, 1 U 13 8 gl 2 P
7
81 1 S
14 7 gl 2 P, 1 S
3
g
9 3 g
1 S
2
g
---------_ ..
1
---------_ ....
s

if

18
19
11
10
16
20
28
11
21
30
1
21
14
4
8
8
17
8
8
13
13

s::

0

or!

0

5
5
8
1
3
3
5
3
3
5
4
8
6
4
4
4
5
5
5
2
1
5
8
7
3

Control Type 2
(Unfarmed)
s::
0

0

or!

(/)
.j.J

or!

.j.J

.j.J

(/)

CJ

III
CJ

Q)

Q)
C/)

z

0

~

30
24

7
2

10
21
20
29
12

1
1
2
1
6

-------

1 U

-------

-------

35
13
10
15
15

7

16

6

-------

7

(/)
Q)

z

-------

15
15
9
8
17
17
17
7
18
18
18
26
8
14
10
10

7

(/)
.j.J

1 U

1 P

4
7
5
2
1
7
7
4
4
4
8
3
4
6
4
4

I

w

I'.)
I

1 A

2 Pa
1 P

Each seeded plot represents 1f4 mile of roadside.
~/ Seeded stand marginal to poor.
1/ Each 1/2 mile of section periphery was numbered in consecutive order, starting at the northeast corner and
running clockwise around the section to locate the study plot. Letter abbreciations are presented as follows:
g = grass, 1 = legume, A = abandoned nest, P = predator destroyed nest, Pa = predation after search, U =
undetermined nest fate, and S = successful nest.
~/ Grass stands seeded in previous years by resident farmers.

�-33-

Table 2. Characteristics
of pheasant
plots during the summer of 1970.

Part 1.

Fate of 29 pheasant

Successful

nests

nesting

attempts

in roadside

found along roadsides

4

--------------------------------------

Predator

destroyed

prior to search--------------

13

Predator

destroyed

after search ----------------

3

Abandoned

(primarily

Unknown

Part 2.

due to human

of nest attempts

Plot Type

Seeded

roadside

Farmed roadside

plots
controls
controls

(Cl)

Unfarmed

roadside

Part 3.

Cover type locations

Seeded

4

disturbance)--

5

----------------------------------------

Plot location

(C2)

and successes

No. Plots

Nest Attempts

Nests/Plot

48

22

0.46

28

2

0.07

23

5

0.22

of nest attempts

and successes

Cover Type

Number

grass dominant

4

Seeded alfalfa

dominant

Mixture of green and residual
cheat grass

3
weeds

study

or

22

�-34-

Characteristics

of Old Grass Stands for Pheasant

Production

Several plots, seeded by farmers in previous years, were utilized in the
study (Table 1). Two of these contained a western-crested
wheatgrass
combination; two contained dense stands of smooth brome; ten were composed of intermediate wheatgrass, and one contained Russian wild rye.
Most possessed near pure stands of the species planted and were attractive
in general appearance.
Pheasant production on these plots was not encouraging.
One successfully
hatched nest was located in an open stand of intermediate wheatgrass.
One
predator-destroyed
nest was found under a canopy of Russian wild rye.
Empirical evaluation indicated the brome, Russian wild rye and the crestedwestern wheatgrass combination generally did not possess adequate heights
to attract pheasants although densities were suitable.
In contrast, intermediate wheatgrass did not provide adequately dense foliage after becoming
root-bound.
The above mentioned grasses, like most species tend to decrease in height as
they increase in density and competition for moisture over a period of years.
Therefore, new stands of crested or western wheatgrass and brome may provide
attractive nesting cover only for two or three years after they are planted.
Intermediate wheatgrass may consistently produce adequate heights, but leaf
foliage decreases as plant density increases.
These and other species also
lodge under winter snows, diminishing their value for winter cover and spring
nesting cover (Fig. 1). Tall wheatgrass, which possesses a stiff strong
stem, does not lodge as readily (Fig. 2). It may prove to be a desirable
species where it will not hold snow to block roads.
Alfalfa, by itself,
provides favorable spring nesting cover, but loses its foliage in late
summer and is flattened by winter snow (Fig. 3). Present observations indicate that alfalfa-tall wheat mixtures produce a good cover combination,
but more evaluation is needed.
Considerable variation in soil and moisture conditions exits along Northeast
Colorado roadways.
Therefore, the same species may provide poor to excellent
nesting cover depending on the site. Roadsides along newly graded roads
lack humus and other nutrients characteristic of productive soils.
Germination and growth of seeded plants are reduced, so it may be preferable
to let weeds grow for a year to two before seeding newly elevated roadways.
Early spring fertilization could also be used advantageously here.

Loss of Roadside
While this study progresses, many roadside nesting and survival areas continue to be eliminated by back sloping of grader ditches and farming to the
road shoulder.
This occurs on old roadways as well as newly elevated ones.
Figure 4 illustrates elimination of roadside on a newly elevated road where,
in the foreground, wheat has already been seeded.
The stubble field will
be worked to the road shoulder in spring.
Most county roads in Phillips and
Sedgwick counties do not contain any roadside cover.

�-35-

Fig. 1. An old naturally reseeded roadside. Snows have lodged grass in
the grader ditch, however, weeds along the fence provide limited winter
cover for pheasants.

�-36-

Fig. 2. Tall wheatgrass holding snow along a seeded roadside.
Little
winter cover for pheasants is available to pheasants under these conditions.

�-37-

Fig. 3. Alfalfa along a roadside provided cover during the fall of 1970
before snows flattened it.

�-38-

Fig. 4. A newly elevated road showing back-sloping of the grader ditch
in preparation for farming to the road shoulder.
Pheasants have no place
to live or nest where this condition exists.

�-39-

In Phillips County, phone lines are presently being put underground,
eliminating roadside poles and providing additional areas for farmers
to eliminate roadside cover. In some instances poles bordering seeded
areas already have been eliminated. Hopefully, these plots can be
retained for use during the remainder of the study.

LITERATURE

CITED

Snyder, W. D. 1969. Pheasant roadside cover evaluation study. Colo.
Div. of Game, Fish and Parks. Game Research Report. April.
pp , 37-45.
1970. Pheasant roadside cover evaluation study. Colo. Div.
of Game, Fish and Parks. Game Research Report. April. pp.
109-114.

Prepared

by

U.r1t.,J
Warren D. Snyder
~,.)

Wildlife Researcher

��-41April, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RA~D~O~
__~----

Project No.

W-37-R-24

Work Plan No.
Job Title

Period Covered:
Personnel:

Game Bird Survey

3

Job No.
8a
Effects of Sagebrush Control
on Distribution and Abundance of Sage Grouse

April 1, 1970 to September 30, 1970

Warren Snyder, Larry Roper, Terry May, Bruce Sigler, Jerry
Whittaker, George Bock, David Croonquist, Don Gore, Courtney
Crawford, John Monarch and Howard Funk.

ABSTRACT
Counts of strutting grounds on the North Park study area showed peak numbers
of male sage grouse present about mid-April, 1970, about average for strutting
ground activity for the area. Numbers present peaked at 274 males, significantly below 1969, but otherwise the highest total since 1964. Data from check
stations operated at Cowdrey and Walden indicated hunting pressure was again
high with 564 hunters checked. However, harvest dropped to about half of the
1969 total with only 266 birds brought through the stations. Immature birds
made up 45 percent of the bag, an increase over the 30 percent of 1969, and
about average for the period since 1955 when check station activity was initiated in North Park.

��-43-

EFFECTS OF SAGEBRUSH CONTROL
ON DISTRIBUTION AND ABUNDANCE OF SAGE GROUSE
Howard D. Funk

An investigation of sage grouse abundance and distribution near Lake John
in North Park has been in effect since initiation in 1963. The job relates
to various specific study topics in various degrees of intensity.
More
recent study has been with regard to the immediate effects of a sagebrush
spraying project, accomplished in 1965, on sage grouse populations with a
study on the long-term effects of this spray project scheduled for a twoyear period beginning the spring of 1973. In the interim period, efforts
are to be limited to collection of study area strutting ground data,
banding samples of birds on these grounds, and operating check stations
near Walden and Cowdrey to obtain harvest data.
This report covers these
items.

P

S. OBJECTIVE

To determine the effects of 2,4-D spraying of sagebrush en: (1) sage grouse
abundance, (2) sage grouse distribution, and (3) vegetative composition and
density.

SEGMENT
the seasonal

OBJECTIVES

1.

To investigate
is controlled.

distribution

2.

To investigate sage grouse numbers
to the area treated.

of sage grouse after sagebrush

by age and sex classes

METHODS

AND MATERIALS

Strutting

Ground Counts

on and adjacent

Wildlife Conservation Officers conduct counts on a number of strutting
grounds in North Park annually.
Usually, three early morning counts are
made on each ground.
Data from these counts are gathered from the Regional
Biologist, high counts for each ground being reported herein.

Check Stations
Two check stations near Walden and Cowdrey are operated the first two days
of the season to collect various data on harvest such as harvest success,

�-44-

age, and sex ratios, banded bird information, and hunter participation for
a check on the year in comparison with previous years.
Project personnel
are responsible for the station with Conservation Officers also contributing
data from field checks.
Main emphasis is on checking harvest within the
study area.
Age and sex of birds are determined by methods described in
previous reports.

RESULTS

AND DISCUSSION

Strutting

Ground Counts

Results of strutting ground counts are displayed in Table 1. Peak counts
occurred about mid-April on most areas, which is about average since strutting
ground activity is subject to snow conditions in the area. Data on peak
numbers of females and total birds were not gathered for this report, but
should be obtained from the field men for continuity of data when preparing
for the final report.

Table

l.

Strutting

Peak counts of male sage grouse on strutting
1/
Ground-

SG

1

0

SG

2

8

SG

3

0

SG

4

SG

1970.

High Count Da te

April

14

66

April

14

5

57

April 21

SG

6

19

April

l3

SG

7

0

SG

8

7

April

14

SG

9

97

April

15

SG

10

16

April

15

SG

11

4

April 8

SG

12

0

Total

1/

Number Males

grounds,

Locations

274

of grounds

shown in previous

reports.

�-45-

Comparative data on annual high counts by strutting ground are presented in
Table 2. Most grounds show a definite decline in 1970 in numbers of males
present with the total males on all grounds about 25 percent below the 1969
total. However, the 1970 total remains the highest count on record, with
the exception of 1969, since 1964 when 337 males were observed. The year
of the indicated "crash" in population numbers was 1965 when males totaled
only 166. This was the year of the spray project, but spraying did not
take place until after the spring counts.

Table 2. Comparison of 1960 through 1970 strutting ground counts of male
sage grouse.
Strutting
Ground

1960

1961

1962

1963

Year
1964 1965

1966

1967

1968

1969

1970

SG

1

10

5

0

0

0

0

0

0

0

0

0

SG

2

43

4

7

20

26

9

9

10

2

8

8

SG

3

14

0

0

0

0

0

0

0

0

0

0

SG

4

130

96

219

216

120

53

47

69

58

82

66

SG

5

19

2

52

17

65

52

54

59

52

82

57

SG

6

5

1

8

9

3

0

2

12

36

36

19

SG

7

0

7

5

0

0

0

0

1

3

2

0

SG

8

5

0

0

0

0

o·

6

5

2

6

7

SG

9

167

109

71

85

99

52

97

81

71

117

97

SG

10

17

77

28

19

17

0

6

9

0

16

16

SG

11

12

50

5

11

7

0

13

15

6

5

4

SG

12

14

12

0

0

0

0

0

0

1

3

0

446

363

395

377

337

166

234

265

231

357

274

Total

Banding
Due to lack of manpower and available time, banding of samples of birds on
strutting grounds was not accomplished during the segment.

�Table 3.

Comparison of North Park sage grouse hunter check information, 1963-1970. 11

Total
Birds

Birds
Per
Hunter

Hours
Hunted
Per Bird

58

506

1.03

4.86

45

41

179

.82

3.49

26

26

51

116

.77

5.40

56

45

71

44

263

.86

4.67

67

42

46

33

267

.89

4.41

Year

Bag
Limit

Hunters
Checked

Hours
Hunted

Adult
Males

1963

3

492

2,460

62

150

42

113

181

1964

2

217

624

25

81

59

28

2

150

626

27

30

49

1966

2

306

1,227

31

116

1967

2

300

1,177

50

127

1965

u

Adult Percent
Females Adults

Juvenile
Males

Juvenile Percent
Females Juveniles

1968

2

546

2,604

80

135

42

156

141

58

512

.94

5.09

1969

2

662

2,936

121

277

70

57

114

30

569

.86

5.16

1970

2

564

2,617

58

89

55

53

66

45

266

.47

9.84

11 Based on Walden and Cowdrey check station data during opening weekends.

II A one day season; hence only a one day check station.

I
.po

0\
I

�-47-

Check Station

Results

Project personnel checked 266 sage grouse through check stations on September
11-12, 1970, the first two days of the season.
This was slightly less than
half the number checked in 1969. Number of hunters also decreased but not
to the extent of the harvest, thus increasing greatly the number of hours
spent per bird in the bag.
Percent immatures in the bag increased to about
45 percent, which was about average over the years.
No valid reason can be given for the sharp decline in harvest in 1970 or
the sequence of events over the past three years.
In 1968, the high count
on grounds was 231 birds, indicated havest was the highest since 1963 at
512 birds, and the bag contained 58 percent young, again the highest since
1963. Count results on the grounds increased to a high of 357 males in
1969, harvest increased to 569 birds, but the harvest contained only 30
percent young.
Then, 1970 count data on grounds produced 274 males, lower
than 1969 but higher than 1968, while harvest dropped to 266 birds but
immatures made up 45 percent of the bag. Adult females were harvested very
heavily in 1969
(48.6 percent of the bag), quite heavily in 1970 with 33.5
percent, and rather lightly in 1968 with only 26.4 percent.
Hunting pressure
was fairly even through these years with 546, 662, and 564 hunters per year
since 1968. Probably a variety of factors have been responsible for this
confusing sequence of data, including poor sampling in one or more areas of
data collection, including check stations.
Only two bands were turned in at check stations (1662-R466 and 1562-R498) as
compared to 23 the previous year.
The former was banded as a subadu1t male
on April 25, 1969 and the latter as an adult male on April 10, 1968, both
on strutting ground 9 (SG9). Both birds were shot north of Lake John the
first weekend of season.

Prepared

by
Howard D. Funk
Section Chief, Small Game Research

��-49-

April, 1971

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~---------

Project No.

Game Bird Survey

W-37-R-24

Work Plan No.

10

Job Title

Study of Hungarian Partridge Adaptability

Period Covered:
Personnel:

Job No.

1

April 1, 1970 to March 31, 1971

Donald M. Hoffman, Bruce C. Sigler, Dale W. Stahlecker, John F. Corey
and Charles E. Brown.

ABSTRACT
Six hundred and seventy four more pen raised Hungarian partridge were released
in Moffat County at 3 sites on April 24, 1970. This brings the total number of
Huns released in Moffat and Routt counties to 1,788 during the period of 19641970
0

Intensive field searches were made by W-37-R Project personnel during the months
of June, August and September, 1970. Only 10 Huns were observed by Project
personnel, these near 2 of the 3 1970 release sites. Some of the released Huns
were seen in the vicinity of all 3 release sites from May through August, 1970
by local residents. Although previously planted areas were searched, no reports
or observations of Huns could be secured, indicating little or no carryover populations and little or no increases in populations from previous years.
One brood of a hen with 2 chicks was reported by a rancher in the Jubb Creek
area in late August, 1970 in the vicinity of one of the 1970 releases. Reports
of Huns being seen distances of 4 and 5 miles from 2 of the 1970 release sites
were secured.

��-51-

HUNGARIAN

PARTRIDGE

ADAPTABILITY

Donald M. Hoffman

P. S. OBJECTIVE
To determine the ability of Hungarian partridge to establish
populations in mixed brushland-wheatland
habitat in selected
Colorado.

SEGMENT

OBJECTIVES

1.

Introduce

Hungarian

partridge.

2.

Determine

presence

and dispersal

METHODS
Field releases of pen-raised
April 24, 1970.

reproducing
sites in

of Huns and success

of previous

plant.

AND Y~TERIALS

stock were made in 3 areas in Moffat

County

on

Intensive field searches were made by Project W-37-R personnel during the
periods June 15-19; August 25-28; and September 14-17, 1970.
In addition,
persons living and working in or near the release sites were interviewed
and observations were recorded.

RESULTS

AND DISCUSSION

On April 24, 1970, 674 additional pen-raised Huns from the Fort Collins
Wildlife Research Station and Little Hills Experiment Station holding pens
were released in 3 areas of Moffat County.
Figure 1 shows a release made on
Johnson Gulch south of Craig and Figure 2 shows general habitat types of
this area.
Table 1 summarizes all field releases made in Colorado since
1964. A total of 1,923 Huns have been released with most (1,774) being
released in Moffat County, 135 in Larimer County, and 14 in Routt County.
Figure 3 shows the map location of the Johnson Gulch release with numbers
of birds released indicated, and Figure 4 shows the map locations of the
Price Creek and Jubb Creek releases, with the numbers of birds released
indicated.
Table 2 summarizes all Hungarian partridge reports and observations for
Moffat County for 1970. All reports and observations during 1970 were from
the vicinities of the April 24, 1970 releases except for a report of a
single Hun approximately 5 miles north of one of these 1970 release sites
and another approximately 4 miles north of another 1970 release site.
Only
10 Huns were observed by Project personnel, all of which were in 2 of 3
release areas for 1970. Photographs of several of these Huns were secured.

�I

VI
N
I

Fig. 1. Hungarian partridge release, John Gulch, south of Craig in Moffat County, April 24, 1970.
(D. Domenick, photo)

�I
I.J1

W
I

Fig. 2. Habitat in the vicinity of the Johnson Gulch release site, Moffat County. Vegetative types are
cultivated wheatlands bordered by big sagebrush, with a few scattered junipers on hillsides. (D. Domenick,
nhnrn)_

�-54Table 1.

Hungarian partridge field releases since 1964.
Number
Released

Source of
Stock

Release Area

Jan. &amp; Feb., 1964

135*

Oregon &amp; Idaho
(wild trapped)

Larimer County, Douglas
Lake Sec.l, T8N, R69W

Sub-total

135

March 28, 1964

40

Idaho
(wild trapped)

Moffat County, Round
Bottom Sec. 34, T6N, R92W

March 23, 1965

124

Oregon
(pen raised)

Moffat County, Round
Bottom Sec. 33, T6N, R92W

April 18, 1967

71

FCWRS
(pen raised)

Moffat County, Round
Bottom Sec. 21, T6N, R92W

April 30 and
May 1, 1968

236

FCWRS
(pen raised)

Moffat County, Round
Bottom Sec. 29, T6N, R92W

April 30, 1968

14

FCWRS
(pen raised)

Routt County, Northeast of
Hayden Sec. 35, T7N, R87W

April 19, 1969

389

FCWRS (374) &amp;
CSU (15)
(pen raised)

Moffat County, Isles
Mountain Sec. 24 &amp; 25, T5N,
R92W

April 19, 1969

240

FCWRS
(pen raised)

Moffat County, Isles Mtn.
Sec. 16, T5N, R92W

April 24, 1970

170

FCWRS
(pen raised)

Moffat County, Johnson Gulch
Sec. 15, T6N, R91W

April 24, 1970

234

FCWRS
(pen raised)

Moffat County, Jubb Creek
Sec. 35, T5N, R93W

April 24, 1970

270

FCWRS
(pen raised)

Moffat County, Price Creek
Sec. 30 &amp; 31, T4N, R95W

Sub-total

1,788

Date

Total

1,923

* An additional 22 birds (11 pairs) were taken to Rocky Ford Experimental
Bird Farm from this shipment. On October 6, 1965, 51 (pen raised) birds were
received from Oregon and were taken to the Fort Collins Wildlife Research
Station for experimental breeding purposes. On February 19, 1970, 13 1969hatched Huns from eggs secured from wild-trapped birds in England were received
from Winchester Farms, Alton, Illinois and were used for experimental breeding
purposes in 1970 and later years.

�R 92W

-55-

R 91W

"

\

It

;.

••

Fig. 3. Location and numbers of Hungarian partridge released, Johnson Gulch,
Moffat County, 1970.

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�-57-

Table 2. Summary of Hungarian partridge reports and observation, Moffat
County, 1970.
Date(s)

Number Huns

Area

Observed By

April, 1970

3 groups of
approx. 8 each

Jubb Creek, vicinity
of release sites

L. Kendall

1

Price Creek Road,
approx. 5 miles N. of
upper Price Creek
release site

May 21, 1970

B. L. M.
Personnel

Late May, 1970

2

Approx. 3 miles S. of
Jubb Creek release
sites

G. Hanson

June 16, 1970

3 (pair &amp;

Jubb Creek, vicinity
of release sites

B. Sigler

Price Creek, vicinity
of release sites

B. Sigler

single)
June 17, 1970

2 (pair)

June 18, 1970

5 (2 pair &amp; 1
single)

Jubb Creek, vicinity
of release sites

B. Sigler

May to early
August, 1970

Few

Price Creek, vicinity
of release sites

Stoddard

Late August, 1970

1 hen with 2
chicks

Jubb Creek, vicinity
of release sites

C. Hebblthwaite

Late August,
1970

Few

Yampa River, approx. 4
miles N. of Upper Jubb
Creek release site

C. Hebblthwaite

May to late
August, 1970

Few

Vicinity of Johnson
Gulch release site

Barnes

�-58-

Although areas of releases made prior to 1970 in Moffat County were searched,
reports and observations for these areas were nil, indicating an absence of
carry-over populations and little if any increase in populations.
One Hungarian partridge brood of a hen with 2 very small checks was reported
by rancher C. Hebblthwaire in the vicinity of the Jubb Creek release site
in late August, 1970.
Period of Release
Specific experiments have not been run to determine the best time for field
releases of Hungarian partridge in Colorado.
Traditionally,
the early
spring period has been selected because ~f the advantages of ample food
conditions with growing green grass and forbs and most of the severe winter
weather periods are past.
A field release time during the last half of April is a compromise between
the near approach of the egg laying season (early May under penned conditions)
and late snows which may occur as late as early May in areas selected for
Hungarian partridge releases.
Birds held in the large holding pens begin
to pair as early as February each year, and most are believed to be paired
by release time. By releasing all birds from a single holding pen at one
site, they have the opportunity to select the same mate upon release.
Handling of the birds is known to cause stress as measured in weight loss
and transporting the birds undoubtedly adds to this stress.
This may delay
the nesting activities of the birds somewhat; however, there should still
be ample time for the birds to establish territories and nest following
release.
Upon release from the crates, most of the Huns stay in a rather loose
group and appear to start pairing almost instantaneously.
Whether they pair
with the same birds as in the large holding pens is not known.
Birds observed
later at the release site or at a distance from the release site are usually
observed in pairs, so there appears to be no problem in the birds finding
mates following release.

Prepared

by

iJ~?r7~~

Donald M. Hoffman
Wildlife Researcher

~

�-59April, 1971

JOB PROGRESS REPORT

State of

...;C:..;O;;,.:L::;:O;.:.RAD=.;:.O
_
Game Bird Survey

W-37-R-24

Project No.

10

Job Title

Experimental Breeding of Hungarian Partridge

Period Covered:
Personnel:

Job No.

2

Work Plan No.

April 1, 1970 to March 31, 1971

Lawrence A. Webster, Robert L. Schmidt: John F. Corey, Donald M.
Hoffman and Dr. Harry D. Muller.

ABSTRACT
A test in which 12 pairs of qungarian partridge breeders fed a standard 22 percent
protein level game bird breeder feed compared with an equal number fed a similar
feed but containing 16 percent protein, indicated the 16 percent protein feed was
significantly better at the 10 percent level in numbers of fertile eggs laid (293
compared with 158). Egg fertility was also higher with the 16 percent protein
feed (86.94 percent compared with 73.49) resulting in higher numbers of chicks
hatched (234 compared with 130). Both levels of protein tested (22 percent and
16 percent) showed excellent results in the 7-day survival tests.
In a test where 6 pairs of early mating pairs were compared with an equal number
of late mated pairs, the late mated pairs produced more fertile eggs (129 compared
with 55) and a slightly higher percent hatch of fertile eggs was found with the
late mated pairs (80.62 percent compared with 78.18).
In another test which compared fertile eggs produced by 3 pairs rece~v~ng 2 extra
hours of artificial light each A.M., 3 pairs receiving 2 extra hours of artificial
light each P.M., and 3 pairs receiving no artificial light, it was found the
artificial lighting of the pens resulted in bringing breeder pairs into production
earlier than those which received no artificial light. However, total egg production and total fertile egg production were not increased with the use of artificial lights.
Production cost records were maintained in 1970. These indicated it costs a
minimum of $9.45 per Hun produced for field release using present methods, facilities, and equipment, including labor costs. If labor costs are excluded, a cost
of $2.20 per bird was calculated.

�-60-

RECOMMENDATIONS
One year's data have been collected in comparing 22 percent protein level
game bird breeder feed with 16 percent. Although results showed that the
16 percent protein feed was better for Hungarian partridge in 1970,
comparisons should be made for at least one more year. Extra lighting
tests started in 1970 should also be continued for at least one more year.
Progeny from highest producing pairs in 1970, Winchester Farms stock,
and the highest producing 1969-hatched adult breeders have been marked and
it is recommended that selection tests be started in 1971.
Numbers of Huns placed in rearing and conditioning pens at the Fort Collins
Wildlife Research Station and at the Rocky Ford Game Farm were recorded in
Segment 24. It is recommended that an accurate count of birds removed by
pen be made and a comparison of overwinter survival by various densities be
made in Segment 25.

�-61-

EXPERIMENTAL BREEDING OF HUNGARIAN PARTRIDGE
Donald M. Hoffman
Even though 36.90 percent fewer eggs were laid by the 52 pairs of Hungarian
partridge breeders in 1970, compared with an equal number of breeders in
1969 (1,147 eggs compared with 1,818), several refinements were made in
incubation procedures and the rearing of young. The 655 pen-raised Huns
released in the field in 1971, plus 20 extra birds held for breeders,
compares favorably with the 674 released in the field in 1970. A total of
614 were released in 1969 and 250 in 1968.
Refinements in procedures included (1) replacing the electric heat lamp
brooders with modified electric Jamesway heatring brooders, (2) eggs were
incubated at a humidity of 85.50 F wet bulb temperature in 1970 in place of
84.50 F used previously, (3) eggs were hatched at a temperature of 99.250 F
in 1970 in place of 99.500 F used previously, and (4) chicks were fed 27%
protein turkey starter crumbles in 1970 for the first seven days, 30% turkey
starter crumbles for the following two to four weeks, and 26% turkey grower
crumbles from the fourth week, with improved results. In 1969, 28% protein
game bird starter mash was fed for the first four weeks, followed by 26%
game bird grower crumbles and pellets.
P. S. OBJECTIVE
To develop game farm production techniques for Hungarian partridge.
SEGMENT OBJECTIVES
1.

To measure the effects of 2 levels of protein in the breeder feed
(16% versus 22%) on :
(a) Egg production
(b) Egg fertility
(c) Egg hatchability
(d) Chick survival for the first 7 days.

2.

To compare the numbers of fertile eggs produced by early mating pairs
with those mating late in the breeding season.

3.

To compare the numbers of fertile eggs produced by breeders subjected
to two extra hours of light each 24 hour period during (1) the early
morning and (2) the evening, with breeders receiving no artificial light.

4.

To determine production costs.

�-62-

METHODS

AND MATERIALS

The detailed arrangement of the Hungarian partridge breeding pens for 1969
is shown in Figure 1. Mated pairs for all except the late mated series
were selected and caught in the large Hungarian partridge holding pens
(Fig. 2). Table 1 lists the pen schedule for the various tests run in 1970.
One mated pair of Hungarian partridge was used in each pen.
Pens 1 through
36 are located in a converted nursery shadehouse with ground floors, and pens
37 through 51 are a separate series of wire floored experimental game bird
breeding pens.
Pens A-I, A-2, and A-3 are alleyway pens in the ground
floored series.
Protein Levels

in Feed

Pens 1-6 and 19-36 (24 pens) were used to test the effect of 2 levels of
protein in the feed. Birds in even numbered pens were fed a standard game
breeders feed containing 22 percent protein and those in odd numbered pens
were fed a similar feed, but containing 16 percent protein.
Eggs were
marked and placed in separate incubator drawers to facilitate determination
of (1) fertile eggs produced; (2) number of fertile eggs that hatched, and
(3) numbers of surviving chicks.
Chicks were held separately in the brooder
until they were 7 days of age to detect differences in survival.
Period of Mating
Pens 7-18 (12 pens) were used to compare numbers of fertile eggs produced
by earliest mating pairs with those mated late in the breeding season.
Even numbered pens were used for early mated pairs and odd numbered pens
for late mated pairs.
Six pairs used for earliest mating pairs were
selected as soon as the birds were observed to start pairing in the large
runs (February 6, 1970) and the 6 pairs used for latest mating pairs were
selected from 10 unpaired hens and 10 unpaired cocks held in separate pens
from February 6, 1970 to March 23, 1970. These began to pair almost
immediately when recombined on March 23, 1970 and pairs were selected later
on the same date.
Extra Lighting

Tests

The wire-floored series of pens are well suited for extra lighting tests
and the necessary wiring and automatic timing devices were installed late
in Segment 23. U. S. Weather Bureau sunrise-sunset tables were used to
determine times when lights should go on and off.
Pens 46-48 (3 pens) were lighted with 2 extra hours in the early morning
and pens 49-51 (3 pens) were lighted with 2 extra hours during the late
evening.
Pens 40-42 (3 pens) were used for controls and were treated
identically except no artificial lights were used.
Originally, pens 37-39
were also listed for controls, but the pair in pen number 39 escaped on
June 2, 1970, so this series of 3 pens was used only in comparing earliness
of egg production.
Records were maintained of dates of first eggs, numbers
of eggs laid, numbers of fertile eggs laid, and hatchability of eggs produced.

�-63-

N

IJ ~
-

CD

75'

6'4"

,

If]

'-

If)
Jf\

Ground-f loored Series

3738 394041

42 43 4445 46 47 4E

~ , v_ '-- 'v _ '---'v _
.'

\ V _'---,

v_ '-- v _
\

..
Wire - floored
CD

v

I-

N

A-2 ~

",'

b

-

0

N

N
(\J

01

N-.!.

-,

/

/

49~

.r:.

N

0

~1
A-3

SCALE 8

L

I

/

en

CD

I

I

0' 5' 10'

I

25'

~
L
/0)

Fig. 1.

U)

Detailed Arrangement

=,

an

5_'~

(11

N,

-

Ser

,

Partridge

/N

J'--'6---1

48' ----·-----······1

I:!

CD

l'" '

Breeding

of

Pins.

Hungarian

i,.

_

....l

�-64Covered
Brooder

Shelter

1

Houses

/~
l.--_-1- __

I'-__

I

..l..._---' _-+----L----.J4't-8'~

f

J
-0
t\J

4'

t

N

-in

o

C\I

C\I
C\I

•.4''i
-L__
~ 15.5
,"'15.5-1
---35
---

~_~

~

---l

~------------I05'-----------~
Fig. 2.
Hungarian

Partridge

Holdin q Pens

�-65Table l.

Pen schedule for Hungarian partridge, 1970.

Protein Level
Pen
Nwnber' 22%
16%

T
T

3
4

8x8

T

1
2

Pen Size ~Ft·2

T
T

5

8x16

x

x

x

x

x

x

x

x
x

x

7

x

x

8

x

x

9

x

x

10

x

x

11

x

x

12

x

x

13

x

x

14

x

x

15

x

x

16

x

x

17

x

x

18

x

x

20

T
T

21
22

T
T

23
24

T

25
26
27

T

T
T

Artificial Light
None
AM
PM

x

T

T

Mated
Late

x

6

19

6x33

Pairs
Early

T

x

T
T

x
x

T
T

x
x

T
T

x
x

T
T

x
x

T
T
T

x

x
x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

-----------------------------------------------------------------------------

�Table 1.

-66Pen schedule for Hungarian partridge, 1970.

Pen
Number

Pen Size ~Ft.2
8x8
8x16
6x33

28

Protein Level
22%
16%
T

29
30

T

T

31
32

T
T

33
34

T
T

35

T

Pairs
Early

(continued) .

Mated
Late

Artificial Light
PM
None

AM

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

36

T

37

x

x

x

38

x

x

x

39

x

x

x

40

x

x

T

41

x

x

T

42

x

x

T

43*

x

x

x

44"k

X

X

x

45"~

x

x

x

46

x

x

T

47

x

x

T

48

x

x

T

49

x

x

T

50

x

x

T

51

x

x

T

A-1"~

x

A-3"/( x
No. Pens
on Test 12
12
0
0
0
6
Key-T - Pen on Test
x - Pen not on Test
,.~Ind icates pens with Winchester stock birds.

x
x
6

3

3

3

�-67-

Production

Costs

Costs were estimated, based upon necessary labor, equipment, and facilities
required to produce a number of Hungarian partridge similar to anticipated
production at the Fort Collins Wildlife Research Station for 1971, using
methods now in use. These costs do not reflect the costs related to research
studies which have. been of considerable value in improving methods and
techniques for producing numbers large enough for adequate field releases.
Winchester

Farms Stock

Although not written as a separate test in 1970, 13 birds (5 hens and 8 cocks)
which were survivors from the 18 Hungarian partridge received from Winchester
Farms in Illinois in February, 1970, afforded the opportunity to test fertile
egg production and hatchability in 1970. These were all July, 1969-hatched
from eggs received at the McGraw Wildlife Preserve from wild Huns in England.
This new blood line will be introduced into our resident flock st.arting in
1971.
The Winchester Farms Huns were held in a separate pen and allowed to pair
naturally.
Pairs were placed in pens 43-45, A-I, and A-3. Records were
maintained of numbers of eggs laid, numbers of fertile eggs, and hatchability
of eggs produced.
Pens 43-45 were compared with pens 40-42 (controls).
Progeny from the first 2 hatches of 1970 were Wing-tagged on the left wing
in preparation for crossing with resident stock.
Because of limited numbers
of these progeny,S
adult cocks were also held over for use in 1971 crosses.

RESULTS

AND DISCUSSION

Protein

Levels

in Feed

Table 2 lists a comparison of numbers of eggs produced and percent hatch of
eggs from breeder pairs in 12 pens using a standard 22 percent protein
level feed with 12 breeder pairs using a similar feed but containing 16
percent protein.
A total of 122 more eggs (337 compared with 215) was laid
by the breeders using the 16 percent protein feed in comparison with the 22
percent protein feed. A total of 135 more fertile eggs (293 compared with
158) was laid by the breeders using the 16 percent protein feed in comparison
with the 22 percent protein feed. This difference is significant at the 10
percent significance level. In addition, 86.94 percent of the eggs laid by
breeders on 16 percent protein feed were fertile compared with 73.49 percent
for the 22 percent protein feed. A total of 234 chicks was hatched from
breeders on the 16 percent protein feed compared with only 130 for the 22
percent protein feed.
Excellent survival for the first 7 days was found for both the 16 and 22
percent protein level feeds (98.26 percent survival for the 16 percent feed
and 97.60 percent survival for the 22 percent feed) as listed in Table 3.
A very high survival rate also continued following the 7 day test, reflecting
the strong, healthy condition of the chicks from both levels of protein in
the feed and improvements in brooding methods used in 1970. The electric
heat lamp brooders were replaced by modified electric Jamesway heatring
brooders in 1970 for better results in both survival and condition of the
young birds.

�Table 2. Numbers of eggs produced and hatchability of eggs laid by Hungarian partridge on two levels of protein in
the feed, 1970.

Protein
Level

Number
Pens

Number
Eggs
Produced

Number
Infertile
Eggs

Number
Fertile
Eggs

Percent
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germ

Number
Culls

16%

12

337

34

293

86.94

234

69.44

79.86

9

50

10

22%

12

215

52

158

73.49

l30

60.47

82.28

2

26

5
I

~
co
I

Table 3.

Survival of Hungarian partridge on two levels of protein in the feed, 1970.
Number Chicks Taken
to Brooder House or
Battery Brooder

Protein
Level

Number
Pens

16%

12

230

22%

40

501

Number Chicks
Lost First Seven
Days

Number Chicks
Survived
Seven Days

Percent Survival
First Seven Days

4

226

98.26

12

489

97.60

�-69-

Based upon this one year's results, the 16 percent protein level appeared to
be considerably better than the 22 percent protein level, for Hungarian
partridge.
It is planned to run similar tests in 1971, but with 6 pens on
16 percent protein level feed and 6 pens on 22 percent protein level feed.

Period of Mating
Table 4 lists a comparison of numbers of fertile eggs produced and percent
hatch of fertile eggs from 6 pens with early mating pairs with 6 pens with
late mating pairs, all using 22 percent protein level feed. Considerably
more fertile eggs were laid by the late mating pairs (129 compared with 55)
and a slightly higher percent hatch of fertile eggs was found with the late
mating pairs (80.62 percent compared with 78.18).
This resulted in 104
chicks hatched from the late mating pairs compared with only 43 for the early
mating pairs.
Three of the 6 pens of early mating Huns laid no eggs compared
with only 1 of the 6 pens of late mating Huns which laid no eggs.
It therefore appears that variability of individual birds played a large part in the
results obtained.
No clear advantage was found in selecting earliest mating
pairs and, in fact, this proved to be disadvantageous
in 1970.

Extra Lighting

Tests

A comparison of numbers of eggs laid, numbers of fertile eggs, hatchability
of eggs, and numbers of chicks hatched by 3 pairs of Hungarian partridge
under artificial lighting for 2 extra hours in the A.M., 3 pairs under 2
extra hours during the P.M., and 3 pairs receiving no artificial lighting
(controls) is listed in Table 5. All pens received 22 percent protein level
feed and all were treated as alike as possible in other respects.
Originally,
the experiment was set up using 6 controls and 6 treated pens but a pair of
Huns in pen number 39 escaped on June 2, 1970 so this series of 3 pens
(Numbers 37-39) was not used except to compare earliness of egg production.
More eggs were produced by 3 pairs receiving no extra lighting or controls
(102) compared with those receiving 2 extra hours in the A. M. (83) and
those receiving 2 extra hours in the P. M. (71). In comparing numbers of
fertile eggs, both the pairs in 3 pens receiving extra morning light and
the 3 pens of controls laid 77 fertile eggs each, while the 3 pens receiving
extra evening light laid only 48 fertile eggs.
Considering hatchability of
fertile eggs, the controls ranked highest with 90.91 percent, followed by
those receiving 2 extra hours during the P. M. (87.50 percent) and those
receiving 2 extra hours during the A. M. (70.13 percent).
Numbers of chicks
hatched were 70 for the controls, 54 for the pens receiving extra morning
light, and 42 for the pens receiving extra evening light. Of the 9 pens
used in this test, the minimum number of eggs laid was 14 from pen Number 46,
(lighted for 2 extra hours in the A. M.), and the maximum number was 51 from
pen Number 42 (a control with no extra lighting).

�Table 4.

A comparison of eggs produced by earliest mated pairs of Hungarian partridge with those mated late, 1970!/.

Treatment

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Early Mated

6

64

Late Mated

6

141

Number
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

6

55

43

67.19

78.18

1

11

3

9

129

104

73.76

80.62

0

25

3

1/ A standard 22% protein game bird breeder feed was used in all pens.

Table 5. A comparison of eggs produced by Hungarian partridge pairs receiving two extra hours of light with those
receiving no extra light. !/
I
....•

0
I

Treatment

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Number
Fertile
Eggs

Two Extra
Hours A. M.

3

83

3

Two Extra
Hours P. M.

3

71

Controls

3

102

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

77

54

65.06

70.l3

5

18

3

23

48

42

59.15

87.50

1

5

0

20

77

70

68.63

90.91

1

6

5

1/ A standard 22% protein game bird breeder feed was used in all pens.

�Table 6.

I.

An estimate of minimum costs involved in producing Hungarian partridge. 1/

Capitol Investment
Breeding Pens - Set of 50 size 8' x 16' each
Brooder Houses - Set of 3 with shelters size 26' x 10'
Holding Pens - Set of 4 size 36' x 222'
Incubator
Brooders, feeders, and waterers
Small tractor with mower
Work facilities with egg storage room

$

7,500.00
3,000.00
10,000.00
2,000.00
500.00
1,000.00
2,500.00
$26,500.00

Sub-total
Forty year life expectancy - Cost per year
II.

Feed and Utilities
Feed for breeders and young per year
Utilities (heat, light, and water) per year

$

600.00
500.00
1,100.00

Sub-total
III.

662.50

$

Labor
Conservation Aide level, 2nd Step, for 5 months;
1 part time for 7 months per year
Retirement, hospitalization, etc. (7%) per year

5,400.00
400.00

Sub-total

5,800.00

TOTAL

$7,562.50

Estimate can produce 800 Huns per year with these facilities and labor.
Estimate of cost per Hun produced

=

$7.562.50
800

=

=

$1,762.50

Estimate of cost per Hun produced, labor excluded

1/ Costs do ~

800

=

$

9.45

s

2.20

include the following:
a. Land investment since value can probably be returned at end of use period plus interest, based
upon present trends.
b. Cost of breeding stock
c. Interest required on capitol investment.
d. Expenses incurred due to unforeseen losses.

I

"-J

I-'
I

�Table 7.

Source

A comparison of eggs produced by Winchester Farms stock Hungarian partridge with resident stock, 1970.

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Number
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

I

-..,J
N
I

Winchester
Farm, Illinois

3

83

26

55

46

55.42

83.67

1

8

2

FCWRS

3

102

20

77

70

68.63

90.91

1

6

5

�-73-

Even though the use of artificial lighting did not increase the total egg
production in 1970, it was found that the lights did bring the breeding
pairs into production earlier than the breeding pairs without artificial
lighting.
On May 15, 1970, all 6 of the lighted pens were in production and
had produced 22 eggs, compared with only 4 of the 6 original controls being
in production and having produced only 13 eggs.
The bringing of breeders
into production early may be of value, particularly when the next year's
breeders are selected from the first hatch of the previous year as is the
case at the Fort Collins Wildlife Research Station.
Because of the small samples possible in 1970, it is planned to rerun this
test for at least one more year, using the same number of pens in each sample.

Production

Costs

Table 6 lists estunates of minimum costs involved in producing Hungarian
partridge based upon estimates of capitol investment, feed and utilities,
and labor required to produce approximately 800 Huns, using procedures now
in use at the Fort Collins Wildlife Research Station.
A cost of $9.45 for
each Hun released in the field was calculated from this cost analysis,
including labor costs.
Most cost estimates on game bird propagation do not
include labor costs, which are the largest single annual cost item.
If
these labor costs are excluded, a cost of $2.20 per bird released in the
field was calculated.
Methods to reduce labor costs of propagating Huns
through rearing of young under semi-natural conditions are being tested.

Winchester

Farms Stock

Table 7 compares numbers of eggs laid, numbers of fertile eggs, hatchability
of eggs, and numbers of chicks hatched from 3 pens of pure Winchester Farms
stock with 3 pens of resident Huns. All birds received 22 percent protein
level feed and were treated as much alike as possible.
The 3 pens of Winchester Farms stock produced fewer numbers of eggs (83
compared with 102), fewer numbers of fertile eggs (55 compared with 77), and
fewer numbers of chicks hatched (46 compared with 70) in comparison with 3
pens of controls (resident stock).
In addition, hatchability of fertile
eggs was lower for the Winchester Farms stock (83.64 percent compared to
90.91) in comparison with the resident stock. Since the Winchester Farms
stock is only one generation removed from the Wild, these results were not
unexpected.
The Winchester Farms stock should be of considerable value in
introducing new bloodlines and may help prevent excessive future inbreeding.

Prepared

by

£~~~

Wildlife

Researcher

��-75April, 1971

JOB PROGRESS REPORT

State of

...;C::;.:O::,:L:;;O;,,;.RAD=;,;::O
_

Project No.

W-37-R-24

Work Plan No.

Game Bird Survey

12
Job No.
--=1.,:.0-,_
Relationships of the Productivity and Distribution of
~W~i~l~d~Tu~r~k~e,y~s~o~n~t~h~e~U~n~c~o~m~p~a=h~g~r=e~P~l~a~t~e~a~u~t~o~t~h~e~
_

Job Title
Period Covered:

April 1, 1970 to March 31, 1971

P.S. OBJECTIVES
1.

To determine the size, distribution, and productivity of the wild turkey
population on the Uncompahgre Plateau.
20 Record movements of wild turkeys and determine how environmental factors
affect these movementso
3. Determine the sex and age structure of the turkey kill on the Uncompahgre
Plateau to provide estimates of total harvest and size of the turkey
population.
40 Determine the vegetative types present on the wild turkey summer and winter
range and the species composition and abundance of food producing species
within each type. Also determine the elevation, describe topography, and
record certain climatic factors on part of the better turkey summer and
winter range.
SEGMENT OBJECTIVE
To compile, analyze, summarize, and publish information.
Progress
Field work was completed on October 20, 1967. All data have been summarized,
and most data have been analyzed. A manuscript containing these data and results
from Work Plan 12, Job 14 is being edited and revised.
Future Plans
The manuscript should be completed in Segment 25.
in the form of a technical bulletin.

c" "

(1

rt1, ~_

Prepared by: &lt;~~
oJ II~
Gary T. Myers
Wildlife Researcher

.'

Publication will probably be

��-77April, 1971

JOB PROGRESS REPORT

State of

C.=..O.=..L;;;,O;;;.;RA=D;;..;O~__:..
_

Project No.

W-37-R-24

Game Bird Survey

12

Work Plan No.
Job Title

Job No.

14

Use of Food Plots to Concentrate Wild Turkeys

Period Covered:

April 1, 1970 to March 31, 1971

P.S. OBJECTIVES
Determine the effects of small food plots on fall concentrations and harvest
of wild turkeys.
SEGMENT OBJECTIVE
To compile, analyze, summarize, and publish information.
Progress
Field work was completed December 2, 1968. Data were analyzed, written up,
and incorporated in a manuscript containing information gathered under W-37-R
Work Plan 12, Job 10. This manuscript is being edited and revised.
Future Plans
The manuscript should be completed in Segment 25.
be in the form of a technical bulletin.

Prepared by:

&lt;.p2"'-~~)
/J",~-f...4..
Gary T .'-yers
Wildlife Researcher

Publication will probably

��-79-

April, 1971

JOB PROGRESS REPORT

State of

~CO~L~O~RA~D~O~

Project No.

W-37-R-24

Work Plan No.
Job Title

Game Bird Survey
Job No.

2

Study of Mountain Quail Adaptability

Period Covered:
Personnel:

15

_

April 1, 1970 to March 31, 1971

Ronald B. Arant, Tom W. Barnes and Donald M. Hoffman.

ABSTRACT
Twenty-five surplus breeders, including eight 1969-hatched and seventeen
1967-hatched or older, were released on the South Fork of Mesa Creek in
Mesa County on June 23, 1970. The release site was approximately .6 mile
below the first crossing of the South Fork of Mesa Creek and approximately
5.7 miles above the last mountain quail sighting location. Birds released
included 14 hens and 11 cocks, which had produced no eggs during 1970.
Field trips to search release sites and surrounding areas on the Uncompahgre
Plateau were made in June and September, 1970. No mountain quail were seen,
no field sign found, and no new reports of sightings of the species were
secured.
Although mountain quail have been most difficult to propagate in captivity,
several problems have been solved. The species holds definite potential
for establishing a new game bird in the extensive areas of mountain shrub
and ponderosa pine zones on both the western and eastern slopes of Colorado.
If experimental propagation efforts are successful in 1971 and following
years at the Fort Collins Wildlife Research Station, it is recommended that
trial introductions be made in the Spanish Peaks area of Las Animas County
starting in 1972.

��-81-

STUDY OF MOUNTAIN

QUAIL ADAPTABILITY

Donald M. Hoffman

P. S. OBJECTIVE
To determine the ability of mountain quail to establish reproducing
populations in areas of mixed shrub and pine in selected sites in Colorado.

SEGMENT

OBJECTIVES

1.

Introduce mountain

quail.

2.

Determine

and dispersal

presence

METHODS

of mountain

quail.

AND MATERIALS

Twenty-five surplus mountain quail breeders from holding pens at the Fort
Collins Wildlife Research Station were banded with aluminum butt end leg
bands, transported to the South Fork of Mesa County in Mesa County in a
State pickup, and released on June 23, 1970.
Two separate field
of the Uncompahgre
living and working
any mountain quail

searches of the release sites and surrounding areas
Plateau were made in June and September, 1970. Individuals
in and near the areas were contacted and asked whether
had been observed.

RESULTS AND DISCUSSION
An observation of 20 mountain quail made by Conservation Aide Tom Barnes of
Nucla, Colorado on the South Fork of Mesa Creek reported in the Job Progress
Report for Segment 23 of this job for early 1970, was actually made in late
December, 1969. These were observed approximately one-half mile above the
old Bogus Mine in a pinon-cedar vegetative community.
Within their native
ranges, mountain quail that summer at higher altitudes frequently migrate
to lower levels in winter.
These were seen only one time, so they may have
moved into this lower area of South Fork of Mesa Creek from some unknown
higher area.
Twenty-five surplus mountain quail breeders, including eight 1969-hatched
and seventeen 1967-hatched or older birds, were released on the South Fork
of Mesa Creek in Mesa County on June 23, 1970. The release site was
approximately
.6 mile below the first creek crossing and approximately 5.7
miles above the last mountain quail sighting.
Birds released included 14
hens and 11 cocks, none of which produced eggs in 1970. Table 1 lists all
field releases of mountain quail and sources of stock being held at the
Fort Collins Wildlife Research Station for experimental propagation purposes.

�Table 1.

Summary of mountain quail field releases.
Number
Birds

Source of
Birds

Release
Area

Aug. 19 and 25, 1965

143*

California
(wild -trapped)

Mesa County, Indian Creek
Sec. 27, TS1N, R17W

Aug. 20, 1966

163

California
(wild-trapped)

Mesa County, Indian Creek
approx. 4 miles west of 1965
releases

Jan. 25, 1968

66

Oregon
(wild -trapped)

Mesa County, Indian Creek
approx. 1 mile east of west
Uncompahgre National Forest
boundary

Date

I

00

June 23, 1970

25

Total

397

FCWRS
(surplus breeders)

Mesa County, South Fork Mesa
Creek, Sec. 17, T49N, R16W

* Sixteen additional mountain quail from these shipments were taken to Fort Collins Wildlife Research
Station for experimental propagation studies. On January 18, 1970, 12 Wild-trapped mountain quail from
Oregon were added to the breeding stock at the Fort Collins Wildlife Research Station. These constitute
the source of our breeding stock.

N
I

�-83-

Field trips to search for mountain quail in the release areas and surrounding
areas on the Uncompahgre Plateau were made in June and September, 1970. No
mountain quail were seen, no field sign found, and no new reports of
sightings were secured from individuals contacted.

Prepared by

J/}~P2~
Donald M. Hoffman
Wildlife Researcher

��-85April, 1971

JOB PROGRESS REPORT

State of

C~O~L~·O~RAD~~O~_

Project No.

W-37-R-24

Work Plan No.
Job Title

Job No.

3

Experimental Breeding of Mountain Quail

Period Covered:
Personnel:

15

Game Bird Survey

April 1, 1970 to March 31, 1971

Lawrence A. Webster, Robert L. Schmidt, John F. Corey and Donald M.
Hoffman.

ABSTRACT

A new set of 10 mountain quail breeding pens was completed on April 13, 1970
and used for the 1970 season, but only one egg, which was infertile, was laid
by the entire group of 32 breeders of mixed sexes placed in the breeding pens
and no eggs were laid by 38 surplus birds of mixed sexes placed in holding
pens. The catching and sorting of the mountain quail breeders on April 14,
1970 apparently stressed these tempermental birds and resulted in almost no
egg production in 1970. This lateness in moving the breeders was due to a
delay in completing the new set of breeding pens.

��-87-

EXPERrnENTAL BREEDING OF MOUNTAIN QUAIL
Donald M. Hoffman

P. S. OBJECTIVE
To develop game fann production techniques for mountain quail.
SEGMENT OBJECTIVE
To measure the relationship of pair mating, selected group mating and flock
mating to fertile egg production.
METHODS AND MATERIALS
A new set of 10 mountain quail breeding pens was completed on April 13,
1970 (Fig. 1). It was planned to have these completed by mid-March but
inclement weather plus a manpower shortage delayed the completion.
Pairs of mountain quail were placed in pen numbers 1, 5, 7, and 9; trios
(2 hens and 1 cock) in pen numbers 2, 4, 6, and 10; and a community pen of
8 hens and 4 cocks was placed in the combined pen numbers 3, 8, and alleyway, on April 14, 1970. All birds were fed a standard 22 percent protein
level game bird breeder feed and all were 1969-hatched birds. Numbers of
fertile eggs produced were to be recorded by individual pens.
Ten wild-trapped mountain quail of mixed sexes received from Oregon in
Segment 23 wereplaced in combined holding pen Numbers 1 and 2, eleven
1969-hatched birds including 2 hens and 9 cocks not needed to fill the
experimental breeding pens were left in combined holding pen numbers 3 and
4, and seventeen 1967-hatched or older breeders of mixed sexes were placed
in combined holding pen Numbers 5 and 6. Figure 2 shows the arrangement
of holding pens.
RESULTS AND DISCUSSION
Although procedures for the 1970 mountain quail tests were carried out as
planned, except for being approximately I month late in completing a set
of 10 experimental breeding pens, only 1 egg was laid. This was laid on
May 1, 1970 in the community pen (combined pen Numbers 3, 8, and alleyway)
and was infertile.
It was found in previous year's tests that mountain quail stress easily.
Catching and sorting the breeders at approximately the time when they should
have started to lay may have been the reason for almost no egg production
in 1970.

�-88-

N

~---------------30'--~
1F-----12'--~

~

~--

12' ---41

o

SCALE 8

.I
·0'

t

I.

5'

10'

Fig. 1.
Petallod

Arranoornont

of

Mou"toln

Quail BrlGdhiQ

Pont.

�-89-

I(

. 20'

~

lE-a'-)E-8'-)f~

f

10

N
~

5

I

3

1

-

\
4

6

2
.

Scole:

o
60' ----------------~
Fig. 2.
Mounialn

Quail

Holding

Pens.

I": 15 ft.

15

30

�-90-

No eggs were laid by ten wild-trapped mountain quail of mixed sexes received
from Oregon in Segment 23, eleven surplus 1969-hatched birds (2 hens and
9 cocks), or seventeen 1967-hatched or older breeders of mixed sexes held
in separate mountain quail holding pens in 1970. It was necessary to catch
and move these surplus breeders in order to select breeders used in the
experimental breeding pens and to utilize holding pens with the best cover.
After it became apparent that surplus breeders held in holding pens were not
producing eggs, 25 (seventeen 1967-hatched or older and eight 1969-hatched)
were banded and released on the west side of the Uncompahgre Plateau on
June 23, 1970. As of February 9, 1971, there were 26, 1969-hatched and
9 wild-trapped mountain quail from Oregon on hand for use as breeders in
the 1971 tests. A total of 7 birds, including 6 1969-hatched and one wildtrapped, died while in wintering holding pens, prior to February 9, 1971.

Prepared

by

!l~'M~~
onald M.offman
Wildlife Researcher

�-91April, 1971

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~----------

Project No.

W-37-R-24

Game Bird Survey

Work Plan No.

17

Job Title

Inventory of Selected Ptarmigan Populations

Period Covered:
Personnel:

Job No.

2

April 1, 1970 to October 15, 1971

Clait E. Braun and Terry A. May.

ABSTRACT
Investigations of white-tailed ptarmigan (Lagopus leucurus) populations in
Colorado initiated in 1965 were continued in 1970 with major emphasis upon
populations levels, and relationships of nesting success and production to
climatic conditions. Breeding density decreases ranged from .6 to 6.0 birds
per square mile on all study areas from levels,observed in 1969. The observed
decreases were primarily the result of poor nesting success and production in
1969 on all areas and excessive harvests in 1969 at Independence Pass and Mto
Evans. Observed densities in 1970 ranged from 5.2 at Mt. Evans to 24.8 in
RMNP. Weather during periods of egg deposition and incubation in 1970 was warm
and dry, and nesting success ranged from about 50 to 75 percent. Survival of
chicks to September 1 was similar to all previous years studied. Average brood
size to September 1 was 4.5 chicks. Present data continue to indicate that
nesting success and production are closely related to amount of precipitation
and number of frost-free nights in June.

��-93-

INVENTORY OF SELECTED PTARMIGAN POPULATIONS
C1ait E. Braun

Knowledge concerning population levels of white-tailed ptarmigan and factors
influencing observed densities is essential for proper management. Substantial data are available relating to this small grouse in Colorado (Braun
1971; Braun and Rogers 1971; and Braun and Schmidt 1971), but additional
information is necessary for management programs. It has been hypothesized
by many workers that all species of grouse (family Tetraonidae) are cyclic,
exhibiting either short-term (3-5 years) or long-term (10-12 years) fluctuations. Present data concerning grouse inhabiting northern areas support
this hypothesis, but few long-term studies have been conducted on species
of grouse at the southern edge of their range. It has not been demonstrated
that any species of grouse is cyclic in Colorado. Relationships between
weather conditions and population levels of grouse have been suggested by
Siivonen (1957), Neave and Wright (1969), and others, but have not been
statistically examined for white-tailed ptarmigan. Campbell (1968) and
Francis (1967; 1970) have demonstrated close relationships between certain
weather parameters and populations of scaled (Callipepla squamata) and
California quail (Lophortyx ca1ifornicus). This project was undertaken to
examine long term population trends and effects of climatic conditions on
white-tailed ptarmigan in Colorado. Following recommendations of Braun and
Rogers (1970) all studies were discontinued at Mesa Seco.
P. S. OBJECTIVE
To test the hypotheses that (1) populations of white-tailed ptarmigan in
Colorado are not cyclic and (2) production and fall population densities
of this species in Colorado are related to temperature and moisture conditions
during June of the same year.
SEGMENT OBJECTIVES
1.

To estimate breeding population levels of ptarmigan in five study areas
(Mt. Evans, Crown Point, Independence Pass, Mesa Seco, and Rocky Mountain
National Park).

2.

To estimate nesting success and production in the above areas.

3.

To obtain weather data from a representative alpine area for use in
determining correlation between spring weather conditions and fall
ptarmigan populations.

4.

To compile data and prepare progress report.

�-94-

METHODS

AND MATERIALS

Techniques used were essentially those developed under Work plan 17, Job 1
and reported in detail by Braun and Rogers (1971).
Ptarmigan were censused
by traversing study areas using tape-recorded calls to locate territorial
males, breeding pairs, and hens with chicks.
Efforts were made with telescoping noose poles to catch all ptarmigan not banded previously.
All birds
caught were banded with serially numbered gold anodized aluminum leg bands,
with all adults being additionally banded with individually numbered blue
plastiC bandettes.
Green bandettes with black numerals were used to replace
any bandettes from previous years that had become too worn for recognition
of individual birds.
Weather data were obtained from the Institute of Arctic and Alpine Research
(INSTAAR), University of Colorado, for their D-l, Niwot Ridge Site.
This
site is located at 12,300 feet between the Rocky Mountain National Park
(RMNP) and Mt. Evans study areas.
Data collected at this site and in RMNP
have been previously ahalyzed and compared (Braun 1969), and differences
were found to be insignificant.
Weather data at this site have been systematically collected using standard instruments since 1952 (Marr 1967a; Marr
et ale 1968a and b).

DESCRIPTION

OF STUDY AREAS

Areas intensively investigated, with one exception, are those studied since
1966 under Work Plan 17, Job 1, and described in detail by Braun and Rogers
(1971). As suggested by Braun and Rogers (1970) at the cessation of the
initial 5-year project on inventory of ptarmigan populations, all work was
terminated at the Mesa Seco site.
The four current study areas are presented
in Figures 1-5.

RESULTS AND DISCUSSION
Breeding

Densities

Censuses of breeding birds were initiated on May 2 with few males being
observed on known territories.
The first breeding pair was observed on
May 4 in RMNP but most pairing did not occur until mid-May.
Weather conditions during the 1970 breeding period were similar to those experienced in
1968 and were colder and drier than in 1969. Snow levels in early May were
slightly less than in previous years studied except 1966. Timing of breeding
activities was earlier than in 1969 but generally was similar to previous
years aild suffered no major disruptions froni adverse weather.
Densities
observed are presented in Table 1.
Breeding
observed

densities decreased on all areas studied
in earlier years (Table 2).

in 1970 from levels

�-95-

7

-

(:
---..,,,

//~- ..

/

Fig.

1.

\
Mt. Evans study area.

T EVANS

ARE

COLORA

0

Stydl1 Area ~()unC»OJrlf

�-96-

.'.:./

CROWN

POINT AREA

COLORADO
-

Study Area Boundary

/-.--~--.
"/0..

Fig. 2.

Crown Point study area.

"...
.".~/

�-97-

ROCKY MOUNTAIN

NATIONAL

Pj\RK

SUNDANCE UNIT
It!

J

-

COLORADO
Study Area Boundary
SCALE

1111MILES

bE=:r::::=::JE3E3:::=J:E3~'3/=2=EE3=:r::::=::JE3===::::j:

Fig. 3.

Rocky Mountain National Park, Sundance Unit study area.

~ILE

�.--------.

Fig. 4.

.,
.~
C

ra

,.

Rocky Mountain

National

-98-

U)

t-

Z
:,:)

«r ~

~
c(

0..

-' &lt;tZ

c(

••••
::l

:z

&gt;L..

c

0

"0

::)
0

en

l!

(It

&lt; )., '&gt;

~

tal

-

::!

:i

"0

&lt;II

u

w
..,j

..I

IU

~~

0

I

-' -

0

...
~
,..

Z
0

.,

~
c( 0 0
0
Z
...J ~
0:
Z
~ ..,J 0
::)
&lt;C

c(

en

0

0

I- ~
Z

:&gt;

::l 0::I

0
~

&gt;~

study area.

/0
1&gt;

Mountain

LL

~ 0:
0 ...J
0 ...J
0=

Park, Fall River-Toll

�-99-

INDEPENDENCE
N

ASS AREA

COLORADO

I T .•..•

Study Aroo Boundory
o••

0,,0

0.000

~==E:~~~1/~2E:~~e+3:3~1
o~
e-3 F3
E3
:+

Fig.

5.

Independence

Pass

s tudy area.

�-100-

Table 1. White-tailed ptarmigan breeding population densities, all areas,
1970.

Square
Miles

No. of
Breeding
Pairs

Unmated
Birds

Total
Breeding Season
Population

Birds per
Square Mile

Tombstone RidgeSundance Mountain

1.25

8

5 males

21

16.8

Toll Memorial

.19

4

3 males

11

57.9

Fall River Pass

.70

9

3 males

21

30.0

2.14

21

11 males

53

24.8

Crown Point

1.93

8

4 males

20

10.4

Mt. Evans

1.54

3

2 males

8

5.2

Independence Pass

1.12

4

4 males

12

10.7·

Study Area
RMNP

Total RMNP

Table 2.

White-tailed ptarmigan breeding population densities, 1966-70.

1966

Birds eer Sguare Mile
1967
1968
1969

1970

RMNP

29.0

25.2

29.4

30.8

24.8

Crown Point

14.5

21.2

18.1

14.5

10.4

Mt. Evans

7.8

7.1

7.1

5.8

5.2

Independence Pass

18.7

18.7

21.4

17.9

10.7

Study Area

Decreases in breeding population levels were greatest at Independence Pass
and lowest at Mt. Evans. Reasons for the observed decreases are related to
the generally poor nesting success and production of young in 1969 at all
aceas and the continued heavy exploitation of fall populations by hunters

�-101~

at Independence Pass and Mt. Evans in 1969 (Braun and Rogers 1971). The
decreases at RMNP and Crown Point can only be attributed to poor nesting
success and production as no hunting is allowed in RMNP and no birds were
reported harvested at Crown Point in 1969. All decreases were _the result of
poor recruitment of young birds into the breeding population in 1970.

Nesting Success and Production
Nesting activities were initiated on all areas in 1970 between June 10 and
June 20. Incubation of one nest under observation was initiated on June 22.
Weather conditions during June and early July 1970 were warm and dry and
were similar to those in 1966 and 1968 when nesting success and production
were good to excellent. Estimated peak of hatching in 1970 was between
July 15-20 with the one nest under observation hatching on July 15-16.
This nest in RMNP contained six eggs and all successfully hatched.
The primary molt of 15 successful hens was examined in order to calculate
hatching dates. The earliest calculated hatching date was July 12 and the
latest was July 28. Eleven of the 15 successful females hatched their
clutches between July 15-20 while only one hen was calculated to have
hatched her clutch after July 25.
In 1970, 32 different hens were observed during the brood period (July 12September 15) with 27 of these being successful. While successful females
are more easily seen than those without chicks, the number of unsuccessful
females observed was low in 1970. When the five unsuccessful hens seen in
1970 are compared with the 22 brood less females reported in 1969 (Braun and
Rogers 1971), it is apparent nesting success was substantially better in
1970 than in 1969. Estimated nesting success in 1970 was 75 percent in RMNP,
65 percent at Crown Point, and ~ 50 percent at Mt. Evans and Independence
Pass.
Initial size of broods in 1970 upon hatching was six chicks. Brood size to
August 1 averaged 5.3 chicks, while each brood averaged 4.5 chicks on
September 1 (Table 3). This was an increase of 1.6 chicks per brood over
the 2.9 chicks per successful hen recorded to September 1, 1969.
Fall Densities
Estimates of densities of white-tailed ptarmigan on September 1, while
useful in illustrating population gain through production, are difficult to
accurately derive due to a number of variables. Estimated fall densities
for each area studied are presented in Table 4. These densities were
calculated following the three basic assumptions discussed in detail by
Braun and Rogers (1971).
Data presented in Table 4 present a good picture of actual populations at
RMNP and Crown Point. This is not true of the data for Mt. Evans and
Independence Pass. All that can really be said of the fall densities at
these two locations is that they were low.

�-102-

Table 3.

Number of broods and average brood size, all areas, 1970.-1/
Number of
Broods Observed

Average Number of
Chicks per Brood

July 1-15

0

0.0

July 16-31

7

5.3

August 1-15

4

4.0

August 16-31

15

4.5

September 1-30

7

3.1

Total

33

Date

Average

4.3

1/ Only distinct broods are included.

Hunting further reduced fall densities at Crown Point as hunters in the fall
of 1970 harvested 15 of 29 banded birds in the population. At Independence
Pass, only 1 of 12 banded birds present in the study area was harvested,
while the Mt. Evans area was closed to all hunting of grouse in 1970.
Hunter check stations were not operated in 1970 so total harvest on the
three study areas open to hunting is not known.

Relationships of Weather:Nesting Success and Production
Weather data obtained from the 12,300 level of Niwot Ridge, through the
courtesy of INSTAAR for the 1966-70 period are presented in Table 5.
Detailed analysis and statistical correlations will be completed for the
final report in 1974-75. The close correlation between June weather
conditions and ptarmigan nesting success and production previously suggested
from work conducted under Work Plan 17, Job 1, continued in 1970. In
general, weather conditions were warm and dry in June and July, 1970, and
nesting success and production improved over 1969 when the early Summer
weather was colder and wetter.

�Table 4.

Estimated fall densities of white-tailed ptarmigan, 1970.

Area

Estimated
Percent
Nesting
Success

Average
Brood Size
on
September 1

Total
Production

Total
Breeding
Population

Total
Population
on
September 1

Birds
per
Square
Mile

Percent
Gain
I

I-'
0
w

RMNP

75

4.3

64

53

112

52.3

57.1

Crown Point

65

5.0

25

20

43

22.3

58.1

Mt. Evans

50

3.0

5

8

12

7.8

41. 7

Independence Pass

50

3.0

6

12

17

15.2

35.3

I

�Llb1e 5.

~.Jeatherdata, Niwot Ridge, 12,300 ft., May-July, 1966-1970.
TemEerature

Mean Daily
Max.
Min.

Number Days
Minimum
Temperature
Less Than 32° F

(Marr 1967b; Clark 1970).
PreciEitation
Number Days
Relative
Ppt/Month
Humidity
(Inches)
90 or Above

Ave. WindSpeed/
Month

Year

Month

Max.

Min.

1966

May

57

6

38.1

26.4

20

1.7

16 1.1

13.2

June

58

22

48.6

33.3

12

1.8

19

11.0

July

65

38

58.2

43.5

0

2.6

25

8.2

May

54

-7

32.5

21.5

25

3.1

25

15.1

June

58

24

44.6

30.4

18

3.7

29

12.3

1967

1968

1969

1970

(mph)

July

62

32

54.7

40.7

0

2.9

27

10.0

May

50

2

33.5

20.2

29

2.6

27 Jj

16.6

June

61

11

50.2

33.7

12

0.65

18

13.9

July

62

28

54.1

39.6

2

2.1

25

9.4

May

57

13

41.5

28.0

22

5.7

23

11.0

June

57

15

40.9

28.8

19

5.7

29

13.8

July

62

33

55.2

41.0

0

2.5

17

10.6

May

51

4

40.0

25.0

25

0.55

22

14.5

June

65

15

47.6

33.0

11

2.5

19

12.6

July

62

30

55.7

41.2

1

2.1

19

9.9

1/ Data not available for 4 days.
1/ Data not available for 1 day.

I
•....

0
.po.
I

�-105-

LITERATURE CITED

Braun, C. E. 1969. Population dynamics, habitat, and movements of
white-tailed ptarmigan in Colorado. Ph.D. Thesis. Colorado State
Univ., Fort Collins. 189 pp.
1971. Habitat requirements of Colorado white-tailed ptarmigan.
Proc. Western Assoc. State Game &amp; Fish Comma 51: In press.
______ , and G. E. Rogers. 1970. Inventory of Ptarmigan populations.
Colo. Div. Game, Fish and Parks, Game Res. Rep., Fed. Aid Proj.
W-37-R. April. pp. 181-183.
, and
1971. The white-tailed ptarmigan in Colorado.
------Div.
Game, Fish and Parks. Tech. Bull. 27: In press.

Colo.

------ , and R. K. Schmidt, Jr.

1971. Effects of snow and wind on wintering populations of white-tailed ptarmigan in Colorado. pp. 238-250.
In Haugen, A. O. (Ed.). Proc. Snow and Ice Symposium, Iowa State Univ.,
Ames. 280 pp.

Campbell, H.
Mexico.

1968. Seasonal precipitation and scaled quail in eastern New
J. Wildl. Mgmt. 32(3):641-644.

Clark, J. M. 1970. Unpublished data.
Colo., Boulder.

Inst. Arctic and Alpine Res., Univ.

Francis, W. J. 1967. Prediction of California quail populations from
weather data. Condor 69(4):405-410.
1970. The influence of weather on population fluctuations in
California quail. J. Wildl. Mgmt. 34(2):249-266.
Marr, J. W. 1967a. Data on mountain environments. I. Front Range, Colorado,
sixteen sites, 1952-53. Univ. Colo. Studies, Sere BioI. No. 27. 110 pp.
1967b. Unpublished data.
Colo., Boulder.

Inst. Arctic and Alpine Res., Univ.

, J. M. Clark, W. S. Osburn, and M. W. Paddock. 1968. Data on
------mountain
environments. III. Front Range, Colorado, four climax
regions, 1959-1964.

Univ. Colo. Studies, Sere BioI. No. 29.

179 pp.

______ , A. W. Johnson, W. S. Osburn, and O. A. Knorr. 1968. Data on mountain
environments. II. Front Range, Colorado, four climax regions, 19531958. Univ. Colo. Studies, Sere BioI. No. 28. 169 pp.

�-106-

Neave, D. J., and B. S. Wright. 1969. The effects of weather and DDT
spraying on a ruffed grouse population. J. Wild1. Mgmt. 33(4):
1015-1020.
Siivonen, L. 1957. The problem of the short-term fluctuations in
numbers of tetraonids in Europe. Finnish Papers Game Res. 19:
44 pp.

Prepared by

_~tU~!:::..::-_,_y=--.:....

--.:..)~~o&lt;:..::.-

Clait E. Braun
Asst. Wildlife Researcher

_

�-107April, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RAD~~O~ _

Project No.

W-37-R-24

Work Plan No.
Job Title

Job No.

1

Experimental Breeding of Tinamou

Period Covered:
Personnel:

19

Game Bird Survey

April 1, 1970 to March 31, 1971

Lawrence A. Webster, Robert L. Schmidt, John F. Corey and Donald M.
Hoffman.

ABSTRACT
A total of 478 eggs was laid by the 36 pale spotted tinamou (24 hens and 12 cocks)
on hand in May, 1970. Of these 50 (10.5 percent) were culls, 132 (27.6 percent)
were infertile, and 296 (61.9 percent) were fertile.
A normal incubation relative humidity of 55-56 percent (84~-85~Owet bulb) was
considerably better than a test incubation relative humidity of 50 percent
(82-3/40wet bulb) both in the hatching of total eggs set and in the hatching of
fertile eggs. In addition, fewer dead germs and pipped eggs which did not hatch
occurred with the normal incubation relative humidity of 55-56 percent in comparison with a test incubation relative humidity of 50 percent.
Higher numbers of total eggs were laid by breeders held in 4 separate pens with
selected groups of 2 hens and 1 cock each (248 eggs) compared with a community
of 8 hens and 4 cocks (62 eggs). In addition, 190 fertile eggs were laid by the
4 selected group pens of breeders compared with only 32 for the community pen.
More fertjle eggs were produced by breeders fed a standard 22 percent protein
level feed (190) compared with a similar feed containing 16 percent protein
level feed (74). A higher percent of the fertile eggs were also hatched from
the birds fed the 22 percent protein level feed (70.00 percent) compared with
the 16 percent protein level feed (51.35 percent).
This study was scheduled to run through 1972-73 but poor overwinter survival
under penned conditions has prevented the releasing of suitable numbers in the
field. A final field release of all remaining stock (82 birds) was made in
Baca County, south of Vilas, on April 30, 1971.

��-109-

EXPERIMENTAL BREEDING OF TINAMOU
Donald M. Hoffman
Good progress was attained in refining techniques for rearing pale
spotted tinamou in captivity at the Fort Collins Wildlife Research
Station in 1970, but overwinter losses were again heavy. From the
36 breeders (24 hens and 12 cocks) held for 1970 tests~ there were
187 chicks hatched but only 62 survived through the fall and winter
periods. A final field release consisting of 59 1970-hatched tinamou
plus 23 surviving 1969-hatched adults was made in Baca County, south
of Vilas, on April 30, 1971.
Although a few tinamou (exact numbers unknown) are known to have escaped
from the holding pens at Rocky Ford, it appears that overwinter losses
approximated 50 percent9 with heaviest losses occurring during and
immediately following the heavy snow and severe cold temperatures in
early January, 1971. In addition, many of the surviving tinamou showed
the effects of having toes frozen during the winter period.
P. S. OBJECTIVE
To develop game farm production techniques for pale spotted tinamou.
SEGMENT OBJECTIVES
1.

To compare two levels of relative humidity (50% versus 55%) during
incubation on egg hatchability.

2.

To measure the relationship of flock mating and selected group mating
to total egg production and fertile egg production.

3.

To measure the effects of two protein levels in the feed (22% versus
16%) on fertile egg production and egg hatchability.

METHODS AND MATERIALS
The detailed arrangement of tinamou breeding pens is shown in Figure 1.
The pen schedule for tinamou for 1970 is shown in Table 1.
Selected groups of 2 hens and 1 cock were placed in pens 1, 2, 4, 5, 6, 7,
9, and 10 and a community flock of 8 hens and 4 cocks was placed in the combined pen 3,8, and alleyway on April 10, 1970. Thus, a total of 36 birds
was used in breeding tests in 1970. All breeders were dusted with louse
powder and primaries on both wings were clipped prior to releasing in the
breeding pens. Two birds died while in the breeding pens and 34 were recovered on September 28, 1970. Numbers of eggs produced and numbers of
fert:ileeggs produced were compared for the selected groups in pens 2, 4, 6,
and 10 and the community group in pens 3,8, and alleyway combined, on all
22 percent protein level feed.

�-110-

N

~------------30'
I'
i"

,

-en

~

12

~

12

-"-

-

LO

0

sr

to

7

.'"
s»

1

to

v

r0

C\J

~

/

\

.

(j)

,.,
SCALE

Fig. 1.

Detailed

Arrangement

a

L~ __ I

I

O'

10'

5'

of Tlnamou Breeding Pens.

�-111-

Table l.

Pen schedule for tinamou, 1970.

Pen
Number

Protein Level
16%
22%
T

1
2

T

3

x

4

T

x
T

T
T
T

5

x

T

6

T
T

7
x

8

x
T

T

9
10

T

Alleyway

x

Number Pens on
Test

4

Key:

Mating
Flock*
Selected Groups
(8 Hens and 4 Cocks) (2 Hens and 1 Cock)

T = Pen on Test;

x
T
T

4

1

4

x = Pen not on Test·

* Pens numbe r 3, 8, and alleyway was used as 1 combined pen for flock mating.

Following normal culling procedures, settings of tinamou eggs were made
alternately (except for setting number 1) in the test and control incubators
every 7 days insofar as possible in 1970. Relative humidity in the control
incubator was held at the standard 55 percent relative humidity (8~0 wet
bulb) for the first 2 settings and then increased to 56 percent relative
humidity (85~0 wet bulb) based upon recommended procedures for Robbins
incubators. In the test incubators, eggs were incubated at a relative
humidity of 50 percent (82-3/40 wet bulb) using a Humidaire Model 50 for
the.first 3 settings and a Robbins Model C-l for the remainder. Temperature
in all incubators was maintained at a normal 99.750 F. Setting number 7
was placed in the control incubator rather than the test incubator because
of problems encountered in maintaining 50 percent relative humidity in the
Humidaire incubator and in anticipation,of securing a new Robbins Model C-l
incubator.

�-112-

Birds in even numbered pens 2, 4, 6, and 10 were fed a standard game
bird breeder feed containing 22 percent protein level. Those in odd
numbered pens 1, 5, 7, and 9 were fed a similar feed but containing 16
percent protein.
Eggs produced were marked to facilitate determination
of fertile eggs produced and numbers of fertile eggs that hatched.

RESULTS AND DISCUSSION
Table 2 lists a comparison of pale spotted tinamou production at the Fort
Collins Wildlife Research Station for the period 1967 through 1970, and
field releases made from 1969 through 1971. A gradual increase in numbers
of fertile eggs and numbers of chicks hatched is evident during the period,
using approximately equal numbers of breeders from 1968 through 1970.

Incubation

Humidity

Test

Table 3 lists hatching success of eggs in the test incubators (50 percent
relative humidity) in comparison with eggs in the control incubator (55
to 56 percent relative humidity).
Results showed a 6.38 percent better
hatch in the control incubators when considering percent of total eggs
hatched (46.58 % compared with 40.20%) and a 9.08 percent better hatch in
the control incubators when considering fertile eggs hatched (67.28%
compared with 58.20%).
Higher percentages of dead germs (21.64% for the
test incubators, compa~ed with 19.23% for the control incubator) and of
pipped eggs (6.70% for the test incubators, compared with 1.28% for the
control incubator) showed the incubation humidity was too low for this
species.
Results indicated the normal incubation relative humidity of 55-56 percent
(84~-85~0 wet bulb) was therefore better than a test relative humidity of
50 percent (82-3/40 wet bulb).

Flock Mating

Versus

Selected

Group Mating

Test

Table 4 lists a comparison of numbers of eggs produced and numbers of
fertile eggs produced by 8 hens and 4 cocks in a community pen with the
same number of birds in 4 pens of 2 hens and 1 cock each. All birds were
fed a standard 22 percent protein feed and treated as alike as possible.
A much larger number of total eggs was laid by the tinamou held in 4
separate pens with selected groups of 2 hens and 1 cock each (248 eggs)
compared with a community pen of 8 hens and 4 cocks (62 eggs).
In addition,
190 of the eggs laid by birds in the 4 selected group pens were fertile as
compared with only 32 for the community pen. Results obtained in 1970 were
similar to those secured in 1969.

�Table 2.

A summary of pale spotted tinamou production,

Item
Number of Breeders
Sexes

Total eggs laid

1967-1970, and field releases made 1969-1971.

1967

1968

Year
1969

1970

1971

17

37

36

36

0

24H;12C

24H; 12C

493

478

mixed
(connnunity pen)
177

mixed
(conununity pen)
335

1/

I

Total eggs set

166

316

434

I-'
I-'
W

428

I

Number fertile eggs

74

211

268

296

Percent fertile eggs

44.6

66.8

61. 8

69.2

Number chicks hatched

44

108

137

187

Percent fertile eggs hatched

59.5

51. 2

51.1

63.2

Number released, Baca County

None

None

42

40

82

1/ Original source of breeding stock provided by Foreign Game Introduction Program of United States Department
of the Interior included 20 birds received on June 25, 1966 and 17 birds received on March 8, 1967, all from
Argentina, South America.

�Table 3. A comparison of hatchability of pale spotted tinamou eggs under approximately 55 percent relative
humidity (normal) versus approximately 50 percent relative humidity, 1970.

Item

On Test

1/

Hatch
Number

Date
Set

Number
Set

1

5-18
6-1
6-15
7-6
7-20
8-3
8-17
8-31

12
14
21
43

3
5
8

10
12
14
16

3
4
4

19
8
8

Dead
Germs

Number
Pipped

4
7
6
5
5

o

6

2

2
2

o
1

4
2

Number
of
Cripples

Number
of
Good Hatch

Percent Hatch
Total
Fertile
Eggs
Eggs

1

4
3

9
8

33.33
21.42
47.61
34.88
54.54
40.74
42.85
34.78

44.44
30.00
58.82
62.50
72.00
57.89
60.00
53.33

78 (40.20)

40.20

58.20

10
12
15
20
13

76.92
66.66
53.57
54.05
41.93
30.43
U.93
14.81
57.69

76.92
80.00
83.33
71.42
81.25
43.75
56.52
25.00
88.23

o
o
o
o
o
o
o

10
15
18

27
21
23

6

4

8

5

194

60 (30.92)

42 (21.64)13 (6.70) 1 (0.51)

13
18
28
37
31
23
31
27
26

o

o
o
o
o

7
8

3
3
3
8
2
8

10

o

11
9

8
2

o

Sub-Total

234

72 (30.76)

45 (19.23) 3 (1.28) 3 (1.28) 109 (46.58)

46.58

67.28

Totals

428

132 (30.84)

87 (20.32)16 (3.73) 4 (0.93) 187 (43.69)

43.69

63.17

Sub-Total
Control Jj

33

Number
of
Infertile

2

5-25

4

6- 8

6
7
9
11
13
15
17

6-22
6-29
7-13
7-28
8-11
8-24
9-8

3

10
9

15

1
1
1

o
o
o
o
o
o
o
3

o

11

7

13
4

15

1/ Eggs incubated at 82-3/40 F wet bulb reading (50% reI. hum.) and temperature of 99.750 F. Setting numbers
1, 3, and 5 incubated in Humidaire Model 50 incubator and the remaining settings on test were incubated in a
Robbins Model C-l incubator.
~/ Eggs in settings number 2 and 4 incubated at 84~0 F wet bulb reading (55% reI. hum.) and the remaining
controls were incubated at 85~0 F wet bulb reading (56% reI. hum.) and temperature of 99.750 F. All settings
were incubated in a Robbins Model H-7 incubator.

I

I-'
I-'
+'
I

�Table 4. A comparison of numbers of pale spotted tinamou eggs laid by a flock consisting of 8 hens and
4 cocks with four selected groups of 2 hens and 1 cock each. 1/
Number
of
Infertile

Number
Fertile
Eggs

Number
of
Good Hatch

Pen
Number

Total
Eggs

Number
of
Culls

Number
Set

Flock

62

3

59

27

16

32

Sub-Total

62

3

59

27 (45.76)

16 (27.11)

32 (54.23) 16 (27.11)

Dead
Germs

16

Percent Hatch
Fertile
Total.
Eggs
Eggs Set
27.11

50.00

•...
•...
I

2

45

3

42

10

13

32

19

45.23

59.37

4

73

7

66

10

17

56

39

59.09

69.64

6

40

6

34

13

9

21

12

35.29

57.14

10

90

6

84

3

18

81

63

75.00

77.77

Sub-Total

248

22

226

36 (15.92)

57 (25.22) 190 (84.07)133 (58.84) 58.84

70.00

Totals

310

25

285

63

73

1/ All pens were fed 22 percent protein feed.

222

149

V1
I

�-116-

22 Percent

Protein

Versus

16 Percent

Protein

Feed Test

Table 5 lists a comparison of numbers of fertile eggs produced and numbers
of fertile eggs that hatched by tinamou breeders fed 16 percent protein
level feed with those fed a 22 percent protein level feed.
Selected groups
of 2 hens and 1 cock each in pen numbers 1, 5, 7, and 9 were fed the 16
percent protein feed and birds in pen numbers 2, 4, 6, and 10 were fed a
standard 22 percent protein level feed.
More fertile eggs were produced by the breeders fed the standard 22 percent protein level feed (190) compared with a similar feed containing 16
percent protein level (74). A higher percent of the fertile eggs hatched
were also from the birds fed the 22 percent protein feed (70.00%) compared
with the 16 percent protein feed (51.35%).
From these comparisons, it is evident that the standard 22 percent protein
level game bird breeder feed was much better than the 16 percent protein
level feed for pale spotted tinamou egg production and hatchability during
1970.

East Pens Versus West Pens
Although not planned as a separate test, a comparison of egg production by
east versus west pen location has shown the importance of providing adequate
natural cover in the breeding pens.
In 1969, a comparison of egg production
by 4 pens on the west side, each containing 7 or 8 established tall wheatgrass clumps, with 4 pens on the east side with no grass clumps, showed
highest numbers of total eggs were produced in the west pens (235 compared
to 193) and 162 fertile eggs were produced in the west pens compared with
only 73 fertile eggs in the east pens.
Prior to the 1970 egg laying season, a minimum of 8 clumps of tall and/or
crested wheatgrass were planted in the east pens and wire cages installed
to protect most of these growing clumps.
The established cover was left
in the west pens as well, so all pens had growing natural cover for 1970.
Table 6 lists a comparison of numbers of eggs laid and fertile eggs laid
by tinamou in the west pens with the east pens in 1970. Total eggs produced was 209 for the 4 west pens and 207 for the 4 east pens.
Numbers
of fertile eggs produced was 143 for the 4 west pens and 121 for the 4
east pens.
Location of pens plus a lack of adequate cover in the east
pens apparently caused this variation in egg production in 1969 but this
variation was considerably reduced in 1970 with the replanting of adequate
natural cover in the form of growing grass clumps.
The variation in
numbers of fertile eggs produced would probably be further reduced in
the future as the newly planted grass clumps mature but may never be
entirely eliminated because of a tall juniper windbreak on the west side

�Table 5. A comparison of numbers of pale spotted tinamou eggs produced and number hatched using 16 percent and 22 percent protein levels in the feed, 1970.

Pen
Number

Total
Eggs

Number
of
Culls

Number
Set

1

28

0

28

8

7

5

63

12

51

16

7

26

5

21

9

51

8

Sub-Total

168

25

Item

On Test
(16% Protein
Feed)

Control
(22% Protein
Feed)

Number
of
Infertile

Number
of
Good Hatch

Percent Hatch
Total
Fertile
Eggs Set
Eggs

Percent
Fertile
Eggs

13

46.42

65.00

71.42

22

13

25.49

37.14

68.62

5

6

10

47.61

62.50

76.19

43

40

1

2

04.65

66.66

06.97

143

69 (48.25)

36 (25.17) 38 (26.57)

26.57

51. 35

51. 74

Dead
Germs

,...
,...
•....•
I

I

2

45

3

42

10

13

19

45.23

59.37

76.19

4

73

7

66

10

17

39

59.09

69.64

84.84

6

40

6

34

13

9

12

35.29

57.14

61. 76

10

90

6

84

3

18

63

75.00

77.77

96.42

Sub-Total

248

22

226

36 (15.92)

57 (25.22)133 (58.84)

58.84

70.00

84.07

Total

416

50

369

105 (25.24)

93 (22.35)171 (46.34)

46.34

64.77

63.93

�Table 6. A comparison of numbers of pale spotted tinamou fertile eggs produced and number of fertile
eggs hatched by 4 west and 4 east pens, 1970.

Item
West Pens

Pen Number

Protein Level

Culls

Laid

Egg Totals
Set
Infertile

Fertile

Hatched

1

16%

0

28

28

8

20

13

2

22%

3

45

42

10

32

19

4

22%

7

73

66

10

56

39

5

16%

12

63

51

16

35

13

Sub-Total

22

209

187

44

143

84
I

t-"
t-"

East Pens

6

22%

6

40

34

13

21

12

7

16%

5

26

21

5

16

10

9

16%

8

51

43

40

3

2

10

22%

6

90

84

3

81

63

Sub-Total

25

207

182

61

121

87

Totals

47

416

369

105

264

171

00
I

�-119-

of the breeding pens, which provides additional shelter and shade to
the west pens. Natural cover in the form of growing grass clumps provides both shade and shelter for the breeders as well as for eggs laid.
By alternating pens by east and west location in all tests run in 1969
and 1970, bias due to pen location should have been eliminated.

��April, 1971

-121-

JOB PROGRESS REPORT

State of

~C~O~L~O~RAD~~O _

Project No.

W-37-R-24

Work Plan No.
Job Title

Job No.

2

Study of Tinamou Adaptability

Period Covered:
Personnel:

19

Game Bird Survey

April 1, 1970 to November 30, 1970

Donald M. Hoffman, Dale W. Stahlecker and Warren D. Snyder.

ABSTRACT
Forty pale spotted tinamou, Nothura darwinii salvadorii, were released in
May, 1970. This was the second release on U. S. Forest Service rangeland in
southern Baca County, Colorado. As in 1969, follow-up searches and checks
on reported sightings failed to provide evidence that tinamou remained, survived or reproduced in the vicinity of the release site.

��-123-

STUDY OF TINAMOU ADAPTABILITY
Warren D. Snyder
P. S. OBJECTIVE
To determine the ability of pale spotted tinamou to establish reproducing
populations in areas of mixed tall and mid-grasses, sandsage and yucca
on the Comanche National Gr~sslands, Baca County, Colorado
SEGMENT OBJECTIVES
1.

Introduce pale spotted tinamou.

2.

Determine dispersal of pale spotted tinamou and success of plant.
METHODS AND MATERIALS

Reference is made to Snyder (1969) for review of methods and materials used
in this study.
RESULTS AND DISCUSSION
Pale spotted tinamou raised at the Fort Collins Wildlife Research Station
under Work Plan 19, Job 1, were released on May 1, 1970 by Donald M.
Hoffman. This release included 15 males and 25 females. The location of
the release, along the Sand Arroyo in southern Baca County, was the same as
that of the previous year (Snyder 1969).
The vicinity of the release and surrounding localities were searched on
June 11-12, August 17-19 and November 1 and 2, 1970. The birds were not
observed, nor were there any indication of their presence in the area.
Bert Widhalm, District Conservation Officer, and Carrizo District personnel
of the U. S. Forest Service were contacted concerning sightings or reported
sightings. One local resident reported seeing the birds frequently during
the summer in a location approximately ten miles east of the release. Contact of this person left doubt as to the accuracy of his observations. His
description of the bird and its habits did not fit the tinamou.
In summary, there was no evidence of survival or reproduction of any of
the tinamou released either in 1969 or 1970. One additional release is
planned in 1970 with follow-up surveys to be made.

�-124-

Fig. 1. Pale spotted tinamou at release site in Baca County.
Don Domenick.

Photo by

�-125-

LITERATURE CITED
Snyder, W. D. 1969. Study of tinamou adaptability. Colorado Div. of
Game, Fish and Parks. Game Res. Rpt. April, pp. 177 -181.

Prepared

by

!f!.~yd'it~
Wildlife Researcher

�</text>
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                  <text>July, 1971
-1-

JOB PROGRESS

State of

REPORT

COLORADO
----~~--------------------

Project No.

W_-_l_0_l_-_R_-_1_3

Work Plan No.

------------~------~~

Job Title

_

Game Range Investigations

1
Job No.
Effects of Different In-t-e-n-s~i~t-i~e-s----------------------~o~f~~S~i~m_u~l~a~t_e~d
__B~r~ow~s~i_n~g~o_n~K~e~y~B~r_o~w~s_e~~S~p~e~c~i~e~s _

1

Period Covered:

April

1, 1970 to March 31, 1971

Personnel:

Harold R. Shepherd

ABSTRACT
A typed final copy of a manuscript entitled "Effects of Clipping on Key
Browse Species in Southwestern Colorado" was prepared for delivery to
Don Art Printers, Inc" to be printed as Colorado Division of Game,
Fish and Parks Technical Publication Number 28.

��-3EFFECTS
OF SIMULATED

OF DIFFERENT INTENSITIES
BROWSING ON KEY BROWSE SPECIES
Harold

R. Shepherd

P. S. OBJECTIVE
To publish

results

of completed

field work.

SEGMENT
1.

To make revisions

2.

To submit manuscript
cation.

OBJECTIVES

in the manuscript
to publisher

RESULTS

as necessary.
and follow

through

to final publi-

AND DISCUSSION

A first-draft manuscript was revised to conform to the suggestions of
five reviewers.
A typed copy of the revision was submitted to editor,
Dr. Lee E. Yeager, who obtained bids for publishing the manuscript and
made additional revisions.
His revisions were edited, and a typed
final copy of the manuscript was prepared, in Technical Publication
format, for delivery to Don Art Printers, Inc., the firm selected to
print the manuscript as Colorado Division of Game, Fish and Parks
Technical Publication Number 28.

Prepared

by

L&lt;, «1tJ4i.,~

v
Harold Ro Shepherd
Wildlife Researcher

""

��July, 1971

-5-

State of

COLORADO
----------------------------

Project No.

W~-~10~1~-~R~-~1~3~ _

4

Work Plan No.

Game Range Investigations
Job No.

la

Job Title

Inventory of Range Manipulation Projects in Colorado

Period Covered:

April 1, 1970 through March 31, 1971

Personnel:

Roland C. Kufeld, Richard M. Bartmann, and Regional Game
Biologists.

ABSTRACT
An inventory was made of all range type-conversion projects completed
during 1970, in the mountain half of Colorado, on lands administered
by the U.S. Forest Service and Bureau of Land Management. Acreages
treated were: Forest Service - 6,250 acres, and Bureau of Land Management - 3,422 acres; Total - 9,672 acres.

��-7-

INVENTORY

OF RANGE MANIPULATION

PROJECTS

IN COLORADO

Roland C. Kufeld

P. S. OBJECTIVE
To collect information which describes the location, extent, environmental conditions, land use practices, type of treatment and effects
of treatment on the range, livestock, and wildlife, for all completed
range manipulation projects in Colorado that are located within the
ranges of deer, elk and sage grouse, and develop a processing system
for information which describes proposed range manipulation projects.

SEGMENT OBJECTIVES
1.

To collect information which describes proposed and completed
vegetation modification projects that are located in Colorado
their effects on the range, wildlife and livestock.

2.

To compile,

3.

To provide desired IBM listings
agencies upon their request.

codify, process

METHODS

and analyze

inventory

of inventory

range
and

data.

data to cooperating

AND MATERIALS

An inventory of all range vegetation modification projects completed
through 1969, in Colorado, west of Interstate Highway 25, on lands
administered by the U. S. Forest Service, Bureau of Land Management
and Bureau of Indian Affairs has been completed, and the results reported by Kufeld (1968 and 1970). This report concerns vegetation
modification projects completed during 1970, however, it does not
include projects located on Indian Reservations, and they will be
deleted from future inventories.
Data were collected using procedures outlined
Game, Fish and Parks Administrative
Directive
Vegetation Modification Projects".

in Colorado Division of
No. 24, entitled "Range

Inventory data were transferred from original
Reader Forms, and then to IBM cards.

data sheets

to Mark Page

Procedures outlined in Administrative
Directive No. 24, call for
evaluations to be made on each vegetation modification project at the
end of the 2nd, 5th and 10th years, following treatment.
During 1970,
2 year post-treatment evaluations were made on 17 vegetation modification

�-10-

"("

__ ._-----

---_ ..

Hor« Unc l;y

---------_.
An irrm I
Specic!&gt;

-:-.------~-~--.-•./.-;.
l.e~f; Use f,y

••

-----.-----.

-----,---

__ .._-_._-- ---_.-._-----

Hcne-

De t r L»

f i.c i.n I

m~ntDl

CiO)

(51)

(52)

No
IUtcd

Un-

How

kno.zn

-------------

Dc t e r'm inc cl

(53)
Ar;cncies

(5t~)
Da t o

-----------------------------------------------------

Remarks

Form CompLe t ed By .
_
A-gene J.C s_:...-.
(ShOH

name,

title

and Agency

rr::-.:-=-

'l'it le

--,

_

Date
of all

persons

pa r t LcLpa t Lug in completion

of f01111)

�-11-

Table 1. Acreages of range land treated during 1970, in Colorado by the
U. S. Forest Service.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grass

Seed only

1

70

70

Spray

1

75

0

Pitting

1

75

75

3

220

145

Spray

5

2860

0

Pitting

2

810

810

7

3670

810

Spray

1

1300

0

Chain~./

2

850

0

Hand cutting

1

60

0

4

2210

0

1

150

150

1

150

150

15

6250

1105

Sagebrush

BrowseY

Pinyon-juniper

Seed only

All Vegetative Types Grand Total

l/ All vegetation modification projects conducted during 1970, in the
"browse" vegetation type were in the "oakbrush" subtype.

1/ Chaining includes chaining, cabling, railing, and bulldozing.

�-12-

Table 2. Acreages of range land treated during 1970, in Colorado by the
U. S. Bureau of Land Management.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Sagebrush

Seed only

1

900

900

Browse 11

Chain '2:..1

1

350

350

Pinyon-Juniper

Chain '2:..1

5

1972

1818

Abandoned lands

Seed only

1

200

200

8

3422

3268

All Vegetative Types Grand Total

II The vegetation modification project conducted in the "browse" vegetation
type was in the "oakbrush" subtype.
1/ Chaining includes chaining, cabling, railing and bulldozing.

�-13-

Table 3. Acreages of range land treated during 1970, in Colorado by the
U. S. Forest Service and U. S. Bureau of Land Management.
Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grass

Seed only

1

70

70

Spray

1

75

o

Pitting

1

75

75

3

220

145

Seed only

1

900

900

Spray

5

2860

o

Pitting

2

810

810

8

4570

1710

Spray

1

1300

o

Chain '1:..1

3

1200

350

Handcutting

1

60

o

5

2560

350

Seed only

1

150

150

Chain '1:..1

5

1972

1818

6

2122

1968

1

200

200

1

200

200

23

9672

4373

Sagebrush

Browse 11

Pinyon-juniper

Abandoned lands

Seed only

All Vegetative Types Grand Total

11 All vegetation modification projects conducted during 1970, in the "browse"
vegetation type were in the "oakbrush" subtype.

~I Chaining includes chaining, cabling, railing, and bulldozing.

�-14-

Table 4. Acreages of range land treated during 1970, within each National
Forest in Colorado.
National
Forest

Vegetative
Type

Kind of
Treatment

Grand Mesa

Sagebrush

Spray

Routt

San Juan

White River

Browse

Grass

Acres
Treated

Acres Seeded as
Part of Treatment

2

1440

0

2

1440

0

1

60

0

1

60

0

1
1
1

70
75
75

70
0
75

3

220

145

Hand Cutting

Seed only
Spray
Pitting

Sagebrush

Pitting

2
2

810
810

810
810

Browse 1./

Chain Jj

1

600

0

1

600

0

Pinyon-Juniper

Seed only

1
1

150
150

150
150

All Veg. Types

All Treatments

7

1780

1105

Spray

3

1420

0

3

1420

0

1
1

1300
250

0
0

2

1550

0

Sagebrush

Browse 1./

Total Al1
Forests

No. of
Projects

Spray
Chain 2:../

All Veg. Types

All Treatments

5

2970

0

Total Al1 Veg.
Types

Total Al1
Treatments

15

6250

1105

1/ All vegetation modification projects conducted during 1970 in the "browse"
;egetation type were in the "oakbrush" subtype.
2:../

Chaining includes chaining, cabling, railing and bulldozing.

�-15-

Table 5. Acreages of range land treated during 1970, within each Bureau
of Land Management District in Colorado.
BLM
District
Craig

Montrose

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Abandoned Lands

Seed only

1

200

200

1

200

200

1

350

350

I

350

350

4

1818

1818

4

1818

1818

Browse 1./

Pinyon-Juniper

Canon City

Grand Junction

Total A1l
Districts

Chain ])

Chain ])

Acres Seeded as
Part of Treatment

All Veg. Types

All Treatments

5

2168

2168

Pinyon-Juniper

Chain

I

154

o

I

154

o

1

900

900

I

900

900

8

3422

3268

Sagebrush

Total All Veg.
Types

Seed only

Total All
Treatments

1/ The vegetation modification project conducted in the "browse" vegetation
type was in the "oakbrush" subtype.

1/ Chaining includes chaining, cabling, railing and bulldozing.

��July, 1971
-17-

JOB PROGRESS REPORT

COLORADO
------~~~~--------------

State Of

Project No.

W-101-R-13

Work Plan No.

Game Range Investigations

Job Title.

~4~~~ __~ __~~
Job No.__~~~~2~~------------------Experimental Improvement of Oakbrush
o~n~D~e~e~r~a~n~d~E~l~k~W~i~n~t~e~r~R~a~n~~g~e~s
__-~B~e~a~v~e~r~C~r~e~e~k~ _

Period
Covered:
~

April 1, 1970 through March 31, 1971

Personnel:

Roland C. Kufe1d

ABSTRACT
Visual observations one year following treatment indicated an approximate
80 to 85 percent kill on Gambe1 oak (Quercus gambe11ii), and 90 percent
kill on snowberry (Symphoricarpos sp.) as a result of spraying by helicopter with 2 pounds of 2,4,5-TP and 10 gallons of water per acre. A
marked increase in grass production was observed.
Study plots along the boundary of the spray and control areas were
classified as to whether they had been sprayed. Photos were retaken
at photo points showing vegetation conditions one year after spraying.
The first post-treatment evaluation involving measurements of vegetation
and deer and elk use is scheduled for the summer of 1971.

��-19-

EXPERIMENTAL

IMPROVEMENT

ON DEER AND ELK WINTER

OF OAKBRUSH

RANGES - BEAVER CREEK

Roland C. Kufeld

P.S. OBJECTIVE
To determine
brush winter

if deer and elk carrying capacity can be increased on oakgame ranges by spraying with 2,4,5-TP to induce sprouting.

SEGMENT OBJECTIVES
1.

To measure percentage composition and cover of all plant species
on the study area prior to treatment and periodically following
treatment to determine vegetative changes.

2.

To measure deer and elk use on the study area prior to treatment
and periodically following treatment to determine changes in game
use.

METHODS

AND MATERIALS

The work accomplished during this segment was necessary for fulfillment
of the two segment objectives, however, it represents only a portion of
the work needed to complete post-treatment measurements of vegetation
and deer and elk use. Additional procedures will be implemented in
future segments to complete post-treatment measurements.
All vegetation measurement and pellet group plots were ref lagged
during May, 1970, so they could be relocated later in the summer when
foliage was full grown.
Vegetation measurement
and pellet group plots located along the boundary
of the spray and control areas were classified during late August, 1970,
as to whether they had been sprayed.
Photos were retaken at photo points during late August and early
September, 1970, to show vegetation conditions one year after spraying
by helicopter with 2 pounds of 2,4,5-TP and 10 gallons of water per
acre. These are directly comparable to photographs taken from the same
locations in late August and early September, 1968, showing vegetation
conditions one year before treatment (Kufeld, 1970).

�-20-

RESULTS AND DISCUSSION
Kill on Gambel oak (Quercus gambellii) one year following treatment
was estimated at about 80 to 85 percent on the basis of visual observations.
There was some basal sprouting of oak but sprouts were not
particularly numerous.
Sprouts are expected to become more abundant
during the second year following treatment.
Approximately 90 percent
of the snowberry (Symphoricarpos sp.) appeared to have been killed
by spraying.
Prior to spraying snowberry was the most abundant understory species.
A marked increase in grass production was observed in
the spray area. Photos showing vegetation conditions one year before
and one year after treatment are shown in Figs. 1 and 2.
The first post-treatment evaluation involving measurements of vegetation and deer and elk use is scheduled for the summer of 1971.

LITERATURE

CITED

Kufeld, Roland C. 1970. Experimental improvement of oakbrush on
deer and elk winter ranges - Beaver Creek.
Colo. Div. of Game,
Fish and Parks, Game Res. Rept. July, 1970, Pt. 1, Fed. Aid
Proj. W-lOl-R-12, p. 95-106.

Prepared

by~C/~
Roland C. Kufeld
Wildlife Researcher

�-21-

Fig. 1. Vegetation conditions
Area one year before treatment

on the Beaver Creek Oak Control
(Sept. 2, 1968).

Study

Fig. 2. Vegetation conditions on the'Beaver Creek Oak Control Study
Area one year after spraying by helicopter with 2 pounds of 2,4,S-TP
and 10 gallons of water per acre.

��July, 1971

-23-

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~-----------------

Project No.

W_-_l_0_l_-_R_-_l_3

Work Plan No.

_

Game Range Investigations

Job Title

4
Job No.
3
Experimental Improvement of Oakbrush on Deer
Elk and Cattle Ranges - Hightower Mountain

Period Covered:

April 1, 1970 through March 31, 1971

Personnel:

~~---~---~~------------------------

Roland C. Kufeld, Richard Fentzlaff,
Freddy, Mattias Kniesel.

Charles Cesar, David

ABSTRACT
seventy-five one meter square vegetation measurement plots and 10 permanent
photo point stations were established in each of the 8 habitat improvement
units.
Vegetation plots were located randomly, and their locations surveyed
with a K &amp; E compass and steel tape.
Pre-treatment vegetation measurements were made between July 10 and August
13, 1970. The measurement technique involved the use of a meter square
plot in which green weights of all grasses and forbs rooted inside the plot,
and portions of all shrub species which occur or protrude into the plot up
to a height of 5 feet above ground level were determined.
Green weights
were later converted to air dry weights.
Green weights of grasses and
forbs were determined by clipping and weighing by species to the nearest
gram.
Green weights of browse species were measured using a variation of
the weight estimate technique originally conceived by Pechanec and Pickford
(1937) and further described by Shoop and McIlvain (1963).
During a 10 day training period observers became very proficient in estimating weights of current annual growth of shrub vegetation.
In all cases
average observer weight estimating error was less than 10 percent, and in
most cases less than 5 percent.
Average estimated-actual weight correlation
coefficients for all observers were Gambel oak (Quercus gambellii),
0.93;
Serviceberry (Amelanchier alnifolia), 0.90; chokecherry (Prunus virginiana),
0.88; snowberry (Symphoricarpos sp.), 0.87; and big sage (Artemisia tridentata),
0.82.

�-24-

ABSTRACT

(continued)

Total air dry weights of forage ranged from 1,621 1bs/ac. in Unit 4 to 1,885
1b/ac. in Unit 3, and averaged 1,717 1bs/ac. Average plant composition for
all units was forbs, 31 percent or 532 1bs/ac.; grasses, 22 percent or 376
1bs/ac.; and shrubs, 47 percent or 810 1bs/ac.
The most common forbs were
lupine (Lupinus argenteus), 135 1bs/ac.; aspen peavine (Lathyrus 1eucanthus),
90 Ibs/ac.; and various species of Aster and Erigeron,
801bs/ac.
Most
common grasses were Kentucky bluegrass (~pratensis),
287 1bs/ac. and
wheatgrass (AgrOP ron sp.), 48 1bs/ac.
Gambel oak was the most common shrub
averaging 255 lbs ac. Other common shrubs were snowberry, 243 lbs/ac.;
serviceberry, 107 Ibs/ac.; chokecherry, 93 1bs/ac.; and big sage, 541bs/ac.

7

Pre-treatment cattle use as reflected by numbers of cow chips deposited in
pellet plots was unevenly dispersed among the 8 habitat improvement units.
By far the most use occurred in Unit 8 which had 18.26 cow days use per
acre. No use was recorded in Units 3 and 4. Deer and elk pellets were
removed from pellet plots on September 3, 1970. Accumulated pellets will
be counted in May, 1971 to obtain a measurement of pre-treatment deer and
elk use.
An herbicide for spraying oak and species for reseeding the study area were
selected on the basis of a literature review, and advice solicited from
professional people experienced in chemical spraying and range reseeding.
The herbicide, 2,4,5-TP (Silvex) will be applied by helicopter at a rate of
2 1bs/ac. mixed with 4 gallons of water and 1/2 gallon of number 2 diesel
fuel per acre. A liquid detergent soap will be added at a rate of 0.5% of
the total spray and carrier volume to facilitate mixing of the water, chemical and diesel fuel.
The following plants and seeding rates were selected:
Fairway crested
wheatgrass (Agropyron cristatum), 2 lbs/ac.; Lincoln smooth brome (Bromus
inermis),
4 lbs/ac.; amur intermediate wheatgrass (Agropyron intermedium),
3 lbs/ac.; chinook orchard grass (Dactylis glomerata),
1 lb/ac.; hard sheep
fescue (Festuca ovina duriscula),
1 lb/ac.; Nomad, Rambler and Ladak varieties
of alfalfa (Meticago sativa),
2 1bs/ac. (total); yellow sweet clover
(Melilotus officinalis),
1 lb/ac.; chickpea milkvetch (Astragalus cicer),
1 lb/ac.; bitterbrush (Purshia tridentata),
2 lbs/ac.; and mountain mahogany
(Cercocarpus montanus), 1 lb/ac.

�-25-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH ON DEER,
ELK AND CATTLE RANGES - HIGHTOWER MOUNTAIN
Roland C. Kufeld

P. S. OBJECTIVE
To determine the extent to which deer) elk and cattle forage production and
game use can be incresed and maintained by chaining, spraying and controlled
burning on overage Gambel oak winter game ranges.
SEGMENT OBJECTIVES
1.
2.
3.
4.

To install permanent vegetative measurement plots in each of the 60 acre
habitat improvement units.
To establish permanent photo points in each of the 60 acre habitat
improvment units.
To make pre-treatment vegetation and game use measurements, take photo
point pictures and assess any preference which cattle might exhibit
toward certain habitat improvement units.
To make a literature review.
METHODS AND MATERIALS

Layout of the study area and methods used in locating and establishing
eight 60 acre habitat improvement units and pellet plots within those units
have been described by Kufe1d (1970). A map showing the habitat improvement units is shown in Fig. 1. Treatments scheduled for these habitat
improvement units are as follows:
Randomized
Block Number
1

Habitat Improvement
Unit Number

3
4

Burn
Spray
Chain
Control

5
6
7
8

Spray
Control
Burn
Chain

1
2

2

Scheduled
Treatment

Detailed maps such as those in Figs. 2 and 3) which show unit boundaries
and locations of 60 permanent .05 acre circular pellet group plots established in each unit) have been made for each of the 8 habitat improvement
units.

�-26Vegetation

Measurement

Plot Establishment

Seventy-five vegetation measurement plots were established in each habitat
improvement unit. Sample size computations based on 80 preliminary meter
square plots read during 1969, indicate 75 plots are sufficient to permit
detection of treatment induced changes of 10 percent in all vegetation, 15
percent in browse, 25 percent in forbs and 25 percent in grasses, as being
significant at the 95 percent confidence level (Kufeld 1970).
Vegetation measurement plots were laid out with a K and E compass set to
allow for 14 degrees of east declination, and a steel tape. When measuring
distances no allowance was made for slope. The tape was always pulled tight.
In order to avoid trampling vegetation in the plots when surveying plot
locations all plots were offset four feet to the left of the original plot
survey line.
Habitat improvement Units 1, 2, 3, 5, 7, and 8 each contain a 30 acre area
in the center of the unit, and it is this center portion that will be treated.
The outside 30 acres will be left for wildlife cover. No 30 acre units were
established in the centers of the 2 control units (Units 4 and 6). Vegetation measurement plots were located only in the 30 acre center portion of
Units 1, 2, 3, 5, 7, and 8. In Units 4 and 6 they were distributed in the
entire 60 acre area.
Vegetation measurement plots were established along randomly located parallel
lines. One side of the inside 30 or outside 60 acre unit (depending on
whether the unit was to be treated or used for control purposes) was used
as a base line. The starting point of each vegetation plot line was established randomly along the base line and marked with a vegetation plot stake.
The distance between the starting point stake and the first vegetation plot
on the line was selected at random and varied from 0 to 50 feet. The distance between each vegetation plot on a line is 50 feet. Detailed maps were
made of each habitat improvement unit showing the exact location of each
vegetation measurement plot (Fig. 4).
Each vegetation plot was marked with two orange 3/8" diameter, reinforcement bar, steel stakes set one meter apart and on line. A numbered metal
tag was fastened to the first stake of each plot. Each tag bears the
designation
"v" indicating it is a vegetation plot, the line number, and
the plot number.
Strips of red plastic surveyor's flagging were tied to
the tagged stake and to an overhanging tree branch near the stake.

Photo Point Establishment
Ten photo points were established and pre-treatment photos were taken
between July 23, and July 28, 1970. Certain vegetation measurement plots
were selected for photo point stations.
An 11 x 16 inch blackboard with
the unit, line, and vegetation plot numbers written in chalk was suspended
from a nearby branch.
This became the photo center and focusing point.
The
compass bearing and distance from the plot stake to the center of the blackboard, and the date and time of day the photo was taken were recorded.
Photo
point centers could be either tpe first or second vegetation plot stake. A
record was made of the stake used for each photo point so future photos can
be taken from exactly the same spot.

�-27-

The camera was held directly over the stake and two photos were taken
at each station.
A black and white photo was taken with a Ciroflex
camera using 120 plus X film, and a 35 rom color slide was made with a
Honeywell Pentex Spotmatic camera using Kodachrome II film. Because
of differences between the two cameras in the type of viewfinder black
and white photos were taken at chest level and slides at eye level.
A sample photo point picture is shown in Fig. 5.

Pre-treatment
Pre-treatment vegetation
August 13, 1970.

Vegetation

measurements

Measurements

were made between

July 10, and

The technique used to measure vegetation during the pre-treatment
evaluation phase of the study, and which will be used to make posttreatment evaluations involves the use of a meter square sampling unit in
which green weights of all grasses and forbs rooted inside the plot, and
portions of all shrub species which occur or protrude into the plot up to
a height of 5 feet above ground level are determined.
Green weights are
later converted to dry weights.
A vegetation measurement method based
on forage weights rather than frequency of occurrence was preferred, because it was felt that treatment induced vegetation changes described in
terms of forage weight would be more meaningful to wildlife and livestock
managers than would changes in frequency of occurrence of plant species.
Green weights of grasses and forbs are determined by clipping and weighing
by species to the nearest gram.
Green weights of browse species are estimated using a variation of the weight estimate technique originally conceived by Pechanec and Pickford (1937) and further described by Shoop and
McIlvain (1963).
Shrub vegetation is measured by the weight estimate
technique rather than clipping because clipping would severely damage
shrub vegetation in the permanent study plots.
The decision was made to clip and weigh grasses and forbs by species because preliminary tests of the weight estimate technique during 1969 revealed weights of individual grasses and forb species could not be estimated
as accurately as shrubs.
This is in contrast to findings of Pechanec and
Pickford (1937) and Shoop and McIlvain (1963) who reported success in estimating forb and grass w.eights. The main reason for this difference can
probably be attributed to the relatively numerous and diverse species of
forbs and grasses and the greater abundance of these plants in the Gambel
oak type of Hightower Mountain as compared to vegetation conditions in areas
where the weight estimate technique was tested by these other workers (76
different forbs and 15 grasses were encountered in vegetation measurement
plots on the Hightower Mountain area).
In this study it was desired that
the relative abundance of each species be determined as accurately as
possible.
Because of this large number of diverse species the time required
to acquire and maintain an acceptable level of proficiency in estimating
weights of each species, and for collecting data from which to compute
correction factors for observer weight estimating error was prohibitive.
Clipping was considered to be the most accurate method for determining
green weights of individual grass and forb species.
The effects of clipping
on grasses and forbs in permanent study plots should be minimal because
plots will only be clipped every third year for the duration of the 12 year
study.

�-28-

Use of the Weight Estimate

Technique

Before vegetation measurements could begin, it was necessary to train the
observers to use the weight-estimate
technique.
A ten day period was
allowed for training observers.
Observers trained on one species at a time.
Since Gambe1 oak (Quercus gambe1lii), snowberry (Symphoricarpos sp.), chokecherry (Prunus virginiana), serviceberry (Amelanchier a1nifo1ia), and big
sagebrush (Artemisia tridentata) are the most abundant species on the area
each observer practiced until he became proficient in estimating green
weights of thosespecies.
Once proficient in estimating weights of these
five plants little extra effort was needed to attain proficiency in estimating green weights of other less abundant shrubs.
Weights of all current annual growth including leaves but not including
acorns, stalks on big and silver sagebrush (Artemisia cana) and berries and
flowers were estimated.
Chrysotharnnus flowers were included in the estimate,
however, because they contributed little to the total plant weight, and
because it was easier to include than to exclude them.
The observers learned to recognize units of 10 grams or fractions of 10 grams
of current annual growth of each shrub species.
This was done through numerous trials where the observer estimated units of 10 grams or less, and
then checked his estimate by clipping and weighing the vegetation to the
nearest gram.
A German-made "Feine Federwaage, Tragkraft" ZOO gram capacity scale was used
for weighing vegetation.
The scale is 6 inches long and 3/4 inch in diameter,
and has a ring by which it can be conveniently hung from a nearby branch.
Scales were obtained from the Oxwall Tool Co., Ltd., 133-10 3Znd Avenue,
Flushing, New York 11354. Clipped vegetation was placed in a "lunch bag"
size paper sack and weighed by hanging the sack on the scale hook by means
of a "binder type" paper clip attached to the sack.
After the observers learned to recognize vegetation units of ten grams or
less they practiced estimating the weights of forage on a limb or entire
shrub by counting the number of 10 gram units starting from the bottom of
each branch and working to the top until the entire shrub or limb was covered.
They then clipped and weighed the material to check their estimates.
When
counting the number of 10 gram units on a limb or shrub the east unit was
often a fraction of a unit. Some observers found that they could estimate
weights of some shrub species most accurately by estimating the weight of
each current annual growth twig encountered and keeping the running total
in mind rather than counting the number of 10 gram units.
However, in no
case were attempts made to estimate in units larger than 10 grams.
It is not possible to estimate exactly the weight of current annual growth
on a limb or shrub every time or even most of the time. Some estimates will
be too high and some too low, but these are compensating errors.
On a series
of plots, some with high and some with low estimates the compensating nature
of these errors should result in an estimate by most observers of the total
weight of vegetation that is accurate to within 10 percent.
If the average
error an observer makes when estimating the weight of a certain shrub
species is known, his weight estimate data can be adjusted to correct for
his error.

�-29-

Due to the large amount of work involved in measuring vegetation on the
Hightower Mountain Oak Control Study Area it was necessary to employ several
student assistants to make vegetation measurements.
Since there are differences among individual observers in their ability to estimate weights of
browse it was necessary to adjust weight estimate data collected by each
observer to correct, for individual estimating error.
When observers had attained proficiency in estimating weights of current
annual growth of each of the five aforementioned shrub species a set of .
data was collected for each observer to determine his average estimating
error on each shrub species. Since there were five observers, five shrub
samples of the same or of different species were tagged. A single shrub
sample might consist of one plant or could be several plants or branches,
but all of the plants in an individual sample were of the same species.
Each observer then estimated and recorded his estimate of the weight of each
sample. One observer was assigned to clip the current annual growth from
each sample. In this manner a set of estimated and actual weights for each
of the five major shrub species was obtained for each observer. A ratio (R)
was computed for each observer and each of the five major shrub species as
follows:

The variance of this ratio was computed using
...••.

V (R)

£y2

_ 2 ~ ~Xy

+ 'R2~X2

X2

(n - 1)

(h)

formula:

The number of samples needed by each observer to estimate his weight estimating error on each shrub species within 10% of his true mean error at
the 95% confidence level was determined through application of the formula:
N

:t.1

where:

V

(R)

n

d2

d

allowable deviation from the true ratio.
The allowable deviation is expressed as a
percent of 'R.

Approximately one day per week during the entire vegetation measurement
period was devoted to collection of estimated and actual weight data by all
observers. Thus, by the end of the vegetation measurement period each observer had many more samples of each of the five shrub species than he needed
as indicated by sample size computations. However, it was felt that observers needed this continued practice in order to maintain a high level of
estimating accuracy. Also, as the summer progressed, vegetation weights
changed as vegetation became mature and drier, so it was necessary for the
observers to practice in order to retain their concepts of vegetation
weight units.
Ratios ~) of estimated to actual weights were computed for all estimated
and actual weight data collected during the vegetation measurement period.
These were applied to weight estimates recorded for the 5 major shrub
species in meter square plots in order to correct for observer weight estimating error. Meter square plot weight estimate data for other less

�-30-

abundant shrub species were not corrected for observer weight estimating
error, because time did not permit collection of data from which to compute
correction factors for those species.
A

.

Correction factor (R) data were derived from individual tagged plants
rather than by estimating and clipping vegetation within meter square plots
in order to insure that exactly the same pieces of vegetation would be estimated by all observers, and to insure that only estimated vegetation would
be clipped.
A meter square plot was actually an imaginary box 1 meter
square and 5 feet tall, and it was up to the observer to decide whether a
piece of overhanging vegetation was inside or outside the plot boundary.
Thus, in a meter square plot one could not be sure that all observers would
include exactly the same pieces of vegetation in their estimates, or that
the vegetation clipped would be exactly that which was estimated.

Plant Identification

Training

Obviously, before vegetation measurements could be made it was necessary
that observers be schooled in identification of all plant species on the
study area.
This was accomplished in a three day period with the aid of
mounted plant specimens collected and identified during 1969 (Kufeld 1970),
and color photographs of plants taken when they were collected.
New species
which were encountered during vegetation measurements were collected,
assigned temporary numbers and later identified by personnel of the Colorado
State University Botany Department.

Measurement

of Vegetation

in Meter Square Plots

A meter square plot was selected as the sampling unit for two reasons: (1)
A relatively large plot was desired because the larger the plot the greater
the area per amount of perimeter.
Thus, the chance of observer
error in
judging whether a piece of overhanging shrub vegetation was in or out of
the plot would be reduced.
(2) By increasing plot size the chance of encountering an individual plant species is increased and the number of plots
needed to adequately sample an area is reduced.
The meter square plot, however, was not too large to be easily measured in the Gambel oak vegetation
type.
The plot frame was constructed of 1/4 inch steel rod. All four sides of
the frame were separate.
A 1-1/2 inch section of 1/8 inch pipe was welded
at a 90 degree angle to each end of two of the sides.
The plot was assembled
by inserting the ends of the other two sides into the pipe sections.
Both
corners of one side to which pipe sections had been welded were equipped
with 9/16 inch diameter metal rings, and each ring was slipped over one of
the two plot stakes.
The frame was assembled so it always pointed 90 degrees
to the left of the observer as he stood on the plot line facing down the line
from the first to the last plot. Thus, the frame can always be replaced in
exactly the same position at a later date. One exception to the standard
plot placement procedure occurred in Habitat Improvement Unit Number 5 where
the frame was mistakenly placed to the right on all plots on lines 1, 2,
and 3, and plots 1 through 7 on line 14. The frame was always kept horizontal.
On a steep hill telescoping forked rods were used to prop up the
down hill side of the frame to maintain a level position.

�-31-

Articles of equipment needed for measuring vegetation are shown in Fig. 6.
They include the meter square plot frame, scales, grass shears, paper sacks,
binder type paper clips, a small bucket, clipboard and supply of data forms.
All grasses and forbs which are rooted inside the plot and shrubs which
occur or extend into the plot up to a height of 5 feet above ground level
were listed on the data form (Fig. 7). This means 5 feet above the ground
at any given point within the plot. Browse species that were rooted in the
plot but had no current annual growth extending into the plot below 5 feet
were not listed.
All portions of forbs and grasses (except Pachistima which was estimated
rather than clipped because it is semi-woody and might be damaged by clipping)
including flowers and fruits which grew during the current growing season,
and which were rooted inside the plot were clipped by species and weighed
to the nearest gram. Plants were clipped to a level of 1 inch above the
ground and plants that were less than one inch tall were excluded. Clipped
material was stored in the bucket until all material for a species had been
removed from the plot. Material for that species was then weighed and discarded and clipping began on the next species.
After all forbs and grasses were clipped and weighed weights of current
annual growth of shrubs (and Pachestima) which occurred in the plot were
estimated using the same techniques employed during the training phase. It
was found easier to estimate the least abundant browse species in the plot
first and the most abundant species last. During the course of measuring
vegetation the observers periodically estimated and clipped 10 gram units of
various species of browse in order to maintain their vegetation weight unit
concepts. This was usually done as the observer began to estimate browse
.vegetation within a plot, and was done outside the plot to avoid damaging
vegetation in the plot.
Conversion of Green Weights to Air Dry Weights-~To facilitate direct
comparison of the relative abundance by weight of various species it was
necessary to convert green weights of all plant species encountered on meter
square vegetation measurement plots to air dry weights because: (1) a great
deal of variation in moisture content exists among plant species. (2) Moisture contents of individual species change drastically during the summer as
plants mature and dry out. One hundred grams of green forage at the beginning of the summer may contain only 25 grams of dry material, whereas, at
the end of the summer the dry matter in 100 grams of green forage of the
same species may weigh 50 grams.
Three green weight samples of 26 species of forbs, grasses, and shrubs were
collected in paper bags at approximately 10 day intervals during the vegetation measurement period. Weights of each sample ranged from 50 to 110
grams. These were weighed to the nearest gram and were hung indoors on
wires to dry. They were reweighed at the end of approximately one month
and again after about two months. All sample weights remained·unchanged
during the last two weighings so they were considered to be air dry.
The 26 species were chosen because they are the most abundant species on the
study area, they are present throughout the entire summer, and they range in
moisture content from the wettest to the driest plants on the study area.

�-32-

Hereafter, they will form the basis for converting green weights of all
plants on the area to air dry weights.
There must be no additions or
deletions in the moisture indicator plant list during the duration of the
study.
Moisture indicator plants are listed in the Results and Discussion
section of this report.

Pre-treatment

Deer, Elk, and Cattle Use Measurements

Accumulated deer and elk pellet groups were removed from all pellet plots
on September 3, 1970. Groups deposited during the winter of 1970-71 will
be counted in May, 1971.
When plots were cleared a record was kept of the number of cow chips in each
plot in order to provide an index to cattle use. Only those chips which
could be identified as having been deposited during the summer of 1970 were
counted.

Literature

Review

A review of recent literature was made to determine the most effective and
practical chemical herbicide for spraying Gambel oak, and the most suitable
plant species for use in reseeding the Hightower Mountain Oak Control Study
Area.
Advice was solicited from a number of professional people experienced
in chemical spraying and range reseeding.

RESULTS

Pre-treatment
The Weight

Estimate

AND DISCUSSION

Vegetation

Measurements

Technique

Observers became very proficient in estimating weights of current annual
growth of shrub vegetation.
In all cases average observer weight estimating
error was less than 10 percent, and in most cases less than 5 percent
(Table 1). Individual observer estimates averaged high on Some species and
low on others, but the usual tendency seemed to be for all or most observers
to estimate either high or low on the same species.
Close correlations were found between estimated and actual weights for all
observers on all five browse species sampled.
Correlation coefficients of
estimated to actual weights ranged from 0.79 to 0.95 and averaged 0.88
(Table 1). Average correlation coefficients by species were Gambel oak,
0.93; serviceberry, 0.90; chokecherry, 0.88; snowberry, 0.87; and big sage,
0.82.
These correlation coefficients closely approximate those reported by
Shoop and McIlvain (1963) which averaged 0.87 and ranged from 0.74 to 0.96.
The main difference, however, is that the Shoop and McIlvain (1963) data
were based on weight estimates of grasses and forbs rather than shrubs.
A relatively low number of samples were needed by each observer to estimate
his mean weight estimating error for each shrub species within 10 percent of
his true mean error at the 95 percent confidence level. The required number
for anyone
observer and any species ranged from 7 to 36 and averaged 18
(Table 1).

�-33-

Ideally, the size of individual estimated and clipped samples taken
for a shrub species should cover the entire range of weights of that
species that could be expected to occur in the meter square vegetation
measurement plots.
This was not always possible for two reasons: (1)
The maximum weights of each shrub species that occurred in meter square
plots were not known until all meter square plot measurements were completed.
(2) The size of estimated and clipped samples was governed by
the amount of time that could be allotted for clipping.
Clipping to
determine actual weights was very time consuming especially on big
sage, snowberry and serviceberry.
Thus, the size of individual estimated and clipped samples must be established beforehand, through
consideration of the maximum green weight of each species expected in
meter square plots, and the amount of time available for clipping.
The percentages of meter square plots in this study where the five
major shrubs occurred, and where weight of a particular species exceeded
the range of estimated and clipped samples was 1 percent for Gambel oak,
12 percent for serviceberry, 10 percent for chokecherry, 16 percent for
snowberry, and 17 percent for big sagebrush.
Because of the close
correlations between estimated and clipped weight data (Table 1), however, it was believed that the data are adequate to provide a basis
for correcting observer weight estimating error.

Measurement

of Vegetation

in Meter Square Plots

Plot Measurement Time--The average time required to measure vegetation
in a meter square plot based on all 600 plots was 50 minutes.
Including
time for walking from the road to a plot area and for traveling between
plots an average of 6 to 7 plots could be measured per individual per 8
hour day. In a plot which supported fairly heavy stands of the three
plant types about half of the measuring time was required for clipping
grasses and forbs, and about half for estimating weights of shrubs.
Conversion of Green Weights to Air Dry Weights--Moisture
indicator plants
were classified on the basis of their relative moisture contents into
categories ranging from dry to very wet for the three major plant types;
forbs, grasses and shrubs (Tables, 2, 3 and 4). Moisture contents were
expressed as air dry weight per 100 grams of green weight (ADW!lOOg GW),
derived by dividing the air dry weight of a sample by the corresponding
green weight.
Green weights of moisture indicator species encountered in meter square
plots were converted to air dry weights by multiplying by the ADW!lOOg
GW value established for that species during the ten day interval in
which the plot was measured.
It was also necessary to convert green
weights of non-moisture indicator plants encountered in meter square plots
to air dry weights, however, it was impossible to collect periodic moisture
content samples of all species on the area due to time limitations.
Thus,
non-moisture indicator plants were assigned to the various categories
described in Tables 2, 3 and 4 by comparing the texture and fleshyness
of each species with the moisture indicator plants in each category.
Green
weights of non-moisture indicator species were then converted to air dry
weights by multiplying by the mean ADW/lOOg GW value established for the
corresponding category during the ten day interval in which the plot was
measured.

�-34-

Moisture category assignments of non-moisture indicator plants are
shown in Table 5. These assignments must remain permanent for the
remainder of the study. Any new plants encountered on the study area
should be assigned to the appropriate category.
It is expected that
the percentage moisture content of the various categories will
fluctuate from year to year because new ten-day moisture content samples
must be collected for each moisture indicator species every time posttreatment vegetation measurements are made.

Forage Production by Weight--Plant species collected on the Hightower
Mountain oak control Study area are listed in Table 6. Not all of these
species were encountered while measuring vegetation, however.
Weights of vegetation in each habitat improvement unit as projected from
meter square plot measurements are shown in Tables 7 through 16. The 8
units were relatively homogeneous in total amount of vegetation produced.
Total air dry weights of forage ranged from 1621 lbs./ac. in unit 4 to
1885 lbs./ac. in unit 3, and averaged 1717 lbs./ac.
Percent composition
by air dry weight of·forbs, grasses and· shrubs also varied relatively
little among the 8 units. Forb composition among units ranged from 22
to 40 percent and averaged 31 percent or 532 lbs./ac.
Composition of
grasses varied between 11 and 33 percent with an average of 22 percent
and weight of 376 lbs./ac.
Shrub composition ranged from a low of 36
to a high of 59 percent and averaged 47 percent or 810 pounds of air dry
forage per acre.
Forage production here was somewhat higher than reported for the Gambel
oak vegetation type by several other workers.
Moinat (1956) found between 179 and 470 pounds of air dry grass forage and 28 to 322 lbs./ac.
of forb herbage on grazed Gambel oak range in southwestern Colorado.
The lower yields were found under oak clumps and the higher yields were
recorded in grass parks between oak stands.
Moinat (1956) also found
198 pounds of dry shrub forage per acre growing up to a height of 4 feet
in oak clumps.
Jefferies (1965), also working on grazed Gambel oak range
in southwestern Colorado reported 144 pounds of dry grass and 110 pounds
of dry forb forage per acre in open oak type parks, and 110 pounds of grass
and 80 pounds of forb forage per acre growing under the oaks. Brown (1958)
reported 346 pounds of air dry grass forage per acre in the Gambel oak
type of west-central Colorado.
The main vegetational differences among habitat improvement units were
in species composition within forb, grass and shrub categories.
Some
units supported more of a particular species than others, but the lack
of a particular species in a unit was offset by abundance of other species
within the same category whether forbs, grasses or shrubs.
Gambp.l oak,
the most abundant shrub varied from 79 lbs.1 ac , air dry weight in unit 2
to 540 lbs./ac. in unit 3, and averaged 255 1bs./ac. on all units.
It is
important to remember, however, that these data represent only vegetation
which occurred within 5 feet above the ground.
On the basis of visual
observation units 2 and 3 appeared more similar in oak production than
the data indicate.
In unit 2, however, Gambel oak trees were larger, and
less of the oak vegetation grew within 5 feet of the ground.
This was

�-35-

also the case in several other units. Production of Gambel oak here
was considerably higher than in northern Arizona, where Reynolds,
Clary and Ffolliott (1970) measured 90 pounds of oak f orage rpe r acre
up to a height of 7 feet.
.
Next to Gambel oak the most common shrub species were snowberry, which
averaged 243 lbs./ac. air dry weight in the 8 units; serviceberry, 107
lbs./ac.; chokecherry, 93 lbs./ac.; and big sage, 54 lbs./ac. The most
common forbs were lupine (Lupinus argentues), 135 lbs./ac.; aspen peavine
(Lathyrus leucanthus), 90 lbs./ac.; various species of Aster and Erigeron,
80 lbs./ac.; western yarrow (Achillea lanulosa), 30 lbs./ac.; Fendler
meadowrue (Thalictrum fendleri), 20 lbs./ac.; bedstraw (Galium sp.), 19
lbs./ac.; Ligusticum sp., 17 lbs./ac.; chickweed (Stellaria sp.), 12 lbs./
ac.; and thistle (Circium canescens), 11 lbs./ac.
Kentucky b Luegrass (Poa Eratensis) was by far,thelIl?,st"eommongrass
averaging 287 p~ou114sof air dry forage per\acr~'~(:'A1:though
all species
of Poa were lumped under the generic name when vegetation measurements
were made, and are designated generically in the tables in this report,
species of Poa other than pratensis probably contributed less than one
pound per acre to the total weight. Moinat (1956) and Jefferies (1965)
also mention Kentucky bluegrass as the most common grass in the Gambel
oak type. Brown (1958), however, reported 25 to 204 pounds of elk sedge
(Carex geyeri) and only 52 to 137 pounds of Kentucky bluegrass. He found
the highest production of elk sedge under oak clumps and Kentucky bluegrass was most abundant in openings. On the Hightower area, however,
Carex produced less than 0.01 pounds of dry forage per acre (Table 15).
Various species of wheat grasses (Agropyron ~.) were second in abundance
to Kentucky bluegrass producing 48 lbs./ac. Production of nodding brome
(Bromus anomalus) averaged 15 lbs./ac., and needlegrasses (Stipa sp.)
produced 11 lbs./ac.
Pre-treatment Cattle Use Measurements
Cattle use was unevenly dispersed among ,the 8 habitat improvement units.
By f~r the most use occurred 'in unit'8~hich had 18:26 cow days. use per
acre (Table 17). No use was recorded in units 3 and 4 .
.Literature Review
Chemical Herbicide Selection
The herbicide, 2,4,5-TP (Silvex) was selected for the spraying treatment.
It will be applied by helicopter at a rate of 2 pounds per acre mixed with
4 gallons of water and 1/2 gallon of number 2 diesel fuel per acre. A
liquid detergent soap will be added at a rate of 0.5% of the total spray
and carrier volume to facilitate mixing of the water, chemical and diesel
fuel. This chemical and application rate was recommended for use on Gambel
oak by Dunster (1970), Heikes (1970), and Marquiss (1970).

�-36-

Silvex was tested on Gambel oak near Rifle, Colorado, by Amchem Products, Inc. and compared with seven other herbicides.
Four of these
herbicides were actually mixtures of 2,4,5-T; 2,4,5-TP; 2,4-DP and
dicamba.
The other three were 2,4,5-T; 2,4-DP; and Amchem Brushkiller
170. After 2 years, 2,4,5-TP kill on oak was exceeded by only one other
chemical and that happened to be 2,4,5-TP with an additive which is not
commercially available (Dunster 1970).
In comparisons of 2,4,5-T, 2,4-5TP, and 2,4-DP, by Marquiss and Norris (1967 and 1968), 2,4,5-TP at the
2 pound per acre rate has provided the best defoliation of Gambel oak.
Lillie (1963) found 2,4,5 -TP to be generally slightly superior to
2,4,5-T in controlling Arizona chaparral, primarily turbinella oak
(Quercus turbinella).
In a test of 2,4,5-T oil soluble amine, 2-4-5TP and 2,4,5-T iso-octyl ester applied at a rate of 2 lbs./ac. with the
Stull Bi-fluid method, however, Pearl (1965) found better top-kill and
less sprouting on Gambel oak with 2,4,5-T amine.
The herbicide 2,4-D has been found relatively ineffective in controlling
most oak species.
Tests of 2,4-D, 2,4,5-T and ammonium sulfamate on 14
species of oak showed ammonium sulfamate to be most effective, 2,4,5-T
second and 2,4-D least effective (Anom. 1961). Ammonium sulfamate was
not considered for use on Hightower Mountain because it is a non-selective
herbicide.
It is used extensively to kill all vegetation (grasses, forbs
and shrubs) growing along roadways.
Several other herbicides which are more effective than 2,4,5-TP for controlling oak were considered but not accepted for use during this study
for various reasons.
Bovey, et al. (1969) reported that picloram (also known as tordon) was a
superior herbicide for controlling live oak (Quercus virginiana) in south
Texas when compared with 2,4-DP, 2,4,5-T or 2,4,5-TP.
Marquiss (1968 and
1969) found picloram or tordon superior to these same three chemicals for
controlling Gambel oak in southwestern Colorado.
Picloram was eliminated
from consideration for use on Hightower Mountain, however, because: (1)
It is retained in the soil from one to several years.
(2) It is not
licensed for use on croplands, and rangeland is considered a crop (Heikes
1970).
(3) The cost is prohibitive.
Tordon 212 applied at a 2 lb. lac.
rate would cost approximately
$40.00 per acre for the spray material.
Fenuron (3-phenyl - 1, 1 - dimethylurea) has been successfully used to
control turbinella oak (Wagle and Schmutz 1963), and Lillie (1962).
Davis and Pase (1969) found fenuron more effective than picloram (tordon)
on turbinella oak and Palmer oak (Quercus chrysolepsis) in Arizona.
Fenuron was rejected for use on this study, however, because no studies
could be located in the literature which had evaluated the effects of
fenuron on Gambel oak, and because fenuron is a soil sterilant.
Monuron was found equally effective as fenuron on turbinella oak at the
8 lb./ac. rate but inferior to fenuron at lower rates by Lillie (1962).
Monuron was found quite effective on five oak species in Iowa by Brinkman
(1959).
Brinkman (1959), however, indicated that areas treated with
monuron remained void of plant cover of any kind for 6 to 7 years, and

�-37-

that materials for the cheapest concentration used cost about $190 per
acre. Thus, monuron was eliminated from consideration for use on Hightower Mountain.
The second week of July was the time decided upon for spraying on the
Hightower Mountain Oak Control Study Area. This period was recommended
by Marquiss (1970), because carbohydrate storage in the roots is increasing rapidly at that time, and this would facilitate better movements
of the herbicide within the plant (Marquiss,1969). July 9, 1969 applications of 2,4-DP, 2,4,5-T, 2,4,5-TP, and a mixture of 2,4,5-TP and
2,4,-DP resulted in better Gambel oak control than August 23, 1969 applications on an area near Crawford, Colorado (Dunster, 1970).

Selection of Species for Reseeding
Species and seeding rates that have been selected for reseeding the
Hightower Mountain Oak Control Study Area are shown in Table 18. The
recommendations of Plummer, Christensen and Monsen (1968) and Plummer
(1970) were closely followed in making these selections, because their
work represents the most comprehensive effort to date on the subject of
reseeding western big game ranges. Most of the species selected have been
tested by others and found suitable for reseeding in the Gambel oak or
"mountain brush" vegetation type.
Crested wheatgrass (Agropyron cristatum) has been recommended for reseeding
in the mountain brush type by Doran (195lb), Hull et a1. (1958), Marquiss
and Everson (1969), and Plummer et al. (1955). It was recommended for the
nearby pinyon-juniper and sagebrush types by Doran (195la), and for the
ponderosa pine zone, which also borders on the mountain brush zone, by
Hull and Johnson (1955).
Smooth brome (Bromus inermis) is recommended for the mountain brush type
by Bleak (1968), Doran (195lb), Hull et al. (1958), Marquiss and Everson
(1969), and Plummer et al. (1955). Doran (195la) recommended smooth brome
for the sagebrush and pinyon-juniper types, and it was found to produce
well in the ponderosa pine type by Hull and Johnson (1955).
Those recommending intermediate wheatgrass (Agropyron intermedium) include
Bleak (1968), Doran (195lb), Hull et al. (1958), Marquiss and Everson (1969),
McGinnies et al. (1963), and Plummer et al. (1955). Hull and Johnson (1955)
also recommended it for the ponderosa pine zone.
Three studies reported orchard grass (Dactylis glomerata) could be successfully reseeded in the mountain brush zone, Doran (195lb), Hull et al. (1958),
and Plummer et al. (1955).
Sheep fescue (Festuca ovina) has been approved for the mountain brush zone
by Hull et al. (1958), and Plummer et al. (1955), and for the ponderosa pine
zone by Hull and Johnson (1955).
Various varieties of alfalfa (Meticago sativa) have been recommended for
reseeding in the mountain brush or Gambel oak type. Marquiss and Everson
(1969) recommended Nomad, Rambler and Ladak. Ladak has been recommended
by Bleak (1968), Hull et al. (1958), and Plummer et al. (1955). Hull and

�-38-

Johnson (1955) suggested that alfalfa be reseeded in the ponderosa
pine zone but made no reference to a particular variety.
Of the remaining forbs yellow sweet clover (Melilotus officinalis)
has been recommended by Plummer et al. (1955), and chickpea milkvetch
(Astragalus cicer) was recommended by Bleak (1968), and Plummer et al.
(1955). Chickpea milkvetch was mentioned for reseeding in the sagebrush and pinyon-juniper types by Doran (195la).
Relatively little research has been done on shrub reseeding in the
mountain brush zone except for that by Plummer, Christensen and Monsen
(1968), who recommended bitterbrush (Purshia tridentata) and mountain
mahogany (Cercocarpus montanus). Bitterbrush was also suggested by
Plummer et al. (1955).

LITERATURE CITED

Anon.

1961. Chemical control of brush and trees.
Farmers Bull. 2158. 23pp.

U.S. Dept. Agr.

Bleak, A. T. 1968. Growth and yield of legumes in mixtures with
grasses on a mountain range. J. Range Mgmt. 21(4):259-261.
Bovey, R. W., S. K. Lehman, H. L. Morton, and J. R. Baur. 1969.
Control of live oak in south Texas. J. Range Mgmt. 22(5):315-318.
Brinkman, K. A. 1959. Killing oakbrush a tricky business.
For. Expt. Sta. Tech. Pap. 165. 9 p , December.

Cent. States

Brown, Harry E. 1958.
39(2):317-327.

Ecology.

Gambel oak in west-central Colorado.

Davis, Edwin A., and Charles P. Pase. 1969. Selective control of
brush on chaparral watersheds with soil applied fenuron and
picloram. U.S.D.A. For. Servo Rocky Mtn. For. and Range Expt.
Sta. Res. Note. RM-140. 4 p.
Doran, Clyde W. 1951a. Reseeding trials in a sagebrush-pinyon-juniper
area in western Colorado. U.S.D.A. For. Servo Rocky Mtn. For. and
Range Expt. Sta. Res. Note No.9.
4 p.
1951b. Guide for reseeding summer rangelands on Colorado's
western slope. U.S.D.A. For. Servo Rocky Mtn. For. and Range Expt.
Sta. Paper No.6.
18 p.
Dunster, Ken W. 1970. Research and Development, Agricultural Chemicals
.Division, Amchem Products, Inc., Ambler, Pa. Personal Communication.
Harrington, H. D. 1954. Manual of the plants of Colorado.
Denver, Colorado. 666 p.

Sage Books,

�-39-

Heikes~ Eugene P~ 1970. Extension Professor of Botany and Plant
Pathology, Weed Laboratory, Colorado State University, Ft.
Collins. Personal Communication.
Hull, A. C. Jr., D. F. Hervey, Clyde W. Doran, and W. J. McGinnies.
1958. Seeding Colorado rangelands. Colo. State Univ. Expt.
Sta. Bull. 498-S. 46 p.
Hull, A. C. Jr., and W. M. Johnson. 1955. Range seeding in the
ponderosa pine zone in Colorado. U.S.D.A. Circ. No. 953. 40 p.
Jefferies, Ned W. 1965. Herbage production on Gambel oak range
in southwest Colorado. J. Range Mgmt. 18(4):212-213.
Kelsey, Harlan P., and William A. Dayton. 1942. Standardized
plant names. J. Horace Mc Farland Co., Harrisburg, Penn.
2nd Ed. 675 p.
Kufeld, Roland C. 1970. Experimental improvement of oakbrush on
deer, elk and cattle ranges - Hightower Mountain. Colo. Div.
of Game, Fish and Parks. P-R Project W-lOl-R-12, WP-4, J-3,
Game Research Rept. July. pp. 113-126.
Lillie, D. T. 1962. Control of shrub live oak (Quercus turbinella
Greene) with granular and pelleted herbicides. Western Weed
Control Conf. Res. Prog. Rept. p. 16.
1963. Control of Arizona chaparral with 2,4,5-T and Silvex.
J. range Mgmt. 16(4):195-199.
Marquiss, Robert W. 1968. Progress in chemical control of Gambel's
oakbrush rangelands. In Papers of the Ann. Meet. Am. Soc. Range
Mgmt. Albuquerque, New Mex. pp. 28-29.
1969. Studies on Gambel's oak at the San Juan Basin Station.
Colo. State Univ. Exp , Sta. Prog. Rept. PR 69-38. 2 p ,
1970. Supt. San Juan Basin Branch Expt. Sta., Hesperus. Colo.
State Univ., Fort Collins. Personal Communication.
Marquiss, R. W., and A. C. Everson. 1969. Performance tests of dryland pasture grasses and alfalfa at the Hesperus Station. Colo.
State Univ. Exp. Sta. Prog. Rept. PR 69-42. 3 p.
Marquiss, R. W., and J. J. Norris. 1967. Management and improvement
of Gambel oakbrush ranges. Colo. State Univ. Exp. Sta. Prog.
Rept. PR 231. 2 p.
and
ranges.

1968. Management and improvement of Gambel oakbrush
Colo. State Univ.Exp. Sta. Prog. Rept. PR 68';"7, 4 p ,

McGinnies, W. J., D. F. Hervey, J. A. Downs and A. C. Everson. 1963.
A summary of range grass seeding trials in Colorado. Colo. State
Univ. Expt. Sta. Tech. Bull. No. 73. 81 p.

�-40Moinat, A. D. 1956. Comparative yields of herbage from oak shrub
and interspersed grassland in Colorado. Ecol. 37:852-854.
Pearl, Robert W. 1965.
Weed Control Conf.

Gambel oak control in New Mexico.
Res. Prog. Rept. p. 40.

Western

Pechanec, J. F., and G. D. Pickford. 1937. A weight estimate method
for the determination of range or pasture production. J. Am.
Soc. Agron. 29:894-904.
--

Plummer, A. Perry. 1970. Range Scientist, U.S.D.A. For. Servo
Intermtn. For. and Range Expt. Sta., Ephraim, Utah. Personal
Communication.
Plummer, A. Perry, Donald R. Christensen and Stephen B. Monsen. 1968.
Restoring big game range in Utah. Utah Div. of Fish and Game.
Pub. No. 68-3. 183 p.
Plummer, A. Perry, A. C. Hull, Jr., George Stewart and Joseph H.
Robertson. 1955. Seeding rangelands in Utah, Nevada, southern
Idaho, and western Wyoming~ U.S.D.A. For. Ser. Agric. Handbook
No. 71. 71 p ,
Reynolds, Hudson G., Warren P. Clary, and Peter F. Ffolliott. 1970.
Gambe1 oak for southwestern wildlife. J. For. 68(9):545-547.
Shoop, M. C., and E. H. McIlvain. 1963. The micro-unit forage
inventory method. J. Range Mgmt. 16(4):172-179.
Wagle, R. F., and E. M. Schmutz. 1963. The effect of fenuron on
four southwestern shrubs. Weeds. 11:149-157.

Prepared by

t/drud C~/ 10t!14

Roland C. Kufeld
Wildlife Researcher

�I
\}
\

Fig. 1. General study area map showing locations of habitat improvement

units.

�-42-

UNIT 1
BOUNDARY MAP

388'

412'

1

19

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1

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498'

18

-

5:

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15
~
17
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2221

~16

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v

(Y)

Z

2

14

°

f'..

0
r-,

C\J

V

CONTROLLED BURN
BOUNDARY A:63·5 Acres
B:30·0 Acres

~

°

"t

(Y)"

C\J

2

16

"t

270'

C\J
C\J

N
&lt;.J)

13

LD

3 56'
15

S66
12

260'
~
(Y)

0

v

3 3
11 .

"r

0
(Y)

~

&lt;D

267'

LD
LD

r-.

263'

4

5

..-"t

6
~

N66°E

Q)

C\J

4

5

6

26 8'

~

7

268'
Q)

W

(Y)

(Y)

(Y)

°
~

0

0
~

0

co
,',

o 100

,

300
feet

J

500

8

..--

5:

If)

7

0

(Y)

C\J

520'

"t

0

9

~

C\J

z

o
(Y)

11
&lt;0
(X)

8

10

..--

534'

Fig. 2. Boundary map of habitat improvement unit number 1.
SEPT

RAF

19/0

�-4j-

UNIT

1·

PELLET PLOT MAP

20

1

N28°W,151'
from post
1-A-1I2

19

16

1

Start stk. to plot 1 = 67 ft.
Distance between plots = 100 ft.
Line bearing = N 6 6°E
P-1-8

18

P-1-8

L

LLtotal

2
15
S24°E, 280
from post
1-A-12

S24°E, 205
from post
1-A-3
S24°E, 280
from post
1-A-3

,

17

16

2
P- 2-10

14

13

15
12

, 3 3

11

14

P-3 -16
P-4,..15

,
5

4

6

/
-

10
4

plots

line number
pellet line

13

5
12

7

6
S24°E,153'
from post
1-A-6

P-5 ':'11

7
9

8

11

o 100
Fig.

300
feet

500

8

9

10

J. Pellet plot map of habitat improvement ur~t number 1.
S£PT 1970
RAF

�-44-

UNIT

1

VEGETATIVE PLOT MAP

20
1r-------------~----~19

1

16

S24°E , 132'
from post -- ~
1-8-1
2

Start stk. to plot 1 = 22 ft._
D'IStance between plots = 50 ft.
V-3 -20
Ltotal plots
line number
vegetative line

.:L

18

N656E
"

V-1-10
15
17

S24°E,150'
from post"
1-8-lm2
S24°E,247'
from post
1-8-2

16

2
E-+ 14
&lt;, N65°E ~1(-N67°

13

12

V-2-10
.?'

V-3-20

3 3/

N65°E

18·8 ft., not

N24°W,125'
from post -- ~
1-8-4

22ft.
11

14

N66°E
V-4-15

5
S24°E,97'
4
-------?
from post - - 1-8-6
4
5

6

6

N65°E
V-5-10

10

7

S24°E,105'
-- ~
from post1- 8-7
7

N66°E
V-6-10

8

9
11

o 100

300
feet

500

8

9

1

Fig. 4. Vegetation measurement plot map of habitat improvemen~ unit number 1.
SEPT

1')7,'

�-45-

Fig. 5. Ten permanent
improvement unit.

Fig. 6.

Equipment

photo points were established

needed for measuring

vegetation.

in each habitat

�For~ Designed Dec. 8, 1969

-46-

HIGlfl'OHEn

Habitat

Improvement

l'10TlNfAIN
VEGETA'.l'IVE l·ll,:t\SU1U~NENT FIELD

Unit No.

Line No.

Time Start

Observer

Plot No.

--------------------

Plant

Species

"Time End

FOmi

--------------------------Date

-----------------------

------~-----Tota 1 TiTI)c ---------------~
Green
Height

Adjusted
Green Height

Grasses

Forbs

Br ows e

Fig. 7. Hightower Mountain vegetation measurement field data form.

Air

Dry

Weight

�Table 1.

Observer proficiency in estimating current annual growth green weights on five shrub species.
Serviceberry

Chokecherry

Snowberry

Big Sage

19
No. samples needed 1/
67
No. samples taken
135g
act. wt. of samples
570 g
Size of largest sample
8,751 g
Tot. est. wt. of samples
9,017 g
Tot. act. wt. of samples
-2.9%
% error est.-act. wt. 2/
.94
r value est.-act. wt. - A
1.0304
Error correction factor (R) 11

19
32
60 g
121 g
2,009 g
1,935 g
+3.8%
.89
.9632

14
47

182 g
5,205 g
5,170 g
+0.1%
.86
.9933

13
56
57 g
107 g
3,432 g
3,202 g
+7.2%
.85
.9330

33
59 g
152 g
1,962 g
1,953 g
0.0%
.83
.9954

Observer
No. 2

No. samples needed 11
26
80
No. samples taken
121 g
x act. wt. of samples
410 g
Size of largest sample
9,444 g
Tot. est. wt. of samples
9,678 g
Tot. act. wt. of s amp les
-2.4%
% error est.-act. wt. 21
.88
r value est.-act. wt. - A
1.0248
Error correction factor (R) 1/

28
46
53 g
121 g
2,450 g
2,454 g
0.0%
.85
1.0016

17
55
103 g
182 g
5,302 g
5,662 g
-6.4%
.86
1.0679

36
70
51 g
107 g
3,834 g
3,596 g
+6.6%
.79
.9379

15
47
59 g
152 g
2,655 g
2,775 g
-4.3%
.86
1.0452

Observer
No. 3

21
No. samples needed ~/
81
No. samples taken
127 g
x act. wt. of samples
570 g
Size of largest sample
10,202 g
Tot. est. wt. of samples
10,248 g
Tot. act. wt. of samples
0.0%
% error est.-act. wt. 2/
.94
r value est.-act. wt. - A 3
.1.0045
Error correction factor (R) _I

10
46
53 g
121 g
2,608 g
2,454 g
+6.3%
.95
.9410

7

9

55
103 g
182 g
5,752 g
5,662 g
+1.6%
.94
.9844

70
51 g
107 g
3,897 g
3,596 g
+8.4%
.94
.9228

19
47
59 g
152 g
2,854 g
2,775 g
+2.8%
.81
.9723

Oak

Observer
No.1

x

no g

9

----------------------------------------------------------------------------------------------------------

,

.j:--

,

-....J

�Table 1. Observer
(continued).

proficiency

in estimating

current

annual growth green weights

Oak

Serviceberry

Chokecherry

Snowberry

Big Sage

18
46
53 g
121 g
2,500 g
2,454 g
+1.9%
.91
.9816

14
55
103 g
182 g
5,329 g
5,662 g
-5.9%
.88
1.0625

20
70
51 g
107 g
3,723 g
3,596 g
+3.5%
.88
0.9659

18
47
59 g
152 g
2,629 g
2,775 g
-5.3%
.80
1.0555

Observer
No.4

No. samples needed l/
No. samples taken
x act. wt. of samples
Size of largest sample
Tot. est. wt. of,samples
Tot. act. wt. of samples
% error est.-act. wt. 2/
r value est.-act. wt. - A
Error correction factor (R)

Observer
No. 5

No. samples needed 24
No. samples taken
81
x act. wt. of samples
127 g
Size of largest sample
570 g
Tot. est. wt. of samples
10,233 g
Tot. act. wt. of samples
10,248 g
% error est.-act. wt. 2/
0.0%
r value est.-act. wt. - A
.93
1.0015
Error correction factor (R) 1/

16
81
127 g
570 g
9,790 g
10,248 g
-4.5%
.95
1/
1.0468

I

1/

l/ Number

22
46
53 g
121 g
2,471 g
2,454 g
+0.1%
.89
.9931

of samples needed by an observer to estimate
of his true mean error at the 95% confidence level.

~/ Correlation

1/ Correction
observer

coefficient

of estimated

factor used to adjust
weight estimating error.

on five shrub species

his weight

17
55

103 g
182 g
5,649 g
5,662 g
0.0%
.86
1.0023

18
70
51 g
107 g
3,739 g
3,596 g
+'1·.0%
.89
.9618

17
46
58 g
2,668 g
2,683 g
-0.1%
.82
1.0056

error on each shrub species within

meter square vegetation

measurement

I

l52g

and actual weights.

each observer's

+=-ex&gt;

plot data for

10%

�-49-

Table 2. Permanent forb moisture content categories and moisture contents
of indicator plants collected in 1970.
Mean ADW/IOO g GW
Mean ADW/100 g OW
of 3 Samples
of 9 Samples
on Each Collection Date
(SununerMean)
7-14-70 7-24-70 8- 7 -70

Forb
Moisture
Category

Moisture
Indicator
Plants

Dry

Grindelia aphanactis

37.5

40.8

42.6

40.3

Lupinus argentues

30.9

36.2

50.7

39.3

Category mean 1/

34

38

47

Lathyrus 1eucanthus

29.5

35.6

40.3

35.1

Achillea 1anu1osa

30.3

34.0

38.2

34.2

Tha1ictrum fendleri

30.3

32.6

37.8

33.6

Iris missouriensis

29.0

31.3

32.9

31.1

Galium boreale

31.4

27.8

33.3

30.8

Agastache urticifolia

23.3

31.7

35.1

30.0

Cirsium canescens

24.2

27.8

31.6

27.9

Ligusticum porteri

23.5

26.3

33.1

27.6

Category mean 1/

28

31

35

He1enium hoopesii

23.1

24.0

27.8

25.0

Rudbeckia montana

22.2

22.6

28.8

24.5

Category mean 1/

23

23

28

Swertia radiata

19.5

21.1

20.4

Category mean 1/

19

21

20

Moderate

Wet

Very Wet

l/ Mean of all species in the category.

20.3

�-50-

Table 3. Permanent grass moisture content categories
of indicator plants collected in 1970.
Grass
Moisture
Category

Dry

Moisture
Indicator
Plants

Agropyron

Mean ADW/IOO g GW of 3 Samples
On Each Collection Date
7-14-70
7-24-70
8-7-70

smithii

contents

Mean ADW/IOO g GW
of 9 Samples
(Summer Mean)

39.9

45.0

48.8

44.5

35.3

42.2

49.0

42.2

38

44

49

Elymus glaucus

30.4

35.3

41. 2

35.7

Bromus

30.2

36.2

37.7

34. 7

30

36

39

Poa pratensis
Category
Wet

and moisture

1:./

Mean

anomalus

Category

1:./

Mean

1:./ Mean of all species in the category.

Table 4. Permanent
of indicator plants

Shrub
Moisture
Category

Dry

Moisture'
Indicator
Plants

Amelanchier
Quercus

alnifolia

gambel1ii

Artemisia

Wet

shrub moisture content categories
collected in 1970.

tridentata

and moisture

contents

Mean ADW/IOO g GW of 3 Samples
On Each Collection Date
7-14-70
7-24-70
8-7-70

Mean ADW/IOO g GW
of 9 Samples
(Summer Mean)

46.4

51. 2

53.9

50.5

45.2

48.5

50.7

48.1

44.7

45.6

50.3

46.9
46.9

Symphoricarpos

sp.

43.8

47.9

49.1

Category

Mean

1:./

45

48

51

Artemisia

cana

40.0

41.0

45.2

42.1

Ch rys othamnus
vicidiflorus

37.8

41.9

43.4

41.0

Prunus

38.4

38.8

39.9

39.0

Rosa nutkana

37.3

36.7

39.2

37.7

Ch rysothamnus
nauseosus

35.6

37.1

40.0

37.6

38

39

42

virginiana

Category

Mean

1:./

1:./ Mean of all species. in category.

�-51-

Table 5. Moisture content category assignments for plants encountered on
meter square plots in 1970, but for which no moisture content samples are
collected.

Type of
Plant

Moisture
Category

Forbs

Dry

"

"

"

"
"

"

"

"
"
"
"

"

"

"
"
"
"
"
"

"
"

"

"

"
"

"

"

"

"
"
II

"

"
"

"

"
"

"

"

"
"
"

"

"

Moderate

"
"

"

"

"

"

"

"
"

"

"

"
"

"
"
"
"

"
"

"
"
"
"

"

"
"
"
"
"
"

"

"

Symbol

Alac
Anse
Aradr
Arhi
Arco
Artdr
Arlu
Cabu
Cali
Chbe
Coum
Deri
Erum
Eppa
Giag
Lare
Lile
MagI
Mudi
Oriu
Pamy
Pecr
Pest
Poav
Podo
Popu
Aggi
Aster-Erigeron
Basa
Bere
Cagu
Dene
Deoc
Ditr
Fram
Gepa
Hale
Lifi
Mefr
Mopa
Phhe
Pofo
Rain

Scientific

Name

Allium acuminatum
Androsace septentrionalis
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia dracunculus
Artemisia ludoviciana
Capsella bursa-pastoris
Castilleja linariaefolia
Chenopodium berlandieri
Comandra umbel lata
Descurainia richardsonii
Eriogonum umbellatum
Epilobium paniculatum
Gilia aggregata
Lappula redowskii
Linum lewisii
Madia glomerata
Musineon divaricatum
Orthocarpus luteus
Pachistima myrsinites
Penstemon crandallii
Penstemon strictus
Polygonum aviculare
Polygonum douglasii
Potentilla pulcherrima
Agoseris glauca
All Aster and Erigeron Species
Balsamorhiza sagittata
Berberis repens
Calochortus gunnisonii
Delphinium nelsoni
Delphinium occidentale
Disporum trachycarpum
Fragaria americana
Geranium parryi
Hackelia leptophylla
Ligusticum filicinum
Mertensia franciscana
Moldavica parviflora
Phacelia heterophylla
Polemonium foliosissimum
Ranunculus inamoenus

----------------------------------------------------------------------------

�-52-

TableS.
Moisture content category assignments for plants encountered on
meter square plots in 1970, but for which no moisture content samples are
collected (continued).

Type of
Plant

Moisture
Category

Forbs

Moderate

"

"
"

"
"
"
"
"
"

"
"

"

"
"
"

"

"

"

II

"
"
"

"
"
"
"
"
"
"

"
"

Wet

"

"

"

"

"

"

"

"

"

"

"

Very Wet

Grasses

Dry

"

"
"
"
"
"

"

"
"
"
"
"
"
"
"
"

Wet

Shrubs

Dry

"

"

"

"

"

"
"
"

"

Symbol

Scientific

Seam
Sein
Sime
Smst
Sosp
Stellaria
Taof
Trpr
Urdi
Vebi
Vimu
Viru
Vish

Senecio ambrosioides
Senecio integerrimus
Silene menziesii
Smilacina stellata
Solidago sparsiflora
All Stellaria Species
Taraxacum officinale
Tragopogon pratensis
Urtica dioica
Veronica biloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii

Gehe
Hyfe
Lasc
Opuntia
Osmorhiza
Pegr
Rucr
Vaoc
Wyam

Gentiana heterosepala
Hydrophyllum fendleri
Lactuca scariola
Opuntia sp.
All Osmorhiza Species
Pedicularis grayi
Rumex crispus
Valeriana occidentalis
Wyethia amplexicaulis

Name

No Plants in Category
Agropyron
Brte
Feth
Kocr
Sihy
Stipa

All Agropyron Species
Bromus tectorum
Festuca thurberi
Koeleria cristata
Sitanion hystrix
All Stipa Species

Carex
Brin
Dagle
Hobr
Juba
Phpr

Carex sp.
Bromus inermis
Dactylus glomerata
Hordeum brachyantherum
Juncus balticus
Phleum pratense

Acgl
Cemo
Chde

Acer glabrum
Cercocarpus montanus
Chrysothamnus depressus

�-53-

Table 5. Moisture content category assignments for plants encountered on
meter square plots in 1970, but for which no-moisture content samples are
collected (continued).

Type of
Plant

Moisture
Category

Shrubs

Dry

"

"

II

Wet

"

II

Symbol

Crer
Putr
Rice

Scientific

Name

Crataegus erthroEoda
Purshia tridentata
Ribes cereum
No Plants

in Category

�-54-

Table 6. Plants collected on the Hightower
during the summers of 1969 and 1970.

Mountain

Oak Control Study Area

FORBS

Herbarium
Collection
Number 1/

K~25
K-137
K-138

K-27
K-14l
K-28

K-29
K-142
K-140
K-33
K-3l
K-33
K-34
K-35
K-36
K-37
K-38

K-39
K-40
K-4l

K-42
*K-43
K-145
K-144
K-47

K-48
~'&lt;K-49

K-90

Symbol

Acla
Agur
Aggi
Alac
Anse
Aqco
Aradr
Arhi
Arco
Arbi
Artdr
Arlu
AsterErigeron
Asha
Basa
Bere
Cagu
Cabu
Cali
Chbe
Cica
Coli
Coum
Dene
Deoc
Deri
Ditr

K-5l
K-50

AsterErigeron

K-52
K-53
K-54
K-55

K-56
K-57
~"K - 58

K-59

Erum
Eppa
Fram
Galiurn
Gehe
Gepa
Giag

2/

Scientific

Name

1/

Common

Name

!l../

Achillea lanulosa Nutt.
Agastache urticifolia Kuntz
Agbseris glauca (Pursh) D. Dietr.
Allium acuminatum Hook.
Androsace septentrionalis L.
Aquilegia coerulea James
Arabis drummondi Gray
Arabis hirsuta (L.) Scop.
Arenaria congesta Nutt.
Artemisia biennis Willd.
Artemisia dracunculus L.
Artemisia ludoviciana Nutt.
(Aster adscendens Lindl.
(Aster rubrotinctus Blake
Astragalus haydenianus A. Gray
Balsamorhiza sagittata (Pursh) Nutt.
Berberis repens Lindl.
Calochortus gunnisonii Watson
Capsella bursa-pastoris
(L.) medic.
Castilleja linariaefolia Benth.
Chenopodium berlandieri Moq.
Cirsium canescens (Nutt.)
Collomia linearis Nutt.
Comandra umbellata (L.) Nutt.
Delphinium nelsoni Greene
Delphinium occidentale (Wats.) Wats.
Descura in ia r ichard son ii (Swce t ) O. E. Schultz
Disporulil trachycarpulll(S.Wats.)Benth.&amp; Hook
(Erigeron flagellaris A. Gray
(
(Erigeron fornlosissium Greene
(
(Erigeron speciosus (Lindl.) DC

Oregon Fleabane

Eriogonulll umbellatum Torr.
Epilobiulllpaniculatum Nutt.
Fragaria americana (Porter) Britt.
Galillill
boreale L.
Gentiana heterosepala Engelrn.
Geranium parryi (Engclm.) Heller
Gilia aggregata (Pursh ) Spreng.

Sulfur Eriogonum
Autumn Willowweed
American Strawber~y
Northern Bedstraw
Annua l Cen tian
Parry Geranium
Skyrocket Gilia

Western Ya rrow
Nettleleaf Giant Hyssop
Pale Agoseris
Tapertip Onion
Rockjasmine
Colorado Columbine
Drummond Rockcress
Hairy Rockcress
Ballhead Sandwort
Biennial Wormwood
Tarragon
Louisiana Sage brush
Common Aster
Aster
Hayden Poisonvetch
Arrowleaf Balsamroot
Creeping Mahonia
Gunnison Mariposa
Shepherds Purse
Wyoming Paintcdcup
Pitseed Goosefoot
Thistle
Gilia
Common Comandra
Menzies Larkspur
Duncecap Larkspur
Richardson Tansey~ustard
Wartberry Fairybells
Trailing Fleabane
Fleabane

------------------------------------------------------------------------------------

�-55Table 6. Plants collected on the High-tower Mountain
during the summers of 1969 aud1970
(continued).
Forbs

Herbarium
Collection
Number II

K-60
K-44
K-6l
K-62
K-63

Symbol ~/

Oak Control Study Area

(Continued)

Scientific

Name

1/

Grap
Grindelia aphanactis Rydb.
Hale
Hackelia leptophyUa
(Rydb.) Johnston
Hebo
lielenium hoopesii A. Gray
Hyfe
Hydrophyllum fcndleri (Gray) Heller
Irmi
Iris lIlissouricllsisNutt.
K-64
Lasc
Lactuca scariola L.
K-45
Lare
Laprula redowskii (Hornem.) Greene
K-136
Lale
Lathyrus leucanLhus (Rydb.)
K-65
Lede
~id
ium dens iflorulIlSchrad.
K-46
·
{J
t'
(Ligusticum
filicinum S. Wats.
L l.g s acurn
K-66
(Ligusticum porteri Coult. &amp; Rose
K-67
Lile
Linum lewisii Pursh
K-68
Luar
Lupinus argentues Pursh
K-149
MagI
Madia glomerata Hook.
K-69
Mefr
Mertensia franciscana Heller
K-26
Mopa
Moldavica parviflora (Nutt.) Britt.
~'~K70
Mudi
Musineon divaricatum (Pursh) Nutt.
K-71
Oeca
Oenothera caespitosa Nutt.
No Specimen Opun t i a Opuntia sp.
Orlu
*K-72
Orthocarpus luteus Nutt.
K-73
(Osmorhiza chilensis Hook &amp; Arn.
Osmorhiza
K-74
(Osmorhiza occidentalis (Nutt.) Torr.
K-30
Pamy
Pac1~.~~timamyrsinites (Pursh) Raf.
K-75
Pegr
Pedicularis grayi A. Nels.
K-76
Pecr
Penstemon crandallii A. Nels.
K-77
Pest
Penstemon strictus Benth.
K-78
Phhe
Phacelia heterophylla Pursh
Ph10
K-79
Phlox 10ngifolia Nutt.
K-80
Pofo
Polemonium foliosissimum A. Gray
K-15l
Poav
Polygonum aviculare L.
K-147
Podo
Polygonum douglasii Greene
K-8l
Popu
Potentilla pulcherrima Lehm.
K-146
Rain
Ranunculus inamoenus Greene
K-152
Rucr
Rumex crispus L.
K-82
Rumo
Rudbeckia montana Gray
K-83
Seam
Senecio ambrosioides Rydb.
K-84
Sein
Senecio integerrimus Nutt.
K-85
Sese
Senecio serra Hook.
K-143
S ime
Silene lllenziesiiHook.
K-86
Smst
Smilacina stellata (L.) Desf.
K-87
Sosp
Solidago sparsiflora A. Gray
K-88
(g~]
a r i.:1j amC!s i.:1
n3 Torr.
Stellaria
~\-K-89
(8 t e.L1 a r i ,I J on \~i. [() Li a Huh 1.
K-91
Swra
S\vc::.rI:raciLlta
i.:1
(Ko Ll og g) Kuntze
K-92
Taof
l'.:lra&gt;s~:!.!.!
nffi.cin.JleHiggars
;'~K-93
Thfe
TlJillJ:.0rtl1l1fenclll~riEuge Lm ,
K-94
Trdu
]'r~~~(')()n dllb~
Scop.
No Specimen
Trpr
Tr[q~()pni?on
E.:1Lens is L.

Common

Name

~I

Gumweed
Stickseed
Orange Sneezeweed
Fendler Waterleaf
Rocky Mountain Iris
Prickly Lettuce
Sticksced
Aspen Peavine
Prairie Pepperweed·
Privet
Porter Ligusticum
Lewis Flax
Silvery Lupine
Cluster Tarweed
Franciscan Bluebells
Dragonhead
Musineon
Tufted Eveningprimrose
Pricklypear
Yellow Owlclover
Sweetroot
Sweetanise
Myrtle Pachistima
Grays Pedicularis
Crandall Penstemon
Rocky Mountain Penstemon
Varileaf Phacelia
Longleaf Phlox
Leafy Polemonium
Prostrate Knotweed
Douglas Knotweed
Beauty Cinquefoil
Buttercup
Curly Dock
Coneflower
Ragweed Groundsel
Lambstongue Groundsel
Butterweed Groundsel
Mensies Silene
Starry Solomonplume
Threenerve Goldenrod
Ch ickweed Ch ickweed
Showy Frasera
Common Dandelion
Fendler Me.:1dowruc
Sa Ls Lf y

Meadow

Sa Is ify

----------------------------------------------------

�-56-

Table 6. Plants collected on the Hightower Mountain
during the summers of 1969 and 1970 (continued).
Forbs

Herbarium
Collection
Number 1/

Symbol

K'-95
K-Y6
K-97
K-153
K-98
K-99
K-139
K-IOO

Urdi
Vaoc
Veam
Vebi
Vimu
Viru
Vish
Wyam

])

Oak Control

Study Area

(Continued)

Scientific

Name

1/

Urtica dioica L. var. procera (Muhl.)Wedd.
Valeriana occidentalis
Heller
Veronica americana Schwein.
Veronica biloba L.
Viguiera multiflora
(Nutt.) Blake
Viola rugulosa Greene
Viola sheltonii Torr.
Wyethia amplexicaulis
(Nutt.) Nutt.

Common

Name

!i/

Big Sting Nettle
Western Valerian
American Speedwell
Speedwell
Showy Goldeneye
Cheyenne Violet
Shelton Violet
Mulesears Wyethia

GRASSES

Herbarium
Collection
N um b er -1/

Symbol

2/

Scientific

Name

1/

K-101
A
(Agropyron smithi Rydb.
K-102
gropyron(Agropyron
trachycaulum
(Link) Malte.
K-103
Bran
Bromus anomalus Rupr.
K-104
Brin
Bromus inermis Leyss.
No Specimen
Brte
Bromus tectorum L.
K-154
Carex
Carex sp.
K-105
Dagl
Dactylus glomerata L.
K-I06
Eigi
Elyrnus glaucus Buckl.
K-1l5
Feth
Festuca thurberi Vasey
K-I07
Hobr
Hordeum brachyantherum
Nevski
K-108
Juba
Juncus balticus Willd.
K-I09
Kocr
Koeleria cristata (L.) Pers.
K-110
Drhy
Drzyopsis hymenoidcs
(R. &amp; S.) Ricker
x-r i i
Php r
Phleum pratense L.
K-1l2
(Poa ampia Herr.
K-113
Poa
(Poa canbyi (Scr ibn.) Piper
K-1l4
(Poa pratensis L.
K-1l6
Sihy
Sitanion hystrix (Nutt.) J. G. Smith
K-l17
(Stipa columbiana Macoun.
Stipa
K-1l8
(Stipa lettermani Vasey

Common

Name

4/

Bluestem Wheatgrass
Slender Wheatgrass
Nodding Brome
Smooth Brome
Cheatgrass Brome
Sedge
Orchardgrass
Blue Wildrye·
Thurber Fescue
Meadow Barley
Baltic Rush
Prairie Junegrass
Indian Ricegrass
Timothy
Big Bluegrass
Canby Bluegrass
Kentucky Bluegrass
Bottlebrush Squirreltail
Subalpine Needlegrass
Letterman Needlegrass

-------------------------------------------------------------------------------------

�-57-

Table 6. Plants collected on the Hightower Mountain
during the summers of 1969 and 1970 ·(continued).

Oak Control

Study Area

BROWSE

Herbarium
Collection
Number 1/

Symbol ~/

K-150
K-ll9
*K-120
K-12l
K-l22
K-123
K-148
K-124
*K-125
*K-126
K-127
No Specimen
K-128
K-129
K-130
K-13l
K-132
K-133
~"K-134
K-135

Acgl
Amal
Arca
Artr
Cefe
Cemo
Chde
Chvi
Chna
Crer
Loin
Potr
Pofr
Prvi
Putr
Quga
Rhsm
Rice
Ronu
Sym.

Scientific

Na~e

1/

Common Name

Acer glabrum Torr.
Amelanchicr alnifolia Nutt.
Artemisia canaPursh
Artemisia tridentata Nutt.
Ceanothus fendleri Gray
Cercocarpus montanus Raf.
Chrysothamnus depressus Nutt.
Chrysothamnus viscidiflorus
(Hook) Nutt.
Chrysothamnus nauseosus (Pallas) Britt
Crataegus erythropoda Ashe
Lonicera involucrata (Rich.) Banks
Populus tremuloides Mich.
Potentilla fruiticosa L.
Prunus virginiana L.
Purshia tridentata (Pursh) DC.
Quercus gambellii Nutt.
Rhamnus smithi Greene
Ribes cereum Dougl.
Rosa nutkana Presl.
Symphoricarpos sp.

!!../

Rocky Mountain Maple
Saskatoon Serviceberry
Silver Sagebrush
Big Sagebrush
'Fendl~r Ceanothus
True Mountain Mahogany
Dwarf Rabbitbrush
Douglas Rabbitbrush
Rubber Rabbitbrush
Cerro Hawthorn
Bearberry Honeysuckle
Aspen
Bush Cinquefoil
Common Chokecherry
Antelope Bitterbrush
Gambel Oak
Smith Buck thorn
Wax Gooseberry
Nootka Rose
Snowberry

1/ Plant

specimens are filed in Harold Shepherd's and Bert Baker's Herbarium in
Fort Collins.
The Herbarium collection numbers listed here designate specimens
that were collected as part of the Hightower Mountain Oak Control Study.

1/ Symbol used on vegetation
1/ Scientific
Colorado.

measurement

forms.

names are from Harrington, H. D. 1954.
Sage Books, Denver, Colorado.
666 pp.

Manual

of the Plants

!!../

Cornmon names are from Kelsey, Harlan P., and William A. Dayton.
Standardized Plant Names.
J. Horace McFarland Co., Harrisburg,
2nd Ed. 675 pp.

*

Specimens of all plants except those prefixed
H. D. Harrington.

of

1942.
Pennsylvania.

by * have been annotated

by

�-58-

Table 7.
Pre-treatment
Unit No. 1.

Type of
Plant

Forbs

Scientific

p roduc tLon of vegetation

Namell

Achillea 1anulosa
Agastache urticifo1ia
Agoseris glauca
Allium acuminatum
Androsace septentrionalis
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia dracunculus
Artemisia ludoviciana
Aster-Erigeron
Balsamorhiza sagittata
Berberis repens
Calochortus gunnisonii
Capsella bursa-pastor is
Castilleja linariaefolia
Chenopodium ber1andieri
Cirsium canescens
Comandra umbe11ata
Delphinium occidentale
Descurainia richardsonii
Disporum trachycarpum
Eriogonum umbel1atum
Epilobium paniculatum
Fragaria americana
Galium sp.
Gentiana heterosepala
Geranium parryi
Gil ia aggrega ta
Grindelia aphanactis
Hacke1ia 1eptophylla
Helenium hoopesii
Hydrophyllum fendleri
Iris missouriensis
Lactuca scariola
Lappula redowskii
Lathyrus leucanthus
Ligusticum sp.
Linum lc\visii
Lupinus argcntues
Madia glomerata
Mcrtcnsia franciscana
Musincon divarcatum
Opuntia sp.

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

1.64
0.00
0.04
0.22
0.00
0.01
0.00
0.31
0.00
0.02
5.78
0.03
0.00
0.09
0.01
0.19
0.01
.2.06
0.12
0.00
0.01
0.00
0.46
0.00
0.00
0.25
0.00
0.12
0.00
'0.55
0.00
0.01

O.Ol
0.00
0.00
0.00
2.61
0.09
0.86
10.85
0.00
0.00
0.11
0.61

27.00
0.00
0.71
3.69
0.00
0.24
0.00
5.12
0.00
0.36
94.81
0.48
0.00
1.42
0.24
3.09
0.12
33.78
2.02
0.00
0.24
0.00
7.49
0.00
0.00
4.04
0.00
1.90
0.00
9.04
0.00
0.24
0.24
0.00
0.00
0.00
42.82
1.43
14.04
177.96
0.00
0.00
1.78
9.99

30.27
0.00
0.80
4.13
0.00
0.27
0.00
5.73
0.00
0.40
106.27
0.53
0.00
1.60
0.27
3.47
0.l3
37.87
2.27
0.00
0.27
0.00
8.40
0.00
0.00
4.53
0.00
2.13
0.00
10.13
0.00
0.27
0.27
0.00
0.00
0.00
48.00
1.60
15.73
199.47
0.00
0.00
2.00
11.20

�-59-

Table 7.
Unit No.1

Type of
Plant

Pre-treatment production of vegetation in habitat improvement
(continued).

Scientific Name }j

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Forbs

Orthocaq~us luteus
Osmorhiza sp.
Pachistima myrsinites
Pedicularis grayi
Penstemon crandal1ii
Penstemon strictus
Phacelia heterophy11a
Polemonium foliosissimum
Polygonum aviculare
Polygonum douglasii
Potent ilIa pulcherrima
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambros ioides
Senecio integerrimus
Silene menziesii
Smilacina stellata
Solidago sparsif10ra
-Stellaria sp.
Swertia radiata
Taraxacum officina1e
Tha1ictrum fendleri
Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Va1eriana occidentalis
Veronica biloba
Viguiera multiflora
Viola rugulosa
Viola she1tonii
Wyethia amp1exicaulis
Total Forbs

0.02
0.00
0.00
0.00
0.49
0.15
0.03
0.00
0.00
0.10
0.34
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.51
1.02
0.00
0.28
0.06
0.01
0.01
0.00
0.04
0.00
0.17
D.Ol
0.51
0.22
31.04

0.36
0,00
0.00
0.00
.-8.09
2.50
0.48
0.00
0.00
1.67
5.59
0.00
0.00
0.00
0.00
0.00
0.00
0.00
8.33
16.77
0.00
4.64
0.95
0.12
0.12
0.00
0.71
0.00
2.85
0.24
8.33
3.57
509.61

0.40
O~OO
0.00
0.00
9.07
2.80
0.53
0.00
0.00
1.87
6.27
0.00
0.00
0.00
0.00
0.00
0.00
0.00
9.33
18.80
0.00
5.20
1.07
0.13
0.13
0.00
0.80
0.00
3.20
0.27
9.33
4.00
571.21

Grasses

Agropyron s p ,
Bromus anoma1us
Bromus inermis
Bromus tectorum
Carex sp.
Dactylus glomerata
E1ymus glaucus
Festuca thurberi
Hordeum brachyantherum
Juncus balticus

4.20
0.44
0.00
0.50
0.00
0.00
0.00
0.06
0.00
0.00

68.99
7.26
0.00
8.21
0.00
0.00
0.00
0.95
0.00
0.00

77.33
8.13
0.00
9.20
0.00
0.00
0.00
1.07
0.00
0.00

---------------------------------------------------------------.------------

�-60-

Table 7.
Unit No.1

Pre-treatment
(continued).

Type of
Plant

Scientific

production

Name

11

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Grasses

Koeleria cristata
Phleum prc1tense
Poa sp.
Sitanion hystrix
Stipa sp.
Total Grasses

0.33
0.00
14.60
0.34
0.72
21.19

5.35
0,.00
239.69
5.59
"11.78
347.82

6.00
0.00
268.67
6.27
l3.20
389.87

Shrubs

Acer glabrum
Amelanchier alnifolia
Artemisia cana
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus gambellii
Ribes cereum
Rosa nutkana
Symphoricarpos
sp.
Total shrubs

0.00
6.75
0.88
11.93
0.04
0.00
O.U
1.56
0.00
0.14
0.00
r2.62
0.00
0.39
13.35
47.77

0.00
UO.87
14.39
195.92
0.71
0.00
1.78
25.57
0.00
2.26
0.00
207.22
0.00
6.42
219.12
784.26

'0.00
124.27
16.13
219 60
0.80
0.00
2.00
28.67
0.00
2.53
0.00
232.27
0.00
7.20
245.60
879.07

100.00

1,641.69

1,840.15

Total of All Species

11 All plants

0

which occur in any of the eight habitat improvement units are
listed even though some species may not occur in this particular unit.

-;

�-61-

Table 8.
Unit No.2.

Type of
Plant

Forbs

Pre-treatment

Scientific

production

Name

II

Achillea
lanulosa
Agastache
urticifolia
~goseris glauca
Allium acuminatum
Androsace
septentrionalis
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia
dracunculus
Artemisia
ludoviciana
Aster-Erigeron
Ba1samorhiza
sagittata
Berberis repens
Calochortus
gunnisonii
Capsella bursa-pastoris
Castilleja
linariaefolia
Chenopodium
berlandieri
Cirsium canescens
Comandra umbel lata
Delphinium
occidentale
Descurainia
·richardsonii
Disporum trachycarpum
Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium sp.
Gentiana heterosepala
Geranium parryi
Gilia aggregata
Grindelia
aphanactis
Hackelia leptophylla
Helenium hoopesii
Hydrophyllum
fendleri
Iris missouriensis
Lactuca scariola
Lappula redowskii
Lathyrus leucanthus
Ligusticum
sp.
Linum le,visii
Lupinus argentues
Madia g10merata
Mertensia
franciscana
Musineon divarcatum
Opuntia sp.

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

2.27
0.02
0.01
0.00
0.00
0.00
0.01
0.03
0.93
0.08
4.11
0.00
0.00
0.l3
0.00
0.04
0.00
~.82
0.00
2.06
0.02
0.00
0.01
0.00
0.00
2.40
0.00
0.60
0.04

G.OO
0.01
0.01
0.00
0.00
0.00
0.00
7.09
1.15
0.20
8.11
0.07
0.01
0.00
0.00

37.23
0~36
0.12
0.00

oioo
0.00
0.12
0.48
15.23
1.31
67.33
0.00
0.00
2.14
0.00
0.71
0.00
29.74
0.00
33.78
0.36
0.00
0.12
0.00
0.00
39.26
0.00
9.75
0.59
0.00
0.24
0.12
0.00
0.00
0.00
0.00
116.10
18.79
3.21
l32.87

1.19
0.12
0.00
0.00

---------------------------------------------------------------------------

41. 73
0.40
0.13
0.00
0.00
0.00
0.13
0.53
17.07
1.47
75.47
0.00
0.00
2.40
0.00
0.80
0.00
33.33
0.00
37.87
0.40
0.00
0.13
0.00
0.00
44.00
0.00

10.93
0.67
0.00
0.27
0.13
0.00
0.00
0.00
0.00
130.l3

21.07
3.60

148.93.
1.33
0.13
0.00

0.00

�-62-

Table 8.
Unit No.2

Type of
Plant

Forbs

Grasses

Pre-treatment
(continued).

Scientific

production

Name

!f

Orthocarr~us luteus
Osmorhiza sp.
Pachistima myrsinites
Pedicularis grayi
Penstemon crandallii
Penstemon strictus
Phacelia heterophylla
Polemonium fo1iosissimum
Polygonum avicu1are
Polygonum douglasii
Potentilla pu1cherrima
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Silene menziesii
Smilacina stellata
Solidago sparsiflora
Stellaria sp.
Swertia radiata
Taraxacum officinale
Thalictrum fendleri
Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
Veronica biloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii
Wyethia amplexicaulis
Total Forbs

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

0.00
0.37
0.00
0.00
0.00
0.36
0.00
0.00
0.00
0.04
1.22
0.01
0.00
0.09
0.00
0.00
0.00
0.00
'0.00
1.05
0.32
0.73
2.33
0.11
0.00
0.00
0.46
0.00
0.01
0.28
0.45
1.02
40.14

0.00
6,.07
0.00
0.00
0.00
5.83
0.00
0.00
0.00
0.59
19.98
0.12
0.00
1.43
0.00
0.00
0.00
0.00
0.00
17.25
5.23
12.01
38.18
1.78
0.00
0.00
7.61
0.00
0.12
4.64
7.38
16.77
657.21

Agropyron sp.
0.54
8.92
Bromus anoma1us
0.48
7.85
Bromus incrmis
0.00
0.00
Bromus tectorum
0.00
0.00
Carex sp.
0..00
0.00
Dactylus glomerata
0.20
3.21
E1ymus glaucus
0.33
5.35
Festuca thurbcri
0.00
0.00
Hordeum brach~antherum
0.00
0.00
Juncus balticus
0.00
0.00
---------------------------------------------------------------------------

0.00
6'.80
0.00
0.00
0.00
6.53
0.00
0.00
0.00
0.67
22.40
0.13
0.00
1.60
0.00
0.00
0.00
0.00
0.00
19.33
5.87
13.47
42.80
2.00
0.00
0.00
8.53
0.00
0.13
5.20
8.27
18.80
736.65
10.00
8.80
0.00
0.00
0.00
3.60
6.00
0.00
0.00
0.00

�-63-

_Tal:iI~:8 Pre-treatment production of vegetation in habitat improvement
Unit No.2 -(continued).

Type of
Plant

Scientific Name l/

Air Dry Forage Weights
Percent Pounds
Kilograms
Composition
Per Acre
Per Hectare

Grasses

Koeleria cristata
Phleum pratense
Poa sp.
Sitanion hystrix
Stipa sp.
Total Grasses

0.00
0.04
19.05
0.00
0.46
21.10

0.00
0..
71
312.02
0.00
_..
-7~49
345.55

0.00
0.80
349.73
0.00
8.40
387.33

Shrubs

Acer glabrum
Amelanchier a1nifolia
Artemisia ~
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus gambe11ii
Ribes cereum
Rosa nutkana
Symphoricarpos sp.
Total shrubs

0.00
4.29
0.00
0.00
0.00
0.00
0.00
0.72
0.00
10.41
0.00
4.81
0.00
1.54
16.99
38.76

0.00
70.30
0.00
0.00
0.00
0.00
0.00
11.78
0.00
170.46
0.00
78.75
0.00
25.22
278.12
634.63

0.00
78.80
0.00
0.00
0.00
0.00
0.00
13.20
0.00
191.07
0.00
88.27
0.00
28.27
311.73
711.34

100.00

1,637.39

1,835.32

Total of All Species

!/ All plants which occur in any of the eight habitat improvement units are
listed even though some species may not occur in this particular unit.

�-64-

Table 9.
Unit No.3.

Type of
Plant

Forbs

Pre-treatment

Scientific

production

Name

II

Achillea
1anu1osa
Agastache
urticifo1ia
Agoseris glauca
Allium acuminatum
Androsace
scptentriona1is
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia
dracuncu1us
Artemisia
1udoviciana
Aster-Erigeron
Balsamorhiza
sagittata
Berberis repens
Ca1ochortus
gunnisonii
Capse11a bursa-pastoris
Casti11eia
1inariaefo1ia
Chenopodium
ber1andieri
Cirsium canescens
Comandra urub e Ll.at a
Delphinium
occidentale
Descurainia
richardsonii
Disporum trachycarpum
Eriogonum umbe11atum
Epilobium panicu1atum
Fragaria americana
Galium sp.
Gentiana heterosepa1a
Geraniu~ parryi
Gilia aggregata
Grindeliaaphanactis
Hackelia 1eptophylla
He1enium hoopesii
Hydro~ly11um
£endleri
Iris missouricnsis
Lactuca scariola
Lappula reduwskii
Lathyrus leucanthus
Ligusticllm sp.
Liuulll le\v i.s ii
~.nus
argcntues
Nadia c~l()ll1r;rat.:l
Mertensi.:l fcanciscana
Musincon divarcaluill
0PUI1tj:1
sp.

----

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

0.44
0.00
0.02
0.03
0.03
0.03
0.02
0.04
0.00
0.05
1.97
0.00
0.00
0.01
0.00
0.26
0.00
0.30
0.00
0.00
0.00
0.00
0.03

8.21
0.00
0.36
0.59
0.59
0.48
0.36
0.83
0.00
0.95

9.20
0.00
0.40
0.67
0.67
0.53
0.40
0.93
0.00
1.07

37.11
0.00
0.00
0.24
0.00
4.88
0.00
5.71
0.00
0.00
0.00
0.00
0.48

41.60
0.00
0.00
0.27
0.00
5.47
0.00
6.40
0.00
0.00
0.00
0.00
0.53

0.00
0.00
0.00
0.00
0.00
0.00
0.00
8.01

0.00
0.00
.18.68
0.00
6.07
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
151. 07

0.00
0.00
20.93
0.00
6.80
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
169.33

2.08
0.00
1. 58
0.00
0.00
0.00
0.00

39.14
0.00
29.74
0.00
0.00
0.00
0.00

43.37
0;00
33.33
0.00
0.00
0.00
0.00

0.00
0.00
0.99
0.00
0.32

b.oo

---------------------------------------------------------------------------

�-65-"

Table 9 • Pre-treatment
Unit No. 3 (continued).

Type of
Plant

Scientific

production

Name

il

of vegetation

in habitat

improvement

Air Dry Forage Weights
Kilograms
Pounds
Percent
Per Hectare
Composition
Per Acre

Forbs

Orthocaq~us luteus
Osmorhiza sp.
Pachistima myrsinites
Pedicularis grayi
Penstemon crandallii
Penstemon strictus
Phacelia heterophylla
Po1emonium foliosissimuin
Polygonum aviculare
Polygonum douglasii
Potentilla Eulcherrima
Ranunculus inamoenus
Rumex crisEus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Silene menziesii
Smilacina stellata
Solidago sEarsiflora
Stellaria sp.
Swertia radiata
Taraxacum officinale
Tha1ictrum fendleri
TragoEogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
Veronica biloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii
Wyethia amplexicaulis
Total Forbs

0.00
0.42
1.37
0.00
0.00
0.00
0.03
0.00
0.00,
0.00
0.09
0.00
0.00
0.00
0.00
0.01
0.02
0.00
'0.00
0.58
0.00
0.09
1.39
0.05
0.00
0.00
0.00
0.00
0.01
-0.01
0.28
1. 76
22.32

0.00
7..97
25.81
0.00
0.00
0.00
0.59
0.00
0.00
0.00
1.78
0.00
0.00
0.00
0.00
0.12
0.36
0.00
0.00
10.94
0.00
1.67
26.17
0.95
0.00
0.00
0.00
0.00
0.12
0.24
5.23
33.19
420.63

0.00
8'.93
28.93
0.00
0.00
0.00
0.67
0.00
0.00
0.00
1.20
0.00
0.00
0.00
0.00
0.13
0.40
0.00
0.00
12.27
0.00
1.87
29.33
1.07
0.00
0.00
0.00
0.00
0.13
0.27
5.87
37.20
470.67

Grasses

Agropyron sp.
Bromus anomalus
Bromus inermis
Bromus tectorum
Carex sp.
Dactylus glomerata
Elymus glaucus
Festuca thurberi
Hordeum brachyantherum
Juncus balticus

0.30
0.04
0.04
0.01
0.00
0.00
0.13
0.00
0.00
0.00

5.71
0.83
0.71
0.24
0.00
0.00

6.40
0.93
0.80
0.27
0.00
0.00
2.80
0.00
0.00
0.00

-

2.50
0.00
0.00
0.00

---------------------------------------------------------------------------

�-66-

Table
No.3

9. Pre-treatment
(continued).

Type of
Plant

Scientific

production

Name

1/

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Grasses

Koe1eria cristata
Ph1eum pratense
Poa sp.
Sitanion hystrix
SUpa
sp ,
Total Grasses

0.00
0.00
17.74
0.00
0.08
18.34

0.00
0.00
334.14
0.00
.- 1.55
345.68

0.00
0,.00
374.53
0.00
1. 73
387.46

Shrubs

Acer glabrum
Ame1anchier alnifo1ia
Artemisia cana
Artemisia tridentata
Cercocarpus montanus
Chrysotha~~us depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus gambe11ii
Ribes cereum
Rosa nutkana
Symphoricarpos
sp.
Total shrubs

0.00
9.91
0.00
0.51
0.00
0.00
0.00
0.00
0.00
9.74
0.00
28.67
0.00
1.69
8.82
59.34

0.00
187.00
0.00
9.64
0.00
0.00
0.00
0.00
0.00
183.55
0.00.
540.41
0.00
31.88
166.30
1,118.78

0.00
209.60
0.00
10.80
0.00
0.00
0.00
0.00
0.00
205.73
0.00
605.73
0.00
35.73
186.40
1,253.99

100.00

1,885.09

2,112.12

Total

of All Species

1/ All

plants which occur in any of the eight habitat improvement units are
even though some species may not occur in this particular unit.

listed

�-67-

Table 10. Pre-treatment
Unit No.4.

Type of
Plant

Forbs

Scientific

production

Name

II

Achillea lanulosa
Agastache urticifolia
Agoseris glauca
Allium acuminatum
Androsace septentrionalis
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia dracunculus
Artemisia ludoviciana
Aster-Erigeron
Balsamorhiza sagittata
Berberis repens
Calochortus gunnisonii
Capsella bursa-pastoris
Castilleja linariaefolia
Chenopodium berlandieri
Cirsium canescens
Comandra umbel lata
Delphinium occidentale
Descurainia richardsonii
Disporum trachycarpum
Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium sp.
Gentiana heterosepala
Geranium parryi
Gilia aggregata
Grindelia aphanactis
Hackelia leptophylla
Helenium hoopesii
Hydrophyllum fendleri
Iris missouriensis
Lactuca scarioia
Lappula redowskii
Lathyrus Ieucanthus
Ligusticum sp.
Linum IC,visii
Lupinus argentues
Had ia£lomerata
Mertcnsia franciscana
Musincon divarcatum
Opuntia sp.

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
,Composition
Per Acre
Per Hectare

1.44
0.21
0.00
0.02
0.01
0.02
0.02
0.15
0.00
0.05
3.01
0.00
0.02
0.04
0.00
0.03
0.00
D.65

0.00
0.42
0.00
0.00
0.00
0.00
0.00
0.95
0.00
0.41
0.00
'0.43
0.10
0.48
0.25
0.00
0.00
0.00
8.85
1.47
0'.05
6.33
0.46
0.98
0.01
0.00

23.32
3'.45
0.00
,,_ O.~36
0.12
0.36
0.36
2.50
0.00
0.83
48.77
0.00
0.36
0.59
0.00
0.48
0.00
10.47
0.00
6.78
0.00
0.00
0.00
0.00
0.00
15.46
0.00
6.66
0.00
6.90
1.67
7.73
4.04
0.00
0.00
0.00
143.22
23.79
0.83
102.42
7.49
15.94

0.24
0.00

26.13
3.87
0.00
0.40
0.13
0.40
0.40
2.80
0.00
0.93
54.67
0.00
0.40
0.67
0.00
0.53
0.00
11. 73
0.00
7.60
0.00
0.00
0.00
0.00
0.00
17.33
0.00
7.47
0.00
7.}3
1.87
8.67
4.53
0.00
0.00
0.00
160.53
26.67
0.93
114.80
8.40
17.87
0.27
0.00

---------------------------------------------------------------------------

�-68-

Table 10.
Unit No.4

Type or
Plant

Forbs

Grasses

Pre-treatment
(continued).

Scientific

production

Name

II

Orthocarr~us luteus
Osmorhiza sp.
Pachistima myrsinites
Pedicularis grayi
Penstemon crandallii
Penstemon strictus
Phacelia heteroEhylla
Polemonium foliosissimum
Polygonum aviculare
Polygonum douglasii
Potentilla Eulcherrima
Ranunculus inamoenus
Rumex crisEus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Silene menziesii
Smilacina stellata
Solidago sj2arsiflora
.Stellaria sp.
Swertia radiata
Taraxacum officinale
Thalictrum fendleri
TragoEogon dubium
TragoEogon Eratensis
Urtica dioica
Valeriana occidentalis
Veronica biloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii
Wyethia amElexicaulis
Total Forbs
AgroEyron sp.
Bromus anomalus
Bromus inermis
Bromus tectorum
Carex sp.
Dactylus glomerata
Elymus glallclls
Festuca tburberi
Hordeum brach:iantherllm
JUl1CllSbalticlls

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

0.00
1.25
0.15
0.00
0.30
0.01
0.00
0.01
0.00
0.00
0.34
0.00
0.00
0 ..25
0.00
0.00
0.00
0.01
0.01
1.03
0.00
0.27
2.73
0.00
0.00
0.15
0.23
0.00
0.07
·0.12
0.29
0.26
34.34

0.00
20.22
2.38
0.00
4.88
0.24
0.00
0.12
0.00
0.00
5.47
0.00
0.00
4.04
0.00
0.00
0.00
0.12
0.12
16.77
0.00
4.40
44.25
0.00
0.00
2.50
3.81
0.00
1.19
2.02
4.64
4.28
556.59

0.00
22.67
2.67
0.00
5.47
0.27
0.00
0.13
0.00
0.00
6.13
0.00
0.00
4.53
0.00
0.00
0.00
0.13
0.13
18.80
0.00
4.93
49.60
0.00
0.00
2.80
4.27
0.00
1.33
2.27
5.20
4.80
623.86

3.66
2.53
0.02
0.07
0.00
0.00
2.00
0.00
0.51
0.01

59.36
41.04
0.36
1.19
0.00
0.00
32.36
0.00
8.21
0.24

66.53
46.00
0.40
1.33
0.00
0.09
36.27
0.00
9.20
0.27

------------------------------------------------------------------------------

�-69-

Table 10.
Unit No.4

Type of
Plant

Pre-treatment
(continued).

Scientific

production

Name

l/

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Grasses

Koe1eria cristata
Ph1eum pratense
Poa sp.
Sitanion hystrix
Stipa sp.
Total Grasses

0.01
0.00
10.15
0.00
1.66
20.62

0.12
0.00
164.51
0.00
--26.88
334.27

0.l3
0.00
184.40
0.00
30.l3
374.66

Shrubs

Acer glabrum
Ame1anchier alnifolia
Artemisia ~
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidif10rus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus g2mbe11ii
Ribes cereum
Rosa nutkana
Symphoricarpos
sp.
Total shrubs

0.00
8.31
0.00
0.00
0.12
0.00
0.00··
0.10
0.00
4.06
0.22
17.33
0.79
1.11
13.00
45.04

0.00
134.78
0.00
0.00
1.90
0.00
0.00
1.55
0.00
65.78
3.57
280.97
12.85
17.96
210.79
730.15

0.00
151. 07
0.00
0.00
2.l3
0.00
0.00
1.73
0.00
73.73
4.00
314.93
14.40
20.13
236.27
818.39

1,621.01

1,816.91

Total

100.00

of All Species

l! All plants which
listed

occur in any of the eight habitat improvement units are
even though some species may not occur in this particular unit.

-;

�-70-

Table 11. Pre-treatment
Unit No.5.

production

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Type of
Plant

Forbs

of vegetation

Achillea lanulosa
Agastache urticifolia
Agoseris glauca
Allium acuminatum
Androsace septentrionalis
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia dracunculus
Artemisia
ludbviciana
Aster-Erigeron
Balsamorhiza
sagittata
Berberis repens
Calochortus
gunnisonii
Capsella bursa-pastoris
Castilleja
linariaefolia
Chenopodium berlandieri
Cirsium canescens
Comandra umbellata
Delphinium occidentale
Descurainia
richardsonii
Disporum trachycarpum
Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium sp.
Gentiana heterosepala
Geranium parryi
Gilia aggregata
Grindelia aphanactis
Hackelia leptophylla
Helenium hoopesii
Hydrophyl1um
fendleri
Iris missouriensis
Lactuca scario1a
Lappul! redowskii
Lathyrus leucanthus
Ligusticum sp.
Linum 1c\,/Lsii
Lupinus ar~~cntues
Madia glolllcrata
Mertensia
franciscana
Musineon divarcatllm
Opuntia sp.

0~84
0.00
D.-DO
0.06
0.26
0.01
0.00
0.00
0.00
0.00'
3.30
0.00
0.02
0.06
0.00
0.48
0.01
-0.01
0.00
0.09
0.00
0.02
0.03
0.00
0.02
0.49
0.03
0.16
0.00
0.13
0.00
0.00
0.Q8
0.00
.0.00
0.00
5.08
0.98
0.13
13.58
0.00
0.81
0.00
0.00

14.16
0.00
0.00
0.95
4.40
0.12
0.00
0.00
0.00
0.00
55.55
0.00
0.36
1.07

15.87
0.00
0.00
1.07
4.93
0.13
0.00
0.00
0.00
0.00
62.27
0.00
0.40
1.20

0.00
8.09
0.24
0.24
0.00
1. 55
0.00
0.36
0.48
0.00
0.36
8.21
0.48
2.62
0.00
2.26
0.00
0.00
1.48
0.00
0.00
0.00
85.53
16.53
2.14
228.99
0.00
13.68
0.00
0.00

0.00
9.07
0.27
0.27
0.00
1.73
0.00
0.40
0.53
0.000.40
9.20
0.53
2.93
0.00
2.53
0.00
0.00
1.60
0.00
0.00
0.00
95.87
18.53
2.40
256.67
0.00
15.33
0.00
0.00

------------------------------------------------------------------------------

�-71-

Table 11.
Unit No.5

Type of
Plant

Pre-treatment
(continued).

Scientific

production

Name

II

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Forbs

Orthocarpus luteus
Osmorhiza sp.
Pachistima myrsinites
Pedicularis grayi
Penstemon crandallii
Penstemon strictus
Phacelia heterophylla
Polemonium foliosissimum
Polygonum aviculare
Polygonum douglasii
Potentilla pulcherrima
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Silene menziesii
Smilacina stellata
Solidago sparsiflora
Stellaria sp.
.
Swertia radiata
Taraxacum officinale
Thalictr'um fend1eri
Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
Veronica biloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii
Wyethia amplexicaulis
Total Forbs

0.00
0.18
0.63
0.28
0.18
0.03
0.00
0.04
0.00
0.00
0.01
0.00
0.00
0.11
0.04
0.00
0.00
0.02
'0.20
0.47
0.04
0.08
1.37
0.01
0.00
.0.17
0.53
0.00
0.00
.0.38
0.04
0.51
32.00

0.00
3.09
10.59
4.64
2.97
0.48
0.00
0.59
0.00
0.00
0.24
0.00
0.00
1. 90
0.71
0.00
0.00
0.36
3.33
7.85
0.59
1.43
23.08
0.12
0.00
2.85
8.92
0.00
0.00
6.42
0.71
8.56
539.28

0.00
3.47
11.87
5.20
3.33
0.53
0.00
0.67
0.00
0.00
0.27
0.00
0.00
2.13
0.80
0.00
0.00
0.40
3.73
8.80
0.67
1.60
25.87
0.13
0.00
3.20
10.00
0.00
0.00
7.20
0.80
9.60
604.40

Grasses

Agropyron sp.
Bromus anomalus
Bromus inermis
Bromus tectorum
Carex sp.
Dactylus glomerata
Elymus glaucus
Festuca thurberi
Hordeum brachyantherum
Juncus baltLcus

1.17
0.45
0.00
0.00
0..00
0.00
0.49
0.00
0.00
0.00

19.75
7.61
0.00
0.00
0.00
0.00
8.33
0.00
0.00
0.00

22.13
8.53
0.00
0.00
0.00
0.00
9.33
0.00
0.00
0.00

----

�-72-

Table 11.
Unit No.5

Pre-treatment
(continued).

Type of
Plant

Scientific

production

Name

l/

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Grasses

Koeleria cristata
Phleurn pratense
Poa sp.
Sitanion hystrix
Stipa sp.
Total Grasses

0.10
0.00
9.24
0.01
0.18
100.00

1.67
0.00
155.71
0.12
...2.97
1,685.17

1.87
0.00
174.53
0.13
3.33
1,888.79

Shrubs

Acer glabrurn
Arnelanchier a1nifo1ia
Artemisia cana
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus
depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus gambellii
.Ribes cereum
Rosa nutkana
Sym:ehoricarpos sp.
Total shrubs

0.03
11. 97
0.00
5.12
0.00
0.08
0.00
0.23
0.00
1. 79
0.00
r9.84
1.33
1.26
14.71
56.36

0.48
201. 75
0.00
86.24
0.00
1.31
0.00
3.93
0.00
30.21
0.00
334.38
22.36
21.17
247.90
949.73

0.53
226.13
0.00
96.67
0.00
1.47
0.00
4.40
0.00
33.87
0.00
374.80
25.07
23.73
277.87
1,064.54

100.00

1,685.17

1,888.79

Total

of All Species

1/ All plants

which occur in any of the eight habitat improvement units are
listed even though some species may not occur in this particular unit.

�-73-

Table 12.

Pre-treatment

production

of vegetation

in habitat

improvement

Unit No.6.

Type of
Plant

Forbs

Scientific

1/

Name -

Achillea 1anu1osa
Agastache urticifo1ia
Agoseris glauca
Allium acuminatum
Androsace seEtentrionalis
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia dracuncu1us
Artemisia 1udoviciana
Aster-Erigeron
Balsamorhiza sagittata
Berberis reEens
Ca1ochortus gunnisonii
CaEse11a bursa-Eastoris
Castilleja 1inariaefo1ia
ChenoEodium ber1andieri
Cirsium canescens
Comandra umbe11ata
De1Ehinium occidentale
Descurainia richardsonii
DisEorum trachycarEum
Eriogonum umbe11atum
EEi10bium Eanicu1atum
Fragaria americana
Galium sp ,
Gentiana heteroseEa1a
Geranium Earryi
Gilia aggregata
Grindelia aEhanactis
Hacke1ia 1eEtoEhylla
He1enium hooEesii
HydroEhy11um
fend1eri
Iris missouriensis
Lactuca scario1a
LaEpula redowskii
Lathyrus 1eucanthus
Ligusticum sp.
Linum lewis ii
Lupinlls argentues
Nadia glomerata
Mertensia franciscana
Musineon divarc.:ttum
0Euntia sp.

Air Dry Forage Weights
Kilograms
Pounds
Percent
Per Hectare
Per Acre
Composition

1.37
0.31
0.-00
0.01
0.01
0.00
0.00
0.03
0.00
0.00
4.31
0.00
0.00
0.10
0·.00
0.12
0.08
·0.22
0.20
0.39
0.00
0.00
0.05
0.00
0.00
1.20
0.01
0.33
0.00
'0.00
0.00
0.12

O.QO
0.00
·0.00
0.00
5.80
1. 68
0.35
8.84
0.00
0.00
0.01
0.00

24.98 .
5.59
0.00
0.24
0.24
0.00
0.00
0.48
0.00
0.00
78.51
0.00
0.00
1. 78
0.00
2.14
1.43
4.04
3.69
7.02
0.00
0.00
0.83
0.00
0.00
21. 77
0.24
5.95
0.00
0.00
0.00
2.14
0.00
0.00
0.00
0.00
105.63
30.45
6.42
160.60
0.00
0.00
0.12
0.00

28.00
6.27
0.00
0.27
0.27
0.00
0.00
0.53
0.00
0.00
88.00
0.00
0.00
2.00
0.00
2.40
1. 60
4.53
4.13
7.87
0.00
0.00
0.93
0.00
0.00
24.40
0.27
6.67
0.00
0.00
0.00
2.40
0.00
0.00
0.00
0.00
118.40
34.13
7.20
179.9-9
0.00
0.00
0.13
0.00

-----------------------------------------------------------------------------

�-74-

Table 12.
Unit No.6

Type of
Plant

Forbs

Grasses

Pre-treatment
(continued).

Scientific

production

Name

l/

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Orthocarpus luteus
Osmorhiza sp.
Pachistima myrsinites
Pedicularis grayi
Penstemon crandallii .
Penstemon strictus
Phacelia heterophylla
Polemonium foliosissimum
Polygonum aviculare
Po1ygonum douglasii
Potentilla pulcherrima
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Si1ene menziesii
Smilacina stellata
Solidago sparsiflora
Stel1aria sp.
SWertia radiata
Taraxacum officinale
Thalictrum fendleri
Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
Veronica b iloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii
Wyethia amplexicaulis
Total Forbs

0.00
0.65
0.41
0.00
0.07
0.05
0.00
0.00
0.00
0.00
0.08
0.00
0.00
0.00
0.00
0.00
0.00
0.10
·0.20
0.56
0.00
0.07
1.17
0.00
0.00
0.00
0.05
0.00
.0.36
0.00
0.07
0.01
29.39

3.69
10.11
0.00
1.31
21.17
0.00
0.00
0.00
0.83
0.00
6.54
0.00
1.19 .
0.24
534.11

0.00
13.33
8.27
0.00
1.47
0.93
0.00
0.00
0.00
0.00
1.60
0.00
0.00
0.00
0.00
0.00
0.00
2.l3
4.13
11.33
0.00
1.47
23.73
0.00
0.00
0.00
0.93
0.00
7.33
0.00
1.33
0.27
598.64

Agropyron sp.
Bromus anomalus
Bromlls inermis
Bromus tectorum
Carex sp.
Dactylus glomerata
Elymus glallcus
Festllca thurberi
Hordeum brachyan thcrum
JllllCllS
balticus

6.03
1.77
0.00
0.00
0.00
0.00
0.00
0.01
0.00
0.00

109.68
32.12
0.00
0.00
0.00
0.00
0.00
0.12
0.00
0.00

122.93
36.00
0.00
0.00
0..00
0.00
0.00
0.13
0.00
0.00

0.00
11. 90
7.37
0.00
1.31
0.83
0.00
0.00
0.00
0.00
1.43
0.00
0.00
0.00
0.00
0.00
0.00
1.90

-----------------------------------------------------------------------------

�-75-

Table 12.
Unit No.6

Type of
Plant

Pre-treatment production of vegetation in habitat improvement
(continued).

Scientific Name l/

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Grasses

Koeleria cristata
Ph1eum pratense
Poa sp.
Sitanion hystrix
Stipa sp.
Total Grasses

0.02
0.00
15.33
0.00
0.29
100.00

0.36
0.00
278.71
0.00
...5.23
1,817.28

0.40
0.00
312.40
0.00
5.87
2,036.89

Shrubs

Acer glabrum
Amelanchier alnifolia
Artemisia ~
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus gambellii
Ribes cereum
Rosa nutkana
Symphoricarpos sp.
Total shrubs

0.00
6.23
0.12
0.79
0.00
0.00
0.03
0.14
0.00
7.82
0.00
i8.50
0.06
1.59
11.88
47.16

0.00
113.13
2.14
14.39
0.00
0.00
0.59
2.62
0.00
142.03
0.00
336.17
1.07
28.91
215.90
856.95

0.00
126.80
2.40
16.13
0.00
0.00
0.67
2.93
0.00
159.20
0.00
376.80
1.20
32.40
241.99
960.52

100.00

1,817.28

2,036.89

Total of All Species

l/ All plants which occur in any of the eight habitat improvement units are
listed even though some species may not occur in this particular unit.

�-76.

Table 13. Pre-treatment
Unit No.7.

Type of
Plant

Forbs

Scientific

production

Name

~I

Achillea lanulosa
Agastache urticifolia
Agoseris glauca
Allium acuminatum
Androsace septentrionalis
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia dracunculus
Artemisia ludoviciana
Aster-Erigeron
Balsamorhiza sagittata
Berberis repen$
Calochortus gunnisonii
Capsella bursa-pastoris
Castilleja linariaefolia
Chenopodium berlandieri
Cirsium canescens
Comandra umbellata
Delphinium occidentale
Descurainia richardsonii
Disporum trachycarpum
Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium sp.
Gentiana heterosepala
Geranium parryi
Gilia aggregata
Grindelia aphanactis
Hackelia leptophylla
Helenium hoopesii
Hydrophy1lum
fend1eri
Iris missouriensis
Lactuca scariola
Lappu1a redowskii
Lathyrus leucanthus
Ligusticum sp.
Linum lcw i s ii
--Lupinus argentues
Mad ia .8lomcr3.ta
Mertensia franciscana
Musineon divarcatum
Opuntia sp.

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

3.50
0.01
0.00
0.12
0.00
0.04
0.04
0.05
0.21
0.39
7.22
0.00
0.00
0.04
0.06
0.48
0.02
0.05
0.02
0.00
0.00
0.00
0.28
0.02
0.01
1.12
0.00
0.07
0.00
0.44
0.00
0.11
0.00
2.25
0.00
0.00
1.80
0.15
0.23
5.42
0.00
0.00
0.00
0.08

58.76
0;12
0.00
2.02
0.00
0.71
0.59
0.83
3.57
6.54
121.10
0.00
0.00
0.71
0.95
8.09
0.36
0.83
0.36
0.00
0.00
0.00
4.64
0.36
0.24
18.79
0.00
1.19
0.00
7.38
0.00
1. 78
0.00
37.71
0.00
0.00
30.21
2.50
3.81
91.12
0.00
0.00
0.00
1.31

65.87
0.13
0.00
2.27
0.00
0.80
0.67
0.93
4.00
7.33
135.73
0.00
0.00
0.80
1. 07
9.07
0.40
0.93
0.40
0.00
0.00
0.00
5.20
0.40
0.27
21. 07
0.00
1.33
0.00
8.27
0.00
2.00
0.00
42.27
0.00
0.00
33.87
2.80
4;27
102.13
0.00
0.00
0.00
1.47

-----------------------------------------------------------------------------

�-77-

Table 13. Pre-treatment
Unit No. 7 (continued).

Type of
Plant

Forbs

Grasses

Scientific

production

Name

1/

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Orthocaq~us luteus
Osmorhiza sp.
Pachistima myrsinites
Pedicularis grayi
Penstemon crandal1ii
Penstemon strictus
Phacelia heterophylla
Po1emonium foliosissimum
Polygonum aviculare
Polygonum douglasii
Potent illa pulcherrima
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambros ioides
Senecio integerrimus
Silene menziesii
Smilacina stellata
Solidago sparsiflora
Stellaria sp.
Swertia radiata
Taraxacum officina1e
Tha1ictrum fend1eri
Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
Veronica biloba
Viguiera multiflora
Viola rugu10sa
Viola sheltonii
W~ethia amplexicaulis
Total Forbs

0.01
0.21
0.00
0.00
0.00
0.19
0.01
0.00
0.21
0.15
0.45
0.00
0.01
0.35
0.00
0.00
0.00
0.00
0.00
0.55
0.00
1.70
0.34
0.03
0.00
0.00
0.00
0.04
'0.50
·0.07
0.04
0.06
29.15

0.24
3.57
0.00
0.00
...
0.00
3.21
0.12
0.00
3.57
2.50
7.49
0.00
0.24
5.83
0.00
0.00
0.00
0.00
0.00
9.16
0.00
28.55
5.71
0.48
0.00
0.00
0.00
0.59
8.33
1.19
0.59
1.07
489.02

0.27
4'.00
0.00
0.00
0.00
3.60
0.13
0.00
4.00
2.80
8.40
0.00
0.27
6.53
0.00
0.00
0.00
0.00
0.00
10.27
0.00
32.00
6.40
0.53
0.00
0.00
0.00
0.67
9.33
1.33
0.67
1~20
548.15

Agrop~ron sp.
Bromus anoma1us
Bromus inermis
Bromus teetorum
Carex sp.
Dactylus glomerata
E1~mus glaucus
Festuca thurberi
Hordeum braehyantherum
Juncus ba1ticus

4.23
0.76
0.00
0.03
0.19
0.00
0.00
0.94
0.36
0.23

71.02
12.73
0.00
0.48
3.21
0.00
0.00
15.70
6.07
3.81

79.60
14.27
0.00
0.53
3'.60
0.00
0.00
17.60
6.80
4.27

-

-----------------------------------------------------------------------------

�-78-

Table 13.
Unit No.7

Type of
Plant

Pre-treatment
(continued).

Scientific

production

Name

11

of vegetation

Koeleria cristata
Phleum pratense
Poa sp.
S Hanion
hystrix
Stipa sp.
Total Grasses

0.23
0.04
16.70
0.04
1.07
100.00

Shrubs

Acer glabr-um
Ame1anchier alnifo1ia
Artemisia ~
Artemisiatridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus gambellii
Ribes cereum
Rosa nutkana
Symphoricarpos
sp.
Total shrubs

0.00
0.16
0.28
3.86
0.00
0.00
0.01
0.28
0.01 .
1.28
0.00
'8.51
4.81
3.10
23.73
46.03

of All Species

!I All plants which
listed

even though

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Grasses

Total

in habitat

100.00

3.93
0.71
280.26
0.71
17.96
1,678.11
0.00
2.62
4.64
64.83
0.00
0.00
0~24
4.76
0.24
21.53
0.00
142.75
80.77
51. 98 .
398.14
772.50
1,678.11

4.40
(}.80
314.13
0.80
20.13
1,880.95
0.00
2.93
5.20
72.67
0.00
0.00
0.27
5.33
0.27
24.13
0.00
160.00
90.53
58.27
446.27
865.87
1,880.95

occur in any of the eight habitat improvement units are
some species may not occur in this particular unit.

'.

�-79-

Table 14. Pre-treatment
Unit No.8.

Type of
Plant

Forbs

Scientific

production

Name

II

Achillea lanulosa
Agastache urticifolia
Agoseris glauca
Allium acuminatum
Androsace septentrionalis
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia drac~nculus
Artemisia ludoviciana
Aster-Erigeron
Balsamorhiza sagittata
Berberis repens
Calochortus gunnisonii
Capsella bursa-pastoris
Castilleja linariaefolia
Chenopodium berlandieri
Cirsium canescens
Comandra umbellata
Delphinium occidentale
Descurainia richardsonii
Disporum trachycarpum
Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium sp.
Gentiana heterosepala
Geranium parryi
Gilia aggregata
Grindelia aphanactis
Hackelia leptophylla
Helenium hoopesii
Hydrophyllum fendleri
Iris missouriensis
Lactuca scariola
Lappula redowskii
Lathyrus leucanthus
Ligusticum sp.
Linum le\visii
Lupinus argentues
Madia glomerata
Mertensia franciscana
Nusineon divarcatum
Opuntia sp.

of vegetation

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
.Composition
Per Acre
Per Hectare

2.56
0.28
0.00
0.04
0.00
0.01
0.01
0.08
0.00
0.19
7.61
0.00
0.00
0.04
0.00
0.16
0.02
.0.10
0.29
0.12
0.00
0.00
0.48
0.00
0.00
1.54
0.00
0.36
0.00
.0.03
0.15
0.33
0.11
0.81
0.01·
0.00
2.40
0.34
0.10
8.91
0.00
0.07
0.00
0.00

45.44·
5.00
0.00
0.71
0.00
0.24
0.12
1.43
0.00
3.33
134.78
0.00
0.00
0.71
0.00
2.85
0.36
1.78
5.23
2.14
0.00
0.00
8.56
0.00
0.00
27.24
0.00
6.42
0.00
0.59
2.74
5.83
2.02
14.51
0.12
0.00
42.47
6.07
1.90
157.85
0.00
1.19
0.00
0.00

50.93
~.•.
60
0.00
0.80
0.00
0.27
0.13
1. 60
0.00
3.73
151.07
0.00
0.00
0.80
0.00
3.20
0.40
2.00
5.87
2.40
0.00
0.00
9.60
0.00
0.00
30.53
0.00
7.20
0.00
0.67
3.07
6.53
2.27
16.27
0.13
0.00
47.60
6.80
2.13
176.93
0.00
1.33
0.00
0.00

-----------------------------------------------------------------------------

�-80-

Table 14. Pre-treatment
Unit No. 8 (continued).

.Type of
Plant

Forbs

Grasses

Scientific

production

Name

II

Orthocaq~us luteus
Osmorhiza sp.
Pachistima myrsinites
Ped icular is grayi
Penstemon cranda11ii
Penstemon strictus
Phacelia heteroJ2hylla
Polemonium foliosissimum
Polygonum aviculare
Polygonumdouglasii
Potentilla Eulcherrima
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Si1ene menziesii
Smi1acina stellata
Solidago sJ2arsiflora
-Stellaria sp.
Swertia radiata
Taraxacum officinale
Thalictrum fendleri
TragoJ2ogon dubium
TragoJ2ogon J2ratensis
Urtica dioica
Valeriana occidenta1is
Veronica biloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii
Wyethia amJ21exicaulis
Total Forbs
AgroJ2yron sp.
Bromus anomalus
Bromus inermis
Bromus tectorum
Carex sp.
Dactylus glomerata
Elymus glaucus
Festuca thurberi
Hordeum brachyantherum
JunCllS balticus

of vegetation

in habitat

improvement

Air.Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

0.01
0.24
0.00
0.00
0.04
0.02
0.00
0.27

-0.02
0.16
0.06
30.81

0.24
4.28
0.00
0.00
0.71
0.36
0.00
4.88
2.62
0.24
9.16
0.00
0.00
3.09
0.00
0.00
0.00
0.00
0.00
4.64
0.00
6.66
10.11
2.62
0.00
8.45
0.00
0.00
2.74
0.36
2.85
1.07
546.71

2.93
0.27
10.27
0.00
0.00
3.47
0.00
0.00
0.00
0.00
0.00
5.20
0.00
7.47
11.33
2.93
0.00
9.47
0.00
0.00
3.07
0.40
3.20
1.20
612.81

2.15
0.40
0.00
0.06
0.00
0.00
0.10
0.00
0.30
0.00

38.07
7.14
0.00
1.07
0.00
0.00
1.78
0.00
5.35
0.00

42.67
8.00
0.00
1.20
0.00
0.00
2.00
0.00
6.00
0.00

0.15
0.01

0.52
0.00
0.00
0.17
0.00
0.00
0.00
0.00
0.00
0.26
0.00
0.38

0.57
0.15
0.00
0.48
0.00
0.00

0.15

0.27
4.80
0.00
0.00
0.80
0.40
0.00

5.47

-----------------------------------------------------------------------------

�-81-

Table 14.
Unit No.8

Type of
Plant

Pre-treatment production of vegetation in habitat improvement
(continued).

Scientific Name !/

Grasses

Koe1eria cristata
Phleum pratense
Poa sp.
Sitanion hystrix
Stipa sp.
Total Grasses

Shrubs

Acer glabrum
Ame1anchier a1nifo1ia
Artemisia cana
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidif10rus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus gambe11ii
,Ribes cereum
Rosa nutkana
Symphoricarpos sp.
Total shrubs

Total of All Species

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

0.00
0.00
29.~n
0.03
0.66
100.00

0.00
0.00 .
528.99
0.59
..'11.66
1,774.44

0.00
0.00
592.93
0.67
13.07
1,988.95

0.00

2.04
0.84
3.39
0.00
0.00
0.00
0.11
0.60
7.11
0.00
'6.95
0.00
3.11
11.53
35.68

0.00
36.16
14.99
60.19
0.00
0.00
0.00
1.90
10.59
126.21
0.00
123.24
0.00
55.20
204.60
633.08

0.00
40.53
16.80
67.47
0.00
0.00
0.00
2.l3
11.87
141.47
0.00
138.l3
0.00
61.87
229.33
709.60

100.00

1,774.44

1,988.95

l/ All plants which occur in any of the eight habitat improvement units are
listed even though some species may not occur in this particular unit.

�-84-

Table 15.
Units No.1

Type of
Plant

Pre-treatment production of vegetation
through 8 combined (Continued).

Scientific

Name

li

in habitat

improvement

Air Dry Forage Weights
Percent
Pounds
Kilograms
Composition
Per Acre
Per Hectare

Grasses

Koe1eriacristata
Ph1eum pratense
Poa sp.
Sitanion hystrix
Stipa sp.
Total Grasses

0.08
0.01
16.70
0.05
0.62
21.88

1.43
0.18
286.75
0.88
10.69
375.86

1.60
0.20
321.42
0.98
11. 98
421. 30

Shrubs

Acer glabrum
Ame1anchier a1nifo1ia
Artemisia cana
Artemisia tridentata
Cercocarpus montanus
Chrysothanmus deprcssus
Chrysothanmus nauseosus
Chrysothamnus viscidif10rus
Crataegus erythropoda
Prunus virginiana
Purshia tridentata
Quercus gambellii
.Ribes cereum
Rosa nutkana
Symphoricarr~os sp.
Total shrubs

T
6.23
0.26
3.14
0.02
0.01
0.02
0.38
0.08
5.40
0.03
14.88
0.85
1.74
14.13
47.17

0.06
107.07
4.52
53.90
0.33
0.16
0.33
6.51
1.35
92.75
0.45
255.48
14.63
29.84
242.60
809.98

0.07
120.02
5.07
60.42
0.37
0.18
0.37
7.30
1. 52
103.97
0.50
286.37
16.40
33.45
271. 93
907.94

100.00

1,717.46

1,925.14

Total

of All Species

1:./ All plants which
listed

occur in any of the eight habitat improvement units arc'
even though some species may not occur in this particular unit.

~/ T (Trace) is less than 0.01% composition.

�-81-

Table 16. Summary of 1970 pre-treatment vegetation measurement data for
habitat improvement Unit Numbers 1 through 8.
Green Wt.
Pounds
Kilograms
Per Ac. Per Hectare

Dry Wt.
Po.unds
Kilograms
Per Ac.
Per Hectare

31.04
47.77
21.19

1572.58
1664.30
856.59

1762.67
1865.47
960.13

509.61
784.26
347.82

571.21
879.07
389.87

Total

100.00

4093.47

4588.27

1641.69

1840.15

Forbs
Shrubs
Grasses

40.14
38.76
21.10

2128.58
1462.55
889.19

657.21
634.63
345.55

736.65
711.34
387.33

Total

100.00

4480.32

2385.87
1639.33
996.67
5021. 87

1637.39

1835.32

Forbs
Shrubs
Grasses
Total

22.32
59.34
18.34
100.00

1412.23
2504.95
921.90

1582.93
2807.73
1033.33
5423.99

420.60
1118. 78
345.68
1885.09

470.67
1253.99
387.46
2112.12

Forbs
Shrubs
Grasses
Total

34.34
45.04
20.62
100.00

1841.90
1595.90
884.79
4322.59

2064.53
1788.80
991. 73
4845.06

556.59
730.15
334.27
1621.01

623.86
818.39
374.66
1816.91

Forbs
Shrubs
Grasses

32.00
56.36
11.64

1367.03
1906.49
420.15

1532.27
2136.93
470.93

539.28
949.73
196.16

604.40
1064.54
219.85

Total

100.00

3693.67

4140.13

1685.17

1888.79

6

Forbs
Shrubs
Grasses
Total

29.39
.47.16
23.45
100.00

1425.08
1807.16
925.23
4157.47

1597.33
2025.60
1037.07
4660.00

534.11
856.95
426.22
1817.28

598.64
960.52
477.73
2036.89

7

Forbs
Shrubs
Grasses
Total

29.15
46.03
24.82
100.00

1292.45
1604.70
873.01
3770.16

1448.67
1798.67
978.53
4225.87

489.02
772.50
416.59
1678.11

548.15
865.87
466.93
1880.95

8

Forbs
Shrubs
Grasses
Total

30.81
35.68
33.51
100.00

1417.59
1367.63
1247.60
4032.82

1588.93
1532.93
1398.40
4520.26

546.71
633.08
594.65
1774.44

612.81
709.60
666.54
1988.95

Total
of All
Units

Forbs
Shrubs
Grasses
Total

30.95
47.17
21.88

1557.14
1739.17
877.29
4173.60

1745.39
1949.43
983.35
4678.17

531. 62
809.98
375.86
1717.46

595.90
907.94
421.30

Habitat
Unit
Number

1

2

3

4

5

Type of
Plant

% Composition
By Dry Wt.

Forbs
Shrubs
Grasses

100.00

4839.08

1925.14

�-86-

Table 17. Pre-treatment cattle use on Hightower Mountain Oak Study Area
(Summer, 1970).

Habitat Improvement
Unit.No.

Cow Days
Use/Ac.

1

4.35

2

8.70

3

0.00

4

0.00

5

3.04

6

1.30

7

9.57
18.26

8

Table 18. Species and seeding rates that have been selected for reseeding
Hightower Mountain Oak Control Study Area.

Seed SEecifications
Purity Fill Germination

Pounds
Per Ac.

AgroEyron cristatum (Fairway strain)

95%

90%

2

Bromus inermis (Lincoln strain)

95%

90%

4

AgroEyron
intermedium (Arnurstrain)
,

95%

90%

3

Dactylis glome rata (Chinook strain)

95%

90%

1

Festuca ovina duriscula

90%

85%

1
11

Meticago sativa (Nomad variety)

99%

90%

Meticago sativa (Rambler variety)

99%

90%

Meticago sativa (Ladak variety)

99%

90%

Melilotus officinalis
Astragalus cf.ce r

99%
90%

99%

Purshia tridentata

95%

95%

2

CercocarEus montanus

95%

95%

1

Species

Grasses

Forbs

Shrubs

)
)
)
)

2

1
1
4

3

total

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                  <text>July, 1971

- 87 -

JOB FINAL REPORT

State of

COLORADO
--~--~~~~~-----------

Project No.

W-38-R-25

Work Plan No.

2

Job Title:

White River Elk Study - Publication of Results

Deer-Elk Investigations
Job No.

7 A

Period Covered: April 1, 1970 through March 31, 1971
Personnel:

Raymond J. Boyd

ABSTRACT
All editing, proof-reading and lay-out was completed. The manuscript was
given to the printer and publication completed. Copies of the publication
("Elk of the White River Plateau, Colorado," Technical Publication No. 25)
were sent to the Federal Aid office in Albuquerque and others sent to names
on the Division mailing list.

��- 89 -

WHITE RIVER ELK STUDY - PUBLICATION OF RESULTS

Raymond J. Boyd

P.S. OBJECTIVE
To formulate a management plan for the elk and their habitat on the White
River area.

SEGMENT OBJECTIVE
To publish all the research findings resulting from the White River elk
study through 1965.

METHODS AND MATERIALS
Publish as a Colorado Division of Game, Fish and Parks Technical Publication.
RESULTS AND DISCUSSION
The completed manuscript was taken to the printer and delivery of the
completed publication was accepted on February 1, 1971. The required
number of copies were sent to the Federal Aid office in Albuquerque and
persons on the Division's mailing list were also furnished with copies of
the completed publication.

Raym
Wildlife Researcher

��July, 1971

- 91 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-25

Work Plan No.

2

Job Title

Deer-Elk Investigations
Job No.

8

Hunter Harvest Surveys

Period Covered:

April 1, 1970 through March 31, 1971

Personnel:

Raymond J. Boyd, Pat Fuqua, Shirley Dunn, Kathy Edwards,
Karen Jackson, Velma Merkle and Charlotte Gardner.

ABSTRACT
Five different elk license types were surveyed by the 1970 random, with
a total of 26,104 questionnaires sent out. The total return was 18,779
usable cards for an average return of 71.9 percent. Kill estimates were
also determined for Sportsman's License holders in High Country Quality
Deer Seasons, Regular Deer Seasons, Primitive Weapons Seasons, 2nd Deer
License Purchasers, Archery Elk Hunters and all elk license hunters. All
kill statistics are reported in the 1970 Colorado Big Game Harvest Resume.

��- 93 -

HUNTER HARVEST

SURVEYS

Raymond J. Boyd

P. S. AND SEGMENT OBJECTIVE
To estimate the annual State kill of all big game by species, sex, age,
game management unit, County, residence of hunters and type of license,
season, period of season and to obtain information on man-days of recreation involved.

METHODS

AND MATERIALS

All big game hunters except deer, will be randomly surveyed by mail
questionnaire and one follow-up to obtain management information.
Returned cards will be coded by hand for IBM keypunch procedures which
will be used in computerized preparation of summaries of the data
supplied by returned questionnaire cards. The continuous sheet reports
will be manually processed by the clerical staff to obtain reports needed
by Game Planning.
Complete instructions for coding, keypunching,
are detailed in Boyd (1970).

etc. for the IBM system

RESULTS AND DISCUSSION
The Colorado Division of Game, Fish and Parks had within its big game
license framework, during 1970, five different kinds of elk licenses
available to the hunting public.
These five licenses are further broken
into two additional categories -- residents and nonresidents.
In effect,
then, 10 .different elk license types were surveyed for 1970 big game
season kill data.
Table 1 indicates the license types surveyed,
sent out, the number of usable questionnaires
return on each type of questionnaire.

Kill Statistics,

the number of questionnaires
returned and the percent

1970 Big Game Season

The total harvest summary for all big game species and special seasons
involved in the 1970 random survey are reported in the 1970 Colorado Big
Game Harvest Resume, a copy of which is attached as an appendix to this
report.
Table 2 details only the total kill estimates for species and
special seasons surveyed in 1970 by the random survey.

�July, 1971

- 97 -

JOB PROGRESS REPORT

State of

C~O~L~O~RAD~~O~

Project No.

W-38-R-25

Work plan No.

Job Title

6

_
Deer-Elk
Job No.

Investigations
11

Evaluation of the Effects of SpringSummer Grazing by Deer on Alfalfa

Period Covered:

April 1, 1970 through March 31, 1971

Personnel:

David F. Gordon, James A. Ives, Steven F. Steinert,
Julius J. Klein, Donald E. Speers, David A. Gordon,
Robert B. Gordon, Rifle Gap Conservation Unit inmates and Richard M. Bartmann.

ABSTRACT
All-night deer counts were made on the Rifle Gap study field from the
first week in April through the last week in September.
The spring peak
in mean deer numbers in 1970 (82.1) was nearly twice that in 1969 (42.2).
Future deer counts on the study field in 1971 will be made only at 9:00,
10:00, and 11:00 p.m. , MST, instead of all night as in previous years.
This decision was based on correlation analyses of the pooled 1968, 1969,
and 1970 deer count data. Lower hay yields were measured on grazed plots
than on ungrazed plots during both harvests.
This amounted to an estimated
1,216 pounds and 581 pounds difference per acre for the first and second
cuttings, respectively.
Differences were significant (P&lt;.Ol) only for
the first cutting.
Production differences cannot be related to mean deer
densities until the exact acreage upon which deer are counted is determined.
Results of proximate analyses of hay samples showed no significant difference
(P&gt;.05) in those factors tested between grazed and ungrazed plots for both
harvests.
At Little Hills, an II-acre portion of the study field was fenced for the
controlled phase of the study and 30 captive deer allowed free access into
it. Evidence of deer presence was later found over the entire field. A
heavy ·infestation of mustard plagued the first hay growth.
This delayed
the first cutting and precluded the second harvest and plot pairing.

��- 99 EVALUATION OF THE EFFECTS
OF SPRING-SUMMER GRAZING BY DEER ON ALFALFA
David F. Gordon and Richard M. Bartmann

P. S. OBJECTIVE
To determine
alfalfa.

the effect of spring-summer

SEGMENT
1.
2.
3.
4.

grazing by deer on production

of

OBJECTIVES

To determine the number and time of night deer use the study field.
To determine if there are differences in production of alfalfa between
grazed and ungrazed plots on the study field at Rifle Gap.
To test for factor(s) that determine a deer's preference for mature
over immature alfalfa.
To fence the control field, establish paired plots, and trap deer for
the controlled phase of the study at Little Hills.

METHODS

AND MATERIALS

- RIFLE GAP

Deer Counts
Deer were counted using the method outlined in the Segment 23 report
(Gordon 1969). This year, weekly counts were made from the first of April
through the end of September, except for the third week of August which was
the opening of archery season.
At each count, the general location of
groups of one or more deer were marked on a blank map of the study field.
A composite of these maps was made to illustrate deer distribution.

Irrigation
the study field was sprinkler irrigated twice during
only. The irrigation schedule was as follows:
Began
Terminated
Began
Interrupted
Resumed
Terminated

the first hay growth

April 16
May 3
May 11
May 18
June 1
June 8

Irrigation was begun while night-time temperatures still dropped below
freezing and a minor problem occurred with ice forming on the plot fences
(Fig. 1). This was easily removed by beating the fences with a shovel.
However, one morning the wind blew before the ice was removed and flattened
one fence on each of two plots and caused a third to sag. Repairs were
completed the same day.

�- 100 -

Fig. 1. Ice formed on plot fences due to sprinkler
irrigation during below freezing temperatures on the
Rifle Gap alfalfa damage study field, 1970.

�- 101 Problems were also encountered with the irrigation pump engine.
After the
irrigation season it was checked by a mechanic who indicated it was underpowered for the job. Consequently, a new pump and engine were purchased
near the end of the segment.,

Hay Harvest
The hay was cut, baled and stacked by the contractor, W. F. Scarrow of
Rifle, Colorado, using the same equipment as last year. Baling of the
second cutting was delayed nine days by machinery breakdown and wet weather.
The harvest schedule was as follows:
Cut
First Cutting
Second cutting

Baled

July 13 &amp; 14
September 4 &amp; 7

July 17 &amp; 18
September 20

A new method of marking the last bale from each plot was used. A sheet of
newspaper was tucked under the twine of the last bale from each plot as it
was ejected from the baler.
A crew following the baler weighed the hay to
the nearest pound with a Chatillion spring scale of 100-pound capacity.
After the baled hay was removed from the field, the edges of each fenced
plot were clipped using a Montgomery Ward 14 horsepow~r hydrostatic tractor
with a cutterbar attachment.
At first cutting the clippings were removed
from the field. At second cutting they were sparse enough to leave on the
field.

Proximate Analysis
The last bale from each plot was labeled with plot number, treatment and
cutting.
These bales were stored in the Rifle Gap barn until both cuttings
were complete and then taken to the Research Center in Fort Collins for
proximate analysis.

Preference

Factor(s)

Segment Objective 3, "to test for factor(s) that determine a deer's preffor mature over immature alfalfa" was deemed unattainable in this study.
There is little information to indicate what nutritional qualities deer
might seek in forage, and we could only surmise from the results.
Therefore, this objective will be deleted in the future.

RESULTS AND DISCUSSION

- RIFLE GAP

Deer Counts
The mean number of deer counted per night for 1969 and 1970 is compared
Fig. 2. The spring peak in 1970 (82.1) was nearly twice that in 1969
(42.2). There is also an indicated difference in the migration pattern

in

�- 102 between years.
In 1969, the spring deer counts dropped in late April;
while in 1970, they did not falloff
until late May.
The reason for this
is unknown.
The same observer counted each year and weather conditions,
at least during count nights, were not substantially different.
An index to deer use on the study field (mean number of deer per acre)
cannot be computed at this time. A question has arisen concerning the
actual area upon which deer are counted.
This will be answered during the
coming Segment.
The 1968, 1969, and 1970 deer count data are shown in Table 1. Linear
regression and correlation analyses were run on the pooled data to determine
what hour or combination of hours might be used in lieu of all-night counts.
Only the evening hours, through midnight, which generally produced the
highest counts were considered.
All correlation coefficients (r) were high,
0.903 to 0.988 (Table 2). Too, standard errors (Sy.x) were small, ranging
from 3.24 to 8.89. Very slight improvement in both rand Sy.x values
occurred with an increased number of hours and later counts.
On the basis
of these analyses, future deer counts at Rifle Gap will be made from 9:00
thru 11:00 P.M., MST (Fig. 3). An average of several counts is pre.ferred
to a single observation to reduce the effects of possible "odd" counts.
Figures 4 and 5 indicate the pattern of deer distribution observed during
nightly counts.
Major access points are at each end and at the middle of
the field across from the road. Again, as last year, there was a noticeable reduction of deer opposite the observatory indicating some deterring
effect by the close proximity of the counting station to the field.
It is
assumed deer frequent this area on nights when the observer is gone. This
will be checked in 1971.

Hay Harvest
The first cutting of hay yielded 2.3 tons per acre.
Plot yields revealed
an average 67 pounds less hay on grazed plots for a 1,216 pound difference
per acre (Table 3). A student's paired "t" analysis showed this difference
to be highly significant (P&lt;.Ol).
The second cutting of hay was not watered and yielded only 0.4 ton per acre.
The grazed plots yielded 32 pounds less hay on the average for a 581 pound
difference per acre (Table 4). This difference was not significant (P&gt; .05).
These production differences cannot be related to deer densities until
exact field acreages are determined as explained under "Deer Counts" above.

Proximate

Analysis

Results of the proximate analyses performed on hay samples from the first
and second cuttings are summarized in Table 5. None of the differences between grazed and ungrazed plots were significant (P&gt;.05) for any of the
factors tested.
Even had significance been shown, it could not be definitely
attributed to deer grazing, as plot pairing was on a production rather than
nutritional basis.
Therefore, it is doubtful that any meaningful results
will come from this portion of the study or that it will be continued.

�- 103 -

70
--·I9M

•

-1970

60

~

•
•
o

c:

:I"

23451234123412345123412345
Apr
May
Jun
Jul

-+-

-+

Montt.

-+-

+- Aug + Sep

a Week

Fig. 2. Mean number of deer counted each night on the Rifle Gap alfalfa
damage study field, 1969 and 1970.

�- 104 -

Table 1. Number of deer counted on the Rifle Gap alfalfa damage study field,
1968, 1969, and 1970.
Date of
Count 1900

2000

Hour of Count {Mountain Standard Time)
2100 2200 2300 2400 0100 0200 0300

0400

0500 Mean

26
12
62
42
12

66
58
96
89
110

48
41
64
87
78

41
59
39
68
67

35
46
18
72
65

33
43
27
67
17

27
10
16
35
49

9
8
29
46
70

8
19
22
52
54

4
16
24
52
84

2
7
18
l3
2

27.2
29.0
37.7
56.6
55.3

nc
40
nc
7
0
0
0
0
0
0
0
0
0
0
1
4

17
42
52
29
40
13
7
0
0
2
5
0
2
0
13
nc

9
27
47
45
32
8
9
6
6
4
4
6
6
7

22
38
56
33
16
16
4
2
2
0
4
6
6
14
16
14

15
47
54
25
10
6
13

9
49
39
32
25
14
2
3
1
0
5
4
8
10

9
28
47
15
l3
9
8
10
0
2
9
6
1
4
13
10

10
37
21
34
8
8

13

l3
28
46
14
10
9
10
3
5
0
4
6
2
6
12
21

7
38
31
15
6
3
5
0
0
0
0
1
2
2
17
19

6
14
29
2
0
8
0
0
0
0
0
0
0
0
2
6

11.7
35.3
42.2
22.8
14.5
8.5
6.3
3.1
1.4
1.8
3.9
3.5
2.8
4.7
10.7
11.2

19
56
72
23
17
36
37
11

0
104
93
72
73
98
118
45
1
4
0
0
0
0

21
55
71
34
54
69
82
52
17
6
4
11
8
6

0
48
49
17
57
58
84
42
20
6
4
6
4
5

0
55
55
1
35
71
88
53
l3
9
1
3
4
6

0
35
58
13
42
55
105
44
14
5
5
3
2
1

0
28
55
22
40
54
91
65
7
5
0
2
1

0
15
48
5
18
9
4
2
0
0
0
0
1
1

3.7
58.5
66.9
28.8
45.6
67.1
82.1
39.6
10.6
4.9
2.8
4.6
3.3
3.9

1968
4-10
4-17
4-24
5-1
5-8
1969
4-2
4-9
4-16
4-23
4-30
5-7
5-12
5-19
5-26
6-2
6-9
6-30
7-21
7-28
8-11
8-18

II

6

5

0
6
5
6
3
5
10
14

II

II

5
1
6
7
4
1
4
12
16

1970
3-31
4-6
4-14
4-22
4-28
5-4
5-11
5-18
5-25
5-31
6-8
6-16
6-22
6-28

0
0
0

0
0
0

0

93
52
50
39
99
95
51
14
6
4
14
7
II

1
88
87
42
61
96
116
45
17
6
4
2
4
4

0

67
96
38
66
93
83
26
14
7
9
10
5
9

0

-----------------------------------------------------------------------------nc = no count.

�- 105 -

Table 1. Number of deer counted on the Rifle Gap alfalfa damage study field,
1968, 1969, and 1970 (continued).
Date of
Count 1900

2000

Hour of Count {Mountain Standard Time}
2100 2200 2300 2400 0100 0200 0300

0400

0500

Mean

7-6
7-l3
7-20
7-27
8-4
8-11
8-25
8-30
9-7
9-13
9-20
9-27

8
12
0
14
7
11
5
12
27
7
4
32

14
19
10
23
20
16
20
19
8
8
12
34

5
8
0
7
4
10
4
9
3
9
l3
25

0
2
0
0
2
2
0
1
2
1
3
13

5.2
9.3
3.5
11.5
l3.5
12.5
12.1
16.0
11.1
9.7
12.8
23.4

0
1
0
0
5
3
0
2
0
4
1
22

7
14
1
24
24
18
18
26
8
20
8
30

12
7
15
18
23
24
24
20
14
16
16
28

1
15
5
16
19
14
29
28
15
8
21
22

0
11
2
15
17
11
17
22
16
14
28
19

3
7
2
9
21
15
14
15
16
11
16
15

7
6
3
1
7
13
2
22
l3
9
19
17

METHODS AND RESULTS - LITTLE HILLS
No data have come from the Little Hills portion of the alfalfa damage study
during the Segment. Therefore, progress will be reported under the combined METHODS AND RESULTS heading.
Deer Counts
Preliminary counts were made to ascertain relative numbers and use patterns
of wild deer on the western one-half of the study field at Little Hills.
Counting procedures were the same as at Rifle Gap. Four counts were made
at weekly intervals during April after which they were discontinued because
of a lack of deer. No deer were seen on the field the first two weeks.
The third week, 31 deer left the field early in the evening when the observer arrived. Three deer returned later that night and the same number
was observed the following week. If substantially more deer are not observed in 1971, the wild-deer portion of the Little Hills study will be
deleted.
Fencing an II-acre field for the controlled phase of the study was completed in early May. A herd of 30 deer (10 bucks, 8 does, and 12 fawns)
that had been confined in an adjacent 90-acre pasture were then allowed
free access to the field. Evidence of deer presence was later found over
the entire field and during one spot check, 27 deer were counted on the
area. Alfalfa hay and a pelleted concentrate mixture were fed on the study
field all winter to supplement a potential forage shortage in the pasture
and to encourage the animals to keep coming to the field.

�- 106 -

Table 2. Linear regressions and correlations relating the mean number of
deer counted at various hou~s (X) and the mean number of deer counted per
night (Y) on the Rifle Gap alfalfa damage study field using 1968, 1969,
and 1970 pooled data.

MST Time (PM)

n

8

46

9

47

10

47

11
12

r

r2

Sy·x

4.11 + 0.54 X

0.948

0.899

6.64

0.71 + 0.71 X

0.936

0.876

7.26

0.63 + 0.71 X

0.979

0.958

4.20

0.33 + 0.78 X

0.953

0.908

6.27

-0.48 + 0.89 X

0.903

0.815

8.89

2.03 + 0.64 X

0.956

0.9l4

6.08

-0.10 + 0.74 X

0.958

0.9l8

4.45

-0.02 + 0.76 X

0.985

0.970

4.23

-1.40 + 0.89 X

0.964

0.929

5.51

1.08 + 0.68 X

0.976

0.953

4.50

-0.63 + 0.78 X

0.985

0.970

3.59

-1.21 + 0.84 X

0.979

0.958

4.21

0.31 + 0.72 X

0.986

0.972

3.57

-1.25 + 0.84 X

0.986

0.972

3.43

-0.05 + 0.83 X

0.988

0.976

3.24

Prediction Equation
1\

Y
l\

Y

1\

47

Y

.1\

Y

1\

47

Y
1\

8+9
9+10

46

Y =

.1\

47

Y
1\

10+11

47

Y
1\

11+12

47

8+9+10

46

9+10+11

47

10+11+12

47

8+9+10+11

46

9+10+11+12

47

7+8+9+ 10+11+ 12

44

Y
1\

Y
.1\

Y
1\

Y
1\

Y
1\

Y
1\

Y

Irrigation
The entire field was sprinkler irrigated twice, once for each growth of hay.
The first irrigation ran from May 25 through June 16, and the second from
August 11 through September 5. Problems exist with the current irrigation
procedure in that about a three-week time lag occurs between the start and
finish of one complete irrigation. This creates an added variable in plant
growth that becomes especially acute during the relatively short growth
period of the second crop. New procedures will be considered in 1971 to
help alleviate this situation.

�- 107 -

100

90

-

•

80

~

GI

Z 70

n= 47
"'"

Sy.x = 3.57

•

•

•

•

•

.

Ya-0.63 + 0.18 X
r = 0.986

•
•
•

Mean Number of Deer at 9 a 10 a II PM, MST
Fig. 3. Relationship between the mean number of deer counted from 9:00
through .11:00 P.M. and the mean number of deer counted per night on the
Rifle Gap alfalfa damage study field based on 1968, 1969&gt; and 1970 pooled
data.

�- 108 -

-~- )c_II_J(_t~_)t~"
»&gt;

...-

,..--,

.

,.....•
~. , ••
\•• \
,/

•••'

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......"
.:-.o" ..\

• ,t.

y'

~\'\
1\.'

• I

0:", ••••• I ."

•

I •••••

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..~t~":'
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~
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•••••••

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'

•••

If

••••

"

•

I, • •

•

Ie

Q

•

.,

II

I,

• ,11

II

\

..' I '."'.,
I

~II'

11

II

• ,

,

'.

,

I

It

I ••
••
I
I •

"

e

0

.

••

,

\

,

I

I.

1

"

•

•

••

0

• ,
"

11

0

'

I

\.

.' •
•• •

&amp;

I
I •
I
\

••

•
'

I •

I
I

e ., It

"

••

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,
.J(

•I

•

., '..'. ..: r:,--,-, I
.,

:

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'.

•

K

)..

' ••
'~ ••
• 'Ii.

t···· . . .•.......
I

'.','1(

I

• .....
"'w

•

•
o

Fig. 4.

Observed

Gap alfalfa

damage

distribution

of groups of one or more deer on the Rifle

study field during the first hay growth, 1970.

�- 109 -

•

,_
.
..
....
• •••
•
\.:.....\ ..'. ·/~
1

..... . • •••
,
••••

,.
r

. ."

.'o .,0
• •

.0

\

•

•••••••

o ••

o

•

o

'\
\

o •

•

, •

•
•

I

.,.•• .. ':..:
.. •••

\

0

\

\

\

z

0

l

.0

•
o

'\

I•
.f
I

•

•
•S·

•
.' :. •
.,., .... .':•
, . ••
., •
·'" ....•,...~}
•
• •
•
.." . • •
.••••
•
••
,I.,..:. •
•

I • ••
\

o

....

....

•

....

.... ;-

....

..'

I
I

I.

01. •
o

I

!
/

.....• • ,. • '.. I
IC

o

J(

I

,
,

.•••••.• •. . I
.

,

0·.

I

"

I

I

)(

I

I •
,

\\
1

I

•

~

I

•

••

;I(

• I

••
•

I

I•
,..........
\

J

It

••

I

;I(

••

' •••
' ~.
.....

• ••••
.~

",

. /'&lt;
J&lt;

l
Fig. 5.

Observed

Gap alfalfa

distribution

of groups of one or more deer on the Rifle

damage study field during the second hay growth, 1970.

�- 112 -

Table 5. Results of proximate analysis of hay samples from the Rifle Gap
alfalfa damage study field, 1970. (n = 14)
First Cutting
Grazed Ungrazed

Second Cutting
Grazed Ungrazed

Analysis

(%)

(%)

Calculated-II
"t".05, 13df

Protein

8.25

7.94

0.608

14.81.

15.89

0.553

Fat

2.57

2.49

0.285

3.92

3.66

1.386

Fiber

49.92

51.19

0.619

41.90

42.10

0.073

Ash

6.09

6.09

0.007

7.27

8.07

1.348

Nitrogen
Free Extract 33.16

32.01

0.717

32.09

30.28

0.725

11 Tabular "t".05, 13 df

(%)

(%)

Calculated11
"t".05, 13df

2.160.

LITERATURE CITED
Gordon, D. F. 1969. Evaluation of the effects of spring-summer grazing
by deer on alfalfa. P. 25-43. In Game Research Report. Colorado Div.
Game, Fish and Parks, Denver. 3{Part 1):1-139.

Prepared by

_
David F. Gordon
Assistant Wildlife Researcher

{/~-L_/ 7/~~
Richard M. Bartmann~
Assistant Wildlife Researcher

�July, 1971

- 113 -

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~---------

Project No.

W-38-R-25

Work Plan No.

llC

Job Title

Deer-Elk Investigations
Job No.

1

White River Elk Population Components

Period Covered:

April 1, 1970 through March 31, 1971

Personnel:

Richard M. Bartmann, George D. Bear, Raymond J. Boyd,
Richard N. Denney, Donald M. Hoffman, Richard M. Hopper,
Robert E. Keiss, Gary T. Myers, Thomas M. Pojar, Edgar J.
Prenzlow, William H. Rutherford, Robert L. Schmidt,
Warren D. Snyder, Steven F. Steinert, Michael R. Szymczak
and Harold M. Swope. Edgar J. Prenz10w had been in charge
of this job since 1966, but in mid-1970 he was transferred
to Denver and the writer was reassigned to the job in order
to complete the investigation which will end with the
termination of Segment 26.

ABSTRACT
This is the fifth annual progress report of a five-year investigation to
determine the effects of specified permit seasons (limited numbers of both
antlered and antlerless permits available to hunters) on the White River
elk herd in northwestern Colorado. Estimated herd size, determined by
formulas using pre and post season sex and age ratios in combination with
kill levels, has decreased from about 6,800 to approximately 6,300 from
1966 to 1970. Preseason sex ratios (bulls per 100 cows) increased from
about 29 to about 33. Age ratios (calves per 100 cows) decreased from 58
to about 55. The changes in sex ratios resulted from issuing more antlerless and fewer antlered permits to hunters in an attempt to stabilize the
female segment of this elk herd. Procedures used to determine kill rates,
sex and age ratios and age structures of the herd are compared and evaluated.
Data from check stations and random surveys are also included.

��- 115 -

WHITE RIVER ELK HERD - POPULATION COMPONENTS
Raymond J. Boyd
P. S. OBJECTIVE
To develop a harvest formula for the White River elk herd.
SEGMENT OBJECTIVE
Test a candidate elk harvest formula for the White River elk herd.
METHODS AND MATERIALS
Data collection (aerial and ground sex and age ratio counts, check stations
and mail questionnaires to determine harvest, age composition, etc.) for
this project have not varied appreciably since its initiation in 1966
(Prenzlow 1967, 1968, 1969, and 1970). Procedures for determining total
elk populations have also remained the same.
DESCRIPTION OF AREA
Complete descriptions of the area covered by the White River elk herd are
included in Harris (1963) and Boyd (1971). Figure 1 locates the study area
in general and Area E (Game Management Units 23 and 24) specifically. Other
descriptions of the study area can be found in Prenzlow (1967 and 1968).
RESULTS AND DISCUSSION
Sex and Age Composition
Aerial Surveys
Pre-season Sex and Age Ratios--The 1970 pre-season sex and age ratio counts
of the elk population in Area E were made with the aid of a helicopter on
September 21 through 26, 1970. These classification counts were confined
to elk observed only in Area E (Table 1). A total of 854 elk was classified,
resulting in a ratio of 32.97 bulls (inculding spikes) per 100 cows and 54.7
calves per 100 cows. Summaries of pre-season sex and age ratio counts for
the period 1961 through 1970 are shown in Table 2; while Figures 2 and 3
illustrate the 95 percent confidence limits around these ratios:

�116 -

N

~--I
•

AREA

LEGEND

Fig. 1. Elk Management Area E (Game Management Units
23 and 24) showing boundaries, drainage patterns and
highway systems.
Solid circles are active check
station locations.

Check Stations

STUDY

GAME

MANAGEMENT

UNIT

BOUNDAR'\'
HIGHWAY

",=-7

SECONDARY

tt

RIVER

-

CREEK

~-::-

PEAK

J;,

LAKE

••

ROAO

-

---

�- 117 .,..
Table 1.

Pre-season sex and age ratio classifications in Area E, 1970.

Location

Mature
Bulls

Highway 13 to Yellowjacket

Young
Bulls

Spike
Bulls

Cows

Calves

Total

1

2

10

3

16

Coal Creek

1

1

Lost Creek

1

1

Lost Park area

4

5

12

45

23

89

Snell Creek, Ripple Creek

3

1

14

39

18

75

Anderson Res. , Skinnyfish Cr.

5

4

4

39

27

79

6

3

9

6

3

10

43

27

89

20

9

29

3

3

7

9

3

14

Crooks Park
1

Ute Creek
Marvine Creek

2

3

14

Big Ridge
1

Big Fish Creek
Trappers Creek

2

South Fork

7

6

16

62

36

127

Park Creek

2

5

2

41

20

70

Lost Solar Creek

1

2

2

28

14

47

Fowler Bench

5

3

37

21

66

2

6

6

14

3

3

22

11

40

2

5

23

15

45

3

14

7

26

Johnson Park
Burro Mt., S. side S. Fk.

1

Patterson Creek
Oak Ridge

2

Total

34

34

82

455

249

854

Percent

4.0

4.0

9.6

53.3

29.1

100.0

�Table 2.

Summaries

of pre-season

sex and age ratios from Area E, 1961-1970.

Bulls

Cows

._-

Calves
%

Total

Bulls

:

Ratio
Cows

:

Calves.

216

33.1

653

41.4

:

100

:

69.9

47.4

298

30.5

976

46.8

:

100

:

64.5

298

47.5

204

32.5

627

41. 9

:

100

:

68.5

17.5

363

52.9

203

29.6

686

33.1

:

100

:

55.9

233

17.3

699

51.8

416

30.9

1,348

33.3

:

100

:

59.5

86

15.4

299

53.7

172

30.9

557

28.8

:

100

:

57.5

Year

No.

%

No.

1961

128

19.6

1962

216

1963

%

No.

309

47.3

22.1

462

125

20.0

1964

120

1965
1966

I-'
I-'

00

1967

134

16.8

415

52.2

247

31.0

796

32.3

:

100

:

59.5

1968

135

18.7

366

50.7

221

30.6

722

36.9

:

100

:

60.4

1969

324

23.9

645

47.5

389

28.6

1,358

50.2

:

100

:

60.3

1970

150

17.6

455

53.3

249

29.1

854

32.9

:

100

:

54.7

Average

165

19.2

431

50.2

261

30.6

858

38.3

:

100

:

60.6

�- 119 -

60

SO
••
~
0

U

~

--

co

0
0

•.. 30
.!!
:&gt;

co

20

0
1961

62

63

65

64

66

67

68

69

70

Years

Fig. 2 •. Confidence intervals. on pre-season sex ratios, Area E, 1961- 1970

10

••• 70
~
0

U
0 60
0
•....

1

--..?

•.. 50

II

0

U 40

:
0

)961

62

63

64

6S

66

67

68

69

70

Years

Fig. 3. 'Confidence .intervals on pre-season age ratios (ca1ves:100 cows),
Area E, 1961- 1970.

�- 120 -

Post-season Sex and Age Ratios'--Post-season sex and age ratio classifications
of the elk population in Area E were conducted, by helicopter, on January 5
through January 8, 1971. A total of 2,406 elk were classified resulting in
a ratio of 8.56 bulls (including spikes) per 100 cows and 63.05 calves per
100 cows (Table 3). Summaries of post-season classifications for the period
1961 through 1970 are included in Table 4, while Figure 4 indicates the 95
percent confidence limits around these post-season ratios.

Table 3.

Post-season sex and age ratio classifications

in area E, 1970.

Mature
Bulls

Location

Young
Bulls

Spike
Bulls

Cows

Calves

Total

Hogback

6

1

5

54

26

92

Highway 13 to Yellowjacket

4

3

8

152

80

247

3

2

76

45

126

Uranium Peak area
Coal Creek, Little Beaver

4

3

88

59

154

Oak Ridge - North

2

6

135

90

233

Oak Ridge - South

4

3

10

262

174

453

Big Beaver, Fawn Creek

2

3

4

127

80

216

22

15

37

3

28

13

53

Marvine Creek

1

14

11

26

Ute Creek to S. Fork

2

72

55

129

15

326

206

555

8

46

30

85

Fawn Creek to Lost Creek
Lost Creek to Snell Creek

9

5

South Fork

3

Miller Creek

1

Total

35

18

67

1,402

884

2,406

Percent

1.4

0.8

2.8

58.3

36.7

100

�Table 4.

Summaries of post-season sex and age ratios from Area E, 1961-1970.

%

No.

%

Total

Bulls

:

Ratio
Cows

:

Total

Calves

Cows

Bulls
Year

No.

%

No.

1961

106

10.6

465

46.5

430

43.0

1,001

22.8

:

100

:

92.5

1962

125

13.8

446

49.2

335

37.0

906

28.0

:

100

:

75.1

1963

45

6.1

413

55.7

283

38.2

741

10.9

:

100

:

68.5

1964

117

6.0

1,130

58.3

692

35.7

1,939

10.4

:

100

:

61.2

1965

85

5.2

930

57.1

613

37.7

1,628

9.1

:

100

:

65.9

1966

128

6.1

1,245

58.9

739

35.0

2,112

10.3

:

100

:

59.4

1967

214

9.2

1,318

56.4

806

34.5

2,338

16.2

:

100

:

61.2

1968

148

8.2

951

52.9

699

38.9

1,798

15.6

:

100

:

73.5

1969

166

9.0

1,012

55.0

661

36.0

1,839

16.4

:

100

:

65.3

1970

120

5.0

1,402

58.3

884

36.7

2,406

8.6

:

100

:

63.0

Average

125

7.5

931

55.7

614

36.8

1,670

13 .4

:

100

:

66.0

t-'
N
t-'

�- 122 -

so
'" 40

~
o

U

30

o
o

..:::.. 20
w&gt;

o L----1-9~6-'--~-6-2------6~3-----6·-4------6~S----~6~6-----6~7-----6~8~--~6~9~--~70
Years

Fig.

Ground

I,.

Confidence

intervals

on post-season

sex ratios,

Area E, 1961-

1970.

Surveys

During the years 1967 through 1970, a ground classification of elk in Area E
was made each July.
This was done to estimate yearling-adult cow ratios,
which cannot be accurately obtained during aerial classifications.
These
data also serve as independent estimates of sex and age structures.
It
becomes very important to determine the proportion of yearling females in
the herd and thus their influence on overall herd productivity.
Usually
only a small fraction of the yearling females breed, but there is considerable conflicting literature on this particular subject.
Observers on horseback, riding during the early morning and evening hours,
classified elk in different sections of the Flat Tops Primitive Area.
In
three days, four teams of two or more men classified a total of 969 elk
(Table 5). Many more elk were seen on which no classifications were possible.
The average size of elk groups observed was 32.3 animals.
Check Station Surveys
Number of Elk Checked--Four special research check stations (Meeker, New
Castle, Deep Creek, and Ripple Creek) were operated during the first nine
days of the 1970 big game season.
Research personnel were also stationed
at the permanent management check station at Rifle to check elk from Area E.
A total of 1,154 elk was checked from Game Management Units 12, 13, 23, 24,
25, 26, 33, and 34. Data were collected from six additional units around
Area E (Units 23 and 24) because of their importance in relation to Area E
elk. Table 6 lists number of elk checked from the White River area 1964
through 1970.

�- 123 -

Table 5. Composition of 969 elk classified during July ground counts in
Area E, 1970.
'Mature Yearling
Bulls
Bulls

Item

Mature
Cows

Yearling
Calves
Cows

Total

Number Observed

52

65

440

133

279

969

Percent Observed

5.4

6.7

45.4

13.7

28.8

100.0

Frequency in Groups 1/

16

17

28

22

20

30

Mean Group Composition ~/

1.7

2.2

14.7

4.4

9.3

32.3

1/ Frequency in groups (i.e. , mature bulls were observed in 16 of 30 total
groups).
~/ Mean group composition determined by percent of elk observed times 32.3.

Table 6. Number of elk checked through stations from White River elk herd,
(Game Management Units 12, 13, 23, 24, 25, 26, 33, and 34), 1964-1970.
Station

1964

1965

1966

1967

1968

1969

1970

Average

Meeker

553

506

591

518

725

637

713

606

Deep Creek

338

227

220

303

308

139

189

246

New Castle

321

196

166

227

290

78

164

206

Rifle

137

138

124

92

83

73

51

100

Idaho ~prings

32

137

159

112

120

129

Ripple Creek

227

56

37

1,381

1,204

1,384

114
4

4

Hamilton
Total

110

1,381

1,633

983

1,154

1,303

�Table

9.

Sex and age determinations

of 452 elk checked

from Area E, 1970.

Age

~:lears2 ]j

k2

l~

2~

3~

4~

5~

6~

7~

8~

9+

'I'o
t.al :

Number

19

202

70

11

11

2

4

3

0

1

323

Percent

5.9

62.5

21.7

3.4

3.4

0.6

1.2

0.9

0

0.3

100

4.2

44.7

15.5

2.4

2.4

0.4

0.9

0.7

0

0.2

71.4

Sex

Males

Percent

Total

Females
t-'

N

Number

36

14

17

20

13

8

12

3

2

4

129

Percent

27.9

10.8

13.2

15.5

10.1

6.2

9.3

2.3

1.6

3.1

100

8.0

3.1

3.8

4.4

2.9

1.8

2.6

0.7

0.4

0.9

28.6

55

216

87

31

24

10

16

6

2

5

452

12.2

47.8

19.2

6.8

5.3

2.2

3.5

1.3

0.4

1.1

100

Percent

Total

Total Elk
Number
Percent

1/ Ages

Total

assessed

by Quimby

and Gaab

technique.

0'\

�Table 10.

Sex and age determinations of 452 elk checked from Area E by unit of kill, 1970.
Age ~years2 1/
5lz
4lz

6lz

7lz

8lz

9+

Total

4

1

1

1

0

0

115

3.48

3.48

0.87

0.87

0.87

8.36

1.24

1.24

0.31

0.3i

0.31

126

43

7

7

1

3

2

0

1

208

8.65

60.58

20.67

3.36

3.36

0.48

1.44

0.96

-

0.48

5.57

39.01

l3 .31

2.17

2.17

0.31

0.93

0.62

8

4

5

5

4

1

4

1

0

0

32

Percent of Total

25.00

12.50

15.62

15.62

12.50

3.12

12.50

3.12

Percent of all
Females

6.20

3.10

3.88

3.88

3.10

0.78

3.10

0.78

28

10

12

15

9

7

8

2

2

4

97

Percent of Total

28.87

10.31

12.37

15.46

7.22

8.25

2.06

2.06

4.12

Percent of all
Females

21. 70

7.75

9.30

11.63

5.43

6.20

1.55

1.55

3.10

k2

llz

2lz

3lz

1

76

27

4

Percent of Total

0.87

66.19

23.48

Percent of all Males

0.31

23.53

18

Percent of Total
Percent of all Males

Unit and Sex

23 - Males
Number

24 - Males
Number

•....
N
-...J

23- Females
Number

24 - Females
Number

1/ Ages assessed by Quimby and Gaab technique.

9.28

6.98

�- 128 -

Table II. Percent error in aging 210 yearling
White River area units, 1970.

Station

Dental
Cementum

Meeker

11

Field
Aged

1/

elk at check stations,

all

Percent
Correct

Percent
Error

152

149

98.03

1.97

Ripple Creek

2

2

100.00

0.0

New Castle

35

35

100.00

0.0

Deep Creek

l3

l3

100.00

0.0

Rifle

8

8

100.00

0.0

Total

210

207

98.57

1.43

II Number of teeth checked

in the laboratory

using the cementum

technique.

~I Number of correct ages determined at check stations using tooth wear and
replacement

techniques.

Table 12. Percent error in aging 242 elk older than yearlings
all White River units, 1970.

Station

Dental
Cementum

Meeker

11

Field
Aged

,{I

at check stations,

Percent
Correct

Percent
Error

155

79

50.97

49.03

Ripple Creek

9

7

77.78

22.22

New Castle

42

22

52.38

41.62

Deep Creek

31

22

70.97

29.03

Rifle

5

0

0.0

Total

242

130

53.72

II Number of teeth checked

in the laboratory

using the cementum

46.28

technique.

II Number of correct ages determined at check stations using tooth wear and
replacement

techniques.

�- 129 Table 13. Percent error between dental cementum and tooth replacement and
wear aging tehcniques by age class of 471 elk from the White River study
area, 1970.

Dental
Cementum

Age
Class

1/

Field
Age ~/

Percent
Correct

Percent
Error

1

213

207

97.2

2.8

2

95

80

84.2

15.8

3

79

23

29.1

70.9

4

33

12

36.4

63.6

5

17

4

23.5

76.5

6

13

3

23.1

76.9

7

7

1

14.3

85.7

8

5

o

o

100.0

9+

9

5

55.6

44.4

471

335

71.12

28.9

Total

1/ Number of teeth checked in the laboratory using the dental cementum
procedure.
~/ Number of ages correctly
replacement procedures.

assigned

at check stations

Table 14. Distribution of assigned ages in relation
dental cementum procedures, Area E only, 1970.

Cementum
Age Class

Check Station Assigned
34567

using tooth wear and

to ages determined

by

Ages
8

9+

Total

1

2

1

119

1

1

2

1

60

24

3

2

7

49

4

2

9

17

3

1

5

3

14

1

6

3

6

9

1

2

7

1

1

1

5

1

1

10

1

4

1

6

7

7

10

369

4

121
1

86

2

64

8

34

2
1

9+
Total

122

70

83

28

24

15

9

8

1

20
21

�- 130 -

Of the 1,900 hunters sampled, 1,419 (74.68%) returned usable cards. The
estimated total elk kill in Area E was 1,360 (C. I. ± 134; Z = 1.96), with
38.4 percent of the active hunters successful. Table 15 gives results of
the survey, including kill by type of license, while Table 16 lists active
(those who actually hunted elk in 1970) hunters and success ratios.

Table 15.

Results of the 1970 random survey of 3,810 Area E permittees.
Antlered
Res.
N. .:~:.cs•

Ant1erless
N. Res.
Res .

Total

Permits Issued

2,159

846

633

172

3,810

Permits in Sample

1,069

431

318

82

1,900

Percent in Sample

49.51

50.94

50.24

47.67

49.87

Sample Returned

748

299

294

78

1,419

Percent Returned

69.97

69.37

92.45

95.12

74.68

Percent of Total Permits

34.65

35.34

46.44

45.35

37.24

Reported "No Hunts"

63

17

17

2

99

Estimated "No Hunts"

168

45

46

6

265

Reported "Kills"

214

108

134

50

516

Estimated Total "Kills"

571

295

357

137

1,360

Table 16. Number of hunters participating during the 1970 Area E elk hunt
and success ratios per license and active hunter. Projected from random
survey data.

Active Hunters

1,991

801

587

166

3,545

Success (kill)/Lic.

26.45

42.20

46.60

79.65

35.70

Success (kill)/Hunter

28.70

44.60

50.30

82.50

38.40

Projections from the random survey indicated that 265 licenses were issued that
were not used (7.0%). The number of "no hunts" in Area E have averaged 7.0
percent since 1966.

�- 131 -

Harvest

in Relation

to Area and Time

Location of Kil1--Ki1l by Game Management Unit was projected
random survey and appears in Table 17. These data indicated
percent of the 1970 Area E kill occurred in Unit 24.

Table 17.
Management

Composition
Unit.

of the 1970 estimated

Unit

Antlered
Number

Antlerless
Number
%

%

from the
that 54.12

elk kill in Area E by Game

Total
Number

%

23

395

29.04

229

16.84

624

45.88

24

471

34.63

265

19.48

736

54.12

Total

866

63.68

494

36.32

1,360

100.00

Date of Kill--Dates of kill by hunters in Area E are shown in Table 18.
Approximately 74.8 percent of the kill occurred during the first five days
of the season, with 90.15 percent occurring during the first 11 days of the
season.

Hunter Effort--Elk hunting in Area E provided 20.954 man-days of hunting
recreation in 1970. This was an increase of 7,845 recreation days compared
with 1969. This increase was due, in part, to a 36 percent increase in the
number of bull permits in Area E compared to 1969. Mean number of days
spent hunting per active hunter increased from 4.6 in 1969 to 5.9 in 1970.

Estimates

of Mortality

Other Than Legal Kill

Wounding Loss--Estimates
of net wounding loss (number of elk reported
wounded minus the number of wounded elk killed by other hunters) are presented in Table 19, while in Table 20, a comparison is presented between
wounding loss reported by hunters with antlered licenses and those having
a antlerless license.
It is assumed these wounding loss figures represent
a minimum loss because all hunters probably would not admit wounding an elk,
and there are elk wounded that hunters are not aware of. Conversely,
hunters who did not wound an elk, most likely would not respond to the
question positively.
The 5.8 percent loss figure from Table 20, is about
one-half that reported in 1969 from Area E.

�Table 18.

Numbers of elk killed, by day of season, Area E, 1970.
October

November

Unit

17

18

19

20

21 1/ 22

23

24

25

26

27 ~/ 28

29

30

31

1

2

3

4

5

6

23

164

85

90

68

37

31

20

8

31

8

14

14

6

3

11

8

-

6

6

6

8

24

189

133

llO

82

59

37

17

14

11

6

11

6

17

6

8

31

-

-

-

-

6

I-'

W
N
I

Total

354

218

200

150

96

68

37

22

42

14

25

20

23

9

19

39

0

6

6

6

1/ 74.8 percent of the total estimated elk kill occurred during the first five days of the open season.
1/ 90.15 percent of the total estimated elk kill occurred during the first eleven days of the open season.

14

�Table 19.

Wounded elk salvaged by hunters and net wounding loss, Area E, 1970.
Wounding Loss
Net Loss
Wounded Killed

Number of Elk
Reported Killed

No. Wounded

Antlered

322

24

7

17

5.3

Ant1er1ess

184

6

2

4

2.2

Total

516

30

9

21

4.1

License Type

Percent Loss

t-'

Table 20.

w
w

Area E wounding loss - comparison of antlered and ant1er1ess hunters, 1970.

License Type

No. of
Hunters Reporting

No. of Elk
Rept. Wounded

% of Hunters
Wounding Elk

No. of Elk
Rept. Killed

% Loss
of Kill

Proj.
Elk Loss

Antlered

1,047

24

2.3

322

7.4

49

Ant1er1ess

372

6

1.6

184

3.3

8

1,419

30

2.1

516

5.8

57

Total

�- 134 -

Population

Total Population

Estimates

and Projections

Estimates

Formulas were used to estimate the total number of elk present in Area E
prior to the 1970 elk hunt.
Detailed descriptions of the method are
presented in Prenzlow (1968) with examples of how ratios of antlered to
antlerless elk from kill, pre- and post-season classification counts,
were used in a formula devised by Dr. David Bowden of Colorado State
University, to estimate a pre-hunt elk population.

Pre-Season Population Estimate, 1970--An estimated 6,347 (C.l. = 162, ~
1.96, q(= .05) elk were present in Area E just prior to the opening of
the 1970 elk hunt (Table 21). This compares to 6,751, 6,313, 6,216 and
5,829 estimated in 1966, 1967, 1968 and 1969, respectively.
Since the
research and management objective in this area was to hold the elk herd
stable, the estimated pre-hunt elk population seemed reasonable.

Table 21. Pre-season population
and age in Area E, 1970.

estimate

and proportions

of elk by sex

Item

Estimated
Number

Expected
Percent II

Observed
Percent 11

Bulls

1,117

20.99

17.56

Cows

3,383

51.08

53.28

Calves

1,847

27.93

29.16

Total

6,347

100.00

100.00

II Percentages

expected

11 Percentages

observed when making

from 1969 post-season

population

1970 pre-season

data.

elk classifications

by

helicopter.

Also, the expected percentages of bulls, cows and calves, based upon 1969
post-season population estimates, were not significantly different from percentages observed when flying in September on sex and age classification
counts in 1970.

�- 135 -

Post-Season Population Estimate, 1970--An estimated 4,987 elk survived the
1970 elk hunt in Area E (Table 22). This population was determined by
subtracting the elk kill, as determined by the random survey, from the
estimated pre-season population.

Table 22. Post-season population
sex and age in Area E, 1970.

estimates

and proportions

of elk by

Item

Pre-Season
Population

Kill

Bulls

1,117

866

251

5.03

4.98

Cows

3,383

453

2,930

58.75

58.27

Calves

1,847

41

1,806

36.22

36.75

Total

6,347

1,360

4,987

100.00

100.00

number

of elk (4,987) that

1/

1/ Based upon 1970 random survey.
1/ Proportions by class of the total estimated
remained

Post-Season Population
No.
Exp. % 1/ Obs. % 2/

after the close of the 1970 elk hunt.

2/ Proportions

actually observed when flying the 1970 post-season
age classifications by helicopter.

sex and

Total post-season elk populations in Area E declined from 5,336 in 1966 to
4,506 in 1968, then increased slightly to 4,646 in 1969 and 4,987 in 1970.
The decline was largely the result of an increased kill in the antler less
segment of the elk population.
Allowing more antler less permits during
these years was necessary, however, to change the composition of the
wintering herd to meet the objective of narrowing the sex ratio (more bulls
per 100 cows). Proportions of bulls in the post-season elk population have
increased from 6.1 percent in 1966 to 9.2, 8.2, and 9.0 in 1967, 1968 and
1969 respectively, then decreased in 1970 to 4.98 percent.

Population

Projections

Each year, based upon the previous years post-season population estimate,
projection is made for the next year.
This procedure is described in
Prenzlow (1968) in detail using 1967 data as examples.

a

Population Projection for Area E, 1971--Immediately
prior to the opening of
the 1971 elk hunt in Area E, a projected elk population of 7,084 animals will
be available to hunters in this area (Table 23). This figure may change
slightly, however, if productivity differs significantly from 54.7 calves
per 100 cows.

�- 136 -

Table 23.

Predicted

Item

1971 pre-season

elk population,

1970 Post-Season
Population

Bulls

251

Cows

2,930

Calves

1,806

Total

4,987

1971 Pre-Season
No.

1/

that of 1,806 calves, 50 percent
(903) are females.
by adding the remaining

POEulation
Exp. % 2/

u

16.29

3,833 ]j

54.11

2,097 ~/

29.60

7,084

100.00

1,154

1/ Assuming

]j Calculated

Area E.

(903) are males and 50 percent

bulls in 1970 (251) and 803 male calves

from 1970.

1/ Calculated

by adding the remaining
calves from 1970.

cows in 1970 (2,930) and 903 female

~/ 'Calculated by multiplying the number of cows in 1971 (3,833) by 54.7
percent expected productivity.

2/ Expected

proportions of total projected
the 1971 hunting season.

numbers

of elk (7,084) prior to

In previous years, population projections have been high because no losses
other than the estimated hunting mortality were considered.
Annual herd
losses of 17.5, 11.5 and 4.3 percent were determined for 1967, 1968 and 1970,
respectively.
These percentages were calculated by subtracting a particular
years popuLat Lon estimate from the projection (Table 24).

Recommendations
for the 1971 Area E Elk Hunt--To fulfill objectives 3a and
3b of this study, it is our recommendation that 2,150 antlered and 1,110
antlerless permits be issued for the 1971 elk hunt in Area E. Table 25
illustrates the composition of the 1971 pre-season Area E elk population if
the projection is reduced by 10 percent to account for over-winter herd
losses.
The resulting post-season population is projected also based on
no significant change in hunter success.

�- 137 -

Table 24.

Estimates

of annual herd losses in Area E, 1966-1970.

Year

Population
Estimate

1966

6,751

1967

6,313

7,654

1,341

17.5

1968

6,261

7,075

814

11.5

1969

5,829 1/

5,829

1970

6,347

6,632

285

4.3

of obvious sampling

errors.

1/ Corrected

because

Population
Projection

Difference
Number
Percent

Table 25. Modified 1971 Area E pre-season population projection and resulting post-season population with a kill level determined by 2,000 antlered
and 1,400 antlerless permits.

Item

Modified 1971 PreSeason POEulati0n
Number
Exp. %

Bulls

1,039

16.30

560

479

9.63

Cows

3,450

54.11

748

2,702

54.30

Calves

1,887

29.59

92

1,795

36.07

Total

6 ,376 1/

100.00

1,400 ]j

4,976

100.00

1/ Modified
theoretical

projection
projection

1971 Post-Season
Projected
Kill

resulted from subtracting
of 7,084 in Table 26.

POEulation
Number
Exp. %

708 elk (10%) from the

1/ Projected

kill is based on 28 percent success on 2,000 antlered
and 60 percent success on 1,400 antlerless permits.

The 479 bulls and 2,702 cows in the post-season
18.39 bulls per 100 cows.

population

permits

give a ratio of

�- 138 -

As a check on the above projections, ratios of bulls, cows and calves
observed during classification counts will be compared to expected ratios;
kill data will again be determined by random survey; and pre-season population estimates will be calculated by formula.
It is important to note
that these comparisons will be independent estimates of the same population
parameter, and if there is close agreement, it would indicate that the
above described method is valid for estimating the elk population in Area

E.

LITERATURE

CITED

Boyd, R. J. 1971. Elk of the White River plateau, Colorado.
Fish and Parks Div., Denver.
Tech. Pub. No. 25. 126 p.
Harris, J. T.
Colorado.

Colo. Game,
illus.

1963. Population dynamics of the White River elk herd,
Ph.D. Thesis.
Univ. Mich. Ann Arbor, Mich.
190 p.

Keiss, R. E. 1969. Comparison of eruption-wear patterns and cementum
annuli as age criteria in elk. J. Wildl. Mgmt.
33(1):175-180.
Prenzlow, E. J. 1967. Population components - White River elk.
Job Completion Report, W-38-R-21.
p. 251-276.

Colorado.

1968. White River elk population components.
Progress Report.
W-38-R-22.
p. 383-421.

Colorado.

Job

1969. White River elk population components.
Progress Report.
W-38-R-23.
p. 179-234.

Colorado.

Job

1970. White River elk population components.
Progress Report.
W-38-R-24.
p. 257-286.

Colorado.

Job

Quimby, D. C., and J. E. Gaab. 1957.
indicator in Rocky Mountain elk.

Prepared by

Mandibular dentition as an age
J. Wildl. Mgmt.
21(4):435-451.

�- 139 -

APPENDIX A
POST CARD SURVEY - AREA E HUNTERS

Please list results from your own elk hunting effort and
not information from those with whom you hunted in 1970.
1. Did you hunt elk in your specified area? Yes

2. What Game Mana ement Unit within the AREA did
# of
nit

County

Drainage

3. Did you harvest an elk? Yes
No
Calf
Bull
Cow
5. List unit of harvest:
6. List county of harvest:
7. Please give date of harvest:
8. How many elk were hit that you weren't able to get?
9. Was your elk wounded before you killed it? Yes
No

4. Did you harvest a:

DIV
USE
ONLY

��July, 1971

- 141 -

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W ....
38-R-25

Work Plan No.

14

Job Title

Deer-Elk Investigations
Job No.

1

Middle Park Deer Study - Population Distribution

Period Covered:

April 1, 1969 through March 31, 1971

Personnel:

R. Bruce Gill, D. W. Reichert, Olof C. Wallmo, Robert L.
Schmidt, Julius J. Klein, Willard Travnicek, Steven Horn,
Don Merrimen, Dave Hoart, Paul Gilbert, and Laren A. Roper.

ABSTRACT
A complete revised list of deer trapping and marking data is presented for
1969, 1970 and 1971. The total deer trapped in five separate areas for
these years are 279, 205 and 193.

��- 143 -

MIDDLE

PARK DEER STUDY - POPULATION

DISTRIBUTION

Laren A. Roper

P. S. OBJECTIVE
To delineate deer concentration areas in Middle Park and relate changes
in deer distribution in time and space to accumulation and physical
properties 'of snow.

SEGMENT OBJECTIVES
1.

Define winter range boundaries of the Middle Park deer population
under mild, moderate, and severe winter conditions.

2.

Define sub-unit boundaries.

3.

Delineate

4.

Relate changes in deer distribution
properties of snow.

major deer concentration

METHODS

areas.
to accumulation

and physical

AND MATERIALS

See Gill (1969).

RESULTS AND DISCUSSION

Trapping

Trapping

and Marking

Results

Trapping was conducted within five suspected deer population sub-units
was reported by Gill (1970). These units are Muddy Creek, Blue River,
Williams Fork River, Troublesome Creek and Beaver Creek.

as

Field records for the 1969, 1970, and 1971 deer trapping data have been
reexamined and summarized on automatic data processing cards. As a result
several errors were found that were reported in the July 1970 report.
This
summary supersedes the previous data.
Trapping success has increased each year of the trapping operation, however,
the total deer tagged each year has decreased due to re-catches of previously tagged deer. In 1969 a total of 279 deer were trapped and tagged,

�- 144 -

205 in 1970 and 193 deer in 1971. These represent newly tagged deer.
Table I is a summary of the total deer tagged in each sub-unit.

Table l. The number of deer tagged in 5 areas suspected
populations of deer in Middle Park.

Year

Muddy

Blue

1969

102

36

18

1970

13

56

1971

49

Total

164

Williams

Fork

Troublesome

to be discrete

Beaver

Total

102

21

279

42

94

0

205

47

43

36

18

193

l39

103

232

39

677

Tables 2 through 15 provide revised lists of deer tagged with either eartags and/or neckbands for the 3-year period.
All trap locations have been coded for automatic data processing.
16 provides a list of these sites and their code number.
A complete analysis
the next segment.

Neckbanded

of these data is in progress

Table

and will be reported in

Deer Observations

These data are currently being analyzed utilizing an automatic data processing program that has been developed this past segment to summarize all
trapping data.

�- 145 -

Table 2. Deer tagged' and marked in the Muddy sub-unit of Middle Park,
Arapahoe National Forest, 1969.

Date
Tagged

1-16-69
1-16-69
1-17-69
1-17-69
1-18-69
1-18-69
1-20-69
1-24-69
1-26-69
1-26-59
1-26-69
?

2-5-69
2-5-69
2-5- 69
2-5-69
1-22-69
1-23-69
1-25-69
1-29-69
1-29-69
1-30-69
2-1-69
2-2-69
2-2-69
2-2-69
2-2-69
2-3-69
2-4-69
2-4-69
2-4-69
2-4-69
2-5-69
2-5-69
2-6-69
2-6-69
2-7-69
2-7-69
2-7-69
2-8- 69
2-9-69

Age

Tag No.

Neck nand Color
and Symbol

Mature
F
Fa\\TJ1
F
Nature
F
Hature
F
Hature
F
Hature
H
Yr l.g ,
H
Hature
F
1'1 Yrlg.
Fawn
F
1'1 Fawn

MP-2
1vlP5
1'1P-8
MP-9
NP-11
MP-12
MP-23
1'1P-28
MP-33
1'1P-35
1'1P-36
MP-43
MP-47
MP-48
HP-49
MP-50
HP-54
Y1P-56
MP-58
MP-64
MP-65
MP-66
MP-76
HP-79
MP-80
MP-81
MP-82
MP-85
MP-86
MP-91
MP-92
MP-93
MP-94
MP-95
1'1P-102
HP-103
HP-108
MP-109
HP-110
HP-119
MP-124

Red no number
Red no number
Red no number
Red no number
Red no number
Red no number
Red no number
Red ;': 20
Red * 12
Red 7"\ 7
Red j" 15
Red 'k 8
Red "k 10
Red I': 4
Red ;': 11
Red * 5
Red -k 1
Red --}, 6
Red -'- 18
Red * 9
Red I': 32
Red -k 31
Red ~: 13
Red i': 14
Red "k 28
Red 7: 29
Red i': 30
Red
* 16
Red * 27
Red ..,': 21
Red -k 3
Red * 1
Red -Ie 26
Red "k 23
Red * 24
Red -k 33
Red ~'e 35
Red ofe 40
Red ~'c 34
Red 'Ok 36
Red "k 37

'Sex

F

?

1'1 1'1ature
Mature
F
1'1 Mature
Mature
F
Mature
F
Mature
F
F ,Mature
1'1ature
F
Mature
F
Mature
F
1'1 Fawn
1'1ature
F
1'1 Yr1g.
1'1 Yrlg.
Mature
F
Mature
F
1'1 F'awn
1'1 Yrlg.
1'1ature
F
Fawn
F
Fawn
F
Yr1g.
1'1
Mature
F
Mature
F
1'1 F'awn
1'1 Yrlg.
Fawn
F
Mature
F
1'1 F&lt;1\"n

Remarks

- - ., - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

�- 146 'Table 2. Deer tagged and marked in the Muddy sub-unit of Middle Park,
Arapahoe National Forest, 1969 (continued).

Age

Tag No.

Neck Band Color
and Symbol

M
2-11-69
2-11-69
F
M
2-12-69
M
2-12-69
Unk
2-13-69
2-13-69
F
2-13-69
F
F
2-15- 69
F
2-16-69
2-10-69
F
F
2-10-69
F
2-11-69
2-17-69
F
F
2-17-69
F
2-18-69
2-19-69
M
2-19-69
F
F
2-19-69
F
2-19- 69
F
2-21-69
2-20-69
F
2-20-69
F
F
2-20-69
2-20-69
M
F
2-20-69
M
2-27-69
F
2-22-69
2.-22-69 F
F
2-22-69
2-23-69
F
F
2-23-69
2-23-69
M
F
2-23-69
F
2-24-69
M
2-24-69
F
2-25-69
F
2-25-69
F
3-2-69
3-2-69
F
3-2-69
F
3-2-69
F
3-3-69
F
3-3-69
F

Eawn
Fawn
Mature
Fawn
Fawn
Mature
Mature
F a\17Il
Fawn
Fa\l7Il
Mature
Mature
Mature
Fa\l7Il
Fa\l7Il
Fa\l7Il
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fa\l7Il
Mature
Yr1g.
Fa\l7Il
Fa\l7Il
Mature
Mature
Fa\l7Il
Fa\l7Il
Fawn
Mature
Yr1g.
Mature
Yrlg.
Mature
Mature
Mature
Fml7Il
Fawn
Mature

MP-126
MP-127
MP-135
MP-136
MP-137
MP-138
MP-139
MP-148
MP-149
MP-153
MP-154
MP-159
HP-161
MP-162
MP-165
MP-176
MP-l77
MP-178
HP-179
MP-181
MP-183
MP-184
MP-185
MP-186
MP-187
MP-196
MP-201
MP-202
MP-203
MP- 209
MP-210
MP- 211
MP-212
MP-213
MP- 214
MP-217
MP-218
MP-227
MP-228
MP-229
MP- 230 .
MP-232
MP-233

Red-{( 27
no neckband
Red no number
no neckband
no neckband
Red -{(42
Red * 45
no neckband
no neckband
Red
* 38
Red ok 39
Red 1: 41
Red ~'(44
no neckband
no neckband
no neckband
Red * 46
Red * 41
no neckband
Red
* 57
Red 1: 48
Red * 49
Red * 50
no neckband
Red no number
Red 1: 29
no neckband
no neckband
Red * 54
Red * 53
no neckband
no neckband
no neckband
Red ..,'c 68
Red 1( 22
Red * 51
Red * 52
Red )'(61
Red * 56
Red 1( 58
no neckband
no neck band
Red * 60

----

-------------------------------.

Date
Tagged

Sex

Remarks

�- 147 'Table 2. Deer tagged and marked in the Muddy sub-unit of Middle Park,
Arapahoe National Forest, 1969 (continued).

Date
Tagged

Sex

Age

Tag No.

Neck Band Color
and Symbol

3-8-69
3-8-69
3-5-69
3-5-69
3-5-69
3-7-69
3-7-69
3-7-69
3-8-69
3-12-69
3-12-69
3-10-69
3-10- 69
3-10-69
3-11-69
3-11-69
3-11-69
3-11-69

Fawn
Mature
Mature
F
Unk Fawn
F .1'1ature
Mature
F
Yrlg.
F
Mature
F
Fm-m
F
Mature
M
Fm-m
M
17
1'1ature
1'1 Fawn
1'1 Fawn
Fawn
F
Fawn
F
Fawn
1'1
Mature
F

MP-239
MP-240
MP-242
MP-243
MP-244
MP-247
MP- 248
MP- 249
MP-251
MP-252
MP-253
1'1P-260
MP-26l
MP-262
MP-265
MP-266
MP-267
MP-270

no neckband
Red -{(64
Red ok 63
no neckband
Red ,'(25
Red ok 55
Red 'Ok 67
Red ,',72
no neckband
no neckband
no neckband
Red &gt;'( 69
no neckband
no neckband
no neckband
no neckband
no neckband
Red * 66

*

F
F

Indicates diamond ~

symbol on neckband

Remarks

�- 148 Table 3. Deer tagged and marked in the Blue River sub-unit of Middle
Park, Arapahoe Na t LonaI Forest, 1969.

Date
Tagged

1- 20-69

1-20-69 .
1-20-69
1-20-69
2-4-69
2-4-69
2-6-69
2-7-69
2-7-69
2-7-69
2-7-69
2-8-69
2-8-69
2-8-69
2-11-69
2-11-69
2-11-69
2-l3-69
2-13-69
2-14-69
2-15-69
2-15-69
2-9-69
2-9-69
2-10-69
2-18-69
2-18-69
2-21-69
2-20-69
2-21-69
2-21-69
2-25-69
2-28-69
2-28-69
3-1-69
3-6-69

*

Sex

Age

Tag No.

F
F
M
M
11
F

Fawn
Mature
Mature
Mature
Yrlg.
Mature
Mature
Fawn
Fawn
l1ature
Fawn
Mature

MP-18
MP-19
MP-20
MP-22
MP-87
MP-88
l1P-l07
MP-115
MP-116
MP-117
MP-118
MP-12l
MP-122
MP-123
MP-l3l
MP-132
MP-133
MP-141

M

F
M
F
F
F
F
F
F
F
M
F
1'1

M
M
M
F
M
M
N

M
M

M
F
M
N
F
M

F
F

Indicate

Fmm

Mature
Nature
Mature
Fawn
l1ature
Hature
Yrlg.
Fawn
Mature
Ma.ture
Mature
Mature
Mature
Yrlg.
Mature
Eawn
Mature
Nature
Fawn
Mature
Fawn
Nature
Mature

}1P~142

MP-144
MP-146
MP-147
MP-151
MP-152
MP-158
MP-170
MP-l71
MP-180
MP-189
MP- 205
MP- 206
MP-219
MP- 221
MP-222
MP-226
MP-237

Neck Band Color
and Symbol

Blue no number
Blue no number
Blue no number
Blue no number
Blue
7
Blue ,'(4
Blue * 9
Blue 1~ 5
no neckband
Blue ~k 2
Blue
* 3
Blue i', 19
Blue "k 8
Blue "k 17
Blue 'k 9
Blue ;': 11
no neckband
Blue * 12
Blue -}:8
Blue * 20
no neckband
Blue ;'~ 10
Blue * 16
Blue no number
Blue no number
Blue no number
Blue no number
Blue 'k 21
no neckband
Blue -}(26
Blue * 22
no neckband
Blue ,'(30
no neckband
Blue * 13
Blue '/(14
71t~

d i amond .• symbol on neckband.

Remarks

�- 149 Table 4. Deer tagged and marked in the Williams Fork sub-unit of
Middle Park, Arapahoe National Forest, 1969.

Date
Tagged

Sex

M
F
1-16- 69
F
1-20-69
F
1-24-69
F
1-25-69
1-26-69 Unk
F
1-25-69
F
1-25-69
M
1-30-69
F
2-5-69
F
2-7-69
F
2-10-69
F
2-17- 69
F
2-21-69
M
2-25-69
F
3-7-69
F
3-9-69
F
3-9-69
?

*

Age

Tag No.

Fawn
Mature
Mature
Fawn
Yrlg.
Unknown
Fawn
Fawn
Fawn
Mature
Mature
Hature
Fawn
Hature
Yrlg.
Fawn
Mature
Fawn

MP-3
MP-7
MP-25
MP-29
HP-32
MP-39
MP-40
HP-4l
MP-62
HP-101
HP-114
MP-157
MP-164
MP-204
MP-220
MP-238
MP-258
MP-259

Indicates diamond ~

Neck Band Color
and Symbol

Yellow no number
Yellow no number
Yellow no number
Yellow ~'r 4
yellow -k 1
Yellow -/~2
Yellow * 6
Yellow -/,7
Yellow ~',8
Ye LLow -;, 19
Yellow 7~ 9
Yellow * 20
no neckband
Yellow 7, 31
?

no neckband
Yellow * 33
no neckband

symbol on neckband.

Remarks

�- 150 Table 5. Deer tagged and marked in the Troublesome
Middle Park, Arapahoe National Forest, 1969.

Datc
Tagged

Sex

1-16-69
1-18-69
1-20-69
1-23-69
1-23-69
1-25-69
1-25-69
1-26-69
1-26-69
1-27-69
1-27-69
1-22-69
1-22-69
1-24-69
1- 28-69
1-28-69
1-28-69
1-31-69
2-1-69
2-1-69
2-1-69
2-1- 69
2-1-69
2-2-69
2-3-69
2-5-69
2-5-69
2-5-69
2-5-69
2-6-69
2-6-69
2-7-69
2-7-69
2-8-69
2-9-69
2-11- 69
2-11-69
2-11-69
2-13-69
2-15-69
2-16-69
2-10-69

Yr1g.
MP-6
Hature
MP-13
1'1atllre MP-24
F
1'1ature MP-26
F
1'1atllre MP-27
F
F
1'1ature 1'1P-30
Hature
M
1'1P-31
Fawn
F
1'1P-37
Fawn
M
HP-38
1'1 Fawn
MP-4S
F
Fawn
MP-46
Fawn
M
MP-S2
Mature
F
MP-S3
Fawn
F
MP-57
F
Fawn
MP-S9
1'1ature MP-60
F
Fawn
M
MP-61
Mature
F
MP-67
Fawn
F
MP-69
Mature
1'1P-70
F
Mature
F
1'1P-71
Fawn
F
MP-72
Unknown MP-73
F
Mature
F
MP-78
·F
Mature
MP-84
Yrlg.
M
MP-89
Yrlg.
MP-96
M
Fawn
M
MP-97
Mature
M
MP-98
Fawn
MP-104
F
Mature
}fP-10S
F
Yr1g.
M
MP-112
Mature
M
HP-113
Mature
M
MP-120
Mature
M
MP-12S
Yr1g.
M
MP-128
Mature
M
MP-129
Mature
F
MP-130
Mature
F
MP-140
Fawn
MP-145
M
;'vi
Fawn
MP-150
Mature
F
MP-1S5

- - - -

M

F

Age

Tag No.

Neck Band Color
and Symbol

sub-unit of

Remarks

White no number
White no numl&gt;er
White no number
White no numl&gt;er
White no number
White -k 4
White
* 10
White 7( 9
White 7( 5
White "k 12
White ,~ 2
l-lli ite ,'(11
White
* 1
White * 3
White ~~ 18
White ,'\ 17
White
* 13
v;rhite 15
* 14
White
* 20
White
*
? 19
?
no neckband
White
* 6
White no number
White * 8
White no number
White
* 6
White i~ 21
White ,'(2·2
White
* 26
White no number
White
* 30
White -,'c 24
White ~~ 23
White no number
White 1r 45
Hhite 7\ 48
White -,'r 46
White -l( 20
no neckband
no neckband
White
* 25

- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

�- 151 Table 5. Deer tagged and marked in the Troublesome sub-unit of Middle
Park, Arapahoe National Forest, 1969 (continued).

Date
Tagged

Sex

Age

2-10-69
F
Fawn
2-16-69
M
Fawn
2-17-69
F
Mature
2-18-69
M
Fawn
2-18-69
F
Mature
2-18-69
M
Mature
2-19-69
F
Fawn
2-21-69
F
Mature
2-21-69
M
Yr1g.
3-4-69
F
Fawn
2-20-69
F
Fa,VIl
2-21-69
F
Fawn
2-25-69
F
Mature
2-25-69
F
Fawn
2-25-69
F
Mature
2-25-69
M
Fawn
2-26-69
F
Mature
2-28-69
F
Mature
3-6-69
F
Mature
2-23-69
F
Mature
2-24-69
F
Fawn
2-24-69
F
Mature
3-1::69 ,':co, ""'F"i""Ma
ture
3-1":'69 F
Mature
3-1-69'
M
Mature
3-26':'69 F
Fawn
,3-3-69
M
Yr1g.
3-4-69
F
Fawn
3-4-69
M
Mature
3-7-69
F
Mature
3-12-69
M
Mature
3-8-69
M
Fawn
3-9-69
M
Fawn
3-9-69
F
Fawn
3-9-69
F
Mature
3-10-69
?
?
3-10-69
M
Fm·m
3-11-69
F
Yrlg.
3-11-69
M
Fawn
3-11-69
M
Mature
3-12-69
M
Mature

Tag No.

Neck Band Color
and Symbol

Remarks

no neckband
MP-156
no neckband
MP-160
White * 38
MP-163
no neckband
MP-166
White no number
MP-167
White no number
MP-168
no neckband
HP-l73
White 'k 47
MP-174
White * 20
MP-175
White * 36
l1P-182
no neckband
MP-188
no neckband.
MP-190
White
* 53
MP-191
no
neckband
MP-192
White ~'(54
MP-193
no neckband
MP-194
White
~':'59
MP-195
1-11'-198 \'vTIitc * 32
White .;:37
MP-200
White * 27
MP- 207
no neckband
l1P-215
White * 52
MP-216
Wh'it'e'"
*,. 51,::·''',,:
MP- 223
Whitei: 40'"
MP- 224
White * 49
MP-225
no neckband
MP-231
White no number
MP-234
no neckband
MP- 235
White no number
MP- 236
White ~'(56
MP-250
Whitei: 11
MP-254
no neckband
MP-255
no neckband
MP-256
no neckband
MP-257
White
* 31
no ta:;s
White
-Ie 34
HP-263
no
neckband
MP-264
no neckband
HP- 268
no neckband
MP-269
no neckband
MP-271
no neckband
MP-272

--------------

- - - - - - - - -

�- 152 Table 5. Deer tagged and marked in the Troublesome sub-unit of Middle
Park, Arapahoe National Forest: 1969 (continued).

Date
Tagged

3-12- 69
3-12-69
3-13-69
3-15-69
3-15-69
3-15-69
3-15-69
3- 26-69
3- 26-69
3- 26-69
3-26-69
3- 27-69
3-28-69
3-28-69
3-29-69
3-30-69
3-30-69

*

Sex

Age

Tag No.

Neck Band Color
and Symbol

F
M
M
M
M

Mature
Mature
Fawn
Yrlg.
Mature

MP-273
MP-274
MP-275
MP-276
MP-277
MP- 278
MP- 279
HP-280
MP-28l
MP- 282
MP-283
MP-284
MP-285
MP-286
MP-288
MP- 289
HP-290

no neckband
no neckband
no neckband
no neckband
White .,&lt;c 42
White * 7
v,,1l1ite43
*
White * 28
no neckband
no neckband
White "/(16
no neckband
no neckband
no neckband
no neckband
White * 41
no neckband

F

Fmm

M
F
F
M
F
F
M
F
M
F
F

Mature
Mature
Fawn
Mature
Hature
Fawn
Fawn
Hature
Fawn
Mature
Fawn

Indicates diamond.

symbol on neckband.

Remarks

�- 153 -

Table 6. Deer tagged and marked in the Beaver Creek sub-unit of
Middle Park, Arapahoe National Forest, 1969.

Date
T3gged

1-14-69
1-17-69
1-18-69
1-19-69
1-19-69
1-25-69
1-25-69
1-21-69
1-22-69
1-30-69
2-3-69
? -69
2-2-69
2-3-69
2-5-69
2-5-69
2-6-69
2-14-69
2-18-69
2-28-69
2-23-69

*

Age

Tag No.

Neck Band color
and Symbol

Mature
Fawn
Fawn
Yr1g.
Mature
Mature
Mature
Yr1g.
Mature
Mature
Fawn
Unk
Mature
Yr1g.
Mature
Fmro
Fawn
Fawn
Hature
J:v1 Mature
Yr1g.
1'1

MP-1
MP-10
MP·-14
MP-16
MP-17
HP-42
J:vlP-44
MP-51
MP-55
MP-63
MP-68
MP-74
MP-77
MP-83
MP-99
MP-100
MP-106
MP-143
MP-169
MP-199
MP-208

no neckband
Pink no number
Pink no number
Pink no number
Pink no number
Pink ~~ 12
Pink * 16
Pink * 5
no neckband
Pink * 19
Pink #': 13
Pink 'k 10
Pink ~~ 24
Pink * 15
Pink 'f( 25
no neckband
Pink no number
no neckband
Pink no number
Pink no number
no neckband

Sex

1'1
M
1'1
1'1
1'1
N.
1'1
1'1
1'1
1'1
1'1
1'1
1'1
1'1
1'1
1'1
1'1
1'1
1'1

Indicates diamond ~

symbol on neckband

Remarks

�- 154 -

Table 7. Deer tagged and marked in the Muddy sub-unit of Middle Park,
Arapahoe National Forest, 1970.

Date
Tagged

Sex

1-17-70
F
1-20-70 . F
17
1-31-70
F
2-1-70
2-2-70
M
2-6-70
M
F
2-7-70
F
2-15-70
2-22-70
M
F
2-26-70
M
3-4-70
F
3-7-70
F
3-9-70

*

Age

Tag No.

l'iD-ture MP-333
Nature
MP-342
l'iD-ture MP-352
Mature
MP-353
Yrlg.
MP-355
Fawn
MP-368
MD-ture MP-376
}1ature MP-397
MP-418
Fawn
MP-430
Mature
FaHn
MP-441
Mature
MP-447
Fawn
MP-453

Indicates circle

@

Neck Band Color
and Symbol
Red ole 14
Red "1: 2
Red ;': 12
Red -1. 6
Red -;e 5
7
Red )';;
Red "k 58
Red * 4
11
Red
Red "l: 17
Red .,': 1
Red "k 16
no neckband
~'(

symbol on neckband ,

Remarks

�- 155 Table 8. Deer tagged and marked in the Blue River sub-unit of Middle
Park, Arapahoe National Forest, 1970.

Date
Tagged

1-8-70
1-8-70
1-9-70
1-9-70
1-10-70
1-10-70
1-10-70
1-10-70
1-10-70
1-10-70
1-11-70
1-11-70
1-11-70
1-11-70
1-11-70
1-12-70
1-12-70
1-12-70
1-12-70
·1-13-70
1-14-70
1-14-70
1-15-70
1-15-70
1-15-70
1-15-70
1-16-70
1-16-70
1-17-70
1-18-70
1-19-70
1-19-70
1-21-70
1-25-70
1-26-70
2-5-70
2-14-70
2-17-70
2-22-70
2-23-70
2-25-70
2-25-70
2-25-70

--- -

Sex

Age

Tag No.

Neck Band Color
and Symbol

F
M
M
M
H

Mature
Yrlg.
Yrlg.
Mature
Mature
Hature
Fawn
Fawn
Mature
Yrlg.
Yr1g.
Mature
Yrlg.
Fawn
Fawn
Yrlg.
Fawn
Mature
Mature
Mature
Fawn
Mature
Ma·ture
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Mature
Yrlg.
Yrlg.
Mature
Hature
Hature
Mature
Mature
Mature
Mature
Mature
Hature
Hature
Mature

MP-29l
MP-292
MP-293
MP-294
MP-295
MP-296
MP-297
MP-298
MP-299
MP-300
HP-301
MP-302
MP-303
MP-304
MP-305
MP-307
MP-308
MP-309
MP-310
MP-312
MP-313
MP-314
MP-317
MP-3l8
MP-319
MP-320
MP-324
MP-325
MP-332
MP-336
MP-337
MP-338
MP-343
MP-350
MP-35l
HP-3G2
MP-396
MP-404
MP-417
MP-423~
MP-427
MP-428
MP-429

Blue * 14
Blue -/,9
Blue * 3
Blue "/e 1
Blue -/e 5
Blue ofe 10
Blue ofe 11
Blue ofe 3
Blue "/e 2
Blue no number
Blue
* 6
Blue * 4
Blue
7
*
Blue * 15
Blue ,'c 12
Blue * 18
Blue * 3.0
Blue * 24
Blue * 27
no neckband
Blue ok 16
Blue
* 28
Blue
* 29
Blue * 28
Blue * 8
Blue ,'c 31
Blue * 34
Blue * 41
Blue -/, 40
Blue ,'e 20
Blue * 23
Blue ~: 39
Blue ,'c 52
Blue * 50
Blue ;'~ 51
Blue ..,,':: 53
Blue -1\ 45
Blue * 49
Blue * 59
Blue '1, 19
no ncckband
no neckband
no neckband

F

F
M
F
M
F
M
F
F
M
M
M
F
F

M
M
F
F
M
M
F
M
M
F
M
M
M
M
M
M
F
F
F
F
M
F
F

F

- - - - - - - -

Remarks

---- - -- --- - - - - - - - - - -

�- 156 -

Table 8. Deer tagged and marked in the Blue River sub-unit of Middle Park,
Arapahoe National Forest, 1970 (continued).

Deer
Tagged

Sex

Age

Tag No.

Neck Band Color
and Symbol

2-26-70
2-26-70
2-28-70
3-4-70
3-4-70
3-13-70
3-19-70
3-19-70
3-21-70
3-22-70
3- 23-70
3- 26-70
3-26-70

F
M
F
F
F
F
M
F
F
F
M
F
F

Mature
MP-432
F'awn
MP-433
Mature
MP-437
Mature
MP-439
MP-440-50
Fawn
Fawn
MP-468
Mature
MP-479
MP-480
Fawn
Mature
MP-489
Fawn
MP-492
}iature MP-495
Fawn
MP-500
Fawn
MP-502

Blue ~~ 57
Blue ~'~44
Blue ~'c 60
Blue ~~47
Blue * 46
Blue ~~ 26
Blue * 22
Blue
48
Blue 1~ 37
no neckband
Blue no number
no neckband
no neckband

*

Indicates circle.

*

symbol on neckband.

Remarks

�- 157 Table 9. Deer tagged and marked in the Williams Fork sub-unit of Middle
Park, Arapahoe National Forest, 1970.

Date
Tagged

Sex

1-13-70
1-14-70
1-14-70
1-15-70
1-15-70
1-16-70
1-16-70
1-16-70
1-17-70
1-17-70
1-17-70
1-18-70
1-18-70
1-20-70
1-20-70
1-21-70
1-22-70
1-23-70
1-24-70
2-6-70
2-5-70
2-7-70
2-7-70
2-7-70
2-9-70
2-14-70
2-14-70
2-15-70
2-20-70
2-22-70
2-22-70
2-24-70
2-27-70
3-6-70
3-7-70
3-9-70
3-10-70
3-14-70
3-17-70
3- 20-70
3-21-70
3-23-70
3-26-70

MP-311
Mature
M
MP-315
Nature
M
MP-3l6
1'1 Yr1g.
HP-321
Fawn
F
MP-323
1'1 Nature
MP-326
Eawn
F
MP-327
Yr1g.
M
MP- 328
Mature
M
MP-329
Mature
F
MP-330_
Fawn
M
HP-33l
Mature
M
MP-334
Mature
F
MP-335
Mature
F
MP-339
Mature
M
MP-340
1'1ature
M
MP-344
Mature
M
MP-346
Fawn
F
MP-348
Mature
M
MP-349
Fmvu
H
Fawn
MP-357
F
MP-36l
Mature
M
MP-373
Mature
F
MP-374
Fawn
F
l1ature Y1P-375
F
MP-380
Fawn
F
MP-392
Fawn
F
HP-393
Mature
F
MP-398
Eawn
H
HP-411
Hature
H
MP-419
Fawn
M
MP-420
Hature
M
MP-426
Mature
F
MP-434
Mature
F
MP-444
Hature
F
Fa\,"I1 MP-448
F
MP-454
Mature
F
HP-458
Fawn
F
MP-469
Fawn
M
MP-472
Fawn
M
MP-487
Fmvu
F
MP-490
Fnwn
F
MP-496
Mature
M
no tags
Mature
M

'1c

Age

Indicntcs circle •

Tag No.

Neck Band Color
and Symbol

Yellow ~'(5
Yellow 'k 9
Yellow "/&lt; 11
Yellow ~~ 16
Ye l.Low ~'&lt; 13
Yellow 'k 4
Yellow "/&lt; 1
Yellow i': 14
Yellow "/c 2
Yellow * 3
Yellow * 35
Yellow * 6
Yellow "/,14
Yellow * 17'
Yellow ~'&lt; 19
Yellow "/,7
Yellow * 8
Yellow * 18
Yellow * 26
Yellow * 24
Ye LLow * 21
Yel10\\I* 34
Ye'LLow ~~ 12
Yellow * 31
Yellow * 28
Yellow * 46
Yellow * 44
Yellow "/,23
Yellow * '29
Yellow * 22
Yellow * 33
Yellow ~'c 39
Yellow * 45
Yellow ~'c 10
Yellow 7( 47
Yellow * 53
no neckband
no neckband
no neckband
no neckband
no ncckband
no neckband
Yellow -;,20

symbol on ncckb and ,

Remarks

�- 158 Table 10. Deer tagged and.marked in.the Troublesome sub-unit of Middle
Park, Arapahoe National Forest, 1970.

Date
Tagged

1-22-70
1-~3-70
2-4-70
2-4-70
2-4-70
2-5-70
2-5-70
2-6-70
2-6-70
2-6-70
2-6-70
2-6·-70
2-7-70
2-7-70
2-7-70
2-7-70
2-9-70
2-9-70
2-9-70
2-10-70
2-10-70
2-11-70
2-11-70
2-11-70
2-11-70
2-12-70
2-12-7.0
2-13-70
2-13-70
2-13-70
2-14-70
2-14-70
2-15-70
2-15-70
2-16-70
2-17-70
2-17-70
2-18-70
2-19-70
2-19-70

-- - --

Sex

Age

Tag No.

Neck Band Color
and Symbol

F
F
F
H
F
M

Mature
Mature
Mature
Mature
Fawn
Fawn
Fa,VIl
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Nature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn

MP-345
MP-347
HP-354
MP-356
HP-358
MP-359
MP-360
MP-363
MP-364
MP-365
MP-366
MP-367
MP-369
MP-370
MP-371
MP-372
MP-377
MP-378
MP-379
MP-381
HP-382
MP-383
MP-384
MP-385
MP-386
MP-387
MP-388
MP-389
MP-390
MP-391
MP-394
MJ?-395
MP-399
MP-400
MP-40l
MP-402
MP-403
MP-405
MP-407
MP-408

White ~~ 3
White ok 4
l'lhi
te ~',5
White * 2
White -k 30
White 'k 1
no neckband
White -k 12
White ~': 20
White
* 14
White ,;', 13
v.Thite;': 9
White ..,'( 15
White ok 23
White
19
White ..,'( 7
White ;'( 10
White ~. 11
White ..,r: 18
White
* 21
White "k 25
White
* 816
White
* 22
White
* 6
White
*
White ok 39
White of, 17
White of, 29
White
* 33
White ~',26
White of, 42
White
* 34
White -k 30
White * 24
White
* 36
White * 37
White
* 28
White -k 43
1rJhite"I, 52
no neckband

M

F
M
M
M
M
F
F
M
M
M
F

M
F
M
M
F
F
F
F

M
M
M
F
M
M
M
F
F
F

M
F
F

M

------

Remarks

"'!(

-----------------------

�- 159 Table 10. Deer tagged and marked in the Troublesome sub-unit of Middle
Park, Arapahoe National Forest, 1970 (continued).

Date
Tagged

Sex

2-19-70
2-19-70
2-20-70
2-21-70
2-21-70
2-21-70
2-22-70
2-22-70
2-23-70
2-23-70
2-26-70
2-28-70
2-28-70
3-2-70
3-4-70
3-4-70
3-5-70
3-6";70
3-6-70
.3-7-70
3-7-70
3-8-70
3-10-70
3-11-70
3-12-70
3-12-70
3-13-70
3-13-7.0
3-13-70
3-13-70
3-14-70
3-16-70
3-16-70
3-16-70
3-17-70
3-17-70
3-17-70
3-17-70
3-19-70
3-18-70
3-19-70
3-20-70

M
M
M
F
M
M
M
M
F
M
M
F
F
F
F
M
F
F
F
F
F
F
F
F
F
F
F
F
F
F
F
F
M
M
F
M
F
F
F
M
F
F

--- -

Age

Tag No.

Neck Band Color
and Symbol

Remarks

White ,~31
MP-409
Mature
White "/,40
.Fawn
MP-410
White * 42
Mature
MP-412
White
HP-414
Mature
* 49
no neckband
MP-415
Fawn
no neckband
MP-416
Fawn
White
"/,45
MP-421
Mature
no
neckband
MP-422
Mature
J:.1P-42If White ,~ 38
Mature
J:.1P-425 no neckband
Hature
53
White
MP-431
Mature
*
White
MP-435
Mature
* 57
MP-436
White * 35
Fawn
White
MP-438
Mature
* 58
no neckband
MP-440-50
Fawn
White ,,:56
MP-442
Mature
no neckband
MP-443
Fawn
White
MP-445
Mature
* 46
White * 47
t-1ature HP-446
MP-449
White * 27
Fawn
no neckband
MP-451
Fawn
MP-452
White * 51
Mature
White
MP-457
Mature
* 44
White
MP-459
Mature
* 60
White * 50
MP-460
Mature
no neckband
MP-461
Fawn
no neckband
MP-462
Fawn
White * 67
Mature
MP-463
no neckband
MP-464
Fawn
White * 68
MP-465
Mature
no neckband
MP-466
Fawn
White
* 59
Mature
MP-467
no
neckband
HP-470
Mature
White * 54
MP-471
Mature
White * 61
MP-473
Mature
no neckband
MP-474
Mature
White
"/,70
MP-475
Mature
no neckband
MP-476
Fawn
no neckband
MP-477
Eawn
no neckband
MP-478
Fawn
no neckband
MP-481
Fawn
no neckband
Mature
MP-482

- - - - - - - -

- - - - - - - - - - - - - - - - - - - - - -

�- 160 -

Table 10. Deer tagged and marked in the Troublesome sub-unit of Middle
Park, Arapahoe National Forest, 1970 (continued).

Date
Tagged

Sex

Age

Tag No.

Neck Band Color
and Symbol

3-19-70
3-19-70
3-19-70
3-19-70
3- 21-70
3-21-70
3-22-70
3-26-70
3-23-70
3-23-70
3-24-70
3-27-70

M
M
F
M
F
M
F
M
H
F
M
H

Fawn
Fawn
Fawn
Hature
Fawn
Hature
Mature
Mature
Mature
Fawn
Mature
Hature

MP-483
MP-484
MP-485
MP-486
HP-488
MP-491
MP-493
MP-494
MP-497
MP-498
MP-499
MP-501

no neckband
no ne ckb and
no neckband
no ne ckb and
no neckband
no neckband
White ~.(63
no neckband
no neckband
no neckband
no neckband
no neckband

*

Indicates circle •

symbol on neckband.

Remarks

�- 161 -

Table 11. Deer tagged and marked in the Muddy sub-unit of Middle Park, Arapahoe
National Forest, 1971.
Neck Band Color
and Symbol

Date
Tagged

Sex

Age

1-11-71
1-12-71
1-19-71
1-24-71
1-27-71
1-27-71
1-28-71
1-29-71
1-29-71
1-29-71
2-3-71
2-13-71
2-l3-71
2-l3-71
2-13-71
2-l3-71
2-14-71
2-14-71
2-15-71
2-16-71
2-17-71
2-21-71
2-22-71
2-25-71
2-25-71
2-25-71
2-26-71
2-26-71
2-26-71
2-27-71
3-1-71
3-3-71
3-3-71
3-4-71
3-4-71
3-7-71
3-9-71
3-9-71
3-11-71
3-12-71
3-12-71
3-16-71
3-16-71
3-16-71
3-17-71
3-17-71
1-15-71
2-22-71
2-25-71

F
F
M
F
F
F
M
M
F
M
M
F
F
F
F
F
F
F
F
F
F
F
F
F
F
F
F
M
F
M
M·
F
F
M
F
F
F
F
F
F
F
F
M
F
F
F
F
F
F

Yr1g. MP 515 Red no number
Mature MP 520 Red no number
Mature MP 540 Red X 9
MatureMP 550 Red X 1
MP 552 Red X 11
Fawn
Mature MP 553 Red Xl3
MP 554 Red X 15
Fawn
MP 557 Red X 7
Fawn
Mature MP 558 Red X 14
Mature MP 559 Red X 8
Removed for res. at Ft.Co11in~, died 2-28-71
Mature MP 564 ·Red X 18
Mature MP 588 Red X- 22
MP 589 Red X 5
Fawn
MP 590 Red X2
Fawn
Mature MP 591 Red X 21
Mature MP 592 Red .X 23
Mature MP 595 Red X 12
Mature MP 596 Red X 31
Mature MP 601 Red X 35
Mature MP 602 Red X 34
MP 603 Red X 32
Fawn
MP 609 Red X 36
Fawn
Mature MP 613 Red X 29
Mature MP 619 Red X 37
MP 620 Red X 30
Fawn
Mature MP 621 Red X 38
Fawn .MP 625 Red X 39
MP 626 Red X 27
Fawn
Trap mortality 2-26-71
Mature None
None
Mature MP 632 Red X 24
MP 637 Red X 25
Fawn
MP 644 Red X 46
Fawn
Mature MP 645 Red X 44
Mature MP 646 Red X 16
Mature MP 647 Red X 48
Mature MP 659 Red X 43
Mature MP 661 Red X 41
Mature MP 662 Red X 40
MP 671 Red X 47
Fawn
Mature MP 673 Red X 50
Mature MP 674 Red X 42
Mature MP 681 Red X 45
Trap mortality 3-16-71
None
None
Fawn
Trap mortality 3-16-71
None
None
Mature
Trap mortality 3-17-71
None
None
Mature
Trap mortality 3-17-51
Mature None
None
mortality 1-15-71
Trap
None
None
Fawn
mortality 2-22-71
Trap
None
None
Hature
Trap mortality 2-25-71
None
None
Fawn

Tag No -.

Remarks

�- 162 .Table 12. Deer tagged and marked in the Blue River sub-unit of Middle Park,
Arapahoe National Forest, 1971.

Date
Tagged

1-12-71
1-15-71
1-16-71
1-16-71
1-16-71
1-16-71
1-16-71
1-21-71
1-24-71
1-29-71
1-29-71
2-7-71
2-8-71
2-10-71
2-11-71
2-14-71
2-15-71
2-15-71
2-17-71
2-17-71
2-18-71
2-21-71
2-23-71
2-24-71
2-25-71
2-25-71
2-26-71
2-26-71
2-27-71
2-27-71
3-1-71
3-2-71
3-3-71
3-6-71
3-7-71
3-7-71
3-8-71
3-9-71
3-9-71
3-9-71
3-10-71
3-17-71
3-20-71
3-20-71
3-23-71
3-25-71
3-25-71

Sex

Age

Tag No.

Neck Band Color
and SYmbol

M
F
M
M
M
M
M
M
M
M
M
F

Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Hature
Mature
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Hature
Mature

MP 521
MP 528
MP 529
MP 530
MP 533
MP 534
MP 535
MP 543
MP 549
MP 560
MP 561
MP 573
MP 574
MP 580
MP 581
MP 597
MP 599
MP 600
MP 604
MP 605
MP 606
MP 610
MP 614
MP 616
MP 622
MP 623
MP 627
MP 628
MP 633
MP 634
MP 638
MP 640
MP 643
MP 652
MP 657
MP 658
MP 660
MP 663
MP 664
MP 665
MP 669
MP 684
MP 688
MP 689
MI' 691
HP 695
MP 696

Blue no number
Blue nO number
No neckband
No neckband
Blue no number
.No neckband
Blue no number
Blue X 15
Blue X 8
Blue X 14
Blue X 9
Blue X 2
Blue X 3
Blue X 12
Blue X 11
Blue X 17
Blue it 28
Blue X 21
Blue X 16
Blue X 18
Blue X 25
Blue X 24
Blue X 23
Blue :I. 22
Blue X. 4
Blue X 26
Blue X 34
Blue X 05
Blue X 36
Blue X. 19
Blue X 35
Blue X 31
Blue X 30
Blue X 39
Blue X 37
Blue X 40
Blue X 53
Blue X. 52
Blue X 33
Blue Z 29
Blue X 50
Blue X 51
Blue X 58
Blue X 42
Blue X 1+3
Blue X 56
Blue X 45

F

F
F
M
F
M

M
M
M
M
M
M
F
F
M
F
F
F
M
M
F
F
F
F
M
M
M
M
F
M
M
F
F
M
F

Remarks

�- 163 Table 13. Deer tagged and marked in the Williams Fork sub-unit of Middle Park,
Aripahoe National Forest, 1971.

Date
Tagged

Sex

Age ....Tag

Mature
Mature
Mature
Fawn
Mature
Mature
Mature

Yellow no number
Yellow X 1
Yellow no number
Yellow no number
Yellow! 2
Yellow no number
No neckband
No neckband
Yellow no number
Yellow no number
Yellow ~. 3
Yellow no number
Yellow no number
Yellow no number
Yellow no number
Yellow X 4
Yellow X 5
Yellow no number

Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn

MP539
MP 541
MP 544
MP 545
MP 548
MP 551
MP 555
MP 563
MP 565
MP 566
MP 567
MP 568
MP 582
MP 583
MP 584
MP 607
MP 611
MP 612
MP 615
MP 617
MP 670
MP 676
None
None

Yellow Z. 6
Yellow X. 9
yellow X 8
Yellow X 11
Ye1l6wX 14
Yellow! 21 .
Yellow X 16
Yellow X 23
Yellow X 27
Yellow X 26
Yellow X 28
Yellow X 15
Yellow X 19
Yellow X 29
Yellow X 20
Yellow X 24
Yellow X 30
Yellow X 33
Yellow X 40
Yellow X 17
Yellow X 42
Yellow X 47
None
None

Mature
Mature
Mature
Fawn
Mature
Mature
Fawn.
Mature
Mature
Fawn

F
F

2-§.'C71

2-1'1-71
2-12-71
2-12-71
2-20-71
2-21-71
2-21-71
2-23-71
2-24-71
3-10-71
3-12-71
1-16-71
1-25-71

F

M
M
M
F
M
M
M
M
M
F
M
F
M
M
F
F
F
F
M
F
F

Neck Band Color
and Symbo 1

MP 505
MP 506
MP 507
MP 508
MP 509
MP 510
MP 511
MP 512
MP 516
MP 517
MP 518
MP 519
MP 523
MP 524
MP 525
MP 526
MP 527
MP 532

1-9-71
M
1-9-71
F
1-'9-71
M
1-10-71
M
1..,10-71 F
1-10 ....
71
M
.1':"10-71 M
1:";'10-71 M
1~1l-71
M
1-11-71
M
1-12-711' F
1-l2-71
M
1-15-71
M
1-:15-71 M
1-15-71
M
1-15-71
F
1-15-71
F
1-16-71
F
1-19-71
1-20-71
1-21-71
1-22-71
1-24-71
1-26-71
1-28-71
2....
2-71
2-4-71
2-6-71
2-6-71

No.

Fawn

Remarks

Removed for research at Fort Collins,
Died 3-9-71.

Trap mortality 1-16-71
Trap mortality 1-25-71

�ill

- 164 -

•Table 14 .. Deer tagged and marked in the Troublesome sub-unit of Middle Park,
Arapahoe: National Forest, 1971.
Date
Tagged

1-17-71
1-20-71
1-28-71
2-6-7J.
2-7-71
2-8-71
2-10-71
2-10-71
2-12-71
2-12-71
2-14-71
2-24-71
2-27-71
2-27-71
2-27-71
3-1-71
3-1-71
3-2-71
3-3-71
3-3-71
3-4-71
3-4-71
3-4-71
3-8-71
3-7-71
3-7-71
3-9-71
3-9-71
3-10-71
3-14-71
3-15-71
3-15-71
3-16-71
3-18-71
3-21-71
3-23-71

Age

M
M
M

M
F

M
M
F

M
F
F
F
F

F
F
F
F

F
F

M
F
F

F
F
M
M

M
F
F
M
F
F
F
M
F

F

Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
M&lt;1ture
Fawn
Fawn
Fawn
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn

.Tage No.

MP 536

.MP 542
MP 556
MP 570
MP 572
MP 575
MP .577
MP578
MP585
MP 586
MP 598
MP 618
MP 629
MP 630
MP 631
MP 635
MP 636
MP 639
MP 642
None
MP 648
MP 649
NP 650

MP 651
MP 655
MP 656
MP 666
MP 667
MP 668
MP 678
MP 679
MP 680
MP 682
MP 685
MP 690
MP 692

Neck Band Color
a.ndSymbol

White X 6
White X 7
White X 10
White X 9
White X 4
White X 14
White X 24
White X 19
White X 22
White X 20
White X 01
White X 17
White X 26 .
White x 27
White X 29
White X 03
White X 02
White X 18
White X 28
None
l.JhiteX 16
White X 30
No neckband
White X 41
White X 23
White X 12
White X 13
White X 36
White X 37
White X 35
White X 44
White :x 26
White X 40
White X 15
White X 39
White X 42

Remarks

Died 2-15-71

Trap mortality 3-3-71

�165 -

Park,
T'able15 . Dee'r taggeJ and marked in the Beaver Creek sub-unit of Midd Le
Arapahoe National Forest, 1971.

Date
Tagged

1-1""71
1-15-71
1-16-71
1-17-71
1-17-71
1-24-71
1-31-71
2-6-71
2-9-71
2-10-71
2-11-71
2 -14-71
2-14-71
2-25-71
3-3-71
3-6-71
3-7-71
3-l3-71

Sex

Age

Tag No.

Neck Band Color
and Symbol

M
F
M
M
F
M
M
M
F
M
F
M

Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Mature

MP 513
MP 522
MP 531
MP 537
MP 538
MP 546
MP 562
MP 569
MP 576
MP 579
MP 587
MP 593
MP 594
MP 624
MP 641
None
MP 654
MP 677

No neckband
Pink no number
Pink no number
No neckband
No neckband
No neckband
Pink no number
Pink ! 16
Pink X. 2
No neckband
Pink X 10
No neckband
No neckband
Pink:X: 13
Pink X. 05
None
Pink Z.
Pink X 07

M

F
F
M
M
F

Remarks

Trap mortality 2-22-71

Trap mortality 3-6-71

�- 166 -

Table 16. Location of trap areas, trap number and automatic data
processing code number.
Sub'Unit

1
1
1
1
1
1

1
1

1
2
2
2

2
2
2
2
2

3
3
3
3
3
5

4
4
4
4
4
4
4
1
1

Trap
Location

Legal
Description

Trap
Location No.

Wolford
S28 T2N R80W
Martin Ranch
S23 T2N RS1W
S, E. Martin Ranch
S25 T2N RSIW
Redl1tn., south
S10 TIN RSOW
Red Mtn., north
SlO TIN RSOW
City Reservoir
S11 TIN RS1W
Wo1.ford, upper
S28 T2N RSOW
·Wo1ford, lower
~33 T2N RSOW
Wolford, west
828 T2N R80W
Williams Peak Road
S35 TIS R80W
Harshy Gulch .
S15 TIS R80W
Shane Gulch
S27T2S R79W
Spring' Cr.eekRoad
S34 TIS RSOW
Cottonwood Gulch
s17 T28 R79W
Horse Gulch'
S7 T2S R79W
King Bulch'
S23 TIS R80W
Junction Butte
S16 TIN RSOW
Cedar Ridge, west
S16 TIN R79W
Cedar Ridge, north . Sl3 TIN R79W
Ce dar Ridge, sec. IS S18 TIN R79W
Cedar Ridge, dam
road
S14 TIN R79W
Cedar Ridge, east
boat ramp
S24T1N'R79W
Beaver
S15 TINR7SW
Rifle Range
S16 TIN R7SW
Parshall Divide
S17 TIN R78W
Corral Creek, lower Sll TIN R79W
Corral Creek, upper S2 TIN R79W
Sulphur Gulch,
lower
S8 TIN R79W
Sulphur Gulch,
upper
S29 T2N R79W
Scholl Ranch
S4 TIN R79W
BLM Corrals
S4 TIN RSOW
Kremmling Dump
S9 TIN RSOW

10
11
12
13
14
15
16
17
18
19

28210S0
2321081
252l0S1
·1021080
1021080
1121081
2821080
3321080
2821080
3522080
1512080
2722079
3412080
1722079
0722079
2312080
16110S0
1611079
1311079
1811079

21

1411079

22

23
24
25
26
32

2411079
1511078
1611078
1711078
1111079
0211079

27

0811079

33

2921079
0411079
0411080
0911080

1
2

3
4
5

6
7
8

29
9

28
30
31

Sub-Unit

ADP Code

Muddy
Blue
Williams Fork
Troublesome
Beaver

1
2

3
4
5

ADP
Code

�- 167 -

LITERATURE

CITED

Gill, R. B. 1969. Middle Park deer study - population distribution.
Colo. Div. Game, Fish and Parks.
Game Res. Div. Federal Aid
Proj. W-38-R-23.
Game Res. Rept., July, Part 1, pp. 79-104.
Gill, R. B. 1970. Middle Park deer study - population distribution.
Colo. Div. Game, Fish and Parks.
Game Res. Div. Federal Aid
Proj. W-38-R-24.
Game Res. Rept., July, Part 3, pp. 287-309.

Prepared

by

JiM 0.4--

Laren A. Roper
Asst. Wildlife Researcher

��- 169 July,

JOB PROGRESS

State of

1971

REPORT

COLORADO
--------~~~~------------

Project No.
Work Plan No.
Job Title

W-38-R-25

Investigations
2

Job No.

14
Middle

Deer-Elk

Park Deer Study - Population
10, 1970 through March

Density

and Structure

Period Covered:

October

7, 1971

Personnel:

R.B. Gill, L.A. Roper, G.L. Brown, L.H. Carpenter,
H.R. Shepherd, R.C. Kufeld, W.T. McKean, D. Baker
A.E. And~rson, C.E. Braun, J.F. Corey and B.D. Baker.

ABSTRACT

The estimate of the 1971 Middle Park winter deer population was 5,730 deer.
This represented a 20.5 percent decline from 1970 and a 46.1 percent decline
from the 1968 high of 10,640 deer. Preseason classifications
indicated
ratios of 42 bucks:100 does and 72 fawns:100 does in the Middle Park deer
population~
Postseason ratios were estimated to be 46 bucks:100 does and
41 fawns:100 does. Examinations of the hunter harvest revealed that the
1969 and 1970 kill contained a higher proportion of does and fawns and a
,lower proportion of bucks than did the 1967 and 1968 harvests.

��~ 171 -

MIDDLE PARK DEER STUDY POPULATION DENSITY AND STRUCTURE
R. Bruce Gill

P. S. OBJECTIVE
To estimate the density and sex and age structure of the Middle Park deer
population in order to harvest this population more efficiently.

SEGMENT OBJECTIVES
1.

Estimate

the size of the total winter

2.

Estimate

the sex and age structure

METHODS

population

of the Middle

of deer in Middle

Park.

Park deer population.

AND MATERIALS

See Gill (1969) for a detailed presentation of methods and materials.
Something which has not been discussed in previous reports, concerns the ground
rules that we established before the first counts were conducted in 1968.
We decided that the perimeters of each section would be flown first so that
decisions could be made as to whether peripheral animals were to be included
or excluded from the quadrat tally. We decided that to be included in the
tally a deer must be within the boundaries of the quadrat when first observed.
Every other deer seen along the perimeter was excluded even though fresh
tracks might reveal that the animal had just exited from the quadrat.
Unless
this rule is strictly adhered to, the tendency is to count every animal near
the quadrat boundary, thus inflating density estimates.
After flying the perimeter, the section is flown in strips, determined by
topographical features of the quadrat.
For example, if the quadrat is bisected by three draws, each draw and adjacent ridge is covered during one
strip.
The width of each strip and the number of strips per section are
determined by the amount of area that can be observed on each strip, and this
depends upon variations of terrain within each quadrat.
Quadrats have been numbered in the order in which they are flown each year.
This counting order was set up to maximize counting efficiency and thus
minimize cost. The counting sequence by sub-unit is depicted in Figs. 1 and
2.

�- 172 -

Fig. 1.
River,

N

Quajrat count sequences in the Muddy Creek. Blue River, Williams Fork
and Troublesome

Creek Sub-units.

�- 173 N

t

Fig. 2.

Quadrat count sequence in the Granby Sub-unit.

�~ 174 -

RESULTS AND DISCUSSION

Density

Timing of the Count
The 1971 Middle Park deer census was conducted much later in the winter
than previous years.
This year the count was not conducted until March
6 and 7, 1971, while in previous years it was conducted in January.
There
were two reasons for this late count in 1971:
(1) the past winter was
exceptionally mild and even though some snow fell in January we did not
believe it was sufficient to force deer out of the lower timber zones and
into the count areas.
(2) snow background was not adequate to insure
accurate tallies of deer until early March.
Prior to this time the detection of deer was difficult.
The effect of the late count period on the density estimates was judged
to be negligible because deer distribution was still widespread during
March, 1971. However, the entire question of the effects of deer concentrations upon density estimates projected from fixed quadrat locations and
based on assumed widespread distribution of deer is still unresolved.
This
question is scheduled for investigation beginning the winter of 1972-73.

Density

Estimates

The 1971 quadrat counts indicated a mean density of 9.75 ± 3.02 deer per
square mile over the 587.7 square miles of winter range. This is projected
to a total population estimate of 5,730 ± 1,775 deer. Comparing the 1971
estimates with the projected means from previous years, the 1971 deer population has declined 20.5 percent from 1970 and 46.1 percent from the 1968
peak populations of 10,640 deer (Table 1).
An examination of the projected estimates by sub-units over the four-year
count period reveals that apparent drops in the Muddy Creek and Troublesome
Creek sub-units have contributed most to the overall 1971 decline (Table 2).
We speculate the reasons for this decline lie in harvests exceeding net
fawn crops in 1969 and 1970.

Validating

the Estimates

Generally, in making estimates of free-ranging wild animal populations, one
is never sure if estimates are correct.
One way to validate the estimate is
to consider all the major birth and death factors affecting the population
in question and calculate the expected population level before conducting
population census.
This expected population level can then be compared with
the census estimate, and if the two are compatible over a series of years,
confidence in the census estimate is bolstered.

�- 175 -

Table 2.

Population

Year

Muddy
Creek

Biue
River

1968

4,101

4,074

843

1,622

1969

2,994

3,531

774

1,679

134

9,112

1970

1,429

3,290

271

1,910

306

7,206

1971

794

3,335 .

571

891

139

5,730

estimates

by sub-unit

Sub-Units
Williams Fork
River

in Middle Park, 1968-71.

Troublesome
Creek

Granby

Totals

not counted

10,640

We tried this approach with the Middle Park deer population estimates beginning with the 1968 census year as a base.
The formula used to calculate
expected population levels was: P2 = PI + b - ~ - mw where P = expected
n
2
population level; PI = actual population estimate from the previous year;
bn = net productivity derived from pre-season classification counts; mh =
harvest mortality; and ffiw = winter mortality.
Thus, for 1971 the expected population level is: P2 = 7,206 + 2,560 - 2,801 1,135 or P2 = 5,830.
The actual quadrat census estimate was 5,730 for a
difference of 1.7 percent.
Similar close agreement was evident for past
years (Table 3), where the maximum difference was 8.4 percent.

Table 3. Comparisons of expected population
census estimates in Middle Park, 1969-1971.

levels with actual quadrat

Year

Expected
Population

1969

9,519

9,112

4.3%

1970

7,862

7,206

8.3%

1971

5,830

5,730

1. 7%

Census
Estimate

Percent
Difference

�Table 1.

Numbers of deer counted per quadrat within the eight sampling strata, Middle Park, Colorado, 1968-1971.

1971

N.C.
0
N.C.
0
N.C.
0
N.C.
N.C.

6
0
0
0
0
0
0
0
0
0
15
0

0
1
0
30
0
0
0
0
0
0
11
0

0
0
0
0
0
0
0
0
0
0
0
0

14
25
0
76
31
4
9
134
49
71
124
28

19
0
1
36
88
20
4
102
15
105
58
17

14
0
0
24
77
18
3
20
83
136
49
6

145
54
7
0
34
43
67
104
0

1971

0
0
0
0
0
9
6
18
99
39
66
46
87
31
6
12
0
0
22
37

0
0
0
0
0
9
34
9
20
6
2
2
42
28
5
12
0
0
5
1

0
12
4
0
0
0
0
0
28
68
2
0
22
0
0
0
0
0
0
0

56
N.C.
0

2
0
0
0
0
8
64
32
124
58
30
32
*N.C.
44
N.C.
20
27
0
1
N.C.

Blue River
High Density
1970
1969
1968

Muddy Creek
Low Density
1970
1969
1968

Muddy Creek
High Density
1970
1969
1968

0

Blue River
Low Density
1970
1969

1971

1971

1968

47
0

4
10
0
30
32
0
0
31
0

8
10
0
0
35
0
0
36
29

0
17
0
37
34
0
0
26
0

0
13
0

0

0

15
0
0
20
0

--

t =

442

478

175

136

56

21

42

0

565

465

430

501

107

118

114

48

Y=

26.0

23.9

8.8

6.8

9.3

1.8

3.5

0.0

47.1

38.8

35.8

41.8

11.9

13.1

12.7

5.3

--------------------------------------------------------------------------------------------------------------------N.C. = No Count.

,.....

-....I

0'\

�Table 1. Numbers of deer counted per quadrat within the eight sampling sf-rata, Middle Park, Colorado, 1968-1971
(continued).
Williams Fork River
High Density
1968
1969
1970

0
2
26
73
4

i=

40
0
45
42
4

0
0
25
21
0

1971

3
0
58
31
0

Williams Fork River
Low Density
1968
1969
1970
1971
0
6
14
0
0

0
0
3
0
0

0
0
1
0
0

0
4
0
0
0

Troublesome Creek
High Density
1968
1969
1970

Granby
1971

43
25
0
35
50
65
0
9
0
26

5
41
0
39
76
44
15
36
0
6

11
60
0
51
42
90
7
8
0
29

0
8
0
0
43
36
14
21
10
7

105

131

46

92

20

3

1

4

253

262

298

139

y = 21.0

26.2

9.2

18.4

4.0

0.6

0.2

0.8

25.3

26.2

29.8

13.9

1968

1969

1970

1971

=

20.75

15.52

12.29

9.75

v(yst)=

11.35

4.25

2.55

3.29

yst

1968

z

0
r-t

CJ
0

c::l

r-t
(1)

0-

N.C.

1969

1970

1971

7
2
2
0
0
0
N.C.

6
23
0
0
0
0
0

0
0
13
0
0
0
0

11

29

13

1.8

4.1

1.9

•.....

-...J
-...J

�- 178 -

Sex and Age Composition

Field Sex and Age Composition

Counts

Pre-season Counts--Pre-season
sex and age classifications were obtained
from a helicopter during the period October 10-12, 1970. Count conditions
were excellent since the counts immediately followed a fresh and abnormally
heavy October snowfall which concentrated deer on upper winter ranges in
the sagebrush-aspen
interfaces.
Seven hundred seventy-one deer were
classified, yielding a buck:doe ratio of 42:100 and a fawn:doe ratio of
72:100 (Table 4). The buck:doe ratio was very similar to that of 1969,
but the fawn:doe ratio was approximately 23 percent lower. The drop in
the fawn:doe ratio was attributed to lower fawn production and survival
compared with 1969 (Table 5).

of 1969 and 1970 pre-season

Table 4. Comparisons
Middle Park.

in

deer classifications

Fawn:Doe
Ratio

Buck:Doe
Ratio

Year

Bucks

Does

Fawns

Total

1969

195
(18.3%)

451
(42.4%)

419
(39.3%)

1,065
(100.0%)

43

100

93

100

1970

152
(19.7%)

361
(46.8%)

258
(33.5%)

771
(100.0%)

42

100

72

100

Table 5. Comparisons
and 1970.

of gross and net fawn production

in Middle Park, 1969

Net**
Fawn
Production

Year

Gross*
Fawn
Production

1969

200

100

125

100

1970

167

100

93

100

* Gross fawn productlon
. = fetuses per 100 does based upon collections

of

32 does each year.

** Net fawn production
surveys.

= fawns:lOO mothers based upon aerial classification

�- 179 -

Post-season Counts--No post-season aerial sex and age classifications
were attempted in 1971 because these are usually made coincidentally
with the quadrat census survey, and in 1971 the quadrat census was
not conducted until March when bucks were well into antler shedding.
The only sex and age data available were obtained by ground counts
from snowmobiles and from afoot.
Results from previous years indicated close agreement between aerial classifications and ground
classifications
(Table 6). The trend in annual fawn:doe ratios has
been steadily downward since 1968 (Fig. 3). Possible factors influencing
this decline are discussed in the Job Progress Report for WP 14, Job 3.

Table 6. Comparison of aerial and ground winter
data for Middle Park, 1967-68 through 1970-71.

count classification

Years

Aerial Counts
Buck : Doe
Fawn : Doe
Ratios
Ratios

1967-68

56

100

90

100

37

100

93

100

1968-69

53

100

85

100

54

100

87

100

1969-70

45

100

77

100

48

100

82

100

46

100

41

100

1970-71

Sex and Age Composition

No Counts

Ground Counts
Buck : Doe
Fawn : Doe
Ratios
Ratios

of the Hunter Harvest

Sex and Age Composition of the Season Harvest--Checks were made of the 1970
Middle Park legal deer harvest at two stationary check stations (Idaho
Springs and Dillon) and by one checker circulating through Middle Park
checking deer at hunter camps.
Check stations were operated for nine days
and proved far superior to the roving checker in terms of time efficiency
per deer checked.
The roving checker examined 40 deer in the nine day
period while six men manning the two stationary check stations examined
1,012 deer. This amounted to 4.4 deer checked per man-day of effort using
a roving checker vs. 18. 7 deer checked per man-day of effort using
stationary check stations.
The sex and age composition of the 1970 harvest is presented in various
formats in Tables 7-10. Comparisons of the 1970 harvest data with those
from 1967, 1968, and 1969 reveals an increase in the doe harvest over the
past two years and a decrease in the buck harvest while the fawn harvest
has remained essentially the same (Table 11).

�100

90

80
(fJ
Q)

70

0

~

0
0

r-I

E-&lt;
(fJ

§
C\l

~

'H

0
(fJ
0

60

S
0

u

0

00

0

\

50

§
~

I-'

\

40

'M
-I.J
C\l

p::j

30

20

10

0
1967-68

Fig. 3.

Trend

in post-season

1968-69

1969-70

fawn:doe ratios obtained

1970-71

from ground count surveys,

1967-68

- 1970-71.

��Table 8.

Sex and age classes of deer checked from Middle Park in 1970, excluding headless deer.

GM
Unit

Adult
Males

Yearling
Males

Male
Fawns

Adult
Females

Yearling
Females

Female
Fawns

Totals

15

9
18. 7%

8
16.7%

6
12.5%

13
27.1%

12
25.0%

0
0.0%

48
100.0%

18

60
15.4%

100
25.7%

28
7.2%

106
27.3%

60
15.4%

35
9.0%

389
100.0%

27

45
17.6%

60
23.5%

29
11.3%

70
27.3%

27
10.5%

25
9.8%

256
100.0%

28

20
15.2%

38
28.8%

13
9.8%

31
23.5%

21
15.9%

9
6.8%

132
100.0%

37

27
21.6%

32
25.6%

21
16.8%

18
14.4%

16
12.8%

11
8.8%

125
100.0%

Totals

161
16.9%

238
25.1%

97
10.2%

238
25.1%

136
14.3%

80
8.4%

950
100.0%

I-'
co
N

�Table 9. Number and percent of deer checked from Middle Park in 1970 in each age category by sex classes,
excluding headless deer.

GM
Unit

Adult
Males

Yearling
Males

Male
Fawns

Totals

Adult
Females

Yearling
Females

Female
Fawns

Totals

15

9
39.1%

8
34.8%

6
26.1%

23
100.0%

13
52.0%

12
48.0%

0
0.0%

25
100.0%

18

60
31.9%

100
53.2%

28
14.9%

188
100.0%

106
52.7%

60
29.9%

35
17.4%

201
100.0% .

27

45
33.6%

60
44.8%

29
21.6%

134
100.0%

70
57.4%

27
22.1%

25
20.5%

122
100.0%

28

20
28.2%

38
53.5%

13
18.3%

71
100.0%

31
50.8%

21
34.4%

9
14.8%

61
100.0%

37

27
33.8%

32
40.0%

21
26.2%

80
100.0%

18
40.0%

16
35.6%

11
24.4%

45
100.0%

TOTALS

161
32.4%

238
48.0%

97
19.6%

496
100.0%

238
52.4%

136
30.0%

80
17.6%

454
100.0%

.
t-'

00
LA)

�- 184 -

Table 10'. Number and percent adults and yearlings in the Middle Park deer
check, 1970.
GM
Unit

Adult
Males

Yearling
Males

Total
Males

Adult
Females

Yearling
Females

Total
Females

15

9
52.9%

8
47.1%

17
100.0%

13
52.0%

12
48.0%

25
100.0%

18

60
37.5%

100
62.5%

160
100.0%

106
63.9%

60
36.1%

166
100.0%

27

45
42.9%

60
57.1%

105
100.0%

70
72.2%

27
27.8%

97
100.0%

28

20
34.5%

38
65.5%

58
100.0%

31
59.6%

21
40.4%

52
100.0%

37

27
45.8%

32
54.2%

59
100.0%

18
52.9%

16
47.1%

34
100.0%

161
40.4%

238
59.6%

399
100.0%

238
63.6%

136
36.4%

374
100.0%

Totals

Table II. Comparison of sex and age structure of the Middle Park deer harvest,
1967-1970.
Year

Males

Females

Fawns

Totals

1967

375
46.1%

260
31.9%

179
22.0%

814
100.0%

1968

342
47.6%

235
32.7%

141
19.7%

718
100.0%

1969

605
40.9%

583
39.4%

291
19.7%

1,479
100.0%

1970

399
42.0%

374
39.4%

177
18.6%

950
100.0%

Table 12 presents a summary of the year specific age composition of the 1970
deer harvest in comparison with the three previous years.

�Table 12.

Year

Comparisons of the year specific age composition of the Middle Park deer harvest, 1967-1970.

Fawns

1 Yr.

2 Yr s ,

3 Yrs.

Numbers and Percent in Each Age Class
4 Yrs. 5 Yrs.
6 Yrs.
7 Yrs.
8 Yrs.

9 Yrs.

10 Yrs.

11+ Yrs.

Total

Males
L.O

8.6%

18
3.9%

12
2.6%

7
1.5%

5
1.1%

1
0.2%

3
0.7%

1
0.2%

463
100.0%

52
12.3%

24
5.4%

14
3.3%

8
1.9%

3
0.7%

4
1.0%

3
0.7%

1
0.2%

2
0.5%

424
100.0%

159
21.2%

92
12.3%

38
5.1%

13
1. 7%

11
1.5%

9
1.2%

5
0.7%

4
0.5%

0
0.0%

2
0.3%

748
100.0%

86
17.6%

32
6.5%

12
2.5%

9

1. 8%

2
0.4%

1
0.2%

6
1.2%

3
0.6%

3
0.6%

1
0.2%

490
100.0%

99
21.4%

187
40.4%

45
9.7%

45
9.7%

1968

82
19.3%

181
42.7%

51
12.0%

1969

160
21.4%

255
34.1%

1970

97
19.8%

238
48.6%

1967

I

I-'

00
V1

Females
1967

80
22.9%

84
24.0%

39
11.1%

45
12.9%

39
11.1%

22
6.3%

16
4.6%

7
2.0%

4
1.1%

3
0.9%

4
1.1%

7
2.0%

350
100.0%

1968

59
21.1%

58
20.7%

40
14.3%

60
21.4%

20
7.1%

8
2.9%

8
2.9%

8
2.9%

5
1. 8%

7
2.5%

3
1.0%

4
1. 4%

280
100.0%

1969

131
18.8%

142
20.3%

192
27.5%

93
13.3%

51
7.3%

20
2.9%

13
1.9%

14
2.9%

19
2.7%

8
1.1%

3
0.5%

12
1. 7%

698
100.0%

1970

80
18.1%

136
30.8%

128
29.0%

45
10.2%

17
3.9%

9
2.1%

7
1.6%

5
1.1%

2
0.5%

2
0.5%

5
1.1%

5
1.1%

441
100.0%

�2,-

- 18

Sex and Age Composition of the Daily Check--One occasionally hears the
criticism that check station sex and age composition data are biased
because they are only gathered during the first nine or ten days of the
hunting season instead of season-long.
These critics argue that a greater
proportion of does and fawns are taken home by hunters during the first
half of the season, whereas more bucks are taken home during the latter
half.
Examination of Figures 4 and 5 reveal no consistent trends in
the Middle Park deer check within any single year or over a period of
years.
We conclude that the nine day check station period provides an
accurate sample of the sex and age composition of the Middle Park deer
harvest.

LITERATURE

CITED

Gill, R. B. 1969. Middle Park deer study - population density and
structure.
Colo. Div. Game, Fish and Parks.
Game Res. Div.
Fed. Aid. Proj. W-38-R-23.
Game Res. Rep. July, Part 1. pp. 105-122.

Prepared

by __ ~~ •.......
~_"}:...:::J~_-\.='Y-""""-""~---'-\.4-' -"--~=~::.....:'
R. Bruce Gill
Wildlife Researcher

=-_

�- 187 --'---i"

I
I

II

i.60

1967

50
M
0
fr.l

tn

0

40
/
.
..1
;).~
- __
.- -'- ~-7'
- ~ - - - -.
--% DO;5 in Total

~

H
&lt;11

!::)

•

30

rx.
0
f-I

:z;
fr.l
0

~
0...

20

...•.

It- - .

.".

..•...
...
: '.

.....•..

-.

••••••••••••••••.•••••••••••.•••

...•....

:!"II ~

¢heck

F1Iwns in Total

Check

% Does in Total

Check

-70

•••••••••

••

10

'.'

· •••••••••.••••
'.
'.'

:

0
1

2

4

7

5
DAY OF CHECK

3

1968

60

50
:x::
0
l"&lt;,

::T~

u

1.•0

..•.

?~
1-4

~-.:
(.::)

.,..

, _ .•...
_ _

I~

...

,

/..........

_ -t -

30

n,

10

rc:;
f-tl

••• ~.~

•...

.•.

-

_..-

•.. : ."

• • • • •••••

•.

••••

---...--

..•..

.....

C~

20

.

'II'

tz.
E-&lt;
;.:.0:;
bJ
U

..,..-

•••••.••••

•.....

• ••

_ ..•.
-

')..,- - -

.•.....•
~ •••

-.

• ~ • "".:

..

'It

'

••

d
]0

Fawns in Total

0

1

2

3

4

5

6

7

8

checks

at big game check

9

DAY OF CHECK

Fig. 4.
Sex and age composition of daily hunter
stations, October 1967 and October 1968.

Check

�- 188 -

1969

60
50
:.:::

u

~

p::

u

40

~

H
H

«

0

30

~

0
E-4

:z:
u

20

~

P:!
~
p..

10
0

:3

2

1

4
5
DAY OF CHECK

6

8

7

9

1970

60
50
:.:::

u

~

p::

u

40

;-~~~~~--~~~--------------~----~-

---

~

H

«
0

30

~

0
E-4

:z:
~
u

ffi
n,

,~Bucks in Total Check

.

20

•• ::

...

.•

••••

D" ••••••••••

..... .
0

•

'

-).'~ Does- in Total

.

.'
G

.

•••••
... -..

~.~'
•••• '.:
.•••••••• -~ Fawns

..

•.

10

in Total Check

•••••••

0
1

2

3

4

5
DAY OF CHECK

6

Fig. 5. Sex and age composition of daily hunter
stations, ()ctober 1969 and October 1970.

Check

7

8

9

check at big game check

-

�- 189 July, 1971

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~---------

Project No.

W-38-R-25

Work Plan No.

14

Deer-Elk

Investigations
3

Job No.

Job Title

Middle Park Deer Study - Productivity

Period Covered:

October

PersonnE!l:

L.H. Carpenter, L.A. Roper, L.M. Brown, D. Baker, R. Hoffman, P.F.Gilbert,
G.L. Brown, W. Brandes, R.E. Keiss,
D.E. Benson and R.B. Gill.

10, 1970 through August

and Mortality

5, 1971

ABSTRACT
Estimates of prenatal productivity were derived from two sources:
counts
of corpora lutea and fetus counts.
The average number of corpora lutea of
pregnancy for 51 does examined was 1.73 C.L./doe.
The average number of
fetuses from the same sample was 1.61 fetuses/doe.
The sex ratio of the
1971 fetus sample did not differ significantly from those of 1969 and 1970,
but there were significantly more males than females in the combined three
year sample (68 percent males vs. 32 percent females).
Conception and parturition dates were calculated from estimated fetal ages. The majority of the
1971 fetuses were conceived during the period November 21 to December 5. The
majority of the parturition dates would have occurred during the period June
11-20. Estimates of postnatal fawn survival were calculated for both the
1969 and 1970 fawn crops. Approximately 51 percent of the 1969 fawn crop
was estimated to have survived from conception to midwinter (January 15) of
the following year, while 46 percent of the 1970 fawn crop survived over a
comparable period.
Projections of the 1970-71 winter mortality were derived
from dead deer surveys.
Estimates were that 489 ± 213 deer died. Highway,
railroad,and other known miscellaneous losses totaled 115 deer, bringing the
total number of deer estimated to have died during the 1970-71 winter to 604.
Estimates of the legal harvest for Middle Park were derived from two separate
sources:
a ratio method involving check station figures and Game Management
Unit harvest report card estimates; and a regression prediction based upon
the relationship between the Idaho Springs check station tally of Middle Park
deer and the Grand and Summit Counties report card deer estimates.
These
estimates were 2,097 and 2,436 deer, respectively.

��- 191 -

MIDDLE PARK DEER STUDY - PRODUCTIVITY

AND MORTALITY

R. Bruce Gill

P. S. OBJECTIVE
To estimate increments and losses to the Middle Park deer population
order to formulate more efficient harvest regulations.

in

SEGMENT OBJECTIVES
1.

Estimate

productivity

rates of Middle Park deer.

2.

Estimate
range.

mortality

3.

Estimate

the magnitude

rates of Middle Park deer over the entire winter

of the hunter harvest.

METHODS AND MATERIALS
See Gill (1969) for details.

RESULTS AND DISCUSSION

Productivity

Pre-natal

Productivity

Corpora Lutea Counts--During the period January-May, 1971, 51 sets of mule
deer ovaries were collected and macroscopically
examined for the presence
of corpora lutea of pregnancy.
Does were separated into two age classes,
yearlings and older aged does. The average number of C.L.P./doe for eleven
yearlings was 1.27 compared to 1.85 C.L.P./doe for older age animals.
The
average number of C.L.P./doe for all does combined was 1.73. These figures
are comparable to data from 1970 does, but are slightly lower than those
observed in 1969 (Table 1). Although these data have not been tested for
statistical significance, there appears to have been a decline in productivity
in 1970 and 1971 from 1969.
The proportion of the fertilized ova developing into viable embryos and
fetuses also appeared to decline in 1970 and 1971. In 1969 95.5 percent of
the fertilized ova developed into viable embryos and fetises as evodemced
from comparisons of C.L.P. and fetus counts.
In 1970 and 1971 the proportions
were 93.2 percent and 93.1 percent, respectively.
Thus, not only did initial
fertility appear to decline, but also survival from the blastocyst stage to
the embryonic and fetal stages of pre-natal development (Fig. 1).

�Table 1. Comparisons of numbers of corpora lutea of pregnancy and fetuses occurring in samples of mule deer reproductive
tracts collected from Middle Park, Colorado, 1969-1971.

Year

All Does
No. of
No.
of
No. of
Does
Fertilization
Corpora
Does
Fertilization
Corpora
Does
Fertilization
Corpora
Examined
Rate
Fetuses
Lutea
Examined
Rate
Fetuses
Lutea
Examined
Rate
Lutea Fetuses

1969

1.50

1.00

66.7%

2

2.05

1.97

96.1%

39

2.02

1.93

95.5'%

·41

1970

1.29

1.29

100.0%

7

1.86

1.71

91.9%

35

1.76

1.64

93.2%

42

1971

1.27

1.09

85.8%

11

1.85

1.75

94.6%

40

1.73

1.61

93.1%

51

Total

1.30

1.15

88.5%

20

1.92

1.82

94.8%

114

1.83

1.72

94.0%

134

Adult Does

Yearling Does

~

\0
N

�II

i

,

i

J.

i----i---

.

i

ADULT DOES
...

n ='39·

)

2

!

r •

o
1969

1970

1971

All Years

1970

1971

All Years

n = 41

2

1

o
1969

Fig. 1. Comparison of corpora lutea and fetal rates from female mule
deer collected from Middle Park, Colorado, 1969-71.

�- 194 -

Accessory Corpora Lutea--Various
authors have described the occurrence of
of luteinized ovarian structures histologically and morphologically
similar
to primary corpora lutea of pregnancy, only smaller in size (Robinette et
al. 1955; Morrison 1960; Simkin 1965; Douglas 1966; and Markgren 1969).
Similar structures were noted in the samples of Middle Park deer ovaries.
Robinette et al. (1955) suggested that these "accessory" corpora lutea
could be distinguished from C.L.P. by their smaller size and lack of a
rupture site.
In this current study ovaries were not examined microscopically so accessory corpora lutea were distinguished by size alone.
Two measurements were made of all the luteinized ovarian structures found
in the 1971 ovaries.
These were the maximum diameter and the minimum
diameter of the structure.
The maximum diameter was the greatest distance
across the largest section of the structure.
The minimum diameter was
taken perpendicular to and at a point midway along the axis of the maximum
diameter (Fig. 2). Those structures which were judged to be primary cor~
pora lutea of pregnancy generally exceeded 5x7 mm while the accessory
corpora lutea measured 5x7 mm or less (Fig. 2).
Of the 101 structures measured, 6.9 percent were classed as accessory corpora lutea of pregnancy.
Most of the accessory corpora lutea (71.4 percent)
occurred in the right ovary.

Multiple Ovulation--Brambell
(1948) lists three sources of error encountered
when counts of C.L.P. are used as indexes to productivity: (1) a follicle
may develop into a corpus luteum without liberating an ovum (accessory
corpus luteum); (2) a follicle may liberate more than a single ovum; and
(3) counting errors may occur.
Suspected examples of case 1 were discussed
in the previous section.
One example of case 2 was noted from specimen
69-3 collected January 17, 1969. This particular doe contained four healthy
embryos, each enclosed within a separate amnion, and one additional amnion
containing amniotic fluid, but not a fully developed embryo (Fig. 3).
Examination of this doe's ovaries revealed only three corpora lutea of
pregnancy and was interpreted evidence that multiple ovulation from a single
follicle had occurred.

Litter

Sizes and Fetal Sex Ratios

Litter Sizes--Comparisons
of the average litter sizes of does examined over
the three year collection period (1969-1971) indicated a decrease in the
proportion of does bearing triplets, and an increase in the percent of nonpregnant does in 1970 and 1971 from 1969 (Table 2). This provided further
evidence of a general drop in productivity for 1970 and 1971 from the 1969
fawn crop.

�Fig. 2.

Scatter diagram of sizes of luteinized ovarian structures

mule deer reproductive
May, 1971.

tracts collected

in Middle Park, Colorado,

from
January

-

�- 196 -

Fig. 3. Embryos in amniotic sacs recovered
in Middle Park, Colorado, January 17,-1969.

from mule deer doe number

69-3

�- 197 -

Table 2. Litter size frequencies
Park, Colorado, 1969-1971.

of mule deer does examined

from Middle

Litter Sizes
2
3

4

n

10
24.4

25
61.0

5
12.2

1
2.4

41
100.0

5

7

28

2

0

42

Percent

11.9

16. 7

66.7

4.7

0.0

100.0

1971

No.
Percent

1
2.0

19
37.2

30
58.8

1
2.0

0
0.0

51
100.0

Totals

No.

6

36

83

8

1

134

Percent

4.5

26.9

61.9

6.0

0.7

100.0

Year

1969

No.
Percent

1970

No.

0

1

0
0.0

Fetal Sex Ratios--Each year that samples of fetuses have been collected from
Middle Park male fetuses have dominated the sample (Fig. 4). Chi-square
tests failed to detect significant year differences so data for all three
years were pooled and tested against the hypothesis that male and female
fetuses occurred with equal frequency (i.e. the samples were drawn from a
population comprised of 50 percent male fetuses and 50 percent female
fetuses).
The probability that these samples were drawn from such a population w,as less than .005. Therefore it was concluded that a significantly
greater number of male fetuses were produced over the three year period.

Conception and Parturition Dates--Forehead-rump
lengths were measured on
all undamaged fetuses in the 1971 sample.
These measurements were then
converted to estimated ages using the growth curve presented in Hudson and
Browman (1959). Dates of conception for each fetus were calculated by
backdating from collection dates.
These data are presented in Fig. 5,
with comparable data for 1969 and 1970. The majority of fetuses (65-80
percent) examined over the three year period were conceived between November
21 and December 5 of each year.
Assuming a gestation period of 202 days (Robinette and Gashwiler 1950)
parturition dates were calculated from fetal ages by foredating from collection
dates of the 1971 sample.
Approximately 50 percent of the projected parturition
dates would have occurred within the period June 11-20 (Fig. 6). Estimated
dates of parturition for the entire sample ranged from June 4 to July 4.

�.•198 -

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.

n ~69

~
...

60

.40

30
20

10

o
1969

1970

1971

All Years

Fig. 4. Proportions of males and females in samples of mule deer fetuses
from Middle Park, Colorado, 1969-1971.

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o,

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11/21-25

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11/21-25

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11/26-30

11/26-30

12/ 1-5

12/ 1-5

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�- 200 -

60
1971i

-1

i

50
(/)'

::r::

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- ...

40

~
Hi
m:

.:

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••

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.••

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-- -- -- ''''-- .-t

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&lt;o

DA.TES OF--~ARTURITION

Fig. 6. Estimate
dates of parturition for Middle Park does based on
fetal ages and an assumed gestation period of 202 days.

�~ 201 -

Post-natal

Fawn Survival

Fawn survival rates were calculated from time of conception through midwinter (January 15) for the 1969 and 1970 fawn crops. Estimates of the
number of fawns conceived per 100 does of producing age (2+ years) were
derived from counts of corpora lutea of pregnancy.
Estimates of the
number of fawns born were derived from fetus counts (Table 1). Estimates
of fawns surviving to hunting season were based on pre-season classification
counts, and estimates of fawn survival to January 15 were based on postseason classification counts (Table 3). Both the pre- and post-season
classification data were corrected so does that had not conceived their first
fawn were excluded from calculations of young:lOO does.

Table 3. Figures used in c~mputing numbers of fawns per 100 does preand post-season, corrected bOy excluding yearling does.

No. of
Does
Counted

Percent
Yearlings
in Doe
Population

Corrected
No. of
Does

No. of
Fawns
Counted

1969

451

24.4

341

419

123

100

1970

368

36.4

237

258

109

100

1969-70

471

24.4

356

363

102

100

1970-71

777

36.4

494

317

64

100

Ratio of
Fawns : 100
Producing Does

Pre-season
Classifications

Post-season
Classifications

Survival rates for the 1969 fawn crop for the 59 week period from conception
to mid-winter of the following year was approximately 51 percent, while the
survival rate for the 1970 fawn crop was 46 percent over a comparable period
(Fig. 7). Thus, not only were there fewer fawns produced in 1970, but a
smaller percentage survived, resulting in a double depressant on ensuing
population levels.

�- 202 -

Fig. 7.

Fawn survival rates for Middle Park mule deer, 1969-1970.

�- 203 -

Mortality

Winter Mortality
The projected estimate for 1971 deer winter mortality amounted to 489
deer, or 8.5 percent of the estimated post-season population of 5,730
deer. Ninety percent confidence levels about this estimate indicated
the true population winter loss lay somewhere between 274 and 704 deer.
Comparisons of 1971 losses with those of previous years are presented
in Fig. 8. The estimated winter loss figure was projected from only
15 carcasses encountered on 93 transects.
The sex and age composition
of 13 of the 15 deer which could be classified was: bucks - 38.4 percent; does - 30.8 percent; and fawns - 30.8 percent.
These data are
too limited to hazard extrapolation to total population losses by sex
and age classes.
The pooled data for the past four years may provide
a better estimate of the sex and age specific mortality (Table 4).
Fawns comprised the greatest proportion of the combined four-year
sample (45.1 percent), followed by does (29.2 percent), and lastly,
by bucks (25.7 percent).

Table 4. Numbers of dead deer by sex and age classes encountered
winter mortality survey, 1968-1971.
Survey
Year

1968

1969

Bucks

Does

Fawns

Total

No.

14

11

26

51

Percent

27.4

21. 6

51.0

100.0

6

5

8

19

31. 6

26.3

42.1

100.0

4

13

13

30

13.4

43.3

43.3

100.0

5

4

4

13

Percent

38.4

30.8

30.8

100.0

No.

29

33

51

113

Percent

25.7

29.2

45.1

100.0

No.
Percent

1970

No.
Percent

1971

Totals

in

No.

�- 204 -

3000
2800
2600
2400
2200

. ~---

2000
1800

&amp;3
~

1600

p..
0
(I)

e5
~
z

1400

TTfj

1200
1000
800
600
400
200
0
1967-68

1968-69

Fig. 8. Middle Park deer winter mortality estimates,
percent confidence limits, 1967-68 to 1970-71.

1969-70

bracketed

1970-71

by the 90

�- 205 -

Highway, Railroad, and Miscellaneous Mortality
During the period July 1, 1970 and June 30, 1971, WCO's in Middle Park
reported 57 deer deaths attributable to highway accidents. Searches
of the railroad rights-of-way intersecting major deer winter concentration sites yielded nine deer presumed to have been killed in deertrain collisions (Table 5).

Table 5.

Known highway and railroad losses, Middle Park, 1963-70.

Year

Highway
Losses

Railroad
Losses

1963-64

90

1964-65

385

1965-66

7

Totals

1966-67

56

l32

188

1967-68

54

60

114

1968-69

77

56

133

1969-70

67

38

105

1970-71

57

9

66

In addition to losses previously discussed, 33 deer were collected for reproduction studies and 16 were known to have perished from injuries sustained during trapping operations. Combining all aforementioned losses
with the dead deer survey estimate, 604 deer were estimated to have died
from all identified sources in the 1970-71 winter.
Harvest Mortality
In the past, three separate estimates of the Middle Park deer harvest have
been available. These were: (1) county report card estimates for Grand
and Summit counties; (2) a ratio estimate projected from the check and
estimated harvest from Game Management Units 18, 28, and 37; and (3) a
regression formula prediction based upon the relationship between the
Idaho Springs check of Middle Park deer and the Grand and Summit county
report card estimates (Gill 1970). Only two of these estimates were
available for the 1970 harvest, since the county of kill was not requested

�- 209 July,

JOB PROGRESS

REPORT

State of

....;C:...:O:..::L:...:o~RAD==...;;.O
_

Project No.

W-38-R-25

Work Plan No.

14

Job Title

Middle

1971

Deer-Elk

Investigations

4

Job No.

Park Deer Study - Food Habits and Physical

Period Covered:

April

1, 1970 through March

Personnel:

R. Bruce Gill, Willard Travnicek,
Paul Gilbert and Laren A. Roper.

Characteristics

31, 1971
Robert

E. Keiss,

Joe Griess,

ABSTRACT
Data on kidney fat and bone marrow fat percentages in mule deer in Middle
Park are presented.
Regression lines for kidney fat are linear for 1969
and 1970 and show a downward trend from January 15 - April of each year.
The 1971 kidney fat percentages show a similar linear trend for the FebruaryApril period, however, unusually low values in January results in a polynomial regression.
To date, no biological significance of this phenomena is
apparent.
Blood counts of collected deer are reported for 1971. An analysis
of variance for yearling buck antlers collected in 1968-1970 suggest no
significant difference exists between years for the antler weights.
The number of points occur in similar proportions each of these years.

�- 211 -

MIDDLE PARK DEER STUDY FOOD HABITS AND PHYSICAL CHARACTERISTICS
Laren A. Roper

P. S. OBJECTIVE
To measure selected physical characteristics related to the assessment of
physical condition of deer and to determine forage preferences of mule deer
in Middle Park.

SEGMENT OBJECTIVES
1.
2.
3.

Estimate forage preferences of deer from stomach content analyses.
Estimate forage preferences of deer from tame deer observations.
Measure selected physical characteristics of deer believed indicative
of the state of physical condition.

METHODS

Forage Preferences
Procedures

- Stomach Content Analyses

are reported by Gill (1969).

Forage Preferences
Procedures

AND MATERIALS

- Tame, Trained

Deer Observations

are reported by Gill (1969).

Physical

Characteristics

Procedures are reported by Gill (1969). The procedures reported by Roper
(1970) for blood samples have been changed.
During the 1971 collection period
blood smears taken from heart blood as soon after death as possible
were used.
Standard procedures for obtaining counts of segmented neutrophils, eosinophils, basophils, lymphocytes and monocytes were used. All
other procedures are reported by Roper (1970).

RESULTS AND DISCUSSION
Forage Preferences

- Stomach Content Analyses

Samples from 84 stomachs are now being analyzed
ported in the next segment.

and results will be re-

�- 212 -

Forage Preferences

- Tame, Trained Deer Observations

These data will be reported

in conjunction

with the Job 5 report.

This report includes data that may be indicative of the physical condition of deer collected from January 15, through May 21, 1969, 1970
and 1971. Data on kidney fat supersedes that reported by Roper (1970)
due to refinement of the statistical treatment.
Some information gained
from the collection of yearling buck antlers and does killed from highway
accidents and miscellaneous causes is included.

Kidney Fat and Bone Marrow Fat Indices
Deer were collected each winter commencing on January 15 and continuing
through April of 1969, 1970 and 1971. Tables 1, 2 and 3 summarize these
data. Kidney fat estimates were determined for deer killed by autos
and other miscellaneous causes when possible.
Data from collected deer
is analyzed separately as we suspect differences from deer killed by
other causes.
This hypothesis will be tested and reported in the final
report.
Kidney fat percentages are obtained using the method reported by Anderson
and Medin (1965). In collected deer for 1969, 1970 and 1971 the right
kidney has significantly (P&lt;.05) higher fat percentages than the left.
In order to compare to the results reported by other workers these data
are pooled.
The right femur bone is collected and quick frozen, then the entire
marrow content is mixed and analyzed for fat content using the ether
extract method.
In 1971 the percentage
year.

of kidney fat was lower in January

than any other

Regression lines for 1969 and 1970 are linear and appear quite different
than the polynomial regression obtained for 1971 (Fig. 1); however, most
of the polynomial regression in 1971 results from low January values.
At present the biological implications, if any, are not clear. Values
for February through April are comparable for each year (Fig. 2). Even
though these data suggest differences for January, Bartlett's test
(Snedecor and Cochran 1968) for homogenity of variances for each month
suggests there is no significant differences between the variances (P&lt;.25)
when all 3 years are pooled.
An F test indicates no significant difference between the means.
Apparently the values for January are too variable to be significant even though the data for 1969 and 1970 fit a linear
regression while the 1971 data fit a polynomial regression.
Plans are to
analyze all the kidney fat data by separate age classes using a multiple
regression analysis to try and account for some of the variation.

�- 213 -

Table 1. Kidney and bone marrow fat percentages for deer collected
in Middle Park in 1969.

Collection
Date

Day
No.

Right
Kidney

1-15
1-16
1-17
1-21
1-21
1-22
1-23
1-23
2-4
2-5
2-6
2-7
2-7
2-17
2-18
2-18
3-4
3-5
3-6
3-6
3-7
3-l3
3-20
3-20
4-8
4-9
4-10
4-10
4-10
4-14
4-14
4-15

1
1
3
7
7
8
9
9
21
22
23
24
24
34
35
35
49
50
51
51
52
58
65
65
84
85
86
86
86
90
90
91

11.9
9.6
59.7
38.8
44.0
30.3
32.8
30.8
22.5
46.6
39.5
42.8
32.1
25.7
30.3
9.8
10.2
26.2
10.8
46.0
26.0
14.0
18.8
16.0
8.8
5.3
14.0
9.0
20.6
15.3
7.0
4.3

Percent Fat
Left
Kidney

6.7
79.8
29.6
23.8
32.0
26.9
30.3
25.8
48.5
49.6
33.0
31.9
30.4
22.7
7.9
14.5
19.5
8.0
37.7
23.9
10.4
16.3
11.1
7.5
9.0
4.7
20.6
10.7
7.4
3.8

Bone
Marrow

85
92
90
86
84
87
91
84
91
94
91
89
92
9
91
85
88
93
92
84
91
87
56
60
88
78
84
89
51
11

�- 214 -

Table 2. Kidney and bone marrow fat percentages for deer collected
in Middle Park in 1970.

Collection
Date

Day
No.

Right
Kidney

1-26
1-26
1-27
1-29
2-5
2-9
2-9
2-10
2-12
2-12
2-13
2-25
2-26
2-27
2-28
3-12
3-19
3-20
3-23
3-23
3-23
3-25
3-27
4-7
4-7
4-9
4-13
4-13
4-14
4-14
4-15

12
12
13
15
22
26
26
27
29
29
30
42
43
44
45
57
64
65
68
68
68
70
72
83
83
85
89
89
90
90
91

17.6
22.8
40.4
26.9
32.7
23.0
38.5
23.8
28.8
.04
20.0
14.9
10.5
13.7
41.1
28.9
15.2
5.6
18.0
4.6
14.4
7.9
3.7
7.2
14.2
9.2
4.8
6.4
.03
9.5

Percent Fat
Left
Kidney

55.1
18.8
51.5
23.8
31.0
41.0
20.2
29.3
.04
20.0
15.2
5.1
25.7
17.7
14.1

6.8
18.4
6.7
3.1
7.4
21.8
6.6
7.5
10.5
7.3
.02
11.4

Bone
Marrow

90
89
76
84
91
91
95
92
74
3

85
89
53
86
88
81
85
66
86
66
79
61
33
82
85
30
72
33
40
2
72

�- 215 -

Table

3.

Kidney

fat percentages

for deer collected

in Middle

1971.
Collection
Date

1-16
1-19
1-22
1-22
1-25
1-26
1-27
1-28
1-28
2-5
2-9
2-10
2-16
2-17
2-23
2-26
3-4
3-8
3-11
3-16
3-19
3-22
3-26
3-30
4-1
4-5
4-7
4-12
4-19
4-21
4-23

Day
No.

2
5
8
8
11
12
13
14
14
22
26
27
33
34
40
43
49
51
56
61
64
67
71
75
77
81
83
88
95
97
99

Percent
Right Kidney

22.5
15.8
23.0
8.2
18.9
27.4
23.8
38.9
23.6
27.3
24.7
34.1
24.1
37.5
37.8
22.7
21.1
17.3
19.1
10.4
20.2
6.0
19.3
17.7
17.4
5.8
12.8
12.1
10.8

Park in

Fat
Left Kidney

15.3
14.3
19.4
23.3
5.2
17.4
19.5
15.6
30.4
19.9
16.5
26.9
32.0
26.3
26.4
45.3
17.5
17.6
15.0
16.2
9.5
26.5
7.6
16.5
20.1
5.8
10.7
5.2
16.7
18.4
10.5

�- 216 -

REGRESSION EQUATIONS

1969

n = 30

Y = 42.57914 + -.32565X
X = 28.95
r
.6561
r2
.4304
S.E.
23.3

1970

n = 26

Y = 37.72300 + -.31327X
X = 33.25
r = .7190
r2 = .5170
S.E.
17.3

1971

n = 30

100

CI.l

50

:&gt;-&lt;

~

z
A

H

::.::

45

E-!

r:r..

~

H

40

X = 30.015
r
.4425
r2
.1958
15.5
S.E.

A

~
E-!

::r::

'"

35

0
H

p::

r:r..
0
E-!

Y = 25.85995 + -.10869X

.

30

&lt;:
r:r..
:&gt;-&lt;

~

25

z
A

H

::.::

20

E-!

Z
~
u
p::

~
P-&lt;

1971
15

1969

z
&lt;:
~

::&lt;::

1970

10
5
0
0

10

20

30

40

50

60

70

80

90

100

COLLECTION DAYS COMMENCING JANUARY 15

Fig. 1. Regression of pooled right and left kidney fat percentages in deer
collected in Middle Park commencing January 15 for 1969, 1970 and 1971.
A test for equality of the three regression lines is significant (P&lt;.025)
and indicates there is no significant difference between years.

�- 217 100

75

70
Upper
~Confidence

Limit

65
60

55

50

45

40

Mean

35

30

25

20

.15
10

5

~

o
JFMA

1969

Lower Confidence Limit

J F M A

J F M A

1970

1971

MONTH AND YEAR

Fig. 2. Mean kidney fat percentage by month for collected deer in Middle
Park during the 1969, 1970 and 1971 winters.
(95 percent confidence
interval).

�- 218 -

Data from animals killed from causes other than collections have not
been analyzed to date. Tables 4 and 5 summarize the kidney fat percentages estimated in these deer in 1969 and 1971.
Analyses for bone marrow fat content in 1971 are in progress and will
be reported in subsequent reports.
Summaries of bone marrow fat for
1969 and 1970 are shown in Tables 1 and 2.

Table 4. The percent of fat on the kidneys and in the bone marrow
deer killed by autos and other causes in Middle Park in 1969.

Collection
Date

Day
No.

Right
Kidney

2-3

20

35.1

2-5

22

21. 6

2-11

28

2-28

45

3-2

Percent Fat
Left
Kidney

of

Bone
Marrow

36.3
88
52.0

43

20.0

26.5

74

47

10.0

4.8

51

3-3

48

26.5

14.6

88

3-4

49

19.0

11. 6

67

3-5

50

7.9

3.1

25

3-17

62

3.7

2.2

3-17

62

48.8

45.5

Blood Counts
A blood smear was made for each collected deer to provide possible indications of blood composition changes that may be related to tbeir physical
condition.
These data are summariz~d in Table 6 and graphed in Fig. 3.
Analyses will be made in a subsequent report.

�- 219 -

Table 5. The percent of fat on the kidneys
other causes in Middle Park, 1971.

of deer killed by autos and

Percent

Fat

Collection
Date

Day
No.

1-27

13

27.7

23.6

2-1

18

25.6

24.0

2-4

21

14.4

15.6

2-9

26

2-15

32

13.3

14.6

2-22

39

12.8

11. 5

2-22

39

3.0

2.9

2-26

43

5.2

5.9

3-4

49

35.2

15.6

3-11

56

9.6

5.1

and Number of Points in Yearling

Bucks

Antler Weights

Right Kidney

Left Kidney

8.7

We suspect that the weights and/or number of points of antlers on yearling
bucks could be indicative of the previous winters severity (Roper 1970).
The left antler of yearling bucks was sawed off just above the burr at
hunter check stations.
Weights of the left antlers from bucks taken in the 1968, 1969 and 1970
hunting seasons show no significant difference between the years when the
data are subjected to an analysis of variance test (Table 7). In Fig. 4,
the mean weight, confidence interval and the maximum and minimum weights
show a wide range for each year.

�Table 6.

Summary of blood counts on collected deer in Middle Park, 1971.
Blood Count

Collection
Date

Day
Number

Deer
Number

1-22
1-22
1-26
1-27
2-5
2-16
2-17
2-23
3-4
3-8
3-11
3-16
3-19
3-26
3-30
4-1
4-5
4-7
4-12
4-19
4-21
4-23
4-26

8
8
12
13
22
33
34
40
49
53
56
61
64
71
75
77
81
83
88
95
97
99
102

71-3
71-4
71-6
71-7
71-10
71-13
71-14
71-15
71-17
71-18
71-19
71-20
71-21
71-23
71-24
71-25
71-26
71-27
71-28
71-29
71-30
71-31
71-32

Segmented
Neutrophi1s

40
26
38
38
26
53
30
34
38
30
66
75
80
81
40
53
75
42
72
60
54
20
62

Eosinophils

0
16
16
2
0
6
6
10
0
10
10
3
0
0
29
13
8
23
2
0
14
8
0

Basophi1s

Lymphocytes

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0

60
56
46
60
66
37
64
54
62
58
24
22
20
19
27
34
17
31
26
38
30
70
38

Monocytes
0
2
0
0
8
4
0
2
0
2
0
0
0
0
4
0
0
4
0
2
2
2
0

N
N
0

�-~p~"~.iI~~~~--~~~~--~-

100

'" ,+""

~~

- --........---

I ------

~___
=z-

)

........--

90-

~

~

c

~

A

Mo~ocytes

s

~

80_
Lymphocytes

70_
60_
H

S
o

N
N

50_ u
~
o

t-'

H

40_Lx.:]z
~
~

Lx.1

30_

'tooSi~ot;nilS

20_
Segmented

Neutrophils

10_.

o~

,---

10

_.

-------.20

30

40
COLLECTION

Fig. 3. Blood composition data from collected
Note:. Deer 71-3 not shown.

50

60

DAY NUMBER

deer in Middle Park in 1971.

70

80

90

100 10:

�- 222 -

240
N=99

200

160

120

80

40

Minimum

o
1968

1969

1970

YEAR

Fig. 4. Antler weights in grams of the left antler of yearling bucks in
Middle Park (1968, 1969, 1970).

�- 223 -

30

-

· ....
·
·
·· ·
··
·· ··
·· ··
··

25 -

l

20

-

15

-

... ·

(f.l

z
0

0

~

0

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...

S

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-

1--

"-

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- - (·

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10 -

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~

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·

E-l

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I
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r-I
C'.I

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-er

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0
00
I
r-I

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0
r-I
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r-I
00

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C'.I
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1
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0
00
r-I
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\0

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0
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C'.I
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00
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C'.I

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C'.I
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C'.I
C'.I

CLASS OF WEIGHTS IN GRAMS
Year:
1968
1969
1970

.....

Fig. 5. Frequency distribution of I-year old buck antler weights from
Middle Park.

�- 224 -

Table 7. Analysis of variance of the antler weights of yearling bucks
taken during the 1968, 1969 and 1970 hunting seasons.

Source of Variation

Degrees

of Freedom

Mean Square

3.00

248 (N-l)

Total
Years

(between-among)

Error

(within-residual)246

2 (K-l)

909.38

(N-K)

1318.88

.6895

To provide a base for comparisons these data have been pooled and will be
used to represent winters of light stress on the deer. The mean of the
pooled data is 74.2 ± 4.5 grams (standard error = 2.2985).
A frequency distribution of the number of points on yearling buck antlers
suggests that each years data is similar (Fig. 5). A contingency test
shows that each years data is independent and that there is no significant
difference between years.
We could expect on the basis of this test that
the same proportion of 1, 2, 3 and 4 point antlers should occur each year.

LITERATURE

CITED

Anderson, A. E., and D. E. Medin.
1965. Two condition indices of the
Cache la Poudre mule deer herd and their application to management .•
Outdoor Facts, Game Information Leaflet No. 23. Colo. Game, Fish
and Parks Dept.
3 p.
Gill, R. B. 1969. Middle Park ·deer study -- food habits and physical
characteristics.
Colo. Div. Game, Fish and Parks.
Game Res. Div.
Federal Aid Proj. W-38-R.
Game Res. Rept., July.
Roper, L.A.
1970. Middle Park deer study -- food habits and physical
characteristics.
Colo. Div. Game, Fish and Parks.
Game Res. Div.
Federal Aid Proj. W-38-R.
Game Res. Rept., July.
Snedecor, G. W., and W. G. Cochran.
19'68. Statistical
Iowa State Press.
6th ed. 2nd printing.
593 p.

Prepared

byck 0 t?~&lt;---1kr

Laren A. Roper
Asst. Wildlife Researcher

methods.

The

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                  <text>- 225 July, 1971

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~~--------

Project No.

W-38-R-25

Work Plan No.

14

Job Title

Middle Park Deer Study - Range Fertilization

Deer-Elk
Job No.

Investigations
5

Period Covered:

June 15, 1970 through March 15, 1971

Personnel:

R. Bruce Gill, Paul F. Gilbert, Laren A. Roper, O. C. Wallmo,
Don W. Reichert, Wayne L. Regelin, Robert Davies, Charles
Grand Pre, Robert Glaze and Len H. Carpenter.

ABSTRACT
First year fertilizer and herbicide treatment response data were gathered
on th~ee study areas in western Grand County, Colorado.
Herbage yield
measurements were made on each of ten treatments at each study area. A
general linear increase in vegetation production was obtained with each
increased increment of nitrogen fertilizer from 30 pounds nitrogen per
acre to 120 pounds nitrogen per acre. Fertilizer and herbicide treatments
together generally resulted in equal production from treatment to treatment.
Botanical composition measurements were made at each study area with no
significant measurable changes being noticed.
Eight tame, trained mule
deer were grazed at each study area during the winter months.
A substantial
increase in shrubs and grasses was noted in the diet from pretreatment data
to the posttreatment data. A corresponding decrease in the forb percentage
of the diet was also noted in this period.
Treatment preference data showed
a definite preference by the deer for the control and fertilizer only treatments over the herbicide only or the herbicide and fertilizer combination
treatments.
A comparison of the 1969 and 1970 monthly and annual precipitation data with the thirteen year average was made.

��- 227 -

MIDDLE PARK DEER STUDY - RANGE FERTILIZATION
Len H. Carpenter

P. S. OBJECTIVE
To test the effects of fertilizers on deer forage and deer feeding intensity
to fertilization of critical winter ranges.

SEGMENT OBJECTIVES
1.

To test the effects of nitrogen and 2,4-D on forage production
lected deer winter range areas.

2.

To evaluate

deer foraging responses

on se-

to treated areas.

METHODS AND MATERIALS
Methods and materials
by Carpenter (1970).

employed in this segment have been detailed previously
Certain changes have been made.

Vegetation

Measurements

Herbage Yield
Herbage yield measurements were made with an electronic capacitance instrumentmodel
18-1000 in place of model 101-X used previously.
A total of 75
subplots was read for each 100' X 100' treatment plot. A ratio of five
mete.r read plots to one clipped plot was used resulting in a total of 15
clip plots per treatment.
The location of these 75 sUbplots was chosen in
a restricted random fashion so that they would cover the entire treatment
in a representative manner.
The procedure used for a clipped plot was as follows:
first, the immediate
vegetation around the edge of the meter that was not part of the subplot
was clipped and removed to avoid interference; next, plastic golf tees were
placec under the corner probes of the instrument so the exact sUbplot location
could be relocated when the instrument was removed to do the clipping.
After
the golf. tees were located, a first meter reading (MR A) was mad-e. This recorded the total vegetation on the subplot.
The meter was then removed and
the leafy shrub components of the subplot clipped and sacked sE::paratelyby
shrub species.
The meter was then replaced on the subplot and a second
meter reading made and recorded.
The meter was then removed and the forb
and grass components of the subplot were clipped and sacked sefarately.
In
addition, all woody material on the sUbplot was clipped and sacked separately

�- 228 -

by species for analysis of its effect on the meter reading.
A residual
meter reading was not taken for previous information indicated it was
not necessary.
All production data were computed on an oven-dry weight
basis (1000 C. for 24 hours).

Botanical

Composition

Considerable difficulty was encountered in delineating and counting the
number of grass plants per unit area on the permanently established plots.
To facilitate and improve this situation a grid of 25 2.4 inch squares
was placed on the square foot plot and the number of grids in which a particular grass species occurred was recorded.

Deer Grazing Measurements
In an attempt to maximize data obtained from the tame mule deer certain
changes were made in procedures.
The length of the grazing period at each
study area was lengthened from four to six days. One grazing period was
held at each study area during the winter months.
The number of deer used
for these measurements was increased from four to eight and the deer were
all of equal age (6-8 months).
Also, in order to maximize the data obtained,
each grazing method was alternated daily. That is, if the grazing trial
for day one was the plant preference or plant selection trial, then day two
would be the treatment preference or tower observation trial. The sequence
of these intervals was randomly chosen at each area. This system allowed
the observers to watch the deer for longer continual time periods and allowed
the deer to develop more standard daily patterns.
At approximately three
hour intervals the deer were rotated between observers in both grazing methods each day.

Treatment

Preference

Measurements

To increase the information obtained from the observation tower the number
of deer observed for each trial was increased from four to six. Each observer
was responsible for observing three deer. The time spent on each treatment
plot was recorded for all three deer by clock minutes while the grazing seconds
were obtained for only one deer by using a cumulative stop watch as that
particular deer grazed. All six deer were used throughout the day and rotated
between observers.

RESULTS AND DISCUSSION

Vegetation

.Herbage

Measurements

Yield

All herbage production data were analyzed using a double sample computer
program.
Production estimates were obtained for each one of the ten treatments

�- 229 -

at each study area. In order to develop a more accurate predictive
equation the data from the five fertilizer treatments (control included)
and the data from the five fertilizer and herbicide treatments (control
included) were combined into one equation resulting in a small sample
(clip) N of 75 and a large sample (meter read only) N of 375. The data
indicated that this combination was feasible.
This predictive equation
was then used to obtain the production estimates for each of the five
individual treatments in that group for each study area. Predictive
equations were developed for: (1) total leafy vegetation (Weight E)
which consisted of all the shrub leaf and current growth plus all forbs
and grasses; (2) shrub leaf and current growth only (Weight C). These
equations were developed using the first meter reading or (MR A) and the
corresponding above mentioned herbage weights.
The above mentioned two
components of the vegetation gave the best relationship between meter
readings and herbage weight in most cases. In certain cases a more
precise estimate of shrub leaf and current growth weight could be obtained
by relating the second meter reading minus the first meter reading (MR C)
against the shrub leaf and current growth weight.
However, the first
method in most all cases gives an accurate prediction of the shrub current
growth yield.
The above mentioned methods were applied to the 1969 pretreatment data to obtain comparable values to the first year treatment
response data. Tables 1, 2, and 3 present these data. In order to obtain
an estimate of forb and grass production the clipped weights of each group
were used. These data for all three study areas combined are presented in
Figures 1 and 2. No real differences are evident in the grass production
for each treatment.
There is a slight increase in grass production on
the combination treatments as compared with the fertilizer only treatments.
The forb data suggest a lower production on the herbicide plots which would
be expected.
No real differences in forb production are apparent on the
fertilizer only treatments.
The forb and grass responses should be quite
evident in future measurements when the full treatment effects are present.
Figures 3 and 4 show the linear response obtained from the nitrogen fertilizer only application on shrub leafy material and on total leafy material.
An analysis of the data for main effect of fertilizer, and the interaction
effect of fertilizer and herbicide, shows a significant (p = .01) linear
response to fertilizer in both shrub and total leafy material.
Tests for
quadratic, cubic and quartic regressions showed that these contributed little
to the sums of squares in the analysis of variance.
Although there is a
significant linear response there seems to be a decrease in production at
the 90 pound nitrogen treatment.
This is difficult to understand, but it
may be that certain components of the vegetation are not responding to this
particular nitrogen level. No explanation of why this might be is available
at this time.
Figure 5 illustrates the production response obtained in total leafy material
at each study area as a function of increased nitrogen levels, with 2,4-D
herbicide in combination.
In general, this herbicide and fertilizer combination tended to result in rather uniform productivity in all treatments.
This is probably due to a somewhat uniform decrease in the shrubby leaf
material as a result of the herbicide application.
Figure 6 shows the average
percent change in the total leafy material produced for each nitrogen level
from the control treatments (0 nitrogen and 0 herbicide) for all three study
areas. This graph illustrates the general linear increase with increased
nitrogen levels and the unexplained drop at the 90 pound nitrogen rate.

�- 230 Table l. Pre-treatment and first year herbage production response data for
Junction Butte study area. Values are in pounds per acre on a dry weight
basis.
Year and Weight Component
1969
1970
1969
1970
Shrub Current Growth Weight
Total Leafy Vegetation

(Lbs. Per Acre)
Treatment

o lb. N. 0 lb. 2,4-D

404

582

649

905

30 N

"

236

739

443

1099

60 N

"

464

957

723

1316

90 N

"

392

940

637

1278

120 N

"

477

1254

732

1603

o N 2 lb.

"

249

365

454

709

30 N

"

251

530

461

905

60 N

"

427

624

675

1001

90 N

"

337

424

585

781

120 N

"

485

549

744

915

2,4-D

Table 2. Pre-treatment and first year herbage production response data for
F1ight1ine study area. Values are in pounds per acre on a dry weight basis.

(Lbs , Per Acre)
Treatment

o lb. N. 0 lb. 2,4-D

Year and Weight ComEonent
1969
1970
1969
1970
Shrub Current Growth Weight
Total Leafy Vegetation
467

606

1177

1519

"

419

497

1187

1399

"

424

861

1134

1852

704

741

1526

1704

822

1625

1620

2633

663

393

1463

1212

696

412

1463

1222

60 N

"
"

802

582

1593

1437

90 N

"

404

290

1147

1059

120 N

"

398

357

1090

1162

30 N
60 N
90 N
120 N

"
"

o lb. N. 2 lb. 2,4-D
30 N

�- 231 -

Table 3. Pre-treatment and first year herbage production response data
for Corral Creek study area. Values are in pounds per acre on a dry
weight basis.

(Lbs , Per Acre)
Treatment

o lb. N. 0 lb. 2,4-D

Year and Weight ComEonent
1969
1970
1969
1970
Shrub Current Growth Weight
Total Leafy Vegetation
435

446

1214

1143

30 N

"

722

713

1494

1512

60 N

"

513

651

1274

1416

90 N

"

632

747

1404

1577

120 N

"

422

562

1173

1332

589

188

1398

990

o lb. N. 2 lb. 2,4-D
30 N

"

600

233

1374

1034

60 N

"

331

225

1115

1082

90 N

"

414

227

1173

1080

120 N

"

297

225

1077

1107

�- 232 -

350

r-I

o
!-&lt;

+J
C

o

O~

u

o
I
o
C"')

o
I
o

\0

__ ~ __ ~~~
Control
Only

o
I
o
0'1

~

o
I
o

N

r-I

and Fertilizer

Fig. 1. Forb production
combined.

by treatment

N

I

o

~

N
I

o
C"')

N
I

N

\0

0'1

o

I

o

N
I

o
N

~r-I

Fertilizer and Herbicide
Combination

for all three study areas

_

�- 233 -

"""
CIJ
H
Cj

~

H
CIJ

p...

rn

'"d

t::

;::l
0

p...

'-'
t::
0
,,-1

..,
u
;::l

-e

200

--I'

0
H
p...

!
rl

100

o

..,H
t::

o

u

o
I
o
C"')

o
I
o

\0

o
I
o
0"1

N
I

o

N
I

o
C"')

N

I

o

\0

N

I

o
0"1

N

I

o

N

rl

0
Control and Fertilizer
Only

Fig, 2. Grass production
combined.

Fertilizer and Herbicide
Combinations

by treatment

for all three study areas

�- 234 -

2800
Flightline

Junction
Butte
Corral Creek

o~ __ -.

~

o

30

~

~

~

60

90

120

Pounds of Actual Nitrogen

Fig. 3. Total leaf material
of nitrogen.

produced

Per Acre

as a function of increasing

levels

_

�- 235 -

1800
Flightline

Junction

Butte

Corral Creek

o

30

60

90

120

Pounds of Actual Nitrogen Per Acre
(Ammonium Nitrate 335%)

Fig. 4. Shrub leaf material
of nitrogen.

produced

as a function of increasing

levels

�- 236 -

1800

Flight line
____------------------~::::~------~~~~~~~----

Corral Creek
Junction Butte

0/2

30/2

60/2

Pounds of Elemental Nitrogen
Pounds of 2,4-D Herbicide

90/2

120/2

Per Acre With 2
Per Acre

Fig. 5. Total leafy vegetation produced as a function of increased
levels of nitrogen with 2,4-D herbicide at three study areas.

�- 237 -

30

60

90

Pounds of Elemental

Nitrogen

Fig. 6. Percent change in production
the mean of the control treatments.

120
Per Acre

of total leafy vegetation

from

�- 238 -

Botanical

Composition

The botanical composition measurements were made in late July, 1970.
The herbicide was applied in early June, 1970 which left a relatively
short period of time between application and measurement.
This short
time period made it difficult to assess herbicide effect. Mortality
rates of affected plants should be evident in the 1971 measurements.
Fertilizer was applied in late October, 1969 resulting in less than a
full year response, which made it unlikely noticeable botanical composition changes would be observed.
Due to these facts only general
comments on botanical composition changes will be made in this report.
In general, the kill of sagebrush due to the herbicide ranged from about
25 percent to 85 percent over the three study areas. No pattern of
enhanced or decreased sagebrush kill could be seen with regard to a particular fertilizer and herbicide treatment.
The Junction Butte area
resulted in the lowest percent kill of sagebrush on nearly all herbicide
treatments.
Soil moisture stress and type of growth form of the sagebrush probably accounted for this fact. A definite decrease of most
forb plants was recorded on the herbicide plots.
Nitrogen applications
seemed to encourage abundance of western wheatgrass (Agropyron smithii),
bluebunch wheatgrass (Agropyron spicatum), and squirreltail grass
(Sitanion hystrix) noticeably.
The 1971 measurements should elucidate
these factors.

Deer Grazing Measurements

Plant Preference

Data

A summary of the diets of the eight tame, trained mule deer for each
study area for both pre-treatment (1969-70) and the first year's treatment data (1970-71) is given in Tables 4, 5, and 6. Table 7 gives all the common
and scientific names of the plants used in this report.
Table 8 summarizes the diet by plant components for all three study areas for both years.
It can be seen from this summary that shrubs and grasses increased substantially in the diet from the pre-treatment data to the post-treatment
data. Correspondingly,
there is a considerable decrease of forbs in the
diet from pre-treatment to post-treatment data. Tables 9, 10, and 11
summarize the diet by plant components for each study area. The greatest
increase in shrub and grass percentages of the diet from pre-treatment to
post-treatment occurred at the Corral Creek area. In addition, a dramatic
drop in the forb percentage was shown at this area. This forb decrease
was probably due to two main factors.
One, snow depths during the posttreatment grazing trials were greater than during the pre-treatment trials.
The snow depth was approximately 12 inches as contrasted with about a three
inch average during pre-treatment trials. This increased snow depth precluded use of many of the forb plants that were available in the more
shallow snow of the year before.
Secondly, the forb decrease can be partially accounted for by the kill obtained with the herbicide applications.
This can be better illustrated by the Flightline study area data shown in
Table 10. The snow conditions for this area were similar both winters yet

�- 239 -

there still was approximately a 11 percent decrease in the forb component
of the diet. The increase in the grass percentage of the diet at all
three study areas is quite impressive.
Over all three areas the increase
averaged nearly eight percent from pre-treatment to post treatment grazing.
Most of this increase in the grass component came with such species as
bluebunch wheatgrass, squirreltail grass and western wheatgrass.
Each of
these species seemed to be first to respond to the treatments.
Table 12
gives a cOlnparison of selected plant species in the diet for both winters
over all three study areas. This summary shows that sagebrush remained a
small portion of the tame deer diets and changed very little from pretreatment to post-treatment grazing.
Green rabbitbrush (Chrysothamnus
viscidiflorous) which seemed to respond favorably to the fertilizer showed
a significant increase in the diet from pre-treatment grazing to posttreatment grazing.
Bitterbrush (Purshia tridentata) still comprised a
large percentage of the diet and did not change appreciably from pretreatment grazing.
Table 13 gives a summary of the total number of bites of all species of
plants taken on each treatment at each study area for both winter grazing
trials.
This summary shows that about 60 percent of the bites were taken
on the control or fertilizer only treatments while 40 percent were taken
on the herbicide or herbicide and fertilizer combination treatments.
In
comparison, 46 percent of the bites during pre-treatment trials were taken
on the control or fertilizer only plots and 54 percent on the herbicide or
fertilizer and herbicide combinations.
Table 14 illustrates the corresponding decrease in the forb and shrub
component of the diet and the increase in the grass component with the
herbicide or fertilizer and herbicide combinations.
This table shows
about 36 percent of the grass was taken on the control and fertilizer
only treatments compared with 64 percent on the combination treatments.
Ninety-one percent of the forbs were taken on the control and fertilizer
plots and only nine percent on the combination treatments.
Correspondingly,
64 percent of the shrubs were taken on the control or fertilizer only
treatments while 36 percent were taken on the herbicide or fertilizer and
herbicide treatments.

Treatment

Preference

Data

Tables 15, 16 and 17 summarize the treatment preference data for both
winter's grazing trials for all three study areas. Table 15 is a summary
of the total observed deer minutes spent per treatment plot. This summary
also indicates a greater preference by deer for the control and fertilizer
only treatments.
Nearly 64 percent of total deer minutes was spent on the
control or fertilizer only treatments and about 36 percent on the herbicide
or herbicide and fertilizer treatments.
These data contrast with the pretreatment data where the time spent per each group was roughly half and half.

�- 240 -

Table 4. Junction Butte study area grazing observation data and diet composition for the 1969-70 and 1970-71 winters.
1969-70

1970-71
Total Bites

Percent
of Diet

Plant Species

Total Bites

Percent
of Diet

Serviceberry

830

36.5

725

22.5

B1uebunch wheatgrass

375

16.5

326

10.1

Snowberry

288

12.6

210

6.5

Pine need1egrass

143

6.3

204

6.3

Snakeweed

131

5.7

Blue gramma

105

4.7

184

5.7

Buckwheat

92

4.0

294

9.1

Big sagebrush

64

2.8

85

2.6

Phlox bryoides

63

2.8

427

l3.3

Bitterbrush

62

2.7

22

0.7

Mutton grass

45

2.0

73

2.3

Fringed sage

27

1.3

Rabbitbrush
(Chrysothamnus spp.)

17

0.8

Lichens

12

0.5

Oregon grape

6

0.2

Penstemon cyathoporus

4

0.2

Unknown grass

4

0.1

52

1.6

Western wheatgrass

2

0.1

263

8.2

Phlox multiflora

2

0.1

Junegrass

2

0.1

5

0.1

Green rabbitbrush

290

9.0

Needle and thread grass

14

0.4

Bluebell

5

0.2

Bott1ebrush squirre1tai1
grass

18

0.6

Currant

7

0.2

Indian ricegrass

15

0.5

3,222

100.0

Totals

2,276

100.0

�- 241 -

Table 5. F1ight1ine study area grazing observation data and diet composition
for the 1969-70 and 1970-71 winters.
1970-71

1969-70

Total Bites

Percent
of Diet

Plant Species

Total Bites

Percent
of Diet

Bitterbrush
Buckwheat

2,075
462

57.2
12.7

4,051
493

53.1
6.4

Snowberry

290

8.0

299

3.9

Pine need1egrass

176

4.9

359

4.7

B1uebunch wheatgrass

122

3.3

28

0.4

Mutton grass

95

2.6

545

7.1

Mat penstemon

95

2.6

12

0.2

Phlox bryoides

84

2.3

Unknown grass

58

1.6

523

6.8

Serviceberry

45

1.2

299

3.9

Paintbrush

43

1.2

30

0.4

Green rabbitbrush

14

0.4

593

7.8

Rose
Penstemon spp.

13
11

0.4
0.3

38
4

0.5
0.05

Phlox multiflora

11

0.3

46

0.6

Sedge

9

0.2

18

0.2

Unknown forbs

6

0.2

5

0.1

Bottlebrush squirre1tail
grass

5

0.1

147

1.9

Penstemon cyathoporus
Big sagebrush
Junegrass
Poa spp.

4

0.1
0.1
0.1

12
34

0.2
0.4

63

0.8

Knotweed
Rothrock sage
Toad flax
Arrow1eaf ba1samroot
Lupine
Astragalus conva1larius
Oregon grape
Rockcress
Bluebell

2
1

19
5
5
5

0.2
0.1
0.1
0.1
0.05

Totals

4

3

0.1
0.1

4

1
1
3,628

100.0

7,629

T

T
100.0

�- 242 -

Table 6. Corral Creek study area grazing observation
for the 1969-70 and 1970-71 winters.
1969-70

data and diet composition

1970-71

Total Bites

Percent
of Diet

Total Bites

Percent
of Diet

Bitterbrush

892

33.6

2,294

32.3

Buckwheat

756

28.5

37

0.5

Mat penstemon

375

14.1

Mutton

231

8.7

Pine need1egrass

71

2.7

Pusseytoes

70

2.6

Serviceberry

60

2.3

1,050

14.8

B1uebunch

57

2.2

1,599

22.4

Snowberry

45

1.7

863

12.2

Phlox multiflora

45

1.7

9

0.1

Green rabbitbrush

16

0.6

808

11.4

Unknown

forbs

6

0.2

19

0.3

Senecio

spp.

6

0.2

Senecio mu1ti1obatus

6

0.2

Gilia

5

0.2

Aster spp.

3

0.1

Sedge

3

0.1

juniper

2

0.1

190

2.7

conva11arius

2

0.1

11

0.1

1

0.1

Plant Species

grass

wheatgrass

Rocky Mountain
Astragalus
Senecio

integerrimus

Unknown

grass

104

1.5

Big sagebrush

45

0.8

21

0.3

19

0.3

11

0.1

6

0.08

Paintbrush

5

0.07

Poa spp.

4

0.05

7,095

100.0

Bott1ebrush

squirre1tai1

grass

Horsebrush
Chrysothamnus
Penstemon

Totals

nauseosus

cyathoporus

2,652

100.0

�- 243 -

Table 7.

Common and scientific

Scientific

Common Name

Arrowleaf balsamroot
Aster
Astragalus (Vetch)
Beardstongue
Big sagebrush
Bitterbrush
Bluebell
Bluebunch wheatgrass
Blue gramma
Blue grass
Bottlebrush squirreltail
Buckwheat
Currant
Fringed sage
Scarlet gilia
Green rabbitbrush
Horsebrush
Indian ricegrass
Junegrass
Knotweed
Lupine
Mat penstemon
Mutton grass
Needle and thread grass
Oregon grape
Paintbrush
Penstemon
Phlox
Pine needlegrass
Pusseytoes
Rabbitbrush
Rockcress
Rocky Mountain juniper
Rose
Rothrock sage
Sedge
Senecio (Groundsel)

Serviceberry
Snakeweed
Snowberry
Toad flax
Western wheat

names of plants used in the report.

grass

grass

Name

Balsamorhiza sagittata
Aster spp.
Astragalus convallarius
Pensterilon spp.
Artemisia tridentata
Purshia tridentata
Mertensia lanceolata
Agropyron spicatum
Bouteloua gracilis
Poa spp.
Sitanion hystrix
Eriogonum umbellatum
Ribes cereum var. inebriums
Artemisia frigida
Gilia candida
Chrysothamnus viscidiflorus
Tetradymia canescens
Oryzopsis hymenoides
Koeleria cristata
Polygonum sawatchense
Lupinus argenteus
Penstemon caespitosus
Poa fendleriana
Stipa comata
Berberis repens
Castilleja flava
Penstemon cyathoporus
Phlox bryoides
Phlox multiflora
Stipa pinetorum
Antennaria parviflora
Chrysothamnus spp.
Arabis drummondii
Juniperus scopulorum
Rosa woodii
Artemisia tridentata spp. rothrockii
Carex spp.
Senecio spp.
Senecio integerrimus
Senecio multilobatus
Amalanchier alnifolia
Gutierrezia sarothrae
Symphoricarpos oreophilus
Comandra umbellata
Agropyron smithii

�- 244 -

Table 8. Percent diet composition by plant components
areas combined for the 1969-70 and 1970-71 winters.

for all three study

1969-70
Percent of Diet

1970-71
Percent of Diet

Shrubs

55.5

66.4

Grasses

17.6

25.5

Forbs

26.9

8.1

Totals

100.0

100.0

Plant Component

Table 9. Summary of Junction Butte study area diet composition
ponents for the 1969-70 and 1970-71 winters.

by plant com-

1969-70
Percent of Diet

1970-71
Percent of Diet

Shrubs

56.7

41. 6

Grasses

29.7

35.8

Forbs

13.6

22.6

Totals

100.0

100.0

Plant Component

Table 10. Summary of F1ight1ine study area diet composition
for the 1969-70 and 1970-71 winters.

by plant components

1969-70
Percent of Diet

1970-71
Percent of Diet

Shrubs

67.3

69.5

Grasses

12.9

22.1

Forbs

19.8

8.4

Totals

100.0

100.0

Plant Component

�- 245 -

Table 11. Summary of Corral Creek study area diet composition by plant components for the 1969-70 and the 1970-71 winters.

Plant Component

1969-70
Percent of Diet

1970-71
Percent of Diet

Shrubs

38.3

74.4

Grasses

13.6

24.4

Forbs

48.1

1.2

Totals

100.0

100.0

Table 12. Comparison of certain selected plant species in the diet for the
1969-70 and 1970-71 winters, for all three study areas.
1970-71
Percent of Diet

Plant Species

1969-70
Percent of Diet

Bitterbrush

35.4

35.5

Serviceberry

10.9

11.5

Snowberry

7.3

7.6

Sagebrush

0.8

0.9

Green rabbitbrush

0.4

9.4

B1uebunch wheatgrass

6.5

10.9

Pine need1egrass

4.8

3.1

Mutton grass

4.4

3.4

Blue gramma

1.2

1.0

Bott1ebrush squirre1tai1 grass

t

1.0

Western wheatgrass

t

1.5

Buckwheat

15.4

4.6

Phlox

1.7

2.4

�Table 13. Total number of bites of all species of plants taken on each treatment at each study area during the
1969-70 and 1970-71 winters. 11
Treatment
(lbs. per acre)

o lb. N. 0 lb. 2,4-D

Jct. Butte
1970-71
1969-70

Flight line
1970-71
1969-70

Corral Creek
1969-70 1970-71

Totals
1969-70 1970-71

Percent of Total
1969-70 1970-71

722

354

1,127

1,443

461

214

2,310

2,01l

27.0

16.6

30

N.

"

104

293

173

182

6

61

283

536

3.3

4.4

60

N.

"

235

0

81

216

67

255

383

471

4.5

3.9

90

N.

"

197

147

84

53

86

1,833

367

2,033

4.3

16.7

120

N.

"

330

843

155

1,365

126

4

611

2,212

7.1

18.2

46.2

59.8

Subtotals

N

~

o lb. N. 2 lb. 2 ,4-D

18

20

881

655

15

27

914

702

10.7

5.8

30

N.

"

18

29

231

17

391

60

640

106

7.5

0.9

60

N.

"

261

152

126

1,645

591

745

978

2,542

11.4

20.9

90

N.

"

299

85

741

185

120

279

1,160

549

13.6

4.5

120

N.

"

92

143

29

99

787

740

908

982

10.6

8.1

53.8

40.2

100.0

100.0

Subtotals

Totals

2,276

2,066

3,628

5,860

2,650

4,218

8,554

12,144

11 Total bite numbers will not compare exactly to diet composition data because of bites that were taken on the
treatment margins.

'"

�- 247 -

Table 14. Grass, forb, and shrub component percentages of the diet by treatment plot for all three study areas for the 1970-71 winter.
(lbs. per acre)
Treatment

o lb. N. 0 lb. 2,4-D

Total Bites
Grass

Percent
Grass

Total Bites
Forbs

Percent
Forbs

Total Bites
Shrubs

Percent
Shrubs

83

2.8

525

46.0

1,403

17.4

30

N.

"

151

5.1

262

23.0

123

1.5

60

N.

"

95

3.2

92

8.1

284

3.5

90

N.

"

122

4.1

2

0.2

1,909

23.7

120

N.

"

619

21.0

156

13.7

1,437

17.9

36.2

Subtotals

o lb. N. 2 lb. 2,4-D

91.0

64.0

17

0.6

7

0.6

678

8.4

30

N.

"

61

2.1

6

0.5

39

0.5

60

N.

"

854

28.9

80

7.0

1,608

20.0

90

N.

"

385

13.0

10

0.9

154

1.9

120

N.

"

566

19.2

0

0.0

416

5.2

63.8

Subtotals
Totals

2,953

100.0

9.0

1,140

100.0

36.0

8,051

100.0

�Table 15. Total deer minutes spent on each treatment plot obtained during plant preference measurements at
each study area for the 1969-70 and the 1970-71 winters.
Corral Creek
1969-70 1970-71

Totals
1969-70 1970-71

Percent of Total
1969-70 1970-71

Treatment
(lbs. per acre)

Jet. Butte
1970-71
1969-70

o lb. N. 0 lb. 2,4-D

151

80

187

159

47

25

385

264

18.4

14.9

Flight line
1969-70 1970-71

30

N.

11

19

40

62

58

6

24

87

122

4.2

6.9

60

N.

11

69

2

48

32

29

54

146

88

7.0

5.0

90

N.

11

63

49

97

28

17

183

177

260

8.5

14.7

120

N.

11

116

210

70

195

50

7

236

412

11.3

23.3

49.4

64.3

Subtotals

N

~
co
I

o lb. N. 2 lb. 2,4-D

7

8

103

47

12

2

122

57

5.8

3.2

30

N.

11

21

12

82

19

135

12

238

43

11.4

2.4

60

N.

11

65

46

63

184

109

65

237

295

11.4

16.7

90

N.

11

97

17

167

39

10

35

274

91

13.1

5.1

120

N.

11

51

66

42

30

93

43

186

139

8.9

7.8

50.6

35.7

100.0

100.0

Subtotals
Totals

659

530

921

791

508

450

2,088

1,771

�Table 16. Total deer grazing seconds obtained from the observation tower for each treatment at each study
area for the 1969-70 and 1970-71 winters.
Flightline
1969-70 1970-71

Corral Creek
1969-70 1970-71

Totals
1969-70 1970-71

Percent of Total
1969-70 1970-71

Treatment
(lbs. per acre)

Jet. Butte
1970-71
1969-70

o lb. N. 0 lb. 2,4-D

1,643

2,366

1,700

1,331

325

1,402

3,668

5,099

15.0

10.8

30

N.

"

0

40

287

1,080

129

81

416

1,201

1.7

2.5

60

N.

"

3,609

163

574

2,712

0

503

4,183

3,378

17.1

7.1

90

N.

"

1,003

724

801

1,562

1,291

9,060

3,095

11,346

12.7

23.9

120

N.

"

496

4,700

2,285

7,172

0

10

2,781

11,882

11.4

25.1

57.9

69.4

Subtotals

N
.p\0

o lb. N. 2 lb. 2,4-D

0

230

876

498

30

24

906

752

3.7

1.6

30

N.

"

0

78

94

294

0

336

94

708

0.4

1.5

60

N.

"

1,462

711

777

2,493

1,787

1,791

4,026

4,995

16.5

10.5

90

N.

"

1,186

3,751

28

266

0

347

1,214

4,364

5.0

9.2

120

N.

"

0

1,272

653

901

3,359

1,543

4,012

3,716

16.5

7.8

42.1

30.6

100.0

100.0

Subtotals

Totals

9,399

14,035

8,075

18,309

6,921

15,097

24,395

47,441

�Table 17. Total number of deer minutes spent standing per treatment plot at each study area during the 1969-70
and 1970-71 winters obtained from the observation tower.
Treatment
(Lbs, per acre)

Jct. Butte
Flight line
1970...,.711969-70 1970-71
1969-70

o lb. N. 0 lb. 2 ,4-D

150

321

126

222

17

115

293

658

15.5

12.6

Corral Creek
1969-70 1970-71

Totals
1969-70 1970-71

Percent of Total
1969-70 1970-71

30

N.

"

1

26

24

131

8

28

33

185

1.7

3.5

60

N.

"

410

66

79

265

24

64

513

395

27.2

7.6

90

N.

"

23

103

60

150

84

798

167

1,051

8.9

20.1

120

N.

"

49

502

187

564

0

13

236

1,079

12.5

20.6

65.8

64.4

Subtotals

N
VI

0

o lb. N. 2 lb. 2 ,4-D

0

51

55

65

11

19

66

135

3.5

2.6

30

N.

"

0

62

17

56

3

52

20

170

1.1

3.3

60

N.

"

130

107

88

309

113

144

331

560

17.5

10.7

90

N.

"

92

354

8

40

1

54

101

448

5.4

8.6

120

N.

"

0

270

57

59

70

214

127

543

6.7

10.4

34.2

35.6

100.0

100.0

Subtotals
Totals

855

1,862

701

1,861

331

1,501

1,887 .

5,224

�Table 18. Comparison of the 1969 and 1970 monthly and annual average precipitation data with the thirteen
year average for the Green Mountain, Kremmling *, and Hot Sulphur Springs weather stations.l/

Year

Jan.

Feb.

Mar.

Apr.

May

Month
June

13-year average

0.89

0.76

0.91

0.99

1.04

1.55

1.24

1.66

1969

1.79

0.43

0.33

1.00

0.82

4.65

0.86

1970

0.76

0.31

1.17

1.22

0.13

1.41

0.85

*

-

l/ Data from Colorado Climatological

annual reports.

Annual Total

Dec.

1.61

1.14

0.80

0.97

13.56

1.23

1.50

3.56

0.49

1.23

17.90-

1.82

2.00

2.01

1.38

0.80

13.86

Aug. Sept.

Kremmling we athe r records are incomplete for this time period.

-

Oct. Nov.

July

~

V1
I-'

�- 252 -

This same trend of treatment preference is also seen in the tower observation data. Table 16 gives a summary of the deer grazing seconds obtained
from the observation tower for each treatment at each study area for both
winter's grazing trials.
These data show that about 70 percent of the
grazing time was spent on the control or fertilizer only treatments.
The
pre-treatment data also showed a slight preference for the control and
fertilizer only treatments.
Table 17 summarizes the total number of deer minutes spent standing on each
treatment plot at each study area for both winters as obtained from the
observation tower. Total standing minutes were used to remove the bias of
long periods of time accumulated on certain treatments when the deer bed
down in commonly used bedding sites. These data show a preference towards
the control and fertilizer only treatments also but the same trend was noted
in the pre-treatment data. It is felt that this was due mostly to the time
spent by deer near areas they prefer to bed in, such as corners, and had
little to do with their preference towards grazing.
With regards to individual treatment preferences no particular trends were
evident from the data. This may be due to the fact that with the narrow
increments of 30 pounds of nitrogen the deer are unable to select a treatment difference.
However, it is interesting to note that at the Junction
Butte and the Flightline study areas the 120 pound nitrogen treatment with
no herbicide was most preferred over all individual treatments.
Over all
three study areas this treatment received about 25 percent of the total
grazing seconds and total time spent by the deer.
An interesting observation was made with regards to the noted preference
for the control or fertilizer only treatments.
It was observed while
following the deer during the grazing trials that snow crust conditions were
quite different in the herbicide plots as contrasted to the fertilizer only
plots. With the removal of the shrub crown cover on the herbicide treatments the snow tended to make a more uniform, crusted blanket as opposed to
the broken, faster melting snow cover on the fertilizer only treatments
where the shrub crown cover remained.
The deer preferred to graze in and
around breaks in the snow cover adjacent to and under the shrubs.
In addition, the deer seemed to dislike traveling in the more crusted snow of the
herbicide treatments and avoided those areas.

Other Measurements

Weather

Information

Using data obtained from weather stations at Kremmling *, Green Mountain,
and Hot Sulphur Springs, a 13 year average of annual monthly precipitation
was obtained (Colorado Climatological data annual reports).
The 1969 and
1970 data were compared with this summary.
Table 18 presents these data.
The year the fertilizer treatments were applied (1969) was very moist, receiving almost 4.5 inches more moisture than the 13 year average.
October,
the month the fertilizer was applied, received about 3.5 inches of moisture
as contrasted to the 13 year average of 1.1 inches of moisture.
June of
1969 received 4.6 inches of rain compared with a norm of 1.55. This added

* Kremmling weather records are incomplete at this time period.

�- 253 -

moisture should have assured good infiltration and use of the nitrogen
fertilizer.
The first year of growth response (1970) was fairly normal
in total precipitation.
The 13-year average total precipitation was
13.56 inches, compared with 13.86 inches received in 1970.

Photographs
The permanent photo hubs were photographed with black and white and color
film at the end of the growing season for a permanent record.

LITERATURE

CITED

Carpenter, L. H. 1970. Middle Park deer study - range fertilization.
Colo. Div. Game, Fish and Parks.
Game Res. Div., Fed. Aid Prog.
W-38-R-24.
Game Res. Rept. July, Part 3. pp. 371-391.
Environmental Science Services Administration.
1965-1970.
data, Colorado.
U. S. Dept. Commerce.
70-75 (13).
U. S. Weather Bureau.
Dept. Commerce.

1958-1964.
70-75 (13).

Prepared
Carpe ter
Student Assistant

Climatological

Climatological

data, Colorado.

U. S.

��- 255 July,

1971

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R~25

Work Plan No.

14

Job Title

Deer-Elk

Investigations

Job No.
6 (Second Year)
Middle Park Cooperative Deer Study Physiology and Prevention of Deer Starvation

Period Covered:

June, 1970 through June, 1971

Personnel:

Julius G. Nagy, Principal Investigator; Gene G. Schoonveld,
Graduate Student; and David S. DeCalesta, Graduate Student.

ABSTRACT
Approximately 60 .mule deer were maintained for experimental use. Rumen
fistulation was unsuccessfully attempted with 12 deer, further attempts
with tame deer are planned.
Projected metabolic trials were postponed.
A simulated malnutrition diet fed 90 days did not appear to induce semistarvation in deer. Rumen-reticular contents of deer fed the malnutrition
diet and then fed dehydrated alfalfa hay exhibited differences in pH, VFA
concentrations, molar proportions of acetate:propionate,
and protozoa
numbers.
Fawns fed the dehydrated alfalfa hay were more prone to digestive
upsets than adults and exhibited elevated serum urea levels.
Total starvation of deer indicated: .(a) rumen-reticular numbers of bacteria and digestive effectiveness of these bacteria may not be seriously reduced by starvation;
(b) twenty-two days of starvation may be close to the point where
supplemental feeding cannot prevent deer losses;
(c) does and fawns may
succumb sooner than adult males to starvation.
Deer fed poor quality alfalfa
suffered abomasal hemorrhagic ulcerations and omasal food compaction.
Goats and sheep, with larger rumen-reticulums,
omasums and abomasums relative
to body size than deer, exhibited no upsets when fed the same ration.

��- 257 -

PHYSIOLOGY

MIDDLE PARK DEER STUDY AND PREVENTION OF DEER STARVATION
Julius G. Nagy

P. S. OBJECTIVE
To ascertain what takes place in deer physiology when starvation occurs,
and to investigate the economic and biologic feasibility of preventing
starvation.

SEGMENT OBJECTIVES
1.

Maintain

2.

Refine techniques for rumen content sampling
for feeding, handling and examining deer.

3.

Conduct metabolic

4.

5.

an experimental

herd of 15 to 20 mule deer.
and improve methodology

trials:

a.

To measure

b.

To determine the critical ambient temperature where
crease their rate of metabolism to combat cold.

Conduct

basic metabolic

starvation

rate.
deer must in-

trials:

a.

To determine
starvation.

changes in deer physiology

b.

To investigate
in vitro.

effects of starvation

c.

To determine the "critical" point of no return in deer starvation,
beyond which they will not recover.

d.

To investigate methods
point is passed.

Prepare

during the process

of

upon rumen microorganisms

of aiding deer recovery

before

the "critical"

report.

METHODS AND MATERIALS

Maintenance

of Experimental

Deer

Facilities for maintaining and handling experimental deer were described
previously (Nagy 1970). The experimental herd now numbers 60 deer. Additional deer are necessary for continued starvation trials and investigations

�- 258 -

into physiological
hay diets.

effects

of feeding deer high-fiber

alfalfa

or native

Rumen Fistulation
Rumen fistulation has been unsuccessfully attempted with 12 mule deer.
Although comparable surgical techniques used and rumen cannula developed
have been successful in fistulating goats, all deer died within 10-21 days
after distulation.
The apparent cause of death was infection within the
oblique abdominal wall muscles, predisposed by excitability and excess
movement of fistulated wild deer. Further fistulation attempts are planned
using tame deer.

Metabolic

Trials

Determination of deer basic metabolic rate was postponed in light of recent
work of Silver et al. (1971) and in preference for more detailed investigations into the apparent inability of deer to digest alfalfa hay.

Experimental

Trials

Deer Malnutrition
Nutritional changes of penned mule deer were monitored during a period of
prolonged malnutrition and upon subsequent feeding with alfalfa.
Five mule
deer does and four fawns were fed a formulated "malnutrition" diet (Table 1)
for 90 days from February to May and subsequently fed good quality dehydrated
alfalfa.
Daily food intake was measured, periodic body weight changes noted
and rumen contents and blood samples obtained.
Rumen contents were analyzed
for total volatile fatty acid (VFA) concentrations, molar proportions of
acetate to propionate, pH and protozoa concentrations.
Blood was analyzed
for serum protein and urea. These parameters may reflect physiological
changes occurring in wild deer prior to a period of complete starvation and
affect deer ability to favorably respond when suddenly offered supplemental
feed.
Total Starvation

Trials

Two pilot trials were
hensive trials.

conducted

to provide

direcf\on

for later, more compre-

The first trial investigated effects of starvation on deer rumen bacteria.
At intervals of one, two, three, and four weeks of starvation one deer was
sacrificed.
Viable bacterial numbers and ability of rumen bacteria to digest
two selected substrates (ground alfalfa leaves and ground concentrate ration)
in vitro from these animals were compared with those of a non-starved deer
in excellent condition.
Viable bacteria numbers were obtained following
anaerobic techniques of Hungate (1966). Digestive ability was obtained

�- 259 -

according to procedures outlined by Tilley and Terry (1963). The second
trial attempted to establish the "critical" point during starvation after
which deer cannot recover even if supplemental feed is offered.
Groups
of three deer were starved for la, 15 and 22 days and subsequently fed
a supplemental ration (Table 2) ad libitum.

Alfalfa

Feeding Trial

Preliminary investigations were conducted into the apparent inability of
deer to digest alfalfa or native hay when offered as emergency supplements.
Two physical forms of poor quality alfalfa, pelleted and cubed, were fed
to eight mule deer and four domestic sheep and goats.
Comparisons among
these animals were made in: (a) ability to digest the rations fed; and
(b) relative sizes of digestive organs.

RESULTS AND DISCUSSION

Malnutrition

Trial

During the trial all deer increased daily food intake (Fig. 1) and
exhibited only minor weight changes (Fig. 2). Fawns fed dehydrated alfalfa
were more susceptible to digestive upsets than were adults; within 2 weeks
one fawn died and two had lost 12 and 17 percent of their pre-trial body
weights.
Rumen-reticular pH decreased during the trial but increased when
alfalfa was fed, becoming alkaline in most deer (Table 3). Rumen-reticular
total volatile fatty acids (VFA) concentrations were 58-74 mM/l during the
trial and increased to 92-111 mM/l when alfalfa was fed. Corresponding
molar proportions of acetic to propionic acids were 4.8:5.8 and 3.7:4.3
respectively (Table 3). Rumen-reticular protozoa numbers decreased during
the trial and were significantly reduced or completely eliminated in rumen
contents of deer fed alfalfa (Fig. 3). Serum protein and urea levels were
within the range of "normal" values in adult deer. Serum urea was significantly higher (30-60 mg/IOO ml) in fawns fed dehydrated alfalfa (Table
4) •

Based upon phsyical appearance and weight change data, the formulated diet
fed during this trial did not simulate natural malnutrition and appears to
have provided minimum nutritional requirements for maintenance.
It is
unlikely that natural forages with nutrient levels similar to the "malnutrition" diet would provide minimum nutritional requirements for deer. Unlike
natural forages, the diet fed was ground into small particles (3 mm or less),
a factor that possibly permitted increased dry matter digestion and faster
rate of food passage.
Digestive upsets, death and moribund conditions in fawns fed dehydrated
alfalfa hay are unexplainable at present and warrant further study.

�Table 1. Chemical composition (oven dry basis) of the compounded ration and dehydrated alfalfa
hay cubes fed during Trial I.

Diet

Crude
Protein

Ether
Extract

Chemical Analysis (Percent)
Cell Wall
Moisture
Ash
Constituents

ADF 1/ Lignin

-Compounded ration
(Diet III) 1/

5.5

4.9

8.5

5.2

74.3

52.8

15.1

Dehydrated alfalfa cubes

15.9

4.0

7.2

7.6

53.0

38.6

6.8

1/ Composition -- 75 percent cotton seed hulls, 15 percent wheat straw and 5 percent each of wheat
bran and 32 percent protein supplement.
2/ Acid-detergent fiber.

I

N
0'1

0

�- 261 -

Table 2.

Supplemental

diet fed to totally starved

Component

Diet Composition

Barley

25%

Corn

5%

Wheat bran

10%

Milo

5%

32% supplement

20%

Cottonseed

35%

Expected

deer.

hulls

Proximate

Analysis

Crude protein

15%

Crude fiber

30%

Nitrogen

50%

free extract

�Table 3. Average rumino-reticu1ar fluid pH, VFA concentration (mM/1) and molar proportions of acetic:
propionic acids (C2:C3) in adult and fawn mule deer during Experiment 1 (observed ranges in parentheses).
Week of
Trial

N

pH

Adults
VFA Cone.

VFA C2:C3

pt 1/

N

pH

Fawns
VFA Cone.

VFAC2:C3

4

6.4 (6.3-6.5) 39 (31-60)

3.4 (2.7-3.8)

2

5

6.7 (6.4-6.9)

66 (44-101) 4.5 (3.7-5.1)

4

6.8 (6.5-6.9) 62 (45-74)

5.5 (5.4-5.7)

4

5

6.7 (6.5-6.9)

74 (55-87)

4

6.6 (6.6-6.7) 70 (50-79)

5.2 (5.1-5.4)

4

6.6 (6.3-6.9) 73 (52-84)

5.4 (5.0-5.8)

5.1 (4.0-6.2)

6
8

4

6.6 (6.5-6.9) 69 (64-74)

5.1 (4.7-5.4)

4

6.3 (6.1-6.6) 58 (49-75)

5.4 (5.1-5.7)

10

4

6.6 (6.3-7.1)

69 (61-81)

4.9 (4.3-5.9)

4

6.2 (6.2-6.4) 67 (57-76)

5.6 (5.3-6.0)

12

3

6.7 (6.3-7.2)

71 (64-75)

5.2 (4.6-6.4)

3

6.2 (6.2)

71 (67-75)

5.8 (5.4-6.4)

13

3

6.7 (6.1-7.2)

64 (50-76)

4.8 (4.6-5.5)

4

6.2 (6.0-6.4) 71 (53-82)

5.8 (5.5-6.4)

15 J)

3

7.1 (6.8-7.3) 103 (98-111) 3.7 (3.5-3.8)

4

7.0 (6.8-7.1) 94 (91-103) 3.8 (3.5-4.3)

1/ Pre-trial, fed concentrate diet ad libitum.
1/ Fed dehydrated alfalfa cubes ad libitum.

N

'"

N

�Table 4. Average
Trial I (observed

Week of
Trial

serum urea (mg/IOO ml) and protein
ranges in parentheses).

(g/IOO ml) in adult and fawn mule. deer during

Fawns

Adults
N

Serum Urea

Serum Protein

pt 1./

N

Serum Urea

Serum Protein

3

21. 6 (15-35)

7.9 0.2-8.5)

-----------------------------------~----------------------------------------------------------------2

3

6.6 (5-10)

8. 4 (8. 3- 8. 5)

3

21. 6 (20- 30)

3

20

6.6

(6.1-7.2)

4
6

8

3

21.6 (20-25)

(15-25)

7.1 (6.6-8.2)
N
0\
W

7.5 0.0-8.5)

10

4

21. 2 (20-25)

6.5 (6.0-7.0)

12
l3

3

21.6 (20-25)

7. 1 (6. 4- 7. 8)

4

21. 2 (20-25)

6.4

15 J./

3

30.0 (25-35)

8.1 (8.0-8.5)

4

42.5 (30-60)

6. 7 ( 5 . 3- 8. 3)

1/ Pre-trial, fed concentrate diet ad libitum.
1/ Fed dehydrated alfalfa cubes ad libitum.

(6.0-7.0)

�- 264 -

'Cone. t-----'------

Diet

III -------I&gt;I-4-Alf.+

2500

I

I
I

/X__*
. ".

/x

2000

:

~

.

x.-x (3)

I

~

~1500

(4)

CIS

•••
-tIO

I

0..,...,.0-0--0.

~)(-x

/"

/

/

,-)(

X

:
, /
I

K

x( 3)

x

~

o

C)

~

.•..
CIS

"

.5 1000

wtv

~x~

I

"

"I

I
500

I

-"-

X;-XAdults
0-0

Fawns

o~~--~~~~~~~~-L--L-~~~~~--~-L~
pt

1

2

3

4

5

6

7

8

9

10

11 12

13 14

15

Weeks

Fii;. 1.
:::ean dai.Ly food intake of concentrate
(p t = pretrial),
Diet
IL~ and dehydrated
alfalfa
cubes by adu Lz an c f avn muLe deer during
'lrial
I.
Figures
in parentheses
indicate
numbers of animals remaining
en trial.

�- 265 -

Diet

III

-----~I&gt;;t-rt-AII.-+

..

1

-

~ (-)1

s::

I
I

~
..r::

o (-)2

)C (3)

--d&gt;

0-

~
...•
~

~~

(-)3

/~

(- )4

I

I

.

X-XAdu1tsl

o

(-)5

I
I

0-0

I

\

lj4)

Fawnsl

12.6

o~-L--~~--~~--J-~--J-~--J-~--~~~~~
1

2

3

4

5

6

7

8

9

10

11

12

13 14 15

Weeks

Fig. 2.
of adult
indicate

Cumulative weight changes (percent
of initial
and f awn mule deer during Trial
1.
Figures
numbers of animals remaining on trial.

body weLght )
in parentheses

�- 266 -

600

500

s-

.~

G)

400

Pc

...,
0
...•
~

•

0

-..•
~

300

s::

0

.•..
CIf
~
.•..
s::

o

~o~

.

G)

o

s::

200

0

0

Fig. 3. Protozoa concentrations in rumino-reticular fluid of adult
and fawn mule deer fed concentrate Cpt = pretrial) niet III and
dehydrated alfalfa cubes during Trial I. Figures in parentheses
indicate numbers c f animals remaining on trial.

�- 267 -

Total Starvation

Trials

Viable bacterial numbers and digestion coefficients from starved deer
are presented in Table 5. These data suggest that numbers of viable
bacteria and digestive capabilities of these bacteria are not seriously
reduced when deer are totally starved.
Two deer, starved 22 and 23
days, died with numbers of viable bacteria in their rumens not greatly
different from those of non-starved deer.
Fates of deer starved and refed are ~resented in Table 6. All deer
that died during this trial were necropsied.
Rumen fluid parameters
measured were recorded for two deer from this trial: one died after 23
days of starvation and the other died after being starved 22 days and
then fed the supplemental ration for two days. All deer starved and
refed survived except one adult doe starved 22 days which died after
two days of refeeding.
At death this animal had the greatest number
of viable bacteria in its rumen of all deer examined (Table 5).
The data presented in Table 6 should be interpreted with caution.
These
animals were captured at Little Hills Experiment Station of the Colorado
Game, Fish and Parks Division, kept there in pastures for several weeks,
then transported 200 miles to Fort Collins.
At the time of their arrival
they were in poor, possibly semi-starved condition.
It seems that, at
least for deer in poor condition, three weeks of starvation may be close
to the point where supplemental feeding is no longer effective in preventing death.
Pregnant does and fawns succumbed much sooner than adult males to total
starvation.
Does dying of starvation carried healthy appearing fawns;
does surviving starvation with refeeding bore healthy appearing fa,vns.
Likely hypotheses resulting from these trials are: (a) starving deer
retain adequate numbers of viable bacteria in their rumens; and (b)
mortality of starved, refed deer is not due to non-functioning or absent
rumen bacteria but rather to failure of deer metabolic processes.
Future
trials will be mainly concerned with investigating these deer-oriented
failures.
Although an estimate has been made of the "critical" point during
starvation, after which supplemental feeding fails to prevent death, more
research is required investigating variables that may affect this point.
Physiological conditions associated with this point should be understood
and methods of determining when deer are at the point must be achieved.

�Table 5.

Rumen fluid parameters from starved deer.

0

7

14

Per ml rumen fluid

2.lxl07

4.4x166

4.lxl05

Total number per rumen

1.2xlOll

1.3xl010

Alfalfa

0.69

Concentrate

Da~s Starved 1/
21

22

23

28

----

3.8xl09

2.5x107

4.9xlO 5

1.6xl09

----

4.8xlO13

1.6xl010

9.8xlO

0.65

----

0.74

----

----

0.66

0.76

0.71

----

0.63

----

----

0.58

Rumen volume contents (ml)

5535

3050

3840

2210

1240

650

2000

Rumen dry matter contents (g)

235

86

263

25

376

527

50

Viable bacteria counts
e

9

Digestion coefficients

1/ One deer sacrificed at each date.

N
0\
00

�Table 6.

Fate of starved deer.
Days Starved

Status

Adult

female

6

Died 6th day of starvation.
before dying.

Appeared

malnourished.

Adult

female

10

Died 10th day of starvation.
normal appearing fawns.

Appeared

slightly malnourished.

Adult

females

10

Fed supplemental

male

11

Died 11th day of starvation.

Appeared

malnourished.

female

12

Died 12th day of starvation.
appearing fawns.

Appeared

malnourished.

Carried

12

Died 12th day of starvation.

Appeared

malnourished.

Infested with lice.

13

Died 13th day of starvation.
normal appearing fawns.

Both obviously

Yearling
Adult

Yearling
Adult

(3)

male

females

(2)

feed.

Each gave birth to healthy

Passed 2 dead fawns

Carried

2

twins.

malnourished,

2 normal

both carried

2
N
0\
1.0

male

14

Died 14th day of starvation.

female

15

Fed supplemental

feed. Gave birth to 1 healthy

15

Fed supplemental

feed.

16

Died 16th day of starvation.
appearing fawns.

22

Fed supplemental feed.
One died after 2 days on feed. Grossly malnourished and carrying 2
normal appearing fawns.
Two survived.
Both appeared grossly malnourished, one bore a healthy
fawn, the other bore none.

male

23

Died 23rd day of starvation.

Grossly malnourished.

female

27

Died 27th day of starvation.
appearing fawn.

Grossly malnourished.

28

Died 28th day of starvation.

Yearling
Adult

Adult males

(2)

Adult

female

Adult

females

Yearling
Adult

Adult male

(3)

Appeared

malnourished.
fawn.

Grossly malnourished.

Carried

Carried

2 normal

1 normal

�- 270 -

Table 7. Mean body weights and weight, volume, percent dry matter and
pH of rumino-reticular contents of mule deer and domestic sheep and
goats fed cubed alfalfa stems (number of animals in parentheses).

Mean
Body
Wt. (kg)

Animal

Sheep
Goats

(4)
(4)

Deer
normal digestion &gt;2)
omasum impacted 1 (2)
abomasum upset 11 (2)

Rumino-reticular Contents
Weight
Dry
(percent
Vol.
Matter
body wt.)
(1)
(percent) pH

51. 7
37.2

19.0
18.5

13.3
7.6

13.3
14.6

6.4
6.6

54.5
60.8
59.0

10.6
12.8
10.4

9.9
12.9
7.1

14.1
2.7
6.1

6.2
7.1
7.1

1/ Deer did not eat for 3-8 days prior to sacrifice;

rumen contents

very fluid.

•

Table 8. Mean size of the omasums and weight, percent dry matter and
pH of omasal contents of mule deer and domestic sheep and goats fed
cubed alfalfa stems (number of animals in parentheses).

Omasum Size
(with conte~ts)

Animal

Sheep
Goats

(4)
(4)

Deer
normal digestion &gt;2)
omasum impacted 1 (2)
abomasum upset 11 (2)

Vol. Cir. L.
W.
(ml) (mm) (mm) (mm)

Omasal Contents
Dry
Wt. Matter
(g) (percent) pH

861
546

271
297

189
155

125
101

757
530

17.5
19.8

7.2
6.2

239
1132
126

154
350
138

88
205
100

50
109
49

314
980
107

20.6
18.6
13.0

6.1
7.0
6.8

11 Deer did not eat for 3-8 days prior to sacrifice.

�Table 9. Mean size of the abomasums and weight, percent dry matter, and pH of abomasal contents
deer and domestic sheep and goats fed cubed alfalfa stems (numbers,of animals in parentheses).

Abomasum

Animal

Vol.
(mm)

Cir.
(mm)

Size ~with contents2
Inside Dia.
Pyloric
W.
Sphincter
L.
(mm)
(mm)
(mm)

Abomasal

Wt.
(g)

of mule

Contents

Dry
Matter
(percent)

pH
N
--.J

I-'

Sheep
Goats

(4)
(4)

Deer
normal digestion 52)
omasum impacted 1 (2)
abomasum upset 1/ (2)

1570
843

280
260

437
338

l34
121

61
52

1134
759

12.7
l3.3

3.5
3.4

448
215
661

206
174
222

267
244
277

82
71
95

48
44
52

521
212
633

12.4
15.7
17.0

3.3
5.8
6.5

1/ Deer did not eat for 3-8 days prior

to sacrifice.

�GOAT

SHEEP

DEER

NORMAL

DEER
OMASUM
IMPACTED

Or
10

I

20
30
(/)

ex:
N

~ 40

"'-I

w

N

z

;::
50
z
','

u

60
70
80
90

Figure 4.

Relative sizes of goat, sheep and deer stomach complexes

(after a photo by Nagy).

�GOAT

SHEEP

DEER
NORMAL

DEER
OMASUM

IMPACiED

:~
r

12

16 -

24·
N

~
(/)

0::

LV

30

W
!-

w

::E 36
!Z
J..J

&lt;&gt;

42
48

54
j
I

I

60
66

Figure 5.
72

Relative sizes of goat, sheep and deer omasums and abomasums

(after a photo by Nagy).

�- 274 -

Alfalfa

Feeding Trials

...

Mule deer fed either pelleted or cubed poor-quality alfalfa (7.9 percent crude protein, 44.2 acid detergent fiber, 9.3 percent lignin and
56.1 percent cell wall constituents) during two feeding trials experienced
varying degrees of digestive orders.
Four domestic sheep and four goats
fed these rations did not exhibit symptoms of digestive disorders.
Two
of eight deer fed pelleted alfalfa and four of eight deer fed cubed
alfalfa suffered severe digestive disorders and were sacrificed.
Necropsy
revealed varying degrees of hemorrhagic ulceration in the abomasum and
small intestine of all deer. Two deer, fed cubed alfalfa, also had
greatly distended omasums due to food impaction.
Figures 4 and 5 show
relative sizes of the stomach complexes of goat, sheep, healthy deer
and deer after developing impaction.
Comparisons among rumen-reticulums,
omasums and abomasums of deer, domestic sheep and goats indicated significant size differences (Tables 7-9). Related to body size, deer have
smaller rumen-reticulums,
omasums and abomasums than either sheep or
goats.
Deer stomachs do not appear morphologically adapted for digesting
high-roughage alfalfa hay diets.

LITERATURE
Hungate, R. E.
New York.

CITED

1966. The rumen and its microbes.
533 p .

Academic

Press,

Nagy, J. G. 1970. Middle Park Cooperative Deer Study - physiology
and prevention of deer starvation.
Colo. Div. Game, Fish and
Parks.
Game Res. Div. Fed. Aid Proj. W-38-R-24.
Game Res.
Rept., July, Part III. pp. 397-402.
Silver, H., J. B. Holter, N. F. Colovos, and H. H. Hayes.
1971.
Effect of falling temperature on heat production in fasting
white-tailed deer. J. Wildl. Mgmt.
35(1):37-46.
Tilley, J. M. A., and R. A. Terry.
1963. A two-stage technique for
in vitro digestion of forage crops. J. Brit. Grassland Soc.
18: 104-111.

Prepared

by __~/~~~/~~~
__~~~~?~.~~
..
~~~~
Julius G. Nagy /j [I
A~sistant Professor,

_

Wildlife

Biology

�- 275 July, 1971

JOB FINAL REPORT

State of

COLORADO
----------~~~~-----------

Project No.
Work

Plan No.

Deer-Elk

W-38-R-25

1

Job No.

15

Job Title:

Investigations

Deer-Auto

Accident

Investigations

Period Covered:

January

15, 1968 through March 31, 1971

Personnel:

Dale F. Reed, Dennis L. Money, Kenneth C. Dillinger,
William B. Zimmerman, Gary T. Myers, Bruce C. Giunta,
Jerome J. Cebula, Charles L. Merrell, Thomas M. Pojar.

ABSTRACT
A total of 727 deer-vehicle collisions were documented on two twenty-mile
segments of highway in western Colorado during 1968, 1969 and 1970. The
number of deer frequenting the area within approximately ~ mile on either
side of the highways. per month was estimated by weekly roadside spotlight
counts.
The three year total of the monthly mean counts on the Highway 13
study area was 5,046 deer and the total deer kill for the same period was
338 deer. The mean annual traffic volume was 243,322 vehicles.
On the
Highway 82 study area the three year 'total of the monthly mean counts was
2,569 deer with a total deer kill of 389. The mean annual traffic volume
was 1,291,825 vehicles.
The highest number of deer-vehicle accidents
occurred in October on Highway 13 and February and March on Highway 82.

��- 277 -

DEER-AUTO

ACCIDENT

INVESTIGATIONS

Thomas M. Pojar

The number of deer killed on Colorado highways has risen steadily since
1966 (Table 1). The problem of deer-vehicle collisions as a drain on
the wildlife resource and as a ha~-:.,'~d
to motorists (Fig. 1) has been
investigated in many states (Michigan, Allen 1969; several states,
Dickerson 1939; Kansas, Hlavachick unpublished; MDine, Howe 1964;
Nebraska, McClure 1951; Pennsylvania, Peek and Bellis 1969). Two areas
in Colorado where the number of deer-vehicle collisions were consistently
high were selected as study areas. An effort was made to determine the
circumstances related to deer-vehicle collisions.
In addition to information from the two intensive study areas, Colorado State Safety Patrol
reports on deer-vehicle collisions from the entire state for 1965-1969
were analyzed for factors related to such accidents.
This information
is tabulated in the Appendix.

P. S. OBJECTIVE
Determine factors which cause deer-auto accidents on Highway 13 south of
the White River bridge in Rio Blanco County and on Highway 82 between
Basalt and Glenwood Springs.

METHODS

AND MATERIALS

Number and Location

of Deer-Vehicle

Collisions

The number of vehicle-killed deer found in the study areas of Highways 13
and 82 was recorded.
Each study area was checked by vehicle for dead deer
at least three times a week during periods when the kill was high. During
summer months when the deer kill was very low the highways were usually
checked twice a week. Occasionally evidence of a deer-vehicle collision
was found (skidmarks, blood smears and deer hair) with the carcass missing.
These apparent kills were included in the dead deer tally. To further
facilitate the location of dead deer, the Highway Division and State Patrol
instructed their field men to leave all deer killed by vehicles at kill
sites.
State Patrolmen and the State Patrol Dispatcher cooperated by
notifying project personnel when investigating a deer-auto collision or
when a dead deer was seen along the highway.

�Table 1. Number of dead deer removed from Colorado highways by Wildlife Conservation Officers during the
period 1965-1970.
Month
July

August

Sept.

Oct.

Nov.

Dec.

Total

Year

Jan.

Feb.

Mar.

Apr.

May

June

1965

246

298

356

372

162

104

99

90

82

120

162

163

2,254

1966

163

126

143

105

112

120

99

93

106

194

213

220

1,694

1967

131

235

234

166

156

160

159

150

108

172

200

140

2,011

142

244

245

248

2,367

N

1968

190

216

283

214

179

142

156

108

1969

189

196

397

270

164

137

157

144

171

225

267

222

2,539

1970

270

206

299

284

222&gt;'&lt; 227

223

225

193

282

343

373

3,147

Totals

1,189

1,277

1,712

1,411

995

890

893

810

802

1,237

1,430

1,366

14,012

Avg.

198

213

285

235

166

148

149

135

134

206

238

228

Percent

8.5

9.1

12.2

10.1

7.1

6.4

6.4

5.8

5.7

8.8

10.2

9.7

* Data from NW Region not included.

100.0

'-l

o:

�N
-....I
\0

Fig. 1. Occasionally more than one deer is involved in a deer-vehicle collision, increasing the danger
to the motorist and adding to the drain on the resource.

�- 280 -

The locations of deer killed by vehicles were marked in relation to
reflective numbers placed at 1/4 mile intervals adjacent to highways
in both study areas. The Highway 82 study area was numbered 1 through
74 beginning at the south edge of Glenwood Springs and continuing to
the north edge of Basalt.
The Highway 13 study area was numbered 1
through 80 beginning at the Rio Blanco and Garfield County line and
continuing to the junction of Highways 13 and 64 near Meeker.

Deer Density Estimates
The number of deer frequenting the area within approximately 1/4 mile
on either side of the highway was estimated using the method described
by Reed (1969). The number of deer seen adjacent to each quarter mile
of highway was recorded.
One estimate was made each week on both highways
except when counting conditions were poor due to rain or snow. The estimates were discontinued on Highway 82 in the summer because of the very
low deer density in this area during this time of year.

Traffic Volume
Traffic volume was measured hourly with Streeter Amet electric recorders
activated by a vehicle passing over a magnetic loop buried in the roadbed.
Recorders and loops were provided and installed by the Colorado Division
of Highways.
Vehicle traffic volume was recorded hourly on each highway
since February, 1968.

Terrain Adjacent

to Study Areas

The area within 1/4 mile on each side of the Highway 13 and Highway 82
study areas was divided into about 40-50 acre fields on countour maps.
The highway side of each field corresponded to marked quarter mile intervals
of highway used to record the location of deer crossings, sightings, and
kills.
One diagonal line was drawn across each field. The number of 40
foot contour intervals intersected by each diagonal provided a quantitative
measure of roughness of terrain for that field. The terrain indices for
fields adjacent to the same length of highway were added to get a measure
of terrain roughness next to each quarter mile length of highway.
This
index was used to place areas into five general categories.
Fields having
indices from 1 to 10 were placed in the flat category, 11 to 20 - gentle
rolling, 21 to 30 - rolling, 31 to 40 - medium rough, 41 to 50 - rough,
and 51 to 60 - very rough.

DESCRIPTION

OF AREA

Highw~
This study area begins at the Rio Blanco-Garfield County line and continues
north to the junction of Highways 13 and 64 near Meeker.
The highway in
this area is two-lane with a speed limit of 60 mph for about 13 miles and 70
mph for the remaining 7 miles.
The highway surface is mixed bituminous material.

�- 281 -

The south end of the study area is about 7,350 feet elevation while the
north end is approximately 6,300 feet. Vegetation consists primarily
of sagebrush, browse, and pinyon-juniper types with some grassland and
a small amount of cultivated land mostly in small grain (Table 2).
Both cattle and sheep are grazed within the study area. In the general
vicinity of the study area the human population density is about 1.5
people per square mile (Baker 1970).

Table 2. Acreage and percent of various vegetative
1/4 mile of the Highway 13 study area.

types present within

Acres

Percent

Sagebrush

3,112

42.7

Browse

1,607

22.1

Pinyon-Juniper

722

9.9

Grassland

563

7.7

Cultivated

543

7.5

Other

740

10.1

Vegetative

Type

The area is covered with deep washes and steep hills.
The Grand Hogback,
a ridge as high as 1,500 feet above the valley floor, and as wide as three
miles, parallels the east side of the study area. The Little Hills Triangle
lies to the west of the study area with a maximum elevation of about 8,300
feet. Most of this area is usable winter range for deer and elk (Baker
1970). Highway 13 follows a valley that is approximately one-half to one
mile wide.
The bottomland adjacent to the highway is predominately big
sage, hayland and grain fields. During the three year study period (April
1968 - March 1971) the extreme temperatures were 900 F and -310 F and the
total annual precipitation ranged from 13.17 to 17.13 inches (Table 3).

Highway

82

The Highway 82 study area lies between Glenwood Springs and Basalt.
In 1968
when this study was started, there were about 5 miles of four-lane highway
from Glenwood Springs to the southeast.
By the end of June, 1969, this fourlane was extended to 12 miles.
The remaining portion of highway is two lane.
All of the highway has a mixed bituminous surface.
The speed limit is 60 mph
except on 3 miles of highway where it is reduced to 50 mph. The study area
lies between 5,746 feet elevation at Glenwood Springs and 6,624 feet elevation near Basalt.

�- 282 -

This study area contains a large amount of land that has been cleared
of native vegetation and cultivated at some time during recent years.
Native vegetation includes sagebrush and pinon-juniper types. Cultivated land is used primarily to raise alfalfa hay or to produce domestic
pasture for livestock (Table 4). The Roaring Fork River parallels the
southwest side of Highway 82 throughout the length of the study area.
Thus, most of the southwest side of the highway is relatively flat with
occasional draws.
The other side of the highway is mountainous.
Temperatures between April 1968 and March 1971, ranged from 980 F and
-200 F at a point in the study area about 7 miles southeast of Glenwood
Springs (Table 5). Total annual precipitation during the three year
study ranged from 9.75 inches to 14.74 inches.

RESULTS AND DISCUSSION

Deer Kill, Spotlight

Counts and Accident

Locations

Knowledge of when and where deer-vehicle accidents occur is essential to
any program aimed at reducing the frequency of deer-vehicle collisions.

Highway

13

During the three year study period (1968, 1969 and 1970) there was a total
of 338 known road-killed deer on Highway 13. This represents about 7 percent of the total number of deer observed near the highway during the spotlight counts for the same period (Figs. 2 and 3). Thirty percent of the
kill occurred during October for the highest kill of any single month.
This is three times the kill of any other single month.
The kill was mostly
concentrated in quarter mile sections 45 through 80 (Fig. 4). Deer distribution during spotlight counts related closely to the location of deer kills
(Fig. 5).

Highway

82

A total of 389 dead deer were found on the Highway 82 study area during
1968, 1969 and 1970. This represents about 15 percent of the total number
of deer observed near the highway during the spotlight counts (Figs. 6 and
7). Over 50 percent of the kills occurred during the months of February and
March.
The months of May through September accounted for only 7.5 percent
of the highway deer mortality.
The total live deer count for 1970 was down 37 percent from the average of
the total counts for the two previous years and the total kill was down 40
percent.
It is hypothesized the decrease in numbers of deer along the highway in 1970 was due to relatively mild weather during the critical winter
months.
Mild winter conditions resulting in lower snow depth probably leave
more forage available to the deer in the higher elevations.

�Table 3.

Climatological data collected near Highway 13.
April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Jan.

Feb.

March

65
5
35
1.70
2.90

74
6
44
0.95
3.30

85
22
57
0.20
3.54

85
29
62
1.09
4.83

78
31
58
2.78
2.78

80
24
51
1.33
3.94

68
13
43
1.37
3.82

50
2
28
1.67
3.56

40
-24
15
0.47
2.04

44
0
27
1.12
2.74

44
-4
23
0.71
1.16

57
-6
27
0.33
4.88

April 1968 to March 1969
Maximum Temperature (OF)
Minimum Temperature (OF)
Average Temperature (OF)
Total Precipitation (In.)
Ave. Wind Velocity (mph)
April 1969 to March 1970
Maximum Temperature (OF)
Minimum Temperature (OF)
Average Temperature (OF)
Total Precipitation (In.)
Ave. Wind Velocity (mph)

N
00

70
17
42
1.34
1.35

80
28
52
0.41
4.81

82
26
54
2.76
3.68

86
40
64
0.73
3.97

90
42
64
1.32
3.19

78
31
56
2.48
4.91

65
8
35
3.57
3.28

56
-13
29
1.20
3.57

60
8
40
2.07
3.4

78
20
56
0.43
3.8

87
22
62
1.15
3.5

86
34
71
1.65
2.7

90
42
68
0.29
2.5

77
18
57
2.14
3.2

75
10
43
2.28
3.0

53
5
37
0.69
3.1

48
-10
24
0.61
2.59

26
1.02
4.26

47
-1
27
0.40
3.26

53
-31
27
0.27
3.1

52
-11
29
0.90
3.8

65
-11

April 1970 to March 1971
Maximum Temperature (OF)
Minimum Temperature (OF)
Average Temperature (OF)
Total Precipitation (In.)
Ave. Wind Velocity (mph)

50
-11

28
0.46
3.3

49
3
27
1.29
3.19

w

�- 284 -

Table 4. Acreage and percent of various vegetative
within 1/4 mile of the Highway 82 study area. '

types present

Acres

Percent

Grassland

2,459

42.1

Pinyon-Juniper

1,367

23.4

742

12.7

397

6.8

Browse

350

6.0

Other

527

9.0

Vegetative

Type

Sagebrush
Broad1eaf

Trees

�Table 5.

Climatological data collected near Highway 82.
April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

Jan.

Feb.

March

71
18
42
0.79
4.22

84
24
51
0.63
4.18

93
34
62
0.15
3.91

92
33
67
1.51
2.44

86
33
63
1.32
2.57

88
30
57
0.30
2.76

83
18
51
0.93
2.75

59
5
35
1.01
2.37

57
-14
28
0.92
2.16

45
-10
21
1.33
2.45

45
-14
20
0.64
2.32

56
0
26
0.22
2.58

April 1968 to March 1969
Maximum Temperature (OF)
Minimum Temperature (OF)
Average Temperature (OF)
Total Precipitation (In.)
Ave. Wind Velocity (mph)
April 1969 to March 1970

N

co

Maximum Temperature (OF)
Minimum Temperature (OF)
Average Temperature (OF)
Total Precipitation (In.)
Ave. Wind Velocity (mph)

78
21
46
0.49
3.84

86
30
57
0.50
3.51

87
32
58
2.71
3.29

92
40
68
0.63
2.60

93
43
67
0.54
2.45

85
32
57
1.38
2.40

71
22
40
2.97
2.10

59
8
33
0.30
1.37

54
4
28
1.23
1.50

50
-12
26
0.51
1.90

56
10
34
0.60
2.40

60
10
34
1.02
3.30

70
14
33
1.92
3.1

85
23
49
0.04
1.5

96
30
52
2.74
3.6

98
38
64
1.28
4.1

94
41
65
2.07
4.3

84
24
51
1.12
2.3

80
16
38
1.68
2.4

59
12
34
1.14
4.8

53
-3
26
0.59
1.5

58
-20
27
0.59
3.2

60
0
29
1.06
3.2

73
5
35
0.51
3.7

April 1970 to March 1971
Maximum Temperature (OF)
Minimum Temperature (OF)
Average Temperature (OF)
Total Precipitation (In.)
Ave. Wind Velocity (mph)

V1

�ROAD KILLED
HIGHWAY 13

40

= 118
= 109
= II I
TOTAL = 338

1968.
1969 D
1970 ~

~ 35
...J
...J

-~ 30

DEER

TOTAL
TOTAL
TOTAL

3 YEAR

tv
00
0'\

u..

o
a::
lLJ

CD

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=&gt;
z

5

Fig.

JAN

FEB

MAR

APR

MAY

JUN

JULY

AUG

SEP

OCT

NOV

DEC

0.3"0

6.20/0

8.9%

5.3%

7.7%

3.6%

5.6 %

7.1%

7.4%

30.2%

10.7%

5.3%

area

by month

2.

The total

number

of deer

killed

on the

Hdghway 13 study

during

1965

1969,

and

1970.

�SPOTLIGHT COUNT
HIGHWAY 13
TOTAL = 1922
1968 •
1969 0 TOTAL = 1487
TOTAL = 1637
1970 ~

o

LLJ

&gt;
0::

LLJ

en

-

TOTAL = 5046

3 YEA R

m

o
0::

N

co

LLJ
LLJ

-.J

o
U.

o
0::
lJJ
m
~

::&gt;

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z
&lt;t

w

::E

Fig.

3.

JAN

FEB

MAR

APR

MAY

JUN

JUL

AUG

SEP

OCT

NOV

DEe

7.8Ofo

12.5Ofo 16.1%

14.0%

6.3%

I. 2 Cfo

2.6%

4.8"0

7.2%

8.8%

8.8%

9.9%

The mean number

of deer

counted

per

month

adjacent

to Hf.ghway 13 study

area

during

1968,

1969,

and

1970.

�NUMBER OF DEER KILLED
1968,

BY SECTION

13

- HIGHWAY

1969 AND 1970

20 ·
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I

I
I

I
1

••••

I

~

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,

"

':1

-I

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~

:.,j

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~
!Oil

.•...

.:..-

- -

,..

,..

,

!5

. ~

.•.:

,.;.

MILE

SECTION

NUMBER

The 1968, 1969, and 1970 total kill by quarter mile section of Highway 13 study area.

801
I
I

�366

Q

..

THREE

•

Q

LLI

~

HIGHWAY 13 SPOTLIGHT COUNT BY SECTION

300 ..

YEAR AVERAGE 1968, 1969, 1970

••

..

~ 250·

••
••

~

s 200...

~

0::
LLI
~

·•

I"'"

.•
.•

.•

~

150:

0::

..

L&amp;J

I"'"

•••
r-

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,...
I-

..

ii 100.'

r-

r-

I"'"

I •..

I

Z

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LU

2

•••

·
·

••••••

,..

·
50..

•••
I-

·
••

'0_
.' 4

Fig. 5.
area.

N
00
\0

.-

••••
I-

Ubhf~
,

28.

36

~

hm12

20

44

52
60
QUARTER MILE SECTION NUMBER

The three year mean of the number of deer observed

by quarter mile section

68

16

of the Highway

13 study

�46

45
ROAD KILLED
HIGHWAY 82

40
a 35

1968 III
1969 0
1970 ~

34

w
...J
...J

~ 30

TOTAL
TOTAL
TOTAL

3 YEAR TOTAL

.,,

a::
w

DEER

= 146
= 155
=

88

= 389
N
\.0

~ 25

o

LL.

21

o
a:: 20

w

16

CD

~ 15

z

to
5

FEB

262%

Fig. 6.

MAR
24.9%

APR

10.5%

MAY
1.0%

JUN
2.6%

AUG
0.5%

SEP
1.5%

OCT
7.7%

NOV
9.3%

3.6%

The total number of deer killed on the Higllway 82 study area by month during 1968, 1969, and 1970.

�C\J

0

v

400:
0

w

350

&gt;
a::
w

en

aJ

300

C\J

"-

C\J

0

0:::

w
w

0

I

3 YEAR TOTAL = 2569

.1

250

0
LL

SPOTLIGHT COUNT
HIGHWAY 82
1968 • TOTAL = 990
1969 0 TOTAL = 963
1970 fZJ TOTAL = 616

200

~II II~

I

I

N

a::
w

aJ

:E

150

:::&gt;

z
z

100

&lt;t

w

:E

50

Fig. 7,

JAN

FEB

MAR

APR

MAY

JUN

6.7%

24.2%

37.2 %

17.9%

2.6%

0.2%

The mean number of deer counted per month adjacent

0.2%

AUG

SEP

OCT

NOV

DEC

0.2%

0.6"0

2.80/0

3.5"0

3.8"0

to the Highway

82 study area for 1968, 1969, and 1970.

N
\0

t-'

�- 292 -

The general patterns of kill and count by quarter mile section do not
resemble each other as closely as expected (Fig's. 8 and 9). Section
numbers 6-9 show a disproportionately
large kill and sections 25-28
show a disproportionately
large count.
In sections 6-9 deer move from
good sagebrush and pinyon-juniper
cover on the east side of the highway,
across the highway to feed in hay fields and on hay stacks.
Therefore,
during severe winters when deer were forced into the Roaring Fork River
Valley for forage, there were numerous deer crossings in this area resulting in a greater number of deer being killed than in other areas.
In sections 25-28 a field of crested wheat grass (Agropyron desertorum)
is adjacent to the highway.
Crested wheatgrass is one of the earliest
grasses to green up in the spring and large numbers of deer have been
observed grazing in this field at that time. Deer were seldom observed
on the opposite side of the highway from the crested wheatgrass field.
This, along with track counts indicates that although many deer frequent
the area, few cross the highway.
This minimizes the road kill in proportion to the number of deer observed.

Traffic Volume
Traffic volume on Highway 82 increased at a rate of about 12 percent per
year during the three year study period.
Total traffic volume was 1.14,
1.29 and 1.44 million vehicles in 1968, 1969, and 1970, respectively.
Every month showed an increase over the same month of the previous year
(Fig. 10). Highway 13 traffic volume increased 1.7 percent from 1968 to
1969 and 5.4 percent from 1969 to 1970. The total traffic volume was 236,
240 and 253 thousand vehicles in 1968, 1969 and 1970 respectively.
Although the total traffic volume increased each year, it showed decreases
during some months (Fig. 11). Traffic volume on Highway 82 was over 5
times that on Highway 13. However, the general pattern by month was
similar with the months of July and August showing the heaviest traffic
on both highways (Fig. 12). October showed an increase over other fall
and winter months on Highway 13 probably because of hunter traffic during
the big game seasons.
March had a heavier traffic volume on Highway 82
than other fall and winter months.
This may be due to increased ski
traffic to the local ski areas.

Terrain

as Related

to Deer-Vehicle

Collisions

The terrain type on the Highway 82 study area did not appear to have a
significant effect on deer kill, with two exceptions.
Approximately 30
percent of the terrain was classed "Flat" but less than 15 percent of the
mortality occurred in these areas. On the other hand about 15 percent of
the area was rated "Rough", while almost 30 percent of the deer kills
occurred adjacent to spch terrain (Table 6).
Findings were similar on the Highway 13 study area where about 57 percent
of the deer kill occurred on rough and medium rough terrain -- which made
up less than 33 percent of the adjacent terrain (Table 7). These results
indicate that there may be some factor associated with the rougher terrain
that causes more deer-vehicle collisions to occur.

�39

NUMBER OF DEER KILLED

Q

,.

BY SECTION - HIGHWAY 82

1968 t 1969 AND 1910

~

25-

o
W

..J 20..J

~

a::
IJJ
IJJ

,..,

~

o 15u,

o
~-ex:
~
~

z

N

.

\0
W

1°1

III

~III

"h
5-1

,...
I"'T"I
P""

,..,

r-

P""

~

,...

I-

•••

r-P""

=

...

P""

I-

81

~

""

P""

4!

n

~

IIIIIII

rr

•.•...

12!

rrrn

161 20. 241 281 32' 361

11 Lr1-1

I-

~

..

nmh
40~ 44i 48: 52 56i 50: &amp;4~-.~ 12i 76~ 801
r

H

~

i

I

QUARTER
Fig. 8.

The combined

MILE

SECTION

NUMBER

1968, 1969, and 1970 deer kill by quarter mile section of Highway

82 study area.

�404
~

300-

"

THREE YEAR AVERAGE 1968,
1969, 1970

c
~

0::

HIGHWAY 82 SPOTLIGHT COUNT
BY SECTION

250-

IJJ

en
m
o 2000::
IJJ
IJJ

r-,

1
N
-0

C

u,

o

.j::-

150-

r-

r-

0::
IJJ

m
~

100r-

==&gt;
z

rr-

Z

&lt;t

r-

50-

I-

~

r-

25~

Fig. 9.
area.

t--,

hil-

I

I

4

12

r-

r-

I-

~n.

I'"

r-

r-

L...r

~ ~ lmI1 b, [rI

I

20
28
36
44
52
60
QUARTER MI LE SECTION NUMBER

The three year mean of the number of deer observed

by quarter mile section

Hf,~
68
of the Highway

82 study

�1968, 1969 AND 1970
2

,.

...JI

)

:J:

~

-

010

·

~

z
«
00

1

8
5
·
:l:
I-

.
~
~

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~

~
~

••

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\Jl

I

iG~

JAN

~
~
~
~

~
""
~

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~
~

~

~'"

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~
~
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~......"
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~

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f,

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.

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~

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c~

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:::

....:....
;;.
.;,.

~

~

~

~

~

~

1Ij• ..,..

~ •••
~

FEBMAR

~
"
~

~

~

~
~
~

r-

~

-

· ~~

-

\0

•••
I"'"

·..

N

~
~

~

u,

VOLUME

HIGHWAY 82
_
1968
roJJ 1969
c::J 1970

en
I.JJ

o

TRAFFIC

".

a

•..
~

~
~

...""::

...

"

~""
~"~
~"~",,
"::
~,,~

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,,~~

~

H

C"'~
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"

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.•.

,01

~

~

~

~

~

~

~

~ ••~

~
~

~

~

~

~
~

~

...
~

I"'"

"

"~~,,

~

APR MAY JUN

~
~

-

~
T

JUL AUG SEP

~

•

"

~

~

~

~

~

~

~

~
~
~

~

""

"

"~

:
""

..•
~

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~

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E:l •• ~

dlljilj

OCT NOV DEe

MONTH
Fig, 10,

The traffic volume by month for 1968, 1969, and 1970 on the Highway

82 study area.

�1968, 1969 AND 1970 TRAFFIC
CI)
LLJ

150

VOLUME

HIGHWAY13

--'o

_

1968

rzm

1969
1910

N

-

Z
LLJ

t:::J

&gt; 100

\.0
0'\

IL.

o
CI)

i&lt;i:

CI)
:::::&gt;

o
z

IJAN

Fig,

lL

FEB MAR APR MAY JUN JUL

The traffic

volume by month for

1968,

AUG SEP OCT NOV DEe

1969,

and 1970 on the Highway 13 study

area.

�THREE YEAR MEAN (1968,1969,1970)

TRAFFIC

VOLUME

200~------------------------------------~
III
o

fJ)

LLI
...J

HIGHWAY 13
HIGHWAY 82

16
144

o

-::I:

LAJ

115

&gt;

~

IVV

J

113

100
91

(/)

0

Z

~

:::&gt;

0

z

h

MONTH

Fig, 12,

The three year mean traffic volume by month on the Bighway 82 and Highway 13 study areas.

I

~
.~

-....J

�- 298 -

Table 6. Types of terrain within 1/4 mile of either side of the Highway 82 study area and the total 3-year road ki~l by terrain type.

Type of
Terrain

Terrain
Index

Quarte~ Mile Sections of Highway
No. of
No. of
Sections
Percent
Deer Killed

Percent

Flat

1-10

22

30.1

58

14.9

Gentle
Rolling

ll-20

17

23.3

120

30.8

Rolling

21-30

14

19.2

63

16.2

Medium
Rough

31-40

7

9.6

35

9.0

Rough

41-50

11

15.1

110

28.3

Very Rough

51-60

2

2.7

3

0.8

Table 7. Types of terrain within 1/4 mile of either side of the Highway 13 study area and the total 3-year road kill by terrain type.
Quarter Mile Sections of Highway
Type of
Terrain

Terrain
Index

No. of
Sections

Percent

No. of
Deer Killed

Percent

Flat

1-10

5

6.2

4

1.2

Gentle
Rolling

ll-20

23

28.8

55

16.3

Rolling

21-30

26

32.5

.84

24.9

Medium
Rough

31-40

17

21.3

134

39.6

Rough

41-50

9

11.2

61

18.0

�- 299 -

Sex and Age Distribution

of Deer Killed by Vehicles

The sex and age ratios of deer killed on each study area were tested
to see if there was a difference between study areas.
Since there was
no difference in these ratios (P&gt; .10) between study areas, the data
from both areas were combined (Table 8).'·
The sex ratios in neither the fawn nor yearling age classes differed
significantly from 50:50 (P&gt; .50 and P&gt; .25, respectively).
However, the
matures showed a significant (P &lt; .005) divergence from a 50: 50 sex ratio.
Bellis and Graves (1971) obtained essentially the same results in the sex
and age composition of road-killed white-tailed deer in Pennsylvania.
If it is assumed that the sex ratio in the fawn population is 50:50, it
can be concluded that there is no difference in the rate of deer killed
by vehicles between sexes in this age group.
It may be reasonable to
assume a 50:50 sex ratio in the fawn age class. The sex ratio of 164
fetuses examined was 89 males to 100 females (Table 9) which is not significantly different (P&gt;.25) from a 50:50 ratio. This close sex ratio plus
the fact that hunter selectivity is highly unlikely for or against either
sex would support the hypotheses that the fawn age class would have an even
sex ratio. However, differential natural mortality between sexes of this
age group is a possibility.
Taber and Dasmann (1954) found that blacktailed deer in Northern California have a mortality rate of 139 males to
100 females for deer under 18 months of age.
The deer killed by vehicles in the yearling age class show a ratio of 84
males to 100 females.
This age class would have gone through one hunting
season when males can be distinguished from females in the field which
should result in higher hunting mortality of the males.
If there was a
significantly greater natural and hunting mortality in males than in females
one would expect a somewhat lower ratio of males to females than 84:100.
A lower ratio of males to females in the population than in the road-killed
deer would indicate a higher susceptibility of male yearlings to deervehicle collisions than female yearlings.
Taber and Dasmann (1954) found
that 47 percent of yearling does accompanied adult does and that only 16
percent of the yearling bucks accompanied adult does. Most yearling bucks
traveled independently which would leave them without an experienced animal
for leadership.
Being with an experienced animal when crossing a highway
may be beneficial.
In the mature age class, the population of mature animals cannot reasonably
be assumed to be 50:50 because of hunter selectivity for mature males and/
or a higher rate of natural mortality for males.
Therefore, the divergence
seen in the sex ratio from 50:50 may be due to an actual difference in the
ratio in the population and/or a different behavior and movement pattern
for mature deer which makes the females more vulnerable or the males less
vulnerable to deer-vehicle collisions.

�Table 8.

The number of vehicle-killed deer by sex and age class from Highway 13 and 82 study areas.

Month

Combined Data from 1968 II, 1969, and 1970
Male
Female
Male
Female
Fawn
Fawn
Yearling &amp; Mature
Yearling &amp; Mature

Jan.

12

17

8

29

4

4

3

2

0

7

Feb.

33

29

10

43

23

22

2

3

5

24

Mar.

29

29

9

52

17

14

2

8

4

31

Apr.

18

9

2

25

11

6

0

2

1

10

May

2

11

5

8

2

9

1

0

3

6

Jun.

6

1

4

9

2

1

3

4

0

3

Combined Data from 1969 and 1970
Male Female
Male
Female
Male Female
Fawn
Fawn
Yearling Mature Mature
Yearling

0
0

Jul.

0

3

5

15

0

2

4

8

0

4

Aug.

4

4

5

10

1

3

3

5

1

2

Sep.

4

6

5

14

4

3

4

2

0

3

Oct.

26

25

12

35

17

13

8

7

2

17

Nov.

14

13

15

28

6

9

9

4

2

8

Dec.

8

II

3

10

4

6

2

4

0

3

Total

152

158

83

278

91

92

41

49

18

ll8

Males:
.Females

94:100

w

30:100

99:100

84:100

1/ The yearling age class was not distinguished from the mature age class during 1968.

15:100

�- 301 -

Table 9. Fetal rate of pregnant does and sex ratios of fetuses collected
from Highway 13 and 82 study areas and the general area of Glenwood
Springs during 1969, 1970 and 1971. 1/

Year

No. of
Does

Fetal
Rate

No. of
Males

No. of
Females

Males:lOO
Females

1969

49

1.55

33

43

78

1970

29

1. 79

27

25

108

1971

24

1.54

17

19

89

Total/Average

102

1.62

77

87

89

February

and March.

1/ Includes

data only from January,

Physical

Condition

of Deer Killed by Vehicles

Although physical condition of deer may not be directly associated with
deer-vehicle collisions this data was collected.
The kidney fat index
was determined on 226 deer using the method described by Anderson and
Medin (1965) (Table 10). As would be expected, the low means occurred
in late winter and early spring, and the high means occurred in late fall
and early winter.
The individual high kidney fat index was 104.91 which
was recorded for a mature male killed in November, 1968. The individual
low index of 1.84 occurred in a female fawn in October, 1969.

CONCLUSIONS
The original study objective "to determine factors which cause deer-auto
accidents" has proven to be very difficult to attain.
The factors causing
automobiles and deer to collide are numerous and have complicated interactions.
However, three of the most obvious measureable parameters involved
(number and location of deer kill, deer count or densities, and traffic
volume) were investigated.
Deer density adjacent to the Highway 82 study area was significantly (P&lt;.Ol)
correlated (r2 = .79) with the number of deer killed.
Estimates of deer
density are important but are not sufficient to describe the cause and
effect relationship of deer-vehicle accidents.
The monthly traffic volume
showed a significant (P &lt;.01) negative correlation with the number of deer
killed per month.
This can be explained by the fact that deer are not
present adjacent to the highway during the summer months when traffic
volume is at its peak.

�- 302 -

Table 10. Kidney fat index of deer killed along the Highway 82 and Highwav
13 study areas during 1968,1969 and 1970.

Fawn

Female
Yearling

Mature

Fawn

Male
Yearling

January

x
n

2
6.31

1
23.15

3
25.49

3
15.37

2
8.79

February

x
n

11
6.70

3
21.03

20
20.44

7
6.31

2
10.06

.March

x
n

5
6.04

6
12.48

21
10.49

6
6.25

April

x
n

1
3.92

1
11.17

6
12.55

1
10.83

May

x
n

1
4.98

1
15.72

1
3.82

June

x
n

2
8.04

July

x
n

2
9.74

August

x
n

September

x
n

October

x
n
x

November

n
December

x

n

2
23.31

1
12.50

Mature

4
6.75
4
5.24

1
6.83
1
12.60
1
8.04

1
6.51

1
19.22

2
13.27
1
43.50

1
24.21

1
11.82

2
45.16

6
17.17

5
66.70

11
28.92

14
12.87

5
68.16

1
96.86

5
21.66

5
49.90

l3
38.52

7
13.14

3
54.93

1
104.91

8
15.23

3

37.95

4
27.18

4
14.57

1
30.30

�- 303 -

Neither the number of deer counted (P&gt;. 25) nor traffic volume (P &gt;.05)
were significantly related to the number of deer killed on the Highway
13 study area. This would indicate that entirely different situations
exist on the two study areas. Perhaps the relatively low traffic volume
on Highway 13 is below the threshold necessary to allow the patterns
seen in the Highway 82 data to be detected.
The traffic volume on Highway 82 was five times that on Highway 13.
There were approximately twice as many deer observed adjacent to Highway
13 as Highway 82, yet the number of deer killed on each highway was nearly
the same. If it is assumed the number of deer crossing the highway where
they could be struck by a vehicle is always proportional to the number of
deer observed adjacent to the highway, then it could be concluded that
increased traffic volume results in increased deer kill.
At this point, the parameters of deer movement in relation to the highway become important.
It is believed there are three basic types of
deer crossing areas that can be defined by gross examination of the monthly
frequency distribution of deer-vehicle collisions.
(1) Terminal wintering
area: The Roaring Fork River valley is an example of this type. The
frequency of deer-vehicle accidents is concentrated in the late winter
and early spring months (Fig. 6). In these areas the deer may cross the
highway several times in route to and from feeding and watering sites.
(2) Migration route area: In the Vail area, an interstate highway severs
the route of a migratory deer herd. The highway deer kill frequency peaks
in late spring and fall with minimal kill during the rest of the year.
(3) A combination of terminal wintering and migration route areas: The
Highway 13 study area falls into this general category.
The frequency
distribution of deer-vehicle accidents by month (Fig. 2) shows peaks in
late winter, late spring and fall. It is speculated the winter peak is
caused by deer using this locality as a terminal wintering area. The
spring and fall peaks may be caused by migrations of the Piceance deer
herd across Highway 13. The extremely high frequency of deer-vehicle
collisions in October may be due to a combination of factors.
First,
the fall migration occurs in October and November; second, the Colorado
big game season begins in October, which results in increased vehicle
traffic on Highway 13 and probably increased deer movement caused by
hunter harassment.
The procedures or devices needed to help alleviate deer-vehicle accident
problems varies with the type of problem di$cussed above.
The terminal
wintering problem can be mitigated by providing for the needs of deer on
one side of the highway, making it unnecessary for them to cross, or by
constructing a physical barrier to prevent the movement of deer across
the highway.
It must be determined that there is no migration through
the area before a physical barrier is constructed.
The migration route
problem must be solved by providing some means for the deer to cross the
highway safely.
This problem can be complicated by migrations that are
not restricted to relatively short segments of highway.
The difficulty
of alleviating the terminal wintering area and migration route problem is
the greatest.
Few methods, other than education of the motorist, are
feasible in terms of cost effectiveness for reducing deer-vehicle accidents
in terminal wintering and migration areas involving long segments of
highway.

�- 304 -

Other jobs under Work Plan 15 emphasize the design and evaluation of
various devices and procedures to deal with the above mentioned problem
areas.

LITERATURE

CITED

Allen, Ross E. 1969. A study of deer-car accidents
M. S. Thesis.
U. of Michigan.
66 p.

in southern Michigan.

Anderson, A. E., and D. E. Medin.
1965. Two condition indices of the
Cache la Poudre mule deer herd and their application to management.
Game Information Leaflet No. 23. Colo. Game, Fish and Parks Dept.
3 p.
Baker, Bertram D. 1970. Survey, inventory, and analysis of deer and elk
winter ranges. Appendix B. Big game winter range analysis, Game Unit
22-Piceance.
pp. 31-58. In Game Res. Rep. July, Part I. Colo.
Game, Fish and Parks Div. 455 p. (processed).
Bellis, E. D., and H. B. Graves.
1971. Deer mortality on a Pennsylvania
interstate highway.
J. Wildl. Mgmt.
35(2):232-237.
Dickerson, L. M. 1939. The problem of wildlife
traffic.
J. Wildl. Mgmt.
3(2):104-116.
Howe, W. Sidney.
1964.
Fed. Aid W-37-R-13.

Experimentation
3 p.

destruction

with reflecting

by automobile

devices.

Maine

McClure, H. Elliot.
1951. An analysis of animal victims on Nebraska's
highways.
J. Wildl. Mgmt.
15(4):410-420.
Peek, Frank W., and Edward D. Bellis.
1969. Deer movements and behavior
along an interstate highway.
Highway Res. News.
36:36-42.
Reed, Dale F. 1969. Techniques for determining potentially critical deer
highway crossings.
Game Information Leaflet No. 73. Colo. Dept.
Natural Resources, Div. of Game, Fish and Parks.
3 p.
Taber, R. D., and R. F. Dasmann.
young Columbian black-tailed

1954.
deer.

,tkk
·
~t-6?~-)?/J lap.)

Prepared by Thomas M. Pojar
Wildlife Researcher

Candidate

A sex difference
J. Wildl. Mgmt.

in mortality in
18(3):309-315.

�- 305 -

A P PEN

D I X

�Table 11.

Number of vehicle accidents involving deer and reported to the Colorado State Patrol by month.
1968
Percent
No.

1969
No. Percent

Month

No.

1965
Percent

No.

1966
Percent

No.

1967
Percent

January

20

5.1

33

6.0

34

5.0

40

5.0

39

February

19

4.9

21

3.8

29

4.2

60

7.5

March

14

3.6

25

4.5

38

5.6

73

April

20

5.1

26

4.7

54

7.9

May

22

5.6

33

6.0

58

June

20

5.0

45

8.2

Total

Percent

5.5

166

5.3

43

6.1

172

5.5

9.2

67

9.5

217

6.9

77

9.7

53

7.5

230

7.4

8.5

58

7.3

44

6.2

215

6.9

69

10.1

49

6.1

56

7.9

239

7.6

69

8.7

70

9.9

323

10.3

w

July

36

9.2

69

12.5

79

11.6

August

38

9.7

40

7.3

53

7.8

37

4.6

45

6.4

213

6.8

September

25

6.4

43

7.8

40

5.8

57

7.2

45

6.4

210

6.7

October

44

11.3

67

12.2

81

11.9

92

11.5

71

10.0

355

11.3

November

70

17.9

96

17.4

96

14.0

104

13.0

97

13.7

463

14.9

December

63

16.1

53

9.6

52

7.6

81

10.2

77

10.9

326

10.4

Total

391

100.0

551

100.0

683

100.0

797

100.0

707

100.0

3,129

100.0

0

'"

�Table 12.

Number of vehicle accidents involving deer and reported to the Colorado State Patrol by day.

Day

No.

1965
Percent

No.

1966
Percent

No.

1967
Percent

No.

1968
Percent

No.

1969
Percent

Total

Percent

Sunday

81

20.7

107

19.4

126

18.4

133

16.7

124

17.6

57l

18.3

Monday

41

10.5

62

11.2

84

12.3

115

14.4

83

11.8

385

12.3

Tuesday

48

12.3

59

10.7

88

12.9

90

11.3

78

11.1

363

11.6

Wednesday

49

12.5

72

13.1

72

10.5

102

12.8

96

13.6

391

12.5

Thursday

46

11.8

55

10.0

92

13.5

76

9.5

83

11.8

352

11.2

Friday

60

15.3

93

16.9

III

16.3

130

16.4

130

18.4

524

16.8

Saturday

66

16.9

103

18.7

110

16.1

150

18.9

111

15.7

540

17.3

Total

391

100.0

551

100.0

683

100.0

796

100.0

705

100.0

3,126

100.0

w
0
'-l

�Table 13.

Time of reported deer-vehicle accidents in Colorado.

Hour by
Military
Time

No.

1965
Percent

No.

1966
Percent

No.

1967
Percent

No.

1968
Percent

No.

1969
Percent

Total

Percent

0100
0200
0300
0400
0500
0600
0700
0800
0900
1000
1100
1200
1300
1400
1500
1600
1700
1800
1900
2000
2100
2200
2300
2400

10
5
13
6
14
16
14
7
2
10
3
4
4
0
5
7
38
37
46
49
37
33
18
13

2.6
1.3
3.3
1.5
3.6
4.1
3.6
1.8
.5
2.5
.8
1.0
1.0
0
1.3
1.8
9.7
9.5
11.8
12.5
9.5
8.4
4.6
3.3

22
12
16
17
31
22
9
8
6
1l
4
2
5
3
6
6
50
86
79
51
32
32
20
20

4.0
2.2
2.9
3.1
5.6
4.0
1.6
1.5
1.1
2.0
.7
.4
.9
.5
1.1
1.1
9.1
15.7
14.4
9.3
5.8
5.8
3.6
3.6

27
17
9
28
27
34
15
13
8
3
4
4
3
4
6
9
57
86
106
99
50
30
28
15

4.0
2.5
1.3
4.1
4.0
5.0
2.2
2.0
1.1
.4
.6
.6
.4
.6
.9
1.3
8.4
12.6
15.5
14.5
7.3
4.4
4.1
2.2

25
24
19
17
27
40
20
14
16
8
4
9
2
6
6
1l
58
115
97
107
64
45
41
20

3.1
3.0
2.4
2.1
3.4
5.0
2.5
1.8
2.0
1.0
.4
1.0
.3
.8
.8
1.4
7.3
14.5
12.2
13.5
8.1
5.7
5.2
2.5

24
21
1l
10
31
32
22
12
12
1l
6
4
6
4
4
10
61
97
75
83
76
52
25
16

3.4
3.0
1.6
1.4
4.4
4.5
3.1
1.7
1.7
1.6
.8
.5
.8
.6
.6
1.4
8.7
13.8
10.6
11.8
10.8
7.4
3.5
2.3

108
79
68
78
130
144
80
54
44
43
21
23
20
17
27
43
264
421
403
389
259
192
132
84

3.5
2.5
2.2
2.5
4.2
4.6
2.6
1.7
1.4
1.4
~7
.6
.6
.5
.9
1.4
8.5
13.5
12.9
12.5
8.3
6.1
4.2
2.7

Total

391

100.0

550

100.0

682

100.0

795

100.0

705

100.0

3,123

100.0

w
0
co

�Table 14.
sunset.

Number and percent of reported deer-vehicle accidents in Colorado in relation to sunrise and

Deer-Vehicle Accidents
1967
1968
1966
No.
No.
Percent
No.
Percent
Percent

No.

Time Interval

No.

1965
Percent

I-Jithin3rd hour
before sunset

4

1.0

1

.2

9

1.3

8

1.0

5

Within 2nd hour
before sunset

5

1.3

10

1.8

10

1.5

11

1.4

At sunset or
w.Lt hf.n 1st hour
before sunset

11

2.8

17

3.1

27

4.0

38

4.8

1969
Percent

Total

Percent

.7

27

.9

6

.8

42

1.3

32

4.5

125

4.0
I

w

Within 1st hour
after sunset

58

14.9

120

21.9

168

24.5

200

25.3

164

23.2

710

22.8

Within 2nd hour
after sunset

74

19.0

88

16.0

120

17.6

115

14.5

127

18.0

524

16.8

Within 3rd hour
after sunset

51

13.0

42

7.6

51

7.5

55

6.9

68

9.6

267

8.5

Within 4 hour
after sunset

24

6.1

37

6.7

30

4.4

40

5.0

39

5.5

170

5.4

Other times of
night

75

19.2

96

17.5

115

16.8

148

18.7

106

15.0

540

17.3

0

--------------------------------------------------------------------------------------------------------------

1.0

�Table 14. Number and percent of reported deer-vehicle accidents in Co1orado"in relation to sunrise and
sunset ( continued).

No.

1966
Percent

No.

1967
Percent

No.

1968
Percent

No.

1969
Percent

Total

Percent

Time Interval

1965
Percerrt
No.

Within 3rd hour
before sunrise

7

1.8

14

2.5

19

2.8

18

2.3

11

1.6

69

2.2

Within 2nd hour
before sunrise

6

1.5

14

2.5

13

1.9

20

2.5

19

2.7

72

2.3

Sunrise or within
1st hour before
sunrise

14

3.6

37

6.7

33

4.8

44

5.5

42

6.0

170

5.4

w

I-'
0

Within 1st hour
after sunrise

17

4.3

17

3.1

28

4.1

24

3.0

16

2.3

102

3.3

Within 2nd hour
after sunrise

9

2.3

8

1.5

15

2.2

14

1.8

16

2.3

62

2.0

Within 3rd hour
after sunrise

7

1.8

6

1.1

12

1.8

15

1.9

17

2.4

57

1.8

All other times
of day

29

7.4

43

7.8

33

4.8

43

5.4

38

5.4

186

6.0

Total

391

100.0

550

100.0

683

100.0

793

100.0

706

100.0

3,123

100.0

�Table 15.

Number of Reported deer-vehicle accidents occuring in Colorado under various light conditions.
Deer-Vehicle Accidents
1967
1968
No.
Percent
No.
Percent

No.

1969
Percent

Total

Percent

17.8

122

17.9

123

16.0

117

17.2

536

17.5

9.8

60

8.8

87

11.3

57

8.4

301

9.8

Degree
of
Light

No.

1965
Percent

No.

1966
Percent

Daylight

76

19.4

98

Dusk or
Dawn

43

11.0

54

Dark Unlighted

w

t-'
t-'

265

67.8

393

71.3

494

72.3

544

71.0

504

74.3

2,200

71.6

Other

7

1.8

6

1.1

7

1.0

l3

1.7

1

.1

34

1.1

Total

391

100.0

551

100.0

683

100.0

767

100.0

679

100.0

3,071

100.0

�Table 16.

Localities in which 3,056 reported deer-vehicle accidents occurred in Co1Qrado.

Locality
of
Accident

1965
Percent
No.

1966
Percent
No.

No.

1967
Percent

No.

1968
Percent

1969
Percent
No.

Total

Percent

-Open
Country

164

42.1

221

40.1

273

40.0

277

36.1

195

29.3

1,130

37.0

Mountainous

154

39.4

221

40.1

280

41.0

365

47.6

296

44.5

1,316

43.0

Farming

68

17.4

94

17.1

117

17.1

112

14.6

157

23.9

548

18.0

Other

5

1.3

15

2.7

13

1.9

13

1.7

16

2.3

62

2.0

Total

391

100.0

551

100.0

683

100.0

767

100.0

664

100.0

3,056

100.0

w
,....
N

�Table 17.

Game crossing sign control associated with reported deer-vehicle accidents in Colorado.

Game
Crossing
Sign

No.

1965
Percent

No.

1966
Percent

No.

1967
Percent

No.

1968
Percent

No.

1969
Percent

Total

Percent

Present

13

11.0

28

23.7

53

45.0

22

18.6

2

1.7

118

100.0

Absent

9

12.3

19

26.0

28

38.4

17

23.3

0

.0

73

100.0

Unknown

368

12.8

501

17.4

598

20.7

722

25.1

693

24.0

2,882

100.0

Total

390

12.7

548

17.8

679

22.1

761

24.8

695

22.6

3,073

100.0

Vol
to-'
Vol

�- 314 -

Table 18. Speed factors associated with reported deer-vehicle accidents
in Colorado during 1965-1969.
Speed
Miles per
Hour

Estimated SEeed
Percent of
No. of
Accidents
Accidents

0-29

52

1.7

30-39

145

4.8

40-49

522

17.2

50-59

973

32.0

60-69

1,099

36.0

70-79

245

8.1

80-89

5

.2

Total

3,041

100.0

�Table 19.

Weather conditions associated with 3,066 reported deer-auto accidents in Colorado.

Weather
Condition

1965
Percent
No.

No.

1966
Percent

No.

1967
Percent

No.

1968
Percent

No.

1969
Percent

Total

Percent

Clear

283

73.2

455

83.0

467

68.5

533

69.7

651

95.0

2,389

77.8

Cloudy

86

22.2

80

14.6

18!+

27.0

190

24.8

0

0

540

17.6

I
W

Rain

7

1.8

6

1.1

15

2.2

18

2.4

20

2.9

66

2.2

Snow

9

2.3

5

.9

12

1.8

18

2.4

4

.6

48

1.6

Other

2

.5

2

.4

4

.6

5

.7

10

1.5

23

.8

Total

387

100.0

548

100.0

682

100.0

764

100.0

685

100.0

3,066

100.0

t-'

VI

I

�- 316 -

Table 20. Number of reported deer-vehicle accidents per mile of state
highway during 1965-1969 in ten Colorado counties where deer-vehicle
accidents were most concentrated.
Deer Kill
Per
Mile

Total Miles
of State
Highway

Total
Kill

La Plata
Garfield
Archuleta

149
165
94

317
249
132

Ouray

61

84

1.40
1.38

Chaffee

109

147

1.35

Clear Creek

95

83

.87

Douglas

130

112

.86

Rio Blanco

124

103

.83

Eagle

153

126

.82

Moffat

206

l38

.67

County

2.13
1.51

Table 21. Number of reported deer-vehicle collisions in ten counties whe re
most of 3,127 reported accidents occurred in Colorado during 1965-1969.

County

Total Kill

Percent of
Total Accidents

La Plata

317

10.1

Garfield

249

8.0

Chaffee

147

4.7

El Paso
Moffat
Archuleta

147
138
132

4.7
4.4
4.2

Eagle

126

4.0

Douglas

112

3.6

Rio Blanco

103

3.3

Gunnison

102

3.3

3,127

50.3

Total

�- 317 Table 22. Miles of state highway per annual average reported accident during
1965-1968 where the miles per annual reported accident is less than 5.000.

State Hwy.
No.

County

Length
(Miles)

82

Garfield

18

115

El Paso

10 (160) }j

Number of Deer-Vehicle
Accidents Reported
1965 1966 1967 1968

Total

Miles Per
Annual Average
Reported Accident

6

13

23

25

67

1.075

20

9

8

16

15

48

1.667

La Plata

52

14

34

36

40

124

1.677

82

Eagle

6

6

2

o

6

14

1. 714

19 (550)

La Plata

45

14

29

21

23

87

2.069

185 (1-25)

Douglas

12

7

5

7

4

23

2.087

15 (285)

Chaffee

29

14

9

12

20

55

2.109

6 (50)

Saguache

9

o

3

5

7

15

2.400

19 (550)

Ouray

37

14

11

16

20

61

2.426

10 (160)

Archuleta

44

9

11

17

31

68

2.588

13

Rio Blanco

41

12

13

18

18

61

2.689

1 (1-25)

El Paso

46

15

20

14

13

62

2.968

1 (85)

Douglas

37

8

14

17

10

49

3.020

10 (160)

Costilla

31

5

8

13

11

37

3.351

78 (6) ]j

Eagle

21

3

5

10

7

25

3.360

2 (40)

Clear Creek

33

6

12

11

8

37

3.568

4 (1-70)

Garfield

89

13

26

27

29

95

3.747

167

Crowley

1

1

o

o

4.000

Jefferson

3

2

1

3

4.000

149

Rio Grande

5

o
o

o
o

1

93

o

2

3

5

4.000

91 (1-70) 3/

Clear Creek

22

1

5

7

8

21

4.190

145

Montrose

17

4

6

2

4

16

4.250

13

Garfield

17

2

2

8

3

15

4.533

2 (40)

Routt

54

8

12

12

13

45

4.800

4 (24)

Chaffee

33

6

6

6

9

27

4.889

1/ Corresponding federal highway number.
2/ Dowd Junction east over Vail Pass.
3/ Georgetown west over Loveland Pass.

�__-1

-"--r------------------

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-e
c:::
cC
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t- --. -1"I~ G A

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-&lt;
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-;-.':.-

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,..J

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t-'
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r/J

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-c

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.. __ L_ .. __ ._ .. _._

J1 •••.•

•••-

NEW

________
..

I
,

"S

~

~.T·__·__··~·
10)-

j

IS·

~eported by the Colorado

Highway Patrol

in 1969.

~I!-)7·
I'·

IOKLAHOMA

MEXICO

Fig. 13. The location of 679 deer-vehicle accidents
black dot represents one deer-vehicle accident.

I

Each

�- 319 July, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RAD~~O~

Project No.

W-38-R-25

Work Plan No.

15

Job Title

_
Deer-Elk
Job No.

Investigation

Investigations
3

of One-Way Deer Structures

Period Covered:

April 1, 1970 through March 31, 1971

Personnel:

Steven F. Steinert, Gary T. Myers, William
Thomas M. Pojar and Dale F. Reed.

B. Zimmerman,

ABSTRACT
Several types of one-way gates that permit deer to move through deer-proof
fences from only one direction were tested under controlled conditions at
the Little Hills Experiment Station paddocks.
Tests indicated that over
90 percent of the deer placed in the experimental paddocks used two of the
one-way gates.
The total frequence~ of use (preference) on these two gates
were not significantly different (X =0.105,P&gt;0.50).
Positions used in
2
the deer proof fence did not show a significant difference in use (X =0.421,
P&gt;0.50).
One-way gates were also tested under field conditions adjacent
to Interstate 70 near Vail and adjacent to Highway 82 south of Glenwood
Springs.
A total of 152 passages have occurred through the eleven one-way
gates under the varying field conditions.
Ninety-three percent of all
field passages were positive (in the direction of the funneling or cone
principle).
In addition, two one-way fence structures did not meet the
one-way criteria during continued preliminary tests. Of a total of only
five crossings, three were negative.
They were rejected for further testing.

�- 320 -

RECOMMENDATIONS

1.

When deer-proof fencing adjacent to high speed highways is used·
to reduce or prevent deer-vehicle accidents, one-way structures
should be considered for installation.
Deer that move around
the ends of the fence, or from the side of the road not fenced,
will be able to pass through a one-way gate rather than be
trapped on the highway and likely killed.

�- 321 -

INVESTIGATION

OF ONE-WAY DEER STRUCTURES

Dale F. Reed

P. S. OBJECTIVE
Evaluate the effectiveness of several types of one-way gates and two
fence structures to permit one-way movement of mule deer through
existing deer-proof fences at Little Hills Experiment Station and
elsewhere.

SEGMENT OBJECTIVES
1.

Test the effectiveness

of each structure

under controlled

2.

Test those structures deemed effective under controlled
for effectiveness under field conditions.

conditions.

conditions

METHODS AND MATERIALS

Testing Under Controlled

One-way

Conditions

Gates

Two types of one-way gates were tested with deer placed in the Little Hills
paddocks.
Tests of preference (frequencies of use) were made with two gates
in several positions between two paddocks.
The location of each gate was
changed in order to eliminate the use of gates because of position.
With
two gates (I and IIIb) being tested in two possible positions, one sequence
was used (if A and C are the positions; and I and IIIb are the gates): AI,
Clllb, Alllb CI. Thus, the position of the gate was reduced or eliminated
as a possible bias and deer preference was tested.
Gate I was tested with
Gate II during the last segment.
It was felt that for a deer to prefer one gate over the other the animal
must have experienced the use of both types. Therefore, small groups of
deer or individual deer (Table 1) were oriented to each of the two gates
for a period of four days. Only one gate was used at a time and was changed
each day with the other so that the deer could experience one gate one day,
another the next, etc. Some deer were oriented in small groups rather than
individually to reduce the number of days required for the testing.

�- 322 -

Table 1. Group, Classification,
and IIIb preference tests.

and number of deer used during Gate I

Group

Does

Fawns

J

1

K

2

Bucks

Total

1

2
2

L

1

M

2

1

N

3
1

0

1

p

1

Q

6

1
1

1

Total

1

2

1

1

4

12

Once the group was oriented, all except one deer were returned to a holding
pen to await individual testing.
The deer to be tested was left in a paddock
'not containing food. Both gates were then placed in position between the
paddock not containing food and the paddock with hay and pellets, thus beginning the sequence.
Each deer was tested for 4 days with the sequence
being repeated (i.e. AI Clllb, Alllb CI, and AI ClIlb, Alllb CI). This
procedure was repeated until each deer of the group was tested.

One-way

Fences

The number of deer crossing one-way fences I and II (overhanging and slanting
respectively) was to be counted each day in a manner similar to that of the
one-way gates. Only preliminary tests were conducted and thus, procedures
for preference testing were not developed.

Testing Under Field Conditions
The one-way gates deemed effective under controlled conditions were to be
tested in the field.
Since there was not a significant difference between
the use of Gate I and IIIb under controlled conditions, both types were

�- 323 -

utilized during the field tests. Gate IIIb was basically the same as
Gate I except that the bottom was installed at ground level allowing
deer to walk through.
Gate I was installed approximately 12 inches off
of the ground, necessitating the animal to jump through as previously
reported.
All installation sites were suitable for the 12 inch above
ground level except one. This was a steep slope adjacent to Highway
82 where no level ground was available.
By utilizing Gate IIIb's characteristic at this location, the need for a seemingly unreasonable jump
was eliminated:
Three one-way gates were installed in a quarter mile
of eight foot fence adjacent to Highway 82. In addition, eight gates
were contracted for and later installed in the eight foot fences
associated with the deer underpass at Vail. Four of the eight one-way
gates were placed in off-sets (Fig. 1) in the south Vail eight foot
fence, while the remainder were strategically located at eight foot
fence junctions or areas where deer had frequented the fence on the
wrong side.

RESULTS AND DISCUSSION

Controlled

One-way

Conditions

Gates

One-way Gates I and IIIb were tested with 12 deer placed in the Little
Hills paddocks.
These deer of various sex and age classes were used in
testing for gate preference.
Gates I and IIIb were evaluated simultaneously with each of the 12 individual deer in orientation groups J through Q (Table 2). Eleven of
the 12 deer made a total of 20 passages through Gate I and a total of
18 positive and one negative through Gate IIIb (Fig. 2). The frequencies
of use (preference) were not significantly different (X2 = 0.105, P&gt;O.sO).
Hence, both gates were field tested. Positions A and C used in the deerproof fence did not show a significant difference in use (X2 = 0.421,
P&gt;O.sO).

One-way

Fences

Preliminary tests of the one-way fences were considered unsuccessful.
The
few positive crossings that occurred with the variable height posts in
their lowest positions (Fig. 3) were outnumbered by negative crossings
(Table 3); consequently, the one-way fences did not meet the previously
established one-way criteria.
Structures not meeting this criteria were
to be modified or rejected.
In this case, modifications did not appear
feasible and the structures were rejected for further testing.

�- 324 -

FENCE CONTINUES

~

I
X

~

ONE-HAv

DEE'

OATE

I
I
I

~,,-~

'~D---I

)&lt;.
nEER-PROO~

I

FENCE POST

I

't..

(HICm-IAY SIDE)

I

I
DEER-PROOF

I

FENCE

APPROXTI1ATEL Y

100'

I
I

'I.

I
I
I

I

I
'I.

o

~\
I

ONE-'~AY DEER GATE

~

Fig. 1. One of several
Vail eight foot fence.

one-way deer gate off-sets

located

in the south

�- 325 -

Fig. 2. Positive passage of deer
through one-way gate IIIb during
controlled conditions at the Little
Hills Experiment Station (Photo by
author) •

Fig. 3. One-way Fence II with
variable height posts in their
lowest positions (Photo by author).

�- 326 -

Table 2. Frequencies of use (preference) of one-way Gates I and IIIb and of
one-way gate positions.
Doe
No. Group

Buck
No. Group

A

C

M

.25

.75

.25

.75

1

J

0

.25

.75

.25

.75

2

.25

0

.25

.75

.50

.25

0

0

0

Gate
I IIb

A

C

1

J

.33

.67

.67

.33

1

2

K

.75

.25

.25

.75

2

3

K

.75

.25

.75

4

M

0

.25

5

M

0

6

P

0

Table 3.
and II.

POSe

Gate
I
lIb

POSe

Fawn
No. Group

Gate
I UIB

POSe
C

A

a

.50

.50

L

.75 .25

.75

.25

3

N

.50 .50

0

1.00

4

Q

.67 .33

.67

.33

1.00

Passages and days elapsed during preliminary tests with one-way Fences I

Days Elapsed
Without
POSe Passage

Days Elapsed
Without
Neg. Passage

Ht. of
Fence
(In.)

Length of
Angle
(In.)

Group or Deer

Crossings
POSe Neg.

Group A (5 deer)

0

5

46

70

Group B (2 deer)

0

7

46

70

Buck (2 pt. from
Group A)
Buck (3 Pt.)

-1

7

46

70

0

3

34

65

Buck (4 P't . )

1

0

1

4

34

65

Doe

1

-1

4

a

34

65

Doe (from Group M)

0

-1

3

a

34

65

4

34

65

34

65

a

Fawn
Fawn

0

Total

2

4

-3

24

18

�- 327 -

Field Conditions
Of eleven one-way gates installed under field conditions, eight were installed adjacent to Interstate 70 and the deer underpass located in the
Vail area. Gates designated 70N-4 through 70S-II (Table 4) were installed
in both the north and south eight foot fences.
(The first number in the
designation stands for the highway adjacent to which the gate is installed.
The letter N or S signifies whether the gate was installed in the eight
foot fence north (N) or south (S) of the highway, and the last number the
sequential number of the gate installed).
Some of the gates received many
more passages than others.
For instance, gate 70N-5 received 84 passages
while 70S-II received none.
This is understandable if the physical layout is considered.
Gate 70N-5 was located in a "V" junction near good deer
cover and approximately one quarter of a mile from the highway, and may have
been used by resident animals throughout the summer months.
On the other
hand, gate 70S-II, as well as gates 70S-8, 70S-9, and 70S-l0, was not installed until after the peak of the spring migration during 1970 when the
greatest number of passages could have been expected.
It was also located
in an off-set with gate 70S-l0 approximately 100 feet from the highway
roadbed.
In short, gates 70S-II and 70N-5 may not be comparable.

Table 4. The number of one-way gate passages, percent of positive passages,
and number of tracks on the negative side when no passages occurred during
field tests.

Gate Designation
and Type

Passages During
Field Tests
No.
No.
POSe
Neg.

82-1 (lIb)

2

0

82-2 (1)

2

82-3 (I)

Percent
Positive

No. of Tracks
on Neg. Side
When no Passage

Type Fence
Installation
and Comment

100

0

-1

67

0

Straight fence
line and trail
Off-set

2

0

100

0

Off-set

70N-4 (I)

44

-3

93

54

"V" junction

70N-5 (1)

84

-2

98

28*

"V" junction

70N-6 (I)

4

0

100

0*

Straight

70N-7 (1)

8

-3

73

78

Straight fence line
and cliff near underpass

70S-8 (I)

1

-1

50

159

Off-set

70S-9 (I)

2

0

100

124

Off-set

70S-l0 (I)

3

0

100

218

Off-set

70S-11

0

0

303

Off-set

152

-10

(I)

Total
* Inaccurate

93

since deer trails pass negative

964
end of gates.

fence line

�- 328 -

The effectiveness of the gates in terms of "one-way passages" is indicated by the few negative passages that occurred.
Most of the nine
negative passages-(against
the funneling or cone principle) according
to the size of the tracks, were caused by new born fawns resulting
from May and June parturition in the vicinity of gates 70N-4 and 70N-5.
In addition, the one negative passage for gate 70S-8 was caused by
trying to drive several deer in front of a vehicle at night with a spotlight. Observation of this negative passage indicated that it was not
easily negotiated despite the animal's running impact.
The gate was
visibly bent as a result _of this passage.
Another indicator of the
"one-way passage" characteristic was the number of tracks counted on
the negative side when neither positive or negative passages occurred.
A total of 964 tracks indicated activity on the negative sides of the
gates where negative passages did not occur; these gates were effective
in preventing deer from getting on the 4-lane Interstate Highway (Fig.

4).
One-way gates 82-1 through 82-3, located in a quarter mile fence adjacent to Highway 82 south of Glenwood Springs, received few passages.
The number of deer in the immediate area has been low during the last
two years (1969 and 1970).

~
Prepared

~~

bY~C{\~
Dale F. Reed
Asst. Wildlife

Researcher

�- 329 -

Fig. 4.
negative

Arrows point to several of the 964 tracks imprinted on the
sides of the one-way gates at Vail.
(Photo by Don Domenick)

��- 331 July, 1971

JOB FINAL REPORT

State of

COLORADO

Project No.

W-38-R-25

Work Plan No.

15

Deer-Elk

Investigations

4

Job No.

Job Title:

Evaluation

of Devices

to Prevent

Deer-Auto

Period Covered:

February

Personnel:

Gary T. Myers, Dale F. Reed, William
Thomas M. Pojar.

Accidents

1, 1968 through March 31, 1971
B. Zimmerman

and

ABSTRACT
Brush was removed from the borrow pits and roadside along two segments of
Highway 13, each one and three-quarter mile long south of Meeker, Colorado.
It was hypothesized that clearing of the roadsides would reduce the frequency
of deer-auto accidents on the highway because of increased visibility for
the motorists and reduced cover for wildlife.
An examination of the ratio
between the number of deer killed by deer-auto accidents per 100 deer counted
during roadside spotlight counts before and after the removal of roadside
brush does not provide sufficient evidence to support the hypothesis.

�- 332 -

RECOMMENDATIONS

1.

Evaluation of brush removal
on this project.

2.

If brush removal is to be evaluated again at a later date, the
following procedures are recommended.

3.

should be relegated

to low priority

a.

Obtain quantitative pretreatment data concerning height,
and species composition of the vegetation to be removed.

b.

Keep study area free of all forbs and other plants that may
obscure vision.

c.

Provide for quantitative
shrubby species.

d.

Provide a method for determining the number of deer crossing
the highway.
This is necessary so that successful and unsuccessful (deer killed) crossings can be compared for
pretreatment and treatment time periods.

evaluation

of a re-vegetation

density

by

Removal of roadside brush for the specific purpose of reducing the
number of deer-auto accidents cannot be recommended as a management
practice on the basis of this study.

�- 333 -

EVALUATION

OF DEVICES TO PREVENT DEER-AUTO

ACCIDENTS

Thomas M. Pojar

The first phase of the Deer-Auto Accident Study (Job 1) was concerned with
the gathering of data related to the circumstances of deer-auto accidents.
Such information as location, time and frequency of deer-auto accidents
was obtained.
The second phase of the Deer-Auto Accident Study is to
evaluate procedures and devices that will reduce the number of deer-auto
accidents.
The evaluation of fencing along Highway 82 was initially an
objective of this job, however, the fence was not installed during this
segment.
The fence evaluation has been made a separate Job (10) and will
be reported on for Segment 26. The remaining objective of this job is to
investigate the effect of roadside brush removal on the number of deerauto accidents.
Tall shrubby species growing in the borrow pits and along
the road obscure the vision of the motorist and provide cover for wildlife.
It was hypothesized that the clearing of roadsides would reduce the frequency
of deer-auto accidents.

P. S. OBJECTIVE
Determine if a fence built adjacent to a portion of Highway 82 and brush
removal in strategic locations adjacent to Highway 13, south of the White
River bridge in Rio Blanco County, affects the numbe~ of deer killed by
vehicles in the areas involved.

METHODS

AND MATERIALS

The time, date and location of all deer-auto accidents in the study area
were documented.
Roadside spotlight counts were conducted weekly using the
method described by Reed (1969). The spotlight counts were used as an index
to the population of deer frequenting the area within approximately onefourth mile on either side of the highway.
The brush (mostly big sage, Artemisia tridentata) was removed from the highway right-of-way by the Colorado Division of Highways.
A road grader was
used to remove sagebrush plants at ground level when the ground was frozen.
The pre- and post-treatment data were not subjected to statistical analysis.
Since the relationship between the kill and count has not been established,
a ratio between them is not a true proportion; therefore, the chi-square
test is inappropriate.
A t-test comparing mean ratios is inconclusive
because only one degree of freedom is available.

DESCRIPTION

OF AREA

Two segments of Highway 13, each one and three-quarter mile long, located
south of Meeker, Colorado, were selected as study areas.
Area I (quarter

�- 334 -

mile sections 65-71) and Area II (quarter mile sections 58-64) began 2.50
and 4.25 miles south of the junction between Highway 64 and Highway 13,
respectively.
The highway follows a valley approximately one-half to one
mile wide.
The Grand Hogback, a ridge up to 1,500 feet above the valley
floor and as wide as three miles, parallels the east side of the study area
(Myers 1969). The bottomland is predominantly Big sage and grain fields.
The east and west facing slopes adjacent to the study areas support browse
type vegetation as characterized by Baker (1970). The pinyon-juniper
community dominates the upper slopes and the mesas above the rim.
A dense stand of big sage occupied much of the highway right-of-way prior
to removal (Fig. 1). Appearance following brush removal is shown in Figure 2.
This general area is traversed by the migratory Piceance deer herd during
spring and fall migrations.
It is also used by mule deer as a wintering
area (Baker 1970).

RESULTS AND DISCUSSION
Pre-treatment data (roadside spotlight counts and number of deer killed by
deer-auto accidents) for Area I was collected from January 1, 1968 to
January 1, 1969. During January 1969 the brush was removed from both sides
of the road.
Post-treatment data collection began February 1, 1969 and was
terminated January 1, 1971.
Area II pre-treatment data was collected from January 1, 1968 to November
30, 1969 and post-treatment data from January 1, 1970 to January 1, 1971.
Brush removal from both sides of the road was completed during December 1969.
Data from the months when brush was being removed were disregarded
of distrubance caused by machinery.

because

In Area I the ratio of the number of deer killed per 100 deer counted
decreased slightly the first year after brush removal (Fig. 3). However, a
sharp decrease, from 2.16 to 1.31, occurred the second year after brush
removal (Table 1). If brush removal had a real effect on the ratio of deer
killed per deer observed, the data from 1969 should have shown a closer
relationship to data from 1970. There is no reason to believe brush removal
would have a latent effect on the kill/count ratio. Therefore, it seems
very unlikely that brush removal was an influencing factor on the kill/count
ratio in Area I.
The kill per 100 deer counted ratio for Area II varied from 2.18 to 2.99
(Table 2). This range was observed for the two pre-treatment years, which
would indicate a relatively wide range of ratios due to random variation or
other factors not measured.
There are several inherent problems that may lessen the effect of brush
removal as a method of reducing deer-auto accidents.
First, by clearing
the areas of shrubby species, the primary invaders are usually forbs.
The forbs may reach heights that obscure vision as effectively as the shrubby
species that were removed, unless measures are taken to control them.

�Table 1. Count and kill data for Area I of Highway 13, south of Meeker, Colorado, where roadside brush
was removed in an attempt to reduce ORp.r-auto collisions.

Month

No. of
Counts*

1968
Total
Count

Total
Kill

No. of
Counts

1969
Total
Count

Total
Kill

No. of
Counts

1970
Total
Count

Total
Kill

Jan.

2

37

0

**

**

**

4

39

0

Feb.

4

188

1

4

135

3

4

100

1

Mar.

4

314

5

4

290

1

4

143

1

Apr.

3

183

5

264

1

4

277

May

5

81

°
0

4

15

3

4

53

°1

June

4

7

July

4

Aug.

4

3

1

4

3

0

9

°1

5

3

0

5

17

0

5

10

1

4

19

0

4

16

1

Sept.

4

16

3

4

17

0

5

35

1

Oct.

5

84

10

5

43

8

3

15

2

Nov.

4

49

1

4

61

2

4

35

3

4

56

-2

5

28

3

31

0

48

1,034

24

48

878

°
19

48

764

10

Dec.
Total

-

Ki11/100 deer ccounted

2.32

2.16

* Count refers to a roadside spotlight count as described by Reed (1969).

** Brush was removed during this period.

1.31

w
w

VI

�Table 2. Count and kill data for Area II of Highway 13, south of Meeker
was removed in an attemnt to reduce deer-auto collisions.

g

Colorado, where roadside brush

No. of
Counts

1969
Total
Count

Total
Kill

No. of
Counts

1970
Total
Count

1

5

107

0

4

1+4

1

55

1

4

165

3

4

78

2

4

224

4

4

178

5

4

104

1

Apr.

3

111

2

5

90

1

4

106

3

May

5

130

1

4

11

1

4

45

2

Month

No. of
Counts*

1968
Total
Count

Total
Kill

Jan

2

25

Feb.

4

Mar.

Total
Kill

Vol
Vol

June

4

16

0

4

8

2

4

17

2

July

4

22

1

5

25

0

5

27

1

Aug.

5

61

2

4

19

0

4

45

2

Sept.

4

41

4

4

22

1

5

60

0

Oct.

5

111

4

5

91

9

3

66

2

Nov.

4

68

0

4

121

3

4

67

0

Dec.

4

55

0

-

**

**
--

**
-

3

79

-2

Total

48

919

20

48

837

25

48

738

18

Kill/100 deer counted

2.18

2.99

* Count refers to a roadside spotlight count as described by Reed (1969).
** Brush was removed during this period.

2.44

0\

�- 337 -

Figure 1. Quarter-mile section number 60 of Area II along Highway 13 in
1969 before roadside brush was removed.

Figure 2. Quarter-mile section number 60 of Area II along Highway 13 in
1970 after roadside brush was removed.

�ROADSIDE
o

BRUSH
I

w
~ 3.00
:::&gt;

REMOVAL - HIGHWAY

13

I PRE-TREATMENT

t:'~;jPOST-TREATMENT

o

o
0::

~ 2.00

o

o
o

w

w

..

00

0::

a.. 1.00
W

..J
..J

-

~

1968

1969
AREA I

1970

1968

1969

1970

, AREA]I

•
Figure 3. Pre-treAtment and p'Ost-treatment kill:count ratios for two areas of Highway
13, south of Meeker, Colorado where roadside brush was removed.

�- 339 -

Forbs may also act as an attractant to deer as a food source, especially
in early Summer (Smith 1952), thus increasing the probability of a deer
entering the roadway.
Second, periodic removal of the shrubby species
would probably be necessary because of re-growth from roots and re-invasion
from neighboring areas. New shrub growth is attractive to deer as a food
source, possibly because of increased succulence (Krefting 1941) and nutritional values (Reynolds and Sampson 1943, Hanson and Smith 1970).
The evidence obtained from this study indicates that roadside brush
removal has no obvious effect on the ratio of deer killed on the roadway
by motor vehicles to deer counted during nighttime spotlight counts.
Since
these data do not lend themselves to statistical evaluation, a gross difference in either a positive or negative direction would be necessary for
a sound recommendation concerning roadside brush removal as an effective
measure to reduce deer-auto accidents.

LITERATURE

CITED

Baker, Bertram D. 1970. Survey, inventory, and analysis of deer and elk
winter ranges. Appendix B. Big game winter range analysis, Game Unit
22 - Piceance.
pp. 31-38.
In Game Res. Rep., July Part I. Colo.
Game, Fish and Parks Div. 455 p. (Processed).
Hanson, W.O.,
and Justin G. Smith.
1970. Significance of forage quality
as a tool in wildlife management.
pp. 25-31.
In Range and wildlife
habitat evaluation, a research symposium.
U. S. Dep. of Agr. Forest
Service.
Misc. Publ. No. 1147. 220 p.
Krefting, Lauzits W.
Mgmt.4(1):95-l02.

1941.

Methods

of increasing

deer browse.

J. Wildl.

Myers, Gary T. 1969. Deer-auto accident investigation.
pp. 147-178.
In
Game Res. Rep., July Part II. Colo. Game, Fish and Parks Div. 393 p.
(Processed).
Reed, Dale F. 1969. Techniques for determining potentially critical deer
highway crossings.
Game Information Leaflet No. 73. Colo. Dept. of
Nat. Res., Div. of Game, Fish and Parks.
3 p.
Reynolds, Hudson G., and Arthur W. Sampson.
1943. Chaparral
as browse for deer. J. Wildl. Mgmt. 7(1):119-122.
Smith, Justin G. 1952.
16(2):148-155.

Prepared

Food habits of mule deer in Utah.

by
Thomas M. Pojar
Wildlife Researcher

Candidate

crown sprouts

J. Wildl. Mgmt.

��- 341 July, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RAD~~O~

Project No.

W-38-R-25

Work Plan No.

15

Job Title

_
Deer-Elk

6

Job No.
Deer Underpass

Investigations

Evaluation

Period Covered:

April

1, 1970 through March 31, 1971

Personnel:

William B. Zimmerman, Gary T. Myers,
M. Pojar and Dale F. Reed.

Paul F. Gilbert,

Thomas

ABSTRACT
The use of a deer underpass 10 feet by 10 feet and 100 feet long that
permits deer to move under a four-lane interstate highway to and from
their summer range has been evaluated near Vail. An electro-optical
detection system with two Streeter Amet recorders was installed at each
end of the underpass.
By examining the recorder readings and the number
of deer tracks it was estimated that 157 and 297 deer moved through the
underpass during the spring and fall of 1970 respectively.
According to
the hourly Streeter Amet printout, approximately half of the deer during
the spring and fall moved through the underpass between 12:00 and 5:00 a.m.
The underpass was lighted during nighttime hours.
During the spring, an
index of entrance versus exit activity was determined by dividing the daily
counter readings on the north end of the underpass by that of the south.
Totally, about 18 times as much activity was recorded by the counter at the
entrance as at the exit.

��- 343 -

DEER UNDERPASS

EVALUATION

Dale F. Reed

There are many areas in Colorado where migratory deer cross highways.
When these highways carry high speed, high-volume traffic, the potential
danger to deer and motorists becomes especially great. Therefore, it is
essential to keep the animals off the highway.
Underpasses with specified
dimensions and characteristics located at strategic points within migration
routes may provide adequate means for these necessary deer movements.
This
study purports to evaluate the use of a specific concrete deer underpass 10
feet by 10 feet and 100 feet long located in the Vail area.

P. S. OBJECTIVE
Determine if deer migrating from winter range on one side of Interstate
70 to summer range on the opposite side utilize an underpass constructed
the area.

in

SEGMENT OBJECTIVE
Measure

deer use of the underpass.

METHODS AND MATERIALS
A nighttime lighting system and an electro-optical
detection system with
two Streeter Amet recorders were installed at the underpass.
One hundred
twenty volt power was provided.
Two 150 watt outdoor flood lights were
mounted at each end of the structure and two equally spaced sockets with
100 watt incandescant bulbs were installed inside the underpass.
The lights
were baffled to reduce glare and simulate moonlight conditions.
The lighting
system was activated by a photo-cell.
The electro-optical system and the counters have been described previously
(Myers 1969). The Streeter Amet counter records on adding machine tape
(3M-carbonless paper rolls) the number of times the near-infrared beam is
broken by deer.
It prints hourly the number of counts accumulated and the
hour of the day. During the spring migration of 1970 such a system was
installed at both the entrance (north end) and the exit (south end) of the
underpass.
During the fall migration only the exit (north end) detection
system was activated.
In addition, before the spring migration, dirt was spread over a portion of
the floor (2/3 of floor width for entire length) constructed of a wooden 2X4
lattice partially submerged in concrete.
This raised dirt-covered portion
was used as a track bed. The remaining 1/3 concrete portion was left clear
for water drainage.

�- 344 -

The track bed and entrance to the underpass was checked periodically
for deer tracks.
As deer activity increased and when deer began using
the underpass, the counter and the number of deer tracks were checked
and recorded daily.
The number of deer passing through the underpass
was estimated by carefully examining the number of tracks, and comparing
them with the counter's reading at the exit. The counter's reading on
the entrance was also recorded and compared with that at the exit. During
each check the detection system was reset at zero and the tracks raked out
by hand.

DESCRIPTION

OF AREA

The deer underpass is located in Eagle County 4.3 miles west of Vail,
Colorado, on Interstate Highway 70. The 3.5 mile section of interstate
that includes the deer underpass and associated eight foot fences was
accepted for completion by the Federal Highway Administration on August
26, 1970. The location utilizes well-established
deer migration trails
and a natural drainage referred to as Mud Springs Gulch (Fig. 1). Approximately 3.0 miles of associated eight foot fencing generally parallels the
highway in both directions from the underpass.
The eight foot fencing on
both sides of the highway joins six foot chain-link fences near the West
Vail Interchange, located about 1.5 miles from the deer underpass (Fig. 2).
One-way deer gates and one-way deer gate off-sets are strategically located
in the eight foot fences and have been described separately (Reed 1970).

RESULTS AND DISCUSSION

Spring and Summer Use of the Underpass
It was estimated that 157 deer moved south through the underpass during
the spring and summer migration of 1970. The first passage (south to the
summer range) occurred on May 26 and the last on September 23 (Fig. 3).
The greatest use of the underpass occurred during June with a maximum of
thirteen deer passing through during one night on June 6th. Deer used the
underpass nightly from June 11th through June 24th with periodic use thereafter.
Only nine passages occurred after July 18th and these may have been
resident deer.
An index of entrance versus exit activity was determined by dividing the
daily counter reading on the north end of the underpass by that on the south.
This index (Fig. 4) shows a reluctance on the part of deer to use the underpass. Overall, approximately 18 times as much activity was recorded by the
counter at the entrance as at the exit. The index would be unity, if each
deer that came to the entrance went through and was counted once at the
entrance and once at the exit. Observations, made from some distance away,
bear out the hypothesis that the deer spent a great deal of time at the
entrance before, and possibly without, passing through the structure.
Most
deer were visibly wary of the structure's entrance.
Through examination of
tracks made in the track bed leading into and out of the underpass, it was
estimated that almost all deer used the dirt portion of the floor (1/3 of the
floor is concrete).

�- 345 -

Fig. 1.

Mud Springs Gulch from south of Interstate

foreground.

Highway

70 with Gore Creek in

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.

One-way deer
gate off-set

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8' woven wire fence

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eight foot fences, one-way

Deer underpass

gates, and the West Vail Interchange.

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Fig. 3. Numb~r of deer and dates they moved south through the underpass during the 1970 spring-summer
migration.

..

�50.

INDEX OF DEER ACTIVITY
(NORTH COUNT/
SOUTH COUNT) AT THE VAIL UNDERPASS

I

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readings on the north end of the underpass by th~t on the south, indicates
the part of deer to use the underpass.

the daily counter
a reluctance on

.,

�- 349 -

According to the hourly Streeter Amet printout, 46 percent of 151 deer
moved south through the artifica11y lighted underpass between midnight
and 5:00 AM (Table 1).

Table 1. Estimated times of passages through the underpass as recorded
by the exit Streeter Amet recorders during the spring-summer and fall
migrations.
Spring-Summer
Passages

Percent

Fall
Passages

Percent

Total

Percent

2

1.3

5

1.7

7

1.5

7:00-8:00

o

o

15

5.1

15

3.3

8:00-9:00

2

1.3

7

2.4

9

2.0

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13

8.3

24

8.1

37

8.2

10:00-11:00

3

1.9

12

4.0

15

3.3

11:00-12:00

6

3.8

16

5.4

22

4.9

6

3.8

35

11. 8

41

9.0

1:00-2:00

6

3.8

30

10.1

36

7.9

2 :00-3:00

16

10.2

30

10.1

46

10.1

3:00-4:00

14

8.9

17

5.7

31

6.8

4:00-5:00

28

17.8

24

8.1

52

11.5

5:00-6:00

25

15.9

18

6.1

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9.5

6:00-7:00

25

15.9

15

5.1

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8.8

7:00-8:00

5

3.2

13

4.4

18

4.0

88.1 ]j 412

90.8

Hour

6:00-7:00

12:00-1:00

PM

AM

96.1 }/ 261

Total

151

1/ Passages

occurred when the counter was inoperative

not included
8:00 AM and 6:00 PM.

or between

�THE NUMBER

AND DATE DEER MOVED THRU
THE UNDERPASS

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the 1970 fall migration.

�- 351 -

Sunrise varied from 5:37 to 5:41 AM MDT during June. Fifty two percent
of the 151 recorded passages occurred between 4:00 and 7:00 AM, when light
conditions associated with sunrise may have been a factor. Decreased deer
activity, and an increase in traffic volume and human activity probably
account for the decline in passages throughout the remaining morning hours.

Fall Use of the Underpass
It was estimated that 297 deer moved north through the underpass during the
fall migration of 1970. The first passage in the fall (north to the winter
range) occurred on September 29 and the last on January 6th. The greatest
use of the underpass occurred between October 8th and November 1,2th, with
only sporadic use thereafter (Fig. 5). Only 12 passages occurred after
December 17. Fifty two percent of 261 deer recorded moved north through
the lighted underpass between 12:00 and 5:00 AM (Table 1). Sunrise ranged
from 7:02 to 7:33 AM MDT during October.
Only 18 percent of the 261 recorded passages occurred between 5:00 and 8:00 AM when morning light may
have inhibited their passage.
The estimate of 297 passages during the fall is an increase of 89 percent
over that of the spring-summer use. The reasons for such an increase are
not known.
However, herd increment, a shift in overall migration patterns,
and the lower hunter harvest are probable factors.
The percent of passages
occurring between 12:00 and 5:00 AM in the fall, as compared with the spring,
were very similar.
However, fewer deer (18% in the fall compared to 52% in
the spring) passed through when morning light at the underpass may have been
a factor.

LITERATURE

CITED

Myers, Gary T. 1969. Development of an electronic deer counting device.
Colo. Div. Game, Fish and Parks.
Game Res. Div., Fed. Aid Project
W-38-R-23.
Game Res. Rep. July, Part 2, pp. 235-247.
Reed, Dale F. 1970. Investigation of one-way structures.
Colo. Div.
Game, Fish and Parks.
Game Res. Div., Fed. Aid Project W-38-R-24.
Game Res. Rep., July, Part 3, pp. 439-449 .

~...

[/

"d~,'
Prepared by ,,o:;;tx l\-i¥-~~
Dale F. Reed
Asst. Wildlife

./

Researcher

��- 353 July, 1971

JOB PROGRESS REPORT

State of

COLORADO
--------------------------

Project No.

W-38-R-25

Work Plan No.

15

Job Title

Deer-Elk Investigations

Job No.
7
Effects of Highway Lighting
~O~n~N~u=m=b~e=r~o=f_=D~e~e=r~K=i=l=l~e~d~b~y~V~e=h=i=c=l=e=s
_

Period Covered:

April 1, 1970 through March 31, 1971

Personnel:

William B. Zimmerman, Gary T. Myers, Thomas M. Pojar
and Dale F. Reed.

�- 354 -

EFFECTS OF HIGHWAY LIGHTING
ON NUMBER OF DEER KILLED BY VEHICLES
Dale F. Reed

P. S. OBJECTIVE
Determine if highway lighting
on a portion of Colorado 82.

affects the rate of deer-vehicle

accidents

SEGMENT OBJECTIVES
1.

Record the number of deer killed by vehicles at each end of, and within a 3/4 mile lighted portion of Colorado 82.

2.

Estimate deer densities
by the Ugh ting .

3.

Compare pre-treatment
count data.

4.

Gather additional information which may be needed to understand
deer-auto accident rates are or are not affected by lighting.

adjacent

to the portion

of highway

kill and count data to post-treatment

affected

kill and

why

RESULTS AND DISCUSSION
A joint decision was made by the Colorado Divisions of Highway and Game,
Fish and Parks to delay installation of the illuminaries.
The attached
appendix, a brief report to the Highway Division, discusses the reduced
number of deer in the study area during 1970 and 1971, possible reasons
for it, and recommendations
for further delay.

Prepared

bY~&lt;ni
Dale F. Reed
Asst. Wildlife

Researcher

'}

�- 355 -

APPENDIX

RESEARCH
1971 Deer-Highway

Lighting

Project

Report

Since the approval of the highway lighting project on September 11, 1969,
the installation of the high intensity illuminaries on Highway 82, south
of Glenwood Springs, has been delayed several times.
Initially, a
shortage of lighting materials delayed installation during the fall of
1969. Secondly, the number of deer and the number of deer-vehicle
accidents in the Jammaron study area showed a marked decline during
January, February, and March of 1970 compared to the previous two years.
Hence, a joint decision was made between the Highway Division and the
Game, Fish and Parks Division to further delay installation until 1971
January-March surveys could be made to determine deer numbers and the
occurrence of deer-vehicle accidents.
During the 1971 period, the number of deer (count) and the number of deervehicle accidents (deer kill) were again low (Fig's.l and 2). Several snowshoe trips by project personnel were made into the higher mountainous
terrain adjacent to the study area during February and March of 1971. It
was speculated that the deer wintering in this area would move down into
the immediate study area if snow depths limited the availability of natural
forage and the animals mobility.
Numerous deer were found wintering there
during the middle of March and snow depths ranged from zero to 16 inches
with most of the southern exposures free of snow.
Weather, then, is suspected of influencing the movement of deer toward the
study area. Weather has been monitored near the Highway 82 study area
continuously since 1969. The average minimum temperature for January through
March increased greatly from 1969 to 1970 and again slightly for 1971 (Fig.
3). While this increase in the average minimum temperature occurred, the
total amount of precipitation per year declined from 2.19 inches in 1969 to
1.52 inches in 1971. Thus, less precipitation and higher temperatures during
this period may mean that more winter forage remained available to the animals
(less snow cover) and that only a few moved down to the highway area. With
acceptance of this hypothesis and by considering the ten year U. S. Weather
Bureau average for the city of Glenwood Springs (average minimum temperature
15.40 F, total average precipitation 3.14 inches) the probability of having
colder temperatures and greater snow accumulations, and, consequently, deer
moving into the study area in relatively large numbers is high.
Recommendations
Another delay in the installation of the illuminaries is hereby requested.
It is felt another year of delay is worthwhile to make sure that deer return
in sufficient numbers so that possible reductions in deer-auto accidents can
be detected.
Meanwhile, a recently installed Animated Deer Crossing Sign
will be evaluated in the study area.

�HIGHWAY 82 SPOTLIGHT COUNT DATA
(JAMrv1l~RON ARE;~)
~------------------------~
---

130
120

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JAN

----:-;--

Fig. 1.

Number

----

"'0_ - _ \91
..,.,.
.. -- ---.".,. ~ ---- ---.,..,---

- - -- ---_- 1971

- ---------------------------~--~-~--=--~-_-_-_--_-_------iw
---

_

_

---~
FEB
I

MAR
of deer counted adjacent

to the Jammaron

study area south of Glenwood

Springs.

�.~

30r-

Ii

~

HIGHWAY 82 ROt~D' ~(ILL DAT.L\·
(JA~.~MARON A,REA)
=••
-

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1971

.. ----- ----------

\91.~-----

Fig.J~~Number

--

-------=-----_----

-----

_"-'~...,.-----

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-----------

of deer killed by vehicles on the JammaroJFs~u~y area south of Glenwood Springs.

f\~AR

�HIGHWAY 82 PRECIPITATION

JANUARY,

!\ND TEMPERATURE

FEBF~Ut;RY AND rv1A,RCH

PRECIPITATION
TEMPERATURE

'--

---

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-

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1Io---r-------------,r-." -----------..---'
1969

1971

1970

Fig. 3. Average minimum temperature and total precipitation
Springs during January, February, and March.

for the Jammaron

study area south of Glenwood

~

w

V1
00

�- 359 July, 1971

JOB PROGRESS REPORT

State of

COLORADO
----------~~--~----------

Project No.

W-38-R-25

Work plan No.

15

Job Title

Deer-Elk

Investigations

Job No.
--=8
Effect of Lighted Deer Crossing
Signs on Number of Deer Killed by Vehicles

Period Covered:

April 1, 1970 through March 31, 1971

Personnel:

William

B. Zimmerman,

_

Dale F. Reed and Thomas M. Pojar

ABSTRACT
A lighted deer crossing sign was evaluated for its effect on the speed of
traffic on State Highway 82 near Glenwood Springs, Colorado.
Vehicle speeds
were recorded with a Newmeyer Model NH-1 Vehicle Speed Recorder.
Sixteen
days of pretreatment data and twenty eight days of treatment data were
collected.
The mean speed of vehicles passing the recording station during
the pretreatment period was 54.52 mph. The mean speed dropped to 53.03 mph
during the treatment period, which is a significant (P&lt;.05) reduction.
The
value of this small average reduction in speed in preventing deer-vehicle
collisions was not determined during this segment.

��- 361 -

EFFECT OF LIGHTED DEER CROSSING
SIGNS ON NUMBER OF DEER KILLED BY VEHICLES
Thomas M. Pojar

Although segment objectives 1 and 2 were accomplished, their value is
limited.
As explained in the Job 7 Progress Report, W-38-R-25, few
deer were present in the study area. Therefore, the sample of deer
killed and deer observed is too small to be meaningful.
Although deer
were present only in small numbers, evaluation of the effect of the
sign on the speed of traffic was still possible.

P. S. OBJECTIVE
Determine
accidents

if lighted deer crossing
on a portion of Colorado

signs affect the rate of deer-vehicle
82.

SEGMENT OBJECTIVES
1.

Record the number of deer killed by vehicles within
highway marked by lighted deer crossing signs.

2.

Estimate

3.

Compare pre-treatment
count data.

4.

Gather additional information which may be needed to understand why
deer-auto accident rates are or are not affected by lighted deer
crossing signs.

deer densities

adjacent

a mile portion

to the signed portion

of

of the highway.

kill and count data to post-treatment

kill and

METHODS AND MATERIALS
The lighted deer crossing sign consisted of a reflectorized yellow, diamondshaped background with the words "Deer Xing" centered on the sign and lighted
with neon tubing (Fig. 1). The tubing was covered with a 1/4" sheet of plexiglass to prevent breakage by vandals.
Vehicle speeds were recorded with a Newmeyer Model NH-l Vehicle Speed Recorder
(Newmeyer Electronics, 4521 West Claremont, Glendale, Arizona).
Timers and
magnetic loops were located 800 feet behind the deer crossing sign. The sign
was turned on at 6:30 PM and off at 10:00 PM every day during test periods.
The speeds of nearly all the vehicles that passed the station in the right
lane were recorded.
Eighty vehicle speeds were randomly sleeted from this
sample for tabulation.

�- 366 -

LITERATURE CITED
Yeager, L. E. (ed.). 1969. Colorado game research review. Big game
research. Published by Colo. Div. of Game, Fish and Parks. 35 p.

7;i'
~~::4J @ R~ ,

Prepared by

Thomas M. Pojar
Wildlife Researcher Candidate

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                  <text>-1-

October, 1971

JOB PROGRESS REPORT

State of

C~O~L~O~RA~D~O~ _

Project No.

W-88-R-16

Work Plan No.
Job Title
Period Covered:

1

Migratory Bird Investigations
Job No.

1

Waterfowl Production Surveys
April 20, 1970 to June 30, 1970

Personnel: C. Bryant and Staff, Monte Vista National Wildlife Refuge;
R. Pearson, Brown's Park National Wildlife Refuge; C. Hayes and J. Randall,
Bureau of Sport Fisheries and Wildlife; G. Robinson, Colorado State University; Delwin Benson, C. Braun, W. Carpenter, H. Funk, J. Grieb, J. Hatfield, C. Hector, R. Hopper, J. Houston, W. Russell, S. Steinert, J. Wheeler
and M. Szymczak, Colorado Division of Game, Fish and Parks.

ABSTRACT
Weather and water conditions were somewhat variable through major breeding
areas in the state, yet generally good in the state as a whole. The total
estimated number of 57,389 duck breeding pairs was 16.9 percent below the
1969 total and 6.1 percent below the 16 year average. Major reductions in
breeding ducks were noted in North Park and Cache 1a Poudre Valley, while
total numbers of pairs in other areas remained near the 1969 figures. The
mallard continues to be the major breeding species in Colorado. Production
of Canada geese in northwest Colorado declined 16.7 percent from the 1969
level but the total number of birds observed in the area w:as nearly the
same as in 1969. A major reduction from 1969 in both goslings produced and
total number of adult birds in the Denver area resulted in 22.7 percent
decline in the total number of geese in north-central Colorado.

��-3-

WATERFOWL PRODUCTION SURVEYS
Michael R. Szymczak
P. S. OBJECTIVE
To formulate waterfowl harVest regulations.
SEGMENT OBJECTIVES
To determine, through statistically reliable sampling techniques, the
number of duck and goose breeding pairs, by species, in each major Colorado waterfowl breeding ground.

METHODS AND MATERIALS
Present duck breeding-pair and production surveys consist of a breeding
pair inventory of only major production areas.
The 1970 duck surveys were conducted during the period May 11 to May 31.
Ground counts were made in the Yampa Valley and Brown's Park. Aerial
surveys were conducted in North Park and the South Platte and Cache la
Poudre Valleys. Intensive aerial counts combined with air-ground comparison studies were conducted in the San Luis Valley. Species visibility
ratios attained in the San Luis Valley were also applied to North Park data.
All survey methods and sample areas remained the same as in previous years.
Flying was accomplished with a Cessna 185 aircraft. Areas ampled by
section or block were flown with one observer, while two observers were
used in sampling by transect.
Ground surveys of Canada goose production were conducted within the period
of April 20 to June 19. Production estimates in the Yampa Valley-Brown's
Park area were made from as complete a count as possible of breeding pairs,
goslings, and eggs. Gosling production and population size in the DenverBoulder-Fort Collins area were determined by conducting a count on June 19,
1970 of all geese on moulting and brood rearing areas and classifying birds
as to whether they were adults or go~lings.
On the basis of these studies, a report is submitted to the Bureau of Sport
Fisheries and Wildlife, which constitutes Colorado's part in the annual
continental cooperative breeding ground survey.
RESULTS AND DISCUSSION
Weather and water conditions for waterfowl nesting were somewhat variable
through the major breeding areas in the state. A heavy winter snowpack,
late spring snows and a retarded run-off were factors affecting all major
waterfowl breeding areas except the San Luis Valley. Water was extremely

�-6-

Springs making the search for nests impossible. Therefore, number of
nesting pairs and total goslings recorded is quite possibly an underestimate
for the Yampa River (Table 3).

Table 3. Number of Canada geese observed and estimated production, Moffat
County, Colorado, 1970.

Area

Nesting
Pairs

NonNesting
Pairs

Total
Adults

Estimated
Number
Goslingsl/

Total
Birds

Yampa River y
Craig to Juniper Spgs.
Juniper to Cross Mtn.
Lilly Park

14
12

164
97
133

174
125
157

20
67
53

194
192
210

YAMPA TOTALS

31

394·

456

140

596

Green River
(Brown's Park)

31

106

168

117

285

Little Snake River
Above upper bridge
Below upper bridge

4

5

25

137

13
187

10
87 l!

23
274

Total

29

142

200

97

297

GRAND TOTAL

91

642

824

354

1,178

5

1/ Calculated using average brood size and number of successful nests.

2/ High water inundated nesting islands making it difficult to estimate
number of nesting pairs and goslings.

1/ Includes 43 goslings hatched from eggs taken from nests before final survey.
The number of birds observed on the Yampa River in 1970 increased by 150 in
comparison to the 1969 total (Table 4). The total numbers on the Green and
Little Snake Rivers declined about 25 percent. Over-all, an increase was
recorded in the total number of adults in the survey area, but lower production estimates, as indicated in Table 5, resulted in a total population
similar in size to that of 1969.

�-7Table 4.

Total Canada geese observed, Moffat County, Colorado, 1970.
No. Geese Counted
1956-1969
1969 Average
1970

Area

Percent Change
From 1956-1969
Average
From 1969

Yampa River

596

456

376

+30.7

+ 58.5

Green River

285

375

111

-24.0

+156.8

Little. Snake River ]J

297

391

229

-24.0

+ 29.7

1,178

1,222

716

- 3.6

+ 64.5

Total

1/ Not included in survey until 1962~

Table 5.
1970.

Estimated number of Canada goose goslings, Moffat County, Colorado,

Area

No. of Goslings
1956-1969
1970
1969
Average

Percent Change
From 1956-1969
Average
From 1969

Yampa River

140

172

140

-18.6

0.0

Green River

117

144

40

-18.8

+192.5

Little Snake River ]J

97

109

72

-11.0

+ 34.7

Total

354

425

252

-16.7

+ 40.5

1/ Not included in survey until 1962.

Results of the June 19, 1970 census of Canada geese in the Fort Co11insBoulder, Denver area are presented in Table 6. Goslings that were subsequently removed for transplant purposes are included in totals for the
respective areas. In the Wellington production area, geese utilized 17
water areas in 1970 compared to only eight areas in 1969. Geese in the
Fort Collins, Loveland, Boulder and Denver areas used essentially the same
nesting areas in 1970 as in 1969.

�-8-

Table 6.

Results of north-central Colorado goose census, June 19, 1970.

Production
Area

No.;
Broods

Total No.
Goslings

Total No.
Adults and
Yearlings

Total No.
of Birds

2
3
0
3
3
1
1

84
7
7
0
11
13
4
4

210
4
6
2
16
45
2
3

294
11
13
2
27
58
6
7

12
7
36
5
32
11
50
12

12
45
109
13
2
38
5
86
16

24
52
145
18
8
70
16
136
28

301

614

915

28
56
27
5
2
28
17
20
9
6
36
4

37
196
21
7
4
30
33
22
13
8
63
'9

65
252
48
12
6
58
50
42
22
14
99
13

238

443

681

18
8
37

56
8
41

74
16
78

63

105

168

Wellington

Water Area
Terry Lake
Deines Reservoir
Launer Pond
Wood Pond
Douglas Lake
North Poudre No. 1
North Poudre No. 3
North Poudre No. 5
Bureau of Standards
Pond No. 1
Divide No. 8
Elder Reservoir
Annex No. 8
Long Pond
Cobb Lake
Hinkley Reservoir
Watson Lake
Curtis Lake

3
2
7
1
1
6
2
4

Sub-total
Fort Collins

Herring Lake
College Lake
Dean Acres
Andrijeski Marsh
Claymore Lake
Sterling Gravel Pits
Linderuneier Lake
Anderson Pond
Parkwood Lake
Kitchel Lake
Timnath Reservoir
Romily Gravel Pit

5
1
1
6
6
2
1
9
1

Sub-total

6

Loveland
Boedecker Reservoir
Flatiron Gravel Pits
Welch Reservoir
Sub-total

5
3

----------------------------------------------------------------------------

�-9-

Table 6. Results of north-central Colorado goose census, June 19, 1970.
(continued).

No.
Broods

Total No.
Goslings

Total No.
Adults and
Yearlings

Total No.
of Birds

0
0
0
4
1

0
0
0
18
4

41
10
110
37
2

41
10
110
55
6

9
1
2
1
6
3

34
4
6
5
20
11

144
14
26
123
18
200

178
18
32
128
38
211

102

725

827

124
35
13
11
29
2
2
13
18

328
41
9
4
36
2
2
8
17

452
76
22
15
65
4
4
21
35

Sub-total

247

447

694

GRAND TOTAL

951

2,334

3,285

Production
Area
Denver

Water Area
So. Colo. Blvd.
and Quincey
Mohn Estate
Denver City Park
Sloan's Lake
Hyatt's Lake
Cline &amp; Kountee
Reservoir
Federal Center
Clairfield
Kings Pond
Upper Tule Lake
Bowles

Sub-total
Boulder

Valmont Reservoir
Terry Lake
Crystal Lake
Ish Lake
Faivre Ponds
Hayden Lake
Angus Ranch
Boulder Valley Farm
Peaceful Valley Farm

Increases in production from the 1969 level were recorded in the Wellington
and Boulder areas (T~ble 7). However, a major decline in production in the
Denver area resulted in a 15 percent reduction in total goslings produced.
A reduction of approximately 54 percent in total geese in the Denver area
resulted in a decline of around 23 percent in the total population (Table 8).

�-10-

Table 7. Total number of Canada goose goslings observed in north-central
Colorado production trend areas, 1969-1970.

Area

Number of Goslings
1970
1969

Percent Change
From 1969

Wellington

301

257

+ 17.1

Ft. Collins

238

256

7.0

Loveland

63

37

+ 70.3

Boulder

247

149

+ 65.8

Denver

102

426

- 76.1

Total

951

1,125

- 15.5

Table 8. Total number of Canada geese observed in north-central Colorado
production trend areas, 1969 and 1970.

Area

Number of Geese
1970
1969

Percent Change
From 1969

Wellington

915

945

- 3.2

Ft. Collins

681

749

- 9.1

Loveland

168

140

+20.0

Boulder

694

630

+10.2

Denver

827

1,784

-53.6

Total

3,285

4,248

-22.7

�-11-

Fall Flight Prediction
A late season run-off should provide excellent conditions for brood sruvival
in most areas. 'However, the size of the fall flight of ducks from Colorado
will be smaller than the previous two years.
The Canada goose flight from northwestern Colorado will be similar in size
to the 1969 flight. The north-central goose flock fall flight will also
be similar to 1969 in all areas except those influenced by reduced production in the Denver area.

Prepared by

J?'hJ.~17P.

--4,,,

Michael R. Szymczak
Assistant Wildlife Researcher

��-13October,

JOB PROGRESS

S tate of
Project
Work

1971

REPORT

C.=.O.=.L::.O::.;RA=D::.,:O::-_
No.

Plan No.

Job Title

Period Covered:

.Migratory

.W-88-R.,.16
1
Trapping

Bird Investigations

Job No.
and Banding

June 25, 1970 to March

2

Ducks and Geese

31, 1971

Personnel:
Charles Hayes, Dale Horne and Jack Randall, Bureau of Sport
Fisheries and Wildlife;
Delwin Benson, George Bock, C1ait Braun, Bill
Carpenter, Dale Coven, Gurney Crawford, David Croonquist, Howard Funk,
Larry Green, Jack Grieb, Bob Kitzmiller, Mike Robinson, Bill Rutherford,
Mike Szymczak, Ken Wagner and Richard Hopper, Colorado Game, Fish and
Parks Division.

ABSTRACT
Over 9,600 ducks, including 12 species, were banded and released at six
locations in Colorado during Segment 16. Mallards contributed 8,278 or 86
percent of the total.
The pintail was the next most common species in the
banded sample, with 871 or 9 percent of the total.
Preseason banding in the
San Luis Valley and South Park produced 2,503 and 1,179 ducks, respectively.
Postseason banding (winter) in eastern Colorado accounted for the remaining
5,937 ducks in the banded sample.
Goose trapping resulted in the banding of 2,293 Canada geese during the
Segment.
Most of these (1,864) were banded postseason in the Arkansas and
Cache 1a Poudre Valleys.
The remaining 429 geese were banded as part of the
annual gosling transplant program.
These transplant birds were released in
North Park and the San Luis, South Platte and Dolores Valleys during June
and July.

�-14-·

RECO:MMENDA TrONS

1.

Continue preseason duck banding in the San Luis Valley and South Park.
Reduce the mallard quota in the San Luis Valley from 4,000 to 1,000
birds.
Place add Lt.Lona I emphasis on the banding of other species of
ducks, preferably up to 500 of each major species in each area.
Attempt to obtain equal age and sex ratios in the sample by species.

2.

Initiate preseason duck banding in North Park with quotas of 1,000
mallards and up to 500 of each of the other major species.
Attempt
obtain equal age and sex ratios in the sample by species.

to

3.

Initiate a postseason mallard banding program in the UncompahgreGunnison-Colorado
River complex.
Obtain a quota of up to 1,000 birds,
divided approximately equally by age and sex.

4.

Continue goose banding
for Segment 16.

program

as outlined

in the W-88-R

Job Description

�-15-

TRAPPING

AND BANDING DUCKS AND GEESE
Richard M. Hopper

This report summarizes all waterfowl banding activities of Federal Aid
Project W-88-R-16 for the Segment year April 1, 1970 to March 31, 1971.
Analysis of band recovery data occurs as a separate job (Work Plan 1, Job 3),
and work of this type is not included here. This report merely presents a
tabulation of numbers of ducks and geese banded by species and location
during the Segment.

P. S. OBJECTIVE
To formulate waterfowl

harvest

regulations.

SEGMENT

OBJECTIVES

1.

To trap and band ducks and geese for the purpose
tion and annual mortality data.

2.

To report species and numbers of ducks winter banded in the South
Platte Valley, Arkansas Valley, San Luis Valley, and Bonny Reservoir
part of Work Plan 3, Job 6.

METHODS

of accumulating

migra-

as

AND MATERIALS

Methods and materials employed during Segment 16 were the same as in previous Segments (Hopper 1968), except that use of the corral trap for drivetrapping ducks during the mid-summer molting period was discontinued.
North
Park was dropped from the list of duck trapping locations in Segment 16,
while the Dolores Valley, being a transplant site, was included as a goose
banding area. All other locations and times of trapping and banding
remained the same as in Segment 15 (Hopper 1970).

RESULTS AND DISCUSSION
Ducks
Numbers and species of ducks banded during Segment 16 are listed in Table 1
by location.
Over 9,600 ducks, including 12 species, were banded and released
in six locations.
As in past years, the mallard was by far the most common
species in the banded sample, contributing 8,278, or 86 percent of the total.
Pre-season banding in the San Luis Valley and winter banding in eastern Colorado accounted for most of the mallard total. The pintail was the next most
abundant species in the banded sample, with 871, or 9 percent of the total.
Less than 100 birds of each of the remaining species were banded, except for
green-winged teals and redh~ads which yielded 174 and 136 birds, respectively.

�Table 1.

Number of ducks banded by species, location and period of year, 1970-71.

Species
Mallard

Pre-season'!'!
San Luis
South
Valley
Park

Cache 1a
Poudre Valley

South Platte
Valley

Post-seasonY
Bonny
Arkansas
Reservoir
Valley

San Luis
Valley

Total

537

8,278

1,628

767

596

3,064

754

9

5

--

--

--

3

--

--

--

--

Green-winged teal

45

129

--

--

--

174

Blue-winged and
cinnamon teal

46

40

--

--

86

Shoveler

2

--

Gadwall
American widgeon

Pintail
Redhead
Lesser scaup
Ring-necked duck
Ruddy duck
Total

1/ August-September
1/ January-February

--

932

14
52

--

--

55

2

634

237

--

--

133

1

--

--

1

--

--

--

1

1

--

--

--

1

1

--

--

--

1

2,503

1,179

596

3,064

--

871
2

754

986

--

537

136

9,619

I

I-'
0\
I

�-17-

Pre-season efforts in the San Luis Valley resulted in the banding of 2,503
ducks. Only 1,628 mallards were banded in the northern portion of the
Valley, short of the usual 2,000,bird quota. A good sample of pintails
(634) was also banded. The final early experimental duck season was held
in the San Luis Valley in 1970, and as a result, the "Cooperative Mallard
Investigation" will terminate following the 1971 breeding population survey.
Pre-season banding should cdntinue as part of Work Plan 1, Job 2, but with
a reduced mallard quota. At least 1,000 mallards, divided equally by age
and sex, should be banded pre-season in the Valley annually, with 500 north
and 500 south of the Rio Grande River. Additional emphasis should be
placed on the banding of other species of ducks in all areas.
Pre-season banding was continued in South Park for the third consecutive
year. A banded sample of 1,179 ducks was attained, with mallards comprising
767 of this total. Pintai1s and green-winged teals contributed 237 and 129
banded birds, respectively.
Trapping and banding was not attempted in North Park during Segment 16.
Past work involved the drive-'trapping of molting ducks, which yielded poor
samples of certain species, sexes, and ages of ducks. Pre-season baittrapping should be initiated in Segment 17 in favor of mid-summer drivetrapping. This would results in pre-season banding being conducted at
approximately the same time in all of Colorado's three major high mountain
production areas. This is desirable to better understand the relationships
of birds among the three areas, and to obtain mortality estimates by species
and age. Thus, it will be necessary to band good samples of adult and
immature birds of all important species.
Winter (post-season) mallard banding quotas were reduced significantly in
Segment 16. Quotas were dropped from 1,000 mallards in each of seven areas
to 600 in each of nine areas. The Arkansas Valley was divided into two
areas and the San Luis Valley was added the previous Segment, accounting for
the increase in number of areas. The result was the banding of 5,883 mallards
during January and February, 1971, or about 500 birds above the overall
quota. Work Plan 3, Job 6 progress report covers this work in greater detail.
Geese
Colorado's goose trapping program during the 1970-71 Segment resulted in
the banding of 2,293 Canada geese (Table 2). Post-season banding of wintering goose populations in the Arkansas and Cache 1a Poudre Valleys accounted
for most of this total (1,864). The remaining 429 geese were banded as
part of the annual gosling transplant program. Thirty-six goslings, hatched
from eggs collected on the Yampa and Little Snake rivers west of Craig, were
released on Totten Reservoir in the Dolores Valley. Goslings transplanted
in the other three areas originated from eggs collected in the Denver and
Fort Collins areas and from birds trapped in the Denver-Boulder-Fort Collins
complex.

�-18-

Table 2.
1970-71.

Number of Canada geese banded by location and period of year,

Number Banded
Summer
Transplantsll
Post-Season£/

Location

Total

Arkansas Valley

873

873

Cache la Poudre Valley

991

991

South Platte Valley

19

19

San Luis Valley

68

68

North Park

306

306

Dolores Valley

36

36

Total

429

1,864

2,293

11 June-July
'1:/ January

LITERATURE CITED
Hopper, R. M. 1968. Trapping and banding ducks and geese. Colo. Game,
Fish and Parks Div., Game Research Rept. Oct., p. 7-12.
Hopper, R. M. 1970. Trapping and banding ducks and geese. Colo. Game,
Fish and Parks Div., Game Research Rept. Oct., p. 11-16.

Prepared

by:~it~Ju~·
""-'CJ}~~",-. ,.....L-.:...-..:~~~

',l-1..~ __

Richard M. Hopper ~
Wildlife Researcher

�-19October,

1971

JOB FINAL REPORT

State of

COLORADO

Project No.

~W_-_8_8_~~R~-_1_6
_

Work Plan No.

I

Job Title:

Analysis

Period Covered:
Personnel:

April

Higratory

Bird Investigations
3

Job No.
of Waterfowl

Banding

Data

1, 1970 through March 30, 1971

Ed Kautz, Dale Coven and Michael

R. Szyrmczak

ABSTRACT
From 1954 through 1969, a total of 14,753 bands were placed on thirteen
species of waterfowl and the American coot in the North Park area of Colorado.
The bandings resulted in a sufficient number of recoveries enabling
varying degrees of analyses of the mallard, pintail, green-winged teal, and
American widgeon populations.
Apprnximately 84 percent of all mallard
recoveries were taken within the Central Flyway portion of the United States,
69.9 percent within Colorado.
The Pacific Flyway portion of Colorado
accounted for an additional 8.5 percent of the total recoveries.
Forty-four
percent of all mallard recoveries within Colorado were taken in the northcentral portion of the state, while 27.9 percent were taken in the San Luis
Valley.
Adult male mallards-had an overall first year recovery rate of
.063. The annual mortality rates for mallards calculated for each sex and
age" group using the composite dynamic-method w.ere as follows:
adult male,
.387; adult females, .456; local males, .488; local females, .500. Adult
male mallards in North Park have very similar mortality character.istics to
their counterparts in the San Luis Valley.
Approximately 29 percent and
25 percent of all pintail recoveries were taken in the states of Texas and
Louisiana, respectively.
The west coastal area of Mexico and the Sacramento and San Joaquin Valleys of California are additional pintail recovery
areas of note.
Pintails recovered in Gulf Coast areas seem to follow a
more direct migration path rather than utilizing the "counter-clockwise"
migration route described by Low (1949). First year recovery rates for
adult male pintai1s in years which had more than 300 birds banded averaged
.013. Annual mortality rates for adult male and female pintai1s calculated
by the composite dynamic method are .347 and .331, respectively.
Thirty-six
percent of the adult male green-winged teal recoveries were taken in California, mainly in the San Joaquin and Imperial Valleys.
An additional 32
percent of the adult male green-winged recoveries were recovered in Texas and
Louisiana, generally in the Gulf Coast region.
Adult male green-wings had
a first year recovery rate of .013 and an overall recovery rate calculated
by the composite dynamic method of .383. Approximately 29 percent of the
American widgeon recovered were reported taken in Mexico with the west coast
state of Sinaloa the major recovery area.
The remainder of the widgeon
recoveries were generally scattered with the Pacific Flyway the major recovery
area.
The first year recovery rate of widgeon banded as locals was .030
compared to .011 for adults.

�....
20-

RECOMMENDATIONS

1.

Duck banding should be continued in No r t h Park, Methods of capture
should be employed which will result in a balanced sex and age distribution of the target species.

2.

Because of their relative abundance in North Park, the following species
should be considered target species:
mallard, pintail, green-winged
teal, blue-winged teal, cinnamon teal, American widgeon
and gadwall.
Because of their current importance on a continental basis, bands
should also be placed on the canvasback and redhead.

3.

The list of target species should be reviewed periodically to determine
if additional species should be added to the list or possibly some
species deleted.

4.

No more thanl,200birds,
300.-of each sex and. age group, Of any: species
should be banded during the first year of banding.
In subsequent years
the size of the banded sample should be de.termined on the basis of
information received concerning cohorts banded in previous years.

5.

Information concerning banded birds recaptured in North Park and
North Park birds recaptured elsewhere should be examined in an attempt
to determine the-origin of birds in the North Park moulting population.

6.

The results of the completed banding analysis in this repOrt and
additional information r'eeeLved from orr-go Lng' Nor th Park- banding
should be compared with the results of similar analyses for populations
banded in South Park and the San Luis Valley in order to determine if
duck populations in Colorado I s high mount.aLn parks ha've similar characteristics.

�-21-

ANALYSIS

OF WATERFOWL

Michael

BANDING

DATA

R. Szymczak

North Park is an intermountain valley located in Jackson County in northcentral Colorado.
The elevation of the park is around 8,000 feet.
It is
one of Colorado's major duck breeding areas.
Duck banding in North Park
began in the summer of 1954 ~nd has continued on an annual basis through
1969.
Drive trapping of youhg and flightless adults has been the method
employed in all but the first two years of banding.
These efforts resulted
in 14,753 bands being placed on ducks, coots and mergansers.

P. S. OBJECTIVE
To formulate waterfowl

harvest

regulations.

METHODS AND MATERIALS
All data were obtained from computer tapes received from the Bird Banding
Laboratory of the Migratory Bird Populations Station.
The tapes contained
all bandings from North Park and all recoveries from those banding through
the 1969-70 hunting season.
The tapes were subjected to automatic data
processing programs which were constructed to obtain the desired information.
The resulting printouts were used in the analysis.
The scientific names for the birds mentioned
in the Appendix.

in this report are presented

RESULTS AND DISCUSSION
Bands were placed on 13 species of waterfowl and the American coot. The
number of birds banded and recoveries received are presented by species in
Table 1.
Substantial numbers of mallards, pintail, widgeon, and the three species
of teal were banded during the 1954 through 1969 period.
A relatively
small number of each of the other species were banded.
Because of the
small banded sample of some species and low recovery rate of others, information regarding only the mallard, green-winged teal, pintail, and American
widgeon will be considered.

Mallard
Distribution

of Recoveries

Approximately 91 percent of the recoveries of male mallards banded as
locals and 85 percent of those banded as adults were taken in the Central

�-22-

Flyway portion of the United States (Tables 2 and 3). About 9 percent of
the local male recoveries and 12 percent of the adult male recoveries
came from the Pacific Flyway.
Colorado was the only recovery area of
significance in the, Central Flyway for either locals or adult. Approximately
74 percent of the Pacific 'Flyway recoveries of adult male mallards were
also taken in Colorado (Table 2). Scattered recoveries of adult males in
New ~~xico and the panhandle area of Texas indicate some movement of birds
south and southeast out of Colorado.'

Table 1. Number of birds banded by species in North Park, 1954-69, and
recoveries received through the 1969-70 hunting season.

No. Birds
Banded

No. Bands
Recovered

Percent
Recovered

3,130

369

11.8

3,687

110

3.0

Pintail

3,390

120

3.5

American widgeon

1,557

62

3.0

1,538

23

1.5

Gadwall

410

14

3.4

Shoveler

298

3

1.0

275

4

1.5

147

5

3.4

135

6

4.4

duck

99

7

7.1

Common merganser

47

4

8.5

'Canvasback

40

5

12.5

Species

1/

Mallard

Green-winged

Blue-winged

American
Lesser

teal

and cinnamon

teal

coot

Scaup

Redhead
Ring-necked

1/ Includes

two birds of unknown

age at time of banding.

�-23-

Table 2. Distribution of recoveries
North Park, Colorado, 1954-69. );/

from local male mallards

banded

in

Total

No.

%

Indirect
No.

Wyoming

1

5.3

0

Nebraska

1

5.3

2

15.4

3

9.4

Colorado

17

89.4

6

46.1

23

71.9

New Mexico

0

1

7.7

1

3.1

Oklahoma

0

1

7.7

1

3.1

10

76.9

29

90.6

Direct
Area

%

No.

%

1

3.1

United States
Central

Flyway

Sub-total

Pacific

19

100.0

Flyway

Idaho

o

1

7.7

1

3.1

Washington

o

1

7.7

1

3.1

Colorado

o

1

7.7

1

3.1

o

3

23.1

3

9.3

19

13

Sub-total

Total

l/ Includes

a limited number of birds recovered

32

which were banded

as immatures.

�-24-

Table 3. Distribution of recoveries
North Park, Colorado, 1954-69.

Direct
No.

Area

from adult male mallards

banded

in

%

Indirect
No.

%

a

0.0
0.0

4
3

2.7
2.0

4

3

1.4
1.1

a

0.0

7

4.7

7

2.5

2
1
1

1.5
0.8
0.8

a
a

0.0
0.0

2

0.7

1

0.4

5
1

3.4

6

2.2

0.7
1.4
68.9

1
4
200
12

0.4
1.4
71.9

9

3.2

Total
No.

%

Canada
Alberta
Saskatchewan

·0

Sub-total

(Canada)

United States
Central

Flyway

Montana
North Dakota
Wyoming
South Dakota
Nebraska
Colorado
New Mexico
Texas

o

Sub-Total

2

1.5

2

98

75.4

5
4

3.8
3.1

102
7

113

86.9

5

4.7
3.4

122

82.5

235

84.5

1

1

0.4
0.4
0.4
0.4

4.3

Pad fic Flyway

a
a

Utah
Oregon
Idaho
Washington
Colorado
New Mexico
Wyoming

1

14
2

11
1
2

0.7
1.4

2

9.0
1.1
0.7

7.4

1
1
1
25
3

17

13.1

17

11.6

34

12.4

a

0.0

1

0.7

1

0.4

a

0.0

1

0.7

1

0.4

130

100.0

140

94.6

270

97.1

o

0.0

1

0.7

1

0.4

Flyway

Sub-total
Sub-total

Total

1

10.8
1.5

a

Ohio

Mexico

a

o

Sub-total
Mississippi

0.8

0.7
0.7
0.0
0.7

1

(U. S.)

130

148

278

�-25-

There was no substantial difference between the distribution of adult males
recovered the first season after banding, or directly, and those recovered
in following years or indirectly (Table 3). Although based on only a
small number of recoveries, birds banded as locals did exhibit some variation
in direct and indirect recovery areas (Table 2).
Recoveries from both local and adult females showed a similar distribution
to that of the males with Colorado being the maj or recovery area (Tables 4
and 5). It is interesting that no male recoveries were received from the
southern Pacific Flyway states of Nevada and Arizona, while three of a
total of 18 local female recoveries were taken in those states.
Male
recoveries from the Pacific Flyway, with the exception of Colorado, came
for the most part, from the northwestern states and were almost exclusively
indirect.
These data for males indicate that drakes migrating from North
Park directly into the Pacific Flyway may winter exclusively in Colorado.
The lack of Canadian recoveries of North Park banded mallards indicate the
population is not related to breeding populations utilizing the prai.rie
provinces.
The recovery distribution of mallards taken within Colorado is presented
in Table 6. There is essentially no difference between the distribution of
adults recovered directly or indirectly.
The immatures show some possible
variations, but any analysis is again hampered by sample size of any group
other than the adult males.
These data in Table 6 indicate that the foothills area of north-central
Colorado is the most important recovery area for North Park banded birds.
The San Luis Valley is also of major importance.
There are indications
that North Park females move into the north-central area at a greater rate,
percentage-wise
than males. However, four years of sex oriented regulation
in the San Luis Valley designed to put hunter pressure on the male and reduce
pressure on the female may have distorted the true distribution of birds
between the two major recovery areas.
There is some possibility that some
males which breed in the San Luis Valley may migrate to North Park to moult,
then return to the San Luis Valley in late summer and early fall.
Substantial numbers of North Park mallards apparently do not move into the
northeastern part of the state where hunting pressure is relatively high.
The west slope does account for a substantial portion of recoveries, particularly adults.
If recoveries were weighted simply on the basis of t he :number
of active duck hunters, the west slope would be as important a recovery
area as north-central
Colorado for North Park banded mallards.
However, any
weighting procedure should also consider such things as season lengths, bag
limits and reporting rates and that type of analyses will not be considered
for these data.

�-26-

Table 4. Distribution
North Park, Colorado,

of recoveries

from local

female mallards

banded in

1954-69. }j

Area

Direct
No.
%

Indirect
No.
%

Total
No.

0

1

14.3

1

5.6

0

1

14.3

1

5.6

14.3
42.9
14.3

1
10
1
1

5.6
55.6
5.6
5.6

%

Canada
Alberta
Sub-total

(Canada)

United States
Central

Flyway

South Dakota
Colorado
Kansas
Texas

0
7
0
1

63.6
9.1

1
3
1
0

Sub-total

8

72.7

5

71.5

13

72.4

0
2

1
0

14.3

18.2

1
2

5.6
11.2

2

18.2

1

14.3

3

16.8

1

9.1

0

1

5.6

1

9.1

0

1

5.6

11

100.0

6

17

94.4

Pacific

Flyway

Nevada
Arizona
Sub-total
Mississippi

Flyway

Louisiana
Sub-total
Sub-total

(U. S. )

Total

1/ Includes

11

a limited

number of birds

7

85.7

18

recovered which were banded as immatures.

�-27-

Table 5. Distribution of recoveries
North Park, Colorado 1954.,..69.

Direct
%
No.

Area

from adult female mallards

banded

Indirect
No.
%

Total
%
No.

in

United States
Central Flyway
Wyoming
Colorado
New Mexico
Texas

4
10
1
0

20.0
50.0
5.0

0
13
2
1

72.2
11.1
5.6

4
23
3
1

10.S
60.S
7.9
2.6

Sub-total

lS

7S.0

16

88.9

31

81.S

3

lS.O

2

11.1

5

13.2

3

lS.O

2

11.1

S

13.2

2

10.0

0

2

S.3

2

10.0

0

2

S.3

18

38

Pacific

Flyway

Colorado
Sub-total
Mississippi

Flyway

Louisiana
Sub-Total

20

Total

Recovery

and Hortality

Rates

Recovery rates for mallards are presented by age and sex in Table 7.
Recovery rates for adult male and female mallards banded during pre-season
(August lS-September 20), 1963 through 1969 in the San Luis Valley were
very similar to North Park, being 13.2 percent and 8.3 percent, respectively, in the latter area (unpublished data).
First hunting season recovery rates for adult males banded in North Park
averaged .063 for the 16 year, 19S4 through 1969 banding period and .066
for the 1963 through 1969 period.
Bandings in the San Luis Valley resulted
in similar first year recovery rates, averaging .063 for the 1963 through
1969 period (Table 8).

�Table 6.

Distribution

of recoveries

in Colorado from mallards

banded in North Park.

Adult Male
Direct
% Indirect

%

Local Male
Direct
%
Indirect

%

Adult Female
Direct
%
Indirect

%

Local Female
Direct
% Indirect

%

North Park

5

4.7

2

1.9

0

-

0

-

0

-

0

-

0

-

2

67.7

South Park

4

3.8

0

-

0

-

0

-

0

-

0

-

0

-

0

San Luis Valley

30

28.3

38

35.2

4

23.5

1

12.5

1

7.7

2

13.3

1

16.7

0

West Slope

14

13.2

11

10.2

0

-

2

25.0

3

23.1

2

13.3

0

-

0

North Central

43

40.6

46

42.6

8

47.1

2

25.0

9

69.2

9

60.0

4

66.7

0

Northeast

3

2.8

2

1.9

2

11.8

0

-

0

-

1

6.7

0

-

1

Southeast

.7

6.6

9

8.3

3

17.6

3

37.5

0

-

1

6.7

1

16.7

0

Area

Total

106

I

108

17

8

13

15

6

3

-

33.3

N

Cfj
I

�-29-

Table 7. Recovery rates by sex and age through
for mallards banded in North Park, 1954-69.

the 1969-70 hunting

season

Percent
Recovered

No. Birds
Banded

No. Bands
Recovered

Adult

2,077

281

13.5

Local

240

26

10.8

Immature

39

6

15.4

516

38

8.4

209

14

6.7

47

4

8.5

Sex and Age

Male

Female
Adult

Immature

Mortality rate estimates were calculated by sex for adults and locals for the
entire banding period using the composite dynamic method.
Only computation of
the adult data are presented in table form (Tables 9 and 10). Estimates for
all categories except adult males are based on very small banded samples and
thus are questionable.
However, despite the small samples, mortality rates
calculated for North Park birds are similar to those rates computed by the
same method for birds banded pre-season in the San Luis Valley (Table 11).
Mortality rates for adult males banded in North Park and the San Luis Valley
remain similar regardless of method of calculation or time period.
However,
if all adult male bandings in North Park and the associated recoveries prior
to 1963 are eliminated, the composite dynamic analysis results in somewhat
higher mortality rates (Table 12) than when all banding and recoveries are
used (Table 9). The mortality is slightly higher than that experienced by
San Luis Valley adult males during the same period.
Conversely, by using
the relative recovery rate method of analysis the annual mortality rate of
adult males banded in North Park is slightly lower, .365 (Table 13), than
those males banded during the same period in the San Luis Valley, .386
(Unpublished data). Taking all methods of analyses into account, it is
apparent that these two adult male populations have very similar mortality
characteristics despite having different recovery patterns.

�-30-

Table 8. First hunting season recovery rates of adult male mallards banded
in North Park.

Recovery Rate
North Park San Luis Valley 1/

Year

Number Banded

Number Recovered

1954

40

4

.100

1955

78

2

.026

1956

18

1

.056

1957

7

1

.143

1958

25

1

.040

1959

15

1

.067

1960

15

2

.133

1961

60

0

.000

1962

47

2

.043

1963

115

10

.087

.043

1964

128

6

.047

.091

1965

220

17

.on

.062

1966

646

41

.063

.076

1967

225

13

.058

.058

1968

295

23

.078

.062

1969

143

7

.049

.049

Total

2,On

131

.063

.063

�Table 9. The composite dynamic method of estimating mortality rate of adult male mallards banded in North
Park, 1954-69.

Year of
Banding

1954
1955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969

Number
Birds Banded

40
78
18
7
25
15
15
60
47
115
128
220
646
225
295
143

1

2

4
2
1
1
1
1
2
0
2

2
3
0
0
0
1
1
0
0
7
4
4
19
11
9
0

10
6
17
41
13
23

7

Recoveries b~ Hunting Season
3
6
4
5

1
4
0
0
3
0
0
3
1
4
3
6
18
4
0

1
3
1
0
2
0
0
1
1
0
2
5
11
0

Total
Total
Banded
2,077 Recovered
131
61
47
27
Banded birds availab1e
2,077
1,934
1,639
1,414
Recoveries/1,000 birds banded
63.1
~31.5
28.7
19.1

•

Alive going into period
Mortality rate

164.6

,101.5

.383

7

8

9

0
0
0
0
1
0
0
0
0

0
0
1
0
0
0
0
0
0
1
5
1
0

1
0
0
0
0
1
0
0
0
0
1
0

0
0
0
0
0
0
0
1
0
0
0

0
0
0
0
0
0
0
0
0
0

8
768
10.4

3
548
5.5

1
420
2.4

0
305
0

11.8

6.3

I
W
I-'
I

1
258
3.9;

164.6

3.9;

425.5

101.5
70.0

41.3

22.2
260.9
.389

3.9

.387

�Table 10. The composite dynamic method of estimating mortality of adult female mallards banded in North
Park, 1954-69.

Year of
Banding

Number of
Birds Banded

1954
1955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969

96
l38
12
2
16
12
10
19
4
41
4
14
119
11
15
3

1

2

8

1
3

8
0

0

0
0
0
0
0
0
0
0

0
0
0
0
0

0
3
0
0
1

°4
0
0
0

°0

Recoveries b~ Hunting Season
3
4
5
6
0
2
0
0
0
0
0
0
0
1
0
0
2
0
0

2
0
0
0
0
0
0
0

0
0
0
0
0
0

0

0
0

8

0
0

0

0
1

0

0
0
0
0
0
0

0
0
0
0
0
0
0
0
0

0

°

0

0
0
0

0
0
0

0
0

0

0

1
309
3.2;

77.1

3.2;

169.0

I

W
N
I

0

Total
Total
Banded
516 Recovered 20
Banded birds available
516
Recoveries/1,000 birds banded
38.8

8
513
\15.6

5
498
10.0

2
487
4.1

Alive going into period

,38.3

22.7

12.7

Mortality rate

0
2
0
0
0
0
0
0
0
0

7

77 .1
.503

2
0
368
354
5.4
0
38.3
8.6
3.2
91.9
.417

350
0
3.2

.456

�-33-

Table 11. Mortality rates of North Park and San Luis Valley banded
mallards calculated by the composite dynamic method.

Sex and Age

First Year
N. Park
S.L.V.

All Years
Excluding First
N. Park
S •L .V.

All Years
N. Park
S.L.V.

Males
Adult

.383

.379

.609

II

.404

.387

.394
.488

.411

.511

Immature
Local

.389

.423

.372 1/

.402

.488 1/

.411

.561

.417

.507

.456

.536

Female
Adult

.503

Local

II Includes

.499

.393 1./

classified

as immatures

.607 );./
some birds

.592

.553

.619

Immature

.422

.500 }j

.457

when banded.

Pintails
Distribution

of Recoveries

The Gulf Coast states of Texas and Louisiana are themajor recovery areas for
pintails banded in North Park (Tables 14 and 15). Approximately 29 percent
and 25 percent of all recoveries were taken in Texas and Louisiana, respectively.
The remainder of the recoveries were generally scattered with only
the west coastal area of Mexico and the Sacramento and San Joaquin Valleys
of California worthy of mention.
Low (1949), in analyzing continental pintail banding data concluded that a
"substantial portion" of pintails west of the Mississippi undergo a "roundrobin, counterclockwise migration".
He described this movement as follows:
"originates in the breeding grounds; moves southward via the prairie provinces, ~IDntana, and North Dakota; swings westward into California, continues
south into Mexico; crosses over to the Gulf coast; and returns north in the
spring up through the Central and Mississippi Flyways.
He further stated
that "this flight passes through California in the early fall and reaches
the Gulf coast in November and December".
Ballou (1954) reported that young
of the year pintails banded in Wyoming dispersed widely in their southern
movements while adult birds seemed to follow the routes described by Low
(1949).

�-34-

Table 12. The composite dynamic method of estimating mortality rate of
adult male mallards banded in North Park from 1963 through 1969.
Year of
Banding

Number of
Birds Banded

1

1963

115

10

7

4

0

1964

128

6

4

3

1965

220

17

4

1966

646

41

1967

225

1968
1969
Total Banded

Recoveries bl Hunting Season
2
3
4
5

6

7

1

0

0

2

5

1

6

5

1

19

18

11

13

11

4

295

23

9

143

7

1,772

117

54

35

18

7

1

0

1,772

1,629

1,334

1,109

463

243

115

Banded birds available
Recoveries/l ,000 birds
banded

66.0

,33.1

26.2

16.2
94.7

15.1

4.1

0.0,

160.7

Alive going into period

160.7

\94.7

61.6

35.4
215.0

19.2

4.1

0.0,

375.7

Mortality rate

.411

.440

.428

�-35-

Table 13. The relative recovery rate method of estimating the mortality
rate of adult male mallards banded in North Park from 1963 through 1969.

Year
Banded

Number
Banded

Number
l-n

1963

115

CD

12

1964

128

21

15

1965

220

33

1966

646

1967

Recoveries
2-n

Recovery
l-n

Rate
2-n

Survival
Rate

.1043

.636

.364

.1641

.1172

.781

.219

16

.1500

.0727

.528

.472

89

48

.1378

.0743

.597

.403

225

28

15

.1244

.0667

.615

.385

1968

295

32

9

.1085

.0305

.622

.378

1969

143

7

CD

.0490
.7338

.4657

Survival

rate

.635

.630

Mortality

rate

.365

.370

Recoveries from North Park pintails, banded during their moult, do not for
the most part, seem to fit the same patterns as those discussed by Low (1949)
and Ballou (1954).
For classification, only 2 of 118 recoveries of North
Park banded birds were recovered directly as immatures.
The counter-clockwise
migration for the major portion of the North Park population is not adapted
by this writer for the following reasons: (1) More than 50 percent of the
total recoveries corne from Texas and Louisiana; California only accounts for
10 to 15 percent.
This recovery proportion is much different than that
expressed for most counter-clockwise migrating populations.
(2) Eight of 12
birds recovered in Mexico were taken after January 1, two in December and two
in September.
All but four recoveries were taken along the west coast; two
in the inland state of Durango in February and March; one in the state of
Mexico in January and one in the east coast state of Vera Cruz in February.
No late fall west to east movement through Mexico is indicated by these
recoveries.
(3) The pattern of direct, as well as indirect recoveries from
North Park pintails indicates a general southeast movement through Colorado
and the Texas p.anhandle into Gulf coastal areas.

�-36-

14.

Table
Park,

Distribution
of
Colorado,
1954-69.

recoveries

from male pintails

Direct
Area

banded

in North

Indirect
No.

%

No.

%

0

2

3.4

2

2.2

0

2

3.4

2

2.2

1
1
2
0
17
2

1.7
1.7
3.4
28.8
3.4

1
3
2
1
25
2

1.1
3.4
2.2
1.1
28.1
2.2

No.

%

Total

Canada
Alberta
Sub-total
United

(Canada)

States

Central

Flyway

South Dakota
Colorado
Kansas
New Mexico
Texas
North Dakota

0
2
0
1
8
0

Sub-Total

11

36.7

23

39.0

34

38.1

0
1
2
1

3.3
6.7
3.3

1
3
6
0

1.7
5.1
10.2

1
4
8
1

1.1
4.5
9.0
1.1

4

13.3

10

17.0

14

15.7

1
0
9
0

3.3
30.0

0
1
15
1

1.7
25.4
1.7

1
1
24
1

1.1
1.1
27.0
1.1

33.3

17

28.8

27

30.3

0

1

1.7

1

1.1

0

1

1.7

1

1.1

6

10.2

11

12.4

Pacific

6.7
3.3
26.7

Flyway

Montana
Utah
California
Arizona
Sub-Total
Mississippi

Flyway

Missouri
Tennessee
Louisiana
Arkansas

10

Sub-Total
Atlantic

Flyway

North

Carolina

Sub-Total

(U.S.)

Hexi co

5

Total

30

16.7

59

89

�-37-

Table 15. Distribution of recoveries from female pintails banded in North
Park, Colorado, 1954-69.

No.

%

No.

%

0

1

4.8

1

3.5

0

1

4.8

1

3.5

No.

Area

Total

Direct

Direct
%

Canada
Saskatchewan
Sub-Total (Canada)
United States
Central Flyway
Colorado
Texas
Kansas
Sub-Total

0
4
0

50.0

2
5
1

9.5
23.8
4.8

2
9
1

6.9
31.0
3.5

4

50.0

8

38.1

12

41.4

2
2
2
1

9.5
9.5
9.5
4.8

2
4
2
1

6.9
13.8
6.9
3.4

Pacific Flyway
Utah
California
Arizona
Washington

0
2
0
0

25.0

2

25.0

7

33.3

9

31.0

0
2

25.0

1
3

4.8
14.3

1
5

3.4
17.2

Sub-Total

2

25.0

4

19.1

6

20.7

Sub-Total (U.S.)

8

100.0

20

95.3

28

96.6

4.8

1

3.4

Sub-Total
Mississippi Flyway
Arkansas
Louisiana

Mexico

0

1

Total

8

21

29

�-38-

A portion of the North Park population may very well follow the counterclockwise migration.
The birds recovered in MeXico probably arrive there
by way of California and may migrate north through the Central Flyway in the
spring.
However, it is doubtful, based on the information available that
the birds recovered on the Gulf coast arrived there via California.
Recovery

and Mortality

Rates

Recovery rates for pint ails banded in North Park (Table 16) are much lower
than those previously presented for the mallards (Table 7). Again, as in
the mallard, the pintail data are strongly orierited to adults banded during
the moult.

Table 16. Recovery
for pintails banded

rates by sex and age through
in North Park 1954-69.

the 1969-70 hunting

season

No. Birds
Banded

No. Birds
Recovered

Percent
Recovered

Adult

2,287

87

3.8

Local

55

4

7.3

Immature

3

o

0.0

Adult

965

23

2.4

Local

72

4

5.6

Immature

8

2

25.0

Sex and Age

Male

Female

First hunting season recovery rates for adult males in years which had more
than 300 birds banded varied from .009 to .019 and averaged .013 for all
bandings.
Adult males banded during the pre-season period in the San Luis
Valley from 1962 through 1969 had a .020 first year recovery rate (unpublished data).
Calculations of mortality rate~ for pintails may not be reliable because
of variability in the banded sample and unusual distribution of recoveries
through the years. Overall rates for adult males and females calculated by
the composite dynanuc method are .347 and .331, respectively (Tables 17
and 18). Using only data since 1961 which contained generally larger annual

�-39-

banded samples, particularly for adult males, the mortality rates calculated
by the composite method are. 359 and .303 for males and females, respectively.
Pint ails banded during the August 15 through September 20 period in the San
Luis Valley exhibited mortality rates calculated by the same method of .405
and .436 for males and females respectively.
An estimate of .227 was calculated for adult males using the relative recovery rate method for the 1961
through 1969 data. Unrealistic mortality estimates resulted for adult
females when using either the normal or "pooled recovery" (Geis et a1. 1971)
relative recovery rate methods.
Because of inadequate banded samples,
mortality rates for immatures.

Green-winged
Distribution

no attempt was made to estimate

annual

Teal

of Recoveries

Adult male green-winged teal recoveries showed a general scattering from
North Park banding sites with California and Texas the major recovery states
(Table 19). Most of the Texas and California recoveries come from locations
designated by Moisan (1967) as major wintering areas for the continental
green-winged population.
Birds migrating into Texas apparently move southeast through the Texas
panhandle, generally terminating migration in the gulf coastal areas of
Texas and Louisiana.
There are two major recovery areas in California, the ImperiaL Valley in
the extreme southern part of the state and the San Joaquin Valley of central
California.
Direct band recoveries present no clear-cut migration route
into California.
Recoveries suggest one· possible route through northern
Utah and another directly south from North Park down through New Mexico and
across Arizona into the Imperial Valley area.
Moisan (1967) suggested that
birds moving through the Salt Lake Valley of Utah fly not only to the central Valley (San Joaquin Valley area) of California but also straight south
to the Imperial Valley.
The only significant variation in direct and indirect recoveries occurs within
the state of California.
Twenty of the 36 California recoveries were
recovered in two specific latitude and longitude degree blocks.
One block
in the San Joaquin Valley and one in the Imperial Valley.
Six of the eight
total recoveries in the Imperial Valley block were taken directly while only
three of the twelve recoveries in the San Joaquin block were taken during
the first year after banding.
These data seem to indicate a somewhat
stable migration pattern by North Park adult male green-wings into the San
Joaquin block.
Along these same lines it is interesting to note that
Moisan's (1967) data indicated that no gr~en-winged teal banded in the central
Valley of California during the winter were subsequently recovered in the
Imperial Valley, whereas large numbers of birds winter banded in the Imperial Valley were subsequently recovered in the Central Valley.

�-40-

Table 17.
The composite dynamic method of estimating
male pint ails banded in North Park, 1954-69.
Year of
Banding

Number of
Birds Banded

1

4
4
5
3
44
57
15
523
101
49
99
92
593
345
312
41

0
0
0
0
0
0
0
5
0
0
1
1
11
5
6
1

1954
1955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969

Total
Banded

Total Re2,287 covered
30

Banded birds

available

Recoveries/I,
banded

000 birds

Alive

going into

Mortality

rate

period

2,287

Recoveries b~ Hunting
2
3
4

mortality

rate

Season
5

6

7

of adult

0
0
0
0
2
2
0
2
1
0
0
3
6
2
2

0
0
0
0
1
0
0
3
2
1
1
0
3
2

0
0
0
0
0
0
0
8
0
1
0
2
2

0
0
0
0
0
1
0
3
1
0
0
0

0
0
0
0
0
0
0
2
0
0
0

0
0
0
0
0
1
0
0
0
0

20

13

13

5

2

1

2,246

1,934

1,589

996

904

805

13.1

\8.9

6.7

8.2
32.2

5.0

2.2

1. 2,

45.3

\32.2

23.3

16.6
85.1

8.4

3.4

1. 2{ 130.4

.289

.378

45.3

.347

�-41-

Table 18. The compsite dynamic method of estimating mortality rate of
adult female pintai1s banded in North Park, 1954-69.
Year of
Banding

Number of
Bi rds Banded

1.
0
0
0
1
'0
0

1954
1955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969

11
14
7
4
21
55
21
177
57
69
33
30
331
45
·84
6

Total
Banded

Total Re965 covered
6

Banded birds available

Recoveries b~ Hunting Season
2
3
4
5

1

0
0
0
0
0

0
0
0
0
0
0
0
1
0
0
0
0
1
0

0
0
0
0
0
1
0
4
0
0
0
0
0

7

2

5

2

1

959

875

830

499

469

0

1
0
0
1
0
0
4
0
1

0

0
1
1
0
1
0
0
1

965

6

0
0
0
0
0
0
0

0
0
2
0
0

0

0
0
0
0

0
0
1
0
0
0

Recoveries/1,000 birds
banded

6.2

\7.3

2.3

6.0
21.7

4.0

2.1 I

27.9

Alive going into period

27.9

,21.7

14.4

12.1
56.4

6.1

2.1

I

84.3

Mortality rate

.222

.385

.331

�-42-

Table 19. Distribution of recoveries from adult male green-winged
banded in North Park, Colorado, 1954-69.

Direct

teal

Indirect
No.
%

No.

%

2.1
12.8
4.3
21.3

0
4
1
18

7.5
1.9
34.0

1
10
3
28

1.0
10.0
3.0
28.0

19

40.5

23

43.4

42

42.0

1
4
15
2

2.1
8.5
31.9
4.3

0
1
21
3

1.9
39.6
5.7

1
5
36
5

1.0
5.0
36.0
5.0

22

46.8

25

47.2

47

47.0

1
1
1
0
1

2.1
2.1
2.1
2.1

0
0
0
1
3

1.9
5.7

1
1
1
1
4

1.0
1.0
1.0
1.0
4.0

Sub-Total

4

8.4

4

7.5

8

8.0

Total (U.S.)

45

95.7

52

98.1

97

97.0

Mexico

2

4.3

1

1.9

3

3.0

Total

47

Area

No.

%

1
6
2
10

Total

United States
Central Flyway
Nebraska
Colorado
New Mexico
Texas
Sub-Total
Pacific

Flyway

Colorado
Utah
California
Arizona
Sub-Total
Hississippi

Flyway

Michigan
Minnesota
Missouri
Arkansas
Louisiana

53

100

�-43Recovery

Rates and Mortality

Banding efforts in North Park resulted in bands being placed on primarily,
adult males (Table 20). The recovery rate for adult males and females
banded in North Park is slightly higher for the same yearly period of
banding than the 2.3 and 1.2 percent recorded for males and females, respectively, which were banded in the San Luis Valley.
.

Table 20. Recovery rate by sex and age through the 1969-70 hunting
for green-winged teal banded in North Park 1954-69.

season

No. Birds
Banded

No. Birds
Recovered

Percent
Recovered

Adult

3,314

104

3.1

Local

43

o

0.0

Immature

o

o

Adult

276

6

2.2

Local

54

o

0.0

Immature

o

o

Sex and Age

Male

Female

Adult males banded in North Park exhibited a first year recovery rate of .013.
First year recovery rates calculated for adult males banded during the summer
in specific continental reference areas varied from .022 in Alberta to .121
in the Imperial Valley of California (Moisan 1967).
The Colorado-Wyoming
reference area, which includes North Park and is excluded from the above
reference, has a first year recovery rate of .008, the lowest rate for any
of the continental summer reference areas designated by Moisan (1967).
Moisan (1967) reported that adults banded in the Colorado-Wyoming
sun®er
reference area had a kill rate, adjusted for band reporting rate and crippling
loss of .048, the lowest rate for all reference areas.
Applying ~bisan's
band-reporting rate and crippling loss correction to North Park adult males
results in an average annual kill rate of .044. Combining the adult males
with the meager adult female data results in the same kill rate for the adult
green-wing population.
The annual mortality rate calculated by the composite dynamic method for
adult male green-winged teal summer banded in North Park is .383 (Table 21).

�-44-

Table 21.
The composite dynamic method of estimating
male green-winged teal banded in North Park, 1954-67.
Year of
Banding

rate of adult

Number of
Birds Banded

1

0
27
1
28
63
60
36
260
207
283
270
98
967
392
491
131

0
0
0
0
1
1
0
2
1
0
3
1
18
6
6
5

0
1
0
0
0
0
0
0
0
5
5
2
7
8
0
0

0
0
0
2
1
1
0
0
3
1
1
0
5
1
0

0
0
0
0
0
0
0
2
0
0
0
0
4
0

0
0
0
0
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
0
2
0
0
0
0

0
0
0
0
1
0
0
1
0
0
0

Total Recovered
44

28

15

6

0

2

2

2,692

2,300

1,333

1,235

965

682

1954
1955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
Total
Banded

3,314

Banded birds

available

Recoveries/l,OOO
banded

rate

3,183

Recoveries
2
3

by Hunting Season
4
5
6

7

birds

Alive goins into period
Mortality

mortality

13.8

,10.4

6.5

4.5
26.4

0.0

2.1

2.:..2.t

40.2

40.2

\ 26.4

16.0

9.5
64.8

5.0

5.0

2.91

105.0

.343

.407

Average annual mortality rates for adult males banded in 16 specific reference areas throughout the continent and calculated by the composite methods
ranged from .454 for summer banded birds in Alberta to .693 for summer banded
birds in the Imperial Valley of California.
Moisan (1967) found that average
annual mortality rates for summer banded populations varied directly with
average first year recovery rates and thus, kill rates.
As a brief summary, adult male green-winged teal apparently undergo one of
the lowest rates of harvest for green-winged populations in North America.
The low rate of harvest results in a low annual mortality rate, theoretically
lower mortality than would be expected on the average, for a population
undergoing no hunting mortality.

.393

�-45-

American Widgeon
Smith (1949) in analyzing continental band recoveries of American widgeon
defined two major migration routes.
One route moved southward through the
Pacific states and inter-mountain areas, while a second flight moved through
the interior portion of the Central Fl.yway, According to the recoveries
of widgeon banded in North Park, most birds move into the Pacific Flyway
area in the fall with a substantial number of birds recovered in Mexico
(Table 22).
The recoveries indicate two possible migrational paths.
One route has birds
moving into northenlUtah,
then southwesterly into the San Joaquin Valley
of central California or into the Imperial Valley or coastal area of extreme
southern California.
The other route shows birds moving generally south
through Colorado, New Mexico, and into southeastern Arizona.
The latter
route is most likely followed by birds which winter in Mexico.
The majority
of the recoveries in Mexico are reported taken in the coastal state of
Sinaloa with the adjacent inland state of Durango being the area next in
importance.
Recovery

and Mortality

Rates

The banded sample of widgeon, unlike the other species, included a good
percentage of birds banded as locals (Table 23). Males and females were
recovered at essentially the same rate. Locals were recovered at a greater
rate than adults.
The first year recovery rate for locals was .030 compared
to .011 for adults.
Annual mortality rates were calculated by the compo sLte dynamic method for
both sexes of locals and adult males .. Widgeons banded as local males had an
overall mortality rate of .615 compared to .516 for adult males. Local
females exhibited a mortality rate of .536.
Conclusion
The population of mallards banded in North Park is contributing birds to
the harvest throughout the state of Colorado.
Annual indicated mortality
rates are in most cases equal to, or greater than populations of mallards
banded in the San Luis Valley.
These mortality rates have been attained
without the North Park population experiencing early season hunting pressure
in the immediate banding area, as has the San Luis Valley population.
Therefore, based on this analysis, North Park mallard should not be subjected to any additional hunting pressure at this time without a complete
evaluation of the significance additional hunting mortality would have on
the population.
North Park pintails are currently subject to comparatively low first year
recovery rates and annual mortality rates. According to this analysis,
additional hunting pressure leading to increased mortality rates should
not be a detriment to the North Park pintail population.

�-46-

Table 22. Distribution
of recoveries
Park, Colo.rado, 1954-69.

Direct

from American widgeon banded in North

Indirect
No.
%

No.

%

3.6
10.7
3.6
7.1
3.6

0
2
0
3
0

0.0
6.7
0.0
10.0
0.0

1
5
1
5
1

1.7
8.6
1.7
8.6
1.7

8

28.6

5

16.7

13

22.4

2
0
3
2
4

7.1
0.0
10.7
7.1
14.3

1
2
3
3
3

3.3
6.7
10.0
10.0
10.0

3
2
6
5
7

5.2
3.4
10.3
8.6
12.1

11

39.3

12

40.0

23

39.7

0
0
1

0.0
0.0
3.6

1
1
2

3.3
3.3
6.7

1
1
3

1.7
1.7
5.2

Sub-Total

1

3.6

4

13.3

5

8.6

Sub-Total
(U.S.)

20

71.4

21

70.0

41

70.7

Mexico

8

28.6

9

30.0

17

29.3

Total

2.8

Area

No.

%

Nebraska
Colorado
.Oklahoma
New Mexico
Texas

1
3
1
2
1

Sub-Total

Total

United States
Central

Pacific

Flyway

Flyway

Colorado
Oregon
Utah
Arizona
California
Sub-Total
~lississippi

Flyway

Minnesota
Tennessee
Louisiana

30

58

�-47-

Table 23. Recovery rates by sex and age through the 1969-70 hunting
for American widgeon banded in North Park, 1954-69.

season

No. Birds
Banded

No. Birds
Recovered

Percent
Recovered

Adult

818

25

3.1

Local

310

17

5.5

Sex and Age

Male

Immature

0

Female
Adult

175

6

3.4

Local

251

14

5.6

3

0

0.0

Immature

Green-winged teal populations of North Park are currently undergoing one of
the Jawest harvest rates for green-winged populations in North America.
Additional hunting pressure on North Park green-winged teal should not be
detrimental to the population.
Throughout this analysis it has been noted that small banded samples of
some species and disproportionate banded samples by sex and age of others
have hampered attempts to determine various characteristics of the North
Park populations.
The methods and time period of trapping used throughout
the banding period have been selective toward various species and age groups
within species.
These bandings and the subsequent analyses have resulted in
some valuable information.
However, I conclude that the population characteristics obtained from these data should only be considered a starting
point.
The information is, for the most part, barely adequate for making
basic management decisions on some species and totally inadequate for the
modern day decisions involving species management~
If North Park duck populations are considered vital to waterfowl management
in Colorado, then banding programs designed to obtain specific objectives
should be initiated and carried out.

�-48-

LITERATURE

CITED

Ballou, R. M. 1954. Live-trapping and banding of waterfowl.
Wyo. Game
and Fish Comm. Fed. Aid in Wildl. Restoration Quart. Rep. W-50-R.
p , 28-38.
Geis, A.. D., R. I. Smith, and J. P. Rogers.
harvest characteristics, and survival.
Spec. Sci. Rep. Wildl. No. 139.

1971. Black duck distribution,
U. S. Fish and Wildl. Servo

Low, S. H. 1949. The migration of the pintail.
In: Migration of some
North American waterfowl.
U. S. Fish and Wildl. Servo Spec. Sci.
Rep. Wildl. No. 1:13-16.
Moisan, G. 1967. The green-winged teal: it's distribution, migration, and
population dynamics.
U. S. Fish and Wildl. Serv., Spec. Sci. Rep.
Wildl. No. 100.
Smith, A. G. 1949. Migration
North American waterfowl.
Rep. Wildl. No 1:11-12.

Prepared

by

of the baldpate.
In: Migration of some
U. S. Fish and Wildl. Servo Spec. Sci.

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Wildlife

Researcher

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�-50-

APPENDIX

Scientific

and common names of birds mentioned

Common Name

in this report.

Scientific

Name

Canvasback

Aythya

Coot, American

Fulica americana

Gadwall

Anas strepera

Mallard

Anas platyrhynchos

Merganser,

Common

valisineria

Hergus merganser

Pintail

Anas acuta

Redhead

Aythya

Ring-necked

Duck

americana

Aythya collaris

Scaup, Lesser

Aythya affinis

Shoveler

Spatula elypeata

Teal, Bl ue-winged

An as discors

Cinnamon

Anas cyanoptera

Green-winged

Anas carolinensis

Widgeon,

American

Mareca americana

�-51October,

1971

JOB PROGRESS REPORT
State of

~C~O~L~O~RAD~~O

Project No.

W-88-R-16

Work Plan No.

1

Job Title
Period Covered:

_
Migratory

Personnel:
Velma Merkle,
Howard D. Funk.

5

Job No.
Waterfowl

October

Bird Investigations

Kill Survey

1, 1970 through July 1, 1971
Velma Fredrickson,

Raymond

Boyd, Robert

Tully and

ABSTRACT
Duck stamps sold in Colorado in 1970 numbered 37,972, a substantial increase
over the 34,281 sold in 1969, but still not of the n~gnitude of 1957 or
1958 when sales reached almost 42,000.
Even with relaxation of season
length and bag limits, estimated number of duck hunters on the east slope
increased only slightly over the 1969 level to 27,410 participants.
Regular
duck season was 90 days with harvest limits based on a point-system, with a
limit of 100+ points, allowing from 2 to 10 and from 4 to 20 ducks in the
daily bag and in possession, respectively.
Point allocations for male and
female mallards were 20 and 90 points, respectively, allowing from 2 to 5
daily and from 4 to 10 in possession if mallards were the only species taken.
Regular season east slope harvest was estimated at about 207,000 ducks, an
increase of almost 27,000 over the 1969-70 harvest and again the highest
since the 1958-59 season.
Duck harvest and hunting pressure on the west
slope both increased somewhat over the 1969 levels with almost 27,700
birds
taken by about 3,900 hunters.
About 2,900 hunters took part in the final
experimental San Luis Valley hunting season, also on a point-system, and
bagged an estimated 23,000 ducks.
Harvest during the special early teal
season was estimated at about 5,600 ducks by almost 1,200 hunters in specified east slope areas.
Species composition estimates for all seasons
combined throughout the State again placed the mallard as the most important
species at about 73 percent of the total harvest.
The number of east slope goose hunters increased to over 18,000 in 1970-71,
an increase of about 5,000 over the previous year. Much of this increase
was due to about 11,000 hunters taking part in the north-central Colorado
goose season as compared to only about 6,000 in 1969-70.
However, estimates
by area suggested that goose hunting pressure also increased in the Arkansas
Valley from approximately 5,300 hunters a year ago to almost 7,900 this year.
Numbers of geese present during hunting season were at all-time highs in both
the Arkansas Valley and north-centaal areas and harvest for each area also
peaked in 1970-71 with about 40,000 geese taken between the two. West slope
hunter participation increased about 28 percent with 375 hunters taking about
770 geese, an increase of about 200 percent.

��-53-

WATERFOWL

KILL SURVEY

Howard D. Funk'

This survey of Colorado small game harvest is a cooperative effort between
personnel of Federal Aid Projects W-88-R and W-38-R, and the Game Management Section of the Colorado Division of Game, Fish and Parks.
Most data presented herein result from two separate major surveys:(l) the
regular annual questionnaire survey limited to purchasers of combination
small game hunting and fishing, resident small game hunting, and non-resident
small game hunting licenses; and (2) the separate questionnaire survey of
holders of the increasingly popular sportsman's license which also allows
small game hunting.
Other data contributing to this report are gathered
from other Federal Aid surveys specific to a particular season such as the
special north-central Colorado or San Luis Valley goose seasons, all of which
require a permit to hunt.

P. S. OBJECTIVE
To formulate waterfowl

harvest

regulations.

SEGMENT OBJECTIVES
To estimate
by species,

the State harvest of waterfowl for the 1970-71 hunting
county and interval of the season.

METHODS

season

AND MATERIALS

Techniques utilized were similar to those employed in the past with hunter's
names drawn in a mechanical random fashion from duplicate license sales
stubs.
The cutoff dates for license sales figures used in projections were
March 31, 1971 for the regular survey and March 1, 1971 for the sportsman's
license survey.
One follow-up letter was sent to non-reporting hunters
after an interval of approximately three weeks.
A survey questionnaire
similar to the revised form utilized for the 1969 survey was again used in
an attempt to simplify reporting by hunters and ultimately obtain higher
reporting rates and more valid harvest information than in past years.
The sportsman's questionnaire was again somewhat more simplified than the
regular survey form. Harvest data collected on this survey under Federal
Aid Project W-38-R (Boyd 1971) were added to results of the regular survey.

�-54-

RESULTS

AND DISCUSSION

Regular survey questionnaires were sent to 12,518 randomly selected 1970
license buyers about January 15, 1971. A total of 7,448 responded for a
return of 59.5 percent.
This was the lowest response rate since the survey
was initiated, even with the more simplified questionnaire.
Of the 7,448
returns, 3,953 reported hunting, 3,364 reported they did not hunt, and 131
were not usable.
Percentages of license buyers actually hunting were calculated from the survey sample by license type (Table 1). The largest percentage of license buyers not utilizing the opportunity to hunt was found
in the combination hunting and fishing license category; these people using
their fishing privilege only. Total sales of regular type licenses
permitting small game hunting during 1970, other than sportsman's licenses,
amounted to 165,322.
It is estimated that 96,158 of these actually hunted
one or more species of small game, including waterfowl.

Table 1. License sales, percent
license type, 1970 season.

License Type

Resident Combination
Resident Small Game
Non-resident Small Game

Sub-total
Sportsman's License
Resident
Non-resident

Sub-total

TOTAL

use and projected

estimates

of hunters by

Sales

Percent Used for
Hunting Small Game

Projected
Number Hunters

121,716
40,830
2,776

49.8
81.6
80.2

60,615
33,317
2,226

165,322

15,404
7,897

96,158

64.3
22.3

9,905
1,761

23,301

11,666

188,623

107,824

Survey questionnaires were sent to 4,140 sportsman's license buyers.
A
total of 2,375 responded and supplied usable information for a response rate
of 57 percent.
Of the 2,375 returns, 1,399 reported hunting small game,
while 976 reported they did not.
Percentages of respondents indicating
they hunted were calculated separately for residents and non-residents.
Thus,

�-55-

of the total sale of sportsman's licenses in 1970 (23,301), it is estimated
that 11,666 of the buyers of this type of license took part in hunting one
or more species of small game, including waterfowl (Table 1).
Total estimated number of regular and sportsman's license buyers hunting
one or more species of small game in 1970 was 107,824 which compares with
105,347 for 1969, 134,606 for 1968, and 98,618 in 1967.
Colorado duck stamp sales are listed in Table 2. Numbers sold in 1970
(37,972) indicate the continuing annual increase in waterfowl hunters from
the low of approximately 18,000 sold in 1962. However, sales still are not
of the magnitude experienced in 1957 and 1958 when almost 42,000 were sold
during the most recent peak period of waterfowl populations.
Table 2.

Duck stamp sales for Colorado, 1954-1970.

Year

Number of
Stamps Sold

Percent Change From
Previous Year

1954
1955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970

32,450
39,107
36,303
41,794
41,897
31,431
30,592
24,854
17,701
22,940
25,282
20,537
29,377
31,064
31,218
34,281
37,972

+ 20.5
+ 20.5
7.2
+ 15.1
+
.02
- 24.9
2.7
- 18.8
- 28.8
+ 29.6
+ 10.2
- 18.8
+ 43.0
+ 5.7

+
.5
+ 9.8
+ 10.8

Duck stamp buyers are classified in Table 3 by the type of waterfowl hunting
in which they participated in 1970 and the previous 16 years. There was a
slight increase in numbers of duck hunters, the highest since 1958, while
estimated goose hunters increased about 5,500 for an all time high. As
will be discussed later, much of the increase in goose hunting activity
took place in north-central Colorado.

�-56-

Table 3. Estimated number of duck and goose hunters, average number of days
hunted, season length, and bag and possession limits, by year, 1954-1970.

Year

Number
of Duck
Hunters

Number
of Goose
Hunters

Average Number
Days Hunted For
Ducks
Geese

1954
1955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970

31,834
37,816
34,793
37,166
38,773
29,060
29,480
22,920
13,918
17,989
19,189
15,374
23,635
25,347
26,528
30,436
31,326

12,136
17,634
12,477
12,057
14,705
13 ,647
14,107
11 ,245
9,159
10,841
13,678
11 ,344
15,807
13,748
13 ,467
13,282
18,774

7.64
8.87
7.37
6.52
5.78
5.70
6.05
4.40
5.27
5.66
6.30
5.62
5.96
6.49
6.18
5.94
6.54

Season Length
{Days}
Ducks
Geese
60
75
75
75
90
50
60
30
25 ]j

6.64
7.37
5.53
6.25
6.43
6.48
6.45
6.74

35

II

36

II

33

II

40 11
60 11
60 11
33 11
90 1./

60
60
60
60
60
75
75
60
75
75
75
75
75
75
75
75
79

Bag and
Possession
Limit
Ducks
Geese
5-10
5-10
5-10
5-10
4- 8
4- 8
3- 6
3- 6
2- 4 41
4- 8 21
4- 8 21
4- 8 31
3- 6 21
3- 6 31
3- 6 21
4- 8 21

"fl

2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2

2-4 51

2-4 1./

II Hunting regulations for East Slope.

West Slope had a general season of 75
days with 4 ducks in bag, and 8 in possession in 1962; and 90 days, 4 in bag
and 8 in possession in 1963; 1964 and 1965; and 90 days,S in bag and 10 in
possession in 1966 and 1967; and 86 days,S in bag and 10 in possession of which
3 and 6, respectively, could be mallards in 1968; and 86 days,S in bag and 10
in possession in 1969; and 93 days, 6 in bag and 12 in possession in 1970.

11 Two mallards allowed in bag, and four in possession on East Slope.
11 One mallard and pintail allowed in bag and two in possession on East Slope,
three mallards or three pintails on West Slope.

il One mallard in bag and two in possession on East Slope.
il One goose per day and two in possession on West Slope.
~I Point-system allowing from two to ten ducks daily and four to twenty
possession.

in

See text.

21 One goose per day and two in possession on West Slope except for Brown's
Park where both bag and possession were two birds.

�-57-

Duck Harvest

Duck hunting statistics for the 1970 season are tabulated and compared
with results of past years in Table 4. Total retrieved harvest of 234,514
ducks was significantly greater than in 1969, an increase of approximately
15 percent, and the largest harvest since 1958. Increased numbers of
hunters, relaxed hunting seasons, favorable publicity, and special seasons
offering a variety of hunting opportunity were probably all responsible for
this increase.
The estimate on wounding losses obtained through the questionnaire this year was 15 percent of the total bag or about 41,000 birds.
An estimate of species composition in the harvest was obtained from the
Bureau of Sport Fisheries and Wildlife wing collection survey for the
regular season and is presented in Table 5 along with data from the previous
15 years, 1969 excluded.
These data show emphasis in 1970 was again oriented
strongly toward the,mallard and that total duck harvest was about 61 percent greater than the fifteen-year average.
Duck harvest by intervals and season types is illustrated in Table 6 for
both the east and west slopes of the State, the Central and Pacific Waterfowl Flyways, respectively.
Regular seasons will be discussed here with
special season results in following sections.
The regular season on the
east slope was governed by a point-system regulation which has been tested
in the past in early October San Luis Valley seasons and several states in
the Central Flyway during late winter periods.
The Bureau of Sport Fisheries
and Wildlife granted Colorado and seven other Central Flyway states, and
two states in each of the Mississippi and Atlantic Flyways the opportunity
to further test the regulation on a broad scale during the regular season
period.
Thus, Colorado was allowed a 90-day season based on a 100-point
daily bag regulation.
Hen mallards, wood ducks, redheads, canvasbacks, and
hooded mergansers plus a few other species of ducks uncommon in the State
were assigned 90 points per bird while drake mallards, green-winged teal,
hen pintails, ringneck and mottled ducks had a point value of 20. All
other species and sexes not mentioned above were designated as 10-point
birds.
The daily bag limit was reached when the points for the last bird
taken caused the accumulated points for all birds bagged to reach or exceed
100 points.
Possession limit was two legal daily bag limits.
Thus, bag
limits could range from two to ten ducks daily, depending on the skill and
desire of the hunter to identify birds in flight.
Numbers of hunters, while the greatest since 1958, did not increase to any
great degree during the regular season over that of 1969, even with the
long season.
Harvest was indicated to have increased 60 percent over 1969
from about 111,000 birds to 178,000 for 1970 with greatest pressure and
harvest occurring during the October 3l-November 20 period.
However,
harvest was indicated to have been fairly equal throughout the season even
though the majority of ducks seemed to migrate south later than usual.
Further reports on the point-system season in Colorado and other states,
complete with hunter performance survey information and harvest estimates,
will be forthcoming from the Bureau of Sport Fisheries and Wildlife later
in 1971. Preliminary results indicated the majority of hunters conducted
themselves well under the point-system regulation and pressure was placed
on the mallard drake and other species and sexes which could stand additional harvest, and that pressure was relaxed on high point birds such as
the mallard hen.

�-58Table 4.

Duck harvest statistics, 1954-1970.

Hunters

Average
Seasonal
Bag

Total
Estimated
Harvest

Wounding Loss
Percent
Numbe r

Total
Estimated
Hunting
Mortality

31,834
37,816
34,793
37,166
38,773
29,060
29,480
22,920

5.6
6.7
5.9
6.8
6.1
4.2
5.0
3.8

179,856
253,367
185,737
254,587
236,515
122,924
147,400
86,408

14.5
13.1
16.3
14.1
12.3
15.5
13 .1
21.4

30,396
38,182
36,195
41,679
33,088
22,417
22,257
23,608

210,252
29l ,549
221,932
296,266
269,603
145,341
169,667
110,016

Numbe r

of

Date

Slope

1954
1955
1956
1957
1958
1959
1960
1961
1962

East
West
TOTAL

11,349
2,569
13,918

2.6
3.5
2.8

29,507
8,992
38,499

13.5
11.7
13 .1

4,603
1,187
5,790

34,110
10,179
44,289

1963

East
West
TOTAL

15,627
22362
17 ,989

5.1
3.6
4.9

80,167
88,670

10.7
9.7
10.6

9,636
916
10,552

89,803
92419
99,222

1964

East
West
TOTAL

16,311
2,878
19,189

4.0
4.0
4.0

65,244
11 2512
76,756

10.6
10.4
10.6

7,764
1,335
9,099

73,008
12,847
85,855

1965

East
West
TOTAL

12, 747
22627
15,374

5.0
4.3
4.9

64,245
11 2322
75,567

18.7
15.4
18.2

12 ,007
12744
13,751

76,252
13 ,066
89,318

1966

East
West
TOTAL

19,494
42141
23,635

5.0
4.4
4.9

97,860
182345
116,205

16.4
20.4
17.1

19,278
42715
23,993

117,138
232°60
140,198

1967

East
West
TOTAL

22,432
2,9l5
25,347

6.5
3.7
6.2

146,032
10,698
156,730

12.0
22.9
12.9

20,006
3,178
23,184

166,038
13 ,876
179,914

1968

East
West
TOTAL

23,006
32522
26,528

6.2
7.4
6.4

142,578
252954
168,532

10.9
10.9
10.9

17,466
3,179
20,645

160,044
29,133
189,177

1969

East
West
TOTAL

27,058
32378
30,436

6.7
7.0
6.7

179,848
232781
203,629

- ]}

- ]}

East
West
TOTAL

27,410
32916
31,326

7.5
7.1
7.5

206,841
272673
234,514

36,181
52169
41,350

243,022
322842
275,864

1970

81503

1./ Data not requested in survey questionnaire.

- ]}
14.9
15.7
15.0

�-59-

Table 5.

Species composition of the bag, 1954-70. 1/

1970

15-year Average
1954-1968
NlUDber
Percent
Harvested
of Total

Percent
Change 1970
from 15-year
Average Harvest

Number
Harvested

Percent
of Total

Mallard

172,133

73.4

99,840

68.5

+

Pintail

5,863

2.5

4,941

3.4

+ 18.7

Green-winged Teal

12,195

5.2

12,622

8.7

3.4

938

0.4

5,509

3.8

83.0

Baldpate

13,602

5.8

1,720

1.2

+ 690.8

Gadwall

16,650

7.1

4,279

2.9

+ 289.1

Shoveler

3,283

1.4

1,615

1.1

+ 103.3

Scaup

704

0.3

1,139

0.8

38.2

Redhead

3,752

1.6

1,154

0.8

+ 225.1

938

0.4

1,062

0.7

11.7

4,456

1.9

11,821

8.1

62.3

234,514

100.0

145,702

100.0

+ 61.0

Species

Blue-winged and
Cinnamon Teal

Canvasback
Others and Unknown
TOTAL

72.4

1/ No species composition estimate for 1969.
Hunting pressure and harvest on the west slope increased over 1969 levels
under standard regulations with six birds in the bag and twelve in possession for a 93-day season. Greatest harvest of ducks was indicated to
have occurred during the November 30-December 28 period but again relatively
similar throughout all other periods, including the last period which was
shorter but even more productive than the others in terms of success per
hunter (Table 6). Total estimated west slope harvest increased about 16
percent from the 1969 figure of 23,781 to 27,673 ducks.
Special Teal Season.--An optional nine-day early teal season was again
granted the states in the Mississippi and Central Flyways in 1970. Colorado
chose to take the season from September 5-13 but restricted the open area to
North Park (Jackson County), Lake and Chaffee counties, and the portion of
the State east of Colorado Highway 71, U. S. Highway 350, and U. S. Interstate Highway 25. Estimated harvest was 5,585 birds by 1,179 hunters or
approximately 2.7 percent of the total east slope duck harvest by 4.3 percent of the total east slope duck hunters (Table 6).

�-60Table 6. Ducks bagged and hunting pressure by season type, area, and
interval of the 1970 season.

Dates

Estimated Harvest
% of
No. of
Tc&gt;ta1
Ducks

Estimated
Hunting Pressure
No. of
% of
Hunters
Total

EAST SLOPE
Teal Season (Special)
Sept. 5-13

5,585

2.7

1,179

4.3

San Luis Valley (Special)
Oct. 1-18

22,959

11.1

2,933

10.7

I

Regular Season (Point-system)
Oct. 17-0ct. 30
Oct. 31-Nov. 20
Nov. 21-Dec. 11
Dec. 12-Jan. 1
Jan. 2-Jan. 14

34,542
44,264
39,093
34,129
26,269

16.7
21.4
18.9
16.5
12.7

206,841

100.0

7,593
8,853
8,579
6,441
4,303

27.7
32.3
31.3
23.5
15.7

1,335
1,735
1,578
736

34.1
44.3
40.3
18.8

WEST SLOPE
Regular Season
Oct. 10-Nov. 1
Nov. 2-Nov. 29
Nov. 30-Dec. 28
Dec. 29-Jan. 10

7,306
6,835
8,828
4,704

26.4
24.7
31.9
17.0

27,673

100.0

San Luis Valley Experimental Season.--This was the eighth and final year of
experimental duck hunting seasons in the San Luis Valley. Bag limit was
based on a point-system regulation exactly the same as for the regular
season previously described. Results of the season obtained through special
field observations and surveys will be reported in a Migratory Bird Populations
Station Administrative Report in 1971 (Geis, et a1. 1971). However,
information on the season gathered from this survey are presented here.
Survey data indicate 2,933 hunters bagged an estimated 22,959 ducks during
the October 1-18 season (Table 6). This compares with 1969 figures of 2,706

�-61-

hunters harvesting 23,021 birds which indicated harvest remained quite
similar between years. However, numbers of hunters listed above are rather
meaningless and are no doubt somewhat deflated since they refer to successful hunters rather than estimates of total hunters. On the other hand,
estimated harvest is probably somewhat inflated because total harvest is
projected from data from the successful hunters. The special Federal survey,
because of its intensity, will produce more valid estimates of hunter participation and success.
Goose Harvest
Results of the 1970 goose season presented in Table 7 indicate 18,774
hunters bagged an average of 2.2 birds each per season for a total of
40,574 geese harvested in the State. This indicates an increase of about
126 percent in harvest and a 41 percent increase in hunters from the previous
year, most of which took place on the Central Flyway east slope. Thus,
goose harvest and hunting pressure in 1970 was at an all-time high for the
l7-year period of survey. Data collected in the questionnaire survey indicated an average crippling loss estimate of about 18 percent. Species
composition of geese in the 1970 bag, as in other years, remained about 99
percent Canada geese. The remainder is composed of snow geese and an even
smaller portion of white-fronted geese.
Arkansas Valley.--Food and water conditions were excellent during the 1970
Arkansas Valley goose season. Census figures indicated goose populations
numbered about 80,000 to 90,000 birds through the majority of hunting
season, the highest number on record. Approximately 62,000 geese were
counted on January 19, 1971, which is about equal to record numbers counted
in January, 1970 for Colorado. Major concentrations were present in the
Eads Lakes, Two Buttes, John Martin, and Meredith Reservoir areas. Hunting
hours from October 31 through December 11, 1970 were from one-half hour
before sunrise to noon, only. From December 12 through January 17, 1971,
full day shooting was allowed. Birds were present in the area on opening
day. This factor, plus half-day shooting which allowed birds to settle in
the area and establish feeding patterns, coupled with increased hunting
pressure and full-day shooting later in the season, all seemed to contribute
to the increase in hunting success. Harvest by area will be discussed in
a later section.
North-central Colorado.--Numbers of geese wintering in the Denver-GreeleyFort Collins area also increased significantly in 1970. Approximately
50,000 birds were censused in both late November and December with huntable
numbers present in early November prior to opening date on November 10.
At least 38,000 birds were counted in early January, 1971, the bulk of them
probably remaining through hunting season which ended January 17. The
high count in 1970-71 is the highest on record and represents an increase of
over 10,000 birds from 1969-70.

�-62-Table 7.

Goose harvest statistics,

1954-1970.

Total
Estimated
Hunting
Mortality

Number
of
Hunters

Average
Seasonal
Bag

Total
Estimated
Harvest

12,136
17,364
11,541
12,057
14,705
13 ,647
14,107
11,245

0.7
1.0
1.0
1.2
1.3
1.6
1.1
1.3

8,168
17,711
11,310
14,589
19,704
21,972
15,659
14,056

22.8
18.3
21.6
23.5
22.3
17.8
20.7
24.5

2,410
3,884
3,116
4,473
5,655
4,730
4,087
4,568

10 ,578
21,248
14,426
19,062
25,359
26,702
19,746
18,624

East
West
TOTAL

8,828
331
9,159

1.5
1.3
1.5

13,241
430
13 ,671

19.5
0.0
19.1

3,218

16,459
430
16,889

East
West
TOTAL

10,462
379
10,841

1.7
0.3
1.6

17,785
114
17 ,899

1502
0.0
17.8

3,184
3,184

20,969
114
21,083

1964

East
West
TOTAL

13 ,295
383
13,678

1.8
0.5
1.7

23,931
192
24,123

19.0
0.0
18.9

5,624
0 1/
5,624

29,555
192
29,747

1965

East
West
TOTAL

10,941
403
11,344

1.2
1.0
1.2

13 ,239
419
13,658

24.6
27.0
24.7

3,257
113
3,370

16,496
532
17,028

1966

East
West
TOTAL

15,443
364
15,807

1.9
1.1
1.9

29,613
420
30,033

16.2
0.0
16.2

5,745
87
5,832

35,358
507
35,865

1967

East
West
TOTAL

13,598
150
13,748

1.7
1.7
1.7

23,117
255
23,372

19.8
19 8
19.8

5,710
63
5,773

28,827
318
29,145

1968

East
West
TOTAL

13,050
417
13 ,467

1.5
0.7
1.4

19,088
291
19,379

16.6
16.6
16.6

3,806
58
3,864

22,894
349
23,243

1969

East
West
TOTAL

12,990
292
13 ,282

1.4
0.9
1.4

17,691
248
17,939

?J

Y

?J

8,831
180
9,011

48,634
951
49,585

Year

Slope

1954
1955
1956
1957
1958
1959
1960
1961
1962

1963

1970

East
West
TOTAL
Seventeen-year

Wounding Loss
Number
Percent

e.

18,399
2.2
39,803
18.2
375
2.1
771
18.9
18,774
2.2
40,574
18.2
average goose harvest (1954-70) is 19,048.

1/ No cripples reported on the west slope.

"fj Data not requested in survey questionnaire.

0 1/

3,218
0 1/

�-63-

For the sixth consecutive year, goose permits with a six-bird season limit
and other special regulations were in effect in most of the area between
Denver, Fort Collins and east of Greeley.
In 1969, the permit area was
enlarged but the limit was removed from number of permits which could be
issued. A relatively large area between Fort Collins and Interstate Highway 25, closed to hunting in previous years, was opened in 1970 and 13,611
hunting permits were issued. A special random survey sample drawn from
these permittees indicated that 11,187 hunters actually hunted geese one or
more times in the entire area and bagged 12,112 geese for an average bag of
1.08 birds per hunter.
Thus, active hunters increased about 82 percent
over 1969-70 and harvest increased about 190 percent.
Distribution of harvest
was greatest in Larimer County with almost 7,500 birds, while Weld County
was second with about 3,500 geese bagged in the area.
It is believed that
results of this special survey are more reliable than those of the comprehensive small game survey because the survey was specific to the north-central
season and the sample size was larger. Results of the regular survey in
terms of hunter participation and harvest by county are presented in a later
section.
San Luis Val1ey.--In 1970-71, the western portion of the San Luis Valley
formerly closed to goose hunting was opened to a limited permit hunt. Two
hundred permits were allowed and issued and a special survey of permittees
suggested 164 took part in the season.
Estimated harvest was 107 birds with
the majority taken in Rio Grande County.
Numbers of geese present during
fall and winter ranged from about 1,000 to 1,500 with the majority tallied
on the Monte Vista Refuge.
Total hunter participation and harvest results
in the Valley as estimated by the regular small game survey will be presented
in the section relating to goose hunting by county.

Waterfowl

Harvest by County

The reader is cautioned that information presented in this section is subject
to a great deal more error in accuracy than estimates in previous sections,
since the original sample has been broken down to county, thus decreasing
the size of sample on which to base estimates.
This is probably even more
true of geese than ducks because there are many more duck hunters.
Consequently, it is realized that in some counties, both duck and goose harvest
has been over-estimated, and in others, under-estimated.
However, despite
this error, these data represent the most accurate information possible at
the present time except for special survey data already mentioned.
Tables 8 and 9 compare the 1970 duck and goose harvests, respectively, with
averages of previous seasons by county within each waterfowl region.
These
regional divisions of the State were located on the basis of waterfowl
migration, location, and topography, and permit a closer evaluation of annual
harvest, yearly changes in harvest patterns, and the effect of different
types of hunting seasons in various portions of Colorado.
Grossly, it appears that the greatest duck harvest again occurred in the
Central Region, although the percentage of total State bag for this area
again decreased somewhat from the previous year. The harvest increased

�-64about 2,000 birds from about 79,000 bagged in 1969 to 81,000 in 1970.
Greatest percentage increases took place in the Northeast and Southeast
Regions with an estimated 10,000 bird increase in each totaling 49,000 and
37,000 birds, respectively. Although many hunters still do not seem willing
to travel to the eastern portions of the State, there is an increasing trend
in use of these areas. The east slope again contributed about 87 percent of
the total state duck harvest in 1970. Greatest harvest again took place in
Weld County. Number of ducks bagged increased in most areas on the west slope.
As indicated in previous sections, the 1970 State goose harvest increased
in volume by 126 percent over 1969 figures to over 40,500 birds. The Southeast Region was again indicated by the regular small game harvest survey to
be the high harvest area with just over 30,000 geese taken out of the total
of almost 40,000 bagged on the entire east slope. Only about 9,000 geese
were indicated to have been harvested in the Central Region, which takes in
the majority of the north-central goose hunting permit area discussed previously in which about 12,000 birds were suggested to have been bagged
through results of the special survey. Thus, a discrepancy of about 3,000
birds exists between the two surveys. The special survey figure of 12,000
is probably the most correct and it is possible the 30,000 goose figure for
the Southeast Region is somewhat inflated. In the Southeast Region, which
supposedly accounted for almost 75 percent of the State harvest, Baca, Kiowa,
Prowers, Crowley, and Bent counties were the top harvest areas in that order.
Larimer and Weld counties were indicated to have been high harvest areas in
the Central Region and third and fifth in order in the State, respectively.
Total harvest in the San Luis Valley was suggested to have been 279 birds.
The estimated west slope harvest increased from 248 birds in 1969 to 771 in
1970, the majority of which was taken in Grand County.
Waterfowl Management Units
Again this year, harvest information has been gathered on the basis of
waterfowl management units (Fig. 1). The purpose of this is a better alignment of data to provide information about specific flocks of ducks and geese.
For example, Unit 1 encompasses the area utilized by wintering ducks at
Jumbo Reservoir, Unit 5 is North Park, and Unit 15 is the San Luis Valley.
In many cases, duck and goose flock boundaries transcend county lines and
it has been difficult to put together county information so that we could
look at the influence of hunting pressure and harvest on separate flocks.
We had hoped that management units would help solve this. However, there
are still problems with interpreting results which have to be examined by
year. Results of this portion of the survey are displayed in Tables 10 and
11 for ducks and geese, respectively. Estimated numbers of hunters and
harvest for the 1970 season are shown by management unit and compared with
the six-year averages from 1964-1969. No other comments will be made at
this time.

�..

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�-66-

LITERATURE CITED

Boyd, R. J. 1971. Hunter harvest surveys. Colo. Div. of Game, Fish and
Parks, Game Res. Rept. July. In press.
Geis, A. D., E. M. Martin, R. M. Hopper, and H. D. Funk. 1971. Progress
report - 1970 experimental duck hunting season in the San Luis Valley
of Colorado - an evaluation of the point system in regulating harvest.
Migr. Bird Pop. Sta. Admin. Rept. No. 210. Laurel, Md. July 6. 14 pp.

~:J~-(£_c'..._~_~

Prepared by......J..o:ak~;;;;....::;:::~~,
~t~~.
Howard D. Funk
Section Chief, Small Game Research

_

�-67Table 8.

Duck harvest by Region and County.

Waterfowl
Region
and County

1970 Duck Harvest
Percent
Number
Harvested of Total

Estimated 1970
Hunting Pressure
Nwnber
Percent
Hunters of Total

Sixteen-year Average
1954-1969
Percent
Number
of Total
Harvested

EAST SLOPE
NORTHEAST
Cheyenne
Kit Carson
Lincoln
Logan
Morgan
Phillips
Sedgwick
Washington
Yuma
NORTHEAST TOTAL

620
1,241
14,065
22,546
207
2,689
2,689
52171
49,228

0.0
0.3
0.6
6.8
10.9
0.1
1.3
1.3
2.5
23.8

27
192
164
1,699
2,823
109
411
4ll
630
6,468

0.1
0.7
0.6
6.2
10.3
0.4
1.5
1.5
2.3
23.6

401
1,590
1,035
7,461
9,721
233
3,717
2,097
32287
29,542

0.3
1.2
0.8
5.7
7.5
0.2
2.9
1.6
2.5
22.7

SOUTHEAST
Baca
Bent
Crowley
Huerfano
Kiowa
Las Animas
Otero
Prowers
Pueblo
SOUTHEAST TOTAL

1,241
9,928
3,103
414
4,344
414
5,170
4,344
7 653
36,611

0.6
4.8
1.5
0.2
2.1
0.2
2.5
2.1
3.7
17.7

329
932
521
55
767
llO
630
603
576
4,523

1.2
3.4
1.9
0.2
2.8
0.4
2.3
2.2
2.1
16.5

1,194
3,306
2,760
559
1,686
1,099
2,272
3,752
3 327
19,955

0.1
2.5
2.1
4.3
1.3
0.9
1.7
2.9
2.6
15.4

CENTRAL
Adams
Arapahoe
Boulder
Douglas
Elbert
E1 Paso
Jefferson
Larimer
Weld
CENTRAL TOTAL

15,100
1,034
9,722
207
827
2,068
414
14,272
372438
81,082

7.3
0.5
4.7
0.1
0.4
1.0
0.2
6.9
18.1
39.2

1,891
329
1,371
82
55
384
219
3,125
52564
13 ,020

6.9
1.2
5.0
0.3
0.2
1.4
0.8
11.4
20.3
47.5

7,544
1,718
7,607
211
291
1,103
1,468
ll,885
282456
60,283

5.8
1.3
5.9
0.2
0.2
0.8
1.1
9.2
21.9
46.4

SAN LUIS VALLEY
Alamosa
Conejos
Costilla
Rio Grande
Saguache

12,617
2,482
2,482
15,513
3 2930

6.1
1.2
1.2
7.5
1.9

1,069
27
164
1,015
521

3.9
0.1
0.6
3.7
1.9

4,670
2,507
882
5,756
3 2928

3.6
2.0
0.7
4.4
3.0

SAN LUIS VALLEY
TOTAL

37,024

17.9

2,796

10.2

17,743

13.7

-----------------------------------------------------------------------------------

�-68Table 8.

Duck harvest by Region and County, continued.

Waterfowl
Region
and County

HIGH COUNTRY (E)
Chaffee
Clear Creek
Cus ter
Fremont
Gilpin
Jackson
Lake
Park
Teller
HIGH COUNTRY TOTAL

1970 Duck Harvest
Number
Percent
Harvested of Total

620
207

Estimated 1970
Hunting Pressure
Numhe r
Percent
Hunters of Total

Sixteen-year Average
J 954-1969
Number
Percent
Harvested
of Total

1,242

0.3
0.1
0.0
0.6

110
27
27
248

0.4
0.1
0.1
0.9

504
81
337
780

0.4
0.1
0.2
0.6

620

0.3

137

0.5

0
207
2,896

0.0
0.1
1.4

27
27
603

0.1
0.1
2.2

216
126
276
41
2,361

0.2
0.1
0.2
0.0
1.8

WEST SLOPE
NORTHWEST
Garfield
Moffat
Rio Blanco
Routt
NORTHWEST TOTAL
WEST CENTRAL
Delta
Mesa
Montrose
Ouray
W. CENTRAL TOTAL
SOUTHWEST
Archuleta
Dolores
Hinsdale
La Plata
Mineral
Montezuma
San Juan
San Miguel
SOUTHWEST TOTAL
HIGH COUNTRY (W)
Eagle
Grand
Gunnison
Pitkin
Summit
HIGH COUNTRY TOTAL

2,795
443
1,494
138
4,870

10.1
1.6
5.4
0.5
17.6

403
133
161
27
724

lO.3
3.4
4.1
0.7
18.5

1,637
533
721
818
3,709

8.5
2.8
3.8
4.3
19.4

3,321
7,002
5,313

12.0
25.3
19.2

540
893
541

13.8
22.8
13.8

15,636

56.5

1,974

50.4

3,163
4,587
2,371
168
10,289

16.5
24.0
12.4
0.8
53.7

830
194

3.0
0.7

161
27

4.1
0.7

1,660

6.0

243

6.2

941

3.4

188

4.8

0
3,625

0.0
13.1

27
646

0.7
16.5

119
28
66
1,475
87
864
14
219
2,872

0.6
0.1
0.3
7.8
0.5
4.5
0.1
1.1
15.0

470
581
2,214
277

1.7
2.1
8.0
1.0
0.0
12.8

82
161
220
82
27
572

2.1
4.1
5.6
2.1
0.7
14.6

801
441
672
273
93
2,280

4.2
2.3
3.5
1.4
0.5
11.9

3,542

-----------------------------------------------------------------------------------

�-69Table 8.

Duck harvest by Region and County, continued.

1970 Duck Harvest
Number
Percent
Harvested of Total

Waterfowl
Region
and County

Estimated 1970
Hunting Pressure
Number
Percent
Hunters of Total

Sixteen-year Average
195L~-1969

Number
Harvested

Percent
of Total

Summary by Region
NORTHEAST
SOUTHEAST
CENTRAL
SAN LUIS VALLEY
HIGH COUNTRY (E)
NORTHWEST
WEST CENTRAL
SOUTHWEST
HIGH COUNTRY (W)
TOTAL OF REGIONS

49,228
36,611
81,082
37,024
2,896
4,870
15,636
3,625
3,542
234,514

20.7
14.4
41.6
8.9
1.9
2.3
6.3
2.1
1.8
100.0

6,468
4,523
l3 ,020
2,796
603
724
1,974
646
572
31,326

20.7
14.4
41.6
8.9
1.9
2.3
6.3
2.1
1.8
100.0

29,542
19,955
60,283
17,743
2,361
3,709
10,289
2,872
2,280
149,034

19.8
l3.4
40.5
11.9
1.6
2.5
6.9
1.9
1.5
100.0

EAST SLOPE
WEST SLOPE

206,841
27,673

88.2
11.8

27,410
3,916

87.5
12.5

129,884
19,150

87.2
12.8

Table 9.

Goose harvest by Region and County.

Haterfow1
Region
and County

1970 Goose Harvest
Number
Percent
Harvested of Total

Estimated 1970
Hunting Pressure
Percent
Number
Hunters of Total

Sixteen-year Average
1954-1969
Percent
Number
of Total
Harvested

EAST SLOPE
NORTHEAST
Cheyenne
Kit Carson
Lincoln
Logan
Morgan
Phillips
Sedgwick
Washington
Yuma
NORTHEAST TOTAL

0
0
0
80
0
119
0
40
239

0.0
0.0
0.0
0.2
0.0
0.3
0.0
0.1
0.6

74
18 .
147
644
19
55
55
110
1,122

0.4
0.1
0.8
3.5
0.1
0.3
0.3
0.6
6.1

24
26
14
145
534

0.1
0.1
0.1
0.8
3.1

48
97
84
972

0.3
0.6
0.5
5.6

-----------------------------------------------------------------------------------

�-70-

Table 9.

Goose harvest by Region and County, continued.

Waterfowl
Region
aridCounty

1970 Goose Harvest
Number
Percent
Harvested of Total

Estimated 1970
Hunting Pressure
Number
Percent
Hunters of Total

Sixteen-year Average
1954-1969
Number
Percent
Harvested
of Total

SOUTHEAST
Baca
Bent
Crowley
Huerfano
Kiowa
Las Animas
Otero
Prowers
Pueblo
SOUTHEAST TOTAL

13,055
2,229
3,025
0
7,005
120
557
3,980
358
30,329

32.8
5.6
7.6
0.0
17.6
0.3
1.4
10.0
0.9
76.2

2,613
975
1,067
19
1,619
110
276
975
221
7,875

14.2
5.3
5.8
0.1
8.8
0.6
1.5
5.3
1.2
42.8

5,083
1,913
716
113
3,098
120
359
2,125
183
13,710

29.1
11.0
4.1
0.6
17.7
0.7
2.0
12.1
1.0
78.3

159
80
159

0.4
0.2
0.4

349
74
442

1.9
0.4
2.4

0
0
0
5,214
3z344
8,956

0.0
0.0
0.0
13 .1
8.4
22.5

18
74
18
4,931
3z109
9,015

0.1
0.4
0.1
26.8
16.9
49.0

297
160
88
4
6
28
150
832
927
2,492

1.7
0.9
0.5
T
T
0.2
0.8
4.8
5.3
14.2

80

0.2

111

0.6

80
80
39

0.2
0.2
0.1

55
203
18

0.3
1.1
0.1

9
137
117
2
2

0.1
0.8
0.6
T
T

279

0.7

387

2.1

267

1.5

3
36

T
0.2

21

0.2

3

T

63

0.4

CENTRAL
Adams
Arapahoe
Boulder
Douglas
Elbert
E1 Paso
Jefferson
Larimer
Weld
CENTRAL TOTAL
SAN LUIS VALLEY
Alamosa
Conejos
Cos tilla
Rio Grande
Saguache
SAN LUIS VALLEY
TOTAL
HIGH COUNTRY (EAST)
Chaffee
Clear Creek
Custer
Fremont
Gilpin
Jackson
Lake
Park
Teller
HIGH COUNTRY TOTAL

-----------------------------------------------------------------------------------

�-71-

Table 9.

Goose harvest by Region and County, continued.

Waterfowl
Region
and County

1970 Goose Harvest
Number
Percent
of Total
Harvested

Estimated 1970
Hunting Pressure
Number
Percent
Hunters of Total

Sixteen-year Average
1954-1969
Percent
Nwnber
of Total
Harvested

WEST SI.OPE
NORTHWEST
Gar fie1d
Moffat
Rio Blanco
Routt
NORTHWEST TOTAL
WEST CENTRAL
Delta
Mesa
Montrose
Ouray
W. CENTRAL TOTAL
SOUTHWEST
Archuleta
Dolores
Hinsdale
La Plata
Mineral
Montezuma
San Juan
San Miguel
SOUTHWEST TOTAL
HIGH COUNTRY (WEST)
Eagle
Grand
Gunnison
Pitkin
Sunnnit
HIGH COUNTRY TOTAL

179
19
198

0.0
23.2
2.5
25.7

25
175
25
225

1
115

0.6
75.7

60.1

2
118

1.3
77 .6
2.7
0.6
1.3

6.7
46.7
6.7

19

2.5

25

6.7

4
1
2

19

2.5

25

6.7

7

4.6

110

15.4

75

20.0

2.0
2.0

119

15.4

75

20.0

3
3

435

56.4

50

13.2

24

15.8

435

56.4

50

13.2

24

15.8

972
13,710
2,492
267
63
118
7
3
24
172656
17,504
152

5.5
77.7
14.1
1.5
0.4
0.7
0.0
0.0
0.1
100.0
99.1
0.9

Sunnnary by Region
NORTHEAST
SOUTHEAST
CENTRAL
SAN LUIS VALLEY
HIGH COUNTRY (E)
NORTHWEST
WEST CENTRAL
SOUTHWEST
HIGH COUNTRY (W)
TOTAL OF REGIONS
EAST SLOPE
WEST SLOPE

239
30,330
8,956
279

0.6
74.8
22.1
0.7

1,122
7,875
9,015
387

6.0
41.9
48.0
2.1

198
19
119
435
402574

0.4
T
0.3
1.1
100.0
98.1
1.9

225
25
75
50
182774
18,399
375

1.2
0.1
0.4
0.3
100.0
98.0
2.0

39,803

771

�-72-

Table 10.

Estimated duck hunters and harvest by management unit, 1964-1970.

1970 Season
Percent Number
of Total Bagged

Six-year Average
1964-1969
Number
Percent Number
Hunters of Total Bagged

Management
Unit

Number
Hunters

1

1,817

5.8

EAST SLOPE
15,712
6.7

1,425

6.2

7,89l

5.9

2

2,036

6.5

14,305

6.1

1,392

6.1

7,471

5.6

3

4,980

15.9

31,425

13.4

3,444

15.1

19,451

14.6

4

3,446

11.0

17 ,589

7.5

2,165

9.5

11 ,498

8.7

5

125

0.4

704

0.3

102

0.4

562

0.4

6

5,545

17.7

42,9l6

18.3

4,200

18.5

25,360

19.2

7

282

0.9

938

0.4

272

1.2

1,116

0.8

8

658

2.1

4,456

1.9

383

1.7

1,847

1.4

9

846

2.7

6,332

2.7

506

2.2

2,696

2.0

10

1,159

3.7

7,739

3.3

611

2.7

2,810

2.1

11

344

1.1

1,407

0.6

224

1.0

682

0.5

12

1,598

5.1

11 ,022

4.7

1,200

5.3

6,367

4.8

13

1,128

3.6

12,898

5.5

779

3.4

4,049

3.0

14

313

1.0

1,407

0.6

257

1.1

1,673

1.3

15

3,133

10.0

37,99l

16.2

2,830

12.4

22,313

16.8

Sub-total

27,410

87.5

206,841

88.2

19,790

86.8

115,787

87.1

16

3,916

12.5

WEST SLOPE
11.8
27,673

3,007

13.2

17,116

12.9

GRAND TOTAL 31,326

100.0

234,514

22,797

100.0

132,903

100.0

Percent
of Total

100.0

Percent
of Total

�-73-

Table 11.

Management
Unit

Estimated goose hunters and harvest by management unit, 1964-1970.

Number
Hunters

1970 Season
Percent Number
of Total Bagged

Percent
of Total

Six-year Average
1964-1969
Number
Percent Number
Hunters of Total Bagged

Percent
of Total

EAST SLOPE
1

244

1.3

122

0.3

248

1.8

124

0.6

2

357

1.9

203

0.5

490

3.6

482

2.3

3

2,459

l3 .1

2,150

5.3

1,856

13.7

1,957

9.1

4

5,388

28.7

6,289

15.5

1,838

l3.6

1,940

9.1

40

0.3

110

0.5

5
6

1,483

7.9

730

1.8

1,212

8.9

1,100

5.1

7

94

0.5

41

0.1

39

0.3

19

0.1

8

l31

0.7

81

0.2

80

0.6

141

0.7

9

188

1.0

41

0.1

66

0.5

53

0.2

10

1,859

9.9

6,208

15.3

1,913

14.1

4,703

21.9

II

2,967

15.8

15,824

39.0

2,420

17.9

5,358

25.0

12

1,671

8.9

3,165

7.8

2,065

15.2

3,783

17.7

13

1,108

5.9

4,625

1l.4

620

4.6

1,030

4.8

14

131

0.7

81

0.2

126

0.9

130

0.6

15

319

1.7

243

0.6

246

1.8

195

0.9

18,399

98.0

39,803

98.1

13 ,258

97.8

21,125

98.6

Sub-total

WEST SLOPE
375

2.0

771

1.9

297

2.2

291

1.4

GRAND TOTAL 18,774

100.0

40,574

100.0

13,555

100.0

21,416

100.0

16

��-75October,

JOB PROGRESS

State of

~C~OL~O~RA~D~O~

Project No.

W-88-R-16

Work

Plan No.

1

1971

REPORT

_

Job No.

Job Title

San Luis Valley Cooperative

Period Covered:

May 11, 1970 to October

12
Mallard

Investigation

18, 1970

Personnel:
Charles Hayes and Jack Randall, Bureau of Sport Fisheries and
Wildlife;
Bill Adrian, Ron Arant, Ron Blumberg, George Bock, Clait Braun,
Harold Burdick, Bob Clark, Don Crane, David Croonquist, John Ellenberger,
Larry Finnell, Howard Funk, John Goettl, Joe Griess, Don Horak, Tom Lines
Daryl Luce, Bill Olmstead, Jr., Frank Rinella, Wayne Russell, Bruce Sigler,
Mike Szymczak, Ken Wagner, Allen Whitaker, Allen White, Charles Woodward
and Richard Hopper, Colorado Game, Fish and Parks Division.

ABSTRACT

The duck breeding population in the San Luis Valley was estimated at 29,410
pairs for 1970, including 13,704 pairs of mallards.
This represented increases of nearly 1,600 pairs for all species combined and about 1,500 pairs
of mallards over 1969. The 1970 mallard estimate was slightly above the sixyear average (1964-1969).
Gadwalls increased significantly in 1970, while
the three teals dropped sharply below the 1969 level.
The point-system regulation was again tested in the San Luis Valley during
the October 1-18, 1970 experimental season.
This season was similar to the
two previous seasons except that the daily bag limit was 100 rather than
70 points, the number of 10 point ducks was increased, and the middle point
category was decreased from 30 or 35 to 20 points.
Mallard drakes were assigned
20 points with mallard hens being 90 points.
Colorado personnel conducted 65 hunter performance surveys during the 1970
experimental season, including 16 on private land and 49 on public land.
Survey cards were sent to the Bureau of Sport Fisheries and Wildlife for
analysis.

��-77-

SAN LUIS VALLEY COOPERATIVE

MALLARD

INVESTIGATION

Richard M. Hopper

An intensive study of the mallard population in the San Luis Valley was
started in 1963 as a cooperative investigation between the Bureau of
Sport Fisheries and Wildlife and the Colorado Game, Fish and Parks
Division.
A major feature of this study has been an annual experimental
duck hunting season held during the period October 1-18. The objectives
listed below refer to the collection of data designed to evaluate the
experimental season and include only those for which the Colorado Division
had primary responsibility.
Results presented here pertain only to these
stated objectives.
A complete analysis of the 1970 experimental season
and related job activities are found in Administrative Report No. 210
(Geis et al., 1971) prepared by members of both agencies and distributed
by the Bureau of Sport Fisheries and Wildlife, Branch of Wildlife Research,
Migratory Bird Populations Station.

P. S. OBJECTIVE
1.

To develop a harvest

formula

for the San Luis Valley mallard

2.

To determine

reaction

to various

hunter

types of hunting

population.

regulations.

SEGMENT OBJECTIVES
This study was designed to determine the influence of various types of
hunting regulations and bag limits on the local breeding population of
mallards in the San Luis Valley.
Colorado's part of the study is covered
under the following segment objectives:
1.

Determine the size of the 1970 duck breeding
San Luis Valley.

2.

Assist in determining the ability and/or willingness of hunters
to abide by specific regulations and determine wounding loss
during the 1970 experimental season.

3.

Evaluate the results of the 1970 experimental season and write
reports in cooperation with the personnel from the Bureau of
Sport Fisheries and Wildlife.

METHODS

population

in the

AND MATERIALS

Methods for conducting the 1970 breeding population survey, air-ground
comparison study, and hunter performance survey remained the same as those
in 1967 (Segment 13). A discussion of these procedures appears in a previous report (Hopper and Rutherford, 1968) and will not be repeated here.

�-78-

RESULTS AND DISCUSSION

Breeding Population Survey
Air-Ground Comparison Study
Air and ground counts of ducks are compared by species in Table 1 for
122.5 miles of selected transect. The proportion identified from the
air (visibility ratio) for the mallard for example, indicates that
the air crew identified, as mallards, 25.5 percent of the mallards
actually present on the air-ground comparison transects. A high proportion of the pintails were observed from the air (0.370), while the
visibility ratio for cinnamon teals was quite low (0.060). An unusually high ratio was calculated for green-winged teals (0.375), perhaps because of the rather small sample size. Visibility ratios were
applied to the regular air transect data as correction factors in
calculating estimates of total breeding pairs for Russell Lakes and
the major portion of the Valley, excluding Monte Vista National Wildlife Refuge and the total-count areas.

Table 1. Air-ground comparison of ducks counted on 122.5 miles of
transect in the San Luis Valley. 1970.

Species

Estimated Breeding Pairs
Ground
Air
Pairs
Percent
Pairs
Percent

Mallard

474

48.0

121

59.3

0.255

Gadwall

130

13.2

12

5.9

0.092

Pintail

100

10.1

37

18.1

0.370

Shoveler

27

2.8

6

2.9

0.222

Blue-winged Teal

16

1.6

1

0.5

0.062

Cinnamon Teal

151

15.3

9

4.4

0.060

Green-winged Teal

40

4.1

15

7.4

0.375

American Widgeon

20

2.0

1

0.5

0.050

Redhead

11

1.1

2

1.0

0.182

Other Divers

18

1.8

0

0.0

0.000

Totals

987

100.0

204

100.0

0.207

Proportion
Identified
From the Air

�-79-

Duck Breeding

Population

Estimate

Amount of habitat, sample sizes, and estimated number of breeding pairs
of ducks are shown in Table 2 for the six areas included in the 1970
survey.
The 1969 estimate plus the average for the period 1964-69 are
included for comparison.
The 1970 survey produced an estimated 29,410
breeding pairs of ducks, or about 1,500 pairs above the 1969 estimate
and about 2,000 pairs more than the 6-year average (1964-1969).
The
estimate for 1970 was the second highest since intensive surveys began
in 1964. The largest figure, 32,762 pairs, was obtained in 1964.
Species composition of the 1970 San Luis Valley breeding population is
presented in Table 3, along with 1969 figures and the 6-year averages.
The mallard estimate for 1970 reached 13,704 pairs, an increase of nearly
1,600 pairs over 1969 but only about 150 pairs above the 6-year average.
As normal, mallards constituted in the vicinity of 50 percent of the
total duck breeding population.
Gadwalls were next in importance, contributing 6,402 pairs in 1970, or about 22 percent of the total. This
represented a significant increase over 1969 and the 6-year average.
Pintail and shoveler numbers remained generally stable, while the three
species of teal experienced a large decline in 1970 from the 1969 level.
The 1970 sampling error, figures at ±17.3 percent, was less than for any
other year of the study.

Harvest

Information

The 1970 experimental duck hunting season (October 1-18) in the San Luis
Valley was again regulated by the point system.
This marked the third
year for testing this type of regulation.
The point system tested in 1970
differed from those in 1968 and 1969 in that the daily bag limit was 100
instead of 70 points, more ducks were placed in the 10 point category, and
the middle point category was reduced from 30 or 35 to 20 points.
Mallard
drakes were allocated 20 rather than 10 points, with mallard hens, redheads,
canvasbacks, and hooded mergansers assigned to a 90-point category.
All
other species and sexes were assigned 10 points.
The object was to attempt
to shift some of the shooting pressure away from mallard drakes and toward
10-point birds.
State personnel helped in gathering hunter performance data during the 1970
experimental season in an effort to determine hunter compliance and reaction
to the point system regulation in effect.
Sixty-five hunter performance
surveys were conducted by State men, including 16 on private land and 49
on public land. Survey cards were mailed to the Bureau of Sport Fisheries
and Wildlife for tabulation and analysis along with those submitted by
Federal personnel.
Results of this survey, as well as Bureau harvest,
wing collection, and hunter opinion surveys, are presented in the joint
report mentioned earlier (Geis et al., 1971).

�Table 2. Breeding pairs by density type as estimated from the San Luis Valley regular air transects, and
nesting transects, 1970, with 1969 and 6-year average for comparison.

Square Miles
Habitat

Percent
Sample

Estimated No. Breeding Pairs 1/
6-Year Average 1/
(1964-1969)
1969
1970

Monte Vista NWR

22

4.03

4,367

4,420

3,815

Russell Lakes

6

33.33

489

876

1,188

San Luis Lakes

7

100.00

456

813

268

Mishak Lakes

4

100.00

263

313

315

Adams Lake

1

100.00

19

103

38

Remainder of Valley

1,265

17.95

23,816

21,354

21,846

Totals

1,305

17.80

29,410

27,879

27,470

Type

I

C»
0
I

1/ Determined from nesting transects on Monte Vista National Wildlife Refuge and from aerial transects on
all other areas.

1/ Only 5-year averages (1965-1969) for San Luis Lakes and Mishak Lakes and a 4-year average (1966-1969)
for Adams Lake.

�Table 3. Species composition of the San Luis Valley duck breeding population, 1970, with 1969 and 6-year
average for comparison.

Species

1970
Pairs

%

1969
Pairs

%

Pairs

%

Mallard

13,704

46.5

12,139

43.5

l3,557

49.4

Gadwall

6,402

21. 7

2,735

9.8

2,543

9.2

Pintail

2,813

9.6

2,397

8.6

3,393

12.4

554

1.9

2,889

10.4

1,540

5.6

Blue-winged/Cinnamon Teal

2,874

9.8

5,496

19.7

3,182

11.6

Shoveler

1,647

Green-winged Teal

I

5.6

00

1,080

3.9

1,270

4.6

American Widgeon

785

2.7

o

0.0

56

0.2

Redhead

520

1.8

1,031

3.7

1,652

6.0

Other Divers 1./

111

0.4

112

0.4

277

1.0

Totals

29,410

100.0

27,879

100.0

27,470

100.0

± .05 Standard Error ~/

+ 17.3%

±

22.8%

l/ Includes lesser scaups, canvasbacks, ruddy ducks, and common mergansers.
~/ Excludes Monte Vista NWR and Russell Lakes.

± 24.9%

I-'
I

�-82-

LITERATURE CITED
Geis, A. D., E. M. Martin, R. Hopper, and H. Funk. 1971. Progress
report: 1970 experimental duck hunting season in the San Luis
Valley of Colorado - an evaluation of the point system in regulating the harvest. U.S.D.I., Bur. Sport Fisheries and Wildl.,
Migratory Bird Pop. Sta., Admin. Rpt. No. 210. 14 pp.
Hopper, R. M., and W. H. Rutherford. 1968. San Luis Valley cooperative
mallard investigation. Colo. Game, Fish and Parks Div., Fed. Aid
Game Res. Rpt. Oct. pp. 33-42.

Prepared by

tt.:.u Jz,z JJo/!1C--f..

Richard M. Hopper
Wildlife Researcher

�-83October,

1971

JOB FINAL REPORT

State of

COLORADO

Project No.

W-88-R-16

Work Plan No.
Job Title:

1
Job No.
13
Determination of Method for Developing and
Managing Waterfowl Habitat - Shallow Impoundment

Period Covered:

April

Personnel:

__

.~~ratory

Bird Inves.tigations

Study

1, 1967 to March 31, 1971

Ralph Baker, Delwin Benson, Robert

Oakleaf and Richard

Hopper.

ABSTRACT

A shallow impoundment study was initiated in 1967 to investigate plant succession in relation to various types of waterfowl use. Unfortunately, the study
had to be abandoned because of the lack of sufficent water to meet the demands
of the present study design.
A literature review, covering wetland ecology and
methods of sampling wetland vegetation, was conducted.
The west end of Bonny
Reservoir in extreme east central Colorado was selected as the study area.
Pre-impoundment
studies of vegetation in the native meadow portion of impoundment No. 1 in 1969 showed three species of grass with the highest frequencies
(percent of total quadrats of occurrence), inland saltgrass (Distichlis stricta),
bluestem wheatgrass (Agropyron smithii),
and sand dropseed (Sporobolus crytandrus).
Inland saltgrass, bluestem wheatgrass, and cheatgrass brome (Bromus
tectorum) had the greatest average densities (stems/quadrat).
Summercypress
(Kochia scoparia) and cammon Russian thistle (Sa1soli kali)
had the highest
frequencies and densities of the species found in the previously farmed area
of impoundment No.1.
Impoundment No. 1 was flooded immediately following termination of pre-impoundment studies.
Post-impoundment
studies of vegetation were conducted one year
after flooding in 1970. Most species that dominated in frequency and density
in 1969 also dominated in 1970. Nearly all species dropped in frequency from
1969 to 1970 on both the native meadow and farmed portions of the study area.
Overall density increased on the native meadow portion from 1969 to 1970, due
mostly to a large increase by inland saltgrass.
Pre and post impoundment soil samples indicated some differences after only one
year of flooding.
Conductivity, potassium and iron increased in the samples
folloWing flooding, while pH and phosphorus decreased.

�-84-

RECOMMENDAT IONS

1.

Terminate the current investigation as presently outlined because
the lack of sufficient water to meet demands of the study design.

of

2.

Initiate a study designed to evaluate waterfowl food crops planted in
two of the three impoundments.
A portion of the data collected for
Work Plan 1, Job 13 would be of value in such a study, particularly
that dealing with soils.

�-85-

SHALLOW

IMPOUNDMENT

STUDY

.Richard M. Hopper

The shallow impoundment study was initiated in 1967 (Segment 13) as a longterm investigation designed to acquire knowledge in the development and
management of waterfowl habitat.
The object was to construct three impoundments and to permanently flood them at two-year intervals, thereby creating
three different stages of plant succession for study and comparison with
various types of waterfowl use. Drawdown studies were also planned on two
additional impoundments for comparison with the permanent impoundments.
The first year of study involved a review of literature and the selection
of a study area (Hopper 1968), while the second year entailed the design
and construction of the dikes and ditching systems for three permanent
impoundments (Hopper 1969).
The third (Hopper 1970) and fourth years were
concerned mainly with pre-impoundment vegetative and soil studies of the
first shallow impoundment to be flooded.
Unfortunately,
this study had to.be terminated after completion of the
fourth year of work because of the lack of sufficient water to meet the
demands of the study design.
Two wells drilled in 1968 and 1970 simply
would not yield enough water to fill and maintain even two of three shallow
impoundments.
Thus, this report constitutes a "Job Final Report" for Work
Plan 1, Job 13 and includes all information collected through the fourth
year of study.

P. S. OBJECTIVE
To determine the effects of (a) shallow, permanent impoundments; and (b)
shallow, drawdown impoundments on waterfowl use and harvest at Bonny
Reservoir.
METHODS

AND MATERIALS

Literature

Review

Pertinent bibliographic abstracts and references were searched for books,
articles, papers, etc. covering the subjects of wetland ecology and methods
of sampling wetland vegetation.
This subject matter was retrieved, read,
and reviewed.
Information important to the study was abstracted on index
cards and a bibliographic
list was compiled for later reference.

Selection

of Study Area

Potential development sites in the Bonny Reservoir and San Luis Valley
areas were investigated from the standpoint of which would constitute the
best study area. Items considered in making a decision included: (1) amount
and reliability of water supply; (2) slope of land; (3) amount of land available in a continuous block; and (4) ability of land to maintain surface water.

�-86-

...
CD

:J

U

:J
•...

-

.•..

ro

U')

I'rt)

=

Fig.

1.

.L

Dike and ditch

system

of three shallow.

Dermanent

imDoundments.

Bonny.Reservoir.

�-87-

Design and Construction

of Impoundments

Dike ";;mdDitch Work
A contour map (2-foot interval) was used as a base for designing the dike
and ditch systems for the three shallow, pe11manent impoundments.
Dike and
ditch locations were drawn on an overlay of this map, keeping in mind that
thre three resulting impoundments should be as nearl~ equal in size as the
topography of the site would allow.
Cubic yards (yd ) of earthwork
required, including 10 percent for settlement were computed using the height
and length of each lOO-foot section of dike.
These sections were then
added together to determine total yardage of earthwork for each impoundment's dike system.
Locations of the dikes were staked on the ground with
laths.
The following

1.

dike and ditch specifications

were outlined:

Dike work
- 3:1 (both faces)

(a)

Side slopes of dikes

(b)

Top width of dikes

(c)

Source of fill for dikes - Fill material should be obtained
from the upstream side of dikes.
The borrow area should be
sufficiently wide (50 feet or more) ,to produce a gently
sloping and shallow depression adjacent to the upstream toe
of the dikes.
Excavation of fill,material should not exceed
one foot below the original ground level. This criteria, will
determine the actual width of the borrow area. Material for
the deeper fills (over 3.5 feet) of Some dike sections in
impoundment Nos. 2 and 3 should be taken from the area
immediately north of No.3 on the edge of the bench directly
above the existing pond (Fig. 1).

(d)

Clearing dike locations and barrow areas - The area to be
covered by the bases of the dikes should be scraped and cleared
of all vegetation, including roots, to a depth of four inches
below the original ground level. Mowing and burning is also
recommended to reduce the bulk of debris.
Similar removal of
vegetation in the borrow area should be to a depth of at least
two inches below the original ground level. Cleared material
is to be deposited over the edge of the bench east of impoundment No.3.

(e)

Compaction of dikes - Suitable compaction should be attained
by traveling over the dikes w'ith heavy rubber-tired equipment
or similar machinery periodically during construction.
Compaction of this nature should occur after the addition of each
layer of fill material on the dike.
The fill material should
be wetted if it is otherwise too dry for desirable compaction.

- 6 feet

�-88-

2.

(f)

Installation of water control structures -Three water control
structures of the flash-board design were purchased and delivered to the construction site. The contractor need only
install these structures.
The horizontal tube should be placed
a little below the original ground level and in line with the
ditches to be constructed across impoundment Nos. 2 and 3.

(g)

Emergency spillway - An earthen spillway
only for impoundment No.3.

is to be constructed

Ditch work
(a)

Location of ditches - Ditches are to be constructed across
impoundment Nos. 2 and 3. The ditch across impoundment No.
2 should extend in a straight line from the water. control
'structure on impoundment No.1
to the water control structure
on impoundment No.2.
The ditch across impoundment No. 3
should extend in a similar manner between water control
structures of impoundment Nos. 2 and 3. These ditches are
necessary to transfer water across the impoundments prior to
the time they are to be flooded.

(b)

Type of ditches - The ditches are to be well trenched farmtype ditches (deep vee) with excavated soil pulled away. Water
flowing in ditches should be largely restricted to below
original ground level to reduce seepage.

The contract for construction of the dikes and ditches was awarded to the
Hale Soil Conservation District.
They, in turn, subcontracted the work to
Virgil Brueggeman of Wray, Colorado.
Only the three permanent impoundments were constructed.
It was decided not
to construct the two drawdown impoundments because of the absence of a
sufficient quantity of water to flood all five. As a result, drawdown
studies were temporarily abandoned as part of the overall investigation.
The three impoundments constructed were to be utilized for drawdown studies
following termin~tion of studies involving permanent water levels.

Well and Pump Work
Permits to drill two wells were obtained from the State Engineer's office,
one in 1967 and one in 1969. Specifications called for each well to have
a casing of 14 inches and an estimated yield of 900 gallons per mintue
(2 c.f.s.).
Fifteen horsepower electric pumps were recommended for
installation over the wells.
Layne-Western Company of Denver was named the
drilling contractor.

Pre- and Post-impoundment

Studies

Vegetation
Permanent quadrats were selected as one means for describing vegetation
associated with the impoundment sites before and after flooding.
Forty-

�-89-

eight, ~-square-meter
(~ m. x 1 m.) quadrats were established in impoundment No. 1 (Fig. 1) on the area to constitute the shallow-water and moist
soil portion of the impoundment.
A different method of vegetative sampling
was planned for the deep'-water portion of the impoundment (borrow area)
adjacent ·to the upstream face of the dikes where the fill material for the
dikes was obtained.
Locations of the 48. quadrats were determined through stratified random
sampling.
The ditch crossing impoundment No •.1 separates two portions of
contrasting vegetation and .it was necessary to stratify the impoundment
area for. sampling purposes.
The area to the north of the ditch covers
about two-thirds of impoundment No. 1 and consists of native meadow-type
vegetation that has been hayed and grazed in the past but not broken by
the plow. Land south of the ditch was farmed prior to 1969, with small
grains as the predominant crop. As a result, 32 quadrats were placed in
the north area and 16 in the' south area.
Twelve reference points were randomly selected and permanently marked with
steel posts along the main (east) dike of impoundment No.' 1 (X - Z, Fig. 1),
with eight points in section X - Y and four in Y - Z. Corresponding points
were permanently marked at the opposite (west) side of the impoundment by
running 11 compass-line perpendicular to the main dike and across the impoundment from each of the 12 points on the main dike.
These reference points
were designated A through L.
Four quadrats were established along each compass-line within a 300-ft.
distance west of the west edge of the borrow area (deep water portion).
This 300-ft. distance was stratified in an east-west direction into four,
75-ft. segments, with one quadrat being randomly placed in each segment
along each compass line. These segments were designated as zones 1 through
4. This would insure sampling of vegetation in various depths of water and
on moist. and dry soils, since conditions will become progressively drier in
an east-west direction upon flooding of the impoundment.
The northwest and
southeast corners of each quadrat were permanently marked with 18 inch
wooden stakes.
An aluminum tag, identifying a quadrat by reference point
and number was placed on the northwest stake. Designation A-2 on an aluminum
tag, for example, refers to the quadrat in the second 75 ft. zone west of
the borrow area along compass line A.
Information collected at each quadrat included:
(1) date, (2) quadrat
number, (3) list of species present, and (4) number of individual plants or
stems of each species.
It was difficult to distinguish individual plants
of some species, especially many grasses and grass-like plants, thus stems
originating at the soil surface were counted.
Analysis of vegetative data
involved the determination of frequency (percentage of quadrats in which a
species occurred) and density (average number of stems or indivudals per
quadrat).
Photographs were taken along compass-lines A through L in an east-west
direction.
Twelve quadrats were also randomly selected for photographing.
Quadrat photos were taken 10 feet from the southeast stake in an east-west
direction, with a yard stick placed behind the northwest stake.

�-90-

Soils
Nine soil samples were taken during the pre-impoundment phase (August 18,
1969) and nine following the flooding of impoundment No. 1 (September 10,
1970). A core of the top six inches of soil was collected adjacent to five
randomly selected vegetative quadrats and along four compass lines in the
deep-water portion of impoundment No 1, with the aid of a soil auger.
These
samples were sent to the Colorado State University Soil Testing Laboratory
for analyses of the following items: pH, conductivity, lime, organic matter,
phosphorus, potash, zinc, iron, and texture.

Flooding

of Impoundment

Sites and Waterfowl

Use

Plans called for initial filling of each permanent impoundment from wells in
September and maintaining stable water levels on a year-around basis.Waterfowl use of the impoundments by species was to be observed and recorded at
two-week intervals from September until freeze up.

DESCRIPTION

OF AREA

The study area is located on the Bonny Reservoir State Recreation Area in
extreme east-central Colorado, about 25 miles north of Burlington.
Bonny
Reservoir is a Bureau of Reclamation facility devoted primarily to controlling
flood waters of the South Fork of the Republican River and Landsman Creek.
The specific study area lies on a bench above the bottom land of the South
Fork of the Republican River at the far west end of the Bonny Reservoir
property (Fig. 1). It covers about 75 acres and occurs on the south side
of the river above the inlet to the reservoir and directly south of the
pothole blasting study area (Work Planl,Job
14). The northern two-thirds
of the study area supported native, meadow-type vegetation (sedges, rushes,
grasses) that had been hayed and grazed in the past but not broken by the
plow.
The southern one':'thirdwas farmed prior to 1969, with small grains
as the predominant crop.
The study area is characterized by 0.3-0.5 percent slopes, well within the
less than one percent slope recommended in marsh development work.
Soil
Conservation Service soils maps and soil auger samples indicated a fairly
heavy soil capable of retaining surface water for extended periods.

RESULTS AND DISCUSSION
Literature

Review

A list of references searched and reviewed as part of this study are
presented in bibliographic form at the end of this report (Appendix A).
Most of these were abstracted on index cards. An additional 200 references
relating to waterfowl habitat preference, a major aspect of this study,
were abstracted and summarized in a "Review of Literature" in 1962 and will
not be included here (Hopper 1962). A firm background in this subject now
exists, but updating of the literature is needed.

�-91-

Selection

of Study Area

Bonny Reservoir was selected as the study area, largely because it is statecontrolled property and dev~lopment could begin almost immediately.
The
San Luis Valley site is on iand administered by the Bureau of Land Management.
This federal agency was receptive to our Division doing research
on a portion of the land on which they were planning to do waterfowl development work.
However, it was later determined that it would be about five
years before development funds would be available to them for this work.

Dike and Ditch Work
The dike and ditch work for the three shallow, permanent impoundments was
completed in March, 1969. Figure 1 comprises a map of the resulting dike
and ditch systems, while Table 1 presents a summary of some of the physical
features.
The construction work involved 7,755 feet of dike requ~r~ng 9,345 cubic
yards of earth fill for all three impoundments combined.
The dike system
for impoundment No. 1 was longer and contained more volume· of earth than
those of the other two impoundments.
The dike forming impoundment No.3
was much shorter than those for the other two impoundments, but required
practically as many cubic yards of earth fill as impoundment No. 1 because
of its greater height.
An earthen, emergency spillway was constructed for
impoundment No.3.
Construction costs for the three impoundments amounted to $4,901.00 including
dike and ditch work, spillway construction, and installation of water control structures.
Dike construction was figured at $0.50 per cubic yard of
earth work, while ditches were constructed at the rate of $0.15 per linear
foot. Spillway construction and installation of water control structures
were included as part of the dike construction costs.

Well and Pump Work
Well drilling, pump installation, and pumphouse construction for well No.1
were completed in September, 1968, but several corrections had to be made
by the contractor in October, 1968. Yield of the well was about 450 gallons
per minute (1 c.f.s.), only one-half of the desired capacity.
As a result,
a 7.5 horsepower pump was installed in place of the anticipated IS horsepower pump. Total cost of this facility was about $5,000.00.
Well No. 2 was completed during the fall of 1970. Its capacity and cost
was similar to that of well No.1.
It was fitted with a pumphouse in the
spring of 1971.

�-92-

Table

1.

Physical

Impoundment
No.

characteristics

of three permanent

Length

Dikes
Earth Fill

(Ft.)

impoundments.

(Cu. Yds.)

]J

Ditch Length

1

3,850

3,320

0

2

2,275

2,729

550

3

1,630

3,296

975

7,755

9,345

1,525

Total

1J Includes

10 percent

for settlement.

Pre- and, Post-impoundment
Pre-impoundment

(Ft.)

Studies

Vegetation

Forty-five species of plants were encountered in the 48 ~-square-meter
quadrats studied in impoundment No.1 during the period August 13-16, 1969
(Tables 2-3). We were unable to identify two of these plants, mainly
because of the lack of sufficient vegetative and flowering parts to work
with in keying them. Forty-four of the 45 species were found in the 32
quadrats on the native meadow north of the ditch that crosses impoundment
No.1
(Table 2), while only seven species occurred in the 16 quadrats
located on the previously farmed area south of the ditch (Table 3). All
seven species found on the farmed area, except green bristlegrass (Setaria
viridus) were also found on the native meadow portion.
Tables 2 and 3 show the frequency (percent of the total number of quadrats
of occurrence) and density (average number of stems per quadrat) for each
plant species encountered on the native meadow and farmed areas, respectively.
Only three plant species, all grasses, occurred in over 75 percent of the
32 native meadow quadrats.
These were inland saltgrass (Distichlis stricta)
(93.8%), bluestem wheatgrass (Agropyron smithii) (84.4%), and sand dropseed
(Sporobolus cryptandrus) (78.1%). Aster (Aster commutatus) was found in
56.2 percen~ of the quadrats, while western ragweed (Ambrosia coronopifolia)
and summercypress
(Kochia scoparia) each occurred in 50 percent of the
quadrats.
Six additional species were found in 25 to 50 percent of the
quadrats.
Thus, only 12 of the 44 species encountered on the native meadow
occurred in 25 percent or more of the 32 quadrats.
Inland saltgrass, bluestem wheatgrass, and cheatgrass brome (Bromus tectorum)
showed the greatest densities in the native meadow quadrats with averages of
168.5, 62.2, and 50.6 stems per quadrat, respectively.
Two other grasses,
sand dropseed and blue grama (Boutcloua gracilis) each had densities of about
31 stems per quadrat.
All other species exhibited average densities of less
than 25 stems per quadrat.

�Table 2. Frequency and density data for plant species encountered in 32 quadrats on the native
meadow portion of inpoundment No. 1, 1969-70.

Species 1/
Agropyron smithii
Ambrosia coronopifo1ia
Ambrosia e1atior
Andropogon gerardi
Aristada longiseta
Artemisia dracuncu1us
Artemisia fi1ifo1ia
Artemisia 1udoviciana
Asclepias incarnata
Asce1epias pumi1a
Asclepias speciosa
Aster commutatus
Aster spp.
Astragalus spp.
Bidens spp.
Boraginaceae family
Boute1oua gracilis
Boute1oua spp.
Bricke11ia ob1ongifo1ia
Bromus tectorum
Carex spp.
Cirsium spp.
Conyza canadensis
Danthonia spp.
Delphinium spp.
Distich1is stricta
E1ymus canadensis
Hap10pappus phy11ocepha1us

Frequency (Percent of Total
No. of Quadrats of Occurrence)
1969
1970
84.4
50.0
0

3.1
9.4
25.0
6.2
25.0
3.1
6.2
6.2
56.2
0
3.1
0
0
34.4
9.4
0
34.4
21.9
6.2
12.5
3.1
18.8
93.8
3.1
3.1

He1ianthus annuus

0

Hesperoch1oa spp.
Hordeum iubatum

0
0

84.4
25.0
3.1
3.1
6.2
21.9
6.2
15.6
3.1
6.2
6.2
34.4
12.5
3.1
3.1
12.5
31.2
6.2
3.1
25.0
25.0
0
6.2
0
3.1
100.0
3.1
3.1
6.2
3.1
3.1

Densi ty
Ave. No. of Stems/Quadrat
1969
1970
62.2
4.6
0
0.6
13.1
6.9

88.4
3.8
T
3.3
11.0
7.0

3.8
T 1/
0.6
0.1
5.8
0
T
0
0
30.6
8.7
0
50.6
17.2
0.1
0.7
0.2
0.5
168.5
0.1
T

2.9
1.6
0.3
0.3
3.4
0.9
T
T
1.2
40.8
7.9
0.1
30.8
36.5
0
0.3
0
T
425.7
1.6
0.1

0

0.1

0
0

T
4.1

-------------------------------------------------------------------------------------------------

I
1.0

w
I

�-96-

Sunnnercypressoccurred
in all 16 quadrats on the previously farmed area,
while common Russian thistle (Salsoli ,kali) was found in 87.5 percent and
lettuce (Lactuca ludoviciana) in 81.2 percent (Table 3). Ground cherry
(Physalis longifolia) was the only other species of the seven encountered
that occurred in more than one quadrat.
Sunnnercypress also had by far the
greatest density, averaging 1,070 stems, or in this case, individual plants
per quadrat.
These plants were noticeably stunted, as might be expected
with the extremely high density.
The density of connnon Russian thistle
(269 stems per quadrat) amounted to only about one-fourth that of summercypress but was greater than any other species in the native meadow during
the pre-impoundment phase of the study. All other species in the farmed
area had relatively low densities.
Species:area curves, as described by Oosting (1956) were applied to the
native meadow and farmed samples to determine the minimum number of
~-square-meter quadrats required for adequate sampling of vegetation on
each area. This procedure indicated that the numbers of quadrats used
(32 and 16, respectively) were sufficient for the present plant composition.

Post-impoundment

Vegetation

Flooding of impoundment No. 1 began on August 10, 1969, but as noted previously only partial filling resulted because of an insufficient quantity
of well water.
A stable level of water was maintained in the impoundment
throughout the remainder of 1969 and in 1970; however, because of the
lower than desired water level only a portion. of the vegetative study area
was subjected to increased moisture conditions.
The 48 vegetative quadrats
were again read during the periods August 19-20 and September 8-10, 1970.
Fifty-two species of plants were found in the 48 quadrats in 1970, an
increase of seven species over 1969 (Tables 2-3. These seven new species
occurred only in quadrats lying within the flooded area of the impoundment.
Apparently, they found moisture conditions more favorable in 1970 than in
1969. Fifty of the 52 species were located in the 32 native meadow quadrats,
with six observed in the 16 quadrats on the previously farmed area.
As in 1969, inland saltgrass, bluestem wheatgrass, and sand dropseed had
the greatest frequencies in the native meadow area, occurring in 100, 84.4,
and 53.1 percent of the 32 quadrats, respectively (Table 2). Other species
with frequencies of 25 percent or more included aster (34.4%), blue grama
(31.2%), sunnnercypress (31.2%), western
ragweed (25.0%), cheatgrass brome
(25.0%), sedge (Carex spp.) (25.0%), and prairie coneflower (Ratibida
columnifera) (25.0%).
Inland saltgrass, bluestem wheatgrass, and sedge were
the only species of the top 10 that did not decrease in frequency of
occurrence from 1969 to 1970. Most remaining species also showed a decrease
in frequency during this period.
Inland saltgrass and bluestem wheatgrass were again the donimnat species in
the native meadow quadrats in terms of denSity, averaging about 426 and 88
stems per quadrat, respectively.
Blue grama, sedge, and cheatgrass brome
e~hibited average densities of 40.8, 36.5, and 30.8 stems per quadrat.
All
of the above five species, except cheatgrass brome, showed increases in
density from 1969 to 1970, with the greatest increases occurring for inland
saltgrass and sedge.

�-97-

Summercypress,
lettuce, and common Russian thistle were the major species
in the farmed area in regard to frequency of occurrence in 1970, just as
they were in 1969 (Table 3). They occurred in 93.8, 87.5, and 56.2 percent
of the 16 quadrats, respectively; representing a decrease for all except
lettuce from 1969 to 1970. Two moist-environment
plants, sedge and softstem bulrush (Scirpus va1idus) made their appearance in 1970.
Density changes were quite apparent on the farmed area from 1969 to 1970.
Summercypress maintained its dominance in this regard, but was reduced by
nearly one-half from 1969. The greatest decrease in density was noted for
common Russian thistle, going from 269 stems per quadrat in 1969 to only
about 28 stems per quadrat in 1970. A tremendous increase in density was
recorded for lettuce in 1970. This species increased from an average of
only 3.6 stems per quadrat in 1969 to 541.6 stems per quadrat in 1970.
I
suspect the changes in density noted above for the farmed area had more to
do with the natural process of plant succession than to increased moisture
brought about by flooding the impoundment.
Few conclusive statements can be made regarding changes in plant frequency
and density from 1969 to 1970. This represented only bne year of flooding,
and only partial flooding at that. Nearly all species dropped in frequency
of occurrence from 1969 to 1970 on both the native meadow and farmed portions
of the study area.
Overall density increased on the native meadow from 1969
to 1970, due mostly to the large increase by inland saltgrass.
This species
no doubt reacted favorably to increased soil moisture caused by flooding of
the impoundment.
Natural plant succession was believed to be the reason
for density changes on the farmed area.

Soils
Results of the analyses of soil samples are shown in Tables 4-5. The 1970
samples (post-impoundment) were taken from within a few feet of the 1969
samples (pre-impoundment) and differences in soil properties between the
two years mayor
may not reflect changes resulting from the flooding of
impoundment No.1.
Sample number AO, EO, HO, and KO were taken from the
borrow area (deep water area) where a foot or more of the top soil had been
removed and used for construction of the dike. These four subsurface
samples were mostly clay loam in texture.
All other samples were taken
from the original soil surface and consisted of lighter textures of loams
and sandy loams. Generally, it seems that surface soils of impoundment No.
1 are light to medium textured, but that .at least some of the area is underlain with heavy soils.
Pre-impoundment samples collected in 1969 showed subsurface soils to be
similar to the surface samples in all respects except for lime and organic
content (Table 4). The clay loams were all high in lime content, whereas
the lighter surface samples ranged from low to high in this regard.
Organic
matter content was generally lower in the clay loam samples (0.8 - 1.3%)
than in the loam and sandy loam samples (1.8 - 3.6%).
This difference in
organic matter content would be expected when comparing surface and subsurface soils.

�Table 4. Pre-impoundment soil test results of nine samples collected in impoundment No. 1 on August
18, 1969.
Organic
Matter

Phosphorus

Potassium

(ppm)

(ppm)

Zinc
(ppm)

Iron

(%)

Conductivity
(salts)
(mmhos/cm)

High

1.3

0.6

8.7

417+

0.59

7.2

8.1

High

0.9

0.5

9.2

281

0.29

7.9

Clay Loam

7.9

High

1.1

0.5

7.4

417+

0.41

8.5

KO

Clay Loan

8.5

High

0.8

0.5

2.2

417+

1.38

6.5

B1

Loam

7.9

High

2.7

0.5

8.1

385

1.40

12.5

F1

Loam

7.5

Low

2.9

0.5

5.2

417+

0.92

6.0

Texture

pH
(paste)

Lime

AO

Clay Loam

7.9

EO

Clay Loam

HO

Sample
No.

(ppm)

I

\0
00
I

G2

Loam

7.7

Low

3.6

0.5

8.5

417+

1.90

4.8

J1

Loam

7.8

Low

2.5

0.3

4.1

417+

0.42

2.7

K2

Sandy Loam

7.9

High

1.8

0.4

3.3

417+

0.50

2.5

�Table 5. Post-impoundment soil test results of nine samples collected in impoundment No. 1 on September
10, 1970.

Texture

pH
(paste)

Lime

Organic
Matter
(%)

AO

Sandy Loam

7.7

Medium

1.3

EO

Clay Loam

7.6

Low

HO

Clay Loam

7.8

KO

Clay

Bl

Sample
No.

Conductivity
(salts)
(mmhos/em)

Phosphorus

Potassium

Irori

(ppm)

(ppm)

Zinc
(ppm)

"(ppm)

1.0

7.5

500+

0.52

24.2

1.0

0.7

4.3

500+

0.32

12.2

Medium

1.0

0.6

2.5

500+

0.46

14.2

7.8

High

1.2

0.7

2.0

500+

0.32

11.4

Clay Loam

7.6

Medium

2.3

0.8

15.0

500+

1.66

40+

Fl

Loam

7.4

Low

2.1

1.1

5.8

345

1.58

40+

G2

Loam

7.1

Low

2.2

0.7

6.5

418

1.26

·40+

Jl

Loam

7.7

Low

2.2

0.5

2.5

500+

0.42

4.8

K2

Loam

7.8

Medium

1.6

0.5

2.3

500+

3.54

4.2

I
\0
\0
I

�-100-

Hydrogen-ion concentrations
(pH) of all soil samples in 1969 ranged from
7.5 to 8.5, or slightly to moderately alkaline.
Samples taken on the
previously farmed portion of the impoundment (Jl and K2) showed smaller
amounts of phosphorus, zinc, and iron than did the samples collected from
the native meadow.
Many years of cropping on farmland can be responsible
for declines in these elements.
Samples taken in 1970, after one year of partial flooding of impoundment
No. I, indicated some differences from 1969 pre-impoundment samples (Table
5). Texture changes from 1969 to 1970 occurred for three of the nine
samples:
AO (clay loam to sandy loam), Bl (loam to clay loam), and K2
(sandy loam to loam). These changes may have been the result of siltation.
Conductivity, potassium,and
iron increased in the samples following
flooding; while pH and phosphorus decreased.
Lime content decreased in the
subsurface samples (borrow area) but remained the same for the surface
samples from 1969 to 1970. Sampling error, rather than induced changes
through flooding may have contributed greatly to the above differences.
All other soil properties tested were generally consistent between the two
years.

, Flooding

of Impoundment

Site

An attempt was made in mid-August, 1969 to fill impoundment No. 1 from the
initial well.
It was obvious after several days that the well did not
produce enough water to meet this demand.
A sufficient quantity of water
was yielded to fill the borrow area and to begin spreading over the flats,
but at least an additional six vertical inches were needed to obtain the
desired amount of flooded area.
The second well was operated in conjunction with the first well starting
in the fall of 1970. Together, they increased the water level in impoundment No. I, but failed to yield any excess water for impoundment No.2 and
3. This situation is completely contrary to study plans and no other
sources of water are presently available for use on the study area. Thus,
it is recorrnnended that this study be dropped and, if deSirable, another one
proposed which can make use of the existing impoundments and quantity of
water.

Waterfowl

Use

Intensive waterfowl use observations were scheduled to begin in the late
surrnnerand early fall of 1969 following initial filling of impoundment No.
1 to the desired level. However, only partial filling occurred and systematic waterfowl counts were delayed in anticipation that a desired water
level would eventually be reached.
Since the desired water level was never
realized, only general statements can be made regarding waterfowl use of
impoundment No.1.
Waterfowl began using impoundment No. 1 irrnnediately upon initial flooding
in August of 1969. Varying numbers of ducks were seen using the impoundment during the fall, with a peak of 43 observed on October 9, 1969. The
impoundment froze over on November 13, 1969, except a sizable area round the
inlet.
This open water area was maintained throughout the winter and spring

�-101-

by water flowing from the wells at temperatures ranging from 48-550 F.
Heaviest duck use occurred during this period, with excess of 200, mostly
mallards, observed at one time. Duck use of the impoundment continued
through the spring and summer of 1970, including breeding birds that
established territories. TJo or three broods of mallards were seen on the
impoundment during the evening of July 6, 1970.
Harvest checks were made on impoundment No. I during the fall of 1970.
Duck hunting pressure is generally light in the Bonny Reservoir area and
hunters were difficult to contact. Only 18 ducks, mostly teal, were checked
during six days of contactsi however, this number is believed to be only a
small fraction of the total number harvested in the 1970-71 season.

Prepared by

~/£t;; m~l
. Richard M. Hopper
Wildlife Researcher

�-102-

APPENDIX A
LITERATURE CITEb

Archibald, E. E. A. 1949.
J. Ecol. 37:260-288.
Ashby, E. 1948.
14:222-234.

The specific character of plant communities.

Statistical ecology.

II.

A reassessment.

Bot. Rev.

Borman, F. H. 1953. The statistical efficiency of sample plot size and
shape in forest ecology. Ecology. 34:474-487.
Bourdeau, P. F. 1953. A test of random versus systemati~ sampling.
Ecology. 34:499-512.
Cain, S. A.

1938.

The species-area curve.

Am. MidI. Nat.

19:573-581.

1943. Sample-plot techniques applied to alpine vegetation in
Wyoming. Am. J. Bot. 30:240-247.
Cochran, W. G.
206-233.

1963.

Sampling techniques.

Coker, R. E. 1954. Streams lakes ponds.
Press, Chapel Hill. 327p.

John Wiley and Sons, N. Y.

The University of North Carolina

Cottam, G., and B. W. Hale. 1953. Some sampling characteristics of a
population of randomly dispersed individuals. Ecology. 34:741-757.
Curtis, V. T., and R. P. McIntosh. 1950. The interrelations of certain
analytic and snythetic characters. Ecology. 31:434-455.
Cutler, D. D. 1955. A permanent plot system of survey for continuous
inventory of ponderosa pine stands in the southwest. J. For.
53(2):186-189.
Daubenmire, R. 1959. A canopy-coverage method of vegetative analysis.
Northwest Sci. 33(1):43-64.
Deming, W. E.
602p.

1950.

Some theory of sampling.

John Wiley and Sons, N. Y.

Evans, F. C. 1952. The influence of size of quadrant on the distributional
patterns of plant populations. Contr. Lab. Vert. BioI. Univ. Mich.,
54:1-15.
Goodall, D. W. 1952. Quantitative aspects of plant distribution.
Rev. Cambridge Philos. Soc. 27:194-245.

BioI.

�-103-

Grieg-Smith, P. 1952. The use of random and contiguous quadrants in
the study of the structure of plant communities.
Ann. Bot. NS.
16 (62):293-316.
1957. Quantitative plant ecology.
Publications, London.
198p.
Hall, O. F. 1959. The contribution
precision of growth estimates.
Hanson, H. C.

1950.

Scientific

of remeasured sample plots to the
J. For. 57(11):807-811.

Ecology of the grassland.

Harrington, H. D. 1954.
Denver.
666pp.
Hopper, R. M.
grounds.

Butterworths

II.

Bot. Rev.

Manual of the plants of Colorado.

16:283-360.

Sage Books,

1962. Habitat preference of waterfowl on the breeding
Unpubl. Review of Literature.
Colo. State Univ.
69pp.

1968. Determination of methods for developing and managing
waterfowl habitat"'--shallow impoundment study.
Colo. Game, Fish and
Parks Div., Fed. Aid Game Res. Rpt., Oct. pp. 43-49.
1969. Determination of methods for developing and managing
waterfowl habitat--shallow
impoundment study.
Colo. Game, Fish and
Parks Div., Fed. Aid Game Res. Rpt. Oct. pp. 65-71.
1970. Determination of methods for developing and managing
waterfowl habitat-shallow impoundment study.
Colo. Game, Fish and
Parks Div., Fed. Aid Game Res. Rpt. Oct. pp. 49-60.
Husch, B.

N. Y.

1963. Forest mensuration
474p.

and statistics.

Kelsey, H. P., and W. A. Dayton.
1942. Standardized
Horace McFarland Co., Harrisburg, Pa. 675pp.

The Ronald Press Co.,

plant names.

J.

Kuchler, A. W. 1955. A comprehensive method
Assoc. of Amer. Geog. 65:404-415.

of mapping vegetation.

Odum, E. P.
546p.

W. B. Saunders

1959.

Fundamentals

of ecology.

Oosting, H. J. 1956. The study of plant .communities.
Co., San Francisco.
440p.
Osborne, J. G.
cover-type

Annals.

Co., Philadelphia.

W. H. Freeman

and

1942. Sampling errors of systematic and random surveys
areas. J. Am. Stat. Assn.
37:256-264.

of

Pechanee, J. F., and G. Stewart.
1940. Sagebrush-grass range sampling
studies: size and structure of sampling unit. J. Am. Soc. Agron.
32:669-682.
Penfound, W. T. 1945. A study of phytosociological
relationships
of aggregations of colored cards. Ecology.
26:38-57.

by means

�-104-

Phillips, E. A. 1959.
Inc. 107p.

Methods

of vegetation

study.

Henry Holt and Co.,

Rice, E. L. 1952. Phytosotiological analysis of tall-grass
Marshall County, Oklahoma.
Ecology.
33(1):112-116.

prairie

___

Ecology.

., and R. W. Kelting.
36: 7-11.

1955.

The species-area

curve.

in

Schumacher, F. S., and R. A. Chapman.
1948. Sampling methods in forestry
and range management.
Duke Univ. School of Forestry Bull. 7, rev.
ed. 22lp.
Spurr, S. H.
340p.

1948.

Aerial photographs

in Forestry.

New York Ronald Press.

Swindale, D. N., and J. T. Curtis.
1957. Phytosociology of the larger submerged plants in Wisconsin lakes. Ecology.
38(3):397-407.
Vestal, A. G. 1949. Minimum
BioI. Monog.
30:1-129.

areas for different

vegetations.

Ill.

Weller, M. W., and C. S. Spatcher.
1965. Role of habitat in the distribution and abundance of marsh birds.
Lowa State Univ., Sci. and
Tech., Agr. and Home Econ. Exp. Sta., Spec. Rpt. No. 43. 31p.
Yates, F. 1953. Sampling methods for censuses
Publ. Co ,, New York.
p , 230-233.

and surveys.

Hafner

�-105-

APPENDIX

Scientific

and common names of plants

Scientific

Name

1/

Agropyron smithii
Ambrosia coronopifolia
Ambrosia elatior
Andropogon gerardi
Aristada longiseta
Artemisia dracunculus
Artemisia filifolia
Artemisia ludoviciana
Asclepias incarnata
Asclepias pumila
Asclepias speciosa
Aster commutatus
Aster spp.
Astragalus spp.
Bidens spp.
Boraginaceae family
Bouteloua gr&amp;C1Lis
Bouteloua spp.
Brickellia oblongifolia
Bromus tectorum
Carex spp.
Cirsium spp.
Conyza canadensis
Danthonia spp.
Delphinium spp.
Distichlis stricta
Elymus canadensis
Haplopappus phyllocephalus
Helianthus annuus
Hesperochloa spp.
Hordeum jubatum
Juncus spp.
Kochia scoparia
Lactuca ludoviciana
Liatrus punctata
Melilotus alba
Muhlenbergia asperifolia
Opuntia spp.
Panicum capillare
Physalis longifolia
Plantago ~.

B

referred

to in the text.

Common Name ]j

Bluestem wheatgrass
Western ragweed
Common ragweed
Big bluestem
Red threeawn
Sagebrush
Sand sagebrush
Louisiana sagebrush
Swamp milkweed
Plains milkweed
Showy milkweed
Aster
Aster
Milkvetch
Beggartick
Borage family
Blue grama
Grama
Mohave brickellia
Cheatgrass brome
Sedge
Thistle
Horseweed fleabane
Danthonia
Larkspur
Inland saltgrass
Canada wild rye
Goldenweed
Common sunflower
Spikefescue
Foxtail barley
Rush
Summercypress
Lettuce
Dotted gayfeather
White sweetclover
Alkali muhly
Prickly pear
Common witchgrass
Groundcherry
Plantain

�-106-

APPENDIX

B (Continued)

Scientific

and common names of plants

Scientific

Name

11

Polygonum aviculare
Ratibida columnifera
Salsola kali
Scirpus validus
Setaria viridus
Solanum rostratum
Solidago graminifolia
Sporobolus airoides
Sporobolus crytandrus
Stellaria media
Stipa comata
Tragopogon porrifolius

II From Harrington (1954).
11 From Kelsey and Dayton (1942).

referred

to in the text.

Common Name

1./

Prostrate knotweed
Prairie coneflower
Common Russian thistle
Softstem bulrush
Green bristlegrass
Buffalobur nightshade
Grassleaf goldenrod
Alkali sacaton
Sand dropseed
Chickweed
Needle and thread
Salsify

�-107October,

JOB PROGRESS

REPORT

COLORADO

State of
Project

1971

Migratory

W-88-R-16

No.

Bird Investigations

1
Job No.
14
Determination
of Methods for Developing and Managing
Job T it1 e __ W_a_t_e_r_f_o_w_l_H_a_b_i_t_a_t_-_A_mm_o._n_i_u_m_N_~_.
t_r_a_t_e_P_o_t_h_o_l_e_B_l_a_s_t_~_·
n"",g",--_S_t_u_d",-y
__

Work Plan No.

Period Covered:
Personnel:

April

Delwin

1, 1970 to March

Benson

and Richard

31, 1971

Hopper

ABSTRACT

Submerged aquatic vegetation was sampled in 36 potholes (9 of each charge
size) in September, 1970. Data from these samples were compared with those
collected from the same 36 potholes in 1968.
Submergents were found in 31
(86%) of the 36 potholes in 1968 and in 3S (97%) in 1970.
Six species were
encountered, with muskgrass (Chara spp.), longleaf pondweed (Potamogeton
nodosus),
and sago pondweed (R. pectinatus)
dominating in both frequency of
occurrence and density during both years.
Plant densities more than doubled
from 1968 to 1970 for each charge size except the ISO-lb. size, which produced
similar densities during both years.
Sago pondweed showed the greatest increase
in density from 1968 to 1970. Coontail (Ceratophyllum demersum) and longleaf
pondweed also increased in density from 1968 to 1970, while decreases were noted
for leafy pondweed (R. foliosus) and horned-pondweed
(Zannichellia palustris).
A manuscript entitled "Waterfowl Use in Relation to Size and Cost of Blasted
Potholes" was prepared and submitted to the Journal of Wildlife Management in
September, 1971.
If accepted for publication,
this paper will satisfy Federal
Aid reporting requirements for a portion of this study. Major points brought
out in this paper include:
(1) The 2S- and 7S-lb. charges produced more square
feet of surface area/lb. than the SO- or ISO-lb. sizes; (2) Total cost/lOO
sq. ft. of surface area was lowest for7S-lb.
charges, followed by the ISO-lb.
charges; (3) most duck use occurred in the spring; (4) 10 species of ducks
used the potholes with mallards (Anas platyrhynchos)
and blue-winged
teals (Anas
discors) contributing most of the duck visits; (S) Potholes produced by 7Sand ISO-lb. charges yielded the largest number of average duck visits/hr. of
the four charge sizes; and (6) The 7S-lb. charge was the most efficient in
regard to cost/duck visit and number of duck visits/100 sq. ft. of surface
area, followed by the ISO-lb. size.

�-108-

RECOMMENDATIONS

I.

Both 75 lb. and 150 lb. charges are recommended over 25 lb. or 50 lb.
charges for use in pothole blasting.

�-109-

AMMONIUM

NITRATE

POTHOLE

BLASTING

STUDY

Richard M. Hopper

P. S. OBJECTIVE
To evaluate various size potholes blasted with ammonium nitrate in terms
of (a) life expectancy, (b) plant succession, (c) soil and water characters,
(d) waterfowl use and hunting potential, and (e) cost.

SEGMENT OBJECTIVES
1.

Collect data pertaining to the vegetative,
associated with the potholes.

2.

Obtain pothole measurements.

3.

Collect waterfowl

use and harvest

METHODS

soil, and water

data on and adjacent

characteristics

to the potholes.

AND MATERIALS

Methods and materials employed during Segment 16 remained
used in Segment 14 (Hopper 1969) and will not be repeated

the same as those
here.

RESULTS AND DISCUSSION
Vegetative,

Submerged

Vegetation

Soil and Water Characteristics

in Potholes

Submerged aquatic vegetation was sampled in 36 potholes (nine of each charge
size) during the period September 9-10, 1970. Sampling consisted of dragging
a garden rake through each pothole the following number of times according to
charge size:
25 lb. - 3; 50 lb. - 4; 75 lb. - 6; 150 lb. - 8. One drag of
the rake through a pothole constituted one sample; thus 189 samples were taken
in the 36 potholes.
Samples obtained in 1970 were compared by charge size of
potholes with those collected from the same 36 potholes in 1968 to give an
indication of plant succession (Tables 1-2).
Six species of submergents were found in the sampling process during the two
years of vegetative study, five in 1968 and the same five plus coontail
(Ceratophyllum demersum) in 1970. Vegetation was encountered in 31 (86.1%)
of the 36 potholes in 1968, with one 25-lb., three 50-lb., and one l50-lh.
potholes being void of submerged
aquatics in the samples.
All but one (150
lb.) of the 36 potholes (97.2 percent) produced vegetation in the samples in
1970.

�Table 1. Comparison of frequency of occurrence of submerged aquatic vegetation among charge sizes, as
estimated from 189 samples in 36 potholes.

25
Species

Freguency of Occurrence (Percent) 1/
Charge Size (lbs.)
50
75
150
1968
1970
1970
1968
1968

Total
1970
1968

1968

1970

0

0

0

·5.6

0

7.4

0

23.6

0

12.2

Chara spp.

44.4

33.3

5.6

41. 7

55.6

55.6

68.0

16.7

49.2

34.9

Potamogeton fo1iosus

25.9

3.7

25.0

8.3

53.7

24.1

55.6

13.9

45.0

14.3

Potamogeton nodosus

33.3

44.4

11.1

19.4

31.5

48.1

40.3

45.8

31.2

41. 3

Ceratophy11um demersum

1970

I

Potamogeton pectinatus

0

66.7

44.4

72.2

24.1

72.2

19.4

47.2

22.8

61.9

Zanniche1lia pa1ustris

7.4

3.7

0

0

18.5

0

0

0

6.3

0.5

Total

77.8

100.0

66.7

91. 7

92.6

100.0

88.9

84.7

84.1

92.6

1/ Percentage of total number of samples in which a species occurred.

I-'
t-'

c
~

�Table 2. Density comparisons of submerged aquatic vegetation among charge sizes, as estimated from 189
samples in 36 potholes.

25
Species

Mean Density 1/
Charge Size ~lbs.2
50
75
1968
1970
1968
1970

1968

1970

0

0

0

0.3

0

Chara spp.

1.9

3.0

0.2

2.7

Potamogeton fo1iosus

2.1

0.1

1.6

Potamogeton nodosus

2.0

3.5

0.4

Ceratophy11um demersum

Potamogeton pectinatus

0

7.4

2.7

150

Total
1968
1970

1968

1970

0.3

0

2.4

0

1.1

3.1

5.2

6.1

1.3

3.5

2.9

0.9

3.6

3.0

4.0

0.6

3.1

1.2

1.7

1.3

4.5

1.6

3.1

1.4

3.3

7.8

1.1

7.5

1.4

6.0

1.3

7.0
T ];./

,....I
,....
,....
I

Zanniche11ia pa1ustris

0.2

0.2

0

0

1.0

0

0

0

0.3

Total

6.2

14.2

4.9

13.4

10.1

20.5

13.1

13.4

9.6

1/ Average number of rake teeth covered per sample.
];./Less than 0.1.

15.5

�-115October,

JOB PROGRESS

State of
Project

1971

REPORT

C~O~L~O~RAD~~O~~ _
No.

Work Plan No.
Job Title
Period Covered:

Migratory

W-88-R-16
2

Bird Investigations

Job No.
Experimental Studies on
Improving Status of Canada Goose Populations
April

2

1, 1970 to March 31, 1971

Personnel:
D. Horn, Bureau of Sport Fisheries and Wildlife; Delwin Benson,
W. Carpenter, C. Crawford, G. Crawford, G. East, H. Gresh, D. Hopper, D. Potts,
J. Monarch, C. Slonaker and M. Szymczak of Colorado Division of Game, Fish and
Parks.

ABSTRACT

In the spring of 1970, the first nesting attempts by Canada geese were noted
in the goose transplant site area near Masters, Colorado.
Sixty goslings
were released on the K-4 Ranch in August, 1970, bringing the total number
of birds released in three years to 346. Band recoveries indicate that most
of the birds released at Masters are remaining in Colorado throughout the
year. As many as 80 geese were observed at the San Luis Lake transplant site
in the spring of 1970. Non-breeding birds from the San Luis Valley population,
including transplanted birds from all years since 1967, were members of a
moulting population captured at Wheatland Reservoir in Wyoming in June of 1970.
Band recoveries indicate that birds released at San Luis Lake may be establishing a fall migration pattern into southwest New Mexico.
Geese began returning
to the North Park transplant area on April 11, 1970. Birds were sighted intermittently throughout the Park through the spring period.
In July, 246 goslings
were released at Lake John Annex.
After the release, geese were commonly observed on the Annex, at Case Flats, at Hebron Sloughs and on Walden and
McFarlane Reservoirs.
Band recoveries indicate that the birds released in
North Park in 1969 may have had a low survival rate through the first winter.
Geese released in 1970 in North Park were recovered primarily in Yuma County,
Arizona, exhibiting a similar migration pattern to those released in 1969.
Approximately
10 percent of the 1970 North Park transplants were reported recovered by hunters.
In June, 1970, 36 goslings were released on Totten Reservoir east of Cortez in Montezuma County.
The birds remained on Totten or at
nearby Narraguinnep Reservoir throught the fall and winter period.

�RECOMMENDATIONS

1.

If goslings are available, release birds at North Park, San Luis Lakes,
Jumbo Reservoir and Totten Reservoir in 1971.- -

�-117-

EXPERIMENTAL STUDIES ON
IMPROVING STATUS OF CANADA GOOSE POPULATIONS
Michael R. Szymczak

P. S. OBJECTIVE
To investigate the success of attempts to establish breeding populations
of Canada geese in suitable habitat where they do not currently exist in
Colorado.

SEGMENT OBJECTIVES
1.

To prepare a technical bulletin on the technique of establishing a
breeding population of Canada geese.

2.

To expand the breeding flocks of Canada geese in the South Platte
Valley, the San Luis Valley and North Park.

3.

To begin efforts to establish a breeding flock of Great Basin Canada
geese on Totten Reservoir, south of Dolores in Montezuma County.
METHODS AND MATERIALS

Canada goose goslings for transplant purposes were obtained.from four different sources: (1) goslings and some eggs from the metropolitan Denver area;
(2) goslings and some eggs from the Fort Collins area; (3) goslings from
Valmont Reservoir in the Boulder area and; (4) eggs from nests in danger of
destruction from high water along the Yampa and Little Snake Rivers in Moffat
County. All eggs were transported to the Fort Collins Wildlife Research
Station for incubation and subsequent raising. All goslings live-trapped
were transported as soon as possible to the release site. Birds obtained
from eggs collected in Moffat County were released on Totten Reservoir south
of Dolores. Birds collected at other locations were released either at San
Luis Lakes, Lake John Annex or the K-4 Ranch.
Migration and mortality information were obtained from recovery cards received from the Migratory Bird Population Station.
RESULTS AND DISCUSSION
Preparation of Technical Bulletin
Very little progress was made on preparing a technical bulletin on the technique of establishing a breeding popUlation of Canada geese during Segment 16.

�-118-

Extensive work on this bulletin is planned for Segment 17.

South Platte Valley - K-4 Ranch
Spring and Summer Status
In early April birds were encountered at the release site on the K-4 Ranch,
along the Platte River near Masters, Colorado, in addition to other nearby
locations. A number of nesting attempts were recorded with the result of
at least four broods being hatched. A production survey on June 27 recorded one brood of three on Empire Reservoir, one brood of four on the
Gallagher Ranch and two broods with a total of seven young on the K-4
Ranch. Additional broods may have been present in the area but were not
encountered. Nesting pairs may have been two-year-01d birds which had
survived since the original transplant made in the summer of 1968 or possibly
migrant birds which had stayed in the area.
On August 2, 60 goslings (30 females and 30 males) were released on the K-4
Ranch bringing the total number of geese released in that area in three years
to 346.
Hunting season recoveries of the transplanted geese continue to indicate that
most of the birds are remaining within the state throughout the year (Table 1).
Birds transplanted in the summer of 1968 and 1969 were recovered at a greater
rate during the 1970-71 season than they had been in previous years.

Table 1. Hunting recovery locations of Canada geese released on the K-4
Ranch near Masters, Colorado in Weld County.
Year
No. Released

1968
217
No. Recovered
Direct
Indirect

1969
69
No. Recovered
Direct
Indirect

Morgan Co.

5

0

1

0

0

N.A.

Weld Co.

0

5

1

2

1

N.A.

Adams Co.

1

2

0

2

1

N.A.

Larimer Co.

0

4

0

1

1

N.A.

Kansas

0

1

0

0

0

N.A.

Oklahoma

0

1

0

0

0

N.A.

Total

6

13

2

5

3

N.A.

Area Recovered

1969
60
No. Recovered
Direct Indirect

Colorado

�-119-

The times and locations of the recoveries indicate that many of the birds
had dispersed from the release site prior to the start of the hunting
season. Most of these bird~ apparently joined other concentrations of
geese in the Larimer and Weld County area. The movement of birds away
from the transplant site, as indicated by band recoveries, may be an
indication of rejection of the transplant site by some of the birds.
On the other hand the surviving birds may return to the transplant area
the following spring to breed, thus undergoing a short b~t important
east-west migration. Continuation of this type of migration, if occurring,
would definitely be detrimental to efforts to obtain a wider distribution
of wintering geese in north-central Colorado.
Approximately 300 square miles around the transplant site remained closed
to goose hunting during the 1970-71 season in order to protect the transplanted birds and encourage them to remain in the area.

San Luis Valley ~ San Luis Lakes
In July of 1969, 160 Canada goose goslings were released on San Luis Lake
in the San Luis Valley. Birds were placed at San Luis in an attempt to
expand the breeding goose population into the northeastern portion of the
Valley. First year recoveries of released birds were reported on by
Szymczak, 1970. The transplant site was not extensively surveyed for
returning birds in spring of 1970. However, as many as eighty birds were
observed on the release area during that spring. Many of these were probably 1969 transplant birds. An additional 7B,goslings were released on
the site in the summer of 1970.
In June of 1970, five of the 1969 transplants were captured in a goose
drive trapping operation on Wheatland Reservoir north of Bosler in Albany
County, Wyoming. This reservoir has traditionally been used by non-breeding
birds of some populations of Pacific Flyway Great Basin Canada geese as a
moulting area. In addition to the 1969 plants, seven of the goslings
released in 1967 and 1968 in the San Luis Valley on the Alamosa National
Wildli·fe Refuge were also captured. These .recap tures suggest that birds
of the San Luis Valley population may be beginning to use Wheatland for
moulting purposes. However, it is also possible that the birds recaptured
at Wheatland may have drifted into Great Basin goose population areas and
subsequently have become members of those popUlations. The recovery of a
1967 San Luis Valley transplant along the Green River in Utah during the
1970-71 hunting season adds to the validity of the latter hypothesis.
The recovery locations of goslings released at San Luis Lake are presented
in Table 2. Only eight recoveries of 1969 released birds were reported
during the 1970-71 reporting year. Two of these were taken during the
1969-70 season. The recoveries of birds during the 1970-71 season outside
the San Luis Valley, although small in number, substantiate the variation
in recovery locations of birds released on different dates in July, 1969
that was suggested by Szymczak, 1970. Most birds released on July 22 were

�-120-

recovered in Pacific Flyway areas while those released on July 8 seemed
to have migrated into New Mexico, wintering in the Roswell area of Chaves
County in southeast New Mexico. Only one bird of the 1969 plant was
actually bagged in the Roswell area during the 1970-71 season. However,
an additional eight birds from that release were recaptured on January
1, 1971 during goose trapping operations on the Bitter Lake National
Wildlife Refuge near Roswell. Of the nine birds known to be in the
Roswell area during the winter of 1970-71, only one was a member of the
July 22, 1969 group.

Table 2. Hunting season recovery locations of Canada geese released in
1969 and 1970 at San Luis Lake, Alamosa County, San Luis Valley, Colorado.
1969
Date Released
No. Released

1970

July 8

July 22

80

San Luis Valley
Chaffee Co.

80

June 29
29

July 2
39

2

3

o

2

1

o

o

o

Chaves Co.

9

o

1

San Miguel Co.

o

1

o

o
o

o

1

o

o

o

6

o

o

o

1

o

o

1

o

o

o

13

12

1

2

Area Recovered

United States
Colorado

New Mexico

Arizona
Cochise Co.
Mexico
Sonora
Southwest
Chihuahua
Unknown
Canada
Saskatchewan
Southwest
Total

�-121-

Thirty of the birds released in 1970 on San Luis were also present in the
Roswell area during the winter of 1970-71. Twenty-nine of the thirty were
also recaptured at Bitter Lakes while one was bagged by a hunter in the
vicinity.
The Roswell area/was the only location outside of the San Luis
Valley where 1970 transplants were harvested or recaptured.
The presence
in the Roswell area of 1969 transplants for two consecutive winters along
with 1970 releases during the winter of 1970-71 indicates that a portion
of these transplanted birds may be establishing a migration pattern into
southeast New Mexico.
The entire San Luis Valley with the exception of the San Luis Lakes-Head
Lake transplant area was open to goose hunting during the 1970-71 season.
The western portion of the Valley was open to hunting,by permit only, with
a maximum of 200 geese the allowable harvest.
The additional open season
apparently increased the hai~est of transplanted birds in the Valley.
However, the increased harvest should not be detrimental to the restoration
program at this time.
San Luis Lake proper lacks natural nesting sites for Canada geese.
Therefore, a number of artificial nesting structures, both pole-type and floatingtype were placed at San Luis prior to the return of geese in the spring of

1970.

North Park - Lake John Annex
In the summer of 1969, 194 goslings were released on Lake John Annex eight
miles west and 2 miles north of Walden.
The majority of these birds migrated
into the Pacific Flyway generally terminating their southern movement in
southwest Arizona and adjacent Mexico (Szymczak, 1970).

Spring Returns
Because of the variation in band reporting rates by geographic area, it is
difficult to estimate the number of birds which survived throughout the 196970 winter and thus were potentially eligible for return to North Park the
following spring.
Since many of the birds apparently move into Mexico, where
reporting rates are very low and hunting regulations are either non-existent
or not abided by, there is a good chance that a large number of the 1969
releases did not survive.
Some of the birds recovered in Mexico were caught
by hand indicating that they were either unwary or possibly unhealthy.
On April 11, 1970 the first returning birds were observed in North Park.
These 11 birds were sighted on Lake John Annex approximately three days after
the ice had begun to break up. Three of the birds were neckbanded.
Fourteen
geese were observed on the Annex on April 13 and 17, on April 20 and again
April 25. On April 20, the 17 birds were observed at close range and only
14 of the birds bore legbands. Assuming that the transplants did not lose
their legbands, the unbanded birds may have been members of the remnant
population of geese in North Park (Szymczak, 1970) or birds from other populations that accompanied the transplants into the Park.

�-122-

On May 6, 20 geese were observed on the Annex with 19 there on May 18, but
still only 3 birds with neckbands. Since 80 percent of the birds released
were neckbanded the geese apparently did not retain their neckbands to the
degree expected. One neckbanded bird, which was recovered in Arizona
during the 1969-70 hunting season, definitely was not wearing a neckband
when bagged.
In late May, 34 birds were observed on the Riley Ranch along the North
Platte River, 5 miles east of Lake John Annex. This was the largest group
of birds sighted in the spring.

New Releases
On July 2, 1970, ten goslings were released at the transplant site on Lake
John Annex. These ten birds were joined by an additional 113 birds on July
24 and 123 more on July 31. Throughout the remainder of the summer and
early fall birds were commonly sighted on Lake John Annex, Case Flats and
Hebron Sloughs. Birds were also observed at Walden and MacFarlane Reservoirs.
An occasional sighting was made on the Delaney Butte Lakes, Boettcher Lakes
and Fisher Lake. All areas on which geese were sighted are shown in Fig. 1.
A group of 79 birds, the largest flock sighted throughout the late summer
and fall, was observed in the Lake John area on October 15, 1970. After
that time 19 were sighted on the Annex on October 31 with eight, eight and
five birds sighted in the same area on November 24, 27 and 30, respectively.
No geese were observed in the Park after November 30.
Band Recoveries and Sightings
1969 Transplants--Only five of the 1969 transplants were reported recovered
during the 1970-71 hunting season (Table 3). An additional three neckbanded
birds were observed at Two Buttes Reservoir in southeast Colorado during the
season. Only two of the 1970-71 recoveries were taken in the Pacific Flyway. One bird was taken in Mesa County, Colorado and the other in Mohave
County, Arizona. The bird taken in Arizona had been recaptured at Wheatland
Reservoir, Wyoming in June of 1970 and when taken was apparently accompanied
by a 1970 transplant which was also bagged. This Arizona recovery probably
indicates a return of at least a portion of the surviving 1969 transplants
to their major 1969-70 wintering area.
Two of the other three recoveries were taken in eastern Montana and the
final recovery in Emmons County, North Dakota. The bird taken in North
Dakota had been recaptured the previous winter at two Buttes Reservoir in
southeast Colorado.

�-123-

z
o

..•

;;:r

-n
o

Fd schez,

c&gt; Lake

('!!.
Delaney
But te Ls.

-=_",:"!,,,",~D Wolden

l.,I OE

\)! .
S.

o

Case 4!J"
Flats

Peterson
Pond

()~

a

Hebron
00 ##
SioughsoMe forlone
Res.
SCALE

o
Fig. 1.

2

4 Mi.

Areas at which Canada geese have been sighted in North Park.

�Table 4. 1970-71 hunting season recoveries of North Park birds released in 1970 plotted according to time
and approximate distance from release site.

Area Recovered

1-10

October
11-20

21-31

Period of Recovery
November
December
1-10 11-20
21-30
1-10
11-20

1

2

1

21-31

January
1-10
11-20

United States
Wyoming
Carbon Co.
Arizona
Mohave Co.

1

I

t-'

Yuma Co.

2

11li

1

California
San Bernardino Co.

1

Imperial Co.

1

1

Mexico
Chihuahua
Northwest

1

l/ Eight birds recovered during the hunting season, Nov. 26-Jan. 10, exact dates unknown.

1

N
0\
I

�-127-

Table 5. First year hunting recovery locations of Canada geese released
in 1970 at Lake John Annex, Jackson County, Colorado.

Date Released
No. Released

July 2
10

July 24
113

July 31
123

Area Recovered

No. n.ecovered

No. Recovered

No. Recovered

United States
Wyoming
0

3

2

Mohave Co.

O·

1

0

Yuma Co.

0

8

6

San Bernardino Co.

0

1

0

Imperial Co.

0

2

0

0

1

1

o

16

9

Carbon Co.
Arizona

California

Mexico
Chihuahua
Northwest

Total

LITERATURE CITED
Szymczak, M. R. 1970. Experimental studies on improving status of Canada goose
populations. Colo. Div. Game, Fish and Parks, Game Res. Rep., Fed. Aid
Project W-88-R. October. 81-93 pp.

Prepared by

~~£1?

~

Michael R. Szymczak ~
Asst. Wildlife Researcher

~

-'-

�-129october,

1971

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-88-R-16

Work Plan No.

2

Migratory

Bird Investigations
5

Job No.

Job Title

Arkansas

Valley Canada Goose Flock Management

Period Covered:

October 31, 1970 to March

Studies

31, 1971

Personnel:
Velma Fredrickson, Ann Leckler, Marie Vendeville, M. Robinson,
G. Nugent, H. Moorhead, R. Cooper, F. Roth, W. Piper, M. Hickey, J. Wheeler,
B. Widhalm, A. Heins, B. Goetze, L. Green, D. Bogart, R. Kitzmiller, G. Bock,
D. Croonquist, R. Forbes, P. Olsen and M. Szymczak.

ABSTRACT
Water levels of most reservoirs in southeast Colorado were near or at
capacity throughout the fall and winter period.
Food conditions in southeast Colorado were generally considered to be good. The hunting season was
characterized by above normal temperatures and below normal precipitation. A
small movement of geese into southeast Colorado was noted about October 15,
with a substantial movement on October 30, followed by a major arrival on
November 17. Goose numbers in southeast Colorado varied from near 90,000 in
late November and early December to 62,720 on January 19. The change from
morning only hunting, to full day hunting on December 12 did not stimulate
an immediate, large movement of geese out of the state. The mid-winter inventory of the entire short grass prairie population totaled 145,349.
Generally,
geese left reservoirs to feed twice a day regardless of moon phases or the
hunting regulation in effect.
Extreme cold temperatures seemed to delay or
completely eliminate morning feeding flights. According to check station data,
approximately 61 percent of the geese bagged were immatures.
There were no
indications that feeding flights, in respect to time of day, were age-specific.
An estimated record 30,329 geese were bagged by 7,875 hunters throught the
1970-71 hunting season in southeast Colorado.
At the Two Buttes Management
area 610 hunters bagged 919 geese during the half-day portion of the season,
while 788 hunters bagged 814 geese during full-day hunting.
At the LamarEads Management area 705 hunters bagged 304 geese throughout the season.
Bands
were placed on 536 geese at Two Buttes Reservoir and 300 geese at Verhoeff's
Pond during postseason trapping operations.
Approximately 36 percent of the
birds banded were immatures.
Birds trapped at Verhoeff's weighed an average of
approximately one pound more than those trapped at Two Buttes.
Sixty-one
percent of southeast Colorado banded birds reported recovered during the 19691970 season were bagged in Canada.
An abnormally low twenty percent of banded
birds taken during the 1969-1970 season were reported taken in Colorado.
The
annual hunting mortality rate of all age classes combined continues to be
approximately 26 percent.

�-130-

RECOMMENDATIONS

1.

Reduce the number of aerial surveys of geese in southeastern
conducted under this job.

2q

Discontinue the collection, under this job, of information concerning
hunting pressure, wounding loss, and hunter success at the Two Buttes
and Lamar-Eads Management Areas.

3.

Discontinue
season.

observation

of feeding

flight patterns

during

Colorado

the hunting

�-131-

ARKANSAS

VALLEY

CANADA GOOSE FLOCK MANAGEMENT
Michael

STUDIES

R. Szymczak

The Arkansas Valley goose study continues to emphasize the evaluation of
the effects of experimental harvest regulations on the distribution of
the short grass prairie Canada goose population and the general behavior
of the southeast Colorado flock. During the 1969-70 hunting season,
morning only shooting showed promise a.s a tool to increase the percent
of the total short grass population wintering in Colorado.
However, the
many undesirable aspects of this regulation, mainly pertaining to the
hunting public, necessitated alteration of the regulation the following
year.
During the 1970-71 season, the framework for the goose season in
southeast Colorado was as follows: morning-only hunting from October 31,
1970 through December 11, 1970; full day hunting from December 12, 1970
through January 17, 1971. An evaluation of the regulation along with the
results of the collection of other data pertinent to the management of the
short grass prairie goose population is presented in this report.

P. S. OBJECTIVE
To collect management information on the wintering short grass pralrle
Canada goose population and evaluate new approaches to goose management
in southeast Colorado.

SEGMENT OBJECTIVES
1.

To determine the time of movement of geese into the Arkansas
and the size and distribution of the wintering flock.

2.

To obtain an estimate

of the age composition

3.

To determine
loss.

pressure

4.

To determine the annual mortality
pattern of the flock.

5.

To evaluate the effects of hunting from one-half hour before sunrise
until noon from October 31, 1970 through December 11, 1970, followed
by one-half hour before sunrise to sunset hunting from December 12,
1970 through January 17, 1971 on population numbers distribution,
harvest and flight patterns of Canada geese in southeast Colorado.

hunting

Valley

of the flock.

on the flock and the associated

and the migration

wounding

and/or harvest

�-132-

METHODS

AND MATERIALS

Periodic aerial counts of Canada geese were conducted in southeast Colorado.
The December 11 and January counts were coordinated with ground and aerial
counts made by various state and federal personnel throughout the wintering
grounds of the short grass prairie Canada goose population.
These counts
were made for the purpose of determining the size and distribution of the
wintering goose population.
Check stations were operated at the Two Buttes Management Area and the
Lamar-Eads Management Area to collect information on hunting pressure and
harvest.
Tail fans from geese bagged were collected at all check stations
with some additional fans obtained throughout southeast Colorado.
The
weights of bagged birds were also recorded.
Goose feeding flights were observed at various locations throughout southeast Colorado with information on time of flight, numbers of geese and
weather conditions recorded.
A total of 836 geese were trapped and banded at Two Buttes Reservoir and
Verhoeff's Pond after the close of the hunting season to obtain information
on age ratios, mortality rates and migration patterns.
The effects of past hunting season regulations
ations for the 1971-72 season were made.

were evaluated

and recommend-

RESULTS AND DISCUSSION

Water,

Food and Weather

Conditions

Reservoirs utilized by geese as resting areas in southeast Colorado presented
a variety of water levels during the 1970-71 season.
Most reservoirs continued to be near capacity, as during the 1969-70 season, in spite of generally
below average precipitation throughout southeast Colorado (Table 1). However,
John Martin, which in many years is drained completely for irrigation purposes,
and Two Buttes, fed only by local precipitation and runoff, showed water
levels lower than those present during the 1969-70 season.
The major reservoirs in the Eads area were characterized by extreme high
water.
The water eliminated the possible use as a resting area of the south
bar on Nee Noshe which had been used steadily by the geese the previous season.
Geese staying on Nee Noshe generally used the north shore area for resting.
It is speculated that the extreme high water in reservoirs in the Eads area
may have, in part, resulted in fewer birds using the area during the 1970-71
season than during the previous year.

�-133Table 1. Yearly precipitation
totals for selected southeast Colorado
stations for 1970 and the water status of associated reservoirs.
Station Location
and
Associated Reservoirs
Rocky Ford

1970 Precipitation
(In.)
Departure
Total
from Norm
9.03

Near
Near
Near
Near
Near
12.86

Horsecreek
Blue Lake

Reservoir

Eads

12.25

capacity
capacity
capacity
capacity
capacity

No
No
No
No
No

open
open
open
open
open

bars
bars
bars
bars
bars

Same as
1969-70

Some open, bars
available

Higher than
1969-70

No open bars

Much lower
than 1969-70

Some open bars
available

Same as
1968-69

Open shoreline
available; no bars

Water at
capacity

No open bars

-1.53

Swede Lake
Sweetwater

Areas

+0.61

Reservoir

John Martin

Resting

-3.28

Dye Reservoir
Holbrook Reservoir
Cheraw Reservoir
Meredith Reservoir
Henry Lake
Las Animas

Water Level

weather

Reservoir

Nee Granda

Very small pool Shoreline

Flamingo

Lower than
1969-70

Some open bars
available

At capacity

No open bars

Lake

Nee Noshe

open

Upper Queens

Reservoir

At capacity

No open bars

Lower Queens

Reservoir

At capacity

No open bars

Near capacity

No open bars

Lower than
1969-70

Open bars
available

Lamar
Thurston

13.55

-0.65

Reservoir

Springfield
Two Buttes

14.28
Reservoir

-0.45

�-134-

John Martin Reservoir was nearly drained completely during the summer of
1970 and provided apparently excellent habitat for the geese during the
fall period.
In addition to open bars, water running into the reservoir
provided continuous floodi.ng of extensive. stands of smartweed (Polygonum
sp.) and cocklebur (Xanthium sp.) which were apparently attractive to the
birds.
The decline in the water level at Two Buttes, resulting from below average
rainfall, produced a series of open bars, void of vegetation and extremely
attractive to the geese.
Food conditiomin
southeast Colorado during the 1970-71 season were variable
and probably not as good as during the previous year.
The dryland milo was
hampered by the lack of spring and early summer moisture.
However, adequate
fall moisture resulted in the establishment of good winter wheat crops, the
mainstay of the diet of geese in southeast Colorado.
A summary of weather conditions
throughout the hunting season is presented
in Table 2. In general, temperatures were above normal and precipitation
below normal; conditions generally not believed to be most conducive to
successful goose hunting.

Migrational Movements
and Distribution of the Wintering

Population

Canada geese normally arrive en masse in southeast Colorado sometime between
November 5 and November 15, depending on weather conditions on the Canadian
staging areas (Grieb 1970; Rutherford 1970). The 1970 fall migration of the
short grass prairie population from staging to wintering areas was definitely
atypical.
Some birds began arriving in southeast Colorado as early as
October 15 followed by a substantial movement into the area about October 30.
On November 9, at least 25,000 geese were present.
About November 17, the
major portion of the migrant birds moved into the area with the first aerial
survey on November 23 recording about 90,000 birds (Table 3). Early season
movement by a portion of the population may have been the result of a cold
period on the Alberta-Saskatchewan
staging areas (Alex Dzubin, pers. comm.)
Normally the first cold period on the staging area moves the birds from the
lakes and reservoirs to the rivers, but not to southern wintering areas.
An effort was made to detect any large movement of birds out of southeast
Colorado stimulated by a change from morning-only hunting to full day hunting.
Aerial surveys were conducted on December 11, the day before the change, and
again on December 23, after the geese had been subjected to two weekends of
full day hunting.
A decline in the population between these two flights was
recorded, but not considered significant (Table 3). An early January count
was scheduled but not conducted because of prohibitive flying weather.
However, comparing the results of the December coordinated inventory with the
January inventory, a definite movement into Texas is indicated sometime between
these two surveys (Table 4). Apparently a substantial number of birds moved
out of Colorado into Texas in late December or early January, a period which
encompassed unusually cold weather with some locally heavy snows. A combination
of continued full day hunting, along with the weather condition may have resulted in this movement.

�-135-

Table 2. Summary from selected southeast Colorado weather stations, .
November 1970 through January 1971.
Temperature
Average
Departure from
Maximum Minimum
Normal Daily Ave.

Precipitation
Monthly
Departure
Total 1/
from Norm

Rocky Ford

58.2

26.0

2.4

0.56 (4)

0.10

Las Animas

56.5

26.4

0.9

0.51 (5)

0.03

Eads

55.5

24.0

0.12 (1)

-0.37

Lamar

55.9

26.3

0.7

0.03 (1)

-0.53

Springfield

57.7

28.2

---

1.47 (3)

Rocky Ford

54.8

15.9

2.9

0 (0)

-0.32

Las Animas

54.4

16.7

2.1

0 (0)

-0.35

Eads

52.0

16.8

T (0)

-0.30

Lamar

51.6

18.1

0 (0)

-0.34

Springfield

53.1

19.8

Rocky Ford

49.7

12.5

1.1

0.17 (3)

-0.26

Las Animas

49.3

13.6

1.0

0.64 (3)

0.16

Eads

47.8

14.7

0.20 (1)

-0.19

Lamar

46.6

15.5

0.29 (2)

-0.21

Springfield

48.9

18.5

Location

November

December

2.1

T (0)

January

1.2

0.20 (1)

1./ Number of days of measurable precipitation in parentheses.

�-136-

Table 3.
1970-71.

Results of Canada goose aerial surveys, southeast Colorado,

Date
December 11 December 23

Area

November 23

Meredith Reservoir

9,700

9,350

10,720

Holbrook Reservoir

o

o

150

o

Horsecreek Reservoir

500

o

o

o

o

o

9

o

Mud Lake

o

1,400

o

25

Swede Lake

o

3,000

o

o

Sweetwater Res.

220

o

120

300

Flamingo Lake

7,300

9,000

1,700

o

Nee Noshe Res.

5,300

11,600

3,800

600

330

400

5,700

1,500

o

o

100

o

Blue Lake

January 19
4,695 ]../

Eads Group:

Upper Queens Res.
Thurston Reservoir
Two Buttes Reservoir

29,000

24,600 1/

20,400

32,200

Turk's Pond

12,000

21,200 ])

20,000

13,400

John Martin Reservoir

23,700

10,000

14,750

9,000 1/

Verhoeff Pond

350

o

1,100

o

Bonny Reservoir

1,000 1/

o

1,700

1,000 1/

90,550

80,249

62,720

Total

1/ Estimate from ground.

89,400

�-137-

Table 4. Results of Short Grass Prairie Canada goose coordinated
inventories, hunting season, 1968, 1969, and 1970.

State

1968

December
1969

1970

1969

January
1970

1971

Nebraska

11,740

10,512

8,000

7,000

8,140

8,000

Colorado

28,633

60,700

90,550

29,201

63,444

63,720

Oklahoma

3,000

1,300

4,000

3,500

1,500

3,000

New Mexico

9,195

7,531

13,842

7,848

10,640

4,654

,

Texas

20,100

26,387

33,800

64,850

63,690

65,975

Totals

72,668

106,930

150,192

112,399

147,414

145,399

The distribution of geese within southeast Colorado exhibited some distinct
changes from the 1969-70 season.
Meredith Reservoir held about 10 percent
of the geese through most of the season.
This is in contrast to the normal
early season build-up in the Meredith area, followed by a decline to between
1,000 - 2,000 birds by the end of the season (Rutherford 1970, Szymczak 1970).
Geese did not use Horsecreek Reservoir or Blue Lake during the early portion
of the hunting season as they did during the 1969-70 season.
During the 1969-70 season, an average of about 35 percent of southeast
Colorado geese used the Eads Lakes area as compared to only 14 percent
during the 1970-71 season.
A substantial decline in use of the Eads area
was noted after the switch to full day hunting (Table 3). The increased
hunting pressure along with the lack of open bars for resting, as discussed
previously, may have resulted in reduced use of that area.
For the second year in succession Two Buttes Reservoir maintained large numbers
of geese throughout most of the hunting season (Table 3). There continues
to be an inter-change of geese between Two Buttes and Turk's Pond.
For the
past two years the geese have not abandoned Two Buttes completely and moved
to Turk's for long periods of time as they had in previous years.
Ideal
resting situations and the removal of one of the two rows of pits at Two
Buttes along with the half-day hunting regulations have all had some influence
on this behavioral change.
At the same time, hunting pressure at Turk's has
become more intense as hunting pits on the south side of the reservoir have
been moved closer to the water.
The status, in terms of numbers, of the post season populations in Colorado
and throughout the entire short grass prairie wintering range is presented

�-138-

in Tables 5 and 6, respectively. Colorado's 1970-71 post-season population was about 63,000; essentially the same as the 1970 totals and far
above population levels recorded for any year previous to 1970. Colorado
has definitely retained more birds in the state since the initiation of
half-day hunting. The total short grass prairie population numbers in
1970-71 remained quite similar to those recorded in 1969-70. This is
unusual in view of the apparent excellent production experienced by this
population during the spring of 1970. Reduced numbers are definitely,
in part, the result of diffusion of short grass prairie geese into other
wintering populations in Colorado, New Mexico and possibly Oklahoma.

Table 5. January inventory of Canada geese, Arkansas Valley, Colorado,
1948-1971.
Year

Goose Count

Year

Goose Count

Year

Goose Count

1948

4,798

1956

24,212

1964

33,750

1949

12,286

1957

24,617

1965

37,693

1950

13,170

1958

35,894

1966

38,635

1951

19,320

1959

44,660

1967

29,835

1952

30,463

1960

37,394

1968

42,682

1953

20,236

1961

31,360

1969

29,201

1954

20,280

1962

40,250 1/

1970

63,444

1955

25,110

1963

35,889

1971

62,720

]j

1/ Inventory of February 7, 1962 substituted for January, 1962 inventory.

1/ Inventory of February 15, 1966 substituted for January, 1966 inventory.

Table 6. Post-hunting season status of Short Grass Prairie Canada goose
population, 1960-71, data generally from regular mid-winter inventories.
Year

Number of Birds

Year

Number of Birds

1960

77,709

1966

110,4851/

1961

103,355

1967

111,452

1962
1963
1964
1965

80,133
1968
127,903
93,940
1969
112,399
81,221
1970
147,414
103,435
1971
145,349
1/ Inventory of Feb. 15, 1966, substituted for unsatisfactory January
inventory.

�-139-

Analysis

of Flight Pattern Data

The characteristics of feeding flights were recorded throughout the 197071 season in an effort to determine the response of the birds, if any,
to variations in hunting pressure dictated by hunting regulations.
From
1966 through 1969, the earliest opening date was November 16 and the
latest November 22. Geese normally arrive in large numbers in southeast
Colorado around November 10. In 1970, the season opened to morning hunting
only on October 31 and, as previously mentioned, there were large numbers
of birds already in southeast Colorado.
The early opening, prior to the normal arrival of geese, resulted in reduced hunting pressure compared to normal opening dates. There was a
gradual increase in hunting pressure as the season progressed through
November.
The progressive increase in hunting pressure may have solicited
a different goose response in terms of feeding flights than would a normal
opening with immediate heavy hunting pressure (Szymczak 1970).
Throughout
the entire month of November, geese flew to feed twice daily, morning and
afternoon, in all major southeast Colorado goose concentration areas.
The
light of the moon period preceding and following the full moon on November
13, had very little noticeable effect on feeding flights.
In previous
years during full moon phases morning flights were retarded for a period
of up to 10 days. It is possible that inclement weather on November 13
and 14, during the peak of the full moon, stimulated feeding flights on
those two days, whereas with good weather conditions the birds may not
have flown twice a day. In 1969, geese, after experiencing two-a-day
feeding flights during the pre-season period. reverted exclusively to
late afternoon-evening
flights following opening day which occurred one
day prior to a full moon (Szymczak 1970).
The geese responded somewhat differently to the December 12 full moon
period.
There was a definite decline in the size of the morning flight on
December 11. On December 12, the day in which full day hunting began, only
a small number of geese fed in the morning.
Morning flights did not occur
again until December 18 in the Ead's Lakes area and December 20 at Two Buttes.
Afternoon flights did continue during this period in spite of moon and
weather conditions being conducive to exclusive night feeding.
Afternoon
flights on December 12 and 13 were somewhat reduced at Two Buttes, as many
flocks returned to the reservoir after being fir~d upon.
Morning flights from late December through the remainder of the season were
much smaller in comparison with morning flights prior to December 12. Portions
of late December and early January were characterized by overnight temperatures
of 100 F or less. As reported below, these relatively cold temperatures have
a definite effect on retarding goose movements during morning hours.
In general large afternoon feeding flights continued daily from mid-December
throughout the remainder of the season.
The light of the moon period
occurring around January 4 had no discernable effect on retardation of daytime goose feeding flights.
Afternoon flights during cold periods, when no
morning flight occurred, would normally begin early in the afternoon.
On
some days with normal temperatures, birds would leave reservoirs in late
afternoon and not return until after dark.

�-140-

Effects

of Cold Temperature

on Canada

Goose Feeding Flights

During the two-year study of feeding flight patterns it was consistently
noted that daily minimum temperatures affected the occurrence and timing
of Canada goose feeding flights.
Information collected daily at Two
Buttes Reservoir revealed that if the minimum daily temperature recorded
near Springfield, Colorado, 15 miles south of Two Buttes, was 100 F or
less, geese would not, for the most part, undergo feeding flights until
early afternoon.
This occurrence was also recorded in other areas in
southeast Colorado but daily feeding flight pattern information was not
consistently recorded in other areas.
Table 7 shows Springfield temperatures, goose flight times and moon phases
for a IS-day period during 1969-70 and a 26-day period during 1970-71.
Morning-only hunting was in effect during the 1969-70 period and all-day
hunting during the 1970-71 period.
Afternoon flight times are incomplete
during the 1969-70 period.

Age Composition
Tail fans were collected throughout the hunting season at the Two Buttes
and Lamar-Eads Management Areas and in the Meredith Lake area. The age
composition of the harvest on the two management areas indicate an increase
over the 1969-70 season in the percent of immatures in the harvest (Table
8). Nearly 60 percent of birds bagged at Two Buttes were classified as
immatures - the highest figure recorded since the 1959-60 season.
A decline
in percent immatures from the 1969-70 level was recorded in the Meredith
Lake area. A sample of 87 tail feathers collected at Turk's Pond, a firing
line area, had only 40.2 percent immatures or a 1:067 adult to immature
ratio. This sample, however was collected in late December and early
January and thus compares favorably to age data collected at Two Buttes
during the same general period (Table 9).
During the 1969-70 season, age ratios of birds bagged at the Lamar-Eads
and Two Buttes Management Areas exhibited some unusual trends as the season
progressed.
Immatures during that morning-only hunting season seemed to
increase in vulnerability during the first half of the season and then
decreased after mid-season (Szymczak 1970). This trend is contrary to a
general decrease in vulnerability as the season progresses which was described by Rutherford (1968) as characteristic of the southeast Colorado
flock. These unusual age ratio trends were attributed to the half-day
hunting season.
The large sample of tail fans collected at Two Buttes
during the half-day portion of the 1970-71 season provided adequate information to examine age ratio trends.
Although not conclusive, the data
presented in Table 9 indicates a general decreasing immature vulnerability
through the half day season at Two Buttes.
Lamar-Eads data show a definite
decreasing trend.
However, the validity of the Lamar-Eads results are
questionable because of the small sample size. After the switch to full day
hunting, the Two Buttes vulnerability data shows .a normal trend. Consistent
with previous years findings, immatures were taken at a greater rate in
the decoy area than on the firing line.

�-141-

Table 7. Goose feeding flight times at Two Buttes Reservoir in comparison
with Springfield, Colorado temperatures for selected periods during the
1969-70 and 1970-71 goose seasons.

Date

Daily TemEerature
Minimum
Maximum

Time of Morning
Feeding Flights

1969-70
December 28
29
30
31
January
1
2
3
4
5
6
7
8
9
10
11

17
12
5
7
9
7
7
8
5
-4
1
-2
1
28
25

26
24
28
34
30
31
37
45
23
22
25
20
46
57
48

9:00 AM
9:30 AM
9: 30 AM (Snow) 1/
No Flight (3:00 AM)
No Flight
No Flight
No Flight
No Flight
No Flight (12:30 PM)
No Flight
No Flight (3:00 PM)
No Flight (2:30 PM)
No Flight
10:00 AM (Small Flight)
7:45 AM

1970-71
December 17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
January
1
2
3
4
5
6
7
8
9
10
11

26
20
7
23
18
20
5
15
3
20
28
19
15
20
16
23
17
10
0
-12
-9
-10
0
16
25
24

67
49
39
54
48
54
37
54
41
43
56
52
53
42
59
46
34
20
10
13
12
25
43
52
59
47

No Time (Small Flight)
7:35 AM
No Flight (2:40 PM)
8:00 AM
8:16 AM
7:15 AM
No Flight (2:00 PM)
No Flight (3:00 PM)
No Flight
No Flight (3:15 PM)
7:45 AM
8:20 AM
7:00 AM
11:00 AM
No Flight (3:15 PM)
8:15 AM
No Flight
No Flight (1:45 PM)
11:50 AM
No Flight (12:10 PM)
No Flight
No Flight (12:45 PM)
No Flight (1:30 PM)
8:35 AM
Small Flight

Moon Phase

Last Quarter

New Moon

Last Quarter

New Moon

First Quarter

Full Moon

1/ Time of afternoon flight, if known, or pertinent comments in parentheses.

�Table 8.

Age composition of the goose harvest in selected areas, southeast Colorado.

Year

Two Buttes (Firing Line)
Iunnature
Adult
No.
Percent
No.
Percent

1951-52

356

1957-58

46.8

404

53.2

Lamar-Eads (Decoy)
Adult
Immature
No.
Percent
No.
Percent

Meredith Area
Immature
Adult
Percent
Percent
No.
No.

No Data

1958-59

929

47.9

1,010

52.1

1959-60

377

36.5

655

63.5

1960-61

612

45.5

732

54.5

1961-62

527

55.7

419

44.3

I

.j::''"""
N
I

1962-63

204

48.9

213

51.1

1963-64

377

52.1

346

47.9

1964-65

442

63~1

259

36.9

1965-66

571

62.7

340

37.3

1966-67

217

45.3

262

54.7

1967-68

245

59.9

164

40.1

1968-69

136

61. 8

84

38.2

79

51. 3

75

48.7

48

66.7

24

33.3

1969-70

110

55.3

89

44.7

143

35.9

255

64.1

18

40.0

27

60.0

1970-71

630

40.6

922

59.4

77

27.4

204

72.6

42

45.7

50

54.3

�-143-

Table 9. Weekly age ratios of Canada geese brought through check stations,
southeast Colorado during the 1970-71 season.

Date

Lamar-Eads (Decoy)
Ratio
Adult: Immature
Number

Two Buttes (Firing Line)
Ratio
Adult: Immature
Number

10/31-11/6

1:5.64

73

1: 1. 82

93

11/7-11/13

1:8.80

49

1:2.88

66

11/14-11/20

1:0.86

13

1:1. 75

77

11/21-11/27

1:1.17

13

1:1.93

281

11/28-12/4

1:0.17

7

1:2.25

179

5

1:1. 39

146

12/5-12/11
Total (Half Day)

1:3.57

160

1: 1. 90

842

12/12-12/18

1:2.86

27

1:1.27

209

12/19-12/25

1: 1. 42

29

1:1. 95

162

12/26-1/1

1:2.13

25

1:0.91

204

1/2-1/8

1: 1. 25

9

1:0.42

17

1/9-1/15

1: 1. 63

21

1:0.51

77

1/16-1/17

1:2.33

10

1:0.75

41

Total (Full Day)

1: 1. 88

121

1: 1. 09

710

Season Total

1:2.65

281

1: 1. 46

1,522

�-144-

An age-specific feeding flight was hypothesized as a possible reason for
unusual age-ratio trends during the half-day season of 1969-70. It was
thought that morning flights may have been composed of immature birds
and family groups with afternoon flights made up of sub-adults. During
the 1970-71 season, the time of harvest was recorded for each bird taken
on the two state-controlled.management areas. Table 10 presents the age
composition of the harvest by time period. No trends or period preferences
are evident during any part of the season. In fact, age ratios for total
morning harvest and total afternoon harvest during the full day hunting
periods are essentially the same at Two Buttes. Unfortunately the 1970-71
full day season period occurred late in the year and did not coincide with
the period in which the unusual age ratios were evident the previous season.

Table 10. Age composition of the harvest by time period
on two state controlled management areas in southeast Colorado, 1970-71.

Hourly Period

Lamar-Eads
Half Day
Full Day

Two Buttes
Half· Day
Full Day

6-7

1:4.67 (34)

(0)

1:3.21 (80)

(1)

7-8

1:6.25 (58)

(1)

1:1.95 (327)

1:3.00 (12)

8-9

1:1. 77 (36)

1:1.50 (5)

1:1.80 (258)

1:1.21 (73)

9-10

1:3.80 (24)

1:6.00 (7)

1:1.38 (114)

1:1.25 (72)

10-11

1:1.33 (7)

1:0.67 (10)

1:1.91 (32)

1:0.94 (64)

11-12

(1)

1:3.00 (4)

1:2.44 (31)

1:0.50 (21)

1:1.70 (27)

1:1.90 (842)

1:1.11 (243)

Total (AM)

1:3.57 (160)

12-1

1:2.00 (3)

1-2

(0)

1:1.11 (19)

2-3

1:1.00 (4)

1:1. 37 (64)

3-4

1:2.00 (33)

1:1.25 (189)

4-5

1:2.00 (54)

1:0.88 (186)

Total (PM)

1:1.94 (94)

1:1.08 (467)

1:0.5

(9)

�-145-

Birds were trapped and banded at Two Buttes Reservoir and at Verhoeff's
Pond near John Martin Reservoir.
The sex and age composition of birds
captured in the cannon-net operation is presented in Table 11. Overall,
about 36 percent of the birds captured were immatures.
Age composition
of the total catches at both areas were essentially the same with 35.4
percent immatures at Two Buttes compared to 36.6 percent at Verhoeff's.

Table 11. Sex and age composition
southeast Colorado, 1971.
Location
and
Date

of Canada geese trapped post-season

Males
Adult
Immature

Females
Adult
Immature

in

Total

Percent
Immatures

Two Buttes 1/24/71

7

1

11

3

22

18.2

Two Buttes 1/25/71

34

14

38

28

114

36.8

Two Buttes 1/26/71

10

3

24

6

43

20.9

Two Buttes 1/27/71

77

31

64

47

219

35.6

Two Buttes 1/28/71

38

29

42

30

139

42.4

Verhoeff

2/1/71

54

15

44

14

127

22.8

Two Buttes 2/2/71

1

0

2

0

3

0.0

Verhoeff

2/8/71

38

50

56

29

173

45.7

Verhoeff

2/13/71

5

1

4

6

16

43.8

264

144

285

163

856

35.9

Total

A summary of all age data collected in southeast Colorado is presented in Table
12. In total, immaturesmade
up approximately 61 percent of the check station
sample and 36 percent of the post-season trapped sample.
It is interesting to
note that the percent of the harvest composed of immatures taken at Two Buttes
during January was 35.9 percent and thus compares favorably to the 35.4 percent immatures in the Two Buttes banded sample trapped directly after the
season.
In total, the available age ratio data indicate that 1970 was an excellent
production year for short grass prairie Canada geese.

�-146-

Table 12. Age composition of southeast Colorado Canada geese, 1970-71,
as estimated by check station and trapping results.
No. of
Adults

Percent
Adults

No. of
Young

Percent
Young

Total
Birds

Lamar+Eads

77

27.4

204

72.6

281

Two Buttes

630

40.6

922

59.4

1,552

Meredith Lake

42

45.7

50

54.3

92

Total

749

38.9

1,176

61.1

1,925

Check Station Sample

Adult/Immature

1:1.57

Trapped Sample
Two Buttes

350

64.6

192

35.4

542

Verhoeff's

199

63.4

115

36.6

314

Total

549

64.1

307

35.9

856

Adult/Immature

1:0.56

Hunting Pressure and Harvest
A record number of Canada geese were harvested in southeast Colorado during
the 1970-71 season. An estimated 30,329 geese were bagged in southeast
Colorado (Table 13, Funk, In Press). An additional estimated 5,459 birds
were lost because of crippling making the total flock hunting lose 35,788
birds. The high harvest and increased average season bag (Table 13) was
the direct result of goose availability. Large numbers of birds were in
southeast Colorado throughout the entire hunting season as was discussed
earlier. After a one-year absence, Baca County regained its prominence as
the leader in goose harvest in southeast Colorado. The harvest in all
counties considered major goose areas exceeded the 16 year average (Table
14).

�-147-

Table 13.

Goose hunting season statistics, 1954-70.
Arkansas Valley
Estimated
Average
Goose Hunters
Season Bag

Year

Dates of Season

Stamp
Sales

1954

11/1 - 12/30

32,450

7,071

1.04

7,372

1955

11/1 - 12/30

39,107

9,054

1.54

13,904

1956

11/9 - 1/7

36,303

9,833

1.05

10,276

1957

11/2 - 11/31

41,794

9,113

1.39

12,656

1958

11/17 - 1/15

41,897

10,082

1.51

15,205

1959

10/26 - 1/8

31,431

8,888

1.61

14,309

1960

10/26 - 1/8

30,592

9,838

1.39

13,629

1961

11/10 - 1/8

24,854

7,577

1.68

11,724

1962

10/31 - 1/13

17,701

6,021

1.58

9,495

1963

11/2 - 1/15

22,940

6,668

2.17

14,444

1964

11/2 - 1/15

25,282

8,016

2.30

18,474

1965

11/2 - 1/15

20,537

6,313

1.52

9,613

1966

11/19 - 1/15

29,377

9,357

2.59

24,269

1967

11/18 - 1/14

31,064

6,975

2.23

15,558

1968

11/16 - 1/15

31,218

6,668

1.66

11,046

1969 1/

11/22 - 1/15

34,281

5,327

1.99

10,597

10/31 - 1/17

37,972

7,875

3.85

30,329

1970

Y

1/ Morning hunting only throughout the season.
~I Morning hunting only through December 11; full day hunting for the
remainder of the season.

Estimated
Kill

�-148-

Table 14. Comparison of the 1970-71 southeast Colorado goose harvest,
by county, with the l6-year average, 1954-69, based on the results of the
random small game survey.

County

Lakes and Reservoirs

Number and Percent of Geese Bagged
1970-71
16 Yr. Average
No.
Percent
No.
Percent

Baca

Two Buttes and Turk's

13,055

43.0

5,083

37.1

Kiowa

Eads and Blue

7,005

23.1

3,098

22.6

Prowers

Two Buttes and Eads

3,980

13.1

2,125

15.5

Crowley

Meredith

3,025

10.0

716

5.2

Bent

John Martin, Blue
and Horsecreek

2,229

7.3

1,913

14.0

Horsecreek, Cheraw,
Dye and Holbrook

557

1.8

359

2.6

Pueblo

358

1.2

183

1.3

Las Animas

120

0.4

120

0.9

Huerfano

o

0.0

116

0.8

Otero

Total

Two Buttes and Lamar-Eads

30,329

Management

13,710

Areas

Hunting activity and harvest at Two Buttes Management area increased dramatically in 1970-71 over recent years (Table 15). The number of hunters utilizing
the Management Area was the highest since the 1965-66 season and the size of
the harvest had been exceeded only by that recorded during the 1958-59 seasons.
Hunting success, by Two Buttes standards, was phenomenal with participants
averaging better than 0.5 birds per day and approximately 1.2 birds per
season.
Hunters during the.half-day season had better success, on the average,
than those hunting during the full day period (Table 15).
Hunting pressure at Two Buttes was well distributed throughout the season
(Table 16). Hunting pressure was heaviest during the last full week in
November, the first full week following the change to full day hunting, and
the last full week in December.
Hunting success was greatest during the first
week of the season and from November 21, through December 4. Hunting success
for any given weekly period during the half day season exceeded success for
all weekly periods during the full day season except the December 19 to
December 25 interval (Table 16).

�-149-

Table 15. Goose harvest, hunting pressure, and hunter success, Two
Buttes Management Area.
No. of
Individual
Hunters

No. of
Hunter
Days

No. of
Geese
Bagged

Ave. Bag
per
Hunter

Ave. Bag
per
Hunter Day

1961-62

2,392

4,758

945

0.40

0.20

1962-63

1,479

3,178

418

0.28

0.13

1963-64

1,750

3,659

728

0.42

0.20

1964-65

1,996

3,946

721

0.36

0.19

1965-66

2,596

5,264

941

0.36

0.18

1966-67

1,257

2,413

493

0.39

0.20

1967-68

840

1,554

433

0.52

0.28

1968-69

792

1,309

243

0.31

0.19

1969-70 1/

659

1,182

248

0.38

0.21

Half Day

610

1,384

919

1.51

0.66

Full Day

788

1,786

814

1.03

0.46

1,398

3,170

1,733

1.24

0.55

Year

1970-71

Total

1/ Morning only hunting throughout the season.

Hunter activity on the Lamar-Eads Management area was similar to the 1969-70
season, but hunting success declined (Table 17). Hunting success was high
during the first week of the season and declined thereafter (Table 18).
After the first week, the geese discontinued use of the management area
and began using fields to the north.

�-150-

Table 16. Hunter activity and goose harvest by weekly intervals, Two
Buttes Management Area, 1970-71.

Interval

Number of
Hunter bays

10/31 - 11/6

143 (4.5)

68

102

0.71

11/7 - 11/13

124 (3.9)

50

68

0.55

11/14 - 11/20

143 (4.5)

60

83

0.58

11/21 - 11/27

419 (13.2)

195

304

0.73

11/28 - 12/4

258 (8.1)

137

204

0.79

12/5 - 12/11

297 (9.4)

124

158

0.53

634

919

0.66

Half-day Total

1,384

Successful
Hunter Days

No. of
Geese Bagged

Ave.Bag Per
Hunter Day

12/12 - 12/18

464 (14.6)

162

243

0.52

12/19 - 12/25

296 (9.3)

127

189

0.64

12/26 - 1/1

575 (18.1)

158

220

0.38

1/2 - 1/8

158 (5.0)

20

25

0.16

1/9 - 1/15

171 (5.4)

62

82

0.48

1/16 - 1/17

122 (3.8)

39

55

0.45

Full-day Total

1,786

568

814

0.46

Season Total

3,170

1,202

1,733

0.55

�-151-

Table 17. Goose harvest, hunting pressure, and hunter success, LamarEads Management··Area.
No. of
Individual
Hunters

No. of
Hunter
Days

No. of
Geese
Bagged

1968-69

598

1,062

160

1969-70

703

1,361

512

0.73

0.38

1970-71

705

1,128

304

0.43

0.27

Year

Ave. Bag
per
Hunter

Ave. Bag
per
Hunter Day
0.15

Table 18. Hunter activity and goose harvest by weekly intervals, LamarEads Management Area, 1970-71.
Week
Interval

Number of
Hunter Days

10/31 - 11/6

127 (11.3)

11/7 - 11/13

Successful
Hunter Days

Number of
Geese Bagged

Ave. Bag per
Hunter Day

52

72 (23.7)

0.57

144 (12.8)

32

49 (16.1)

0.34

11/14 -11/20

64 (5.7)

11

14 (4.6)

0.22

11/21 - 11/27

101 (9.0)

12

14 (4.6)

0.14

11/28 - 12/3

70 (6.2)

6

6 (2.0)

0.09

12/4 - 12/11

49 (4.3)

4

7 (2.3)

0.14

12/12 - 12/18

105 (9.3)

26

34 (11.2)

0.32

12/19 - 12/25

99 (8.8)

25

32 (10.5)

0.32

12/26 - 1/1

193 (17.1)

23

25 (8.2)

0.13

1/2

- 1/8

68 (6.0)

10

11 (3.6)

0.16

1/9

- 1/15

65 (5.8)

18

29 (9.5)

0.44

43 (3.8)

8

11 (3.6)

0.26

304

0.27

1/16 Totals

1/17

1/

1,128

1./ Percent of total in parentheses.

227

�-152-

Banding Investigations
Trapping efforts post-season in southeast Colorado resulted in 836 Canada
geese being banded (Table 19). Three hundred birds were banded at
Verhoeff's Pond near Hasty, Colorado and 536 at Two Buttes Reservoir.
Again, as during the 1970 banding operations, about 47 percent of the birds
captured were males.

Table 19. Sex and age composition of Canada geese banded post-season in
southeast Colorado, 1971.
Location
and
Date

Males
Adult Immature

Females
Adult
Immature

Total

Two Buttes

1/24/71

7

1

11

3

22

Two Buttes

1/25/71

30

13

36

27

106

Two Buttes

1/26/71

10

3

23

6

42

Two Buttes

1/27/71

76

30

62

47

215

Two Buttes

1/28/71

38

29

41

30

138

Verhoeff

2/1/71

53

15

43

14

125

Two Buttes

2/2/71

1

0

2

0

3

Verhoeff

2/8/71

36

47

55

29

170 1/

Verhoeff

2/13/71

5

1

3

6

15

256

139

276

162

836

Total

]j Includes two females of unknown age and 1 goose of unknown age and sex.

Weights
The mean weights of all geese harvested on the two state-controlled management areas and trapped post-season are presented in Table 20. As in all
past years, except 1969-70, the mean weights of birds bagged at Two Buttes
exceeded mean weights of birds trapped there post-season.

�-153-

Table 20. Comparison of weights, in pounds, of geese from trap and check
station samples, southeast Colorado, 1970-71.
Adult Ave.
Wt.

Source

Range

Imm. Ave.
Wt.

Range

Total Ave.
Wt.

Range

Two Buttes
Check Station

5.47 (673)1/ 3.00-9.50

4.92 (949) 2.75-8.75

5.15 (1622) 2.75-9.50

Trapping

5.08 (332)

3.50-8.63

4.59 (192) 3.25-8.38

4.90 (524)

3.25-8.63

5.68 (8)

3.50-8.88

4.99 (2l0) 3.00-7.50

5.18 (288)

3.00-8.88

6.30 (215)

4.00-9.50

5.63 (128) 3.75-9.50

6.05 (343)

3.75-9.50

Lamar Mgmt.Area
Check Station

Verhoeff's

Pond

Trapping

1/ Sample size in parentheses.

Weights of birds bagged plotted by time period (Table 21) indicate that birds
taken during the full day hunting season, which began on December 12, averaged
approximately 0.2 lb. less than birds taken earlier during the half-day season.
A similar weight loss was recorded when comparing weights for the same time
periods for birds bagged during the 1969-70 season with a half-day regulation
in effect throughout the season (Szymczak 1970). However, during 1969-70,
birds trapped post-season weighed more than birds bagged, indicating the geese
may have maintained better body condition during the half-day season and
thus were able to increase in weight qUickly soon after the season (Szymczak
1970) •
Birds captured post-season at Verhoeff's Pond were, on the average, much
larger than those captured at Two Buttes (Table 20). The weights of birds
captured are presented by catch, and age and sex group in Table 22.

Distribution

of Harvest

Alberta, Saskatchewan and Colorado continue to be the major harvest areas
of short grass prairie geese (Table 23). The High Prairie-Peace River area
of Alberta, located north of 53 degrees latitude, is becoming increasingly
important as a harvest area. Therefore, recoveries above 53 degrees latitude
are now being delineated for analysis purposes.
In the provinces below 53
degrees, Saskatchewan reportedly harvested more banded birds than Alberta
for only the second year s~nce 1951-52 with the 1967-68 season being the
first year.

�-154-

Table 2l. Average weights of geese bagged by time interval on two state
management areas in southeastern Colorado during the 1970-71 hunting
season.

Lamar-Eads .;
Adults
Immatures

Two Buttes
Adults
Immatures

Oct. 31-Nov. 6

5.92 (1) 1.1

4.92 (67)

5.53 (36)

4.85 (57)

Nov. 7-Nov. 13

5.65 (5)

5.01 (43)

5.52 (18)

5.09 (48)

Nov. 14-Nov. 20

6.00 (7)

4.92 (6)

6.04 (29)

5.10 (54)

Nov. 21-Nov. 27

6.05 (7)

5.54 (7)

5.55 (98)

5.05 (193)

Nov. 28-Dec. 4

6.19 (6)

3.50 (1)

5.49 (60)

4.88 (l30)

5.l3 (6)

5.50 (64)

4.73 (87)

Intervals

Dec. 5-Dec. 11
Dec. 12-Dec. 18

5.56 (6)

5.33 (20)

5.54 (99)

4.82 (120)

Dec. 19-Dec. 25

5.60 (l3)

5.30 (18)

5.46 (59)

4.94 (113)

Dec. 26-Jan. 1

5.44 (8)

4.62 (17)

5.27 (114)

4.80 (99)

Jan. 2-Jan. 8

4.44 (4)

4.94 (4)

4.69 (l3)

4.65 (5)

Jan. 9-Jan. 15

5.47 (8)

4.55 (14)

5.40 (52)

5.14 (25)

Jan. 16-Jan. 17

5.75 (3)

5.12 (7)

5.54 (31)

5.04 (18)

1./ Sample size in parentheses.

The percent of banded birds reported recovered in Colorado declined to its
lowest annual level during the 1969-70 season. This decline was primarily
a reflection of the reduction in hunting pressure and harvest during the
1969-70 season (Szymczak 1970).
Only 105 bands were reported recovered throughout the 1969-70 season in
spite of the addition of over 1,000 banded birds to the population after
the 1968-69 season. The reduction of the number of bands reported being
recovered along with a very low first-year recovery rate may have been
a direct result of a major portion of the short grass population remaining
in Colorado and thus undergoing a lower harvest rate because of reduced
pressure.

�-155-

Table 22. Weights of Canada geese captured with cannon-nets in southeast
Colorado, 1971.

Date

Location

Grouping

Number

Ave. Weight

Range

January 24

Two Buttes

AM

7
1

AF
IF
Subtotals

11
22

5.88
5.25
4.96
3.96
5.13

5.00-7.00

1M

4.25-6.25
3.25-5.00
3.25-7.00

AF
IF
Subtotals

30
14
34
28
106

5.25
4.73
4.58
4.39
4.74

4.38-6.50
4.00-5.25
3.50-5.25
3.38-5.50
3.38-6.50

AM

10

1M

3

AF
IF
Subtotals

23
42

6.13
5.09
5.33
4.04
5.32

4.25-7.50
4.88-5.38
4.00-8.63
3.75-4.50
3.75-8.63

76
31
62
47
216

5.39
4.95
4.79
4.46
4.95

4.25-7.25
3.75-7.00
3.50-6.50
3.50-5.88
3.50-7.25

38
29
41
30
138

5.37
4.94
4.43
4.32
4.77

4.50-8.13
3.63-8.38
4.00-5.50
3.38-6.38
3.38-8.38

AF
IF
Subtotals

52
13
41
13
119

6.75
6.52
6.33
5.62
6.46

4.50-9.38
5.13-9.50
4.38-9.50
4.25-7.00
4.25-9.50

AM

10

1M

8

AF
IF
Subtotals

11
36

7.53
6.69
6.51
6.04
6.74

5.13-9.50
4.25-8.88
5.38-8.50
4.13-7.63
4.13-9.50

34
46
49
23
152

6.34
5.42
5.37
4.99
5.54

5.25-8.00
4.25-7.00
4.00-6.75
3.75-6.38
3.75-8.00

January 25

Two Buttes

AM
1M

January 26

January 27

Two Buttes

Two Buttes

AM
1M

AF
IF
Subtotals
January 28

Two Buttes

AM
1M

AF
IF
Subtotals
February 1

Verhoeff Pond

AM
1M

February 3

February 8

Verhoeff Pond

Verhoeff Pond

AM
1M

AF
IF
Subtotals

3

6

7

-----------------------------------------------------------------------

�Percentages of total band recoveries, Arkansas Valley Eost-season bandings, by area and year of recovery, all banding years combined.
Percentage
Recovery Year
of Total
Total No.
Five Year Averages
Recoveries
Recoveries
1951-1955 1956-1960 1961-62 1962-63 1963-64 1964-65 1965-66 1966-67 1967-68 1968-69 1969-70
Area
Far North Above 530
1.0
2.1
---N.W. Territories
-----7.1
12.4
---Alberta
-----0.7
---Saskatchewan
------9.4
7.5
9.9
13.4
254
15.2
10.4
15.5
4.7
9.8
7.1
7.9
6.3
Total
Table 23.

--

--

Provinces Below 530
Alberta
Saskatchewan
B. C., Manitoba
Ontario
Central Fl.yway
Montana
North Dakota
Hyoming
South Dakota
Nebraska
Colorado
Southeast and Other
North-central
Total

28.6
18.3

35.3
10.5

34.9
9.3

33.3
19.4

0.4

6.5

5.4

3.1

---

---

---

25.5

30.9

31. 5

0.4
0.9
1.5

0.3
0.3
0.7

0.8

2.3
2.0
0.8
5.1

2.8
2.2
0.2
5.2

5.4

Pacific Flyway

2.4

2.2

Mississippi

0.1

Flyway

26.8
8.5

27.9
10.9

0.1
0.8
0.1
0.7
0.1
5.4

Kansas
Oklahoma
New Mexico
Texas
Panhandle
\.JaggonerRanch
Gulf Coas t
Total

28.4
17.4

Mexico

0.1

Total Number of
Recoveries

748

0.7

1.5

0.7

0.7

0.7

5.5

6.7

2.0
1.4
8.8

0.7
0.7
9.7

--

--

---

---

30.9

21.7

--30.1

35.4

---

32.4

0.7
1.4

0.7

0.5

1.0
0.5
1.0

0.6

5.4

4.6
0.8
0.8
6.2

4.0
2.0
1.0
6.0

7.3
2.4
0.6
10.3

4.8

0.8

1.5

1.2

129

14.2
24.6

0.6

0.6

677

19.4
16.4

129

201

164

1.4
0.7
0.7

21. 9
25.7

1.9

9.0

4.3

1.9
4.8

27.6
1.5
29.1

29.1
2.8
31. 9

19.0
1.0
20.0
1.0

2.7
2.0

7.5
0.7

4.7

8.2

147

23.4
17.0

134

785
416

29.0
15.4

4

0.1

17
2
15
6
166

0.6

773

28.5
0.4
0.6
0.8

T

0.6
0.2
6.1

1.4

1.0

10
15
23

2.2
9.7

5.0
0.7
1.4
7.1

6.7
1.0
1.0
8.7

107
45
17
169

3.9
1.7
0.6
6.2

0.7

1.4

50

1.8

0.7

0.7

3

0.1

1

T

1.5
0.7
7.5

134

141

105

2709

I
•.....
Ln

C1:&gt;
I

�Table 24. Composite dynamic analysis of band recoveries from Canada geese banded as juveniles, Arkansas Valle" , Colorado, 1951-1969.

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1&lt;160-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69

300
628
578
0
0
0
0
176
298
167
248
123
399
251
198
0
285
270
184

Recoveries by Year Following Bandin~
11
12
10
8
9

13

14

15

16

17

18

19

0
5
1

0
2
3

1
1

1
0
1

0
0
0

0
1
2

1
0

0

X
X
X
X

X
X
X
X

X
X
X
X

X
X

X

0
1

0

1

2

3

4

5

6

7

31
58
52

25
31
25

11
33
25

11
20
24

4
16
20

5
8
11

4
7
11

7
6
9

1
4
2

4
4

0
0
4

X
X
X
X

X
X
X
X

X
X
X
X

X

X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

21
25
16
15
10
40
16
15

16
16
10
5
12
24
11
7

X
X
X
X
6

16
3
4
10
9
11
8

4
8
14
7
0
14
13
1

4
7
8
8
1
6
5
0

1
3
0
7
1

4
0
0
3
1
2

0
1
1
2
1

3
1
0
1

1
2
0

X

2
2

1

1
X
X
X

I
•....
VI

'"
I

X

X

X

18
16
9

10
9

5

342

201

141

116

79

40

32

27

12

12

5

6

5

3

2

0

3

1

0

Banded Birds
Eligible

4105

3921

3651

3366

3366

3168

2917

2518

2395

2147

1980

1682

1506

1506

1506

1506

1506

928

300

Recoveries per
1,000 Banded

83.3

51.3

38.6

34.5

23.5

12.6

11.0

10.7
207.5

5.0

5.6

2.5

3.6

3.3

2.0

0.0
291.9

2.0

1.11

0.0

37.1
818.3
.254

26.4

3.1
- 1111.3
.263

3.1

1.1 (

0.0

--Total
Recoveries

4105

Alive Going
Into Period
Mortality Rate

291.9 ,208.6
.285

157.3

118.7

84.2

60.7

48.1

1.3
£.

21.4

15.8

13.3

9.7

6.4

=

4.4
~.

�Table 25. Composite dynamic analysis of band recoveries from Canada geese banded as adults, Arkansas Valley, Colorado, 1951-1969.

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963 -64
1964-65
1965-66
1966-67
1967-68
1968-69

Total
Recoveries

Recoveries b~ Year Following Banding
12
11
10
8
9

13

14

15

16

17

18

19

2
7
2

0
4
4

1
2
2

1
0
0

0
1
1

1
0
1

0
0
0

0
0

0

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X

X
X

X

2
1
1

0
0

0

1

2

3

4

5

6

7

344
650
900
0
0
0
0
347
217
250
306
334
369
335
780
0
389
646
905

45
71
85

23
37
55

19
24
50

10
23
31

7
21
25

12
6
20

5
11
9

3
6
12

3
9
10

2
7
6

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

33
21
27
29
26
35
24
55

27
9
16
13
18
20
12
40

17
7
6
18
12
14
15
20

8
4
12
12
5
4
10
22

5
4
8
4
13
11
4
12

8
2
4
7
3
5
4

2
1
6
3
2
2

2
1
3
2
2

2
0
2
3

X

X

X

X

29
35
22

26
17

9

6772

537

313

211

141

114

71

41

31

29

19

11

8

5

1

2

Banded Birds
Eligible

6772

5867

5221

4832

4832

4052

3717

3348

3014

2708

2458

2241

1894

1894

1894

Recoveries per
1,000 Banded

79.3

,53.3

40.4

29.2

23.6

17.5

11.0

9.3
214.3

9.6

7.0

4.5

3.6

2.6

0.5

1.1

Alive Going
Into Period

293.6 '\214.3..1(jl.0_120~

39.3
828.7

30.0

Mortality Rate

.270

91.4

67.8

I
•....
Q'I

0
I

50.3

.259

0

0

0

1894

944

344

1.1

0.0

0.0 I

0.0

1.1

0.0

0.01

0.0

2

1894

~- - 293.6
20.4

13.4

8.9

5.3

2.7

2.2

~- = 1122.3

.262

�Table 26. Composite dynamic analysis of band recoveries from Canada geese banded in all age classes, Arkansas Valley, Colorado, 1951-1969.

1

2

3

4

5

6

7

Recoveries by Year Followin~ Bandin~
11
12
10
8
9

644
1278
1478
0
0
0
0
523
515
417
554
457
768
586
978
0
674
922
1089

76
129
137

48
68
80

30
57
75

21
43
55

11
37
45

17
14
31

9
18
20

10
12
21

4
13
12

3
11
10

2
7
6

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

54
46
43
44
36
75
40
70

43
25
26
18
30
44
23
47

23
23
9
22
22
23
26
28

12
12
26
19
5
18
23
23

9
11
16
12
14
17
9
12

9
5
4
14
4
7
6

6
1
6
6
3
4

2
2
4
4
3

5
1
2
4

3
3
1

X

X

X

X

47
51
31

36
26

14

10883

879

514

352

257

193

III

73

58

41

31

16

14

10

4

4

2

3

1

0

Banded Birds
Eligible

10883

9794

8872

8198

8198

7220

6634

5866

5409

4858

4438

3923

3400

3400

3400

3400

3400

1922

644

Recoveries per
1,000 Banded

80.8

,52.5

39.7

31.3

23.5

15.4

11.0

9.9
211.1

7.6

6.4

3.6

3.6

2.9

1.2

1.2
- 292.5

0-"-6

0.8

0.5,

0.0

Alive Going
Into Period

292.5 ,211.7

159.2

119.5

88.2

64.7

49.3

38.3
823.6

28.4

20.8

14.4

10.8

7.2

4.3

3.1

1.9

1.3

0.51

0.0

Year
Banded

Number
Banded

1950-51
1951-52
1952 -53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69

Total
Recoveries

Mortality

Rate

.276

13

14

15

16

17

18

19

0
9
5

1
4
5

2
1
1

1
1
2

1
0
1

0
1
2

1
0

0

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X

X
X

X

0
1

0

....
....
I

(J\

I

.256

~

z .. = 1116.1

.262

�Table 27. Relative recovery rate analysis of band recoveries from Canada geese banded as juveniles, Arkansas
Valley, Colorado, 1951-1969.

Winter
Banded

Number
Banded

1950-51

300

cu 1/

72

--

1951-52

628

196

138

.3121

1952-53

578

195

CU

.3374

1957-58

176

CU

39

1958-59

298

80

1959-60

167

52

Number Recoveries
1 - n
2 - n

Survival
Rate

Mortality
Rate

.2400

.769

.231

.2197

.651

.349

--

.2216

.825

.175

55

.2685

.1846

.593

.407

36

.3114

.2156

1.028

--

Recover:l Rates
2 - n
1 - n

1960-61

248

52

37

.2097

.1492

.510

.490

1961-62

123

36

26

.2927

.2114

.870

.130

1962 -63

399

97

57

.2431

.1429

.618

.346

1963-64

251

58

42

.2311

.1673

1.068

1964-65

198

31

cu

.1566

1966-67

285

CU

15

--

.0526

.568

.432

1967-68

270

25

9

.0926

.0333

.681

.319

1968-69

184

9

--

.0489
--

4,105

1.1 Cannot use.

2.5041

1.8382

Average Survival rate

.734

Average Mortality

.266

rate

I

I-'

0'\
N
I

�Table 28. Relative recovery rate analysis of band recoveries from Canada geese banded as adults, Arkansas
Valley, Colorado, 1951-1969.
Winter
Banded

Number
Banded

1950-51

344

CD 1/

89

--

1951-52

650

229

158

.3523

1952-53

900

313

CD

.3478

1957-58

347

CD

73

1958-59

217

50

1959-60

250

85

1960-61

306

91

62

.2974

.2026

.835

.185

1961-62

334

81

55

.2425

.1647

.668

.342

1962-63

369

91

56

.2466

.1518

.737

.246

1963 -64

335

69

45

.2060

.1343

.703

.303

1964-65

780

149

CD

.1910

1966.,67

389

CD

35

--

.0900

1.118

1967-68

646

52

17

.0805

.0263

1.082

1968-69

905

22

--

.0243

Number Recoveries
1 - n
2 - n

Recovery Rates
1 - n
2 - n

Survival
Rate

Mortality
Rate

.2587

.734

.266

.2431

.699

.301

--

.2104

.913

.087

29

.2304

.1336

.393

•602~

58

.3400

.2320

.780

.207

I

t-'

0\
W

6,772

1/ Cannot use.

2.5588

1.8475

Average Survival Rate

.722

Average Mortality Rate

.278

I

�Table 29. Relative recovery rate analysis of band recoveries from Canada geese banded in all age classes,
Arkansas Valley, Colorado, 1951-1969.

Winter
Banded

Number
Banded

1950-51

644

1951-52

Number Recoveries
1 - n
2 - n

Recovery Rates
2 - n
1 - n

Survival
Rate

Mortality
Rate

CD 1/

161

--

.2500

.752

.248

1,278

425

296

.3326

.2316

.674

.326

1952-53

1,478

508

CD

.3437

1957-58

523

CD

112

--

.2141

.848

.152

1958-59

515

130

84

.2524

.1631

.496

.504-

1959-60

417

137

94

.3285

.2254

.873

.127

1960-61

554

143

99

.2581

.1787

.698

.302

1961-62

457

117

81

.2560

.1772

.724

.276

1962-63

768

188

113

.2448

.1471

.679

.321

1963-64

586

127

87

.2167

.1485

.807

.193

1964-65

978

180

CD

.1840

1966-67

674

CD

50

--

.0742

.889

.111

::'967-68

922

77

26

.0835

.0282

.989

.011

1968-69

1,089

31

--

.0285

-lO,883

11 Cannot use.

2.5003

1.8381

Average Survival Rate

.735

Average Mortality Rate

.265

I

I-'

&lt;l'
.po
I

�-165October,

JOB PROGRESS

REPORT

S t ate of

...::C...::o..:;;L;.;;o~RAD=...::O'____
_

Project No.

W-88-R-16
2

Work Plan No.
Job Title

Studies

Period Covered:

of Canada

April

1971

Migratory

Bird Investigations
6

Job No.
Goose Populations

in Colorado

Transplant

Areas

1, 1970 to March 31, 1971

Personnel:
C. Brown, E. Cochran, D. Coven, G. Crawford, E. Downer, H. Funk,
J. Grieb, J. Hatfield, J. Hobbs, R. Hopper, T. Lynch, S. Palm, J. Pogorelz,
C. Roberts, W. Russell, J. w~eeler and M. Szymczak.

ABSTRACT
During the 1970-71 hunting season 179 birds banded in areas outside of Colorado were reported recovered in north-central Colorado.
Summer banding locations contributing a significant number of banded birds were:
EdmontonCamrose area, Alberta; Dowling Lake area, Alberta; Cypress Hills area, Saskatchewan; Phillips and Garfield counties, Montana.
Birds banded as flightless young and recovered in north-central Colorado originated from all of the
above areas except Garfield County, Montana.
Approzimately
30 percent of the
birds banded in January, 1968 have been recovered in southwest Saskatchewan
compared to about 45 percent in north-central Colorado.
The recovery birds
banded as locals in Phillips County, Montana and Dowling Lake, Alberta, definitely classify these breeding areas as contributors to the Hi-Line population.
The recovery of birds banded as locals in southwest Saskatchewan in addition
to the recovery in southwest Saskatchewan, of birds banded during the winter in
north-central Colorado, indicates that area of Saskatchewan may be a major
contributor of geese to the Hi-Line population.
Based on trapped samples,
approximately 15 percent of the winter population in north-central is composed
of small Canada geese which are presumable breeding members of the arctic nesting short grass prairie population.
Approximately
90 percent of the Hi-Line
population wintered in Colorado during 1970-71.
An estimated 11,187 hunters
harvested 12,112 geese in north-central Colorado during the 1970-71 hunting
season, for an average season bag of 1.08 per hunter.
Only four percent of the
active hunters achieved the season maximum of six birds.
Approximately
62 percent of all birds harvested in the north-central permit area. were reported taken
in Larimer County.
A limited Canada goose hunting season in the San Luis Valley
resulted in an estimated 107 birds being harvested by 164 active goose season
permit holders for an average season bag of 0.65 birds per hunter.
Seven birds
were reported recovered which were banded outside the San Luis Valley; five of
these birds were banded in Hi-Line population breeding areas.
Goose counts
in the San Luis Valley totaled 1,490 birds on November 23, 1970, 1,050 on December 24, 1970, and 1,241 on January 5, 1971. Throughout the winter the large
majority of the geese in the San Luis Valley remained on the Monte Vista National
Wildlife Refuge.

��-167-

STUDIES OF CANADA GOOSE POPULATIONS
MichaelR.

IN COLORADO

TRANSPLANT

AREAS

Szymczak

Colorado has. had continuing success in establishing Canada goose nesting
populations through the use.;of gosling transplants in various selected parts
of the state. These programs have resulted in hlintable populations in
north-central Colorado and the San Luis Valley.
Data must be collected
continuously in order to determine the information needed to establish
sound hunting regulations.
In addition, certain ecological factors which
are pertinent to the long term management of these populations must also
be examined.
This study, which is in its first year, was established to
meet those objectives.
P. S. OBJECTIVE
To collect management da ca on Canada goose populations
from transplant.programs·within
Colorado.

SEGMENT
1.
2.
3.

which

have resulted

OJBECTIVES

To examine migration routes and/or harvest patterns and distribution of
Canada geese wintering in north-central Colorado and the San Luis Valley.
To estimate'hunting pressure on, and hunter harvest of Canada geese in
north-central Colorado and the San Luis Valley.
To make recommendations
for establishing a limited Canada-goose hunting
season in the San Luis Valley of Colorado and continuing the hunting
season on the north-central Colorado goose population.

METHODS

AND MATERIALS

Complete information on all banded Canada geese which have been reported
recovered in north-central Colorado has been requested from the Migratory
Bird Population Station.
This information has not yet been received.
Therefore, recovery data for north-central Colorado as well as the San Luis
Valley were hand tabulated and analyzed utilizing the periodic reports
distributed by the Migratory Bird Population Station.
Recovery data was
sorted as to banding location, age at time of banding and estimated breeding
area, if possible.
Canada goose tail fans collected during the 1969-70 hunting season were
measured in an attempt to determine the subspecific classification of the
harvest, and thus,possibly
delineate breeding areas.
Canada geese in north-central Colorado were counted on November 4 and 23,
December 22 and January 6 during the 1970-71 season.
Geese in the San
Luis Valley were censused on November 23, December 24 and January S.

�-168-

A mechanically random sample or 'hunters issued permits to hunt geese in the
north-central Colorado "Special Permit" goose hunting area were mailed
questionnaires
inquiring about their hunting activity and success.
All of
the hunters holding special permits to hunt geese in the San Luis Valley
were mailed similar questionnaires.
All data received were tabulated and
statistically analyzed.
Reconmenda t Lons were formulated for the 1971-72 goose season in the San Luis
Valley and north-central Colorado
and submitted to management personnel.

RESULTS AND DISCUSSION
North-Central
Breeding

Colorado

- The Hi-Line

Population

Range

The wintering Hi-Line population, which now generally terminates its southern
inigration in north-central Colorado, has undergone a substantial annual
increase 'in size since 1968. During this period the post-season wintering
population increased from approximately
17,000 to 45,000.
Low 'harvest rates,
expansion of the breeding range in northeastern Montana and the innnigration
of Canada geese of the short grass prairie population into Hi-Line wintering
areas are known factors that have contributed, in some degree, to the population increase.
In addition, breeding populations in southeast Alberta
and southwest Saskatchewan are believed to contribute 'a significant number
of geese to the Hi-Line population.
Breeding' areas of these Canadian, and
possibly other unidentified population segments must be specifically located
in order to monitor reproductive success, and thus provide a basis for
establishing hunting season regulations.
Foreign Recoveries.--During
the 1970-71 hunting season 179 foreign bands
were reported recovered in north-central Colorado.
An additional 36 foreign
banded birds were recaptured during post-season banding operations.
Both
types of recoveries are classified as to general area, if in a Province, or
County of banding if in the United States, in Tables 1 and 2, respectively.
The term direct recoveries refers to birds 'taken the first hunting season
after
banding.
All birds recovered that were banded in Nebraska, New Mexico, Oklahoma,
Texas, and Illinois were members of wintering populations.
All other birds
were banded during the sunnner or pre-season period.
Based on knowledge of the current distribution of the Hi-Line population,
the small number of indirect recoveries of Pacific Flyway birds banded at
scattered locations do not represent any specific breeding populations which
contribute significantly to the Hi-Line wintering flock at this time. These
birds were most likely transients, an example of the type of birds that occurs
in any population.
The birds from Big Horn County, Montana and Fremont
County, Wyoming are additional examples of misplaced Pacific Flyway birds.

�-169-

Table 1. Banding areas outside Colorado of Canada geese recovered in
north-central Colorado during the 1970-71 hunting season.

Area
Keewatin
TheIon River .
Alberta
Edmonton-Camrose Area
Dowling Lake Area
Brooks Area
Saskatchewan
North Battleford Area
Moose Jaw Area
Cyprus Hills Area

Direct

Indirect

o

1

6

6
7·
3

15

o

o

1
2

o

8

Montana
Lake County

o

2

Wyoming
Lincoln County

o

1

Rich County

o

1

California
Modoc County

o

1

3
0

0

o

Pacific Flyway

Utah

Central Flyway
Montana
Teton
Blaine
Phillips
Fergus
Garfield
Big Horn

20
0
0

3
39
1
37

0

1

0

3

1
1

Nebraska
Keith County

1

1

New Mexico
Mora County
Socorro County

1
5

0
5

Texas
Potter County

0

3

54

lis

Wyoming
Fremont
Albany

Total

.---~--

------

�-170-

+ab1e 2. Banding areas outside Colorado of Canada geese recaptured, postseason in north-central Colorado, January 1971.
Area

Direct

Indirect

1

Alberta
Edmonton-Camrose Area
Dowling Lake Area
Cypress Hi lIs

1

5
2

0

1

Saskatchewan
Swift Current -S• Saskatchewan R. Area
Moose Jaw Area
Cypress Hills Area

0
0
0

1
1
4

Montana
Blaine County
Phi llips County
Fergus County
Garfield County

0
3
0
0

1
4

Oklahoma
Alfalfa County

0

1

Texas
Potter County

0

2

New Mexico
Socorro County

0

2

Washington
Benton County

0

1

Nevada
Washoe County

0

1

0

1

5

31

Central Flyway

1
3

Pacific Flyway

Mississippi Flyway
Illinois
Alexander County
Total

�-171-

The geese banded in Teton and Fergus counties, Montana, were birds hand
reared and released in the respective areas. Most transplanted goslings
are chronic wanderers, particularly during their first migration season;
thus, they may end up as members of any wintering population.
The birds
from the North Battleford and Moose Jaw areas of Saskatchewan, as well as
birds from one banding location in the Edmonton-Camrose
area of Alberta,
were also tranplants.
Probably the best indication of breeding areas is obtained from examining
recoveries of birds wild-trapped and banded as locals, the term used for
flightless young, or possibly innnatures during the sllIlimer
period.
Most
likely birds classified as immatures captured during surrunerdrive-trapping
operations were actually locally raised birds which had attained the
capability but not the inclination to fly. All locals or summer banded
innnatures recovered or recaptured in north-central Colorado during the
1970-71 season are categorized by location and year of banding in Table 3.
Again, Pacific Flyway banded local birds as well as those from Big Horn
County, Montana and Fremont County, Wyoming, are most likely transients
that were produced in Great Basin Canada goose population breeding areas.
Locals originated from five specific areas in Alberta.
The Dusty Lake
area east to northeast of Camrose looks promising as a potential Hi-Line
breeding ,area. Since the number of banded birds in that population is not
known, the comparative numerical value of a single recovery canriot be
weighted.
The Dowling Lake area is a definite contributor to the Hi-Line
Population.
During the summer of 1970, 177 flightless geese were banded
at Dowling (Weaver et al., 1970).
Sixteen of these birds were recovered or recaptured (Tables 1 and'2) in
north-central Colorado during the 1970-71 season.
Nine of these were
locals (Table 3). Two locals from the Brooks area were recovered in Colorado. However, none of the 145 flightless b~rds banded near Brooks during
the summer of 1970 were recovered in north-central Colorado.
The final local banded by Alberta and recovered in 'north-central Colorado
during the 1970-71 season came from within the area which I have termed the
Cypress Hills area.
This artificially termed area is depicted in Figure 1
and is generally bordered by the South Saskatchewan River on the north,
Swift Current, Saskatchewan on the east, Medicine Hat, Alberta on the west,
and the 49th parallel
on the south. According to recoveries of both
summer and winter banded birds, this area is apparently quite important to
Hi-Line Canada geese.
Twelve geese that were banded in the Cypress Hills
area of Saskatchewan were recovered or recaptured in north-central Colorado
during 1970-71 (Figure 1). Ten of the 12 were banded as locals.
All of
the birds were banded in 1965 or before.
Apparently no summer banding of
Canada geese has been done in that part of Saskatchewan since 1965. The
Significance of Cypress Hills area as a Hi-Line population breeding area
should definitely be explored further.
A total of 59 banded geese recovered during the 1970-71 season in northcentral Colorado originated from Phillips County, Montana (Table 1).
Phillips County, which contains Bowdoin National Wildlife Refuge, has
historically been a major contributor of Canada geese to the Hi-Line population.
Garfield County contributed 37 banded birds to the harvest in
Colorado (Table 1). However, only qne of these birds was a "young of the
year" (Table 3). Apparently birds banded in Garfield County were captured
in an area used predominan~ly for moulting by non-breeding birds.

�-172-

Table 3. Banding locations of Canada geese classified as wild-trapped
locals which were recovered or recaptured in north-central Colorado during
the winter of 1970-71. 1/

Location

Before 1965

Alberta
Edmonton-Camrose Area
10 mi. E of Camrose
Dusty Lake
Dowling Lake Area ...
Brooks Area

Year of Banding
1965 1966 1967 1968

1970

1/

2, (2)-1/
(1)2

1

1

Cypress Hills
40 mi. NE of Medicine Hat
Saskatchewan
Cypress Hills
Supreme Area
Crane Lake
Maple Creek Area
Many Island Lake
Cypress Lake
10 mi. N. of Eastend

1969

1

4
1
1
1
2
1

Central Flyway
Montana
Blaine County
Phillips County
Garfield County
Big Horn County

2
7

1

7

13

4, (5)1/19,(1)1/
(1)1/
1

Wyoming
Fremont

1

Pacific Flyway
Montana
Lake County

2

Wyoming
Lincoln County

(1)

Utah
Rich County

1

1/ Numbers in parentheses indicate geese classified as immatures when
banded during the summer period.

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.-

-

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-

-

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~

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-

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-

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-

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Iv: '

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o

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GVLL
LAKe:

c:c.P Bute r

\'/j~'-:

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as a Locol

G!I

Bonded

as on Ad uII

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Experimental

I

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Banding Loca t Lons

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t-;Iie~'jp,;.

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in ~OTth--ee.rrtTaJ:---E:olotado-dttrin? 197(:)-71.

r,
(

.

�-174Winter Bandings - North-Central Colorado.--In January of 1968, 478 Canada
geese were banded in north-central Colorado.
Saskatchewan and Colorado have
been the major recovery area for these geese (Table 4). A significant
number of these. birds have been reported taken in the previously defined
Cypress Hills area (Fig. 2). There are three possible explanations for the
Cypress Hill area being a major recovery area for these Hi-Line birds:
(1) these birds are members of populations which breed farther north and
move into the Cypress Hills area prior to the start of any Canadian waterfowl season, (2) these are non-:-breed.ingsub-adult birds which move out of
the primary eastern Montana breeding area into the Cypress Hills area or
possibly farther north to moult, then are harvested on their return south,
(3) these are birds which breed in the Cypress Hills area. There is Some
evidence supporting each of·the above hypotheses.
In support of (1) birds
from Dowling Lake, Alberta are known to pause in their southern migration
in the Medicine Hat area of Alberta (Weaver et al. 1971), (2) there has
been an annual decrease in the nUITber of 1968 post-season banded birds
recovered in the Cypress Hills area (Table 4 Saskatchewan).
This decrease
would be considered normal had not the total number of recoveries remained
stable because of an annual increase in· the number of bands recovered' in
the other major harvest area, north-central Colorado.
However; increased
total harvest in Colorado definitely complicates analysis of these recovery
data.
(3) As discussed previously, there has been a significant number of
recoveries in no r t h+ce nt r.aI Colorado of birds .t.hat;
were banded as locals
in the Cypress .Hills area.

Winter Banding - Southeast Colorado. --The shift of Some arctic nesting
short grass prairie Canada geese into Hi-Line wintering areas was first
noted during the 1967-68 season when two birds previously banded in .the
Colorado's short grass wintering range were recovered in north-central
Colorado.
Short. grass banded birds have been taken in the Hi-'Line area
annually since that time, not only' from southeast Colorado bandings, but
also from Texas.
The significance of short grass birds, in terms of numbers,
in the Hi-Line area will be examined later in this report.

Summary.--Analysis
of band recoveries of Canada geese reported harvested
or recaptured in north-central Colorado has given preliminary indications
of Some specific breeding areas of Hi-Line Canada geese.
The Dowling Lake
area of Alberta joins Phillips County, Montana as a known reproduction
area for Hi-Line birds.
Some populations in the Edmonton-Camrose
area also
seem to be contributing
to this population.
Specifically, birds in the
Dusty Lake area and a newly developing transplanted population at Oliver
Lake, Alberta are migrating into the Hi-Line winteririg area.
The Cypress Hills area of Saskatchewan and adjacent Alberta. may possibly be
an important reproductive area for Hi-Line birds.
The recovery of birds
banded in the winter. in north-central Colorado in the Cypress Hills add
importance to that area •

�J

°1

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111

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Island

r'--.~
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Lake
f"_
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yl

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MAPLE

G'
.;

-

I

-

--

a CnEEK

I

e
,

Firs! Year-Bonded

as Imm.

Fitst Year-Bonded

as Adult

e

Other

Years-Bonded

as Imm.

I

X

Other Years-Bonded

0&amp; Adult

I

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I

t

I

I

,

20 mil es

I
~

ra!~o\'/ki
1.0 k e

~

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Fig.

2.

Recovery

I

locations

Jn

-

o

I

X

E:'STEND

tbl Cypress Hi] 15 area of C.mlad~ geese 'bimcled-in

II

(

I

I

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..

-

X

La,e

I
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�-176-

Table 4. Distribution of recoveries
in north-central Colorado, i967-68.

of Canada geese banded post-season

1968-69
No. Percent

RecoveE2: Year
1969-70
1970-71
No. Percent
No. Percent

No.

British Columbia

0

0.0

1

3.7

0

0.0

1

1.3

Alberta

1

4.2

3

11.1

0

0.0

4

5.3

Saskatchewan

11

45.8

9

33.3

3

12.0

23

30.3

Montana

4

16.7

0

0.0

2

8.0

6

7.9

Idaho

1

4.2

1

3.7

0

0.0

2

2.6

South Dakota·

0

0.0··

1

3.7

0

0.0

1

1.3

Wyoming

0

0.0

1

3.7

0

0.0

1

1.3

Nebraska

0

0.0

1

3.7

0

0.0

1

1.3

North Central

5

20.8

10

37.0

19

76.0

34

44.7

Southeast

1

4.2

0

0.0

1

4.0

2

2.6

Arizona

1

4.2

0

0.0

0

0.0

1

1.3

Total

24

Area

Total
Percent

Colorado

Migration

Routes or Harvest

27

25

76

Patterns

The recovery distribution of birds banded in north-central Colorado postseason, 1968 were presented in Table 4. These data indicate Saskatchewan
is the major harvest area north of the terminal wintering area in Colorado.
Generally, banded birds reported harvested in Saskatchewan were taken from
late September to mid-October.
Only six birds banded in 1968 have been
taken in Montana and these show no specific trends for period of harvest.
Recoveries indicate that a good portion of the 1968 banded birds were
definitely present in north-central Colorado by November 10.
Analysis of foreign banded birds recovered in north-central Colorado during
the 1970-71 season indicate some variation in movement of birds into Co1o~
rado. Recovery information indicated that at lea.st a portion of the birds

�-177-

from Phillips and Garfield counties in Montana were in Colorado prior to
the beginning of the season on November 10. Dowling Lake birds apparently
did not move into the area until late November-early December.

Wintering

Flock Composition

As was mentioned previously, some birds of the short grass prairie population
have been m,oving.irito Hi-L:Lne popuiation wintering areas.
These birds are
arctic nesters and much smaller in. size than the prairie nesters of the HiLine population.
In an effort to .obt a i.na minimal estimate of the number o-f short grass birds
in the Hi-Line wintering area, a sample of hunters were asked to send in
complete tail fans from the f Lr.s t two geese which they bagged duri.ng the
1969-70 season.
The center rectrices were measured and classified according
to the method developed by Dzubin (unpublished data) which separate "large"
type Canada geese from small geese.
Of 143 usable tails, only six, or 4.2
percent were classified as being small geese. Assuming there was no variation
in migration chronology between the large and small birds, which would
affect early season,availability,
a projected 175 small geese would have been
harvested during the 1969-70 season in north-central Colorado.
Hunter selection for the larger geese negates the possibility of estimating
what portion of the entire wintering population consisted of small birds.
The number of band recoveries of small Canada geese in north-central Colorado have been inconsistent in reference to the size of the harvest.
Continued recoveries annually only indicate that small birds of the short
grass prauie are indeed present in the Hi-Line area.
From the 1967-68
season through the 1970-71 season, there have been three, seven, four, and
nine short grass prairie banded birds taken during the respective seasons in
north-central Colorado.
One of the birds taken during the 1970-71 season
was an adult male of intermediate size when banded (7 lbs., 80z.),
atypically large fora short grass prairie bird.
An estimate of the percent of small birds in the 1970'-71 post-season HiLine population was attained during trapping and banding operations.
Of
the 990 birds banded, 147 or 14.8 percent were considered small birds.
These birds were all captured at one time in a very large walk-in type trap,
thus the trapped birds may be representative of the post-season population.

Distribution.--Coordinated
inventories throughout the te~inal
wintering
range of the Hi-Line population during the winter of 1970-71 revealed that
approximately 90 percent of the population wintered in Colorado Crable 5).
The distribution of geese within Colorado is presented in Table 6 ~ith
comparative totals for previous years available in Table 7. The increase
in the number of birds utilizing the Longmont-Denver-Boulder
area was the
only major shift .recorded during the 1970-71 season.
This change probably
resulted from hunting pressure in areas previously closed to goose hunting
in the Ft. Collins-Longmont
area.
'

�-178-

The decline in the number of geese present in the area between the December
count and the January count may have been the result of birds dispersing
from the area during a late December early January heavy snow and cold
period.
It is also possible that the late December count may have been too
high and the total drop in the number of birds from November to January
was an indication of harvest.

Table 5.

Results

of Hi-Line Canada goose population

inventories,

1970-71.

Nov. 24

Dec. 22

Jan. 5

Location

Nov. 4

Saskatchewan

1,800

Montana

12,622

1,812

1,335

800

Wyoming

1,770

1,990

2,635

2,710

Colorado

14,650

48,050

51,753

37,870

307

1,074

2,644

3,332

31,149

52,926

58,367

44,712

New Mexico

Totals

Hunting

Pressure

and Harvest

An estimated 11,187 hunters harvested 12,112 geese in north-central Colorado during the 1970-71 season (Tables 8 and 9). Larimer County continues
to be the major area of hunter activity and goose harvest (Table 10). The
total number of hunter trips and geese bagged increased over 1969-70 levels
in all counties except Morgan (Table 10). The large increase in the number
of birds in Boulder County through the 1970-71 season (Table 7) was
generally not reflected in harvest and hunting success in that County
(Table 10).
The majority of the active hunters continue to be unsuccessful (Table 11).
Only about four percent of the active hunters achieved the season maximum
of six birds.

San Luis Valley
The entire San Luis Valley was open to goose hunting during the 1970-71
season for the first time since Canada goose restoration efforts began in
the Valley.
Two hundred permits were issued for the taking of one goose
per permit in the counties of Hinsdale, Mineral, Rio Grande, and Saguache
east of the continental divide; Costilla and Alamosa counties north ofU. S.
160 and west of U. S. 285; and Conejos County west of U. S. 285. Goose
hunting in the remainder
of the Valley
was subject to the general Central
Flyway bag limits of two geese per day and four in possession.
Significant
closures to goose hunting were present in the San Luis Lakes area, and on
portions of the Monte Vista and Alamosa National Wildlife Refuges.

�-179Table 6.

Results of north-central Colorado Canada goose surveys, 1970-71.

Area
Ft. Collins-Loveland Area
Lindenmeier Lake
Reservoir No. 8
Reservoir No. 8 Annex
Elder Reservoir
Reservoir No. 5
Reservoir No. 6
Bureau of Standards
Douglas Reservoir
Rocky Ridge Reservoir
Cement Plant Reservoir
Terry Lake
Mountain Supply' Reservoir
College Lake
Herring Lake
Claymore Lake
S terling Pond
Watson Lake
Dean Lake
Anderson I s Pond
Greenwalt (Nelson) Lake
Fossil Creek Reservoir
Boyd Lake
Horseshoe Lake
Hollister Lake (Windsor)
New Windsor Reservoir
Woods (Eaton) Lake
Timnath Reservoir
Cobb Lake
Black Hollow Reservoir
Parkwood Lake
Boxe1der Reservoir
Sub-total
Boulder-Longmont Area
Ish Lake
Terry Lake (Longmont)
McIntosh Lake
Foothills Reservoir
Faivre Ponds
Swede Lake
Dodd Lake
Gaynor Lakes
Baller Lake
Boulder Reservoir
Va1mont Reservoir

Sub-total

Nov. 4

Nov. 23

Dec. 22

Jan. 5

.2,500
720
320
460
170
1,020

2,200
150
46

25'.,
1,700

250
275

o

o
o

197
63
250
140

164

830
200
125

o

o

o
o

425

155
35

o
280
290

o
2,100
170
1,750

o

o
o
220

o
12

o
o

o
o

o
600

o
520
1,480

o
o
o

o
o

o
o

o

o
o
20
1,700

o

o

4,200
120
.50
1,025
1,025
450

10,947

13 ,500

o

o
o

o
o

6,000
1,400

400
400

4,500
6,000
75
250
225
7,000
6

850
155
63

o

o

15
50
150

25

o
150

o

o

o

o

450
7,000
535
52
4,500

100
2,100
175
220
400

o

o

o
o

70
2,300

o
o

12,612

29,970

36,034

19,879

170
90

55
9,300
1,225

o
o

o

o

o

o

4,200
330
185
368

2,095

o
o

o
o

o
16
205

o

o

530

163
30
12
35
22
175
3,400

725
5,500

11,233

790

14,417

11 ,308

13 ,549

o

o
o

o
o

o

o

o

-------------------------------------------------------.---------------------

�-180-

Table 6. Results. of north-central Colorado Canada goose surveys, 1970-71
(continued).
Area

Nov. 4

Nov. 23

Dec. 22

Jan. 5

Denver Area
C leav iew Reservo ir
Mohn La.ke
Sloans Lake
Stand ley Lake .
Bowles Lake Area
Marston Reservoir
Grant's Reservoir
Bonfil 's Es tate
Denver City Park
Cherry Creek Reservoir
Denver Metro Area
Ricketson's Pond

0
0
0
0
0
0
0
0
0
0
.900
0

5
66
109
155
1,200
75
300
120
290
23
0
0

0
0
105
0
933
0
0
0
240
38
0
40

0
0
135
0
1,282
0
0
0
350
250
0
0

Sub-total

900

2,293

1,356

2,017

190
40
0
18
0
0
100
0
0
0

340
400
400
25
15
35
60
35
60

150
400
400
0
0
650
0

0

400
30
500
350
85
610
425
475
0
50

0
0

0
0

130
0

800
25

348

1,370

3,055

2,425

14,650

48,050

51,753

37,870

Brighton-Greeley-Fort Morgan Area
Barr Lake
Horsecreek Reservoir
Prospect Reservoir
Latham Reservoir
Milton Reservoir
Empire Reservoir
Riverside Reservoir
Jackson Reservoir
Bijou Reservoir
K-4 Ranch
South Platte River, Greeley Ft.
Morgan
Mile High Duck Club
Sub-total
Grand Total

0

0
0

�-181-

Table 7.

Winter inventories of the Colorado Hi-Line Canada goose population.

Count Date

Fort Collins,~ove1and

Longmont-Bou1derDenver

1967
January 9

9,739

2,883

991

13 ,613

1968
January 10

12,217

4,029

678

16,924

1968-69
November 20
December 19
January 2 &amp; 13

15,848
20,905
19,693

3,461
4,236
4,874

2,667
1,170
775

'21,976
26,311
25,342

1969-70
November 5
November 28
December 23
January 6

8,737
31,350
18,522,
30;650

2,255
3,782
5,668\
5,060

390
1,374
1,259
1,914

11 ,382
36,506
25,499
37,624

1970-71
November 4
November, 23
December 22
January 6

12,612
29,970
36,034
19,879

1,690
16,710
12,664
15,566

348
1,370
3,055
2,425

14,650
48,050
51,753
37,870

Brighton~Gree1ey-For t Morgan

Total

Table 8,. Hunter activity and success in permit area.
Number
Permits Issued

Est. Number
Active Hunters

Average
Days Hunted

1964-65

1,608

1,427

6.1

0.48

1965-66

2,335

1,578

6.5

0.53

1966-67

3,996

2,910

5.1

0.41

1967-68

3,000

2,256

6.7

0.55

1968-69

3,000

2,379

9.2

1.11

1969-70

8,342

6,149

7.4

0.68

1970-71

13 ,611

11,187

6.9

1.08

Year

Average
Seasonal Bag/Hunter

�-182-

Table 9.

Distribution of harvest, by county, in the permit area.

Year

Larimer

Weld

Boulder

Morgan

Adams

Total

"'"""'-

1964-65

504

181

1965-66

665

144

29

838

1966 -67

764

409

11

1,184

1967-68.

944

265

37

1,246

1968-69

1,584

886

161

2,631

1969-70

2,431

1,112

383

146

100

4,172

1970-71

7,486

3,544

620

93

370

12,112

Table 10.
1970-71.

685

Hunting pressure and harvest, by county, in north-central Colorado,

Larimer County 1/

Estimated Number
1969-70

Estimated Number
1970-71

Total Individual Hunters

3,535 (±

6,977 (±

Total Hunter Trips
Total Geese Bagged

226)

Percent Change
From 1969.,.70

328)

+ 97.4

24,139 (±2,722)

43,046 (±3,851)

+ 78.3

2,431 (± 411)

7,541 (±

+210.2

868)

Ave. Hunter Trips/Hunter

6.8

6.2

8.8

Ave. Bag/Hunter

0.69

1.08

+ 56.5

Ave. Bag/Hunter Trip

0.101

0.175

+ 73.3

1/ Larimer County permit area enlarged in 1970-71.
.

.

----------------------------------------------------------------------------.

,

�-183-

Table 10. Hunting pressure and harvest, by county, in north-central.
Colorado, 1970-71 (continued).

Weld County·

Es t Imat.edNumber
1969-70

Estimated Number
1970-71

Percent Change
From 1969-70

Total Individual Hunters

2,461 (±

212)

4,293 (+ 308)

+ 74.4

Total Hunter Trips

12,742 (±1,867)

22,557 (±2,677)

+ 77.0

Total Geese Bagged

1,112 (±

3,497 (±

+214.5

281)

595)

1.9

Ave. Hunter Trips/Hunter

5.2

5.3

+

Ave. Bag/Hunter

0.45

0.81

+ 80.0

Ave. Bag/Hunter Trip

0.087 ..

0.155

.+ 78.2

Boulder County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged

728 (±

131)

4,483 (±1,215)
383 (±

179)

186)

+ 69.1

7,543 (±2,207)

+ 68.3

620 (± 229)

+ 61.9

1,231 (±

Ave. Hunter Trips/Hunter

6.2

6.1

1.6

Ave. Bag/Hunter

0.53

0.50

5.7

Ave. Bag/Hunter Trips

0.085

0.082

3.5

Morgan County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged

575 (± 231)

759 (±

147)

+ 32.0

2,300 (± 639)

2,147 (±

580)

6.7

93 (±

·64)

- 36.3

146 (±

65)

Ave. Hunter Trips/Hunter

4.0

.2.8

- 30.0

Ave. Bag/Hunter

0.25

0.12

- 52.0

Ave. Bag/Hunter Trip

0.063

0.043

- 31.7

-----------------------------------------------------------------------------

�-184-

Table. 10. Hunting pressure and harvest, by county, in north-central
Colorado, 1970-71 (continued).

Adams County
Total Individual Hunter
Total Hunter Trips
Total Geese Bagged

Estimated Number
1969-70

Estimated Number
1970-71

Percent Change
From 1969-70

337 (± 212)

592 (±

131)

+ 75.7

1,730 (± 645)

2,294 (±

818)

+ 32.6

361 (±

147)

+261.0

100 (±

80)

Ave. Hunter Trips/Hunter

5.1

3.8

- 25.5

Ave. Bag/Hunter

0.30

0.60

+100.0

Ave. Bag/Hunter Trip

0.058

0.157

·+170.7

Entire Permit Area
Total Individual Hunters

6,149 (±

201)

11,187 (± 254)

+ 81.9

Total Hunter Trips

45,394 (±3,598)

77 ,587 (2:4 ,403)

+ 70.9

Total Geese Bagged

4,172 (±

12,112 &lt;±1,005)

+190.3

583)

Ave. Hunter Trips/Hunter

7.4

6.9

6.8

Ave. Bag/Hunter

0.68

1.08

+ 58.8

Ave. Bag/Hunter Trip

0.092

0.156

+ 69.6

�-185Table 11. Estimated distribution of season bag for active hunters in the
north-central Colorado permit area.

1968-69
Number
Percent

Hunting Season
1969';'70
Number'
Percent'

1,299

54.6

4,263

1

420

17.6

2

259

3

,Number

1970-71
Percent

69.3

6,588

,58.9

836

13 .6

1,620

14.5

10.9

452

7.4

1,073

9.6

142

6.0

238

3.9

564

5.0

4

103

4.3

153

2.5

528

4.7

5

78.

3.3

138

2.2

342

3.1

6

78

3.3

69

1.1

472

4.2

Total

2,379

100.0

6,149'

'100.0

11,187

100.0

Season Bag

°

Foreign Band Recoveries
Seven banded birds were reported recovered in the San Luis Valley during the
1970-71 season that were banded outside of the Valley (Table 12). The birds
from Alberta and Montana, as well as those from north-central Colorado,
are from Hi-Line population breeding areas. The San Luis Valley lies along
the route which Hi-Line population birds might be expected to take to their
historical Wintering grounds along the lower Rio Grande River in New Mexico.
However, the time of recovery of the birds banded at Dowling Lake, early
January, is much later than the normal migration period for Hi-Line birds.
It is possible that the Dowling Lake birds had moved out of north-central
Colorado during the late December heavy snow and cold weather period. An
additional bird from Dowling was taken in early January in the Salida area,
north of the San Luis Valley. A small influx of birds was noted in the
San Luis Valley between the December 24 and January 5 inventories and could
have been missed from earlier counts (Table 13).
The bird from Wyoming was banded at Wheatland Reservoir, a traditional
moulting area for birds of Great Basin Canada goose populations. As was
noted under Work.Plan 2, Job 2 of this Federal Aid Report, sub-adults from
the San Luis Valley may be beginning to use Wheatland Reservoir for moulting
purposes.
Distribution and Harvest
Aerial surveys conducted during the 1970-71 hunting season indicated that
the majority of the birds utilized the Monte Vista National Wildlife Refuge
during the season ,(Table 13). The majority of the 107 birds harvested in
the permit area was extrac~ed from the population utilizing the Monte Vista
Refuge in Rio Grande County (Table 14).

�-186Table 12. Banding areas outside the San Luis Valley of birds recovered
within the San Luis Valley during the 1970-71 hunting season.
Area

Number of Recoveries

Recovery Dates

2

1-2-71
1-5-71

1

11-14-70

1

1-12-71

2

11-6-70
.11-14-70

1

11-6-71

Alberta
Dowling Lake
~~•.
mtana
Phillips County
Wyoming
Albany County
Colorado
North-central
Texas
Waggner Ranch-Panhandle

Table 13. Res~lts of Canada goose surveys during the 1970-71 hunting
season in the San Luis Valley.

Location
Monte Vista Refuge

November 23

Number of Geese
December 24
January 5

1,125

975

1,000

285

72

30

Russell Lakes

o

o

16

Smith Reservoir

o

3

160

o

o

35

80

o

o

1,490

1,050

1,241

Rio Grande River
De 1 Nor te-ALames a

Conejos River
McIntire Springs-Rio Grande
Sanchez Reservoir
Total

�-187-

Table 14.
area.

Distribution

of harvest,

Year

Rio Grande

Alamosa

Saguache

74

31

2

1970-71

by county

in the San Luis Valley

Conejos

o

permit

Total

107

It was estimated that 164 of the 200 permit holders actually hunted geese
during the special season in the San Luis Valley (Table 15). Rio Grande
and Alamosa counties received the greatest amount of hunting pressure but
success was much greater in Rio' Grande (Table 16).
Since the majority of the harvest was expected to be extracted from locally
raised birds that had not been hunted previously, the first part of the
season was expected to be the heavy harvest period.
However, the maJor
harvest period in the permit area extended from late November to late
December (Fig. 3). Possibly the hunting pressure was not as great as
expected because the opening week ran concurrently with the last few days
of the big game season.

Table IS.

Year

1970-71

Hunter

activity

Number
Permits Issued

200

and success

in the San Luis Valley

permit area.

Est. Number
Active Hunters

Average
Days Hunted

Average
Seasonal Bag/Hunter

164

3.4

0.65

LITERATURE
Weaver, H., C. Lacey, and D. Neave.
Committee Annual Report, 1970.

CITED
1971.

Alberta Waterfowl

54 p.

Prepared by ---IZe-'_J../-f_:t.-:-c...;... A-,-:l =d..L;;;.....[""". , 6--' _"'_X_;~-:-I_+, /.,..,';:/;&lt;....-r'-'}-'-Z~c~.),~d;.=..:;;;,L:;..-/_
M:J-chaelR. SzymczaY
'{_"j
&lt;~(.:;(
Assistant Wildlife Researcher
v

zr

Technical

�-188Table 16. Hunting pressure ·andharvest,
Valley special goose season, 1970-71.

Rio Grande County

Total Individual HUnters
Total Hunter Trips
Total Geese Bagged
Average Hunter Trips/Hunter
Average Bag/Hunter
Average Bag/Hunter Trip
Alamosa

Es timated Number

107
278
74
2.6

0.69
,0.27

82

237
.31
2.9
0.38
0.13

10
28
2

2.8
0.20

0.07

County

Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Average Hunter Trips/Hunter
Average Bag/Hunter
Average Bag/Hunter Trip
Entire

the San Luis

County

Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Average Hunter Trips/Hunter
Average Bag/Hunter
Average Bag/Hunter Trip
Conejos

during

County

Total Individual Hunters
Total Hun t.er Trips
Total Geese Bagged
Average Hunter Trips/Hunter
Average Bag/Hunter"
Average Bag/Hunter Trip
Saguache

by county,

6
13

o
2.2

o

o

Permit Area

Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Average Hunter Trips/Hunter
Average Bag/Hunter
Average Bag/Hunter Trip

164

(± 3)

556 (±20)
107 (± 4)
3.4
0.65
0.19

1970-71

�-189-

.

25

20
-I..l
(I)

Q)

::-J.I
C\1

::r:
IH
0

15

-I..l

c::
Q)

0

J.I

Q)

p..,

10

5

Oct. 31
to
Nov. 13

Nov. 14
to
Nov. 27

Nov. 28
to
Dec. 11

Dec. 12
to
Dec. 25

Dec. 26
to
Jan. 8

Jan. 9
to
Jan. 17

Time Period
Fig. 3. Percentage distribution by time period of Canada goose harvest in the
San Luis Valley permit area during the 1970-71 hunting ~eason.

��-191October,

1971

JOB PROGRESS REPORT

State of
Project

COLORADO
No.

W-88-R-16

Work Plan No.
Job Title

3
Investigation

Period Covered:

December

Migratory

Bird Investigations

Job No.

6

of Mallard Management

Units of Eastern

1, 1970 through April

1, 1971

Colorado

Personnel:
Richard Hopper, Mike Szymczak~ Robert Kitzmiller, Larry Green,
Dale Coven, Jack Corey, Michael Robinson, C1ait Braun, Velma Fredrickson,
J. Edward Kautz and Howard Funk of the Division of Game, Fish and Parks;
Jack E. Randall of the Bureau of Sport Fisheries arld Wildlife.

ABSTRACT
Winter trapping efforts in eastern Colorado produced a total of 5,884 mallards
banded between mid~January through February, 1971. An excellent year of production on breeding grounds resulted in some of the highest winter census
figures for Colorado since 1957 and 1958, the last period of peak production.
A late November census indicated about 470,000 birds wintering in Central Flyway Colorado, about 430,000 in late December, and about 310,000 in early January.
Sex ratio counts again suggested about 60 percent of the Wintering mallard population was composed of males.
The point-system regulation in effect
in most Central Flyway states, which directs pressure toward the mallard drake,
evidently had minimum effect on altering sex composition of the flocks, especially in Colorado.
Harvest in Colorado increased substantially even though
estimated number of hunters was up only slightly from the 1969 season.
Greatest
increases in hunting pressure and harvest occurred in the north-central and
southeast portions of the state.
Programming and machine processing of all banding and recovery data gathered since 1963-64 was accomplished during the segment
but examination of printout material will not be completed until the next segment.

�-192-

RECOMMENDATIONS

1.

Recovery data for winter banded mallards in the San Luis Valley are
similar to those obtained during preseason periods and the Valley
does not seem to be as important a wintering area for northern birds
as other Central Flyway Units.
Thus, winter banding in the San Luis
Valley should be terminated.

2.

Banding quotas should remain at between 6 to 8 hundred mallards per
study unit, depending upon success of trapping efforts.
Every effort
should be made to obtain at least 600 birds in each of the eight study
areas.

3.

Analysis
1970-71

of all IBM tapes, including
season, should be completed

the updating tape which
in the 1971-72 segment.

covers the

�-193-

INVESTIGATION OF MALLARD
MANAGEMENT UNITS OF EASTERN COLORADO
H6ward D. Funk

Winter banding of mallards was continued in 1970-71 for the purpose of
investigating populations of birdswinteririg
in the Central Flyway portion of Colorado and dividing the State into management units if feasible.
Main criteria utilized in defining groups and areas are differences in
mortality and migration characteristics
of the birds.
The study was
begun in 1963-64 and expanded in recent years to a cooperative Central
Flyway investigation of management units.
A report entitled "Justification of the Central Flyway High Plains Mallard Management Unit" (Funk
et al. 1971) includes some of the data cdllected in Colorado through
1969-70. However, Colorado data are now being analyzed more intensively
by the study units designated upon initiation of the investigation.
This
analysis will also include data collected through the hunting season of
1970-71 and should be completed in the 1971-72 segment.
Thus, this report
will cover only results of banding efforts and sur'veys conducted within
the designated reporting period.

P. S. OBJECTIVE
To develop

a harvest

formula

for Colorado

mallard

management

units.

SEGMENT OBJECTIVES
1.

To trap and band mallards after hunting season in the following eight
study areas for the purpose of obtaining migration, life history and
annual mortality data:
(a) South Platte Valley Denver to Greeley,
(b) South Platte Valley Greeley to Fort Morgan; (c) South Platte Valley
Fort Morgan to Sterling, (d) South Platte Valley Sterling to Julesburg,
(e) the Fort Collins area, (f) Bonny Reservoir, (g) Arkansas Valley
Pueblo to Rocky Ford, and (h) Arkansas Valley Rocky Ford to the Kansas
line. Feasibility of winter banding in the San Luis Valley will also
be investigated.

2.

To conduct surveys for the purpose of obtaining winter popUlation estimates of mallards; age and sex ratios in the hunter's bag, wintering
populations, and trap samples and; hunter pressure and harvest data by
study area.

3.

To gather and tabulate band recovery data from the Bird Banding Office
from mallards banded in Colorado since 1963-64 for use in making preliminary estimates on migration and harvest routes and annual mortality
for use in final analysis during a later segment.

�-194-

4.

To conduct further special mallard seasons designed to take advantage of surplus birds where feasible.

METHODS AND MATERIALS
Trapping methods and materials were described in earlier segment reports
with cage traps being the main type utilized in 1970-710 Trapping began
immediately after hunting season, about mid-January, 1971, and continued
through February. A sample of birds was again obtained in the San Luis
Valley, making a total of nine study areas.
Periodic aerial censuses were again conducted to estimate numbers of ducks
and movement by time period and area. Ground sex ratio counts were conducted where and when possible to allow a continuing annual account of
possible changes in sex composition of mallards in the respective study
units. Hunter performance checks were conducted by Division personnel in
an effort to evaluate hunter attitude and willingness to abide by pointsystem regulations in effect during the season. The latter data were
combined with those gathered in other states having similar regulations to
allow an overall analysis.
Tapes containing mallard banding and recovery data for Colorado since initiation of this study were obtained from the Bird Banding Laboratory at
Patuxent Wildlife Research Station. Programs for tabulating, sorting and
complete analysis of data by machine processing were constructed by Mr.
J. Edward Kautz, Wildlife Biology graduate from Colorado State University.
Program analysis included examination of migration routes utilized by the
various age and sex classes of birds banded, movement between banding sites
or study areas within Colorado, fall movement down the Flyways by age and
sex by time periods, and mortality and survival rates of the various age
and sex classes. These programs are designed so that, with a minimum of
alteration, either card or tape records of recoveries can be used to update
any of the analyses as more harvest information is received.
RESULTS AND DISCUSSION
Trapping and Banding
Nine study areas were again sampled in winter banding efforts with results
displayed in Table 1. Quotas of birds to be banded in each unit (Fig. 1)
remained at 600 birds with as equal proportions by age (adult and immature)
and sex as possible. These quotas were reached or exceeded in six of the
nine units with a total of 5,884 birds banded. As always, adult males were
most prevalent in trap samples. Usually the adult female quota is most
difficult to obtain but immature females were also in short supply in samples
in 1970-71. It should be noted that percentages listed in Table 1 by age
and sex are not representative of trapping ratios because many trapped males
were released unbanded.

�Table 1.
Numbers and percentages of mallards in the banded sample by sex, age and area, eastern
Colorado 1970-71.
Adult
Percent

Female
No.
Percent

Immature
Male
Female
No. Percent
No. Percent

156

26.2

145

24.3

159

26.7

136

22.B

664

164

24.7

112

16.9

215

32.3

173

26.1

BOO

240

30.0

191

23.9

156

19.5

213

26.6

BOO

227

2B.4

iB7

23.4

23B

29.7

··148

18.5

800

242

30.3

201

25.1

221

27.6

136

17.0

Bonny Reservoir

754

198

26.3

130

(12) Rocky Ford-Lamar

468

133

28.4

127

27.1

87

18.6

121

25.9

465

151

32.5

128

27.5

117

25.2

69

14.8

537

135

25.1

124

23.1

137

25.5

141

26.3

5,884

1,646

28.0

1,345

22.8

1,539

26.2

1,354

23.0

Management Unit

No.
Banded

No.

(4)

Fort Collins

596

Denver-Greeley
Greeley-Ft. Morgan

(6)
(3)

(2)
(1)
(9)

Ft. Morgan-Sterling
Sterling-Julesburg

Male

I

(13) Pueblo-Rocky Ford
(15) San Luis Valiey:

Totals

17.2

I-'

209

27.7

217

2B.8

1.0
VI
I

�Ul

"
'"

W
Z

•••••••
~

~

M
c.:I
;
~~
~~

:5;
t-;;

I

"
~
0
I

"

0

u
&gt;(

~

w

;:
w
Z

�-197-

Tagging and banding in the San Luis Valley had to again be concentrated
on the Monte Vista National Wildlife Refuge because of a lack of other
suitable areas. Retraps were quite numerous, however almost all were
from pre-season bandings in the San Luis Valley and South Park. This
factor plus examination of recoveries from San Luis Valley winter
bandings in other areas has prompted the decision to terminate winter
banding efforts in the Valley and rely on pre-season banding data for
the San Luis Valley population.
Winter Sex Ratio Counts
"

Sex ratio counts were conducted in a number of eastern Colorado locations
in November, December and January. Lumped data by month on samples of
about 5,300, 8,200 and 2,800 mallards produced percentages of about 61,
60 and 59 drakes, respectively. This information suggests annual hunting
pressure in Colorado is not sufficient to greatly alter sex ratios in the
popUlations through the hunting season even though the point-system
regulation directs pressure toward the drake.
Sex ratio data obtained after hunting season by unit of study are.shown
in Table 2. Only five un~ts were sampled during this period but other
areas should have been similar, approximating 60 percent drakes overall
which is similar to results of previous efforts since 1963-64.
Table 2. Mallard post-season sex ratio ground counts by study unit, winter
of 1970-71.

Date and Location

No. Ducks Counted
Male
Female
Total

Percent
Males

292
293

214
232

506
525

57.7
55.8

135
216

111
114

246
330

54.9
65.5

313

267

580

54.0

120
157
142
304
332

56
95
87
214
176

176
252
229
518
508

68.2
62.3
62.0
58.9
65.4

141
161
178

119
139
122

260
300
300

54.2
53.7
59.3

Ft. Morgan-Sterling (Unit 2)
1-21-71
1-21-71

South Platte River
"
"
"

Ft. Collins Area (Unit 4)
1-28-71
1-28-71

College Lake
Lindenmaier Lake

Bonny Reservoir Area (Unit 9)
1-16-71

Bonny Reservoir

Arkansas Valley (Units 11 &amp; 12)
1-23-71
1-25-71
1-26-71
2-2-71
2-8-71

Two Buttes Reservoir

"
"

"

"
"

"

John Martin Reservoir

"

"

"

San Luis Valley (Unit 15)
2-19-71
2-19-71
2-19-71

Monte Vista NWR

"

"

"
"

"
"

�-198-

Winter Aerial Surveys
Periodic aerial surveys were again conducted to obtain data on numbers
of ducks present by area and time period. Another good year of reproduction was reflected in high numbers of birds censused, especially
in the November and December counts (Table 3). However, there was a
suggested decline through the winter period, the most drastic decrease
in numbers occurring from late December to early January. The high
count in 1969-70 was the January Inventory (January 5, 1970) when about
430,000 ducks were censused. In 1970-71, November and December counts
were comparable to that figure with 470,000 and 430,000 birds censused,
respectively, while the January Inventory was only 310,000, much below
that of the previous year. Weather, counting conditions and movement
of birds has to affect counts on any specific date so average figures
for the winter of 1970-71 have to be utilized to a certain degree when
comparing numbers of birds present in Colorado between years.

Table 3. Aerial duck counts by interval and study unit, eastern Colorado,
1970-71.

Area and Study Unit

Number Ducks Counted by Date
Nov. 23
Dec. 21

Jan. 5

(1)

Sterling-Julesburg

26,250

27,000

17,000

(2)

Ft. Morgan-Sterling

64,100

35,400

23,900

(3) Greeley-Ft. Morgan

43,000

70,700

59,000

(4)

Ft. Collins

88,400

56,850

9,950

(6) Denver-Greeley

57,000

50,230

59,525

(9)

Bonny Reservoir

57,300

51,930

48,000

(1/) Arkansas Valley

75,885

87,160

28,505

(15) San Luis Valley

57,170

52,200

64,060

Grand Totals

469,105

431,470

309,940

1/ Includes Units 10, 11, 12 and 13.

�-199-

Wing Surveys
Information on age and sex ratios in the harvest, as in previous years,
was gathered by The Bureau of Sport Fisheries and Wildlife wing survey.
These data are presented in Administrative Reports by The Bureau and
will be utilized in the final analysis and report in a future segment.
Hunting Pressure Surveys
Duck hunting pressure and harvest by unit was again estimated by unit
with data illustrated in Table 4. The hunting season was 90 days in
length, the longest since 1958, but estimated number of hunters (about
31,000 statewide) was not up to the total estimated for 1958 (almost
39,000). Bag limit was based on the point-system, allowing from 1 to
10 birds in the bag (Funk 1971). Species composition of the bag was
estimated at 73 percent mallards on an overall basis. Figures in Table
4, however, are total birds and not restricted to mallards.

Table 4. Hunter pressure, harvest and population survey data for comparison between management units and areas, 1970-71 duck season.

Area 1/

Estimated
Duck
Hunters ]:./

Estimated
Ducks
Bagged 2:..1

Average
Bag Per
Hunter

Number
Number Hunters
Ducks
Present Per 100
Censused 11 Ducks Censused

Northeast
Unit 1
Unit 2
Unit 9

1,817
2,036
846

15,712
14,305
6,332

8.6
7.0
7.5

27,000
35,400
51,930

6.7
5.8
1.6

Total

4,699

36,349

7.7

114,330

4.1

-------------------------------------------------------------------------------North Central
Unit 3
Unit 4
Unit 6
Total

4,980
3,446
5,545
13,971

31,425
17,589
42,916
91,930

6.3
5.1
7.7
6.6

70,700
56,850
50,230
177,780

7.0
6.1
11.0
7.9

-------------------------------------------------------------------------------Southeast
Und.t; 10
Unit 11
Unit 12
Unit 13

1,159
344
1,598
1,128

7,739
1,407
11,022
12,898

6.7
4.1
6.9
11.4

- il

Total

4,229

33,066

7.8

87,160

11 See Fig. 1 for management unit locations.

2:..1 Funk 1971.
11 Data from Table 3. December 21 count data utilized.

il Inventory data for Arkansas Valley usable only as total.

4.9

�-200-

In comparing data from the 1969 season (Funk 1970) with those in Table
4, hunting pressure in the northeast area was down somewhat in 1970 with
harvest about the same. Hunting pressure and harvest were up in 1970
in both the north-central and southeast areas. Average bag per hunter
was up somewhat in all areas.
As estimated by comparing numbers of hunters per each 100 birds censused,
hunting pressure was indicated to have been greatest in the Denver-Greeley
area (Unit 6) and lowest at Bonny Reservoir (Unit 9). Most other units
were similar in hunting pressure at between 6 and 7 hunters per 100 birds
censused, with the exception of the Arkansas Valley at about 5 hunters per
100 birds.
The latter, however, was an increase over the estimate for the
1969 season, indicating a continuing added increase in duck hunting
interest in the area.

Evaluation

of the Point-system

Duck Season

Colorado personnel cooperated in the hunter blind performance effort conducted in all states having the point-system season.
Data collected were
submitted to The Bureau of Sport Fisheries and Wildlife for analysis and
reporting.
Reaction of the hunters to the point-system regulation and
acceptance of it by them were again judged to be good, based on results of
the performance checks and comments by hunters on a harvest questionnaire.

Analysis

of Band Recovery

Data

All banding and recovery data on tapes from the initial 1963-64 banding
effort through the hunting season of 1969-70 were processed through the
various programs constructed for analysis.
Due to lack of time, examination
of the program results was not completed during the segment but will be
forthcoming in the next segment report, including an updating of information
through the 1970-71 season.

LITERATURE

CITED

Funk, H. D., J. R. Grieb, D. Witt, G. F. Wrakestraw, G. W. Merrill, J. Sands,
T. Kuck, D. Timm, T. Logan, and C. D. Stutzenbaker.
1971. Justification
of the Central Flyway High Plains Mallard Management Unit.
Cent. Fly.
Tech. Comm. Rept. March.
48 pp.
Funk, H. D. 1970. Waterfowl kill survey.
Colo. Div. of Game, Fish and
Parks Game Res. Rept. Oct. pp. 19-40.
1971. Waterfowl kill survey.
Game Res. Rept. Oct. In press.

Prepared

d{.

t:2J7

L

Colo. Div. of Game, Fish and Parks

by _~
__ ~~~~U~/~I~~,~~ __ L__._~
____~_~_,_._~
~ce
_
Howard D. Funk
Section Chief
Small Game Research

�-201October,

1971

JOB PROGRESS REPORT

State of
Project No.

COLORADO
~W_-~8~8_-~R~-~1~6
_

Work plan No.

4

Job Title

Trapping

Period Covered:

April

Migratory

Bird Investigations
3

Job No.
and Banding

Doves

1, 1970 through March 31, 1971

Personnel:
Charles Hayes, Dale Horne and Jack Randall, Bureau of Sport Fisheries and Wildlife; Clait Braun, William Carpenter, Howard Funk, Larry Green,
Gary Robinson, Michael Robinson, Steve Steinert and Al White, Colorado Division
of Game, Fish and Parks.

ABSTRACT

Efforts initiated in 1964 to trap and band samples of mourning doves (Zenaidura
macroura) in Colorado were continued in 1970. Cooperative Federal and State
efforts resulted in 2,883 birds being newly banded.
Of this total, 1,359 were
immatures; 821 were adult males while 703 were adult females.
Assigned quotas
were accomplished for the Eastern Slope of the Rocky Mountains, but not for western Colorado or extreme eastern Colorado.
Wing molt data from trapped and harvested immatures indicated three hatching peaks.
Peak hatching periods were
late May-early June, late June-early July and late July-early August.
A sample
of 229 hunter-killed birds in southeastern Colorado gave a young-to-old ratio
of 5.2:1, more than double the age ratio obtained in the same area and time
period in 1969. Wing molt data from harvest immatures in this sample indicated
that most early hatched young migrated from Colorado prior to September 1.
Fifty-three band returns were reported in 1970 from doves banded by Division
personnel.
Thirty-six (67.9 percent) of the bands reported were from doves killed
in Colorado.
Apparent hunting mortality rates for doves banded in Colorado continues to be less than 3 percent.

��-203-

MOURNING

DOVE TRAPPING

AND BANDING

Clait E. Braun

Intensive efforts to trap and band mourning doves in Colorado initiated
in 1964, were continued in 1970 in cooperation with personnel of the
U. S. Fish and Wildlife Service.
Data presented in this report are those
collected in 1970, the seventh year of this continuing investigation.

P. S. OBJECTIVE
To investigate migration patterns and mortality
banded in Colorado by age, sex, and area.

SEGMENT

rates of mourning

doves

OBJECTIVES

1.

To trap and band mourning doves in three selected areas of Colorado
for the purpose of obtaining migration, life history, and annual
mortality data.

2.

To estimate

harvest

size and hunter

METHODS

success.

AND MATERIALS

Methods and materials used in 1970 were similar to those described in
earlier reports and summarized by Braun (1970). In addition, less than
100 doves were trapped using cannon nets in 1970 during trapping operations
for band-tailed pigeons (Columba fasciata).

DESCRIPTION

OF TRAPPING

AREAS

Trapping sites in East Slope areas were similar to those described earlier
(Braun 1970). An additional East Slope site was intensively trapped in
1970. This area in the San Luis Valley was an abandoned cultivated field
formerly planted to field peas (Pisum sativum) and barley (Hordeum vulgare) •
Mourning doves were also trapped at an additional West Slope location in
1970. This site, near Durango, was located in an area dominated by small
cultivated fields of barley and alfalfa (Medicago sativa) interspersed with
pinon pine (Pinus edulis).

RESULTS AND DISCUSSION
Trapping
Cooperative

and Banding

Federal and State trapping and banding efforts resulted in
Of this total, 2,101 were banded by

2,883 doves being banded in 1970.

�-204-

Division personnel.
Doves were banded at six major sites on the West
Slope.
In addition, a few doves were banded at other scattered locations
throughout the mountainous areas of Colorado in conjunction with bandtailed pigeon trapping activities.
The banding goal of 4,000 birds was
not accomplished as no birds were banded in extreme eastern Colorado.
Only 527 of the 1,000 quota were banded in western Colorado.
Sex and age
distribution of birds banded by area are presented in Table 1.
As in earlier years of this study, more adult doves were trapped than
immature, although the disparity was not great in 1970. The small
difference between numbers of adults and immatures trapped is probably a
function of time of season that traps were operated.
W11en adult:immature
ratios are compared by t Lme period, large differences are apparent.
Trapping at Meeker was conducted in May and early June with mostly adults
being caught.
At the major Denver trapping site, traps were operated
from June into August.
Only 21.7 percent of the birds trapped at this
site in June were immatures, this percentage increased to 30.4 in July,
and 64.3 in August.
Thus, trapping conducted in May, June, and early
July results in high adult:young ratios, while trapping in late July and
August after most young are flying reverses ratios of birds in trap
samples.
Similar trends were noted in trap samples from Durango, Monte
Vista, and Fort Morgan.
Numbers of adult males and females trapped in
1970 were similar, unlike earlier years when substantially more males
than females were caught.

Hatching

Data

Wing molt data were available for 1,253 of the 1,359 immatures trapped and
banded in 1970. Estimated hatching dates (Allen 1963) are presented in
Table 2. Examination of data in Table 2 indicates that few mourning doves
trapped in Colorado hatched prior to May 1, with most hatching between
mid-May and mid-July.
Two definite peaks of hatching about 1 month apart
(late May-early June, and late June-early July) are apparent from data
presented in Table 2. While few of the doves trapped hatched prior to
May 1 or after July 23, progeny of late nesting doves are not represented
in Table 2 as trapping ceased on August 20. It should be recognized that
percentages of young hatching in each time period are not absolute as
trapping effort and success throughout the period was not uniform and that
distribution of adults and immatures throughout the State was also not
uniform.
Upon tabulating

the number

of immature

doves molting

p r'LmarLes 1 through

10, it was apparent that there was a constant turnover of young doves
available for trapping in the populations sampled.
Approximately 65
percent (65.5) of the immatures trapped had not molted primary 1 or were
molting primaries 1 or 2. In contrast, only 4 percent (3.6) of the young
trapped had molted primary 6 or beyond.
These data vary only slightly
from those collected in previous years of this investigation, again
suggesting that immature doves leave nesting areas within 30 to 40 days
of fledging.

�-205-

Table 1. Sex and age distribution of mourning doves banded by area in
Colorado, 1970.
Adult Male
Number
Banded Percent

Adult Female
Number
Banded Percent

Immature
Number
Banded Percent

Total

Denver l/

431

30.7

342

24.4

631

44.9

1,404

Honte Vista 'l:../

74

23.4

64

20.3

178

56.3

316

Vineland 1/

25

10.2

50

20.4

170

69.4

245

Fort Morgan

55

26.0

59

28.0

97

46.0

211

Fort Collins i/

51

32.9

21

13.6

83

53.5

155

Rocky Ford

1

4.0

o

0.0

24

96.0

25

Sub-total

637

27.0

536

22.8

1,183

50.2

2,356

Durango 2/

70

22.8

72

23.5

165

53.7

307

Meeker 2../

114

51.8

95

43.2

11

5.0

220

Sub-total

184

34.9

167

31. 7

176

33.4

527

Total All Areas

821

28.5

703

24.4

1,359

47.1

2,883

Area
Eastern Slope
Rocky Mountains

Western Slope
Rocky Mountains

1/ Includes 14 birds banded in the Evergreen-Aspen Park area.

2/ Includes 1 bird banded near Salida.
3/ Includes 1 bird banded near Rye.
4/ Includes 9 birds banded near Estes Park.
5/ Includes 1 bird banded in Sanborn Park.

i/ Includes 1 bird banded at Buford, 6 at Craig, and 4 at Lay.

�-206-

Harvest
No effort was made to collect wings from hunter-killed doves throughout
Colorado during the 1970 hunting season. Wings of 229 birds harvested
September 1-3 in southeastern Colorado were available for analysis. Of
this total, 37 were from adult doves while 192 were from immatures, giving
a young-to-o1d ratio of 5.2:1. This is approximately double the 2.4:1
ratio obtained from a sample of 172 birds harvested in this same area in
early September 1969 (Braun 1970). Approxi~~te1y 80 percent (81.2) of
the young harvested had molted five or fewer primaries, indicating they
were less than 63 days of age. This supports data collected in previous
years of the study which indicated that most early hatched young (May-June)
migrated from Colorado prior to September 1. Molt data from harvested
birds also indicate that substantial production occurs after July 23
(Table 2) as over 30 percent (33.3) of the birds shot between September 1-3
hatched after July 23. These young possibly represented the progeny of a
third nesting attempt.
Harvest data obtained through use of a mail survey of hunters after the
hunting season indicated that 24,149 hunters havested 301,562 doves in
Colorado in 1970 (Funk and Tully 1971). Fifty-three band returns were
received from doves shot in 1970 that were banded by Division personnel.
One of these birds had been initially banded in 1967, 4 were banded in
1968. 20 in 1969, while 28 had been banded in 1970. Apparent hunting
mortality rate continues to be less than 3 percent. Of the 53 bands
reported, 36, (67.9 percent) were recovered in Colorado, 10 in Mexico,
4 in New Mexico and 1 each in Texas, Arizona, and Oklahoma. All but four
were shot recoveries.

LITERATURE CITED
Allen, J. M. 1963. Primary feather molt rate of wild immature doves in
Indiana. Ind. Dept. Conserv., Game Res. Sect. Circ. No 4, Indianapolis. 4 p.
Braun, C. E. 1970. Mourning dove trapping and banding. Colo. Div.
Game, Fish, and Parks, Game Res. Rept., Fed. Aid Proj. W-88-R.
Oct. pp 143-149.
Funk, H. D., and R. J. Tully. 1971. Colorado small game hunter harvest
survey, 1970. Colo. Div. Game, Fish and Parks, Fort Collins.
:t-tlmeo
Rept. In prep.

Prepared by

~rU.::::'
cPo· /?

C/~

t/,~

,

--------------~-------------------Clait E. Braun
0

Assistant Wildlife Researcher

�Table 2.

Estimated hatching dates for wild-trapped mourning doves, 1970.
I

Number
hatching

N

Prior to
Hay 1

1-7

8-14

15-21

22-28

29-June 4

5-11

12-18

June
19-25

26-Ju1y 2

3-9

10-16

Ju1
17-23

24-30

3

5

14

77

124

178

86

98

137

187

191

93

59

1

Ha

0

~
I

��-209October, 1971
JOB PROGRESS REPORT
State of

COLORADO
--------~~~~~-------

Project No.

W-88-R-16

Migratory Bird Investigations

Work Plan No.

4

Job Title

Band-tailed Pigeon Investigations

Period Covered:

Job No.

4

April 1, 1970 through March 31, 1971

Personnel: Jack Randall, Bureau of Sport Fisheries and Wildlife; personnel
of the U.S. Forest Service; Foster M. Barker~ Gene W. Bassett, Clait E.
Braun, William Carpenter. Arch Cowan, Velma Fredrickson, Howard D. Funk, Dan
Gravestock, Charles Hector, Richard Hopper, Wayne E. Kniselyp Wilbur Ladd,
John Leslie, Richard McDonald, Charles Reichert, Nathan Riggs, Charles Roberts,
Michael Robinson, Walter Schuett, Michael Szymczak, Marie Vendeville, Kenneth
Wagner, J. Allen White and other personnel of the Colorado Division of Game,
Fish and Parks.

ABSTRACT
Investigations of band-tailed pigeons (Columba fasciata) initiated in Colorado
in 1969 in conjunction with the Four Corners States Cooperative Study (Arizona,
Colorado, New Mexico, Utah) were continued in 1970. Major emphasis was placed
on delineating distribution, ascertaining relative abundance, trapping and banding samples of pigeons throughout the State and documenting breeding and nesting phenology. In 1970, band-tails occurred throughout the forested mountains
of Colorado from late April into December.
Changes in abundance from 1969
were not detected. Apparent densities were highest in southern and southwestern
Colorado. Pigeons were trapped at 13 different locations in 1970 with 3,292
bi~d~ (2,744 adults and 548 immatures) being newly banded. Thirty-four birds
were retrapped outside of their original banding area, while 34 birds banded in
1969 returned to their initial banding locations. During the experimental hunting season~ 364 permits were issued with 182 individuals actually hunting. Only
77 hunters were successful, each harvesting an average of 5.9 pigeons. Total
estimated harvest (including crippling loss) projected from the questionnaire
survey (92.3% response) was 541 birds. Crippling loss approximated 15 percent
of the birds shot and retrieved. Wings were received from 68 successful hunters.
Immature and subadult pigeons comprised 32.2 percent of the 394 wings received.
Field checks of 144 (89 adults, 55 immatures) hunter killed birds indicated
that 52.3 percent of the adults were still involved with nest activities. Recoveries were received from 34 pigeons banded in Colorado. Twenty-six were from
birds recovered in Colorado, 6 from Mexico and one each from Utah and New Mexico.
At least 6.3 percent of the birds harvested during the experimental season in
Colorado were banded. Data from collected and hunter-killed pigeons indicated
that band-tails nested in Colorado from early May to mid-September in 1970. Calculated hatching dates for 658 immature pigeons ranged from mid-April to midAugust with a major peak in mid-June to early July. Helminth infections continue
to be low (10.4%) but a survey of blood parasites revealed high levels in both
adults (93.6%) and immatures (73.0%).

��-211-

BAND-TAILED PIGEON INVESTIGATIONS
Clait E. Braun

Intensive ecological studies of wild band-tailed pigeons which nest and
reside in Colorado from late April to early November, initiated in 1969,
continued in 1970. Investigations in Colorado represent a portion of the
regionwide Four Corners Cooperative Band-tailed Pigeon Investigation which
was initiated in Arizona in 1967.
.

P. S. OBJECTIVE
To investigate life history and ecological facts pertinent to band-tailed
pigeons in Colorado as follows: range; relative numbers; relation between
numbers of birds present by area and such physical factors as weather,
food, vegetative cover, elevation, and salt; breeding and nesting
chronology; migration patterns and chronology; harvest patterns; and
mortality rates.

SEGMENT OBJECTIVES
1.

To investigate distribution and relative numbers of band-tailed
pigeons in Colorado by area.

2.

To investigate the correlation between abundance and distribution of
band-tailed pigeons aridphysical features of the environment.

3.

To investigate the breeding and nesting chronology of bandtails in
Colorado.

4.

To trap, band, age, and sex as many birds as possible in selected
areas of Colorado to investigate migration patterns and chronology
of Colorado bandtails throughout their range.

5.

To conduct an experimental hunting season, gather harvest data, and
evaluate results.

METHODS AND MATERIALS
Most methods and materials used in 1970 were similar to those used in
1969 and described by Braun (1970). Reporting procedures were slightly
modified in 1970 as all flocks of 50 or more pigeons were to be reported
directly by telephone to the Game Research Center of Fort Collins. In
addition, all monthly reports of field personnel in the Southwest Region
were sent directly to the Regional Biologist who summarized them and forwarded the summary sheets to Fort Collins. This procedure worked fairly well.

�-212-

In addition to the one 30- by 75-foot cannon net with l~-inch mesh used
in 1969, three 30- by 60-foot three-cannon nets with l~-inch mesh and
24-inch "skirts" were utilized in 1970. These nets were more efficient
than .the large net as they were lighter, projected faster, and had smaller
mesh which reducedentaqg1ementof
the birds. Cloacal inspection for
determination of sex was discontinued in 1970 and all adult pigeons trapped
were sexed using external plumage characters described and tested by
Braun (1970). In adaition to the three major grains used to bait pigeons
for trapping in 1969
(barl§!y (Hordeum vulgare), wheat (Triticum aestivum),
and mi10 (Sorghum vulgare», large numbers of pigeons were trapped in 1970
utilizing field peas (Pisurilsat! vum) and corn (~
mays) ~
In order to evaluate the experimental hunting season in 1970, all hunters
were required to obtain a free individually numbered permit. In addition,
each hunter was issued a map of the area open to hunting and a packet
containing five numbered wing envelopes. Following the experimental
season; iil1hunters were surveyed by questionnaire concerning their hunting
activities. One fo11owup letter was sent to all nonrespondents to the
initia11etter.
Copies of all materials used for the experimental hunting
season. are attached (Appendix).
Pigeons collected or killed in trapping operations were processed as in
1969 (Braun 1970), except that all crop gland and gonadal material was
preserved in Bouin's Fixative.

DESCRIPTION OF TRAPPING AREAS
Pigeons were trapped at locations near Dolores, Stoner, Bayfield, Durango,
Del Norte,· Buford', Evergreen, Sanborn Park, Longmont, Salida, Rye, Monte
Vista, and Estes Park in 1970. Trapping sites were primarily in fields in
which barley (Dolores, Durango, Del Norte-Davie's, Buford, and Salida),
wheat (Bayfield, Sanborn Park, and Rye), or field peas (Del No rt.e-Bauer 's
and Monte Vista) had been planted in 1969. These sites were characterized
by having an abundance of waata grain scattered on the ground. The trapping
site at Longmont consisted of a trench silo where rolled corn with molasses
was stored. Areas where pigeons were trapped in the vicinity of EvergreenConifer-Bergen Park and Estes Park were those where residents of these
communities put out grain for pigeons and other birds. The trap site at
Stoner was located at the bottom of a ski tow. Barley was spread in this
area in order to attract pigeons as the previous owner fed birds at this
site in the early 1960's. All trap sites were adjacent to areas which had
trees suitable for perching. No preference for tree type was observed as
pigeons commonly roosted in Ponderosa pine (Pinus ponderosa), cottonwoods
(Populus spp.), Gambe1's oak (Quercus gambe1i), Engelmann spruce (Picea
engelmannii), Douglas fir (Psuedotseuga menziezii), and aspen (Populus
tremuloides) • At Monte Vista and Buford, pigeons commonly would sit on
power lines before feeding on the baited sites. Areas preferred for feeding
were not in crop production in 1970 although most had been cultivated in
1969.

�-213-

RESUI,TS AND DISCUSSION
Distribution and Abundance
The distribution of band-tailed pigeons in Colorado in 1970 (Fig. 1) was
prepared from reported observations of field and project personnel.
Differences in distribution between 1969 (Braun 1970) and 1970 were slight
as no real changes were detected. As in 1969, pigeons were found to be
widely but unevenly distributed throughout the forested mountainous areas
of the State. They continue to be absent (not observed) in extreme northwestern Colorado and in the North Park, South Park, and Gunnison Basin
regions.
In 1970,·399 different observations of pigeons were reported from April
into December. Number of reported sightings by month were: April - 6;
May - 85; June- 97; July - 72; August - 69; September - 41; October - 17;
November - 11; and December - 1. These data indicate that in 1970
pigeons started arriving in Colorado in late April and started migrating
south in late August, with the peak of southward migration occurring in
September. Only scattered observations of smal L groups of pigeons were
reported in October, November, and .December.
Data concerning observations of bandtail occurrence by habitat type were
available for 363 different sightings. Of these sLghtLngs , 94 (25.9%)
were aaaocd at.edwith cropland, primarily grainfields. Eighty-nine sightitfgs
(24.5%) were in areas dominated by oak, 65 (17.9%) were from along stream
courses dominated primarily by cottonwoods, 59 (16.3%) were in mixed
coniferous areas (primarily spruce-fir-aspen), 34 (9.4%) were in areas
dominated primarily by ponderosa pine, 21 (5.8%) were in pinyon-juniper
(Pinus edulis-Juniperus spp.) areas, while only one observation was from
an alpine area. Observations are undoubtedly influenced by access and
distribution of observers, but reported observations probably give an
adequate picture of those areas in which pigeons feed. Few data are
available concerning abundance of pigeons in Colorado. From observer
reports, it would appear that numbers in 1970 were similar to those in
1969.
Abundance of pigeons continues to be highest in areas where mast- ,.
producing trees and shrubs are abundant. Thus, densities appear to be
highest in southern Colorado and lowest north of a line from Colorado
Springs to Grand Junction. This is not to imply that substantial numbers
of birds do not occur north of this line. Possibly the concentrations of
pigeons near Evergreen and Longmont are fairly recent in origin and cof.n-«cide with the urbanization of foothill areas (Evergreen) and the increase
in livestock feeding (Longmont).
Banding
Banding and trapping activities were intensified in 1970 with pigeons
being trapped in 13 different areas, including all five of the 1969
trap sites. As a result of these efforts, 3,292 pigeons were newly
banded (Table 1).

�COLORADO
MOFFAT

/fOU"

WUI)

S£O(;WIClf

LOGAN

.PHILLIPS

MORGAN
WASHINGTON

I rUMA

RIO 8LANCO

GARF/CLIJ
lOT GAIISON

LINCOLN
MESA

I

N'
f-I

~
I

CHUlNNE

1:L'3.

ICROWLEY

I..

IKIOWA

SAGUACHE
SENT

I.A.S ANIMAS

Figure 1.

Band-tailed pigeon distribution, 1970.

IPROWERS

I SACA

�Table 1.

Number

Area
Dolores

Bayfield

Durango
Stoner

0f

b and-tailed pigeons banded by area in Colorado, 1970.

Dates

Adults
No.
Banded
Percent

Immatures
No.
Banded
Percent

-

Totals

Estimated No. of
Birds Using
Trapping Sites

198
93

300+
200+

448
50

1,000+
200+

May 11-17
June 12

198
91

100.0
97.8

-

May 18-31
June 1-11

448
49

100.0
98.0

-

-

1

2.0

May 18-29
June 1-15

137
196.

100.0
97.5

5

2.5

137
200

300+
300+

June 16-24
July 7

85
16

100.0
100.0

-

-

85
16

100i50+

2

2.2

-

I

~
....
VI
I

Del Norte

June 16-18

168

98.8

2

1.2

170

200+

Buford

June 22

58

100.0

-

-

58

.200+

Evergreen

June 27-30
July 1-17

66
237

97.1
90.5

2
25

2.9
9.5

68
262

100+
300+

Sanborn Park

July 5-6

116

96.7

4

3.3

120

300+

Longmont

July 8-12
August 5

298
3

92.8
6.7

23
42

7.2
93.3

321
45

400+
60+

Salida

July 10-11

150

94.9

8

5.1

158

250+

Rye

July 13-19

88

89.8

10

10.2

98

1.50+

Monte Vista

July 27-30
August 1-30

149
181

48.9
40.7

156
264

51.1
59.3

305
445

700+
1,000+

Estes Park

August 7

10

71.4

4

28.6

14

50+

2,744

83.4

548

16.6

3,292

5,86Ot

�-216-

Upon examination of data presented in Table 1, it is apparent that number
of immatures trapped increased from late June to late August. Only 12
immatures were trapped in June, 226 in July, while 310 were trapped in
August. As in 1969, trap data suggest that some pigeons in Colorado
hatched in late May-early June, but that peak of fledging occurred in midJuly and early August.
Sex of most adult pigeons handled in 1970 was determined from external
plumage characters. Of the 2,734 adults newly banded for which sex was
determined, 65 percent (1,776) were males and 35 percent (958) were
females. Resasons for this disparity are not adequately known, but may
be related to time of day that trapping was conducted.
A total of 295 previously banded pigeons were retrapped in 1970. Fiftytwo of these had been banded in Colorado in 1969 , five had been banded by
private banders in Colorado as early as 1965 (one) and 1967 (four), while
one had been banded in Utah. The remaining 237 were banded in Colorado
in 1970 and then retrapped. Thirty-four birds were retrapped outside the
original banding area (foreign returns), while 34 banded in 1969 returned
to their initial banding location (returns).
Experimental Hunting Season
The first hunting season for band-tailed pigeons in Colorado since 1944
was conducted in 1970 as part of a 3-year experimental harvest program.
Hunting was allowed from September 12 through September 20, with a daily
bag limit of five birds and a possession limit of 10. During the experimental season only the southern portion of the State was open (Appendix).
Hunting was by free permit obtainable at five Division offices with no
limit on the number of permits issued. In 1970, only 364 permits were
issued.
Hunter Questionnaire Survey
The experimental hunting season for bandtails closed at sunset on September
20. On September 21, letters containing a self-addressed postage paid
return card were sent to all permittees (Appendix). Only one (.3%)
questionnaire was returned for lack of sufficient address. On October 5
a followup letter was sent to the 93 permittees (excluding the one
permittee for whom we had insufficient address) who had failed to respond
to the first letter. Results of the questionnaire survey are presented
in Table 2. Data for the 28 non-respondents were calculated using
average values of the respondents to the followup survey.
It is apparent from data presented in Table 2 that the initial experimental season for band-tailed pigeons in Colorado was not eve rly successful
in attracting hunters or productive for those who obtained permits. Only
182 (50%) of those with permits went hunting, and only 77 (42.3%) of
these hunters actually bagged one or more pigeons. However, those hunters
who were successful harvested about six birds (5.9) each. Total harvest
including those pigeons crippled and lost was calculated to be 541 birds,
substantially less than originally expected. The reported crippling loss
of 15.3 percent is not felt to be excessive but is deplorable.

�Table 2.

Experimental band-tailed pigeon season, harvest statistics, 1970. 1/
Response
First
Letter

Response
Second
Letter

Number of permittees responding

270 (74.2)

66 (18.1)

336 (92.3)

Number of permittees hunting

149 (55.2)

23 (34.8)

172 (51.2)

182 (50.0)

121 (44.8)

43 (65.2)

164 (48.8)

182 (50.0)

63 (42.3)

10 (43.5)

73 (42.4)

77 (42.3)

Number of permittees not hunting

Sum
First and
Second Letters

Projected
All (364) Permittees

",

Number of successful hunters
Number of hunter days
Days per hunter
Number of pigeons bagged

I

312

43

2.1
363

355

1.9
66

374

429

2.1
458

5.8

6).6

5.9

5.9

Pigeons per hunter

2.4

2.9

2.5

2.5

Number of pigeons crippled and lost

74

6

80

83

.5

.3

e .

5

.•5

Total harvest (bagged + crippled and lost)

437

72

509

541

Percent crippling loss

16.9

8.3

15.7

15.3

1/ Values in parentheses are percentages.

"
I

2.1

Pigeons per successful hunter

Pigeons crippled and lost per hunter

.1'.)
t-'

�-218-

The questionnaire was devised in order to obtain LnformatLon concerning
date of harvest. It was found that 36.8 percent of the total harvest
occurred on the opening weekend, 30.7 percent during the next 5 days,
while 32.5 percent occurred on the second and last weekend. These data
suggest that a longer season would result in a larger harvest.
Concentrations of hunters did not occur in 1970, and hunters were widely
scattered in the area open. Of 168 hunters reporting their hunting
locations 39 (23.2%) had hunted near Durango and 23 (13.7%) had hunted on
Bear Creek near Salida. Of the other 28 hunting areas listed, none had
been hunted by 10 percent or more of the hunters. On a regional basis, 91
of 168 hunters (54.1%) hunted in Eastern Slope areas, while 77 hunters
reporting hunting west of the Continental Divide. Forty-seven of 91 East
Slope hunters (50.6%) were successful, while only 35 percent of the West
Slope hunters reported killing at least one bird. Reasons for this
disparity are not known but were certainly not related to pigeon abundance.
Many written comments were received on questionnaires with many permittees
(20) reporting that the season was too late and that.the pigeons had
migrated south before the season. This was partially true in 1970 as cool
wet we.ather in late August and the opening weekend of the season apparently
affected distribution and number of pigeons in hunting areas.
Results from the 1970 Small Game Hunter Survey (Funk and Tully 1971) were
compared with those obtained from the special questionnaire survey of
pigeon hunters. Projected data from the 1970 Small Game Survey indi~ate
that 1,041 pigeon hunters havested 5,063 pigeons. However, only 364
hunters actually had pigeon permits (all pigeon hunters were required to
have permits), and these hunters killed a total of 541 birds. The difference between the two surveys is highly significant and suggests that either
(1) many hunters hunting pigeons did not have permits, or (2) hunters
responding to the Small Game Survey reported hunting and killing pigeons
when they actually did not, or both. Data from the small game survey are
based on projections from 54 hunters who actually reported hunting pigeons.
Names and addresses of 20 of these hunters were available for direct
comparison with the special survey of pigeon hunters. Only two of these
20 hunters actually had permits for pigeon hunting. Both of these hunters
reported harvesting the same number of birds on both surveys. Examination
of the addresses of the other 18 hunters who reported hunting pigeons
revealed that one-half of them lived in areas far removed from the pigeon
hunting area. It is believed that most or all of these people did not
hunt pigeons and that possibly they confused pigeons with mourning doves
(Zenaidura macroura) or rock doves (Columba livia) in reporting.
Wing Survey
Packets containing five wing envelopes were issued to each pigeon permit
holder in 1970. Envelopes containing 394 wings (excluding three rock dove
wings) were received from 68 hunters. Additional data were received
concerning hours hunted (343), birds crippled and lost (65), location of
kill, and banded birds harvested (25).

�-219-

Major harvest areas were Bear Creek near Salida (26.1%), Durango (18.7%),
Wetmore-Florence (13.7%), and Dolores (12.2%). These data compare favorably
with those from the questionnaire
survey, but suggest that hunters near
Salida were more successful than those hunting near Durango. Also, percent
harvest at Wetmore-Florence and Dolores was higher than the percent of
hunters hunting in these' areas, suggesting that hunting was good at these
sites.
Crippling loss calculated from the wing survey was slightly
lower
(14 •.2%as compared to 15.3%) than that calculated
from the questionnaire
survey.
Differences were not signifieant
though.'
.
Twenty-five of the 394 band-tailed
pLgeons reported harvested on the wing
survey were, banded (6.3%). Percent of birds harvested that were banded.
varied from 12.5 at Dolores,12.2
at Durango, 9.7 at Sal.Lda, to zero for all
other hunting areas •. Substantial
numbers (Table 1) of pigeons had been
banded at other locations but no recoveries were reported.
Comparable data were available from both hunt e r questionnaire
and wing
surveys concerning birds bagged and birds crippled and lost by 65 individual hunters.
A reporting difference of 3.6 percent (379 in wing surveys
versus 392 in questionnaire
survey) was found in number of birds killed,
and 8.8 percent (62 versus 68) in numb.er of birds crippled and lost 'for
those hunters who sent in wings and also. responded to the questionnaire.
These differences
are slight,
but do suggest that the questionnaire
survey
gives slightly
higher values for total birds bagged and birds crippled and
lost than actually occurred.
All wings received were measured (carpal and all 10 primaries) and
examined in order to obtain age (immature, subadult, and adult) structure
of the harvest.
Wings were re ceLved from 267 adults, (67.8%), 22 definite
sub-adults (5.6%), and 105 inuna.tures (26.6%). SubadUlts were those
molting juvenile primaries 5 through 10, while immatures were those molting
juvenile primaries 1 through 4 or had not molted primary number 1. The
percent of immatures (birds in their first year of life)
in the harvest
(32.2 when subadults are added) is higher than' that reported in other
studies and is probably a reflection
of better aging techniques rather than
higher production.
. However, from all appearances , reproductive success
and production were high in Colorado in 1970. Caution should be exercised
in using wing surveys to estimate production because several factors may
be involved such as differential
migration and vulnerability.

Hunter Field Checks
During the 1970 experimental season, emphasis was placed in contacting
pigeon hunters in the field in order to examine harvested birds for crop
gland activity..
By examining adequate samples of birds shot during hunting
seasons, the percentage of birds still
involved with rearing of young can
be calculated.
The premise is that season dates in succeeding years can
be adj usted accordingly.

�-220-

Crops of all adult pigeons checked were classified as being active (crop
gla~ds with curds), stimulated (gland apparent but no curds), or inactive.
In addition, sex by gonadal inspection was obtained for as many birds as
possible. A total of 144 birds were field checked.
Eighty-nine adults (61.8%) and 55 immatures (38.2%) were field checked,
with 46 (52.3%) of the adults having either active (28.5%) or stimulated
(22.~7%) crops. Sex of 84 adults was determined with 40 (54.l%) being
males and 34 (45.9%) females. While sample sizes are sm~ll, a higher
percentage of males (65.0) had active or stimulated crops than did females
(47.6). Comparisons between areas could not be made because of inadequate
~amples, but no apparent large differences were found~
Mortality
Recoveries of 34 band-tailed pigeons initially trapped and banded in Colorado were reported in 1970. Twenty-two of these were initially banded in
1970, while 12 were initially banded in 1969. While numbers of recoveries
is inadequate for detailed analyses some general observations can be made.
Twenty-four of 'the recoveries were from the 1970 experimental pigeon
season in Colorado and all were taken within 25 air miles of original
banding location. Two recoveries were from other states {Utah and New
Mexico) during their 1970 experimental seasons. One of these birds was
banded at Dolores and was shot near Mason Spring, Utah, less than 50 air
miles west of the banding location. The other bird was banded near Bergen
Park (west of Denver) and was shot in Chloride Canyon near Truth or
Consequences, New Mexico. Both birds were recovered the same year banded.
Six recoveries were from Mexico with,S being shot recoveries, while cause
of death of the other was unknown. These birds were recovered in the
Mexican atates of Chihuahua (3) and"Durango (3). Recovery of the remaining
two birds was in Colorado with one being a trap casualty, while the other
was found dead where it had flown into a stationary object. One pigeon,
banded as an adult near Milwaukie, Oregon on 20 April, 1968 was shot by a
hunter 15 miles north of Durango, Colorado on 20 September, 1970. This
recovery is the first documented movement between the Coastal· and Interior
populations of band-tailed pigeons.
Based on the few recoveries to date, it appears that known mor~ality is
concentrated in breeding areas (Colorado) and wintering areas (Mexico)
with little mortality occurring during migration., At present, causes and
magnitude of natural mortality are unknown. It also:appears that bandtailed pigeons, which are summer residents in Colorado, have a great
~idelity to'specific areas. as shot recoveries away from general banding
areas were not reported in 1970.

Breeding Phenology
pata on breeding phenology were obtained through crop and gonadal inspection of birds accidentally lost in trapping operations and those systematically collected. Additional data were collected from 89 hunter-killed

�-221-

adult pigeons examined from September 12 through 20. Excluding the 89
hunter-killed birds, 135 birds were available for detailed examintion.
Of this number, 118 were adults (62 males and 56 females), while the
remaining l7.were immatures. The sample included birds taken from 10 May
through 9 October with the following monthly totals: May - 29, June - 28,
July - 39, August - 27, September - 11, and October - 1.
As in 1969, data in crop activity and gonadal condition were tabulated
but no analyses were. conducted. These data will be utilized by a.graduate
student and will be compiled and analyzed in a M. S. thesis at Colorado
State University..Cursory examination, of the gonadal data indicates that
male and female pigeons were in breeding condition (enlarged testes and
egg follicles) from early May until mid-August. Testes of some males
began regression by 25 August but two collected females still had enlarged
follicles (up to 5.0 mm) on the same date. Crop gland development in
collected birds was observed from 12 June to 20 September. Data collected
during the hunting season indicated that about 50 percent (52.3) of the
adult pigeons (46 of 89) were about to feed young, still feeding young or
had recently been feedingyotmg.
Thus, nesting of bandtai1s in Colorado
in 1970 occurred. from early May to mf.d-Bept.embe
r , It is probable that
many. pigeons had two nesting attempts in this period. Success of these
attempts is not known but wing survey data suggests that production was
good. Actual b reedfng age of w.ild bandtai1s has not been reported.
Examination of collected birds thought to be subadul.ts (with at least
juvenile primaries 10 and 9) revealed that all were either approaching
breeding condition or had successfu1y bred (evidence of crop gland development). It is certain that most, if not all, subadult pigeons are capable
of breeding in'their first year of life (second calendar year) •
Wing Measurement and Molt Data
Data concerning length of primaries, rectrices, total body 1ength,carpa1
length and bill length were obtained from all pigeons collected in 1970.
Once adequate samples are available, these data will be analyzed to examine
differences between age classes of both male and female pigeons. It
appears that sex of pigeons will beie1iab1y determinable from measurements of wing characters.
Prior to dissection of collected birds, plumage characters of each adult
specimen were examined and recorded. Each bird was then classified as
either male or female. Upon dissection, gonads were examined and actual
sex of each bird was compared with the classification based on plumage
appearance. Comparative data were available for 117 adults (62 males and
55 females). Of the 62 adult males compared, one (1.6%) was classified
incorrectly, while three (5.5%) of 55 females were classified as males
from plumage examination. Total error in classifying pigeons to sex from
plumage examination alone was 3.4 percent. These percentages are similar
to those reported by Braun (1970) for a sample of 119 adult band-tailed
pigeons and illustrate the accuracy with which an experienced observer
can sex pigeons using plumage alone.

�-222-

Primary molt of most pigeons examined in 1970 was systematically recorded.
Molt of adult pigeons is irregular and does not connnence in most birds
until late July or early August. Most birds molting primaries in May and
June are subadults. While the molt of subadults appears to be sequential
until juvenile primaries 9 or 10 are molted, molt of adults is n9t.
It is not uncommon to find adult pigeons molting two or even three primaries
at one time in no apparent sequence. All data on molt patterns, timing and
race will be incorporated in a M. S. thesis. at Colorado State University
and will not be further discussed at this time.
Information concerning primary molt was available for 658 Lmmatures (545
banded and released, 16 collected and 97 from the hunter wing survey).
Of the 658 innnatures, 293 had not shed primary 1, 204 were molting PI, 87
were moltingP2, 38 were molting P3; 18 were molting P4, while 18 were
moltingP5 or 6. Hatching dates for these birds were estimated using
unpublished data collected by H. M. Wight and his students (Personal
conununication, H. M. Wight). Number of days used to backdate for each
primary class were: 0 = 36, 1 = 43, 2 = 54, 3 = 72, 4 = 81, 5 = 96,
6 ""121. Estimated hatching dates for all immatures for.which data are
available are presented in Table 3.

Table 3.

Estimated hatching dates for wild band-tailed pigeons, 1970 •.

Time Period,

Number Hatching

Time Period

Number Hatching

April 26 &lt;

2

June 22-28

182

April 27-May 3

2

June 29-July 5

115

May 4-10

8

July 6..;.12

33

May 11-17

2

July 13-19

5

May 18-24

8

July 20-26

8

May 25-31

23

July 27-August 2

9

June 1-7

59

August 3-9

20

June 8-14

38

August 10-16

16

June 15-21

128

August 17 &gt;'

0

�-223-

As in 1969 (Braun 1970) hatqhing and fledging of band-tailed pigeons was
spread over a wide time period (April 13 to August 15). One obvious peak
occurred in 1970 between June 15 and July 5. If bandtai1s were consistently
attempting to nest twice, more than one hatching peak would be expected.
Multiple broods could occur and not be recognized as trapping operations
normally cease in mid-August when pigeons disperse to feed on native foods.
Thus, progeny of second nesting attempts would not be available for trapping.
However, wings from progeny of late nests could be expected in the hunt.er
wing survey. This did 'occur in 1970 as 45 of 97 (46.4%) of the wings
received from immature pigeons during the September hunting season were
from birds hatching after .JttlY27. The failure to receive an even higher
percentage of late hat chfng birds in the September harvest could be'
partially related to differential migration. This reasoning is not
presently accepted as data collected in 1969 and 1970 suggest that immatures
from early nestings and adults comp1~ting their nesting activities in late
August are the first to migrate south. All immatures calculated to have
hatched in April and May were handled in trapping operations in June and
.July. While numbers up to May 25-31 are small; they are puzzling. This
is especially true, for the four.b Lrds calculated to have hatched in April.
Since present data indicate that bandtai1s do not arrive in Colorado until
mid- to late April, these birds cannot reasonably be, assumed to be the
result of nesting at.tempts in Colorado. At the present time, delayed molt
of immatures has not been conclusively proven. Thus, young calculated to
have hatched in April and early May undoub t.edLy are the progeny of pigeons
nesting south of Colorado in late winter. Obviously, nesting in wintedng
areas is not commontm.1ess progeny of these nests have a high mortality
or fail to migrate north in their first year of life.
During trapping operations, 33 individual immature pigeons which had
molted at least one primary since initial capture were retrapped. Data
from these birds' indicated that about 11 days are required between molting
of individual primaries, 0 !:o1, 1 to 2, and 2 to 3. However, length of
time reqt,rl.red
by individual birds was quite variable. This was expected
as some birds were undoubtedly about to replace a primary when initially
trapped while others had probably just molted prior to initial capture.
Parasitic Load
Information on helminths was available for 133 band-tailed pigeons (116
adults and 17 immatures) necropsied. Fourteen (13 adults, 1 immature) of
the 133 had helminth infections. The infection rate of 10.4 percent is
low and only one bird (immature female) was considered to be heavily
infested. Six of 55 adult females (14.5%) harbored helminths, while only
5 of 61 (,8.3%)adult males were infected. These data are extremely
similar to that reported for a sample of 144 bandtai1s examined by BralUl
(1970). Tapeworms were found in 12 birds (4 males, 8 females) while
nematodes were found in only two birds (one adult male, one immature female).
Worms were found in pigeons collected as early as 18 May and as late as 9
October. No deleterious effects of the observed parasitism were noted.
All worms were preserved for later identification.

�-224-

Blood films were taken from 346 pigeons handled in 1970 representing 201
adults (111 males and 90 females) and 145 immatures with samples from May
through September. Bone marrow of an additional 18 birds (unclassified)
was also examined. All examinations identification of parasites were by
Dr. R. M. Stabler of Colorado College. Four protozoan parasites,
Leucocytozoon marchouxi, Haemoproteus columbae, ~. sacharovi, and
Trypanosoma' avium and the mfcrofd.Lardae of an unidentified nematode were
reported. Of the 201 adults examined, 93.6 percent were positive for at
least one parasite, while only 73 percent of the immatures were infected.
Leucocytozoon marchouxi and Haemoproteus sacharovi were most commonly
found in both adults and immatures (66 and 67%; 64 and 68%) with less than
five percent of all birds having microfilariae in the blood. No differences
in parasitic load were found between adults and subadults(reported
as
adults) and males and females. It is not surprising that immatures were
less heavily infected than adults. The actual effect that blood parasites
have on most birds is unknown, but it has been well documented that young
of some birds (especially waterfowl) heavily infested with hematozoa
suffer extremely high (up to 100%) losses. The finding of Trypanosoma
avium in band+t ad.Led pigeons constitutes a new host record.

LITERATURE CITED
Braun, C. E. 1970. Band+t.af.Ledpigeon investigations. Colo. Div. Game,
Fish and Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct.
pp 151-171.'
Funk, H. D., and R. J. Tully. 1971. Colorado small game hunter harvest
survey, 1970. Colo. Div. Game, Fish and Parks, Ft. Collins.
Mimeo Rept. In prep.

Prepared by _-.,..,"¥::~:l£.s.&lt;:r;JI::i:::::"'---,,:.r:~. _. ~~....L.'!:::k::..:;.::....~_.
Clait E. Braun
Assistant Wildlife Researcher

_

�-225-

APPENDIX

��-227-

INSTRUCTIONS FOR ISSUING EXPERIMENTAL
BAND-tAILED PIGEON HUNTING SEASON PERMITS

Hunters may obtain a special permit for hunting band-tails by appearing
in person, or by submitting a request by phone or mail. While we have a
place for the hunting license number, I see no reason why a person without
his license in his possession carmot also receive a permit. This means
persons can also pick up permits for dthers if he supplies complete names
and addresses. If you can get the license number, even over the phone,
get it. Permits may be issued from the time you receive them, throughout
the hunting season, which extends from September 12-20, ·1970. Please
follow the instructions below carefully.
1.

The permits are numbered, so please issue them in sequence.

2.

The permits are in duplicate. Please type them when at all possible,
or if not, write legibly and with enough force so the carbon copy is
clear.

3.

The original or top card of the permit is the State Reference Card
and is to be retained by us for our files. Please file them in order
of issue.· The carbon or bottom card goes to the hunter and is to be
carried on his person.while hunting.
There is a space for his
signature on the back of the permit.

4.

Disregard the line for wing collection envelope packet number until
you receive further instructions. We don't have any idea as to how
many applicants we will have this initial year of hunting so we will
draw our sample of hunters to send envelopes to at a later date.
We will be in touch with you regarding this later.

5.

Utilize the special window envelopes supplied for mailing the permits.
Give each applicant one of the sheets, also supplied, showing the open
area and information on hunting hours and bag limits.

6.

Hold all our copies of the permits (State Reference Cards) until you
hear from us.

7.

If you have questions, please call Clait Braun, Jack Grieb or me.

Howard D. Funk
Section Chief
Small Game Research

..

'

�-228(State Reference Card)
COLORADO SPECIAL BAND-TAILED
Val id

PIGEON HUNTING SEASON PERMIT

--:Hunting Season

No.

(Year)
Issued to:
(Name)
(Mailing Address)
(City)
(State)
(Zip Code)
Wing Collection Envelope
Packet No. :
_
Issued by:
Date:

Hunting License No.:

Top -- sheet Nuinber 1.

-----------------------~-------.

,

SPECIAL BAND-TAILED
Valid

PIGEON HUNTING SEASON PERMIT

Hunting Season

No.

(Year)
Issued to:

Hunting License No.: --.....:---Issued by:
Date:
Top -- sheet number 2.

_

�-229NOTE. TO ISSUING AGENT
Please be sure that the hunter's name and complete mailing address,
including zip code, are shown on this permit. Give a packet of.wing
collection envelopes to the hunters whose permits end with certain
numbers as specified in your letter of special instruction on issuing
permits and write the packet number on the S tate Reference Card in
:the space provided.

SPECIAL BAND-TAILED

PIGEON HUNTING SEASON PERMIT

This permit authorizes the person whose name and address appear
on the reverse side and whose signature appears below, to participate
in the special Band-tailed Pigeon hunting season in Colorado in
accordance with the proviSions of both Federal and State regulations
governing the special season •. Refer to State regulations for State
hunting license requirements.
The permittee's cooperation, when requested, in State and Federal
data-gathering activities (questionnaire survey, wirig collections,
etc.) will help determine the feasibility of future special seasons.
SIGNATURE OF HUNTER
(Not valid unless signed)

�-232-

r
L.
1''PGI

THE COLOHADO
WILDLIFE

ARE STUDYING

f:'-;,

.4

&lt;

d~~, ~~.)

OF GAME, FISH jHTD

DIVISION

TInS FRO!.I PIGEON

D"
PARKS lIND

THE Yr:~lnLY PRODUCT'ION

THE BUHr:r.U

OF BAND·TAILED

OF SPORTS

PIGEONS.

F'SHERIES

WE CAN

l,ND

DETEIWEi'U:;

vn: HEED, YOU~ HELP TO Mf.KE THIS STUDY SUCCESSFUL.

WINGS.

C'.':~n, ·C.....
t :J}
r~d ~I C-·"
,}f1r S '
I'N

L HeInov!? one complete wing !!:::~_each bird you
h:lg. D:;, not WIO.P wings in .i.")lu~;ti(', foil or other
material as this causes spoilopo.
EUV:':LOPE. us: A
SEPAHATE ENVELOPE FOR EliCH DAY'S HFHT.
DO NOT ENCLOSE VlINGS rnox ANOTHER

2. PLI.•.
CE WINGS

BUNTEH'S

IN THIS

BAG.

3. Mc il this envelope
POSTAGE

as soon as possible.

NO

REQUUED.

4. Continue

to send

us your wings

ecich time

you

go pigeon hunting.
5, Pl~m:/)fill out the quastionnoire below.

,r.lIIJIiUIt:,:s:.uu....~·£~-::~~~~~~

...
:.,~::t.tsII••
Siil2 ~

P LEAS
l. Deta of this hunt.

E

COM

Month.

.•
T('
.....
"':C~~.'";lo~~"Vi.~.i~~:~\.·,.v~.:;;

•..
~~ ...~t"Iff..~v:::~

P LET E
Day-

..tr~:;:.~x.

'PI.C&lt;l:.e de&gt; no~_wrire

__ ' _Year ..

t:

_

i

-;_..._.. _. _':::-:.;._:

in this l:~c~:. ,;

r

A

.~':

2. Location(s) where birds were shot.
Loco lity,

Direction
Norrie
3, Number

_

and distcnce from nearest

of town,

of hours you hunted

town

_

todcy.,

4. Eo w m:my bcridto ils din you knock down within
siUllt but could not retrieve i.,
.
_

.

,..; •...
, I
.

.. 'a ...· ~.. ,
'.

! -',

.--.,~ "-'~

-....----

- -. - - .- - -_.

---

--

-.

�-233STATE

OF

COLORADO

DEPARTMENT

OF

John

NATURAL

A. Love,

Governor

RESOURCES

R. Woodward.

OF NATURAL

T. W. Ten Eyck,

DIVISION OF GAME, FISH AND PARKS
Harry

DEPARTMENT
RESOURCES

Director

Executive

Director

GAME, FISH AND
COMMISSION

PARKS

Floyd Getz, President
LeRoy Robson, Vice President
Harrv Combs, Ser re tarv
C. M. F urneau x, Member
Ore-s t Ger baz. Member
John f. Holden, Member
O.

K. Niess,

September 21, 1970

Dear Band-it.a
Ll.edPigeon Hunce r ;

a

By obtaining
permit for the Colorado experimental band~tailed pigeon
hunting season from September 12 through September 20, 1970, you have
indicated an interest in pigeon hunting and the future of the sport.· We
respectfully ask your assistance in evaluating this experimental season by
completing the enclosed self-addressed questionIJ,aireand returning it as
soon as possible. Information you supply will be instrumental lit determin~ng the potential and feasibility of this type of season in the future.
Please read the questionnaire carefully and fill in all requested information completely and accurately. If you did not hunt during the experimental season, we ask that you complete Question No. 1 and return the
questionnaire because negative information is also important in our evaluation. If you did participate, please report only on your experimental
season hunting activities and not those of your friends or other members
of your family.
SpeCial Note: Some of you may also receive another questionnaire concerning your activities on hunting all small game species. Please fill
out and return that questionnaire as well as this one because the two surveys are completely independent of each other.
Thank you for your assistance.

CEB:al
Enclosure

Member

W. Robinson,
Ford Su-onq, Member
Dven Suttle, fAember

\A.'i::lu'll

MeJl1bH

�-234-

BAND-TAILED PIGEON HUNTER REPORT
1. Did you hunt band-tailed pigeons during the 1970 season?
Yes
No
2. County or area most frequently hunted
3. How many days did you hunt during the pigeon season? c=J

~~----------~-----

4. How many pigeons did you bag during the entire season?c=J
5. How many pigeons did you knock down within sight but
could not retrieve?

0

6. Please indicate in the appropriate boxes below the number of
pigeons killed each day you hunted. Put an "0" for the days
you did not .get any birds. Leave days you did not hunt blank.
Is t
t
2nd 2nd
Sat. Sun. MOn. Tues. Wed. Thurs. Fri. Sat. Sun.

-rs

D 0 o 0

o o ODD

Band-tailed Pigeon Hunter Survey
Game Research Center
P. O. Box 567
Fort Collins, Colorado 80521

�-235STAT:E

OF

COLORADO

DEPARTMENT

OF

John

NATURAL

A. Love.

DEPARTMENT
RESOURCES

Governor

RESOURCES

T. W. Ten Eyo'.
Execchve

DIVISION OF GAME, FISH AND PARKS
Harriy

R. Woodward.

OF NATURAL

Director

GAME. FISH AND
COMMISSION

Director

PARKS

Floyd Gctz , President
LeRoy

Robson,

Harry

Ccmhs

C. M. ;

Vice

Pr e sidvnt

Scc-c rorv

,

-neau-

t\~'-':~h(~

Or esr ::'(,rL.).:, t.~( . b'r
john
O.

Game Research Cen~~~~
P. O. Box 567
Fort Collins, Colorado 80521
October 5, 1970

E. Holoeo .
K. Niess,

Wrllrun
Fc.rc

.

.

Thank you!
Sincerel~

CEB:a1

Mt:-lI1ht::

Dean Su n.e. N,(: ~.I,,'

We recently sent you a questionnaire pertaining to your hunting
activities during the experimental band-tailed pigeon hunting
season in southern Colorado from September 12 through September
20, 1970. So far we have. not received your reply. Please assist
us by completing the questionnaire and returning it to us as
quickly as possible. Information you supply, ~
though you did
~
participate, would enable us to fully evaluate the special
season and would assist us in determining the potential and
feasibility of such a season in the future.

~y

~N. Robino:.or,

~'lrr:·nq,

Dear Band-tailed Pigeon Hunter:
.

A."l'!'

R. Woodward, Director

h

Mt.:'n'.:.N
M

r-,

�-236BAND-TAILED PIGEON - HUNTER BAG CHECK
INSTRUCTIONS - USE ONE SHEET PER HUNTING PARTY
Crop Condition:
.Active:

- Be sure to examine interior of crop •

Loose curds (white to yellow) of pigeon milk present in crop.

Stimulated:
Inactive:

Convoluted glartdular material
side of the inside crop wall.

Normal, thin crop wall without glands.

Innnature: No white neck ring present
gray appearance.
Adult:

Locat.Lon
Age.

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30

(may be very faint), unf.form slate

Definite white neck ring present, bird has purplish brown tint to
breast and head.

Date
Bird No.

(appears like brains) on either

(A or I)

------------~--------------------------------------

Crop Condition
Active
Stimulated
Inactive

..

.

Sex·
(If Gonads were Checked)

Band No.
if Banded

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                  <text>-1-

January, 1972

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~--------

Project No.

w-4l'-R-2l-22

Work Plan No.

1

Bighorn Sheep &amp; Mountain Goat Investigations
Job No.

12

Job Title

Location and Distribution of Bighorn Sheep Herds in Colorado

Period Covered:

June 1, 1970 to May 31, 1972

Personnel:

George D. Bear

ABSTRACT

The report is presently being edited and should be published during the summer
of 1972.

��-3-

January,

1972

JOB FINAL REPORT

S ta te of
Project
Work

.::.CO.::.L::.O::::.:RA=D::..:O=-_
No.

W-4l-R-2l-22

Bighorn

Plan No.

1

Job No.

Job Title:

Evaluation

Period

June

Covered:

Personnel:

William

of Proposed

Sheep

&amp; Mountain

Goat

Investigations

14
Bighorn

Sheep

Transplant

Sites

1, 1969 to May 31, 1972

H. Rutherford

ABSTRACT

Game Information Leaflet No. 93, entitled "Guidelines
for Evaluating
Bighorn
Sheep Transplanting
Sites in Colorado", is the final report for this job.

��-5-

EVALUATION OF PROPOSED BIGHORN SHEEP TRANSPLANT SITES
William H. Rutherford

P. S. OBJECTIVE
To situate bighorn sheep transplant opportunities.
METHODS AND MATERIALS
Habitat characteristics of a wide variety of bighorn sheep-occupied areas
were observed and measured. Primary consideration was given to: Vegetative
types, competition of other ungulates with bighorns, topographic features,.
snow depth, and human influence. This infor~mation was interpreted and
'
applied toward a set of guidelines for evaluating proposed transplant sites.
RESULTS AND DISCUSSION
A set of general guidelines for evaluating potential bighorn sheep transplant sites was developed. These guidelines were published as a Game
Information Leaflet (Rutherford 1972), and made available to management
personnel in this form. Thus, the formal publication becomes the final
report for this job.

LITERATURE CITED
Rutherford, W. H. 1972. Guidelines for evaluating bighorn sheep
transplanting sites in Colorado. Colo. Div. of Game, Fish and
Parks, Game Info. Leaflet No. 93. April, 1972.

Wildlife Researcher

��-7-

January, 1972

JOB FINAL REPORT

State of

COLORADO

Project No.

w-41-R-21

Bighorn Sheep &amp; Mountain Goat Investigations

Work Plan No.

1
------~-----------

Job No.

15

Job Title:

Mortality Factors in Declining Bighorn Sheep Herds

Period Covered:

June 1, 1969 to May 31, 1971

Personnel:

Thomas N. Woodard, Ralph J. Gutierrez, William H. Rutherford

ABSTRACT

A Master of Science thesis (Woodard 1971) is the final report for this job.

��-9-

MORTALITY FACTORS IN DECLINING BIGHORN SHEEP HERDS
William H. Rutherford

P. S. OBJECTIVE
To identify factors responsible for, or contributing to, decline in
bighorn sheep herds in Colorado.
METHODS AND MATERIALS
Observation of habits, movements, behavior, and composition of the ewelamb herd inhabiting the Sand Creek area in the Sangre de Cristo Range,
from early summer through fall during two separate years, and interpretation of the observation data thus obtained, was done. Necropsy of two
lambs collected the second year of study was done by staff members of the
College of Veterinary Medicine, Colorado State University, with the results made available to the principal investigator.
RESULTS AND DISCUSSION
A Master of Science thesis (Woodard 1971) was prepared, based on the data
and information obtained from this study. This thesis, copies of which
are on file at the Game, Fish and Parks Division Research Center in Fort
Collins and at the Colorado State University Library, becomes the final
report for this job.
LITERATURE CITED
Woodard, Thomas N. 1971. Bighorn sheep lamb production, survival, and
mortality in the Sangre de Cristo Mountains, Colorado. M. S. Thesis,
Colorado State University, Fort Collins. 84 pp. (unpublished).

Prepared by

-id(£,. 11 &amp;:-d;;;;~
William H. Rutherford
Wildlife Researcher

!

��January,

-11-

1972

JOB FINAL REPORT

State of

COLORADO

Project No.

W-4l-R-2l-22

Work Plan No.

Bighorn

2

Sheep

4

Job No.

Job Title:

Location

Period Covered:

June 1, 1965 to May 31, 1972

Personnel:

&amp; Mountain Goat Investigations

and Distribution

L. Dale Hibbs, William

of Mountain

H. Rutherford,

Goat Herds in Colorado

Thomas N. Woodard

ABSTRACT

Game Information Leaflet No. 90, entitled "Status of Mountain Goats in Colorado",
is the final report for this job. In addition, detailed written descriptions
for access to specific herd areas, recommendations for census procedures, and
forms for recording census and harvest data have been prepared for management
personnel.

��-13-

LOCATION AND DISTRIBUTION OF MOUNTAIN GOAT HERDS IN COLORADO
William H. Rutherford

P. S. OBJECTIVE
To determine numbers, composition, seasonal range, and location of
mountain goat herds in Colorado.

METHODS AND MATERIALS
Aerial and ground observations of mountain goats were made at all seasons
of year, in all known areas of occupancy, in cooperation with Regional
Game Biologists. Maps and forms were prepared based on the information
thus obtained.
RESULTS AND DISCUSSION
The current (1972) status of mountain goats in Colorado is reported in
Game Information Leaflet No. 90 (Rutherford 1972). Detailed written
descriptions for access to specific herd areas, recommendations for
census procedures, and forms for recording census and harvest data have
been prepared. These have all been made available to management personnel, and constitute the final report for this job.

LITERATURE CITED
Rutherford, W. H. 1972. Status of mountain goats in Colorado. Colo.
Div. of Game, Fish and Parks. Game Info. Leaflet No. 90. April,
1972.

Wildlife Researcher

�-14-

Descriptions of access routes and census recommendations for mountain
goat herds in Colorado, submitted to Game Management personnel as a
part of the requirements for Work Plan 2, Job No.4, W-4l-R.

MOUNT EVANS MOUNTAIN GOAT AREA
Access:
Access to the primary goat range, the Owl's Head Burn on the south-facing
slope of Porcupine hill, can be either from the end of the road on Bear
Creek or from the Echo Lake-Lincoln Lake trail. The only mode of travel
into the area actually frequented by the goats is by foot or horseback.
If the access route from the valley bottom in Bear Creek is chosen,climbing
will be steep, rough, and rapid. It is suggested that travel by vehicle
be made to the Echo Lake Campground by traveling southwest from Idaho
Springs on State Highway 103, which will accomplish a great deal of altitudinal gain. From here, it is a fairly easy hike along the Lincoln Lake
trail to the top of the ridge south of Vance Creek. At this point, leave
the trail and head nearly due east along the south side of the ridge until
the burn area is entered. Goats may be found anywhere in this area.
Access to the alpine areas to the south and west of the Owl's Head Burn
can be from the Mount Evans Highway and then by foot around Mount Evans,
by foot or horseback along the Echo Lake-Beartracks Lake trail to the
Mount Rosalie-Mount Epaulet area, and from the Grant-Guanella PassGeorgetown road and then by foot around Mount Bierstadt.
Census:
Because of the difficulty in locating goats in the standing dead timber of
the burn area, census by fixed-wing aircraft is almost impossible. Aerial
census should be done by helicopter, but even this technique has limitations
when sex-age classification data are desired. The most accurate herd composition census is undoubtedly from the ground, by an observer using
binoculars and spotting scope. It is suggested that foot travel be made
from the Lincoln Lake trail, as described above, to one of the promontories
at the edge of the Owl's Head Burn. The goats can then be located by using
binoculars, and the observer can use his judgment to either stay where he
is or to move to another vantage point. The suggested time for conducting
these census observations is late summer and fall.

�WP 2, J 4, W~4l-R

-15-

COLLEGIATE RANGE MOUNTAIN GOAT AREA

Access:
Access to the Mount Shavano-Jones Peak area is best made from the east
side. From the junction of U. S. Highways 50 and 285, north of Poncha
Springs, proceed west by vehicle on Highway 50 for 1.7 miles, then turn
north on county road. Proceed north for 1.2 miles, at which time the
county road becomes a BLM dirt road. Follow this dirt road northwest
about 6.0 miles to Blank Cabin, then about another 0.3 miles to the
beginning of the Shavano Trail. Then proceed by foot or horseback up
the trail to Mount Shavano. Tabeguache Peak and the west side of
Mount Shavano are best reached by hiking up McCoy Creek, which is a
tributary of the North Fork of the South Arkansas River. Leave U.S.
Highway 50 at Maysville and proceed northwest by vehicle on the North
Fork road about 6.0 miles to the mouth of McCoy Creek. The Mount Aetna
area can be reached by hiking up Hunkydory Gulch, another tributary
which enters the North Fork of the South Arkansas River about 2.5 miles
upstream from the mouth of McCoy Creek.
Access to the Mount Antero-Carbonate Peak area is best made by vehicle
travel up the Chalk Creek road from Nathrop, on U.S. Highway 285, about
11.0 miles to the mouth of Baldwin Gulch, which enters Chalk Creek from
the south, and then hiking up Baldwin Gulch.
Access to the Mount Princeton area is also made from the Chalk Creek
Road. Directly opposite the mouth of Baldwin Gulch, on the north side
of Chalk Creek, proceed by foot up the Grouse Canyon trail to the west
side of Mount Princeton.
The Gladstone Ridge-Sheep Mountain area is best reached from South
Cottonwood Creek. A climb by foot directly north from the Cottonwood
Lake Campground is the shortest way to reach Sheep Mountain. From this
point, hiking directly west will enable the traveler to reach Gladstone
Ridge; or alternatively, Gladstone Ridge can be reached by vehicle travel
up the South Cottonwood Creek road to a point 5.0 miles upstream from the
Cottonwood Lake Campground, and then hiking north up Grassy Gulch.
Census:
Aerial census of mountain goats in the Collegiate Range is entirely feasible
and practical. Census from the ground with binoculars and spotting scope,
using the access routes to specific herd areas previously described, can be
quite accurate if it is done thoroughly, but such a census is too timeconsuming for an area as large as this one. Any census attempt in the
Collegiate Range should be planned for collecting herd composition data
as well as ordinary enumeration of goats, and for this reason the helicopter is a better choice.

�WP 2, J 4, W-4l-R

-16-

It is suggested that the three main herd areas are the only ones in which
aerial census is likely to be productive. A systematic coverage of the
alpine areas of Mount Shavano-Jones Peak, Mount Antero-Mount Princeton,
and Gladstone Ridge, plus the below-timberline areas of Sheep Mountain,
should provide adequate numerical trend and herd composition data. These
counts are best made in late summer Or fall.

NEEDLES MOUNTAINS MOUNTAIN GOAT AREA
Access:
There is no access to any portion of the Needles Mountains except by foot
or horseback. In either case, the trip should not be made without preparation
for spending at least two nights camped out. Horseback access is best made
by departing from the trail head above Vallecito Reservoir on Vallecito
Creek. Approximately eight miles up Vallecito Creek, the trail forks at
the mouth of Johnson Creek which comes in from the west. Take the Johnson
Creek trail for a distance of about five miles to the top of Columbine
Pass, and then another two miles down into Chicago Basin, at the head of
Needle Creek, where good campsites can be found. Most of the access to
specific areas such as Mount Aeolus, Sunlight Peak, Windom Mountain,
Pigeon Peak, and Turret Peak is by hiking only, as horses cannot be used
in the steep cliffs and rockslides.
Access from the west side, strongly recommended in case of backpack hiking
without horses, is from the narrow gauge railway in Animas Canyon between
Durango and Silverton. Departure from the railroad can be made at the
mouth of Needle Creek, Ruby Creek, or No Name Creek, according to the wishes
of the hiker. Needle Creek has the best trail.
Census:
The problems in obtaining systematic trend-type census data on goats in the
Needles Mountains are almost insurmountable. The best that can be done is
to make one or two observation flights per year by fixed-wing aircraft to
locate goats and ascertain whether kids are present. A flight in late
summer and one in mid-winter should suffice. Until the herd grows larger,
the expense of making helicopter flights can hardly be justified, unless
such flights can be combined with winter elk census operations in the
Animas River Valley.

�WP 2, J 4, W-4l-R

-17-

GORE RANGE MOUNTAIN GOAT AREA

Access:
Access to the goat release site on Black Creek is by vehicle up the Black
Creek Road. The creek comes into the Blue River from the southwest at the
upper end of Green Mountain Reservoir. The road extends up the creek for
approximately four miles to the boundary of the Gore Range-Eagles Nest
Wild area. From this point, the headwater area of Black Creek is reached
by hiking.
Census:
While the goat herd is small and still becoming established, the only
purpose in conducting census operations is to check on distribution and
reproduction.
This is best done with aircraft, either fixed-wing or
helicopter. A fixed-wing flight in late summer, and a helicopter flight
in conjunction with winter elk census, should be made.

��-19-

January, 1972

JOB FINAL REPORT

State of

COLORADO

Project No.

W-41-R-2l-22

Work Plan No.

2
--------~---------

Bighorn Sheep &amp; Mountain Goat Investigations

Job No. 5

Mountain Goat Habitat Requirements

Job No.6

Evaluation·of Proposed Mountain Goat Transplant Sites

Period Covered:

June 1, 1965 to May 31, 1972

Personnel:

L. Dale Hibbs, William H. Rutherford, Thomas N. Woodard

ABSTRACT

Game Information Leaflet No. 91, entitled "Guidelines for evaluation of Mountain
Goat Transplant Sites", is the final report for these two jobs.

��-21-

MOUNTAIN GOAT HABITAT REQUIREMENTS
EVALUATION OF PROPOSED MOUNTAIN GOAT TRANSPLANT SITES
William H. Rutherford

P. S. OBJECTIVE
Job No.5:

To determine habitat characteristics of presently occupied
mountain goat range in Colorado.

Job No.6:

To situate mountain goat transplant opportunities.
METHODS AND MATERIALS

Habitat characteristics of all known mountain goat-occupied areas were
observed and measured. Primary consideration was given to: Vegetative
types, competition of other ungulates with mountain goats, topographic
features, snow depth, and human influence. This information was inter~
preted and applied toward a set of guidelines for evaluating proposed
transplant sites.

RESULTS AND DISCUSSION
A set of general guidelines for evaluating potential mountain goat
transplant sites was developed. For publication purposes, Jobs 5 and 6
were combined, since both jobs were used as input in developing these
guidelines. Publications as a Game Information Leaflet (Rutherford 1972)
was done, and made available to management personnel in this form. Thus,
the formal publication becomes the final report for these jobs.
LITERATURE CITED
Rutherford, W. H. 1972. Guidelines for evaluation of mountain goat
transplant sites. Colo. Div. of Game, Fish and Parks, Game Info.
Leaflet No. 91. April, 1972.

---7

Prepared by

UIk" Ii !fr:;~~

William H. Rutherford
Wildlife Researcher

~~V~

��-23January,

1972

JOB PROGRESS REPORT

Sta te of

~CO::::.;L=_O::::.;RAD:::::::::.:O::..._
_
An Ecological Investigation
of
the
Cache la Poudre Deer Herd, Colorado
W-lOS-R-l2

Project No.

Job No.

Work Plan No.

3

Job Title

Ecological

Period Covered:
Personnel:

March 1, 1971 to February

Robert D. Roughton

in press is listed.

(Eleventh Year)

History of Key Browse Species

ABSTRACT

A manuscript

6

28, 1972

��-25-

ECOLOGICAL HISTORY OF KEY BROWSE SPECIES
Robert D. Roughton
P. S. OBJECTIVES
1.

Investigate and refine suitable age determination techniques for individuals of selected browse species.

2.

Determine the age structure of selected browse populations on the Cache
la Poudre deer winter range.

3.

Draw ecological inferences such as pattern of species reproduction,
successional relations between species, treatment influences, and individual longevity by relating age structures to known site histories and
species association.

SEGMENT OBJECTIVE
Prepare manuscript from M. S. Thesis.

RESULTS AND DISCUSSION
Progress
A manuscript was submitted, revised and finally accepted by the Editor of
Ecology.
Manuscript in Press
Roughton, R. D. 1972. Shrub age structures on a mule deer winter range in
Colorado. Ecology (In Press).

Prepared by

~I

e~

Allen E. Anderson
Wildlife Researcher

��-27January, 1972

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~--------

Project No.

W-105-R-12

Work Plan No.

4

Job Title

Job No.

1

(Eleventh Year)

Population Density and Structure

Period Covered:
Personnel:

An Ecological Investigation of
the Cache la Poudre Deer Herd, Colorado

March 1, 1971 to February 28, 1972

Allen E. Anderson

ABSTRACT

A publication and a manuscript in press are listed.

��-29-

POPULATION DENSITY AND STRUCTURE
Allen E. Anderson

P. S. OBJECTIVE
1.

Estimate population density on each of five selected study locations
believed to be representative of the lower~inter, middle-winter, upperwinter, transitional, and summer ranges of the herd to: (a) relate density
to changing environmental conditions (Work Plan 3, Jobs 1 and 3), and
(b) elucidate density-elevational relationships, particularly between
years.

2.

Estimate sex and age structure (composition) of the population to provide
basic information on herd dynamics.

SEGMENT OBJECTIVE
Tabulate, summarize, analyze, and interpret collected data, review recent
literature, and prepare manuscripts for publication.
RESULTS AND DISCUSSION
Progress
Estimation of herd structure and population densities on selected areas was
completed prior to June 1, 1965. Analyses of herd structure samples-timeweather relationships have been completed.
Publications:
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1972. Mule deer fecal group
counts related to site factors on winter range. J. Range Mgmt. 25:66-68.
Manuscripts in Press:
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1972. Mule deer numbers and
shrub-yield utilization on winter range. J. Wildl. Mgmt. 36 (April).
Future Plans
Prepare brief manuscript on the herd structure samples-time-weather

Prepared by

e

~
~4""1
Allan E. Anderson
Wildlife Researcher

relationships.

��-31January, 1973

JOB PROGRESS REPORT

State of

COLORADO
------------~-------------

Project No.
Work Plan No.

An Ecological Investigation
of the Cache la Poudre Deer Herd, Colorado

W-105-R-12
5

Job Title

Job No.

1

(Eleventh Year)

Physical Characteristics

Period Covered: March 1, 1971 to February 28, 1972
Personnel: Allen E. Anderson

ABSTRACT

Publications, manuscripts in press, and manuscripts in preparation are listed.

��-33-

PHYSICAL CHARACTERISTICS
Allen E. Anderson

P. S. OBJECTIVE
Measure relevant physiological and morphological characteristics of deer collected seasonally so that: (a) the response of the herd to its environment
can be more adequately interpreted. (b) establish "physiological norms", and
(c) provide basic morphological data as related to sex and age classes.
SEGMENT OBJECTIVE
Tabulate, summarize, analyze. and interpret collected data, review recent literature, and prepare final report and publication manuscripts.
RESULTS AND DISCUSSION
Progress
Physiological and morphological studies on four deer collected each month,
1961-1965 were completed April 27, 1965. The number totaled 192 regular collections plus 33 from supplementary sources. All data were placed on punchcards and blood and tissue chemistry data were completed during Segment 12.
The review of literature was continued and about 1,000 references were obtained
for project files during Segment 12.
Publications
Anderson, A. E •• D. E. Medin, and D. C. Bowden. 1971. Adrenal weight in a
Colorado mule deer population. J. Wildl. Mgmt. 35(4):689-697.
Anderson, A. E., and D. E. Medin. 1971. Antler phenology in a Colorado mule
deer population. Southwestern Naturalist 15(4) :485-494.
Nicolls, K. E. 1971. A light microscopic study of nuclear and cytoplasmic
size of the aggregate acidophil population in the hypophysis cerebri pars
distalis of the mule deer Odocoileus hemionus hemionus relative to seasons
of the photoperiod and antler cycles. Z. Zellforsch Mikros. Anat. 115:
314-326.

~~nuscripts in Press
Anderson, A. E •• D. E. Medin, and D. C. Bowden. 1972.
a population of mule deer. J. Mammal. 53 (May).

Total serum protein in

�-34-

Anderson, A. E., D. E. Medin, and D. C. Bowden. 1972. Indices of carcass
fat in a Colorado mule deer population. J. Wild1. Mgmt. 36 (April).
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1972. Carotene and vitamin A
in the liver and blood serum of a Rocky Mountain mule deer (Odocoileus
hemionus hemionus) population. Compo Biochem. Physiol. 40 p. (April).
Manuscripts in Preparation
Anderson, A. E., D. E. Medin, and D. C. Bowden. Biometrics of the organs, glands,
and carcass of Rocky Mountain mule deer. Wildl. funograph.
Anderson, A. E., D. E. Medin, and D. C. Bowden. Seasonal variation in the
weight and color of the pelage of mule deer. J. Mammal.
Future Plans
Complete and submit final manuscripts to appropriate Journals.

�-35January,

1972

JOB PROGRESS REPORT

Sta te of

.:;;.CO.:;;.L;;:;,O,;;.;RA=D=-O:::.._
An Ecological Investigation
W-105-R-12
of the Cache la Poudre Deer Herd, Colorado

Project No.
Work Plan No.

Job No.

5

Job Title

Reproductive

Period Covered:
Personnel:

March 1, 1971 to February

Allen E. Anderson

ABSTRACT

A manuscript

in preparation

is described.

2
Studies

28, 1972

(Eleventh Year)

��-37-

REPRODUCTIVE STUDIES
Allen E. Anderson
P. S. OBJECTIVE
Determine the reproductive pattern of the deer herd to provide data on:
(a) morphology of the reproductive organs as related to age and season.
(b) gross and net productivity between years, and (c) tentatively, the relationship of productivity to measured factors of the environment (Work Plan 3).
SEGMENT OBJECTIVE
Tabulate. summarize, analyze, and interpret collected data, review recent
literature, and prepare final report and publication manuscripts.
RESULTS AND DISCUSSION
Progress
Reproductive data gathering from collected deer was completed April 27, 1965.
Ovarian collections from hunter killed deer were completed with the 1964
hunting season. Growth curves of certain male and female reproductive organs
were calculated and plotted.
Publications
Markwa1d, R. R., R. W. Davis, and R. A. Kainer. 1971. Histological and histochemical periodicity of cervine Leydig cells in relation to antler growth.
Gen. Com. Endocrino1. 16:268-280.
Short, Catherine. 1970. Morphological development and aging of mule and
white-tailed deer fetuses. J. Wi1d1. Mgmt. 34(2) :383-388.
Manuscripts in Preparation
Portions of this material will be included in a monograph on growth of organs
and glands as described in Work Plan 5, Job 1, Physical Characteristics.
Future Plans
Complete and submit monograph manuscript to Wildlife Monographs.

Prepared by

e ~~

~
Allen E. Anderson
Wildlife Researcher

��-39January, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-105-R-12

Work Plan No.

5

Job Title

Job No.
Harvest Analysis

Period Covered:
Personnel:

An Ecological Investigation
of the Cache la Poudre Deer Herd, Colorado

March 1, 1971 to February 28, 1972

Allen E. Anderson

ABSTRACT

Publication plans are outlined.

3

(Eleventh Year)

��-41-

HARVEST ANALYSIS
Allen E. Anderson

P. S. OBJECTIVES
1.

Determine the age structure of the deer herd kill to provide estimates
of (a) net productivity (percent female yearlings in the kill), and (b)
the effects of hunting regulations.

2.

Locate the distribution of the kill by sub-unit to relate age and sex
structure of the kill to elevational levels and harvest intensity.

3.

Measure relevant physical characteristics of the kill to provide a
possible index of herd response to winter food quality.
RESULTS AND DISCUSSION
Progress

Check station sampling of the fall harvest was completed with the 1965 hunting
season. Collected data have been tabulated and statistical treatment has been
completed. Most of the literature has been reviewed.
Publications
None.
Manuscripts in Preparation
None.
Manuscripts Submitted
None.
Future Plans
Incorporate findings relative to Objectives 1 and 2 in a Department Special
Report. The first draft will be completed during Segment 13.

Prepared by

~e"",.,. ~ ~~
Allen E. Anderson
Wildlife Researcher

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JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-37-R-25
1

Work Plan No.
Job Title

16

Job No.
Pheasant Nest Site Selection

Period Covered:
Personnel:

Game Bird Survey

Study

April 1, 1971 to March 31, 1972

Warren D. Snyder, Carroll J. Grand Pre, Lawrence A. Webster,
Robert L. Schmidt, John F. Corey, George G. Harrington, Larry
Crooks, John Ellenberger, Thomas Lines, Dale Stahlecker, Rusty
Kozacek, Dr. David Bowden and Donald M. Hoffman.

ABSTRACT
Each of the four quadrats were stocked with a minimum of 13 hens and 3 cocks by
January 28, 1971(Segment 24) in preparation for nesting studies in 1971. Fortyeight pheasant nests were found in the 48 nesting plots, representing six vegetative species or combinations of species.
Eight nests were found in intermediate
wheatgrass mowed plots; six nests were found in alfalfa unmowed plots; five nests
each were found in alfalfa mowed, intermediate wheatgrass unmowed, tall wheatgrass
unmowed and alfalfa-crested wheatgrass unmowed plots; and four nests each were
found in smooth bromegrass unmowed and crested wheatgrass unmowed plots.
Fewer
numbers of nests were found in other vegetative types and treatments represented.
A Chi-square analysis of data showed that a significant preference (P&lt; 0.05) for
cover types or treatments could not be shown in 1971. Seventy-seven percent of
all nests in 1971 were established 15 feet or less from plot perimeters.
Average
clutch size of 19 nests, where it was possible to determine this, was 9.1. Field
work on this study was concluded in 1971 with the removing of the wing-clipped
wild pheasants and windrowing of vegetation in all plots on July 23, 1971.

��-3-

PHEASANT NEST SITE SELECTION STUDY
Donald M. Hoffman

P. S. OBJECTIVE
To compare pheasant nesting use of, and success in (1) winter wheat; (2)
alfalfa; (3) crested wheatgrass; (4) hairy vetch; (5) white sweet clover;
(6) alfalfa-crested wheatgrass mixture; (7) smooth bromegrass; (8) tall
wheatgrass; and (9) intermediate wheatgrass.

SEGMENT OBJECTIVES
1.

To maintain plots.

2.

To measure nesting preference.

3.

To determine nesting success by cover type.
METHODS AND MATERIALS

Fifty-nine wild pheasants were captured by 4 ~enusing hand nets, hand spotlights, and 2 4-wheel drive vehicles in Phillips and Sedgwick counties in
only 6 trap nights during Segment 24. In addition, 6 wild pheasants were
captured with wire cloverleaf traps at the Fort Collins Wildlife Research
Station. All 4 quadrats were stocked with a minimum of 13 hens and 3 cocks
by January 28, 1971. All pheasants were wing clipped on one wing and leg
banded prior to release in the quadrats. Considerable effort was again
expended to control predation on the wing clipped wild pheasants. Three
great horned owls and several feral cats were taken in control operations.
Phenology measurements were again secured in 1971 to ascertain readiness of
the various vegetative species and cover types for use by nesting pheasants.
Three separate, complete nest searches were again made in 1971. An initial
nest search in 1971 was made during the period June 3 through 10, with 8 men.
Each plot was systematically searched using ropes to divide the plots into
narrow segments. Nests were marked, recorded, and removed to encourage
renesting. A second nest search was made during the period June 23 through
28 by 3 men searching each plot systematically in strips outlined by lath
markers. A third and final search was made by 4 men using methods similar
to the second search during the period July 19 through 23, 1971, preceding
and following the windrowing of all cover plots on July 23, 1971.
Distances were measured from each nest to the nearest edge and to the apex or
base of the triangular plot, whichever was closest, with a steel tape. Each
nest could later be located on a scaled diagram of the quadrats. Number of
eggs, fate of the nest and estimated age of embryos were recorded. In addition,
vegetative heights and vegetative species in the 4 cardinal directions from
each nest were recorded.

�-4-

Prior to the windrowing of vegetation on all plots on July 23, 1971, most of
the wing clipped pheasants were caught with hand nets. Thirty two of these
were released three miles west of Holyoke in Phillips County and the remainder
were released outside the quadrats at the Fort Collins Wildlife Research
Station. Only one incubating hen was killed by the windrowing machine in 1971.
Nesting data gathered in the 5 year period from 1967 through 1971 were coded
for computer analysis and submitted to Dr. David C. Bowden in October, 1971.
Results and interpretations of the statistical data will be included in a final
report.
Plot Composition
Plot Composition for 1971 (Treatment Following 1970 Nesting Season)
The changing of vegetative types used very little, or not practical to
establish along roadsides, started in 1968, was completed in the fall of 1969.
No additional changes were made during 1970.
Plot Number 1.

Originally Winter Wheat, Now Smooth Bromegrass

The 4 original winter wheat plots seeded to smooth bromegrass in the fall of
1968 were left unmowed in the summer of 1970. The 4 original winter wheat
plots, seeded to smooth bromegrass in the fall of 1969, were mowed in the
summer of 1970. A residual cover check was secured in 1971.
Plot Number 2.

Alfalfa

Four of these were mowed and 4 were left unmowed in 1970 allowing a residual
cover check in 1971. Two of the alfalfa plots including Number 2, Interior,
NW Quadrat and Number 2, Exterior, NE Quadrat, are approaching alfalfacrested wheatgrass mixtures due to invasion by crested wheatgrass.
Plot Number 3.

Crested Wheatgrass

Four of these were mowed and 4 were left unmowed following the 1970 nesting
season, allowing a residual cover check in 1971 on half of the plots. Composition for 1971 was therefore comparable to that of 1970.
Plot Number 4.

Orignially Hairy Vetch, Now Intermediate Wheatgrass

All of these plots were reworked and seeded to intermediate wheat grass following
the 1968 nesting season. Four of these intermediate wheatgrass plots were
mowed and 4 were left unmowed following the 1970 nesting season, allowing a
residual cover check in 1971 on half of the plots.
Plot Number 5.

Originally White Sweet Clover, Now Tall Wheatgrass

The 4 original white sweet clover plots seeded to tall wheatgrass in the fall
of 1968, were left unmowed in the summer of 1970. The 4 original white sweet
clover plots seeded to tall wheatgrass in the fall of 1969, were mowed in the
summer of 1970. A residual cover check was secured in 1971.

�-5-

Plot Number 6.

Alfalfa-Crested Wheatgrass

Four of these were mowed and 4 were left unmowed following the 1970 nesting
season, allowing a residual cover check in 1971 on half of the plots.
Composition for 1971 was therefore comparable to 1970.

RESULTS AND DISCUSSION
Stocking of Pheasants for 1971
Only 6 trapnights, using two 4-wheel drive vehicles, 4 hand spotlights, and
2 men per vehicle with hand nets were requred to capture 59 wild pheasants
in Phillips and Sedgwick counties in January, 1971. Six additional wild
pheasants were captured in wire cloverleaf bait traps at the Fort Collins
Wildlife Research Station. All 4 quadrats were stocked with a minimum of 13
hens and 3 cocks by January 28, 1971. All pheasants were leg banded and wing
clipped prior to releasing in the quadrats.
Predator control was again accomplished in 1971 after 1 hen was killed by a
great horned owl in early February, 1971. The removal of 3 great horned owls
and several feral domestic cats prevented undue losses of the wing clipped
pheasants prior to the first nest search period. At least 5 hens were killed
later in June and July, 1971 by a feral cat which entered the southwest quadrat
through a small hole in the perimeter fence, mistakenly left open by Station
personnel. The cat thought to have been responsible was trapped but most of
the surviving pheasants in the southwest quadrat also escaped through this
hole in the perimeter fence.

Location of Nests
Forty eight pheasant nests were found in the 48 triangular shaped nesting
plots representing 6 vegetative species or combination of species. Figures 1
through 4 show the approximate locations of nests in the various quadrats.
Table 1 lists pheasant nests found by vegetative cover types and treatments in
1971.
By number of nests, intermediate wheatgrass mowed (8 nests) ranked highest
for nest establishment in 1971, followed by alfalfa unmowed (6 nests); alfalfa
mowed, intermediate wheatgrass unmowed, tall wheatgrass unmowed, and alfalfacrested wheatgrass unmowed (each with 5 nests); and smooth bromegrass unmowed
and crested wheatgrass unmowed (each with 4 nests). Fewer nests were found
in other vegetative types and treatments represented. Numbers of plots by
cover type and treatments were equal with 4 of each kind in 1971, so a separate
ranking by nests per plot was not required.
In a Chi-square analysis of data, significant preferences for individual cover
types or treatments (P&lt;0.05) could not be shown in 1971. This result was in
contrast to the years 1969 and 1970 when significant preferences were indicated. Two severe hailstorms occurred during the early portion of the 1971

�-6-

nesting season on May 22 and June 1. Three nests appeared to have been
abandoned following these hails, but more importantly, it also appeared that
a number of the hens did not recycle. The 48 nests were probably less than
would have been recorded without adverse weather plus feral cat depredations
described earlier.
Seventy seven percent of all nests in 1971 were established 15 feet or less
from plot perimeters, as shown in Table 2. An inspection of plot diagrams
indicates that on an average, 70 percent of the total plot area is contained
in a perimeter strip 15 feet wide so this may not indicate a significant edge
effect. During 1971, 28 nests were found in interior plots and 20 in exterior
plots. The terms "interior" and "exterior" refer to the location of the individual plots in relation to a central point where all 4 quadrats join. These
are indicated in the margins of Figures 1 through 4.
Clutch Sizes
TWenty nine nests contained unknown clutch sizes. These were nests where the
hen was still laying, or nests which were abandoned for one reason or another
without incubation having started. The average clutch size of the remaining
19 nests, which included both incubating and hatched nests was 9.1 eggs,
ranging from 4 to 17 (Table 3).
Considerable variation was again found in clutch sizes between the various
cover types and treatments, as shown in Table 3. Highest average numbers of
eggs per clutch was found in the intermediate wheat grass unmowed plots (1 nest
with 13 eggs), intermediate wheatgrass mowed plots (4 nests averaged 11.8 eggs),
crested wheatgrass unmowed plots (1 nest with 10 eggs), alfalfa unmowed (4 nests
averaged 9.2 eggs), and tall wheatgrass mowed plots (1 nest with 9.0 eggs).
Fate of Nests
Forty eight nests were found and recorded. Of these, 32 were determined to
be in an active state of laying or incubation, 5 were hatched successfully,
and 11 were apparently abandoned for one reason or another (Table 4). All
nests were recorded and removed to encourage renesting. This also prevented
the recording of the same nest more than once.
Of the 5 nests which hatched successfully, one each was in alfalfa mowed,
alfalfa unmowed, intermediate wheatgrass mowed, tall wheatgrass unmowed, and
alfalfa-crested wheatgrass unmowed plots. Of 31 total eggs in the 5 nests
which had successfully hatched, 28 (90.32 percent) of the eggs hatched.
Of the 11 apparently abandoned nests, 3 showed extensive hail damage, 2 were
destroyed by nest predators with the hen also killed (probably by a domestic
feral cat), and 6 were abandoned for unknown reasons. All three nests thought
to have been abandoned because of hail were found in early June, 1971.

�-7-

370'

3
Agcr

(Ill)

12'

5
Agel

hi)

12'

12'

Detailed Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan 1, Job 16

Pig. 1.

Nest Site

Location of pheaaant neata in northwest quadrat, 1971.

�-8-

370'

2

o

Agc:r (Ill)

7

5

3
Agel

(Ill)

f

/
6

Mesa-Agcr

&lt;Ill&gt;

,
Br1n hll&gt;

12'

12'

Detailed

1'1g. 2.

Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan 1, Job 16

Nest Site

Location of pheasant nests in northeast quadrat, 1971.

�-9-

370'

\\

5
0

4
Ag1.n

12'

&lt;.)

7

2

/

Mesa h.)

\JIll)

f

10/
0

\ \ II /
.&gt;
.&gt;
WLiU/ .&gt;
kin

~

&lt;.&gt;

~

-.

~

-------

~

~

"--

Mesa-Agcr

~

0

6
~

o Ag1n &lt;

\\

0

7/

4

(.)

»&gt;

~,~0

»&gt;

2

0

//

=./
Agc:r

&lt;.)

36'

V

C
i
?7i\~~~~

~

3

Q46'
Water

Agcr

\\10'

/I
I

12'

6

\
I

Brin (u.)

Detailed Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan 1, Job 16

Location of pheasant nest.

0

~

5
Agel (ua)

12'

l"1ge 3.

(ua)

Nest Site

in southwest quadrat, 1971.

.,..

&lt;:)

eN

�-10-

370'

2

3

Mesa (UIl)

.\gin (m)

o

§ 4
~

Aq1n

(,,111)

12'

12'

Detailed

Pig. 4.

Arrangements of Individual Quadrat, Pheasant
Selection Study, Work Plan 1, Job 16

Nest Site

Location of pbeasant nests in southeast quadrat, 1971.

�Table 1. Number of pheasant nests found in 48 plots representing various vegetative types and treatments, 1971.
Plot
Number

Type and Treatment

NW

NE

SW

SE

Total

Number
Plots

Nests/
Plot

1

Smooth bromegrass (mowed)

0

0

1

0

1

4

0.25

1

Smo ot.h bromegrass (unmowed)

2

1

0

1

4

4

1.00

2

Alfalfa (mowed)

0

2

3

0

5

4

1.25

2

Alfalfa (unmowed)

3

-1

0

2

6

4

1.50

3

Crested wheatgrass (mowed)

0

0

0

1

1

4

0.25

Quadrats

I

3

Crested wheatgrass (unmowed)

2

1

1

0

4

4

1.00

4

Intermediate wheatgrass (mowed) 1

1

4

2

8

4

2.00

4

Intermediate wheatgrass
(unmowed)

2

0

2

1

5

4

1.25

5

Tall wheatgrass (mowed)

0

0

1

1

2

4

0.50

5

Tall wheatgrass (unmowed)

1

4

0

0

5

4

1.25

6

Alfalfa-crested wheatgrass
(mowed)

0

0

1

1

2

4

0.50

6

Alfalfa-crested wheatgrass
(unmowed)

2

1

1

1

5

4

1.25

13

11

14

10

48

48

Total
Average

1.00

t-'
t-'
I

�-12-

Table 2.

Distance of pheasant nests from plot perimeters, 1971.

Distance
From Nearest
Edge

Number of
Nests

Percent

0 - 5'

6

12.5

5 - 10'

18

37.5

10 - 15'

13

27.0

15 - 20'

3

6.3

20 - 25'

6

12.5

25 - 30'

2

4.2

30 - 35'

0

0.0

35 - 40'

0

0.0

48

100.0

Total

Periods of Nest Establishment
Table 5 lists periods of nest establishment based upon aging egg embryos
according to procedures outlined in Game Information Leaflet Number 15
(Sandfort 1965).
Only one peak in nest establishment occurred in 1971. This was during the
period May 16-31 when 17 nests out of the 48 recorded were established.
Phenology
Heights of vegetation measurements were made on April 15 and 27; May 6 and 19;
June 2, 14, and 25; and July 5 and 15, 1971. Table 6 summarizes average
heights of vegetation on April 15, May 6, June 14, and July 15, 1971. Heights
of the new growth vegetation on May 6, 1971 was at least one week ahead of that
found in early May, 1970. Suitable nesting cover was considered to be available in early May, 1971.
As found in 1970, most vegetative species continued to grow in height throughout the nesting season of 1971. Good early soil moisture has been maintained
for the 1969, 1970, and 1971 nesting season by complete plot irrigations
during the preceding fall periods.

�Table 3.

Clutch sizes found in 19 incubating or hatched pheasant nests in various vegetative types, 1971.

Clutch
Size

1
Brin
(m)

1
Brin
(urn)

2
Mesa
(m)

2
Mesa
(urn)

3
Agcr
(m)

3
Agcr
(um)

Plot Number and Treatment
4
4
5
5
Agel
Agin
Agin
Agel
(m)
( 111)
(urn)
(um)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(urn)

Total

1
2
3

4

1

1

5

2

1

3

6

1

1

7

2

8

1

1

2

1

9

1

10

1

1

11

3

1

1

2
1

12

1

13

1

1
1

2

14
15
16

1

1

17

1

1

18
19
20
Total

0

1

2

4

0

1

Ave.C1utch 0.0

8.0

7.0

9.2

0.0

10.0

4
11.8

1

1

2

1

2

19

13.0

9.0

6.0

6.0

8.5

9.1

•...
I

w
I

�Table 4.

Fate of 48 pheasant

nests

in various

vegetative

types,

1971.

Plot

Type

2

2

3

Mesa
(um)

Agcr

(urn)

~sa
(m)

3

4

4

1

1

1
Brin

1
Brin

(m)

Active
Laying or
Incubating
Hatched
Abandoned
or Unsuccessful

1

1

1

(m)

Number and Treatment
3
4
4
Agcr
Agin
Agin
(um)
(m)
(um)

5

5

Agel
(m)

Agel

5

1

3

3

1

1

1

2

(um)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(um)

3

2

4

32

1

5

1

2

1

Total

11

1

I
.....

.pI

Total

1

4

5

6

1

4

8

5

2

5

2

5

48

�Table 5.

Estimated period of nest establishment in various vegetative types, 1971.1

Period

1
Brin
(m)

1
Brin
(um)

2
Mesa
(m)

April 16-30

2
Mesa
(um)

3
Agcr
(m)

Plot Number and Treatment
3
4
4
Agcr
Agin
Agin
(um)
(m)
(um)

5
Agel
(m)

5
Agel
(um)

6
Mesa-Agcr
(m)

6
Mesa-Agcr
(um)

1

1

May 1-15

2

1

May 16-31

1

2

June 1-15

1

June 16-30

1

Total

1

1
3

2

4

1

1

2

1

5
17

2
2

3

8
2

1

I

I-'

V1

July 1-15

I

July 16-30
Unknown

1

1

Total

1

4

5

2

1

1

3

2

1

2

6

1

4

8

5

2

5

2

1

15

5

48

1 Period of nest establishment based upon aging of embryos according to Game Information Leaflet No. 15 (Sandfort 1965) .
First nest search was made June 3-4, 1971, second made June 23-28, 1971, third made July 19-21, 1971.

�-16-

Table 6.

Average height of vegetation in pheasant nesting plots, 1971.

Plot
Number

Vegetative Type
and Treatment

4-15-71

Average Height (In.)
5-6-71
6-14-71
7-15-71

1

Smooth bromegrass (um)

6

12

18

14

1

Smooth bromegrass (m)

4

8

31

30

2

Alfalfa (urn)

3

11

19

21

2

Alfalfa (m)

4

11

21

25

3

Crested wheatgrass (um)

7

9

9

15

3

Crested wheatgrass (m)

5

9

14

13

4

Intermediate wheatgrass (urn)

8

13

17

17

4

Intermediate wheat grass (m)

6

12

22

38

5

Tall wheatgrass (um)

9

14

19

17

5

Tall wheatgrass (m)

4

8

17

17

6

Alfalfa-crested wheatgrass (um)
Alfalfa

1

4

1

Crested wheatgrass

6

11

6

16

Alfalfa

3

10

18

12

Crested wheatgrass

4

10

15

22

6

Alfalfa-Crested wheatgrass (m)

Height of Vegetation Surrounding Nests
Nests found prior to windrowing of all the cover plots on July 23, 1971 provided an opportunity for obtaining vegetative height measurements surrounding
46 nests. Table 7 lists average heights and species of vegetation. An average height of 16.13 inches was found surrounding 30 nests found during the
first nest search period of June 3-10. This compares with an average vegetation height of 17.73 inches surrounding 11 nests found in the second nest
search period of June 23-28, and 18.80 inches surrounding 5 nests found in the
third nest search during July 19-23, 1971. An overall average of 16.80 inches
was calculated for vegetation surrounding the 46 nests measured.

�Table 7.

Nest
Number

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30

Heights and composition of vegetation surrounding 48 nests, 1971.

Quadrat

Plot
No.

Date
Measured

SE
SE
SE
SE
SE
SW
SW
SW
SW
SW
SW
SW
SW
NW
NW
NW
NW
NW
NW
NW
NE
NE
NE
NE
NE
NE
SW
SW
SW
SE

2-1
1-1
4-E
3-E
5-E
2-E
2-E
6-E
4-E
4-1
4-1
3-1
6-1
6-1
6-1
3-1
3-1
4-1
4-1
l-E
2-E
5-1
3-1
2-1
1-1
5-1
5-1
4-E
4-E
2-1

6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-3-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-4-71
6-9-71
6-10-71
6-10-71
6-14-71

Composition

Alfalfa
Smooth bromegrass
Intermediate ~heatgrass
Crested wheatgrass
Tall wheatgrass
Alfalfa
Alfalfa
Alfalfa-crested wheatgrass
Intermediate wheatgrass
Intermediate wheatgrass
Intermediate wheatgrass
Crested wheatgrass
Alfalfa-crested wheatgrass
Alfalfa-crested wheatgrass
Alfalfa-crested wheatgrass
Crested wheatgrass
Crested wheatgrass
Intermediate wheatgrass
Intermediate wheat grass
Smooth bromegrass
Alfalfa
Tall wheatgrass
Crested wheatgrass
Crested wheatgrass
Smooth bromegrass
Tall wheat grass
Tall wheatgrass
Intermediate wheat grass
Intermediate wheatgrass
Alfalfa

Ave. Height
(In.)

Ave. Height
For Period (In.)

15
16
15
16
14
18
18
15
22
12
13
12
17
14
13
13
16
15
14
16
20
16
12
17
16
15
15
22
27
20

I

to-'

~
I

16.13

-------------------------------------------------------------------------------------------------------------

�Table 7.

Heights and composition of vegetation surrounding 48 nests, 1971 (continued).

Nest
Number

Quadrat

SE
SE
SW
SW

Plot
No.

Date
Measured

Composition

Ave. Height
(In.)

6-23-71
6-23-71
6-24-71
6-24-71
6-24-71
6-24-71
6-24-71
6-24-71
6-28-71
6-28-71
7-5-71

Intermediate wheat grass
Alfalfa-crested wheatgrass
Intermediate wheatgrass
Smooth bromegrass
Alfalfa
Alfalfa
Tall wheatgrass
Intermediate wheatgrass
Alfalfa-crested wheatgrass
Tall wheatgrass
Alfalfa

15
19
13
17
19
19
17
20
14
20
22
16
13
19
31
15

31
32
33
34
35
36
37
38
39
40
41

NE
NE
SW

2-1
6-E
4-E
I-I
2-1
2-1
5-E
4-E
6-1
5-1
2-E

42
43
44
45
46

SE
NW
NE
NE
NE

6-1
1-E
2-E
4-E
5-1

7-19-71
7-20-71
7-21-71
7-21-71
7-21-71

Alfalfa-crested wheatgrass
Smooth bromegrass
Alfalfa
Intermediate wheat grass
Tall wheatgrass

47
48

SE
NW

4-E
2-1

7-23-71
7-23-71

Intermediate wheatgrass
Alfalfa

NW

NW
NW
NW

Ave. Height
For Period (In.)

17.73

I

18.80

Average height all measurements

I

I-'

00

16.80

�-19-

KEY TO ABBREVIATIONS
Figures 1 through 4 and Tables 3 through 5
Plants
Symbol

Scientific Name

Common Name

Agcr

Agropyron cristatum

Crested wheatgrass

Agel

Agropyron elongatum

Tall wheatgrass

Agin

Agropyron intermedium

Intermediate wheat grass

Brin

Bromus inermis

Smooth bromegrass

Mesa

Medicago sativa

Alfalfa

Treatment
Following 1970 nesting season:
(m)

Mowed

(um)

Unmowed - residual cover check in 1971

LITERATURE CITED
Sandfort, W. W. 1965. Aging pheasant embryos. Colo. Game, Fish and Parks
Dept. Outdoor Facts, Game Information Leaflet No. 15. 2 p.

Prepared by

---!tf2~L~'~
---+RV_m
" ~q~"
~m.!-/~~
__
Don~offman~
Wildlife Researcher

��-21April,

1972

JOB PROGRESS REPORT

State of

COLORADO
--------~~--------------

Project No.

W-37-R-25

Game Bird Survey

Work Plan No.

1

Job Title

pheasant Roadside

Cover Evaluation

1, 1971 to March

31, 1972

Period Covered:
Personnel:

April

Russell

Kozacek,

Job No.

Carroll

Grand Pre and Warren

18
Study

Snyder.

ABSTRACT

Abundant winter and spring moisture stimulated early spring growth and greatly
improved the quality of seeded grass and legume stands in 1971.
Fourteen
additional quarter-mile plots were seeded in late spring to conclude seeding
study efforts.
Nest searching, conducted on 67 seeded plots, 38 farmed controls
and 38 unfarmed controls, yielded 45 nesting attempts.
Of these, 12 were
successful, 22 were predator destroyed, 4 were abandoned and the fate of 7 remained uncertain.
Pheasants attempted significantly more nests in seeded plots
than in farmed controls on an acres per nest basis.
Both 1970 and 1971 data
were combined in the analysis.
The unfarmed controls yielded significantly more
nests during the two years than the farmed controls did. Discussion of nest
association with proximal cover species and adjacent field cover types are
provided.
Data concerning life-form characteristics of the study plots are
presented.

��-23-

PHEASANT ROADSIDE COVER EVALUATION STUDY
Warren D. Snyder

P. S. OBJECTIVE
To compare pheasant production under natural roadside conditions with the
following cover types to be established along roadsides: (1) grass, and
(2) grass-legume mixtures.
SEGMENT OBJECTIVES
1.

To establish grass and grass-legume cover on roadside test plots in
northeastern Colorado.

2.

To

3.

To measure life-form characteristics of roadside cover.

measure pheasant production in test and control plots.

METHODS AND MATERIALS
Methods and materials used in selection of plots, establishment of seeded
plots and measurement of pheasant production are presented in Snyder (1969,
1970, and 1971).
Measurement of Life-form Characteristics of Roadside Cover
Height and density measurements were obtained during the second week of May
and the second week of June, 1971. Ten groups of plots were included in
the sample. Fifty measurements each for height and density were obtained
per plot on 20 seeded plots, 9 unfarmed controls, and on 7 farmed controls
that contained green wheat. Five measurement sites, each containing ten
measurement points were spaced by use of vehicle speedometer readings.
The ten measurement points were obtained while walking at an angle across
the roadside plot. Measurements were taken one ~ace apart and located
six inches to one foot in front of the right foot. Vegetative height was
recorded by use of a yard stick. Density levels at heights six inches or
more above the ground were obtained by ocular estimates. A scale of 1 to
3 was used with the latter estimate being the highest.
RESULTS AND DISCUSSION
Establishment of Grass and Grass-legume Cover
Past Seedings
Seeded stands, especially those containing grass showed marked improvement
in growth. Natural spreading of the grasses was enhanced by excellent winter

�-24-

snows and spring and early summer precipitation.
Some alfalfa stands were
so rank and tangled by mid-June that nest search efforts were nearly
impossible.
Pitch forks were finally used to lift and separate this dense
growth.
The excellent growing conditions permitted an increase in the number of
seeded plots utilized in the study from 48 in 1970 to 67 in 1971.
Accomr
panyiug natural unfarmed (C2) controls and farmed (Cl) controls increased
the total number of quarter-mile plots from 99 in 1970 to 143 in 1971.
Approximately 36 linear miles or about 76 acres were included in the 1971
study results.
Seeding

Efforts

in 1971

The near continuous wet, rainy weather during the spring of 1971 curtailed
planting efforts, but helped insure good to excellent grass and/or legume
stands on those that were completed.
Fourteen additional quarter-mile plots,
separated into seven groups, will be available for nest search in 1972.
Various combinations of tall wheatgrass (Agropyron elongatum), switchgrass
(Panicum virgatum, var. Nebr. 28) and rhyzoma alfalfa (Medicago sativa) were
used in these plots.
Burnet (Sanquisorba minor), a member of the Rosacae
family, was tried in several of the plots.
Its initial growth did not
compare to that of alfalfa in the roadsides, but more time is needed for a
complete evaluation of the species.
The 1971 spring and early summer
plantings were the final attempts to establish roadside cover as part of
this study.
Measurement

of Pheasant

Production

Nest Fate
Results of 1971 nest searches were generally similar to those obtained in
1970.
Table 1 shows the number and fate of nests for both years.
Hatching
success was higher in 1971 but the number was too small to say a sigificant difference occurred.
Predators continued to destroy a large percentage
of the nests.
Nest search disturbance caused the abandonment of one nest.
Second and third checks to determine nest fate were done cautiously only
after several days had elapsed.
Nest Density

Comparisons

A compilation of statistics concerning the number of plots, average plot
size, total acreage per type, total nests, acres-per-nest and acres-persuccessful nest is illustrated in Table 2. The acres-per-nest columns best
illustrate the attractiveness of the three cover types for nesting.
Results,
which were similar for both 1970 and 1971, indicate seeded stands were
proportionally
utilized more than either unfarmed (C2) or farmed (Cl) control
plots.
Nesting attempts on the C2 controls most closely compared to those
on seeded roadside plots.
Chi-square analyses were conducted based on total nesting attempts per total
acres of each roadside cover type. Acres of each cover type were used

�-25-

instead of number of plots due to slight differences in average plot width
between seeded and C2 controls. Widths of farmed controls were considered
the same as those of the seeded plots, so their acreages were directly
proportional (Table 2).
Table 1. Fate of pheasant nests found along roadside study plots during the
summers of 1970 and 1971.
1970

1971

Number

Percent

Number

Percent

Successful

4

13.8

12

26.7

Predator destroyed

16

55.2

22

48.9

Abandoned

4

13.8

4

8.9

Unknown

5

17.2

7

15.5

Total

29

Nest Fate

45

Chi-square values for combined 1970-71 data indicate pheasant nesting
attempts in seeded plots were not significantly greater than those in
unfarmed (C2) controls (Table 3). In turn, the unfarmed plots yielded a
highly significantly greater number of nests per total acres than the farmed
(C1) controls did. Additional Chi-square values comparing seeded plots with
Cl and C2 type controls for the respective years, 1970 and 1971, are also
presented in Table 3. Similar comparisons based on numbers of successful
nests per cover type were not attempted because of inadequate sample sizes.
A higher number of nests were found in 1971 than during the previous summer.
This resulted primarily from an increase in the number of plots, not an
increase in nesting effort. In general, the results in Table 2 indicate
one nesting attempt was made along approximately every 0.5 mile of seeded
roadside, every 5 miles of farmed roadside, and every 1 mile of unfarmed
roadside.
Nest Association with Cover Types
Proximal Cover Association--Nest searches in 1970 showed that most pheasant
nests were associated with green and residual weed cover, even in the
seeded plots (Table 4) where grasses and legumes were dominant. The 1971
results, shown in the table, indicate increased use of seeded grasses and
marked decrease in use of annual forbs. This may, in part, be due to
increased spreading and growth of the seeded stands following excellent
spring moisture.

�-26Table 2. Statistics concerning roadside study plots and pheasant nesting
for 1970 and 1971.
Cover
Type

No. of
Plots

Acres/:!.! No. of
Plot
Acres

Nesting
Attempts

Acres/
Nest

Successful
Nests

Acres/
Suc. Nest

6.66

1970
Seeded

48

.5549

26.6337

22

1.21

4

Farmed
Control

28

.5549

15.5372

2

7.77

0

Unfarrned
Control 23

.5072

11.6656

5

2.33

0

53.8365

29

1.86

4

13.46

5.13

Total

99

1971
67

.5364

35.9358

32

1.12

7

Control

38

.5364

20.3832

2

10.19

0

Un farmed
Control

38

.5231

19.8768

11

1.81

5

3.98

Total

143

76.1958

45

1.69

12

6.35

Seeded
Farne d

11 The average farmed control plot contained approximately 0.14 acres of
~nfarmed cover along the road shoulder. Total plot width and acreage was
considered equal to that of seeded plots for direct comparison purposes.
Table 3. Results of Chi-square analyses comparing nesting attempts on seeded,
unfarmed (C2) controls and farmed (Cl) controls during 1970 and 1971.
Chi-square Values
1971
1970-71 Comb~ned

D. F.

1970

Seeded vs. Farmed (C1)

1

8.38**

13.52**

21. 89**

Seeded vs. Unfarmed (C2)

1

1.84

1.88

3.57

Farmed (C ) vs. Unfarmed (C2)
l

1

-- y

6.46*

8.88**

Comparison

!I Insufficient sample size.

* Significant at 0.05 level.
** Significant at 0.005 level.

�-27-

Table 4. Utilization of roadside cover types by nesting pheasants during
the summers of 1970 and 1971.

Plot Type

Cover Type

Number of Nests
1970
1971

Seeded Roadside
Seeded grass

4

9

Grass-alfalfa

1

1

Seeded alfalfa

1

1

Grass-downy brome

0

3

Alfalfa-downy brome

0

4

Straight downy brome

1

6

Grass and residual weeds

0

2

Grass, alfalfa and weeds

2

1

Grass, alfalfa and downy brome

0

2

Alfalfa, weeds and downy brome

1

0

Green weeds

0

2

Weeds and residual

11

1

Weeds, residual and downy brome

1

0

Sub-total

22

32

Unfarmed (C2) Roadside
Native grass

o

1

3

Straight downy brome

o
o

2

Downy brome and residual

1

o

Green weeds

2

1

Weeds and residual

2

2

Straight residual cover

o

2

Sub-total

5

11

Farmed (C1) Roadside
Downy brome and residual

o

1

Green weeds and residual

Native grass and downy brome

2

1

Sub-total

2

2

Total

29

45

�-28-

The winter snows and early spring moisture, which favored seeded stands,
was ideal for the growth of doWny brome or cheat grass (Bromus tectorum).
As noted in Table 4, many of the nests found in 1971 were in partial to pure
stands of downy brome. Because of its early growth and maturity characteristics it competed extensively with new seeded perennial grasses and alfalfa.
It was considered one of the primary factors suppressing seeded stand establishment. In dryer years its growth form is often too short to provide good nesting
cover, but usually it is one of the best available cover types. Farmers,
objecting to its invasion of their fields, frequently burn it as it matures
in late spring or early summer. One hen, nesting in downy brome, was observed
attempting to resume incubation after her nest had been burned over in July
1971.
Downy brome can potentially be controlled by use of the chemical, Dacthal.
Application must be made prior to summer or early fall germination of the
seed, but tests probably should be run to determine the best time and amounts
to use in application.

Relationship to Adjacent Field Cover Type--Nest occurrences were compared to
plot occurrences adjacent to summer fallow, green wheat, and the combination
of other field types, such as corn, grain sorghum, etc. Combined 1970 and
1971 data show that 35 nests were found in 100 plots adjacent to fallow; 24
nests were found in 106 plots adjacent to green wheat and 14 nests were
found in 36 plots adjacent to other cover. The data do not show any clearcut evidence that pheasants selected nesting sites with regard to adjacent
field cover types. The presence of wheat stubble, instead of fallow, at
the onset of some nesting attempts further complicates the issue.
Life-form Characteristics of Roadside Cover
An attempt was made to index growth characteristics of seeded roadsides,
unfarmed (C2) controls and farmed (Cl) controls containing green wheat.
Three indices, height, density and height X density were obtained during May
and again in June. Results of 1971 measurements, summarized in Table 5,
indicate heights of seeded stands generally exceeded those of unfarmed (C2)
roadsides. As would be expected, green wheat in farmed controls exceeded
the other two cover types in height.
The farmed and seeded roadside covers were similar in density according to
ocular estimates. Cover in unfarmed roadsides was somewhat less dense. Some
of the unfarmed plots contained dense matted stands of kochia or other cover
which were suppressed in height because of competition for moisture. These
were given a lower index rating because the cover was not high enough to hide
a nesting hen pheasant.
Question arises as to why did pheasants fail to utilize the farmed controls
if height and density characteristics were similar to those on the seeded
plots? One reason to consider is that 15 of the 38 Cl plots were summer
fallowed to the road shoulder. This greatly reduced potential nesting cover.
In addition, green wheat stands lacked litter and old vegetation close to
the ground. Pheasant nest bowls usually are formed of litter and frequently
are positioned so that residual vegetation breaks up the outline of the hen.
Additional life-form variables must be measured before attempting to relate
nesting attempts to vegetative life-form.

�-29-

Table 5. Height and density measurements and a product index of the two
comparing vegetative growth characteristics on samples of seeded roadsides,
unfarmed natural roadsides, and farmed roadsides containing green wheat.

Date

Seeded
Roadside

Unfarmed
Natural

Farmed
(Green wheat)

Height
May, 1971

8.71

6.08

14.18

June, 1971

16.31

13.06

26.72

Density
May

1.65

1.35

1.66

June

2.23

2.01

2.19

Height X Density
May

14.93

8.33

23.99

June

37.19

27.31

58.98

Sample Size;!!
May

20

9

8

June

20

9

6

!/

Number of plots sampled.
obtained on each plot.

Fifty height and density measurements were

LITERATURE CITED
Snyder, W. D. 1969. Pheasant roadside cover evaluation study. Colo. Div.
of Game, Fish and Parks. Game Research Report. April. pp. 37-45.
1970. Pheasant roadside cover evaluation study. Colo. Div. of Game,
Fish and Parks. Game Research Report. April. pp. 109-114.
1971. Pheasant roadside cover evaluation study. Colo. Div. of Game,
Fish and Parks. Game Research Report. April. pp. 27-39.

��-31-

JOB PROGRESS

S ta te of
Project

April,

1972

REPORT

C.::.O:;,;L=.,O:;,;RA=D::..;O::.-_

3

Job No.
8a
Effects of Sagebrush Control
on Distribution and Abundance of Sage Grouse

Work Plan No.
Job Title
Period Covered:
Personnel:

Game Bird Survey

W-37-R-25

No.

April

1, 1971 to September

30, 1971

Warren Snyder, Russell Kozacek, Don Gore, Courney
Ellenberger, John Monarch and Howard Funk.

Crawford,

John

ABSTRACT

The peak periods of sage grouse strutting ground activities in the North Park
study area again centered around the middle of April, 1971.
There were increases
in numbers of males observed on almost every ground with the total of all high
counts on each ground being 326, as compared to 274 in 1970. Data from check
stations operated the first two days of season at Walden and Cowdrey suggested a
decrease in hunter participation
in 1971 with only 357 hunters tallied as compared
to 564 the previous year.
Number of sage grouse bagged by these hunters was 264,
which was similar to the year before, despite the lower number of hunters.
About
49 percent of the bag was composed of immature birds, which was up slightly from
1970 and about average over the years.

��-33-

EFFECTS OF SAGEBRUSH CONTROL
ON DISTRIBUTION AND ABUNDANCE OF SAGE GROUSE
Howard D. Funk

An investigation of sage grouse abundance and distribution near Lake John
in North Park has been in effect since initiation in 1963. The job relates
to various specific study topics in various degrees of intensity. More
recent study has been with regard to the immediate effects of a sagebrush
spraying project, accomplished in 1965, on sage grouse populations with a
study on the long-term effects of this spray project scheduled for a twoyear period beginning the spring of 1973. In the interim period, efforts
are to be limited to collection of study area strutting ground data,
banding samples of birds on these grounds, and operating check stations near
Walden and Cowdrey to obtain harvest data. This report covers these items.
P. S. OBJECTIVE
To determine the effects of 2,4-D spraying of sagebrush on: (1) sage grouse
abundance, (2) sage grouse distribution, and (3) vegetative composition and
density.
SEGMENT OBJECTIVES
1.

To investigate the seasonal di~tribution of sage grouse after sagebrush
is controlled.

2.

To investigate sage grouse numbers by age and sex classes on and adjacent
to the area treated.

METHODS AND MATERIALS
Strutting Ground Counts
Wildlife Conservation Officers conduct counts on a number of strutting
grounds in North Park annually. Usually, three early morning counts are made
on each ground. Data from these counts are gathered from the Regional
Biologist, high counts for each ground being reported herein.
Check Stations
Two check stations near Walden and Cowdrey are operated the first two days of
the season to collect various data on harvest such as harvest success, age,
and sex ratios, banded bird information, and hunter participation for a check
on the year, in comparison with previous years. Project personnel are

�-34-

responsible for the station, with Conservation Officers also contributing
data from field checks. Main emphasis is on checking harvest within the
study area. Age and sex of birds are determined by methods described in
previous reports.
RESULTS AND DISCUSSION
Strutting Ground Counts
Results of strutting ground counts are displayed in Table 1. Peak counts
occurred about mid-April on most areas, which is about average, since
strutting ground activity is subject to snow conditions in this area. Data
on peak numbers of females and total birds were not gathered for this report,
but should be obtained from the field men for continuity of data when preparing for the final report.
Table 1.

Peak counts of male sage grouse on strutting grounds, 1971.

Strutting Ground 1:./

High Count Date

10

April 15

'1:./

SG

1

SG

2

SG

3

SG

4

88

April 24

SG

5

81

April 15

SG

6

25

April 12

SG

Total

Number Males

!:.l

7 '1:./

SG

8

5

April 15

SG

9

96

April 13

SG

10

21

April 23

SG

11

0

SG

12 Y
326

1/ Locations of grounds shown in .previous reports.
2/ Counts discontinued~

•

�-35-

Comparative data on annual fluctuations and high counts by strutting ground
are presented in Table 2. }bst grounds showed increases over 1970 in numbers
of males present, with the total number of males on all grounds increasing
about 19 percent over 1970 results • Thus, the trend has definitely been
upward since the recent low in 1965 when only 166 males were observed on all
grounds.
Table 2.
grouse.

Comparison of 1960 through 1971 strutting ground counts of male sage

Strutting
Ground
1960

1961

1962

1963

1964

Year
1965 1966

1967

1968

1969

1970

1971

SG

1

10

5

0

0

0

0

0

0

0

0

0

y

SG

2

43

4

7

20

26

9

9

10

2

8

8

10

SG

3

14

0

0

0

0

0

0

0

0

0

0

y

SG

4

130

96

219

216

120

53

47

69

58

82

66

88

SG

5

19

2

52

17

65

52

54

59

52

82

57

81

SG

6

5

1

8

9

3

0

2

12

36

36

19

25

SG

7

0

7

5

0

0

0

0

1

3

2

0

Y

SG

8

5

0

0

0

0

0

6

5

2

6

7

5

SG

9

167

109

71

85

99

52

97

81

71

117

97

96

SG

10

17

77

28

19

17

0

6

9

0

16

16

21

SG

11

12

50

5

11

7

0

13

15

6

5

4

0

SG

12

14

12

0

0

0

0

0

0

1

3

0

Y

Total

446

363

395

377

337

166

234

265

231

357

274

326

Y Count discontinued.
Banding
Due to lack of manpower and available time, banding of samples of birds on
strutting grounds again was not accomplished.

�Table 3.

Comparison of North Park sage grouse hunter check information, 1963-1971. !!

Hunters Hours
Checked Hunted

Adult
Males

Adult Percent
Females Adults

Juvenile
Males

Juvenile Percent
Females Juveniles

Total
Birds

Birds
Per
Hunter

Hours
Hunted
Per Bird

Year

Bag
Limit

1963

3

492

2,460

62

150

42

113

181

58

506

1.03

4.86

1964

2

217

624

25

81

59

28

45

41

179

.82

3.49

1965 'l:..,/

2

150

626

27

30

49

26

26

51

116

.77

5.40

1966

2

306

1,227

31

116

56

45

71

44

263

.86

4.67
I

1967

2

300

1,177

50

127

67

42

46

33

267

.89

4.41

1968

2

546

2,604

80

135

42

156

141

58

512

.94

5.09

1969

2

662

2,936

121

277

70

57

114

30

569

.86

5.16

1970

2

564

2,617

58

89

55

53

66

45

266

.47

9.84

1971

2

357

1,802

28

67

51

54

39

49

264

.74

6.83

!! Based on Walden and Cowdrey check station data during opening weekends.
'l:..,/ A one day season; hence only a one day check station.

w

0\
I

�-37-

Check Station Results
Project personnel again manned the check station for the first two days of
season. There were 357 hunters checked through the period, which was the
lowest total since 1967, and down sharply from the 564 hunters in 1970
(Table 3). However; 264 sage grouse were checked, which was almost equal to
the 1970 total, thui producing a higher bird-per-hunter ratio than that in
1970 (.74 compared to .47 birds-per-hunter).
Also, amount of time spent
hunting per bird in the bag was lower in 1971 (6.83 hours) than in 1970
(9.84 hours), but remained higher than the average of previous years. Juvenile
birds increased to 49 percent of the total bag, 'a slight increase over the
previous year, but these ratios are not considered good estimates of the
actual age ratio in the population.

Prepared by _~~:::.-..::::..".::::~:..:~==-:;;;::';;;...;:,;~~~~.....;'
:;..;~=..:..::::'=-_
Howard D. Funk
Section Chief, Small Game Research

��-39April,

JOB PROGRESS

State of

~CO~L~O~RA~D~O~

Project No.

_

10

Job Title

Game Bird Survey
Job No.

Study of Hungarian

Period Covered:
Personnel:

REPORT

W~-~3~7_-R~-2~5~ _

Work Plan No.

April

1972

1, 1971 to March

Partridge

1
Adaptability

31, 1972

Carroll J. Grand Pre, John F. Corey, George G. Harrington, Gordon P.
East, Daniel F. Potts, Walter H. Schuett, Robert V. Clark and Donald
M. Hoffman.

ABSTRACT
Only four reports of isolated sightings of from two to four Hungarian partridges
were recorded for Moffat County in Segment 25. These were observed in the spring
of 1971. A total of 1,774 Huns were released in Moffat County from 1964 through
1970.
Six hundred and fifty-five pen raised Hungarian partridges were released at three
sites in the San Luis Valley on April 23, 1971. This constitutes the first of a
planned three-year stocking effort to attempt establishment of the species in
south-central Colorado.
A total of seven broods ranging in size from one to fourteen chicks were observed
by Project personnel or interested landowners on or near the three release sites
in the San Luis Valley in August, 1971. In addition, one nest containing eight
eggs was destroyed by a farm tractor and one hatched nest with twelve eggs was
found by Project personnel.
Pairs of Huns were reported at distances of four
to eight miles from two of the release sites but pairs are known to have remained
near all three release sites as well.
A minimum of 28 Huns was known to be
ranging in the vicinity of the three release sites during the period December
9-11, 1971. One flock of 16 was observed l~ miles west of Hooper, Colorado
on December 11, 1971.

�-40-

RECOMMENDATIONS

Trial field releases are recommended for the San Luis Valley in southcentral Colorado for two more years.
Intensive field surveys will
also be required to evaluate these stocking efforts.

�-41-

STUDY OF HUNGARIAN PARTRIDGE ADAPTABILITY
Donald M. Hoffman

P. S. OBJECTIVE
To determine the ability of Hungarian partridges to establish reproducing
populations in mixed brushland-wheatland habitat in selected sites in
Colorado.

SEGMENT OBJECTIVES
1.

Introduce Hungarian partridges.

2.

Determine presence and dispersal of Huns and success of previous plants.

METHODS AND MATERIALS
On April 23, 1971, 655 pen raised Hungarian partridges from holding pens
at the Fort Collins Wildlife Research Station and the Rocky Ford Bird Farm
were released at 3 sites in the San Luis Valley.
Intensive field searches of all release sites in the San Luis Valley were
made by Project personnel during the periods June 7-11, July 14-15, August
2-6, September 13-16, and December 9-11, 1971. Persons living or working
in or near the release sites were interviewed and observations were recorded.
Reports of Huns from Moffat County were investigated during the period June
29-July 2, 1971.

RESULTS AND DISCUSSION
Table 1 summarizes the 1971 field releases of Hungarian partridges. These
were made at 3 sites in the San Luis Valley on April 23, 1971, and consisted
of a total of 655 Huns. Figure 1 shows a release on 'the Robert Young property northeast of Alamosa, Colorado. Figures 2, 3, and 4 show the map
locations and numbers of Huns released in 1971.
Table 2 summarizes Hungarian partridge reports and observations from Moffat
County during Segment 25. Isolated sightings of 2 to 4 adults were reported
in the spring of 1971, indicating only a few have survived in northwestern
Colorado.

�-42-

Table 1.

1/
Hungarian partridge field releases, San Luis Valley, 1971.Number
Released

Source of
Stock

April 23, 1971

286

FCWRS
(pen raised)

Alamosa County, Robert
Young farm. Sec. 7,
T38N, RIlE.

April 23, 1971

200

FCWRS
(pen raised)

Conejos County, Hot
Creek Management Area.
Sec. l5~ T35N, R7E.

April 23, 1971

169

FCWRS
(pen raised)

Saguache County, Ernest
Stoeber farm. Sec. 32,
T4lN, RIOE.

Total

655

Date

Release Area

1/
- See Job Progress Report for FA Project W-37-R, Work Plan 10, Job 1, April,
1971 for field releases made in Larimer, Moffat·, and Routt counties from 1964
through 1970 and sources of stock.

Table 2. Summary of Hungarian partridge reports and observations, Moffat
County, 1971.
Period

Number of Huns

Area

Observed By

April-May, 1971

2 (pair)

Two miles north of Highway 13 on Milk Creek.

G. Hiltoll

April, 1971

3

Three miles north of Lay
Postoffice.

W. Roland

April-May, 1971

4

Along Highway 13 on south- G. Hilton
west side of Isles Mountain.

Several

Two miles east of Gossard
L. Kendall
Ranch release site on Wagner
Ranch.

Spring, 1971

�I

+:-

w
I

Fig. 1. Hungarian partridge release, Robert Young farm, Alamosa County, April 23, 1971.
(D. Hoffman photo).

�II

••

~2=+-6-4-----",25:"'-"'1,,----30

\

2.

2.

29

2.

~

28

~~ f.. ..

i(~

.~

i
~.

3.

32

32

3'

II

--.,\0_'0

LocatiOn """

..-ra

t;~===Al\ K

~

Hungar10n partrl.dge

nl

I~.

\)

I

I
1

~,\

i
~

---lL~.1.': .,:. ~:

of!

3

/~

:

i

3!5.36

3.

r---f..-------.........

L-+----l--L-I

F1g. 4.

33

(

'II

"_

\~~¥ '

sed in Saguache County,

1971..\

I

I

,

i'

•••

-&lt;"J

R9E.
A
.-- .. -

...

~

;
~

L

A

-----..

...- ....

~

RIOE.
M
0

s

A

�-47-

Table 3 summarizes reports and observations in the San Luis Valley
following the 1971 field releases. Pairs were formed shortly after
releasing from the crates and some remained on or near the release
sites selected. Movements of pairs of Huns were also recorded at
distances of 4 to 8 miles from the Robert Young and Ernest Stoeber
release sites. Pairs released on the Hot Creek Management Unit probably moved less than a mile, based upon field observations.
Six
broods were observed by Project personnel and another brood was reported by an interested landowner. All broods were on or near the
release sites. One nest with 8 eggs was reportedly destroyed by a
farmer in farming operations and 1 hatched nest with 12 eggs was
observed by Project personnel. This is the highest number of broods
and numbers of nests found in anyone year since field releases were
started in 1964. Broods began to appear during early August, 1971
and ranged in size from 1 to 14 chicks.
In September, the young could no longer be readily distinguished from
the adults. Groups of 1 to 6 birds were observed in 2 of the 3 release sites during this period.
At least 28 Huns were known to be ranging in the vicinity of the 3
release sites during the period December 9-11, 1971. One flock of
16 was observed 1-1/2 miles west of Hooper, Colorado on December 11,
1971. Tracks of 2 groups of 2 and 4 were observed on the Hot Creek
Management Unit during this period and a report of a group of approximately 6 Huns'was secured from the vicinity of one of the Robert Young
releases.

Prepared by

£~JtYi¥~'~
Wildlife Researcher

�Table 3.

Summary of Hungarian partridge reports and observations, San Luis Valley, 1971.
Observed By

Date

Number of Huns

Area

May, 1971

Several pairs

Area from Mosca to U. S. Highway
160 (from R. Young releases)

G. East and area
residents

May, 1971

Several pairs

Area from 4 miles west of Hooper
to town of Hooper (from E. Stoeber
release)

E. Stoeber and area
residents

May 26, 1971

Several

R. Young release site

R. Young

Early June, 1971

1 nest with 8 eggs
(destroyed by tractor)

E. Stoeber release site

E. Stoeber

June 8, 1971

4 pair

R. Young release site

R. Young's hired hand

I

~

00
I

June 9, 1971

7 pair and 2 singles

Vicinity of Hot Creek Management
Area release sites

D. Hoffman

June 9, 1971

5 pair and 3 singles

Vicinity of E. Stoeber release site

D. Hoffman, E. Stoeber
and sons

June 10, 1971

5 pair

Vicinity of Hot Creek Management
Area release sites

D. Hoffman

July 15, 1971

2 pair and 3 singles

Vicinity of Hot Creek Management
Area release sites

D. Hoffman

August 3, 1971

1 pair with 3 chicks

1 mi. west of E. Stoeber release site

C. Grand Pre

August 3, 1971

1 pair with 14 chicks

1-1/2 mi. west of E. Stoeber release
site

C. Grand Pre

August 4, 1971

1 pair with 10 chicks

1 mile southwest of first release site,
R. Young farm

C. Grand Pre

-------------------------------------------------------------------------------------------------------------

�Table 3.

Summary of Hungarian partridge reports and observations,

Date

Number of Huns

San Luis Valley, 1971 (continued).

Area

Observed By

August 4, 1971

1 pair

Vicinity of second release site,
R. Young farm

C. Grand Pre

August 4, 1971

1 pair

1/2 mi. northwest of second release
site, R. Young farm

C. Grand Pre

August 4, 1971

1 nest with 12
hatched eggs

1 mile west of first release site,
R. Young farm

C. Grand Pre

August 4, 1971

1 pair with 1 chick

1-1/2 miles west of first release
site, R. Young farm

C. Grand Pre

August 5, 1971

1 pair with 10 chicks

1/2 mi. southwest of first release
site, Hot Creek Management Area

C. Grand Pre and W.
Schuett

August 5, 1971

1 pair

1/4 mi. east of first release site,
Hot Creek Management Area

C. Grand Pre

August 5, 1971

1 pair with 3 chicks

1/2 mi. south of first release site,
Hot Creek Management Area

C. Grand Pre and R.
Clark

August 5, 1971

2 pair

Vicinity of first release site, Hot
Creek Management Area

C. Grand Pre

August 5, 1971

1

1/2 mi. south of first release site,
Hot Creek Management Area

C. Grand Pre

August 5, 1971

3

Vicinity of first release site, Hot
Creek Management Area

c. Grand Pre

August, 1971

1 pair with 8 chicks

Vicinity of Rogers headquarters,
southwest of Hooper

Rogers

September 14, 1971

1

Vicinity of first release site, Hot
Creek Management Area

1 mi.

C. Grand Pre

-------------------------------------------------------------------------------------------------------------

I
.po

\0
I

�Table 3.

Summary of Hungarian partridge reports and observations,

Date

Number of Huns

San Luis Valley, 1971 (continued).

Area

Observed By

September 14, 1971

5

1/4 mi. southwest of first release site,
Hot Creek Management Area

C. Grand Pre

September 14, 1971

1

Vicinity of second release site, Hot
Creek Management Area

C. Grand Pre

September 14, 1971

4

1/4 mi. west of first release site,
Hot Creek Management Area

C. Grand Pre

September 14, 1971

4

1/4 mi. west of first release site,
Hot Creek Management Area

C. Grand Pre

September 15, 1971

6

1/2 mi. southwest of first release
site, R. Young farm

C. Grand Pre

September 15, 1971

4

1/2 mi. west of second release site,
R. Young farm

C. Grand Pre

September 15, 1971

1

Vicinity of second re1ease.I'3Lte,R.
Young farm

C. Grand Pre

September 15, 1971

1

1 mi. west of first release site, R.
Young farm

C. Grand Pre

December, 1971

1 group of 6 Huns

Vicinity of R. Young's south head~
quarters

R. Young's hired hand

December 11, 1971

Tracks of groups of
2 and 4 Huns

Vicinity of first release site, Hot
Creek Management Area

D. Hoffman

December 11, 1971

1 group of 16 Huns

Abandoned farmhouse 1/4 mi. southwest
of release site, E. Stoeber farm

D. Hoffman

,

I.n

o
,

�-Sl-

April,

1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-37-R-2S

Game Bird Survey

Work Plan No.

10

Job No

Job Title

Experimental

Breeding

Period Covered:
Personnel:

2

o

of Hungarian

Partridge

April 1, 1971 to March 31, 1972

Lawrence A. Webster, John F. Corey, Robert L. schmidt, George
Harrington, Larry R. Crooks, Carroll J. Grand Pre, Donald M.
Hoffman and Dr. Harry D. Muller.

G.

ABSTRACT
In a test in which egg production from six pairs of Hungarian partridge breeders
fed a standard 22 percent protein level game bird breeder feed was compared with
an equal number of pairs fed a similar feed but containing 16 percent protein,
results indicated the 22 percent protein was the better of the two based upon
total numbers of eggs produced (114 compared to 79), numbers of fertile eggs
produced (82 compared to 69) and numbers of chicks hatched (70 compared to 62).
Survival for the first seven days was excellent on both levels of feed.
Thus,
results were opposite those obtained in 1970, when the 16 percent protein level
feed appeared to be the better of the two.
In another test which compared numbers of fertile eggs produced by three pairs
of Huns receiving two extra hours of artificial light in the morning, three pairs
receiving two extra hours in the evening, and three pairs receiving no artificial
light (controls), it was found that results were similar to tests run in 1970.
The artificial lighting resulted in bringing breeder pairs into production earlier
than those receiving no artificial lighting but total eggs produced were not
increased using the artificial lighting.
In a series of tests in which over-winter survival was compared with various
bird densities in holding pens during 1970-71, it was found that excellent survival resulted with densities varying from 41.16 square feet of ground space
per bird to SOl.24 square feet of ground space per bird.
Similar results were
obtained in tests completed in March, 1972 with densities varying from 13.0S
square feet of gro~nd space per bird to 109.7 square feet of ground space per
bird.
However, hysteria was observed in a group of 73 Huns held in the pen with

(Abstract continued

on reverse

side)

�-52-

only 13.05 square feet of ground space per bird from the time they were
approximately 7 weeks of age until they were 15 weeks of age. Over-winter
survival in this particular group of birds was 100 percent, indicating they
become accustomed to the crowded condition, and the hysteria did not result
in losses.
In comparing total eggs produced by six pairs each of various ages and
sources of breeders, it was found that six pairs of 1969-hatched highest
producing adults held over from 1970 tests produced the highest number of
total eggs (213), followed by 1970-hatched run-of-the-mill pairs (199), and
1970-hatched females of pure Winchester Farms stock crossed with 1970-hatched
males from pens three to nine (109). In considering numbers of fertile eggs
produced, the 1970-hatched run-of-the-mill pairs with 175 fertile eggs ranked
highest, followed by 1969-hatched adults with 160, and 1970-hatched Winchester
Farms females crossed with 1970-hatched males from pens three or nine (96).

RECOMMENDATIONS
Based upon tests results comparing various protein levels in the breeder
it is recommended that the standard 22 percent protein game bird breeder
be used in the future.

feed,
feed

One year's data have been collected comparing fertile egg production and hatchability of eggs by source and ages of breeders.
It is recommended that these
selection tests be continued in 1972. Two years' data have been collected
in
determining over-winter survival Cif Huns in holding pens with various bird densities.
Additional tests are needed to compare survival with higher densities of
birds than used in tests completed in April, 1971 and March, 1972.
With completion of field work connected with a pheasant nesting study, it is
recommended that 12 pairs of wing-clipped 1971 hatched breeders be placed in
one quadrat and 15 pairs of similar Huns be placed in a second quadrat in April,
1972. This will allow for an evaluation of rearing young Hungarian partridges
under semi-natural conditions in these large open topped quadrats.

�-53-

EXPERIMENTAL BREEDING OF HUNGARIAN PARTRIDGE
Donald M. Hoffman

Total egg production from the 51 pairs of Hungarian partridge breeders in 1971
was 70 eggs higher than the total produced by 52 pairs of breeders in 1970
(1,217 eggs in 1971 compared with 1,147 in 1970). The 530 pen raised Huns
released in the San Luis Valley on April 25, 1972, plus 58 extra Huns held for
breeders in 1972 tests (total 588) is 67 birds less that the 655 released in
1971. This lower number of young Huns produced for field releases in 1972
resulted from lower percentage hatch of all eggs set (59.65% in 1971, compared
with 68.08% in 1970) and lower percentage hatch of fertile eggs set (78.21%
in 1971 compared with 81.31% in 1970).
P. S. OBJECTIVE
To develop game farm production techniques for Hungarian partridge.
SEGMENT OBJECTIVES
1.

To measure the effects of 2 levels of protein in the breeder feed (16%
versus 22%) on:
(a)
(b)
(c)
(d)

Egg production
Egg fertility
Egg hatchabili ty
Chick survival for the first 7 days

2.

To compare the numbers of fertile eggs produced by breeders subjected to
two extra hours of light each 24 hour period during (1) the early morning,
and (2) the evening, with breeders receiving no artificial light.

3.

To compare the numbers of surviving Huns held overwinter in holding pens
with various bird densities.

4.

To compare the numbers of fertile eggs, and hatchability of eggs produced
by breeders, consisting of:
(a)

Six pairs of 1969 hatched adults which were the highest producing
pairs in 1970.
(b) Six pairs of 1970 hatched progeny consisting of Winchester Farms
strain females crossed with males from highest producing pairs
in 1970 (pen numbers 3 or 9).
(c) Six pairs of 1970 hatched progeny consisting of females from a
high producing pair in 1970 (pen number 31) crossed with males
from highest producing pairs in 1970 (pen numbers 3 or 9).

�-54-

(d)

Six pairs of 1970 hatched progeny consisting of run-of-the-mill
females and males (1970 first hatch minus progeny from highest
producing pens).

METHODS AND MATERIALS
The detailed arrangement of the Hungarian partridge breeding pens for 1971
tests is shown in Figure 1. Individual marked birds of desired sources and
ages needed for selection tests were placed in separate pens and allowed to
choose mates before placing in the various breeding pens. Mated pairs of runof-the-mill Huns were selected and captured in the large holding pens (Fig. 2).
Table 1 lists the pen schedule for the various tests run in 1971. One mated
pair of Hungarian partridge was used in each pen. Pens 1 through 36 are located in a converted nursery shadehouse with ground floors, and pens 37 through
51 are a separate series of wire floored experimental breeding pens.
Protein Levels in Feed
Pens 7 through 18 (12 pens) were used to compare the effects of 2 levels of
protein in the feed. All females used in this test were from 1970 hatched
progeny from highest producing pens 3 or 9 and all males were of pure Winchester Farms stock. Pairs in even numbered pens were fed a standard game
bird breeder feed containing 22 percent protein and those in odd numbered
pens were fed a similar feed, but containing 16 percent protein. Eggs were
marked and placed in separate incubator trays to facilitate determination of
fertile eggs produced and numbers of fertile eggs that hatched. Chicks were
held separately in the brooder until they were 7 days of age to detect
differences in survival.
Extra Lighting
The wire floored series of pens were again used for extra lighting tests to
recheck results obtained in 1970. U. S. Weather Bureau sunrise-sunset tables
were again used to determine times when artificial lights were turned on and
off.
Pens 46-48 (3 pens) were lighted for 2 extra hours in the early mornings and
pens 49-51 (3 pens) were lighted for 2 extra hours in the evening. Starting
on March 12, 1971, pens 40-42 (3 pens) were used for controls and were not
lighted artificially. Sixteen percent protein game bird breeder feed was used
in all pens. All birds used in these tests were 1970 hatched run-of-the-mill
stock. Records were maintained of dates of first eggs, numbers of eggs laid,
numbers of fertile eggs laid, and hatchability of eggs.
Survi val in Holding Pens
Numbers of Huns placed in holding pens at the Fort Collins Wildlife Research
Station and at the Rocky Ford Game Farm were recorded by dates in 1970 and
1971. An accurate count of birds recovered by pens was made and percentage
survival for the 1970-71 and 1971-72 fall and winter periods was calculated.

�-55-

N

IJ
v32 33~1.'34 35 11/36 1

~

75'

6'4"

l' ~

f
If)

."
Ground':floored

Series

3738 39 4041 42 43 4445 46 47 4S

.......
'v_'-- ,l.--_'-- 'v _

6'4'

A- 2

\v _'--\ v_ ,--' ~-

19'
Wire-floored

Series

l-

"L..//-----'---i

(\J
(\J

49 ~

-'

5_'

..L.-..JII

(\J

en

SCALE 8

,

I

I

I

0' 5' 10'

25'

A-3 / Q)
FIG.I.
Oetailed ArranQement of
PartridQe

/-

o

-T
CD
t--

16' ----0

If-----

.l.

L-16"----j

48'------ti

BreedinQ Pens.

HunQarian

�-56Covered Shelter

1

Brooder Houses

/~
3

·0

'"
4

4'

6

f

J
4t-8'~

4

2

N

-

.'
o

I()

C\I

C\I
C\I

SCALE

~----~
"'15.5 -

~4'.•

----~----~

o

,•...
15.5-1

---35 ----------I05~-·---

15

30

-- •I

Fig. 2.

Hungarian

Partridge

Holdino Pens,

Fort Collins Wildlife Research Station

�Table 1.
Pen
Ntnnber
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30

Pen schedule for Hungarian partridge,
Protein Level
22%
16%

x
x
x
x
x
x

T
T
T
T
T
T
T
T
T
T
T
T
x
x
x
x
x
x

x
x
x
x
x
x

1971.

Artificial Light
AM

PM

None

Hatch

x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x

1969(1)
1970(1)
1970(1)
1970(1)
1969(1)
1970(1)
1970(1)
1970(1)
1970 (1)
1970(1)
1970(1)
1970(1)
1970 (1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970 (1)
1969(1)
1970(1)
1970(1)
1970 (1)
1969(1)
1970(1)
1970 (1)
1970(1)
1969(1)
1970(1)

Source

Highest producing adults
F-Win; M- 3 or 9
F- 31; M- 3 or 9
Run-of-the-mill
Highest producing adults
F-Win; M- 3 or 9
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 3 or 9; M- Win.
F- 31; M- 3 or 9
Run-of-the-mill
Highest producing adults
F- Win; M- 3 or 9
F- 31; M- 3 or 9
Run-of-the-mill
Highest producing adults
F- Win; M- 3 or 9
F- 31; M- 3 or 9
Run-of-the-mill
Highest producing adults
F- Win; M- 3 or 9

------------------------------------------------------------------------------------------------------------

I
U1

-....J
I

�Table 1.

Pen schedule for Hungarian partridge, 1971 (continued).
Protein Level
16%
22%

Number
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51

x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x

Number Pens
on Test
Key:

6

T = Pen on Test.

6

AM

Artificial Light
PM
None

x
x
x
x
x
x
x
x
x

T
T
T
x
x
x

T
T
T
T
T
T

3

x = Pen not on Test.

3

3

Hatch

Source

1970(1)
1970(1)
1969 (1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)
1970(1)

F- 31; M- 3 or 9
Run-of-the-mill
Highest producing adults
F- Win; M- 3 or 9
F- 31; M- 3 or 9
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill
Run-of-the-mill

I

VI
00
I

�-59-

Egg Production

and Hatchability

by Source

and Age of Breeders

Pens 1-6 and 19-36 (24 pens) were used to measure egg production and hatchability of eggs by various sources and ages of breeders.
Six mated pairs each
of (1) 1969 hatched highest producing adults held over from 1970 tests, (2)
1970 hatched females from pure Winchester Farms stock crossed with 1970
hatched males from highest producing pens 3 or 9, (3) 1970 hatched females
from high producing pen 31 crossed with 1970 hatched males from highest producing pens 3 or 9, and (4) 1970 hatched females and males of run-of-the-mill
stock.
These were alternated throughout the 24 pens to reduce or eliminate
effect of pen position in so far as possible.
All pens were fed a standard
16 percent protein game bird breeder feed and were treated alike as much as
possible.
For comparison purposes, similar data for 6 pens with 1970 hatched females
from highest producing pens 3 or 9 and males from pure Winchester Farms stock
on 16 percent protein feed in the Protein Levels in Feed Test series were also
calculated.

RESULTS AND DISCUSSION
Protein

Levels in Feed

In 1970, a total of 122 more eggs (337 compared with 215) were laid by 12 pairs
of breeders on 16 percent protein feed in comparison with 12 pairs on 22 percent protein feed. A total of 135 more fertile eggs (293 compared with 158)
were laid by the breeders using 16 percent protein feed in comparison with the
22 percent protein feed. This difference was significant at the P&lt;O.lO level.
Table 2 lists a comparison of numbers of eggs produced and percentage hatch
of eggs from breeder pairs in 6 pens using a 16 percent protein level game
bird breeder feed with 6 pens using a standard 22 percent protein level feed
in 1971. A total of 35 more total eggs (114 compared with 79) were laid by
the breeders fed the 22 percent protein feed.
In addition, 13 more fertile
eggs (82 compared with 69) were laid by the breeders on 22 percent protein
feed and 70 eggs from pens fed the 22 percent protein feed hatched compared
with 62 from the 16 percent protein feed pens.
Excellent survival for the
first seven days was found for both the 16 and 22 percent level feed chicks
(96.77 percent survival for the 16 percent feed, and 98.46 percent survival
for the 22 percent feed). These results are listed in Table 3.
Results obtained in the protein level tests in 1971 were therefore quite
different from the 1970 tests.
In 1970 the 16 percent protein level feed was
much better than the 22 percent protein level feed and in 1971 the 22 percent
protein level feed showed better results than the 16 percent protein level
feed. The standard 22 percent protein level game bird breeder feed will be
used for all pens in 1972 tests.
Extra Lighting
A comparison of numbers of eggs laid, numbers of fertile eggs, hatchability
of eggs, and numbers of chicks hatched by 3 pairs of Huns under artificial lights
for 2 extra hours in the morning, 3 pairs under 2 extra hours in the evening,
and 3 pairs without artificial lighting (controls) is listed in Table 4.
Generally, results in 1971 were similar to 1970 tests.

�Table 2. Numbers of eggs produced and hatchability
protein in the feed, 1971.

of eggs laid by Hungarian partridge on two levels of

Protein
Level

Number
Pens

Number
Eggs
Produced

Number
Infertile
Eggs

Number
Fertile
Eggs

Percent
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

16%

6

79

10

69

87.34

62

78.48

22%

6

114

32

82

71.92

70

61. 40

Table 3.

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germ

Number
Culls

89.85

2

5

10

85.36

2

10

23

Survival of Hungarian partridge on two levels of protein in the feed, 1971.

,
0-

,

0

Protein
Level

Number
Pens

Number Chicks Taken
to Brooder House or
Battery Brooder

16%

6

589

19

570

96.77

22%

6

65

1

64

98.46

Number Chicks
Lost First Seven
Days

Number Chicks
Survived
Seven Days

Percent Survival
First Seven Days

�Table 4. A comparison of eggs produced by Hungarian partridge pairs receiving two extra hours of light
with those receiving no extra light, 1971. 1/

Treatment

Number
Pens

Number
Eggs
Laid

Number
Infertile
Eggs

Number
Fertile
Eggs

Two Extra
Hours A.M.

3

70

13

Two Extra
Hours P.M.

3

51

Controls

3

77

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

57

28

40.00

49.12

-

29

25

25

15

29.41

60.00

2

8

25

52

41

53.24

78.84

-

11

Number
Culls

2
(1 unknown)
2

I

at-'
I

1/
- A 16 percent protein game bird breeder feed was used in all pens.

�-62-

More total eggs were laid by the breeder pairs in the 3 pens receiving no
extra lighting or controls (77) compared with those receiving 2 extra hours
of artificial lighting in the morning (70) and those receiving 2 extra hours
in the evening (51). In comparing numbers of fertile eggs laid, the 3 pairs
receiving 2 extra hours of lights in the morning were highest (57), followed
by the controls (52) and the 2 extra hours of lights in the evening (25).
Percentage hatch of fertile eggs was highest for the controls (78.84%) followed
by extra lighted pairs in the evening (60.00%), and the extra lighted pairs in
the morning (49.12%). Numbers of chicks hatched was highest for the controls
(41) followed by extra lighted pairs in the morning (28) and extra lighted
pairs in the evening (15).
The extra lighting brought the breeding pairs into production earlier than
the breeding pairs without artificial lighting. The extra lighting was started
on March 12,1971 and the first egg was laid in pen number 48 (lighted for 2
extra hours in the early morning) on April 14, 1971. Thus, 33 days, or approximately 5 weeks were required to stfmulate this hen into laying eggs. The
earliest date which breeders without artificial lighting laid in 1971 was April
29, or 15 days after pen number 48 came into production. On May 8, 1971, all
6 of the extra lighted pens were in production and had laid a total of 32 eggs
in comparison to no eggs laid by the controls with no extra lighting. Two of
the extra lighted pens were in production in mid-April 1971. Early egg production in 1971 proved to be disadvantageous because enough total eggs were
not secured for an incubation setting until May 25, 1971. This caused many
of the very early eggs to be held in storage too long for good hatchability.
Survival in Holding Pens
Table 5 lists results of the overwinter survival tests completed in April,
1971. Densities of birds varied from 41.16 square feet of ground space per
bird to 501.24 square feet of ground space per bird. Overwinter survival was
excellent under all densities tried for holding pens located both at the
Fort Collins Wildlife Research Station (Fig. 2) and Rocky Ford Game Farm
(Fig. 3).
Similar results were obtained in overwinter survival tests completed in
March, 1972 (Table 6). All birds were held in holding pens at the Fort Collins
Wildlife Research Station. In one group of 73 Huns (4th hatch, 1971) wintered in a holding pen with 953 square feet of ground space (13.05 square feet
per bird), overwinter survival was excellent (100 percent). Hysteria was however observed within this group of birds from October 5, 1971 through November,
1971, after which they appeared quite normal. Thus, the hysteria occurred
from the time these birds were approximately 7 weeks of age until they were
approximately 15 weeks of age. Hysteria was negligible in other densities of
birds tested during the period.
Poorest overwinter survival in tests completed in March, 1972 occurred in a
group of 14 highest producing, 1970 hatched adults held over from 1971 tests.
Three females from this group died in the holding pen, resulting in 78.57
percent survival. Percentage survival in all other groups tested was over
90 percent.

�Table 5. Survival of Hungarian partridge during fall and winter periods according to various densities
of birds and sizes of pens, 1970-71.

Pen and
Hatch

Size
(Ft.)

Total
Sq. Ft.

Date Birds
Counted into
Pen

Number
Birds
In

Sq. Ft. /
Bird

Date Birds
Recovered

Number Birds
Recovered

Percent
Survival

FCWRS
A-I, 1969
(Win.)

l6x48

768

9-27-70

5

153.60

2-23-71

5

100.00

A-2, A-3
1969 (1)

16x96

1,536

9-27-70

14

109.71

2-23-71

14

100.00

Pens 1 &amp; 2
1970 (3 &amp; 4)

35x240+
1lxl1

8,521

12-17-70

207

41.16

4-22-71

199

96.13
I

0"1
Vol
I

Pens 3 &amp; 4
1970 (2)

35x240+
llx11

8,521

9-27-70

17

501. 24

3-1-71

17

100.00

Pens 5 &amp; 6
1970 (1)

35x240+
llx11

8,521

12-17-70

195

43.70

3-1-71

192

98.46

Pen 6
1969 (1)

50x75

3,750

9-28-70

81

46.29

4-22-71

78

96.29

Pens

4(50x75)

15,000

9-28-70

286

52.45

4-22-71

281

98.25

Rocky Ford

3, 5,
7 &amp; 8
1970 (2)

�...••.
~~'

4'

.""'"

~

.

x

~

x
3

10

8

6

4

2

I

--

x
5

N

-o

x
7

12
~

9

II

T~'
..,

•
I'-

I

&lt;l'

~
I

SCALE

~---------------300'
Fig.

3

Holding Pens, Rocky Ford GameFarm

~50~1

o

.0·

l

100'

�Table 6. Survival of Hungarian partridge during fall and winter periods according to various densities
of birds and sizes of pens, 1971-72. !/

Pen and
Hatch

Size
(Ft.)

Date Birds
Total
Counted into
Age
Sq. Ft.
Pen
(approx.)

Number
Birds
In

Sq. Ft./
Bird

Date Birds
Recovered

Numb er Birds
Recovered

Percent
Survival

A-I
1969 (1)

l6x48

768

9-22-71

Adults
(1969 hatch)

12

64.00

3-6-72

11 ~/

91. 67

A-2, A-3
1970 (1, 2)

16x96

1,536

9-22-71

Adults
(1970 hatch)

14

109.71

3-6-72

11 ~/

78.57

Pens 1 &amp; 2
1971 (2)

35x240+ 8,521
1lxll

9-22-71

10 wks.

198

43.03

3-24-72

192

96.97
I

0'1
VI
I

Pens 3 &amp; 4
1971 (3)

35x240+ 8,521
1lxl1

9-22-71

8 wks.

201

42.39

3-24-72

193

96.02

Pen 5
1971 (4)

20x35+
11x11
12x12

953

9-22-71

5 wks.

73

13.05

3-24-72

73

100.00

Pen 6
1971 (1)

35x200

7,000

.9-22-71

13 wks.

136

51.47

3-17-72

132

97.06

North Pen

53x105

5,565

9-22-71

Adults
(1970 hatch)

86

64.70

3-2-72

82

95.35

Totals

719

Average
1/
- All Huns were held in holding pens at the Fort Collins Wildlife Research Station.
1/on1y females lost in these two groups of birds.

694
96.52

�-66-

Egg Production and Hatchability by Source and Age of Breeders
Table 7 lists a comparison of eggs produced and hatchability of eggs by Huns
of various ages and sources. Highest numbers of total eggs (213) were produced by the 6 pairs of 1969 hatched highest producing adults held over from
1970 tests, followed by 1970 hatched run-of-the-mill pairs (199), and 1970
hatched females from pure Winchester Farms stock crossed with 1970 hatched
males from pens 3 or 9 (109). In considering numbers of fertile eggs laid,
the 1970 hatched run-of-the-mill pairs were highest (175) followed by 1969
hatched adults (160), and 1970 hatched females from pure Winchester Farms
stock crossed with 1970 hatched males from pens 3 or 9 (96). Numbers of chicks
hatched ranked in the same order as numbers of fertile eggs produced.
Percentage hatch of all eggs was highest (78.48%) for the 6 pairs of females
from pens 3 or 9 crossed with males from pure Winchester Farms stock, followed
by 6 pairs of run-of-the-mill stock (71.85%) and 6 pairs of highest producing
adults held over from 1970 tests (62.91%). Percentage hatch of fertile eggs
was highest (89.86%) for the 6 pairs of females from pens 3 or 9 crossed with
males from pure Winchester Farms stock. The next highest percentage hatch of
fertile eggs (83.75%) occurred with the 6 pairs of highest producing adults
held over from 1970 tests, followed by 6 pairs of run-of-the-mill pairs
(81.71%) and 6 pairs of females from pen 31 crossed with males from pens 3 or
9 (77.55%).
These results showed the value of holding exceptionally high producing pairs
for an additional year's production, considering numbers of fertile eggs
laid and hatchability. Additional tests will be made in 1972 to compare the
third year's production with the second and first years production. Results
of the first year's selection tests were somewhat disappointing but additional
tests will be required before the value of selection can be fully evaluated.

Prepared by

tb:1.(rrL~~
Wildlife Researcher

�Table 7.

A comparison of eggs produced by Huns of various ages and sources, 1971.1/
Number
Eggs
Laid

Number
Infertile
Eggs

Number
Fertile
Eggs

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

Number
Pipped

Number
Dead
Germs

Number
Culls

Source

Hatch

No. of
Pens

Highest
Producing
Adults

1969 (1)

6

213

53

160

134

62.91

83.75

1

25

3

F-Win.
M-3 or 9

1970 (1)

6

109

13

96

63

57.79

65.62

1

32

1

F-31
M-3 or 9

1970 (1)

6

73

24

49

38

52.05

77 .55

-

11

4

I

0\
......s
I

Run-ofthe-mil1

1970 (1)

6

199

23

175

143

71.85

81.71

5

27

F- 3 or 9
M- Win.

1970 (1)

6

79

10

69

62

78.48

89.86

2

5

1/
- A 16 percent protein game bird breeder feed was used in all pens.

9
(1 unknown)

10

�I

0\
00
I

Fig. 4. A group of lO-day old Hungarian partridge chicks explore a rearing pen at the Fort Collins
Wildlife Research Station. (D. Hoffman photo).

�-69April,

JOB PROGRESS

S tate of

1972

REPORT

--=.CO:::.;L=..O:::.;RA=D:..:O:....._
Game Bird Survey

W-37-R-25

Project No.

10
Job No.
12
Relationships
of the Productivity and Distribution of
Wild Turkeys on the Uncompahgre Plateau to the Environment

Work Plan No.
Job Title

Period Covered:

April

1, 1971 to March

31, 1972

P.S. OBJECTIVES
1.

To determine the size, distribution and productivity
population on the Uncompahgre Plateau.

of the wild turkey

2.

Record movements of wild turkeys and determine
affect these movements.

3.

Determine the sex and age structure of the turkey kill on the Uncompahgre
Plateau to provide estimates of total harvest and size of the turkey
population.

4.

Determine the vegetative types present on the wild turkey summer and
winter range and the species composition and abundance of food producing
species within each type. Also determine the elevation, describe topography and record certain climatic factors on part of the better turkey
summer and winter range.

how environmental

factors

SEGMENT OBJECTIVE
To compile,

analyze,

summarize

and publish

information.

Progress
Field work was completed on October 20, 1967. All data have been summarized,
and most data have been analyzed.
A manuscript containing these data and results
from Work Plan 12, Job 14 is being edited and revised at this time.
Future Plans
The manuscript should be completed
been completed.

Prepared

in Segment

26.

!?:-L~~:::.::::.==" :::-- __

bY_ct~/.;:;....:::~~~~--=,
Howard D. Funk
Section Chief, Small Game Research

Publication

plans have not

��-71April,

JOB PROGRESS

S ta te of
Project
Work

REPORT

C.=.O::..:L=..O::..:RAD=::..;O=--_
Game Bird Survey

W-37-R-2S

No.

Job Title

Use of Food Plots

Covered:

April

14

Job No.

12

Plan No.

Period

19'72

1, 1971 to March

to Concentrate

Wild

Turkeys

31, 1972

P.S. OBJECTIVE
Determine the effects
wild turkeys.

of small food plots

SEGMENT
To compile,

analyze,

summarize

on fall concentrations

and harvest

of

OBJECTIVE

and publish

information.

Progress
Field work was completed December 2, 1968. Data were analyzed, written up and
incorporated
in a manuscript containing information gathered under W-37-R
Work Plan 12, Job 10. Due to changes in work assignments
of personnel,
the
final product has been delayed but the manuscript
is being edited and revised at
present.

Future
The manuscript
completed.

will

be completed

Plans

in Segment

26.

Publication

- .._---_ .._------

Prepared

irulI~[)~lA~~~._

by:.----.:L::::.....:::.!.·/~/
Howard D. Funk
Section Chief, Small Game Research

plans have not been

---_ ... __ ._---_._ ... _-----

��-73...,
April, 1972

JOB PROGRESS REPORT

COLORADO

State of
Project No.

W-37-R-25

Work plan No.

15

Job Title

Job No.

2

Study of Mountain Quail Adaptability

Period Covered:
Personnel:

Game Bird Survey

April 1, 1971 to March 31, 1972

Ronald B. Arant, Tom W. Barnes, Dwight E. Owens and Donald M. Hoffman.

ABSTRACT
No additional field releases of mountain quail have been made in Colorado
since June, 1970. A small covey of eight mountain quail was located approximately two miles northeast of Gateway, Colorado by Wildlife Conservation Officer
Dwight Owens (ltr. D. Owens to J. Grieb, 1/18/72). The location of these
mountain quail was reported in early December, 1971 by late season deer hunters.

��-75-

STUDY OF MOUNTAIN QUAIL ADAPTABILITY
Donald M. Hoffman

P. S. OBJECTIVE
To determine the ability of mountain quail to establish reproducing
populations in areas of mixed shrub and pine in selected sites in
Colorado.
SEGMENT OBJECTIVES
1.

Introduce mountain quail.

2.

Determine presence and dispersal of mountain quail.
METHODS AND MATERIALS

An inquiry was made in July, 1971 (pers. comm. D. Hoffman to R. Arant) to
determine whether any new leads on locatio~s of mountain quail on the
Uncompahgre Plateau had been received sIncc Segment 24. A negative reply
was received on July 21, 1971.
With no new leads secured from western Colorado, time scheduled for field
searching for mountain quail was used for inventory work concerned with
Hungarian partridge releases.

RESULTS AND DISCUSSION
A total of 372 wild-trapped and 25 pen-raised mountain quail (total 397)
have been released in Mesa County on the west side of the Uncompahgre
Plateau since 1965. The last field release was made in June, 1970 and
consisted of 25 surplus pen raised mountain quail from the Fort Collins
Wildlife Research Station.
In early December, 1971, late season deer hunters reported seeing chukar
partridges with topknots in an area approximately 2 miles northeast of
Gateway, Colorado to Wildlife Conservation Officer Dwight Owens. A small
covey of 8 mountain quail were later found by Dwight Owens (ltr. D. Owens
to J. Grieb, 1/18/72). This is the first sighting of mountain quail since
December, 1969 when approximately 20 were observed on the South Fork of
Mesa Creek by Conservation Aide Tom Barnes.

�-76-

Within their native ranges, mountain quail that summer at higher elevations frequently migrate to lower levels in winter. Whether this has
been the case with the last 2 observations is not known at this time.

Prepared by

~~&gt;rz~~

Don~offman:
Wildlife Researcher

�-77April,

JOB PROGRESS

State of

COLORADO

Project No.

W-37-R-25

Work Plan No.
Job Title

Personnel:

REPORT

Game Bird Survey

15

3

Job No.
Experimental

Period Covered:

1972

Breeding

of Mountain

Quail

April 1, 1971 to March 31, 1972

Lawrence A. Webster, John F. Corey, Robert L. Schmidt,
Harrington, Larry R. Crooks and Donald M. Hoffman.

George G.

ABSTRACT
Good success was experienced with the experimental propagation of mountain
quail in 1971 with 344 eggs laid (culls excluded), of which 209 were fertile.
A total of 173 good chicks were hatched from these eggs. A total of 28 breeders
(18 hens and 10 cocks) were used in four pairs (1 hen and 1 cock), four trios
(2 hens and 1 cock), and two quads (3 hens and 1 cock) in ten separate pens on
test in 1971. The sorting of breeders in early February, 1971 and disturbing
the breeders as little as possible appeared to be of considerable value in
securing egg production.
In a test to compare egg production by various sex ratios of breeders, numbers
of fertile eggs laid per hen was highest with the trios (12.50) followed by
the quads (11.00) and pairs (10.75). The best hatching of fertile eggs occurred
with the trios (92.00%) compared with 83.72 percent for the pairs and 72.72
percent for the quads.
Numbers of chicks hatched was also highest for the trios.
The clipping of the flight feathers on both wings of breeders in half of the pens
calmed these easily stressed birds considerably but also resulted in 20.34 percent
fewer fertile eggs being produced (94 for the five pens with clipped flight
feathers compared to 118 for the five pens with normal wings).
Excellent survival was experienced with the first three hatches for the first
seven days (average 94.80%) and good survival was experienced for the first 30
days (average 83.24%) during 1971. No birds were hatched in a fourth setting
of 13 eggs. Over-winter survival in pens with various bird densities and ages
ranged from 86.67 percent in a holding pen with 26.40 square feet of ground space
per bird containing 60 1971-hatched quail to 100 .•00 percent in a holding pen
with 144.00 square feet of ground space per bird containing 11 1971-hatched quail.
The average over-winter survival was 91.62 percent for the five groups of mountain
quail on test.

�-78-

RECOMMENDATIONS

An unanswered_question in mountain quail propagation is whether it is
possible to secure egg production from young of the previous year. A
test to determine this is recommended for 1972.
A test to determine the value of providing 2 extra hours of lighting
in the early morning hours on total and fertile egg production is planned
for 1972. In addition, tests to compare survival of mountain quail in
holding pens containing various bird densities, started in 1971, are
recommended for continuation in 1972.

�-79-

EXPERIMENTAL BREEDING OF MOUNTAIN QUAIL
Donald M. Hoffman

Total egg production (culls excluded) from the 28 mountain quail breeders
(18 hens and 10 cocks) was 344 compared with only 1 infertile egg laid by
32 breeders of mixed sexes in 1970. A total of 173 good chicks were
hatched from the eggs in 1971. Eighty seven surplus mountain quail were
released on the Spanish Peaks Management Area in Las Animas County on
April 15, 1972 and 66 were held for breeders for the 1972 test series.
P. S. OBJECTIVE
To develop game farm production techniques for mountain quail.
SEGMENT OBJECTIVE
To measure the relationship of pair mating versus group matings of 2 hens
and 1 cock and 3 hens and 1 cock to fertile egg production.
METHODS AND MATERIALS

Fertile Egg Production by Various Sex Ratios
A new set of 10 experimental breeding pens was completed in April, 1970
(Fig. 1). Table 1 lists the pen schedule for mountain quail for 1971 tests.
Pairs (1 hen and 1 cock) were placed in pens 1, 5, 7, and 9; trios (2 hens
and 1 cock) were placed in pens 2, 4, 6, and 10; and quads (3 hens and 1
cock) were placed in pens 3 and 8 on February 9, 1971.
All breeders were fed a standard 22 percent protein level game bird breeder
feed from March 6, 1971 until removed from the breeder pens on July 23, 1971.
Primaries and secondaries of both wings were clipped on all breeders in pens
4, 5, 8, 9, and 10. All birds were 1969-hatched or older and all but 4 were
pen-raised at the Fort Collins Wildlife Research Station. The 4 were wild
trapped mountain quail received from Oregon in January, 1970.
All pens were fed and watered only twice a week and eggs were collected at
these times to reduce the disturbance factor. Eggs laid were marked with
the date collected and pen number to facilitate determination of total eggs
laid, fertile eggs laid, and numbers of chicks hatched by pens. Adequate
cover in the form of growing clumps of tall wheatgrass were planted in
April, 1970. Perches were installed in the spring of 1971 on the advice
of a successful game bird breeder in California.

�-80-

N·

30'---"-

JE----

~

1E----12'--~

12

-

0

CD

'\

ex&gt;
~

.

-

\

m
SCALE 8

Fig. 1.
Ootall.d

ArranGement

of Moulttaln

.I

t

I.

·0'

5'

10'

0"011 Broadlng

Pent.

�-81-

Table l.

Pen schedule for1ll.ountainquail, 1971.

Pen
Number

Pairs
(1 Hen &amp; 1 Cock)

1

T

Trios
2 Hens &amp; 1 Cock)

T

2

T

3
T*

4

T*

5

T

6

T

7
8

T*

9

T*
T*

10
Number of Pens
on Test
Key:

QuadruElets
3 Hens &amp; 1 Cock

4

4

2

T = Pen on test.
* = Primaries will be clipped on both wings.

Survival Tests
Although not specifically written as planned tests for 1971, records of
survival of chicks for the first 7 and 30 day periods and for overwinter
survival in holding pens were maintained.
Numbers of day old mountain quail placed in the brooder houses after removal from the hatcher were recorded and losses experienced were marked
on brooder house cards with causes of death when this could be determined.
Young removed were also recorded and percentage survival calculated. Figure
2 shows a group of 12-day old mountain quail at the Fort Collins Wildlife
Research Station.
Birds counted into holding pens (Figs. 3 and 4) in either July, 1971 (adult
breeders) or in September, 1971 (1971-hatched young) were recorded. Birds
recovered in March and April, 1972 were also recorded and survival rates
calculated.

�I

00

tv
I

Fig. 2. A group of 12-day old mountain quail chicks leave a brooder house at the Fort Collins
Wildlife Research Station. (D. Hoffman photo).

�-83.,
',1

"

,,.

"

-&gt;

..
!

'.'

-.-

10'

~4.!lI

9

8

N

"~i

...

::

7

-•....
N

SCALE
'7---"---'

Jt-------

O'

30'

5'

F 16. 3.
MOUNTAlN

QUAIL

H'OLDING

PENS,

WEST

SECTION

10'

�(

-84-

I

I(
1E-8'~

f

-.

20'
zj
E-- 8'---7f~

I

·co

N

!
I

"

1

5

I

3

I

1

\
6

4

2

.

-co
It)

Stole:

o
GO' ----------------~
F1g.4.
Mountain

Ouoll

Ho l d in q Pens,

EAST

SEOTION

,":

15 ft.
15

30

�-85-

RESULTS AND DISCUSSION

Fertile Egg Production by Various Sex Ratios
Due to the extreme difficulty in sexing mountain quail by external
characteristics, it is possible that errors in the sex ratios of pens
6 and 8 were made. In each case, one or more birds originally called hens
may actually have been cocks. Birds were not sacrificed for examination
of internal organs.
Table 2 lists a comparison of eggs produced and hatched by mountain quail
in pens with 3 different sex ratios of breeders. Because numbers of hens
varied, numbers of total eggs and numbers of fertile eggs per hen were
calculated. Highest numbers of fertile eggs per hen were produced by
the trios (12.50) followed by the quads (11.00) and pairs (10.75). The
best hatch of fertile eggs occurred with the trios (92.00%), compared
with 83.72 percent for the pairs and 72.72 percent for the quads. Numbers of chicks hatched was highest for the trios (92 chicks). Based upon
these comparisons, it appears that trios (2 hens and 1 cock) produced
better than either pairs (1 hen and 1 cock) or quads (3 hens and 1 cock).
In comparing numbers of fertile eggs produced by 5 pens of breeders with
clipped flight feathers on both wings with an equal number of pens and
breeders with normal wings, the pens of breeders with normal wings produced 118 fertile eggs compared with 94 (20.34% less) for the pens of
breeders with clipped wings. Thus the advantage of considerably calming
the breeders down must be weighed with the disadvantage of fewer fertile
eggs being produced.
Table 3 lists information relating to the 4 separate settings made in the
automatic forced air incubators. Of 344 eggs set, 135 were infertile, 25
were dead germs, 8 pipped but did not hatch, 3 produced cripples, and 173
produced good chicks. About half (51.16%) of the total eggs set hatched
and 84.21 percent of the fertile eggs hatched. Hatchability of the last
setting of 13 eggs was zero with 11 being infertile and 2 dead germs.
Survival Tests
Table 4 lists survival percentages of mountain quail by separate hatches
in 1971. Excellent survival was experienced for the first 7 days (average
94.80%), and good survival was experienced through the first 30 days
(average 83.24%). A number of young mountain quail from the first hatch
developed sore eyes, apparently from faulty brooding procedures. Eight of
the total of 44 chicks taken to the brood from this hatch were lost due to
total blindness and 2 additional were lost from other causes. Brooding
procedures were corrected for the second and third hatches and no further
losses from this cause were experienced.

�Table 2.

A comparison of eggs produced and hatched by mountain quail in pens with various sex ratios, 1971.

Sex
Ratio
1H:1C
(pairs)

Pen
Number

Number
Good
Eggs
Laid

1
5*
7
9*

40
35
6
17

Sub-total

2H:1C
(trios)

98
-2
4*
6
10*

3H:1C
(quads)

3
8*

24.50

17.75

89
15

Sub-total

104

TOTAL

344

AVERAGE

Hen

36
28
0
78
142

Sub-total

No.1

17.33

19.11

*Primaries on both wings clipped.

Number
Infertile
Eggs

Number
Fertile
Eggs

10
35
6
4
55

43

3
22
0
17

33
6
0
61

42

100

37
1

52
14

38

66

135

209

Number
Hatched

Percent
Hatch
All Eggs

Percent
Hatch
Fertile Eggs

30

24

60.00

80.00

a
a

a

-

-

0
12

70.58

36

No.1
Hen

13
10.75

12.50

11.00

Number
Pipped

Number
Dead
Germs

Number
Culls

92.30

-

3
0
0
0

3
0
0
1

2
1
2
1

36.73

83.72

3

4

6

30
6
0
56

83.33
21.42

90.90
100.00

a

91.80

2
0
0
2

a

71. 79

1
0
0
3

00
0'

4

I

92

64.78

92.00

4

4

9

37
11

41.57
73.33

71.15
78.57

1

a

14
3

8
1

48

46.15

72.72

1

17

9

25

24

-

-

-

8

176
11.61

51.16

84.21

5

I

�Table 3.

Mountain quail hatches, 1971.

Hatch
Number

Date and
Time Set

Number
Set

1

5-10-71
6:00 PM

93

6-5-71
11:00 AM

44

1

6

38

4

48.38

83.33

2

6-2-71
11:00 PM

111

6-28-71
11:00 AM

58

1

5

46

1

53.15

90.76

3

6-23-71
11: 49 PM

127

7-19-71
10:00 AM

71

1

12

40

3

56.69

82.75

Date and
Time to
Brooder

Hatched
Cripples
Good Chicks

DG's

Not Hatched
Inf. Pipped

Percent Hatched
Fertile Eggs
Total Eggs

I

00

"
I

4

TOTAL
AVERAGE

7-17-71
2:00 AM

13

344

8-12-71
Hatch
Terminated

0

0

2

11

0

173

3

25

135

8

0.00

0.00

51.16

84.21

�Table 4.

Survival of young mountain quail, 1971.

Hatch
Number

Date

Number to
Brooder

1

6-5-71

2

% Survival

Number Surviving 7-Days
Date
Number

% Survival

7-Days

Number Surviving 30-Days
Number
Date

44 1./

6-12-71

41

93.18

7-5-71

34

77 .27

6-28-71

58

7-5-71

55

94.82

7-28-71

49

84.48

3

7-19-71

71

7-26-71

68

95.77

8-19-71

61

85.91

4

8-12-71

0

30-Days

I

00
00

TOTAL

173

94.80

AVERAGE

1/ A number of young from this hatch developed
were lost due to blindness.

I

144

164

sore eyes from faulty brooding methods.
This was corrected in the brooding of later hatches.

83.24

Eight out of 10 listed

�-89-

Table 5 lists survival of mountain quail during the fall and winter
periods while held in holding pens with various densities and ages of
birds. Survival ranged from 86.67 percent in a pen with 26.40 square
feet of ground space per bird containing 60 1971-hatched mountain quail
to 100.00 percent in a pen with 144.00 square feet of ground space per
bird containing 11 1971-hatched mountain quail. The average overwinter
survival was 91.62 percent for 5 groups of mountain quail. Densities
of birds tested,of from a low of 144.00 to a high of 20.57 square feet
of ground space per bird in holding pens overwinter were evidently not
high enough to affect overwinter survival rates of mountain quail,
based upon data gathered. Total numbers of birds held in holding pens
may affect survival rates of mountain quail to a greater extent than the
amount of ground space per bird which is provided.

Prepared by
Wildlife Researcher

�Table 5. Survival of mountain
sizes of pens, 1971-72.

quail during fall and winter periods by various densities of birds and

Date Birds
Released in
Pen

No. of
Birds

Sq. Ft.1
Bird

Date Birds
Recovered

No. of Birds
Recovered

Percent
Survival

Pen and
Hatch 1/

Size
(Ft. )

Total
Sq. Ft.

1 &amp; 2

1971-2

2Ox76+
8x8

1584

9-22-71

11

144.00

3-7-72

11

100.00

3 &amp; 4

1971-3

2Ox76+
8x8

1584

9-22-71

60

26.40

4-14-72

52

86.67

5 &amp; 6

--

2Ox76+
8x8

1584

--

0

--

--

0
I

\0

0

7 (adult
breeders)

lOx 72

720

7-23-71

29

24.83

3-7-72

27

93.10

8

1971-2

lOx 72

720

9-22-71

35

20.57

3-7-72

34

97.14

9

1971-1

lOx 72

720

9-22-71

32

22.50

3-7-72

29

90.63

TOTALS
AVERAGE

11

- See Figures 2 and 3.

167

153
91.62

I

�-91-

April,

1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-37-R-25

Work Plan No.

17

Job Title

Continued

Period Covered:
Personnel:

G?me Bird Survey

2

Job No.
Inventory

of Selected

Ptarmigan

Populations

April 1, 1971 to October 15, 1971

C1ait E. Braun and Terry A. May.

ABSTRACT

Investigations of white-tailed ptarmigan (Lagopus 1eucurus) populations in
Colorado initiated in 1965 were continued in 1971 with major emphasis in
ascertaining population levels and relationships of nesting success and production to climatic conditions.
Breeding densities decreased slightly (1.4
birds per square mile) at Rocky Mountain National Park, and significantly
(4.7 birds per square mile) at Crown Point, and increased significantly at
both Mt. Evans and Independence Pass (5.8 and 10.7 birds per square mile, respectively).
The decrease at Crown Point was related to heavy hunter harvest
in 1970 and poor recruitment of immature birds.
The significant increases at
Mt. Evans and Independence Pass were the result of no or little harvest in
1970 and good recruitment of immature birds into the 1971 breeding populations.
Weather during periods of egg deposition and incubation in 1971 was warm and
dry, and nesting success ranged from about 50 to 65 percent.
Survival of chicks
to September 1 was similar to most previous years studied, and was 4.3 chicks
per successful female. Present data continue to indicate that nesting success
and production are closely related to amount of precipitation and number of
frost-free nights in June.

��-93-

CONTINUED

INVENTORY

OF SELECTED

PTARMIGAN

POPULATIONS

Clait E. Braun

Knowledge concerning population levels of white-tailed
ptarmigan and factors
influencing observed densities is essential for proper management.
This
project was undertaken to examine long term population trends and effects of
climatic conditions on white-tailed
ptarmigan in Colorado.
The initial 5
years of research on this grouse have been previously reported by Braun and
Rogers (1971).
This report presents data collected during the second year
of the last 5-year segment.
Data collected during the initial year of the
second 5-year segment were presented by Braun (1971).

P. S. OBJECTIVE
To test the hypotheses that (1) populations of white-tailed
ptarmigan in
Colorado are not cyclic, and (2) production and fall population densities of
this species in Colorado are related to temperature and moisture conditions
during June of the same year.

SEGMENT

OBJECTIVES

1.

To estimate breeding population levels of ptarmigan in four study areas
(Mt. Evans, Crown Point, Independence Pass, and Rocky Mountain National
Park) •

2.

To estimate

3.

To obtain weather data from a representative
alpine area for use in
determining correlations between spring weather conditions and fall
ptarmigan populations.

4.

To compile

nesting

success

data and prepare

METHODS

and production

progress

in the above areas.

report.

AND MATERIALS

Techniques used were essentially those developed under Work Plan 17, Job 1,
and reported in detail by Braun and Rogers (1971), and updated by Braun (1971).
In 1971, white bandettes with black numerals were used to mark all newly
banded birds and those whose older bandettes had become too worn for individual recognition.
Weather data were obtained from the Institute of Arctic
and Alpine Research (INSTAAR), University of Colorado, for reasons previously
explained (Braun 1971).

�-94-

Table 2.

White-tailed ptarmigan breeding population densities, 1966-71.
Birds Per Sg,uare Mile
1968
1969
1970

Study Area

1966

1967

1971

Rocky Mountain National Park

29.0

25.2

29.4

30.8

24.8

23.4

Crown Point

14.5

21.2

18.1

14.5

10.4

5.7

Mt. Evans

7.8

7.1

7.1

5.8

5.2

11.0

Independence Pass

18.7

18.7

18.7

17.9

10.7

21.4

Breeding population levels in Rocky Mountain National Park have remained
markedly stable over the 6-year period studied, varying about seven birds per
square mile. At Crown Point, the trend in breeding population is steadily
downward, apparently partially because of increased hunter activity at this
site coupled with poor production. At Mt. Evans, breeding populations were
at low levels when the study was initiated, with no increase being recorded
until after the experimental closure of the area to hunting in 1970. The
situation at Independence Pass remained stable until 1970, when after 2
successive years of heavy hunter harvests, the breeding population significantly decreased. Hunting at this site was very poor in the fall of 1970
(Braun 1971), and the population rebounded after only 1 year. Although
nesting success and production were average or better on all areas in 1970
(Braun 1971), recruitment of subadult birds into the breeding population was
good only at Independence Pass and Mt. Evans in 1971. Reasons for the low
recruitment of subadult birds at Rocky Mountain National Park and Crown
Point are not known.
Nesting Success and Production
As in most previous years, nesting activities were initiated on all areas
between June 10 and June 20, with events at Mt. Evans slightly earlier than
those at the other areas. Weather conditions during June were similar to
those experienced in 1970, although existing snowfields were much more extensive in 1971. No major snowstorms were recorded after June 1. Estimated
peak of hatching in 1971 was between July 13 and July 17. While nesting
success was anticipated to be good in 1971, only 20 different successful
females were observed on the study areas, while 12 unsuccessful females were
observed. This compares with 27 successful females and 5 unsuccessful females
observed in 1970. Estimated nesting success was 65 percent in Rocky Mountain
National Park, &gt; 50 percent at Mt. Evans, and about 50 percent at Independence
Pass. Due to the removal experiment at Crown Point (WP 17, J3) no hens were
observed after the breeding period.
Initial brood size in 1971 upon hatching was about seven chicks, although no
nests were located. By early August this number had decreased to about 4.5,

�-95-

DESCRIPTION OF STUDY AREAS
Areas intensively investigated have been described in detail by Braun and
Rogers (1971) and have been presented by Braun (1971).

RESULTS AND DISCUSSION
Breeding Densities
Censuses of breeding birds were initiated on May 6 in Rocky Mountain National
Park, with few males being observed on known territories. Due to heavy snow
accumulations in subalpine areas, breeding surveys were not initiated until
May 14 at Mt. Evans, June 2 at Crown Point, and June 13 at Independence Pass.
Most pairing was completed at Rocky Mountain National Park and Mt. Evans by
May 20, and by at least June 1 at Crown Point and Independence Pass. Timing
of breeding events in 1971 was later than in 1970 and resembled that
observed in 1967 and 1968. Densities observed are presented in Table 1.
Table 1.

White-tailed ptarmigan breeding population denSities, all areas, 1971.

Square
Miles

No. of
Breeding
Pairs

Unmated
Birds

Total
Breeding Season
Population

Birds per
Square Mile

Tombstone RidgeSundance Mtn.

1.25

8

3

19

15.2

Toll Memorial

.19

4

3

11

57.9

Fall River Pass

.70

8

4

20

28.6

Total RMNP

2.14

20

10

50

23.4

Crown Point

1.93

4

3

11

5.7

Mt. Evans

1.54

6

5

17

11.0

Independence Pass

1.12

10

4

24

21.4

Study Area
Rocky Mountain
National Park

Breeding densities at Independence Pass and Mt. Evans increased significantly
over 1970 levels while the density at Crown Point continued to decline.
Numbers of breeding birds observed at Rocky Mountain National Park were essentially unchanged from those recorded in 1970 (Table 2).

�-96-

with average brood size to September 1 of about 4.2 chicks per successful
female (Table 3). This is approximately the same number of chicks per successful female observed in all other years except 1969.
Table 3.

Number of broods and average brood size, all areas, 1971. 11

Date

Number of
Broods Observed

Average Number of
Chicks per Brood

July 1-15

0

0

July 16-31

1

7

August 1-15

8

4.5

August 16-31

16

4.2

September 1-30

4

4.0

Total

29
4.3

Average

1./ Only distinct broods are included.
Fall Densities
Estimates of densities of white-tailed ptarmigan on September 1, while useful
in illustrating population gain through production, are difficult to accurately
derive due to a number of variables. Estimated fall densities for each area
studied are presented in Table 4. These densities were calculated following
the three basic assumptions discussed in detail by Braun and Rogers (1971).
Data presented in Table 4 generally depict the situation at all areas.
In 1971, all alpine areas of the State were open to ptarmigan hunting from
September 11 through September 30, except in southern and southwestern Colorado,
where the season was open from September 11 through October 11. Thus, the
Crown Point area was open from September 11 through September 30, while the
Independence Pass area was open from September 11 through October 11. No
ptarmigan hunting was allowed during any of the early big game seasons, and
the area ~ mile on either side of the Mt. Evans highway remained closed for
the second year to all grouse hunting. Bag and possession limits for ptarmigan
alone were 3 and 6.
During and following the hunting season no ptarmigan bands were received from
Mt. Evans or Crown Point, indicating that the closure at Mt. Evans and the
removal experiment (WP 17, J3) at Crown Point were successful.

�-97A minimum of 18 birds, of which 7 were banded, were harvested at Independence
Pass in 1971. The seven banded birds harvested represented about 17 percent
of the 41 banded birds available to hunters.

Relationships of Weather:Nesting

Success and Production

Weather data obtained from the 12,300 level of Niwot Ridge, through the
courtesy of INSTAAR for 1971 are presented in Table 5. Detailed analysis and
statistical correlations will be completed for the final report in 1975-76.
The close correlation between June weather conditions and ptarmigan nesting
success and production previously suggested continued in 1971. In general,
weather conditions were warm and dry in June and July 1971, and nesting
success and production were similar to 1970 when weather conditions were
similar, and better than 1969 when the early summer weather was colder and
wetter.

LITERATURE CITED
Braun, C. E. 1971. Continued inventory of selected ptarmigan populations.
Colo. Div. Game, Fish and Parks, Game Res. Rept., Fed. Aid Proj. W-37-R.
April. pp 91-106.
______
~' and G. E. Rogers. 1971.
Div. Game, Fish and Parks.

a~U~·

The white-tailed ptarmigan in Colorado.
Tech. Pub1. No. 27:80 pp.

Prepared by _ •••
r~,_~...;;....,.
C1ait E. Braun
Wildlife Researcher

_

Colo.

�Table 4.

Estimated fall densities of white-tailed ptarmigan, 1971.
Estimated
Percent
Nesting
Success

Average
Brood Size
on
September 1

Total
Production

Total
Breeding
Population

Total
Population
on
September 1

Birds Per
Square Mile

Percent
Gain

Ro cky Moun tain
National Park

65

4.4

57

50

102

52.3

55.9

Crown Point

o 1:../

0

0

11

0

0

0

Mt. Evans

50

3.0

9

17

24

15.6

37.5

Independence Pass

50

4.5

22

24

44

38.4

50.0 .

Area

I

1:../

\0
00

Due to experiment conducted under WP17, J3.

Table 5.

I

Weather data, Niwot Ridge, 12,300 ft., May-July, 1971.

Number Days
Minimum
Temperature
Less than 32° F

Ppt/Month
(Inches)

Number Days
Relative
Humidity
90 or Above

Average
Winds peed/
Month
(mph)

Month

Max.

Min.

Mean Dail~
Max.
Min.

May

50°F

7°F

34.70F

20.7°F

31

3.04

19

12.9

June

61°F

22°F

49.loF

35.loF

12

0.39

12

12.8

July

63°F

33°F

53.6°F

39.20F

0

2.43

15

9.5

�-99-

April,

JOB PROGRESS

S ta te of
Project

1972

REPORT

_=C;.::o:.=L;.::o~RAD=_=O~
_
No.

W-37-R-25

Work Plan No.

Game Bird Survey
Job No.

Job Title Experimental

Removal

Period

1, 1971 to September

Covered:

Personnel:

April

of a Breeding

Population

3
of White-tailed

Ptarmigan

30, 1971

Clait E. Braun and Terry A. May.

ABSTRACT
All known white-tailed ptarmigan (Lagopus leucurus) occupying the Crown Point
study area were removed in early June, 1971. This involved nine birds, six
males, and three females.
Locations of all birds were recorded and all males (5)
that had been banded previously on the study area (1967-69) were on the same
territories where initially banded.
Data on vegetative composition of the study
area collected in 1967-68 are presented for descriptive purposes.

��-101-

EXPERIMENTAL REMOVAL OF A
BREEDING POPULATION OF WHITE-TAILED PTARMIGAN
Clait E. Braun
Management of wild animals is dependent upon knowledge about population
levels, reproductive capacity and success, survival and mortality rates, and
habitat requirments. Without adequate habitat for any given species there
would be no need to investigate other aspects of the biology of a given
animal. In addition to knowledge about the particular habitat requirments of
a species knowledge must also be available on the pioneering abilities of a
species and age of pioneers.
During preliminary descriptive studies of white-tailed ptarmigan from 1965-69
(Braun and Rogers 1971) all study areas were vegetatively mapped. This work
was reported by Braun (197lb). Analyses of ptarmigan movements in relation
to vegetative communities revealed that ptarmigan seek and utilize certain
areas depending upon season and that the same patterns prevailed throughout
the alpine areas of ,Colorado (Braun 1969). Since number of breeding territories controls breeding densities, potential productivity, and fall densities,
an experiment was designed to investigate the exact requirments for breeding
territories. Crown Point was chosen as the site for the experiment, as it is
somewhat isolated from adjacent alpine areas, thus reducing the possibility
of ingress between the breeding period and the early fall. This area had been
studied from 1966-70 (Braun and Rogers 1971; Braun 1971a), and locations of
all previously occupied territories were known. Additionally, hunters removed
about 50 percent of the fall population in 1970 (Braun 1971a), and it was
anticipated that the population would be low in the spring of 1971. This
would reduce the number of birds to be removed. This study was initiated in
1971 and is scheduled to terminate in 1974-75.
P. S. OBJECTIVE
To test the hypotheses that (1) ptarmigan select sites for breeding territories which contain readily available bushes of Salix spp. (&gt; 10 em. in
height) and snowfree locations, (2) immature birds initially pioneer new or
empty habitats, and (3) number of occupied territories is affected by age
class of male ptarmigan.

SEGMENT OBJECTIVES
1.

To remove the breeding population of ptarmigan from the established Crown
Point study area.

2.

To map occupied breeding territories at Crown Point prior to removal of
breeding birds.

3.

To obtain blood samples from all birds removed for eventual establishment
of blood parameters for resident and pioneering ptarmigan.

�102-

4.

To compile data and prepare progress report.

METHODS AND MATERIALS
Ptarmigan were located during the breeding season through use of taperecorded male challenge calls as described by Braun et al. (1973). Once
birds were located they were shot with a .410 shotgun. Immediately after
death, blood from each bird was obtained, labeled, and placed on ice for
transport to the laboratory where it was centrifuged and frozen. All blood
samples were transferr~d to .the University of Alberta for examination of
blood serum proteins through starch gel electrophoresis. Location of all
birds removed were plotted on standard U. S. Geological Survey topographic
maps. The area was systematically surveyed five times in 1971, with both
male challenge and chick distress calls following procedures described by
Braun et al. (1973), but additional birds were not found after the initial
removals. All vegetation mapping was accomplished in 1967-68 following
procedures outlined by KUchler (1955).
DESCRIPTION OF STUDY AREA
This area was described in detail by Braun and Rogers (1971) and will be
redescribed in the final report. Extent of the area investigated is shown
in Figure 1.

RESULTS AND DISCUSSION
Removal
During the removal period, a total of nine birds were located and removed.
This total was comprised of six males and three females. Age of the birds
was as follows:
6+ = 1 bird
4+ = 1 bird
4- = 1 bird
3- :z: 3 birds
2+ == 1 bird
2 birds
1Although a total of five searches were made of the area, no other birds were
located. It is possible that some birds were missed, but this number could
not have exceeded two. For practical puposes, the area was considered to
be completely void of ptarmigan from early June until late September. Hunters
reported killing no ptarmigan in the Crown Point area in the fall of 1971.
Location of Breeding Territories
Six of the nine birds] ocated and removed were previously banded. Five of
these birds were males and all were on the same territories they occupied in

�-103-

"~""'.~'

~

,

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00

.. .'

10600

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"',

... ;

.;::,

:

. '" ,~,

Bd.l, /;~

" j

\'

,0

,

,; g,
I!

"

i
I

I/

point,:

, •.\

i:::;

I

I

: /
\

J I
,

",''/:

~

.-

/

,,'

I

I

I

~

"

7'

!

)

-,

CROWN

AREA

COLORADO

H

o

POINT

t

-

Study Area Boundary
SCALE 'N MILES

1/2

' Mile

Ee=+3==~==e==3~~==~~~~==Ee==3==~==Ee=+3==~~
/

Fig. 1.

Crown Point

study

area

(X's mark occupied

\

breeding

territories

1971).

�-104-

1970. One male was on the same territory where initially located in 1967,
while the other four males were on the same territories where initially
banded in 1968 (1) and 1969 (3). Location of all territories occupied are
marked on Figure 1.

Description of Breeding Territories
The vegetative description of the Crown Point area is given in Fig. 2 and
Table 1. All vegetative mapping was a.ccomplished during 1967-68 under Work
Plan 17, Job 1.

Blood Parameter Examination
Blood samples from the nine birds collected were sent to the University of
Alberta for Examination. No results have been received or are necessary
until samples are available from pioneering birds. It is possible that the
results from blood testing will be negative, due to small sample size and
insufficient knowledge about blood proteins in ptarmigan and grouse in general.

LITERATURE CITED
Braun, C. E. 1969. Population dynamics, habitat, and movements of white-tailed
ptarmigan in Colorado. Ph. D. Dissertation. Colorado State Univ., Ft.
Collins. 189 pp,
1971a. Continued inventory of selected ptarmigan populations. Colo.
Div. Game, Fish and Parks, Game Res. Rept., Fed. Aid Proj. W-37-R. April.
pp. 91-106.
_____ • 1971b. Habitat requirments of Colorado white-tailed ptarmigan.
Western Assoc. State Game and Fish Comms. 51:284-292.

Proc.

, and G. E. Rogers. 1971. The white-tailed ptarmigan in Colorado.
----Colo.
Div. Game, Fish and Parks Tech. Publ. No. 27:80 pp.
, R. K. Schmidt, Jr., and G. E. Rogers. 1973. Census of Colorado whitetailed ptarmigan with tape-recorded calls. J. Wildl. Mgmt. 37(1): in press.
K~chler, A. W. 1955. A comprehensive method of mapping vegetation.
Assoc. Amer. Geogr. 45:404-415.

Prepared by
Clait E. Braun
Wildlife Researcher

Ann.

�-104- a

Fig.

2. Vegetation units, Crown Point.'
C-l

Salix-~-Carex

Alpine Meadow

C-2

!2!-Carex Alpine Meadow

C-3

Picea-Salix Krummholz

C-4

Salix-Poa Alpfne Meadow

C-5

Trifo1ium-~-Sa1ix

C-6

Trifolium-Poa Alpine Meadow

C-7

Silene-Trifolium-Carex-Poa

C-8

Dryas-Picea-Sa1ix

C-9

Dryas Turf

C-lO

Pinus-Carex Krummholz

C-ll

Carex Subalpine Headow

C-l2

Dryas-Pinus Subalpine Meadow

C-13

Sibba1di~ Sn0W Accl_l1J1,-,l~tf(ll1 Ar::~::1

C-l4

Salix-Poa-Betu1a

C-l5

Sa1ix-Picea-Trifolium-Poa

C-l6

Artemisia-Trifolium

.

Alpine Meadow

Fellfield

Rock Krummholz

Alpine Meadow
Krummholz

Snow Accumulation Area

�CROWN POINT

I

I-'
o

V1
I

'.

SCALE IN MILES
I

1/2

£3~:=:EE.===

F{ __

0

�-106-

Table

1.

Description of vegetation units, Crown Point.

UNIT
C-1

DESCRIPTION
Sa Hx-Poa-Ca'rex Alpine Meadow:
Salix spp , and Poa spp.
each co;prise 5-25 percent of the total coverag;:-with
Carex spp. accounting for at least 20 percent of the
total cover. Shrubs are less than 1 m in height, with
herbaceous vegetation being less than 1/2 m tall.
Plant density is primarily continuous. Rocks cover at
least 5 25 percent of the areap with small areas approaching 75 percent rock cover. Rocks larger than 12
inches in diameter are dominant. Lichens are about
equally green and black and cover up to one-half the
surface of rocks. Percent slope is variable and averages about 10 percent.
G

C-2

Poa-Carex Alpine Meadow: Between 25 and 50 percent of
the area is covered with Poa spp., with Carex spp.
covering 5-25 percent of the total area. The vegetation is ahort, less than 1/2 m in height, and density
is mostly continuous. Rocks cover less than 25 percent
of the total area and are primarily in the 6- to 12inch diameter size class. Green and black lichens predominate, each covering 5-25 percent of the exposed
rock surface. Slopes are gentle and average about 7
percent.

C-3

Picea-Salix Krummholz: Picea engelmannii covers 25-50
percent of the area, while Salix spp. covers 5-25 percent of the total area. Appearance of this unit is
ragged, as some shrubs attain a height of 4 m. Most of
the vegetation is less than 2 m tall, and the density
is mostly continuous. The amount of rock is variable,
with small areas covering 75 percent of the total area.
Rocks larger than 12 inches are dominant. Lichens are
primarily green and black, with each covering up to 5
percent of the exposed rock surface. Slopes are variable but average 16-20 percent.

C-4

Salix-Poa Alp:i.neMeadow: More than 75 percent of the
area i~overed
with Salix spp., with species of Poa
amounting to about 5 percent of the total coverage.
Plant density in this unit is continuous, and average
height of the vegetation is 2-3 m. Few rocks are
present and slope averages about 7 percent.

C-5

Trifo1ium-Poa-Sa1ix Alpine Meadow: Each of the three
dominant genera comprises 5-25 percent of the coverage
of the area. Most plants in this unit are less than
1/2 m tall; although the shrubs exceed this height,

�-107-

Table
UNIT

1.

Cont.
DESCRIPTION
they do not attain 1 m. Density of the vegetation is
continuous. Rocks cover less than 25 percent of the
area and are primarily larger than 6 inches in diameter.
Green and black lichens dominate, and each covers up to
25 percent of the exposed rock surface. Slope gradient
averages about 10 percent.

C-6

Trifolium-Poa Alpine Meadow: Trifolium spp. and Poa
spp. comprise 5-25 percent each of the total cover;ge
of the area. The dominant clover is 1. dasyphyllum in
contrast to!. parryi in C-5. Vegetation in this unit
is less than 1/2 m in height, and the density is continuous. Rocks cover 5-25 percent of the total area
and are primarily in the 6- to l2-inch diameter class.
Lichens are predominately green and black and cover up
to 50 percent of the surface of exposed rocks. Average
percent slope is about 15.

C-7

Silene-Trifolium-Carex-Poa Fellfield: Rocks dominate
this area and account for about 75 percent of the total
coverage. Rocks are primarily larger than 6 inches in
diameter. Four plant genera comprise less than 5 percent each of the coverage. Plants are short and occur
singly or in small patches. Green and black lichens
each covers up to 25 percent of the exposed rock surface. Slopes are smooth and average about 30 percent.

C-8

Dryas-Picea-Salix Rock Krummholz: Dryas octopetala
comprises 25-50 percent of subunit C-8a, with Picea
engelmannii and Salix spp. each covering up to 5 percent of the total area. Appearance of this unit is
irregular as some Picea bushes attain a height of 4 m.
Density is variable, but is mostly continuous or in
small patches. Rocks cover at least 5-25 percent of
the area with no one size class dominating. Lichens
are few but are mO$tly green and black. Slope gradient varies up to 60 percent but averages about 20
percent. Subunit C-8b differs from C-8a in having
about equal amounts of the three dominant genera.
Picea engelmannii dominates the overs tory of the
Krummholz, while Dryas octopetala dominates the
interspersed windswept areas.

C·~9

Dryas Turf: Mats of Dryas octopetala cover 5-25 percent of the total area, with rocks covering an equal
area. Vegetation of this unit is less than 1/2 m in
height, although scattered Salix bushes attain a
height of 1 m. Density of plants is mostly continuous

J

�-108-

Table
UNIT

1.

Cont.
DESCRIPTION
although many plants occur in small patches. Rock size
is variable, but most rocks are in the 0-6 and larger
than 12 inches diameter size classes. Green and black
lichens each COvers 5-25 percent of the exposed rock
surface. Slopes average 10-12 percent. This unit is
similar to R-l in RMNP.

C-10

Pinus-Carex Krummholz: Pinus flexilis and Carex spp.
each covers 5-25 percent of the area, with rocks
amounting to 25~50 percent of the total coverage. Some
shrubs attain a height of 3-4 m, but much of the vegetation is less than 1/2 m tall. Plant growth is primarily interrupted, although small patches do occur.
One large boulder outcrop is included, and rocks are
primarily larger than 12 inches in diameter. Lichens
are primarily green and black and cover up to 50 percent of the exposed rock surface. Slopes are variable
and average 15-20 percent.

c-i i

Carex Subalpine Meadow: Different species of Carex
cover 5-25 percent of this area. Other subdominants
are Poa spp. and Sibbaldia procumbens. Plant density
is mostly continuous, and the majority of the plants
present are less than 1/2 m tall. Scattered shrubs
attain a height of 3-4 m. Rocks comprise less than 25
percent of the entire cover9 with all diameter classes
being about equal in cover. Lichens are uncommon.
The slope gradient averages 17-18 percent.

C-12

Dryas-Pinus Subalpine Meadow: Dryas octopeta1a covers
25-50 percent of this area, with Pinus flexi1is covering up to 5 percent. Appearance of this unit is ragged,
with most plants being shorter than 1/2 m. Density of
the vegetation is principally continuous, but areas of
interrupted growth are present. Total rock coverage is
5-25 pe rcent, with all diameter classes being equally
represented. Coverage of exposed rock surfaces is less
than 25 percent, with green and black being the dominant colors. Slopes average 12 percent.

C-·13

Sibbaldia Snow Accumulation Area: This area is dominated by rocks which cover 50-75 Percent of the entire
area. No one rock diameter class predominates.
Sibbaldia procumbens is the dominant plant present,
covering 5-25 percent of the area. Vegetation in this
unit is short, less than 1/4 m in height, and the density is interrupted as plants occur singly or in small
groups. Lichens are uncommon on rocks, but a gray
lichen covers up to 5 percent of the exposed soil. The
average slope gradient is about 15 percent.

�-109-

Table

1.

Cont.

UNIT

DESCRIPTION

C-l4

Salix-Poa-Betula Alpine Meadow:
This area is similar
to C-4 except Salix spp. covers 5-25 percent of the
area and Betula glandulosa occurs only in this unit.
Most shrubs in this unit are less than 1 m tal19 but
scattered conifers extend to a height of 2 m. Density
of the vegetation is mostly continuous.
Rocks larger
than 12 inches in diameter cover 5-25 percent of the
area.
Black and green lichens are common, with each
covering up to 5 percent of the exposed surface of
rocks.
Slopes are gentle and average 5-7 percent.

C-15

Salix-Picea-Trifolium-Poa
Krummholz:
Salix spp. covers
25-50 percent of the area9 with the other three dominant genera each amounting to 5-25 percent of the total
cover. Vegetation of this unit is irregular as some
groups of conifers reach a height of 3 m~ but much of
the vegetation is shorter than 2 m. Plant density is
primarily continuous.
Rocks of all size classes are
common but make up less than 25 percent of the coverage. About 50 percent of the exposed surface of stones
is covered with green and black lichens.
Slope gradient averages 12 percent.

C-16

Artemisia-Trifolium
Snow Accumulation Area:
Species of
Artemisia (principally ~. scopulorum and ~. norvegica),
and Trifolium parryi each covers 5-25 percent of the
area, with Artemisia occurring singlY9 while !. parryi
is grouped in small patches.
plants in this unit do
not exceed 1/2 m in height, and the density is frequently interrupted.
Stones of all size classes are
common and cover at least 25 percent of the area.
Lichens are uncommon, with green and black being the
most prevalent colors.
Slope gradient averages 10-12
percent.

��-111April,

JOB FINAL REPORT

State of

C~O~L~O~RA==D~O~

Project No.

W-37-R-25

Work plan No.

19

Job Title:

Game Bird Survey
1

Job No.
Experimental

Period Covered:
Personnel:

_

Breeding

of Tinamou

April 1, 1971 to March 31, 1972

Donald M. Hoffman.

ABSTRACT

An article

covering

results

of this study was published

as follows:

Hoffman, Donald M.
1971. Raising pale spotten tinamou.
Game Bird Breeder, Aviculturists, Zoologists, and Conservationists Gazette.
Gazette, Salt Lake City, Utah.
May. pp , 17-21.

1972

��-113-

EXPERIMENTAL

BREEDING OF TINAMOU

Donald M. Hoffman

P. S. OBJECTIVE
To develop game farm production techniques for pale spotted tinamou.
SEGMENT OBJECTIVE
Compile all information, analyze data and prepare a final report.
lish in an appropriate journal.

Prepared by

/()~'
/Jd!J.

CJ11

~'

Donald M. offman
Wildlife Researcher

..~

Pub-

��-115April,

1972

JOB FINAL REPORT

State of

~CO~L~O~RA~D~O~

Game Bird Survey

W-37-R-25

Project No.

Job Title:

Study of Tinamou

Period Covered:

December

Warren

2

Job No.

19

Work plan No.

Personnel:

_

Adaptability

1, 1968 through March 31, 1972

D. Snyder

ABSTRACT

During the consecutive springs of 1969, 1970 and 1971, a total of 164 pale
spotted tinamou, Nothura darwinii, were released in national grasslands along
the Sand Arroyo drainage in southern Baca County.
To date, positive evidence
of their survival, reproduction, or existence in the release vicinity or
elsewhere has not been obtained.
Unless new information is obtained at a
future date it will be concluded that the introduction attempt was unsuccessful.

��-117-

STUDY OF TINAMOU
Warren

ADAPTABILITY

D. Snyder

P. S. OBJECTIVE
To determine the ability of pale spotted tinamou to establish
populations in areas of mixed tall and mid-grasses, sandsage,
the Comanche National Grasslands, Baca County, Colorado.

METHODS

reproducing
and yucca on

AND MATERIALS

1.

Selection

of release sites on Comanche National

Grasslands

2.

Determine

presence

tinamou.

and dispersal

of pale spotted

in Baca County.

(a)

Assistance was asked of the Forest Service personnel responsible for
management of the area to secure their help in observing tinamou.
They also maintained close liaison with persons residing in the
vicinity of the releases and with persons leasing the rangelands for
grazing concerning the tinamou releases.

(b)

Assistance of the area Wildlife Conservation
checking likely areas for tinamou.

(c)

Follow up verification
made.

checks of reported

Officer was asked in

tinamou sightings

were

RECOMMENDATIONS
Additional trial releases of this species should not be made using existing
release methods.
If future tinamou releases are attempted, a large, predator
proof pen, approximately 5 or 10 acres in size should be used to permit
species adaptation to the site, food, etc. Protected natural reproduction
should be permitted by retaining a few of the adult pairs within the pen
through the summer.
If a species of tinamou exists that is better adapted to cold and snow
conditions, as they exist in eastern Colorado, it should be tried in preference
to the pale spotted tinamou.
Use of radio telemetry tracking equipment on a
few of the birds after release is also recommended.

DESCRIPTION

OF AREA

The release sites were located along the Sand Arroyo drainage in southern
Baca County in the extreme southeast corner of Colorado.
There, within the

�-118-

Carrizo District of the U. S. Forest Service, rangelands provide some of the
best comparative tinamou habitat available in the state.
Cover types vary widely within the region from straight short grass and
reseeded mid-grass prairie through many densities and interspersions with
sandsage (Artemisia filifolia) and yucca (Yucca glauca). In most parts of
the County rangelands and pastures are interspersed with private farmland.
The reader is referred to Hoffman (1965) and Snyder (1967) for more detailed
discussions of the region's habitat.
Bump and Bohl (1965) reported a number of characteristics of the pale spotted
tinamou's native range. These characteristics are summarized in following
paragraphs along with comparative descriptions of the transplant site in
Colorado.
The native environment of the pale spotted tinamou includes dry grasslands,
pastures, open thorny scrub, savannah-like woodlands, cultivated fields and
open weedy areas in arid to semi-arid regions. Grass is considered a prime
requisite. Southern Baca County rangelands fit well into this type of habitat
description.
The species occupies level, rolling, or hilly country in South America.
Elevations range from 400 feet to 12,000 feet above sea level. Southeast
Colorado plains are level to rolling. The general elevation of the region
is about 4,000 feet.
At the release site soils are primarily sandy and alkaline in pH. Soils in
the native range vary from sandy to clayish loam. The pH level is about 6
to 8.
Precipitation varies from 4~ to 26 inches annually within the species native
range. In semi-arid regions moisture is received throughout the year. Lowest
monthly accumulations are during winter and early spring. This pattern of
dry winters becomes more pronounced in the arid regions. Drought periods are
reported as common and snow is uncommon. Southeast Colorado precipitation fits
well within this range. Annual precipitation averages around fifteen inches.
Winters are predominately dry. Most rainfall is received in late spring and
summer. Snow and blizzards are occasional and snow seldom covers the ground
for longer than a week.
o

0

0

0

Temperatures average around 85 to 93 F at maximum in summer, and 30 to 40
F at minimum in winter in the species native habitat and in Baca County.
Occasionally more severe cold periods of potential detriment to the species do
occur in Baca County. However, protective cover, wind chill indices, and other
factors have to be taken into consideration when comparing winter extremes.
The pale spotted tinamou does not require open water. Foods in native range
include seeds, fruits, flower heads, and leaves in winter. Some insects are
included in the diet during summer and fall. Waste grain and alfalfa are also
consumed. The many successional stages of range revegetation in southern Baca
County would provide a variety of similar foods for consumption. Interspersion
of farmland adds to this selection.

�-119-

RESULTS AND DISCUSSION
Three releases were made during the consecutive springs of 1969, 1970 and
1971.
Table 1 summarizes the information concerning numbers released, dates,
and locations.
All were released at guzzler sites previously installed for
scaled quail along the Sand Arroyo drainage.

Table 1. Releases of pale spotted
in southern Baca County, Colorado.

tinamou on the Comanche

National

Grasslands

Release
Number

Date of
Release

Number Released
Males
futal
Females

1

4-8-69

25

17

42

13

32S

46W

2

5-1-70

15

25

40

18

32S

45W

3

4-30-71

41

41

82

36

31S

45W

81

83

164

Total Released

Location Released
Section
Township
Range

Almost no information concerning these birds has been obtained since they were
released.
Bert Widhalm, Wildlife Conservation Officer, reported finding feathers
about two weeks after the initial release.
Personnel of the U. S. Forest
Service visited the release sites several times after the releases were made.
Searches by the writer, using a dog, were completed in and around the release
sites in June, 1969; December, 1969; June, 1970; November, 1970; and November,
1971.
Tinamou were not observed, nor was evidence of their survival indicated
by any of the observers.
U. S. Forest Service personnel received only two or three reports of possible
sightings from local residents in 1969 and 1970. Subsequent searches of the
locations were conducted, but no confirmation of the sightings could be made.
An interview of one of the observers instilled doubt as to his correct identification of the species.
His description of the bird and its habits more
closely fit the resident bobwhite quail.
No reports were received in 1971.
It is regretful that more information concerning the species could not be
obtained.
At present we do not know if the birds perished immediately,
survived until winter, attempted to reproduce, moved out of the region or are
still in existence.
The hiding abilities of the species make follow up
searches near futile.
Unless the species is sighted and positive identification
is made in the future it must be assumed that they did not survive and introduction efforts were not successful.

�-120-

LITERATURE CITED
Bump, G., and W. H. Bohl. 1965. Some tinamous of Argentina and Chile.
Interim report. U. S. Dept. of the Interior. Bureau of Sport Fisheries
and Wildlife. 15 p. Mimeo. (Illust.).
Hoffman, D. M. 1965. The scaled quail in Colorado.
and Parks Tech. Pub. No. 18. 47 p.

Colo. Div. of Game, Fish

Snyder, W. D. 1967. Experimental habitat improvement for scaled quail.
Div. of Game, Fish and Parks Tech. Pub. No. 19. 65 p.

Prepared by

1:d~e(J~
Wildlife Researcher

Colo.

�-121-

April, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-37-R-25

Work Plan No.

20

Job Title
Period Covered:
Personnel:

Game Bird Survey

1
Job No.
Investigation of Population
Status of Bobwhite Quail in Eastern Colorado

April 1, 1971 to March 31, 1972

Russell A. Kozacek, Michael Robinson, Jack Vayhinger,
Marvin Gardner, Howard D. Funk and Warren D. Snyder.

Frank Stumpf,

ABSTRACT
Whistling counts of male bobwhite quail, conducted in June and July, 1971,
were used to inventory quail in eastern Colorado.
A mechanical whistle was
used to induce calling as a supplement to natural calling.
A direct linear
relationship between the two census techniques was observed, however, the
natural calling method was deemed adequate for determining general trends.
Routes were established and all known or potential quail range was inventoried
along the South Platte drainage.
Inventory counts were also initiated in
the Republican River drainages and in southeast Colorado.
Intensive study of
bobwhite quail population dynamics were initiated on the Tamarack Management
Area along the South Platte River in northeast Colorado.
Information gathered
at the Tamarack check station indicated that 531 fall banded quail represented
25.1 percent of the total early fall population of 2,114 birds on the area.
Crippling loss of approximately 14.8 percent was added to the 486 known harvest
to project a total hunting season mortality of 558 quail or 26.4 percent of the
early fall population.
Lack of a total hunter check on the east part of the
Tamarack was considered the primary weakness in estimates of harvest.
Density
estimates indicated the early fall population exceeded one bird per two acres on
the study area. Adults comprised approximately 19.2 percent of the early fall
population, 19.3 percent of the hunting season sample and 21.1 percent of the
winter banded sample.
This latter sample included 68 fall retraps and 155 newly
banded quail. Data were calculated separately for the east and west segments
of the Tamarack Management Area.
Past harvest data indicated 1971 was a high
production year. Additional information on sex ratios, weights, movements, hunter
success and other variables was compiled.
Additional fall banding of 450 bobwhite
quail along the South Platte River above Sterling, Colorado provided supplemental
information concerning hunting pressure, harvest and population densities and production.
This phase of study has been terminated because band return and harvest
information were too unreliable to be of use. Preliminary work was initiated
on a third phase of study concerning environmental limitations to bobwhite but
no quantitative data are presented.

�-122-

RECOMMENDATIONS

1.

To improve fall population estimates two items are needed:
(1) larger
sampling on the Tamarack Management Area to approximate one-third of
the fall population; an~ (2) a more uniformly distributed banded sample
within the population.

2.

To improve harvest data on the Tamarack, regulations should be implemented making it a misdemeanor to hunt on the area without first reporting through the check station.
Signs should be placed at key locations
where hunters gain access to the Tamarack Management Area along the north
side of the river.

�-123-

INVESTIGATION OF POPULATION
BOBWHITE QUAIL IN EASTERN
Warren

STATUS OF
COLORADO

D. Snyder

P. S. OBJECTIVE
To investigate (1) the distribution and relative density of bobwhite quail
in eastern Colorado; and (2) population structure and level, and rate of
harvest, and identify key factors that limit bobwhite on the Tamarack Management Area.

SEGMENT

OBJECTIVES

1.

To investigate
Colorado.

the distribution

and density of bobwhite

2.

To investigate population structure and level, and rate of harvest
bobwhite quail on the Tamarack Management Area.

3.

To identify
Area.

key factors limiting

METHODS

bobwhite

in eastern

on the Tamarack

of

Management

AND MATERIALS

1. Counts of whistling calls made by male bobwhite in late spring and summer
were used to determine species distribution and relative densities (Kabat
and Thompson, 1965).
Initially, it was thought that because population levels
were marginal in some areas, additional stimulus to induce calling would be
needed.
A female call, recorded and played on a cassette tape player
attracted and excited males at close range, but failed to stimulate calling
at distances needed in inventory.
A mechanical whistle, obtained from
Herters, Inc. did provide the volume needed and often induced response from
otherwise silent males.
As a result, the implemented technique entailed
(1) tally of natural calling male bobwhite for two to four minutes, then
(2) induced calling of additional males was attempted by use of the whistle
for a similar period of time.
The call used to obtain whistling response from male bobwhite could not be
accurately identified by name in Stoddard (1931).
It appeared to be a form
of locational or inquiry call which was often heard in the wild.
It consisted
of five or six replicates spaced at one second intervals.
Each call started
at a constant level and ascended sharply at the end so as to appear in
diagram

as

~~.-J

would be represented

For comparison
as __

/

(bob white)

or

the regular bobwhite
/

call

(ah bob white).

�-124-

Males would respond with either of the two illustrated calls.
If the
caller remained hidden they often would approach either on foot or wing to
within a few feet.
Frequently, after initial responses to the call their
approach would be silent.
Routes were ea t ab Ld.sh ed in, or adjacent to river bottom flood plains which
contained quail habita~. and in sandsage range lands of southeast Colorado.
Listening stations wer~ spaced at least one-half mile apart.
Routes were
established at intervals along the rivers and mapped prior to census using
state highway maps.
Listening stations were pinpointed on these maps.
Counts starting at sunrise on r~latively calm mornings were used. At least
two replicates per route were obtained along the South Platte with additional
counts planned for the following year.
Forenoon or evening counts frequently
were obtained, but usually were not comparable in tally to those obtained
during the early morning.
Therefore~ they were usually not used in density
evaluations.
Irrigation motors, interstate truck traffic and other noises
frequently interfered with inventory.
Kabat and Thompson (1963) recommended call transects be conducted between
June 15 and July 4. In Colorado, most inventory counts were conducted
between June 8 and July 3. For evaluation purposes, weekly replicates were
conducted on the west Tamarack route from late May through mid-July.
These
indicated high counts could be obtained into the mid-July period in northeast Colorado.
In marginal quail ranges of the upper South Platte River census efforts were
supplemented with landowner and Conservation Officer contacts to obtain
information on possible existence of bobwhite.
Increased use of the whistle
to induce calling was also attempted in marginal quail habitat.
2. Population structure and level, and rate of harvest were determined
use of the following techniques:

by

Early fall trapping and banding was initiated in mid-September.
A majority
of the trap sites were cleared of vegetation with a tractor-mounted
rototiller and pre-baited with whole corn and milo.
The combination of cleared
edge area, cultivated ground, and bait was considered of value in attracting
quail coveys.
Trap sites were located along the property as shown in Fig. 1.
Time limitations and lack of access to some areas on the north side of the
river prevented trap distribution throughout the entire river bottom.
Fluorescent flagging tape was used to mark trap locations.
Pre-baiting for a few
days to a week or more was employed in nearly all instances.
Small welded wire funnel type bait traps were used. These, in part, were
replaced as the trapping season progressed, with circular collapsible nettopped traps designed to reduce the occasional scalping problem.
All traps
were covered with tree limbs,'leaves, etc. to reduce avian predation.
Traps
were usually checked between 8:30 and 10:00 A. M. and again starting approximately one-half hour before sunset.
Quail were removed from the trap and
held in a gunny sack until banded and released.

�-125-

-'
&lt; ~

w

V)

u

o

w
ex::

«
«

Z

C)

w
ex::

w

\--

~

:s:
o
«

D

w
L

Fig. 1. The Tamarack Management Area river bottom showing vegetative cover
types, and fall (1971) trap sites.

�-126-

Consecutively numbered aluminum leg bands were placed on all quail.
Immatures were distinguished from adults by the appearance of the upper
coverts (Taber 1960). Yearling adults could frequently be distinguished
from older birds during the fall, prior to the completion of molt.
Yearling adults contained both buffy-tipped old feathers and new solid
gray replacements, whereas both the old and new feathers of the upper
coverts were solid gray on the older birds.
Aging of juveniles by the primary feather molt technique (Petrides and
Nestler 1943) was attempted d~ring the late fall, however, molt had
progressed beyond the point where meaningful data on peak of hatch could
be obtained.
This effort will be expanded in subsequent years of fall
trapping.
Quail that were not retrapped at the immediate initial banding location
were recorded for movement.
Trap sites were plotted with aid bf aerial
photos for future reference.
Bag checks of hunters were obtained throughout the 1971 hunting season
by Division personnel at the Tamarack check station.
Specific locations
of harvested birds could not be obtained since hunters frequently hunted
more than one area. Age 9 sex, weight, crippling loss, crop samples and
band numbers were obtained along with numbers of hunters and hours hunted.
Since pheasants, rabbits, ducks, squirrel and quail could all be hunted,
effort was not directed to delineate the actual hours consumed in quail
hunting.
A few hunters, who did not report through the Tamarack check
station, turned in bands and were queried by mail or phone to determine
the number of banded and unbanded quail they had harvested.
Pre-hunting season trapping and banding was also conducted on two public
and three privately owned areas along the South Platte river between
Sterling and Hillrose.
The location, size, and other characteristics of
these will be listed in the Results section.
Trapping and banding
techniques followed those used on the Tamarack except a tractor was not
employed in site preparation.
Roving checks of hunters were conducted on the sites during the first
and second weekends of the 1971 quail season.
Hunter success, age, sex
and band return information was obtained along with information on
hunting pressure.
Mail in returns provided most of the information concerning quail harvest from the sites.
Winter trapping and banding was initiated on the Tamarack in mid-February,
1972, and terminated in late March.
Techniques were repetitious of
those used in fall except site preparation was not necessary.
Special
attention was directed toward retrapped fall banded birds and in extending the trapping effort into locations not previously trapped.
Direct proportional and change-in-ratio techniques described by Petrides
(1954), Selleck and Hart (1957) and others, were used in calculation of
population levels, rate of harvest and qther variables.
Robson and
Regier (1964) was used in interpreting the adequacy of sample sizes.
Dr.
David C. Bowden and Howard D. Funk reviewed analytical procedures.

�-127-

3. Key fact~rs limiting
methods:
Comparison
made.

of trapping

bobwhite were identified

success

by use of the following

on grazed and ungrazed

river bottom were

Comparisons of sex ratios of adult quail were made as a possible
indicating extensive predation on nesting hens.
Quail utilization
observed.

of habitat

types at various

DESCRIPTION

clue

times of the year was

OF AREA

;:;enera1Species Rarlge in Colorado
Bobwhite quail in Colorado are confined by climate and habitat limitations
to two major cover types, (1) eastern Plains river bottoms and (2) sandsage (Artemisia fi1ifo1ia) grasslands with or without interspersed farmland along the eastern edge of the state.
Some of the better river bottom
flood plain habitats contain population densities equal to those in the
primary quail range of southeastern United States.
But most bobwhite
range in eastern Colorado is marginal.
Most of the sandsage rangelands
contain only scattered coveys confined to localities by old homestead
plum thickets or similar cover which can sustain them through the winter
months.
It is worth noting that winter cover requirements are not as
critical in the southeast corner of the state as in the northeast.
Again,
winter climate seems to make the difference, so the highest rangeland
populations are found in the Southeast.
Both the Arkansas and South Platte rivers, which flow east across Colorado
from the mountains, support varying bobwhite densities at lower elevations
within their unfarmed flood plains.
Along the South Platte, this habitataverages between one-fourth and one-half mile in width.
Along much of the
Arkansas, intensive farming has pushed this into an even more narrow strip
in many areas. The Tamarack Management Area, along the South Platte river
near Crook, Colorado typifies excellent ungrazed river bottom in the northeast part of the State.
Scattered cottonwoods (Populus sargentii) give
much of the area a savanna like appearance.
Snowberry (Symphoricarpos sp.),
wild rose (Rosa sp.), willow (Salix sp.), poison ivy (Rhus radicans),
golden currant (Ribes aureum), indigo bush (Amorpha fr~cosa)
and others
provide essential shrub and brush cover.
Cordgrass (Spartina pectinata) provides the primary ground cover in some
areas. Phragmites communis, of tall reed like character forms dense cover
in local patches.
Numerous annual and perennial forbs of varying densities
provide food and cover.
Occasional flooding retains early successional
vegetative stages in much of the bottomland.
Vegetative composition along the Arkansas river is markedly different.
Cottonwoods and willow are abundant, but tamarisk or salt cedar (Tamarix
sp.) is the primary woody plant of the flood plain.
Greasewood

�-128-

(Sarcobatus vermiculatus) and four-wing saltbush (Atriplex canescens) are
often found in unfarmed areas adjacent to the flood plain. Salt grass
(Distichlis stricta) and a variety of other grasses and forbs provide
ground cover.
In the Arkansas valley and its tributaries, low densities of bobwhite are
found throughout much of the irrigated farmland and along the irrigation
canals which feed this farmland. Therefore, the species range may be
several miles in width in places. With few exceptions, bobwhite are nearly
exclusively confined to the unfarmad flood plain along the South Platte.
Bobwhite populations along the headwater tributaries of the Republican
River drainage in eastern Colorado usually are Low in density as a direct
result of deficiences of brushy vegetation. Flood plains along these
streams are narrow and grazed. Some brush cover such as skunk bush
(Rhus trilobata) , exists along the breaks bordering the North Fork and
Arikaree drainages. Quail densities are increased as a direct result.
Moderate quail densities, both scaled quail and bobwhite, are found in
sandsage rangelands of Baca County in extreme southeast Colorado. The
Cimarron River, which cuts through the corner of the state, is included in
this rangeland area which comprises some of the best general quail range in
the state.
Tamarack Management Area
This public hunting area, along the South Platte river, was utilized for
intensive study of population dynamics. Highway 25, south of Crook bisects
the area so that approximately 8.25 miles of river bottom lies to the
southwest and 5.25 miles lies to the northeast. Additional Tamarack
property continues east of Hunting Area 11 East (Fig. 1), but access
problems excludes this from hunting and use in this study. One mile of
river bottom, just east of Highway 25, remained closed to hunting in 1971
and received moderate grazing through the year. The remaining land in
the east and west segments was respectively divided into 11 and 25 hunting
areas. Most were approximately one-fourth mile wide and consecutively
numbered (Fig. 1). All were open to hunting except areas 16 and part of
17 in the west segment during 1971.
Hunters were required to check in and out through a central check station
and selected their hunting areas on a first come basis. One hunting party
was permitted per area. During the 1971 small game season hunters could
hunt adjoining areas on foot after 9:00 A. M.
The majority of the overall hunting effort was directed toward ducks.
Quail, pheasants, mule deer, white-tailed deer, doves, rabbits and squirrels
rank in approximately that order of importance as secondary species.

�-129-

RESULTS AND DISCUSSION
Distribution

and Density of Bobwhite

Quail

The whistling call of bobwhite, used as an index, appeared to be an effective technique for determining distribution and relative density of bobwhite
in eastern Colorado.
Comparison of natural calling and induced calling
indices, Fig. 2, indicates either or both can be used, but the natural
calling index is adequate for determining general popuiation trends.
Some
benefits of the induced calling method may be obtained where existence of
quail in a locality is extremely low or uncertain.
Table 1 summarizes counts of whistling males in 1971 conducted throughout
the known range of the species along the South Platte River.
As the table
indicates, population levels were consistently higher between the Nebraska
state line and Fort Morgan, than those above Fort Morgan.
Call counts failed
to indicate the existence of bobwhite upriver from Hardin, Colorado in eastern
Weld County (Fig. 3).

Table 1. A summary of bobwhite quail whistling
South Platte River during 1971.

counts completed

along the

Location

No. of
Counts

Number
Stops

Natural
Tally

Mean

Induced
Tally

Mean

Stateline-Ovid

2

20

78

3.90

107

5.35

Red Lion-East
Tamarack

2

20

67

3.35

96

4.80

West Tamarack

6

75

219

2.92

227 };)

3.50

Atwood-Merino

2

20

108

5.40

130

6.50

Balzac

3

28

55

1.96

67

2.40

Brush-Fort

Morgan

2

20

77

3.85

101

5.05

Schaeffer

Ranch

2

20

37

1. 85

49

2.45

Goodrich-Orchard

2

20

23

1.15

31

1.55

Masters-K-4

2

8

9

1.13

13

1.63

Hardin-Kersey

2

20

0

1/ The total number

0

of stops for the induced tally was 65 instead of 75.

�-130-

7

a:

Atwood-

o Lyon

6

Purgatorie-Ft.

~Stateline-ovid
~rush-FtoMorgan
c:)ast TamarackRed Lion
0Campo EastCimarron

0West

Schaeffer
~

Tamarack

Ranch

Balzac

Masters K-4

E. pro~s
County

0

GoodrichOrchard

1

O

South Fork
of Republican R.

o

1

2
Natural

Fig.2. The relationship
by male bobwhite quail
in 1971.

3

4

Bobwhite Quail Whistling

5
Rate

between natural and induced rates of whist! ing
per station along Eastern Colorado census routes

6

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SOUTHPLATTECENSUSROUTE;S
StatelineOvid
8.
Gooc1tlich - Orchard
Red Lion
9. Masteirs K-4
West Tamarack
10.
Hard£n - Kersey
Atwood - Merino
11.
Mil1*en
Balzac:
12.
P1at~-eville
Brush
Con Schaeffer

j

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�-132-

Personal communications with Charles Brown (Wildlife Conservation Officer),.
local farmers, and others, indicate a few bobwhite do survive along the South
Platte in the Hardin-Kersey area east of Greeley. Earl Downer (Area Supervisor) reported a local small population along the river south and west of
Platteville in southern Weld County in past years, but no sightings during
the past two years.
Lack of adequate access along the river in the Eagles Nest Ranch vicinity
between Hardin and Masters in eastern Weld County prevented adequate inventory, but reports indicate low to moderate quail densities do exist there.
Completed summaries of counts obtained in the Republican, Arkansas and
Cimmaron drainages were considered inadequate for presentation at this time.
Range boundaries and relative densities in these areas are pending until
surveys are completed during the forthcoming year.
Projected birds-per-square mile density estimates were made for the intensively studied Tamarack Management Area as will be shown later in Table 6.
Attempts to relate these estimates to call indices and densities in various
parts of the species range in Colorado will not be attempted at this time.
Population Structure and Level, and Rate of Harvest
Tamarack Management Area
Fall Banding-- Table 2 summarizes the fall banding results wherein 317 bobwhite were banded along the south side of the South Platte on the west
Tamarack and 214 were banded along both sides on the east Tamarack. Observed
trapping mortality was very low, with subsequent survival of banded birds
considered nearly equal to that of unbanded birds for analysis purposes.
Coveys undoubtedly existed in every hunting area during the fall trapping
period even though no quail were caught in a few areas.
Harvest--The 1971 quail season in..
.nort he as t; Colorado ran from November 27
through December 5 and again, from December 22 through January 2, 1972, in
direct conjunction with pheasant season. Sunrise to sunset shooting hours
were the rule, except for the noon opening on the first day of season. Bag
and possession limits were eight and sixteen, respectively. Harvest of quail
by age, sex, and day of season are presented in Table 3. Harvest of other
small game is also provided for comparison purposes. Hunting seasons for
rabbits, squirrels, and watetfowl extended beyond the dates given for quail.
Harvest Sample--Hunters reported 371 bobwhite through the Tamarack check
station during the season. Follow-up inquiries from mailed band returns
indicated at least 115 additional bobwhite were taken on, or adjoining the
Tamarack, but not reported through the check station. This unchecked harvest
primarily occurred at two locations: (1) along the north side of the river
east of Crook in all areas (1 through 11), and (2) in the vicinity of the
Proctor river bridges at the west end of the Tamarack. A few additional quail
were undoubtedly taken and not reported. Efforts will be made to curtail
unchecked hunting on the area in future years.

�-l33Table 2. A summary by area of fall banded bobwhite, hunter use, banded harvest
recoveries and total harvest on the west and east sections of the Tamarack
Management Area in 1971.

Area

Number Quail
Banded

Hunter
Hunt e rs !I Hours 1/

Number

Total Quail
Harvest y

Banded Quail
Harvest '}j

West Tamarack
87
106
78
82
81
62
60
63
67
95
156
98
132
81
132

3
3
0
3
9
3
7
4
0
8
8
5
14
1
23

2
0
0
0
1
0
2
3
0
1
7
0
0
0
3

30
46
43
57
59
47
46
58
23

85
174
133
250
276
176
210
203
104

0
10
17
27
21
22
17
18
32

0
0
3
11
3
13
5
1
7

872

2,991

255

62

11
19
7
8
11
30
11
34
21
3
1

3
7
2
6
4
13
3
9
8
2
0

12
13
14
15
16
17
18
19
20
21
22
23
24
25 ~/

22
7
6
25
35
4
6
22
15
0
0
0
0
0
27
11
0
4
0
24
14
21
17
13
45

25
27
32
26
31
23
26
30
37
39
37
38
31
26
35

Total

317

1
2
3
4
5
6
7
8
9
10
II

-- ~/

East Tamarack
1
2
3
4
5
6
7
8
9
10
11

31
1
8
6
9
37
13
51
34
0
0

47
49
35
33
33
47
28
34
45
30
29

ll8
142
106
69
77
132
69
164
125
93
101

�-134-

Table 2. A summary by area of fall banded bobwhite, hunter use, banded
harvest recoveries and total harvest on the west and east sections of the
Tamarack Management Area in 1971 (continued).

Area

Number Quail
Banded

Number
Hunter
Hunters 11 Hours 1/

Total Quail
Harvest ~/

Banded Quail
Harvest 1/

East Tamarack (Continued):
Braddock
Pasture
Armstrong
Ranch

24

O§j

o

o

0

o

75

4

Total

214

410

1,196

231

61

Combined
Total

531

1,282

4,187

486

123

1/ Includes only known hunters and hours reported through check station.
Some hunting probably went unrecorded.
1/ Quail were not necessarily shot in the exact area to which they are
assigned in the table.
4/ Figures include area 25 and land adjoining Proctor Road.
S/ Area 16 and most of 17 were closed to public hunting.
~/ Braddock land closed to public hunting.

2/ Includes only known harvest.

Wounding Loss--Most of the hunters were queried concerning the number
of birds wounded but not retrieved. Responses, if correct, indicate 49
were lost for 330 bagged. This percentage loss was 14.8 percent. Direct
proportional projection would indicate that 72 were not retrieved, bringing
total harvest mortality to 558 birds.
Hunting Effort and Harvest by Area--Table 2 provides a general index
of hunting effort and quail harvest by hunting area. The data must be
considered indicative rather than accurate for several reasons. First,
many hunters covered more than their one assigned area in pursuit of quail.
Therefore, many of the quail were not necessarily shot in the area to
which they are assigned in the table. Second, differences in duck hunting
qualities among the areas was the primary reason some received much more
hunting pressure than others. Breakdown of hunting hours by species on an
area basis was not attempted. We can, therefore, only assume that man hours
of hunting efforts for quail were, in part, in direct proportion to the
total hours of effort per area.

�-135Table 3. Harvest by day during the 1971-72
Tamarack Management Area. !/

Ducks2/ Rabbits Squirrels

quail and pheasant

Adult .Adult
Male
Female

season on the

Bobwhite Quail
Imm.
Imm.
Male Female
uncI. Total

Date

Pheasants

11-27

17

22

6

o

5

2

21

9

16

53

11-28

18

13

2

1

8

6

15

20

5

54

1

2

o

o

1

2

10

9

9

31

11-30

1

1

1

o

o

o

7

4

o

11

12-1

1

1

1

1

3

4

4

6

o

17

12-2

o

o

o

7

1

1

4

3

o

9

12-3

4

15

o

o

3

1

21

10

o

35

12-4

9

13

9

o

4

1

13

14

16

48

12-5

4

7

1

o

1

3

9

13

13

39

Extended

Season

12-22

8

12

1

o

1

1

1

5

o

8

12-23

6

9

o

o

3

4

10

6

o

23

12-24

1

7

4

o

o

2

2

3

2

9

12-25

o

o

o

o

o

o

o

o

o

0

12-26

2

6

o

o

o

o

o

o

2

2

12-27

o

22

o

o

o

o

3

o

1

4

12-28

4

14

1

o

o

o

3

4

8

15

12-29

4

25

2

o

o

1

5

8

o

14

12-30

5

15

2

o

1

2

3

7

1

14

12-31

o

4

o

o

1

o

1

1

3

6

1-1

1

2

o

o

o

o

1

o

6

7

1-2

1

6

o

o

o

o

1

4

7

12

Total

87

196

30

9

32

30

134

126

89

411 3/

1/ Seasons

for waterfowl, rabbits, and squirrels extended beyond those listed here.
seventy of the harvested ducks were mallards.
Late repo rted-harves t data along Armstrong Ranch is not; included.

2/ One-hundred

II

�-136-

A breakdown of hunter effort by species will probably be attempted in the
future.
But it must be realized that any attempt can.be only partially
accurate because of variable types of hunting pursuit.
Four general catagories listed as follows might be most informative:
(1) exclusive quail
hunter, (2) upland game hunter, (3) exclusive waterfowl hunter, and (4)
combined upland and waterfowl hunter.
The average hunter expended approximately 6.2 hours per animal bagged.
Since the exact hours of effort per species is unknown the average time
required to bag one quail or one mallard is also unknown.
However, observations indicated bobwhite quail probably yielded the greatest return in
birds per hunter effort.
Comparison with Previous Years of Harvest--Harvest
information
presented in Fig. 4 indicates 1971 was an excellent production year for
bobwhite quail on the Tamarack.
The 1971 harvest was indicated to have
been surpassed only twice during the previous 19 years.
Information fot
1965 is lacking but production failure and poor harvest were known to
follow extensive June flooding of the river bottom that year.
Northeast region total harvest and average season bag projections taken
from Colorado's Small Game Hunter-Harvest Surveys are illustrated in
Fig. 4. These data, along with information from the Tamarack, suggest
that quail populations in northeastern Colorado fluctuate dramatically
from year to year.
Projected Small Game Hunter-Harvest Survey data show an upward trend in
hunting pressure on bobwhite in northeast Colorado.
Hunter numbers
averaged 1,600 per year from 1956 to 1960, 1,672 during the 1961-65
period, and 2,022 during the 1966-70 period.
This is in direct contrast
to a downward trend in hunting pressure on pheasants in Colorado during
the same time interval.
Winter Banding--Trapping
and banding efforts, commencing in midFebruary and terminating in late March, resulted in banding of 155
bobwhite and retrapping of 68 fall banded birds.
Mild snow-free weather
with an apparent ample supply of natural food made trapping more difficult than expected.
The birds also were increasingly wary following
hunting season.
Trapping efforts were distributed along both sides of
the river in a fairly intensive coverage of the area. Reduced size of
covey wintering home ranges also made intensified trapping efforts
essential.
Fig. 5 illustrates the locations of coveys known to exist
along the Tamarack during the late winter.
Other coveys may have been
present.
The Fall Population Estimate--Fall banding and harvest data were
used in deriving fall population estimates for the west Tamarack (1,304)
and the east Tamarack (810) as shown in Table 4. The estimates from the
two sections were then combined to provide a total estimate of 2,114 quail.
Direct calculation from the last column of figures in Table 4 results in
a slightly different estimate, 2,098 quail, because of sample size differences on the east and west sections of the study area.

�•.l37Harvest

Bobwhite Quail
I-'
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0

0

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0

0

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0

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,s:..
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1952
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1956

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1959
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1962

w

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1966
1967

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Thousands of Bobwhite Quail Harvested

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MANAGEMENT

AREA

~~

111 (
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LEGEND
1

HUNTING AREA

_

LOCATION
OF
~~- 1971-72 COVEYS
.

~

~UNFARMED
RIVER BOTTOM
~
AND RIVER CHANNELS.

o

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,

�-139-

Table 4. A summary of fall, 1971 bobwhite quail banding
information from the Tamarack Management Area.

and harvest

West
Tamarack

Tamarack

Total
Area

317

214

531

Known harvest

255

231

486

Crippling

38

34

72

Item

Number

of quail banded

Percent

loss (projected)
crippling

Projected

14.8

loss

293

265

558

(harvest)

62

61

123

band recoveries

71

70

141

total harvest

Band recoveries
Projected

EMt

mortality

Percent

harvest

22.5

32.7

26.4

Percent

banded

24.3

26.4

25.1

Projected

fall population

1,304

810

2,114

Different fall populations could be derived by combining winter trapping
information with harvest samples.
On the west hunting areas the fall estimate would be increased from 1,304 birds using only the harvest sample to
1,338 birds using the combined harvest-winter
sample.
In contrast, on the
east study group, the population estimate would drop from 810 to 697 when
the winter banded sample was added to the harvest sample.
Or the total
fall population estimate for the Tamarack would be 2,035 quail based on
combined harvest-winter
trapping samples.
Estimates presented in Table 4, which exclude the winter trapping sample,
are considered more accurate for the following listed reasons:
(1) The
fall banded sample was not randomly distributed in the population prior to
hunting season nor prior to winter trappin$ because fall trapping was
segmented and nearly all coveys remained sedentary from fall to winter.
This made it more essential that recapture samples be as randomly taken
as possible.
(2) Hunting, especially on the east group of areas, was
believed more evenly distributed or more random than winter trapping was.
For example, hunters gained access along the Armstrong ranch where neither
fall nor winter trapping was attempted.
Winter trapping was probably
biased toward areas where coveys had been banded in fall. Therefore,
hunters probably more uniformly sampled the total population.

�~140~

Confidence Limits and Sample Size--A formula presented by Davis
(1960) was used to obtain confidence limits for the fall population estimates.
These are presented in Table 5. Since the Tamarack population
could be considered finite, a finite correction (Snedecor, 1956) was
applied reducing the confidence limits (Table 5).

Table 5. Fall population estimates for the Tamarack
confidence intervals at the 95 percent level.

Management

Area with

Population

Population
Estimate

Confidence
Interval

Confidence Interval
with Finite Correction

West Tamarack

1,304

± 288

± 258

East Tamarack

810

± 178

± 150

2,114

± 327

± 286

Total area

The primary weakness contained within the 1971 fall population estimate
appeared to lie within the randomness of sampling.
More even distribution
of fall banding would reduce this error and make subsequent recapture
sampling on a completely random basis less essential.
A second weakness
resulted from inadequate sample size when harvest data alone were used
as the recapture sample.
Robson and Regier (1964) state that sample sizes
should be sufficient to prevent errors in population estimates greater
than 10 percent with 95 percent accuracy in population dynamics studies
of the type presented here.
According to information presented in their
text, fall banding and harvest samples in this study were only sufficient
to prevent errors greater than 25 percent with 95 percent accuracy in
estimating the population.
Fall banding of a least a third of the
population would be needed in future years of study.
Projected Fall Density--Aerial photo measurements indicated that the
Tamarack river bottom from the east edge of Hunting Area 11 East to the
Proctor bridges occupied approximately 13.5 linear miles (Table 6). Unfarmed
flood plain habitat, which seldom exceeded a half mile in width, occupied
approximately 6 square miles.
Bobwhite quail were confined to the timbered
bottom land by farm lands and short grass pastures along the entire west
part of the Tamarack (Fig. 5). A few birds were observed moving into the
sand sage rangelands and up along tree plantings bordering the east part
of the Tamarack during the spring, summer and fall. But in winter, coveys
were again confined to the timbered, brushy river bottom.
Feeding in
adjacent corn or beet fields was restricted to the edge of those fields.
Therefore, the density-per-square-mile
figures presented in Table 6 are
believed fairly accurate.
This density, exceeding one bird per two acres
during the fall of 1971, compares favorably with quail densities in the

�-141-

species primary range in southeast United States.
Fall densities there
of a bird per acre or more have been attained, but usually occur only
under careful management.

Table 6. Basic statistics
quail habi tat.

on Tamarack

Management

West Tamarack

Item

Area river bottom

East Tamarack

Total Area

Length in miles

8.25

5.25 1./

13.5

Average width in miles

0.44

0.46

0.45

Approximate

3.601:./

2.44

6.04

square miles
estimate

1,304

810

2,114

per square mile

361

332

350

Fall population
Bobwhite

-1/ Includes

one mile of unhunted

pasture

-2/ Includes river channel which averages

east of Highway

25.

200 to 250 feet in width.

Natural Fall Mortality--The percentage harvest shown in Table 4 is
based on population levels existing during the period of fall banding.
But natural mortality occurred between the period of banding and the
start of hunting season.
On the west hunting areas, 66 days elapsed from
the mean banding date, September 22, to the start of quail season, which
was on November 27. On the east hunting areas the time lapse was 47 days
(October 12 to November 27). The actual rate of mortality during the fall
is unknown.
Kabat and Thompson (1963) used a 2 percent mortality rate per
week for bobwhite up to 16 weeks of age. Rosene (1969) found a nest to
hunting season mortality rate of slightly over 3 percent per week.
Klimstra {195~calculated
a 4 percent weekly loss in birds up to eight
weeks of age. Robel (1965) found evidence that fall to winter mortality
of young increased in December when snows and cold temperatures put
greater stress on the population.
Nearly all young quail exceeded eight
weeks of age prior to banding and hunting season opened prior to the
December stress period in 1971 on the Tamarack.
Therefore, a weekly
mortality rate of 2 percent was hypothesized for the banding to harvest
interval.
A hypothetical model was constructed, based on the above information, to
illustrate fall population change from banding to harvest.
This information is presented in Table 7. As shown, 224 birds or 17.2 percent of the
early fall population would have been lost prior to hunting season on the
west part of the Tamarack.
On the east part, 106 quail or 13 percent of

�-14:2-

the original

population

would have died.

This hypothetical

removal

of

330 birds on the Tamarack would leave 1,784 alive at the start of hunting
season.
Thus. projected harvest mortality'WCulcl
aGtu~lly
percent of the population surviving just prior to hunHnl

h~ve removed'lL
SHM.i'llH'.

3

(l'~l:lle&gt;1).

Table 7. An illustration of population decline from the mean early fall
banding date to the start of hunting season based on a hypothetical
natural mortality rate of two percent per week.

Tamarack
Item

Mean banding

date

Interval to season opening
(Days)
(Weeks)
Starting

population

Natural

mortality

Population

Management
East

Area
Total

Sept. 22

Oct. 12

Sept. 29

66
9.4

47
6.7

West

removal

at season opening

Rate of projected total harvest
Based on early fall population
Based on season opening

1,304

810

2,114

224

106

330

1,080

704

1,784

22.4%
27.1%

32.7%
37.6%

26.6%
31.3%

Age Ratios--A summarization of age ratio data obtained during the fall
and winter banding and during the hunting season on the Tamarack is presented
in Table 8. This table leaves a couple of questions unanswered.
First, does
hunting provide an unbiased sample of the existing population?
According
to Leopold (1933), Robel (1965), Rosene (1969) and others, it does. Assuming
that the harvest sample is unbiased, a second question arises:
Why is there
a highly significant lower percentage of adults in the fall banded sample
(Table 8) than in the harvest and winter samples (Chi sq. = 10.44 df = I)?
Either biased fall sampling existed or differential mortality following the
banding period removed a greater proportion of young than were present in
population.
Highly significant divergence of age ratios between the banded
and unbanded segments of the harvest sample indicate that early fall trapping probably was selective toward juveniles (Chi sq. = 8.89 df = 1). The
banded harvest sample was similar in ratio to the fall banded sample,
whereas the unbanded harvest sample showed a much higher proportion of
adults.
Little evidence of differential juvenile mortality was indicated
during the pre-hunting season period.
Instead it is hypothesized that the
fall population actually contained approximately 19.2 percent adults instead
of 11.5 percent as shown in the fall banded sample.

�-143-

Table 8. Bobwhite quail age ratio information obtained
Management Area during the fall and winter of 1971-72.

on the Tamarack

Number
Adults

Number
Young

Percent
Adults

Adu1t:Yearling
Ratio

sample

61

470

11.5

1:7.705

season sample

62

260

19.3

1:4.194

Banded harvest

15

114

11.6

1:7.600

Unbanded

harvest

47

146

24.4

1:3.106

fall age ratio

112

470

19.2

1:5.217

47

176

21.1

1:3.745

Item

Fall banded
Hunting

Projected

Winter banded

sample

Age bias in trapping was not evident in late winter when all quail had
reached maturity.
Some indication of differential mortality is illustrated
in Table 8 by the reduced percentage of young in the population.
But
changes in age ratios from harvest to late winter were not significant
(Chi sq. = 0.40 df = 1). If age bias toward young did occur during winter
trapping then differential mortality of young would be higher than indicated.
Sex Ratios--Kabat and Thompson (1963) provided data and cited numerous
references showing that while juvenile sex ratios were usually near equal,
adult males consistently exceeded females in the population.
In most cited
studies approximately 60 percent of the adult population was composed of
males.
Stress on the hen during the reproductive period was repeatedly
listed as the probable reason for this divergent adult sex ratio.
Sex ratios, obtained during the fall and winter studies in Colorado
(Table 9), suggest some excess of males but samples are inadequate to
be certain at this date. Adult males did not reach the 60 percent level
shown in many other studies.

Table 9. Sex ratios of adult and immature
fall and winter banding and harvest.

quail obtained

during

Inun. Male

Inun. F emal e

28

179

216

31

28

139

128

banding

25

22

94

82

79
46.7%

412

426

ratio

89
53.5%

Sample Period

Adult Male

Fall banding

33

Hunting

season

Winter
Total
Adult

bobwhite

Adult Female

�-144- .

Weights of Bobwhite Quail--A large sample of bobwhite quail taken on
the Tamarack Management Area were weighed at the check station.
Results,
illustrated in Table 10 indicate juveniles were near equal in size to
adults and a size difference between sexes was not evident during the
hunting season.
Sinc~ the specimens were taken by hunters at various
times of the day and contained varying amounts of shot, blood and food,
the tabular data should be considered proximal rather than highly accurate.

Table 10.
Management

Weights of 163 bobwhite quail collected
Area during the 1971 hunting season.

UncI.

Sample

size

Mean weight
(in grams)

Adult Male

Adul t Female

on

the Tamarack

Imm. Male

Imm. Female

37

IS

14

69

6S

193.4

202.7

203.7

202.1

199.S

Fall and Winter Movements--Habitat
restrictions bordering the Tamarack
prevented quail movements except in upstream or downstream directions.
In
other words, the Tamarack population was isolated except at the upper and
lower ends. This reduced ingress and egress movements, so that the existing
population could be considered more finite 'than if it was bordered by buffer
quail populations on all sides.
Accuracy of population estimates was
believed improved as a result.
Numerous studies of bobwhite quail movements have been published.
Therefore,
it was not deemed essential to conduct additional intensive movement studies.
Fall and winter trapping, banding, retrap and harvest data generally support
data from previous studies.
Preliminary conclus.ions reached in this year
of work include:
(1) most of the fall shuffle, if there was one, occurred
prior to commencement of all trapping; (2) nearly all coveys banded in fall
were still present in the same vicinity in late winter; (3) covey territories became smaller as winter approached; (4) .sustained hunting pressure
through the season did not shift coveys into new permanent home ranges
to any noti ceable degree; and (S)late
fall or winter movements of coveys
to areas adjoining cornfields were not in evidence.
Such movements might
have occurred if snow accumulations had created a food stress.
Fall banding to fall retrap data did not produce evidence of movements
exceeding one mile.
Fall banding to harvest data were not of much value
because hunters did not necessarily stay in the specific area to which they
were assigned.
However, significant quail movements of a mile or more
could have been detected, but were not. Winter retrapping of 68 fall
banded quail showed nearly all quail were stable in the area from fall
through late winter.
One quail had moved from Area 8 West to Area IS West
or approximately
two miles.
Some coveys occasionally to frequently crossed

�-145the river channel(s).
Other coveys seldom, if ever, crossed to the other
side. To illustrate the latter point, trapping efforts failed to produce
quail in fall along the south side of the west Tamarack in Areas 10 through
14. Late winter trapping in Areas 9 through 13, across the river on the
north side, resulted in banding of 34 quail from at least six small coveys.
None of these birds had been banded previously indicating that they probably
were sedentary in their territories throughout the fall and winter.
Late winter baiting and trapping efforts failed to locate quail in several
areas where they were known to exist during the fall. In some areas quail
may have existed in small interior territories, but this was not believed
to be the general case. Searches for tracks were usually futile due to
lack of snow. Winter banding produced some evidence that these coveys had
shifted a half mile or so either upstream or downstream.

Upriver

Study Results

Site Selection--The Davies easement and the Jones property, both Division
administered properties, were utilized for fall banding-harvest
studies.
Additional properties owned by Keith Propst, John Bianco and Mr. Lutes
were also utilized.
These private landowners posted their land and
reportedly permitted little or no hunting.
It was hoped, that these
private lands would provide measurements of differences in population
characteristics between areas with little or no hunting and those which
received more intensive hunting pressure.
Locations and approximate sizes
of the sites are illustrated. in Fig. 6.
Fall Trapping, Banding and Harvest--Summer whistling counts above Sterling
along the Atwood-Merino and Balzac routes (Table 1) indicated quail populations were as high or higher in some areas there than they were on the
Tamarack Management Area. Fall trapping efforts confirmed this with one
exception.
The Lutes Property, which received moderate to heavy grazing
was deficient in brushy habitat and lacking in quail.
Only one or two
small coveys were found along the area. Trapping efforts were cut short
due to lack of quail on the property.
Table 11 summarizes the results to trapping efforts conducted from midOctober to mid-November.
Age and sex ratio information for the sites is
summarized in Table 12. Hunter contacts, conducted the first and second
weekends of season, provided only a partial sample of weekend hunting as
hunters on all plots could not be contacted at the time they terminated
hunting on a particular area. Hunting and harvest on the sites through
weekday periods went unobserved.
The majority of the quail bands were
mailed in. A return letter was sent to the hunter informing him concerning when and where his quail was or were banded and thanking him
for sending in the band number.
~ercentage Harvest--Discussions
with numerous hunters revealed
information indicating some discarded the bands rather than turn them
in. The percentage return was unknown so percentage harvest also remained
uncertain.
The Davies easement was heavily hunted.
On the opening weekend one to two dozen vehicles were consistently parked around this area
which contained approximately a section of land. Yet information on only
eight bands was obtained (Table 11). In contrast a much higher band

�(IJ

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CD

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�-147-

return rate was recorded on the Propst land which sustained a much lower
rate of hunting pressure.
Only one or two parties were obserVed using
the property at anyone
time. The Jones property which contains less
than 160 acres of timbered bottom land apparently sustained the highest
rate of harvest with nearly half of the 34 bands returned.
Spring movements of quail from adjoining less intensively hunted river bottom would
replenish the breeding population on this small area. Fear of overharvest therefore is probably unwarranted.

Table 11. A summary of bobwhite quail banding, band return and harvest
information from locations banded above Sterling during the fall of 1971.

Banded

to Unbanded
Ratio

Percent
Banded

Location

Number
Banded

Number of
Returns

Percent
Return

Bianco

202

6 to 50+

23.3

47 of 70+

50+

Jones

34

15

44.1

5 of 23

25-30

Propst

127

22

17.3

10 of 22

4.0-50

Davies

79

8

10.1

No data

Table 12. Age and sex ratio information
Sterling during the fall of 1971.

on quail banded

Imm. Males

upriver

Imm. Females

from

Total

Area

Adult Males

Adult Females

Bianco

13

10

84

95

202

Propst

10

8

54

55

127

Jones

3

3

16

12

34

Davies

2

3

44

30

79

Lutes

1

0

6

1

8

Total

29

24

204

193

450

Questionnaire letters were sent to two persons contacted on, and known to
have repeatedly hunted, the Bianco property.
In addition, Mr. Bianco kept
a rough running total of quail taken on his property by these men.
The

�-148-

questionnaire returns indicated only 11 quail were taken. However,
Mr. Bianco reported 70+ quail killed by these men. More than half, 47,
were banded.
This points out the failure of some hunters to accurately
report.
Returns in addition to those reported by Mr. Bianco indicate
that a least one-fourth of the quail banded on the Bianco property were
harvested.
Population Estimates--The banded-unbanded
ratios obtained from
harvest samples (Table 11) provide a general basis .for population estimates.
But, .any attempt to accurately determine populations on the banded
sites could not be completed for several reasons.
First, ingress and
egress of quail before and during the hunting season makes it difficult
to define the population.
Second, some hunters covered adjoining areas
during their hunt so that some of the quail were taken from other popUlations.
In other words, the sites were not adequate in size for making
accurate population or density calculations.
Available information indicates the 202 quail banded on the Bianco site
represented slightly over one-half of the total population.
The trapping area was approximately a quarter-mile wide by a mile in length.
The
unfarmed river bottom was less than a half mile in width.
Projections
indicate densities of a bird-per-acre or more may have existed in this
localized "hot spot."
Approximately a mile and one-half of river bottom was included in the
Propst trapping site. Projections from Table 11 again indicate a fall
density in excess of one quail-per-two acres on land which was grazed
in winter.
Data on the Jones and Davies sites were considered inadequate
for even proximal density estimates.
Studies on the upriver properties have been terminated for two reasons.
First, hunting pressure and harvest were higher than expected on the
Bianco and Propst sites.
Second, difficulties in obtaining accurate
harvest data and population estimates do not justify the time and effort
needed.

Identification

of Key Factors Limiting

Bobwhite

Quail

This phase of study primarily involved measurement of habitat characteristics with respect to their impact on bobwhite population levels.
Time limitations prevented collection of data quantifying habitat characteristics.
Numerous observations were made during other phases of study
concering possible habitat deficiencies.
Some of these observations are
briefly summarized here but more complete information will be forthcoming
in future reports.
Grazed

and Ungrazed

River Bottom

Trapping success on the Lutes property compared to other lightly to
ungrazed sites point
out the impact of sustained livestock grazing on
river bottom.
Grazing, especially in spring and summer, destroys much
of shrub and brush cover and suppresses forb production and the maturation of essential forb seeds. Nesting cover is also probably inadequate.

�-149-

When grazing is restricted to only winter use, as on the Propst site,
forbs and brush have a chance to recover during the growing season and
the impact on quail habitat is not nearly as critical.
The Braddock land, within the Tamarack complex, was lightly pastured
through the fall and winter and possibly yearlong.
Shrubs and forbs made
significant summer growth and provided fairly attractive fall food and
cover. A few quail were trapped on the site in early fall. Trap destruction by livestock prevented winter trapping, but continued use by livestock
through the winter opened the vegetation until it appeared unattractive as
quail cover. It is not known whether any coveys existed there or not in
winter.
Lack of shrubby cover along the South Fork of the Republican River
seemingly curtailed the potential of that drainage for bobwhite.
Whistling counts totaled only a fraction of those along the lower South
Platte.
Again, extensive grazing is practiced along the Republican.

Sex Ratio Comparisons
As previously cited in the section on Tamarack sex ratios,numerous
authors have considered the predominance of adult males in the population
as evidence of female loss during the reproductive period.
Robel (1965)
provided uncertain data concerning a higher than average ratio of hens
and speculated that abundant nesting cover was the reason why. Since
Tamarack data indicate hens were proportionally more abundant in the adult
population than expected, one might speculate that adequate nesting cover
was available and/or predation on nesting hens was not too important.

Visual Inspection

of Quail Utilization

of Habitat

Types

Late winter trapping on the Tamarack provided evidence
coveys favored locations which contained above average
resulting from early successional vegetative growth in
More stable sites containing dense continual. stands of
snowberry were not utilized as much.

that wintering
forb supplies
disturbed sites.
cordgrass or

Plotting of wintering covey locations on a map (Fig. 5) provided little
evidence of mass movements to locations adjacent to corn fields. However,
those coveys living next to corn fields frequently fed there. More data,
which may provide some quantitative support to these observations, will
be forthcoming in future reports.

�...150-

LITERATURE CITED

Davis, D. E. 1960. Estimating the numbers of game populations. pp. 5.15.27. In H. S. Mosby (Editor) Manual of Game Investigational Techniques.--The Wildlife Society, Washington, D. C.
Kabat, C., and D. R. Thompson. 1963.
dynamics and habitat management.
136 pp.

Wisconsin quail, 1934-1962 population
Wise. Conser. Dept. Tech. Bull. 30.

Klimstra, W. D. 1950. Bob-white quail nesting and production in southeastern Iowa. Iowa State ColI. J. Sci. 24(4):385-395.
Leopold, A.

1933.

Game Management.

C. Scribner's Sons, New York.

481 p ,

Petrides, G. A. 1954. Estimating the percentage kill in ringnecked
pheasants and other species. J. Wild1. Mgmt. 18(3): 294-297.
_____ , and R. Nestler. 1943. Age determination in juvenile bobwhite
quail. Amer. Mid. Nat., 30:773-782.
Robel, R. 1965. Differential winter mortality of bobwhites in Kansas.
J. Wildl. Mgmt. 29(2) :261-266.
Robson, D. S., and H. A. Regier. 1964. Sample size in Peterson markrecapture experiments. Trans. Amer. Fish. Society 93(3):215-226.
Rosene, W.
N. J.

1969. The Bobwhite Quail.
418 p.

Rutgers Univ. Press, New Brunswick,

Selleck, D. M., and C. M. Hart. 1957. Calculating the percentage kill
from sex and age ratios. Calif. Fish and Game 43(4):309-316.
Snedecor, G. W. 1956.
Ames. 534 p.

Statistical Methods.

The Iowa State ColI. Press,

Stoddard, H. L. .1931. The Bobwhi te Quail: Its Habi ts, Preservation
and Increase. C. Scribner's Sons, New York. 559 p. + 69 plates.
Taber, R. D. 1960. Criteria of sex and age. pp 6.1-6.60. In H~ S.
Mosby (Editor) Manual of Game Investigational Techniques. The
Wildlife Society, Washington, D. C.

Prepared by
Warren D. Snyder
~~
Wildlife Researcher

.

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                  <text>-71July,

JOB PROGRESS

State

Work

4

Job Title

Inventory

Covered:

Personnel:

Game Range

W-lOl-R-14

No.

Plan No.

Period

REPORT

COLORADO

of:

Project

1972

April

Roland

Investigations
la

Job No.
of Range Manipulation

1, 1971 to March

C. Kufeld,

Regional

Projects

in Colorado

31, 1972
Biologists.

ABSTRACT

An inventory of range-type conversion projects located on u.S. Forest Service
and Bureau of Land Management lands in Colorado was continued.
Since not all
data from projects scheduled for evaluation in 1971 have been received from the
Game, Fish and Parks Regions, presentation of 1971 data will be delayed until
the 15th segment report.

��-73-

INVENTORY

OF RANGE MANIPULATION

PROJECTS

IN COLORADO

Roland C. Kufeld

P. S. OBJECTIVE
To collect information which describes the location, extent, environmental conditions, land use practices, type of treatment and effects
of treatment on the range, livestock, and wildlife, for all completed
range manipulation projects in Colorado that are located within the
ranges of deer, elk and sage grouse, and to develop a processing
system for information which describes proposed range manipulation
projects.

SEGMENT OBJECTIVES
1.

To collect information which describes proposed and completed
range vegetation modification projects that are located in
Colorado and their effects on the range, wildlife and livestock.

2.

To compile,

3.

To provide desired IBM listings
agencies upon their request.

codify, process

METHODS

anq analyze

inventory

of inventory

data.

data to cooperating

AND MATERIALS

An inventory of range-type conversion projects located on U. S. Forest
Service and Bureau of Land Management lands in Colorado was continued
using procedures outlined in Colorado Division of Game, Fish and Parks
Administrative Directive No. 204, entitled "Range Vegetation Modification
Projects".
Procedures outlined in Administrative Directive No. 204 call for evaluations to be made on each vegetation modification project prior to completion and at the end of the 2nd, 5th and 10th years following treatment.
During 1971, 2 year post-treatment evaluations were scheduled to be
made on 31 vegetation modification projects completed in 1968.
Five requests for retrieval of range vegetation modification project
inventory data were processed during the 14th segment.
Requests were
made by the following: (1) Game, Fish and Parks Commissioner William
Robinson; (2) Colorado Coop. Wildlife Research Unit; (3) U. S. Bureau

�-74-

of Land Management, Colorado State Office; (4) Future Studies
Organization, Pioche, Nevada; and (5) U. S. Forest Service Rocky
Mountain Regional Office.

RESULTS AND DISCUSSION
Inventory data sheets for some type-conversion projects completed in
1971, and some scheduled for 2 year post-treatment evaluations in
1971, have not yet been received from the Game, Fish and Parks Regions
who are making the inventories and evaluations.
Therefore, presentation
of 1971 data will be delayed until the 15th segment report.

Prepared

by
Roland C. Kufeld
Wildlife Researcher

�-75July,

JOB PROGRESS

State of
Project
Work

REPORT

COLORADO
No.

W-lOl-R-14

Investigations

2

Job No.

Experimental Improvement of Oakbrush
on Deer and Elk Winter Ranges - Beaver Creek

Job Title
Covered:

Personnel:

Game Range

4

Plan No.

Period

1972

April

Roland

1, 1971 through March

C. Kufeld,

William

31, 1972

C. Osborne,

Jr., Martin

C. Brandt

ABSTRACT

Two year post-spraying
vegetation measurements were made on permanent pointquadrat plots established in 1968.
Analysis of data has been delayed until
literature reviews on forage value of oak study area plants for elk and deer
are completed, so benefits of vegetation changes caused by spraying can be
determined.
Forty-eight food habits studies were combined to determine what plants are
normally eaten by elk, and the relative value of these plants from a manager's
viewpoint based on the response elk have exhibited toward them.
Photographs showing vegetation conditions in the spray and control areas were
retaken between August 30 and September 1, 1971 at 67 permanent photo points.
All 115 permanent .001 acre pellet
groups on Septbember 1, 1971.

plots were

cleared

of deer and elk pellet

��-77-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH
ON DEER AND ELK WINTER RANGES - BEAVER CREEK
Roland C. Kufeld

P. S. OBJECTIVE
To determine if deer and elk carrying capacity can be increased on
oakbrush winter game ranges by spraying with 2,4,5-TP to induce
sprouting.

SEGMENT OBJECTIVES
1.

To measure percentage composition and cover of all plant species
on the study area following treatment to determine vegetation
changes that have occurred as a result of spraying with 2,4,5-TP
in 1969.

2.

To measure deer and elk use on the study area following treatment
to determine changes in game use that have occurred as a result
of spraying with 2,4,5-TP in 1969.

3.

To summarize and analyze pre-and
game use measurement data.

post-treatment

vegetation

and

METHODS AND MATERIALS
Vegetation measurements were made on permanent point-quadrat plots
established in 1968 (Fig. 1). Procedures for measuring vegetation
were exactly the same as those used in 1968, when pre-treatment
vegetation measurements were made (Kufeld 1970). Measurements began
August 6, 1971 and were completed August 26.
Photographs showing vegetation conditions in the spray and control
areas were retaken between August 30, and September 1, 1971, at 67
permanent photo points established in 1968.
All 115 permanent, .001 acre, circular pellet plots were cleared of
deer and elk pellet gro~ps on September 1, 1971. Accumulated pellet
groups will be counted in May, 1972, to determine game use changes
due to spraying.
An extensive review of literature was made to determine the forage
value of oak study area plants for elk. Forty-eight food habits
studies were combined to determine what plants are normally eaten
by elk, and the relative value of these plants from a manager's viewpoint based on the response elk have exhibited toward them.

�-78-

Fig. 1.

Measuring

vegetation

on a point quadrat plot.

�-79-

RESULTS AND DISCUSSION
A decision was made to delay the analysis of post-treatment vegetation measurement data until the literature review on forage value
of oak study area plants for elk and a review for deer are completed.
Thus, the vegetation measurement data are not presented herein.
Elk forage plants in the 48 food habits studies were categorized by
highly valuable, valuable and least valuable.
This information will
be used to determine the benefits of vegetation changes caused by
spraying.
Elk and deer forage value data will be entered on I.B.M.
cards with vegetation measurement data when pre- and post-treatment
vegetation data are analyzed.
The elk food habits paper has been
completed, and will soon be submitted for publication in a scientific
journal or as a Division of Game, Fish and Parks Special Report.

LITERATURE

CITED

Kufeld, Roland C. 1970. Experimental improvement of oakbrush on deer
and elk winter ranges - Beaver Creek.
Colo. Div. of Game, Fish
and Parks.
P-R Project W-10l-R-12, WP-4, J-2, Game Research Rept.
July. Pt. 1. pp. 95-112.

'j/, , ;.
';i,,-L(l,-,-cl
C
/t~'-,Lll tv,
Prepared by
'I
i'

.

•

'-, /

Roland C. Kufeld
Wildlife Researcher

",

��July,

-81-

JOB PROGRESS

State

of

Project
Work

REPORT

COLORADO
No.

W-lOl-R-14
4

Plan No.

Job Title
Period

1972

on Deer,

Covered:

Personnel:

April

Roland

Game Range

Investigations

3

Job No.

Experimental Improvement of Oakbrush
Elk and Cattle Ranges - Hightower Mountain
1, 1971 through March

31, 1972

C. Kufeld

ABSTRACT

Pre-treatment
game use measurements
showed an average of 19 deer and 14 elk
per square mile on the study area between September 3, 1970 and May 5, 1971.
Units 2 and 5 were sprayed by helicopter on July 12, 1971 with 2,4,S-TP at
2 pounds per acre mixed with 4 gallons of water and ~ gallon of No. 2 diesel
fuel per acre.
Firelanes were bulldozed during August around the portions of
units 1 and 7 which were to be burned.
These units were burned on October 7
and 8, 1971 by personnel of the U.S. Forest Service with assistance from Game,
Fish and Parks employes.
Units 3 and 8 were double anchor chained on November 16, 17 and 18, 1971.
All six treated units were seeded by helicopter in
November between the first and second chainings with a mixture of grasses,
forbs and shrubs at a rate of 18.9 pounds per acre.
Forty-eight food habits studies were combined to determine what plants are
normally eaten by elk and the relative value of these plants from a manager's
viewpoint based on the response elk have exhibited toward them.

��-83-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH
ON DEER, ELK AND CATTLE RANGES - HIGHTOWER MOUNTAIN
Roland C. Kufeld

P. S. OBJECTIVE
To determine the extent to which deer, elk and cattle forage production and game use can be increased and maintained by chaining,
spraying and controlled burning on overage Gambel oak winter game
ranges.

SEGMENT OBJECTIVES
1.

To make pre-treatment

game use measurements.

2.

To implement selected methods for improving
in the 60 acre habitat improvement units.

3.

To continue

a literature

deer and elk habitat

review.

METHODS AND MATERIALS
Pre-treatment

Deer and Elk Use Measurements

Deer and elk pellet groups were removed from all 480, 0.05 acre pellet
plots on September 3, 1970, and accumulated groups were counted May 5,
1971. In this study a pellet group is a definable group with more than
5 pellets.
Pre-treatment measurements of cattle use and forage production were reported by Kufeld (1971).

Implementation

of Habitat

Improvement

Measures

On July 12, 1971, 30 acre areas in the centers of habitat improvement
units number 2 and 5 were sprayed by helicopter with 2,4,5-TP herbicide
at a rate of 2 pounds per acre mixed with 4 gallons of water and 1/2
gallon of No.2 diesel fuel per acre (Fig. 1). A liquid detergent soap
was added at a rate of 0.5 percent of the total spray and carrier volume
to facilitate mixing of the chemical, water and diesel fuel. The remaining 30 acre areas within both units were left untreated for cover.
In August, 1971, firelanes were constructed around the inside 30 acre
perimeters of units 1 and 7. Lanes were about 25 feet wide and bulldozed
down to mineral soil (Fig. 2).

�-84-

Unit 7 was burned on October 7, 1971, and Unit 1 on October 8.
Fuels were not prepared before burning because of the additional
cost involved.
Vegetation was burned in its natural state (Fig.
3). Burning was supervised and conducted by personnel of the Grand
Mesa-Uncompahgre
National Forest.
Game, Fish and Parks employees
assisted with the burning.
Each area was burned in sections for
safety reasons.
Two pumper trucks and a bulldozer were on hand in
case the fire got out of control.
The first fire line was ignited on the downwind side of the unit and
allowed to burn to the bulldozed firelane.
The next fire line was
set upwind from the first, and this procedure repeated until the
entire unit was burned.
Ignition crews used propane brush burners
and fuses.
Diesel fuel pumped from hand garden sprayers was used to
increase the fire intensity.
On November 16, 17, and 18, 1971, the 30 acre centers of units 3 and
8 were anchor chained (Fig. 4). A 270 foot chain with 14 inch steel
crossbars welded to each link in the center 90 feet was used (Fig.
5). Each foot of chain equipped with crossbars weighed 70 pounds,
and the chain had a total weight of 16,570 pounds.
The center modified chain was attached to the two 90 foot, smooth leader chains with
swivels weighing 500 pounds each. The chain was pulled by two D-8,
46-A Caterpillar tractors.
The units were chained twice with the
second chaining in the opposite direction from the first. About 8
inches of snow was present when the chaining was done.
Between the first and second chainings all six 30 acre treated areas
were seeded by helicopter at a rate of 18.9 pounds per acre (Fig. 6).
Species used and seeding rates are presented in Table 1.
Units 4 and 6 were left untreated
sprayed, burned and chained.

for purposes

of comparison with those

An extensive review of literature was made to determine the forage value
of oak study area plants for elk. Forty-eight food habits studies were
combined to determine what plants are normally eaten by elk, and the
relative value of these plants from a manager's viewpoint based on the
response elk have exhibited toward them.

RESULTS AND DISCUSSION
Pre-treatment

Deer and Elk Use Measurements

Pre-treatment deer and elk use measurements are shown in Table 2. Deer
density ranged from 7 deer per square mile in unit 8 to 30 in unit 1 with
an average of 19 for all units. The relatively low deer density for the

�-85-

September 3, through May 5, period is because deer are absent from
the area during mid-winter.
The study area is primarily a springsummer-fall deer range.
Elk density ranged from 8 elk per square mile in unit 4 to 22 in
unit 3 with an average of 14 for all units.
The study area is
primarily a fall-winter-spring
elk range and normally receives a
higher degree of use than was recorded for this particular year.
Many elk which usually winter in the study area summer in the Muddy
Pass area to the southeast.
An early snowstorm can catch many of
the herd on the east side of the divide and force them into the Muddy
Creek drainage for the winter.
Aerial elk population trend counts
made annually by Colorado Division of Game, Fish and Parks Northwest
Regional personnel indicate this happened in the fall of 1970 (White
1971). The lower overall use should not affect pre- and post-treatment
elk use comparisons, however, because it is the relative degree of use
per unit that is of primary importance.

Implementation

of Habitat

Improvement

Measures

Spraying
Gambel oak in units 2 and 5 appeared to be about 80 percent top killed
5 weeks after spraying.
The kill appeared to be fairly uniform in
both units with the exception of one or two small spots that appeared
to have been missed.
The spray was kept within the prescribed boundaries
with very little if any adjacent drift damage.

Burning
Fall burning of Gambel oak can be done only after the leaves have turned
brown and begun to drop which happens about October 1. Unless the summer
has been particularly dry and fire danger very high a good burn cannot be
obtained in green vegetation.
A second factor which must be considered is the U.S. Forest Service
National Fire Danger Buildup Index, which is based on daily weather data
and reflects the ignitability of fuels. The fire danger buildup index on
the Grand Mesa-Uncompahgre National Forest is considered low from 0-30,
moderate 31-55, high 56-90, very high 91-140, and extreme over 140 (U. S.
Forest Service 1969). Burning of standing Gambel oak should only be
considered when the buildup index is over 40 and preferably between 60
and 75. Above 75 the danger becomes too high for safe prescribed burning.
Fall burning is thus confined to a relatively short period between October
1, and the first snowfall during which the buildup index must be at least
40 and no more than 75. Since these conditions do not always prevail there
are some years when fall burning is not possible.
U. S. Forest Service

�-86-

weather data from a site 4 miles northwest of the study area shows
the potential oak burning period which existed during each of the
last 6 years as follows: 1966 - 11 days; 1967 - 53 days; 1968 - 32
days; 1969 - no burning period; 1970 - 3 days; 1971 - 11 days.
On October 7 and 8, 1971, the days units 7 and 1 were burned, the
fire danger buildup index was 46 and 49 respectively.
Weather data
for those days are shown in Table 3.
Fire danger in 1971, was above the previous 6 year average during
June 20-30, July and August, but below average for September (Table
4). Thus, the vegetation was drier than average during the growing
season, but wetter weather in September probably made 1971 about
an average year for burning in October.
Neither unit burned uniformly.
Results varied from almost total
incineration of all vegetation with just a few dead snags left standing
(Fig. 7), to only partial removal of the understory with little or no
damage to the larger oaks (Fig. 8). On much of the area most of the
understory was removed with partial destruction of the overstory (Fig.
9). Figs. 7, 8 and 9 can be compared with Fig. 10, which shows typical
unburned vegetation adjacent to the burned area.
The extent to which vegetation was consumed by the fire varied with
wind velocity and brush density.
Strong wind gusts caused the fire
to crown and burn intensely resulting in a good burn. When the wind
subsided only the understory was consumed.
The heavier the brush
the more wind was required to penetrate it and fan the flames.
Thus,
some of the heaviest thickets were relatively undamaged.
Since weather conditions prior to and during the burning appeared to
be about average, it is felt that results are typical of the type of
burn which one can expect to obtain during the fall in the Gambel oak
type. For purposes of big game habitat improvement a non-uniform burn
such as this is probably better than a complete burn because patches
of cover are left intact.
Thus, fall prescribed burning can be used
to open large areas of Gambel oak habitat without destroying all of
the cover.

Anchor Chaining
Immediate effects of double chaining are shown in Fig. 11. Results of
chaining were uniform throughout beth units. Most large trees were
snapped off at the base while a few were uprooted.
Some smaller
serviceberry, chokecherry and oakbrush whips sprang back after the chain
passed, but chaining was very effective in opening the areas. A great
deal of slash remains (Fig. 12), but most of this should be concealed
from view within 2 or 3 years by new vegetation.

�-87-

Aerial Seeding
Several units were checked on the ground and by helicopter immediately
after seed application.
Seeds were visible on the snow and appeared
to be uniformly distributed.
Original project plans called for a survey to determine the rodent
population level, and implementation of a control program prior to
reseeding to reduce seed loss if a high rodent population was found
to exist. Nelson, Wilson and Goebel (1970) compared rodent depredations
on seeds on untreated plots and on plots where rodents were controlled
by broadcasting zinc phosphide treated wheat 1 week prior to seeding.
Within 6 weeks after fall seeding, rodents, particularly Peromyscus
maniculatus, had caused 98 percent depredation of all broadcast wheatgrass seeds in plots not protected by poison grain. Little or no
damage was observed on poisoned plots.
In mid-September, 1971, a trapline was established on the Hightower
Mountain study area through cooperation of the U. S. Bureau of Sport
Fisheries and Wildlife with a resulting catch of 30 rodents per 100
trap nights.
Eighty percent were deer mice (Peromyscus maniculatus)
and 20 percent were chipmunks (Eutamias sp.). According to the B.S.F.
&amp; W. biologist this population was high enough to warrant control of
rodents.
However, due to contemporary action by the Colorado Game,
Fish and Parks Commission prohibiting use of poisons to control predators where state funds are involved the rodent control work was not
permitted.

Literature

Review

Elk forage plants in the 48 food habits studies were categorized by
highly valuable, valuable and least valuable.
This information will
be used to determine the benefits of vegetation changes caused by
spraying, burning, chaining and reseeding.
The information will be
submitted for publication in a scientific journal or as a Division of
Game, Fish and Parks Special Report.

�-88-

LITERATURE

CITED

Kufeld, Roland C. 1971. Experimental improvement of oakbrush on
deer, elk and cattle ranges - Hightower Mountain.
Colo. Div.
of Game, Fish and Parks. P-R Project W-10l-R-13, WP-4, J-3,
Game Research Rept. July. Pt. 1. pp. 23-86.
Nelson, Jack R., A. M. Wilson, and Carl J. Goebel.
1970. Factors
influencing broadcast seeding in bunchgrass range. J. Range
Mgt. 23(3) :163-170.
U. S. Forest Service.
1969. National fire danger rating system
handbook.
F.S.H. 5109.11, Reg. 2. Supplement No.1.
June, 1969.
Amendment No.5.
White, Claude E. 1971. Principal Game Biologist, Northwest Region.
Colo. Div. Game, Fish and Parks, Grand Junction, Colo. Personal
communication.

I

Prepared by

J

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).
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Roland C. Kufeld
Wildlife Researcher

�Table 1. Species and seeding rates used to reseed units 1, 2, 3, 5, 7 and 8 on trre Hightower
oak study area on November 17 and 18, 1971.

Purity

Species

Grasses

Seed Specifications
Percent
Germination

Mountain

Pounds
Per Acre

Agropyron

cristatum

(Fairway strain)

95%

90%

2.22

Agropyron

intermedium

(Amur strain)

95

90

3.33

95

90

4.44

95

90

1.11

95

90

Bromus inermis

(Lincoln strain)

Dactylis

glomerata

(Potomac strain)

Festuca

ovina duriscula

Sub-total

1.11
12.21

I

co

Forbs

Shrubs

Meticago

sativa

(Ladak variety)

99%

49%

0.75

Meticago

sativa

(Nomad variety)

99

80

0.75

Meticago

sativa

(Rambler variety)

99

73

0.75

Melilotus

officinalis

99

85

Cercocarpus

montanus

Purshia

II

tridentata

Seed collected near Woodland

11

Seed collected near Maybell,

Sub-total

!I

11

Park, Colorado,
Colorado,

98%

85%

99

85

at an elevation

at an elevation

of 8,000 feet.

of 6,000 feet.

1.11
3.36
1.11

Sub-total

2.22
3.33

TOTAL

18.90

&lt;o
I

�-90-

Table 2.

. s . -1/
Pre-treatment deer and elk on 8 habitat improvement um.t

Animal

Unit
No.

Total
Pellet
Groups

Pellet
Groups
Per Ac.

Animal
Days Use
Per Ac .J) ,1/

1

18

150

11.5

30

2.7

2

10

83

6.4

17

4.7

3

15

125

9.6

25

3.2

4

8

67

5.2

14

5.7

5

9

75

5.8

15

5.3

6

15

125

9.6

25

3.2

7

9

75

5.8

15

5.3

8

4

33

2.5

7

11.4

Ave.

11.0

92

7.1

19

4.2

1

7

58

4.5

12

6.6

2

7

58

4.5

12

6.6

3

13

108

8.3

22

3.6

4

5

42

3.2

8

10.0

5

8

67

5.2

14

5.7

6

6

50

3.8

10

8.0

7

11

92

7.1

19

4.2

8

11

92

7.1

19

4.2

Ave.

8.5

71

5.5

14

5.7

~
~
~

Animal
Density
Per Sq. Mi.

Acres per
Animal Eer
Month _/

t=l

~
H

~

1/
Period of use was from 9-30-70 to 5-5-71.

u Defecation rate for deer and elk = 13 groups per day.
1/ Accumulation period = 244 days or 8.026 months.
4/ One month = 30.4 days.

�-91-

Table 3.

Weather data on days when units 7 and 1 were burned.
(Unit 7)
October 7, 1971

(Unit 1)
October 8, 1971

Temperature: Max.

670

680

Min.

270

280

Precipitation

0

0

Ave. Cloud Cover

0%

10%

Relative Humidity

25%

25%

Weather
Data

Wind Direction

N, then W, then SW

Wind Velocity

o to 15 mph

NW

0 to 10 mph

Table 4. Fire danger buildup index data for 1966-1971 inc. , at Curriers
Ranch, 4 miles northwest of the Hightower Mountain oak control study area.
Average Fire Danger Bui1duE Index
Sept.
Aug.
July

Summer

Year

June 20-30

1966

103

127

58

61

85

1967

28

33

60

52

46

1968

85

68

18

71

56

1969

13

42

74

35

47

1970

38

38

47

21

36

1971

89

96

61

35

67

Ave. All Years

59

67

53

46

56

�-92-

Fig. 1. On July 12, 1971, 30 acre areas in the centers of Units 2 and
5 were sprayed with 2,4,5-TP.

Fig. 2. In August, 1971, firelanes were constructed around the inside 30
acre perimeters of Units 1 and 7.

�-93-

Fig. 3.

Units

7 and 1 were burned on October

7 and 8, 1971.

Fig. 4. On November 16, 17 and 18, 1971, the 30 acre centers
Units 3 and 8 were double anchor chained.

of

�-94-

Fig. 5. The center 90 foot section of the chain used in Units 3 and
8. This section weighs 70 pounds per foot.

Fig. 6. In November, 1971, between the first and second chainings
all six 30 acre treated areas were seeded by helicopter with grasses,
forbs and shrubs at a rate of 18.9 pounds per acre.

�-95-

Fig. 7. Most effective burn obtained in burned units. Vegetation
was almost completely burned with just a few dead snags left standing.

Fig. 8. Least effective burn obtained in burned units. Only part of
the understory was removed with little or no change to larger oaks.

�-96-

Fig. 9. On much of the burned area most of the understory
removed with partial destruction of the overstory.

Fig. 10.

Typical

unburned

vegetation

adjacent

was

to the burned area.

�-97-

Fig. 11.

Edge of the chained area one day after double chaining.

Fig. 12. A great deal of slash remains in the chainings but most of
this should be concealed from view within 2 or 3 years by new vegetation.

��-99July,

JOB PROGRESS

State

of

Work

w-10l-R-14

Plan No.

5

Job Title
Period

REPORT

COLORADO
No.

Project

1972

April

Harold

Sheep Range

1, 1971 to March

Investigations

1

Job No.
Bighorn

Covered:

Personnel:

Game Range

Inventory

31, 1972

R. Shepherd, Daane A. Crook, James R. Hayward

ABSTRACT

Work preliminary to a range inventory of the Trickle Mountain and Buffalo Peaks
bighorn sheep study areas was performed during the period of this report.
This
included acquisition of color aerial photographs of the study areas, preparation
of field herbaria as plant identification aids, photography of plant species and
preparation of photo aids for plant identification,
the design and fabrication
of an improved Daubenmire canopy-coverage
frame, and the field testing of two
range-plant inventory devices.

��-101-

BIGHORN SHEEP RANGE INVENTORY
Harold R. Shepherd

P. S. OBJECTIVE
To inventory, contrast and evaluate certain range attributes
and an unproductive bighorn sheep habitat.

of a productive

SEGMENT OBJECTIVES
1.

Contract color aerial photography of the two study areas to a scale
of 1:12000 on Kodak Aero-Negative film, providing 60% lineal series
overlap and 30% overlap of adjacent flight lines.
Secure 3 sets of
9x9-inch color stereo-pair prints from above negatives and begin
delineating vegetative types on prints for the preparation of vegetative-type maps.
Delineate types on stereo-pair photographs with
mirror stereoscope as described in "Forester's Guide to Aerial Photo
Interpretation, Agricultural Handbook No. 308" and "The Manual of
Photogrammetry" of the American Society of Photogrammetry.

2.

Collect, identify, press and preserve plants representative of the
species composition of the vegetation found within the summer and
winter bighorn sheep ranges of the Trickle Mountain and Buffalo Peaks
areas.
Prepare herbarium mounts of plants collected, following
standard procedures described by H. D. Harrington and L. W. Durrell
(1957). Use Colorado State University herbarium facilities and the
assistance of its personnel in the identification of plants.

3.

Take in situ color photographs of range plants.
Obtain 2-l/2"x3-l/2"
color prints and assemble in several sets for the later use of field
crews in identifying plants encountered along transects, in typemapping, and in food habits investigation.

4.

Prepare several sets of plants mounted between plastic on 8-l/2"xll"
herbarium-grade paper and arrange systematically in loose-leaf notebooks as a part of the equipment to be used by student crews in transecting and type mapping and by other personnel in food habits and
nutrition studies.

5.

Arrange for the manufacture
(Daubenmire 1959).

6.

Test two types of transecting to learn which is best suited to the
vegetational and topographic features of the Trickle Mountain and
Buffalo Peaks areas and to the constraints imposed by the size of
the areas to be sampled and the objectives of the comprehensive
bighorn study.

of Daubenmire

canopy-coverage

frames

�-102-

(1)

The point-contact method
1967, Anderson 1971).

(2)

Daubenmire's

(Drew 1944, Fayle 1959, Loveless

canopy-coverage

method

(Daubenmire

1959, 1970).

RESULTS AND DISCUSSION

Color Aerial Photography

of Study Areas

The Trickle Mountain, Sheep Mountain and Buffalo Peaks areas were
vertically photographed on Kodak Color Aero-Negative film, at a scale
of 1:12000, by Kucera and Associates.
The photography was performed
August 17, and September 7 and 9, for the Trickle Mountain area, August
17 for the Sheep Mountain area, and September 7 for the Buffalo Peaks
area.
The photography resulted in three sets of 9x9-inch color and one
set of black and white stereograms.
rn addition to the aerial photography, many color photographs were taken
of selected vegetation types to serve as ground-truth information and
aids in the interpretation of aerial photo features and in the delineation
of vegetational types.

Preparation

of Field Herbaria

Two hundred and sixty-nine species of plants representative of those
occurring on summer and winter bighorn sheep ranges within the three
study areas were collected.
Nine specimens of each species collected
were pressed.
Identifications
to species were made in the field and
through use of Colorado State University Herbarium collections and
facilities.
The identifications were later checked by Dr. William M.
Klein and Dr. Richard G. Walter of the Botany and Plant Pathology
Department of Colorado State University.
One specimen of each species collected was donated to Dr. Richard M.
Hansen, Department of Range Science, Colorado State University and used
to prepare reference microscope slides used in bighorn sheep food habits
work.
A second specimen of each species was donated to Colorado State
University for inclusion in the University herbarium, and a standard
herbarium mount was prepared of a third specimen of each species for
inclusion in the Colorado Division of Game, Fish and Parks, Game Research
Center Range Project herbarium.
The remaining specimens were mounted on 8-l/2xll-inch, heavy, herbarium
paper and encased in laminating film. These were then used to prepare
five sets of field herbaria organized in loose-leaf binders according to
flower color, family, genus and species.
The herbaria will be used by
student transecting crews as plant identification aids in transecting
and vegetation type-mapping work and by other bighorn project personnel
concerned with food habits and nutrition studies.

�-103-

Preparation

of Photo Plant

Identification

Aids

Color photographs were made of most of the species of plants collected,
showing gross, vegetative and floral characteristics,
habitat and
associated species.
Originals '·'ere35mm Ektachrome transparencies.
From these, three sets of 2-l/2x3-l/2-inch
color prints were made and
labeled with the name of the plant and pertinent collection data.
These color prints were then inserted into transparent plastic holders
and assembled in three 8-l/2xll-inch
loose-leaf binders, organized
according to flower color, family, genus and species.
The books of
photographs will be used in conjunction with the field herbaria as
plant identification
aids.
The collection of transparencies will be
used, together with field herbaria and color prints, in intensive
plant identification
training for student field crews prior to and
during field work.
The transparencies will also be used in preparation
of slide talks describing the study.

Design

and Manufacture

of Canopy-Coverage

Frames

A Daubenmire canopy-coverage
frame, patterned after descriptions by the
originator (Daubenmire 1959) was made and field tested during the summer
of 1971.
Results of the field trials suggested several improvements.
Working drawings were prepared for an improved model, and two frames were
made.

Testing

Transecting

Methods

To learn which of two kinds of frames were best suited to range inventory of bighorn sheep habitat, field tests were made in three different
vegetation types, using the Daubenmire canopy-coverage
frame, and a
point-quadrat
frame (Loveless 1967).
Forty of each kind of plot were
established and read at seven-foot intervals along a 300-foot transect
in each vegetation type.
The vegetation types were alpine meadow,
Douglas fir-ponderosa pine, and pinyon-juniper.
Reading time for each
plot was recorded as well as canopy-coverage
data for each plant species
and percentages of bare ground, rock and litter.
In each vegetation type tested, the Daubenmire canopy-coverage
method
required the least time, less than half the time per plot required by
the point-quadrat method for alpine meadow and pinyon-juniper
types
(Table 1). An examination of data from the two methods indicated that
many more point-quadrat plots than Daubenmire canopy-coverage
plots
would be required for the same degree of accuracy.
A statistical test
was conducted to determine the number of Daubenmire canopy-coverage
plots
required per 300-foot transect in each of the three types transected to
estimate within 95 percent of the mean at the 95 percent confidence
level.
The mlnlmum number of plots indicated at the stated accuracy was
143 and the maximum 181, depending upon the species involved.

�-104-

Table 1. Average number of minutes
and read Daubenmire canopy-coverage
vegetation types.

Vegetation

Alpine
Douglas

Type

Point-Quadrat

meadow
fir-ponderosa

Pinyon-juniper

per plot required
and point-quadrat

pine

(A)

to establish
plots in three

Canopy-Coverage

(B)

A/B

7.9

3.6

2.2

4.8

2.5

1.9

7.6

3.5

2.2

The point-quadrat
frame weighed several times as much as the Daubenmire
canopy-coverage
frame, and was cumbersome to transport, and required two
men and a heavy hammer to set up.
Plots whic~ fell among rocks and trees
were sometimes impossible to establish at the exact spot desired because
supporting stakes struck solid rock.
In such instances the plot had
either to be shifted a few inches or feet along the transect line or
propped up with boulders or tree branches.
This was time consuming,
frustrating and conducive to subjective bias.
In contrast, the Daubenmire
canopy-coverage
frame was light and small enough to be easily carried
in a backpack, an important feature when working at high altitudes and
in steep, rocky terrain.
Taking into consideration
all of the advantages discussed, the Daubenrnire
canopy-coverage
frame is clearly the better of the two methods tested for
the bighorn sheep range inventory.

�-105-

LITERATURE

CITED

American Society of Photogrammetry.
The manual of photogrammetry.
Amer. Soc. Photogrammetry.
Menasha, Wis.
876 p.
Anderson, Allen E., Dean Medin, and David C. Bowden.
1971. Mule
deer fecal group counts related to site factors on winter range.
(Unpublished manuscript).
Daubenmire, R. 1959. A canopy-coverage method of vegetational
Northwest Science.
33(1):43-64.
analysis.

1970. Steppe vegetation
Sta. Tech. Bull. 62. 131 p.
Drew, W. B.
analysis

of Washington.

Wash. Agr. Exp.

1944. Studies on the point-quadrat method of botanical
of mixed pasture vegetation.
J. Agr. Res. 69:289-297.

Fayle, D. C. F. 1959. The point contact method as a three-dimensional
measure of ground vegetation.
The Forestry Chronicle.
35(2):

135-141.
Harrington, H. D., and L. W. Durrell.
Sage Books.
203 p.

1957.

How to identify plants.

Kuchler, A. W. 1955. A comprehensive method of mapping vegetation.
Annals Ass. of Amer. Geog.
65:404-415.
Loveless, Charles M. 1967. Ecological characteristics of a mule
deer winter range.
Colo. Game, Fish and Parks Dept., Tech. Pub.
No. 20. 124 p ,
U. S. Forest Service.
1969. Forester's guide to aerial photo
interpretation, Agr. Handbook No. 308. 40 p.

Prepared by

::-&lt;
. "", ")
~·'h""'.f.-(u:",'1; K.

I,'

fj.J-f.J;'-.

Harold R. Shepherd'
Wildlife Researcher

L,.",.i
fL.L"'·-\"

��-107-

JOB PROGRESS

State of
Project
Work

1972

REPORT

COLORADO
No.

W-10l-R-14

Plan No.

6

Job Title
Period

July,

Personnel:

LeRoy

April

Investigations
1

Job No.

Pilot

Covered:

Game Range

Plan for Game Management

1, 1971 to March

Carlson,

Bertram

Unit

Inventory

31, 1972

Baker, William

McKean

ABSTRACT

Deer and elk distribution and abundance data in map overlay form were obtained
from Allen Anderson and land-ownership
status information was collected from
Archuleta County Clerk records, respectively,
for application to Poudre River
and Pagosa Springs pilot areas computer mapping test operations.
Wildlife and allied data were compiled and tabulated for Wildlife Management
Units 61 (West Uncompahgre) and 62 (East Uncompahgre)
following selection of
those units for consideration
this segment.
Principal items of information
that were compiled for both units included areal extents, human population
characteristics
of corresponding
political entities, physiographic-climatic
descriptions,
land-ownership
and land use acreage summaries, wildlife species
checklists by game, nongame, bird and mammal categories, narrative and map
descriptions of game species distributions
and abundances, game harvests and
seasons, game species introductions records and management problems checklists.

��-109-

PILOT PLAN FOR GAME MANAGEMENT

UNIT INVENTORY

Bertram D. Baker and William

T. McKean

P. S. OBJECTIVE
To compile broad descriptive and wildlife
by wildlife management unit.

species management

information

SEGMENT OBJECTIVES
1.

Describe
physical

Wildlife Management Units by name, number, boundaries,
features, and climate.

2.

Inventory by unit the cultural and physical characteristics including
landownership status, land use status, and human population.

3.

Inventory by unit wildlife characteristics including species lists,
distribution and abundance, harvest data, introductions and/or past
species records, census areas and routes, and research and/or
management studies bibliographies.

4.

Inventory habitat (range) characteristics of vegetation types, vegetation condition and trend, restoration projects, and transect records,
all from existing sources.

5.

Inventory

6.

Test application of any item of information compiled or recorded in
steps 1.-5. above in pilot computerized map project now in operation.

present

and future management

problems

size,

by unit.

RESULTS AND DISCUSSION
Activities under this job were temporarily interrupted March 1, 1972, when
both authors were assigned duties on Project FW-lO-R~l.
Therefore, completion
of Unit 61 and 62 inventories will not be completed until late in 1972 or
early 1973 after work obligations have been met on the "Planning Project".
At that time a report, similar to the one compiled for Unit 22, will be
submitted.

Prepared

by

(
"
/,
/0'
j,'
(c
i&gt;."
I')
cd-}; V
Bertram D. Baker
Asst. Wildlife Researcher
U

,', -'-

��-111-

July, 1972

JOB FINAL REPORT
State of
Project No.

COLORADO

6

Work Plan Noo
Job Title:
Period Covered:

Deer-Elk

W-38-R-26

April,

Investigations

11
Job No.
Evaluation of the Effects of
Spring-SurrunerGrazing by Deer on Alfalfa

1968 through March,

1972

Personnel:
Richard M. Bartmann, Marion C. Coghill, John F. Corey, Kenneth C.
Dillinger, David A. Gordon, David F. Gordon, Robert Bo Gordon, James Ao Ives,
Julius J. Klein, Orville E. Lance, Ruel A. Loucks, Dale F. Reed, Jerry L.
Robinson, Donald Eo Speers, Steven Fo Steinert, Harold M. Swope, Fred C.
Vanderau, Collidge Woodard, Rifle Falls Hatchery personnel and Rifle Gap Conservation Unit inmates.

ABSTRACT
The two portions of the alfalfa damage study, one at Rifle Gap and the other
at the Little Hills Game Experiment Station, were terminated at the end of
Segment 26. Major reasons for termination at Rifle Gap include greater
levels of deer use than are found elsewhere in the state, unknown effects of
moisture levels on plant responses when interacted with grazing, the limited
need for the information and legal questions concerning applicability of results
to other areas. Reasons for discontinuing the Little Hills portion include
too little deer use on the unfenced field, the unnatural reactions of confined
deer on the fenced field during spring and the same last three reasons mentioned
above for Rifle Gap. At Rifle Gap, significant (p&lt;.Ol) reductions in hay
yields of 1216, 1109 and 835 pounds per acre were recorded for the first cutting
in 1970 and 1971 and the second cutting in 1971, respectively.
Total deer-nights
use corresponding to these losses were 4560, 5224 and 1669
Deer densities during
an earlier study of spring damage at Billy Creek were 2.7 deer per acre in both
1958 and 1959. The highest density recorded on a spring damage claim was 2.6
deer per acre. In both cases, deer were present for only about a one month period
during spring. Densities for a comparable period at Rifle Gap were 3.6 and 4.0
deer per acre. The loss of air-dry forage on a deer-night basis was estimated
at 6.6 and 5.2 pounds for the first cuttings and 14.0 and 12.4 pounds for the
second cuttings.
No significant (p) .05) differences were found in percentages
of protein, fat, fiber, ash and nitrogen free extract between grazed and ungrazed
hay samples taken in 1970
0

0

�-112-

RECOMMENDATIONS

1.

Discontinue all research activities related to investigation
damage to growing hay crops.

2.

When necessary, continue to settle claims of deer damage to growing hay
crops on the basis of three pounds of air-dry hay per deer-night.

3.

Continue efforts to gain more favorable revisions to the wildlife damage
law and, ultimately, to gain repeal of the State's liability for damages
to private property incurred by wildlife.

of deer

�-113-

EVALUATION OF THE EFFECTS OF
SPRING-SUMMER GRAZING BY DEER ON ALFALFA
Richard M. Bartmann

This report covers only the current phase of the alfalfa damage study
which began in April, 1968. The initial effort under Work Plan 6, Job
11, occurred from April, 1961, through March, 1965, at Rifle Gap, and
was an attempt at assessing the effects of summer-long grazing by deer
on alfalfa production (Boyd 1963; Boyd 1964; Boyd 1965a; and Boyd 1965b).
Numerous problems were encountered during the four-year study duration
and only token data were collected for the first cutting of the final
year. Results from five fenced and five unfenced plots, each 30 by 200
feet, indicated slightly more hay produced on areas grazed by deer than
on ungrazed areas.
Information on deer use was limited to five "spot
checks" by the local Wildlife Conservation Officer during the month
immediately preceding hay harvest.
A second attempt at evaluating the effects of deer grazing on alfalfa
production was made from April, 1965, through March, 1968, at the Little
Hills Game Experiment Station (Boyd 1966; Gordon 1967; and Gordon 1968).
A series of paired plots, each 9.6-square feet in area, were located on
two different hayfields; one was to be irrigated and the other unirrigated.
Results from the final year indicated nearly equal production on grazed
and ungrazed plots. Deer counts showed negligible use on one field and
light use on the other, and then only for about a one month period.
Numerous uncontrolled variables were cited as responsible for the lack of valid
results.
The third phase of the alfalfa damage study was a final attempt to answer
questions concerning the effects of deer grazing on alfalfa production
(Gordon 1969; Gordon 1970; and Gordon and Bartmann 1971). The study was
conducted on West Rifle Creek and data were collected for two seasons.
The study was terminated at the end of Segment 26 and the following is a
final report of this effort.

P. S. OBJECTIVE
To determine
alfalfa.

the effect of spring-summer

DESCRIPTION

grazing by deer on production

of

OF AREA

In 1968, the Division of Game, Fish and Parks purchased the Charles W.
Clark ranch on West Rifle Creek, about 10 miles north of Rifle, Garfield
County, for development as part of the Rifle Gap State Recreation Area.
This property had a history of spring and summer deer grazing on alfalfa
fields and, since it was now Division property, was considered an appropriate
study site.

�-114-

The Rifle Gap study field is located along West Rifle Creek (Fig. 1).
It contains 25.7 acres II and is oriented in a general east-west direction (Fig. 2). The field slopes downward from west to east at about
a four percent grade. A road borders the north side behind which are
steep hills with pinyon (Pinus edulis)-juniper
(Juniperus osteosperma)
overstory.
West Rifle Creek borders the south side of the field immediately
beyond which are steep hills with pinyon-juniper overstory interspersed
with open sagebrush (Artemisia tridentata) flats.

METHODS AND MATERIALS

Study Field Preparation
During April, 1968, soil samples were taken from the study field and sent
to the Soil Testing Laboratory at Colorado State University to determine
fertilizer requirements.
The following May, about 350 pounds per acre of
45 percent phosphate fertilizer were spread on about a 20-acre portion of
the field.
These 20 acres were then plowed, disced, and seeded with Ladak
alfalfa at 12 pounds per acre and a nurse crop of Colorado 37 oats at 24
pounds per acre by a private contractor.
A poor seeding job by the contractor, problems with the irrigation
and little rain until late summer created a poor stand of alfalfa.
bare areas were reseeded with a hand-operated seeder the following
but cheatgrass (Bromus tectorum) was still abundant.

system,
The
spring,
,
,

The field had originally been flood irrigated, but it was decided that'
better water distribution could be accomplished by sprinkler irrigation.
A sprinkling system was set up with water being pumped out of the original
main irrigation ditch.
Irrigation, once started, was continued around the clock until completed.
Sprinkler lines were moved every 12 hours to get the newly seeded field
started, and every 24 hours thereafter.
The field could be irrigated once
in eight days when lines we re moved every 24 hours.
Precipitation from April through August, as measured at the nearest weather
station in Rifle, was slightly below normal in 1968, slightly above normal
in 1969 and 1970, and about 25 percent below normal in 1971 (U. S. Department of Commerce 1968, 1969, 1970, 1971). The norm for this period was
4.69 inches.

Study Design
A plot size of 12 by 200 feet was selected to sample alfalfa yields on
grazed and ungrazed areas. The width conformed to a standard l2-foot

l/previous reports listed the field size at 16 acres. Recent calculations
show the total acreage to be 25. 7 acres of which nearly 20 acres were reseeded for the study. Accordingly, all data in this final report are considered in relation to the revised acreage figure.

�-115-

cutterbar on a self-propelled swather. The 200-foot length was convenient to the width of the field and would, hopefully, provide a
sufficient sample of hay to minimize variability.
Accordingly, the corners of 42 possible 15 by 200 foot plots, spaced 30
feet apart were marked with steel fence posts. The IS-foot width allowed
room for the swather and baler to travel through the plots after they were
fenced (Figs. 3 and 4). The plots were aligned across the short length of
the field to facilitate moving irrigation pipe.
Hay production on all plots was measured at both first and second cuttings
in 1969. The hay was first cut the long length of the field across the
ends of the plots.
Care was taken to cut as close to each plot as possible.
Hay on the plots was harvested by cutting a swath down the center.
The
hay adjacent to each plot was then cut with the corner markers being removed and replaced as the swather approached and passed.
Baling was accomplished in the same pattern as cutting.
The baler was
tripped before entering the first plot and again after the last hay from
each plot was picked up. The bales were marked as they emerged from the
baler.
As soon as baling was complete on the plots, the bales were weighed
to the nearest pound on a chatillion spring scale (Fig. 5).
After the plots were fenced, the swather left a small strip of hay along
each side of the fences. This residual hay was mowed with a small garden
tractor fitted with a four-foot cutterbar attachment.
During the first
cutting this loose hay was hauled off the field. At second cutting it was
sparse enough to leave on the field.

Sample Plot Selection

and Fencing

Comparison of hay yields on grazed and ungrazed areas was by paired plots.
Plot pairing was on a production basis using the 1969 second cutting data
(Table 1). There were fewer weeds in the second crop and more pairs could
be selected.
For purposes of plot palrlng, the field was stratified into three sections;
northwest, northeast, and southwest.
Only plots within each strata were
paired.
A five-pound difference was chosen as the maximum allowable between
plot pairs. Whenever three or more plots fell within this limit, the two
closest to each other in the field were used. Fourteen pairs of plots were
selected with a coin toss determining which one would be protected and which
one unprotected from deer use.
Plot fencing was completed in September, 1969 (Fig. 6). Wooden posts were
set at each corner and a 10-foot steel post driven every 10 feet between
corners.
Two courses of 39-inch woven wire were strung on the posts to produce a 7-l/2-foot high fence. Gates for the plot ends were made of two
courses of woven wire fastened to steel posts. The gates were closed at the
beginning of the growing season, about April 1, and left open after the
second cutting each year.

�-116R93W

Study Field

~
10

~I/I'

II

HOGBACK

U.S. 6-24

Fig.

1.

Vicinity

map of Ri fle

Gap alfalfa

damage

study

field.

�N

Countinc;»

~

~t~tl(H'I~~t!If!£J!P

\~--

__ -------

Road

;-.--=-"'7..,

•••••••

•.....•~
,

",
\

,

I

\

"

'\
)
(

Reseeded area

\

Freid boundary

\

\
\
\

\.

"' .•..

Fenced plot
Unfenced plot

Fig. 2.

I

Layout of the Rifle Gap study field and paired plot locations.

(

,

,
)

I

~

~
-...J
I

�I

t-'
t-'
00
I

Fig. 3.

Swathing hay on a fenced plot on the Rifle Gap alfalfa field.

R. M. Bartmann photo.

�I

I-'
I-'
-0
I

Fig. 4.

Baling hay on a fenced plot on the Rifle Gap alfalfa field.

R. M. Bartmann photo.

�I

I-'
N

o
I

Fig. 5. Weighing baled hay to measure production on plots grazed and ungrazed by deer on the
Rifle Gap alfalfa field. R. M~ Bartmann photo.

�-121-

Table l. Hay yields on potential sample plots on the Rifle Gap alfalfa damage
study field, 1969, and plot pairs selected from second cutting data.

First Cutting
Plot No.
Yield (lbs)

Second Cutting
Yield (lbs)
Plot No.
Northwest

12
13
10
14
6
9
11
7
2
8
5
4
3
1

Northeast
28
33
16
19
17
15
22
23
18
21
32
29
20
34
31
27
30
26
24
25

294
288
280
278
276
275
251
143

176
172
158
154
145
140
128
127
102
86
70
68
65
40

3 Fenced
3 Unfenced
4 Unfenced
4 Fenced
5 Unfenced
5 Fenced
2 Fenced
2 Unfenced

1 Fenced
1 Unfenced

Section of Field
15
16
24
19
29
28
32
27
17
34
23
31
18
25
22
33
26
30
21
20

321
312
292
286
280
270
264
258
256
242
237
236
232
224
220
194
173
169
158
152
Southwest

40
38
39
41
37
42
36
35

Section of Field
8
12
10
13
6
14
7
11
9
5
2
1
3
4

346
326
318
312
294
294
290
289
286
283
278
266
262
250

Plot Pairs

142
142
134
118
115
114
108
107
104
102
98
98
96
85
84
84
76
76
71
No weight

6 Unfenced
6 Fenced

9 Unfenced
9 Fenced
11 Unfenced
11 Fenced
10 Unfenced
10 Fenced

7 Fenced
7 Unfenced
8 Fenced
8 Unfenced

Section of Field
36
41
37
38
40
35
42
39

179
168
164
164
164
162
160
141

13 Fenced
12 Fenced
13 Unfenced
12 Unfenced
14 Unfenced
14 Fenced

�I

i-'
N

N
I

Fig. 6.

Plots fenced to exclude deer grazing on the Rifle Gap alfalfa field.

Harold M. Swope photo.

�-123-

Deer Counts
Deer on the field were counted during the growing season to enable comparison of deer use with differences in hay yields on paired plots.
A
counting station was located across the road along the north side of the
field about one-third the distance from the west end. The observer arrived
at the counting station about one-quarter to one-half hour prior to the
first count and remained there until the last count was made.
Counting was accomplished with a hand-held spotlight and 7-power binoculars.
With few exceptions, the deer were not spooked by the observers presence
or the light. Deer were counted on the entire 25.7-acre field as it was
impossible to differentiate the old from the newly seeded area at night.
In 1970, weekly counts were made beginning about the first of April and
ending at second cutting about the first of September.
A count was made
every hour from 7 P.M. to 5 A.M., MST.
Weekly counts were again made in 1971, but only three hourly counts were
made per night from 9 P.M. to 11 P.M., MST. This reduction in the number
of hourly counts was promoted by analysis of previous years' data (Gordon
and Bartmann 1971).

Proximate

Analysis

There was some concern over the effects of grazing on the nutritional
quality of alfalfa.
An attempt at approaching this matter was made in
1970. The last bale from each plot was labeled with plot number, treatment,
and cutting.
After second cutting, all bales were taken to the Research
Center in Fort Collins for proximate analysis.

RESULTS

Hay Harvest
Hay harvest data for 1970 and 1971 are shown in Tables 2 and 3. Grazed
plots averaged less hay than protected plots all four times. These differences were significant (p &lt;.01) for the first cutting both years and for
the second cutting in 1971, while the difference for the second cutting in
1970 was not (p&gt; .05). In most cases, alfalfa within the fenced plots was
noticeably taller than that outside (Fig. 7).
The plot yield data were projected
acre basis. These estimated are:

to estimate production

1970

losses on a per

1971

First cutting

1216 pounds

1109 pounds

Second cutting

581 pounds

835 pounds

�-124-

Table 2. Hay yields on grazed and ungrazed
alfalfa damage study field, 1970.

Plot
Pair

Yield
Grazed

plots at the Rifle Gap

(Lbs)
Ungrazed

Difference

First Cutting
1
2
3
4
5
6
7
8
9
10
11
12
13
14

236
308
332
323
298
312
319
219
335
320
235
290
257
212

294
319
272
454
308
394
356
304
392
360
412
388
307
374

58

- 11
+ 60
- 131
10
82
37
85
57
- 40
- 177
- 98
50
- 162

x difference

- 67 pounds

Second Cutting
1
2
3
4
5
6
7
8
9
10
11
12
13
14

51
208
192
100
84
88
58
74
112
92
72_
122
62
30

63
117
118
216
164
101
123
92
138
120
34
123
134
149

12

+ 91
+ 74
- 116
80
- 13
65
18
26
28
58
1
72
- 119

x difference

32 pounds

�-125-

Table 3. Hay yields on grazed and ungrazed plots at the Rifle Gap
alfalfa damage study field, 1971.

Plot
Pair

Yield (Lbs)
Grazed

Difference

Ungrazed

First Cutting
1
2
3
4
5
6
7
8
9
10
11
12
l3
14

224
344
287
290
233
246
248
263
268
344
359
235
227
203

- 148
- 105
40
88
92
66
+ 20
56
72
+ 57
- 42
31
39
- 154

312
324
343
356
325
318
402
302
299
287
301
275
332
351

61 pounds

x difference

Second Cutting
1
2
3
4
5
6
7
8
9
10
11
12
13
14

17
112
110
95
30
114
47
68
82
129
79
43
29
28

30
3
23
80
- 117
- 45
- 104
- 90
- 25
+ 65
- 10
82
63
- 43

47
109
133
175
147
159
151
158
107
64
89
125
92
71

+

x difference

-

46 pounds

�•...
I

N

0'
I

Fig. 7. Differences between grazed and ungrazed alfalfa were usually noticeable to varying degrees.
The above photo represents an extreme contrast immediately prior to the second harvest.
The poor
appearance of the alfalfa is a result of not being irrigated during the second growth period.
R. M. Bartmann photo.

�-127-

Deer Counts
The highest count for the hours of 9 P.M. to 11 P.M., MST, are used to
calculate deer-nights use on the study field (Fig. 8). The highest
number of deer seen on the field, 132, was in the spring of 1971. Both
years, deer numbers declined sharply beginning the third week in Mayas
the migrant deer left.
The estimated 5224 total deer-nights of grazing during the first cutting
in 1971 was about 15 percent more than the estimated 4560 deer-nights in
1970. This difference would have been greater had not there been a 15day longer period of deer exclusion (plot gates closed) in 1970; 102 days
in 1970 and 87 days in 1971.
Deer use of the second crop also was greater in 1971. There were 1669
deer-nights use estimated that year compared to 1021 deer-nights in 1970.
The period of deer exclusion was slightly longer the second year; 44 days
in 1970 and 50 days in 1971.
Mean deer densities on the entire 24.7-acre unfenced portion of the field
were estimated for both cuttings both years using the maximum hourly counts
from each week.
The deer per acre estimates are:
1970

1971

First cutting

1.8

2.4

Second cutting

0.9

1.4

The pattern of deer distribution on the study field in previous years was
observed to be non-uniform (Gordon 1970; and Gordon and Bartmann 1971).
An area around the counting station had noticeably fewer deer than the
rest of the field.
The same situation was noted again during counts and
also during several "spot checks" on other nights in 1971.
Inspection of hay weight data does not reveal any gross irregularities in
paired plot yields from that portion of the field around the counting station
and those from the remainder of the field.
It is probable that deer numbers
increased on this area later in the night or on nights when the observer was
not present.
Alternatively, hay yields may be reduced by lesser numbers of
deer, or changes in productivity between plot pairs may have occurred naturally.

Proximate

Analysis

Results of the proximate analyses performed on hay samples from the first
and second cuttings in 1970 are summarized in Table 4. None of the differences between grazed and ungrazed plots were significant (p&gt;. 05) for any
of the factors tested.

�-128-

140
130
120
110
100

~'1

90

0::

I

\

,"

,

,

II

I

\\
\

,

\

I

\

~ 80

,

I.L

o 70

,

,

~ 60

,

::::&gt;

\ I

,

40
30
20
10

I

,,, \,,I I
, "

0::

W

50

1971

\

W

z

1970

,I

,,,
,

V

,,,

,,,
,,,
,,

..

~- -..

I

'"I

I

,,

2 345
APR

1 234

.1~

MAY

1 234

.1~

JUN

MONTH

I 2345123412

.1~
a

JUL

--I ~

AUG-+SEP

WEEK

Fig. 8. The high number of deer counted per night during weekly counts on
the Rifle Gap alfalfa damage study field, 1970 pno 1971.

�-129-

Table 4. Results of proximate analyses of hay samples from the Rifle
Gap alfalfa damage study field, 1970. (n = 14).
First Cutting
Grazed
Ungrazed

Calc. J:./

Second Cutting
Grazed
Ungrazed

Calc. J:./

(%)

(%)

"t" .05
13 df

0.608

14.81

15.89

0.553

2.49

0.285

3.92

3.66

1.386

49.92

51.19

0.619

41.90

42.10

0.073

Ash

6.09

6.09

0.007

7.27

8.07

1.348

Nitrogen
Free Extract

33.16

32.01

0.717

32.09

30.28

0.725

Analysis

(%)

(%)

"t" .05
13 df

Protein

8.25

7.94

Fat

2.57

Fiber

))

Tabular

"t" .05, 13 df

2.160.

DISCUSSION
The relationship between deer numbers and loss in hay production for the
first cutting was inverse.
Estimated hay Los'swas 9 percent less in 1971,
while estimated deer use increased about 15 percent.
For the second cutting,
hay loss and deer use increased 44 and 63 percent, respectively.
Estimates of forage loss on a deer-night basis were calculated from the
Rifle Gap data to compare with the three pounds per deer-night figure used
in settling claims of deer damage to growing hay. These estimates are:

1970

1971

First cutting

6.6 pounds

5.2 pounds

Second cutting

14.0 pounds

12.4 pounds

The above values are not indicative of forage consumption by deer. Rather,
they represent the response of alfalfa to deer grazing interacted with
growing conditions.
An evaluation of results to date, of problems involved in continuing the
study, and of benefits to be derived was made after conclusion of field work
and data analysis in Segment 26. It was decided that little could be gained
by continuing and the study wa s terminated.
Some of the considerations influencing this decision are discussed below.
Another portion of the study,
to be done at the Little Hills Game Experiment Station, was also terminated.
A brief summary of this work is included at the end.

�-130-

The results from the Rifle Gap field are the first since damage studies
were begun in Colorado to show reduced hay production on deer grazed areas.
One possible explanation for this relates to intensity and seasonal duration
of deer use. Estimated deer densities during the study at Billy Creek
(calculated from raw data) were 2.7 deer per acre in both 1958 and 1959.
However, this was only for a one month period from mid-April to mid-May.
Densities at Rifle Gap for the same period were 3.6 and 4.0 deer per acre
in 1970 and 1971, respectively.
Calculated densities based on acreage and
deer count figures given in claims submitted for spring damage to alfalfa
range from 0.1 to 2.6 deer per acre. About 75 percent figured less than
1.0 deer per acre and nearly all were for only a one to two month period
in early spring prior to deer migration.
No comparison of deer densities for the second cutting can be made as only
incomplete data from a previous study are available for this period.
A few
damage claims were submitted, but nearly all were for irrigated hay. Calculated densities on these areas ranged from 0.04 to 1.0 deer per acre,
except for one eight-acre field which had 4.4 deer per acre. Dates of
damage varied from August through October.
There were 0.9 and 1.4 deer
per acre on the unirrigated second hay crop at Rifle Gap during 1970 and
1971, respectively.
The s~asonal duration of deer use at Rifle Gap differed somewhat from that
at B~ly Creek.
The pattern of deer use was similar through mid-May, but
at Ri~le Gap, a resident herd maintained relatively light but constant
grazing pressure through harvest.
How much this contributed to the recorded loss in hay production is not known.
The method of irrigation could play an important role in hay response to
deer grazing.
The efficiency of sprinkler irrigation at Rifle Gap was
probably less than with the flood irrigation at Billy Creek. According to
water out-put tables for the sprinkler system, 12 inches of water should
have been put on the field with two irrigations; whereas only 6 inches were
actually measured.
Evaporation, particularly during the hotter daylight
hours, was probably one reason for this difference.
Other factors that could be equally or more important than the ones already
mentioned above relate to edaphic and climatic conditions, and alfalfa
variety.
The effects of deer grazing intensity on alfalfa production still remain
largely unanswered.
It has been shown at Rifle Gap that, given enough
deer, production can be affected.
However, deer densities there were considerably above those encountered on any previous study field or on areas
where damage claims have been paid. Because of restrictive hunting seasons
in 1971, there is a good chance of even higher densities at Rifle Gap in
1972. Thus, the applicability of the results can become even farther removed from other situations.
Another consideration in terminating the study is the limited need for information on deer grazing-alfalfa production relationships.
The Billy Creek
data, since completion of that study in 1959, have been used twice in arbitration proceedings (Boyd 1972). The rancher won the settlement both times.

�-131-

In neither case was the validity of the study design or the results
questioned.
The main point of contention was that the study was done
in an area removed from that of the claimant's.
Additional studies
would not remedy this situation as each rancher could still consider
his field a special case.
The scope of the problem also appears quite limited.
Payments for deer
damage to growing alfalfa from 1963 through 1970 have averaged about
$2900 per year. However, fieldmen's time investigating claims, and processing of claims would involve considerable additional expense.
Twothirds of the total amount of claims paid for alfalfa damage in the past
eight years has been to six individuals and three-fourths of the amount
has been paid in four small geographical areas.
In several of these
cases, recommendations for payment were partly based on public relations
reasons; a situation that research could not ,tackle anyway.
The 1971 revision of the wildlife damage law by the Colorado Legislature
restricted conditions under which damage to private property by wildlife
can be claimed.
The law has not been in effect sufficient time to fully
evaluate its effectiveness, but in principle the people of the State of
Colorado should benefit.
However, indications are that problems may arise
because of some undefined terminology in the law.

Little Hills
It was thought desirable to conduct the alfalfa damage study at two
locations.
The second area selected was the Little Hills Game Experiment
Station about IS-air miles west of Meeker.
It was also thought desirable
to conduct one part of the study using known numbers of confined deer.
Again, Little Hills proved suitable because of a series of already existing
deer-tight pastures ranging from 76 to 20S acres that could be used to hold
deer. Preparations were made for this study but it was terminated before
any results were obtained.
For the record, a brief synopsis of progress
up to termination, and reasons for termination are presented.
In 1968 and 1969, an abandoned 60-acre field was plowed, fertilized,
disced, and seeded the same as at Rifle Gap. A well was drilled to provide
water for a sprinkler irrigation system that was set up. The alfalfa was
established and an II-acre area containing 8 acres of alfalfa was fenced
for the controlled phase of the study. Deer were trapped from the
surrounding area, placed in a 90-acre pasture connected to the field, and
allowed free access to the field all year.
At this point, the decision was made to discontinue the study. Major
reasons for this decision relate to problems of getting sufficient deer
use on the field open to natural deer use, and of getting some semblance
of "natural" deer use on the fenced field during spring.

�-132-

The lower, or west, one-half of the field was open to use by freeranging deer.
Twenty-one deer were the most ever seen on the 30-acre
field at one time and they were gone by the fourth week in April, nearly
a month earlier than occurs in most other areas. Also, deer use was
limited to only one small portion of the field.
Casual observations of deer on the fenced field were made in 1970, and
several "spot checks" were made during spring and summer in 1971. All
of the confined deer came into the fenced area during spring, but they
tended to walk around the fences in a large group and few were actually
seen on the alfalfa field.
From about mid-May until early July, deer
were rarely seen in the fenced area. After this time they were observed
on the field regularly through the rest of the summer.
Other reasons for discontinuing the study at Little Hills are the expected
low precision of the data due to the expected high variability and the
small number of plots we could reasonably establish, the problems with
application of the results to other situations, and the need for the time
and money for more important research projects.

LITERATURE

CITED

Boyd, R. J. 1960. An evaluation of spring grazing on alfalfa by deer
in western Colorado.
Proc. Western Assoc. Game and Fish Commissioners.
40: l30-l47.
1963. Methods of evaluating deer and elk use of alfalfa under
summer-long use. P. 39-42.
In Game Research Report.
Colorado Dept.
Game and Fish, Denver.
1:1-115.
1964. Methods of evaluating deer and elk use on alfalfa under
summer-long grazing.
P. 42-46.
In Game Research Report.
Colorado
Dept. Game, Fish and Parks, Denver.
2(Part 1):1-69.
1965a. Methods of evaluating deer and elk use on alfalfa under
summer-long grazing.
P. 69-73.
In Game Research Report.
Colorado
Dept. Game, Fish and Parks, Denver.
3(Part 1):1-126.
1965b. Methods of evaluating deer and elk use on alfalfa under
summer-long grazing.
P. 309-332.
In Game Research Report.
Colorado
Dept. Game, Fish and Parks, Denver.--3(Part
111):251-410.
1966. Evaluation of deer use on alfalfa under different irrigation
rates. P. 201-206.
In Game Research Report.
Colorado Dept. Game,
Fish and Parks, Denver.
3(Part 11):95-343.
1972.

Personal

communication.

�-133-

Gordon, D. F. 1967. Evaluation of deer use on alfalfa under different
irrigation rates. P. 175-178.
In Game Research Report.
Colorado
Dept. Game, Fish and Parks, Denver.
3(Part 11):73-310.
1968. Evaluation of deer use on alfalfa under different irrigation
rates. P. 309-322.
In Game Research Report.
Colorado Div. Game,
Fish and Parks, Denver.
3(Part 111)209-421.
1969. Evaluation of the effects of spring-summer grazing by
deer on alfalfa.
P. 25-43. In Game Research Report.
Colorado
Div. Game, Fish and Parks, Denver.
3(Part 1):1-140.
1970. Evaluation of the effects of spring-sun®er grazing by
deer on alfalfa.
P. 217-249.
In Game Research Report.
Colorado
Div. Game, Fish and Parks, Denver.
3(Part 11):127-286.
______ , and R. M. Bartmann.
1971. Evaluation of the effects of springP. 97-112.
In Game Research
summer grazing by deer on alfalfa.
3(Part II):
Report.
Colorado Div. Game, Fish and Parks, Denver.
87-224.
U. S. Department of Commerce, Environmental
1968. Climatological data, Colorado.
1969.
219-231.

Climatological

Science Services Administration.
Annual summary.
73(13):211-222.

data, Colorado.

Annual

summary.

74(13):

U. S. Department of Commerce, National Oceanic and Atmospheric Administration.
1970. Climatological data, Colorado.
Annual summary.
75(13):217-229.
1971.
213-224.

Prepared

Climatological

data, Colorado.

by
Richard M. Bartmafin
Asst. Wildlife Researcher

Annual

summary.

76(13):

��-135-

July, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project NOD

W-38-R-26

Work Plan No.
Job Title

Investigations
1

Job No.

White River Elk Experimental

Period Covered:
Personnel:

11 C

Deer-Elk

Management

Study

April 1, 1971 through March 31, 1972

Richard M. Bartmann, Raymond J. Boyd, William
G. Smith and Steven F. Steinert.

H. Rutherford,

Donald

ABSTRACT

This annual report summarizes the effect of a spike bull protection regulation
in 1971 upon an intensively managed elk herd and continues the evaluation of
specified permits on herd size, sex and age structure, hunter success and
pressure.
Estimated herd size, determined by formulas using pre and postseason
sex and age ratios in combination with kill levels decreased from about 6,800
in 1966 to 6,300 in 1971 and increased to about 6,400 in 1971. Preseason sex
ratios (bulls per 100 cows) increased from about 29 in 1966 to about 31 in 1971.
Age ratios (calves per 100 cows) increased from 58 in 1966 to 62 in 1971. Changes
in sex ratios resulted from issuing more antlerless and fewer antlered permits
to hunters in an attempt to stabilize the female segment of this herd.

��-137-

WHITE RIVER ELK EXPERIMENTAL
Raymond

MANAGEMENT

STUDY

J. Boyd

INTRODUCTION

Originally the 1971 pre-season classification counts were to be the
last field data collected on this jobo However, in February, 1971,
the Colorado Game, Fish and Parks Commission established new elk hunt
regulations for the entire state which included an across-the-board
25 percent reduction in cow elk licenses and made the spike bull an
illegal animal.
Because of the great amount of background data available on elk in
Area E, this job was amended to include check station work and postseason aerial classification counts.
Since we knew from past years
work that over 30 percent of the yearling bulls in Area E have extra
points, and thus would be legal under the new hunt regulation, attempts
were to be made to classify branch-antlered
spike bulls in both the preand post-season counts.
Data were also to be collected at check stations
on numbers of abandoned elk, by sex and age and the number of points on
antlered of abandoned bulls.

P.S. OBJECTIVE
To develop

a harvest

formula

for the White River elk herd.

SEGMENT OBJECTIVES
10

Test a candidate

2.

Publish
study.

elk harvest

the research

formula

findings

METHODS

for the White River elk herd.

resulting

from the White River elk

AND MATERIALS

Data collection (aerial and ground sex and age ratio counts, check stations
and mail questionnaires to determine harvest, age composition etc.) for this
project have not varied appreciably since 1966 (Prenzlow 1967, 1968, 1969
and 1970, Boyd 1971). Procedures for determining total elk populations
have also remained the same.
DESCRIPTION

OF AREA

Complete descriptions of the area covered by the White River elk herd
are included in Harris (1963) and Boyd (1971). Figure 1 locates the
study area in general and Area E (Game Management Units 23 and 24) specificallyo
Other descriptions of the study area can be found in Prenzlow
(1967 and 1968)0

�-138-

N

• STUDY AREA

LEGEND

Figure 1 ~ Elk :';anaeementArea E (Game Hanae:ement Units
23 and 24) showing boundaries, drainage patterns and
highway systems. Solid circles were active check
stations in 1971.

GAME MANAGDI£lIT
IIO\NIAI!Y
HtGHWI\y

--:::?

S£co~

ROAD ----

IIIYIIICll!EIC

Check ,stations

UNIT

•

LAlli

~~

_

�-139-

RESULTSAu~

DISCUSSION

Sex and Age Composition

Aerial

Surveys

Pre-season Sex and Age Ratios--The 1971 pre-season sex and age ratio
counts of the elk population in Area E were made with the aid of a helicopter on September 27 through 30, 1971. These classification counts
were confined to elk observed only in Area E (Table 1). A total of
1,047 elk were classified resulting in a ratio of 31.2 bulls (including
spikes) per 100 cows and 62.3 calves per 100 cows. Summaries of preseason sex and age ratio counts for the period 1961 through 1971 are
shown in Table 2; while Figures 2 and 3 illustrate the 95 percent confidence limits around these ratios.

Table 1.

Pre-season

Location

sex and age ratio classifications

in Area E, 1971.

Mature
Bulls

Young
Bulls

Spike
Bulls

Cows

Calves

Total

2
2

1

5
3

23
14

15
10

7
6
6
20
85
129
13
35
14
46
11
2
63
7
60

6
4
6
14
56
72
12
23
8
26
9
3
36
6
31

46
29
1
14
12
13
39
174
238
31
64
31
82
21
8
124
16
100
4

541
51.67

337

1,047

32.19

100.00

Sleepy Cat Area
Lost Park Area
Skinnyfish Creek
Anderson Res. Area
Trappers Lake Area
Shingle Peak
Trappers Peak
South Fork
Park Creek
Lost Solar Creek
Marvine Creek
East Marvine Creek
Big Ridge
Big Fish Creek
Burro Mountain
S. Side of South Fork
Patterson Creek
Oak Ridge
Ellison Mt.

3
11
6
1
4
4
4
7

5
1
1

3
13
2
8
4

Total

46

30

93

Percent

4.39

2.86

8.88

1

1
1

1
1
2
6
7
1

3
1

16
24
4
2
5
3

During these flights, an attempt was made to count branch-antlered
spike
bulls, but high winds throughout the entire count period prevented us from
flying low enough to accurately observe antler branching on yearling bulls.

�Table 2.

Summaries of elk pre-season sex and age ratio counts, Area E, 1961-1971.

Year

Bulls
No.
%

Cows
No.

1961

128

19.6

309

47.3

1962

216

22.1

462

1963

125

20.0

1964

120

1965
1966

Calves
No.

%

Total

Bulls

Ratios
: Cows : Calves

216

33.1

653

41. 4

:

100

:

69.9

47.4

298

30.5

976

46.8

:

100

:

64.5

298

47.5

204

32.5

627

41.9

:

100

:

68.5

17.5

363

52.9

203

29.6

686

33.1

:

100

:

55.9

233

17.3

699

51. 8

416

30.9

1,348

33.3

:

100

:

59.5

86

15.4

299

53. 7

172

30.9

557

28.8

:

100

:

57.5

%

1967

134

16.8

415

52.2

247

31.0

796

32.3

:

100

:

59.5

1968

135

18.7

366

50.7

221

30.6

722

36.9

:

100

:

60.4

1969

324

23.9

645

47.5

389

28.6

1,358

50.2

:

100

:

60.3

1970

150

17.6

455

53.3

249

29.1

854

33.0

:

100

:

54.7

1971

169

16.1

541

51. 7

337

32.2

1,047

31. 2

:

100

:

62.3

Average

165

18.9

441

50.4

268

30.7

874

37.4

:

100

:

60.8

I
I-'
.p.
0
I

�-141-

(I)

60

~

0
0

50

0
0
r-I

40

s...
(J)

~

30

(I)

r-I
r-I
;=j

20

CQ

0

64

61
Fig.

CJ)

~

2.

66

7C

68

95 % ccnf'Ld ence Lnt er-va Ls on IJre-season sex r'ltios

t

71

Area E, 1961-1971.

(&gt;.0

0
:)

8
r-I
s...

70
60

(!)

0..

5C

(JJ
(J)

&gt;
,.-1
:.:i
0

40
0

61
Fig. 3.

62

64

95 fo co;:firience intervals

Area Et 196J-1971.

66
on pre-season

68

70

71

ag(~ r.::.tios (calves/IOO cows) t

�-142-

Post-season Sex and Age Ratios--Post-season
sex and age ratio classifications of the elk population in Area E were conducted on December
6 through December 8, 1971. A total of 1,936 elk were classified resulting in a ratio of 14.5 bulls (including spikes) per 100 cows and
69.9 calves per 100 cows (Table 3). Summaries of post-season classifications for the period 1961 through 1971 are included in Table 4,
while Figure 4 indicates the 95 percent confidence limits around the
post-season sex ratios.

Table 3.

Post-season

Mature
Bulls

Location

West Miller
Middle

Creek

Miller

East Miller

sex and age ratio classifications
Young
Bulls 1:../ Spikes

1

7

Branched
Spikes

Cows

Calves

Total

11

76

53

148

9

5

14

1

15

13

29

Creek

Creek

Hwy. 13 to Yellowjacket

1

12

4

123

82

222

2

32

8

141

98

283

8

123

99

248

93

76

175

28

19

51

1

3

7

30

23

53

240

171

430

Coal Creek Area

2

Big Beaver Creek

2

16

Burro Mt. Area

3

3

South Fork Canyon

2

Park Creek

3

Bloomfield

in Area E, 1971.

2

Bench

South Fork-Campground
down to Buford

1

Oak Ridge

2

Total

10

Percent

0.5

1:../ Typical antler formation

3

15
10

1

171

92

276

7

100

35

1,050

734

1,936

0.4

5.2

1.8

54.2

37.9

100.0

(4-5 points),

but light beam.

The most obvious facet of these counts was the very apparent lack of large,
mature bulls.
Of the 17 bulls seen that were older than yearlings, only two
were large 6-point bulls, the rest being small 5- and 6-point males.

�Table 4.

Summaries of elk post-season sex and age ratio counts, Area E, 1961-1971.

No.

%

No.

%

No.

%

Total

Ratios
Bulls : Cows : Calves

1961

106

10.6

465

46.5

430

43.0

1,001

22.8

:

100

:

92.5

1962

125

13.8

446

49.2

335

37.0

906

28.0

:

100

:

75.1

1963

45

6.1

413

55.7

283

38.2

741

10.9

:

100

:

68.5

1964

117

6.0

1,130

58.3

692

35.7

1,939

10.4

:

100

:

61.2

1965

85

5.2

930

57.1

613

37.7

1,628

9.1

:

100

:

65.9

1966

128

6.1

1,245

58.9

739

35.0

2,112

10.3

:

100

:

59.4

1967

214

9.2

1,318

56.4

806

34.5

2,338

16.2

:

100

:

61.2

1968

148

8.2

951

52.9

699

38.9

1,798

15.6

:

100

:

73.5

1969

166

9.0

1,012

55.0

661

36.0

1,839

16.4

:

100

:

65.3

1970

120

4.9

1,402

58.3

884

36.8

2,406

8.6

:

100

:

63.0

1971

152

7.9

1,050

54.2

734

37.9

1,936

14.5

:

100

:

69.9

Average

128

7.6

942

55.6

625

36.8

1,695

13.6

:

100

:

66.3

Year

Calves

Cows

Bulls

I

I-'
.pw
I

�-144-

(I,)

~

0
0
0
0
r-I
H

40
30

&lt;1&gt;

c,
(I,)

r-I
r-I
:;$

P=l

20
10
0

61

62

6=~

64

65

66

67

68

69

70

71

Years

Fig. 4. 95% confidence intervals on post-season sex ratios, Area
E, 1961-1971.

An examination of Table 5 shows the reduction in mature bulls per
100 cows that occurred as a direct result of the 1971 elk hunt regulation. In one year, the mature male segment of this herd was
reduced back to levels present in Area E at the time specified permits were suggested as one means of increasing this age segment of
the male portion of the herd.

Table 5.
1971.

Year

Comparison of bull elk ratios, post-season, Area E, 1964-

All Bulls

Bulls Per 100 Cows
Mature Bulls

Yearling Bulls

1971

14.5

1.6

12.9

1970

8.6

3.9

4.7

1969

16.4

2.9

1968

15.6

3.8

13.5
11.8

1967

16.2

12.6

1966

10.3

3.6
1.1

1965

9.1

2.8

6.3

1964

10.4

1.6

8.8

9.2

�-145-

Another item of interest is the number of branch-antlered spike bulls
observed.
This was the first year that a separation of this kind was
attempted and several important facts were learned.
I feel a snow
background is mandatory for this type of classification and that time
spent in pre-season counts trying to delineate branch-antlered spikes
would be a waste of time and money.

Ground Surveys
During the years 1967 through 1971, a ground classification of elk in
Area E was made each July. This was done to estimate yearling-adult
cow ratios which cannot be accurately obtained during aerial classifications.
These data also serve as independent estimates of sex and
age structures.
It becomes very important to determine the proportion
of yearling females in the herd and thus their influence on overall
herd productivity.
Usually only a small fraction of the yearling
females breed, but there is considerable conflict in the literature
on this particular subject.
Observers on horseback, riding during the early morning hours, classified elk in different portions of the Flat Tops Primitive Area.
In
three days, four teams of two men each classified a total of 1,039 elk
(Table 6). Many more elk were seen that could not be accurately classified.
The average size of elk groups observed was 23.1 animals.

Table 6. Composition
in Area E, 1971.

Item

of 1,039 elk classified

during July ground counts

Mature
Bulls

Hature
Cows

Yearling
Bulls

Yearling
Cows

Calves

Total

Number Observed

62

82

498

142

255

1,039

Percent Observed

6.0

7.9

47.9

13.7

24.5

100.0

25

22

37

24

28

45

Mean Group Composition~./ 1.4

1.8

11.1

3.2

5.6

23.1

Frequency

in Groups ])

1/ Frequency

in groups

(i.e. mature bulls were observed

in 25 of 45 total

groups) .

2/
-

Mean group composition

23.1

determined

by percent

of elk observed

times

�-146-

Check Station Surveys

Number of Elk Checked--Only one special research check station was operated during the 1971 elk season. No research personnel were stationed
at any of the regular management check stations to collect data from
Area E elk. A total of 280 elk were checked out through the Meeker
station. Table 7 lists the number of elk checked through Meeker from
the White River area, 1965 through 1971.

Table 7. Number of elk checked through Meeker, 1965-1971.

1970

Average

1971

Percent Change
from
5-year Average

Day

1966

1967

1968

Year
1969

1

15

11

11

10

19

l3

8

- 38.5

2

45

46

46

21

36

39

l3

- 59.6

3

75

69

117

71

105

87

32

- 61.9

4

115

71

l34

152

no

116

37

- 64.5

5

122

9l

182

l38

129

l32

51

- 63.7

6

91

94

99

123

l31

108

43

- 62.8

7

61

76

93

64

80

75

49

- 59.1

8

43

36

31

38

64

42

23

- 58.2

9

24

29

27

20

39

28

24

- 56.3

Total

59l

523

740

637

713

640

280

- 56.3

Table 8 indicates the daily elk check-out from the two Area E units through
the Meeker check station in 1971.

Sex and Age Composition of Elk Checked--The percent composition (bulls, cows
and calves) of elk checked in 1971 from Area E is shown in Table 9. Sex and
age ratios of the 206 elk from Area E were 52.43 percent bulls, 39.32 percent
cows and 8.25 percent calves. This compares with 61.5 percent bulls, 31.6
percent cows and 6.9 percent calves in 1970. During the first three years of
the experimental management study, a constant number of antlered permits were
issued for Area E (2,500). In 1969, however, antlered permits were reduced

•

�-147-

to 2,200; 1970, 3,000 antlered permits and 1971, 2,200 branch-antlered
only permits were sold. Ant1er1ess permits varied from 1,000 in 1966
to 1,500 in both 1967 and 1968, down to 1,000 in 1969, 800 in 1970 and
600 in 1971. These changes in permit numbers account, in part, for the
changes in bulls to cows to calves in the yearly harvest.

Table 8.

by day of season, Area E, 1971.

Elk check-out

Unit and Sex

Day of Season
4
5
6

1

2

3

Males

1

1

1

4

4

Females

0

4

1

7

Sub-total

1

5

2

Males

0

5

Females

3

Sub-total

7

8

9

Total

3

5

3

1

23

10

0

3

4

1

30

11

14

3

8

7

2

53

12

9

17

9

19

8

9

88

3

10

5

14

13

9

1

7

65

3

8

22

14

31

22

28

9

16

153

Males

1

6

13

13

21

12

14

11

10

111

Females

3

7

11

12

24

13

12

5

8

95

Total

4

13

24

25

45

25

26

16

18

206

Percent of
Total Check

1.9

6.3 11.6 12.1 21.8 12.1 12.6

7.8

8.7

Unit 23

Unit 24

Area E

�-148-

Table 9. Composition
from Area E, 1971.

of elk checked through the Meeker

check station

Bulls

%

Cows

%

Calves

%

Total

Unit 23

22

41.5

26

49.0

5

9.5

53

Unit 24

86

56.2

55

35.9

12

7.9

153

Total

108

52.4

81

39.3

17

8.3

206

Age Structure of the Harvest--Of the 206 elk checked in 1971 from Area E,
126 were aged using techniques described by Quimby and Gaab (1957). Table
10 lists these ages, by sex, while Table 11 details similar information by
game management unit.
One incisor tooth was collected from each elk that came through the station with a jaw present.
These teeth were taken to the Research Center
Laboratory in Fort Collins where they were aged according to the dental
cementum technique reported by Keiss (1969). Assuming that ages determined by dental cementum were correct, this comparison revealed an error
of only 4.6 percent in assigning ages to 2-l/2-year old elk, while the
error associated with aging elk older than 2-l/2-years was 64.7 percent
(Table 12).
Table 13 indicates the distribution of error by age class. Overall
error was 41.1 percent.
This compares with 60.4 percent in 1967, 43.1
percent in 1968, 40.1 percent in 1969 and 28.9 percent in 1970. Of 126
check station assigned ages from Area E, 56 were comparatively aged by
the dental cementum technique.
Of these 56 animals aged by the two
methods, 17 (30.4 percent) were field aged within one year on either side
of the actual age as determined by dental cementum procedures.

�Table 10.

Sex and age determinations of 126 elk checked from Area E, 1971.
Age (years) 1/
5-1/2
4-1/2
3-1/2

6-1/2

7-1/2

8-1/2

9+

Total

4

0

1

1

0

0

72

6.9

5.6

0

1.4

1.4

0

0

100.0

19.0

4.0

3.2

0

0.8

0.8

0

0

57.2

5

9

7

3

2

0

1

2

54

9.2

16.7

l3.0

5.6

3.7

0

1.8

3.7

100.0

7.9

4.0

7.1

5.6

2.4

1.6

0

0.8

1.6

42.8

17

45

29

14

11

3

3

1

1

2

126

l3.5

35.7

23.0

11.1

8.7

2.4

2.4

0.8

0.8

1.6

100.0

1/2

1-1/2

2-1/2

Number

2

35

24

5

Percent

2.8

48.6

33.3

Percent "I'o
t aL

1.6

27.8

Number

15

10

Percent

27.8

18.5

Percent Total

11.9

Sex
Males

Females
I

I--'

~

&lt;o
I

Total Elk
Number
Percent Total

-!/ Ages assigned by techniques described by Quimby and Gaab (1957).

�Table 11.

Sex and age determinations of 126 elk aged from Area E, by unit of kill, 1971.

Unit and Sex

Age {years~ 1:/
4-1/2
5-1/2

6-1/2

7-1/2

8-1/2

9+

Total

0

0

0

1

0

0

17

11.8

0

0

0

5.9

0

0

100.0

8.3

2.8

0

0

0

1.4

0

0

23.6

18

3

4

0

1

0

0

0

55

1/2

1-1/2

2-1/2

3-1/2

Number

0

8

6

2

Percent of Total

0

47.0

35.3

Percent of all Males

0

11.1

2

27

Unit 23 - Males

Unit 24 - Males
Number

I

I-'

Percent of Total

3.6

49.1

32.7

5.4

7.3

0

1.8

0

0

0

100.0

Percent of all Males

2.8

37.5

25.0

4.2

5.6

0

1.4

0

0

0

76.4

5

5

1

6

2

2

1

0

0

2

24

Percent of Total

20.8

20.8

4.2

25.0

8.3

8.3

4.2

0

0

8.3

100.0

Percent of all Females

9.2

9.2

1.8

11.1

3.7

3.7

1.8

0

0

3.7

44.4

10

5

4

3

5

1

1

0

1

0

30

Percent of Total

33.3

16.7

13.3

10.0

16.7

3.3

3.3

0

3.3

0

100.0

Percent of all Females

18.5

9.2

7.4

5.6

9.2

1.8

1.8

0

1.8

0

55.6

Unit 23 - Females
Number

Unit 24 - Females
Number

V1
0
I

�-151-

Table 12. Percent error between dental cementum and tooth replacement
and wear aging techniques, by age class, of 56 elk from Area E, 1971.
Age
Class

Dental
Cementum

Calf

1/

1

1/

2

Field
Age ]j

Percent
Correct

Percent
Error

22

21

95.4

4.6

3

8

6

75.0

25.0

4

11

3

27.3

72.7

5

7

1

14.3

85. 7

6

2

0

0

100.0

7

2

0

0

100.0

8

1

0

0

100.0

9+

3

2

66.7

33.3

Total

56

33

58.9

41.1

1/

1/

- Number of teeth checked in the laboratory
procedure.
~/ Number of ages correctly assigned
wear and replacement procedures •

using the dental

cementum

at the check station using tooth

. _3/ Calves and yearlings were not aged by the dental cementum technique.

�-152-

Table 13. Distribution of assigned ages in relation
mined by dental cementum procedures, Area E, 1971.
Cementum
Age Class

2

2

22

3

3

4
5

Check Station Assigned·Ages

3

4

5

6

7

9+

Total

23

8

2

13

1

4

11

1

1

2

3

7

2

7

1

14

2

4
2

1

14

19

Mortality

9

4
1

9+

27

18

1

2

8

Harvest

8

1

6

Total

to ages deter-

5

3

2

1

3

4

2

3

82

Estimates

Estimates

Random Survey--Because of annually decreasing hunter report card returns, no
report cards were attached to the 1971 elk license and a random survey was
run on selected elk license buyers to determine the 1971 elk harvest estimate.
While elk hunters with Area E permits were surveyed along with all
other elk hunters, an extra large sample (1,370 out of 2,743 permittees,
a 49.9 percent sample) was taken from this hunter population to correspond
with random samples drawn from hunters in this area since 1968. Of the
1,370 hunters sampled, 1,318 (96.20%) returned usable cards. The estimated
1971 total elk kill in Area E was 632 (C. I. ± 81; ~ = 1.96), with 25.29 percent of the active hunters being successful.
Table 14 gives results of the
1971 random survey, including kill by type of license, while Table 15 lists
active (those who actually hunted elk in 1971) hunters and success ratios.
Projections from the random survey indicated that 244 licenses were issued
that were not used (8.9%). The number of "no hunts" in Area E have
averaged 7.0 percent since 1966.

�-153-

It should be pointed out, here, that data shown in Table 14 does not
correspond with kill estimates, numbers of hunters or success ratios
calculated and published as the 1971 harvest estimates by the Game
Management Section.
Data published by Game Management is based on a
random card return of 1,246, while Table 14 is based on a card return
of 1,318.
There are two possible reasons for this difference: (1) The 72 additional cards counted by personnel of this project could have come into
the Denver Office after the cut-off date and were placed in the file
by mistake where they were counted by hand after the computer run, or:
(2) When follow-up letters were sent to non-reporting hunters, new
questionnaires were included and some cross-over could have occurred.
According to Denver Office personnel there is no way we can eliminate
duplicate card returns for 1971. This has been remedied for 1972.

Table 14.

Results

of the 1971 random survey of 2,743 Area E permittees.
Antlered
Res.
N. Res.

Item

Permits

Issued

Permits
Percent

Antlerless
Res.
N. Res.

Total

1,482

6S6

499

106

2,743

in Sample

740

329

248

S3

1,370

in Sample

49.93

SO.lS

49.70

SO.OO

49.94

711

322

237

48

1,318

Sample Returned
Percent

Returned

96.08

97.87

9S.S6

90.S7

96.20

Percent

of Total Permits

49.79

49.08

47.49

4S.28

48.0S

Reported

"No Hunts"

81

20

l3

3

117

Estimated

"No Hunts"

169

41

27

7

244

Reported

Kills

98

61

114

29

302

Estimated

Total Kill

204

124

240

64

632

�-154-

Table 15. Number of hunters participating during the 1971 Area E
elk hunt and success ratios per license and active hunters.
Projected from random survey data.

Res.

Item

Active

Hunters

Antlered
Non. Res.

Antlerless
Res.
Non. Res.

Total

1,313

615

472

99

2,499

Success

(kill) Per Lie.

13.76

18.90

48.10

60.38

23.04

Success

(kill) Per Hunter

15.54

20.16

50.85

64.65

25.29

Harvest

in Relation

to Area and Time

Location of Kill--Kill by Game Management Unit was projected from the
random survey data and appears in Table 16. These data indicated that
60.44 percent of the 1971 elk kill in Area E occurred in unit 24.

Table 16.
Management

Composition
Unit.

Unit

Bulls
%
Number

of the 1971 estimated

elk kill in Area E by Game

Cows
Number

%

Calves
Number

%

Total
Number

%

Unit 23

120

48.00

124

49.60

6

2.40

250

39.56

Unit 24

208

54.45

143

37.43

31

8.12

382

60.44

Total

328

51.90

267

42.25

37

5.85

632

100.00

Date of Kill--Dates of kill by hunters in Area E are shown in Table 17.
Approximately 90.3 percent of the kill occurred during the first five
days of the season.

Hunter Effort--Elk hunting in Area E provided an estimated 11,877 mandays of hunting recreation in 1971. This was a decrease of 9,077 recreation days compared to 1970. This decrease was due, primarily, to a 25
percent reduction in the number of antlerless permits compared to 1970
and further, a 26.67 percent reduction in antlered permits and the further
restriction making spike bulls illegal.
Mean number of days spent elk
hunting decreased from 5.9 days in 1970 to 4.76 days in 1971.

�-155-

Table 17.

Numbers

of elk killed, by day of season, Area E, 1971.

1

2

Day of Season
3
4
5
6

Unit 23

17

7

9

11

4

Unit 24

58

24

14

22

Total

75

31

23

33

Estimates

of Mortality

7

8

9

Total

1

2

2

0

53

20

1

6

4

4

153

24

2

8

6

4

206

Other Than Legal Kill

Wounding Loss--Annual estimates of wounding losses have been gathered in
Area E since 1966. In 1971, the random questionnaire was designed and
sent out by the Game Management Section in Denver and questions concerning
wounding of elk and retrieval of wounded elk were not included on the
questionnaire.
No comparable data for 1971 are thus available.

Population

Total Population

Estimates

and Projections

Estimates

Formulas were used to estimate the total number of elk present in Area E
prior to the 1971 elk hunt. Detailed descriptions of the method are presented in Prenzlow (1968) with examples of how ratios of antlered to
antlerless elk from kill, pre- and post-season classification counts,
were used in a formula devised by Dr. David Bowden of Colorado State
University, to estimate a pre-hunt elk population.

Pre-Season Population Estimate, 1971--An estimated 3,358 (C. I. = 85,
g = 1.96, ~ = .05) elk were present in Area E just prior to the opening
of the 1971 elk hunt (Table 18). This compares to 6,751, 6,313, 6,216,
5,829 and 6,347 estimated in 1966, 1967, 1968, 1969 and 1970, respectively.

�-156-

Table 18. Pre-season population
sex and age in Area E, 1971.

Item

Estimated
Number

Bulls

estimate

and proportions

of elk by

Expected
Percent 1./

Observed
Percent ']j

542

16.29

16.14

Cows

1,735

54.11

51.67

Calves

1,081

29.60

32.19

Total

3,358

100.00

100.00

1./
Percentages

expected

from 1970 post-season

population

data.

2/
- Percentages actually observed when making
classifications
from a helicopter.

the 1971 pre-season

elk

Until 1969 and again in 1971, total population data were believed to be
reasonable estimates of actual elk population levels in Area E. For
instance, predictions for the percentages of bulls, cows and calves
made after the post-season counts were within five percent of what was
actually observed during pre-season counts the next year during all years
of the study (1961-1971) with the exception of 1969. In 1969, the expected
and observed percentages of bulls differed almost 20 percent.
This was a
critical difference, because the change in ratios of bulls from pre- to
post-season is a major component of the population estimate formula used
to estimate the pre-season elk population.
However, as Prenzlow (1970:
280) explained, early severe September snow storms in 1969 probably caused
an abnormal movement of the elk herd and samples drawn in 1969 were not
taken from the "normal" herd usually found in Area E.
The 1971 pre-season population estimate of 3,358 appears to be extremely
low for four reasons: (1) 4,271 elk were actually counted in Area E
during the 1972 trend count which exceeds the total pre-season population
estimate by 913 animals, plus the fact that an additional 632 animals
were harvested, resulting in a minimum pre-hunt population of 4,903
animals; (2) The 3,358 elk estimated to be in Area E does not fit the
trend in population numbers over the past years and it is less than onehalf the number of elk that should have been in this area based on the
1970 post-hunt population estimates for this herd; (3) The estimated
harvest of 632 could not have reduced the population to this low level,
even if an additional 10 percent wounding loss is added, and (4) The
bull kill estimate is low due to an extremely high spike bull kill that
went unreported in 1971, because of the new spike bull protection
regulation.
Estimates of the loss of spike bulls in Area E for the 1971

�-157 -

range from 63 to 139 which if added to the estimated kill would increase the population estimate some 1,600 animals.
This unrecorded
loss of yearling bulls would have a great depressing effect upon the
formula population estimate.
Therefore, to accomplish the objective of testing an elk harvest formula, it became necessary to use the 1970 projected population figure
of 6,572 (Boyd 1971) rather than the 1971 estimate of 3,358. This
was thought justified because past projections, after being corrected
for over-winter loss, have been close approximations of total elk
population estimates derived by formula.

Post-Season Population Estimates, 1971--An estimated 5,940 Area E
elk survived the 1971 hunting season (Table 19). This was calculated
by subtracting the elk harvest, as determined by the random survey, by
class of animal from the pre-season population.

Table 19. Post-season population
sex and age in Area E, 1971.

estimate

and proportions

of elk by

Post-Season POEulation
Exp. % J:./
Obs. % 1/
No.

Item

Pre-Season
Population

Bulls

1,038

328

710

12.00

7.85

Cows

3,450

267

3,183

53.58

54.24

Calves

2,084

37

2,047

34.42

37.91

Total

6,572

632

5,940

100.00

100.00

Harvest 1/

1/
Based upon the 1971 random

survey.

2/
- Proportions, by class, of the total estimated number
that remained following the 1971 elk hunting season.

3/

- Proportions actually observed when making
fications by helicopter.

of elk (5,940)

the 1971 post-season

classi-

�-158-

Total post-season elk populations in Area E declined from 5,336 in
1966, 4,945 in 1967, to 4,506 in 1968, then increased slightly to
4,646 in 1969, 4,987 in 1970 and increased rather strongly to 5,744
in 1971. The decline in the post-season population during the early
years was primarily the result of issuing more antlerless permits
in order to change the composition of the wintering herd to meet one
of the objectives of narrowing the sex ratio (more bulls per 100 cows).
Proportions of bulls in the post-season elk populations have increased
from 6.1 percent in 1966 to 9.2, 8.2, 9.0 in 1967, 1968 and 1969, decreased to 5.0 in 1970 and again increased to 7.8 percent in 1971.

Population

Projections

Each year, based upon the previous years post-season population estimate, a projection is made for the next year.
This procedure is
described in Prenzlow (1968) in detail.

Population Projection for Area E, 1972--Immediately
prior to the opening
of the 1972 elk hunt in Area E, a projected elk population of 8,441
animals will be available to hunters in this area (Table 20). This figure
may change somewhat, however, if productivity differs significantly from
59.5 calves per 100 cows.

Table 20.

Predicted

1972 pre-season

elk population,

1971 Post-Season
Population

Item

Area E.

1972 Pre-Season
No.

Population
Exp. % ~/

1/

20.53

3,183

4,206 1/

49.83

Calves

2,047 1/

2 ,502 4/

29.64

Total

5,940

8,441

100.00

Bulls

710

Cows

1/

1,733

.

- Assumlng that of 2,047 calves, 50 percent
cent are females.

(1,023) are males and 50 per-

~/Calculated by adding the remaining
calves from 1971.

bulls in 1971 (710) and 1,023 male

1/ Calculated

cows in 1971 (3,183) and 1,023

female

by adding the remaining
calves from 1971.

4/ Calculated by multiplying the number of cows in 1972 (4,206) by 59.5
percent expected productivity.
5/ Expected proportions of total projected
to the 1972 elk hunting season.

numbers

of elk (8,441) prior

�-159-

In previous years, population projections have been high because no
losses other than the estimated hunting mortality were considered.
Annual herd losses of 17.5, 11.5 and 4.3 percent were determined
for 1967, 1969 and 1970 respectively.
These percentages were
calculated by subtracting a particular years population estimate
from the projection (Table 21).

Table 21.

of herd losses in Area E, 1966-1971.

Estimates

Year

Population
Estimate

1966

6,751

1967

6,313

7,654

1,314

17.5

1968

6,261

7,075

814

11.5

1969

5,829 1/

5,829

1970

6,347

6,632

285

4.3

1971

6,572 ].)

6,376

II

Corrected

because

Population
Projection

of obvious sampling

Difference
Number
Percent

error.

~I Corrected because of large increase in illegal kill of spike bulls.

Consequences of 1972 Area E Elk Hunt--In 1971, the management of the
Area E elk herd was turned back to Regional control and the Research
Section made no more season recommendations.
In order to assess the
impact of the 1971 spike bull protection regulation, this project was
continued, but data included in the balance of this report will be based
upon the 1972 hunt regulations proposed by the Northwest Region personnel.
The 1972 Area E hunt recommendation was recommended to be 2,500 antlered
permits, with spike bulls being legal, and 800 antlerless permits.
Table
22 illustrates the composition of the 1972 pre-season Area E elk population if the projection is reduced by 10 percent to account for overwinter herd losses.
The resulting post-season population is projected
based upon no significant change in hunter success in 1972.

�-160-

Table 22. Modified 1972 Area E pre-season population projection and
resulting post-season population with a kill level determined by
2,500 antlered and 800 antlerless permits.
Modified 1972 PreSeason Population

1972 Post-Season

Item

Number

Exp. %

Projected
Kill

Bulls

1,560

20.53

625

935

14.49

Cows

3,786

49083

442

3,344

51.83

Calves

2,251

29.64

78

2,173

33.68

Total

7,597 1/ 100.00

1,145 2/

6,452

100.00

1/ Modified

POEulation
Number
Exp. %

projection resulting from subtracting
the theoretical projection of 8,441 in Table 20.

844 elk (10%) from

~/ Projected kill is based upon 25 percent success on 2,500 antlered
permits and 65 percent success on 800 antlerless permits.

The 935 bulls and 3,344 cows in the estimated
give a ratio of 27.96 bulls per 100 cows.

post-season

population

As a check on the above projections, ratios of bulls, cows and calves
observed during classification counts will be compared to expected
ratios; kill data will again be determined by random survey; and preseason population estimates will be calculated by formula.
It is
important to note that these comparisons will be independent estimates
of the same population parameter, and if there is close agreement, it
would indicate that the above described method is valid for estimating
the elk population in Area E.
A first draft of the final report on the effect of specified permits
on the sex and age structure of the White River elk herd is nearly
complete and should be in the hands of the Project Leader by June 15,

1972.

�-161-

LITERATURE

CITED

Boyd, R. J. 1971a. Elk of the White River plateau, Colorado.
Colo.
Game, Fish and Parks Div., Denver.
Tech. Publ. No. 25. 126 p.
illus.
1971b. White River elk herd - population
Job Progress Report, July, No.2. W-38-R-25.
Harris, J. T.
Colorado.
190 p.

components.
p. 113-138.

Colo.

1963. Population dynamics of the White River elk herd,
PhD Thesis. Univ. of Michigan, Ann Arbor, Mich.

Keiss, R. E. 1969. Comparison of eruption-wear patterns and cementum
annuli as age criteria in elk. J. Wildl. Mgmt.
33(1):175-180.
Prenzlow, E. J. 1967. Population components - White River elk.
Colorado.
Job Completion Report, W-38-R-2l.
p. 251-276.
1968. White River elk population components.
Progress Report. W-38-R-22.
p. 383-421.

Colorado Job

1970. White River elk population components.
Progress Report. W-38-R-24.
p. 257-286.

Colorado Job

Quimby, D. C., and J. E. Gaab. 1957.
indicator in Rocky Mountain elk.

Prepared by __ ~~~~~~~~~~~,
Raymo d J. Boyd
Wildlife Researcher

Mandibular dentition as an age
J. Wildl. Mgmt.
21(4):435-451.

��-163-

July, 1972

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No.

W-38-R-26

Work Plan No.
Job Title

Investigations
1

Job No.

Middle Park Deer Study - Population

Period Covered:
Personnel:

14

Deer-Elk

Distribution

April 1, 1971 through March 31, 1972

Greg Ericks, R. Bruce Gill and Laren A. Roper.

ABSTRACT

All deer trapping records and marked deer observations from 1969-1972 were
coded for an automatic data processing program which will be used to summarize these data so that deer population sub-unit boundaries can be defined.

��-165-

MIDDLE

PARK DEER STUDY - POPULATION

DISTRIBUTION

Laren A. Roper

P. S. OBJECTIVE
To delineate deer concentration areas in Middle Park and relate changes
in deer distribution in time and space to accumulation and physical
properties of snow.

SEGMENT OBJECTIVES
1.

Define deer population

See Gill

sub-unit

boundaries.

METHODS

AND MATERIALS

RESULTS

AND DISCUSSION

(1969).

Trapping

and Harking

An automatic data processing program was completed and all deer trapping
records and marked deer observations from 1969-72 were coded.
During the
next segment the program will be debugged and the data will be summarized.
A department publication describing deer population sub-unit boundaries
will be completed for the final report.

:'-)
Prepared

,T

_'~.~~a:"",,,,-~~.-"-__ -. _

by ---',d--=-""",,'L~;/~~"--"~L--,C,,-,-l-=--,
'taren A. Roper
Asst. Wildlife Researcher

��-167-

July, 1972

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~---------

Project No.

W-38-R-26

Work Plan No.

14

Deer-Elk

2

Job No.

Job Title

Middle Park Deer Study - Population

Period Covered:

October,

Personnel:

Investigations

1971 through January,

Density

and Structure

1972

R. Bruce Gill, Laren A. Roper, Gary L. Brown, Len H. Carpenter
Paul F. Gilbert.

and

ABSTRACT

Winter range density of the Middle Park deer population was estimated from
Estimated density
helicopter counts of deer on square mile sample quadrats.
population
projection
for 1972 was 9024
3.26 deer per square mile for a total
the
Blue
River
subof 5,429 + 1,913 deer. Empirical evidence suggested that
Park
deer
population
population was greatly underestimated and the true Middle
numbered approximately 7,000 deer.

±

Pre and postseason classification counts indicated fawn production in 1971 was
similar to 1970 but lower than 1969. Comparisons of aerial and ground classifications for the past five winters revealed no significant differences between
fawn:doe ratios obtained from each method, but buck:doe ratios differed significantly in two of the five years.

��-169-

MIDDLE PARK DEER STUDY
POPULATION DENSITY AND STRUCTURE
R. Bruce Gill

P. S. OBJECTIVE
To estimate the density and sex and age structure of the Middle Park
deer population in order to harvest this population more efficiently.

SEGMENT OBJECTIVES
1.

Estimate
Park.

the size of the total winter population

of deer in Middle

2.

Estimate

the sex and age structure

Park deer population.

METHODS
Methods and materials
Gill (1971).

of the Middle

AND MATERIALS

have been detailed

previously

in Gill

(1969) and

RESULTS AND DISCUSSION

Density
Estimates of the 1971-72 Middle Park winter deer population level indicated
a total population of 5,429 ± 1,913 deer, or 9.24 ± 3.26 deer per square
mile of winter range (Table 1). Empirical evidence suggested this figure
was conservative.
Three of the five sub-unit population sample areas
exhibited population increases compared with 1970-71 levels while two
appeared to decline (Table 2). The apparent decline in the Granby subunit estimate is of little significance to the total population, but the
Blue River decline is of considerable importance.
There was no logical
explanation why the Blue River population should have declined by approximately 1,100 deer from the 1970-71 estimate.
In fact, it should have
either remained stable or increased in about the same proportion that
the Muddy Creek, William Fork River, and Troublesome Creek sub-unit populations did. Winter loss estimates for the 1970-71 winter were low (489
deer) as was the 1971 harvest (348 deer) and even if all of the mortality
associated with these two sources had occurred within the Blue River there
still should have been an increase of 0.4% in the Blue River population.
On the other hand, if the Blue River population had actually increased the
same as the other four sub-unit populations did (32.4%), the Blue River
population should have been 4,414 bringing the total estimated Middle Park
population to 7,586 deer. This would have been considerably closer to the
expected or calculated population estimate (Gill 1971) of 7,072 deer (Table
3) •

�Table 1.

Numbers of deer counted per quadrat within eight sampling strata, Middle Park, Colorado, 1968-1972.

Mudd~ Creek High Densit~
1968 1969 1970 1971 1972
2
0
0
0
0
8
64
32
124
58
30
32
NC
44
NC
20
27
0
1
NC

0
0
0
0
0
9
6
18
99
39
66
46
87
31
6
12
0
0
22
37

0
0
0
0
0
9
34
9
20
6
2
2
42
28
5
12
0
0
5
1

0
12
4
0
0
0
0
0
28
68
2
0
22
0
0
0
0
0
0
0

0
0
0
0
0
1
8
11
24
39
0
5
35
6
1
1
0
0
0
27

~ = 442

478

175

136

158

Y = 26.0 23.9

8.8

6.8

7.9

Mudd~ Creek Low Densit~
1968 1969 1970 1971 1972

Blue River High Densit~
1968 1969 1970 1971 1972

Blue River Low Densit~
1968 1969 1970 1971 1972

56

14
25
0
76
31
4
9
134
49
71
124
28

4
10
0
30
32
0
0
31
0

NCll
0
0
NC
0
NC
0
NC
0
NC
NC

6
0
0
0
0
0
0
0
0
0
15
0

0
1
0
30
0
0
0
0
0
0
11
0

0
0
0
0
0
0
0
0
0
0
0
0

0
2
0
0
0
0
0
0
0
0
0
6

19
0
1
36
88
20
4
102
15
105
58
17

14
0
0
24
77
18
3
20
83
136
49
6

47
0
0
145
54
7
0
34
43
67
104
0

9
0
6
19
82
41
29
15
22
64
52
0

8
10
0
0
35
0
0
36
29

0
17
0
37
34
0
0
26
0

0
13
0
0
15
0
0
20
0

0
0
0
0
30
0
0
2
0
I
I-'
-....J

0
I

56

21

42

0

8

565

465

430

501

339

107

118

114

48

32

9.3

1.8

3.5

0.0

0.7

47.1

38.8

35.8

41.8 28.3

11.9

13.1

12.7

5.3

3.6

-------------------------------------------------------------------------------------------------------------------------

�Table 1. Numbers of deer counted per quadrat within the eight sampling strata, Middle Park, Colorado, 1968-1972
(continued).
l&lt;lms.Fork River High Densitz
1968 1969 1970 1971 1972

Wms. Fork River Low Densitz
1968 1969 1970 1971 1972

Troublesome Cr. Hi~h Density
1968 1969 1970 1971 1972

0
2
26
73
4

0
6
14
0
0

43
25
0
35
50
65
0
9
0
26

40
0
45
42
4

0
0
25
21
0

3
0
58
31
0

9
0
52
71
0

0
0
3
0
0

0
0
1
0
0

0
4
0
0
0

0
0
0
0
0

5

41
0
39
76
44
15
36
0
6

11
60
0
51
42
90
7
8
0
29

0
8
0
0
43
36
14
21
10
7

3
24
2
70
20
40
27
21
0
12

1968
z

0
r+

n

0

.::
::s
ro
c,

r+

Granbz Low Densitz
1969 1970 1971 1972
7
2
2
0
0

0
NC

6
23
0
0
0
0
0

0
0
13
0
0

0
0

1
0
1
0
0
0
0

I

t-'

-...J

I: = 105

131

46

92

132

20

3

1

4

0

253

262

298

139

219

11

29

13

2

26.2

9.2

18.4

26.4

4.0

0.6

0.2

0.8

0.0

25.3

26.2

29.8

13.9

21.9

1.8

4.1

1.9

0.3

-

Y = 21.0

1968

1969

1970

1971

1972

= 20.75

15.52

12.29

9.75

9.24

var (yst) = 11.35

4.25

2.55

3.29

1.96

yst

1/
- NC = No count.

t-'
I

�-172-

Table 2.

Deer population

estimates

by sub-units

in Middle

Park, 1968-

1972.

Year

Muddy
Creek

Blue
River

Williams
River

1968

4,101

4,074

1969

2,994

1970

Fork

Troublesome
Creek

Granby

Total

843

1,622

no count

10,640

3,531

774

1,679

134

9,112

1,429

3,290

271

1,910

306

7,206

1971

794

3,335

571

891

139

5,730

1972

1,002

2,259

742

1,404

22

5,429

Table 3. Comparisons of expected population
census estimates in Middle Park, 1968-1972.

levels with actual quadrat

Year

Expected
Population

Census
Estimate

Percent
Difference

1969

9,519

9,112

+ 4.3% 1/

1970

7,862

7,206

+ 8.3%

1971

5,830

5,730

+ 1.7%

1972

7,072

5,429

+ 30.3%

1/
Indicates the amount that the expected population
census estimate.

differs

from the quadrat

There was evidence that significant numbers of deer still remained above the
winter range sample areas of the Blue River at the time the census was conducted.
Fresh deer tracks were abundant in timbered areas above 9,500 feet
in the Ute Pass vicinity which is considered fall and spring transitional
range.
None of the other four sub-unit areas exhibited similar abnormal
deer distribution patterns.
Therefore it was concluded that the actual
Middle Park deer population level approximated 7,000 deer, or 11.9 deer per
square mile of winter range.

�-173-

Sex and Age Composition

Classification

Counts

Pre-season Counts--The 1971 pre-season deer classifications were conducted
from helicopter flights on October 13 and 14 after most of the aspen leaves
had fallen.
These counts were generally unproductive, and only 91 deer
were classified compared with 771 in 1970 and 1,065 in 1969 (Table 4).
Lack of snow cover during the 1971 classifications was a major factor contributing to the large decline in numbers of deer observed.

Table 4.
Park.

Comparisons

of 1969-1970

pre-season

deer classifications

in Middle

Buck:Doe
Ratio

Fawn:Mother
Ratio ])

FaW'n:Doe
Ratio

Year

Bucks

Does

Fawns

Total

1969

195
(18.3%)

451
(42.4%)

419
(39.3%)

1,065
(100.0%)

43

100

93

100

125

100

1970

152
(19.7%)

361
(46.8%)

258
(33.5%)

771
(100.0%)

42

100

72

100

112

100

1971

20
(22.0%)

41
(45.0%)

30
(33.0%)

91
(100.0%)

49

100

71

100

103

100 ]j

l/Fawn:Mother Ratio = Fawn:Doe Ratio recalculated after removing
estimated non-producing yearlings from the doe category.

the number of

2/

- Assumes 28.9 percent of the doe population were non-producing yearlings,
based on the average percent of yearling females in the doe harvest from
1967-1970.

The ratio of 49 bucks:lOO does estimated from the 1971 sample compares favorably with 42 bucks:lOO does and 43 bucks:lOO does estimated in 1970 and 1969,
respectively.
The ratio of 71 fawns:lOO does was not different from the 1970
ratio, but it was approximately 24'percent below the fawn:doe ratio of 1969.
The fetus:doe ratios for 1969, 1970, and 1971 (Table 5) supported the conclusion that the observed differences in pre-season fawn:doe ratios were
real (Gill 1972).
Post-season Counts--Post season classification data were obtained in two ways,
helicopter counts and ground counts.
Comparisons of these data for the years
1967-68 through 1971-72 are presented in Table 6. Chi-square analyses revealed no significant differences between helicopter and ground count estimates of fawn:doe ratios for any year.
Buck:doe ratios computed from helicopter
and ground data were significantly different in 1967-68 and 1971-72.
Because

�-174-

of the consistency of the helicopter classification data among years it
was concluded that bucks were under-sampled in the 1967-68 ground counts
and over-sampled in 1971-72. Buck:doe ratios did not change over the
five year period, but fawn:doe ratios showed a steady decline from 196768 through 1970-71 and then an upturn in 1971-72 (Fig. 1).

Table 5.

Comparisons

of gross and net fawn production

in Middle

Park,

1969-1971.

Year

Gross Fawn
Production 1/

Net Fawn
Production

1969

193

100

125

100

1970

164

100

112

100

1971

161

100

103

100 }j

1/
- Gross fawn production
collected does.

= fetuses

1/

per 100 does in samples of road-killed

and

2/
- Net favm production
fawns per 100 producing does based upon aerial
classifications of deer from helicopter surveys in October of each year.

3/
- 1971 net fawn production
the doe population

was calculated by assuming
were non-producing yearlings.

Table 6. Comparison of aerial and ground winter
Middle Park, 1968-1972.

Year

Bucks

1968

HelicoEter
Does

Surveys
Fawns

that 28.9 percent

classifications

of

of deer in

Ground Surveys
Does
Fawns

Total

Bucks

316

558
501
1,375
57 Bucks
100 Does
90 Fawns : 100 Does

147

396
363
906
37 Bucks
100 Does
92 Fawns : 100 Does

1969

261

490
418
1,169
53 Bucks : 100 Does
85 Fawns : 100 Does

319

582
505
1,406
55 Bucks : 100 Does
87 Fawns : 100 Does

1970

211

471
45 Bucks
77 Fawns

363
1,045
100 Does
100 Does

104

231
178
513
45 Bucks : 100 Does
77 Fawns : 100 Does

1971

------

No Counts

----------

356

1972

218

244
55 Bucks
100 Does
61 Fawns : 100 Does

783
321
1,460
45 Bucks : 100 Does
41 Bucks : 100 Does
390
261
927
71 Bucks
100 Does
67 Fawns : 100 Does

398

860

276

Total

�100
90
80
III
Q)

CIl

Cl

~
g
o ~
u

70

o

60

..

§
~ 50
o co
~

rz..

d
~
P:1
E-&lt;

~
0

Z

0

III

40

I

r-'
-...J
V1

~ j
~

I

30

20
10

o
1967-68

Fig. 1.

Trend in post-season

1968-69

1969-70

1970-71

1971-72

fawn: doe ratios obtained from ground count surveys, 1967-68 - 1971-72.

�-176-

Sex and Age Composition

of the Hunter Harvest

The 1971 deer season in Middle Park restricted hunters to taking antlered
deer.
For this and other reasons the harvest was one of the lowest recorded for Middle Park. Only 107 deer were checked during nine consecutive
checking days at Idaho Springs big game check station.
This was approximately 90 percent less than the total deer check of 1970 and nearly 73
percent lower than the 1970 buck check. The proportions of yearling bucks
and adult bucks are summarized in Table 7.

1/
1971.-

Table

7. Age classes of bucks harvested

GM
Unit

Adult
Males

Yearling
Males

Total

15

8
(50.0%)

8
(50.0%)

16
(l00.0%)

18

14
(70.0%)

6
(30.0%)

20
(100.0%)

27

16
(45.7%)

19
(54.3%)

35
(100.0%)

28

11
(9l.7%)

1
( 8.3%)

12
(100.0%)

37

13
(54.2%)

11
(45.8%)

24
(100.0%)

62
(57.9%)

45
(42.1%)

107
(100.0%)

Totals

in Middle

Park, October,

1/
- The 1971 deer season in Middle
antlered deer.

Park was restricted

to harvesting

only

�-177-

LITERATURE CITED

Gill, R. B. 1969. Middle Park deer study - population density and
structure. Colo. Div. Game, Fish and Parks. Game Res. Div.
Fed. Aid Proj. W-38-R-23. Game Res. Rep., July, Part 1. pp.10s122.
1971. Middle Park deer study - population density and structure.
Colo. Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. W-38R-2s. Game Res. Rep., July, Part 2. pp. 169-188.
1972. Productivity studies of mule deer in Middle Park, Colorado.
Mule Deer Workshop, Proc. 2: 8 pp.

Prepared by
R. Bruce Gill
Wildlife Researcher

��-179July, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.
Work Plan No.
Job Title

14
Middle

Period Covered:
Personnel:

Deer-Elk

W-38-R-26

October,

Investigations

Job No.

3

Park Deer Study - Productivity

and Mortality

1971 through August,

1972

R. Bruce Gill, Laren A. Roper, Paul Fo Gilbert, Gary L. Brown, Bruce
Sigler, David S. DeCalesta, Dan L. Baker, Lonnie M. Brown, Jan L.
Wassink and Len H. Carpenter.

ABSTRACT

Counts of corpora lutea of pregnancy in ovaries from does collected in Middle
Park averaged 1.95 CLP per doe, the second highest rate in the four year collection period.
Fetuses averaged 1.87 per doeo Fecundity rates were lowest for
yearling does, highest for 2 year-olds, followed in order by prime age does and
old does. All does examined were pregnant with 87.2 percent producing twins and
12 8 percent containing singletons. No triplets or quadruplets were found in the
1972 sample. Fetal sex ratios were nearly equal in 1972 (107 males:lOO females)
in contrast to the previous three years when they were unbalanced in favor of
males.
Mean breeding date for Middle Park does was estimated to be November 28
with the earliest breeding occurring on November 13 and the latest on January 110
Mean parturition date was June 17.
0

Over-winter mortality rates were estimated by searching 93 dead deer transects.
Only two deer were found for a projected mortality total of 50 deero Additionally,
101 deer died as a result of collisions with autos and trains and from collections.
The 1971 deer harvest totaled 374 bucks, the lowest total since harvest records
have been kept for Middle Park.

��-181-

MIDDLE PARK DEER STUDY
PRODUCTIVITY AND MORTALITY
R. Bruce Gill

P. S. OBJECTIVE
To estimate increments and losses to the Middle Park deer population
order to formulate more efficient harvest regulations.

in

SEGMENT OBJECTIVES

1.

Estimate

productivity

2.

Estimate
range.

mortality

3.

Estimate

the magnitude

rates of Middle

rates of Middle

See Gill

Park deer over the entire winter

of the Middle

METHODS

Park deer.

Park hunter harvest.

AND t-1ATERIALS

(1969) and Gill (1970) for details of methodology.

RESULTS

AND DISCUSSION

Productivity

Pre-natal

Productivity

Corpora Lutea of Pregnancy--Thirty-two
deer were collected by shooting
during the period January - April, 1972. In addition, eight does which
died from deer-auto collisions were examined.
Forty sets of mule deer
ovaries were sectioned and examined macroscopically
for counts of corpora
lutea of pregnancy (CLP). Overall the sample averaged 1.95 CLP per doe
for 1972. This figure exceeded the averages for 1970 and 1971 and was
only slightly below the 1969 high of 2.02 CLP per doe (Table 1). CLP were
found in at least one ovary from every doe examined yielding a conception
rate of 100 percent.
Right ovaries contained more CLP than left ovaries in
the 1972 sample, whereas in all other years left ovaries contained greater
numbers of CLP (Table 2).

�Table I. Comparisons of numbers of corpora lutea of pregnancy and fetuses occurring in samples of mule deer reproductive
tracts collected from Middle Park does, 1969-1972.
Yearling Does

Adult Does
No. of
Does
Examined

All Does
No. of
Does
Examined

No. of
Corpora
Fertilization
Does
Lutea Fetuses
Rate
Examined

Year

Fertilization
Corpora
Rate
Lutea Fetuses

1969

1.50

1.00

66.7%

2

2.05

1.97

96.1%

39

2.02

1.93

95.5%

41

1970

1.29

1.29

100.0%

7

1.86

1.7l

91.9%

35

1.76

1.64

93.2%

42

1971

1.27

1.09

85.8%

11

1.85

1. 75

94.6%

40

1.73

1.61

93.1%

51

1972

1.67

1.67

100.0%

6

2.00

1.91

92.6%

33.Y

1.95

1.87

93.6%

39 ~/

Corpora
Fertilization
Lutea Fetuses
Rate

I
f-'

00
N
I

Totals 1.38

1.27

91. 7%

26

1. 94

1.84

II
= 34 sets of ovaries were examined, 33 uteri were examined.
~I = 40 sets of ovaries were examined, 39 uteri we re examined.

21 = 148 sets of ovaries were examined, 147 uteri were examined.
!il

= 174 sets of ovaries were examined, 173 uteri were examined.

94.7%

147 ]j

1.86

1. 75

93.8%

173 !il

�-183-

Table 2. Number of corpora lutea of pregnancy (CLP) in right and left
ovaries collected from Middle Park deer, 1969-1972.

Year

CLP
Right Ovaries
No.
%

CLP
Left Ovaries
No.
%

Total
CLP
No.
%

1969

41

49.4

42

50.6

83

100.0

1970

34

45.3

41

54.7

75

100.0

1971

40

44.9

49

55.1

89

100.0

1972

44

56.4

34

43.6

78

100.0

All
Years

159

48.9

166

51.1

325

100.0

Accessory Corpora Lutea--The presence of accessory corpora lutea (CLA) has
been noted previously (Gill 1971). Accessory corpora lutea were found in
samples from all four collection years with 1972 samples containing the
greatest proportion relative to CLP and 1970 samples containing the least
(Table 3). CLA of the type encountered and tallied in this study apparently
arise when small follicles rupture about the same time as primary follicles
which ultimately develop into CLP. According to Thomas (1970), these
smaller follicles rarely shed viable ova and are most cornmon in younger and
older aged deer. The main distinction between CLP and CLA is the relatively
smaller size of CLA. CLA are usually less than one-quarter the size of the
largest CLP in the sample (Thomas 1970).
The significance, if any, of differences in proportions of CLA to CLP among
yearly samples is not known.
They do not appear to be consistently correlated
with the number of CLP per doe (Table 3).

Table 3. Accessory corpora lutea (CLA) and corpora
found in ovaries of Hiddle Park does, 1969-1972.

lutea of pregnancy

(CLP)

Year

No.
CLA

CLA per
Doe

No.
CLP

CLP per
Doe

Ratio
CLA/CLP

1969

5

0.12

83

.06

1

1970

2

0.05

75

2.02
1.76

.03

1

1971

7

0.13

89

1.73

.08

1

1972

7

0.18

74

1.95

.10

1

�-184-

Fetus Counts--Thirty-nine
does were examined in 1972 to estimate Middle
Park pre-natal fawn production.
All were pregnant, yielding 73 fetuses
for an overall fetus:doe ratio of 1.87 (Table 1). Yearlings averaged
1.67 fetuses per doe and adults (2+ years) averaged 1.91. The 1972
overall fetus:doe ratio was the second highest recorded in the four
year doe collection program.
Extrapolating from fetus data, 1972
should experience excellent fawn production approaching that of 1969.
The combined 1972 sample indicated that right and left uterine horns
contained approximately equal numbers of fetuses.
This was similar to
1969 while in 1970 a slightly greater proportion of the fetuses was found.
in the right uterine horn, and in 1971 the pattern was reversed (Table 4).

Table 4.
collected

Number of fetuses in right and left uterine horns from does
in Middle Park, 1969-1972.

Horn

Year

Fetuses
Right Uterine
No.

1969

41

50.6

40

49.4

81

100.0

1970

39

53.4

34

46.6

73

100.0

1971

38

46.9

43

53.1

81

100.0

1972

37

50.7

36

49.3

73

100.0

All
Years

155

50.3

153

49.7

308

100.0

%

Fetuses
Left Uterine Horn
No.
%

Total
Fetuses
No.
%

Age Specific Fecundity--Productivity
of does within each age class was estimated by pooling samples from all four collection years into composite tables.
Sample sizes were increased in this manner, but potential year differences
were masked.
Age-specific sample sizes were so small within individual
collection years that only very large year to year differences would have
been significant anyway, and so pooling was justified.
Perusal of individual age classes revealed no consistent patterns except that year
classes 1, 9, 11, and 12 appeared to have lower fetal rates than other year
classes (Table 5). When age classes 9 and older were grouped into a single
9+ category a general pattern was apparent. Yearlings had the lowest
fecundity, age classes 2 through 8 had highest fecundity rates, and age
class 9+ was intermediate between yearlings and ages 2-8 (Table 6). Grouping age classes into four broad categories [yearlings, 2 year-olds, prime
(3-7), and old (8+) Jas Robinette et al. (1955) did for Utah mule deer,
even more distinct patterns were apparent.
Yearlings produced fewest numbers
of fetuses per doe, followed in order of increasing productivity by the old
category, prime does and 2 year-olds (Table 7).

�-185-

Table 5. Age specific fecundity rates of Middle Park does collected
from 1969 to 1972.

Age at
Breeding

No. of
Does

No. of
eLP

eLP
per Doe

No. of
Fetuses

Fetuses
per Doe

Percent
of Does
Pregnant

1.5

26

36

1..38

33

1.27

92.3

2.5

18

38

2.11

38

2.11

100.0

3.5

26

51

1.96

48

1.85

100.0

4.5

21

41

1.95

41

1.95

100.0

5.5

16

31

1.94

28

1. 75

100.0

6.5

17

35

2.06

31

1. 82

94.1

7.5

8

15

1.88

15

1.88

100.0

8.5

11

20

1.82

20

1. 82

100.0

9.5

9

15

1.67

15

1.67

88.9

10.5

9

18

2.00

17

1.89

100.0

11.5

4

7

1. 75

6

1.50

100.0

12.5

2

4

2.00

2

1.00

100.0

13.5

5

10

2.00

9

1.80

100.0

Table 6. Age specific fecundity rates of Middle Park does with age classes
9.5+ grouped into a single age class.

No. of
eLP

eLP
per Doe

No. of
Fetuses

Fetuses
per Doe

Percent
of Does
Pregnant

26

36

1.38

33

1.27

92.3

2.5

18

38

2.11

38

2.11

100.0

3.5

26

51

1.96

48

1.85

100.0

4.5

21

41

1.95

41

1.95

100.0

5.5
6.5

16
17

31
35

1.94
2.06

28
31

1.75
1.82

100.0
94.1

7.5
8.5
9.5+

8

15
20
54

1.88
1.82
1.86

15
20
49

1.88

100.0

1.82
1.69

100.0
96.6

Age at
Breeding

No. of
Does

1.5

11

29

�-186-

Table 7. Fecundity rates of Middle Park does which have been grouped
into four broad age categories.

Age
Category

No. of
Does

No. of
CLP

CLP
per Doe

No. of
Fetuses

Fetuses
per Doe

Percent
of Does
Pregnant

Yearlings

26

36

1.38

33

1.27

92.3

Two year-olds

18

38

2.11

38

2.11

100.0

Prime (3.5-7.5)

88

173

1.97

163

1.85

98.9

Old (8.5+)

40

74

1.85

69

1.72

97.5

Litter Sizes and Fetal Sex Ratios

Litter Sizes--In 1972 no does were examined which contained more than two
fetuses. Additionally, no does were examined which did not contain at least
one fetus. The 1972 sample contained the highest percentage of twins (87.2%)
and the lowest proportion of singletons (12.8%) (Table 8). This increased
proportion of twins accompanied by a decreased percentage of singletons and
no does which were not pregnant was sufficient to maintain a high reproductive
rate similar to 1969 even though the 1972 sample lacked triplets or quad-.
ruplets.

Table 8.

Litter size frequencies of Middle Park does, 1969-1972.

Year

0

Litter Sizes
1
2

3

4

Total
Sample Sizl:':

1969

No. Litters
Percent

0
0.0

10
24.4

25
61.0

5
12.2

1
2.4

41
100.0

1970

No. Litters
Percent

5
11.9

7
16.7

28
66.7

2
4.7

0
0.0

42
100.0

1971

No. Litters
Percent

1
2.0

19
37.2

31
60.8

a

a

0.0

0.0

51
100.0

No. Litters
Percent

0
0.0

5
12.8

34
87.2

0
0.0

0.0

39
100.0

No. Litters
Percent

6
3.5

41
23.7

118
68.2

7
4.0

1
0.6

173
100.0

1972

All
Years

a

�-187-

Fetal Sex Ratios--Fetal sex ratios of the previous three years have been
unbalanced in favor of males (Fig. 1). In contrast, the 1972 sample
yielded approximately equal numbers of male and female fetuses (Table 9).
Consequently, the 1972 fawn crop should supply a greater proportion of
producers to future populations than any of the three previous years,
provided the 1972 sample adequately reflects the total Middle Park doe
population.

Conception

Dates and Parturition

Dates

Conception Dates--Conception
dates of Middle Park deer were calculated
by backdating from collection dates a number of days equivalent to each
estimated fetus age. Fetal ages were estimated from a mule deer growth
curve constructed by Hudson and Br owman (1959). Each fetus was aged
separately regardless of sex or litter size, and it was assumed that
differences between sexes or among individual fetuses within litters were
attributable to different conception dates.
In 1972, 64 fetuses were aged. The mean breeding date for this sample
was computed according to the method of Jackson and Hesselton (1971).
Mean breeding dates were calculated by starting from the date on which
the first fetus was conceived and counting consecutive days up to the
date the last was conceived.
Each day was multiplied by the number of
fetuses conceived on that day, and these figures were summed over all
days. This total was divided by the total number of fetuses to arrive
at the mean breeding date.
Mean breeding dates were:
1969 collections - November 28, 1968; ·1970
collections - December 1, 1969; 1971 collections - November 28, 1970;
and 1972 collections - November 24, 1971 (Fig. 2). No significant differences were found among breeding dates for the four years so all years were
combined and a pooled mean breeding date was calculated.
This date was
November 28, with the earliest conception occurring on November 13 and
the latest on January 11 (Fig. 3).

Parturition Dates--Parturition
dates were computed by adding 202 days to
each conception date. An average gestation period of 202 days was assumed
for Middle Park deer (Robinette and Gashwiler, 1950; Jackson and Hesselton,
1971). Computations were made only for the pooled sample. The mean date
of parturition for Hiddle Park was estimated to be June 17 with parturition
dates ranging from June 3 to August 1 (Fig. 4).

�-188-

o
II

= Males
= Females

Fig. 1.

Comparative proportions
Park does, 1969-1972.

of males and females in litters

of ?'~iddle

�-189-

40
1968-69
30
20
10
0
40
1969-70
30
(I)

z

0

20

H

t

f.:&lt;l

(.)

z

10

0

(.)

H

&lt;
E-t
0

E-t
I%..

0
40
1970-71

0
E-&lt;

~

30

u

&amp;1
n,
20
10
0
40
1971-72
30
20
10

o
NOV. NOV. NOV. NOV. DEC. DEC. DEC. DEC. DEC. DEC. JAN. JAN. JAN.
11-15 16-20 21-25 26=30 1-5
Fig. 2.

Histogram

6-10 11-15 16-20 21-25 26-31

1-5

6-10 11-15

of Miodle Park deer breeding dates for each of four sample years.

�-190-

~
o

40

~
~

)0

H
f:.-4

o

(.)

H

&amp;S 20
o

E-t

~

10

NOV. NOV. NOV~ NOV. DEC. DEC. DEC. DEC. DEC. DEC. JAN. JAN. JAN.
11-1§-16-20 21-25 26-)0

Fig.).

1-5

6-10 11-15 16-20 21-25 26-)1

1-5

6-10 11-15

Histogram of Middle Park deer breeding dates pooling data from

1969-1972.

�-191-

40_
CIl

::r:
E-;
p::
H

P=l

30 _

H
&lt;l!

8
0
8

20 _

r.x..
0
8

10 _

Z
1%1
U

~

A.

0_
JUNE

JUNE

1-5

6-10 11-15 16-20 21-25 26-30 1-5

Fig. 4.

JUN3

JUNE

JUNE

JUNE

JULY

JULY

Histogram of Middle Park deer parturition

1969-1972.

JULY

JULY

JULY

JULY

6-10 11-15 16-20 21-25 26-31

dates, pooling data from

A.UG.

1-5

�-192-

Table 9. Proportions of males and females
does collected from 1969 to 1972.

in litters of Middle

Males

Park

Females

Ratio of
: 100 Females

Year

No.

%

No.

%

1969

43

63.2

25

36.8

172

100

1970

47

67.1

23

32.9

204

100

1971

43

57.3

32

42.7

134

100

1972

29

51.8

27

48.2

107

100

All
Years

162

60.2

107

39.8

151

100

Males

Fawn Survival
Fawn survival rates were graphed from ratios of fawns per 100 producing does.
Four temporarily separate ratio calculations were used to graph changes:
estimates of the number of fawns conceived per 100 does were obtained from
road-killed deer and doe collections; fawns per 100 does at birth were
obtained from the same sources, fawns per 100 does prior to the October regular deer season were calculated from helicopter survey results, and fawns
per 100 does in January were computed from ground classification data.
Ratios obtained during fawn:doe counts in October and January were corrected
to fawn:producing doe ratios by subtracting the proportion of yearling
(non-producing) does from the total doe sample.
Estimates of the proportion
of yearlings in the doe population were obtained from check station data
concerning age structure of the doe harvest.
In 1971 only bucks were harvested in Middle Park so the yearling doe proportion was approximated by
pooling harvest data from 1967-70 and computing the average percent of
yearlings (Table 10).
Changes in survival rates over the 60 week period are presented in Figure
5. Fewer fawns were conceived and born in 1970 and 1971 than in 1969, but
survival rates were similar through 46 weeks in all three years.
Survival
from 46 to 58 weeks was similar in 1969 and 1971 but somewhat lower in

1970.

�-193-

Table 10.

Percent of yearlings

in the Middle

Park doe harvest,

1967-

1970.

Year

Yearlings
No.

1967

Adult

Totals

%

No.

%

No.

%

84

32.2

176

67.7

260

100.0

1968

58

24.7

177

75.3

235

100.0

1969

142

24.4

441

75.6

583

100.0

1970

136

36.4

238

63.6

374

100.0

All
Years

420

28.9

1,032

71.1

1,452

100.0

Mortality

Over-winter

Mortality

Estimated over-winter mortality of deer in 1971-72 was lowest of any year
since sampling was initiated in 1967-68. Only two dead deer of the year
(both old bucks) were found on 93 transects for a total mortality projection of 50 deer (Fig. 6). Subjective appraisal of the 1971-72 winter by
Area Supervisor Paul F. Gilbert indicated it was one of the mildest winters
in Middle Park since 1965-66 and little winter deer mortality was expected.
Low mortality combined with indicated high fawn production and survival
should result in a large increase in the 1973 deer winter population level.
Populations should number from 9,000 to 10,000 deer by January, 1973, provided the October deer harvest does not exceed 1,000.

Highway,

Railroad,

and Collection

Mortality

Deer losses resulting from deer-auto collisions were summarized from Wildlife Conservation Officer records.
Losses resulting from collisions with
trains were obtained from counts conducted by Colorado State University
wildlife management students in May, 1972. Totals from these two mortality
sources were 48 deer perishing from deer-auto collisions and 21 dying from
collisions with trains.
In addition, 32 does were collected to estimate
productivity (Table 11). Therefore, the total non-harvest mortality
(minimum) for Middle Park in 1971-72 was estimated at 151 deer.

�-194-

100

z
0
H
F-&lt;

1969

H

~
8

50

eJ

z

0...

0
Ul

e58

~

;.2~

Ul
I

~

~

0
0

10

20

3D

40

50

60

WEEKS FROM CONCEPTION
100

93.2%

Z

H

0

&lt;

H

~
:&gt;

H

f5
Ul

8

1970

g5
8
p::

0:

50

8

Z
0
Ul

~

~

,

U

p::
I%l
A.

Ul

&amp;i

~
0...

~
~

0
0
100

10

20

3D
93.1%

40

50 '

60

�1800

-I

1706

•

1600 1400 1200 -,

\

&amp;1

1135
~

w

Q

~

1000 -

0

~
w
~

Z

800 600

-.

v

-I

639
400 -

I
I-'
\.0

V1
I

I

200 01967-68

Fig. 6.

Over-winter

mortality

1968-69

of Midd1e Park deer,

1969-70

1967-68 through 1971-72.

1970-71

1971-72

�-196-

Table II. Summary of sources of recorded deer mortality other than overwinter mortality and harvest mortality for Middle Park, 1963-64 to 1971-72.

Railroad
Mortality

Trap Mortality
and
Collections

Total
(Minimum)

1963-64

90

0

90

1964-65

385

0

385

1965-66

7

0

7

Year

Highway
Mortality

1966-67

56

132

0

188

1967-68

54

60

0

114

1968-69

77

56

51

184

1969-70

67

38

40

145

1970-71

57

9

49

115

1971-72

48

21

32

101

Harvest Mortality
Three different methods were used to estimate harvest mortality (exclusive
of wounding loss) in Middle Park for the 1971 hunting season.
These methods
have been described previously (Gill 1969). In general, all three methods
have been in agreement with the regression equation method usually yielding
slightly higher estimates than the county report card and ratio-deer check
methods.
In 1971 deer hunters were restricted to taking only antlered deer
and deer and elk seasons were held at separate times. The result was the
lowest deer harvest recorded since 1947 when data were first available
(Table 12). Estimates of the 1971 harvest ranged from 348 deer (ratiodeer check method) to 516 deer (regression equation method) (Table 13).
The county report card ratio-deer check methods probably were more accurate
since the regression equation method projected beyond the range of data used
in its computation.

�-197-

Table 12.
Year

Summary of deer seasons and harvests in Middle Park, 1947-1971.
Type of Season

Deer
Harvest

1947

Either sex - 1 deer

2,639

1948

Either sex - 1 deer; 1st half season
Antlered only - 1 deer; 2nd half season

2,314

1949

Either sex - 1 deer

2,247

1950

Either sex - 1 deer; 1/2 area
Antlered only - 1 deer; 1/2 area

2,537

1951

Either sex - 1 deer

4,462

1952

Either sex - 1 deer

1,800

1953

Either sex - 1 deer

2,300

1954

Either sex - 1 deer; 1st half season
Either sex - multiple licenses; 2nd half

4,656

1955

Either sex - multiple licenses

3,322

1956

Either sex - 2 deer; 1st half season
Does only; 2nd half

4,009

1957

Multiple licenses - 2 deer per license

2,747

1958

Either sex - 1 deer

2,170

1959

Either sex - 1 deer

2,881

1960

Either sex - 1 deer

2,386

1961

Either sex - 1 deer

2,468

1962

Either sex - 2 deer

4,338

1963

Either sex - 2 deer

4,719

1964

Either sex - 2 deer

5,012

1965

Either sex - 2 deer

2,392

1966

Either sex - 2 deer

5,264

1967

Either sex - 2 deer

3,510

1968

Either sex - 1 deer; 1st half season
Either sex - 2 deer; 2nd half

2,104

1969

Either sex - 1 deer; 1st half season
Either sex - 2 deer, 2nd half

4,151

1970

Either sex - 1 deer

2,097

1971

Antlered only

374

�-198-

Table l3. Comparison of three different methods for estimating Middle
Park deer harvests, 1967-1971.
Regression
Equation
Estimate

Year

County Report
Card Estimate

Ratio-Deer
Check Estimate

1967

3,510

3,494

X2

161.95 + 3.348 Xl

3,066

1968

2,104

2,237

X2

321.41 + 3.140 Xl

2,909

1969

4,151

4,321

X2

52.00 + 3.329 Xl

4,503

1970

Not Available

2,097

X2

165.90 + 3.184 Xl

2,436

1971

374

348

X2

165.90 + 3.184 Xl

516

Regression
Equation

LITERATURE CITED
Gill, R. B. 1969. Middle Park deer study - productivity and mortality.
Colo. Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj.
W-38-R-23. Game Res. Rep. July, Part 1. pp. 105-122.
1970. Middle Park deer study - productivity and mortality. Colo.
Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. W-38-R-24.
Game Res. Rep. July, Part 3. pp. 337-354.
1971. Middle Park deer study - productivity and mortality. Colo.
Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. W-38-R-25.
Game Res. Rep. July, Part 2. pp. 189-207.
Hudson, P., and L. G. Browman. 1959. Embryonic and fetal development of
the mule deer. J. Wildl. Mgmt. 23(2):295-304.
Jackson, L. W., and W. T. Hesselton. 1971. Breeding and parturition dates
for various age classes of female white-tailed deer (Odocoileus
virginianus borealis) in New York. NE Sect. Wildl. Soc., Trans. 28:21-35.
Robinette, W. L., and J. S. Gashwiler. 1950. Breeding season, productivity,
and fawning period of the mule deer in Utah. J. Wildl. Mgmt. 14(4):
457-469.
Robinette, W. L., J. S. Gashwiler, D. A. Jones, and H. S. Crane. 1955.
Fertility of mule deer in Utah. J. Wildl. Mgmt. 19(1):115-136.
Thomas, D. C. 1970. The ovary, reproduction, and productivity of the female
Columbian black-tailed deer. PhD Thesis. Univ. British Columbia,

prepa:::c:~ver~2~~~.
~
R. Bruce Gill
Wildlife Researcher

�-199-

July, 1972

JOB PROGRESS REPORT

State of

~C~O~L~O~RA~D~O~

Work Plan No.

Investigations
4

Job No.

14

Middle Park Deer Study - Physical

Period Covered:
Personnel:

Deer-Elk

W-38-R-26

Project No.

Job Title

_

Characteristics

and Food Habits

April 1, 1971 through March 31, 1972

R. Bruce Gill, William Adrian,
Sigler and Laren Ao Roper.

David DeCalesta,

Robert Keiss, Bruce

ABSTRACT

Kidney fat and bone marrow fat indices were estimated from mature doe mule deer
collected and accidently killed in Middle Park from 1969-1972.
Findings are
presented in tables. Analyses will be reported in the final report due this
segment.
Blood counts and plasma analyses will also be included in the report.
In the 1971 buck only deer hunt, 16 right antlers were collected for comparisons
with antlers taken from deer killed in Middle Park in 1969 and 1970. Weights and
numbers of points are shown
0

��-201-

PHYSICAL

MIDDLE PARK DEER STUDY
CHARACTERISTICS AND FOOD HABITS
Laren A. Roper

P. S. OBJECTIVE
To measure selected physical characteristics related to the assessment
of physical condition of deer and to determine forage preferences of
mule deer in Middle Park.

SEGHENT OBJECTIVES
1.

Heasure selected
to be indicative

2.

Estimate

physical characteristics
of the state of physical

forage selections

HETHODS

Forage Preferences
Procedures

are reported by Gill

Physical

of deer which are believed
condition of deer.

of wild deer from stomach

content analysis.

AND MATERIALS

- Stomach Content Analyses

(1969).

Characteristics

All procedures are reported by Gill (1969) and Roper (1970 and 1971) except
for those initiated during this segment for collecting blood samples and
analyzing blood plasma.
Approximately 40 ml of blood was taken directly
from the heart as soon after death as possible and placed into tubes containing .20 grams of sodium heprin.
These samples were centrifuged for 10
minutes at 30,000 RPH and the plasma was drawn off with a pipet.
The
procedure for analyzing these samples will be detailed in the final report
as the analysis is yet to be completed.

RESULTS AND DISCUSSION

Forage Preferences

- Stomach Content Analyses

Samples from rumen contents were taken from 32 collected
will be analyzed at a later date.

deer.

The data

�-202-

Condition

Kidney

Fat and Bone Marrow

Indices

Fat

Deer were collected each winter commencing in mid January and continuing
through April of 1969, 1970, 1971 and 1972. In 1969, 1970 and 1971 one
deer was collected in May. Kidney fat and bone marrow fat estimates were
obtained from road killed deer and some deer accidently killed during
trapping operations whenever this was feasible.
In preparation for the
final report which will be forthcoming this segment all data for kidney
fat estimates have been checked for accuracy and are summarized in Tables
1-7. Ages are shown as determined by counts of dental cementum annuli
(Erickson and Seliger 1969).
Starting January 1 each day was numbered
consecutively throughout the collection period.

Blood Counts and Plasma Analyses
Blood smears were taken from each collected deer.
Counts of segmented
neutrophils, monocytes, lymphocytes and eosinophils are being taken. The
data will be presented in the final report.
Blood plasma from a majority of the collected deer will be analyzed for
blood urea nitrogen, glucose, ornithine carbamyl transferase, volatile
fatty acids, glucocorticoides,
lactic dehydrogenase ratios, aldolase
and ketone bodies.
These data will be compared with similar samples
drawn from experimental deer that were starved and refed in conjunction
with Job 6.

Antler Weights

and Number of Points in Yearling

Bucks

During the 1971 deer season the right antler was collected from yearling
bucks.
These antlers were weighed and tabulated according to the number
of points (Table 8). Due to the bucks only deer regulation the total
harvest was very low. As a result our sample size is much smaller than
previous years.
Within
points

the sample of antlers 28 percent were spikes, 55 percent had 2
and 17 percent 3 points.

�-203-

Table 1. Physical condition indices for mature doe deer collected in
Middle Park in 1969.

Deer
No.

Collection
Date

Day
No.

Right
Area ~/ Kidney

69-1
69-2
69-3
69-4
69-5
69-6
69-7
69-8
69-9
69-10
69-11
69-12
69-13
69-14
69-15
69-16
69-17
69-18
69-19
69-20
69-21
69-22
69-23
69-24
69-25
69-26
69-27
69-28
69-29
69-30
69-31
69-32
69-33

1-15
1-16
1-17
1-21
1-21
1-22
1-23
1-23
2-4
2-5
2-6
2-7
2-7
2-17
2-18
2-18
3-4
3-5
3-6
3-6
3-7
3-13
3-20
3-20
4-8
4-9
4-10
4-10
4-10
4-14
4-14
4-15
5-17

15
16
17
21
21
22
23
23
35
36
37
38
38
38
39
39
63
64
65
65
66
72
79
79
98
99
100
100
100
101
101
102
137

M
M
B
T
T
W

w
B
B
B
T
T

w
M

M
T
M
W
T
T
M

w
B
B
T
M

B
B
T
W

w
M

M

11.9
9.6
59.7
38.8
44.0
30.3
32.8
30.8
22.5
46.6
39.5
42.8
32.1
25.7
30.3
9.8
10.2
26.2
10.8
46.0
26.0
14.0
18.8
16.0
5.3
14.0
9.0
20.6
20.7
7.0
4.3

Left
Kidney

6.7
79.8
29.6
23.8
32.0
26.9
30.3
25.8
48.5
49.6
33.0
31.9
30.4
22.7
7.9
14.5
19.5
8.0
37.7
23.9
10.4
16.3
11.1
8.8
7.5
9.0
4.7
20.6
10.7
7.4
3.8

Percent Fat
Mean
Kidney
Bone
Fat
Marrow

8.1
69.8
34.2
33.9
31.1
29.9
30.5
24.2
47.6
44.6
37.9
32.0
28.0
26.5
8.9
12.3
22.9
9.4
41.9
25.0
12.1
17.6
13.5
6.4
11.5
6.8
20.6
15.7
7.2
4.0

85
92
90
86
84
87
91
84
91
94
91
89
92
9
91
85
88
93
92
84
91
87
56
60
88
78
84
89
51
11

Age By
Dental
Cementum

8
9+
5
2
1
1
2
13
8
2
Mature
8
2
6
6
3
7
3
7
5
2
5
6
4
2
2
5
3
5
4
6
3
7

Pooled Mean Percent Kidney Fat = 32.5
1/
- Areas: M = Muddy Creek; B = Blue River; W = Williams Fork River; T =
Troublesome Creek.

�-204-

Table 2. Physical condition indices for mature doe deer collected in
Middle Park in 1970.

Deer
No.

Collection
Date

Day
No.

70-1
70-2
70-3
70-4
70-5
70-6
70-7
70-8
70-9
70-10
70-11
70-12
70-13
70-14
70-15
70-16
70-17
70-18
70-19
70-20
70-21
70-22
70-23
70-24
70-25
70-26
70-27
70-28
70-29
70-30
70-31
70-32
70-33

1-23
1-26
1-26
1-27
1-29
2-5
2-9
2-9
2-10
2-12
2-12
2-13
2-25
2-26
2-27
2-28
3-12
3-19
3-20
3-23
3-23
3-23
3-25
3-27
4-7
4-7
4-9
4-13
4-13
4-14
4-14
4-15
5-23

23
26
26
27
29
36
40
40
41
43
43
44
56
57
58
59
71
78
79
82
82
82
84
86
97
97
99
103
103
104
104
105
143

1 Right
Area _I Kidney

T
T
T
M
M
B
B
B
W
W
W
M
M
B
W
T
M
T
M

W
W
B
B
T
M
M
B
T
T
W
W
B
B

17.6
22.8
40.4
26.9
32.7
23.0
38.5
23.8
28.8
.04
20.0
14.9
10.5
13.7
41.4
28.9
15.2
5.6
17.9
4.6
14.4
7.9
3.7
7.2
14.2
9.2
4.8
6.4
.03
9.5

Left
Kidney

Percent Fat
Mean
Kidney
Bone
Fat
Marrow

26.2
18.8
51.5

21.9
20.8
45.9

23.8
31.0
41.0
20.2
29.3
.04
20.0
15.2
5.1
15.2
58.0
17.7
14.1
5.5

28.2
27.0
39.7
22.0
29.0
.04
20.0
15.0
7.8
14.4
49.7
23.3
14.6
5.5

6.8
18.4
6.7
2.5
7.4
21.8
6.6
7.5
10.5
7.3
.02
11.4

5.7
16.4
7.3
3.1
7.3
18.0
7.9
6.1
6.8
.02
10.4

Age By
Dental
Cementum

90
89
76
84
91
91
95
92
74
3
85
89
53
86
88
81
85
66
86
66
79
61
33
82
85
30
72
33
40
2
72

Pooled Mean Percent Kidney Fat
l.lAreas: M = Muddy Creek; B = Blue River; W = Williams Fork River; T =
Troublesome Creek.

13
12+
5
1
3
6
10+
2
4
10
6
1
1
8
10
3
6
6
10+
6
4
3
3
4
3+
3+
3+
3+
3
4
3
3+
2
30.3

�-205-

Table 3. Physical condition indices for mature doe deer collected in
Middle Park in 1971.

Deer
No.

Collection
Date

Day
No.

71-1
71-2
71-3
71-4
71-5
71-6
71-7
71-8
71-9
71-10
71-11
71-12
71-13
71-14
71-15
71-16
71-17
71-18
71-19
71-20
71-21
71-22
71-23
71-24
71-25
71-26
71-27
71-28
71-29
71-30
71-31
71-32
71-33

1-16
1-19
1-22
1-22
1-25
1-26
1-27
1-28
1-28
2-5
2-9
2-10
2-16
2-17
2-23
2-26
3-4
3-8
3-11
3-16
3-19
3-22
3-26
3-30
4-1
4-5
4-7
4-12
4-19
4-21
4-23
4-26
5-21

16
19
22
22
25
26
27
28
28
36
40
41
47
48
54
57
64
68
71
76
79
82
86
90
92
96
98
103
110
112
114
117
142

Right
Area 1/ Kidney

Left
Kidney

22.5
15.8

15.3
14.3
19.4
23.3
5.2
17.4
19.5
15.6
30.4
19.9
16.5
26.9
32.0
26.3
26.4
45.3
17.5
17.6
15.0
16.2
9.5
26.5
7.6
16.5
20.1
5.8
10.7
5.2
16.7
18.4
10.5
8.2
8.2

M
W
T
B
M
W
T
B
B
M
B
T
H

W
T
W
M
W
T
B
H

W
T
B
M
T
W
B
M
T
W
B
M

23.0
8.2
18.9
27.4
23.8
38.9
23.6
27.3
24.7
34.1
24.1
37.5
37.8
22.7
21.1
17.3
19.1
10.4
20.2
6.0
19.3
17.7
17.4
5.8
12.8
12.1
10.8
9.3
9.2

Percent Fat
Mean
Kidney
Bone
Marrow
Fat

18.9
15.0
23.1
6.6
18.1
23.4
19.7
34.7
21.8
21.9
25.8
33.0
25.2
32.0
41.6
20.1
19.4
16.1
17.7
10.0
23.4
6.8
17.9
18.9
14.0
5.5
14.8
15.2
10.2
8.8
8.7

Age By
Dental
Cementum

80.1
92.7
85.1
89.6
89.2
91.3
85.1
75.7
89.1
82.2
80.8

1
Mature
6
3-4
8
1
1
11
6
5
3
3
1
2
13
5
3
5
1
7
8
3
9
4
4
4
7
10
12
6
1
3
9-10

91.3
83.2
88.8
94.0
88.6
91.1
87.9
94.4
75.3
74.4
70.6
85.2
78.6
65.1
70.1
22.7
80.0
30.6
89.6
25.4
66.4

Pooled Mean Percent Kidney Fat

18.0

1/
- Areas: M = Muddy Creek; B = Blue River; W = Williams Fork River; T =
Troublesome Creek.

�-206-

Table 4. Physical condition indices for mature doe deer collected
in
Middle Park in 1972.

Deer
No.

Collection
Date

Day
No.

72-1
72-2
72-3
72-4
72-5
72-6
72-7
72-8
72-9
72-10
72-11
72-14
72-13
72-14
72-15
72-16
72-17
72-18
72-19
72-20
72-21
72-22
72-23
72-24
72-25
72-26
72-27
72-28
72-29
72-30
72-31
72-32

1-14
1-14
1-15
1-15
1-20
1-21
1-24
2-1
2-1
2-3
2-8
2-10
2-15
2-17
2-22
2-28
3-1
3-1
3-7
3-8
3-14
3-16
3-24
3-24
4-3
4-11
4-11
4-19
4-19
4-20
4-20
4-21

14
14
15
15
20
21
24
32
32
35
39
41
46
48
53
59
60
66
66
67
73
75
83
83
93
101
101
109
109
110

Right
Area 1/ Kidney
M
M
T
T
W
B
W
B
M
T
W
B

M
T
W
B
M
T
W
B
M
T
W
B
M
T
T
W
W
B

III

B

112

M

45.8
12.6
52.9

Left
Kidney

Percent Fat
Mean
Kidney
Bone
Fat
Marrow

31.1
41.5
43.6

48.7
17.4
44.8
36.8
24.4
28.8
45.2
21.8
38.4
41.1

28.1
40.0
42.4

20.1
39.2
31.3
23.1
22.5
34.8
21.6
19.5
26.8
24.8
31.2
40.5
28.9
18.7
9.1
21.4
13.1
17 .9
8.0
16.9
7.0

20.4
39.2
22.5
22.7
23.7
28.2
16.7
13.5
32.22
23.7
21.0
27 .1
21.9
12.1
6.6
16.4
13.2
14.8
7.5
12.9
10.1

20.2
39.2
26.9
23.0
23.1
30.6
19.1
16.5
29.5
24.3
24.9
33.8
25.4
15.4
7.9
18.9
13.1
15.4
7.9
15.2
8.5

25.4
39.6

47.2
15.0
48.8
24.9
34.2

92.7
83.4
81.7
86.4
79.4
87.5
86.4
88.0
84.0
76.4
86.2
68.7
89.1
87.0
90.6
90.9
82.5
86.3
67.5
88.7
78.8
90.0
78.8
82.4
82.8
50.7
72.5
73.2
80.3
50.3
86.0
67.3

Age By
Dental
Cementum
11
10
9
3
1
4
17
2
8
8
5
9
4
6
10
6
3
5
1
2
9
3
9
4
2
1
4
1
4
8
7
9

Pooled Mean Percent Kidney Fat = 32.6
1/
-Areas: M = Muddy Creek; B = Blue River; W - Williams Fork River; T =
Troublesome Creek.

�-207-

Table 5. Physical condition indices for hiway and railroad killed mature
doe deer in Middle Park in 1969.

Deer
No.

Collection
Date

69-l-RK
69-4-RK
69-5-RK
69-6-RK 2.../
69-7-RK
69-8-RK
69-l0-PlZ
69-11-RK
69-l2-RK
69-l4-RK
69-l5-RK

1-16
2-3
2-5
2-13
2-11
2-28
3-2
3-3
3-4
3-17
3-17

Day
1\10.

Right
Area }j Kidney

Left
Kidney

16
34
35
44
42
59
62
62
63
76
76

M
T
T
M
M
T
M
T
T
T
M

21.2
36.3

1/
- Areas: M = Muddy Creek; B
Troublesome Creek.

10.8
35.1
21.6
.1
20.0
10.0
26.5
19.0
36.9
48.8

Percent Fat
Mean
Kidney
Bone
Marrow
Fat

Age By
Dental
Cementum

16.0
35.7
88

.1
52.0
16.7
4.8
14.6
11.6
21.9
45.5

.1
18.9
5.4
20.6
15.3
29.4
47.1

43
74.
51
88
67

1
1
1
3

Blue River; W = Williams Fork River; T

2/
- Trap mortality.

Table 6. Physical condition indices for hiway and railroad and accidentally
killed mature doe deer in Middle Park in 1971.

Deer
No.

Collection
Date

7l-2-RK
71-3-RK
71-l-A Jj
71-5-RK
71-5-A JJ
7l-6-A Jj
71-7-A
71-9-RK
71-l0-RK
71-S-A Jj
71-l2-ARK
71-11-A Jj
71-l4-A
71-15-A ]j

u

u

71-16-A 1/

71-ll-RK
7l-l2-RK

1-27
2-1
2-4
2-9
2-22
2-22
2-26
3-4
3-4
3-4
3-11
3-11
3-11
3-19
3-19
5-18
5-20

Percent Fat
Mean
Kidney
Bone
Fat
Marrow

Day
No.

Right
Area 1/ Kidney

Left
Kidney

27
32
35
40
53
53
57
64
64
64
71
71
71
79
79
139
140

M
B
M
T

27.7
25.7
7.5

25 .t.

H
M
M

l2.S
3.0
5.2

23.6
24.1
15.6
8.7
11.5
2.S
5.9

B
B
T

35.1

15.6

25.3

24.9
11.5
12.1
2.9
5.5

M

T
H

M
M
T
T

9.6

5.1

7.3

90.4
83.4
91.3
S.O
39.0
61.2
90.0
78.5
84.4
63.0
2.6
8.7
18.3
44.5
71.8

1/ Areas: M = Muddy Creek; B = Blue River; W = Williams Fork River; T
Troublesome Creek.
~/ Trap mortality.

Age By
Dental
Cementum
5
9
1
1
7
11
1
8
1
3
6
1
4
6
1
4
1

�-208-

Table 7. Physical condition indices for hiway and railroad killed mature
doe deer in Middle Park in 1972.

Deer
No.

Collection
Date

72-1-RK
72-2-RK
72-4-RK
72-5-RK
72-6-RK
72-7-RK
72-8-RK
72-9-RK
72-10-RK
72-11-RK

1-2
1-8
1-12
1-12
1-20
1-20
2-8
2-12
3-30
4-13

Day
No.

Right
Area 1/ Kidney

2
8
12
12
20
20
39
43
89
103

M
B
T
B

Left
Kidney

72.4

52.8

Percent Fat
Mean
Kidney
Bone
Fat
Marrow

Age By
Dental
Cementum

1
3
1
4
2
7
4
1
1
3

62.6

B
M
T
B

82.2
83.2
75.8
81.2

1/
- Areas: M = Muddy Creek; B - Blue River; W = Williams Fork River; T
Troublesome Creek.

Table 8. Right antler weights of yearling bucks taken in Middle Park and
checked at the Idaho Springs Check Station in 1971.

Number of Points

CIl

§
!-I

1

2

3

21.3

52.1

87.9

26.4
48.1

59.9
75.6

107.0
127.7

52.0

85.3

74.1

92.1

c.!l

I=l
OM
.j.J

..c:

eo

OM
(iJ

!3

92.2
101.1
109.7
116.4
l32.5

N
Mean

18
81.2

4

�-209-

LITERATIJRE CITED

Gill, R. B. 1969. Middle Park deer study - physical characteristics
and food habits.
Colo. Div. Game, Fish and Parks.
Game Res. Div.
Fed. Aid Proj. W-38-R.
Game Res. Rept., July.
Roper, L. A. 1970. Middle Park deer study - physical characteristics
and food habits.
Colo. Div. Game, Fish and Parks.
Game Res. Div.
Fed. Aid Proj. W-38-R.
Game Res. Rept., July.

1971. Middle Park deer study - physical characteristics and
food habits.
Colo. Div. Game, Fish and Parks.
Game Res. Div.
Fed. Aid Proj. W-38-R.
Game Res. Rept., July.
Erickson, J. A., and W. G. Seliger.
1969. Efficient sectioning of
incisors for estimating ages of mule deer.
J. Wildl. Mgmt.
33(2):384-388.

Laren A. Roper
Asst. Wildlife Researcher

��-211July, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-26

Work Plan No.
Job Title

Job No.

Investigations
5

Middle Park Deer Study - Range Fertilization

Period Covered:
Personnel:

14

Deer-Elk

June 15, 1971 through March

15, 1972

R. Bruce Gill, Paul Fo Gilbert, Laren Ao Roper, O. C. Wallmo,
Don W. Reichert, Wayne L. Regelin, Lonnie M. Brown, Dan L. Baker,
Richard W. Hoffman and Len Ho Carpenter.

ABSTRACT

Second year fertilizer and herbicide treatment response data were gathered
on three study areas in western Grand County, Colorado.
The herbage yield
data showed a significant linear response to increasing increments of
nitrogen fertilizer.
The main effect of fertilizer was realized at the 30
pound nitrogen level. Significant responses in vegetation yield were also
obtained with the herbicide and fertilizer treatments, where forb production
was decreased and grass production increased.
Adequate sampling was obtained
in most cases for a stated degree of precision of total vegetation yield.
Six tame, trained mule deer were grazed at each study area during the winter
months.
A definite increase in grass and a corresponding decrease in forbs
in the diet was noted from the pretreatment period to the second year posttreatment period.
Treatment preference data at each study area revealed no
preferences by the tame deer for the control or fertilizer only treatments over
Additional fertilizer response information on
the combination treatmentso
a fourth study area is presented.

��-213-

MIDDLE

PARK DEER STUDY - RANGE FERTILIZATION
Len H. Carpenter

P. S. OBJECTIVE
To test the effects of fertilizers
tensity responses to fertilization

on deer forage and deer feeding inof critical winter range areas.

SEGMENT OBJECTIVES
1.

To test the effects of nitrogen and 2,4-D on forage production
composition on the selected study areas.

2.

To evaluate

3.

To evaluate the quality of big sagebrush
influenced by fertilizer treatment.

deer foraging

responses

METHODS

and

to treated areas.
as a mule deer food as

AND MATERIALS

Methods and materials employed in this segment have been detailed previously by Carpenter (1970) and (1971). Certain additions and changes
have been made.

Vegetation

Measurements

Herbage Yield
No changes made from 1971.

Botanical

Composition

No changes made from 1971.

Sagebrush

Samples

A separate study area was established to evaluate the influence of fertilizer upon the essential oil content and composition and the digestibility
of big sagebrush.
This study area is in critical mule deer winter range
located on Junction Butte Mountain, 3 miles southwest of Kremmling,
Colorado.

�-214-

Ammonium nitrate fertilizer, at the rate of 120 pounds of elemental
nitrogen per acre, was applied by hand on March 17, 1971. The fertilizer was applied in three strips 330 feet long by 176 feet wide with
a 66 foot control strip between each treatment.
The stripping procedure was done so that paired samples could be obtained from treated
and untreated adjacent strips.
Thirty 200 gram samples of the terminal-inch of big sagebrush stem,
plus leaf material, was collected on February 11 and 12, 1972. These
thirty samples were obtained from fifteen paired plots.
The fifteen
paired plots were made up of five lines with three sets of paired plots
each. The location of each paired sample was chosen randomly.
Once
the random location was chosen the plots on the treated and untreated
areas were paired as nearly as possible regarding soil conditions,
topography, and sagebrush plant size. The 200 grams of plant material
were taken from an average of ten sagebrush plants and from all vertical and lateral locations on those plants.
Plant samples were immediately
placed in plastic bags and frozen for future analysis.
In a separate experiment, common epsom salts (magnesium sulfate - MgS04)
were applied to plots on the same Junction Butte study area. Fifty
pounds of magnesium per acre were applied on March 31, 1972 to six
randomly selected square chain plots.
Three of the plots were located
on strips previously treated with nitrogen and three were located on
untreated or control strips.
Magnesium was chosen because it is present
in the chlorophyll molecule and is important in the photosynthetic process.
It was thought the presence of additional magnesium elements
would result in more efficient plant photosynthesis, producing fewer
essential oils which are by-products of plant metabolism.
Some justification for this hypothesis was found in Powell's (1968) work where
he found a significant inverse relationship between magnesium content
of the soil and volatile oil content in sagebrush.
Sagebrush samples
from these magnesium treatments have not yet been collected.
They will
be obtained in December or January in a manner similar to that described
above.
To determine oil content 100 grams of plant material will be taken from
each frozen sample and placed in a steam distillation apparatus and
distilled for 6 hours.
Previous studies (Ward and Nagy 1966) have
demonstrated that 95 percent of the oils are obtained during this period.
Total oil content for each sample will be determined on a dry weight
basis.
At the end of the distillation process, oils will be drained
into glass vials and stored in a refrigerator.
Analytical determination of the specific compounds present in each of
the 30 volatile oil samples will be done in a Varian Aerograph Model
1740 chromatograph with hydrogen flame detector.
Column oven temperatures
will be linear programmed from 600 to 2300 C. at 4°C./min.
Nitrogen gas
glow will be 28 ml/min.
A 10 foot by 1/8 inch column packed with 5 percent polyetholene glycol (PEG) on chromosorb "w" with a mesh of 80-120
will be used. A varian aerograph model 20 recorder and model 477 integrator

�-215-

with attached digital printer will be connected to the chromotograph.
The digital printout value for each peak will be divided by the total
for the sample to express the amount of each terpenoid as a percentage
of the total. Tentative identification of the individual terpenoids
will be done by the relative retention time. These times will be
compared with previous times of terpenoids identified by Ward and Nagy
(1966) and Paintner (1971).

Digestion

Pr9cedures

If the above analyses indicate that differences exist between treated
and untreated samples in volatile oil content or composition, tests
will be made to determine if these differences affect certain digestion
parameters.
The interest of this study lies in the effect of nitrogen fertilizer on
the food values of big sagebrush to mule deer during the winter period,
when their diet has a higher fiber content.
At this time the role of
cellulose digesting bacteria becomes most important.
As a result, some
measure of the comparative antibacterial effects of treated and untreated
volatile oils on rumen cellulo1ytics in vitro will provide information on
the comparative food value of treated and untreated big sagebrush.
Measuring in vitro cellulolytic activity requires a Rumen Fluid Cellulose
Broth to culture rumen organisms.
Composition of this broth is described
by Bryant and Burkey (1953) and Bryant and Robinson (1961) with modifications described by Ward and Nagy (1966). The rumen fluid to be incorporated into the media will be obtained from a rumen fistulated goat
or a tame mule deer receiving an ad libitum diet of concentrate and alfalfa
hay.
Total volatile oils from three controls and three treatments will be
tested for antibacterial action on rumen bacteria (Paintner 1971).
Commercially obtained individual terpenoids that may differ (if any do)
from the treated and untreated samples will also be tested.
Total volatile
oils and selected terpenoids will be dispensed into test tubes by means of
a microliter syringe in amounts of 0, 3, 6, 12, and 15 microliters for
culturing with RFCB. According to Ward and Nagy (1966) these levels should
demonstrate a definite antibacterial effect.
The culturing broth will be pipetted in 9 ml amounts into test tubes containing the volatile oils or selected terpenoids.
The broth will be
constantly stirred during dispension and will be maintained at a pH of
6.8. '
Rumen inoculum will be taken from tame deer by stomach pumping.
These deer
will be maintained on a known concentrate diet. Rumen contents will be
immediately taken to the laboratory, strained through cheesecloth, and 1
ml will be placed in an aerobic diluting fluid.
Composition of this fluid

�-216-

is described b Bryant and Burkey (1953). Rumen inoculum will be
diluted to 10- by this aerobic solution.
This diluted inoculum will
then be dispensed in 1 ml amounts to test tubes containing the 9 ml
of broth.
Duplicates will be run on all oil and terpenoid levels.

3

Cultures will be incubated at 390 C. for three days. After this period, the remaining cellulose in each tube will be weighed to determine
percent of cellulose digested.
Weights from each test tube culture
duplicates will be averaged and subtracted from the mean of the control.
This value will be expressed as a percentage of cellulose digested.

Deer Grazing Measurements
No changes made from 1971.

RESULTS

AND DISCUSSION

Vegetation

Herbage

Measurements

Yield

Methods for obtaining total herbage yield estimates are detailed by
Carpenter (1971). Yearly comparisons from 1969 to 1971 for the various
weight components at each study area are presented in Tables 1, 2, and 3.
First year vegetation responses to 120 pounds of nitrogen per acre applied
to the 4.6 acre sagebrush volatile oil study area are summarized in Table
4. It is interesting to note that the nitrogen treatment resulted in an
81 percent production increase in total leafy material over the untreated
areas.
Figures 1 and 2 show average forb and grass production by treatment for the 1971 data for all three study areas combined.
Definite decreases in forb production are shown in Figure 1 on combination
treatments compared to fertilizer only treatments.
Mean forb production
for all three study areas for all fertilizer only treatments is 136 pounds
per acre. Mean forb production for all three study areas for all combination treatments is 84 pounds per acre. Figure 2 shows the increase in
grass production of the combination treatments as compared to the fertilizer only treatments.
Mean grass production for all three study areas
for all fertilizer only treatments is 370 pounds per acre, while mean
production of grass for combination treatments is 641 pounds per acre.

�-217-

Table 1. Pre-treatment, first and second year herbage production response
data for Junction Butte study area. Values are in pounds per acre on a
dry weight basis.
(Lbs. Per Acre)
Treatment

Year and Weight Com:eonent
1971
1969
1970
1971
1969
1970
Shrub Current Growth Weight
Total Leafy Vegetation

o lb. N. 0 lb. 2,4-D
30 N
"

404
236

582
739

614
713

649
443

905
1099

792
961

464

957

853

723

1316

1074

90 N

"
"

392

940

991

637

1278

1231

120 N

"

477

1254

1102

732

1603

1346

249

365

388

454

709

782

505
625
492

461
675
585

905
1001
781

890
1069
902

637

744

915

1077

60 N

o N 2 lb. 2,4-D
30 N
60 N
90 N

"

251

"
"

427
337

530
624
424

120 N

"

485

549

Table 2. Pre-treatment, first and second year herbage production response
data for F1ight1ine study area. Values are in pounds per acre on a dry
weight basis.
(Lbs , Per Acre)
Treatment

o lb. N. 0 lb. 2,4-D
30 N
"
60 N
"
90 N
"
120 N

"

o N 2 lb. 2,4-D

Year and Y-JeightCom:eonent
1970
1971
1971
1969
1969
1970
Total Leafy Vegetation
Shrub Current Growth Weight

467

606

480

1177

1519

1078

419

497

506

1187

1399

1172

424

861

501

1134

1852

1097

704

741

700

1526

1704

1328

822

1625

1043

1620

2633

1773

633

393

227

1463

1212

1034

30 N

"

696

412

265

1463

1222

1110

60 N

"

802

582

489

1593

1437

1369

90 N

"

404

290

341

1147

1059

1256

120 N

"

398

357

373

1090

1162

1209

�-218-

Table 3. Pre-treatment, first and second year herbage production response
data for Corral Creek study area. Values are in pounds per acre on a dry
weight basis.

(Lbs. Per Acre)
Treatment

Year and Weight ComEonent
1969
1970
1971
1969
1970
1971
Shrub Current Growth Weight
Total Leafy Vegetation

o lb. N. 0 lb. 2,4-D

435

446

477

1214

1143

1148

30 N

"

722

713

687

1494

1512

1385

60 N

"

513

651

846

1274

1416

1576

90 N

"

632

747

846

1404

1577

1558

120 N

"

422

562

812

1173

1332

1538

589

188

247

1398

990

1040

o N 2 lb. 2,4-D
30 N

"

600

233

353

1374

1034

1323

60 N

"

331

225

270

1115

1082

1172

90 N

"

414

227

190

1173

1080

1093

120 N

"

297

225

310

1077

1107

1247

Table 4. First year herbage production response data for the volatile oil
study area. Values are in pounds per acre on a dry weight basis.

Treatment

Weight ComEonent
Shrub Current Growth Weight
Total Leafy Vegetation

No nitrogen
(Control)

378

730

120 1bs. N per acre

964

1324

�-219-

rl
0

b~
0

0

0
I

c-

0
I

I

'-0

0

o

Control and Fertil.iz,er
Cnly

Fertili z,er and :Ierb::'cide
Combination

Fig. 1. AV8rage forb pr oduc t.Lcn by tre8t;;'lent for
areas combined.

1971 date, "or all

three

study

�-220-

800

s:::
o

·rl

+'
V

~

M
0

'D

o

!5 0

~
c,

s:::

0

0
0

6

r&lt;"\

Control

0
I

0

&lt;o

0

6

0"&gt;

0
I

0
C\I

~

and Fertilizer
Only

Fig. 2. Average grass production
three study areas combined.

C\I
I

0

C\I
I

0

r&lt;"\

C\I

6

&lt;o

C\I

6

0"&gt;

C\I
I

0
C\I

~

and Herbicide
Fertilizer
Combina.tion

by treatment

for

1971 data

for all

�-221-

Figures 3, 4, 5 and 6 depict responses of shrub leafy material and total
leafy material to increasing levels of nitrogen fertilizer with and without 2,4-D herbicide at each study area. The general linear increase in
production with increased levels of nitrogen noted the first year after
treatment is still evident.
Figures 5 and 6 point out the general
leveling effect that herbicide treatments in combination with increasing
increments of nitrogen have on vegetation production.
Figures 7 and 8
show the average percent change in total leafy material production relative to control treatments.
Data are presented for each herbicide
and each fertilizer treatment for all three study areas combined, 19691971 where 1969 is the pre-treatment year. A randomized block analysis
of variance was used for both shrub leaf production and total leaf
production values for all treatments at each study area. First year
response data for shrub leaf weight showed a significant (P~.09) response
for the main effect of fertilizer and a highly significant response
(p &lt;.01) for herbicide effect.
Total leaf production showed a significant
(P~.07) response for the main effect of fertilizer and a highly significant response (P&lt;.Ol) for the herbicide effect.
A breakdown of the main effect of fertilizer into single degree of freedom
components allowed tests of significance to be made on each increment of
fertilizer.
This breakdown showed the main effect of fertilizer on first
year shrub leaf responses was received at the 30 pound level.
None of the
additional increments had a significant effect (P~.05) on production, but
each increment did result in increased production.
The 30 pound level
also gave the only significant response (P&lt;.05) in total leafy material.
However, a test for linearity for all levels of nitrogen showed a significant (P&lt;.05) linear response for both weight components with each added
increment of nitrogen.
Tests for quadratic, cubic and quartic relationships showed that these relationships contributed little to the sum of
the squares.
Analyses of second year post-treatment data produced similar results.
Shrub leaf components showed highly significant (P&lt;.Ol) responses to both
the main effect of fertilizer and herbicide.
Total leafy material also
showed a highly significant (P &lt; .01) response to both the main effect of
fertilizer and herbicide.
Breakdown of the main effect of fertilizer into single degree of freedom
components again pointed out that the main effect of fertilizer was received at the 30 pound level. None of the additional increments resulted
in a significant (P~.05) effect.
This held true for both shrub leaf and
total leaf production.
However, the test for linearity once again showed
a significant linear response for both weight components with each added
increment of nitrogen.
Table 5 presents average increases in total leafy material produced per
pound of nitrogen applied for 1970 and 1971. It is obvious that initial
rates result in the greatest return per pound of nitrogen applied.

�-222-

«&lt;;

(v
h
C)

&lt;;:!

s.,
Q;

o.
(fJ
~j

§
0

c ..

~

0
0.-,

+'
()

;:i
"0
0
H

rOf

0

,
·,',r:::-:j.S

o f ~i tro:::en, 1971.

�-223-

LTun~"'::.ior:. "DU+t.e

J'~ir.:h tline

�-224-

.Junc t.i on But.t.e

~

o

.r!

+-'

o

::&lt;

'6.....•

n..

0/2

30/2

60/2

~'(I;,ds
of ::::le:neY)talNi t·ogen
Pouid s of ?,Lj.-D tIel~()i.ci-Je

?L:-_ 5.
levels

of

Fer ,kre

".dth 2

~)er Acre

Total leafy v2::"st.ation
"!:'ocj.j':,ed as a function
of Lr.cr-e as ed
ni troe';er. id t.h 2,1.j.-~ her-b ic i de for alJ. three s tudy areas, 1971.

�-225-

,-..
(l)

r...•

1600

0

"'--r:

~
(l)

P-.
Ul
'0

c

::J

0

r4

c

0
-r1

Junction

+)

Butte

(.

::J

-o
o
~
o,

Fli[fhtline
Corral Creek
0

0/2

30/2

60/2

90/2

120/2

POUDrlsof~~J_8J!.ental ntro.ren
Fer Acre with 2
?ounds of 2,1-~-D fi:erb:i.cide Per Acre
Fir:o 50 Shrub leaf material
nr oduced as a --"lJ.rJ:tion of .incr-eased levels
of nitro~~en l...rith2,lj--D herc'cide,
1971.

�-226-

1971
Q)

b.lJ

~
C!i
..c

u

1969
1970

+'

s::
Q)
o
$..,
Q)

c,

0/2

30/2

90/2

120/2

Fig. 7. ::~vera-"epercsr:t ohan=e in total leafy production 0: each
herbicide treatment relative
to th':3 ccmtro'l tro"trr,ents for all three
study areas co:,~bined "'or the years 1969, 1970, 1971.

�-227-

1970
1971

1969

"7it!. (;. ,\V8raze per-cen t. chan-e in total leafy pr-c+uctd on of ea.ch
fertilL'8r
treatment rAICl-;:,i V(," to the ~0n+rol
tr,"'8t"l)ents for 2.11
t'::ree study areas oomb.ined for the years 1969, 1970, and 1971.

�-228-

Table 5. Average increase in pounds of total leafy material
of nitrogen applied for all three study areas combined.

Treatment
(Lbs. Per Acre)

per pound

Increased Production
Per Pound of Nitrogen Applied
1970
1971

30 N

3.9

5.3

60 N

4.5

4.1

90 N

1.9

2.8

120 N

3.2

3.2

Sample size requirements
for total vegetation production estimates were
determined by using a double sampling computer program.
The degree of precision was set so sample size requirements would result in an estimate
within 10 percent of the true population mean 95 percent of the time.
Table 6 presents actual samples taken and numbers of both large (meter
read plots only) and small (clip plots) samples that would be required
to meet the stated degree of precision for estimating total leafy material
at each study area for years 1969, 1970, and 1971. Data presented represent
the combined units of five fertilizer treatments and five herbicide treatments at each study area for years 1970 and 1971. This results in a large
sample size of 375 (75X5) and a small sample size of 75 (15X5) where the
ratio between large and small samples is 5:1. The 1969 pre-treatment
data
represent all 10 treatments combined.
The sample required to meet the
stated degree of precision for estimating total leafy material has been
met in every instance except the small sample taken in 1970 for Junction
Butte herbicide treatments and the large 1970 sample of Junction Butte
fertilizer treatments.

Botanical

Composition

Botanical composition measurements were made in late July, 1971. All
measurement
data for years 1969, 1970, and 1971 have been punched on data
processing cards for computer analysis.
Covariance analysis of variance
will be performed on these data to assess treatment effects.

�-229-

Table 6. Actual number of samples taken and the number of samples required to sample total leafy material produced within 10 percent of the
true population mean 95 percent of the time at all study areas for the
years 1969, 1970, and 1971.

Year
Area and
Treatments

Junction

Type
of Sample

1969 1/
Actual

1971

1970

Required

Actual

Required

Actual Required

Butte

Fertilizer

Herbicide

Large 1/
Small 1/
Large
Small

500
100

381
81

375
75

416 2:/
52

375
75

365
59

375
75

369 /
971.

375
75

365
72

375
75

262
30

375
75

303
58

375
75

258
58

375
75

299
49

375
75

252
34

375
75

252
35

375
75

204
64

375
75

192
57

Flightline
Fertilizer

Large
Small

Herbicide

Large
Small

500
100

317
40

Corral Creek
Fertilizer

Large
Small

Herbicide

Large
Small

1/

- For the 1969 pre-treatment
not apply.

2/

- Denotes

3/

adequate

500
100

310
65

data the fertilizer

and herbicide

breakdowns

do

sample not taken.

- Large sample represents meter readings
plots.

only; small sample represents

clip

�-230-

Deer Grazing Measurements

Plant Preference

Data

A summary of diets of tame, trained mule deer for each of the three study
areas for the 1969-70, 1970-71, and 1971-72 winters is presented in Tables
7, 8, and 9. Table 10 gives all cammon and scientific names of plants used
in this report.
Table 11 summarizes plant components in diets for all
three study areas for three years.
It is evident from this table grasses
increased and forbs decreased dramatically in 1971-72 compared with 196970 pre-treatment diets. Shrub components in the diets have not changed
noticeably.
Changes in the grass and forb composition of the diet are
believed to result from vegetation composition changes following herbicide treatments.
Tables 12, 13, and 14 summarize the diet by plant components for each study area. These tables point out large decreases in
forbs consumed by deer at Flightline and Corral Creek for 1969-70 to 197071. This decrease is not noticed at Junction Butte primarily because
Phlox bryoides (a low growing, woody-based plant classed as a forb and
readily eaten by deer) was not severely reduced by herbicide treatments.
Increases in grass percentages can be noted at each study area. This
increase is most noticeable at Corral Creek. Greatest grass responses
to herbicide treatments also occurred there. For some unknown reason
total shrub percentages in the diet did not decrease following herbicide
treatments.
However, it is suspected that increased growth of shrub
species, resulting from nitrogen combined with herbicide, counteracted
the decrease (Fig. 6).
Table 15 presents a comparison of certain selected plant species in diets
for the three winters for all three study areas.
Note that most shrub
species percentages remained fairly constant for the three years.
However,
green rabbitbrush (Chrysothamnus viscidiflorus) showed a significant increase from pre-treatment to post-treatment grazing.
This species also
increased following fertilizer-herbicide
treatments.
Also, bluebunch
wheatgrass (Agropyron spicatum) increased after chemical treatment and
and in the diet.
Buckwheat (Eriogonum umbellatum), the most prominent
forb in the pre-treatment diet, decreased to an insignificant portion
of post-treatment diets but was killed extensively by herbicide treatments,
however, snow cover in the 1971-72 winter somewhat precluded deer use of
this forb. Phlox bryoides, the other forb listed for comparison, received
a slight increase in deer use from pre-treatment to post-treatment.
As
previously mentioned, it seemed to be unaffected by herbicide treatments.
Table 16 summarizes total number of bites of all species of plants taken by
deer on each treatment at each study area for three winters.
Greater numbers of bites were taken on control and fertilizer-only plots compared with
fertilizer-herbicide
combinations in 1970-71. This was not evident in
1971-72. In fact, the 1971-72 data are almost identifical to the 1969-70
pre-treatment data.
Carpenter (1971) reported snow crust conditions in 197071 differed between herbicide treated plots and those which received no herbicide. Herbicide treated plots generally had harder, thicker crusts than nonherbicide plots.
In contrast, there were no apparent differences between
snow conditions on herbicide and non-herbicide areas in 1971-72, and this
similarity in snow environment may explain the similarity in diets from
herbicide and non-herbicide treatments for 1971-72.

�-231-

Table 7. Junction Butte study area grazing observation data and diet
composition for deer during the 1969-70, 1970-71, 1971-72 winters.

Plant Species
Serviceberry
B1uebunch wheatgrass
Snowberry
Pine need1egrass
Snakeweed
Blue granuna
Buckwheat
Big sagebrush
Phlox bryoides
Bitterbrush
Mutton grass
Fringed sage
Rabbi tbrush
(Chrysothamnus spp.)
Lichens
Oregon grape
Penstemon cyathoporus
Unknown grass
Western wheatgrass
Phlox multiflora
Junegrass
Chrysothamnus Earryi
Green rabbitbrush
Needle and thread
grass
Bluebell
Bott1ebrush
squirre1tai1 grass
Currant
Indian ricegrass
Poa spp.
Unknown forb
Totals

1969-70
Total
Percent
Bites
of Diet

1970-71
Total Percent
Bites of Diet

1971-72
Total Percent
Bites of Diet

830
375
288
143
l31
105
92
64
63
62
45
27

36.5
16.5
12.6
6.3
5.7
4.7
4.0
2.8
2.8
2.7
2.0
1.3

725
326
210
204

22.5
10.1
6.5
6.3

184
294
85
427
22
73

5.7
9.1
2.6
l3.3
0.7
2.3

614
1135
53
2
67
143
4
83
675

14.1
26.1
1.2
T
1.6
3.3
T
2.0
16.0

80
386

1.9
9.0

17
12
6
4
4
2
2
2
2

0.8
0.5
0.2
0.2
0.1
0.1
0.1
0.1

4

T

24
90
232

0.1
2.2
5.4

2,276

100.0

3
52
263

0.1
1.6
8.2

5

0.1

290

9.0

729

17.0

14
5

0.4
0.2

8
2

T
T

18
7
15

0.6
0.2
0.5

2

T

21
1

0.1
T

4,355

100.0

3,222

100.0

�-232-

Table 8. Flight line study area grazing observation data and diet
composition for deer during the 1969-70, 1970-71, 1971-72 winters.

Plant Species

1969-70
Total
Percent
Bites
of Diet

1970-71
Total Percent
Bites of Diet

1971-72
Total Percent
Bites of Diet

Bitterbrush
2,075
Buckwheat
462
Snowberry
290
Pine need1egrass
176
B1uebunch wheatgrass
122
Mutton grass
95
Mat penstemon
95
Phlox bryoides
84
Unknown grass
58
Serviceberry
45
Paintbrush
43
Green rabbitbrush
14
Rose
13
Penstemon spp.
11
Phlox multiflora
11
Sedge
9
Unknown forbs
6
Bott1ebrush squirre1tai1
grass
5
Penstemon c~athoEorus
4
Big sagebrush
4
Junegrass
3
Poa spp.
Knotweed
2
Rothrock sage
1
Bastard toadf1ax
Arrow1eaf ba1samroot
Lupine
Astragalus conva11arius
Oregon grape
Rockcress
Bluebell
Penstemon watsoni
Alkali bluegrass
Fowl bluegrass
Western wheatgrass

57.2
12.7
8.0
4.9
3.3
2.6
2.6
2.3
1.6
1.2
1.2
0.4
0.4
0.3
0.3
0.2
0.2

4,051
493
299
359
28
545
12

53.1
6.4
3.9
4.7
0.4
7.1
0.2

5,719
5
1,368
94
1,048
129

55.0
0.1
13.1
0.9
10.1
1.2

523
289
30
593
38
4
46
18
5

6.8
3.9
0.4
7.8
0.5
0.05
0.6
0.2
0.1

208
475

2.0
4.6

747
18

7.2
0.2

14

0.1

6

0.1

0.1
0.1
0.1
0.1

147
12
34

1.9
0.2
0.4

5

0.1

120

1.2

63

0.8

21

0.2

19
5
5
5
4
1
1

0.2
0.1
0.1
0.1
0.05
T
T

13

0.1

10
167
38
182

0.1
1.6
0.4
1.7

Totals

100.0

10,387

100.0

3,628

0.1
0.1

7,629

100.0

�-233-

Table 9. Corral Creek study area grazing observation data and diet
composition for deer during the 1969-70, 1970-71, 1971-72 winters.

Plant Species

1969-70
Total
Percent
Bites
of Diet

Bitterbrush
892
Buckwheat
756
Mat penstemon
375
231
Mutton grass
71
Pine need1egrass
Pusseytoes
70
60
Serviceberry
B1uebunch wheatgrass
57
45
Snowberry
Phlox multiflora
45
16
Green rabbitbrush
6
Unknown forbs
6
Senecio spp.
6
Senecio mu1ti1obatus
5
Gilia
3
Aster spp.
3
Sedge
2
Rocky Mountain juniper
2
Astragalus conva11arius
1
Senecio integerrimus
Unknown grass
Big sagebrush
Bott1ebrush squirre1tai1
grass
Horsebrush
Chrysotharnnusnauseosus
Penstemon cyathoporus
Paintbrush
Poa spp.
Junegrass
Western wheatgrass
Fowl bluegrass
Alkali bluegrass
Lupine

33.6
28.5
14.1
8.7
2.7
2.6
2.3
2.2
1.7
1.7
0.6
0.2
0.2
0.2
0.2
0.1
0.1
0.1
0.1
0.1

2,652

100.0

Totals

1970-71
Total Percent
Bites of Diet

2,294
37

32.3
0.5

1971-72
Percent
Total
of Diet
Bites
2,213
28

17.9
0.3

54

0.5

1,050
1,599
863
9
808
19

14.8
22.4
12.2
0.1
11.4
0.3

1,557
5,911
449

12.6
47.8
3.6

1,034
29

8.4
0.3

190
11

2.7
0.1

174
11

1.4
T

104
45

1.5
0.8

58
260

0.5
2.1

21
19
11
6
5
4

0.3
0.3
0.1
0.08
0.07
0.05

39

0.3

5
10

T
T

28
112
360
18
12
2

0.3
0.9
2.9

12,364

100.0

7,095

100.0

O·f
T
T

�-236-

Table 14.
components

Summary of Corral Creek study area diet composition
for the 1969-70, 1970-71 and 1971-72 winters.

Plant Component

by plant

1969-70

1970-71

1971-72

Percent of Diet

Percent of Diet

Percent of Diet

Shrubs

38.3

74.4

46.0

Grasses

13.6
48.1

24.4
1.2

53.3

Forbs
Totals

100.0

100.0

100.0

0.7

Table 15. Comparison of certain selected plant species in the diet for
the 1969-70, 1970-71, and 1971-72 winters, for all three study areas.
Plant Species

1969-70

1970-71

1971-72

Percent of Diet

Percent of Diet

Percent of Diet

Bitterbrush

35.4

Serviceberry

Green rabbitbrush

10.9
7.3
0.8
0.4

Bluebunch wheatgrass

6.5

10.9

29.9

Pine needlegrass

4.8

3.1

0.4

Mutton

grass

4.4

3.4

1.0

Blue gramma

1.2

1.0

0.5

Bottlebrush squirreltail grass

T

1.0

0.2

Western wheatgrass

T

1.5

2.9

Buckwheat

15.4

4.6

0.1

Phlox bryoides

1.7

2.4

2.5

Snowberry
Sagebrush

35.5
11.5
7.6
0.9
9.4

29.3
9.8
6.9
1.7
9.3

�Table 16. Total number of bites of all species of plants taken by deer on each treatment at each study area
during the 1969-70, 1970-71, and 1971-72 winters. !/

Treatment
(Lbs. Per Acre)

Jct. Butte
69-70 70-71 71-72

Flightline
69-70 70-71 71-72

Corral Creek
69-70 70-7l 71-72

722
104
235
197
330

1127
173
81
84
155

461
6
67
86
126

Totals
69-70 70-71 71-72

Percent of Total
69-70 70-71 71-72

2310
283
383
367
611

27.0
3.3
4.5
4.3
7.1

16.6
4.4
3.9
16.7
18.2

17.8
1.6
0.6
16.8
10.2

46.2

59.8

47.0

o lb. N. 0 lb.
2,4-D
30 N
60 N
90 N
120 N

"

"
"
"

354
293
0
147
843

663
99
17
1048
489

1443
182
216
53
1365

2230
46
75
24
1570

214
61
255
1833
4

760
176
26
2383
32

2011
536
471
2033
2212

3653
321
118
3455
209l

Subtotals

I
N

o lb. N. 2 lb.
2,4-D
30 N
60 N
90 N
120 N

"
"
"
II

18
18
261
299
92

20
29
152
85
143

628
398
97
311
145

881
231
126
741
29

655
17
1645
185
99

642
326
2603
539
3

15
391
591
120
787

27
60
745
279
740

686
116
1545
1058
1771

914
640
978
1160
908

702
106
2542
549
982

1956
840
4245
1908
1919

Subtotals
Totals

!/

2276

2066

3895

3628

5860

8058

2650

4218

8553

8554

12144 20506

10.7
7.5
11.4
13.6
10.6

5.8
0.9
20.9
4.5
8.1

9.5
4.1
20.7
9.3
9.4

53.8

40.2

53.0

100.0 100.0 100.0

Total bite numbers will not compare exactly to diet composition data because of bites that were taken on the
treatment margins.

w
-...J
I

�-238-

Table 17 presents grass, forb, and shrub component percentages of diets
by treatment for all three study areas combined.
A larger percentage
(77.8%) of grass was taken on fertilizer and herbicide combination treatments than was taken on control or fertilizer only treatments.
In addition, a larger proportion (63.5%) of shrubs was taken on control or
fertilizer only treatments as compared with combination treatments.
However, in contrast to the 1970-71 winter where 91 percent of the forbs
were eaten on the control or fertilizer treatments, only 19 percent of
the forbs were taken on these treatments during the 1971-72 winter.
This change was attributed to the previously mentioned fact that Phlox
bryoides made up over 90 percent of the forb component of the diet, and
it was relatively unaffected by herbicide treatments.
This resulted in
a large percentage of the forb diet being taken on the herbicide treatments.

Table 17. Grass, forb, and shrub component percentages of deer diet by
treatment plot for all three study areas for the 1970-71 and 1971-72
winters.
Percent Grass

Treatment
(Lbs. Per Acre)

Percent

Shrubs

1970-71

1971-72

1970-71

1971-72

1970-71

1971-72

0 lb. N. o lb.
2,4-D

2.8

8.8

46.0

13.8

17.4

23.6

Percent Forbs

30 N

"

5.1

1.6

23.0

1.7

1.5

1.6

60 N

"

3.2

0.2

8.1

0.0

3.5

0.5

90 N

"

4.1

8.4

0.2

2.3

23.7

22.9

120 N

"

21.0

3.2

13.7

1.2

17.9

14.9

Subtotals

36.2

22.2

91.0

19.0

64.0

63.5

0 lb. N. 2 lb.
2,4-D

0.6

9.2

0.6

44.8

8.4

8.3

30 N

"

2.1

7.6

0.5

34.6

0.5

0.8

60 N

"

28.9

24.9

7.0

0.6

20.0

20.7

90 N

"

13.0

14.4

0.9

0.3

1.9

2.8

120 N

"

19.2

21.7

0.0

0.7

5.2

3.9

Subtotals

63.8

77.8

9.0

81.0

36.0

36.5

Totals

100.0

100.0

100.0

100.'0

100.0

100.0

�-239-

Treatment

Preference

Data

Tables 18, 19, and 20 summarize deer preference data by treatment for the
three winters for all three study areas. Data were recorded from the observation tower for total standing minutes per treatment, bedding minutes and
seconds spent grazing. Table 18 summarizes total minutes of deer grazing
observed on each treatment plot during species preference grazing trials
where the deer are accompanied by the observers.
The strong preference
shown for control or fertilizer only treatments over fertilizer-herbicide
combination treatments in 1970-71 is not evident in 1971-72. Differences
in snow crusting conditions mentioned previously at least partially accounted
for this change.
This same change in preference is also noted in deer
grazing seconds obtained from observation towers for each treatment at each
study area for the three winters (Table 19). In contrast to the 1970-71
winter in which about 70 percent of the gra~ing time was spent on control
or fertilizer only plots, grazing time was split about half and half in
1971-72. In addition, no definite preference in grazing time per individual treatment was noted in 1971-72.
Summaries of the number of minutes deer spent standing in each treatment
at each study area for all three winters as obtained from the observation
tower, show the same treatment preference change that was mentioned previously comparing the 1970-71 and 1971-72 winters (Table 20). Data in
1971-72 show almost 50 percent of the time spent on control and fertilizer only treatments in contrast to 65 percent of the time being spent on
these treatments in 1970-71. Again, no definite individual treatment
preferences are noted in these data.

Additional

Deer Grazing Measurements

A pasture 610' X 330' was fenced to obtain additional information regarding
deer grazing responses to nitrogen fertilizer.
This was equivalent to 4.6
acres. Within the fenced area three strips containing about 136,000 square
feet were treated with nitrogen, while 65,340 square feet were untreated.
The untreated strips were 66 feet wide by 330 feet long alternated between
the treated areas. The treatments were well marked to make them obvious
during subsequent grazing experiments.
To measure deer grazing responses on this study area six tame mule deer were
placed in the pasture for a 30 day period from January 19 to February 17,
1972. One observer followed one deer through each grazing trial. Deer food
choices and grazing activities were observed morning and evening each day
for the entire period.
Each grazing trial averaged about 74 minutes.
In
addition, each time an observer arrived at the study area, location and
activity of each deer was noted. During grazing observation trials the
location and activity of each deer was recorded every half hour for the
duration of the trial. This location and activity record was maintained
to provide additional data regarding treatment choices.
All six deer were
rotated in order during each succeeding grazing trial so that all deer were

�-240-

observed equal amounts of time. Observers began each trial from a randomly assigned starting point to minimize bias due to starting location.
Deer grazing activity was recorded in the same manner described by
Carpenter (1970). Deer were weighed at the beginning of their pasture
confinement and at weekly intervals thereafter through the duration of
the experiment to assess changes in relative physical condition.
Weights were taken with a Chatillon scale attached to a tripod and hooked
to a suspended deer holding crate (Fig. 9). Deer were weaned from their
standard concentrate ration of approximately 2 pounds per deer per day
at the start of the experiment, to no concentrate over a 10 day period.
The first six days of weaning were conducted before the 30 day trial
began.
The last four days of the weaning schedule coincided with the
first four days of the 30 day trial. A post-treatment ration schedule
was also followed and consisted of an increase from 200 grams of concentrate per deer per day from the 25th day of the trial to 500 grams
of concentrate per deer per day on the 30th day of the trial.
Because the three treated areas were larger than the three untreated,
they were further subdivided into 66' X 310' tracts with colored markers
placed on the pasture perimeter fence.
This partitioned the pasture into
10 subdivisions of nearly equal size and configuration.
Subdividing the
pasture in such a manner facilitated ultimate comparisons of treated and
untreated pasture preferences by deer.
Snow measurements were taken throughout the 30 day period.
Eighty snow
measurements were taken on 8 transects systematically located throughout
the pasture.
Snow measurements were taken only when snow conditions had
changed noticeably from previous measurements.
Snow depth and crust
conditions were recorded for each plot. Air temperature was recorded at
the beginning and end of each grazing observation period.
At the termination of the 30 day period following snow melt, a pellet
group survey was conducted over the entire area in an additional attempt
to measure deer use on each treatment area. Twenty-six transects of 10,
.001 acre plots were located in the pasture.
Both "old" and "new" pellet
groups were recorded.
A record of the old pellet groups represented previous wild deer use of the pasture area.

Plant Preference

Data

A total of 72,531 bites was gathered during the 30 day period during a total
observation time of 5,881 minutes (98.017 hours). A total of 79 deer trials
was conducted throughout this period for an average of 2.6 trials per day and
an average of 918 bites per trial. Deer diets for the 30 day period are
summarized in Table 21. Grasses comprised 70.2 percent of the combined diets.
The main grass species eaten was bluebunch wheatgrass which made up 43.4 percent of the diet.
Big sagebrush comprised a considerably larger portion of
the diet than had been noted previously in other grazing trials in this study.
Sagebrush percentages in the diet increased steadily from the beginning of
the 20 day trial until the end (Fig. 10). Grass percentages increased until
midway through the 30 day period and then decreased (Fig. 11). Forbs made
up a relatively small percentage (9.1) of the diet while shrubs made up 20.7
percent of the total diet for the 30 day period.

�Table 18. Total deer minutes spent on each treatment while obtaining plant preference measurements at each
study area for the 1969-70, 1970-71, 1971-72 winters.
Treatment
(Lbs. Per Acre)

Jet. Butte
69-70 70-71 71-72

Flightline
69-70 70-71 71-72

Corral Creek
69-70 70-71 71-72

Totals
69-70 70-71

71-72

Percent of Total
69-70 70-71 71-72

500
66
39
441
301

18.4
4.2
7.0
8.5
11.3

14.9
6.9
5.0
14.7
23.3

20.0
2.6
1.6
17.6
12.0

49.4

64.3

53.8

o lb. N. 0 lb.
2,4-D
30 N
"
60 N
"
90 N
"
120 N
"

151
19
69
63
116

80
40
2
49
210

157
31
16
163
156

187
62
48
97
70

159
58
32
28
195

267
14
14
7
137

47
6
29
17
50

25
24
54
183
7

76
21
9
271
8

385
87
146
177
236

264
122
88
260
412

Subtotals

1
N

+--

r-'
1

o lb. N. 2 lb.
2,4-D
30 N
"
60 N
"
90 N
"
120 N
"

7
21
65
97
51

8
12
46
17
66

92
72
13
30
93

103
82
63
167
42

47
19
184
39
30

72
75
234
101
8

12
135
109
10
93

2
12
65
35
43

47
12
82
73
154

122
238
237
274
186

57
43
295
91
139

211
159
329
204
255

Subtotals

Totals

659

530

823

921

79l

929

508

450

753

2088

1771

2505

5.8
11.4
11.4
13.1
8.9

3.2
2.4
16.7
5.1
7.8

8.4
6.4
13.1
8.1
10.2

50.6

35.7

46.2

100.0 100.0 100.0

�Table 19. Total deer grazing seconds obtained from the observation tower fo~ each treatment at eaeh study
area for the 1969-70, 1970-71, 1971-72 winters.

Treatment
(Lbs . Per Acre)

o lb. N. 0 lb.
2,4-D
30 N
"
60 N
"
90 N
"
120 N
"

Jet. Butte
69-70 70-71 71-72

1643
0
3609
1003
496

2366
40
163
724
4700

1432
1626
845
995
2202

Flight1ine
69-70 70-71 71-72

Corral Creek
69-70 70-71 71-72

69-70

1700
287
574
801
2285

325
129
0
1291
0

1183
0
978
8122
273

3668
416
4183
3095
2781

1331
1080
2712
1562
7172

3301
1700
2169
110
5856

1402
81
503
9060
10

Totals
70-71 71-72

5099
1201
3378
11346
11882

5916
3326
3992
9227
8331

Subtotals

Percent of Total
69-70 70-71 71-72

15.0
1.7
17.1
12.7
11.4

10.8
2.5
7.1
23.9
25.1

9.8
5.5
6.6
15.3
13.8

57.9

69.4

51.0

I

N

.pN
I

o lb. N. 2 lb.
2,4-D
30 N
"
60 N
"
90 N
"
120 N
"

0
0
1462
1186
0

230
78
711
3751
1272

3028
648
1678
5149
1358

876
94
777
28
653

498
294
2493
266
901

3355
422
2090
6742
826

30
0
1787
0
3359

24
336
1791
347
1543

951
1039
327
121
1934

906
94
4026
1214
4012

752 7334
708 2109
4995 4095
4364 12012
3716 4118

Subtotals
Totals

9399

14035

18961

8075

18309

26571

6921

15097

14928

24395

47441 60460

3.7
0.4
16.5
5.0
16.5

1.6
1.5
10.5
9.2
7.8

12.1
3.5
6.8
19.8
6.8

42.1

30.6

49.0

100.0 100.0 100.0

�Table 20. Total number of deer standing minutes from observation towers per treatment plot at each study area during
the 1969-70, 1970-71, 1971-72 winters.
Treatment
(Lbs . Per Acre)

Jet. Butte
69-70 70-71 71-72

Flightline
69-70 70-71 71-72

150
1
410
23
49

126
24
79
60
187

Corral Creek
69-70 70-71 71-72

69-70

Totals
70-71 71-72

Percent of Total
69-70 70-71 71-72

o lb. N. 0 lb.
2,4-D
30 N
"
60 N
"
90 N
"
120 N
"

321
26
66
103
502

179
175
77
75
257

222
131
265
150
564

277
171
142
7
457

17
8
24
84
0

115
28
64
798
13

105
10
30
813
15

293
33
513
167
236

658
185
395
1051
1079

561
356
249
895
729

Subtotals

15.1
1.7
27.2
8.9
12.5

12.6
3.5
7.6
20.1
20.6

10.4
6.6
4.6
16.6
l3.5

65.8

64.4

51.7

,
N

+:w

,

o lb. N. 0 lb.
2,4-D
30 N
"
60 N
"
90 N
"
120 N
"

0
0
130
92
0

51
62
107
354
270

290
114
108
396
225

55
17
88
8
57

65
56
309
40
59

328
21
161
589
62

11
3
113
1
70

19
52
144
54
214

94
73
58
8
73

66
20
331
101
127

l35
170
560
448
543

712
208
327
993
360

Subtotals
Totals

855

1862

1896

701

1861

2215

331

1501

1279

1887

5224

5390

3.5
1.1
17.5
5.4
6.7

2.6
3.3
10.7
8.6
10.4

l3.2
3.9
6.1
18.4
6.7

34.2

35.6

48.3

100.0 100.0 100.0

�-244-

Fig. 9.

Device used in deer weighing

procedures.

�-245-

Table 21. Junction Butte volatile oil study area grazing observation
data for the 30 day period from January 19 to February 17, 1972.
Plant

Species

Bluebunch

wheatgrass

Total Bites

Percent

31,473

43.4

Big sagebrush

9,334

12.9

Unknown

grass

7,400

10.2

Phlox bryoides

6,277

8.7

Pine need1egrass

3,995

5.5

Western

wheatgrass

2,325

3.2

Mutton

grass

2,023

2.8

Serviceberry

1,967

2.7

Snowberry

1,402

2.0

Green rabbitbrush

1,394

1.9

Blue gramma

1,173

1.6

869

1.2

777

1.1

569

0.8

Junegrass

444

0.6

Needle

387

0.6

188

0.3

180

0.3

103

0.1

75

0.1

48

T

30

T

29

T

Indian ricegrass

9

T

Mat penstemon

2

T

Others

58

T

Totals

72,531

100.0

Poa spp.
Fringed

sage

Bott1ebrush

squirrel tail grass

and thread grass

Sandberg

bluegrass

Buckwheat
Penstemon
Ground

spp.

lichen

Greasewood
Unknown
Penstemon

forb
cyathoporus

of Diet

�-246-

25-26-27
31-1-2
22-23-24
28-29-30
3-4- 5

19-20-21

-Janusr'y

12- .1.)-14-

9-10-11

15-16-17

February

Fig. 10. Percent bif: sat~e'orush in diets of tame mule deer by three
day periods for Junction 3utte vclatile
oil study area, 1972.

�-247-

19-20-21
25-21)-27
31-1-2
22-2)-2'-!2[&lt;,-29-30

6-7-8
j-~}-5

12-1j-14
9-10-11
15-16-17

Fi::r. 11. Percent
cr-ass in diets of t.arne 'TIule deer
for Junction
:::utte votat.i.l.e oil s t.udy ar-ea , 1972.

by three day "geriods

�-248-

Treatment

Preference

Data

A thorough analysis of the data has not been completed at this time.
However, data summarized at this point indicate no uniform preference
for fertilized areas in contrast to unfertilized areas. A chi-square
analysis of the total number of bites observed for all treatments showed
there was a significant (P=.05) preference for the fertilized treatments,
but a closer analysis of the data revealed that this difference was
attributable to only one treatment strip and not to all fertilizer strips.
In fact, the other two fertilizer treatments had fewer observed bites
than would be expected if all treatments had been equal.
It was felt the
one strip was preferred by deer because of biological factors, such as
preferred bedding sites, location in the pasture, and basic deer behavior,
and did not represent an actual preference for treated forage.
A complete analysis of total minutes observed per treatment strip has not
been completed, but these data seem to follow the same pattern as the
total bites data with no apparent preference being shown for the fertilizer
treatments.
An analysis will also be made of deer bites on sagebrush,
bluebunch wheatgrass and total grasses on fertilized and untreated plots.
Summaries of deer locations and activities throughout the 30 day grazing
period did not show any differences between fertilized and unfertilized
strips.
The pellet group survey which was conducted on April 5, 1972, also failed
to reveal significant (P=.05) differences in pellet densities on treated
versus untreated strips.
A total of 147 new pellet groups was counted on
the 260 plots, with 82 groups (56%) on fertilized strips and 65 (44%) on
control strips.
An analysis of old pellet groups on the strips showed an
even larger percentage (71%) on fertilized strips as opposed to control
strips (29%). The difference may indicate that wild deer even prior to
the establishment of the pasture favored locations where fertilizer was
subsequently applied.

Deer Weight

Changes

Deer gradually lost weight during the 30 days they were entirely dependent
upon native pasture forage (Fig. 12). The maximum loss in weight was recorded at the February 9 weighing.
The gain in weight from February 9-13
is not fully understood.
Deer began recelvlng small amounts of concentrate
after February 13 and again showed a loss from that weighing period to the
terminal weighing period on February 17.

Other Measurements

Precipitation

Records

Table 22 presents comparisons of 1969, 1970, and 1971 precipitation data
with 13 year averages for Kremmling, Green Mountain, and Hot Sulphur Springs
weather stations.
Precipitation in 1971 was lowest since the study began.
Total precipitation in 1971 was about 1.1 inches below the 13 year average.

�-249-

+8

(I)

•..

~:Ij
,....

I.::

..c
u
+-'
...c

0

till
·rl

&lt;l'

~:
+-'

~

(J)

o

1-&lt;

(I)

p.,

1-19

1-26
Janl18.ry

1!ig.

2-9

2-17

2-13
Febr-uar-y

12. Aver:::l:~e pro:r?SSlVS ~;,:8iz:ht ch:'":lr..-~e of six +ams mul.e deer on a
4.6 s8~ebrl1sh nas t.ur-e "'8rlrti.31J.~,:t tre.":1"Sed i.,ri t.h ni,t.ro,;.:en f'er t.i.I izer dur mz
t.he 30 - dayoed_od
from J anuar y 19 to ?88r,18ry 17, 1972.

�Table 22. Comparison of the 1969, 1970, and 1971 monthly and annual average precipitation data with
the thirteen year averages for the Green Mountain, Kremmling !/, and Hot Sulphur Springs weather
stations. ]:./

Year

Jan.

Feb.

Mar.

Apr.

May

Month
July
June

Aug.

Sept.

Oct.

Nov.

Dec.

Annual
Total

13-year Average

0.89

0.76

0.91

0.99

1.04

1.55

1. 24

1. 66

1. 61

1.14

0.80

0.97

13.56

1969

1. 79

0.43

0.33

1.00

0.82

4.65

0.86

1.23

1.50

3.56

0.49

1.23

17.90

1970

0.76

0.31

1.17

1.22

0.13

1.41

0.85

1.82

2.00

2.01

1.38

0.80

13.86

1971

0.83

0.70

0.96

1.33

1.38

0.37

0.93

1.05

1.78

0.57

0.99

1.56

12.45
I

!/Kremmling weather records are incomplete for this time period.
llData from Colorado Climatological annual reports.

N
V1
0
I

�-251-

Photographs
The permanent photo hubs were photographed with black and white and
color film at the end of the growing season.

LITERATURE

CITED

Bryant, M. P., and L. A. Burkey.
1953. Cultural methods and some characteristics of some of the more numerous groups of bacteria in the
bovine rumen.
J. Dairy Sci. 36:205-217.
Bryant, M. P., and 1. M. Robinson.
1961. An improved non-selective
culture medium for ruminal bacteria and its use in determining diurnal
variation in numbers of bacteria in the rumen.
J. Dairy Sci.44:l446-l456.
Carpenter, L. H. 1970. Middle Park deer study - range fertilization.
Colo.
Div. Game, Fish and Parks.
Game Res. Div., Fed. Aid Prog. W-38-R-24.
Game Res. Dept. July, Part 3. pp. 371-391.
1971. Middle Park deer study - range fertilization.
Colo. Div.
Game, Fish and Parks. Game Res. Div., Fed. Aid Prog. W-38-R-2S.
Game
Res. Dept. July, Part 3. pp. 225-253.
Environmental Science Services Administration.
1965-1971.
data, Colorado.
U. S. Dept. Commerce.
70-76 (13).

Climatological

Paintner, W. W. 1971. Volatile oil content and composition of juniper and
its effect on deer rumen microorganisms.
M.S. Thesis, Colo. State Univ.
Ft. Collins.
40 pp.
Powell, J. 1968. Site factor relationships with volatile
brush.
PhD. Diss. Colo. State Univ.
82 pp.
U. S. Weather Bureau.
Dept. Commerce.

1958-1964.
63-69 (13).

Climatological

oils in big sage-

data, Colorado.

U. S.

Ward, G. M., and J. G. Nagy.
1966. Bacterial inhibition by the essential
oils of sagebrush.
Final report to the National Sci. Foundation, Grant
No. GB 1507. Colo. State Univ., Ft. Collins.
54 pp.

-F

«&gt;. .s.»:

Prepared by __ f-::~7)':"/':;:·-"':":~%7·" ...;'..:,.' :=--'-_-_'_c.....,.ci-,f'Len H. Carpenter
Student Assistant

L.::..

_

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                  <text>-253July, 1972

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~---------

Work Plan No.
Job Title
Period Covered:
Personnel:

Deer-Elk

W-38-R-26

Project No.

Investigations

7
Job No.
Middle Park Cooperative Deer Study Remote Sensing of Deer Population Parameters

14

June 1, 1971 to May 31, 1972

Ro Bruce Gill, R. S. Driscoll, Ho Do Parker, Jro (Graduate Student),
T. Ho Watson, Po Lo Shaw, Paul Fo Gilbert, Gary Brown, Larry Roper,
Dan Baker, Lonnie Brown, Rick Hoffman, F. W. Knott, R. E. Francis,
CSU Wildlife Biology Students.

ABSTRACT
An airborne, thermal infrared scanner was flown over three study sites in
Middle Park, Simultaneously with black and white aerial photographic coverage.
AI"though intended to provide a check on the number of deer detected by the
thermal scanner, the photographs failed to detect any animals due primarily to
minimum light conditions under which the photography 'was flown.
The thermal
scanner did detect deer, cattle and elk. However, without the aerial photographs,
the only positive identifications of animals in the thermal imagery were those
on animals visually spotted from the aircraft during the scanning operation.
Further analysis, including electronic imagery enhancement procedures of the taped
video data, is planned for 1972-1973.
Aerial color infrared (CIR) photographs
at two scales were obtained on July 2-3, 1971 of three transects 1 mile long
by 1/10 mile wide located in the Middle Park deer winter range.
The hypothesis
of this experiment was that this kind of aerial photography could be used to
estimate over-winter mule deer mortality.
On the average, eight interpreters
detected and identified 57 percent of the carcasses in the transect areas using
1:2000 scale photographs and verified by ground truth. Five interpreters with
previous training using CIR aerial photographs detected and identified an average
of 68 percent of the carcasses.
The other three interpreters with minimum training using CIR photographs identified an average of 36 percent of the carcasses.
Interpretation should improve significantly by flying early in the spring after
snow melt but before advanced carcass erosiono

��-255-

MIDDLE PARK DEER STUDY
REMOTE SENSING OF DEER POPULATION PARAMETERS
R. S. Driscoll and R. Bruce Gill

P. S. OBJECTIVE
To test the capabilities of aerial thermal and color infrared remote
sensing techniques to estimate: densities of live and dead deer.

SEGMENT OBJECTIVES
1.

Test the capability of an airborne thermal infrared scanner to
detect numbers of live deer occupying a selected study area in
Middle Park.

2.

Compare ground search estimates of simulated deer winter mortality
with estimates obtained from aerial color infrared photographic
imagery.

METHOD::;AND MATERIALS

Thermal Scanning Test for Live Deer Detection
The thermal scanner used was a Texas Instruments Model FFS-l, owned by
the U.S.D.A. Forest Service, Region 1, Missoula, Montana. It was a
modified Texas Instruments ~[odel RS-7, specifically designed for detection
of forest fires. The scanner was flown in a Forest Service Beechcraft
Kingair, modified and equipped for the purpose. Characteristics of the
scanner were as follows:
Instantaneous Angular Field of View (IFOV):
Total Angular Field of View:
Detectors:

2.0 milliradians

120°

Indium antitnonide (Channel A) and mercury-cadmium-telluride
(Channel B)

Thermal Sensitivity:

2°C (0.2°C with 20x overscan)

�-256-

Scanning Speed (t):

.005 sec. per scan line

Wavelength Response:

Channel A - 3.0 - 4.1 micrometers
Channel B - 8.5 - 11.0 micrometers

Data Output Mode:

127 mm filmstrip, 1/2 inch electrical tape

Further discussion of the system may be found in Wilson, et al. (1971).
On-board processing of the filmed imagery was provided by an Ansco Model
KD-14 rapid film processor and camera system. This system provides black
and white film imagery within a few seconds after scanning. The video
was also recorded on electromagnetic tape, by an Ampex Model FR 1300 analog
tape recorder, operating at 60 inches per second tape speed. Video from
both channels was recorded on tape; only Channel B was recorded on film.
Simultaneous black and white aerial photographs of the test areas was
provided by a Zeiss, Model RMK A 21/23 9 X 9-inch aerial camera, using
Kodak Tri-X Aerographic (Type 2403) film. The camera system was flown
in a North American Rockwell Aerocommander 500B owned by the U.S.D.A.
Forest Service, Region 5, Berkeley, California. The Aerocommander flew
above the Kingair, at an altitude between 10,000 and 11,000 feet.
The test was conducted on January 25, 26, and 27, 1972, in three missions
over one intensive test site and two extensive sites. Each mission was
flown as early in the morning as possible, consistent with flying safety.
Both aircraft were hangared at Denver, Colorado, about 30 minutes flying
time from Middle Park. They departed from Denver each morning before
daylight and data gathering from the aircraft occurred between 0700 and
0800 hours. There was approximately 90 percent low cloud cover during
all three mission days; ceiling approximately 4,000 feet above ground
datum.
Three areas were selected from which thermal scanner imagery would be
obtained: Jensen Creek, Cedar Ridge, and Junction Butte. These areas
are described in the next section.
The Jensen Creek area was divided into four flight lines 1,500 feet
apart. They were all flown in the same direction at a magnetic heading
of 229°. Coverage was planned at two altitudes, approximately 650 and
1,000 feet above the ground. The 120° sweep of the scanner provided
imagery overlap between adjacent flight lines of 375 feet at the 650-ft.
altitude, and about 577 feet at the 1,000-ft. altitude. The 1,000-ft.
altitude was flown first, to minimize disturbance of deer in the area.

�-257-

The Cedar Ridge area was divided into eight flight lines for a total
distance of about 22 miles. The magnetic heading for all flight lines
was 271°. Only one altitude, 650 feet, was flown on the Cedar Ridge
ar~a. The height above ground was only approximate, however, due to
topographic variation.
A deer pasture at Junction Butte was flown at three altitudes, 300,
600, and 1,000 feet above the ground. Only one pass at each altitude
was required to cover the area.
The initial points of all flight lines on the Jensen Creek and Cedar
Ridge areas were marked with either fluorescent orange panel markers
or red flares. The panel markers were arranged into a "T" shaped
pattern. The flares used were standard, IS-minute highway flares.
Although intended to be supplementary to the panel markers, the flares
proved to be the primary means of locating the flight lines from the
air. The panel markers were almost invisible under the 10~1 light
conditions prevailing.
The use of flares necessitated the presence of ground personnel to
ignite them at each of the test areas during the flights. The panel
markers were laid out the day before each mission and were anchored
to the ground with steel rods.
Two Barnes PRT-5 infrared radiometers were used to record radiant
temperatures of vegetation, snow, and rocks during all missions. On
the morning of the Junction Butte mission, radiant temperatures of
six tame deer were also recorded.

Aerial Color Infrared Photography for Deer Mortality
At each of three locations described in the next section, a belt
transect 1 mile long by 1/10 mile wide was established. These represented sample units used to obtain ground sample estimates of mule
deer over-winter mortality (Gill, 1969). Each transect was divided
into 640 one-chain-square blocks. Ten blocks were randomly selected
from each transect and a deer carcass was placed in the center of
each block.
Each carcass was placed flat on the ground to simulate the appearance
of a naturally fallen deer. For example, no carcasses were draped
over bushes. After placement, each carcass was described as to whether
it was an adult or fawn, relative stage of decomposition, and the degree
to which it was obscured by surrounding vegetation. The carcasses,
resulting from natural death, had been previously stock-piled so there
would be an adequate number for the experiment.

�-258-

Aerial photographs of the transect areas were obtained on July 2-3,
1971, between 0900 and 1100 hours MST. This represented a 3-week
delay in original planning due to logistics problems in securing an
airplane to the area with the required photographic equipment. A
Forest Service Cessna 180, in which two 70 mm Maurer KB-8A cameras
equipped with 150 mm Schneider Xenator lenses had been mounted, was
used for the mission. Two film types were used, one in each camera;
Ektachrome Infrared Areo, type 8443 and Aerochrome Infrared, type 2443.
The CIR film type was selected because previous research not specifically designed for deer mortality estimates indicated it to be superior
to normal color film to detect and identify deer carcasses (Driscoll,
1971). The type 2443 film was used because it was reported to have
improved color balance (Driscoll, 1969).
These are both "false color" reversal films and differ from ordinary
color film in that the three dye layers are sensitive to green, red,
and infrared radiation instead of the usual blue, green, and red used
for normal rendition of the visible spectrum. However, since all three
layers are also sensitive to blue radiation, a yellow Wratten 12 filter
was used in front of the lenses to absorb that part of the spectrum.
When exposed and processed properly, this combination of sensitizations
and dyes produces false colors for most natural objects. For example,
rapidly growing green vegetation produces an image in shades of magenta
or red. Bare soil or mature drying vegetation appears as shades of cyan
or blue. Heller (1970) describes in more detail the light/film physics
involved when using this film type.
Two photo scales were planned: 1:1500 and 1:3400. The smallest scale
was selected because previous work indicated that it might be near the
maximum for deer carcass detection and identification (Driscoll, 1971).
The largest scale was selected to determine if it might be the optimum
for the objective. Severe ground turbulance at the time of the mission
precluded securing these photo scales precisely; the resultant scales
averaged 1:2000 and 1:4000 over the three transects. Even then, three
of the carcasses were not photographed because airplane drift could not
be controlled. Consequently, 27 of the total carcass population of 30
were photographed.
The laboratory procedure consisted of developing a photointerpretation
test which was subsequently presented to eight photointerpreters with
varying amounts of photointerpretation experience. Since all transects
were not completely photographed at the largest scale, all stereo frames
in which deer carcasses were imaged, validated by ground truth and carcass location maps, were selected for the test. Therefore, some carcasses
images were represented more than once, but in different stereo pairs,
in the test. In total, 47 imaged carcasses formed the population for
the photointerpretation test.

�-259-

DESCRIPTION OF AREA

Thermal Scanning Test for Live Deer Detection
The intensive test area was on the southwest slope of Junction Butte.
A 330- X 6l0-ft. deer enclosure, built by the Colorado Division of
Game, Fish and Parks, was used to hold six tame mule deer, three
bucks and three does, while the area was scanned. The deer, all 19
months old, weighed 90, 96, 116, 124, 152, and 155 pounds on the
morning of the flight. They were put into the pen on January 19.
The first of the extensive sites flown was an area of about 2 square
miles, along Jensen Creek, north of Parshall, Colorado. The area
was selected because, historically, large numbers of mule deer concentrate on it during part of the winter. Also, the area has little
topographic relief (about 400 ft.) compared to much of Middle Park,
and there is no coniferous overstory. Most of the Jensen Creek area
is open, slightly rolling terrain, characterized by big sagebrush
(Artemisia tridentata) and other shrubby species, with a few small
pockets of aspen (Populus tremuloides).
The largest of the two extensive sites was Cedar Ridge. This area,
southwest of Parhsall, is about 7 square miles in size, and is characterized by considerable topographic variation. Elevation varies from
approximately 7,600 to 8,400 feet above mean sea level. Bounded on
the north by the Colorado River, on the south by a county road, and
on the east by the Williams Fork River, the area was described by
Gilbert and Grieb (1957) as follows:
"The terrain consists of high rolling ridges with numerous rocky
outcrops. There are pockets of dense Douglas-fir (Pseudotsuga menziesii
var. glauca) timber on the north-facing exposure and the south-facing
slopes support big sagebrush (Artemisia tridentata) with an open overstory of juniper (Juniperus monosperma). Patches of aspen (Populus
tremuloides var. aurea) are scattered throughout."
The area was selected because it was to be drive-censused on the
Saturday following the thermal scanning mission (January 29), and a
comparison between the two census methods could be made. The Cedar
Ridge area is drive-censused each winter by the Colorado Division
of Game, Fish and Parks with the help of wildlife biology students
from Colorado State University. Also, the rough terrain and juniper
overstory provided conditions for testing which were similar to those
which would be encountered in practical applications of the technique
for deer census.

�-260-

Aerial Color Infrared Photography for Deer Mortality
The transects for the research in this segment objective were located
in the big sagebrush (Artemisia tridentata) type in Middle Park near
Kremmling. The areas selected were within that strata of the winter
range where previous mortality estimates were greatest (Gill, 1969).
The specific transect locations were purposefully selected to exclude
a tree overs tory since the aerial camera lens does not "see" through
a tree canopy. A mixed shrub community was common to all locations
and was represented by a very dense to very sparse canopy cover.
Specifically the three transects that were selected occurred at the
following locations:
1.

Little Wolford: Wl/2, El/2, S2l, T2N, R8W; immediately
east and south of Little Wolford Mountain.

2.

Cow Gulch: Center S32, T2N, R8W; approximately 2 miles
north of Kremmling.

3.

Elliott Creek: Center S32, TIN, R8W; lower northwest flank
of Lawson Ridge and south of Elliott Creek.

RESULTS AND DISCUSSION

Thermal Scanning Test for Live Deer Detection
The most comprehensive set of radiant temperature data was obtained
at the Junction Butte site on January 26. These measurements of snow,
vegetation, bare soil, and tame deer were made between 6:00 and 6:30 a.m.
Thermal contrast between deer and sagebrush (~Ts) averaged 1.6°C, with
a minimum potential ~Ts of O.SoC. Average ~Tn (deer vs. snow) was
6.9°C, and average ~Tsoil (deer vs. bare soil) was 4.0°C. Air temperature was 1°C. Radiometer malfunction rendered ERT measurements on
the Jensen Creek and Cedar Ridge sites unusable.
The photographs were generally unusable due to very poor photographic
light conditions when the data were obtained. Even under clear skies,
the early morning missions with diffuse light would not be optimum
for obtaining high quality photographs. However, we needed to know
not only numbers but location of animals to provide validation of
data obtained by the scanner. Aerial photographs provided the only
possible way to secure this information.

�-261-

The photographic
scale varied from about 1:3000 to 1:5000 as camerato-ground distance varied with terrain elevation.
Flight altitude
of the Aerocommander
was constant at 10,050 feet on the Jensen Creek
site, and 10,800 feet over Cedar Ridge.
These scales were inadequate
to visually detect deer in the photos under the prevailing poor light
conditions.
Neither could cattle present along the Colorado River
be seen.
Scale, however, was near the minimum of 1:5000 in the low
areas bordering the river.
Both the Jensen Creek and Cedar Ridge missions produced thermal imagery
in which some animals were apparent.
However, since the corroborating
aerial photography was not obtained, the only positive identifications
were those which were made on animals visually noted from aircraft.
The deer in the pen at Junction Butte could not be distinguished
from
the vegetation in the original thermal imagery.
Although spots were
noted in the imagery at the positions in the pen recorded by ground
observers as deer locations, their presence would not have been detected
without prior knowledge.
The Jensen Creek site was spe.c Lf Lca Ll.y chosen for its topographic
uniformity and history of deer use. During an aerial check of the
sites about 6 weeks prior to the tests, an estimated 200 to 300 deer
were observed in the Jensen Creek area.
No deer were observed from
the aircraft in the Jensen Creek test site on the day of the mission,
January 26. Consequently,
the mission was aborted when half completed.
However, a small band of elk (Cervus canadensis) was seen just prior
to the beginning of flight lin~on
Jensen Creek, and they were
imaged by the scanner.
Air temperature during the Jensen Creek mission
was about -6°C and wind was less than 5 m.p.h.
Only two sightings of deer were made visually from the scanning aircraft on the Cedar Ridge site. At least 25 deer were ide:ntified in
the original imagery as verified by the actual sighting.
Numerous
cattle along the Colorado River were successfully
imaged by the
scanner at the end of several flight lines.
Again, SinCE! no photography was available to compare with the thermal imagery, there was
no way of knowing how many deer were actually detected by the scanner.
Many spots were apparent in the Cedar Ridge imagery which could have
been deer.
The drive census. by the Colorado Game, Fish and Parks
Division, in cooperation with Colorado State University wildlife
management students, on January 29, produced a count of 487 deer.
During the Cedar Ridge mission, air temperature was -5°C, and winds
were gusty at an estimated 10 to 20 m.p.h.
Considerable
distortion was apparent in the original thermal imagery
because of problems with the on-board film processing system.
This
accounted for the difficulty in identifying deer in this imagery.

�-262-

The video taped data recorded in-flight was taken to the NASA Manned
Spacecraft Center, Houston, Texas, where the equipment required for
optimum tape-to-film conversion eliminating distortion was available.
Additional film imagery was obtained which was better than the original
imagery in some respects, but inferior in others. Limited time precluded attempts at electronic imagery enhancement techniques which
may improve the imagery considerably, from the standpoint of seeing
deer on the film.
Additional work is planned for 1972, to apply these electronic
enhancement techniques to the taped data, and to attempt development
of automatic (computer) recognition procedures.

Aerial Color Infrared Photography for Deer Mortality
Detailed examination of both scales of photographs by two experienced
photointerpreters showed that the 1:4000 scale imagery was too small
to obtain reliable estimates of deer mortality. In many cases, the
carcasses were in such advanced stages of erosion or had been so severely
disturbed by scavangers, that potential identifiable image characteristics could not be detected in that scale of imagery. This was due
primarily to the fact that when the mission was flown in July, preceding
hot weather and scavangers had reduced many of the carcasses to
non-descript masses when viewed from a distance. Consequently, only
the 1:2000 scale photographs were used for the interpretation test.
A total of 47 carcasses were imaged in the photographs and used in the
interpretation for reasons previously explained. The number of correct
identifications ranged from 12 to 35 for an average of 57 percent
(Table 1). Commission errors, identifying an imaged object as a deer
carcass when in fact it was not, ranged from 0 to 19. However, with
two exceptions, interpreters C and F, commission errors were less than
17 percent. Errors of omission, not classifying any image as a peer
carcass, ranged from 25 percent to 75 percent.
Interpreter results can be grouped into two classes: interpreters A
through E and interpreters F through H. The first group of interpreters
represented individuals with considerable experience with CIR photographs. This group identified 68 percent of the imaged carcasses
correctly. The other group identified only 36 percent of the carcasses
correctly. This group represented the more cautious type of interpreter
requiring very positive evidence before calling an image a carcass.
For example, they apparently needed to see a leg, a head, or the absolute
body configuration in the photography before they would commit to an
interpretation.
The first group of interpreters used associated
evidences such as color and relative shape and position to other objects
in the scene to commit their interpretation of an image.

�-263-

Table 1.

Photointerpretation test results.

Photo interpreters
A

B

C

D

E

F

G

H

1/ Correct
(47 possible)

35

33

32

31

29

22

19

12

Error of
Commission

8

6

19

6

3

13

0

2

Total
Identified

43

39

51

37

32

35

19

14

Error of
Omission

12

14

15

16

18

25

28

35

�-264-

Commission and omission errors are believed to be related to three
hypotheses. Some of the carcasses were so severely decomposed that
even detection on the ground was difficult except by a very close
look. To compound this problem, the carcasses had been moved while
in a state of decomposition to the transect areas and this further
aggravated the disturbance problem.
Groups of rocks and piles of dead shrubby material also presented
confusion to the interpreters. Images of these objects closely
resembled piles of bones which were the de facto representation of
some carcasses. This problem can be minimized by planning photo
missions earlier in the year, preferably immediately after snow melt.
Bare, light-colored soil areas, frequent in some of the transect
areas, resulted in the most frequent misclassification of the deer
carcasses. In most cases, these areas were about the same relative
size as a carcass. Consequently, both commission and omission errors
~'ere made by classifying the bare soil areas as carcasses or not
c:lassifying them at all. These errors should be significantly reduced
by photo mission planning prior to serious decomposition.
Time to interpret the photography versus time to ground search for
carcasses in the three areas was used to develop an eatfmate of
lcelative efficiency comparing the two techniques. On the average,
,each interpreter required 6 hours or 3/4 man-days to interpret the
:photography from the three t ransect; areas. It required a four-man
ground search party 8 hours, or 4 man-days, to ground search the
same areas. This represents an approximate 5:1 efficiency ratio in
favor of CIR aerial photointerpretation for estimating deer mortality.
This does not include cost of the airplane, equipment, or flying
time. Such items could be prorated for other jobs for multi-purpose
mission planning.
The results of this segment objective were very encouraging considering
the constraints imposed by the photographs obtained. We will test the
hypothesis that estimates of deer mortality can be improved by obtaining
1:2000 CIR aerial photographs in May, 1972. This mission is planned.

LITERATURE CITED
Driscoll, R. S. 1969. Personal communication with N. L. Fritz,
Photographic Research Division, Research Laboratories, Eastman
Kodak Company, Rochester, New York.
Driscoll, R. S. 1971.
range management.
U p ,, iUus.

Color aerial photography--a new view for
U.S.D.A. Forest Servo Res. Pap. RM-67.

�-265-

Gilbert, P. F. and J. R. Grieb. 1957. Comparison of air and
ground deer counts in Colorado. J. Wildl. Manage. 21:33-27.
Gill, R. B. 1969. Middle Park deer study--population productivity
and mortality. Colo. Div. Game, Fish and Parks. Game Res.
Div. Fed. Aid Proj. W-38-R-23. Game Res. Rep. July. Part 1.
p. 105-122.
Heller, R. C. 1970. Imaging with photographic sensors. p. 35-72.
In Remote Sensing with Special Reference to Agriculture and
Forestry. Wash., D.C. Nat. Acad. Sci.
Wilson, R. A., S. N. Hirsch, F. H. Madden, and B. J. Losensky. 1971.
Airborne, infrared forest fire detection system; final report.
U.S.D.A. Forest Servo Res. Pap. INT-93. 102 p., illus.

~;f~~~'~~~
__
~~_~_~
'__
.

Prepared by__

_

~ S. Driscoll
Project Leader
Rocky Mountain Forest and Range Experiment Station

��-267-

July, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project Noo

W-38-R-26

Work Plan No.
Job Title
Period Covered:
Personnel:

15

Deer-Elk

Investigations

la
Job No.
Monitor Potentially Critical
Deer-Vehicle Accident Areas Statewide

April 1, 1971 through March 31, 1972

Thomas No Woodard, Dale F. Reed, Marion Lowery, Russ Mason,
Richard McDonald (and all Wildlife Conservation Officers in
his area), Larry Green, J. Kris Moser, Thomas M. Pojar and
William B. Zimmerman
0

ABSTRACT

The number and location of deer-vehicle accidents were monitored in five
general areas of Colorado.
Recommendations for installation of devices
to alleviate deer-vehicle accidents were submitted to the Division of
Highways where sample size (number of deer killed by vehicles) and other
conditions permit reasonable evaluation of the devices.

��-269-

MONITOR POTENTIALLY CRITICAL
DEER-VEHICLE ACCIDENT AREAS STATEWIDE
Thomas M. Pojar

P. S. OBJECTIVE
To establish statewide, the location and number of deer-vehicle
in areas appropriate to the evaluation of devices.

accidents

SEGMENT OBJECTIVES
1.

Examine potentially critical
management and as determined

deer-vehicle
by Job 1.

accident

areas reported

2.

As appropriate,

record all deer found killed

3.

As appropriate,

estimate

deer densities

in each selected

highway

4.

Measure deer activity along the roadside
selected highway area when feasible.

or in the median

of each

5.

Summarize the Highway Department's
highway area as needed.

in selected highway

traffic volume

by

area.
area.

data for each selected

METHODS AND MATERIALS
The locations of vehicle-killed deer were recorded to the nearest 0.1 mile
from some identifiable structure or land mark (such as bridges, roads or
prominent topographic features).
On the areas where quarter-mile section
markers have been installed, the kills were recorded in relation to these
markers.
Spotlight
conducted

counts, to estimate density of deer adjacent
using the method described by Reed (1969).

DESCRIPTION

Highway

to highways,

were

OF AREAS

6-24 (1-70) Rifle West

This area begins at the west edge of Rifle and extends to five miles west
of Grand Valley (18 miles total).
In general, this is used as a wintering
area by mule deer. Highway 6-24, a 2-lane highway, follows the Colorado
River Valley on the north side of the river.
The Book Cliffs are adjacent
to the Colorado River Valley to the north and Grand Mesa to the south. At
this writing the development of the interstate highway, which will replace
Highway 6-24, is in the preliminary planning and design stages.

�-270-

Highway

1-70 Eagle East

During this segment, highway construction crews completed the grading
of the 1-70 roadbed from Eagle to a point 8.5 miles east. The new roadbed follows the north side of the Eagle River Valley at the base of southfacing slopes.
The south-facing slopes are typically sagebrush and pinyonjuniper communities and are used as a wintering area by deer.
The fields
on the south side of the new roadbed are irrigated hay land.

1-70 Avon-Wolcott
A l4-mile segment of Interstate 70 beginning at the Eagle River bridge
2.5 miles east of the Avon interchange to 1.5 miles west of the Wolcott
interchange constitutes this study area. This segment of interstate was
completed and opened to traffic October 1, 1971.

Durango

Area

The Durango area includes Highway 550, 15 miles both north and south of
Durango, and Highway 160 from Mancos to Pagosa Springs.
A total of 118
miles of highway are included in this area, all of which is two-lane.
This general area is inhabited by deer year-long with some winter concentration and possibly some migration through the area.

Highway

82 Carbondale

East

This study area extends from the junction of Highways 133 and 82 near
Carbondale, southeast on Highway 82 for 1.75 miles to the Crystal Springs
Road.
The highway is adjacent to south-facing slopes that are covered
with either sagebrush or pinyon-juniper
communities.
This area is used by deer during midwinter.
south facing slopes are irrigated hayfields
in the fall and early spring.

Above and to the north of the
that are used heavily by deer

The highway is currently 2-lane but the Division
place this with 4-lanes during the next segment.

RESULTS

Highway

of Highways

plans to re-

AND DISCUSSION

6-24 (1-70) Rifle West

A total of 321 known road-killed deer have been documented on this l8-mile
segment of highway since November 1, 1969. The months of January, February
and March account for 75 percent of the kill, which indicates this area is
used principally as a wintering area by deer. Approximately
75-80 percent
of the total kills have occurred on two 3-mile and one 2-mile segment of
this highway.

�-271-

Coordination with the Division of Highways is continuing so provlslons
for reducing deer-vehicle accidents can be incorporated in the original
highway design.

Highway

1-70 Eagle East

There was no road-kill on this area because it was not completed and was
not open to vehicular traffic.
However, this situation permitted estimating
deer density adjacent to the roadbed and deer movement in relation to the
roadbed without the influence of vehicular traffic.
Mean monthly spotlight
count (Fig. 1), location of deer counted (Fig. 2), and location of deer
crossings based on track counts (Fig. 3) have been obtained for this area.
It appears that most of this segment of 1-70 is a potentially serious deervehicle accident area. However, until one year of data is gathered with
the highway complete and open to traffic, identification of critical areas
and actual magnitude of the problem cannot be determined.

Highway

1-70 Avon-Wolcott

Three areas of this l4-mile segment of interstate
deer kill areas.

have proven to be critical

The first area, from the Avon interchange east to the Eagle River bridge
(quarter-mile section number 1-9) is 2.25 miles long and has sustained a
deer kill of 18 animals since October 1, 1971. Installation of 8-foot
fencing on the north side of the highway for this distance has been recommended to the Division of Highways.
The second area extends from the Edwards interchange westerly for 2.25 miles
(quarter-mile section number 25-34).
Fourteen deer have been killed in this
area since October 1, 1971. Recommendations
for 8-foot fencing on the north
side of the highway have been submitted to the Division of Highways.
The third critical area begins 0.5 miles west of the Eagle River bridge east
of Wolcott and extends one mile in a westerly direction towards Wolcott.
Fifteen deer have been killed on this segment of highway.
The exact nature
of the deer movement on and across this segment is not fully understood, but
it is suspected that this is both a wintering area and migratory route.
No
devices have yet been recommended for this area because further information
is needed on the deer movement patterns.

Durango Area
The number and location of road-killed deer have been documented for one
year. A total of 446 deer and 29 elk have been killed on the 118 miles
of highway that constitute the Durango area. The kill is spread fairly
evenly over the entire area with few real concentration areas. Unless
devices that do not require highway construction to install are deemed

�-272-

feasible, recommendations
construction is proposed.

will be limited to those areas where highway

Highway

82 - Carbondale

East

Road kill and spotlight count data have been collected on this segment
of highway since 1968. A 5-year total of 98 deer kills have been recorded.
This segment of highway is scheduled for conversion to 4-lanes during the
next segment.
Recommendations have been submitted to the Division of Highways to include 8-foot fencing along the north side of the highway in the
construction project.

LITERATURE

CITED

Reed, Dale F. 1969. Techniques for determining potentially critical deer
highway crossings.
Game Information Leaflet No. 73. Colo. Dept.
Natural Resources, Div. of Game, Fish and Parks.
3 p.

/?
Prepared

j

by -c,;~/,~~/,""i.c-:~,-,'
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...•...
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If}/)
. "',
Thomas M. Poj ar
'J
Asst. Wildlife Researcher
n,,· .•··j

�-273-

100~----------------------~

80

-

78.3
~

IZ
;:)

860 -

w

50.5
~

(!)

«

:5 40 -

4~0
3~3

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. DEe

JAN

FEB

MAR

APR

Fig. 1. The mean number of deer counted per month adjacent
to 1-70 from 1.15 miles west of Wolcott Interchange to Eagle
(Dec. 1971 - Apr. 1972).

�~140

·

...-

0

~ 120 ·

-

&gt;

0::100 ·
w
(f)
CD 80

r--

r--

0

ffiw 60

I

- -

c 40
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~ 20
0

••••

r-

r--

r--

r-~

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r--

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10
15
20
25
QUARTER MILE SECTIONS

N

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Fig. 2. Total number of deer observations adjacent to 1-70 roadbed from 1.15 miles west
of Wolcott Interchange to Eagle (Dec. 1971 - Apr. 1972).

35

�-275-

440

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..

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~200
en

en

o

,......r--

-

a:: 150 ..
u
..

r--

ri--

r--

r--

I--

.. r-

r-

t--

100 ..

..
50 ..

I--

I--

r-f--

r-f--

n5

~

,......

!--

u
10

15

QUARTER MILE

20

25

30

SECTIONS

Fig. 3. Total number of deer crossings counted from 1.15 miles west
of Wolcott Interchange to Eagle during winter of 1971-1972.
Based on
track counts in November, 1971 (n=3); December, 1971 (n=12); January,
1972 (n=3); and February, 1972 (n=4).

35

��-277July, 1972

JOB PROGRESS REPORT

COLORADO

State of

Work Plan No.
Job Title

15

Job No.
Investigation

Period Covered:
Personnel:

Deer-Elk

W-38-R-26

Project No.

Investigations
3

of One-Way Deer Structures

April 1, 1971 through March 31, 1972

Thomas N. Woodard,
Dale F. Reed.

William

B. Zimmerman,

Thomas M. Pojar and

ABSTRACT
A number of one-way gates that permit deer to move through deer-proof fences
from only one direction were evaluated under field conditions adjacent to
Interstate 70 near Vail and adjacent to Highway 82 south of Glenwood Springs.
A total of 413 passages have occurred through thirteen one-way gates under the
varying field conditions.
Ninety-seven percent of all field passages were
positive (in the direction of the funneling or cone principle).

��-279-

INVESTIGATION

OF ONE-WAY DEER STRUCTURES
Dale F. Reed

P. S. OBJECTIVE
Evaluate the effectiveness of several types of one-way gates and two
fence structures to permit one-way movement of mule deer through existing deer-proof fences at Little Hills Experiment Station and elsewhere.

SEGMENT OBJECTIVE
1.

Test those structures deemed effective under controlled
for effectiveness under field conditions.

conditions

METHODS AND MATERIALS

Testing Under Field Conditions
The one-way gate deemed most effective under controlled conditions was
further tested in the field. Three of these one-way gates were evaluated
in a quarter mile of 8-foot fence adjacent to Highway 82 and eight in the
8-foot fences associated with the deer underpass near Vail.
Four of the
eight one-way gates at Vail were in off-sets (Reed 1971) in the south 8foot fence, while the others were strategically located at fence junctions
or areas where deer have frequented the fence on the wrong side.
In
addition, two one-way deer gates were installed in a 1.1 mile section of
Uni-strut 8-foot fence adjacent to Highway 82.

RESULTS AND DISCUSSION

Field Conditions
Of the l3-one way gates being evaluated under field conditions, the eight
located adjacent to Interstate 70 and the deer underpass in the Vail area
received the greatest use. However, some of these gates received many more
passages than others.
For instance, gate 70N-S has received an accumulation
of 261 passages while 70S-II has received only six (Table 1). Gate 70N-S
was located in a "V" junction near good deer cover approximately one quarter
of a mile from the highway, and was used by resident animals that moved
around the end of a wing fence during the summer months.
On the other hand,
gate 70S-II was located near the highway with little cover and could be used
only by animals escaping the immediate highway right-of-way.

�-280-

Table 1. The accumulated number of one-way deer gate passages, percent
of positive passages, and number of tracks on the negative side when no
passages occurred during 1970 and 1971 field tests.

Gate
.
D.
eSlgnatlon

1/

-

Passages during
field tests
No.
No.
Po s ,
Neg.

Percent
Positive

No. of Tracks
on Neg. Side
When No Passage

Type Fence
Installation
and Comment

82-1

2

o

100

o

Straight fence
line &amp; trail

82-2

3

-1

75.0

o

Off-set

82-3

2

o

100

o

Off-set

70N-4

76

-4

95.0

504

"v" junction
(R-O-W) ]j

"v" junction
wing fence

70N-5

261

-2

99.2

70N-6

21

o

100

Straight fence
line (R-O-W)

70N-7

9

-9

50.0

Fence line/changed
in '71 to half offset (R-O-W) 1/

70S-8

9

-1

90.0

322

70S-9

10

o

100

192

70S-l0

14

100

441

103

)

-,

Off-set

(R-O-W)

Off-set

(R-Q-W)

70S-11

6

o
o

82-12

o

o

12

Half off-set

82-13

'0

o

3

Half off-set

Total

413

-17

~
100

97.1

in

754

2331

1/

- The first number designates the highway adjacent to the gate location, the
N (north) or S (south) the side of the highway, and the last number the sequential number of the gate installed.

J:../"v" junction

refers to two 8-foot fences which join to form an apex or funnel
effect.
Deer tend to move along the fences to the apex. R-O-W refers to the
gate being located off the highway right-of-way.

l/Half

off-set

refers to half of the off-set previously

described

(Reed 1971).

�-281-

The effectiveness of the gates in terms of "one-way passages" is indicated
by the few negative passages that occurred.
Most of the negative passages
occurring during 1971 (against the funneling or cone principle) resulted
from human interference (leaving gate open). Gate 70N-7 received so much
human use and interference that it was decided to relocate the structure
to a different area of the 8-foot fence. Another indicator of the "oneway passage" characteristic was the number of tracks counted on the negative side when neither positive or negative passages occurred.
A total
of 2331 tracks indicated activity on the negative sides of the gates where
negative passages did not occur. These gates were effective in preventing
deer from getting on the 4-lane Interstate Highway while providing a means
of escape for deer that were trapped on the highway side of the fence.
According to the track counts an estimate of 145 deer have escaped the
highway rights-of-way since the structures were installed.
One-way gates 82-1 through 82-3, located in a O.25-mile fence section
adjacent to Highway 82 south of Glenwood Springs, received few passages.
The number of deer in the immediate area has been low during the last three
years (1969 through 1971). Gates 82-12 and 82-13 were installed in a 1.1
mile fence section also adjacent to Highway 82. These gates received no
passages although several deer moved along the highway side of the 8-foot
fence.

LITERATURE

CITED

Reed, Dale F. 1971. Investigation of One-Way Deer Structures.
324. In Game Research Report, July, Part 3, Federal Aid.
Div. Game, Fish and Parks.

--~Prepared

by

i

·--/0 /'

)~i{lf!-~et\ ,Sa
Asst. Wildlife

(

Researcher

p. 323Colo.

��-283-

July, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W_-~3_8_-_R_-_2_6 _

Work Plan No.

Investigations
6

15
Job No.
----------~~---------

Job Title

Deer Underpass

Period Covered:
Personnel:

Deer-Elk

Evaluation

April 1, 1971 through March 31, 1972

J. Kris Moser, Thomas M. Pojar, Thomas N. Woodard,
Zimmerman and Dale F. Reed.

William

B.

ABSTRACT

The use of a concrete box deer underpass 10 feet by 10 feet and 100 feet long
that permits deer to move under Interstate 70 to and from their summer range
has been evaluated.
The structure is located 4.3 miles west of Vail and utilizes a well-established
deer migration trail and a natural drainage referred to
as Mud Springs Gulch. According to the number of passages recorded by the
electro-optical detection system, an increase of 31 percent occurred from 1970
to 1971. The greatest use of the underpass occurred during the June and the
October-November periods with maximums of 16 and 28 deer per days, respectively.
An index of entrance versus exit activity has indicated a behavioral reluctance
on the part of the animals to use the underpass.

��-285-

DEER UNDERPASS

EVALUATION

Dale F. Reed

There are many areas in Colorado where migrating deer cross highways.
When these highways carry high speed, high-volume traffic, the potential
danger to deer and motorists becomes especially great.
Therefore, it is
essential to keep the animals off the highway.
Underpasses with specified dimensions and characteristics located at strategic points within
migration routes may provide adequate means for these necessary deer movements.
This study purports to evaluate the use of a specific concrete
deer underpass 10 feet by 10 feet and 100 feet long located in the Vail
area.

P. S. OBJECTIVE
Determine if deer migrating from winter range on one side of Interstate
70 to summer range on the opposite side utilize an underpass constructed
in the area.

SEGMENT OBJECTIVE
Measure

deer use of the underpass.

METHODS

AND MATERIALS

A nighttime lighting system and an electro-optical detection system with
two Streeter Amet recorders (previously described by Myers 1969, and Reed
1970) were maintained at the underpass.
During the spring migration of
1970 and 1971, and during the fall migration of 1971, the detection system
was activated at both the entrance (north end) and the exit (south end) of
the underpass.
However, during the fall migration of 1970 only the exit
(north end) detection system was used.
Before the spring migration, dirt was spread over two-thirds of the floor
width of the underpass for the entire length.
This raised dirt-covered
portion was used as a track bed.
The track bed and entrance to the underpass were checked periodically for
deer tracks.
As deer activity increased and when deer began using the
underpass, the counters and the number of deer tracks were checked and
recorded daily.
The number of deer passing through the underpass was
estimated by carefully examining the number of tracks, and comparing them
with the counter's reading at the exit. The counter's reading on the
entrance was also recorded and compared with that at the exit. During
each check the detection system was reset at zero and the tracks raked out
by hand.

�-286-

During the 1971 fall migration a weather station housing containing a
hygrothermograph
and maximum and minimum thermometers was installed,
along with a Belfort rain gage, anemometer, and snow depth measuring
stake.
In addition, acoustical housings were constructed and installed
for the two Streeter Amet recorders.

RESULTS AND DISCUSSION

Spring and Summer Use of the Underpass
It was estimated that 227 deer moved south through the underpass during
the spring and summer migration of 1971. The first passage, south toward higher summer range, occurred on May 18. The greatest use of the
underpass occurred during June with an estimated maximum of 16 deer
passing through during one night •• Use of the underpass slackened during
July as it had during the spring and summer of 1970 (Fig. 1). Compared
with 1970, the number of passages during the spring and summer of 1971
increased by 44.6 percent.
This may have been caused by an increase in
the population or a shift in the migration pattern.
In addition, an estimated 42 negative passages (north toward winter
range) occurred during July through September when relatively dry conditions existed.
Some animals appeared to be passing through the underpass in both directions on a nightly basis.
It was hypothesized that
resident deer may have used Gore Creek as a water source.
An index activity was determined by dividing the daily counter reading
at the entrance of the underpass by that at the exit. This index (Fig.
2) illustrates an apparent reluctance on the part of deer to use the
underpass.
This index of activity would equal 1.0, if each deer that
came to the entrance went through and was counted once at the entrance
and once at the exit. The greater the index, the greater the number of
counts at the entrance as compared with the exit, and the greater the
amount of time and energy expended there by deer. Observations, made
from some distance away, bear out the hypothesis that most groups of
deer spend a great deal of time at the entrance before, and possibly
without, passing through the structure.
This provides a possible
explanation for the sharp peaks represented in Fig. 2.

Fall Use of the Underpass
It was estimated that 367 deer moved north through the underpass during
the fall migration of 1971. The first passage recorded for the fall
migration occurred on October 2 and the last on December 18. However,
passages in both directions occurred during September.
The greatest
use of the underpass occurred during October and November with a mean
of 10.6 deer passages per day during the first week of November (Fig.
1). A maximum of 28 deer passed through the underpass during one night.

�-287-

MEAN

PASSAGES

so

CD

MAY 15-3 1

0

-- \

\

JUN 16-31

./'

./

./'
./
./

/

,,

JUL 16-31

/
/

w W

/

~

~

0

I
/

0

l&gt;

-&lt;

/

AUG16-31

en

SEP 16-

OCT 16-31

//

--

--- --

---:::-.

/
/
".,/

.,;

DEC 16-31
Fig. 1. The mean number of passages per day is represented for each
2-week period for both the spring-summer
and fall migrations
for 1970
and 1971.

�50------------------------------------------~
1970
197 I ------

A

,

&gt;-

,
I

,,
,,I,,,
,,
, I

••••

I

"

&gt;
••••
o

I

&lt;t

o

A

/1

II
! I

X
W
C

,~ ,, II

'

I I:

\

,I
I'

, ,,

, I'

I

\

Z

, I

1\
I, '
II ,

u,

,I AI'"

I

10

1'1
,I ,
t I,

I

I J
\,..-/

\

,~ I,

I

\
\

I' \

\

00
00

I
I

I ~

,: Il',~
\ ,, . "I,

I

I

I
N

\

\I

I

0
4

8

12

CD

ro

(\J

UJ
Z

&gt;c:t

::E

~

-:&gt;

32

36 40 44 48

ex&gt;

CD

v

(\J

Z

Z

UJ
Z

~

-:&gt;

-:&gt;

~
~

-:&gt;

16 20
0
UJ
~

24

UJ
~

28

(\J

~

&gt;...J

-:&gt;

-:&gt;

52

56
0
(\J

&gt;...J
~

-:&gt;

Fig. 2. An index of activity, determined by dividing the daily counter readings at the entrance of
the underpass by that at the exit, indicates a reluctance on the part of deer to use the underpass during the spring and summer of 19?O and 1971.

�-289-

The estimate of 367 passages during the fall is an increase of 61.7
percent over that of the 1970 spring-summer use. The reasons for such
an increase are not known.
However, herd increment, a shift in overall migration patterns, and the lower hunter harvest are probable
factors.
The variation in seasonal use of the underpass occurring
between 1970 and 1971 fall migrations (Fig. 1) may be due to weather
conditions, primarily the timeliness and severity of snowfalls.
Fifteen snowfalls were recorded during October and November of the 1971
fall migration period.
The mean temperatures (F) when passages occurred
during October, November, and December were 28.0, 21.0, and 10.3, respectively.
One hundred nineteen deer were recorded moving north through
the underpass during clear weather (Table 1). In addition 120 deer were
recorded moving north through the structure during snowfalls or within
24 hours following snowfall cessation.
Additional weather collection
and data analysis will be conducted during the next segment.

Table 1. The number of passages through
November, and December, 1971 in relation
conditions.

the underpass during October,
to snowfalls and weather

In Relation

to Snow:

Oct.

Nov.

Dec.]/

Total

Following

snow 1/

33

73

14

120

26

140

14

180

Not following

snow ]j

Unknown

0

67

Total

367

In Relation

to Weather

Conditions:

Cloudy and snowing

18

14

11

43

Cloudy to partly cloudy

6

18

5

29

Clear

20

88

11

119

Clear and moonlight

4

79

0

83

Unknown

14

93

Total
l/Within

367
snowfall

period or 24 hours following

cessation.

]j Not during or&gt; 2lf hours following cessation of snows.
3/

- Data only from December

1-10.

�··290-

The index of entrance versus exit activity for the 1971 fall migration
was much lower than the spring-summer indexes previously discussed.
A comparison by year will be made during the next segment.

LITERATURE CITED
Myers, Gary T. 1969. Development of an electronic deer counting
device. p. 235-247. In Game Research Report, July, Part 2.
Colo. Game, Fish and Parks Div. Federal Aid.
Reed, Dale F. 1971. Deer underpass evaluation. p. 341-351.
In
Game Research Report, July, Part 3. Colo. Game, Fish and Parks
Div. Federal Aid.

Prepared

�-291July, 1972

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~---------

Work Plan Noo
Job Title
Period Covered:
Personnel:

Deer-Elk

W-38-R-26

Project No.

15

Investigations

7
Job Noo
Effects of Highway Lighting
on Number of Deer Killed by Vehicles

April 1, 1971 through March 31, 1972

William B. Zimmerman, Gary T. Myers, Thomas Mo Pojar, Dale Fo Reed
and Thomas N. Woodard.

ABSTRACT

��-293-

EFFECTS OF HIGHWAY LIGHTING
ON NUMBER OF DEER KILLED BY VEHICLES
Dale F. Reed

P. S. OBJECTIVE
Determine if highway lighting
on a portion of Colorado 82.

affects

the rate of deer-vehicle

accidents

SEGMENT OBJECTIVES
1.

Record the number of deer killed by vehicles at each end of, and within a 3/4 mile lighted portion of Colorado 82.

2.

Estimate deer densities
by lighting.

3.

Compare pre-treatment
count data.

4.

Gather additional information which may be needed to understand
deer-auto accident rates are or are not affected by lighting.

adjacent

to the portion

of highway

kill and count data to post-treatment

RESULTS

affected

kill and

why

AND DISCUSSION

A joint decision was made by the Colorado Division of Highways and Game,
Fish and Parks Division to delay installation of the illuminaries.
The
attached appendix discusses the number of deer and the number of deervehicle accidents in the study area, and the recommendations
for installation.

Prepared

byCZAi14-1c':~Q
Dale F. Reed
Asst. Wildlife

Researcher

�-294-

APPENDIX

A

RESEARCH
1972 Deer-Highway

Lighting

Project Report

Since the deer-highway lighting project's approval on September 11, 1969,
the installation of the high intensity illuminaries on Highway 82, south
of Glenwood Springs, has been delayed several times. The number of deer
and the number of deer-vehicle accidents in the Jammaron study area has
shown a marked decline since 1970 (Figs. 1 and 2). However, during the
1972 period, the number of deer-vehicle accidents (deer kill) was relatively high in January (Fig. 1) but declined during February and March.
Weather continued to be suspect in terms of influencing the movement of
deer toward the study area.
Snow depths adjacent to and above the study
area approximated 14.0, 13.2, 7.0, and
inches for December, January,
February, and March, respectively.
Hence, the relatively moderate to
severe winter conditions in December diminished during mild weather in
January, February, and March.
The average minimum temperature for
January through March increased in 1972 (Fig. 3), while the total amount
of precipitation declined to 1.36 inches.

°

Recommendations
A cooperative Deer-vehicle Accident Research Proposal submitted to the
Highway Division in May, 1972 included a time schedule for the highway
lighting project.
A request for approximately 0.25 miles of lighting
will be submitted

in January,

1973.

.~.

'&gt;?(J;

.'

~- ./
~A-

~I(J:t_ (j

\~./

,......

-.

/
.,

Dale F. Reed
Assistant Wildlife

Researcher
. ~.r )

Thomas M. Pojar
Assistant Wildlife
COLORADO

Researcher

DIV. OF GAME, FISH AND PARKS

�HIGHWAY 82 ROAD KILL DATA
(JAMMARON AREA)

30~----------------~-------------------------------'
025
w
..J
..J

&lt;,

~

&lt;,

&lt;,

0::: 20

w
w

o

&lt;,

/

lL..

o 15

\OJ~

0:::'

&lt;,

I

tv
'-D

&lt;,

-&gt;

W

rn

~"l

~

~ 10

00

•

00

0

o~

0
000

..J

00000

0

o •
•

-----

------ ----

---- --------

\91&lt;L---

--

JAN
fig. 1.

0

o

-- --

&lt;,

00.

o

'97'

&lt;,

......... .........

/

/

U1
I

1972

/

~
~

&lt;,

Number of deer killed by vehicles

------_
-------_---

--

0

o

0

o

0

o

··0.
--=......•.•.~~:....~
-- ..-0

~,.':"---00

study area south of Glenwood

00

MAR

FEB
on the Jammaron

_

Springs.

�HIGHWAY 82 SPOTLIGHT COUNT DATA
(JAMMARON AREA)

IJ...

I

o

N

-0

a::

-------

lLJ

CO

~

~

\9t;l!J~ ~

::&gt;

z-:

z

0\
I

--------

~~
~
~

1972

- -.-

... ........ .... .....•. ..•....
. ..... -..
.
..
0
~.. . . . . . .. .. .. .
\9~ - - .....
- 1971 ---_----...
-- ---- ------.------- -- -....-.::: .",. ."...:-_-------JAN
Fig.

---

2.

-

-

Number of deer counted adjacent

-- - - - --------- --J~~

FEB
to the Jammaron

study area south of Glenwood

_

••.,. ••••••

....

MAR
Springs.

�HIGHWAY 82

PRECIPITATION AND TEMPERATURE

JANUARY, FEBRUARY, AND MARCH
~
(J)

-PRECIPITATION
---TEMPERATURE

LaJ

::J:
o

-

~2PO

o

-

=&gt;0

//

••••••

//

«
0-

I-

16

///

U

zLaJ

i~

I

(Do::
«LaJ

//

12

.//

10

ffi&gt;LaJ~

« I-

~

g
1969

1970

1971

I

tv
\D

LaJ~

/

LaJ

0...J

~-

/._--------_._---------_.

z

0::

20 :i.-.
u,

1972

Fig. 3. Average minimum temperature and total precipitation for the Jammaron study area south
of Glenwood Springs during January, February, and March.

-....J

��-299July, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-26

Work Plan No.

Personnel:

Investigations

15

Job Noo
8
Effect of Lighted Deer Crossing
Signs on Number of Deer Killed by Vehicles

Job Title
Period Covered:

Deer-Elk

April

_

1, 1971 through March 31, 1972

Larry Green, Thomas N. Woodard,
and Thomas M. Pojar.

Dale F. Reed, William

B. Zimmerman

ABSTRACT
Lighted, animated deer crossing signs were tested to determine their effect
on the rate of deer-vehicle accidents and their effect on the speed of traffico
Vehicle speeds were monitored at three distances behind the signs.
The differences in mean vehicular speed was 2 88, 1042 and 1016 miles per hour at 0.15,
0065, and 1.50 miles behind the signs, respectively.
The mean speeds were
significantly (P&lt;0005)
lower at all three stations with the signs on. The
number of estimated crossings per kill increased 43.6 percent with the signs
on, compared with the estimated crossings per kill with the signs offo However,
this was not a significant difference (P)O.50)
in ratios according to chisquare analysiso
0

��-301-

EFFECT OF LIGHTED DEER CROSSING
SIGNS ON NUMBER OF DEER KILLED BY VEHICLES
Thomas M. Pojar

P. S. OBJECTIVE
Determine
accidents

if lighted deer crossing
on a portion of Colorado

signs affect the rate of deer-vehicle
82.

SEGMENT OBJECTIVES
1.

Record the number of deer killed by vehicles within
highway marked by lighted deer crossing signs.

2.

Estimate

3.

Compare pre-treatment
count data.

4.

Gather additional information which may be needed to understand why
deer-auto accident rates are or are not affected by lighted deer
crossing signs.

deer densities

adjacent

a mile portion

to the signed portion

of the highway.

kill and count data to post-treatment

METHODS

of

kill and

AND MATERIALS

In early April, 1971, two lighted, animated deer crossing signs were installed on the same Highway 82 study area where lighted deer crossing signs
were evaluated previously (Pojar 1971). The animated deer crossing signs
used in this study (Fig. 1) were described by Pojar, Reseigh and Reed (1972).
Estimates of the number of deer crossing the highway were made by making
nightly spotlight counts.
The total number of deer seen in the fields
southwest of the highway was multiplied by two, assuming that each deer in
the field had crossed the highway once and would cross again that same night
to return to the pinyon-juniper cover on the northeast side of the highway.
For every deer killed on the highway, one crossing was arbitrarily added to
the total crossings for that night.
The deer crossing signs were activated and turned toward traffic for a I-week
period, then deactivated and turned away from traffic the following week.
This routine of alternating on a weekly basis the treatment and pre-treatment
effect of the signs on both the deer kill and the speed of traffic was continued until the number of deer in the area diminished to near zero.
Vehicle speeds were recorded with an automatic vehicle speed recorder at
stations located 0.15, 0.65, and 1.50 miles behind the signs to monitor the
effect of the signs on the speed of traffic.
The speeds of nearly all the
vehicles that passed the speed stations between 6:00 and 10:00 P.M. were recorded.
Eighty vehicle speeds were randomly selected from this sample for
tabulations.

�-302-

Fig. 1. The animated deer crossing sign shows deer silhouettes
in sequence from right to left (photo by Don Domenick).

lighted

�-303-

RESULTS AND DISCUSSION
Deer occupied the study area for four weeks, allowing two I-week periods
of data collection with the signs off and two I-week periods with the
signs on. The number of estimated crossings per kill increased 43.6
percent with the signs on compared with the estimated crossings per kill
with the signs off (Table 1). According to chi-square analysis there is
no significant difference (P&gt; 0.50) in the ratios.

Table 1. The estimated deer crossings and total kill on Highway 82 study
area during two I-week periods with the animated deer crossing sign off
and two I-week periods with the sign on.

Estimated

crossings

Sign Off

Sign On

227

163

6

3

37.8

54.3

Total kill
Crossings/kill

The results of the vehicular speed evaluation are presented in Fig. 2. The
mean speeds were significantly (P&lt;0.05) lower at all three speed stations
with the signs on. The differences in mean vehicular speed was 2.88, 1.42,
and 1.16 miles per hour at 0.15, 0.65, and 1.50 miles behind the sign,
respectively.

LITERATURE

CITED

Pojar, Thomas M. 1971. Effect of lighted deer crossing signs on number
of deer killed by vehicles.
p. 359-366.
In Game Research Report, July,
Part 3. Colo. Game, Fish and Parks Div. Federal Aid.
Pojar, Thomas M., T. C. Reseigh, and Dale F. Reed. 1972. "Deer Crossing"
signs may prove valuable in reducing accidents and animal deaths.
Highway Research News, Winter.
No. 46:20-23.

Prepared

by __ i_,",_""_" c:_~/;~1_./---"-_"_"/.~:i.:
_
Thomas M. Pojar
Asst. Wildlife Researcher

�58

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:I:
a.. 57
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I I I I I I I I I I I I I I I I I I I

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15

(1.50 MILES)

(0.65 MILES)

(0.15 MILES)

5

o

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53

SPEEDS 1972

•
••

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~

VEHICLE

I I I I I

20

25

0

5

10

o

sign.

Each circle or dot

5

DAYS
Fig. 2. Vehicular speed at three distances behind
represents a mean of 80 randomly selected speeds.

the animated

deer crossing

10

�-305July,

JOB PROGRESS

State of

~C~O~L~O~RA==D~O

REPORT

_
Deer-Elk

W-38-R-26

Project No.

1972

Investigations

10
Job No.
An Evaluation of Deer-proof Fence Length
Job Title __~R~e~q~u~~~'r~e~d~t~o~P~r~e~v~e=n~t~D~e~e~r~M~o~v~e=m~e~n~t~o~n~o~r~A~c~r~o~s~s~H=i~g~
_

Work Plan Noo

Period

Covered:

Personnel:

15

April 1, 1971 through March 31, 1972

Thomas N. Woodard, Dale F. Reed, Larry Green, William
Thomas M. Pojar, J. Kris Moser and Russ Mason.

B. Zimmerman,

ABSTRACT

A 1.1 mile 8-foot fence was erected along Highway 82 near the Diamond S
Ranch to prevent deer movement across the 4-lane highway.
Prior to installation of the 8-foot fence, the mean number of deer crossings per 24 hour
period during March-May 1971 was 12.8 (n=29) with two deer killed on the
highway and a mean spotlight count of 47 6 (n==ll) In the vicinity of the
V:lil underpass-fencing
complex, sightings of marked deer were used to monitor
their movements in relation to the fenceo
0

0

��-307-

AN EVALUATION OF DEER-PROOF
FENCE LENGTH REQUIRED TO PREVENT DEER
MOVEMENTS ON OR ACROSS HIGH SPEED HIGHWAYS
Thomas M. Pojar

P. S. OBJECTIVE
To evaluate length of deer-proof
or across high speed highways.

fence in relation

to deer movement

on

SEGMENT OBJECTIVES
1.

Install deer-proof fence along appropriate
determined by Job la.

2.

Evaluate the effectiveness of various
in affecting deer movements.

METHODS

sections

lengths

of highway

of deer-proof

as

fence

AND MATERIALS

Track counts in the median of the 4-lane highway were used to estimate
the number and location of deer crossings.
Conditions were normally
favorable for maintenance of soil track beds during March, April, and
May.
During winter, when the soil was frozen, track counts were made
in the snow.
Deer density adjacent to the highway was estimated using the method
described by Reed (1969) and the locations of all vehicle-killed
deer
were documented in relation to quarter-mile section markers.
Location
and length of the 8-foot fence was determined by the concentration of
kills and/or crossings.
Deer were marked with numbered neck bands or automatic tagging devices
(Siglin 1966), both in the Vail underpass-fencing
complex (Reed 1971)
and on the winter range of the migratory Vail deer herd. A drop-net and
automatic tagging devices were used during fall and spring when deer
frequented areas behind the 8-foot fence that leads to the underpass.
Clover traps were used on the winter range.
Sightings of marked deer in
the vicinity of the 8-foot fence were used to monitor their movements in
relation to the fence.

DESCRIPTION

Highway

OF AREA

82 - Diamond

S

The Highway 82 study area has been described by Pojar (1971).
The segment
of Highway 82 near the Diamond S Ranch is 1.5 miles long (quarter-mile

��-309-

sections 24-30).
A field of crested wheatgrass (Agropyron desertorum)
is located on the east side of the highway and is one of the earliest
grasses to green up in the spring (Reynolds and Springfield 1953).
Sagebrush is abundant around the perimeter of the crested wheatgrass
field and is replaced by pinyon-juniper type as elevation increases to
the east. Alfalfa fields line the west side of the highway.
Deer concentrate in the crested wheatgrass field and sagebrush
the highway in late winter and early spring.
The Vail underpass-fencing
Reed (1971).

complex and adjoining

east of

area is described

by

RESULTS AND DISCUSSION

Highway

82 -Diamond

S

The existing 42" Uni-strut telespar fence along the east highway rightof-way boundary was extended in height to eight feet during October 1971.
The 8-foot fence is 1.1 miles long and is roughly centered on the quartermile section where the greatest number of deer kill and crossings occur.
A soil track bed was established in the median of the 4-lane highway with
a 6-foot wide spring tooth harrow.
This track bed extended past the ends
of the 8-foot fence so that deer coming around the ends of the fence and
across the highway could be detected.
In addition, small track beds about
5 feet wide by 10 feet long were established at each end of the fence.
Mean crossings between quarter mile section markers 25 to 30 during MarchMay 1971 (before 8-foot fence was installed) was 12.8 (n=28) with two deer
killed on the highway and a mean spotlight count of 47.6 (n=ll).
The comparable 1972 data are currently being obtained.
Although extensive track count data (crossings) was not gathered prior to
1971, accurate kill and spotlight count information is available since
1968. The March-May mean number of deer observed on spotlight counts and
the number of road-killed deer between quarter-mile section markers 25 to
30 are: 1968, 47.8 (n=13) and 1 kill; 1969, 77.4 (n=lO) and 5 kill; and
1970, 63.7 (n=ll and 8 kill.

Highway

1-70 - Vail Area

A total of 39 deer have been marked.
Twenty-three were marked on winter
range and 16 were originally marked near the Vail underpass-fencing
complex.
Preliminary results on the movements of four deer in relation to the 8-foot
fence are available at this time. Deer number l66b traveled north perpendicular to the north Vail fence a minimum of 1.5 miles.
Deer number 70b remained in the vicinity of the fence for a minimum of five days and traveled
parallel to the fence a minimum of 0.5 miles.
Deer number 22 and 190b
traveled parallel to the fence 0.4 and 0.7 miles, respectively.

�-310-

LITERATURE

CITED

Pojar, Thomas M. 1971. Deer-auto accident investigations. p. 275-318.
In Game Research Report.
July, Part 3. Colo. Game, Fish and Parks
Division.
Federal Aid.
Reed, Dale F. 1969. Techniques for determining potentially critical
deer highway crossings.
Game Information Leaflet No. 73. Colo.
Dept. of Nat. Res., Div. of Game, Fish and Parks.
3 p.

1971.
Report.

Deer underpass evaluation.
p. 341-351. In Game Research
July, Part 3. Colorado Game, Fish and Parks Div. Federal Aid.

Reynolds, H. G., and H. W. Springfield.
1953. Reseeding southwestern
range lands with crested wheatgrass.
Farmers' Bulletin No. 2056.
U. S. Department of Agriculture.
U. S. Government Printing Office.

20 p.
Siglin, Roger J. 1966. Marking mule deer with an automatic
J. Wildl. Mgmt.
30(3):631-633.

/
Prepared

.z:-,-_..',,-;

~i;.....&gt;_·:'+i
by -"(,-"/I",,,-'~-,-~t,,;,,;:"-"'k,,,-I!.:....~_·
Thomas M.I Po j ar .;
Asst. Wildlife Researcher

-

tagging device.

�-311July, 1972

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~------------

Work Plan No.
Job Title

Investigations

11
Job No.
Effects ~f 8-foot Fence Angle
~i~n~D~i~v~e~r~t~1~·n~g~D~e~e~r~~f~r~o~m~t~h~e~i~r~E~s~t~a~b~l~i~s~h~e~d~D~.~ir~e~c~

Period Covered:
Personnel:

Deer-Elk

W-38-R-26

Project No.

15

April

1, 1971 through March 31, 1972

Thomas Mo Pojar, William
Dale Fo Reed.

B. Zimmerman,

ABSTRACT

Thomas N. Woodard

and

��-313-

EFFECTS OF 8-FOOT FENCE ANGLE IN DIVERTING
DEER FROM THEIR ESTABLISHED DIRECTION OF MOVEMENT
Dale F. Reed

P. S. OBJECTIVE
Evaluate the effectiveness of three angles of 8-foot fence to divert
deer from their established direction of movement at Little Hills
Experiment Station and elsewhere.

SEGMENT OBJECTIVES
1.

Design and construct a movable
closure at Little Hills.

8-foot fence section

in a deer en-

2.

Test the effectiveness

3.

Evaluate those angles deemed effective under controlled
for effectiveness under field conditions.

of each angle under controlled

conditions.
conditions

RESULTS AND DISCUSSION
A decision was made to modify the Segment 26 Job Description to include
only simulated fence angles tested under field conditions.
The photo
detectors (DSI Type 1050) necessary for the original plan failed to
function adequately under low light and morning light conditions.
The
modified plans will be implemented during the next segment.

Prepared
Asst. Wildlife

Researcher

��-315July, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-26

Work Plan No.

16

Job Title

Piceance

Period Covered:
Personnel:

December

Deer-Elk
Job No.
Deer study - Population

Investigations
1

Distribution

1, 1971 through March 31, 1972

R. M. Bartmann,
F. C. Vanderau,

So F. Steinert, J. Jo Klein, Jr., D. C. Hoart,
D. E. Merriam and G. E. Lewis.

ABSTRACT

From January 3 to March 9, 1972, 378 deer were trapped and marked on the
Piceance winter range.
This included 99 deer in Area 1, 207 in Area 2,
4 in Area 3 and 78 in Area 4. The sex and age composition of the total was
70 male fawns, 58 female fawns, 67 mature males, 182 mature fema.les, and one
female of unknown age. An aerial deer distribution survey was made in December and again in January.
Little change in the upper winter range boundaries
was noted from December to January, presumably because of the relatively mild
and open winter.
No survey was made in February because of insufficient new
snow.

��-317-

PICEANCE

DEER STUDY - POPULATION

DISTRIBUTION

Richard M. Bartmann

P. S. OBJECTIVE
To develop methodology for the systematic collection of data relating
to mule deer population status on pinyon-juniper rangeland for use in
making annual harvest recommendations.

SEGMENT OBJECTIVES
1.

To delineate deer sub-population
on the Piceance winter range.

boundaries

and concentration

areas

(a) Define the winter range boundaries of the Piceance deer population under mile, moderate, and severe winter conditions.
(b) Define deer sub-population
winter range.

boundaries

(c) Delineate major deer concentration
winter range.

METHODS

within

the Piceance

areas within

the Piceance

AND MATERIALS

Deer Trapping
The deer winter range in Game Management Unit 22 was sub-divided into seven
areas for trapping purposes (Fig. 1). Tentative winter range boundaries
were those described by Baker (1970:35-39) based on browse form and age
class and relative pellet group density data gathered from 1965 through
1967.
Descriptions of trapping area boundaries are given below.
Descriptions
start at the northwest corner of each area and proceed in a clockwise
direction.

Area 1
Boise-Barcus--That
area bounded by the Game Management Unit 22 boundary
along Colorado Highway 64, Yellow Creek, Main Barcus Creek, the upper deer
winter range limit, and the Game Management Unit 22 boundary along Boise
Creek.

Area 2
Barcus-Ryan--That

area bounded by the Game Management

Unit 22 boundary

�-318-

along Colorado Highway 64, Piceance Creek, Ryan Gulch, the upper deer
winter range limit, Main Barcus Creek, and Yellow Creek.

Area 3
Ryan-Stuart--That
area bounded
range limit, and Ryan Gulch.

by Piceance

Creek,

the upper deer winter

Area 4
White River-Dry Fork--That area bounded by the Game Management Unit 22
boundary along Colorado Highways 64 and 13, the upper deer winter range
limit, Timber Gulch, Dry Fork, and Piceance Creek.

Area 5
Dry Fork-Greasewood--That
area bounded by Dry Fork, Timber Gulch, the
upper deer winter range limit, the divide between Dry Fork-Timber Gulch
and upper Piceance Creek, Greasewood Gulch, and Piceance Creek.

Area 6
Greasewood-Dudley
Bluffs--That area bounded by Greasewood Gulch, the divide
between Hatch Gulch and upper Piceance Creek, the divide formed by the
Dudley Bluffs, and Piceance Creek.

Area 7
Dudley Bluffs-Fourteenmile--That
area bounded by the divide formed by the
Dudley Bluffs, the divide between Hatch Gulch and upper Piceance Creek,
the divide between Dry Fork-Timber Gulch and upper Piceance Creek, the upper
deer winter range limit, and Piceance Creek.

Neckband

colors and the number of deer to be marked

in each area are:
No. of Deer

Color

Area

Neckband

1

Blue

140

2

White

220

3

Pink

100

4

Green with white

stripe

250

5

Yellow with red stripe

110

6

Blue and white

(~ &amp; ~)

100

7

Red with white

stripe

130

�N

t
Scale

o I 2 3 4 5 miles
E _- ~ r--l

(-

I

I
W

•.....

1

Upper winter
range limit
(Baker. 1970)

G'I \

\

'"
I

~ /.:£,-;

\

\

\\~,

,,
~

'\

')

,

L~__
,
Fig. 1. Location
trapping areas on of seven deer
the Piceance
winter range.

"'-""'\,
Unit

"'22

~

I

I \
(A"" _"
••••••
'\,

~_
bo •
un OIj

r-"'

\

--~,)

�-320-

A quota of 1,000 deer was originally set for the entire Unit 22 with
area quotas based on relative pellet group densities given by Baker
(1970:56) and the size of each area. No area was to have less than
100 marked deer. Adjustments were later made in some areas based on
observations during the winter of 1971-72 raising the unit total to
1,050 deer.
Deer were trapped with Clover traps
used only in Area 4. Each deer was
in both ears. A number was painted
identification of animals.
The red
served only as an "attention-getter"
easily seen in poor light.

and box traps, the latter being
neckbanded, and tagged and streamered
on each neckband for individual
streamer attached with each eartag
as some neckband colors were not

Sightings and recoveries of marked deer will be recorded from all available sources.
In addition, fixed-wing airplane flights will be made
during the spring migration to observe marked deer as they disperse from
the winter range.

Deer Distribution
Deer winter range boundaries and concentration areas were plotted at
monthly intervals from December through February.
Two methods were used
to gather this information.
The first was snowmobile transects located
throughout the winter range. Mechanical problems with machines, rapidly
changing snow conditions due to the highly variable topography, and the
relatively long interval required to adequately cover the entire winter
range forced discontinuance of this approach.
However, snowmobiles were
still used to check snow conditions on the ground.
Subsequently, fixed-wing airplane flights were made immediately after a
snowfall of at least four inches.
This amount was usually sufficient to
cover old deer tracks and, if cold weather followed, allow completion of
the survey before the south slopes became bare.
Each aerial survey took two days. Flight patterns followed natural
terrain features, usually ridges.
Two observers made separate notes of
relative deer track abundance (little or none, moderate, or heavy) on
field maps. Notes were later compared and a composite deer distribution
map overlay drawn.

RESULTS AND DISCUSSION

Deer Trapping
Deer trapping was done only in Areas 1, 2, 3, and 4 during the 1971-72
winter.
Each area was further sub-divided into smaller trapping locales
with all of the traps being set in several of these. As the desired number

�-321-

of deer were caught, or trapping success dropped, all traps in a particular locale were moved to a new one. It was thought that this procedure would improve trapping efficiency by reducing the number of recatches and the time necessary to check the traps.
Sixty Clover traps were used during most of the 66-day trapping period
from January 3 to March 9. In addition 20 box traps were used at odd
times and accounted for nearly all the deer marked in Area 4.
In all, 378 deer were marked: 99 in Area 1, 207 in Area 2, 4 in Area 3,
and 78 in Area 4 (Table 1). The total sex and age composition was 70
male fawns, 58 female fawns, 67 mature males, 182 mature females, and
1 female of unknown age. No records were kept of recatches because of
the trapping procedure used, but these records will be kept next year.
Quotas of marked deer were not reached in any area because of the
unusually mild and open winter.
During the March 23-24, 1972, meeting of the Colorado Game, Fish and
Parks Commission, a portion of the west boundary of Game Management Unit
22 was moved eastward about 2 miles from Hammond Draw to Boise Creek.
This places about 11 square miles of Unit 22 into Unit 21. About 35
deer were marked on these 11 square miles.
All figures in this report
show the old boundary as the change came into effect after the winter
fieldwork.

Deer Distribution
Only two aerial surveys were made during the winter of 1971-72;
December and one in January.
Insufficient new snow in February
that month's flight.

one in
precluded

From 10 to 18 inches of powder snow were present during the December survey on the 16th and 17th. Deer were generally scattered over the entire
winter range with high concentrations occurring only along the extreme
east and north sides of the unit (Fig. 2).
Cattle were still present on some BLM grazing allotments on the west side
of Piceance Creek from Black Sulphur Creek to Greasewood Creek.
Cattle
and deer tracks intermixed in the deep snow created some difficulty in
assessing relative deer track densities from the air. The problem would
have been reduced somewhat had this area been flown first before an extra
day's accumulation of tracks occurred.
The entire area was classified as
moderate deer density to acknowledge that deer were present over most of
the area.
Wild horses, also present throughout this western portion of the unit,
were found in small localized groups and their tracks were usually easily
recognized.
Elk were found at the upper limits of the deer winter range
in the southeast part of the unit from Stuart Gulch to Cow Creek.
Some

�-322-

confusion of tracks occurred there but was considered minimal as the
area is open brushland and the animals could usually be seen.
Upper elevational limits of deer winter range during December varied
by area. In the Piceance triangle, upper limits averaged around 7,500
to 7,600 feet. However, the steep heavily timbered northerly exposures
below these levels were generally void of deer. In the south and western
portions of the unit, upper limits were more variable ranging from about
7,300 to 7,800 feet.
By the last week in December, nearly all the snow had melted at the
lower elevations and on south slopes at higher elevations.
Little new
snow fell until the four to six inches received immediately prior to the
second aerial survey on January 14 and 15. Accumulated snow depths at
that time varied from four inches at the lowest elevations to 12 to 16
inches at the higher elevations.
Mean upper elevational limits of deer use decreased only slightly from
that observed in December (Fig. 3). Although snow depths were generally
less, varying degrees of crusting were found. More concentration areas
were located within the winter range as well as scattered areas of little
or no use. The problems with cattle experienced previously were almost
lacking.
Casual observations indicate that the spring migration to summer range
occurred earlier than normal.
The usual build-up of deer on open sagebrush parks and on meadowlands along Piceance Creek and other drainages
during late March and early April did not occur; and few deer were seen
on the meadows by the end of March.
Some deer were reported on summer
ranges along Cathedral Bluffs and the Roan Plateau shortly after April 1.

LITERATURE

CITED

Baker, B. D. 1970. Survey, inventory, and analysis of deer and elk
winter ranges.
p. 15-58.
In Game Research Report.
Colorado Div.
Game, Fish and Parks, Denver.
3(Part 1):1-126.

/"'/

/~.

Prepared

by

I

.

.: /).:-,
-.:.,-&amp;'., .•... .r
.
-c ..

-+:~7~...
;.~:
...
"",~_.~
__
,._

--

Richard M. Bartmann
Asst. Wildlife Researcher

�Table 1.

Date
(1972)

Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72.

Sex

Age

Eartag Numbers

Neckband
No.

Tnp.

Location
Rg.

Sec.

2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N

100W
100W
100W
100W
100W
100W
100W
100W
100W
100W
100W
100W
lOOW
100W
100W
100W
100W
100W
100W
lOOW
100W
100W
100W
100W
100W
100W
100W
100W
100W
100W

11
11
10
10
11
11
2
11
10
10
10
10
11
23
11
2
24
24
10
14
14
11
11
2
12
24
1
36
14
11

Remarks

Area 1 -- Blue Neckbands
1-4
1-4
1-5
1-5
1-5
1-5
1-5
1-5
1-6
1-6
1-6
1-6
1-6
1-6
1-6
1-6
1-6
1-6
1-7
1-7
1-7
1-7
1-7
1-7
1-7
1-7
1-7
1-7
1-8
1-8

Female
Male
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Male
Female
Female
Female
Male
Female
Female
Male
Female
Female
Male

Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn

L-1084
L-1086
L-1088
L-1090
L-1101
L-1103
L-1105
L-1107
L-1109
L-1116
L-1111
L-1113
L-1115
L-1119
L-1121
L-1123
L-1151
L-1153
L-1125
L-1127
L-1129
L-1131
L-1133
L-1135
L-1154
L-1158
L-1160
L-1162
L-1137
L-1139

L-1085
L-1087
L-1089
L-1091
L-1102
L-1104
L-1106
L-1108
L-1110
L-1117
L-1112
L-1114
L-1118
L-1120
L-1122
L-1124
L-1152
L-1155
L-1126
L-1128
L-1130
L-1132
L-1134
L-1136
L-1157
L-1159
L-1161
L-1163
L-1138
L-1140

1
5
8
3
9
2
11
4
6
26
28
27
23
24
22
25
16
17
48
15
49
45
19
14
31
30
18
36
44
61

-------------------------------------------------------------------------------------------------------------

I

w
N
w
I

�Table l. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-9

1-9
1-9
1-10
1-11
1-11
1-11
1-11
1-11
1-12
1-12
1-12
1-12
1-12
1-12
1-12
1-12
1-12
1-12
1-12
1-13

Sex

Age

Male
Male
Male
Male
Female
Male
Female
Male
Female
Male
Male
Male
Female
Female
Female
Male
Male
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Male
Female
Male
Male

Fawn
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn

Eartag Numbers
L-1141
L-1143
L-1149
L-1145
L-1164
L-1147
L-1156
L-1168
L-1170
L-1174
L-1172
L-1167
L-l177
L-1179
L-1181
L-1183
L-1185
L-1137
L-1189
L-1191
L-1193
L-1195
L-1197
L-1199
L-1201
L-1228
L-1230
L-1232
L-1234
L-1236
L-1238
L-1205

L-1142
L-1144
L-1150
L-1146
L-1165
L-1148
L-1166
L-1169
L-l171
L-1175
L-1173
L-1176
L-1178
L-1180
L-1182
L-1184
L-1186
L-1138
L-1190
L-1192
L-1194
L-1196
L-1198
L-1200
L-1202
L-1229
L-1231
L-1233
L-1235
L-1237
L-1239
L-1206

Neckband
No.
67
68
65
62
20
66
21
60
29
63
64
12
13
32
33
82
7
34
47
46
35
38
71
39
77
40
41
42
10
50
74
79

Tnp.
2N
2N
2N
2N
2N
2N
2N
2N
2N
3N
3N
3N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
3N
2N
2N
2N
2N
2N
?

Location
Rg.
100W
100W
100W
100W
100W
100W
100W
100W
100W
99W
99W
991,;1

100W
100W
100W
99W
99W
99W
98W
98W
99W
99W
99W
99W
99W
99W
98W
98W
98W
98W
98W
99W

Sec.

Remarks

11
11
12
11
13
24
24
24
13
36
34
36
13
14
24
1
11
12
8
9
1
12
12
12
12
3
8
8
8
8
8
?

I

w
N

.pI

Steadman Mesa

-----------------------------------------------------------~------------------------------------------------

�Table l. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

Sex

Age

Eartag Numbers

1-l3
1-l3
1-13
1-13
1-l3
1-13
1-14
1-14
1-14
1-14
1-15
1-15
1-15
1-15
1-15
1-15
1-15
1-16
1-16
1-16
1-16
1-16
1-16
1-17
1-17
1-18
1-19
1-19
1-19
1-22
1-23
1-24

Male
Female
Female
Female
Female
Male
Female
Female
Male
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Male
Female
Male
Male
Male
Male
Female
Female
Male
Female
Female
Female

Fawn
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Mature
?
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Fawn

1-1215
1-1203
1-1207
1-1209
1-1211
1-1213
1-1217
1-1219
L-1221
1-1223
1-1277
1-1279
1-1281
1-1283
1-1285
1-1287
1-1289
1-1295
1-1297
1-1299
1-1301
1-1303
1-l305
1-1240
1-1242
1-1323
1-1244
1-1246
1-1248
1-1262
1-l361
1-1367

1-1216
1-1204
1-1208
1-1210
1-1212
1-1214
1-1218
1-1220
1-1222
1-1224
1-1278
1-1280
1-1282
1-1284
1-1286
1-1288
1-1290
1-1296
1-1298
1-l300
1,-1302
1-1304
1-1306
1-1241
1-1243
1-1324
1-1245
1-1247
1-1249
1-1263
1-1362
1-1368

Neckband
No.

Tnp.

94
69
70
72
53
80
43
51
91
54
83
87
84
86
52
55
104
90
88
56
106
89
108
111
110
81
99
37
116
97
32
98

?
3N
2N
?
3N?
?
3N
?
2N
2N
3N
2N
3N
2N
2N
2N
2N
2N
2N
2N
3N
3N
3N
2N
2N
2N
2N
2N
2N
2N
2N
2N

1ocation
Rg.

Sec.

Remarks

?
99W
36?
99W
12
99W
?
99W
36?
99W
?
99W
99',;r 36?
?
99W
4
99W
4
99W
34
99W
1
99W
34
99W
4
99W
4
99W
4
99W
12
99W
99W
9
12
99W
12
99W
34
99W
36
99W
31
99W
12
99W
12
99W
11
99W
99W
9
99W
9
9
99W
33
99W
34
99W
34
99W

Steadman Mesa
Steadman Mesa
Steadman Mesa
Steadman Mesa
Steadman Mesa
Steadman Mesa
Steadman Mesa

I

w

N
V1
I

Has a white neckband

------------------------------------------------------------------------------------------------------------

�Table 1. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

Sex

Age

Eartag Numbers

1-26
1-27
1-28
1-28
1-28

Female
Male
Female
Female
Hale

Fawn
Fawn
Fawn
Mature
Fawn

L-1375
L-1417
L-1423
L-1425
L-1427

Summary:

19 male fawns;

19 female fawns;

L-1376
L-1418
L-1424
L-1426
L-1428

Neckband
No.

Tnp.

Location
Rg.

Sec.

100
75
102
57
117

2N
2N
2N
2N
2N

99W
99W
99W
99W
99W

34
34
34
34
34

15 mature males;

45 mature females;

Rema rks

1 female of unknown age.

Area 2 -- White Neckbands
1-12
1-15
1-16
1-16
1-17
1-17
1-17
1-17
1-17
1-18
1-19
1-19
1-19
1-19
1-20
1-20
1-20
1-20
1-20
1-20
1-21
1-21
1-21
1-21

Female
Female
Female
Female
Male
Male
Female
Female
Female
Female
Female
Female
Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
Male
Female

Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fmvn
Mature
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature

L-1226 L-1227
L-1225 L-1276
L-1291 L-1292
L-1293 L-1294
L-1307 L-1308
L-1309 L-1310
L-1311 L-1312
L-1313 L-1314
L-1319 L-1320
L-1321 L-1322
L-1325 L-1326
L-1327 L-1328
L-1329 L-1330
L-1331 L-1332
L-1250 L-1251
L-1252 L-1253
L-1254 L-1255
L-1256 L-1257
L-1258 L-1259
L-1260 L-1261
L-1333 L-1334
L-1335 L-1336
L-1337 L-1338
L-1341 L-1342

1
15
2
3
22
21
17
18
6
25
13
12
9
14
8
16
19
29
30
33
35
36
7
50

2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
IN
2N
2N
2N
IN
2N
2N
2N
2N
2N
2N
2N
2N
2N

98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
97W
98W
98W
98W
97W
98W
98W
98W
98W
98W
97W
97W
97W
98W

10
14
14
14
14
14
11
11
10
14
15
14
14
10
15
14
11
11
10
10
18
32
32
10

-----------------------------------------------------------------------------------------------------------

I

w
N

0"\
I

�Table l. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

Neckband
No.

Tnp.

Location
Rg.

L-1340
L-1344
L-1346
L-1348
L-1350
L-1352
L-1354
L-1356
L-1358
L-1360
L-1265
L-1267
L-1269
L-1271
L-1273
L-1275
L-1364
L-1366
L-1370
L-1372
L-1374

40
37
38
5
11
10
31
20
23
24
49
51
52
75
53
76
28
65
4
44
73

2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N

98W
98W
98W
97W
97W
97W
97W
98W
98W
98W
98W
98W
98W
98W
97W
97W
97W
97W
97W
97W
97W

L-1402
L-1404
L-1406
L-1408
L-1410
L-1414
L-1416
L-1422
L-1420
L-1430
L-1432

·77
60
42
39
74
57
26
58
124
86
61

IN
2N
IS
2N
IN
IS
2N
IN
IS
IS
IN

97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W

Sex

Age

Eartag Numbers

1-21
1-21
1-21
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-23
1-23
1-23
1-23
1-23
1-23
1-24
1-24
1-24

Female
Female
Female
Female
Male
Male
Female
Male
Male
Male
Female
Female
Female
Male
Female
Male
Male
Male
Female
Female
Male

Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn

L-1339
L-1343
L-1345
L-1347
L-1349
L-1351
L-1353
L-1355
L-1357
L-1359
L-1264
L-1266
L-1268
L-1270
L-1272
L-1274
L-1363
L-1365
L-1369
L-1371
L-1373

1-24
1-25
1-26
1-26
1-26
1-27
1-27
1-28
1-28
1-29
1-29

Male
Female
Female
Female
Male
Female
Male
Female
Male
Female
Female

Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature

L-1401
L-1403
L-1405
L-1407
L-1409
L-1413
L-1415
L-1421
L-1419
L-1429
L-1431

Sec.

Remarks

10
14
14
18
29
29
32
14
23
14
10
10
I
w
14
N
--.J
11
I
32
33
18
29
18
19&gt;
Caught in same trap and
19.both
chewed on slightly
by dog
10
18
5
31
8
5
31
28
17
5
15

------------------------------------------------------------------------------------------------------------

�Table l. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued).
Date
(1972)

1-30
1-30
1-30
1-30
1-30
1-31
1-31
1-31
1-31
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-2
2-3
2-3
2-3

Sex

Age

Female
Female
Female
Female
Male
Female
Female
Female
Female
Male
Female
Male
Male
Male
Female
Male
Male
Female
Male
Male
Male
Female
Female
Female
Female
Male
Female
Female
Female
Male
Male
Male

Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Mature

Eartag Numbers

Neckband
No.

Tnp.

Location
Rg.

Sec.

L-1433
L-1435
L-1437
L-1439
L-1441
L-1443
L-1445
L-1447
L-1449
L-1504
L-1506
L-1508
L-1510
L-1512
L-1514
L-1516
L-1518
L-1520
L-1522
L-1524
L-1526
L-1528
L-1530
L-1532
L-1534
L-1536
L-1538
L-1540
L-1542
L-1450
L-1544
L-1546

45
43
82
89
80
55
59
54
56
71
41
122
123
79
47
67
121
48
81
78
98
96
85
88
87
103
90
92
91
97
99
100

IS
IS
IS
IN
IN
IS
IS
IS
IN
2S
2S
2S
2S
2S
2S
2S
IN
2N
2N
2N
IS
IS
IS
IS
IS
IS
IS
IN
IN
IS
IS
IS

98W
98W
97W
97W
97W
98W
97W
97W
97W
98W
98W
98W
97W
97W
97W
97W
97W
97W
98W
98W
98W
98W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W

35
35
19
28
15
25
8
5
10
12
12
19
17
8
8
8
15
8
1
1
35
35
19
8
5
5
5
28
28
5
5
8

L-1434
L-1436
L-1438
L-1440
L-1442
L-1444
L-1446
L-1448
L-1501
L-1505
L-1507
L-1509
L-1511
L-1513
L-1515
L-1517
L-1519
L-1521
L-1523
L-1525
L-1527
L-1529
L-1531
L-1533
L-1535
L-1537
L-1539
L-1541
L-1543
L-1451
L-1545
L-1547

Remarks

------------------------------------------------------------------------------------------------------------

I

w
N

00
I

�Table 1- Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

Sex

Age

Eartag Numbers

2-3
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-7

Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Male
Female
Male
Female
Female
Female

Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Fawn
Mature
Mature
Mature
Fawn
Mature
Fawn
Fawn
Mature
Mature

L-1548
L-1550
L-149l
L-1493
L-1495
L-1497
L-1499
L-1551
L-1553
L-1555
L-1557
L-1558
L-1569
L-1570
L-1571
L-1572
L-1573
L-1574
L-1559
L-1560
L-1561
L-1562
L-1563
L-1564
L-1565
L-1575
L-1566
L-1567
L-1568
L-1576
L-1577
L-1607

L-1549
L-1490
L-1492
L-1494
L-1496
L-1498
L-1500
L-1552
L-1554
L-1556
L-1557
L-1558
L-1569
L-1570
L-1571
L-1572
L-1573
L-1574
L-1559
L-1560
L-1561
L-1562
L-1563
L-1564
L-1565
L-1575
L-1566
L-1567
L-1568
L-1576
L-1577
L-1607

Neckband
No.

Tnp.

95
68
84
109
125
46
34
112
111
62
105
104
118
129
126
128
113
114
63
64
66
101
69
70
71
72
102
93
119
83
94
146

IS
IS
IS
IS
IS
1S
1S
2S
2S
1S
1S
2S
1S
1S
1S
IS
1S
IS
IN
IS
IS
1S
IS
IS
IS
2S
IS
IS
IS
IS
IS
IS

Location
Rg.
97W
97W
97W
97W
97W
97W
98W
98W
98W
97W
97W
98W
97W
97W
97W
97W
97W
98W
97W
97W
97W
97W
97W
97W
98W
98W
98W
98\-J

98W
97W
97~-J
97~-J

Sec.
8
5
5
8
19
19
25
3
3
32
8
12
29
29
32
32
31
18
22
5
5
5
8
19
25
4
18
18
35
32
32
31

Remarks

Possible neck injury

------------------------------------------------------------~----------------------------------------------

I

LV
N

-o
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

2-7
2-7
2-7
2-7
2-7
2-7
2-7
2-8
2-8
2-8
2-8
2-8
2-8
2-8
2-9
2-9
2-9
2-9
2-9
2-9
2-9
2-9
2-9
2-9
2-9
2-10
2-11
2-11
2-11
2-11
2-11
2-11

Sex

Age

Male
Female
Female
Female
Male
Female
Female
Female
Female
Male
Male
Female
Female
Female
Female
Male
Female
Male
Female
Female
Female
Male
Female
Male
Male
Female
Female
Male
Male
Female
Male
Female

Fawn
Fawn
Mature
Fawn
Fawn
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature

Eartag Numbers

Neckband
No.

Tnp.

L-1579
L-1601
L-1602
L-1603
L-1604
L-1605
L-1606
L-1608
L-1609
L-1610
L-1611
L-1612
L-1580
L-1581
L-1613
L-1614
L-1615
L-1616
L-1582
L-1583
L-1584
L-1585
L-1586
L-1587
L-1588
L-1589
L-1590
L-1591
L-1592
L-1593
L-1594
L-1595

120
116
117
127
134
130
145
107
110
140
139
115
108
155
147
135
148
138
156
159
157
142
158
141
131
149
106
137
144
151
143
150

2S
IN
IN
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IN
IS
IS
IS
IS
2S
2S
2S
IS
IS
IS
IS
IS
IS

L-1579
L-1601
L-1602
L-1603
L-1604
L-1605
L-1606
L-1608
L-1609
L-1610
L-1611
L-1612
L-1580
L-1581
L-1613
L-1614
L-1615
L-1616
L-1582
L-1583
L-1584
L-1585
L-1586
L-1587
L-1588
L-1589
L-1590
L-1591
L-1592
L-1593
L-1594
L-1595

Location
Rg.

98W
97W
97~J

97W
97W
98W
97W
97W
97W
97W
97W
98W
97W
97W
97W
99W
99W
98W
97W
97W
97W
97W
97W
98W
98W
97W
97W
97W
98W
98W
98W
98W

Sec.

Remarks

4
28
28
5
8
35
31
29
32
31
31
18
5
5
32
10
10
18
28
5
8
8
8
4
4
25
5
8
13
13
25
18

----------~-----------------------------------------------------~------------------------------------------

I

w
w
0
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

Sex

Age

Eartag Numbers

Neckband
No.

Tnp.

Location
Rg.

2-11
2-11
2-11
2-12
2-12
2-12
2-12
2-l3
2-13
2-l3
2-14
2-14
2-15
2-15
2-15
2-16
2-16
2-17
2-17
2-17
2-17
2-17
2-17
2-17
2-17
2-17
2-18
2-18
2-18
2-18
2-18
2-18

Male
Male
Female
Male
Male
Male
Female
Female
Male
Female
Female
Male
Female
Female
Female
Female
Male
Female
Female
Female
Male
Male
Male
Female
Female
Female
Female
Male
Male
Hale
Female
Male

Mature
Fawn
Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Hature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Hature
Mature
Fawn

L-1596 L-1596
L-1597 L-1597
L-1598 L-1598
L-1599 L-1599
L-1600 L-1600
L-1617 L-1617
L-1618 L-1618
L-1619 L-1619
L-1620 L-1620
L-1621 L-1621
L-1622 L-1622
L-1623 L-1623
L-1651 L-1651
L-1652 L-1652
L-1624 L-1624
L-1625 L-1625
L-1626 L-1626
L-1672 L-1672
L-1673 L-1673
L-1627 L-1627
L-1628 L-1628
L-1629 L-1629
L-1630 L-1630
L-1631 L-1631
L-1632 L-1632
L-1674 L-1674
L-1633 L-1633
L-1634 L-1634
L-1635 L-1635
L-1636 L-1636
L-1637 L-1637
L-1638 L-1638

l36
l33
152
169
172
171
179
153
170
154
160
168
176
177
161
184
167
181
183
162
l32
164
165
163
178
185
182
193
166
174
180
173

IS
IS
IS
IS
IS
2S
IS
IS
IS
IS
IS
IS
2S
2S
IS
2S
2S
2S
2S
IN
IN
IN
IN
IS
IS
2S
2S
IS
IS
IS
IS
IN

97W
97W
97W
97W
98W
98W
97W
97W
98W
99W
97W
97W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
99W
98W
98W
98W
98W
98W
98W

Sec.

Remarks

31
32
32
19
25
3

32
32
19
11
29
32
4
3
19
18
18
4
8
32
20
33
33
4
18
l3
8
17
18
3
3
33

------------------------------------------------------------------------------------------------------------

I

w
w
t-'
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued).
Date
(1972)

Sex

Age

2-19
2-19
2-20
2-20
2-21
2-22
2-23
2-23
2-24
2-24
2-24
2-25
2-26
2-27
2-27
2-27
2-27
2-27
3-2
3-2
3-6
3-7
3-9

Female
Female
Male
Male
Male
Male
Female
Female
Female
Male
Male
Male
Male
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female

Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature

Summary:

35 male fawns ; 28 female fawns;

Eartag Numbers

L-1653
L-1654
L-1655
L-1656
L-1657
L-1658
L-1659
L-1713
L-1660
L-1661
L-1662
L-1663
L-1701
L-1707
L-1706
L-1705
L-1704
L-1703
L-1709
L-1708
L-1691
L-1694
L-1696

L-1653
L-1654
L-1655
L-1656
L-1657
L-1658
L-1659
L-1713
L-1660
L-1661
L-1662
L-1663
L-1702
L-1707
L-1706
L-1705
L-1704
L-1703
L-1709
L-1708
L-1691
L-1694
L-1696

Neckband
No.

Tnp.

186
187
175
191
195
192
188
200
199
198
120
197
196
201
204
202
203
205
189
206
190
194
208

IN
IS
IN
IN
2S
IN
IN
2S
IS
2S
2S
2S
IN
2S
2S
IS
IS
IN
2S
2S
IS
IS
IN

39 mature males;

Location
Rg.
Sec.

98W
98W
98W
98W
98W
98W
98W
99W
98W
99W
99W
98W
98W
99W
98W
98W
98W
98W
99W
99W
98W
98W
98W

Remarks

28
19
29
20
8
27
33
13
18
13
13
8
32
13
4
18
2
32
13
13
3
2
32

105 mature females.

Area 3 -- Pink Neckbartds
3-2
3-3
3-3
3-6
Summary:

Female
Mature
L-1710
Male
Fawn
L-1711
Female
Mature
L-1712
Female
Mature
L-1692
1 male fawn; 3 mature females.

L-1710
L-1711
L-1712
L-1692

1
12
2
5

2S
2S
2S
2S

97W
97W
97W
97W

32
29
32
29

------------------------------------------------------------------------------------------------------------

I

w
w
N
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

Sex

Eartag Numbers

Age

Neckband
No.

1ocation
Rg.
Tnp.

Sec.

Remarks

Area 4 -- Green with White Stripe'Neckban ds
1-23
1-23
1-23
1-23
1-25
1-25
1-25
1-25
1-26
1-26
1-26
1-26
1-26
1-27
1-27
1-28
1-28
1-29
1-29
1-29
1-29
1-30
1-30
1-31
1-31
1-31
1-31
1-31
2-1
2-2
2-2
2-2

Male
Male
Female
Male
Male
Female
Female
Female
Female
Female
Male
Female
Male
Female
Female
Male
Male
Male
Female
Female
Female
Male
Female
Female
Female
Female
Female
Male
Female
Male
Female
Female

Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature

1-1377
1-1379
1-1381
1-1383
1-1385
1-1387
1-1389
1-1391
1-1393
1-1395
1-1397
1-1399
1-1411
1-1452
1-1454
1-1456
1-1458
1-1466
1-1460
1-1462
1-1464
1-1468
1-1470
1-1472
1-1474
1-1476
1-1478
1-1480
1-1482
1-1484
1-1486
1-1488

1-1378
1-1380
1-1382
1-1384
1-1386
1-1388
1-1390
1-1392
1-1394
1-1396
1-1398
1-1400
1-1412
1-1453
1-1455
1-1457
1-1459
1-1467
1-1461
1-1463
1-1465
1-1469
1-1471
1-1473
1-1475
1-1477
1-1479
1-1481
1-1483
1-1485
1-1487
1-1489

11
12
1
13
15
2
3
4
17
5
14
6
16
7
8
18
20
22
28
26
27
None
10
9
19
21
23
32
24
30
25
29

IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN

96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

28
28
34
20
28
21
34
20
28
21
20
20
20
28
20
28
28
20
28
20
20
21
21
28
34
34
20
20
34
28
34

34

------------------------------------------------------------------------------------------------------------

I

w
w
w
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

Sex

Age

Eartag Numbers

2-15
2-15
2-15
2-15
2-15
2-16
2-16
2-16
2-16
2-16

Male
Female
Male
Female
Male
Female
Female
Female
Female
Female

Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature

L-1639
L-1640
L-1641
L-1642
L-1643
L-1644
L-1645
L-1646
L-1648
L-1649

L-1639
L-1640
L-1641
L-1642
L-1643
L-1644
L-1645
L-1646
L-1648
L-1649

2-16
2-16
2-16
2-17
2-17
2-17
2-17
2-17
2-17
2-18
2-18
2-19
2-21
2-21
2-21
2-23
2-24

Female
Female
Male
Female
Male
Female
Female
Female
Female
Male
Female
Male
Male
Male
Female
Male
Male

Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature

L-1664
L-1650
L-1665
L-1666
L-1667
L-1668
L-1669
L-1670
L-1671
L-1675
L-1676
L-1677
L-1678
L-1679
L-1680
L-1681
L-1682

L-1664
L-1650
L-1665
L-1666
L-1667
L-1668
L-1669
L-1670
L-1671
L-1675
L-1676
L-1677
L-1678
L-1679
L-1680
L-1681
L-1682

Neckband
No.

Location
Tnp.
Rg.

Sec.

31
37
38
36
39
35
34
33
42
43

IN
IN
IS
IS
IS
IN
IN
IS
IS
IS

96W
96W
96W
96W
95W
96W
96W
96W
96W
96W

34
34
3
11
18
34
34
3
12
12

45
44
40
41
49
47
48
50
53
46
54
51
52
73
55
75
76

IS
IS
IS
IN
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IN
IN
IN

96W
96W
95W
96W
96W
96W
96W
96W
95W
96W
96W
96W
95W
96W
96W
96W
96W

13
13
18
34
11
11
2
13
18
3
12
11
19
11
34
36
26

Remarks

Collar may be #34 also
Velvet covered antler
nubbin on right side
which was broken at the ~
w
skull
.p.

------------------------------------------------------------------------------------------------------------

I

�Table l. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1971-72
(continued) .
Date
(1972)

Sex

Age

Eartag Numbers

2-26
2-27
2-27
2-27
2-27
3-1
3-4
3-6
3-7

Female
Female
Male
Male
Male
Female
Male
Male
Female

Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature

L-1683
L-1684
L-1685
L-1686
L-1687
L-1688
L-1689
L-1690
L-1693

Sununary: 15 male fawns;

11 female fawns;

L-1683
L-1684
L-1685
L-1686
L-1687
L-1688
L-1689
L-1690
L-1693

Neckband
No.
56
64
77
78
57
58
83
85
59

13 mature males;

Location
Rg.
Tnp.

Sec.

lS
IN
lS
lS
IS
lN
lS
lN
lN

6
36
12
13
19
36
6
36
36

95W
96W
96W
96W
95W
96W
95W
96W
96W

Remarks

29 mature females

I
Vol
Vol

V1
I

Grand Summary:

70 male fawns;

58 female fawns; 67 mature males;

182 mature females;

1 female of unknown age

�N

t
\
II

Scale

o

-.--.--....
I

2

3

4

5 ml les

»J/ .

t"
I

-.

.

. ,.,.

I

W
W
0'\

1
\\

\

\

\\~,
~OI

IEGEND

~

W#iWi'JhUXI Heavy use

,

."

..

I -~

,

,

,.

, Moder
Little

or no use

r

,

""-,-,..,.,.,.,
Fig. 2. Deer distribution
on the
Piceance winter range. December 16
and 17. 1971.

"-~
Unit

"'-

cc'

(I" -,

"""""',
\

I
'\,1

)

.-I

' ;:darY

�o
i

I

Scale
2 3

;;;;;-,.

4

5 miles

?rw-'I

,/

J

(-

I

r&lt;')
I

1

Heavy use

I ... . ···. . . ·1.

Moderate use

I

'-J
I

1:1

~

1£

\
\
~\
~
~~ '\
%. ,

ffi'@mmml]

W

.C
01

&lt;B \
~

LEGEND

w.

&gt;.

J

"4..--,_,_~
)

I Little or no use~

Fig. 3. Deer distribution on the
Piceance winter range, Janua~ 14
and 15, 1972 •

Unif

~"'

c;&lt;

_
Ir-'~don'

••••••••
~

.'~\

..,~j
1

bell

\
{~

"'\'

...•...""".../

,\'111\.'1-

~,

��-339July, 1972

JOB PROGRESS REPORT

State of

~C~O~L~O~RA==D~O _
Deer-Elk

W-38-R-26

Project No.

Job No.

Work Plan No.

16

Job Title

Piceance

Period Covered:
Personnel:

November

Investigations

Deer Study - Population

2
Density

1, 1971 through March 31, 1972

R. M. Bartmann, R. B. Gill, J. J. Klein, Jr., S. F. Steinert
and F~ C. Vanderau.

ABSTRACT

Experimental helicopter counts of deer in three of the Little Hills pastures
and on 1/16-square mile and 1/4-square mile quadrats on open range were made
during February to assess the feasibility of such counts on pinyon-juniper
winter range. All three aerial counts in the pastures were in agreement with
ground counts.
Subjective evaluation of the two quadrat sizes favored the
1/4-square mile area as probably the most efficient of the two. Other conditions that were recognized include the need for adequate quadrat corner
marking, the use of two observers and the impracticality of classifying
animals during counts.

��-341-

PICEANCE

DEER STUDY - POPULATION

DENSITY

Richard M. Bartmann

P. S. OBJECTIVE
To develop methodology for the systematic collection of data relating to
mule deer population status on pinyon-juniper rangeland for use in making
annual harvest recommendations.

SEGMENT OBJECTIVE
1.

To establish a deer density estimate
yon-juniper winter range.

technique

(a)

for estimating

Evaluate the use of a helicopter
pinyon-juniper winter range.

METHODS

for the Piceance

pin-

deer density on

AND MATERIALS

The use of a helicopter for aerial counting of deer on the Piceance pinyonjuniper winter range was tested February 4, 1972. Mr. R. Bruce Gill, who
has had five years experience in deer quadrat census from a helicopter in
Middle Park, Colorado, did the counting.
Two aspects of helicopter counting on pinyon-juniper winter range were
checked.
One was a test of accuracy of counts.
Deer, already present in
three of the Little Hills pastures, were counted and results compared to
ground tallies.
A conscious effort was made to count each pasture as an
actual quadrat and not take advantage of the fenced boundaries.
The other aspect was to subjectively evaluate two sizes of quadrats for
counting; 1/16-square mile and 1/4-square mile.
Three quadrats of each
size were located in four different situations representative of various
terrain and overstory conditions present on the winter range.
These included (1) lightly broken terrain with scattered stands of pinyon-juniper,
(2) moderately steep terrain with moderate to heavy pinyon-juniper overstory, (3) extremely steep terrain with variable pinyon-juniper overstory,
and (4) extremely steep terrain with variable pinyon-juniper overs tory and
with Douglas fir stands on northerly exposures.
The fourth situation
comprises only a small proportion of the total winter range area.
Selection of quadrat sites was entirely subjective.
In addition to sampling
the above situations, they were located in areas of suspected high deer
density.
Quadrat corners were plotted on aerial photos and later marked
in the field with surveyors flagging.
The time to count each quadrat and
the number of deer seen were recorded.

�-342-

RESULTS AND DISCUSSION
Snow cover for the experimental helicopter counts was less than desired
for good counting conditions, but was probably typical of conditions that
would be encountered during some years.
Patchy snow cover occurred on
south slopes with mostly complete cover on north exposures.
Aerial and ground counts of deer in the pastures agreed in all three instances (Table 1). Grount counts in Pastures 4 and 7 were completed within 4 and 12 days, respectively, of the aerial counts. Thus the possibility
of ingress, egress, or mortality was small. Pasture 3 did not lend itself
to ground counting and the deer had to be trapped.
Gates were opened into
an adjacent group trap and the deer baited out with alfalfa hay. Fourteen
deer were removed in this manner from March 8 through March 17. A drive
through the pasture was made April 5 and one deer was seen for a total of
15 deer believed present.
The possibility of population change was greater
than in the other pastures because of the longer period between the aerial
count and final ground check. Despite only three pastures being counted,
the artificiality of the pasture situation, and the possibility for population changes between the times of the air and ground counts, it is
encouraging that the results were in agreement all three times.

Table 1. Results of experimental helicopter
Hills pastures, February 4, 1972.

counts of deer in the Little

Pasture

Acres

Number of Deer
Air
Ground

3

162

15

15

6:20

6:15

4

144

10

10

5:55

6:35

7

121 "l/

12

12

4:35

6:05

Includes

about 30 acres in an adjacent

Time to Count (minutes and seconds)
Actual
Adjusted to 160 Acres

"l/
gulch.

The results of deer counts on the two sizes of quadrats are shown in Table 2.
Because the quadrat locations were non-randomly selected, little can be
extrapolated from the results.
However, the data do suggest considerably
higher variability associated with the smaller quadrats.
The 1/4--square
mile quadrat was selected for continued evaluation because it presents a
larger area for counting which does not seem unwieldy, and it should be
less variable which would result in smaller sample sizes. The possibility
of further increase in quadrat size is not ruled out and will be considered
during the next segment.

�-343-

Table 2. Results of experimental helicopter counts of deer on two sizes
of quadrats on the Piceance pinyon-juniper winter range, February 4, 1972.
No. of
Deer

Quadrat

1/16-square

Time to Count
(min. &amp; sec. )

Quadrat

No. of
Deer
1/4-square

mile

Time to Count
(min. &amp; sec.)
mile

1

4

3:30

2

14

4:00

3

24

2:35

4

11

3:35

5

3

1:55

6

19

4:35

7

0

1:45

8

24

3:30

9

3

1:50

10

12

5:15

11

0

1:55

12

3

5:45

13

0

2:10

14

4

6:15

15

13

2./

16

18

5:15

17

0

2:00

18

0

6:15

19

0

1:45

20

4

5:55

21

0

2:50

22

0

3:45

23

0

1:55

24

3

6:15

3.9

2:10

9.3

5:00

x =

1/ Starting

x =

time not recorded.

Several problems were identified during the course of the preliminary
helicopter counts.
A major one was adequate marking of quadrat corners.
Much of the flagging on open ridgetops had been removed by the wind, and
flagging in other areas was difficult to see. It is estimated that only
about 10 percent of the marked corners were actually seen during the counts
even though aerial photos with quadrat boundaries marked on them were referred to during flight.
An associated problem is keeping oriented on quadrat boundaries.
Many
corners will not be seen until almost above them because of the rough
terrain and the overstory.
Placing of markings high in trees whenever
possible and reference to aerial photos during counts should help alleviate
much of the problem.

�-344-

Deer visibility, as long as the animals were moving, was good. Although,
extra care was required to keep track of animals as they darted under
trees and were momentarily out of sight. An extra observer, in addition
to keeping track of quadrat boundaries, can help locate and count deer.
If the animals are not moving, they could very easily be missed.
Since
nearly all deer seem to run at the approach of a helicopter, and with
two observers and the relatively small quadrat size, it is hoped this
error will be minimal.
A good job of sex and age classification does not seem possible during
the counts.
It is a full-time job for two people just to keep track of
deer on the quadrat.
Such classifications may have to be made separately
either from the air or on the ground.
Ground speed, altitude, and counting pattern will be dictated
terrain and cover conditions present on each quadrat.

Prepared

by the

�-345July, 1972

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~~--------

Project No.

W-38-R-26

Work Plan No.

16

Job Title

Job No.

Picenace Deer Study - Productivity

Period Covered:
Personnel:

Deer-Elk

Investigations
3

and Mortality

April l~ 1971 through March 31, 1972

R. M. Bar tmann , J. J. Klein, Jr., S. F. Steinert and F. C. Vanderau.

ABSTRACT

Seventeen does were collected from March through June, 1971 and 20 does from
January through April, 1972. Twenty-three were killed along highways,
11 were
trapping mortalities and three died from other causes. Prenatal productivity
data for these deer in 1971 show 1.65 fetuses:doe with a sex ratio of 87 males:
100 females.
In 1972, they averaged 1.45 fetuses:doe with a sex ratio of 123
males:lOO females.

��-347-

PICEANCE

DEER STUDY - PRODUCTIVITY

AND MORTALITY

Richard M. Bartmann

P. S. OBJECTIVE
To develop methodology for the systematic collection of data relating
to mule deer population status on pinyon-juniper rangeland for use in
making annual harvest recommendations.

SEGMENT OBJECTIVE
1.

To estimate

increments

and losses to the Piceance

(a) Estimate productivity

rates for the Piceance

deer population.
deer population.

METHODS AND YillTERIALS
Prenatal productivity data were recorded for female mule deer of reproductive age during the 1971 and 1972 gestation periods.
Major sources
of animals were road-kills along Colorado Highways 13 and 64 and the
Piceance road, and trapping mortalities.
Ovaries were preserved in 10 percent formalin.
They were later sectioned
and examined macroscopically
for corpora lutea of pregnancy.
The number
and sex of fetuses was recorded.
Forehead-rump measurements were taken in
millimeters and converted to age in days according to the fetal growth
curve of Hudson and Browman (1959). Conception dates were estimated by
back-dating from date of death.
The first incisor on the left mandible was preserved in formalin for later
age estimation by counts of dental cementum annuli (Erickson and Seliger,
1969).

RESULTS AND DISCUSSION
Seventeen does were examined from March through June, 1971; 14 died along
highways, 2 died while being removed from one of the Little Hills pastures,
and 1 died from predation.
From January through April, 1972, 20 does were
examined; 11 died from trapping injuries and 9 died along highways.
How representative the above animals are of the entire population with
respect to productivity parameters is not known.
Yearly sample sizes are
admittedly small, but these are the first prenatal productivity data
collected for the Piceance herd.

�-348-

All does in 1971 and all but one in 1972 were pregnant
sex ratios favored females in 1971 and males in 1972.

(Table 1).

Fetal

Estimated conception dates based on forehead-rump measurements of 21
fetuses in 1971 and 27 fetuses in 1972 range from November 24 to December
29 the first year, and from November 14 to December 3 the second year
(Fig. 1). The majority of breeding activity, based on both years data,
occurred between November 16 and December 5.

LITERATURE

CITED

Erickson, J. A., and W. C. Seliger.
1969. Efficient sectioning of
incisors for estimating ages of mule deer. J. Wildl. Mgmt.

33(2):384-388.
Hudson, P., and L. G. Browman.
1959. Embryonic and fetal development
of the mule deer. J. Wildl. Mgmt.
23(3): 295-304.

/i

'

:/./ ?/

Prepared

~/

hi

/

.r=::»:--------

by r/c f-,,?,: __ ,._'0
/'
/ -'-::::::::;;t:-o
1'/.•.,.;_ ._,.
.••
..~
Richard M. Bar~n
Asst. Wildlife Researcher

�-349-

Table 1. Prenatal productivity data for mule deer does dying from various
causes in and around Game Management Unit 22, 1971 and 1972.
Collection
Number

Estimated
Age 1/

Date
Killed

How
Killed

No. of Fetuses
Female
Male

Corpora Lutea
of Pregnancy

Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Predator
Vehicle
Hit fence
Hit fence
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle

0
0
1
1
2
0
1
1
1
1
0
1
1
1
0
1
1

2
2
3
2
2
2
2
1
1
2
2
2
1
1
2
2
One ovary missing

1971
P-l
P-3
P-4
P-5
P-7
P-I0
P-ll
P-12
P-15
P-16
P-17
P-22
P-23
P-26
P-27
P-28
P-29

Mature
Mature
Mature
Yrlg.
Mature
Mature
Mature
Yrlg.
Yrlg.
Mature
Mature
Mature
Yrlg.
Yrlg.
Mature
Mature
Mature
165 fetuses:l00

3-12
3-24
3-30
4-1
4-1
4-5
4-5
4-6
4-9
4-9
4-8
4-15
4-18
5-2
5-12
5-12
6-14

P-50
P-51
P-52
P-53
P-54
P-55
P-56
P-57
P-58
P-59
P-60
P-61
P-62
P-63
P-64
P-66
P-67
P-68
P-69
P-70

Mature
Mature
Mature
Mature
Yrlg.
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Yrlg.
Mature
Mature
Yrlg.
Mature
Mature
Yrlg.
Yrlg.

1.20 CLP:yearling

does

87 male fetuses:l00

female fetuses
1972
1-21
1-21
1-31
2-3
2-6
2-8
2-9
2-10
2-15
2-16
2-20
2-27
3-7
3-8
3-8
4-2
4-2
4-5
4-6
4-14

Trap
Vehicle
Trap
Trap
Trap
Trap
Trap
Trap
Trap
Trap
Trap
Vehicle
Vehicle
Trap
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle

145 fetuses:l00 does
123 male fetuses:l00 female fetuses

1/ Age estimated

2
2
1
0
0
2
1
0
0
1
2
1
0
0
2
0
1

by tooth replacement.

2.09 CLP:mature

2
0
1
1
0
1
1
1
0
1
1
1
1
1
1
0
1
1
1
0

doe
doe

0
2
0
1
1
1
1
1
0
0
0
1
0
1
1
1
0
1
0
1

1.00 CLP:yearling doe
1.87 CLP:mature doe

2
2
1
2
1
2
2
2
1
2
1
2
1
2
2
1
3
2

1
1

�-350-

_1971

t:::J 1972
III

s::
0

.r-!

+'
o,
&lt;D

o

s::

7
6

5

0

4

4-l

3

(.)

0

,..
&lt;D

~
~

2
1
\.f\
0..-1

&gt;0..-1I
Z..-I

0
oN
I
O~
Z..-i

&gt;

\.f\

oN

&gt;0..-1I
ZN

0
o C'\
I
O~
ZN

&gt;

\.f\
0

()\.f\

&lt;D

I

~..-i

00
()..-I
Q) I

~~

o ..-i

o
I
Q)..-I
~..-I

0
oN

o

I

Q)~
~..-I

\.f\

oN

o

I

Q)..-I
~N

0
o C'\
I
Q)~
~N

o

Date
Fig. 1..Estimated

conception dates of Piceance deer fetuses,

1971 and 1972.

�-351July, 1972

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~----------

Project No.

W-38-R-26

Work Plan No.

16

Job Title

Investigations

Job No.

4

Piceance Deer Study - Food Habits

Period Covered:
Personnel:

Deer-Elk

April 1, 1971 through March 31, 1972

R. M. Bar tmann , J. J. Klein, Jr., S. F. Steinert and F. C. Vanderau.

ABSTRACT
Twenty-three deer rumen content samples were collected from March through May,
1971. Grass and big sagebrush (Artemisia tridentata) were the most important
food items comprising 41 and 33 percent, respectively, of the total volume.

��-353-

PICEANCE

DEER STUDY - FOOD HABITS

Richard M. Bartmann

P. S. OBJECTIVE
To develop methodology for the systematic collection of data relating to mule deer population status on pinyon-juniper rangeland
for use in making annual harvest recommendations.

SEGMENT OBJECTIVE
1.

To evaluate

deer forage use on pinyon-juniper

winter range.

(a) Estimate forage use by deer from stomach content analyses.

METHODS AND MATERIALS
Rumen content samples were collected from deer found freshly killed
along Colorado Highways 13 and 64, and the Piceance road. After
thoroughly mixing contents of the rumen and reticulum, about one
quart of food material was squeezed to remove excess rumen fluid
and then frozen for later analysis.
Another 103 rumen content samples were made available from Work
Plan 15, Deer-Auto Accident Investigations.
These samples were
collected yearlong from deer killed along Colorado Highway 13 during
1969 and 1970. Samples were preserved in 10 percent formalin.
In preparation for analysis, samples were washed over an 8-millimeter
mesh screen. The material was air-dried and thoroughly mixed.
Analysis was by the point method described by Chamrad and Box (1964).
Shrubs and forbs were identified to species when possible, but no
attempt was made to do the same for grass and grass-like plants.

RESULTS AND DISCUSSION
Twenty-three rumen content samples were collected from March through
May, 1971 (Table 1). The two most important food items were grass
and big sagebrush.
Together they comprised 74 percent of the total
food volume.
Pinyon pine (Pinus edulis) and Utah juniper (Juniperus
osteosperma) each contributed 4 percent.
Total forb volume was
small, comprising only 7 percent.

�-354-

Sixteen rumen content samples were collected during November, 1971,
and one each December, 1971, and January, 1972. These samples have
not been analyzed nor have most of those collected under the deerauto accident study.

LITERATURE

CITED
A point frame for sampling

Chamrad, A. D., and T. W. Box. 1964.
rumen contents.
J. Wildl. Mgmt.

Prepared

~

//

/. a.- ~'/

28(3) :473-477.

.

by I~-/.:/~~'_',f//!~c
j;.. £••
,~~-.-,.;..-,-Richard M. Bartiitinn
Asst. Wildlife Researcher

�-355-

Table 1. Food items found in 23 rumen content samples collected from mule
deer killed along highways within and around Game Management Unit 22 during
March, April, and May, 1971.
Food Item

Mean Volume

(Percent)

Frequency

Browse
Artemisia tridentata
Pinus edulis
Juniperus osteosperma
Symphoricarpos tetonensis
Cercocarpus montanus
Berberis repens
Amelanchier utahensis
Populus angustifolia
Atriplex confertifolia
Prunus virginianus
Sarcobatus vermiculatus
Quercus gambellii
Pachystima myrsinites
Unidentified browse

33
4
4
3
2
2
1
1
1
tr.
tr.
tr.
tr.
1

Total Browse

52

Eriogonum umbellatum
Phlox caespitosa
Gutierrezia sarothrae
Eriogonum tristichum
Cryptantha sp.
Medicago sativa
Penstemon caespitosus
Unidentified forbs

3
2
1
tr.
tr.
tr.
tr.
1

Total Forbs

7

Grass and Grass-like

41

0.96
0.39
0.43
0.17
0.22
0.26
0.17
0.09
0.09
0.04
0.04
0.04
0.04
0.43

Forbs
0.35
0.17
0.04
0.09
0.09
0.04
0.04
0.43

1.00

Index

��-357July, 1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-26

Work Plan No.

14
Middle
Physiology

Job Title

Period Covered:
Personnel:

Deer-Elk

Investigations

Job No.
Park Cooperative Deer Study and Prevention of Deer Starvation

6

June, 1971 - June, 1972

Julius G. Nagy, Principal Investigator;
David S. DeCalesta,
Student, David E. Reeder, Graduate Student.

Graduate

ABSTRACT

Six male mule deer dropped their rate of food intake 47-98 percent two to nine
days subsequent to an abrupt change from a concentrate ration to a ration of
50 percent concentrate, 50 percent alfalfa hay.
It was hypothesized that the
maximum alfalfa hay concentration in a ground, pelleted ration that deer can
survive on is 50 percent.
Six deer and six sheep exhibited similar feed intake
rates and mean retention times when fed concentrate diets containing no alfalfa
hay.
Increasing percent composition of alfalfa in the rations fed to sheep and
deer resulted in decreasing mean retention times and higher food intakes in sheep
but not in deer. Retention times under these circumstances were statistically
significantly different between deer and sheep. Rumen fluid of four deer starved
to death exhibited no statistically significant differences in counts of viable
bacteria or in vitro digestion of three rations than that from three non-starved
deer. These results suggest that rumen bacteria in starved deer neither die nor
lose the ability to digest feeds. Lack of significant differences in the parameters between starved deer permitted to eat soil and feces infected with rumen
bacteria and starved deer not provided with dirt and feces indicated that starving
deer maintain large numbers of viable rumen bacteria without reinfecting their
rumens by eating bacteria in soil or feces.
Starv~d deer exhibited little differences in weight change upon refeeding, whether refed a high-protein, low-fiber
diet or a high-fiber, low protein diet. Neither diet caused observable pigestive
upsets.
There may have been slightly better (but not statistically significant)
weight gains in deer refed the high-protein, low-fiber diet. Comparison of fawns
born from non-starved does and does starved 10-33 days revealed no statistically
significant differences in mean birth weights.
Fawns born from starved does
suffered 40 percent mortality, whereas those born from non-starved does suffered
no mortality.

IThanks are due to Dr. James A. Bailey for supervision
during my absence.

of graduate

students

��-359-

MIDDLE PARK COOPERATIVE DEER STUDY PHYSIOLOGY AND PREVENTION OF DEER STARVATION
Julius G. Nagy
Mass starvation of deer during severe winters is a reoccurring problem
in northern and western United States. It was estimated, for example,
that approximately 40 percent of the Middle Park (Colorado) deer herd
died of starvation and causes connected with starvation during the last
hard winter in 1964-65. Many of these animals died by haystacks, their
digestive organs full of undigested hay particles. During the past
decades several explanations have been offered of why and how deer die
during these stress periods with their stomachs full of food. All
explanations have been based on speculation rather than on clinical evidence.
This annual report is part of a cooperative study on the Middle Park
deer herd in which Colorado Division of Wildlife, U.S.D.A. Forest Service,
Rocky Mountain Forest and Range Experimental Station, Colorado State
University Department of Fishery and Wildlife Biology are working together
to learn more about this herd through basic and applied research.
In the segment of study reported here we have investigated the effect of
quantity of fiber on digestion, how long rumen bacteria remain functional
during starvation, length of time deer can survive starvation, and acceptable kinds of supplemental feed that can be utilized by deer during prolonged nutritional stress.
P. S. OBJECTIVE
To ascertain what takes place in deer physiology when starvation occurs
and to investigate the economic and biologic feasibility of preventing
starvation.
SEGMENT OBJECTIVES
1.

2.

To publish reports on:
a.

Responses of mule deer to alfalfa feed, and

b.

Responses of rumen microflora in mule deer to complete starvation.

To measure physiological responses of mule deer starved in pens to
refeeding with:
a.

A moderately digestible high-protein diet, and

b.

A poorly digestible low-protein diet.

�-360-

3.

To test diets for refeeding deer kept in field pens and malnourished
to simulate natural conditions during a severe winter.

4.

Prepare

report.

METHODS AND MATERIALS

Effects

of Abrupt Diet Changes

on Food Intake

Six male mule deer were placed in separate pens. They had been on an
alfalfa hay and concentrate diet ad libitum for three to four months previously.
For the first nine days of the trial a standard concentrate
ration was offered ad libitum and daily intake measured.
On the tenth day
the diet was changed abruptly to a ground, mixed and pelleted ration consisting of 50 percent concentrate and 50 percent alfalfa hay (good quality,
third cutting hay) diet. Animals were fed this diet for 11 days and daily
intake measured.

Rate of Food Passage

Through

the Digestive

Tract

Six yearling mule deer and six yearling crossbred sheep were used for the
study.
Five trials were conducted using the radioisotope chromium-51 and
a red stain, basic fuchsine, as inert food markers.
Four pelleted rations
were formulated (Appendix A) with 0, 15, 35, and 50 percent alfalfa hay
mixed in the diet. Rations were fed to deer and sheep in order 6f increasing alfalfa hay content.
The feeds were offered ad libitum and daily
food intake measured.
Each trial consisted of feeding a different ration
to all animals.
The ration with zero percent alfalfa hay was fed first
and last. A five day adjustment period was allowed for each diet and five
days for feces collection and sampling for the markers.
Mean retention
times were calculated by using cumulative excretion curves after Castle
(1956).

Effects

of Starvation

on Rumen Microbial

Activity

Nine adult deer were starved in pens until death. Three control deer were
kept on their normal ration and sacrificed at 20, 30 and 40 days after the
start of the experiment.
Deer were divided into four groups (Table 1).
Upon death of a deer viable rumen microorganisms were cultured and counted,
and in vitro digestion trials were conducted as in 1971.

�-361-

Table 1.

Division of deer into four starvation groups.

Group

N

I

4

1-60' x 40' pen, dirt floor

II

2

2-20' x
floor

6' pens, concrete

Water, sterilized soil

III

3

3-20' x
floor

6' pens, concrete

Water

IV

3 (control)

1-100' x 100' pen, dirt floor

Intake

Holding Facility

Water, soil, feces

Water and feed

Comparison of counts of viable bacteria and in vitro digestion from rumen
bacteria and in vitro digestion from rumen fluid from deer in groups I and
IV tested the hypothesis that rumen bacteria in starving deer die or lose
their ability to digest foods. A comparison of these two parameters among
groups I, II and III further tested the hypothesis that starving deer maintain large numbers of viable rumen bacteria by eating dirt and feces. If
higher counts of viable bacteria and higher in vitro digestion were exhibited
in group I deer compared to deer in groups II and III, eating soil and feces
might be a mechanism for preserving viable rumen bacteria during starvation.
If these parameters for groups I and II are similar or higher than those
for group III deer, eating dirt and feces might sustain rumen bacteria during
starvation by providing a nutrient source. If no discernable differences in
bacteria-population parameters exist among group I, II and III deer, other
mechanisms must account for maintenance of high counts of viable, functioning
rumen bacteria in starved deer.

Refeeding Trial
Beginning February 29, 1972, thirteen fawns and thirteen does were divided
into two groups, starved for 10 to 61 days (Appendix G) and refed. One group
was refed with a low fiber, high protein diet (Diet I) the other with a high
fiber, low protein diet (Diet II) (Appendix B). All deer were weighed
immediately before starvation began, the first day of refeeding and ten days
after refeeding. Daily food and water intakes were recorded for each refed
deer during the first ten days of refeeding. Condition and weights of fawns
born to starved does that survived refeeding were recorded after parturition.
RESULTS AND DISCUSSION
Effects of Abrupt Diet Changes on Food Intake
During the first day six male mule deer readily consumed the abruptly changed
diet (50% alfalfa hay, 50% ~oncentrate) b~t between the second and ninth day,

�-364-

Effects of Starvation

on Rumen Microbial

Activity

Results of in vitro digestion trials indicate that no statistically
significant differences in in vitro digestibility exist between deer
in groups I and IV and deer in groups I-III.
Counts of viable bacteria
were also not statistically significantly different between deer in
groups I and IV, and between deer in groups I and II. These counts were
statistically significantly different between deer in groups II and III and
I and III (Table 2). These data indicate:
a) that rumen bacteria in
starving deer do not die nor lose the ability to digest food; b) that
deer do not require ingestion of dirt or feces infected with rumen bacteria
to maintain large numbers of viable rumen bacteria.
It is believed that
contamination and poor technique may have led to the high counts of viable
bacteria recorded for group III deer.
To better understand effect of length of starvation on counts of viable
bacteria and in vitro digestion, data from group I, II and III deer were
pooled.
This pooling was felt justified by the lack of significant differences between group I-III in vitro digestion data and group I and II counts
of viable bacteria.
The uncertain reliability of group III count data may
justify including group III deer in this pooling.
Results of in vitro digestion trials with rumen fluid obtained from deer
starved 16-47 days and from non-starved deer are presented in Fig. 2.
Buffer solution digestion represents non-bacterial digestion whereas in
vitro digestion by starved versus non-starved deer rumen fluid represents
impairment of bacterial digestion due to starvation.
Digestion of dry
matter, cell contents and cell wall contents by starved deer rumen fluid
declined slightly as starvation length increased but remained above levels
of digestion by buffer solution.
Slopes of regressions of percent feed
digestion with length of starvation are slightly negative, but 90 percent
confidence intervals of these slopes include zero. Thus, in vitro digestion
by rumen bacteria of starved deer declined slightly, but not statistically
significant, from that of non-starved deer.
Counts of viable rumen bacteria cultured from non-starved deer were statistically significantly higher than those from deer starved 16, 20, 22 and
47 days. Counts from deer starved 29, 31 and 37 days were significantly
higher than those of non-starved deer (Appendix F). This latter finding
is difficult to understand.
Counts for the deer starved 37 days were known
to be contaminated and the correction factor used to derive the counts may
have led to the inflated counts.
Counts for deer starved 29 and 31 days may
have been contaminated and escaped detection.

Refeeding

Trial

One fawn and doe were refedDiet
I and one fawn and doe refed Diet II after
10, 17, 31 and 38 days of starvation with the assumption that after approximately 20 days of starvation refeeding would not prevent death.
Failure of
any deer to appear weak or die after 24 days of starvation and refed led us
to redesign the refeeding schedule (Appendix G).

�-365-

Table 2.

Parameters from starved and non-starved deer.
Starvation Regime
I

II

III

IV

Number of Deer

4

2

3

3

Days Starved

20, 22
22, 29

16, 22

31, 37
47

o

372.3

142.0

4191. 0

350.0

Alfalfa

37.2

43.1

29.7

36.0

Concentrate

57.3

60.4

53.5

61.5

Pelleted Feed

34.8

39.4

34.8

42.9

Alfalfa

80.8

86.3

75.9

88.4

Concentrate

93.3

90.8

90.7

94.8

Pe11eted Feed

90.4

87.6

85.1

91. 6

Alfalfa

63.8

69.3

58.2

65.7

Concentrate

81.4

80.5

78.6

84.9

Pe11eted Feed

61. 8

64.7

63.6

66.4

Mean Counts of
Viable Bacteria Per
m1 Rumen Fluid
(X 106)
Mean In Vitro
Digestion (Percent)
Cell Wall

Cell Contents

Dry Matter

�0

-:

Does

6 Fawns

-:
-:

40

Y = 7.3 + 0.61X (fawns)

CJl
CJl

/

0
...;I

~
..c::
..-lec
Q)

~

35

6

30

1M

"'C

0

~

25

0

..-l0

6

-:

~
t1I

~
t1I
~
CJl
I

Q)

~
p.,
~
~
Q)
u

~
Q)

p.,

,/

D 0/

&gt;-

!Xl

00

20

15
1°0

6

/

Y = 6.0 + 0.44x (does)
I
\.oJ

0

0\

0

6/
-:

l

/'

/~:
~

0

0

5

10
Figure 3.

20

30
40
50
60
Days Starved Before Refed
Percent pre-starvation body weight loss associated with length of starvation.

00
I

�-369-

SUMMARY AND CONCLUSIONS
As expected, the nutritional physiology of deer is somewhat different
from that of other ruminants.
Failures in emergency feeding programs
obviously were due to the fact that they did not meet the nutritional
needs of the deer - not that starving deer cannot be successfully refed.
Most feeding programs in America were necessarily based on experience
with sheep and cattle.
During the past two years we have shown that although deer rumen bacteria
are able to digest hay, their digestive organs are apparently not as
efficient in handling alfalfa hay fiber as those of sheep and goats.
Deer not in a starved condition and given a good quality (leafy) hay may
choose between the leafy and the stemmy parts and survive without complication or digestive upsets.
Healthy deer should be able to survive
on leafy alfalfa hay, since it is digestible and nutritious to other
ruminants (Johnson 1972).
Poor quality hay (mostly stems) causes severe
digestive upsets even if the hay is ground.
Reaction of the animal to
sudden diet stresses is affected strongly by previous nutritional history
of deer. Deer previously in prime condition can adjust to the stress and
survive much longer than those in poor condition.
In order to survive
the hardships of winter deer evidently need a better quality (low fiber)
forage than e.g., sheep which are able to increase their food intake and
rate of passage of ingesta through the digestive tract.
We have shown in the starvation trials that deer rumen bacteria can survive long periods of starvation.
This finding is contrary to our previous
hypothesis that if deer are subjected to complete starvation rumen microorganisms will die or pass down the digestive tract.
It seems now that
viable and functional rumen microorganisms will remain in the rumen as
long or possibly longer than the starved animal is able to sustain life.
Our studies with high fiber diets suggest that deer can be eating but
passing food through the digestive tract too slowly to extract the nutrients needed.
How long deer are able to live without food would depend again on previous
condition and very likely on other factors such as environmental temperature, wind and other stress (harrassment, etc.).
We did not study these
factors.
It should be emphasized that although we were able to starve deer
in 1972 for very long periods, in 1971 when condition of the experimental
animals was poor, deer started to die much sooner after the start of the
experiment.
Starving deer therefore should be provided with forage as
early as possible.
Fawns of does subjected to severe starvation during gestation can be
expected to suffer greater mortality than fawns of does kept on a goodto-fair nutritional plane, as previously documented by Verme (1963). This
loss therefore should be added to any winter mortality in which severe
starvation was the cause of death.
Although many starved does will survive, their fawns will be subjected to a high rate of mortality after birth.

�-370-

LITERATURE CITED

Castle, Elizabeth J. 1956. The rate of passage of foodstuffs through the
alimentary tract of the goat. 1. Studies on adult animals fed on
hay and concentrates. British J. Nutrition 10(1):15-23.
Johnson, R. R. 1972. Feedstuffs utilized by ruminants. Pages 9-34. In
D. C. Church (Editor), Digestive physiology and nutrition of rumin;,nts.
Vol. 3, Practical nutrition. O.S.U. Bookstores, Inc., Corvallis,
Oregon. 350pp.
Nagy, J. G., G. G. ,Schoonveld, and D. S. DeCalesta. 1971. Middle Park
Cooperative Deer Study--Physiology and Prevention of Deer Starvation.
Colo. Div. Game, Fish and Parks, Game Res. Div. Fed. Aid Proj.
W-38-R-24. Game Res. Rept., July. 22pp.
Schoonveld, G. G. 1971. Feeding mule deer high-fiber diets.
Colorado State University. 96pp.

M. S. Thesis.

Verme, L. J. 1963. Effect of nutrition on growth of white-tailed deer
fawns. Trans. N. Amer. Wildl. and Nat. Res. Conf. 28:431-443.

Prepared by_~-..,~J
__ ~\_~
__ ...,Dp.).,..' __ ~_ A_ ~--IEn,--~
~G.Na{;
~
Associate Professor,
Wildlife Biology

�-371-

Appendix A.
trials.

Chemical analysis of four diets fed during rate of passage

Chemical Analysis
Ether
Crude
Extract
Fiber

Diet

Percent
Alfalfaa

Protein

I

0

13.8

2.4

18.6

52.9

II

15

13.8

2.3

20.1

51.0

III

35

13.4

2.1

22.8

47.7

IV

50

13.5

2.0

24.9

45.3

NEF

b

a

The remaining portions of the compounded rations consisted of 40 percent
barley, 30 percent cottonseed hulls, 20 percent of 32 percent protein
supplement and 10 percent corn.
b

Nitrogen free extract.

Appendix B.

Feeds used in refeeding and effect of previous diet trials.
Chemical Analysis
Crude
Protein

Fat

Crude
Fiber

NFE

1

13.99%

3.97%

9.65%

57.11%

lIb

3.78%

1.32%

41.26%

39.28%

Diet
8

a

Diet I composition by percent dry weight: rolled barley = 12.5; rolled
corn
20.0; corn chop = 12.5; rolled oats = 10.0; beet pulp = 2.5; brewers
pulp = 35.0; molasses = 7.5.
b

Diet II composition by percent dry weight: cottonseed hulls
straw = 60.0.

40.0; barley

�-374-

Appendix E. Mean daily food consumption (g) of deer and sheep feeding
on diets with varying percentages of alfalfa hay.
Trial I
0% Alfalfa

Trial II
15% Alfalfa

Trial III
35% Alfalfa

Trial IV
50% Alfalfa

Trial V
0% Alfalfa

Deer 1

841

901

910

1172

788

2

1095

1598

1513

1306

500

3

488

820

1027

1062

458

4

250

474

473

721

779

5

827

951

1013

1075

798

6

685

954

1018

326

706

Means

698

933

992

944

672

Sheep 1

1313

1176

819a

2

1579

1678

1739

1881

1591

3

774

1329

1301

1464

1190

4

1469

1636

1447

1839

1609

5

1201

1319

1426

1531

1279

6

1325

1582

1550

1588

1397

Means

1277

1453

1493

1660

1413

a

Animal ceased to eat and subsequently died.

�-375-

Appendix F.

Counts of viable bacteria cultured from deer rumen fluid.
Bacteria per ml
Rumen Fluid (x105)

Total Bacteria
in Rumen (x109)

Days to Death

Group

Oa

IV

350.0

98.7

16

II

35.0

10.0

20

I

110.0

22.0

22

I

181.0

28.9

22

I

22.7

4.1

22

II

250.0

16.0

29

I

960.0

130.0

31

III

8300.0

1500.0

37

III

2400.0

850.0

47

III

82.0

3.6

a

Mean of three control deer.

�Appendix G.

Fate of starved deer.
Body Weisht Chanses
PrePostRefeedinga Refeedingb

Deer
Number

Age
Class

Days
Starved

Diet
Refed

5

FC

10

I

-19.3

+ 3.0

SSRd

8

F

10

II

- 9.7

+10.1

SSR

24

De

10

I

-10.4

+ 0.8

SSR

18

D

10

II

- 7.4

- 1.5

SSR; bore 2 fawns, both alive doing well

2

F

17

I

-19.8

+11.4

SSR

10

F

17

II

-14.6

+ 2.6

SSR

Final Disposition

I

22

D

17

I

- 7.5

+ 2.0

SSR; bore 2 fawns, both alive doing well

w

--.I
0I

15

D

17

II

-24.6

+ 17.0

SSR; bore 2 fawns, one died next day, other
doing well

7

F

24

I

-24.0

+ 4.0

SSR

3

F

24

II

-11.9

- 5.6

SSR

23

D

24

I

- 7.1

0.0

SSR

20

D

24

II

-14.9

-13.6

SSR; both fawns born died next day

1

F

33

I

-27.4

Died

Died 3.5 hours after being refed

12

F

33

II

-19.8

+ 4.1

SSR

9

F

33

None

-29.5

-

Died before could be refed

26

D

33

I

-15.3

+ 2.6

SSR; bore 2 fawns, one died 22 days age,
other missing at 19 days age

----------------------------------------------------------------------------------------------------------

�Appendix G.

Fate of starved deer (continued).
BodX Weight Changes
PrePostRefeedinga
Refeedingb

Deer
Number

Age
Class

Days
Starved

Diet
Refed

19

D

33

II

-16.4

- 1.6

SSR

4

F

34

II

-33.9

Died

Died same day refed; ate 157g feed

11

F

34

II

-28.1

Died

Died same day refed; ate unknown amount

13

F

35

I

-30.0

+ 7.5

SSR

6

F

35

II

-28.6

+ 5.3

SSR

14

D

36

II

-29.7

+ 5.4

SSR

17

D

54

None

-37.0

-

16

D

55

I

-26.0

+ 6.0

21

D

56

II

-36.3

-

25

D

61

II

-24.2

-6.0

a

Final Disposition

Died before could be refed; carried 2
normal appearing fawns
SSR
Died 5th day of refeeding; carried 2
healthy appearing fawns
SSR

Percent body weight change from pre-starved weight to day of refeeding weight.

bpercent body weight change from day of refeeding weight to 10th day of refeeding weight.
c

Fawn.

d
e

Survived starvation and refeeding.

Doe.

I
\..V

-..J
-..J
I

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                  <text>October) 1972

-1-

JOB PROGRESS REPORT

State of __

---::::C.:::.,OL:::,O:;:,;RA:.;:,::D::.,:O::......
_

Project No.
Work Plan No.
Job Title
Period Covered:

W-88-R-17
1

Migratory Bird Investigations
Job No.

1

Waterfowl Production Surveys
April 19, 1971 to June 25, 1971

Personnel: C. Bryant and staff, Monte Vista National Wildlife Refuge;
C. Hayes, D. Horne and J. Randall, Bureau of Sport Fisheries and Wildlife;
F.N. Folks, Utah Division of Fish and Game; C. Braun, D. Coven, G. Crawford,
H. Funk, R. Hopper, E. Kautz, R. Lowry, W. Miller, D. OWens, W. Russell,
K. Wagner and M. Szymczak, Colorado Division of Game, Fish and Parks.

ABSTRACT
Weather and water conditions were considered adequate to good for all major
breeding areas of the State except the San Luis Valley which had the driest
conditions observed during the past eight years. The total estimated number
of 60,691 breeding pairs was 5.8 percent above the 1970 total and 2.3 percent
above the long term average. The mallard continues to be the major breeding
species in Colorado. Production of Canada geese in northwest Colorado increased
approximately 15 percent from the 1970 level but the total post-nesting
season population size remained essentially the same for the third successive
year. A Canada goose production trend route initiated in west-central Colorado
resulted in 50 adults and 60 goslings observed. Production of Canada geese
in north-central Colorado increased 40.3 percent over the 1970 level, while
the total population increased to approximately 4,000 birds, 23.9 percent
above the 1970 level.

��-3-

WATERFOWL

PRODUCTION.SURVEYS

Michael

R. Szymczak

P. S. OBJECTIVE

To formulate

waterfowl

harvest

regulations.

SEGMENT

OBJECTIVES

To determine, through statistically
reliable sampl±ng techniques, the number
of duck and goose breeding pairs. by species, in each major Colorado waterfowl breeding ground.

METHODS

AND MATERIALS

Present duck breeding-pair
and production
inventory of only major production areas.

surveys

consist

of a breeding

pair

The 1971 duck breeding pair surveys were conducted during the period of May
10 to June 25. Surveys in North Park and the Cache la Poudre and South
Platte Valleys were conducted exclusively
from the air.
Ground counts were
made in the Yampa Valley and Brown's Park.
In the San Luis Valley intensive
aerial counts were adjusted for visibility by air-ground comparison studies.
Pair estimates for the Monte Vista National Wildlife Refuge in the San Luis
Valley were obtained from nesting transects.
All survey methods and sample
areas for ducks remained the same as in previous years.
Surveys of Canada goose production were conducted within the period of
April 19 to June 18. Production estimates for Moffat County were made from
as complete a count as possible of hatched or active nests and brood size.
Population estimates in north-central
Colorado were obtained from counts of
goslings and adults conducted from the ground during the period in which the
birds were flightless.
The west-central
Colorado production survey was
conducted from the air.
Results from the latter survey is presented in
this report for the first time.
All flying was accomplished with a Cessna 185 aircraft.
Areas sampled by
section or block were flown with one observer, while two observers were
used in sampling by transect.
On the basis of these studies. a report is submitted
Fisheries and Wildlife, which constitutes Colorado's
continental cooperative breeding ground survey.

to the Bureau of Sport
part in the annual

�-4-

RESULTS AND DISCUSSION
Weather and water conditions were variable through the major breeding areas
in the state. Light snowfall on mountain ranges surrounding the San Luis
Valley combined with essentially no winter or spring moisture on the Valley
floor produced the driest conditions observed for the past eight years.
Heavy winter snows in the mountains around North Park produced excellent
water conditions in the Park. Heavy spring rains in the Cache la Poudre
and South Platte Valleys produced fair water conditions. However, these
same late rains retarded the initiation of irrigation of farmlands, and
thus delayed the development of irrigation drainage ditches and seepage ponds
which are normally available for duck use. Unseasonably cool weather
delayed the melting of very heavy snowpacks draining into the Yampa and
Little Snake Rivers, thus allowing geese and ducks to complete their nesting
cycle prior to high water.
The total number of duck breeding pairs on Colorado's major production areas
increased slightly over the 1970 level (Table 1).
Brown's Park and the San
Luis and South Platte valleys had increases in duck populations with North
Park, the Cache la Poudre and Yampa Valleys registering declines. The 1971
breeding population was within two percent of the long term average.
Table 1. Summary of Colorado duck breeding ground population estimates in
selected areas, 1971, with 1970 and long-term averages for comparison.

Area

Total Estimated Breedin~ Pairs
Long Term
1971
1970
Averagel/

.Percent Chan~e
From
From Long
1970
Term Average

San Luis Valley

30,272

27,059

27,462

+ 11.9

+ 10.2

North Park :?:.l

14,711

15,356

19,081

4.2

- 22.9

South Platte Valley

8,672

6,912

5,852

+ 25.5

+ 48.2

Cache la Poudre Valley

3,115

4,360

3,017

- 28.6

3.2

Yampa Valley

2,340

2,350

2,898

0.4

- 19.3

Brown's Park

1,581

1,352

996

+ 16.9

+ 58.7

Totals

60,691

57,389

59,306

+ 5.8

+ 2.3

1/ San Luis Valley and North Park averages are based on results of 1964 through
1970 and 1968 through 1970 surveys, respectively, because of changes in survey
methods utilized prior to those dates. Figures for other areas are 17 year

averages.
1/ Aerial counts corrected by species fro~ visibility ratios obtained in the
San Luis Valley.

�-5-

The mallard breeding population increased slightly in total numbers, but
declined in terms of composition of the total breeding population in comparison with 1970 figures (Table 2). Significant increases were noted in
the number of blue-winged-cinnamon teal, shoveLers , and redheads. Gadwall
was the only species to register a major decline in comparison with 1970
figures. All species, with the exception of the mallard and green-winged
teal, were above the l7-year average in terms of numbers.

Table 2. Species composition of Colorado's 1971 duck breeding population,
with 1970 and 17 year average for comparison.

Number of Breeding Pairs
1954-1970
1971
1970 Averagel/

Percent Species Composition
1954-1970
1971
1970
Average

Mallard

24,150

23,255

28,355

39.8

40.5

57.9

Blue-winged and
Cinnamon Teal

7,901

4.893

4,574

13.0

8.5

9.3

Gadwall

9,884

14,989

4,877

16.3

26.1

10.0

Pintail

3,862

3,736

3,393

6.4

6.5

6.9

Green-winged Teal

1,923

2,587

2,316

3.2

4.5

4.7

Shoveler

4,354

3,461

1,940

7.2

6.0

4.0

Redhead

4,276

1,141

1,668

7.0

2.0

3.4

American Widgeon

2,640

2,636

748

4.3

4.6

1.5

Other Divers

1,701

691

1,109

2.8

1.2

2.3

Total

60.691

57.389

48,980

Species

!/ Species composition computed from data from all areas for the 17 year
period regardless of changes in survey methods.

A 1971 post-nesting season population of 1,518 geese was estimated for trend
areas for the three major river systems in Moffat County (Table 3). Included
in this total for the first time were estimates for populations utilizing
the Green River in the Dinosaur National Monument. A preliminary survey
of the Monument in 1970 indicated a significant number of geese were utilizing
the canyon-type habitat for nesting.

�-6-

Table 3. Number of Canada geese observed and estimated production, Moffat
County, Colorado, 1971.
Nesting
Pairs

Area

Non-nesting
Birds

Total
Adults

Estimated No.
Goslings 1:../

Total
Birds

Yampa River
Craig to Juniper Spgs.

14

178

206

62

268

Juniper to Cross Mtn.

13

85

111

67

178

Lily Park

14

80

108

44 2:./

152

41

343

425

173

598

28

93

149

101

250

33

187

253

136

389

Sub-total

61

280

402

237

639

Little Snake River

31

87

149

132 !!l

281

GRAND TOTAL

133

710

976

542

Sub-total
Green River
Brown's Park
Dinosaur Nat' 1. Monument

1/

1,518

!/ Calculated using average brood size and number of successful nests.

lJ Includes 20 goslings hatched from eggs taken from nests before final survey.
1/

Area first surveyed in 1970; data supplied by F. Neil Folks, Utah Division of
Fish and Game.

4/
- Includes 29 goslings hatched from eggs taken from nests before final survey.

�-7-

Excluding geese in the Monument, the MOffat County post-nesting population
remained at approximately the same level as recorded the previous two
years (Table 4). A 15 percent increase in gosling production was noted
(Table 5), but a four percent decline in adult birds negated any possible
gain in total population.

Table 4.

Total Canada geese observed, MOffat County, Colorado, 1971.

Area
Yampa River

No. Geese Counted
1956-70
1971
1970 Average

Percent Change
From 1956-1970
From 1970
Average

598

596

391

+ 0.3

+ 52.9

Brown's Park

250

285

123

-12.3

+103.3

Dinosaur Nat'1 Monumentl

389

400

281

297

234

- 5.4

+ 20.1

1~129

1,178

748

- 4.2

+ 50.9

Green River

Little Snake River2
Total3
1

- 2.8

Area first surveyed in 1970.

2 Not included in survey until 1962.
3 Dinosaur Nat' 1. MOnument not included in total.

The results of an aerial survey of a recently developed Canada goose nesting
population in west-central Colorado are presented in Table 6. This population
is composed of Great Basin Canada geese and was developed through Colorado's
restoration program, Table 6 presents the results of an aerial trend route
and is definitely not a total count of birds in the area. High water during
the census made counting difficult.
Results of the June 15, 1971 census of Canada geese in the Ft. CollinsBoulder-Denver area are presented in Table 7. Successfully hatched eggs or
goslings that were subsequently removed for restoration programs in other
parts of the state are included in totals for the respective areas.
Gosling production increased in 1971 over 1970 levels in all production trend
areas except Wellington where production remained stable (Table 8). Increases
were substantial in the Loveland, Boulder, and Denver areas.

�-8-

Table 5.
1971.

Estimated number of Canada goose goslings, MOffat County, Colorado,

No. of Gos1in~s
1956-70
1971
1970 Average

Area
Yampa River

Percent Change
From 1956-1970
From 1970
Average

173

140

140

+ 23.6

+ 23.6

Brown's Park

101

117

45

- 13.7

+124.4

Dinosaur Nat'1 Monument 1

136

90

132

97

74

+ 36.1

+ 78.4

406

354

259

+ 14.7

+ 56.8

Green River

Little Snake River

2

Total
1
2
3

+ 51.1

Area first surveyed in 1970.
Not included in survey until 1962.
Dinosaur Nat'l. Monument not included in total.

Table 6.

Canada goose production survey, west-central Colorado, 1971.

Area

No. of Canada Geese Observed
Adults
Goslings

Colorado River
Silt to Grand Junction

17

21

Grand Junction to Utah Border

26

28

7

11

50

60

Gunnison River
Delta to Whitewater
Total

�-9-

Table 7.

Results of the north-central Colorado goose census, June 15, 1971.

Production
Area

Wellington

Water Area

Terry Lake
Launer Pond
Douglas Lake
North Poudre No. 1
Dry Creek Reservoir
North Poudre No. 3
North Poudre No. 2
North Poudre No.5
Bureau of Standards Pond
No. 1
Divide No. 8
Elder Reservoir
Annex No. 8
Van Sant Pond
Cobb Lake
Hinkley Reservoir
Dale Pond
Watson Lake
Curtis Lake

No. Total No.
Total No.
Total No.
Broods Goslings Adults &amp; Yrlgs. of Birds

3
2
3
0
0
1
2
2
5
5
0
1
2
1
2

Total
Fort Collins

Peterson Pond
He rring Lake
College Lake
Dean Acres
C1aymo re Lake
Sterling Gravel Pits
Lindenmeier Lake
Grey Lakes
Novak Reservoir
Winick Ponds
Anderson's Pond
Timnath Reservoir
Romily Gravel Pit
Parkwood Lake

1

3
4
2
1
1
1
2
1

Total
Loveland

Total

Flatiron Reservoir
Boedeaker Reservoir
Flatiron Gravel Pits
McNeil Reserv9ir
Welch Reservoir

1
2
4

87 1/
14
6
10
0
0
5
8

93
6
9
39
11
3
2
4

180
20
15
49
11
3
7
12

6

38
10
3
62
7

9
45
72
17
2
83
4
11
63
48

15
66
90
17
8
121
14
14
125
55

301

521

822

5
18
108 !/
28
18
25
4
2
5
3
19
11
4
5

2
12
146
21
60
61
41
2
2
2
18
51
12
5

7
30
254
49
78
86
45
4
7
5
37
62
16
10

255

435

690

3
36
10
58
18

2
60
4
48
10

5
96
14
106
28

125

124

249

21
18
0
6

----------------------------------------------------------------------------------

�-10-

Table 7.

Results of the north-central

Production
Area

Water Area

Boulder

Colorado goose census, June 15, 1971 (cont.).

No. Total No.
Total No.
Total No.
Broods Goslings Adults &amp; Yrlgs. of Birds

Valmont Reservoir
Terry Lake
Crystal Lake
Ish Lake
Faivre Ponds
Boulder Valley Farm
Saw Hill Ponds

Total
Denver

Total
Grand Total

S. Colo. Blvd. &amp; Quincy
B lackme r Lake
Reservoir No. 3
Denver City Park
Sloans Lake
Kountze Reservoir
Federal Center Pond
Clairfield Reservoir
Gallup Reservoir
Columbine Country Club
Bowles Lake
Kings Pond
Lowe r Tule Lake
Marston Reservoir
Grant B Reservoir
Grant C Reservoir

o

191
53
10
10
34
22
37

360
32

551

9

30
36
15
23

19
40
70
37
60

357

505

862

o

49
26
23
212
50
110
13
24
89

49
35
32
252
90
110
53
63
107

9
3

o
1

4

o

9

40
40

o
40 ]:/
39
18

o

85

5

5

o

234
105

57

o

o

5

58

4

17
22

71
24
57

291
105
63
71
41
79

296

1,150

1,446

1,334

2,735

4,069

o
1

1/ Includes goslings produced from eggs collected on the area.

A definite reduction was noted in the number of adult and sub-adult birds in
the Wellington and Ft. Collins areas. However, the total number of Canada
geese observed throughout the area in 1971 was approximately 24 percent above
the 1970 total (Table 9).

�-11-

Table 8. Total number of Canada goose goslings produced in north-central
Colorado production trend areas, 1971.

Area

No. of Goslings
1969-70
1971
1970 Average

Percent Change
From 1969-70
From 1970
Average

Wellington

301

301

279

Fort Collins

255

238

247

+

Loveland

125

63

50

+ 98.4

+150.0

Boulder

357

247

198

+ 44.5

+ 80.3

Denver

296

102

264

+190.2

+ 12.1

Total

1,334

951

1,038

+ 40.3

+ 28.5

0.0

+

7.9

7.1

+

3.2

Table 9. Total number of Canada geese observed in north-central Colorado
production trend areas, 1971.

No. of Geese
1969-70
1971
1970 Average

Percent Change
From 1969-70
From 1970
Average

Wellington

822

915

930

- 10.2

- 11.6

Fort Collins

690

681

715

+

1.3

3.5

Loveland

249

168

154

+ 48.2

+ 61. 7

Boulder

862

694

662

+ 24.2

+ 30.2

Denver

1,446

827

1,286

+ 74.8

+ 12.4

Total

4,069

3,285

3,747

+ 23.9

+

Area

8.6

�-12-

FALL FLIGHT PREDICTION

Dry conditions in the major duck breeding area, the San Luis Valley, will
definitely retard Colorado's total production. The results should be the
smallest fall flight of ducks from Colorado since 1966. Canada goose
populations in northwest Colorado and north-central Colorado remain
essentially stable.

Prepared by -7lLU"-L~;:::::':....!. ~:...-....:·7&lt;:L..::~_L~'-'9=~:...cz:l
-~_
£/:L-__
Michael R. szym~
Assistant Wildlife Researcher

�October, 1972

-13-

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~---------

Project No.
Work Plan No.
Job Title
Period Covered:

W~-_8~8_-~R~-~1~7
_
1

Migratory Bird Investigations

Job No.

2
-------------------------------------

Trapping and Banding Ducks and Geese
April 1, 1971 to March 31, 1972.

Personnel: J. Randall, Bureau of Sport Fisheries and Wildlife; Dr. R. Ryder,
Colorado State University; M. Babler, C. Braun, R. Clippinger, J. Corey,
D. Coven, C. Crawford" G. Crawford, R. Desilet, J. Ellenberger, G. Eyre,
H. Funk, T. Henry, R. Kitzmiller, W. Ladd, Jr., C. Leonard, J. Lorentzson,
T. Lytle, B. Petersen, T. Rauch, M. Robinson, Wo Rutherford, J. Seidel,
D. Snyder, M. Szymczak, D. Todd, K. Wagner, J. Wolfe, R. Hopper, Colorado
Game, Fish and Parks.

ABSTRACT
Trapping activities during Segment 17 resulted in the banding of 12,630 ducks
of 10 species at seven locations in Colorado. Mallards were again the major
species, contributing 9,420, or 75 percent of the total. Pre-season efforts
involved the initiation of a new study aimed at investigating relationships
among the three major high country production areas (North Park, South Park,
San Luis Valley) in regard to migration routes, harvest areas, and mortality
rates. Information derived from this study will help us to evaluate the
justification for establishing duck hunting season dates in Colorado based
on elevation. Banded samples for these high country areas in 1971 amounted to
2,286 in North Park, 1,593 in South Park, and 3,101 in the San Luis Valley.
Post-season banding (Winter) yielded 6,564 mallards to the banded sample.
Nearly 2,600 Canada geese were banded during Segment 17. Goslings, transplanted to various locations during the summer, contributed 504 to this
total. The rema1n1ng birds were banded post-season in the Arkansas Valley
(1,032) and the Cache 1a Poudre Vall~y (1,043).

�-14-

RECOMMENDATIONS

1.

Continue pre-season duck banding in North Park, South Park and the San
Luis Valley, Quotas should include 1,000 mallards, 1,000 pintails and
at least 500 each of the other major species in each of the three areas.
Attempt to obtain equal age and sex ratios in the sample by species.
Experiment with new capturing techniques, such as night lighting in an
effort to increase sample sizes of species other than mallards and
pintails.

2.

Initiate a post-season mallard banding program in the UncompahgreGunnison-Colorado River complex. Obtain a quota of up to 1,000 birds,
divided about equally by age and sex.

3.

Continue goose banding program as outlined in the W-88-R Job Description
for Segment 17.

�-15-

TRAPPING AND BANDING DUCKS AND GEESE
Richard M. Hoppe r
This report summarizes all waterfowl banding activities of Federal Aid
Project W-88-R-17 for the Segment year April 1, 1971 to March 31, 1972. Band
recovery data are analyzed separately under another job (Work Plan 1, Job 3),
and work of this nature is omitted from this report. This report simply
presents a tabulation of numbers of ducks and geese banded by species and
location during the Segment.

P. S. OBJECTIVE
To formulate waterfowl harvest regulations for Colorado.
SEGMENT OBJECTIVES

.

1.

To provide adequate samples of banded ducks and geese in Colorado's major
breeding and harvest areas for purposes of accumulating migration and
annual mortality data.

2.

To report species and number of ducks winter banded in the South Platte
Valley, Arkansas Valley, and Bonny Reservoir as part of Work Plan 3, Job 6.
METHODS AND MATERIALS

The overall banding program remained similar to past years with operations
limited to three phases: (1) mid-summer goose transplant banding, (2) late
summer (pre-season) duck banding, and (3) winter (post-season) duck and goose
banding. Goslings for transplant purposes are hand-reared and also obtained
from the wild through drive-trapping or by using baited walk-in traps prior
to the time the birds are capable of flight. Pre-season duck banding is
conducted in August and September and utilizes baited Salt Plains traps
(cage-type) to capture birds in North Park, South Park, and the San Luis
Valley. Post-season banding (January-February) presently limited to the
eastern slope, employs baited Salt Plains traps and cannon-nets for duck
trapping, and baited cannon-nets and a large walk-in trap for goose trapping.
RESULTS AND DISCUSSION
Ducks
Numbers and species of ducks banded during Segment 17 by State personnel are
listed in Table 1 by location. The banded sample amounted to 12,630 ducks
of 10 species in seven locations involved. This was an increase of about
1,000 ducks over the previous segment. largely because of the addition of North
Park to the list of banding sites and the increase of quotas during postseason banding.

�Table 1.

Number of ducks banded by species, location, and period of year, 1971-72.

Species

North
Park

Pre-season!'!
South
San Luis
Park
Valley

Cache 1a
Poudre Valley

Post-season!.!
South Platte
Bonny
Valley
Reservoir

Arkansas
Valley

Total

1,333

9,420

Mallard

1,105

1,079

672

831

3,400

1,000

Gadwall

1

1

17

-

-

-

American Widgeon

13

3

-

-

-

Green-winged Teal

231

163

413

-

-

-

807

33

100

102

-

-

-

235

Blue-winged

19
16

and

Cinnamon Teal

...•
I

0\
I

Pintail

879

245

950

-

-

-

2,074

Redhead

24

0

33

-

-

-

57

Canvasback

-

1

-

-

-

Wood Duck

-

1

-

-

-

-

-

1

2,286

1,593

2,187

831

3,400

1,000

1,333

12,630

Total

!/ August-September.
l/ January-February.

1

�-17-

The mallard, again, was the major species represented, contributing 9,420, or
75 percent of the total. Only mallards were banded during the post-season
period, and this species was also the most prominent during pre-season
efforts. Pintai1s and green-winged teals were the only other species that
contributed significantly to the banded sample, with 2,074 and 807, respectively.

High Country Study
Pre-season banding efforts in 1971 marked the beginning of an overall investigation of high country duck populations in North Park, South Park, and the San
Luis Valley. Samples of ducks will be banded annua11y'in all three areas
during the same period for the next five years. Attempts will be made to
band good samples of each major species, with approximately equal numbers by
age and sex. The objective is to determine relationships among the three
areas in regard to migration routes, harvest areas, and mortality rates.
Information derived from this study will allow us to evaluate the justification
for establishing duck hunting season dates in Colorado based on elevation.
This can be termed "altitudinal zonation" of hunting season dates. This could
provide earlier and separate season dates for the high country, thereby
yielding additional duck hunting opportunity where little occurred before.
Tables 2-5 show the age and sex composition by species for 6,980 ducks banded
in the three high country areas during the first year of work. The San Luis
Valley figures include bandings by both State and Federal personnel. The
Bureau of Sport Fisheries and Wildlife is cooperating in this study by banding
a portion of the Valley quota on the Monte Vista and Alamosa National Wildlife
Refuges. Results of the first years work were not completely satisfactory
because of the lack of adequate banded samples for species other than the
mallard and pintail. Presently our traps are selective toward the mallard
and pintail, but means must be found to capture other species in significant
numbers if this study is to be wholly successful. Other species for which
good samples are particularly needed include gadwall, American widgeon, and
the three teal. Night lighting will be attempted in North Park during next
Segment as a possible technique for capturing these species.
Winter Banding
Post-season mallard banding was continued as part of the investigation of
eastern Colorado mallard management units (Work Plan 3, Job 6). Trapping
conditions were good, reSUlting in the banding of 6,564 mallards during a
30-day period from mid-January to mid-February (Table 1). Quotas were reached
or slightly exceeded in all but one of the eight banding areas.
Geese
Nearly 2,600 Canada geese were banded during Segment 17 (Table 6). Summer
transplant geese accounted for 504 of this total. These goslings were banded

�-18-

and released at Totten Reservoir in the Dolores Valley (38), Lake John Annex
in North Park (241), San Luis Lake in the San Luis Valley (123), and Jumbo
Reservoir Annex in the South Platte Valley (102). Work Plan 2, Job 2 covers
the transplant program in more detail.
Two thousand and seventy-five Canada geese were banded post-season in the
Arkansas and Cache la Poudre Valleys. These birds were banded at Turks
Pond and Two Buttes Reservoir in the Arkansas Valley (1,032) and at New
Windsor Reservoir in the Cache la Poudre Valley (1,043).

Wildlife Researcher

�-19-

Table 2.
1971.

Sex and age composition of ducks banded pre-season in North Park,

AM

1M

Age and Sex
AF

IF

UNK.

Total

Mallard

245

311

237

312

0

1,105

Gadwall

0

0

0

1

0

1

American Widgeon

0

9

1

3

0

13

Green-winged Teal

61

79

41

50

0

231

0

14

2

17

0

33

Pintail

205

239

223

210

2

879

Redhead

5

3

13

3

0

24

Total

516

655

517

596

2

2,286

Species

Blue-winged and
Cinnamon Teal

Table 3.

Sex and age composition of ducks banded pre-season in South Park, 1971.

AM

1M

Age and Sex
AF

IF

UNK.

Mallard

411

203

373

92

0

1,079

Gadwall

0

1

0

0

0

1

American Widgeon

0

1

2

0

0

3

Green-winged Teal

75

28

26

34

0

163

Cinnamon Teal

29

34

2

34

1

100

Pintail

70

72

28

74

1

245

Canvasback

0

1

0

0

0

1

Wood Duck

1

0

0

0

0

1

586

340

431

234

2

1,593

Species

Total

B1ue-winged and

Total

�-20-

Table 4. Sex and age composition of ducks banded pre-season
in the San Luis
Valley, 1971.

AM

1M

Age and Sex
AF

IF

UNK.

Mallard

312

353

305

254

2

1,226

Gadwall

0

4

4

12

0

20

241

84

41

55

0

421

Cinnamon Teal

39

49

14

30

0

132

Pintail

359

286

346

258

0

1,249

Redhead

3

24

5

21

0

53

Total

954

800

715

630

2

3,101

Species

Green-winged Teal

Total

Blue-winged and

Table 5. Sex and age composition
of ducks banded pre-season in the three highcountry areas combined, 1971.

AM

1M

Age and Sex
AF

IF

UNK.

Total

Mallard

968

867

915

658

2

3,410

Gadwall

0

5

4

13

0

22

American Widgeon

0

10

3

3

0

16

Green-winged Teal

377

191

108

139

0

815

Cinnamon Teal

68

97

18

81

1

265

Pintail

634

597

597

542

3

2,373

Redhead

8

27

18

24

0

77

Canvasback

0

1

0

0

0

1

Wood Duck

1

0

0

0

0

1

2,056

1,795

1,663

1,460

6

6,980

Species

Blue-winged and

Total

�-21-

Table 6.

Number of Canada geese banded by location and period of year, 1971-72.

Number Banded
Location
Arkansas Valley
Cache la Poudre Valley
Dolores Valley

Summer
Transplants

°
°

38

North Park

241

San Luis Valley

123

South Platte Valley

102

Total

504

1/ June-July.

IJ January

1./

Post-season J:./

Total

1,032

1,032

1,043

1,043

°
°
°
°

38

2,075

241
123
102

2,579

��October, 1972

-23-

JOB PROGRESS REPORT

State of

-=-C,;;:.OL::.O:::.;RAD=:::..:O::...._

Project No.
Work Plan

W-88-R-17

1
Job No.
-----------~---------

Job Title

3

Analysis of Waterfowl Banding Data

Period Covered:
Personnel:

Migratory Bird Investigations

April 1, 1971 to March 31, 1972.

Dale Coven, Ed Kautz, Michael Szymczak and Richard M. Hopper.

ABSTRACT

Banding and recovery tapes were obtained from the Bird Banding Laboratory
for all banding from North Park, South Park and the San Luis Valley and
all recoveries from those bandings through the 1969-70 hunting season.
Programs were written and the desired data were retrieved from the tapes
through computers.
Preliminary tabulations were prepared for much of the San Luis Valley data,
but none of this information is currently ready for presentation. Work in
Segments 18 and 19 will include preparation of final and updated analyses
for all three high mountain areas.

�-24-

RECOMMENDATIONS
1.

Continue pre-season duck banding in North Park, South Park and the
San Luis Valley, with existing quotas by species, sex and age.

�-25-

ANALYSIS OF WATERFOWL BANDING DATA
Richard M. Hopper

P. S. OBJECTIVE
To formulate waterfowl harvest regulations for Colorado.

SEGMENT OBJECTIVES
To investigate migration, mortality, recovery distribution, and relationships among populations of ducks banded in Colorado's high country,
specifically in North Park, South Park, and the San Luis Valley.
METHODS AND MATERIALS
All data were obtained from computer tapes supplied by the Bird Banding
Laboratory of the Migratory Bird Populations Station, Bureau of Sport
Fisheries and Wildlife. These tapes contained all bandings from North
Park, South Park, and the San Luis Valley and all recoveries from those
bandings through the 1969-70 hunting season. Programs were written and
the desired information was retrieved from the banding and recovery tapes
through the use of automatic data processing machines.
RESULTS AND DISCUSSION
The following information was obtained on print-out sheets (tabs) of
specific format from the banding and recovery tapes:
1.

Number of birds banded by year and species (species included mallard,
gadwall, American widgeon, green-winged teal, blue-winged teal,
cinnamon teal, blue-winged or cinnamon teal, shoveler, pintail, redhead, lesser scaup, and ring-necked duck).

2.

Number of recoveries by year, species, age, and sex.

3.,

Recovery tables by year of banding and season of recovery after banding
for each species, sex and age (some species).

4.

Recovery locations by degree block, species, sex, and age (some species)
for first year and all year recoveries.

5.

Banding locations of recoveries for some species by degree block, age,
and sex for pre-season bandings~ including first year and all year
recoveries during the following periods: Oct. 1-10, 11-20, 21-31, Oct.Jan., and Nov.-Jan.

�-26-

6.

Banding locations of Colorado recoveries (by location) for some
species by degree block, age, and sex for pre-season bandings,
including both first year and all year recoveries.

7.

Banding locations of Colorado recoveries (by location) for some
species by degree block, age, and sex for winter bandings~ including both first year and all year recoveries.

8.

Percentage of recoveries occurring at each recovery location by
banding location, species, sex, and age (some species) for both
first year and all year recoveries.

9.

Summary of Colorado recoveries of bandings from everywhere by
species and recovery location.

An analysis of North Park banding data was recently completed by Szymczak
(1971). This analysis was done through use of the print-outs listed above.
Preliminary tabulations have been prepared for much of the San Luis Valley
banding and recovery data on the print-out sheets, but none of this information
is currently ready for presentation. Final and updated analyses will hopefully
be conducted for all three high mountain areas in Segments 18 and 19.
LITERATURE CITED
Szymczak, M. R. 1971. Analysis of waterfowl banding data. Colo. Div.
Game, Fish and Parks, Fed. Aid Game Res. Rpt., Oct. pp. 19-50.

Prepared by --~~~i~~~~~a~·~~~~~~·~~·~~-o-pp~~~r~·~~~L~~~
Wildlife Researcher

�October, 1972

-27-

JOB PROGRESS REPORT

State of
Project No.
Work Plan No.
Job Title
Period Covered:

~CO.;;;.L;;:.O.;;;.RAD=:...;O:..._
_
W-88-R-17
1

Migratory Bird Investigations
Job No.

12

San Luis Valley Cooperative Mallard Investigation
April 1, 1971 to March 31, 1972

Personnel: D. Horne and J. Randall, Bureau of Sport Fisheries and Wildlife;
C. Bryant and Staff, Monte Vista National Wildlife Refuge; C. Braun, E. Kautz,
W. Russell, M. Szymczak, K. Wagner and R. Hopper, Colorado Division of
Wildlife.

ABSTRACT
The duck breeding population in the San Luis Valley was estimated at 30,176
pairs for 1971, including 12,311 pairs of mallards. This represented an
increase of about 850 pairs for all species combined, and a decrease of
about 1,300 pairs of mallards from 1970. The mallard estimate was about
1,250 pairs below the 7-year average. Gadwalls and pintails showed significant decreases in 1971 from 1970, but shovelers and the three species of
teal were up over 1970.
Progress is being made in evaluating the 8-year study, with both Bureau
and State personnel contributing to all phases of the data analysis and
preparation of the final report.

��-29-

SAN LUIS VALLEY COOPERATIVE MALLARD INVESTIGATION
Richard M. Hopper

An intensive study of the mallard population in the San Luis Valley was
initiated in 1963 as a cooperative investigation between the Bureau of
Sport Fisheries and Wildlife and the Colorado Division of Wildlife. An
important part of this study has been an annual experimental hunting
season held during the period October 1-18. The eighth and final experimental season was conducted in 1970. Preliminary results of these seasons,
as well as all phases of the study, were presented in Administrative Report
No's. 42, 49, 79, 120, 130, 158, 175, 195, and 210 prepared by members of
both agencies and distributed by the Bureau of Sport Fisheries and Wildlife,
Branch of Wildlife Research, Migratory Bird Populations Station.
Segment 17 (1971) marked the final year of field work for the overall
investigation. This involved conducting the last intensive survey of the
breeding population of ducks in the spring of 1971; thus, completing the
assessment of breeding numbers following each experimental season. Also,
work during Segment 17 included initial steps in evaluating the results of
the overall cooperative study in preparation for writing final job report.
P. S. OBJECTIVE
1.

To develop a harvest formula for the San Luis Valley mallard population.

2.

To determine hunter reaction to various types of hunting regulations.
SEGMENT OBJECTIVES

1.

Estimate the size of the 1971 breeding population of ducks in the San
Luis Valley.

2.

Evaluate results of the experimental hunting seasons since 1963 and
begin preparation of final reports covering all phases of the investigation in cooperation with the Bureau of Sport Fisheries and Wildlife.
METHODS AND MATERIALS

Methods for conducting the 1971 breeding popUlation survey and air-ground
comparison study remained the same as those employed since 1967 (Segment
13). A discussion of the procedures appears in a previous report (Hopper
and Rutherford 1968) and will not be repeated here.

�-30-

RESULTS AND DISCUSSION
Breeding Population Survey
Air-Ground Comparison Study
Air and ground counts of ducks are compared by species in Table 1 for
122.5 miles of selected transects. The proportion identified from the
air (visibility ratio) for the mallard for example, indicated that the
air crew identified, as mallards, 23.5 percent of the mallards actually
present on the air-ground comparison transects. These ratios were applied
to the regular air transect data as correction factors in projecting
estimates of total breeding pairs by species for Russell Lakes and the
major portion of the Valley, excluding Monte Vista National Wildlife Refuge
and the total-count areas.

Table 1. Air-ground comparison of ducks counted on 122.5 miles of transect
in the San Luis Valley, 1971.

Species

Estimated Breeding Pairs
Ground
Air
Pairs
Percent
Pairs
Percent

Mallard

434

46.9

102

50.7

0.235

Gadwall

114

12.3

22

10.9

0.193

Pintail

52

5.6

17

8.5

0.327

Shoveler

44

4.8

8

4.0

0.182

Blue-winged Teal

44

4.8

3

1.5

0.068

Cinnamon Teal

139

15.0

13

6.5

0.094

Green-winged Teal

46

5.0

9

4.5

0.196

American Widgeon

5

0.5

2

1.0

0.400

Redhead

32

3.4

4

2.0

0.125

Other Divers

16

1.7

21

10.4

0.333 1/

Totals

926

100.0

201

100.0

0.217

1/
- Estimate based on previous years.

Proportion
Identified
From the Air

�-31-

Duck Breeding Population Estimate
Amount of habitat, sample sizes, and estimated number of breeding pairs
of ducks are shown in Table 2 for the six areas included in the 1971
survey. The 1970 estimate plus the 1964-1970 averages are included for
comparison. The 1971 estimate amounted to 30,176 pairs of breeding ducks,
or about 850 pairs above the 1970 estimate and about 2,300 pairs more
than the 7-year average (1964-1970). The estimate for 1971 was the
second highest since intensive surveys began in 1964. The largest figure,
33,035 pairs, was obtained in 1964.
Species composition of the 1971 San Luis Valley breeding population is
presented in Table 3, along with 1970 figures and the 7-year averages.
The mallard estimate for 1971 was 12,311 pairs, a decrease of about 1,300
pairs from the 1970 figure of 13,662 pairs. As usual, the mallard was
the dominant breeding species, contributing about 41 percent to the total
population estimate. Gadwalls were next in importance, with almost 5,000
pairs, or 16.5 percent of the total. This represented a significant
decrease from 1970, but was well above the 7-year average. Pintail
numbers decreased in 1971, while shovelers and the three species of teal
all showed increases. Sampling error, expressed as a percentage (coefficient
of variation) was slightly larger in 1971 (11.8%) than in 1970 (10.0%), but
remained in line with the 7-year average (12.3%).

Preparation of Fi;nal Report
Work was initiated in Segment 17 on tabulations and analysis of data covering
all phases and years of the cooperative study. Both State and Bureau personnel were assigned to work on various segments of the data and report.
A report outline was prepared and sent to the contributors for approval.
All persons concerned are now actively engaged in the preparation of this
final report. We are hopeful that this paper can be published as a Wildlife
Monograph.

LITERATURE CITED
Hopper, R. M., and W. H. Rutherford. 1968. San Luis Valley cooperative
mallard investigation. Colo. Game, Fish and Parks Div., Fed. Aid
Game Res. Rpt., Oct. pp. 33-42.

Prepared

by

i;&lt;kj)??7 2bJjL'chard M. Hopper
Wildlife Researcher

�Table 2. Breeding pairs by density type as estimated from the San Luis Valley regular air transects,
and nesting transects, 1971, with 1970 and 7-year averages for comparison.
Estimated No. of Breeding Pairs 11
7-Year Average
1971
1970
(1964-1970)

Square Miles
Habitat

Percent
Sample

Monte Vista NWR

22

2.01

3,633

4,367

3,894

Russell Lakes

6

33.33

1,428

489

1,088

San Luis Lakes

7

100.00

722

456

300

Mishak Lakes

4

100.00

180

263

306

Adams Lake

1

100.00

58

19

34

Type

27

I

w

N

Remainder of Valley

1,265

18.02

24,155

23,734

22,235

Totals

1,305

18.57

30,176

29,328

27,857

11 Determined from nesting transects on Monte Vista National Wildlife Refuge and from aerial transects
on all other areas.

11 Only 6-year averages (1965-1970) for San Luis Lakes and Mishak Lakes and a 5-year average (1966-1970)
for Adams Lake.

I

�Table 3. Species composition of the San Luis Valley duck breeding population, 1971, with 1970 and
7-year averages for comparison.

1971

%

7-Year Average
~1964-1970~
Pairs
%

Species

Pairs

%

1970
Pairs

Mallard

12,311

40.8

13,662

46.6

13,583

48.9

Gadwall

4,986

16.5

6,379

21. 7

3,094

11.2

Pintail

2,136

7.1

2,804

9.6

3,315

11.9

Green-winged Teal

1,076

3.6

557

1.9

1,457

5.2

Cinnamon Teal

2,788

9.2

2,597

8.8

2,282

8.2

880

3.0

276

0.9

796

2.9

2,305

7.6

1,642

5.6

1,326

4.8

38

0.1

782

2.7

160

0.6

3,049

10.1

518

1.8

1,494

5.4

607

2.0

III

0.4

254

0.9

30,176

100.0

29,328

100.0

27,761

100.0

Blue-winged Teal

I

Shoveler
American Widgeon
Redhead
Other Divers 1/
Totals
Coefficient of Variation

l/

11.8

10.0

-1/ Includes lesser scaups, canvasbacks, ruddy ducks, and common mergansers.
1/ Excludes Monte Vista NWR and Russell Lakes.

12.3

w
w
I

��October, 1972
-35-

JOB PROGRESS REPORT

State of

COLORADO
------------~~~~~-----------

Project No.

W-88-R-17

Work Plan No.
Job Title

1
Job No.
14
Determination of Methods for Developing and Managing
Waterfowl Habitat - Ammonium Nitrate Pothole Blasting Study

Period Covered:
Personnel:

Migratory Bird Investigations

April 1, 1971 to February 28, 1972

Richard M. Hopper

ABSTRACT
A paper entitled '~aterfowl Use in Relation to Size and Cost of Potholes"
was published in the April, 1972 issue of The Journal of Wildlife Management
36(2):459-468. This publication constitutes the Job Final Report for items
(d) and (e) under the P.S. Objective and satisfies Federal Aid requirements
for those portions of the study.

�-36-

RECOMMENDATIONS
1.

Either the 7S-lb. or ISO-lb. charge of ammonium nitrate-fuel oil mixture
is recommended for use in pothole blasting. Both were far superior to
either the 2S-lb. or 50-lb. size in terms of efficiency.

�-37-

AMMONIUM NITRATE POTHOLE BLASTING STUDY
Richard M. Hopper

P. S. OBJECTIVE
To evaluate various size potholes blasted with ammonium nitrate in terms
of (a) life expectancy, (b) plant succession, (c) soil and water characters,
(d) waterfowl use and hunting potential, and (e) cost.

SEGMENT OBJECTIVES
Compile and analyze all data collected during the course of the study and
prepare a final report covering all phases except that related to "life
expectancy" of the potholes.

METHODS AND MATERIALS
Dr. David Bowden, statistician at Colorado State University, was consulted
regarding the appropriate statistical tests to apply to the pothole blasting
data. These tests were conducted and the overall results were evaluated. A
final report was prepared, following The Journal of Wildlife Management
format and standards.

RESULTS AND DISCUSSION
A manuscript entitled ''Waterfowl Use in Relation to Size and Cost of Potholes"
was published in the April 1972 issue of The Journal of Wildlife Management
36(2):459-468. This publication constitutes the Job Final Report for items
(d) and (e) under the P. S. Objective for Work Plan 1, Job 14, and thereby
satisfies Federal Aid requirements for those portions of the study. All
other items will be covered in the next segment.

Prepared

by

~r

k

~

Wildlife Researcher
Migratory Bird Investigations

��October,

-39-

1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.
Work

W-88-R-17

Migratory

2

Plan No.

Job No.

Job Title Experimental

Studies

Period Covered:

1, 1971 to March 31, 1972

April

on Improving

Bird Investigations
2

Status of Canada Goose Populations

Personnel:
C. Hayes and D. Horne, Bureau of Sport Fisheries and Wildlife,
D. Coven, J. Corey, D. Crane, C. Crawford, G. Crawford, G. East, J. Ellenberger, H. Gresh, R. Hopper, T. Lines, J. Monarch, T. Rauch, S. Steinert,
L. Webster and M. Szymczak, Colorado Division of Game, Fish and Parks.

ABSTRACT
In the spring of 1971, at least five nests were noted in the K-4 Ranch transplant area near Masters, Colorado.
The transplant program at Jumbo Reservoir
in the South Platte River Valley in extreme northeast Colorado was initiated
in 1971, with the release of 102 Canada goose goslings.
Only one of the
birds released at Jumbo was reported recovered during the 1971-72 hunting
season.
Some Canada goose pairs were noted to be frequenting artificial
nest structures at San Luis Lake but no broods were observed in the area.
One hundred and twenty three goslings were released at San Luis Lake in 1971,
raising the total number of birds released at that site to 351. A portion
of the birds released at San Luis Lakes continue to use the Bitter Lakes
National Wildlife Refuge in southeast New Mexico during the winter.
The first
group of Canada geese observed in the spring of 1971 in North Park were
sighted on Lake John Annex on April 10. The largest group of geese, approximately 50 birds, was sighted in late May on the Annex.
Wheatland Reservoir,
north of Laramie in Wyoming, continues to be used as a molting area by birds
released at Lake John Annex.
An additional 241 goslings were released on the
Annex in July of 1971. Riverside and Imperial counties of extreme southern
California and adjacent Yuma County, Arizona were the major recovery areas
during the 1971-72 season for geese released at Lake John Annex.
About 19
percent of the birds released at Lake John Annex in 1971 were reported
recovered during the 1971-72 season.
In July, 1971, 38 goslings were added
to the Totten Reservoir flock near Cortez in Montezuma County.
In mid-December
the majority of the birds released at Totten left the area and apparently
moved straight south, terminating migration on the Gila River in Graham
County, where a total of eight birds were reported taken by hunters.

��-41-

EXPERIMENTAL STUDIES ON
IMPROVING STATUS OF CANADA GOOSE POPULATIONS
Michael R. Szymczak
P. S. OBJECTIVE
To investigate the success of attempts to establish breeding populations of
Canada geese in suitable habitat where they do not currently exist in Colorado.

SEGMENT OBJECTIVES
1.

To prepare a technical bulletin on the technique of establishing a
breeding population of Canada geese.

2.

To expand the breeding flocks of Canada geese in the South Platte Valley,
the San Luis Valley, and North Park.

3.

To continue efforts to establish a breeding flock of Great Basin Canada
geese on Totten Reservoir, south of Dolores in MOntezuma County.
METHODS AND MATERIALS

Canada goose goslings for transplant purposes were obtained from four different
sources: (1) goslings from the metropolitan Denver area; (2) goslings and
some eggs from the Fort Collins area; (3) goslings from Valmont Reservoir and
Fairvre Ponds in the Boulder area, and (4) eggs from nests in danger of destruction from high water along the Yampa and Little Snake Rivers in MOffat
County. All eggs were transported to the Fort Collins Wildlife Research
Station for incubation and subsequent raising. All goslings live-trapped, were
transported as soon as possible to the release site. Birds obtained from
eggs collected in Moffat County were released on Totten Reservoir south of
Dolores. Birds collected at other locations were released either at San Luis
Lakes, Lake John Annex or the K-4 Ranch.
Migration and mortality information were obtained from recovery cards received
from the Migratory Bird Population Station.

RESULTS AND DISCUSSION
Preparation of Technical Bulletin
Because of an increasing work load no progress was made on preparIng a technical bulletin on the technique of establishing a breeding population of Canada
geese.

�-42-

South Platte Valley-Jumbo Reservoir Annex
A total of 346 Canada goose goslings were released from 1968 through 1970
on the K-4 Ranch along the South Platte River near Masters, Colorado. At
least five nests were noted in the K-4 transplant area in the spring of 1971.
Goose transplant efforts in the South Platte Valley were moved to Jumbo Reservoir Annex, a Division controlled property in extreme northeast Colorado,
with the release of 102 goslings in June and July of 1971. Sixty two birds
were released on June 22 and an additional forty on July 13. An area of
approximately 250 square miles was closed to goose hunting around the transplant site during the 1971-72 season. Generally, the geese remained in the
area throughout the winter. They were joined by as many as 5,000 migrant
Canada geese of the short grass prairie population during late fall and winter.
Only one of the birds released was reported taken during the 1971~72 hunting
season. That bird was harvested near the south boundary of the closed area,
directly south of the release site.

San Luis Valley-San Luis Lakes
A total of 228 Canada goose goslings were released on San Luis Lake in the
San Luis Valley in June and July of 1969 and 1970. In the spring of 1971
variable numbers of geese were observed at San Luis Lake. Some pairs were
noted to be frequenting the floating type artificial goose nesting structures
but no broods were observed in the area. An additional 123 goslings were
released on San Luis Lake in June and July of 1971, the last scheduled release
for that area.
In June of 1971, two of the birds released on San Luis Lakes in 1969 were
captured in a goose drive trapping operation on Wheatland Reservoir north of
Bosler in Albany County, Wyoming. Five of the 1969 transplants were captured
on that same reservoir in June, 1970, along with seven goslings released in
1967 and 1968 on the Alamosa Refuge in the San Luis Valley (Szymczak 1971).
Szymczak (1971) postulated that either geese from the San Luis Valley were
using Wheatland as a molting area, or the Valley birds captured at Wheatland
had drifted into Great Basin Canada goose population area and subsequently
became members of those populations. Recoveries during the 1971-72 period
provided only a limited amount of evidence as to whether Wheatland Reservoir
is important to the San Luis Valley birds. Only one of the San Luis Valley
transplants recaptured at Wheatland has been recovered. That bird, a 1968
Alamosa Refuge transplant, was recovered in Sevier County in central Utah in
a Great Basin population area.
The number of reported recoveries of birds released on San Luis Lakes in 1970
and 1971 has been somewhat limi~ed (Table 1). However, recoveries along with
recapture data continue to indicate that a portion of the San Luis Lake
birds are using the Bitter Lakes National Wildlife Refuge in Chaves County,
New Mexico in the winter. During the 1971-72 Winter, two birds from San Luis
Lakes were harvested in Chaves County, one released in 1970 and one in 1971,
and six birds were recaptured on the Bitter Lake Refuge, three from 1969 and
three from 1971. The bird harvested in Chaves County from the 1970 group had
been recaptured at Bitter Lakes the previous year. Three birds were reported

�-43taken in southeast Colorado during the 1971-72 season, one each in the
counties of Bent, Prowers, and Otero (Table 1). The bird taken in Otero County
was released in the San Luis Valley in 1971, indicating an eastern movement
out of the Valley. The bird taken in Prowers County had been recaptured in
January, 1971 at the Bitter Lake Refuge.

Table 1. Hunting season recovery locations of Canada geese released at San
Luis Lakes, Alamosa County, San Luis Valley, Colorado.
Year Released
Number Released
Area Recovered

1969
160
No. Recovered
Direct Indirect

1970

1971

68

123

No. Recovered
Direct Indirect

No. Recovered
Direct Indirect

United States
Colorado
San Luis Valley

2

3

2

o

3

N. A.

Chaffee County

o

1

o

o

o

N. A.

Bent County

o

o

o

1

o

N. A.

Prowers County

o

o

o

1

o

N. A.

Otero County

o

o

o

o

1

N. A.

Chaves County

9

o

1

1

1

N. A.

San Miguel County

1

o

o

o

o

N. A.

o

1

o

o

o

N. A.

5

1

o

o

o

N. A.

1

o

o

o

o

N. A.

o

1

o

o

o

N. A.

18

7

3

3

5

New Mexico

Arizona
Cochise County
Mexico
Sonora
Southwest
Chihuahua
Unknown
Canada
Saskatchewan
Southwest
Total

�-44-

The entire San Luis Valley, with the exception of the San Luis Lakes-Head
Lake transplant area was open to goose hunting during the 1971-72 season.
The western portion of the Valley was open to hunting by permit only, with
a maximum of 350 geese the allowable harvest.

North Park-Lake John Annex
During the summers of 1969 and 1970 a total of 440 Canada goose goslings
were released on Lake John Annex, eight miles west and 2 miles north of
Walden. Band recoveries indicate that these birds migrated to the southwest
and spent the winter along the lower Colorado River bordering Arizona and
California (Szymczak 1971).
Spring Returns
Although an additional 246 birds were added to the transplant population in
the summer of 1970, only a relatively small number of returning birds were
observed in North Park in the early spring of 1971. Ten birds were first
observed in North Park on April 10, 1971, along the Illinois River just
west of Walden. About the same time, 27 birds were found at Lake John Annex.
On April 19, 25 birds were counted on Lake John Annex. Among the 25 birds
were: two single geese, which may have been defending territories; 2 specific
pairs, of which three birds wore neckbands identifying them as birds released
in 1969; one distinct group of three birds; and the remaining 16 birds, one
of which wore a neckband.
The largest group of birds in North Park during the spring period was approximately 50 observed on the Annex in late May. No coordinated survey of
waters in North Park was conducted.
It was reported that one brood of three goslings was produced on Lake John
Annex.

Molting
On June 22, 1970, 16 birds released on Lake John Annex in the summer of 1969
were recaptured at Wheatland Reservoir north of Laramie, Wyoming. The significance of Wheatland Reservoir as a molting area for various Great Basin
Canada goose populations was discussed in an earlier section. Only one of
the 16 birds was subsequently recovered during the 1970-71 hunting season.
That bird was taken along the lower Colorado River in Arizona (Szymczak 1971).
On June 21, 1971, four birds from the 1969 release and four from the 1970
release were recaptured at Wheatland. All four of the 1969 birds were
present at Wheatland the year before. It is possible that all of the North
Park transplants at Wheatland in 1971 were not identified as such, because
not all of the banded birds recaptured were recorded (G. Wrakestraw, personal
communication). From all indications, it seems that Wheatland may become a
major molting area for North Park geese.

�-45-

New Releases and Summer Observations
On July 22, 1971, 241 goslings were released on Lake John Annex. Throughout
the remainder of the summer and early fall the birds frequented most of the
large water bodies, as had the birds released the previous year (Szymczak
1971). The largest daily number of geese observed between August 14 and
September 9 was a total of 189 sighted on Walden Reservoir, Lake John Annex,
MacFarlane Reservoir and Hebron Sloughs on August 27.
At least 20 geese were still present in North Park on October 20, the last
recorded observation date during the fall period.
Only one goose with a neckband (1969 release) was observed during the 1971
summer and fall period. That bird, a member of a pair, was observed on the
Boettcher Lakes on September 5, 1971.
Band Recoveries
Only five of the birds released in 1969 and six of the birds released in 1970
were reported recovered during the 1971-72 hunting season. These birds are
recorded in the indirect recovery column for their respective years in Table
2. Seven of the 11 total recoveries came from the lower California counties
of Riverside and Imperial. Fifteen of the 45 first year recoveries of birds
released in 1971 also came from Riverside and Imperial counties. Of the 25
recoveries from the three southern California counties (Table 2) during the
three year period 22 were reported taken during the 1971-72 season (15 direct
and 7 indirect recoveries) indicating the importance of California as a
recovery area during that season.
Yuma County, Arizona was also an important recovery area in 1971-72, but only
for birds released in 1971. In fact, all 46 of the recoveries from Yuma
County, Arizona during the three year period have been direct (Table 2),
indicating the birds have not returned to that area in subsequent winters.
The birds released in 1971 exhibited the same general cohesiveness during
the fall and winter period as birds released in 1970. Only two, one each
from Graham County, Arizona, and Santa Fe County, New Mexico~ of the 45 birds
recovered were taken outside the expected migration route and winter area.
The birds released in 1971 and taken in Yuma County were harvested earlier
(Table 3) than were their counterparts released in 1970 and harvested during
the 1970-71 season (Szymczak 1971). This was probably the result of an
earlier opening date in 1971 (November 13) than in 1970 (November 26).
The recoveries recorded by time period in Table 3 seem to indicate a movement of birds from the Yuma County, Arizona area into the Imperial Valley
of California. The goose season in the Imperial Valley did not open until
November 27. Hunting pressure applied in mid-November in Arizona may have
moved the birds across the border into California. Only one bird was taken
in Yuma County Arizona after December 10, even though the goose season did
not end until January 9.

�-46-

Table 2. Hunting season recovery locations for all years
of Canada geese
released at Lake John Annex, Jackson County,
Colorado.
Year Released
No. Released

1969
193

Area Recovered

No. Recovered
Direct Indirect

1970
246

·1971
241

All Years
680

No. Recovered ...No ~ .Recovered
Direct Indirect Direct Indirect

No. Recovered
Direct Indirect

United States
North Dakota
Emmons Co.

0

1

0

0

0

N.A.

0

1

Custer Co.

0

1

0

0

0

N.A.

0

1

Rosebud Co.

0

1

0

0

0

N.A.

0

1

Montana

W:l!:0mins
Park Co.
Albany Co.

0

1

0

0

0

1

0

0

1

0

N.A.
N.A.

0

0

0

1

Carbon Co.

2

0

5

0

0

N.A.

7

0

1

0

0

0

2

3

0

0

1

0

0

0

0

1

1

0

2

0

Colorado
Jackson Co.!/
Larimer Co.
Baca Co.
Moffat Co.

1

0

0

0

0

2

0

0

0

0

N.A.
N.A.
N.A.
N.A.

Mesa Co.

0

1

0

0

0

N.A.

0

1

1

0

0

0

0

0

0

0

1

0

N.A.
N.A.

1

0

0

1

0

0

0

0

1

N.A.

1

0

1

0

0

0

0

N.A.

1

0

Utah
Emery Co.
Cane Co.
New Mexico
Santa Fe Co.
Texas
E1 Paso Co.

----------------------------------------------------------------------------------

�-47-

Table 2. Hunting season recovery locations ·for all years of Canada geese
released at Lake John Annex, Jackson County, Colorado (continued).
Year Released
No. Released

1969
193

Area Recovered

No. Recovered
Direct Indirect

''1911
241

1970
246
No. Recovered
Direct Indirect

All Years
680

.No. Recovered
Direct Indirect

No. Recovered
Direct Indirect

Arizona
Yavapai Co.

1

0

0

0

0

N.A.

1

0

Gila Co.

1

0

0

0

0

N.A.

1

0

Graham Co.

0

0

0

0

1

N.A.

1

0

Mohave Co.

0

1

1

0

0

N.A.

1

1

Yuma Co.

6

0

14

0

26

N.A.

46

0

San Bernardino
Co.

0

0

1

0

0

N.A.

1

0

Riverside Co.

0

2

0

4

6

N.A.

6

6

Imperial Co.

0

1

2

0

9

N.A.

11

1

Northwest

1

0

0

0

0

N.A.

1

0

Unknown

1

0

0

0

0

N.A.

1

0

0

0

2

0

0

N.A.

2

0

2

0

0

0

0

N.A.

2

0

2

0

0

0

0

N.A.

2

0

22

10

25

6

45

92

16

California

Mexico
Baja

Chihuahua
Northwest
Coahuila
West-central
Sinaloa
Northwest

Totals

1/
- Area closed to goose hunting.

�-48-

Table 3. 1971-72 hunting season recoveries of North Park birds released in
1971 plotted according to time period and area of harvest.

Area Recovered

November
1-10 11-20 21-30

Period of Recovery
December
1-10 11-20 21-31

January
1-10 11-20

Colorado
Jackson County II

2

New Mexico
Santa Fe County

1

Arizona
Graham Coun ty
Yuma County

1
6

14

5

1

Riverside County

1

2

1

Imperial County

1

3

5

California

!I

1

1

Area closed to hunting.

Mortality
Forty five or 18.7 percent of the birds released in 1971 were reported
recovered. Comparative first year recovery rates for 1969 and 1970 transplants were 11.4 and 10.2 percent, respectively. The earlier season in
Arizona may very well be responsible for the increased harvest of the birds
released in 1971 as compared to the 1970 group. As mentioned before, both
groups showed similar migration route and wintering area tendencies.
Totten Reservoir
The first release of Canada goose goslings was made at Totten Reservoir east
of Cortez in Montezuma County, in June, 1970. These birds, 36 in number,
stayed in the area throughout the winter, using both Totten and nearby
Narraguinnep Reservoir (Szymczak 1971).

�-49-

On July 2, 1971, 38 goslings were added to the Totten Reservoir flock.
These birds also used Narraguinnep and Totten through the summer and fall,
along with the previous year's plants. However, sometime in mid- to late
December the entire flock left the area. Approximately one month later, 23
birds returned to Totten.
According to band recoveries the majority of the birds apparently moved
almost straight south and terminated migration on the Gila River in Graham
County near Stafford in southeast Arizona. Two of the birds released in 1970
and six of those released in 1971 were reported harvested in the Stafford area.
In addition, the earliest bird reported recover.d (12-18-71) was taken upriver
on the Gila from Stafford near the Arizona-New Mexico border. To complete
the recovery picture, one bird was found dead near the release site.

LITERATURE CITED

Szymczak, M. R. 1971. Experimental studies on improving status of Canada
goose populations. Colo. Div. Game, Fish and Parks, Game Res. Rep.,
Fed. Aid Project W-88-R. October. 115-127 pp.

f~._.-~~~_l~
__

Prepared by --.77Z~~---&lt;-~-127...p...f~gCo....:.::-"g?
•...•-"-,,,.~
Michael R. SZymCz~UV~
Assistant Wildlife Researcher

��-51-

October,

1972

JOB PROGRESS REPORT

State of

COLORADO
---------------~~--~--------

Project No.

W-88-R-17

Work Plan No.

2

Migratory

Bird Investigations

Job No.

Job Title

Arkansas

Valley

Canada Goose Flock Management

Period Covered:

November

1, 1971 to March 31, 1972

5
Studies

Personnel:
Ann Leckler, D. Clippinger, R. Forbes, A. Heins, T. Henry,
R. Kitzmiller, W. Ladd, C. Leonard, T. Lytle, H. Moorhead, P. Olson,
Potts, R. Rosette, R. Velarde, B. Widhalm and M. Szymczak.

n.

ABSTRACT
Forty thousand geese were present in southeast Colorado by November 3, 1972.
By November 9, 1972, 80,200 geese were in the area and from that time through
early January, aerial surveys indicated that numbers varied from 65,050 to
87,750.
The total number of geese in the Two Buttes-Turk's Pond area built
throughout the season reaching a high of 59,000 by December 21, 1972. The
mid-winter inventory for the entire short grass prairie population totaled
134,500 birds. According to the tail fans collected in southeast Colorado
during the season, approximately 44 percent of the geese bagged were
immatures.
An estimated 29,114 geese were bagged by 9,468 hunters in southeast Colorado during the 1971-72 season.
Bands were placed on 1,032 geese
during post-season trapping operations at Two Buttes Reservoir and Turk's
Pond. Approximately 33 percent of the birds banded were immatures.
For
the first time since the Lamar-Eads check station was established in 1968,
the average weight of geese harvested at Lamar-Eads did not exceed the mean
weight of geese harvest at Two Buttes. Weights of birds harvested plotted
by time period, indicated the birds did not lose weight as the season
progressed.
The record harvest of Canada geese in southeast Colorado
during the 1970-71 season was reflected in the distribution of band recoveries during that season.
The annual mortality rate of all age classes
combined, as reflected in band recoveries, is 26 percent when calculated by
the composite dynamic method, and about 27 percent when calculated by the
relative recovery rate method.

��-53-

ARKANSAS VALLEY CANADA GOOSE FLOCK
MANAGEMENT STUDIES
Michael R. Szymczak

After two years of intensive evaluation of half-day goose hunting in southeast Colorado, the Arkansas Valley Canada Goose Flock Management Study geared
down to the primary function of monitoring the annual status of the short
grass prairie popUlation. The collectiort of information concerning hunting
pressure, wounding loss, and hUnter success at the Two Buttes and LamarEads Management area was discontinued under this job. The observation of
goose feeding flight patterns was also discontinued. A greater emphasis
was placed on obtaining a better distribution of tail fans collected from
harvested birds across Colorado's wintering range. Banding operations
continued as a major part of the study.
P. S. OBJECTIVE
To investigate the status of Canada geese wintering in southeast Colorado.
SEGMENT OBJECTIVES
1.

Estimate the size of the wintering flock.

2.

Estimate the age composition of the flock.

3.

Estimate the hunting pressure on, and harvest of the flock under
varying types of harvest regulations.

4.

Investigate the annual mortality and the migration and/or harvest pattern
of the flock under varying types of harvest regulations.
METHODS AND MATERIALS

Coordinated inventories were conducted throughout the short grass prairie
populations wintering range in Nebraska, Colorado, Oklahoma, Texas, and New
Mexico on December 13 and again in early January. In addition, periodic
counts were made in Colorado throughout the hunting season. Although these
periodic counts are not included in the procedures, result will be reported
as a part of this job.
Tail fans from geese bagged were collected at the Two Buttes and Lamar-Eads
Management Area check stations. In addition, some fans were collected near
Turk's Pond, and in the John Martin and Meredith Reservoir areas.
Hunting pressure and harvest information was obtained from the regular Colorado Small Game Harvest Survey.
A total of 1,032 Canada geese were banded during trapping operations at Turk's
Pond and Two Buttes Reservoir after the close of the hunting season in order
to obtain information on age ratios, mortality rates and migration patterns.

�-54-

RESULTS AND DISCUSSION
Size and Distribution of Wintering Population
A total of seven aerial surveys were conducted in southeast Colorado during
the fall and winter of 1971-72 (Table 1). These surveys indicated Colorado
wintered approximately the same number of birds in 1971-72 as during 1970-71.
In comparison with the 1970-71 season, there was a definite decline in the
number of birds utilizing Meredith Reservoir. Blue Lake held a significant
number of geese throughout November and late December, whereas in 1970-71,
essentially no geese used Blue Lake (Szymczak 1971). For the second consecutive year, John Martin Reservoir held comparatively large numbers of geese
throughout the season. Also, for the second year in succession, the number
of birds in the northern areas (Meredith, Blue, Eads group), excluding those
on John Martin, declined from mid-December through January. This decline
seemed to begin about the time the change over from half-day morning-only
hunting to full-day hunting occurred.
The Two Buttes-Turk's population continued to build throughout the season,
reaching a peak of 59,000 in late December. Approximately 75 percent of the
record number of geese recorded for the regular January lnventory in southeast Colorado (Table 2) were located in the Two Buttes-Turk's Pond area.
Surveys conducted on December 13 and January 4 were coordinated with counts
in other states throughout the short grass prairie popUlations wintering
range in order to determine the total number of birds in the population. In
December, a total of 148,570 birds were recorded. The January figure of
134,500 was the lowest total recorded since 1969 (Table 3). The degree to
which short grass prairie birds are utilizing other wintering areas outside
their historical range is not known. However, there is definite evidence
that substantial numbers of birds are mixing with the Hi-Line population
wintering in north-central Colorado (Szymczak 1971). Therefore, the results
presented in Table 3 include only birds utilizing the traditional range.
Age Composition
Goose tail fans were collected from harvested birds throughout the 1971-72
season at a number of locations in southeast Colorado. Tail fans collected
at Turk's Pond and John Martin Reservoir supplemented samples obtained
from the normal collection points of the Two Buttes and Lamar-Eads Management Areas and the Meredith Lake area.
Collection from all locations indicated a much larger percent of adults in
the harvest in 1971-72 than in the 1970-71 season (Table 4). The change in
age distribution of the harvest does not necessarily indicate a decline 1n
production in 1971, but may in fact, be a sign of a large number of subadults in the population resulting from excellent production in 1970
(Szymczak 1971). This type of a distribution change has been noted before,
and is generally expected.

�Table

l.

Results of Canada goose aerial surveys, southeast Colorado, 1971-72.

Nov. 3

Nov. 9

Nov. 22

Date
Dec. 6

Dec. 13

Dec. 21

Jan. 4

Meredith Reservoir

3,400

13,000

1,300

5,200

3,000

250

1,500

Horsecreek Reservoir

600

3,800

600

0

0

0

Blue Lake

100

6,000

0

7,500

7,000

300

°
°

Sweetwater

0

0

1,400

3,000

0

0

0

Flamingo

0

4,100

8,000

8,000

0

Nee Noshe

0

15,000

12,000

6,000

°

0
12,000

0

Area

Eads Group:

5,000

Upper Queens

10,600

2,700

50

1,500

Lower Queens

3,400

0

0

0

Two Buttes Reservoir

11,000

14,000

16,400

6,700

18,500

25,000

6,000

Turk's Pond

8,300

16,000

17,300

32,000

25,000

34,000

51,000

John Martin Reservoir

2,000

5,600

8,000

15,000

22,500

2,200

9,500

600

--

--

--

--

300

80,200

65,050

85,000

87,750

Bonny Reservoir

Total

40,000

60O

85,500

4,000

14,000

°

0

8,100

°

76,400

I

VI
VI
I

�-56-

Table 2 • January Inventory of Canada geese, Arkansas Valley, Colorado,
1948-1972.
Year

Goose Count

Year

Goose Count

Year

Goose Count

1948

4,798

1957

24,617

1965

37,693

1949

12,286

1958

35,894

1966

38,635 II

1950

13,170

1959

44,660

1967

29,835

1951

19,320

1960

37,394

1968

42,682

1952

30,463

1961

31,360

1969

29,201

1953

20,236

1962

40,250 II

1970

63,444

1954

20,280

1963

35,889

1971

62,720

1955

25,110

1964

33,750

1972

76,400

1956

24,212

!/Inventory of February 7, 1962 substituted for January, 1962 inventory.
IIInventory of February 15, 1966 substituted for January, 1966 inventory.

Table 3. Post-hunting season status of Short Grass Prairie Canada goose
population, 1960-72, data generally from regular mid-winter inventories.
Year

Number of Birds

Year

Number of Birds

1960

77,709

1967

111,452

1961

103,355

1968

127,903

1962

80,133

1969

112,399

1963

93,940

1970

147,414

1964

81,221

1971

152,734

1965

103,435

1972

134,500

1966

110,485 :!/

:!/ Inventory of Feburary 15, 1966 substituted for unsatisfactory January
inventory.

�Table 4.

Age composition of the goose harvest in selected areas, southeast Colorado.

Year

Two Buttes (Firing Line)
Immature
Adult
No.
Percent
Percent
No.

1951-52

356

46.8

404

53.2

Lamar-Eads (Decol)
Immature
Adult
Percent
No.
Percent
No.

Meredith Area
Immature
Adult
Percent
No.
Percent
No.

No Data

1957-58
1958-59

929

47.9

1,010

52.1

1959-60

377

36.5

655

63.5

1960-61

612

45.5

732

54.5

1961-62

527

55.7

419

44.3

1962-63

204

48.9

213

51.1

1963-64

377

52.1

346

47.9

I
\J1

1964-65

442

63.1

259

36.9

I

1965-66

571

62.7

340

37.3

1966-67

217

45.3

262

54.7

1967-68

245

59.9

164

40.1

1968-69

136

61. 8

84

38.2

79

51.3

75

48.7

48

66.7

24

33.3

1969-70

110

55.3

89

44.7

143

35.9

255

64.1

18

40.0

27

60.0

1970-71

630

40.6

922

59.4

77

27.4

204

72.6

42

45.7

50

54.3

1971-72 !/

804

63.5

463

36.5

128

33.7

252

66.3

63

51.6

59

48.4

.....•

!/ Data for Two Buttes also includes some birds bagged in the Turk's Pond area.
some birds bagged in the John Martin Reservoir area.

Data for Meredith area includes

�-58-

The overall ratio of adults to immatures in the harvest in southeast Colorado
was 1:0.78. This ratio in comparison with the age ratios of geese harvested
on the Canadian prairies reflects the general accepted trend of a decline in
immatures in the harvest as the season progressed. Age ratios in the prairie
harvest were 1:1.18 in western Saskatchewan and 1:1.35 in eastern Alberta
(A. Dzubin, personal communication).
The age composition of Canada geese trapped during post-season banding operations varied considerably by individual catch (Table 5). The percent of
immatures in individual catches ranged from 15.3 percent to 55.2 percent at
Two Buttes and averaged 29.2 percent or 1:0.41 adult/immature. Birds
captured in the one catch at Turk's Pond were composed of 37.4 percent immatures
1:0.59 adult/immature.
A summary of all age ratios obtained in southeast Colorado is presented in
Table 6.

Table 5. Sex and age composition of Canada geese trapped post-season in
southeast Colorado, 1972.

Location and Date
Two Buttes

Total

Male
Immature

Adult

Female
Immature

Total

Percent
Immature

1-20-72

80

15

69

12

176

15.3

1-21-72

50

5

29

12

96

17.7

1-23-72

47

17

41

31

136

35.3

1-25-72

14

4

8

4

30

26.7

1-29-72

11

9

3

1

25 ~/ 41. 7

1-30-72

14

11

13

4

42

35.7

2-1-72

15

33

24

15

87

55.2

231

94

187

79

592

29.2

166

85

119

85

455

37.4

397

179

306

164

1,047 ~/

32.8

Sub-total
Turk's Pond

Adult

1-28-72

~/ Includes one female of unknown age.

�-59-

Table 6. Age composition of southeast Colorado Canada geese, 1971-72, as
estimated by check station, field checks, and trapping results.
No. of
Adults

Percent
Adults

No. of
Young

Percent
Young

Total
Birds

Lamar-Eads

128

33.7

252

66.3

380

Two Buttes

474

68.4

219

31.6

693

Turkts Pond

330

57.5

244

42.5

574

John Martin-BlueMeredith

63

51.6

59

48.4

122

Total

995

56.2

774

43.8

1,769

Check Station-Field
Check Sample

Adult/Immature

1:0.78

Trapped Sample
Two Buttes

418

70.7

173

29.3

591

Turk's Pond

285

62.6

170

37.4

455

Total

703

67.2

343

32.8

1,046

Adult/Immature

1:0.49

Hunting Pressure and Harvest
Goose hunters enjoyed generally excellent success in southeast Colorado during
the 1971-72 hunting season. An estimated 9,468 hunters bagged 29,114 birds
during the 1971-72 season (Table 7) (Riffel and Tulley, personal cormnunication).
The total harvest in the area was nearly identical to the record harvest
taken during the 1970-71 season. An estimated additional 5,299 birds were
reported crippled, resulting in a total flock loss of 34,413 geese.
Baca County continues to be the major harvest area with over 45 percent of
the birds taken in that area (Table 8). The distribution of total harvest
by county during the 1971-72 season closely paralleled that observed during
the 1970-71 season.

�-60-

Table 7.

Goose hunting season statistics, 1954-71.

Southeast Colorado
Estimated
Average
Estimated
Goose Hunters
Season Bag
Kill

Year

Dates of Season

Stamp
Sales

1954

11/1 - 12/30

32,450

7,071

1.04

7,372

1955

11/1 - 12/30

39,107

9,054

1.54

13,904

1956

11/9 - 1/7

36,303

9,833

1.05

10,276

1957

11/2 - 11/31

41,794

9,113

1. 39

12,656

1958

11/17 - 1/15

41,897

10,082

1.51

15,205

1959

10/26 - 1/8

31,431

8,888

1.61

14,309

1960

10/26 - 1/8

30,592

9,838

1. 39

13,629

1961

11/10 - 1/8

24,854

7,577

1.68

11,724

1962

10/31 - 1/13

17,701

6,021

1.58

9,495

1963

11/2 -

1/15

22,940

6,668

2.17

14,444

1964

11/2 -

1/15

25,282

8,016

2.30

18,474

1965

11/2 - 1/15

20,537

6,313

1.52

9,613

1966

11/19 - 1/15

29,377

9,357

2.59

24,269

1967

11/18 - 1/14

31,064

6,975

2.23

15,558

1968

11/16 - 1/15

31,218

6,668

1.66

11,046

1969 };./

11/22 - 1/15

34,281

5,327

1.99

10,597

1970 'f:../

10/31 - 1/17

37,972

7,875

3.85

30,329

1971 1./

11/1 - 1/16

43,743

9,468

3.07

29,114

1/ Morning hunting only throughout the season.
'f:../ M:&gt;rning hunting only through December 11, full day hunting for the
remainder of the season.

1/ Morning hunting only through December 11, full day hunting for the
remainder of the season.

�-61-

Table 8. Comparison of the 1971-72 southeast Colorado goose harvest, by
county, with the l7-year average, 1954-70, based on the results of the
random small game survey.

Number and Percent of Geese Bagged
1971-72
l7-Yr. Average
No.
Percent
No.
Percent

County

Lakes and Reservoirs

Baca

Two Buttes and Turk's

13,264

45.6

5,552

37.8

Kiowa

Eads and Blue

6,039

20.7

3,328

22.7

Prowers

Two Buttes and Eads

2,686

9.2

2,234

15.2

Bent

John Martin, Blue and
Horsecreek

2,620

9.0

1,932

13.2

Meredith

2,305

7.9

852

5.8

880

3.0

120

0.8

734

2.5

371

2.5

Pueblo

534

1.8

193

1.3

Huerfano

52

0.2

109

0.7

Crowley
Las Animas
Otero

Total

Horsecreek, Cheraw, Dye
and Holbrook

29,114

14,691

Banding Investigations
Trapping efforts post-season in southeast Colorado resulted in 1,032 Canada
geese being banded (Table 9). All birds banded were captured either at Two
Buttes Reservoir or at Turk's Pond. Very few geese were using reservoirs
in the John Martin or Ead's Lakes areas so no attempt was made to capture
geese in those areas.
Forty-three percent of the birds captured were females. This compares with
53 percent females captured during the previous two years of trapping.
Weights
The mean weights of all geese harvested on the two state controlled management areas and trapped post-season are presented in Table 10. Some weights
from geese bagged at Turk's Pond are also included.

�-62-

Table 9. Sex and age composition of Canada geese banded post-season in
southeast Colorado, 1972.

Location and Date

Two Buttes

Turk's Pond

Adult

Males
Immature

Females
Adult
Immature

Total

1/20/72

80

15

68

12

175

1/21/72

50

5

29

12

96

1/23/72

47

17

41

31

136

1/25/72

14

4

8

4

30

1/29/72

11

9

3

1

25 1:/

1/30/72

14

11

13

4

42

2/1/72

15

32

24

15

86

1/28/72

161

83

116

82

442

392

176

302

161

1,032 1:/

Total

1:/ Includes one female of unknown age.
Table 10. Comparison of weights, in pounds, of geese from trap and check
station samples, southeast Colorado 1971-72.
Adult
Ave. Wt.

Range

Check
Station

5.49(126)1

4.00-7.75

Trapping

5.49(695)

5.50(126)

Source

Immature
Ave. Wt.

Range

Total
Ave. Wt.

Range

5.12 ( 79)

4.00-7.00

5.35

4.00-7.75

3.25-9.88

5.37(343)

3.38-9.38

5.45

3.25-9.88

3.38-9.20

4.78(254)

3.13-7.00

5.02

3.13-9.20

Two ButtesTurk's Pond

Lamar-Eads
Check
Station
1

Sample size in parentheses.

�-63-

Geese trapped post season in the Two Buttes-Turk's area weighed, on the
average, slightly more than those harvested in that same area. Only in
1969-70 have trapped birds weighed more than harvested birds. However, in
1969-70 the trapped birds weighed about .5 pounds more than harvested birds.
The difference in 1971-72 is only .1 of a pound and is a result only of
differences in the immature category.
For the first time since the Lamar-Eads check station was established, in
1968-69, the average weight of geese harvested at Lamar-Eads did not exceed
the mean weight of geese harvest at Two Buttes (Table 10).
Weights of birds harvested plotted by time period (Table 11) indicated no
distinct weight loss as the season progressed as had occurred during 1969-70
and 1970-71. The validity of the analysis is hampered by generally small
samples during the latter part or full day portion of the season. However,
the fact that birds trapped post season in the Two Buttes-Turk's area
weighed, on the average, more than birds bagged there (Table 10), seems to
indicate that geese may not have lost weight during the 1971-72 season as
they normally had during past seasons.
The groups of geese captured on January 29 at Two Buttes were unusually
heavy and those captured on January 30 were slightly heavier than the average
(Table 12). The average weights by age and sex groups for other catches
were considered "normal".
Distribution of Harvest
The record harvest of Canada geese in southeast Colorado during the 1970-71
season (Szymczak 1971) was reflected in the distribution of band recoveries
during that season (Table 13). Approximately 38 percent of the bands reported
recovered were taken in Colorado. Of the major harvest areas, only Saskatchewan failed to record a significant decline in the percent of bands recovered
in response to the heavy Colorado harvest.
For the second consecutive year, and for the third year out of the past four,
the percent of bands recovered below 530 in the Provinces was greater in
Saskatchewan than in Alberta.
Hunting Mortality Estimates
Mortality estimates calculated by the composite dynamic method for juveniles,
adults and both age classes combined continue to be approximately 26 percent
(Tables 14, 15, and 16). The first year mortality rate for juveniles declined
from .285 calculated for recoveries through the 1968-69 season (Szymczak 1971)
to .283 calculated for recoveries through the 1969-70season(Tab1e
14). The first
year recovery rate for adults took a greater drop from .270 to .265.
Mortality rates calculated by the relative recovery rate method for juveniles,
adults, and all age classes (Tables 17, 18, and 19) are all slightly higher
than the rates calculated by the composite dynamic method.

�-64-

Table 1l. Average weights of
geese bagged by time interval on two major
harvest areas in southeastern Colorado during the 1971-72 hunting season.
Lamar-Eads
Adults
Immatures

Time Interval

Two Buttes-Turk's Pond
Adults
Immatures

Nov. 1 - Nov. 7

5.49( 56)1

4.78(152)

5.56(

Nov. 8 - Nov. 14

5.66( 16)

4.97( 35)

5.80( 24)

5.09( 17)

Nov. 15 - Nov. 21

5.76( 20)

4.75(19)

5.43( 16)

5.07( 17)

Nov. 22 - Nov. 28

5.03( 13)

4.48( 27)

5.63( 22)

4.80( 14)

Nov. 29 - Dec. 6

6.08(

4)

4.33(

Dec. 7 - Dec. 10

4.75(

2)

Dec. 11 - Dec. 17

6.13(

2)

5.25(

2)

4.78(

Dec. 18 - Dec. 24

5.15(

4)

4.91(

5)

5.24 ( 25)

5.33( 10)

Dec. 25 - Dec. 31

4.57(

3)

4.97(

6)

5.48( 25)

5.09( 11)

Jan. 1 - Jan. 7

5.80(

2)

5.20(

1)

Jan. 8 - Jan. 16

5.76(

4)

5.06(

1)

5.25(

5.38(

Average

5.50(126)

1

8)

5.30(

5)

6)

4.78(254)

4)

2)

5.49(126)

6.50(

2)

5.25(

1)

2)

5.12( 79)

Sample size in parentheses.

Table 12. Weights of Canada geese captured with cannon-nets in southeast
Colorado, 1972.
Date

Location

Grouping

Number

Ave. Weight

Range

January 20

Two Buttes

AM

74

5.29

4.13-7.25

1M

15

5.13

4.38-6.63

AF

69

4.74

3.25-6.50

IF

12

4.67

3.50-6.75

Subtotal

170

5.01

3.25-7.25

----------------------------------------------------------------------------

�-65Table 12. Weights of Canada geese captured with cannon-nets in southeast
Colorado, 1972 (continued) •
Date

Location

Grouping

Number

Ave. Weight

Range

January 21

Two Buttes

AM

50

5.27

4.00-7.25

1M

5

6.10

5.25-8.00

AF

29

4.88

3.63-6.25

IF

12

4.54

3.63-5.25

Subtotal

96

5.11

3.63-8.00

AM

47

5.79

4.00-7.38

1M

17

5.37

4.25-6.63

AF

40

5.12

3.50-8.38

IF

31

4.74

3.50-6.00

Subtotal

135

5.30

3.50-8.38

AM

13

6.08

4.63-8.00

1M

4

5.76

4.88-6.38

AF

8

4.99

4.50-5.75

IF

4

5.19

4.25-5.88

Subtotal

29

5.61

4.25-8.00

AM

11

7.14

6.63-7.75

1M

9

7.35

6.13-9.38

AF

3

7.88

7.25-8.75

IF

1

7.13

Subtotal

24

7.31

6.13-9.38

AM

14

6.36

4.25-9.88

1M

11

6.23

4.25-9.13

AF

13

5.49

4.00-7.00

IF

4

6.16

4.75-6.88

Subtotal

42

6.04

4.00-9.88

January 23

January 25

January 29

January 30

Two Buttes

Two Buttes

Two Buttes

Two Buttes

------------------------------------------------------------------------------

�-66-

Table 12. Weights of Canada geese captured with cannon-nets in southeast
Colorado, 1972 (continued).
Date

Location

Grouping

Number

Ave. Weight

Range

February 1

Two Buttes

AM

15

5.79

4.25-7.25

1M

33

5.25

3.50-6.63

AF

24

5.16

3.38-6.63

IF

15

5.19

3.88-6.00

Subtotal

87

5.31

3.38-7.25

AM

224

5.63

4.00-9.88

1M

94

5.63

3.50-9.38

AF

186

5.01

3.25-8.75

IF

79

4.91

3.50-6.88

Total

583

5.34

3.25-9.88

AM

166

5.92

4.13-7.88

1M

85

5.72

3.50-7.00

AF

119

5.40

3.63-7.38

IF

85

5.14

3.38-6.25

Total

455

5.60

3.38-7.88

AM

390

5.75

4.00-9.88

1M

179

5.68

3.50-9.38

AF

305

5.16

3.25-8.75

IF

164

5.03

3.38-6.88

Total

1,038

5.45

3.25-9.88

Average Two Buttes
All Dates

January 28

Turk '9 Pond

Average Two Buttes-Turk's
Pond, All Dates

�Table 13.

Percentages of total band recoveries, Arkansas Valley post-season bandings, by area and year of recovery, all bandings.

Recovery Year
Area

Five Year Averages
1951-1955 1956-1960

Percentage
Total No. of Total
1970-71 Recoveries Recoveries

1961-62

1962-63

1963-64

1964-65

1965-66

1966-67

1967-68

1968-69

1969-70

--

--

1.0
12.4

0.5
6.1

13.4

6.6

267

9.2

21.9
25.7

15.7
24.7

816
465

28.1
16.0

4

0.1

17
2
16
7
175

0.6
0.1
0.6
0.2
6.0

848
12
15
27

29.2
0.4
0.5
0.9

Far North Above 530
N. W. Territories
Alberta
Saskatchewan
Total

--

--

----

----

----

----

--

9.8

7.1

7.9

6.3

15.5

28.6
18.3

35.3
10.5

34.9
9.3

33.3
19.4

28.4
17.4

--

--

--

--

---

----

15.2

4.7

10.4

7.5

2.1
7.1
0.7
9.9

26.8
8.5

27.9
10.9

19.4
16.4

14.2
24.6

23.4
17.0

0.7

1.5

0.7

--

Provinces Below 530
Alberta
Saskatchewan
B.C., Hanitoba,
Ontario

0.1

Central F1~
Hontana
North Dakota
Wyoming
South Dakota
Nebraska
Colorado
Southeast and Other
North-central
Total
Kansas
Oklahoma
New Hexico

0.8
0.1
0.7
0.1
5.4

--

0.4

0.6

0.7

0.7

2.0
1.4
8.8

0.7
0.7
9.7

0.5

0.6
6.5

5.4

3.1

5.5

6.7

---

---

--

---

30.1

35.4
0.7
1.4

32.4

--

--

--

--

--

---

25.5
0.4
0.9
1.5

30.9
0.3
0.3
0.7

31.5
0.8

30.9

21.7
1.0
0.5
1.0

0.6

--

0.7

1.4
0.7
0.7

1.9

1.9
4.8

0.5
0.5
4.5

9.0

4.3

27.6
1.5
29.1

29.1
2.8
31.9

19.0
1.0
20.0
1.0

34.3
3.0
37.9
1.0

1.5
0.7

1.4

1.0

2.0

---------------------------------------------------------------------------------------------------------------------_._------------------------------------

I

a-

-..J
I

�Table 13.

Percentages of total band recoveries, Arkansas Valley post-season bandings, by area and year of recovery, a11bandings

Recovery Year
Area

Five Year Averages
1951-1955 1956-1960

1961-62

1962-63

1963-64

1964-65

1965-66

1966-67

1967-68

1968-69

1969-70

5.4

4.0
2.0
1.0
6.0

7.3
2.4
0.6
10.3

2.7
2.0

7.5
0.7

7.5

4.7

8.2

2.2
9.7

5.0
0.7
1.4
7.1

6.7
1.0
1.0
8.7

1.5

1.2

0.7

1.4

0.7

0.7

(continued).

Percentage
Total No. of Total
1970-71 Recoveries Recoveries

Central F1ywaX (continued)
Texas
Panhandle
Waggoner Ranch
Gulf Coast
Total

2.3
2.0
0.8
5.1

2.8
2.2
0.2
5.2

5.4

4.6
0.8
0.8
6.2

Pacific Flyway

2.4

2.2

4.8

0.8

Mississipp~ Flyway

0.1

Mexico

0.1

Total Number
Of Recoveries

748

2.5
3.0
5.5

112
51
17
180

3.9
1.8
0.6
6.2

1.0

52

1.8

3

0.1

1

0.1

I
0\

00
I

677

129

129

201

164

147

134

134

141

105

198

2,907

�Table 14.

Composite dynamic analysis of band recoveries from Canada geese banded as juveniles, Arkansas Valley, Colorado, 1951-1969.

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69
1969-70

300
628
578
0
0
0
0
176
298
167
248
123
399
251
198
0
285
270
184
241

Recoveries bl Year Following Banding
11
10
12
13
8
9

14

15

16

17

0
2
3

1
1
1

1
0
1

0
0
0

0
1
2

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X

X
X

X

0
1

0

Ts----i9

1

2

3

4

5

6

7

31
58
52

25
31
25

11
33
25

11
20
24

4
16
20

5
8
11

4
7
11

7
6
9

1
4
2

1
4
4

0
0
4

0
5
1

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

21
25
16
15
10
40
16
15

16
16
10
5
12
24
11
7

6
16
3
4
10
9
11
8

4
8
14
7
0
14
13
1

4
7
8
8
1
6
5
0

1
3
0
7
1
2
2
3

4
0
0
3
1
2
8

0
1
1
2
1
4

3
1
0
1
1

1

0
1
1

X

43

40

31

13

14

6

7

5

3

2

0

3

1

4,346 4,105 3,921 3,651 3,366 3,366 3,168 2,917 2,518

2,395

2,147

1,980

1,682

1,506

1,506

1,506

1,506

1,506

5.8
209.5
21.1
26.3
822.9
.255

2.8

3.5

3.0

2.0

1.3

0.0

2.0

O.J..J

0.0

6.0
4.0
""-= 1,115.2
.262

2.7

2.7

0.7 /

0.0

X

X

X

X

18
16
9
18

10
9
9

5
8

10

360

210

149

126

2

0
2

1
0
0

0
0

I

co
'"'
I

Total
Recoveries 4,346
Banded Birds
Eligible
Recoveries per
1,000 Banded
Alive Going
into Period
Mortality Rate

82.8 \51.2

38.0

34.5

292.3,209.5 158.3 120.3
.283

79

23.5
85.8

12.8
62.3

12.6
49.5

10.6
36.9

5.2

0

928

£ =292.3

15.3

12.5

9.0

�Table 15.

Composite dynamic analysis of band recoveries from Canada geese banded as adults, Arkansas Valley, Colorado, 1951-69.

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69
1969-70

344
650
900
0
0
0
0
347
217
250
306
334
369
335
780
0
389
646
905
358

X

29
35
22
21

Total
Recoveries 7,130

558

351

--

Recoveries bl Year Following Banding
11
12
13
10
9

14

15

16

17

18

19

1
2
2

1
0
0

0
1
1

1
0
1

0
0
0

0
0
0

0
0

X
X

X
X

X
X

X
X

X

X

X
X

X
X

X
X

X
X

X

0
0
1

0

1

1

2

3

4

5

6

7

8

45
71
85

23
37
55

19
24
50

10
23
31

7
21
25

12
6
20

5
11
9

3
6
12

3
9
10

2
7
6

2
7
2

0
4
4

X
X

X
X

X

X

X
X

X
X

X

X
X

X

X

X
X

X

X
X

X
X

X
X

X
X

X
X

X

X

X

X
X

X
X

X
X

X
X

X
X

X
X

33
21
27
29
26
35
24
55

27
9
16
13
18
20
12
40

17
7
6
18
12
14
15
20

8
4
12
12
5
4
10
22

5
4
8
4
13
11
4
12

8
2
4
7
3
5
4
12

2
1
6
3
2
2
11

2
1
3
2
2
5

2
0
2
3
4

2
1
1
1

X

X

X

X

26
17
38

9
28

9

239

150

83

62

36

33

20

12

8

6

1

2

2

0

0

0

7,130 6,772 5,867 5,221 4,832 4,832 4,052 3,717 3,348

3,014

2,708

2,402

2,152

1,935

1,588

1,588

1,588

1,588

1,588

9.9

6.6
217.1
20.2
856.1
.254

4.4

3.3

1.3
0.5
295.4
3.1
2.6
5.9
£ = 1,151.5
.257

1.3

0.0

0.0/

0.0

1.3

0.0

0.0/

0.0

Banded Birds
Eligible
Recoveries per
1,000 Banded
Alive Going
into Period

295.4,217.1 165.312~.6

Mortality Rate

.265

78.3 ,51. 8

40.7

28.7

114

23.6
95.9

17.2
72.3

15.3
55.1

9.7
39.8

0

I

-J

0
I

30.1

2.8

1£ =

13.6

9.2

�Table 16.

Composite dynamic analysis of band recoveries from Canada geese banded in all age classes, Arkansas Valley, Colorado, 1951-1969.

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69
1969-70

644
1,278
1,478
0
0
0
0
523
515
417
554
457
768
586
978
0
674
922
1,089
599

Total Recoveries
11,482
Banded Birds
Eligible
Recoveries per
1,000 Banded
Alive Going
into Period
Mortality Rate

Recoveries b~ Year Following Banding
13
11
12
14
10

15

16

Ii

18

19

2
1
1

1
1
2

1
0
1

0
1
2

1
0
0

0
0

X
X
X
X

X
X
X

X
X

X

1

2

3

4

5

6

7

8

9

76
129
137

48
68
80

30
57
75

21
43
55

11
37
45

17
14
31

9
18
20

10
12
21

4
13
12

3
11
10

2
7
6

0
9
5

1
4
5

X
X
X
X

X
X
X
X

X
X
X
X

X

X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

54
46
43
44
36
75
40
70

43
25
26
18
30
44
23
47

23
23
9
22
22
23
26
28

9
5
4
14
4
7
6
15

6
1
6
6
3
4
19

2
2
4
4
3
9

5
1
2
4
5

3
3
1
3

0
1
2

0
1

1

26
19
5
18
23
23

9
11
16
12
14
17
9
12
X

126

92

67

46

34

18

15

11

4

4

2

3

1

0

11,482 10,883 9,794 8,872 8,198 8,198 7,220 6,634 5,866

5,409

4,855

4,438

3,923

3,400

3,400

3,400

3,400

3,400

1,922

3.7
212.1
20.4
14.1
829.9
.256

3.4

2.8

1.2

1.2

0.6

0.9

0.3 /

0.0

1.8

1.2

0.3 /

0.0

12
12

X

X

X

X

47
51
31
39

36
26
47

14
36

19

918

561

388

276

80.0 \51.5

39.6

31.1

292.1\212.1 160.6 121.0
.274

193

23.5
89.9

15.4
66.4

12.7
51.0

10.1
38.3

I
-.J

•...
I

7.8
28.2

6.3

~ = 292.1

10.4

7.0

4.2

3.0

£ = 1,122.0

.260

�Table 17. Relative recovery rate analysis of band recoveries from Canada geese banded as juveniles, Arkansas
Valley, Colorado, 1951-1969.
Winter
Banded

Number
Banded

1950-51

300

cJ

72

1951-52

628

196

1952-53

578

1957-58

Number Recoveries
1 - n
2 - n

Rates
2 - n

Survival
Rate

Mortality
Rate

--

.2400

.769

.231

138

.3121

.2197

.651

.349

195

CU

.3374

176

CU

39

--

.2216

.815

.185

1958.;.59

298

81

56

.2718

.1879

.592

.408

1959-60

167

53

37

.3174

.2216

1.018

0.000

1960-61

248

54

39

.2177

.1573

.523

.477

1961-62

123

37

27

.3008

.2195

.867

.133

1962-63

399

101

61

.2531

.1529

.582

.418

1963-64

251

66

50

.2629

.1992

1.160

0.000

1964-65

198

34

CU

.1717

1966-67

285

CU

25

--

.0877

.718

.282

1967-68

270

33

17

.1222

.0630

.644

.356

1968-69

184

18

9

.0978

.0489

.655

.345

1969-70

241

18

--

.0747

4.346

1 Cannot use.

Recove~
1 - n

2.7396
2.0193
Average Survival Rate
Average Mortality Rate

.737
.263

....,I
I\.)
I

�Table 18. Relative
recovery rate
Valley, Colorado, 1951-1969.

analysis

of band recoveries

from Canada geese banded as adults.

Arkansas

Winter
Banded

Number
Banded

1 - n

2 - n

1 - n

2 - n

Survival
Rate

Mortality
Rate

1950-51

344

cJ

89

--

.2587

.734

.266

1951-52

650

229

158

.3523

.2431

.699

.301

1952-53

900

313

CU

.3478

1957-58

347

CU

74

--

.2133

.926

.074

1958-59

217

50

29

.2304

.1336

.388

.612

1959-60

250

86

59

.3440

.2360

.785

.215

1060-61

306

92

63

.3007

.2059

.809

.191

Number Recoveries

Recove!1: Rates

1961-62

334

85

59

.2545

.1766

.679

.321

1962-63

369

96

61

.2602

.1653

.692

.308

1963-64

335

80

56

.2388

.1672

.810

.190

1964-65

780

161

CU

.2064

1966-67

389

CU

44

--

.1131

.914

.086

1967-68

646

80

45

.1238

.0697

1.051

0.00

1968-69

905

60

38

.0663

.0420

.716

.284

1969-70

358

21

--

.0587

7,130

2.7839

I

2.0245

Average Survival
Average Mortality

1 Cannot use.

I

""

W

Rate
Rate

.•727
.273

�Table 19. Relative recovery rate analysis of band recoveries from Canada geese banded in all age classes,
Arkansas Valley, Colorado, 1951-1969.
Winter
Banded

Number
Banded

1950-51

644

CUI

161

--

1951-52

1,278

425

296

.3326

1952-53

1,478

508

CU

.3437

1957-58

523

CU

113

1958-59

515

131

1959-60

417

1960-61

l:inmb~I ~!:'Q:ll~I1~a
1 - n
2 - n

Recove!1 Rates
1 - n
2 - n

Survival
Rate

~rta1ity
Rate

.2500

.752

.248

.2316

.674

.326

--

.2161

.849

.151

85

.2544

.1650

.495

.505

139

96

.3333

.2302

.874

.126

554

146

102

.2635

.1841

.690

.310

1961-62

457

122

86

.2670

.1882

.734

.266

1962-63

768

197

122

.2565

.1589

.638

.362

1963-64

586

146

106

.2491

.1809

.907

.093

1964-65

978

195

CU

.1994

1966-67

674

CU

69

--

.1024

.835

.165

1967-68

922

113

62

.1226

.0672

.939

.061

1968-69

1,089

78

47

.0716

.0432

.664

.336

1969-70

599

39

--

.0651

I

~
.J:'I

11,482

1

Cannot use.

2.7588
2.0178
Average survival rate
Average mortality rate

.731
.269

�-75-

LITERATURE CITED

Szymczak, M. R. 1971. Arkansas Valley Canada goose flock management studies.
Colo. Div. Game, Fish and Parks, Game Res. Rept., Fed. Aid Project
W-88-R. Oct. pp. 129-164.

��October,

-77-

JOB PROGRESS

REPORT

State of

COLORADO
----------~~~~----------

Project

No.

W-88-R-17

Work Plan No.
Job Title
Period

Migratory

2
Studies of Canada

Covered:

April

1972

Bird Investigations

Job No.
Goose Populations

6
in Colorado

Transplant

Areas

1, 1971 to March 31, 1972

Personnel:
D. Benson, C. Braun, C. Brown, E. Cochran, J. Corey, D. Coven,
G. Crawford, E. Downer, G. East, H. Funk, B. Goetze, J. Grieb, J. Hatfield
J. Hobbs, R. Hopper, T. Lynch, J. Monarch, S. Palm, J. Pogorelz, C. Roberts,
W. Russell, L. Searle and M. Szymczak.

ABSTRACT
During the 1971-72 hunting season, 208 geese banded outside Colorado were
reported recovered in north-central Colorado.
The recoveries indicated that
Big Horn County, in south-central Montana, is a probable additional breeding
area for Hi-Line Canada geese.
Phillips County, Montana and the EdmontonCamrose.and Dowling Lake areas of Alberta, as well as the Cypress Hills area
of Saskatchewan were well represented in the list of banding areas contributing geese to the north-central Colorado population.
During the 1971-72
hunting season Alberta was an important recovery area for birds banded in
January, 1971 in north-central Colorado, particularly for those birds judged
to be' small, and thus members of the short grass prairie population.
Alberta
has not been an important recovery area for previous bandings in north-central
Colorado.
During the 1971-72 season Colorado accounted for' 62.1 percent and
43.0 percent of the recoveries from post-season bandings in north-central
Colorado in January, 1968 and January, 1971 respectively.
Band recoveries
indicate that birds believed to be members of the Arctic nesting short grass
prairie population that are present in north-central Colorado's Hi-Line population area are not consistent in their migrational habits and may return to
their historical winter range in subsequent years.
Coordinated counts
throughout the Hi-Line population range totaled 61,054 on November 3, 1971;
62,552 on November 23, 1971; 61,236 on December 21, 1971; and 37,113 on
January 10, 1972. Between 85 and 90 percent of the population was located
in Colorado from late November through mid-January.
There was a substantial
percentage increase in the number of geese in the Brighton-Gree1ey-Ft.
Morgan
area during 1971-72 hunting season, as compared to the 1970-71 hunting season.
continued

�-78-

An estimated 12,920 hunters harvested 14,771 geese in north-central Colorado.
Approximately 52 percent of the birds harvested were reported taken in
Larimer County. A limited goose season in the San Luis Valley resulted in
an estimated 296 hunters bagging 177 geese. Analysis of banded birds recovered
in the San Luis Valley disclosed evidence indicating sub-adult birds from the
San Luis Valley breeding population are using Wheatland Reservoir in Albany
County, Wyoming for molting purposes. Aerial surveys indicated there was
approximately 1,200 to 1,500 Canada geese in the San Luis Valley during
the 1971-72 season, with the large majority of the birds located on the
Monte Vista National Wildlife Refuge. A limited goose hunting season in
west-central Colorado was initiated in 1971 with an estimated 112 hunters
harvesting 33 geese. Approximately 55 percent of the birds harvested were
taken within five miles ·of High1ine Reservoir, northwest of Grand Junction.
Aerial counts in the west-central permit area resulted in a total of 485
birds observed on November 24, 1971 and 989 birds counted on January 5, 1972.

�-79-

STUDIES OF CANADA GOOSE POPULATIONS IN COLORAOO TRANSPLANT AREAS
Michael R. Szymczak
P. S. OBJECTIVE
To investigate the status of re~ident and migrant Canada goose flocks and
their inter-relationships in areas in which populations have been established
through transplant programs in Colorado.

SEGMENT OBJECTIVES
La , To examine migration routes and/or harvest patt.erns and distribution of
Canada geese Wintering in north-central Colorado and the San Luis Valley.
lb.

To determine breeding areas of Canada geese wintering in north-central
Colorado.

2.

To estimate hunting pressure on, and hunter harvest of Canada geese in
north-central Colorado, the San Luis Valley and west-central Colorado.

3.

To make recommendations for establishing a limited Canada goose hunting
season in west-central Colorado and continuing the hunting season on the
north-central Colorado and San Luis Valley goose populations.

METHODS AND MATERIALS
Band recoveries listed on computer printouts provided periodically by the
Migratory Bird Populations Station were sorted and plotted as to banding and
recovery location, sex, and age, at time of banding, and estimated breeding
area for birds associated with the San Luis Valley and north-central Colorado.
Canada geese in north-central Colorado were counted on November 3, November 23,
and December 21, 1971; and January 10, 1972. In the San Luis Valley birds
were counted on October 31, November 23, and December 21, 1971; and January
6, 1972.
A mechanically random sample of hunters issued permits to hunt geese in the
north-central Colorado "Special Permit" goose hunting area were mailed
questionnaires inquiring about their hunting activity and success. All of
the hunters holding special permits to hunt geese in the San Luis Valley and
westcentral Colorado were mailed similar questionnaires. All data received
were tabulated and statistically analyzed.
Recommendations concerning regulations for the 1972-73 goose seasons in northcentral Colorado, the San Luis Valley, and west-central Colorado were formulated and submitted to management personnel.

�-80-

RESULTS AND DISCUSSION
North-Central Colorado-The Hi-Line Population Breeding Range
Foreign Recoveries
During the 1971-72 hunting season 208 foreign bands were reported recovered in
north-central Colorado. An additional 36 foreign-banded birds were recaptured
during post-season banding operations. Both types of recoveries are classified as to general area of banding if in a Province, or County if in the United
States, in Tables 1 and 2, respectively.
Analysis of the banding areas presented in Tables 1 and 2, along with the
banding locations of recovered wild-trapped locals presented in Table 3
revealed only one potential breeding area for Hi-Line Canada geese not discussed
by Szymczak, 1971a. Big Horn County in south-central Montana may be an area
occupied by breeding birds migrating into Hi-Line population areas. A total of
five birds banded in Big Horn County were reported recovered in north-central
Colorado during the 1971-72 season (Table 1). All five birds were banded as
wild-trapped locals, two in 1969 and three in 1971.
Phillips County, Montana and the Edmonton-Camrose and Dowling Lake area of
Alberta were well represented in the list of banding areas contributing birds
to the north-central Colorado population (Table 1). In addition, 11 birds
banded as locals during the summer of 1971 in the Cypress Hills area of Saskatchewan were reported taken or were recaptured in north-central Colorado
during the winter of 1971-72 (Table 3). These latter recoveries further substantiate that breeding Canada geese in Cypress Hills area are members of the
Hi-Line goose popualtion. All 15 of the 1971-72 recoveries or recaptures from
the Cypress Hills area as defined by Szymczak 1971a, were banded within one
degree block, 49 degrees latitude and 109 degrees longitude.
Winter Banding-North Central Colorado
Substantial numbers of Canada geese were banded in north-central Colorado in
January 1971. This was the first major banding operation since January 1968.
A total of 991 birds were banded during the 1971 operation. The distribution
of the birds recovered during the 1971-72 season north of Colorado are plotted
in Figures 1 and 2.
Alberta was an important recovery area for birds banded in January 1971. This
was generally not true for the cohort banded in 1968. The recoveries of
"large" birds (size 8 bands) in Alberta, were from scattered locations from
530 latitude south and are not particularly useful in pinpointing additional
unknown breeding areas in Alberta which contribute birds to the Hi-Line population.
The recovery distribution of "small" birds (size 7 bands) in Alberta follows
the same pattern expected for birds of the short grass prairie population
(Grieb 1970). It is of interest that only one "small" bird was reported
recovered in Saskatchewan, and that an atypically large 7 pound immature male
taken south of the major staging area for short grass prairie geese.

�-81-

Table 1. Banding areas outside Colorado of Canada geese recovered in northcentral Colorado during the 1971-72 hunting season.
Area

Direct

Indirect

12

3

Dow.l Ln g Lake Area

18

9

Brooks Area

o

1

3

2

Swift Current-South Saskatchewan
River Area

o

1

Cypress Hills Area

9

3

La Fleche Area

o

2

Teton County

o

1

Blaine Coun ty

4

1

Phillips County

32

39

Fergus County

o

4

Garfield County

o

19

McCone County

o

1

Big Horn County

3

2

o

1

Alberta
Edmonton-Camrose

Area

Saskatchewan
Moose Jaw Area
Eyeb row Lake

CENTRAL FLYWAY
Montana

North Dakota
Ward County

---------------------------------------------------------------------------

�-82Table 1. Banding areas outside Colorado of Canada geese recovered in northcentral Colorado during the 1971-72 hunting season (continued).
Area

Direct

Indirect

o

1

3

3

2

o

1

o

4

o

4

5

3

o

1

5

o

1

o

1

o

1

Davis County

o

1

Weber County

o

1

Wayne County

o

1

99

109

South Dakota
Hughes County
Wyoming
Albany County
Kansas
Phillips County
New Mexico
Mora County
San Miguel County
Socorro County
Chaves County
Texas
Randall County
PACIFIC FLYWAY
Montana
Lake County
Oregon
Lake County
Nevada
Washoe County
Utah

Total

�-83Table 2. Banding areas outside Colorado of Canada geese recaptured, postseason in north-central Colorado, January 1972.
Area

Direct

Indirect

2

2

Dowling Lake Area

o

3

Brooks Area

o

1

Eyebrow Lake

1

o

Cypress Hills Area

2

1

Blaine County

o

1

Phillips County

4

4

o

1

1

1

Mora County

4

2

Socorro County

1

1

Chaves County

1

1

o

1

o

1

16

20

Alberta
Edmonton-Camrose

Area

Saskatchewan
Moose Jaw Area

CENTRAL FLYWAY
Montana

South Dakota
Charles Mix County
Wyoming
Albany County
New Mexico

Texas
Randall County
PACIFIC FLYWAY
Montana
Lake County
Total

�-84Table 3. Banding locations of Canada geese classified as wild-trapped locals
which were recovered or recaptured in north-central Colorado during the
winter of 1971-72.
Year of Banding
Location

Before
1966

1966

1967

1968

1969

1970

1971

2

5

19

Alberta
Edmonton Camrose Area
10 miles E. of Camrose

1

Dowling Lake Aeea
Brooks Area

1

1

Saskatchewan
Cypress Hills Area
Supreme Area

2

Maple Creek Area
Cypress Lake

7

2

4

Moose Jaw Area
Eyebrow Lake

1

CENTRAL FLYWAY
Montana
Blaine County

2

Phillips County

3

Garfield County
Bighorn County

3
3

6

7

8
1

20, (12)

(1)
2

3

North Dakota
Ward County

(1)

PACIFIC FLYWAY
Oregon
Lake County

1

-------------------------------------------------------------------------------

�-85-

Table 3. Banding locations of Canada geese classified as wild-trapped
locals which were recovered or recaptured in north-central Colorado during
the winter of 1971-72 (continued).

Year of Banding
Location

Before
1966

1966

1967

1968

1969

1970

1971

Utah
Weber County

1

Davis County
Wayne Coun ty

1
1

1/ Numbers in parentheses indicate geese classified as immatures when banded
during the summer period.

The most recent harvest distribution for short grass geese on the Canadian
prairies shows an approximate 50-50 split for the two provinces (Szymczak
1971b). As is indicated in the combined Figs. 1 and 2. six small birds
were reported taken in Alberta, compared to the one mentioned above, in
Saskatchewan. This distribution suggests the small geese wintering in northcentral Colorado may be closely associated to the Alberta portion of the
short grass prairie population and do not use the Saskatchewan side to any
great degree. Grieb (1970) suggested that birds of the short grass prairie
population using predominantly Alberta as a staging area were generally
members of a western segment of that population which breed along the
Mackenzie River, and on south into northern Alberta. Additional recovery
years are definitely needed to substantiate any migrational boundaries.
All recoveries in Saskatchewan were south of the South Saskatchewan River
and within the defined Cypress Hills area (Fig. 2). a suspected breeding
concentration area for Hi-Line Canada geese (Szymczak 1971a). The distribution of recoveries in Saskatchewan for 1971 bandings was very similar to
the distribution of the 1968 banded cohort (Szymczak 1971a).
Four of the eight recoveries in Montana for 1971 banded birds were taken
along the Yellowstone River and adjacent areas in Big Horn and Treasure
counties. This area was discussed earlier as a potential breeding area for
Hi-Line geese and these recoveries of winter banded birds lend support to
that claim.
Color Marking--Approximately 130 geese were color-marked during January 1971
post-season banding operations in Colorado and New Mexico. It was hoped
that sightings of these marked birds would help define breeding areas in
the Canadian Provinces. Only one bird was observed. That bird was sighted
in the Cypress Hills area of Saskatchewan, a suspected Hi-Line breeding area.

�,

;
I

'

I

~

/' II I,i

I

!

,

---1-1-tt1' 7·
-T'I

!

Ii,

' , ,----

; ! :

I ! I

!

• Banded as 1mm.

X Banded as Adult

o Banded as lrnrn, (size 7)
e Banded as Adult (size 7)
S Experimental

i

Fig. 1. Distribution of recoveries reported taken outside of Colorado during
the 1971-72 hunting season of birds banded in north-central Colorado,
January 1971.

...J

�I'-

o

Co

o

'J

en
o

-- ~

x

x

-87-

J

I

\-

I

l0 ;

to

o

- - -

I'N

&lt;I)

~

E
&lt;{

"0

"3

E

E

.:!!-

'"

. -

"0

&lt;I)

-- - -

I/)
0
OJ

®

&gt;&lt;
w.
&lt;D

a.

QJ.

c:

0

E
•...

0

"0

~

III

&lt;I)

X

0

•

x

&lt;I)
~ -"0-~c:
c:
c:
0
0
0
en m (Q

::c

~x·

~ .&gt;&lt;&lt;0

....,

;1;

- -)(-

Q

x

Fig. 2. Distribution of recoveries reported taken in the Cypress Hills area and
the two adjacent degree blocks to the west during the 1971-72 hunting season of
birds banded in north-central Colorado, January 1971.

(J')

o
V

�-88-

SUMMARY
Band recoveries during the 1970-71 season presented additional evidence that
the Edrnonton-Camrose and Dowling Lake areas in Alberta, and the Cypress
Hills area of Saskatchewan are Hi-Line population production areas. Further
analysis indicated that the Yellowstone River and adjacent areas in Big
Horn and Treasure counties may be Hi-Line production areas. In addition,
the small Canada geese wintering in north-central Colorado may be members of
the western segment on the short grass prairie population as described by
Grieb 1970.
Migration Routes or Harv~st Patterns
The distribution of harvest during the 1971-72 season of birds banded in
north-central Colorado in January 1968 was essentially the same as during
the 1970-71 season (Table 3). Colorado continues to be the major recovery
area in the south, with Saskatchewan and Montana the northern harvest areas.
Table 3. Distribution of recoveries of Canada geese banded post-season in
north-central Colorado, 1967-68.

Area

Recovery Year
1969-70
1970-71
No. Percent
No. Percent

1971-72
No. Percent

No.

Percent

British Columbia

0

0.0

1

3.7

0

0.0

0

0.0

Alberta

1

4.2

3

11.1

0

0.0

a

0.0

Saskatchewan

11

45.8

9

33.3

3

12.0

3

10.3

Montana

4

16.7

a

0.0

2

8.0

4

13.8

Idaho

1

4.2

1

3.7

a

0.0

a

0.0

South Dakota

0

0.0

1

3.7

a

0.0

1

3.4

Wyoming

a

0.0

1

3.7

a

0.0

2

6.9

Nebraska

a

0.0

1

3.7

a

0.0

1

3.4

North Central

5

20.8

10

37.0

19

76.0

18

62.1

Southeast

1

4.2

a

0.0

1

4.0

a

0.0

Arizona

1

4.2

0

0.0

a

0.0

a

0.0

Total

24

Colorado

27

25

29

�-89-

The first year recovery distribution of birds banded in January 1971 presented a somewhat different picture than the 1968 bandings (Table 4).
Alberta became a recovery area of some significance for "large" birds wearing
size 8 bands, as well as for small birds, as was discussed in a previous
section. Alberta accounted for 18.5 percent of the total number of recoveries, 40 percent of the small birds, 12.7 percent of the large, and 33.3
percent of the experimentals, which were also large. Saskatchewan was the
major northern recovery area for 1971 banded "large" birds as it generally
has been for the 1968 bandings. A smaller percent of the recoveries from
the 1971 banding were taken in Colorado during the 1971-72 season than from
the 1968 bandings. Additional recovery years will be needed to substantiate
any population distributional changes that may have occurred.
Of significant interest is the apparent erratic migration and resulting
winter distribution pattern of small Canada geese9 supposedly members of the
short grass prairie population, which are sometimes present in the Hi-Line
population area of Colorado. In January 1971. 148 birds subjectively judged
to be "small", were banded in north-central Colorado. During the first
recovery year, five of those small birds were taken in Colorado, three in the
north-central portion of the state and two in the normal short grass prairie
population range in the southeast part of the state. Conversely, from
bandings in January and February, 1971 in the southeast part of the state,
one bird was recovered and three recaptured during the winter of 1971-72 in
north-central Colorado. From the 1967-68 season through the 1971-72 season
there have been 3, 7, 4, 9, and 5 short grass prairie banded birds from
southeast Colorado reported taken during the respective seasons in northcentral Colorado. This continued erratic migrational pattern makes it
totally impossible to predict the number of supposedly short grass prairie
birds that can be expected in the Hi-Line area of Colorado during anyone
year.
Distribution
Coordinated counts throughout the Hi-Line population range during the fall
and winter period of 1971-72 indicated a slight increase in total population
numbers over 1970-71 levels (Table 5). The distribution remainded essentially
the same as the previous year, with the bulk of the birds in Colorado.
Within the state of Colorado there were some mentionable shifts in the distribution (Table 6). There was a definite increase in the number of birds
using the Brighton-Greeley-Ft. Morgan area. During the 1970-71 season an
average of four percent of the geese in the Hi-Line area were located in
the Brighton-Greeley-Ft. Morgan area. During the 1971-72 season an average
of 16 percent used that same area. Another population shift occurred
within the Longmont-Boulder-Denver area resulting in a substantial increase
in geese in the Metropolitan Denver area and a reduction in numbers in the
Longmont area, in comparison to the 1970-71 distribution. The distribution
by water area is presented in Table 7.
Again in 1971-72, as during the previous two years, there was a substantial
decrease in the number of geese recorded in the Colorado Hi-Line area
between the late December and early January counts (Table 6). The reason
for this consistent yearly decline is still unknown.

�-90Table 4. Distribution of recoveries of Canada geese banded post-season in
north-central Colorado, 1970-71.

Area
Alberta

Band Size 8-746
No.
Percent

Band Size 7B-148
No.
Percent

Experimental
(Lar~e)-97
No.
Percent

No.

Total
Percent

9

12.5

6

40.0

2

33.3

17

18.3

Saskatchewan 13

18.1

1

6.7

2

33.3

16

17.2

Montana

6

8.3

2

13.3

0

0.0

8

8.6

South Dakota

2

2.8

1

6.7

0

0.0

3

3.2

Wyoming

2

2.8

0

0.0

0

0.0

2

2.2

Nebraska

2

2.8

0

0.0

O.

0.0

2

2.2

NorthCentral

35

48.6

3

20.0

2

33.3

40

43.0

Southeast

0

0.0

2

13.3

0

0.0

2

2.2

Arizona

1

1.4

0

0.0

0

0.0

1

1.1

Pennsylvania

1

1.4

0

0.0

0

0.0

1

1.1

Unknown

1

1.4

0

0.0

0

0.0

1

1.1

Total

72

Colorado

Percent Recovered

15
9.7

6
10.1

93
6.5

9.4

�-91-

Table 5.

Results of Hi-Line Canada goose population inventories, 1971-72.

Location

Nov. 3

Saskatchewan

9,415

Montana

Nov. 23

Dec. 21

Jan. 10

12,902

5,629

323

481

Wyoming

3,900

2,032

3,335

1,150

Colorado

34,419

53,778

55,517

33,520

418

1,113

2,061

1,962

Totals

61,054

62,552

61,236

37,113

Comparati ve Totals 1970-71

31,149

52,926

58,367

44,712

New Mexico

Hunting Pressure and Harvest
An estimated 12,920 hunters harvested 14,771 geese in north-central Colorado
during the 1971-72 season (Tables 8 and 9). Substantial increases in
harvest on a percentage basis were recorded in Weld, Boulder, and Morgan
counties (Table 10). The harvest in Larimer County remained essentially
the same, with the Adams County harvest declining slightly (Table 10). The
large percentage increase in harvest in Boulder County was probably the
result of a decrease in the size of the closed area in that county.
Again the majority of the active hunters did not harvest any geese (Table
11). However, this majority is decreasing in size on a yearly basis.
San Luis Valley
Foreign Band Recoveries
The number of foreign bands reported recovered in the San Luis Valley during
the first two San Luis Valley Special goose seasons are presented in Table 12.
The recoveries show very little consistency in terms of the area of banding
between the two years. Birds from Hi-Line areas (Garfield County) are
represented in the 1971-72 season recoveries but defnitely not to the degree
exhibited in 1970-71 (Dowling Lake, Phillips County, MOntana; north-central
Colorado) .

�-92-

Table 6.

Winter inventories of the Colorado Hi-Line Canada goose population.

Count Date

Fort Co11insLoveland

LongmontBou1derDenver

BrightonGree1eyFort Morgan

9,739

2,883

991

13,613

12,217

4,029

678

16,924

15,848
20,905
19,693

3,461
4,236
4,874

2,667
1,170
775

21,976
26,311
25,342

8,737
31,350
18,522
30,650

2,255
3,782
5,668
5,060

390
1,374
1,259
1,914

11,382
36,506
25,499
37,624

12,612
29,970
36,034
19,879

1,690
16,710
12,664
15,566

348
1,370
3,055
2,425

14,650
48,050
51,753
37,870

25,699
31,072
31,516
19,117

1,815 1/
9,181
19,525
10,742

6,905
13,525
4,476
3,661

34,419
53,778
55,517
33,520

Total

1967
January 9
1968
January 10
1968-69
November 20
December 19
January 2 &amp; 13
1969-70
November 5
November 28
December 23
January 6

1970-71
November 4
November 23
December 22
January 6
1971-72
November 4
November 23
December 21
January 10

1/ Denver area not included.

�-93-

Table 7.

Results of north-central Colorado goose surveys, 1971-72.

Area

Nov. 3

Nov. 23

Dec. 21

Jan. 10

3,500
2,450
340
2,000

2,360
170

2,100
19

1,195
11

8
1

o

o

107

10
253

Ft. Collins-Loveland
Lindenmeier Lake
Reservoir No. 8
Reservoir No. 8 Annex
Elder Reservoir
Reservoir No. 5
Reservoir No. 6
Bureau of Standards
Douglas Reservoir
Rocky Ridge Reservoir
Cement Plant Reservoir
Terry Lake
Mountain Supply Reservoir
College Lake
Herring Lake
Claymore Lake
Sterling Pond
Watson Lake
Dean Lake
Greenwalt (Nelson) Lake
Fossil Creek Reservoir
Boyd Lake
Horseshoe Lake
Hollister Lake
New Windsor Rservoir
Woods (Eaton) Lake
Timnath Reservoir
Cobb Lake
Black Hollow Reservoir
Park Wood Lake
Boxelder Reservoir
Warren Reservoir
Dixon Reservoir
Reservoir No. 15
Owl Creek Reservoir
Newel Lake
Reservoir No.4
Demmel Lake
Sub-total

o

o

1,325

63
31
127

o

o

o

10

o
2,400
130
1,000

o
1,200
156
320
311

o
1,900
320
21

o
3,000

o
1,050
350
1,050
273
850
18
575
550
600

o

o
o
25,699

475
2,728

o
2,140
2,395
1,590
2,559
1,138
1,456

o

o

o
o

o

426

200

o

o

o
o

4,270

757

o

o

3,252

5,150

o

o

651
2,610
1,045
661

3

o

1,500
200
1,000
500
5,500
250
63
218
84
425
2,510
665

2,500
3,500

411
656
1,195
242

o
75
3,700
300
130
3,087
300
51
517
95
340

o
150
8,750
350
51
530

o
2

323

4

o

o

o

335
178
400

154

o

o

o
o

o

o
o

310

o
128

o

65

o

31,072

31,516

19,117

----------------------------------------------------------------------------

�-94-

Table 7.

Results of north-central Colorado goose surveys, 1971-72 (cont.).

Area

Nov. 3

Brighton-Gree1ey-Ft.

Dec. 21

Jan. 10

850
500
525
15
0

2,500
1,955
50
0
20

1,450
0
525
0
0

2,476
0
460
0
0

6,905

13,525

4,476

3,661

34,419

53,778

55,517

33,520

Morgan Area (continued)

Empire Reservoir
Riverside Reservoir
Jackson Reservoir
Pond near Cornish
K-4 Ranch
Sub-total
Grand Total

Table 8.
area.

Nov. 23

Hunter activity and success in north-central Colorado goose permit

Number
Permits Issued

Est. Number
Active Hunters

Average
Days Hunted

Average
Seasonal Bag/Hunter

1964-65

1,608

1,427

6.1

0.48

1965-66

2,335

1,578

6.5

0.53

1966-67

3,996

2,910

5.1

0.41

1967-68

3,000

2,256

6.7

0.55

1968-69

3,000

2,379

9.2

1.11

1969-70

8,342

6,149

7.4

0.68

1970-71

13,611

11,187

6.9

1.08

1971-72

14,847

12,920

7.3

1.14

Year

�-95-

Table 7.

Results of north-central Colorado goose surveys, 1971-72 (cont.).

Area

Boulder-Longmont

Nov. 3

Nov. 23

Dec. 21

Jan. 10

450
430
45
0
0
0
0
0
0
850
20
20
0
0
0

33
4,450
52
29
25
0
300
300
150
2,450
22
0
44
25
150

44
4,625
360
0
25
0
50
150
0
9,000
0
0
18
0
0

28
1,286
930
12
0
500
100
0
100
4,500
0
0
77
0
0

1,815

8,030

14,272

7,533

0
0
0
0
0
0
0
0
0
0
0
0
0
0

0
0
57
0
107
0
0
84
3
63
11
43
783
0

133
95
1,970
382
0
37
'0
0
0
74
0
462
2,100
0

153
291
1,870
0
257
0
256
0
0
0
0
0
0
382

0

1,151

5,253

3,209

260
2,800
150
0
1,805

1,200
1,500
3,000
3,000
300

900
700
1
500
400

350
275
0
100
0

Area

Ish Lake
Terry Lake (Longmont)
McIntosh Lake
Foothills Reservoir
Faivre Ponds
Swede Lake
Dodd Lake
Baller Lake
Boulder Reservoir
Valmont Reservoir
Crystal Lake
Highland Lake
Union Reservoir
Steel Lake
Lugerman Lake
Sub-total

".

Denver Area
Sloans Lake
Standley Lake
Bowles Lake Area
Marston Reservoir
Denver City Park
Great Western Reservoir
Cherry Creek Reservoir
Washington Park
Rocky Mountain Arsenal
Tule Lake
Lower TUle Lake
Kings Pond
Green Gables Country Club
Englewood Reservoir
Sub-total
Brighton-Greele~-Ft.

Morgan Area

Barr Lake
Horsecreek Reservoir
Prospect Reservoir
Latham Reservoir
Milton Reservoir

----------------------------------------------------------------------------

�-96-

Table 9. Distribution of harvest, by county, in the north-central Colorado
goose permit area.
Year

Larimer

Weld

1964-65

504

181

1965-66

665

144

29

838

1966-67

764

409

11

1,184

1967-68

944

265

37

1,246

1968-69

1,584

886

161

2,631

1969-70

2,431

1,112

383

146

100

4,172

1970-71

7,486

3,544

620

93

370

12,112

1971-72

7,723

4,484

1,804

464

296

14,771

Boulder

Morgan

Adams

Total
685

Table 10. Hunting pressure and harvest, by county, in north-central Colorado, 1971- 72.

Larimer County

Estimated Number
1970-71

Estimated Number
1971-72

Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

6,977 (+ 328)
43,046 (+3,851)
7,541 (± 868)
6.2
1.08
0.175

6,760 (+ 285)
41,535 (+3,217)
7,723
693)
6.1
1.14
0.186

4,293 (+ 308)
22,557 (±2,677)
3,497 (+ 595)
5.3
0.81
0.155

5,911 (+ 280)
31,914 (+3,126)
4,484 (± 514)
5.4
0.76
0.141

(±

Percent Change
From 1970-71

+
+

+

3.1
5.5
2.4
1.6
5.6
6.3

Weld County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

+
+
+

+

37.7
41.5
28.2
1.9
6.2
9.0

----------------------------------------------------------------------------

�-97-

Table 10. Hunting pressure and harvest, by county, in north-central
1971-72 (continued).

Colorado,

Estimated Number
1970-71

Estimated Number
1971-72

1,231 (+ 186)
7,543 (+2,207)
620 (+ 229)
6.1
0.50
0.082

2,296 (+ 206)
14,961 (+1,962)
1,804 (+ 353)
6.5
0.79
0.121

+ 86.5
+ 98.3
+ 191.0
+
6.6
+ 58.0
+ 47.6

759 (+ 147)
2,147 (+ 580)
93 (±
64)
2.8
0.12
0.043

1,104 (+ 149)
3,774 (+ 738)
464 (+ 166)
3.4
0.42
0.123

+ 45.5
+ 75.8
+ 398.9
+ 21.4
+ 250.0
+ 186.0

592 (+ 131)
2,294 (+.: 818)
361 (+ 147)
3.8
0.60
0.157

545 (+ 107)
2,288 (+ 681)
296 (± 152)
4.2
0.54
0.129

7.9
0.3
18.0
10.5
10.0
17.8

11,187 (+ 254)
77,587 (+4,403)
12,112 (±:1,005)
6.9
1.08
0.156

12,920 (+ 189)
94,472 (+4,027)
14,771 (+ 850)
7.3
1.14
0.156

Pe rcen t Change
From 1970-71

Boulder County 1/
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip
Morgan County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip
Adams County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

+

Entire Permit Area
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

1/ Boulder County permit area enlarged in 1971-72.

+
+
+

+
+

15.5
21.8
22.0
5.8
5.6

o

�-98-

Table 11. Estimated distribution of season bag for active hunters in the
north-central Colorado permit area.

Season Bag

Total

1969-70
Number
Percent

Hunting Season
1970-71
Number
Percent

1971-72
Number
Percent

0

4,263

69.3

6,588

58.9

7,006

54.2

1

836

13.6

1,620

14.5

2,209

17.1

2

452

7.4

1,073

9.6

1,428

11.1

3

238

3.9

564

5.0

768

5.9

4

153

2.5

528

4.7

633

4.9

5

138

2.2

342

3.1

371

2.9

6

69

1.1

472

4.2

505

3.9

6,149

100.0

11,187

100.0

12,920

100.0

The most informative recoveries are those from Wheatland Reservoir in Albany
County, Wyoming. Evidence that sub-adult geese from the San Luis Valley have
been migrating to Wheatland to molt has been accumulating since goose
restoration efforts began at San Luis Lake in the San Luis Valley in 1969.
Most of this evidence has been documented by Szymczak, 1970 and Szymczak 1971c.
The five recoveries of Wheatland banded birds is the most conclusive positive
evidence uncovered to date. Three of the Wheatland birds recoverd were
banded in 1970 and two in 1971.
Distribution and Harvest
Aerial surveys conducted in the San Luis Valley during the 1971-72 season
indicated that for the second consecutive year the majority of the birds
were utilizing the Monte Vista National Wildlife Refuge (Table 13). After
the original count on October 31, the birds not located on the Refuge were
found on the rivers. There were generally about the same number of geese in
the San Luis Valley during the 1971-72 season as during the 1970-71 season
(Table 14).
A total of 350 permits were issued authorizing the taking of one goose for
the 1971-72 special San Luis Valley season. According to the special goose
harvest survey, an estimated 296 hunters harvested 177 geese for an average
season bag of 0.60 (Tables 15 and 16). Approximately 60 percent of the birds
harvested were reported taken within two miles of the Monte Vista National
Wildlife Refuge. The majority of the remainder of the harvest was taken
along the Rio Grande River between Monte Vista and Alamosa.

�-99-

Table 12. Banding areas outside the San Luis Valley of geese recovered
within the San Luis Valley during the 1970-71 and 1971-72 hunting seasons.

1970-71

Area

Number of
Recoveries

Recovery
Date

2

1-2-71

1971-72

Number of
Recoveries

Recovery
Date

2

12-2-71

Alberta
Dowling Lake

1-5-71

Montana
Phillips County

1

11-14-70

Garfield County

12-11-71

Wyoming,
Albany County

1

1-12-71

5

(Wheatland Reservoir)

11-27-71
12-17-71
1-15-72
1-16-72
3-4-72 1/

Fremont County (Ocean Lake)

1

11-4-71

1

11-23-71

1

11-24-71

Colorado
North-central

2

11-6-70
11-14-70

1

11-6-71

Texas
Waggner Ranch-Panhandle
New Mexico
Soccoro County
Wisconsin
Columbia County

1/ Found dead.

Assumed shot during the season.

�-100-

Table 13. Results of Canada goose surveys during the 1971-72 hunting season
in the San Luis Valley.

Area

October 31

Number of Geese
November 23 December 21

January 6

Monte Vista National
Wildlife Refuge

1,000

400

980

1,100

125

131

Rio Grande River
Del Norte to Monte Vista

10

Monte Vista to Alamosa

200

483 );/

140

0

Alamosa South

45

0

0

0

0

0

10

55

San Luis - Head Lakes

300

0

0

0

Smith Reservoir

15

0

0

0

Pond North of La Jara

0

30

0

0

1,570

913

1,255

1,286

Conejos River

Total

!/ Includes birds from Del Norte to Monte Vista, if present.
Table 14. Comparison of 1970-71 to 1971-72 Canada goose winter inventories
in the San Luis Valley.

Year

October

1970-71
1971-72

1,570

Number of Geesell
November
December

January

1,490

1,050

1,261

913

1,255

1,286

!/ Compared monthly, inventories for the two years were taken within five
days of each other.

�-101-

Table 15.

Hunter activity and success in the San Luis Valley permit area.

Year

Number
Permits Issued

Est. Number
Active Hunters

Days Hunted

1970-71

200

164

3.4

0.65

1971-72

350

296

4.2

0.60

Average

Average
Seasonal Bag/Hunter

Table 16.
area.

Distribution of harvest, by county, in the San Luis Valley permit

Year

Rio Grande

Alamosa

Saguache

Conejos

Total

1970-71

74

31

2

o

107

1971-72

121

50

1

5

177

Harvest and hunter activity for the entire permit area for both the 1970-71
and 1971-72 seasons are presented in Table 17.

Table 17. Hunting pressure and harvest in the San Luis Valley goose permit
area, 1971-72.

Estimated Number
1970-71

Estimated Number
1971-72

Total Individual Hunters

164 (+ 3)

296 (+ 5)

Total Hunter Trips

556 (+ 20)

1,236 (+ 36)

Total Geese Bagged

107 (+ 4)

177 (+

Entire Permit Area

Average Hunter Trips/Hunter

3.4

4.2

Average Bag/Hunter

0.65

0.60

Average Bag/Hunter Trip

0.19

0.14

6)

�-102-

West-central

Colorado

A limited Canada goose hunting season was initiated in west-central Colorado in 1971. The west-central Colorado population was established through
restoration efforts along the Colorado Ri'Terbelow Grand Jucntion which
began in 1967. The resulting breeding population has been essentially nonmigratory and birds from other populations have begun using the area during
the fall and winter period.
Hunter Activity and Harvest
One hundred and fifty permits were issued authorizing the taking of one
Canada goose within the west-central goose permit area. The season ran from
November 27, 1971 to December 19, 1971. The permit area included all of Mesa
and Garfield counties. An estimated 112 hunters harvested 33 geese for an
average seasonal bag of 0.29 birds per active hunter (Tables 18 and 19).

Table 18. Hunter activity and success in the west-central Colorado goose
permit area.

Year

Number
Permits Issued

Est. Number
Active Hunters

Average
Days Hunted

Average
Seasonal Bag/Hunter

1971-72

150

112

2.9

.29

Table 19. Hunting pressure and harvest in the west-central Colorado goose
permit area, 1971.
Total Individual Hunters

112 (+ 4)

Total Hunter Trips

329 (+ 20)

Total Geese Bagged

33 (+ 4)

Average Hunter Trips/Hunter

2.9

Average Bag/Hunter

0.29

Average Bag/Hunter Trip

0.10

Approximately 55 percent of the birds harvested were taken within five miles
of Highline Reservoir. Another 28 percent were taken along the Colorado
River below Loma.

�-103-

The results of pre- and post-season aerial inventories of birds within the
permit area are presented in Table 20.

Table 20.

Canada goose inventories, west-central Colorado.

Date
November 24, 1971
January 5, 1972

Area
Highline Lake

365

53

Silt to Palisade

49

259

Grand Junction to Fruita Bridge

o

6

Fruita Bridge to Utah Stateline

71

583

Colorado River

Gunnison River
Black Canyon to North Fork

21

Confluence North Fork and Gunnison

35

Delta to Grand Junction

32

Total

485

989

LITERATURE CITED
Grieb, J. R. 1970. The short grass prairie Canada goose popUlation.
Monogr. No. 22. 49 pp.

Wild!.

Szymczak, M. R. 1970. Experimental studies on improving status of Canada
goose populations. Colo. Div. Game, Fish and Parks, Game Res. Rept., Fed.
Aid Project W-88-R. October. 81-93 pp.
Szymczak, M. R. 1971a. Studies of Canada goose popUlations in Colorado
transplant areas. Colo. Div. Game, Fish and Parks, Game. Res. Rept., Fed.
Aid Project W-88-R. October. 165-189 pp.
Szymczak, M. R. 1971b. Arkansas Valley Canada goose flock management studies.
Colo. Div. Game, Fish and Parks, Game Res. Rept., Fed. Aid Proj. W-88-R.
October. 129-164 pp.
Szymczak, M. R. 1971c. Experimental studies on improving status of Canada
goose populations. Colo. Div. Game, Fish and Parks, Game Res. Rept., Fed.
Aid Project W-88-R. October. 115-127 pp.

Prepared by

711.;.&amp;,.. P aZ

4r G-L

Michael R. Szymczak
Assistant Wildlife Researcher

��October,

-105-

1972

JOB PROGRESS REPORT

State of

C~O~L~O~RA~D~O~~ _

Project No.

W-88-R-17

Work Plan No.
Job Title

December

of Mallard

Management

1, 1971 through April

Richard Hopper, Michael Szymczak,
Dale Coven and Howard Funk.

Bird Investigations

6

Job No.

Investigations

Period Covered:
Personnel:
Kitzmiller,

3

Migratory

Units of Eastern

Colorado

1, 1972

Clait Braun, Jack Corey, Robert

ABSTRACT
Post-season mallard trapping and banding efforts for the winter of 1971-72
were again successful.
Division personnel banded 6,564 mallards in 9 study
units in eastern Colorado from mid-January through most of February, 1972.
Winter census figures peaked higher in 1971-72 than the previous year to a
new high since 1957 and 1958. Early November, 1971 produced a high of
422,000 birds in eastern Colorado, 453,000 were censused in late November,
and peak numbers of 566,000 were observed in mid-December.
The January
count dropped somewhat to 344,000 birds.
Sex ratio counts averaged about
58 percent males in January and February populations, similar to those of
previous years, again indicating the point system drake mallard-oriented
regulations had little effect on altering sex ratios of wintering mallards
in the western Central Flyway.
Hunting pressure was indicated to have
increased substantially in 1971-72 over the previous year and harvest increased
correspondingly.
Greatest increase in pressure and harvest occurred in
north-central Colorado.

��-107-

INVESTIGATIONS

OF MALLARD MANAGEMENT UNITS OF EASTERN COLORADO

Howard D. Funk

Banding of wintering populations of mallards post-season in eastern Colorado,
initiated in 1963-64, was continued in 1971-72. Purpose of the effort is
to contribute data to the cooperative Central Flyway program to determine
the feasibility of splitting the Flyway into two management units. Recognition of the western or High Plains Unit has been progressing well in the
past few years. Also, more intensive analysis is being accomplished within
Colorado to see if management can be refined even more so on a unit basis
within Colorado. Further, mallards banded in Colorado since initiation of
the study have been aged (adult or immature) according to wing characteristics, which is a very reliable method. Thus, differential mortality and
survival estimates, as well as differing migration routes will be discernable over the years by age as well as sex. Monitor banding will continue
in Colorado until the initial cohort of birds banded in 1963-64 are eliminated from the population. This report mainly covers the results of banding
and survey data collected during the segments.

P.

S.

OBJECTIVE

To develop a harvest formula for eastern Colorado management units.
SEGMENT OBJECTIVES

1.

To analyze for each of nine eastern Colorado study areas of wintering
mallards, all data collected from 1963-64 through the 1970-71 hunting
season on the discreteness of populations and publish results in an
appropriate journal.

2.

To determine the feasibility of requesting special hunting regulations
for individual mallard management units within Colorado.

3.

To trap, band, age, and sex samples of mallards during winter in each
management unit, based on results of final analYSis, of sufficient size
to allow adequate monitoring of each population.

4.

To investigate the ability and/or willingness of hunters to abide by the
point-system regulation established for the 1971-72 duck hunting season.
METHODS AND MATERIALS

Cage traps were again the main type of trap utilized although some birds
were captured with cannon-nets. Trapping was again conducted from immediately after season, mid-January, until almost the end of February. Because

�-108-

data have shown the San Luis Valley no longer seems to be an important
wintering area for northern ducks, no birds were trapped in that area.
study areas were again reduced to eight units.

Thus,

Periodic aerial censuses were again conducted to estimate movements and
numbers of birds present in the State. Ground sex ratio counts were conducted
as described in previous segment reports.
A tape containing updated information on band recoveries through the 1970-71
season was obtained from the Bird Banding Laboratory. Data were extracted
and made ready for analysis during the segment.

RESULTS AND DISCUSSION
Trapping and Banding
Banding quotas of six to eight hundred mallards were set for each of the eight
study areas, and quotas were reached or exceeded in all but one area (Table 1).
Management Units 11 and 12 are listed separately in Table 1, but for this
study are considered one unit. Good weather conditions for trapping were
conducive to excellent results, with the trapping effort being completed
prior to the end of February.
Attempts to band nearly equal numbers of birds by age (adult and immature)
and sex were quite successful (Table 1). As usual, adult male mallards were
the easiest to obtain, followed by immature males, immature females and adult
females. Many birds, especially adult males, were released unbanded so as
not to cheapen the band recovery rates by excessive numbers banded for any
age or sex group.

Winter Sex Ratio Counts
Results of sex ratio ground counts are displayed in Table 2 by area. A
total of 11,484 birds were observed with an overall rate of 58.1 percent
males present. This is close to rates observed in past years, those
averaging about 60 percent males. There was some variation observed among
areas and dates of observation, but this can be expected. The important
factor is that ratios continue to be high toward males, suggesting males
continue to have higher survival rates than females even though regulations
have been aimed toward greater harvest of the drake. The data also suggest
that Colorado, or even western Central Flyway hunting pressure is not sufficient
to have greatly altered the sex ratios under drake-oriented regulations in
recent years.
Winter Aerial Surveys
Dates and results of the periodic aerial counts are shown in Table 3 by
area. Birds were in good supply over most of eastern Colorado by early
November and peaked to near record high levels in mid-December. Numbers
dropped by early January, which usually occurs in most years. Bonny Reservoir had record numbers of birds from mid-November through the January count;
about twice the average wintering population in comparison with other years.
Ample food in the area, a relatively mild winter, and little hunting pressure
all probably contributed to this event.

�Table 1.
1971-72.

Numbers and percentages of mallards in the banded sample by age, sex, and area, eastern Colorado,

Adult
Female
Male
Percent
No.
Percent

Immature
Male
Female
Percent
No.
Percent

No.
Banded

No.

(1) Sterling-Julesburg

900

252

28.0

169

18.8

255

28.3

224

24.9

(2) Fort Morgan-Sterling

800

200

25.0

239

29.9

193

24.1

168

21.0

(3) Greeley-Fort Morgan

800

207

25.9

161

20.1

209

26.1

223

27.9

(4) Fort Collins

831

205

24.7

155

18.6

217

26.1

254

30.6

(6) Denver-Greeley

900

232

25.8

132

14.7

228

25.3

308

34.2

Management Unit

No.

I

I-'

(9) Bonny Reservoir

1,000

257

25.7

250

25.0

247

24.7

246

24.6

(11) Two Buttes Reservoir

487

122

25.0

175

35.9

66

13.6

124

25.5

(12) Rocky Ford-Lamar

400

197

49.2

98

24.5

42

10.5

63

15.8

(13) Pueb1o~Rocky Ford

446

180

40.4

73

16.4

89

20.0

104

23.2

6,564

1,852

28.2

1,452

22.1

1,546

23.6

1,714

26.1

Totals

0

\0
I

�-110-

Table 2.

Mallard post-season sex ratio counts by study unit, winter of 1971-72.

Managemen t Uni t and Date

No. Ducks Counted
Male
Female
Total

Percent
Males

Ster1ing-Julesburs ~Unit 1)
2-17-72 South platte River
2-18-72 Jumbo Reservoir

312
322

197
192

509
514

61. 3
62.6

Fort Morgan-Ster1ins (Unit 2~
2-17-72 South Platte River
2-18-72 South Platte River

303
302

210
236

513
538

59.1
56.1

Gree1e~-Fort Morgan (Unit 3)
2-17-72
South Platte River
2-18-72
South Platte River

252
194

180
156

432
350

58.3
55.4

Fort Collins Area (Unit 4)
2-7-72
Woods Lake
2-18-72 Hollister Lake

289
308

229
229

518
537

55.8
57.4

Denver-Gree1e~ (Unit 6~
2-17-72 South Platte River
2-18-72 South Platte River

309
286

199
217

508
503

60.8
56.8

Bonn~ Reservoir Area (Unit 9)
1-15-72 Bonny Reservoir
1-16-72 Bonny Reservoir
1-17-72 Bonny Reservoir
1-17-72 Bonny Reservoir
1-17-72 Bonny Reservoir
1-18-72 Bonny Reservoir
1-19-72 Bonny Reservoir
2-11-72 Bonny Reservoir
2-11-72 Bonny Reservoir
2-12-72 Bonny Reservoir

310
310
299
270
280
295
303
260
288
295

204
231
239
248
229
207
205
240
226
239

514
541
538
518
509
502
508
500
514
534

60.3
57.3
55.6
52.1
55.0
58.8
59.6
52.0
56.0
55.2

Arkansas Val1e~ (Units 11 and 12)
1-15-72 John Martin Reservoir
1-15-72 Verhoeff Reservoir
1-16-72 Turk's Pond
1-16-72 Two Buttes Reservoir
2-3-72
Two Buttes Reservoir

75
54
171
259
327

49
47
64
211
127

124
101
235
470
454

60.5
53.5
72.8
55.1
72.0

6,673

4,811

11,484

58.1

Total

�Table 3.

Aerial duck counts by date interval and study unit, eastern Colorado, 1971-72.

Management Unit

Nov. 3-4

Number of Ducks Counted
Nov. 22-24
Dec. 20-22

Jan. 4, 6

(1)

Sterling-Julesburg

64,500

55,100

60,030

21,800

(2)

Fort Morgan-Sterling

43,200

34,800

78,100

34,600

(3)

Greeley-Fort Morgan

53,400

64,800

81,200

50,250

(4)

Fort Collins

144,820

82,055

58,200

31,850

(6)

Denver-Greeley

65,260

63,700

116,350

76,000

(9)

Bonny Reservoir

30,000

63,200 !/

77 ,000

69,000

Y

71 ,400

35,000

(10-13)
(15)

Arkansas Valley

San Luis Valley

Total

!/ December 6, 1971.
~/ Not censused.

1/ San Luis Valley not included.

21,100

69,700

-- 'fj

19,635

24,148

25t940

452,990

566,428

344,440

422,280

]j

I

I-'
I-'
I-'
I

�-112-

Wing Surveys
As in previous years, age and sex ratios of mallards harvested in Colorado
were obtained through the annual Wing Bee conducted by the Bureau of Sport
Fisheries and Wildlife. These data are presented in Bureau administrative
reports and will be utilized in final analyses and reports.

Hunting Pressure Surveys
Duck hunting pressure and harvest by unit was again examined (Table 4).
Number of hunters and harvest estimated from the small game harvest survey
were paired with estimated numbers of birds censused by unit. As the year
before, the mid-December count data were utilized in the comparison. This
method has to be considered quite crude and comparisons are good only for
the year and not between years, mainly because of varying numbers of birds
present. However, the comparison continues to show relative hunting pressure
is greatest nearer the more highly population cities. The Bonny Reservoir
area (Unit 9) continues to have fewest hunters present per available birds.
Unit 1 also dropped quite low in relation to other areas. This is partially
explained by an indicated drop in numbers of hunters from 1970 while all
other areas increased in numbers as estimated from the harvest survey. The
Arkansas Valley also increased in comparative. pressure.
Evaluation of the Point-system Duck Season
Colorado again participated in gathering hunter performance data to evaluate
the ability and/or willingness of hunters to abide by point-system regulations.
Data were gathered and submitted to Patuxent Research Station for analysis
by a group of State and Federal personnel. A total of 28 usable hunter checks
were obtained in the State. Findings from all participating states were
reported in the Bureau's Administrative Report No. 215 (Kimball et al.). As
per previous years, data indicated the assigned point values tended to direct
hunting pressure away from high point birds, such as the mallard hen, and
toward lower point birds, including mallard drakes. Behavior of hunters was
indicated to have been good with relatively few violations directly related
to point-system regulations observed.
Analysis of Band Recovery Data
Considerable time was spent in analysis of band recovery data through the
1969-70 hunting season. Also, the Bureau furnished an update tape which was
utilized for extracting data for the 1970-71 season. These data are in the
process of being combined.
One of the major efforts during the next segment will be to ascertain which
of the management units contain duck populations with like migration and
harvest characteristics. Preliminary analyses suggest Units 1, 2, and 9 are
similar, Units 3, 4, and 6 seem closely related, and that Units in the
Arkansas Valley are all closely related. This information will be utilized
mainly for distinguishing areas of monitor banding in the continuing cooperative study of the High Plains Mallard Management Unit.

�-113-

Table 4. Hunter pressure, harvest and population survey data for comparison
between management units and areas, 1971-72 duck season.

.;»

Estimated
Estimated
Duck
Ducks
Hunters '];/ Bag.ged J:../

Average
Bag Per
Hunter2/

Number
Number Hunters
Ducks
Present Per 100
Censused3/ Ducks Censused

Northeast
Unit 1

1,652

12,400

7.5

60,030

2.8

Unit 2

3,273

27,292

8.3

78,100

4.2

Unit 9

1,038

6,634

6.4

77 ,000

1.3

5,963

46,326

7.8

215,130

2.8

Unit 3

6,265

54,033

8.6

81,200

7.7

Unit 4

4,248

25,988

6.1

58,200

7.3

Unit 6

6,438

49.716

7.7

116,350

5.5

16,951

129,737

7.7

255,750

6.6

Unit 10

1,069

6,113

5.7

Unit 11

297

811

2.7

Unit 12

2,013

14,312

7.1

Unit 13

1,434

12,342

8.6

Total

4,813

33,578

7.0

71,400

6.7

Total
North Central

Total
Southeast

l/ See Fig. 1 for management unit locations.
J:../ Figures from 1971-72 Colorado small game harvest survey.

1/ Data from Table 3, December 21 census.
~/ Inventory data for Arkansas Valley usable only as total.

LITERATURE CITED
Kimball, C. F., R. Bridges, C. D. Crider, J. H. Dunks, R. M. Hopper, D. D.
Kennedy, J. McDaniels and W. Okamoto. 1972. Results in the hunter performance survey in the point regulation test states, 1971. Mig. Bird
Pop. Sta. Admin. Rept. No. 215. Bur. of Sport Fish. &amp; Wi1d1., Laurel,
Md. June 28. 21 pp.
Prepared by

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�October,

-115-

1972

JOB PROGRESS REPORT

State of

COLORADO

Project No.
Work Plan No.
Job Title
Period Covered:

Migratory

4

Bird Investigations

Job No.
Trapping

3

and Banding Doves

April 1, 1971 through March 31, 1972

Personnel:
Charles Hayes, Dale Horne and Jack Randall, Bureau of Sport
Fisheries and Wildlife; Clait E. Braun, Howard D. Funk, J. Edward Kautz,
Wilbur Ladd, Brett Petersen, Michael Robinson, Steve Steinert and J. Allen
White, Colorado Division of Wildlife.

ABSTRACT
Efforts initiated in 1964 to trap and band samples of mourning doves
(Zenaidura macroura) in Colorado were continued in 1971. Cooperative Federal
and State efforts resulted in 5,233 birds being newly banded.
Of this total
2,470 were immatures;
8 were unsexed adults; 1,565 were adult males; while
1,190 were adult females.
Assigned quotas were accomplished for all areas
except extreme eastern Colorado.
Wing molt data from trapped and harvested
imrnatures suggested three hatching peaks in 1971. Peak hatching periods were
late May-early June, late June-early July, and late July-early August.
A
sample of 310 hunter-harvested birds, mostly from southeastern Colorado,
gave a young-to-old ratio of 11.9:1, more than double that obtained in 1970.
Wing molt data from harvested immatures in this sample indicated that most
early hatched young migrated from Colorado prior to September 1. Thirty-six
band recoveries were reported in 1971 from doves banded by Division personnel.
Nineteen (52.8 percent) of the bands I'eported were from doves killed in Colorado.
Apparent hunting mortality for doves ~anded in Colorado continues to be less
thant 3 percent.
.

��-117-

MOURNING DOVE TRAPPING AND BANDING
Clait E. Braun

Intensive efforts to trap and band mourning doves in Colorado initiated
in 1964 were continued in 1971, in cooperation with personnel of the
U. S. Fish and Wildlife Service. Data presented in this report are those
collected in 1971, the seventh year of this continuing investigation.
P. S. OBJECTIVE
To investigate migration patterns and mortality rates of mourning doves
banded in Colorado by age, sex, and area.

SEGMENT OBJECTIVES
1.

To trap and band mourning doves in three selected areas of Colorado for
the purpose of obtaining migration, life history, and annual mortality
data.

2.

To estimate harvest size and hunter success.

METHODS AND MATERIALS
Methods and materials used in 1971 were similar to those described in
earlier reports and summarized by Braun (1970). In addition, less than
100 doves were trapped in cannon nets in 1971 during trapping operations
for band-tailed pigeons (Columba fasciata).

DESCRIPTION OF TRAPPING AREAS
Most trapping sites in Eastern Slope areas were similar to those described
earlier (Braun 1970). Two new areas were utilized in 1971 east of the
Rocky Mountains in addition to those also trapped in 1970. These were near
Bonny Reservoir and at Fort Carson Military Reservation. At Bonny Reservoir,
trapping was conducted near plantings of introduced trees adjacent to grainfields (primarily wheat, Triticum aestivum), while trapping at Fort Carson
was accomplished primarily near established cultivated food plots (primarily
wheat and barley, Hordeum vulgare). West Slope trap sites were essentially
those also used in 1970, with the exception of one site. This site in
Unaweep Canyon was a livestock feeding area surrounded by fairly dense stands
of oak (Quercus gambelli) and sparse pinyon pine (Pinus edulis).

�-118-

RESULTS AND DISCUSSION
Trapping and Banding
Cooperative Federal and State trapping and bartding efforts resulted in
5,233 doves being newly banded in 1971. Of this total, 4,401 were banded
by Division personnel. One hundred and twenty-two of the immature doves
caught were additionally banded with reward bands as part of a cooperative
Central Management Unit study to ascertain band reporting rates. Doves
were banded at eight major locations on the Eastern Slope and at four
major sites west of the Continental Divide. Additionally, some doves were
banded at five other scattered locations in conjunction with band-tailed
pigeon trapping activities. The banding goal of 4,000 birds was accomplished
with over 1,000 being banded in western Colorado and over 2,000 banded along
the eastern slope of the Rockies. The 1,000 quota for extreme eastern
Colorado was not met as only 602 birds were trapped at two different locations. Sex and age distribution of birds banded by area are presented in
Table 1.
As in earlier years of this study, more adult doves were trapped than immature,
although the disparity was not great in 1971. Differences in numbers of
adults and immatures trapped is undoubtedly a function of time of season of
trapping effort. Trapping at Meeker, Durango, and Fort Collins was accomplished primarily in June and less than 25 percent of the doves caught were
immatures. On the other hand, trapping at Denver and Fort Garland was
principally in August and over 60 percent of the birds trapped were immatures.
However, trapping in July near Fort Morgan resulted in about 90 percent of
the doves caught being young-of-the-year.
This illustrates the infeasibility
of using age ratios in trapped samples to estimate productivity. While
significantly more doves were banded in 1971 than 1970 (5,233 versus 2,883)
percentages of males (29.9 to 28.5), females (22.8 to 24.4) and immatures
(47.3 to 47.1) banded were not significantly different.

Hatching Data
Wing molt data were available for 2,467 of the 2,470 immature doves trapped
and banded in 1971. Estimated hatching dates (Allen 1963) are presented in
Table 2. Examination of data in Table 2 indicates that few mourning doves
trapped in Colorado hatched prior to May 1, with most hatching in late May
and late June. Definite hatching peaks are difficult to ascertain from
data in Table 2, but two peaks are suggested about 20-30 days apart. Some
bias is certainly present in Table 2 as trapping effort and success and
distribution of adults and immatures were not uniform. Progeny of latenesting doves are not represented in trap samples since all dove trapping
ceases by August 20.

�-119-

Table 1. Sex and age distribution of mourning doves banded by area in
Colorado, 1971.
Adult Male
No.
Banded Percent

Area

Adult Female
No.
Banded Percent

Immature
No.
Banded Percent

Total

Eastern Colorado
Fort Morgan

27

5.1

21

4.0

480

90.9

528

Bonny Reservoir

8

10.8

1

1.4

65

87.8

74

Subtotal

35

5.8

22

3.7

545

90.5

602

Fort Garland

140

21.5

57

8.8

454

69.7

651

Monte Vista

227

37.5

155

25.6

223

36.9

605

Denver

78

15.0

121

23.3

320

61. 7

519

Fort Collins

172

33.9

212

41. 7

124

24.4

508

Fort Carson

160

37.5

132

30.9

135

31.6

427 1./

Vineland

95

76

57.0

398

5

19.1
41. 7

227

Missouri Park

23.9
41. 7

2

16.6

12

LaVeta

7

70.0

3

0

7

77.8

1

1

0.0
11.1

10

Evergreen

30.0
11.1

Woodland Park

1

12.5

7

87.5

0

0.0

8

Subtotal

892

28.3

769

24.4

1,486

47.2

3,147

Durango

252

50.1

180

35.8

71

14.1

503 5:../

Craig-Hayden

115

28.8

73

18.2

212

53.0

400

Unaweep

125

40.2

66

21.2

120

38.6

311

Meeker

135

54.6

77

31.2

35

14.2

247

Stoner

11

73.3

3

20.0

1

6.7

15

638

43.2

399

27.0

439

29.7

1,476

1,565

29.9

1,190

22.8

2,470

47.3

5,225

Eastern Slope

5

9

Western SloEe

Subtotal
Total All Areas

1/ Total does not include 7 AHY-U's.
2/

Total does not include 1 AHY-U.

�Table 2.

Estimated hatching dates for wild-trapped
Prior
to
May 1

1-7

8-14

Ma:z:
15-21 22-28

Number
Hatching

3

20

57

155

Percent
of
Total

0.1

0.8

2.3

6.3

mourning doves, 1971.

29-June 4

5-11

June
12-18

19-25

26-Ju1y 2

3-9

Ju1:z:
10-16

17-23

24-30

337

281

313

292

286

308

202

193

20

0

13.7

11.4

12.7

11.8

11.6

12.5

8.2

7.8

0.8

0

I
•....

N

0
I

�-121-

Harvest
No effort was made to collect wings from hunter-harvested doves throughout Colorado during the 1971 hunting season. Wings from 310 doves harvested between September 1 and September 11 from southeast Colorado (272),
southcentral (17), and northwest Colorado (21) were available for analysis.
Of this total, only 24 were from adults, while 286 were from immatures,
giving a young-to-old ratio of 11.9:1. This is approximately double the
5.2:1 ratio obtained from a sample of 229 birds checked in 1970 (Braun
1971). Obviously, the young-to-old ratio in this small sample does not
relate to production but rather is a function of flocking and use patterns
of the birds. Eighty-five percent of the immatures examined had replaced
five or fewer juvenile primaries, indicating they were less than 63 days
of age. This supports data collected in previous years of the study which
indicated that most early hatched young (May-June) migrated from Colorado
prior to September 1. Molt data from harvested birds also indicate that
substantial production occurs after July 23 (Table 2) as over 30 percent
(33.9) of the birds shot and examined in early September hatched after
this date. These young may represent progeny of a third nesting attempt.
Harvest data obtained through use of a mail survey of hunters after the
hunting season indicated that 22,033 hunters harvested 298.767 doves in
Colorado in 1971 (Tully et ale 1972). Thirty-six band recoveries (dead)
were received in 1971 that had been banded by Division personnel. One
of these birds had been initially banded in 1967, one was banded in 1968,
nine in 1969, eight in 1970. while 17 had been banded in 1971. Apparent
hunting mortality continues to be quite low (less than 3 percent). Of
the 36 bands reported, 19 (52.8 percent) were recovered in Colorado, 7 in
Mexico, 3 each in Arizona and Texas, and 2 each in California and New
Mexico. All but four were shot recoveries.

LITERATURE CITED
Allen. J. M. 1963. Primary feather molt rate of wild immature doves in
Indiana. Ind. Dept. Conserv., Game Res. Sect. Circ. No.4.
Indianapolis.

4 p.

Braun, C. E. 1970. Mourning dove trapping and banding. Colo. Div. Game,
Fish and Parks. Game Res. Rept •• Fed. Aid. Proj. W-88-R. Oct. pp. 143149.
1971. Mourning dove trapping and banding. Colo. Div. Game, Fish
and Parks, Game Res. Rept., Fed. Aid. Proj. W-88-R. Oct. Pp.20l-207.
Tully, R. J •• H. Riffel. and H. D. Funk. 1972. Colorado small game hunter
harvest surveys -- 1971 and 1972. Colo. Div. of Wildl. Denver. In
press.

Prepared by

_--=eg~U-_~f:.:...
-'-~...w:..~~_
Clait E. Braun
Wildlife Researcher

��October,

-123-

JOB PROGRESS

S ta te of
Project
Work

REPORT

-:::.Co:::.L:::::.o~RA~D:::.:O:::.._
_
No.

Plan No.

Job Title
Period

1972

Covered:

W-88-R-17

Migratory

4

Bird Investigations

4

Job No.
Band-tailed

Pigeon

Investi~ations

April 1, 1971 through March 3l~ 1972

Personnel:
Ken Baer and Jack R8ndall, Bureau of Sport Fisheries and Wildlife;
personnel of the U.S. Forest Service; John Arthur, Del Benson, Clait Braun,
Herb Browning, Frank Cordova, Les Denton, Howard Funk, Richard Hopper,
J. Edward Kautz, Bill Kent, Wilbur Ladd, Ann Leckler, Tom Lynch, Richard
McDonald, Brett Petersen, Charles Roberts, Robert Rouse, Marvin Smith, Marie
Vendeville, Ken Wagner, Pat Waters, J. Allen White, Mike Zgainer, Wildlife
Conservation Officers and other personnel of the Colorado Division of Wildlife.

ABSTRACT
Investigations of band-tailed pigeons (Columba fasciata) initiated in Colorado
in 1969 in conjunction with the Four Corners States Cooperative Study were
continued in 1971. Major emphasis was placed on ascertaining relative abundance,
trapping and banding samples of pigeons throughout the State, and conducting
an experimental hunting season.
Changes in abundance from 1969 and 1970 were
detected in southwest Colorado with numbers being down at least by half in
1971. Pigeons were trapped at 15 different locations in 1971 with 4,006 birds
(3,718 adults and 288 immatures) being newly banded.
Over eight hundred (801)
pigeons were retrapped in 1971, including 14 banded outside of Colorado.
During
the experimental hunting season, 524 permits were issued with 344 individuals
actually hunting.
Two hundred and one hunters were successful, each harvesting
an average of 7.6 pigeons.
Total estimated harvest (including crippling loss)
projected from the questionnaire survey (91.0 percent response) was 1,723 birds.
Crippling loss approximated 11 percent of the birds shot and retrieved.
Wings were received from 154 successful hunters.
Immature and sub-adult pigeons
comprised 32 percent of the 1,113 wings received.
Field checks of 428 (324 adults,
104 immatures) hunter-killed birds indicated that 28.7 percent of the adults
were still involved with nest activities.
Recoveries were received from 157
pigeons banded in Colorado.
The majority of these (120) were recovered in
Colorado, with 25 being recovered in Mexico and 12 in New Mexico.
At least
9.3 percent of the birds harvested during the experimental season in Colorado
were banded.
Calculated hatching dates for 617 immature pigeons ranged from
mid-April to early September.
Helminth infections continue to be low (18.3
percent) with females being more frequently parasitized than males.

��-125-

BAND-TAILED PIGEON INVESTIGATIONS
C1ait E. Braun

Intensive ecological studies of wild band-tailed pigeons which nest and
reside in Colorado from late April to early November, initiated in 1969,
continued in 1971. Investigations in Colorado represent a portion of the
regionwide Four Corners Cooperative Band-tailed Pigeon Investigation which
was initiated in Arizona in 19670 Data presented in this report are those
collected in 1971, the third year of this continuing investigation.

P. S. OBJECTIVE
To investigate life history and ecological facts pertinent to band-tailed
pigeons in Colorado as follows: range; relative numbers; relation between
numbers of birds present by area and such physical factors as weather, food,
vegetative cover, elevation, and salt; breeding and nesting chronology;
migration patterns and chronology; harvest patterns; and mortality rates.
SEGMENT OBJECTIVES
1.

To investigate distribution and relative numbers of band-tailed pigeons
in Colorado by area.

2.

To investigate the correlation between abundance and distribution of
bandtails and physical features of the environment.

3.

To trap, band, age, and sex as many birds as possible in selected
areas of Colorado to investigate migration patterns and chronology
of Colorado bandtails throughout their range.

4.

To conduct an experimental hunting season, gather harvest data, and
evaluate results.

METHODS AND MATERIALS
Most methods and materials used in 1971 were identical to those used in 1969
and 1970 (Braun 1970; 1971). Reporting procedures for field personnel were
standardized in all regions with all monthly reports being sent to the
regional biologists for tabulating and forwarding to Fort Collins. Trapping
methodology in 1971 was that used in 1970 (Braun 1971). Evaluation of the
experimental hunting season and disposition of collected birds were the same
as in 1970 and described by Braun (1971).

�-126-

DESCRIPTION OF TRAPPING AREAS
In 1971, pigeons were trapped at seven locations also utilized in 1969
or 1970. These locations were near Stoner, Bayfield, Del Norte, Evergreen,
Longmont, Salida, and Monte Vista, and have been previously described
(Braun 1970; 1971). In addition to these sites, pigeons were trapped near
Unaweep Divide, LaVeta, Fort Garland, Aspen-Basalt area, Minturn, Manitou
Springs, Woodland Park, and Be~lah. These sites were typically old grainfields (LaVeta, Fort Garland, Beulah), livestock feeding areas (Unaweep
Divide, Aspen-Basalt area, Minturn), or in residential areas where residents
fed a variety of birds (Manitou Springs, Woodland Park). As in previous
years, all trap sites were immediately adjacent to readily accessible trees
or power lines used for perches. Trees used were pri~ari1y ponderosa pine
(Pinus ponderosa), Gambe1's oak (Quercus gambe11i), Pinyon pine (Pinus
edu1is), cottonwoods (Populus spp.), and Engelmann spruce (Picea engelmannii).
It was difficult to ascertain feeding preferences as about equal numbers of
pigeons were trapped in fields where they were feeding on waste barley
(Hordium vulgare) and wheat (Triticum aestivum).
RESULTS AND DISCUSSION
Distribution and Abundance
The distribution of band-tailed pigeons in Colorado in 1971 was essentially
that reported in 1969 and 1970 (Braun 1970; 1971). No real differences were
detected in distribution between any of the three years studied. Pigeons
continue to be absent (not observed) in extreme northwestern Colorado, North
Park, South Park, and in the Gunnison Basin. The lack of observations in
all of these areas except northwestern Colorado may be related to the lack
of grain production and the scarcity of livestock feeding operations utilizing
grain. Oak is absent from all of these areas except northwestern Colorado.
In 1971, 520 different observations of pigeons were reported from April into
November. Number of reported sightings by month were: April - 18; May - 130;
June - 73; July - 95; August - 118; September - 77; October - 5; and November 4. These data indicate that in 1971 pigeons started arriving in Colorado in
late April and started migration south in September, with the peak of southward migration occurring in September. Only scattered observations of small
groups of pigeons, primarily at feeders in residential areas, were reported
in October and November.
Data concerning observations of band-tail occurrence by habitat type were
available for 472 different sightings. Of these sightings, 157 (33.3 percent) were associated with cropland, primarily grainfields. One hundred and
twenty sightings (25.4 percent) were in areas dominated by oak, 58 (12.3 percent) were from along stream courses dominated primarily by cottonwoods, 47
(10.0 percent) were in mixed coniferous areas (primarily spruce-fir-aspen),
61 (12.9 percent) were in areas dominated primarily by ponderosa pine, while
the remaining observations were scattered in orchards (7), pinyon pine-juniper
(15), open rangeland (6), and in the alpine zone (1). Observations are undoubtedly influenced by access and distribution of observers, but reported
observations probably give an adequate picture of those areas in which pigeons
feed.

�-127-

Few data are available concerning abundance of pigeons in Colorado. From
observer reports it would appear that numbers in 1971 were similar to
those in 1969 and 1970. This was not true as numbers of pigeons in
central and southwestern Colorado were substantially less than in 1969
or 1970. In the San Juan Basin numbers appeared to be down by at least
one-half to two-thirds. This is illustrated by the fact that only 168
pigeons were banded in this area in contrast to 1,228 in 1970. This is
a decrease of 86.3 percent. Reasons for the apparent decrease are not
fully known, but may include poor food conditions compounded by extensive
drought extending into New Mexico and Mexico. Trapping effort was
essentially the same in this area in both 1970 and 1971. Abundance of
pigeons continues to be highest in areas where mast producing trees and
shrubs are abundant.

Banding
Banding and trapping activities were comparable to that of 1970. Trapping
was conducted at 15 sites including three of the five 1969 trap sites and
seven of the thirteen 1970 sites. In all, 4006 pigeons were newly banded
(Table 1).
While numbers of immatures banded were exceedingly low in 1971 (288), it is
apparent that numbers in trap samples increased from June (2), to July (72),
and August (148) and decreased in September (66). In 1971, few pigeons
hatched prior to June, with the peak of fledging occurring in late July and
August.
Sex of most adult pigeons banded in 1971 was determined from external plumage
characters. Of the 3,718 adults (including subadults) newly banded, 48.1
percent (1,789) were males and 51.9 percent (1,929) were females. Reasons
for the slight disparity are not fully understood but may be related to time
of day that trapping was conducted. Time of day of trapping was recorded
for all but three pigeons banded in 1971. These data are presented in Table
2.
It is obvious from data in Table 2 that sex ratios of birds available for
trapping changed during the daylight hours. This change is undoubtedly due
to differences between males and females in incubation and squab brooding
and feeding periods. Also, it is apparent from Table 2 that more than half
of the trapping effort or at least trap success occurred after 1000 MDT.
This is probably the major reason that more females than males were banded
in 1971.

�Table 1.

Number of band-tailed pigeons banded by area in Colorado, 1971.
Adults
No.
Banded
Percent

lnunatures
No.
Banded
Percent

Totals

Estimated No. of Birds
Using Trapping Sites

0.0
2.1

66
97

100+
200+

0
0
1

0.0
0.0
2.0

10
5
51

40+
10+
100+

0
0
1
48

0.0
0.0
5.9
13.8

404
175
17
349

600+
600+
50+
500+

Area

Dates

Unaweep

May 9-20
June 16-27

66
95

100.0
97.9

0
2

Monte Vista

May 12-13
July 12-16
August 5

10
5
50

100.0
100.0
98.0

LaVeta

May 20-25
June 11-12
July 3
August 18-22

404
175
16
301

100.0
100.0
94.1
86.2

....
I

N

00
I

Del Norte

May 27
June 9-10
August 6-29

52
112
72

100.0
100.0
72.7

0
0
27

0.0
0.0
27.3

52
112
99

100+
200+
200+

Longmont

June 4
July 11
August 7 &amp; 20

119
123
16

100.0
96.1
94.1

0
5
1

0.0
3.9
5.9

119
128
17

200+
200+
50+

Salida

June 2-16
July 9-14

254
70

100.0
97.2

0
2

0.0
2.8

254
72

400+
100+

Bayfield

June 8-Ju1y 7

153

100.0

0

0.0

153

200+

Fort Garland

June 30-Ju1y 7
August 4-17

424
220

99.5
90.5

4
23

.5
9.5

426
243

600+
300+

Aspen-Basalt

June 28-29
July 12-21

27
105

100.0
91. 3

0
10

0.0
8.7

27
115

50+
200+

-------------------------------------------------------------------------------------------------------

�Table 1.

Number of band-tailed pigeons banded by area in Colorado, 1971 (continued).

Area

Dates

Adults
No.
Banded
Percent

Minturn

July 18-23

173

94.0

11

Stoner

July 25

10

66.7

Evergreen

July 14-27
September 28

174
13

Woodland Park

July 21-Aug. 4
September 16

Immatures
No.
Banded
Percent

Totals

Estimated No. of Birds
Using Trapping Sites

6.0

184

250+

5

33.3

15

50+

91.1
40.6

17
19

8.9
59.4

191
32

200+
50+

107
54

85.6
79.4

18
14

14.4
20.6

125
68

150+
100+

Manitou Springs Aug. 5-17
September 13-15

171
92

79.2
73.6

45
33

20.8
26.4

216
125

300+
200+

Beulah

57

96.6

2

3.4

59

100+

3,718

92.8

288

7.2

4,006

6,400+

I
t-O

N
\0
I

Totals

August 19

�-130-

Table 2.

Time of day of band-tailed pigeon trapping, 1971. !I

Time (MDT)

Males

Females

Percent
of Total
Adults

0600-0959

67.4

32.6

35.B

29.7

1000-1559

36.6

63.4

57.B

59.B

1600-

45.0

55.5

6.4

10.5

Adults

Percent of
Total
Juveniles

l/All values are in percent.

A total of BOI previously banded pigeons were retrapped in 1971. All but
14 of these had been banded in Colorado, with 377 of this total being recaptured out of the original degree block of banding or were banded in
earlier years. Four hundred and nine were retrapped in the same degree
block and same year as originally banded, while one, banded by a private
bander in 1967, also was recaptured in the degree block of original banding. Of the 14 out-of-state-bandings retrapped in Colorado, 11 were
originally banded in New Mexico, 2 in Utah, while 1 was originally banded
in Arizona. Data on all recaptures through 31 December 1971 were analyzed
in detail and were reported by Braun (1972).

Experimental Hunting Season
1971 was the second year of the 3-year experimental harvest program initiated
in 1970. Hunting was allowed from September 4 through 26, with a daily bag
limit of five birds and a possession limit of 10. During the experimental
season only the southern portion of the State was open (Appendix). Hunting
was by free permit obtainable at seven Division offices with no limit on
the total number of permits to be issued. In 1971, 524 permits were issued,
an increase of 160 (44.0 percent) over 1970.
Hunter Questionnaire Survey
The experimental hunting season for bandtails in 1971 closed at sunset on
September 26. On September 27, letters containing a self-addressed, postagepaid return card were sent to all permittees (Appendix). Only two (.4 percent) questionnaires were returned for lack of sufficient address. On
October 11, a followup letter was sent to the 130 permittees (excluding the
two permittees for which insufficient addresses were available) who had
failed to respond to the initial letter. Results of the questionnaire survey are presented in Table 3. Data for the 47 non-respondents were calculated
using average values of the respondents to the fol10wup survey.

�Table 3.

Experimental band-tailed

pigeon season, harvest statistics, 1971. -1/
Response
First
Letter

Response
Second
Letter

Sum
First and
Second Letters

Number of Permittees Responding

392 (74.8)

85 (16.2)

477 (91.0)

Number of Permittees Hunting

259 (66.1)

55 (64.7)

314 (65.8)

344 (65.6)

Number of Permittees Not Hunting

133 (33.9)

30 (35.3)

163 (34.2)

180 (34.4)

Number of Successful Hunters

152 (58.7)

32 (58.2)

184 (58.6)

201 (58.4)

Number of Hunter Days

630

143

773

851

Days Hunted Per Hunter

2.4

Number of Pigeons Bagged

1,214

2.6
209

2.5
1,423

Projected
All (524) Permittees

2.5
1,537

Pigeons Per Successful Hunter

8.0

6.5

7.7

7.6

Pigeons Per Hunter

4.7

3.8

4.5

4.5

Number of Pigeons Crippled and
Lost

138

Pigeons Crippled and Lost Per
Hunter
Total Harvest (Bagged + Crippled
and Lost)
Percent Crippling Loss

.5
1,352
10.2

II
Values in parentheses

31

are percentages.

169
.6

240
12.9

186
.5

1,592
10.6

.5
1,723
10.8

I

t-'
t-'I
w

�-132-

Interest in pigeon hunting increased substantially (44 percent) in 1971
over 1970. Along with the increase in hunter interest, increases were
also recorded in hunter success (59 percent versus 42 percent) and in
total harvest (1,723 versus 541). However, as in 1970, a substantial
number of permittees did not hunt (34.4 percent). Possibly some hunters
do not know how to hunt pigeons or they just obtain a permit because of
the novelty or on the chance they may see pigeons while hunting other
game animals.
Average number of days hunted per hunter increased slightly in 1971 from
1970 (2.1 to 2.5) as did the nUmber of pigeons bagged per successful
hunter (5.9 to 7.6). Percent crippling loss (total number of pigeons
reported lost divided by total harvest (bagged + crippled and lost» was
10.8 in 1971, substantially less than the 15.3 percent reported in 1970.
Undoubtedly reported crippling loss is minimal and actual crippling loss
may represent as much as 20 percent of the birds actually retrieved.
The questionnaire was devised in order to ascertain the proportion of
the harvest that occurred in given time periods. It was found that 29.3
percent of the total harvest occurred on the opening weekend, 17.4 percent on the second weekend, 12.3 percent on the third weekend, and only
7.6 percent on the final weekend. Some hunting occurred during the week
as 13.1 percent of the total harvest occurred during the first week, 13.9
percent during the second week, but only 6.4 percent during the third
week. Thus harvest was well spaced during the 1971 season, suggesting
that hunting remained productive throughout the season.
Pigeon hunters were clumped in 1971 with three areas, Fort Carson (22.7
percent), Durango (15.9 percent), and LaVeta (12.6 percent) receiving more
than 50 percent of the total pressure. In all, 309 hunters reported
hunting at 25 different locations. Other hunter concentrations occurred
near Monte Vista (7.8 percent), Arboles (7.1 percent), Salida (6.8 percent),
and Dolores-Mancos (6.1 percent). None of the remaining 18 locations received more than 4 percent of the total hunter pressure. On a regional
basis, 158 of the 309 hunters (51.1 percent) hunted in southeastern
Colorado, while 137 (44.4 percent) hunted in southwestern Colorado, and
only 14 (4.5 percent) hunted in northwestern Colorado. There were no
significant differences in hunter success between regions in 1971 as 58.2
percent of the hunters in the southeast were successful, 57.1 percent in
the northwest, while 56.9 percent were successful in the southwest region.
Many written comments were received from hunters with the majority of the
comments relating to a lack of pigeons, poor acorn crops, or to bad weather
early in the season. All of these comments had some validity as many
pigeons had left areas where they had been all summer, the acorn crop was
extremely spotty, and some areas of the State did have wet, cool weather
during the opening weekend of the season.
Wing Survey
Packets containing five wing envelopes were issued to each pigeon permit
holder in 1971. Envelopes containing 1,113 wings were received from 154

�-133-

hunters. Additional data were received concerning hours hunted (834.5),
birds crippled and lost (118), location of kill and banded birds harvested (103). Major harvest areas were near LaVeta (18.1 percent),
Fort Carson (16.4 percent), Arboles-Pagosa Springs (13.4 percent), Monte
Vista (12.1 percent), and Dolores-Mancos (10.1 percent). These data are
somewhat different than the distribution of hunters obtained from the
questionnaire survey. Thus LaVeta with 18 percent of the total harvest
had only 13 percent of the total hunters, while Fort Carson with 23 percent of the total hunters had only 16 percent of the total kill. Hunting
was also better than average near Arboles (13 percent of the harvest and
7 percent of the hunters), Monte Vista (12 percent of the harvest and 8
percent of the hunters), and Dolores-Mancos (10 percent of the harvest
and 6 percent of the hunters). In contrast, Salida had 7 percent of the
hunters and only .9 percent of the harvest indicating that hunting was
extremely poor in that area in 1971. Crippling loss calculated from the
wing survey was slightly lower (9.6 percent as compared to 10.8 percent)
than that calculated from the questionnaire survey but the slight difference is not significant.
In all, 103 of the 1,113 band-tailed pigeons reported harvested on the wing
survey were banded (9.3 percent). Percent of birds harvested that were
banded varied from 19.1 (N = 183) at Fort Carson, 11.5 (N = 52) at Dolores,
9.7 (N = 196) at LaVeta, 7.4 (N = 135) at Monte Vista, 4.6 (N = 153) at
Durango, to 2.1 (N = 149) at Arboles-Pagosa Springs. Bands were reported
from seven other locations but sample sizes of wings received were too
small (less than 40) for meaningful comparisons. None of the banded birds
shot were harvested at trap sites. It is therefore possible that for the
areas from which substantial numbers of wings were available, the above
percentages of banded birds in the harvest may give an adequate picture
of the percent of the populations that were banded. It is believed that
the 9.3 percent banded birds in the harvest is representative and that
possibly 10 percent of the band-tailed pigeons present in Colorado in
fall, 1971, were banded.
Comparable data were available from both hunter questionnaire and wing
surveys concerning numbers of birds bagged and those crippled and lost by
136 individual hunters. A reporting difference of 13.1 percent (978 in
wing surveys versus 1,126 in questionnaire survey) was found in number of
birds killed, and 12.5 percent (98 versus 112) in number of birds crippled
and lost for those hunters who sent in wings and also responded to the
questionnaire.
The difference in birds harvested and retrieved could be
related to the fact that hunters only received five wing envelopes each
and could thus send in only 25 wings. Some hunters reported killing as
many as 69 birds. The difference in crippling loss can also be attributed
to the number of wing envelopes each hunter received as the number of pigeons
reported lost on wing envelopes was less than on the questionnaire survey.
Since both percentages are similar, (13.1 and 12.5), it is assumed that
reporting differences were slight between the questionnaire and wing surveys.
All wings received were measured (carpal and all 10 primaries) and classified as immature, subadult, and adult in order to derive age structure of
the harvest. Wings were received from 779 adults (68.0 percent), 122 subadults (10.6 percent), and 245 immatures (21.4 percent). Percent of adults

�-134-

in the harvest was similar to that recorded in 1970 (68.0 versus 67.8
percent), while percent immatures decreased from 1970 levels (21.4 versus
26.6). If the subadult (birds less than 1 year of age but having molted
past P5) age class is added to the immature class, data for all birds less
than 1 year of age are comparable between the two years (32.0 in 1971 and
32.2 percent in 1970). Age ratios in the harvest were different than
expected from trapping results. It was expected that substantially fewer
immatures (percent) would be harvested in 1971 than in 1970 as all indications were that production was substantially less in 1971 than in 1970.
Reasons why the expected decrease did not occur are not known, but may be
related to differential migration and differences in vulnerability.
It
is also possible that production was higher than that predicted from
trapping results.

Hunter Field Checks
During the 1971 experimental season emphasis was again placed on contacting
pigeon hunters in the field in order to examine harvested birds for crop
gland activity. Crops of all adult pigeons checked were classified as being
active (crop gland with curds), stimulated (gland apparent but no curds), or
inactive. In addition, sex by gonadal inspection was obtained for as many
birds as possible. In all, 428 birds were checked with 104 being immatures
(24.3 percent), 320 birds being adults (74.8 percent), and 4 being subadults
(0.9 percent). Of the adults, 13.9 percent were classified as having active
crop glands while 14.8 percent were classified as being stimulated. Percent of crops showing glandular activity (active + stimulated) was significantly less in 1971 than in 1970 (28.7 versus 52.3) even though the hunting
season was 1 week earlier. This suggests that either nesting was earlier
in 1971 than in 1970, or that fewer pigeons had second or late nests.
None of the field data collected support the theory that nesting was earlier
in 1971. These data thus support the theory that only one nesting attempt
was most common in 1971.
Data on sex were available for 300 adults. Of this number 58.0 percent
were males and 42.0 percent were females. Of the males, 33.9 percent
exhibited some glandular development while only 23.3 percent of the females
showed development of the crop gland. While samples of adults examined by
area were not high, there were some interesting differences between areas
in crop gland development. At Durango, 55.5 percent of the adults exhibited
gland development (N = 27), it was 44.9 percent at Monte Vista (N = 69),
20.0 percent at Arboles (N = 40), 19.7 percent at LaVeta (N = 71) and only
9.6 percent at Fort Carson (N = 73).

Mortality
Recoveries of 157 band-tailed pigeons initially trapped and banded in
Colorado were reported in 1971. Eleven of these were initially banded in
1969, 45 were banded in 1970, while 101 were initially banded in 1971. All
but six were shot recoveries. Of the 151 shot recoveries, 117 were shot in
Colorado during the 1971 experimental season, 12 were shot in New Mexico

�-135-

during their 1971 experimental season, while 22 were shot in Mexico,
principally in the States of Durango (10) and Sinaloa (7)'. Four pigeons,
banded outside of Colorado, were reported shot in 1971 in Colorado. Three
of these were from New Mexico and one was from Utah.
In a preliminary analysis of all bandings and recoveries from 1969 through
31 December 1971, Braun (1972) found that the reported hunting mortality of
Colorado bandtails was less than 3 percent per year and that most hunting
mortality was concentrated in breeding (77.6 percent) and wintering areas
(14.8 percent).

Breeding Phenology
Data on breeding phenology were obtained through crop and gonadal inspection
of birds accidently lost in trapping operations and those systematically
collected. Additional data were collected from 324 hunter-killed adult
pigeons examined from September 4 through 26. Excluding the 324 hunterkilled birds, 177 birds were available for detailed examination. Of this
number, 150 were adults (68 males and 82 females) while the remaining 27
were immatures. The sample included birds taken from 7 May through 28
September with the following monthly totals: May - 13, June - 19, July 22, August - 35, and September - 88. Five additional birds were available
that died in the aviary during the October-March period. Data from the
above collected birds are reported elsewhere (Braun and Benson 1972).
Wing Measurement

and Molt Data

Data concerning length of primaries, rectrices, total body length, carpal
length, and bill length were obtained from all pigeons collected in 1970.
Once adequate samples are available, these data will be analyzed to examine
differences between age classes of both male and female pigeons. Prior to
dissection of collected birds, plumage characters of each adult specimen
were examined and recorded. Each bird was then classified as either male
or female. Upon dissection, gonads were examined and actual sex of each
bird was compared with the classification based on plumage appearance.
Comparative data were available for 126 adults (56 males and 70 females).
Of the 56 adult males compared, two (3.6 percent) were classified incorrectly,
while one (1.4 percent) of 70 females was classified as a male from plumage
examination. Total error in classifying pigeons to sex from plumage examination alone was 2.4 percent. These percentages are similar to those reported
by Braun (1970; 1971) for samples of 119 and 117 adults.
Information concerning primary molt was available for 617 immatures (284
banded and released, 234 from the hunter wing survey, 72 trapped and placed
in the aviary, and 27 collected). Of the 617 immatures, 329 had not molted
primary 1, 133 were molting PI, 75 were molting P2, 41 were molting P3, 33
were molting P4, and 6 were molting P5. Hatching dates for these birds
were estimated using unpublished data from White (1972). Estimated hatching
dates for all immatures for which data are available are presented in Table 4.

�-136-

Table 4.

Estimated hatching dates for wild band-tailed pigeons, 1971.

Time Period

Number Hatching

Time Period

Number Hatching

April 12-21

2

July 1-10

70

April 22-May 1

4

July 11-20

82

May 2-11

12

July 21-30

58

May 12-21

26

July 31- Aug. 9

47

May 22-31

35

August 10-19

55

June 1-10

49

August 20-29

37

June 11-20

69

Aug. 30-Sept. 8

2

June 21-30

69

As in earlier years (Braun 1970; 1971), hatching and fledging of band-tailed
pigeons were spread over a wide time period. It is questionable if pigeons
estimated to have hatched in April were progeny of nesting attempts in
Colorado. If the few pigeons arriving in early April nested as soon as they
arrived, it is possible that the few young estimated to have hatched could
have been produced in Colorado. Progeny of late nests (after September 8)
have little chance of being shot or trapped and thus are not represented in
Table 4.

Parasitic Load
Information on helminths was available for 120 band-tailed pigeons (106
adults and 14 irnmatures) necropsied. Twenty-two (all adults) of the 120
birds examined had helminth infections. The infection rate of 18.3 percent,
while higher than that found in 1969 (13.2 percent) and 1970 (10.4 percent),
is low and none of the infected birds was heavily parasitized. As in other
years, more adult females (26.2 percent) harbored helminths than did males
(13.3 percent). Tapeworms were found in 16 birds while 7 birds had nematodes
(one bird had both tapeworms and nematodes). Worms were found in pigeons
collected as early as 4 June and as late as 26 September. No deleterious
effects of the observed parasitism were noted. All worms were preserved
for later identification.

�-137-

LITERATURE

CITED

Braun, C. E. 1970. Band-tailed pigeon investigations.
Colo. Div. Game,
Fish and Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct.
pp. 151-171.
1971. Band-tailed pigeon investigations.
Colo. Div. Game, Fish
and Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. pp. 209236.
1972. Movements and hunting mortality of Colorado band-tailed
pigeons. Trans. No. Am. Wildl. &amp; Nat. Resour. Conf. 37:326-334.
______ , and D. E. Benson. 1972. Band-tailed pigeon nesting and breeding
phenology investigations.
Colo. Div. Wildl., Game Res. Rept., Fed.
Aid Proj. W-88-R. Oct. pp. In press.
White, J. A. 1972. Plumage studies of band-tailed pigeons. Colo. Div.
Wildl., Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. pp. In press.

Prepared by

f!t;/ c. ~~
Clait E. Braun
Wildlife Researcher

oaf'

�APPENDIX

EXPERIMENTAL BAND·TAILED PIGEON SEASON .1971

GENERAL OPEN SEASON
SEPTEMBER 4 through SEPTEMBER 26, 1971, in all lands west of U. S. Interstate
Highway No. 25 from the New Mexico state line to its intersection with U. S.
Highway No. 24 at Colorado Springs; south and west of U. S. Highway No. 24 to
its intersection with U. S. Interstate Highway No. 70 at Dowd Junction; and
south of U. S. Interstate Highway No. 70 to the Utah state line.
HUNTING HOURS:

Sunrise to sunset

DAILY BAG AND POSSESSION LIMITS
DAILY BAG LIMIT - five (5).
POSSESSION LIMIT - ten (10).
SPECIAL PERMIT REQUIRED
All hunters participating in this season must first obtain and have in his
possession while hunting band-tailed pigeons a special hunting permit.
Permits unlimited in number and issued free of charge will be available from
Division of Game, Fish and Parks offices at Denver, Fort Collins, Colorado
Springs, Grand Junction, Durango, Glenwood Springs, and Montrose. Permits
will be available starting AUGUST 16 and will be issued upon request
throughout the season.

�-139STATE OF COLORADO

.John A. Love, Govarnor

DEPARTMENT OF NATURAL

RESOURCES

DIVISION OF GAME, FISH AND PARKS
Harry

R. Woodward,

6060 Broadway

Director

Denver, Colorado 80216 • 825-1192

September 27, 1971

Dear Band-tailed

DEPARTMENT OF NATURAL
RESOURCES

r. W. Ten Eyck,
Executive

Harry Combs, Chairman
Dean Sullie, Vice Chairman
William W. Robinson, Secretary
Wilton W. Cogswell, Jr., Member
R. Withers Cool, Member
Charles A. Gebauer, Member
Oresl Gerbaz, M"mbcr
Lonnie Pippin, Member
LeRoy Robson, Member
Ford SIrong, Member

Pigeon Hunter:

By obtaining a permit for the Colorado experimental band-tailed
pigeon hunting season from September 4 through September 26, 1971,
you have indicated an interest in pigeon hunting and the future
of the sport. We respectfully ask your assistance in evaluating
this experimental season by completing the enclosed self-addressed
questionnaire and returning it as soon as possible.
Information
you supply will be instrumental in determining the potential and
feasibility of this type of season in the future.
Please read the questionnaire carefully and fill in all requested
information completely and accurately.
If you did not hunt during
the experimental season, we ask that you complete Question No. 1
and return the questionnaire because negative information is also
important in our evaluation.
If you did participate, please report
only on your experimental season hunting activities and not those
of your friends or other members of your family.
Thank you for your assistance.

CEB:jc
Enclosure

Director

GAME, fiSH AND PARKS
COMMISSION

�-140-

BAND-TAILED PIGEON HUNTER REPORT
1. Did you hunt band-tailed pigeons during the 1971 season?
Yes
No,
_
2. County or area most frequently hunted,
3. How many days did you hunt during the pigeon season?

_

c:::J

4. How many pigeons did you bag during the entire season? I~_---I
5. How many pigeons did you knock down within sight but
could not retrieve?
6. Please indicate in the appropriate boxes below the nu~f
pigeons killed each time period you hunted. Put an "0" for
the periods you did ~
get any birds. Leave periods you did
not hunt blank.
First
First
Second
Second
Third
Third
Fourth
Weekend
Week
Weekend
Week
Weekend
Week
Weekend
4 &amp;5
6-10
11 &amp; 12
13-17
18 &amp; 19
20-24
25 &amp; 26

r---l

c:J c:J c:J

Cl

CJ

I

I

Paul Revere

Band-tailed Pigeon Hunter Survey
Wildlife Research C~nter
P. O. Box 567
Fort Collins, Colorado 80521

I

I

�-141STATE OF COLORADO

DEPARTMENT OF NATUkAl
RESOURCES

.John A. Love, Govel'nor

DEPARTMENT OF NATURAL RESOURCES

DIVISION OF GAME, FISH AND PARKS
Harry

R. Woodwal'd,

Dlrectol'

6060

Broadway

Denver, Colorado 80216 • 825-1192

Wildlife Research Center
P. O. Box 567
Fort Collins, Colorado 80521
October

Dear Band-tailed

T. W. Ten Evck,
Executive Director

GAME, FISH AND PARKS
COMMISSION
Harry Combs, Chairman
Dean Suttle, Vice Chairman
William W. Robinson, Secretary
Wilton W. Cog~well, Jr., Membor
R. Withers Cool, Member
Charles A. Gebau~r, Member
O,elt G.,baz,
Member
Lonnie Pippin, Mambar
LeRov Robson, Member
Fo,d Strong, Member

11, 1971

Pigeon Hunter:

We recently sent you a questionnaire pertaining to your hunting
activities during the experimental band-tailed pigeon hunting
season in southern Colorado from Septe~er
4 through September
26, 1971. So far we have not received your reply. Please assist
us by completing the questionnaire and returning it to us as
quickly as possible.
Information you supply, ~
though you did
~
participate, would enable us to fully evaluate the special
season and would assist us in determining the potential and
feasibility of such a season in the future.
Thank you!

CEB:jc

��-143-

October, 1972

JOB PROGRESS REPORT

State of

~CO~L~O~RAD~~O~

Project No.

W-88-R-17

Work Plan No.

4

_
Migratory Bird Investigations·
Job No.

5

Job Title

~B~a~n~d~-_t~a~i~l~e~d~P~i~g~e~o~n~I~n~v~e~s~t~i~g~a~t~L~·~o~ns~:
__~P~l~u~m~a~g~e~S~t~u~d~i~e~~s
_

Period Covered:

April 1, 1971 to March 31, 1972

Personnel: Clait E. Braun, Jack Corey, J. Edward Kautz, Breet Petersen,
Julie M. White, J. Allen White.

ABSTRACT
This aviary study of band-tailed pigeons (Columba fasciata) was attempted in
order to develop techniques suitable for care, maintenance and breeding of
captive bandtails; document progression of molts and subsequent plumages from
juvenile to adult age classes; and investigate the validity of plumage
characters as aids in accurately distinguishing between juvenile, sub-adult,
and adult age classes. Results indicate the bandtails are not difficult to
keep and rear if adequate housing, minimum disturbance and patience are used.
Progression of postjuvenile molt in all feather tracts were studies. It was
found that captive pigeons complete this molt at about 8 months of age.
Primaries are molted and replaced in sequence from the inner primary (PI)
to outer primary (PlO), a primary being lost and replaced about every 20 days.
Subsequently, it was ascertained that certain plumage characters are useful
in age and sex determination. Color of the head and upper breast is at
least 95 percent accurate in determining sex in bandtails over 65 days of
age. Wing characters, including primary measurements, are of little value
in determination of sex. Juvenile bandtails are eaSily distinguished by
gross plumage examination, while sub-adults can be identified by buffy tipping
in the upper wing coverts or the presence of juvenile primaries or secondaries
until they are about 8 months of age.

��-145-

PLUMAGE STUDIES OF BAND-TAILED PIGEONS
J. Allen White
In recent years, interest in many so-called ''minor''game species has
intensified, partly because of increased demands for added recreational
opportunities and partly to fill voids in present knowledge. The band-tailed
pigeon is a minor game species, even though having a wide distribution in
the western hemisphere. In order to obtain sufficient knowledge upon which
to base management programs, research on this native Columbid has intesified in the past few years. In the management of any species, estimates of
popUlation trends, annual production, and survival rates are essential. However the manager must first be able to quickly classify individual animals
into age and sex categories. Unfortunately, the available methods of age and
sex determination for bandtails were for the most part inadequate for application in game management programs. This study, involving the documentation
of moLts and subsequent plumage and time required for each change, was undertaken to further the knowledge needed to manage band-tailed pigeons. The
objectives of this study are: (1) develop techniques suitable for care,
maintenance, and breeding of wild pigeons in captivity, (2) document progression of molts and subsequent plumages from juvenile to adult age classes,
and (3) investigate the validity of plumage characters as aids in accurately
distinguishing between juvenile, subadult, and adult age classes and between
sexes.
P. S. OBJECTI VE
To investigate life history and ecological facts pertinent to band-tailed
pigeons in Colorado as follows: range; relative numbers; relation between
numbers of birds present by area and such physical factors as weather, food,
vegetative cover, elevation, and salt; breeding and nesting chronology; migration patterns and chronology; harvest patterns; and mortality rates.
SEGMENT OBJECTIVES
1.

To trap and keep in captivity band-tailed pigeons.

2.

To document timing and duration of molts and subsequent plumages of bandtailed pigeons from immature to the adult age class.

3.

To compile data and prepare progress report.

REVIEW OF LITERATURE
The band-tailed pigeon is a widely distributed game bird found in most mountainous regions of western North America, Central America, and northwestern
South America (Goodwin 1967). There are several recognized subspecies
including two which occur in the United States (A.O.U. 1957). The larger

�-146-

coastal pigeon (C. f. moni1is) winters in southern California and Baja
California, migrating north as far as southern British Columbia during warmer
seasons. The smaller interior bandtai1 (C. f. fasciata) winters in central
Mexico and seasonally migrates into Arizona,-Colorado, New Mexico, and Utah.
Fitzhugh (1970) compiled the most complete literature review of the band-tailed
pigeon to date. He foUnd that most of the available references are of
limited scientific value. However, some of the more recent works are worth
noting. These include Neff (1947), Glover (1953), Houston (1963), Mace and
Batterson (1961), Macgregor and Smith (1955), Silovsky (1969), Smith (1968),
Silovsky et al. (1968), and Wight et al. (1967), among others.
Unfortunately, few individuals have been successful in keeping and rearing
band-tailed pigeons, and no published material is available on this subject.
M. R. Blankenship of Walnut Creek, California (personal communication), is
probably the most successful person to raise bandtails, a hobby he has been
working with for nearly 20 years. Levi (1969) wrote extensively on care and
breeding of domestic pigeons (Columba livia), but makes no mention of wild.
breeds. However, much of his information on housing, feeding, and disease
prevention are applicable to keeping wild birds.
An attempt has been made to study post-juvenile primary molt in bandtails by
H. Wight (unpublished data). Levi (1969) studied the post-juvenile molt of
primaries, secondaries, and rectrices in domestic pigeons. Both of these
studies dealt with very small sample sizes.
Methods of age determination for bandtails during fall and early winter have
been suggested by Silovsky et al. (1968). This method has some obvious
limitations for use in spring and midsummer banding operations, although some
of the characters discussed are useful. Miller and Wagner (1955) detailed a
method of sex determination for Columbids using cloacal characters. Their
method is good, but proved to be slow when large numbers of birds are to be
quickly processed.

METHODS AND MATERIALS
Ten pens were used during this study to house pigeons. Six were in two banks,
three pens per bank, and each pen was 6 feet high, 6 feet wide, and 12 feet
deep. Other pens measured 6 feet high, 6 feet wide, and 6 feet deep; and 5
feet high, 8 feet wide, and 5 feet deep. A goose brooderhouse of redwood
construction measuring 18 feet long, 10 feet wide, and 9 feet high was used to
house sick birds during cold weather. All pens were provided with adequate
perches and shelter suitable for keeping pigeons.
During August and September, 76 immature band-tailed pigeons were trapped,
individually marked with yellow plastic bandettes, and placed in the aviary
for plumage studies. Four of these birds were obtained from F. John Ward,
Pebble Beach, California, including one fawn-colored (imperfect albino or
schizochromic mutation) bird. The other 72 pigeons were trapped by Colorado
Division of Wildlife personnel at Evergreen, Longmont, Manitou Springs, Fort
Garland, Monte Vista, Del Norte, and La Veta.

�-147-

All pigeons had free access to food and water with grit being available from
the fine gravel which was provided on the floor of each pen. The grain
mixtures fed to pigeons during the plumage investigations varied slightly to
ascertain any preferences. Among the grains tested were whole and cracked
dent corn (Zea mays), field peas (Pisum sativum), whole wheat (Triticum
aestivum), whole milo (Sorghum vulgare), whole barley (Hordeum vulgare), and
oats (Avena sativa). The basic food mixture fed consisted of six parts
whole milo, three parts whole corn, and one part whole wheat. Water was
provided in 3-gallon galvanized pans, while food was placed on the gravel
floor.
During October, November, and December, ten birds became sick and seven of
these subsequently died. One dead bird was sent to the Diagnostic Laboratory
at Colorado State University and the cause of death was determined to be
peritonitis. Ano,tner sick bird was taken to the Colorado State University
Small Animal Clinic for examination and treatment. The diagnosis was probable
Aspergillus fum!gatus, a fungal infection caused by damp conditions and moldy
grain. Symptoms were similar in all sick pigeons and were characterized by
a slow decline in weight following infection, until the birds became extremely
thin and weak. Shortly before death, breathing became labored and traces
of blood became apparent in the saliva. At death, hemorrhaging in the respiratory tract occurred and the lungs were filled with mixed saliva and blood.
During post-juvenile molt of primaries, each pigeon was examined at least
three times each month, with four examinations being preferred. Data concerning each bird were tape-recorded or taken by an assistant for later transcription onto individual data sheets. Data taken during examinations include
the following 29 categories:
1 through 10. Timing and replacement with measurement of each primary as
it was replaced until maximum length was attained. .In addition, the length and
width of P8, P9, and PIO were taken before and after post-juvenile molt for
comparative puposes.
11.

Primary edging

12.

Secondary wear

13.

Molt and replacement of secondaries

14. Replacement of rectrices. In addition, length and width of outer
and center rectrices were originally taken for comparative purposes.
15.

Tail covert edging

16.

Upper wing covert edging

17.

Alular edging

18.

Alular covert edging

19.

First and second greater secondary coverts edging

20.

Color of crown and upper breast

�-148-

21.

Crescent appearance

22.

Nape color

23.

Eye color

24.

Cuticle color

25.

Feathering of nares (completion of juvenile plumages)

26.

Bill color

27.

Foot and leg color

28.

Weight

29.

Other

The pigeons were disturbed as little as possible, even though they did not
generally become alarmed when being handled. All extensive cleaning of pens
was done after pigeons had been caught and placed in covered holding boxes
prior to routine examination. Feeding and watering took only a short time
each day, and was done without entering the pens. Subdued clothing colors
were essential, as bright colors, whites, and some plaids caused the birds
to become nervous, and at times, to panic. Dark greens, browns, or faded
camouflage attire was best, and resulted in the birds being much calmer,
reducing the chance of feather damage.
As the post-juvenile molt was completed, individual birds were either released,
placed in a large breeding pen, or transferred to other aviaries. Two pairs
were transferred to Dr. R. M. Stabler at Colorado College in Colorado Springs
for breeding studies. These were later released. One pair was transferred
to Mary Gilbert, a graduate student at Colorado State University for breeding
and behavior studies. One pair was transferred to Dr. H. D. Muller, Avian
Science Department, Colorado State University, for breeding studies. An
additional 13 pigeons were released from the aviary to reduce crowding.
RESULTS AND DISCUSSION
Care and Maintenance
M. R. Blankenship is probably the most successful breeder of band~tailed
pigeons in the United States, having over 20 years experience and unfailing
results in raising bandtails, as well as many other species of wild doves
and pigeons. Oddly enough, his techniques are simple. According to Blankenship (personal communication), the three most important considerations in
raising any wild pigeon are adequate facilities, providing a minimum of disturbance, and patience. Care should be kept to a minimum, set time period
each day, and should be restricted to feeding, watering, and cage cleaning.
If the keeper is present at the same time each day the pigeons will learn to
expect him and remain calm. Normally, the best time is around noon, but never

�-149in late afternoon. It is also desirable to wear the same subdued colored
clothing each time the pen is entered. Pigeons, like most birds, appear to
"recognize" their caretaker and wearing the same clothing each day causes
less disturbance.

Feeds
Levi (1969) and Blankenship agree that grain mixtures are adequate rations
for pigeons. Levi (1969) suggests a mixture of whole corn, milo, wheat, and
field peas for domestic pigeons. Blankenship (personal communication) has
fed only whole corn and mi10 for 20 years with excellent success. During this
study over 60 immature pigeons were kept on a mixed ration of whole corn,
whole milo, and wheat, with no apparent problems. It is doubtful, however,
that grain mixtures would be the ideal ration for breeding pairs or young
birds. Commercial pigeon rations and gamebird breeder rations are available
and are probably better than grain mixtures.

Housing
Another factor limiting success in raising bandtails is the use of inadequate
facilities. ~vi (1969) discusses types of housing used for domestic pigeon
breeds. The bandtai1, however, will not tolerate crowding and conditions
which are optimum for domestic birds. While it is possible to keep '~arn
pigeons" in very small cages, bandtai1s require cages at least 6 feet high, 6
feet wide, and 12 feet deep. Such a cage will hold a maximum of 10 nonbreeding birds. In all such cases, the size of the pen determines the amount
of stress and injury caused by routine care. In large pens, the birds simply
do not panic when approached.
All pens should be covered with 1-inch mesh poultry netting. Smaller mesh is
too expensive and 2-inch mesh wire will cause injury. Bandtai1s are strong
fliers and can easily reach speeds over 60 miles per hour (Fitzhugh, personal
communication). At such speeds (nearly double that of most gallinaceous
birds) 1-inch wire is bad, but 2-inch wire results in badly cut crops and dead
birds.
Several other requirements must be met in pen construction. All pens should
provide shelter from storms, wind protection, shade, privacy from outside disturbance, and adequate perches. Bandtai1s, like any other pigeon, prefer to
perch well above ground level and require a roof for protection from storms.
Wind protection can be provided with existing vegetation, snow fencing, or
burlap. Blankenship (personal communication) prefers a natural windbreak of
native plants which he maintains around each pen. Although this restricts
cleaning activities, it looks good (reducing complaints from neighbors), and
provides shade and wind protection. A suitable temporary measure to provide
privacy is the application or burlap. Means of providing perches are endless,
but the best perches are made from 2-inch lumber. Straight tree limbs work
and are inexpensive, but they are hard to clean and maintain. Blankenship
(personal communication) uses a combination of the first two types of perches,
plus some brush piles and small live trees.

�-150-

Disease Problems
Apparently, disease problems do not occur frequently in bandtails if normal
precautions are taken. Pens should always be arranged so that sunlight enters
each pen. Normal cleaning is, of course, required. The only problem to
occur in this study was an infection of Aspergillus fumigatus caused by moist
conditions and possibly moldy grain.
Tetracycline (Terramycin) was used during this study for treatment of
Aspergillosis with poor success. In this case, a tetracycline solution was
used to replace the normal water supply. DeLaRonde and Greichus (1972) had
good results treating Aspergillosis-infected double-crested cormorants
(Phalacrocorax auritus) with a fog of tetracycline. Birds were treated three
times the first week, and once each of two following weeks. This treatment
should also work well for band-tailed pigeons.

Post-juvenile Molt
Primary Molt and Replacement
Molt of primaries is in sequence from innermost (PI) to outermost (PlO). At
about 45 days of age, primary molt begins and progresses at an uninterrupted
interval of one primary every 20 days. Individual primaries are fully
replaced in about 25 days, although some minor growth may occur for 4 or 5
weeks.
Table 1 shows the average number of days for bandtails to attain selected
stages of primary development during post-juvenile molt. For this table, primaries were visually classified as empty (E), one-fourth replaced (1/4), onehalf replaced (1/2), three-quarters replaced (3/4), or fully replaced (F).
The table shows that all juvenile primaries are replaced at 248 days or 8 months.
Primary Tipping or Edging
Both immature and adult bandtails show some light edging on the primaries. In
adult pigeons the edging is a fine white or cream colored line, while immatures
normally have a slightly wider brown line. Edging appears to be less evident
on the outer primaries. Thus, edging should be of minor value as an age
determination technique. In outer primaries the immature edging is difficult
to distinguish from adult edging of the innermost primaries. Age determination
while pigeons still retain juvenile primaries can be more easily and reliably
accomplished using primary molt, secondary examination, and covert examination.
Secondary Wear
No wear, as it occurs in wild-caught pigeons, has been observe~ in the captive
pigeons.. Some damage and accidental loss has occurred, but noticeable or
extensive wear have not occurred to date.

�-151Table l. Primary replacement of band-tailed pigeons, in days. (Ntnnbers in parentheses are sample sizes). Upper numbers represent means. Lower numbers represent
one standard deviation.
MOLT
E

Primaries
1

1/4
(N)

1/2
(N)

45*

52

(N)
60

(72)
16
2

64
13

3

19

84
17

4

108

114
(67)

20
5

18

125

133
(66)

18
6

19

145
20

7

25

164
20

8

22

183
18

9

23

202
21

10

24

221
23

*

Assume 45 days at loss of Pl.

27

(51)
22

241
(49)

22

226
(51)

26

235
(49)

(56)
26

221
(52)

23

226
(49)

200
(58)

23

214
(54)

(66)
20

199
(59)

19

206
(55)

190
(66)

20

195
(60)

(66)
24

182
(66)

26

189
(60)

165
(66)

19

179
(66)

(66)
19

163
(66)

21

171
(66)

150
(66)

22

155
(66)

(67)
23

143
(66)

20

151
(66)

126
(67)

19

138

(67)
20

125
(67)

(66)

110
(67)

21

20

(69)
19

104

119
(67)

89
(69)

(67)
19

(71)
25

25

97

16

68

82
(70)

(68)

(N)

(71)
20

20

92
(68)

65

75
(70)

F

(N)

(72)
22

70
(70)

3/4

248
(49)

24

(49)
25

�152-

Secondary Replacement
Most captive pigeons began post-juvenile molt of secondaries after P6 had
been molted, and continued until after the primary molt was completed (Fig. 1).
About 80 percent of the pigeons molted in sequence from outer (1) to inner (10).
Some pigeons exhibited irregular molt of secondaries in a sequence from 1 to
6 and then 10 to 7, or from 1 to 6 and simultaneously from 7 to 10. Timing
and initiation of molt is irregular and replacement is rapid. This character
is, therefore, of little use in calculation of hatching dates.
Examination of secondaries 1 through 5 or 6 appears to be useful for age determination purposes. Juveniles secondaries have rounded margins at the tip.
Secondaries 1 through 5 or 6 are replaced by square or slightly emarginated
secondaries in the post-juvenile molt. The presence of rounded, juvenile secondaries 1 through 5 or 6 would indicate a juvenile pigeon. Molt data indicates
that about 60 percent of juveniles retain S4, S5 or S6 after PlO is replaced.
Examination of secondaries, with practice, might allow recognition of juveniles or sub-adults after primary molt is completed in no more than 60 percent
of the birds.
Molt and Replacement of Rectrices and Tail Coverts
Tail characteristics are of little value in age determination because rectrices
and coverts are easily damaged and accidently lost. It is probable that this
is also true for wild-trapped pigeons. Molt of rectrices was not documented
in this study; however, Levi (1969) found that molt is in an irregular sequence
which occurs between 100 and 225 days of age (Fig. 1) in rock doves. Scattered
data were collected concerning molt of tail coverts and subsequent loss of
tail covert edging. The findings are summaried in Fig. 1. Molt of tail coverts
occurs between fledging and complete replacement of P4.
Upper Wing Covert Edging
Molt of upper wing coverts (including alula, alular coverts, and greater
secondary coverts) is summaried in Fig. 1. Wing coverts are lost continuously
from 0 molt until after PlO is completely replaced. About 20 percent of the
bandtails retained one or two buffy-tipped coverts after PlO was completely
replaced. About 30 percent retained buffy-tipped coverts after P9 was
replaced. As a result, wing covert edging is one of the best indicators of a
juvenile pigeon. This characteristic, in combination with primary and secondary examination allows accurate determination of age, and in some cases,
hatching date.
Color of Crown and Upper Breast
The chronology of the molt of the crown and upper breast feathers is expressed
in Fig. 1. Little replacement occurred until PJ was molted. At this time, a
fairly heavy molt occurred in this region, as well as on the nape. By the
time P7 was molted, most imma~ures have completed molt of the breast feathers.
At about 8 weeks of age (assuming 45 days at PIE) it is possible to distinguish sexes using the breast color as the criteria. As part of this study,

�Character
Loss of Natal
Feathering

Down*

of Nares

Wing Coverts
Alula
Alular

Coverts

Greater

Secondary

Upper Breast,

Coverts

Crown and Throat

Crescent
I
t-I

Nape

VI
IN

•

I

Secondaries
Eye
Cuticle
Tail

Coverts

Rectrices**

Fig.

*

1.

Molt sequence

Estimated
** After Levi (1969)

Fledging

o

of juvenile

band-tailed

PI

P2

pigeons

P3

P4

from fledging

P5

P6

to completion

P7

P8

of primary

P9

molt.

PIO

�-154-

it waadetermined
that, with practice, sex determination based primarily upon
breast color is at least 95 percent accurate. This figure is baaed on gonadal
examination of about 400 band-tailed pigeons of all age and sex classes. This
is baaed on the evidence that adult males have a purple hue on the crown,
throat, and breast, while females are more brown or gray.
Crescent and Nape
Immature bandtails have no crescent or irridescent nape as observed in adult
birds. MOst 0 molt pigeons have a uniform gray nape, although a few have a
very light crescent or irridescence. After P3 haa molted, a moderately heavy
molt occurs on the head and neck. By the time P6 or P7 have been molted, most
birds have completed molt of the nape and crescent (Fig. 1). These "subadults" are indistinguishable from adults; i.e., males have a Wide, clear
white crescent and well developed irridescence, while females have a thin or
mottled crescent and pale irridescence. This characteristic, used in combination with breast color is a useful indicator of sex.
Eye Color
Pigeons at 0 primary molt have a brownish-pink iris. Iris color rapidly
changes to reddish-brown after PI molt, and finally, to a blood red, characteristic of adults, by P4 molt (Fig. 1).
Cuticle Color
The color of the eye ring, or cuticle, in immatures is highly variable.
Colors observed included pinkish-yellow, pinkish-brown, gray, orange, yelloworange, and brownish orange. These colors changed rapidly after P3 waa molted
and by molt of P4 most pigeons had the dark pink adult cuticle (Fig. 1).
Nares
Feathering of the nares completes the juvenile plumage in band-tailed pigeons.
This generally occurs after fledging and by molt of PI is completed (Fig. 1).
Loss of Natal Down
Several immatures were captured each summer which still retained some of the
yellow natal down. This is probably lost just at fledging (Fig. 1) or before.
Bill, Foot, and Leg Color
Most immatures have pale, light colored bills, feet and legs. Most adults
normally have much darker pigmentation in these areas, although some do not.
By the time of P3 molt, all immatures were similar to adults in bill, foot and
leg color. Levi (1969) and other domestic pigeon raisers suggest that this
character may be used as an indicator of health. Presumably, dark or bright
colors indicate good health.

�-155-

Weight
Weight gain was rapid in captive immatures until a peak was reached in midOctober (Fig. 2). At capture, weights of immatures ranged from 190 to 290
grams. By mid-October the average weight was 345 grams. Since then the
average weight has dropped to about 330 grams and has remained remarkably
stable. The increase and subsequent slight decrease may have been related to
normal pre-migratory fat deposition with a return to "normal" after the migratory period passed. Stability of observed weights is probably attributable to
the apparent good health of all birds.
Sex Determination Using Wings Alone
While all available data has not been analyzed, some progress is being made in
this area. Using 1969 pigeon collection data, it was determined that about 25
percent of all pigeons could be classified correctly as to sex, using measurements of the outer primaries. However, more data from 1970 and 1971 should be
analyzed before making any final conclusion.
During examination of about 250 known sex wings from pigeon collections, it
was discovered that a large portion of the adult wings could be correctly
classified as to sex, using wing covert characteristics.
In a test, just over
90 percent of adult males were correctly classified as to sex, while only 80
percent of the females were correctly classified. However, sub-adults and
immatures could not be correctly classified as to sex. It was noted that
adult male coverts show a light metallic blue hue, while females present a
much darker gray with a faint touch of irridescent green.
In the near future, attempts will be made to improve methods of sex determination using wings alone. Perhaps, with practice, sex determination by
covert color, in combination with measurements of primaries, will yield a
usable method.
Abnormal Plumage in the Band-tailed Pigeon
The incidence of abnormal plumage in birds varies greatly within different
species. Gross (1965) reported albinism in 276 species of North American birds
with only 28 species compriSing 51.54 percent of all recorded albinos. In
general, species with close associations with man show a higher incidence of
abnormal plumage because of the increased chance of observation. Because of
this, the robin (Turdus migratorius) , house sparrow (Passer domesticus),
common crow (Corvus brachyrhychos), and red-winged blackbird (Agelaius
phoeniceus) share 20.01 percent of the recorded albinos (Gross 1965). In
addition, many game birds show a high incidence of albinism. Bobwhites
(Colinus virginianus), mallards (Anas platyrhynchos), ring-necked pheasants
(Pnasianus colchicus), Canada geese (Branta canadensis), ruffed grouse (Bonasa
umbellus), and California quail (Lophortyx californicus) total 9.01 percent
of recorded albinos (Gross 1965).
During recent investigations of band-tailed pigeons in Arizona, California,
and Colorado, several individuals displaying abnormal plumage have been
encountered. Three of these birds were partial albinos with patches of white
in the primaries. Lee Fitzhugh of the Arizona Cooperative Wildlife Research

�~eight
(In Grams)

425
400
375
345

350
325

I

340

'.
310

300

'(69)
285

I

275

I

f45

1335t30 t35
,

I

t340

330

(66)

(55)

(66) (66)

(49)

I

•••

(60)

VI

Q\

(67)

I

(70)

250;
I

I

(73)
230

225
200

(76)
175

o

1

2

3

4
Primary

5

6

7

1

1

r

8

9

10

Molt

Fig. 2. Weights of juvenile band-tailed pigeons at replacement of each primary.
vertical bars indicate ranges, numbers in parentheses i.ndicate sample sizes).

(Horizontal bars indicate means,

�-157-

Unit has observed two of these. Both were banded and released. One, a juvenile, was shot by a hunter afterward, and the wing was returned to State
Game and Fish personnel. The wing was loaned by Lee Fitzhugh for examination.
It is completely normal except that the two outer primaries are white. The
other partial albino was collected on June 6, 1970 at Stoner, MOntezuma County,
Colorado. This bird shows a pure white spot in the center of the primaries.
This pigeon is presently being mounted at the Denver Museum of Natural History.
Two bandtails have been encountered with imperfect albinism. One that was
noticeably lighter than normal was banded and released by J. Edward Kautz and
Brett Peterson in southern Colorado in 1971. The second imperfect albino was
captured near Carmel, California by F. John Ward on August 19, 1971. This
fawn-colored juvenile was sent to us for examination and at pr~sent it is
still alive and healthy. It appears normal in every way except for color. The
fawn color is apparently the result of a lack of dark eumelanin which produces
gray, black, and dark brown colors. However ,phaeomelanin, which causes light
brown and yellow-brown is still present and causes the light brown color of
this bird (Sage 1970).
Two melanistic bandtails have been observed by F. John Ward near Carmel, California. One was captured, but later escaped.
These seven birds have been encountered during the handling of over 17,000
band-tailed pigeons. This seems to indicate the rarity of abnormal plumage
in band-tailed pigeons. The occurrence of abnormality in plumage color is
probably at least one in 5,000 individuals based on trapping in Colorado,
Arizona, and California.

LITERATURE CITED
American Ornithologists Union. 1957. Check-list of North American birds.
5th ed. Lord Baltimore Press, Inc. Baltimore. 691 pp.
DeLaRonde, G. G., II, and Yvonne A. Greichus. 1972. Care and behavior of
penned double-crested cormorants. Auk 89(3):644-650.
Fitzhugh, E. L. 1970. Literature review and bibliography of the band-tailed
pigeons of Arizona, Colorado, New Mexico, and Utah. Spec. Rept., Arizona
Game and Fish Dept. 53 pp.
Glover, F. A. 1953. A nesting study of the band-tailed pigeon (Columba i.
fasciata) in northwestern California. Calif. Fish and Game 39(3) :397-407.
Goodwin, D. 1967. Pigeons and doves of the world.
Hist., London. Publ. No. 663. 446 pp.

British Museum Nat.

Gross, A. o. 1965. The incidence of albinism in North American birds.
Banding 36:67-71.

Bird-

Houston, D. B. 1963. A contribution to the ecology of the band-tailed pigeon
Columba fasciata, Say. M. S. Thesis, Univ. Wyoming, Laramie. 74 pp.

�-158-

Levi, W. M. 1969.
667 pp.

The pigeon.

Levi Publ. Co., Sumter, South Carolina.

Mace, R. U., and W. M. Batterson. 1961. Results of a band-tailed pigeon
banding study at Nehalem, Oregon. Proc. Ann. Conf. Western Assoc. State
Game and Fish Comms. 41 :151-153.
Macgregor, W. G., and W. M. Smith. 1955. Nesting and reproduction of the
band-tailed pigeon in California. Calif. Fish and Game 41(4) :315-326.
Miller, W. J., and F. H. Wagner. 1955. Sexing mature Columbiformes by
cloacal characters. Auk 72(3):279-285.
Neff, J. A. 1947. Habits, food and economic status of the band-tailed
pigeon. U. S. Fish and Wi1d1. Serv., No. Amer. Fauna 58. 76 pp.
Sage, B. L.

1970.

Albinism and melanism.

Birds

3(6):130-133.

Si1ovosky, G. D. 1969. Distribution and mortality of the Pacd Hc .Coast
band-tailed pigeon. M. S. Thesis. Oregon State Univ., Corvallis. 70 pp.
, H. M. Wight, L. H. Sisson, T. L. Fox, and S. W. Harris. 1968.
---Methods
of determining age of band-tailed pigeons. J. Wild1. Mgmt.
32(2):421-424.
Smith, W. A. 1968. The band-tailed pigeon in California.
and Game. 54(1):4-16.

California Fish

Wight, H. M., R. U. Mace, and W. M. Batterson. 1967. Mortality estimates of
an adult band-tailed pigeon population in Oregon. J. Wildl. Mgmt.
31(3):519-575.

Prepared by

_-=-::L~~~QlV~·-=--_f._
.

..L..~L.L.::..==';~

Clait E. Braun
Wildlife Researcher

_

�October,

-159-

JOB PROGRESS

State

of

Project
Work

~CO~L~O~RAD~~O~
No.

W-88-R-17

Plan No.

4

Job Tit Ie
Period

Covered:

April

Personnel:
Del Benson,
J. Allen White.

1972

REPORT

_
Migratory

Bird

Investigations

Job No.
~6
_
Rand-tailed
l'igeon Investigations
.::B:.::r...:;e:.:::;e;..:::d:,:i;.:;n:,cgt....:::a::,;n;..:::d:.....:.N:,:;e
_
1, 1971

to March

31, 1972

Clait

E. Braun,

J. Edward

Kautz,

Brett

Petersen

and

ABSTRACT

Investigations
were initiated
in 1971 to locate areas where band-tailed
pigeons
(Columba fasciata)
breed and nest in order to intensively
study
these facets of pigeon biology.
Efforts were continued
to collect pigeons
in order to study ovarian and testicular
cycles and their relationship
to
body condition
and crop gland activity.
Intensive
field investigations
in four different
areas suspected
of being utilized
for breeding
and
nesting by bandtails
failed in locating active nests.
Only one inactive
nest was found.
It is apparent
that nesting densities
of pigeons in
Colorado are extremely
low, possibly no more than three pairs per square
mile.
Examination
of gonadal materials
and crop glands indicate that bandtailed pigeons are inbreeding
condition when they arrive in Colorado
in
late April and are capable of breeding
into late August and early September.
While one egg is most commonly
laid, a Rmall percentage
of females may
produce two eggs per clutch.
All c o Ll cc t cd tna.t:crlftl/'J ~u~geF:Jr-- 1hi'll 114ml! 4111'l
may attempt two to three nests per breeding season.
Sub~tidult pf-geollB
definitely
attempt to nest, but they may initiate breeding
activities
later
than adults.

�-160-

RECOMMENDATIONS
1.

It is recommended that the field portion of this study concerning
locating and observing free-living band-tailed pigeons in breeding
and nesting areas be discontinued.

2.

The gonadal and crop gland investigation of this project should be
continued through 31 March 1973 with most analyses to be completed
by Ralph Gutierrez, Graduate Student, University of New Mexico,
Albuquerque.

�-161-

BAND-TAILED PIGEON INVESTIGATIONS
BREEDING AND NESTING CHRONOLOGY STUDIES
Clait E. Braun and Delwin E. Benson

The band-tailed pigeon, a native forest-dwelling Columbid, occurs throughout the southern Rocky Mountains and along the west flank of the Sierra
Nevada and Cascade Mountains in Western North America. Little is known
about the life history, biology, or ecological relationships of this
species as few detailed studies have been undertaken. The lack of systematic studies is primarily related to the secretive nature of bandtails
away from feeding sites, poor access throughout most of their range, and
purported low numbers of pigeons in some regions. As part of a larger,
comprehensive study of band-tailed pigeons seasonally resident in Colorado,
this study was initiated to investigate nesting habits and breeding chronology of these pigeons living in Colorado.

P. S. OBJECTIVE
To investigate life history and ecological facts pertinent to band-tailed
pigeons in Colorado as follows: range; relative numbers; relation between
numbers of birds present by area and such physical factors as weather,
food, vegetative cover, elevation, and salt; breeding and nesting chronology; migration patterns and chronology; harvest patterns; and mortality
rates.

SEGMENT OBJECTIVES
1.

To locate several breeding and nesting areas of free-living band-tailed
pigeons suitable for intensive studies.

2.

To document breeding and nesting chronology from field observations,
gonadal, and crop gland examination of pigeons systematically collected
in Colorado.

3.

To compile data and prepare progress report.

REVIEW OF LITERATURE
Two major populations of band-tailed pigeons occur north of Mexico with

~. i· monilis found from southern California north through Oregon, Washington,
into south-central British Columbia, primarily west of the summit of the
Sierras and Cascade Mountains. The interior population, ~. i. fasciata,
appears to be more restricted in range as it occurs from Utah and Colorado
south through New Mexico and Arizona, wintering in north-central Mexico
(A.O.U. 1957). Other subspecies occur south of the International Boundary

�-164-

La Veta - Wahatoya Creek; 8,000-9,000 feet in ponderosa pine, Gambel oak
(Quercus gambeli), and Douglas-fir.
Salida - Methodist Mountain; 8,000-10,000 feet in Engelmann spruce (Picea
engelmannii), limber pine (Pinus flexilis), and bristlecone pine (Pinus
aristata).
Longmont - Flying over ponderosa pine and Douglas-fir at 7,000 feet.
Castle Rock - Bear Canyon and adjoining areas in the Rampart Range; 8,0009,000 feet in lodgepole pine (Pinus contorta), Douglas-fir, limber pine,
and quaking aspen.
At this time, no specific vegetation type or elevation can be correlated
with nesting.
Following pigeons to nesting areas from feeding sites worked for a portion
of the distance. As the vegetation became thicker and the terrain more
diversified, selecting correct observation points became difficult. Pigeons
did not follow exact drainage routes and once feeding flight patterns were
lost, it took many trial and error locations to again find the flight path.
Spiral nuptial flights were not observed. Birds were found cooing from sunrise to 9:30 a.m., MDT, and from 4:00 p.m., MDT, to sunset. These areas
were searched for nests, but none ~as found. Cooing activity was observed
mainly in Bear Canyon and surrounding areas in the Rampart Range west of
Castle Rock. This area was intensively studied because of published reports
of nests found there in the late 1940's. Cooing was also commonly heard and
observed at pigeon feeding sites.
Ground reconnaissance surveys in probable nesting areas did not prove
productive. Band-tailed pigeons were flushed and observed flying, but the
magnitude of vegetation and land area to support nesting birds was too great
to adequately cover.
Analyses of all collected materials have not been completed at this time.
Cursory examinations of gonadal materials and crop glands indicate that
band-tailed pigeons are in breeding condition (enlarges testes and ova)
when they arrive in Colorado in late April. Testes remain enlarged throughout the summer and start regressing in late August and early September.
There is some suggestion that testes may slightly regress (about 1 to 2 mm)
during the period of late incubation and feeding of young. Females show
ovarian activity from late April until early September with up to six ova
being larger than 5.0 mm at one time. Examination of ovarian. materials
indicates that up to 30 percent of the females in some years may lay clutches
of two eggs. However, one egg clutches would appear to be most common.
There is little doubt that most pigeons in Colorado attempt at least two
nests each year and possibly three nests in extremely early years or when
feeding conditions are good. Crops exhibiting glandular development have
been examined from late May until late September. It could be argued that
this finding just suggests that pigeons are asynchronous nesters, allowing
individual birds to be in various stages of one nesting attempt. This

�-165-

argument is not reasonable as it does not explain those pigeons that
have been collected with "active or stimulated" crops and having enlarged testes or a developing follicle indicative of breeding readiness.
While two nesting attempts were most common in 1969 and 1970, probably
only one nesting attempt occurred per pair of pigeons in 1971. This
was undoubtedly the result of extremely poor food conditions in that
year.
Only one adult or subadu1t pigeon has been collected that apparently
had not been in breeding condition during the time interval prior to
death. All field data indicate that subadu1t pigeons (less than 1 year
of age) breed. However, this age class does appear to nest later than
adults. While most pigeons are capable of breeding, not all males
appear to be successful in attracting a mate.
Present Status
Graduate Research Assistant Delwin E. Benson terminated his contract
with the Colorado Division of Wildlife by mutual consent on December 31,
1971. The nesting and breeding phenology segment was held inactive from
that time until May 1, 1972. At that time an agreement was reached with
Ralph Gutierrez, a graduate student at the University of New Mexico,
Albuquerque, to continue and complete the study using primarily the
collected gonadal materials. All materials, measurements, descriptions,
etc., were turned over to him for analyses. Additionally, it was agreed
that fresh materials would be provided through 1972. The final report
will be compiled primarily from the M.S. thesis of Gutierrez with additional data from the aviary studies of White (1972).

LITERATURE CITED
Abbott, C. G. 1927. Notes on the nesting of the band-tailed pigeon.
Condor 29(2):121-123.
Allen, W. I. 1941. Nesting of band-tailed pigeons at Altadena,
California. Condor 43(3):156-157.
American Ornithologist's Union. 1957. Check-list of North American
birds. 5th ed., Lord Baltimore Press, Inc., Baltimore. 69lpp.
Brandt, H.

1937.

Brown, W. W., Jr.

Some Arizona bird studies.
1909.

Correspondence.

Auk 54(1):62.

Condor 9(4):143.

Burleigh, T. D. 1929. Notes on the bird life of northwestern
Washington. Auk 46(4):502-519.
Cooke, W. W. 1898. Further notes on the birds of Colorado.
Expt. Sta., Ft. Collins, Bull. 44:147-176,179-239.

Agric.

�-166-

Cottam, C. 1941. Indigo bunting and band-tailed pigeon in Utah.
Condor 43(2):122.
Davis, J. M. 1938. Nesting dates from the Humboldt Bay region.
Condor 40(4):182-183.
Dawson, W. L. 1923. The birds of California. South Moulton Co.,
San Diego, Los Angeles, San Francisco. Vol. 3 of 4, 2l2lpp.
Fitzhugh, E. L. 1970. Literature review and bibliography of the
band-tailed pfgeons of Arizona, Colorado, New Mexico and Utah.
Spec. Rept., Arizona Game and Fish Dept. 53pp.
Gilman, M. F. 1903. More about the band-tailed pigeon (Columba
fasciata). Condor 5(5):134-135'.
Glover, F. A. 1953. A nesting study of the band-tailed pigeon
(Columba f. fasciata) in northwestern California. Calif. Fish
and Game 39(3):397-407.
Grinnell, J. 1938. September nesting of the band-tailed pigeon.
Condor 30(1):126-127.
______ , H. C. Bryant, and T. I. Storer. 1918. The game birds of
California. Univ. California Press, Berkeley. 642pp.
Hargrave, L. L. 1935.
Condor 37(6):285.

Nine new birds from Williams, Arizona.

Houston, D. B. 1963a. A contribution to the ecology of the bandtailed pigeon, Columba fasciata, Say. M. A. thesis, Univ. Wyoming,
Laramie. 74pp.
1963b. The reproductive ecology of the band-tailed pigeon,
Columba fasciata. Jour. Colo.-Wyo. Acad. Sci. 5(4):42.
Huey, L. M. 1913. With the band-tailed pigeon in San Diego County.
Condor 15(4):151-153.
Jensen, J. K. 1923. Notes on the nesting birds of northern Santa Fe
County, New Mexico. Auk 40(3):455.
Jewett, S. G. 1941.
43 (1) : 78.

Late nesting of the band-tailed pigeon.

Condor.

Johnson, N. K. 1965. The breeding avifaunas of the Sheep and Spring
Ranges in southern Nevada. Condor 67(2):93-124.
Lamb, C. C.

1926.

The viosca pigeon.

Condor 28(6):262-263.

Macgregor, W. G., and W. M. Smith. 1955. Nesting and production of
the band-tailed pigeon in California. Calif. Fish and Game.
41(4):315-326.

�-167-

March, G. L., and R. M. F. S. Sadleir. 1970. Studies on the bandtailed pigeon (Columba fasciata) in British Columbia. I. Seasonal
changes in gonadal development and crop gland activity. Canadian
Jour. Zool. 48(6):1353-1357.
Matteson, C. P. 1948. Life history (wild and aviary).
Game and Fish Fed. Aid Rept. October. pp. 20-22.

Colo. Dept.

1949. Life history (natural and aviary). Colo. Dept. Game
and Fish. Fed. Aid Rept. October. pp. 36-37.
Mearns, E. A., M.D. 1890. Observations
Arizona. Auk 7(1):52-53.

on the avifauna of portions of

Michael, C. W. 1928. Nesting time of the band-tailed
Valley. Condor 30(1):127.
Morse, W. B. 1949. The band-tailed
Bull. 4(7):1,5,7.

pigeon.

pigeons in Yosemite

Oregon State Game Comm.

Neff, J. A. 1947. Habits, food and economic status of the band-tailed
pigeon. U. S. Fish and Wildl. Ser., No. Amer. Fauna 58, 76pp.
___

, and J. C. Culbreath. 1947. Status of the band-tailed pigeon
in Colorado, season of 1946. Colo. Dept. Game and Fish, Mgmt. Div.
Fed. Aid Sec. Rept. 24pp.

____ ~, and R. J. Niedrach. 1946 •. Nesting of the band-tailed
in Colorado. Condor 48(2):72-74.
Peeters, H. J. 1962. Nuptial behavior of the band-tailed
the San Francisco Bay Area. Condor 64(6):445-470.

pigeon

pigeon in

Pemberton, J. R., and H. W. Garriger. 1915. A partial list of the
summer resident land birds of Monterey County, California.
Condor 17(5):189-201.
Peyton, S. B. 1937. Minutes of Cooper Club Meeting - Southern Division.
Condor 39(2):95.
Pierce, W. M. 1913.
15(5) :227.

Nesting of the band-tailed

pigeon.

Condor

Racey, K. 1926. Notes on birds observed in the Alta Lake region,
British Columbia. Auk 43(3):319-325.
1948. Birds of the Alta Lake region, British Columbia.
Auk 65(3):383-401.
Rockwell, R. B. 1908. An annotated list of the birds of Mesa County,
Colorado. Condor 10(4):152-180.

�-168-

Rowley, J. s. 1934. Notes on the nesting of band-tailed pigeons.
Condor 36(5):216-217.
Schwi11ing, M. D. 1950. More band-tails nesting in Co10~ado.
Unpubl. Note. Files of Colo. Div. Wi1d1., Ft. Collins.
Sharp, C. S. 1919. Nesting of the band-tailed pigeon in San Diego
County, California. Condor 21(1):40.
Smith, A. P. 1917.
19(5):161-165.

Some birds of the Davis Mountains, Texas.

Condor

Stager, K. E. 1954. Birds of the Barranca De Cobre region of southwestern Chihauhua, Mexico. Condor 56(1):21-32.
Stephens, F. 1913.
15(3):129.
1914.
Swarth, H. S.
1908.

Early nesting of the band-tailed pigeon.

Arizona records.
1900.

Condor

Condor 16(5):259.

Avifauna of a 100-acre ranch.

Condor 2(1):16.

Some fall migration notes from Al-izona. Condor 10(3):

108.
Thayer, J. E.

1909a.

Letter to the editor.

Condor 11(4):142-143.

190,9b. Some rare birds and sets of eggs from the Cape region
of Lower California. Condor 11(1):10-11.
Vorhies, C. T. 1928. Band-tailed pigeon nesting in Arizona in September.
Condor 30(4):253.
White, J. A. 1972. Plumage studies of band-tailed pigeons. Colo. Div.
Wi1d1., Game Res. Rept., Fed. Aid Proj. W-88-R. October. In Press.
Willard, F. C. 1913. Some late nesting notes from the Huachuca Mountains,
Arizona. Condor 15 (1):41.
1916. Nesting of the band-tailed pigeon in southern Arizona.
Condor 18(3):110-112.
Ziegler, D. L. 1971. Crop4mi1k cycles in band-tailed pigeons and
losses of squabs due to hunting pigeons in September. M.S. thesis.
Oregon State Univ., Corvallis. 48pp.

Prepared by
i

_~~~",-,fJ~~J:
~,-- _
Clait E. Braun
Wildlife Researcher

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                  <text>January,

-1-

1973

JOB FINAL REPORT

State of

.COLORADO
------~~~~~--------------

Work Plan NO'.
Job Title:

2

Investigations
3

Job No.
Physiological

Period Covered:
Personnel:

Antelope

W-40-R-13

Project No.

Studies

May 1, 1965 through April 30,

George D. Bear, Allen E. Anderson,
Florence Fields.

1971
John P. Goettl, Robert Keiss,

ABSTRACT

Seventy-five antelope were collected and necropsied during this study. Buck
and doe antelope were collected at monthly intervals for a one-year period in
Moffat County and Saguache County; also wounded or dead animals were collected
following the hunting season. Measurements were recorded in the following
categories for each animal:
external measurements
- (shoulder height, total
body length, tail length, body length, head length and width, ear lengths,
hindfoot lengths, hoof lengths, horn lengths, girth, neck circumference and
carcass weights);
measurements of organs
- (heart, liver, brain, spleen, kidney,
eyeballs, lungs); measurements of glands
- (thyroid, adrenal and pituitary);
measurements of the male reproductive system
(testes, prostate gland, scrotum
and penis); measurements of the female reproductive system
(ovaries, uterine
horn, vagina, cervix, uterine body and embryos);
blood analysis
erythrocytes,
leucocytes, neutrophils, lymphocytes, monocytes, eosinophils,
basophils, hemoglobin and hematocrit);
stomach weights;
body temperatures; and body condition
indices
(thoracic fat, visceral fat, kidney fat, bone marrow, and rump fat).
These data were grouped into age and sex categories:
fawns (sexes combined),
yearlings (buck or doe), and mature (buck or doe). Single mean values were
computed for various measurements and entered in tables under the sex and age
categories;
except for the measurements on the reproductive systems, which
were presented by monthly intervals in an effort to establish seasonal trends
or changes.

�-2-

Body, organ and gland measurements increased most rapidly during the period
from the fawn to the yearling age class.
Increase in these measurements were
less pronounced between the yearling and mature age categories.
Individual
variations and small sample sizes obscured differences due to sex, if such
differences do occur.
Measurements
of the female reproductive system showed seasonal changes; whereas, data for the male reproductive system was erratic and did not show pronounced changes.
.Changes in the female reproductive system were closely
associated with .fetus,development, which accelerated in the latter four months
of pregnancy.
..
.....
.
.
Mean muscle and rectal temperatures taken soon after the death of the animals
were 38.7 and 38.4 degrees centtgrade, respectively.
Heat loss from the
carcass was most pronounced during the first three hours
after the animal
was eviscerated.
Measurements on body fat deposits were reported in a technical publication
entitled "Seasonal trends in fat levels of pronghorns, Antilocapra americana,
in Colorado"; J. Mammalogy 52(3):583-589.
Data from this study were also
printed in the following publication, "The annual testicular cycle and horn
casting in the pronghorn (Antilocapra american)";
J. Mammology 52(3):537-544.

�-3-

PHYSIOLOGICAL

STUDIES

George D. Bear

P. S. OBJECTIVE
To measure physiological and morphological characteristics of antelope
collected seasonally to establish "physiological norms" and to provide
basic morphological data as related to sex and age classes.

METHODS

AND MATERIAL S 1/

Antelope were collected at monthly intervals on study areas near Maybell
in Moffat County, Colorado and near Saguache in Saguache County, Colorado;
also, a few dead or wounded antelope were picked up following the 1964
hunting season in Moffat County.
A total of 75 antelope were collected
and necropsied during this study. With the exception of the data concerning
female reproductive systems, there appeared to be little difference in the
data cQllected from each of the two study areas, thus the data were combined
for firlal analysis.

Field Procedures

Shooting

of Antelope

Location of Shot--When
the neck and shoulder.

possible,

the antelope was shot at the junction

of

Observations Recorded--The following observations were recorded: behavior of
the antelope before and after shooting, time of shooting, time of death,
estimated distance of shot in yards, weather conditions, and number and behavior of associated antelope.

Collection

of Blood Samples

Method of Entering Heart--Blood samples were collected within 15 minutes
of shooting.
The blood was aspirated from the heart with a syringe and
needle.
The heart was entered from the left side between the fourth and
fifth ribs and about two inches above the sternum.
The index finger was

1/

- Field and laboratory
Anderson

and Medin,

procedures

1962.

were patterned

after those developed

by

�-4-

used to push aside the pelage at the point of needle entry until the point
was inside the thoracic cavity. Both hands were then used to introduce
the needle into the heart; the syringe plunger being slowly withdrawn when
contact was made with the heart.

Anti-Coagulant Samp·le--When about 20 cc of blood had been obtained, the
syringe was detached from the needle and about 7 cc of blood was gently
flushed into each of two anti-coagulant tubes. The blood was flushed
gently and allowed to trickle down the tube walls. Immediately after
filling, each tube was stoppered, held downward momentarily, and its top
moved gently through an arc of 1800 for about two minutes.
Serum Sample--After the anti-coagulant sample had been obtained, two
samples of about 50 cc each were aspirated in the same manner and flushed
into each of the two 60 ml centrifuge tubes.
Female Reproductive Tract
Vulva Measurements--The maximum length and width of the vulva were measured
with a transparent rule to the nearest millimeter as soon as possible after
death.
External Body Measurements

General Methods--Three additional external body measurements were made as
soon as possible after death. They were taken with a flexible steel tape
to the nearest centimeter with the carcass lying on a flat surface.

Measurements
Total Length--The distance measured from the tip of the nose to the distal
end of the last tail vertebrae. The carcass is laid on its right side
with head extended forward so that the nose is brought into as nearly a
straight line with the back as possible, the tape being passed from the
end of the nose over the top of the head directly to the top of the
shoulders and along the backbone to the distal edge of the last coccygeal
vertebrae.
Shoulder Height--The distance measured from the posterior edge of the left
hoof to the top of the scapula.
Girth--The circumference of the body within the first inch posterior to the
edge of the scapula. The tape is held firmly but without undue tension.

�-5-

Laboratory

Blood Analyses

Procedures

and Chemical

Determination

The blood samples w.ere analyzed for the following:
erythrocyte count,
leucocyte count, differential leucocyte count (neutrophils, lymphocytes,
monocytes, eosinophils and basophils), hemoglobin, and hematocrit.
This
work was done at the Game, Fish and Parks Division Research Center, Fort
Collins, Colorado, by the laboratory technicians.

External

Body Measurements

General Methods

All external body measurements were made in the metric system at three to
24 hours from the time of death.
The measurements, instruments used and
their precision were described as follows:

Measurements

Ear Lengths--Intertragic
notch to the tip of the auricle of both ears, not
including hair, with a transparent rule to the nearest millimeter.
Head Length--Occiput

to the incisors with a caliper to the nearest millimeter.

Inter-orbital Width--The maximum distance between the zygomatic
measured with a caliper to the nearest millimeter.

arches

Neck Circumference--Two
measurements: oue (a) made at a point approximately
10 centimeters posterior to the occiput and the other (b) at the extreme
lower base of the neck. Both measurements were made with a steel tape and
measured to the nearest millimeter.
The tape was held snugly but without
excessive tension.

Hoof Lengths--Measured
on all four feet as the maximum distance between the
tip of the outside claw to its posterior edge. These measurements were made
with a vernier caliper to the nearest millimeter.

Hind Feet Lengths--Measured
on both limbs from the tuber calcis (calcaneum)
of the fibular tarsal bone to the tip of the hoof (outside claw) with a tape
to the nearest millimeter.

�-6-

Tail Length--Measured from the perianal region to the distal end of the
last coccygeal vertebrae with a transparent rule to the nearest one-half
centimeter. The tail was held straight in relation to the spine and its
ventral surface laid along the edge of the rule.

Mammary gland--The maximum length and width were measured from the hair
line, or point of swelling, and the maximum depth of a medial, longitudinal
inclSl0n. A transparent rule was used in all cases, the length and width
being measured to the nearest centimeter and also measured with a transparent rule to the nearest millimeter while their basal diameters were
measured with a vernier caliper to the longitudinal axis of the body.
Male Reproductive Organs--Measurements include the maximum length, width,
and depth of the scrotum and the length of the penile sheath from the
scrotum to the distal end. These were measured with a transparent rule
to the nearest one-half centimeter.

Eviscerated Carcass--Weighed with all viscera and visceral fat removed.

Gland Methodology

General Methods
Immediately after dissection the glands were placed in individual, labeled,
beakers containing physiological saline (prepared with tap water and either
non-iodized table NaCl or reagent grade NaCl) until all fat and extraneous
tissue could be removed. After blotting, measurements including longitudinal, transverse, and height diameters were made with a vernier caliper
to the nearest 0.1 millimeter. Fresh weights were then obtained on a triplebeam balance scale to the nearest centigram. Volumes were calculated using
their weight in air and submerged weights in water in the following formula
based on Archimedes' principle and as illustrated by White (1956:219):
gland + apparatus
Volume in cc

(wt. in air - wt. in H2O)
apparatus

- (wt. in air - wt. in H2O)
Measurements, Weights, Volumes, Special Treatments

Adrenals--Longitudinal,
were measured.

transverse, and vertical diameters of each adrenal

�-7-

Thyroid--The maximum longitudinal, transverse and vertical
the weight and volume of each lobe were obtained.

diameters

and

Pituitary--Following
dissection of the brain, this gland was removed from
the sella turcica by cutting the dura mater at the rim of the sella turcica
with a scalpel, cutting the infundibular stalk and gently removing the
encapsulating dura mater with a probe.
The maximum and minimum diameters
were measured and its weight and volume determined.

Testes and Epididymides--Each
testis was removed from the scrotum by
severing the spermatic cord at the caudad portion of the ductus deferens
as welt as the tunica vaginalis where attached to the testes.
The tunica
albuginea was then carefully stripped from the testes exterior and the
epididymis is removed by severing the coni vasculosi at their junction with
the testes.
The longitudinal and transverse diameters of each testes were
measured with a vernier caliper to the nearest millimeter; their individual
weights and volumes were recorded.

Prostate--The prostate
bladder and its weight

gland was dissected from the neck of the urinary
and volume determined.

Ovaries--The maximum longitudinal, transverse, and vertical diameters were
measured to the nearest 0.1 millimeter with a vernier caliper; fresh w~ights
and volumes were also obtained.

Organ Methodology

General Methods
After dissection, each organ was washed in tap water and placed in physiological saline.
Paired or fragile organs were placed in separate marked containers while the larger, discrete organs were placed in a common container.
Following the removal of fat and extraneous tissue the organs were blotted,
measured, and weighed on a triangle or double beam balance scale to the
nearest 0.1 gram of I gram as stated below.
Volumes were determined as
described for glands.

Dissection,

Measurements,

Weights,

Volumes

Special Treatments

Liver--The maximum longitudinal (left to right lobes), transverse (right
lobe at right angles to the longitudinal axis of the liver), and height
(along the transverse bisection) diameters were measured to the nearest
millimeter.
A caliper was used to measure the longitudinal and transverse
diameters and a transparent rule for the vertical diameter measured along
the transverse bisection.

�-8-

Kidneys--Each kidney was removed by first cutting the fat anterior and
posterior to the kidney and at right angles to its longitudinal axis.
A cut was then made parallel to its longitudinal axis and immediately
adjacent to the kidney capsules thereby severing the fat, ureter pelvis,
and the renal arteries and veins.
The kidney was weighed to the nearest
0.1 gram with fat after removal of the ureter pelvis and the renal
arteries and veins at the point of their junction with the hilum.
The
attached fat was then removed and the kidney was weighed without fat.
The maximum longitudinal, transverse (at the hilum), and vertical (opposite
the hilum) diameters were measured with a caliper to the nearest millimeter.
The kidney fat index (percent) for each kidney was computed by subtracting
the kidney weight without fat from the kidney weight with fat and dividing
the difference by the kidney weight without fat. The result was multiplied
by 100. Total kidney fat index was computed the same way from the summed
values of both kidneys.

Heart--The pericardium was removed, coagulated blood in the chambers washed
out with tap water, and all vessels cut at their junction with the heart.
Preparatory to measurement, the heart was placed so the left side was facing
the measurer.
The maximum longitudinal diameter was measured from the left
atrium (auricle) between the aortic arch and superior vena cava to the apex.
The maximum transverse diameter was measured at the point of greatest heart
breadth.
Both measurements were taken with a caliper and read to the nearest
millimeter.
The weight was recorded to the nearest gram, and the volume was
taken.

Spleen--Weight to the nearest gram and volume were recorded
also the longitudinal and transverse diameters.

for this organ,

Lungs--The left and right lungs were separated
gram. No measurements or volumes were taken.

to the nearest

and weighed

Brain--It was removed by first making three cuts with a bone saw; two along
the edge of the frontal and parietal bones and one at right angles to the
longitudinal axis of the skull and at, or just above the post-orbital process
of the frontal bone. A rongeur forceps was then used to remove the frontal,
parietal, and interparietal bones thus completely exposing the dorsal surface
of the brain from the olfactory bulbs to the medulla oblongata.
A probe
and blunt scissors were used to separate the meninges from the brain case and
to sever the cranial nerves and the infundibulum stalk at the ventral surface
of the cranium.
When the brain was completely free in the cranium it was
moved forward and the spinal cord was severed at the extreme posterior portion of the rhomboid fossa or level of the obex. This appeared as the
posterior point of a shallow depression on the dorsal surface of the medulla
oblongata.
The described point of severance corresponds roughly to the
junction of the medulla oblongata and spinal cord. Following removal of the
meninges the maximum longitudinal diameter was measured with the brain on its

�-9-

ventral surface from the frontal to the occipital lobes of the cerebrum
immediately parallel to the longitudinal fissure.
The maximum transverse
diameter of the brain was measured at a selected point on the temporal
lobes. All measurements were made with a caliper to the nearest millimeter;
weight was recorded to the nearest gram •

.Eyeballs--Both eyeballs were removed from their orbits by first cutting the
conjunctiva (the mucous membrane which lines the eyelids) completely around
the eyeball then pressing the eyeball gently outward and to the side.
The
extrinsic nerves, muscles, and vessels were severed with scissors and the
eyeball withdrawn from its socket.
Two external maximum diameter measurements were made with a vernier caliper to the nearest 0.1 millimeter as
follows:
Longitudinal, from the cornea to the extreme posterior wall;
transverse, from wall to wall at a right angle to the longitudinal measurement.
Weights were taken to the nearest 0.1 gram.

Stomach and Contents--The stomach was removed by severing at the cardia and
pylorus and tying each end with string to retain the contents.
It was
weighed to the nearest gram on a balance scale.
The omasum and abomasum
were removed, then the rumen and reticulum and their contents were weighed.
The entire stomach without the contents was then weighed.

Measurements

of Teeth and Bones

Mandibular and Maxillary Teeth Row Series--The series were measured with a
vernier caliper to the nearest 0.1 millimeter.
Measurements included (1)
the alveolar distance from the anterior border of the anterior premolar to
the posterior border of the posterior molar.

Length of Mandible--It was measured with a caliper to the nearest millimeter.
The maximum length of the left ramus was measured, not including any part of
a tooth that extends forward from the anterior end of the bone.

Length of Mandibular Diastema--The distance from the posterior-most margin
of the alveolus of the lower incisor to the anterior-most margin of the first
fourth cheek tooth was measured with a caliper to the nearest millimeter.

Left Femur and Tibia--Before dissection, the femur was cracked, marrow from
the central portion of the shaft was removed, placed in a crucible and
macerated into a homogenous mass. Marrow color was immediately classified
using the color charts of Munsell (1929-1960). The marrow consistency was
rated in one of three categories, (a) solid, (b) putty-like, and (c) gelatinous.

Right Femur and Tibia--No measurements were taken.
The bones were fairly
well cleaned, labeled, and placed in a labeled polyethylene bag and frozen
immediately.
After storage the marrow was removed for analysis of fat at
the laboratory.

�-10-

Female Reproductive

Tract

General Methods--Immediately
after dissection the intact tract was placed
in physiological saline.
After draining, blotting, and trimming all fat
and the bladder (at its junction with the urethra), the tract was measured
and weighed as described below.

Measurements--Allmeasurements
were made with the tract lying on its dorsal
surface and include the length and width of the uterine horns and body,
vagina, and cervix.
All measurements were made with a transparent rule to
the nearest millimeter.

Weight--Weights
were obtained with balance scales to either the nearest one
or ten grams depending on its size. In the gravid uterus, the weight of the
tract with its uterine fluids and membrane was found by subtracting the
weight of the embryo(s) or fetus(es).

Measurements

of Fetuses

and Embryos

External measurements of the fetuses were made as for the larger adult
animals.
No internal measurements were made.
Only a rump-crown measurement
was made on very small embryos.

Fat Deposition
Records of fat deposition on the brisket, rump, and visceral fat were made
in addition to the kidney fat index and the marrow fat indices.
Depth of
the brisket fat was measured on the sternum at the third rib, when a cut
was made parallel to the longitudinal axis of the carcass.
Depth of the
rump fat was measured at the maximum fat depth along a cut approximately
12 inches long made at a 450 angle to the backbone starting at the base
of the tail and proceeding anteriorly across the right side of the rump.
These measurements were made with a transparent rule to the nearest millimeter.
Fat in the visceral cavity, including mesenteric fat and kidney
fat, was removed and weighed to the nearest gram.

RESULTS

External

AND DISCUSSION

Body Measurements

There was a general increase in the external body measurements of the
antelope as related to the three age classes, fawn, yearling, and mature
(Tables 1-5). The only fawns necropsied were ones found after the 1964
hunting season.
These animals were approximately three to four months old.
The yearling age class included animals from eight months to twenty-four
months old; while the mature class was all animals over twenty-four months

�-11-

old. Growth or increase in body measurements was much more pronounced
from the fawn class to yearling, than from yearling to mature age class.
It is difficult to make a reliable comparison between the groups due to
the small sample sizes and distribution of the animals within each of
the broader age categories.
However, these data indicated there were
minor differences in the body measurements due to sex. Mature bucks were
slightly heavier, had slightly larger girth measurements,
and larger head
measurements than mature does. Percent decrease in body weight following
evisceration was as follows:
fawns 25 percent, yearling bucks 21 percent,
yearling does 28 percent, mature bucks 27 percent, and mature does 31
percent.

Organ Measurements
Organs generally increased in size in relation to the general growth
pattern, with the increase being most pronounced from the fawn age group
to the yearling age group (Tables 6-10). Mean organ measurements for
yearlings and mature bucks were slightly greater than similar measurements
for yearling and mature does. However, variations in each of these measurements within each sex and age class were quite large, and larger samples
would be needed to determine the validity of the above statement.
Measurements for the left and right eyeball and left and right kidney were very
similar for each of the animals necropsied, so a mean figure was computed
for each animal.
The left lung weighed less than the right lung in all
animals necropsied.

Gland Measurements
As in previous measurements, the gland measurements showed a greater increase in size from the fawn class to yearling than yearling to mature age
class (Tables 11-15).
Any differences in gland measurements due to sex of
the animals are likely due to small samples and the method of grouping the
data.

Reproductive

Sy~

Measurements on the reproductive systems of yearling animals were combined
with measurements of the mature age class, since the yearling animals were
reproductively mature.
Data from the fawn age classes are presented in
Tables 16 and 17.
There appeared to be a slight increase in testes size and the longitudinal
measurements for bucks in August and October; but other measurements on
the prostate gland, scrotum and penis did not indicate a seasonal change
(Table 18).
The female reproductive system showed seasonal changes.
Ovarian and vaginal
measurements did not show significant changes while the remainder of the
reproductive system changed with fetal development (Tables 19 and 20). Data
collected in Moffat County was presented in a table separate from data

�-12-

collected in Saguache County because fetal development in the Saguache
County does appeared to be one month behind the Moffat County does.
The uterine horns and uterine body greatly increased in size from
November through Mayas
the fetus developed; then decreased abruptly
in June when parturition occurred.
Fetal development was very pronounced during the last four months of pregnancy; weight of the fetuses
nearly doubled each. month during this period.

Blood Analysis
Data obtained on blood chemistry showed considerable variation, although
for does and bucks the mean values were similar (Tables 21 and 22). The
data were examined on a monthly basis but the trends were very erratic.

Stomach Measurements
Increase in'stomach size was most pronounced between the fawn and yearling
age classes, which is the most active period of growth (Table 23). There
was little difference in mean stomach weights for bucks and does in the
same age class.

Body Temperatures
There was little difference in the muscle temperature and rectal temperature of the antelope collected with means of 38.7 degrees and 38.4 degrees
Centigrade (Table 24). There was always a few minutes lag after the animal
was shot before the temperatures could be taken, thus these mean values
are likely to vary from temperatures of living animals.
The muscle temperature recorded for the antelope after they were eviscerated showed a marked
decrease during the first three hours, after that heat loss was much slower.
These temperatures were collected under field conditions in which the air
temperatures were extremely variable.
Therefore, a study with stricter
controls on air temperatures would yield more reliable data.

Body Condition

Indices

Seasonal trends in the fat levels of males and females were traced using
thoracic fat, visceral fat, kidney fat, bone marrow, and rump fat measurements.
The results of these findings were reported in a technical publication entitled "Seasonal trends in fat levels of pronghorn, Antilocapra
americana, in Colorado", published in the Journal of Mammalogy, 52 (3):
583-589, 1971.

�-13-

LITERATURE·CITED

Anderson, A. E., and D. E. Medin. 1962. An ecological investigation
of the Cache la Poudre deer herd, Colorado -- physiological studies.
Colorado Quarterly Report, July 1962, pp. 253-275.
Munsell, A. H. 1929-1960. Munsell book of color, pocket edition.
Munsell Color Co., Inc. Baltimore, Maryland.
White, H. E. 1956. Modern college physics.
New York. 824 pp.

.r H"
:

Prepared by

/

_/~v_~,.-~1-;;./___ ',-~/

)

~-;
,~~,-{

George j{'. Bear
Wildlife Researcher

4

.

D. Van Nostrand Co., Inc.

�-14-

Table
three

1.
to

Body measurements
four months old.

(centimeters)

for

fawn antelope

approximately

Measurement

Mean

Range

Sample
Size

Shoulder

height

75.3

87.1- 66.5

7

6.6

Total

body length

113.0

125.0-107.5

8

6.3

Tail

length

10.3

15.2-

6.5

8

2.5

Body length

102.8

109.8- 97.3

7

4.7

Head length

22.5

24.2- 21.3

6

1.0

10.3

10.6-

9.9

6

0.2

Ear lengths

14.2

15.2- 12.2

8

0.9

Hindfoot

37.4

39.7- 34.2

8

1.9

5.8

6.2-

4.8

8

0.5

5.2-

4.7

8

0.4

Interorbital

width

lengths

Standard
Deviation

Hoof lengths
Front
Rear
Horn length

2.9

4.5-

1.2

9

1.6

Girth

74.6

83.8- 67.2

8

5.0

Neck
Point

a

30.0

32.7- 22.5

8

3.4

Point

b

41.4

47.8- 33.8

8

4.3

Bled

51.3

68.0- 37.4

8

10.0

Eviscerated

36.9

41.1- 29.0

6

4.7

Carcass

weights

(lbs)

�-15-

Table 2.

External body measurements for yearling buck antelope.

Measurement

Mean

Range

Sample
Size

Standard
Deviation

Shoulder height

86.7

95.0- 78.0

17

5.0

Total body length

134.4

147.0-112.7

17

10.2

Tail length

12.2

15.2-

8.2

17

2.4

Body length

122.2

134.2-102.5

17

9.9

Head length

27.6

29.6- 24.2

15

1.6

Interorbital width

12.8

14.1- 11.3

17

1.0

Ear length

14.5

17.2- 12.2

17

1.0

Hind foot length

43.7

45.5- 37.5

17

2.1

Front

6.8

7.9-

5.7

17

0.6

Rear

6.5

7.3-

5.3

17

0.6

Basal circumference

11.1

14.9-

5.7

16

2.1

Length

17.5

25.4-

4.3

17

4.5

Prong length

2.3

7.0-

0.0

17

2.2

Spread (tip to tip)

17.1

21.0-

6.8

15

4.9

91.2

106.0- 74.6

17

7.5

Point a

40.5

52.5- 22.5

14

8.6

Point b

52.7

63.2- 42.6

15

6.6

Bled

92

113

- 68

17

17.1

Eviscerated

73

88

- 58

11

12.1

Hoof length

Horns

Girth
Neck

Carcass weights (lbs)

�-16-

Table 3.

External body measurements for yearling doe antelope.

Measurement

Mean

Range

Sample
Size

Standard
Deviation

Shoulder height

90.0

96.7- 84.5

8

4.1

Total body length

139.0

151.0-126.0

7

7.9

Tail length

13.3

15.6- 10.7

9

1.6

Body length

124.2

136.9-113.6

5

8.7

Head length

27.3

30.1- 25.1

9

1.9

Interorbital width

12.6

13.2- 12.9

7

0.4

Ear length

14.4

16.5- 13.4

16

1.0

Hind foot length

37.9

45.4-

4.9

18

9.0

Front

6.4

7.7-

5.2

18

1.0

Rear

6.0

7.0-

4.7

18

0.7

Horn length

1.7

5.1-

0.1

18

1.9

Girth

87.5

74.0- 58.0

8

3.6

Point a

34.3

39.0- 30.7

7

3.2

Point b

47.2

53.3- 39.6

7

4.9

Bled

94.4

112.0- 79.0

9

11. 7

Eviscerated

68.4

79.0- 59.0

9

7.9

Hoof length

Neck

Carcass weights (lbs)

�-17-

Table 4.

External body measurements

(centimeters)

for mature buck antelope.

Standard
Deviation

Measurement

Mean

Range

Sample
Size

Shoulder height

90.9

97.0- 82.0

15

4.3

Total body length

139.1

146.0-132.8

15

3.7

Tail length

13.0

15.6- 10.5

15

1.4

Body length

126.1

l34.5-117.2

15

4.4

Head length

28.7

31.1- 24.7

15

1.6

14.2

14.7- 12.8

15

0.4

Ear lengths

14.6

17.2- 12.3

15

1.3

Hindfeet

41.8

45.5- 36.8

15

2.0

Front

6.6

7.5-

5.4

15

0.6

Rear

6.1

7.1-

5.3

15

0.5

Length

27.3

37.3- 18.1

15

4.7

Prong length

5.6

11.0-

0.0

15

2.6

Spread

(tip to tip)

16.8

25.7-

3.7

15

6.0

Spread

(maximum inside)

22.3

33.6- 16.0

15

4.3

14.4

20.5- 11.6

15

2.1

93.5

99.0- 85.7

15

4.5

Point a

38.6

49.0- 37.2

12

5.2

Point b

56.3

63.0- 47.5

13

5.0

Bled

111

123 - 89

15

10.4

Eviscerated

81

92

15

7.6

Interorbital

width

lengths

Hoof length

Horns

Basal circumference
Girth
Neck

Carcass weights

(lbs)

- 64

�-18-

Table

5.

External

body me as uremerrt s (centimeters)

for

mature

doe antelope.

Standard
Deviation

Measurement

Mean

Range

Sample
Size

Shoulder

height

88.9

98.0- 77.0

32

5.3

Total

body length

139.7

150.~-1l4.2

31

7.9

Tail

length

12.0

17.1-

7.4

32

2.3

. Body length

127.8

138.2-103.6

31

5.9

Head length

25.9

31.0- 26.3

31

1.1

11.1

13.7- 10.1

30

0.8

14.9

16.4- 13.4

32

0.6

41.8

45.0- 36.5

31

2.0

Front

7.0

8.8-

5.4

31

1.0

Rear

6.9

13.4-

5.4

31

1.4

Horn Lengths

2.5

6.3-

0.0

31

1.9

Girth

90.8

82.0- 98.2

30

4.7

Interorbital

width

Ear lengths
Hind feet

lengths

Hoof lengths

Neck
Point

a

34.1

40.4- 24.6

26

3.8

Point

b

49.1

55.7- 42.1

24

3.6

Bled

102

125

- 79

32

12.2

Eviscerated

70

86

- 63

31

7.1

Carcass

weights

(lbs)

�-19-

Table 6. Measurements (millimeters), weights (grams) , and volwnes (cubic
centimeters) of organs of fawn antelope approximately three to four months old.

Range

Sample
Size

Standard
Deviation

111
81
198

119 - 96
95 - 72
284 - 259

7
7
7

8.1
7.3
43.9

256
l30
31
541

284 - 222
160 - 116
38 - 23
1010 - 390

7
7
7
7

20.1
15.7
6.0
220.2

Organ

Mean

Heart
Longitudinal
Transverse
Weight
Liver
Longi tudinal
Transverse
Vertical
Weight
Brain
Longitudinal
Transverse
Vertical
Weight
Volume
Spleen
Longitudinal
Transverse
Vertical
Weight
Volume
Kidney
Longitudinal
Transverse
Vertical
Weight
Volume

1
1

121
70
43
49

58

29

7
1

11.7

71
45
30
56

81 53 49 102 -

64
30
11
41

14
14
14
14

5.6
5.3
10.2
15.8

2
2
2
2

0.3
0.1
0.4
0.0

6
6

35.4
61.1

Eyeball
Longitudinai
Transverse
Weight
Volume

30
34
17
17

Lung Weights
Left
Right

192
258

230 - 130
340 - 200

�-20-

Table 7. Measurements (millimeters), weights (grams), and volumes (cubic
centimeters) of organs of yearling buck antelope.

Organ

Mean

Range

Sample
Size

Standard
Deviation

Heart
Longitudinal
Transverse
Weight
Voltnne

131
103
348
348

153 - 103
122 - 79
497 - 236
467 - 224

12
12
12
8

14.5
13.2
84.9
89.1

Liver
Longitudinal
Transverse
Vertical
Weight
Volume

276
156
39
831
829

313 - 233
182 - 124
50 - 25
1430 - 462
1324 - 432

13
13
13
13
8

26.9
19.5
6.9
267.9
293.7

Brain
Longitudinal
Transverse
Vertical
Weight
Volume

76
66
39
108
104

83 78 42 127 122 -

68
59
37
91
88

5
5
4
7
7

5.9
7.3
2.6
13.6
12.9

Spleen
Longitudinal
Transverse
Vertical
Weight
Volume

134
92
12
77
72

155 - 120
109 - 70
14 - 19
13353
94 - 50

8
8
6
13
6

13.7
13.2
1.7
22.4
15.9

Kidney
Longitudinal
Transverse
Vertical
Weight
Volume

82
46
47
95
91

94 61 61 155 147 -

70
37
24
66
63

22
22
22
25
17

6.4
7.2
9.8
22.9
24.8

Eyeball
Longitudinal
Transverse
Weight
Volume

32
35
19
19

33 37 22 21 -

29
34
17
16

19
19
19
19

1.1
1.0
1.8
1.7

Lung Weights
Left
Right

262
394

370 - 180
563 - 240

8
8

55.8
106.0

�-21-

Table 8. Measurements (millimeters), weights (grams), and volumes (cubic
centimeters) of organs of yearling doe antelope.

Organ

Hean

Range

Sample
Size

Standard
Deviation

Heart
Longitudinal
Transverse
Weight
Volume

128
97
340
324

143 - 116
117 - 82
421 - 269
405 - 254

9
9
9
8

7.9
10.3
47.5
49.8

Liver
Longitudinal
Transverse
Vertical
Weight
Volume

280
151
42
825
757

302 - 249
173 - 136
55 - 34
1230 - 575
1152 - 535

8
8
8
8
7

17.8
12.4
6.6
202.0
202.0

Brain
Longitudinal
Transverse
Vertical
Weight
Volume

73
65
41
107
101

7774 47 122 108 -

69
58
37
97
93

6

7

3.1
5.9
3.4
8.6
6.1

Spleen
Longitudinal
Transverse
Vertical
Weight
Volume

138
94
12
72
67

192 - 117
105 - 78
17 9
105 - 56
99 - 53

9
9
7
9
8

23.5
8.3
3.0
17.7
17.6

Kidney
Longitudinal
Transverse
Vertical
Weight
Volume

78
42
47
83
76

95 48 56 120 114 -

72
35
37
63
60

18
18
18
17
15

6.1
3.8
5.1
16.0
15.5

Eyeball
Longitudinal
Transverse
~.iTeight
Volume

32
35
20
19

35 37 22 21 -

30
31
18
17

15
14
15
13

1.6
1.9
1.3
1.2

Lung Weights
Left
Right

205
388

268 - 155
580 - 245

5
5

50.2
137.0

6

6
8

�-22-

Table 9. Measurements (millimeters), weights (grams), and volumes (cubic
centimeters) of the organs of mature buck antelope.

Organ

Mean

Range

Sample
Size

Standard
Deviation

Heart
Longitudinal
Transverse
Weight
Voltnne

140
109
423
403

150 - 127
124 - 91
524 - 352
508 - 334

14
14
14
14

8.2
8.4
49.4
42.9

Liver
Longitudinal
Transverse
Vertical
Weight
Volume

320
185
48
1144
1067

335 - 274
197 - 150
54 - 37
1500
705
1392 - 655

14
14
14
14
14

18.6
13.6
5.6
218.9
203.8

Brain
Longitudinal
Transverse
Vertical
Weight
Volume

77
68
39
114
111

83 - 72
80 - 60
43 - 32
138- 100
13494

9
9
8
12
12

4.0
6.1
4.0
12.4
12.6

Spleen
Longitudinal
Transverse
Vertical
Weight
Voltnne

153
106
13
98
92

175 - 134
13691
15 - 10
122 - 80
116 - 76

15
15
11
15
15

12.4
15.1
1.3
16.5
15.5

Kidney
Longitudinal
Transverse
Vertical
Weight
Volume

85
48
53
114
112

96 60 63 178 171 -

71
37
45
86
82

15
15
15
15
15

8.0
5.9
4.8
30.7
29.3

Eyeball
Longitudinal
Transverse
Weight
Volume

33
36
21
21

37 38 24 23 -

32
33
20
19

14
14
14
14

1.3
1.2
1.3
1.7

Lung Weights
Left
Right

303
422

415 - 185
580 - 240

10
10

82.2
121.5

~"

�-23-

Table 10. Measurements (millimeters), weights (grams), and volumes (cubic
centimeters) of the organs of mature does.

Organ

Mean

Range

Sample
Size

Standard
Deviation

Heart
Longitudinal
Transverse
Weight
Volume

l33
101
355
340

151 - 115
ll3 - 87
430 - 270
404 - 273

29
29
29
24

9.2
6.8
40.2
32.8

Liver
Longitudinal
Transverse
Vertical
Weight
Voltnne

293
170
46
1013
982

324 - 249
201 - 143
55 - 25
1440 - 665
l343 - 615

25
25
25
25
20

22.0
18.6
22.2
224.7
214.1

Brain
Longitudinal
Transverse
Vertical
Weight
Volume

75
65
39
108
103

84 72 47 l33 129 -

68
37
31
85
82

16
16
15
19
19

4.0
8.4
5.4
12.5
14.5

Spleen
Longitudinal
Transverse
Vertical
Weight
Volume

146
100
11
82
80

168 - 125
122 - 79
16 7
113 - 20
106 - 64

24
24
16
29
24

13.0
12.1
2.5
19.0
12.7

85
47
50
104

99 56 59 289 149 -

76
39
27
74
71

30
30
30
30
25

5.3
5.6
8.7
39.1
20.5

Eyeball
Longi tudinal
Transverse
Weight
Volume

33
37
23
22

35 39 26 25 -

31
34
20
19

22
22
22
19

0.9
1.0
1.4
1.3

Lung Weights
Left
Right

248
362

340 - 180
500 - 260

16
16

86.4
100.8

Kidney
Longitudinal
Transverse
Vertical
Weight
Voltnne

III

�-24-

Table 11. Gland measurements (millimeters), weights (grams), and volumes
(cubic centimeters) for fawn antelope approximately three to four months
old.
Gland and
Measurement

Mean

Range

Thyroid (Left)
Longitudinal
Transverse
Vertical
Weight
Volume

25.1
11.8
5.0
0.9
1.2

26.5
12.6

Thyroid (Right)
Longitudinal
Transverse
Vertical
Weight
Volume

5.0
0.9
0.9

Sample
Size

Standard
Deviation

33.3 - 2.3
14.0 - 10.5
6.5 - 3.8
1.4 - 0.9

7
7
7
7
1

10.7
1.4
1.0
0.3

32.5 - 19.0
15.0 - 11.1
6.1 - 4.0
1.2
0.5

7
7

4.7
1.2
0.9
0.4

7

1

Adrenal (Left)I
Longitudinal
Transverse
Vertical
Weight
Voltnne

27.4
10.1
5.3
1.0
0.9

30.6 - 24.8
11.4 - 9.9
6.9 - 3.7
1.60.6

Adrenal (Right)
Longitudinal
Transverse
Vertical
Weight
Volume

26.9
11.0
4.8
1.0
1.0

35.5 - 20.2
17.5 - 7.5
6.7 - 3.2
1.70.6

Pituitary
Longitudinal
Transverse
Weight
Volume

7

7

2.6

7
7

0.8
1.4
0.4

7
1

7

5.4

7
7
7
1

3.4
1.2

0.4

�-25-

Table 12. Gland measurements (millimeters),weights
(cubic centimeters) for yearling buck antelope.

(grams), and volumes

Gland and
Measurement

Mean

Range

Sample
Size

Standard
Deviation

Thyroid (Left)
Longitudinal
Transverse
Vertical
Weight
Volume

35.6
11.7
7.2
1.7
1.6

43.8 - 27.9
16.7 - 9.9
9.4 - 4.1
2.5 - 1.6
2.3 - 1.7

8
8
8
8
7

5.5
2.4
1.5
0.5
0.4

Thyroid (Right)
Longitudinal
Transverse
Vertical
Weight
Volume

33.4
12.6
6.4
1.6
1.6

44.6 - 19.0
19.1 - 9.9
9.0 - 3.8
2.7 - 0.8
2.5 - 0.9

10
10
10
10
8

7.1
2.9
1.5
0.7
0.6

Adrenal (Left)
Longitudinal
Transverse
Vertical
Weight
Volume

33.4
11.1
5.2
1.2
1.2

38.7 - 26.8
16.9 - 9.1
7.1 - 2.6
2.1 - 0.9
2.0 - 0.9

13
13
13
13
9

4.2
2.1
1.9
0.5
0.4

Adrenal (Right)
Longitudinal
Transverse
Vertical
Weight
Volume

31.8
12.0
5.5
1.5
1.3

36.9 - 26.3
18.9 - 9.7
6.2 - 4.9
2.3 - 0.9
2.2 - 1.3

11
11
11
11
9

3.9
3.1
0.9
0.4
0.4

Pituitary
Longitudinal
Transverse
Weight
Volume

10.4
8.3
0.5
0.4

11.5 10.1 0.5 0.4 -

5
5
7
7

1.1
1.9
0.1
0.1

8.8
5.2
0.6
0.5

�-26-

Table 13. Gland measurments (millimeters), weights (grams) , and volumes
(cubic centimeters) for yearling doe antelope.
Gland and
Measurement

Mean

Range

Sample
Size

Standard
Deviation

Thyroid (left)
Longitudinal
Transverse
Vertical
Weight
Volume

34.0
11.9
6.9
1.5
1.4

38.0- 31.8
14.3- 9.5
8.7- 5.4
2.4- 0.9
2.0- 0.8

7
7
7
7
6

2.5
1.7
1.3
0.5
0.4

Thyroid (right)
Longitudinal
Transverse
Vertical
Weight
Volume

29.3
l3.9
6.4
1.5
1.4

40.3- 19.4
16.3- 11.4
8.6- 4.9
2.3- 1.0
2.1- 0.9

8
8
8
8
7

5.9
1.5
1.2
0.5
0.4

Adrenal (left)
Longitudinal
Transverse
Vertical
Weight
Volume

34.5
12.3
6.7
1.7
1.6

41.1- 28.8
16.0- 10.3
9.7- 5.1
2.4- 1.3
2.2- 1.2

8
8
7
8
7

4.3
2.1
1.6
0.3
0.3

Adrenal (right)
Longitudinal
Transverse
Vertical
Weight
Volume

28.9
13.8
6.3
1.4
1.3

34.7- 21.3
18.3- 9.1
8.7- 4.6
2.1- 1.4
1.4- 0.9

7
7
6
7
6

4.4
3.0
1.4
0.4
0.4

Pituitary
Longitudinal
Transverse
Weight
Volume

10.0
8.5
0.5
0.4

10.69.60.60.4-

6
6
7
6

0.6
0.8
0.2
0.2

8.9
7.1
0.3
0.2

�-27-

Table 14. Gland measurement (millimeters), weights
(cubic centimeters), for buck (mature) antelope.

(grams) , and volumes

Sample
Size

Standard
Deviation

Glands and
Measurements

Mean

Range

Thyroid (left)
Longitundinal
Transverse
Vertical
Weight
Volume

35.5
15.2
7.4
2.4
2.2

38.8- 28.5
19.8- 10.9
9.9- 5.2
3.2- 1.2
2.8- 1.0

13
13
13
l3
13

5.4
3.1
1.4
0.8
0.8

Thyroid (right)
Longitudinal
Transverse
Vertical
Weight
Volume

34.6
14.6
8.1
2.2
2.0

42.8- 25.5
19.0- 10.9
9.8- 4.2
3.2- 1.0
2.8- 0.9

12
12
12
12
12

5.8
4.5
1.4
0.6
0.6

Adrenal (left)
Longitudinal
Transverse
Vertical
Weight
Volume

35.0
13.4
7.1
2.0
1.8

42.5- 30.9
15.5- 10.3
8.9- 5.5
2.9- 1.5
2.7- 1.4

14
14
14
14
14

9.0
1.4
1.1
0.4
0.4

Adrenal (right)
Longitudinal
Transverse
Vertical
Weight
Volume

30.1
l3.3
7.5
1.8
1.6

34.8- 29.7
17.2- 10.3
9.8- 5.1
2.5- 1.3
2.3- 1.2

11
11
11
11
11

2.2
2.4
1.7
0.4
0.3

Pituitary
Lon gitudin al
Transverse
Weight
Volume

10.4
9.8
0.5
0.4

12.611. 50.80.7-

8
8
9
9

1.5
1.0
0.2
0.3

9.6
9.9
0.2
0.1

7

�-28-

Table 15. Gland measurements (millimeters) , weights (grams), and volumes
(cubic centimeters) for mature doe antelope.
Gland and
Measurement

Mean

Range

Sample
Size

Standard
Deviation

Thyroid (Left)
Longitudinal
Transverse
Vertical
Weight
Volume

41.4
16.1
7.6
2.8
2.6

71.9 - 31.5
23.8 - 11.5
11.9 - 4.4
6.0 - 1.3
5.8 - 1.5

27
27
27
27
24

9.6
3.2
1.8
1.1
1.1

Thyroid (Right)
Longitudinal
Transverse
Vertical
Weight
Volume

35.5
15.6
7.3
2.4
2.3

51.6 - 27.3
36.6 - 10.6
12.1 - 2.1
5.4 - 1.1
5.2 - 1.3

21
21
21
21
18

6.6
3.4
2.7
1.3
1.3

Adrenal (Left)
Longitudinal
Transverse
Vertical
Weight
Volume

36.7
12.9
6.6
2.0
1.9

46.5 - 28.3
19.0 - 10.0
8.8 - 2.0
3.9 - 1.3
3.7 - 1.2

27
27
27
27
22

4.3
1.4
1.6
0.4
0.3

Adrenal (Right)
Longitudinal
Transverse
Vertical
Weight
Volume

32.7
13.2
7.2
1.9
1.8

42.4 - 24.7
18.1
9.6
12.2 - 4.9
3.1 - 1.1
2.7 - 1.3

25
26
26
26
23

4.1
2.4
1.6
0.4
0.3

Pituitary
Longitudinal
Transverse
Weight
Volume

11.5
9.0
0.6
0.6

14.0 10.8 1.00.9 -

15
14
16

1.3
1.1
0.2
0.2

9.2
6.8
0.4
0.3

16

�-29Table 16. Measurements (millimeters), weights (grams), and volumes (cubic
centimeters) for the reproductive system of male antelope fawns (3-4 months
old).

Measurement

Mean

Range

Sample
Size

Standard
Deviation

Testes
Longitudinal
Transverse
Vertical
Weight
Volume

22.3
16.0
13.7
2.7
2.3

24.5 - 18.6
18.0 - 13.2
18.0 - 11.2
3.3 - 1.5
2.3 - 2.3

10
10
10
10
2

2.2
1.9
1.5
0.8

Prostate
Weight

4.1

11.8 -

0.3

3

7.7

Scrotum
Length
Width
Height

4.1
3.6
1.6

5.5 4.5 2.0 -

3.0
2.5
1.5

5
5
5

1.0
0.7
0.4

Penis
Length

11.0

13.6 - 10.5

5

1.4

Table 17. Heasurements (millimeters), weights (grams), and volumes (cubic
centimeters) for the reproductive system of female antelope fawns.
Sample
Size

Standard
Deviation

1.6
0.8

3
3

2.0
0.6

13.3 8.3 4.4 0.3 -

8.5
5.4
2.2
0.1

6

6

1.6
1.2
0.9
0.1

3.4
1.0

4.0 1.0-

2.8
0.9

6
6

0.6
0.1

9.5
1.5

10.5 1.8-

9.0
1.6

3
3

0.9
0.3

2.7
0.9
1.5
2.4
16.7

3.1 - 2.5
1.00.8
1.4
1.82.6 - 1.8
17.4 - 15.4

3
3
3
3
3

0.4
0.1
0.3
0.5
1.1

Measurement

Mean

Range

Mammary
Length
Depth

3.7
1.4

5.5 2.0 -

Ovary
Longitudinal
Transverse
Vertical
Weight

11.5
7.0
4.0
0.2

Uterine Body
Uterine Horns
Longitudinal
Transverse
Vagina
Longitudinal
Transverse
Cervix
Longitudinal
Transverse
Median Septum Length
True Length
Weight of Uterine Body

6

6

�Table 18. Means of measurements (millimeters), weights
reproductive system of buck antelope.

(grams), and volumes (cubic centimeters) for the

Measurement

Jan.

Feb.

Mar.

Apr.

May

Month
Ju1.
June

Aug.

Sept.

Oct.

Nov.

Dec.

Bucks Collected

2

2

2

2

2

2

3

2

1

2

1

2

Testes
Longitudinal
Transverse
Vertical
Weight
Volume

33.3
22.6
23.7
10.4
9.7

33.3
22.8
21.8
8.5
8.1

35.7
24.0
22.6
10.4
9.9

39.9
27.3
23.7
13.5
12.8

42.4
28.0
27.6
17.7
17.0

41. 7
28.4
26.3
16.4
15.8

41. 8
28.6
23.7
17.2
16.4

44.3
33.0
30.0
23.9
22.7

42.3
28.3
25.9
17.4
16.7

50.0
31.0
27.9
24.5
23.4

36.3
23.5
22.5
10.5
9.8

43.0
31.5
16.6
14.3
13.7

Prostate
Longitudinal
Weight
Volume

11.1
1.2
1.1

17.3
1.4
1.3

15.7
1.5
1.4

19.7
1.9
1.7

18.2
2.5
2.4

1.6
1.6

-

17.6
1.9
1.6

24.1
1.9
1.8

-

-

22.0
1.7
1.6

14.0
1.8
1.6

17.0
1.2
1.1

Scrotum
Length
Height
Width

42
48
38

40
51
42

64
47
30

50
48
45

53
53
38

60
53
33

58
53
38

58
55
58

60
55
60

70
50
43

60
45
35

45
53
40

Penis
Length

125

155

135

135

150

137

138

153

140

143

150

155

I
Vol

0
I

�Table 19. Measurements (millimeters) and weights (grams) for the reproductive tracts, ovaries, and
embryos from doe antelope collected in Moffat County.
Month
Ju1.
June

Aug.

Sept.

Oct.

Nov.

Dec.

2

2

7

2

2

2

16.3
13.2
8.1
1.0
0.1

16.9
13.2
8.1
1.1
0.1

13.1
10.9
7.6
0.7
0.1

15.9
13.0
7.6
1.1
0.1

18.4
14.2
10.2
1.5
0.1

16.8
13.4
9.6
1.2
0.1

18.6
13.8
l1.6
1.3
0.1

450
197

l17
58

75
14

55
14

61
21

67
18

103
46

125
62

180
26

195
30

151
28

153
32

150
33

135
34

145
30

155
29

158
30

90
18

102
12

87
25

55
20

50
15

61
15

60
15

65
l1

70
17

43
169
3203
433

45
85
230
170
5390 12565
795 l165

33
43

24
33

25
25

29
32

27
33

36
40
1249
153

684
242
332
248

1443
348
430
304

20
64

Measurement

Jan.

Feb.

Mar.

Apr.

May

Does Collected

1

2

2

1

1

2

Ovaries
Longitudinal
Transverse
Vertical
lveight
Volume (cc)

17.6
16.7
9.0
1.7
0.2

19.1
16.6
9.5
1.6
0.1

16.9
13.1
7.6
1.3
0.1

18.1
14.0
9.5
1.3
0.1

22.9
18.1
10.8
2.4
0.1

Uterine Horn
Longitudinal
Transverse

235
103

251
126

227
142

240
223

Vagina
Longitudinal
Transverse

165
20

175
20

189
23

Cervix
Longitudinal
Transverse

l10
15

93
18

63
19

Uterine Body
Median Septum Length
True Uterine Length
Weight with Young
lveight without Young

35
60
1590
270

60
113
2630
500

159
163
175

365
227
261
172

Embryos 1/
~]eight
Rump-Crown Length
Total Length
Shoulder Height

1/
- 22 embryos were measured.

-

3538

630
485

-

-

-

-

-

204

94

68

95

104

57
35
259
107

-

,-

-

-

-

4
35

I

W

.....
I

�Table 20. Measurements (millimeters) and weights (grams) for the reproductive tracts, ovaries, and
embryos from doe antelope collected in Saguache County.

Feb.

Mar.

Apr.

May

Month
June

Ju1.

Aug.

Sept.

Oct.

Nov.

Dec.

Measurement

Jan.

Does Collected

1

1

1

1

1

1

1

1

1

1

2

1

Ovaries
Longitudinal
Transverse
Vertical
Weight
Volume (cc)

16.4
11.6
8.9
0.9
0.1

16.1
15.1
8.3
1.2
0.1

16.6
11.8
9.0
1.3
0.1

16.4
13.2
9.6
1.6
0.2

23.7
15.4
9.4
2.0
0.2

12.3
11.4
6.5
0.6
0.1

15.6
l3.3
9.2
1.2
0.1

15.1
9.8
9.2
1.0

l3.1
8.7

-

15.1
11.6
7.4
1.0
0.1

0.8
0.1

16.7
13.6
9.9
1.6
0.2

21.2
16.4
9.2
2.0
0.2

Uterine Horn
Longitudinal
Transverse

153
80

232
119

219
104

145
79

335
183

91
25

70
32

74
14

53
12

44
14

72
23

165
78

I

-

I

W
N

Vagina
Longitudinal
Transverse

150
22

150
25

155
21

161
29

210
20

150
20

160
36

l34
25

140
32

132
29

149
19

140
20

Cervix
Longitudinal
Transverse

75
12

65
17

90
10

66
15

105
22

40
18

71
18

60
15

65
15

66
12

75
12

75
12

44
28
1055
150

42
104
1720

60
72
1345
215

42
83
1720
240

70
180
8765
920

32
36

23
25

27
29

20
15

26
16

32
23
88
81

49
28
725

99
137
164

102

186
174
203
123

605
243
400
198

2575

1.1
21.3

24.5
89.9

Uterine Body
Median Septum Length
True Uterine Length
Weight with Young
1.veightwithout Young
Embryos
Weight
Rump-Crown Length
Total Length
Shoulder Height

-

197

-

520
383

-

-

-

-

-

104

83

77

68

62

-

-

-

-

-

-

I

�-33Table 21.

Blood analysis

for buck antelope.

Mean

Erythrocytes

M/cu. mm

Leucocytes

per cu. mm

9.47

Range

12.25-

Sample Size

7.50

10

1830

4000 - 1100

10

Neutrophi1s
Segs/percent
Immature

29

32
4

- 16
0

9
9

Lymphocytes

46

66

- 24

9

18

32

2

9

6

16

0

9

2

0

9

Monocytes

- percent

- percent

Eosinophi1s

- percent

.4

Basophi1s

- percent

.2

Hemoglobin

'1,/100 m1

16.5

22.5 - 13.7

10

Hematocrit

- percent

49

58

10

Table 22.

Blood analysis

for doe antelope.

Mean

Erythrocytes

M/cu. mm

Leucocytes

per cu. mm

- 42

9.48

Range

12.65-

Sample Size

5.60

19

1756

3100 - 570

19

Neutrophi1s
Segs/percent
Immature

40
0.2

56
3

- 20
0

16
16

Lymphocytes

46

65

- 40

17

14

36

0

17

2

8

0

17

0

17

Monocytes

- percent

- percent

Eosinophils

- percent

Basophils

- percent

0.1

1

Hemoglobin

g/100 m1

16.5

29.0 - 11.0

19

Hematocrit

- percent

49

75

19

- 33

�-34-

Table 23.

Stomach weights

Weights

(grams) for antelope.

Mean

Range

Sample
Size

2469

3470 - 1510

7

748

2103
450

3020 - 1090

7
1

749

4638

5150 - 4330

7

322

3968
646

4600 - 3430
730 - 550

7
4

425
95

4725

6460 - 3250

6

1291

3907
608

5680 - 2405
755 - 515

6
6

1250
109

5678

7030 - 3970

14

,905

4910
838

6260 - 3540
1020 - 670

14
13

759
145

4995

9620 - 2670

30

1610

4171
852

8190 - 2300
1300 - 530

30
25

1459
252

Standard
Deviation

Fawns
Entire stomach and
contents
Rumen and reticulum
and contents
Stomach without contents
Yearling males
Entire stomach and
contents
Rumen and reticulum
and contents
Stomach without contents
Yearling females
Entire stomach and
contents
Rumen and reticulum
with contents
Stomach without contents
Mature males
Entire stomach and
contents
Rumen and reticulum
with contents
Stomach without contents
Mature females
En tire stomach and
contents
Rumen and reticulum
with contents
Stomach without contents

�-35-

Table 24.

Mean body temperatures (degrees centigrade) of antelope.

Time
(hrs.)

Mean

Range

Sample

Muscle

38.7

41.0 - 37.5

48

Rectal

38.4

40.5 - 37.0

49

1

3.4

5.0 -

1.0

18

2

5.5

8.5 -

1.5

14

3

11.0

12.5 -

5.0

13

4

11.3

15.5 -

9.0

7

5

11.4

17.0 -

7.5

7

6

13.4

18.0 - 10.0

5

7

16.9

22.0 - ll.O

6

8

18.7

30.0 - 12.0

5

At death

Degrees decrease in muscle
temperature after evisceration

9
10

14.0

16

19.2

17

20.0

1

18

22.3

2

20

25.0

1

1

25.5 - 12.5

3

��January,

-37-

JOB FINAL REPORT

State of

COLORADO
------~~~~~-------------

Project No.

W-40-R-13

Work Plan No.

1

Job Title:

Job No.

Investigations
2

Food Habits of Antelope

Period Covered:
Personnel:

Antelope

May 1, 1965 through May 31, 1972

George Do Bear, James Olterman,

Jerrry Robinson.

ABSTRACT

Study areas were established on three range types in Colorado, (a) bitterbrush,
(b) sagebrush and (c) rabbitbrush.
A total of 76 antelope rumen samples
were obtained from these areas at monthly intervals for a one-year period and
from hunter harvested animals.
Also, seasonal and daily feeding activity
patterns were established for antelope on the study areas.
There were seasonal variations in the antelope diets, and the diets varied
between range-types.
Browse plants were the major food items eaten, forbs
secondary and 'grasses minor.
Forbs increased in the diet and browse decreased
during the spring and summer periods.
Forbs were more important in the
antelope diet on the sagebrush-type than on the other areas.
Shrubs in the
samples from the bitterbrush range did not decrease during the summer to the
extent they decreased in the rumen samples from the other ranges.
Vegetation
surveys indicated antelope selectively sought forbs to eat. A few plants
were selected over others, but in general a large variety of plant species
were eaten in relative proportion to their availability on the range.
There were seasonal variations in the daily feeding pattern of antelope.
They
were most active in early morning and late evening hours.
Feeding activity
increased in the fall-winter period, showing less fluctuation during mid-day.

1973

��-39-

FOOD HABITS OF ANTELOPE
George D. Bear

P. S. OBJECTIVE
To determine

antelope

forage preferences

LITERATURE

and use.

REVIEW

Information concerning food preferences and plants eaten by herbivores
are of major importance when managing rangelands.
Year-long data is
needed since feeding behavior varies throughout the year. Hoover, Till,
and Ogilvie (1959) studied the food habits of antelope in the plains
region in eastern Colorado with limited work in the mountain bunchgrass
type and the intermountain shrub type. Percent composition of grasses
in the rumen samples collected in the shortgrass plains was: winter, 4.6
percent; spring, 20.3 percent; summer, 1.3 percent; and fall 1.3 percent.
Forbs comprised the following portion of the diet: winter, 26.0 percent;
spring, 50.5 percent; summer, 65.6 percent; and fall 22.1 percent.
Comparable figures for browse were: winter, 54.2 percent; spring, 24.7
percent; summer, 22.3 percent; and fall, 71.6 percent.
Cacti comprised
15.1, 4.3, 10.7, and 5.0 percent of the rumen samples during the winter,
spring, summer, and fall seasons, respectively.
Rumen samples from
August to September only were analyzed for the South Park, Colorado
mountain bunchgrass type. Shrubs were the more important forage class
eaten, then forbs, next grasses.
Cacti were eaten only in minor amounts.
Hoover et ale (1959) also examined rumen samples collected in September
on the mountain brush type and intermountain desert shrub types in Moffat
County, which are comparable to the bitterbrush and sagebrush ranges used
for this study, respectively.
Browse made up 93.7 percent of the diet
in the mountain brush area, followed by forbs, grasses, and cacti in
respective order of abundance.
For the intermountain desert shrub type
forbs were most important (60.1 percent), followed by shrubs, cacti, and
grasses.
Antelope food habits studies in Utah, Nevada, Oregon, Montana and Wyoming
indicated similar patterns in the antelope diets (Beale and Scotter 1968;
Mason 1952; Cole and Wilkins 1958; Severson, May, and Hepworth 1968;
and Bayless 1968). Browse was the most important forage class in the
winter and fall, with forbs increasing in the diet during the spring and
summer months.
Grasses were minor food items in the antelope diets.
Mason (1952) listed Artemisia sp., Purshia tridentata, Chrysothamnus sp.,
Phlox sp., Polygonum sp., Iva axillaris, Trifolium sp., Erigeron austinae
and Bromus tectorum,as
the more important plants eaten by antelope in
Nevada-Oregon.
Severson et ale (1968) listed Chrysothamnusviscidiflorus
and Artemisia tridentata as major food items in Wyoming.
Bayless (1968)

�-40-

listed Artemisia tridentata, Chrysothamnus nauseosus, Opuntia polycantha,
and Artemisia frigida as important food items in Montana.
There were deviations in the dietary pattern mentioned above. Cole and
Wilkins (1958) found browse to be the major food item in the fall, winter,
and spring diets with forbs being the major item in the summer diet.
This occurred on the silver sagebrush-grassland
and the big sagebrushgrassland range types. However, on grassland range forbs were the most
important food item in the spring, summer, and fall; while browse was
most important in the winter diet. Beuchner (1950) found forbs to be
the most important forage class throughout the year in Texas, followed
by browse.
Grasses were of minor importance but its consumption was
highest in the autumn when plants were actively growing.
In Kansas
antelope consumed a larger amount of grasses and forbs than browse
(Hlavachick 1966). The area occupied by antelope is primarily grassland
with little shrub cover. Opuntia macrorhiza was also a major food item
during the summer months.
Russel (1964) found forbs were of major importance in antelope diets in New Mexico, browse was secondary, and grasses
were taken in minor amounts.
This pattern varied in some of the different areas under study. Therefore, browse was not the major forage class
in the diets of antelope in Kansas, New Mexico, and Texas that it was in
the other states cited above.
Several authors found antelope to be most active in early morning and
late evening hours (Kautz 1942, Prenzlow 1968, Beuchner 1950, and
Einarsen 1948). Kautz (1942) reported there was an extended period of
resting or lying down during the warmer part of the day; lasting just a
few hours in April, but extending from 8 or 9 A.M. to about 5:00 P.M.
during July.
Beuchner (1950) reported the following average daily percentages of feeding time for antelope: autumn - 72.8 percent, 77.8 percent for winter, 63.0 percent in spring, and 67.0 percent for summer.
He commented that antelope on poor ranges spent more time feeding than
. antelope on good ranges. Also, antelope spent more time feeding in the
winter period because food was scarcer and the daylight hours were
shorter.

METHODS AND MATERIALS

Rumen Analysis
Antelope were collected at monthly intervals in Moffat and Saguache
counties during the period from June 1, 1965 to April 30, 1967 (one
year in each locality).
Three animals were collected each month in
Moffat County, where two were taken on a rangeland characterized by
sagebrush (Artemisia tridentata) and the third on a rangeland characterized by bitterbrush (Purshia tridentata).
Two animals were collected
each month in Saguache County on rangeland characterized by a rabbitbrush (Chrysothamnus sp.) overstory.
Additional rumen samples

�-41-

were obtained from the bitterbrush rangeland during the hunting season
in September.
Two-quart rumen samples were taken from each antelope
and preserved in formalin.
In the laboratory rumen samples were washed through graduated sieves
and air dried on blotters.
These samples were hand sorted (using
forceps) to individual plant species.
Plant species were identified
with the use of a binocular microscope and reference specimens collected
in each area, identified, pressed, dried, and placed on cards. Plants
were weighed to the nearest gram and the percent composition for each
species computed for each rumen sample.

Vegetation

Surveys

Vegetation surveys were conducted in the locality where each antelope
was collected to determine plant availability to the animal and the
relative abundance of each plant species present.
A temporary linetransect was run in each locality with 50 circular plots (three square
feet in size) established on each transect.
The plots were placed at
125-foot intervals.
Percent crown coverage was recorded for each plant
species occurring on each plot. Percent coverage was determined by
visual projection of the plant canopy onto the plane surface of the
ground and related to the total area within the plot.

Feeding Activity
An effort was made to determine seasonal and daily fluctuations in
:.antelope feeding activity.
Antelope activities were classified into
two broad categories: (1) inactive or lying down, and (2) active or
feeding.
Activity of all undisturbed antelope encountered on the study
areas was recorded when the animals were first observed.
Time-of-day
and date were recorded with each observation.

RESULTS

Rumen Analysis

and Vegetation

Surveys

A total of 76 rumen samples were analyzed.
They were grouped into three
seasonal categories: (1) spring (April-May-June), (2) summer (July-AugustSeptember), and (3) fall-winter (October through March).
These categories
were based on the general phenology of the plants in the area. Spring was
the period during which plants were actively growing; summer the period
when plants were maturing and developing seeds; and fall-winter the dormant, curing-out period.

�-42-

Sagebrush

Rangeland

Spring--Forbs were the most abundant food item in the diet during spring.
They comprised 64 percent of the diet, shrubs 34 percent and grasses 2
percent (Table 1). Vegetation surveys indicated shrubs were most important (59 percent); then grasses (32 percent), and forbs least abundant
(9 percent) (Table 2). Forbs most abundant in the diet were: Cilia
laxiflorus, Caura sp., Sphaeralcea coeeirtea; Erigeron sp.,Chertop6dium
sp., Astragalus sp., Cirsium sp., and Eriogortum sp. Artemisia tridentata,
Chrysothamnus viscidiflorus, Artemisia frigida, and Purshia tridentata
were the more abundant shrubs in the diet. Three species of grasses
were eaten in trace amounts.
In general, plant species occurred in the spring rumen samples in the
relative proportion they were available on the range. A wide variety
of forbs and shrubs were available and many were eaten by antelope.
However, numerous species of grasses were available but only three
species were consumed.
Another discrepancy was that Caura sp. and Cilia
laxiflorus were very important food items but were not encountered on the
vegetation surveys.

Summer--Forbs were the most important class of vegetation in the summer
diet. They comprised 79 percent of the diet, shrubs 20 percent, and
grasses 1 percent (Table 3). Vegetation surveys indicated shrubs were
the most abundant class of plants in the area (64 percent), followed by
grasses (23 percent), and forbs (12 percent) (Table 4). The most important
forbs in the rumen samples were Cilia laxiflorus, Astragalus desperatus,
Sphaeralcea coccinea, Lygodesmia sp., Catira sp., and Erigeron sp. Purshia
tridentata was the most abundant shrub in the diet. Bromus tectorum and
Stipa comata occurred in most samples, but comprised very little of the
diet.
In general, there was a wide variety of plants available and many of them
were eaten by antelope.
Numerous grasses were present but the antelope
fed only on two species.
Cilia laxiflorus was a major food item, but was
not encountered on the vegetation survey; and Astragalus desperatus was a
major food item but was minor in forage abundance.
Artemisia tridentata
was the major shrub encountered on the vegetation surveys, however, Purshia
tridentata was the most important shrub in the diet. Other plants occurred
in the diet in relative proportion to their availability.

Fall-Winter--Shrubs
were the major class of vegetation in the fall-winter
diet. They comprised 90 percent of the diet, forbs 9 percent, and grasses
1 percent (Table 5). The vegetation surveys indicated shrubs were the
most abundant forage class on the area (85 percent), followed by grasses
(12 percent), with forbs least abundant (2 percent) (Table 6). The more
abundant shrubs in the rumen samples were: Artemisia tridentata, Artemisia
cana, and Atriplex canescens.
The more important forbs were: Erigeron sp.,

�-43-

Eriogonum sp., Chenopodium sp., Gaura sp., Leptodactylon pungens, and
Sphaeralcea coccinea.
Agropyron sp., Bromustectorum,
andStipa
comata
were the grasses identified in the rumen samples.
Plant species generally occurred in the same relative abundance in the
rumen samples as on the vegetation surveys.
Artemisia tridentata was
the most important item in the diet (69 percent) and was the most
abundant species encountered on the range surveys (64 percent).
Other
species of plants made up minor portions of the diet as well as the
overall available forage on the range. Several species of grasses
were represented in the range surveys, and three species were eaten
by antelope.

Bitterbrush

Rangeland

Spring--Shrubs were the most abundant class of forage in the spring
diet of the antelope on this range-type.
They comprised 68 percent
of the diet, forbs 23 percent, and grasses 6 percent (Table 7).
The vegetation surveys indicated shrubs were the most abundant forage
class (75 percent), followed by grasses (21 pe r cent ) , and forbs (4 percent) (Table 8). The more important shrubs in the rumen samples were:
Artemisia cana, Purshia tridentata, and Artemisia frigida.
The most
abundant forbs in the diet were: Cleome sp., an unidentified legume, and
Eriogonum sp. Bromus tectorum was the most abundant grass in the samples.
There was some correlation between the availability of plant species on
the range and their relative abundance in the rumen samples.
Bromus
tectorum was the more abundant grass in the diet and on the rangeland.
Artemisia cana and Purshia tridentata were important species in the diet
and on the area. However, a large variety of forbs occurred in trace
amounts in the diet but only a few species were represented on the range
surveys.
Also, Chrysothamnus species were abundant; in the area, but had
relatively low occurrence in rumen samples.

Summer--Shrubs were a major forage class in the summer diet of antelope.
They comprised 84 percent of the diet, forbs 14 percent, and grasses 1
percent (Table 9). The vegetation surveys indicated shrubs were most
abundant (76 percent), followed by grasses (16 percent), and forbs (7 percent) (Table 10). The most abundant shrubs in the rumen samples werePurshia
tridentata and Artemisia cana. The most important forbs in the samp~es
were: Salsola kali, Psoralea sp., Erigeron sp., Chenopodium sp.,
Eriogonum sp., and Polygonum sp. Several species of grasses occurred in
trace amounts, but Bromus tectorum occurred in the rumen samples with
greater frequency.
In general, the plant species occurring in the diet were common in the
area. Artemisia cana and Purshia tridentata were the most common plants
in the area and in the diet. However, Opuntia was common in the area, but
was not eaten by the antelope.
Salsola kali was the most abundant forb
in the rumen samples, yet it was not encountered on the vegetation surveys.

�-44-

Fall-Winter--Shrubs
were the most abundant class of plants in the fallwinter antelope diet. They comprised 96 percent of the diet, forbs 3
percent, and grasses 1 (Table 11). The vegetation surveys indicated
shrubs were most abundant in the area (86 percent), followed by grasses
(13 percent), and forbs (1 percent) (Table 12). The most important shrubs
in the diet were: Purshia tridentata, Artemisia tridentata, and Artemisia
cana. Erigeron sp., Leptodactylon pungens, Stipa comata and Bromus
tectorum were the most common forbs and grasses eaten, but only comprised
a minor portion of the diet.
There were some correlations between plant availability and relative
abundance in the diet. Purshia tridentata, Artemisia cana, Artemisia
tridentata, Stipa comata, and Bromus tectorum were common in the area
and were the more common plants in the antelope rumen samples.
However,
Chrysothamnus species and Opuntia sp. were fairly common to the area,
but eaten only in trace amounts.

Rabbitbrush

Rangeland

Spring--Shrubs were the most abundant forage class eaten by antelope in
the spring.
They comprised 67 percent of the diet, forbs 31 percent, and
grasses 1 percent (Table 13). Vegetation surveys indicated grasses were
most abundant in the area (61 percent), followed by shrubs (28 percent),
and forbs (3 percent) (Table 14). Artemisia frigida was the most abundant
species in the diet (37 percent), followed by Opuntia sp. (7 percent),
Eriogonum sp. (5 percent), and Eriogonum alatum (4 percent).
A wide variety of forbs, shrubs, and grasses were consumed in lesser amounts.
In general, plants were consumed in amounts relative to their occurrence
on the range, with two exceptions.
Bouteloua gracilis was the most
common plant species on the area, but it occurred only in trace amounts
in the rumen samples.
The reverse was true for Artemisia frigida.

Summer--Shrubs were the major class of vegetation in the antelope summer
diet. They comprised 53 percent of the diet, forbs 44 percent, and
grasses 1 percent (Table 15). The vegetation surveys indicated grasses
were most abundant (60 percent), then shrubs (22 percent), and forbs (9
percent) (Table 16). The more abundant shrubs in the diet were:
Symphoricarpos sp., Artemisia dracunculus, Rhus trilobata, Opuntia sp.
and Artemisia frigida.
The most important forbs were: an unidentified
composite, Sphaeralcea coccinea, Eriogonum sp., Chenopodium sp., and
Erysimum aspermum.
Bouteloua gracilis was eaten in minor amounts, but
occurred in most rumen samples.
There was general correlation between the quantity of a given plant
species in the rumen samples and its relative abundance in the area,
however, there were several exceptions.
Artemisia dracunculus,

�-45-

Symphoricarpos sp., and Rhus trilobata were important food items, however, they were not common in the area. Boutelouagracilis
was the
most common plant species in the area, but was a minor food item.

Fall-Winter--Shrubs
were the most abundant class of plants in the rumen
samples, comprising 94 percent of the total.
Forbs made up 3 percent,
grasses 1 percent, and miscellaneous items 1 percent (Table 17). The
vegetation surveys indicated grasses were most abundant (50 percent),
then shrubs (34 percent), next forbs (9 percent), and miscellaneous
items least abundant (6 percent) (Table 18). Opuntia sp., Artemisia
frigida, Chrysothamnus nauseosus, Atriplex sp., and Artemisia dracunculus were the most common food items. Most plants appeared in the
diet in the same relative proportions they occurred on the area, however, Bouteloua gracilis and Chrysothamnus viscidiflorus were minor
food items, but were fairly common in the area.

�-46-

Table 1. Analysis of three antelope rumen samples collected on the
sagebrush study area in Moffat County during the period April-June.

Plant Species

Percent
Frequency

Percent
Composition

67
33
67

T

Grasses
Bromus tectorum
Festuca ovina
Stipa comata
Unid. grass fragments

T
T

2
Sub-total

2

Forbs
Agoseris sp.
Astragalus sp.
Campanula sp.
Chenopodium sp.
Cirsium sp.
Erigeron sp.
Eriogonum sp.
Gaura sp.
Gilia laxiflorus
Lappula redowski
Lepidium sp.
Leptodactylon pungens
Penstemon sp.
Solanum nigrum
Sphaeralcea coccinea
Trifolium gymnocarpum
Zygadenus elegans
Unid. forb fragments

33
67
33
67
33
33
100
100
100
33
33
33
33
33
67
33
33

T

1
T

3
1

4
T

11
22
T
T

T
T
T

7
T
T

13
Sub-total

64

Shrubs
Artemisia cana
Artemisia frigida
Artemisia tridentata
Chrysothamnus viscidiflorus
Juniperus scopulorum
Purshia tridentata
Rhus trilobata
Unid. shrub fragments

T

33
33
67
100
33
67
33

1

24
2
T

1
T

5

Sub-total

34

Miscellaneous
Insects
Antelope hair

T
T

33
33
Sub-total

Total

T

100

�-47-

Table 2. Summary of the vegetation surveys on the sagebrush study area
in Moffat County during the period April-June.

Plant Species

Percent
Frequency

Percent
Composition

30
23

3
2
2
3
T
1
4
T
T
2
3
12

Grasses
AgroEyron sp.
AgroEyron smithii
Boute1oua gracilis
Bromus tectorum
Carex sp.
Festuca sp.
Koe1eria cristata
Muh1enbergia torreyi
Oryzopsis hymenoides
Poa sp.
Sitanion hystrix
StiEa comata

7

14
1
27
53
2
1
24
37
61
Sub-total

32

Forbs
Agoseris sp.
Allium sp ,
Antennaria sp.
Asc1eEias sp.
Erigeron sp.
Eriogonum sp.
LaEEu1a redowski
LeEtodacty1on Eungens
Lesguere11a montana
Linum 1ewisi
LUEinus sp.
Oenothera trichoca1yx
OxytroEis sp.
Penstemon sp.
Phlox sp.
PseudocymoEterus montanus
SEhaera1cea coccinea
Trifolium gymnocarEum
Unidentified forbs

2
19
16
1
16
23
22
25
13
11
8
2
2
2
32
13
21
18

T
T
T
T
1
1
1
2
T
T
T
T
T
T
2
T
1
T
1
Sub-total

9

Shrubs
Artemisia frigida
Artemisia tridentata
Atrip1ex canescens
Atrip1ex confertifo1ia
Chrysothamnus nauseosus
Chrysothamnus viscidif10rus
Eurotia 1anata
Gutierrezia sarothrae
0Euntia sp.
Purshia tridentata
'I'e
t radvmf a canescens

5
75
4
2
3
20
1
30
17
9
1

T

40
3
1
1
4
T
2
4
4
T
Sub-total

59

Total

100

�-48-

Table 3.
sagebrush

Plant

Analysis of five antelope rumen samples collected on the
study area in Moffat County during the period July-September.

Species

Percent
Frequency

Percent
Composition

100
60

1
T

Grasses
Bromus tectorum
Stipa comata

Sub-total

1

Forbs
Astragalus desperaterus
ChenoEodium
sp.
Cryptantha jamesii
Erigeron sp.
Eriogonum sp.
Gaura sp.
Gilia laxiflorus
Lappula sp.
Lepidium sp.
Lesquerella montana
LYl?;0desmia sp.
Mertensia sp.
Polygonum sp.
Psoralea. sp.
sp.
Pseudoc~oEterus
Salsola kali
Sisymbrium sp.
SEhaeralcea coccinea
Tral?;°E°l?;°n
Trifolium g~nocarpum
Zygadenus elegans
Unid. ComEositae
Unid. Cruciferae
Unid. forb fragments

21
1
T
1
T
2
34

60
40
20
80
40
60
100
60
20
20
20
20
20
20
20
20
20
80
20
20
20
40
40

T

T
T
2
T
T

1
T
T
T
2
1
T
T
T
T

14

Sub-total

79

Shrubs
--Artemisia cana
Artemisia dracunculus
Artemisia tridentata
Chrysothamnus viscidiflorus
Purshia tridentata
Unid. shrub fragments

T
1
T
T
18
1

20
60
60
40
100

Sub-total

20

Miscellaneous
Insects
Antelope

hair

T
T

20
20

Sub-total

Total

T

100

�-49-

Table 4. Summary of the vegetation surveys on the sagebrush
in Moffat County during the period July-September.

Plant Species

Percent
Frequency

study area

Percent
Composition

Grasses
Agropyron sp.
Agropyron spicatum
Agropyron ttachycaulum
Bouteloua gracilis
Bromus tectorum
Carex sp.
Koeleria cristata
Oryzopsis hymenoides
Sitanion hystrix
Stipa comata

4
19
32
6
21
7
40
18
46
56

T
4
2
T
2
T
2
2
4
7

Sub-total

23

Forbs
Allium sp.
Arenaria fendleri
Aster sp.
Astragalus sp.
Chenopodium sp.
Chrysopsis villosa
Erigeron sp.
Eriogonum a1atum
Eriogonum umbe11atum
Erysimum asperum
Lappu1a sp.
Linum 1ewisi
Lesquere11a montana
Lupinus sp.
Lygodesmia sp.
Oenothera trichoca1yx
Penstemon sp.
Phlox sp.
Sa1so1a ka1i
Sphaera1cea coccinea
Trifolium gymnocarpum
Unidentified forbs

15
1
1
1
3
1
17
19
24
1
1
1
1
3
1
1
1
15
1
18
12

1

T
T
T
T
T
1
2
2
T
T
T
T
T
T
T
T
1
T
1
1
3

Sub-total

12

Shrubs
Artemisia frigida
Artemisia tridentata
Chrysothamnus sp.
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Eurotia 1anata
Gutierrezia sarothrae
Leptodacty1on sp.
Opuntia sp.
Purshia tridentata
Tetradymia canescens

5
83
1
3
24
4
22
15
25
7
9

T
51
T
T
5
T
2
1
2
2
1

Sub-total

64

Total

99

�Table 5. Analysis of eleven antelope rumen .samples collected on the
sagebrush study area in Moffat County during the period October-March.

Plant species

Percent
Frequency

Percent
Composition

18
27
27

T
T
T

Grasses
Agropyron sp.
Bromus tectorum
Stipa comata
Unid. grass fragments

1

Sub-total

1

Forbs
Chenopodium sp.
Cryptantha jamesii
Erigeron sp.
Erigonum sp.
Gaura sp.
Gilia laxiflorus
Koshia sp.
Leptodactylon pungens
Oxytropis sp.
Phlox sp.
Polygonurn sp.
Salsola kali
Sphaeralcea coccinea
Unid. forb fragments

27
9

T

36

T

45

T

18
9

T

9

T

27
9
9

1
T
1
T

1

1

9

18
27

T

1
4

Sub-total

9

Shrubs
Artemisia cana
Artemisia dracunculus
Artemisia frigida
Artemisia tridentata
Atriplex canescens
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Opuntia sp.
Purshia tridentata
Unid. shrub fragments

18
18
18
82

11

36

3
T

1
T

69

27
64
18
27

1
T
T

5

Sub-total

90

Miscellaneous
Insects
Antelope hair
Rocks

55

9

T
T

9

T

Sub-total

Total

T

100

�-51-

Table 6. Summary of the vegetation surveys on the sagebrush
in Moffat County during the period October-March.

Plant Species

study area

Percent
Frequency

Percent
Composition

20
14
1
3
14
8
14
44

2
1
T
1
1
1
1
5

Grasses
Agropyron sp.
Agropyron smithii
Agropyron trachycaulum
Bromus tectorum
Koeleria cristata
Oryzopsis hymenoides
Sitanion hystrix
Stipa comata

Sub-total

12

Forbs
Antennaria sp.
Aster sp.
Chenopodium sp.
Erigeron sp.
Eriogonum sp.
Leptodactylon pungens
Phlox sp.
Salsola kali
Sphaeralcea coccinea
Unidentified forbs

3
1
1
1
3
4
6
1
2

T
T
T
T
T
T
1
T
T
1

Sub-total

2

Shrubs
Artemisia cana
Artemisia frigida
Artemisia tridentata
Atriplex canescens
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Eurotia lanata
Gutierrezia sarothrae
Opuntia sp.
Purshia tridentata
Tetradymia canescens

3
2
80

T
T
64
6
1
9
T
2
3
T
T

5

5
23
1
12
16
1
2

Sub-total

85

Total

99

�-52-

Table 7. Analysis of five antelope rumen samples collected on the
bitterbrush study area in Moffat County during the period April-June.

Plant Species

Percent
Frequency

Percent
Composition

Grasses
Bromus tectorum
Festuca ovina
Oryzopsis hymenoides
Stipa comata
Unid. grass fragments

5

100
20
20
20

T
T
T
1

Sub-total

6

Forbs
Agoseris sp.
Astragalus sp.
Chenopodium sp.
Cleome sp.
Erigeron sp.
Eriogonum sp.
Gaura sp.
Lappula redowski
Lepidium sp.
Leptodactylon pungens
Lupinus sp.
Lygodesia sp.
Oenothera sp.
Polygonum sp.
Salsola kali
Sisymbrium sp ,
Sphaeralcea coccinea
Zygadenus elegans
Unid. Compositae
Unid. legume
Unid. forb fragments

T
T
1

40
20
40
20
40
80
20
20
20
20
20
20
20
40
40
20
20
20
20
20

9
1
T
T
T
T
T
T
T
T
T
T
T
T
T
T

6
6

Sub-total

23

Shrubs
Artemisia cana
Artemisia dracunculus
Artemisia frigida
Artemisia tridentata
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Purshia tridentata
Unid. shrub fragments

27

60
80
80
40
60
40
100

1

5
T
2
T

16
17

Sub-total

68

Miscellaneous
Insects
Rocks
Antelope

hair

T
T
T

60
20
40

Sub-total

Total

T

100

�-53-

Table 8. Summary of the vegetation surveys on the bitterbrush
area in Moffat County during the period April-June.

Plant Species

study

Percent
Frequency

Percent
Composition

14
96
2

19

Grasses
Agropyron sp.
Bromus tectorurn
Oryzopsis hymenoides
Sitanion hystrix
Stipa comata

T

2

T
T

42

2

Sub-total

21

Forbs
Chrysopsis villosa
Delphinium sp.
Lappula sp ,
Phlox sp.
Unidentified forbs

4
8
2
2

T
T
T
T
4

Sub-total

4

Shrubs
Artemisia cana
Artemisia frigida
Artemisia tridentata
Chrysothamnus viscidiflorus
Chrysothamnus nauseosus
Opuntia sp.
Purshia tridentata

24
8
2
60
14
52
56

14
T
2
12
9
5
33

Sub-total

75

Total

100

�-54-

Table 9. Analysis of twenty-one antelope rumen samples collected on the
bitterbrush study area in Moffat County during the period July-September.

Plant Species

Percent
Frequency

Percent
Composition

5
33
5
19

T
T
T
T
1

Grasses
Agropyron sp.
Bromus tectorum
Oryzopsis hymenoides
Stipa comata
Unid. grass fragments

Sub-total

1

Forbs
Astragalus sp.
Chenopodium sp.
Cryptantha jamesii
Erigeron sp.
Eriogonum sp.
Gaura sp.
Lappula redowski
Lupinus sp.
Lygodesmia sp.
Oxytropis sp.
Penstemon sp.
Polygonum sp.
Psoralea sp.
Rumex sp.
Salsola kali
Senecio
Sisymbrium sp ,
Sphaeralcea coccinea
Verbena sp.
Unid. Cruciferae
Unid. forb fragments

sP:-

T
T
T
T
T
T
T
T
T
T
T
T
2
T
6
T
T
T
T
T
6

5
29
10
52
29
5
10
5
5
5
10
29
57
10
71
5
5
5
5
5

Sub-total

14

Shrubs
Artemisia cana
Artemisia dracunculus
Artemisia frigida
Artemisia tridentata
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Eurotia lanata
Purshia tridentata
Unid. shrub fragments

21
1
1
T
3
2
T
44
12

86
24
24
10
48
62
5
95

Sub-total

84

Miscellaneous
Antelope

hair

10

T

Sub-total

T

Total

99

�-55-

Table 10. Summary of the vegetation surveys on the bitterbrush study
area in Moffat County during the period July-September.

Plant Species

Percent
Frequency

Percent
Composition

Grasses
Agropyron trachycaulum
Bouteloua gracilis
Bromus tectorum
Carex sp.
KOleria cristata
Oryzopsis hymenoides
Sitanion hystrix
Sporobolus cryptandrus
Stipa comata

22
2

T
T

45

4

2

T

9
8

T

8

T

11
94

12

T

T

Sub-total

16

Forbs
Agoseris sp.
Allium sp ,
Antennaria sp.
Chenopodium sp.
Chrysopsis villosa
Erigeron sp.
Eriogonum alatum
Er~ogonum umbellatum
Erysimum asperum
Lappula sp.
Lupinus sp.
Lygodesmia sp.
Oenthera trichocalyx
Phlox sp.
sph&amp;eralcea coccinea
Zygadenus elegans
Unidentified forbs

2
2
1
42

T
T

T

2

4
1

T
T

10

1

6
1

T

T
1
1

29
9
1

T
T

15
2

5

T
T

1

T

2

Sub-total

7

Shrubs
Artemisia cana
Artemisia IiIifolia
Artemisia frigida
Artemisia tridentata
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Gutierrezia sarothrae
Opuntia sp.
Purshia tridentata
Tetradymia canescens

53
5
13
3
15

14
T

1
T

4
8

48

8
64

T

10
39

59
2

T

Sub-total

76

Total

99

�-56-

Table 11. Analysis of six antelope rumen samples collected on the bitterbrush study area in Moffat County during the period October-March.

Plant Species

Percent
Frequency

Percent
Composition

67
17
67

1
T
T

Grasses
Bromus tectorum
Oryzopsishymerioides
Stipa corriata
Unid. grass fragments

T

Sub-total

1

Forbs
Chenopodium sp.
Cryptantha janiesii
Erigeron sp.
Eriogonum sp.
Leptodactylon pungens
Mentzelia sp.
Unid. forb fragments

17
17
33.
17
33
17

T
T

1
T

1
T

1

Sub-total

3

Shrubs
Artemisia cana
Artemisia dracunculus
Artemisia frigida
Artemisia tridentata
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Opuntia sp.
Purshia tridentata
Unid. shrub fragments

83
17
33
67
17
50
17
100

19
T

1
26
T
T
T

35
15

Sub-total

96

Miscellaneous
Antelope
Rocks

hair

17
17

T
T

Sub-total

Total

T

100

�-57-

Table 12. Summary of the vegetation surveys on the bitterbrush
area in Moffat County during the period October-March.

Plant Species

Percent
Frequency

Percent
Composition

2
2
70
2
2
12
1
3
63

T
T
5
T
T
1
T
1
6
T

Grasses
Agropyron sp.
Agropyron trachycaulum
Bromus tecto:tum
Carex sp.
Koleria cristata
Oryzopsis hymenoides
Sitanion hystrix
Sporobolus cryptandrus
Stipa comata
Unidentified grasses

Sub-total

13

Forbs
Chenopodium sp.
Chrysopsis villosa
Eriogonum sp.
Lappula redowski
Lupinus sp.
Oenothera trichocalyx
Phlox sp.
Senecio sp.
Tragopogon porrifolius
Zygadenus elegans
Unidentified forbs

3

T
T

1

12
1

1

T
T
T
T
T
T
T
T

1

5
2
1
1
1

Sub-total

1

Shrubs
Artemisia cana
Artemisia frigida
Artemisia tridentata
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Gutierrezia sarothrae
Opuntia sp ,
Purshia tridentata

study

30
16
4
11
35
7
39
65

7
1

4
4
9

T
5
56

Sub-total

86

Total

100

�-58-

Table 13. Analysis of six antelope rumen samples collected on the rabbitbrush study area in Saguache County during the period April-June.

Plant Species

Percent
Frequency

Percent
Composition

33
17
17
17
17

T
T
T

Grasses
Boute1oua gracilis
Muh1enbergia montana
Poa fend1eriana
Sitanion hystrix
Stipa comata
Unid. grass fragments

T
T

1

Sub-total

1

Forbs
Arenaria fend1eri
Chenopodium sp.
Chrysops is vi110sa
Cryptantha jamesii
Eriogonum sp.
Eriogonum ~
Gaura coccinea
~sp.
Lappu1a sp.
Leucocrinum montanum
Lygodesmia sp.
LapiDus

Sp.

Oenothera trichoca1yx
Oxytropis sp.
Potentilla sp ,
Pseudocymopsis
Sa1so1a kali
Senecio sp.
Sisymbrium sp .
Sphaera1cea coccinea
Unid. forb fragments

T
T

50

33
17
17

T
T

5
4
1
T
T
1
T

50

33
17
17
17
17
17
17
17
17
17
17
33
17
17
33

T
T

T
T

1
T

1
T

1

17

Sub-total

Artemisia dracuncu1us
Artemisia frigida
Artemisia tridentata
Chrysothamnus nauseosus
Chrysothamnus parryi
Chrysothamnus viscidif10rus
Opuntia sp.
Quercus gambe11i
Rhus tri10bata
Ribes cereum
------Sarcobatus vermicu1atus
Symphoricarpos sp.
Unidentified vine
Unid. shrub fragments

31

2

17
67
17
33
17

37
T

1
3
2
7
T
1
3
4

50
50

17
17
17
17
17
17

T

1
6

Sub-total

67

Miscellaneous
Insects
Lichen
Antelope hair

T
T

33
33
17

T

Sub-total
Total

T
99

�-59-

Table 14. Summary of the vegetation surveys on the rabbitbrush study area in
Saguache County during the period April-June.

Plant Species

Percent
Frequency

Percent
Composition

2

T
T

Grasses
Agropyron smithii
Agropyron trachycaulum
Aristid..'!
sp.
Boute1oua gracilis
Muh1enbergia richardsonis
Muhlenbergia torreyi
Oryzopsis hymenoides
Sit anion hystrix
Sporobo1us crytandrus
Stipa comata
Stipa lettermani

1
1

T

82
2
22

49

1

T

9

1
2
2
1

T

6

14
14
3

Sub-total

61

Forbs
Allium sp.
Chenopodium sp.
Chrysopsis vi110sa
Cryptantha jamesii
Erigeron sp.
Eriogonum sp.
Gaura coccinea
Hymenoxys richardsoni
Lappula sp.
Lupinus sp.
Oenothera trichoca1yx
Phlox sp.
Potentilla sp.
Sa1so1a ka1i
Sphaera1~coccinea
Unidentified forbs

6

T
T

4

T

2
2

1

T
T

7
2

T
T

1
15
1
1
5
1

T

1
T
T
T
T

1
1

17
7

T

Sub-total

3

Shrubs
Artemisia dracuncu1us
Artemisia frigida
Atriplex canescens
Atrip1ex sp.
Chrysothamnus nauseosus
Chrysothamnus viscidif10rus
Echinocactus sp.
Eurotia 1anata
Gutierrezia sarothrae
Leptodactylon sp.
Opuntia sp.
Quercus gambe11i
Ribes cereum
------Symphoricarpos sp.
Yucca sp.

1

T
2

32
5
1
16
31

T
T

2

T

10

3
T

7
8

1
1
13
1
1

T

6
1
1
T
T

1

1

Sub-total

28

Mis ce11aneous
Lichen

8

63

Total

100

�-60-

Table 15. Analysis of six antelope rumen samples collected on the rabbitbrush study area in Saguache County during the period July-September.

Plant Species

Percent
Frequency

Percent
Composition

87
17
17
17
17

T
T
T
T
T
1

Grasses
Boute1oua gracilis
Carex sp.
MUhIenbergia montana
Muh1enbergiatorreyi
Oryzopsis hymenoides
Unid. grass fragments

Sub-total

1

Forbs
Allium
~ia
fend1eri
Aster co1oradoensis
CfieriOpodiumsp ,
Cryptantha jamesii
Erigeron sp.
Eriogonum sp.
Erysimum aspermum
Hymenoxys richardsoni
Lappu1a sp.
Lupinus sp.
Lygodesmia sp ,
Mentzalia sp.
Oenothera trichoca1yx
Penstemon sp.
Phlox sp.
P"Otentil1a s p ,
Polygonum sp ,
Sa1so1a ka1i
Sedum sp-.-SiSYIDbrium sp ,
Sphaera1cea coccinea
Unid. Compositae
Unid. forb fragments

17
33
17
87
33
87
87
17
17
17
17
33
17
17
17
17
17
17
17
17
17
17
17

3
T
T
2

T
1
6
5
T
T
T
T
T
T
T
T
1
1
T
1
T
9
15
3
Sub-total

44

Shrubs
.Art emf.sLa dracuncu1us

Artemisia frigida
Artemisia tridentata
Atrip1ex confertifo1ia
Cercocarpus sp.
Chrysothamnus nauseosus
Chrysothamnus parryi
Chrysothamnus viscidif10rus
Opuntia sp.
Rhus tri10bata
Ribes cereum
Rosa sp-.-$Yiiij?horicarposs p ,
Unid. shrub fragments

33
50
17
33
17
33
50
50
33
33
17
17
50

10
3
T
2

T
2

1
T
4

9
T
T
18
4

Sub-total

53

Miscellaneous
Insects
Lichen
Rocks
Antelope hair

T
T
1
T

33
17
17
33
Sub-total

1

Total

99

�-61Table 16. Summary of the vegetation surveys on the rabbitbrush study area
in Saguache County during the period July-September.

Plant Species

Percent
Frequency

Percent
Composition

9
91
16

1
T
52
1

1
2

T
T

Grasses
Agropyron smithii
Aristida sp.
Bouteloua gracilis
Carex sp.
~ria
cristata
Muh1enbergia richardsonis
Muhlenbergia torreyi
Poa fend1eriana
Sitanion hystrix
Sporobo1us crytandrus
Stipa comata
Stipa 1ettermani
Unidentified grasses

2

19

3

2

T

8
5

T
T
3
T
T

14
1

Sub-total

60

Forbs
Allium sp .
Arenaria fend1eri
Castilleja sp .
Chenopodium sp.
Crvptantha jamesii
Erigeron sp.
Eriogonum sp.
Hymeno~s richardsoni
Lappu1a sp ,
Lupinus sp.
Lygodesmia sp.
Oenothera trichoca1yx
Penstemon sp.
Phlox sp .
Potentil1a sp ,
Sa1so1a ka1i
Senecio sp ,
Sphaera1cea coccinea
Unidentified forbs

1
6
1
14
1

T
T
T
1
T
T
1
T

9

12
5

6
6
1
1
1
11
1
12
4
21

T

2
T
T
T
1
T
2
T
1
1
Sub-total

9

Shrubs
Artemisia dracunculus
Artemsia frigida
Chrysothamnus nauseosus
Chrysothamnus ~
Chrysothamnus vis cidifl0 rus
Echinocactus sp.
Eurotia 1anata
Gutierrezia sarothrae
Opuntia sp.
Quercus gambe11i
Rhus t r i.Lob at.a
Ribes cereum
------SymEhoricaq)os sp.
Yucca sp.

T
3
3
T
4
T
1
T
9

4
26
11
1
16
1
10
4
34
1
1
1
1
2

T

1
T
1
T
Sub-total

22

Mis ce11aneous
Lichen

8

55

Total

99

�-62-

Table 17. Analysis of thirteen antelope rumen samples collected on the
rabbitbrush study area in Saguache County during the period October-March.

Plant Species

Percent
Frequency

Percent
Composition

8

T
T
T
T
T
T
T
T
1

Grasses
Agropyron smithii
Bouteloua gracilis
Festuca sp.
Muhlenbergia sp.
Muhlenbergia richardsoni
Sitanion hystrix
Sporobolus cryptandrus
Stipa comata
Unid. grass fragments

77

8
8
8
23
8
15

Sub-total

1

Forbs
Antennaria parva
Arenaria fendleri
Astra!\a1us sp.
Chenopodium sp.
Eris;eron sp.
Erios;onum sp.
Erysimum aspermum
Lupinus sp.
Oenothera trichocalyx
Oxytropis sp.
Salsola kali
Sedum sp-.-Sphaeralcea coccinea
Unid. Compositae
Unid. Cruciferae
Unid. forb fragments

T
T
T
T
T
1
T
T
T
T
1
T
T
T
T
1

8
15
15
23
8
46
8
8
8
8
15
15
54
8
8

Sub-total

3

Shrubs
Artemisia dracunculus
Artemisia frigida
Artemisia tridentata
Atriplex sp .
Chrysothamnus nauseosus
Chrysothamnus parryi
Chrysothamnus viscidiflorus
Juniperus scopulorum
Opuntia sp.
Pinus ponderosa
Rhus trilobata
Sarcobatus vermiculatus
Symphoricarpos sp.
Unid. vine
Unid. shrub fragments

5
23
T
9
15
2
T
T
35

69
77

8
54
46
31
38
8
100
8
15
8
8
8

T

T
T
T
T
5
Sub-total

94

Miscellaneous
Lichen
Insects
Rocks
Antelope hair

T
T

23
8
15
23

1
T

Sub-total

1

Total

99

�-63-

Table 18. Summary of vegetation surveys on the rabbitbrush study area in
Saguache County during the period October-March.

Plant Species

Percent
Frequency

Percent
Composition

11
1
90
2
4
1
3
1
15
5

1
T
43
T
1
T
1
T
1
1
2

Grasses
Agropyron smithii
Aristida sp.
Bouteloua gracilis
Buchloe dactyloides
Carex sp.
Muhlenbergia richardsoni
Muhlenbergia torreyi
Oryzopsis hymenoides
Sitanion hystrix
Sporobolus cryptandrus
Stipa~
Unidentified grasses

7

T

Sub-total

50

Forbs
Came1ina microcarpa
Castilleja sp ,
Chenopodium sp.
Chrysops is villosa
Erigeron sp.
Eriogonum sp.
Helianthus sp.
Hymenoxys richardsoni
Lappula sp.
Leptodactylon sp.
Lupinus sp.
Lygodesmia sp.
Oenthera trichocalyx
Phlox sp.
POt"elltillasp ,
Salsola kali
Sphaeralcea coccinea
Thermopsis sp.
Unidentified forbs

1
1

T
T

6

T

1
1
12
1
2
9
1
1
1
1
2
1
14
9
1

T
T
2
T
T

1
T
T
T
T
T
T
5
1
T
T
Sub-total

9

Shrubs
Artemisia dracunculus
Artemisia frigida
Atriplex canescens
Atriplex sp.
Chrysothamnus nauseosus
Chrysothanmus parryi
Chrysothamnus viscidiflorus
Echinocactus sp.
Eurotia lanata
Gutierre~rothrae
Opuntia sp.
Quercus sp.
Rhus trilobata
Ribes cereum
SyTilj?ho
os sp •
Tetradymia canescens
Yucca sp.

ric;u:p

3
38
3

1
4
1
1
4
T
9
T
2
T
11
1
T
T
T
T
T

2

l3
1
34
1
l3
3
26

1
1
1
1
3
1
Sub-total

34

Miscellaneous
Lichens

6

Sub-total
Total

6

99

-----

�-64-

Feeding

Activity

Observations from the three areas were combined and classified by the
three seasons of the year used above. A total of 7,958 observations
were recorded.
There were fluctuations in the daily and seasonal
feeding patterns of antelope; however, activity patterns during the
spring and summer seasons were very similar.
Most antelope observed
from daybreak to 10 A.M. were active (71-78 percent), with activity
decreasing until noon (Fig. 1). Activity increased again from noon to
2 P.M., then decreased again until 4:00 P.M. Activity increased again
until dark.
The antelope generally were more active during the fallwinter period than during the other two seasons.
In the fall-winter
season nearly all antelope observed from daybreak to 8:00 A.M. were
actively feeding (98 percent).
During this season feeding activity
decreased earlier in the day and remained more constant during mid-day
than it did during spring or summer.
Antelope activity in the fallwinter season increased from 2:00 P.M. until dark. During the later
part of this time period, 4:01 P.M. - dark, 99 percent of the animals
observed were feeding.
Antelope are active at night, however, the only successful night-time
observations were made near a waterho1e during the summer when there
was bright moonlight.
The antelope activity at the waterho1e peakedout at approximately 10:00 P.M. and again at 1:00 A.M.

DISCUSSION

Diets
The basic antelope diet on the three range types in this study is comparable to findings in eastern Colorado and states in the northern
extension of the antelope distribution.
Shrubs were the major food items
during the fall-winter periods; forbs were then secondary in importance,
increasing during the spring and summer periods.
Grasses were a minor
food item throughout the year. There appeared to be a certain degree of
selectivity in the antelope feeding behavior, particularly during the
summer periods.
Vegetation surveys indicated forbs were the least abundant
class of 'forage on the three areas, yet were major food items. Grasses
were abundant on the areas, but were of minor importance.
There were differences in the relative amounts of forbs and shrubs in the
rumen samples from the three areas. Forbs were more prominent in the
spring and summer diets on sagebrush ranges than on bitterbrush or rabbitbrush range types (Fig. 2). Browse was most prominent in spring and summer
diets on bitterbrush range. The rabbitbrush range was intermediate.
Palatability and availability of vegetation undoubtedly plays an important
role in antelope feeding behavior.
As the forbs become green and succulent
during the spring and summer months antelope seek them out, even though
forbs are less abundant than other classes of vegetation on the range.

�-65-

Grasses were never very abundant in the rumen samples; however, grass
consumption increased during the spring of the year when many of these
plants were actively growing.
Antelope use on grasses and forbs decreased as the plants matured and cured-out.
Snow accumulation during
winter months often made grasses and forbs unavailable to antelope,
thus they were forced to feed on the shrubs available above the snow.
However, shrubs were preferred foods in the late fall and early winter
peTiods, before snow accumulated on the areas.
Shrubs were not dormant
at this time and were likely more palatable than the cured forbs and
grasses.
Chemical content of the plants probably affected antelope
selectivity also, but this was beyond the scope of the study.
Antelope ate a wide variety of plants which were available.
Generally
plants occurred in the rumen samples in the same relative abundance
vegetation surveys indicated they occurred on the range. However, a few
species were preferred and sought by antelope.
Plants comprising a major
portion of the rumen samples in the sagebrush type were: Artemisia
.tridEmtata, Purshia tridentata, Artemisia cana, Gilia laxiflorus,
Sphaeralcea coccinea, Astragalus sp., and ~a
sp. Artemisia tridentata
was the most common· plant on the area and was a major plant in the diet.
Gilia laxiflorus and Gaura sp. were not common on the area, but were
important food items. The more important plants in the diets of antelope
on the bitterbrush type were: Purshia tridentata, Artemisia cana,
Chrysothamnus sp., Erigeron sp., Eriogonum sp., Sphaeralcea coccinea,
Bromus tectorum, and Stipa comata. Purshia tridentata was the most common
plant on the area and most common plant in the diet. Bromus tectorum and
Stipa comata were the most common grasses on the area and in the rumen
samples.
The more important plants in the antelope diets on the rabbitbrush area were: Artemisia frigida, Artemisia dracunculus, Chrysothamnus
sp., Opuntia sp., Sphaeralcea coccinea, Erigeron sp., Eriogonum sp., and
Bouteloua gracilis.
Bouteloua gracilis was the most common plant on the
range, but eaten in minor amounts.
In general the plant species eaten by
the antelope was different for each of the three range types.
There are a few interesting observations concerning plant species eaten by
antelope.
Artemisia sp. is a common plant species in the diets of antelope
in many areas. Also, Sphaeralcea sp. is a common plant listed for many areas.
It.was interesting to note Opuntia sp. was common in all three areas studied
here , but was only an important food item on the rabbitbrush area, which
was more of a grassland range than the other two areas. There is some
'similarity in that Opuntia sp. were important food items in the grasslands of eastern Colorado and Kansas (Hoover et. al 1959; Hlavachick 1966).
There is inadequate information on this relationship, but cacti on these
grassland areas may have been a substitute for woody plants.
Cacti may also
have been eaten for succulence.
In evaluating the food habits data for
bitterbrush and sagebrush ranges in Moffat County, shrubs greatly decreased
in the diet during the summer and spring on the sagebrush area more than on
the bitterbrush area. This difference may be due to the species involved.
Artemisia tridentata was the more important browse species on the sagebrush
area and may be less acceptable when green forbs are present than the
Purshia tridentata and Artemisia cana on the bitterbrush area. Deming (1963)
in discussing antelope food habits stated, "It may well be that the high
incidence of sagebrush in the diet of antelope today may be through availability rather than palatability."
These discrepancies and similarities are
interesting, but much information is lacking concerning the causes.

�-66-

100

15
bO

s::

.~ 50
Po
U)

25

100~

15

~I

100-

.

I

r---"'"""""

r---

,

r---

15
,.----

~
'.

!

'.

"""

1······

25

l-

I

I

!

o
AM-8

Fig. 1.

8-10

10-12

12-2

2-4

4-PM

Time of Day

Daily and seasonal activity patterns

for antelope.

�-67Range-Types
Sagebrush
Bitterbrush
Rabbitbrush

100

80

60
:z;
0

I/)

H.o
~
~ 0
(f)~

40

0

~

-:"'""'j

0

u

~

ILl

u

,

0::
ILl
p..

.

9j':!
":c:iJ

,

100

rl
1
1

80

t

!

~

I

,

"

I/)

.0

60

:,

.. '1

.•..

~
~
.c

,
'.

f

!

~.: j

II

.'

(f)

40

i

;

"

"

. "

!

.

!

I ;: I

·l·:'.:
-:

t '.
j'
j'

~.'.-J

"'1

{"

j

Spring

'0'

"

"
•

4

-

~ ..

~.

Summer

Fall-Winter

SEASON OF YEAR
Fig. 2. Relative abundance of shrubs, forbs, and grasses in the antelope
rumen samples collected on three range types.

�-68-

Activity
Activity patterns established in this study were very similar to findings
in other studies.
Antelope were most active in early morning and late
evening hours yearlong.
Antelope were more active during mid-day in the
winter than during spring or summer.
Beuchner (1950) reported the following average daily percentages of feeding time: autumn 72.8, winter 77.8,
spring 63.0, and summer 67.0. The average daily percentages of feeding
time on this study were: fall-winter 79, spring 68, and summer 65. Rest
periods during the day were followed by short periods of activity in other
studies (Beuchner 1950; Einerson 1948). This may account for the increased
activity shown from noon to 2:00 P.M. during spring and summer in this
study.

SUMMARY
1.

Study areas were established in three range types in Moffat and Saguache
counties; (a) sagebrush, (b) bitterbrush, and (c) rabbitbrush.

2.

A total of 76 antelope rumen samples were obtained
at monthly intervals for one year.

3.

Vegetation surveys were conducted in the area when each antelope was
collected to determine plant species available to the antelope.

4.

Antelope
feeding,

5.

Antelope diets on this area were comparable
other states.

6.

Browse was a major forage class in antelope diets and forbs were
secondary.
Forbs increased in the diet during spring and summer
periods.
Grasses were minor food items throughout the year.

7.

Plant species were generally found in rumen samples in the same
relative proportions as they occurred on the range; although the
antelope were selective for a few species, and forbs in general.

8.

There were some differences
areas studied.

9.

Palatability and availability of the plants likely influenced
foraging habits of the antelope.

10.

Important plant species in the rumen samples from the sagebrush range
were: Artemisia tridentata, Purshia tridentata, Artemisia cana, Gilia
laxiflorus, Sphaeralcea coccinea, Astragalus sp., and Gaura sp.

11.

Important plant species in the rumen samples from the bitterbrush
range were: Purshia tridentata, Artemisia cana, Chrysothamnus sp.,
Erigeron sp., Eriogonum sp., Sphaeralcea coccinea, Bromus tectorum,
and Stipa comata.

from these areas

activities were recorded in two categories,
and (b) lying down or inactive.

(a) active or

to those reported

in the food habits between

in

the three

the

�\

-69-

12.

Important plant species in the rumen samples from the rabbitbrush
range were: Artemisia frigida, Artemisia dracunculus, Chrysothamnus
sp., Opurttia sp., Sphaeralcea coccinea, Erigeron sp., Eriogonum sp.,
and Bouteloua gracilis.

13.

Antelope

14.

Antelope were more active during mid-day through the fall-winter
than during either the spring or summer periods.

were most active during early morning

LITERATURE

and late evening hours.
period

CITED

Bayless, S. 1968. Food habits, range use and home range of pronghorn
antelope in central Montana during winter.
Proc. Antelope States
Workshop.
3:104-114.
Beale, D. M., and G. W. Scotter.
1968. Seasonal forage use by pronghorn
antelope in western Utah. Utah Science, p. 3-6, 16.
Beuchner, H. K. 1950. Life history, ecology, and range use of the pronghorn antelope in Trans-Pecos Texas. The Amer. Midland Nat.
43(2):
257-354.
Cole, G. F., and B. T. Wilkins.
1958. The pronghorn antelope.
Its range
use and food habits in central Montana with special reference to wheat.
Mont. Fish and Game Dept., Tech. Bull. No.2.
39 pp.
Deming, O. V. 1963. Antelope
Conf. 1963. p. 55-60.·

and sagebrush.

Trans. Interstate

Antelope

Einerson, A. S. 1948. The pronghorn antelope and its management.
pole Co., Harrisburg, Pa. 238 pp.

Stack-

Hlavachick, B. D. 1966. Some preferred
Antelope States Workshop.
2:60-65.

Proc.

foods of Kansas antelope.

Hoover, R. L., C. E. Till, and S. Ogilvie.
1958. The antelope of
Colorado.
Colo. Game and Fish Dept., Tech. Bull. No.4.
110 pp.
Kautz, L. G. 1942. Report of the antelope
Dept., Fed. Aid Report W-4-R.

survey.

Mason, E. 1952. Food habits and measurements
J. Wildl. Mgmt.
16(3):387-389.
Prenzlow, E. J., D. L. Gilbert,
patterns of the pronghorn.
Report No.7.
15 pp.

Colo. Game and Fish

of Hart Mountain

antelope.

and F. A. Glover.
1968. Some behavior
Colo. Dept. Game, Fish and Parks.
Spec.

�-70-

Russell, T. P. 1964. Antelope of New Mexico.
Game and Fish, Santa Fe, N.M. 103 pp.

New Mexico Dept. of

Severson, K., M. May, and W. Hepworth.
1968. Food preferences,
carrying capacities, and forage competition between antelope and
domestic sheep in Wyoming's Red Desert.
Univ. Wyo. Agri. Exper.
Sta., Laramie,. Wyo. Sci. Mono. No. 10. 51 pp •

Prepared

by

.~~--tf.'~
Ceo rge . Bear
Wildlife Researcher

�-71January,

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-4l-R-22

Bighorn

Sheep &amp; Mountain

Work Plan No.

1

Job No.

12

Job Title

Location

Period Covered:
'Personnel:

and Distribution

Goat Investigations

of Bighorn Sheep Herds in Colorado

June 1, 1971 to May 31, 1972

George D. Bear

ABSTRACT

A final report- has been comp Le't ed and is in the last editing stage. The report
wili be published in the. Game Research Report format but under separate cover
in a spe~ial binder to~provide a readily usable source of information for
wildlife managers.
.

1973

��-73-

LOCATION AND DISTRIBUTION
OF BIGHORN SHEEP HERDS IN COLORADO
George D. Bear

P. S. OBJECTIVE
To determine numbers, composition, seasonal
of bighorn sheep herds in Colorado.

range and distribution

SEGMENT OBJECTIVE
To publish

final report manuscript.

METHODS

AND MATERIALS

This is strictly a publications job which during the past segment has
been concerned with editing and final printing.
It is expected the
publication will be completed and ready for distribution by April 1,
1973.

RESULTS AND DISCUSSION
This report, which will appear under separate cover, is a compilation
of all available information for each bighorn sheep herd in Colorado.
It is hoped wildlife managers will not only find this publication useful but also will add to it as additional information becomes available
for a particular sheep herd.

Prepared

by

~.v"

_~c"'-"L.

'f ..C:.

/.. .

~.•••.•.
/.-

~.;.:.j~
. _"
,
_

Geojfge D. Bear
Wildlife Researcher

1_"

i

..

��-75January,

JOB PROGRESS

State

of

Project
Work

REPORT

COLORADO
No.

W-4l-R-22

Bighorn

Plan No.

1

Job No.

Job Title
Period

Evaluation

Covered:

Personnel:

June,

Robert

of the Nutritional
1971 to May,

L. Schmidt,

Sheep

&amp; Mountain

Goat

Investigations

16
Requirements

of Bighorn

Sheep

1972

Robert

E. Keiss

and Gene G. Schoonveld

ABSTRACT
First year collections of bighorn sheep fecal samples and forage plants
were obtained from the Trickle Mountain, Buffalo Peaks and Pikes Peak
ranges.
Collections were made monthly June-September
and thereafter
during November, January and March.
Bighorn sheep food-habits were
determined using the fecal microanalysis
technique.
Grass and grasslike forages comprised the major proportion of yearly sheep diets, however
increased utilization of browse species occurred during early fall.
Use
of forbs was insignificant by sheep on all three ranges - totals of all
species used being less than 4 percent of their diets.
Two l2-day feeding trials were conducted to measure accuracy of fecal
microanalysis
for identifying plant species and estimating percent realtive
density of forages ingested by an animal.
A fistulated domestic goat
(Capra sp.) was force fed through the fistula two formulated diets compounded
with varying percentages
of seven plant species.
Fecal samples collected
after the animal had been fed the test diets for 10 and 12 days were analyzed
by the microanalysis
technique for identifiable plant residues.
For unknown
reasons the majority of residues were unidentifiable
woody-stem material
and no determination
of species composition in the diets could be made.
Unfed samples of the test diets were also analyzed by the microanalysis
technique.
All plant species in both diets were identified.
However, two
plant species not in the diets were also estimated to be present in minor
amounts.
Ratios of percentages of known composition
to estimated compossition varied from 0.1 to 4.2.
Percentages of Artemisia frigida
were highly
overestimated while Purshia tridentata and Cercocarpus montanus underestimated
in both diets.

1973

��-77-

EVALUATION

OF THE NUTRITIONAL

REQUIREMENTS

OF BIGHORN

SHEEP

Gene G. Schoonveld

P. S. OBJECTIVE
Develop methods: (a) for evaluation of the nutritional requirements
bighorn sheep under natural conditions, and (b) to measure adequacy
selected ranges to meet these requirements.

of
of

SEGMENT OBJECTIVES
1.

To estimate the nutritional
bighorn sheep.

quality of forage species utilized

by

2.

To evaluate if the nutritional level of selected bighorn sheep ranges
is adequate to meet nutrient requirements of bighorn sheep during all
seasons of the year.

(AMENDMENT )
To measure accuracy of fecal sample microanalysis
for identifying plant
species and estimating percent relative density of forages ingested by
a ruminant.

DESCRIPTION

OF STUDY AREAS

Three bighorn sheep ranges were selected for nutritional investigations.
Study areas chosen were considered indicative of differing quality sheep
ranges based upon trend of the population and estimated numbers of sheep
currently using the range.
Study Area No.1,
Trickle Hountain, Saguache County, Colorado, is a low
altitude (8,400-12,100 ft.) primarily pinyon-juniper
type bighorn sheep
range.
Quality of this range was arbitrarily considered good in that the
bighorn sheep population has been generally increasing.
Area No.2,
the
Buffalo Peaks, Park County, Colorado, is primarily an alpine (11,600-12,900
ft.) sheep range.
Quality of this range was considered marginal as the
bighorn population has been declining.
Area No.3,
Pikes Peak, El Paso
and Teller counties, Colorado, is both alpine and sub-alpine (10,20014,100 ft.) bighorn sheep range.
Since a major sheep die-off during
1953-54, the bighorn population has been steadily increasing which is
thought to be directly related to mineral supplementation.

�-78-

METHODS

Nutritional

AND MATERIALS

Quality of Forage Species Used by Bighorn Sheep

Bighorn sheep fecal samples (for food-habits analysis) and forage samples
were collected from the three study areas beginning June 1971. Collections
were made monthly, June-September and thereafter during November, January
and March.

Food-habits

Analyses

Estimates of forage species used by bighorn sheep were made using microanalysis techniques to identify residues of plant tissues in fecal samples.
Samples were collected after first observing sheep and subsequently collecting
only fresh material from the area where sheep had been observed.
This procedure insured fecal samples collected were definitely from bighorn sheep
and also representative of the season.
A minimum of 10 pellet groups were
collected during each period.
Individual groups were placed in envelopes
and labeled as to study area and date. Efforts were made to collect the
entire pellet group, however, this was not always done due to snow conditions
or elimination while the animal had been moving.
Preparatory to analysis samples were placed in a 4mm soil sieve and agitated
to remove foreign matter (gravel, plant parts, etc.).
If dust or soil
particles remained,the sample was rinsed using minimal quantities of water
and air-dried 10-14 days. Sub-samples of 6-8 pellets from each group were
composited and ground in a Wiley mill using a 2 mm screen.
Determinations
of bighorn sheep foob-habits from these samples were subsequently made by
the Composition Analysis Laboratory, Department of Range Science, Colorado
State University, using microanalysis techniques.
Procedures used for
preparing microscope slides, identifying tissue residues and estimating
percent relative density of forage species comprising sheep diets were
those used by Sparks and Malechek (1968). Twenty systematically located
fields were observed on each of 10 slides using 100 X magnification.
Residues of plant species were identified and percent relative density of
forage estimated.

Forage Quality Analyses
Representative forage samples were also collected from areas in which sheep
were observed to have been feeding.
Initially, all forage plants in an
area were collected, however,' as food-habits became known only those forages
used by sheep were collected.
Grasses and grasslike species were collected
by clipping the entire plant as close to the ground as possible; browse by
clipping current years growth.
Clipped material was placed in paper bags
and labeled as to tentative identification, date, and study area.
Prior to chemical analysis forage species collected were definitely identified using Bighorn Sheep Herbarium Reference Manuals (Shepherd 1972) or
Manual of the Plants of Colorado (Harrington 1954).
Samples collected were
air dried 2-3 weeks, ground in a Wiley mill using 1 mm screen, and stored
in 4 oz sample jars until chemically analyzed.

�-79-

Forage species were analyzed for nutritional quality using standard laboratory procedures for determination of percent dry matter, crude protein,
crude fat and ash (AOAC 1965), cell-wall constituents (Van Soest and Wine
1967), acid-detergent fiber (ADF) and lignin (Van Soest 1962). Trace
minerals (calcium, potassium, sodium, magnesium, manganese, copper, zinc
and iron) were determined using a Perkin-Elmer 303 atomic absorption spectrophotometer.
Phosphorus content was estimated using the vanadate-molybdate
procedure and analyzing content with a Spection 20 spectrophotometer.

Forage Digestibility
Digestion coefficients and total digestible nutrients (TDN) of bighorn
sheep diets were estimated by simulating diets (based on percent relative
density of forages in actual diets determined by fecal microanalysis) and
chemically analyzing diets and fecal material for proximate constituents.
Chemical analyses used are those previously described.
Differences in
percent lignin between simulated diets and fecal samples was used as an
indirect indicator of digestion and used to estimate diet digestibility
and TDN.

Evaluate Nutritional Level of Ranges in
Furnishing Nutrient Requirements of Bighorn Sheep
Nutrient requirements of bighorn sheep are not known.
Nutritional data,
obtained through Procedure One for the three study areas, will therefore
be compared for indications of possible nutrient deficiencies.
Such
comparisons will not define nutrient requirements of bighorn sheep but
will permit nutritional evaluation of different ranges supporting sheep.

Accuracy of Fecal Microanalysis
Technique in Determining Food-Habits

Feeding Trials
Two consecutive l2-day feeding trials using a fistulated domestic goat
were conducted.
The animal was put into an isolation pen (6.1 X 1.8 m,
with cement floor) 2 weeks prior to the first trial to accustom it to
confinement and measure its normal daily food intake.
Daily intake of its
regular feed was used as a basis for the amount of test diet fed during the
trials.
Ad libitum intake was measured by offering a weighed amount of
feed and weighing back uneaten amounts each following day. Prior to feeding
a test diet, the animal was fasted 48 hours to aid in voiding its digestive
tract of previously ingested food. During the l2-day trials, test diets
were force-fed twice daily by introducing 250g of the diet through the
fistula directly into the rumen.
No other food was available to the
animal.
After each feeding the most recent fecal group was collected,
placed in a paper sack and labeled as to date and time of collection.
The pen was then swept clean of all remaining fecal pellets.

�-80-

Test Diets
Test diets fed were formulated with true mountain mahogany (Cercocarpus
montanus), bitterbrush
(Purshia tridentata), big sagebrush (Artemisia
tridentata), fringed sagebrush (Artemisia frigida), rubber rabbitbrush
(Chrysothamnus nauseosus), snow willow (Salix nivalis) and alfalfa
(Medicago sativa).
Plant species, except Salix and Medicago, were
collected during late winter by hand-clipping the previous year's growth
from plants along the front range near Fort Collins.
Salix was collected
from the Pikes Peak range during June when the species was in full leaf
with mature catkins present.
Alfalfa used was an exceptionally leafy,
baled, third-cutting from the previous season.
Each plant species was
ground individually in a hammer mill with a 3 mm screen resulting in
particles varying from extremely small to 3 em in length.
After grinding,
material was stored in plastic sacks at -10 C.
Two test diets were formulated with varying percentages of individual plant
species (Table 1). Species used were sampled to determine moisture content, compounded, thoroughly mixed and stored in plastic bags at 0 C until
fed.

Table 1. Composition (percent dry weight
fed during feeding trials.

basis) of two formulated

ComEosition

(Percent)
Diet II

Plant Species

Diet I

Artemisia

tridentata

9.3

5.0

Artemisia

frigida

14.1

10.0

9.7

25.0

0

9.9

Cercocarpus

montanus

Chrysothamnus

nauseosus

Medicago

sativa

35.5

10.7

Purshia

tridentata

9.3

24.9

22.1

14.5

Salix nivalis

Microanalysis

diets

of Fecal SamEles

Fecal samples were analyzed by the Composition Analysis Laboratory, Department of Range Science, Colorado State University, using micro-techniques
identical to those outlined for Procedure One.

�-81-

The laboratory technician who prepared and read the slides had extensive
training and experience in micro-techniques.
She did not have prior
knowledge of the diet-compositions but was furnished a complete list of
all browse species that could be expected but were not necessarily included in the diets.

RESULTS AND DISCUSSION
Nutritional

Quality of Forage Species Used by Bighorn

Sheep

Food-habits
Primary forage species identified in bighorn sheep fecal samples collected
from the three study areas are listed in Table 2. Comparisons of data
among the three areas indicate sheep using the Trickle Mountain range are
utilizing greater numbers of different plant species than sheep on either
Pikes Peak or the Buffalo Peaks.
Grasses and grass-like forage comprise
the major proportion of bighorn sheep diets (Fig. 1), however, there was
increased utilization of browsel species during early fall (Fig. 2). Use
of forbs2 by sheep on all three ranges was insignificant - totals of all
species used being less than 4 percent of the diets.
Comparisons of food-habits data between summers 1971 and 1972 show extreme
variation in all study areas. Variability of data may be due to actual
differences in food-habits of sheep between years due to different climatic
conditions or errors in identifying and estimating percent relative density of plant species in sheep diets.
Extremely high utilization of
browse species, primarily Artemisia frigida by bighorn sheep on Trickle
Mountain during November 1971 and January 1972 are thought to be overestimates (See Amendment results).

Forage Quality
Chemical analysis of forage samples collected during this segment have not
been completed or data obtained to date analyzed.
Results will therefore
not be presented until all aspects have been completed.

Forage Digestibility
Chemical analysis of forage and fecal samples needed to estimate digestion
coefficients and compute TDN are continuing.
Results will be reported in
future segments.
Evaluate Nutritional Level of Ranges in
Furnishing Nutrient Requirements of Bighorn Sheep
Range evaluations are dependent upon comparisons of nutritional data
obtained from Procedure One. Complete evaluations cannot adequately be
done until after all samples and data have been analyzed at which time the
final report for this study will be written.
1

Browse - a broad-leafed

woody plant, shrub, bush or tree of small stature.

2
Forb - a broad-leafed

herbaceous

plant.

�-82-

Table 2. Primary forage species identified in fecal samples of bighorn
sheep collected from the Buffalo Peaks, Pikes Peak and Trickle Mountain
ranges, June 1971 - September 1972.

Plant Species
Agropyron spp.

Buffalo Peaks
0.6 -

7.4

Percent (Range)
Pikes Peak
Trickle Mountain
0.8 - 10.3

Bouteloua gracilis
Carex spp.

1.1 - 29.9
2.7 - 53.6

7.8 - 91. 8

Danthonia parryi

0.1 - 6.7

Deschampsia caespitosa

6.0 -

Festuca spp.

13.0 - 66.8

Muhlenbergia spp.

0.1 -

6.8
0.5 - 57.6

5.6

0.7 - 52.9
0.1 - 56.6

Oryzopsis hymenoides
Poa spp.

4.2 - 19.2

0.2 - 11.0
2.4 - 10.5

0.5 - 14.7

3.5 - 15.2

Stipa spp.

2.5 - 10.7

0.1 -

Artemisia frigida

0.1 -

7.7

0.2 - 75.4

0.2 -

7.4

0.1 - 14.1

Cercocarpus montanus

12.3

Holodiscus dumosus
Pinus spp.
Salix spp.

0.7 - 42.0
0.4 -

6.8

0.1 - 28.8

27.6

35.1

Yucca spp.
Shrub (not identified)

5.4

0.5 -

9.9

0.3 - 15.7
8.4 - 53.4

0.2 - 68.3

0.7

40.4

�GRASSES
100
90

\

\\

&gt;- 80
••••

-.. --

\

\

Q

W

60
50

&gt;
-

•.••40

&lt;t

..J
W

,"

,,
,,
,,
,
.',
,

\

It'
\

\ I

\

\

II

~:
I

\

I

•........ ' ....

.•..•..,,.,"

"---- _, ,,
....••.•...••.._--, "

,;
;

,,\"

"

,,'9
"

I
I
I

\

I

'\

••.••••
J.

30

;"

\

20

\V /

I
I

Buffalo

Pk.

Pikes Pk.

/

------

Trickle

Mtn. ---

June

July

10

June
1971

July

.

Aug

Sept

Nov

-.on

I

00
L,..)
I

I

a::
~
0

,

..... ...

\

,

GRASSLIKE

r+r=&gt;:
.,'4';
.•.
,
I
......I
'

l,

\
\"

•..

~

\

-/) 70 \
C
Z
W

,,,"""
,,"
,,"',

\

and

Mar

Aug

Sept

1972

N;ONTH
Fig. 1. Use of grasses and grass-like forages by bighorn sheep on Trickle Mountain, Buffalo Peaks
and Pikes Peak ranges, June 1971 - September 1972.

�SHRUBS
100

90

r-.

&gt;- 30

f-

I
I

C/) 70

z

~

60

I
I
I
I
I

W 50

&gt;
I-

&lt;!

--

,~

'

I

/\
/

/

a::

~ 20]
o

10

/
I

I
I
I

Pikes

\

Trickle

\\

I

0:&gt;
.l:"I

I
"-

,

,", \
,
,

,,
,,
.•...••.

•..
.-------------,-------------,---------

1971

Mtn. ---

\ ..
\/ ',,

"-

Aug

---

\

\ \

"-

July

---

\

''''''

June

Pk.

\

\

'I •..

/

Pk.

\

\

~\

/

»-&gt;:

\

\
\

I
I

-1
W 30

Buffalo

I

I

40

I

--.

.,

"-

Sept

Nov

Jon

Mar,

June

••••
-r••••••••••••
.,---- ••------ ••
Aug
Sept
July

1972

MONTH
Fig. 2. Use of browse species by bighorn sheep on Trickle Mountain, Buffalo Peaks and Pikes Peak
ranges, June 1971 - September 1972.

�-85-

Accuracy of Fecal Microanalysis
Technique in Determining Food-Habits
Force feeding test diets directly into the rumen through a fistula proved
to be an effective means of feeding a diet of known composition to an
animal.
The method eliminated any selectivity of food intake by the
animal which might have occurred if the diets were offered free-choice or
in a cafeteria type feeding trial. During the trials rumination appeared
normal, the animal remained in good flesh and quite often was observed
contentedly chewing cud. Feces eliminated were a normal size and consistency.
Fecal samples collected after the animal had been fed the test diet for 10
and 12 days during each trial, contained little identifiable plant-residues a few Artemisia trichomes and a few unattached trichomes which may have been
from Salix, Cercocarpus or Purshia.
The bulk of material was unidentifiable
woody-stem material (Hanson, letter or 20 Sept , 1972).
Similar results
were reported by Zyznar and Urness (1969), who fed 17 tree, shrub and
herbaceous plant species separately to captive mule and white-tailed deer.
They found only low percentages of recognizable plant residues in fecal
samples using microanalysis techniques.
Unfed samples of the test diets were also analyzed by microanalysis techniques as an additional test of the method's accuracy in identifying plant
species in the diets and estimating percent composition.
All species in
the diets were identified.
However, microanalysis results include 2.1 percent Chrysothanmus sp. and 0.2 percent Pinus sp. in Diets I and II respectively (Table 3). These species were not formulated in the respective diets.
Estimates of the relative percentages of some plant species in the diets
were also highly over or underestimated
Crable 3). Stewart (1967) questions
validity of the microanalysis technique for estimating relative percent of
plant species in an animal's diet due to some forage plants fragmenting
more easily than others. We found, after feeding a number of perennial
grasses to African ungulates, that some plants easily fragmented and consequently appeared more important in their diets although intake was the
same or less than that of other species.
These results question usefulness and accuracy of fecal microanalysis
techniques for detenning food-habits of ruminants.
Reasons why no identifiable plant residues were found in fecal sanlples from these trials are
unknown.
Other investigators (Regg 1961, Kiley 1966, Lay 1965, Stewart
1967, Zyznar and Urness 1969) have used the method, with varying success,
to identify plant residues in feces from other ruminants.
Differences in
digestive capabilities among various species of ruminants studied during
these investigations and the domestic goat used during these trials may
account for differences of results obtained.
Digestion may be more complete in ruminants possessing relatively large capacity digestive organs
such as the goat, compared to ruminants with small organs.
Easily digestible
plants or plant parts may have been completely digested by the goats
digestive system leaving only undigestible wood-stem material to be eliminated in feces.

�-86-

Table 3. Composition
(percent dry weight basis) of two test diets
compared to the estimated relative composition determined by microanalysis.
(y/x is ratio of known percent (y) to estimated percent
(x ) ; values&gt; 1 are overestimates
and-C 1 underestimates).
I

\\

Plant Species

Compo

Diet I
Estimated

y/x

Compo

Artemisia

tridentata

9.3

6.4

0.7

5.0

10.7

2.1

Artemisia

frigida

14.1

42.1

3.0

10.0

41.6

4.2

9.7

1.3

0.1

25.0

5.8

0.2

0

2.1

9.9

8.6

0.9

Cercocarpus

montanus

Chrysothamnus

nauseosus

Diet II
Estimated

y/x

Medicago

sativa

35.6

25.6

0.7

10.7

11. 7

1.1

Purshia

tridentata

9.3

2.1

0.2

24.9

10.0

0.4

22.1

20.4

0.9

14.5

11.4

0.8

0

0

0

0.2

Salix nivalis
Pinus sp ,

Accurate food-habits data are essential for nutritional studies involving
any species of animal.
Based on results of these trials, application of
fecal microanalysis
techniques for determining food-habits of bighorn sheep
in our current studies cannot be reliably used.
However, determining foodhabits of free-ranging wild animals is extremely difficult and to date
fecal microanalysis
offers the best approach to the problem.
It is recommended therefore, that use of the technique be continued and further investigations be made to clearly define its application and limitations.

LITERATURE

CITED

Association of Official Agricultural Chemists.
analysis.
10th Ed. Wash., D.C.
957 p.
Harrington, H. D. 1954. Manual
Denver, Colo.
652 pp.

1965.

Official

of the plants of Colorado.

methods

of

Sage Books,

Hegg, O. 1961. Analyses of big-game droppings to determine their dietary
composition in Swiss National Park.
Revue Suisse de Zoologie 68 (12):
156-165.
(Transl. by J. J. Stransky, U.S.D.A. Forest Service, Southern
Forest Exp. Sta., 9 pp.).

�-87-

Kiley, M. 1966. A preliminary investigation into the feeding habits
of the waterbuck by faecal analysis.
Jour. E. Africa Wildl.

4:153-157.
Lay, D. W. 1965. Fruit utilization by deer in southern
J. Wildl. Mgmt.
29(2):370-375.

forests.

Shepherd, H. R. 1972. Game range investigations.
Colo. Div. of
Wildlife. Game Res. Div., Fed. Aid Proj. W-lOl-R.
Game Res.
Rept. ,'July.
Sparks, D. R., and J. C. Malechek.
1968. Estimating percentage dry
weight in diets using a microscope technique.
J. Range Mgmt.

21(4):261-265.
Stewart, D. R. M. 1967. Analysis of plant epidermis in faeces; a
technique for studying the food preference of grazing herbivores.
J. Appl. Exol.
4(1):83-111.
Van Soest, P. J. 1963. Use of detergents in the analysis of fibrous
feeds.
II. A rapid method for the determination of fiber and
lignin.
J. Assn. Official Agr. Chem. 46(5):829-835.
______ , and R. H. Wine.
1967. Use of detergents in the analysis of
fibrous feeds.
IV. Determination of plant cell-wall constituents.
J. Assn. Official Agr. Chem. 50(1):50-55.
Zyznar, E., and P. J. Urness.
1969. Qualitative identification of
forage remnants in deer feces. J. Wildl. Mgmt.
33(3):506-510.

Prepared

by

,Z:::··&lt;L-dfJ}H??U"t«·tl!

Gene G. Schoonveld
Asst. Wildlife Researcher

��January,

-89-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-4l-R-22

Bighorn

Work Plan No.

1

Job No.

Job Title

Manipulation

Period Covered:
Personnel:

Sheep

&amp; Mountain Goat Investigations
17

of Vegetation

on Bighorn

Sheep Ranges

June 1, 1971 to May 31, 1972

Thomas N. Woodard

and George D. Bear

ABSTRACT

Test plots were established on two study areas, Cebolla Creek and alpine
range.
Nitrogen and phosphorous fertilizers were applied to designated
plots.
Nitrogen was applied at the rates of 0, 30, 60 and 90 pounds per
acre. Phosphorus was applied at the rate of
and 30 pounds per acre.
A herbicide (2,4-D) will be applied in the next work segment.

°

Preliminary sampling indicated a 16-inch quadrat should be used for the
vegetative frequency sampling on Cebolla Creek, while a split-sampling
system (I-inch and 8-inch quadrats) should be used in the alpine •
.A Neal Electronics Herbage Meter was used to determine herbage production.
Sampling indicated a minimum sample of 40 plots per treatment should be
used in the alpine and 39 plots on Cebolla Creek. There was a strong
correlation between green weight of the forage and the meter readings;
.95 in the alpine and .81 on Cebolla Creek.
There was very little bighorn use on the study areas; likely due to minor
snow accumulation on the winter ranges.

1973

��-91-

MANIPULATION

OF VEGETATION

ON BIGHORN

SHEEP RANGES

George D. Bear

P. S. OBJECTIVE
Improve the herbage
on selected bighorn

yield, vegetative density
sheep ranges in Colorado.

and vegetative

composition

SEGMENT OBJECTIVES
1.

Alter the quality and quantity of forage on specific study areas with
varied treatments of chemical fertilizers and herbicides.

2.

Determine the effect of each treatment
and chemical content of plant species.

3.

Determine grazing preferences
to each treatment.

METHODS

on the composition,

of wild bighorns

production,

on the area in respect

AND MATERIALS

The study area is located about 35 miles south of Gunnison, Colorado.
The
resident bighorn herd winters in two areas: in a low (8,000 feet elevation)
bunchgrass and ponderosa pine type, and in a high (11,000 feet elevation)
alpine type. The following work was or will be conducted on each winter
range:
1. Bighorn preference and response of the vegetation will be measured
on plots treated with 2,4-D (herbicide), nitrogen, and phosphorus.
Two
levels of treatment (0 and 2 lbs./acre) will be evaluated for 2,4-D; four
levels of treatment (0, 30, 60, and 90 lbs./acre) for nitrogen; and two
levels of treatment (0 and 30 lbs./acre) for phosphorus.
These will be
evaluated in all possible combinations, or a 2x4x2 factorial for a total
of 16 treatments, as outlined in the following table:

�-92-

2,4-D

Treatment
Nitrogen

Phosphorus

1

0

0

0

2

0

0

30

3

0

30

0

4

0

30

30

5

0

60

0

6

0

60

30

7

0

90

0

8

0

90

30

9

2

0

0

10

2

0

30

11

2

30

0

12

2

30

30

13

2

60

0

14

2

60

30

15

2

90

0

16

2

90

30

Treatment
Number

Treatment plots were established, measuring 50 feet by 100 feet.
Each corner was marked with a metal fence post. These markers were numbered
to assist in identifying plots when bighorns are grazing on them. Three
replications were randomly established on each study area. Each replication
consisted of the 16 treatment plots listed above.
Plots in each replication
were arranged in a rectangular area eight plots long and two plots deep
(see following diagram).
Randomized selection of treatment for each plot
was determined by using a table of random numbers (Huntsberger 1961). The
arrangement is as follows, with the corresponding treatment number (from
preceding table) listed in each plot:

�-93-

Replication

No. 1

Replication

No. 2

Replication

No. 3

Ammonium nitrate and treble super phosphate were used as sources
for nitrogen and phosphorous treatments.
Approximately 0, 90, 180, and 270
pounds of ammonium nitrate were applied to obtain corresponding nitrogen
treatment levels of 0, 30, 60, and 90 pounds per acre. These compounds
were applied in the "pellet form" last fall before the ground froze, using
a whirling broadcaster.
The herbicide will be applied in the spring of 1972, as soon as
the ground thaws (next work segment).
This chemical will be applied with a
back-pack sprayer.
2. Plant Identification.
Representatives of all plant species occurring
on the study area were collected and dried.
Collections were made during the
flowering and fruiting stages to assist in identification.
Permanent mounts
of each species were made on cards measuring 8-1/2 X 11 inches.
Frequency Sampling.
Transects will be established on each plot to
determine any changes in vegetative composition which may result from the
various treatments.
Work here was directed toward establishing the sampling
procedures needed to detect changes in vegetative composition.
Temporary
transects were established in several plots to test for the proper quadrat
size to be used. The following quadrat sizes were tested on the Cebolla Creek
winter range:
4, 8, 12, and 16 inches square. Quadrats measuring 1, 2, 4, and
8 inches square were tested on the alpine winter range.

�-94-

Frequency percentages were derived for all plant species encountered
on the quadrats.
To be considered as occurring on a given quadrat, the
center of an individual plant or one-half of its basal area must be inside
the quadrat.
An "individual" for bunchgrasses shall be a clump; for a single
stem species, such as wheatgrass, it will be individual stems; for cactus it
shall be a lobe; and for shrubs and forbs it will be the basal area of all
stems.
Frequency (p) is defined p = mln, where m is the number of quadrats
containing a given species and n is the total number of quadrats observed in
each treatment.
Yield Plots.
Quantity of forage produced on the treatment plots
shall be determined by using a Neal Electronics Herbage Meter.
The herbage
meter measures an area 24 inches by 12 inches, which will be designated as
a yield plot. A series of meter readings were taken on each replication to
determine the variability to be expected and the sample sizes needed.
In addition to the herbage meter readings, the vegetation on every
fifth plot was clipped and weighed.
All green vegetation on these plots is
clipped off at the root crown and placed in a labeled paper sack. These
samples are taken to the laboratory, oven dried, and then weighed to the
nearest gram.
The weights are then used to compute regression equations
for converting herbage meter readings into pounds of forage per acre. Again,
work in this segment is being done to check for variability.
Chemical Analysis.
Vegetative samples shall be analyzed for phosphorus, potassium, and protein content.
This work shall be done at the Fort
Collins Research Center Laboratory.
Since the study areas are bighorn sheep
winter ranges, the vegetation should be sampled at the completion of the
growing season, again in mid-winter, and in late winter.
Samples of the most
common plant species will be collected from the treated areas.
Since the
fertilizers were applied in September, following the growing season, the
chemical composition of the plant could not be changed, thus plant collections
were not made in this segment.
3. Bighorn preference or selection of treatment plots shall be determined.
Numbers of bighorn sheep using the study areas are to be recorded
during scheduled observation periods.
Again, since the vegetation was not
affected by the late application of the fertilizers, observations were limited
to times when other work was being conducted in the area.

RESULTS

AND DISCUSSION

The replications and treatment plots were surveyed and marked with metal
fence posts, as outlined in the procedures.
Fertilizer was applied on the
designated plots.
In using plant frequency percentages to detect changes in vegetative
composition the quadrat must be large enough to adequately represent the less
abundant plant species, yet the most abundant species must have frequency
percentages somewhat below 100 percent to detect increases that may occur.

�-:95-

Visual evaluation of the data in Tables 1-3 indicated the l6-inch quadrat
to be the proper size for the Cebolla Creek study area. Data in Tables
4-6 indicated a split-sampling scheme should be used on the alpine; a
quadrat measuring 1 inch x 1 inch for Trifolium nanum and Carex sp., and
a l6X16-inch quadrat for all other species.
It was decided that four transects should be established on each treatment
plot.
The transects are to be parallel to the base of the plot with
approximately a 10-foot spacing between transects.
A total of 15 quadrats
are to be located at 5-foot intervals along each transect, or a total of 60
quadrats for each treatment plot.
Each quadrat is to be permanently marked
with a metal stake.
Data collected with the herbage meter indicated 40 quadrats were needed on
each treatment plot in the alpine study area for a statistically valid (P
.10) sample, and 30 quadrats were needed on the Cebolla Creek study area.
The 60 quadrats established on each treatment to determine frequency will
also be used to locate yield plots.
Obviously the 60 plots is considerably
greater than the recommended numbers given above; however, the vegetation
may be changing due to treatment and larger samples will be needed as
variation increases.
There was a strong correlation between herbage meter readings and forage
(weight) on the plots (Fig's. 1-4). The correlation coefficient for green
weight to meter readings in the alpine was .95 and the dry weight correlation
coefficient was also .95; the regression equations were: green weight = -19.53
+ 3.24 (meter reading), and dry weight = -3.11+ 1.06 (meter reading).
The
correlation coefficient for green weight to meter readings on Cebolla Creek
was .81 and the dry weight correlation coefficient was .76; the regression
equations were: green weight= -13.96 + 1.71 (meter reading), and dry weight
= -7.38 + 1.00 (meter reading).
The Cebolla Creek study area is very rocky,
thus the correlation coefficients were lower than those obtained on the
alpine.
Sheep-use on the study areas appeared to be extremely light during the 1971-72
winter.
Bighorns fed on the adjacent areas near the alpine plots, but were
never observed on the plots.
Snowfall was below normal in this area and the
entire south-exposure of the ridge below the plots was relatively free of
snow all winter.
Five to seven inch drifts had accumulated on plots 7 and
15 (Replication 1) and on plots 10 and 11 (Replication 2) in late February;
otherwise, the study area was relatively free of snow accumulation.
The
Cebolla Creek area was also free of snow accumulation
(except immediately
following a storm) throughout the winter.
Three rams were observed on plot
10 (Replication 1) in late May.
The plots were fertilized after the growing
season last fall so there could not have been a change in the vegetation
to attract the sheep.
Snow depths were so minimal sheep were not forced
to concentrate in the area where plots are located.

�Table l.

Frequency of plant species (percent) on the Cebolla Creek Study Area, Replication No.1,

1
4

8U

Plot Number and Quadrat Size (Inches)
3
4
5
4
8
12
4
8
12
4
8
12

4

2
8

12

8

11

15

3

5

-

-

8

-

-

-

-

-

3
50

4
15

4
34

7
54
1
70

4
16
1
1
8

-

-

-

-

3

3

4

-

3
1

3
1

-

-

-

-

-

1971.

6
8

12

4

7
8

12

48

12

-

-

7
1
26

8
4
57

18

4
67

3
1
18

15

1
45

-

12

1
18

5

16

24

15
3

32
3

47
3

18
4

36
9

51
12

33
1

43
4

-

-

1

-

3
1

54
4
1
7
3

4

8

Grasses
Agropyron sp.
Bouteloua gracilis
Carex sp.
Festuca arizonica
Festuca sp.
Koe1eria cristata
Muh1enbergia montana
Muhlenbergia richardsoni
Oryzopsis hymenoides
Sitanion hystrix
Stipa comata

7 13
3
4
- 4
5 22

-

-

-

37

54

-

-

-

-

18
8
8
34

1
65

-

1

3
3

1
3
4

-

-

-

-

-

15

30

-

-

22

1
38

1
57

-

-

1
1

1
3

-

-

-

-

-

-

1

-

-

-

-

25

-

-

53

-

-

-

-

4
7
34
1
1
22

-

5
8
51
3
5
30

18

34

-

-

4
3
18

4
4
33

4
5
45

-

4
53

-

-

-

-

-

1

1

4

-

-

1

1
1

-

1

-

-

-

1

-

-

-

-

-

-

-

-

-

-

1

-

-

1

1

1

1

1

-

Forbs
Androsace sp.
Antennaria parva
Cruciferae
Erigeron sp.
Eriogonum sp.
Geranium sp.
Hymenoxys richardsoni
Mertensia sp.
Penstemon sp.
Potentil1a sp.
Senecio sp.
Unid. forbs

-

-

1

-

1

1
-

3

5

-

-

3
1

3

-

-

1

-

-

8

-

-

4

-

-

1

1

1

-

-

-

-

1

-

-

9

-

-

-

1
3

..,

1

11
1

-

-

-

-

1

1

1

-

-

-

1
1
1

1

-

-

1

-

3

-

-

-

-

1

3

-

-

-

4

8

-

-

-

5

1

4

-

-

13
1

-

-

-

-

-

-

-

-

-

1

-

-

-

-

-

-

-

1

-

1

1
1

-

-

3

-

-

4

4

-

-

-

-

1

3

-

-

-

-

-

1

1

4

-

-

1

1
1
1

1
1

-

-

1
1
1

-

-

-

-

-

-

3

-

-------------------_._----------------------------------------------------------------------------------------------------------------------

,
,'"

'J:)

�Table 1. Frequency of plant species (percent) on the Cebolla Creek Study Area, Replication No.1,
Plot Number and ~adrat
4
3
12
8
4
12
8
4

4

2
8

12

-

-

-

-

-

-

-

-

37
5
1

16
3
4

28
4
7

40
9
11

15
3

24
4

34
4

16
4

28
4

-

-

-

-

-

-

-

-

-

1

1

1

1

4
1

4
1

4
1

1

1

-

-

4

1
8

12

-

-

-

8
3
1

21
3
1

Size (Inchesl
5
12
8
4

1971 (continued).
6

7
8

8
8

12

12
1

1
32
3

1
46
4

-

-

3

12

4

33
4

3
46
5

-

-

4

8

12

4

-

1
28
5

1
38
15

-

-

12
1

15
1

1

1

1

-

Shrubs
Artemisia dracuncu1us
Artemisia ~rigida
Chrxsothamnus parryi
Leptodacty1on pungens
Physocarpus sp.
Rhus trilobata
Ribes cereum
-----

1

-

-

-

-

-

-

-

-

-

3

1
45
5
1
3

-

-

1

3

-

-

18

-

34
4

47
5

1
1

1
3

1
4

-

-

-

-

-

I

'"-..j
I

�Table 2.

Frequency of plant species (percent) on the Cebolla Creek Study Area, Replication No.2,
1
4

812

4

2
8

11
1
4

15
1
9

15
1
22

l3

21

-

-

-

12

-

-

5

-

41

62

-

71

25

-

40

-

58

-

-

5

5

-

-

-

-

1

-

9

12

28

-

-

12

28

Plot Number and Quadrat Size (Inches)
3
4
5
4
8
12
4
8
12
8 12
16

1971.
6

8

12

16

8

7
12

8
1
20

11
1
26

15
1
42

4
3
28
12
7
4

8
5
45
21
16
5

-

-

1
22

1
30

1
4

1
8

16

8
SIT

16

G~
Boute1oua gracilis
Carex sp ,
Festuca arizonica
Koe1eria cristata
Muh1enbergia montana
Muh1enbergia torreyi
Oryzopsis hyrnenoides
Poa secunda
Sitanion hystrix
Forbs
Antennaria parva
Cruciferae
Erigeron sp.
Eriogonum sp.
Geranium sp.
Hyrnenoxysrichardsoni
Lappu1a sp ,
Penstemon sp.
Potent::t11a
sp.
Senecio sp.
Unid. forbs

-

-

-

-

1

1

-

-

1

-

-

-

1

1

-

20

1

-

-

1
1
l3

1
3
25

-

-

-

-

9

22

34

4

-

-

-

-

-

1
1

1
1
1

-

-

-

-

-

-

-

1

1
1
1

-

1

1
1
1
1

-

-

1
21
4

1
30
5

-

-

16
3

21
7

38
9

-

-

-

-

-

-

-

-

-

12
8
50
4
57

-

-

-

-

3
9
3
3
20 34
3
16 38

-

-

-

-

-

-

1

1

-

-

5

8

-

-

17
8

30
1
20

-

-

1
5

5
8

-

-

-

-

9
1
48
8
41

-

8
18

-

8

-

34

-

-

-

-

-

46

59

75

24

37

57

-

-

-

-

5
1

-

1

-

-

-

-

-

-

1

1

-

-

-

1

1

-

1

-

-

-

-

-

-

-

-

-

-

-

-

-

17

22

1
3
1

-

8

-

43

-

-

-

-

-

-

-

-

12

17

16

21

30

1

1
1

-

-

-

1

4

4

-

-

-

21

-

-

1

3
3

1

3

3

-

-

-

-

-

-

-

-

1

1

3

3

3

-

1
59
4

1
78
5

36
1

57
4

74
5

1
3

1
3

-

-

1

-

-

-

-

-

-

-

3

-

-

-

-

-

-

-

1

-

3

-

-

-

9
1
51
3
33

1
11
5

-

-

8
4
7
9
1
1
65 24 38
25
- 3
18 18 24
9
- 1 1
1
43 11 16

Shrubs
Artemisia dracuncu1us
Artemisia frigida
Chrysothamnus parryi
Chrysothamnus viscidif10rus
LeE!Qdacty1on pungens

-

7
3

-

-

-

-

-

-

-

-

-

-

16

-

38

-

-

-

-

62
1

21

37

54

42

68

-

87

-

-

-

38
3

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

58
4

78
5

36

-

-

-

-

-

1

4

4

3

I

'!)

ex&gt;
I

�Table 3. Frequency of plant species (percent) on the Cebolla Creek Study Area, Replication No.3,

1

Grasses
Boute1oua gracilis
~sp.
Danthonia parryi
Festuca arizonica
Koe1eria cristata
Muh1enbergia montana
Muh1enbergia richardsoni
Muh1enbergia torreyi
Oryzopsis hymenoides
Sitanion hystrix

8 12--r6

8

45
40

21
4

25
30

34
34

-

-

-

-

16

25
1
49

30
1
59

15
3
32

29

1971.

2
12

16

8

Plot Number and Quadrat Size (Inches)
5
4
3
8
16
8 12
16
8 12
16
12

29
5
1
20
3
43

38
7
1
28
3
54

15
8
1
21

21
22
3
37

26
24
3
45

-

-

22

32

-

1
9
18

-

-

-

-

-

-

4

4

1

5

-

-

4

-

-

-

16

18

29

15

21

7
1
36

-

-

4
5

-

4
12

16

32
9
3
58

17

21

26

11

13

15

1
20

1
33

17

29

-

-

4
29

-

-

-

43

51

55

-

-

-

-

30
1

53
1

62
1

33

16

22

29

5

15

29

1

-

-

-

43

40

53

67

-

-

15

25

-

12

24
8
3
42

21

-

8

17
3

36
20
3
41

12

13

-

-

-

15

-

-

-

8

16

20

29
17
1
29

21
12

7

6
12

-

812

16

-

-

53

66

-

-

-

-

-

59

72

80

-

-

-

66
7

80
8

1
42
1
87
9

12

13

18

3

12

16

33

I

'-D
'-D

Forbs
Cruciferae
Erigeron sp.
Lappu1a sp.
Potenti11a sp.
Senecio sp.
Unid. forbs

I

1

-

1

3

3

3

-

-

-

1
1

3
1

4
3

-

-

-

30
3

50
4

68
5

-

-

-

-

-

-

-

-

-

1

1

1

5
8
1

7
9
1

1
66
3

3
84
3

4

-

-

1

-

1
1

-

1

-

7

1

-

-

4

3
7
7
1
1
1

3

- 3
- - 9 16
- -

1

1

1

-

1

-

-

-

1

-

-

-

-

24
1

1

3
1

3
1

8

1
47
13
1
1
7
9

1

9
7

1
29
7

1
46

1

1

1

3

-

5

1

3

5

-

1

~

13

13

1

1

1
1
1

1
9

1
15

1
21

-

11

-

-

-

26
3

1
43
4

5
59
4

3

1

1

1

3

3

3

-

-

Shrubs
1
Artemisia dracunculus
45
Artemisia frigida
3
Chrysothamnus parryi
Chrysothamnus viscidif1orusEchinocactus sp.
Leptodacty1on pungens
Rhus tri10bata
RibesCer~
-

-----

-

-

-

-

-

1
24
4

1
30
8

1

-

-

1

3

4

3

4

7

21

22

37

13

36

53

�Table 4.

Frequency of plant species (percent) on the Alpine Study Area, Replication No.1,

1

2

4

8

1

rIot Number and Quadrat Size (Inches)
2
3
15
2
4
8
1
2
4
8
1
2
4

20
8
15

30

49

13

62
37
57
15
7
7

18
7
11

22
12
30

45
20
51

-

1

3

1

1971.

16
8

1

2

4

8

45
5
25

71
7
38

87
13
45

87
18
55

-

-

-

1

Grasses
Carex sp.
Festuca ovina
Poa sp.
Sitanion hystrix
Trisetum spicatum
Unid. grasses

1

-

22
18 26
1
1
5
3
- 1

-

-

67
34
79
3
4

-

8

1
9

3
1

-

9
7
18
4

24

49
29

59

81

-

40
7
1
3

-

74
9
11

-

96

11

4

7

20

99
1
33

-

-

-

-

- -

3

Forbs
Achillea 1anu1osa
Anemone sp.
Androsace sp.
Artemisia scopu1orum
Caltha 1eptosepa1a
Cerastrium beeringianum
Dryas octopetala
Erigeron sp.
Geum turbinatum
Hymenoxys grandif10rus
Mertensia bakeri
Phlox caespitosa
Po1ygonum bistortoides
Po1emonium viscosum
Potentil1a diversifo1ia

I

t-'

-

-

-

-

-

9
1
4

26
1
11

-

-

17

26
3

-

-

-

2

-

-

-

-

-

-

-

-

-

1

1

-

-

-

44
3
22
3

1
68
15
36
12

-

-

-

-

40
8
1
5

59
20
1
24

24
3

38
4

3

4

-

-

9
2
9
1

-

17
8
21
1

3
46
15
34
3

0
0

1.

1

3

4·

-

1

1

4

1

3
1

-

-

1

4
3

-

-

-

-

3
14
11
5

1
3
3
1

-

1
4
7
4

-

-

49
17

4
66
33
43
7
1
62
40

9
1

12
3

9

22

1

1

I

1

-

1

5
1
1
5
34

1

3

-

-

11
1
1
11
58

22

1

1

3

-

-

3

-

3

7

1
16

11
22
75

4
18

5
46

20
71

30
92

3

-

-

60
29

-

-

18
8

4
4
25
20
9
1
25
26

70
59

83
88

88
96

45
13

3
45
34

4
49
70

5
50
88

11

26

-

-

-

-

-

-

-

3

4
13

-

------------------------------------------------------------------------------------------------------------------

�Table 4.

Frequency of plant species (percent) on the Alpine Study Area, Replication No.1,

1
1

2

4

8

1

1
9

4 16
22 30
1
3
15 24
1
1
11 16
4
4
75 82

3
17
1

Plot Number and Quadrat Size (Inches)
15
3
2
2
4
1
2
8
1
4
4
8
2

1971 (continued).

8

1

16
4
2

-

-

-

3
1
16
57

8

Forbs (continued)
Potenti11a nivea
Pseudoc~oEterus sp.
Rununcu1us inamoenus
Saxifraga sp ,
Sedum sp.
Si1ene acau1is
Solidago sp.
Trifolium nanum
Trifolium Earryi
Tro11ius a1bif1orus
Unid. compositae
Unid. forbs

1

1
15
1
9

-

-

1

7
3
57

45

-

-

-

-

1

1

1

3
3

-

- -

-

-

-

-

3
4

7
12

17
22

36
30

1
1
12

1
1
25

1
4
41

-

-

-

-

-

-

-

1

-

16
6
13

32
17
29

40
34
55

47
53
76

7
1
3

16
3
4

32
3
8

38
4
25

3
1

-

- -

-

-

-

1

1

5
25
1
12

13
41
1
24

36
51
3
36

4
17
9

15

-

-

-

-

-

15
1
51

20
3
72

29
4
84

41
8
90

3

5

-

-

30

46

13
37
1
20
1
15
1
67

-

-

-

-

-

-

-

22
54
4
34
3
20
4
83
1

1
3

1
5

- - 1

3
1

-

-

5
25

- -

- -

I

t-'
0
t-'
I

�Table 5.

Frequency of plant species (percent) on the Alpine Study Area, Replication No.2,
Plot Number and Quadrat Size (Inches)
1
2
4
8
1
2
4
8
1

1

2

55

70

-

-

15
18

26
25

-

-

88
1
59
38
1

1971.

3
2

4

8

Grasses
Carex sp.
Deschampsia caespitosa
Festuca ovina
Poa sp.
Trisetum spicatum

90
4
75
55
9

36

53

66

71

55

76

91

91

29
11
3

45
18
4

65
26
7

91
42
22

16
13
3

29
21
7

42
36
17

63
51
26

-

-

4

5

8

-

-

-

21
17

-

-

5
1
61
57

-

8
9

5
1
37
34

-

-

17

25

44

-

-

78

-

-

21
9
1
20
1

37
22
1
36
1

1

1

-

4

-

-

13
1
7

28
1
8
3
25

54
3
15
3
41

-

-

-

36
20
1

8
5
25
25
1
34

12
13
41
49
3
66

Forbs
Achillea 1anu1osa
Androsace sp.
Artemisia scopu1orum
Cerastrium beeringianum
Erigeron sp.
Erigeron pinnatisecta
Eritrichium e10ngatum
Gentiana sp.
Geranium sp.
Geum turbinatum
Hymenoxys grandif10ra
Lycopodium sp.
Mertensia sp.
Phlox caespitosa
Po1emonium viscosum
Po1ygonum bistortoides
Potenti11a nivea
Pseudoc~opterus sp.
Rununcu1us inamoenus
Saxifraga sp ,
Sedum sp.
Si1ene acau1is
Trifolium nanum
Trifolium parryi
Tro11ius a1bif1orus

1

5

5

7

-

-

-

-

21
7
15

32
12
1
21

45
37
8
36

-

66
62
12
50

-

-

-

4

5

7

-

13

-

-

-

13

4
1

25
1
1
5
1

38
4
3
7
3

-

-

11

-

51
8
7
7
9

-

-

12

-

-

-

4

11
4
7
1
18
3
3
24
15
28

36
17
8
5
21
8
9
45
18
47

-

1

3
1

11
8

-

7

5
1
7

-

-

8

12
1
1
18
9
16

1
13
4
7

5

4

4
3
11
7
1
8

-

5
3
12
12
1
13

12
4

-

-

5

13

-

-

21
1

66
49
5
49
3
1
1
30
3

4
9
3
1
3

8
16
1
3
3
4

13
30
5
13
7
15

26
49
8
37
17
18

1

4

11
1

21
7
1
58
4
71

11
1

-

-

-

-

-

15

26

42
1
40

-

-

7

20

I
•....•

0

N
I

�Table 6.

Frequency of plant species (percent) on the Alpine Study Area, Replication No.3,

1971.

4

1
2

4

8

1

2

4

8

1

2

4

8

1

2

4

8

1

58
5
9
1
3

80
11
21
3
5

93
17
32
3
12

99
29
63
5
25

55
4
16

71
9
25

-

-

4

5

90
21
38
3
11

91
50
76
13
37

71
3
12
4
5

90
8
18
5
8

96
24
29
8
25

97
54
47
20_
58

51
5
5
5
4

72
5
11
11
9

80
24
20
22
17

86
54
33
51
33

-

-

-

-

-

42

59

1
15
11

3
7

12
20

25
30

7

-

-

-

-

-

1

12

22

1
8
47
3

-

-

9

-

-

3
5

7 15
1
1
7 13
1 9
21 32

30

-

7
1
1
1

17

-

7

13
1
43
50

-

-

3
1
24
24

-

8

16

-

-

-

-

1
1
1

1
3
4
7

1
3

1
5

4
7

28
9

Grasses
Carex sp.
Festuca ovina
Poa sp.
Trisetum spicatum
Unid. grasses
Forbs
Artemisia scopu1orum
Arenaria fend1eri
Cerastrium beeringianum
Dryas octopeta1a
Erigeron sp.
Erigeron pinnatisecta
Eritrichium e10ngatum
Geum turbinatum
Phlox caespitosa
Po1ygonum bistortoides
Potentilla diversifo1ia
Potenti11a nivea
PseudocymoEterus sp.
Saxifraga sp.
Sedum sp.
Si1ene acau1is
Trifolium nanum
Trollius a1biflorus
Unid. compositae
Unid. forbs

8
1

-

-

-

1

-

-

-

-

-

1
1

3
3
1

9
4
1

-

-

-

-

28
5
8
1

3
5
17
13

9
3

13
4
3

20
16
9

26
24
28

5
1

3
5
8

5
9
12

-

-

-

3
63
18

3
82
29

9
9
18
1
3
87
54

1

1

1

-

-

-

1
1
32
1

-

-

-

1
4

3
8

9
9

1
7

4
13

12
18

-

-

-

28
16

5
5
38
8
5
34
20

-

-

-

-

-

-

1
37
7

1
54
12

4
96
30
1

1
38
5

1
50
18

1
75
29

1
93
42

38
16

63
29

82
51

91
62

3
42
9

-

3
80
20
1

-

-

-

-

-

-

-

-

- -

-

1

-

-

- -

30
1

-

1
1

-

-

- - - - - - - - - -

-

I

~

0

w
I

�-104-

80

70

60

30

20

10

.

•

,

O~------~--------~-------L----~~----~_
10

20

Fig. 1.

Relationship

Cebolla

Creek Study Areat

30
METER READING

40

50

of meter readings to green weight of the forage,
1971.

�-105-

45
•

40

35

30

1.5
•

•
•

•••

•

10

•
••

••

• •
•

5

•
•
••

•

•

•

•

•
•

o

•

10

•

20

30

40

50

METER READING

Fig. 2. Relationship of the meter readings to the dry wejght of the forage,
Cebolla Creek Study Area, 1971.

�-106-

350

300

250
,-..
III

S
&lt;:"J

~

d

&lt;:»

~
;:r::

200

CJ

H

~
Z
r.il
r.il

~

CJ

150

100

50

25

75

100

125

METER READING
Fig. 3. Relationship of meter readings to green weight of the forage,
Alpine Study Area, 1971.

�-107-

•

•

50

---CI)

s
Ct!
~

C

'-"
E-&lt;

::r:
c 40

H

~
:&gt;-I
0::
Q
&lt;II

;z;

~

0

30

10

Fig. 4.

20

40
MEI'ER READING

30

50

60

Relationship of meter readings to dry weight of the forage, Alpine study

Area, 1971.

70

�-108-

LITERATURE CITED

Huntsberger, D. /1961. Elements of statistical inference.
Bacon, Inc., Boston. 291p.
(

Prepared by

~_~_'_~_~~~~~~--.--c-/-·
'__/_:_~_-~_~_'
_." "
George D. Bear
Wildlife Researcher

Allyn and

�January,

-109-

JOB PROGRESS REPORT

State of

COLORADO
W-4l-R-22

Project No.
Work plan No.
Job Title

Job No.
Control of Lungworm

Period Covered:
Personnel:

1

Bighorn Sheep &amp; Mountain

Goat Investigations

18
in Bighorn Sheep

June 1, 1971 to May 31, 1972

Robert Lange, Steven Steinert, Julius Klein, George Bear, Robert
Keiss, Charles Hibler.

ABSTRACT

Five bighorn sheep were captured near Glenwood Springs and placed in a
363 acre pasture at Colorado Division of Wildlife Little Hills Experiment
Station near Meeker, Colorado.
Two adult ewes were successfully treated
with Tramisol to test this drug's therepeutic effect against Protostrongylid lungworms.
Tramisol did not reduce lungworm larvae output in fecal
pellets of treated sheep. The experiment will be repeated under contractual agreement with Colorado State University personnel, with some modifications.

1973

��-111-

CONTROL OF LUNGWORM

IN BIGHORN

SHEEP

Robert Lange and George Bear

P. S. OBJECTIVE
Evaluation of potentially
bighorn sheep.

useful drugs in the control of lungworm

in

SEGMENT OBJECTIVES
1.

Establish

lungworm larvae output prior to treatment

2.

Determine the effect of therapeutic
orally to bighorn sheep.

METHODS

with drugs.

doses of the drugs administered

AND MATERIALS

Five bighorn sheep were transplanted from a semi-domesticated herd near
Glenwood Springs and placed in a 363 acre pasture at Colorado Divlsion
of Wildlife Little Hills Experiment Station near Meeker, Colorado.
These
sheep were used as experimental animals for this study.
This study was conducted by contractual agreement with Colorado State
University College of Veterinary Medicine personnel after it had been
approved for implementation in this project.
Therefore project personnel
(George Bear) duties were limited to assisting with initial capture of
the sheep, treating sick sheep during the summer, and assisting in administering Tramisol to the sheep. Robert Lange (student in Pathology) and
Steven Steinert (Research Assistant, Little Hills Experiment Station) conducted all research in this study.
1. The initial job proposal stated that the captive bighorns were
to be monitored for lungworm larvae output prior to treatment with drugs;
however, this portion of the study was modified to be compatible with Mr.
Lange's schedule at Colorado State University.
The sheep were divided into
two groups; those treated with the drug and those untreated.
The larvae
output of treated sheep was compared with that of untreated sheep and pretreatment output was compared with post-treatment output.
2. The initial proposal stated two drugs, Tramisol and Hygromycin B,
were to be evaluated; however, an insufficient number of bighorn sheep were
available for this study, therefore only Tramisol was evaluated.
Therapeutic
doses of Tramisol (8 mg./lb. B.W.) were administered orally in gelatin capsules to three ewes after they were tranquilized with M-99.
One yearling
ewe died five hours post-treatment.
The effects of treatment on the other
two ewes (adults) was estimated by larvae output in the feces.

�-112-

Fecal pellets were collected only when they were positively known to
have been dropped by an individual sheep under observation.
Each
pellet group was placed in individual envelopes and labeled with the
bighorn's name and date of collection.
Samples were then air-dried
and stored until they could be analyzed in the laboratory.
The
Baermann funnel technique was used to determine the lungworm larvae
per gram of feces.

RESULTS AND DISCUSSION
Two treated adult ewes and two untreated ewes, a yearling and an adult,
were monitored through the winter and spring.
No effects of Tramisol
on larvae output are evident (Table 1).

Table 1. Larvae output in treated and untreated
Hills Experiment Station, 1971-72.

Pre-treatment
Date
Collected

bighorn

sheep, Little

Larvae Output (larvae/gm. feces)
and Post-treatment
Adult Ewe
Adult Ewe
Adult Ewe
Yearling Ewe
(Boss)
(Smoky)
(Mom)
(Blackie)

11-5-71
11-15-71
11-16-71
11-29-71

147
204

12-11-71
12-12-71
12-13-71
12-14-71
12-18-71
12-19-71
12-20-71
12-21-71
12-30-71

58
42

treated

142

128
32
244

12
131

0
20

52
36
25

41
10

21
26

151

22

treated

2
170
159

42
14
45

56
84

1-6-72
1-13-72
1-27-72

40
103
10

245
113

234
2

39

2-5-72
2-14-72
2-29-72

59
230
60

27
128
133

222
387
153

16
43
65

81

-----------------------------------------------------------------------------

�-113-

Table 1. Larvae output in treated and untreated
Hills Experiment Station, 1971-72 (continued).

Pre-treatment
Date
Collected

sheep, Little

Larvae Output (larvae/gm. feces)
and Post-treatment
Yearling Ewe
Adult Ewe
Adult Ewe
Adult Ewe
(Boss)
(Smoky)
(Mom)
(Blackie)

3-4-72
3-5-72
3-6-72
3-7-72
3-8-72
3-9-72
3-10-72
3-13-72
3-14-72
3-15-72
3-16-72
3-18-72

508

290

153
49
18
86
1
61
230
81
123

36
112
153
24
33
105
186
5

4-5-72
4-17-72
4-19-72

13

5-13-72

bighorn

177
66
122
102
176
64
265
205
25
16
41

67
16
35
32
21
5
27
51
32
27

77

38
4

59

33

0

8

34

There are several possible reasons for the poor results.
(1) Inactive or
irregular quality of commercial Tramisol preparation, (2) Refractivity of
lungworms to Tramisol, (3) Inability of Tramisol levels (8 mg/lb. B.W.) to
penetrate lungworm lesions in adult sheep, and (4) Experimental errors such
as improper administration or collection of fecal pellets from wrong sheep.

Pr epared by __ -;::/.:..~_.:...._:'/'_L.,f~/::,.'~_~_:....&lt;_'_/_~-_~_'.A_/
__
George D. Bear
Wildlife Researcher

��January,

-115-

JOB PROGRESS

State

of

Project
Work

REPORT

COLORADO
No.

W-4l-R-22

Bighorn

Plan No.

1

Job No:

Job Title
Period

Mineral

Covered:

Personnel:

November

William

&amp; Mountain

Goat

Investigations

19

Block Experimentation

1, 1971 to April

H. Rutherford

Sheep

30, 1972

and Dale W. Stahlecker

ABSTRACT
.

.

Two separate brands of range livestock mineral blocks, the Moorman Special
Range Mineral Block and the Carey Range-Min Block, were placed on bighorn
sheep winter range at Tarryall and Georgetown.
Usage of these blocks was
noted by observation and by periodic weighing of the blocks.
Loss of weight
due to weathering was assessed by the use of controls.
The CArey block, containing about 50 percent salt (NaCl) , was invariably
selected by bighorn sheep in preference to the Moorman block, which contains
only about 20 percent salt.
Blocks placed in sites where sheep naturally
move received the most use, and blocks placed in well-drained
sites showed
the least deterioration
by weathering.

1973

��-117-

MINERAL

BLOCK EXPERIMENTATION

William H. Rutherford

Previous attempts to improve the welfare of bighorn sheep by providing
mineralized salt blocks have indicated that such dietary supplementation
has a place as a management technique, and should be continued.
Hopefully, studies of nutritional needs and deficiencies on bighorn sheep
range will eventually provide information on precisely what elements
need to be supplemented, and in what proportions.
At the present time,
however, it appears that mineralized salt blocks standardized by the
livestock industry for general range use offer the best available source
of supplementary minerals.
It was recognized that information on acceptance and use of such blocks by bighorns, and on efficient location and
placement of blocks, was lacking.
The study reported here is thus
limited in scope to an investigation of acceptance, use and placement,
and is intended only to provide guidelines for efficiency in providing
supplementation.

P. S. OBJECTIVE
To assess factors involved

in use of mineral

blocks by bighorn

sheep.

SEGMENT OBJECTIVES
1.

Determine relative
by bighorns.

2.

Determine

rate of use of mineral

3.

Determine

best location

preference

of various mineral

block formulae

blocks by bighorns.

and placement

of mineral

blocks.

METHODS AND MATERIALS
Materials consisted of mineralized salt blocks from two separate manufacturers, Carey and Moorman.
Initially, it was intended to test several
different mineral block formulations, but limitations on availability
dictated that the standard range mineral block supplied by each of these
two manufacturers would be the only two that could be tested.
The major differences between the two blocks are that the Carey Range-Min
Block contains from 50.0 to 53.0 percent salt (NaCl) while the Moorman
Special Range Mineral Block contains only from 17.5 to 21.0 percent salt;
and the Moorman block contains from 15.0 to 18.0 percent calcium while
the Carey block contains only from 7.5 to 9.0 percent calcium.
Levels of
other minerals, particularly the trace elements, are approximately equal
in the two blocks.

�-118-

Blocks were placed in several dissimilar types of terrain on two
separate bighorn sheep winter ranges during the winter of 1971-72.
Generally, the various locations consisted of open rocky ridge-tops,
open flat areas, timbered or brushy hillsides, and bottoms of small
watercourses, all of which showed past usage by bighorns.
In addition,
a location was selected on each of the two ranges where blocks could
be placed out of reach of all ungulates, but still exposed to weathering.
Usage of the blocks was noted at frequent, but irregular, intervals by
observers residing in the vicinity of the two winter ranges.
Each of
the ranges was visited four different times during the winter, and all
blocks were weighed with a set of spring scales accurate to 1/4 pound.

DESCRIPTION

OF AREAS

Winter ranges in the Tarryall and Georgetown areas were selected as sites
for this study.
Both are easily accessible, and bighorn sheep are usually
in full view of observers who travel the adjacent roads.

RESULTS AND DISCUSSION
According to some Wildlife Conservation Officers who have used the Moorman
block at various times in the past, acceptance by bighorn sheep has been
rather indifferent.
In a few cases it would be lightly used, but typically
the block could be placed where sheep would pass by it every day and it
would not be used during an entire winter.
Its main attribute seemed to
be that it withstood weathering quite well.
Accordingly, the Moorman block was used with some skepticism in this study,
and was not placed at every location selected as a station.
Rather, the
Carey block was placed at every station, and after a period of use by
bighorns, a Moorman block was placed next to the Carey block at the one station
on each winter range that showed the greatest use.
Usage of the blocks is presented

as tabular data with comments in Table 1.

Past experience has indicated that the higher the salt (NaCl) content of a
block, the more attractive and palatable it is to bighorns, to the point
that if bighorns are offered a pure white salt block they will use it to
the complete exclusion of any mineralized block if a choice exists.
Thus,
it appears that if mineral blocks are to be of value in bighorn dietary
supplements, the formula must be a compromise between salt content and
mineral content.
In the current study, when the sheep were offered a choice
between a block containing approximately 20 percent salt and a block containing approximately 50 percent salt, the block with the lower salt content
was not used at all even though it was situated next to a block which was
used.
The Carey Range-Min Block appears to offer the best compromise between acceptability and mineral content.
Bighorns will use this block if
it is the only one provided for them.

�Table 1.

Weights of mineral blocks placed on bighorn sheep winter ranges winter, 1971-72.
Initial
Weight,
(Pounds)
l2/13/7l

Weight
(Pounds)
1/29/72

Weight
(Pounds)
2/19/72

Weight
(Pounds)
3/25/72

North Control

50.25

50.00

50.00

50.00

South Control

50.00

50.00

50.00

50.00

Ute Creek West

50.00

49.00

47.50

Ute Creek East

49.50

49.00

47.50

46.75 )
)___ Used by approx. 12 sheep, no deer.
46.50 )

Hay Creek West

49.00

49.00

47.50

Hay Creek East
Hay Creek East 1/

49.50
50.00

48.00
50.00

44.50
50.00

47.25 )
)___ Used by approx. 20 sheep, no deer.
Lf3.00 )
50.00
Never used.

Quartz Mine

50.25

50.25

50.00

49.75

Never used.

Spruce Grove Upper

49.75

49.75

49.75

49.75

Never used.

Spruce Grove Lower

50.00

50.00

50.00

49.75

Never used.

Tarryall Townsite

49.75

49.75

48.75

48.75

Used some by deer, never by sheep

1/21/72

2/26/72

3/8/72

3/25/72

Control

50.00

49.75

49.75

49.75

St. George Upper

49.75

49.50

49.50

49.50

Never used.

St. George Lower

50.00

48.75

46.00

44.50

Used by approx. 8 sheep, no deer

St. George Lower 1/

50.00

50.00

50.00

50.00

Never used.

Block Location

Comments

Tarryal1 Range:

Georgetown:

1/
- Moorman Special Range Mineral Block.

All other blocks are Carey Range-Min Blocks.

I

t-'
t-'

1.0
I

�-120-

Blocks placed on open rocky ridge-tops, and on open flat areas close
to rocky escape cover, receive more use than blocks placed in depressions, brushy areas, or open flat areas away from escape cover.
This simply corroborates what is known to be generally true regarding
bighorn habitat preferences.
In selecting locations for placing
mineral blocks, it seems more logical to choose those sites to which
sheep will naturally move, rather than attempting to use the blocks
as attractants to less desirable locations, assuming that the primary
purpose is to provide mineral supplements.
If, in certain instances,
it is desirable to use blocks for their attractant value (e.g., keeping
sheep away from heavily-traveled
highways) it would be better to use
the plain white block than any mineral block.
Finally, the observation that blocks placed on open rocky ridge-tops
are not subject to deterioration from accumulated moisture is in order.
In the current study, the control blocks were placed where drainage was
perfect.
No weathering or sloughing was evident during the entire
winter period.
In contrast, the block placed at the Quartz Mine in the
Tarryall Range was situated where some moisture could collect.
Even
though never used, it lost a half-pound in weight during a three~onth
period.

Wildlife

Researcher

�January,

-121. I

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-4l-R-22

Work Plan No.
Job Title

1

&amp; Mountain Goat Investigations

Sheep

Job No.
Collection

Period Covered:
Personnel:

Bighorn

of Bighorn Lambs

June 1, 1971 to September

Robert L. Schmidt,

20

30, 1971

Gene G. Schoonveld

and William

H. Rutherford

ABSTRACT

Bighorn sheep lambs were collected from four separate areas in Colorado
during 1971,and were turned over to the Department of Pathology, College
of Veterinary Medicine, Colorado State University, for necropsy examination.
Three' lambs and one yearling were collected on Buffalo Peaks, seven lambs
were collected on Trickle Mountain, three lambs and one yearling were
collected in the Sangre de Cristo Range and five lambs were collected on
Pikes Peak. In addition, one lamb was found dead in the Sangre de Cristo
Range.

1973

��-123-

COLLECTION
William

OF BIGHORN LAMBS
H. Rutherford

P. S. OBJECTIVE
To collect bighorn lambs as specimens
to be done on outside contract basis.

for necropsy

study;

such study

SEGMENT OBJECTIVE
1.

Collect bighorn lambs at time intervals prescribed by contractual
agreement from herds having a history of high lamb mortality, and
from a productive herd to serve as a control.

METHODS

AND MATERIALS

Bighorn lambs were collected by hunting them on foot on Buffalo Peaks,
Pikes Peak and Trickle Mountain, and from a helicopter in the Sangre
de Cristo Range.
Collection attempts using the Cap-Chur gun were unsuccessful.
All lambs collected were taken with conventional firearms.

DESCRIPTION

OF AREAS

Bighorn lambs were collected on Buffalo Peaks, lying on the west side
of South Park between Fairplay and Buena Vista; on Pikes Peak, lying
just west of Colorado Springs; on Trickle Mountain, lying in the northwest part of the San Luis Valley between Saguache and Cochetopa Pass;
and in the Sangre de Cristo Range, lying between the Wet Mountain
Valley and the San Luis Valley in south-central Colorado.

RESULTS
Bighorn sheep collected during this segment are listed in Table 1,
following.
All sheep collected were turned over to the Department of
Pathology, College of Veterinary Medicine, Colorado State University,
for necropsy study by contractual agreement.
The results of this study
will be reported by the contractor as a part of the contract obligation.

Prepared

by ~~~~~~~~~~~~
William H. Rutherford
Wildlife Researcher

~

�-124-

Table 1.

Bighorn sheep collected for necropsy study, 1971.

Area

Buffalo Peaks

Trickle Mountain

Sangre de Cristo

Pikes Peak

1/ Found dead.

Date

Sex

Age

Weight (kg.)

6/16

F

Lamb

12.5

6/29

F

Lamb

9.0

8/10

F

Yr1g.

38.6

8/23

H

Lamb

28.2

6/17

F

Lamb

4.8

6/30

F

Lamb

11.5

7/27

F

Lamb

20.0

7/28

M

Lamb

23.0

8/19

F

Lamb

27.3

9/3

F

Lamb

19.1

9/3

M

Lamb

26.0

8/26

M

Lamb

12.71/

8/28

M

Lamb

8/30

F

Yr1g.

49.0

9/2

M

Lamb

14.6

9/2

M

Lamb

18.2

9/13

F

Lamb

21.0

9/13

F

Lamb

19.0

9/16

M

Lamb

17.3

9/16

M

Lamb

14.0

·9/16

M

Lamb

18.2

�-125-

January,

JOB PROGRESS

State

of

Work

W-4l-R-22

Plan No.

1

Job Title
Period

REPORT

COLORADO
--------~~~~---------No.

Project

1973

J~
Spontaneous

Covered:

Personnel:

Bighorn

July

Sheep

Goat

Investigations

CONTRACT

~.

Diseases

&amp; Mountain

in Bighorn

Sheep

1, 1971 to June 30, 1972

Game, Fish and Parks:
H. Rutherford, Robert

Gene Schoonveld,
E. Keiss.

Robert

L. Schmidt,

William

Colorado State University:
J. L. Adcock and C.P. Hibler,
Co-Principal Investigators;
G. C. Solomon, Pathology;
J. B. Parks, Virology; Harold Breen, Bacteriology;
L. J.
Rich, Hematology; R. E. Pierson, Clinical Pathology.

ABSTRACT

Twenty-two bighorn sheep (20 lambs and 2 yearlings) were presented to the
Department of Pathology, Colorado State University for post-mortem examination and collection of tissues, fluids and contents. Weights of lambs (9F
and 11M) ranged from 4.8 kg. to 28.2 kg.
The two female yearlings weighed
38.6 kg. (#7) and 49 kg.
(#11). Animals collected from the Buffalo Peaks
and Trickle Mountain herds were in good to excellent physical conditioD.
No
severe lung lesions were observed until August 26, 1971.
Ten of the eleven
animals from the Pikes Peak and the Sangre de Cristo herds were diagnosed
grossly as pneumonia of varying degrees as well as lungworm lesions.
The
female yearling (#11) from the Sangre de Cristo herd was in excellent
condition although affected with lungworms.
The Histopathological
examination revealed few lesions in tissues except for the lungs.
The bacteriological results suggested no consistent pattern especially for pathogenic
bacteria.
The hematological
results were reasonably normal in hemoglobin,
platelet, total protein and mean corpuscular hemoglobin concentration
values.
A wide variation existed in the blood cell counts and these should not be
used for interpretation
at this time.
The virological results were revealing
in that positive antibody titers against PI3 virus was demonstrated.
The
parasitological
examination revealed that all animals had infections with
Protostrongylus
stilesi.
Some had light infections, many were severe.
A
few sheep had light infections with gastrointestinal
nematodes and coccidia.
Transplacental
transmission of lungworm was observed in a ewe from the Pikes
Peak herd.
Snails have been established in the laboratory, infected with
Protostrongylus
larvae, and experimental
infections are currently in progress.
Treatment of sheep with tramisol furnished inconsistent results.

��-127-

SPONTANEOUS

DISEASES

IN BIGHORN SHEEP

P. S. OBJECTIVE
To determine causes, nature and effects of spontaneous diseases in
Colorado bighorn sheep with particular reference to mortality in lambs,
and study efficiency of chemotherapeutic agents on Protostrongylus in
bighorn sheep.

SEGMENT OBJECTIVE

#1

Obtain bighorn sheep from the commission in such numbers
locations as indicated in procedures below.

and from such

Twenty-two Rocky Mountain bighorn sheep were collected for studies on
lamb mortality and necropsied by the personnel in the College of Veterinary Medicine and Biomedical Sciences.
A summary of these collections,
including dates, sources of the animals, and gross necropsy findings are
summarized in Table 1. Details are included in an appendix to this report.

SEGMENT OBJECTIVE
Conduct

chemical

and necropsy

examinations

#2
of bighorn

sheep.

PROCEDURES
1.

Live animals will be delivered to Colorado State University where they
will undergo thorough clinical evaluation by veterinary personnel.
Based upon the clinical findings the decision will be made whether to
maintain the animal for further evaluation or subject it to postmortem
examination.

2.

Necropsy examinations of animals killed or dead when submitted will be
conducted by a veterinary pathologist and will include gross and histologic evaluation of all organs and tissues as well as collection of
appropriate specimens, as indicated, for nutritional, parasitologic,
bacteriologic, virologic and toxicologic studies.

�-128-

Table 1.

Summary of necropsy findings on 22 bighorn sheep.
Date

Wt. (Kg.)

Sex

Area 1/

1

6-16

12.5

F

B.P.

2

6-17

4.8

F

T .M.

NVL

3

6-29

9.0

F

B.P.

NVL

4

6-30

11.5

F

T.M.

5

7-27

20

F

B.P.

LWL

6

7-28

23

M

T .M.

NVL

7

8-10

38.6

F

B.P.

NVL yr1g.

8

8-19

27.3

F

T.M.

LWL

9

8-23

28.2

M

B.P.

LWL

10

8-26

12.7

M

S.C.

LWL+P

11

8-30

49

F

S.C.

LWL yrlg.

12

8-28

M

S.C.

LWL+P

13

9-2

14.6

M

S.C.

LWL+P

14

9-2

18.2

M

S.C.

LWL+P

15

9-3

19.1

F

T .M.

NVL

16

9-3

26

M

T .N.

LWL

17

9-13

21

F

P.P.

LWL+P

18

9-13

19

F

P.P.

LWL+P

19

9-16

17.3

M

P.P.

LWL+P

20

9-16

14

M

P.P.

LWL+P

21

9-16

18.2

M

P.P.

U1L+P

22

9-16

27.2

F

RMPT.

LWL+P

Gross Lesions

II

- B.P.=Buffalo Peaks; T.M.=Trick1e Mt.; S.C.=Sangre de Cristo; P.P.=Pikes
Peak; RMPT.=Rampart Range.
l/NVL=No visible lesions except for gunshot wound and attendant pathology.
LWL=Lungworm lesions. LWL+P=Lungworm lesions + pneumonia.

�-129-

RESULTS
The only live animal delivered to the University was a 2 or 3 day old
bighorn male lamb. This animal was captured at Pikes Peak on May 15,
1972. A complete clinical, bacteriological, parasitological and virological examination was conducted on this animal.
Questionable results
on the virological tests of nasal washings and the blood suggests that
add i t Lona L tests are needed to clarify the initial results.
Lungworms
would not be detected before mid-July, 1972. If the animal is normal
and' since it is needed in Meeker, Colorado in the semi-tame herd, it
will be raised in captivity and eventually released in the pasture with
the other animals.
In the meantime, regular examinations will be conducted on this animal to obtain as much baseline data as possible.

NECROPSY

EXAMINATION

The necropsy examinations revealed that the respiratory system was the
one most commonly affected.
No grossly visible lesions were obtained in
four lambs (#1, 2, 3, 4) and one yearling (#7). For a brief summary, see
-Table 1.
The lesions

in the respiratory

(1) No visible

lesions

(2) Mild to moderate

(3) Marked

(WL)

system varied:
- 5 animals - 4 lambs (ffl, 2, 3, 4)
- 1 yearling (#7)

lesions - 7 animals - 6 lambs (#5, 6, 8, 9, 15, 16)
- 1 yearling (#11)

to severe lesions - 10 animals

Mild to moderate

lesions were characterized

by:

(1) Presence of lungworm nodules primarily
surface of the diaphragmatic lobes.
(2) Few to no microabscesses
(3) Few to no yellowish
Marked

on the posterio-dorsal

in lung parenchyma.

to cream colored foci.

to severe lesions wer e characterized

(1) Fibrino-purulent

(#10, 12, 13, 14, 17, 18, 19,
20, 21, 22)

by:

exudate in the pleural

cavity.

(2) Adhesions between lobes of lungs, between lobes and costal pleura,
between lobes and the diaphragm and between pericardium and pleura.
(3) Consolidation of lung tissue with a meaty texture and plum coloration especially in the ventral 1/2 to 1/3 of the lobes of the lung.

�-130-

(4) Purulent exudate expressed
parenchyma.
(5) Enlarged

from the cut surface of lung

and edematous bronchial

(6) Granular appearance
development.
Summary of individual

and mediastinal

lymph nodes.

of many lobes due to lymphofollicular

necropsy

examinations:

#1

Female lamb approximately 21 days old. 12.5 kg. Buffalo Peaks.
Excellent condition.
No grossly visible lesions observed.

#2

Female lamb. Youngest lamb collected.
4.8 kg. Trickle Mountain.
Excellent condition.
No grossly visible lesions.

#3

Female lamb. 9.0 kg. Buffalo
grossly visible lesions.

#4

Female lamb. 11.5 kg. Trickle Mountain.
grossly visible lesions.

#5

Female lamb.
20 kg. Buffalo Peaks. Good condition.
Few nodules
observed on the posterio-dorsal surface of the right diaphragmatic
lobe.

#6

Male lamb. 23 kg. Trickle Mountain.
nodules present on the posterio-dorsal
lobes.

#7

Female yearling.
38.6 kg.
grossly visible lesions.

#8

Female lamb. 27.3 kg. Trickle Mountain.
Excellent condition.
Several yellowish, circumscribed nodules (2-3 mm) on the posteriodorsal surface of both diaphragmatic lobes. Edema in bronchial
lymph nodes.

#9

Male lamb. 28.2 kg. Buffalo Peaks.
Good condition.
Several
elevated nodules (1 to 4.5 mm) on the posterio-dorsal surface of
the diaphragmatic lobes. Whipworms present in cecum. Mesenteric
lymph nodes moderately enlarged.

#10

Male lamb. 12.7 kg. Sangre de Cristo.
Very poor condition.
Rough hair coat. Enophthalmus.
Fibrino-purulent
exudate in pleural
cavity.
Adhesions between ventral one-half of lobes and costal
pleura.
Mediastinal and hilar lymph nodes enlarged.
Ventral portions of all lobes consolidated with a meaty plum colored texture.
Numerous nodules posterio-dorsal surface of diaphragmatic lobes
and intermediate lobe. Microabscesses present.
Adhesions between

Peaks.

Excellent

condition.

Excellent

No

condition.

No

Excellent condition.
Few
surface of the diaphragmatic

Buffalo Peaks.

Excellent

condition.

No

�-131-

diaphragm and diaphragmatic lobes; also between pericardium
and pleura. Mesenteric nodes enlarged.
Small thymus. Peyer's
patches of ilium dark red. Fluid in tarsal joints.
Purulent
exudate in frontal sinuses.

#11

Female yearling.
49 kg. Sangre de Cristo.
Excellent physical
condition.
Post mort5n autolysis very marked throughout.
Nodules present on posterio-dorsal surface of diaphragmatic lobes.

#12

Male lamb. No weight.
Sangre de Cristo.
Poor condition.
Adhesions of right diaphragmatic and cardiac lobes to costal
pleura. Adhesions of cardiac to apical lobe. Purulent exudate
expressed from cut surface.
Hilar lymph nodes edematous.

#13

Male lamb. 14.6 kg. Sangre de Cristo.
Poor condition.
Adhesions
between lobes of lung and costal pleura. Ventral 1/2 of right
diaphragmatic intermediate, and cardiac lobes consolidated and
meaty in texture. A large abscess was present in right apical lobe.

#14

Male lamb. 18.2 kg. Sangre de Cristo. Poor condition. Rough
hair coat. Thymus very reduced in size. Hilar lymph nodes
hemorrhagic and edematous.
Adhesions of liver to diaphragm
(1-2 rom) consisted of a capsule and reddish colored fluid within the capsule (a cyst). Adhesions between apical lobes and
costal pleura. Ventral ~/3 or all lobes we re consolidated, meaty
and plum colored in texture.
Purulent exudate was expressed from
the cut surface of the lung tissue.
Small granular wh Lte foci
(0.5 mm to 1 mm) were present t'':-rroughout
the consolidated areas.
Posterio-dorsal surface of diaphragmatic lobes contained several
circumscribed and organized yellowish foci.

fi15

Female lamb. 19.1 kg. Trickle Mountain.
Excellent condition.
Large thymus. Few elevated nodules on the posterio-dorsal surface
of diaphragmatic lobes.

#16

Male lamb. 26 kg. Trickle Mountain.
Good cond it Lon , Few elevated nodules on pos t.er i o+doz sa.L sur f ace of di2pl'1rag~tlatic
lobes.
Two microabscesses in dLaph r agrnat Lc lobe.

#17

Female lamb. 21 kg.
Pikes Peak. Fair condition.
Fibrin tags
and adhesions from borders and edges of all Lob es to the costal
and diaphragmatic pleura. Ventral 1/3 of all lobes were consolidated ~ meaty in texture and p Lum colored.
Purulent exudate was
expressed from cut surfaces.
Hilar and mediastinal lymph nodes
were enlarged and edematous.
Several elevated nodules were present on the posterio-dorsal d i.aphr agmat.Lc lobes. Hucoid sinusitis
of frontal sinuses.
Kidneys 'were pale, mottled, and contained
dark pigment.

I·

�-132-

#18

Female lamb. 19 kg. Pikes Peak. Fair condition.
Fibrin tags
and adhesions between lobes and costal pleura. Ventral 1/3 of
all lobes were consolidated, meaty in texture, and plum colored.
Purulent exudate expressed from the surface.
Hilar and mediastinal lymph nodes were enlarged and edematous.
Elevated
nodules were present on the posterio-dorsal surface of the
diaphragmatic lobes. Mucoid sinusitis of frontal sinuses.
Kidneys were mottled and pigmented.
Milk curd in abomasum
and rumen was full.

#19

Male lamb. 17.3 kg. Pikes Peak. Good condition.
Rough hair
coat. Curds of milk in abomasum.
Pebbles in rumen which was
full. Palatine tonsil enlarged.
Hilar lymph nodes edematous.
Elevated nodules were present on posterio-dorsal surface of
diaphragmatic lobes. Ventral 1/3 of right cardiac lobe and
right apical lobe consolidated with a meaty texture and plum
coloration.
Lymphofollicular development marked.
Opacity
of left lens.

#20

Male lamb. 14 kg. Pikes Peak. Very poor condition.
Milk
curd in abomasum.
Pebbles and vegetable matter in rumen. All
lobes of both lungs consolidated, meaty in texture, and plum
colored.
Numerous granular greyish-white foci (lymphofollicular
nodules) present throughout the consolidated areas. Numerous
elevated nodules and microabscesses were present on the posteriodorsal surface of the diaphragmatic lobes. Hilar lymph nodes
were enlarged and edematous.
Purulent exudate was expressed
from cut surfaces of lung tissue.

#21

Hale lamb. 18.2 kg. Pikes Peak. Very poor condition.
Chewing
lice present around penis and testicles.
Ventral 1/2 of all
lobes grey in appearance.
Lobules demarcated by these granular
areas. Adhesions between lobes of the lung and costal pleura.
Purulent exudate expressed from cut surface.
Several elevated
nodules were present on the posterio-dorsal surface of diaphragmatic lobes (4-5 rom). Mediastinal, tracheal, and hilar lymph
nodes enlarged.
Healing fractures of ribs #8 and 11. Pleura
thickened.
Mesenteric lymph node enlarged.

#22

Female lamb. 27.2 kg. Rampart Range.
Good condition. This animal
was observed the day before it died walking very slowly and in
a lateral motion.
It was found dead the following day. The hair
coat was smooth.
Autolysis was extensive.
Blood colored fluid
present in thoracic cavity.
Few nodules on posterio-dorsal
surface of diaphragmatic lobes. Ventral 1/3 of all lobes were
dense, meaty, and plum colored.
Purulent material could be
expressed from the cut surface.
Abscesses were present in the
right apical and intermediate lobes.

�-133-

Summary and interpretation of gross findings:
Weights of lambs ranged
from 4.8 kg. to 28.2 kg. The yearlings weighed 38.6 kg. (#7) and 49
kg. (#11). The animals collected from the Buffalo Peaks and Trickle
Mountain herds were in good to excellent physical condition including
the two collected from Trickle Mountain September 3, 1971. The two
yearlings (#7 from Buffalo Peaks and #11 from Sangre de Cristo) were
in excellent condition.
The four lambs from the Sangre de Cristo
range were in poor condition while only two of the six lambs (#20 and
#21) from the Pikes Peak and Rampart Range were in poor condition.
Milk curds were present in the abomasum of many lambs including #20
which was collected from the Pikes Peak herd September 16, 1971. The
rumen of each, except #2 which was still on a milk diet, was full of
plant material.
Hair-balls in the abomasum and small pebbles in the
rumen were observed in several lambs.
No severe lung lesions were observed until August 26, 1971 at which
time a male lamb (#10) was found dead in the Sangre de Cristo area.
Of the eleven animals collected from the Sangre de Cristo (5) and
Pikes Peak or Rampart Range (6) herds each of the lungs contained
nodules in the posterio-dorsal area of the diaphragmatic lobes.
In
addition, all animals except the yearling from the Sangre de Cristo
area (#11) had a severe pneumonia in the pendant (lower) portions of
the lung. This varied from the ventral portion of a single lobe being
affected to a large percentage of all lobes affected.
The gross
characterization of the affected lobes was dense, meaty and plum
colored, accompanied by numerous foci of small granular greyish-white
areas believed to be lymphoid follicles.
Purulent exudate could be
expressed from the cut surface of the lungs from several animals.
In
those from which no exudate could be expressed the changes were
suggestive of a "viral" pneumonia.

VIROLOGICAL
Virus isolation attempts:
Thus far, thirty different tissues from six
animals (71 BHL - 13, l4~ 15. 19, 20, 21) were inoculated onto bovine
embryonic kidney. bovine emb ryorrLc Lung and lamb kidney tissue cultures.
Three blind passages were made from all Lno cu Iums . All results were
negative.
Serology:
anti PI3

71 BHL 5
7
17
18
19
20

1:4
1:8
1:4
1:64
1:16
1:8

72 BHL 1
2

0
0

anti IBR

0
0
0
0
0
0

anti Bluetongue

0
0
0
0
0
0

�-134-

Tissue Culture Initiation--A bighorn lamb foetus was collected aseptically
and primary tissue cultures were started from heart, lung, adrenal,
thyroid, spleen and kidney.
All tissues are presently in their fifth
passage and still growing well.
Early passages of each tissue have been
frozen and should supply cells for at least one year.
Most primary cells show spontaneous degeneration at about the third
passage.
These cells have already been passed long enough to classify as
cell strains and there may be a possibility of establishing a cell line
(infinite subpassage capacity) from one or more cell types.
All of the bighorn lamb cell cultures except heart are very susceptible
to the strain of PI3 isolated from the Wyoming bighorn herd. The virus
exhibits a more marked cytopathic effect and grows to a higher titer in
the bighorn lamb tissue than it does in the domestic lamb lung which was
used for the initial isolation.
The bighorn lamb tissue cultures may be an excellent
isolation attempts.

substrate

for future

Significance of Findings--The negative results encountered in viral isolation attempts from improperly collected field tissue and the positive
isolations from usable field tissue stresses the importance of proper
procedures for collection of material and the uselessness of indiscriminate
wholesale sampling, without correct equipment.
Serological studies indicate a widespread (85%) incidence of exposure to
PI3. The absence of antibody titers to IBR and bluetongue may mean that
these diseases do not occur in bighorn sheep in this area, but the small
number of serum samples must be taken into consideration.
A large number
of serum samples might be more indicative.
The isolation of PI3 virus from the sheep at Sybille substantiates earlier
indirect (serologic) evidence that PI3 viral infections occur in this
species.
At the time of their capture, 4 sheep had HI titers to PI3 virus,
which indicates that the virus was not acquired from adjacent animals while
in captivity.
After 1 month in captivity, all sheep had HI titers to PI3
virus.
The close confinement might have precipitated the spread of PI3
virus from sheep that were infected in their natural habitat to the remainder of the flock while in captivity and the PI3 virus, isolated from
3 of the sheep, could have contributed to the deaths of all 10 sheep.
The isolation of PI3 virus indicates that bighorn sheep are definitely
capable of supporting infection with this agent. Parainfluenza-3 virus
has been isolated from domestic sheep and has been shown to cause experimental virus pneumonia in domestic lambs.
The next area of investigation
should be the role that PI3 virus plays in the pneumonia complex of bighorn sheep.

�-135-

The initiation of tissue culture cells from the bighorn lamb fetus may
provide a substrate that is mere Se~sitive ror the isolation of additional viral agents.
It also provides an in vitro method for testing
the susceptibility or bighorn sheep to various viral agents.
If bighorn cells in tissue culture are susceptible to a virus, it is likely
that the animal will be susceptible also.

BACTERrOLOG reAL

Procedure:

Swabs and/or tissue were inoculated

(a)

thioglycollate

(b)

agar plate media

into (onto);

broth

(1) B+ (Blood agar)
(2) EMB (eosin methylene blue)
(3) BG (brilliant green)
(4) NaAz (sodium azide - crystal violet)
After inoculation, the tubes I/leTeincubated aeriobically at; 37°C and plates
were incubated at 370C under aerobic, CO2 and anaerobic environments
(complete replicate sets of inoculated media for each of the environments).
Readings

and re-inoculations

were carried on as follows:

(a)

Thioglycollate broth tubes - after 24 hours incubation, were
re-inoculated onto the respective plate media and incubated
under the three different enviro~ments for 24 more hours and
read.

(b)

The direct inoculations onto the plate media (all 3 environments)
were incubated at 37°C for 48 hours, read, reincubated for 24
hours and read again.

(c)

Reading consisted of observing gross morphology, pigment, hemo Iy t Lc
pattern and other distinctive, recognizable. identifying colony
characteristics and gram staining.

(d)

Where indicated for purposes of typing (or process of elimination)
recognized biochemical tests were run (i.e. iron milk, sugars,
etc.) and/or serological tests using Salmonella polyvalent 0 anti
sera.

�-136-

Interpretations of Findings:
Of all the specimens sampled, assuming that
all external contaminants were eliminated, the samples where various
bacterial growth would be expected are: abomasum, eyes, intestine and
nasal cavity; the other tissues in healthy, disease-free animals routinely
reveal no growth unless there is a localized bacterial infection or a
systemic bacteremia, or depending on interval between death and necropsy,
agonal invasion and spread.
The most consistent and most widespread bacteria were Staphylococcus (nonhemolytic) and Alpha Hemolytic Streptococcus
(Table 2). Ordinarily, unless
associated with suggestive tissue changes, these organisms would be considered
questionable pathogenic significance.
The Bacil~us species (practically
all~. subtilis) are common environmental inhabitants.
The finding of
Clostridium perfringens and!. coli in the gastrointestinal tract is considered normal in many or most ruminants.
The V = unidentLfied coliform
mostly was classed as a non-lactose ferment or that was negative on the
Salmonella polyvalent 0 anti serum test; its significance is not evaluated,
but noted that it appeared in 11 of the composite summaries (samples)
(!. coli appeared in 10); 9 of these composite summaries (samples) had both
!. coli and the unidentified coliform, while!.
coli appeared one time without it, and the unidentified coliform appeared twice without !. coli.
Of the other organisms found, a diphtheroid (Corynebacterium) was found once,
Proteus once, Pseudomonas three times, Gamma Streptococcus once. Only the
Pseudomonas and/or Proteus would ordinarily be incriminated, if lesions or
infection (i.e. cystitis or nephritis) were present.
As is commonly found in domestic animal (bacterial) disease study, the
presence of various bacteria at varying levels are not considered significant
primary etiologic agents.
However, should there be other initiating causes
of localized tissue or systemic pathology, many bacteria can and do act as
secondary invaders and are associated with various lesions and disease
entities.

HEMATOLOGY
Hematologic
Table 3.

data obtained

from the lambs collected

in 1971 are presented

in

A total of eight complete hematological examinations were made.
Six partial
examinations were made and no samples were collected from seven animals.
From one animal (#7), only serum was available (Table HI). All serum samples
were given to the virologist and the results are included in his report.
Hematologic values determined from the field collections are difficult to
assess. All except two (#7 and #11) are lambs and it is difficult to determine the true variation between lambs of different ages and sex and the
difference between lambs and adults.
It appears that the hemoglobin value
falls within or close to the range of values determined by Franzmann (1971)
on captive and wild adult bighorn sheep except for animal #4 with a value
of 9.3.

of

�Table

2.

Summary of

B= Bacillus

Bacteriological

s p . ; C= Clost.

p= Pseudomonas

sp.;

findings.

:=..= Corynebacterium

perfr.;

S= non

hem.

staph.;

~=

alpha

s p . ; E=J;.

strep;

N"" no growth;

colLi

G&gt;=~mma

strep.;

@=protein;

U= undetermined

BHL if
1

Specimen
Abomasum

Blood
Brain

Cerv,

2

3

4

I

I
l

6

C
E
S

B
C
E

7

8

9

10 11 12 13 14 15 16 17 18 19 20 21

E
s

C

.

1\
S

B

E

s
B

Co~oosite

I

B

C

N

s

B

N

Summar
E

S

s

S

s

U

s

N

i

&amp; CNS

5

s

s

s

s

N

I

s

TI

N
S

I

l·~

I

,
I

N

I

~

w

Eye(s)

I N

t:

S

Intest.

Joints

S
S

I

,
I

1
S
s
B
C
E

I\'

E

F

N

B
E

E

B

s

s

B

S
s

E

B
C
E

s

s

1;

1\

C

s

C

B

s

S

'IT
U

S

1\'

IT 'IT

u
s

E
U

U

N

N

s

S
Ki.dney

i

N

K

N

(j?)

S

S

S
s

s
Liver

,
I

N

N

N

N

S
S

S

N

s
U

B
E

B

B

S

U

E
P

i

B

E

N

S

S

I B

C

E

S

s

U

E

N

S

s

U

B

E

New

@l
U

I

N ~

I

P
U
S

I

B

E

N

P

U

S U
s

S

TI
,
-----------------------------------------------------------------------------_._----------------------

o@u

"
I

�Table

2.

Summary

B= Bacillus

sp.;

p= Pseudomonas

of

Bacteriological

C== Clost.
sp.;

findings

S:..~.= Corynebacterium

perfr.;

S== non

hem.

(continued).

staph.;

~== alpha
BHL

Soecimen

,

1

3

4

5

6

7

8

9

E.o::J;. _coIL;

N" no

G&gt; "" gCimma s trep.;

strep;

N

S
s

Lymphnodes

S

B
N

N

cavity

@==protein;

U== unde termined

-if

10 11 12 13 l!t 1516

17 18 19 20 21

E
N

B
N

S
s

-

G) ~
B
s

s

U

B
S

B B B s
E EP Ep U
-sU SsU Ss

S

S
U

Hesentery
Nasal

growth;

Co~posite

E
S

~

E
S
s

B
E
s

B
P
s

s

E

-

pancreas

N

s
U

s
-

U

U

IB

E

N

P

S

S
U

IB

N

S

s

U

E

P

S

N

N

B
s
U

B

00
I

N

Peritoneum

N

N

B

B

N

N

N

U

S

S
s

S
s

B
E
s

S
s

S
s

s

- -

Urine

I s

I

N

E
s

s

S
s

c

N
N

B
s

U
I

N

N

-s

w

N

N

U

N

N

Spleen

s

!-'

N

Pericardium

Trachea

S1.llr'J:\arv

i

I Ss

Lung

2

sp.;

S
s

-

s
U

S
s
U

B

E

N

S

s

U

B
s
U

IB

c

E

S

s

U

IN

�/

Table 3.

Hb

Hematological

data obtained from bighorn sheep lambs examined in 1971.

Platelets

Total Protein

PCV

(MCRC) )~
%

53
6.3
OK
18
28
8.0
OK
10.1
40
5.3
OK
13.5
27
4.6
OK
9.3
43
5.0
OK
5 15.2
43
4.7
OK
6 15.2
7 only serum available
6.3
41
8 15.0
6.3
41
9 15.0
10 no blood sample (lamb was found dead)
11 no blood sample
7.6
46
12 16
5.5
53
13 19.4
14~8
6.1
38
14
13.2
5.5
32
15
no
blood
sample
16
17
18 no blood sample
19 clotted - no determination
20
21 clotted - no determination
22 no blood sample
1
2
3
4

Hb
1

RMBR

l8.1±3

29
3. 7

29
29
29
29

WBC
Segs
(103)
4.1
30
7.6
3.2
5.4

23
23

27
27

25

84
35
40

Lymphs

16
70
61
60

7.6

42
39

2.0
2.5

too few cells
40

58

61

39

2.7

28.5

Bands

16.9
3.2
0.87

Total Protein

PCV

+
6.5~L5

+
45.2-6.6

Eosinos

1

3

57

56

71
23
cells disintegrated
cells disintegrated
too fe,v cells
71

Monos

23

2

4

2

\0
I

6

WBC

6.4-11.7
54-56
3.1±10.6
18-22.
2 Stone Sheep
1, Franzmann~ Albert W. 1971. Comparative Physiologic Values in Captive and Wild Bighorn Sheep,
;!.. \-lilctUfeDis., Vol. 7:105-108.
2. Franzmann, Albert W. 1971. Physiologic Values of Stone Sheep, ;!.. Wildlife Di.s , , Vol. 7 :139-141.
i'HCRC :::.
mean corpuscular hemoglobin concentration, which is the relationship between Hb in mean
corpuscular volume in cubic microns (%). Reduced in microcytic anemias, iron deficiency, and
chronic blood loss.

I

t-'

W

�-140-

Total protein values are rather consistent and agree generally with values
determined by Franzmann (1971).· The packed cell volume (PCV) except for
animals 1/2, 1/4, and 1/15 appear to be acceptable.
The mean corpuscular
hemoglobin concentration (MCRC) is consistent and suggests that there is
not microcytic anemia, iron deficiency, or chronic blood loss.
The complete blood count (CBC) is the most inconsistent and difficult to
evaluate.
The white blood cell count (WBC) varies so much that additional
samples under more ideal conditions should be collected.
The very low
white blood cell counts in animals 1/8, 9, 12 and 15 mayor may not be
significant.
The high WEC for animal 1/13 is not reliable since many
cells were disintegrated prior to counting.
The variation in the numhers
of neutrophiles and lymphocytes and the ratio between them is too great to
evaluate accurately.

SEGMENT OBJECTIVE

1/3

Survey bighorn sheep for the incidence of infection and level of infection
with ectoparasites and endoparasites and evaluate the significance of these
parasites on the health and well-being of the animal.

PROCEDURE
After the pathologist had examined the lambs, and obtained all the tissues,
cultures, etc., they were then examined for ectoparasites and endoparasites.
Lungs were opened and the air passages examined for Protostrongylus rushi;
the parenchyma was examined for Protostrongylus stilesi.
Following this
examination, the lungs were finely chopped and placed in Baermann funnels
for 24 hours to recover first stage larvae of Protostronglyus.
The abomasu~,
small intestine and large intestines were opened, the contents flushed
.
through screens, and examined for nematodes and tapeworms.
In addition,
fecal flotations were done on all animals to determine presence or absence
of parasites.

RESULTS
The results of this examination

are given in Table 4.

None of the 23 lambs examined were infected with Protostrongylus rushi.
All were infected with Protostrongylus stilesi.
For an interpretation of
the severity of infection, read pathology section (Segment Objective 1/2),
and compare.
Animals numbered 1, 7, 10, 12, 17, 19, 20, 21, 22 and 23 all
possessed first stage larvae of Protostrongylus in numbers sufficient to be
considered detrimental to the host. These numbers of larvae are indicative
of a heavy infection with adult parasites.
Animals 17 through 20 and 22 had extremely low infections with a species of
Nematodirus not yet identified, and animals 7,11, 16 and 22 each had 2-7

�I ~

-141-

Marshallagia marshalli.
The numbers of both parasites were insignificant.
Bighorn sheep #22 had a very low infection with Trichuris ovis. No other
nematodes or tapeworms were found in these animals.
All of the species of coccidia found in the lambs were very few in number.
The range was 25 to 200 oocysts per grmn of feces, which consistutes an
insignificant infection.
Only one bighorn sheep lamb was infested with ectoparasites,
animal had a few Linognathus sp., a suckling louse.

and this

�Table 4.

Endoparasites found in lambs collected during 1971.

Sheep
Identification

Protostrongylus
larvae

Nematodirus
sp.

Marshallagia
marshalli

7lBHL - 1
?lBI-IL
- 2
7lBHL - 3
7lBHL - 4

19,937
Patent Lnfec t Lon'
43
300

-

-

E. crandallis
E. arloingi

7lBHL - 5
7lBHL - 6
7lBHL - 7

16,333

-

+

7lBHL - 8
71BHL - 9
71BHL - 10
71BHL - 11

1,699
2,120
151,515
2,748

-

-

E. faurei
E. arloingi
E. faurei

-

+

?lBHL - 12
7lBHL - 13
7lBHL - 14
71BHL - 15
71BHL - 16
71BHL - 17

14,092
2 ,117
5,250
1,596
1,560
9,998

-

-

+

+
-

71BHL - 18

6,333

+

-

71BHL - 19

17,575

+

-

7lBHL - 20

401,750

+

-

7lBHL - 21

337,500

-

-

65,585
23,400

+

+

7lBHL - 22
7lBHL - 23
l"

.

Eimeria
species

E. crandallis
E. arloingi
-

Other

-

~
~
I

-

E. intricata
E. iaurei
E. faarei
E. arloirtgi
E~ arloingi
E. faurei
E. ah-sa-ta
E. arloirtgi
E. arloingi
E. faarei
E. crandallis
E. faurei

First stage larvae too few in number to obtain an accurate count.

Trichuris ovis

�-143-

SEGMENT OBJECTIVE
Elucidate

the biological

114

cycle of Protostrongylus

in Colorado bighorn

sheep.

PROCEDURE
Establish a colony of land snails in the laboratory and expose these snails
to infection with the larvae of Protostrongylus by placing larvae with the
snails.
Once snails are infected, and then infective (about 60 days later)
they will be fed to domestic lmnbs. Domestic lambs with and without immune
suppressant compounds will be used. Bighorn sheep lambs will be infected
as known uninfected lmnbs become available at Little Hills Experiment
Station.

RESULTS

Culturing

Snail Intermediate

Hosts

During the past year, Vallonia pulchalla. a suitable intermediate host for
Protostrongylus, was successfully established in the laboratory.
They are
cultured in four inch diameter glass fil1.gerbowls covered with a glass plate.
The substrate consists of a coarse basal layer of sand covered with a fine
sand surface.
The sand is sterilized in an autoclave and is kept moistened
by adding an eyedropper-full of water every few days.
Condensation should
not be allowed to accumulate within ..
he dish or on the underside of the
glass plate.
Snails are fed a mixture of pulverized oatmeal mixed with
calcium carbonate, ground calf-starter pellets or cottonwood leaves.
Some
mold growth can be expected but this appears to be acceptable to the snails
in moderate amounts and, in fact, is eaten by them. Maintaining cultures in
a cool (700F) fairly dark area is beneficial.
Culture dishes may be changed
every month or so. When cultures are initiated, many eggs will be deposited
by the adult snails.
It is best to transfer these adults to another culture
dish to prevent cannibalism.
Eggs will hatch in 14-21 days. Young snails can be maintained in nursery
cultures until they reach maturity. about 3-4 months.
We have had greater
success in raising snails from eggs than we have had maintaining adults
.
collected in the field. This may be related to pathogens carried by ·fieldcollected snails, changes in feed, or age of snails.

Infecting

Snails With !,ung,.;rorin
La·rvae

Methods of successfully infecting snails in ·[he laboratory
loped this past year.
Excellent success has been obtained

have been deve-'
using two methods:

The simplest method is to place infected pellets in a circle on sand contained
within a culture chamber. He use 12-16 pellets from samples that have about

.' .

�-144-

100-300 larvae per gram of feces.
Pellets are soaked in water for 5-10
seconds before being placed in a circle.
Snails can be activated when
placed in a drop of water.
They are then put onto the pellets.
If snails
are reintroduced onto the pellets every half hour or so for 5-6 hours,
infections of 1-8 larvae per snail will be obtained.
The other method is more suitable for heavier infections.
First stage
larvae are collected from pellets suspended in a Baermann Apparatus.
The
supernatant is removed from the collecting tube and discarded.
The remaining fluid contains the first stage larvae.
This is placed on a glass
plate.
The fluid is then mixed with pulverized oatmeal, or a similar food
and the result is a thin film of food containing first stage larvae.
Snails
are activated and placed on the plate.
They become infected as they feed.
The plate can be kept from drying by placing it upside down over a culture
dish. This technique often produces undesirably heavy infections; consequently, it has not been used often.
Infected snails become less active after they are infected and are much more
subject to changes in culture moisture, humidity, temperature and food.
They must be carefully attended.
Often they become dormant but the larvae
will continue to develop.
This is a critical time and they must not be
allowed to become too dry or too damp. After about three we.eks, black
cuticles, characteristic of third stage infective larvae, begin to appear.
In another two weeks larvae are ready for infection experiments.

Infection

Experiments

A pregnant domestic ewe was fed 150 infective lungworm larvae by trickle
doses over a four week period.
This ewe subsequently gave birth to twin
lambs.
One lamb was sacrificed and the critical tissues (liver, lungs and
lymph nodes) run through the Baermann Apparatus to check to transplacental
transmission of the infective larvae.
The ewe and her remaining lamb are
being checked for evidence of infection.
Feces are being collected biweekly and run through the Baermann Apparatus.
The results of this experiment are not yet known.
Similar infection experiments using bighorn sheep will be initiated at
Little Hills within the next few weeks.
The objective of these infections
will be to investigate the possibility of Protostrongylus spp. as a
triggering mechanism resulting in pneumonia.

Transplacental Transmission of
Protostrongylus ~. in Bighorn Sheep
During the course of investigations into the causes of excessive lamb mortality among bighorn sheep in Colorado, a 9-10 year old pregnant ewe was
collected from the Pikes Peak herd on May 16, 1972. The purpose of the
collection was to obtain foetal tissues for use in tissue culture media,
and to examine tissues for evidence of transplacental or transmammary
transmission of Protostronglyus spp.

�-145-

A complete postmortem examination of this aclimal revealed that she was
in excellent pLysical condition fOT a ewe ::::o·lected
in early spring.
Nodules caused by infection wid'! Protostrongylus stilesi wer-e evident
in the diaphragmatic lobes of both lungs but these were few in number.
No additional changes were detected.
Following postmortem examination, two cotyledons, the mamma.ry gland, the,
mesenteric, hilar and bronchial lymph nodes from the ewe; the mesenteric,
hilar and bronchial lymph nodes f rom the foetus; and a Sill 11 piece of
foetal lung from one d Laph r agma t i.c lobe (approxi-mately 5 x 50 rom) were
Baermannized separately for 24 hours and examil'ceGf or Laxv ae of Protostrongylus ~.
One third stage larva was recovered from the cotyledons, 11 third stage
larvae from the foetal liver, and one from the foetal lung. Except for
size, these 13 third stage larvae are morphologically
identical to the
infective larvae of Protostrongylus found in the snail host. Vallonia
pulchella.
They measure 532 to 606 microns long and 33 to 40 microns wide.
Third stage from snails infected 33 to LW days previously average 505
microns long and 30 microns wide.
The majority of the foetal lung was -USed for the tissue culture medium,
and the remainder of the cotyledons 'were discarded; o t herwf se , a greater
number of third stage larvae -might have been recovered.
Post (personal communication) and a number of other inVestigators have
excellent circumstantial evidence that P:cotostrongylus s p , was transmitted
transplacentally; however, this is the first time that t-hird stage infective
larvae have been isolated from the foetal tissues.
Monson and Post (J. Parasitol., 1972. 58:29-33) have shown that infection
of an animal can occur by ingestion of the third stage larva with the snail.
Presumably, infection of the foetus occurs when the ewe eats infected snails,
releasing the third stage to cross the placenta.

SEGMENT OBjECTIVE

Study the epidemiology

of Protostrongylus

If 5
in biglorn

sheep.

PROC:EDURES
The epizootiology of Protostrongylus in bighorn sheep will be accomplished
by the study of a herd of six semi-t:filr:e
sheep in a 200-acre pasture at Lft t Le
Hills Experiment Station.
These sheep '(,]e:ce
put in a clean pasture to establish a herd for future use. Lungteo rm larval output will be determined at
bi-monthly intervals for each sheep. The date that lambs from these sheep
begin producing larvae will also be determined.
The longevity and dispersal

�-146-

of the larvae from sheep feces will also be determined.
These data will
be obtained by extracting and counting larvae f~om the feces and/or soil
using the Baermann Apparatus.
The species, number and distribution of
snails will be determined, and the level of their infection evaluated
by periodic collection and examination for lungworm larvae.

RESULTS

Monitoring

Program

Several hundred samples have been collected from the small herd of bighorn sheep at Little Hills and analyzed for lungworm larvae. At no time
have parasite loads, as measured by larval output, been high. They have
varied between 0 and 350 larvae per gram of feces during the past year
(Table 5). Seasonal fluctuations occur, with the highest larvae output
occurring in the winter; the lowest in the summer.
Levels of infection
appear to be stable.
An uninfected ram was put into the pasture at
Little Hills a year ago and he is still uninfected.
Last year's lamb
shows less than 20 larvae per gram of feces which is about what she showed
last summer.
These observations indicate that the sheep are not picking
up infected snails from the range within the pasture.

Intermediate

Host Species

The following land snails have been recovered from within the pasture at
Little Hills:
Oreohelix ~.,
Vallonia cyclophorella, Pupilla blandi,
and Vertigo~.
The last three of these snails were reported by Pillmore
to be suitable intermediate hosts.
Only a few of the snails collected
have been alive and none of them have been infected.
All of them, with
the exception of Oreohelix ~.,
are rare in the pasture.
Vallonia cyclophorella and Pupilla blandi are found widely in duff under oakbrush, pinon
pine, and mountain mahogany or in moss that grows under sagebrush which
grows in the bottoms of the canyons.
Vertigo~.
is found on grass roots
and under rocks in moist, shaded areas in the bottoms of the ravines.

�-147-

'Tab1e 5.

Bighorn

Date
Smokey**
6/8"/71
25
6/9/7"J6/21/71
227
6/22/716/27/71
62
7/8/71
7/11/71
7/14/71
7/15/71
6
7/16/71
7/24/71
9
7/28/71
2
8/1/71
8/5/71
1
8/18/71
8/13/71
8/15/71-4
8/20/71
5
9/1/71
10/2/7,1
11
10/18/71
11/5/71
11/15/7132
11/16/71
- 11/29/71
244
52
12/11/71
36
12/12/71
25
12/13/71
12/19/71
42
12/20/71
14
12/21/71
12/30/71
45
1/6/72
245
1/13/72
113
1/27/72
27
2/5/72
128
2/14/72
133
2/29/72
290
3/4/72
3/5/72
3/6/72
3/7/72
3/8/72
3/9/72
3/10/72
3/13/72
3/14/72
,'.

&amp;.~~'.

*i;*

Sheep Larval
Mom

211

Monitoring

Boss*

B'l ack i e

19

14
37

(larvae/gm.

feces)

B1ondie&gt;'&lt;*&gt;'&lt;
. Ram

Lamb

Program

13
31

171
0

80

12

35

0
0

130
0
4
199
22
8
12
8
106
30
11
67
51
233
128
12
131
41
10
151
56
84
23~'
2
222
387
153
177
66

20
20

10
31
23

17
31
3

66

11

24

45
8
21
59
70
147
204

7
8
10
21
22

16
6
12

59

lLf2

58
42
2
170
159
40
103
10
59
230
60
508

0
20
21
26
22

81
39
16
4·3
65

0
0
0
0
0
0

14
8

0
0

4·1
13
41
23

0
0
1

1
0
1
0
3
0

67
153

Boss given tramiso1 on 12/18/71Smokey given tramiso1 on 12/14/71.
Blondie died on 12/14/71.

0
0
0
0
0
0
0

7
5
2
0
0
2
2_
9
T-I
1
1
17
1
1
0
1
2

0
1
0
1
2
0

�-148-

Larvae Longevity

on Range at Little Hills

In early July, 1971, caged pellet groups collected from individual bighorn sheep were placed on the ground at various locations in the pasture
at Little Hills.
These groups were sampled periodically and have been
analyzed for larvae levels since that time. Larvae, at the same level
as when the study was initiated, are still being isolated from the pellets
(Table 6). During most of the winter the pellets were encased in ice
beneath 12-18 inches of snow. Temperatures reached 30 below zero. During
last summer ground temperatures exceeded 110 degrees Fahrenheit and larvae
were subject to severe dessication.
Apparently, the larvae possess the
ability to resist severe environmental conditions.

Table 6.

Larval longevity.
July

August

48.7
47.5
58.7

Ledges
Slope
Bottom

Ledges
Slope
Bottom

(All figures are larvae per gram of feces).
Sept.

Oct.

Nov.

Dec.

Feb.

59
65.5
65.2

55
21
13

21.7
9.4
3.6

25.5
56
9

44
17
6.7

42.5
27.5
6.7

25
35.9
48.3

109.7
58.3
110

33.3
25.6
11.1

30.4
·26
23

1.8
34.5
26.7

30
45.8
16

36.4
35.8
14

8.8
96.4
12.3

71.4
46
177

72.2
·88.5
30

28.8
35.4
9.4

37.3

56.2
72.1
28.3

86.4
37.5
20

12.9
13.3
20

27.3
18.2
25.8

26.4
30.5
33.3

o

7.8
9.1

31.3
60

8

lost

Boss
Ledges*
Slope **
Bottom***
Smokey

Mom

lost

12.2

Blondie
Ledges
Slope
Bottom

6

6

6.9

12
13.3

*Ledges:

Are the rock ledges found on the sides of hills in the pastures.
They are the driest areas in the summer and are blown free of snow most of
the winter.
The pellets were under snow for 3-4 months.

** Slope:

Forming the sides of the hills below the ledges.
most of the winter but it is less snow than in the bottoms.

*** Bottom:

They hold snow

Forms the ravines draining the pasture.
During the winter these
areas carry snow all of the time. In the summer these areas are the coolest
and dampest parts of the pasture.
Intermediate host snails are most easily
found here.

�-149-

SEGMENT OBJECTIVE
Develop

a procedure

for controlling

116

Protostrongylus

in bighorn

sheep.

PROCEDURE

I.
I'

Once the lungworm larval output for each sheep at Little Hills has been
established, one-half will be fed a vermifage (Tramisol) known to have
some efficacy against lungwonns.
Reduc t Lon in lungworm larval output will
be used to evaluate efficacy of this treabnent.
If the compound proves
effective, a means of administration will be evaluated using these animals.

Background

1"°- ••

Tramiso1 is a parasiticide, produced by knerican Cyanamid, and said to be
effective against lungworm (Dictyocaulus) and many gastrointestinal nematodes of cattle.
In this study it is noped that its efficacy against
Protostrongyline
infections of bighorn sheep ca.n be determined.
George
Bear has used it with some. success at Glenwood Springs where a dose of
2-3 mgm./lb. body weight suppressed larval output in bighorn sheep lambs
for up to 12 weeks, in a yearling for about 5 weeks, and in an adult for
3-4 weeks. This study is a continuation of that effort. It is hoped that
oral dosages, administered at a known level by gelatin capsules can be
related to specific decreases in layval output.
Also, it is hoped that
the technique of an oral oblet, capsule or dosage at a known level can be
utilized in bighorn sheep transplanting progrffiTIs.The next step is to incorporate tramisol into feed and to monitor its effects in treated sheep
versus controls.

Dosages with Tramisol--Earlier
tre.atTIe~Ti:s
at Glenwood Springs were made
with an effective dose of between 2 2nd 3 mgl. tramisol/lb. body weight.
This study was designed for delivery of a dose of about 8 mgm , effective
tramisol/lb. body weight.
Dr- Bergstrom of the Wyoming State Veterinary
Service in Laramie says Drs. Tho:C!.l.e ar:.d.
Harley have used this dose and
higher ones up to 16 mgm ,
body we:::'ght
in treating big}-,o';.l.!.
sheep ,
The
sheep selected for use in this study had Deen showing 12:cvae output greater
than the other sheep. The following tables give a description of the treated
(Table 7) and contro~ (Table 8) sheep and the dosages used for treated sheep.

ns.,

Table 7.

Treated

sheep.

'I'r amf.so L

Estimated Wt.
Sheep
Smokey
Boss
B10ndie

Description
ewe; 3 yrs. old
ewe; at least 3 years
ewe; yearling

Dose

(Ibs. )

(mgm. )

180

1440

200

1600
1040

130

Highest Larval
Output in Last
Hanth
(W!gm. feces)
244
204
40

�-150-

The control sheep were those remaining
in Table 8.

Table 8.

Descriptions

of them are

Control sheep_

Sheep

Mom
.·Blackie
Ram
Lamb

sheep.

Description

ewe;
ewe;
ram;
ewe;

at least 3 yrs.
yearling
yearling
lamb

Estimated wt.
(lbs.)

Highest Larval Output
ln Last Month
(lllgm. feces)

180

233

140
130
90

21

The dosages were encapsulated in Nmaber 11 gelatin
the Wyoming State Veterinary Service in Laramie.

o
17

capsules obtained

from

Pretreatment Monitoring Program--In the past six months over 100 fecal
pellet samples have been collected and analyzed for larvae level from this
group of sheep_
These samples have been dried and stored in envelopes.
Then a quantity of them is weighed, crushed with a mortar and pestle and
suspended for 24 hours in a Baermann setup.
The number of larvae collected
in the collecting tube is counted and related to one gram of feces as a
standard.
It is recognized that this technique is not the ideal method
of relating larvae output to infection level. No suitable substitute has
been found.
The monitoring program has established that larvae output in this group of
sheep is quite low compared to other Colorado and Montana counts.
Prior
to treatment this group of sheep was more intensively monitored than it
had been in the past months.
Between Dec. 11 and 14 daily samples were
collected from most of the sheep_
Table 5 shows the results of the monitoring program so far.

Treatment Techniques--Treated
sheep were tranquilized with 2 cc. of M-99 in
a physiological saline carrier shot in a dart syringe from a tranquilizer
gun. The tramisol was administered orally in a Number 11 gelatin capsule
given by using a balling gun. Nasal swabs were taken and preserved for Dr.
John Parks who is doing viral work on bighorn sheep.
Each of the treated
sheep were given 3 cc. of PDC, a penicillin-dihydrostreptomycin
preparation,
1M in the rump. Each of the sheep was given 2 cc. of M 50-50, the antidote
to M--99, IM in the rump when the work was completed.
Each of the sheep,
except Boss, was ear tagged for identification purposes.

�-151-

It was hoped that Tuesday, Dec. lL;, ';Jo-:.Jld be che day ';;JteI'" all the test
sheep cou Ld be treated.
OL odds &lt;:1ey t'viO sheep,
S;:,jOkeYa~'!.i BlancHe, ~.Jere
treated.
One of these
sheep" Blo~cdie, diee:.
This sheep -was more excitable
and younger t han the 0'LheT ":_~JO
_
~est sheep 2'-"10 she was trea'c:ed
under adverse
conda t Lons iYl d2e~) s; __
(/v.l or; E.. s-;:c2:P side!-l.ill..
;~{he Last
test
sheep was treated
on Su~day&gt; ~ec. IS.
The two sheep r~naining
show no ill
effects
and appear to De heelthy.

Post-Treat':!ent
Monitorin;;: ProgTeL.1--Af-i:er
Ltce fi::st
treatments
{Dec. 1-4)
the sheep were not coming down into the elfalfa
field
and would not let
people approach.
Since treatL"1g tll"" last
sheep ~ Boss&gt; (Dee. 18) they
have been easier
to approach a,-",:; f e ca L pellet
samples have been collected
(see 'I'ab Le 5)..
The Tesul-ts o~E tl~.:.is -~~i-=-~ i~ldicate tl"1at Tr.arnisol does no t
have any efficacy
against
=--,"-:C1.g\'lO~iTi in 2c-ult
b Lgho-rri sheep.
Dr" Thorne
(Wyoming Game and Fish) a::1':: :Jy. Pc s z , in i:2.depenacLlt
experiments
obtained
similar
r e su.l t s ,
This failu:ce)l af-eel: past succes ses , cou l d be due to
inconsistent
quality
cont rc i s he the p:coacct.

�-152-

ADDENDUM

1.

General

2.

Histopathological

and Special Protocol

(Bighorn
3.

Histopathological

of Tissues

Description

Investigations

at the Northern

from the 71

on the Epidemiology

in Bighorn

in Colorado.

Wild Sheep Council

Some Observations

Sheep in Colorado.

A report

during April 1971

of "The Lungworm-Pneumonia
A progress

report sub-

to the Commission.

Isolation
Sheep.

from the 71 BHL

Sheep) Collection.

Park, Canada.

mitted

and/or Necropsy.

of Lung Sections

at Jasper National

Complex"

6.

for Examination

Description

Bighorn Lamb Mortality
presented

5.

REPORT

Sheep) Collection.

BHL (Bighorn
4.

TO 1971 PROGRESS

of Parainfluenza

A publication

Veterinary

Medicine.

3 Virus from Rocky Mountain

submitted

Bighorn

to the American Journal of

�-153BIGHORN

PROJECT

1971-72
Department of Pathology
Colorado State University

General Protocol
Examination and/or Necropsy
Drs. Solomon &amp; Adcock
Field Collection of Bighorn Sheep
Dept. of G, F, &amp; P Personnel

1

History

i

i
W

\I

Live

Dead

1
Field &lt;:Necropsy

CSU

.-

...-- ...---

.------

.

1
,

!

II

1

csu

,t
Clinical Examination
and Evaluation
(Dr. Pierson)

I
I

J
.J./

MTCE

------- --'-....\
of Animal

.:::&gt;.

r-

Termlnatlon

or

/

A'
1
nlma

I

Necropsy

(G,F &amp; P Dept.)

----~------~-----------.~----------------~(----~---------y--~
Virology
(Hold Frozen)
(Dr. Parks)

Histopathology
and Impression
(Dr. Solomon)

• Toxicology
(Hr. Keiss)

Smears

~
Bacteriology
and Hycology
(Dr. Breen)

Parasitology
(Dr. Hibler)

/

"

1

Assay
(Mr. Keiss)

Clinical

! Pathology
j
I

(Dr. Rich)

I
j

~
Immunology
(Dr. Larson)

��-155-

Histopathological Description
of Tissues from the 7lBHL (Bighorn Sheep)
Collection

Dr. G.C. Solomon

��-157 -

BIGHORN Pl~OJECT
1971-72
DepaJCCiTLen'c
of Pathology
Colorado St az.e Urri.ver s Lt y

The following
individuals
spective
responsibilities
diseases
of b Lgho rn sheep

2.-::.3./or2ge::"1'::::les
are listed
with their
reor -:o:ces L~ t.ne investigation
of spontaneous
il".. 201orado fr.:&gt;m July 1&gt; 1971 - June 30, 1972.

Project
Leaders - Dr s , Jer-:y Accodz a:.-1 Charles
Hibler
Field Collection
of Bighorn S:-.ee? - DE:pt" of G&amp;u.e&gt;Fish &amp; Parks Personnel
Transportation
of Animals to 2SU - :Jcp':::" of Came , Fish &amp; Parks Personnel
Clinical
Examination
and Evaluation
- Dr Pf e r son , Dept. Clinical
Science
Clinical
Pathology
- Dr, Lon Ric~1.9 Dept. of Clinical
Science
Bacteriology
and Mycology - Dr. Baro16 Ereen, Dept. of Pathology
Virology
- Specimens f:cozeu for iutu:ce study
Parasitology
- Dr. Hibler 9 Dep': _ of ?c:L:c:ology
Histopathology
- Dr. Solomon.&gt; Dep;:, of Pathology
Toxicology
(selected)
- "i'lr. Robe r t Reiss&gt; DeV;:. of Game&gt; Fish &amp; Parks
Assay (selected)
- Mr. Robe'rc Reiss,
D2f-t. of Came , Fish &amp; Parks
Immunology (selected)
- Dr , Ke::-l Lar son , Dept , of Microbiology
Photography
- Mr. Pat H.::::Nar[~2:=c&gt;
De?t. or Pathology
0

�-158Bighorn Project
1971-72
Department of Pathology
Colorado State University
Special Protocol

I.

Field collection of Bighorn Sheep - Dept. of Game, Fish and Parks
Personnel
A. Complete history
1. Date, location, time of collection
2. Flock and site (area) observations
B. Live or dead collection
1. Method
2. Time
C. Photographs
1. Area
2. Individual lamb or adult
3. Flock

II.

Transportation of animals to CSU - Dept. of Game, Fish and Parks
Personnel
A. Method of transportation
B. Time enroute
C. Observations enroute

III.

Live animals
A. Clinical examination and evaluation - Dr. Pierson
L
Physical examination
2. Complete records
B. Clinical pathology - Dr. Lon Rich
1. Blood
a. Smears - Thick and thin - save slides
b. CBC
c. Hemoglobin and hematocrit
d. Platelet enumeration if indicated
e. Sedimentation rate if indicated
f. Parasites
g. Bone marrow biopsy if indicated
2. Serum
Serum sample will be divided"into two aliquots and frozen
a. Icterus index
b. Immunology - Dr. Ken Larson
1) Total protein
2) A/G ratio
3) Enzymes, ego SGOT if indicated
3. Urinalysis - routine
4. External lesions
a. Scrapings
b. Culture
c. Biopsy for histopathology
5. Fecal specimens
a. Parasites - Dr. Rubin
b. Gross observations

?

�-1:':9-

6.
IV.

Necropsy

Cerebral
-

spinal

fluid

if i~Jicn:2~

Dr. Solomon

If a Bighorn Sheep is to be t£:rminated

following the clinical

examination and evaluation,
electrocution
is tte mathod of choice.
Dead animals
transported
to CSU \-1i2..1 be :,acr0psiec. as soon as
possible
after
General

arrival.

A.

1.

2.
3.

,
4.

CompLet e ne c rc ps y xe c or ds by:.:rac'"scripc::'oI;'
from tape recordings
Piio t og r aph.i c equipment
available
~ 11::. Pat Ik:Namara
Equipment,
supplies
and re2.ge::ts
for c0I:':plete necropsy
a.
Fixatives
- Zenke rs ~ 5o,-~L~s, 3..."'lc. 10% buffered
neutral
fo rma Li.n ,
Unless 3~2'::i£i.2d~ 9.2.2..tissues
will
be

Virology
Samples selected
f.or ·vir0~0g~c:s..l st~C2-2:S \-1:"11 be removed
as soon as possible,
pl2.cad ::..,_w~:irl-,?acks&gt; labeled ~ and
stored
frozen
at -JOe c.
ao
Routine
co Ll.e c t Lcri
L)
Lung tissue
~ p2~ip~·~2~21 P\)=~i0:1. of e ach lobe (7)
and 2 dorsal portions

b.

2)
FeCeS ~ ~aeco-colic ~~23
Other tissues
if'indicaced

1)
2)
3)

4)

Blood

5)

Kidney
Liver

6)
7)

5.

Lymph nodas -- ~ila.r
.and :1.1ese:rit2ric
Sr.-leeL'Nerve 'Cissue:
PO:cti0::1S of bYa.:i.n and spinal

cord

Joints and synovia
Bacteriology
and my co Logy
Samples selected
for bacteriological
and mycological
studies
will
be take::1 as soon as possible.
Cauterization
of the
external
surface
w i.Ll. be emp Loye d and eseptic
techniques
for collectioc
will
be tol19wec.
The presence of pathogenic
ae r co Lc and ar.ae rob Lc bacte::i2.&gt;::nyccplas~11a1
and fungal
organisms'
will
be de te rmi.ne d ,
Pos i.t i.ve cul.t ures will
be s aved for further
study.
=r0:2 each qces t i.onab Le lesion
and/or
tissue will be madcv
80
Ro u t Lne co Ll e ct.Lor;
1) Respiratory
t r ac t ~ n as aL mucos a , Larynx, trachea ,
bronchi 9 Lungs
2)
fillomasal contents
3)
Sections
of ileum
b,
Other
t=-SSCleS if i"Ci3.:'C2t:2G.
1)
Ly~ph nodes ~ hilar and mesenteric
SDJec~1.
2)
3)
spinal cord
L) )

Bleod

5)
6)

~(idr~2y
L:i:Ve:':-

7) Joints and synovia
Co

Mjcopl.u.S~·.-~2
r: :"J2.~gc: 2\Jab S~~')i..:ld:,)2 ',:JscC to

suf f Lc Len t ma t e r i a L
fo::: C\.:::";:::c::e,. Tl:'12: 2\;J.c.b s:_0uld be ~le2.d in the refrigerator
cverll-:igh~ 0~ l(ept =::-oz~~':i£ cult~!'£.:1.g 'Wil:t.. be delayed
beyond
o'bta2.11

�-160-

12-18 hours.
impression smears - covered with spraycite
1. Nasal and conj unctival mucosa
2.
External lesions
3.
Trachea and lung
LI•
Bone marrow
5. Spleen
C. Integument and subcutis
1.
Description of external lesions
2.
Scrapings, cult ure, and spe cf.men for histopathology
3. Eyes
a. Remove eyes, eyelids, and lachrymal apparatus.
Remove
as much of 'the optic nerve as possible so that the
orbital rete is present
l&gt;. Trim orbital fat and conjunctiva.
Leave rectus muscles
attached
c. Place eye and trimmed portion in Zenker's fixative for
two hours.
Dilute fixative 50/50 with tap water and
maintain eyes for 2 hours.
Wash overnight.
The following
morning place eye in 70% ethyl alcohol for 24 hours.
Gross
and cut eye, optic nerve, and trimmed portions.
D. Cardiovascular system
1. Pericardial fluid - test for glucose and save for, culture
2. Blood - culture
3. Heart
a. Anterior and posterior papillary muscles; left ventricular wall
b. Right ventricle, atrium and pulmonary artery
c. Coronary and aortic vessels
d. Valves
E. Lymphohemopoietic
system
1. Spleen - samples for, impression smear, vt rologf.cal,and
bacteriological
studies, and histopathology
2.
Lymph nodes and tiss ue
a. Hilar and mesenteric especially
b. Others - Peyer's patches
3.
Tonsils
a. Lingual or faucial
b.
Pharyngeal
4. Hone marrow - impression smear and sample for histopathology
5. Thymus
F. Digestive system
1. Lips
2. Teeth - incisor and a portion of the mandible
3. Tongue - several sections
4. Soft palate - nasopharynx
5. Pharynx
6. Salivary glands
7. Esophagus
8. Stomach, intestine, caecum, colon, rectum, anus
9. Liver and gall bladder
10. Pancreas
11. Peritoneum

B.

�-161-

G.

H.

1.

J.

K.

Rcspi.r a t o ry system
1. Nasal mucosa
2. Turhinates
J.
Larynx
4.
Trachea and bronchi
s. LW1gS - fixed in Bouin' s and fo rmalin
i1.
Peripheral section from each lobe (7)
b.
Dorsal portion from diaphragmatic
lobe (2)
6.
Pleura
Musculoskeletal
system
1. Df.aph r agrn and heavy musculature
- gluteal region
2.
Skelcton--'
a.
Calvaria
0.
iZiglltfemur - wrapped in Saran l-]r.:;.f.; for Hr. Ke Ls a
c. Left femur
One half frozen; other half in formalin
d. Lumbar vertebra
1) One wrapped in Saran Vlrap for Hr. Keiss
2)
One half of a vertebra frozen; other half in formalin
e. 'i'empo raI region to Ln cLude ear canal, middle and inner ear
Open bulla slightly to permit penetration
of fixative
3. Ligaments
)
4. Joints, tendons, and tendon sheaths
i:l.
Tarsal joints
D.
Antlanto-occipital
joint
Urogc:nit:al system
1.
Urine for urinalysis and test stix
2. Kidney, ureters, large vessels
3.
i\ Lndde r and tirethra
Ii.
CUil ;JUS,
accessory glands ~ genitalia
5. rmmma ry glands and contiguous lymph nodes
G.
Fetus and placenta
Endocrine system
1. TIlyroids and parathyroids
2.
Adrcnals
3.
4.

Pancreas

1.

Brain
a. Remove cornp Le tely and hcmt se c t , Place one half in formalin.
From the other half remove sections from the medulla
ob Loug a tLon , ce rel.eLl.um, nri d=b raf.n region and telecephalon
for virological
and bacteriological
studies.
Place remainder

2.

Spinal cord
Hcn i.n ge s

Pituitary - remove pituitary
in ~;itll with
sphclloiJ 2nd other structures
5.
Pineal
~ervous system

.i.n f or'ma l Ln

3.
4",

Pe

r i.ph e ra I. rie

xve

&lt;&gt;

SeC£lGTI from sciatic

adjacent

parts

of

�\

-162-

Samples of tissues for histopathological
examination were collected
from different organs from all anatomical systems and from the majority
of animals.
Selected tissues were taken from animals which were markedly
autolyzed.
The most severe and consistent changes were observed in
sections from lungs. These changes were characterized primarily by perivascular cuffing of lymphocytes, lymphofollicular nodules, lungworms,
pleuritis, edema, the presence of neutrophils, hyperplasia of respiratory
epithelium, healing lesions, and necrosis and/or abscesses (Table Ml).
These changes seldom occurred singly.
Perivascular cuffing (accumulation
of lymphocytes around blood vessels) was observed in 20 of the 21 lung
samples.
Lymphofollicular nodules (group of lymphocytes) were observed
in 19 of the 21 lung samples.
In a normal healthy lung, these nodules
are very difficult to observe.
In the effected animals, the nodules were
very obvious and randomly scattered throughout the substance of the lung
section.
The six animals in which lungworms wer-e not observed in tissue
sections were from Trickle Mountain (4 of 6) and Buffalo Peaks (1 of 5).
However, these results do not agree with the results of parasite recovery
from the lungs by the parasitologist.
Edema and pleuritis (inflammation
of the thin membrane covering the thoracic wall, diaphragm, and lungs)
were quite common and are considered to be reactive processes to the concurrent pathology in the lungs.
The occurrence of healing lesions in 3
of 21 lung samples were characterized by the localization, death, and
removal of parasites and cellular debris by macrophages.
Necrosis or
death of tissue and/or abscess formation in 3 of 21 lung samples was/
were suggestive of bacterial infection and the presence of neutrophils
in the alveoli and bronchioles of several lung sections was probably related to the presence of bacteria.
Major histopathological
findings in
lungs by geographic area is presented (Table M2).
In several lung sections there were many areas characterized by the presence of numerous alveolar mononuclear cells (macrophages), thickened
alveolar walls, and hyperplasia of bronchiolar epithelium.
These areas
were not directly associated with lungworm, bacterial, or necrotic lesions.
These changes were similar to those described as "viral" pneumonia in other
species and are suggestive that agents other than lungworm and bacteria
may be involved.
A complete narrative description
is included in this report.

of the histopathology

for lung sections

A histopathological
summary of the examination of tissues except lung shows
that the lymphatic system is the one most commonly affected (Table M3).
It
is difficult in young animals to determine the growth and maturation of
the lymphatic system especially if these animals are exposed to an agent
which causes the lymphatic system to respond.
It seems that with the
animals collected through July 10, 1971, the lymph nodes and spleen were
not mature.
These were compared with animal #7, a yearling, in which the
nodes and spleen appear morphologically
to be mature.
Maturity or reaction
of the lymph nodes and spleen were based primarily upon the appearance of
the germinal centers.
Immaturity was characterized by a deficiency of mature
lymphocytes an~ rather well organized activity of immature lymphocytes and
zones of maturation from the center of the germinal center outward to the
periphery.
A reactive node was characterized by not only a deficiency of
mature lymphocytes but rather disorganized activity with no zones of maturation from the center of the germinal center outward to the periphery.

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1._..-

'1

Ii

r;

~l~

-f"

+

.!-

-I-

rj_

+

h.:~

I\")

+

e
1"''0

+

I;J

10

+

r':;

I"~

~!:

~j

+

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bj

~~ ~~~:~~

,

I-J.j

e

I~
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~

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I-'
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1-'

c»

r-'

NOW

.l&gt;-

r:x;

•

I-'

i·....

I-'

\.0

N

1'.)
-...J

N
0

N
f--'

(Kg,)

Autolysis

Mineraliza t i.on

Healing

Necrosis or
Abscess

Epithelial
Hyperplasia

Pleuritis

Edema

Ne ut roph i.l.s

Lymphoid
Follicles

Perivascular
Cuffing

Lun gwo rms

Area

Sex

VIt.

Animal No.

o

c;
::::l

[j)

OQ

I-'

'&lt;:

ti

III

~
:::.

c

C{)

o
,,'"
o
){)
'&lt;:

::r'

f"i·

f"

'0

(J

r+

[j)

/-'-

~{j

f••••
\D
-...J
1-"

r+

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(\l

w.

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Ii

:::l

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/-'-

/:).J

.

I-'

ro
~

0I-'

&gt;-'l
OJ

I

W
I

0\

....

�Table M2. Bighorn Project 1971.
Lung Sections.
FINDINGS
Perivascular cuffing
Lymphofollicular nodules
Lungworms
Pleuritis
Healing
Necrosis

Histopathological

findings by area -

T .N.

B.P.

S. C.

P.P. &amp;
RMPT.

+

+

-

+

+

5
5
4

0
0
1

4
4
4
4
2

2

1

3

-

5
4
2
2

2
4
4

0
0

6
6

1

2
4
1 4
3

1

0
0
0
0

-

6 0
6 0
6 0
6 0
3 3
1 5

I
I-'
(j'\

T.M. = Trickle Nt.
B.P. = Buffalo Peaks
RMPT. = Rampart Range

s.c. ::::
Sangre de Christo
P.P. ::::
Pike's Peak

+'
I

�I-'

N

Iv
0

Iv

-.J

(,.l

.J&gt;-

,

0

0

\.0

I-'

0

I-'

N

-.J

~

'"'"

7'

~~

I':Ij
1'Ij

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...."

,•...)

t"Xj

.

~s:

N

I-'

I-(J

,

"J

I'D

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,

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ti:J

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)-.7"

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0

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N

I-'
0

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N
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+

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+
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+
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______

+

+

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."

+

+
-I-

+

(l)

I'i

ru

+

+

q--------------_.
(I)

(fJ

+

+
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+
CfJ
I-'----,,-~---(I)

.J&gt;-

N

I-'
N

I-'

H

I':Ij

V1

t.d

I':Ij

0

I"'~

I':Ij

CXJ

b:J

J-rj

V1

+

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+

~

I Sex

I Wt.

,::£:

'0

No.

(Kg.)

I Animal

..

+

+

~.

+

+

+

Kidney

+

X

(1)

t:d

&gt;-0

Eosinophiles

I Thymus

I Eyes

Inclusion-like bodies
in lymphatics

Nematodes
Other tissues

t-i
'&lt;!

Ul

(1)

c

CfJ
CfJ

1-"

rt

1

~
III

CfJ

I-'

III

o

1-"

OQ

0
I-'
0

::r'

r+

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pJ

0

r+

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r"

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I...•
1..0

r+

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(D

w.

0

t-i

::l

0
t"i

:.T

1-"
()Q

Nematodes in
Lymph nodes

()Q

::l

I-'V-&gt;

~

(1)

t::

I-'
(1)

o-

1--'3
III

'0
rt

(')

Liver

~-=1

..
----

Heart

Spleen
Reactive

Lmma t ur e

:~::~ve

Immature
Lymph nodes

-. Lymph nodes
+-=------_
+
+

,

"d

----.-------------.-~----------....-

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()

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+

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.

+

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. · . . . I Area
·
t.d

I':Ij

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I-'

+:-

. . . . . · . . .
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I-'

---- .........-.,..,...,.--_._--------

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+'-

I-'

I-'
V-&gt;

.~-----...--.-""-..-.-------...
------,,-.--_&amp;_.~~..
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17;

..

1'-:&gt;

I-'
,

co
,

\!)

0

.

I-'

+'-

I......• I-'

I-'
Ul

0&gt;

N

I-'
0&gt;

. . . .

I-'

I-'

I-'
CXJ

------~~~,-------------~

.co

N

.

I-'

N
-.J

\.0

I-'

1-••.•

-----------

N
N

I

0'\
V1
I

~

�-166-

The thymus in each of the young animals through July _0, 1971 was very
extensive being observed from the midcervical region to the base of the
heart. Morphologically,
the thymus was mature and active. Within the
medulla of different lobes of the thymuses of 6 ardma.I s , nests of
eosinophils were present.
The significance of these are not known.

Heart--The heart sections from two animals (117 and #9) was observed to
have minor pathological alterations.
Sarcosporidia and a focal myoca~dial
degeneration were observed in the sections from animal #7, a female yearling. A focal area of inflm,~ation characterized by edema and lymphocytic
infiltration was observed in one heart section from animal #9.

Kidney--A very focal and minor glomerulosclerosis
and interstitial nephritis
were observed in kidney sections from animal #7, a female yearling.
Sections from animal #9 showed a focal toxic tubular necrosis and interstitial
nephritis.
A focal interstitial nephritis was observed in sections from
animal #18. These findings in the kidney sections from three animals
appeared to be very limited and minor and would not be considered significant in the functioning of the kidney.

Liver--Minor liver pathology was observed in six animals.
One focus of
degeneration was observed in liver section from animal #3. Hypertrophy of the
reticulo-endothelial
cells in the liver section from an.ima.Ls 119 and 10 was
noted.
Focal hepatitis (undetermined cause) was observed in a section from
animals #12 and #17. Chronic infl&amp;~nation characterized by focal tibro,sis
of the capsule, focal portal fibrosis, bile duct proliferation, and the
presence of a yellowish-brown pigment was observed in a section from animal

1114.
Nematode Larvae--Nematode
larvae were observed in sections of lymph nodes
from animals #8 and 1116. The ones (3) in animal lis appeared to be viable
while six in #16 were encapsulated and dead or dying.
In addition to
lymph nodes, degenerate larvae were observed in the submucosa of the o:.cal
mucocutaneous junction and the peril1.odaltissue of a non-mesenteric lymph
node of animal #16. In a section of thymus from animal #19, degenerate
larvae appear to be present in the medulla of two lobes.

Inclusion-like Bodies--In five of the animals, "inclusion-like" bodies were
observed in several of the reticular and mononuclear cells of different
lymph nodes and the spleen.
These were characterized by being nuclear,
acidophilic to basophilic, disorganization of the cytoplasm, and the presence of a halo around the body. These are called "inclusion-like" "because
they are only suggestive and must be confirmed by additional observations
and experimentation.
These were observed in animals #10, 12, 13, 17; and
20.

Miscellaneous Observations--Focal
from both eyes from animal 113.

conjunctivitis

A small focal area of mineralization
brain from animal 114.

was obServed

in sections

was obs€:.cved.
in a section of the mi':'='-

�-167-

Histopathological
of Lung

Sections

Description

from the 71BHL
Collection

Dr. G.C. Solomon

(Bighorn

Sheep)

��-169-

A histopathological
description of the lung seC::lons from 7lBHL-l, a
female lamb approximately
21 cays old.
Weight 12.5 kilograms.
Collected from Buffalo Peaks on 16 June 1971.
The lamb was in excellent
physical condition and no gross lesions were observed.
Microscopically,
there is marked congestion or all vessels, including the interalveolar
capillaries, the venules, arterioles and larger vessels.
Lymphatics
are not remarkable.
There is separction of lobules (slide #14) which
suggests interstitial edema.
Sections (slides #15 and 16) are 2~~~~cterized by marked congestion of
all vessels.
All stages (eggs, Larvae and adults) of Lungwo rm parasites
are present in the alveoli.
Ad"uILS were observed in the Lumf.na of some
bronchioles.
Accumu.lar Lons of. lymphocytes are present&gt; especially perivascularly.
Focal areas of epithelialization
are present, which may have
originated from the mucosa of terminal bronchioles.
Septal cells
(macrophages) are present and abundant in many alveoli.
Small foci of
polymorphonuclear
leucocytes are present.
The pleura adjacent to the
par as Lt e+af f e ct.ed lung is cuboidal rz the'r than squamous.
Lymphatics are
apparent in the subpleural area.
'l'hereappears to be hyperplasia and
hypertrophy of the epithelium of the bronchioles.
Lymphofollicular
development is present in focal areas (slide #16).
These appear to be
adjacent to the bronchioles and/or in lung parenchyma not associated with
the perivascular
lymphocytic accumulations.
Slides illS and 1116 are from
the dorsal region of the diaphragmatic
lobes of the lungs.
Following is a histopathological
description of lung sections from animal
7lBHL-2, a female lamb which weighed 4.8 kg.
Collected from Trickle Mountain
17 June 1971.
The lamb was in excellent physical condition and no gross
lesions were observed.
All 18 sections of lung were characterized by congestion of all vessels and focal areas of atelectasis.
There was no microscopic evidence of lymphofollic.ular development nor the presence of any
stage of parasites in the sections.
This animal was the youngest one of
the 22 which wer e selected.
Marked lobulation of lung tissue is evident
and probably nonnal.
Histopathological
description of lung sections from animal 71BHL-3:
A
female lamb we Lgh Lng 9&gt;.0 kg. co Ll.ect.ed from Buffalo Peaks 29 June 197L
The animal was in excellent physical condition and no gross lesions were
observed.
There is marked lobulation of. lung tissue which is probably
normal.
Congestion of all vessels is apparent.
There are focal areas of
blood in alveoli and focal areas of atelectasis.
There are also areas of
intra-alveo_ar
edema.
Perivascular accumulations of lymphocytes are present around one arteriole (slide #17).
A focus of peribronchiolar
lymphocytic follicular development is seen in slide #19.
Focal interstitial
edema is present {slide #26).
There is no microscopic
evidence of parasites
in lung sections.
Histopathologicel
de s cr Lpr Lon of lung sect:ions from animal 7lBHL-4:
A
female lamb which ~.veighed11. 5 kg. collected. 30 June 1971 from Trickle
Mountain.
The lamb was in excellent physical condition and no gross
lesions were observed.
There is mC!.:cKed
lobulation of lcng tissue.
There is congestion of all
vessels and focal areas of atelectasis.
There is focal thickening of

�-170-

pleura (slide #22). Many lymphocytes and mononuclear cells are present
in the lumen of one arteriole (slide #21). There are focal areas of
perivascular accumulation of lymphocytes.
No lungworms were observed.
Histopathological
description of lung sections from animal 7lBHL-S:
Female lamb which weighed 20 kg. collected from Buffa 0 Peaks 27 June
1971. The animal was in good physical condition and nodules were observed in the right posterior diaphragmatic lobe. These nodules varied in
size from 1-2 mm. and were cream-colored in consistency.
There are numerous areas of perivascular lymphocytic accumulations (slide
#19). One large focus in this lung section is characterized by marked
lymphofollicular development, perivascular cuffing, atelectasis of adjacent
lung tissue and caseation necrosis of alveolar cells. Within the center
of the necrotic area are several giant cells, fibroplasia and parts or
sections of dead parasites.
The necrotic center is surrounded by mononuclear macrophages, lymphocytes and eosinophils, and epithelialization
of alveolar and/or terminal bronchiolar epithelium is evident.
A small
subpleural focus is characterized by epithelialization and the presence
of lymphocytes, eosinophils and macrophages.
There is congestion of all vessels within this lung section (slide #10).
The pleura is normal except for slight thickening adjacent to the areas
of infection.
Lungworms are present in the lung sections.
The eggs
especially are visible with few larvae and few adults throughout the
tissue.
The most striking thing about this section is the rrumerous and
marked areas of lymphofollicular development.
In addition to this, there
are numerous foci of perivascular lymphocytic accumulat:ons around most
of the arterioles.
Within the large focus of lymphocytic development,
there are several giant cells. Areas of fibroplasia, atelectasis of
lung tissue within and adjacent to this large focus, and small pockets
or nests of neutrophils are present.
Th70 foci in the lung sections are
characterized by a necrotic area and mineralization and the presence of
dead sections of parasites. primarily larvae, which were trapped and
killed.
Surrounding these areas are accumulations of lymphocytes and
some fibroplasia that has taken place. Huch of the response of the lung
tissue to the infection in these described areas appears to be the
accumulation or presence of many alveolar macrophages, the presence of
neutrophils either in nests or singly and in addition, the epithelialization of the terminal bronchiolar epithelium or alveolar epithelium.
There appears to be hypertrophy and hyperplasia of some of the epithelium
in the proximal regions of the bronchioles.
Histopathological
description from the lung sections from animal 71BHL-6:
A male lamb collected 28 July 1971 from Trickle Mountain.
The estimated
weight was 23 kg. and the necropsy was performed by Dr. Hibler and no
visible lesions were apparent and the recording of the gross necropsy
was by tape. The tape was garbled, so there is no good record of the
necropsy, but apparently the lamb was in excellent physical condition and
there were no gross lesions observed.
Sections are characterized by marked
congestion of most of the vessels within the lung tissue.
In addition,
there is scattered throughout the lung tissue free blood within the alveolar

�-171-

spaces

and accumuLat.Lons of Lymphocy tes a::ound most or the arterioles.
areas W'1'}icl1Seer:l. to "[,2 ·~2:~.i·.c~i·:~·.lgp2:·.:Ci£·Dllic~21ar lY.:'ilPh,Qcytic
development close to broc,d!.i01es.
There is ,"0 evidence of parasites
in any of the s eccLons , There are focal areas of atelectasis
and some
which are probably emphysematous.
Ac:ctmmlation of blood in large
numbers of the alveoli
in the different
sections
may result
in difficulty in observing
parasites
if they ssec e present.

There are

Histopathological
descziptioC'l of hi.o_g sectio:"s
f::OYl1
animal 71BHL-7:
Collected
10 August 1971.
F'sD.ai2 yes.:::::'L"lgweighed 38.6 kg. ·which was
collected
from Bu:::':falo Feaks.
S·::-l'~ l':"...cd been dead about 14 hours prior
to necropsy&gt; but; she appea:,ced 'co be L"..eycel1e::-.t physical
condition
and there was no gross lesio~l.s c;bserved exc epr; those associated
with
the gunshot w011l1.d iri t·~'1e: Yt11d.C2.T\lic21 Te.gio~1.~
I.

There is evid.ence of ms:cke:C!
2."Jl:olysis wil::h all of the blood being hemolyzed withiTl the vesse:_s and witidn most of the alveoli.
·Where there
is no hemolysis
bLood , d"'..ere is ederaa in many of the alveoli.
The
ep i.t.heLf.um of the bronchf.o Les has autolyzed
to the point that normal
distinctive
morphology wes not retained.
There is however (slide
#8)
a focus in which there: is the preseEce or all stages of parasites
(eggs. larvae and adu Lt ) witI1.Ll. the: alveoli.
SurroUl_dii.....
g this area
and ad j ac enc to t':J.e b:colichioles .ifi thin t·bis ar e a , ar e ll1..rmerousfoci
of lynlphofollicular
nodule:s.
Also pre:se:nt is marked perivascular
lymphocytic accumu.Lat.Lon,

or

Histopathological
description
of lung sections
from animal 7lBHL-8:
A female bighorn lamb collected
19 August 1971 f rom Trickle Mountain.
The animal weighed. 27.3 kg.
She appeared to be in excellent
physical
condition,
although the posterior
borders OT both diaphragmatic
lobes
contained
yellowish.
circGY!scribed nodu Les , approximately
2-3 mm. in
diameter.
There was, in addition,
SO"f!1e pathology
that resulted
from
the gunshot; wound. hi terms of 1-_emo::.:Thage
and in che vent r aL anterior
cervical
region.
There is marked cOl"..ges
tion of all vessels.
hemolysis
of blood contained
Ln the ve s s e Ls (s::'..ide 118).
Ln addition,
there is
free blood iT" many of the alveoli,
"itich is fte:molyzed and there is
evidence of lYIT;.phoiollicular CE::'Je:10plT,ei.i:c
adjacent
to many of the
bronchioles
and in addition:::b,::re
is perivascular
ac cumu.La
t Lon of lymphocytes around t.he, smaller a:cterio12s"
Atelectasis
is present
in many of
the areas of this section.
LobuLat t.on of the lung tissue
was very
evident in those se:ctions which are not from the di2pnragmatic
lobes.
In addition
to the conge:stion of the vessels,
the presence of free
blood in the alveoli,
all of which is hemolyzed, and the atelectasis
in focal areas,
there are 2 major foci in w·hich there are all stages
of parasites
present
(slide. #11) a LL ·~Jithin the alveolL
These stages
are r ep r e.sen t.ed by eggs, la.:cvae, ard ad.ults and in addition
there is
the presence of adults
in the br0nchio~es
in one of the areas.
The
striking
thing about ·both of these elcas is that there is marked Lympho+
follicular
development,
especially
adjacent
to bronchioles.
as well as
perivascular
lymphocytic 2ccUi.Trulation. vJithin the lesions
themselves

�-172-

there is atelectasis and epithelialization
and the presence of many
septal cells or macrophages.
There appears not to be any mineralization or the presence of dead or dying parasites.
Adjacent to the
pleura in one of the affected areas there is also the presence of
these lymphofollicular nodules as well as the presence of accumulations
of parasites in the subpleural space.
A section contains parasitic stages with eggs, larvae and also adults
in the alveoli (slide #7). The most striking thing about this section
is the very marked lymphofollicular nodules.
Many of these are located
close to the bronchioles and in addition there is marked perivascular
accumulation of lymphocytes.
The atelectasis is quite marked in the
area where the parasites are observed and in addition there is a great
accumulation of septal cells or macrophages and epithelialization
occurring.
In longitudinal section of a bronchiolar lymph node, it is
characterized by edema in the peripheral zone of the lymph node as well
as in and among the germinal centers.
The microscopic appearance of
this node is one of a reactive node. There are pale germinal centers
and the presence of lymphoblasts and at the periphery more mature lymphocytes.
Congestion of the vessels is very prominent.
Cross sections of
adult lungworms are present in the sinuses of the zone (slide #20).
The histopathological
description of animal 7lBHL-9: A ram or male
bighorn lamb collected 23 August 1971 from Buffalo Peaks which weighed
28.2 kg. There were elevated nodules in both diaphragmatic lobes,
especially on the caudal posterior borders.
They ranged in size from
I mm. to 4-5 mm.
Some of these nodules were pale, with greenish-black
centers and others were plum-colored.
Whipworms were present in the
cecum.
In sections from the left diaphragmatic lobe there are foci
in the peripheral portion of the section characterized by the presence
of many large lymphofollicular nodules and the presence of lungworms
in and among these nodules with all stages represented (eggs, larvae
and adults of both sexes (slide #12). Most of these appear to be
located in alveoli.
There are numerous giant cells observed throughout
the parenchyma.
Septal cells or macrophages are present in the alveoli.
There are numerous neutrophils and free lymphocytes present in alveoli
and within nests throughout the affected tissue. Perivascular accumulation
of lymphocytes around the smaller arterioles is present especially in the
areas in which there are no lungworms.
There are foci of epithelialization
and it appears that these cells are originating from the epithelium of the
terminal bronchioles.
A section of lung from the left diaphragmatic lobe
characterized by atelectasis is prominent throughout the section (slide
#13). The vessels are congested and in the center of the section there is
a rather large focus in which all stages of the lungwonn parasite are
evident (eggs, larvae and adult) and all seem to be contained within
alveolar spaces.
The lymphofollicular development is not as marked as in
slide #12, but there are foci of these nodules and in addition there is
perivascular accumulation of lymphocytes around many of the arterioles.
Section of the lung tissue around the parasites is characterized by
atelectasis, the presence of many alveolar macrophages, the presence of
lymphocytic nodules as well as many lymphocytes in and around the parasites

�-173-

and in addition there are several foci of eosinophils in the stroma
surrounding the parasites.
The parasites appear to be alive at the
time the tissue was taken. There appears to be S0me epithelialization
and these cells appear to origiL3.te from the epithelium lining the
terminal bronchioles.
In addition&gt; in the Lar ge'r b ronchf oLes , there
appears to be some hypertrophy and ~yperplasia associated with the
lining epithelium.
This seems to be especially true or in close to
the area in which the parasites are present.
A section from the left
diaphragmatic lobe is characterized primarily by atelectasis, the
presence of much free blood in many of the alveoli, congestion of
vessels and a large focus in whidl different stages of the lungworm
are present (slide #14).
The pleura is not r emar kab Le , a.lchough
there is distention of many of t:1"'~&lt;2 Lymphat.Lcs that are located in the
subpleural :regions. Perivascular accu.mulation of lympr:ocytes is the
prominent

feature

in. this

secti2."i~ and again

there

is

not; tl~e marked

lymphofollicular development that there was in slide ff12.
All the
tissue sections are taken from the same lobeo Focal epithelialization
is present and again appears to originate fro~ the terminal portions
of the terminal bronchio1es.
Alveolar macrcphages are very numerous
and very large and fill many of the alveolar spaces within the affected
area of this lung section"
Accurnu Lat.Lons of Ileutrophils are present
throughout the affected portions of the lung. Cross section of several
adult lungworms are seen in alveoli which are some dista~ce from the
focus of extreme reaction.
Once again, the bronchiolar epithelium,
especially among the bronchi, in and close to the affected area, appears
to exhibit extreme degrees of hyperplasia and hypertrophy (slide #15).
Section of lung tissue from the right diaphragmatic lobe is characterized
by many areas or atelectasis and the presence of many lY:fiphofo_licular
nodules.
There is perivascular accumulation of lympho.::ytesaround some
of the arterioles.
The larger bronchioles 2;'".::1 smaller broneh i contain
within their lumina massive numb ers of cells
some or which are identifiable as alveolar or mononuclear mac:copl,ages and some of the cells can be
identified as epithelial" p robab Ly origL-lating from the .Lining epithelium
of the b roncha or b-ro ach Lo Les , Neut::-ophils are present in the alveoli,
but this is restricted to a few small f oc L. ,\)0 Lungwc rms were observed
in this section.
Ther e are foci of e;:,-;:ihysema,
especially in the subpleural areas. The sections from the diaphrzgmatic lobe are no t as
lobulated, if one compar-es ttis
"CO 'che lo'b"lations of the
other lobes of
the lung.
A section of lung tissue from t.he right: diaphragmatic lobe was taken from a
nodular area that was observed grossly (slide #16). This section is
characterized by marked atelectasis, some areas of 2mphysema, free blood
in a Iveo Lf and the

presence

of

3-4

smai Ler foci

in whi.ch Lungworms are

present.
There is not the marked presence of lymphofollicular nodules in
this section
as was previously seen in other sections containing lungworms.
however, there is rnuch perivascu:lar ac cum i.l.at Lon of lymphocytes
around many of the arterioles.
In the large affected focus, only eggs and
larvae of the lungworm are observed.
Section of the lung tissue in this
focus is sIm'iLa'r to that of other sections which contain the lungworm
larvae with the presence of nU::'Ti2rOUS al\Y2:o1armac rophage s , lymphocytes,

�-174-

neutrophils and few eosinophils.
Epithelialization
is also present and
again the cells seem to originate from the epithelium of the terminal
bronchioles.
In the smaller localized foci there are larvae which are
present and the reaction in and around these parasites of these larvae
are similar to that described for the large focus but not to the same
degree.
A section of lung from the right diaphragmatic lobe which upon gross
examination contained a focus which appeared to be greyish-white and
firm in consistency (slide #17). This section is characterized by the
presence of widespread atelectasis, the presence of perivascular
accumulations of lymphocytes around many of the vessels, but the most
marked characteristic of this section is the very numerous and large
lymphofollicular
nodules present in and around a large focus which contains within it all stages of the lungworm.
It is interesting to note
that there appears to be a chain of lymphofollicular
nodules just
beneath the pleura at the periphery of this lung section.
And in addition, there is, in a section removed from the large focus, the presence
of adult parasites in the alveoli.
The reaction of the lung tissue is
quite typical as has been described previously in that there are numerous
alveolar macrophages, lymphofollicular
nodules, as well as free lymphocytes in the affected area, small nests of neutrophils and the presence
of focal epithelialization
within the mass itself.
The extent of
necrosis of the alveolar walls within focus is difficult to determine;
however, it appears that there is atelectasis and the great accumulations
of the cells described previously.
A section of lung from the right diaphragmatic lobe which upon gross
inspection contained a greyish-white focus and was firm in consistency
(slide #18).
Changes present in this section are very similar to those
described for slide #17.
Sections from the hilar lymph node and changes are characterized by the
presence of free blood in the sinuses focally and a reaction in which
the germinal centers contain quite immature lymphocytes and scant mature
ones at the edge. There are along with the lymphocytes and in the
sinuses a number of large mononuclear round cells filled with an eosinophilic granular material.
A section of tissue from the right apical lobe
characterized primarily by the presence of free blood in almost every
alveoli, so much so that the normal architecture of the alveoli is very
difficult to discern (slide #12). In addition, there are several foci of
perivascular accumulations of lymphocytes around the smaller vessels.
No lungworm parasites were observed in this section.
Sections from the right cardiac lobe are characterized by the presence of
free blood in many of the alveolar spaces, focal atelectasis, emphysema,
and the presence of perivascular accumulation of lymphocytes around the
smaller arterioles (slides 23, 24, and 25). No lungworms were observed in
any of these three sections.

�-175-

Histopathological
description of aUDual 71BHL-IO:
Male lmnb weighing
12.7 kg. collected 26 August 1971 from the Sangre de Cristo Mountain
range. The animal was in very poor condition and the major lesions
were found in the lungs: lun.gworm lesions and pneumonia.
Sections from the r Lght; cardiac Lcb e are characterized by congestion of
all vessels and hemolysis of many of the erythrocytes indicating postmortem changes (slides 4, 5, 6). In addition, there are several areas
of atelectasis.
EpitheliDn1 of many of the smaller bronchi and bronchioles
has been sloughed away from the basement; membr-ane and in. the Lumf.na of
some of the bronchioles, there are cross sections of adult w rms and
larvae.
In one large focus within the lung parenchyma are the presence
of lungworms in all stages (eggs, larvae and adults) which are located
in the ~lveolar spaces.
In addition to the presence of numerous alveolar
macro phages and some epithelialization of bronchiolar epithelium there
is the presence of foci which is indicative of a bacterial infection.
Around these small foci are numerous ne.utrophils, lymphocytes and mononuclear macrophages with little or no connectiVe tissue.
There are many
foci of neutrophil accumulations in alveoli in which there doesn't seem
to be bacteria present and likewise there are many parasites in which
there is no netruophil accumulation around these that are located in the
alveolar spaces.
In another smaller foci within this S&amp;ile section, it
appears that there are many alveolar macrophages whose cytoplasm contains
blueish granular ma t.erLa L which pr esumab Ly is bacteria that has been
phagocytized.
In other smaller foci in the same section there appears
to be necrosis of cells and these might be the alveolar macrophages which
have been necrotized by the bacteria "which were engulfed" leaving
karyorrhectic nuclei and granular cytoplasm with ill-defined cytoplasmic
borders and these being in alveoli in the form of a granular debris.
In addition to showing marked congestion and desquamation of much of the
lining epithelium of the smaller bronchi and bronchioles, there are several
foci of different sizes, each characteristically
showing early to late
coagulative necrosis (slide 116). Foci in the section are characterized
by coagulative necrosis and at the periphery, blue granular material which
contains bacteria and peripheral to that are primarily mononuclear cells
with some neutrophils and all of this deliTnited by a cOl1.:r-.cective
tissue
capsule so that in effect it is a microabscess.
Smalley and earlier foci
show coagulative necrosis without the capsule hut with the presence of
mononuclear cells and some neutrophils.
There is some perivascular
accumulation of lymphocytes around the smaller vessels" but there is a
marked absence of lymphofollicular nodules.
In many areas the alveoli
contain alveolar macrophages and lymphocytes, neutrophils and plasma cells.
Sections from the right apical lobe are characterized prilnarily by the
appearance of relatively small to large foci of early to marked coagulative
necrosis (slides 1, 2, 3). The microscopic picture or the large areas of
coagulative necrosis are the srune as described for slide #6.
The pleura in some areas appears to be quite thickened anc lymphatics in
the pleura seem to be enlarged (slide HI). A section of pleura, in addition to being thickened, also shows inflammation in that many mononuclear
cells and neutrophils are present among the collagenous fibers.
These

�-176-

sections from the intermediate lobes are similar to the ones described
previously and characterized by a complete disruption in the microscopic
architecture of the lung (slides #7 &amp; 8). There is marked congestion
of all the vessels in the section.
There is desquamation of much of
the epithelial lining of the smaller bronchioles and bronchi.
There is,
in some foci, the presence of lymphofollicular nodules especially around
the bronchioles.
The disruption in the architecture observed grossly
which may have been described as consolidation, is due in part to
atelectasis of many of the alveoli.
In addition the presence of cells
within alveoli: alveolar macrophages, lymphocytes, and in some foci,
neutrophils add to the overall picture of consolidation.
There are in
these sections foci of coagulative necrosis varying from small beginning
or early necrosis to larger ones in which necrosis is well advanced and
also delimited by the presence of mononuclear cells and connective tissue
capsule.
In many foci, it appears that many of the alveolar macrophages
have coalesced into a mass of dark and probably degenerating nuclei.
These are within alveolar spaces and within the lumina of the bronchioles.
The pleura in these slides is not markedly affected, although there is
enlargement of some of the lymphatic vessels.
Lobulation of this particular section is quite marked and again this is quite different from that
seen in the diaphragmatic lobes, in which lobulation is either not very
well marked or completely absent.
Sections from the right diaphragmatic lobe present a somewhat different
microscopic picture than ones previously described from this animal
(slides 9, 10, 11, 12). There is a pleuritis and this is evident in the
pleura which does remain in the section.
In addition, there are at least
very small to large necrotic foci scattered throughout the section.
There
is a disruption in the architecture of the tissue in that alveoli are very
difficult to find. These alveoli are filled, as in previous sections, with
many alveolar macrophages, lymphocytes, and neutrophils, but in addition
there are several foci in which lungworm parasites are seen, some of these
different stages of the parasite are observed in relatively normal areas
or areas free from the necrotic foci.
There is also in a large bronchus
numerous parasites with larvae being the predominant stage of the parasite
found in the lumen.
There are, however, stages of the lung parasite (eggs,
larvae and adults) within the necrotic foci.
Some of these parasites seem
to be dying or dead and others seem to be quite healthy and existing amidst
the surrounding focus of necrotic tissue.
There are very few, if any,
lymphofollicular
nodules in any of the sections and likewise there is very
little perivascular lymphocytic accumulations around the vessels.
Again,
the consolidated appearance that might have been observed grossly between
and among the necrotic foci is probably due in part to atelectasis as well
as the accumulation of these different cells within the existing spaces
between the alveolar walls.
The presence of the accumulations of blue
granular material are probably colonies of bacteria.
These are also present
in the areas where there is early necrosis present as well as at the edge
of the larger foci of coagulative necrosis.
Sections from the left apical lobe are characterized by marked congestion
of all vessels, a focal pleuritis and the presence of small to large
necrotic foci (slides #13 &amp; 14). The accumulations of blue granular
material is probably bacteria.
The septa which separate these lobules is
somewhat edematous.
The architecture of the lung tissue is disrupted in

�-177-

the area where there is the Lecrotic
feci 2nd agaiL £he consolidation
which.
might
have
bec!l obs erve d g r os s Ly is
dUe in
part
to a t.e Lec t as Ls
and in part to the co LLect f.on or acc:wr,~la.tio-':l or ce.LLs it •.the alveoli
and especially
the alveolar
mecrophag2s.
The-:e is a section
of relatively
normal lung in this particular
slide in which the a~veoli are
quite distinct.
There is scattered
-::hrocgl':out this section
Lungwo rms
of different stages
(eggs. leyv2.e:, and adults).
These Ior the most
part are present
within
the alveolar
spaces.
Desquamiition of much
of the epi rhe Li.um c f t~le sl"aaller
bro:':--ichi and t.he brorichf.o.Les
is evf den t.
The microscopic
picture
of t.he s ec t.Lon f r om t~le left ca:cd..:'..aclobe is
very s iraf.Lar to that de s cx fb ed f or tile left
apical
lobe, except; that
lungworms were not observed in this s ect Lon (slide iIlS)_
f r cm
t.he
left dia'ptil:."agfD.2tic lobe
sugge
s t t'b.at the Inic.Toscopic
for ·(hese sections
is very similar
'to ·c~.at des cr fb ed fOT the Tight
diaphragmatic
lobe (slides
1f16,17,18;19). There is a. marked infection
of the lengwolil1 parasite with ell stages present and tneY2 are also larvae
that iire observed. in the smaLLer bTOnC~1.iana the hrmcchioles.
There are
also foci of early and late necrotic
focL
Se

c t Loris

p Lccirre

I

10

Hf s t opat ho.LogLcaL description
of tissues
from animal
71BHL-ll:
Lf9 kg"
Female yearling
bighorn collected
from the Sangre de Cristo Mountain
r ange 28 August 1971.
The animal was in excellent
physical
condd t Lon
grossly,
but had been dead about 36 hours before po st.Lug , Therefore&gt;
po stmo r t em autolysis
was very marked t.hroughouL
Sect:ions f rom the
ap i.ca.L and caro Lac and intermediate
Lob es showed primarily
hemolysis
of
free blced which was preSeil.t in ".::h2 alveoli.
Wf.ere lung tissue
is di8ce rriab Le , che re is present
SO·~Tle
revl to ·iuoderate numbers of Lymphof oLki ca.Lar
nodules and there is some perivascular
lymphocytic
ace'illuulation
around the
smaller vas s e Ls , Desquamation of most of the lining
epithelilnn
of the
respiratory
tract
is marked (slides
/15,0,7&gt; B, 9,10,11).
Section
from the
right
diep:'ycegms.tic lobe is from a Les i on described
grossly
as a smaLl,
greyish-white
area (slide
1112).
This section
is characterized
by v2ry
little,
if any, normal lung tissue.
There are several
foci where lymphofolliCular
nodules are present,
especially
adjaCent or near some or the
pre-existing
bronchioles_
TherE: are several
l-:lngwoymparasites
present,
especially
La'rvae and adu l.t s .
There is a.I.so present
'througOrwut 'c::his
lesion
much connec t Lve tissue
comprised
of yOU11g co LLagenous fibers
and
fibroblasts
and mature connec1:ive tissue,
There is a Tiiarked absence of
the alveolar
macrophages and other cells
'within the collapsed
alveolar
spaces _ There is marked hyperplasia
of bronchiolar
2pithelium
thro'u.ghout
the lesion.
Therefore,
this lesion
iT!. my opinion
represents
a healing
stage of t.he disease
r at.he r tl12i1. 2:.11. active
one ..

A sffiall greyish-white

area (slide
#13) was grossly
observed on a section
from the right
diaphragmatic.
lobe.
Changes observed in this section
are
similar
to those described
for slide
#12. except t.hat; the area of fibroplasia,
or healing,
is much mcre marked than that in slide
ff12"
There
is also rhe presence
of eggs, larvae,
ana adults
in the alveoli
which are
still
fur.c t Loria L. A s Lmf.La
r s"[£13.11grE:yisb.-'{olhite
ar e a was also grossly
observed on the left
dLaphr agmat.Lc lobe (slide
H14). Grossly,
tr_2: e:f,aEges

�-178-

were similar to those described for tissues used for slides #12 and 13
in that there is a large area which has been replaced by fibrous connective tissue.
In the sections from this particular animal, there is
an abundance of saprophytic bacteria.
There is also in this slide the
presence of lungworm parasites in all stages.
Some are in the area where
fibroplasia has occurred and there are also different stages of the parasite in the alveoli which appear to be more normal.
Lung sections are missing at this time from animal
was posted by Mr. Keiss and Dr. Post.

7lBHL-12:

This animal

Histopathological
description of lung sections taken from animal 7lBHL-13:
A male bighorn lamb collected 2 September 1971 from the Sangre de Cristo
Mountain range.
Ventral portion of the lobes of the right lung were consolidated and numerous adhesions between the costal pleura and the lung surface
were observed.
An abscess was also grossly observed on a section of lungs
(slide #1). Microscopically,
there is marked congestion of all vessels.
There is also free blood in alveoli of part of the sections.
The outstanding feature of the section is the appearance of most of the tissue to be
consolidated and the marked hypertrophy and hyperplasia of the lining
epithelium of the bronchioles and smaller bronchi and the foci of epithelialization which is present throughout the section.
The consolidated appearance is due in part to partial atelectasis of the alveoli and the presence
of numerous macrophages, neutrophils, lymphocytes within the spaces.
Perivascular lymphocytic accumulation is moderate around most of the smaller
arterioles and there is not much evidence of lymphofollicular
development.
There are numerous nests of neutrophils.
These nests are within the
alveolar spaces, although partially collapsed, and within the lumen of the
bronchioles.
There is no evidence of any stage of the lungworm parasite
in this section.
A section of lung from the posterior portion of one of
the diaphragmatic lobes is shown in slide #2. The changes observed here
are similar to those described for slide #1, except there is the presence
of parasite (eggs and larvae) within some of the alveoli.
In these foci
where the parasites are located there is consolidation and there appears
to be fibroplasia as well as the presence of the alveolar macrophages and
lymphocytes.
This focus is probably indicative of an early healing lesion.
In this section of the lung, it seems that the picture of consolidation is
due to a less chronic process than the one just described.
In this particular focus, there is the presence of numerous alveolar macrophages and
some lymphocytes, but the alveolar walls are still quite discernable with
a minimum of fibroplasia occurring.
One of the most outstanding features
of this section as in the previous slide is the hypertrophy and hyperplasia of the lining epithelium of the bronchioles and the epithelialization
which is evident.
A section from a diaphragmatic lobe and the changes seen
here are very similar to the ones described in slide #2, except that there
are numerous nests of neutrophils within the partially collapsed alveolar
spaces of the tissue (slide #3). Also lymphofollicular
development as
well as perivascular accumulation of lymphocytes is present.
The appearance
of this section is also less chronic than that described previously in that
there is numerous alveolar macrophages, lymphocytes and neutrophils.
There are few stages of lung parasites seen in this section.
Some of eggs
and adults appear to be dead or dying.
One area within this focus of

�-179-

consolidation is cha ract.e.rd
z.ed by the presence of dead paras Lt.e s and
fibroplasia which is in progress.
This wO;Jld s-ugges-cthat
there are
smaller foci of nealing or fibroplasia withir&lt; the larger focus of
consolidation whLch is less ch ror.Lc chan toat of che smaller foci. A
section of lung from one of the diaphragmatic lobes and again the microscopic picture is very similar to that described for slide #3, except
that there are more numerous nests of neutrophils in alveoli and
bronchioles (slide #4). There are present many larvae within the
alveoli but many of these appear to be dead or dy Lng , Th2"re is marked
fibroplasia within these areas of consolidation and in one or two foci
it appears that there is some mineralization
occurring.
There are
several lymphofollicular nodules as well as perivascular ac cumu La t Lons
of lymphocytes around the SInaller arterioles.
There is, as in the previous slide, marked hyperplasia and hypertrophy of the lining epithelium
of the bronchioles and epithelialization
which is evident throughout the
section.
Histopathologic~l
description for animal 71BHL-14: A male bighorn lamb
co Ll.e cz ed 2 September 1971 from the Sangre de Cristo Mountain range.
He weighed 18.2 kg. The general condition of the lamb was poor and the
major lesions observed grossly were in the lungs.
There were adhesions
of both apical lobes to the thoracic wall and the ventral third of all
lobes appeared to be meaty a~d consolidated.
There are numerous lymphofollicular nodules as well as perivascular accumulation of lymphocytes
around the arterioles (slide #1). The large focus in the section appears
to be one of consolidation and composed primarily of partially atelectic
alveoli and numerous alveolar macrophcges, lymphocytes and neutrophils,
and other macrophages wienie, the alveolar spaces (slide #1).
Epithelialization originated from the bronchiolar epithelium as well as hypertrophy
and hyperplasia of the b ronchf.o Lar epithelium itself.
No lungworm
parasites were observed in this section.
A section of the hilar lymph
node suggests a reactive node and the presence of edema at the periphery
or i:..1.
the cortex of the node (slide 1f2).
Erythrocytes are present: in the
sinuses of medulla but this probably is the result of "the gunshot ~mund..
A section from the right cardiac lobe and the microscopic changes in this
section are very sblilar to those described in slide III (siiae #3).
A
se c t Lcn

f:CCi:1~

the left

car df.ac lc~€: 2:~'1d nucr oscopf,c

changes

ir-l this

slide

are vei.-Y simi12:;'to those
GescTibed in slides 113 and 1 (slide #4).
A
sec t i.on rron, "c"he left diap"~~::-2g-:flatic
lobe a:.d the changes in ":::his
seetieD.
are char aet er i zed by marked consolidation rhz oughou c the Section and again
this consc Lxdat.Lcn
is due partially to atelectasis, but pr Imar Ll.y the
ac cumu Laz Lon of a.Lveo Lar raac rophages , lymphocytes, and neutrophils
in
alveol2Y s pace s (slice #5).
There ar e numerous Lympho fo Ll.LeuLar nodules
pr es errc ~ Theye is hype;~trop[,y and. hype rp Las La of the lining epitheli1.~m
of the bronchi and bTonchioles and also foci where epithelialization
is
evident.
There are I2"7 Lungwo rms in this section and those that are
observed appear to be adults.
They are located in the alveoli.
Congestion of all vessels is apparent.
These changes would suggest a rather
acute pneumonic process.
C~langes in these s ec t.Lorrs are similccT to those
described for sections from slide 5 (slides #6 &amp; 7). In addition, there
are
mrne'r o us Ei8.St cells
wh Lch are
pr e sent; witt! GeiiTiSa. strain
and ~tler2
ar e s eve.r a.L 2C-C~.-: lu~ng'Cv'orl113
lo~ated
:II"! t:j:le alveo~j_..
Thes e Ere '::O".!.1.fil'l2d
to Oi1.e s:nall section. of ti'12 SJ..lC:·2;" ,(,;Ji·cI-;;.ir_ SO~11e of t~1.esma.t l e r b~0nchi

�-180-

and bronchioles, there appears to have been a great proliferation of
the lining epithelium and the appearance of degeneration of many of
these lining cells into a rather homogenous pinkish material, which
lie primarily near the edge or within the actual lumen of these
altered bronchioles.
A section from the left apical lobe and the
changes in this section are similar to those described for slide #1,
a section of the right apical lobe, and slides #3 and 4, the right
and left cardiac lobes (slide #8). In certain areas of the pleura,
there is thickening and there is one focus of pleuritis which may
have been a point of adhesion to the thoracic wall.
No lungworms
were observed in this particular section.
A section from the left
diaphragmatic lobe and changes observed in this section are similar
to those described in slides #5, 6 and 7 (slide #9). A section from
the caudal border of the right diaphragmatic lobe is shown in slide
#10. The changes in this section are similar to those described for
the left diaphragmatic lobe, with the exception that no lungworms were
observed in this particular section.
Histopathological
description of lung tissues from animal 7lBHL-15:
A female bighorn lamb collected from Trickle Mountain 3 September 1971.
The weight of the animal was 19.1 kg. Grossly, the animal was in very
good condition and no visible lesions were observed.
Sections of lung
from all lobes and in addition included in this number are two sections
from hilar lymph nodes (slides #5-l5)~
The hilar lymph nodes are normal.
Sections of lung are characterized by the fairly normal except
for some perivascular lymphocytic accumulations around the arterioles
and few lymphofollicular
nodules near or adjacent to the bronchioles.
There is congestion of most of the vessels and there are foci of atelectasis.
Lungworms were not observed in any of the sections, although
one might be suspicious with the presence of the lymphofollicular
nodules and the perivascular cuffing.
Histopathological
description from lung sections from animal 7lBHL-16:
A male bighorn lamb collected 3 September 1971 from Trickle Mountain.
The weight of the lamb was 26 kg. The lamb was in generally good
physical condition and there were lungworm lesions observed grossly
on the caudal borders of the diaphragmatic lobes.
Sections of lung
from lobes other than the diaphragmatic lobes are characterized by
being relatively normal in appearance (slides #5-10).
There are focal
areas of atelectasis and smaller foci of emphysema.
The small arterioles
show perivascular lymphocytic accumulations and the vessels for the most
part are congested.
No lungworms were seen in any of these sections.
A section from one of the diaphragmatic lobes is characterized by congestion of all vessels, the presence of numerous lymphofollicular nodules
and a rather small focus in which numerous lungworm parasites are observed
and all stages are present (eggs, larvae and adults) (slide #11). There
is a cellular reaction as determined by the presence of alveolar macrophages, lymphocytes and some eosinophils in the alveoli and adjacent and
around the parasites.
There is perivascular lymphocytic accumulation
around the arterioles.
A section from one of the diaphragmatic lobes is
characterized by changes similar to that described for slide #11 (slide
#12).
However, there is no focus of lungworm parasites and the changes
associated with those parasites as described in slide #11. A section
from the diaphragmatic lobe which grossly appeared as a nodule and the

�-181-

changes in this section
are very similar
to those described
in slide
II
11 (slide
#13). A section Iron a diaphragmatic
lobe through a nodule
that was grossly
observed is shown in slide
1114. The changes in this
slide
are similar
to those described
in slides
#11 and 13. A section
from one of the diaphragmatic
lobes and the changes in this section
are
primarily
ones of ccnges t t on of all vessels,
rather
large areas of
atelectasis&gt;
perivascular
accumulations
of lymphocytes around arterioles,
a few lymphofollicular
nodules present
but no lungwonns were observed in
this particular
section
(slide
#15).
Histopathological
des.cr Lpt.Lon of the Lung sections
from anama.l 7lBHL-17:
A female bighorn lamb collecteD. 1.3 Septem"!:)er1971 f rom Pikes Peak area.
The weight of the lamb was 21 kg.
Grossly.
there were lesions
observed
in the lungs,
particularly
along the border of the posterior
caudal
border of the diaphragmatic
10'0e in "jhich several
nodules of varying
sizes were present.
On the cut surface
or some of the lobes a purulent
exudate was expressed
and fib-'::lL tags m~Te present
on the borders
and
edges of all lobes&gt; some of "ll1.ic~1. wec:e a t t ached to the pari.etal
pleura.
A section
from the right
diaphragr:,atic
lobe and this 5ec:t10'-1.'Nas t aken
from the gross sample in whic~r a ye:llowish area of quite firm. consistency
was obse rved (slide
1/5). This seC'1.:io:"iis characterized.
generally
as one
with moderate congestion
of all vessels,
the presence
of nuaerous lymphofollicular
nodules&gt; numero us Ties::s or neutrophils
in alveoli&gt;
some mild
to moderate perivascular
aCC;.ii"ula·cion of Lymphocyt es around the arterioles,
the presence
of many larvae
tr.i-.r£)"aghoutthe section
and the consolidated
appearance
of the lung tissue.i·herE:
are large number's of multicellular
macrophages which contain
at least
15 nuclei and completely fill,
with
the cytoplasm,
the alveolar
space.
L_ 3.G.Ctition, there 31.-e lymphocytes
and nests of rieut r ophf Ls . There is c.lso epithelializ3.tio:C':..
originating
from the lil'1in.g 2pi~r!.eliull1 of the ·b:,:-c"{.!.(:'~2.io12s 8.1'1(1ill t1'_e larger
bronchioles
and smaL'l.e r bronchi.
T::.c-.ceappears to be some hypertrophy'
and hyperplasia
of lini"ng e:"?i-cheliUiiL :vi2-~lY of the larvae s w:hich are
observed in the section
seem to be healthy.
while numer-ous at'_lers thrcughout the section
appear to De G.egeEeratingo
and in cross section
many of
the disintegrated
Larvae can be observecL
The pleura appears to be somewhat thickened
in some areas.
but no evideEce at acute Lnf Lammat.Lo
n is
suggested.
The changes observed in t{-,is section
are veTY similar
"Cothose
described
for slide
115 (slide If6).
T:ais section
is from the right
diaphragmatic lobe also.
The microscopic
pic:t-ure in a section
of lung from the
left
diaphragmatic
lobe, grossly
referred
to as lymphoid-like
foci,
is
very similar
to that described
for slides
#5 and 6 (slide #7).
The gross
description
is probably
true L_ '.::ILat there are many lymphofollicular
nodules as well as many large nests of neutrophils,
which would give it
the appearance
of a lymphoid-like
appearance ,
Changes in a section
from
the left
diaphragmatic
lobe wer e very similar
to those de s cr Ib ed for the
section
on slide
117 (slide
118).
Section f r om a portion
of lobe other
than the diaphragmatic
lobe and '.::1:12 chaug es iI"~ this section
are characterized
primarily
by numerous 1 npho f oLl.LcuLa'r nodules,
per tvas cuLar accufflulations
of lymphocytes ar ound arterioles,
edema, and numerous nests of neut roph Ll.s ,
many of which are in the lumina of the smaller bronchioles
and smallEr
bronchi
(slide
#9).
The appearance
of coriso Lf.dat Lon is not as marked as
ill. ~;~evio·i.2s s~:Lde3:1 but "c:ner2 ere :ca~tr:!.2r l:2.-.cge J.~-urnbers of a~l\..Ye&lt;)laT mac r o-:
pha.ges,
=-·.:l•.
)ypr~cc'yteE: ann T.te-otrc,pl1.ils ~LTl. t.l~.e a~-v'2,:,:"i ~ but:: t~h.e ar ch Ltee 'CUTe

�-182-

of most of the alveoli in this section can be discerned"
Edema is
present in the interlobular septae, which is very marked.
The pleura
of that which is intact does not appear to be abnormal.
No lung parasites were observed in this section.
A lung section from a lobe other
than diaphragmatic lobe and the changes observed in this section are
similar to t.ho se observed. in se c t Lcn If9 (slide illO).
In addition, there
are changes in the lining epithelium of these large bronchioles and the
smaller bronchi and this is in the form of hypertrophy and perhaps some
hyperplasia of the ciliated columnar epithelium.
A section from a lobe
other than the diaphragmatic lobe and the changes in this section are
similar to those described for slide #10, except the consolidation
appears to be more ma-,,:-l.(2':,iLl. ti:is
3e;::oti0T"i b ecaus e of the larger numbe-r
of a Lveo Lar mac rophages which are o'Dsc:rved in the alvecJ,.i&gt; but the
alveoli themselves are still disc21:nable
even 'with the presence of the
alveolar macrophages (slide #11). A section from a lobe of the lung
other than the diaphra~natic lobes and the changes here are very similar
to those described in slide #11 (slide #12). A section from a lobe
other than diaphragmatic lobe and the most outstanding feature in this
section is the presence of lyraphofo1licular nodules and perivascular
accumulation of lymphocytes around the arterioles (slide #13). There
are several nests of neutrophils in alveoli and in the l~~ina of some
of the bronchioles, but there is much less consolidation as judged by
the number of cellular elements in alveoli.
There are several foci of
atelectasis and few smaller foci of
physema.
No lung parasites were
observed in this section.
Sections or lung from lobes other than the
diaphragmatic lobe have numer-ous ly'iflphJfollicu12rnodules present; also
many nests of neutrophils and alveoli in the Imuiua of the bronchioles
(slides 1114 &amp; 15). The outstanding feature of these sections is the
alveolar edema which is present in almost every alveoli in this section
and accompanying this are moderate numbers of
veolar macrophages and
lymphocytes which are also present in the alveoli.
There is some
interstitial edema. The pleura, which has remained, is somewhat thickened
and there seems to be an enlargement of the lymphatics in the subpleural
region.
No parasites were observed in this particular section.
Histopathological
examination of lung sections from animal 71BHL-18:
Female bighorn lamb collected on 13 September 1971 from the Pikes Peak
area.
She weighed 19 kg. Nodules were present on the posterior caudal
surface of the diaphragmatic lobe grossly.
The lobes of the other portions of the lung were consolidated and purulent exudate COl d be
expressed from the cut surface.
Fibrin tags were present on the surface
of the visceral pleura and the bronchial and mediastinal l~nph nodes were
enlarged and edematous.
Section of lung from the diaphragmatic lobe which
grossly was observed as a circumscribed area and microscopically
the
outstanding feature about this section is the massive number of neutrophi Is in the alveoli and in the bronchi and bronchioles (slide 114).
In
addition, there are numerous lymphofollicular nodules, some of which have
been exhausted.
There is also marked congestion of all the vessels.
There is, in addition to the neutrophi1s present, large numbers of alveolar
macrophages and some lymphocytes in the alveoli.
There is also alveolar
edema and in some alveoli, the presence of granular fibrillar material.
Interstitial edema is present and in areas where the pleura is still intact there is inflammation of the pleura and enlargement of the lymphatic

�-183-

spaces in the subpleural regions.
Lu~gwonns were observed in this
section.
A section of lung from the diaphragmatic lobe and grossly
observed to be a nodular area is sho~" in slide #5. The changes in
this section are similar to 'chose described in s Lf.de !f4. &gt; except there
is not the degree and number of r:.aut::ophilsin alveo Lar spaces, but
there are many neutrophils in the bronchi and "bronchioles. TLere are
many more alveolar macrophages and neutrcphils in the alveoli.
There
are foci which are relatively normal in appearance in and around a large
focus of consolidation.
There are nume rcus lymphofollic-u:lar nodules&gt;
some or 1ihich appear to be reactive a:C!'Q
some at a point or having been
exhausted.
There are foci of atelectasis.
No Lungwo rms were observed
in this particula:c section.
A section from a diaphragmatic Lobe (slide
In) shows numerous Lymphofo Ll acu Lar r.od·:.::les
present t.hr oughout;
In
addition, there seems to be a marked change in the epithelium of t.he
br-orich
Lo Las which suggest hyperchromatic nuclei wit:, some hyperplasia
and hypertrophy of this epithelium.
The large focus of consolidation
appears to be one of a healing lesion in which there aTe not only
numerous mac rophages in the alveoli, but also fibrin and SOi e collagen.
Many slooth muscle fibers appea:c to be very evident in tbe lesion and
there are very few neutrophils present.
One isolated focus within the
lesion appears to contain dead lungworms.
In addition there is the
presence of multinuclear macrophages and this focus of dead lungworms
is surrounded and infiltrated with nmnerous lymphocytes.
In many areas
of the section dead parasites can be identified.
A section of lung from
one of the diaphragmatic lobes and the changes in this section are similar
to those described for slide #7 except no parasiteS, live or dead, were
observed in this section (slide #8). A section of lung irom a diaphragmatic
lobe showed varied changes (slide #9). Thera aTe foci of rather normal
lung tissue, foci of atelectasis and emphysema, marked congestion of all
vessels, the presence of some lymphofollicular nodules s.djacent to or near
bronchioles.
There is perivascular lymphocytic accumu.Lat.Lon around the
smaller arterioles.
There is one large focus which is very similar to
that described in slides 7 ffild8, in which it appears more of an active
healing lesion.
Throughout the whole section the lining epithelium of
the bronchioles and smaller bronchi is hyperchromatic and appears to be
hyperplastic. and hypertrophied.
In anoth8r focus, few Lung parasites are
observed, but these appear to be dead or dying and in this particular
focus there are numerous alveolar macrophages and some lymphocytes.
There
is a striking absence of neutrophi_s throughout the section.
There is
slight to moderate interstitia~ edema and the pleura, which has been preserved, does not show any marked changes.
A section of lung from the
apical lobe and changes in this section are characterized by the presence
of a few 1~fiphofol1icular nodules, perivascular cuffing around the smaller
a::terioles, congestion of most of the vessels, hyperc:nron1c:.tism
of the
nuclei of the lining ep Lt.heLf.um '(.7ith
hypertrophy and hyperplasia of this
epf t he l.Lum, the presence of many parasites of all stages, but each one
appears to be degenerating or dead (slide #10). The consolidated .appearance of many areas in the section appear to be that of a healing lesion
with the presence of alveolar macrophages, some neut ropha Ls , some lymphocytes, and fibroplasia that appears to be occurring.
There are sec&lt;::ions
in the lung which appear rather nonnal.
There are many areas of atelecLasis
and emphysema.
u

�-184-

Histopathological
description of lung sections from animal 7lBHL-19:
Collected 16 September 1971 from the Pikes Peak region.
A male weighing
17.3 kg. Grossly, several nodules were observed on the posterior caudal
borders of both diaphragmatic lobes and in addition the ventral third
of the right cardiac lobe and right apical lobe contained areas with
what appeared to be lymphofollicular nodules.
The general condition of
the lamb was good, as determined by fat deposition in the coronary grooves
and around the kidneys.
Sections of lung from the right apical lobes
characterized primarily by the presence of numerous lymphofollicular
nodules, many of which indicate or suggest a reactive state (slides #7
and 8). The center of these nodules contain many immature lymphocytes,
while the periphery does show mature lymphocytes but they are relatively
scarce in numbers.
There are focal areas of atelectasis and congestion
is present in all of the vessels that are observed.
There is no indication of any pneumonic process and no lung parasites were observed.
Sections of lung from the left apical lobes characterized by a relatively
large area involving more than one lobule in which there are lung parasites of different stages present (eggs, larvae and adults), and marked
consolidation of the alveoli in which there are numerous alveolar macrophages, lymphocytes, and nests of eosinophils (slides #9 and 10). Congestion of all vessels is marked.
There are several lymphofollicular
nodules and many of the smaller arterioles show perivascular accumulation
of lymphocytes.
There are sections of relatively normal lung and some
of the bronchiolar epithelium appears to be hyperplastic and hypertrophic.
Pleura in many areas is somewhat thickened and the mesothelial cells are
cuboidal in nature, rather than squamous.
Sections of lung from the right
cardiac lobe and the changes in this section are very similar to those
described in slides 7 and 8 in that there are numerous lymphofollicular
nodules, foci of atelectasis, congestion of vessels and some perivascular
cuffing around the smaller arterioles, but no consolidation nor lung
parasites were observed (slides #11 and 12). In slide #12 there are
numerous bronchioles and smaller bronchi whose lumina contain numbers of
neutrophils.
Much alveolar macrophages are present in many of the alveoli,
along with few lymphocytes, but the alveoli themselves are still discernable. Thus, there is not the consolidation and disruption of the architecture of the lung, as in many of the previous sections which have been
observed.
Sections of lung from the left cardiac lobes characterized by
the presence of numerous lymphofollicular nodules, many of which are
reactive, marked perivascular lymphocytic accumulations around small
arterioles and large confluent foci of consolidation in which there is
marked epithelialization
of the lining epithelium of terminal bronchioles,
the presence of numerous large alveolar macrophages and lymphocytes and
partial atelectasis and thickening of the alveolar walls (slides #13 and
14). Congestion is marked throughout the sections and there is hyperplasia
and hypertrophy of the bronchiolar epithelium.
The pleura in some areas
appears to be thickened and there is also enlargement of many of the
lymphatic vessels in the pleura and subpleural areas.
Sections of lung
from the intermediate lobe and these sections are characterized by the
presence of a few lymphofollicular nodules, congestion of most vessels,
some epithelialization
of the lining epithelium of terminal bronchioles,
a rather small area of consolidation in which there is partial atelectasis,
the presence of large alveolar rnacrophages in the alveolar spaces and some
thickening of the alveolar walls (slides #15 and 16). Parasites were

�-135-

observed

in both sections.
tl1.icke:::''1i~{'ig and ed.2·.:::~3. in tl'!.e pleu:ra..
S2c·~io:..1.s of Lung
from the right
diaphragmatic
~obe :::.re c~-2:C3.c'cerized.by re2.atively
large areas of consolidation
in w~!.ic"!:':
~:'"2::ceare GUl;lero-usLyrapho+
f o Ll.Lcu Lar l'loa:ules)I
SO;Ti.e.of 'wr~ic:~\2::-2 -:\:::~c.·Li:ve:; 2:.11 s'tag2s of Lung
parasites
pr es enr. (eggs, l&amp;rvb.s &amp;.:-.:.d 2c'~lt) i::.:.. ':=":lC 2.~\?eoli:; :''''::.·~_!inerous
large a1v201a:::-macr ophage.s a:.d lyap-:.;)c:y-::::es (slides
ill? s 18 3.1."&lt;1
19).
There appears to be a conspict:.ous absence
Le.utrophils"
There is
partial
at e Lect.as Ls or collapse
of the alveoli
and. h1. 80"[,120 an~as ,..rithin the Lazge ai"re.ct2G. r eg Lorr i'c 2P~)c&amp;''':s t~~at tricTE: is Su'lf~e fi~jYQPlasia
takil1.g
place..
-TrIere
a15;:) t!ie pr,sSE'!:'1.C2
fibers
'V.Thich aye -'\;f2~y appa.:ce::c.:.'~
~ "9JC'o·.:')2.-t~y arot:cA.1.C trlc ·::2r~T:.::':::..-.•.a L ~b:rOIj.cr...ioles
There is much epi l:heiializat:ion
ci:ld ':::h2 2.ii.1.ingcpithelh:,,'ii_ of '1:-£le larger
bronchioles
2nd sn aIle".::"bronc"j".i ap?ear to be hyperplastic
2n6 hypertrophic.
A large
nuabe r of lY111phofollicular nodu.Le s are app2rent just
beneath the pleura which is somewhat thickened
and congested.
Giant
cells
or multinucleated
cells
are prese •..
t within
alveoli
in certain
areas,
and some of the parasites
appaar to be degener at Ing or dead
while others
appear to be alive
and functionaL
Sections
from a hilar
lymph node which are characterized
by ederr.a in the co:ctical
regions,
reaction
of tr:e Lymph node il~ vihicI-:. t-:-le gerfflil'1al cent.er i:''1cicates stages
of immature lymphocytes with more mature lymphocytes at the periphery
and in the interstitial
area among the nodes the:ce is erythropoiesis
occurring
with the presence of few megakaryocytes
and nucleated
red
blood cells
(slides
1120 and 21). Sections from the left diaphragmatic
lobe which are characterized
by the presence
of a few lymphofollicular
nodules,
congestion
of vessels,
very little
perivascular
cuffing
of
smaI Ler 2!rterio:2s.
and a :cather clrrorrl.c picture
of s cor._solidated lung
section
in whic~-l there is pao:--sfal atelectasis,
the presence
of alveolar
macrophages and some lymphocytes
(slides
#22 and 23).
There are no
neutrophils
and the alveol.::!.r walls appear to be thickened
with in some
areas fibroplasia
and the marked presence
of smooth muscle at the terminal portion
of the bronchioles.

there

is

SO:TLe

or

or

hea.l

fng

is

of

s-..~:.oot1J.

Hi.stopat.ho Log.tcaf description

mus

cLe

0

of lung sections
f r om amma.l 71BffL-20:
Collected
16 September 1971 f r om the Pikes ?eek ar ea , Male bighorn
Lamb weig~,iEg 2.4 kg.
The condition
of t.1is lamb was ex t r eraeLy poor and
all Lobas c f bo t.h Lungs were af f ect ed to the degree that ve:cy little
f1.ln.c:~iorlal z Ls sue Yif8..S Lef t , Th e \j'ent~al -half of each lobe was dark red,
markedly cons;)lidated
3::ld cO:C1.t:aL-:ed
-:lillile:cousf ocf. of ·what appeared to be
Lymphof oLl.Lcu Ler nodules gzos s Ly , N-umerous nodules arid abscesses
were
present
-::hroughout most of the diaphra.gmatic
lobes.
Sections
of lung
from the left
apical
lobes 3re characterized
by -ehe presence
of numerous
lymphofollicular
nodu Les , many or which are reactive
(slides
#7 and 8).
Large areas of consolidation
indicate
a partial
collapse
of alveoli,
the
presence
numerous and large alveolar
macrophages,
lymphocytes and
neut.r oph.i Ls present
in a:-'..veo::'L M"G1Y or -the 2.lr,11iYla t~1.ebronchioles
contain masses of neutrophils.
There is marked congestion
of all vessels.
The pleura is t.hickened,
congested
and infla~~ation
is present
in localized
foci.
Interstitial
edema is moder at e between lobules.
There is marked
reaction
of the lining
epitheliu:.Tl of tIle bronchioles
in that there is
hyperplasia
and hypertrophy
end ex~e~sive projections
of vi~~i.
Lymphofollicular
no duLes are observed ill. ~t~1.is
particular
section
and t~'1ere is

or

0=

�-186-

also marked perivascular accumulation of lymphocytes.
Sections of lung
from the right cardiac lobe and the changes in these sections are very
similar to those described in slides #7 and 8 (slides #9 and 10).
Sections from the left cardiac lobe and the changes in these sections
are very similar to those described for the previous four slides (slides
#11 and 12). Sections of lung from the intermediate lobe and the changes
in these sections are similar to those described for the previous slides
(slides 1113. 14 and 15). Sec t Loris from the right diaphragmatic lobe are
characterized. by a marked pleuritis, numerous lymphofollicular nodules,
some of which are very reactive. p2rivascular cuffing of small arterioles,
foci in which lungworms are present and large areas of consolidation in
which there are partially collapsed alveoli or atelectasis, numerous
large alveolar macrophages with l}~lphocytes and nests of neutrophils
present (slides 1116, 17 and 18). There 2.:::\2' neutrophils present in many
of the lumina of bronchioles.
There is hyp erpLas La and hyperplastic
reaction of bronchiolar epithelium; epithelialization
is not as marked
in these sections as others.
There are localized areas of relatively
normal lung tissue. A section of lung tissue from the left diaphragmatic
lobe is shown in slide #19 and appeared grossly to be a nodule.
This
section is characterized by the presence of a number of lymphofollicular
nodules and a large area in which numerous lungwoTms are observed and are
primarily larvae.
In another section of this lung adults are observed
and in another section, eggs and larvae are observed.
The large area
in which many larvae are seen appears to be a healing lesion in that
there are many degenerating or dead larvae present.
There are numerous
alveolar macrophages, some lymphocytes and some neutrophils.
There does
appear to be fibroplasia in progress in some of these areas.
In contrast
to this in the area where the eggs and larvae are seen, there appears to
be more of an acute reaction with the presence of alveolar macrophages,
lymphocytes and some neutrophils.
The eggs and larvae in this section
appear to be alive and functional.
There is marked congestion of all
vessels and in the consolidated areas where the lungworms are present
there is partial atelectasis and thickening of the alveolar walls and
the presence of cells within the alveoli.
A section of lung from the
left diaphragmatic lobe through a nodule was observed grossly.
The
changes in this section are similar to those described for slide #19
(slide #20). A section from the left diaphragmatic lobe in which there
appeared to be lymphofollicular areas was grossly observed (slide #21).
This section is characterized by marked focal pleuritis. the presence
of many lymphofollicular nodules, most of which are reactive, and large
areas of consolidation in which there are partial atelectasis, accumulation
of large numbers of alveolar macrophages, lymphocytes and in focal areas,
and nests of neutrophils.
There are also neutrophils present in the lumina
of many of the bronchioles and small bronchi.
There are areas of relatively
normal lung present in the section.
Congestion is marked in all vessels.
There is some perivascular accumulation of lymphocytes around the smaller
arterioles.
No parasites were observed in this particular section.
Histopathology of lung sections from animal 7lBHL-2l collected 16 September
1971 from the Pikes Peak area: Male bighorn lamb weighing 18.2 kg. Thin
and in poor general condition.
The pathology that was seen grossly was

�-187-

primaTily
that in the lungs in which there was pL2urnonic involvement
of the right
lung. early fibrous
2-2:1esio':cs chat for-Heo. bet1Afeenthe
ventral
border or the ap Lca.I cazd i ac :;'obes 0.;::21.:h2 pari2"Cal pleura.
The vent.r a.L half of the whole lung -;lias rather
g:;:ey Ln appearance
and
these individual
lobules were demar car ed grey&gt; somewhat gr anu l.ar and
between each of these lobules was a distinct
hyperemic area.
Demarcation was relatively
sharp across the midportions
of '.::l':.eLur.g, 'I'here
were numerous scattered
nodu Les , presci"ably
containing
lungworm larvae
and adults.
A section of lung from one of the apical lobes characterized by a raar ke d pleuritis,
the p:cesel,-ce iIl areas or fi'brir. within
the
alveoli
and in other areas the presence of serum in the alveoli
is
shown in slide
1f7.
There is llYterstitial
involvement
and the presence
of fibrin
thin lymphatic ve s s eLs 'which are extremely
enlarged.
The
lining
epithelium
of the smaller bronchi and bronchioles
are hyperplastic
and in some cases hyper~rophied.
There aTe n~~2rous lymphofollicular
nodules present,
many of whi ch are reactive"
There are
numerous nests of neut roph i.Ls , congestion
of all '\i8sse2-s is marked
and some of the smaller
arterioles
appear to have hyalinized
vessel
walls.
There are four different
ioci ranging from small to very large
of basophilic
areas that appear to be oat cell ';o.acTOpC2.ges
which have
proliferated
and have "Deen fon:,2C1 iru::o rather
ci::ec::uler i..•
:2.SSes.
"("Jithin
some areas of these masses appear 'co be coLorn.es of bacteria.
There
are free macrophages within 'l,any of the alveoli
ani. in some foci there
is fibrosis
of the lung t Lssue itself
and especially
'"here there has
been involvement
and a telectas is.
No Lurigworms ,Iere observed in this
s e ct Lon, A section
from one oE the a~ical
lobes 2.::.0' the cnange s in
this section
&lt;:ire very similar
to those described
in #7, except that
there is much rcor e marked oat cell macrophage proliferation
than in
slide
if7 (slide
ji8),
Otherwise&gt; the changes are very siIr:ilar,
Lung+
worms were obs erved in this part LcuLax see t Lon,
See;tion of lung from
one of the apical
lobes is ShOWL in slide
#9.
In this pa:-:-ticular section there is relatively
normaL l'CTlg t:iSS"ll2 and an a:::-2:aJ.n wl-:ich almost
all of the lung tissue
is affectei".
Tl-:e demar car Lon is \l2ry sharp
followin.g along t.ne iIlte".:lol::lJ.lar
sep·~G..Ihoy,je~s
~=t~:_e"Ye is SOIne interstitial
LnvoIvercent; as well 28 a pleuritis:7
especialljJ
i:c. tl~·c pleura
that is covering
the affected
ar eas . \i:fithin the section
2::::efew eggs.
larvae
and adults
or
I-uIlg·worI!:[,s..
T:rLf:::::-_~e aYe n·i.lI!}.el~O·US 2_YTil~~:h.of.;)11icular
nodules,
some of 'which s eem to be qui'ce reacti"'le v 'i:i:'E:.reis c.iso
bronchiolitis
of varying degrees ar ound the different
b:c0nchioles.
Conge s t.Lon is very mar ked ,
Ther e are rie s r s of neu1:Top'hils"
ri'he lil1.irig
epithelium
of the bronchioles
is hyperp Last Lc and/or hypertrophic.
There is atelectasis
with the presence of numerous macrophage s ,
'I'her e is thickening
of many of the alveolar
septae 9 and in some cases
a fibrosisv
Epithelialization
of the lining
epitheliruu
or the terminal
bronchioles
is present
and in some cases this gives 'c"heappearance
"that
there are large multinucleated
cells
present.
There are neutrophils
in

,,"i

v

Some of the smaller vessel walls appear to be undergoing hyalinization.
In the more normal portion
of the section
there are lYil,-phofol1icular
nodules,
congestion
of vessels,
hyperplasia
and smn2 nypertrophy
of the
lining
epithelium
of the broTichioles.
Presence of neutrophils
is in
the lumen of the Dronchioles,
and in several
of the alveoli
there are
numerous alveolar
macrophages and neut rophf Ls present
arid so .e of the

�-188-

walls of the small arterioles seem to be degenerating.
A section of
one of the apical lobes containing rather normal lung tissue and lung
tissue which is markedly affected is shown in slide #10. The changes
that are present in this section are very similar to those changes
described for slide #9, except there is present a rather large focus
of oat cell macrophages which appears to be melted off by inflammatory
elements and surrounding this appears to be a conSiderable amount of
fibrosis of the lung parenchyma.
Parasites, especially larvae, are
very numerous in this section.
A section from one of the apical lobes
and the changes in this section are similar to those described for slide
#10 except no parasites were observed in this section and there were no
oat cell macrophages observed (slide #11). There is also no adjacent
or contiguous normal lung tissue which was present in slide #10. A
section from one of the apical lobes and the changes observed in this
section are similar to those described for slide #11, except the changes
are less marked than those in slide #11 (slide #12). A section from one
of the cardiac lobes characterized by relatively normal lung tissue in
terms of the ability to discern specific alveoli is shown in slide #13.
There are numerous lymphofollicular nodules and bronchiolitis is present
in almost every bronchiole.
There is marked congestion of all the vessels
and focal atelectasis.
No lung parasites were observed.
There is within
the lumen of one of the smaller bronchi strands of fibrin and a mixture
of cellular elements among which are macrophages and neutrophils.
Lining
epithelium of many of the bronchioles is hyperplastic.
There is some
perivascular accumulation of lymphocytes around some of the smaller
arterioles and in some of alveoli there are few to numerous alveolar
macrophages.
A section from one of the cardiac lobes characterized
by the presence of numerous lung parasites of all stages is shown in
slide #14. There is an area or focus of this section in which there are
numerous eggs as well as adults and larvae, and congestion of vessels is
marked.
There is much atelectasis and along with this thickening of the
alveolar walls.
There are several lymphofollicular nodules, and bronchiolitis is present.
There are small foci where in addition to the thickening
of the alveolar walls, there is epithelialization
of the lining epithelium
of the terminal bronchioles and fibrosis.
There are numerous alveolar
macrophages and lymphocytes present, especially within the areas or adjacent
to the areas in which lungworms are found.
Bronchiolitis is present and
the lining epithelium is hyperplastic and perhaps hypertrophic.
Neutrophils
are few in this particular section.
The pleura is somewhat thickened in
some of the regions of this section.
A section from the cardiac lobe was
examined and the whole section was affected (slide #15). Microscopically,
it gives one the appearance of consolidation and the absence of distinct
architecture of the lung tissue.
There are numerous lymphofollicular
nodules, some of which are very reactive.
Congestion is well marked.
There is bronchiolitis and the lining epithelium is hyperplastic and hypertrophic.
There is epithelialization,
resulting from the reaction of the
lining epithelium of the terminal bronchioles.
There are numerous foci
of neutrophils, both within alveoli and within the lumina of the bronchioles.
There are numerous alveolar macrophages and lymphocytes within the alveoli.
The consolidation is due partially to atelectasis and thickening of the
alveolar walls, but in addition, the presence of numerous cellular elements
within spaces between the alveolar walls.
There is pleuritis in areas or

�-189-

regions of the pleura which remain attached ~o the IUTi):! tissue.
No
parasites were observed in this particular section.
A section from
one of the diaphragmatic lobes is shown in slide #16. There is congestion of all vessels.
There is a large area within the section which
is relatively normal and the architecture is quite discernable.
In
another area of this particular section is the presence of supposedly
what was observed grossly as a nodule and &lt;Nithin this area there are
all stages of the lungworm (egg, larvae. and adults).
Most of these
are present within the alveoli and within this area appe ar s to be foci
in which most of the parasites are either degenerating or dead.
In
another focus the parasites seem to be rather healthy and alive at the
time of death of the animal.
In this nodular area, then~ is atelectasis
and. the presence of alveolar macr-ophages , In t.hf.s small fo cus in which
the parasites appear to be degenerating or dead there s eems to be f ewe r
macropZlages than in the area where they appear to be alive.
The alveolar
walls have become thickened and there is some fibrosis present.
The
smooth muscle of the terminal bronchioles in this particular area is
very easy to d:j.scernand it appears that there may be hypertrophy of
the smoot h musc Le , There is bronchiolitis present in all the bronchioles
-withL,- this particular section.
There are numerous lymphofollicular
nodules
acme of wh i ch appear to be very reactive.
There is epithelialization and there are small areas in which multinucleated
giant cells are
forming or have formed and these seem to be originating from some of the
lining cells of the epithelium or the alveolar lining cells. The pleura
appears 'Co be rather nOr!nal even over the area where the nodule is located.
Lining epitbelium of the bronchioles and smaller b ron ch.I appear to be
somewhat; hypertrophic and hyperplastic.
A section from one of the
diaphragmatic lobes is shown in slide 1f17. The changes in this section
are similar to slide 1116 in which there is a relatively normal lung
tissue and areas in which are probably those described grossly as nodules
and within these nodules there are numerous lung parasites of all stages
present.
~nthin these nodules are many parasites which are degenerating
or dead and the reaction of the s'Jrrounain-s tissue is one primarily of
healing in which there are numerous macrophages and mahY of these
macropnages do contain pigment which probably origi:n&amp;tes from the degenerating parasites within the alveoli.
There is epi'(belialization
of the lining epithelium of the tenni:-:.al bronchioles.
The smooth muscle
fibers stand out very mar~edly in these areas anQ there is a rather marked
absence of neutrophils.
The consolication as i..t.t previous slides is due
Ln part to collapse of the alveolar waL'l , or at.eLeccas Ls ,
The thickening
and in some foci, fibrosis. of the alveolar wall and the presence of the
macrophages within the spaces are partially responsible for the consolidation.
There are ll.tmlerouslymphofollicular nodules, some of which appear to be
quite reactive.
Bronchiolitis is present and the lL1.ing epithelium of the
bronchioles is reactive and appears to either be hyperplastic or hypertrophic.
There is a focal pleuritis.
A section from the diaphragmatic lobe and the
changes in this slide are very similar to those described for slide 1f17
(slice #18). Sections
rom one of the diaphragmatic lobes and the changes
observed in these sections aTe similar to those described for slide #18,
but less marked than in that particular slide (slides #19 and 20).
9

�-190-

Histopathological
description of lung sections from animal 7lBHL-22:
Collected 26 September 1971 from Pikes Peak.
Female bighorn lamb
weighing 27.2 kg. It was found dead and before dying it was observed
the day before walking with a lateral motion and very slowly.
Autolysis was extensive throughout.
The animal seemed to be in generally
good condition.
The hair coat was smooth. Upon posting, the thoracic
cavity contained quite a volume of blood colored fluid.
The lungs
contained 3-4 palpable nodules in the caudal dorsal region of the
diaphragmatic lobes.
The ventral third of all lobes were consolidated
and purulent material could be expressed from the cut surface.
A
large abscess of approximately I cm.3 was present in the right apical
lobe. More abscesses were present in the ventral third of the intermediate lobe.
Section of lung tissue from an unidentified lobe
characterized by congestion of all vessels, widespread atelectasis,
very little change in the lining epithelium of bronchioles, and no
lymphofollicular
nodules nor perivascular cuffing of arterioles was
shown (slide #1). In one region of the section it appears that there
is a pleuritis and an organization of fibrin in alveoli deep to the
pleura with the presence of many mononuclear cells and this may have
been adjacent to or close to an area in which a small abscess was
described in one of the anterior lobes. A section of lung from one of
the diaphragmatic lobes is shown in slide #2. In general the whole
lung section is affected.
The most striking thing about this section
is the tremendous number of parasites of all stages (eggs, larvae and
adults) but especially the large number of eggs that are present in
the alveoli.
There are numerous lymphofollicular nodules, some of
which are very reactive.
There is perivascular cuffing of some of the
smaller arterioles.
The pleura has been thickened greatly and some
areas show a marked infiltration of lymphocytes especially within the
pleura and subpleural space.
Bronchiolitis is present and there is
some evidence of reaction of the lining epithelium in terms of hyperplasia and perhaps hypertrophy.
The alveolar walls have been thickened
and has been partially collapsed.
There are numerous alveolar macrophages and lymphocytes present.
There is very few, if any, neutrophils
present.
A section of lung from the diaphragmatic lobe and the changes
in this section are very similar to those described in slide #2 (slide
#3). A section of lung from one of the diaphragmatic lobes is shown
in slide #4. This section is characterized primarily as consolidation
with very few alveolar spaces recognizable.
There is a marked pleuritis.
There are several paribronchial areas in which there is an accumulation
of lymphocytes and a chronchiolitis.
The lining cells of/the bronchi
and bronchioles have reacted and appear to be somewhat ~yperplastic or
hypertrophic.
The consolidated appearance of the lung tissue is due to
the thickening of the alveolar walls, the partial collapse of many of
these walls and the presence of alveolar macrophages and lymphocytes.
There is also hypertrophy of many of the lining cells although they are
still attached to the alveolar wall.
There are numerous nests of neutrophils, both within the alveoli and within the lumina of bronchioles.
Few lungworm larvae are present in the alveoli.
A section of lung from
an unidentified lobe which appears microscopically
to be a section from

�-191-

the diaphragmatic lobe is shown in slide #5. Marked congestion of all
vessels is present.
There is diffuse or widespread atelectasis or consolidation of lung tissue.
The bronchi and bronchioles appear to be
quite normal.
There are few lymphofollicular nodules present.
No
lungworms were observed in this section and again the consolidated
appearance of the lung tissue appears to be due to partial thickening
of alveolar walls and accumulation of macrophages within the spaces,
and hypertrophy of some of the lining cells which are still attached
to the wall.
Section from a hilar Lymph node is shown l:i slide #6.
There is marked congestion of all vessels in the lymph node, The node
presents a picture of being a very reactive one in which there is
exhaustion of many of the geL'il1incl
cerrt.e rs with very rei.)" 1l1at-Grelymphocytes at the periphery of the cent.er , The sinus of the node and the
spaces around the trabeculae contain a mixture of cells, most of which
are plasma cells, but also scattered among these are neut r ophf.Ls .
Reticular cells are quite prominent throughout the tiSSUe and there are
also immature lymphocytes of the Lymphob Last Lc series present.
A section
from the right apical lobe in which an ab sce sswas observed grossly is
shown in slide #7, This sec t tcn is characterized by marked bronchiolitis,
marked congestion of all vessels, interstitial edema, diffuse atelectasis,
and the presence of many alveolar macrophages within the spaces.
No
parasites were observed in this section.
There is no visible evidence
of an abscess within this section, although along the pleura in one region
it appears that there was an extremely inflammatory r-es pons e and there is
a marked pleuritis.
Two lungworm larvae were observed in the alveoli of
this section.

�-192-

BIGHORN

LAMB MORTALITY

Colorado Division
University,

INVESTIGATIONS

IN COLORADO

of Game, Fish and Parks and Colorado

College of Veterinary

Medicine

and Biomedical

State
Sciences

The Sangre de Cristo mountain range in south-central Colorado is a long,
narrow chain of rugged peaks and steep rocky slopes.
The portion lying in
Colorado is over 100 miles long, and rarely more than 12 miles wide at any
point along its length.
This range contains 8 of Colorado's 54 mountain
peaks over 14,000 feet high. Rocks are largely of sedimentary origin, not
highly mineralized, and for this reason the area has not suffered the impact
of mining activity so common in other parts of the state.
Only a very few
roads exist, and with the exception of one major east-west highway, all are
quite primitive.
The earliest explorers documented the presence of bighorn sheep in this range,
and throughout the latter part of the 19th Century, up to the present date, it
has been the location of one of the major bighorn herds of the state. At one
time, sheep could be found the entire length of the range, but during the
past two or three decades, the herd has become more or less concentrated in
one portion not over 15 miles long. With this concentration has come a
decrease in total numbers, but as recently as ten years ago the herd was
judged to be healthy on the basis of population age-class structure.
Starting some time in the early 1960's, a drastic change in age-class
structure began to occur.
Survival of lambs to reach the yearling age-class
diminished to a very low level, reSUlting in almost no recruitment and
replacement.
The herd continued to decline in numbers, and of course became
dominated by old age-class members.
By 1968, it became obvious that the
annual lack of herd increment was a persistent and recurrent problem that
showed no indication of abatement.
The first investigation aimed at identifying the cause of the problem was
a two-year study done by Woodard, then a graduate student in the Dept. of
Fishery and Wildlife Biology, Colorado State University, and initiated in
the spring of 1969. Objectives of this study were to: (1) ascertain herd
composition, (2) determine breeding success in terms of conception and live
births, (3) follow the herd from time of lamb drop through the summer and
fall, and document lamb survival, and (4) assess the importance of predation,
trauma, weather, and disease in mortality of lambs.
It became evident early in the investigation that the problem did not involve
lack of conception or failure to bear live lambs. By mid-June of 1969, most
breeding-age females in the herd were accompanied by seemingly healthy and
active lambs. The same pattern was repeated in 1970. Also, during both
years the ewe:yearling ratio was exceptionally low (100:11 and 100:17,
respectively) reflecting the lack of lamb survival from the previous year.

�-193-

According to previous experience, the classic symptoms of respiratory illness
should have appeared by mid-July in lambs frcm Euffalc Peaks.
When they did
not appear, it was decided by memb ers of the ream that collections should
immediately begin in the Sangre de Cristo Range, in spite of the handicap of
almost impossibly difficult terrain.
Ultimately, all collections
made in
the Sangre de Cristos in 1971 were done by helicopter.
By early August, lamb
mortality began to appear on Pikes Peak, approximately 75 miles east of
Buffalo Peaks.
The Pikes Peak herd had previously been considered to be
a very healthy herd, having a high rate of recrui tmen t and replacement.
The Sengre de Cristo and the Pikes Peak herds thus became the collection
locations fer sick lambs. By the end of Septe,£i;:'er.
five lambs had been
taken from. each of these tsao herds&gt; 2.::dJ.lorD3;.li ty of remail:!.inglambs was
almost complete.
Lambs collected from the Buffalo Peaks and Sagua che Creek-Trickle 110untain
herds were essentially healthy i~ appearance and of nonnal weight for their
age.
Postmortem examinaLlon of these lambs did not reveal any significant
pathological changes (Tables 1 an~ 2).
Five of the 18nbs were infected
with Protostrongylus
s t Ll.esI , bur the :':"evel
of i'Clfectior.
"JaS quite low and
no significant lesions compatible with "verminous pneumonia" were present.
Lambs collected from the Sangre de Cristo and Pikes Peak herds were
obviously ill, small for 1:1:.eir
age and light in color.
Coughing and
dsypnea were frequently observed.
Pos tmo r rem examination revealed that
all animals, except a yea.rling accidentally killed in the Sangre de Cristo
herd, had pneumonia compatible with "verminous pneumon.ta'".
Protostrongylus
stilesi wa.s numerous in a_I of these animals (Tables 3 and 4).
One lamb frOB ~he Sangre de Cristo he~d and one from the Pikes Peak herd
had a species or Pasteure-Jla and a gram rieg a t Lve diplococcus, chough t to
be a species ih the genus "Neisseria., in their lungs.
An adciitiona.l lamb
from Pikss Peak also had this gram nega.tive diplococcus in the lungs.
Bacteria.l results are incoThp12te as yet, but: thus far the results have
not revealed any add.itior.&amp;lLambs infected with psthOgei::ic species.
Virological studies are '::ime-consuming and consequently, incomplete at rhi.s
time. Low titers (1:2 to 1:64) aga.inst bovine PI-3 virus 'was demonstrated
in two Lambs from Buffalo Peaks, a.ni f ou r lambs f rom Pikes Peak.
Gastro-intestioilal parasitism was not a problem in any ci these lambs.
Very lig'nt infections \.JithMarshalla.gia n1arshal1 -1, Nematodirus spp. and
six sp ec Les of Eimeri a '.Jerefound in 10 of 20 Lamb s .
This rese2.rcD.is currently being i.nv2s tiga'c:edby a team. cons is t i.ng of
William H. Ruthe:::-ford.Wildlife Researcher; Thomas N. Woodard, Gradual:e
Student, 'C,.0T,J Senior Research 'I'e chn Lc.Lan; Gene Schoonveld, Wildlife
Researcher; Robert Schmidt, Wildlife Technician; George Bear, Wildlife
Researcher; Robert Keiss, Wildlife Researcher; Gordon Solomon, Associate
Professor of Pathology; John Parks, Assistant Professor of Virology;

�-194-

Loss of lambs in the herd began to occur generally in late July and early
August during both summers.
The rate of loss reached peaks in early September
of both years, and by mid-September most lambs had succumbed.
The ewe:lamb
ratio dropped from 100:83 on July 3, 1969 to 100:17 on September 10, 1969;
and from 100:72 on June 30, 1970 to 100:22 on September 18, 1970.
At no time during the course of this two-year study could any lamb mortality
be ascribed to predation or accidents.
Physical appearance and observed
coughing in lambs, beginning in late July, 1969, indicated that disease was
the most likely factor in mortality, and when the pattern was repeated in the
summer of 1970, it was decided to collect lambs in an attempt to learn something about this phenomenon.
Accordingly, two lambs were collected, one on
August 6, 1970 and another on September 9, 1970. The first one collected
exhibited early stages of two different types of pneumonia, a verminous
proliferative type and a purulent bacterial bronchopneumonia.
Although not
clinically ill at the time of collection, it appeared to be host to elements
that could bring about its eventual demise.
The second lamb collected exhibited the classic pattern of advanced
respiratory illness, diagnosed as verminous pneumonia.
It was in extremely
poor body condition, and at the time of collection appeared not to have the
strength to accompany the herd in its flight from danger.
It stood without
moving while the rest of the herd ran out of sight.
The peak in lamb losses during both summers, but particularly during the
summer of 1969, appeared to coincide with a period of cold, wet weather,
more or less typical of the Colorado Rockies during August.
Critical
analysis of the combination of circumstances and conditions encountered by
these bighorns, seemed to support the conclusion that high lamb mortality
was the result of respiratory disease caused by lung parasitism, enhanced
by inclement weather at critical periods, with possible nutritional
deficiencies or imbalances contributing.
This general conclusion formed the basis for a greatly expanded research
effort, initiated in the spring of 1971, involving a team approach by personnel in the Departments of Pathology, Microbiology and Anatomy, College of
Veterinary Medicine, Colorado State University, and personnel of the Colorado
Divison of Game, Fish and Parks.
This investigation aimed at precise
identification of the factors or combination of factors responsible for a
high rate of mortality of lambs in certain bighorn sheep herds in the state.
A collection schedule, to take lambs from a herd known to have a high rate of
lamb mortality and from a herd known to have a high rate of lamb survival, at
two-week intervals through the summer and fall, was established.
Initially,
the Buffalo Peaks herd, located in approximately the geographic center of
the state, was selected as the herd having a high mortality rate. This
selection was based on work done three years previously by graduate student
Robert G. Streeter, and also on the fact that the terrain lends itself to
ease of collection.
The herd selected for the control was the Saguache CreekTrickle Mountain herd, located about 60 miles southwest of Buffalo Peaks.
Five animals were taken from the Buffalo Peaks herd, and five from the
Saguache Creek-Trickle Mountain herd.

�-195-

Harold Breen, Professor of Pathology (Bacteriology); Jerry Adcock, Professor
of Pathology; and Charles P. Hibler, Professor of Parasitology.
Considerable
assistance has come from George Post, Associate Professor of Microbiology.

�-196-

Table 1.

Bighorn sheep collected from the Buffalo Peaks herd.

I.ESIO~S
DATE

HT. (kr;)

SEX

6/16
6/29
7/27
8/10
8/23

12.5
9.0
20.0
38 . 6;'~
28.2

F

Viral
Parasitic
.~~--~~~--~~~~~

Racterial

Pro ros t:rongylus

F
F

Protostrongylus

F

:r-[

Protostrongylus

~':yearling

Table 2. Bighorn sheep collected from the Saguache Creek - Trickle
Mountain herd.

LESIONS
DATE

HT.

6/17
6/30
7/28
8/19
9/3
9/3

4.8
11.5
23.0
27.3
19.1
26.0

(kg)

SEX

Bacterial

Viral

Parasitic

F
F

N
F
F

Protostrongylus

N

Protostrongylus

�-197-

Table 3.

Bighorn sheep collected from the Sangre de Cristo herd.

DATE

HT.

8/26
8/28

12.7

8/30

49 .oli,.6
18.2

LESIONS':~"'"

9/2
9/2

(kg)

.,'~ yearling
,,&lt;-{,
All animals
pneumonia"

SEX

Bacterial

Viral

Parasitic
Protostrongylus
Protostrongylus
Protostrongylus
Protostrongylus
Protostrongylus

M
H
F
H
M

except yearling

Table 4.

Bighorn sheep collected

DATE

HTo

9/13
9/13
9/16
9/16
9/16

21.0

had the classical

"verminous

from the Pikes Peak herd.

LESIONS"':

19.0
17.3
14.0
18.2

(kg)

SEX
F
F
N
H

Ba c t e r i al

Viral

Parasitic
Protostrongylus
Protostrongylus
Protostrongylus
Protostrongylus
Protostrongylus

�-198-

sons OBSERVATIONS
"THE LUNGWORl'1-PNEUMONIA

ON THE EPIDEHIOLOGY

OF

COllLPLEX" IN BIGHORN SHEEP IN COLORlillO

Lamb mortality among bighorn sheep in some Colorado herds, often claims 90
to 100% of the animals born eVery year. The cause of this mortality is primarily pneumonia.
Ir.vestigato:cs c~:.:cocghout
the ·(,J2sterE
Uai::::ed
States and
Canada have spent their lifetir:,esar remp t t ng to d.etermine the disease agents
responsible for this die-off.
At the time rife ini tiated a study in Colorado
we had some of the facts; 'VIe also had many of th e fallacies to contend with
and overcome; in addition, we have enough umanswe red que srLons to frighten
away even the most seasoned. sciei.1.tific
investigators.
The lungworm, Protostrongylus stilesi, is said to be primarily responsible
for the pneumonia in bighorn sheep, and it may well "be; hcweve r , it may also
be just a part of the total picture.
The biological cy;::leof the lungworm is
extremely complex and since much of the problem revolves around it, we should
discuss this cycle, at least briefly.
Adult Lungwo rms liVe in the tissues or
air sacs in the lungs of their host an.lma l , Their life Sp::L1 is unknown, but
they probably live several years. Adult males and females mate, and the
females begin producing a multitude of eggs wh i ch they scatter about themselves in the lung tissue. These eggs develop and an extremely small larval
form, the "first stage larva", emerges.
This larva then makes its way into
the air passages, eventually crawling up the trachea and into the mouth.
The
larva is then s'Wal10wed, and goes through the gastro-intestinal tract, passing
out with the fecal pellets as the sheep defecates.
This 2.arva is unbelievably
resistant to envt ronment al. conditions.
Fecal pellets can be dried or frozen
for months and the larvae will all be alive, active, and ready to further
develop.
Before the larva is infective to the bighc:cn sheep, it must undergo
at least 40 to 60 days of development in a small land snail. Vallonia
pulchella, an ubiquitous snail in the Rocky Mountain Region, is one of the
most important species in Colorado.
This snail is rather tiny, about the size
of the head of a pin. \\flieD.
the f i rs t stage larva of the Lungwo rm penetrates
the foot of this snail, it begins a period of grovJth and development.
As it
grows the outer skin (cuticle) is shed; a process called "molting", 2nd. the
larva is then known as a "second stage larva". "With further growth and
development another molt occurs, ane. the larva is then a "third stage larva".
This third stage does not deve Lop further, for it is the stagevihich :'s
infective for the bighorn sheep. Once the: Larv a is infective, it remains in
the snail, living as long as the snail and snails g ene raIly survive about
two years.
Richard Pillmore, a biologist formerly with the Colorado Division of Game,
Fish and Parks was the first investigator to take the biological cycle
through the sna.iL, He vJas unable to put the ·worms back LOLe the sheep
because uninfected sheep were not available.
Dr. George· Post, an Associate
Professor at CSU, and Ruth Monson, a Cz aduar;e S·tuder~t"iiE211y succeeded in
completing the cycle. They fed infected snails to Mouflon-bighorn sheep

�-199-

crosses and recovered adult parasites 6 to 8 weeks later.
This finally
proved what appeared to be so obvious:
the bighorn sheep becomes infected
with lungworm after eating infected snails.
Yet several questions remained
unanswered.
Lambs frequently b ecame ill "with "ve rn.Lnous pneurr.onia" in midJuly, at an age of 6 to 8 weeks.
At this age they were far too young to
have acquired very many lungworms by eating snails because in mid-July
they are no t eating very much grass.
Mas t of us wo rk i.ng wi th the problem
believed that these lambs were born with their lungworms (transplacental
transmission) or acquired them shortly after birth (transmammary
transmission).
In the course of our investigation on the causes of excessive lamb mortality
among bighorn sheep in Colorado we showed that transplacental
transmission
of lungworm infective larvae does indeed o ccuz , a discovery which filled a
rather large gap in our know Le dg e about the "Lungwo rm-P'neumonLa Complex".
We found infective larvae in the placenta. foetal liver and foetal lungs,
which is about all the proof we need to make the statement that lambs are
born with Lungwo rm .
What we can say about the cycle from this po Lnt; on is theoretical, and will
require several years of hard work to prove, but ?roo£ is now within our
grasp.
Theoretically,
the pregnant ewe obtains "third stage (infective) larvae
with snails Which she accidentally ingests while grazing.
When these larvae
are released in her gastro-intestinal
tract, apparently they enter her circulatory system and cross the placenta, entering the foetal circulation.
The
first stop is the foetal liver.
Eventually,
they make their way to the foetal
lungs.
When the la~b is born, infective larvae are already present, ready
to continue their development.
In the lungs the infective larvae grow, develop,
and molt to the fourth stage.
They continue to develop. finally molting to
the fifth and last stage, the adult.
Adult parasites reach sexual maturity,
mate, and begin producing eggs.
First stage larvae begin to appear in the
feces 6 to 8 weeks later.
There are a number of reasons why this mode of transmission is extremely
important, and all of them are impor~ant to the management of a bighorn sheep
herd; consequently,
the problem must be thoroughly studied and all of its
implications evaluated.
When the ewe eets snails, and acquires infective larvae, some of these larvae
may be destined to cross the placenta and enter the foetus, and some may be
destined to enter her lungs and mature.
However, since she already has lungworm,
the odds are gooa that she is immune to a higher level of infection, especially
if she is healthy; thus, the majority of the larvae are probably destined to
cross the placenta and enter the foetds.
Another very good possibility
to be
considered is that Larvae acqua re d by the ecre i:::l the: course of grazing
throughout the year may go into her body tiSSUeS and become: dormant.
Here
they could accumulate, often in great ll.mabers, to be released during pregnancy yec..r after
year,
even \;Jitl-: !1.0 f;:;.rther Lnt ake of Larvae ,
7his is not
too rar-fetched, for the dog asca:cid hs.s an identical cycle.
The infective
larvae accumu Lat e in the bi t ch and a hormonal t.r
Lgge r ma cr.anf.sm s t Lmu Laz es
larvae to cross the placenta and ence r the foetal liver on about tl'ce 54th day
of gestation (abou t one vleek b ef ore b Lr ch) . They z emaf.n in the LLve r un t i.L
birth, tL2:'1. complete their cycle _ A "Di-;::d. can Lnf ect; "her pups year after year
with no further intake of larvae.

�-200-

Perhaps
one important
management problem which
that lungworm free herds cannot be established
snail-free
environment.

;-

.;l

.

•..
0 rm.na 1S
comes iIT~ediately
by p Lan t Lng Lambs raised in a

Another significant
implication
of transplacental
transmission
of lungworm is
the potential
problems which could. (and probably
do) develop on poor and
overgrazed
sheep ranges.
All par2sites
require
certain
factors
to insure
perpetuation
of their
own kind;
crowding and malnutrition
are the avo most
important.
During the pas t 50 to 70 years we have seen the bighorn
sheep range
become smaller
and smaller,
forcing
anir:;als :::'.0.1:0 a crmhlad s Lt uat Lcn , When
sheep are c rowd ed together
this means more snails
per dHi t area are being
infected,
furnishing
a better
chance for ewes to accumulate
g re ar e r numb
e r s of
infective
larvae
for their
offspring.
Cr'owd mg prorno t e s overgrazing,
and this
too means more snails
ingested
because
grasses
arc being rac r e closely
cropped.
If the Bighorn Sheep is on poor, ove rg r azed range,
.ir is undoubtedly
in a
marginal
or suboptimal
plane of nutrition.
Most likely
the lambs born of
these ewes are in the same condition.
The ewe gives birth
to a weak lamb,
and does not furnish
adequate milk for the l&amp;mb. Thus the lamb has several
strikes
ag a i.nst it at: birth.
HovY iOes the p.ar as f t.e fit into
eLis picture?
If ewes are crowded and in an overgrazed
area,
they will be passing
a greater
numbe r of 13:L-v",e to the foetus"
If they are in a poor p Lane of nutrition,
the number may he even greater.
For ex ampLe , a healthy
animal can "throw
off" heavy infections
of parasites,
but a ewe in poor co nd Lt.Lc n probably
loses
some of her in~~unity.
Some of the parasites
she might have thrown off
probably
develop in her,
pulling
her down further,
and a greater
number may
cross
the placenta.
When these factors
are all put: together,
and the parasite
added as the final
ingredient,
the result
is that the lambs are born
predestined
to die.
How and when do the lambs die?
Tom Woodard (MS Thesis,
CSU) studied
the
die-off
in the Sangre de Cristo
herd f o r two years.
Tom observed
that lambs
begin coughing about the middle of July (6 to 8 weeks after
b i r th) wh i.ch
coincides
perfectly
with the maturation
of the lungwo:cms.
Also,
the summer
rains
begin in the high country
at thE: same ·time.
This added environmental
s tress
is extremely
important&gt;
for it seems co "kf ck-o ff " the p ne'cmon i a .
Field biologists
have observed
that the ill animals
(those
destined LO di.e ? )
are light
in \-leight,
Lf.ght in co Lo r , \weak; "snoety-nosed"
aed ::iave f r equ en t;
paroxysms of cough i.ng ; t.he ryp i c a.l "1,LT'.g,,;rOI.-m-PneClmonia'"
synd rcme • By ·che
time summer' r a i.ns have ended rh es e Lamb s have seV2:;:-2. pneumcn i.a ,
Ir~. Augcst
or early
September lamb mortality
is essentially
complete.
given us some answers,
b'Jt
The discovery
of transplacental
transmission
h'
we do not by any means feel
that we can say unequivocally
that; we have them
all.
It is important
to understand
that several
agents have been blamed as
(or suggested)
causes of mor t a.Lf ty.
First,
the lUc,g\JOZIn is quire
capable or
causing pneumonia.
Second,
the Lungworm may furnish
an avenue for pathogenic
bacteria
to become es t ab Lt she d L. lung tiSS1':c.
Tliird,'::
virus may be a
factor.
~Jirh all these pc s s Lbl I i r i es now in mind, let us orice again exp Lore
lamb mor t a Lf ty .
If ewes in a poor plane of nutrition
givE:' -bL~th to unhealthy
lambs ,.j:hicll have Lungworm 'Alhen they are bo rri , and the Lamo s are s tressed
by
summe r rains
about the time they have developed
a "verminous
pneumon i a'",
t.he s e ~,reakc:ccedanimals are then h::ig1:-iljsucep t tb Le to invas::'on by pa·chogenic
bacteria
which are frequently
found in the lungs of ·0ighorn sheep.
A species

�-201-

of Pasteurella is the primary pathogen involved.
It is not unusual for us
to find a bighorn sheep lamb showing pnaumond c lesions prLla:cily of bacterial
origin and secondarily of Lungworm origin; however, the ill Lamb next to it
may have lesions primarily of lungworm origin and secondarily of bacterial
origin.
The lambs collected from the Pikes Peak and Sangre de Cristo herds
during 1971 had pneumonic lesions attributable primarily -co lungworm.
Last, but not least by any means) Dr. John Parks, a virologist who is a
member of the research tea~ has isolated a virus called Parainfluenza-3.
This virus is quite capable of causing all the problems we have seen, without
the help of bacteria and/or lungworm. }'lanyhave attempted viral isolation,
but Dr. Parks of CSU was the first to succeed.
Thus far no lesions in the
lungs attributable to a virus has been seen; however, these lesions could be
masked by lungworm and/or bacterial i:;}ctuced
lesions.
Viral titers were found
in ill lambs examined from the Pikes Peak herd in 197:. ':£'"'18 significance of
the virus, and its role in lamb mortality must be carefully considered this
year and in coming years. We need to have blood and nasal swabs from wi:d
bighorn sheep to determine how many have titers against PI-3. We also need
to experimentally infect lambs with this virus, and we have plans to do so
this year.
One other aspect of the transplacental transmission of Lungwo rm and the
isolation of PI-3 virus which bothers all of us is the knmvledge rha t in.
other animals (swine, for exmnple) viral agents have been transmitted by
Lungwo rms (swine cholera and Lungco rm) . The possibili ty that this situation
exists must be f&lt;llly explored.
This last year we acquired the ability to
raise snails in the laboratory and infect these snails with lungworm larvae.
Dr. Parks can isolate and culture the virus if one is presen t; in the lungworm
larva. This coming year we will attempt to LsoLa'ce a virus from the third
stage (infective) larva. If we succeed, then our problem has truly become
complex:
This is the problem as we see it. What caused the problem?
Nutrition is
probably the answe r . Disease kills the lambs, but if these lambs were
healthier most of them would have survived the diseases wh i ch they faced
early in life.
Where do we go from here?
fraught

with

The road is Long and rough; moreover,

tr... dang er s of public.

opi~c:..iono..f:d p re s sur e .

·2

Lots

of

it is
t Lrae ,

effort and quite a little h aar tbreak were ::eqc:'r2dto gec us this far) and
we could not have done so without: the effort of many hard\vorking men and
women. We are not going to solve the question of bighorn sheep lrunbmoYtali t.y this year, nor will sse have the answers nexc year, but we will make
h eadway this year, and. next year--eveDtually we w'ill ob t aLn those answers.
There 2.r2 a l1.&lt;.L.nber
of aspects to be Lnve stigated &gt; and :?riorit Les to be
assigne~; thus far this has not been done and I can only give you a list
at present.
1.

Collect more pregnant
t r s.nsmt ss Lon of

ewes and study the tr2.nsplacental

luri.gT~or~

0&gt;

�-202-

2.

Study the transplacental transmission
tame herd at Meeker, Colorado.

3.

Attempt transplacental transmission
sheep (currently underway).

4.

Experimentally infect bighorn
to produce pneumonia.

5.

Experimentally infect bighorn sheep vlith PI-3 virus and
attempt to produce the pnelliilonia.

6.

Experimentally infect bighorn sheep Hith the species of
Pasteurella said to cause pneumonia.

7.

Experimentally infect b i.ghorn sheep wi th a comb i.nat.Lon of
parasites, bacteria and virus.

8.

Develop a treatment for lungvlorm in bighorn sheep. A product
which will eliminate both zhe adults and larvae is required.

9.

Develop a vaccine for PI-3 virus
be responsible for mortality.

of lungworm using the

of Lungworm

to domestic

sheep with lungworm

:i.f

and attempt

this agen t is shown to

More could probably be added to this lis t and no doubt \.Jillbe. Some of
this research is basic, some applied, and throughout, of course, there are
academic aspects to it. Eventually, however, everything we do must be aimed
at management.
How do we get this done'? We use a research team composed of the specialists
at the universities, game biologists and game management personnel.
This is
to insure that our goals are directed at production of bighorn sheep and not
research for the sake of research.
This research has progressed satisfactorily, and has accOll1plished the things
we talked about because of teamwork by the following personnel (listed in
alphabetical order);
Dr. Jerry Adcock, Professor of Pathology at CSU; George
Bear, Wildlife Researcher with Game and Fish; Dr. Harold Breen, Professor of
Pathology at CSU: Dr. Charles Hibler, Professor of Parasitology at CSU;
Robert Ke Lss , Wildlife Researcher \vith Game arid Fish; Robert Lange, Graduate
Student in Parasitology at CSU; Carol Metzger, Research Assistant in Parasitology at CSU; Dr. John Parks, Assistant Professor of Virology at CSU;
Dr. George Post, Associate Professor of Microbiology at CSU; William
Rutherford, vJildlife Researcher TN'ithGame and Fish; Robert Schmidt, Wildlife
Researcher li7ith Game and Fish; Gene Schoonveld, Wildlife Researcher with
Game and Fish; Dr. Gordon Solomon, Associate Professor of Pathology at CSU;
and Tom Woodard, Wildlife Researcher wi th Game and Fish ..

�-203-

ISOLATION

OF PARAINFLUENZA

J. B. Parks,

D.V.M.,

Department
Biomedical

Ph.D.;

3 VIRUS

George Post,

of Microbiology,

Sc.f.erice s , Colorado
and

the

FROH

B.S.,

College

State

1Alyoming

ROCZY

M.S., Ph.D.;

of Veterinary

University,

G-.31i--:c

BIGHOR..."N'
SHEEP

HGUj:~IAIN

Fort

and Ef.sh

Tom Thorne,

Hedicine

Collins,

D.V.M.

and

Colorado

80521

D2pa.rti.TIC1.1.i:

Parainfluenza
3 (PI-3) virus -itJ2Sisolated
from nas al. "(.yashings of 3 of 10 Rocky
Mountain bighorn sheep (Ovis c anade ns Ls ) a-t the Sybille
Big Came Research Unit
near
am.i
wyomiEg.
-':-;:1e
tiTlE:
t:-"C 2,-,:::'::12ls
had been in captivity for 1 month.
Several
animals were febrile
and exhibite&lt;i
clinical
signs
of a respiratory
disease.
Hemagg Lu t i nat Lori inhibition
(HI) titers
to PI-3
virus
increased
markedly during the conf Lneraerrt period
in 9 of the 10 animals.
All 10 of the bighorn sheep eventually
died.
Lar

e

,

At

of

s

amp

l

rng

,

INTRODUCTION

7
The bighorn sheep pneumonia complex ,22
is suspected
of being che primary
cause for a decline
in the populacion
of bighorn sheep and many diseasecausing organisEs
have been incriminated
as etiological
agents.
The lungworm,
Protostrongylus
stilesi,was
initially
i",;;&lt;Licated as a:" irritant
factor
predisposing
sheep to bacterial
infections8•17&gt;
20.
The bacterid
mosr ofte?
i~oIated from the respiratory
tracts
of bighorn sheep were CorYLebacterium
,1. ,
? 21
'D' i'lploco.:::cuS-»-'.
-=&lt;?C ,
-..
P
j
prllT,ary
p
i ty h_as not
been direc.r.ly at t.r Ibut ab Le "LO t.h em, 1~vco~Jla;sI!1c. .:=_::cgi~'"'.iir!.:_. was isolated
f rom
pne~lli110ni~ ~·u.ngs of ·Digl~_QY-;"1.
S:1.22P i!.1d.i2a·:ir~g
"t}-lC"tt~is
o=:ga:"i.:'sI:.: !nay be
pathogenlc
.
1 -

ast

eure

1

La-&gt;-

-

and

but

a

tnogena

c

a

rway

s

With the lack of ag re emeri c as to ch e ac t.ua.I cause of the pneuraonLa complex,
viruses
we r e sugges ted as a possibility.
At =-'east: 4 viral
agents have been
associ~ted
with biZhorn sheep,
ei~her in _~apt,i:ity
or ~n their _natural
habitat.
Con t ag i ous e cthyma " \oJas detected
In Canad i.ar, b Lgho rn srieep , but: has not been
2b.-wa~s
Imp
Lf, ca t.•..e d
",'Q ~. caus az(....L
i ve ~b!
az en t'- __
-i r-,
-;-"\--_c
-;~
R1-·,pto'-,o-ue
.1
U
~......,
L..
'-~L'"-.• '''''~'''-''T''~''
.!:-' s: _c....:;._ :.U.l._ ....•....
.......;.r-~-.wn
p ~ "-""'l....
..•....'-.1.-0
shown to be the cause of death in a young desert
bighorn
ram by inoculation
of
a suspension
of its lung -[issue Lito a susceptible
dorae s t f,c ::'amb in VJ~ich it
caus e d typical
bluetongue
symp t oms and s t imu La t ed tl-:e prod-:.;;ction of bluetongue
ant.Lbod Le s .
Epizootic
hemorrhagic
d Ls eas e (ERD) was diagrcosed in a bighorn
s haap at -tr:.2 San Di2g0 zoo on tIl2 b as i s of e:.lill.:'cal S::.g:'1S._ 2T!C pathological c
lesions
during an outbreak
of ERD involving
7 sp e ci es oJ:: exotic
ruminants..).
Finally
P~-3 virus has been incriminated
becaus~ of serological
eVidence14.
The virus has never been isolated,
bUL several
negative
:e.ttempts have been
-~r
'8.:L
'&lt;'1--.-',
-'n""'"
.:l~-.~~-=-,~",
.,.1-·,
-',-. l •. ---;nPT_'"
f ror
Rocky
reporLea.
.1..
.•.
l.-LS pC-pc
.•... Uc:0l.-~...!....LH::::v
•.....
.:.._c:~ 1.:::.O_Ci.._..;...UL!
O.f .l.
..2.
. ..) '"_._-""
v.~l.u::;:'
..
rvlll
K '-_
'-"iJ •.••

Co.

'j --

Hount.a.Ln

-..

bigl"lo:cn

sheep

and discEsses:::he

s Lgnf f i.cance

of

that

isolation.

�-204-

Y.•.
ATERIALS AND r-ffiTHODS

Animals
Ten Rocky Mountain bighorn sheep were taken into captivity from their
natural habitat.
The animals ranged in age from 7 to 8 months to 2 years.
The animals had not coming led with domestic animals on their ranges. During
their captivity period the entire group of 10 bighorn sheep was penned
together wi th no other animals present.
Hov;rever,domestic sheep, elk, mule
deer and pronghorn antelope could have ranged up to the pen and were
separated iroTI the bighorn sheep only by a stockade fence.
Tissue Culture
Primary cell cultures of domestic lamb lung were pre-prepared from trypsinized
lung cell suspensions and grown in minilT::C:Hl
essential medium (Jvf-EM)
containing
10% fetal calf SerlliTI
(FCS) as descriced by Madin 15
Isolation

Technique

Nasal washings 'were accomplished by irriga Ling rhe nasal cavi ties wi th 8 ml
of MEM and collecting the effluence wi.th a funnel and tube combination.
Samples were taken one month after the date of c pture.
Tubes were immediately frozen at -70 C until tissue cultures could be inoculated.
Samples
were thawed, clarified at 1,000 g for 10 min. at 4 C and 0.1 ml from each
sample was inoculated onto each of S Lubes of secondary lamb lung cell
cultures.
Incubation was at 37 C in a stationary position.
Tubes were
examined daily for evidence of cytopathic effects (CPE). Cultures were
maintained with MEM containing 2% FCS and antibiotics,
Identificat:Lon
.

1

Identification of viral agents causing CPE was accomplished by HI and serum
neutralizati9n19
using specific ant.ise cum prepared in rabbits against the
2J
SF-4 strain
of PI-3 virusa.
An iserum was produced by inoculating 107
TCIDSO I s of PI-3 virus. intravenously tHice 2 week for 3 weel,:sand waiting
2 weeks after the fiEal injection to collect the hype:ci::::mune
serum.
Serology
Detection of PI-3 viral an t.Lbo d.ies in t.he bighorn sheep was accomplished
by HII. Serum was obtained on the date of viral sampling. 2 weeks before
sampling and L;weeks prior to samp Li.ng 'V;rhich
coincided with the date of
capture.

aStrain SF-4 of PI-3 virus was obtained from Dr. A. Bo
Microbiology, Colorado State University.

Roerlein,

Dept. of

�-205-

?2SDD:'S

The 'results
obtained
in this iE\f8stigBcioCl, a:ce SUlTi£,:a:ciZed
if, 'I'abI,e 1.
Parainfluenza
3 virus was recovered
from 3 of tree Ie bighorn sheep sampled.
Each animal had relatively
low HI titers
to PI-3 at the t Lme of capture.
At
the second serum sampling,
2 weeks later,
titers
had risen in 5 of the animals
while declining
in 1.
The third serlli~ sample taken coincideuLal
with viral
sampling showed marked elevations
of titers
in 9 of the bighorn sheep and a
fall
in titer
in 1 individual.
Body temperatures
were recorded for each bighorn sheep on the day of viral
sampling.
Sever21 2ni-;Eals were febrile
and many
showed clinical
signs of respiratory
disease.
Nasal 'W'&amp;shingsof some animals
were filled
with thick muco-purulent
nasal exudaces.
Coughing and dyspnea
was

COTIl.lT •.on ~

The first
mortality
occurred
2 days aiLer viral sampling and all 10 bighorns
died within a f.ew weeks . Post-mortem investigatiOlLs
have not been completed,
but gross examinations
revealed
pneumonia present
in ell.

DIscussrm:;
The first
isolation
of PI-3 virus from Rocky Hountain b Lgho r n sheep reported
herein substantiates
previous
yeports indicating
se~ological
evidence of PI-3
viral
infections
in this species14" 18.
Negative attempts
to isolate
PI-3
virus from bighorn shesp have bee-C!,T2portec.8 ,14, but the authors qualified
their data by s t at.Lng thE..z; ·~iSSt:2S used for iSQls..t~Jll a-ct.2T.:'p"CS "had been
stored at -IS C for several
months befoO:E:tes-cingS and t:la~- paramyxoviruses
at that 'temperature 'will lose infectivity
in a £e':J days.
He also encountered
negative
results
in virel
isolation
attempts
from r-ume rcus s amoLe s collected
in the field ~,]ithout proper handling proc~dur2s18.
The posit:!.:~e results
obtained
in 6_i8 study were probably faeilita'ted
by the i'irillediate freezing
of nasal washings at -70 C, a terr:perature at which paralllyxoviruses
may survive for Several months and to the fo r t.unat.e 'ciming of sample collection
whi.ch happened to be eluring the -viral shedding s -cage of cliIi.ical
p::lelli~cmia.
Parainfluenza
3 infection
is rather
transienc
and tbe virus can be recovered
for only a few days following
experiraental
inocula'.::io'il16.
This could explein
the failure
to isolate
the virus
from more ':han 3 of the i0 animals.
Serum vlas collected
from all 10 bigho:cn she ep at t.he t Lme of their
capture.
Four had HI t Lt.e r s to PI-3 virus,
which indica'Ces thatc'c,e
t.nf ec t t on was a
nat.uz a I one; occ'urring Ln the w:iId ar40. IJ.Ot fY"OTI.l adjac2r.l.t
a:Liyaals 'vihi2-e in
captivity.
After 1 month in captivity,
all 10 bighorn steep had HI Liters
to PI-3 virus and the titers
were elevated
in 9 of the animals.
One interpretation
is -:::hc:.tthe clOSE: conf t.nemenc precipitated.
the spr e.ad of PI-3
virus
f r om those rha t were infected
in their
natural
h&amp;bitat
to the remainder
of the herd while in cc:.ptivi-cy and that the PI-3 virus,
isolated
from 3 of
the animals,
could have been instrumental
in the ells-uir:g dea ths of all 10
bighorn

sheep.

�-206-

Table l. Parainfluenza
bighorn sheep.

3 isolation

results

arid HI titers

in captive

Body Temp.
Day 29

Isolation

Day 29

Day of
Death

PI-3
HI AntiboG:Y Titers
Day 0* Day IL, Day 29

Animal
Number

Age

1

1

1:32

1:16

1:16

105.0

pas.

84

2

1

1:16

1:32

1:64

103.1

neg.

82

3

lamb

0

0

1:4

105.0

pas.

48

4

lamb

0

0

l:8

102.3

pos.

31

5

1

1:8

1:32

1:64

103.6

neg ,

43

6

1

0

1:8

1:32

105.4

neg.

44

7

2

0

0

1:4

103.1

ne"'.
c»

44

8

2

0

1:4

1:32

105.0

neg.

73

9

lamb

0

0

1:8

105.8

neg.

35

10

1

1:4

1:8

1:64

102.5

neg.

44

*

Days

from date of capture.

�-207-

Previous serological studies indicated that bighorn sheep had been in contact
with PI-3 virus.
The primary isolation of PI-3 virus repo rred he rei.n shows
that they are definitely capable of supporting infection with the agent.
Parainfluenza-3 virus has been isolated from domestic sheeplO.12 and has
been shown to cause experimental virus pneumonia in domestic lambsl1.
The
next question to be answered is, "'(AThat
is the role of PI-3 virus in the
pneumonis. complex of bighorn sheep'?"·

ACKNOWLEDGMENTS

The authors would like to thank Mr. Peter Nash of the Depa:ctment of
Microbiology, Colorado State university, and Hr. Floyd Blunt of the
Wyoming Game and Fish Department for their assistance in obtaining samples
for this work.
Support for this research was funded by the Colorado
Division of Game, Fish and Parks.

�-208-

REFEP..ENCES

1. Abianti, F. R., A. B. Hoerlein, R. L. Watson, and R. J. Huebner.
1961.
Serological studies of iliy:w']iruspara-influenza
3 -virus infection
in cattle and the prevalence of antibodies in bovines.
J. Immunology
86: 505-511
2.

Al-Aubaidi, J. M., ~L D. Taylor, G. R. Bub ash , and A. H. ;}arcliri.
1972.
Ider~Lification and characterization of ~':y'-&gt;'::IL)lClS;Ul
Clrgillini
AJVR 33: 87-90.
from bighorn sheep (Ov:L~ canadens" s) and goats.

3.

Bingham, D. A. 1962.
Study of paste-c:::-,e:llt;sis in bighorn sheep.
Fed. Aid Div., Quarr. Rept., Wyo. Dept. of Game and Fish. p. 20.

4.

Blood, D. A. 1971.
Contagious ecthyma in Rocky Mountain
sheep.
J. Wildl. Mgmt. 35: 270-275.

5.

Griner, L. A. 1970. Hemorrhagic
zoo. JAvl 1A 157: 600-603.

disease

bighorn

in exotic ruminants

in a

v

6.

Forrester, D. J. 1964. A survey of lungworm infection
sheep of Montana.
J. Wildl. Mgmt. 28: 481.

in bighorn

7.

1971. Bighorn Sheep Lungworm-Pneumonia
Complex.
In Parasitic
Disease of Wild Animals.
Edited by J. W. Davis and R. C. Anderson.
The Iowa State University Press, Ames: 158-173.

8.

, and E. M. Wada.
1967. An attempt to isolate viruses from lung
tissue and lung nematodes of bighorn sheep.
Bull. Wildlife
Disease Assn. 3: 74-76.

9.

Hadlow, W. J., and W. L. Jellison.
1962.
bighorn sheep.
JAVMA 141: 243.

Amyloidosis

10.

Hore, D. E. 1966.
Isolation of Ovine Strain of Parainfluenza
Serologically Related to Type 3. Vet. Rec. 79: 466.

Virus

11.

, and R. G. Stevenson.
1967.
Lambs.
Vet. Rec. 80: 26-27.

12.

1968.
Isolation of PI-3 viruses from the Lungs and Nasal
Passages of Sheep showing Respiratory Disease.
J. Compo Path.
78: 259.

13.

Experimental

in Rocky Mountain

Virus Pneumonia

Howe, D. L. 1964. Etiology of pneumonia in bighorn
Div. Quart. Rept., Wyo. Dept. of Game and Fish.

sheep.
p. 68.

in

Fed. Aid.

�-209-

14.

Howe, D. L., G. T. Woods, and G. Narquis.
1966. Infection of bighorn sheep (Ovis canadensis) wit:h Jv1yxovirusparainfluenza-3 and
other respiratory viruses.
Results of Serologic tests and
culture of nasal swabs and lung tissue.
BulL Wildlife Disease
Assoc. 2: 34-37.

15.

Madin, S. H., P. C. Andriese, and N. B. Darby.
1957. The in vitro
cultivation of tissues of domestic and lab animals.
AJVR 18: 932.

16.

Marshall, R. G., and G. H. Frank.
1971. Neutralizing antibody in
serum and nasal secretions of calves exposed to parainfluenza-3.

17.

Marsh, H.

18.

Parks, J. B.

19.

Paul, J. R., and J. L. Melnick.
and Rickettsial diseases.
New York, N. Y. 53 p.

1938. Pneumonia
19: 214-219.
1971.

in Rocky Mountain

Unpublished

bighorn

sheep.

J. Ma~TIal.

data.

1956. Diagnostic procedures for Virus
2nd. ed. Am. Public Health Assoc. Press,

20. Pillmore, R. E.

1961. Study of lung nematodes of bighorn aheep ,
Aid Div. Quart. Rept., Colo. Dept. of Gmae and Fish, p. 69.

Fed.

21.

Post, G., and K. B. Winter.
1967. Life cycle of lungworms.
Div. Quart. Rept., Wyo. Dept. of G@me and Fish. p. 68.

22.

Post, G. 1971. The Pneumonia Comp Lex in bighorn
Am. Wild Sheep Conf.
1: 98-102.

23.

1959. A Myxovirus (SF-4)
Reisinger, R., K. Heddleston, and C. Manthei.
JAVMA 135: 141-152.
associated with Shipping Fever or cattle.

24.

Robinson, R. M. 1967. Bluetongue
Wildlife Mgmt.
31: 165-168.

Fed. Aid

sheep. Trans. North

in the desert "bighorn sheep.

J. of

��January,

-211-

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~--------

Project No.

W-105-R-13

Work Plan No.

4

Job Title

1

Job No.
Population

Period Covered:
Personnel:

An Ecological Investigation
of the Cache la Poudre Deer Herd

Density and Structure

March 1, 1972 to February

28, 1973

Allen E. Anderson.

ABSTRACT

The population density portion of this job has been published (Anderson et
al. 1972 J. Wildlife Mgmt. 36(2):571-578).
The population structure data
were partially analyzed and the literature reviewed.
Tentatively, this
material will be incorporated into a Division Special Report •

...... ..:.,." .
~.
Prepared by: r.A.--I.-f-,;:;}"~ 'i.::; //.. ;:-'~/{I.';, .1('7''--'
Allen E. Anderson
Wildlife Researcher
-~

.'

1973

��January,

-213-

JOB PROGRESS REPORT

State of

COLORADO
------~~~~----------

Project No.

Job Title

Physical

Period Covered:

March

1

Job No.

5

Work Plan No.

Personnel:

An Ecological Investigation of
the Cache la Poudre Deer Herd

W-105-R-13

Characteristics

1, 1972 to February

28, 1973

Allen E. Anderson.

ABSTRACT

The first draft of a long manuscript on the growth and morphometry of the
carcass, and selected organs and glands of mule deer was about 3/4 complete
as the segment ended.
It is planned that the final version will be summitted to Wildlife Monographs.

Prepared by:

/"

:"'"

,_( ( t, r;. C." {, ,:
Allen E. Anderson
Wildlife Researcher
('.'

/,',,~
?',r(

/

,: :.

('/

,
••

1973

��January,

-215-

JOB PROGRESS REPORT

State of

COLORADO
----~~~~~----------

Project No.

W-105-R-13

Work Plan No.

5

Job Title
March

2

Job No.
Reproductive

Period Covered:
Personnel:

An Ecological Investigation
of the Cache la Poudre Deer Herd

Studies

1, 1972 to February

28, 1973

Allen E. Anderson.

ABSTRACT

Histological studies of ovarian material begun during 1965 by Dr. Schoultz
of the University of Colorado Medical School were continued by Dr. Bruce
Criley
of Illinois Wesleyan University, Bloomington, Illinois.
Morphometric
studies on the reproductive tract of both sexes were in progress as the
segment ended.
The morphometry of the reproductive organs will be included
in the Work plan 5, Job 1 (Physical Characteristics) manuscript to be
submitted to Wildlife Monographs during Segment 14.

Prepared by:

(/{{;;t:.~~.
~-?~(;~~.&lt;r,"(:;1"/"
Allen E. Anderson
Wildlife Researcher

7\

;'

1973

��January,

-217-

JOB PROGRESS REPORT

State of

COLORADO

Project No~

w-105-R-13

Work Plan No.

5

An Ecological Investigation
the Cache la Poudre Deer Herd

Job Title
Period

Harvest Analysis

Covered:

Personnel:

3

Job No.

March

1, 1972 to February

28, 1973

Allen E. Anderson.

ABSTRACT

All analyses were virtually completed •. The current literature was
reviewed.
Tentatively, this material will be incorporated into a
Division Special Report.

Prepared by:

/~,
( ••

-{

for

·c",.? ....,_.

r::

(.".

I

,~-,

L. ~ t 'rt (~'',.{/-'-4;·-Z __/

Allen E. Anderson
Wildlife Researcher

of

1973

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                  <text>-155-

April, 1973

JOB FINAL REPORT

State of

COLORADO
------~~~~~-----------

Project No.

W-37-R-26

Work Plan No.

1

Job Title:

Job No.

16

pheasant Nest Site Selection Study

Period Covered:
Personnel:

Game Bird Survey

April 1, 1972 through March 31, 1973

Many individuals have contributed to the completion of this study.
A complete listing is found in Hoffman (1973b).

ABSTRACT

A publication covering results of this study was published as follows:
Hoffman, D.M. 1973b
pheasant nest site selection.
Colorado Division of Wildlife, Denver. 27 p.
o

Special Report No. 32.

��-157-

PHEASANT

NEST SITE SELECTION

i
STUDY

Donald M. Hoffman

P. S. OBJECTIVES
During the early years of the study (1964-1966), objectives were to establish
the necessary facilities and plant selected species.
During 1967 and 1968,
objectives were to compare pheasant nesting use of, and success in (1) winter
wheat, (2) alfalfa, (3) crested wheatgrass, (4) hairy vetch, (5) white sweet
clover, and (6) alfalfa-crested wheatgrass mixture.
Following the 1968 nesting season, change over of three types (winter wheat,
hairy vetch, and white sweet clover), which were used very little for nesting
cover, or not practical to maintain along roadsides, was started, and three
new cover types (smooth bromegrass, intermediate wheatgrass, and tall wheatgrass) were establi~hed.
Objectives for 1969 through 1971 therefore included
comparing nesting use of, and success in these three additional cover types,
plus the older cover types.

RESULTS AND DISCUSSION
The objectives of this study have been completed, and manuscript written, and
results were published in the Division's Special Report series as follows:
Hoffman, D. M. 1973b.
Pheasant nest site selection.
Colorado Division of Wildlife, Denver.
27 p.

Prepared

by --",tu"",-:-~",'L' ~""""''-!I!Q._·__?r/--::-:~'::'~--L~MI---C~
~Hoffman
Wildlife Researcher

_

\

\

Special

Report No. 32.

��April,

-159-

1973

JOB PROGRESS REPORT

State of

~C~O=L~ORA~D~O~

Project No.

W-37-R-26

Work Plan No.

1

Job Title

.Game Bird Survey

April

18

Job No.
Pheasant

Period Covered:
Personnel:

_

Roadside

Cover Evaluation

study

1, 1972 through March 31, 1973

Thomas Lederhos,

Carroll Grand Pre and Warren

Snyder.

ABSTRACT

Pheasant nest searches
of 76 seeded plots, 42 farmed controls and 42 unfarmed
natural roadsides yielded 58 nesting attempts and 16 successful nests in 1972.
Nest density per acre on seeded stands was significantly greater, and approximately twice that found on either of the controls.
Combined data for the
three years of study produced similar results, except that in 1970 and 1971,
roadsides farmed to the shoulder contained significantly fewer nests than
either of the other cover types.
Predators continued to destroy over half
the nests. Approximately 24 percent of the nests hatched successfully over
the three year period.
Nesting success increased as the season progressed
from spring to summer.
New growth and residual cover, remaining over winter,
were both rated good to excellent in 1972, and provided better nesting cover
than in previous years.
Spring, 1972 applications of nitrogen fertilizer
greatly increased growth of old stands of intermediate wheat grass and smooth
brome, but had little noticeable effect on new seeded stands.

��-161-

PHEASANT

ROADSIDE

COVER EVALUATION

Warren

STUDY

D. Snyder

P. S. OBJECTIVE
To compare pheasant production under natural roadside conditions with the
following cover types to be established along roadsides: (1) grass, and (2)
grass-legume mixtures.

SEGMENT OBJECTIVES
1.

To measure

pheasant

production

2.

To measure

life-form

characteristics

METHODS

in test and control plots.
of roadside

cover.

AND MATERIALS

Reference is made to Snyder (1969, 1970, 1971 and 1972) concerning methods
and materials used in this study.
A supplement to these is added as follows:
Dry pelletized 33 percent nitrogen fertilizer was broadcast onto linear
segments of a number of new and old seeded roadside stands in March 1972. A
cyclone seeder was used to apply the nitrogen at approximat~ly 110 pounds per
acre, bulk rate, or 35 pounds of active ingredient per acre. Both ends of
the test strips within the seeded plots were marked for later inspection and
comparison of growth characteristics.

RESULTS AND DISCUSSION
Measurement
Nest Density

of Pheasant

Production

Comparisons

Seeded roadsides contained one nest per 1.06 acres of 39 nests on 76 quarter
mile plots during the summer of 1972 (Table 1). This compared with one nest
per 2.37 acres on 42 natural unfarmed (e2) plots and one nest per 2.29 acres
on farmed (Cl) roadside control plots.
Nest establishment on seeded plots
was significantly greater than on either of the control acreages (Chi-square
= 4.92*, 1 d.f.). No significant difference was detected in nesting attempts
between natural and farmed control acreages.
Combined nest search results for 1970, 1971 and 1972 are illustrated in Table
2. Seeded plots showed nest densities that were greater than those on farmed
controls with high significance (Chi-square = 24.61**, 1 d. f.). The

�-162-

difference in nest densities between seeded and natural roadsides was also
highly significant (Chi square = 8.23**, 1 d. f.). Natural controls yielded
a Significantly higher nest density than farmed controls over the three year
period (Chi square = 4.42*, 1 d. f.) even though farmed controls contained
slightly higher nest densities in 1972 (Table 1). An abundant growth of
downy brome (Bromus tectorum) potentially attracted nesting hens to farmed
roadsides in 1972.

Table 1. Densities of nesting attempts and successful nests in roadside
study plots in 1972.
Number
of Plots

Total
Acres

x Plot
Size

Number
of Nests

Seeded

76

41.52

0.546

39

1.06

9

4.61

Un farmed Contno1

42

21.30

0.516

9

2.37

4

5.32

Farmed Control

42

22.94

0.546

10

2.29

3

7.65

Cover Type

Acres/ Success. Acres/
Nest
Nests
Sue. Nest

Chi square Values
Seeded vs. farmed

4.92*

Seeded vs , un farmed = 4.92*
Farmed vs. unfarmed = 0.004

A slight increase in nest density on the seeded plots has been indicated each
year. In 1970 there were 1.21 acres per nest. In 1971 the figure was 1.12
acres per nest and in 1972 the average was 1.06 acres per nest on seeded plots.
On all plots (seeded and controls) one nest was found per 1.86, per 1.69, and
per 1.48 acres in 1970, 1971 and 1972, respectively. Whether these results
can be attributed to increased attractiveness of the plots for nesting hens,
increased pheasant populations, or increased efficiency in nest search,
remains unknown.
Higher nesting success tended to compensate for fewer nests in the unfarmed
natural control plots when compared with nesting in the seeded plots (Table 2).
Nine of 25 nests, or 36 percent, were successful in tqe unfarmed controls,
whereas approximately'21 percent were successful in the seeded and farmed
control plots. Therefore, on the basis of acres per successful nest, natural
controls were nearly as productive as seeded plots.

�-163-

Table 2. Densities of nesting attempts and succ~ssiul nests in roadside
study plots during three years, 1970, 1971, and 1972 combined.
Number of
Plots

Total
Acres

Nesting
Attempts

Acres/
Nest

Successful
Nest

Acres /Succeas f

Seeded

191

104.09

93

1.12

20

5.20

Unfarmed Control

103

52.84

25

2.11

9

5.87

Farmed Control

108

58.87

14

4.20

3

19.62

Cover Type

Nest

Chi square Values
Comparisons

Nesting Attempts

Successful Nests

Seeded vs. Farmed Controls

24.61**

5.31*

Seeded vs. Unfarmed Controls

8.23**

0.09

Farmed vs. Unfarmed Controls

4.42*

3.68

Chi square values indicate that seeded roadsides produced significantly more
successful nests per acre than did farmed controls (Chi square = 5.31, 1 d.f.:
There was no difference between seeded and unfarmed controls, nor was the
difference between farmed and unfarmed controls significant (Table 2).
Fate of Pheasant Nests Along Roadsides
Nesting Success--Table 3 itemizes nest fate in'all plots for the three years
of nest search and provides an overall average. Predation destroyed about
half or more of the nests each year. Some, probably most, of the unknown
fate could be added to this category.
Success of about one-fourth of the nesting attempts is evident. How does
this hatching success compare with information from other similar studies?
In general, it is about the same, In Illinois, Joselyn (1970) found, on the
average, approximately 27 to 29 percent of pheasant nests hatched successfully
during the years 1963 to 1970. A slightly lower percent, 23-28 percent, were
abandoned, and between 40 and 50 percent were destroyed by predators. In
South Dakota, Trautman (1960) recorded 20 and 28.8 percent hatching success
along roadsides in 1958 and 1959, respectively. In Nebraska 19.3 percent of
the roadside nests succeeded (range 11.8 to 45.4 percent) compared to a 24.8
percent mean success rate in adjoining wheat fields (Linder,et al. 1960).

�-164-

Nest Fate Relative to Time--In both 1971 and 1972, higher predation was
noted on early nests found during the first nest search prior to July 15
(Table 4). Nesting success rates were much higher during the second search.
No difference was noted in 1970 when only four successful nests were found.
The low sample did not permit a valid comparison.

Table 3. Fate of pheasant nests found within
the summers of 1970, 1971 and 1972.

roadside study plots during

No.

1970
Percent

No.

1971
Percent

No.

1972
Percent

No.

Total
Percent

Successful

4

13.8

12

26.7

16

27.6

32

24.2

P~edator

16

55.2

22

48.9

33

56.9

71

53.8

Abandoned

4

13.8

4

8.9

5

8.6

13

9.9

Unknown

5

17.2

7

15.5

4

6.9

16

12.1

Nest Fate

Destroyed

Table 4. Early and late season nest success
1970, 1971 and 1972.

in the roadside study areas,

Year

Period

Total Nests

Number
Successful

Percent
Successful

1970

Before July 15

14

2

14.3

After July 15

14

2

14.3

Before July 15

22

4

18.2

After July 15

23

8

34.8

Before July 15

41

7

17.1

After July 15

17

9

53.0

1971

1972

�-165-

Proximal

Cover Associations

Seeded grasses, alfalfa and their combinations with weeds, downy brome and
residual cover were used extensively by pheasants for nesting in 1972 (Table
5). Weeds and residual cover, which ranked high in use in 1970 were utilized
much less in 1972. Downy brome continued to rank as secondary, but important,
nesting cover in the seeded plots. These results probably reflect declines
in quantities of weeds and residual blown in cover in the seeded roadsides.
Meanwhile, downy brome has continued to compete with seeded stands in many
of the seeded plots.
The farmed roadside controls contained an abundance of downy brome which
spread, in many instances, into the edge of the green wheat in 1972. Ample
spring moisture permitted higher than normal growth which apparently attracted
nesting hens.
Six of ten nests found in farmed controls were associated with
downy b rome •
There is some evidence indicating pheasants primarily utilize cover along
road edges of farmed controls when cover is present in the farmed portion of
the plot. Lesser use is received when summer fallow is adjacent.
Nine of
14 nests were found in plots adjacent to green wheat fields during the past
three years.
The remaining five were adjacent to summer fallow when found,
but actually may have been started when wheat stubble was still present.
One
of these was located in a 10 to 12 foot wide atypical road shoulder.
Most
road shoulders average only two to four feet in width, in farmed controls,
and the remainder acreage is farmed.
During the three years of nest search, nests have not been fOlmd in green
wheat, although this cover occupies about 35 percent of the farmed controls.
Winter wheat often possesses height and above ground canopies equal or
better than grasses, but it lacks the residual camouflage cover and litter
possessed by untilled cover types (Table 6).
Past studies in Nebraska (Linder et ale 1960), South Dakota (Trautman 1960)
and other states all indicate that nest densities in small grain fields are
low in comparison to those in natural roadsides.
However, because of large
acreages and relative safe nesting conditions in small grain lands, a majority
of the pheasants produced come from these fields.
Since wheat fields comprise even a greater percent of the available nesting cover in Northeast
Colorado than in Nebraska and South Dakota study areas, it can be assumed
with confidence that these fields are the key to pheasant production here.
Roadside cover approximates one-half of one percent of the land area, or
about three acres in every 640 acre section (Snyder 1970).
Green wheat
occupies about 250 o.r mo re acres out of every 640.
Pheasants also used a wide variety of cover types for nesting in the unfarmed .
natural controls.
Weeds and residual cover ranked high.
Native grasses,
downy brome and a variety of others, were also included (Table 5).

�-166-

Table 5.
pheasants

Utilization of vegetation within roadside cover types by nesting
during the summers of 1970, 1971 and 1972.

Plot Type

Seeded

Cover Type

Combined
Total

Roadside

Seeded Grass
Grass-Alfalfa
Seeded Alfalfa
Grass-Downy Brome
Alfalfa-Downy Brome
Straight Downy Brome
Grass and Residual Weeds
Grass, Alfalfa and Weeds
Grass, Alfalfa and Downy Brome
Alfalfa, Weeds and Downy Brome
Green Weeds
Weeds and Residual
Weeds, Residual and Downy Brome
Grass, Green Weeds and Residual
Subtotal
Unfarmed

Number of Nests
1970
1971
1972

4
1

9
1

8
6

1

1

o

o

2

6

9

o

3
4
6
2

2

1

o

1

2
2

1
1
2

1

o

o
1

1

o
11
1

o
22

Roadside

o
o
o

21
8

1

5

4

11
4
4
3

2

3
4

2

4
12
4

4

32

39

1

1

1

3

1

2

2

4
4
2
4
6
3

93

(C2)

o
o
o

Native Grass
Native Grass and Downy Brome
Straight Downy Brome
Downy Brome and Residual
Green Weeds
Green Weeds and Residual
Straight Residual Cover
Downy Brome, Weeds and Residual

o
o

2

o

2
1
1

Subtotal

5

11

9

25

o
o

o

1

o

1
2

1
3

3
3

2

Farmed Roadside

o

2

1
2

2

1
1

1

(Cl)

Native Grass
Downy Brome and Residual
Green Weeds and Residual
Straight Downy Brome
Green Weeds
Downy Brome and Green Weeds
Green Weeds, Residual and Downy
Brome

Subtotal

1

2

o

o
o
o
o

2

2
2

o

o

1

1

2

2

10

14

2

o

o

�-167-

Table 6. Comparisons
study plot types. l/

of height

and density

for 1971 and 1972 on roadside

June Measurement
Density
Height
1972
1971
1972
1971

Plot Type

May Measurement
Density
Height
1972
1971
1972
1971

Seeded

8.71

l3.l8

1.65

2.14

16.31

19.00

2.23

2.51

Control

6.08

8.07

1. 35

1.71

13.06

15.10

2.01

2.50

Farmed Control~/

14.18

17.33

1.66

1.95

26.72

25.53

2.19

2.43

Unfarmed

1I Ocular

estimates of density were based on a scale of 1 to 3 using the
latter figure for high density.
Height is represented as inches above ground.

~/ Only plots containing

Life-Form

small grains were sampled.

Characteristics

of Roadside

Cover

Height and Density Measurements
Mid-May and late June height and density measurements were completed on 15
seeded plots, 10 unfarmed controls, and 10 farmed controls.
In the latter
group, only those containing stands of small grain were sampled, both in
1971 and 1972.
Density estimates on all groups were higher than in the
preceding year (Table 6). In seeded and unfarmed controls, residual cover
remained standing because of lack of snow over winter.
Spring moisture
also prompted excellent new growth on all plots.
Cover in seeded and unfarmed controls was also slightly taller than were samples obtained in
previous years.
Fertilization

of Seeded Stands

Little observable effect of nitrogen applications was noted by visual
inspection of new seeded stands (4 years or younger).
Two old seeded stands
showed readily observable effects of nitrogen application.
The first was a
pure seeded stand of intermediate wheat grass (Group 10-7-45) seeded by the
farmer in approximately
1965. When the stand was initially seeded, it was
dense and quickly became root bound and suppressed in overall vigor.
By
1970, the stems were open in placement and not considered attractive to
nesting pheasants.
In late July, 1972, height and weight samples were randomly obtained on
the sample area fertilized in March, 1972, and proximal unfertilized portions
of the plot. Twelve height measurements averaged 29.3 inches tall on the
fertilized section compared to 21.2 inches on the unfertilized sample.

�-168-

Twelve square foot wet weight samples averaged 100.4 grams on the fertilized
section, compared to 57.6 grams in the unfertilized samples.
Both height
and weight were significantly greater on the fertilized strip (Table 7).

Table 7. A comparison of vegetative height and weight
unfertilized old grass stands, July 1972.

Grass Type

Treatment

Sample
Size

Mean
Height

X Sample
Weight

Intermediate
Wheatgrass

Fertilized

12

29.3

100.4

Untreated

12

21.2

57.6

Fertilized

12

22.7

131.4

Untreated

12

16.1

67.S

Brome Grass

on fertilized

and

t Value
of Height

t Value
of Weight

3.66**

5.32**

3.43**

5.70**

A second test was made on an old stand of pure brome grass.
The age of the
stand is unknown.
It contained high stem density but height was considered
inadequate for good nesting cover in all but wet years.
Again statistically
significant growth results were obtained by use of the nitrogen (Table 7).
Visual inspection also showed dramatic differences between treated and untreated portions of the plot.

LITERATURE

CITED

Joselyn G. B. 1970. Management of roadside cover for nesting pheasants.
Job Completion Rpt. Illinois Proj. 10-66-R-S.
Job No.3.
41 p.
Linder, R. L., D. L. Lyon, and C. P. Agee. 1960. An analysis of pheasant
nesting in South-central Nebr. Trans. N. Am. Wildl. Conf. 25:214-230.
Snyder, W. D. 1969. Pheasant
of Game, Fish and Parks.

roadside cover evaluation study.
Colo. Div.
Game Res. Rpt. April.
pp. 37-45.

1970a.
Pheasant roadside cover evaluation study.
Colo. Div. of
Game, Fish and Parks.
Game Res. Rpt. April.
pp. 109-114.
1970b.
Pheasant Hen Harvest Investigation - Final Report.
Colo.
Div. of Game, Fish and Parks.
Game Res. Rpt. April.
pp. 3-81.

�-169-

Snyder, W. D. 1971. Pheasant roadside cover evaluation study. Colo. Div.
of Game, Fish and Parks. Game Res. Rpt. April. pp. 27-39.
1972. Pheasant roadside cover evaluation study. Colo. Div. of
Game, Fish and Parks. Game Res. Rpt. April. pp. 21-29.
Trautman, C. G. 1960. Evaluation of pheasant nesting habitat in Eastern
South Dakota. Trans. N. Am. Wildl. Conf. 25:202-213.

,o/Lalz/teJi- &amp;. .d-rt~/rI:Uc./'
~

Prepared by

)/,}

Warren D. Snyder
Wildlife Researcher

r

~

/
f

I

��April, 1973

-171-

JOB PROGRESS REPORT

State of

COLORADO
------~~~~-------------

Project No.

W-37-R-26

Work Plan No.

3

Game Bird Survey

Job Title

Job No ..
Sa
Effects of Sagebrush Control
on Distribution and Abundance of Sage Grouse

Period Covered:

April 1, 1972 through March 31, 1973

Personnel:

Robert Streeter, Russell Kozacek and Jack Gustafson, Cooperative
Research Unit, Warren Snyder, Don Hoffman, Tom Lederhos, Clait
Braun, Don Gore, Courtney Crawford, John Ellenberger, Don Benson,
Mike Szymczak and Howard Funk, Division of Wildlife.

ABSTRACT

Low numbers of sage grouse observed on strutting grounds in spring of 1973
seem to indicate peak periods of activity were missed by the observers.
Only one ground increased in numbers counted, while the remainder declined
Significantly. Other factors also seem to contribute to making the total
counts for the grounds invalid. Hunter participation during the season
increased over 1971, but birds bagged per hunter remained stable. Thus,
hunter success decreased. A low percentage of immatures in the bag suggested
1972 production was almost at a record low. Trapping efforts were revived in
the spring of 1973 with 288 birds banded on Wintering and strutting ground areas.

��-173-

EFFECTS OF SAGEBRUSH CONTROL
ON DISTRIBUTION AND ABUNDANCE OF SAGE GROUSE
Howard

D. Funk

This job has been active since 1963, with an initial 2-year study of sage
grouse abundance and distribution in the Lake John area northwest of Walden.
The study produced recommended patterns for spraying for sagebrush control,
with spraying accomplished to design specifications in 1965.
An investigation of the immediate effects of the spray project on the sage grouse
population was initiated in 1965 for a 2-year period.
This was followed by
an additional two years of banding arid instrumenting birds with telemetry
equipment for movement studies.
Investigation of the long term effects of
the spray project was scheduled for 1973-74 and 1974-75.
During the interim
period between 1969 and 1973 efforts have been limited to collection of
study area strutting ground data, banding of birds when possible, and
operating check stations near Walden and Cowdrey to obtain harvest data.
This report is limited to coverage of the latter,it~s
for the 1972-73
period.
. ;
.

P. S. OBJECTIVE
To determine the effects of 2,4-D spraying of sagebrush on: (1) sage grouse
abundance, (2) sage grouse distribution, and (3) vegetative composition and
density.

SEGMENT
the seasonal

OBJECTIVES

1.

To investigate
is con trolled.

2.

To investigate sage grouse numbers
to the area treated.

METHODS

distribution

of sage grouse after sagebrush

by age and sex classes

on and adjacent

AND MATERIALS

Wildlife Conservation Officers in North Park conduct annual counts on a
number of strutting grounds including those in the study area.
Usually,
three early morning counts are made on each ground.
Data from these counts
were gathered from the Regional Biologist, high counts for each ground being
reported herein.
Two check stations near Walden and Cowdrey were again operated the first two
days of season to collect various data on harvest such as harvest success,
age and sex ratios, banded bird information, and hunter participation
for a

�-174-

check on the year in comparison with other years.
Project personnel were
responsible for the station, with Conservation Officers also contributing
data from field checks.
Main emphasis was on collecting information from
the study area. Age and sex of bagged birds were determined by methods
described in previous reports.
In early March, 1973, two temporary employees hired by the Colorado Cooperative Wildlife Research Unit began trapping, banding and color marking sage
grouse in the Walden area under the direction of Assistant Unit Leader Dr.
Robert G. Streeter.
Most birds were captured with hand nets and night
lighting equipment.
Standard leg bands were placed ,on the birds, along with
colored bandette-type leg bands to assist in recognition of birds by sex
from specific wintering or strutting ground areas.

RESULTS AND DISCUSSION
Strutting

Ground Counts

Strutting ground count data are shown in Table 1. Peak counts seem to have
occurred from mid- to late April, which is somewhat later than average.
Usually high counts are obtained about mid-April, depending upon snow and
weather conditions.
However, results also seem to indicate peak of strutting
activity was missed by the observers, thus negating the value of the counts
in comparison with other years.
The MOnahan Draw ground (SG5), for example,
dropped from a high of 81 cocks present in 1971 to 10 in 1972, with the high
total bird count also being 10 birds, which were probably all cocks.
Both
items strongly suggest invalid data, especially when the high count date was
as late as April 29. All areas, with the exception of the Alkali Lake
ground (SG4), showed declines in numbers (Table 2), which could occur in a
given year.
However, this was probably not the case, because of late counts,
for whatever reason, and peak activity probably was not observed.
Strutting ground data by year are presented in Table 2, illustrating annual
high count figures for cocks and relative annual fluctuations.
As shown,
there was a drastic decrease in numbers of cocks seen in 1972 as compared to
1971.
Also, number observed in 1972 was the lowest since 1965, when only
166 cocks were recorded on four grounds.
There is a possibility that some of the birds had moved to new grounds in
1972, which would have some effect on numbers observed on the 12 known grounds.
The reason for this statement is that three new grounds were observed in, or
adjacent to the study area in spring of 1973.
If these grounds were active
in spring of 1972 as well, numbers
of birds present on the known grounds
might be expected to drop.
This is only a possibility, but it has to be
considered as at least part of the reason for the decline in numbers.
Data
on the new grounds for 1973 will be presented in the next segment report,
along with those from the 12 known grounds.
Check Station

Results

The Cowdrey and Walden check stations were again operated by project personnel for the first two days of season, September 9 and 10, 1972.
Number of

�-175-

hunters checked increased over 1971, but this was still the second lowest
indicated level of participation since 1967 (Table 3). Birds per hunter
(.56) decreased from the 1971 level making 1972 the second lowest year on
record for success since 1963. Only 1970 was lower with only .47 birds
per hunter.
Hours hunted per bird also increased over 1971, which made 1972
the second highest year on record since 1963 in this category.
Juvenile
birds in the bag decreased to 33 percent, almost a record low, which suggests
a relatively poor production year in 1972. However, sage grouse check
station results are probably not the best estimates of production, especially
in North Park.

Table 1.

Peak counts of male and total sage grouse on strutting

Strutting

Ground Jj

SG

1 ]j

SG

2

SG

3 2:./

SG

Number

Males

Total Birds

grounds,

High Count Date

o

o

4

96

127

-1/

SG

5

10

10

April 29

SG

6

4

4

April 20

SG

7 '!:../

SG

8

o

o

- 1/

SG

9

62

86

April 18

SG 10

7

12

-1/

SG 11

No count

No count

SG 12 2:./

Total

!/ Locations

179

of grounds shown in previous

'!:../ Counts discontinued.

1/ No date given.

239
reports.

1972.

�Table 2.

Comparison of 1960 through 1972 strutting ground counts of male sage grouse.

Strutting
Ground

1960

1961

1962

1963

1964

1965

Year
1966

1967

1968

1969

1970

1971

1972

SG

1

10

5

0

0

0

0

0

0

0

0

0

1/

11

SG

2

43

4

7

20

26

9

9

10

2

8

8

10

0

SG

3

14

0

0

0

0

0

0

0

0

0

0

1./

1/

SG-

4

130

96

219

216

120

53

47

69

58

82

66

88

96

SG

5

19

2

52

17

65

52

54

59

52

82

57

81

10

SG

6

5

1

8

9

3

0

2

12

36

36

19

25

4

SG

7

0

7

5

0

0

0

0

1

3

2

0

1/

1/

SG

8

5

0

0

0

0

0

6

5

2

6

7

5

0

SG

9

167

109

71

85

99

52

97

81

71

117

97

96

62

SG

10

17

77

28

19

17

0

6

9

0

16

16

21

7

SG

11

12

50

5

11

7

0

13

15

6

5

4

0

Y

SG

12

14

12

0

0

0

0

0

0

1

3

0

11

1/

Total

446

363

395

377

337

166

234

265

231

357

274

326

179

1/ Count discontinued.
'!:.../

No coun t ,

I

~
..•..•
Q'\
I

�Table 3.

Comparison of North Park sage grouse hunter check information, 1963-1972.

1/

Year

Bag
Limit

Hunters
Checked

Hours
Hunted

Adult
Males

Adult
Females

Percent
Adults

Juvenile
Males

Juvenile
Females

Percent
Juveniles

Total
Birds

Birds
Per
Hunter

Hours
Hunted
Per Bird

1963

3

492

2,460

62

150

42

113

181

58

506

1.03

4.86

1964

2

217

624

25

81

59

28

45

41

179

.82

3.49

2

150

626

27

30

49

26

26

51

116

.77

5.40

1966

2

306

1,227

31

116

56

45

71

44

263

.86

4.67

1967

2

300

1,177

50

127

67

42

46

33

267

.89

4.41

1968

2

546

2,604

80

135

42

156

141

58

512

.94

5.09

1969

2

662

2,936

121

277

70

57

114

30

569

.86

5.16

1970

2

564

2,617

58

89

55

53

66

45

266

.47

9.84

1971

2

357

1,802

28

67

51

54

39

49

264

.74

6.83

1972

2

452

2,377

50

69

67

40

18

33

253

.56

9.40

1965

u

I

t-'
"'-I
"'-I

!.J Based on Walden and Cowdrey check station data during opening weekends.
~/ A one day season, hence only a one day check station.

I

�-178-

Trapping

and Banding

As mentioned previously, trapping and banding efforts were again initiated
in North Park during the spring of 1973.
Because the majority of work was
accomplished after the close of this segment period, complete results will
be listed in the next segment report.
Briefly, Cooperative Wildlife Research
Unit personnel, with the cooperation of local Conservation Officers, banded
109 female and 179 male sage grouse from March 26 through May 24, 1973.
Some were trapped on winter areas off the study area to obtain information
on dispersal to strutting grounds.
The remainder were trapped on study area
grounds.
Mr. Thomas Beck was chosen for, and accepted the graduate assistantship with
the Unit, and joined the project as the principal investigator in June, 1973.

Prepa red by __

L

J

r,

-'I

7

,_,_,_J_(.(_,J".C_(.' ._)G_i!_ .u{
__':-~
_::-:_'_&lt;_7_' _ LL_«(_/_1!'_'l._J_~
.»

_

Howard D. Funk
Section
Chief, Small Game Research

�April, 1973

-179-

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No.

W-37-R-26

Work Plan No.

8

Job Title

Job No.

2

Inventory of Lesser Prairie Chickens

Period Co~ered:
Personnel:

Game Bird Survey

April 1, 1972 through March 31, 1973

Ann Leckler and Donald M. Hoffman.

ABSTRACT

A complete census of known lesse~ pra~r~e chicken booming grounds was
attempted in 1972 following a 10-year period when no counts were conducted
except for limited counts on a few grounds by u. S. Forest Service personnel
and others. Weather conditions in the spring of 1972 were unfavorable for
censusing work during approximately one-half of the mornings when attempts
were made. This was due to high winds, rain and/or thunderstorms.
A recheck of 13 known booming grounds discovered prior to 1962 showed 7 have
apparently been abandoned for various reasons and the remaining 6 are still
being used. Three new grounds were located in 1972. In 1972, a total of
110 lesser prairie chickens were counted on 9 active grounds compared with
130 on the 13 active grounds in 1962. This amounts to an apparent reduction
in population of 15 percent although numbers of birds counted per ground was
12.22 in 1972, compared with 10.00 in 1962. Because of unfavorable weather
conditions in 1972, caution should be exercised in comparing 1972 population
counts with those of 1962.
Early morning listening checks in areas in Lincoln. County and Baca County
where releases of wild-trapped lesser prairie chickens from Kansas were made
failed to disclose the locations of booming grounds. No releases have as
yet been made in El Paso County so this area was not checked.

�-180-

RECOMMENDATIONS

Southeast Regional personnel have expressed a desire to assume population
counts of lesser prairie chickens in Colorado.
Census activities by
Project W-37-R personnel should, however, be continued during the spring
of 1973 (Segment 27) to instruct Regional personnel on ground locations and
methods of census.
It is recommended that this study be completed with the
write up of final results following the spring, 1973 inventory.

�-181-

INVENTORY OF LESSER PRAIRIE CHICKENS
Donald M. Hoffman

P. S. OBJECTIVE
To determine the population status of lesser prairie chickens in Colorado.

SEGMENT OBJECTIVES
1.

To inventory booming grounds located during the period 1959 through
1962 in Baca and Prowers counties.

2.

To locate additional booming grounds in Baca, Prowers, Lincoln, and
El Paso counties and obtain population status data.
METHODS AND MATERIALS

Ntnnbers of males and female lesser prairie chickens using known booming
grounds were determined through early morning counts during the period
mid April through late May, 1972. Census work was conducted in alternate weeks during the period. Flushing counts were secured after completing counts on both accessible and inaccessible grounds.
Early morning listening checks were made along roads and trails in
occupied ranges to locate additional display grounds. Persons living
and working within lesser prairie chicken ranges were interviewed to
secure leads on the locations of new flocks.
Early morning listening checks were conducted in the vicinities where
lesser prairie chickens were released in (1) Lincoln County, south of Hugo
(1968) and Co Baca County, southeast of Springfield (1972). The proposed
release for EI Paso County had not materialized so no checks were conducted
in this area.
Maps of locations of new booming grounds were drawn from field sketches
made and will be included in the final report (Segment 27).

RESULTS AND DISCUSSION
During January and February, 1972, 49 additional lesser pra1r1e chickens
were received from Kansas through a wildlife trade agreement for antelope.
Thirty-six of these (18 hens and 18 cocks) were used for a new release on
the Comanche National Grasslands southeast of Springfield, Colorado, and
13 excess cocks were used to add new bloodlines to existing populations
southeast of Campo, Colorado. All birds released were leg banded prior to
release (Fig. 1). A view of the sand sagebrush-grassland habitat where the
36 lesser prairie chickens were released along Sand Arroya in Baca County is
shown in Fig. 2. Table 1 summarizes all lesser prairie chickens received
from Kansas to date and results of past releases made.

�Table 1.

Lesser prairie chickens received from Kansas.

Date

·Hens

No. Birds
Cocks
Total

March 23, 1968

10

12 1/

22

April 2, 1968

4

2

6

Sub-totals

14

14

28

January 20, 1972

8

13

21

January 29, 1972

8 2:./

2

10

February 24, 1972

2

3

5

--

--

--

18

18

36

January 29, 1972

0

13

13

TOTALS

32

45

77

Sub-totals

Area Released

Results to Date and Remarks

Lincoln County, Big Sandy
Creek, 7 mi. south of Hugo

"

"

"

Baca County, Sand Arroya,
11 mi. southeast of Springfield.

"

"

One band recovered by W.C.O. Lengel
from road-killed lesser near Boyero
(15 miles southeast of release site).
Group apparently moved down Big Sandy
Creek.

One band recovered from bird killed
by flying into fence on May 28, 1972
on Dale Schroder farm, approximately
2 mi. southwest of release site.

"

Baca County, Dye, Glover
pasture, 12 mi. southeast
of Campo.

1/
- One of these cocks died enroute to release site.

2/
- One of these hens could not fly so sacrificed and used for study skin.

I

I-'
+:00
I

These were excess cocks released in
an occupied lesser range to add new
bloodlines to existing flocks.

�-185-

Early morning listening checks conducted in the vicinities where lesser
prairie chickens received from Kansas in a wildlife trade agreement including (1) Lincoln County, south of Hugo and (2):Baca County, southeast
of Springfield, ,failed to disclose the locations of booming grounds.
"Table 2 lists a comparison of highest numbers of lesser prairie chickens
counted on booming grounds in 1962 and 1972. Of 13 known grounds censused
in 1962, 7 have apparently been abandoned for various reasons and 6 are
still being used. Three new grounds were located in 1972. A total of 110
lesser prairie chickens were counted on 9 active grounds in 1972, compared
with 130 on 13 active grounds in 1962. This amounts to an apparent reduction in population of 15 percent although numbers of birds counted per
ground was 12.22 in 1972, compared with 10.00 in 1962. Fig. 3 shows 2
cocks on an active booming ground in Baca County in 1972.
Weather conditions were often unfavorable for censusing lesser prairie
chickens in 1972. During alternating weeks during the period from April
11 to May 26, 1972, counts were attempted in Baca and Prowers counties
with approximately one-half of the mornings determined to be unsuitable
because of high winds, rain, and/or thunderstorms. For this reason,
caution should be exercised in comparing 1972 population levels with those
of 1962 until additional census work is completed.

Prepared by

J)~~~

DonaldlM.~offman
Wildlife Researcher

...,

�I
•.....

00

~
I

Fig. 3. Two lesser prairir chicken cocks pause during display activities on a ground in Baca County,
1972. (Photo by D. Hoffman)

�Table 2. A comparison of highest numbers of lesser prairie chickens counted on booming grounds, 1962
and 1972.
1972
-Unc.Iass ,
Hens

Display Ground

County

Cocks

Hens

1962
Unclass.

Total

Cocks

C. Low (a) 11

Prowers

5

1

0

6

0

T. Stalford (a)

Baca

10

0

3

13

Dye, Glover (a)

Baca

12

0

3

15

11

T. Stalford (c) ~I

Baca

13

0

3

16

0

Hanes (a)

Baca

12

0

8

20

Abandoned

Perkins, Tanner (a)

Baca

4

0

2

6

Abandoned

C. Low (b) 11

Prowers

7

0

1

8

Total

8

8

1

0

12

0

23

23

20

20

0
Abandoned

0

0

I

t-'
00
.....•

Dye, Glover (b)

Baca

5

0

1

6

Abandoned

T. Stalford (d) 11

Baca

2

0

0

2

Abandoned

Lowder (a)

Baca

13

0

3

16

10

0

5

15

Dye, Glover (c)

Baca

7

0

0

7

15

1

0

16

Chick, Dow (a)

Baca

9

0

1

10

Abandoned

Shell Oil (a)

Baca

5

0

0

5

Dye, Glover (e) 11

Baca

5

Abandoned
0

0

5

Shell Oil (b)

Baca

4

0

3

7

Dye, Glover (f) !I

Baca

4

0

0

4

49

2

59

110

I

Totals
Average Per Ground

104

1

25

130
10.00

12.22

--

II

- Letters a, b, c, etc. refer to first, second, third, etc. ground located on various properties.
(d) was a ground discovered in 1963 by Warren D. Snyder and was abandoned prior to 1972.

Dye, Glover

��April, 1973

-189-

JOB FINAL REPORT

State of

COLORADO
------~~~~~-----------

Project No.

W-37-R-26

Work Plan No.

10

Job Title:

Job No.

1

Study of Hungarian Partridge Adaptablity

Period Covered:
Personnel:

Game Bird Survey

April 1, 1972 through March 31, 1973

Charles Brown, Gary Brown, Robert Clark, John Corey, Gurney Crawford,
Corrin Davidson, David de Calesta, Ronald Desilet, Harry Dobbs, Gordon East, Roger Evans, Mark Frasier, Carroll Grand Pre, Aden Greer,
George Harrington, James Houston, Roger Lowry, Ted McKinney, Jon
Moser, Daniel Potts, Dale Reed, William Roland, Tom Rauch, Wayne
Sandfort, Robert schmidt, Walter Schuett, Bruce Sigler, Warren Snyder,
Dale Stahlecker, Harold Swope, Henry Wilson, Michael Zgainer, Ann
Leckler and Donald Hoffman.

ABSTRACT

From 1964 through 1973, a total of 4,299 Hungarian partridges were released in
Colorado in 3 main areas
A total of 135 were released in Larimer County,
1,744 in Moffat County and 2,376 in the San Luis Valley (Alamosa, Conejos and
Saguache counties). Fourteen were also released in Routt County during this
period. By sources, 175 of these were wild-trapped birds from Oregon and Idaho
and the remainder (4,124) were pen-raised by the Colorado Division of Wildlife.
o

Although some reproduction followed the release of 135 wild trapped Huns in the
Douglas Lake area of Larimer County in early 1964, this population had completely
disappeared by 1965. Limited reproduction followed the releasing of 40 wild
trapped and 1,734 pen raised Huns in Moffat County from 1964 through 1970, but
the species failed to become established. Results of trial releases of 2,376 pen
raised Huns in the San Luis Valley (Alamosa, Conejos and Saguache counties)
from 1971 through 1973 are very questionable as to the success in establishing
reproducing populations. Late fall surveys in December, 1971 and December, 1972
indicated only small nuclei populations of Huns had survived but no follow up
checks were made following the final releases made in March, 1973. The probability that the species will become established in the San Luis Valley is slim.

�-190-

RECOMMENDATIONS

The decision was made during the Small Game Project Review meeting in January,
1973 to release all remaining pen raised Hungarian partridges during the spring
of 1973, because development from all preceding releases was far less than had
been hoped for at the start of the study.
Wildlife Conservation Officers and other Division of Wildlife personnel who
work in the areas of past releases have been requested to submit future sightings of Hungarian partridges to the Research Section.
It is recommended that this job be terminated with this final report and any
further introduction of exotic game bird species await the outcome of these
Hungarian partridge releases.

-r

�-191-

STUDY OF HUNGARIAN PARTRIDGE ADAPTABILITY
Donald M. Hoffman
INTRODUCTION
Early introductions of Hungarian or gray partridges (Perdix perdix perdix)
into Colorado were unsuccessful in establishing permanent populations.
Twenty-five pairs of Huns were initially imported in 1910 and released along
the foothill areas in the eastern part of the State. This effort was followed
by subsequent releases in many areas of Colorado over several years, involving a total of around 5,000 birds (McKinney unpublished).
Even though these early introductions failed. the Colorado Department of
Game, Fish and Parks in 1959 again became interested in trying to establish
this desirable exotic game bird species. Evans and Sandfort (1960) concluded
the climates and habitats of the Fort Collins, Craig, Del Norte, Montrose, and
Norwood areas were suitable for the Hungarian partridge.
An initial trade agreement was made between the States of Colorado and Oregon
whereby 20 wild turkeys from Colorado would be traded to Oregon for 80 to 100
wild trapped Hungarian partridges (Sandfort 1962). Additional wild trapped
and pen raised Huns were later secured from Oregon and Idaho through wildlife
trades. In 1964, eleven pairs of wild trapped Huns were held for artificial
propagation purposes and 51 pen raised Huns from Oregon were added to this
stock in 1965, since it was deemed desirable to release relatively large
groups of birds and the potential for securing wild trapped stock was low.
The study was initiated by Wayne W. Sandfort from September, 1961 to March,
1963, continued by Harold M. Swope from April 1963 to July 1968, continued
and concluded by Donald M. Hoffman during the period August 1968 to March 1973.
Throughout the entire period of the study (1961-1973) the two main objectives
were to release Hungarian partridges and determine the success of Ln t roduc t Lons .
P. S. OBJECTIVE
To determine the ability of Hungarian partridges to establish reproducing
populations in mixed brushland~heatland habitat in selected sites in Colorado.
METHODS AND MATERIALS
Methods and materials used in this study have varied by years and by individual investigator. These are summarized with the source of data given in
parentheses after the year as follows:
1961 (Sandfort 1962)
A trade agreement was made between the states of Colorado and Oregon whereby
20 wild turkeys from Colorado would be traded to Oregon for 80-100 wild

�-192-

Hungarian partridges. The wild turkeys were shipped to Oregon during February 1961. Preliminary to release of the Huns, surveys were made within
several areas suggested by Evans and Sandfort (1960). Two initial sites
selected for release of the wild birds were the Round Bottom Area of Moffat
County and the Douglas Lake Area of Larimer County.
1962 (Sandfort 1963)
Procedures of the study, in the event birds are received, will involve gentle
release from a holding pen in the excellent ~over around Douglas Lake, north
of Fort Collins. Several birds will be held in captivity in attempts to
establish a sizable breeding flock, making large scale trial releases eventually possible. Following introduction, periodic field surveys will be
conducted to determine reproductive success, disperson, general survival and
adaptability.
1963 (Swope 1964)
About half the birds received from Oregon and Idaho were held in small pens
at the release sites for several days prior to liberation. They were fed,
watered and allowed to leave the pens voluntarily. Feed and water were placed
near the release sites through the late winter months. Release areas were
covered on foot and nearby roadsides were driven in an effort to observe the
Huns. Local Department personnel assisted in these surveys. People working
and living in the vicinity of the releases were interviewed regarding their
observations.
1964 (Swope 1965)
It was found that standard roadside census techniques were of little use in
observing Huns. The intensive search of large areas of cover on foot produced
very few observations. Most birds were seen by accident while observers
were working or passing through the area. Local ranchers were again interviewed. Release techniques were similar to those used a year ago. About half
the birds were held in a small pen for several days while the remainder were
released in the same area. Feed was scattered around the release area until
the snow disappeared. A graduate student (Ted D. McKinney) from Colorado
State University was assigned to a study of the Hungarian partridge in the
Douglas Lake area. His work included the compilation of basic life history
data and an evaluation of the Larimer County release.
1965 (Swope 1966)
Roadside census methods and searches of release site vicinities on foot
proved ineffective. Interviews of local residents, and observations by
resident Game, Fish and Parks Department personnel were also made without
success.
1966 (Swope 1967)
Wildlife Conservation Officers and ranchers in the vicinities of the release
sites have been asked through newspaper articles and personal contact to be

�-193-

on the lookout for Huns and to record number, location, and observation date.
These persons were interviewed by a mailed questionnaire to check on the
results of their observations and some were contacted personally.
1967 (Swope 1968)
Followup personal contacts were made on a landowner questionnaire circulated
in 1966. Locally assigned Department personnel were asked to obtain and
report information on Hun sightings. Hungarian partridge produced at the Fort
Collins Wildlife Research Station as a result of a study to develop production
techniques were released in the Craig area.
1968 (Hoffman 1969)
Twenty followup mail questionnaires were sent in December 1968 to landowners
in Moffat County who answered the 1966 mail questionnaire.
Due to personnel
changes, no landowner contacts were made during 1968.
1969 (Hoffman 1970)
Field releases of pen raised stock were made with the use of a helicopter to
fly the crated birds from Round Bottom on the north side of the Yampa River
to Isles Mountain south of the river. Intensive field searches were made by
Project W-37-R personnel during the periods June 11-13. July 14-17, August 5-8,
18-22, 26-29 and September 2-5, 1969. In addition, persons living and ,working_
in or near the release site were interviewed and observations reported were
recorded. A winter search of the release sites and surrounding areas in
MOffat County by 12 Division employees including 2 Researchers, 6 w.c.o
Trainees, 3 W.C.O.'s, and 1 Area Supervisor was made during the last week of
February 1970.
1970 (Hoffman 1971)
Field releases of pen raised Huns were made in 3 areas an Moffat County on
April 24, 1970. Intensive field searches were made by Project W-37-R
personnel during the periods June 15-19; August 25-28; and September 14-17,
1970. In addition, persons living and working in or near the release sites
were interviewed and observations were recorded.
1971 (Hoffman 1972)
On April 23, 1971, 655 pen raised Hungarian partridges from holding pens at
the Fort Collins Wildlife Research Station and the Rocky Ford Bird Farm were
released at 3 sites in the San Luis Valley. Intensive field searches of all
release sites in the San Luis Valley were made by Project personnel during
the periods June 7-11, July 14-15, August 2-6, September 13-16, and December
9-11, 1971. Persons living or working in or near the release sites were interviewed and observations were recorded. Reports of Huns from Moffat County
were investigated during the period June 29-July 2, 1971.
1972 (Hoffman 1973)
On April 25, 1972, 530 pen raised Hungarian partridges from holding pens at
the Fort Collins Wildlife Research Station were released at the same 3 sites

�-194-

in the San Luis.Valley as the 1971 releases. On July 24, 1972, 86 breeders
used in the 1972 test series were transported to the Hot Creek Management
Area site in the San Luis Valley and added to the earlier releases. Intensive field searches of all release sites in the San Luis Valley were made by
Project personnel in June, July. and December, 1972. Persons living or
working in or near the release sites and surrounding areas were interviewed
and observations were recorded. During the period March 20 through March 24,
1973 additional field releases were made of 1,105 pen raised Huns at 5 sites
in the San Luis Valley. These were the remainder of the pen raised stock
held at the Fort Collins Wildlife Research Station.

RESULTS AND DISCUSSION
Table 1 lists all field releases of Hungarian partridges in Colorado for the
period 1964 through 1973. IX total of 135 Huns were released in Larimer County
in 1964 at a site shown in Fig. I, 1.774 in Moffat County at sites shown in
Fig. 2, 1,928 in Alamosa and Saguache counties at sites shown in Fig. 3, and
448 in Conejos County at a site shown in Fig. 4. The map location of a release
of 14 on April 30, 1968 in Routt County is not shown because the exact location is unknown. Thus, a total of 4,299 Hungarian partridges were released
in Colorado in 3 main areas from 1964 through 1973. By source, 175 of these
we re wil d trapped birds from Ore gon and Idaho and the remainde r (4 ,124) we re
pen ~aised by the Colorado Division of Wildlife. Table 2 lists Hungarian
partridge reports and observations for Moffat County for the years 1969 through
1971, and Table 3 lists similar information for the San Luis Valley (Alamosa,
Conejos, and Saguache counties) for 1971 and 1972. Findings are summarized
by years with the source of data 3hown in parentheses as follows:
1961 (Sandfort 1962)
Attempts to trap wild Hungarian partridge during the winter of 1961-62 in
Oregon, were unsuccessful. Trapping efforts will be resumed during the fall
and winter of 1962-63.
1962 (Sandfort 1963)
No birds were received during this segment.
1963 .(Swope 1964)
The following is a record of the receipt and dispOSition of the Hungarian
partridges received from Idaho and Oregon.
120 - birds received from Oregon (83 live birds
received, one
died later)
120 - birds received from Idaho (115 live birds
received, 131
shipped)
22 - birds taken to Rocky Ford Experimental
Bird Farm (paired)
135 - released at Doug.LasLake release site
40 - released at Round Bottom release site.
In the Round Bottom area only 1 authenticated report of a Hun observation
was received in the several months following release. They were released near
the end of March, 1964. A local rancher (Lorence Ellgen) saw 2 birds at the
release site 1 week later.

�-195-

Table 1.

Hungarian partridge field releases, all areas, 1964-1973.

Date
Jan. &amp; Feb., 1964

Number
Released

Source of
Stock

Release Area

135*

Oregon &amp; Idaho
(wild trapped)

Larimer County, Douglas
Lake Sec. 36, T9N, R69W

Subtotal

135

March 28, 1964

40

Idaho
(wild trapped)

M::&gt;ffatCounty, Round
Bottom Sec. 34, T6N, R92W

March 23, 1965

124

Oregon
(pen raised)

MOffat County, Round
Bottom Sec. 33, T6N, R92W

April 18, 1967

71

FCWRS
(pen raised)

Moffat County, Round
Bottom Sec. 21, T6N, R92W

April 30 and
May 1, 1968

236

FCWRS
(pen raised)

MOffat County, Round
Bottom Sec. 29, T6N, R92W

April 30, 1968

14

FCWRS
(pen raised)

Routt County, Northeast
of Hayden

April 19, 1969

389

FCWRS (374) &amp;
CSU (15)
(pen raised)

Moffat County, Isles
Mountain Sec. 24 &amp; 25,
T5N, R92W

April 19, 1969

240

FCWRS
(pen raised)

MOffat County, Isles Mtn,
Sec. 16, T5N, R92W

April 24, 1970

170

FCWRS
(pen raised)

MOffat County, Johnson
Gulch Sec. 15, T6N, R91W

April 24, 1970

234

FCWRS
(pen raised)

MOffat County, Jubb Creek
Sec. 35, T5N, R93W

April 24, 1970

270

FCWRS
(pen raised)

Moffat County, Price Creek
Sec. 30 &amp; 31, T4N, R95W

Subtotal

1,788

April 23, 1971

286

FCWRS
(pen raised)

San Luis Valley, Alamosa Co.,
Robert Young-farm Sec. 7,
T38N, RllE

April 23, 1971

200

FCWRS
(pen raised)

San Luis Valley, Conejos
Co., Hot Creek Management
Area Sec. 15, T35N, R7E

�-196-

Table 1.

Hungarian partridge field releases, all areas, 1964-1973 (cont.).
Number
Released

Source of
Stock

April 23, 1971

169

FCWRS
(pen raised)

San Luis Valley, Saguache
Co., Ernest Stoeber farm
Sec. 32, T4lN, RlOE

April 25, 1972

103

FCWRS
(pen raised)

San Luis Valley, Alamosa
Co., Robert Young farm
Sec. 7, T38N, RlIE

April 25, 1972

162

FCWRS
(pen raised)

San Luis Valley, Conejos
Co., Hot Creek Management
Area Sec. 15, T35N, R7E

April 25, 1972

265

FCWRS
(pen raised)

San Luis Valley Saguache
Co., Ernest Stoeber farm
Sec. 32, T41N, RlOE

July 24, 1972

86

FCWRS
(pen raised)

San Luis Valley, Conejos Co.,
Hot Creek Management Area
Sec. 15, T35N, R7E

March 20, 1973

328

FCWRS
(pen raised)

San Luis Valley, Alamosa Co.,
Ernest Stoeber farm Sec. 25,
T40N, R9E

March 21, 1973

180

FCWRS
(pen raised)

San Luis Valley, Alamosa
Co., Robert Young farm
Sec. 7, T39N, RlOE

March 21, 1973

180

FCWRS
(pen raised)

San Luis Valley Saguache
Co., Ernest Stoeber farm
Sec. 22, T4lN, RlOE

March 24, 1973

240

FCWRS
(pen raised)

San Luis Valley, Alamosa
Co., William McKinley farm
Sec. 35, T40N, RlOE

March 24, 1973

177

FCWRS
(pen raised)

San Luis Valley, Alamosa
Co., Eric Ecklund farm
Sec. 18, T40N, RlOE

Subtotal

2,376

Grand Total

-4,299

Date

Release Area

* An additional 22 birds (11 pairs) were taken to Rocky Ford Experimental
Bird Farm from this shipment. On October 6, 1965, 51 (pen raised) birds were
received from Oregon and were taken to the Fort Collins Wildlife Research
Station for experimental breeding purposes. On February 19, 1970, 13 1969
hatched Huns from eggs secured from wild trapped birds in England were
received from Winchester Farms, Alton, Illinois and were used for experimental
breeding purposes in 1970 and later years.

�Fig. 1. Portion of Larimer County showing location with numbers of Hungarian partridge released during 1964.
ofuL_

~n_~~~
R.68W

I

_ ~ -!l." _~ I_.ft!

~
~

�..

,

R91W

SHEET

3

T.6 N

N

\

..

T5N.

10

II

,.

,.

Fig. 2. Portions of Moffat County showing locations with numbers of
Hungarian partridge released from 1964 through 1970.

�-199A

u

G

c

A

E

H

10

I.

18

17

I.

I.

17

I.

12

~

15
"

T39N.,

;;;
23

2.

20

"

21

0

2.

2'

20

21

22

23

30

2.

28

27

2.

31

32

"

30

~

2.

35

2.

28

32

33

27

I

i

.!

Fig. 3. Portions of Alamosa and Saguache counties showing locations
numbers of Hungarian partridge released from 1971 through 1973.
FAP

I:

• ,".~I~I
__ ~'~.__ ~I_.

..

with

~lP~ __

~~_p
__~_~_"_V__...

�-200-

--------_ .. - .~-

o

G

R

A

N

E

0

C

0

U

T

y '"~ f5'-

N

i? ~

Line

I
I

g

~

&lt;3

10

...

B
.4.0
10

18

i

I.

17

0

----15
13

!T36N.
2'

0

12

,.

18

Ai

15

22
2'

'9

21

19

•

30
MORGAN

I

~35N .

•...
Ul
Ul

:x:

(/)

&gt;....

z
:::&gt;
o
o

(

10

12

12

(/)

o
..,
Ul
Z

o

u

CONEJOS

COUNTY

II
II

/1
I

I.

I

T.34N.

~'.-;.==:::=

23

'9

2'

22

0'

o.

OJ

r{

2'

l
II

"

.,

1/
SCALE

Fig. 4.

Portion

,

29

OF MILES

t

of Conejos County showing locations
Hungarian partridge released during 1971 and 1972.

28

27

II

/I

Q)

2.

·5

o

\\

with numbers

25

"0

of

30

I

�-201-

Table 2. Summary of Hungarian partridge reports and observation, Moffat
County, 1969-1971.
Date

Number Huns

Area

Observed By

Late Winter 1969

Several

Round Bottom area

L. Ellgen

Spring 1969

2 (1 pair)

Isles Mountain, BLM
catchment

G. Hilton

4 (2 pair)

Isles Mountain, saddle south G. Hilton
of BLM catchment

Few

Round Bottom, Fuhr Gulch

J. Counts

2 (1 pair)

Isles Mountain, south of
Meade Ranch Headquarters

Meade Brothers

6 (3 pair)

Isles Mountain, north of
Meade Ranch Headquarters

Meade Brothers

2 (1 pair)

Isles Mountain area

Meade Brothers

4

Isles Mountain, Bowers
fields

B. Bowers

2 (1 pair)

Isles Mountain, Meade
Brothers property

D. de Ca1esta

July 16, 1969

2 (1 pair)

Isles Mountain, Meade
Brothers property

D. de Ca1esta

August 5, 1969

4 chicks

Isles Mountain, Bowers
fields

D. de Ca1esta

Summer 1969

July 15, 1969

(~ grown)
August 6, 1969

3 broods

Isles Mountain Bowers
fields

B. Bowers

August 7, 1969

2 (1 pair)

Isles Mountain, Bowers
fields

D. de Ca1esta

August 19, 1969

1 adult,
male

Isles Mountain, Meade
Brothers property

D. de Calesta

October 20, 1969

60 to 80

Cedar Mountain

Hunters report
to D. Reed

April, 1970

3 groups of
approx. 8
each

Jubb Creek, vincinity
of release sites

L. Kendall

---------------------------------------------------------------~------------

�Table 3.

SummarY of Hungarian partridge reports and observations, San Luis Valley, 1971 and 1972 (continued).

Date

Number of Huns

Area

Observed By

August 4, 1971

1 pair

~ mile northwest of second release site, R. Young
farm

C. Grand Pre

August 4, 1971

1 nest with H?:
hatched eggs

1 mile west of first release site, R. Young farm

C. Grand Pre

August 4, 1971

1 pair with 1 chick

1~ miles west of first release site, R. Young farm

C. Grand Pre

August 5, 1971

1 pair with 10 chicks

~ mile southwest of first release site, Hot Creek
Managemen t Area

W. Schuett

August 5, 1971

1 pair

C. Grand Pre and

~ mile east of first release site, Hot Creek
Management Area

C. Grand Pre

August 5, 1971

1 pair with 3 chicks

~ mile south of first release site, Hot Creek
Management Area

C. Grand Pre and
R. Clark

August 5, 1971

2 pair

Vicinity of first release site, Hot Creek Management Area

C. Grand Pre

August 5, 1971

1

~ mile south of first release site, Hot Creek
Management Area

C. Grand Pre

August 5, 1971

3

Vicinity of first release site, Hot Creek Management Area

C. Grand Pre

August, 1971

1 pair with 8 chicks

Vicinity of Rogers headquarters, 1 mile southwest
of Hooper

Rogers

September 14,
1971

1

Vicinity of first release site, Hot Creek Management Area

C. Grand Pre

September 14,
1971

5

~ mile southwest of first release site, Hot Creek
Management Area

C. Grand Pre

------------------------------------------------------------------------------------------------------------------

I
N

0
.pI

�Table 3.

Summary of Hungarian partridge reports and observations, San Luis Valley, 1971 and 1972 (continued).

Date

Numbe r 0 f Huns

Area

Observed By

September 14,
1971

1

Vicinity of second release site, Hot Creek
Management Area

C. Grand Pre

Septembe r 14,

4

~ mile west of first release site, Hot Creek
Management Area

C. GEand Pre

September 15,
1971

6

~ mile southwest of first release site, R. Young
farm

C. Grand Pre

September 15,
1971

4

~ mile west of second release site, R. Young farm

C. Grand Pre

,
o
,

N

September 15,
1971

1

Vicinity of second release site, R. Young farm

C. Grand Pre

September 15,
1971

1

1 mile west of first release site, R. Young farm

C. Grand Pre

December, 1971

1 group of 6 Huns

Vicinity of R. Young's south headquarters

R. Young's hired
hand

December 11, 1971

Tracks of groups of
2 and 4 Huns

Vicinity of first release site, Hot Creek
Management Area

D. Hoffman

December 11, 1971

1 group of 16 Huns

Abandoned farmhouse ~ mile southwest of release
site, E. Stoeber farm

D. Hoffman

June 20, 1972

8 (3 pair and 2 singles)

Hot Creek Managment Unit vicinity of release sites

C. Grand Pre

June 2 1, 1972

5 (1 pair and 3 singles)

Robert Young property vicinity

release sites

C. Grand Pre

June 22, 1972

12 (5 pair and 2
singles)

West of Hooper within 1.5 miles of release sites

C. Grand Pre

July 25, 1972

3 (1 pair and 1 single)

Hot Creek Management Unit vicinity of release sites

C. Grand Pre

---------------------------------------------------------~--------------------------------------------------------

V1

�Table 3.

Summary of Hungarian partridge reports and observations,

Date

Number of Huns,

San Luis Valley, 1971 and 1972 (continued).

Area

Observed By

July 26, 1972

10 (adults)

Robert Young property vicinity of release sites

C. Grand Pre and
R. Young

July 27, 1972

Approx. 45 (8 pair,
25-30 chicks and 1
single)

West of Hooper, within 2 miles of release sites

C. Grand Pre, E.
Ekl und and Ro ge rs

Summer, 1972

Brood of pair and 12
chicks

Farber farm, vicinity of Hooper release site

Farber

September, 1972

Group of 22

Robert Yo ung property vicinity of release sites

G. York, reported
I
to J. Corey
N

October, 1972

Few groups

November, 1972

0f

5 or 6

o
~

Hot Creek Management Area vicinity of release sites

Deer hunters reported to R.
Desilet

4 (2 pairs)

E. Stoeber farm, 6 miles S. of Hooper release site

E. Stoeber

November, 1972

12

McEwen farm, 3 miles S. of Hooper release site

McEwen

December, 1972

Groups of 7, 3, and 2
(total 12 birds)

Vicinity R. Yowg

R. Young

December 11, 1972

5

Whittier

December 12, 1972

Groups of 5 or 6

Vicinity of Hooper release site

Farber, Eklund
and others

December 12, 1972

6 (adults)

Vicinity of Hooper release site

D. Hoffman

December 13, 1972

Tracks of groups of 5
and 10

Vicinity of Hot Creek release sites

D. Hoffman

December 13, 1972

Tracks of 7

Vicinity of R. Young release site

D. Hoffman

release site

farm, 2 miles SW of Hooper release site

Whittier

I

�-207-

From 4 to 10 birds were observed on numerous occasions in the Douglas Lake
release area during February and March by Wildlife Conservation Officer
Crawford. I too, saw these birds several times during the same period. No
observat~ons were reported by local residents, though a ditch rider (Glen
Corbin) did see a few Huns in the immediate vicinity of the release site.
1964 (Swope 1965)
No recorded Hungarian partridge observations were made from March 31, 1964
until July 30, 1964, when 15 young and 1 adult were flushed by Swope near the
Douglas Lake release site. The young were about two thirds grown. During
August and September at least 3 brood observations were made in the Douglas
Lake area. Conservation officer Gurney Crawford saw a covey of 16 Huns, 10
of which were young birds nearly grown, on September 3, 1964. Several ranchers
reported rare to frequent observations of these birds ranging from 1 to 300.
Some of these reports were surely valid, but others are questionable. Graduate
student Ted McKinney observed a covey of 6 Huns near Douglas Lake on January
9, 1965.
In the Round Bottom area southwest of Craig even better reproductive success
was reported. Conservation officer Charles Brown saw about 15 Huns, mostly
young, while checking deer hunters on August 22. About the same time a
rancher (Charles Counts) combining wheat in the area flushed approximately 30
Huns from his grain field. He said most of these birds were young ones.
The Craig area experienced one of the severest winters on record, from the
standpoint of snow depth in 1964-65. Snow depths of from 2 to 5 feet persisted
over most of the area for at least 3 months. The effect of these deep,
persistent snows on the Huns has been undetermined, but heavy losses or total
annihilation are feared. In contrast the Douglas Lake area experienced a very
mild winter.
On March 23, 1965, 130 Huns (6 of these died in transit) were received from the
Oregon State Game COmmission. These were immediately transported to the Round
Bottom site and about half of them released. The remainder were held for
several days and also released. This plant was delayed 6 weeks from initial
arrangements because of the deep snow persisting in the area. Even on the
release date there was still 2 feet of snow covering much of the ground, but
the south facing slopes were bare. Food was scattered around the release site
until the snow melted, however, heavy wet snows continued well into April and
may have been detrimental to these pen raised birds. As happened after the
1964 releases, the birds just seemed to disappear.
Ted McKinney asked 16 states and Canadian provinces for information on Hun
partridge census methods. None of them reported a specific method for observing and counting the birds. Several reported that they count Huns while
making surveys for other upland game species. Searching for Huns in the vast
release site areas certainly gives the field worker the feeling he is hunting
for the proverbial needle in the haystack.
1965

(Swope 1966)

No Huns have been released in the Douglas Lake vicinity since March, 1964.
There were no reported sightings of these birds during the period covered by

�-208-

Segment 19. Numerous roadside checks and walking searches were made within
a 3 mile circle of the release site. Tracks were observed in fresh snow
approximately 2 miles north of the release site that could have been made by
Hungarian partridges, but no verification was possible. Discussions with
several persons who live and work in the area failed to yield evidence of ~un
survival.
Following the March, 1965 release of 124 pen raised Huns from Oregon in the
Craig area, feed was scattered around the release site and checked every few
days by local conservation officers. They reported occasional sightings of
these birds very close to the release site until June. No other authenticated
observations were made, however, a rancher did report a brood he thought to be
Huns, sighted during wheat harvest operations in August.
Because of the relatively small number of Huns released in such a vast area
of homogeneous habitat, with no known concentration area, intensive searches
were not attempted.
1966 (Swope 1967)
Contacts with landowners, ditch riders, and Game, Fish and Parks Department
personnel worki~g in the Douglas Lake area have failed to reveal any Hungarian partridge sightings during 1966. Complete attention has since been
focused on releases made in the Craig area.
Several unconfirmed Hun sightings were reported from the Craig area. A landowner questionnaire was sent to 41 persons in the release vicinity. Seven of
the 24 who responded claimed to have seen Hungarian partridge during 1966.
Though 192 Huns were reported, it is likely that some of these birds were
chukar partridge or sharp-tailed grouse. Personal contacts will be made with
those reporting Hun observations.
1967 (Swope 1968)
The 1966 mail questionnaire resulted in the follOWing information: Lorence
Ellgen who owns land where initial releases were made,observed approximately
6 Huns about the middle of the summer. He felt hawks had taken a heavy toll
of birds released in 1965.
Jim Wilson saw 20 Huns the middle of October in a brushy draw on a lower
north slope of Isles Mountain. This is approximately 2 miles south of the
site of the initial release in 1964.
Bud Bowers reported consistently seeing 3 bunches of Huns on top of Isles
Mluntain during the summer and fall of 1966.
These observations totaled about 100 birds. During the past summer, Mr.
Bowers has frequently seen 50 Huns in his grain fields on Isles Mountain. He
attributes the decline in numbers observed to the taller, denser vegetation in
1967. I made follow up checks on this report. No birds were located, but
tracks and sign that were very probably made by Huns were observed. This
mountain appears to be an ideal location for future releases.

�-209-

George Counts, who farms the land immediately north of Lorence Ellgen claimed
to have seen 8 to 10 Hungarian partridge about the middle of the summer. This
was close to the 1967 release site. Near the first of September, his hired
man saw a brood of 10 Huns approximately 1 mile west of the release site. This
appeared to be a fairly "hot" lead, so I searched the brushy draw between two
wheat fields, near where these birds were reported. A Hungarian partridge
nest, with the intact caps f~om 17 hatched eggs was found. The nest was
located under 2 live sagebrush plants about 25" tall. The draw along which
this nest was found contained intermittent water. A stock pond full of water
was .2 mile to the north. This area also appeared to be another good potential
release site.
Lowry Seely reported seeing several Huns in Axle Basin during the summer.
Huns were seen last spring (1967) in Maudlin Gulch by Elton Gent.

Two

I. P. Bicket, who reported seeing 1 Hungarian partridge during the 1966 grain
harvest, 15 miles north of the initial release site, has seen none since.
Wildlife Conservation Officer Bill Roland and Area Supervisor Harry Dobbs
received reports from land owners in the Round Bottom area that Hungarian
partridge were frequently seen on top of the snow along the main access road
during February and March, 1967. No details were given. Deep, persistent snow
occurred during late winter and early spring of 1967, so it was gratifying to
learn that birds did survive until the warming up period commenced.
Seventy-one pen raised Huns from stock produced at the Fort Collins Wildlife
Research Station were released in Round Bottom area of Moffat County on April
18, 1967.
1968 (Hoffman 1969)
Two-hundred and thirty-six more Hungarian partridge produced at the Fort
Collins Wildlife Research Station were released in the Round Bottom area of
Moffat County on April 30, and May 1, 1968, and 14 from the same source were
released in Routt County northeast of Hayden on April 30, 1968. These releases
were made by Harold M. Swope and other Division personnel.
Twenty follow up questionnaires were mailed in December 1968 to landowners in
Moffat County who answered the 1966 mail questionnaires. The names of the landowners who were sent the 1966 questionnaires had been obtained by Harold M.
Swope from the County Assessor's office in Craig. A sample copy of the personal
letter and questionnaire can be found in Hoffman (1969). These were sent with
a copy of a photograph, a brief description of a Hun and an addressed, postage
paid envelope.
Thirteen of these questionnaires were returned with 4 individuals reporting
seeing approximately 130 Huns during 1968 in groups ranging from a few to over
100 birds. The report of one-hundred Huns is from an area in northwest Moffat
County, which is northeast of Dinosaur National Monument. This area is at
least 55 airline miles from the Round Bottom release sites, so it is extremely
doubtful that these were Hun sightings. The other 30 plus birds reported were
from areas near the release sites.

�-210-

1969 (Hoffman 1970)
On April 19, 1969, 629 additional pen raised Huns from Fort Collins Wildlife
Research Station and Little Hills Experiment Station holding pens were
released at 3 sites on Isles MOuntain, southwest of Craig, Colorado in
Moffat County. Although only 11 Huns were observed during the intensive
field searches conducted in the summer of 1969 in Moffat County, a brood of
at least 4 chicks was located by Division personnel.
Past reports of peaks of Hun numbers sighted by land owners on Isles MOuntain
have coincided closely with wheat harvests. This occurs normally in late
August in MOffat County. Many of these reports of Huns may have been confused with sharp-tailed grouse based upon observations made in 1969. From
27 to 29 sharp-tailed grouse in 4 different groups were observed by D. de
Calesta on or near wheat fields during the period August 16-29, 1969. Three
of these locations were the same or nearly the same as where groups of Huns
had been reported in recent years, mostly during wheat harvests. Only 1 Hun
was observed by de Calesta during this same period. It appears that sharptailed grouse congregate in the wheat fields at harvest time, and may be
difficult to distinguish from Huns by untrained observers. There exists a
strong possibility that many of the past Hun reports on Isles MOuntain were
actually sharp-tailed grouse.
One report of 60 to 80 Huns seen in the Cedar Mountain area northwest of Craig
about October 20, 1969 was secured from hunters by Dale F. Reed. A winter
search of the release sites and surrounding areas including Cedar Mountain
was made by 12 Division employees during the last week of February, 1970. No
Huns were observed, no field sign found and no new leads were secured from
land owners interviewed.
1970 (Hoffman 1971)
Six hundred and seventy-four more pen raised Hungarian partripge were released
in MOffat County at 3 sites on April 24, 1970. This brought the total number
of Huns released in Moffat and Routt counties to 1,788 during the period of
1964-1970.
Intensive field searches were made by W-37-R Project personnel during the
months of June, August and September 1970. Only 10 Huns were observed by Project
personnel, these near 2 of the 3 1970 release sites. Some of the released
Huns were seen in the vicinity of all 3 release sites from May through August
1970 by local residents. Although previously planted areas were searched,
no reports or observations of Huns could be secured, indicating little or no
carry over populations and little or no increases in populations from previous
years. One brood of a hen with 2 chicks was reported by a rancher in the Jubb
Creek area in late August, 1970 in the vicinity of one of the 1970 releases.
Reports of Huns being seen distances of 4 and 5 air miles from 2 of the 1970
release sites were secured.
Specific experiments have not been run to determine the best time for field
releases of Hungarian partridge in Colorado. Traditionally, the early spring
period has been selected because of the advantages of ample food conditions
with the growing green grass and forbs and most of the severe winter weather

�-211-

periods are past. A field release time during the last half of April is a
compromise between the near approach of the egg laying season (early May
under penned conditions) and late snows which may occur as late as early May
in areas selected for Hungarian partridge releases. Birds held in the large
holding pens begin to pair as early as February each year, and most are
believed to be paired by release time. By releasing all birds from a single
holding pen at one site, they have the opportunity to select the same mate
upon release. Handling of the birds is known to cause stress as measured in
weight loss and transporting the birds undoubtedly adds to this stress. This
may delay the nesting activities of the birds somewhat, however, there should
still be ample time for the birds to establish territories and nest following
release. Upon release from the crates, most of the Huns stay in a rather
loose group and appear to start pairing almost instantaneously. Whether they
pair with the same birds as in the large holding pens is not known. Birds
observed later at the release site or at a distance from the release site are
usually observed in pairs, so there appears to be no problem in the birds
finding mates following release.
1971 (Hoffman 1972)
Only 4 reports of isolated sightings of from 2 to 4 Hungarian partridges were
recorded for Moffat County in 1971. These were observed in the spring of
1971. A total of 1,774 Huns were released in MOffat County from 1964 through
1970.
Six hundred and fifty-five pen raised Hungarian partridges were released at
3 sites in the San Luis Valley on April 23, 1971. This constitutes the first
of a planned 3 year stocking effort to attempt establishment of the species in
south-central Colorado.
A total of 7 broods ranging in size from 1 to 14 chicks were observed by
Project personnel or interested landowners on or near the 3 release sites in
the San Luis Valley in August, 1971. In addition, 1 nest containing 8 eggs
was destroyed by a farm tractor, and 1 hatched nest with 12 eggs was found
by Project personnel. Pairs of Huns were reported at distances of 4 to 8
miles from 2 of the release sites, but pairs are known to have remained near
all 3 release sites as well. A minimum of 28 Huns was known to be ranging in
the vicinity of the 3 release sites during the period December 9-11, 1971.
One flock of 16 was observed l~ miles west of Hooper, Colorado on December 11,
1971.
1972 (Hoffman 1973)
Sightings of both adults and broods in 1972 were far less than observed in
1971. A severe drought occurred in this area in the spring and summer of
1972, followed by heavy snows and severe cold during the fall period. These
conditions apparently affected both reproduction and survival, based upon
numba ra of birds observed.
Although movements of Huns of up to 6 miles from the Hooper release site, 3
miles from the Robert Young farm release site, and 1 mile from the Hot Creek
Management Area release site were recorded, birds apparently did not locate
away from the actual release sites. Field surveys made in December, 1972
indicated a small nuclei population existed in all 3 release site areas,

�-212-

including 12 near the Hooper site, 12 near the Robert Young farm release
site, and 15 near the Hot Creek Management Area site. This minimum of 39
Huns for December, 1972 is only slightly more than the minimum of 28 Huns
known to be ranging in the vicinity of the same 3 release sites in December,
1971. Fig. 5 shows a pair of Huns near the release site west of Hooper in
Saguache County in December, 1972.
Five hundred thirty additional pen raised Huns were released in the San Luis
Valley on April 25, 1972 at the same 3 sites where the 1971 releases were
made. Eighty six pen raised surplus breeders were also released at the Hot
Creek Management Area site on July 24, 1972.
1973 (Hoffman, Final)
All remaining pen raised stock held in pens (1,105 Huns) was released at 5
sites in the San Luis Valley from March 20, 1973 to March 24, 1973. Two of
the 3 previously stocked sites received additional Huns but 1 site (Hot
Creek Management Area) could not be reached because deep snow drifts blocked
nhe access road. No follow up checks were made by Project W-37-R personnel
during 1973.
SUMMARY AND CONCLUSIONS
From 1964 through 1973, a total of 4,299 Hungarian partridges (175 wild
trapped, and 4,124 pen rais~d birds) were released in Colorado in 3 main areas.
A total of 135 were released in Larimer County in 1964, 1,774 in Moffat County
from 1964 through 1970, and 2,376 in the San Luis Valley (Alamosa, Conejos
and Saguache counties) from 1971 through 1973. Fourteen were also released
in Routt County in 1968. Results of releases in the 3 main areas are discussed separately as follows:
Larimer County
Frequent sightings of Huns in 1964 in the Douglas Lake area were made following
the release of 135 Huns in January and February, 1964. One brood of 15 young
was observed by Harold M. Swope and at least 3 broods were observed by Gurney
Crawford in the summer of 1964. During 1965 and later years, no further
sightings of these birds were secured, indicating complete disappearance.
Movements of up to 15 miles from the release site were recorded in 1964 by
McKinney (unpublished). This trial release involving wild trapped stock
failed to survive.
Moffat County
Frequent sightings of Huns in MOffat County following releases totaling 1,774
birds from 1964 through 1970 occurred. At least 4 broods were observed or
reported in 1969 and 1 brood was reported in 1970 so some reproduction is
known to have occurred. MOvements of Huns from 4 to 5 miles from the release
sites were recorded.
A mid-winter search of the release sites and surrounding areas made by 12
Division employees indicated few, if any, of these Huns were surviving through
the winter period with no birds observed, no field sign found, and no new

�I

N
t-l

W
I

Fig. 5. A pair of Huns in the San Luis Valley on December 13, 1972. These were part of a group of 12
ranging near the release site, 1 mile west of Hooper, Colorado. (D. Hoffman, photo).

�-214-

leads secured from landowners. During 1971, only 4 reports of isolated
sightings of from 2 to 4 Huns were recorded for Moffat County. Trial releases
involving 40 wild trapped and 1,734 pen raised Huns in Moffat County also
failed to establish reproducing populations.
San Luis Valley
Frequent sightings of Huns were also secured in the San Luis Valley during
1971 and 1972 following releases made. No follow up checks were made in 1973
following final field releases made in March, 1973, because of a decision to
abandon the study. During 1971, 35 separate sightings and observations were
recorded (Table 3) including 7 broods ranging from 1 to 14 chicks. During
1972, 17 separate sightings and observations were recorded, including several
broods (Table 3). Movements of up to 6 miles from the release sites were
recorded for 1971 and 1972.
Late fall surveys made in December, 1971 and December, 1972 indicated only
small nuclei populations of Huns had survived from these respective releases
in the viCinity of the release sites and none in the surrounding areas.
Results of trial releases of 2,376 pen raised Huns in the San Luis Valley
from 1971 through 1973 are very questionable as to the success in establishing
reproducing populations, even though results appear to be better than experienced in the Larimer County and Moffat County releases. Future surveys will
be required to properly evaluate these releases, but the probability that
the species will become established in the San Luis ·Valley is slim.

LITERATURE CITED
Evans, R. L., and W. W. Sandfort. 1960. Ecological appraisal of Hungarian
partridge habitat with reference to adaptation of this species in Colorado. Colo. Dept. of Game and Fish, Denver. Mimeo. 2~8p.
Hoffman, D. M. 1969. Hungarian partridge adaptability. Job Progress Rpt.,
Fed. Aid Proj. W-37-R-22, Work Plan 10, Job 1. Colo. Div. of Game,
Fish and Parks, Denver. p. 87-96.
1970. Stu4y of Hungarian partridge adaptability. Job Prog. Rpt.,
Fed. Aid Proj. W-37-R-23,.WP 10, .Job iL,
Colo. Dfrv,of Game, Fish and
Parks, Denver. p. 139-148.
1971. Study of Hungarian partridge adaptability. Job Prog. Rpt.,
Fed. Aid Proj. W-37-R-24, WP 10, Job 1. Colo. Div. of Game, Fish and
Parks, Denver. p. 49-58.
1972. Study of Hungarian partridge adaptability. Job Prog. Rpt.,
Fed. Aid Proj. W-37-R-25, WP 10, Job 1. Colo. Div. of Wildl., Denver.
p. 39-50.
McKinney, T. D. 1966. Survival, distribution, and reproduction of an introduced gray partridge population. M. S. Thesis, Colorado State University.
Fort Collins. 55 p. Unpub.

�-215-

Sandfort, W. W. 1962. Study of Hungarian partridge adaptability. Job
Comp1. Rpt., Fed. Aid Proj. W-37-R-15, WP 10, Job 1. Colo. Dept. of
Game and Fish, Denver. p. 133-134.
1963. Study of Hungarian partridge adaptability. Job Comp1. Rpt.,
Fed. Aid Proj. W-37-R-16, WP 10, Job 1. Colo. Dept. of Game, Fish and
Parks, Denver. p. 239-240.
Swope. H. M. 1964. Study of Hungarian partridge adaptability. Job Comp1.
Rpt., Fed. Aid Proj. W-37-R-17, WP 10, Job 1. Colo. Dept. of Game, Fish
and Parks, Denver. p. 237-239.
1964.

The Huns are coming.

Colo. Outdoors 13(6) :24-26.

1965. Study of Hungarian partridge adaptability. Job Comp1. Rpt.,
Fed. Aid Proj. W-37-R-18, WP 10, Job 1. Colo. Dept. of Game, Fish and
Parks, Denver. p. 85-87.
1966. Study of Hungarian partridge adaptability. Job Comp1. Rpt.,
Fed. Aid Proj. W-37-R-19, WP 10, Job 1. Colo. Dept. of Game, Fish and
Parks, Denver. p. 61-64.
1967. Hungarian partridge adaptability. Job Comp1. Rpt., Fed. Aid
Proj. W-37-R-20, WP 10, Job 1. Colo. Dept. of Game, Fish and Parks,
Denver. ~ 157-162.
1968. Study of Hungarian partrdige adaptability. Job Prog. Rpt.,
Fed. Aid Proj. W-37-R-21, WP 10, Job 1. Colo. Div. of Game, Fish and
Parks, Denver. p. 65-68.

Prepared by

4.~~

nald M. Hoffman
Wildlife Researcher

��-217-

.April, 1973

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~-------

Project No.

W-37-R-26

Work Plan No.

10

Job Title

Job No.

2

Experimental Breeding of Hungarian Partridge

Period Covered:
Personnel:

Game Bird Survey

April 1, 1972 through March 31, 1973

Lawrence Webster, John Corey, Larry Crooks, Carroll Grand Pre,
David Bowden, Ann Leckler and Donald Hoffman.

ABSTRACT

During 1972, a total of 2,010 eggs were produced by 52 pairs of Huns, of
which 1,640 were fertile, and 1,410 hatched. This was the highest number
of eggs and chicks produced in any single year for the 7 year period from
1966 through 1972. Only one group of 5 pens of selected breeders produced
significantly (P~0.05) more total eggs per hen (X = 53.60) than the 12 pens
of random bred controls (X = 39.00) for a gain of 14.60 eggs per hen. All
other matings tried in 1972 produced nonsignificant (P«0.05) results in
regard to differences in total eggs produced, percent fertility of eggs,
and percent hatchability of eggs. Excellent survival for the first 7 days
(98.56) percent) and for the first 21 days (96.89 percent) was recorded for
the young Huns in 1972. Overwinter survival of Huns was also excellent
(91.44 percent) with 1,121 live birds tallied in March, 1973, out of the 1,226
released into the holding pens in August, 1972. All remaining breeders and
young were used for field releases during the period March 20 through March 24,
1973. Twenty seven pairs of wing-clipped Huns were released in 2 large open
topped quadrats and allowed to incubate their eggs in order to study nesting
success under semi-natural conditions. A total of 15 nests were located,
of which 6 hatched (40 percent). Clutch sizes range from 1 to 19 with an
average of 11.2 eggs.

�-218-

RECOMMENDATIONS

With all rema~n~ng pen raised Hungarian partridges released in the San Luis
Valley in March, 1973, it is recommended that this job be terminated and data
summarized in a final report during Segment 27.

�-219-

EXPERIMENTAL

BREEDING

OF HUNGARIAN

PARTRIDGE

Don ald M. Ho ffman

Total egg production from 52 pairs of Hungarian partridge breeders in 1972
was 793 eggs higher than the total produced by 51 pairs in 1971 (2,010 eggs
in 1972 compared with 1,217 in 1971). Numbers of eggs laid and numbers of
birds raised until field release time reached a peak in 1972 for the 7 year
period 1966 through 1972. The 1,105 pen raised Huns released in the San
Luis Valley during the period March 20 through 24, 1973, plus 86 older
breeders released on July 24, 1972 (total 1,191) terminated Colorado's
efforts t·o establish populations through use of pen raised stock.

P. S. OBJECTIVE
To develop

game farm production

techniques

for Hungarian

part ridge.

SEGMENT OBJECTIVES
1.

To compare numbers of Huns surviving overwinter with numbers
in holding pens containing various bird densities.

2.

To compare numbers of total eggs laid, percentage fertility of eggs, and
percentage hatchability of eggs produced by breeders consisting of:

3.

released

a.

Twelve pairs of 1971 hatched young from pens 5 and 25 in 1971.
These are the Fl progeny from the highest producing adults (1969
hatched) .

b.

Twelve pairs of 1971 hatched young from 1971 pens 11, 14; 17 or 18.
These are F2 p ro geny produced by crossbreeding hens from 1970 pens
3 or 9 with Winchester Farm Cocks.

c.

Twelve pairs of 1971 hatched

random bred controls.

To determine the value of rearing Huns under semi-natural conditions,
using wing clipped, 1971 hatched random bred stock in large open topped
pens with adequate nesting cover.

METHODS
Survival

AND MATERIALS'

Tests

The detailed arrangement of the Hungarian partridge holding pens (rearing
and conditioning runs) is shown in Fig. 1. With the large number of Huns
on hand, holding pens Ql, Q2, Q5 and Q6, formerly used for mountain quail

�-220Covered
Brooder

Shelter

1

Houses

/~

L--.-...l-_-f-_--''===~ ,12_' -+--"-' --...I._---J4'~8'-1
3

6

r

J

-0
N

4

2

;

.~"
"

N

o
N
N

SCALE
•.4'1

~ 15.5

="- ,- -'5-5--j--'--- .

---35

o

-----

t----------.--I05~--------Fig. 1.

Hungarian

"'::

Por tr idqe

'

HoldinCJ Pens

1

15

30

�-221-

(Fig. 2) were also used for holding Huns. Numbers of Huns placed in the
various holding pens on August 22, 1972 were tallied and an accurate count
of birds recovered from the various pens on March 19-23, 1973 was made.
Percentage of overwinter survival was then calculated.

Hatchability

Egg Production, Fertility of Eggs and
of Fertile Eggs by Various Sources of Breeders

Fig. 3 shows the pen arrangement for the 1972 test series and Table 1 lists
the pen schedule for 1972. Egg production was compared by 3 main sources
of stock (12 pens each), and then the 3 main sources of stock were subdivided
into 5 known sources (with varying numbers of pens) for statistical analyses.
Standard t-tests were used by Dr. David C. Bowden to test for significant
(P&lt; 0.05) differences in total eggs laid, percentage fertile eggs produced,
and percentage hatchability of fertile eggs by known sources of breeding
stock. Pens of the 3 main sources of stock were alternated through the
entire group of 36 ground floored pens to reduce the effect of pen location.
All birds were fed a standard 22 percent game bird breeder ration.

Rearing Huns Under Semi-natural

Conditions

Fifty four (27 pairs) of 1971 hatched pen raised Huns of random bred source
were wing clipped, leg banded and released in 2 large open topped quadrats.
These quadrats each measured 370 ft by 405 ft each. The east quadrat was
stocked with 15 pairs of Huns on March 13, 1972 and the west was stocked
with 12 pairs on March 15 and 16, 1972.
Each quadrat contained 12 vegetative plots of 6 different species or combination of species left over from a pheasant nesting study.
These included 2
each of alfalfa, crested wheat grass , intermediate wheatgrass, tall wheatgrass, bromegrass, and alfalfa-crested wheatgrass mixtures (Fig. 4). All
cover plots were windrowed after the nesting season to facilitate nest
searching.
Prior to windrowing, all birds were removed from the quadrats
and released in the surrounding areas to prevent unnecessary death loss.
Efforts were made to control predators, particularly feral cats and horned
owls.
Two separate nest searches were completed to evaluate nesting success.
The
initial nest search was made during the period July 10 through 13. Each plot
was systematically searched by use of laths used to divide each plot into
narrow strips.
The final nest search was made on July 17, prior to windrowing
the vegetation in the quadrats.
Nests were marked, recorded as to total
number of eggs, fate of nest, and estimated age of embryos.
These were then
removed to avoid duplication of effort.

RESULTS AND DISCUSSION
Table 2 lists a summary of Hungarian partridge propagation by the. Colorado
Division of Wildlife from 1966 through 1972.
Improvement has been accomplished both in increasing numbers of chicks hatched and rearing of young

�-222-

I(

20'

~

~e'-)re'---7f~

i

-Q)

'N

\.
5

3

I

1

\
6

4

2
,

-

Q)
It)

Scole:

o

I": 15 ft.
15

GO'

Fig. ::&gt;.
M0 u n t a In
partridges

0 u a i I H0 I din Q Pen S ( Pens 1,2,5,
during 1972-73).

and 6 were used for Hungarian

30

�-223-

N

31

32 33

34 35

~

O£ 62 82

L2

92

75'

6'4"

J

I'

,

eo

-

Ii)
J'/l

Ground -f loored Series

37E813914041 42 43 4445 4647 4c
w _ v __ .'\v -'--'\11' _'-, l&gt;

'-

Wire-floored

~)
rt)\

A-2

1"/

N
N

-

L

\

49,9C 51

/

'--

~

0

01

N--",

,

&lt;D
-

_v.

I

(7)

-

\j./

It

SCALE 8

/

&lt;X)
I"-

-

~
.:

N

en
-

(J)

Q

N

N

-

-v

/

Series

I

I

I

O' 5' 10'

25'

-.J
/

"-

A-3 /

m

Fig. 3.
Detailed ArranQement

&lt;D
/

....

Partridge

/-

to

-

0

v
-

/

....

-

(N

rt)

t-16'----o

16 ---i

48'

I

cD
.!,

Breeding

of

Pens.

HunQorian

,

�-224-

until field release time. Numbers of chicks hatched has increased from 236
in 1966 to 1,410 in 1972, and percentage of survival of young birds until
field release time has been increased from a low of 49.32 percent in 1967 to
a high of 91.44 percent in 1972.

Table 1.

Pen schedule

for Hungarian

Pen
Number

Year
Hatched

Hatch
Number

1

1

34
35
36
37

1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1971
1969

1
1

38

1969

1

2
3
4
5

6
7
8
9

10
11

12
13
14
15
16
17
18
19
20
21

22
23
24
25
26
27
28
29
30
31

32
33

1
1
1
1
1
1
1

1
1

1
1

2
1
1

1 and 2
1
1
2
1
1

2

1 and 2
1
2
1 and 2
1

2
1 and 2
1
2

1 and 2
1

2
1 and 2

partridge,

1972. !/

Source

Pens 5 and 25
Pens 11, 14, 17 and 18
Run a f the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run of the mill
Pens' 5 and 25
Pens 11, 14,:17 and 18
Run of the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run of the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run of the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run of the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run a f the mill
Pens. 5 and 25
Pens 11, 14, 17 and 18
Run of the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run of the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run of the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run a f the mill
Pens 5 and 25
Pens 11, 14, 17 and 18
Run a f the mill
High producing adults from Pen
3 in 1970
High.producing adults from Pen
9"in 1970

----------------------------------------------------------------------------

�-225-

Table 1.

partridge,

1972l../(continued) .

Pen schedule

for Hungarian

Pen
Number

Year
Hatched

Hatch
Number

39
40

1971
1969

1
1

Run of the mill (spare)
High producing adults from Pen 31 in

41

1969

1

High producing

Source

1970
adults from Pen 32 in

1970
42

1970

1

High producing

adults from Pen 4 in

1971
43

1970

1 and 2

44

1970

1

45

1970

1

46
47
48
49
51

1971
1971
1971
1971
1971
1971
1971

2
2
1
2
1
1 and 2
1 and 2

High producing adults from Pen 14
in 1971
High producing adults from Pen 24
in 1971
High producing adults from Pen 36
in 1971
Pens 5 and 25 (spare)
Pens 11, 14, 17 and 18 (spare)
Run of the mill (spare)
Pens 5 and 25 (spare)
Run of the mill (spare)
Pens 25 and 18 (spare)
Pens 5 and 17 (spare)

McCarty
McCarty

(W)

l/ Birds

in all pens were fed a standard

(E)

22 percent game bird breeder

ration.

Numbers of eggs produced per hen have shown increases from a low of 22.06 in
1966 to a high of 38.65 in 1972. These accomplishments have been secured
through, (1) the use of modern game bird farm techniques, (2) improvements to
the environments, and (3) application of artificial selection pressures
during the period 1970 through 1972.

Survival

Tests

Table 3 lists survival of young Hungarian partridges, ages 1 to 21 days, for
the 5 hatches in 1972. Excellent percentage survival for the first 7 days
(98.56 percent) and for the first 21 days (96.89 percent) was recorded.
Table 4 lists results of the overwinter survival tests completed in 1973.
Densities of birds varied from 13.54 square feet of ground space per bird to
102.40 square feet of ground space per bird. Highest survival (98.53 percent)
occurred in a group of 68 Huns from hatch number 5 in 1972 held in a pen with
14.01 square feet of ground space per bird. Lowest survival (66.67 percent)
occurred in a group of 15 adult breeders, 4 of which were 1969 hatched and 11
were 1971 hatched.
Of these 15 adult breeders, 2 1969 hatched and 3 1971
hatched Huns died ove rwtnt e r, Overall survival of 91.44 percent was the
highest recorded for the 7 year period 1966 through 1972.

�-226·-

Plot Number

Species of Vegetation

Planted

I

Smooth bromegrass

(Bromus inermis)

2

Alfalfa

sativa)

3

Crested wheat grass (Agropyron

4

Intermediate

5

Tall wheat grass (Agropyron

6

Alfalfa-crested

Fig. 4. Diagramatic
enclosures.

(Medicago

arrangement

cristatum)

wheat grass (Agropyron

intermedium)

elongatum)

wheat grass

of the vegetative

plats in the east and west

�Table 2.

Summary of gray partridge
(fJ

.~
cU

"t:l

Q
(l)
::I::

....:l

~

(l)
"t:l
(l)
(l)

~
cU

(l)
:&gt;&lt;

(fJ

00
00
I'tl

~
~

.

.-i

z

H

cU

.j..J

0

0

(fJ

00
00
I'tl

.
z
.

propagation,

1966-1972.
(l)
.-i

(fJ

-e

.j..J
(l)

~

U)

(fJ

.-i

(fJ

,-...

(fJ

.~
.j..J

(l)
.-i
'.-4
.j..J

&amp;

.c ~

H

&amp;

.j..J
Q
(l)
(J

0

0

(l)
.-i

.~
.j..J
~

(l)
4-l
Q

~

:J
&amp;
U 0

00
00
I'tl

&lt;tl::I::

~ ~

~;§

0

Z

Z

Z

~
(l)
e,

0

.~
Q

.

~

.

(fJ

"t:l 0.
(l) '.-4
(JU
.j..J

cU
::I::.-i
(J
• Q

OH

.c

"5.j..J
cU

(fJ

::I:: 00
00
.j..J1'tl
Q
(l).-i
(J cU
~ .j..J
(l) 0

(J (fJ
.j..J 00
cU 00
::I::1'tl
.j..J (l)
Q.-i
(l) -.-4
(J .j..J
(l)

Z'-'

p..,~

(l)

-e

cU

(l)
.-i
0.
0.
•.-4

p..,

~

U

0

0

0

"t:l
(l)
0.
0.
'.-4

.

~ ~

p..,H

~
~
-e

.

"t:l
(l)
.-i
0.
0.
-.-4

~

U

~

.

Z

Z

Z

'";i1

:::;1
"t:l
(l)
(fJ

:&gt;
.~
~
;:I

U)

.j..J
Q
(l)
(J

-.-4

.j..J

~

(l)
(J

(l)

0

Z

p..,

~

p..,

.

P

~

(l)

19662:,/ 341/

750

22.06

--

699

255

444

63.52

236

33.76

53.15

--

--

--

--

121

51.27

1967

50

1203

24.06

46

1157

281

876

75.71

511

44.17

58.33

35

264

64

12.52

252

49.32

1968

51

1783

34.96

64

1719

376

1343

78.13

1003

58.35

74.68

34

306

11

1.10

614

61.22

1969

52

1818

34.96

28

1793

344

1449

80.81

945

52.71

65.22

53

451

64

6.77

674

71.32

1970

52

1147

22.06

35

1112

181

931

83.72

757

68.08

81.31

22

152

34

4.49

675

89.17

1971

51

1217

23.86

64

1153

274

879

76.24

689

59.76

78.38

16

176

35

5.08

584

84.76

1972

52

2010

38.65

77

1933

293

1640

84.84

1410

72 .94

85.98

65

165

90

6.38

1121

91.44

l/ Survival

of young birds until

following

2/ Data deleted from reports where blanks

spring period when

field releases were made.

are shown

3/ Sex ratio 1:1 except for 1 community pen with 11H:2C.

Sex ratio of 1:1 used from 1967 through

1972.

I

N
N

"
I

�Survival

of Hungarian

Hatch
Number

Date
Hatched

Number Chicks
Battery
BrooderBrooder
House

1

6-12-72

159

29

0

2

0

0

188

186

100.00

98.94

2

6-27-72

326

25

10

15

0

0

341

336

97.15

95.73

3

7-11-72

377

0

6

10

-

-

371

367

98.41

97.35

4

7-31-72

336

0

3

13

-

333

323

99.11

96.13

5

8-15-72

68

0

0

1

-

-

68

67

100.00

98.53

Table

3.

partridges,

ages 1 to 21 days,

1972.

Number Chicks Lost
Battery
House
21 da
7 da
21 da
7 da

Number
Chicks Survived
21 days
7 days

First

Percent Survival
First 21 days
7 days

I

tv
tv
00
I

Totals
Average

1266

54

19

41

0

0

l301

1279
98.56

96.89

�Table 4. Survival of Hungarian partridge during late summer, fall, and winter periods according to various densities
of birds and sizes of pens, 1972-73.

Pen and
Hatch

Size
(Ft.)

Total
Sq. Ft.

Date Birds
Counted In

Age
(Approx.)

Number
Birds In

Sq. Ft.
Per Bird

Date Birds
Recovered

Number
Birds Out

Percent
Survival

A-2 &amp; A-3
(adults)

16 X 96

1,536

8-22-72

Adults

15

102.40

3-23-73

10

66.67

1 &amp; 2
1972 (4)

35 X 240 +
llX11

8,521

8-22-72

3 weeks

315

27.05

3-19-73

265

84.13

3
1972 (5)

35 X 20 +
llX11+
12 X 11

953

8-22-72

1 week

68

14.01

3-19-73

67

98.53

35 X 220

7,700

8-22-72

7 weeks

286

26.92

3-20-73

273

95.45

5 &amp; 6
1972 (3)

35 X 240 +
llXll

8,521

8-22-72

6 weeks

351

24.28

3-23-73

326

92.88

Q1 &amp; Q2
1972 (1)

20 X 76 +
8 X 8

1,584

8-22-72

9 weeks

117

13.54

3-23-73

109

93.16

Q5 &amp; Q6
1972 (1&amp;2)

20 X 76 +
8 X 8

1,584

8-22-72

9 weeks

74

21.41

3-23-73

71

95.95

4

I

N
N
\0 .

1972 (2)

Totals
Average

1,226

1,121
9l.44

I

�-230-

Egg Production, Fertility of Eggs, and
Hatchability of Fertile Eggs by Various Sources of Breeders
Table 5 lists egg production, fertility of eggs, and ,hatchability of fertile
eggs by 3 main groups of breeders on test during 1972. The main groups
involved
known but combined ancestries.
Group A consisted of 1971 hatched
Fl progeny from pens 5 and 25 (and reciprocal crosses).
These were the 2
highest producing pens from the closed flock maintained at the Fort Collins
Wildlife Research Station.
Group B consisted of 1971 hatched F2 progeny
strain crosses between selected high producing birds from the closed flock
maintained at the Fort Collins Wildlife Research Station and the Winchester
Farm st rain. Group C consisted of 1971 hatched F2 progeny of the random bred
birds used for controls.
Group A birds produced 44.33 eggs per hen in 1972
compared with 38.42 for Group B, and 39.00 for Group C. Percent fertility
and percent hatchability was lower for Group A birds than either Group B or
C (which were about equal).
Birds in the various groups indicated in Table 5 were then sorted by individual ancestry as indicated in Table 6, and placed into Source Codes 9 through
14. These source code numbers were those assigned for computer analysis for
the 1972 tests. Pens 5, 25, 11, 14, 17 and 18 listed in Table 6 are the same
as pens 5, 25, etc. listed in Table 5.
Results of t-tests by Dr. David C. Bowden, Department of Statistics, Colorado
State University, comparing total eggs laid, numbers of fertile eggs, and
numbers of fertile eggs which hatched by source code of stock (Table 7)
indicated that significantly (P&lt; 0.05) more total eggs were laid by only 1
group of breeders (5 pens) on test, in comparison with the random bred
controls (12 pens).
These 5 pens laid an average of 53.60 eggs per hen
compared with only 39.00 eggs per hen for the controls for a gain of 14.60
eggs per hen. All other matings tested in 1972 produced nonsignificant
(P&lt; 0.05) results in regard to differences in total eggs produced, percent
fertility of eggs, and percent hatchability of eggs.
Rearing of Huns Under Semi-natural

Conditions

A total of 15 Hun nests were located in the open topped pens where 27 pairs
of wing clipped breeders were released in March 1972.
Fourteen of these nests
were found on the first nest search and 1 on the second.
By quadrat, 9 nests
were found in the east quadrat and 6 in the west (Fig. 4).
Of the 15 nests recorded, 6 were hatched successfully, (40 percent) and 9
were abandoned or unsuccessful (60 percent) as indicated in Table 8. Three
of the 6 successfully hatched nests contained a total of 37 eggs, of which 24
hatched, for an average hatchability of 64.86 percent.
The remaining three
of the 6 successfully hatched nests contained an unknown number of eggs. Of
the 9 abandoned or unsuccessful nests, 6 were abandoned for unknown reasons,
and 3 were thought to have been abandoned as a result of human disturbance.
Three of the 15 nests found contained unknown clutch sizes because the nests
had deteriorated prior to discovery ..The average clutch size of the remaining
12 nests was 11.2 eggs, with a range of 1 to 19 eggs.

�-231Table

5.

by source,

Egg production

Source

Total
Eggs

Pen
Number

Group A - Crosses
Pens 5 X 25
and

25 X 5
( Reciprocal)

Number
Fertiles

(of

small

Percent
Fertile

Number
Fertiles
Hatched

Percent
Fertiles
Hatched

size)

within

a closed

flock

43
37
14
60
52
46
18
61
48
53
55
45

42
37
13
59
48
45
17
58
46
51
54
42

33
33
8
59
46
26
14
54
38
42 .
46
39

78.57
89.19
61.54
100.00
95.83
57.78
82.35
93.10
82.61
82.35
85.19
92.86

29
31
1
57
42
10
13
43
22
38
38
31

87.88
93.94
12.50
96.61
91.30
38.46
92.86
79.63
57.89
90.48
82.61
79.49

532
44.33

512
42.67

438
36.50

83.45

355
29.58

75.30

46
53
23
34
20
39
20
49
15
62
66
34

44
51
23
34
20
38
19
46
15
58
65
33

36
50
21
22
17
34
18
44
15
57
62
32

81.82
98.04
91.30
64.71
85.00
89.47
94.74
95.65
100.00
98.28
95.38
96.97

31
45
21
17
15
26
15
39
13
43
48
28

86.11
90.00
100.00
77.27
88.24
76.47
83.33
88.64
86.67
75.44
77.42
87.50

461
38.42

446
37.17

408
34.00

90.95

341
28.42

84.76

28
30
32
40
35
41
28
34
52
68
31
49

27
30
31
40
34
39
26
34
52
67
31
45

19
30
31
39
33
28
25
31
51
60
31
32

70.37
100.00
100.00
97.50
97.06
71.79
96.15
91.18
98.08
89.55
100.00
71.11

18
27
27
34
32
13
24
28
41
58
28
31

94.74
90.00
87.10
87.18
96.97
46.43
96.00
90.32
80.39
96.67
90.32
96.88

468
39.00

456
38.00

4lO
34.17

90.23

361
30.08

87.75

crosses

02
05
08
11
14
17
20
23
26
29
32
35

Subtotals
Averages
Group C - Random Bred

03
06
09
12
15
18
21
24
27
30
33
36
Subtotals
Averages

Total
Eggs
Set

01
04
07
10
13
16
19
22
25
28
31
34

Subtotals
Averages
Group B - Strain
Pens 11 or 14
X 17 or 18
and
17 or 18
X 11 or 14
(Reciprocal)

1972.

Controls

�Table

6.

Ancest ry of Hungarian

part ridges

on progeny

testing

program,

1972.

Ancestry

Year

Source
Code

Number
Pens

1972

9

12

10

Year
Hatched

Cocks

Year
Hatched

F2 - FCWRS (Rb ).!.I

1971

F2 - FCWRS (Rb)

1971

7

F1 - Pen 1125 (FCWRS)

1971

F1 - Pen 115 (FCWRS)

1971

11

5

F1 - Pen 115 (FCWRS)

1971

F1 - Pen 1125 (FCWRS)

1971

12

6

F2 - Pen 1111 (H-FCWRS X
C-Win. )

1971

F2 - Pen 1117 or 18
(H-FCWRS XC-Win.)

1971

Hens

I

N

4

13
Subtotal'

1/

Rb

34

random

bred

controls.

F2 - Pen 1114 (H-FCWRS X
C-Win. )

1971

F2 - Pen 1118 (H-FCWRS
XC-Win. )

1971

w

N
I

�Table 7. Results of t-tests comparing total eggs laid, numbers
hatched by source of breeding stock. 11

Source
Code

Number
Pens

Source
Code

Number
Pens

of fertile eggs, and numbers

Significant Difference
Indicated In

of fertile eggs

Conclusion

9

12

Compared with 10

7

9

12

Comp ared with 11

5

9

12

Compared

with 12

6

Non-significant

9

12

Compared with 13

4

Non-significant

Non-significant
Total eggs laid
t = -3.6591 (15df)

Source Code 11 (X = 53.60)
laid significantly
(P 0.05) more total eggs
than Source code 9
(X = 39.00)
I

N
W
W
I

1/ See Table 6 for ancestry

of breeding

stock by Source

Code.

�Table 8.
Nest
Number

Clutch sizes found in 15 hatched or unsuccessful Hungarian partridge nests, 1972.

1

2

3

4

5

6

1

7

8

9

10

Clutch Size
11
12

13

14

15

16

17

18

19

20

Unk.

1H

Total
Eggs

7

2

1H

9

3

lU

9

4

IH

1

5

lU

6

14

lU

10

7

I

lU

8

19

+:--

lU

9

11
lU

12

10

lU

11

lU

12

lU

14

13

lH

14

IH

12

15

Total

IH
1

0

0

0

0

Average Clutch Size 11.2
Key:

H = Hatched = 6 (40%)
U = Unsuccessful = 9 (60%)

0

7

0

18

10

11

N
W

24

0

28

0

16

16
0

0

19

0

--

134

I

�-235-

As of July 31, 1972, a total of 17 casualties were found in and near the
quadrats, including 4 adult hens, 4 adult cocks, 2 unclassified adults, and
7 young Huns. Nine of these dead birds were found in the east quadrat, 6
in the west, and 2 on the adjacent grounds.
In each case, the cause of death
was unknown, since the carcasses had badly deteriorated.
Routine observations by station personnel indicated approximately 20 to 30
Huns resulting from this study ranged on or near the Fort Collins Wildlife
Research Station from late July through September, 1972. By late January,
1973, a group of 8 Huns were still being observed intermittently on the
Station grounds.

Prepared

by
Wildlife

Researcher

��-237-

State of

April, 1973

COLORADO
------~~~~~~---------

Project No.

W-37-R-26

Work Plan No.

15

Job Title:

Job No.

2

Study of Mountain Quail Adaptability

Period Covered:
Personnel:

Game Bird Survey

April 1, 1972 through March 31, 1973

Ronald B. Arant, Tom W. Barnes, john F. Corey, Alexander Hamilton,
Donald E. Lengel, Gary T. Myers, Dwight E. OWens, Robert W. Paintin,
Glenn E. Rogers, John Serra, Thomas J. Sherrill, J. Allen White,
Ann Leck1er and Donald M. Hoffman.

ABSTRACT

A total of 574 mountain quail were released in 2 different areas of Colorado
from 1965 through 1973. Of these, 372 were wild-trapped birds received from
California and Oregon in wildlife trades and 202 were pen-raised at the Fort
Collins Wildlife Research Station. By area, 397 were released in Mesa County
on the west side of the Uncompahgre Plateau and 177 were released in Las Animas
County on the Spanish Peaks Management Area.
One confirmed movement of 25 airline miles distance from a release site was
recorded after three mountain quail were confiscated from hunters in Mesa
County by Gary Myers. Other reported movements of 15, 7, 3~, 3 and 1 miles
from release sites were recorded during the period 1966 through 1972.
The status of mountain quail in Colorado is very questionable because (1)
numbers observed and reported since 1966 have been small relative to numbers
released, (2) no instance of reproduction has been recorded, and (3) consistent
observations for anyone locality are lacking. The probability that the species
will become established in Colorado is slim.
Game farm propagation of mountain quail was terminated and all remaining stock
released on March 20, 1973.

�-238-

RECOMMENDATIONS

Because relatively poor success was attained in propagating mountain quail
at the Fort Collins Wildlife Research Station since 1967, due to traits which
appear to be inherent with the species, game farm propagation was terminated
and all remaining stock released on March 20, 1973.
If further releases of
mountain quail are made, wild-trapped
stock should be obtained through wildlife trade agreements with other states.
Wildlife Conservation Officers and other Division of Wildlife personnel working
in the areas of past releases have been requested to submit future sightings
and observations of mountain quail to the Research Section.
It is recommended
that this job be terminated with this final report.

�-239-

STUDY OF MOUNTAIN

QUAIL ADAPTABILITY

Donald M. Hoffman

During the 1960's, the State of Colorado became interested in extending
populations of upland game birds by introducing exotic species.
An
appraisal of habitat by Rogers (1965) indicated selected areas of Colorado
contained suitable food, terrain, and climate for establishing mountain
quail populations.
The area selected for initial releases was the Indian
Creek drainage in Mesa County on the west side of the Uncompahgre Plateau
but he concluded that several other mountainous areas would also be available.
Releases of 397 mountain quail on Indian Creek in Mesa County were made
with 372 wild-trapped birds from California and Oregon secured through
wildlife trade agreements and 25 surplus breeders held at the Fort Collins
Wildlife Research Station from 1965 through 1970. Two releases of 87 and
90 mountain quail in 1972 and 1973 were made on the Spanish Peak Management
Area in Las Animas County from pen-raised stock reared at the Fort Collins
Wildlife Research Station.
This study was initiated in 1967 to (1) introduce mountain quail in selected
areas, and (2) determine success of introductions in establishing reproducing
populations.
The study was conducted from 1966 through 1968 by Glenn E.
Rogers and by Donald M. Hoffman from 1969 through 1972.

P. S. OBJECTIVE
To determine the ability of mountain quail to establish reproducing populations in areas of mixed shrub and pine in selected sites in Colorado.

METHODS

AND MATERIALS

An appraisal of mountain quail habitat (Rogers 1965) indicated that selected
areas of Colorado appeared suitable in regard to food, terrain, and climate
for mountain quail.
Prior to an initial release of 143 mountain quail on Indian Creek in Mesa
County and release of 16 in holding pens at the Fort Collins Wildlife Research
Station for propagation purposes, a job was initiated to (1) release mountain
quail in selected areas and (2) to determine the success of these releases
(Rogers 1966).
All field releases, with the exception of 90 birds released on March 20,
1973, consisted of transporting birds to the selected release site in State

�-240-

vehicles and releasing the birds as quicky as possible in hard type releases.
Scratch grain was scattered prior to releasing 25 birds on June
23, 1970 in Mesa County and 87 on April 15, 1972 in Las Animas County in
an attempt to hold the birds in the release areas.
There was no indication
that the birds used this, however.
On March 20, 1973, 30 mountain quail
were placed in a temporary holding pen for one week in Las Animas County
and 60 were released outright, following the scattering of scratch grain
in the vicinity.
Early methods to determine success of releases consisted of (1) use of
educational media (radio and TV) to inform people in the vicinity of the
releases and request sighting information, (2) contacting of Wildlife
Conservation Officers, ranchers, sportsmen and personnel of State and
Federal agencies requesting information on numbers and locations of birds
observed, and (3) field searches of release sites and areas where sightings
were recorded.
All reports and observations were recorded on maps and
in tables to evaluate the degree of success in establishing the species
(Rogers 1966).
From 1969 on, methods for evaluation of success were quite
similar except the use of educational media to inform people of releases
was not used since little earlier response was secured.

RESULTS

AND DISCUSSION

Field Releases
A total of 574 mountain quail were released in 2 different areas of Colorado
from 1965 through 1973. Table 1 summarizes all releases by date, numbers of
birds, source of birds, and release areas.
Fig. 1 shows the map locations
and numbers of mountain quail released in Mesa County from 1965 through 1970
and Fig. 2 shows similar information for Las Animas County for 1972 and 1973.
Three hundred seventy-two wild-trapped and 25 pen-raised mountain quail
(total 397) were released in Mesa County on the west side of the Uncompahgre
Plateau from 1965 through 1970 (Table 1). In addition, the State of Utah
released approximately 100 mountain quail in Willow Basin on the north end
of La Sal Mountain in August, 1968 (pers. comm. D. Nish to D. Hoffman).
The Willow Basin site in Utah is approximately 15 to 20 miles southwest of
Gateway, Colorado (Fig. 1).
Eighty-seven pen-raised mountain quail were released in Vigil Canyon on April
15, 1972, and all remaining (90) mountain quail were released in Burro Canyon
on the Spanish Peaks Management Area in Las Animas County on March 20, 1973
(Fig. 2). Fig. 3 shows the habitat in the vicinity of Vigil Canyon site
and Fig. 4 shows the release in Burro Canyon.

Success of Introductions
Figures 1 and 2 show map locations and numbers
through 1972. Results of surveys to determine
summarized (Table 2) by years as follows:

of birds observed from 1965
the success of releases are

�Table 1.

Summary of mountain quail field releases.

Date

Number of
Birds

Source of
Birds

Release
Area

Reference

Aug. 19 and 25, 1965

1431:./

California
(wild-trapped)

Mesa County, Indian Creek
Sec. 27, T51N, Rl7W.

Rogers (1966),

Aug. 20, 1966

163

California
(wild-trapped)

Mesa County, Indian Creek,
approx. 4 miles west of 1965
releases.

Rogers (1967)

Jan. 25, 1968

66

Oregon
(wild-trapped)

Mesa County, Indian Creek
approx. 1 mile east of west
Uncompahgre National Forest
boundary.

Rogers (1968)

Mesa County, South Fork Mesa
Creek, Sec. 17, T49N, R16W

Hoffman (1971)

FCWRS
(surplus breeders)

June 23, 1970

25

Sub-total

397

April 15, 1972

87

FCWRS
(pen raised)

Las Animas County, Vigil Canyon,
Spanish Peaks Management Area
Sec. 11, T32S, R67W.

March 20, 1973

90

FCWRS
(pen raised)

Las Animas County, Burro Canyon
Spanish Peaks Management Area
Sec. 7, T32S, R66W.

Sub-total

177

TOTAL

574

I

N

~
•....

l/ Sixteen additional mountain quail from these shipments were taken to Fort Collins Wildlife Research Station
for experimental propagation studies. On January 18, 1970, 12 wild-trapped mountain quail from Oregon were
added to the breeding stock at the Fort Collins Wildlife Research Station. These constitute the source of our
breeding stock.

I

�Table 2.

Summary of mountain quail reports and observations, 1965-1972 (continued).

Date

Numbers of
Mountain Quail

Area Observed

Distance (airline miles) and
Direction from Release Site

Observed By

April 20, 1972

3

Las Animas County,
R. Dochter tract
hdq., Burro Canyon

3 miles SE of Vigil Canyon release site.

R. Paintin

Early May, 1972

4

Las Animas County,
B. Haught Ranch
hdq ,, Wet Canyon

3 miles SW of Vigil Canyon release site.

B. Haught

Mid May, 1972

5

Las Animas County,
Oberhausler tract
hdq., Burro Canyon

3-1/2 miles SE of Vigil Canyon
release site.

R. Paintin

Las Animas County,
Vigil Canyon

1 mile NE of Vigil Canyon re~
lease site

I

N
.p-

-...]
I

Mid August, 1972

1

!/ Willow Basin was the site where Utah released approximately 100 mountain quail in August, 1968.

J. Serra

�-248-

1965
One indefinite report of a mountain quail sighting of 14 mountain quail
in August, 1965 was secured from a California hunter following an initial
release of 143 wild-trapped birds in Mesa County (Rogers 1966).
In the
late fall of 1965, Mr. J. D. Sloggett of Grand Junction, Colorado sighted
an unknown number of mountain quail on Blue Mesa (3 miles southeast of
the 1966 Indian Creek release site (Rogers 1967).
1966
On November 21, 1966, Gary T. Myers confiscated 3 mountain quail that had
been killed by hunters on November 19, 1966.
The hunters stated there were
12 birds in the covey and more were heard calling in the area.
These quail
were killed on the East Fork of Escalante Creek, about one and one-half
miles above Pickett Corral and 25 airline miles southeast of a release site
on Indian Creek (Rogers 1967).
In October, 1966, Mr. High, a State mine inspector, reported seeing 12 to
14 mountain quail at a spring about one mile below (southwest of) the site
where birds were released in 1966 according to Rogers (1967).
1967
On June 6, 1967, Wildlife Conservation Officer Tom Sherrill observed 4
mountain quail on South Fork of Mesa Creek, about 1 mile west of the Uncompahgre National Forest boundary and 11 miles southeast of the 1966 Indian
Creek release site. On October 28, 1967, deer hunters from California reported to Wildlife Conservation Officer Sherrill of seeing 42 mountain quail
in the same vicinity (Rogers 1968).
1968
Four to 6 mountain quail were reportedly seen by a Mr. Smith, one-fourth
mile below Blue Creek Ranch Headquarters in the spring of 1968.
This
location is approximately 6 airline miles southwest of the 1966 Indian Creek
release site (Hoffman 1969). All mountain quail sightings and reports during
the period 1966 through 1968 were summarized in Hoffman (1969).
1969
Twenty mountain quail were observed by Conservation Aide Tom Barnes on the
South Fork of Mesa Creek in December, 1969.
These were observed approximately
one-half mile above the old Bogus Mine (Hoffman 1971).
This location is
approximately 10 airline miles southeast of the 1966 Indian Creek release site.
1970
No additional

mountain

quail sightings

were recorded

for 1970

(Hoffman 1971).

�-249-

1971
In early December, 1971, late season deer hunters reported seeing 8 mountain
quail in an area approximately 2 miles northeast of Gateway, Colorado.
These
were observed intermittently until the early spring of 1972 by W.C.O. Dwight
Owens (Table 2). The Gateway Postoffice (elevation 4,570 feet) is approximately 9 airline miles northwest of the 1968 Indian Creek release site in
Colorado and approximately 15 to 20 airline miles northeast of Utah's Willow
Basin release site. Whether this small flock of 8 birds moved from Indian
Creek or Willow Basin is not known.
These birds apparently moved during
the early spring of 1972, probably to higher summer ranges.

1972
The 1972 release of 87 birds constituted the initial release of mountain
quail on the eastern slope of Colorado.
Thirteen mountain quail were observed
following the 1972 release and ali but 1 of these birds were seen at sites at
least 3 airline miles from the release site (Table 2). One bird was seen
approximately 1 mile from the release site in Vigil Canyon.
Field searches
and interviews with Division personnel and landowners indicate that most of
the 87 mountain quail released in 1972 dispersed in at least 3 different
directions (NE, SE, SW) shortly after the release was made (Table 2). These
dispersed in small groups of 1 to 5 or 6.

1973
Follow-up checks were not made in 1973 following the releasing of .all remaining pen-raised mountain quail (90) on March 20, 1973 on the Spanish
Peaks Management Area, because of a decision to abandon the study.

SUMMARY AND CONCLUSIONS
The status of mountain quail in Colorado is very uncertain at this time
because (1) numbers observed and reported since 1966 have been small relative to numbers released, (2) no instance of reproduction has been recorded to date, and (3) consistent observations for anyone
locality are
lacking.
In view of these findings, past releases of mountain quail must
be considered as questionable as to success in establishing populations
in Colorado.

�-250-

LITERATURE CITED

Hoffman, Donald M. 1969. Study of mountain quail adaptability. Job
Progress Report, Federal Aid Project W-37-R-22, WP 15, J2. Colo.
Div. of Game, Fish and Parks, Denver. pp. 155-158.
1970. Study of mountain quail adaptability. Job Progress
Report, Federal Aid Project W-37-R-23, WP 15, J2. Colo. Div. of
Game, Fish and Parks, Denver. pp. 167-170.
1971. Study of mountain quail adaptability. Job Progress
Report, Federal Aid Project W-37-R-24, WP 15, J2. Colo. Div. of
Game, Fish and Parks, Denver. pp. 79-83.
1972. Study of mountain quail adaptability. Job Progress
Report, Federal Aid Project W-37-R-25, WP 15, J2. Colo. Div. of
Game, Fish and Parks, Denver. pp. 73-76.
Rogers, Glenn E. 1965. Appraisal of mountain quail habitat.
Compl. Report, Federal Aid Project W-37-R-18, WP 15, Jl.
Dept. of Game, Fish and Parks, Denver. pp. 217-225.

Job
Colo.

1966. Study of mountain quail adaptability. Job Compl. Report,
Federal Aid Project W-37-R-19, WP 15, J2. Colo. Dept. of Game, Fish
and Parks, Denver. pp. 125-133.
1967. Study of mountain quail adaptability. Job Compl. Report,
Federal Aid Project W-37-R-20, WP 15, J2. Colo. Dept. of Game, Fish
and Parks, Denver. pp. 215-218.
1968. Study of mountain quail adaptability. Job Prog. Rept.,
Fed. Aid Proj. W-37-R-21, WP 15, Job 2. Colo. Div. of Game, Fish
and Parks, Denver. pp. 113-116.

Prepared by

;Y~m~

Donald M. Hoffman
Wildlife Researcher

�-251-

April, 1973

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~---------

Project No.

W-37-R-26

Work Plan No.

20

Job Title
Period Covered:
Personnel:

Game Bird Survey

Job No.
1
Investigation of Population
Status of Bobwhite Quail in Eastern Colorado

--~~~--~-------------------

April 1, 1972 through March 31, 1973

Tom Lederhos, John Corey, Bruce MCCloskey and Warren Snyder.

ABSTRACT

Bobwhite quail whistling counts obtained on the Tamarack Management Area from
mid-June to mid-July, 1972 were higher in number than in 1971. Corresponding
fall populations and harvest also were higher in 1972. In general, counts
throughout the South Platte Drainage were higher than those of the previous
year. Counts also were completed on routes established within the Arkansas,
Cimarron and Republican River drainages. A distribution and density map based,
in part, on whistling counts was prepared. Approximate square miles of various
range densities were tallied. The 1971 fall population estimate was revised downwar
downward to 1,745 quail based on additional band return information, exclusion
of the northeast corner fo the Tamarack from the sample, and minor corrections
for fall emigration. Approximately 27 percent of that population was removed
by hunting and 16.3 percent was lost to natural fall and winter mortality so
that 988 quail remained by March, 1972. Spring and summer mortality subsequently
removed 34.9 percent of the original fall population leaving 227 males and 152
females, or 379 adults in early fall, 1972. The fall to fall mortality totaled
78.3 percent. Significantly higher mortality of females than males was noted
over-summer. Production was high as a result of tenacious nesting efforts even
into August, resulting in a fall population estimate of 2,319 quail. Of these,
802 were fall banded and 34 previously banded adults were retrapped. Hunters
removed approximately 548 quail plus a crippling loss of about 15.7 percent
of that number. Total harvest mortality approximated 26.7 percent. Severe
cold and snow conditions in November and December, 1972 in unison with hunting
is believed to have substantially increased mortality over the previous year.
Only 47 bobwhite were banded and 31 previously banded retraps were taken in .
February, 1973 as evidence of a much reduced population. Vegetation, food
habits, and climatic conditions as limiting factors to bobwhite on the Tamarack
were discussed.

��-253-

INVESTIGATION OF POPULATION
STATUS OF BOBWHITE QUAIL IN EASTERN

COLORADO

Warren D. Snyder

P. S. OBJECTIVE
To investigate (1) the distribution and relative density of bobwhite
quail in eastern Colorado; and (2) population structure and level, and
rate of harvest, and identify key factors that limit bobwhite on the
Tamarack Management Area.

SEGMENT OBJECTIVES
1.

To investigate
Colorado.

the distribution

and density

2.

To investigate population structure and level~ and rate of harvest
bobwhite quail on the Tamarack Management Area.

3.

To identify
Area.

key factors limiting

METHODS

bobwhite

of bobwhite

in eastern

on the Tamarack

of

Management

AND MATERIALS

Methods and materials used in this investigation were previously listed by
Snyder (1972). Amendments to this listing are presented as follows:
1.

Distribution

and density

of bobwhite

quail in Eastern Colorado.

Inventory was expanded to include additional routes along the Arkansas
Valley.
Replicate counts were also completed on all routes previously established in eastern Colorado.
Wildlife Conservation Officers assisted in inventory counts along the South Platte River.
The Red Lion - East Tamarack route
was extended along the meadow, west of the ranch headquarters,
to more
thoroughly census the East Tamarack Management Unit.
Because of high calling rates obtained the previous year, use of the induced call technique was reduced or eliminated along many of the census routes
in 1972.
2.

Population

structure,

level and rate of harvest.

Quail trapping on the Tamarack study
and terminated on October 31, 1972. Low
trapping over a greater part of the area
bution.
Trap sites were prepared by use

area was initiated on September 25
water levels permitted expansion of
to attain increased sampling distriof shovel and rake instead of using

�-254-

a tractor and rototiller.
Signs were placed at key access areas along the Tamarack boundaries to
inform the hunting public that check station sign-up was required to hunt
in this controlled access hunting unit.
The Tamarack was used exclusively for population studies in 1972.
During the previous year, some additional properties, up-river, were also
included in study efforts.
3.

Key limiting

factors.

Ground cover samples measuring the occurrence of grass, brush and vines,
forbs, dead vegetation, litter and bare ground were obtained at six locations
along the Tamarack and at one pastured location (Lutes Property) southwest
of Merino.
Brush, vines and forbs were distinguished by species.
Over-story
canopies of cottonwoods were not included in the samples.
A metal ring, approximately three inches in diameter, held at arm's
length a foot in front of the right leg, projected a field of view for sampling
vegetation.
The predominate cover type or species observed within the observation field was recorded.
Samples were obtained at one or two pace intervals
while walking somewhat at random through the sample areas.
One person sampled
while a second person recorded.
This technique permitted obtaining a high
number of samples widely distributed over a large area in a relatively short
period of time.
Sampling was not completely random, but effort was directed
toward wide dispersal of sampling as opposed to sampling intensively within
a small area.
Vegetation height was obtained by use of a yardstick.
Vegetative density
within the metal ring field of view was measured by ocular estimate on a
scale of 1, 2, or 3 representing sparse, medium or dense cover.
The amount and duration of snow cover along with temperature comparisons
for the November-February
period were obtained from U. S. Bureau of Weather
records.
Data'obtained at a station five miles south of Sedgwick, Colorado
were utilized as most typical of conditions along the Tamarack.
Natural mortality estimates for winter and summer were obtained by use
of banded-unbanded
ratios to project populations alive during the fall, late
winter and subsequent fall periods.
Comparisons with snow cover and winter
andsurnmer weather conditions were made.
Hunting season removal, natural winter mortality and subsequent
duction figures were related to check for evidence of over-harvest.

pro-

Late hatched quail potentially were subject to higher mortality than
earlier broods under fall weather conditions in 1972. Ages of banded quail,
harvested in December or retrapped in Feburary, were compared to the overall
age structure of fall banded juveniles to determine if differential mortality
did occur.

�-255-

RESULTS AND DISCUSSION

Bobwhite Distribution and Density in Eastern Colorado
Distribution and density of bobwhite in eastern Colorado are illustrated
in Figures 1 and 2. Approximate square miles of various range densities
per county and region are summarized in Table 1. Although these data are
not considered highly accurate, they provide a base on which to revise and
improve. As these data indicate, habitat of fair to high quality for bobwhite is not abundant in Colorado.
South Platte River Drainage
Whistling count indices of male bobwhite indicated breeding popuiations
along the South Platte were generally higher in 1972 than during the previous
year. Comparisons of 1971 and 1972 natural (not induced) tally indices are
presented in Table 2. The overall average for all routes was 2.91 calling
males per stop in 1971 and 4.37 calling males per stop in 1972. The SterlingIliff route was excluded in this comparison. The figures for 1972 may be
skewed slightly above those for 1971 since induced call indices were not used
on all counts during the latter year. This reduced the time needed per station and route permitting coverage of more stations within the optimum listening
time interval. Induced calling was of little value when natural counts of 6,
8, or more males per stop made it difficult to distinguish calls of additional
males.
Indices representing high counts obtained per route are included for comparison (Table 3). Averaging nine routes along the South Platte River, they
project a 55 percent increase in calling rate for 1972 over 1971. The average
of all counts (Table 2) resulted in a 50 percent increase in calling rate for
1972 over 1971.
The Atwood-Merino area continued to provide the highest calling rate attained
along the river (Tables 2 and 3). Markedly higher tallies were also obtained
in the Brush-Fort Morgan area and upriver along the Con Schaefer Ranch near
Weldona. The State-line-Ovid route was believed, in part, to rank lower
because of high noise interference from proximal highway traffic. Other
variables such as location of listening stations, weather conditions, and
calling variance among mornings undoubtedly influenced the data. Calling
peaked in late June and continued high through the first two weeks in July.
Bobwhite quail hunter survey harvest information is not yet available for
comparison with call indices. However, an increased fall population and
increased harvest on the Tamarack over 1971 figures (presented in subsequent sections) correspond favorably with increased call indices obtained
there in 1972. Reference is made to Ellis, Thomas and Moore (1972), who
provide one of the best examples of the relationship between call indices
and pre-hunt population levels available in recent literature.

�W E L D

WAS

H I N G TON

I

N
V1

'"
I

FIG. L
BOBWHITE DISTRIBUTION AND DENSITY IN
NORTHEASTCOLORADOBASED ON 1971-72 SURVEYS.

Range
Classification

Whistling Birds/Sq.Mi.
Count Index
(Spring)

~

Isolatedto Scattered

None

0 to 4

~

ContinuousLow

0.5 - 0.9

5 to 9

® Fair to Moderate

1.0 - 2.4

10 to 24

~

2.5 - 3.9

25 to 39

4.0 +

40 +

Moderate to High

&lt;CD High to Very High

�LINCOLN

I

N
VI
'-I
I

FIG. 2.
BOBWHITEDISTRIBUTION ANDDENSITY IN
SOUTHEASTCOLORADO
BASEDON 1971-72 SURVEYS·
Whistling
Count Index

Range
Classification
~

Isolated

® Continuous
~

Fair

Low

0.5

to Moderate

1.0

@ Moderate
~

to Scattered

to High

High to Very High

2.5

BirdS/Sq.
Mile
(S~rin51)

None

0

to

4

- 0.9

5

to

9

10

to 24

25

to

- 2.4

- 3.9

4.0 +

40 +

39

�-258-

Table 1. The approximate square miles of bobwhite quail range by
density
classification per county and region in eastern Colorado, 1972.

County

Isolated
to Scattered

Densit~ Classification
Continuous
Fair to
Moderate
Low
Moderate
to High

High to
Very High

Total

Southeast Region
Baca

600

Prowers

360

Bent

137

Otero

118

Crowley

38

Pueblo

5

18

23

Kiowa

137

20

157

1395

147

Sub-total

105

183

52

940

18
4

378

13

8

25

187

6

7

9

140
38

220

67

34

1863

Northeast Region
Republican River Drainage
Kit Carson
Yuma
Phillips
Sub-total

65

6

1070

54

71
12

5

1141

85
1220

85
60

12

5

1297

South Platte Drainage
Sedgwick

15

Logan

35

3

Washington

3

2

Morgan

10

Weld

10

4

4

73

9

12

12

57

163

Sub-total

1293

69

24

17

57

1460

Grand Total

2688

216

244

84

91

3323

Sub-total

15

30

1

28

67

4

3

12

11

36

3

12

18

�-259-

Table 2. A comparison of 1971 and 1972 bobwhite quail whistling counts
completed along the South Platte River.

Route

Year

Number of
Counts

Number of
Stops

Natural
Tally

Mean

Stateline - Ovid

1971
1972

2
2

20
20

78
65

3.90
3.25

Red Lion E. Tamarack 1971
1972

2
4

20
45

67
202

3.35
4.49

West Tamarack

1971
1972

6
7

75
79

219
312

2.92
3.95

Sterling-Iliff

1972

2

19

63

3.33

Atwood - Merino

1971
1972

2
2

20
20

108
182

5.40
9.16

Balzac

1971
1972

3
2

28
21

55
72

1.96
3.43

Brush - Ft. Morgan

1971
1972

2
2

20
20

77
163

3.85
8.15

Con Schaefer

1971
1972

2
2

20
13

37
56

1.85
4.31

Goodrich - Orchard

1971
1972

2
2

20
23

23
32

1.15
1.39

Masters K-4

1971
1972

2
2

8
9

9
8

1.13
0.89

Overall Average!/

1971
1972

231
250

673
1,092

2.91
4.37

1/
- The Sterling-Iliff route was excluded in this comparison.

�-260-

Table 3. A comparison of high counts obtained on bobwhite quail census
routes in eastern Colorado during 1971 and 1972.
Route

1971

1972

South Platte Drainage
Stateline - Ovid

5.3

3.4

East Tamarack - Sonneburg

4.0

6.1

West Tamarack

4.1

8.2

Sterling-IlHf

4.3

Atwood - Merino

6.7

9.1

Balzac

2.5

5.8

Brush - Ft. Morgan

3.9

8.3

Con Schaefer

2.2

4.3

Goodrich-Orchard

1.4

2.3

Masters - K-4

1.2

1.0

Hardin - Kersey

0.0

0.0

Republican Drainage
South Fork - Bonny

1.4

0.5

Arikaree - Blk. Wolf Creek
North Fork (Laird - Wray)

1.4

2.8
1.9

Arkansas Drainage
E. Prowers County

0.8

1.8

Purgatoire - Ft. Lyon

5.2

9.4

La Junta - Las Animas

6.1

La Junta - Rocky Ford

3.5

Manzanola

0.9

Boone - E. Pueblo Co.

0.4
Cimarron Drainage

Campo East - Cimarron

3.8

5.5

�-261-

Arkansas River Drainage
Whistling counts were generally lower along the Arkansas River than along
the South Platte (Tables 2, 3 and 4). One exception was observed near
the town of Las Animas where the Purgatoire joins the Arkansas. Moderate
to high counts were also obtained along the Arkansas upstream to Rocky
Ford. Extension of low densities was noted to the east edge of Pueblo.
Bobwhite are confined to a narrow flood plain along the Arkansas east of
John Martin Reservoir by intensively farmed irrigated land and short-grass
pastures. The flood plain, itself, is grazed in most locations resulting
in fairly low populations of bobwhite (Table 4) (Fig. 2).

Table 4. A summary of bobwhite whistling counts completed in southeast
Colorado and the Republican River Drainage in 1971 and 1972.
Location and
Route

Year

Number of
Counts

Number of
Stops

Natural
.Tally

Mean

E. Prowers County

1971
1972

1
3

10
30

8
33

0.8
1.1

Purgatoire - Ft.
Lyon

1971
1972

1
2

10
24

52
194

5.2
8.1

Las Animas - La
Junta

1972

2

27

152

5.6

La Junta - Rocky
Ford

1972

2

23

76

3.3

Rocky Ford Manzanola

1972

2

18

8

0.4

Fowler Boone E. Pueblo

1972

3

34

9

0.3

1971
1972

1
2

10
28

38
145

3.8
5.2

Arkansas River Valley

Cimarron River VaHey
S. E. Baca County

ReEub1ican River Drainage
South Fork - Bonny

1971
1972

3
2

30
27

21
12

0.7
0.4

Arikaree

1972

1

11

31

2.8

North Fork

1971
1972

1
1

7
9

10
17

1.4
1.9

�-262-

Induced calling inventories were conducted along the Huerfano River,
Fountain Creek, Apishapa River and accessible sections of the Purgatoire
River northeast of Trinidad and south of La Junta.
Low bobwhite quail
densities apparently extend only a few miles up the Huerfano and Apishapa
Rivers (Fig. 2). Quail were not located along Fountain Creek north of
Pueblo.
Alex Hamilton, Area Supervisor, and John Stevenson, Wildlife
Conservation Officer,
reported that they were unaware of any populations
above Pueblo either on the Arkansas River or Fountain Creek.
Carl Leonard,
Area Supervisor, reported observing quail in the Higby area along the
Purgatoire River.
Farmlands exist along the Purgatoire in that area.
Low scattered bobwhite popUlations exist throughout the irrigated areas
along the Arkansas River from Pueblo east to Kansas.
Some of these coveys
may winter proximal to the river while others probably winter along irrigation ditches, in farm yard shelterbelts, and at other suitable locations.
Range boundaries to the north and south are abruptly cut short by semi-arid
short-grass rangelands in most areas.
Bobwhite show much closer reliance
on irrigated farmland interspersions with cover than do scaled quail also
resident along the Arkansas Valley.
Winter weather conditions are observably less severe in extreme southeast
Colorado than in the northeast corner of the State. As an apparent consequence, higher densities of bobwhite can survive in sand sage rangelands
of southeastern Colorado (Fig. 2). Bobwhite and scaled quail coveys often
intermingle in southern Baca County, but the latter species usually is
dominant in number.
Populations of both species fluctuate widely from year
to year.
Call indices along the southeast Baca County census route (Table 4) were
considered disproportionately
higher than those obtained along river
drainages.
Therefore, spring densities were adjusted downward in proportion to calling indices (Fig. 2). Census along rivers usually sampled
a semi-circle of quail habitat.
The remainder of the circle was farmland
or pasture.
Proximal highway traffic and heavy tree growth impeded tally
of distant calling males.
In the rangelands of Baca County, whistling bobwhite were usually distributed in all directions rather than in a semicircle.
Neither tree growth or traffic noise interfered with listening
for calls although winds presented an increased obstacle at times. As a
result, it was assumed that about twice as much bobwhite habitat was censused in rangelands compared to river-bottom census.

Republican

River Drainage

Lack of brush cover along the South Fork, North Fork, Frenchman and
Arikaree drainages limits quail densities in east-central Colorado.
The
Arikaree, with adjacent sand sage and scattered brushy draws, is the best
among this group of drainages (Table 4). Bobwhite exist along the Arikaree
up river to near the town of Cope in southeast Washington County (Fig. 1).

Other Rangeland
Along

Areas

the east edge of Colorado

most of the sand sage rangelands,

especially

�-263-

where
integrated with farmland, contained scattered to isolated coveys
as shown in Figs. 1 and 2. These coveY$ winter in plum or current
thickets at old homestead sites, and in windbreaks and other coverts
around occupied farm and ranch yards.
In spring they disperse into the
rangelands for reproduction.
Bobwhite occur across eastern Colorado at other isolated areas.
Reports
have been received from numerous straight, dryland farm areas along the
east edge of the state.
In addition, reports have been received from
Karval, Fort Collins, Estes Park, Vail and near Denver.
In many cases
these are stocked 'birds which mayor
may not persist in the area.
Such
populations are considered too scattered to merit placement on the distribution map.

Quail Harvest

in Relation

to Distribution

and Density

As indicated in Table 1 the southeast region contains slightly more bobwhite range classed as isolated to scattered than the northeast region,
which includes Kit Carson County.
But range classes from continuous low
to high comprise approximately 167 square miles in the northeast region
compared to 468 square miles in the southeast region.
These statistics
would mean that more bobwhite exist in the southeast part of the state
and harvest should be accordingly higher.
However, Small Game HunterHarvest Survey statistics, averaged for 1968 through 1971, fail to agree.
As Table 5 shows, apparently more quail were bagged by slightly more hunters
in the northeast region.
Reasons for these discrepancies are uncertain.
However, availability of
bobwhite is probably a primary reason.
Moderate to very high densities
along the Arkansas River reside primarily in salt cedar (Tamarisk) brush.
Hunting these coveys is difficult to impossible in this tall, dense brush.
Many of the bobwhite found elsewhere in southeast Colorado winter around
occupied farm residences where they are not available to the hunter.
Those
dwelling in the rangelands are more difficult to find than scaled quail.
Census of the two species in the early 1960's in southern Baca County rangelands revealed a ratio of bobwhite to scaled quail of 1 to 4.25.
Hunters
checked in the locality bagged approximately one bobwhite to eleven scaled
quail (Snyder 1967).
In northeast Colorado, harvest occurs mainly along the South Platte River
where high to very high densities exist. Rose bushes, buck brush and other
brush, grasses and weeds provide excellent huntable cover. Much of the
river is leased and nearly every mile is hunted to some degree.
The percentage harvest seems much higher.

�-264-

Table 5. Bobwhite quail harvest in eastern Colorado per region based
on the 1968-1971 average.
Management
Unit

Location

Hunters

Harvest

Average
Season Bag

1 and 2

South Platte, Arikaree
&amp; N. Fork of the
Republican

2,049

9,188

4.48

14

S. Fork of Republican
and Kit Carson Co.

197

788

4.00

2,246

9,976

Total:

Northeast Region

12, 13 &amp; 14

Arkansas Valley

1,511

5,112

3.38

26

Southern Baca Co.

415

2,047

4.93

1,916

7,159

4,172

17,135

Total:

Southeast Region

State Total

4.11

Relating Whistling Counts to Populations Resident on the Tamarack
An estimated 400 male bobwhite resided on the Tamarack in June, 1972 based
on change-in-ratio projections and mortality data presented later in the
text. Or, there were approximately 29 males per linear mile of river. This
compares to an average call index of 4.2 whistling males per stop or 8.4 males
per linear mile since stops were at one-half mile intervals. The average
maximum distance for hearing a whistling bobwhite is unknown but along the
river it is assumed to approximate one-fourth mile. If so, then less than
half of the calling quail per linear mile of river were heard since the river
averaged .45 mile wide and census was taken along the south edge. Past
experience with induced calling indicates that additional quail usually were
resident, but not calling except on peak census mornings.
Table 6 relates average calling indices on the Tamarack with late-winter
and subsequent fall population densities per 100 acres. Other reference
materials are included in the table for general comparison. Direct comparisons can not'be made due to differences in habitat and census conditions,
among other variables.

�-265-

Table 6. Comparisons of calling males with previous
subsequent fall population levels.

Mean Calling Males
Per Stop

Source

3.0 to 3.3
,
4.0 to 4.5

Tamarack

Illinois
(Edwards 1972)

Quail Density Per 100 Acres
Subsequent
Previous
Fall
Late Winter

47 1:../

(1971)
(1972)

26

1.0 to 1.3

3.8 to 6.0

1.5 to 2.1

7.5 to 8.0

0.8

Indiana
0.6
(Hoekstra &amp;
Kirkpatrick 1972)
Illinois
(Ell is &amp; Thomas
and Moore 1972)

late winter and

59

2

1.5

9.4

2.3

14.2

3.9

18.2

4.2

23.3

5.0

26.1

5.2

33.3

5.5

36.4

1:../ Habitat and census conditions are probably not comparable between Illinois
and Colorado.
Therefore,
in a general way.

Population

the Illinois data are applicable

Structure,

to Colorado

only

Level and Rate of Harvest

Some of the band return information obtained during this segment supplements
information obtained in the previous segment (Snyder 1972). This new information necessitates reassessment, and in some cases, correction of information previously presented.
These changes are presented in the following
pages before proceeding to statistics compiled during the 1972 harvest.

�-266Movements

and Emigration

of Banded Quail

Fall Movement - 1972--Habitat barriers on either side of the river greatly
restrict banded quail egress from the Tamarack.
But, during the fall of
1972, band returns from locations, both up and down river, gave evidence
that some emigration of banded quail had occurred.
Some of these move-.
ments covered long distances.
To illustrate, 5 bands with close consecutive numbers were obtained by hunters southeast of Sedgwick on November
11, 1972.
They came from quail banded on Area 7-West in late September
(Fig. 3). Linear movement of these quail downstream approximated 19 miles.
Two quail, banded in October on Area 9-West, moved upstream for approximately 30 miles to the John Bianco farm near Atwood where they were taken
on November 11, 1972. Clayton Wetherill, Wildlife Conservation Officer,
made a followup check on this report which verified its accuracy.
Three
additional quail bands, returned by hunters, were reportedly taken in the
Iliff-Proctor region.
Locations were not pinpointed so it is possible one
or more could have been taken on the west end of the Tamarack.
One of these
quail was banded on Area 9-West and two were banded on Area 25-West.
Review of trap, retrap and harvest information promotes some interesting
speculation concerning the previously described movements.
This information
is summarized as follows:
1. Evidence of high quail densities on Areas 7-West and 9-West is indicated
by the high numbers of quail banded on these areas, (64 and 56 respectively),
(Table 16).
2. Relative low harvest returns on these areas
many of the banded quail moved elsewhere.

(Table 16) lend evidence

that

3. These movements are confirmed by fall retrap information which showed some
of the quail on Area 9-West moved rapidly upstream where successive retraps
were made in West Areas 19, 21 and 22 within a few days after initial banding.
Harvest recoveries indicate at least one, and possibly more, remained on the
west end of the Tamarack while others of the covey continued on.
Banded quail, taken near Sedgwick, initially were found on Area 6-West.
They moved to Area 7-West, then to Area 9-West before reversing directions
and heading downstream.
One retrap of a member of this covey was recorded
in the Braddock pasture on the East Tamarack.
Retrap information indicates one other covey initially banded on Area
l2-West, moved upstream but remained on the study area through the hunting
season.
4. Additional fall and winter retrap and harvest return information (Table
7) shows nearly all other coveys remained resident where banded in fallon
the Tamarack.
5. Available evidence indicates greater movement occurred during the fall of
1972 than in 1971. A much higher density (2,319 compared to 1,745) existed
on the Tamarack the second fall. Possibly increased movement. was a result of
increased density.
Whether the quail leaving Areas 7 and 9-West were resident
and exceeding the carrying capacity, which induced movement, or were already
shifting locations when trapped, remains unknown.
Trap disturbance, itself,
may have induced movement, but it is evident that higher than average densities of quail were banded in the two or three hunting areas.

�TAMA RACK MANAGEMENT

AREA

~ tS1

~RE~S

I
N

-..J
'"
I

tv

/
EAST

AREAS

o

SC ALE
y.

Mi.

LEGEND
1

HUNTING

~UNFARMED
~
AND

AREA

RIVER

RIVER BOTTOM
CHANNELS

li;oiIGRASSLAND
DHAY

MEADOW

~.

AL.FALFA

~

ROW CROPS

LD

PLANTING

Fig. 3. Bobwhite habitat within the Tamarack Management Area along the South Platte River in
northeastern Colorado.

�-268-

Table 7. Approximate movements of bobwhite quail between various time
intervals on the Tamarack Management Area.

Period

0

1/4

1/2

Distance Moved 'irtMiles
3/4 1
1-1/2 2 2-1/2 3

3-1/2

4

4+

Number of Quail Moving Duting 'Fall'Periods
Fall-71to
Harvest-7l

51

28

23

4

9

1

2

0

0

0

1

0

Fall-71 to
Winter-72

33

20

10

1

1

1

1

0

0

0

0

0

Fall-72 to
Harvest-72

57

70

28

3

3

1

5

0

4

2

1

10

Fall-72 to
Winter-73

6

13

3

2

0

0

0

0

0

0

0

0

Movements Over Longer Time Intervals
Fall-71 to
Fall-72

6

7

2

2

1

0

0

2

2

2

0

0

Fall-71 to
Harvest-72

4

2

8

5

6

1

1

3

0

1

2

1

Winter-72 to
Fall-72

5

2

2

0

1

0

0

0

0

0

0

0

Winter-72 to
Winter-73

0

2

0

0

0

1

0

0

1

0

0

0

Fall-71 to
Winter-73

0

1

0

1

1

0

0

0

0

0

0

0

4

7

1

2

2

0

0

0

1

0

0

0

Winter-72 to
Harvest-72

Number of Quail Leaving the Tamarack-Fall, 1972--Available information indicates two or possibly three coveys left the Tamarack, while nearly all,
if not all, the other coveys remained resident. If thii,is correct, then
potentially 40 to 60 bobwhite left the study area, bas~~~n general covey
sizes in early fall. Since portions of these coveys were trapped and retrapped more than once, at least half probably were banded. If so, then
20 to 30 banded quail potentially left the study area.
Hunters reported bands from 7 to 10 quail taken outside the Tamarack during
the fall of 1972. Assuming randomness of selection, a 20 percent harvest
and 100 percent reporting, then approximately 50 banded quail left the
Tamarack.

�-269-

Since little confidence can be placed in either of the above listed calculations, a systematic approach to fall emigration of banded quail was
constructed and has been used for calculating egress in both 1971 and
1972. Although its accuracy is also questionable, it does provide a
relative base for comparing movement between and among years and for
making subsequent corrections in fall population estimates.
One important weakness, inherent in this method, is that quail usually
move as a covey in fall, not as individuals. In addition, since hunters
walk from area to area while hunting, many of the harvest recoveries were
not taken in the area which the hunter initially selected for hunting.
Therefore, harvested quail were frequently tallied in one area when they
were actually killed in another nearby hunting area. So, on the average
they were probably taken slightly closer to their initial trap site than
is depicted in Table 7. However, due to compensating errors, actual movements were probably only slightly less than those illustrated in the table.
The method applied and illustrated in Table 8 determines the percentage of
banded quail, based on movement data, which would leave the Tamarack from
the upstream or downstream ends of the Tamarack. Areas proximal to the
two ends would lose the highest percentage of banded quail. Those near the
center of the Tamarack would lose few, if any. Usually, only half of the
quail would be lost among those moving distances great enough to put them
out of the Tamarack. The other half would simply move inward and remain
on the Tamarack.

Table 8. Banded bobwhite quail leaving the Tamarack Management Area based
on movement-recovery data from fall to harvest and fall to winter, 1972-73.
Movement
in Miles

Percent of
Quail Moving

o to .25

70.2

None

None

0.50

14.9

25-W

0.75

2.4

1.0 to 1.75
1.50 to 1.75

Areas Influenced
West
East

Total Quail
Banded

Number Quail
Leaving !/

ll-E

56

8.3

24-w

10-E

39

2.9

1.4

23-W

9-E

56

3.5

0.5

22-W

8-E

51

2.8

2.00 to 2.25

2.4

21, 20

7, 6

96

5.1

2.50 to 2.75

0

19, 18

5, 4

60

2.4

3.00 to 3.25

1.9

17, 16

3, 2

72

3.0

3.50 to 3.75

1.0

IS, 14

1

27

0.8

4.00 to 4.25

0.5

13, 12

1/2 Brad.Past. 37

1.0

Greater Distances

4.8

ll-W to Brad Past.

Total Quail

308

14.8

802

44.6

~/Assumes one-half of the quail were leaving via each end of the Tamarack
except for the 308 quail in the center of which 4.8 percent would leave by
either end. Movement distances by other quail would not put them out of the
l':Tnrlv

~"P::l_

�-270-

Impact of Movement onChange-in~Ratio
Ca1cu1ations--Based
on calculations
in Table 8, 45 banded quail emigrated from the study area following fall
banding in 1972, compared to 12 lost the previous fall. We can only assume
that a directly proportional number of unbanded quail also left the Tamarack.
If there was no immigration of unbanded quail, the ratio of banded to unbanded would not change and adjustments prior to calculation of fall population estimates would not be needed.
However, it must be assumed, without
evidence to the contrary, that directly proportional immigration of unbanded
birds did occur.
Therefore, necessary corrections have been made in estimates
of resident fall populations on the Tamarack (Tables 9 and 14).

Movement - Spring and Summer,1972--The
number of quail banded in late winter,
1972 which were retrapped the subsequent fall (Table 7) is too small to use
as a base for movement calculations.
All other data in the table include at
least one, if not two fall movement intervals.
Fall movements might tend to
compensate summer movements or be additives.
Therefore, fall movement data
can not be subtracted from total movement data to derive summer movement.
Rosene (1969) reported that quail usually moved only short distances in
early spring after covey breakup.
Results of most studies that he cited
confirmed this. Available evidence from the Tamarack indicates summer movement upstream or downstream was not of significance.
First, hunters did not
report bands of adult quail in adjacent river areas.
Second, the percentage
of 1971 fall banded quail in the population did not decline over-summer.
Samples indicated a slight increase (Table 9). If banded quail moved out of
the study area over-summer and unbanded quail moved in we would expect a percentage decline of banded quail.

Population

Levels and Mortality

Rates From 1971 Through

1972

Proportion Banded in the 1971 Fall Population--Band
recoveries obtained during
harvest, 1971 indicate that approximately 25 percent of the fall population
had been banded prior to hunting season (Snyder 1972). Winter sampling indicated that 30.5 percent were banded.
Subsequent samples obtained in fall,
1972 and winter, 1973 indicated that 33.3 percent were banded of that original
population.
Since there was no basis for assuming a higher mortality rate
among unbanded quail, a reassessment of the discrepancies was in order.
Review of information obtained during harvest, 1971 indicated that of 411
quail originally included in the harvest sample, 119, or 29 percent were
banded.
It was only the July report of one party, hunting in the vicinity of
the Armstrong Ranch, that altered the projected sample.
That party reported
four banded quail among approximately 75 quail taken.
Inclusion of this
information into the total harvest greatly increased the projected total fall
popUlation estimate of quail resident on the east part of the Tamarack.
The exact location of harvest by this hunting party is unknown.
Some hunting
apparently was on, or proximal to, the Armstrong property along the north
edge of the East Tamarack.
Additional quail probably were taken to the east
on Division property along the north side of the River (Fig. 3). In general,

�-271-

this latter location is not sampled by hunters repprting through the check
station and hunting from the south side of the'Riv'er., Because of access
difficulties trapping was not attempted in the location except in the winter
of 1972. Then only five quail were banded there.
Only minor harvest return
information came from the region during 1972.
Based on the above information, it seems justified to exclude this portion
of the Tamarack from the study.
The one late report of 75 quail will also
be excluded from the 1971 harvest sample for purposes of analysis in the
following sections of this report.

Revised Fall Population Estimates, 1971--Table 9 summarizes revised 1971
fall population estimates including corrections for quail lost to emigration.
Two population estimates are presented in the table.
The first projects an
estimate of 1,849 quail based only on corrected 1971 harvest returns.
Inclusion of samples from winter, 1972, fall, 1972 and winter, 1973 samples
reduced the estimate further to 1,745 quail as shown at the bottom of Table
9. This latter estimate, based on a sample of 817 quail handled, is considered most accurate and will be used in subsequent derivations of natural
mortality and population turnover.
Based on data from Robson and Regier
(1964) this sample size was sufficient to prevent errors greater than 10
percent with 95 percent accuracy.

Natural Mortality - Fall to Late Winter, 1971-72--The ratio of fall, 1971
banded quail to winter, 1972 banded quail that were recovered in subsequent
samples permits an estimate of natural mortality from fall to late winter.
Change-in-ratio
computations used in deriving the late winter population
are presented in Table 10. The recovery of 61 banded quail from fall, 1971
and 32 banded quail from winter, 1972 does not provide a large sample on
which to place high confidence of accuracy.
However, because of high annual
mortality rates, little additional information will be available in subsequent samples.
If the late winter population estimate of 988 quail is near accurate, natural mortality approximated only 16.3 percent based on a fall population
estimate of 1,745 quail.
For comparison another method of calculation
was used. Natural and harvest mortality were added to surviving quail
(based on 30.3 percent banded).
A 1971 fall population estimate of 1,776
quail was derived, which compares favorably with the previous estimate of
1,745.
Therefore, we can conclude that banded (fall - winter) recovery
ratios obtained in fall, 1972 and winter, 1973 were closely proximal to
the actual.
Combined natural mortality and harvest mortality approximated 43 percent
for the fall through winter period.
Rosene (1969) recommended a 45 percent harvest plus winter loss mortality as maximum for northern regions of
the species range.

�-272-

Table 9. Revised fall population estimates for 1971 based on (1) adjusted fall~ 1971 harvest samples and (2) addition of subsequent samples
obtained through the winter ·of1972.,.,.73.
West Tamarack

East Tamarack

Total Area

Number Banded

317

214

531

Marked Lost to Emigration

10

2

12

Banded Rema ining

307

212

519

Adjusted Harvest

255

156

411

Crippling Loss

38

23

61

Item

14.8

Percent Crippling Loss
Projected Total Harvest Mortality 293

179

472

Band Recoveries

62

57

119

Projected Band Recoveries

71

64

135

Percent Harvest

23.1

30.2

26.0

586
Fall Population Estimate
1~263
+
(Based on Harvest Returns
Alone)
Subsequent Sample Additions
Sample/Banded (Winter, 1972)

=

77/35

146/33

223/68
30.5

Percent Banded
Sample/Banded (Fall &amp; Winter,
1972-73)

183/61

50/21

133/40

33.3

Percent Banded

817/248

283/113

Sample/Banded (Combined Totals) 534/135
Percent Banded

25.3

39.9

30.3

Percent Harvest

24.1

33.7

27.0

Fall Population Estimate
(Based on Combined Samples)

1,214

+

531

=

1,745

�-273-

Table 10.
1971-72.

Derivation of natural mortality during the fall and winter of

West Tamarack

East Tamarack

Total

Fall Banded Quail Remaining
(After Emigration)

307

212

519

Projected Harvest Removal

71

64

135

Banded Quail Subject to Nat.
Mortality

236

148

384

Retraps, Fall-72

15

9

24

Recoveries, Harvest-72

23

11

34

Retraps, Winter-73

2

1

3

Total

40

21

61

113

42

155

7

3

10

Recoveries, Harvest-72

13

5

18

Retraps, Winter-73

4

o

4

Total

24

8

32

Fall Banded Alive at End of
Winter-72

188

llO

298

Fall Banded Natural Mortality

48

38

86

Total Natural Mortality

190

95

285

15.6

17.9

16.3

293

179

472

Percent Harvest Hortality

23.1

30.2

27.0

Natural Plus Harvest Mortality

483

274

757

Percent Total Mortality
Quail Surviving on March 5, 1972

39.8
731

51.6
257

43.3
988

Item

Fall, 1971 Retraps or Band
Recoveries

Quail Banded Late Winter, 1972
Winter, 1972 Retraps or Band
Recoveries
Retraps, Fall-72

Percent Natural Mortality
Total Harvest Mortality

�-274-

Weekly Rate of Natural Mortality - Fall to Late Winter, 1971-72--A hypothetical model was presented by Snyder (1972) to calculate the number of
quail alive at the start of the 1971 hunting season. A,mortality rate
of two percent per week was used as a base for these calculations. If
information just presented on fall to winter natural mortality is near
correct then the actual weekly rate of mortality was slightly less than
one percent per week. Based on a standard one percent per week loss
approximately 133 quail died of natural causes prior to the start of
hunting season in 1971 (Fig. 4).

Natural Mortality - Adults Through Spring 'and 'Sunlmer,"1972--Previous projections indicated there were approximately 988 quail on the Tamarack in
March, 1972. Of these, 520 were males and 468 were females (Table 11).
By October, 1972 their total number had dropped to approximately 379 or
a decline of 61.6 percent. The ratio of adult males to adult females,
combined from fall trapping and harvest, indicates 60 percent, or 227,
were males. Forty percent, or 152 hens, were still alive.
Spring-summer mortality of males equalled 56.3 percent whereas hens declined in number by 67.5 percent from the late winter population level.
Based on the original fall population estimate of 1,745 quail, spring and
summer mortality equalled 34.9 percent compared to 27.0 harvest mortality
and 16.3 percent fall and winter natural mortality. In total, 1,366, or
78.3 percent, of the original population were lost from early fall, 1971
to early fall, 1972.

Fall Banding--The fall trapping effort along the Tamarack resulted in
banding of 802 bobwhite and retrap of 34 previously banded adults. Trapping commenced on September 25 and was terminated on October 31 when heavy
snowfall shut down operations. Table 19 summarizes the composition of the
trapped sample by age and sex. Net top traps, used for the most part,
resulted in minimal scalping. Two quail, severely scalped in one trap were
released on September 25. Both when retrapped a few days later were
rapidly healing. One was bagged later in the same vicinity of November 16,
apparently without evidence of the previous scalping. A few quail were lost
to predation during the fall trap efforts, but the number was not over one
or two percent of the total number banded.
Low water levels in the river, dry ground conditions and additional time
permitted trapping to be extended into more areas along the Tamarack than
during the preceding fall. Therefore, a much better distribution of sampling
was attained.

�-2752,600

\
I
I

2,500

I

2,400

,

\
\

I

\
\

,,

,

I

2,300

2,319
Quail alive
Oct. 8, 1972

2,200
2,100
r-t

2,000
~
II
~ 1,900
~
.v 1,800
ftI

1,745 Quail alive

r-t

5. 1,700

Oct. 1971

0

0.

~

1,600

ftI

6 1,500

Hunting Season
Removal

ftI

~ 1,400

i 1,300
! 1,200
ftI

Hunting Season
Removal

~ 1,100

M 1,000
~

a

900
800
700
600
500
400

335 Adults

Adults

300
200
100

o
Oct.

Nov.
1971

Dec.

Jan.

Feb. Mar.

Apr.

May
1972

Fig. 4. Projected population levels of bobwhite quail
1972 on the Tamarack ManagementArea.

June

July

Aug.

from October,

Sept.

Oct. Nov.

1971 to October,

�-276-

Table 11. Natural mortality of bobwhite quail through the spring and
sunnner of 1972 on the Tamarack Management Area •.
Item

Males

Females

Total

Projected Quail Alive March, 1972

520

468

988

Projected Alive, October, 1972

227

152

379

Natural Mortality, Spring and Summer

293

316

609

Percent Composition of Survivors

60.0

40.0

100.0

Percent Mortality (Spring Base)

56.3

67.5

61.6

Percent Mortality (Previous Fall Base)

34.4

35.4

34.9

Percent Annual Mortality (Fall to Fall)

75.7

83.0

78.3

Projected Period and Peak of Hatch. 1972--As illustrated in Fig. 5, the hatch
of quail during the summer of 1972 was spread across the entire summer. The
initial and main peak of hatch occurred in late June and early July. A
second peak of lower impact occurred in mid-August. A few late quail were
hatched in early September. Information, illustrated in Fig. 5 is based on
aging of 768 quail during the fall trapping period. Ages were not precisely
determined, but it is believed they tended to average out within various age
intervals.

Harvest, 1972--The 1972 quail season in Small Game ,Management Unit 1 began
at noon on Saturday, November 11 and continued through November 19. An
extended season opened on the 9th of December and continued through the 31st.
The,resultant 31.5 day season compared with 20.5 days for the previous year.
Harvest by age, sex and day of season are presented in Table 12. Harvest of
other small game species is also included in the Table for comparison. Rabbit,
squirrel and waterfowl seasons extended beyond the dates given for quail.
Therefore, the totals in Table 12 do not reflect the total season bag for
those species. Table 13 compares the harvest of species or groups of
species between 1971 and 1972. Waterfowl harvest increased most significantly,
due primarily to weather conditions. Pheasants, quail, and cottontail harvests also were proportionally above those of the previous year.

�120

110

100

~)'t?~~

Per iod and Peak of Hatch

90

l~~~~~~~l

Winter Survivors

80

by Age

t=l

&lt;

s 70
ell

~
~

ro

sr...
0

I
N

60

~
~
I

50

a:
w

~
~

40

30

20
10

0
14-21
Mf-\.Y

. 22-29 30-5

I

6-13

14-21 22.-29 30-7

JUNE

,

8-15

16-23 24-31 1-8

JULY

I

9-16

17-24 25-2 3-10

t.UGUST

I

1,'iEEKLY INTcf&lt;V!.L

Fig. 5. The period and peak of bobwhite hatch on the Tamarack based on 768 aged quei.l.in 1972.
The suppl ernent al. small graph depicts ages of qua Ll, surviving in December and Februory.

SEPT.

�-278Table 12. Harvest of small game per day on the Tamarack Management Area
during the 1972 quail and pheasant season. 1/

Date

Pheasants

Ducks llRabbits

Squirrels

Ad.
Male

Bobwhite Quail
Ad. Imm. Imm.
Fem-~Male Fem •.Unc1. Total

First Season
11-11
12
13
14
15
16
17
18
19

16
29
17
4
6
3
4
11
0

27
14
42
25
29
29
25
17
15

8
13
1
0
0
1
8
26
5

0
0
0
2
0
0
0
0
0

8
6
2
0
1
1
1
4
5

2
3
2
0
1
1
1
1
2

28
13
6
5
7
15
7
12
11

35
18
9
6
9
7
8
6
8

1
2
0
0
0
0
0
2
3

74
42
19
11
18
24
17
25
29

Sub-total 90

223

62

2

28

13

104

106

8

259

5
9
0
1
3
10
9
15
10
1
1
5
0
3
4
0
0
0
1
5
1
6
2

1
1
0
7
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0

24
27
4
17
23
31
12
46
20
3
4
10
1
9
8
0
0
1
1
14
4
13
3

Extended Season
12-09
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31

6
2
0
1
1
4
0
13
6
0
5
2
1
1
1
1
1
2
0
5
1
1
0

68
54
9
18
17
59
39
33
11
0
11
13
16
6
9
9
0
21
9
18
18
38
5

12
5
1
3
0
6
4
19
4
1
7
2
0
5
0
3
0
2
4
5
1
4
6

0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0

3
2
1
4
3
5
1
7
1
0
0
0
1
2
2
0
0
0
0
1
1
0
1

1
1
2
2
5
3
0
5
4
0
0
2
0
0
0
0
0
1
0
0
0
1
0

14
14
1
3
12
13
2
19
5
2
3
3
0
3
2
0
0
0
0
8
2
6
0

Sub-total 53

481

94

2

35

26

113

91

10

275

Total

704

156

4

63

39

217

197

18

534

143

----

II

- Seasons for waterfowl, rabbits and squirrels extend
beyond dates listed here.
llInc1uded 518 mallard drakes, 139 mallard hens and 47 others of mixed species.

�-279-

Table 13. A compar ison of small game harvest
quail and pheasant seasons.

Year

Length of
Season

Quail

1971

20.5 days

411

1972

1s t 20. 5 days

1972

31. 5 days

during the 1971 and 1972

Ducks

Rabbits

Squirrels

87

196

30

9

480

130

555

126

4

548

143

704

156

4

Pheasants

Harvest Sample--Hunters removed 533 bobwhite from the Tamarack in 1972 as
reported through the check station.
Information from mailed-in returns
indicated at least 15 additional
quail were removed, but not reported through
the check station bringing the total check to 548. In 1971, 371 quail were
checked in 21 days of hunting.
The first
21 days of the 1972 seasons produced 480 quail in the bag. Total hunter use per day on the Tamarack was
approximately equal dur ing both years.
Posting of access areas to the lamarack with controlled access signs was only
partially
successful in curtailing
unchecked lrunting along the north side of
the river and in the Proctor Bridge vicinity.
A regulation is needed making
it a misdemeanor to trespass on controlled access areas such as the Tamarack
where hunting access is regulated through a check station.

WoundingLoss-Hunters questioned concerning crippling loss reported 78
wounded and not recovered.
Inclusion of a few parties missed in the inquiry,
proj ects the total to 86 bobwhite lost.
This represents 15.7 percent of the
knownmortality compared to 14.8 percent reported lost in 1971. Knownharvest
plus crippl ing loss brings the proj ected total harvest mortality to 634 quail
during the 1972 season (Table 14).

Use of Hunting Dogs--Most quail hunters use dogs as indicated by a sample of
those checked with quail at the check station.
Hunting parties with dogs
totaled 120 or 71.4 percent, contrasted to 48 parties without dogs (Table 15).
Amongsuccessful hunting parties,
the quail bagged per hunter averaged higher,
1.42 quail, for those using dogs, than for those without dogs, where the
average was 0.82 quail per hunter.
Despite more shots and a higher bag
for hunters using dogs, the crippling loss was slightly less, 0.197 cripples
per hunter, contrasted to 0.217 cripples per lrunter without dogs (Table 15).
Most significant
is the ratio of crippling loss per quail bagged. The ratio
is nearly 2 to 1 in preference toward hunters with dogs. Unsuccessful hunting
parties are not included in these samples.

�-280-

Table 14. A summary of fall, 1972 bobwhite quail banding and harvest
information from the Tamarack Management Area.
West
Tamarack

East
Tamarack

Total
Area

Number of Fall Banded Quail

490

312

802

Number of Retrapped Adults

22

12

34

512

324

836

29

18

47

483

306

789

113

'62

175

8

2

10

Total Band Recoveries

121

64

185

Crippling Loss (Projected)

63

23

86

Percent Crippling Loss

16.5

13.8

15.7

Known Harvest

381

167

548

Projected Total Harvest

444

190

634

Projected Total Band Recoveries

138

73

211

Percent Banded

31.75

38.32

33.75

Percent Harvest (Projected Total)

28.57

23.86

26.74

798

2,319

Item

Total Marked
Marked Lost to Emigration
Marked Remaining
Band Recoveries - Fall Banded
Retrapped Adults

Fall Population Estimate

1,521

+

As illustrated in Table 15 the proportion of those hunting without dogs was
much higher during the first nine-day season than during the second season.
Two reasons for this are possible. First, more of the hunters participating
in the second season may have possessed dogs, or hunters with dogs were more
successful in finding and bagging quail during the second season than those
without dogs. Both variables probably are true to a degree.

�-281-

Table 15. Quail hunter success and crippling loss among hunters with
and without hunting dogs during the 1972 Tamarack quail season 1/.
Hunters Using Dogs
Number of
Number of
Quail
Hunters

Hunters Without Dogs
Number of
Number of
Number of
Cripples
Hunters
Quail

Number of
Cripples

First Season
112

174

81

66

21

39

32

5

120

98

26

29

Second Season
162

214

25

Combined
274

Total

54

388

Quail Per Hunter

0.82

0.197

Cripples

Per Hunter

0.217

0.139

Cripples
Bagged

Per Quail

0.265

142

1/
- Sample includes

information

only from successful

hunting

parties.

Harvest and Hunting Effort by Hunting Area--Utilization
of hunting areas or
zones along the Tamarack property is indicated in a general way (Table 16).
Since hunters could walk into other areas after 9 AM, much of the quail harvest assigned to these areas may actually have occurred in adjoining areas.
But the tabular data do provide a general index of hunter use and success in
the various areas or groups of areas.
It must be noted that much of the
selectivity for specific areas was for duck hu~ting rather than for quail
hunting.
A breakdown of hunter effort by species was ~ttempted, but soon abandoned
as futile.
After the early morning duck hunting effort, nearly all hunters
were opportunists and attempted to bag any available species of game.

�-282-

Table 16. A summary by hunting area of fall banded bobwhite, hunter use,
total harvest and banded harvest recoveries on the Tamarack Management
area in 1972.

Area

Number Quail .Number of
Banded
Hunters !/

Hunter
Hours !/

Total Quail
Harvest 'l:../

Banded Quail
Harvest 3/

28
54
26
14
18
28

16
22

West Tamarack
1
2
4

16
6
21
24

5
6

5
32

7
8

64
11
56
10

43

30

7
0
5

3

9

10
11
12
13

14
15
16
17
18
19
20
21
22

11
18

57
49
39

181
133

126
82
198
170

31

61
39
57
52

163

5

14
10

60

166
120
199
194

34

126

57
48
67

173
157
261

35

!!../

12
21
7
9
3

7

1
6

10
3
8
5

5

10
1
2

2

4

2

7
21
4
21

2

0

16
22

53

312
263

8

45

209

29
22
28
20

55

318

44

19

7

62

8
21

4

43

46

188
319
343
192

10

27

13

Sub...,.tota1
504

1,175

4,593

381

149

23

24
25

71

70

3
4
6

East Tamarack
Braddock P. 63
1
11
2

44

3
4

10
16

5

6

6
7
8

4/
63
45
47
48
41
68
59

196

4

3

139

20

11

13
8

6

14
27
10
26

4

9

23
36
29
28

10

19

63
44
30

11

13

73

168
117
126
225
198
214
170
109
223

298

581

Grand Total 802

1,756

Sub-total

1/

2/

3/
4/

8
18

7

15
9

14

19

4
1
4

1,885

167

78

6,478

548

227

Includes only known hunters and hours reported through the check station.
Includes only known harvest. Some hunting probably went unrecorded.
Approximates the location where quail were shot. Exact location unknown.
Area closed to public hunting.

�-283-

Comparison With Previous Years of Harvest--Approximately 4.3 hours were
expended per animal bagged in 1972 by the average hunter compared to 6.2
hours per animal in 1971. Duck hunting during the period yielded a greater
return than quail hunting did in contrast to the previous year. Direct
comparisons in Table 13 show that 1972 was a significantly better year
for harvest of most species.
Quail harvest on the Tamarack in 1972 was second highest on record since
1952 according to information presented by Snyder (1972). Only in 1958 was
a higher harvest of quail recorded.
Hunting Success with Season Progression--Some decline in hunting success
was noted with season progression both in 1971 and 1972 (Table 17). The
most significant decline occurred during the last eight days of the extended 1972 season when very few quail were taken. This probably reflects
a combination of factors including a declining quail population due to
.previous harvest and increased wariness of remaining quail~ It is also
possible that significant natural mortality occurred prior to the last week
in December, 1972 as a result of severe weather in combination with prolonged snow cover and hunting stress.

Table 17. Hunter success ratios on bobwhite quail with the progression of
hunting seasons in 1971 and 1972.
Interval

Number of Hunters 1/

Number of Quail

Hunters/Quail

1971 Season
1st 8 Days

638

258

2.47

2nd 8 days

363

100

3.63

Last 5 Days

258

53

4.87

1972 Season
1st 8 Days

606

230

2.63

2nd 8 Days

386

167

2.31

3rd 8 Days

369

100

3.69

4th 8 Days

392

36

10.89

1:./
Includes all hunters using the Tamarack and not just quail hunters.

�-284-

Winter Bartding--Trapping was conducted through the month of February,
1973, with generally poor success. Only 47 quail were banded and 31
retraps were taken. Retraps included 24 fall, 1972 banded quail,
4 winter, 1972 banded, and 3 quail banded in fall, 1971.
The trapping effort and success indicated a marked drop in the number
of wintering quail compared to the previous year. Approximately half
as many man-days of effort were utilized in 1973 as in 1972 winter
trapping, but less than a third as many quail were trapped. Considerably
fewer coveys were located and most were much smaller in number, averaging
only 5 to 10 birds.
Muddy trails and high water conditions in 1973 hindered trapping operations.
Notably higher cottontail populations resulted in more frequent catches of
this species. As a consequence, use of metal-topped traps was mandatory.

Age Ratios - Fall to Winter, 1972--A highly significant difference in adult
to young ratios existed between banded and unbanded samples in the total
harvest sample (Table 18) •(Chi-square = 24.33** 1 d.f.). As previously
described (Snyder 1972) this indicates trap bias toward young during fall
trapping. It is assumed that harvest provides an unbiased sample of age
and sex in bobwhite quail.
Corrections to determine the actual proportion of adults in the early fall
sample are complicated by discrepancies between banded and unbanded samples
between the early (November) season and the second (December) season (Table
18). The banded samples show no evidence of disproportionately greater
juvenile mortality between the two seasons. The unbanded samples, containing much higher samples, show considerable evidence of first season to
second season differential loss of young. A chi-square analysis of the
total (banded + unbanded) ratios between the two seasons show significance
at the 0.05 level for a 1 tailed test (Chi-square = 3.63 with 1 d.f.).
This would indicate differentially higher juvenile mortality did occur between the two hunting seasons. It is supported by ratios obtained in late
winter samples (Table 18).

•

Apparently, the percentage adults in the early fall population lay somewhere
between 15.36 and 19.76. Assuming little or no differential mortality between fall banding and the first season and significant differential mortality between the two harvest periods, 16.33 will be used as the percentage of adults originally present in the fall population.
Robel (1965) in Kansas found that differentially greater mortality of young
came when snows, sub-freezing temperatures and lower food supplies were first
encountered in December. On the Tamarack, these conditions prevailed in
close unison with the November and December hunting seasons in 1972. Assuming
adults comprised 16.33 percent of the early fall population of 2,319 quail,
then there were 379 adults within that population at the mid-point of fall
trapping.

�-285-

Table 18. Quail age ratio information obtained on the Tamarack Management
Area during the fall and winter of 1972-73.

Total

Percent
Adults

Adult: Young
Ratio

776

836

7.18

1 : 12.93

8

84

92

8.70

33

126

159

20.75

41

210

251

16.33

Banded

7

85

92

7.61

Unbanded

54

119

173

31.21

61

204

265

23.02

Banded

15

169

184

8.15

Unbanded

87

245

332

26.20

102

414

516

19.76

Banded

0

24

24

0

Unbanded

19

35

47

40.43

19

59

78

24.35

Number
Adults

Number
Young

60

Banded
Unbanded

Item
Fall Banded Sample
Hunting Season Sample
November Season

Total

1

5.12

1

3.34

1

4.06

1

3.10

December Season

Total
Combined Seasons

Total
Winter Trapped

Total

Age Ratio Comparison With the Previous Year--The banded retrapped and banded
adults in the 1972 fall population represent only 7.18 percent of the total
sample compared to 11.5 percent banded adults in the 1971 fall banded sample.
Best estimates indicate that 19.2 percent and 16.3 percent of the fall populations were adults respectively in 1971 and 1972. If these ratios are
correct, then 1972 was a better production year. The higher fall population
present in 1972 provides support to that s t atiement;
,

�-286-

The 1972 Fall Population Estimate--Projections based on fall banded minus
movement samples indicate there were approximately 1,521 quail on the west
section of the Tamarack and 798 quail on the east section (Table 14) in
early October, 1972. The combined total of 2,319 quail is proximal to an
estimate of 2,337 obtained by direct calculation in the last column of
Table 14. If immigration of unbanded quail was not considered, then the
total population estimate for 1972 would equal 2,457 birds. These estimates may again be revised following collection of additional band returns
during the fall and winter of 1973 and 1974.
During the early fall of 1972 there were approximately 384 quail per square
mile or 60 quail per 100 acres resident on the Tamarack river bottom. This
compares to 289 quail per square mile or 45 quail per 100 acres found on the
Tamarack in the fall of 1971.

Sex Ratios--The higher rate of mortality among hens than among males over
summer was previously illustrated and discussed (Table 11). This phenomenon
routinely occurs in bobwhite populations as shown by numerous references in
the literature. A summary of sex ratios for adult and immature quail obtained
in fall and winter, 1972-73 is provided in Table 19.

Table 19. Sex ratios of adult and immature quail obtained during fall and
winter trapping and harvest, 1972-73.
Sample

Adult Male

Adult Female

Immature Male

Immature Female

Fall Trapped

36

22

375

313

Hunting Season

63

39

217

197

Winter Trapped

11

8

27

32

Total

110

69

619

542

61.45

38.55

53.32

46.68

Percent of Totals
By Age Class

Confidence limits calculated from a formula provided by Davis (1960) are
presented in Table 20. Finite corrections for this finite population are
also included (Snedecor 1956).

�-287-

Table 20. Fall population estimates for 1972 with confidence limits at
the 95 percent level for the Tamarack Study Area.
Fall Population
Estimate

Population

Confidence
Limits

Conf. Interval With
Finite Correction

West Tamarack

1,521

±

228.4

±

197.8

East Tamarack

798

±

156.8

±

139.5

2,319

±

279.7

±

244.4

Total Area

Robson and Regier (1964) recommend that samples should be large enough to
prevent errors in population estimates greater than 10 percent with 95
percent accuracy for the type study presented here. Comparison of the
1972 fall banded sample and harvest recovery information with charts included in Robson and Regier (1964) show that sample sizes were adequate
to achieve this degree of accuracy. Examination of additional members of
the population for bands in the future will further improve accuracy.
Factors Limiting Bobwhite Quail on the Tamarack
Habitat Conditions as Limiting Factors
The high densities attained both in 1971 and 1972 provide evidence that
conditions along the ungrazed South Platte flood plain are quite well
suited to bobwhite. Apparently brush, food, nesting cover and the other
requirements needed by bobwhite are generally in good supply. Periodic
floods along the river may temporarily disrupt nesting and curtail reproduction, but in the long run, they insure numerous forbs and early successional vegetation needed by bobwhite.

Vegetation Sampling
Late summer inventories show a wide variety of annual and perennial forbs
abound along the South Platte (Table 21). By general occurrence these forbs
occupy about 30 to 40 percent of the sampled sites within the Tamarack
(Table 22). Grasses usually rank second in abundance with varied lesser
amounts of brush and trees, dead vegetation, litter and bare ground. Overstory cottonwoods were not included in the samples. General height and density measurements are provided in Table 23.

�Table 21.

Forb occurrence on six sites along the Tamarack Management Area in late summer, 1972.
6-7
West

14-15
West

3-6
East

Braddock
111 '!:..I

Braddock

Genera or species
Per. Ragweed
Lamb's Quart.
Clover
Kochia
Horseweed spp.
Sunflower
Annual Ragweed
Wild Lettuce
Four-O'Clock
Milk Purslane
Purslane
Curly Cock
Ground Cherry
Poison Hemlock
Dandelion
Gumweed
Crotons
Snow-on-the-Mountain
Pigweed spp.
Milkweed
Indian Hemp
Mustard spp.
Mullein
Licorice
Vervain spp.
Goldenrod
Mint
Plantain
Horsetail

24
45
15
54
8
14
8
3
3
5
2
7
4
10
0
0
8
0
2
3
1
5
1
1
0
0
0
0
1

85
36
36
17
18
39
7
8
23
10
3
5
19
1
0
5
5
2
2
12
5
5
0
7
0
1
2
1
0

101
l3
35
0
17
7
45
35
25
3
3
3
5
1
0
1
1
2
0
9
18
1
2
1
1
2
0
0
0

33
4
45
2
24
11
0
10
3
40
16

9
0
6
3
0
1
0
0
2
0
0
0
1
6
2
4
0
0
0
0
0
0
1
0
0
1
4
0
0

6

0
3
30
12
1
0
1
0
0
3
5
14
22
0
0
12
1

112

3-6
Sand Baril
0
1
5
4
5
7
1
5
0
0
0
3
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0

1/

Total

Percent
Occurrence

252
99
142
80
72
79
61
61
56
58
24
24
29
22
32
22
15
4
5
24
24
14
9
23
23
4
6
l3
2

17.97
7.06
10.13
5.71
5.14
5.63
4.35
4.35
3.99
4.11
1.71
1.71
2.07
1.57
2.28
1.57
1.07
0.29
0.36
1.71
1.71
1.00
0.64
1.64
1.64
0.29
0.43
0.93
0.14

----------------------------------------------------------------------------------------------------------

I
N
(1:)
(1:)

I

�Table 21.

Forb occurrence on six sites along the Tamarack Management Area in late summer, 1972. l/ (continued)

Genera or species
Prickly Pear
Cocklebur
Russian Thistle
Canadian Thistle
ClannnyWeed
Tall Eve. Primrose
Prostrate Primrose
Butterfly Guara
Goatsbeard
Asparagus
Pro Coneflower
Gilia
Knotweed
Smart Weed
Unid. Composites
Sunnnation

6-7
West

14-15
West

3-6
East

Braddock
III '!:.../

Braddock
fI2

3-6
Sand Bar

o

o
o
o

15

o

1

o

1

o
o
o
o
o
o

o
o
o
o
o
o

o

o

o

1

o
o
o
o

o

o
o
o
o
o

1

o

3

o
o
o

1
1

o
1

o

6
3
1

o

o
o
o
o

o
7

1

o

o
o

30

o
o
o
o
o
o
1

o
o

11
2

o
1

1

o

o

o
o
o

o

Total
16
13
3
31
2
4
6
3
1
1

1
7

1

2

o

12

2

2

3

14

o

o

12
21

227

356

354

333

72

60

1,402

Percent
Occurrence
1.14
0.93
0.21
2.21
0.14
0.28
0.43
0.21
0.07
0.07
0.07
0.50
0.14
0.86
1.50

!/

Different sample sizes were obtained on different sites, therefore direct comparisons between or among
sites should not be made.

'!:.../

Braddock #1 sample was obtained in timbered flood plain.
cover along the south side of the flood plain.

}j

A frequently flooded site with the channel system of the river where annual forbs alone were resident.

Braddock #2 was taken in open grass-brush

I
N
CI)
1.0
I

�-290-

Table 22. Percentage occurrence of vegetation,
on several sites within the Tamarack Management
property near Merino, Colorado.

litter and bare ground
Area and on the Lutes

Location

6-7

14-15

3-6

Cover Type

West

West

East

Braddock
Pasture

Lutes
Property

Grass

38.24

32.35

31.75

23.70

62.17

Brush and Vines

5.84

11.17

21.30

14.51

7.95

Forbs

34.87

37.16

31.66

40.26

21.20

Dead Vegetation

6.30

3.76

3.04

0.24

2.41

Litter

7~07

5.64

8.05

4.72

4.10

Bare Ground

7.68

9.92

4.20

16.57

2.17

,

Table 23. Comparisons of height and density measurements
habitats of the Tamarack and Lutes properties.
Location

Site

Mean Height

in Inches

obtained

in quail

Mean Density

Tamarack

6-7 West

16.40

1.70

14-15 West

22.58

2.12

Braddock

Pasture

12.62

1.19

Braddock

Pasture-South

15.08

1.35

21.08

2.09

12.13

1.99

3-6 East

Lutes Property

Density is based on a scale of 1 to 3 with the latter representing
highest density.

the

1/

�-291-

Quail Winter Use in Relation

to Vegetation

On the Tamarack different areas were selected for measurement on the basis
of their occupancy by quail coveys during the previous winter.
Areas 14
and 15 on the West Tamarack contained several small coveys during the winter
of 1971-72, whereas no quail were found in Areas 6 and 7 in that section,
nor in Areas 3 to 6 on the East Tamarack.
The Braddock pasture, which is
lightly grazed yearlong, provided another situation for comparison.
The
Lutes property, which lies up-river near Merino (Snyder, 1972) receives
moderate to heavy grazing.
Quail winter use of specific locations along the Tamarack was not consistent
between winters of 1971-72 and 1972-73.
As a consequence, comparisons with
site vegetation cannot be made.
Little use of Areas l4-West and l5-West was
observed during the winter of 1972-73, however, a covey was resident on Area
7-West.
Undoubtedly the variables within hunting and weather play an important role as to the location of wintering coveys on the Tamarack.
Markedly fewer forbs were tallied on the Lutes property (Table 22), where
grazing evidently had an impact on quail populations.
Limited trapping in
that location during the fall of 1971 showed very few quail were resident
there (Snyder 1972).. Nearby locations, less heavily grazed, showed densities
comparable to those found on the Tamarack.
Therefore, it seems probable that
grazing was important in reducing quail density on the Lutes property.
Crouch (1961) observed significantly more bobwhite on his ungrazed Tamarack
Study A~ea than on his grazed plot located a few miles upstream.
The greatest
contrast was in summer and fall. No quail were observed on the ungrazed
section in winter, but observations are often difficult in heavy cover sites
after hunting season.
Crouch also recorded a significantly higher occurrence
of forbs on his ungrazed plot than on bis grazed study area.

Quail Movement

in Relation

to Vegetation

The previously mentioned Braddock pasture, lying east of the Crook River
Bridges (Fig. 3), has been lightly grazed.
As a result, brush cover is
observably open in many locations over-winter and apparently less attractive
to wintering quail.
Fall banding had been conducted within this pasture both
in 1971 and 1972. Harvest recoveries and winter retraps indicate many of
these quail apparently move either east or west to better cover sites for
winter.
Most recoveries have been obtained in Areas I-West and 2-West, indicating this is the primary direction of movement.
Since the Braddock pasture is not heavily grazed, numerous forbs are able
to reproduce each year.
Therefore, the site appears to be an attractive
brood rearing and feeding location through late summer and early fall.

Food Habits

in Relation

to Forb Occurrence

Wassink (1972) conducted an analysis
during the November-December
hunting

of 50 bobwhite quail crops collected
season in 1971 on the Tamarack.
Bait

�-292-

grain, corn and milo, was still present from trapping operations which
ended just prior to harvest. This undoubtedly biased results as 52 percent of the consumed food by volume was comprised of corn and milo.
Important natural food included knot weed, Polygonum aviculare, which
comprised 20 percent by volume and 80 percent by occurrence (Table 24).
Penny cress, Thaspia arvense, occurred in 58 percent of the crops and made
up 5 percent of the volume. Giant ragweed, Ambrosia trifida, occurred in
68 percent of the crops and totaled 5 percent by volume. Other lesser
species are illustrated in Table 24 as adapted from Wassink's report.
Quantities of forbs vary greatly from year to year depending on growing
conditions. Therefore, comparisons of 1971 crop analysis with 1972 forb
abundance can only be related in a general way.
Natural Mortality, Weather and Sex Ratio Changes
Information presented in previous sections of this report indicate that
natural mortality was quite low over-winter in 1971-72, but relatively high
during the spring and summer of 1972. Preliminary data indicate higher
natural mortality occurred in late fall and early winter, 1972-73.

Winter Mortality and Weather--Other studies conducted in northern and central parts of the species range show that winter mortality is directly proportional to the duration of snow cover on the ground over-winter. Roseberry
and Klimstra (1972) in Illinois, Latham and Studholme (1952) in Pennsylvania,
Kabat and Thompson (1963) in Wisconsin are among the primary examples. It
seems probable that this same stress factor applies to quail in Colorado.
If so, then low mortality in winter, 1971-72 and high mortality in winter,
1972-73 are to be expected. Table 25 shows that the first winter had only
16 days of snow cover compared to 75 days in 1972-73. Temperatures in
November and December of 1972 also ranged well below those for the same months
in 1971.

Spring and Summer Mortality--Mortality in spring and/or summer, 1972 was
relatively high, but apparently was not a significant detriment to productivity since the fall popUlation was higher in 1972 than in 1971. POSSibly,
this rate of mortality is to be expected. But why this mortality occurred,
just when it occurred, or how it occurred, remains unknown. It was significantly higher among hens than males (Chi-square = 5.83 with 1 d.f.). Numerous
authors have cited this higher loss of hens as probably attributable to hen
efforts in nesting and/or brood rearing. The prolonged period of hatch
illustrated in Fig. 5 indicates most hens probably made more than one nesting
attempt. Undoubtedly successive egg laying and nesting attempts induced stress
on hens and increased their vulnerability to predation, parasitism and other
mortality factors.

�-293-

Table 24. Principal foods of a sample of bobwhite quail collected on the
Tamarack during the late fall of 1971. !/

Percent by

Food Item

Occurrence

Volume

Polygonum aviculare

80

20

Giant Ragweed

Ambrosia trifida

68

5

Field Penny-cress

Th1aspia arvense

58

5

Sunflower Spp.

Helianthus spp.

58

4

Corn

Zea mays

48

32

Curled-leaf Dock

Rumex crispus

44

T

Wild Buckwheat

Polygonum convolvulus

40

4

Halberd-leaved
Orache

Atriplex patula hastata

38

1

Milkweed

Asclepias spp.

30

1

Snow-on-the-Mt.

Euphorbia marginata

22

T

Kochia

Kochia scoparia

20

1

Sorghum

Sorghum vulgare

18

18

Pigweed Spp.

Amaranthus spp.

16

T

Green Foxtail

Setaria viridis

14

3

Sweet Clover

Melilotus spp.

14

T

Buffalo Burr

Solanum rostratum

4

T

Poison Ivy

Rhus radicans

4

T

Witchgrass

Panicum capil1are

4

T

Vervain

Verbena stricta

2

T

Smart Weed

Polygonum pennsy1vanica

2

T

Russian Olive

E1eagnus angustifolia

2

T

Night Shade

Solanum spp.

2

T

Common Name

Scientific Name

Knot Weed

!/

Data revised from Wassink (1972) include samples from 50 bobwhite.

�-294-

Table 25. Average monthly temperature, snowfall and days with snow on
ground during the winters of 1971-72 and 1972-73 !/.
1971-72

1972-73

Snowfall

Days of
Snow Cover

50.1

3.3

1

November

39.1 l:/

1.0

December

29.2

January

Snowfall

Days of
Snow Cover

50.0

0.5

1

2

29.8

29.0

25

3.0

5

19.6

6.0

23

24.4

5.0

8

24.6

13.5

9

February

36.4

T

0

31.7

3.0

9

March

45.8

0

0

39.6

6.0

4

12.3

16

58.0

75

Month

X Temp.

October

Total
X Temp.

37.5

X Temp.

32.5

1/
- Data from U. S. weather station located 5 miles south of Sedgwick,
Colorado.
2/
- Underscoring denotes large temperature differences between the two
winters.

Harvest as a Limiting Factor
Quail in the bag plus crippling loss approximated 26 to 27 percent of the
original fall population in both 1971 and 1972. This harvest appears relatively
low and not of significant impact in itself when contrasted to other studies.
Rosene (1969) recommended that combined harvest and winter loss not exceed 45
percent of the fall population in northern states. The combined 1971-72 loss
approximated 43 percent on the Tamarack.
Vance and Ellis (1972) presented data showing that harvest rates of 60 to 70
percent, excluding crippling loss, were not detrimental to subsequent population levels over a seven year period. They also found that low winter losses
occurred after high harvest rates and high winter losses followed lower harvest
rates. They concluded that 70 percent harvest was a safe level in southern
Illinois, but stated that this rate of harvest would not be attained during
years of low population. After their presentation at the First National Bobwhite Quail Symposium at Stillwater, others questioned the possibilities of
immigration of surrounding quail to potentially help sustain these study area
populations.

�-295-

Roseberry and Klimstra (1972) reported an average harvest of about 44
percent on another southern Illinois population without apparent detriment. November to April mortality totaled 66 percent of the original
population on the hunted area compared to 54 percent decline on a nearby
unhunted population.
Hunting stress may be additive in increasing mortality when combined with
snow cover and extreme cold. These conditions occurred on the Tamarack
during the late fall of 1972, but over-winter mortality rates are not yet
certain. Frequent covey disturbance, which often occurred on the Tamarack,
would undoubtedly have much greater impact than only occasional covey
disturbance on less intensively hunted sites. Rosene (1969) stated that
bobwhite have a low number of feathers and are quite susceptible to cold
and exposure when their feathers become wet.
Snows and cold weather came unusually early in the fall of 1972. Short
days, long nights, snow and severe cold usually are not a significant problem until mid-December in Colorado. Therefore, it is recommended that
quail seasons begin earlier, preferably around the first of November,
and run c9ntinuous to termination. Most years this would put harvest
ahead of the main stress period, and also permit harvest of some quail
otherwise lost to natural mortality.
Early Hatched Versus Late Hatched Survival
The stress of November snow and severe cold and snow cover that persisted
into December, 1972 potentially induced increased mortality among late
hatched quail. Some of these birds were not fully grown or feathered at
the start of the hunting season on November 11. A sample obtained from
banded quail taken during the December (extended) season and winter trapping
operations is illustrated in Fig. 5. As shown in the illustration, there
is little indication of excessive mortality among younger juveniles. The
data show that these birds survived the adverse weather about as well as
the older juveniles did.

LITERATURE CITED
Crouch, G. L. 1961. Inventory and analysis of wildlife populations and
habitat, South Platte River Valley. Colo. Game and Fish Dept. Fed.
Aid Proj. W-l04-R. Job Compl. Rpt. July. 68 p. Mimeo.
Davis, D. E. 1960. Estimating the numbers of game populations, pp. 5.15.27. In H. S. Mosby (Editor) Manual of Game Investigational Techniques.--The Wildlife Society, Washington, D. C.
Edwards,W. R. 1972. Quail, land use, and weather in Illinois, 1956-70.
National Bobwhite Quail Symposium. 1:174-183.

�-296-

Ellis, J. A., K. P. Thomas, and P. Moore. 1972. Bobwhite whistling
activity and population density on two public hunting areas in
Illinois. National Bobwhite Quail Symposium. 1:282-289.
Hoekstra, T. W., and C. M. Kirkpatrick. 1972. The bobwhite quail on
the Crane Naval Ammunition Depot, Indiana -- 25 years of protection
and plant succession. National Bobwhite Quail Symposium. 1:184-194.
Kabat, C. D., and D. R. Thompson. 1963. Wisconsin quail, 1834-1962,
population dynamics and habitat management. Tech. Bull. No. 30.
Wisc. Cons. Dept., Madison. 136 p.
Latham, R. M., and C. R. Studholme. 1952. The bobwhite quail in
Pennsylvania. Pennsylvania Game News. Special Issue 4, 95 p.
Robson, D. S., and H. A. Regier. 1964. Sample sizes in Peterson markrecapture experiments. Trans. Amer. Fish Society. 93(3):215-226.
Robel, R. 1965. Differential winter mortality of bobwhite in Kansas.
J. Wildl. Mgmt. 29(2):261-266.
Roseberry, J. L., and W. D. Klimstra.
of a hunted bobwhite population.
1:268-282.

1972. Some aspects of the dynamics
National Bobwhite Quail Symposium.

Rosene, W.
N. J.

Rutgers Univ. Press, New Brunswick,

1969. The Bobwhite Quail.
418 p.

Snedecor, C. W. 1956.
Ames. 534 p.

Statistical methods.

The Iowa State ColI. Press,

Snyder, W. D. 1967. Experimental habitat improvement for scaled quail.
Colo. Dept. of Game, Fish and Parks. Tech. Bull. No. 19. 65 p.
1972. Population status of bobwhite quail in eastern Colorado.
Colo. Div. of Game, Fish and Parks. Game Res. Rpt. April. pp. 121-150.
Vance, D. R., and J. A. Ellis. 1972. Bobwhite populations and hunting on
Illinois public hunting areas. National Bobwhite Quail Symposium.
1:165-174.
Wassink, J. L. 1972. Fall food habits of bobwhite in eastern Colorado.
Colo. State Univ., Dept. of Fishery and Wildl. Biology. Mimeo Rpt.
9 p.

~J

I

)

I

~;

)

Prepared by _-.:"
/:"::-::-':.../_/_~/:...('-=,_) _"-:I):-1_/-::-_/_J_, _I '-I' _~&amp;_''Zh_,-,_
_
Warren D. snyd~r
Wildlife Researcher

J

�April, 1973

-297-

JOB PROGRESS REPORT

State of

COLORADO
------~~--~-------------

Project No.

W-37-R-26

Work Plan No.

17

Job Title
Period Covered:
Personnel:

Game Bird Survey

Job No.
4
Migration of White-tailed
Ptarmigan to and from a Major Wintering Area

----~~~~------------------------

April 1, 1972 through March 31, 1973

Richard W. Hoffman, Clait E. Braun and Terry A. May.

ABSTRACT

Studies on migration of white-tailed ptarmigan (Lagopus leucurus) were initiated
in Colorado in the spring of 1972. Major emphasis was to document movements by
sex and age class from and to a major wintering area in order to ascertain the
importance of such areas to surrounding breeding sites and the distances and
directions involved in annual movements. Guanel1a Pass was the major wintering
area investigated. Of 147 birds individually identified on the wintering area
from October 1, 1972 to March 31, 1973, 114 (77.4 percent) were females, while
42.3 percent were young of the year. Movement data were available for 58
ptarmigan. These data indicated females of all age classes moved significantly
greater distances than males, with subadult females moving longest distances to
breeding or summering sites. Both adult and subadult males were relatively
.sedentary, showing a high fidelity to natal areas and generally moving less than
2,air miles. However, males became more mobile upon cessation of breeding
activities. Average female movements were 3.62 air miles, but 3 subadults moved
over 10 air miles to breeding sites. Wintering areas for females are believed
to be limited; thus forcing hens to move greater distances to breeding sites.
The longer movements of subadults was attributed to their wandering in search
of available breeding areas. Significantly more birds (16 of 58 recoveries)
~ved northwest from Guanella Pass than in any other direction. In addition,
the longest movements were also recorded in this direction. Seventy percent
(11) of the birds moving northwest were subadults, primarily females. Only
7 birds moved in any other single direction. Birds could move over 15 air
miles northwest across continuous alpine habitat, which was probably the primary
reason for documenting significantly more movements in this direction. Approximately 44 percent of the birds banded each winter returned the following winter.
It was concluded that Guanel1a Pass attracted major numbers of ptarmigan,
especially females, from 15 to 20 square miles of surrounding breeding habitat.

�-298-

RECOMMENDATIONS

1.

Efforts should be increased to trap and band chicks in order to document
movements from natal areas to wintering and breeding sites.

2.

Major wintering areas for ptarmigan should be protected from motor vehicle
use and livestock grazing.

3.

Additional efforts should be made to investigate other wintering areas
in the State to fully assess sex ratios in differing types of habitats.

�-299-

MIGRATION OF WHITE-TAILED
PTARMIGAN TO AND FROM A MAJOR WINTERING AREA
Richard W. Ho ffman
White-tailed ptarmigan, the smallest member of the Tetraonidae, live
primarily in alpine habitats of western North America. It occupies
suitable habitat from south-central Alaska, the Yukon, British Columbia,
western Alberta, and Vancouver Island sout.hwa'r
d to Mt. Ranier and the
Cascade Mountains of Washington, east to western Montana, and south in the
Rocky Mountains to northern New Mexico (Aldrich 1963). The distribution of
ptarmigan in Colorado correlates with the distribution of alpine tundra
from 11,000 to over 14,000 ft in elevation, an area of approximately 3,750
square miles (Braun and Rogers 1971). Studies of ptarmigan have been conducted at Guanella Pass since 1966 and relate primarily to bird collection,
description of habitat and general winter ecology (Braun and Schmidt 1971).
Some banding of this winter population had been accomplished as about 350
individuals were marked prior to the initiation of this study. Only 11 band
recoveries were documented prior to 1972.
The intensively studied Mt. Evans area (Braun and Rodgers 1971) lies 3 miles
east of Guanella Pass. Prior to 1972, only one bird banded at Guanella was
recovered on Mt. Evans, while two birds from Mt. Evans were recaptured at
Guanella during the winter. Most of the other bands recovered (mainly biy
hunters) indicated birds wintering at Guanella Pass moved northwest with
females novang the greatest distances. The extent of the movements was
not known, although banded females had been recovered 6 and 10 miles away.
Previous banding studies at Guanella indicated that approximately 80% of
the 200-300 birds wintering there were females. It is known that breeding
densities are remarkably stable between years, averaging 15 to 20 birds per
square mile, of which one half are females. Thus, initial indications were
that Guanella Pass must be supplying breeding females to 15 to 20 square
miles of surrounding breeding areas (Braun and Schmidt 1971).
The question investigated concerned the annual migration of ptarmigan,
principally females, to and from a major wintering area and involved understanding the importance of such an area to surrounding breeding sites.

P. S. OBJECflVE
To document lOOvements of white-tailed ptarmigan by age and sex class from
and to a major wintering area in order to evaluate importance of such areas
and distances involved in annual migrations.
SEGMENT OBJECfIVES
1.

Individually identify at least 100 different birds during the winter
period at Guanella Pass.

�-300-

2.

Locate winter banded birds on breeding
of established census techniques.

3.

Compute distances and directions involved in normal
from wintering areas by age and sex class.

4.

To compile

data and prepare

progress

and summering

areas through use

migrations

to and

report.

In addition, attempts were made to: (1) document departure and arrival
dates, (2) record and analyze flock sizes, (3) describe the distribution of
birds on the wintering area, (4) determine the affinity of birds for the
wintering area, and (5) individually identify at least 50 chicks on summering
areas in order to document their movements to wintering sites.
Limited observations were made at four smaller wintering sites in the Georgetown-Montezuma
area. This was done in order to document movements of birds banded at Guane11a Pass to other winter use sites, locate chicks banded during the summer,
estimate numbers of birds using these sites, and estimate sex and age composition of winter flocks utilizing the areas.

REVIEW OF LITERATURE
Intensive studies of white-tailed ptarmigan in Colorado (Braun 1969, Haskins
1969, Schmidt 1969. and May 1970, 1973) have provided extensive data on general ecology, population dynamics, movements, habitat, seasonal foods,
behavior and parasites.
Weeden (1959, 1967) and Choate (1960, 1963a and b)
have also studied this grouse.
Other than these studies, the literature
contains limited scientific references to this species.
Seasonal movements have been described in general terms by several authors.
Packard (1945), Racey (1948), Bailey (1927), Bent (1932), and Gabrielson
and Lincoln (1959) describe the species as spending the summer (breeding)
above timberline descending to or below timberline in winter.
Choate (1960)
suggested that some flocks remain above timberline in winter, while others
descend to areas dominated by willow (Salix spp.) and poplar (Populus spp.)
below timberline.
Weeden (1964) indicated male (and some female) rock
ptarmigan (L. mutus) spend the winter at the upper fringe of timber, moving
higher to alpine habitats for breeding.
He found that females (and some
males) primarily wintered along stream courses dominated by willow 10 to 20
miles from the nearest alpine breeding areas.
Irving et al. (1967) extensively monitored the migration of willow ptarmigan (L. lagopus) at
Anaktuvuk Pass in the Brooks Range of Alaska.
It was documented that willow
ptarmigan, expecially females, may move 200 miles south of their northern
breeding range to winter in willow dominated stream bottoms below timberline.
Adult males predomianted in winter in their northern range near breeding
areas.
Evidence indicated each sex and age category moved according to its
own program.
Braun (1969) documented movements of white-tailed ptarmigan throughout the
year, however, he suggests his data for the fall and winter periods are
limited and may not represent true situations.
His studies indicated that

�-301-

males are relatively sedentary, usually moving less than one mile in a
given year, while females are highly mobile with some birds moving 10 to
15 miles from wintering to breeding areas. He also found that subadults
move farther than adults, with subadult females moving the greatest
distances.
Techniques for locating, capturing, aging and sexing white-tailed ptarmigan
have been described by Braun and Rogers (1971)', and Braun et al. (1973).
The telescoping noose pole used to capture ptarmigan was adapted from
Zwickel and Bendell (1967).
Braun and Schmidt (1971) intensively studied wintering populations of whitetailed ptarmigan.
Objectives were to describe the habitat and environmental
features typical of wintering areas, determine the composition of winter
flocks, and delineate behavior and movements of individual birds and flocks
in relation to changes in environmental conditions.

Study Areas
Guanella Pass was chosen for study principally due to an abundance of
wintering ptarmigan at this site, year round accessibility, and availability
of information on this site from prior studies.
Four additional smaller
wintering areas were briefly investigated.
These were Waldorf, Naylor Lake,
Stevens Gulch and Horseshoe Basin.
The Guanella
Pass area is approximately 5 air miles south of Georgetown,
Colorado, in the Pike and Arapaho National Forests, Clear Creek and Park
counties, T5S, R74W.
Topography of the area is gentle, with slopes varying
from 5 to 30% and elevation extremes of 11,400 to 11,990 ft above sea level.
Total area investigated was 3.5 square miles (Fig. 1).
Surrounding breeding and summering sites, referred to as the MbntezumaGeorgtown study area, were searched for birds banded at Guanella Pass.
This
area encompasses approximately 113 square miles (Fig. 2), however, probably
only 25 to 30 square miles are suitable as breeding and summering sites.
Alpine areas searched included parts of T4S, R75 and 76W; T5S, R73, 74, 75
and 76W; and T6S, R73, 74, 75 and 76W in Clear Creek, Park and Summitt
counties.
Topography of the area is irregular, varying from sharp peaks
over 14,000 ft to gently rolling expanses of hundreds of acres.
This area
included the four additional wintering areas that were briefly investigated.

METHODS

AND MATERIALS

Locating

Ptarmigan

Winter flocks of ptarmigan were located by concentrated searching on foot
with periodic stops to scan surrounding areas for birds or their sign.
During the breeding and brood seasons birds were located by the use of a

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-302-

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Fig. 2.

MOntezuma-Georgetown

study area.

�-304-

portable cassette-type tape recorder utilizing
"distress" calls (Braun et al. 1973).

male "challenge"

and chick

The male "challenge" call was used during the breeding season to locate
territorial males. Non-territorial males usually did not respond, but sat
quietly or fled from the area, consequently many were probably not located.
Once territorial males were located, their females could usually be found
nearby.
At times, upon hearing the call, the male would leave the female
and fly to vantage points on the boundary of his territory where he would
commence calling.
MOst frequently, males would return to their hens in 10
to 15 minutes, but on several occasions an hour or more was ~pent before
a male would return to his mate.
Use of tape recorded chick "distress" calls resulted in the location of
females with broods.
OccaSionally, females without chicks responded, and
it is believed these hens had recently lost their broods.
Intensity with
which females responded varied with age of chicks, and closeness to the
source of the distress call. In general, as chicks got older, females were
less likely to respond.
Males and unsuccessful females congregated in flocks along rocky ridge tops
following the breeding s.eason (Bz atm 1971).
These groups were located by
intensively searching edges of rocky areas along ridge tops. Once birds
were located, they were observed through 7 X 50 binoculars, classified as
banded or unbanded, and during the winter, the number of birds comprising
each flock was determined.
Unbanded birds located during the winter were
pursued until caught or flushed and not relocated.
Only unbanded females
(and chicks) were captured and banded during the spring and summer.
Attempts
we~e made to identify previously banded birds by reading the bandette
number.
If not readable, efforts were made to recapture the bird.
In
Winter, band numbers would frequently be obscured by leg feathers or snow.
Only 2 banded ptarmigan were observed away from wintering areas that could
not be identified or caught.
Each observation was recorded at the time on
standardized observation cards.
All locations were subsequently plotted
on 7.5 minute U.S.G.S. topographic maps.

Capturing

and Banding

Ptarmigan

Ptarmigan were captured with a 16 ft telescoping noose pole described by
Zwickel and Bendell (1967). Nooses were made from 60 pound nylon covered
leader wire.
For chicks older than 2 weeks, 18 pound test was more suitable.
Known mortality due to this technique was less than one percent.
Upon
capture, birds were aged and sexed by techniques described by Braun and
Rogers (1971). Each bird was banded with a gold, aluminum, seriallynumbered size 8 band.
Chicks were banded on the right leg only, with this
band.
In addition, adults were banded with numbered, colored, plastic
bandettes (no. 6).

�-305-

Delineating

Distribution

of Ptarmigan

Distribution of ptarmigan on the wintering area was determined by periodically traversing the entire area and from reports of sightings by reliable
observers.
Sites were classified as being utilized if signs such as
droppings, tracks, roosts, or the birds themselves were observed.
Data
were available for all winters since 1966. Although all locations on the
wintering areas were not equally searched, it is believed that during the
6 years for which data were available, all sites being utilized by ptarmigan
were noted.

Documenting

Movements

of Ptarmigan

Movements of birds banded at Guanella and recovered on breeding or summering
areas were plotted on 7.5 minute USGS topographic map. A straight line was
drawn from the banding site on the wintering area to the recovery or reobservation location.
Thus, all movements represent minimum distance traveled
in air miles.
Directions were determined from the wintering area to the
recovery or reobservation site.
All initial ages were taken as the age of
the bird when banded on the wintering area. Additional information recorded
included: (1) date banded and recovered, (2) band numbers, (3) sex, and (4)
years returning to the wintering area.
If a bird banded at Guanella in 1969 was reobserved on the wintering area in
1972 it was assumed this bird also returned in 1970 and 1971 and will return
in following winters until it is Los't from the population.
Birds banded at
Guanellaand
reobserved on breeding or summering areas, but not reobserved
at Guanella, were assumed to have dispersed from the area and were designated
as utilizing other wintering sites.
Extensive efforts were made to contact hunters in the area primarily to
obtain information on locations where banded birds were harvested.
Data
were also available from hunter recoveries, collections, and observations
from years prior to the initiation of this study through the courtesy of
C. E. Braun, Colorado Division of Wildlife.
Arrival and departure
study area in October

Analysis

dates were documented
and April.

through periodic

search of the

of Data

Statistical methods employed in the analysis of data were tested at the .05
level of significance through use of chi square and students t tests.
RESULTS AND DISCUSSION
Composition

of the Wintering

Population

Six hundred and sixty eight individual bd r ds were identified on the Guanella
Pass study area from 1966 through 1973 (34 in 1966-67, 66 in 1967-68, 43 in
1968-69, III in 1969-70, 125 in 1970-71, 142 in 1971-72, and 147 in 1972-73).

�-306-

Winter on the alpine was arbitrarily designated as the period starting in
late October when ptarmigan became entirely white through mid- to late April
when the prenuptial molt of males was initiated.
Present estimates indicate
200 to 300 birds utilize this site each winter.
Males comprised 21.7% of
the birds handled since 1967 (27.3 in 1967-68, 18.6 in 1968-69, 21.6 in 196970, 19.2 in 1970-71, 21.1 in 1971-72, and 22.6 in 1972-73).
Percent immatures varied considerably with year (45.5 in 1967-68, 30.2 in 1968-69, 21.6
in 1969-70, 38.4 in'1970-7l, 43.7 in 1971-72, and 42.3 in 1972-73) and
averaged 36.9% (Table 1). Thus, Guanella Pass was utilized principally by
females (78.3%) and primarily adults, varying from a high of 78.4% in 1969-70
to a low of 54.5%
in 1967-68.
The number of recaptures (birds banded in previous winters) varied with year
but there was a tendency for percentage of recaptures to increase in succeeding winters (Table 1). This would be expected, as the number of banded birds
in the population increased each year.
Initial investigations in 1973 at Horseshoe Basin located 5 air miles northwest of Guanella Pass, revealed this area was primarily utilized by males.
Twelve birds were identified, of which 10 were males (83.3%).
This was the
reverse of the situation at Guanella Pass. The primary difference between
Guanella Pass and Horseshoe Basin appears to be availability of willow.
Less
wind action in Horseshoe Basin allows snow to accumulate and cover much of
the willow.
Brief observations at the Waldorf wintering site, located 4 air miles northwest of Guanella Pass, indicated no apparent differences in percent of males
and females utilizing the area. Of 11 birds identified, 6 were males (54%)
indicating flocks were comprised of approximately equal numbers of both sexes.
However, sample sizes were small and may not reflect the true situation.
Winter

Flocks

Like many grouse, ptarmigan are gregarious birds.
The basic population unit,
the flock, is fragmented temporarily by territorial activities accompanying
reproduction.
In winter it is known that segregation of sexes, while not
complete, results in the separation of a large proportion of the males and
females by habitat types (Braun and Schmidt 1971). Weeden (1964) reported
similar findings for rock and willow ptarmigan.
Weeden also examined 17
white-tailed ptarmigan shot by hunters from one flock near Summit Lake in
the Alaska Range, and found that 14 were hens.
Choate (1963) suggests a
slight tendency toward sexual segregation by white-tailed ptarmigan in
Montana, but he had little information on winter flocks.
Braun and Schmidt
(1971) documented sex segregation of wintering birds at Guanella Pass.
One hundred and sixty-four flocks were observed on the Guanella Pass study
area from 1966 through April 1973 (16 in 1966-67, 18 in 1967-68, 9 in 1968-69,
29 in 1969-70, 24 in 1970-71, 33 in 1971-72 and 35 in 1972-73.
Winter
flock ,size varied from less than 5 to over 75 birds, but most flocks contained either less than 10 or between 11 to 25 birds (Table 2). Comparison
of flock size by months revealed no significant differences (Table 3).

�-307-

Table 1.

Composition of the Guanella Pass population.
Sex and Age Com,Eosition
Percent
Percent
Percent
Percent
Recaptures
Males
Females
Juveniles

Total
Identified

Winter

Percent
Adults

1967-68

54.5

45.5

72.7

27.3

6.0

66

1968-69

70.0

30.2

81.4

18.6

16.2

43

1969-70

78.4

21.6

78.4

21.6

16.2

III

1970-71

61.6

38.4

80.8

19.2

24.8

125

1971-72

56.3

43.7

78.9

21.1

26.8

142

1972-Apr.11,' 73

57.7

42.3

77.4

22.6

34.3

147

Average

63.0

36.9

78.3

21.7

20.7

1/ The winter of 1966-67 was excluded due to small sample sizes.

Table 2.
to 1973.

Number of
Flocks

Size of flocks observed at the Guanella Pass wintering area, 1966

1-10

11-25

26-40

Flock Size
41-60

61-75

Over 75

74

47

30

7

2

4

Similarly, no significant differences in flock size were apparent between
the winters of 1969-70, 1970-71, 1971-72, and 1972-73 (Table 4). Due to small
sample sizes, data from the winters of 1966-67, 1967-68, and 1968-69 were not
included in the analysis.
Since 1966, only 11 single birds have been observed at Guanella Pass. Eight
of the 11 were males. Flocks of males were usually small (less than 15 birds)
while females generally congregated in flocks of larger size. Possibly there
is an optimum flock size for males and females most beneficial for utilization
of food supplies and escape from predators.

�-308Table 3. Size of winter flocks observed at Guane11a Pass, pooled
data, 1966-1973.

October

Number of Birds Per Flock Per Month
November
December
January
February

March

April

16
14
12
12
18

35
6
3
13
35
12
25-30
40
31+
5
5
15
7
8
35+
20
30+
60+
15+
20
35
15-16
5-6
10
15
9
25
1
35-40

75+
7
5
30+
3
30+
5
40+
5
11
75+
14
7-8
60+
50+
15+
2
40
30
20+
15

8
2
19
2
2
5
23
11
6
4
12
5
42
3
15
7
70
2
1
30
5-6
5-6
9
53
35-40
11
5-6

40
14
6
10
1
9
6
4
12
4
12-15
1
20
5
12
30
25
75

40
1
6
75
2
12
6
25
8
30
3
50-60
3
4
3
5
30
26
21
30
6
13
15-20
4
8
37
35
11
4

58
20
1
15
20
1
60
42
2
1
12
2
4
4
18
5
15
1
1
12
35
8
5
11
2
30
11
20
2
1
17
10
3
3

Total
Flocks

5

29

21

27

18

29

34

Mean

14.4

19.9

25.7

14.7

16.0

18.0

13.3

�-309-

Table 4.

Size of winter flocks by years, Guane11a Pass.

Years
1969-70

1972-Apr.11, '73

1970-71

1971-72

35
15
20
35
5
30
3
2
2
5
23
11
14
6
2
10-14
3
50-60
3
4
3
5
12
2
4
4
18
5
11

12
12
6
3
13
35
15
5
30
5
40
15
11
6
6
4
12
5
42
3
4
12
30
15
18

18
12
25-30
40
10
15
5
10-12
2
40
6
10
1
4
12-15
1
20
6
26
20-22
25-35
6
13
1
2
20
2
11
5
8
35
12
1

31
5
5
15
9
25
1
35-40
75+
14
7-8
30
20
15
15
7
70
2
5
12
30
25
75+
15-20
4
8
35-40
35
4
10-12
1
17
10
3
3

9

29

24

33

35

30.9

12.0

14.2

13.2

19.5

1966-67

1967-68

1968-69

7
8
35+
1
6
30+
5-6
5-6
8
2
19
9
25
8
58+
30+

16
20
30+
75+
7
60
50
9
53
40
1
30
20
1
15
20
1
60

14
60
35-40
35-45
6
75+
40-45
2
1
14

Total
Flocks

15

18

Mean

16.1

28.2

�-310-

Differences in flock sizes of males and females also may possibly be related
to differences in winter habitat utilized.
Females typically winter in
areas where willow is dense, thus, food supplies are abundant and they can
afford to congregate in large groups.
Males. winter at higher elevations
where willow is sparsely distributed.
If males congregated in large flocks
they possibly could over utilize food supplies.
Braun and Schmidt (1971) noted some indication that larger flocks were
comprised of discrete subunits of up to 15 birds all of the same age and sex.
This feature was also noted in the current study, but was not fully investigated.
Individual birds were not always associated with the same flock. Although
numbers can be cited, some ptarmigan banded at Guanella were found several
times throughout the winter associated with flocks of various sizes comprised
of different birds at different locations.
There appeared to be no aggressive behavior towards a new member of a flock. Irving et al. (1967) noted
similar behavior for willow ptarmigan in Alaska.
Birds favored certain
localities on the wintering area, but the same group of birds could not
always be located at these sites.
At Horseshoe Basin flocks of primarily males, were scattered throughout the
area with the largest flock comprised of 14 birds.
This suggests that males
do not congregate in large flocks typical of wintering females.
Due to
the scarcity of exposed willow, this area probably could not support more
than 50 birds.
At Waldorf all birds were found utilizing areas above 12,000 ft
where willow was sparsely distributed and large areas we'f'edevoid of snow.
Although more willow was available at lower elevations, birds or sign were
not observed below 12,000 ft. This area was investigated in mid-April about
the time birds depart from wintering areas to breeding sites.
Possibly
females utilized the lower areas earlier in the winter but had vacated these
areas and moved uphill, joining males in flocks closer'to breeding sites.
Thus, sex segregation may have been evident on the Waldorf wintering area
earlier in the winter.
Visual estimates indicate Waldorf has more willow
available than Horseshoe Basin and this possibly could mean more birds,
especially females, utilize the area. However, both areas encompass only
about 1.5 square miles each and could not support large numbers of birds.

Movements
Movements

of Chicks

Sixty chicks were individually marked from July through September, in order
to ascertain movements to wintering areas. Thirty-five of these were banded
within 2 miles of Guanella Pass, while the others were banded from 2 to 10
miles away. Of the sixty-one juvenile birds identified at Guanella Pass from
late October through April none were banded as chicks the previous summer.
In addition, no banded chicks were observed on other wintering areas investigated.

�-311-

Two birds, a male and a female, banded as chicks at Mt. Evans in 1971, were
recovered at Guanella Pass in 1972-73.
The movement was 4 air miles W for
the female, and 3.46 air miles WNW for the male. Neither bird had been
observed on a territory since banded.
Five banded chicks were known to be harvested by hunters during the 1972
hunting season.
All were shot within a mile of where they were banded
during the summer.
Since chicks suffer a 60 to 70% mortality during their
first year of life, it can be assumed that a considerable number of the
chicks banded were no longer alive by late winter.
Studies by Braun (1969) indicate that after brood breakup in late Septemberearly October, groups of chicks wander widely over the tundra.
Of those
females banded as chicks, only two were located in subsequent years.
One
was located as a transient 6 miles from where originally banded, while the
other was established on a breeding territory approximately 12 miles from
original banding location.
Braun (1969) also found that males banded as
chicks showed a tendency to return to natal areas and no males banded as
chicks were observed during the breeding period more than 1 mile from where
initially banded.
He suggested that undoubtedly some males, banded as
chicks, become established some distance from natal areas but were not
located due to relatively small size of areas studied «3
sq miles).
Contrary to males, females banded as chicks seldom return to natal areas.
This behavior of females allows for gene flow and results in the integrity
of the subspecies of white-tailed ptarmigan in Colorado.
Bendell and Elliot (1967), Gullion and Marshall (1968), Zwickel et al.
(1968), and Robel et al. (1970) have reported increased movements of juvenile grouse in the fall and have interpreted this increased mobility as
dispersal.
If increased movements of juveniles in the fall actually
represents dispersal, this may act as a population regulatory mechanism.
Documented mortalities for white-tailed ptarmigan suggest that mortality
occurs primarily in May and June and in September and October (Braun 1969).
Most ptarmigan lost in the fall are young of the year.
This mortality is
probably attributable to increased mobility of juveniles at this time.
If
such mortality is representative for dispersing juveniles, then dispersal
could be an effective regulatory force.
Locations where ptarmigan banded as chicks spend their first winter are
unknown.
It is known that chicks, especially females, are highly mobile
and wander great distances during fall and early winter.
Possibly some
chicks continue their nomadic life throughout the winter and wander between
wintering areas. This is suggested from data collected at Guanella Pass
during April when small flocks of unbanded juveniles were often encountered.
Possibly competition for space on wintering sites results in few sub adults
becoming established, and forces others to continue to wander.
Larger
sample sizes of banded chicks and increased surveys in other wintering
areas may be necessary to counterbalance high mortality rates and area
involved if chick movements to wintering areas are to be documented.
Directions

Moved

Fifty-eight movements were documented from Guanella Pass to territories or
summering areas (18 adult females, 21 subadult females, 9 adult males, and

�-31210 subadult males).
Directions were determined from banding location on
the wintering area to the summer recovery site.
In addition, three directions were recorded on movements of birds from the Mt. Evans area to breeding
or summering sites in the Guanella Pass area (adult female moving NW, adult
female novfng NE, and adult male moving SW).
Two females banded on territories in 1972 were reobserved at Guanella Pass
during the winter of 1972-73.
Since experienced birds generally return to
the same breeding site each year (Schmidt 1969), it was assumed these birds
would return to the same territory from Guanella in 1973.
Consequently, the
direction recorded was from Guanella Pass to the territory in order that
these movements be comparable with known movements from Guanella Pass.
If a bird was observed on a territory and summering area, two directions were
recorded; one from the wintering area to the territory and the other from
the wintering area to the summering area. Two males (G3 and B99) and one
female (YlOO) were observed on both breeding and summering sites.
Directions moved by sex and age class of birds banded at Guanella Pass and
recovered on breeding or summering areas are given in Table 5. Upon examination of these data it is evident that significantly more birds moved
northwest.
Twenty-eight percent (16) of the 58 known movements were in
this direction.
In addition, 70% (11) of the birds moving northwest were
.sub adults •
Distances moved in each direction are presented in Table 6. As with direction, the longest movements were also to the northwest.
Birds could move
approximately 15 air miles northwest across continuous alpine habitat, while
maximum distance they could IOOve in any other direction without encountering
unsuitable habitat, mainly forest, was 7 miles.
Thus, subadult birds leaving
Guanella Pass in late April were more likely to find vacant breeding territories to the northwest since the largest amount of alpine habitat occurred
in this direction.
The longest movements recorded were by 3 subadult females
moving over 10 air miles northwest to the Loveland Pass area.
No birds were recorded moving east and only one bird was documented to move
in each of the following directions: north-northeast,
east~southeast,
southeast, and south-southeast.
The only significant IOOvements in the easterly
direction was northeast (four birds).
Thus, birds departing from Guanella
Pass in early spring preferred to move in the westerly directions.
Ptarmigan investigations on the Mt. Evans area, which lies 3 miles east of
Guanella Pass, indicated breeding densities of ptarmigan were at a maximum.
in 1972 due to a closure of the area to hunting in 1970 and 1971 (Braun 1973).
It was originally believed that Guanella Pass birds were not moving east
because they were unlikely to find available breeding sites. However,
breeding densities prior to the closure (1966, 1967, and 1968) were the
lowest reported in the state (Braun 1969) and considerable breeding sites
were available.
Only 2 birds banded at Guanella were recorded moving to
the Mt. Evans area prior to the closure, consequently, neither the high
breeding density in 1972 nor the low densities in earlier years were reasons
why few Guanella Pass birds moved east to the Mt. Evans area.

�Table

5.

Directions

moved by ptarmigan

from the

Guane11a Pass wintering

Direction
SSE
S

Sex

Age!./

N

NNE

NE

ENE

E

ESE

SE

Female

2+

1

0

1

0

0

0

0

1

Female

1-

1

1

1

1

0

0

1

Male

2+

0

0

1

1

0

0

Male

1-

1

0

1

1

0

1

area

to territories

WSW W

or summering

SSW

SW

WNW NW

1

0

1

4

2

2

4

0

0

0

1

1

2

3

1

8

1

0

0

1

0

1

1

1

0

1

1

0

0

0

2

0

0

1

1

3

0

3

12

1

sites.

NNW

I
Vol

0

0

1

1

1

1

2

6

5

Females

2

1

Total

Males

1

0

2

2

0

1

0

0

1

2

1

1

2

1

4

Total

Both Sexes

3

1

4

3

0

1

1

1

2

3

3

7

7

4

16

-l/ Age when banded.

2

1

Total

•....

Vol

I

1
,

2

�Table 6.

Distances moved by ptarmigan from Guane11a Pass by directions.1/

N
1.12
1.12
1.68

Mean
N

NNE

NE

ENE

3.31

1.60
2.89
3.52
3.70

.67
2.28
4.50

-

1. 31
3

1

1/ In air miles.

2.92

2.48

4

3

ESE

SE

Direction
SSE
S

4.47

1.50

1.92

E

0

1

1

I

1.43
.94

SSW

SW

WSW

W

WNN

NW

NNW

.79
1.12
.63

1.49
2.10
4.94

5.77
2.l3
.59
1.71
7.76
6.14
2.38

2.25
1.56
2.86
1.00
1.00
.40
2.74

.86
.22
6.95
5.67

10.64
2.60
10.89
.60
10.01
5.40
6. 76
3.20
2.18
8.55
3.l3
4.05
5.27
8.46
3.21
3.84

1. 34
2.92

1.18
2

.85
3

I

w

~
~
I

2.84

3.78

1.69

3.43

5.55

2.13

3

7

7

4

16

2

�-315-

Only one bird (subadult male) banded at Guanella Pass, has been able to
become established on a territory in Mt. Evans (1972). Two additional males
shot and recovered at Mt. Evans in 1972 were located on summering sites and
were known to breed elsewhere. No females banded at Guanella were found to
move to the Mt. Evans area. Thus, mainly males moved east to Mt. Evans,
primarily to summering sites. Three birds banded at Mt. Evans (2 females
and 1 male) were recovered on breeding sites in the vicinity of Guanella
Pass. The adult male moved 4.94 air miles southwest from Mt. Evans to a
territory approximately 2.5 miles south-southeast from the top of Guanella
Pass. This was the longest movement recorded for a male as most males moved
less than 2 miles to a territory. It was concluded that minimal movement
occurred from Guanella Pass east to Mt. Evans, and west from Mt. Evans to
Guanella.
Adult males showed no significant movements in any given direction. Most
adult males occupied territories in the immediate vicinity surrounding
areas where they spent the winter. Unlike males, both adult and subadult
females exhibited significant movements to the)n01ljthwest. In addition,
significantly more females than males moved west,southwest and west. Of 14
birds moving in these directions, 11 were females. Also, more adult females
moved west-southwest than males or juvenile females. Since most adjacent
alpine areas lie to the west, more females were attracted to Guanella from
this area. Consequently, in spring when birds depart from the wintering areas
most females move in the westerly directions, primarily northwest, west and
west--southwest.
I

Movements from Wintering to Breeding or Summering Areas
Four hundred and ninety-seven different ptarmigan have been banded at Guanella
Pass since 1966. Locations were described in detail when birds were initially
banded and upon subsequent reobservation on breeding or summering areas.
During 1972-73, 50 individual movements were documented. In addition, 8
movements were recorded from band recoveries prior to the initiation of
intensive study. Movements were plotted on topographic maps (Figs. 3, 4, 5,
and 6) and distances (air miles) between observations were measured. Tables
7 and 8 summarize movements from wintering to breeding and summering areas,
respectively.
Upon examination of Table 7, three generalizations can be made: (1) females
of all age classes moved farther than males from wintering to breeding sites,
(2) subadult males moved greater distances than adult males, and (3) subadult females moved the greatest distances. Similar results were reported
by Braun (1969).
Observed differences in movements between adult and subadult males were not
signficant, but this may be due to the small sample size of males recovered
on breeding areas. Most subadult males do not become established on territories their first year, and therefore cannot be located with the tape
recorded call. With no prior attachment to breeding sites, subadult males
leave the wintering area in late April in search of places where they can
become established. Adult males are experienced and generally winter at the
lower edge of their breeding site. Thus, adult males moved short distances
(1. 34 air miles) to the territory occupied the previous year. However, on
the average, subadult males only moved .31 air miles further than adults to

�!
W

t-'

0\

e

Fig. 3, Movements of adult female white-tailed ptarmigan from the Guane11a Pass wintering
area (and Mt. Evans) to breeding or summering areas, pooled data from 1966-1972.
(Solid
line = movement to breeding site; dashed line = movement to summering site).

�I

W

I-'
......•
I

Fig. 4. Movements of subadult females from the Guanella Pass wintering area to breeding
or summering areas, pooled data from 1966~72. (Solid line = movement to breeding site;
dashed line = movement to summering site).

�~

=-

\ ..=-

-I::l

--:.-.;

-=::I:""

•...

-=

&lt;

I

....

W

00
I

Fig. 5. Movements of subadult males from the Guanella Pass wintering
or summering areas, pooled data from 1966-72.
(Solid line = movement
dashed line = movement to summering site).

area to breeding
to breeding site;

�u
W

I-'

1.0
I

Fig. 6. Movements of adult males from the Guane11a Pass wintering area (and Mt. Evans) to
breeding or summering areas, pooled data from 1966-72. (Solid line = movement to breeding
site; dashed line = movement to summering site).

�-320-

breeding sites. Since most males return to natal areas, larger sample
sizes may indicate no significant differences in movements of males of all
age classes.

Table 7. Movements of white-tailed ptarmigan from wintering to breeding
areas. 1./

Adu1ts

Females
Subadu1ts

Males
Adults

Subadults

1.00

1.50

1.00

2.89

1.71

1.60

.67

1.12

2.10

2.28

.60

.79

1.43

.87

.59

.40

1.92

1.49

.63

.22

3.52

10.01

4.94

4.50

1.68

1.12

2.18

3.31

8.55

5.40

1.12

6.76

'i

'.94,

1.34

3.20
2.92
7.76
10.64
10.89
N

10

15

8

6

Ave nage

2.52

4.72

1.34

1.65

1./ In air miles.

�-321-

Table 8.
summering

Movements of white-tailed
areas. 1:../

ptarmigan

from wintering

to

Males

Adults

Females
Subadults

Adults

Sub adults

6.95

2.38

3.70

3.13

4.05

3.84

6.14

2.74

4.47

5.27 ~/

8.46

2.60

3.212/

2.86

1.56

2.25

5.67

?J

2.13
5.77
1

N

8

6

Average

4.60

3.79

4

3.51

1/ In air miles.

~I Unsuccessful

females.

Departure from wintering areas in 1972 (spring of 1971-72) was abrupt, with
winter use sites being essentially devoid of ptarmigan after April 25. Adult
males, occupying territories .in the immediate vicinity of the wintering area,
probably moved directly to their breeding site while subadults wandered,
looking for available sites.
Female movements were probably gradual over a
10-15 day period, with short flights between areas. Similar results were
reported by Irving et ale (1967) for movements of willow ptarmigan in
Alaska.
Females undoubtedly c~ossed timbered areas during longer movements
to territories and upon their subsequent return to the wintering area.

Movements

To and On Wintering

Areas

Movements of ptarmigan to wintering areas are gradual and greatly influenced
by climatic conditions (Braun and Schmidt 1971). Birds generally appeared
on the wintering area by late October.
However, moderate weather conditions
during the fall of 1972 delayed the arrival of birds to Guanella Pass by
approximately two weeks.

�-322-

Both adult and sub adult females moved longer distances than males to
wintering areas.
Differences in wintering habitat preferred by males and
females were probably the major factors affecting movements.
Considerable data were available on the affinity of birds for wintering areas.
Braun (1969) noted that once males have successfully competed for breeding
sites, they return to these areas in following years.
Similarly, females
generally return to the same territory, but if the previous territorial male
dies, the female may seek a male on another territory.
It is also believed
that birds must establish themselves on wintering areas similar to their
establishment on breeding sites.
Once established, most birds return to the
same wintering area year after year.
Table 9 presents the analysis of birds returning to the wintering area.
With the exception of 1968-69, approximately 28% of the birds banded during
the previous winter returned to the same area in following winters.
Due to
small sample sizes, data for 1968-69 may not represent the true situation.
Seventy-two percent of the birds banded in most years were not observed
again, and are assumed not to have returned.
Undoubtedly this is an overestimate as some birds, which normally would have returned, died, and all
birds returning were not located.
Approximately 39% of the adult males banded returned, while only 18.5% of
the banded subadult males returned.
Together, about 24% of the banded males
returned.
Unlike males, more subadult (36.5 %) than adult (27.5%) females
returned to the wintering area, with approximately 31.5% of all banded
females returning.
It is evident then that at least 28% of the birds banded at Guanella are
migrating from and to the wintering area. The 72% not returning include:
(1) birds dying, which normally would have returned, (2) birds returning but
not located, (3) birds dying, which would not have returned, and (4) birds
dispersing from the wintering area and becoming established on other sites.
Undoubtedly, the figure for birds returning was underestimated.

of 52 birds

banded

at Guanella

and observed

on breeding or summering areas,
17 were known to return in
winters following initial banding, and 22 were.!not reobserved at Guanella.
With subtraction of known losses, a more accurate estimate (43.6%) of birds
returning is attained.
Forty-five percent of the females returned, while
only 30% of the males returned.
The figure fo~ males returning probably is
underestimated,
as wintering areas for males are not searched as thoroughly
as those where females winter at Guanella Pass.

13 were known to be removed from the population,

Significantly more adult than juvenile males returned to the wintering site
used the previous year.
Since territorial males generally winter at the
lower edge of their territories, only subadult males which become established
on territories in the vicinity of Guanella Pass should return the following
winter.
Those becoming established farther away probably winter at the lower
edge of their territories and do not return.
Most territorial males are
adults, consequently, adult males banded at Guanella probably occupy territories on the breeding areas immediately surrounding the Guanella Pass

�-323-

wintering area and return year after year.
It was concluded that once
established on territories, males of all age classes show a high fidelity
to these sites and adjacent wintering areas.
Since most territorial males
are adults, more adults are observed returning to the same wintering site
6f previous years.
No significant differences were found in the number of adult and subadult
females returning to Guanella Pass.
Females were found to return from
greater distances away (over 8 miles) than males.
Thus, Guanella Pass
attracts major numbers of ptarmigan, primarily females of all age classes,
from at least 8 miles away. This supports the hypothesis that wintering
sites for females may possibly be limited.
Once ptarmigan reached areas suitable for wintering, movements became
restricted.
There were no significant differences in movements between
sexes and age classes during the winter period.
Daily movements varied, but
generally, birds were more sedentary during the earlier months of winter,
with increasing movements in mid- to late winter.
Increased mobility was
also noted in mid- to late April.
It was believed that snow depths directly
affected ptarmigan movements on the wintering area. Increasing snowfall in
mid- to late winter covered much of the willow and birds increased their
daily movements to find available willow.
In addition, soft snow suitable
for night roosting was: often located in large snowfields or near forested
areas some distance from exposed willow.
Consequently, most birds were
believed to more 200 to 300 yards from roosting to feeding sites.
Upon
completion of feeding, birds usually flew to a day roost. Braun and Schmidt
(1971) noted feeding movements were often circular, with much of the same
area covered each day. Individuals were documented to move over 1 mile
between sightings on the wintering area. An adult female, observed on 5
consecutuve weekends, moved 1.2, .5, 1.4, and .75 miles from the location
observed the previous weekend.
Reobservations in mid- to late April indicated some birds banded east of the road moved to areas on the west side.
Reasons for the observed movements to the west side were not definitely known,
but appear related to snow conditions.
Areas suitable for roosting on the
west side may have been limited due to wind action until early spring.
As
weather moderated, in late winter, ptarmigan snow roost less, thus they
could utilize the relatively unused willow exposed as the snow settled west
of the road.

Distribution

on the Wintering

Area

Analysis of observation data revealed no apparent differences in distribution of ptarmigan on the wintering area between years.
However, differences in monthly distribution in winter were documented.
Essentially, all
sites on the wintering area were utilized, with some being highly preferred
to others.
During most months, about 90% of the birds utilizing the Guanella Pass
wintering area were east of the road. This was probably due to differences
in snow quality.
Snow conditions on the wintering area appeared to have a
pronounced influence upon the distribution of birds.
During January and
February, when most snow on exposed willow sites was crusted, birds were
most often observed near timbered areas where soft snow was present.

�Table 9.

White-tailed ptarmigan returning to the Guanella Pass wintering area.11

Winter

Females
Returning

Adult Females
Returning

Subadult Females
Returning

Males
Returning

Adult Males
Returning

1967-68

39.0

28.0

53.3

18.7

50.0

8.4

32.7

1968-69~/

46.7

50.0

40.0

66.0

66.0

0.0

50.0

1969-70

36.7

36.2

38.5

26.3

30.0

22.0

30.4

1970-71

23.6

20.0

28.0

30.0

40.0

26.6

25.0

1971-72

28.7

26.4

27.0

21.7

33.0

17.6

25.5

Subadult Males
Returning

Total Birds
Returning

I

Ave rage-31

32.0

27.7

36.7

24.2

38.0

18.6

II Percent of birds banded previous winter.

21 Due to small sample size (only 18 birds banded) this may not reflect the true situation.

11 Excluding 1968-69.

28.4

w
N
~
I

�-325-

Birds could normally be found along the east side of a low ridge running
southeast from the top. of Guanella Pass.
Significantly more birds were
found along this ridge than any other site on the wintering area. Currently,
no reason can be postulated for this occurrence, but perhaps a combination
of abundant food supply, suitable snow conditions, and protection from the
prevailing westerly winds attracts birds to this site.
Increased hUman activity at Guanella in recent years has appeared to influence ptarmigan distribution.
In earlier years(1966-67, 1967-68, and 196869) birds were often located within 100 yards of the road. Possibly due to
increased traffic and human activity, ptarmigan appeared reluctant to utilize
willow areas near the road in recent years.
Buds and twigs of willow provide the bulk of the food eaten by ptarmigan
during the winter (May 1970).
Thus, ptarmigan were seldom found where
willow was absent.
Sites used early by ptarmigan were devoid of birds later
in the winter, primarily because increasing snow depths covered the willow
in these areas. Thus, there was a change in distribution with increasing
snow depth, birds gradually moved from the outer boundaries of the study area
where willow bushes were generally short and sparsely distributed to the more
central portions where tall dense stands of willow were never completely
snow covered.
In mid- to late April birds were again observed on the outer
boundaries of the wintering area. At this time, birds were preparing to move
to breeding sites.
It was concluded that weather was the primary factor
determining distribution of ptarmigan in winter because it influenced availability of food and location of roosting sites.

LITERATURE
Aldrich, J. W. 1963. Geographic
Wildl. Mgmt. 27(4) :529-545.
Bailey, A. M. 1927.
44(2) :184-205.

CITED

orientation

of American

Notes on the birds of southeastern

tetraonidae.

Alaska.

J.

Auk

Bendell, J. F., and P. W. Elliott.
1967. Behavior and regulation of
numbers of blue grouse.
Canadian Wildl. Servo Rept. Series No.4.
Bent, A. C. 1932. Life histories of North American
U. S. Natl. Museum Bull. 162. 490 p.

gallinaceous

76p.

birds.

Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado.
Ph. D. Dissertation.
Colo. State Univ.
Ft. Collins.
189 p.
1971. Habitat requirements of Colorado white-tailed ptarmigan.
Proc. West. Assoc. State Game and Fish Comm., 51:284-292.
1973. Continued inventory of selected
Div. Wildl. Fed. Aid Rept. In Prep.

ptarmigan

populations.

Colo.

�-326-

Braun, C. E., and G. E. Rogers. 1971. The white-tailed ptarmigan in Colorado. Colo. Div. of Game, Fish and Parks Tech. Publ. No. 27. 80 p.

--- , and R. K. Schmidt, Jr.

1971. Effects of snow and wind on wintering
populations of white-tailed ptarmigan. Pp. 238-250. In Haugen, A. 0.,
(ed.), Proc. Snow and Ice Symposium. Iowa State Univ., Ames. 280 p.

____ , and G. E. Rogers. 1973. Census of Colorado white-tailed
ptarmigan with tape recorded calls. J ..Wildl. Mgmt. 37(1) :90-93.
Choate, T. S. 1960. Observations on the reproductive activities of whitetailed ptarmigan (Lagopus leucurus) in Glacier Park, Montana. M. A.
Thesis. Univ. MOntana, Missoula. 113 p.
1963. Ecology and population dynamics of white-tailed ptarmigan
(Lagopus leucurus) in Glacier National Park, Montana. Ph. D. Thesis.
Univ. MOntana, Missoula. 205 p ,
Gabrielson, I. N., and F. C. Lincoln.
Co., Harrisburg, Pa. 922 p.

1959.

The birds of Alaska.

Stackpole

Gullion, G. W., and W. H. Marshall. 1968. Survival of ruffed grouse in a
boreal forest. Living Bird 7:117-167.
Haskins, A. G. 1969. Endoparasites of white-tailed ptarmigan (Lagopus
leucurus) from Colorado. M. S. Thesis. Colorado State Univ., Ft.
Collins. 147 p ,
Irving, L., G. C. West, L. J. Peyton, and S. Paneak. 1967. Migration of
willow ptarmigan in arctic Alaska. Arttic 20(2) :77-85.
May, T. A. 1970. Seasonal foods of white-tailed ptarmigan in Colorado.
M. S. Thesis. Colo. State Univ., Ft. Collins. 55p.
1973. Bioenergetics of white-tailed ptarmigan.
Univ. of Colorado, Boulder. In Prep.

Ph. D. Thesis.

Packard, F. M. 1945. The birds of Rocky Mountain National Park, Colorado.
Auk 62(3):371-394.
Racey, K.

1948.

Birds of the Alta Lake region, B. C.

Auk 65(4) :383-401.

Robel, R. J., J. N. Briggs, J. J. Cebula, N. J. Silvy, C. E. Viers, and
P. G. Watt. 1970. Greater prairie chicken range, movements and habitat
usage in Kansas. J. Wildl. Mgmt. 34(2):286-306.
Schmidt, R. K., Jr. 1969. Behavior of white-tailed ptarmigan in Colorado.
M. S. Thesis. Colo. State Univ., Ft. Collins. 174 p.
Weeden, R. B. 1959. The ecology and distribution of ptarmigan in western
North America. Ph. D. Dissertation. Univ. British Columbia,
Vancouver. 247 p.

�-327-

Weeden, R. B. 1964.
migan in winter.

Spatial separation of sexes in rock and willow
Auk 81(4):534-541.

ptar-

1957.

Seasonal and geographic variation in the foods of adult
white-tailed ptarmigan.
Condor 69(3):303-309.
F., and J. F. Bendell.
1967.
J. Wildl. Mgmt. 31(1) :202-204.

Zwickel,

___

A snare for capturing blue grouse.

, 1. O. Buss, and J. H. Brigham.
1968. Autunm movements
grouse and their relevance to populations and management.
Mgmt.
32(3) :456-468.

Prepared

by

~~

Richard W. Ho f fman

of blue
J. Wildl.

��April,

1973

-329-

JOB FINAL REPORT

Sta te of __
Project
Work

---.:.C,;:..O.::;L,;:..ORAD=::..O::.._

Plan No.

15

Job Title:
April

3

Job No.
Experimental

Period Covered:
Personnel:

Game Bird Survey

W-37-R-26

No.

Breeding

l~ 1972 through March

of Mountain

Quail

31, 1973

Thomas A. Barber, John F. Corey~ Larry R. Crooks, Carroll J. Grand Pre,
George Harrington, Willis G. Mansfield, Robert L. schmidt, Lawrence A.
Webster, Harold M. Swope) Ann Leckler and Donald M. Hoffman.

~')STRACT
During the period 1967 through 1972 when efforts were made to propagate mountain
quail in captivity at the Fort Collins Wildlife Research Station, fair to good
egg production occurred in 1967, 1969 and 1971 and no or very poor egg production in the alternate years of 1968, 1970 and 1972. Possible reasons include:
(1) an inadequate number of birds were held at the start of the propagation effort,
resulting in significant gains in inbreeding per generation,
(2) adequate breeding
pen facilities were not available until 1970, and (3) the species was found to
stress easily from a variety of causes.
Fertility averaged 63.14 percent for the
7 year period and the hatchability of fertile eggs set was 82.53 percent for the
period.
Survival of young and adults was considered fair to good. Mountain
quail are considered to be one of the more difficult game birds to propagate in
captivity due primarily to their tempermental nature in egg production.
The
study was terminated in March, 1973 and all remaining pen raised stock was released due to the erratic production secured with the species.

�-330-

RECOMMENDATIONS

Any future plans for releasing mountain quail in Colorado should be based
upon an evaluation of the success or failure of past field releases.
Primary
considerations prior to contemplating future exotic game bird propagation
efforts should be advance planning for adequate facilities and adequate
numbers of breeders.

�-331-

EXPERIMENTAL

BREEDING

OF MOUNTAIN

QUAIL

IX:&gt;naldM. Hoffman

Initial releases of mountain quail in Colorado were made from wild trapped
stock from California and Oregon received through wildlife trade agreements.
It became apparent early in the study, if continuing releases of mountain
quail to selected areas of Colorado were to be made an attempt should be
made to propagate the species in captivity.: Only 16 mountain quail from'
159 received initially from California in 1965 were held for experimental
propagation studies.
Twelve wild trapped birds from Oregon were added to
the penned

stock in 1970.

This study was begun in 1967 in order to, (1) propagate mountain quail in
numbers suitable for field releases, and (2) to study techniques in reproducing the species.
The study was conducted by Harold M. Swope during 1967
and 1968, and by Donald M. Hoffman from 1969 through 1972.

P. S. OBJECTIVE
To develop

game farm production

techniques

SEGMENT
Segment

objectives

varied

for mountain

quail.

OBJECTIVES

by year as follows:

1967
To determine the effects of a special high protein diet (with supplemental
calcium) on fertile egg production.
To determine the effects of artificial
light stimu1ighting on fertile egg production.
1968
To measure

the effects

28 percent)

on fertile egg production.

of two levels

of protein

in the feed (22 percent

vs.

1969
To measure
fertile

the relationship

of flock mating

and selected

groups mating

to

egg production.

1970
To measure the relationship of pair mating,
mating to fertile egg production.

selected

group mating

and flock

1971
To measure the relationship of pair mating versus group matings
and 1 cock, and 3 hens and 1 cock to fertile egg production.

of 2 hens

�-332-

1972
To compare numbers of survavang mountain quail with numbers released in
holding pens (rearing and conditioning runs) containing various bird
densities. To compare earliness of egg production and numbers of fertile
eggs produced by breeders receiving 2 extra hours of lighting in the mornings
during each 24 hour period, with breeders receiving only normal daylight. To
compare fertile egg production and hatchability of eggs according to the
following ages of breeders: (a) adults (1969 hatched or older), (b) first
hatch young from 1971, and (c) second hatch young from 1971.

METHODS AND MATE RIALS
From 1967 through 1969 attempts were made to propagate mountain quail even
though very inadequate facilities were available at the Fort Collins Wildlife
Research Station, due to intensive efforts to propagate 2 other upland game
birds (Hungarian partridge and pale spotted tinamou). Because ground floored
breeding pens were unavailable, wire floored pens had to be utilized. These
proved unsuitable because the mountain quail breeders soon became sore footed
and stressed, on the wire floors. A new set of 10 mountain quail breeding
pens was completed on April 13, 1970 (Fig. 1), and a similar set of 10 breeding
pens formerly used for tinamou breeding pens (Figs. 2 and 3) was used in
1972 after tinamou propagation was terminated. Thus, adequate facilities
suitable for experimental mountain quail propagation were aVailable only from
1970 through 1972. Methods and materials used by years are discussed below.
1967 (Swope 1968)
Four hens and one cock were placed in pen number 1 and fed a hen lay feed
containing 20 percent protein. The remaining three hens and one cock were
fed a 22 percent protein game bird feed in pen number 2. Birds were not available to test the effects of stimulighting. The fact that only 9 mountain quail
(7 hens and 2 cocks) were available for the 1967 tests indicates that an inadequate number of these birds were held for propagation purposes at the start
of the study. The gain in inbreeding per year with the number of breeders in
a closed population can be calculated by the formula (MUller, unpublished).
F(gain)
F(gain)
~

=

1
8(M)

+

1
8(F)

1
=

8(2)

+

1
8(7)

= 8 percent gain in inbreeding per generation

(Hoffman 1969)

Forty mountain quail breeders were on hand for 1968 tests. Mountain quail
breeders were placed in breeding pens 1 and 3, and fed the standard game bird
breeder feed containing 22 percent protein. Birds in pen 2 were fed a similar
feed, but with a protein content of 28 percent. Plans were to mark all eggs
so that fertile egg production could be determined by protein level in the feed,
but no eggs were produced.

�-333-

N

30l----"------~

fiE---------

rE----12'---7I

iE----

12 _~-)I
I

o

-

\

SCALE 8

- L,---_-I-l-:--_---li .
-0'
51
10'
Fig- L.

OO~(lilQ" ArrtlnOQmont

of

Moulltoin

Quail 6rctedino

Pens ,South

section,

�-334-

~----------------30'----------------~
IL

7i"

;

I'

,

12

=»

IL

4J

-0)

I

1

-

0

"1"

(0

/

-~

v

ro

-

~

-

-

CX&gt;

N

~

/

-,

-

en

-

){

N

J

12

i;

SCALE 8

L_._I
0'

5'

-----1
10'

Fig_ 2•.
Detail.w

Arrangement of Mountain Quail Breeding Pens t North Section

�-335-

Fig. 3. A set of 10 breeding pens (north section) used for experimental
mountain quail breeding studies in 1972. (D. Hoffman, photo).

�-336-

1969 (Hoffman 1970)
Wire floored pens number Ql and Q3 were divided into 2 approximately equal
size pens. each with a wood framework and 1 inch chicken wire.
Selected
groups of 1967 hatched birds or earlier hatched birds, consisting of 2 hens
and 1 cock were placed in each of these 4 pens of equal size and a flock of
8 hens and 4 cocks were placed in the larger undivided pen Q2 on March 24,
1969.
On April 2,' 1969, after only 10 days confinement on wire, 10 of the
12 mountain quail in pen Q2 had extremely sore and bleeding feet. Several of
these birds on wire had started to develop
a white diarrhea, apparently due
to stress conditions.
Some of the birds in the selected groups of 2 hens and
1 cock each (4 pens) had also started to exhibit these symptoms of stress,
although not as many as in the more crowded community breeding pen. All birds
were then moved to the ground floored quail runs shown in Fig. 4. Pens I, 3,
and 5 contained selected groups of 2 hens and 1 cock each, and pen 6 contained
a community of 6 hens and 3 cocks.
The surplus breeders (5 sore footed hens,
plus 1 cock) were held in pen 4, and 1 hen plus 3 cocks were held in pen 2.
1970

(Hoffman

1971)

A new set of 10 mountain quail breeding pens was completed on April 13, 1970
(Fig. 1). It was planned to have these completed by mid-March, but inclement
weather plus a manpower shortage delayed the completion.
Pairs of mountain quail were placed in pen numbers 1, 5, 7, and 9; trios (2
hens and 1 cock) in pen numbers 2, 4, 6, and 10; and a community pen of 8 hens
and 4 cocks was placed in the combined pen numbers 3, 8, and alleyway, on
April 14, 1970.
All birds were fed a standard 22 percent protein level game
bird breeder feed and all were 1969 hatched birds.
Numbers of fertile eggs
produced were to be recorded by individual pens.
Ten wild trapped mountain quail of mixed sexes received from Oregon in Segment
23 were placed in combined holding pen numbers 1 and 2, eleven 1969 hatched
birds including 2 hens and 9 cocks not needed to fill the experimental breeding
pens were left in combined holding pen numbers 3 and 4, and seventeen 1967
hatched or older breeders of mixeti sexes were placed in combined holding pen
numbers 5 and 6.
1971

(Hoffman

1972)

Pairs (1 hen and 1 cock) were placed in pens 1, 5, 7, and 9 (Fig. 1); trios
(2 hens and 1 cock) were placed in pens 2, 4, 6, and 10; and quads (3 hens and
1 cock) were placed in pens 3 and 8 OIl February 9, 1971.
All breeders were fed a standard 22 percent protein level game bird breeder
feed from March 6, 1971 until removed from the breeder pens on July 23, 1971.
Primaries and secondaries of both wings were clipped on all breeders in pens
4, 5. 8, 9, and 10. All birds were 1969 hatched or older, and all but 4 were
pen raised at the Fort Collins Wildlife Research Station.
The 4 were wild
trapped mountain quail received from Oregon in January 1970.

�-337-

!(

20'

i

1E-8'~ r 8'--) E-~

\

N

i
I

I\.

I

3

15
\

,

\
:

2

4

6
.'

,

o
ll----------

Fig"

60'

4,,,

Moui'l~oln

Ouoil

HoldinCJ

Pens,

EAST

SECTION

15

30

�-338-

All pens were fed and watered only twice a week and eggs were collected at
these times to reduce the disturbance factor.
Eggs laid were marked with
the date collected and pen number to facilitate determination of total eggs
laid, fertile eggs laid, and numbers of chicks hatched by pens.
Adequate
cover in the form of growing clumps of tall wheat grass were planted in April
1970.
Perches were installed in the spring of 1971 on the advice of a successful game bird breeder in California.
Although not specifically written as planned tests for 1971, records of
survival of chicks for the first 7 and 30 day periods and for overwinter survival in holding pens were maintained.
Numbers of day old mountain quail
placed in the brooder houses after removal from the hatcher were recorded and
losses experienced were marked on brooder house cards with causes of death
when this could be determined.
Young removed were also recorded and percentage
survival calculated.
Birds counted into holding pens in either July 1971 (adult breeders), or in
September 1971 (1971 hatched young) were recorded.
Birds recovered in March
and April 1972 were also recorded and survival rates calculated.
1972 (Hoffman 1973)
A set of 10 breeding pens (south section), built specifically for mountain
quail propagation (Fig. 1), plus a set of similar breeding pens (north section)
used previously for tinamou propagation (Figs. 2 and 3) were used for both
extra lighting and egg production by age of breeder tests. Breeding pen
numbe rs 11 through 20 (north section) were wired for lights, and an automatic
timer installed during the winter period, prior to sorting the mountain quail
breeders.
Mountain quail of various ages were selected" and trios (2 hens and
1 cock) placed in each breeding pen on February 7, 1972. Trios were also
placed in 2 McCarty brooder units placed in one of the mountain quail holding
pens (number 8). All breeders in all of the 22 pens used as breeding pens
were fed a standard 22 percent protein game bird breeder feed in 1972.
Table
1 lists the pen schedule used for mountain quail in 1972.
All pens were fed
and watered twice a week and eggs were collected at these times, similar to
procedures used in 1971.
Eggs laid were marked with the date collected and
pen number to facilitate determination of total eggs laid, fertile eggs laid,
and numbers of chicks hatched by pens.
The center portion of the mountain quail holding pens (east section) (pens 3
and 4) shown in Fig. 4, and the 3 holding pens (west section) (pens 7, 8, and
9) shown in Fig. 5 were used for overwinter survival tests. Known numbers of
mountain quail of the different age groups were placed in these holding pens
in July 1972, when collected from breeding pens (1969 hatched or older adults)
or moved from brooding pens (1972 hatched young).
Numbers recovered on March
19, 1973 when banding and crating for field release were determined and overwinter survival calculated.

�-339-

Table 1-

Pen schedule for mountain quail, 1972. 1/
Hatch
Number

Artificial Light
None
A.M.

Year Hatched

1

T

1969 or older

2

T

1971

1

3

T

1971

2

4

T

1969 or older

5

T

1971

1

6

T

1971

2

7

T

1969 or older

8

T

1971

1

9

T

1971

2

10

x

1971

1 (not on test)

Pen
Number

11

T

1969 or older

12

T

1971

1

13

T

1971

2

14

T

1969 or older

15

T

1971

1

16

T

1971

2

17

T

1969 or older

18

T

1971

1

19

T

1971

2

20

x

1971

2 (not on test)

McCarty (8)

1971

2 (H) and 1 (C)

MCarty (N)

1971

2

Number Pens
on Test

9

9

l/ All pens,were fed a standard 22 percent protein game bird breeder feed.
Each pen contained 2 hens and 1 cock (trio).

�-.340-

r----..--:../
'
• ---.----~--10

I(- 4-lI

9

8

N

7

-.N
,...

SCALE

'7----'----.

I{------_

fiG.

5"

30'

0'

5'

10'

�-341-

RESULTS AND DISCUSSION
Results obtained

by years include:

1967 (Swope 1968)
Only nine mountain quail were on hand for research to develop production
techniques.
Four hens fed a 20 percent protein hen lay feed laid 19 eggs.
Three other hens fed. a game bird feed containing 22.percent protein laid
146 eggs.
Sixty-five young quail were hatched from the 165 eggs.
Of the
161 eggs set (four culls) 98 were fertile.
1968 (Hoffman 1969)
The 40 breeders did not produce any eggs.
Station personnel feel that this
may have been due to the poor condition of birds from confinement in wirefloored pens during a relatively severe winter.
Although the breeders were
wintered in these same wire-floored pens successfully the previous winter,
the winter of 1967-68 was more severe.
This resulted in many mountain quail
having sore feet and a general unthrifty condition.
During early 1968, all of the mountain quail were medicated with "Piperazine"
in the drinking water after a few Trichamonas spp. organisms were found in
the digestive tract of a bird which had died. This probably induced additional stress in these birds just prior to the normal laying season.
In
addition, all of the breeders were penned in3 wire-floored community mating
pens during 1968, in contrast to breeding groups of 3 hens with 1 cock or 4
hens with 1 cock in 1967.
1969 (Hoffman 1970)
Good progress was made in the experimental game farm propagation of mountain
quail in 1969, with 100 eggs being laid by 28 breeders of mixed sexes (18
hens and 10 cocks) and 68 young hatched, compared with no egg production in
1968. Of the 68 young hatched in 1969. there are 45 healthy birds to be used
in the 1970 tests, plus 23 older breeders at the Fort Collins Wildlife Research
Station.
In addition, twelve wild trapped mountain quail from Oregon were
added to this breeding stock on January 18, 1970. Ten of these wild trapped
birds survived until the breeding season.
A test to compare fertile egg production by three selected groups of two hens
and one cock each, with a community pen of six hens and three cocks cannot be
considered a valid test, due to stresses induced on the breeders by holding
them in wire-floored pens because of inadequate ground floored facilities.
1970

(Hoffman 1971)

Although procedures for the 1970 mountain quail tests were carried
planned, except for being approximately 1 month late in completing
experimental breeding pens, only 1 egg was laid. This was laid on
1970 in the community pen (combined pen Numbers 3, 8 and alleyway)
infertile.

out as
a set of 10
May 1,
and was

�-342-

It was found in previous years tests that mountain quail stress easily.
Catching and sorting the breeders at approximately the time when they should
have started to lay may have been the reason for almost no egg production in
1970. No eggs were laid by t.enwild trapped mountain quail of mixed sexes
received from Oregon in Segment 23, eleven surplus 1969 hatched birds (2 hens
and 9 cocks), or seventeen 1967 hatched or older breeders of mixed sexes
held in separate mountain quail holding pens in 1970. It was necessary to
catch and move these surplus breeders in order to select breeders used in
the experimental breeding pens and to utilize holding pens with the best cover.

1971 (Hoffman 1972)
Good success was experienced with the experimental propagation of mountain
quail in 1971, with 344 eggs set (culls excluded), of which 209 were fertile.
A total of 176 chicks were hatched from these eggs. A total of 28 breeders
(18 hens and 10 cocks) were used in four pairs (1 hen and 1 cock), four trios
(2 hens and 1 cock), and two quads (3 hens and 1 cock) in ten separate pens
on test in 1971. The sorting of breeders in early February, 1971 and disturbing the breeders as little as possible appeared to be of considerable
value in securing egg production.
In a test to compare egg production by various sex ratios of breeders, numbers
of fertile eggs laid per hen was highest with the trios (12.50), followed by
the quads (11.00), and pairs (10.75). The best hatching of fertile eggs
occurred with the trios (92.00 percent) compared with 83.72 percent for the
pairs, and 72.72 percent for the quads.
Numbers of chicks hatched was also
highest for the trios.
The clipping of the flight feathers on both wings of
breeders in half of the pens calmed these easily stressed birds conSiderably,
but also resulted in 20.34 percent fewer fertile eggs being produced (94 for
the five pens with clipped flight feathers, compared to 118 for the five pens
with normal wings).
Excellent survival was experienced with the first three hatches for the first
seven days (average 94.80 percent) and good survival was experienced for the
first 30 days (average 83.24 percent) during 1971. No birds were hatched in
a fourth setting of 13 eggs.
Overwinter survival in pens with various bird
densities and ages ranged from 86.67 percent in a holding pen with 26.40
square ft of ground space per bird containing 60 1971 hatched quail to 100.00
percent in a holding pen with 144.00 square ft of ground space per bird
containing 11 1971 hatched quail.
The average overwinter survival was 91.62
percent for the five groups of mountain quail on test.

1972
Procedures used in 1972 were similar, for the most part, to those used in
1971 when good egg production was secured from the mountain quail breeders.
These included selecting and sorting breeders in early February, disturbing
the breeders as little as pOSSible, and providing feed and water twice per
week during the egg laying period.' Eggs were gathered when feeding and
watering the birds.
The provision of 2 hours of extra lighting in the early
mornings was started on March 1, 1972.

�-343-

Numbers of eggs produced in 1972 are listed by pens in Table 2. Total egg
production (culls excluded) from the 66 mountain quail breeders (44 hens and
22 cocks) was only 83 eggs, compared with 368 total eggs produced by 28
breeders (18 hens and 10 cocks) in 1971. Thus, 1972 proved consistent in the
trend of securing fair to good egg production from the mountain quail only
in alternate years since 1967.
Breeders (Fig. 6) in 5 pens, including 2 with no extra lighting (numbers 2 and
5). and 3 with extra lighting in the early morning hours (numbers 11, 17 and
20), began producing eggs in early April 1972, and hopes were high that egg
pro duct Lon would continue.
During April 1972, 26 eggs (includes 3 culls) were
laid by breeders in the 5 pens listed above, but by the second week of May
only 1 pen (number 17) was still in production.
Hens in pen number 17 continued to produce eggs until July 10, 1972, but all eggs laid after June 3,
1972 (25 eggs) were infertile.
Reasons why egg production beg&amp;L in 5 breeding
pens and then all but 1 quit early is not known.
Weather factors are not a
likely explanation, since Hungarian partridge breeders at the Station experienced a peak egg production year in 1972. At least 3 factors were different
.in the spring of 1972 compared to 1971, including (1) 3 stray dogs were
observed intermittently and tracks indicated these frequented the breeding
pen area, (2) colonies of Norway rats moved into the breeding pens during the
spring, and (3) vehicular traffic
on Interstate Highway 25 (approximately
100 yards west of the breeding pens) appears to be increasing with time,
although no traffic counts have been made.
It is possible that the mountain
quail breeders were stressed during the early egg production period by 1 or
more of these factors or possibly some unknown factor.
A check on breeder
feed composition ruled out a change of ingredients in the 22 percent protein
game bird breeder feed as a factor.
Earliness

of Egg Production

and Egg Production

by Ages of Breeders

Because egg production began in 5 pens (numbers 2,5,11,17
and 20) but
continued in only 1 pen (number 17), planned tests in 1972 to compare earliness of egg production with extra lighting in the early morning hours, and
compare egg production by various ages of breeders were invalid.
The fact
that limited egg production was secured from both first hatch young from 1971
and second hatch young from 1971 indicates that young of the previous year
are capable of producing eggs.
This was not previously known.
Table 2 lists a comparison of eggs produced in various pens in 1972 and Table
3 lists information relating to the mountain quail hatches in 1972. Although
3 separate settings of eggs were made in 1972, ,all 19 eggs set in the third
setting were infertile, as they were late eggs from pen number 17. An average
of 97.92 percent of all fertile eggs hatched, showing the high hatchability
of fertile mountain quail eggs.
Table 4 lists information relating to survival of mountain quail young for
1972. An average survival rate of 82.98 percent is indicated for the first
7 days and 78.72 percent for the first 21 days.

�Table 2.

A comparison of eggs produced and hatched by mountain quail in various pens, 1972. 11

Artificial
Lights

Number
Good
Eggs
Laid

Hen

Number
Fertile
Eggs

1971-1

None

8

4.00

8

8

100.00

100.00

5

1971-1

None

1

0.50

1

1

100.00

11

1969 or
older

A.M.

1

0.50

0

0

17

1969 or
older

A.M.

69

34.50

39

38

Pen 2 Year and
Number-I Hatch No.

2

No.1

Percent
Number
Hatch
Hatched All Eggs

Percent
Hatch
Number
Fertiles Pipped

Number
Dead
Germs

Number
Culls

0

0

0

100.00

0

0

0

00.00

--

0

0

0

55.07

97.44

0

1

5

I

w
.p.
.p.
I

20

1972-2

.

A.M.

Totals
Averages

11 Sex ratio of all pens was 2H:1C.

4

2.00

83
8.30

0

0

48

47

00.00

56.63

--

0

o

1

0

1

6

97.92

All birds were fed a standard 22 percent protein game bird breeder feed
and none of the wing primaries were clipped. Only pens which produced eggs are listed (17 produced no eggs).

'l:..1 See Figs. 1 and 2.

�-345-

Fig. 6.

An adult mountain quail.

(D. Hoffman, photo).

�Table

Hatch
Number

3.

Mountafn quail

Date and
Time Set

Number
Set

-_._-----_. -.---.-1

2

3

hatches,

5-18-72
7 :00 P. M.

39

6-9-72
9:00
P.M.

19

7-13-72
11:59 P.M.

19

1972.

Date and
Time to
Brooder

------Hatched
Good Chicks
Cripples

DG's

Not Hatched
Inf.
Pipped

Total

Percent
Eggs

Hatched
Fertile

6-13-72
1:00 P.M.

34

0

1

4

0

87.17

97.14

7-5-72
2:00 P.M.

13

0

0

6

0

68.42

100.00

8-8-72
Hatch
Terminated
6:00 P.M.

0

0

0

19

0

00.00

Eggs

I

w
.r:-eI

Totals

77

47

0

1

29

0

Averages
61.04

97.92

�-347-

Table 4.

Survival of young mountain quail, 1972.

Hatch
Number

Date

Number to
Brooder

1

6-13-72

34

28

27

82.35

79.41

2

7-5-72

13

11

10

84.62

76.92

3

8-8-72

0

39

37
82.98

78.72

Totals

Number Survivin~ at End
21 Days
7 days

47

Percent Survival
21 Days
7 Days

Averages

Overwinter Survival by Various Densities in Holding Pens
Table 5 lists information relating to the survival of mountain quail in
holding pens (rearing and conditioning runs) during the period July 1972
through March 1973. An overall survival of 89.11 percent was determined. The
small number of mountain quail (101) in 4 different age groups limited the
effectiveness of this test. None of the density tests in 1972 (highest was
1 bird per 21.06 square ft of ground space) exceeded those tried in 1971 when
excellent survival was demonstrated with densities as high as 1 bird per 20.57
square feet of ground space.
Summary of Mountain Quail Propagation 1967-1972
Segment objectives were seldom attained due to the erratic nature of the
species in producing eggs for reasons which include:
1.

An inadequate nUmber of birds were held at the start of the propagation
e f fort;, resulting in significant gains in inbreeding per generation. A
few (10) wild trapped mountain quail from Oregon were added in 1970.

2.

Adequate breeding pen facilities were not available until 1970.

3.

The species was found to stress easily from a variety of causes. An
early effect of stress in game birds is reduced or termination of egg
production. Possible stress factors recorded during this study include:
(a)

Abrupt changes in diet.

(b)

Poor condition of birds following severe winter period.

(c) Working of breeders just prior to egg laying period.
(d) Holding birds in wire floored pens.

�-348-

(e)

Sorting

and handling

birds just prior to egg laying period.

(f)

Presence

of stray dogs in vicinity

(g)

Infestations

(h)

Noise from vehicles

of Norway

of breeding

pens.

rats in pens.

on a nearby

Interstate

Highway.

Table 6 summarizes mountain quail propagation at the Fort Collins Wildlife
Research Station from 1967 through 1972.
Fair to goo d :egg production was
secured in the alternate years of 1967, 1969, and 1971, and no or very poor
egg production in 1968, 1970 and 1972. This demonstrates an irregularity in
egg production which cannot be fully explained, although the species is known
to stress easily.
An average of 63.14 percent of all mountain quail eggs
were fertile and 82.53 percent of the fertile eggs hatched during the period
1967 through 1972.
Fig. 7 shows a group of 12 day old mountain quail leaving
a brood house at the Fort Collins Wildlife Research Station.
A search of available literature failed to reveal whether years of no, or
very poor reproduction occurs in wild populations of mountain quail.
In this
study, inbreeding could have been a factor, since a gain of 8 percent per
generation would be expected in a closed flock when only 7 hens and 2 cocks
were used initially.

Sex Ratios in Breeding

Pens

In 1967, 7 mountain quail hens laid 165 eggs in 2 community pens.
One pen
had 4 hens and 1 cock, and produced a total of 19 eggs and the other pen
contained 3 hens and 1 cock and produced a total of 146 eggs
During 1971, the only year when comparative tests were made, egg production
was compared among breeders in 4 pens, each containing 1 hen and 1 cock
(pairs), 4 pens, each containing 2 hens and 1 cock (trios), and 2 pens, each
containing 3 hens and 1 cock (quads).
Numbers of fertile eggs laid per hen
was highest with the trios (12.50), followed by the quads (11.00), and pairs
(10.75).
Based upon these results, all breeders used in 1972 tests were
housed as trios and trios are recommended in propagating the species.
Clipping

of Primaries

of Breeders

In 1971, clipping of primary flight feathers on both wings on half of the
breeders calmed these easily stressed birds conSiderably, but 20.34 percent
fewer fertile eggs were produced (94 fertile eggs were produced by breeders
in 5 pens with clipped wing feathers, compared to 118 fertile eggs produced
by breeders in 5 pens with normal wings).
Based upon these results, recommendations are not to clip flight feathers.
Incubation

Procedures

An automatic, forced air incubator was used for hatching mountain quail eggs.
Standard incubation temperatures (99.750 F + .250) and incubation humidites
of approximately 55 percent relative humidity (850 F wet bulb) worked well for
hatching mountain quail eggs.
The average time required to incubate mountain
quail eggs was 24 days + 18 hours, and the young chicks were normally held in
the hatcher for an additional 24 hours prior to moving to the brooderhouse.

�Table 5. Survival of mountain quail
sizes of pens, 1972-73.

during

summer through w Lnt.er period

Pen and
Hatch Numbers 1/

Size
(Ft. )

Total
Sq. Ft.

Date Birds
Released in
Pen

3 &amp; 4, 1972-1

20X76+
8X8

1,584

7, 1972-2

10X72

8, 1971 adults

by various

densities

of birds

and

No. of
Birds

Sq. Ft. /
Bird

Date Birds
Recovered

No. of Birds
Recovered

Percent
Survival

7-4-72

27

58.67

3-19-73

24

88.89

720

7-26-72

10

72.00

3-19-73

7

70.00

10X72+
6X45

990

7-13-72

47

21.06

3-19-73

45

95.74

10X72

720

I

w
.j:-o
\0

9, 1969 or older,
adults

17

101

Totals

42.35

3-19-73

14

82.35

90
89.11

Averages

1/ See Figs.

7-13-72

4 and 5.

I

�Table 6.

Summary of mountain quail propagation, 1967-1972.

Year

Number
Breeder
Hens

1967

7

1968

21 1/

1969

18

1970
1971
1972

Totals
Averages

~

Total
Eggs Ave. No.
Laid Eggs/Hen

165

23.57

0

0.00

100

5.56

No. No. Eggs
Culls
Set

No.
Inf.

No.
Percent
Fert. Fertile

Number
Chicks
Hatched

Percent Hatch
Total Fertile
Eggs
Eggs

63

98

60.87

65

40.37

66.33

0

100

20

80

80.00

68

68.00

85.00

1

11

84.21

8

25

97.92

0

1

0

9

37

3

0.05

0

1

1

0

0.00

0

0.00

18

368

20.44

24

344

135

209

60.76

176

51.16

126
21

1.98

717
120

6

34
5.71

No.
Crippled

161

1

83

No.
DC's

4

20

42

No.
Pips

83

35

48

689

254

435

I

3

(.oJ

VI

0

57.83

47

56.63

356
63.14

52.10

A sex ratio of 1H:1C assumed in 1968 since no record was made of number of hens.

82.53

I

�-351-

Fig. 7. A group of 12 day old mountain
Research Station.
(D. Hoffman, photo).

quail at the Fort Collins Wildlife

�-352-

Brooding
A modified Jamesway electric heat ring brooding unit with an attached fan
worked better than heat lamps for brooding mountain quail chicks.
Temperatures in the brooderhouse were maintained at 950 F for the first week and
gradually lowered thereafter in accord&amp;~ce with usual game bird rearing
procedures.

Feeding
Based upon very limited data gathered in 1967 (Swope 1968) mountain
breeders should be fed a standard game bird breeder feed containing
protein in pellet form.

quail
22 percent

For chicks, a 28 percent protein game bird starter, in crumble form, should
be used for six weeks.
Game bird grower, 26 percent protein, in a one-eighth
inch pellet form~ should be used from six to twelve weeks of age. The birds
may then be changed to a scratch feed, richer in carbohydrates, which will
provide them with more body heat during the winter months.
A suggested
scratch feed or winter maintenance moisture by weight would be 35 percent
whole milo, 30 percent whole wheat, 15 percent cracked corn, 10 percent
cracked barley, and 10 percent cracked oats (Webster unpublished notes).
Survival

of Young

During 1971, excellent survival (94.80 percent) was experienced for the 3
successful hatches for the first 7 days, and good survival (83.24 percent)
continued through the first 30 days.
In 1972, good survial (82.98 percent)
was experienced for the 2 successful hatches for the first 7 days, and fair
survival (78.72 percent) survival continued through the first 21 days.

Overwinter

Survival

in Holding

Pens by Bird Densities

Overwinter survival of mountain quail held in holding pens during the 1971-72
period ranged from a low of 86.67 percent in a pen with 26.40 square ft of
ground space per bird, containing 60 1971 hatched quail to a high of 100.00
percent in a pen with 144.00 square ft of ground space per bird containing
11 1971 hatched quail.
None of the densities tested in 1972 exceeded those
of 1971 because fewer numbers of mountain quail were held.

Facilities

Required

Adequate breeding pen facilities are necessary in mountain quail propagation.
Isolation of breeders is important and breeding pens should have adequate
natur
cover.
Breeders should be selected and moved to individual breeding
pens by early February to allow time for these easily stressed birds to
become accustomed to their new environment.
Wire floored pens proved unsatisfactory for housing mountain quail.
Clumps of tall wheatgrass were used
to provide adequate natural cover in breeding pens.

�-353-

winter holding pens such as those shown in Fig. 4 are adequate for holding
60 birds per each 20' X 76v pen when protection and seclusion is provided,
using bales of hay plus natural weed and grass cover.

LITERATURE CITED
Hoffman, D. M. 1969. Mountain quail production techniques. Job Prog. Rept.,
Fed. Aid Proj. W-37-R-22, WP 15, Job 3. Colo. Div. of Game, Fish and
Parks, Denver. p. 159-160.
1970. Experimental breeding of mountain quail. Job Prog. Rept.,
Fed. Aid Proj. W-37-R-23, WP 15, Job 3. Colo. Div. of Game, Fish and
Parks, Denver. p. 171-179.
1971. Experimental breeding of mountain quail. Job Prog. Rept., Fed.
Aid Proj. W-37-R-24, WP 15, Job 3, Colo. Div. of Game, Fish and Parks,
Denver. p. 85-90.
1972. Experimental breeding of mountain quail. Job Prog. Rept., Fed.
Aid Proj. W-37-R-25, WP 15, Job 3. Colo. Div. of Wildl., Denver.
p. 77-79.
Muller, H. D.

Unpubl.

Inbreeding handout.

Colo. State Univ.

2 p.

Swope, H. M. 1968. Experimental breeding of mountain quail. Job Prog.
Rept., Fed. Aid Proj. W-37-R-21, WP 15, Job 3. Colo. Div. of Game, Fish
and Parks, Denver. p. 117-119.
Webster, L. A.

Unpubl. Notes.

Colo. Div. of Wildl., Ft. Collins.

Wildlife Researcher

��April, 1973

-355-

JOB PROGRESS REPORT

State of

~C~O=L~ORA~t~D~O~

_
Game Bird Survey

W-37-R-26

Project No.
Work Plan No.

17

Job No.

2

Job Title

Continued Inventory of Selected Ptarmigan populations

Period Covered:

Apri11,

Personnel:

1972 through October 20, 1972

Clait E. Braun, Howard Funk, Richard Hoffman, Terry A. May and
Charles Wagner.

ABSTRACT

Investigations of white-tailed ptarmigan (Lagopus leucurus) populations in
Colorado initiated in 1965 were continued in 1972 with major emphasis in
ascertaining population levels and relationships of nesting success and
production to climatic conditions. Breeding densities decreased slightly
at Rocky Mountain National Park and Independence Pass and declined significantly at Crown Point. Reasons for the slight reductions at Rocky Mountain
National Park and Independence Pass were unknown, but were linked to lower
overwinter hen survival and poor recruitment of subadults into the breeding
population. The major cause of the decrease at Crown Point was failure of
birds to emigrate into the area following the 1971 removal. The breeding
density at Mt. Evans increased significantly, the result of closure of the
hunting seasons in 1970 and 1971. Nesting success was higher on all areas
than in 1971 but production to September 1 increased only at Mt. Evans. Survival of chicks to September 1 was similar to most previous years studied.
Weather conditions during the May through July period were warmer in May
and June, 1972 than in 1971, but were cooler in July. While exact moisture
data were not available, indications were that slightly more moisture fell in
June and July, 1972 than in 1971.

��-357-

CONTINUED

INVENTORY

OF SELECTED

PTARMIGAN

POPULATIONS

Clait E. Braun

Knowledge concerning population levels of white-tailed ptarmigan and factors
influencing observed densities is essential for proper management.
This
project was undertaken to examine long term population trends and effects
of climatic conditions on white-tailed ptarmigan in Colorado.
The initial
5 years of research on this grouse have been previously reported by Braun
and Rogers (1971). This report presents data collected during the third
year of the last 5-year segment.
Data collected during the first two years
of the second 5-year segment were presented by Braun (1971, 1972).

P. S. OBJECTIVE
To test the hypotheses that (1:)populations of white-tailed ptarmigan in
Colorado are not cyclic, (2) production and fall population densities of
this species in Colorado are related to temperature and moisture conditions
during June and early July of the same year, and (3) harvest of over 50
percent of the fall populations adversely affects breeding densities the
following spring.

SEGMENT

OBJECTIVES

1.

To estimate breeding population levels of ptarmigan in four study areas
(Mt. Evans, Crown Point, Independence Pass, and Rocky MOuntain National
Park) .

2.

To estimate nesting

3.

To obtain weather data from a representative alpine area for use in
determining correlations between spring weather conditions and fall
ptarmigan populations.

4.

Manipulate hunting seasons at Mt. Evans through closures,
late timing, etc. in order to affect breeding densities.

5.

To compile data and prepare

success and production

METHODS

progress

in the above areas.

reduced

length,

report.

AND MATERIALS

Techniques used were essentially those developed under Work Plan 17, Job 1
and reported in detail by Braun and Rogers (1971), and updated by Braun
(1971).
In 1972 pink bandettes with black numerals were used to mark all
newly banded birds and those whose older bandettes had become too worn for
individual recognition.
Weather data were obtained from the Institute of

�~358Arctic and Alpine Research (INSTAAR),University of Colorado, for reasons
previously explained (Braun 1971). Hunting statistics
were collected
through return of bands from successful hunters (Independence Pass and
CrownPoint) and a check station (Mt. Evans).
Description

of Area

Areas intensively investigated have been described in detail
Rogers (1971) and have been presented by Braun (1971).

by Braun and

RESULTS
ANDDISCUSSION
Breeding Densities
Censuses of breeding birds were initiated
on May2 in R.M.N.P. with only
solitary males being observed. Breeding surveys were initiated
on May16
at Mt. EVans, May 23 at CrownPo.int and May 31 and Independence Pass. While
SlOme
pa.iring was observed as early as April 17 at CoronOaPass and April 18
at Loveland Basin, most pairing did not occur until between May4 and 15.
Timing of breeding events was somewhat earlier than in 1971 and was similar
to 1969 and 1970. Densities observed are presented in Table 1.
Table 1.

White-tailed

ptannigan breeding densities,

Study Area

all

areas,

1972.

No. of
Breeding
Pairs

Unmated
Males

Total
Breeding
Population

Birds per
Square
Mile

Rocky Mountain National Park
Tombstone RidgeSundance Mm.

1.25

6

5

17

13.6

Toll Me~w:)1';,ial

.19

4

4

12

63.2

s

.70

8

3

19

27.1

Total Rocky Mtn.
National Park

2.14

18

12

48

22.4

CrownPoint

1.93

o

2

2

1.0

Mt. EVans

1.54

13

4

30

19.5

Independence Pus

1.12

8

5

21

18.7

Fall R1ver P

1/ In square miles.

. ...

�-359-

Breeding densities were lower on all study areas except Mt. Evans in 1972
from levels observed in 1971. Decreases were not significant at either
R.M.N.P. or Independence Pass, but the low number of birds at Crown Point
was much less than anticipated. Reasons for the observed decline in numbers
of breeding ptarmigan were not apparent although recruitment of subadults
into spring populations was low. While the decrease at Independence Pass
fell within the range observed in prior years (Table 2), the decrease at
R.M.N.P. continued the reduction of numbers of breeding birds that has been
observed since 1969. The increase at Mt. Evans was the second since 1970
and was the result of two years of hunting season closure at that site.
The decrease at Crown Point followed the complete removal (Work Plan 17,
Job 3) of all breeding birds in 1971.

Table 2.

White-tailed ptarmigan breeding densiities, 1966-72.

Study Area

1966

1967

Birds Per Sguare Mile
1968
1969
1970

1971

1972

Rocky Mo lID tain
National Park

29.0

25.2

29.4

30.8

24.8

23.4

22.4

Crown Point

14.5

21.2

18.1

14.5

10.4

5.7

1.0

Mt. Evans

7.8

7.1

7.1

5.8

5.2

11.0

19.5

Independence Pass

18.7

18.7

18.7

17.9

10.7

21.4

18.7

Wnile the trend in breeding densities at R.M.N.P. is downward, breeding
populations appear to have stabilized as witnessed by only the slight
fluctuation from 1970 to 1972. However, recruitment of subadults remains
low, with most of the breeding population being comprised of birds in the
older age classes. At Crown Point, the trend has been downward since 1967
with the low densities in 1970 and 1971 being due, in part, to heavy hunting
pressure in 1969 and 1970. The marked decrease at this site in 1972 was
due in part to the removal of all breeding birds in 1971 as part of an
experiment (Work Plan 17, Job 3). Recovery is slower than expected,
possibly due to the distance this site is from other occupied range, and the
few subadults available in the Mummy Range in the spring of 1972. The population at Mt. Evans continued the increase first observed in 1971 and is the
direct result of the hunting closure at this site in the fall of 1970 and
1971. This increase is due to good survival of all age classes, average to
good production in 1970 and 1971 and excellent recruitment of subadu1ts
into the breeding population. With the exception of 1970, the breeding
population at Independence Pass has remained remarkedly stable since 1966.
Apparently little affects the breeding population at this site except
severe hunting pressure during at least two successive years.

�Table 4.

Estimated fall densities of white-tailed ptarmigan, 1972.

Total
Production

Total
Breeding
Population

Total
Population
on
September 1

Birds Per
Square Mile

Pe rcen t Gain

3.3

43

48

86

40.2

50.0

0

0

0

2

2

1.0

0.0

Mt. Evans

75

4.4

44

30

71

46.1

62.0

Independence Pass

45

4.5

14

21

33

29.5

42.4

Estimated
Percent
Nesting
Success

Average
Brood Size
on
September 1

Rocky Mountain
NationaL Park

75

Crown Point

Area

I

w

0\
N
I

Table 5.

Weather data, Niwot Ridge, 12,300 ft, May-July, 1972.
Number Days
Minimum
Temperature
Less Than 32° F

Pre cipitation/
Month
(Inches)

Number Days
Relative
Humidity
90 or Above

Average
Wind Speed/
M:mth

Month

Maximum

Temperature
Mean Dail~
Maximum
Minimum
Hinimum

May

49° F

- 1° F

35.6° F

22.8° F

29

Y

18

12.5

June

53° F

28° F

48.60 F

34.2° F

7

1/

20

12.5

July

61° F

26° F

53.3° F

.37.7°

F

4

Y

17

11. 3

1/ Data not presently available.

(mph)

�-363-

April, 1973

JOB PROGRESS REPORT

State of

COLORADO
---------~~~~------------

Project No.

·W-37-R-26

Work Plan No.

17

Job Title

Job No.

3

ExperiITenta1 Removal of a Breeding population of White-tailed Ptarmigan

Period Covered:
Personnel:

Game Bird Survey

April 1, 1972 through September 30, 1972

C1ait E. Braun.

ABSTRACT

Despite intensive searching, only two white-tailed ptarmigan (Lagopus 1eucurus)
(both males) were located on the Crown Point study area in 1972. All areas
occupied were sites previously used by territorial males in preceding years.
One male was banded and was a subadult. Behavior of the one unbanded male also
suggested that it was a subadult.

�-364-

.i

RECOMMENDATIONS

1.

It is recommended that examination of blood parameters "be discontinued
due to small samples involved.

�-365-

EXPERIMENTAL REMOVAL OF A
BREEDING POPULATION OF WHITE-TAILED PTARMIGAN
Clait E. Bratlll
Management of wild animals is dependent upon knowledge about population
levels, reproductive capacity and success, survival and mortality rates,
and habitat requirements. This study to investigate habitat preferences and
pioneering ability of white-tailed ptarmigan was initiated in 1971 after
five years of previous study of the Crown Point population (Braun and Rogers
1971, Braun 1971). This report presents data collected during the second
year of a 4 year study. Data collected during the initial year of the study
were presented by Braun (1972).
P. S. OBJECTIVE
To test the hypotheses. that (1) ptarmigan select sites for breeding territories which contain readily available bushes of Salix spp. (&gt;10 cm in
height) and snowfree locations, (2) immature birds initially pioneer new or
empty habitats, and (3) number of occupied territories is affected by age
class of male ptarmigan.
SEGMENT OBJECTIVES
1.

To map and compare number and characteristics of all occupied breeding
territories at Crown Point.

2.

To obtain blood samples from all breeding birds present for eventual
establishment of blood parameters for resident and pioneering ptarmigan.

3.

To compile data and prepare progress report.
METHODS AND MATERIALS

Ptarmigan were located during the breeding season through use of taperecorded male challenge calls as described by Braun et al. (1973). Techniques
used in capturing, observing, banding, etc. were essentially those described
in detail by Bratllland Rogers (1971) and updated by Braun (1972).
Description of Study Area
This area was described in detail by Braun and Rogers (1971) and will be
redescribed in the final report. Extent of the area investigated was shown
in the initial report (Braun 1972).

�-366-

RESULTS AND DISCUSSION
Population Status
The study area was systematically surveyed 3 times during the breeding season
and twice during the late summer brood period. Only two males were located
during the entire period surveyed. One male was captured, banded and
released. This bird was a subadult (less than one year of age). The other
bird was observed and was unbanded. Efforts to capture it were unsuccessful.
This bird exhibited all the characteristics of an unmated, non-territorial
subadult. Ptarmigan P31 was initially observed just above the saddle between
Crown Point and Crown Mountain in mid-May, but subsequently moved south to an
area about 200 yards north of the top of Crown Mountain where it appeared
territorial. Both areas where this male were observed had been occupied by
territorial males in previous years. The unbanded male was also observed in
the area just north of the summit of Crown Mountain on a previously occupied
territory, but this male did not appear to be territorial. Neither of these
two birds had prior experience at Crown POint, but both selected areas which
had been previously occupied. This supports the hypothesis that willow and
snow free areas are essential for breeding territories. At least one bird
was a subadult and behavior of the second male also suggested that it too
was less than one year of age, supporting the hypothesis that subadult
birds tend to pioneer new or vacant habitats.

Description of Breeding Territories
Since all areas occupied or used by the two males in 1972 had been previously
described (Braun 1971). no mapping was done in 1972. Present plans are for
all occupied areas to be remapped and described in 1974.

Blood Parameter Examination
Due to small samples involved (one bird), blood was not obtained for electrophoresis study. It is felt that samples will be too small for analysis of
blood parameters of resident and pioneeering ptarmigan. Consequently, it is
recommended that this aspect of the study be terminated.

LITERATURE CITED
Braun, C. E. 1971. Habitat requirements of Colorado white-tailed ptarmigan.
Proc. Western Assoc. State Game and Fish Comms. 51:284-292.
1972. Experimental removal of a breeding population of white-tailed
ptarmigan. Colo. Div. Wildl., Game Res. Rept. Fed. Aid Proj. W-37-R.
April. p. 99-109.

�-367-

Braun, C. E., and G. E. Rogers. 1971. The white-tailed ptarmigan in
Colorado. Colo. Div. Game, Fish and Parks. Tech. Publ. No. 27. 80 p.
_____ , R. K. Schmidt, Jr., and G. E. Rogers. 1973.
white-tailed ptarmigan with tape-recorded calls.
37 (1):90-93.

Prepared by

_~-::-=-tfa_U
__' __r._o ~~"-Clait E. Braun
Wildlife Researcher

_

Census of Colorado
J. Wildl. Mgmt.

�-368-

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!

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'-"

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.r: '.

.-

'-

'

---- ...•..
~ r
--------.:..

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--. '...
-~ -"'...

.

10600

I

i
I
, I
.•.I.
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I'

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tJ""~

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CROWN

POINT

AREA

COLORADO
-

Fig. 1.

Study Area Boundary

Crown Point study area (x's mark occupied breeding areas, 1972).

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                  <text>-1-

July, 1973

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-lOl-R-15

Work Plan No.

4

Game Range Investigations
Job No.

la

Job Title

Inventory of Range Manipulation

Period Covered:

April 1, 1972 through March 31, 1973

Personnel:

Roland C. Kufeld and Regional

Projects

in Colorado

Game Biologists

ABSTRACT
An inventory was made of all range vegetation-modification
projects completed
during 1971, in the western half of Colorado, on lands administered by the
U.S. Forest Service and Bureau of Land Management.
Acreages treated were:
Forest Service - 6,728 acres and Bureau of Land Management - 2,822 acres.
Total - 9,550 acres.

��-3-

INVENTORY

OF RANGE MANIPULATION

PROJECTS

IN COLORADO

Roland C. Kufeld

F. S. OBJECTIVE
To collect information which describes proposed and completed range vegetation modification projects that are located in Colorado and their effects
on the range, wildlife and livestock; and to provide desired IBM listings
of these data to cooperating agencies upon their request.

SEGMENT OBJECTIVES
1.

To collect information which describes proposed and completed
vegetation modification projects that are located in Colorado
their effects on the range, wildlife and livestock.

2.

To compile,

3.

To provide desired IBM listings
agencies upon their request.

codify, process

METHODS

and analyze

inventory

of inventory

range
and

data.

data to cooperating

AND MATERIALS

An inventory was made of all range vegetation modification projects completed
through 1970, in Colorado, west of Interstate Highway 25, on lands administered by the U.S. Forest Service and Bureau of Land Management
(Kufeld 1968,
1970 and 1971).
Kufeld (1968 and 1970) also covered projects completed on
the Southern Ute and Ute Mountain Indian Reservations through 1969.
Indian
Reservations projects were dropped from the inventory after 1969. This
report concerns vegetation modification projects completed on U.S. Forest
Service and U.S. Bureau of Land Management lands west of Interstate Highway
25 during 1971.
Data were collected using procedures outlined in Colorado Division of Wildlife Administrative Directive No. 24, entitled "Range Vegetation Modification
Projects", and by Kufeld (1968 and 1970).
Inventory data were transferred from original
Reader Forms, and then to I.B.M. cards.

data sheets

to Mark Page

Procedures outlined in Administrative Directive No. 24, call for evaluations
to be made on each vegetation modification project at the end of the 2nd,
5th, and 10th years following treatment.
During 1971, 2 year post-treatment

�-4-

evaluations were made on 23 vegetation modification projects that were
completed in 1969, and data recorded on a form described by Kufeld (1971).
Seven requests for retrieval of range vegetation modification project
inventory data were processed during the 14th segment. These were for
Stearns-Roger Corporation of Denver, Colo.; Future Studies Organization
of Pioche, Nevada; Colorado Cooperative Wildlife Research Unit; Colorado
Wildlife Commissioner William Robinson; U.S. Forest Service Rocky Mountain
Regional Office; and two for the Colorado State Office of the U. S. Bureau
of Land Management.

RESULTS AND DISCUSSION
Acreages treated during 1971, on all lands administered by the U.S. Forest
Service and Bureau of Land Management are shown by vegetation type and kind
of treatment in Tables 1 through 3. These acreages are shown by individual
National Forests and Bureau of Land Management Districts in Tables 4 and
5.
A total of 9,550 acres were treated in 1971, of which 6,728 were treated
by the U.S. Forest Service, and 2,822 acres by the Bureau of Land Management. To date (through Dec. 31, 1971) a total of 578,382 acres have been
treated in Colorado on lands administered by the U.S. Forest Service and
Bureau of Land Management. Of this total 238,875 acres were on Forest
Service and 339,507 acres were on Bureau of Land Management lands.
Since 1971 was only the second year for 2 year post-treatment evaluations
of completed vegetation modification projects, data from post-treatment
evaluations of 1968 and 1969 projects will be kept on file and covered
in a future report when enough 2 year post-treatment evaluations have been
made to provide a basis for a meaningful data analysis.

LITERATURE CITED
Kufeld, Roland C. 1968. Inventory of range manipulation projects in Colorado.
Colo. Div. Game, Fish and Parks. Game Res. Rept. P.R. Project W-lOl-R-lO,
WP 4, Jla, July, Part 1. p. 1-121.
1970. Inventory of range manipulation projects in Colorado. Colo.
Div. Game, Fish and Parks. Game Res. Rept. P.R. Proj. W-lOl-R-12,
WP 4, Jla, July, Part 1. p. 59-94.
1971. Inventory of range manipulation projects in Colorado. Colo.
Div. Game, Fish and Parks. Game Res. Rept. P.R. Proj. W-lOl-R-13,
WP 4, Jla, July, Part 1. p. 1-15.

Roland C. Kufeld
Wildlife Researcher

�-5-

Table 1. Acreages of rangeland
the u.s. Forest Service.

treated during 1971, in Colorado

Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grass

Seed Only

2

390

390

Spray

2

160

160

Pitting

5

455

455

9

1,005

1,005

Spray

8

3,934

0

Pitting

1

150

150

9

4,084

150

Spray

1

60

60

Chain 2:...1

5

859

859

Burn

2

660

60

Hand Cutting

1

60

0

9

1,639

979

27

6,728

2,134

Sagebrush

Browse

1:./

All Vegetation

Types GRAND TOTAL

!/All vegetation modification projects
vegetation type were in the "oakbrush"

conducted
subtype.

2:.../Chaining
includes

railing

chaining,

cabling,

Table 2. Acreages of rangeland treated during
the U.S. Bureau of Land Management.

during

by

1971 , in the "browse"

and bulldozing.

1971, in Colorado by

Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Sagebrush

Seed Only

2

1,222

1,222

Pinyon-Juniper

Chain -

2

1,600

1,600

All Vegetation

Types GRAND TOTAL

4

2,822

2,822

!/Chaining

II

includes

chaining,

cabling,

railing

and bulldozing.

�-6-

Table 3. Acreages of rangeland treated during 1971, in Colorado by
the U.S. Forest Service and u.S. Bureau of Land Management.

-----

Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grass

Seed Only

2

390

390

Spray

2

160

160

Pitting

5

455

455

9

1,005

1,005

Seed Only

2

1,222

1,222

Spray

8

3,934

0

Pitting

1

150

150

11

5,306

1,372

Spray

1

60

60

Chain ]j

5

859

859

Burn

2

660

60

Hand Cutting

1

60

0

9

1,639

979

2

1,600

1,600

2

1,600

1,600

31

9,550

4,956

Sagebrush

Browse 1.1

Pinyon-Juni per Chain'l;./

All Vegetation Types GRAND TOTAL

1/
- All vegetation modification projects conducted during 1971, in the "browse"
vegetation type were in the "oakbrush" subtype.

2/
- Chaining includes chaining, cabling, railing and bulldozing.

�-7 -

Table 4. Acreages of rangeland treated during 1971, within each
National Forest in Colorado.
National
Forest

Vegetation
Type

Kind of
Treatment

Grand Mesa

Sagebrush

Spray

Browse 1./

Routt

San Juan

Browse 1./

Grass

Sagebrush

Browse 1./

Uncompahgre

Sagebrush

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

2

1,235

o

2

1,235

o

Spray

1

60

60

Burn

1

60

60

Chain 2:,./

1

60

60

3

180

180

1

60

o

1

60

o

Seed Only

1

215

215

Pit

5

455

455

6

670

670

1

150

150

1

150

150

1

600

o

1

600

o

4

1,699

o

4

1,699

o

Hand Cutting

Pit

Burn

Spray

�-8-

Table 4. Acreages of rangeland treated during 1971, within
National Forest in Colorado (continued).

each

National
Forest

Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres Seeded as
Part of Treatment

White River

Grass

Seed Only

1

175

175

Spray

2

160

160

3

335

335

2

1,000

0

2

1,000

0

4

799

799

4

799

799

27

6,728

2,134

Sagebrush

Spray

. 2/
Ch aln-

Browse }/

Total all Forests,
and Treatments

Vegetation

Acres
Treated

Types

1/
- All vegetation modification projects
vegetation type were in the "oakbrush"
~/Chaining

includes

chaining,

conducted
sUbtype.

during 1971 in the "browse"

cabling, railing and bulldozing.

Table 5. Acreages of range land treated during 1971, within
Land Management District in Colorado.

each Bureau of

BLM
District

Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres Seeded as
Part of Treatment

Montrose

Sagebrush

Seed Only

1

102

102

1

102

102

1

1,120

1,120

1

1,120

1,120

2

1,600

1,600

2

1,600

1,600

4

2,822

2,822

Grand Junction

Sagebrush

Seed Only

PinyonJuniper

Chain }/

Total all Districts,
and Treatments

Vegetation

1.lChaining includes

chaining,

Acres
Treated

Types

cabling, railing

and bulldozing.

�-9-

July,

1973

JOB PROGRESS

State

Project
Work

COLORADO

of
No.

W-10l-R-1S

plan No.

4

Job Title

Period

Covered:

Personnel:

Game Range

---------

April

Roland

Investigatio_n__s

_

2
Job No.
Experimental
Improvement of Oakbrush on
Deer and Elk Winter Ranges-B~e~a~v~e~r~C~r~e~e~k~

1, 1972 through

March

_

31, 1973

C. Kufeld

ABSTRACT
Elk use between September and May increased 73 percent on a 67 acre Gambel
oak type range two years after spraying with 2,4,S-TP at a rate of 2 lbs. per
acre mixed in 10 gallons of water per acre.
Game use was measured by pellet
group counts made on the sprayed area, and compared with similar counts on
an adjacent 68 acre control.
Deer use on the same areas decreased 41 percent;
however, the sprayed plot received 16 percent more deer use than the control
acreage.
Ninety-nine
food habits studies were combined to determine what plants are
normally eaten by mule deer and the relative value of these plants from a
manager's viewpoint, based on the response deer have exhibited toward them.
A paper summarizing 48 elk food habits studies and ranking plants eaten by
relative value was published in The Journal of Range Management
26(2):106-113.
A decision was made to delay analysis of 2 year post-treatment
vegetation
measurement data until the literature review on forage value of oak study
plants for elk and deer could be completed.

area

��-11-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH ON
DEER AND ELK WINTER RANGES-BEAVER CREEK
Roland C. Kufe1d

P. S. OBJECTIVE
To determine if deer and elk forage production and game use can be increased
on overage Gambe1 oakbrush winter game ranges by spraying with 2,4,S-TP to
induce sprouting.

SEGMENT OBJECTIVES
1.

To measure deer and elk use on the study area following treatment
determine changes in game use that have occurred as a result of
spraying with 2,4,S-TP in 1969.

2.

To review the literature in order to determine
oak study area plants for mule deer and elk.

3.

To summarize and analyze pre- and post-treatment
use measurement data.

vegetation

4.

To write a report of the findings
journal.

in a scientific

to

the forage value of

for publication

and game

METHODS AND MATERIALS
Deer and elk pellet groups which were deposited
plots after the plots were cleared on September
May 8, 1972.

in circular, 0.001 acre
1, 1971, were counted on

An extensive review of literature was made to determine the forage value
of oak study area plants for mule deer. Ninety-nine food habits studies
were combined to determine what plants are normally eaten by mule deer, and
the relative value of these plants from a manager's viewpoint, based on the
response deer have exhibited toward them.

RESULTS AND DISCUSSION
Effects

of Spraying

on Elk

The overall elk population in the vicinity of the study area appears to
have been higher in 1972, the year of 2 year post-treatment evaluation, than
1969, when pre-treatment game use measurements were made.
Aerial trend counts

�-12-

flown in January in the Williams Fork area (Game Management Unit 12, where
the study area is located) showed 1,217 elk during the winter of 1968-69
and 1,540 during the winter of 1971-72 (White 1972).
This 27 percent
increase may have been due to the restrictions on shooting spike bulls and
the reduction of cow permits during the 1971 elk season.
Data showing changes in elk use on the study area two years following
spraying with 2,4,5-TP are presented in Table 1. Prior to spraying the
control area received twice as much elk use as the area designated for
spraying.
Two years after spraying, elk density per square mile on the
spray areas, as reflected by pellet counts, increased 106 percent, while a
19 percent increase was recorded on the control area.
Comparison of ratios
of elk use intensity measured in 1969 and 1972 on the control with the same
ratios on the spray area indicates the increase in elk density per square
mile attributable to spraying is 73 percent.
This is an increase in elk
density of 67 animals per square mile. Elk use on the study area two years
following spraying was quite heavy, averaging 62.5 elk days use per acre.
The 19 percent increase on the control area roughly coincides with the 27
percent population increase reflected by aerial counts. However, part of
the increase on the control area may have been due to it's close proximity
to the spray area. Elk attracted to the spray area would also use the
control because it is adjacent.
Thus, the actual increase due to spraying
could have been higher than 73 percent, but doubtful if much higher.

Effects

of Spraying

on Deer

Deer use on the study area (spray and control areas combined) as represented
by "deer per square mile" was down 41% in 1972 from the 1969 level.
It is
not known whether this represents a drop in the overall deer population
because there were no aerial deer trend counts flown in the Williams Fork
area in January, 1972 to provide pre- and post-treatment population comparisons.
It may be possible, however, that the decline in deer use on the
study area is due to the high elk use which forced the deer to leave.
Deer use in 1972 declined 45% on the control area and 37% on the spray
area from levels recorded in 1969. Since deer use declined less on the
sprayed portion than on the control, comparison of ratios of deer use
intensity before and after treatment on the two areas suggests that spraying,
in effect, resulted in an increase of 16 percent in deer density per square
mile (Table 1).

Literature

Review

Deer forage plants in the 99 food habits studies were categorized by
heavily, moderately or lightly eaten.
This information will be used to
determine the benefits of vegetation changes caused by spraying.
Current
plans call for submitting the information for publication as a U. S. Department of Agriculture Conservation Research Report.
Authors will be Roland

�-13-

C. Kufeld of Colorado Division of Wildlife, and o. C. Wallmo and Charles
Feddema of the U. S. Forest Service, Rocky Mountain Forest and Range
Experiment Station.
A paper entitled "Foods Eaten by the Rocky Mountain Elk" by Roland C.
Kufeld was published in The Journal of Range Management 26(2) :106-113.
This was based on a literature review of elk food habits completed during
the 15th project segment.
Forty-eight food habits studies were combined
to determine plants normally eaten by the Rocky Mountain elk (Cervus
canadensis nelsoni), and the relative value of these plants from a manager's
viewpoint based on the response elk have exhibited toward them. Plant
species are classified as highly valuable, valuable, or least valuable on
the basis of their contribution to the diet in food habits studies where
they were recorded.
A total of 159 forbs, 59 grasses, and 95 shrubs are
listed as elk forage and categorized according to relative value.

Analysis

of Post-treatment

Vegetation

Measurement

Data

A decision was made to delay analysis of 2 year post-treatment vegetation
measurement data until the literature review on forage value of oak study
area plants for elk and deer could be completed.
Thus, the vegetation
measurement data are not presented herein.

LITERATURE

CITED

White, Claude E. 1972. Principal Game Biologist,
Div. Wildlife.
Personal communication.

Prepared

by -L;e_),,--::Cl....:.{"-....·(.:;:,.i.::.;;..l.i..::::;.;{/
__
C_l_, ,_L,-='--:....:..{.~Lr-::.~=t-=j-';/{::::.L·l
Roland C. Kufeld
t
Wildlife Researcher

Northwest

_

Region.

Colo.

�Table 1.

Changes in elk and deer use two years following spraying with 2,4,5-TP on the Beaver Creek oak study area.1/
Animal Days
Use Per Acre
Observed
Change Due 5/
To SprayingI\.
No.
E4/
No.
%

No.

Animal Density /
3
Per Square MileObserved
Change Due
I\.
To Spraying

Acres Per
Animal Pe r Month
Observed
Change Due
To
Spraying
I\.
No.
E
No.
%

Animal

Area

Year

Total
Pellet
Groups~/

Elk

Control

1969

41

55

l36

0.552

1972

47

63

162

0.484

1969

23

31

77

0.975

1972

47

62

Spray

36

+26

+72

159

E

92

No.

+67

%

+73

0.493

0.855

-0.362

-42
I
•.....

Deer

Control

Spray

1969

59

80

198

0.379

1972

31

42

108

0.726

1969

42

56

138

0.544

1972

26

34

29

+ 5

1/ Pre-treatment period of use was from 8-19-68 to 5-5-69.

+17

87

75

+12

+16

0.902

+'"I

1.042

-0.140

-13

Post-treatment period of use was from 9-1-71 to 5-8-72.

2/ Spray area contains 58 .001 acre plots and totals 67 acres. Control area contains 57 .001 acre plots and totals 68 acres.

1/ This reflects percent change more accurately than animals days use per acre, because it takes the length of the pellet

accumulation period into account.
~/ Denotes the amount of use expected on the spray area in 1972 if elk and deer use on the spray area in 1969 and 1972
had occurred in the same proportion as on the control area.
2/ The change in animal use which can be attributed to spraying. This has been adjusted to consider animal use changes
on the control area.

�July, 1973

-15-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-lOl-R-15

Work

Job Title

3
Job No.
Experimental Improvement of Oakbrush
on Deer, Elk and Cattle Ranges - Hightower Mountain

Period Covered:

April 1, 1972 through March 31, 1973

Plan No.

Personnel:

Game Range Investigations

4

Roland C. Kufe1d

ABSTRACT

Post-treatment photos were taken at permanent photo points established prior
to treatment in 1971. These show effects of spraying, burning and chaining
during the first summer following treatment.
Ninety-nine food habits studies were combined to determine what plants are
normally eaten by mule deer, and the relative value of these plants from a
Manager's viewpoint based on the response deer have exhibited toward them.
A paper summarizing 48 elk food habits studies and ranking plants eaten by
relative value was published in The Journal of Range Management 26(2):106-113.

��-17-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH
ON DEER, ELK AND CATTLE RANGES - HIGHTOWER MOUNTAIN
Roland

C. Kufeld

P. S. OBJECTIVE
To determine the extent to which deer, elk and cattle forage production
and game use can be increased and maintained by chaining, spraying and
controlled burning on overage Gambel oak winter game ranges.

SEGMENT OBJECTIVES
1.

To replace study plot stakes destroyed
construction and anchor chaining.

2.

To retake photographs

3.

To review the literature in order to determine the forage value of
oak study area plants for mule deer, elk and cattle.

at established

METHODS

as a result of fire lane

photo points.

AND MATERIALS

Study Plot Stake Replacement
Permanent vegetation measurement and pellet plot stakes destroyed as a
result of fire lane construction and anchor chaining were replaced using
surveying techniques described by Kufeld (1971). In most cases the old
steel stakes were bent over flat against the ground and could be straightened.
In very few cases were the stakes actually missing so the plot locations
had to be resurveyed.

Photo Point Photographs
Post-treatment photos were taken at permanent photo points established
prior to treatment in 1971. These show effects of spraying, burning and
chaining during the first summer following treatment.
A black and white
and a 35 mm color slide was taken at each photo point with the same cameras
used to take pre-treatment photos.

Literature

Review

An extensive review of literature was made to determine the forage value
of oak study area plants for mule deer. Ninety-nine food habits studies

�-18-

were combined to determine what plants are normally eaten by mule deer,
and the relative value of these plants from a manager's viewpoint based
on the response deer have exhibited toward them.

RESULTS AND DISCUSSION

Photo Point Photographs
Photos showing conditions in the chained, sprayed and burned areas before
and the first summer after treatment are shown in Fig's. 1,2 and 3. From
initial appearances chaining seems to have done the best job of opening
the oak stand with burning second and spraying third. The first posttreatment evaluation to determine which of the three treatments was most
successful in increasing forage production and deer, elk and cattle use,
however, will be made during the summer of 1973, which is the second
growing season after treatment.
Sprouting of oak (Quercus gambellii), serviceberry (Amelanchier a1nifolia),
chokecherry (Prunus virginiana), and snowberry (Symphoricarpos sp.) was
profuse in the two burned units (Fig. 4), and occurred to a lesse~ extent
in the chained areas.
Sprouting appeared to be least prominent in the
sprayed units.
Reseeding success during the first summer after treatment was very poor.
Young plants of all seeded grass and forb species listed by Kufe1d (1972)
were observed, but germination was very spotty.
No germination of bitterbrush (Purshia tridentata) or mountain mahogany (Cercocarpus montanus)
was observed.
Poor germination is attributed to the extremely dry winter,
spring and summer of 1972. The best seeding success occurred in the
burned units with intermediate wheatgrass (Agropyron intermedium) (Fig. 5).
Some of the best seeding success with other reseeded grasses and forbs
occurred in the fire lanes constructed around the two burned units.
Hopefully the rest of the seed will remain viable and undisturbed and germinate
during the 1973 growing season.

Literature

Review

Deer forage plants in the 99 food habits studies were categorized by heavily,
moderately or lightly eaten.
This information will be used to determine the
benefits of vegetation changes caused by spraying, burning, chaining and
reseeding.
Current plans call for submitting the information for publication
as a U.S. Department of Agriculture Conservation Research Report.
Authors
will be Roland C. Kufeld of Colorado Division of Wildlife, and O. C. Wallmo
and Charles Feddema of the U.S. Forest Service, Rocky Mountain Forest and
Range Experiment Station.
A paper entitled "Foods Eaten by the Rocky Mountain Elk" by Roland C. Kufeld
was published in the Journal of Range Management 26(2):106-113.
This was
based on a literature review of elk food habits completed during the 15th

�-19-

project segment.
Forty-eight food habits studies were combined to determine plants normally eaten by the Rocky Mountain elk (Cervus canadensis
nelsoni), and the relative value of these plants from a manager's viewpoint based on the response elk have exhibited toward them. Plant species
are classified as highly valuable, valuable, or least valuable on the
basis of their contribution to the diet in food habits studies where they
were recorded.
A total of 159 forbs, 59 grasses, and 95 shrubs are listed
as elk forage and categorized according to relative value.

LITERATURE

CITED

Kufeld, Roland C. 1971. Experimental improvement of oakbrush on deer,
elk and cattle ranges - Hightower Mountain.
Colo. Div. of Game,
Fish and Parks.
P-R Proj. W-lOl-R-13, WP-4, J-3, Game Research
Rept. Part 1. pp. 23-86.
1972. Experimental improvement of oakbrush on deer, elk and
cattle ranges - Hightower Mountain.
Colo. Div. of Wildlife.
P-R
Proj. W-lOl-R-14, WP-4, J-3. Game Research Rept. Part 2. pp. 81-97.

;;

(

/

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I

Prepared

by

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I

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Roland C. Kufeld
Wildlife Researcher

"-1'-

-

,{,I

:....I_·\-::L

L

,:

�-20-

Fig. 1. Vegetation conditions on the chained areas one year before and
the first summer after double anchor chaining.
These photos are identical
views taken from the same permanent photo point.

�-21-

Fig. 2.

Vegetation

conditions

on the sprayed areas one year before and the

first summer after spraying with 2,4,5-TP.
taken from the same permanent

photo point.

These photos are identical views

�-22-

Fig. 3. Vegetation conditions on the burned areas one year before and
the first summer after burning.
These photos are identical views taken
from the same permanent photo point.

�-23-

Fig. 4. Vegetation conditions on the burned area one day after and the
first summer after burning shows profuse resprouting. These photos are
identical views taken from the same photo point.

�-24-

Fig. 5. Seeding success of intermediate wheatgrass in a burned area.
Unfortunately such seeding response was obtained in only a few scattered
areas.

�July, 1973

-25-

JOB PROGRESS REPORT

State of

COLORADO

----~~~~~-------------

Project No.

W-101-R-15

Work Plan No.

5

Job Title

1

Job No.
Bighorn Sheep Range Inventory

Period Covered:
Personnel:

Game Range Investigations

April 1, 1972 to March 31, 1973

Harold R. Shepherd, Jack E. Gustafson,
Rowe and Robert J. Saban

James R. Hayward, Douglas J.

ABSTRACT

A range inventory of the Trickle Mountain Bighorn Sheep Study Area, near
Saguache, Colorado was completed.
This included preparations for and sampling
with the Daubenmire canopy-coverage method an area of 7,270 acres.

\

��-27-

BIGHORN

SHEEP RANGE INVENTORY

Harold R. Shepherd

P. S. OBJECTIVE
To inventory, contrast and evaluate
ductive and an unproductive bighorn

certain range attributes
sheep habitat.

of a pro-

SEGMENT OBJECTIVES
1.

Determine the distribution and extent of vegetation types in relation
to physical features of the habitat on ranges with good (Trickle
Mountain) and poor (Buffalo Peaks) bighorn sheep production.

2.

Estimate percentages plant species composition, frequency, crown cover,
litter, rock and bare ground on ranges with good (Trickle Mountain)
and poor (Buffalo Peaks) bighorn sheep production.

METHODS

Establishing

AND MATERIALS

Study Area Boundaries

Department personnel who, in previous years, had observed the habits and
movements of bighorn sheep in the general Trickle Mountain area had come
to recognize two distinct concentration areas.
These had come to be known
as the Winter Range, and Lambing Range.
A long, narrow strip of country
along East Pass Creek through which sheep traveled be tween the Winter Range
and Lambing Range we named the Intermediate Range.
The locations of these
ranges are shown in Fig. 1. The Winter Range is comprised of 3,241 acres,
the Intermediate Range of 2,130 acres, and the Lambing Range of 1,899 acres.

Preparation

of Photo Mosaics

A photo mosaic of the Trickle Mountain study area, Fig. 2, was prepared
by arranging 9 x 9-inch black and white aerial prints in overlapping flightline patterns and stapling them to a 4 x 8-foot sheet of plywood, painted
white.
Additional large photo numbers were placed on selected photos for
better legibility.
Study areas names of prominent topographic features
and landmarks were supplied.
The mosaic was then photographed by a photogrammetric company and reduced to several 20 x 24-inch prints for laboratory
and field use. The mosaic prints were useful for selection of proper color
prints for recording vegetation type and transect location and for correlation
and identification of photo and topographic map features.

�1

B.L.M.

U.S.F.S.

Nl

---

IGHORN SHEEPSTUDY AREA
Si~guache .County, Colo.

1--

1

UPPER
SAGUACHE
GUARD
. STATION

TRICKLE MOUNTAIN

1 MIL

tv
00

I

�Wieb"ctrCi

'~"

U.S.F. S.

B.L.M.

-.-A,\

•

N

\0

~~~
'-!({'(C' GUlQ£, ..-

'-'''~

NJ

---

UPPER
SAGUACHE
GUARD
, STATION

-

------ ------- ------

TRICKLE MOUNTAIN

BIGHOIRN SHEEP STUDY ARE

1 MILE

.-Fig. 1. Trickle ~ountain bi~horn sheep study area •
_- -_._._._. _._-.-._-------_._- ._--_._--------_ ..;--._-------------_._---_.-

..

--

Saguache County, Cole .

..",

-,

•

�TRICKLE MOUTAI:I
BIGHORN SHEEP STUDY AREA
Soguache
County. Cote
Pho!oqrophed

for

Cola

DiVision

Game,

rl~h

8 P(J'~$

By
Kucera 8 Assoc., Denver, Coo.

Prepared

by H R Shepherd 8

Negative

Scale

1'15840

Aug 1971

J R Hayward

r~i

J

I

W

o
I

Fig. 2. Mosaic of Trickle Mountain bighorn sheep study area.

�-31-

Delineation

of Vegetation

Types

All aerial photographs of the study areas were prepared for vegetationtype mapping and field use in the following manner:
Principal and
coordinate principal points and flight lines were established for each
photo.
To eliminate as much photographic distortion as possible,
effective areas were outlined with red pencil for each photo, using
a method commonly used by the U.S. Forest Service.
If a portion of a
study area boundary occurred in a photo it was shown as a dotted red
line. For easy reference all photos comprlslng a mosaic of a study
area were number coded to show their positional relationships to one
another.
Next, photos were laminated with cold laminating acetate to protect
them from handling, dirt and moisture and to permit erasures of type
line and transect information.
Laminated stereo pairs of photographs were viewed through a mirror
stereoscope, and vegetation types were delineated on the acetate in
black with a Sanford's "Sharpie" marker.
Many different brands of
pencils and pens were tried, but the "Sharpie" was the only one that
would produce a sharp, black line that was waterproof and smearproof
yet erasable on acetate.

Training

Student Crew

A student field crew of four men reported for work June 5. The first
week was a period of intensive training in plant identification, using
the field herbarium of mounted plants collected in the study areas.
The
second week was devoted to training in transecting procedures in the
Winter Range portion of the Trickle Mountain Study Area.

Field Checking

of Vegetation

Type Lines

Before an area of vegetation type delineated on aerial photos was transected,
the location of type boundaries was checked on the ground and, if needed,
type-line boundaries were adjusted on photos.

Transecting

Vegetation

Types

Before vegetation sampling was begun in a vegetation type, the Project
Leader made a cursorial examination of the type, noting kinds, extent and
locations of vegetation stands present.
The type was assigned a number,
and the number was written within the type boundary in black with a "Sharpie"
marker.
In each vegetation stand, transects were located by the Project
Leader to sample parts of the stand most representative of the entire stand.

�-32-

This might require one or many transects, depending on the size and
variability of the stand. When sampling was first begun, an attempt
was made to randomly select sites for transects.
However, this was
abandoned after the first week's work as being impractical, considering
the large size of the area to be sampled, the denseness of the timber
types, the roughness of terrain, and the time constraint imposed of three
months to complete the range inventory of 7,270 acres.
It is recognized
that many American researchers will fault the study because transects
were not located randomly.
However, those more in sympathy with the
European school of vegetation sampling may be more inclined to approve
the subjective method placing more reliance on judgment and experience
and less on blind, mechanical selection of sites to be sampled.
Transects were laid out along a 300-foot steel tape secured between two
steel stakes driven into the ground.
Ends of the transect were marked
with rock cairns.
Transects were run along the contour so that they could
be kept within the same general soil type, and as a consequence, changes
in vegetation composition would be kept to a minimum.
Vegetation was
sampled systematically along the transect every five feet with the
Daubenmire canopy-coverage
frame. When 40 plots had been sampled, speciesarea curves were constructed for species having a canopy coverage of 1 percent or greater.
If more plots were needed for the accuracy desired,
additional plots were sampled, following Daubenmire's suggestions.
After
several types had been sampled, we learned that 60 plots would be sufficient
for the most variable types we would be likely to sample.
So to save time,
we sampled this number of plots as a general rule, testing only for those
types which appeared variable enough to justify the extra work.
The sampling crew of four men worked as teams of two men each.
The composition of teams remained the same through the summer.
However, the men of
a team alternately estimated and recorded data and verified each other's
plant identifications
and canopy-coverage estimates.
On the basis of field testing the point-quadrat frame and the Daubenmire
canopy-coverage
frame from the previous summer, the latter was selected for
use in the range inventory (Fig. 3). Modifications from the frame described
by Daubenmire (1959) were found desirable.
Consequently, a frame was
designed having one end that hinged, permitting placing it around the trunk
of a shrub.
Also, its four pins were made adjustable so that it would rest
securely on steep, rocky slopes.
Quarter sections of the frame edges were
painted contrasting colors and stamped with canopy-coverage percentage
ranges they represented.
Each transect was photographed from one of its ends. A small slate on
which was written with chalk important transect identifying information
usually was propped against a transect stake or cairn. Photos were taken
with an Exakta camera and Kodacolor film.
Both a 3" x 5" color print and
color transparency were made from the color negative.
Prints were later
attached to transect data forms to provide help in the interpretation of
transect data.

�I
Vol
Vol
I

•

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-,

\

-\

,
r

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Fig. J. Daubenmire canopy-coverage frame, modified to permit one end to open and close around the bases of shrubs.

�-34-

The ends of a transect were marked with rock cairns for possible future
reference and its location was marked on an aerial photograph by the
Project Leader at the time he selected the transect site. Using a
rapidigraph pen, a short, red line to scale was marked on the photograph
to indicate the position, length and bearing of the transect.
A numeral
indicating the number of the transect in its type was placed at one end
of the mark designating the transect.
Canopy-coverage
and ground-surface data were recorded by two-man teams,
using the system of Daubenmire (1959). The men of a team would alternately estimate and record canopy coverage, each estimating or recording
for all plots along a transect.
These data were recorded on prepared
forms (Fig. 4). When all of the plots along a transect had been read,
physiognomic data for that transect and its stand in the immediate
vicinity were recorded on a prepared form (Fig. 5). The Braun-Blanquet
system of vegetation analysis and description (Kuchler 1955) was used
to record physiognomic data for each species.
The two men of a team
collaborated in assigning the physiognomic descriptions.
Consequently,
they compared each other's estimates of sociability and coverage, thus
increasing the uniformity and reliability of their estimates.

RESULTS

AND DISCUSSION

Upon termination of transecting, Sept. 8, the crew was brought to the
Research Center where they began the task of compiling and summarizing
the summer's collection of data from 17,434 plots and 292 transects.
For each transect, a form was prepared, summarizing the data for all
canopy-coverage
plots occurring along it (Fig. 6). To the physiognomic
data form was stapled a 3 x 5-in. color photograph, showing the location
of the transect and the vegetation and the characteristics of the ground
surface and topography of the site. All of this information was assembled
into large loose-leaf binders, arranged by vegetation type, for each of
the three study areas (Fig. 7).
The task of relating transect data to acres of vegetation type was begun
but not completed during this segment of the project.
It will be completed during the later part of the next segment devoted to compilation
of data and publication of results.

�-35-

CANOPY-COVERA(;E-PLOTDL\'J'L\

FOltM

Sheet
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Date

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5;/1

)1 J

/~4rn.

ro,./&lt;$

40

_
of

8,,,,v&lt;:
Cover
and
Sociability

CRY/~
ST,tJA J

I
---"------

_

and
Structure

A TeA

No.

=S'---.....:3"--'I'--o

I Physiognomy

Floristic
Composition
(Suecies !_

No.:&lt;

_

_~~g,----=-1-=---o__
Length_IOO
!J CArds No. Plots
G._O'-7- ~:l.
Stand Photo No. 7£ - S ~ Date 7- IS- 72

&amp; Ground Surface
By

I

T~/CkLG ~T£-W/~TL_F_~
__~~A~M_~_E

Study Area. Name
Type No.

No.

Remarks

- ASL7e&lt;..t
J

;.$

J?J:- 80Gf(

:z.
+. /
I.

-

J./

+.1

s/-

-

--

--

--

-

------_._--- 1---------------~L_

--------

----~=____
1___
--

Fig. 5. Physiognomic data field form and sample transect data.

-

�'1HI\.N:&gt;£CT

:jU!',li'iidC{

Pago 1

-37Study Aroa Name
Typo
--(~-----.-

Vog.

Aspect
Phy

s

Slope __

&amp;

&amp;

i.ognomy

Stan'j 110.

.,':u...:0

St.r

uc

7-22
Photo No. -;---1
Tr-ans oc t No. ,------

Soil &amp; Groun~ Surf~co

_

~(.l..:' ,.

7/ 15/7?_, _

Date
2

T[,','iJl

t.ur-e

~_

Cr-ound-Eur f'nc e &amp; Vcreb.tion

Dare -:f
Ground /~

Hock

%

Li t tel'

Point,

S,;timatcs
T Lc hr.n

Total

1150

2775

2075

0

AVe.

19.17

46.25

)4.58

0

Average

No.

of Spec i es y Fl.o t

Vascular
Plant '%

I'-loss

f
f'

4

,)

-

','

Total No. Canopy-Coveraf,e Entries

---------------

0
0

92

1·53
-----------~~----Tot.al
Canopy

Species

Cano py

,A

%

ition

:::overago

A
'0

Cov o r a r-e Oc c ur r-ed

C~;'POS-::]

fiver'age

r.V0rarcc
rotal
Ti:T;cS Fr-equency

.,-C'

GRJ\}iINOIDS
BOGR

992.5

52

86.67

16.54

HU1I;O

1)0.0

12

2.17

-9.07

r,iUFI2

15.0

1

20.00
1.67

0.25

1.05

50.0

5

8.3)

15·0

1

1.67

0.25

1.05

55.0

)

5.00

0.92

J.85

CAEL)
TOTLL GRM-:IKOIDS

.

o.G) I

--+-------JI----L--~--L-I-

--~

FORDS

I

_

BIrr:

,

123.3J

1257.5

.--

0

69.15

'"'
/

17.5

1

2.5

.,

15.0

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•

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I

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Iolal

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.

.

.

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14

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-L __

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Fig. 6. Transect summary form and san.ple canopy -c over-age

92
dat.a,

-.====-__ ==------:-.----1

~~_+___

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2).92

100 Ot_1_]
0

�I

W
00
I

Fig. 7. Loose-leaf binder containing summarized data by transect and vegetation type by range area.

�-39-

BIBLIOGRAPHY
and
LITERATURE

CITED

American Society of Photograrnmetry.
The manual
Amer. Soc. Photograrnmetry.
Menasha, Wis.

of photograrnmetry.
876 p.

Anderson, A. E., D. E. Medin, and D. C. Bowden.
fecal group counts related to site factors
(Unpublished manuscript).

1971. Mule deer
on winter range.

Daubenmire, R.
analysis.

1970.
Sta.

Tech.

1959. A canopy-coverage method of vegetational
Northwest Science.
33(1):43-64.
Steppe vegetation of Washington.
Bull. 62. 131 p.

Wash. Agr. Exp.

Drew, W. B. 1944. Studies on the point-quadrat method of botanical
analysis of mixed pasture vegetation.
J. Agr. Res.
69:289-297.
Fayle, D. C. F. 1959. The point contact method
measure of ground vegetation.
The Forestry

as a three-dimensional
Chronicle.
35(2):

135-141.
Harrington, H. D., and L. W. Durrell.
Sage Books.
203 p.

1957.

Kuchler, A. W. 1955. A comprehensive method
Annals Ass. of Amer. Geog.
65:404-415.

How to identify

of mapping

plants.

vegetation.

Loveless, C. M. 1967. Ecological characteristics
of a mule deer winter
range.
Colo. Game, Fish and Parks Dept., Tech. Pub. No. 20. 124 p.
U. S. Forest Service.
1969.
pretation, Agr. Handbook
U. S. Forest

Prepared

Service.

Forester's guide
No. 308. 40 p.

B.P.I. workbook.

by --'H..:...·~.j:.:o.1..~;.;Jl....&lt;:d..1...1 R...cl..•...••-' S-h"-"
e-(p
•.•.
(:h
'~f'-'r(.",;}
d"'-/
.•...
'....•
'c;.!,'_
Wildlife

Researcher

to aerial

Washington,

D.C.

photo

inter-

47 p.

��July,

-41-

/

/

JOB PROGRESS

State

of

Work

COLORADO
--------~~~~~---No.

W-10l-R-1S

Plan No.

6

Project

Job Title

Period

REPORT

April

Bertram

1, 1972 to March

D. Baker,

LeRoy

Unit

Investigations
1

Job No.
Game Management

Covered:

Personnel:

Game Range

Inventory

31, 1973

C. Carlson

and William

T. McKean

ABSTRACT
Broad descriptive and wildlife species management information is compiled
separately for Wildlife Management Units 61 and 62 (Uncompahgre Plateau).
Subjects discussed included:
Unit Description
(Boundaries, Size, Physical
Features and Climate);
Landownership
Status;
Land Use Status;
Human Populations (by County with projections); Wildlife Species Checklists, Harvests
and Seasons; Narrative and Map Descriptions of Game Species Distribution
and
Abundance;
Introductions of Game Species; Census Areas and Routes; Habitat
Restoration
Projects;
Management Problems Checklist; and Pertinent Research
References by Game Species and Author.

1973

��-43-

GAME MANAGEMENT

UNIT INVENTORY

Bertram D. Baker and William

T. McKean

The Job Progress Report for the previous segment (14) called attention
to a delay in work on this proj'ect due to assignment of both authors to
P~oject FW-lO-R-l.
The special assignment began March 1, 1972; it ended
April 1, 1973 - one year and one month later. Thus, this report summarizes
work planned and very nearly completed under W-lOl-R-14.

P. S. OBJECTIVE
To compile broad descriptive and wildlife
by wildlife management unit.

species management

information

SEGMENT OBJECTIVES
1.

Describe
physical

Wildlife Management Units by name. number, boundaries,
features, and climate.

2.

Inventory by unit the cultural and physical characteristics including
landownership status, land use status, and human population.

3.

Inventory by unit wildlife characteristics including species lists,
distribution and abundance, harvest data, introductions and/or past
species records, census areas and routes, and research and/or management studies bibliographies.

4.

Inventory habitat (range) characteristics of vegetation types, vegetation condition and trend, restoration projects, and transect records,
all from existing sources.

5.

Inventory

6.

Test application qf any item of information compiled or recorded in
steps 1-5 above in pilot computerized map project now in operation.

present and future management

METHODS
Five steps were necessary
Wildlife Uni t :

to produce

problems

size,

by unit.

AND MATERIALS
the desired information

about each

1.

The priorities of units to be studied were determined,
recommendations of field and administrative personnel.

based upon

2.

Working with sources in Fort Collins and Denver, as much data as
possible concerning each item were assembled and condensed (see
APPENDICES A and B).

�-44-

3.

First-draft copies of descriptive material were submitted to
Southwest Regional personnel and to knowledgeable Denver office
persons for review and correction.

4.

Group meetings were arranged with field personnel and others to
make pencilled corrections on 1/2"/mile species distribution maps
and to make refinements to the narrative.

5.

Final revisions to narratives and maps were made and all information was gathered into file binder folders, except for the 1/2"/
mile maps which were reproduced by hand and bound in separate
folders.
Copies of all this material were distributed to the
Southwest Regional office, the Denver office (Game Management
Section), and to the Federal Aid Section.

RESULTS AND DISCUSSION

For the work done on T.VildlifeHanagement Units 61 and 62 final corrections,
collations, and distributions were made as indicated in step five above.
Complete copies of the information accumulated are attached as two
appendices, except for maps of game animal distribution.
APPENDIX A
contains data for Wildlife Management Unit 61; APPENDIX B contains data
for Wildlife Management Unit 62.
No information was obtained concerning habitat (range) characteristics
of
vegetative types, nor about vegetative condition and trend (objective four).
Data gathered under this project should be useful in the pilot computerized
map project now in operation by the Division Planning Section (objective
six), but this has not been realized as yet.

Prepared

by

1f:c tt34,')-'J'l..-}( 2}i' C:/(Jla_1L-O
William T. McKean
Wildlife Researcher

�APPENDIX

A

HILDLIFE MANAGEMENT UNIT 61

nol. UNCOMPAHGRE)

Mesa, Montrose, and San Miguel Counties, Colorado

Information on Unit Description,
Landownership, Land Use, Human
Population; Wildlife Species
Checklists, Harvests, Seasons,
Narrative and Map Descriptions
of Distribution and Abundance;
and Other Allied Data. To:
March 15, 1972.
Compiled by:

W. T. McKean and
B. D. Baker

�lA

DESCRIPTION

. 2/,
B oun d arles-

- WILDLIFE

HANAGEMENT

3/ "hT at portlon
.
-.--

0f

UNIT 61 (W. UNCOHPAHGRE)];./

Mesa County east

0f

th e Dol ores River,

south of State Highway

No. 141 from the town of Gateway to the Uncompahgre

Plateau

the Gunnison

divide between

of MOntrose
Highway

River and Dolores River; that portion

County east of the Dolores

No. 90 from its intersection

section with State Highway

River and north

and east of State

with the Dolores

River to its inter-

No. 145, and north and east of State Highway

No. 145 and south and west of the Uncompahgre
of San Miguel

Size.--Unit
in size.

County north of State Highway

61 was determined
A planimeter

scale Uncompahgre

Physical

River near Uravan,

to be 1,067.7 square miles or 683,272 acres
from a 1/4" = 1 mile

Forest map.

exception

Redvale,

divide; that portion

No. 145."

was used to obtain the acreage

National

Features.--With

Plateau

of included

and Norwood,

areas south of the San Miguel

Unit 61 approximates

the southwestern

1/ Legally termed "Big Game Hanagement Unit" (see footnote 2/ below) but
tentatively called ''Wildlife Management Unit" here to include broader aspects
of animal life present.
~/ Colorado Game, Fish and Parks Division
(p. 12, Chap. 2 - Big Game).

3/ As of Harch 23, 1972, Commission

Laws and Regulations

Hdbk.,

1970

action changed the boundaries of Unit
Involved was a transfer of about 2 to 2~ townships of land to Unit 70,
with the area lost being territory enclosed by the Dolores and San Higuel
Rivers and State Highway 90. Inasmuch as voluminous mapwork
and all tabular
and narrative data had been compiled using boundaries observed here, it was
decided not to revise data to correspond with new houndaries.

61.

�2A

one-half

of the Uncompahgre

Plateau.

The plateau

is a broad structural

uplift that lies within

the Canyon Lands section of the Colorado Plateaus

physiographic

Oldest rocks are Precambrian

exposed

province.

in wes t e rn Unaweep

whe ra in the northern

Canyon and generally

portion of the unit.

stones and Mesozoic

sandstones,

siltstones,

crop out variously

over remaining

crystalline

rocks

at higher elevations

Paleozoic

else-

gray shales and lime-

and shales of several

formations

parts of Unit 61 (U. S. Dept. of Agricul-

ture 1971).

Much of the west side of the plateau
and steeply

sloping mesas.

sloping broad valleys
near Redvale

Minimum

Contrasting

with those features are gently

such as East Paradox

and Norwood

elevation

consists of deep canyons, sharp cliffs,

and gently rolling tablelands

(U. S. Dept. of Agriculture

1971).

is about 4,570 feet where State Highway No. 141 inter-

cepts the Dolores River near Gateway.

Maximum elevations

plateau

divide and range from about 6,900 feet in Unaweep

10,338

feet on Horsefly

are found on the
Canyon up to

Peak and back down to near 8,900 feet at-Dallas

Divide.

Most of the plateau

generally

drop below

crest is above 9,000 feet, but elevations

that figure within short distances

southwestwardly

from the divide.

Unit 61's two principal

streams

are the northwestwardly

and San Miguel

Rivers;

they lie near,

unit's southern

and western

edges.

flowing Dolores

form, or help form boundaries

North to south, in order, and flowing

south and west to the rivers are the following main tributaries:
Mes a, Tabeguache,

Horsefly

at the

and Leopard

Creeks.

Besides

West,

many small stock

�3A

ponds,

the unit has Casto Reservoir

surface

acres at full capacity.

Both detailed
consulting
Norwood

and more general soils information

U. S. Soil Conservation

office or examining

report cited previously

Climate.--Climate

44

0

Service Work Unit personnel

at their

of Agriculture

(1971)

herein.

of Unit 61 is variable

due to differing

climate, having

o

recorded

topography.

Gate-

a mean annual temp0

of about 54 F and average

annual growing season of 194 days (28 F

Uravan and Norwood

freeze threshold).

about 40

can be obtained by

the U. S. Department

way lSW perhaps has the mildest
erature

on Gill Creek that contains

have mean temperatures

of 53.5

o

F and
0

F and growing seasons of about 180 days and 133 days, respectively

F freeze threshold).

Growing season lengths scale downward with increase

in elevation

to an estimated

Precipitation

is variable,

30 days or less on the Uncompahgre

too.

Plateau

Nine inches or less in East Paradox

is the least mean annual amount reported
inches ~ Uravan

12 inches, Redvale

for Unit 61.

15 inches, Norwood

Gateway

15+ inches,

25 inches extends

top.

of Unit 61 receives

extreme

from 12 to 16 inches,

to 20-inch zone.

Norwood

growing season.

Short cloudbursts

However,

20 and

the greatest

followed next by the 16-

gets about 55 percent of its precipitation
characterize

Valley

and Placer-

A narrow belt of between

over the plateau's

crest.

receives 11

ville 14 inches mean annual precipitation.

portion

(28

summer

rainfall

in the

at lower

elevations.

Disc ussions in preceding
ment of Agriculture

paragraphs

were based upcn data f rorn U. S. Dep ar t=-

(1971) and U. S. Department

of Commerce

(Continuing)

�4A

reports.

Besides

Unit 61, possible
Service

the information

other useful records can be obtained

cooperating

just outside

given for those stations located within

stations

at Paradox

U. S. Department

of Agriculture.

Service,

and related land resources,

and Utah.

Coop. Study Rep. of Colo-

of Commerce.

Service.

February.

(Continuing).

and Annual Summaries.

Asheville,

Water

Board and U.S.D.A. Econ , Res. Service,

and Soil Conserv.

U. S. Department

Cited

1971.

River Basin in Colorado

rado Water Conserv.

Monthly

which are located

of the unit.

Literature

Dolores

and Telluride

for National Weather

National

Denver.

Climatological
Climatic

N. C. 28801.

B. D. Baker
February 1972

Forest

(processed).

data, Colorado.

Center, Fed. Bldg.,

�SA

LANDmmERSHIP

STATUS - WILDLIFE MANAGEMENT UNIT 61

Bureau of Land Management

222,176

Acres

Forest Service

283,803

Acres

Private lands excluding lands within
National Forest boundaries

155,576

Acres

Private lands within National Forest
boundaries

19,477

Acres

Colorado Division Game, Fish and Parks

o Acres

Other Federal lands including Park
Service, Indian and Military Reservations, and Bur. Sport Fisheries and
Wildlife

0

Acres

State Land Board Administration lands

640

Acres

Municipal lands

801

Acres

Total

682,473

Acres *

(1066.4 sq. miles)

* Total land acreage differs from total in Unit Description Section
(683,272 A. or 1067.7 sq. miles). Forest Service lands and private
lands within the National Forest were derived by planimetering on
2" = 1 mile Uncompahgre National Forest township plat maps. B.L.M.
lands, Game, Fish and Parks lands and other private land acreages
resulted from planimetering and estimating on 1/2" = 1 mile Uncompahgre
National Forest map. Municipal lands include Nucla and those portions
of Naturita and Norwood within Unit 61.

W. T. McKean and L. C. Carlson
September 1971

�6A

LAND USE STATUS - HILDLIFE MANAGEMENT UNIT 61

Irrigated croplands

21,698

Acres

Non-irrigated croplands

1,599

Acres

Grasslands

26,118

Acres

Sagebrush

75,030

Acres

Brush

110,643

Acres

Woodland - pinyon-juniper,
oakbrush

304,524

Acres

Aspen

21,418

Acres

Corrnnercialtimber

112,947

Acres

River bottom vegetation

1,489

Acres

Miscellaneous

7,007

Acres
,J

Total

682,473

Acres *

(1066.4 sq. miles)

* Cover type categories and acreages were derived from planimetering
on 1/2" = 1 mile S.C.S. Land Use and Cover Type Maps, 1954, for Mesa,
Montrose, and San Miguel counties. Computations were 7% over landownership status total acreage. An adjustment was made to each
category so that the total would conform to the landownership figure.

L. C. Carlson
November 1971

�7A

HUMAN POPULATION

- WILDLIFE

Inasmuch

as Unit 61 involves

counties

and unit bounding

Norwood,

determination

difficult.

Instead,

are presented

spite being
involved

portions

highways

UNIT 61

of Mesa, Montrose,

and San Miguel

split the towns of Naturita

of specifically

applicable

populations

and

would be

rural and urban data and trends for 1960 and 1970

in total for Montrose
(Table 1).

municipalities

MANAGEMENT

comparatively

and San Miguel

counties

Mesa County data are excluded
large in size, that portion

in Unit 61 contains

an indeterminable,

and their

because,

de-

of the county

very small and entirely

rural, human population.

This analysis
sources,

differs

from those of the Bureau of the Census and other

since they consider

lation as being

rural in character.

towns are considered
populations

cities and towns of less than 2,500 popu-

being

as being urban regardless

very slightly

third between

cities and

of size, with remainder

rural.

Data shown on Table 1 indicate
increased

Here, all incorporated

the years

that the population

and that of San Miguel
1960 and 1970.

increased

moderately,

Montrose.

Other municipalities

of Montrose

County

County dropped by one-

Montrose

County urban population

being carried by the strong increase

for the city of

in the two counties varied both ways but

were generally

downward

in populations.

lation showed

a rather sharp decline

of about 19%.

both Montrose

and San Miguel

dropped noticeably,

counties

The San Miguel

County urban popu-

Rural populations

of

with the latter

�SA

declining

793 people

or about 44%.

lations in Mesa, Hontrose,

Additional

and San Miguel

detailed

analyses of popu-

counties can be found by con-

suIting pages 4-5 through 4-8 of the f o Ll.ow i ng report:
U. S. Department

of Agriculture.

1971.

Water and related

land resources,

Dolores River Basin in Colorado

and Utah.

Coop. Study Rep. of Colorado Water Conserve Board and U.S.D.A.
Econ. Res. Service,

Forest Service,

February.

(processed).

Denver.

and Soil Conserve

Table 1. Populations and trends of counties and municipalities
part, w i.t.h Unit 61 1/.
Population
County

1960

1970

Change

Population
Inc. Cities
%
and
Towns
1960 1970 Change

Hontrose

18,286 18,366

+0.5

Montrose

5,044

6,413

+27.1

Naturita

979

820

-16.1

Nucla

906

949

+ 4.7

Olathe

773

756

- 2.2

Totals

7,702

8,938

+16.0

%

-----------------------------------------------------San Hf.gue L

1/ Sources

2,944

1,949 -33.8

involved,

10,584 9,428

Telluride

677

510

-24.7

Norwood

443

408

- 7.9

Totals

1,120

918

-18.7

(urban population).

-10.9

-------------------------------~

is based upon total county population

cities and towns

in

Rural Population 2/
%
1960 1970 Change

1,824 1,031

are the U.S. Bureau of the Census and Colorado Yearbook,

2/ County rural population
incorporated

Service.

1959-61.

less total of

-43.5

�9A

Starting

with a population

also gives advanced

of 10,422 in 1960, the aforementioned

population

projections

to be:

1980 - 10,800;

2000 - 12,300;

Colorado

State Planning

Office

for Montrose
Similarly,

San Miguel

specific

in indicating

2020 - 13,700.

River Basin

Estimates

by the

(pers. comm. 1971) placed populations

County at 18,500 in 1980 and 20,000 by 2010 and thereafter.

Both of these sources
being

for the Dolores

report

County's

population

was given as 2,000 "forevermore".

for county and basin projections

for Unit 61, but the projections
that generally

are far from

are fairly consistent

light gains will occur within

the next 40

to 50 years in and near the unit.

B. D. Baker
July 1971

�lOA

GAME SPECIES

Big game mammals

- WILDLIFE

t£~AGEMENT

UNIT 61 (W. UNCOMPAHGRE)

!I

Black bear (Ursus americanus)
Elk (Cervus canadensis)
Mountain lion (Felis concolor)
Mule deer (Odocoileus hemionus)

Small game mammals

11

Cottontail rabbit (Sylvilagus audubonii; .§..~nuttallii)
Pine (red) squirrel (Tamiasciurus'hudsonicus)
Snowshoe hare (Lepusamericartus)
Abert's squirrel (Sciurus aberti)

Game birds Jj
Migratory

waterfowl

and shorebirds

Great Basin Canada goose'(Branta canadensis moffitti)
Black brant (Brantanigricans)
11
White-fronted goose (Anser albifronsfrontalis)
11
Snow goose (Chen hyperborea hyperhorea)
Mallard (Anas platvrhynchos platyrhyllchos)
Gadwall. (Anas strepera)
,
Pintail (Anas acuta)
Green-winged teal (Anas carolinensis)
Blue-winged teal (Anas discors discors)
Cinnamon teal (Ana~anoptera
septentrionalium)
American widgeon (Mareca amez Lcana)
Shoveler (Spa,tula clypeata)
Wood duck (Aix sponsa)
Redhead (Aythya americana)
Ring-necked duck (Aythya collaris)
Canvasback CAythya valisineria)
Greater scaup (Aythya marila nearctica) 11
Lesser scaup (Aythya affinis)
Common goldeneye (Bucephala clangula americana)
Barrow's golden-eye (Bucephala islandica) 11

_:!/ Nomenclature
Colorado.

according to Lechleitner, R. R. 1969.
Pruett Publishing Co., Boulder.
254 pp.

Wild mammals

-~/Nomenclature according to Bailey, A. E", and R. J. Neidrach.
Pictorial checklist of Colorado birds.
Denver Mus. Nat. Rist.
llunverified in hunter's
game in 1970-71.

bag checks but possible

(

rare migrant

of

1967.
168 pp.
and legal

�llA

Game birds - Migratory

waterfowl

and shorebirds

(continued)

Bufflehead (Bucephala albeola)
Ruddy duck (Oxyura j amaic.ensis rubida)
Hooded merganser (Lophodytes cucullatus)
Common merganser (Mergus merganser americanus)
Red-breasted merganser (Mergus serrator serrator)
Common snipe (Capella gallinago delicata)
American coot (Fulica americana americana)

Upland

game birds

Blue grouse (Dendragapus obscurus obscurus)
Sage grouse (Centrocercus urophasianus urophasianus)
Ring-necked pheasant (Phasianus colchicus)
Chukar (Alectoris graeca)
.
Band-tailed pigeon (Columba fasciata fasciata)
Mourning dove (Zenaidura macroura marginella)
Sharp-tailed grouse (Pedioecetes phasianellus columbianus)
Mountain quail (Oreortyxpicta)
Wild turkey (Meleagris gallopavo merriami)

T. McKean
November 1971

W.

�12A

OTHER MAMMALIAN SPECIES 1/ - VlILDLIFE MANAGEMENT UNIT 61 CW. UNCOMPAHGRE)

Furbearers ]j
Beaver (Castor canadensis)
Muskrat (Ondatra zibethicus)
Ringtail (Bassariscus astutus)
Weasels (Mustela erminea; M.ftenata)
Mink (Mustela vison)
Marten (Hartes amer'Lcana)
Nongame mammals 1/
White-tailed jack rabbit (Lepustownsendii)
Black-tailed jack rabbit (Lepus"califomicus)
Yellow-bellied marmot (Marmota flaviventtis)
White-tailed prairie dog (Cynomys leucutus)
Gunnison's prairie dog (Cynomysgunnisoni)
Rock squirrel (Spermophilusvatiegatus)
White-tailed antelope squirrel (Ammospermophilus leucurus)
Golden-mantled ground squirrel (Spermophilus lateralis)
Least chipmunk (Eutamias minimllS)
Colorado chipmunk (Eutamias"quadrivittatus)
Coyote (Canis latrans)
Red fox (Vulpes fulva)
Gray fox (Urocyon cinereoargenteus)
Kit (swift) fox (Vulpes velox)
Raccoon (Procyon lotor)
American badger (Taxidea taxus)
Spotted skunk (Spilogale putorius)
Striped skunk (Mephitis mephitis)
Bobcat (Lynx rufus)
Porcupine (Erethizon dorsatum)
1/ These species, grouped separately as "Furbearers" and "Nongame mammals"
and outside of "game" categories, follow Chapter 62, Colo. Rev. Statutes
1962 As Amended, in Colo. Game, Fish and Parks Div. Laws and Regulations
Hdbk., 1970 (Art. 1, Item 3, Definitions, p. 327).

1/ Nomenclature according to Lechleitner, R. R.

1969.

Colorado.

254

Pruett Publishing Co., Boulder, Colo.

Wild mammals of
pp.

W. T. McKean
November 1971

�13A

OTHER AVIAN SPECIES

Nongame

birds

1/ - WILDLIFE

MANAGEMENT

UNITS 61 AND 62

1/

Common loon (Gavia immer) Rare migrant.
Horned grebe (Podiceps auritus cornutus) Rare migrant.
Eared grebe (PodiceDs caspicus californicus) Uncommon migrant.
Western grebe (Aechmophorus occidentalis) Possible rare migrant.
Pied-billed grebe (Podilymbus podiceps podiceps) Uncommon migrant 1/ and
possible rare summer resident.
Double-crested
cormorant (Phalacrocorax auritus auritus) Rare
migrant.
Great blue heron (Ardea herodias treganzai) Common summer resident and few
in winter 1/, !!../.
Snowy egret (Leucophoyx thula brewsteri) Uncommon summer resident.
Black-crowned night heron (Nycticorax nyctic0rax hoactli) Uncommon summer
resident.
Least bittern (Ixobrychus exilis exilis) Possible accidental summer vlsitor.
American bittern (Botaurus lentiginosus) Possible rare migrant.
White-faced ibis (Plegadis chihi) Possible rare migrant.
Whistling swan (Olor columbianus) Rare migrant.
Sandhill crane (Grus canadensis canadensis; G. c. tabida) Common migrant.
Virginia rail (R~s
limicola limicola) Pos~ible uncommon summer resident.
Sora (Porzana carolina) Possible uncommon SUmmer resident.
Semipalmated plover (Charadrius semipalmatus) Possible rare migrant.
Snot .•
:ry plover (Charadrius alexandrinus nivosus) Possible accidental migrant.
Killdeer (CharadriliS--vQc-iferus ~ociferus)- -C-;~on summer and uncommon winter
resident 3/.
Mountain plover (Eupoda montana) Possible accidental migrant.
Black-bellied plover (Sguatarola sguatarola) Possible rare migrant.

1/ These species, grouped separately as "Nongame birds" and "Raptores" and
-;-utside of "game" categories, follow Chapter 62, Colo. Rev. Statutes 1963 As
Amended, in Colo. Game, Fish and Parks Div. Laws and Regulations Hdbk., 1970.
(Art. 1, item 3, Definitions, p. 327).

1/ Nomenclature

from Bailey, A. M., and R. J. Neidrach.
19~5. Birds of Colorado.
Denver Mus. Nat. Hist. 2 Vols. 895 pp. Information on occurrence and method for
reporting same are adapted from the foregoing reference and Davis, W. A. 1969.
Birds in western Colorado.
Colo. Field Ornith. 61 pp., with the system of Davis
(1969:6) favored and followed closely throughout.
Presence of term "possible"
is an author judgement connotating that the species could occur in the units
some time even though evidence is presently lacking that it does.
Absence of
term means that evidence by Bailey and Neidrach (1964) and/or Davis (1969) makes
occurrence a near certainty or that sighting(s) have been reported verbally by
anyone
or more of Division personnel H. Burdick, T. Quick, L. Denton
[and others,
as appropriate] or qualified by additional footnotes that follow.

1/ Unpublished

checklist

of birds of the Uncompahgre

Plateau

by Dr. A. S. Hyde,

1971.

4/ Sight record of A. E. Anderson as given in APPENDIX A - PARTIAL CHECKLIST OF
VERTEBRATE FAUNA OBSERVED IN VICINITY OF STUDY AREA, pp. 59-60, of Anderson, A. E.
1960. Effects of sagebrush eradication by chemical means on deer and related wildlife.
Fed. Aid Job Compo Rep. Proj. W-38-R-13, July, Colo. Dept. Game and Fish.
72 pp , (processed).

�Nongame

birds

(continued)

Long-billed curlew (Numenius americanus americanus) Possible rare migrant.
Spotted sandpiper (Actitis macularia) Common Summer resident 2/.
Solitary sandpiper (Tringa solitaria cinnamomea) Common migrant and
occasional summer visitor.
Willet (Catoptrophorus semipalmatus inornatus) Rare migrant.
Greater yellowlegs (Totanus melanoleucus) Possible common migrant.
Lesser yellowlegs (Totanus flavipes) Possible uncommon migrant.
Knot (Calidris canutus rufa) Possible accidental migrant.
Pectoral sandpiper CErolia melanotos) Possible rare migrant.
Baird's sandpiper (Erolia bairdii) Possible uncommon migrant.
Least sandpiper (Erolia minutilla) Possible common migrant.
Long-billed dowitcher (Limnodromus scolopaceus) Possible rare migrant.
Semipalmated sandpiper (Ereunetes pusillus) Rare migrant 2/.
Western sandpiper (Ereunetes mauri) Possible uncommon migrant.
Marbled godwit (Limosa fedoa) Possible rare spring migrant.
Sanderling (Crocethia alba) Possible accidental migrant.
American avocet (Recurvirostra americana) Rare migrant.
Black-necked stilt (Himantopus mexicanus) Possible rare migrant.
Wilson's phalarope (Steganopus tricolor) Common migrant and uncommop
Summer res ident .
.
Northern phalarope (Lobipes lobatus) Possible rare migrant.
Herring gull (Larus argentatus smithsonianus) Possible uncommon migrant.
California gull (Larus californicus) Possible rare migrant.
Ring-billed gull (Larus delawarensis) Uncommon migrant.
Franklin's gull (Larus pipixcan) Cornmon migrant.
Bonaparte's gull (Larus philadelphia) Possible rare migrant.
Forster's tern (Sterna forsteri) Possible rare migrant.
Common tern (Sterna hirundo hirundo) Possible accidental migrant.
Leas i, i.er n (SLeL"UCl a.lui.L~cuH6 d.ti!cild65(5)
Possible: acc i.dent.a Lrni.grunt •
Black tern (Chlidonias niger surinamensis) Possible rare migrant.
Rock dove (Columba livia) Common resident.
Yellow-billed cuckoo (COCCYZU3 americanus americanus) Possible uncommon
summer resident.
Poor-will (Phalaenoptilus nuttallii nuttallii) Common summer resident 4/.
Cornmon nighthawk (Chordeiles minor powelli ; ~. ~. henryi ) Common
Summer resident if.
Black swift (Cypseloides niger borealis) Possible accidental Summer migrant.
White-throated
swift (Aeronautes saxatalis sclateri) Common Summer
resident 2/, if.
Black-chinned hummingbird (Archilochus alexandri) Common summer resident.
Broad-tailed hummingbird (Selasphorus platycercus platycercus) Common
summer resident 2/.
Rufous hummingbird (Selasphorus rufus) Common late Summer and fall migrant.
Calliope hummingbird (Stellula calliope) Rare Summer migrant i/and Possible
rare Summer resident.
Rivoli's hummingbird (Eugenes fulgens aureoviridis) Possible rare summer
visitor.
Belted kingfisher (Megacerle alcyon alcyon) Common resident 3/.
Yellow-shafted
flicker (Colaptes auratus luteus) Possible accidental
migrant.

: \'

�15A

Nongame

birds

(continued)

Red-shafted flicker (Colaptes cafer collaris) Common resident 3/,4/.
Lewis' woodpecker (Asyndesmus lewis) Common resident 11.
-Yellow-bellied
sapsucker (Sphyrapicus varius muchalis) Common summer
resident.
.
Williamson's sapsucker (Sphyrapicus thyroideus nataliae) Common summer
resident.
Hairy woodpecker
(Dendrocopos villosus monticola) Uncommon resident i/.
Downy woodpecker
(Dendrocopos pubescens leucurus) Uncommon resident.
Northern three-toed woodpecker
(Picoides tridactylus dorsalis) Rare resident.
Eastern kingbird (Tyrannus tyrannus) Uncommon summer resident 11.
Western kingbird (Tyrannusverticalis)
Common summer resident 1/·
Cassin's kingbird (Tyrannus vociferans vociferans) Possible uncommon
summer resident.
Ash-thro?ted flycatcher (Myiarchus cinerascens cinerascens) Common summer
resident.
Say's phoebe (Sayornis saya saya) Common summer resident.
Traill's flycatcher (Empidonax traillii) Uncommon summer resident 11.
Hammond's flycatcher (Empidonax hammondii) Uncommon summer resident.
Dusky flycatcher (Empidonax oberholseri) Uncommon summer resident.
Gray flycatcher (Emphidonax wrightii) Common summer resident.
Western flycatcher (Empidonax difficilis hellmayri) Common summer resident.
Western wood peewee (Contopus sordidulus veliei) Common summer resident i/.
Olive-sided flycatcher (Nuttallornis borealis) Uncommon summer resident.
l''T
..l
1.1.ULllCU

'1
1_
..LC1.Lr\.

/l.'
L...!1 __
\.1.:J.LCWUPL1..L..La.

1 __
CL..Lpc:.v

....•
"-_...:
l.-J....1..Ct

1

__
-::-::'

_,..,\

..L.\-,..U,-V.J..CLL.J.UQ./

~ ...........•.•.•...•....
__
o
..;,:t"_r
VLlllllUoV.LL

");/

.L~"""..L.'-'-_"'''''''' .:::...' •

Violet-green swallow (Tachycineta thalassina lepid~) Common summer
resident 3/, 4/.
Tree swallow-(Irfdiprocne
bicolor) Common summer resident 3/.
Bank swallow (Riparia riparia riparia) Uncommon migrant and summer resident.
Rough-winged swallow (Stelgidopteryx ruficollis serripennis) Uncommon
migrant and summer resident.
Barn swallow (Hirundo rustica erythrogaster)
Common migrant and summer
resident 3/.
Cliff swallow (Petrochelidon pyrrhonota pyrrhonota) Common summer resident.
Purple martin (Progne subis subis) Possible accidental summer migrant.
Gray jay (Perisoreus canadensis capitalis) Common resident.
Steller's jay (Cyanocitta stelleri macrolopha) Common resident 1/, i/.
Scrub jay (Aphelocoma coerulescens woodhouseii) Common resident 11.
Black-billed magpie (Pica pica hudsonia) Common resident 11, i/.
Common raven (Corvus corax sinuatus) Common resident 11, i/.
Common crow (Corvus brachyrhynchos brachyrhynchos)
Common resident 1/·
Pinyon jay (Gymnorhinus cyanocephalus) Common resident 11, i/.
Clark's nutcracker
(Nucifraga columbiana) Common resident i/.
Black-capped chickadee (Parus atricapillus garrinus) Common resident 4/.
Mountain chickadee (Parus gambeli gambeli) Common resident
Plain titmouse (Parus inornatus r Ldgway i ) Common resident.
Common bushtit (Psaltriparus minimus plumbeus) Common resident 11, i/.
White-breasted
nuthatch (Sitta carolinensis nelsoni) Uncommon resident i/.

i/.

�16A
Nongame

birds

(continued)

Red-breasted nuthatch (Sittacanadensis)
Rare resident.
Pygmy nuthatch (Sitta pygmaea melanotis) Common resident ii/.
Brown creeper. (Certhia familiaris montana) Uncommon resident.
Dipper (Cinclus mexicanus unicolor) Common resident.
House wren (Troglodytes aedon parkmanii) Common summer resident 3/., 4/.
Winter wren (Troglodytes troglodytes pacificus) Status unknown iil. Bewick's wren (Thryomanes bewickii eremophilus) Common summer and rare
winter resident 1/.
Long-billed marsh wren (Telmatodytes palustris plesius) Uncommon summer
resident 3/.
.
Canyon wren (Salpinctes mexicanus conspersus) Uncommon summer 1/ and
possible rare .winter resident.
.
Rock wren (Salpinctes obsoletus obsoletus) Common.summer and possibl~
rare winter resident.
Mockingblrd (Mimus polyglottos leucopterus) Uncommon·resident.
Catbird (Dumetella carolinensis) Possible rare summer resident.
Brown thrasher (Toxostoma·rufum
longicauda) Possible accidental migrant.
Sage thrasher (Oreoscoptes montanus) Common summer resident ~/.
Robin (Turdus migratorius propinguus) Common resident 1/, ii/.
Hermit thrush (Hylocichla guttata auduboni)
Common summer resident.
Swainson's thrush (Hylocichla ustulata almae) Possible common migrant
and rare summer resident.
Veery (Hylocichla fuscescens salicicola) Common summer resident.
Western bluebird (Sialia mexicana bairdi) Common migrant, uncommon
. summer resident ii/, and possible rare winter resident.
Mountain bluebird (Sialia currucoides) Common migrant and summer resident
1/, ~/, and possible winter resident •
.Townsend's solitaire (Myadestes townsendi townsendi) Uncommon resident 4/.
Blue-gray gnatcatcher (Polioptila caerulea amoenissima) Common summer
resident 1/.
Golden-crowned kinglet (Regulus satrapa amoenus) Possible common summer
and uncommon winter resident.
Ruby-crowned kinglet (Regulus calendula cineraceus) Common migrant 1/ and
summer resident 4/ and possible accidental winter resident.
Water pipit (Anthus-spinoletta
alticola) Common migrant 1/ 'and possible
winter resident.
Bohemian waxwing (Bombycilla garrulus pallidiceps) Common winter migrant.
Cedar waxwing (Bombycilla cedrorum) Irregular visitor.
Northern shrike (Lanius excubitor invictus) Common winter resident 3/.
Loggerhead shrike (Lanius ludovicianus excubitorides) Uncommon summer
and common winter resident 3/.
Starling (Sturnus vulgaris vulgaris) Common resident 1/.
Gray vireo (Vireo vicinior) Uncommon summer resident.
Solitary vireo (Vireo solitarius plumbeus) Common summer resident 1/.
Red-eyed vireo (Vireo olivaceus) Possible rare summer resident.
Warbling vireo (Vireo gilvus swainsonii) Common summer resident.
Tennessee warbler (Vermivora peregrina) Possible rare migrant.
Orange-crowned warbler (Vermivora celata orestera) Possible uncommon
migrant and summer resident.
Nashville warbler (Vermivora ruficapilla rid~]ayi) Possible rare migrant.
Virginia's warbler (Vermivora virginiae) Common summer resident.
Yellow warbler (Dendroica petechia aestiva) Common summer resident 1/.
Myrtle warbler (Dendroica coronata coronata) Uncommon migrant.

;

F

�17A
Nongame birds

(continued)

Audubon's warbler (Dendroica auduboni memorabilis) Common summer
resident 1/, ~/ and possible rare winter resident.
Black-throated gray warbler (Dendroica nigrescens) Common summer
resident 3/.
Townsend's w;rbler (Dendroica townsendi) Uncommon migrant 4/.
Grace's warbler (Dendroica graciae graciae) Rare summer re;ident ~/.
Northern waterthrush (Seiurus noveboracensis notabilis) Rare migrant 1/.
MacGillivray's warbler (Opor~rnis tolmiei monticola) Common migrant and
uncommon summer resident 3/.
Yellowthroat (Geothlypis tri~has occidentalis; G. t. campicola) Uncommon
summer resident 1/.
- Yellow-breasted
chat (Icteria virens auricollis) Common summer resident.
Wilson's warbler (Wilsonia pusilla pileolata) Common migrant and possible
summer resident.
American redstart (Setophaga ruticilla tricolora) Possible rare migrant.
House sparrow (Passer domesticus domesticus) Common resident 3/.
Bobolink (Dolichonyx oryzivorus) Possible rare summer migrant~
Western meadowlark (Sturnella neglecta neglect a) Common summer 1/ and
uncommon winter resident.
Yellow-headed blackbird (Xanthocephalus xanthocephalus) Common summer
resident 3/.
Red-winged blackbird (Agelaius phoeniceus fortis) Common resident 1/.
Orchard oriole (Icterus spurius) Possible accidental summer visitor.
Bullock's oriole (Icterus bullockii bullockii) Common summer resident 1/.
Rusty blackbird (Euphagus carolinus carolinus) Possible late winter visitor.
Brewer's blackbird (Euphagus cyanocephalus) Common resident 1/.
Br own=he aded cowb i rd (:iVlolotnrus
ater ar t em'i.s i.ae ) Common summe r resLd en t ;
Western tanager (Piranga ludovic~)
Common migrant and summer resident ~/.
Scarlet tanager (Piranga olivacea) Possible accidental summer migrant.
Rose-breasted grosbeak (Pheucticus ludovicianus) Possible accidental spring
migrant.
Black-headed grosbeak (Pheucticus melanocephalus melanocephalus)
Common
summer resident.
Blue grosbeak (Guiraca caerulea interfusa) Uncommon summer resident 1/.
Lazuli bunting (Passerina amoena) Uncommon summer resident.
Dickcissel (Spiza americana) Possible accidental migrant.
Evening grosbeak (Hesperiphona vespertina brooks i) Irregular resident.
Cassin's finch (Carpodacus cassinii) Common resident.
House finch (Carpodacus mexicanus frontalis) Common summer and uncommon
winter resident 1/.
Pine grosbeak (Pinicola enucleator montana) Uncommo~ resident.
Gray-crowned rosy finch (Leucosticte tephrocotis tephrocotis; ~. t.
littoralis) Common winter migrant.
Black rosy finch (Leucosticte atrata) Uncommon winter migrant.
Brown-capped rosy fincD (Leucosticte australis) Common winter resident.
Common redpoll (Acanthis flammea flammea) Possible accidental winter migrant.
Pine siskin (Spinus pinus pinus) Common resident ~/.
American goldfinch (Spinus tristis tristis; ~. ~. pallidus) Common summer
1/ and possible uncommon winter resident.
Lesser goldfinch (Spinus psaltria psaltria) Possible uncommon summer
resident" and accidental winter migrant.
Red crossbill (Loxia curvirostra) Irregular visitor.
White-winged crossbill (Loxia leucoptera leucoptera) Possible accidental
winter migrant.

�18A
Non-game

birds

(continued)

Green-tailed towhee (Chlorura chlorura) Common Summer resident 11, ~/ and
possible rare winter resident.
Rufous-sided towhee (Pipilo erythrophthalmus montanus) Uncommon resident 1/.
Lark bunting (Calamospiza melanocorys) Possible uncommon summer resident.
Savannah sparrow (Passerculus sandwichensis nevadensis; f. ~. anthinus)
Possible uncommon migrant and Summer resident.
Grasshopper sparrow (Ammodramus savannarum perpallidus) Uncommon Summer
resident 5/.
LeConte's sp;rrow (Passerherbulus caudacutus) Possible accidental migrant.
Sharp-tailed sparrow (Ammospiza caudacuta nelsoni) Possible accidental
migrant.
Vesper sparrow (Pooecetes gramineus confinis) Common migrant and summer
res ident 11, ~/.
Lark sparrow (Chondestes grammacus strigatus) Common migrant and summer
resident 1/.
Black-throated
sparrow (Amphispiza bilineata deserticola) Common summer
res ident.
Sage sparrow (Amphispiza belli nevaden3is) Common summer resident.
White-winged junco (Junco aikeni) Possible accidental winter visitor.
Slate-colored junco (Junco hyemalis hyemalis;
cismontanus) Possible
rare winter resident.
Oregon junco (Junco oreganus oreganus; L. Q. mearnsi) Common winter
resident 3/.
Gray-headed junco (Junco caniceps caniceps) Common resident ~/.
Tree sparrow (Spizella arborea ochracea) Uncommon winter visitor 1/.
Chipping sparrow (Spizella passerina boreophila) Common summer resident 1/, ~/.
Brewer's spar-row (Spizella breweri breweri) Common summer resident 11, ~/.
narri~' ~parrow \.L.OnOLr~cn~a querUl.a) .LO::;~~Dl.e
rare w~nLer re~~ueni...
White-crowned
sparrow (Zonotrichia leucophrys leuchophrys; ~. 1. gambelii)
Common resident 1/.
White-throated
sparrow (Zonotrichia albicollis) Possible ac~idental migrant.
Fox sparrow (Passerella iliaca schistacea) Rare Summer resident.
Lincoln's sparrow (Melospizalincolnii
alticola) Common migrant and Summer
resident.
Song sparrow (Melospiza melodia) Common resident 11.
Lapland longspur (Calcarius lapponicus alascensis) Possible rare winter
migrant.
Chestnut-collared
longs pur (Calcarius ornatus) Possible accidental winter
migrant.

L. h.

�19A

Raptores

~/

Turkey vulture (Cathartes aura meridionalis) Common summer i/, ~/ and
rare winter resident.
Goshawk (Accipiter gentilis atricapillus) Uncommon resident ~/.
Sharp-shinned hawk (Accipiter striatus velox) Uncommon summer i/ and
winter resident.
Cooper's hawk (Accipiter cooperii) Uncommon summer ~/ and winter
resident.
Red-tailed hawk (Buteo jamaicensis calurus) Common resident i/, ~/.
Swainson's hawk (Buteo swainsoni) Uncommon migrant and summer resident.
Rough-legged hawk (Buteo lagopus ~. johannis) Common winter resident.
Ferruginous hawk (Buteo regalis) Rare summer resident.
Golden eagle (Aquila chrysaetos canadensis) Common resident ~/, 2/'
Bald eagle (Haliaeetus leucocephalus alascanus) Common winter resident 2/.
Marsh hawk (Circus cyaneus hudsonius) Common resident 4/.
Osprey (Pandion haliaetus carolinensis) Rare migrant.
Prairie falcon (Falco mexicanus) Uncommon resident.
Peregrine falcon (Falco peregrinus anatum) Rare resident.
Pigeon hawk (Falco columbarius) Rare winter migrant.
Sparrow hawk (Falco sparverius sparverius) Common summer and uncommon
winter resident 3/, 4/.
Screech owl (Otus asio) Possible rare resident.
Flammulated owl (Otis flammeolus flammeolus) Possible accidental summer
resident.
GrPRr hornpc1 mJl (R1Jb0 Yirgi!li9.!lus)
C0~0!"! ree Lderrt 1/, 5.-./.
Pygmy owl (Glaucidium gnoma californicum) Uncommon resident 3/.
Burrowing owl (Speotyto cunicularia hypugaea) Uncommon summer and possible
accidental winter resident.
Long-eared owl (Asio otus wilsonianus) Uncommon resident.
Short-eared owl (Asio flammeus flammeus) Uncommon winter migrant and resident.
Saw-whet owl (Aeg~s
acadicus acadicus) Uncommon resident.

5/ Golden and bald eagles specifically excluded from statutes defining
"Raptore" as cited in footnote 1:./ but herein listed to avoid omission.

B. D. Baker
February 1972

�20A

DISTRIBUTION AND ABUNDANCE OF BIG GAME HAMHALS
WILDLIFE :MANAGEHENT UNIT 61 (W'oUNCOHPAHGRE)

BLACK BEAR
Black bears occupy approximately the area between 7,000 and 10,000 feet
in Unit 61, though at times they may wander over almost the entire unit
(Gilbert 1953).

Based upon the hunter kill surveys, an average annual kill of 19 bears
occurred 1960-1970 in Unit 61, ranging from 3 to 62 animals. ,During
preseason deer sex ratio counts six bears were counted in 1970 and nine
in 1971 (Burdick pers. comm. 1972).

Southwest Regional office files

contain details of locations observed.

ELK
Big game winter range surveys conducted jointly with the BLM and Grand
Hesa-Uncompahgre National Forest from 1961 through 1963 failed to produce much information on distribution of elk in Unit 61.

In fact, of

71 browse range-pellet group transects that were read within or near the
wintering deer zone, only five contained any elk groups at all (F.A.
Project W-IOI-R files).

Current information supplied by Southwest Region staff and field personnel
through group and individual interviews in 1972 yielded year-round distribution for elk.

That information has been displayed by mylar overlay (see

accompanying map portfolio).

Interview data regarding distribution also revealed that elk apparently
move upwards to the extreme plateau crest (over 10,000 feet on Horsefly
Peak) in summer and downward to about 7,000 feet in winter.

In addition,

�21A

elk probably
Generally,
boundary

can be found anywhere

the lower limit line corresponds
or southwestward

outside

As is the case with adjoining
61.

in between

Supporting

those elevations

with the National

of it two to four miles.

is the low average

head for the years 1965-70.

Division

are distributed

small groups where

MOUNTAIN
The literature
distribution
occurrence

available

one 2 miles northeast
Dixon

listed, however,

Hunter

collections)

of Norwood,

of the forested

report

summaries

it within

during

of 40

say that herds

found.

to mountain

three specimens

taken

lion habitats

more isolated

at least a minimum

any mountain

of

and one near Gateway.

that the rougher,

inclusive.

lion

(1972) whose records

any of the better

fail to indicate

killed by people other than hunters
recorded

people

reference

indicate

area would harbor

61 for the years 1965-1970

annual harvest

one west of Redvale,

it seems reasonable

in Unit

LION

makes no specific

(1967) does not include

portions

regional

within Unit 61, except Armstrong
(from museum

Forest

Unit 62, elk are not very numerous

this contention

in relatively

any time.

population.

lions taken in Unit

A few lions could easily have been
and no reports

recorded.

Nonewere

deer trend counts.

MULE DEER
Joint BLM-Forest

Service-Division

Project

files) were primarily

W-lOl-R

zone of winter
information

deer use.

aimed at establishing

Data from those investigations

from interviews

an upper winter

big game range surveys of 1961-63

of region personnel

limits line roughly

including

the average

combined

in February

the following

(F.A.

with

1972 yielded
areas

(see

�22A

mylar map
1.

overlay):
At Unit 61's extreme

southeastern

corner, Iron Springs Mesa,

upper Sheep Draw, and all lower and middle
and McKenzie
2.

Creeks.

Lower Craig Draw and lower Clay, Horsefly,

and Cottonwood

Creeks areas inside of the Forest boundary

two to four miles

(northeast
3.

Middle

and north of Norwood).

Coal Canyon

and Tabeguache

Pinto Mesa and The Meadows
4.

Upper middle
(northeast

5.

areas of Goodenough

Campbell,

Creek to include most of

all north of Nucla.

Atkinson,

and Mesa Creeks areas

and north of Uravan).

Lower Pine Mountain

and most of upper Ute Creek southeast

and east of Gateway.
6.

Bottomland

and extreme

Gulch to the divide

The lower limits of winter

Lowe r slopes of West Creek from Turner

in Unaweep

Canyon.

deer distribution

generally

Unit 61's southern

and western

cultivated

near Nucla which would not be included

farmland

boundary.

coincide

An exception

All of Unit 61 is believed

to be occupied

drier portions

light populations.

at higher

supporting

elevations

with

would be the
as winter range.

by deer in summer, with lower,
Heaviest

from about 7,000 feet elevation

summer use occurs

to the crest of the

plateau.

Joint winter

range surveys

high fall-winter
Creek area;
Calamity

yielded

pellet group count indices

deer use in: (1) the Sanborn Park-Craig

(2) the Dry Park-Cottonwood

Creek area.

of fairly

Draw-Goodenough

Creek area; and (3) Calamity }1esa-

The Shavano-Campbell-Spring

Creeks and Blue Hesa-

�23A

North Fork Mesa Creek areas also produced moderately high indices of
winter use.

More currently, deer aerial count data for the Uravan trend area (see
DEER overlay) are available at the Division Southwest Region office in
Montrose.

Total counts of deer have varied between a low of 749 in 1969

and a high of 1,411 in 1971 (Burdick pers. comm. 1972).

Literature Cited

Armstrong, D. M.

1972.

Distribution of Mammals in Colorado.

Kansas Printing Service, Lawrence.

Dixon, K. R.

1967.

pp. 141-164.

415 pp.

Evaluation of effects of mountain lion predation,
In Game Res. Rep., July-Part II.

Fish and Parks.

Gilbert, D. L.

Univ. of

1953.

Colo. Dept. Game,

pp. 73-310.

Fur resources and bear studies.

Quart. Prog. Rep., Jan.

Colo. Dept. Game and Fish.

pp. 59-60.
141 pp.

W. T. McKean and B. D. Baker
February 1972

In

�'24A

DISTRIBUTION AND ABUNDANCE OF SMALL GAME MAMMALS
WILDLIFE MANAGEMENT UNIT 61 CW. UNCOMPAHGRE)
COTTONTAIL
Every square mile in this unit is occupied by cottontails, or capable of
being so, with the possible exception of three or four urban areas.

Based

on reported altitudinal preferences CLech1eitner 1969, Armstrong 1972), both
the desert cottontail and Nuttall's cottontail would be present.

No specific

study of cottontail distribution pertinent to the unit is known.
Shepherd (1965a) indicates a possible density of 150-200 animals per square
mile for most of the better areas of the state excluding cyclic highs or
Lows , This estimate should be applicable to Unit 61.

SNOWSHOE HARE
Shepherd (1965b) indicates that snowshoe hares occupy, or could occupy,
Unit 61 between 8,000 and 10,000 feet elevation in the coniferous forest
for its entire length (see accompanying mylar map portfolio).
Shepherd (1965b) postulates a statewide mean density of 125 hares per section
on the habitable range.

This is believed to be applicable to the higher

areas of this unit.

PINE (RED) SQUIRREL
Pine squirrels are reportedly distributed throughout the upper one-half
(elevationally) of Unit 61 in the coniferous forests.

Armstrong (1972)

indicates, on a distribution map, that they are pres ent; ,
Southwest Regional personnel indicate tnat where pine squirrels are present
at all in the unit, they exist in maximum numbers.

�25A

ABERT'S SQUIRREL

Southwest Regional personnel have seen this squirrel in Unit 61 in widely
scattered sites, but one specific area lies between McKenzie and Clay
Creek drainages.

Lechleitner (1969) states that it occurs in the ponderosa

pine forests at 5,000 to 8,000 feet.

Maps in Armstrong (1972) and Hall and

Kelson (1959) do not show the distribution to extend this far into westcentral Colorado.

Abundance of this squirrel is undocumented for Unit 61, but it is definitely
present (see accompanying map portfolio).

Literature Cited
Armstrong, D. M.

1972.

Distribution of mammals in Colorado.

Kansas Printing Service, Lawrence.

Hall, R. E., and K. R. Kelson.
Ronald Press, N. Y.

Lechleitner, R. R.
Boulder.

1969.

1959.

2 Vols.

415 pp.

The mammals of North America.

The

1083 pp.

Hild mammals of Colorado.

Pruett Publ. Co.

254 pp.

Shepherd, H. R.

1965a.

Colorado long range management plans for game

species, 1965-75, for cottontail rabbits.
Dept. Rep.

1965b.

11 pp.

Colo. Game, Fish and Parks

(Mimeo).

Colorado long range man agemerit; plans for game species,

1965-1975, for snowshoe hares.
11 pp.

Univ. of

Colo. Game, Fish and Parks Dept. Rep.

(Himeo).
W. T. HcKean
February 1972

�26A

DISTRIBUTION At'ID ABm'l'Dk"il'CE
OF SMALL -.:AME BIRDS
WIL:QLIFE--.!1!WAGj:J~NT
mUT 61 (W. UNCOMPi\1-IGl~)

SAGE GROUSE
Southwest

regional

the southeast

Rogers

personnel

quarter

(1964) states

indicate

that sage grouse are dispersed

over

of the unit.

that populations

of only 1-10 birds per square mile,

or less, were found in the areas listed as occupied

range

(see accompanying

map portfolio).

BLUE GROUSE
Rogers

(1968) indicates

in his maps that the blue grouse

Unit 61 is largely within

Uncompahgre

entirely.

habitat

The important

National

distribution

Forest boundaries,

of fir (Abies) species,

in

but not

as well as mixed

brushy

areas and aspen occur on Bureau

of Land Management

private

lands, but not as extensively.

Blue grouse occur at 7,000 to 10,000

feet most

commonly.

A census by Rogers
between

(1968) of 62.5 road miles along the Uncompahgre

Units 61 and 62 in 1961-1963

blue grouse
being

lands as well as

per mile,

conducted

ranging

developed

from 0.025 to 0.097.

count of 0.039

This census route is not

at present.

SHARP-TAILED
Rogers

(1969) reports

grounds

or observed

on the Tongue;

an average

Divide

elsewhere

}~ntrose

of Sams; San Miguel

sma1~ populations

GROUSE
of sharp tails either at dancing

in Unit 61 as follows:

County-on

Horsefly

County-scattered

Mesa Count.y -Out.Law Mesa,

Creek, near Johnson

Springs north

small flocks in the southwest

portion

�27A

of the Uncompahgre
T-Bone Springs
accompanying

Plateau.

More recently,

area, Montrose

County

counts have been made in the

(Arant pers.

comm. 1972).

(See

map portfolio).

Only very small numbers

of birds were observed

at these sites.

CHUKARS
Sandfort

(1965) in surnrnarizin~ chukar distributions,

bird had been introduced
recently,

at many sites within

Southwest

regional

flocks from Gateway

to Nucla.

During

the period 1958-1963

densities
better

varied widely,

personnal

ranging

stated

More

that there are scattered

ones, population

from 47 to 26 birds per square mile in the

1965).

Projected

kill figures show an increase

(total) taken in Small Game Unit 18 in 1968 and 1969 to 498 in

1970 (Funk and Tully 1971).

)

RING-NECKED
Distribution

that this

Unit 61 previously.

in this unit, and adjoining

chukar range (Sandfort

from 82 birds

indicated

coincides

areas, occupying

PHEASh~T

with the extreme west side of the unit in farmland

approximately

36.7 square miles

of fair to poor range

(Swope 1965).

Recent updated

population

data are available

Region office,

including

sex ratios,

success

(Burdick pers.

COnTIno

1972).

and on file in the Southwest

crmv count routes,

and reproductive

�WILD TURKEYS

(1964 through 1967) mapped the appropriate

Hyers

turkeys within Units 61 and 62 for four years.
were widely

observed

He showed that turkeys
of Unit 61.

in Unit 61 in which turkeys were

consistently

throughout

at one or more seasons of the year were:

Tabegauche

Creek, Cottonwood

Turkeys generally

Creek, McKenzie

wintered

(5,500 to 7,000 feet), or near. the pinon-juniper

and summering
and aspen

areas were largely

types; however,

A

Blue' Creek, Spring Creek,

Creek, Clay Creek, Horsefly

Creek, Sanborn Park, and 47 Creek.
elevations

of wild

the forested portions

dispersed

few of the drainages

distribution

at lower

type; breeding

at 8,000-9,500 feet in the ponderosa

they have often been observed

pine

along the top of

the plateau.

Myers

(1964 to 1967) indicated

that the most reliable

census method

several

tried, was the count at artifically

winter.

The data at hand are for the entire Uncompahgre

bining both Big Game Units 61 and 62.
birds,

ranging

numbers

1963 through 1967.

be twe en units.
a considerable

drop in turkey popUlations,

unchecked)

that similar

decreases

have occurred

scattered

throughout

of dense, fruit-producing

in roughly

stations

equal

in Unit 62

it seems probable

(but

in Unit 61.

that band-tailed

both Wildlife

Management

The occurrence 'seems to relate to areas where an abundant
supply

area, com-

PIGEON

(1970, and pers. comm. 1971) indicates

irregularly

in

drop in population

Birds were counted

indicated

Braun

Plateau

a considerable

Since recent counts on similar

BAND-TAILED

food stations

An average feed ground count of 306

from ~40 to 485, indicated

for thie five winters

established

among

pigeons

are

Units 61 and 62.
and consistent

native shrubs exist, as well as an abundance

�29A

of acorns from Gambel' s oak.

Also, valleys wh ere cultivated

grains are

grown have appreciable

numbers

of pigeons.

occurs in the

spruce-fir

pine zones; therefore

and various

is used at all seasons,

Only generalizations
reported

except winter

concerning

observations,

are possible

as one of the places

No differences

in bird densities
Highest

and grain) are readily
cover.

Burdick

Plateau

through April).

pigeon

(Braun 1970).

abundance,

based

The uncompahgre

upon
Plateau

in Colorado where pigeons

are most abundant.

have been detected

Wildlife

observed

available

commonly

the entire Uncompahgre

(December

band-tailed

is mentioned

Units 61 and 62.

Nesting

densities

occurred

and adjacent

(pers. comm. 1972) reported

between
where

to forested

having

Managemep_t

food (berries,

roosting

acorns

and escape

counted in 1970 by helicop-

ter in excess of 2,500 birds within Units 61 and 62.

MOUkNmG

Mourning

doves are distributed

spring,

summer,

Unit

and fall

over the entire Uncompahgre

(Braun pers. comm. 1971).

(Quick pers. comm. 1972).

doves on the Uncompahgre

DOv""ES

Braun

during

A few doves winter

No data exist concerning

Plateau.

Plateau

densities

in the

of mourning

(pers. comm. 1971) stated that higher

concentrations

occur in the pinon-juniper

and farmland

or spruce-fir.

Over three hundred birds were trapped

types than in the pine
in Unaweep

Canyon

during

the summer of 1971.

WATERFOHL
Breeding

population:

of dabbling
Dolores

Hopper

(pers.

ducks are dispersed

comm.197l)

along creeks tributary

Rivers and on those rivers themselves,

irrigated

farmlands

adjacent.

indicated

that low populations

to the San Miguel

and

as well as in the limited

Also, small numbers

nest in isolated

beaver

�30A

ponds and other high elevation wat er areas.

Hopper

(pers. comm. 1971) estimated

the spring population

be about two breeding

pairs per square mile

made over comparable

land in the San Luis Valley.

of 50 square miles of irrigated

cropland

produces

a population

of 200 ducks.

Wintering

Population:

No information

tribution

or abundance

of wintering

numbers

A very rough estimate

and river bottom vegetation

concerning

types

either dis-

ducks in Unit Gl, but obviously

the

are small.

Four transplants

INTRODUCTIONS.

(S.27, R.l7W, T.5lN).

Subsequently,

.

For details

25 birds were planted

1965

see section

on

on South Fork of Mesa

.

(S.17,.T.49N.,

The introductions

R16H).

(See accompnaying

above-mentioned

25 on South Fork of Mesa Creek.
in a flock wintering

totaled

1970.

Res. Rep., Oct.

340 birds on Indian Creek and the
have revealed

up to 25

on South Fork of Hes a Creek (Barnes pers. comm. 1972).

Band-tailed

pigeon

Cited
investigations,

Colo. Div. Game, Fish and Parks.

Funk, H. D., and R. J. Tully.
survey -- 1970.

map portfolio).

Recent observations

Literature
C. E.

QUAIL

of this exotic bird were made into Unit 61 between

and 1970 on Indian Creek

Braun,

(4 ducks), based on counts

is available

MOUNTAIN

Creek

of Unit 61 to

1971.

Colorado

pp. 151-171.

In Game

171 pp.

small game hunter harvest

Colo. Div. Game, Fish and Parks, Game Planning

Services.

�31A

Myers, G. T.

1964 thru 1967.

Game bird surveys.

Colo. Dept. Game, Fish and Parks.

Rogers, G. E.

1964.

No. 16.

Sage grouse investigations in Colorado.

The blue grouse in Colorado.

Game, Fish and Parks Dept.

1969.

132 pp.

Tech. Publ. No. 21.

The sharp-tailed grouse in Colorado.

Sandfort, W. W.

1965.

for pheasants.

Colo.

Tech. Publ. No. 23.

94 pp.

Colorado long range plans for game species, 1965-

1975, for Chukar partridge.

1965.

Tech. Publ.

63 pp.

Colo. Div. Game, Fish and Parks.

Swope , R. H.

Pages variable by years (proc~ssed).

Colo. Game, Fish and Parks Dept.,

1968.

In Game Res. Rep., April.

Colo. Game, Fish and Parks Dept. Rep.

19 pp.

Colorado long range plans for game species, 1965-1975,
Colo. Game, Fish and Parks Dept. rep.

W. T. HcKean
February 1972

54 pp.

�32A
DEER KILLI SEASONS AND HUNTING PRESSURE
HILDLIFE MANAGEMENT UNIT 61
YEAR

HUNTING
PRESS URE~'~ BUCKS

DOES

FA"I-lNS TOTAL

HUNT AND SEASON

1956

959

449

266

46

761

ESM
T

10/15
12/1

10/31
12/31, In Part

1957

1,316

669

433

144

1,246

2DM

10/15

11/17

1958

2,272

793

642

291

1,726

1DM
1DM

10/15
11/3

11/2
12/31, In Part

1959

2,382

900

782

267

1,949

1DM
1DM
2DM

10/17
11/4
12/5

11/3
11/17, In Part
12/31, In Part

1960

1,754

981

714

257

1,952

lDM

10/17

11/30

1961

2,242

1,484 1,277

605

3,366

3D

10/21

11/30

1962

1,751

933

749

280

1,962

1DM

10/20

11/4

1963

1,911

945

902

385

2,232

lDM
1DM

10/19
11/23

11/7
12/15, In Part

1964

2,041

1,049

703

217

1,969

lDM

10/17

11/5

·1965

2,266

946

794

280

2,020

1DM
1DM

10/16
12/4

11/5
12/19, In Part

1966

1,709

487

318

98

903

ES
ES

10/15
12/3

11/6
12/11, In Part

1967

1,509

519

369

56

944

ES

10/21

11/9

1968

1,571

502

413

82

997

ES

10/19

11/7

1969

2,953

969

495

116

1,580

ES

10/18

11/6

1970

2,763

1,094

92

·12

1,198

AO
ES

10/17
10/17

11/6, In Part
11/6, In Part

Based upon combined total resident and non-resident license sales and expressed
in number of licenses.
ES = Either Sex, One Deer.
T
= t.va Deer, Either Sex, One License.
ESM = Either Sex Multiple (Unlimited licenses and deer per individual).
2DM = Two Deer Hultiple, Either Sex (2 licenses and 4 dcer per indivi.dual).
IDM = One Deer Multiple, Either Sex (2 licenses and 2 deer per individual).
3D
Three Deer, One Each Either Sex (1st and 2nd licenses and 3rd deer
coupon with 2nd license).
Note: IDM evolved into 2D, HC or THO Deer, Hunter's Choice (2 licenses and 2
deer per individual) and is synonymous.
-k

B. D. Baker

�33A

ARCHERY DEER KILL AND SEASONS
WILDLIFE MANAGEMENT UNIT 61
YE.AR]),]j .BUCKS

DOES

HUNT AND SEASON

FA\oJNS TOTAL

1954

ES

10/1

1955

ES
T

10/1
10/15

ES
ESM
T

10/1
10/15
12/1

2DM

10/1

ES
1DM
ES

9/1
10/15
11/3

-

1956

1957
1958

5

0

0

5

10/14

-

10/13
10/31
10/14
10/31
12/31, In Part
11/17

-

9/30
11/2
12/31, In Part

1959

0

0

0

0

ES .
1DM
2DM

9/15
10/17
12/5

-

9/30
11/17, In Part
12/31, In Part

1960

9

0

0

9

ES
1DM
2DM

9/10
10/17
12/17

-

9/30
11/30
12/31, In Part

3D
3D

8/26
10/21

-

9/10
11/30

ES
1DM

8/25
10/20

-

9/23
11/4

ES
lDM
1DM

8/17
10/19
11/23

-

9/8"
11/7
12/15, In Part

ES
lDM

8/15
10/17

-

9/13
11/5

1961
1962

1963

2

4
2

2

5

5

10

0

0

4
19

7

21

-

-

1964

14

1965

13

5

0

18

ES
lDM
1DM

8/21
10/16
12/4

1966

16

10

0

26

ES
ES
ES

8/20
iO/15
12/3

-

ES
ES

8/19
10/21

-

ES
ES

8/17
10/19

-

1967 ]j

1"'8
~o
-4/

5

5

7

0

-

4

5

0

0

9

10

-

9/12
11/5
12/19 , In Part
9/18
11/6
12L11 , In Part

9/17
11L9

9/15
110

�34-A

Archery
YEAR

Deer Kill

&amp; Seasons - Wildlife Management Unit 61 (Continued)

1/, Jj BUCKS

DOES

FAWNS

TOTAL

HUNT AND SEASON
----'"-j

1969 2/

39

0

0

39

AO
ES

8/16
10/18

9/14
11/6

1970 !jj

21

0

a

21

AO
ES
AO
ES

8/15
9/1
10/17
10/17

8/31
9/20
11/6, In Part
11/6, In Part

1/ Years

1954-1956 free permits were issued to holders of regular big game
hunting licenses by application only. Years 1957-1960 there was no issue of
special permits; archery hunting was allowed to holders of valid deer licenses.
Years 1961-1971 separate archery license regulations have been in effect.

1/ Inconclusive

kill data are omitted

for years 1954-1957.

1/ 72 total resident

and non-resident

archery

licenses recorded

for unit.

~/ 66 total resident

and non-resident

archery

licenses recorded

for unit.

1/ 1,163

total hunts recorded

for unit.

&amp;/ 101 total hunters recorded for unit.
Note:
AO
ES
ESM
lDM

Hunt symbols

are explained

as follows:

== Antlered

Only, One Deer.
Either Sex, One Deer.
Either Sex Multiple (Unlimited licenses and deer per individual).
==
One Deer Multiple, Either Sex (2 licenses and 2 deer per individual),
which evolved into 2D, HC or Two Deer, Hunter's Choice (2 licenses
and 2 deer per individual) and is synonymous.
lDMA == One Deer Multiple, One Must Be Antlerless (2 licenses and 2 deer per
individual) .
T
== Two Deer, Either Sex, One Licenseo
2DM == Two Deer Multiple, Either Sex (2 licenses and 4 deer per individual).
3D
Three Deer, One Each Either Sex (1st and 2nd licenses and 3rd deer
Coupon with 2nd license).
3DA
Three Deer, One Must Be Antlerless
(2 licenses plus 3rd deer coupon
on 2nd license).

B. D. Baker
.September 1971

�35A

ELK KILL, SEASONS AND HUNTING PRESSURE
WILDLIFE HANAGEHENT UNIT 61

YEARY

;HUNTING
PRESSURE.Y

1957

HUNT AND SEASON

TOTAL

BULLS

COWS

CALVES

l3

2

0

0

2

25 AOP]/

1958

55

9

0

0

9

AO

10/15

1959

29

12

0

0

12

AO

10/17 - 11/3

1960

81

12

0

0

12

AO

10/17 - 11/6

1961

43

20

0

0

20

AO

10/21 - 11/8

1962

36

6

0

0

6

AO

10/20 -.11/4

1963

40

14

0

0

14

AO

10/19 - 11/7

1964

59

13

0

0

13

AO

10/17 - 11/5

1965

98

20

4

2

26

AO + 30 ASOP]/ 10/16 - 11/5

1966

73

20

5

4

29

AO + 50 ASOP]/ 10/15 - 11/3

1967

80

15

3

0

18

AO + 50 ASOPJ./ 10/21 - 11/9

1958

133

16

12

0

28

AO + 50 ASOP]/ 10/19 - 11/7

1969

339

67

11

2

80

AO + 50 ASOP]/ 10/18 - 11/6

1970

329

45

12

3

60

AO + 50 Asopl/

10/15 - 10/31

10/17 - 11/6

1/ Unit was closed to elk hunting within recent historj prior to 1957.
~/ Based upon combined total resident and non-resident license sales
expressed in nwnber of licenses. Years of antlered only seasons, either
singly or combined with cow permits, very probably yield conservative
hunting pressure statistics. Despite difficulties in verifying the
foregoing assumption, unknown numbers of deer hunters also carry elk
licenses hoping for a bull elk. This happens regardless of how low
elk populations might be. Thus, because of low success and tendency
(above s t.a t ewi.d
e nonreporting percentages that are used in projections)
for unsuccessful elk license holders not to report by card, elk present
in Unit 61 probably receive extra pressure over that indicated by card
return projection years of antlered only seasons.

(Footnotes continued on following page) .

11/2

�36A

ELK KILL, SEASOtjS AND HUNTING PRESSURE
WILDLIFE HANAGEHENT UNIT 61, continued

1/ Unit is combined with Unit 62.
Note:
AO
AOP
ASOP

Hunt symbols are explained as follows:

=

Antlered Only.
Antlered Only Permits.
Antlerless Only Permits.

B. D. Baker
September 1971

�37A
ARCHERY ELK KILL AND SEASONS
WILDLIFE MANAGEMENT UNIT 61

HUNTING
PRESSURE ]j BULLS.?! cm~sJj CALVEs'll TOTAI2./

YEAR ]j

HUNT AND SEASON

1965

AO + 30 ASOP

10/16 - 11/5

1966 .

AO + 50 ASOP

10/15 - 11/3

1967 ]/

ES
AO + 50 ASOP

- 9/17
9/2
10/21 - 11/9

1968 !!../

0

0

0

0

ES
AO + 50 ASOP

8/17 - 9/15
10/19 - 11/7

1969

33 1/

0

0

0

0

AO
AO + 50 ASOP

8/16 - 9/14
10/18 - 11/6

1970

17 &amp;./

0

0

0

0

AO
ES

8/15
9/1

- 8/31
- 9/20

1/ 1957 was first elk season in recent history; and, the 25 antlered only regular
elk season permits for Units 61 and 62 combined that year could have been legally
obtained, however unlikely, by archers. For the years of 1958-1964, archery
.elk hunting was permitted for regular elk license holders during regular seasons
under antlered only regulations as well as for 1965-1969 regular seasons under
antlered only-specified antlerless (cow) permit regulations.

1/ Dash denotes no data available.

Zero means no kill(s) recorded.

1/ First year that archery elk licenses were authorized and sold •.
.i/ Does not include harvest by archers during gun season.
5/ Expressed as number of hunts.

&amp;./ Expressed as total participating resident and nonresident license holders.
Note:

Hunt symbols are explained as follows:

AO

= Antlered Only, One Elk.

ES

Either Sex, One Elk.

ASOP

Antlerless Only Permits (For Units 61 and 62 combined).

B. D. Baker
September 1971

�38A

BLACK BEAR HARVESTS AND SEASONS - WILDLIFE MANAGEMENT UNIT 61
YEAR

HARVEST 1/

1955

l7R

8/15 - 10/1; regular deer and regular archery bear seasons.

1956

0

5/15 - 8/15; regular and post deer and regular archery
bear seasons.

1957

3R+4S=7

4/1 through regular deer, elk, and archery seasons ending
11/15/1957.

1958

10R+3S=13

4/1 through regular deer, elk, and archery seasons and
extended deer season, in part, ending 11/15/1958.

1959

5R

4/1 through regular deer, elk, and archery seasons ending
11/1/1959.

1960

3R

4/1 - 9/15; regular deer, elk and archery seasons ending
11/6/1960.

1961

22R+10S=32

4/1 - 9/15; regular deer and elk seasons ending 11/9/196l.

1962

3R+3S=6

4/1 - 9/15; regular deer and elk seasons ending 11/4/1962.

1963

6R

4/1

1964

9R+14S=23

4/1 - 9/15; regular deer and elk seasons ending 11/5/1964.

1965

2R

4/1 - 9/30; regular deer and elk seasons ending 11/5/1965.

1966

12R+3S=15

4/1 - 9/30; regular deer and elk seasons' ending 11/6/1966.

1967

17

4/1 - 9/30; regular deer and elk seasons ending 11/9/1967.

1968

30R+32S=62

4/1 - 9/30 ; regular deer and elk seasons ending 11/7/1968.

1969

16

4/1 - 9/30; regular deer and elk seasons ending 11/6/1969.

1970

11R+1lS=22

4/1 - 9/30; regular deer and elk seasons ending 11/6/1970.

HUNT AND SEASON ]j, jj

9/15; regular deer and elk seasons ending 11/7/1963.

1/ R=Regular Big Game (Deer and/or Elk) License; S= Bear License for Special
Spring, Summer, or Spring-Summer seasons; Unaugmented numbers represent total
animals harvested not differentiated by season and license controls.

11 1955 was the first year of bear license sales and corresponding season regulations. From 1955-1959, 1 bear per bear license and/or 1 bear per bear coupon
on either or both deer and elk licenses were allowed; from 1960-1965, same as
1955-1959, except bear license variously invalid after 9/1 or 9/30, as noted,
when bear coupon on either or both deer and elk licenses covered take and
possession for the regular big game seasons; 1966, same as for 1960-1965, except
one bear, hunter's choice, per person per calendar year was the allowable bag
(footnotes continued on next page)

�39A

BLACK BEAR HARVESTS

AND SEASONS

- WILDLIFE

MANACEMENT

UNIT 61 (Continued)

1/ continued
limit; for 1967 to the present, 1 bear, hunter's choice, per bear or sportsman's license per person per calendar year only during special bear and regular
deer and elk seasons.
See deer and elk gun and archery kill and season summary
tables herein and/or annual regulations for dates and more details of seasons.

1/ Dogs were

permitted

except during bear seasons

concurrent

with deer and

elk seasons.

B. D. Baker
September 1971

�40A

MOUNTAIN LION HARVEST AND SEASONS - WILDLIFE MANAGEMENT UNIT 61

YEAR 1/

HARVEST ]j

HUNT AND SEASON

1965

(0 - Mesa Co. ,1965-66)

10/16/65 - 3/31/66 West ~lope; 10/23/65 _
3/31/66 East Slope; 1 either sex

1966

(5M - Mesa Co ,,1966-67)

1/1/66 - 12/31/66 counties west of
Continental Divide plus Jackson, Conejos,
Alamosa, Mineral, Saguache, Rio Grande,
Costilla, Archuleta, Hinsdale, and San
Juan counties; 3 lions per year.
10/22/66 - 2/28/67 counties east of
Continental Divide except those above;
3 lions per year.

1967

No Data

1968

o

9/1/68 - 9/1/69; 1 either sex per year.

1969

o

--9/1/69 - 3/31/70; 1 either sex per year.

1970

o

9/1/67 - 3/31/68 statewide; 9/1/67 - 5/31/68
west of State Hwy. #13 and north of U.S.
Hwy. #6; 1 either sex per year.

10/17/70 - 11/6/70 concurrent with regular
big game seasons; 1 lion, either sex.

1/ Bounty of $50 per lion was paid within period of May 7, 1929 through March
12, 1965; lions became protected by statute July 1, 1965 when licensed hunting
was authorized.
2/ Data for Mesa County for years 1965-1966 are from Dixon (1967:147). Dixon
(1967:152) also said that lions we re uncommon in Hesa County south of the
Colorado River to indicate that kills originating in the Unit 61 portion of
Mesa County are improbable (but very possible). Data for 1968-1970 are
hopefully accurate under compulsory reporting regulations for hunters, although
the present damage law allows lax reporting of kills of suspected marauders by
offended citizens.

LITEP~~TURE CITED
Dixon, K. R. 1967. Evaluation of effects of mountain lion predation. pp. 147,
152. In Game Res. Rep ,, July - Part II. CcLo , Dept. Game, Fish and
Parks. pp. 73-310.

B. D. Baker
September '1971

�41A

SMALL GAME HARVESTS

AND HUNTING

UNIT 18 1/, ']j

SMALL GANE MANAGEMENT

NUMBER
SPECIES )j

PRESSURE

HARVEST

OF HUNTERS

1970

1968

1969

0

Closed

Closed

0

580

334

845

1,459

545

70

104

76

76

84

498

485

708

681

5,224

7,461

4,525

No Data

326

169

No Data

2,590

905

23

0

61

0

0

153

Pheasant

439

1,104

840

1,759

5,943

1,900

Ptarmigan

0

30

34

0

0

0

148

301

64

328

526

318

25

199

74

0

0

0

71

128

73

115

156

l30

1970

1968

1969

Closed

Closed

Blue grouse

298

Chukar partridge

pigeon

Band-tailed

rabbit

Cottontail
Dove
Gambe1's

quail

Sage grouse
Sharp-tailed
Snowshoe

hare

grouse

1/ See Chap.

3 - Colo. Game, Fish and Parks Division Laws and Regulations
Handbook; Hild1ife Nanagement Unit 61 is wholly contained within Small Game
Management Unit 18 and approximates the latter's northern half.

1/

Colorado Small Game and/or Waterfowl Harvest Surveys reports for 19551970 also present hunting pressure and kill by region and county, including
Nesa, Nontrose, and San Niguel Counties; harvest by Small Game Nanagement
Unit are only available in reports of 1968-1970.

]j

Since July 1, 1969, w i l.d turkey has had legal status as a "game bird"
w i t.h i.n the broader category of "small game."
Prior to 1969, the turkey
wa s treated as "big game."
Because of these irregularities and the bird' s
relatively high ranking as a game species, separate information has been
presented in the table "Turkey Harvests and Hunting Pressure" elsewhere in
th is report.

B. D. Baker
September 1971

�42A

DUCK AND GOOSE HARVESTS AND HUNTING PRESSURE _
SAN MIGUEL COUNTY 1/, 1954-1970

Jj

YEAR

DUCKS
EST. NO. HUNTERS

1954

No Data

522

No Data

0

1955

83

405

27

0

1956

No Data

0

No Data

0

1957

No Data

840

No Data

0

1958

34

186

0

0

1959

81

98

29

0

1960

0

0

0

0

1961

0

0

0

0

1962

33

144

0

0

1963

0

0

0

0

1964

0

0

0

0

1965

13

0

0

0

1966

6 ]/

220

0

0

1967

52

96

25

21

1968

53

337

0

0

1969

64

595

0

0

1970

27

0

0

0

EST. HARVEST

].1

GEESE ]j
EST. NO HUNTERS

,

EST. HARVEST

1/ Data do not have great applicability to Unit 61 but are best onhand.
Although available and involved, Mesa and Montrose County information can
not be interpreted in relation to Unit 61. Information for those counties
are thus omitted.
]j Accuracy of data is questionable, probably due to sampling difficulties
associated with comparatively small populations. Geese data are presented
to indicate that kills are mostly accidental and bird abundance and hunter
numbers so far are very negligible.
3/
Very inaccurate.

Figures are considerably inconsistent w.i t.h other data.
B. D. Baker
September 1971

�43A

TURKEY HARVESTS AND HUNTING PRESSURE
WILDLIFE HANAGE:1'1ENT
UNIT 61 }j

HARVEST
YEAR

TOMS

HENS

POULTS

TOTAL

1954
1955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969 ]./
1970

2
2
3
3
0
l3
27
36
38
38
29
3
7
5
2
2
0

2
0
0
6
2
11
49
88
45
35
36
7
4
0
3
4
0

0
0
0
0
5
11
8
20.
23
22
18
11
11
0
0
2
0

4
2
3
9
7
35
84
144
106
95
83
21
22

HUNTING
PRESSURE 1./
N.D.
N.D.
N.D.
21
11
37
151
126
144
119
156
68
50
7
9
N.D.
21

5
5

8
0

1/ For years 1954-1967; effective 1968, Unit 61 became part of Small
Game Management Unit 18, so data for years 1968-1970 apply to the latter
area.

1./ Expressed as "total hunters" for years of records.

N.D. = No Data.

1/ The legal definition of wild turkey was changed July 1, 1969 from
that of "big game" to that of "game bird", the latter also within the
broader category of "small game."

B. D. Baker
.September 1971

�GAME SPECIES INTRODUCTIONS - WILDLIFE MANAGEMENT UNIT 61
Species
Chukar=

Date

Hales

Females

Young

Total

-

-

-

174

3-26-59
3-29-60
3-16-62
3-21-63

87

134

-

-

221
126
50
150

8-19-65

-

-

-

48

8-25-65
8-20-66

-

-

-

95
163

1-25-68

-

-

-

66

6-23-70

-

-

-

25

12-7-55

-

-

.1&gt;-.J&gt;!J&gt;

Release Area

Source of Stock

1 mt . SE Gateway, E. of
Dolores River.
Paradox Valley.

Game farm.

"

"

11

Sandfort's files

"
"

"

Mesa Creek, Montrose Co.
S5,T49N,R18W; Blue Creek.

Reference

Sandfort's
Sandfort's
Sandfort's
Sandfort's

files
files
files
files

" "
" "
-------------------------------------------------------------------------------------------------------------------------Mountain
quail

S27,T5lN,R17W; Indian
Cr., Mesa County.

"

"

"

"

About 4 mi. W. of 1965
releases.
1 mi.W. USFS Bdy. on
Indian Cr. ,Mesa Co.
S17,T49N,R16W; S. Fork
of Mesa Cr. ,Mesa Co.

Wild-trapped
California

from

Rogers (1966)

"

"

"

Rogers (1966)

"

"

"

Rogers (1967)
Rogers (1968)

Wild-trapped from
Oregon
Game farm.

Hoffman (1971)

-------------------------------------------------------------------------------------------------------------------------lVild
turkey

1934
1935-36
1945

-

1946

-

Unknown

-

Gateway

"

-

Ute Ranger Station

8

Calamity Creek

-

1-8-46
11-22-46
12-4-46

4

6
9
24

1947

-

-

3
3

-

12
28

-

15

4
9

*No records are available for years prior to 1955.

very likely included releases in Unit 61.

"

N.Mex. ,Burget (1957)

"
"

"
"

"

Clay Creek

"

Variable,-Ok1a.,
Texas, Mexico
Unknown

"

Turkey Ranch
f1

"

Tabeguache Basin

"

"

Cottonwood Creek

Unknown

Myers (pers. comm.
1972)

"
"

"
f1

Burget (1960)
Burget (1960)
Burget (1960)
Myers (pers. comm.
1972)

Division transplanting program, using game farm stock in the 1940's,

�45A

Gfu~E SPECIES INTRODUCTIONS - WILDLIFE MANAGEMENT UNIT 61 (Continued)

Literature Cited
Burget, H. L.

1957.

The wild turkey in Colorado.

Hild Turkey

Invest., F.A. Proj. W-39-R, Colo. Game and Fish Dept.
68 pp.

(processed).

1960.
July.

Jan.

Wild turkey development, pp. 11-42.

Colo. Game and Fish Dept.

Hoffman, D. L.

1971.

1966.

(processed).

Summary of mountain quail field releases.

Colo. Div. Game, Fish and Parks.

Rogers, G. E.

110 pp.

In Quart. Rep.,

1 p.

(mimeo).

Study of mountain quail adaptability, pp. 125-133.

In Game Res. Rep., April.

Colo. Game, Fish and Parks Dept.

192 pp.

(pr oces s ed) •

1967.

Study of mountain quail adaptability, pp. 215-218.

Game Res. Rep., April - Part 2.

In

Colo. Game, Fish and Parks Dept.

(processed).
1968.

Study of mountain quail adaptability, pp. 113-116.

Game Res. Rep., April.

Colo. Game, Fish and Parks Dept.

(processed).

B. D. Baker
February 1972

In

131 pp.

�~·6A

---

----------------

.1

BIG GAJvlEMAMMAL RESEARCH
WILDLIFE

REFERENCES

MANAGE~ffiNTUNIT 61

BLACK BEAR
Boyd, R. J. 1970. Hunter harvest survey, pp. 133-213.
In Game Res. Rep.,
July - Part II. Colo. Div. Game, Fish and Parks.
pp. 127-286.
Denney, R. N., and D. L. Gilbert.
1952. Fur resources and bear studies,
pp. 1-31.
In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.
Gilbert,
,
D. L. 1951. Fur resources study and bear investigations, pp. 1-8.
In Quart. Prog. Rep., July. Colo. Dept. Game and Fish.
102 pp.
1951. Fur resources
Quart. Prog. Rep ,, Oct.

study and bear investigations, pp. 1-9.
Colo. Dept. Game and Fish. 117 pp ,

1953. Extent of bear ranges in Colorado, pp. 59-60.
Rep., Jan. Colo. Dept. Game and Fish.
141 pp.
Gilbert, P. E. 1946. Bear studies.
Game and Fish.
4 pp.

In

In Quart. Prog.

In Quart. Prog. Rep., June.

Colo. Dept.

Oct.

1948. Deer-elk-bear investigations, pp. 30-34.
Colo. Dept. Game and Fish.
80 pp.
1952. Checking station survey, pp. 39-43.
Colo. Dept. Game and Fish.
60 pp.

In Quart. Prog. Rep.,

Jan.

1953. Checking station survey, pp. 59-61.
April.
Colo. Dept. Game and Fish.
99 pp.

In Quart. Prog. Rep.,

Gilbert, P. F., and G. E. Rogers.
In Quart. Prog. Rep., July.

In Quart. Prog. Rep.,

1954. Checking station survey, pp. 125-128.
Colo. Dept. Game and Fish.
182 pp.

Hammit, H. C. 1950. Deer-elk-bear investigations, pp. 10-14.
Proj. W-38-R.
Oct. Colo. Dept. Game and Fish. 21 pp.
Hunter, G. N. 1965.
Fish and Parks.

Colorado
41 pp.

big game harvest,

1959-1965.

Fed. Aid

Colo. Dept. Game,

McKean, W. T. 1965. Colorado long range game species management plans,
1975, for black bear. Colo. Dept. Game, Fish and Parks.
6 pp.

1965-

Tigner, J. R., and D. L. Gilbert.
1960. A contribution toward a bibliography
on the black bear.
Colo. Dept. Game and Fish. Tech. Publ. No.6.
43 pp.

�47A

ELK
Boyd, R. J. 1970. Hunter harvest survey, pp. 133-213. In Game Res. Rep.,
July - Part II. Colo. Div. Game, Fish and Parks. pp. 127-286.
Denney, R. N. 1965. Colorado long range game species management plans,
1965-1975, for elk. Colo. Dept. Game, Fish and Parks. 61 pp.
Gilbert, P. F. 1953. Checking station survey, pp. 59-61.
Rep., April. Colo. Dept. Game and Fish. 99 pp.

In Quart. Prog.

Gilbert, P. F., and G. E. Rogers. 1954. Checking station survey, pp. 125128. In Quart. Prog. Rep., July. Colo. Dept. Game and Fish. 182 pp.
Hammit, H. C. 1951. Deer-elk-bear investigations, pp. 16-22.
Prog. Rep., Jan. Colo. Dept. Game and Fish. 33 pp.

In Quart.

Hunter, G. N. 1965. Colorado big game harvest, 1959-1965.
Game, Fish and Parks. 41 pp.

Colo. Dept.

Rogers, G. E. 1951. Checking station survey, pp •.32-33.
Rep., Jan. Colo. Dept. Game and Fish. 33 pp.

In Quart. Prog.

MULE DEER
Anderson, A. E. 1959. Effects of sagebrush eradication by chemical means
on deer and related wildlife, pp , ll3-l28. In Quart. Rep., Jan. Colo.
Dept. Game and Fish. 128 pp.
1960. Effects of sagebrush eradication by chemical means on deer
and related wildlife. Fed. Aid Job CampI. Rep., Proj. W-38-R-13. July.
Colo. Dept. Game and Fish. 72 pp.
1960. Distribution and abundance indices of selected biota in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Univ., Ft. Collins. 130 pp.
1966. An investigation of 2,4-D application to sagebrush within a
mountain shrub-conifer complex, Uncompahgre National Forest. In Game
Res. Rep., July - Part III. Colo. Dept. Game, Fish and Parks. pp, 345420.
1969. 2,4-D, sagebrush, and mule deer-cattle use of upper winter
range. Special Rep., No. 21., July. Colo. Div. Game, Fish and Parks.
21 pp ,
1969. Sagebrush vegetation before and six years after 2,4-D
application. Outdoor Facts, No. 67. Colo. Div. Game, Fish and Parks.
4 pp.
Baker, B. D. 1955. Detailed study of range forage by use of fenced exclosures,
pp. 149-157. In Quart. Prog. Rep., July - Part II. Colo. Dept. Game and
Fish. pp. 117-175.

�48A

1956. Detailed study of range forage by use of fenced exclosures,
pp. 43-64. In Quart. Prog. Rep., July - Part I. Colo. Dept. Game and
Fish. pp. 1-102.
1957. A directory of Colorado big game exclosures concerning their
locations, studies and other general information. Spec. Rep., Fed. Aid
P'ro
j , hT-38-R,Jan. Colo. Dept. Game and Fish. 60 pp.
1961. A directory of Colorado big game exclosures concerning program
history, study techniques, location and other general information. Spec.
Rep. - revised edition, Fed. Aid Proj. W-IOI-R-3, Jan. Colo. Dept. Game
and Fish. 51 pp.
1964
A detailed study of range forage by use of fenced exclosures,
pp. 3-15. In Game Re. Rep., Jan. - Part I. Colo. Dept. Game, Fish and
Parks. pp. 1-121.
1966. Browse transect analysis and application, pp. 53-67. InGame
Res. Rep., July - Part I. Colo. Dept. Game, Fish and Parks. 93 pp.
1970. Survey, inventory, and analysis of deer and elk winter range,
pp. 15-58. In Game Res. Rep., July - Part I. Colo. Div. Game, Fish and
Parks. 126 pp.
Boyd, R. J. 1970. Hunter harvest survey, pp. 133-213. In Game Res. Rep ,,
July - Part II. Colo. Div. Game, Fish and Parks. pp. 127-286.
Carhart, A. H. 1940. Deer-elk survey, Vol. 4.
Game and Fish Commission. 19 pp.

Fed. Aid Proj. W-4-R.

Gilbert, P. F. 1948. Deer-elk-bear investigations, pp. 14-18.
Prog. Rep., April. Colo. Dept. Game and Fish. 54 pp.

Colo.

In Quart.

1948. Deer-elk-bear investigations, pp. 31-34.
Rep., July. Colo. Dept. Game and Fish. 83 pp.

In Quart. Prog.

1948. Deer-elk-bear investigations, pp. 30-34.
Rep., Oct. Colo. Dept. Game and Fish. 80 pp.

In Quart. Prog.

1949. Deer-elk-bear investigations, pp. 42-45.
Rep., Jan. Colo. Dept. Game and Fish. 67 pp.

In Quart. Prog.

1949. Deer-elk-bear investigations, pp. 47-51.
Rep., April. Colo. Dept. Game and Fish. 98 pp.

In Quart. Prog.

1949. Deer-elk-bear investigations, pp. 33-36.
Rep., July. Colo. Dept. Game and Fish. 84 pp.

In Quart. Prog.

1950. Deer-elk-bear investigations, pp. 41-44.
R~p.: Jan. Colo. Dept. Game and Fish. 66 pp.

In Quart. Prog.

1950. Deer-elk-bear investigations, pp. 60-64.
Rep., April. Colo. Dept. Game and Fish. 101 pp.

In Quart. Prog.

�49A

1950. Deer-elk-bear investigations, Three Year Sum. Rep., 19471950. July. Colo. Dept. Game and Fish. 21 pp.

Jan.

1952. Checking station survey, pp. 39-43.
Colo. Dept. Game and Fish. 60 pp.

In Quart. Prog. Rep.,.

1952. Deer-elk investigations, pp. 41-64, 66-67,69-82, 93-97;
Quart. Prog. Rep., July. Colo. Dept. Game and Fish. 171 pp.
1953. Checking station survey, pp. 59-61.
April. Colo. Dept. Game and Fish. 99 pp.

In

In Quart. Prog. Rep.;',

1953. Detailed study of range forage by use of·fenced exclosures,
pp , 47-49. In Quart. Prog. Rep ,, July. Colo. Dept. Game and Fish.
129 pp.
1954. Correlation of aerial trend counts with ground counts, sex ..
·
ratios, pellet group counts, pre-season and post-season sexing, winter
loss, trend checks, trend counts, sex-ratio counts to determine big game.
numbers, pp. 129-143. In Quart. Prog. Rep., July. Colo. Dept. Game and.
Fish. 182 pp.
Gilbert, P. F., and G. E. Rogers. 1954. Checking station survey, pp. 125~128.
In Quart. Prog. Rep., July. Colo. Dept. Game and Fish. 182 pp.
Gilbert, P. F., et ala 1951. Location and extent of summer ranges, pp.l09110. In Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish. 117 pp.
Hammit, H. C. 1950. Deer-elk-bear "Lnvest LgatLo'ns, pp. 10-14.
Proj. W-38-R. Oct. Colo. Dept. Game and Fish. 21 pp.
1951. Deer-elk-bear investigations, pp. 16-22.
W-38-R. Jan. Colo. Dept. Game and Fish. 33 pp.
Hammit, H. C., and P. F. Gilbert. 1950.
35-37. In Quart. Prog. Rep., July.

Fed. Aid

Fed. Aid Proj.

Deer-elk-bear investigations, pp.
Colo. Dept. Game and Fish. 80.pp.

Hunter, G. N. 1965. Colorado big game harvest, 1959-1965. Special Mgmt.
Rep., No.1.
Colo. Dept. Game, Fish and Parks. 41 pp.
Jeep, F. T., and L. E. Riordan. 1947. Deer-elk studies. In Quart. Prog.
Rep., Jan. - Part IV. Colo. Dept. Game and Fish. 7 pp.
Kufeld, R. C. 1967. Inventory of range manipulation projects in Colorado,
pp. 63-71. In Game Res. Rep., July - Part I. Colo. Dept. Game, Fish
and Parks. pp. 1-71.
1968. Inventory of range manipulation projects in Colorado.
Game Res. Rep., July - Part I. Colo. Div. Game, Fish and Parks.
pp. 1-121.

In

1969. Inventory of range manipulation projects in Colorado, pp.
299-304. In Game Res. Rep., July - Part III. Colo. Div. Game, Fish
and Parks. pp , 249-393.
1970. Inventory of range manipulation projects in Colorado, pp. 5994. In Game Res. Rep ., July - Part 1. Colo. Div. Game, Fish and Parks.
pp. 1-126.

�50A

McKean, IV. T. 1958. Location, extent and ownershLp of wi.nt er ranges,
pp . 3-23. In Quart. Rep , , July - Part I. Colo. Dept. Game and Fish.
pp. 1-103.
Medin, D. E. 1965. Colorado long range game species management plans,
1965-1975, for mule deer. Colo. Dept. Game, Fish and Parks. Lf pp ,
Minnich, D. IV.
1969. Vegetative response and pattern of deer use following
chaining of pinon and juniper forests. J. Range Mgmt., Abstracts of
Papers, 22nd Ann. Mtg. Amer. Soc. Range Mgmt. 59 pp.
Riordan, L. E. 1948.
Prog. Rep., Jan.

Deer-elk-bear investigations, pp. 14-30.
Colo. Dept. Game and Fish. 49 pp.

Rogers, G. E. 1951. Checking station survey, pp. 32-33.
W-38-R, Jan. Colo. Dept. Game and Fish. 33 pp.

In Quart.

Fed. Aid Proj.
:.:" .

Shepherd, H. R. 1966. San Miguel project, pp. 91-92.
July - Part I. Colo. Dept. Game, Fish and Parks.

In Game Res. Rep.,
pp. 1-93.

Williams, J. E. 1951. Deer-elk investigations, pp. 68-71.
Rep., July. Colo. Dept. Game and Fish. 102 pp.

In Quart. Prog.

1956. Study of mortality factors - winter loss, starvation, disease,
parasites, highway kills, pp. 1~10. In Quart. Prog. Rep., Jan. Colo.
Dept. Game and Fish. 119pp.
lVilliams, J. E., et al. 1952.
Quart. Prog. Rep., April.

Deer-elk investigations, pp. 57-62.
Colo"."
Dept. Game and Fish. 62 pp.

In

PRONGHORN
Anderson, A. E. 1965. Colorado long range management plans, 1965-1975, for
antelope. Colo. Dept. Game, Fish and Parks. 6 pp.
Douglas, G., and C. E. Till. 1950. Antelope surveys and investigations,
Three Year Sum. Rep., 1947-1950, July. Colo. Dept. Game and Fish. 9 pp.
Elliott, R. R. 1957. Antelope investigations, pp. 1,6-8.
Rep., Oct. Colo. Dept. Game and Fish. '22 pp.
1949. Antelope restoration, pp. 19-24, 83-91.
April. Colo. Dept. Game and Fish. 98 pp.

In Quart. Prog.

In Quart. Prog. Rep.,

Hoover, R. L., C. E. Till, and S. Ogilvie. 1959. The antelope of Colorado.
Tech. Bul., No.4.
Colo. Dept. Game and Fish. 110 pp.

L. C. Carlson
October 1971

�51A

SMALL GAflli
M&amp;~L

RESEARCH REFERENCES

WILDLIFE MANAGEMENT UNIT 61

COTTONTAIL RABBIT
Anderson, A. E. 1959. Effects of sagebrush eradication by chemical means
on deer and related wildlife, pp. 113-128. In Quart. Rep., Jan. Colo.
Dept. Game and Fish. 128 pp.
1960. Effects of sagebrush eradication by chemical means on deer
and related wildlife. Job Completion Rep., July. Colo. Dept. Game
and Fish. 72 pp.
1960. Distribution and abundance indices of selected biota in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Univ., Ft. Collins. 130 pp.
1966. An investigation of 2,4-D application to sagebrush within
a mountain shrub-conifer complex, Uncompahgre National Forest. Game
Res. Rep., July - Part III. Colo. Dept. Game, Fish and Parks. pp.
345-42Q. .',
1969. 2,4-D, sagebrush, and mule deer-cattle use of upper w Lnt.e
r
range. Special Rep., No. 21, July. Colo. Div. Game, Fish and Parks.
21 pp.
Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studLes,
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.
Sandfort, W. W. 1951. Cottontail rabbits spring census, pp. 45-47.
Quart. Prog. Rep., July. Colo. Dept. Game and Fish. 102 pp.

In

Shepherd, H. R. 1965. Colorado long range game species management plans,
1965-1975, for cottontail rabbit. Colo. Dept. Game, Fish and Parks.
12 pp.

SNOWSHOE HARE
Shepherd, H. D. 1965. Colorado long range game species management plans,
1965-1975, for snowshoe hare. Colo. Dept. Game, Fish and Parks. 11 pp.

L. C. Carlson
October 1971

�52A

••

»-:

/

NONGAME MAMMAL RESEARCH REFERENCES
WILDLIFE MANAGEMENT UNIT 61

BOBCAT
Boyd, R. J. 1965. Colorado long range game species management plans,
1965-1975, for predators. Colo. Dept. Game, Fish and Parks. 21 pp.
Burget, M. L. 1957. The wild turkey in Colorado, pp. 53-54.
Game and Fish. January. 68 pp.

Colo. Dept.

Burget, M. L., and D. M. Hoffman. 1951. Wild turkey surveys and investigations, pp. 11-29. In Quart. Prog. Rep., Oct. Colo. Dept. Game
and Fish. 117 pp.
Rutherford, W. H. 1965. Colorado long range game species management plans,
1965-1975, for furbearers. Colo. Dept. Game, Fish and Parks. 12 pp.
COYOTE
Boyd, R. J. 1965. Colorado long range game species management plans, 19651975, for predators. Colo. Dept. Game, Fish and Parks. 21 pp.
Burget, M. L. 1949. Wild turkey surveys and investigations. pp. 91-106.
In Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish. 141 pp.
1957. The wild turkey in Colorado.
January. 68 pp.

Colo. Dept. Game and Fish.

Burget, M. L., and D. M. Hoffman. 1951. Wild turkey surveys and investigations, pp. 11-29. In Quart. Prog. Rep., Oct. Colo. Dept. Game and
Fish. 117 pp.
1956. Wild turkey investigations, pp. 107-131. In Quart. Prog.
Rep., July - Part II. Colo. Dept. Game and Fish. pp. 103-138.
Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies,
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 60 pp.
SKUNK
Boyd, R. J. 1965. Colorado long range game species management plans, 19651975, for predators. Colo. Dept. Game, Fish and ?arks. 21 pp.
Burget, M. L. 1957.
Fish. January.

The wild turkey in Colorado.
68 pp.

Colo. Dept. Game and

�53A

Burget, M. L., and D. M. Hoffman. 1951. Wild turkey surveys and investigations, pp. 11-29. In Quart. Prog. Rep., Oct. Colo. Dept. Game and
Fish. 117 pp.
1954. Population limiting studies - disease, accident, predator
factors, pp. 25-30. In Quart. Prog. Rep., July. Colo. Dept. Game and
Fish. 182 pp.
Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies,
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.
Rutherford, W. H. 1965. Colorado long range game species management plans,
1965-1975, for furbearers. Colo. Dept. Game, Fish and Parks. 12 pp.

L. C. Carlson
October 1971

�FURBEARER RESEARCH REFERENCES
WILDLIFE MANAGEMENT UNIT 61

MINK
Denney, R. N. 1950. Fur resource surveys, pp. 75-80.
Rep., July. Colo. Dept. Game and Fish. 80 pp.
Nemanic, W. M. 1942. Fur resource survey, Vol. 2.
and Fish Commission. 6 pp.

In Quart. Prog.

Jan.

Remington, J. D. 1953. Fur harvest survey, pp. 17-23.
Rep., Oct. Colo. Dept. Game and Fish. 60 pp.

Colo. Game

In Quart. Prog.

Rutherford, W. H. 1965. Colorado.long range game species management
plans, 1965-1975, for furbearers. Colo. Dept. Game, Fish and Parks.
12 pp.

MUSKRAT
Denney, R. N. 1950. Fur resource survey, pp. 93-10l.
April. Colo. Dept. Game and Fish-~ 101 pp.
1950. Fur resource survey, pp. 75-80.
Colo. Dept. Game and Fish. 80 pp.

In Quart. Prog. Rep.,

In Quart. Prog. Rep., July.

Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies,
pp. 1-3l. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.
Nemanic, W. M. 1942.
Fish Commission.

Fur resource survey, Vol. 2, Jan.
6 pp.

Colo. Game and

Remington, J. D. 1953. Fur harvest survey, pp. 17-23.
Rep ,, Oct. Colo. Dept. Game and Fish. 60 pp.

In Quart. Prog.

Rutherford, W. H. 1965. Colorado long range game species management plans,
1965-1975, for furbearers. Colo. Dept. Game, Fish and Parks. 12 pp.
WEASEL
Denney, R. N. 1950. Fur resource survey, pp. 93-101.
April. Colo. Dept. Game and Fish. 101 pp.
1950. Fur resource survey, pp. 75-80.
Colo. Dept. Game and Fish. 80 pp.

In Quart. Prog. Rep.,

In Quart. Prog. Rep., July.

�55A

Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies,
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.
Fur resource survey, Vol. 2, Jan.
6 pp.

Colo. Game and

Remington, J. D. 1953. Fur harvest survey, pp. 17-23.
Rep., Oct. Colo. Dept. Game and Fish. 60 pp.

In Quart. Prog.

Nemanic, H. M. 1942.
'Fish Commission.

Rutherford, W. H. 1965. Colorado long range game species management plans,
1965-1975, for furbearers. Colo. Dept. Game, Fish and Parks. 12 pp •.

L. C. Carlson
October 1971

�56A

GANE BIRD RESEARCH REFERENCES
WILDLIFE HANAGEHENT UNIT 61

BAND-TAILED PIGEON
Braun, C. E. 1970. The band-tailed pigeon, pp. 26-29. Colo. Outdoors,
No.5, Sept.-Oct. Colo. Div. Game, Fish and Parks. 48 pp.
1970. Band-tailed pigeon investigations, pp. 151-171.
Res. Rep., Oct. Colo. Div. Game, Fish and Parks. 171 pp.

In Game

Matteson, C. P. 1950. Band-tail pigeons, location and census, pp. 39-40.
Fed. Aid. Proj. 37-R, Oct. Colo. Dept. Game and Fish. 44 pp.
Neff, J. A., and J. C. Culbreath. 1946. Colorado band-tailed pigeon,
pp. 1-24. Fed. Aid Proj. 4-R. Colo. Dept. Game and Fish. 24 pp.

BLUE GROUSE
Aldrich, J. W., and A. J. Duvall. 1955.. Distribution of American
gallinaceous game birds. U.S.D.I. Fish and Wildl. Ser., Circular No.
34. 23 pp.
Anderson, A. E. 1959. Effects of sagebrush eradication by chemical means
on deer and related wildlife, pp. 113-128. In Quart. Rep., Jan. Colo.
Dept. Game and Fish. 128 pp.
1960. Effects of sagebrush eradication by chemical means on deer
and related wildlife. Job CampI. Rep., July. Colo. Dept. Game and
Fish. 72 pp.
1960. Distribution and abundance indices of selected biota in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Univ., Ft. Collins. 130 pp.
1966. An investigation of 2,4-D application to sagebrush within a
mountain shrub-conifer complex, Uncompahgre National Forest. In Game
Res. Rep., July - Part III. Colo. Dept. Game, Fish and Parks. pp.
345-420.
1969. 2,4-D, sagebrush and mule deer-cattle use of upper winter
range. Spec. Rep., No. 21, July. Colo. Div. Game, Fish and Parks.· 21 pp.
Myers, G. T.
season.

1965. Evaluation of a combined blue g rouse-w.i
Ld turkey hunting
Outdoor Facts, No. 27. Colo. Dept. Game, Fish and Parks. 1 p.

Rogers, G. E. 1962. Blue grouse, pp. 121-131.
I. Colo. Dept. Game and Fish. pp. 1-160.

J~ Quart. Rep., July - Part

1963. Development of blue grouse census techniques, pp. 157-168,191.
In Game Res. Rep., Oct. - Part II. Colo. Dept. Game, Fish and Parks.
pp. 109-241.

�57A
1965. Colorado long range game species management plans, 19651975, for blue grouse, sage grouse, sharp~tailed grouse, ptarmigan.
Colo. Dept. Game, Fish and Parks.
16 pp.
1968. The blue grouse in Colorado.
Dept. Game, Fish and Parks.
63 pp.

Tech. Publ. No. 21.

Sandfort, W. W. 1950. Grouse brood counts, p. 37.
Oct.
Colo. Dept. Game and Fish.
44 pp.

Colo.

Fed. Aid Proj. W-37-R,

CHlITCAR
Aldrich, J. W., and A. J. Duvall.
1955. Dis tribution of American
game birds.
D.S.D.1. Ff sh and \\Tildl.Ser. ,Circular
No. 34.

gallinaceous
23 pp.

Bartmann, R. H. 1963. Chukar partridge, pp. 33-40,45-69.
In Game Res. Rep.,
Oct. - Part l. Colo. Dept. Game, Fish and Parks.
pp. 1-108.
1964. Mapping zones and routes for chukar partridge census, pp.
151-178.
In Game Res. Rep., April - Part II. Colo. Dept. Game, Fish
and Parks.
pp. 71-233.
Evans, R. L., and \-1. W. Sandfort.
1957. -Chuka r partridge, pp. 55-67.
Quart. Rep., Oct.
Colo. Dept. Game and Fish.
116 pp.
Manning,
.1111v

D. D.
.- "PRrt:

1962.
L

Chukar partridge,
r.nln.

TlPl")t.. (;RDlP

RM1

pp. 9-16,25-31.
T&lt;'ish.

In Quart. Rep.,

T'~. ]-160.

Miller, J. 1959. Chukar partridge, pp. 99-107,115-124.
Oct. Colo. Dept. Game and Fish.
124 pp.

In Quart.

Miller, J., and W. H. Sandfort.
1959. Chukar production studies,
In Quart. Rep., Jan. Colo. Dept. Game and Fish.
128 pp.
Sandfort, H. W. 1951. Chukar
Wildl. Conf., Jan. 16-19.

In

Re p . ,

pp. 67-73.

census, pp. 22-31.
Spec. Rep., Glenwood
Colo. Dept. Game and Fish.
85 pp.

1952. Game bird survey, p p , 40-56.
Colo. Dept. Game and Fish.
62 pp.

In Quart. Prog.

1952. Nesting and production studies, pp , 93-95.
Rep., Oct. Colo. Dept. Game and Fish.
105pp.

Rep , ,

Sprgs.

April.

-In Quart.

Prog.

1954. Adaptability, survival, and population checks, pp. 23-32.
Quart. Prog. Rep , , April.
Colo. Dept. Game and Fish.
79 pp.
1955. Game bird survey, pp. 17-18.
Colo. Dept. Game and Fish.
94 pp .

In

In Quart. Prog. Rep., Jan.

1957. Chukar partridge, p p , 29-38, 45-51.
In Quart. Prog. Rep.,
July - Part I. Colo. Dept. Game and Fish.
pp. 1-119.
___

1958. Trapping
and transplanting
chuka r partridge, pp. 131-134.
In
Quart. Rep., July - Part II. Colo. Dept. Game and Fish.
pp. 105-181.

�S8A

1959.
Chulcar partridge,
Colo. Dept. Game and Fish.

pp . 59-65,75-77.
128 pp.

In Quart.

Rep , , Jan.

1959. Trapping and transplanting, pp. 33-36. In Quart. Rep.,
April - Part I. Colo. Dept. Game and Fish. pp . 1-116.
1959. Location of chukar partridge release sites, pp. 109-113.
In Quart. Rep., Oct. Colo. Dept. Game and Fish. 124 pp.
1960. Trapping and transplanting, pp. 91-94.
April. Colo. Dept. Game and Fish. 161 pp.

In Quart. Rep.,

1961. Chukar partridge, pp. 119-126,139-155. In Quart. Rep.,
April - Part II. Colo. Dept. Game and Fish. pp. 97-208.
1962. Chukar partridge, pp. 17-23,33-44. In Quart. Rep., July Part I. Colo. Dept. Game and Fish. pp. 1-160.
1965. Colorado long range game species management plans, 19t51975, for chukar partridge. Colo. Dept. Game, Fish and Parks. 18·pp .

.GAl-fEEL
r S QUAIL
,.

Sandfort, 1-1. W. 1950. Game bird surveys, pp. 17-21.
July. Colo. Dept. Game and Fish. 80 pp.

In Quart. Prog. Rep , ,

1952. Nesting and production studies, pp. 93-95.
Rep., Oct. Colo. Dept. Game and Fish. 105 pp.

In Quart. Frog.

1965. Colo. long range game species management plans, 1965-1975,
for Gambe1's quail. Colo. Dept. Game, Fish and Parks. 8 pp.

MOUNTAIN QUAIL
Hoffman, D. M. 1969. Study of mountain quail adaptability, pp. 155-158.
GarneRes. Rep , , April. Colo. Div. Game, Fish and Parks. 181 pp.
1970. Study of mountain quail adaptability, pp. 167-169.
Res. Rep ; , April. Colo. Div. Game, Fish and Parks. 198 pp.

In

In Game

Rogers, G. E. 1965. Appraisal of mountain quail habitat, pp. 217-225. In
Game Res. Rep., April - Part II. Colo. Dept. Game, Fish and Parks. pp.
175-235.
1967. Study of mountain quail adaptability, pp. 215-218. In Game
Res. Rep ; , April - Part II. Colo. Dept. Game, Fish and Parks. pp. 107241.
1968. Study of mountain quail adaptability, pp. 113-116.
Res. Rep., April. Colo. Dept. Game, Fish and Parks. 131 pp.

In Game

�59A

MOURNING DOVE
Nolting, D., et a1. 1952. Population trends, pp. 99-101.
Rep., Oct. Colo. Dept. Game and Fish. 105 pp.

In Quart. Prog.

Sandfort, W. W. 1950. Mourning dove population trends, pp. 41-42.
Aid Proj. W-37-R, Oct. Colo. Dept. Game and Fish. 44 pp.
1951. Mourning dove spring census, pp. 42-43.
July. Colo. Dept. Game and Fish. 102 pp.

Fed.

In Quart. Prog. Rep.,

1951. Mourning dove population trends, pp. 83-84.
Rep., Oct. Colo. Dept. Game and Fish. 117 pp.

In Quart. Prog.

RING-NECKED PHEASANT
Aldrich, J. H., and A. J. Duvall. 1955. Distr:i.::'ution
of American gallinaceous
game birds. U.S.D.I., Fish and Wildl. Ser., Circular No. 34. 23 pp.
Sandfort, W. W. 1950. Game bird survey, pp. 17-21.
July. Colo. Dept. Game and Fish. 80 pp.

Oct.

1950. Pheasant brood survey, pp. 9-12.
Colo. Dept. Game and Fish. 44 pp ,

In Quart. Prog. Rep ;,

Fed. Aid Proj. W-37-R,

1951. Pheasant pre-nesting study, pp. 9-19.
Julv. Colo. Dent. Game and Fish. 102 pp.
1952. Brood survey, pp. 37-41.
Dept. Game and Fish. 105 pp.

In Quart. Prog. Rep.,

In Quart. Prog. Rep., .Oct.

Colo.

1955. Pheasant brood survey, pp. 1-9'. In Quart. Prog. Rep ,, Jan.
Colo. Dept. Game and Fish. 94 pp.
1955. Game bird survey, pp. 65-69. In Quart. Prog. Rep., July Part I. Colo. Dept. Game and Fish. 115 pp.
______
1960. Game bird survey, pp. 37-55,59-65.
Colo. Dept. Game and Fish. 161 pp.

II.

In Quart. Rep., April.

1961. Pre-nesting study, pp. 97-102. In Quart. Rep., April - Part
Colo. Dept. Game and Fish. pp. 97-208.

1963. Programming computer machine analysis of pheasant census and
hunter check data, pp. 213-223. In Game Res. Rep., Oct. - Part II. Colo.
Dept. Game, Fish and Parks. pp. 109-241.
Swop e , H. M. 1965. Colorado long range game species management plans, 19651975, for pheasant. Colo. Dept. Game, Fish and Parks. 54 pp.

�60A

SAGE GROUSE
Aldrich, J. W., and A. J. Duvall. 1955. Distribution of American gallinaceous
game birds. U.S.D.1., Fish and Wildl. Ser., Circular No. 34. 23 pp.
Anderson, A. E. 1959. Effects of sagebrush eradication·by chemical means on
deer and related wildlife, pp. 113-128. In Quart. Rep , , Jan. Colo.
Dept. Game and Fish. 128 pp.
1960. Effects of sagebrush eradication by chemical means on deer and
related wildlife. Job Completion Rep , , July. Colo. Dept. .Game and Fish.
72 pp ,
1960. Distribution and abundance indices of selected biota in
western Colorado before 'and two months after2,4-D appiication. ~f.S.
Thesis. Colo. State Dniv., Ft. Collins. 130 pp.
19E6. An investigation of 2,4-D application to sagebrush within a
mountain shrub-conifer complex, Uncompahgre National Forest. In Game
Res. Rep., July ~ Part III. Colo. Dept. Game, Fish and Parks. pp. 345420.
1969. 2,4-D, sagebrush and mule deer-cattle use of upper winter
range. Special Rep., No. 21, July. Colo. Div. Game, Fish and Parks.
21 pp.
Rogers, G. E. 1963. Programming computer machine analysis of sage grouse
census and hunter check data, pp. 13-32. In Game Res. Rep., Oct. - Part
I. Colo. Dept. Game, Fish and Parks. pp. I-lOB.
1964. Sage grouse investigations in Colorado.
Colo. Game, Fish and Parks. 132 pp.

Tech. Publ. No. 16.

SHARP-TAILED GROUSE
Aldrich, J. W., and A. J. Duvall. 1955. Distribution of American gallinaceous
game birds. D.S.D.I., Fish and Wildl. Ser., Circular No. 34. 23 pp.
Anderson, A. E. 1959. Effects of sagebrush eradication by chemical means on
deer and related ,vildlife, pp, 113-128. In Quart. Rep ,, Jan. Colo.
Dept. Game and Fish. 128 pp.
1960. Effects of sagebrush eradication by chemical means on deer
and related wildlife. Job Completion Rep., July. Colo. Dept. Game and
Fish. 72 pp ,
1960. Distribution and abundance indices of selected biota in
we s t ern Colorado before and t w o months after 2,4-D application. M. S.
Thesis. Colo. State Univ., Ft. Collins. 130 pp.

..

\

�61A

1966. An.investigation of 2,4-D application to sagebrush within
a mountain shrub-conifer complex, Uncompahgre National Forest. In
Game Res. Rep ,, July - Part III. Colo. Dept. Game, Fish and Parks.
pp. 345-420.
1969. 2,4-D, sagebrush and mule deer-cattle use of upper winter
range. Special Rep., No. 21, July. Colo. Div. Game, Fish and Parks.
21 pp.
Rogers, G. E.
1963. Happing sharp--tailedgrouse range, pp. 187-192.··.In
Game Res. Rep., Oct. - Part II. Colo. Dept. Game, Fish and Park.s.
pp. 109-241.
i

______
1965. Colorado long range game species management plans, 19651975, for blue grouse, sage grouse, sharp-tailed grouse, ptarmigan.
Colo. Dept. Game, Fish and Parks. 16 pp.
1965. S~~rp-tailed grouse, pp. 209-214. In Game Res. Rep., AprilPart II. Colo. Dept. Game, Fish and Parks. pp , 175-235.

Nov.

1969. The sharp-tailed grouse in Colorado.
Colo. Div. Game, Fish and Parks. 94 pp.

Tech. Publ., No. 23,

Rogers, G. E., and F. D. Stearns. 1964 .. Sharp-tailed grouse, pp. 291-341.
In Game Res. Rep., April - Part III. Colo. Dept. Game, Fish and Parks.
pp. 235-397.

HATERFOWL
Funk, H. D. 1970. Waterfowl kill survey, pp. 19-40.
Oct. Colo. Div. Game, Fish and Parks. 17lpp.

In Game Res. Rep.,

Grieb, J. R. 1959. Haterfowl kill survey, pp. 145-155. In Quart. Rep.,
July - Part II. Colo. Dept. Game and Fish. pp. 145-238.
1960. Haterfm.Jl kill survey, pp. 93-103.
Colo. Dept. Game and Fish. 103 pp.

In Quart. Rep., Oct.

1961. Haterfowl kill survey, pp. 9-21.
Dept. Game and Fish. 103 pp.

In Quart. Rep., Oct.

Colo.

1962. Waterfowl kill survey, pp. 15-25;
Dept. Game and Fish. 137 pp.

In -Quart. Rep ., Oct.

Colo.

1965. Haterfowl kill survey, pp, 23-40.
Colo. Dept. Game, Fish and Parks. 199 pp ,

In Game Res. Rep., Oct.

1966. Waterfowl kill survey, pp. 17-31.
Colo. Dept. Game, Fish and Parks. 187 pp.

In Game Res. Rep., Oct.

1967. WaterfoHl kill survey, pp. 15-30.
Colo. De~t. Gmnes, Fish and Parks. 103 pp.

In Game Res. Rep., Oct.

�62A

Oct.

1968. Waterfowl kill survey, pr. 15-31. In Game Res. Rep.,
Colo. Div. Game, Fish and Parks. 135 pp.

Oct.

1969. Waterfowl kill survey, pp. 39-56. In Game Res. Rep.,
Colo. Div. Game, Fish and Parks. 162 pp .

Grieb, J. R., and G. N. Hunter. 1963. Waterfowl kill report, pp. 1-20.
In Game Res. Rep., July. Colo. Dept. Game, Fish and Parks. 51 pp.
1964. Waterfowl kill survey, pp. 65-78.
Colo. Dept. Game, Fish and Parks. 78 pp.

In Game Res. Rep., July.

1968. Wetlands of Colorado, Tech. Pub1., No. 22, March.
Dept. Game, Fish and Parks. 89 pp.

Colo.

WILD TURKEY
Aldrich, J. W., and A. J. Duvall. 1955. Distribution of American gallinaceous
game birds. U.S.D.L, Fish and Wildl. Ser., Circular No. 34. 23 pp.
Burget, M. L. 1946.
Fish. 21 pp.

Colorado wild turkey.

Vol. II.

Colo. Dept. Game and

1948. Wild turkey surveys and investigations, pp. 47-62.
Prog. Rep , , Oct. Colo. Dept. Game and Fish. 80 pp.

In Quart.

1949. Wild turkey surveys and investigations, pp. 91-106.
~rog. Kep.) Oct. Colo. Dept. Game and Fish. 141 pp.

In Quart.

1949. Wild turkey development, pp . 3-18.
Colo. Dept. Game and Fish. 98 pp.

In Quart. Prog. Rep , , April.

1951. Wild turkey rehabilitation in Colorado, pp. 7-17. Spec. Rep.,
Glenwood Sprgs. Wildl. Conf., Jan. 16-19. Colo. Dept. Game and Fish.
85 pp.
1956. Brood counts, pp. 29-38.
Dept. Game and Fish. III pp.

In Quart. Prog. Rep., Oct.

Jan.

1957. The wild turkey in Colorado.
68 pp.

Jan.

1959. Wild turkey development, pp. 25-26,33-38.
Colo. Dept. Game and Fish. 128 pp.

Colo.

Colo. Dept. Game and Fish.

In Quart. Rep.,

1959. Wild turkey development, pp. 71-84. In Quart. Rep., April _
Part I. Colo. Dept. Game and Fish. pp. 1-116.
1960. Wild turkey development, pp. 1-42.
Colo. Dept. Game and Fish. 110 pp.

Oct.

In Quart. Rep., July.

1961. l.JiJ.d
turkey development, pr. 23-43,51-55.
Colo. Dept. GRme and Fish. 103 pp.

In Quart. Rep ,,

�63A
1963. Mapping of ",i1d, turkey ,range, pp.' 71-85.
April.
Colo. Dept. Game and Fish.
109 pp.

In Game Res. Rep.,
.... :.
"

Burget, M. L., and C. Ford.
1951. Wild turkey development, pp. 1-7.
Quart. Prog. Rep., April.
Colo. Dept.Game and Fish.
64 pp.
Burget, M. L., and D. M. Hoffman.
1950. Wild turkey
gations,
Three Year Sum. Rep , , 1947-1950, July.
Fish.
4 pp.
1950. Wild turkey investigations.
Colo. Dept. Game and Fish.
18 pp.

In

surveys and investiColo. Dept. Game and

Fed. Aid Proj.

W-39-R, ..oct.

In

1951. \H1d turkey surveys and investigations,
pp. 11-29.
Quart. Frog. Rep, , Oct.
Colo. Dept. Game and Fish.
117 pp.
1952. 1.Ji1d turkey investigations,
pp , 1-5.
April.
Colo. Dept. Game and Fish.
62 pp.

In Quart.

-w

Prog.

1952. Hild turkey investigations,
pp. 1-9,11-14,21-24.·
Frog. Rep., Oct.
Colo. Dept. Game and Fish.
105 pp.

Rep. ,

In Qua-rt.

33.

1953.
Trapping and transplanting
of Herriam's ",ild turkey,
pp. 19In Quart. Prog. Rep, , April.
Colo. Dept. Game and Fish.
99 pp.

July.

1953. Hild turkey survival
checks, pp. 1-6.
Colo. Dept. Game and Fish.
129 pp.

April.

Colo. Dept.

Game and Fish.

In Quart.

Frog.

79 pp.

1954. Hild turkey s urv Lva l checks, pp , 13-24.
July.
Colo. Dept. Game and Fish.
182 pro

In Quart.

1954.
Food studies,
pp. 39-57.
Dept. Game and Fish.
III pp.

Rep.,

Part

Rep.,

In Quart.

Frog.

1955.
Survival checks, pp . 93-99.
In Quart.
I.
Colo. Dept. Game and Fish.
pp. 1-115.

1955. Hi1d turkey investigations,
Rep., Oct.
Colo. Dept. Game and Fish.

Frog.

pp. 1-8,13-19.
108 pp.

Frog.

Oct.

Rep.,

Colo.

July

In Quart.

Rep.,

-

Prog.

1956.
Trapping and transplanting
Herriam' s w i Ld turkey,
pp. 29-38.
In Quart. Prog. Rep., April.
Colo. Dept. Game and Fish.
102 pp.
1956. Wild turkey investigations,
pp. 107-121,123-131.
In Quart.
Prog. Rep., July - Part II.
Colo. Dept. Game and Fish.
pp. 103-138.
1957. \.Ji1d turkey development, pp. 45-70.
Colo. Dept. Game and Fish.
108 pp.

In Quart.

Rep. , April.

1957.
Brood counts,
Game and Fish.
116 pp.

Rep.,

Colo.

pp. 69-86.

In Quart.

Oct.

Dept.

�1958. Hild turkey investigations, pp. 1/-17:-169.
112 Quart. Rep.,
July - Part II. Colo. Dept. Game and Fish. pp. 105-181.
Evans, R. 1. 1963. Relationships of the environment to w i Ld turkey productivity and distribution, pp. 71-85. In Game Res. Rep., April.
Colo. Dept. Game and Fish. 109 pp.
Hoffman, D. H. 1965. Colorado long range game species management plans,
1965-1975, for wild turkey. Colo. Dept~ Game, Fish and Parks. 11 pp.
Hoffman, D. M., and M. L. Burget. 1959. Study of food plot usage by wild,
turkey, pp. 27-32. In Quart. Rep., Jan. Colo. Dept. Game and Fish ..
128 pp.
Hunter, G. N. 1965. Colo. big game harvest, 1959-1965.
No.1.
Colo. Dept. Game, Fish and Parks. 41 pp.

Special Mgmt. Rep ,,

Myers, G. T. 1963. Relationships of the productivity and distribution of
w LLd turkeys on the Uncompahgre Plateau t c the environment, pp , 173-178.
In Game Res. Rep., Oct. - Part II. Colo. Dept. Game,Fish and Porks.
pp. 109-241.
1964. Relationships of the productivity and distribution of wild
turkeys on the Uncompahgre Plateau to the environment, pp. 243-280. In
Game Res. Rep ;, April - Part III. Colo. Dept. Game, Fish and Parks.
pp. 235-397.
1965. Evaluation of a combined blue grouse-wild turkey hunting
season. Outdoor Facts, No. 27. Colo. Dept. Game, Fish and Parks. I p.
1965. Reliability of turkey sex and age ratio data based on hunter
report card returns. Outdoor Facts, No. 29. Colo. Dept. Game, Fish
and Parks. 2 pp.
1965. Relationships of the productivity and distribution of wild
turkeys on the Uncompahgre Plateau to the environment, pp. 91-174. In
Game Res. Rep., April - Part I. Colo. Dept. Game, Fish and Parks.
pp. 1-174.
1966. Relationships of the productivity and distribution of w i Ld
turkeys on the Uncompahgre Plateau to the environment, pp. 69-84. In
Game Res. Rep., April. Colo. Dept. Game, Fish and Parks. 192 pp.
1967. Wild turkeys, pp. 167-182,205-211. In Game Res. Rep.,
April - Part II. Colo. Dept. Game, Fish and Parks. pp. 107-241.
1968. lhld turkeys, pp. 71,99-110. In Game Res. Rep., April.
Colo. Dept. Game, Fish and Parks. 131 pp.
1969. IHld turkeys, pp. 107,153.
Div. Game, Fish and Parks. 181 pp.

In Game Res. Rep ,, April.

1969. The topic is turke~, pp. 30~34. Colo. Outdoors, No.5,
Sept.-Oct. Colo. Div. Game, Fish and Parks. 48 pp.

Colo.

�65A

1970. Wild turkeys, pp. 161,165. In Game Res. Rcp ,, April.
Colo. Div. Game, Fish and Parks. 198 pp.
Sheats, C. C. 1970. My experience with the turkey, pp. 30-31. Colo.
Outdoors, No. 30, May-June. Colo. Div. Game, Fish and Parks. 48 pp.

L C. Carlson
October 1971

�66A

GENERAL MAMMALIAN REFERENCES
WILDLIFE MANAGEMENT UNIT 61

Armstrong, D. M.

1972.

Distribution of mammals in Colorado.

Kansas Printing Service, Lawrence.

Anthony, H. E.

1928.

Putnam's Sons~

Hall, E. R., and K. R. Kelson.
Ronald Press Co.

Lechleitner, R. R.
Co.

Boulder, Colo.

1959.

The mammals of North America.

2 Vols.

1162 pp.

~.J'ild
mammals of Colorado.

Pruett Publ.

254 pp.

Miller, G. S., Jr., and R. Kellogg.

205.

G. P.

674 pp.

New York.

1969.

recent mammals.

4l5pp.

Field book of North American mammals.

New York.

Univ.

1955.

List of North American

Smithsonian Institution.

Hash. D. C.

Bulletin

954 pp.

Palmer, R. s.

1954.

City, N. Y.

Warren, R. R.

Doubleday &amp; Co., Inc.

Garden

384 pp.

1942.

Norman, Okla.

The mammal guide.

The mammals of Colo.

Univ. of Okla. Press,

330 pp.

L. C. Carlson
February 1972

�67A

GENERAL AVIAN REFERENCES
WILDLIFE ~~NAGEMENT UNIT 61

Bailey, A. M., and R. J. Niedrach. 1965.
of Nat. Rist., Den., 2 Vol. 895 pp.

Birds of Colorado.

1967. Pictorial checklist of Colorado birds.
Rist., Den. 168 pp.
Davis, W. A.
61 pp.

1969.

Birds of western Colorado.

Den. Mus.

Den. Mus. of Nat.

Colo. Field Ornithologists.

Kortright, F. A. 1943. The ducks, geese, and swans of North America.
The Amer. Wi1dl. Institute, Wash. D. C. 476 pp.
Linduska, J. P. 1964. Waterfowl tomorrow.
Wi1d1., Wash. D. C. 770 pp.
Pearson, T. G. 1940.
N. Y. 289 pp.

Birds of America.

U.S.D.I. Bur. Sp. Fish and

Garden City Publ. Co., Inc.,

Peterson, R. T. 1941. A field guide to western birds.
Co., Boston. 240 pp.
Robbins, C. D., B. Bruun, and R. S. Zim.
Golden Press, N. Y. 340 pp.

1966.

Roughton Mifflin

Birds of North America.

L. C. Carlson
February 1972

�68A

---CENSUS AREAS AND ROUTES - WILDLIFE

MANAGEMENT

UNIT 61

BLACK BEAR
Black bear are counted
ratio censuses.

in Unit 61 incidental

to aerial deer pre-hunt

Data are on file at both the Division's

and Denver offices

(Burdick pers.

Southwest

sex

Region

comm. 1972).

BLUE GROUSE
A blue grouse

census route was established

by Rogers

(1963) in 1961 and

repeated

in 1962 and 63.

and 62.

This route was 62.5 miles long and was divided

of approximately
traversed,

12.5 miles each.

elevation,

the feasibility

It lay along the Divide Road between

The sections

and water proximity.

of counting

into five sections

varied as to range type

The purpose was to determine

birds on specific

counts were made all three years;

Units 61

production

routes.

Brood

IIhootingltcounts were attempted

in 1962

along parts of the same route.

Counts on Roger's
currently

route were discontinued

after 1963, and none are conducted

in Unit 61 (Burdick pers. comm. 1972).

CHUKARS
The Gateway

Census Area is in part within Unit 61.

141 east and north of Gateway
Report

for April 1964.

vegetated
dog.

for observing

About 4.3 miles of Highway

are mapped and described

Those areas \-Thichwere too wide or too densely
the birds Here covered on foot with the aid of a

Counts were made at least three' times in August.

for the 1963 count may be found in the report cited.
personnel

have continued

that office

in the Federal Aid

Tabulated
SouthHest

this count and more recent records

as well as in the Denver office.

results

Regional

are available

in

�69A

ELK
Elk numbers

are currently

recorded

incidental

to aerial deer counts in

Unit 61 (Burdick pers. comm. 1972).

MULE DEER
A helicopter

aerial trend area, established

continuously

since then .. The area is called the Uravan

located between
separate

in 1967, has been counted

Blue and Campbell Creeks near Unit 61's north end (under

cover see 1/2-inch

scale map overlay).

Pellet

group count data, discussed

in a foregoing

MAMMAL

DISTRIBUTION

constituted

deer population

AND ABUNDANCE,

locations,

RING-NECY~D
Aid Project W-37-R,

sex ratios,

spring

personnel

This included

procedures

and run annually

around Nucla,
up-to-date.

Roadside

from 1955-1965 with appropriate

sheets being produced

and filled out.

regions.

Finally,

was prepared

routes

field forms

a manual

com-

for each major

For Unit 61 all these counts were

and they have been continued by Southwest

Maps of these routes and tabular records

may be found in the Southwest

for

counts for winter

crowing counts, and summer brood counts.

area within management

established

tables

(B. D. Baker unpublished).

these data, together with instructions,

pheasant

More detailed

the Game Bird Survey, developed

census record in the early 50's.

piling

to establish

PHEASA1~T

a pheasant

and summary

an older attempt

data, and results of this work are available

in the files of Federal Aid Project W-IOI-R

were selected

section on BIG GA}lli

trend data for Unit 61, however meagre.

and maps concerning

Federal

trend and is

Regional

of results

Region and Denver office of the Division.

�70A

SHARP-TAILED
Counts on sharp tails on dancing

grounds were made by Rogers

Unit 61 in the Outlaw 11esa vicinity
on the Tongue,
Location

at Mud Springs,

(Cold Springs),

and at Cillilands

(1969) in

on Horsefly

Creek,

in 1964 and 1965.

maps for three of these sites as well as lists of all by

county appear in Rogers

Dancing
pers.

CROUSE

ground

(1969).

counts are not currently

being made in Unit 61 (Burdick

comm. 1972).

WILD TURKEYS
Myers

(1967 and pers.

at 'which winter

comm. 1971) reports having

feed stations were located

(approx S.l, T.49N.,

R.18W).

Creek

3. Hill Ranch

files of the Southwest
Aid Report

continued

2. Cottonwood

R.12W.).

Regional

cited.

R.13W.),

1. Blue Creek

and 4. McKenzie

for these counts are in

the Denver office,

The feed stations

and in the

and counts have been dis-

since 1967 •.

Burdick

(pers. comm. 1972) reported

Sanborn

Park

(Clay Creek)

that

C. T.

1967.

April - Part 2.

wf.n t er

feed station

counts in

are currently being done.

Literature

Hyers,

Records

office,

as follows:

In

Creek (approx. S.36, T.47N.,

(approx. S.30, T.46N.,

(approx. S.3l, T.45N.,

Federal

feed stations

counts of turkeys were made from 1963 through 1967.

Unit 61 the principle

R.14W.).

established

Cited

Came Bird Survey, pp. 167-182.

In Quart. Rep.,

Colo. Div. Came, Fish and Parks.

(processed).

�7lA

Rogers, G. E.

1963.

Game bird surveys, pp. 157-168.

Rep., Oct. - Part 2.

1968.

Colo. Dept. Game, Fish and Parks.

The blue grouse in Colorado.

Dept. Game, Fish and Parks.

1969.

In Fed. Aid Quart.
(processed).

Tech. Publ. No. 21.

Colo.

63 pp.

The sharp-tailed grouse in Colorado.

Colo. Div. Game, Fish and Parks.

Tech. Publ. No. 23.

94 pp.

B.D. Baker and W.T. HcKean
February 1972

�72A

HABITAT

Detailed
Kufeld

RESTORATION

information

PROJECTS

on projects

at the Research

details

available.

So, depending

Mr. Kufeld
compiled

Simply make your requests

on only one project are needed,

for restoration

UNIT 61

are kept in the office of Mr. Ron

could be conveniently

days work.

If specific

requests

MANAGEMENT

Center, Fort Collins.

that all Unit 61 projects
by about one-half

- WILDLIFE

informed

and summarized
directly

information

to him.

that, too, is readily

upon what data are needed, we recommend
project

us

be sent directly

that

to Mr.

Kufeld.

The above approach
McKean

and I take the attitude

data retrieval
habitat

to handling

this information

that a central repository

system have been established

data current

is alterable,

and easily available.

and efficient

to keep rapidly
Another

but Mr.

middleman

not be necessary.

B. D. Baker
December 1971

changeable
should

�73A

}~NAGEMENT

A.

PROBLEMS

CHECKLIST

- WILDLIFE

~AGEMENT

UNIT 61

Present
1.
Boyd

Deer-auto

collision

losses, State Highway

(1961, 1962, and 1964) reported

killed by auto collisions
10 miles of Highway

Bear Canyon confluences.
drain problem
partially

that a minimum

during the 1961-63

141 in Unaweep

These losses represent

will also increase.

As vehicular
Highway

Assistance

W-38-R

who are working

personnel

the Ute Creek and
a continuing

yearlong

residents

traffic increases,

structures

are needed here.

of 17 deer were

period in approximately

Canyon between

on deer which are partially

migratory.

141.

designed

can be solicited

herdand

this problem

to prevent

collisions

from Federal Aid Project

on similar problems on Interstate

High-

way I 70 and elsewhere.

2.

Need for techniques

to provide

population

of selected wildlife

densities

reliable

periodic

indices

to annual

species by area or

drainage.

The numerous,
present
and elk.

abrupt topographic

special

sampling

problems

Also, census techniques

changes from ridge top to creek bottom
for pellet group count work
for many other wildlife

for deer

species

are

needed.

3.

Agricultural

crop damage.

Alfalfa

standing

crops near Redvale

and Nucla are currently

apparent

damaging

use by deer (Division pers. comm. 1972).

receiving

�74A

B.

Future.
1.

San Miguel Project,

U. S. Bureau of Reclamation.

The San Miguel Project has bee~ proposed by the U. S. Bureau of
Reclamation

to control and better utilize stream flow of the San

Miguel River

(U.S. Bur. Recl. 1966).

An estimated

mule deer range would be lost when project
(U.S. Bur. Sport Fish. and Wildlife

25,000 acres of

lands are cultivated

1966, 1968).

2,752 acres would be lost through inundation

An additional

of reservoir

sites.

Most of these losses of deer range would occur in Wildlife
ment Unit 70 (San Miguel),
in the San Miguel
confluence

but inundation

Canyon between

at the Saltado

Manage-

Reservoir

Placerville

and the Specie Creek

and canal and control structures

in the Mail Box Park

area north of Norwood would

directly

affect wildlife

habitat" in

Unit 61.

In addition
construction,
would

to direct losses caused by reservoir
expansion

also increase

strative

problems

Several methods
suggested
1968).

big-game

cropland

for mitigation

in Mail Box Park

damage to crops and attendant

of losses of big-game

by the U. S. Bureau of Sport Fisheries

Acquisition

control

admini-

for the Division.

of lands for intensive

would be the principal
scheme,

of agricultural

and water

method

either singly or in combination

and Wildlife

management

of mitigation.

range have been
(1966,

by the Division

Nevertheless,

that·

with other contingent

proposals,

�75A

very probably

could not effect full compensation

loss of habitat

2.

Tabeguache

that the project will cause.

Watershed

Project,

U. S. Department

This project

is a potential

improvement

of irrigation

distribution

Cooperative

Ditch Company

CU.S.D.A. 1971).

project

for the permanent

upon wildlife

PL566 activity

of Agriculture.

that would mainly

systems

served by the Colorado

Possible

have not been evaluated

involve

impacts of the

but are assumed

to be

minor.

3.

Naturita

Watershed

Project,

U. S. Department

This proposed

project

involves

a very small acreage and is listed

mainly

4.

as a reminder

of Agriculture.

that plans exist for its development.

Coal strip mining,

There is an estimated

Nucla area.
reserve of three million

tons of Grade A bi-

tuminous

coal in the Dolores River Basin.

Strip-mined

supplies

the power to generate

by the Colorado-Ute

electricity

Association

plant near Nucla.

power would

require an estimated

coal of 60,000 tons a year

Potential
doubling

CU.S.D.A.

increase

coal now

in demand for electric

of current production

1971).

Electric

Depending

rate of

upon where new

strip mines would be located and how much effort is made to adequately
rehabilitate

resulting

spoils areas, wildlife

habitat might well suffer

degradation.

5.

Housing

developments

in existing

natural wildlife

habitat.

�76A

Literature Cited

Boyd, R. J.
82.

1961.

Study of deer losses on Colorado hLghways , pp , 75-

In Fed. Aid Quart. Rep., April - Part I.

Fish.

96 pp.

1962.

Colo. Dept. Game and

(processed).

Study of deer losses on Colorado highways, pp. 93-100.

In Fed. Aid Quart. Rep., October.

Colo. Dept. Game and Fish.

69 pp.

(processed).

1964.

Study of deer losses on Colorado highways, pp. 36-41.

Fed. Aid Game Res. Rep., April - Part I.
Parks.

69 pp.

February.

1966.

San Miguel Project, Colorado.

U. S. Dept. Int.-,Reg. 4 Bur. Reel., Salt Lake City.

63 pp. + 15 p. Addenda.

U. S. Bureau Sport Fisheries and Hildli£e.
Colorado.

Colo. Dept. Game, Fish and

(processed).

U. S. Bureau of Reclamation.
Feasibility Rep.

In

1966.

Bur. Sport Fish. and Wild1. Rep.

and Wildl. Serv., Albuquerque, June 30.

1968.

San Miguel Project, Colorado.

San Miguel Project,

U. S. Dept. Int., Fish

19 pp. + 2 plates (Maps).

Bur. Sport Fish, and Wildl.

Rep. Supply.

U. S. Dept. Int., Fish and Wildl. Serv., Albuquerque.

February 13.

5 pp. + I plate (Map).

U. S. Department of Agriculture.

1971.

Water and related land resources,

Dolores River Basin in Colorado and Utah.

Coop. Study Rep. of Colo.

Hater Conserv. Board and U.S.D.A. Econ. Res. Service, Forest Service,
and soil Conserv. Service.

February.

Denver.

(pp. val:'.
by chapter) +

sep. bound Soil Conserv. Service Appendix 1 - Hatershed Investigations Reps.

W. T. HcKean and B. D. Baker
FphY--ll;:,r'T

1 Q7?

�APPENDIX

B

WILDLIFE MANAGEMENT UNIT 62 (E. UNCOMPARGRE)
Delta, Mesa, Montrose, and Ouray Counties, Colorado

Information on Unit Description,
Landownership; Land Use, Human
Population; Wildlife Species
Checklists, Harvests, Seasons,
Narrative and Map Descriptions
of Distribution and Abundance;
and Other Allied Data. To:
March 15, 1912.
Compiled by: 'W. T. McKean and
B. D. Baker

�IB

DESCRIPTION

Boundaries

- WILDLIFE

J:../.--"Thatportion

No. 141 from the Uncompahgre

County

of Mesa County south and east of State Highway
Plateau

divide to the tmvn of Whitewater,

No. 50 from the town of Whitewater

and west of U.S. Highway

line, and east of the Uncompahgre

Plateau

Delta County south and west of U.S. Highway
line to the Montrose-Delta

County

line; that portion

No. 50 from the Delta-Montrose

of U.S. Highway

No. 550 from Montrose

north and east of the Uncompahgre

its intersection

divide;

of Montrose

of
County

County west

County line to Montrose,

divide;

County

that portion

No. 550 from the Montrose-Ouray

with State Highway

to the Mesa-Delta

that portion

to the Ouray-Montrose

Plateau

south

No. 50 from the Mesa-Delta

of U.S. Highway

County west of U.S. Highway

1/

UNIT 62 (E. UNCOMPAHGRE)

MANAGEMENT

west

line, and

of Ouray

County

line to

No. 62 and north and west of State Highway

No. 62."

Size.--Unit
size.

62 was determined

A planimeter

Uncompahgre

National

to be 1,334.1 square miles or 853,824 acres in

was used to obtain the acreage

from a 1/4" = 1 mile scale

Forest map and a 1/2" = 1 mile scale Colorado

Division

Mesa County map.

Physical

Features.--Unit

62 coincides

approximately

half of the Uncompahgre

Plateau.

uplift

the Canyon Lands section

that lies within

Geologically,

with

State Highway

the northeastern

the plateau

is a broad

of the Colorado

one-

structural

Plateaus

1/ Legally termed "Big Game Management Unit" (see footnote 2/ below) but
tentatively
called "Wildlife Management Unit" here to include broader aspects
of animal life present.
2/ Colo. Game, Fish and Parks Div. Laws and Regulations Hdbk., 1970 (pp. 12-13,
Chap. 2 - Big Game).

�2B

physiographic

province.

granites

to Mesozoic

include

a comparatively

mesas

Parent materials

sandstones,

between

sharply-rimmed

undulating
farmland

toward the Uncompahgre

that support

a substantial

flood plains

1962, 1971).

The Uncompahgre

and Gunnison

Principal

Rivers,

acreage

at Unit 62's eastern

tributaries

of the rivers,

Creeks.

No sizable

lakes or reservoirs

Depending

upon specific

Descriptions

and northern

peri-

Tributary

streamflow

Escalante,

Monitor,

is northeastwardly.

exist in Unit 62.

portion(s)

Montrose.

(U. S. Dept.

in order from north

Dry, and Dallas

Work Unit personnel

of irrigated

They both flow in a northwestwardly

Roubideau,

Service

Rivers

and gentle to moderate

East, Big and Little Dominguez,

information

features

and Gunnison

to south, are as follows:

needed,

Terrain

south and west of Delta and Montrose

are the unit's main streams.
here.

crystalline

gentle to strong sloping parallel

canyons,

at lower elevations

direction

summit,

and shales.

or steep-sided

of Agriculture

meter,

siltstones,

flat plateau

that run northeastwardly

vary from Precambrian

can be.obtained
located

of the unit where soils data might be
by contacting
at offices

U. S. Soil Conservation

in Grand Junction,

and a map of broad soil classifications

in a previously· cited report of the U. S. Department

Delta,

or

can be found

of Agriculture

(1962).

�Climate.--The

climate

wide extremes

resulting

precipitation

varies

Montrose,

to between

of Unit 62 is typical
from variations

and short duration

tation is fairly uniform,

National

Wea~her

spectively,

higher

Service

long growing

records

ward,

depending

seasons

from reports
Department

generally

receiving

a greater

and at Montrose

present,

re-

on the Gunnison

canyon bottoms

of garden,

exposure,

seasons
River

also have comThese warmer

field, and orchard plants

site frost-free

periods

and other factors,

for

scale down-

to about

30 days

divide.

information

of Commerce

Tributary

and mesa-top

was obtained,

(Continuing).

aforementioned

National

station

close to but outside

southeast

of precipi-

distribution

season near Whitewater

of the U. S. Department

fairly

Seasonal

for much of their lower portions.

upon elevations,

climatic

Cloudburst

of 51.00 F and 49.60 F an~ growing

selection

Canyon-rim

or less along the plateau

Preceding

crest.

areas and

for near Delta

Growing

permit wider

crop production.

Mean annual

occur over localized

elevations

to be about 175 days.

microc1imates

features.

in the form of snowfall.

of 146 days and 152 days.

paratively

conditions.

mean annual temperatures

is estimated

slope, with

20 and 25 inches along the plateau

result from summer convective

in winter

in topographic

western

from about 7.75 inches at Delta, about 9 inches at

storms of high intensity

proportion

of Colorado's

Weather

of Grand Junction.

and in some cases interpreted,

of Agriculture

(1962, 1971) and U. S.

The latter agency is parent bureau

Service which

also administers

of the unit's perimeter

to the

a cooperating

six miles

east-

�4B

Literature Cited
U. S. Department of Agriculture.

1962.

Gunnison River Basin - Colorado.

Water and related land resources,

Coop. Study Rep. of Colo. Water

Conserv. Board and U.S.D.A. Econ. Res. Service, Forest Service, and
Soil Conserv. Service.

1971.

November.

Denver.

103 pp.

(processed).

Water and related land resources, Dolores River Basin in

Colorado and Utah.

Coop. Study Rep. of Colorado Water Conserv. Board

and U.S.D.A. Econ. Res. Service, Forest Service, and Soil Conserv. Service.
February.

Denve r,

(processed).

U. S. Department of Commerce.

(Continuing). Climatological data, Colorado.

Monthly and Annual Summaries.
Asheville, N. C.

National Climatic Center, Fed. Bldg.,

28801.

B. D. Baker
August 1971

�5B

LANDOWNERSHIP STATUS - WILDLIFE MANAGEMENT UNIT 62

Bureau of Land Management

319,240

Acres

Forest Service

255,841

Acres

Private lands excluding lands within
National Forest boundaries

249,170

Acres

Private lands within National Forest
boundaries

14,923

Acres

Colorado Division Game, Fish and Parks

7,299

Acres

o

Acres

160

Acres

1,280

Acres

847,913

Acres

Other Federal lands including Park
Service, Indian and Military Reservations, and Bur. Sport Fisheries and
Wildlife
State Land Board Administration lands
Municipal lands
Total

*

(1324.9 sq. miles)

* Total land acreage differs from total in Unit Description Section
(853,824 A. or 1334.1 sq. miles). Forest Service lands and private
lands within the National Forest were derived by planimetering on
2" = 1 mile Uncompahgre National Forest township plat maps. B.L.M.
lands, Game, Fish and Parks lands and other private land acreages
resulted from planimetering and estimating on 1/2" = 1 mile Uncompahgre
National Forest map and 1/2" = 1 mile B.L.M. Glade Park Recreation map.
Municipal lands include those parts of Delta, Olathe, and Montrose
within Unit 62.

w. T. McKean and L. C. Carlson
September 1971

�'J

6B

LAND USE STATUS - WILDLIFE MANAGEMENT UNIT 62

Irrigated croplands

72,428 Acres

Non-irrigated croplands

2,229 Acres

Grasslands

20,708 Acres

Sagebrush

72,710 Acres

Brush

165,404 Acres

Woodland-Pinon-juniper,
oakbrush

297,404 Acres

Aspen

38,014 Acres

Commercial timber

131,420 Acres

River bottom vegetation

8,674 Acres

Miscellaneous

38,922 Acres
Total

847,913 Acres

*

(1324.9 sq. miles)
*Cover type categories and acreages were derived from planimetering on
1/2" = 1 mile S.C.S. Land Use and Cover Type Maps, 1954, for Delta,
Mesa, Montrose, and Ouray counties. Computations were 3% over landownership status total acreages. An adjustment was made to each
category so that the total would conform to the landownership figure.

L. C. Carlson
November 1971

�7B

HUMAN POPULATION

Since Unit 62 involves

- WILDLIFE

portions

counties

and unit bounding

Montrose,

and Ridgway,

Bureau

(Table 1).

of the Census

Instead

of specifically

in total for the counties

This analysis

differs

By comparing

as being rural in character.

populations

of Delta and Ouray counties
of Montrose

of municipalities

showed

about 27% to exhibit
supported

cities

being

as being urban

rural.

1970 figures with those of 1960 in Table 1, it is apparent

that populations
the population

from those of the

cities and towns are considered

of size, with remainder

popu-

and their

in that they consider

and towns of less than 2,500 population

regardless

applicable

urban and rural data and trends

and other sources

Here, all incorporated

and Ouray

split the towns of Delta, Olathe,

determination

for 1960 and 1970 are prese~ted

UNIT 62

of Mesa, Delta, Montrose,

highways

lations would be difficult.

municipalities

MANAGEMENT

County increased

are evident

of Montrose

and 1970, only slightly
County.

Additional

counties

are presented

Montrose's

increase

County dropped noticeably

analyses

of populations

U. S. Department

of Agriculture.

land resources,

Dolores

Light

between

1971.

in Mesa and Montrose

Water Conserv.

February.

(processed).

report:

Water and related

River Basin in Colorado

Forest Service,

1960

in Ouray

4-8 of the following

Econ. Res. Service,
Denver.

evidently

County.

and gained slightly

on pages 4-5 through

Coop. Study Rep. of Colorado

gaining

for both Delta and Ouray counties.

in Delta County,

detailed

while

Populations

with Montrose

of total urban growth for Montrose

losses of urban populations
The rural population

trend.

slightly

very slightly.

changes both directions,

the strongest

the increase

decreased

and Utah.

Board and U.S.D.A.

and Soil Conserv.

Service.

�8B

Table 1. Populations and trends of counties and municipalities involved, in
part, with Unit 62 -1/ •
POEulation
2/
County -

1960

1970

Change

Inc. Cities
Population
and
%
Towns
1960
1970 Change

Delta

15,602 15,286

-2.0

Delta

3,832

3,602

- 6.0

Paonia

1,083

1,161

+ 7.2

Hotchkiss

626

507

-19.0

Cederedge

549

581

+ 5.8

Crawford

147

171

+16.3

Totals

6,237

6,022

- 3.4

Montrose

5,044

6,413

+27.1

Naturita

979

820

-16.1

Nucla

906

949

+ 4.7

Olathe

773

756

- 2.2

Totals

7,702

8,938

Ouray

785

709

- 9.7

Ridgway

254

262

+ 3.1

Totals

1,039

971

- 6.5

Montrose

Ouray

%

18,286 18,366

1,601

1,546

+0.5

-3.4

Rural Population 1/
%

1960

1970

Change

9,365 9,264

- 1.1

+16.0 10,584 9,428

-10.9

562

575

1/ Sources are the U. S. Bureau of the Census and Colorado Yearbook, 1959-61.
~/ Excludes Mesa County because, despite being comparatively large in size, that
portion of the county involved in Unit 62 contains an indeterminable, very small
and entirely rural, human population.

1/ County rural population is based upon total county population less total of
incorporated cities and towns (urban population).

+ 2.3

�9B

The Colorado State Planning Office (pers. comm. 1971) provided the
following advance population projections for the three counties listed·
in Table 1:
1980
Montrose County

18,500

2010 and Thereafter

Forevermore

20,000
15,000

Delta County
1,500
Ouray County

B. D. TIakeI
July 1971

�lOB

GAME SPECIES - WILDLIFE
Big game mammals

MANAGEMENT

UNIT 62 (E. UNCOMPAHGRE)

~I

Pronghorn (Antilocapra americana)
Black bear (Ursus americanus)
Elk (Cervus canadensis)
Mountain lion (Felis concolor)
Mule deer (Odocoileus hemionus)

Small game mammals

11

Cottontail rabbit (~vilagus
audubonii; ~. nuttallii)
Pine (red) squt rre I (Tamiasciurus hudsonicus)
Snowshoe hare (Lepus americanus)
Abert's squirrel (Sciurus aberti)
Game birds ]j
Migratory

waterfowl

and shorebirds

Great Basin Canada goose (Branta canadensis
Black brant (Brant a nigricans) 11
T.TL...:.•..........£:....-,..._+"U

...LL..c:.

J...

LULJ.\...C\..L

..3

__

-.

6vvoIL-

IA_

_

\.C'"l...LLOL.L,

11....:+=_-.-.
Q..l..U..J...L

.LULL':"

1::_

moffitti)

_+-_1.:_\

.L. LULLLOJ....L0I

-s r

.::!....I

Snow goose (Chen hyperborea hyperborea) 11
Mallard (~as platyrhynchos platyrhynchos)
Gadwall (Anas strepera)
Pintail (Anas acuta)
Green-winged teal (Anas carolinensis)
Blue-winged teal (Anas discors discors)
Cinnamon teal (Ana~anoptera
septentrionalium)
American widgeon (Mareca americana)
Shoveler (Spatula clypeata)
Wood duck (Aix sponsa)
Redhead (Aythya americana)
Ring-necked duck (Aythya _collaris)
Canvasback (Aythva valisineria)
Greater scaup (Aythya marila nearctica) 11
Lesser scaup (Aythya affinis)
Common goldeneye (Bucephala clangula americana)
Barrow's golden-eye (Bucephala islandica) 11

11 Nomenclature
Colorado.

according to Lechleitner, R. R. 1969.
Pruett Publishing Co., Boulder.
254 pp.

Wild mammals

11 Nomenclature
Pictorial

according to Bailey, A. M., and R. J. Neidrach.
checklist of Colorado birds.
Denver Mus. Nat. Rist.

11 Unverified

in hunter's
game in 1970-71.

bag checks but possible

rare migrant

of

1967.
168 pp .
and legal

�llB

Game birds

- Migratory

waterfowl

and shorebirds

(continued)

Bufflehead (Bucephala albeola)
Ruddy duck (Oxyura jamaicensis rubida)
Hooded merganser (Lophodytes cucullatus)
Connnon merganser (Hergus merganser americanus)
Red-breasted merganser (~serrator
serrator)
COlumon snipe (Capella gallinago delicata)
American coot (Fulica americana americana)

Upland game birds
Blue grouse (Dendragapus obscurus obscurus)
Sage grouse (Cp.ntrocercus urophasianus urophasianus)
Ring-necked pheasant (Phasianus colchicus)
Chukar (Alectoris graeca)
Band-tailed pigeon (Columba fasciata fasciata)
Mourning dove (Zenaidura macroura marginella)
Sharp-tailed grouse (Pediocetes phasianellus columbianus)
Gambel's quail (Lophortyx gambelii sanus)
Wild turkey (Meleagris gallopavo merriami)
Mountain

quail

(Oreortyx picta)

W. T. McKean
November 1971

�12B

OTHER MA}~IAN

Furbearers

SPECIES

1/ - WILDLIFE

MANAGEMENT

UNIT 62 (E. UNCOMPAHGRE)

Jj

Beaver (Castor canadensis)
Muskrat (Ondatra zibethicus)
Ringtail (Bassariscus astutus)
Weasels (Mus tela erminea; M. frenata)
Mink (Hustela vison)
Marten (Martes americana)

Nongame

mammals

1/

White-tailed jack rabbit (Lepus townsendii)
Black-tailed jack rabbit (Lepus c.all f orn.Lcus)
Yellow-bellied marmot (Marmota flaviventris)
White-tailed prairie dog (Cynomys leucurus)
Gunnison's prairie dog (Cynomys gunnisoni)
Rock squirrel (Spermophilus variegatus)
White-tailed antelope squirrel (Ammospermophilus leucurus)
Golden-mantled
ground squirrel (Spermophilus lateralis)
Least chipmunk (Eutamias minimus)
Colorado chipmunk (Eutamias quadrivittatus)
Coyote (Canis latrans)
Red iox (Vulpes iulva)
Gray fox (Urocyon cinereoargenteus)
Porcupine (Erethizon dorsatum)
Raccoon (Procyon lotor)
American badger (Taxidea taxus)
Spotted skunk (Spilogale putorius)
Striped skunk (Mephitis mephitis)
Bobcat (Lynx rufus)

1/ These species, grouped separately as "Furbearers" and "Nongame mammals"
~nd outside of "game" categories, follm., Chapter 62, Colo. Rev. Statutes
1962 As Amended, in Colo. Game, Fish and Parks Div. Laws and Regulations
Hdbk., 1970 (Art. 1, Item 3, Definitions, p. 327).

1/ Nomenclature
Colorado.

actording to Lechleitner, R. R. 1969. Wild mammals
Pruett Publishing Co., Boulder, Colo.
254 pp.

W. T. McKean
November 1971

of

�l3B

/'

Nongame

OTHER AVIAN SPECIES

birds

1/

WILDLIFE

MANAGEMENT

UNITS 61 AND 62

1/

Common loon (Gavia immer) Rare migrant.
Horned grebe (Podiceps auritus cornutus) Rare migrant.
Eared grebe (Podiceps caspicus californicus) Uncommon migrant.
Western grebe (Aechmophorus occidentalis) Possible rare migrant.
Pied-billed grebe (Podilymbus podiceps podiceps) Uncommon migrant ]/ and
possible rare summer resident.
Double-crested
cormorant (Phalacrocorax auritus auritus) Rare
migrant.
Great blue heron (Ardea herodias treganzai) Common summer resident and few
in winter 3/, 4/.
Snowy egret (Leuc;phoyx thula brewsteri) Uncommon summer resident.
Black-crowned night heron (Nycticorax nycticorax hoactli) Uncommon summer
resident.
Least bittern (Ixobrychus exilis exilis) Possible accidental summer visitor.
American bittern (Botaurus lentiginosus) Possible rare migrant.
White-faced ibis (Plegadis ch Lh i.) Possible rare migrant.
Whistling swan (Olor columbianus) Rare migrant.
Sandhill crane (Grus canadensis canadensis; Q. ~. tabida) Common migrant.
Virginia rail (Rallus limicola limicola) Possible uncommon summer resident.
Sora (Porzana carolina) Possible uncommon summer resident.
Semipalmated plover (Charadrius semipalmatus) Possible rare migrant.
Snowy plover (Charadrius alexandrinus nivosus) Possible accidental migrant.
Killcieer(ClldLadLiu5
v0cifcLUO vvcif~~u3) COu~0u 3ulliill2r olid u~~G~~G~ wi~t~~
resident 3/.
Mountain plo~er (Eupoda montana) Possible accidental migrant.
Black-bellied plover (Sguatarola sguatarola) Possible rare migrant.

ll

1/ These species, grouped separately as IINongame birds" and IIRaptores and
;utside of "game" categories, follow Chapter 62, Colo. Rev. Statutes 1963 As
Amended, in Colo. Game, Fish and Parks Div. Laws and Regulations Hdbk., 1970.
(Art. 1, item 3, Definitions, p. 327).

1/ Nomenclature

from Bailey, A. M., and R. J. Neidrach.
1965. Birds of Colorado.
Denver Mus. Nat~ Hist. 2 Vols. 895 pp. Information on occurrence and method for
reporting same are adapted from the foregoing reference and Davis, W. A.
1969.
Birds in western Colorado.
Colo. Field Ornith. 61 pp., with the system of Davis
(1969: 6) favored and f oLl.owed closely throughout.
Presence of term "possible"
is an author judgement connotating that the species could occur in the units
some time even though evidence is presently lacking that it does. Absence of
term means that evidence by Bailey and Neidrach (1964) and/or Davis (1969) makes
occurrence a near certainty or that sighting(s) have been reported verbally by
anyone
or more of Division personnel H. Burdick, T. Quick, L. Denton
[and others,
as appropriate] or qualified by additional footnotes that follow.

1/ Unpublished

checklist

of birds of the Uncompahgre

Plateau

by Dr. A. S. Hyde,

1971.
4/ Sight record of A. E. Anderson as given in APPENDIX A - PARTIAL CHECKLIST OF
VERTEBRATE FAUNA OBSERVED IN VICINITY OF STUDY AREA, pp. 59-60, of Anderson, A. E.
1960.
Effects of sagebrush eradication by: chemical means on deer and related wildlife.
Fed. Aid Job Compo Rep. Proj. W-38-R-13, July, Colo. Dept. Game and Fish.
72 pp. (processed).

�Nongrune birds

(continued)

Eong+b i.l.Ledcurlew

(Numenius americanus americanus) Poss ible ra re mi.gran t ,
Spotted sandpiper (Actitis macularia) Common summer resident 2/.
$olitary sandpiper (Tringa solitaria cinnamomea) Common migrant and
occasional summer visitor.
Willet (Catoptrophorus semipalmatus inornatus) Rare migrant.
Greater yellowlegs (Totanus melanoleucus) Possible common migrant.
Lesser yellowlegs (Totanus flavipes) Possible uncommon migrant.
Knot (Calidris canutus rufa) Possible accidental migrant.
Pectoral sandpiper (Erolia melanotos) Possible rare migrant.
Ba Lr'd s sandpiper (Erolia bairdii) Possible uncommon migrant.
Least sandpiper (Erolia minutilla) Possible COmmon migrant.
Long-billed dowitcher (L'Lmnodr-omus scolopaceus) Possible rare migrant.
Semipalmated sandpiper (Ereunetes pusillus) Rare migrant 2./.
Western sandpiper (Ereunetes mauri) Possible uncommon migrant.
Marbled godwit (Limosa fedoa) Possible rare spring migrant.
Sanderling (Crocethia alba) Possible accidental migrant.
American avocet (Recurvirostra americana) Rare migrant.
Black-necked stilt (Himantopus mexicanus) Possible rare migrant.
Wilson's phalarope (Steganopus tricolor) Common migrant and uncommon
Summer resident.
Northern phalarope (Lobipes lobatus) Possible rare migrant.
Herring gull (Larus argentatus smithsonianus) Possible unCOmmon migrant.
California gull (Larus californicus) Possible rare migrant.
Ring-billed gull (Larus delawarensis) Upcommon migrant.
Franklin's gull (Larus pipixcan) Common migrant.
Bonaparte's gull (Larus philadelphia) Possible rare migrant.
forster's tern (Sterna forsteri) Possible rare migrant.
Common tern (Sterna hirundo hirundo) Possible accidental migrant.
~east tern (Sterna albifrons athalassos) Possible accidental migrant.
Black tern (Chlidonias niger surinamensis) Possible rare migrant.
Rock dove (Columba livia) Common resident.
yellow-billed cuckoo (Coccyzus americanus americanus) Possible uncommon
summer resident.
roor-will (Phalaenoptilus nuttallii nuttallii) Common summer resident ~/.
Common nighthawk (Chordeiles minor powelli ; f. ~. henryi ) Common
" Summer res ident 41.
"
Black swift (Cypsel;ides niger borealis) Possible accidental Summer migrant.
White-throated
swift (Aeronautes saxatalis sclateri) Common Summer
, resident 3/,4/.
alack-chinned hu~ingbird
(Archilochus alexandri) Common Summer resident.
aroad-tailed hummingbird (Selasphorus platycercus platycercus) Common
: summer resident 2/.
Rufous hummingbird (Selasphorus rufus) Common late s~mmer and fall migrant.
~alliope hummingbird (Stellula calliope) Rare summer migrant ~/and Possible
" rare Summer resident.
~ivoli's hummingbird (Eugenes fulgens aureoviridis) Possible rare summer
" visitor.
Belted kingfisher (Megacerle alcyon alcyon) Common resident 2/.
Ye lLow-cs ha f t ed flicker (Colaptes auratus luteus) Possible accidental
migrant.
t

�15B

Nongame

birds

(continued)

Red-shafted flicker (Colaptes cafer collatis) Common resident 3/, 4/.
Lewis' woodpecker
(Asyndesmus lewis) Common resident 3/.
-Yellow-bellied
sapsucker (Sphyrapicus varius muchalis) Common summer
resident.
Williamson's
sapsucker (Sphyrapicus thyroideus nataliae) Common summer
resident.
Hairy woodpecker
(Dendrocopos villosus monticola) Uncommon resident if.
Downy woodpecker
(Dendrocopos pubescens leucurus) Uncommon resident.
Northern three-toed woodpecker
(Picoides tridactylus dorsalis) Rare resident.
Eastern kingbird (Tyrannus tyrannus) Uncommon summer resident 1/.
Western kingbird (Tyrannus verticalis) Common summer resident 1/.
Cassin's kingbird (Tyrannus vociferans vociferans) Possible uncommon
summer resident.
Ash-throated
flycatcher (Myiarchus cinerascens cinerascens) Common summer
resident.
Say's phoebe (Sayornis saya saya) Common summer resident.
Traill's flycatcher (Ernpidonax traillii) Uncommon summer resident 1/·
Hammond's flycatcher (Ernpidonax hammondii) Uncommon summer resident.
Dusky flycatcher (Empidonax oberholseri) Uncommon summer resident.
Gray flycatcher (Ernphidonax wrightii) Common summer resident.
Western flycatcher (Ernpidonax difficilis hellmayri) Common summer resident.
Western wood peewee (Contopus sordidulus veliei) Common summer resident if.
Olive-sided flycatcher (Nuttallornis borealis) Uncommon summer resident.
R0:!:'n",rl l?yk (Fr",rn_&lt;:mh;1::l
::llnp.~rri~lell;:;'.8p.m.8)
Common resident 3/.
Violet-green
swallow (Tachycineta thalassina lepi~a) Common summer
resident 3/, 4/.
Tree swallow-(Irfdiprocne
bicolor) Common summer resident 1/.
Bank swallow (Riparia riparia riparia) Uncommon migrant and summer resident.
Rough-winged
swallow (Stelgidopteryx ruficollis serripennis) Uncommon
migrant and summer resident.
Barn swallow (Hirundo rustica erythrogaster)
Common migrant and summer
resident 3/.
Cliff swallo; (Petrochelidon pyrrhonota pyrrhonota) Common summer resident.
Purple martin (Progne subis subis) Possible accidental summer migrant.
Gray jay (Perisoreus canadensis capitalis) Common resident.
Steller's jay (Cyanocitta stelleri macrolopha) Common resident 11, if.
Scrub jay (Aphelocoma coerulescens woodhouseii)
Common resident 1/·
Black-billed magpie (Pica pica hudsonia) Common resident 11, !/.
Common raven (Corvus corax sinuatus) Cornmon resident' 11,
Common crow (Corvus brachyrhynchos
brachyrhynchos)
Common resident 1/·
Pinyon jay (Gyrnnorhinus cyanocephalus)
Common resident 1/, ~/.
Clark's nutcracker
(Nucifraga columbiana) Common resident
Black-capped
chickadee '(Paru s atricapillus garrinus) Common resident
Mountain chickadee (Parus gambeli gambeli) Common resident
Plain titmouse (Parus inornatus ridgwayi) Common resident.
Common bushtit (Psaltriparus minimus plumbeus) Cornmon resident 1/, !!..
White-breasted
nuthatch (Sitta carolinensis nelsoni) Uncommon resident if.

i/.

i/.

i/.

i/.

�loB

Nongame

birds

(continued)

Red-breasted nuthatch (Sitta canadensis) Rare resident.
Pygmy nuthatch (Sitta pygmaea melanotis) Common resident ~/.
Brown creeper (Certhia familiaris montana) Uncommon resident.
Dipper (Cinclus mexicanus unicolor) Common resident.
House wren (Troglodytes aedon parkrnanii) Common summer resident 3/, 4/.
Winter wren (Troglodytes troglodytes pacificus) Status unknown
Bewick's wren (Thryomanes bewickii eremophilus) Common summer and rare
winter resident 1/.
Long-billed marsh wren (Telmatodytes palustris plesius) Uncommon summer
resident 3/.
Canyon wren (Salpinctes mexicanus conspersus) Uncommon summer 1/ and
possible rare winter resident.
Rock wren (Salpinctes obsoletus obsoletus) Common summer and possible
rare winter resident.
Mockingbird (Mimus polyglottos leucopterus) Uncommon resident.
Catbird (Dumetella carolinensis) Possible rare summer resident.
Brown thrasher (Toxostoma rufum longicauda) Possible accidental migrant.
Sage thrasher (OreoscoDte~ montanus) Common summer resident ~/.
Robin (Turdus migratorius propinguus) Common resident 1/, ~/.
Hermit thrush (Hylocichla guttata auduboni)
Common summer resident.
Swainson's thrush (Hvlocichla ustulata almae) Possible common migrant
and rare summer resident.
Veery (Hylocichla fuscescens salicicola) Common summer resident.
Western bluebird (Sialia mexicana bairdi) Common migrant, uncommon
summer resident ~/, and possible rare winter resident.
Mountain bluebird (Sialia currucoides) Common migrant and summer resident
]/, ~/, and poss-ib-i-e-~·i-nter-r;sident.
Townsend's solitaire (Myadestes townsendi townsendi) Uncommon resident ~/.
Blue-gray gnatcatcher (Polioptila caerulea amoenissima) Common summer
resident ]..1.

47. -

Golden-crowned kinglet (Regulus satrapa amoenus) Possible common summer
and uncommon winter resident.
Ruby-crowned kinglet (Regulus calendula cineraceus) Common migrant 1/ and
summer resident 4/ and possible accidental winter resident.
Water pipit (Anthus-spinoletta
alticola) Common migrant 1/ and possible
winter resident.
Bohemian waxwing (Bombycilla garrulus pallidiceps) Common winter migrant.
Cedar waA~ing (Bombvcilla cedrorum) Irregular visitor.
Northern shrike (Lanius excubitor invictus) Common winter resident ]j.
Loggerhead shrike (Lanius ludovicianus excubitorides) Uncommon summer
and common winter resident 3/.
Starling (Sturnus vulgaris vulgaris) Common resident 1/.
Gray vireo (Vireo vicinior) Uncommon summer resident.
Solitary vireo (Vireo solitarius plumbeus) Common summer resident 11.
Red-eyed vireo (Vireo olivaceus) Possible rare summer resident.
Warbling vireo (Vireo gilvus s1.J"ainsonii)Common summer resident.
Tennessee warbler (Vermivora peregrina) Possible rare migrant.
Orange-crowned warbler (Vermivora celata orestera) Possible uncommon
migrant and summer resident.
Nashville warbler (Vermivora ruficapilla ridgwayi) Possible rare migrant.
Virginia's warbler (Vermivora virginiae) Common summer resident.
Yellow warbler (Dendroica netechia aestiva) Common summer resident 1/.
Myrtle warbler (Dendroica coronata co ron a t a) Uncommon migrant.·

�l7B
Nongame birds

(continued)

Audubon's warbler (Dendroica auduboni memorabilis) Common summer
resident 3/, 4/ and possible rare winter resident.
Black-throat~d g~ay warbler (Dendroica nigrescens) Common summer
resident 3/.
Townsend's w~rbler (Dendroica townsendi) Uncommon migrant 4/.
Grace's warbler (Den~~aciae
graciae) Rare summer re;ident ~/.
Northern waterthrush (Seiurus noveboracensis notabilis) Rare migrant 1/.
MacGillivray's warbler (Oporornis tolmiei monticola) Common migrant and
uncommon summer resident 1/.
Yellowthroat (Geothlypis trichas occidentalis; G. ~. campicola) Uncommon
summer resident 1/.
Yellow-breasted
chat (Icteria virens auricollis) Common summer resident.
Wilson's wa rbLer (Hilsonia pusilla pileolata) Common migrant and possible
summer resident.
American redstart (Setophaga ruticilla tricolora) Possible rare migrant.
House sparrow (Passer domesticus domesticus) Common resident 1/.
Bobolink (Doli~honyx oryzivorus) Possible rare summer migrant.
Western meadowlark (Sturnella neglecta neglecta) Common summer 1/ and
uncommon winter resident.
Yellow-headed blackbird (Xanthocephalus xanthocephalus) Common summer
resident 3/.
Red-winged blackbird (Agelaius phoeniceus fortis) Common resident 1/.
Orchard oriole (Icterus spurius) Possible accidental summer visitor.
Bullock's oriole (Icterus bullockii bullockii) Common summer resident 1/.
Rusty blackbird (Euphagus carolinus carolinus) Possible late winter visitor.
Brewer's blackbird (Euphagus cyanocephalus) Common resident 1/.
Brown-headed cowbird (Mol~thrus atel:'_
artemisiae) Common summer resident.
Western tanager (Piranga ludoviciana) Common migrant and summer resident i/.
Scarlet tanager (Piranga olivacea) Possible accidental summer migrant.
Rose-breasted grosbeak (Pheucticus ludovicianus) Possible accidental spring
migrant.
Black-headed grosbeak (Pheucticus melanocephalus melanocephalus)
Common
summer resident.
Blue grosbeak (Guiraca caerulea interfusa) Uncommon summer resident 1/.
Lazuli bunting (Passerina amoena) Uncommon summer resident.
Dickcissel (Spiza americana) Possible accidental migrant.
Evening grosbeak (Hesperiphona vespertina brooksi) Irregular resident.
Cassin's finch (Carpodacus cassinii) Common resident.
House finch (Carpodacus mexicanus frontalis) Common summer and uncommon
winter resident 1/.
Pine grosbeak (Pinicola enucleator montana) Uncommon resident.
Gray-crowned rosy finch (Leucosticte tephrocotis tephrocotis; ~. t.
littoralis) Common winter migrant.
Black rosy finch" (Leucosticte atrata) Uncommon winter migrant.
Brown-capped rosy finch (Leucosticte australis) Common winter resident.
Common redpoll (Acanthis flammea flammea) Possible accidental winter migrant.
Pine siskin (Spinus pinus pinus) Common resident ~/.
American goldfinch (Spinus tristis tristis; ~. ~. pallidus) Common summer
1/ and possible uncommon winter resident.
Lesser goldfinch (Spinus psaltria psaltria) Possible uncommon summer
resident and accidental winter migrant.
Red crossbill (Loxia curvirostra) Irregular visitor.
White-winged crossbill (Loxia leucoptera leucoptera) Possible accidental
winter migrant.

�l8B
Non-game

birds

(continued)

v.

Green-tailed
towhee (Chlorura chlorura) Common summe r res ident
~j and
possible rare winter resident.
Rufous-sided
towhee (Pipilo erythrophthalmus
montanus) Uncommon resident '}../.
Lark bunting (Calamospiza melanocorys)
Possible uncormnon Surmner resident.
Savannah sparrow (Passerc~lus sandwichensis nevadcnsis; f. ~. anthinus)
Possible uncormnon migrant and surmner resident.
Grasshopper sparrow (Armnodramus savannarum perpallidus) Uncormnon surmner
res ident 5 I .
LeConte's sp;rrow (Passerherbulus
caudacutus) Possible accidental migrant.
Sharp-tailed
sparrow (Armnospiza caudacuta nelsoni) Possible accidental
migrant.
Vesper sparrow (Pooecetes gramineus confinis) Cormnon migrant and surmner
res ident '}../,
i/.
Lark sparrow (Chondestes grarmnacus strigatus) Common migrant and surmner
res ident ,}j.
Black-throated
sparrow (Amphispiza bilineata deserticola) Cormnon surmner
res ident.
Sage sparrow (Amphispiza belli nevadensis) Cormnon sununer resident.
White-winged
junco (Junco aikeni) Possible accidental winter visitor.
Slate-colored
junco (Junco hyemalis hyemalis; I. h. cismontanus) Possible
rare winter res ident.
Oregon junco (Junco oreganus oreganus; I. ~. mearnsi) Cormnon winter
res ident 3 I .
Gray-headed junco (Junco caniceps caniceps) Cormnon resident i/.
Tree sparrow (Spizella arborea ochracea)Uncormnon
winter visitor 1/.
Chipping sparrow (Spizella passerina boreophila) Common Surmner resident 1/,4/.
Brewer's sparrow (Spizella breweri breweri) Common summer resident 1/, i/.
Har-r+s

' s ps.r row

(7.nnnt:rir.hiR

(JITF&gt;rI11;]) Po s s t b l a

r2rp.

wintF&gt;r

resinF&gt;nt:_

1.

White-crowned
sparrow (Zonotrichia leucophrys leuchophrys; ~.
gambelii)
Cormnon resident 3/.
White-throated
spar~w
(Zonotrichia albicollis) Possible accidental migrant.
Fox sparrow (Passerella iliaca schistacea) Rare summer resident.
Lincoln's sparrow (Melospiza lincolnii alticola) Cormnon migrant and Surmner
resident.
Song sparrow (Melospiza melodia) Common resident 1/.
Lapland longs pur (Calcarius lapponicus alascensis) Possible rare winter
migrant.
Chestnut-collared
longs pur (Calcarius ornatus) Possible accidental winter
migrant.

�19B

Raptores

l:.../

Turkey vulture (Cathartes ~
meridionalis) Common sunnner ;}/, !!/ and
rare winter resident.
)
Goshawk (Accipiter gentilis atricapillus) Uncommon resident ~/.
Sharp-shinned hawk (Accipiter striatus velox) Unconnnon summer ;}j and
winter resident.
Cooper's hawk (Accipiter cooperii) Uncommon summer ~/ and winter
resident.
Red-tailed hawk (Buteo jamaicensis calurus) Common res Lden t ;}../,~/.
Swainson's hawk (Buteo swainsoni) Uncommon migrant and summer resident.
Rough-legged hawk (Buteo lagopus ~. johannis) Common winter resident.
Ferruginous hawk (Buteo regalis) Rare summer resident.
Golden eagle (Aquila chrysaetos canadensis) Common resident if, ~/.
Bald eagle (Haliaeetus leucocephalus alascanus) Common winter resident 5/.
Marsh hawk (Circus cyaneus hudsonius) Common resident ~/.
Osprey'(Pandion haliaetus carolinensis) Rare migrant.
Prairie falcon (Falco mexicanus) Uncommon resident.
Peregrine falcon (Falco peregrinus anatum) Rare resident.
Pigeon hawk (Falco columbarius) Rare wLnt.e r migrant.
Sparrow hawk (Falco sparverius sparverius) Common summer and uncommon
winter resident 3/, 4/.
Screech owl (Otus ;sio) Possible rare resident.
Flammulated owl (Otis flammeolus flammeolus) Possible accidental summer
resident.
Great horned owl (Bubo virginianus) Common resident 3/: 4/.
Pygmy owl (Glaucid~g;-oma"--califo-rnicum)
Uncommon resid-ent 3/.
Burrowing owl (Speotyto cunicularia hypugaea) Uncommon summer and possible
accidental winter resident.
Long-eared owl (Asio otus wilsonianus) Uncommon resident.
Short-eared owl (Asio~mmeus
flammeus) Uncommon winter migrant and resident.
Saw-whet owl (Aeg~s
acadicus acadicus) Uncommon resident.

5/ Golden and bald eagles specifically excluded from statutes defining
"Raptore" as cited in footnote 1) but herein listed to avoid omission.

B. D. Baker
February 1972

�20B

DISTRIBUTION AND ABUNDANCE OF BIG GAME MAMMALS
IHLDLIFE MANAGEMENT UNIT 62 (E. UNCOHPAHGRE)

BLACK BEAR
Black bears occupy approximately the area between 6,000 and 10,000 feet
elevation in Unit 62, though at times they wander over almost the entire
unit.

Their yearlong habitat here is largely within the timbered belt

and canyon country where fruit-bearing shrubs are common.

The only consistent index to black bear abundance is the hunter report
card survey.

Estimated annual black bear kill for the period 1960-1970

inclusive, averaged 24 head.

On a statewide basis, this unit ranked

no less than fourth in kill during seven years out of the past ten
(1960-1970).

In addition, during sex-ratio counts of deer a count of 18 black bears
was made in 1971 (Burdick pers. comm. 1972).

ELK
The 1962 Division-Uncompahgre

National Forest big game winter range

inventory of Unit 62 did not yield much data on elk distribution.
20 widely-distributed

Only

pellet group transects were read that year, with

negligible winter occupancy indicated for all of the areas transected.
Project W-lOl-R files at the Fort Collins Research Center hold transect
data and summaries.

Current information obtained from Southwest Region staff and field
personnel through group and individual interviews in February 1972

�21B

included year-raund elk distributian via map display.

That effort

yielded lawer limits af elk use essentially carresponding with the
Natiana1 Farest baundary, except for the fo1lawing areas af which
the Farest baundary daes not accaunt:
1.

A three- to.five-mile wide zone that extends northward
fram Dallas Divide east of the Hontrase-Ouray Caunty line.

2.

The upper-mid Dry Creek drainage area east af Silesca
Ranger Statian in a zo.netwo.to five miles wide east af
the Farest baundary.

3.

Middle Big Daminguez and Gibb1er Canyan areas east and
north af the Farest baundary.

Also, the lawer elk limit line excludes comparatively small areas of
National Farest an Dry Mesa and near middle Raubideau Creek.

Present Unit 62 elk numbers are not great in comparison with much
larger herds f ound eLsewhere in Colorado.

Nevertheless, from former

nanexistence on the plateau, elk numbers have increased to allow far
the first apen season in 1957 (no animals taken), to. a peak total
harvest of 199 in 1969.

MOUNTAIN LION
The literature makes no specific reference to. mountain lian distributian
within Unit 62.

Dixan (1967) daes not include it within any af the

better lian habitats listed, hawever, the rougher, mare isalated portians af the canyan areas do harbor at least a law populatian.

�22B

Perusal of all available data on hunter reports for the years 19651970 inclusive, failed to reveal conclusively a single lion taken in
Unit 62.

A few lions could have been killed by people other than

hunters.

However, their scarcity in this unit is evident.

MULE DEER
The cooperative Grand Mesa-Uncompahgre National Forest and Division
big game winter range inventory in 1962 yielded a tentative average
upper winter limit line for all National Forest lands plus nonForest areas in T15S, RlOOW, 6 PM; T50 and 5lN, R14W, llifPM;and T49
and 50N, RI3W, NMPM.

The upper limit line corresponds closely with

the National Forest boundary from Unaweep Canyon on the north to
about the Corral Gulch fork of Little Dominguez Creek on the south.
From the latter place, the line traverses alternating corners of non-

caused a gap on lands largely controlled by the BLM.

The line was

again placed to include a very small corner portion of the Forest in
T47N, RlOW, NMPM.

Discontinuance of the line here left the extent of

winter range in the unit's extreme southwest corner undetermined.
The upper winter line varies between extreme elevations of about 6,600
and 8,200 feet but falls more frequently between 7,000 and 7,600 feet.
Although the 1962 survey did not set a lower winter limit line, it is
assumed that a lower boundary would logically include all suitable
habitat west of cultivated cropland from Delta southward in the
Uncompahgre Valley,

More information is needed for locating the lower

line northwest of Delta in Unit 62.

�23B

Extent of occupied

summer deer range is not available

at this writing.

It is a reasonable

assumption,

deer can be

found in all territory

however,

that summering

not under intensive

the latter occasionally.

Heaviest

cultivation,

summer use probably

about 8,000 feet to the crest of the Uncompahgre
(1969) reported
elevations

that the upper Big Dominguez

and even in
occurs from

Plateau.

Anderson

area lying between

of 7,900 and 8,570 feet constituted

upper winter

and lower

summer range for deer.

Pellet group count data obtained
ditions

transect

on the abundance

readings

in 1962

concurrently

(see Elk) produced

of deer in Unit 62.

con-

information
read in

too little data from which

to

conclusions.

More recently,

the Montrose

BLM District

also conducted

counts in Unit 62 (Hollis pers. comm. 1972).
life Biologist

Hollis

data, and possibly

Abundance

meager

With just 20 transects

all of the unit, the small base yielded
draw valid

with browse range

Although

is no longer at Montrose,

a report, were to have been available

ratio and trend counting.

That information

Montrose

office

Perhaps

a rough but valid

concept of general

inferred

from past harvest

data.

that la-year

District

Wild-

he said that study

of deer by year and census area is available

years 1959-68,

pellet group

by late 1972.

from aerial

sex-

is kept at the Division's

(Burdick pers. comm. 1972).

deer abundance

On the basis of total harvest

Unit 62 ranked a strong 5th statewide.

span was 34,548,

can be

or an approximate

annual

for

Total kill for
average

of nearly

�3,455 animals.

As elsewhere in Colorado, deer numbers declined in

the late 1960s.

Despite that skid, Unit 62 ranked 4th in statewide

harvest as recently as 1971 (total yield of 1,588 head taken).

Thus,

it is possible to state that Unit 62 has exhibited good to excellent
deer abundance in spanning high and low population

trend fluctuations

experienced statewide.

PRONGHORN
The only occurrence of pronghorns within this unit is near the extreme
northeast boundary (Douglas and Ogilvie 1953).

Pronghorns commonly

move north and south across Hf.ghway 50 from the site of introduction
(1948-50) in the Wells Gulch area of Unit 411.

They occur between the

Gunnison River and Highway 50 (see 1/2" scale overlay and 1/411 scale
distribution maps).

Population estimates, based upon aerial censuses, over the years for
the small area of occupied range in Unit 62 vary from about six in
1954 (Ogilvie and Hoover 1954) upward to 75.

These animals are only

part time residents due to movements across U.S. Highway 50.

Literature Cited

Anderson, A. E.

1969.

2,4-D, sagebrush, and mule deer-cattle use of

upper winter range.
Fish and Parks.

Dixon, K. R.

1967.

pp , 141-164.

Spec. Rep. No. 21.

July.

Colo. Div. Game,

21 pp.

Evaluation of effects of mountain lion predation,
In Game Res. Rep., July - Part II.

Fish and Parks.

pp. 73-310.

Colo. Dept. Game,

�25B

Douglas, C., and S. Ogilvie.
sites, pp. 123-125.
Game and Fish.

1953.

Survey of antelope at transplant

In Quart. Prog. Rep., July.

pp. 1-129.

Ogilvie, S., and R. L. Hoover.

1954.

Survey of antelope at transplant

sites, pp. 96-97.' In Quart. Prog. Rep., July.
and Fish.

Colo. Dept.

Colo. Dept. Game

pp. 1-182.

w. T. McKean and B. D. Baker
February 1972

�26£

DISTRIBUTION AND ABUNDANCE OF SMALL GAME MAMMALS
WILDLIFE MANAGEMENT UNIT 62 (E. UNCOMPAHGRE)

COTTONTAIL
Every square mile in this unit is occupied by cottontails, or capable of
being so, with the possible exception of three or four urban areas.

Based

on reported altitudinal and habitat preferences (Lechleitner 1969, Armstrong
1972) both the desert cottontail and Nuttall's cottontail would be present.
No specific study of cottontail distribution pertinent to the unit is
known.

Recent large scale chaining operations on pinon-juniper covered.

lands have created heavy concentrations of cottontails, due to improved
brush and herbaceous cover.

Shepherd (1965a) indicates a possible density of 150-200 rabbits per square
milp f0r ~0~r

of

rhe.he.tter ~reas of the.st~te excluding highs or lows.

This estimate should be applicable to Unit 62.
SNOWSHOE HARE
Shepherd (1965b) indicates that snowshoe hares occupy, or could occupy,
Unit 62 between 8,000 and 10,000 feet elevation in the coniferous forests
for its entire length.
A statewide mean density of 125 hares per section on the habitable range
is postulated by Shepherd (1965b).
the higher areas of this unit.

This is believed to be applicable to

Anderson (1959) reported that in 1958,

snowshoe hares were apparently very scarce on the upper Big Dominguez
study area which lies between elevations of 7,900 and 8,570 feet.

�27B

PINE (RED) SQUIRREL
Pine squirrels

are reportedly

distributed

throughout

the upper two-thirds

elevationally

of Unit 62 in the coniferous

forests.

Armstrong

indicates,

on a distribution

map, this fact also.

Southwest

regional personnel

indicate

present

at all in the unit ,they

that where pine squirrels

SQUIRREL

Currently

there are no field or written

occurrence

in Unit 62.

should be found there in the ponderosa
Maps in Armstrong

No data on abundance
seen this squirrel

reported

in Unit 62.

The fact that several

1959.

units, suggest

Colo. Dept. Game and Fish.

D. M.

1972.

Kansas Printing

field men have

Cited

eradication

pp. 113-128.

by chemical

means

In Quart. Rep., Jan.

128 pp.

Distribution

Service,

Colorado.

that it very probably will be

Effects of sagebrush

on deer and related wildlife,

Armstrong,

(1959)

Unit 61, and that no physical

throughout

Literature

fu1derson A. E.

at 5,000 to 8,000

to extend this far into west-central

scattered

exist between

pine forests

(1972) and Hall and Kelson

are available.

barriers

records of this squirrel's

(1969) implies that this squirrel

Lechleitner

do not show the distribution

are

exist in maximum numbers.

ABERT'S

feet elevation.

(1972)

of mammals

Lawrence.

415 pp.

in Colorado.

Univ. of

�28B

Hall, R. E., and K. R. Kelson.
Ronald Press, N. Y.

Lechleitner, R. R.
Boulder.

1959.

2 Vols.

1969.

The mammals of North America.

1083 pp.

Wild mammals of Colorado.

254 pp.

Shepherd, H. R.

1965a.

Colorado long range management plans for game

species, 1965-1975, for cottontail rabbits.
Parks Dept. Rep.

1965b.

Colo. Game, Fish and

11 pp. (Mimeo).

Colorado long range management plans for game species,

1965-1975, for snowshoe hares.
Rep.

Pruett Pub L, .Co ,

11 pp.

Colo. Game, Fish and Parks Dept.

(Mimeo).

W. T. McKean
February 1972

The

�29B

DISTRIBUTION AND ABUNDANCE OF SMALL GAME BIRDS
WILDLIFE MANAGEMENT UNIT 62 (E. UNCOMPAHGRE)

SAGE GROUSE
Rogers (1964) indicated two areas within Unit 62 where sage grouse
populations had been consistently observed.

These were:

Dominguez

Creek-Smith Fork area on the north plateau and the Sims Mesa-Duckett
Drat" area "lest of Colona.

Only light populations (1-10 birds per square mile) were found in the
areas listed as occupied range.

Anderson (1960) also reported ou in-

dices of sage grouse numbers on the northeast portion of Uncompahgre
National Forest.

Southwest Regional personnel report that sage grouse

populations have diminished considerably in this unit over the last
ten years (Burdick pers. comma 1972).

BLUE GROUSE
Rogers (1968) indicates in his maps that the blue grouse distribution
in Unit 62 is largely within the National Forest boundary, but not
entirely.

The important habitat of fir (Abies) species, as well as

mixed brushy areas and aspen occur on Bureau of Land Management lands
as well as private lands, but not as extensively.

Blue grouse occur

at 7,000 to 10,000 feet elevation most commonly.

A grouse census (Rogers 1968) of 62.5 road miles along the Uncompahgre
Divide between Units 61 and 62 in 1961-63 developed an average count
of 0.039 blue grouse per mile, ranging from 0.025 to 0.097.

�30B

SHARP-TAILED GROUSE
Rogers (1969) reports small populations of sharp tails either at
dancing grounds or observed elsewhere in Unit 62 as follows: Mesa
County - near junction of Uncompahgre Divide Road and DOT Cow Camp
Road; Montrose County - near T-Bone Springs on Sawmill Mesa, and on
old Paradox Road; Ouray County - south end of Uncompahgre Plateau
(no definite land marks noted).

None were reported in that portion

of Delta County encompassing this unit.

The only available data on density is that of Rogers (1969) in which
observations on dancing grounds were made on the north end of the
Uncompahgre Plateau for 1963-1965.

Density estimates

high of 3.0 birds per square mile to 0.0 in 1965.

varied from a

Current indications

by Southwest Regional personnel are that only a very light population
remains.

GAMBEL'S QUAIL
Sandfort (1965) states that birds are found along the Gunnison and
Uncompahgre Rivers ~vithin this unit at elevations of 4,500 to 6,200
feet in drier, brushy sites where, "... irrigated farmlands, creek
bottoms, thickets and brushy sides of mesas are found in desirable
combination".

No data on density per unit area within Unit 62 have been found.

Har-

vest surveys do indicate that a few birds have been taken annually
since 1955, at least, in Small Game Unit 19.

HildlifeManagement

Unit

62 comprises approximately the southwestern third of Small Game Unit 19.

�3IB

CHUKARS
Sandfort (1965) in summarizing statewide chukar distributions, indicated that this bird had been introduced at several sites within
Unit 62 previously.

Populations were established by 1965 in

Escalante and Dominguez Creeks, as well as in the Roubideau Creek
area by 1967.

During the period 1958-1963 in this unit, and adjoining ones, population densities varied widely, varying from 47 to 26 birds per square
mile (Sandfort 1965) in the better chukar range.

Fair numbers are

indicated at the present time in Roubideau Creek (Cox pers. comm. 1972)
and light numbers in Escalante Creek (~ink pers. comm. 1972).

RING-NECKED PHEASANT
Fair to poor pheasant populations have existed in the irrigated farmlands of Unit 62, principally along the Gunnison and Uncompahgre Rivers
and their tributaries for many years (Swope 1965).

The area between

Delta and Montrose contains the largest single block of pheasant range.
There are approximately 91.3 square miles of pheasant range within Unit
62.

Recent updated data on abundance are available and on file in the Southwest Regional office, including sex ratios, crow count routes, and reproductive success.

WILD TURKEY
Myers (1964 through 1967) mapped the approximate distribution of wild
turkeys within Unit 61 and 62 for four years.

He showed that turkeys

were widely dispersed throughout the forested portions of Unit 62.
A few of the drainages in Unit 62 in which turkeys were consistently

�observed

at one or more seasons of the year were: Big Dominguez,
/

Dry

Fork Escalante,

Dry Creek, Spring Cre~,

Horse Fly Creek, and Pleasant

Valley

Turkeys

at lower elevations

Creek.

generally~intered

7,000 feet) .within, or near to the pinon-juniper
summering

areas were largely

at 8,000-9,000

pine and aspen types, although
of the plateau.
indicated

Myers

Currently

in winter.

Plateau,

combining

is more restricted

that the most reliable

than above

considerable

for the five winters

Birds were counted in roughly

Recent

counts at similar

stations

100 birds on the entire unit

a population

Their occurrence

commonly

a

units.

of less than

PIGEON

throughout

that band-tailed

both Wildlife

seems to relate

of acorns from Gambel's

native
oak.

grains are grown have appreciable
occurs in the spruce-fir

Management

pigeons
Units 61

to areas where an abundant

supply of dense, fruit-producing

as well as an abundance

feed

(Burdick pers. corom. 1972).

(1970 and pers. carom. 1971) indicates

cultivated

food

1963 through

equal numbers between

indicate

BAND-TAILED

scattered

An average

from 140 to 485, indicated

1967.

and consistent

established

both Big Game Units 61 and 62.

drop in population

are irregularly

census method

The data at hand are for the entire Uncompahgre

ground count of 306 birds, ranging

Nesting

feet in the ponderosa

tried, was the count at artifically

stations

where

and

(Cox and Mink pers. comm. 1972).

among several

and 62.

type; breeding

they had often been seen along the top

distribution

(1964 to 1967) indicated

Braun

(5,500-

shrubs exist,
Also, valleys

numbers

and various

of pigeons.

pine zones;

�33B

therefore the entire Uncompahgre Plateau is used at all seasons,
except winter (Dec. through April).

Only generalizations concerning band-tailed pigeon abundance, based
upon reported observations, are possible (Braun 1970).

The Uncompahgre

Plateau is mentioned as one of the places in Colorado where pigeons are
most abundant.

No differences in bird densities have been detected

between Wildlife Management Units 61 and 62.

Highest observed densities

occurred where food (berries, acorns, and grain) are readily available
and adjacent to forested roosting and escape cover.

Burdick (pers. comm. 1972) reported having counted in 1970 by helicopter
in excess of 2,500 birds within Units 61 and 62.

Quick (pers. comm.

1972) also indicated an increase in Unit 62.

MOUKNING

DOVE

Mourning doves are distributed over the entire Uncompahgre Plateau during
spring, summer and fall (Braun pers. comm. 1971).

A few winter on the

unit (Quick pers. comm. 1972).
No data exist concerning densities of mourning doves on the Uncompahgre
Plateau.

Braun (pers. comm. 1971) stated that higher concentration

occurs in the pinon-juniper and farmland types than in the pine or sprucefir.

Over three hundred birds were trapped in Unaweep Canyon during the

summer of 1971.

WATERFOWL
Breeding population:

Grieb (1965) indicated that fair to high breeding

populations of green-winged teal and coots occur in Unit 62, with fair

�34·B

to low populations

of mallards.

and a few other dabbling

Hopper

(1968) added that mallards

ducks breed along the Uncompahgre

Montrose

to Delta and along the Gunnison

Adjoining

these rivers are irrigated

ponds.

These provide

numbers

also nest in isolated beaver

River from

River from Delta to Whitewater.

farmlands

with their canals and

food and cover for additional

ducks.

ponds throughout

Small

the unit

(Burdick

pers. comm. 1972).

Hopper

(pers. corom. 1971) estimated

to be between

the spring population

two and three breeding

based on counts made over comparable
Planimetered

acreages

of irrigated

types add up to 127 square miles.

of Unit 62

pairs per square mile

(4-6 ducks),

land in the San Luis Valley.

cropland

and river bottom vegetation

An estimated

population

of 500 to 800

total birds results.

Wintering

population:

geese, winter within
sections
bottom

The only known areas where waterfowl,
Unit 62 are in open waters

of the Uncompahgre

lands.

Black Canyon

Large numbers
(Unit 64).

and Gunnison

including

of the above mentioned

Rivers and small ponds of river

of ducks, principally

They rest on the Gunnision

mallards,

winter

in

River but feed in

Unit 62, in corn and waste grain fields.

The river bottom
1968-1971,

duck population

within

based upon aerial censuses

in January

has averaged

2,605 birds ranging

birds ranging

from 32 to 69 (unpublished

by Regional

from 817 to 4,986.

MOUNTAIN
A number of birds were reported

Unit 62 for the four year period

information

personnel,

Geese averaged

57

- Colo. G.F.&amp; P.).

QUAIL

1-1/2 miles above Pickett

Corral ranch

�35B
on East Fork Escalante Creek by Gary Myers and others in 1966.

These

had apparently moved across the Uncompahgre Plateau from the Indian
Creek release site a distance of about 25 miles (Rogers 1967).

As of 1972, the flock at Pickett Corral did not number in excess of
25 head (Burdick pers. corom.1972).

Literature Cited

Anderson, A. E.

1960.

Distribution and abundance indices of selected

biota in western Colorado before and two months after 2,4-D
application.

Braun, C. E.

M.S. Thesis.

1970.

1965.

Colo. Div. Game, Fish and Parks.

Hopper, R. M.

1968.

April.

Wetlands of Colorado.

1964 through 1967.

Colo. Game, Fish and

Tech. Publ. No. 22.

Colo.

88 pp.

Game bird surveys.

Colo. Dept. Game, Fish and Parks.

(processed).

171 pp.

36 pp.

Dept. of Game, Fish and Parks.

Myers, G. T.

In

Colorado long range management plans for game

species, 1965-1975, for migratory birds.
Parks Dept. Rep.

130 pp.

Band-tailed pigeon investigations, pp. 151-171.

Game Res. Rep., Oct.

Grieb, J. R.

Colo. State Univ., Ft. Collins.

In Game Res. Rep.,

pp. variable by year

�36B

Rogers, G. E.

1964.

Pub1. No. 16.

1967.

Sage grouse investigations in Colorado.
Colo. Game, Fish and Parks Dept.

Tech.

132 pp.

Study of mountain quail adaptability, pp. 215-218.

Game Res. Rep., April - Part 2.

In

Colo. Dept. Game, Fish and Parks.

pp. 107-241.

1968.

The blue grouse in Colorado.

Div. Game, Fish and Parks.

1969.
23.

The sharp-tailed grouse in Colorado.

Sandfort, W. W.

1965.

1965.

8 pp.

94 pp.

Colo. Game, Fish and

(Mimeo).

Colorado long range management plans for game

species, 1965-1975, for pheasants.
Dept. Rep.

Tech. Pub1. No.

Colorado long range management plans for game

species, 1965-1975, for Gambe1's quail.
Parks Dept. Rep.

Colo.

63 pp.

Colo. Div. Game, Fish and Parks.

Swope, H. M.

Tech. Pub1. No. 21.

Colo. Game, Fish and Parks

54 pp.

W. T. McKean
February 1972

�37B
DEER KILLl SEASONS AND HUNTING PRESSURE
WILDLIFE MANAGEMENT UNIT 62

YEAR

HUNTING
PRESSURE*

BUCKS

DOES

FAWNS

HUNT AND SEASON

TOTAL

1956

1429

633

308

57

998

1957

2229

1261

863

293

2417

1958

3285

1181

827

381

2389

1959

4159

1789

1277

390

3456

1960

3374

1871

1392

546

3809

1961

4112

2729

2305

983

6017

1962

3201

1538

1313

478

3329

1963

3549

1654

1367

648

3669

1964

4315

2055

1536

475

4066

1965

4546

1701

1585

645

3931

1966

3524

985

681

216

1882

1967

3793

1553

838

227

2618

1968

2449

1060

651

162

1873

1969

4826

1988

1423

406

3817

1970

3794

1603

271

25

1899

ESM
T
ESM
2DM
2DM
1DM
1DM
1DM
1DM
1DM
3D
3D

10/15 - 10/31
12/1 - 12/31, In Part
11/17 - 11/26, In Part
10/15 - 11/17
12/1 - 12/31, In Part
10/15 - 11/2
11/3 - 12/31, In Part
10/17 - 11/3
11/4 - 11/17
10/17 - 11/30
10/21 - 11/30
12/2 - 12/31, In Part

IDM
1DM
1DM

10/20 - 11/4
10/19 - 11/7
11/23 - 12/15, In Part

1DM
1DM
1DM
ES
ES

10/17 - 11/5
10/16 - 11/5
12/4 - 12/19, In Part
10/15 - 11/6
12.fj - 12iJ.l, in .l:'art

ES
ES
ES
IDM
ES
ES
1DM
AO
ES

10/21 - 11/9
10/19 - 11/7, In Part
10/19 - 10/21 and
10/22 -- 11/7, In Part
10/18 - 11/6, In Part
10/18 - 10/20 and
10/21 - 11/11, In Part
10/17 - 11/6, In Part
10/17 - 11/6l In Part

*

Based upon combined total resident and non-resident license sales and expressed
in number of licenses.
ES
Either Sex, One Deer.
T
Two Deer, Either Sex, One License.
ESM
Either Sex Multiple (Unlimited licenses and deer per individual).
2DM
Two Deer Multiple, Either Sex (2 licenses and 4 deer per individual).
1DM
One Deer Multiple, Either Sex (2 licenses and 2 deer per individual).
3D
Three Deer, One Each Either Sex 1st and 2nd licenses and 3rd deer coupon with
2nd license.
Note:1DH
evolved into 2D, HC or Two Deer, Hunter's Choice (2 licenses and 2 deer per
individual) and is synonymous.
B. D. Baker
September 1971

�38B

ARCHERY DEER KILL AND SEASONS
WILDLIFE MANAGEMENT. UNIT 62
YEAR

u .u BUCKS

DOES

FAWNS

TOTAL

HUNT AND SEAS ON

1954

ES

10/1

1955

ES
ES
T

10/1
10/15
10/15

-

10/l3
10/31, In Part
10/31, In Part

ES
ESM
ESM

10/1
10/15
11/17

-

10/14
10/31
11/26, In Part

2DM
2DM

10/1
12/1

ES
1DM
ES

9/1
10/15
11/3

ES
1DM

1956

1957

1958

3

1959

0

4

0

2

3

0

6

10/14

11/17
12/31, In Part

-

9/30
11/2
12/31, In Part

9/15
10/17

-

9/30
11/17

1960

0

4

0

4

ES
1DM
?DM

9/10
10/17
P/P

-

9/30
11/30
)2/3', In PE!r"r

1961

5

2

10

17

3D
3D
3D

8/26
10/21
12/2

-

9/10

ES
IDM

·8/25
10/20

ES
1DM
1DM

8/17
10/19
11/23

ES
1DM

8/15
10/17

ES
1DM
1DM

1962

1963

1964

1965

1966

1967 1.1

2
11

15

25

24

33

14

4

16

40

10

13

0

0

5

7

0

6

16

15

36

52

34

52

11130

12/31, In Part

-

9/23
11/4

-

-

9/8
11/7
12/15, In Part

-

9/l3
11/5

8/21
10/16
12/4

-

9/12
11/5
12/19, In Part

ES
ES
ES

8/20
10/15
12/3

-

9/18
11/6
12/11, In Part

ES
ES

8/19
10/21

-

9/17
11/9

�39B
Archery Deer Kill &amp; Seasons - Wildlife Management Unit 62 (Continued)
YEAR

u .u BUCKS

1968

!if

HUNT AND SEASON

DOES

FAWNS

TOTAL

15

7

11

33

ES
ES
ES
1DM

8/17
10/19
10/19
10/22

9/15
11/7, In Part
10/21 and
11/7, In Part

1969 2/

17

0

0

17

AO
ES
ES
lDM

8/16
10/18
10/18
10/2i

9/14
11/6, In Part
10/20 and
11/11, In Part

1970 ~/

44

18

0

62

AO
ES
AO
ES

8/15
9/1
10/17
10/17

8/31
9/20
11/6, In Part
11/6, In Part

1/ Years 1954-1956 free permits were issued to holders of regular big game
hunting licenses by application only. Years 1957-1960 there was no issue of
special permits; archery hunting was allowed to holders of valid deer licenses.
Years 1961-1971 separate archery license regulations have been in effect.
Jj Inconclusive kill data are omitted for years 1954-1957.

]j 237 total res ident and non-res ident archery licenses recorded for unit.
!if 119 total resident and non-resident archery licenses recorded for unit.

2/ 1,744 total hunts recorded for unit.
~/ 271 total hunters recorded for unit.
Hunt symbols are explained as follows:

.No t e :

AO
ES
ESM
lDM

lDMA
T
2DM
3D

=

3DA

=

Antlered Only, One Deer.
Either Sex, One Deer.
Either Sex Multiple (Unlimited licenses and deer per individual).
One Deer Multiple, Either Sex (2 licenses and 2,deer per individual),
which evolved into 2D, HC or ~vo Deer, Hunter's Choice (2 licenses
and 2 deer per individual) and is synonymous.
One Deer Multiple, One Must Be Antler1ess (2 licenses and 2 deer
per individual).
Two Deer, Either Sex, One License.
Two Deer Multiple, Either Sex (2 licenses and 4 deer per individual).
Three Deer, One Each Either Sex (1st and 2nd licenses and 3rd deer
coupon with 2nd license).
Three Deer, One Must Be Antlerless (2 licenses plus 3rd deer coupon
on 2nd 'license).
B. D. Baker
September 1971

�ELK KILL, SEASONS AND HUNTING PRESSURE
WILDLIFE MANAGEMENT UNIT 62

YE~/

HUNTING 2/
PRES SURE-

BULLS

COWS

CALVES

TOTAL

1957

12

0

0

0

0

1958

116

11

0

0

11

AO

10/15 - 11/2

1959

69

23

0

0

23

AO

10/17 - 11/3

1960

97

18

0

0

18

AO

10/17 - 11/6

1961

170

17

0

0

17

AO

10/21 - 11/8

1962

204

19

0

0

19

AO

10/20 - 11/4

1963

243

33

0

0

33

AO

10/19 - 11/7

1964

212

32

0

0

32

AO

10/17 - 11/5

1965

276

31

6

2

39

AO + 30 Asopl/

10/16-11/5

1966

293

44

2

6

52

AO + 50 ASOP]./

10/15 - 11/3

HUNT AND SEASON
25 AO-rl-/

10/15 - 10/31

~ I

1967

352

47

8

0

55

AO + 50 ASOP~f

10/21 - 11/9

1968

536

50

34

4

88

AO + 50 ASOpl/

10/19 - 11/7

1969

591

182

17

0

199

AO + 50 ASOP]/

10/18 - 11/6

1970

536

161

9

0

170

AO + 50 ASO-rl-/

10/17 - 11/6

1/

Unit was closed to elk hunting within recent history prior to 1957.
Based upon combined total resident and non-resident license sales
expressed in number of licenses. Years of antlered only seasons, either
singly or combined with cow permits, very probably yield conservative
hunting pressure statistics. Despite difficulties in verifying the
foregoing assumption, unknown numbers of deer hunters also carry elk
licenses hoping for a bull elk. This happens regardless of how low
elk populations might be. Thus, because of low success and tendency
(above statewide nonreporting percentages that are used in projections)
for unsuccessful elk license holders not to report by card, elk present
in Unit 62 probably receive extra pressure over that indicated by card
return projection years of antlered only seasons.

2/

(Footnotes continued on following page) .

�41B

ELK KILL, SEASONS AND HUNTING PRESSURE
WILDLIFE MANAGEMENT UNIT 62, continued

1/

Unit is combined with Unit 61.

Note:

Hunt symbols are explained as follows:

AO
AOP
ASOP

=

=

Antlered Only.
Antlered Only Permits.
Antlerless Only Permits.
B. D. Baker
September 1971

�42B

ARCHERY ELK KILL AND SEASONS
WILDLIFE MANAGEMENT UNIT 62
HUNTING /
PRESStJREl BULLd/

YEAR 1/

cowd/

CALVEd/

TOTAL~/

HUNT AND SEASON

1965

AO + 30 ASOP

10/16 - 11/5

'1966

AO + 50 ASOP

10/15

1967 1/

ES
AO + 50 ASOP

9/2
- 9/17
10/21 - 11/9

1968 .!i/

11/3

0

3

0

3

ES
AO + 50 ASOP

8/17 - 9/15
10/19 - 11/7

1969

134 i/

0

0

0

0

AO
AO + 50 ASOP

8/16
9/14
10/18 - 11/6

1970

32 ~/

0

0

5

5

AO
ES

8/15
9/1

- 8/31
- 9/20

1/ 1957 was first elk season in recent history; and, the antlered only regular
elk season permits for Units 61 and 62 combined that year could have been
legally obtained, however unlikely, by archers. For the years of 1958-1964,
archery elk hunting was permitted for regular elk license holders during
--------,--J..C:5U..1..0,L

~-

...•.----

~c:a.OVLLO

..1 .....
ULLUCJ..

4....1

.•...•..•...... ..l

aUL.-LCLC:U

.•..._,

VLL..J....Y

,_

.•.. ..!

LC51.1..Lac.....LUUb

"

Ob

WC.L..L

_

.•...

C1b

c

1nt:c

.1.Ul...

.1..'JU-l-..L/V/

,nt:n

_
.L..C-

gular seasons under antlered only-specified antlerless (cow) permit regulations.
~/ Dash denotes no data availab12.

Zero means no kill(s) reccrded.

1/ First year that archery elk licenses were authorized and sold.
~/ Does not include harvest by archers during gun season.

i/ Expressed as number of hunts.
~/ Expressed as total participating resident and nonresident license holders.
Note:

Hunt symbols are explained as follows:

AO

= Antlered Only, One Elk.

ES

= Either

ASOP

= Ant1erless Only Permits (For Units 61 and 62 combined).

Sex, One Elk

B. D. Baker
September 1971

�43B
BLACK BEAR HARVESTS AND SEASONS - WILDLIFE MANAGEMENT UNIT 62
YEAR

HARVEST 1/

1955

39R

8/15 - 10/1; regular deer and regular archery bear seasons.

1956

8R

5/15 - 8/15; regular and post deer and regular archery
bear seasons.

1957

14R

4/1 through regular deer, elk and archery seasons ending
11/15/1957.

1958

24R+5S=29

4/1 through regular deer, elk, and archery seasons and
extended deer season, in par t , ending 11/15/1958.

1959

17R

4/1 through regular deer, elk, and archery seasons ending
11/1/1959.

1960

13R+3.:3=16

4/i - 9/15; regular deer, elk, and archery seasons ending
11/6/1960.

1961

19R+9S=28

4/1 - 9/15; regular deer and elk seasons ending 11/9/1961.

1962

22R+3S=25

4/1 - 9/15; regular deer and elk seasons ending 11/4/1962.

1963

20R+5S=25

4/1 - 9/15; regular deer and elk seasons ending 11/7/1963.

1964

29R+14S=43

4/1 - 9/15; regular deer and elk seasons ending 11/5/1964.

1965

9R

4/1 - 9/30; regular deer and elk seasons ending 11/5/1965.

1966

18R+21S=39

4/1 - 9/30; regular deer and elk seasons ending 11/6/1966.

1967

18

4/1 - 9/30; regular deer and elk seasons ending 11/9/1967.

1968

16R

4/1 - 9/30; regular deer and elk seasons ending 11/7/1968.

1969

3R+13S=16

4/1 - 9/30; regular deer and elk seasons ending 11/6/1969,
in part~ regular deer season ending 11/11/1969, in part.

1970

15R+16S=31

4/1 - 9/30; regular deer and elk seasons ending 11/6/1970.

HUNT AND SEASON ~/, 3/

1/ R=Regular Big Game (Deer and/or Elk) License; S=Bear License for Special Spring,
Summer, or Spring-Summer seasons; Unaugmented numbers represent total animals
harvested not differentiated by season and license controls.
~/ 1955 was the first year of bear license sales and corresponding seasons regulations. From 1955-1959, 1 bear per bear license and/or 1 bear per bear coupon
on either or both deer and elk licenses were allowed in bag limit; from 1960-1965,
same as 1955-1959, except bear license variously invalid after 9/1 or 9/30, as
noted, when bear coupon on deer and elk licenses covered take and possession for
the regular big game seasons; 1966, same as for 1960-1965, except one bear,
hunter's choice, per person per calendar year was the allowable bag limit; for
1967 to the present, 1 bear, hunter's choice, per bear or sportsman's license per
person per calendar year, only during special bear and regular deer and elk seasons.
See deer and elk gun and archery kill and season summary tables herein and/or
(footnotes continued on next page)

�44B

BLACK BEAR HARVESTS AND SEASONS - WILDLIFE MANAGEMENT UNIT 62 (Continued).

!:..I continued
annual regulations for dates and more details of seasons.

11 Dogs were permitted except during bear seasons concurrent with deer and
elk seasons.

B. D. Baker
September 1971

�45B

MOUNTAIN LION HARVESTS AND·SEASONS - WILDLIFEMANAGEMENT·UNIT·62

YEAR 1/

HARVEST I/

HUNT AND SEASON

1965

(0 - Mesa Co., 1965-66)

10/16/65 - 3/31/66 West Slope; 10/23/65 3/31/66 East Slope; 1 either sex.

1966

(5M - Mesa Co., 1966-67)

1/1/66 - 12/31/66 counties west of
Continental Divide plus Jackson, Conejos,
Alamosa, Mineral, Saguache, Rio Grande,
Costilla, Archuleta, Hinsdale, and San
Juan counties; 3 lions per year.
10/22/66 - 2/28/67 counties east of
Continental Divide except those above;
3·lions per year.

1967

No Data

9/1/67 - 3/31/68 statewide; 9/1/67 - 5/31/68
west of State H~.Jy.1113 and north of U. S.
Hwy. #6; 1 either sex per year.

1968

o

9/1/68 - 9/1/69; 1 either sex per year.

1969

o

9/1/69 - 3/31/70; 1 either sex per year.

1970

o

10/17/70 - 11/6/70 concurrent with regular
big game seasons; 1 lion, either sex.

1/ Bounty of $50 per lion was paid within period of May 7, 1929 through March
12, 1965; lions became protected by statute July 1, 1965 when licensed hunting
was authorized.
2/ Data for Mesa County for years 1965-1966 are from Dixon (1967:147). Dixon
(1967: 152) also said that lions were uncommon in Hesa County south of the
Colorado River to indicate that kills originating in the Unit 62 portion of
Mesa County are improbable (but very possible). Data for 1968-1970 are
hopefully accurate under compulsory reporting regulations for hunters, although
the present damage law allows lax reporting of kills of suspected marauders
by offended citizens.

LITERATURE CITED
Dixon, K. R. 1967. Evaluation of effects of mountain lion predation. pp. 147,
152. In Game Res. Rep., July - Part II. Colo. Dept. Game, Fish and
Parks. pp. 73-310.

B. D. Baker
September 1971

�46B

SMALL GAME HARVESTS

AND HUNTING

UNIT 19 ]j,

SMALL GAME MANAGEMENT

NUMBER
SPECIES ]..1

PRESSURE

Jj

OF HUNTERS

HARVEST

1968

1969

1970

1968

1969

1970

Closed

Closed

44

Closed

Closed

344

Blue grouse

465

663

396

1,036

437

604

Chukar

460

417

418

1,072

404

710

1,499

1,107

2,000

10,447

7,737

15,159

No Data

276

483

No Data

2,914

6,6~4

486

1,032

550

1,237

3,426

1,398

Pheasant

3,196

3,588

4,652

14,363

15,900

14,824

Ptarmigan

0

30

34

0

0

34

Sage grouse

95

274

36

144

767

0

49

249

0

157

236

0

116

280

367

211

1,187

686

Band-tailed

pigeon

partridge

Cottontail

rabbit

Dove
Gambel's

quail

Sharp-tailed
Snowshoe

hare

grouse

1/

See Chap. 3- Colo. Game, Fish and Parks Division Laws and Regulations
Handbook; Wildlife Management Unit 62 is wholly contained whithin Small
Game Management Unit 19 and approximates the latter's sout~western onethird.

1/ Colorado Small Game and/or Waterfowl

Harvest Surveys reports for 19551970 also present hunting pressure and kill by region and county, including
Mesa, Delta, Montrose, and Ouray Counties; harvests by Small Game Hanagement Unit are Only available in reports of 1968-1976. .

1/ Since

July 1, 1969, wild turkey has had legal status as a "game bird"
within the broader category of "small game."
Prior to 1969, the turkey
was treated as "big game."
Because of these irregularities and the bird's
relatively high ranking as a game species, separate information has been
presented in the table "Turkey Harvests and Hunting Pressure" elsewhere in
this report.

B. D. Baker
September

1971

�47B
DUCK AND GOOSE HARVESTS AND HUNTING PRESSURE DELTA, MONTROSE, AND OURAY COUNTIES II, 1954-1970

GEESE 'lj

DUCKS
YEAR

EST. NO. HUNTERS

EST. HARVEST

EST. NO HUNTERS

EST. HARVEST

1954

No Data

14,550

No Data

0

1955

1,119

8,437

83

25

1956
1957

No Data
No Data

7,706
6,238

No Data
No 'Data

0
24

1958
1959
1960
1961

980
862
708
55G

6,798
3,564
4,908
2,661

78
0
78
35

175
0
31
0

1962

650

2,167

99

166

1963

499

1,726

40

0

1964

1,053

4,040

104

76

1965

815

3,963

64

0

1966

107 11

5,265

48

0

1967

1,014

3,958

0

0

1968

982

8,090

13

19

1969

1,082

6,064

0

0

1970

1,081

8,634

25

19

~I Totals for these three counties are best available but only roughly comparable
indicators of harvests and pressures for Unit 62. Mesa County data are.omitted
in assuming that there is negligible pressure on that county's portion of Unit 62.

11 Accuracy of data is questionable and probably due to sampling difficulties
associated with very small populations, geese and goose hunters alike.

11 Obvious inaccuracy.

About 1,000 hunters would be more correct.

B. D. Baker
September 1971

�TURKEY HARVESTS AND HUNTING PRESSURE
WILDLIFE MANAGEMENT UNIT 62 11

HARVEST
YEAR

TOMS

19541955
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969 1/
1970

HENS

POULTS

TOTAL

0

0

0

0

2
0
0
0
2
53
52
35
56
45
18
7
5
2
0
0

2
0

0
4
0
3
2
16
124
100
54
149
135
37
18
8
5
2
0

0

3
2

6
52
31
16
56
58
8
7
3
0
0

0

0

0
8
19
17
3
37
32
11
4
0
3
2
0

1/ For years 1954-1967; effective 1968, Unit 62 became
Game Management
area.

]j

Expressed

Unit 19, so data for years

as "total hunters"

HUNTING
PRESSURE

u

N.D.
N.D.
N.D.

3
11
29
261
203
149
221
268
171
58
18
15
N.D.
14

part of Small

1968-1970 apply to the latter

for years of records.

N.D. == No Data.

1/

The legal definition of wild turkey was changed July 1, 1969 from
that of "big game" to that of "game bird," the latter also within the
broader category of "small game."

B. D. Baker
September. 1971

�GAME SPECIES INTRODUCTIONS

- WILDLIFE MANAGEMENT

UNIT 62

Source of Stock

Species

Date

Males

Females

Young

Total

Release Area

Pronghorn

2-16-49

5

12

0

17

l.;rells
Gulch near U. S. Hwy.
50 NW Delta 1/.

12-30-49

1

5

7

13

" "

"

rr

"

"

2-11-50

6

10

0

16

" "

"

"

"

"

1-29-70

-

-

-

50

" "

II

"

"

"

Wild-trapped; Chico
Basin E. of Pueblo
Wild-trapped; Ingle
Rch.E. of Co10.Spgs.
Wild-trapped; 20 mi.
W. of Grover
Wild-trapped; Chico
Basin

Reference
Elliott (1949)
Bear (1969)
Till (1950)
Bear (1969)
XiII (1950)
Bear (1969)
Mink (pers. comm ,
1972)

-------------------------------------------------------------------------------------------------------------------------Sandfort's files
Wild-trapped from
Kannah Cr.,3/4 mi. from
16
8
8
Chukar 2/ 1-02-52
11-09-52
12-09-52

14
6

16
11

-

30
17

12-18-52
3-03-54
3-24-55
11-09-56

11

7

18
3
99
30

confl. W/Gunnison River.
Dry Creek, Montrose Co.
Kannah Cr.,3/4 mi. from
confl. W/Gunnison River.

"
"

"
"

"
"

Sandfort's files
Sandfort's files

II

II

II

II

"

II

"

"

Sandfort's files
8andfort's files
Sandfort's files
Sandfort's files

-

-

59
15

40
15

-

3-21-63
4-20-64
7-14-64
3-16-67

-

-

-

-

-

-

150
258
55
127

1938

-

-

-

15

Musser's Rch.-Escalante

1938

-

-

-

15

Head Spring Creek

12-30-48

3

9

-

12

Log Hi11 Mesa

1-15-58
3-07-58

2
2

5
1

-

7
3

Big Dominguez Canyon

-

-

"
"

II

"

"

"

II
II

Dry Creek, Montrose Co.
Cottonwood Cr.; 833, T50N,
R13W.
Vicinity of Montrose •.
Escalante Creek.
II

II

Escalante Cr.

Game farm.
Wild-trapped from
Escalante Cr.
Game farm.
II

II

II

"

"

Roubideau Creek.

"

8andfort's
Sandfort's
Sandfort's
Sandfort's

files
files
files
files

--------------------------------------------------~----------------------------------------------------------------------Hild
turkey

l/Transplant

-

"

"

Fks. Unknown

"
Wild-trapped from
Turkey Ranch.

"

"
"

"
"

"

"

Myers (pers. comm.
1972)
Myers (pers . comm.
1972)
Burget (1960)
Burget (1960)
Burget (1960)

actually made in Unit 411 but close enough to be effective for Unit 62.

~/No records are available for years prior to 1952. Division transplanting program9
very likely included releases in Unit 62 and was the origin of trappable populations

using game farm stock in the 1940's,
in Escalante Creek.
.po.
1..0

b:l

�SOB

GAME SPECIES INTRODUCTIONS - WILDLIFE MANAGflfENT UNIT 62 (Continued)

Literature Cited
Bear, G. D. 1969. Antelope transplants in Colorado. Game Inf. Leaflet
No. 70. Colo. Div. Game, Fish and Parks. October. 3 pp.
Burget, M. L. 1960. Wild turkey development, pp. 11-42. In Quart. Rep.,
July. Colo. Dept. Game and Fish. 110 pp. (processed)-.Elliott, R. R. 1949. Antelope restoration, pp. 19-24. In Quart. Prog.
Rept., April. Colo. Dept. Game and Fish. 98 pp. (processed).
Till, C. E. 1950.
Rept., April.

Antelope restoration, pp. 17-21. In Quart. Prog.
Colo. Dept. Game and Fish. 101 pp-.-(processed).

B. D. Baker
November 1971

�51B·

BIG GAME MAMMAL
WILDLIFE

RESEARCH

MANAGEMENT

REFERENCES
UNIT 62

BLACK BEAR
Boyd, R. J. 1970. Deer-elk investigations.
pp. 133-213. In Game Res.
Rep., July - Part II. Colo. Div. Game, Fish and Parks.
pp. 127-286.
Denney, R. N., and D. L. Gilbert.
1952. Fur resources and bear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.
Gilbert, D. L. 1951. Fur resources study and b~ar investigations.
pp. 1-8.
In Quart. Prog. Rep., July.
Colo. Dept. Game and Fish.
102 pp.

1951. Fur resources study and bear investigations.
In Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish.
1953.
Rep., Jan.

Fur resources and bear studies.
pp. 59-60.
Colo. Dept. Game and Fish.
141 pp.

Gilbert, P. E. 1946. Bear studies.
Dept. Game and Fish.
4 pp.

1948.
Rep., Oct.

pp. 1-9.
117 pp.
In Quart. Prog.

In Quart. Prog. Rep., June.

Deer-elk-bear investigations.
pp. 30-34.
Colo. Dept. Game and Fish.
80 pp.

Colo.

In Quart. Prog.

1952.

Checking station survey.
pp • .39-43. In Quart. Prog. Rep.,
Colo. Dept. Game and Fish. 60 pp.

Jan.

pp. 59-61.
Checking station survey,
Colo. Dept. Game and Fish. 99 pp.

1953.
April.
Gilbert,

128.

In Quart. Prog. Rep.,

P. F., and G. E. Rogers.
1954. Checking station survey.
In Quart. Prog. Rep., July. Colo. Dept. Game and Fish.

Hammit, H. C. 1950. Deer-elk~bear investigations.
pp. 10-14.
Proj. \o.1-38-R.
Oct. Colo. Dept. Game and Fish.
21 pp.
Hunter, G. N. 1965. Colorado big game harvest,
Game, Fish and Parks.
41 pp.

1959-1965.

pp. 125182 pp.

Fed. Aid

Colo. Dept.

\0.1. T.
1965. Colorado long range game species management plans,
1965-1975, for black bear. Colo. Dept. Game, Fish and Parks. 6 pp.

McKean,

Tigner, J. R., and D. L. Gilbert.
1960. A contribution toward a bibliography on the black bear.
Colo. Dept. Game and Fish. Tech. Publ. No.
6.
43 pp.

�52B

ELK
Boyd, R. J. 1970. Deer-elk investigations. pp. 133-213. In Game Res.
Rep., July - Part II. Colo. Div. Game, Fish and Parks. pp. 127-286.
Denney, R. N. 1965. Colorado long range game species management plans,
1965-1975, for elk. Colo. Dept. Game, Fish and Parks. 61 pp.
Gilbert, P. F. 1953. Checking station survey. pp. 59-61.
Prog. Rep., April. Colo. Dept. Game and Fish. 99 pp.

In Quart.

Gilbert, P. F., and G. E. Rogers. 1954~ Checking station survey, pp.
125-128. IE. Quart. Prog. Rep , , July. Colo. Dept. Game and Fish.
182 pp.
Hammit, H. C. 1951. Deer-elk-bear investigations. pp. 16-22. In
Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 33 pp.
Hunter, G. N. 1965. Colorado big game harvest, 1959-1965.
Game, Fish and Parks. 41 pp ,

Colo. Dept.

Rogers, G. E. 1951. Checking station survey. pp. 32-33.
Rep., Jan. Colo. Dept. Game and Fish. 33 pp.

In Quart. Prog.

MULE DEER
Anderson, A. E.
Rep , , Jan.

1959. Deer-elk investigations. pp. 113-128.
Colo. Dept. Game and Fish. 128 pp.

In Quart.

1960. Effects of sagebrush eradication by chemical means on deer
and related wildlife. Fed. Aid Job Compl. Rep., Proj. W-38-R-13. July.
Colo. Dept. Game and Fish. 72 pp.
1960. Distribution and abundance indices of selected biota in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Univ., Ft. Collins. 130 pp.
1966. An investigation of 2,4-D application to sagebrush within
a mountain shrub-conifer complex, Uncompahgre National Forest. In
Game Res. Rep., July - Part III. Colo. Dept. Game, Fish and Parks.
pp. 345-420.
1969. 2,4-D, sagebrush/and mule deer-cattle use of upper winter
range. Special Rep., No. 21. July. Colo. Div. Game, Fish and Parks.
21 pp.
1969. Sagebrush vegetation before and six years after 2,4-D
application. Outdoor Facts, No. 67. Colo. Div. Game, Fish and Parks.
4 pp.
Baker, B. D. 1955. Deer-elk investigations. pp. 149-157. In Quart. Prog.
Rep., July - Part II. Colo. Dept. Game and Fish. pp. 117-175.

�53B

1956. Deer-elk investigations, pp. 43-64. In Quart. Prog. Rep.,
July - Part I. Colo. Dept. Game and Fish. pp. 1-102.
1957. A directory of Colorado big game exclosures concerning their
locations, studies and other general information. Spec. Rep., Fed. Aid
Proj. W-38-R, Jan. Colo. Dept. Game and Fish. 60 pp.
1961. A directory of Colorado big game exclosures concerning program
history, study techniques, location and other general information.
Spec. Rep. - revised edition, Fed. Aid Proj. W-10l-R-3, Jan. Colo. Dept.
Game and Fish. 51 pp.
1964. Game range investigations. pp. 3-15. In Game Res. Rep. ,
Jan. - Part I. Colo. Dept. Game, Fish and Parks. pp. l-12l.
1966. Game range investigations. pp. 53-67.
July - Part I. Colo. Dept. Game, Fish and Parks.

In Game Res. Rep. ,
93 pp.

1970. Game range investigations. pp. 15-58. In Game Res. Rep.,
July - Part I. Colo. Div. Game, Fish and Parks. 126 pp.
Boyd, R. J. 1970. Deer-elk investigations. pp. 133-213. In Game Res. Rep.,
July - Part II. Colo. Div. Game, Fish and Parks. pp. 127-286.
Carhart, A. H. 1940. Deer-elk survey. Vol. 4.
Colo. Game and Fish Commission. 19 pp.

Fed. Aid Proj. W-4-R.

Gilbert~ P. F. 1948. Deer-elk-bear investigations. pp. 14-18.
Prog. Rep., April. Colo. Dept. Game and Fish. 54 pp.

In Quart.

1948. Deer-elk-bear investigations. pp. 31-34.
Rep., July. Colo. Dept. Game and Fish. 83 pp.

In Quart. Prog.

1948. Deer-elk-bear investigations. pp. 30-34.
Rep., Oct. Colo. Dept. Game and Fish. 80 pp.

In Quart. Prog.

1949. Deer-elk-bear investigations. pp. 42-45.
Rep., Jan. Colo. Dept. Game and Fish. 67 pp.

In Quart. Prog.

pp. 47-5l.
98 pp.

In Quart. Prog.

1949. Deer-elk-bear investigations. pp. 33-36.
Rep., July. Colo. Dept. Game and Fish. 84 pp.

In Quart. Prog.

1950. Deer-elk-bear investigations. pp. 41-44.
Rep., Jan. Colo. Dept. Game and Fish. 66 pp.

In Quart. Prog.

1950. Deer-elk-bear investigations. pp. 60-64.
Rep., April. Colo. Dept. Game and Fish. 101 pp.

In Quart. Prog.

1949. Deer-elk-bear investigations.
Rep., April. Colo. Dept. Game and Fish.

�Sl:·B

1950. Deer-elk-bear investigati.ons. Three Year Sum. Rep.,
1947-1950. July. Colo. Dept. Game and Fish. 21 pp.
1952. Checking station survey. pp. 39-43.
Rep., Jan. Colo. Dept. Game and Fish. 60 pp.

In Quart. Prog.

1952. Deer-elk investigations. pp , 41-64,66-67, 69-82, 83-84.
In Quart. Prog. Rep., July. Colo. Dept. Game and Fish. 171 pp.
1953. Checking station survey. pp. 59-61. In Quart. Prog.
Rep., April. Colo. Dept. Game and Fish. 99 pp.
1954.
Rep ,. , July.

Deer-elk investigations. pp. 129-143.
Colo. Dept..Game and Fish. 182 pp.

In Quart. Prog.

Gilbert, P. F., and G. E. Rogers. 1954. Checking station survey. pp.
125-128. In Quart. Prog. Rep., July. Colo. Dept. Game and Fish.
182 pp.
Gilbert, P. F., et a1. 1951. Deer-elk investigations. pp. 109-110.
Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish. 117 pp.
Hammit, H. C. 1950. Deer-elk-bear investigations. pp. 10-14.
Proj. W-38-R. Oct. Colo. Dept. Game and Fish. 21 pp.
1951. Deer-elk-bear investigations. p p , 16-22.
W-38-R. Jan. Colo. Dent. Game and Fish. 33 pp.

In

Fed. Aid

Fed. Aid Proj.

Hammit, H. C., and P. F. Gilbert. 1950. Deer-elk-bear investigations.
pp. 35-37. In Quart. Prog. Rep., July. Colo. Dept. Game and Fish.
80 pp.
Hunter, G. N. 1965. Colorado big game harvest, 1959-1965. Special Mgmt.
Rep., No.1.
Colo. Dept. Game, Fish and Parks. 41 pp.
Jeep, F. T., and L. E. Riordan. 1947. Deer-elk studies. In ~uart. Prog.
Rep., Jan. - Part IV. Colo. Dept. Game and Fish. 7 pp ,
Kufe1d, R. C. 1968. Game range investigations. In Game Res. Rep., July _
Part I. Colo. Div. Game, Fish and Parks. pp. 1-121.
1969. Game range investigations. pp. 299-304. In Game Res. Rep.,
July - Part III. Colo. Div. Game, Fish and Parks. pp. 249-393.
1970. Game range investigations. pp. 59-94. In Game Res. Rep.,
July - Part I. Colo. Div. Game, Fish and Parks. pp. 1-126.
McKean, W. T. 1958. Deer-elk investigations. pp. 1-23. In Quart. Rep.,
July - Part 1. Colo. Dept. Game and Fish. pp. 1-103.
Medin, D. E. 1965. Colorado long range game species management plans,
1965-1975, for mule deer. Colo. Dept. Game, Fish and Parks. 4 pp.
Minnich, D. W. 1969. Vegetative response and pattern of deer use following
chaining of pinon and juniper forests. J. Range Mgmt., Abstracts of
Papers, 22nd. Ann. Htg. Amer. Soc. Range Hgmt.

59 pp.

�55B

Riordan, L. E. 1948. Deer-elk-bear investigations. pp. lLf- 30.
Quart. Prog. Rep. , Jan. Colo. Dept. Game·and Fish. 49 pp.
Rogers, G. E. 1951. Checking station survey. pp. 32-33.
Proj. W-38-R, Jan. Colo. Dept. Game and Fish. 33 pp.

In

Fed. Aid

Shepherd, H. R. 1966. Game range investigations. pp. 91-92. In Game
Res. Rep., July - Part I. Colo. Dept. Game, Fish and Parks. pp. 1-93.
Williams, J. E. 1951.
Prog. Rep ,, July.

Deer-elk investigations. pp. 68-71.
Colo. Dept. Game and Fish. 102 pp.

In Quart.

1956. Deer-elk investigations. pp. 1-10. In Quart. Prog.
Rep., Jan. Colo. Dept. Game and Fish. 119 pp.
Williams, J. E., et al. 1952.
Quart. Prog. Rep., April.

Deer-elk investigations. pp. 57-62.
Colo. Dept. Game and Fish. 62 pp.

In

PRONGHORN
Anderson, A. E. 1965. Colorado long range management plans, 1965-1975,
for antelope. Colo. Dept. Game, Fish and Parks. 6 pp.
Douglas, G., and C. E. Till. 1950. Antelope surveys and investigations.
Three Year Sum. Rep., 1947-1950, July. Colo. Dept. Game and Fish. 9 pp.
DcugLas ,': G. ,2nd

S. Ogilvie.

1952.

In Quart. Prog. Rep., July.

July.

Antelope

1953. Antelope restoration.
Colo. Dept. Game and Fish.

Elliott, R. R. 1947.
Prog. Rep., Oct.

r as t or at Lon,

pp.

Colo. Dept. Game and Fish.
pp. 123-128.
129 pp.

168-17L

171 pp.

In Quart. Prog. Rep.,

Antelope investigations. pp. 1, 6-8.
Colo. Dept. Game and Fish. 22 pp.

.1949. Antelope restoration. pp. 19-24, 83-91.
Rep., April. Colo. Dept. Game and Fish. 98 pp.

In Quart.

In Quart. Prog.

Hoover, R. L. 1955. Antelope restoration. pp. 39-48. In Quart. Prog.
Rep., July - Part I. Colo. Dept. Game and Fish. pp. 1-115.
Hoover, R. 1., C. E. Till, and S. Ogilvie. 1959. The" antelope of Colorado.
Tech. Bul., No.4.
Colo. Dept. Game and Fish. 110 pp.
Ogilvie, S., and R. L. Hoover. 1954. Antelope restoration.
In Quart. Prog. Rep.; July. Colo. Dept. Game and Fish.

pp. 94-104.
182 pp.

Till, C. E. 1950. Antelope restoration. pp. 17-21. In Quart. Prog.
Rep ;, April. Colo. Dept. Game and Fish. 101 pp.

1. C. Carlson
October 1971

�56B

SHALL GAME HAMMAL RESEARCH REFERENCES
WILDLIFE MANAGEHENT UNIT 62

COTTONTAIL RABBIT
Anderson, A. E.
Rep., Jan.

1959. Deer-elk investigations. pp. 113-128.
Colo. Dept. Game and Fish. 128 pp.

In Quart.

1960. Deer-elk investigations, effects of sagebrush eradication
by chemical means on deer and related Hildlife. Job Completion Rep_,
July. Colo. Dept. Game and Fish. 72 pp.
1960. Distribution and abundance indices of selected biota in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Univ., Ft. Collins. 130 pp.
1966. An investigation of 2,4-D application to sagebrush Hithin
a mountain shrub-conifer complex, Uncompahgre National Forest. Game
Res. Rep., July - Part III. Colo. De.pt.Game, Fish and Parks. pp.
345-420.
1969. 2,4-D, sagebrush and mule deer-cattle use of upper Hinter
range. Special Rep., No. 21, July. Colo. Div. Game, Fish and Parks.
21 pp.
Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 60 pp.
Sandfort, W. W. 1951. Game bird surveys. pp. 45-47.
Rep., July. 'Colo. Dept. Game and Fish. 102 pp.

In Quart. Prog.

Shepherd, H. R. 1965. Colorado long range game species management plans,
1965-1975, for cottontail rabbit. Colo. Dept. Game, Fish and Parks.
12 pp.
SNm\1SHOEHARE
Shepherd, H. D. 1965. Colorado long range game species management plans,
1965-1975, for snOHshoe hare. Colo. Dept. Game, Fish and Parks. 11 pp.

1. C. Carlson
October 1971

�57B

NONGAME MAMMAL RESEARCH
HILDLIFE

MANAGEMENT

REFERENCES
UNIT 62

BOBCAT

1965. Colorado long range game species management plans,
1965-1975, for predators.
Colo. Dept. Game, Fish and Parks.
21 pp.

Boyd, R. J.

Burget, M. L. 1957. The wild turkey in Colorado.
Dept. Game and Fish. January.
68 pp.

pp. 53-54.

Colo.

Burget, M. L., and D. M. Hoffman.
1951. Wild turkey surveys and investigations.
pp. 11-29. In Quart. Prog. Rep., Oct. Colo. Dept. Game
and Fish.
117 pp.
Rutherford, W. H. 1965. Colorado long range game species management
plans, 1965-1975, for furbearers.
Colo. Dept. Game, Fish and Parks.
12 pp.

COYOTE

1965. Colorado long range game species management plans,
1965-1975, for predators.
Colo. Dept. Game, Fish and Parks.
21 pp.

Boyd, R. J.

Burget, H, L. 1949. Wild turkey surveys and investigations.
pp , 91-106.
In Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish.
141 pp.
: .;.

1957.
January.

The wild turkey in Colorado.
68 pp.

Colo. Dept. Game and Fish.

Burget, M. L., and D. M. Hoffman.
1951. Wild turkey surveys and investigations.
pp. 11-29. In Quart. Prog. Rep., Oct. Colo. Dept. Game and
Fish.
117 pp ,

1956. Wild turkey investigations.
pp. 107-131. In Quart. Prog.
Rep., July - Part II. Colo. Dept. Game and Fish. pp. 103-138.
Denney, R. N., and D. L. Gilbert.
1952. Fur resources and bear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.

SKUNK

1965. Colorado long range game species management plans, 19651975. for predators.
Colo. Dept. Game, Fish and Parks.
21 pp.

Boyd, R. J.

Ourget, M. L. 1957.
Fish. January.

The wild turkey in Colorado.
68 pp.

Colo. Dept. Game and

�S8B

Burget. M. 1., and D. M. Hoffman. 1951. Wild turkey surveys and investig at i.ons ,
pp , 11-29. In Quart. Prog. Rep , , Oct. Colo. Dept. Game
and Fish. 117 pp.
1954. Wild turkey investigations. pp , 25-30.
Ren., July. Colo. Dept. Game and Fish. 182 pp.

In Quart. Prog.

Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.
Rutherford. \\1. H. 1965. Colorado long range game species management plans,
1965-1975, for furbearers. Colo. Dept. Game. Fish and Parks. 12 pp.

L. C. Carlson
October 1971

�59B

FURBEARER

RESEARCH

REFERENCES

WILDLIFE

MANAGE}-llNTUNIT 62

MINK
Denney, R. N. 1950. Fur resource surveys, pp. 75-80.
Rep., July. Colo. Dept. Game and Fish.
80 pp.

In Quart. Prog.

survey. Vol. 2 .. Jan.

Colo. Game

Remington, J. D. 1953. Fur resources and bear studies.
Quart. Prog. Rep. , Oct. Colo. Dept. Game aud Fish.

pp. 17-23.
60 pp.

Nemanic, W. M. 1942. Fur resource
and Fish Commission.
6 pp.

In

Rutherford, W. H. 1965. Colorado long range game species management
plans, 1965-1975. for furbearers.
Colo. Dept. Game, Fish and Parks.
12 pp.

MUSKRAT
Denney, R. N. 1950. Fur resource survey.
pp. 57-66.
Rep ,, Jan. Colo. Dept. Game and Fish.
66 pp.

1950.
April.

Fur resource survey.
pp. 93-101.
Colo. Dept. Game and Fish.
101 pp.

1950.
July.

Fur resource survey.
pp. 75-80.
Colo. Dept. Game and Fish.
80 pp.

1951. Fur resource survey.
Dept. Game and Fish.
8 pp.

In Quart. Prog.

In Quart. Prog. Rep.,

In Quart. Prog. Rep.,

In Quart. Prog. Rep., Jan.

Colo.

Denney, R. N .• and D. L. Gilbert.
1952. Fur resources and bear studies.
pp , 1-3l. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.

60 np ,
Nemanic, W. M. 1942.
Fish Commission.

Fur resource
6 pp.

survey. Vol. 2, Jan.

Remington, J. D. 1953. Fur resources and bear studies.
Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish.

Colo. Game and

17-23.
60 pp.

pp .

In

W. H. 1965. Colorado long range game species management plans,
1965-1975, for furbearers.
Colo. Dept. Game, Fish and Parks.
12 pp.

Rutherford,

WEASEL
Denney. R. N. 1950. Fur resource survey.
pp. 93-101.
April.
Colo. Dept. Game and Fish.
101 pp.

In Quart. Prog. Rep.,

�60B

____

1950.
July.

Fur resource survey.
pp , 75-80.
Colo. Dept. Game and Fish.
80 pp ,

In Quart. Prog. Rep ,,

Denney, R. N., and D. L. Gilbert.
1952. Fur resources and bear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.
60 pp.
Nemanic, H. M. 1942.
Fish Commission.

Fur resource
6 pp.

survey.

Vol. 2 •. .Ian ,

Remington, J. D. 1953. Fur resources and bear studies.
Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish.

Colo. Game and

pp , 17-23.
60 pp.

In

W. H. 1965. Colorado long range game species management plans,
1965-1975. for furbearers.
Colo. Dept. Game, Fish and Parks.
12 pp.

Rutherford,

L. C. Carlson
October 1971

�61B

GAME BIRD RESEARCH REFERENCES
WILDLIFE MANAGEMENT UNIT 62

BAND-TAILED PIGEON
Braun, C. E. 1970. The band-tailed pigeon. pp. 26-29.
No.5, Sept.-Oct. Colo. Div. Game, Fish and Parks.

Colo. Outdoors,
48 pp.

1970. Migratory bird investigations. pp. 151-171.
Rep., Oct. Colo. Div. Game, Fish and Parks. 171 pp.

In Game Res.

Neff, J. A., and J. C. Culbreath. 1946. Colorado band-tailed pigeon.
pp. 1-24. Fed. Aid Proj. 4-R. Colo. Dept. Game and Fish. 24 pp.

BLUE GROUSE
Aldrich, J. 1-1., and A. J. Duvall. 1955. -Distribution of American
gallinaceous game birds. U.S.D.I. Fish and Wildl. Ser .• Circular
No. 34. 23 pp.
Anc1ersono

A. E.

Rep., Jan.

]q,)q.

Deer-elk

investiERtionl':.

Colo. Dept. Game and Fish.

pp.

11.'1-128.

Tn C)llRrt.

128 pp.

1960. Effects of sagebrush eradication by chemical means on deer
and related wildlife. Job Cornp L, Rep., July. Colo. Dept. Game and
Fish. 72 pp .
1960. Distribution and abundance indices of selected biota in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Univ•• Ft. Collins. 130 pp.
1966. An investigation of2,4-D application to sagebrush within a
mountain shrub-conifer complex, Uncompahgre National Forest. In Game
Res. Rep., July - Part III. Colo. Dept. Game, Fish and Parks. pp.
345-420.
1969. 2,4~D, sagebrush and mule deer-cattle use of upper winter
range. Spec. Rep. No. 21, July. Colo. Div. Game, Fish and Parks. 21 pp.
Myers, G. T.
season.

1965. Evaluation of a combined blue grouse-wild turkey hunting
Outdoor Facts, No. 27. Colo. Dept. Game, Fish and Parks. 1 p.

Rogers, G. E. 1962. Game bird survey. pp. 121-131.
Part 1. Colo. Dept. Game and Fish. pp. 1-160.

In Quart. Rep., July -

1963. Game bird survey. pp. 157-168,191. In Game Res. Rep., Oct.Part II. Colo. Dept. Game, Fish and Parks. pp. 109-241.

�62B

1965. Colorado long range game species management plans, 19651975, for blue grouse, sage grouse, sharp-tailed grouse, ptarmigan.
Colo. Dept. Game, Fish and Parks. 16 pp.
196B. The blue grouse in Colorado.
Dept. Game, Fish and Parks. 63 .pp.

Tech. Pub1., No. 21.

Sandfort, W. W. 1950. Game bird surveys. p. 37.
Oct. Colo. Dept. Game and Fish. 44 pp .

Colo.

Fed. Aid Proj. W-37-R.

CHUKAR
Aldrich, J. W., and A. J. Duvall. 1955. Distribution of American gallinaceous
game birds. D.S.D.I. Fish and Wild1. Ser., Circular No. 34. 23 pp.
Bartmann, R. M. 1963. Game bird surveys. pp. 33-40,45-69.
Rep., Oct. - Part I. Colo. Dept. Game, Fish and Parks.

In Game Res.
pp ,

I-lOB ..

1964. Game bird survey. pp. 15l-17B. In Game Res. Rep., AprilPart II. Colo. Dept. Game, Fish and Parks. pp. 71-233.
Evans, R. L., and W. W. Sandfort. 1957. Game bird survey.
Quart. Rep,, Oct. Colo. Dept. Game and Fish. 116 pp ,

pp. 55-67.

In

Hurd, C. L., and W. W. Sandfort.

pp. 85-88.

In

1955.

Game bird survey.

lOR
-

1956. Game bird survey. pp. 75-81.
Colo. Dept. Game and Fish. 101 pp.

nn
rr·

In Quart. Prog. Rep., April.

Manning, D. D. 1962. Game bird survey. pp. 9-16, 25-31. In Quart. Rep.,
July - Part I. .Colo. Dept. Game and Fish. pp , 1-160.
Miller, J. 1959. Game bird survey. pp. 99-107,115-124.
Oct. Colo. Dept. Game and Fish. 124 pp.

In Quart. Rep ,,

Miller, J., and W. W. Sandfort. 1959. Game bird survey. pp. 67-73.
Quart. Rep., Jan. Colo. Dept. Game and Fish. 128 pp.
Sandfort, W. W. 1950. Game bird survey. pp. 51-52.
April. Colo. Dept. Game and Fish. 101 pp.
1950. Game bird survey. pp. 37-39.
Colo. Dept. Game and Fish. 44 pp.

In

In Quart. Prog. Rep.,

Fed. Aid Proj. W-37-R, Oct.

1951. Chukar census. pp. 22-31. Spec. Rep., Glenwood Sprgs. Wildl.
Conf., Jan. 16-19. Colo. Dept. Game and Fish. 85 pp.
1951. Game bird survey. pp. 37-41.
Colo. Dept. Game and Fish. 102 pp.

In Quart. Prog. Rep., July.

1951. Game bird survey. pp. BO-82.
Colo. Dept. Game and Fish. 117 pp.

In Quart. Prog. Rep., Oct.

�63B

1952. Game bird survey. pp. 40~56. In Quart. Prog. Rep.,
April. Colo. Dept. Game and Fish. 62 pp.

Oct.

1952. Game bird survey. pp. 93-95. In Quart. Prog. Rep.,
Colo. Dept. Game and Fish. 105 pp.

1953. Game bird survey. pp. 81-89. In Quart. Prog. Rep.,
April. Colo. Dept. Game and Fish. 99 pp.
1953. Game bird survey. p. 107. In Quart. Prog. Rep.,
July. Colo. Dept. Game and Fish. 129 pp.
1954. Game bird survey. pp. 13-33. In Quart. Prog. Repp.,
April. Colo. Dept. Game and Fish. 79 pp.
1954. Game bird survey. pp. 67-72. In Quart. Prog. Rep.,
July. Colo. Dept. Game and Fish. 182 pp.
1955. Game bird survey. pp. 17-18.
Jan. Colo. Dept. Game and Fish. 94 pp.

In Quart. Prog. Rep.,

1955. Game bird survey. pp. 82-84. In Quart. Prog. Rep.,
July - Part I. Colo. Dept. Game and Fish. pp. 1-115.
1957. Game bird survey. pp. 29-38,45-51. In Quart. Prog.
Rep., July - Part I. Colo. Dept. Game and Fish. pp. 1-119.
1958. Game bird survev. DD. 129-134. In Quart. Rep •. July Part II. Colo. Dept. Game and Fish. pp. 105-181.
1959. Game bird survey. pp. 53-65.
Colo. Dept. Game and Fish. 128 pp.

In Quart. Rep., Jan.

1959. Game bird survey. pp. 33-36. In Quart. Rep., April Part I. Colo. Dept. Game and Fish. pp. 1-116.
1960. Game bird survey.
Dept. Game and Fish. 74 pp.

pp. 49-54.

1960. Game bird survey. pp. 91-94.
Colo. Dept. Game and Fish. 161 pp.

In Quart. Rep., Jan.

Colo.

In Quart. Rep., April.

1961. Game bird surveys. pp. 119-126,139-155. In Quart. Rep.,
April - Part II. Colo. Dept. Game and Fish. pp. 97-208.
1962. Game bird survey. pp. 17-23,33-44. In Quart. Rep., July Part I. Colo. Dept. Game and Fish. pp. 1-160.
1965. Colorado long range game species management plans, 19651975, for chukar partridge. Colo. Dept. Game, Fish and Parks. 18 pp.
Sandfort, W. W., and D. Nolting. 1952. Game bird survey. pp. 89-91.
Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish. 105 pp.
White, C., and J. E. Hi1liams. 1960. Escalante. pp. 21-23.
No.5, Sept.-Oct. Colo. Dept. Game and Fish. 32 pp.

In

Colo. Outdoors,

�64B

GAMBEL'S QUAIL
Sandfort, W. H. 1950. Game bird surveys. pp. 17-21.
Rep , , July. Colo. Dept. Game and Fish. 80 pp.

In Quart. Prog.

Oct.

1950. Game bird survey. pp. 37-39.
Colo. Dept. Game and Fish. 44 pp.

Fed. Aid Proj. W-37-R,

Oct.

1951. Game bird survey. pp. 80-82. In Quart.
Prog. Rep. ,
Colo. Dept. Game and Fish. 117 pp.

Oct.

1952. Game bird survey. pp. 93-95. In Quart. Prog. Rep. ,
Colo. Dept. Game and Fish. 105 pp.

1953. Game bird survey. pp. 81-89. In Quart. Prog. Rep.,
April. Colo. Dept. Game and Fish. 99 pp.
1965. Colo. long range game species management plans, 19651975, for gambe1's quail. Colo. Dept. Game, Fish and Parks. 8 pp.
Sandfort, W. W., and D. Nolting. 1952. Game bird surveys.
In Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish.

pp. 89-9l.
105 pp.

Sandfort, W. W., and H. M. Swope. 1954. Game bird survey.
In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish.

pp. 19-29.
113 pp.

MOUNTAIN QUAIL
,

Hoffman, D. M. 1969. Game bird survey. pp. 155-158.
April. Colo. Div. Game, Fish and Parks. 181 pp.
1970. Game bird survey. pp. 167-169.
Colo. Div. Game, Fish and Parks. 198 pp.

In Game Res. Rep.,

In Game Res. Rep., April.

Rogers, G. E. 1965. Game bird survey, pp. 217-225. In Game Res. Rep.,
April - Part II. Colo. Dept. Game, Fish and Parks. pp. l75~235.
1967. Game bird survey. pp. 215-218. In Game Res. Rep., April Part II. Colo. Dept. Game, Fish and Parks. pp. 107-241.
1968. Game bird survey. pp. 113-116.
Colo. Dept. Game, Fish and Parks. 131 pp.

In Game Res. Rep., April.

MOURNING DOVE
Nolting, D., et a1. 1952. Game bird survey. pp. 99-101.
Rep., Oct. Colo. Dept. Game and Fish. 105 pp.
Sandfort, W. H. 1950. Game bird survey. pp. 41-42.
Oct. Colo. Dept. Game and Fish. 44 pp.

In Quart. Prog.

Fed. Aid Proj. W-37-R,

"

�65B

1951. Game bird survey. pp. 42-ff3. In Quart. Prog. Rep.,
July. Colo. Dept. Game and Fish. 102 pp.
1951. Game bird survey. pp. 83-84. In Quart. Prog. Rep.,
Oct. Colo. Dept. Game and Fish. 117 pp.
1954. Game bird survey. pp. 23-24. In Quart. Prog. Rep.,
Oct. Colo. Dept. Game and Fish. III pp.
1955. Game bird survey. pp. 91-92. In Quart. Prog. Rep.,
July - Part I. Colo. Dept. Game and Fish. 115 pp.
pp. 103-105.
Sandfort, W. W., and D. Nolting. 1952. Game bird survey.
105 pp.
Colo.
Dept.
Game
and
Fish.
In Quart. Prog. Rep., Oct.

RING-NECKED PHEASANT
Aldrich, J. W., and A. J. Duvall. 1955. Distribution of American
gallinaceous game birds. U.S.D. 1., Fish and Wildl. Ser., Circula:r:
No. 34. 23 pp.
Figge, H.
Jan.

1952. Game bird survey. pp. 45-60.
Colo. Dept. Game and Fish. 60 pp.

Figge, H., et al. 1953. Game bird survey.
Rep., Jan. Colo. Dept. Game and Fish.

In Quart. Prog. Rep.,

pp. 75-86.
141 pp.

Sandfort, W. W. 1950. Game bird survey. pp. 17-21.
Rep., July. Colo. Dept. Game and Fish. 80 pp.
Oct.

In Quart. Prog.

In Quart. Prog.

1950. Game bird survey. pp. 1-12. Fed. Aid Proj. W-37-R,
Colo. Dept. Game and Fish. 44 pp.

1951. Game bird survey. pp. 1-7.
Colo. Dept. Game and Fish. 35 pp.

Fed. Aid Proj. W-37-R, Jan.

1951. Game bird survey. pp. 9-20.
Colo. Dept. Game and Fish. 102 pp.

In Quart. Prog. Rep., July.

1952. Game bird survey .. pp. 115-119.
July. Colo. Dept. Game and Fish. 171 pp.

In Quart. Prog., Rep.,

1952. Game bird investigations. pp. 37-41.
Rep., Oct. Colo. Dept. Game and Fish. 105 pp.
1955. Game bird survey. pp. 1-9.
Colo. Dept. Game and Fish. 94 pp.

In Quart. Prog.

In Quart. Prog. Rep., Jan •.

1955. Game bird survey. pp. 49-63,65-69. In Quart. Prog. Rep.,
July - Part I. Colo. Dept. Game and Fish. 115 pp.
1960. Game bird survey. pp. 37-55,59-65.
Colo. Dept. Game and Fish. 161 pp.

In Quart. Rep., April.

�66B

1961. Game bird survey. pp. 97-102. In Quart. Rep., July _
Part II. Colo. Dept. Game and Fish. pp.97-20S.
1963. Game bird survey. pp. 213-223. In Game Res. Rep.,
Oct - Part II. Colo. Dept. Game, Fish and Parks. pp. 109-241.
Sandfort, W. W., and H. M. Swope, 1954. Game bird survey.
In Quart. Prog. Rep.) Jan. Colo. Dept. Game and Fish.
Stiehm, H. A. 1950. Game bird survey. p p . 1-4.
July. Colo. Dept. Game and Fish. SO pp.

pp. 19-29.
113 pp.

In Quart. Prog. Rep.,

Swope, H. M. 1965. Colorado long range game species management plans,
1965-1975, for pheasant. Colo. Dept. Game, Fish and Parks. 54 pp.

SAGE GROUSE
Aldrich, J. W., and A. J. Duvall. 1955. Distribution of American
gallinaceous game birds. U.S.D.I., Fish and Wildl. Ser., Circular
No. 34. 23 pp ,
Anderson, A. E.
Rep., Jan.

1959. Deer-elk investigations. pp. 113-12S.
Colo. Dept. Game and Fish. 12S pp.

In Quart.

1960. Deer-elk investigations, effects of sagebrush eradication
by chemical means on deer and related wildlife. Job Completion Rep. ,
J~ly~ C0lc. Dept. G~~e ~~d Fish. 72 pp.

----

1960. Distribution and abundance indices of selected biota in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Vniva, Ft. Collins. 130 pp.

1966. An investigation of 2,4-D application to sagebrush within a
mountain shrub-conifer complex, Uncompahgre National Forest. In Game
Res. Rep , , July - Part III. Colo. Dept. Game, Fish and Parks. pp. 345420.
1969. 2,4-D, sagebrush and mule deer-cattle use of upper winter
range. Special Rep., No. 21~ July. Colo. Div. Game, Fish and Parks.
21 pp.
Rogers, G. E. 1955.
July - Part II.

Game bird survey. pp. 123~125. "In Quart. Rep.,
Colo. Dept. Game and Fish. pp. 105-lS1.

1963. Game bird survey. pp. 13-32. In Game Res. Rep., Oct. _
Part 1. ~olo. Dept. "Game, Fish and Parks. pp. I-IDS.
1964. Sage grouse investigations in Colorado.
Colo. Game, Fish and Parks. 132 pp.

Tech. Publ., No. 16.

�67B

SHARP-TAILED GROUSE
Aldrich, J. W., and A. J. Duvall. 1955. Distribution of American
gallinaceous game birds. U.S.D.I., Fish and WildIe Ser., Circular
No. 34. 23 pp.
Anderson, A. E.
Rep., Jan.

1959. Deer-elk investigations. pp. 113-128.
Colo. Dept. Game and Fish. 128 pp.

In Quart.

1960. Deer-elk investigations, effects of sagebrush~radication
by chemical means on deer and related Hildlife. Job Completion Rep.,
July. Colo. Dept. Game and Fish. 72 pp.
1960. Distribution and abundance indices of selected biota in
Hestern Colorado before and tHO months after 2,4-D application. M.S.
Thesis. Colo. State Univ., Ft. Collins. 130 pp.
1966. An investigation of 2,4-D application to sagebrush witain
a mountain shrub-conifer complex, Uncompahgre National Forest. In
Game Res. Rep., July - Part III. Colo. Dept. Game, Fish and Parks.
pp , 345-420.
1969. 2,4-D, sagebrush and mule deer-cattle use of upper winter
range. Special Rep., No. 21, July. Colo. Div. Game, Fish and Parks.
21 pp.
Rogers, G. E. 1963.
Oct. - Part II.

Game bird survey. pp. 187-192. In Game Res. Rep.,
Colo. Dept. Game, Fish and Parks. pp.109-24L

1965. Colorado long range game species management plans, 19651975, for blue grouse, sage grouse, sharp-tailed grouse, ptarmigan.
Colo. Dept. Game, Fish and Parks. 16 pp •.
1965. Game bird survey. pp. 209-214. In Game Res. Rep., April Part II. Colo. Dept. Game, Fish and Parks. pp. 175-235.

Nov.

1969. The sharp-tailed grouse in Colorado.
Colo. Div. Game, Fish and Parks. 94 pp.

Tech. Publ., No. 23,

Rogers, G. E., and F. D. Stearns. 1964. Game bird survey. pp. 291-341.
In Game Res. Rep., April - Part III. Colo. Dept. Game, Fish and Parks.
pp. 235-397.

WATERFOHL
Funk, H. D. 1970. Migratory bird investigations. pp. 19-40.
Rep., Oct. Colo. Div. Game, Fish and Parks. 171 pp.

In Game Res.

Grieb, J. R. 1959. Waterfowl surveys and investigations. pp. 145-155. In
Quart. Rep., July - Part II. Colo. Dept. Game and Fish. pp. 145-238.
1960. Waterfmv1 investigations.
Colo. Dept. Game and Fish. 103 pp.

pp , 93-103.

In Quart. Rep ., Oct.

�68B

1961.

Waterfowl investigations.
pp ..9-21.
Colo. Dept. Game and Fish.
103 pp.

Oct.

In Quart. Rcp ,,

1962. Waterfowl surveys and investigations.
pp. 15-25.
Quart. Rep., Oct.
Colo. Dept. Game and Fish. 137 pp.

In
.&lt;

---'.

1965.
Rep , , Oct.

1966.
Rep , , Oct.

1967.
Rep , , Oct.

1968.
Rep , , Oct.

1969.
Rep., Oct.

Migratory bird investigations.
pp. 23-40. In Game Res.
Colo. Dept. Game, Fish and Parks.
199 pp .
Migratory bird investigations.
pp, 17...;.31.
In Game Res.
Colo. Dept. Game, Fish and Parks.
187 pp.
Migratory bird investigations.
pp. 15-30. In Game Res.
Colo. Dept. Game, Fish and Parks.
103 pp.
Migratory bird investigations.
pp. 15-31.
CGlo. Div. Game, Fish and Parks.
135 pp.

In Game Res.

Migratory bird investigations.
pp. 39-56.
Colo. Div. Game, Fish and Parks.
162 pp.

In Game Res.

Grieb, J. R., and G. N. Hunter.
1963. Waterfowl surveys and investigations.
pp. 1-20. In Game Res. Rep., July. Colo. Dept. Game, Fish and Parks.
51 pp.

1964.
Rep., July.

Migratory bird investigations.
pp. 65-78. In Game Res.
Colo. Dept. Game, Fish and Parks.
78 pp.

Hopper, R. M. 1965.
Res. Rep , , Oct.

Migratory bird investigations.
pp. 41-58.
Colo. Dept. Game, Fish and Parks.
199 pp.

In Game

1968. Wetlands of Colorado. Tech. Pub L; , No. 22, March.
Dept. Game, Fish and Parks.
89 pp.
Sandfort, W. W. 1950. Game bird surveys.
pp. 46-51.
April.
Colo. Dept. Game and Fish.
101 pp.

Colo.

In Quart. Prog. Rep.,

WILD TURKEY
Aldrich, J. W., and A. J. Duvall.
1955. Distribution of American
gallinaceous game birds.
U.S.D.I., Fish and Wild~. Ser., Circular
34. 23 pp.
Burget, M. L. 1946.
Fish. 21 pp ,

Colorado wild turkey.

Vol. II.

Colo. Dept. Game and

1948. Wild turkey surveys and investigations.
pp. 47-62.
Prog. Rep., Oct.
Colo. Dept. Game and Fish.
80 pp.
1949.
Rep.

J

Wild turkey surveys and investigations.
Colo. Dept. Game and Fish.
141 pp.

No.

pp. 91-106.

In Quart.

In Quart.Prog.

�69B

\1949. Wild turkey development. pp. 3-lS.ln
Quart. Prog.
Rep.',April. Colo. Dept. Game and Fish. 9S pp0';~.
1951. Wild turkey rehabilitation in Colorado. pp. 7-17. Spec.
Rep. Glenwood Sprgs. Wildl. Conf,, Jan. 16-19. Colo. Dept. Game
and Fish. 85 pp.

Jan.

1957. The wild turkey in Colorado.
68 pp ,

1960. Wild turkey development.
Colo. Dept. Game and Fish. 110 pp.

Colo. Dept. Game and Fish.

pp. 5-:-42.In Quart. Rep., July.
In Quart. Rep.,

1961. Wild turkey
Oct. Colo. Dept. Game
.1963. Wild turkey investigations.'pp. 71-85.
April. Colo. Dept. Game and Fish. 109.pp.

In Game Res. Rep.,

Burget, M. L., and C. Ford. 1951. Wild turkey development. pp. 1-7.
Quart. Prog. Rep., April. Colo. Dept. Game and Fish. 64 pp.

In

Burget, M. L., andD. M. Hoffman. 1950. Wild turkey surveys and investigations.
Three Year Sum. Rep., 1947-1950, July. Colo. Dept. Game and Fish.
4 pp.
1950. Wild turkey investigations.
Colo. Dept. Game and Fish. lS pp.

Fed. Aid Proj. W-39-R, Oct.

"

1951. Wild turkey surveys and investigations. pp. 11-29.
Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish. 117 pp.
1952. \.Jildturkey investigati6ns. pp , 1-5.
April. Colo. Dept. Game and Fish. 62 pp.

In

In Quart. Prog. Rep.,

1952. Wild turkey investigations. pp , 1-9,11-14,21-'24. In Quart.
Prog. Rep., Oct. Colo. Dept. Game and Fish. 105 pp.
1953. Wild turkey investigations. pp. 19-33.
April. Colo. Dept. Game and Fish. 99 pp.
1953. Wild turkey investigations~ pp. 1-6.
July. Colo. Dept. Game and Fish. 129 pp.

In Quart. Prog. Rep. ,

In Quart. Prog. Rep. ,

1954. \-Jildturkey investigations. pp. 13-24.
July. Colo. Dept. Game and Fish. 182 pp.

In Quart. Prog. Rep. ,

1955. Wild turkey investigations.
Rep., Oct. Colo. Dept. Game and Fish.

pp. 1-S,13-19.
108 pp.

1957. Wild turkey investigations.
Colo. Dept. Game and Fish. 116 pp.

pp. 69-86.

In Quart. Prog.

In Quart. Rep., Oct.

�70B

1958. Wild turkey investigations. pp, 1117-169. In Quart. Rep ,,
July - Part II. Colo. Dept. Game and Fish. pp. 105-181.
Evans, R. L. 1963. Wild turkey investigaUons.
pp. 71-85.
Rep , , April. Colo. Dept. Game and Fish. 109 pp.

In Game Res.

Hoffman, D. M. 1965. Colorado long range game species management plans,
1965-1975, for wild turkey. Colo. Dept. Game, Fish and Parks. 11 pp.
Hunter, G. N. 1965. Colo. big game harvest, 1959-1965. Special Mgmt.
Rep. No. 1. Colo. Dept. Game, Fish and Parks. 41 pp.
Myers, G. T. 1963. Game bird survey. pp. 173-178. In Game Res. Rep.,
Oct. - Part II. Colo. Dept. Game, Fish and Parks. pp. 109-241.
1964.
Part III.

Game bird survey. pp. 243-280. In Game Res. Rep., April _
Colo. Dept. Game, Fish and Parks. pp. 235-397.

1965. Evaluation of a combined blue g rouse-wi Ld turkey hunting
season. Outdoor Facts, No. 27. Colo. Dept. Game, Fish and Parks. 1 p.
1965. Reliability of turkey sex and age ratio data based on hunter
report card returns. Outdoor Facts, .No , 29. Colo. Dept. Game, Fish
and Parks. 2 pp.
1965. Game bird survey. pp. 91-174. In Game Res. Rep., April Part I. Colo. Dept. Game, Fish and Parks. pp. 1-174.
1966. Game bird survey. pp. 69-84. In Game Res. Rep., April.
Colo. Dept. Game, Fish and Parks. 192 pp.
1967. Game bird survey. pp. 167-182,205-211. In Game Res. Rep.,
April - Part II. Colo. Dept. Game, Fish and Parks. pp. 107-241.
1968. Game bird survey. pp , 71,99-110.
Colo. Dept. Game, Fish and Parks. 131 pp.
1969. Game bird survey. pp. 107,153.
Colo. Div. Game, Fish and Parks. 181 pp.

In Game Res. Rep , , April.

In Game Res. Rep., April.

1969. The topic is turkey. pp. 30-34. Colo. Outdoors, No.5,
Sept.-Oct. Colo. Div. Game, Fish and Parks. 48 pp.
1970. Game bird survey. pp , 161,165.
Colo. Div. Game, Fish and Parks. 198 pp.

In Game Res. Rep , , April.

Sheats, C. C. 1970. Hy experience w Lth the turkey. pp , 30-31. Colo. Outdoors, No. 30, May-June. Colo. Div. Game, Fish and Parks. 48 pp.

L. C. Carlson
October 1971

�7lB

GENERAL MAMHALIAN REFERENCES
WILDLIFE MANAGEMENT UNIT 62

Anthony, H. E.

1928.

Field book of North American mammals.

Putnam's Sons, New York.

Armstrong, D. M.

1972.

674 pp.

Distribution of mammals in Colorado.

Kansas Printing Service, Lawrence.
Hall, E. R., and K. R. Kelson.
Ronald Press Co.
Lechleitner, R. R.

New York.

1969.

Boulder, Colo.

1959.

The mammals of North America.
1162 pp.

Wild mammals of Colorado.

N. Y.

1955.

List of North American recent
Bulletin ?05.

954 pp.

The mammal guide. Doubleday &amp; Co., Inc. Garden City,

384 pp.

Warren, R. R.
Okla.

Pruett Publ. Co.

254 pp.

mannnals. Smithsonian lnst.,Washington, D. C.

1954.

Univ.

415 pp.

2 Vols.

Miller, G. S., Jr., and R. Kellogg.

Palmer, R. S.

G. P.

1942.

The mammals of Colo.

Univ. of Okla. Press, Norman,

330 pp.

L. C. Carlson
February 1972

�72B

GENERAL AVIAN REFERENCES
WILDLIFE HANAGEMENT UNIT 62

Bailey, A. M., and R. J. Niedrach. 1965.
of Nat. Hist., Den., 2 Vol. 895 pp.

Birds of Colorado.

1967. Pictorial checklist of Colorado birds.
Hist., Den. 168 pp.
Davis, W. A.
61 pp.

1969.

Birds of western Colorado.

Den. Mus.

Den. Mus. of Nat.

Colo. Field Ornithologists.

Kortright, F. A. 1943. The ducks, geese, and swans of North America.
The Amer. Wildl. Institute, Wash. D.C. 476 pp.
Linduska, J. P. 1964. Waterfowl tomorrow.
Wildl., Wash. D.C. 770 pp ,

U.S.D.I. Bur.Sp.

Pearson, T. G. 1940.
N.Y. 289 pp.

Birds of America.

Peterson, R. T. 1941.
Boston. 240 pp.

A field guide to western birds.

Robbins, C. D~, B. Bruun, and H. S. Zim.
Golden Press, N.Y. 340 pp.

Fish and

Garden City Publ. Co., Inc.,

1966.

Houghton Mifflin Co.,

Birds of North America.

L. C. Carlson
October 1971

�73B

CENSUS AREAS AND ROUTES - \.;rILDLIFE
HANAGEMENT UNIT 62

MULE DEER
The pellet group count data, discussed in the foregoing section on
BIG GANE HAMMAL ABUNDANCE, constitute one fairly recent attempt to
establish deer population trend data for Unit 62, however meagre.
More detailed tables and maps concerning locations, dates, and results of this work are available in the files of Federal Aid Project
W-10l-R (B. D. Baker unpublished).

Currently the unit is sampled by four aerial trend areas.

Sex ratio

data, both pre and post hunting, are also gathered throughout the area.

ELK
Incidental to aerial deer pre and post hunt sex ratio counts~ elk are
tallied in Units 61 and 62 (Burdick pers. comm. 1972).
BLACK BEAR
Incidental to aerial deer pre and post hunt sex ratio counts, black
bear are tallied in Units 61 and 62 (Burdick pers. comm. 1972).
PRONGHORN
Annual aerial census of pronghorns in Units 62, 41, and 411 have been
conducted by the Southwest Regional personnel.

For results see that

office.
SAGE GROUSE
Strutting ground counts were made by Rogers (1964) on Sims Nesa and
on Dominguez Creek in 1960.

A brood count route on the latter area was

�14B

established in 1959.

Currently, all sage grouse counting has been

discontinued due to low populations. (Burdick pers. comm. 1972).

BLUE GROUSE
A blue grouse census route was established by Rogers (1963) in 1961
and repeated in 1962 and 1963.
Units 61 and 62.

It lay along the Divide Road between

This route was 62.5 miles long and was divided into

five sections of approximately 12.5 miles each.

The sections varied

as to range type traversed, elevation, and water proximity.

The

purpose was to determine the feasibility of counting birds on specifi~
production routes.

Brood counts were made all three years; "hooting"

counts were attempted in 1962 along parts of the same route.

Currently,

all blue grouse counting has been discontinued (Burdick pers. comm.
1972) .

SHARP-TAILED GROUSE
Counts of sharp tails on dancing grounds were made by Rogers (1969)
in Unit 62 as follows: near the junction of Divide Road and DOT Cow
Camp Road, near T-Bone Springs, Sa'Wffiill
Hesa on old Paradox Road, and
at an unidentified spot on the south end of Uncompahgre Plateau in
Ouray County.

Location maps for two of these sites appear in Rogers

(1969) and lists of all by county.

Currently these counts have been

discontinued (Burdick pers. comm. 1972).

GAMBEL'S QUAIL
During the period 1950-52 Gambel's quail counts were made incidental
to pheasant crowing and brood counts in Unit 62 (Sandfort 1950-1952).

�75B

Routes

covered

probably

varied yearly

as: 1. Delta-Montrose,
Colona.

in length,

but were identified

2. Pea Green Corner-Montrose,

3. Shavano Valley-

No counts have been made since 1952.

CHUKARS
The Escalante

Census Area was established

been censused

annually

long, beginning
River.

since then.

at the confluence

Those portions

densely vegetated

This comprises
of Escalante

of Escalante

for observing

by Sandfort

in 1950 and has

an area eight miles

Creek and the Gunnison

Canyon which were too wide or too

the birds are sometimes

foot with the aid of a bird dog.

Counts usually

covered on

are made at least

three times in August.

The Indian Rock Ranch Census Area was established
(Bartmann

1963).

way, beginning
Gunnison

This comprised

by Sandfort

in 1950

an area about 3.3 miles long, one

at or near the junction

of Dominguez

Creek and the

River.

Maps and descriptions

of these census routes,

may be found in Federal

Aid Reports

together with field forms,

for October

1963 and Apr-il

Tabulated

results

for the 1963 counts are in the April

Southwest

Regional

personnel

have continued

for recents years are available

1964 report.

these counts,

in that office

1964.

and records

as well as in the Denver

office.

RING-NECKED

PHEASANT

Federal Aid Project W-37-R,

the Game Bird Survey,

developed

for a pheasant

in the early 50's.

This included

census record

procedures
counts

�76B

for winter
Roadside

spring

crowing

counts were selected

appropriate
out.

sex ratios,

was prepared

and run annually

field forms and summary

Finally,

a manual

counts, and summer brood counts.

compiling

from 1955-1965

sheets being produced

with

and filled

these data together with instructions,

for each major pheasant

area within management

regions.

For Unit 62 all these counts were established

on a route known as

Delta-Olathe-Montrose

by management

.up-to-date.

and have been continued

Maps of these routes and tabular records

be found in the Southwest

Regional

personnel

of results may

and Denver offices

of the Division.

WILD TURKEY
Myers

(1967 and pers. comm. 1971) reports

stations
1967.

at which winter

Creek

feed stations were located

(approx. S.15, T.15S.,

Corral Ranch on East Escalante
and 3. Spring Creek

Creek

R.100W).

(approx. S.9, T.48N.,

(approx. S.14, T.48N., RlOW).

Records

Regional

and in the Federal Aid Report

The feed stations

cited.

office,

as follows:

2. Pickett

counts are in files of the Southwest

R.16W.),
for these

the Denver

been discontinued

since 1967, except those near Pickett

Escalante

(Burdick pers. comm. 1972).

Canyon

feed

counts of turkeys were made from 1963 through

In Unit 62 the principle

1. Big Dominguez

having established

office,

and counts have
Corral in

WATERFOWL
Wintering

populations

annually

from fixed-wing

Regional

Management

River from Montrose

of ducks and geese in Unit 62 were censused
aircraft

personnel.

in January

1968 through

Areas covered included

to Delta and the Gunnison

1971 by

the Uncompahgre

River from Delta to

�77B

Whitewater.

Results are on file in the Southwest Regional office and

in unpublished records of Federal Aid Project W-88-R, Migratory Bird
Investigations, in Fort Collins.

Literature Cited

Bartmann, R. M.

1963.

Game bird surveys, pp. 61-70.· In Fed. Aid Quart.

Rep., October - Part 1.

Myers, G. T.

1967.

April - Part 2.

Rogers, G. E.

1963.

Game bird surveys.

No. 16.

(processed).

Game bird surveys, pp. 157-168.

In Fed. Aid Quart.

Colo. Dept. Game, Fish and Parks.

Colo. Dept. Game, Fish and Parks.

1950 through 1952.

(processed).

Tech. Pub1.

132 pp.

The sharp-tailed grouse in Colorado.

Colo. Div. Game, Fish and Parks.

Sandfort, W. H.

Tech. Pub1. No.

94 pp.

Game bird surveys.· In Fed. Aid

Quart. Rep., Colo. Dept. Game, Fish and Parks.
year.

In Quart. Rep.,

Sage grouse investigations in Colorado.

1969.
23.

pp. 167-182.

(processed).

Colo. Div. Game, Fish and Parks.

Rep., October - Part 2.

1964.

Colo. Dept. Game, Fish and Parks.

Pages variable by

(processed).

W. T. McKean
February 1972

�78B

HABITAT

Detailed

RESTORATION

information

PROJECTS

on projects

Center,

Unit 62 projects

could be conveniently

day's work.

specific

details

able.

for restoration

Fort Collins.

information

to handling

and I take the attitude

compiled

us that all

and summarized
directly

are needed,

this information

to him.

Another

middleman

If
avail-

that requests

to Mr. Kufeld.

is alterable,

repository

to keep rapidly

by about

that too is readily

be sent dire~tly

that a central

system have been established
and easily available.

UNIT 62

upon what data are needed, we recommend

project

The above approach

Mr. Kufeld informed

Simply make your requests

on only one project

So, depending

MANAGEMENT

are kept in the office of Mr. Ron Kufeld

at the Research

one-half

- WILDLIFE

but Mr. MCKean

and efficient

changeable

habitat

should not be necessary.

B. D. Baker
December 1971

data retrieval
data current

�79B

MANAGEMENT

A.

PROBLEHS

CHECKLIST

- WILDLIFE

MANAGEMENT

UNIT 62

Present.
1.

Deer-auto
Boyd

collision

losses, U.S. Highway

(1961, 1962, and 1964) reports

were killed by auto ~ollisions
roughly

14 miles

recently,

of highway

ten deer deaths per month

on this portion

that a minimum

during

between

of U.S. Highway

herd-drain

problem

dominantly

yearlong

residents.

2.

collisions

similar

problems

are reported

As vehicular

62 and 65
a

are pre--

increases,
designed

to

can be solicited

I 70 and elsewhere.

southeast

portion

winter,

traffic

Assistance

have been common and persistent
of Unit 62, and more
of Unit 65.

from farmers

especially,

Decreased

is alleged because

Damage by deer to stacked

Units

structures

Complaints

and spring grasses

3.

Highway

to farm crops.

portion

to have occurred

The se losses represent

Deer damage

westcentral

Hore

on a group of deer which

are needed here.

on Hi.ghway

period on

Colona and Ridgway.

550, lying between

this problem will also increase.
prevent

of 58 deer

the 1961-1963

(Denton, Cox, Roy pers. comm. 1972).
continuing

550.

in the

in the adjoining

production

of alfalfa

of early grazing by deer.

alfalfa has been fairly common in

as well as to orchards

in this area.

Need for techniques

to provide

population

of selected wildlife

species by area and

topographic

from ridge top to creek

densities

reliable

periodic

indices

to annual

drainage.
The numerous,

abrupt

changes

�bottom present special sampling problems for pellet group
work for deer and elk.

aunt

Techniques for censusing many other wild-

life species are also needed.

4.

Habitat manipulation, federal agencies and private landowners.

Removal of food and cover vegetation over large blocks of land
create serious survival problems for both big game and various
grouse species.

In recent years progress has been made in

cooperative planning and advice-giving on new projects.
B.

Future.
1.

Dallas Creek Project, U. S. Bureau of Reclamation.

Proposed by the U. S. Bureau of Reclamation (1965) is the Dallas
Creek Project which would involve construction of three reservoirs
and several canals.

Principal impoundment would be Ridgway Reser-

voir on the Uncompahgre River and Dallas Creek near Ridgway.
Smaller reservoirs would be the Dallas Divide Reservoir on upper
Pleasant Valley Creek and Sneva Reservoir on a Cow Creek offstream
site.

The project would supply irrigation water for new and

existing agricultural cropland on the Log Hill Mesa area northwest
of Ridgway and supplemental water for other croplands of the
Uncompahgre River Basin between Ridgway and Colona, on both sides
of the river.

Additional benefits would result from improved

domestic rural and municipal and industrial water supplies for the
Uncompahgre Valley from Montrose to Delta.

An estimated 14,690 acres of Log Hill Mesa would be changed from

�8IB

deer and turkey habitat
duction

of range is judged

with an ultimate
hunting

to farmland with the project.
carrying

Also, some 300 man-days

to be lost annually

ments would practically

of deer

eliminate

are

Farm develop-

sage grouse on Sims Mesa,

Sport Fish. and Wildlife

of

of turkey hunting

with the project.

the Log Hill Mesa blue grouse population
(U. S.Bur.

capacity

loss of 600 or more deer and 1,000 man-days

annually.

estimated

to reduce

This re-

and

would be displaced

1966).

Losses of Log Hill Mesa deer habitat

are proposed

in part, by acquisition

Game, Fish and Parks Division

development

and Colorado

of 1,900 acres of land adjoining

Billy Creek Big Game Management
by the project.

would present

losses.

an opportunity

and Wildlife

Division

Area, with all costs to be borne

Establishment

for mitigating

of Ridgway

for developing

area which would be administered
Sport Fisheries

the existing

There are no feasible means

and grouse habitat

to be mitigated,

Reservoir

a waterfowl

and managed

turkey

production

by the Bureau of

(U. S. Bur. Sport Fish. and Wildlife

1966).

2.

PL 566 Watershed

There is one knmm
southwest

Project,

U. S. Department

active small watershed

of Delta.

Updated

project

status of several

jects that have been considered

is not known

but names of them are given as follows:
Creek, Happy Canyon,

of Agriculture.
on Roatcap Wash

other PL 566 pro-

as of this writing,

Horsefly

Creek, Dolores

Spring Creek, Shavano Valley,

Sandy Creek, Sumner Dr aw , and Buttermilk

Dry Creek,

~oJash(U.S .D.A. 1962).

�82B

In view of flood prevention-watershed protection objectives of
these projects, it is assumed that adverse impacts on wildlife
would be minor.

Nevertheless, if such activity as injudicious

brush control is included in practices designed to effect
erosion and f Loodwat.e'r
be reduced.

control, habitat of deer and grouse could

Other wildlife species, as well, could be placed

at a disadvantage depending upon kinds of practices used.

3.

Housing developments in existing natural wildlife habitat.

Literature Cited

Boyd, R. J.
82.

1961.

Study of deer losses on Colorado highways, pp. 75-

In Fed. Aid Quart. Rep., April - Part I.

and Fish.

1962.

96 pp.

(processed).

Study of deer losses on Colorado highways, pp. 93-100.

In Fed. Aid Quart. Rep., October.
pp.

Colo. Dept. G&amp;~e

Colo. Dept. Game and Fish.

137

(processed).

1964.

Study of deer losses on Colorado highways, .pp. 36-41.

In Game Res. Rep., April - Part I.
Parks.

69 pp.

(processed).

U. S. Bureau of Reclamation.
Feasibility Rep.
March.

Colo. Dept. Game, Fish and

1965.

Dallas Creek Project, Colorado.

U. S. Dept. Int., Bur. RecL,

Salt Lake City.

75 pp. + Reps. of Coop. Agencies addenda, 59 pp.

�83B

'U. S. Bureau of Sport Fisheries and Wildlife.
Project, Colorado.

1966.

Dallas Creek

Bur. Sport Fish. and Wildlife Rev. Rep.

U.S.D.l., Fish and Wildl. Ser.

Albuquerque.

July 27.

7 pp. +

Dallas Cr. Proj. Substantiating Rep., 18 pp.

U. S. Department of Agriculture.

1962.

Water and related land

resources, Gunnison River Basin - Colorado.

Coop. Study Rep.

of Colo. Water Conserv. Board and U.S.D.A. Econ. Res. Service,
Forest Service, and Soil Conservation Service, Denver.

November.

103 pp.

W. T. McKean and B. D. Baker
February 1972

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                  <text>July, 1973

-45-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-27

Work Plan No.

llC

Job Title
April

Raymond
William

Investigations
1

Job No.
Experimental

Period Covered:
Personnel:

Deer-Elk

Elk Harvest Regulations

1, 1972 through March

31, 1973

J. Boyd, Walter T. Burkhard, Larry E. Harris,
H. Rutherford and Richard Sherman.

Donald Horak,

ABSTRACT
This annual report summarizes the effect of a 4-point bull restriction in
1972 on an intensively managed elk herd and continued the evaluation of the
impact of specified permits on herd size, sex and age structure, hunter
success and pressure.
Estimated herd size, determined by formulae using
pre and post-hunt sex and age ratios in combination with kill levels,
decreased from about 6,800 in 1966 to 6,300 in 1970 and increased to near
6,400 in 1972 and to approximately 7,400 in 1972. Pre-hunt sex ratios
(bulls per 100 cows) increased from about 29 in 1966 to near 49 in 1972.
Age ratios (calves per 100 cows) increased from 58 in 1966 to 65 in 1972.
The changes in sex ratios were caused by issuing more antlerless and fewer
antlered permits to hunters in an attempt to stabilize the female segment
of this herd.

��-47-

EXPERIMENTAL

ELK HARVEST

REGULATIONS

Raymond J. Boyd

P. S. OBJECTIVE
To determine if modified specified permits fOF elk will enable
gers to achieve a pre-determined kill level on bulls.

game mana-

SEGMENT OBJECTIVES
1.

Continue

review of relevant

2.

Determine accurate sex and age ratios in the elk herd, both pre- and posthunt by classifying at least 1,000 animals in late September and early
December.

3.

Estimate the number of mature bull elk in the 1972 pre-hunt elk population that would have antlers with at least 6 points on one side.

4.

Establish the number of trophy bull permits that could conceivably
remove one-half of the estimated 6 point bull population.

5.

Request the Division of Wildlife Commission to set a trophy bull elk
hunt, with the restriction that a legal bull must have 6 points or more
on at least one antler.

6.

Measure

the kill by age class.

7.

Prepare

annual report.

METHODS

literature

on population

analysis.

AND MATERIALS

The Colorado Division of Wildlife Commission did not accept our recommendations for an experimental 6 point bull elk "trophy" hunt in the White
River for 1972. Instead, they established a regulation for the 1972 elk
season requiring that a legal bull elk had to have at least 4 points on one
antler.
They also stated that this regulation would be in force during the
regular elk season (October 28 through November 12, 1972) and there would be
no special early "trophy" hunt in the study area.
For these reasons, methods
and materials reported below and data analysis in succeeding portions of
this report-will not adhere strictly to the segment objectives listed above.
1.

Check Wildlife Review and other biological and statistical
articles concerning animal population analysis.

reviews

for

�-48-

2.

See Prenzlow (1968).

3.

Using data collected during the 1971 post-hunt classification counts in
Area E, total the number of bulls observed that had 4 points or more on
at least one antler and add to these animals those whose antler conformation would probably develop into a 4 point antler by September, 1972.
Utilize the population projection formula (Prenzlow 1968) to estimate
the total bull population in Area E in September, 1972 and calculate
the number of 4 point or larger bulls that would be available to hunters
during the 1972 elk season.

4.

During the regular elk season in 1972, biologists familiar with the
tooth wear and replacement method of'aging elk were stationed at the
regular Game Management check station at Rifle where they aged all elk
taken from Area E. Aging of these elk took place during the first nine
days of the elk season.

RESULTS AND DISCUSSION
Sex and Age Composition
Aerial Surveys
Pre-hunt Sex and Age Ratios--The 1972 pre-hunt sex and age ratio counts of
the elk population in Area E were made with the aid of a helicopter on
September 25 through September 28, 1972. These classification counts were
confined to elk observed only in Area E (Table 1). A total of 1,103 elk
was classified, resulting in ratios of 64.92 calves per 100 cows, 48.84
bulls (includes all bulls) per 100 cows, 21. 70 branch-antlered bulls (legal
bulls, not branched spikes) and 27.13 yearling bulls per 100 cows.
Summaries of pre-hunt sex and age ratio counts for Area E during the period
1965 through 1972 are shown in Table 2.

Post-hunt Sex and Age Ratios--Post-hunt sex and age ratio classifications of
elk in Area E were flown on December 18 and 19, 1972. A total of 2,290 elk
were classified giving ratios as follows: 68.68 calves per 100 cows, 23.12
bulls (includes all bulls) per 100 cows, 3.77 mature (branch-antlered) bulls
per 100 cows, and 19.35 yearling bulls per 100 cows (Table 3). Summaries of
post-hunt classifications for this area from 1965 through 1972 are shown in
Table 4.

Ground Surveys
During the years 1967 through 1972 a ground classification of elk in Area E
was made each July. This was done to estimate yearling-adult cow ratios,
which cannot be accurately obtained during early fall aerial classification
counts. These data can also serve as independent estimates of sex and age
structure. It becomes very important to determine the proportion of yearling
females in the herd, and thus their influence on overall herd productivity.
Usually only a small fraction of the yearling females breed, however there
is considerable conflicting literature on this particular subject.

�-49-

Table l.

Pre-hunt

Mature
Bulls

Young
Bulls

Spikes

Cows

Calves

Total

1

3

8

20

11

43

Nine-Mile

3

1

21

13

8

46

Sleepy Cat Area

1

9

28

59

36

133

Lost Park Area

2

4

10

33

11

60

Oak Ridge

4

3

8

Cattle Creek

27

20

67

Big Ridge

3

3

7

Trappers

Creek
Peak Area

sex and age ratio classifications

1
4

4

12

1

in Area E, 1972.

Location

to Yellowjacket

1

1

1

10

9

22

Shingle

15

7

10

104

82

218

South Fork to Campground,
north side

6

5

2

38

26

77

Park Creek

2

2

5

11

8

28

Bloomfield

1

13

9

26

Lost Solar Creek

1

2

1

4

Ute Creek

3

Bench

10

2

4

43

24

83

Marvine

2

4

3

34

22

65

East Marvine

5

2

1

8

Burro Mountain

Creek
Creek

2

1

4

10

6

23

S. Side of South Fork

1

2

5

23

12

43

Patterson

3

2

24

15

44

Skinny fish Creek

5

2

20

43

28

98

Anderson

61

51

140

516

335

1,103

Total

5.53

4.63

12.69

46.78

30.37

100.0

Percent

Creek

Reservoir

�T~Q1e _2.

Year

1965
1966
1967
1968

Bulls Counted
Mature Young Spikes
23
17
17

49

161

15

-54

29

17

1969
1970

Summary of pre-hunt aerial sex and age ratio counts in Area E, 1965-1972.

37
113

34

88
81
211

34

1971

46

30

1972

61

51

82
93
140

Ratios Per 100 Cows
Branch-Antlered
Spike
Bulls
Bulls

Cows
Counted

Calves
Counted

Total
Count

Mature
Bulls

Young
Bulls

699

416

1,348

3.3

7.0

10.3

23.0

59.5

299

172

557

5.7

5.0

10.7

18.1

57.5

415

247

796

4.1

7.0

11.1

21.2

59.5

366

221

722

4.6

10.1

14.7

22.1

60.4

645

389

1,358

17.5

32.7

60.3

455

249

854

15.0

18.0

54.7

17.5
7.5

7.5

Calves

541

337

1,047

8.5

5.5

14.0

17.2

62.3

516

335

1,103

11.8

9.9

21.7

27.1

64.9

I
IJ1

0
I

�-51-

Table 3.

Post-hunt sex and age ratio classifications in Area E, 1972.

Mature
Bulls

Young
Bulls

Spikes

Cows

Calves

Total

6

78

46

130

Hiway 13 to Ye1lowjacket

Location

4

4

47

197

113

365

Coal Creek

1

6

43

240

169

459

Oak Ridge, N. Side

17

8

28

Little Beaver

3
2

5

67

236

170

480

Big Beaver

2

1

6

59

45

113

Oak Ridge, S. Side

1

14

93

74

182

Buford to Fawn Creek

1

0

8

27

23

59

Fawn Creek

3

6

3

34

25

71

Fawn Creek to Lost
Creek

5

16

13

34

Marvine Creek

8

27

28

63

Crooks Park Area

1

7

64

44

116

Burro Mountain

2

1

11

59

37

110

South Fork, Park
Creek to Campground

1

1

6

47

25

80

Miller Creek

19

26

231

1,194

820

2,290

Total

0.83

1.14

10.09

52.14

35.80

100.00

Percent

Observers on horseback, during the early morning hours, classified elk in
different sections of the Flat Tops Primitive Area. In three days teams
of observers classified a total of 1,111 elk (Table 5). Many more elk
were observed but classifications could not be completed, so they were not
included in the totals.

�Table 4.

Summary of post-hunt aerial sex and age ratio counts in Area E, 1965-1972.

Year

1965
1966
1967
1968

Bulls Counted
Mature Young Spikes
13
7

7

18
14

1969
1970

13

1971

10

1972

19

Calves
Counted

Total
Count

Mature
Bulls

Young
Bulls

59

930

613

1,628

1.4

1.4

2.8

6.3

65.9 !/

114

1,245

739

2,112

0.6

0.6

1.2

9.2

Y
61.1 Y

30

166

1,318

806

2,338

1.4

2.3

3.7

12.6

22

112

951

699

1,798

1.5

2.3

3.8

11.8

1,012

661

1,839

2.9

13.5

29
35

Ratios Per 100 Cows
Branch-Antlered
Spike
Bulls
Bulls

Cows
Counted

137

2.9

Calves

59.4

Y
65.3 Y
73.5

18

67

1,402

884

2,406

2.5

1.3

3.8

4.8

63.0 '£/

7

135

1,050

734

1,936

1.0

0.7

1.7

12.8

69.9 2/

26

231

1,194

820

2,290

1.6

2.2

3.8

19.3

68.7 ~/

!/ Wide open bull hunting, no restriction

on numbers of bull hunters, but only 650 ant1er1ess only permits.

~/ Specified permit hunting, numbers of bull hunters restricted each year; 1966, 1967 and 1968 had 2,500 bull
only permits; 1969 had 2,200 bull only permits, and 1970 had 3,000 bull only permits. Ant1erless elk hunting
was also restricted with 1,000 permits in 1966; 1,500 in both 1967 and 1968; 1,000 in 1969, and 800 in 1970.

1/ Only 2,200 branch antlered bull permits

(spike bulls were not legal) with 600 ant1erless permits.

~/ Only 2,500,4 point bull permits (only bulls with 4 points or more on at least one antler were legal)
and 800 antlerless only permits.

I

VI
N
I

�-53-

Table 5. Composition
in Area E, 1972.

of 1,111 elk c.l as sLf Led du rLng .lu ly

gl'lltllld

l'llllllt~,

Mature
Bulls

Yearling
Bulls

Mature
Cows

Yearling
Cows

Calves

Total

71

77

486

129

348

1,111

6.39

6.93

43.74

11.61

31.33

100

25

27

43

31

33

47

Me an G roup CompoS1t10n"
2/

1.51

1.64

10.34

2.74

7.19

23.42

1.1 Frequency

(i.e., mature bulls were observed

Item

Number Observed
Percent

Observed

Frequency

. G roups-1/
1n

in groups means
total groups).

~/ Mean group composition
(average group size).

determined

by percent

of elk observed

in 25 of 47
times 23.42

Check Station Surveys
Number of Elk Checked--Check station data from elk harvested in Area E during
the 1972 elk hunt were gathered only at the regular management check station
at Rifle, Colorado.
A total of 300 elk were checked during the first nine
days of the hunt. Table 6 lists the daily elk check out from the two Area E
units through the Rifle station in 1972.
Sex and Age Composition of Elk Checked--The percent composition (bulls, cows
and calves) of elk checked in 1972 from Area E is shown in Table 7. Sex and
age ratios of the 300 elk from the study area were 47.67 percent bulls, 43.00
percent cows and 9.33 percent calves.
This compares to 52.43 percent bulls,
39.32 percent cows and 8.25 percent calves checked out in 1971.
Age Structure of the Harvest--Of the 300 elk checked in 1972 from the study
area, 128 were aged using techniques described by Quimby and Gaab (1957).
Table 8 lists these ages, by sex, while Table 9 details similar data by game
management unit.
One incisor tooth was collected from each elk that was field aged to be older
than 2 1/3 years.
These teeth were taken to the Research Laboratory in Fort
Collins where they were aged according to the dental cementum technique
reported by Keiss (1969). Assuming that ages determined by dental cementum
were correct, comparisons of field ages and dental cementum revealed an error
of 37.50 percent in assigning ages to 3-1/3 year old elk, while the error
associated with aging elk older than 3-1/3 years was 83.33 percent (Table 10).

�-54Table 6.

Elk check out by day of season, Area E, 1972.

Unit and Sex

Da~ of Season
5
6
7

1

2

3

4

0

8

1

13

7

8

1

1

4

9

4

1

9

5

22

0

3

10

1

1

1

Males

8

9

Total

9

19

3

68

16

7

7

1

50

11

24

16

26

4

118

3

13

17

16

10

11

83

9

4

17

20

21

15

11

99

4

19

7

30

37

37

25

22

182

0

11

11

16

20

25

25

29

15

152

Females

2

2

13

13

21

36

28

22

12

149

Subtotal

2

13

24

29

41

61

53

51

27

301

Percent of
Total Check

0.7

4.3

8.0

9.6

13.6 20.3 17.6 16.9 9.0

Unit 23
Males
Females

Subtotal
Unit 24
Males
Females
Subtotal
Area E

100.00

Table 7. Composition
of elk checked through the Rifle check station from
Area E, 1972.

Bulls

Percent

Cows

Percent

Calves

Percent

Total

Unit 23

67

56.78

41

34.75

10

8.47

118

Unit 24

76

41.76

88

48.35

18

9.89

182

Total

143

47.67

129

43.00

28

9.33

300

�Table 8.

Sex and age determinations of 128 elk checked from Area E, 1972

Sex

Age (Years) !/
4-1/3
5-1/3

1/3

1-1/3

2-1/3

3-1/3

Number

8

7

48

7

1

Percent

10.96

9.59

65.75

9.59

Percent Total

6.25

5.47

37.50

5.47

6-1/3

7-1/3

8-1/3

2

0

0

0

0

73

1. 37

2.74

0

0

0

0

100.00

0.78

1.56

0

0

0

0

57.03

9+

Total

Males

Females
Number

20

9

10

9

1

4

1

0

0

1

55

Percent

36.37

16.36

18.18

16.36

1. 82

7.27

1.82

0

0

1. 82

100.00

Percent Total

15.63

7.03

7.82

7.03

0.78

3.12

0.78

0

0

0.78

42.97

Number

28

16

58

16

2

6

1

0

0

1

128

Percent Total

21. 88

12.50

45.31

12.50

1.56

4.69

0.78

0

0

0.78

100.00

Total Elk

l/ Ages assigned by field techniques described by Quimby and Gaab (1957).

I
\.II
\.II
I

�Table 9.

Sex and age determinations of 128 elk aged from Area E, by unit of kill, 1972.

Unit and Sex

1/3

1-1/3

2-1/3

Age (Years) 1/
3-1/3
4-1/3
5-1/3

6-1/3

7-1/3

8-1/3

5

1

1

0

0

0

0

39

12.82

2.56

2.56

0

0

0

0

100.00

6.85

1. 37

1.37

0

0

0

0

53.42

2

0

1

0

0

0

0

34

9+

Total

Unit 23 - Males
Number

1

Percent of Total
Percent of All Males

4

2.56

10.26

1.37

5.48

27
69.23
36.99

Uni t 24 - Males
Number
Percent of Total
Pe rcent of All Males

7

3

20.59

8.82

9.59

4.11

21
61. 76
28.77

5.88

0

2.94

0

0

0

0

100.00

2.74

0

1. 37

0

,0

0

0

46.58

2

1

3

0

0

0

0

24

8.33

4.17

12.50

0

0

0

0

100.00

3.64

1.82

5.46

0

0

0

0

43.64

7"

0

1

1

0

0

1

31

22.58

0

3.23

3.23

0

0

3.23

100.00

12.73

0

1. 82

1.82

0

0

1.82

56.36

I

Unit 23 - Females
Number
Percent of Total
Percent of All Females

9
37.50

5

4

20.83

16.36

9.09

16.67
7.27

Unit 24 - Females
Number
Percent of Total

11

35.48

Percent of All Females 20.00

4

6

12.90
7.27

!/ Ages assigned by field techniques

19.36
10.91

described by Quimby and Gaab (1957).

I

V1
0-,

�-57-

Table 10. Percent error between dental cementum and tooth replacement
and wear aging techniques, by age class, of 20 elk from Area E, 1972.

Dental
Cementum

Age
Class

1/

Field
Age '!:./

Percent
Correct

Percent
Error

Calf
1
2

3

8

5

62.50

37.50

4

6

1

16.67

83.33

5

2

1

50.00

50.00

6

3

o

o

100.00

7

1

o

o

100.00

8

o

o

9+

o

o

Total

20

7

35.00

65.00

!/ Number

of teeth checked in the laboratory

using the dental

cementum

procedure.

'!:.../ Nurilberof ages correctly
and replacement

assigned

at the check station

using tooth wear

procedures.

1/ Calves, yearlings

and 2 year olds were not aged by the dental

cementum

procedure.

Table 11 indicates the distribution of error by age class.
Overall error
was 65.00 percent and this compares with 64.31 percent in 1971.
Of 25
check station assigned ages from elk older than 2-1/3 years, 20 were com·parative1y aged by the dental cementum technique.
Of these 20 animals aged
by the two methods, 12 (60.00 percent) were field aged within one year on
either side of the actual age as determined by dental cementum procedures.

�-58-

Table 11. Distribution of assigned
ages in relation to ages determined by
dental cementum procedures, Area E, 1972.
Cementum
Age Class

3
4

Check Station Assigged Ages

2

3

1

8

3

4

6

1

4

5
6

1

7

5

6

7

8

9+

1

Total

13
11

2

1

3

2

3

6

1

1

8

2
0

0

9+

Total

2

13

9

5

5

1

0

0

0

0

35

Mortality Estimates
Harvest Estimates
Random Survey--Because of annually decreasing hunter report card returns, no
report cards were attached to the 1972 elk licenses. A random survey was
conducted on 31 percent of the elk license buyers to determine the 1972
elk harvest estimate. While elk hunters in the study area were surveyed
along with all other elk hunters, an extra large sample (1,032 hunters out
of 3,300 permittees, a 31.27 percent sample) was drawn from this hunter
_
population to obtain a more accurate estimate of the elk harvest. The 31.37
percent sample drawn in 1972 compares with a 50.00 percent sample drawn from
hunters in this area since 1968. Of the 1,032 hunters sampled, 926 (89.73
percent) returned usable cards. The estimated total elk kill in Area E for
1972 was 976 elk (500 bulls, 413 cows and 63 calves), with 34.01 percent of
the 2,870 active hunters successful. Table 12 gives results of the 1972
random survey, Lnc.Luddrrg kill by type of license, while Table 13 lists
active (those who actually hunted elk in 1972) hunters and success ratios.
Projections from the random suryey indicated that 430 licenses were issued
that were not used (13.03 percent). The number of no hunts in Area E have
averaged 7.0 percent since 1966.

�-59-

Table 12. Results of the 1972 random survey of Area E elk hunt permit
holders. II

Item
Permits Issued

Antlered
NonResident Resident

Antlerless
NonResident Resident

Total

1,773

727

637

163

3,300

Permits in Sample

452

180

323

77

1,032

Percent in Sample

25.49

24.76

50.71

47.24

31.27

Sample Returned

416

161

282

67

926

Percent Returned

92.04

89.44

87.31

87.01

89.73

Percent of Total Permits

23.46

22.15

44.27

41.10

28.06

Reported'"No Hunts"

69

17

23

3

112

Estimated "No Hunts"

294

77

52

7

430

Reported Kills

75

40

159

48

322

Estimated Total Kill

320

180

359

117

976

!I These data will not agree with published figures from the Game Management
Section, as these were compiled from a hand count of the returned survey
cards. Actual count and projected figures are slightly different when the
hand count and the computer run are compared.

Table 13. Number of hunters participating during the 1972 Area E elk hunt
and success ratios per license and active hunters. Projected from random
survey data.

Item

Antlered
Resident Non-Resident

Antlerless
Resident Non-Resident

Active Hunters

1,479

650

585

156

Success (Kill) per License

18.05

24.76

56.36

71.78

Success (Kill) per Hunter

21.64

27.69

61.39

75.00

�--60-

Harvest in Relation to Area and Time
Location of Kill--Kill by Game Management Unit was projected from the random
survey data and appears in Table 14. These data indicate that 65.37 percent
of the 1972 elk kill in Area E occurred in Unit 23. This is a radical
departure from past harvest statistics. The usual pattern is for 60 to 65
percent of the elk harvest to take place in Unit 24. However, because of
the deep snow that occurred in Area E in 1972 many of the elk had moved to
lower ranges in Unit 23 and this was the probable cause of the increased
kill in this unit.

Table 14. Composition
of the 1972 estimated elk kill in Area E by Game
Management Unit.

Unit

Number

Bulls
Percent

Number

Cows
Percent

Unit 23

321

50.31

285

44.67

32

5.02

638

65.37

52.96

128

37.87

31

9.17

338

34.63

51.23

413

42.32

63

6.45

976

100.00

Unit 24

179

Total

500

Calves
Number Percent

Total
Number Percent

Date of Kill--Dates of kill by hunters in Area E are shown in Table 15.
Approximately 81 percent of the kill occurred during the first five days of
the season.

Table 15.
E, 1972.

Numbers of elk killed, by

1
Unit 23
Unit 24
Total
Percent

22
49
71
24.2

2
27
21

48
16.4

day of season, check station data, Area

3

4

Da~ of Season
5
6

7

8

9

Total

15

13

22

7

8

3

0

117

16

30

22

14

14

4

6

176

31

43

44

21

22

7

6

293

10.6

14.7

15.0

7.2

7.5

2.4

2.0

�-61-

Estimates

of MOrtality

Other Than Legal Kill

Wounding Losses--Annual estimates of wounding losses have been made in Area
E since 1966.
In 1971 and 1972 the random questionnaire was designed and
sent out by the Game Management Section in Denver, and questions concerning
wounding of elk and retrieval of wounded elk were not included on the
questionnaire,
thus no comparable data for 1971 and 1972 are available.

Population
Total Population

Estimates

and Projections

Estimates

Formulas were used to estimate the total number of elk present in Area E
prior to the 1972 elk hunt.
Detailed descriptions of the method are presented in Prenzlow (1968) with examples of how ratios of antlered to antlerless
elk from kill, pre- and post-hunt classifications counts were used in a
formula devised by Dr. David Bowden of Colorado State University to estimate
a pre-hunt elk population.
Pre-Hunt Population Estimate 1972--An estimated 3,729 elk were present in
Area E just prior to the opening of the 1972 elk season (Table 16). This
compares to 6,751, 6,313, 6,216, 5,829, 6,347, and 3,358 estimated in 1966,
1967, 1968, 1969, 1970 and 1971, respectively.

Table 16. Pre-hunt population
age in Area E, 1972.

Item

Estimated
Number

estimate

and proportions

of elk by sex and

Expected
Percent 1./

Observed
Percent 1./

Bulls

852

20.53

22.85

Cows

1,744

49.83

46.78

Calves

1,133

29.64

30.37

Total

3,729

100.00

100.00

1/ Percentages
1./ Percentages

expected

from 1971 post-hunt

actually observed
cation counts from a helicopter.

when making

population

data.

the 1972 pre-hunt

elk classifi-

�-62-

Until 1969, and again in 1971 and 1972 total elk populatIon dntn Wl'.'('
believed to be reasonable estimates of actual elk populat Lou Le ve Ls 1n Are a
E. For instance, predictions for the percentages of bulls, cows and calves
made after the post-hunt counts were within five percent of what was actually
observed during pre-hunt counts the next year during all years of the study
(1961-1972), with the exception of 1969, 1970 and 1972. In 1969, the
expected and observed percentages of bulls differed almost 20 percent.
This
was a critical difference, because the change in ratios of bulls from preto post-hunt is a major component of the population estimate formula used to
estimate the pre-hunt elk population.
However, as Prenzlow (1970:280).
explained, early severe September snow storms in 1969 probably caused an
abnormal movement of the elk herd and samples that were counted in 1969 were
not taken from the "normal" herd usually found in Area E.
In 1970, the expected and observed percentages of bulls differed again by
almost 20 percent.
Counting conditions during these flights were very
difficult because of very bad weather conditions and elk that acted very
spooky and were not in areas that elk are normally found during pre-hunt
counts.
In 1972, the observed and expected ratios of bulls differed by 15
percent, caused in all probability by flying at a later date than normal,
and the fact that most of the aspens had not lost their leaves, nor had the
oakbrush lost their leaves.
The flying of inexperienced helicopter pilots
also left much to be desired, as they flew too high and too fast in spite
of repeated requests to do a better job.
The 1972 pre-hunt population estimate of 3,729 appears to be extremely low
for three reasons: (1) 4,440 elk were actually counted in Area E during the
1972 post-hunt trend count, which exceeds the total pre-hunt population
estimate by 711 animals, add to this an additional 976 harvested animals and
there had to be a minimum pre-hunt population of 5,416 elk; (2) the 3,729
elk estimated to be in Area E does not fit the population trend in recent
years, and is less than one-half the number of elk that should have been in
this area based on the 1971 post-hunt population estimate for this herd; and
(3) the bull kill estimate is low due to an extremely high illegal bull kill
that went unreported in 1972 because of a 4 point bull restriction placed
on hunters in this area.
Estimates of the illegal kill of bulls not
retrieved in Area E in 1972 were 447 animals, which if added to the estimated
legal kill, would increase the total pre-hunt population estimate some 3,637
animals.
This unrecorded loss of illegally killed bulls would have a great
depressing effect upon the formula population estimate.
A modified pre-hunt population estimate for Area E can be calculated by
adding in the estimated illegal kill to increase the harvest figures and
recalculating the pre-hunt population by formula (Table 17).
Therefore, to accomplish the objective of testing an elk harvest formula, it
becomes necessary to use the best estimate of the pre-hunt population rather
than the population estimate of 3,729 derived by using legal harvest and
estimates of wounding loss in combination with changes from pre- to posthunt sex ratios.
Recalculating the pre-hunt population using the total herd
drain figure in Table 17 gives an estimated pre-hunt elk population in Area
E in 1972 of 7,366, which appears to fit the herd population trend better
than the 3,729 figure.

�-63-

Table 17. Modified elk harvest data, Area E ,1972,
illegal kill.

Item

Estimated
Legal Kill 1/

including estimated

Wounding
Loss '!:.../

Abandoned
Elk 1/

Total Herd
Drain

Bulls

500

34

447

981

Cows

413

15

64

492

Calves

63

3

Total

976

52

66
511

1,539

!/ From random survey data in Table 12.
Reported by hunters at check stations, wounded elk minus wounded elk
retrieved.

'!:.../

1/ Illegal kill of spikes, 2 points and 3 point bulls.

Post-Hunt Population Estimates, 1972--An estimated 5,827 elk survived the
1972 hunting season (Table 18). This was calculated by subtracting the
total herd drain in Table 17, by class of animal, from the pre-hunt population estimate of 7,366.

Table 18. Post-hunt population estimate and proportions of elk by sex and
age in Area E, 1972.

Post-Hunt POEulation
Expected
Observed
Percent
Percent 1/

Item

Pre-Hunt
Population

Harvest !/

Bulls

1,683

981

702

12.05

12.06

Cows

3,446

492

2,954

50.71

52.14.

Calves

2,237

66

2,171

37.25

35.80

Total

7,366

1,539

5,827

Number

u

1/ From Table 17.

2/ Proportions, by class of animal, of the total estimated number of elk
(5,827) that remained following the 1972 elk hunt. '
1/ Proportions actually observed when making the 1972 post-hunt classifications
by helicopter.

�-64-

During the 1966-1970 period, the management objective for the Area E elk
herd was to hold the post-hunt herd to 5,000 animals. In 1971 and 1972
the herd objective was changed to one of increasing elk with no numerical
limit stated. Table 19 details the trend in post-hunt population estimates
for this elk herd. The reduction in elk numbers during the early years of
the study was primarily the result of issuing more antlerless permits to
narrow the sex ratio of the wintering herd. Data in Table 19 indicates
that management of this herd to a 5,000 animals post-hunt population was
achieved rather well, and the desired herd increase for 1971 was also
achieved.

Table 19.

Post-hunt elk population figures, 1966-1972, Area E.

Year

1966
1967
1968
1969
1970
1971
1972

Post-Hunt Population
Estimate

5,336
4,945
4,506
4,646
4,987
5,744
5,827

Population Projections
Each year based on the previous years post-hunt population estimate, a
projection is made for the next year. This procedure is described in
Prenzlow (1968) in detail.
Population Projection for Area E, 1973--Immediately prior to the opening of
the 1973 elk hunt in Area E, a projected elk popUlation of 8,249 animals will
be available to hunters in this area (Table 20). This figure may change
somewhat, however, if productivity differs significantly from 60 calves per
100 cows.
Consequences of the 1972 Area E Elk Hunt~-In 1971, management of the Area E
elk herd reverted to Regional control, and the Research Section made no more

�-65-

season recommendations. The Wildlife Commission instigated a spike bull
protection regulation for this ~rea in 1971, and a 4 point bull restriction
regulation in 1972. In order to maintain data continuity so evaluation of
the effects of these two special regulations on the bull structure of the
herd could be determined, the Research Section continued to make pre- and
post-hunt classification counts, and to sex and age elk at check stations.

Table 20.

Predicted 1973 pre-hunt elk population, A:J;eaE.

Item

1972 Post-Hunt
Population

Bulls

702

Cows

2,954

Calves
Total

.1973 Pre-Hunt POEulation
Expected
Number
Percent 2/
1,787 J:..I

21.66

4,039

11

48.96

2,171 1/

2,423

!il

29.38

5,827

8,249

!/Assuming that of 2,171 calves, 50 percent (1,085) are males and 50 percent
are females.
J:..ICalculated
by adding the remainin~ bulls in 1972 (702) and 1,085 male
calves from 1972.
11Calculated by adding the remaining cows in 1972 (2,954) and 1,085 female
calves from 1972.
!iICalculated by mUltiplying the number of cows in 1973 (4,039) by the
expected productivity of 60 percent.
ilExpected proportions of total projected numbers of elk (8,249) prior to
the 1973 elk hunt.

When check station data from the 1972 elk hunt were examined, it became
evident that a hunt regulation designed to increase "trophy" antlered bulls
by protecting yearling bulls (by making only bulls with 4 point antlers or
larger legal) placed all bull hunting pressure on the mature bull segment
of the herd and actually reduced chances of bulls living long enough to
develop "trophy" class antlers.
Fig. 1 indicates what happened in 1972 compared with the 5 year period 19661970. By making only bulls with 4 point or larger antlers legal, the 2 year
old bull kill jumped from about 15 to 65 percent of the total bull kill.

�t

AGE

AGE

1
a T

9+

7
6

I

I
55 60

I

I

~~

e

1

I--

7

6

5
4
3
2

I

I

c
I

50

40
%

1966 -1970

30

I

I

20 15 10 5

MALES

DATA

-

I

'"'"

I--

1
J
I

I

I 5 J
14 I
r

I

I

I

5

/0

15 20

%

FEMALES

BASED ON 3,148

n

I

ELK

I

65 60

I

0/0

1972

I

I

I

I

I

I

20 /5

/0

5

5

/0 15 20

MALES

DATA

I

I

I

40

I

c

r
I

3
2

0/0

-

BASED

ON

r28

Fig. 1. Comparison of age pyramids of Area E elk, 1966-1970 and 1972, showing effects of the 4-point bull
restriction on the bull age structure.

I

FEMALES

ELK

I

I

30

�-67-

About 10 percent of the harvest was composed of legal yearling bulls and the
kill of three-year old bulls was nearly tripled.
No bulls older than 5
years of age were taken.
Contrast this with the situation in the left age
pyramid in Fig. 1 where it is evident that older age classes of bulls began
to show up in the harvest after only 5 years of limited bull hunter pressure
caused by the specified permit regulation.
By making all bulls legal, with
reduced hunting pressure, illegal kill of bulls was virtually nonexistent.
When the above data were presented to the Colorado Wildlife Commission in
March, 1973, they voted not to continue the 4 point restriction regulation
and to return to a bull hunt regulation with all age classes of bulls
being legal.
A second draft of the final report on the effect of specified
permits on the sex and age structure of the White River elk herd has been
completed and is now in the hands of the co-author for his review. When
this review is completed, the manuscript will be submitted to the Journal of
Wildlife Management for consideration for publication in the new Bulletin
of the Wildlife Society.

LITERATURE

CITED

Keiss, R. E. 1969.
Comparison of eruption-wear patterns and cementum
annuli as age criteria in elk. J. Wi1d1. Mgmt. 33(1) :175-180.
Prenzlow, E. J. 1968. White River elk population
Job Progress Report. W-38-R-22.
p. 383-421.
1970. White River elk population
Report.
W-38-R-24.
p.257-286.
Quimby, D. C., and J. E. Gaab. 1957.
indicator in Rocky Mountain elk.

components.

components.

Colorado

Colorado

Job Progress

Mandibular dentition as an age
J. Wi1d1. Mgmt. 21(4) :435-451.

��July, 1973

-69-

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~---------

Project No.

W-38-R-27

Work plan No.

14

Job Title

Job No.

1

Middle Park Deer Study - Population Distribution

Period Covered:
Personnel:

Deer-Elk Investigations

April 1, 1972 through March 31, 1973.

R. Bruce Gill and Laren A. Roper.

ABSTRACT
Field work has been completed and a Division Special Report is in preparation
that will be published during the next project segment.

��-71-

MIDDLE PARK DEER STUDY - POPULATION DISTRIBUTION
LarenA.

Roper

P. S. OBJECTIVE
To delineate deer concentration areas in Middle Park and relate changes
in deer distribution in time and space to accumulation and physical properties of snow.

SEGMENT OBJECTIVES
1.

Define deer population sub-unit boundaries.

2.

Compile data and prepare report.

METHODS AND MATERIALS
See Gill (1969).

RESULTS AND DISCUSSION
Field work has been completed and a Division Special Report is in preparation that will be published during the next project segment.
LITERATURE CITED
Gill, R. Bruce. 1969. Middle Park deer study - population distribution.
Colo. Dept. of Game, Fish and Parks, Game Research Report, July,
Part 1. pp. 79-103.

Researcher

��-73-

July, 1973

JOB FINAL REPORT

State of

COLORADO
--------~~~~~---------

Project No.

W-38-R-27

Work Plan No.

14

Job Title:

Job No.

2

Middle Park Cooperative Deer Study - Population Density and Structure

Period Covered:
Personnel:

Deer-Elk Investigations

January 1, 1973 through March 31, 1973

R. Bruce Gill.

ABSTRACT

Several publications have already emanated from this job. Remaining data
are presented in a manuscript currently in preparation for the Journal of
Wildlife Management. Management recommendations will be published in a
Division Special Report.
.

��-75-

MIDDLE PARK COOPERATIVE DEER
STUDY - POPULATION DENSITY AND STRUCTURE
R. Bruce Gill

•

P~ S. OBJECTIVE
To estimate the density and sex and age structure of the Middle Park deer
population in order to harvest this population more efficiently.
SEGMENT OBJECTIVES
1.

Publish the results of this job.

2.

Prepare management recommendations.
METHODS AND MATERIALS

Methods and materials have been detailed previously by Gill (1969 and 1971).
RESULTS AND DISCUSSION
A manuscript titled, "Estimating total numbers of a migratory mule deer
population" is in preparation. This will be submitted to the Journal of
Wildlife Management approximately January 15, 1974.
Recommendations for management are being placed in a Division Special Report
which will be submitted to the printer by March 1, 1974.
Several publications have already emanated from this job, in addition to the
annual Game Research Reports. These are listed in the Bibliography section.
BIBLIOGRAPHY
Gill, R. B. 1969. Middle Park deer study - population density and structure.
Colo. Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. W-38-R-23.
Game Res. Rep., July, Part 1. pp. 105-122.
1969. A quadrat count system for estimating game populations.
Div. of Wildl. Game Info. Leaflet No. 36. 2 pp.
1969.

An apology for a deer season.

____~, o. C. Wallmo, and J. G. Nagy.
Colo. Outdoors.

19(5):14-15.

1970.

Colo. Outdoors.

Colo.

18(6):30-33.

The Middle Park deer study.

�-76-

Gill, R. B. 1971. Middle Park deer study - population density and
structure. Colo. Div. Game, Fish and Parks. Game Res. Div.
Fed. Aid Proj. W-38-R-2S. Game Res. Rept., July, Part 2. pp.
169-188.

----)~:~')-

9!

Prepared by __ \~:'
.•...
,_§...;;;;z ••.·-''J.~....•
L~(:~C'.•.••
&lt;'--'7~~:::::i/.•••.
;;;.;(_
...--f...-=
_
R. Bruce Gill
Wildlife Researcher

�July, 1973

-77-

JOB FINAL REPORT

State of

COLORADO

---------------------------

Project No.

W-38-R-27

Work Plan No.

14

Job Title:

Middle Park Deer Study - Productivity and Mortality

Period Covered:
Personnel:

Deer-Elk Investigations
Job No.

3

April 1, 1972 through March 31, 1973.

R. Bruce Gill, Laren A. Roper, Paul F. Gilbert, Dan L. Baker, Lonnie
M. Brown, Jan L. Wassink ,and Len H. Carpenter.

ABSTRACT

All field data obtained during this segment were reported by Gill, 1972. Publications emanating from this study are listed in the bibliography. A Division
Special Report which will tie Work Plan 14, Jobs 1, 2 and 3 together and provide
management recommendations is in preparation.

��-79-

MIDDLE PARK DEER STUDY PRODUCTIVITY AND MORTALITY
R. Bruce Gill

P. S. OBJECTIVE
To estimate increments and losses to the Middle Park deer population in
order to formulate more efficient harv~st. regulations.

SEGMENT OBJECTIVES
1.

Complete doe collections initiated in the previous segment.

2.

Estimate mortality rates of Middle Park deer over the entire winter
range.

3.

Publish the results of this job.

METHODS AND MATERIALS
See Gill (1969) and Gill (1970) for methodology details.

RESULTS AND DISCUSSION
Doe collections were completed in April, 1972 and these data were reported
by Gill (1972).
Over-winter mortality for 1971-72 was assessed and these data were reported
by Gill (1972).
Several pUblications have already emanated from this job. These are listed
in the bibliography section. A Division Special Report is currently being
prepared that will summarize all of the production and mortality data obtained
during the five year study. Recommendations for management will be an
essential part of this publication.

BIBLIOGRAPHY
Gill, R. B.

1969.

An apology for a deer season.

Colo. Outdoors.

18(6):30-33.

1969. Middle Park deer study - productivity and mortality. Colo.
Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. W-38-R-23.
Game Res. Rep. July, Part 1. pp. 105-122.
1970. Middle Park deer study - productivity and mortality. Colo.
Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. W-38-R-24.
Game Res. Rep. July, Part 3. pp. 337-354.

�-80-

Gill, R. B., O. C. Wallmo, and J. G. Nagy. 1970.
study. Colo. Outdoors. 19(5):14-15.

The Middle Park deer

Gill, R. B. 1971. Middle Park deer study - productivity and mortality.
Colo. Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj.
W-38-R-25. Game Res. Rep. July, Part 2. pp. 189-207.
1972. Productivity studies of mule deer in Middle Park, Colorado.
Mule Deer Workshop, Proc. 2:40-47.
1972. Middle Park deer study - productivity and mortality. Colo.
Div. of Wildl. Game Res. Div. Fed. Aid Proj. W-38-R-26. Game Res.
Rept. July, Part 2. pp. 179-198.
Wallmo, o. C., and R. B. Gill. 1971. Snow, winter distribution and population dynamics of mule deer in the Central Rocky Mountains. Snow
and Ice in Relation to Wildlife and Recreation Symposium, Proc. 1:1-15.

-"---

Prepared by

"----"

,

(.~

,

,,-="'&gt;,(t

~
&gt;,~);") \ \
R. Bruce Gill
Wildlife Researcher

,1

�-81-

JOB

State

of

Project

~CO~L~O~RA~D~O~

plan

Job

Tit Ie :__

Period

1973

REPORT

_
Deer-Elk

14

No.

Job

Investigations
4

No.

.:;.M~i::.:d::.:d::;l::.:e~P::..:a::.r~k::...D~e~e,;;,r.....!;;S~t;;:u:.::d:..2y:....--.-....::P:.
c:::;a:::.:::.l....:::C:::.h::::a,;;,r..::a:;:c:;:t:;:e:.
c:.:s:-;a::.n:,;d::....::..F,.::o,.

Covered:

Personnel:

PROGRESS

W-38-R-27

No.

Work

July,

O.C.
L.R.

April

1, 1972

through

March

Wallmo,
D.W. Reichert,
Carpenter,
W.J. Adrian,

31,

1973.

W.L. Regelin,
R.E. Keiss,

D.L. Baker,
L.A. Roper.

R.B.

Gill,

ABSTRACT

Forage
selections
of tame deer were observed
during
715 deer-hours
of grazing
in nine seasonal
periods
during
the year.
Forbs and shrubs predominated
in
the deer diets April through
September.
In fall the leaves and a small amount
of the current-growth
stems of Populus
tremu1oides,
Ame1anchier
a1nifolia,
Prunus virginiana
and Salix spp. comprised
about 40 percent
of the diet.
Only
in December,
January
and February
were woody stems a significant
component.
Grasses
were a negligible
part of the diet in summer but prominent
in fall,
spring,
and especially
winter.
Crude protein
in the diet was below levels
considered
adequate
from October
through
April.
Apparent
digestibility
was
highest
in August
and lowest in April.

��-83-

MIDDU~ PARK DEER STUDY··
PHYSICAL CHARACTERISTICS AND FOOD HABITS

O. C. Wallmo and R. Bruce Gill

P. S. OBJECTIVE
To measure selected physical characteristics related to the assessment of
physical condition of deer and to determine forage preferences of mule
deer in Middle Park.
SEGMENT OBJECTIVES
1.

Analyze and report data compiled from measured, selected physical
characteristics of deer which are believed to be indicative of their
physical condition.

2.

Identify deer forage items on:
a.
b.
c.

Alpine and alpine margin areas.
Transitional fall ranges.
Transitional spring ranges.

METHODS AND MATERIALS
Procedures were reported by Gill (1969) and Roper (1970, 1971 and 1972).
Under other phases of the Middle Park Deer Ecology program, tame, trained
mule deer have been used to determine their forage preferences in various
seasonal habitat types. During this project segment, seasonal food habits
sampling was completed on spring and fall range, samples of the major foods
in the diet were collected for all seasonal periods, and nutritional analyses
of the samples were initiated.
The food habits sampling method begins with training newborn fawns to be
dependent physchologically on the handlers, to willingly enter a truck and
ride to the study areas, and to permit the handlers to follow them closely
while they graze at will (Reichert 1972). A total of 20 different deer
(10 males and 10 females), including fawns (beginning at 2 months of age)
and adults (up to 42 months age), were used.
Each day during a sampling period, one to four deer - each with an observer were taken to the study area early in the morning and late in the afternoon
and allowed to roam as long as they grazed effectively. Data were recorded
on portable tape recorders as numbers of bites taken of each species and
plant part (procedures in Wallmo et al. 1972). Study areas were subjectively
chosen as being a representative portion of the habitat types used by wild
deer during that period. A minimum of 10,000 bites was arbitrarily set as
the sample goal for each period. Altogether, 243,365 bites were recorded
in 715 deer-hours of grazing during 509 individual deer outings.

�-84-

The habitat types and localities sampled are:
June-July

lodgepole-spruce-fir, 9,500-11,500 feet,
Fraser Experimental Forest

August

lodgepole-spruce-fir, 9,500-11,500 feet,
Fraser Experimental Forest

September

lodgepole-spruce-fir, 9,500-11,500 feet,
Fraser Experimental Forest

October

conifer-aspen-sagebrush ecotone, 8,500-9,300
feet, 4" miles N of Parshall

November

conifer-aspen-sagebrush ecotone, 8,500-9,300
feet, 4 miles N of Parshall

December-January

mid-elevation sagebrush-bitterbrush-snowberry,
8,000-8,400 feet, 2-1/2 miles N of Parshall

January-February

low-elevation sagebrush, 7,800 feet, 3 miles
SE of Kremmling

April

same as October-November, 8,500-8,700 feet

May

same as October-November, 8,700-9,000 feet

At the end of each sampling period from each species approximately 300 g of
plant material representative of that used by the deer were collected from
the 10-15 leading species in the composite diet of all deer. The samples
were frozen for storage, and later oven-dried and ground in a Wiley mill
through a 0.5 mm screen for analysis. In vitro digestions were conducted
in triplicate by the method of Tilley and Terry (1963) with Pearson's (1968)
modification. Rumen fluid for the digestions was obtained from three deer
maintained on a uniform, nutritionally balanced diet; the three samples
were mixed to avoid the influence of individual variations. Crude protein
content was estimated by the microkjeldal method (A.O.A.C. 1955) with
selenium as the digestive catalyst. Cell-wall constituents were determined
by techniques outlined by Van Soest and Wine (1967). Cellulose and lignin
were separated by the procedure of Van Soest (1963).

RESULTS AND DISCUSSION

Physical Characteristics
A Division Game Information Leaflet listing the physical characteristics of
deer in Middle Park is being prepared.
A portion of the physical characteristics data from this job was presented
in July at the Western Association of Game and Fish Commissioners at Salt
Lake City. Title of the paper, "Estimating Winter Losses of Mule Deer in
Northern Colorado Using a Winter Severity Index".

�-85-

Seasonal Forage Selections
Table 1 lists for each season the plants that ranked among the first 20
in total bites eaten and comprised at least 1 percent of the composite
diets.
It includes 77 species.
An additional 117 species were grazed
but are not listed. Many of them (61 species) never comprised as much
as 10 bites in anyone
grazing trial.
Seasonal contribution of trees and shrubs, forbs, and grasses, sedges,
and rushes to the diet is shown in Figure 1. Forbs and subshrubs predominated April through September.
In summer, leaves and terminal portions
of current-growth stems of the low shrubs Vaccinium scoparium and y.
myrtillus comprised two-thirds of the diet. In fall the leaves and a
small amount of current-growth stems of Populus tremuloides, Amelanchier
alnifolia, Prunus virginiana, and Salix spp. comprised about 40 percent of
the diet. Only in December, January, and February were woody stems
(especially of Purshia tridentata, Amelanchier alnifolia and Symphoricarpos
oreophilus) a significant component.
Forbs, primarily Eriogonum umbellatum, made up the largest part of the diet
in April and May, and in a small March sample.
Through summer and fall,
forbs constituted only 20 percent of the intake, but in each of those seasons
63 different forb species were used.
Grasses were a negligible part of the diet in summer but prominent in fall,
spring, and especially winter.
In one 30-day sampling period from midJanuary to mid-February grasses made up 70 percent of th(~ diet in 79 grazing
observations with six deer subsisting exclusively on native forage for that
period.
Agropyron spicatum far outranked all other species of grass used.
The three species that were used most heavily
periods are shown in Table 2.

Nutritional

in various

seasonal

sampling

Analyses

Tables 3, 4, and 5 list the digestible dry matter percentage (in vitro
digestibility),
crude protein content, and cell-wall content for all of
the species that were analyzed.
Figure 2 shows the mean level of digestibility and crude protein content of the diet of the tame deer for each
season (the level of the constituent in the forage times the percentage of
that forage in the diet). Approximately 7 percent crude protein is considered necessary for body maintenance in deer. The diet means were well
below that level October through April.
The cellulose and lignin determinations were not completed in time for this report, and cell-wall
constituents alone are not a useful measure of nutritional limitations so
they are not presented in Figure 2. (The uncompleted portions will be
presented next year in conjunction with the report on nutritional investigations under Job No.7).

�-86-

LITERATURE CITED
Association of Official Agricultural Chemists. 1955.
analysis. 8th Ed. Washington, D.C. 1008 p.

Official methods of

Pearson, H. A. 1968. Digestibility trials: in vitro techniques. Pages
85-93 in Range and Wildlife Habitat Evaluation, A Research Symposium.
USDA For. Servo Misc. Publ. 1147. 220 p.
Reichert, D. W. 1972. Rearing and training deer for food habits studies.
USDA For. Servo Res. Note RM-208, 7 p. Rocky Mt. For. and Range Exp.
Stn., Fort Collins, Colo.
Tilley, V. M. A., and R. A. Terry. 1963. A two-stage technique for the in
vitro digestion of forage crops. J. Br. Grassl. Soc. 18:104-111.
Van Soest, P. J. 1963. Use of detergents in the analysis of fibrous feeds.
II. A rapid method for the determination of fiber and lignin. Assoc.
Off. Agric. Chern. J. 46:829-835.
Van Soest, P. J.,and R. H. Wine. 1967. Use of detergents in the analysis of
fibrous feeds. IV. Determination of plant cell-wall constituents.
Assoc. Off. Anal. Chern. J. 50:50-55.
Wallmo, o. C., W. L. Regelin, and D. W. Reichert. 1972. Forage use by mule
deer relative to logging in Colorado. J. Wildl. Manage. 36:1025-1033.

Prepared by

Q. (/J ?ckL12c&lt;A..A..,r--,

o. C. Wallmo
Principal Wildlife Biologist
Rocky Mountain Forest and Range Experiment Station

�-87-

Table 1.

Major species in seasonal diets.

Species

Dec.-Mar.

Apr.-May

June-Sep.

Oct.-Nov.

Trees and Tall Shrubs

Acer glabrum
Amelanchier alnifolia
Artemisia tridentata

* 1:./

+

+

*
+

Ceanothus velutinus
Chrysothamnus nauseosus
Juniperus scopulorum

+

Lonicera involucrata

+

Pinus contorta

+

Populus tremuloides

*
+

Prunus virginiana
Purshia tridentata

*

+

+

Ribes cereum
Ribes lacustre
+

Rosa acicularis

+
+

Salix spp ,

*

Salix brachycarpa
Sarcobatus vermiculatus

+

Shepherdia canadensis
Symphoricarpos oreophilus

+

---------------------_ .. _-------------------------------------------------------------------

�-88-

Table 1.

Major species in seasonal diets (continued).

Species

Dec.-Mar.

Apr.-May

June-Sep.

Oct.-Nov.

Sub-Shrubs -2/
Arctostaphylos ~-ursi

+

Berberis repens
Chrysothamnus viscidiflorus

*

Pachystima myrsinites

+

Salix anglo rum
Vaccinium scoparium and myrtillus

*
Forbs 2:..1

Arnica cordifolia
Artemisia fr1gida

+

+

Artemisia scopulorum
Aster spp.

+
Balsamorhiza sagittata

+

Castilleja flava
Clematis hirsutissima
Epilobium angustifolia
Eriogonum umbellatum

+
+

*

*

Fragaria ovalis
Geum rossii
Gutierrezia sarothrae

+
+

Lathyrus leucanthus

+
Parmelia chlorochroa

----------------------------.--.-----~~-------------------------------------------------------

�-89-

Table 1. Major species in seasonal diets (continued).

Species

Dec.-Mar.

Apr.-May

June-Sep.

Pedicularis bracteosa

+

Pedicular is racemosa
Penstemon spp.
Penstemon caespitosus

*

+

Penstemon cyathophorus
Phlox bryoides
Phlox multiflora

*
+

Polygonum bistortoides
Potentilla spp.
Potentilla concinna

+

Pyrola asarifolia and/or minor
Saxifraga arguta
Saxifraga bronchialis
Senecio spp.
Senecio crassulus
Sibbaldia procumbens
Streptopus amplexifolius
Taraxacum officinale

+

Trifolium repens

+

Grasses and Sedges
Agropyron sp.
Agropyron desertorum

Oct.-Nov.

�-90Table 1.

Major species

Species

in seasonal

diets

(continued).

Dec.-Mar.

Agropyron

smithii

+

Agropyron

spicatum

*

Bouteloua

gracilis

+

Apr.-May

June-Sep.

Oct.-Nov.

+

Bromus anomalus

----

+

Bromus ciliatus
Carex arapahoensis
Care x geyeri
Deschampsia
Oryeopsis

+
caespitosa

hymenoides

Poa spp.

+

Poa fendleriana

+

Poa juncifolia
Poa secunda
Sitanion

hystrix

Stipa comata
Stipa pinetorum

+

+

*

!/

The symbol
indicates that the plant comprised over 10.0% of the foods consumed in
at least one of the seasonal sampling periods; + indicates 1.0 to 10.0%, and _ indicates
less than 1.0%.

~/

Low shrubs are those generally less than 30 cm. in height.
Forbs include all herbaceous
plants that do not fit in another category and some low shrubs with primarily herbaceous
stems.

�100

80

FORPS;

bOI-j I 11111

SUBSHRUPS

r

r

I

r

I

r

I

I

I&amp;,j

IIIIIIIIIIIIIIIIIIIIIIIIIIII~~

•••••••••••••••••

lOFlI.

I

r

I

r

I

•

_,

I

11111"'i'IIIJIf'IIIIIIIII

I

I

I

I

I

r

r_;

II:!

t-t;;

! IVII I :;0
t%j

(")
t%j

Z

1-3

4o~1 I I I I I I I I I I I I I I I I I I I I I I II

I

IIII II III I I IIII ! II ~

1.0

o
~
t::::I
H
t%j

20R-llllllllllllllilll.IIIIII~1111111-3

JuL

Fig. 1.

AUG

SEP

MAR

Contribution of major forage classes to the diets of tame mule deer in Middle Park.

MAY

";"'

�-92-

Table 2. The three (3) leading forage species in the composite diets of tame deer
during seasonal sampling periods.

Season and
sample size

Species

Major parts

Percent

of plant

of diet

December-January

Purshia tridentata

stems

39.9

47,128 bites

Agropyron spicatum

all

18.8

Symphoricarpos oreophilus

stems

9.5

January-February

Agropyron spicatum

all

40.4

82,199 bites

Artemisia tridentata

leaves &amp; stems

11.6

Phlox bryoides

leaves &amp; stems

9.6

Eriogonum umbellatum

leaves &amp; stems

44.3

All grasses 1/

new growth

24.3

Populus tremuloides

stems

7.3

April-May
20,613 bites

July-September

Vaccinium spp.

leaves &amp; stems

62.2

58,175 bites

Salix brachycarpa

leaves &amp; stems

10.2

leaves,stems,flowers

4.5

Epilobium angustifolium
October-November

Populus tremuloides

leaves

23.6

21,604 bites

Eriogonum umbellatum

leaves &amp; stems

13.4

Amelanchier alnifolia

leaves

7.5

1/

Grasses lumped due to difficulty of identification.

�Table 3.

Percent digestible dry matter of major forage species (mean of 3 determinations).

Collection Dates
.-l
.-l

~I
4J
~

ro

Agropyron

!-I

p..,p..,ro

.-l

Species

.
~

00

"

~

M

III
4J

I

ro

.

.0

Gl

smithii

LS

45.9

48.8

Agropyron spicatum

LSH

38.5

47.5

Ame1anchier

L

--

a1nifo1ia

S
Arctostaphylos

uva-ursi

.

\0
.-l

.
eo

M

.-l
I
00

Po

&lt;.J

~

Gl

U

t/)

M
.-l

r-...

.

0

.

.

Artemisia frigida

LS

Artemisia

LS

tridentata
sagittata - green
dry

Bromus anomalus

LS

Carex geyeri

LS

Ceanothus velutinus

LS

Chrysothamnus

LSH

.

39.1

52.2

20.1

12.8

28.1

17.3

45.9

31.5

37.0

36.3

28.2

28.4

.-I
;j

"

0

&gt;
z
0

42.2

61.4

.I

67.8

66.1

I

37.5
64.2

53.5
70.8

L
L

...;

«

L

Bouteloua gracilis

C\

:&gt;.
~

!-I
Po

~

LS

viscidiflorus

35.1

LS

-----

Arnica cordifo1ia

Balsamorrhiza

36.2

0

00
.-l

24.8

27.3

33.2
39.4
47.2

48.1

49.1

52.0
4L..4

42.6

41.5

-------------------------------------------------------------------------------------------------------------------------

�Table 3.

Percent digestible dry matter of major forage species (mean of 3 determinations)

(continued).

Collection Dates
r-I
r-I

:::;-,
Species

~~lobium

.j.I

r::

CJ)
.j.I

(Ij

1-1

r-I

(Ij

p..p..

angustifolium

Eriogonum umbellatum

....,

(Ij

co

.

.0
Q)

r:r..

0

co

r-I

.

1-1

p..

&lt;

r-I

\0
r-I

:&gt;,
(Ij

)::

7.8

15.8

11.4

'l:../

Lathyrus leucanthus

,.....

M
r-I

r-I

....,::l

eo
~

47.3

52.7

.

57.5

LS

54.4

.

0

M

I

.

co

0\
r-I

p..

.j.I

Q)
Cf.l

u

:&gt;

0

0
Z

17.9

18.7

53.0

LS

Pachystima myrsinites

LS

Penstemon caespitosus

LS

Phlox bryoides

LS

Phlox multiflora

LS

Poa fendleriana

--

LS

Populus tremuloides

L

- from ground

.

49.8

69.0

56.3

LS

Lonicera involucrata

.

51.4

18.6

L

Grasses, mixed, unid.

Prunus virginiana

.
r::

I

LSH
LSH

Fragaria oval is

M

53.4

I

\0

~

64.5

64.5

I

65.5
42.0

55.0

42.7

46.0

39.8

48.1

57.9

16.3

44.7
39.5

S

45.9

44.4

L

20.1

16.7

46.3
34.8

L

51.8

S

34.8

-------------------------------------------------------------------------------------------------------------------------

24.4

�Table 3.

Percent digestible dry matter of major forage species (mean of 3 determinations)

(continued) .

Co llec tion Dates
~I

("")

+J II)
~ +J
t1S

\-I
t1S

.-t

Species

p.,p.,

.

~
CIS
""J

.-t
.-t
I

co

.

co
.-t

.

\0

.-t

.c

\-I

r:..Q)

~

:&gt;.
~

29.9

25.7

36.6

r-..

("")

.-t

Ill)

.

;j

""J

.-t

.

0
("")

.

Po

.;J

U)

Q)

0
.-t
I
00

0\

+J

&gt;0

.

.-t

.

u

0

z

Purshia tridentata

S

Pyro1a spp. 11

LS

36.4

45.8

47.1

Rosa acicu1aris

LS

55.4

53.2

50.3

Salix spp.

L

38.1

26.0

S

43.3

43.0

--

--

I

LS

49.2

Saxifraga arguta

LS

62.9

Senecio crassu1us

LS

Senecio sp.

LS

76.4

Shepherdia canadensis

LS

Stipa pinetorum

LS

Streptopus amp1exifo1ius

LS

Symphoricarpos

S

-

-

or eophLl.us

Taraxacum officina1e
Vaccinium

spp.

il

L
LS

50.4

"

47.5

v

I

45.3

33.7

38.0

33.0

48.0

32.3

67.2

65.4

76.6

74.4

45.2

50.3

53.2

71.4

73.7

72.6

64.7

66.8

69.7

32.9

33.3

33.8

64.3

34.5
35.2

28.4

------------------------------------------------------------------------------------------------------------------------

�J)
L = leaves, S = stems, H = inflorescence or fruiting stalk.
2/
- When unidentifiable,

all grasses used by deer were collected together.

3/
- P. minor or P. asarifolia or both.

!!/
V. scoparium or y. myrtillus or both.

I
\ji)

m
I

�Table 4.

Crude protein content (percent) of major forage species

(mean of 2 determinations).

Collection Dates
0

.-j
.-j

~I

.-j

00

='
&lt;

.

.-j

Po

~

+J

f.)

U)

0

&gt;
0

.

~~

...,

.c

~
~

~

4.6

7.2

6.5

5.2

3.8

4.6

4.5

4.8

.

rz,.~

LS

5.5

3.1

Agropyron

spicatum

L8H

3.0

2.3

L

alnifo1ia

&gt;..

.

I

0\

00

C"')

.
§

smithii

Ame1anchier

.

\0

,...

.-j

C"')

Agropyron

--

.-j

00

0

00

+J (/j
I=l +J
Cd ~
.-j Cd

Species

I

C"')
.-j

S

4.6

4.4

.-j

.

...,='

.

z

3.3

I

\0

-..J

Arctostaphylos

Arnica cordifo1ia

LS

Artemisia

frigida

LS

Artemisia

tridentata

LS

Ba1samorrhiza

3.8

LS

uva-ursi
-----

sagittata - green
- dry

Bromus anoma1us

LS

Carex geyeri

L8

Ceanothus ve1utinus

LS

Chrysothamnus

L8H

viscidif10rus

11. 7

5.6

6.7
8.1

9.3
24.5
4.4

L
L

--

18.2

L

Boute1oua gracilis

I

4.2

5.5

5.3
5.1
4.6

5.4

4.7

4.5
8.9

4.9

6.8

------------------------------------------------------------------------------------------------------------------------

�Table 4.

Crude protein content (percent) of major forage species (mean of 2 determinations) (continued).

�Table 4,

Crude protein content (percent) of major forage species (mean of 2 determinations)

(continued)"

Collection Dates
..-I
..-I

:::;-1

.

00

...,

cu
J:x.&lt;

C")

+J rJ)
~ +J

co •.•

..-I

Species

t1l

p..p..

I _

~

t1l

.

..0

0

..-I
00
..-I

.

C")

\0
..-I

&gt;-

•••
~

~

6.9

7.3

".
..-I

..-I

.

be

0

I

0\
..-I

+J

0

&gt;
z0

.

.

00

cu

(,)

C")

Po.

...,::s

~

en

,

Purshia tridentata

S

Pyro1a spp. 1./

LS

10.9

9.9

9.8

Rosa acicu1aris

LS

16.3

12.2

7.2

Salix spp.

L

5.4

9.4

S

5.6

7.4

--

7.2

LS

18.7

13.2

10.5

Saxifraga arguta

LS

20.6

14.3

9.6

Senecio crassu1us

LS

12.0

8.0

Senecio sp.

L8

Shepherdia canadensis

LS

23.8

16.6

15.5

Stipa pinetorum

L8

Streptopus amp1exifo1ius

LS

21.7

14.9

8.6

Symphoricarpos oreophi1us

S

Taraxacum officina1e

L

20.6

13.8

11.1

Vaccinium spp. fl..!

L8

13.9

10.8

9.4

I

\0
\0
I

-

-'--

--

15.6

6.2

4.5

4.9

5.0

5.4

3.6

10.2

7.3
14.4

5.4

�Table 5.

Percent cell wall constituents of major forage species (mean of 2 determinations)

Collection Dates

:::tl

,.....j
,.....j

.

('f)

+J

=

I'IS +J
J.I
,.....j CIS

Species

~ropyron

CIl

~~

=

I'IS

~

I

00

.
,c
OJ

~

smithii

LS

63.1

68.8

A~ropyron §picatum

LSH

69.5

74.4

Amelanchier alnifolia

L
S

Arctosta~hylos ~-ursi

58.6

60.2

LS

00
,.....j

.

0
,.....j
\0

,.....j

,.....

.

('f)

,.....j

.
eo

0

I

.

('f)

00

.

0\

,.....j

.

J.I

e-.

,.....j

~

~

~

53.2

62.9

50.3

55.9

44.1

40.6

56.4

59.5

54.1

54.8

33.9

27.6

;:J

~

Po.
OJ

+J

U

en

0

:&gt;

0

z

70.5

I

t-'

Arnica cordifolia

LS

Artemisia frigida

LS

Artemisia tridentata

LS

Balsamorrhiza sagittata - green

L

- dry

39.6

L

Bromus anomalus

LS

Carex geyeri

LS

0
0

29.7

I

47.5
28.4

41.9
40.7

L

Bouteloua gracilis

31.7

36.7

38.7

59.3
67.0

Ceanothus velutinus

62.7

62.8

64.9

58.4

LS
24.3

Chrysothamnus viscidiflorus

LSH

50.6

47.0

--------------------------------------------------------------------------------------------------------------------

�Table 5.

Percent cell wall constituents of major forage species (mean of 2 determinations)

(continued).

Collection Dates
...-i
...-i

::;-,
.j..I
~

tlI

C/l
.j..I
101

...-i tlI
~~

Species

Epi10bium angustifo1ium

LSH

Eriogonum umbe11atum

LSH

Fragaria ova1is

"t,

LS

Lathyrus 1eucanthus

LS

Lonicera invo1ucrata

LS

Pachystima myrsinites

LS

Penstemon caespitosus

LS

Phlox bryoides

LS

Phlox multiflora

LS

Poa fend1eriana

LS

Populus tremu10ides

L

Prunus virginiana

tlI

~

36.2

0

00

...-i

.0

101

.

Qj

~

38.1

...-i

00

.

\C

r--

&gt;.
)!l

;:I

...-i

c,

&lt;:

38.7

58.7

.
...-i

M

0

...-i

M

;:I

Qj

.
eo

~

-e

tf.l

29.4

27.6

20.5

40.4

36.0

I

.

00

.
~

32.1
~

L

Grasses, mixed, unid. ];./

- from ground

.
s::

M

I

26.5

.j..I

t)

35.6

32.1

'"

33.1

26.1

34.5

I
•....
0
•....
I

33.6

39.1

~

33.7

43.1

28.8

.

Z

51.2

27.1

...-i

0

46.0

22.5

0\

24.6

23.4

67.9
43.4

S

43.3

45.2

49.5

44.3

L

46.0

51.4

35.5

41.1

L

27.8

S

53.9

----------------------------------------------------------------------------------------------------------------------

�Table 5.

Percent cell wall constituents

of major forage species

(mean of 2 determinations)

(continued).

Collection Dates

....•

•...•

~J
~

II)

=~

CIS 1-1
••••• CIS

Species

.

C"')

Il-IP-&lt;

...,=

CIS

I

co

.

,.0

co
....•

.

1-1

....•
\0

:&gt;.

~

)¥

44.4

53.2

Q)

~

0
....•

.

,....
•...•

C"')

•...•

.

CIO

0
C"')

.

co

I

0\
•...•

Po.
Q)

~
CJ

0

:&gt;
~

43.2

51.2

29.6

44.0

...,:::I

~

CI.l

Purshia tridentata

S

Pyrola spp. 1/

LS

32.3

28.7

28.0

Rosa acicularis

LS

37.1

33.6

22.7

Salix spp.

L

45.6

S
L8

41.0

33.6

.

.

29.9
I

8axifraga arguta

LS

34.8

27.7

I-'
0

22.3

N
I

Senecio crassulus

LS

Senecio sp.

LS

Shepherdia canadensis

LS

Stipa pinetorum

LS

Streptopus amplexifolius

LS

Symphoricarpos

S

oreophilus

Taraxacum officinale
Vaccinium spp. if

L
LS

33.0

26.1

44.7

33.0

28.7

39.2

34.1

28.7

37.9

30.8

25.7

51.5

47.4

40.6

17.3

66.9

59.5

69.9

66.2

61.6

71.2

57.0

60.8
22.4

56.7

�16

42

15

40
/" .•••.
__
I

D~D.M.
-""

I

,

,

14

13

",
,
"" "
,,'
"

v

~-",

~ ,~
;I'

36

'"

"

Protein

12

38

,

,

"
"

I

,
"

,

"

iN!

H

~
E-i
~
~

34

,
I

,

11

..
z

I

,-'

I

32

I

"

~

10

30

9

28

··
~
~·
:E!

I

I-'
0

W
I

8

26

7

24

6

22

5

20

4

18

JUL

AUG

SEP

OCT

NOV

DEC

JAN

FEB

MAR

APR

MAY

JUN

WEIGHTED MEAN OF MAJOR DIET ITEMS
Fig. 2. Seasonal variation in digestible dry matter (D.D.M.) and crude protein content
.of the diets of tame mule deer in Middle Park.

��-105-

July, 1973

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~---------

Project No.

W-38-R-27

Work Plan No.

14

Job Title:

Job No.

5

Middle Park Deer Study - Range Fertilization

Period Covered:
Personnel:

Deer-Elk Investigations

April 1, 1972 through March 31, 1973

R. Bruce Gill, Paul F. Gilbert, Laren A. Roper, O.C. Wa11mo, Dan L.
Baker, Lonnie M. Brown, Jan L. Wassink and Len H. Carpenter.

ABSTRACT
All field work was completed during the project year. A final report is being
prepared and will be published in the July, 1974 Game Research Report.

��-107-

MIDDLE PARK DEER STUDY - RANGE FERTILIZATION
Len H. Carpenter

P. S. OBJECTIVE
To test the effects of fertilizers on deer forage and deer feeding intensity responses to fertilization of critical winter range areas.

SEGMENT OBJECTIVES
1.

Test the effects of fertilizers on forage production and composition
on the selected study areas.

2.

Analyze and summarize all data from all segments of this study and
prepare a final report and a PhD Thesis. Begin work on publication(s)
of study results.

METHODS AND MATERIALS
Methods and materials have been detailed previously by Carpenter (1970 and
1971) .

RESULTS AND DISCUSSION
All vegetation measurements pertinent to this study were completed during
the segment. Data compilation and analysis were initiated. Plans called
for a final report to be written during this segment, however, the complexity and volume of data obtained from this study have made it impossible
to maintain this reporting schedule. Rather than prepare a meaningless,
superficial report at this time it has been deemed advisable to devote all
energies towards completing the final report. This will be prepared as a
PhD thesis and will be published in the July, 1974 Game Research Report.
LITERATURE CITED'
Carpenter, L. H. 1970. Middle Park deer study - range fertilization. Colo.
Div. Game, Fish and Parks. Game Res. Div., Fed. Aid Proj. W-38-R-24.
Game Res. Rept. July, Part 3. pp. 371-391.
1971. Middle Park deer study - range fertilization. Colo. Div.
Game, Fish and Parks. Game Res. Div., Fed. Aid Proj. W-38-R-25.
Game Res. Rept. July, Part 3. pp. 225-253.

Prepared by

��-109-

July, 1973

JOB PROGRESS

State of

COLORADO
----------~~~~---------

Project

No.

Deer-Elk

W-38-R-27

Work Plan No.
Job Title
Period Covered:
Personnel:

REPORT

Investigations

.14
6
Job No.
Middle Park Cooperative Deer Study Physiology and Prevention of Deer Starvation
June, 1972 through June, 1973.

Julius G. Nagy, Principal
Student, David E. Reeder,

Investigator, David S. DeCa1esta,
Graduate Student.

Graduate

ABSTRACT
Eleven adult male mule deer were fed ad libitum concentrate and alfalfa hay
approximately three months pre-trial.
During the trial, six deer were fed an
alfalfa hay diet and five were fed native hay. Two levels (37 and 75%) of
each hay were mixed with a concentrate mixture.
The five deer fed the native
hay diet consumed more of the pelleted ration than the deer fed the pelleted
alfalfa hay ration.
Chemical composition of both rations were similar in
moisture, crude protein, ash, cell wall constituents and acid detergent fiber.
However, lignin composition of the alfalfa hay diet was considerably higher
than that of the native hay diet.
Thirteen mule deer were used to study the effects of starvation and refeeding
upon deer physiology.
During starvation deer catabolized stored fat and muscle
protein, indicated by increased blood levels of free fatty acids and urea
nitrogen.
As a result deer lost weight, which was linearly related to length
of starvation.
Stored fat seemed to be the major source of endogenous energy
used by starving deer. Deer that died as a result of starvation had exhausted
fat reserves and increased catabolism of muscle protein.
Under these conditions
glucose production was inadequate for sustenance.
Starved deer showed marked
hypoglycemia, glucose levels being 24-32.6 mg percent.
Glucose levels in deer
surviving starvation were 73 mg percent or higher.
Refeeding
toxicity,

starved deer either good or poor quality
liver malfunction or high mortality.

diets did not result

in

Analysis of blood from a doe that died during refeeding revealed occurrence of
hypoglycemia.
Other blood parameters were similar to those for deer that
survived with refeeding.
After 10 days of refeeding, blood levels of most
energy-metabolites
were similar to those of non-starved deer, indicating that
metabolism of deer refed for 10 days had normalized.

��-111-

MIDDLE PARK COOPERATIVE DEER STUDY PHYSIOLOGY AND PREVENTION OF DEER STARVATION
Julius G. Nagy

P. S. OBJECTIVE
To ascertain what takes place in deer physiology when starvation occurs
and to investigate the economic and biological feasibility of preventing
starvation.
SEGMENT OBJECTIVES
1.

To publish reports on:
a.
b.

2.

Responses of mule deer to alfalfa feeds,
Responses of rumen microflora of mule deer to complete starvation.

To measure physiological responses of mule deer starved in pens to
refeeding with:
a. A moderately digestible high protein diet,
b .. A poorly digestible low protein diet.

3.

To test diets for refeeding deer kept in field pens and malnourished
to simulate natural conditions during a severe winter.

4.

Prepare report.
METHODS AND MATERIALS

Alfalfa Hay vs. Native Hay as Deer Feed
In previous experiments (Schoonveld 1971) deer experienced low food intakes
and digestive problems when fed diets containing alfalfa stems but did not
evidence these problems when fed a native hay diet. This experiment was
designed to investigate if compounded diets, containing various percentages
of alfalfa or native hay, would result in digestive problems.
Eleven adult mule deer, weighing 60-75 kg and judged to be in good health,
were used. Five were adult castrated males, and six were adult females. All
had been raised in captivity and were fed ad libitum a high quality ration,
including alfalfa hay for 3 months before the experiment. During the trial,
they were confined individually in pens measuring 6.1 X 1.8 X 2.4 m high.

�-112-

Five rations (Table 1), including their normal ration as a control,
were ground and pelleted. The control ration was a concentrate
without hay. For the four test rations, alfalfa and native hay
were each substituted to provide 37 and 75 percent of the diet.
Alfalfa hay used was a leafy, third cutting and the native hay was also
of good quality, containing mostly timothy (Phleum pratense) and sedge
(Carex spp.). Native hay was obtained from North Park, Colorado. All
five rations contained similar levels of protein (Table 2). Except for
lignin content, the diet containing 37 percent alfalfa was similar to
that containing 37 percent native hay and the diet containing 75 percent
alfalfa was similar to that containing 75 percent native hay (Table 2).

Table 1. Compositions of five pelleted rations used
in the alfalfa hay
and native hay feeding trials.
ComEosition (Percent Air-Dr~ Weight)
Corn
Protein Supp •.
!/ Native Hay

Ration

Barley

Control

60

20

20

0

0

N 37

38

12.5

12.5

37

0

N 75

15

5

5

75

0

A 37

38

12.5

12.5

0

37

A 75

15

5

5

0

75

Alfalfa Hay

1/

- 32 percent protein supplement.

Table 2. Chemical composition of five rations used in the alfalfa hay and
native hay feeding trials.

Ration

Cru e
Protein

Control

Com osition
Ash

Constituents

ADF !/

Lignin

14.7

5.2

74.6

14.2

3.6

N 37

14.9

5.4

70.2

24.0

3.5

N 75

15.4

6.4

64.4

32.1

3.4

A 37

14.8

6.5

69.7

23.1

5.4

A 75

15.6

6.9

63.9

32.3

7.3

1/

- Acid-detergent fiber.

�-113-

Deer were divided into two groups with six rece1v1ng the diet containing
alfalfa and five receiving the diet containing native day. A 3 X 3 Latin
Square design was used in the experiment, the rations being randomized
among 3 groups of deer in 3 consecutive 7-day periods (Tables 3 and 4).
In each sequence, the diet was changed after each 7-day period. Food
consumption by each deer was measured daily. Throughout the experiment
water was available ad libitum and intake not measured.

Table 3. Average daily food intake (grams) of six deer fed alfalfa-hay
rations during 7-day periods !/. Rations control (0), A 37 and A 75 contained
0, 37, and 75 percent alfalfa, respectively.
Deer
Number

I

Period
II

III

0

A 37

A 75

5

392

1078

854

6

913

442

1253

Mean

652

760

1053

A 37

A 75

0

1

1466

260

1078

2

1945

226

1522

Mean

1706

243

1300

A 75

0

A 37

3

590

1103

861

10

1102

1028

962

Mean

846

1065

911

!/
0 (control) , overall mean intake = 1006 g.
37, overall mean intake = 1126 g.

A
A

75, overall mean intake

= 714

g.

�-114-

Table 4. Average daily food intake (grams) of five deer fed native-hay
rations during 7-day periods l/. Rations control (0), N 37 and N 75 contained 0, 37 and 75 percent native hay, respectively.
Deer
Number

11

7

8

Mean

4
9

Mean

Period

I

II

III

0

N 37

N 75

972

1851

1948

N 37

N 75

0

298

1182

761

1931

1526

947

1114

1354

854

N 75

0

N 37

1450

138

1542

1371

186

1115

1410

162

1328

II
- 0 (control), overall mean intake = 601 g.
N 37, overall mean intake

1347 g.

N 75, overall mean intake = 1495 g.

Average daily food intakes for the last 5 days of each feeding period, within
each series (alfalfa and native hay), were analyzed using the procedure of
Dixon and Massey (1969:310). Average food intake between the alfalfa-hay
and the native-hay series were analyzed by the method of Dixon and Massey
(1969:175).

�-115-

Physiological Response of Deer Starved and Refed
From January 4 to January 13, 1973, food intake was measured for 13 deer
kept individually in isolation and fed ad libitum a diet containing 14.0
percent protein, 4.0 percent fat, 23.2 percent cell-wall constituents,
and 57.1 percent nitrogen-free extract. Five of the deer were then
selected as controls and the remainder matched to control deer on the
basis of mean feed consumption and body weight. A six-week conditioning
period to simulate malnutrition condition was conducted from January 16 to
March 1, 1973. In the conditioning period the control deer continued on an
ad libitum diet and the treatment deer were given one-fourth of their previous mean intake. At the end of the conditioning period all deer were
deprived of all feed until any three deer died, whereupon remaining deer
were refed the same diet ad libitum for 10 days. Jugular blood samples
and body weights were obtained at initiation of conditioning, after 3 weeks
of conditioning, at initiation of starvation, at initiation of refeeding,
and 10 days post-refeeding. Daily feed intake was recorded during refeeding.

Blood Samples
Forty ml jugular blood samples were withdrawn with a 50 ml syringe. Approximately 0.05 ml of blood was drawn from the syringe tip into a heparinized
capillary tube. Remaining blood was immediately placed in a 50 ml centrifuge
tube containing sodium-flourine heparin. Forty ml blood samples were centrifuged for 5 minutes at 3000 rpm and plasma withdrawn and frozen.

Blood Parameters Examined
Toxicity Indicators--Plasma urea nitrogen levels were determined from blood
samples collected during 1972 (see report for Segment 26) and 1973 spring
trials. Evaluation of plasma ammonia levels was desired as an index of
ammonia toxicity.
Liver Malfunction--Plasma levels of two liver enzymes, glutamic oxalacetic
transaminase (GOT) and ornithine carbamyl transferase (OCT) were determined
from 11 deer in the spring 1972 trial. Deer sampled were two non-starved
deer, four starved deer, four refed deer and the one deer that died during
refeeding.
Energy Metabolism--Levels of plasma glucose, urea nitrogen and free fatty
acids were determined from deer in spring 1972 and 1973 trials.
Hematocrits of deer from spring trials in 1972 and 1973 were determined by
placing centrifuged capillary tubes containing heparinized blood in an
International Micro-capillary Reader (International Equipment Co.) and reading
hematocrit from a sliding scale. Two drops of plasma from each centrifuged

�-116capillary tube were placed onto a Goldberg Tic Refractometer and optical
density recorded. Plasma total protein levels were obtained from conversion tables based on optical density. More detailed information
concerning methods and materials can be obtained from the PhO dissertation
of David S. deCalesta (1973).

RESULTS AND DISCUSSION
Alfalfa VS. Native Hay as Supplemental Deer Feed
No deer on either the alfalfa or native-hay rations showed serious symptoms
of digestive problems. However, some deer evidenced low consumption during
some periods, apparently due to abrupt changes in diets (Tables 3 and 4).
Average daily food intake of deer in both the alfalfa and native-hay series
showed no significant (P&gt; 0.10) effects due to periods (columns), animals
(rows), or to percent of hay in the ration (treatments) (Table 5). However,
sample sizes are small, variation among deer is large and treatment effects
are confounded with the sequence in which rations were fed. Further, the
short feeding periods (7 days) may not have allowed sufficient adjustment
to the rations.

Table 5. Analyses of variance of data from the alfalfa and native-hay
feeding trial.
Source

df

Mean Square

Alfalfa Hay Rations
Total

8

Columns (Periods)

2

151361.8

0.44

Rows (Order)

2

51357.4

0.15

Treatments (% Hay)

2

134469.4

0.40

Residual

2

340280.4
Native-Hay Rations

Total

8

Columns (Periods)

2

55598.1

0.10

Rows (Order)

2

321018.11

0.57

Treatments (% Hay)

2

217153.45

0.38

Residual

2

566183.4

II
- F ratios for columns, rows, and treatments are to be compared with
F.90 (2,2) = 9.00.

�-117-

Throughout the experiment (Tables 3 and 4) deer consumed more (P&lt;O.lO)
of the rations containing native hay than of the rations containing alfalfa.
This difference in consumption was greater for the rations containing 75
percent hay than for the rations containing 37 percent hay. Greater consumption of native hay may be related to the higher concentrations of
lignin in the alfalfa-hay rations (Table 2).
Physiological Response of Deer Starved and Refed
Experimental deer lost an average of 12.0 percent body weight during conditioning, whereas control deer gained an average of 3.7 percent (Table 6).
Mean PUN (plasma urea nitrogen) level for experimental deer at the third
week of conditioning was significantly lower (P~0.05) than when conditioning
began (Table 7). This was contrary to the expectation of an increase in PUN
level as a reflection of catabolism of muscle protein.
During conditioning mean level of plasma FFA (free fatty acids) for the
control deer did not change (Table 8). At the end of conditioning, mean
level of plasma FFA for experimental deer was significantly higher (P&lt; 0.1)
than that for control deer.
Mean level of plasma glucose for control and experimental deer exhibited
large, non-significant (P&gt;O.l) variations during conditioning (Table 9).
Mean hematocrit did not differ significantly (P&gt; 0 .1) between control and
experimental deer during conditioning (Table 10).
Mean level of plasma total protein did not differ significantly (P&gt; 0.1)
between control and experimental deer at the end of the conditioning (Table
11) •

Pre-,Starvation Nutritional Plane and Starvation Performance
Of eight experimental deer starved three died; one at 11, one at 12 and one
at 23 days of starvation. Difference between mortality of experimental deer
(37.8 percent) and control deer was not significant (P~ 0.1).
Percent weight loss during starvation for control deer was not significantly
different (P&gt;O.l) from that for experimental deer (Table 6). Percent weight
loss during the entire trial for experimental deer was significantly higher
(P &lt; 0.01) than that for control deer (Table 6).
A blood sample was collected immediately prior to death, from the experimental
deer dying on the 12th day of starvation. Mean PUN levels, plasma FFA,
plasma glucose, hematocrit and plasma total protein for control deer and
experimental deer surviving starvation were not significantly different
(P&gt; 0.1).

�Table 6.

Body weight changes of starved and refed deer.

Deer

Control

Experimental
deer surviving
starvation
Experimental
deer dying
during
starvation

Percent Weight
Loss During
Conditioning
n

5

5

X

-11. 9 ]j ,]j

Percent Weight
Loss,
Entire Trial

Percent Weight
Gain During
Refeeding

X

X

SD

X

SD

-

SD

+ 3.7 )j,]j

Percent Weight
Loss During
Starvation
SD

3.0

-14.8 Jj

5.6

-13.4 '])

7.5

+5.8

6.5

5.1

-16.4 1/

3.3

-25.9 1/

5.2

+6.7

3.7
I
•....
•....

00

3

.u 1.4

-12.2 ]./

I

-15.8 1/

2.7

1/
- Differences between means in columns significant

(P~0.05)

2/
- Differences between means in columns significant

(P&lt; 0.01).

1/
Differences between means in rows significant

(P&lt;O.l).

-26.7 ])

2.5

�-119-

Table 7. Means and standard deviations of plasma urea nitrogen concentrations 1/, spring 1973 trial.
Control Deer

EXEerimental Deer
SD

Trial Phase

X

SD

n

X

Begin
Conditioning J:.../

21.4 1/ ,§/ ,J../

3.6

4

25.1 1/ ,tE/ ,2./

3.6

8

3 !!;./ Week of
Conditioning 1./

20.4

u. 2./

3.7

5

18.4 §.../,Jj ,10/

1.6

8

Begin
Starvation !!;./

l6.41/,§.../,2../

3.0

5

20 5 1/ ,§.../,J../ ,
• 10/

4.3

7

Begin
Refeeding i/

16.7

8.3

5

22.6

10.9

6

Ten Days
Post-Feeding E.../

17.1

9.9

4

17.2

5.9

5

n

"

1/
- Mg urea nitrogen per 100 ml plasma.
2/
- Beginning of conditioning period, control deer fed ad libitum, experimental
deer fed one-quarter of control deer intake.
3/
- Twenty-third day of conditioning.
4/
- Forty-fourth day of conditioning, beginning of food deprivation, all deer.

Twenty-third day of food deprivation, beginning of refeeding, all deer.
6/
- Tenth day of refeeding.
7/
- Differences between means in rows significant at P&lt;0.05.
8/, 9/
- Differences between means in columns significant at P&lt;0.05.
10/
-- Differences between means in columns significant at P&lt;O.l.

�-120-

Table 8. Means and standard deviations
of plasma free fatty acid concentrations 1.f, spring 1973 trial.
Control Deer
Trial Phase
Begin
Conditioning !:./

SD

n

EXEerimental Deer
X
SD

n

0.29 if

0.08

4

0.25 if

0.03

8

0.26 if

0.05

·5

0.33

if,}./

0.10

8

0.13

5

0.43 ]./,if ,:2./

0.20

7

0.70 if

0.28

5

0.73 if

0.20

6

0.27 if

0.15

4

0.20 if

0.06

5

X

2f

3 - Week of
Conditioning !:./

Begin
Starvation !:./

Begin
Refeeding !:./

Ten Days
Post-Feeding !:./

0.30

]j ,if

!f
Micro-equivalents of free fatty acids per ml plasma.

!:)
See Table 7.

3f
Differences between means in rows significant at P&lt;O.l.

4f
Differences between means in columns significant at P&lt;0.05.

Differences between means in columns significant at P~O.l.

�-121-

Table 9.

Means and standard deviations of plasma glucose concentrations

J) , spring 1973 trial.

Trial Phase

Control Deer
X
SD

n

EXEerimental Deer
SD
X

n

III

Begin
Conditioning ];/ 114.3.

28.4

5

131.5

20.0

8

3 !:...I Week of
Conditioning !:...I

129.1 1/

17.5

5

112.8 1/

19.9

8

Begin
Starvation !:...I

101.5

38.9

5

119.3

54.9

8

Begin
Refeeding !:...I

132.8

46.2

5

108.9 III

26.4

6

Ten Days
Post-Feeding !:.../

138.5

40.9

4

113.4

33.9

5

1/
- Mg glucose per 100 m1 plasma.

21

- See Table 7.
3/
- Differences between means in rows significant at P&lt; 0.1

41

- Differences between means in columns significant at P&lt;0.05.

�-122-

Table 10.

Hematocrit means and standard deviations, spring
1973 trial.
Control Deer
X

SD

n

EXEerimental Deer
X
SD

Conditioning l/

52.6

2.9

5

54.6 2:../

2.5

7

Begin
Starvation 1:../

51.0

3.7

5

52.3 2:../

2.5

7

Begin
Refeeding l/

50.2 2:../

3.0

5

50.8 2/

5.1

6

Ten Days
Post-Feeding 1:../

45.5 2:../

0.7

2

43.8 1/

4.9

5

Trial Phase

n

Begin
Conditioning l/
3 l/ Week of

1/

- See Table 7.
2/
- Differences be tween means in columns significant at P&lt; 0.1.
3/
- Differences between means in columns significant at P&lt;0.05.

�-123-

Table 11. Means and standard deviations of total protein concentration
spring 1973 trial.

l/,

Experimental Deer

Control Deer

I'

Trial Phase

X

SD

n

X

SD

n

Begin
21
Conditioning -

7.6

0.5

4

8.0 §j

0.5

8

3 2:...1Week of
Conditioning 2:...1 7.6 11

0.4

5

8.011,2-1

0.5

7

Begin
Starvation 2:...1

7.6

0.6

5

7.7 2./

0.6

7

Begin
Refeeding 2:...1

7.5

0.5

5

7.3 ~./,!l./ 0.4

6

Ten Days
Post-Feeding 2:...1 8.0!J

0.4

2

7.0!!./

5

I;

0.4

11
- G total protein per 100 ml whole blood.

2:...1
See Table 7.

11
Differences between means in rows significant at P&lt;O.1.
!!../
Differences between means in rows significant at P$.0.5.

2./
Differences between means in columns significant at P$.O.1.

2-1
Differences between means in columns significant at P~ 0.05.

�-124-

Pre-Starvation Nutritional Plane and Refeeding Performance
Neither control nor experimental deer experienced refeeding mortality.
Percent weight gain during refeeding for control deer was not significantly
different (P;&gt;O.l) from that for experimental deer (Table 6).
On the tenth day of refeeding mean PUN levels, plasma FFA, plasma glucose,
and hematocrits for experimental and control deer were not significantly
different (P&gt; 0.1). Mean level of plasma total protein on the tenth day
of refeeding for control deer was significantly higher (P~0.05) than that
for experimental deer.
Experimental deer ate significantly more (P~.005) during refeeding than
controls. Intakes of experimental and control deer approximately doubled
by the third day of refeeding. Experimental deer tripled initial intake
during 6-8 days of refeeding; intake of the control deer appeared to increase at a slower rate and never tripled that of the first day.
Blood Energy-Metabolites from Spring 1972 Trial Deer

Starvation--Mean PUN levels for does and fawns differed significantly
(P~0.025) only at 33-36 days of starvation (Table 12). Three of six fawns
starved 33-36 days died; mean PUN for these deer (42.3 mg percent) was significantly higher (P~0.05) than that for surviving fawns (26.5 mg percent).
Mean PUN level for fawns starved 33-36 days was significantly higher
(P&lt; 0.001) than that for non-starved fawns. Mean levels of plasma FFA for
noU:starved does and fawns were not significantly different (p&gt; 0.1).
During starvation mean levels of plasma FFA for does were significantly
higher (P&lt;0.05) than those for fawns (Table 13). Mean level of plasma
FFA for fawns starved 33-36 days was Significantly higher (P~ 0.1) than that
for non-starved fawns. Mean level of plasma glucose for does starved 55-64
days was significantly lower (P~0.05) than that for non-starved does.
Mean levels of plasma glucose for starved fawns and other starved does were
not significantly different (P&gt;O.l) from each other nor from those of respective age-class non-starved deer (Table 14). Mean hematocrit for fawns
and does were not significantly different (P&gt;O.l) (Table 15). Mean hematocrit for does and fawns starved ten days were significantly higher (P&lt;O.l)
than those for respective age-class non-starved deer.
Refeeding--Levels of blood energy-metabolites on the tenth day of refeeding
were compared respectively between: (1) fawns; and (2) does refed Diet III
and Diet IV for each starvation period. There were no significant differences
(P&gt;O.l) in levels of blood energy metabolites between deer refed Diet III
and deer refed Diet IV.
Assuming levels of blood energy metabolites were not affected by diet refed,
values of each blood parameter on the tenth day of refeeding were regressed
on length of starvation prior to refeeding. This was done to determine
whether or not length of starvation prior to refeeding affected levels of
blood energy-metabolites on the tenth day of refeeding. Hematocrit of does
on the tenth day of refeeding declined Significantly (P~0.05) as length of
starvation increased prior to refeeding (Table 16). Further information concerning results and discussion can be obtained from deCalesta (1973).

�-125-

Table 12. Means and standard deviations of plasma urea nitrogen concentrations "J:./ from deer starved 0-64 days, spring 1972 trial.
Does

Days
Starved

X

SD

n

0

17.7

3.0

10

22.3

17
24

X

Fawns
SD

n

13

17.2

3.0

13

6.7

2

17.0

0.0

2

15.0

1.4

,2 -

20.5

4.9

2

17 .5

1.4

2

20.5

0.7

2

33-36

18.0 ];./

5.2

3

34.4 ];./

11.0

6

55-64

20.5

2.8

3

1/
- Mg urea nitrogen per 100 m1 plasma.
];./Differencesbetw~en means significant at P&lt;0.025.

Table 13. Means and standard deviations of plasma free fatty acid concentrations 1:./ from deer starved 0-64 days, spring 1972 ];./trial.
Days
Starved

X

Does
SD

n

X

Fawns
SD

n

0

0.24 1/

0.04

13

0.26 1/ ,!!./

0.04

13

10

0.53 1/

0.24

2

0.34 1/

0.08

2

17

1.32 1/

0.54

2

0.39 1/

0.01

2

24

0.65 1/

0.16

2

0.37 1/

0.16

2

33-36

0.61 1/

0.42

3

0.29 i/

0.06

6

55-64

0.69 1/

0.61

3

!/Micro-eQUiValents of free fatty acids per m1 plasma.
~/Accumu1ated comparisons of doe and fawn mean values during starvation
significant at P~ 0.05.
3/Means of non-starved deer significantly lower (P~O.025)
starved deer.
4/Differences between means significant at P&lt;O.l.

than those of all

�-126-

Table 14. Means and standard deviations of plasma glucose concentrations
1:.../ from deer starved 0-64 days, spring 1972 trial.
Days
Starved

X

Does
SD

n

X

Fawns
SD

n

0

126.5 l.:/

28.9

13

145.1

35.5

13

10

162.5

48.8

2

183.3

22.9

2

17

97.8

30.1

2

106.0

22.9

2

24

118.8

36.4

2

145.5

2.8

2

33-36

101.5

28.2

3

73.3

49.9

6

55-64

89.7 l.:/

41.9

3

l/Mg glucose per 100 m1 plasma.
2/
- Differences between means significant at P&lt;O.Ol.

Table 15. Hematocrit means and standard deviations from deer starved 0-64
days, spring 1972 trial.
Days
Starved

Does
X

.u

Fawns

SD

n

X

SD

n

2.8

13

50.3 1:.../

4.2

13

0

52.1 1:.../

10

56.5 1/

0.7

2

56.0 1/

1.4

2

17

55.0

1.4

2

53.5

5.0

2

24

54.0

4.2

2

53.5

3.5

2

33-36

53.5 1:.../

0.7

2

50.3

6.1

6

55-64

46.3 l.:/

3.1

3

1/
- Differences between means significant at P&lt;O.l.

-

2/
- Differences between means significant at P&lt; 0.01.

�Table 16. Slopes and associated 90 percent confidence intervals for regression of mean level of blood
parameters (at the tenth day of refeeding) on length of starvation prior to refeeding.

PUN 1/

Does
FFA 2/ Glucose

Slope ]/

O.OS

0.00

Confidence
Level

±0.21

±O.OO

PUN 1/

FFA 2/

Fawns
Glucose

-0.19 !!.../

0.00

0.00

0.61

-0.24

±O.lS

±0.42

±0.01

±2.S1

±0.32

Hematocrit

-0.12

±0.7S

Hematocrit

1/
- PUN = plasma urea nitrogen.
2/
- FFA = free fatty acid.

I

I-'

N

-...J
I

l/Change in mean blood parameter value per day of starvation.

4/
- Slope significantly different from zero (P&lt;O.OS).

�-128-

Energy Intake of Starving and Refed Deer
None of the diets fed to starving deer in this study resulted in observable urea toxicity or acute indigestion. During 1973 trials, deer
voluntarily restricted food intake the first 2 days of refeeding following starvation. Of 40 starved and refed deer, only 2 died during refeeding. Refeeding starved deer Diets II, III and IV (Table 17) did not
result in observable toxicity nor in appreciable mortality.

Table 17.

Diets used in study.
Chemical Analysis
Crude
Protein

Fat

CWC II

NFE 2:..1

Diet I

18.0%

3.7%

26.1%

58.4%

Diet II

15.0%

2.0%

42.0%

51.0%

Diet III

14.0%

4.0%

23.2%

57.1%

Diet IV

3.8%

1.3%

42.0%

39.3%

Alfalfa

15.0%

1.7%

33.5%

38.3%

!/cwC
2:..1 NFE =

cell wall contents.
nitrogen free extract.

Blood levels of two liver enzymes (OCT and GOT) were well below those considered indicative of liver damage or malfunction. These results, coupled
with low feeding mort~lity, refute the hypothesis that liver malfunction
during refeeding is an important mortality agent of starved and refed deer.
Spring 1973 Trial
Deer in two differing nutritional strata were produced during conditioning.
Control deer did not lose weight or exhibit elevational PUN (plasma urea
nitrogen) or plasma FFA (free fatty acid). Experimental deer exhibited
significant weight losses and plasma FFA elevations during conditioning,
indicating that fat stores were mobilized to offset inadequate exogenous
energy intake. Experimental deer PUN did not increase during conditioning.
This result may be interpreted as indicating that catabolism of muscle protein was not occurring in experimental deer during conditioning.

�-129-

Weight loss and levels of blood energy metabolites for the first 3 deer
that died were not significantly different from those deer surviving
starvation.
However, levels of blood energy-metabolites
obtained from
one deer dying during starvation strongly suggest that it succumbed
to inadequate energy metabolism. Likely catabolism of muscle protein
did not provide the degree of gluconeogenesis
required to mai~tain
energy balance.
Extremely low plasma glucose level (24 mg percent) of the
deer dying during starvation suggests that hypoglycemia was the primary
mortality agent.
Surviving starved deer had levels of plasma glucose
greater than 73 mg percent.

I.

Control deer and experimental deer surv1v1ng starvation exhibited similar
levels of blood parameters (excepting total plasma protein which is unexplainable), weight changes and lack of mortality after 24 days of
starvation and 10 days of refeeding.
Mean levels of plasma glucose, FFA,
PUN and total proteins on the tenth day of refeeding for both groups of
deer were similar to those at beginning of conditioning.
Mean hematocrits,
however, were significantly lower than during conditioning, starvation and
those of non-starved deer in the spring 1972 trial, indicating that an
unexplicable hyper-hydration
occurred in spring 1973 trial deer.
These results indicate that: (1) deer starved to the point of death can
be successfully refed (assuming experimental deer surviving starvation were
physiologically
equivalent to experiment deer dying during starvation);
(2) experimental deer surviving starvation were catabolizing stored fat
during starvation and metabolically equivalent to control deer during starvation and refeeding, and (3) deer dying during starvation failed to maintain
levels of blood energy-metabolites
required for sustenance.

Spring 1972 Trial
As in the 1973 spring trial, starving does lost weight and exhibited increasing plasma FFA levels and constant PUN levels.
Plasma glucose levels
remained constant until 55-64 days of starvation when they declined lower
than before starvation.
Weight loss was linearly related to length of
starvation, whereas plasma FFA levels appeared to increase rapidly, peaking
at 17 days of starvation, and dropping to a constant level as length of
starvation increased.
Fawns also lost weight but at a significantly higher
rate than does.
Fawn plasma FFA levels were significantly lower than those
of does during starvation.
PUN levels for starved fawns remained at a prestarvation level until 33-36 days of starvation whereupon they increased
sharply and were significantly higher than those for does starved the same
length of time.
Differences in levels of blood energy-metabolites
between fawns dying and
fawns surviving 33-36 days starvation were of particular interest.
Dying
fawns exhibited significantly higher PUN, lower plasma glucose and nonsignificantly lower FFA and higher weight loss than surviving fawns.
Mean
weight loss of fawns dying during starvation was 30 percent.
Higher rate
of weight loss during starvation for fawns suggest that they catabolized
stored fat and muscle protein at higher rates than does.

�-130-

Levels of blood energy-metabolites for fawns on the tenth day of refeeding were similar to non-starved levels, indicating that fawns, as
did does in the 1973 spring trial, normalized their metabolism by the
tenth day of refeeding.
These results indicate that: (1) starving deer can be refed successfully;
(2) does and fawns are not metabolically equivalent during starvation
and refeeding; and (3) deer dying during starvation and refeeding succumbed
to hypoglycemia.

LITERATURE CITED
deCalesta, D. S. 1973. Starving and refeeding mule deer.
Colorado State University, Fort Collins. 109 pp.

PhD Dissertation.

Dixon, W. J., and F. J. Massey, Jr. 1969. Introduction to statistical
analysis. 3rd ed. McGraw-Hill, New York. 638 pp.
Schoonveld, G. G. 1971. Some nutritional and physiological effects of
feeding mule deer high-fiber diets. M.S. Thesis. Colorado State
University, Fort Collins. 95 pp.

Dan L. Baker for:
Julius G. Nagy

�July, 1973

-131-

JOB PROGRESS

State of

COLORADO

Project No.

W-38-R-27

Work Plan No.

14

Personnel:

Deer-Elk

Investigations

Job No.
7
Middle Park Cooperative Deer Study Remote Sensing of Deer Population Parameters

Job Title
Period Covered:

REPORT

March

1, 1972 through February

28, 1973

Richard S. Driscoll and H. Dennison Parker, Jr., of the Rocky
Mountain Forest and Range Experiment Station.

ABSTRACT
Both V/H correction and frequency filtering were required to process experimental
thermal imagery secured over Cedar Ridge in Middle Park, Colorado, for thermal
detection and identification of wild deer. V/H correction is a function of
aircraft speed (V) and altitude above terrain (H) in relation to scanner speed
and resolution.
A V/H aspect ratio of approximately 1:1 was required to
separate the thermal images of animals in close proximity to each other.
This
provided interpretation of imagery in which contiguous scan lines were presented
rather than "stacked" scan lines. Frequency filtering is a process whereby
either certain frequency bands or frequencies above or below established base
levels read in taped thermal imagery are allowed to be processed.
The various
frequencies relate to certain thermal imaged objects in the ground scene.
Highpass frequency filtering at 20,000 hz (hertz) produced the apparent best imagery
of the deer thermal targets.

��-133-

MIDDLE PARK DEER STUDY
REMOTE SENSING OF DEER POPULATION

PARAMETERS

Richard

Parker,

S. Driscoll

and H. Dennison

Jr.

This is the second and final report for Job 7, Remote Sensing of Deer
Population Parameters, Middle Park Cooperative Deer Study, on using
an airborne thermal scanner for mule deer detection.
A thermal scanner
is a device which detects and records thermal (radiant) energy emitted
by an object.
The scanner, after energy detection, transforms this
information into electrical voltage levels proportional to the magnitude
of the energy impinging on it. The data are recorded in two ways: (1) as
photographs of the energy displayed on a cathode-ray tube or (2) on magnetic tape. The previous report summarized results obtained by visually
interpreting the photographic film products produced by the scanner system
on-board the aircraft (Driscoll and Gill 1972). The original photographic
data was "impure" due to variances in on-board film processing and difficulty in establishing the best image contrasts for display on the cathoderay tube. The current report summarizes the findings obtained by processing
more "pure" magnetic taped data.

P.S. OBJECTIVE
Test the capabilities of aerial thermal remote sensing
estimate live deer densities in Middle Park.

DESCRIPTION

techniques

to

OF THE AREAS

Three areas within Middle Park, Colorado were selected to be overflown to
secure thermal imagery.
All were relatively representative of the total
~1iddle Park deer winter range.
They were:

1.

Junction Butte:
The specific test area was on the southwest
slope of Junction Butte.
It included an enclosure, 330- X
610- feet, within which six tame mule deer were placed.
This
area was selected for control purposes where known numbers of
deer were present.

2.

Jensen Creek:
This was an area approximately 2 square miles in
size located along Jensen Creek north of Parshall, Colorado.
The
area had little topographic relief (about 400 feet per mile) and
was characterized by open, slightly rolling terrain with big
sagebrush (Artemisia tridentata) and other low-growing shrubby
species the primary vegetation.

�-134-

3.

Cedar Ridge: This was an area approximately 7 square miles in
size southwest of Parshall, Colorado. It included considerable
topographic relief in relation to the thermal scanner capabilities
subsequently defined. Mean elevation differences within the area
was approxi~ately 800 feet, but changes were rapid; as much as
400 feet within one-quarter mile. Generally, the vegetation consisted of dense pockets of Douglas-fir (Pseudotsuga menziesii var.
glauca) on the north-facing slopes, big sagebrush and open stands
of Rocky Mountain juniper (Juniperus scopulorum) on the south-facing
slopes, and big sagebrush with associated shrubs across the more
gentle terrain.

METHODS AND MATERIALS

Equipment
The thermal scanner used was a Texas Instruments Model FFS-l, owned and
operated by the U.S. Forest Service. The instrument, a modified Texas
Instruments Model RS-7, had been specifically designed for forest fire
detection and mapping, but had characteristics conducive to detection of
large mammals. The characteristics of the scanner were:
Instantaneous Angular Field of View (IFOV):
Total Angular Field of View:
Detectors:

2.0 milliradians

120 degrees

Channel A; indium antimonide
Channel B; mercury-cadmium-telluride

Wavelength Response:

Channel A; 3.0 - 4.1 micrometers
Channel B; 8.5 - 11.0 micrometers

Thermal Sensitivity:
Scanning Speed:
Data Output Mode:

2° C (0.2° C with 20X overscan)

0.005 seconds per scan line
127 mm film strip
1/2 inch magnetic tape

Wilson et ale (1971) provides a more detailed description of the system.
The instrument was flown in a Forest Service Beechcraft Kingair, modified and
equipped for the purpose. Data gathering missions occurred on January 25,
26, and 27, 1972, and at a time to comply with the constraints set forth by
Parker (1972). Basically, he determined that the highest probability of
success for detecting deer in a cold environment using a thermal scanner would
be when there was no direct beam solar radiation, low air temperatures, and a

�-135-

snow background.
Consequently, the missions were flown as early in the
morning as possible, consistent with flying safety at an altitude of approximately 650 feet above terrain.
This altitude was s~lected to be consistent
with the capabilities of the scanner used to theoretically resolve a ground
target the size of deer in the environments they occurred.
Black and white aerial photographs, using Kodak Tri-X Aerographic film,
were secured at the same time the scanner imagery was flown.
This was done
in an attempt to obtain data on the number and location of deer in the
Jensen Creek and Cedar Ridge area to provide positive validation of the
processed scanner data. The photo missions were flown under severe atmospheric conditions; early morning with complete cloud cover and low solar
angle producing low-intensity diffuse light.
Due to these conditions, the
photographs were unusable for which they were flown.
Consequently, we had
to rely on visual sightings of deer from the air to use for control information in the thermal data processing.
There were two visual sightings,
both in the Cedar Ridge area, of a total of 14 deer. This constituted a
very small sample, 2.9 percent, of the estimated number of deer on Cedar
Ridge (487), determined by a drive count 2 days after the mission.
These
sightings were used as references for processing taped data sequences.

DATA PROCESSING
The processing techniques selected were ones which would potentially electronically enhance the thermal scanner data to facilitate tape-to-film conversion of that data for detection of wild mule deer.
Three techniques were
used to determine the system that would provide the best information.
These
included:

V/H Correction
The term V/H is a ratio factor used to determine the intensity at which a
particular scanner should be adjusted to secure data about the terrain over
which it is flown.
This factor expresses the relationship between aircraft
speed (V), aircraft altitude above the terrain (H), the speed of the scanners
rotating mirror systems, and the instantaneous field of view of the scanner.
A v/H aspect ratio of 1:1 says that the recording mechanism of the scanner
system is recording data as quickly as it is produced.
The scanner used was
capable of providing contiguous terrain coverage at the alti~ude flown.
However, the scanner film recording system, limited by the film processer, was
incapable
of moving film at a sufficient speed to record the data as quickly
as it was produced.
The result was severely compressed or "stacked" imagery,
adjacent imager scan lines superimposed on one another, in the direction of
the flight line. This caused a difficult detection problem in the original
imagery.
Therefore, the taped scanner imagery, which contained all information,
was "stretched" to produce a record of the taped video data at an aspect ratio
of approximately 1:1.

�-136-

Video Signal Frequency

Filtering

This technique involves determining the recorded frequency signal of the
object of interest, deer in this case, and electronically filtering all
unwanted information to highlight that object.
The technique has considerable value for differentiating
objects of near-equal temperature
but different size. For example, under conditions of constant ground
spatial resolution and target (deer) orientation with respect to direction
of scanner movement, it should be possible to isolate the targets (deer)
of interest and subdue ,all other information.
A general explanation of how the technique works requires consideration of
how ground targets are recorded in the imagery.
Figure I shows the relationship between target size, radiant temperature which is detected by the scanner,
and the reSUlting taped signal.
Figure lea) depicts typical scene surfaces
found in the Cedar Ridge test area.
In Figure l(b) the video signal produced
by scanning the scene area is shown in which all surfaces have the same radiant
temperature.
In Figure l(c) the same video signal is illustrated except that
the surfaces scanned have different radiant temperatures.
The variation in
video signal pulse width (Figures l(b) and l(c)) is frequency variation caused
by variation in target size seen from the vertical.
Variation in pulse height
(Figure l(c)) is amplitude variation caused by differences in radiant temperatures of the targets seen from the vertical.
Therefore, the objective in
applying the filtering technique was to attempt to discover a particular bandpass frequency or range of frequencies which would optimize the resulting imagery
for detection of deer.
The frequency filtering is accomplished by designing
and inserting into the data processing circuitry certain resistance networks
which "pass" pulses of certain frequencies only and suppresses all others.

Amplitude

Thresholding

As shown in Figure l(c) variation in radiant temperatures of different objects
in the scene causes variation in the scanner electrical signal levels recorded
on the magnetic tape. As a target is scanned, which has a radiant temperature
greater than the radiant temperature of adjacent surfaces, the resulting electrical pulse has greater amplitude than those preceding and following it. This
change is theoretically exploitable by use of electronic hardware which senses
pulses above a given amplitude.
Thresholding implies a sharp dividing line between two adjacent quantities.
Although it is possible to construct a circuit which would provide this type
of discrete division, such equipment requires custom construction for a specific
purpose and was not available for processing the Middle Park taped imagery.
However, an attempt was made to approximate the thresholding effect by selective manipulation of gain and level adjustments on a film recorder used to record
a scene from the taped data displayed on a cathode-ray tube.
The manner in which gain and level adjustments in the thresholding system affect
displayed imagery is fairly straightforward.
An increase in gain causes an increase in both positive and negative amplitude from an established reference source.

�~

SCANNED

VIDEO

SCENE

---------~
""

SIGNAL

RECORDED

LIJ

o

::J

I-

",

-1

""

o,

""

I
•...•.
(Y')

.-l
I

( B) Constant

ERT

(C) Variable

ERT

2
ct

""

--1

""

LIJ

""

tree

deer

I hrub

roc k

&gt;

LIJ

-1

""

"~

-1

«
z
(!)
CI)

shrub

tre e

ro ck

(A)

---ONE

Figure

SCAN

LlNE----

1 - Relationships between a ground scene (A) and the resulting video
signal under conditions of constant ERT (B) and variable ERT (C).

�....

__ ...._"

.-

- . -- '-'---~--~---."-- - . ..
...

".'.'

,.-

•. .",--- "--., ."
,

-_._"_._-_ ..

,

--.-.---:"

.

.

"

- ...•.

-138-

This reference source is a certain voltage level recorded on the tape and
is set at a "zero" level.
Therefore, a change in level adjustment causes
the reference to change and results in a total shift of the data recorded
on the tape. A positive shift in the level causes more of the signal to
be positive with regard to the zero level of the film recorder.
Sincethe
zero level of the recorder is a true zero voltage level and does not change,
a shift in the input signal results in a change of the whole signal with
respect to the film recorder.
Thus, the recorded film or the latent magnetic tape image can be made to appear lighter or darker, a more negative
or positive electrical amplitude, by gain and level adjustments.
The result
is an enhancement of the desired sc ena target, wild deer in this case, f rom
all other information in the scene.

RESULTS AND DISCUSSION

Junction

Butte

All three enhancement and processing techniques were applied to that imagery
secured of the Junction Butte pens where absolute numbers of deer were located.
Imagery was secured by flying the scanner at three altitudes above terrain:
300, 600, and 1,000 feet. None of the processing procedures described were
successful in isolating the six known deer from other terrain targets.
This
was due primarily to environmental conditions immediately prior to and during
the data collection mission.
Parker (1972) had determined that the highest probability of detecting deer in
a cold environment would occur when there was (1) no direct beam solar radiation
(no direct sunlight), (2) air temperature lower than the radiant temperatures
of deer, and (3) a snow background.
These constraints were not entirely met
at the time of the data mission.
For 2 days prior to the mission, unseasonably
warm weather resulted in snow melt which caused patches of bare ground.
This
resulted in a heat-sink into the soil and higher than usual bare soil radiant
temperatures the morning of the mission.
In addition, measured air temperatures
(1° C) were higher, rather than lower, than the deer radiant temperatures (_20 C)
and the relatively high air temperatures resulted in relatively high radiant
temperature (-3.6° C) of shrubs protruding through the patchy snow cover.
Even
though "hot spots" were detected in the imagery, those believed to be deer could
not be absolutely ascertained by the various data processing techniques without
knowledge of deer location determined by ground observers.
The radiant temperature difference between deer and vegetation (1.4° C) was not within the detection capabilities of the scanner under the conditions it was flown.

Cedar Ridge
Two segments of the Cedar Ridge taped imagery were selected to investigate
the data processing techniques previously discussed.
These include those
areas where deer sightings were made from the scanner aircraft during the
data gathering mission.

�-139-

VIR Correction
The previous discussion of the viR ratio function identified that there was
a probable "stacking" of information in the imagery processed at the time
of the data missiop by the scanner processing unit on-board the aircraft.
The stacking resulted in a compression of the data such that required information about the deer in the taped imagery was lost. The laboratory procedures. required decompression of the taped imagery so that an apparent VIR
aspect ratio of 1:1 was accomplished.
An exact viR value to provide an aspect ratio of 1:1 was difficult to determine
because absolute aircraft ground speed or altitude above the ground were not
known. Rowever, the results of the decompression processing are illustrated
in Figure 2. The apparent number of animals at the arrow in Figure 2(a) is
three. This imagery was produc~d at an aspect ratio of approximately 1:0.3,
that is three adjacent scan lines were stacked on one another, and represents
the kind of imagery produced by the scanner processing unit. Increasing the
laboratory processer speed by a factor of 3 produced the imagery shown in
Figure 2(b). The original group of three deer now appears to contain four,
the number accounted for in the aircraft sighting. This represents taped
data processing at an aspect ratio of approximately 1:1, or processing the
imagery for contiguous scan lines.
It should be understood that the correct data processer viR for one area in
the experimental imagery would not be the same for all areas in the imagery
due to the highly yariable terrain relief of Cedar Ridge. The absolute altitude
of the scanner aircraft was not constant; it varied as much as 40 percent as
the pilot attempted to fly as low as possible within the established 650-ft
maximum altitude above terrain. The required control for total processing of
all the experimental imagery would entail segmenting the imagery into units of
equivalent elevation relative to known aircraft flying height. Some of these
segments could be relatively large, such as over relatively smooth terrain, or
relatively small, such as over terrain where elevation changes rapidly.
Video Signal Frequency Filtering
The video signal from the airborne thermal scanner was very complex in terms
of the frequencies contained in the data relative to the sensed energy responses
of the ground scene. This was caused by a number of factors but was primarily
related to the contents of the ground scene resolved within a single digital
resolution element of the scanner as a function of aircraft flying altitude.
The variable topography of Cedar Ridge caused a considerable amount of apparent
artificial variation in both signal level and frequency. It appeared artificial
in that some of the variation was a function of scene energy integration by the
scanner optical system. The integration effect caused all the energy in a
single digital resolution element to be recorded as though it were emanating
from the entire element. In factp however, warmer parts of the element were
radiating disproportionate amounts of the total energy which obscured other
information in the scene. A real source of variation was due to changes in
elevation of the scanner aboveground.
In this case, the energy detected per
unit of scanner detector area varied according to the distance from the source

�-140-

(a)

~)

Fig. 2. The effects of adjusting vIR to decompress the thermal imagery for
deer detection. Note that in (a) there appears to be three deer at the
arrow (VIR:: 1:0.3). At (b), four deer thermal images can be discriminated
at the arrow rather than three (VIR:: 1:1).

�-l41-

to the scanner detector system. For example, if the altitude (H) is halved
(H/2) as the aircraft passes over a hill, energy (E) at the detector varies
from a value E at H to E2 at H/2.
A base level video signal voltage of 300 hz was established for the frequency
filtering processing. The term hz (hertz) is a unit of electrical frequency
with a periodic process equal to one cycle per second. Although the lowest
frequency component of the scanner taped data was 200 hz, the 300 hz base had
to be established to remove distortion at the lower frequency levels.
Three modes of filtering were applied to the data: low-pass which means allowing
those frequencies below a given frequency to be passed and recorded by the processer; high-pass which refers to the passing and recording of frequencies above
a given frequency; and band-pass filtering which means to allow frequencies to
be passed and recorded in a band or range of frequencies with cut-offs both on
the low and high ends.
Low-pass filtering produced negative results. This would be expected since the
low frequencies corresponded to relatively cool surfaces in the scene such as
snow and partially snow-covered shrubs and rocks.
Band-pass filtering indicated that image sharpness and small detail were lost
when frequencies below approximately 12,000 - 15,000 hz were filtered from the
imagery. This should be expected since a frequency pulse, indicating a change
in detected radiant temperature, which exhibits a relatively large change in
amplitude over a short time or space interval, is a high frequency pulse. This
loss of detail is exemplified in Figure 3 in which the deer images at the arrow
are barely discernible and could be easily confused with other objects in the
thermal scene. Since deer images were of primary interest, it was apparent
that high-pass filtering was required to preserve the deer thermal images.
High-pass filtering was tested at frequencies of 15-, 20-, 25-, 35-, 45-, 50-,
and 100-thousand hz. There was an integration of large detail with each high
frequency step increase. This integration or loss of large detail in the
imagery was created as a result of relative large target surfaces scanned in
the area which appeared as wide pulses in the taped data. Thus, the higher
the frequency the narrower the pulse width and, therefore, the information cut
off by the frequency filter is rendered nondescript.
Most of the high-pass filtered imagery up to 45,000 hz improved or maintained
the thermal signal of known groups of deer. Above 50,000 hz, the signals faded
to non-detectable images. High-pass filtering at 20,000 hz appeared to provide the best overall filtering for deer recognition by their thermal image
(Fig. 4). This was a compromise since the apparent deer image size varied
with terrain relief. Over flat terrain, it would be possible to define the
specific frequency range which would discriminate deer images from all other
objects in the scene. Such inmgery would allow more positive characterization
of deer frequency signal in relation to size (amplitude) of the signal.

�-142-

Fig. 3. Image sharpness was lost by band-pass filtering below approximately
12,000 - 15,000 hz (hertz). Deer images at the arrow are difficult to discern
in this 1,000 - 10,000 hz band-pass filtered image.

�-143-

F~g. 4. High-pass filtering at ,approximately 20,000 hz (hertz) provided
the best recognition thermal signature of deer in the Cedar Ridge area.
Thermal signatures of three deer are at the arrow point.

�Amplitude Thresholding
This processing technique did not provide any improvement in the experimental imagery for detection and identification of deer thermal signals.
This was primarily due to lack of access to the equipment required to
experiment with amplitude thresholding. Theoretically, the technique
might be useful for reasons previously explained.
We did simulate amplitude thresholding empirically by increasing the gain
setting in a film recorder to increase image contrasts among the ground
scene targets. Although the simulation improved identification of some
scene objects, it did not improve deer detection in the imagery.

Jensen Creek
The first two image processing techniques, v/H correction and video signal
frequency filtering, were applied to the taped thermal imagery secured
over the Jensen Creek area. No deer thermal signals were detected in this
imagery. Even though an estimated 200 deer were visually sighted from an
aircraft in the area 6 weeks prior to the scanner mission, there apparently
were none there at the time of the data mission. This was confirmed by the
aircraft ground control crew the morning of the mission. No tracks were
observed leading into or out of the area and no animals could be seen from
vantage points.

CONCLUSIONS
Both v/H correction and frequency filtering of experimental thermal imagery
of Cedar Ridge in Middle Park, Colorado improved detection and identification
of deer thermal signals. Simulated amplitude thresholding was of no value.
A v/H ratio value of 3.0 for the data processer was required to secure an
apparent v/H aspect ratio of 1:1. This technique essentially stretched the
magnetic taped data so that scan lines in the processed imagery were contiguous and not stacked as in the original imagery. This technique provided
for detection and identification of the thermal signals of animals in close
proximity.
High-pass frequency filtering to approximately 20,000 hz (hertz) provided
the best frequency filter for detection and identification of deer by their
thermal signals. Filtering for less than this amount resulted in highlighting
ground scene thermal signals that would result in confused discrimination
between deer and other ground features. Higher frequency filtering resulted
in a fading of the deer targets into nondescript images.
At the present, variable terrain like that of Cedar Ridge presents serious
problems for thermal detection of deer. Rapid changes in relief, 400-500
feet in one-quarter mile, requires that the imagery be segmented into units
of equivalent elevation for data processing. This would be required to develop
data processing functions which must consider the critical values of spatial
resolution and thermal sensitivity of the scanner which are directly related
to scanner altitude above ground.

�-145-

LITERATURE CITED
Driscoll, R. S. and R. Bruce Gill. 1972. Middle Park deer study--remote
sensing of deer population parameters. Colo. Div. Wildl. Game Res.
Sect. Fed. Aid Proj. W-38-R-26. Game Res. Rep. July 1972. Part 3:
253-265.
Parker, H. Dennison, Jr. 1972. Airborne infrared detection of deer.
Thesi&amp;, 186 p. Colo. State Univ., Fort Collins.

Ph.D.

Wilson, R. A., S. N. Hirsch, F. H. Madden, and B. J. Losensky. 1971. Airborne infrared forest fire detection system: final report. USDA For.
Servo Res. Pap. INT-93 , 102 p., illus.

Prepared by:

([j
Richard S. Driscoll
Project Leader
Rocky Mountain Forest and Range Experiment Station

��-147-

JOB PROGRESS

State of

~C~O~L~O~RAD~~O~

REPORT

_
Deer-Elk

W-38-R-27

Project No.
Work Plan No.

Job No.

14

Job Title

Middle

Period Covered:

April

Personnel:

July, 1973

Park Deer Study - Experimental

Investigations
8

Harvest Regulations

1, 1972 through March 31, 1973

W.J. Adrian, D.L. Baker, G.L. Brown, L.M. Brown, L.R. Carpenter,
R.D. Clippenger, P.F. Gilbert, R.B. Gill, R. E. Keiss, J. F.
Lipscomb, D. Luce, L.A. Roper, G.G. Schoonveld, J.L. Wassink
and J.L. Wolfe.

ABSTRACT

Population dynamics information resulted in an estimate of 10,087 ± 2,239
(p&lt;.lO) deer in January, 1973. Prior to this date pre-1972 hunt data suggested
the population consisted of 72 bucks per 100 female deer per 88 fawns.
An
attempt to survey all deer hunters resulted in an 81.6 percent check and gave
a known minimum harvest of 288 deer. An estimated maximum was 356. A random
survey of these same permittees gave an estimated harvest of 336 ± 35 (p&lt; .05)
deer. Following the 1972 hunt post season, sex and age classifications were
64 bucks per 100 females per 82 fawns.
Pre-natal fawn ratios for 1973 were 162
fawns per 100 road killed does. These estimates were used in a life equation
to formulate recommendations
for the 1973 deer hunt.
Included in the life
equation model were winter mortality estimates (.7 percent or 50 deer) for
1972. The harvest objective in 1972 was to achieve a m~n~mum harvest of bucks.
The 1973 objective is to harvest approximately 2900 deer.

��-149-

MIDDLE PARK DEER STUDY - EXPERIMENTAL

HARVEST REGULATIONS

Laren A. Roper

P. S. OBJECTIVE
To adjust a mule deer population

to a specified

density

on the winter range.

SEGMENT OBJECTIVES
1.

Estimate deer density,
Middle Park.

structure,

2.

Recommend season regulations and publicize the procedures for hunters
to obtain permits and participate in the experimental hunt.

3.

Measure and estimate
of hunters.

4.

Determine if the hunting public will participate
to achieve adequate harvests utilizing specified
hunts.

5.

Determine if specified
harvest goals.

the. harvest

permits

METHODS

1.

Structure,

deer density and structure.

Productivity

have been detailed

deer productivity

predetermined

AND MATERIALS

Estimate

Estimate

in

on a continuing basis
permits in post season

are capable of achieving

Deer Density,

and materials

and winter mortality

by sex and age class and the success

Estimate

Methods
2.

productivity

and Winter Mortality

previously

by Gill

(1969 and 1971).

and mortality.

To estimate deer productivity, mature road-killed does obtained from January
through May were necropsied to obtain a doe-fawn pre-natal estimate.
In
addition, a helicopter survey classifying fawns, does and mature bucks was
made prior to the 1972 deer hunting season.
Methods

for estimating

winter mortality

have been detailed

by Gill

(1969).

�-150-

Recommending

and Publicizing

Season Recommendations

Recommendations
for the deer hunting season were designed to provide a
recreational hunt during the regular deer season that would in all probability result in an underharvest of deer. To take the balance of deer
needed to provide an adequate harvest a post-season in mid-December was
recommended using specified numbers of permits.
The recommendations were
formulated with Division Wildlife Conservation Officers and the Area
Supervisor.
They were then presented to the Wildlife Commission for approval.
The Middle Park area was marked with informational signs at 14 highway locations to inform hunters they were with'in'the confines of the experimental
harvest area.
Information was also released to the public through the newspaper, radio and television news medias, special talks to local rod and gun
clubs, and a popularized article in the Colorado Outdoors magazine.
Special hunting permits were delivered to license agents within the Middle
Park area' and to Division offices in Denver and 4 regions.
Permits were
checked out to each agent or office so that control of hunters' names could
be maintained.
Regulations enforcing the special permits and establishing the experimental
harvest regulations were set by the Wildlife Commission.
These regulations
were included in the Division's hunting regulation brochure.

Estimate

the Harvest

and Success

of Hunters

Four procedures to obtain an estimate of harvest were used.
The game management section randomly surveys a percentage of all deer hunters in the State.
This estimate has been reported by Gill (1969, 1970, 1971 and 1972) as the
report card check for Summit and Grand counties.
Three additional estimates were:
1) a regression projection from the 9 day
check of hunters at the Idaho Springs check station, 2) an attempt to survey
all Middle Park hunters (100% check), and 3) a randomly selected 20 percent
survey of all Middle Park hunters.
In order to secure a complete listing of
hunters each was required to obtain a special free permit.
Each permit included a detachable report to be completed by the hunter and either mailed
in a self-enclosed envelope or dropped in a box at one of four locations
within Middle Park.
The boxes were identified as self-service check stations
and located at service stations within Granby, Hot Sulphur Springs, Kremmling
and Breckenridge.
Each hunter was identified by a permit number which was placed on an automatic
data processing program.
This program was developed by James Lipscomb (Division
of Wildlife) to automatically print-out: 1) a total permit listing, and 2) a
list of non-responding permittees.
As hunter reports were returned, the permit number was put into the computer program so that names could be removed
from the current list of non-respondents.
Follow-up

letters were designed

for each survey.

�-151-

To obtain an estimate
sampled by telephone.

of kill from non-respondents

Determine

20 individuals

were

If Hunters Will Participate

A subjective analysis of how many hunters
to the special regulations will be used.

participated

Determine If Specified Permits Are
Capable of Achieving Predetermined Harvest

and their reaction

Goals

The population was monitored and the goal for harvest compared
vest actually achieved according to the 100 percent survey.

RESULTS

to the har-

AND DISCUSSION

Density

Estimates

The 1972-73 winter population estimate of deer for Middle Park was 10,087
± 2239 (p&lt;.lO)
deer or 17.14 deer per square mile of winter range.
Table 1
is a summary of deer counted per quadrat and Table 2 is a list of population
estimates by each deer herd sub-unit.
Table 3 is a presentation of all the
information we have in a life equation (R. B. Gill and L. A. Roper, Div.
Wildl., unpub. data, 1973), and Table 4 is a more specific life equation for
January 1972 through January 1973.

Structure

Pre-season

(Sex and Age Composition)

Counts

From October 8-11, 1972 two observers classified deer throughout Middle Park
using a helicopter.
The weather was clear and windy on four flights, and
rainy on two classification counts.
Count conditions were poor with no snow
background.
The best counting success was during 2 flights of rainy weather.
Total deer observed wa~ 273 by one observer and 267 by the other.
Total
classified was 264. Averages for the two observers were 29 mature bucks,
43 young bucks and 88 fawns per 100 mature doe deer (Table 5). The only
differences between the two observers classifying independently were 2 fawns
and 1 mature buck per 100 females.

Post-season

Counts

A total of 3146 deer were classified
observers walking deer concentration
1972.

in 48 man-days by two independent
areas from November 24-December 29, .

�-152-

Classifications were 495 mature bucks, 323 young bucks, 1281 mature females and 1047 fawns.
Ratios wer-e : 39 rna t ure bu cks r lOt) Ct'.mnlcs:2') \','1Jllp,
bucks: 100 females; 82 f awn s :100 females.
Aerial classifications were also obtained during census flights un January
5 and 6, 1973. Comparisons of post-season deer classifications by ground
observers and aerial observations are presented in Table 6.

Productivity

Estimates

Pre-season fawn:doe estimates prior to the 1972 deer season were 88 fawns:
100 mature doe deer (Table 5). Productivity estimates obtained from JanuaryMay 1973 from road:-killed mature does were 162 fawns per 100 mature doe deer.
Table 7 is a comparison of pre-natal fawns per 100 does from 1969-1973.

Mortality:
Over-winter mortality for the 1971-72 winter was completed during this segment and reported by Gill (1972). The estimate was 0.7 percent or 50 deer
(Table 8).
Highway and railroad loss estimates for the period June 1972-May 1973 were
136 and 56, respectively.
Total losses since 1967-68 are listed in Table 8.

Recommending

and Publicizing

Season Recommendations

During the 1972 deer season the public was informed that Middle Park was an
experimental hunting area. To help accomplish this, permits were required
to hunt in Middle Park and a special information letter was given to each
hunter with the permit.
Apparently the hunter information signs, news
releases, special talks and the popular article in Colorado Outdoors (Roper
1972) were sufficient for Wildlife Conservation Officers reported a high
degree of compliance by hunters in obtaining permits.
Recommendations
for the 1973 deer season were based on a life equation model
of the deer herd. Assumptions used to estimate the harvest needed in the
1973 hunt were:
I} a 10 percent over-winter mortality, 2) a 20 percent
wounding loss in 1973, and 3) a reproductive rate of 49.8 percent or 91 fawns
per 100 does. Using these assumptions~ a harvest of 2900 deer will be required to leave 10,000 deer on the winter range following the 1973 deer season.
Recommendations
to the Wildlife Commission were:
1) antlered only unrestricted buck permits and 2000 either sex permits available by a drawing
for the regular deer season; 2) a post-season to take the balance of the
needed harvest provided 2900 deer are not removed from the herd in the regular
season.
Unsuccessful regular season buck only permit holders will be able
to apply for the post-hunt drawing.
It was also recommended that all hunters

�-153-

obtaining special Middle Park permits in the regular season would be restricted to hunting in the experimental area.· To obtain the estimate of
the regular season kill a projected estimate from the 9-day check at the
Idaho Springs check station will be used. These recommendations were
accepted by the Commission for the 1973 deer seasons.

Estimate the Harvest By Sex
and Age Class and Success of Hunters

Harvest

and Hunter Success Estimates

All hunters were required to have a permit to hunt in Middle Park in 1972.
This provided a total listing of 4109 hunters.
Permits were issued by the
Denver office, 4 Regional offices, the research office in Kremmling and all
license agents within Middle Park. To maintain control of the permits a
check-out list of permit numbers was maintained for each office or agency.
Control efforts were only partially successful as our records showed 593
missing permits.
An attempt was made to contact all hunters to obtain the total harvest.
Hunters were asked to voluntarily return a report card attached to their
Middle Park permit.
Four self-service boxes located in Granby, Hot Sulphur
Springs, Kremmling and Breckenridge were used for collection.
These "selfservice check stations" accounted for only 134 reports.
After the hunt,
surveys were sent to all hunters not returning report cards.
Subsequent to
this letter follow-up reminders were mailed at two-week intervals to nonrespondents.
A computer program updated the list of non-respondents.
Final results were 3524 returns with 288 total deer harvested and a hunter
success of. 8.2 percent.
Since 575 hunters (18.4 percent) did not return
their surveys, 20 were selected at random to be contacted by telephone so
that an estimate of kill could be made for non-respondents.
Only 11 of the
20 could be contacted and one of these was successful.
Since the success
ratio of the 11 hunters (9 percent) was close to 8.2 percent, this rate was
used for non-respondents and resulted in an additional 47 deer.
Seventy-nine of the 1562 voluntarily returned reports were not on the computer
file because permit copies were lost or not turned in. This means that some
of the 593 "missing permits" were obtained by hunters.
Assuming these 79
permits were returned at the same rate (38 percent) as the other 1562
volunteer reports, then 208 additional hunters had Middle Park permits.
This revises the estimate to 4317 permit holders and increases the harvest
by 17 (8.2 percent success rate).
There were 52 undeliverable follow-up surveys due to incorrect or incomplete
addresses.
By applying the 8.2 percent success ratio to these the kill is
augmented by 4 more deer. The total estimated kill from the. 100 percent
survey was a known minimum of 288 and a possible maximum of 356 deer. For
use in the life equation a harvest of 356 was used.

�-154-

Since it is expensive and difficult to obtain a 100 percent check of
hunters each year, a 20 percent random sample of the 4109 hunters was
taken to determine if it would be adequate.
An initial survey letter
and two follow-ups resulted in a 79.9 percent return from 842 mailed
surveys.
Only 790 surveys were deliverable due to incorrect addresses
which made the actual sample size 16.37 percent.
The percent return of
deliverable surveys was 85.2 percent and the success was 55 deer out of
673 hunters or 8.2 percent of permit holders.
The estimated harvest was
336 ± 35 (p &lt;.05) (Snedecor 1968).
On the random 20 percent sample respondents were asked if they hunted in
Middle Park. Only 77.27 percent of the permit holders actually hunted
in the area. This gave an estimate of 3336 ± 53.96 (p&lt;.05) hunters
(Snedecor 1968) and a real ·success ratio of 10.07 percent.
The real
success ratio for the 100 percent survey attempt was 10.67 percent.
Since the known minimum harvest was 288 deer and the maximum was estimated
at 356, it is interesting to note the minimum and maximmn values of the
95 percent confidence interval of the random survey (301 and 369, respecti.vely) are within 4.5 percent of these values (Table 9).
In the future it is planned
to estimate the harvest.

to randomly

sample 20 percent

of the permittees

Another method used to estimate the legal take of deer in Middle Park was
a regression estimate based on the 9-day check at the Idaho Springs check
station (Gill 1969).
This estimate for 1972 was 341.
The Game Management survey used to estimate the deer harvest in all herd
management units in Colorado sampled 11.5 percent of Colorado hunters in
1972. This has been reported in previous years by Gill (1969) as the
County report card estimate.
The 1972 harvest for Middle Park using this
method was 554 deer taken by 3405 hunters.
Their average success was 16.3
percent.
Some hunters with Middle Park permits did not hunt in Middle Park
(22.73 percent in our sample).
I am surmising that these hunters may have
hunted in other units and were successful at a higher rate than Middle Park
hunters.
Some may have reported they did hunt in Middle Park on the
Game Management survey while in fact they may not have.
If this were true
the total kill could be inflated.
This hypothesis cannot be substantiated
but it seems reasonable.
Table 10 is a summary of the four methods to estimate
ratio for hunters in Middle Park in 1972.

Sex and Age Structure

the harvest

and success

of the Harvest

A total of 105 bucks were aged. All were classed as either yearlings or
mature animals using the tooth replacement and wear method (Erickson and
Seliger 1969).
Fifty-four of 55 animals field aged as yearlings were also
classed as yearlings by the dental cementum technique (Erickson and Seliger
1969).
One was judged to be a two-year-01d and the other was not aged using

�-155-

the dental cementum method.
Thirty-three mature animals were aged with
dental cementum annuli counts.
Table 11 is a summary of these data and
shows 52.4 percent yearling males and 47.6 percent bucks over 2 years
old.

Determine

If Hunters

Will Participate

on a Continuing

Basis

To attain this objective data will need to be analyzed as the study continues, however after one year of requiring permits it appears that hunters
will cooperate and obtain a permit to hunt in the area.
Out of 4317 permits 3336 (77.27 percent) permit holders actually did hunt in the area.

Determine

If Specified

Permits

Help Obtain

Harvest

Goals

In 1972 the harvest goal was to have a minimum harvest to allow the deer
herd to increase to an estimated 10,000 deer on the winter range.
The buck
only season was apparently successful in minimizing the harvest.
The census
estimate for January 1973 was 10,087.
The 1973 deer season will be designed
to maintain this population level.
This should provide a test of the achievement of harvest goals using specified permits.

LITERATURE

CITED

Gill, R. B. 1969, 1970, 1971 and 1972. Middle Park deer study - population
density and structure - productivity and mortality.
Colo. Div. Game,
Fish and Parks.
Game Res. Div., Fed. Aid Proj. W-38-R.
Game Res.
Rept., July.
Erickson, J. A., and W. G. Seliger.
1969.
Efficient sectioning of incisors
for estimating ages of mule deer.
J. Wildl. Manage.
33(2):384-388.
Roper, L. A. 1972.
Deer permits:
Sept.-Oct.
p. 20-22.

a look at the future.

Snedecor, G. W., and W. G. Cochran.
1968.
State Univ. Press.
Ames.
593 p.

Prepared

bYJ&amp;~Q~R~
Assistant

Wildlife

Researcher

Statistical

Colo. Outdoors.

methods.

The Iowa

�jIII=

Table 1. Numbers of deer counted per quadrat within eight sampling strata, Middle Park, Colorado, 1968-1973.
Muddy Creek High Density
1968 ·1969 1970

1972

1973

o
12

o
o

o
o

4

o

2

o
o

o

o

o
o
o
o
o

8

9

9

64

6

34

32

18

9

o
o
o
o
o

124

99

20

28

2

o
o
o

58
30

39
66

o
o
o
o
o

6

68

2

Muddy Creek Low Density

1971

1969

56

o

o

0

24

132

Ne

0

46

2

o

5

46

42

22

35

42

31

28

5

o
o

20

12

12

27

o

o
o

o
o

1

22

5

NC

37

1

o
o
o
o
o
o
o

6

6

t ..442

478

175

Y •. 260 23.9

8.8

1

1

o

o
o
o

o
3
3

27

3

136

158

480

6.8'

7.9 24.0

0

o

.0

NC

~5

Ne

o

o
o
o
o
o
o

60

87

o

0

11

NC
44
NC

o
o
o
o

30.

40

52

o

'0

f:

1

O.

8

o

1973

o

o

32

1972

0

1

97

1971

o·

o
o

Blue River High Density

1970

6

NC 1/

NC
00
NC

39

2

1968

o
o

o

o
o
o
o
o
o

11

o

2

o
o
o
o
o
o
o
o
o

1968

1969

1970

1971

14
25

19
0

14
0

Blue River Low Density

1972

1973

47

9

o

25

0

o

10

o

63
50

o

145

30

37

88

77

o

82
41

156

32

35

34

15

9

o
o

19

29

o
o

30

28

o
o

o
o

o
o

o
o

o
o
o
o

4

20

18

9

4

3

o

134

102

20

34

15

78

31

36

26

20

49

15

83

24

22

74

o

2

43

29

o

o

o

3

50

104

52

102

6

28

17

600

565

465

0.7

0.5
47.1

38.8

430
35.8

..

339

I

41.8 28.3

637

107

118

114

48

32

50

53.2

11.9

13.1

12.7

5.3

3.6

5.6

.,

,
.

I

2

501

.I
(I'

,

'

"

/

•

....
\J1

'"

----------------------------------------------------------------------------------------------------------------------------------------------------.. ~

4

o
o

54
7

49

0.0

,0

o

31

58

3.5

o

o

24

124

1.8

o
13

36

64

9.3

o
17

76

67

6

8
10

1973

6

136

8

4

1972

o

105

o

1971

o
o

71

42

1970

6
19

010

o
o
o

21

1969

o
o
o

o

56

1968

�..•
•

,;'

I'

,

.

~
~
,~

I.

I

... ,..

Table 1. Numbers of deer counted per quadrat within eight sampling strata, Middle Park, Colorado, 1968-1971 (continued).
Wms. Fork River High Densitl
1968 1969 1970 1971 1972 1973
0
2

40
0

0
0

26

25
21

58

73

45
42

4

4

0

Wms. Fork River Low Densitl
1968 1969 1970 1971 1972 1973

7
2

6
23

0
0

1
0

3
0

2

0
0

13
0

1
0

11
0

0

0
0

0
0

0

0

0
0
0

'0

5
41

11
60

0
8

3
24

21
40

0
35

0
39

0
51

0
0

2

0

0
0

70

2
70

0

0

50

76

42

43

20

55

65
0

44
15

90
7

36
14

40
27

61
31

9
0
26

36

8

21

21

31

0
6

0
29

10
7

0
12

21
33

253 262

298

139

219

365

11

29

13

25.3 26.2 29.8 13.9 21.9 36.5

1.8

4.1

1.9

0
0

0
6

0
0

0
0

0
4

0
0

45
54

14
0

3
0

1
0

0
0

0

31

52
71

0

0

0

0

0

0

0

0

GranbI'Low Densitl
1968 1969 1970 1971 1972 1973

43
25

9
0

3

Troublesome Cr. High Densitl
1968 1969 1970 1971 1972 1973

0

z
0
rt

C"l
0

0

§
rt
~
~

0
0
NC

0
0

•..
I

t. - 105

131

99

20

Y -21.0

26.2 9.2 18.4 26.4 19.8

4.0

46

92

yst
Sy

1.1
NC - No count.

132

-

1

4

0

0

0.6 0.2

0.8

0.0

0.0

3

V1

--.j
I

1968

1969

1970

1971

1972

1973

20.75

15.52

12.29

9.75

9.24

17.14

3.37

2.02

1.92

1.81

1.64

2.29

2

14

0.3 2.0

�Table 2.

Deer population estimates by sub-units in Middle Park, 1968-1973.
Muddy
Creek

Year

Blue
River

Williams Fork
River

Troublesome
Creek

Granby

Total

1968

4,101

4,074

843

1,622

No Count

1969

10,640

2,994

3,531

774

1,679

134

1970

9,112

1,429

3,291

271

1,910

306

7,207

794

3,335

569

889

137

5,724

1972

1,002

2,259

742

1,404

22

1973

5,429

2,856

4,186

556

2,340

149

10,987

1971

Table 3.

Life equation for deer in Middle Park, winter 1967-68 through 1972-73.

I

t-'

V1
00
I

PreFawn
Pop.

Fawn
ReEroduction
%
No.

PreHunt
Pop.

Harvest
%
No.

8,934

42.65

3,810

12,744

23

639

8,473

53.48

4,531

13,004

15.8

1,135

6,070

45.35

2,753

5,719

8.6

489

5,230

47.36

1971-72

5,429

•7

50

7,038

1972-73

10,087

6.4

647

*

Year

Jan. Pop.
Est.

Winter Loss
%
No.

1967-68

10,640

16.0

1,706

1968-69

9,112

7.0

1969-70

7,205

1970-71

*Data not available this segment.

Wounding
Loss (20%
Harvest)

Other
Known
Losses

2,909

582

51

4,503

901

8,823

28

2,436

487

2,477

7,707

7

516

103

-

52.22

3,675

10,713

3

356

71

*

*

*

*

*

*

Post-hunt POE.
Census
Calculated
Est.
9,253

9,112

7,600

7,206

5,900

5,730

7,088

5,429

101

10,541

10,987

*

*

*

�Table 4.

Life equation for deer in Middle Park from January 1972 through January 1973.

PostSeason
Pop.
Bucks

Harvest

Wounding
Loss
(20% of
Harvest)

PostSeason
Pop.

27.9%
2,909

100.0%
356

100.0%
71

24.9%
2,873

38.5%
4,129

0

0

39.8%
4,129

18.9%
2,021

0

0

19.4%
2,021

Winter
Mort.

PreFawn
Pop.

New
Fawns
89:100

PreSeason
Pop.

24.8%
1,758

27.0%
14

40.0%yr1g.
2,909
--

Does

45.0%
3,190

28.7%
14

23.1%yr1g.
-4,129

B. Fawns

16.6%
1,177

24.4%
12

--

55.0%
2,021

Other
Known
Losses*

Calculated
Pop. Est.

1

t-'

V1

~

-I

D. Fawns

13.6%
963

19.9%
10

--

45.0%
1,654

15.4%
1,654

0

0

15.9%
1,654

Total

100.0%
7,088

100.0%
50

63.1%
7,038

100.0%
3,675

100.0%
10,713

100.0%
356

100.0%
71

100.0%
10,642

1972-73
Census
*
Highway mortality

101

10,541
10,087

= 48; railroad mortality = 21; miscellaneous

mortality

= 32.

�-160-

Table 5.

Pre-season deer classification comparisons
from 1968-1972.

Year

Bucks

Does

Fawns

Total

Bucks:
100 does

Fawns:
100 does

1969

195

451

419

1,065

1970

43

152

93

361

258

771

1971

42

20

72

41

30

91

1972

49

73

71

102

89

264

72

88

Table 6.

Year
1968

1969

1970

1971

1972

Comparison of aerial and ground winter
classifications of
deer in Middle Park, 1968-1973.

Bucks

He1icoEter Surve~s
Does
Fawns

Ground Surve~s
Does
Fawns

Total

Bucks

316

558
501
1,375
57 Bucks
100 Does
90 Fawns : 100 Does

147

396
363
906
37 Bucks
100 Does
92 Fawns: 100 Does

261

490
418
1,169
53 Bucks
100 Does
85 Fawns : 100 Does

319

582
505
1,406
55 Bucks
100 Does
87 Fawns : 100 Does

211

471
45 Bucks
77 Fawns

104

231
178
513
45 Bucks
100 Does
77 Fawns : 100 Does

356

783
321
1,460
45 Bucks
100 Does
41 Fawns : 100 Does

818

1,281
1,047
3,146
64 Bucks
100 Does
82 Fawns : 100 Does

363
1,045
100 Does
100 Does

------- No Counts ------------

362

693
540
1,595
52 Bucks
100 Does
78 Fawns : 100 Does

Total

Table 7. Comparisons of fetuses per 100 mature female deer in Middle Park,
1969-1973, in road killed does examined from January-May.
Year
1969
1970
1971
1972
1973

Fetuses per 100 Females
193
164
161
187
162

100
100
100
100
100

�-161Table 8. Summary of over-winter mortality, highway and railroad kills,
and miscellaneous losses from 1967-1973 in Middle Park.
Winter

Winter Loss
Estimate

Highway
Loss

Railroad
Loss

Miscellaneous
Losses

Total

1967-68
1968-69
1969-70

1,706
639
1,135

54
77
67

132
56
38

0
51
40

1,892
823
1,280

1970-71
1971-72
1972-73

489
50
647

57
48
136

9
21
56

49
32
0

604
151
839

Table 9. Comparison of minimum and maximum estimates of harvest using a 100
percent survey attempt and a 16 percent random survey, Middle Park, 1972.

100 Percent Survey

16 Percent
Random Survey

Diff.

% Diff.

288
356

301*
369*

+13
+13

+4.5
+3.7

Minimum Estimate
Maximum Estimate

* p(301&lt;Mean&lt;

369)

.95, Snedecor 1968.

Table 10. A comparison of four methods estimating total deer harvest, number
of hunters and hunter success in Middle Park in 1972.

Number

Harvest
Estimate

Percent
Success

Permits Issued (known)
Estimate of Total Permits
Issued

4,109

356*

8.7

4,317

356

8.2

Estimate of Total Hunters

3,336**

356

10.7

Min.
Harvest Estimates
1. 100% Survey (81.6% actual)
2. 20% Survey (16.4% actual)
3. Regression Estimate
4. Game Mgmt. Survey Estimate
(No. Hunters = 3,405)

288
301

Mean

336
341
554

Max.

% Success

356
369

8.6-10.7
9.0-11.1
10.2
16.3

*This estimate will be used for projections in the life equation for deer in
Middle Park.

** This estimate is derived from the 20% survey where 77.27% of the permjt holders
reported they actually hunted in Middle Park in 1972. The balance of the permittees
either reported hunting in other areas or did not hunt.

�-162-

Table 11. Summary of the age structure of male deer in the harvest,
Middle Park, 1972.
Age
Class

Number

1

55

2

13

No. Aged By
Dental Cementum

52.4

54

12.4

13

3

10

9.5

10

4

4

3.8

4

5

3

2.9

3

6

1

.9

1

.9

1

.0

o

.0

o

.9

1

.0

o

16.3

o

100.0

87

7

1

8

0

9

0

10

1

11

0

Unclassified Mature

17

Total

Percent

105

�July, 1973

-163-

JOB PROGRESS REPORT

State of

~CO~LO~RA~D~O~

Project No.

W-38-R-27

Work Plan No.

15

Job Title:

Deer-Elk Investigations
Job No.

la

Monitor Potentially Critical Deer-Vehicle Accident Areas Statewide

Period Covered:
Personnel:

_

April 1, 1972 through March 31, 1973.

Don Crane, Larry L. Green, William R. Heicher, Marion Lowery,
RichardF. McDonald (and all Wildlife Conservation Officers in
his area), J. Kris Moser, John W. Seidel, Thomas M. Pojar, Dale
F. Reed and Thomas N. Woodard.

ABSTRACT

The number and location of deer-vehicle accidents were monitored in five general
areas of Colorado. Recommendations for installation of devices to alleviate
deer-vehicle accidents were submitted to the Division of Highways when warranted.

�-164-

RECOMMENDATIONS

1.

Recommendations were developed from this study and made to the
Colorado Division of Highways for erection of the 9-foot fences
and attendant structures referred to in this report.

�-165-

MONITOR POTENTIALLY CRITICAL
DEER-VEHICLE ACCIDENT AREAS STATEWIDE
Thomas N. Woodard

P. S. OBJECTIVE
To establish statewide, the location and number of deer-vehicle accidents
in areas appropriate to the evaluation of devices.

SEGMENT OBJECTIVES
1.

Examine potentially critical deer-vehicle accident areas reported by
management and as determined by Job 1.

2.

As appropriate, record all deer found killed in selected highway area.

3.

As appropriate, estimate deer densities in each selected highway area.

4.

Measure deer activity along the roadside or in the median of each
selected highway area when feasible.

5.

Sunnnarize the Highway Department's traffic volume data for each selected
highway area as needed.
METHODS AND MATERIALS

Methods and materials have been described by Pojar (1972).
DESCRIPTION OF AREAS

Highway 6-24 (1-70) Rifle West
See Pojar (1972). Preliminary planning and design of the interstate highway which will replace Highway 6-24 is completed but construction has not
begun. A total of 8 underpasses and 1 overpass with associated 8-foot
fencing has been incorporated in the original highway design.

Highway 1-70 Eagle East
Pojar (1972) described this area. The 8.5 miles of 1-70 from Eagle east
constituting this area was opened to tnaffic October 5, 1972. A 4.75 mile
8-faat fence will be installed along the north right-of-way during summer,
1973.

�-167-

Highway 1-70 Avon-Wolcott
Pojar (1972) described this area. Two 2.25 mile 8-foot fences were installed adjacent to the highway in this area in 1972.

Durango Area
Pojar (1972) described this area.

Highway 82 - Glenwood-Basalt
Myers (1969) described this area.

RESULTS AND DISCUSSION

Highway 6-24 (1-70) Rifle West
A total of 92 vehicle-killed deer were documented on 18 miles of highway
in the Rifle-West study area during this segment. Eighty-three percent
of these were killed during December, January, and February. Seventy
percent were killed along a 7 mile length beginning approximately 5 miles
west of Rifle.

Highway (1-70) Eagle East
A total of 165 vehicle-killed deer were documented on 8.5 miles of highway
in the Eagle East study area from October 5, 1972, to March 31, 1973. Deer
crossed nightly to the south side to feed on irrigated hayland and back to
the north for cover in the pinyon-juniper communities common on that side.
Deer kill declined in January when snow depth precluded use of the hayland
(Fig. 1).
The majority of deer were killed along the section of highway beginning
1/2 mile east of Eagle Interchange and proceeding east 4.5 miles (Fig. 2).
The 4.75 mile 8-foot fence will be installed along this section of highway.

Highway 1-70 Avon-Wolcott
A total of 131 vehicle-killed deer were documented on 14 miles of highway
in the Avon-Wolcott study area during this segment. Deer kill was reduced
on the portions of highway opposite two sections of 8-foot fence (Woodard
1973) .
Deer kill increased along the section of highway from the Eagle River Bridge,
2 miles east of Wolcott, to approximately lA5 miles east of that point. Two
deer were killed in this area from January through March in 1972 and 13 were
killed in 1973 durt ng the same period. A 14 x 14 foot equipment underpass
is situated near the center of this area. It could be used, with associated
8-foot fencing, as a deer underpass.

�..-54

5 )-

..-44

a::

4 1-

w
w

0
LL

3

-

2

-

0

,.22.

..-28

a::

w

I

00
\0

.-I
I

CD
~

=&gt;

2

•

.--

7

6

OCT

rr:

NOV.

DEC.

JAN.

FEB.

MAR.

MONTH
Fig. 1. The total number of deer killed on I-70 from 1.15 miles west of Wolcott Interchange to
Eagle Interchange (Oct. 5, 1972 - March 31, 1973).

�8- FOOT FENCE TO
BE INSTALLED IN 1973

20
0:
w
lJ..J

0

I.J...

0
I
0"1
\0
.-4
I

·

15 •

•
•
•

a::
lJ..J

·
10 •

CD

..

::E
::::&gt;

z

-

·

~

--

-

I-

.. ~

••••

5 •

-

·•
..

-

~

~

•

~

~

I

5

~

-, 10

-

-I-

"'"

••
15

QUARTER - MILE

20

25

!O

35

SECTIONS

Fig. 2. The total number of deer killed on 1-70 by quarter-mile section from Oct. 5, 1972 - ~arch 31,
1973 from 1.15 miles west of Wolcott Interchange (section 1) to Eagle Interchange (section 35).

�-170-

Durango Area
A total of 356 deer and 52 elk was killed by vehicles on the 118 miles
of highway in the Durango study area during this segment. This total
constitutes a decrease of 90 deer and an increase of 23 elk from the
previous year period. A deer underpass with associated 8-foot fencing
has been incorporated into a highway construction project on Highway
160 west of Durango between Cherry Creek and Mancos. Twelve deer were
killed in the underpass area during this segment.

Highway 82 - Glenwood-Basalt
One hundred sixty~five vehicle-killed deer were documented on 18 miles
of highway in the Glenwood-Basalt study area during this segment.
Seventy-five percent were killed from January through March. March,
with 51 deer, had the highest monthly kill.
Traffic volume has increased annually by approximately 12 percent since
1968. The 1972 traffic volume was 1.73 million vehicles.
A highway construction project which includes 8-foot fencing in the
Carbondale East area (Pojar 1972), 9 miles southeast of Glenwood, will
begin during the next segment. A 1.1 mile 8-foot fence in the Diamond-S
area, 7 miles southeast of Glenwood is currently being evaluated (Woodard
1973).

LITERATURE CITED
Myers, Gary T. 1969. An investigation of deer-auto accidents. Colo.
Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. W-38-R-23.
Game Res. Rep., July, Part 2. pp. 147-178.
Pojar, Thomas M. 1972. Monitor potentially critical deer-vehicle
accident areas statewide. Colo. Div. Wildl. Game Res. Div. Fed.
Aid Proj".W-38-R-26. Game Res. Rep., July, Part 3. pp. 305-310.
Woodard, Thomas N. 1973. An evaluation of deer-proof fence length
required to prevent deer movements on or across high speed highways.
Colo. Div. Wildl. Game Res. Div. Fed. Aid Proj. W-38-R-27. Game
Res. Rep. In progress.

./

"L.

)

r

I .

Prepared by -1 {-n,:/;::.
(J/J.et!llMj!
Thomas N. Woodard
Senior Research Technician

�-171-

July, 1973

JOB FINAL REPORT

State of

COLORADO
--------~~~~~------------

Project No.

W-38-R-27

Work Plan No.

15

Job Title:

Job No.

3

Investigation of One-Way Deer Structures

Period Covered:
Personnel:

Deer-Elk Investigations

April 1, 1972 through March 31, 1973

Thomas N. Woodard, Thomas M. Pojar and Dale F. Reed.

ABSTRACT

Gates designed to allow passage of mule deer (Odocoileus hemionus hemionus)
in only one direction were tested under controlled and field conditions.
Two gate types had significantly (P~0.02) different frequencies of use under
controlled conditions. Eight gates of type deemed most effective were installed
in 8-foot fences adjacent to Interstate Highway 70 near Vail, Colorado. A
total of 406 passages were recorded through these gates during 1970-71 and over
96.2 percent of these were in the one-way direction for which the gate was
designed. Based on track counts, we estimate five of these gates permitted
about 145 deer to escape the immediate highway right-of-way. Two one-way
fences did not meet the one-way criteria. More negative crossings occurred
than positive under controlled conditions. They were rejected for field tests.

�-172-

RECOMMENDATIONS

When deer-proof fencing adjacent to high-speed highways is used to reduce
or prevent deer-vehicle accidents, one-way structures should be considered
for installation.
Deer that move around the ends of the fence, or from
the side of the road not fenced, will be able to pass through a one-way
gate rather than be trapped on the highway and likely killed.

�-173-

INVESTIGATION

OF

DEE.I{ :-;T!\UCTU KES

ONE-\.JAY

Dale F. Reed

P. S. OBJECTIVE
Evaluate the effectiveness of three types of one-way gates and two fence
structures to permit one-way movement of mule deer through existing deerproof fences at Little Hills Experiment Station and elsewhere.

METHODS

AND MATERIALS

One-Way

See manuscript

in preparation

Gates

described

One-Way

below.

Fences

Two one-way fences, similar to the overhanging and slanting fences of Long-.
hurst and Jones (1962) were constructed.
Each was made with variable
vertical sections and with either a variable overhanging or slanting section.
Table 1 lists the characteristics and adjustable features of both fences.
The variable sections were accomplished by use of telescoping square steel
tubing (Unistrut Western, Inc., 601 S. Jason St., Denver, Colorado).

Table 1.

One-Way

Characteristics

Fence

and adjustable

Max. -Min.
Vertical (In. )

features

of one-way

fences I and II.

Max. -Min.
Depth (In. )

Variable
Angle Limits
(from Vertical)

Operational
Principle

I

66-42

64-34

450- 900

Ove rh anging

II

72-48

90-48

1150-1300

Slanting

The number of deer crossing one-way fences I and II (overhanging and
slanting, respectively) was to be counted each day in a manner similar to
that of the one-way gates.
Only preliminary tests were conducted.
Procedures for further testing were not developed.

�-174-

RESULTS AND DISCUSSION
One-Way
Manuscript

Gates

in Pre~ration

Reed, Dale F. and Thomas M. Pojar.
Wildlife Mgmt.

One-Way

Use of one-way

gates by mule deer.

J.

Fences

Preliminary tests of the one-way fences were considered unsuccessful.
The
few positive crossings that occurred with the variable height posts in their
lowest positions were outnumbered by negative crossings (Table 2); consequently, the one-way fences did not meet the previously established oneway criteria.
Structures not meeting this criteria were to be modified or
rejected.
In this case, modifications did not appear feasible and the
structures were rejected for further testing.

Longhurst
William H., Milton
and Maynard W. Cummings.
Circular 514. pp. 7-13.
9

Prepared by

~dL~

B. Jones, Ralph R. Parks, Loren W. Neubauer,
1962. Fences for controlling deer damage.

Dale F. Reed
Wildlife Researcher

�-175-

Table 2. Passages and days elapsed during
preliminary tests with one-way Fences
I and II.

Days Elapsed
Without
Pos. Passage

Days Elapsed
Without
Neg. Passage

Ht. of
Fence
(In.)

Length of
Angle
(In.)

Group or Deer

Crossings
Pos. Neg.

Group A (5 deer)

0

5

46

70

Group B (2 deer)

0

7

46

70

Buck (2 pt. from
Group A)

-1

7

46

70

Buck (3 p t , )

0

3

34

65

Buck (4 Pt.)
Doe
Doe (from Group M)

1

0

1

4

34

65

1

-1

4

0

34

65

0

-1

3

o

34

65

4

34

65

34

65

Fawn
Fawn

Total

0
0

2

4

-3

24

18

��July, 1973

-177-

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~-----------

Project No.

W-38-R-27

Work Plan No.

15

Job Title

Job No.

6

Deer Underpass Evaluation

Period Covered:
Personnel:

Deer-Elk Investigations

April 1, 1972 through March 31, 1973

Thomas M. Pojar, Thomas N. Woodard and Dale F. Reed.

ABSTRACT

The use of a concrete box underpass 10 feet by 10 feet and 100 feet long that
permits deer to move under Interstate 70 to and from their summer range was
evaluated. The structure, located 4.3 miles west of Vail, utilizes a well
established deer migration trail and a natural drainage referred to as Mud
Springs Gulch. According to the number of passages recorded by the electrooptical detection system and video surveillance, deer increased their use
of the structure by 40 percent from 1971 to 1972. The greatest use of the
underpass occurred during June and November with maximums of 28 and 39 net
passages per day respectively. An index of entrance versus exit activity
indicated a behavioral reluctance on the part of deer to use the underpass.

��-179-

DEER UNDERPASS EVALUATION
Dale F. Reed

P. S. OBJECTIVE
Determine if deer migrating from winter range on one side of Interstate
70 to summer range on the opposite side utilize an underpass constructed
in the area.

SEGMENT OBJECTIVE
Measure deer use of the underpass.
METHODS AND MATERIALS
Methods and materials have been described by Reed (1971). In addition,
a video time-lapse surveillance system was used during the 1972 spring
and fall migration periods. This system and its application has been
described separately (Reed, Pojar, and Woodard 1973).
RESULTS AND DISCUSSION
Spring and Summer Use of the Underpass
It was estimated that 286 deer moved south through the underpass during
the spring and summer migration of 1972. The first passage, south toward
higher summer range, occurred on May 18. The greatest use occurred during
June with a maximum of 28 deer passing through during one night. Use
slackened during July to a greater extent than it had during the two previous spring and summer migrations (Fig. 1). Compared with 1971, the
number of passages during the spring and summer of 1972 increased 26.0 percent.
This may have been caused by an increase in the population, a shift in the
migration pattern, or by estimating the number of net passages with the video
system instead of the electro-optical system (Myers 1969).
Indices of activity were determined by dividing counter readings at the
entrance to the underpass by those at the exit (Fig. 2). These indices
would equal 1.0, if each deer that came to the entrance went through and
was counted once at the entrance and once at the exit. The greater the index,
the greater the number of counts at the entrance as compared with the exit,
and the greater the amount of time and energy expended there by deer. The
indices, indicators of apparent reluctance on the part of deer to use the
underpass, decreased during 1972 (Fig. 2). Observations, made via the video
system, confirm th~hypothesis that most deer spend time at the entrance before, and possibly without, passing through the structure.
Results including the number of deer approaches, entrances, exits and passages
recorded by the video system has been described separately (Reed, Pojar, and
Woodard 1973).

�(J)

~

l&gt;

o

16 -31

JUl

. '"

I

,

.

"

....
....•...

-- . "'2:..___

I

_--

"""'-

b
.....,

(JI

-&gt;

~/

.•...•........•.

...•

\

--.
-- ---\
-~
~.
- - ~-:::.-=--.::..-.=::"- _.i'
-- -.
.---- ~~,,.----!B

/

,

" "

I

,,,
.,
./

/

.

,'." "
,.
,

,.•

. _--

-- -+-.
I

I

....

Fig. 1. The mean number of passages per day for each 2-week period,
spring-summer and fall migrations 1970, 1971, and 1972.

DEC

\
DEC

/.

\"~

I

,I

NOV 16-

NOV

OCT

OCT

SEP 16-

SEP I

AUG

AUG

I

JUL

JUN

JUN I

MEAN PASSAGES

-180-

�40 .

.

-"

"

&gt;
&gt;
lo

1970
1911
1972

-------

~

"

I ,
I ,

I

,' ''
I

'

,,
,,
,,

I

~

,
I
I

«

,,

020

,
,,,

,,
,,
,

I

LL

I
I

. ,

I

X

W

,"

c

""",

,•

Z

«
W

:'\"' .

Z 10

•

,/

•

-

0

I

I

I

MAY

-

0

- - \'I

It)

N

CoSiJ

__

.'

CD

It)

\

CD

\

\

N
I

N

,,
,,
,,

,"''' ,
,.,...,.'"
",

"- "

""

•

"

~

•

~

JUNE

TIME

.

,,

&amp;I)

-- -. I

," "

//\
/

.

It)

CD

__

.

1"~"\

,

,,----- ...
"",

:E

,

-• JULY

10

•

0
CD

It)

0

I

~

CD

PERIOD

Fig. 2. Mean indices of activity at Vail underpass during the spring and summer migrations,
1971, and 1972.

1970,

1$

�-182-

Fall Use of the Underpass
It was estimated that 546 deer moved north through the underpass during
the fall migration of 1972. The first passage occurred on October 1
and the last on December 22. The greatest use occurred during October
and November with a mean of 16.9 deer passages per day during the first
15 days of November (Fig. 1). A maximum of 39 deer passed through the
underpass during one night.
The fall use increased by 90.9 percent over spring-summer use. Herd
increment, a shift in overall migration patterns, and low hunter harvest
are probably responsible for such an increase.
The number of passages during the fall of 1972 increased 48.8 percent
over a comparable period in 1971. As with the increase in the spring
migration between 1971 and 1972, this may also have been caused by
estimating the number of passages with the video system instead of with
the electro-optical system.
Variation in seasonal use of the underpass occurring between fall migrations
(Fig. 1) may have been due to weather conditions, primarily the timeliness
and severity of snowfalls.
Approximately 21 rains or snowfalls occurred during October and November
of the 1972 fall migration period. The mean temperatures (OF) when deer
passages occurred during October and November were 30.9 and 21.8, repectively.
Two hundred twenty-eight deer (Table 1) were recorded moving north through
the structure during, or within 24 hours following, precipitation. Additional weather data will be collected and analyzed during the next segment.
The indices of activity for the 1971 and 1972 fall migrations were much
lower than the spring-summer indices previously discussed (Fig. 3).
LITERATURE CITED
Myers, Gary T. 1969. Development of an electronic deer counting device.
p. 235-247. In Game Research Report, July Part 2. Colo. Game, Fish
and Parks Div. Federal Aid.
Reed, Dale F. 1971. Deer underpass evaluation. p. 341-351. In Game
Research Report, July Part 3. Colo. Game, Fish and Parks Div. Federal
Aid.
Reed, D. F., T. M. Pojar, and T. N. Woodard. 1973. A video- time-lapse
system for wildlife surveillance. Colo. Game, Fish and Parks Div.,
Game Inform. Leafl. No. 94. 3 p.

Prepared by

c:o tti&amp;yczl}
Dale F. Reed
Wildlife Researcher

�-183-

Table 1. Passages through the underpass during October, November,
December, 1972 in relation to precipitation and weather conditions.
Oct.

Nov.

Following Snow l/

140

88

228

Not Following Snow:!:../

85

166

251

Unknown

43

17

Dec.2/

Total

In Relation to Snow:

67

7

Total

546

In Relation to Weather
Condi tions':
Cloudy and Snowing

94

35

129

Cloudy and Partly Cloudy

65

140

205

Clear

25

29

54

Clear and Moonlight

38

12

50

Unknown

46

55

Total

7

108

546

1/
- During precipitation or 24 hours following cessation.
2/
- Not during or within 24 hours following cessation of precipitation.
3/
- Data through December 22.

�i~/\

~ 51
&gt;
-

---1972

•

-

I-

1971

~

u,

0

e 1
3

&lt;,

,

•

z 2

",'.~ ./

'.'

.

",.

&lt;t

W

II- •..•

\
\

.

It

I

\

:E

10

0

10

I

I

I

- - CD -

0

N
I

10

N
I

-

If)

It)

I

I

CD
N
OCTOBER
CD

.N

- 0

U')

0

It)

I

I

N

N

t\I
NOVEMBER

I

CD

-I

CD

TIME PERk)O
Fig. 3.

Mean indices of activity at Vail underpass during the fall migration of 1971 and 1972.

�July, 1973

-185-

JOB PROGRESS REPORT

State of

--------~~~~-------COLORADO

Project No.

W-38-R-27

Work Plan No.

15

Job Title

Job No.

7

Effects of Highway Lighting on Number of Deer Killed by Vehicles

Period Covered:
Personnel:

Deer-Elk Investigations

April 1, 1972 through March 31, 1973

Thomas M. Pojar, Thomas N. Woodard and Dale F. Reed.

��-187-

EFFECTS OF HIGHWAY LIGHTING
ON NUMBER OF DEER KILLED BY VEHICLES
Dale F. Reed

P. S. OBJECTIVE
Determine if highway lighting affects the rate of deer-vehicle accidents
on a portion of Colorado 82.

SEGMENT OBJECTIVES
1.

Record the number of deer killed by vehicles at each end of, and
within a 1/4 mile lighted portion of Colorado 82.

2.

Estimate deer densities adjacent to the portion of highway affected
by lighting.

3.

Compare pre-treatment kill and count data to post-treatment kill
and count data.

4.

Gather additional information which may be needed to understand
why deer-vehicle accident rates are or are not affected by lighting.
RESULTS AND DISCUSSION

The decision made by the Colorado Division of Highways and Division of
Wildlife to delay installation of the i11uminaries was continued through
this segment. However, implementation of the agreement to install the
luminaries was initiated February 8, 1973 (Appendix A). The number of
deer, the number of deer-vehicle accidents, and several weather measurements were monitored (Appendix B).

Prepared

bY~4k:{CkQ
Dale F. Reed
Wildlife Researcher

�-188-

February 8, 1973

APPENDIX A

DEER-HIGHWAY LIGHTING PROJECT
(F.H.A. Project 1478)

The Deer-Highway Lighting Project (F.H.A. No. 1478) was approved September
11,1969.
The installation of the luminaries for this study has been delayed for various reasons.
We recommended implementation of the agreement (dated September 19, 1969)
between the Colorado Division of Wildlife and the Colorado Division of
Highways. Installation of the luminaries is to be completed during the
summer of 1973 with the exact location of the lighted area to be coordinated
with the Deer-Vehicle Accident Study personnel in Glenwood Springs.
The following changes are suggested by the Division of Wildlife to update
the agreement.
1)

Light a one-quarter mile section of highway rather than a threequarter mile section as originally proposed.

2)

Under Section 4, Method of Study, add the following to paragraph
one: "In order that pre-treatment and treatment data can be
gathered during the same season, the lights will be turned on
at dusk nightly for a one-week period and then be turned off
and left off for a one-week period during the season of evaluation."

3)

Section 13 wording to be changed to read: "Work on this project
is scheduled for two seasons of evaluation. The final report is
to be completed no later than six months after the final season
of evaluation."

�-189-

APPENDIX B

RESEARCH
1973 Deer-Highway Lighting Project Report

The number of deer-vehicle accidents and the number of deer counted in
the Jammaron study area showed an increase in 1973 over the past two
years (Figs. 1 and 2). The number of deer-vehicle accidents (deer kill)
increased linearly from January to March (Fig. 1).
Snow depths adjacent to and above the study area approximated 10.0, 18.5
18.8, and 9.5 inches for December, January, February, and March, respectively.
Hence, the relatively moderate to severe winter conditions continued during
the January, February, and March period. The average minimum temperature
for January through March decreased in 1973 (Fig. 3), while the total amount
of precipitation increased to 1.69 inches.
Installation
Installation of the luminaries was recommended as of February 8, 1973.

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�HIGHWAY 82 SPOTLIGHT COUNT DATA

(JAMMARON

130

AREA)

120

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Fig. 2. Number of deer counted adjacent to the Jammaron study area south of Glenwood Springs.

MAA

�HIGHWAY 82 PRECIPITATION AND TEMPERATURE

JANUARY, FEBRUARY, AND MARCH

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1969

1970

1971

Fig. 3. Average minimum temperature and total precipitation
of Glenwood Springs during January, February, and March.

1972

1973

for the Jammaron study area south

�July, 1973

-193-

JOB FINAL REPORT

State of

.:;:.CO:::,;L::;,;O::.,:RA=D:,;;O:..._

Project No.

W-38-R-27

Work plan No.
Job Title

Period Covered:
Personnel:

15

Deer-Elk Investigations
Job No.
:--....;8~
Ef.f.ec.t
..o.f.Light.ed.D.e.er
Crossing Signs
on Number of Deer Killed by Vehicles

- __

April 1, 1972 through March 31, 1973

Larry Green, Thomas N. Woodard, Dale F. Reed and Thomas M. Pojar.

ABSTRACT

Two lighted, animated deer crossing signs were installed adjacent to State
Highway 82, south of Glenwood Springs, Colorado, delineating a one-mile segment
of highway where deer-vehicle accidents frequently occurred. The signs were
turned on and off for alternate weekly periods in 1972 and 1973 during the time
when most deer-auto accidents were expected to occur. Numbers of deer crossing
the highway were estimated and all deer-vehicle accidents were documented. The
number of crossings per deer killed on the highway was 56.9:1 and 56.5:1 with
the signs on and with the signs off respectively. There was no difference in the
ratios (P"&gt;0.50). Mean vehicle speeds were lower (P&lt;0.05) with the signs on
but the reduction in speed was less than 3.0 mph. When three deer carcasses
were placed along the highway with the signs on the mean vehicular speed dropped
6.27 mph. With the signs off the difference was 7.85 mph. There was no significant difference in the mean speeds (P&lt;O.OOl) when dead deer were on the highway
whether the signs were on or off. Apparently motorists did see the sign but
their response in the form of speed reduction and/or increased awareness was not
sufficient to affect the crossings per kill ratio.

-

�-194-

RECOMMENDATIONS

1.

In areas where deer-vehicle accidents are especially numerous the
motorist should be given some warning for public relations and
liability reasons. A prototype of the lighted, animated sign cost
$2,000 (1971 prices) and line power (110 volt) was needed. Use
of conventional signs which cost $94 (1973 prices) each for materials,
is more practical.

�-195-

EFFECT OF LIGHTED DEER CROSSING
SIGNS ON NUMBER OF DEER KILLED BY VEHICLES
Thomas M. Pojar

P. S. OBJECTIVE
Determine if lighted deer crossing signs affect the rate of deer-vehicle
accidents on a portion of Colorado 82.

RESULTS AND DISCUSSION
Two seasons of evaluation of a lighted, animated deer crossing sign were
completed by the end of this segment. A manuscript was prepared, edited
and forwarded to the Federal Highway Administration for permission to
submit it to the Journal of Wildlife Management.

Manuscript in Preparation
Pojar, Thomas M., Richard A. Prosence, Dale F. Reed, and Thomas N. Woodard.
Effectiveness of a lighted, animated deer crossing sign. J. Wildl. Mgmt.

Prepared

by--tl;;::::J 1!?,

I~

Thomas M. Pojar
Asst. Wildlife Researcher

��July, 1973

-197-

JOB PROGRESS REPORT

State of

-:;C,;;,OLO::;;;,;RA;;;;:;:D:;,;O::._

Pro.ject No.

W-38-R-27

Wo.rk Plan No.

15
Jo.b No.
~1~0~
~~-------An Bva IuatLon iof Deer'-Proof Fence Length Required to.
Prevent Deer Movements o.n or Acro.ss High Speed Highways

Job Title

Deer-Elk Investigations

Perio.d Co.vered: April 1, 1972 thro.ugh March 31, 1973
Perso.nnel: Dale F. Reed, Tho.mas M. Po.jar, William R. Heicher, Larry L. Green,
J. Kris Moser and Tho.mas N. Wo.o.dard.

ABSTRACT

Deer cro.ssings were reduced 82 percent during the first March-May perio.d
after the 1.1 mile 8-fo.ot fence o.n Highway 82 was installed. Annual deer
kill was reduced by 61 percent. Deer kill was reduced by 72 percent and
71 percent after two. 2.25 mile 8-fo.o.tfences were installed adjacent to.
Interstate 70 near Avon and Edwards, respectively.
Sightings of 6 marked
deer near the Vail underpass-fencing complex indicated mo.vements parallel
to the fence distances o.f 0.25 to. 1.00 miles.

��-199-

AN EVALUATION OF DEER-PROOF
FENCE LENGTH REQUIRED TO PREVENT DE.ER
MOVEMENTS ON OR ACROSS HIGH SPEED HIGHWAYS
Thomas N. Woodard

P. S. OBJECTIVE
To evaluate length of deer-proof fence in relation to deer movement
on or across high speed highways.

SEGMENT OBJECTIVES
1.

Install deer-proof fence along appropriate sections of highway
as determined by Job 1a.

2.

Evaluate the effectiveness of various lengths of deer-proof fence
in affecting deer movements.

METHODS AND ~TERIALS
Methods and materials have been described by Pojar (1972).

DESCRIPTION OF AREA
Highway 82 - Diamond S
The Highway 82 study area has been described by Pojar (1972).
Interstate 70 - Avon Area
The Interstate 70 - Avon study area consists of a segment of the
Interstate from the Avon Interchange east 2.25 miles to the Eagle
River Bridge. This section of highway was opened to traffic October
1, 1971. An 8-foot fence was completed along the north highway rightof-way on October 5, 1972 (Fig. 1).
Open sagebrush areas and various browse species are common on the
north side of the highway. Alfalfa fields are prevalent south of
the highway. Deer inhabit this area from August until snow depth
precludes use, usually by late December.

�-200-

Fig. 1. The Colorado Division of Highways and Division of Wildlife
cooperated in planning and installation of 8-foot fence adjacent to
1-70 near Avon.

�-201-

Interstate 70 - Edwards Area
The Interstate 70 - Edwards study area consists of a segment of the
Interstate from the Edwards Interchange west 2.25 miles. This section
of highway was opened to traffic October 1, 1971. An 8-foot fence
was completed along the north highway right-of-way in July, 1972.
Deer utilize the area adjacent the highway to the north during the
winter. Sagebrush flats and pinyon-juniper communities are common.

Interstate 70 - Vail Area
The Vail underpass-fencing complex and adjoinirig area has been described by Reed (1971).

RESULTS AND DISCUSSION

Highway 82 - Diamond S
Deer concentrate in the crested wheatgrass fields northeast of the
highway in late winter and early spring (Table 1).

Table 1. The March-May mean nUmber of deer observations on spotlight
counts between quarter-mile section markers 25 to 30 for 1968 through
1972 on Highway 82.
Mean Count
Month

1968

1969

1970

1971

1972

134.75 (n=4)

151.25 (n=4)

104.50 (n=4)

66.75 (n=4)

102.25 (n=4)

73.00 (n=4)

34.00 (n=5)

56.00 (n=5)

51.40 (n=5)

4.50 (n=4)

May

6.00 (n=5)

0.00 (n=l)

1.50 (n=2)

0.00 (n=2)

MEAN
TOTAL

66.23 (n=13)

77.50 (n=lO)

63.73 (n=ll) 47.63 (n=ll)

March
April

53.37 (n=8)

Mean deer crossings during March-May periods in 1971 (before 8-foot fence
was installed) and 1972 (after 8-foot fence was installed) were: 1971, 11.7
(n=32); and 1972, 2.1 (n=38). This represents an 82 percent reduction in
crossings in 1972.
The mean deer kill per year before the 8-foot fence was installed (1968-1971)
was 7.75. Three deer were killed in 1972 (after 8-foot fence was installed);
a reduction of 61 percent.

�-202-

Interstate

70 - Avon Area

The mean number of deer observations on spotlight counts during the
August - December periods was 15.5 (n=Lf ) in 1971 and 13.4 (n=21)
in 1972.
Mean daily deer crossings during the period November 12 - December 29
in 1971 was 35.8 (n=24). ~ean daily crossings in 1972 before the 8-foot
fence was completed was 13.6 (n=62) for the period August 7 - October 5.
After the fence was completed in 1972 the mean daily crossings was
3.6 (n=28) for the period October 6 - November 2.
Eighteen deer were killed from October 5, 1971, to March 31, 1972, before
the fence was ins ta1led.
Five were killed from October 5, 1972, to March
31, 1973, after the fence; a 72 percent reduction.

Interstate

70 - Edwards Area

Fourteen deer were killed on this stretch of highway before the fence was
installed from October 5, 1971, to ~farch 31, 1972. Four were killed
after the fence from October 5, 1972, to March 31, 1973; a reduction
of 71 percent.

Interstate

70 - Vail Area

The movements of six marked deer in relation to the Vail fencing have
been monitored.
Two travelled parallel to the south fence a minimum of
0.4 and 0.7 miles.
Four travelled parallel to the north fence a
minimum of 0.25, 0.5, 0.65 and 1.0 miles.

LITERATURE

CITED

Poiar, Thomas M. 1972. An evaluation of deer-proof fence length required
to prevent deer movements on or across high speed highways.
Colo.
Div. Wi1dl.
Game Res. Div. Fed. Aid. Proj. W-38-R-26.
Game Res.
Rep., July, Part 3. pp. 305-310.
Reed, Dale F. 1971. Deer underpass evaluation.
Game Res. Div. Fed. Aid Proj. W-38-R-25.
Julv, Part 3. pp. 341-351.

Prepared

bv _~~~~~ffYK~~a~5~~~~.~~~~~~
Thomas
Senior Research Technician

Colo. Div. Hildl.
Game Res. Rep , ,

�July, 1973

-203-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-27.

Deer-Elk Investigations

Work Plan No.
Job Title

15
Job No. __~
~~~l~l~
Effects .0£ a Simulate.d8;-.footFence Angle
in Diverting Deer from Their Established Direction of Movement

Period Covered:
Personnel:

April 1, 1972 through March 31, 1973

Thomas N. Woodard, Thomas M. Pojar and Dale F. Reed.

ABSTRACT

Time was measured and observations made on mule deer (Odocoileus hemionus) response to five simulated fence angles. Of 19 deer, only 10 successfully moved
through the control time zone. The mean time was 101.3 seconds. Sample sizes
(n) of the angle variables (45, 60, 90, 120 and 135 degrees) were severely
limited.

_

��-205-

Fig. 1. A permanent 8-foot wire deer fence is shown in the foreground.
The simulated fence (netting) is shown installed at the 60° angle
(right center) from the permanent fence. The wires to the left and
the far right are the 90° and 45° angles, respectively.

��-207-

EFFECTS OF A SIMULATED 8-FOOT FENCE ANGLE
IN DIVERTING DEER FROM THEIR ESTABLISHED DIRECTION OF MOVEMENT
Dale F. Reed

P. S. OBJECTIVE
Evaluate the effectiveness of three angles of a simulated 8-foot fence to
divert deer from their established direction of movement.

SEGMENT OBJECTIVES
1.

Install a movable 8-foot section of 4-inch square netting adjacent
to and across heavily traveled deer trails and concentration points.

2.

Test the effectiveness of three angles under field conditions.
METHODS AND MATERIALS

An 8-foot high simulated fence made of 4-inch netting (350 lb. test) wa5
easily installed for a length of 90 feet and at one of three angles (45 ,
600, and 900; plus converse angles of 1200 and 1350) in relation to a
permanent 8-foot wire deer fence (Fig. 1). The permanent wire fence had
been placed across a deer migration route to divert migrating deer. There
was a heavily traveled deer trail along this fence.
The time for a deer to travel the 90-foot length of permanent 8-foot deer
fence, (hereafter referred to as the "control"), and the same length of
simulated fence at each angle (variables 450,600, and 900; plus converse
angles of 1200 and 1350) was measured by making early morning observations
and recording the time each deer spent between two clearly marked points
with a stop watch. The amount of time was recorded to the nearest second
for each animal. If a group approached, the first animal moving into the
marked area (time zone) was chosen for timing. When two or more deer moved
into the zone simultaneously either the "lead" animal or a tagged animal was
used.
Observations were made during early morning hours for 33 days from June 6
to July 19, 1972. The sequence of the simulated fence angles tested was:
0
0 (control), 450, 60°, and 90°. This sequence was repeated throughout the
above period. The simulated fence was installed at the angle specified by
the above sequence during the evening before the day's observation.
Factors such as the wind, light, natural cover, and camouflage clothing were
utilized to advantage during the observations. However, if the observer was
detected by the animals, the day's observations were terminated.

�-208-

DESCRIPTION OF AREA
The observations on deer response to the 8-foot fence angle simulation
were made near Vail, Colorado, adjacent to the north 8-foot fence of
the Vail underpass - fencing complex (Reed 1971). The observation site
was located on the edge of a conifer community with interspersed aspen
(Populus tremuloides) and big sagebrush (Artemisia tridentata). Sunrise occurred behind the observer's location and normal westerly winds
or breezes placed the location on the lee side of most deer during the
observations.
A dirt road paralleled the 8-foot fence, and curved away from it near
the point where the simulated fence was attached to the wire fence.
There was a gradual slope located adjacent to the road opposite the
observation point. The simulated fence angle of 900 was installed across
the road and up this slope. The fence angle of 600 was tangent to the
outer shoulder of the bend in the road. The fence angle of 450 was installed down an incline to the east and 20 to 30 feet from the road and
slope at the closest point.

RESULTS AND DISCUSSION
Time was measured on deer in the simulated fence angle area on 64 occasions. Of 19 deer recorded in the time zone during control measurements,
only 10 were successful in moving through the zone (Table 1). Nine failed
in that they moved into the zone, turned around, and left in the direction
they had come. Sample sizes (n) obtained during the evaluation of the
angle variables were not sufficient to determine a meaningful pattern.
To increase the sample sizes during the next segment, the angle of 600 (and
converse angle of 1200) will be omitted from the evaluation.

LITERATURE CITED
Reed, Dale F. 1971. Deer underpass evaluation. In Game Research Report.
July, Part 3. Colorado Game, Fish and Parks Div., Federal Aid. p. 344.

Prepared

by

A'2~¢r~
Dale F. Reed
Wildlife Researcher

�-209-

Table 1. The mean times (seconds) of successes or f u ll ures of dcor moving
through the zone with either the simulated fence angle removed (control.)
or installed (variables: 45°, 60°, or 90°). Successful animals either
moved 1) along the simulated fence and then around the end, 2) along a
dirt road (during angles of 450 and 600 only), or 3) up an adjacent slope.
Failure animals moved into the zone, turned around, and left in the direction
that they had come.
Successful

Failure

Percent Successful

101.3 (n=lO) i/
61.8 (n= 8)
95.3 (n= 3)

92.1 (n=9) ,
0
74.5 (n=2)

53
100
60

Fence
Road
Slope

0
116.3 (n= 3)
75.0 (n= 1)

52.0 (n=2)
62.0 (n=2)
211. 3 (n=3)

0
60
25

Fence
Road
Slope

0
189.7 (n= 3)
93.0 (n= 4)

0
150.7 (n=3)
0

50
100

Fence
Road
Slope

309.0 (n= 1)
206.0 (n= 2)
126.0 (n= 1)

69.0 (n=L)
0
0

50
100
100

121. 3 (n=4)
0
0

0

Angles
00

(Control)
Fence 1:./
Road ];./
Slope 1/

45 °

60°

90°

120°
0
0
0

Fence
Road
Slope
135 °
Fence
Road
Slope

73.0 (n= 1)
0
51.0 (n= 1)

0
0
0

100
100

1:./
Deer that moved along the trail adjacent to the permanent 8-foot fence.
2/
- Deer that moved along the dirt road.
3/
- Deer that came from the trail adjacent the 8-foot fence or from the road,
and moved up an adjacent slope.
4/
- Number of observations

(n).

5/
- When deer approached the simulated fence angles of 60° and 45° from the
opposite direction (east), they encountered converse angles of 120° and 135°.

��-211-

July, 1973

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~---------

Project No.

W-38-R-27

Work Plan No.

16

Job Title

Job No.

1

Piceance Deer Study - Population Distribution

Period Covered:
Personnel:

Deer~E1k Investigations

April 1, 1972 through March 31, 1973

R. M. Bartmann, S. F. Steinert, J. J. Klein, Jr., L. Crooks, R. Stone
and D. Talcott.

ABSTRACT

From December 12, 1972 to March 9, 1973, 427 deer were neckbanded on the Piceance
winter range and another 90 eartagged only. The sex and age composition of the
marked deer was 65 male fawns, 67 female fawns, 90 mature males and 295 mature
females. Sightings of 196 marked deer were reported and 37 were recovered. Aerial
deer distribution surveys were made in December, January and February. A considerable decrease in occupied winter range occurred from December to January because
of deep snow. Little new snow fell after Janaury 10 and so the February survey
was based on accumulated rather than new tracks. From the air, deer distribution
appeared little different from what it had been in January. Ground checks later
revealed that deer were more concentrated than was indicated by aerial observation of accumulated tracks.

��-213-

PICEANCE DEER STUDY - POPULATION DISTRIBUTION
Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer
population density and structure data on pinyon-juniper winter range
for use in making annual harvest recommendations.
SEGMENT OBJECTIVES
1.

To delineate deer sub-population boundaries and concentration areas
on the Piceance winter range.
a.

Define winter range boundaries of the Piceance deer population
under mild, moderate, and severe winter conditions.

b.

Define deer sub-population boundaries.

c.

Delineate major deer concentration areas.
METHODS AND MATERIALS

See Bartmann (1972).
RESULTS AND DISCUSSION

Deer Trapping
Deer trapping began December 12, 1972, and terminated March 9, 1973.
Trapping conditions the past winter were markedly different from those
of the relatively mild winter the preceding year. Deep snow restricted
vehicular access in many areas and srtowmobiles were used to move and
check traps most of the winter; consequently only 30 to 40 traps were
set at anyone time. Although this was only about half of the number
used the previous year, the mean number of deer marked per day was comparable; 5.9 in 1973 and 5.7 i~ 1972. Marked deer quotas were reached
in five of the seven trapping areas and about 250 deer remain to be
neckbanded in Areas 4 and 5.
Beginning in mid-January, fawns were eartagged only. The condition of
fawns deteriorated noticeably late December to March and many were not
expected to survive the winter.

�-214-

The use of ear-streamers was also discontinued this past winter. From
the comments of hunters and others who observed marked deer, the red
streamers did help attract attention to marked animals. However, there
were several reports during the 1972 big game season of deer with red
streamers and red neckbands. To our knowledge, no deer anywhere near
the White River area were marked in this manner. Most people probably
confused the streamers with neckbands. Since red neckbands were used
this past winter, the ear-streamers were not.
Four hundred twenty-seven deer were both eartagged and neckbanded
during the 87-day trapping period. Another 90 deer, mostly fawns, were
eartagged only for a total of 517 deer marked (Table 1). The sex and
age composition was 65 male fawns, 67 female fawns, 90 mature males,
and 295 mature females. The percent fawns was 26, or 8 percent less
than in the previous year. This decline was offset by an increase in
mature females from 48 to 57 percent. The percent of mature males was
nearly constant, 18 and 17 in 1972 and 1973, respectively.
There was only one recatch of a deer banded the previous year even
though about three weeks were spent in areas trapped in 1972. Another
recatch was made in Area 6 of a deer banded earlier in the winter in
Area 1.

Marked Deer Sightings and Recoveries
Sightings of 196 marked deer were reported through March 15, 1973 (Table
2). Forty-seven were considered summer sightings (May through September)
and 149 winter sightings (October through April). In addition, 37 marked
deer were recovered, most during the deer hunting season in October
(Table 3). Comments and conclusions concerning the results of the trapping
program will be reserved until data from all portions of the winter range
provide a more complete picture of deer movements.

Deer Distribution
The 1972-73 winter was unusually severe. Snow depths ranged from 15 to
30 inches over much of the winter range during January and February and
mean temperatures averaged about 100 F lower than in 1972 according to
Little Hills weather records. Even though little snow fell after January
10, low temperatures prevented much melting and settling of the snow pack
until late February. South slopes, which are usually bare within a week
after most snowfalls, remained snow-covered nearly all winter. Little
snow-crusting occurred until February and then only on south exposures.
Deep powder snow prevented travel by snowmobile over much of the higher
winter range areas and ground checks of snow depths and deer distribution
were kept to a minimum.
During the first aerial distribution survey December 6, 10 and 11, 1972,
snow depths varied from about 12 inches at lower elevations to 20 inches

�-215-

or more at the higher levels. As this was the first large accumulation
of snow for the winter, deer were still fairly well scattered. Several
areas of heavy deer use were noted along Highway 13, adjacent the upper
White River, and along Hunter Creek (Fig. 1). Deer had already vacated
some of the treeless areas and steep timbered north slopes at higher
elevations in the Piceance triangle. Some of the larger open sagebrush
and chained areas in the western portion of the winter range were also
devoid of deer tracks. Although deer were present on the more extensive
south slope areas, track densities indicated only light to moderate use.
During the January 5,6 and 7, 1973 survey, it was difficult to assign
track density ratings. Deep snow, ranging from 18 to 30 inches, had
forced deer to use defined trails in many areas. The upper limits of
deer use had lowered since the previous month and additional concentration
areas were noted (Fig. 2). Deer sign was particularly heavy in the small
sagebrush parks found at the heads of most drainages and deer were beginning to concentrate around haystacks on irrigated meadows. Use on
south slopes was also becoming heavier.
The final survey was flown February 19 and 20, 1973. No attempt was made
to classify track densities as distribution had to be based on accumulated
rather than new tracks. Only 2-1/2 inches of snow fell after January 10
and it was hoped that this small amount of new snow together with windblown snow would cover most of the older tracks. Some southerly exposures
were beginning to bare but snow depths on the rest of the area averaged
only two to three inches less than during the January ,survey.
Upper deer distribution limits had constricted in some areas but not as
much as expected (Fig. 3). Spot checks on the ground later revealed that
the deer were actually concentrated more than indicated by the aerial
survey. Apparently, new and blowing snow failed to cover many older
tracks as assumed. Also, deer use was heavier on south slopes than
estimated from the air. The presence of snow-free patches together with
the slow but continuous melting of snow tended to obscure tracks on those
areas. Deer were concentrating heavily in and around bottomlands and dead
animals, mostly fawns, were largely found around haystacks.

LITERATURE CITED
Bartmann, R. M. 1972. Piceance deer study - population distribution.
p. 315-337. In Game Research Report. Colo. Div. Wildl., Denver.
3(Part 3):253-377.

2

Prepared by .

-:?"'-&lt;..~-~
Richard M. Bartmann
Wildlife Researcher

�Tah1e 1.

Date
(1972)

Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1972-73.

Sex

Age

Eartag
No.

Neckband
No.

Twp.

Location
Range
Sec.

IN
IN
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN

99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W
99W

Remarks

Area 1 - Blue Neckbands
12-13
12-16
12-14
12-14
12-14
12-14
12-15
12-15
12-15
12-15
12-15
12-15
12-15
12-15
12-16
12-16
12-16
12-16
12-16
12-16
12-16
12-16
12-16
12-16
12-17
12-1712-17
12-17
12-17
12-18

Female
Male
Female
Female
Female
Male
Male
Female
Female
Female
Female
Male
Male
Female
Male
Female
Male
Male
Female
Female
Female
Female
Female
Female
Female
Male
Male
Female
Female
Female

Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature

L-1697
L-1698
L-1726
L-1727
L-1728
L-1729
L-1730
L-1731
L-1732
L-1733
L-1734
L-1735
L-1736
L-1737
L-1738
L-1739
L-1740
L-1741
L-1742
L-1743
L-1744
L-1745
L-1746
L-1747
L-1700
L-1714
L-1715
L-1716
L-1717
L-1719

96
115
58
59
85
95
121
76
78
119
120
73
92
124
122
101
93
None
None
103
125
126
None
127
129
114
118
132
131
134

36
11
36
35
35
14
35
35
35
14
14
14
14
14
14
14
14
35
35
14
14
14
14
14
12
12
11
11
11
12

I
f',,)

.....

0'1
I

Found dead 3-17-73

-----------------------------------------------------------------------------------------------------------

�Table 1. Record of deer trapped and marked on the Kinter range in Game Management
(continued) .
Date
(1972)

Age

Eartag
No.

Neckband

Sex

No •

Twp.

12-21
12-21
12-22
12-22
12-22
12-25
12-25
12-27
12-28
12-28
12-24
12-24
12-24
12-29

Female
Male
Female
Male
Female
Female
:1a1e
Female
Female
Female
Female
Female
~1a1e
Female

i'1ature
Hature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature

L-1722
L-1723
L-1725
L-1764
L-1765
L-1749
L-1754
L-1755
L-1760
L-1756
L-1750
L-1751
L-1752
L-1757

128
123
130
None
135
136
105
133
107
112
137
138
113
109

IN
IN
IN
IN
IN
2N
2N
2N
2N
2N
2N
2N
2N
2N

Summary:

7 Hale Fawns;

9_X~m_al_e_Xawn~;

7 Mature Males;

Location
Range
Sec.
99W
99W
99W
99Wo
99W
99W
991&gt;1
99W
99W
991&gt;1
99W
99W
99W
99W

11
11
12
12
11
16
17
17
16
16
16
16
16
16

21 Mature Females

Area 2 - White Neckbands
12--17
12-18
12-20
12-21
12-22
12-22
12-23
12-23
12-24
12-24
12-24
12--25
_____________

Hale
Female
~1a1e
Female
Male
Female
Female
Female
Female
:Female
:Female
Female
0

Mature
Mature
Hature
Hature
Fawn
Mature
Mature
Mature
:v1ature
Hature
l1ature
Fawn

_________________________________________________________________________________________________

L-1718
L-1720
L-1721
L-1724
L-1766
L-1767
L-1768
L-1769
L-1770
L-1771
L-1722
L-1774

214
213
207
218
211
219
210
220
215
216
212
222

IS
18
18
IS
18
IS
IS
IS
IS
IS
IS
IS

Unit 22, winter 1972-73

99W
991i7
99W
99W
99W
99W
99W
99W
99v7

99W
98:~
99W

34
34
34
11
11
34
11
10
10
34
31
34

Remarks

I
N

t-'

"
I

�Table l.
Record of deer trapped and marked on the'Kinter range in Game Management
(continued) .
Date
(1972-73)

Sex

Age

12-25
12-25
12-27
12-28
12-29
12-30
12-31
1-1-73
1-2
1-2
1-2
1-3
1-3
1-3
1-4
1-4
1-4
1-4

Male
Female
Male
Female
Male
Female
Female
Female
Female
Male
Female
Female
Male
Male
Male
Female
Female
Female

Summary:

5 Male Fawns;

Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature

Eartag
No.

Neckband

~~o•

Twp.

L-1773
L-1775
L-1753
L-1759
L-1761
L-1762
L-1814
L-1818
L-1823
L-1824
L-1825
L-1758
L-1807
L-1808
L-1833
L-1834
L-1835
L-1836

225
221
217
223
227
229
230
228
235
226
None
236
224
231
232
240
238
239

IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS

4 Female F~.3

5 Hat ure Hales;

Unit 22, winter 1972-73

Location
Range
Sec.
99w
99W
98W
99W
99W
98W
98w
98W
99W
99W
98W
98W
98W
99w
98W
98w
98W
98W

Remarks

34
34
31
31
34
31
31
31
10
3
31
31
31
33
32
32
31
31

I
N

t-I

00
I

16 Hature Females

Area 3 - Pink Neckbands
12-29-72
12-30
12-31
12-31
12-31
1-1-73
1-1
1-1
1-1
1-2

Female
Female
Female
Female
Male
Male
Female
Male
Female
Female

Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature

L-1801
L-1763
L-1815
L-1816
L-1817
L-1819
L-1820
L-1821
L-1822
L-1802

26
63
27
16
59
'58
28
19
30
29

2S
2S
3S
3S
3S
2S
2S
2S
2S
2S

97W
97W
97W
97W
97W
97W
97T-J
971"
97W
971"

29
29
9
4
9
29
29
29
29
29

May be neckband No. 61
Hay be neckband No.4':;

---------------------------------------------------------------------------------------------------------------

�Table l. Record of deer trapped and marked on the t-;interrange in Game Management Unit 22, winter 1972-73
(continued) .

.Date
(1973)
1-2
1-2
1-2
1-2
1-3
1-3
1-3
1-3
1-3
1-3
1-3
1-4
1-4
1-5
1-6
1-6
1-7
1-7
1-7
1-7
1-7
1-7
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8
1-8

Sex

Age

Eartag
No.

Keckband
"J
.~o•

Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Male
Male
Female
Male
Male
Male
Male
Female
Female
Female
Male
Female
Female
Male
Male
Male
Female
Female
Female
Female
Female

Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn .
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn

L-1803
L-1804
L-1805
L-1806
L-1830
L-1829
L-1828
L-1827
L-1826
L-1831
L-1832
L-1837
L-1851
L-1852
L-1853
L-1854
1-1855
L-1856
L-1857
L-1858
L-1859
L-1860'
L-1863
L-1776
L-1777
L-1778
L-1779
L-1780
L-1781
L-1782
L-1783
L-1784
L-1861
L-1862

53
62
54
31
10
9
8
6
7
11
13
65
41
57
61
34
51
39
24
18
66
64
32
21
15
37
14
22
20
36
33
35

17
25

Twp.

28
28
38
3S
38
28
28
28
28
38
38
38
38
38
38
38
38
38
38
38
38
38
38
38
38
38
38
35
38
38
28
28
38
38

Location
Range
Sec.
97W
97W
97W
97W
97W
97W
97W
97W
97W
97Ti!
97W
97W
97W
97W
97W
96W
97W
97W
97W
97W
96W
96W
96W
96W
97W
97W
97W
97W
97W
97W
98W
98W
96W
96W

29
29
9
9
9
32
29
29
29
9
9
4
4
2
9
21
9
2
2
11
9
9
16
16
2
2
4
4
4
9
24
24
15
15

Remarks

I
N

I-'

\0
I

May be neckband No. 16

May be eartag L-1859
May be eartag L-1858
May be eartag L-1777
May be ear tag L-1863

--------------------~-------------------------------------------~~--------------------------------------------

�Table 1.

Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1972-73
(continued).

Date
(1973)

1-9
1-9
1-9
1-9
1-9
1-9
1-9
1-9
1-9
1-9
1-9
1-10
1-10
1-10
1-10
1-10
1-10
1-10
1-10
1-11
1-11
l-ll

r-ri
l-ll
1-11
l-ll
1-11
1-11
1-11
1-11
1-11

Sex

Female
Female
Male
Female
Female
Male
Male
Female
Female
Female
Female
Female
Female
Female
Female
Male
Male
Female
Female
Male
7emale
Female
Female
Hale
Female
Female
Female
Hale
Female
Hale
Female

Age
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Hature
Fawn
Mature

Eartag
No.
L-1785
L-l786
L-l787
L-1838
. L-1839
L-1840
L-1841
L-1842
L-1843
L-1844
L-1845
L-1846
L-1847
L-1809
L-1810
L-1811
L-1812
L-1813
L-1788
L-1789
L-1890
L-1791
L-1791
L-1793
L-1794
L-1795
L-1796
L-1797
L-1798
L-1848
L-1849

r-:"e
ckband
No.

Twp.

Location
Range
Sec.

73
68
46
70
72
55
44
71
67
74
69
None
None
90
75
96
97
76
77
47
86
89
3
95
91
None
None
None
None
43
82

38
38
38
38
38
38
38
38
28
28
28
28
28
38
38
38
38
38
38
38
3S
38
38
38
38
3S
38
38
38
2S
28

96W
96W
96W
96w
96W
97W
97W
97W
98W
98W
97W
97W
97W
96W
96W
96W
97W
97W
97W
96W
96W
96W
97W
97W
97W
97W
97W
97W
97W
97W
97W

9
15
16
21
9

Remarks

May be neckband No. 59

2

14
4
24
24
7
7
7
15
16
16
14
4
4
9
16
16
2

2
4
4
9
9
9
7
7

-------------------------------------------------------------------------------------------------------------

I

N
N

0
I

�Table 1. Record of deer trapped and marked on the wtnter range in Game Management Unit 22, winter 1972-73
(continued) .
Date
(1973)

Sex

Eartag
No.

Age

Ke ckband

:~o•

Twp.

Location
Range
Sec.

Remarks

1-21
1-24
1-25
1-27
1-27

Female
Female
Female
Female
Male

Mature
Mature
Mature
Mature
Mature

StmlIIlar.r:
14 Male Fawns;

L-1942
L-1946
L-1953
L-1963
L-1968

20 Female Fawns;

40
None
23
None
50
21 Mature Males;

2S
2S
2S
2S
2S

97W
97W
97W
97W
97W

29
29
29
29
29

56 Mature Females

Area 5 - Yellow with Red Stripe Neckband
2-24
2-24
2-24
2-28
2-28
2-28
3-1
3-1
3-1
3-2
3-3
3-4
3-4
3-4
3-4
3-5
3-5
3-5
3-5
3-5

Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Hale
Female
Female

Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature
Hature
Mature
Mature

L-2l82
L-2184
L-2l86
L-230l
L-2302
L-2303
L-2l88
L-2304
L-2305
L-2306
L-2307
L-2308
L-2309
L-23l0
L-23ll
L-2312
L-2313
L-2314
L-2315
L-23l6

1
3
5
14
13
6
12
15
4
2
20
None
18
None
19
None
25
7
16
17

IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS

96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

5
8
15
23
23
26
9
9
23
26
15
26
15
15
16
15
15
22
16
17

------------~-------~---------------~---------------------------------~--~------------------.---~---------------

I

N
N

t-'
I

�Table 1.
Record of deer trapped and marked on the Kinter range in Game Management
(continued).
Date
(1973)

Sex

Age

1-11
1-11
1-11
1-12
1-12

Male
Male
Female
Male
Female

Fawn
Mature
Fawn
Mature
Fawn

1-12
1-12
1-12

Male
Female
Female

Mature
Mature
Mature

1-12
1-12

Female
Female

Mature
Mature

Eartag
No.

Ne ckband

Location
Range
Sec.

Ho.

Twp.

L-1850
L-1864
L-1865
L-1866
L-1867

99
100
81
56
87

2S
2S
2S
2S
2S

98W
98w
98W
98W
98W

12
12
12
12
12

L"'"1799
L-1800
L-1876

52
94

2S
3S
3S

98w
97W
96W

24
2
'9

L-1877
L-1868

80
88

3S
2S

96W
97W

15
7

97W
96W
96W
96W
96W
97W
97W
98W
98W
97W
97W
97W

7
16
16
21
21
12
2
12
12
7
18
18

{j

1-12
1-12
1-12
1-12
1-12
1-13
1-13
1-13
1-13
1-13
1-13
1-13

Female
Male
Male
Female
Female
Female
Male
Female
Female
Hale
Female
Female

Fawn
Fawn
Fawn
Fawn '
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature

L-1869
L-1878
L-1879
L-1880
L-1881
L-1883
L-1882
L-1870
L-1871
L-1872
L-1873
L-1874

79
49
48
None
None
42
60
85
84
38
78

28
3S
38
38
3S
3S
3S
2S
2S
2S
28
2S

1-13
1-13
1-14
1-14

Male
Female
Female
Female

Fawn
Mature
Mature
Mature

L-1875
L-1888
L-1899
L-1898

None
45
None
93

2S
2S
2S
2S

97W
98W
97W
97W

18
24
12
12

L-1889
L-1890
L-1891

83
89
None

3S
3S
3S

97W
97W
97W

2
2
2

1-14
1-14
1-14

Female
Female
Female

Mature
Mature
Fawn

92

Unit 22, winter 19i2-73

Remarks

May be neckband No. 78
or 88

Sex and age could be in
error
May be neckband No. 78
or 87

Hay be neckband No. 93
May be neckband No. 87
or 88

May be neckband No. 92
and ear tag No. L-1896

---------------------------------------------------------------------------------------------------------------

1
N
N

~

I'

�Table 1.

Record of deer trapped and marked on the Kinter range in Game Management Unit 22, winter 1972-73
(continued) .

--,

Date
(1973)

Sex

Age

Eartag
No.

Keckband
No.

Twp.

1-20
1-20

Female
Female

Fawn
Mature

L-1940
L-194l

None
62

28
28

97W
97W

29
29

3-5
3-6
3-6
3-7
3-7
3-7
3-7
3-7
3-8
3-8
3-8
3-9
3-9
3-9
3-9

Female
Female
Female
Male
Female
Male
Female
Female
Female
Female
Female
Male
Male
Male
Female

Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature

L-23l7
.L-23l8
L-23l9
L-2320
L-232l
L-2322
L-2323
L-2324
L-2325
L-2326
L-2327
L-2328
L-2329
L-2330
L-233l

21
34
32
22
24
10
26
29
28
23
36
11
8
38
39

IS
18
18
18
18
18
18
18
18
IS
18
IS
18
18
18

96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

23
15
15
15
15
15
9
23
23
15
23
23
23
15

8ummarx.: 1 Male Fawn;

2 Female Fawns;

7 Ma ture Ha l.es;

Location
Range
Sec.

Remarks

Has red with white
stripe neckband

8

25 Mature Females

Area 6 - Blue and White Neckbarids
2-7
2-8
2-8
2-8
2-10
2-10
2-10
2-10
2-10

Female
Female
Male
Female
Female
Female
Male
Female
Female

Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature

L-2048
L-2035
L-2036
L-2037
L-2068
L-2069
L-2070
L-2071
L-2072

13

11
None
2
14
15
20
16
17

2S
28
28
28
28
28
28
28
2S

97W
97W
97W
97W
97W
97W
97W
97W
97W

10
3
10
10
4
4
10
4
4

------------------_ .._------------------------------------------------------------_._-------..;..------------------------

I

N
N

w
I

�Table l.
Record of deer trapped and marked on the w.l nter range in Game Hanagement Unit 22, winter 1972-73
(continued) .
Date
(1973)

2-10
2-10
2-10
2-11
2-11
2-11
2-11
2-11
2-11
2-11
2-11
2-12
2-12
2-12
2-12
2-12
2-12
2-12
2-12
2-12
2-12
2-12
2-13
2-13
2-13
2-13
2-13
2-13
2-13
2-13
2-13

Sex

Eartag
No.

Ne ckband

Age

Female
Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
Male
Female
Male
Female
Female
Female
Female
Male
Male
Male
Male
Female
Female
Female
Female
Female
Female
Male
Male
Female

Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature

L-2073
L-2074
L-2075
L-208l
L-2082
L-2083
L-2084
L-2085
L-2086
L-2087
L-2088
L-2089
L-2090
L-209l
L-2092
L-2093
L"-2094
L-2095
L-2096
L-2097
L-2098
L-2099
L-2l0l
L-2102
L-2103
L-2l04
L-2l05
L-2l06
L-2107
L-2108
L-2l09

6
10
1

~o•
-..

35
30
29
24
23
None
31
5
None
7
None
12
18
21
26
4
8
None
None
39
40
42
43
41
50
22
23
24

,

Twp.

Location
Range
Sec.

IS
18
IS
28
2S
2S
2S
2S
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
2S
2S
2S
2S
IS
IS
IS
IS
IS
IS
IS
IS
IS

97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W

Remarks

27
27
26
4
4
10
3
3
27
26
24
24
14
22
22
22
26
26
4
10
4
5
13
13
14
14
13
24
24
14
22

---------------------------~---------------------------------~-----------------------------------------------

I

N
N

.pI

�Table 1. Record of deer trapped and marked on the wl.nt er range in Game Hanagement Unit 22, winter 1972-73
(continued) .
Date
(1973)

Age

Eartag
No.

Ne ckband

Sex

Ho.

Twp.

2-13
2-13
2-13
2-13
2-14
2-14
2-14
2-14
2-14
2-14
2-15
2-15
2-15
2-15
2-15
2-15
2-15
2-15
2-16
2-16
2-16
2-16
2-16
2-16
2-17
2-17
2-17
2-17
2-17
2-17
2-17
2-17

Female
Female
Female
Female
.Female
Female
Female
Female
Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
Male
Male
Male
Male
Female
Female
Female
Female
Male
Female
Male
Male
Female
Female

Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Maturw
Mature
Mature
Fawn .
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn ,.
Mature
Mature
Mature

L-2110
L-2111
L-2112
L-2113
L-2076
L-2114
L-2115
L-2116
L-2117
L-2118
L-2119
L-2120
L-2121
L-2122
1-2123
L-2124
L-2125
L-2126
L-2127
L-2128
L-2129
L-2130
L-213l
L-2132
L-2133
L-2134
L-2135
L-2136
L-2137
L-2l39
L-2138
L-2140

25
52
57
53
None
49
33
32
None
54
46
59
58
63
60
55
None
51
37
38
None
None
None
65
67
71
78
68
None
74
64
80

IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
1S
IS
2S
2S
IS
IS
IS
IS
IS
IS
1S
IS
IS
1S
IS
IS
IS
IS

Location
Range
Sec.
97W
97W
97W
97W
97W
97W
97W
96W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97liJ

97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W

Remarks

22
27
26
26
12
12
13
29
14
14
12
13
14
13
14
26 '
4
4
14
14
14
22
26
26
12
12
13
14
13
13
13
22

------------------------------------------------------~------------------------------------------------------

I
N
N
VI
I

�Table 1.
Record of deer trapped and marked on the ~-lnter range: in Game Management Unit 22, winter 1972-73
(continued).
Date)
(1973)
2-17
2-17
2-17
2-18
2-18
2-18
2-18
2-18
2-18
2-18
2-18
2-18
2-18
2-18
2-18
2-19
2-19
2-19
2-19
2-19
2-19
2-19
2-19
2-19
2-19
2-20
2-20
2-20
2-21
2-21
2-21

Sex

Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Male
Female
Female
Female
Male
Female
Female
Female
Male
Female
Female
Male
Female

Age

Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature

Eartag
No.

Keckband
No.

L-2141
L-2142
L-2143
L-2145
L-2146
L-2147
L-2148
L-2149
L-2150
L-2151
L-2152
L-2153
L-2154
L-2155
L-2156
L-2157
L-2158
L-2159
L-2160
L-2161
L-2162
L-2163
L-2164
L-2165
L-2166
L-2167
L-2168
L-2169
L-2170
L-2171
L-2172

62
None
69
73
70
79
None
None
56
66
None
72
44
19
34
96
None
47
None
104
103
83
None
106
107
88
None
94
100
36
82

Twp.

IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
2S
IS
IS
IS
IS
IS
IS
2S
2S
2S
2S
IS
IS
IS
IS
IS
IS

Location
Range
Sec.

97W
97W
97w
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97ft!

97W
97lv
97W

Remarks

22
27
27
13
14
14
13
13
13
13
22
27
26
26
4
14
13
13
22
27
26
4
3
3
3
13
26
26
14
22
26

-------------------------------------------------------------------------------------------------------------

I

N
N
0-.
I

�Table 1. Record of deer trapped and marked on the wtnter range in Game Management Unit 22, winter 1972-73
(continued).
Date
(1973)

Age

Ear tag
No.

Neckband

Sex

i~o•

Twp.

2-22
2-22
2-22
2-22
2-22
2-22
2-22
2-22
2-22
2-23
2-23
2-23
2-23
2-23
2-24
2-24
2-24
2-24
2-24
2-24
2-24
2-24
2-25
2-25

Female
Male
Female
Male
Female
Female
Female
Female
Female
Male
Male
Female
Female
Male
Male
Male
Female
Male
Male
Male
Female
Female
Female
Female

Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature

L-2173
L-2174
L-2175
L-2176
L-2177
L-2178
L-2179
L-2180
L-2181
L-2189
L-2190
L-2191
L-2192
L-1193
L-2194
L-2195
L-2196
L-2197
L-2198
L-2199
L-2200
L-2183
L-2185
L-2187

92
48
None
None
86
87
85
95
97
75
None
90
102
None
None
None
93
45
76
None
105
84
101
77

IS
IS
IS
IS
2S
2S
2S
2S
2S
IS
IS
IS
15
2S
IS
1$
IS
IS
IS
IS
IS
15
IS
IS

Summarz:

18 Male Fawns;

10 Female Fawns;

18 Mature Males;

Location
Range
Sec.
97W
96W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
96W
96W
97W
97W
97W
97W
97W

Remarks

12
19
14
22
4
4
4
3
3
14
22
22
26
3
13
13
13
19
19
14
27
26
12
14

81 Mature Females

Area 7 - Red with White Stripe Neckbands
1-13
1-13

Female
Female

Fawn
Mature

L-1884
L-1885

6
7

35
35

95W
95W

4
4

--~----~---------------------------------------------------------------~----------------------------------------

I

N
N
-...J
I

�Table 1.

Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1972-73
(continued) .

Date
(1973)
1-13
1-13
1-14
1-14
1-15
1-15
1-15
1-15
1-15
1-15
1-15
1-15
1-15
1-14
1-14
1-14
1-14
1-16
1-16
1-16
1-16
1-16
1-16
1-16
1-16
1-17
1-17
1-17
1-17
1-17
1-17
1-17

Sex
Male
Male
Male
Female
Female
Female
Male
Male
Female
Female
Female
Female
Female
Male
Female
Male
Female
Male
Male
Female
Female
Na1e
Male
Female
Female
Male
Female
Female
Female
Female
Female
Male

Eartag
No.

Age
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature

I

L-1886
L-1887
L-1897
L-1896
L-1900
L-1901
L-1902
L-1903
L-1904
L-1905
L-1906
L-1907
L-1908
L-1892
L-1893
L-1894
L-1895
L-1909
L-1910
L-1911
L-1912
L-1913
L-1914
L-1915
L-1916
L-1917
L-1918
L-1919
L-1920
L-1921
L-1922
L-1923

t'eekband
;~o.
5
3
2
10
11
12
22
25
14
15
21
16
13
1
8

4
9
37
36
26
27
38
24
20
None
40
35
30
29
28
33
18

Twp.
38
28
28
28
28
28
28
38
38
28
28
28
28
28
38
38
38
28
28
3S
3S
38
28
28
28
28
28
28
28
28
28
28

Location
Range
Sec.
95W
95W
95W
95W
96W
96W
96W
95W
95W
95W
95W
95W
95W
97W
95W
95W
95W
97W
96W
95W
95W
95W
95W
95W
95W
97W
97W
96W
96W
96w
96W
96w

4
34
34
34
30
30
19
4
'4
343434
34
24
4
4
4
25
30
4
4
4
34
34
34
25
25
30
30
30
30
30

Remarks

May be eartag L-1898

---------------------------------------------------------------------------------------------------------------

I

N
N
00
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1972-73
(continued).

Date
(1973)

Sex

Age

Eartag
No.

Keckband
No.

Twp.

1-17
1-18
1-18
1-18
1-18
1-19
1-19
1-19
1-19
1-19
1-19
1-19
1-19
1-20
1-20
1-20
1-22
1-22
1-22
1-24
1-24
1-24
1-24
1-24
1-24
1-24
1-25
1-25
1-25
1-26
1-26

Female
Female
Female
Female
Female
Male
Female
Female
Female
Male
Female
Male
Female
Female
Male
Female
Male
Female
Female
Female
Male
Male
Female
Female
Female
Male
Female
Female
Female
F:ema1e
Female

Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature

L-1924
L-1925
L-1926
L-1927
L-1928
L-1929
L-1930
L-1931
L-1932
L-1933
L-1934
L-1935
L-1936
L-1937
L-1938
L-1939
L-1943
L-1944
L-1945
L-1947
L-1948
L-1949
L-1950
L-1951
L-1952
L-1964
L-1954
L-1965
L-1966
L-1955
L-1956

31
34
46
32
47
None
51
None
52
23
53
None
54
50
None
19
None
55
72
77
None
None
76
None
56
None
None
79
78
75
None

28
28
2S
28
28
28
28
28
28
28
28
28
28
38
28
28
28
28
28
2S
2S
2S
2S
2S
28
38
28
28
3S
28
28

Location
Range. Sec.
96W
97W
97W
96W
96W
97W
97W
97W
97W
96W
96W
96W
96W
95W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
95W
97W
96W
95W
97W
97H

Remarks

19
25
25
30
30
25
24
24
24
30
30
30
30
11
25
25
25
24
24
25
24
24
24
24
24
12
25
34
12
25
25

--------------~--------------~-----------------------------------~------------------------------------~-------

I

N
N
\0
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management
(continued) .
Date
(1973)
1-26
1-26
1-26
1-27
1-27
1-27
1-28
1-28
1-28
1-28
1-28
1-28
1-28
1-29
1-29
1-29
1-29
1-29
1-29
1-30
1-30
1-30
1-30
1-30
1-30
1-30
1-30
1-30
1-30
1-31
1-31

Sex

Age

Eartag
No.

1\'e
ckband
:~o•

Twp.

Male
Male
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Hale
Female
Male
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Male
Male
Female
Male
Female

Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature

L-1957
L-1958
L-1959
L-1960
L-1961
L-1962
L-1969
L-1970
L-1971
L-1972
L-1973
L-1974
L-1975
L-1976
L-1977
L-1978
L-1979
L-1980
L-198l
L-1982
L-1983
L-1984
L-1985
L-1986
L-1987
L-1988
L-1989
L-1990
L-1991
L-1992
L-1993

45
48
17
74
90
89
None
88
60
73
71
87
68
None
86
39
92
93
None
None
102
None
103
None
104
110
42
None
95
41
112

3S
2S
2S
3S
3S
2S
3S
3S
3S
3S
3S
2S
2S
3S
3S
3S
3S
3S
2S
3S
3S
3S
3S
2S
2S
2S
2S
2S
2S
3S
3S

Unit 22, winter 1972-73

Location
Range
Sec.
96W
96W
96w
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96w
96w
96W
96W
96W
96W
96W
96w
96W
96w
96W
96W
96W
96W
96w

12
34
34
12
12
34
12
12
12
2
2
34
34
12
12
12
2
2
34
12
12
12
12
34
34
34
34
34
34
12
12

Remarks

May be neckband No. 86

I

tv

w

0
I

Maybe

neckband No. 89

---------------------------------------------------------------------------------------------------------------

�Table 1.

Record of deer trapped and marked on the ,",'Lnter
range in Game Management Unit 22, winter 1972-73

(continued).
Location

Date
(1973)

Age

Eartag
No.

Neekband

Sex

No ,

Twp.

-Range

Sec.

1-31
1-31
1-31
1-31
1-31
1-31
1-31
2-1
2-1
2-1
2...;1
2-1
2-1
2-2
2-2
2-2
2-2
2-2
2-2
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-4
2-4

Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Male
Female
Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female

Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Mature

L-1994
L-1995
L-1996
L-1997
L-1998
L-1999
L-2000
L-2014
L-2015
L-2016
L-2018
L-2017
L-2019
L-2020
L-202l
L-2022
L-2023
L-2024
L-2025
L-200l
L-2002
L-2003
L-2004
L-2005
L-2006
L-2007
L-2008
L-2009
L-20l0
L-20ll
L-2012

None
120
118
117
121
119
105
None
111
83
61
80
81
None
None
None
106
113
None
82
None
114
116
None
115
84
58
None
None
None
100

3S
3S
3S
3S
3S
3S
28
38
38
3S
3S
3S
38
2S
2S
3S
2S
3S
3S
3S
3S
3S
3S
3S
2S
2S
2S
2S
2S
3S
3S

96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

2
2
2
2
3
35
34
12
i2
2
3
3
35
34
34
3
34
2
12
12
12
2
3
3
34
34
34
27
27
2
2

Remarks

---------------------------------------------------------------------------------------~-----------~-----------

I

N

(,.0.)

~
I

�Table 1.
Record of deer trapped and marked on the winter range in Game Management Unit 22, winter 1972-73
(continued).
,-.
Date
(1973)
2-4
2-4
2-4
2-4
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-6
2-6
2-7
2-7
2-7
2-7
2-7
2-7
2-7
2-8
2-8
2-8
2-8
2-8
2-8
2-8
2-8
2-8

Sex
Male
Female
Male
Female
Female
Male
Female
Female
Female
Female
Male
Male
Male
Male
Female
Male
Female
Male
Female
Female
Hale
Female
Female
Female
Female
Female
Female
Male
Female
Female
Male

Age
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn

Eartag
No.
L-2013
L-2026
L-2027
L-2028
L-2029
L-2038
L-2039
L-2040
L-204l
L-204~
L-2043
L-2045
L-2044
L-2046
L-2047
L-2050
L-2049
L-2030
L-2031
L-2032
L-2033
L-2034
L-2051
L-2052
L-2053
L-2054
L-2055
L-2056
L-2057
L-2058
L-2059

},'eekband
No.
None '.
85
122
108
64
127
None
None
59
63
124
125
None
None
99
128
3
129
107
96
130
101
None
65
43
57
109
.44
49
97
None

Twp.
3S
38
28
28
38
38
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
38
38
)8

Location
Range
Sec.
96W
96W
96W
96w
96W
96W
96W
96w
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
96W
96w
96w

3
3
34
35
2
3
34
35
15
15
15
14
14
15
14
16
16
16
16
15
14
14
16
16
16
15
15
14
2
2
2

Remarks

I

N

w

N
I

Has a blue &amp; white neckband

---------------------------------------------------------------------------------------------------------------

�Table 1. Record of deer trapped and marked on the ",'Lnterrange in Game Hanagement Unit 22, winter 1972-73
(continued).

-,
Location
Range
Sec.

Date
(1973)

Age

Eartag
No.

Neckband

Sex

j~o.

Twp.

2-8
2-9
2-9
2-9
2-9
2-9
2-10
2-10
2-11
2-11
2-11
2-11

Female
Female
Male
Male
Female
Female
Female
Male
Male
Female
Female
Male

Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature

L-2060
L-2061
L-2062
L-2063
L...;2064
L-2065
L-2066
L-2067
L-1967
L-2078
L-2079
L-2080

98
70
67
69
66
None
91
126
94
None
123
None

3S
2S
2S
2S
2S
28
2S
28
2S
2S
28
28

8ummaEl:

20 Male Fawns;

22 Female Fawns ;

32 Mature Males;

96 Mature Females

65 Male Fawns;

67 Female Fawns;

90 Hature Males;

295 Mature Females

96W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W

Remarks

2
16
16
15
15
14
14
16
15
15
14
14

I

N

w

w
I

Grand
Summarz:

Corrections to 1972 TraEEing Records
(Corrections ~
Underlined)
2-1-72
2-1-72
2-1-72
2-1-72

Male
Female
Male
Male

Fawn
Mature
Mature
Fawn

L-1504
L-1520
L-1522
L-1524

27
48
81
78

2S
IN
IN
IN

98W
97W
98W
98W

12
8
1
1

White neckband
White neck band
White neckband
"White neckband

5
128

2S
IS

97W
97W

29
14

Pink neckband
Blue neckband

&amp; 1505
&amp; 1521
&amp; 1523
&amp; 1525
RecaEtures

12-30-72
2-17-73

Female
Female

Mature
Mature

L-1692
L-I722

�-234-

Table 2. Sightings of deer lleckballdedon the winter
Management Unit 22, through March 15, 1973.
Date
3-16-72
3-16-72
3-16-72
3-16-72
3-16-72
3-16-72
3-16-72
3-16-72
3-16-72
3-16-72
3-25-72
3-25-72
4-03-72
4-05-72
4-06-72
4-07-72
4-09-72
4-10-72
4-10-72
4-10-72
4-11-72
4-13-72
4-15-72
4-21-72
4-21-72
4-24-72
4-25-72
4-25-72
4-25-72
4-27-72
4-28-72
4-28-72
4-28-72
5-10-72
5-14-72
5-20-72
5-20-72
5-26-72
6-04-72
6-04-72
6-08-72
6-08-72
6-08-72
6-11-72
6-11-72
6-21-72

r;lI1ge

Neckband
Color
White
White
White
White
White
White
White
Blue
Blue
Blue
Green w/white stripe
Green w/white stripe
White
Green w/white stripe
White
Green w/white stripe
White
White
White
White
Green w/white stripe
Blue
Pink
White
White
Blue
Blue
Blue
Blue
White
White
Green w/white stripe
White
White
Green w/white stripe
Blue
Blue
Blue
Pink
White
Blue
Blue
Blue
White
Blue
Blue

Ntnnber
17
18
30
40
75
27
53
41
46
72
51
73
179
73

156
12
83

91
32
110

34
38

46
12

36
9

Twp.

2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
IS
IS
IS
IS
3S
2S
2N
IN
IN
IN
IS
IS
5S
2S
2S
IN
2N
IN
IN
4S
IS
2S
5S
2S
2S
2N
2N
6N
4S
2N
3N
3N
3N
2S
3N
2N

:In(:mne

Location
Range

Sec.

98W
98W
98W
98W
98W
97W
97W
98W
98W
98W
96W
96W
97W
96W
98W
95W
98W
96W
96W
96W
96W
95W
96W
97W
97W
97W
99W
97W
97W
96W
95W
95W
96W
94w
90W
93W
93W
90W
95W
93W
94w
94W
94w
91W
94W
88W

3
3
3
3
3
18
18
9
4
4
10
10
28
10
5
2
11
9
9
9
1
22
6
35
35
1
12
8
8
18
35
10
10
14
7
9
9
13
34
15
26
26
26
5
35
1

-------------------------------------------------------------------------------

�-235-

Table 2. Sightings of deer neckbanded on the winter range in Game
Management Unit 22, through March 15, 1973 (continued) .
Neckband
Date
7-01-72
.7-01-72
7-04-72
7-07-72
7-25-72
7-26-72
7-30-72
7-31-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
7- ?-72
8-01-72
8-10-72
8-12-72
8-20-72
8-26-72
8-27-72
8-28-72
8-28-72
8-28-72
8-29-72
9-10-72
9-10-72
9-27-72
9- ?-72
10-6-72
10-6-72
10-6-72
10-6-72
10-10-72
10-10-72
10-10-72
10-14-72
10-14-72
10-14-72
10-14-72
10-14-72
10-14-72

Color
Blue
White
Blue
White
White
Blue
Blue
White
White
White
Blue
Blue
White
White
White
White
White
White
White
White
White
Green w/white stripe
White
White
Blue
Blue
White
White
White
White
Blue
Blue
Blue
Blue
White
White
White
White
Blue
Blue
White
Green w/white stripe
Blue
White
White
Blue
Blue

Number

Twp.

IS
4S

IN
100
81

112
161

12

43

39

IN
IS
3N
3N
3S
4S
4S
2N
5S
55
5S
5S
5S
5S
5S
5S
5S
IN
2S
2S
7S
2N
4N
2N
58
5S
5S
2S
2S
3N
2S
48
5S
6S
48
2N
2N
2N
IN
4S
IS
IS
2S
2S

Location
Range

Sec.

94W
97W
89W
90W
9lW
94W
93W
99W
97W
98W
86W
97W
97W
97W
97W
97W
97W
97W
97W
97W
94W
90W
90W
97W
94W
95W
94W
94W
94\-1
94W
90W
90W
99W
94W
95W
96W
97W
96W
98W
98W
98W
96W
94W
99W
99W
91W
91W

36
27
1
19
7
27
7
22
29
26
4
17
4
15
15
29
21
21
21
21
25
7
23
24
12
27
12
28
29
26
23
23
33
1
23
29
17
29
3
3
2
36
4
30
30
18
18

-------------------------------------------------------------------------------

�-236Table 2. Sightings of deer neckbanded on the winter range
in Game
Management Unit 22, through March 15, 1973 (continued).
Neckband
Date
10-14-72
10-14-72
10-14-72
10-14-72
10-14-72
10-14-72
10-14-72
10-15-72
10-15-72
10-15-72
10-15-72
10-15-72
10,-15-72
10-15-72
10-15-72
10-15-72
10-15-72
10-15-72
10-15-72
10-15-72
10-16-72
10-16-72
10-16-72
10-16-72
10-16-72
10-16-72
10-16-72
10-16-72
10-16-72
10-16-72
10-16-72
10-16-72
10-17-72
10-17-72
10-17-72
10-17-72
10-17-72
10-17-72
10-18-72
10-19-72
10-20-72
10-20-72
10-21-72
10-21-72
10-21-72
10-22-72
10-22-72
10-22-72
10-22-72

Color
White
White
White
White
White
White
White
White
White
White
White
White
Green w/white stripe
White
White
White
White
Blue
Blue
Green·w/white stripe
Green w/white stripe
White·
White
White
White
White
White
White
Green w/white stripe
White
White
Blue
White
Blue
White
White
Blue
White
Green w/white stripe
White
White
Blue
White
Blue
Pink
White
White
White
White

Number

115

39

Twp.

4S
2S
IS
3S
3N
IS
3N
2N
IN
2N
2N
2N
2N
2S
IS
IN
4S
3S
3S
IS
IN
IS
IS
IS
2S
4S
2S
IS
2S
IS
3S
IN
IS
IS
IS
IS
2N
2N
3S
5N
IS
4N
5N
5N
4S
4N
IS
IS
IS

Location
Range

96W
100W
99W
99W
95W
100W
95W
96W
96W
98W
98W
98W
94W
100W
99W
93W
95W
95W
95W
95W
96W
100W
100W
100W
97W
99W
95W
100W
95W
99W
96W
loOW
97W
99W
99W
99W
98W
98W
96W
91W
96W
90W
96W
93W
94W
92W
97W
97W
97W

Sec.

26
36
17
18
18
36
16
24
30
32
32
32
15
35
17
25
34
36
36
10
30
35
35
35
15
2
14
36
28
17
26
14
21
17
17
8
30
10
15
31
3

27
35
3
6
24
16
16
16

--------------------------------------------------------------------------------

�-237Table 2. Sightings of de~r ncckbanded on the winter range in Cnme
Management Unit 22» through March 15, 1973 (c on t Lnuc-d ) •
Neckband
Date

Color

10-22-72
10- ?-72
10- ?-72
11-12-72
11-16-72
11-21-72
11-22-72
11-22-72
11-22-72
11-24-72
11-29-72
12-07-72
12-12-72
12-13-72
12-13-72
12-14-72
12-14-72
12-14-72
12-14-72
12-15-72
12-15-72'
12-22-72
12-22-72
12-22-72
12-23-72
12-23-72
12-26-72
12-27-72
12-27-72
12-27-72
12-27-72
12-27-72
12-27-72
12-27-72
12-27-72
12-27-72
12-27-72
12-27-72
12-28-72
12-28-72
12-28-72
1-06-73
1-06-73
2-03-73

White
White
White
White
Green w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
White
White
Blue
White
Blue
Green w/white stripe
Greenw/white stripe
White
White
Pink
White
White
Green w/white stripe
White
White
Pink
Pink
Green w/white stripe
White
White
White
White
White
White
White
White
White
White
White
Pink
Pink
Pink
White
White
Green w/white stripe

Number

49
47
13
14

44

41
40
12
15

47

Twp.

IS
6S
3S
2N
IS
IS
IN
IN
18
2N
2N
2N
2N
2N
IN
IN
28
28
2S
2S
IS
IS
28
IN
28
28
IN
18
IS
18
18
18
IS
IS
18
IS
IS
IS
2S
2S
2S
2S
IS
IS

Location
Range

Sec.

97W
95W
97W
98W
96W
96W
96W
96W
99W
97W
97W
99W
97W
99W
96W
96W
98W
98l.o1
98W
97W
98W
96W
99W
97W
97W
97W
96W
97W
97W
97W
98W
98W
98W
98W
98W
98W
98W
98W
97W
97W
97W
98W
97W
96W

16
28
21
4
10
14
16
16
21
20
19
11
28
35
28
26
12
12
12
6
8
11
26
21
20
20
28
29
21
16
36
8
8
8
8
8
8
8
29
29
29
5
17
11

------------------------------------------------------------------------------

�-238-

Table 2. Sightings of deer neckbanded on the winter range in Game
Management Unit 22, through March 15, 1973 (continued).
Neckband
Date

Color

2-08-73
2-08-73
2-19-73
2-26-73
2-26-73
2-26-73
2-26-73
2-26-73
2-28-73
3-04-73

Green w/white stripe
Green w/white stripe
Blue
Blue
Blue
Blue
Green w/white stripe
Blue
Blue
White

Ntnnber
13
128
118
131
36
132
54

Twp.

IS
IS
3N
IS
IS
IS
IS
IS
IS
IN

Location
Range

Sec.

96W
96W
98W
97W
97W
97W
96W
97W
96W
97W

4
4
31
14
14
14
11
14
9
14

�-239-

Recoveries
of deer neckbanded
Table 3.
ment UnLt; 22, through Harch 17, 1973.

Neckband
Color

Date of
Recovery

4-6-72
4-27-72
5-25-72
6-21-72
8-29-72
10-14-72
10-19-72
10-22-72
10-19-72
10-15-72
10-14-72
10-15-72
10-15-72
10-14-72
10-17-72
10-18-72
10-16-72
10-16-72
10-14-72
10-15-72
10-14-72
10-16-72
10-14-72
10-14-72
10-17-72
10-?-72
10-?-72
10-?-72
10-?-72
10-?-72
10-?-72
10-?-72
10-?-72
10-?-72
10-?-72
3-9-73
3-17-73

White
White
Green
Green
White
Green
Green
White
White
White
Green
White
White
White
White
Green
White
White
Green
Green
White
Green
White
White
White
Green
White
White
White
White
Blue
White
Blue
White
White
White
Blue

w/white
w/white

stripe
stripe

w/white
w/white

stripe
stripe

w/white

stripe

w/white

stripe

w/white
w/white

stripe
stripe

w/white

stripe

w/white

stripe

on

the

w Ln t.c r

No.

Twp.

Location
Range

Sec.

15
70
37
35
171
76
38
123
143
119
57
25
124
137
68
1
165
11
40
32
79
18
205
97
42
76
90
8
120
76
10
158
71
100
91
54
129

2N
2S
IN
2S
lS
2N
IS
IS
5S
1S
IS
2N
2S
3S
1N
3S
3S
3N
IN
IN
3S
2S
IN
IN
3S
2N
2S
IN
3S
?
5N
IS
2N
2S
3N
IN
1N

98W
97W
96W
95W
98W
94W
95W
97W
98W
98W
95W
98W
97W
97W
97W
96W
97W
96W
93W
96W
95W
95W
99W
97W
95W
94W
96W
94W
99W
?
90W
97W
93W
97W
91W
97W
97W

13
9
26
2
33
28
29
20
2
4
20
32
23
5
29
23
7
30
30
27
30
14
29
31
30
20
16
35
34
?
32
16
22
36
16
14
15

r an ge

in

Came

~tmage-

Remarks

Died soon after
Died soon after
Died soon after

tagging
tagging
tagging

Died soon after

tagging

Died soon

after

tagging

Strawberry

Creek Area

Died soon after

tagging'

�GAME

MANAGEMENT UNIT

22

..... •..

-PICEANCE-

~

l

r
)------

I

I

~
o

\

.f:"

n
u

I

I

\~\
••

LEGEND
=~",",," Heavy use

~:::::::::::;::l Moderate use
!·.·.·.·t·.·.·.·~~
,-----'

i

- I
I

Fig. 1.

Little or no use

Deer distribution

on the Piceance winter range, December 6, 10 and 11, 1972.

�GAME

MANAGEMENT UNIT

22

-PICEANCE-

~-•..•

r
)------

I

r-&gt;
.p0-

I

I-'

~

n
u

I

I

\--\

\ 1.

"I
\

\

''\

LEGEND
Heavy use
~"":lM
t:::.d~ 0d erat e use

'"' "I--\,.
" :r'---•...

-"

c====JLittle or no use

I,

,..•..

Fig. 2. Deer distribution on the Piceance winter range, January 5, 6 and 7, 1973.

�~

.-_ .... - .._.. . ..:..

GAME

MANAGEMENT UNIT

22

-PICEANCE-

l
I

N

.j::-o
N
I

Fig.

J.

Deer distribution

on the Piceance

winter

r-ange , February

19 and 26-, 197J.

�July,

-243-

JOB PROGRESS

S ta te of
Project

REPORT

.;:::C.;:::OL;:::,O::::,;'
RA:.:;.;:;:D::..;O:;.'
:.Deer-Elk

W-38-R-27

No.

16

Job No.

Job Title

Piceance

Deer Study - Population

Period

April

Work

1973

Plan No.

Covered:

Personnel:

1, 1972 through March

Investigations
2

Density

and Structure

31, 1973

R. M. Bartmann, S. F. Steinert, J. J. Klein, Jr., D. Hoart, J. Ellenberger, C. H. Gore, T. A. Lytle, C. W. Reichert, C. H. Roberts and
D. J. Todd.

ABSTRACT

A helicopter count of deer on the Piceance winter range was made on two sizes
of quadrats on December 21 and 22, 1972; ~ square mile and ~ square mile.
The
smaller quadrat was selected for future counts as the loss in precision from the
larger quadrat was not considered great and, subjectively,
it could be counted
more efficiently.
The mean number of deer on 60 ~ square mile quadrats was 7.4,
or 29.6 deer per square mile.
The projected deer population estimate with 90
percent confidence limits was 19,684 + 3,990.
Based on sample size estimates,
120 quadrats will be counted next year.
Combined ground and air sex and age
classification
data gave a buck:doe:fawn
ratio of 28:100:74.
The sex and age
breakdown of the Piceance deer population was estimated at 2,756 adult males,
2,748 yearling females, 6,995 mature females, 3,945 male fawns, and 3,240 female
fawns. A range of sample sizes at different prevision l~vels were calculated
from preliminary pellet group count data from 200 100 ft circular plots per
section on 12 sections of land on the Piceance winter range.
The major problem
anticipated in field application of a large-scale pellet group survey is getting
enough dependab Le manpower.
"

��-245-

PICEANCE

DEER STUDY - POPULATION

DENSITY

AND STRUCTURE

Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer
population density and structure data on pinyon-juniper winter range
for use in making annual harvest recommendations.

SEGMENT OBJECTIVE
To establish mule deer population density and sex and age structure
estimate techniques for the Piceance ~inter range.
la.

Evaluate

the use of a helicopter

for estimating

lb.

Estimate
levels.

quadrat

2a.

Evaluate
density.

the pellet group c9unt technique

2b.

Assist Field Services personnel in establishing a pellet
group survey on the Piceance winter range so that problems
arising in the field application of this technique can be
identified and remedied or considered for further research.

3a.

Evaluate the "key browse" concept of forage production-utilization surveys used for deer pOpulation trend assessment.

3b.

Estimate species composition of forage consumed
by deer from stomach content analyses.

sample size requirements

METHODS

deer density.

at various

precision

for estimating

deer

during winter

AND MATERIALS

Deer Density
In February, 1972, trial helicopter counts of deer on two sizes of quadrats
(1/4-square-mi1e and 1/16-square-mile) were made on the Piceance winter
range.
From subjective evaluation, the larger quadrat was selected for
future counts.
As no work was done with the possibility of using an even
larger quadrat, this was incorporated into the 1972-73 winter count.
Sixty 1/4-square-mile
(1/2 X 1/2-mile) quadrats were randomly selected on
the Piceance winter range (Fig. 1). A second 1/4-square-mile quadrat was
placed adjacent to the first on the cardinal side, within the section, that
would provide the greatest elevational relief.
This produced an over-all
quadrat of 1/2 X I-mile.
The four outside corners and the two common
corners of each set of quadrats were marked with 6 X lO-inch orange markers.
Markers were placed in tree tops when possible or on tall shrubs or fence
posts.

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~,

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Two ~-square-m1.l
IiiiICensus
Quadrats

\

....

... /
,~/

/.......

\ \ •..

C Pellet

/

,,/ff"J
f."/

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:~;:~,;;;---/

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,.~

' l!iii"~------."

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~

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r"~/

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i I
Fig. 1.
Piceance

Locat i.ons of dear census
winter range, 1972.

quadrats

And preliminary

pellet

group count

sections

on the

�-248-

Each set of quadrats was counted as one but deer observed were recorded
separately for each half of the 1/2-mile quadrat.
Deer were assigned
to the half they were first seen in. Two observers were used; one to
count and the other to count and navigate.
Aerial photos of each
quadrat were carried in the helicopter to help insure staying within
the plot bounds.
I

Population

Structure

The composition of the winter deer population was estimated from postseason sex and age classifications made from the ground.
Additional
data from the southern one-third of Game Management Unit 22 were obtained
from post-season helicopter classifications made by Northwest Region
personnel.
Check station data from the 1972 deer season were used to
estimate fawn sex ratios and percent yearling females.

Pellet Group Count
A preliminary deer pellet group count was made on the Piceance winter
range to help Field Services personnel get information upon which to
base sample size projections and to identify problems that might be
encountered in a large-scale survey.
Twelve sections of land were
randomly selected on the winter range (Fig. 1). Each section was divided
into four 1/4 X I-mile segments with the long axes oriented in the cardinal
direction giving the greatest elevational change.
Four one-mile-long
transects were then randomly selected, one in each segment.
Fifty 100 ft2
circular plots were systematically spaced about 100 feet apart, by pacing,
along each transect for a total of 200 plots per section.
Two-man crews
followed the guidelines for pellet group counting given by Smith, Neff,
and McCulloch (1969).

Food Habits
No work was done on this job during the segment.
The student assistant
assigned this work resigned before doing any analyses.

RESULTS AND DISCUSSION

Deer Densitx:
A helicopter count to determine deer density was made December 21 and 22,
1972. This was probably earlier than will occur most years, but heavy
snows pushed deer down within the pre-determined winter range limits set
for the count.
Snow background conditions were considered excellent for
counting.
Four hundred forty-four and 434 deer were counted on each set of 1/4-squaremile quadrats for a mean of 7.4 and 7.2 deer per quadrat (Table 1). Accordingly, the large quadrats contained 878 deer or 14.6 deer per quadrat.
The

�-249-

coefficient of variation decreased an average 19 percent from the larger
This relatively small gain in precision was not
to the smaller quadrat.
considered sufficient to warrant using the larger plot.
Subjective evaluation of counting efficiency also favored the smaller
quadrat.
The rough and varied topography of the winter range made it
difficult to keep oriented in a 1/2-square-mile area which, together
with the tree overstory, increased the chances of missing deer.
The possibility of increasing sampling efficiency through stratification
was considered.
On the basis of two years' deer distribution data, the
winter range was divided into high and low deer density areas.
The
original set of 60 1/4-square-mile quadrats were then assigned to the
proper strata.
Twenty-six were located in the low density area and 34
in the high density area.
Stratification produced no increase in preClslon over the original random
sample.
The standard error of the mean of the stratified sample (0.89)
was the same as that of the random sample (Snedecor 1956:507).
Therefore,
unless additional information shows otherwise, a random sampling design
will be used for density estimates.
Sample size estimates were made using the formula n = t2s2/d2; where t =
the tabular "t" value at a selected confidence level, s = the sample
standard deviation, and d = the allowable error in the estimate (Snedecor
1956:501-502).
Separate sample sizes were estimated using data from each
set of 1/4-square-mile quadrats.
Acceptable precision was ± 15 percent of
x at the 90 percent confidence level.
Sample sizes of 107 and 146 plots
were indicated and a sample size of 120 plots was selected.
Data from the original set of 1/4-square-mile
quadrats yielded an estimate
of 29.6 deer per square mi.le. The total winter range area measured from
USGS topographic quadrangle maps was 665 square miles.
At the 90 percent
confidence level, the total winter population was estimated at 19,684
± 3,990 deer.
This estimate is a minimal one as the probability of missing
animals is considered greater than that of recounting them. This is particularly true of this count as the relatively large 1/2 X I-mile quadrat
proved cumbersome and a greater than "normal" number of animals were probably missed.
The magnitude of this error cannot be estimated at this time.

Population

Structure

Post-season sex and age classifications made from the ground November 1730, 1972, resulted in 1,712 deer being classified.
This work was done
entirely from vehicles and complete coverage of the winter range was not
achieved although most areas with substantial deer concentrations were
sampled.
Post-season helicopter classifications made by Northwest Region
personnel added another 724 deer.
The combined ground and air data gave a buck:doe:fawn ratio of 28:100:74
(Table 2). Check station data for Game Management Unit 22 show that of
328 fawns examined, 54.9 percent were males and 45.1 percent were females.
Of 817 does older than fawns examined at check stations, 28.2 percent were
classed as yearlings by tooth replacement.
Incisors of females older than
fawns were collected for aging by aental cementum (Erickson and Seliger
1969), but this information is still unavailable.

�-250-

Table l. Numbers of deer counted on
\-square-mile quadrats randomly
located on the Piceance winter range during the 1972-73 winter.

Quadrat

Number of Deer
First
Second
Both
Half
Half
Halves

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35

1
11
10
7
1
0
17
0
0
4
10
13
0
0
6
4
0
14
5
1
5
4
11
2
21
14
13
23
13
4
14
17
0
0
0

6
9
10
4
1
4
15
4
14
10
3
2
7
5
2
3
0
0
0
4
2
2
0
12
19
34
26
13
0
17
20
25
0
3
0

7
20
20
11
2
4
32
4
14
14
13
15
7
5
8
7
0
14
5
5
7
6
11
14
40
48
39
36
13
21
34
42
0
3
0

Quadrat

First
Half

Second
Half

Both
Halves

36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60

0
1
0
9
2
26
4
19
6
13
7
18
0
3
16
0
10
4
12
16
1
9
0
14
9

6
4
0
8
12
15
14
3
6
3
0
1
1
1
7
0
3
10
1
25
2
17
17
0

6
5
0
17
4
38
19
33
9
19
10
18
1
4
17
7
10
7
22
17
26
11
17
31
9

444

434

878

~

X

x

=

s
Coeff.
Var. =

2

7.4

7.2

14.6

6.97

7.93

12.07

94%

110%

83%

�-251-

Table 2. Post-season deer sex and age classification
the Piceance winter range during the 1972-73 winter.

count results

on

Type of Count

Date

(1972)

Bucks

Does

Fawns

Total

Ground

Nov. 17-30

267

822

623

1,712

74

384

266

724

341

1,206

889

2,436

Buck:Doe:Fawn
Helicopter

Ratio

32:100:76

Dec. 27
Btick:Doe:Fawn Ratio

19:100:69

All Counts
Buck:Doe:Fawn

A summary
below:

Ratio

28:100:74

of the 1972 deer population

Area of winter

range in Game Management

Total adult males

2,756

Total

9,743

adult females

Yearling
Mature
Total

females
females

fawns

and structure

Unit 22

data is given

665 square miles.

2,748

=

6,995

=

7,185

Male fawns

3,945

Female

3,240

fawns

density

Pellet GrauE

Count

The pellet group count was made from April 24 through May 15, 1972.
The
range in mean pellet group densities was from 0.15 to 1.44 groups per plot
with an overall mean of 0.91 groups per plot (Table 3).
Sample sizes were estimated for both the number of plots per section and
the number of sections.
The formula n = t2s2/d2 was again used in both
cases.
The calculated sample sizes at various prec Ls Lon levels are shown
as follows:

�Number of Plots per Section
± X% of the Hean

__

+-5

10

15

20

95%

I 2945

755

335

188

:&gt;
~ ...~

90%

2078

533

236

l33

u

80%

1258

322

143

81

(!)
()

~
(!)

'""d
-r-i
'H

r-i
OJ

(!)

0

Number of Sections
± X% of the Mean

OJ

o

$:l

~5%

OJ
'""d

.-I

'H

(!)

I

5

10

15

20

243

61

27

16

171

43

19

11

104

26

12

7

I

Ql

or! i&gt;
~ H

90%

u

80%

0

I
I

As expected, estimated sample sizes indicate that a full-scale pellet
group survey in Game Management Unit 22 would be a major efforto
However,
the statistical aspect is not necessarily the main limiting factor.
Rather, experience in conducting the preliminary survey points to dependable rnan+powaj- as the key limitation.
Although it was possible to
get the requested number of people for the start of the survey, no help
was available following a few days delay because of a snowstorm.
This
is, of course, not a fault of the survey itself.
The high pellet group density found in the preliminary survey probably
stems, in part, from locating plots by pacing and mistaking old groups
for new. Locating permanent plot centers with chain and compass would
be a prerequisite to clearing plots which, together, should reduce bias
from both sources to a minimum.
Initial locating and marking of plots
would be a major one-time effort while clearing would be an annual chore.
Ideally, clearing plots should be done in the fall and would at least
double the overall workload and the associated man-power problems.
The starting and ending times were recorded for each 50-plot transect.
This included counting, but not clearing, pellet groups and pacing between plots.
The range in times was from 40 to 165 minutes with a mean
of 75 minutes.
The shorter times are overly fast and are probably
attributable to the general lack of experience by most participants.

�2
Table 3. Results of deer pellet group counts on 200 uncleared 100-ft circular plots per section on
the Piceance winter range during spring, 1972.
Number of Plots Containing X Number of GrouEs
10
8
9
6
7
4
5
3

Section

Total Pellet
Groups/Four Transects

1

30

178

16

4

2

2

85

133

53

11

2

1

3

140

116

50

20

8

4

2

4

170

101

54

28

10

5

2

5

173

113

50

13

11

4

7

1

1

6

185

100

49

28

17

4-

1

-

-

X =

0

1

2

1
I

-

-

1

-

1

2

1

1

4

4

1

1.27

Coefficient Var. =

140%

s = 72 .10

Coefficient Var. =

-40%

7

191

96

59

24

9

6

3

3

-

-

8

194

105

45

21

17

7

3

1

1

-

9

222

87

56

31

15

3

3

4

-

1

10

240

80

59

29

17

6

6

3

11

261

72

57

41

14

9

4

1

-

12

288

80

51

27

17

10

5

6

Totals

2,179

1,261 599

277

139

59

36

19

s =

x pellet groups/plot

=

0.91

x pellet groups/four transects = 181.6

1

N
VI
W
I

�-254-

Clearing plots would, of course~ have added considerable extra time.
A two-man crew might reasonably expect to cover about 100 plots per
day including travel time if plots were both counted and cleared, and
probably two to three times this many if plots were only counted.
No attempt was made to convert pellet group density to deer density
because of problems mentioned above. Methods for making this conversion are described by Smith, Neff, and McCulloch (1969). One of
the key components in this conversion, the mean daily deer defecation
rate, seems warrant of additional research to estimate annual variability and factors affecting it if pellet group data are to be used
for deer management.

LITERATURE

CITED

Erickson, J. A., and W. C. Seliger.
1969. Efficient sectioning of
incisors for estimating ages of mule deer. J. Wildl. Mgmt.

33(2):384-388.
Smith, R. H., D. J. Neff, and C. Y. McCulloch.
1969. A model for
the installation and use of a deer pellet group survey. Arizona
Game and Fish Dept.
Spec. Rept. No. I. 30 p.
Snedecor, G. W. 1956. Statistical
Press, Ames.
534 p.

Prepared by
Richard M. Bartmann
Wildlife Researcher

methods.

The Iowa State College

�July, 1973

-255-

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~--------~----

Project No.

W-38-R-27

Work Plan No.

16

Deer~Elk Investigations
job No.

3

Job Title

Piceance Deer Study - Productivity and Mortality

Period Covered:

April 1, 1972 through March 31, 1973

Personnel:

R. M. Bartmann, S. F. Steinert, J. J. Klein, Jr., L. Crooks, R. Stone
and D. Talcott.

ABSTRACT

Pre-natal productivity data in 1973 were collected, all or in part, from 65 does
killed on highways and during trapping operations in and around Game Management
Unit 22. Productivity was generally similar to that in 1972. Yearlings had
1.0 CLP:doe and 100 fetuses:100 does. Mature animals had 1.9 CLP:doe and 163
fetuses:lOO does. The fetal sex ratio was 96 males:lOO females. The percent of
does pregnant was 86 which is 9 percent lower than in 1972. The peak of conceptions was estimated from December 6-10, 1972. The mean date was December 4. Some
problems are suspected in application of the fetal growth curve to Piceance deer.
Data for 13 does found dead in fences suggest these animals were generally older and
less productive than road-kills and trap mortalities.

��-2':/7-

PICEANCE DEER STUDY - PRODUCTIVITY

AND MORTALITY

Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.

SEGMENT OBJECTIVE
1.

To estimate increments and losses to the Piceance deer population,
a sub-population thereof, to allow evaluation of density estimate
techniques.
a.

To estimate

productivity

METHODS
See Bartmann

or

rates.

AND MATERIALS

(1972).

RESULTS AND DISCUSSION
The 1973 pre-natal productivity data were collected, all or in part, from
65 does killed in and around Game Management Unit 22. Most were road-kills,
some trap mortalities, and two were from miscellaneous sources.
Another
13 does were found dead in fences and are considered separately.
Animals
were aged as yearlings or mature by tooth replacement.
A breakdown by year
class will be available later from ages assigned by the dental cementum technique (Erickson and Seliger 1969).
Ovulation rates were estimated from counts of corpora lutea of pregnancy
(CLP) in ovaries of 59 does. Yearlings averaged 1.0 CLP and mature does
1.9 CLP which is the same as in 1972 (Table 1). The overall conception rate,
based on 65 does, was 86 percent or 9 percent lower than in 1972.
Fetus:doe ratios, also based on 65 does, were about the same as in 1972.
Yearlings had 100 fetuses:lOO does and mature animals 163 fetuses:lOO does.
All yearlings in previous years were pregnant with one fetus each.
In 1973,
11 yearlings had singletons, four had twins, and four were barren.
The fetal
sex ratio was nearly even, 96 males:lOO females.
Most fetuses appeared
skinny compared to those in previous years.
This is purely subjective as no
measurements were taken that would reflect this, but if true, it could be
attributable to poor nutrition during the severe winter.

�-258..

Table l. Pre-natal productivity data for mule deer does dying
from
various causes in and around Game Management Unit 22,
1973.
Call.
No.

Est.
Age

Date
Killed
(1973)

How
Killed

P-72
P-73
P-74
P-75
P-76
P-77
P-78
P-80
P-81
P-82
P-83
P-84
P-85
P-86
P-87
P-88
P-89
P-90
P-91
P-92
P-93
P-94
P-95
P-96
P-97
P-98
P-99
P-100
P-10l
P-102
P-103
P-104
P-105
P-108
P-109
P-1l4
P-115
P-116
P-1l7
P-118
P-119
P-12l
P-123
P-125
P-127
P-128
P-129

Mature
Mature
Mature
Mature
Yr1g.
Mature
Yr1g.
Yr1g.
Yrlg.
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Yr1g.
Mature
Mature
Yrlg.
Yr1g.
Mature
Mature
Nature
Mature
Yrlg.
Mature
Mature
Yrlg.
Yrlg.
Yrlg.
Mature
Mature
Mature
Mature
Mature
Yr1g.
Yrlg.
Yrlg.
Yr1g.
Mature
Mature
Mature
Yr1g.
Mature

1-19
1-21
1-27
1-24
2-01
2-08
2-08
2-19
2-19
2-18
2-15
2-21
2-22
2-22
2-25
3-01
3-05
3-05
3-09
3-14
3-14
3-14
3-14
3-18
3-21
3-21
3-21
3-24
3-24
3-24
3-24
3-28
3-28
3-28
3-28
3-31
3-31
3-31
4-04
4-04
4-04
4-04
4-04
4- 07
4-11
4-11
4-11

Vehicle
Vehicle
Trap
Trap
Vehicle
Trap
Trap
Vehicle
Trap
Trap
Shot
Blind
Trap
Trap
Vehicle
Vehicle
Trap
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle

No. of Fetuses
Male
Female
1
0
2

0
1
0
1?
1?

1
0
1
0
2
1
1
1
1
1
0

1
1
0
1
1
0
0
1
0
0
2
1
0
0
0
0

1
1
0
2

1
1
0
0
0
1
1
0
0
1
0
1
1
2
1
1
1
1
1
2
2
0
0
2
0
1
1
0
1
0
0

0

0

1
0
0
0
0
1
1
0
2
1

1
0
1
0
0

1
1
2
0
1

Corpora Lutea
of Pregnancy

1
2
4
2
1
2
1
1
0
2
2
2
2
2
2
0

4
2
2
2
1
1
2
2
2
1
2
0

1
1
1
2
0

0
1
0
0
2
2
2
2
2

-------------------------------------------------------------------------------

�-259-

Table I. Pre-natal productivity data for mule deer does dying from
various causes in and around Game Management Unit 22, 1973 (continued).

ce u.
No.

Est.
Age

Date
Killed
(1973)

How
Killed

No. of Fetuses
Female
Male

P-13l
P-133
P-134
P-135
P-136
P-138
P-139
P-140
P-14l
P-143
P-144
P-145
P-146
P-149
P-150
P-151
P-152
P-153

Mature
Yrlg.
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Yrlg.
Mature
Mature
Mature
Mature
Yrlg.

4-14
4-14
4-14
4-14
4-14
4-18
4-21
4-24
4-25
4-25
4-25
4-27
4-30
5-04
5-05
5-06
5-08
5-13

Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle
Vehicle

1
1
1
1

100 fetuses:lOO yearling does
163 fetuses:lOO mature does
145 fetuses:lOO does
96 male fetuses:lOO female fetuses

2

1
1
1
1
0
0
1
0
1
0
2
0
1

Corpora Lutea
of Pregnancy

1
1
1
1
0
1
0
1
1
2
2
1
2
1
1
0
0
0

2
2
2
2
2

2
2
2
2

2
2

2
2

3
2
0
1

1.0 CLP:yearling doe
1.9 CLP:mature doe

Percent pregnant

86%

The 13 does found dead in fences appeared older and less productive than road-kills and trap mortalities. Therefore, productivity data for them were
summarized separately (Table 2). These suspicions are strengthened by comparison of data for both groups. All fence-kills were estimated as mature
with many aged as 9 years or older. Also, productivity was lower according
to ovulation, pregnancy, and fetal rates. It seems that inclusion of fencekills with the other sources of mortality could bias productivity data.
Whether or not road-kills and trap mortalities also provide biased information
is not known, but will be checked as soon as additional data become available.
Seventy-three fetal ages were estimated from forehead-rump 'measurements using
Hudson and Browman's (1959) growth curve. Conception dates were estimated by
back-dating from date of death. The range in conception dates was from
November 12, 1972, to January 9, 1973 (Fig. 1). The peak number of conceptions
occurred from December 6 to 10, 1972. The mean date was December 4 which
was about the same as in 1971 and 9 days later than in 1972.

�-263-

July, 1973

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~--------------------

Project No.

W-38-R-27

Deer-Elk Investigations

Work Plan No.

17

Job Title

Systems Modeling Big Game Populations

--------~------------------------

Period Covered:
Personnel:

Job No.

1

September 1, 1972 through March 31, 1973

Allen E. Anderson, Ronald G. Blumberg, Raymond J. Boyd, Harold E.
Burdick, V. Robert Clark, Harvey Donaho, Marcus P. Elkins, R. Bruce
Gill, James F. Lipscomb, Richard L. Norman, Perry D. Olson and
LareriA. Roper.

ABSTRACT

Simulation models were developed for the White River elk herd (Area E) and the
Uncompahgre deer herd. A method was developed and programmed in Fortran IV for
calculating expected values for herd characteristics under sustained yield management based on information gained through simulation modeling. The computer
program is used to produce nomograph plots.

��-265-

SYSTEMS MODELING BIG GAME POPULATIONS
James F. Lipscomb

P. S. OBJECTIVE
To construct workable and usable systems models of big game populations.

SEGMENT OBJECTIVES
1.

Identify and select one deer and one elk herd which have sufficient
information for modeling purposes.

2.

Construct production models which will simulate the output of these
populations.
METHODS AND MATERIALS - RESULTS AND DISCUSSION

Herd Selection Criteria
Population information currently available on Colorado elk and deer herds
was collected from regional biologists and researchers A. E. Anderson,
R. J. Boyd, R. B. Gill and L. A. Roper. The big game herds which were
considered to be candidates for modeling were Middle Park, Poudre and
Uncompahgre deer, and San Juan, Rio Grande and White River elk.
The White River elk herd, with population data collected under Work Plan
11 of this project continuously since 1958, was judged to have the best
information of any herd for the purpose of systems modeling. This herd
was chosen as the first to be modeled.
The Uncompahgre deer herd was selected for modeling based on four years
of population data. The primary reason for this choice was that the data
on this herd had been collected by regional biologists using regular game
management procedures rather than under an intensive research project.
If modeling is to be a usable tool it must be capable of working with the
kinds of data that can be collected on a management basis.
Modeling Procedures
Trial and error adjustment of natural mortality rate estimates for the Area
E elk herd produced the simulation summarized in Table 1, which shows close
agreement with the observed sex ratio values over the period 1965-1972.

�-266-

Table l. Simulated population dynamics of
the White River (Area E)
elk herd - prehunt 1966 to prehunt 1972.

Prehunt 1966

No.
Bulls

No.
Cows

No.
Calves

Total

1,040

3,375

1,944

6,359

31

29

Ma1es/100 Females
Simulated
Observed

Harvest

719

578

118

Posthunt 1966

32l

2,797

1,826

4,944

11

10

3,301

1,964

6,308

32

32

Prehunt 1967

1,042

Harvest

619

623

135

Posthunt 1967

423

2,678

1,829

4,931

16

16

3,191

1,928

6,263

36

37

Prehunt 1968

1,143

Harvest

800

781

174

Posthunt 1968

343

2,410

1,754

4,508

14

16

2,916

1,758

5,707

35

50

Prehunt 1969

1,034

Harvest

610

488

85

Posthunt 1969

424

2,428

1,673

4,524

17

16

2,907

1,592

5,577

37

33

Prehunt 1970

1,077

Harvest

866

453

41

Posthunt 1970

211

2,454

1,551

4,217

9

9

2,891

1,802

5,504

28

31

Prehunt 1971

812

Harvest

429

267

37

Posthunt 1971

383

2,624

1,765

4,772

15

15

3,124

2,028

6,230

35

49

Prehunt 1972

1,078

�-267-

Simulations were limited to this period when it became apparent that
harvest figures for these years (based on a random survey of 50 percent of the hunters) were considerably better than those of previous
years.
This simulation is based on the assumptions that harvest data
and cow:calf counts are accurate and that natural mortality rates are
relatively constant over the period of the simulation.
Given these
assumptions, natural mortality rates and 1965 population size were
varied to obtain a simulation that provided close agreement with the
observed sex ratio data. Annual natural mortality rates used in this
simulation were 0.19 on calves, 0.02 on yearlings, 2 year olds and 3
year olds, 0.03 on 4 year olds, 0.07 on 5 year olds, 0.15 on 6 year
olds, 0.30 on 7 year olds, 0.40 on 8 year olds and 100 percent on 9
year olds. This simulation is believed to be a good approximation
of the population dynamics of the Area E herd.
It agrees well with
herd size estimates made under Work Plan 11, especially for the first
years of the simulation period.
In order to project what could be accomplished with this herd it was
necessary to estimate expected reproductive rates as a function of
winter population size.
This relationship was approximated by a linear
density dependent function.
Between 1966 and 1972 posthunt populations
averaged 4650 while calf crops averaged 60 calves/lOO cows.
From 1958
to 1962 herd size averaged around 3000 (based on a comparison of helicopter trend counts) while average reproduction was 67 calves/lOO cows.
These two points determined the regression
= 0.8 - 0.0000435X, ~vhere
Yis the expected reproductive rate and X is the population size the
previous winter.

Y

It should be noted that this function and any projections based on it
are not likely to be accurate for population sizes greatly differ from
recent levels.
It is, however, a first approximation of what might
happen.
Based on this function and natural mortality rates estimated by simulation, response surfaces were drawn to show long term average expected
levels of herd characteristics for sustained yield management plans.
All combinations of antlered and antlerless harvest rates from 0 to 100
percent were considered, although many of these would be obviously infeasible.
The procedure for computer calculation of these response
surfaces is discussed below.
These were assembled in the form of a
nomograph (ouija board) as described by Gross et al. (1972).
Uncompahgre deer data was treated in the same way using information from
1969 to 1972. Table 2 summarizes simulation results for the Uncompahgre
herd. Mortality rates were 30 percent for fawns and 12 percent for all
adult age classes.
The reproductive function was estimated as Y = 1.6 0.00003X.

�-268-

Table 2. Simulated population
prehunt 1969 to prehunt 1972.

Prehunt

1969

Harvest
Posthunt

Prehunt

1969

1970

Harvest
Posthunt

Prehunt

1970

1971

Harvest
Posthunt

Prehunt

1971

1972

dynamics

of the Uncompahgre

deer herd _

No.
Bucks

No.
Does

No.
Fawns

Total

6,780

13,067

12,153

32,000

52

49

2,580

2,101

580

4,200

10,966

11,573

26,739

38

41

7,747

13,701

9,865

31,313

56

44

2,893

464

111

4,854

13,237

9,754

27,845

37

37

7,686

15,063

11,900

34,649

51

50

1,929

0

0

5,757

15,063

11,900

32,720

38

37

9,231

17,420

12,542

39,193

53

61

Nomograph

Calculation

Males/IOO
Simulated

Females
Observed

Procedures

A computer program was developed to make the calculations necessary for a
nomograph.
Prior to the development of this program, information for a
nomograph had to be derived from a series of long term simulations.
Values
for population characteristics
are calculated for each of 441 combinations
of antlered and antler1ess harvest rates.
These data are graphed as contour
maps to produce the nomograph plots.
The computer program uses the following
logic to perform these calculations.

Basic Principles
1.

and Underlying

Assumptions

There exists a single average expected value for every characteristic
of a population under any sustained yield management plan (a management
plan is defined here as a combination of an antlered harvest rate and
an antlerless harvest rate to be maintained over a period of time).

�-269-

2.

Certain features of a population remain constant (or vary around
a mean value in an unpredictable way).
These features include the
percent males in the young, the distribution of hunting effort by
age class, natural mortality rates by age class, and wounding loss
rates.

3.

Reproductive rates tend to decrease as population size increases.
This relationship may be approximated by expressing reproductive
rates as linear functions of population size at some point in the
year.
For species subjected to a "winter pinch period", the beginning of the winter season is an appropriate time.

Derivation

of Calculation

Procedure

Given: .
a value for the percent bucks in the fawn crop = PCB.
a SQt of age specific natural mortality rates for winter
summer = WMi, SMi•

and

a set of sex and age specific effort rates BEFi, DEF •
i
a wounding loss rate for antlered and antlerless animals WLB, WLD.
a set of reproductive parameters:
Al
fawn reproductive rate intercept.
A2

yearling

A3

adult reproductive

rate intercept.

Bl

fawn reproductive

rate slope.

B2

yearling reproductive rate slope.
adult reproductive ro!lteslope.

B3

reproductive

rate intercept.

a management plan consisting of the percent of the antlered population to be harvested annually and the percent of the antlerless population to be harvested annually = BRR, DHR
Let:
WMSi

1 - WMi

SMSi == 1 - SMi
BHSl

1

DHR*EEFl*

(1 + WLD)

BHS

1

BHR*BEF.*

(1 + WLB),

DHR*DEF.*

(1 + l&lt;!LD)

i

DHSi
F

1

1

1

i :f 1

number of fawns at the time of the preseason

count.

�-270-

Then:
# buck fawns at preseason count = PCB*F
# buck fawns prehunt = PCB*F*SMS

l

= PCB*F*SMS1* BHSl

# buck fawns posthunt

# yearling bucks at preseason count = PCB*F*SMS1*BHS1*WMSl
# yeariing bucks prehunt

PCB*F*SMS1* BHS * WMS * SMS
1
1
2
# yearling bucks posthunt = PCB*F*SMS1*BHS1*WMS * SMS * BHS
1
2
1
In general the number of bucks of age class "n" (n ~ 2) at the preseason count is: n-l

n

PC:S*F*

i=l

==

BHS.*
SMS·* WMS.
111

and the total buck population (~2)

at the preseason count is:

PCB*F*

similarly the number of antlerless animals of age class n (n ~ 2) at the
first reference point (preseason count) is:
n-l
(1 - PCB)*F*

~

DHS *SMS * WMS.
i

i=l

i

1

and the total doe population (~2) at this point is:

(1 - PCB)*F*

t [J

DHSi * SMS *
i

Total fawns at this point of course is "F".
Now the fawn/doe ratio is:

FI

t*(1

- PCB) *

tQ~J

DHS/

J

SMS/ WMSJ

�-271-

the unknown "F" values
computed as:

cancel each other and the ratio can be

By the same reasoning

the buck/doe

to a simplified form by cancelling
numerator and denominator.

ratio may be reduced

from:

out the F's which occur in both

Note that the preseason fawn/doe and buck/doe ratios can be calculated
for any sustained yield management plan without considering reproductive
rates.
The total number of bucks

PCB*F*

(~2)

L [(1 i

BHS *SMS(WMS

and the total number of does

posthunt

iJ

(~2)

*SMYBHS j

posthunt

J

*F*"t. n
j-1

(l - PCB)

is:

.*WMS .
[ . I DHS.1*SMS 1
1

*SMS *DHS
j

j=2 1=

and the total number of fawns posthunt
F*(PCB*SMS

is:

j

is:

*BHS + (1-- PCB)*SMS *DHS )
1
1
I
I·

The ratios can again be calculated

because

the F's can be cancelled.

The sum of the 3 expressions above is the posthunt or prewinter population size. This is the value used in the reproductive functions.
To
save space from now on this sum will be represented as SUMF. Note that
the F can be factored from this sum. Therefore SUMF/F contains no F's.
Call this quantity SUM.

�-272-

The number of does who were fawns at breeding season can be
calculated from expression (1) by setting n = 2. Denote this
quantity as FDI and note that F may be factored out and that
FDI/F contains no Fls. Call this quantity D •
l
The number of yearling does can be calculated
n = 3. Define FD2 and D2 as above.

the same way with

The remainder of the preseason doe population is the number of
adult does. Define FD3 and D3 for these animals.
Now the number of fawns at the preseason
calculated as follows:

Divide

through by F leaving:

Multiply

this out to get:

Collect

count is F and can be

terms:

Rearranging

again:

Since SUMF

= SUM*F:

F ~ (Dl* Al + D2*A2 + D3*A3 - 1) I [:SUM* (Dl* Bl + D2*B2 + D3* B3~
Now F is defined in terms of the reproductive parameters, the Di values,
and SUM. None of these contains any left over F values so this is a
computational formula for F, which in turn can be plugged back into the
earlier formulae to give actual values for such things as total population
pre and posthunt, buck and doe harvests, etc. Any characteristic of the
steady-state population may now be calculated.
It should be noted that all population characteristics
of interest are
either entirely independent of the reproductive parameters (the ratios)
or directly proportional to F. This means that any error in estimating
the reproductive parameters which results in a calculated fawn crop size
10 percent higher than the true value will not affect the ratio characteristics (fawns/doe, bucks/doe, etc.) at all, but will result in
calculated values for the other characteristics
that will be exactly
10 percent too high.

�-273-

It is therefore possible to calculate a relative sensitivity to
error in estimating reproductive parameters for any management
plan. The most straightforward approach is to utilize the population size at the midpoint of the observed data points used to
calculate the reproductive functions.
Any reproductive function
must be forced through this point. A change in slope therefore
must result in a change of intercept as well.
For example if the population size and reproductive rate through
which any function must be forced are X and R respectively, and
the intercept of the current function is A; then if the slope of
a new function is 10 percent less than that of the current function.
the new intercept may be calculated as A' = .9A + .1R since
"
(A - R)/X is the current slope and A' must satisfy (A' - R)/X =
.9 [(A '- R)/X].
In this example - 10 percent was chosen arbitrarily.
For a
sensitivity test a value should be chosen to reflect the highest
reasonable amount of slope variation that might exist.
Now with
the new A' and B' values a new F' can be calculated for any management plan, and the ratio F'/F is a measure of the relative
sensitivity of that management plan to possible error in estimating
the reproductive parameters.
As stated before, the ratio characteristics are unaffected by such errors, but characteristics such as
population or harvest sizes may have true values which differ from
the calculated values by as much as
I - F'/F
100 percent.

I

LITERATURE

1*

CITED

Gross, .J. E., J. E. Roelle, and G. L. Williams.
1972. Progress
report - program ONEPOP and information processor: A systems
modeling and communications project.
Colo. Coop. Wildl. Res.
Unit, Ft. Collins, Colo.
327 pp.

J

·1

Prepared

.

.e:.:

/

•

,

by _'·..J.:4/~· ·,~·&lt;-",,·lut:"':'i..!.(.lo!""_·
.::x:::"-:"':-;"-/";:'~.,JoL...:.J:::.'--=c .;. e'~,&gt;,,~~::.:/~;_-.-_
/James F. Lipscoml{
Wildlife Resear&amp;her Candidate
.j

��July,

-275-

JOB PROGRESS

State

of

Project
Work

REPORT

COLORADO
No.

W-38-R-27

Experimental. Elk Harvest

Covered:

April

Raymond

Investigations

Job No.

Job Title:

Personnel:

Deer-Elk

lIe

Plan No.

Period

1973

1, 1972 through March

2
Regulations

31, 1973

J. Boyd

ABSTRACT

Publication of this job was not
for submission of an article to
be delayed because of personnel
completed and it is anticipated
Segment 29.

completed during this segment.
Plans called
a professional Wildlife Journal but had to
transfers.
The article is about 50 percent
that it can be submitted for publication during

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                  <text>-1-

JOB PROGRESS

October,

REPORT

S tate of

.:::.Co.:::.L::::.o.:::.RA~D::.:O::.._
_

Project No.

W-SS-R-1S

Work Plan No.

1

Job Title
April

Production

16, 1972 through

Bird

Investigations
1

Job No.
Waterfowl

Period Covered:
Personnel:

Migratory

Surveys

June 30, 1972

C. Bryant and Staff, Monte Vista National Wildlife Refuge;
W. Blanchard and Staff, Brown's Park National Wildlife Refuge;
C. Hayes, D. Kirkland and J. Randall, Bureau of Sport Fisheries
and Wildlife; F. N. Folks and C. Jensen, Utah State Division
of Wildlife Resources; C. Braun, J. Corey, D. Coven, G. Crawford,
R. Desilet, W. Dolezal, H. Funk, R. Hopper, J. Houston, J. Lorentzson, R. Lowry, W. Russell, K. Wagner and M. Szymczak,
Colorado Division of Wildlife.

ABSTRACT
Generally dry conditions were found throughout most of Colorado's major
waterfowl breeding areas.
The total estimated number of 47,276 breeding
pairs of ducks was the smallest total recorded for the state since 1964
and was 20 percent below the long term average.
The mallard continued as
the major breeding species and in 1972 constituted nearly half of the entire
breeding population.
The post-nesting season population in Moffat County
declined approximately eight percent below the 1971 level as a result of
production being down nearly 16 percent.
Gosling production in northcentral Colorado was approximately
17 percent below the 1971 level but the
total number of geese observed in the area was unchanged from 1971.

1973

��-3-

WATERFOWL

PRODUCTION

Michael

SURVEYS

R. Szymczak

P. S. OBJECTIVE
To formulate

waterfowl

harvest

regulations

SEGMENT

for Colorado.

OBJECTIVES

To estimate the number of duck and goose breeding pairs,
each of Colorado's major waterfowl breeding areas.

METHODS

by species,

in

AND MATERIALS

Present duck breeding pair and production surveys
pair inventory of only major production areas.

consist

of a breeding

The 1972 duck breeding pair surveys were conducted during the period of
May 8 to June 30. Surveys in North Park and the Cache la Poudre and South
Platte Valleys were conducted exclusively
from the air.
Ground counts were
made in the Yampa Valley and Brown's Park.
In the San Luis Valley intensive
aerial counts were adjusted for visibility by air-ground comparison studies.
Pair estimates for the Monte Vista National Wildlife Refuge in the San Luis
Valley were obtained from nesting transects.
All survey methods and sample
areas for ducks remained the same as in previous years.
Surveys of Canada goose production were conducted within the period of
April 16 to June 16. Production estimates for Moffat County were made from
as complete a count as possible of hatched or active nests and brood size.
Population estimates in north-central
Colorado were obtained fro.m counts of
goslings and adults conducted from the ground during the period in which the
birds were flightless.
The west-central
Colorado survey was conducted,
however, poor results were obtained and are therefore not presented in this
report.
All flying was accomplished with a Cessna 185 aircraft.
Areas sampled by
section or block were flown with one observer, while two observers were
used in sampling by transect.
On the basis of these studies, a report is submitted
Fisheries and Wildlife, which constitutes Colorado's
continental cooperative breeding ground survey.

to the Bureau of Sport
part in the annual

�-4-

RESULTS AND DISCUSSION
Generally dry conditions were found throughout most of Colorado's major
waterfowl breeding areas.
The San Luis Valley, Colorado's most important
duck breeding area continued in a dry cycle which began in 1971. The run
off from the snow pack in the mountains surrounding North Park produced
only fair water conditions.
Marshes and drainage basins in the Cache la
Poudre and South Platte River Valleys were very dry and river flows extremely
low. Low water along the rivers in northwest Colorado was detrimental to
duck nesting, but did insure that island nesting Canada geese would not be
threatened by flooding.
The reduction in the number of breeding pairs in Colorado's two major
breeding areas, the San Luis Valley and North Park, resulted in the smallest
breeding pair total recorded for the state since 1964. The 47,276 breeding
pairs are nearly 20 percent below the long term average (Table 1). Only in
the Cache la Poudre Valley were more breeding birds observed in 1972 than
in 1971.
The mallard, although recording a decrease in total numbers from the 1971
level, increased in terms of the composition of the total breeding population (Table 2). Three species, the gadwall, redhead, and widgeon recorded
major declines from the 1971 level on a percentage basis.
All species,
except the green-winged teal decreased in number.
The post-nesting season
Moffat County population of Canada geese was estimated to be approximately
1,400 birds in 1972 (Table 3). The total is eight percent below the 1971
level (Table 4). Production estimates in Moffat County in 1972 were down
approximately 16 percent from 1971 levels (Table 5). The lower estimates
were mainly a result of a decline in brood sizes on the Yampa and Little
Snake Rivers rather than a reduction in the estimated number of nesting
pairs.
Production on the Green River increased substantially over the 1971
level in the Brown's Park area and remained stable in the Dinosaur National
Monument.
The results of the 1972 Canada goose production census in the Ft. CollinsBoulder-Denver area are presented by individual area in Table 6. Gosling
production was down approximately 17 percent from the 1971 level (Table 7).
Only Fort Collins recorded an increase among the five trend areas. The
greatest decline was measured in the Boulder area. The total number of geese
observed on the trend areas in 1972 was essentially unchanged from both the
1971 total and the three year average (Table 8).

FALL FLIGHT PREDICTION
Dry conditions throughout Colorado's major duck breeding areas in 1972 will
definitely result in a much smaller fall flight than in 1971 and probably
the smallest since 1964. The Canada goose flight from both northwest and
north-central Colorado in 1972 will be very similar to the 1971 flight.

Prepared

by2n:.~,j!L~
R.
Michael
Szymcz~
Assistant Wildlife

Researcher

�-5-

Table 1. Summary of Colorado duck breeding ground population estimates in
selected areas, 1972.

Area

Percent Change
Total Estimated Breeding Paris
From Long
From
Long Term
Term
Average
Average
1:./
1971
1971
1972

San Luis Valley

23,509

30,272

27,813

-22.3

-15.5

North Park 'l:../

8,922

14,711

17,989

-39.4

-50.4

South Platte Valley

7,019

8,672

6,009

-19.1

+16.8

Cache 1a Poudre Valley

4,630

3,115

3,022

+48.6

+53.2

Yampa Valley

1,857

2,340

2,867

-20.6

-35.2

Brown's Park

1,339

1,581

1,029

-15.3

+30.1

Totals

47,276

60,691

58,729

-22.1

-19.5

1/ San Luis Valley and North Park averages are based on results of 1964 through
1971 and 1968 through 1971 surveys, respectively, because of changes in survey
methods utilized prior to those dates. Figures for other areas are 18 year
averages.
Aerial counts corrected by species from visibility ratios obtained in the
San Luis Valley.

'l:../

�-6-

Table 2.

Species composition of Colorado's 1972 duck breeding population.
Number of Breeding Pairs
1954-1971
1972
1971
Average 1/

Percent S2ecies Composition
1954-1971
1972
1971
Average

22,504

24,150

28,121

47.6

39.8

56.7

6,532

7,901

4,759

13.8

13.0

9.6

Gadwall

4,666

9,884

5,155

9.9

16.3

10.4

Pintail

3,073

3,862

3,419

6.5

6.4

6.9

Green -winged Teal

2,094

1,923

2,294

4.4

3.2

4.6

Shoveler

4,152

4,354

2,074

8.8

7.2

4.2

Redhead

1,802

4,276

1,813

3.8

7.0

3.7

American Widgeon

1,331

2,640

853

2.8

4.3

1.7

Other Divers

1,122

1,701

1,142

2.4

2.8

2.3

Total

47,276

60,69l

49,630

Species
Mallard
Blue-winged and
Cinnamon Teal

l/ Species composition computed from data from all areas for the 18 year
period regardless of changes in survey methods.

�-7-

Table 3. Number of Canada geese observed
County, Colorado, 1972.

and estimated

production,

Moffat

Estimated
Goslings

Total
Birds

Nesting
Pairs

Non-nesting
Birds

Total
Adults

Spgs.

13

183

209

47 'l:...1

256

to Cross Mtn.

21

31

73

48

121

Lily Park

5

104

114

122

136

Sub-total

39

318

396

117

513

Area

No.

!.I

Yampa River
Craig to Juniper
Juniper

Green River
Brown's

Park

29

67

125

139

264

Dinosaur

Nat' 1 Manum. 11

29

106

164

136

300

58

173

289

275

564

27

205

259

61 !!!

320

124

696

944

453

Sub-Total
Little

Snake River

Grand Total

11

Calculated

'l:...1Includes

using average brood size and number

12 goslings

hatched

11

20 goslings

hatched

nests.

from eggs taken from nests before

Area first surveyed in 1970, data supplied
Division of Wildlife Resources.

il Includes

of successful

1,397

final survey.

by F. Neil Folks, Utah State

from eggs taken from nests before

final survey.

�Table 6.

Results of the north-central Colorado goose census, June 15, 1972,(continued).

Production
Area

Fort Collins
(continued)

Water Area

No.
Broods

Total
No. Goslings

2
4
1

7
15
2

4
15
2

11
30
4

295

550

845

26
4
44

16
1
58

42
5
102

74

75

149

58
29
6
18
36
11
3
26
6
6
0

264
92
2
8
31
15
15
45
2
2
4

322
121
8
26
67
26
18
71
8
8
4

199

480

679

0
15
0
0

42
28
48
125

42
43
48
125

Parkwood Lake
Kitchel Lake
Horseshoe Lake

Sub-total
Loveland

Boedeaker Reservoir
Mariana Reservoir
McNeil Reservoir

5
1

-

Sub-total
Boulder

Valmont Reservoir
Terry Lake
Crystal Lake
Ish Lake
Faivre Ponds
Boulder Valley Farm
Hayden Lake
Peaceful Meadows
Eddy Lake
Angus Ranch
McCall Lake

-

-

-

Sub-total
Denver

S. Colo. Blvd. &amp; Quincy
Blackmer Lake
Reservoir No. 3
Denver City Park

0

0
0

Total No.
Adults and Yrlgs.

Total No.
of Birds

.....I
0
I

---------------------------------------------------------------------------------------------------------

�Table 6.

Results of the north-central Colorado goose census, June 15, 1972 (continued).
No.
Broods

Total
No. Goslings

Total No.
Adults &amp; Yrlgs.

Total No.
of Birds

0
0

0

0
18
31
50
25
60
37
0
2
9
6
8
0
8
25

105
3
26
30
72
30
336
104
7
108
66
4
40
13
16
33

105
3
44
61
122
55
396
141
7
llO
75
10
48
13
24
58

Sub-total

294

1,236

1,530

Grand Total

1,081

2,853

3,934

Production
Area
Denver
(continued)

Water Area
Sloans Lake
Kountze Reservoir
Federal Center Pond
Clairfield Reservoir
Gallup Reservoir
Columbine Coun try Club
Bowles Lake
Kings Pond
Lower Tule Lake
Marston Reservoir
Grant B Reservoir
Patrick Lake
Pinehurst Country Club
Kendrick Lake
Mohn Estate
Standley Lake

0
1
3
2
2
0
3

-

I
I-'
I-'
I

�-12-

Table 7. Total number of Canada goose goslings produced in north-central
Colorado production trend areas, 1972.

No. of Goslings
1969-1971
1972.
1971
,,1iverage

Percent Change
From 1969-1971
From 1971
Average

Wellington

219

301

286

-27.2

-23.4

Fort Collins

318 1/

255

250

+24.7

+27.2

Loveland

74·

125.

75

-40.8

- 1.3

Boulder

199

357

251

-44.3

-20.7

Denver

294

296

21g,

- 0.7

+ 6.9

1,137

-17.2

- 2.9

Area

".{f

.,'

..

;. ~:

Total

1,104

J

1,334.

,;1

!/ Includes 23 birds raililedat Ft. Colli,ns Wildlife Research Center.
"
0::"

t.:;:-.

Table 8. Total number of Canada geese observed in north-central Colorado
production trend areas, 1972.

No. of Geese
1969-1971
1972
197~
Average

Percent Change
From 1969-1971
From 1971
Average

Wellington

731

822

894

-11.1

-18.2

Fort Collins

868

690

707

+25.8

+22.8

Loveland

149

249

186

-40.2

-19.9

Boulder

679

862

729

-21.2

- 6.9

Denver

1,530

1,446

1,339

+ 5.8

+14.3

Total

3,957

4,069

3,855

- 2.8

+ 2.6

Area

�-13-

October,

1973

JOB PROGRESS REPORT

State of

COLORADO
----~~~~~----------

Project No.

W-88-R-18

Work plan No.

1

Job Title

Job No.
Trapping

Period Covered:
Personnel:

Migratory

Bird Investigations
2

and Banding Ducks and Geese

April 1, 1972 through March 31, 1973

J. Randall, Bureau of Sport Fisheries and Wildlife; R. Baker,
C. Braun, J. Corey, D. Coven, C. Crawford, G. Crawford, W. Dolezal,
J. Ellenberger, H. Funk, A. Hemmert, R. Henry, J. Jackson,
R. Kitzmiller, J. Lorentzson, J. Olterman, B. Peterson, L. Rottman,
W. Rutherford, N. Smith, H. Spear, M. Stromberg, M. Szymczak,
K. Wagner, R. Hopper, Colorado Division of Wildlife.

ABSTRACT
Trapping activities during Segment 18 resulted in the banding of 12,721 ducks
of 10 species at seven locations in Colorado.
The mallard was again the major
species in the banded sample, contributing 8,853, or about 70 percent of the
total. Pre-season efforts continued in the high country production areas,
where a study was undertaken to' investigate migration routes, harvest characteristics and mortality rates of duck populations here.
The banded sample
in these high country areas in 1972 totaled 8,272 ducks, or 2,758 in North
Park, 1,623 in South Park and 3,891 in the San Luis Valley.
Progress was
made in increasing the banded samples of some species other than mallards and
pintails by employing the night-lighting technique and by using smaller funnel
hoops on the Salt Plains traps. Post-season banding (winter) resulted in a
banded sample of 6,021 mallards during the Segment.
Segment 18 goose trapping operations resulted in the banding of 1,169 Canada
geese. Goslings, hand-reared or captured in the wild and transplanted in
various locations during the summer, accounted for 427 of this total.
The
remalnlng geese were banded post-season in the Arkansas Valley (564) and
the Cache 1a Poudre Valley (178).

�-14-

RECOMMENDATIONS

1.

Continue all pre- and post-season duck and goose trapping and
banding operations as outlined in the W-88-R Job Description
for Segment 18. Make additional effort to band a sample of
mallards wintering in western Colorado.

2.

Continue to experiment with new trapping techniques, including
night-lighting, ftrive-trapping at night and funnel modification.

�-15-

TRAPPING AND BANDING DUCKS AND GEESE
Richard M. Hopper
This report summarizes all waterfowl banding activities under Federal
Aid Project W-88-R-18 for the segment year April 1, 1972 to March 31,
1973. Analyses of band recovery data are conducted under a separate
job (Work Plan 1, Job 3) and thus, will not be included in this report.
This report merely presents a listing of numbers of ducks and geese
banded by species and location during th¢ segment.
P. S. OBJECTIVE
To formulate waterfowl harvest re&amp;ulations for Colorado.
SEGMENT OBJECTIVES
1.

To provide adequate samples of banded ducks and geese in Colorado's
major breeding and harvest areas for the purpose of accumulating
migration and annual mortality data.

2.

To report species and numbers of mallards winter banded in the South
Platte Valley, Arkansas Valley, and Bonny Reservoir as part of Work
Plan 3, Job 6.

3.

To report transplant geese banded under Work Plan 2, Job 2.
METHODS AND MATERIALS

The banding program in Segment 18 remained the same as in past years with
operations consisting of three phases: (1) mid-summer goose transplant
banding, (2) late summer (pre-season duck banding, and (3) winter (postseason) duck and goose banding. Goslings for transplanting were handreared and also secured from the wild by drive-trapping or by using baited
walk-in traps prior to the time the birds were capable of flight. Preseason duck banding was conducted in August and September utilizing baited
Salt Plains traps (cage-type) to capture birds in North Park, South Park,
and the San Luis Valley. An additional method, night-lighting, was also
used on an experimental basis to catch ducks in North Park during the preseason period. The night-lighting technique used is best described by
Cummings and Hewitt (1964) and Drewin et al. (1967). Post-season banding
(January-February) employed baited Salt Plains traps and cannon-nets for
duck trapping, and baited cannon-nets and a large walk-in trap for goose
trapping.

�-16-

RESULTS

AND DISCUSSION
Ducks

Numbers and species of ducks banded during Segment 18 by State personnel
are listed in Table 1 by location.
The banded sample was similar to the
previous year, totaling 12,721 ducks of 10 species.
Seven locations were
again involved, with the South Platte Valley contributing the largest number
(3,744) followed by North Park (2,758) and the San Luis Valley (2,269).
As usual, the mallard was the dominant species represented in the banded
sample, constituting 8,853, or about 70 percent of the total.
However, this
percentage was somewhat less than in the previous year, mainly because of an
effort to band greater numbers of other species in the high country areas
during the pre-season period.
Even with this added effort to band other
species, pintails and green~inged
teals were still the only other species
that contributed a significant number of birds to the banded sample, with
2,524 and 835, respectively.

High Country Study

Pre-season banding was continued in North Park, South Park, and the San Luis
Valley as the basis for an overall investigation of high country duck populations in Colorado.
Samples of ducks were banded Simultaneously in all
three areas for the second consecutive year.
This procedure is expected to
continue for at least the next four years.
The objective is to investigate
relationships among the three areas in terms of migration routes, harvest
characteristics,
and mortality rates.
Information resulting from this study
will allow us to evaluate the justification
for establishing duck hunting
season dates in Colorado based on elevation.
This can be termed "altitudinal
zonation" of hunting season dates.
This could provide earlier and separate
season dates for the high country, thereby yielding additional duck hunting
opportunity where little occurred before.
Species composition by age and sex for 8,272 ducks banded in the three high
country areas is shown in Tables 2-5. Figures for the San Luis Valley
include bandings by both State and Federal personnel.
The Bureau of Sport
Fisheries and Wildlife is cooperating in this study by banding a portion of
the Valley quota on the Monte Vista and Alamosa National Wildlife Refuges.
sample of 8,272 ducks for the three areas combined during Segment
above that of the previous Segment (6,980). This
increase was due to larger samples being obtained in North Park and the San
Luis Valley.
Sizable increases were attained for the gadwall and pintail in
North Park and for the gadwall, green-winged teal and blue-winged and
cinnamon teal in the San Luis Valley. Total number of birds banded in South
Park was similar during Segments 17 and 18. The decrease in the mallard
sample for this area in Segment 18 was compensated for by an increase in
number of pintails banded.
The banded

18 was substantially

�Table 1.

Number of ducks banded by species, location, and period of year, 1972-73.

Species

North
Park

Pre-season I7
San Luis
South
Valley
Park

Number of Ducks Banded
Post-season '1.7
Bonny
Cache la
South Platte
Reservoir
Valley
Poudre Valley

Arkansas
Valley

Total

Mallard

1,114

868

800

645

3,744

618

1,064

8,853

Gadwall

123

-

5

-

-

-

-

128

American Widgeon

27

1

-

-

-

-

-

28

Green-winged Teal

247

140

448

-

-

-

-

835
I

B1ue-winged and
Cinnamon Teal

~

'-I

76

84

87

-

-

-

-

247

Pintail

1,103

521

900

-

-

-

-

2,524

Redhead

68

6

29

-

-

-

-

103

Lesser Scaup

-

1

-

-

-

-

-

1

Wood Duck

-

2

-

-

-

-

-

2

2,758

1,623

2,269

645

3,744

618

1,064

Total

l/ August-September.
~/ January-February.

12,721

I

�-18-

Table 2. Age and sex composition of ducks banded pre-season in North
Park, 1972.

AM

Number of Ducks Banded
Age aridSex
1M
AF
IF

Mallard

262

303

285

264

1,114

Gadwall

49

11

46

17

123

American Widgeon

5

10

4

8

27

Green-winged Teal

69

82

31

65

247

Blue-winged and
Cinnamon Teal

8

36

6

26

76

Pintail

280

277

273

273

1,103

Redhead

23

11

25

9

68

Total

696

730

670

662

2,758

Species

Table 3.

Total

Age and sex composition of ducks banded pre-season in South Park,

1972.

AM

Number of Ducks Banded
Age and Sex
IF
AF
1M

250

209

250

159

868

American Widgeon

0

0

1

0

1

Green-winged Teal

45

56

16

23

140

Blue-winged and
Cinnamon Teal

5

42

4

33

84

Pintail

150

125

125

121

521

Wood Duck

0

1

0

1

2

Redhead

0

2

4

0

6

Lesser Scaup

1

0

0

0

1

451

435

400

337

1,623

Species
Mallard

Total

Total

�Table 4,
Age and sex c.omposition
Luis VaI Ley , 1972. 1:./

of

ducks

banded

p re+s e as on in

the

San

--------------_.-_ ...-.-._._--_ .._..._---_ ..._-.----._----_. __ ._-.•._-----._-----_._----_._---------------Number of Ducks Banded
--,.-.---.~-,--~.~~~~-.-~.•.
._._._ .________
Age and=-=S:...:e:..:x~
__

1M.

AF

IF

Total

ME.llard

372

368

338

1,421

Gadwell1

1~4

9

40

94

5

2

2

1

10

3fQ
'+~

153

82

89

673

67

131

28

84

310

o

.1

0

0

1

298

397

234

408

1,337

9

10

44

Species

American

Hidgeon

Green --winged

'I'e 81

Blue-\vi.nged

and CinrUll1JOn

Shoveler
Pintail

Teal

Redhead
Lesse:!: Sc.aup

Total

}j Includes
National

ducks
"\Hldlife

1

o

0

0

1

1.0n

19112

732

970

3,891

banded by Fede xa.l, pe raonneL on Alamosa and Monte Vista
Refuges as part of hLgh+coun t ry duck study.

Progress
vms made in incre2.sin.g
the s amp I e s Lzes of species
other
than mallards
and pLnt.a I Ls ,
(is indicat.ed
abo ve , t.he most notable
i.ncreases
were for gadw al.Ls , gre(::n.--Ij\;-]:Dged t.';:~als, and blu.e--v!iIl.ged and cinnamon teals.
This improvement was due p:cJ_lIw.ril:y to Lnc r'e as ed arfo rt; and time expended by the banders.
The n.I.ght r-Lf.ghti.n g t.echnLque was heLpful, and showed great
p ro mt se ,
Smaller
funnel hoops in the Salt Pla.:Lnstcap!:;
appe a.re d to increase
the selectivity
fo these traps toward tee ..1 and &amp;.'VJaY f rom the larger
species.
However, much
larger
samples of spec:;_.es other
than mall.a rds and pintails
are needed in
order to make the study complete"

�-20-

Table 5. Age and sex composition of ducks banded pre-season
high-country areas combined, 1972. !/

in the three

AM

Number of Ducks Banded
Age and Sex
1M
AF
IF

Total

Mallard

855

884

903

761

3,403

Gadwall

50

5»

55

57

217

10

12

7

9

38

Species

American

Widgeon

Green-winged

Teal

463

291

129

177

1,060

Blue-winged

and Cinnamon Teal

80

209

38

143

470

Shoveler

0

1

0

0

1

Pintail

728

799

632

802

2,961

Wood Duck

0

1

0

1

2

Redhead

36

25

38

19

118

2

0

0

0

2

2,224

2,277

1,802

1,969

8,272

Lesser

Scaup

Total

!/ Includes

ducks banded by Federal personnel on Alamosa and Monte Vista
National Wildlife Refuges in the San Luis Valley as part of the high-country
duck study.

Winter

Banding

Post-season banding of mallards was continued in eastern Colorado under Work
Plan 3, Job 6 (Investigation of Mallard Management Units of Eastern Colorado).
Quotas were difficult to attain in three of the eight banding areas because
of a poor distribution of the duck population.
Quotas were increased in
some of the other areas to make up for this situation, but the total banded
sample of 6,021 mallards was somewhat below the 6,564 birds banded the
previous year (Table 1).
Some effort was made to band a sample of mallards that winter in west-central
Colorado.
This venture was unsuccessful in terms of birds banded, as most
of the few days of effort expended were spent becoming acquainted with the

�-21country and searching for bandable populations.
Highline Lake, northwest
of Grand Junction, showed considerable promise as a trapping site.
More
effort will be made in this area next Segment.

Geese
About 1,200 Canada geese were banded during Segment 18 (Table 6). This
number represented less than one-half of that banded during Segment 17,
mainly because of poor success during the winter or post-season banding
period.

Table 6.

Number of Canada

geese banded by location

and period of year,

1972-73.

Number Banded
Summer
Transplants

Location

Arkansas

Valley

1./

Post-season

2/

Total

0

564

564

0

178

178

32

0

32

North Park

150

0

150

South Park

75

0

75

170

0

170

427

742

1,169

Cache la Poudre
Dolores

Valley

Valley

South Platte

Valley

Total

1/ June-July.

I./ January-February.
Summer transplant geese accounted for 427 of the total banded sample (Table
6). These goslings were banded and released at Totten Reservoir in the
Dolores Valley (32). Walden Lake in North Park (150). Antero Reservoir in
South Park (75), and Jumbo Reservoir (100) and Prewitt Reservoir (70) in
the South Platte Valley.
Numbers of males and females were nearly equal
for the overall composition of transplant birds (Table 7). Work Plan 2, Job
2 discusses the transplant program in more detail.
Only 742 Canada geese were banded post-season in the Arkansas and Cache la
Poudre Valleys (Table 8). The number banded was much below the 1,000 quota

�-22-

desired for each area. A fair sample was obtained at Turk's Pond (564) in
the Arkansas Valley, but an inadequate sample was taken at New Windsor
Reservoir (178) in the Cache la Poudre Valley. A different density and
.distribution of geese from previous years were the main problems encountered
in acquiring a suitable sample in the Cache la Poudre Valley during Segment
18. The age and sex composition of both banded samples was distorted in
favor of adults, espeC~ally males (T~ble 8). Additional information regarding
post-season
banding
for Work
Plan 2, Jobs
5-6.activities can be found in the Job Progress Report

Table 7.

Sex compositib~ of goslings banded for summer transplants, 1972-73.

Number Banded
Sex

Location

I1tle
'~~.

Female

Total

'..-+,'&lt;

Dolores Valley
1;3
North Park

72

South Park

39

South Platte V~l1ey

86

19

32

78

150

36

75

84

170

217

427

Total
210

Table 8. Age and
sex composition
eastern Colorado, 1972-73.
of Canada geese banded post-season

Location

AM

Arkansas Valley

249

Cache 1a Poudre Valley

77

Total

326

r

Number of Geese Banded
Age and Sex
1M
AF
IF
41
23
64

in

UF

Total

24

0

564

2

0

76

178

252

24

76

742

250

�-23-

LITERATURE CITED
Cummings, G. E., and o. H. Hewitt. 1964.
birds at night with light and sound.

Capturing waterfowl and marsh
J. Wildl. Mgmt. 28(1):120-126.

Drewin, R. C~, H. M. Reeves, P. F. Springer, and T. L. Kuck. 1967. Backpack unit for capturing waterfowl and upland game by night-lighting.
J. Wildl. Mgmt. 31(4): 778-783.

��October,

-25-

1973

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~---------

Project No.

W-88-R-18

Work Plan No.

1

Job Title

of Waterfowl

Bird Investigations
3

Job No.
Analysis

Period Covered:
Personnel:

Migratory

Banding Data

April 1, 1972 through March 31, 1973

James Jackson, Robin Henry, Michael

Szymczak

and Richard

Hopper.

ABSTRACT
This job is dependent upon the banding program conducted under Work Plan 1,
Job 2. The current banding analysis phase is concerned with that portion of
the banding program dealing with high country duck populations, specifically
in North Park, South Park and the San Luis Valley.
Pre-season banding was
continued in these three areas during Segment 18 to yield band recovery data
necessary in implementing this analysis job. Pre-season banding in all three
.high country areas as a group has been done for only two years (1971 and 1972),
thus it will probably take at least three more years of banding here to provide
enough data for a reliable analysis.
Banding and recovery tapes were obtained from the Bird Banding Laboratory
prior to last segment.
Programs were written and the desired information was
retrieved from the tapes through computers.
Preliminary work using the resulting print-outs included mapping of recovery locations and calculating recovery
end mortality rate estimates by species, age and sex. Also, all fully punched
recovery cards received have been sorted and filed by species, banding location,
year of recovery and how obtained.

�-26-

RECOMMENDATIONS

1.

Continue pre-season duck banding in North Park, South Park and the San
Luis Valley, with existing quotas by spec1es, age and sex.

�-27-

ANALYSIS OF WATERFOWL BANDING DATA
Richard M. Hopper
This particular job is directly related to and dependent upon the banding
program carried out under Work Plan 1, Job 2. The current segment as well
as the next few segments of Work Plan 1, Job 3 will deal specifically with
the portion of the banding program aimed at high country duck populations.
Pre-season banding was initiated in the three major high country areas
(North Park, South Park, and the San Luis Valley) as a group in 1971, with
at least a five-year program in mind (1971-1975). Recovery data from these
bandings will provide the basis for the band analysis job in question.
P. S. OBJECTIVE
To formulate waterfowl harvest regulations for Colorado.
SEGMENT OBJECTIVES
To investigate migration, mortality, recovery distribution, and relationships among populations of ducks banded in Colorado's high country, specifically in North Park, South Park, and the San Luis Valley.
METHODS AND MATERIALS
Computer tapes containing all bandings from the three high country areas
and all recoveries from those bandings through the 1969-70 hunting season
were obtained from the Bird Banding Laboratory of the Bureau of Sport
Fisheries and Wildlife. During last segment, programs were written and the
desired information was retrieved from the banding and recovery tapes through
the use of automatic data processing machines.
Fully punched recovery cards have also been supplied by the Bird Banding
Laboratory for all recoveries through the 1971-72 hunting season. The Bird
Banding Laboratory will continue to provide these throughout future recovery
years. The tape program mentioned above was modified to handle cards, thus
making annual updating of recovery data a relatively simple task.

RESULTS AND DISCUSSION
Pre-season banding was continued during Segment 18 to supply band recovery
data necessary in the implementations of this job. Results of this 1972 preseason banding effort are presented in a prior section of this issue of the
Game Research Report (Hopper 1973). The only major problem confronted thus
far in the banding program is the difficulty in obtaining good banded samples
of important species of ducks other than mallards and pintails. We hope to
improve these samples in future years.

�-28-

Pre-season banding has been conducted in the San Luis Valley since 1963
and in South Park since 1968, but was not initiated in North Park until
1971. The earlier years in the San Luis Valley and South Park yielded
some valuable data that will be useful in the overall high country
analysis, but information resulting from years of simultaneous banding
in all three areas will give us the best look at the high country situation.
Thus, it will probably take at least another three years of banding in all
three areas, or a total of about five years of recoveries, to supply sufficient data for a reliable analysis.
The types of information obtained on print-out sheets (tabs) from the banding and recovery tapes were listed in the previous segment report (Hopper
1972) and will not be repeated here. Work during Segment 18 involved some
preliminary, but incomplete, mapping of recovery locations, and recovery
and mortality rate estimates by species, age and sex. Also, all fullypunched recovery cards received as of March 31, 1973 have been sorted and
filed by species, banding location, year of recovery, and how obtained.
Plans for next segment include an update of all recovery data through the
1972-73 hunting season from bandings in all three high country areas. This
will involve processing recovery cards through the established programs to
obtain print-outs of recovery data from the 1970-71 through 1972-73 hunting
season. This information will then be combined with that processed earlier
from tapes containing recovery data through the 1969-70 hunting season.
LITERATURE CITED
Hopper, R. M. 1972. Analysis of waterfowl banding data.
Wildl., Game Res. Rept., Oct. pp. 23-26.
1973. Trapping and banding ducks and geese.
Wildl., Game Res. Rept., Oct.

Prepared by

~?=-z.

~jP&amp;.

R. M. Hopper
Wildlife Researcher

Colo. Div. of

Colo. Div. of

�October,

-29-

1973

JOB PROGRESS REPORT

State of

COLORADO
------------------------

Project No.

W.,.88-R-18

Work Plan No.

1

Job Title

April

Bird Investigations

Job No.

San Luis Valley

Period Covered:
Personnel:

Migratory

Cooperative

12
Mallard

Investigation

1, 1972 through March 31, 1973

Dr. Aelred D. Geis, Bureau of Sport Fisheries and Wildlife;
Dr. Lewis Nelson, Jr., University of California; Jack R. Grieb
and Richard M. Hopper, Colorado Division of Wildlife.

ABSTRACT
A final report covering the 8-year study of duck populations and experimental
seasons in the San Luis Valley is progressing smoothly.
The first rough draft
of this report was finished during Segment 18. The final manuscript will be
completed during Segment 19 and submitted to Wildlife Monographs for possible
publication.

��-31-

SAN LUIS VALLEY COOPERATIVE
Richard

MALLARD

INVESTIGATION

M. Hopper

An intensive study of the duck population in the San Luis Valley was
started in 1963 as a cooperative investigation between the Bureau of Sport
Fisheries and Wildlife and the Colorado Division of Wildlife.
A major portion
of this study was an annual experimental hunting season held during the
period October 1-18. The eighth and final experimental season was conducted
in 1970. Preliminary results of these seasons, as well as all phases of the
study, were presented in Administrative
Report Numbers 42, 49, 79, 120, 130,
158, 175, 195, and 210 prepared by members of both agencies and printed and
distributed by the Bureau of Sport Fisheries and Wildlife, Branch of Wildlife
Research, Migratory Bird Populations Station.
Segment 17 (1971) was the last year of field work for the overall study.
Work
during part of Segment 17 and all of Segment 18 included the analysis of data,
evaluation of results, and writing the final job report covering the entire
investigation.
P. S. OBJECTIVE
1.

To develop

a harvest

2.

To determine

hunter

formula

for the San Luis Valley mallard

reaction

to various

types of hunting

population.

regulations.

SEGMENT OBJECTIVES

1. Evaluate results of the experimental hunting seasons since 1963 and begin
preparation of final reports covering all phases of the investigation
in
cooperation with the Bureau of Sport Fisheries and Wildlife.

METHODS

AND MATERIALS

All data gathered during the course of the investigation were compiled and
analyzed.
The Bird Banding Laboratory provided printouts of banding data
including computations of mortality rates, etc. necessary in the overall
analysis.
Preparation of the final report followed the style of Wildlife
Monographs.
RESULTS AND DISCUSSION
All data were analyzed during Segment 18 and the first rough draft of the
final report was finished.
The four coauthors, Jack R. Grieb, Dr. Aelred D.
Geis, Dr. Lewis Nelson, Jr., and Richard M. Hopper were assigned specific

�-32-

segments of the report for which they were responsible.
Work on this report
will continue into Segment 19, with reviewing, editing, and typing being
the major considerations.
The final manuscript will hopefully be completed
by January 1, 1974 and submitted to Wildlife Monographs for possible publication.

Prepared by -...:1{;.~"::~:Joh~f-=rl..t$~d::;.:i2=-.
-~-o+~-'~:"'.e...,¢9JJ~· w~/."-':.lo&gt;-",-_
.1.0.

Wildlife

Researcher

�October,

-33-

1973

JOB PROGRESS REPORT

COLORADO

State of

Work Plan No.
Job Title
Period Covered:
Personnel:

Migratory

W-88-R-18

Project No.

Bird Investigations

2
Job No.
Experimental Studies on
Improving the Status of Canada Goose Populations
2

April 1, 1972 through March

31, 1973

C. Hayes, D. Kirkland and J. Randall, Bureau of Sport Fisheries and
Wildlife; R. Baker, J. Corey, D. Coven, G. Crawford, J. Ellenberger, H. Gresh, R. Hopper, T. Lines, G. Lorentzson, W. Olmstead,
S. Steinert, K. Wagner, L. Webster and M. Szymczak, Colorado
Division of Wildlife.

ABSTRACT
An additional 100 goslings were added to the Jumbo Reservoir restoration
flock in the South Platte River Valley, making the total number released
there in two years, 202. Band recoveries indicate that a portion of the
birds released on Jumbo in 1971 migrated north in the spring of 1972. All
geese moved out of the Jumbo area in response to inclement weather in late
November and early December, 1972. In early spring, 1973, about 100 birds
were present in the Jumbo Reservoir area. Restoration efforts at Prewitt
Reservoir in Washington County in northeast Colorado were initiated, with
the release of 70 goslings in July, 1972. Five of the birds released at
Prewitt were reported taken during the 1972-73 hunting season.
It appeared
that the birds left the Prewitt release site about mid-November.
In North
Park, 150 birds were released on Walden Reservoir, located about 1 mile west
of Walden, Colorado, making a total of 830 birds released in North Park in
the last four years.
A total of 65 adult geese and three broods with a
total of 17 goslings were observed in North Park in late May, 1972. Two
additional nests, which later hatched, were also found in late May. At
least 85 geese were present in North Park in mid-June.
A total of 13 geese
which had been released in North Park during previous years were captured on
Wheatland Reservoir in late June, 1972. Only 12 geese from all North Park
releases were reported recovered during the 1972-73 hunting season, six from
the group released in 1972. Restoration efforts in South Park were initiated
with the release of 76 goslings on Antero Reservoir in July, 1972. Five of
the birds released on Antero were reported recovered during the 1972-73
hunting season.
Restoration efforts at Totten Reservoir near Cortez in
southwest Colorado were concluded
with the release of 32 goslings.
In
early January, 1973 the birds left the Totten area for about two weeks, and
according to band recoveries, moved into eastern yavapai County, Arizona.

��-35-

IMPROVING

EXPERIMENTAL STUDIES ON
STATUS OF CANADA GOOSE POPULATIONS
Michael

R. Szymczak

P. S. OBJECTIVE
To investigate the succes s of attempts to establish breeding
Canada geese in su i.t abLa hab Lt at whe re they do not currently
rado.

populations of
exist in Colo-

SEGMENT OBJECTIVES
1.

To prepare a technical bulletin on the technique
breeding population of Canada geese.

2.

To expand the breeding flocks of Canada geese in the South Platte Valley,
and North Park, to begin efforts to establish a breeding flock of Canada
geese in South Park.

3.

To continue efforts to establish a breeding
geese on Totten Reservoir, south of Dolores

METHODS

of establishing

a

flock of Great Basin Canada
in Montezuma County.

AND MATERIALS

Canada goose goslings for transplant purposes were obtained from four
different sources: (1) goslings from the metropolitan Denver area; (2)
goslings and some eggs from the Fort Collins area; (3) goslings from Valmont
Reservoir and Faivre Ponds in the Boulder area, and (4) eggs from nests in
danger of destruction from high water along the Yampa and Little Snake Rivers
in Moffat County.
All eggs were transported to the Fort Collins Wildlife
Research Station for incubation and subsequent raising.
All goslings livetrapped, were transported as soon as possible to the release site. Birds
obtained from eggs collected in Moffat County were released on Totten Reservoir south of Dolores.
Birds collected at other locations were released
either at Lake John Annex, Prewitt Reservoir, Jumbo Reservoir Annex, or
Antero Reservoir.
Migration and mortality information were obtained from recovery
received from the Migratory Bird Population Station.

cards

RESULTS AND DISCUSSION
Preparation

of Technical

Bulletin

Progress was made on preparing a technical bulletin outlining the foothills
area goose restoration project.
The bulletin will be completed during the
next Segment.

�-36-

South Platte

Valley-Jumbo

Reservoir

Annex

In the summer of 1971, Canada goose restoration efforts were initiated at
Jumbo Reservoir, near Segwick, Colorado, with the release of 102 goslings.
The lack of reported band recoveries during the 1971-72 season indicated
the released birds stayed within the established 250 square mile closed area
throughout the hunting season (Table 1). On April 28, 1972, 44 geese were
observed on the release site.
In June and July of 1972, 48 male and 52
female goslings were added to the Jumbo Annex population.
Banded birds from the 1971 release reported recovered in October in Minnesota and western Nebraska indicate that a portion of the birds released in
1971 moved north during the spring of 1972, probably in accompaniment of
migrant popUlations (Table 1). Heavy snows and extreme cold temperatures
in late November and early December of 1972 stimulated movement by all geese
from the release area.
According to the band recoveries, the birds moved
straight south, with recoveries in Baca County, Colorado, the Panhandle of
Texas and an area about 100 miles west southwest of San Antonio, Texas
(Table 1).
Reports in the early spring 1973 indicate there are approximately 100 birds
in the Jumbo Reservoir area.
About 30 artificial nest sites have been
erected at Jumbo Reservoir Annex and on adjacent small ponds.

South Platte

Valley-Prewitt

Reservoir

In July 1972, 37 male and 33 female Canada goose goslings were released on
Prewitt Reservoir, located near the South Platte River in Washington County,
northeast Colorado.
An area of about 100 square miles was closed to goose
hunting around the release site.
Five of the seventy birds were reported recovered during the 1972-73 season.
Four birds were taken near the release site and one in Randall County in the
Texas Panhandle in a short grass prairie population Wintering area. No
recoveries near the release site came after November 13, 1972, indicating
that the birds may have moved south after that time.

North Park
By late summer 1971, a total of 681 Canada goose goslings had been released
on Lake John Annex, eight miles west and two miles north of Walden.
These
birds had established the lower Colorado River bordering California and
Arizona and the Imperial Valley of California as their wintering areas
(Szymczak 1972).
Some released birds have been recaptured in the summers
of 1970 and 1971 on Wheatland Reservoir, in Albany County, Wyoming during
drive trapping operations on a moulting flock of Canada geese, indicating
some of the birds are using that area for moulting (Szymczak 1972). But
very few of the 440 birds released in 1970 and 1971 had returned to spend
the spring and summer in North Park (Szymczak 1971, 1972).

�-37-

Table 1. Hunting season recovery locations of Canada geese released
Jumbo Reservoir, near Sedgwick, in extreme no rt.he as t e rn Colorado.

--------------------------_._-_
Year Released
Number Released
Area Recovered

1971
102
No. Recovered
Direct
Indirect

nt

_._---_ .. _----

..

1972
No. Recovered
Direct
Indirect

Colorado
Logan County

1

0

0

Baca County

0

0

1

Chase County

0-

1

0

Keith County

0

1

0

0

1

0

County

0

0

1

Uvalde County

0

0

1

Zavala County

0

0

2

1

3

5

Nebraska

Minnesota
Chippewa County
Texas
Hockley

Total

Spring Returns
Geese began returning to North Park in the spring of 1972 as early as March
12, when 16 birds were observed on the North Platte River near Delaney Butte
Lakes.
At that time, short stretches of the rivers provided the only open
water in the Park. On March 15 four geese were observed near the feeder on
Lake John Annex.
A complete survey was made on all major water areas in North Park, with the
exception of the rivers, on May 23 and 24, 1972. The search turned up a
total of 65 adult geese, and sixteen gosling~ in three broods.
There were
eleven definite pairs observed.
One brood was found on Lake John Annex

�-38-

and two broods on Boettcher Lake.
In addition, one active nest was found
on Lake John Annex and one nest was reported on the Arapahoe National Wildlife Refuge.
Both of these active nests were eventually successful.
One
goose was found dead near a nest which had been destroyed.
The dead goose
was banded, however, it was not a bird released at the Annex.
The bird
had been banded at Wheatland Reservoir in Wyoming in June 1970. This
recovery further indicates the Wheatland Reservoir may play a significant
role.
It was later reported that there were at least 85 geese in North Park in
mid-June, with the majority being on Lake John Annex.
Moulting

Some Canada geese released in North Park continue to be recovered on Wheatland Reservoir in Wyoming during their moult.
In June 1970, 16 North Park
birds were captured on Wheatland.
Eight were captured at Wheatland in 1971,
and 13 in 1972. One bird was captured at Wheatland for the three consecutive
years.
Two of the birds captured at Wheatland in 1972 were reported harvested
the following season, one near Yuma, Arizona, and one near Fort Collins,
Colorado.
In addition, one bird was captured as a member of a mou Lt Lng
flock at Turbid Lake in Yellowstone National Park.

New Releases

and Summer Observations

On July 13, 1972, 46 male and 50 female goslings were released on the southeast side of Walden ReserVOir, which is located about 1 mile west of the
town of Walden.
An additional 54 birds, 26 males and 28 females were
released at that same location on July 21. A feeder filled with milo maize
was placed at the release site.
As during the previous four summers the birds moved throughout North Park
using IOOSt of the major water area, but seemed to prefer Hebron Sloughs,
Walden Reservoir and Lake John Annex.
The largest number of birds observed
on anyone
day was 207, 128 at Walden Reservoir and 79 at Hebron.
Band Recoveries

Only 12 birds brom all North Park releases were reported recovered during
the 1972-73 hunting season, 3 from releases in 1969, 1 from 1970, 2 from
1971, and 6 from the 1972 release.
Four birds, representing all years of
release except 1971 were reported taken along the lower Colorado River in
Arizona (3), and California (1). Three birds, 1 from 1971 and 2 from 1972
were reported taken in Moffat County in northwestern Colorado.
One of the
birds taken in Moffat County was recovered in the Brown's Park area, as
was 1 bird reported taken in adjacent Daggett County, Utah.
A summary

of all recoveries

to date is presented

in Table 2.

�Table 2. Hunting season recovery locations for all years of Canada geese released at Lake John Annex,
Jackson County, Colorado.
Year Released
No. Released

1969
193

1970
246

Area Recovered

No. Recovered
Direct Indirect

No. Recovered
Direct Indirect

All Years
830

1972
150

1971
241
No. Recovered
Direct Indirect

No. Recovered
Direct Indirect

No. Recovered
Direct Indirect

United States
North Dakota

--

Emmons Co.

0

0

0

0

-

0

1

0

0
0

0
0

0
0

0
0

-

0

0

0

1
1

0

0
0
0

-

0
0

1
1

0

0
0
0

7

0

0

2

0

0

0

0

0
0
0
2

3

1
1

0
0
0

1

0

0
0

1

0
2
1
1
1
0

1
1
0

0

1

0

0
0

1
1

0
0
2

1
0
0

0

0

0
5

1

1
0
1
2

0

Montana
Custer Co.
Rosebud Co.
Wyoming
Park Co.
Albany Co.
Carbon Co.

0
0

Colorado
Jackson Co.})
Larimer Co.
Baca Co.
Moffat Co.
Mesa Co.
Hinsdale Co.

0
0

0

0

0
0

0

0
0

0

0

0
0

1

-

0
0
0

0
0
0

1
0
0

-

0
0
0
0

1
0

1

0

1
4
0

Utah
Daggett Co.
Emery Co.
Cane Co.

0

1
0

0

0
0

0
0

1

0
0

1

New Mexico
0
1
-----------------------------------------------------------------------------------------------------------Santa Fe Co.

0

0

0

0

1

0

0

I

W
'-0
I

�Table 2. Hunting season recovery locations
Jackson County, Colorado (continued).
Year Released
No. Released

. 1969
193

Area Recovered

No. Recovered
Direct Indirect

for all years of Canada geese,released

1970
246

1971
241

No. Recovered
Direct Indirect

at Lake John Annex,

1972
150

No. Recovered
Direct Indirect

All Years
-830

No. Recovered
Direct Indirect

No. Recovered
Direct Indirect

Texas
El Paso Co.

1

0

0

0

0

0

0

1
1
0
0
6

0
0
0
1
1

0
0
0
1
14

0
0
0
0
2

0
0
1
0
28

0
0
0
0
0

0
0
0
0
1

-

1

0

1
1
1
1
49

0
0
0
1
3

Arizona
Yavapai Co.
Gila Co.
Graham Co.
Mohave Co.
Yuma Co.

-

-

California
San Bernardino
Co.
Riverside Co.
Imperial Co.
Mexico

0
0
0

0
2
1

1
0
2

0
4
0

6

1
1

0
0

0
0

Northwest

0

1

Coahuila
West-central

2

0

-

9

0
0
0

0
1
0

-

1
7
11

0
6
1

0
0

0
0

0
0

0
0

-

1
1

0
0

2

0

0

0

0

-

2

1

0

0

0

0

0

0

-

2

0

2

0

0

0

0

0

0

-

2

0

22

13

25

~8

47

2

6

-

100

23

Baja
Northwest
Unknown
Chihuahua

Sinaloa

--

Northwest
Totals

-------

l/Area closed to goose hunting.

I

~

0
I

�-41-

Mortality
The birds released in 1972 suffered the least amount of first year hunting
mortality of all groups that have been released in North Park. Only 4
percent o f the birds released in 1972 were reported recovered compared to
11.4, 10.2 and 19.5 percent in 1969, 1970, and 1971, respectively.
The overall first year recovery rate average for released birds in North Park is
12 ••0 percent.
South Park-Antero

Reservoir

Canada goose goslings were released on Antero Reservoir in Park County for
the first time in July 1972. A release of 26 males and 22 females was
made on July 10 followed by 13 males and 15 females liberated on July 28.
Restoration efforts in South Park are designed to supplement a small nesting
population which is now present on Antero Reservoir.
An inventory of geese
on Antero in June 1972 found 33 adults and 5 broods
with a total of 17
goslings.
Five of the birds released on Antero Reservoir were reported taken during
the fall of 1972. Two birds were reported taken in Reeves County along the
Pecos River in southwestern Texas.
The other three birds were reported
taken in Colorado, 1 in Fremont County near Canon City, 1 in the San Luis
Valley, and 1 in Pueblo County near Avondale.
Totten

Reservoir

Thirty-six and 38 geese were released on Totten Reservoir in June of 1970
and 1971, respectively.
During the winter of 1970-71 the birds generally
stayed in the release area.
During the 1971-72 winter the birds migrated
generally straight south and stayed on the Gila River near Stafford, Arizona
(Szymczak 1972).
At least 23 birds returned from the 1971-72 excursion.
In ~uly 1972, 32 goslings were added to the Totten Reservoir flock. The
birds stayed in the release area until early January, at which time they
left for about two weeks.
According to band recoveries, the birds migrated
into eastern Yavapai County, Arizona where 3 birds were harvested along the
Verde River.
Again, as in 1972, at least 23 birds returned to the release
site after about a two week absence.
Reports in March 1973 indicate no utilization
erected in the release area.

of artificial

structures

�-42-

LITERATURE CITED

Szymczak, M. R. 1971. ExperLmen t af studies on improving status of Canada
goose populations. Colo. Div. Game, Fish and Parks, Game Res. Rep.,
Fed. Aid Project W-88-R. October. 115-127 p.
Szymczak, M. R. 1972. Experimental studies on improving status of Canada
goose populations. Colo. Div. of Wildl., Game Res. Rep., Fed. Aid
Project W-88-R. October. 39-49 p.

Prepared by

m.:d. ,L l?~,()4

Michael R. Szymc~~
Assistant Wildlife Researcher

�October,

-43-

1973

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~-------Migratory

W-88-R-18

Project No.
Work Plan No.

2

Job No.

Bird Investigations
5

Job Title

Arkansas

Valley Canada Goose Flock Management

Period Covered:

November

1, 1972 through March 31, 1973

Personnel:

Studies

R. Baker, J. Carsella, G. Claasen, G. Eyre, R. Forbes, R. Kitzmiller, P. Olson, S. Porter, D. Potts, R. Rosette and M. szymczak.

ABSTRACT
According to aerial surveys the size of the wintering population of Canada
geese in southeast Colorado reached a high of 47,500 in late November.
Results
of coordinated inventories throughout the short grass prairie Canada goose
wintering flock in early January indicated the total population numbered
257,500 birds.
The age composition of the harvest in southeast Colorado
during the 1972-73 hunting season indicated that short grass prairie geese
had a poor production year in 1972. Trapping operations in southeast ,Colorado in January, 1973 resulted in 564 birds being banded.
During the 1971-72
hunting season the percent of recoveries of birds banded post-season in
southeast Colorado reported taken on the Canadian prairies in Alberta and
Saskatchewan reached a record low of 33.7 percent, the southeast colorado
harvest reached a record 46.5 percent.
Average annual mortality estimates
for Canada geese banded in southeast Colorado remain at approximately 25
percent.

��-45-

ARKANSAS

VALLEY

CANADA GOOSE FLOCK MANAGEMENT
Michael

STUDIES

R. Szymczak

The final report for the Arkansas Valley Canada goose flock management study
is in preparation.
Data presented in this progress report is being incorporated into the final report.
Therefore, this progress report will be
abbreviated in order to prevent needless repetition.

P. S. OBJECTIVE
To investigate

the status of Canada geese wintering

in southeast

Colorado.

SEGMENT OBJECTIVES
1.

Estimate

the size of the wintering

2.

Estimate

the age composition

3.

Estimate the hunting pressure
types of harvest regulations.

4.

Investigate the annual mortality and the migration and/or harvest
of the flock under varying types of harvest regulations.

METHODS

flock.

of the flock.
on, and harvest

of the flock under varying

pattern

AND MATERIALS

Coordinated inventories were conducted throughout the short grass pra~r~e
populations wintering range in Nebraska, Colorado, Oklahoma, Texas, and New
Mexico on December 12 and again in early January.
In addition, periodic
counts were made in Colorado throughout the hunting season.
Although these
periodic counts are not included in the procedures, results will be reported
as a part of this job.
Tail fans from geese bagged were collected
Management Area check stations.
Hunting pressure and harvest information
rado Small Game Harvest Survey.

at the Two Buttes and Lamar-Eads

was obtained

from the regular

Colo-

A total of 564 Canada geese were banded during trapping operations at Turk's
Pond after the close of the hunting season, in order to obtain information on
age ratios, mortality rates and migration patterns.

�-46-

RESULTS AND DISCUSSION
Size and Distribution

of the Wintering

Population

According to aerial surveys, the size of the 1972-73 wintering population
of Canada geese in southeast Colorado was substantially reduced in comparison to 1970-71 and 1971-72 levels.
The largest count in southeast Colorado
was obtained in late November when 47,500 birds were observed (Table 1).
This compares with high counts of 90,550 on December 11, 1971 and 87,750 on
December 21, 1972. Reduced numbers were thought to be largely the result
of continuous inclement weather in the area. Again, as in previous years,
the majority of the birds within the state congregated iri the Turk's PondTwo Buttes area (Table 1).

Table 1.

Results

of Canada goose aerial surveys,

southeast

Colorado,

1972-73.

Date
Area

Nov. 10

Nov. 28

Dec. 12

3,500

1,100

50

200

Henry Lake

o

300

o

o

Blue Lake

400

o

o

o

Nee Noshe

2,000

11,000

3,000

o

Upper Queens

2,200

200

700

100

Two Buttes Reservoir

1,300

11 ,500

10,000

13,000

Turk's Pond

10,900

10,000

18,000

23,000

3,900

13,100

900

5,000

32,650

41,300

Meredith

Reservoir

Jan. 12

Eads Group:

John Martin

Reservoir

Bonny Reservoir

Total

300
24,200

47,500

Results of the coordinated January inventory were surprising.
The survey
totaled 257,500 geese (Table 2), with only 41,300 birds being in Colorado
(Table 3). The large majority of the birds were located in the Texas panhandle
where a much larger area was surveyed than in previous years.
Coordinated
surveys in the same areas during the 1973-74 season should indicate whether
the short grass prairie population has actually reached the 250,000 bird level.

�-47Table 2. Post-hunting season status of Short Grass Prairie
population, 1960-73, data generally from regular mid-winter

Canada goose
inventories.

Year

Number

Number

of Birds

Year

of Birds

1960

77,709

1967

111 ,452

1961

103,355

1968

127,903

1962

80,133

1969

112,399

1963

93,940

1970

147,414

1964

81,221

1971

152,734

1965

103,435

1972

134,500

1966

llO ,485 1/

1973

257,500

1/ Inventory of Feb. 15, 1966 substituted
Table

3.

January

Inventory

for unsatisfactory

of Canada geese, Arkansas

Valley,

January

Colorado

inventory.

1948-73.

Year

Goose Count

Year

Goose Count

Year

Goose Count

1948

4,798

1957

24,617

1966

38,635

1949

12,286

1958

35,894

1967

29,835

1950

13,170

1959

44,660

1968

42,682

1951

19,320

1960

37,394

1969

29,201

1952

30,463

1961

31,360

1970

63,444

1953

20,236

40,250 1/

1971

62,720

1954

20,280

1963

35,889

1972

76,400

1955

25,llO

1964

33,750

1973

41,300

1956

24,212

1965

37,693

1/ Inventory

of February

7, 1962 substituted

for January,

1962 inventory.

1/

of February

15, 1966 substituted

for January,

1966 inventory.

Inventory

Y

�-48-

Age Composition
Goose tail fans collected at the Two Buttes and Lamar-Eads Management areas
during the 1972-73 hunting season indicated that an overwhelming portion of
the harvest was made up of adults (Table 4). Comparative estimates obtained
from tail fans collected through the Bureau of Sport Fisheries and Wildlife's
Parts Collection Survey indicated that 76.2 percent of the harvest in southeast Colorado was composed of adults.
In addition, only 12 percent of 564
Canada geese captured during post-season banding operations at Turk's Pond
in southeast Colorado were classified as immatures (Table 5). All the above
measurements indicate extremely poor production during the spring of 1972.

Hunting

Pressure

and Harvest

The results of the Colorado small game harvest survey for 1972-73 are not
available at this time. The information will be presented in the final
report.

Banding

Investigations

Trapping efforts post-season in southeast Colorado were concentrated in the
Turk's Pond-Two Buttes area.
However, all birds were captured at Turk's Pond.
The sex and age composition of each catch are presented in Table 5. Approximately 48 percent of the birds captured were female.
Distribution

of Harvest

During the 1971-72 season, the percent of recoveries of birds banded postseason in southeast Colorado reported taken on the Canadian prairies in
Alberta and Saskatchewan reached a record low of 33.7 percent (Table 6). The
percent harvest in southeast Colorado, which on a long term basis has
averaged about 30 percent, reached a record high of 46.5 percent.
The high
percentage harvest in southeast Colorado was no doubt the result of large
numbers of geese being present in the state throughout the 1971-72 season
(Szymczak 1972).
Mortality

Estimates

Average annual mortality estimates for Canada geese banded in southeast Colorado calculated by the composite dynamic or relative recovery rate method
remain at approximately 25 percent for any and all age groupings.
Tables
are withheld from this report and will be included in the final report.
LITERATURE

CITED

Szymczak, M. R. 1972.
Arkansas Valley Canada goose flock management studies.
Colo. Div. of Wildl., Game Res. Rept., Fed. Aid Proj. W-88-R.
Oct.
p. 51-75.

l

I }/f
'.
I
by _1,--' t:-&gt;:.i.-'-(-",.'-A-"'("-'(-"" -; "'-'._/_;....;{'-'-.----'~::....-:;""'&lt;;,.;,~"f.. ,.;....f_(..,&lt;.i...:; •...;;·~::.../-43'"""~. ""./•.••
;:••
Michael R. SzyritC'z'!lk
.',
Assistant Wildlife Researcher
I'

Prepared

/.

.;;()

_

�Table 4.

Age composition of the goose harvest in selected areas, southeast Colorado.

Year

Two Buttes (Firing Line)
Adult
Immature
No.
Percent
No.
Percent

1951-52

356

1957-58

46.8

404

Lamar-Eads (Deco~)
Adult
Immature
No.
Percent
No.
Percent

Meredith Area
Adult
Immature
No.
Percent
No.
Percent

53.2

No Data

1958-59

929

47.9

1,010

52.1

1959-60

377

36.5

655

63.5

1960-61

612

45.5

732

54.5

1961..;62

527

55.7

419

44.3

1962-63

204

48.9

213

51.1

1963-64

377

52.1

346

47.9

1964-65

442

63.1

259

36.9

1965-66

571

62.7

340

37.3

1966-67

217

45.3

262

54.7

1967-68

245

59.9

164

40.1

1968-69

136

61.8

84

38.2

79

51. 3

75

48. 7

48

1969-70

110

55.3

89

44.7

143

35.9

255

64.1

1970-71

630

40.6

922

59.4

77

27.4

204

1971-72 1./

804

63.5

463

36.5

128

33.7

1972-73

62

92.5

102

71. 3

5

7.5

I

~

\0
I

24

33.3

18

66.7
40.0

27

60.0

72.6

42

45.7

50

54.3

252

66.3

63

51. 6

59

48.4

41

28.7

1/ Data for Two Buttes also includes some birds bagged in the Turk's Pond area.
some birds bagged in the John Martin Reservoir area.

Data for Meredith area includes

�-50-

Table 5.
southeast

Sex and age composition
Colorado, 1973.

Location

and Date

Turk's

Total

Adult

of Canada geese trapped

Male
Immature

Adult

Female
Immature

post-season

in

Total

Percent
Immature

Pond

1-18-73

101

8

80

3

192

5.7

1-20-73

37

6

38

2

83

9.6

1-31-73

47

18

68

7

140

17.9

2-7-73

20

2

29

9

60

18.3

2-12-73

44

9

32

4

89

14.6

249

43

247

25

564

12.0

�Table 6.

Percentages

of total band recoveries,

Area

1951-55

Arkansas

Five Year Averages
1956-60
1961-65

Valley post-season

1966-67

1967-68

-

-

bandings,

by area and year of recovery,

Recove!2: .Ye ar
1968-69
1969-70

1970-71

all bandings.

1971-72

Total No.
Recoveries

Percentage
of Total
Recoveries

274

8.8

Far North Above 530
N. W. Territories
Alberta
Saskatchewan
Total
Provinces

-

-

9.8

-

-

-

1.0
12.4

0.5
6.1

7.5

2. 1
7.1
0.7
9.9

13.4

6.6

0.5
2.4
0.5
3.3

23.4
17.0

21.9
25.7

15.7
24.7

11.9
21.8

841
511

27.0
16.4

0.5

5

0.2

1.9
1.0
1.4
5.2

21
4
19
7
186

0.7
0.1
0.6
0.2
6.0

44.6
1.9
46.5
1.0
0.5
0.5

946
14
16
28

30.3
0.4
0.5
0.9

-

7.1

10.2

10.4

35.3
10.5

28.9
11.5

19.4
16.4

14.2
24.6

0.7

1.5

0.7

Below 530

Alberta
Saskatchewan
B.C., Manitoba,
Ontario
Central

-

28.6
18.3
0.1

I
VI

Flyway

Montana
North Dakota
Wyoming
South Dakota
Nebraska
Colorado
Southeast and Other
Northcent ra1
Total
Kansas
Oklahoma
New Mexico

0.8
0.1
0.7
0.1
5.4

-

0.4

0.3

0.7

0.6

0.5
0.3
5.8

0.7
0.7
9.7

6.5

-

-

-

25.5
0.4
0.9
1.5

30.9
0.3
0.3
0.7

28.4
0.5
0.4
0.4

32.4
0.7

1.4
0.7
0.7

1.9

1.9
4.8

0.5
0.5
4.5

9.0

4.3

27.6
1.5
29.1

29.1
2.8
31.9

19.0
1.0
20.0
1.0

34.3
3.0
37.9
1.0

1.5
0.7

1.4

1.0

2.0

------~-----------------------------------------------------------------------------------------~-------------------------------------------------------

•...
I

�Table 6.

Percentages

of total

band recoveries,

Arkansas Valley post-season

Five Year Averages
1951-55
1956-60
1961-65

Area

1966-67

1967-68

7.5
0.7

7.5

bandings,

by area and year of recovery,

Recove!}: Year
1968-69
1969-70

all

bandings

(continued) .

Total No.
Recoveries

Percentage
of Total
Recoveries

3.8
1.7
0.5
6.0

1970-71

1971-72

2.5
3.0

2.4
1.0

5.5

3.3

117
53
17
187

1.0

1.4

55

1.8

3

0.1

1

0.03

Cent ra1 Fly_way (continued)
Texas
Panhandle
Waggoner Ranch
Gulf Coast
Total
Pacific

Flyway

Mississippi

Flyway

2.3
2.0
0.8
5.1

2.8
2.2
0.2
5.2

4.6
1.5
0.5
5.5

2.4

2.2

0.8

8.2

0.1

Mexico

0.1

Total Number
Of Recoveries

748

677

795

134

2.2
9.7

5.0
0.7
1.4
7.1

0.7

1.4

0.7

0.7

134

141

6.7
1.0
1.0
8.7

105

198

211

3,118

-

_.-".

I
Vl
N
I

�october,

-53-

1973

JOB PROGRESS REPORT

.COLORADO

State of

2

Work plan No.
Job Title

Studies

Period Covered:
Personnel:

Migratory

W-88-R-18

Project No.

Bird Investigations

6

Job No.

of Canada Goose Populations

in Colorado

Transplant

Areas

April 1, 1972 through March 31, 1973

Charles Bryant and Staff, Monte Vista National Wildlife Refuge;
D. Benson, C. Brown, R. Clark, E. Cochran, J. Corey, G. Crawford,
G. East, J. Frothingham, H. Funk, B. Goetze, J. Hatfield, T. Henry,
J. Hobbs, C. Leonard, J. Lorentzson, T. Lynch, F. Marcoux,
J. Monarch, D. Owens, S. Palm, J. Pogorelz, C. Roberts, W. Russell,
G. Saville, L. Searle, E. Wagner and M. szymczak, Colorado Division
of Wildlife.

ABSTRACT
During the 1972-73 hunting season, 95 geese banded outside Colorado were
reported recovered in north-central Colorado.
As in past years, the Cypress
Hills area of Saskatchewan and Phillips County area of Montana were well represented by banded birds in the north-central Colorado harvest.
On the Canadian
prairies, Saskatchewan was the major recovery area during the 1972-73 hunting
season for birds banded in north-central Colorado in January, 1971. First year
recoveries of birds banded in January, 1972 in north-central Colorado showed an
equal distribution between Alberta and Saskatchewan.
The recovery distribution
of birds banded in north-central Colorado indicated a shift by a portion of
the 1972-73 Hi-Line wintering population from north-central Colorado to the
southeast portion of the State.
Coordinated counts throughout the Hi-Line
wintering range indicated population numbers were below 1971-72 levels.
An
estimated 9,938 hunters harvested 5,270 geese in north-central Colorado during
the 1972 season, a 64 percent decline in harvest compared to the 1971-72 season
level. Five birds banded at Wheatland Reservoir, Wyoming were reported recovered
in the San Luis Valley during the 1972-73 special season.
An estimated 311
hunters bagged 255 geese during the special season in the San Luis Valley. Aerial
surveys in the San Luis Valley indicated there were between 1,000 and 1,600
geese in the Valley during the 1972-73 season.
An estimated 182 hunters harvested
58 geese in the special west-central colorado goose permit area. Approximately
41 percent of the birds harvested in west-central Colorado were taken within
five miles of Highline Reservoir.
Goose inventories in west-central Colorado
totaled 329 on November 21, 1972 and 1,183 on January 11, 1973.

��-55-

STUDIES OF CANADA GOOSE POPULATIONS IN COLORADO TRANSPLANT AREAS
Michael R. Szymczak
P. S. OBJECTIVE
To investigate the status of resident and migrant Canada goose flocks and
their interrelationships in areas in which populations have been established
through transplant programs in Colorado.

SEGMENT OBJECTIVES
la.

To examine migration routes and/or harvest patterns and distribution of
Canada geese wintering in northcentral Colorado and the San Luis Valley.

lb.

To determine breeding areas of Canada geese wintering in northcentral
Colorado.

2.

To estimate hunting pressure on, and hunter harvest of Canada geese in
northcen tral Colorado, the San Luis Valley, and westcen tral Colorado.

3.

To make recommendations for continuing Canada goose hunting seasons on
population in westcentral Colorado, northcentral Colorado and the San
Luis Valley.

METHODS AND MATERIALS
Band recoveries listed on computer printouts provided periodically by the
Migratory Bird Populations Station were sorted and plotted as to banding and
recovery location, sex, and age at time of banding, and estimated breeding
area for birds associated with the San Luis Valley and northcentral Colorado.
Canada geese in northcentral Colorado were counted on November 6, November 22,
and December 20, 1972; and January 10, 1973. In the San Luis Valley birds
were counted on October 25, November 22, and December 18, 1972; and January
8, 1973.
A mechanically random sample of hunters issued permits to hunt geese in the
northcentral Colorado "Special Permit" goose hunting area were mailed questionnaires inquiring about their hunting activity and success. All of the hunters
holding special permits to hunt geese in the San Luis Valley and westcentral
Colorado were mailed similar qQestionnaires. All data received were tabulated and statistically analyzed.
Recommendations concerning regulations for the 1973-74 goose seasons in northcentral Colorado, the San Luis Valley and westcentral Colorado were formulated
and submitted to management personnel.

�-56-

RESULTS
Northcentral

Colorado-The

AND DISCUSSION
Hi-Line

Population

Breeding

Range

Foreign Recoveries
During the 1972-73 hunting season 95 geese wearing foreign bands were
reported recovered in northcentral Colorado.
An additional 15 foreign
banded birds were recaptured during the post-season banding operations.
Both
types of recoveries are classified as to general area of banding if in a
Province,
or County if in the United States in Tables 1 and 2, respectively.
Banding locations of wild trapped locals recovered in northcentral Colorado
during the 1972-73 season are presented in Table 3.
AnalYSis of banding areas presented in Tables 1, 2 or 3 did not reveal any
new possible production areas for Hi-Line Canada geese.
There continues to
be significant numbers
of recoveries of birds banded in the Cypress Hills
area of southeast Saskatchewan, and from Phillips County, Montana where
annual banding programs are in progress.
A renewed banding effort in southern
Alberta during the summer of 1972 produced only one recovery in the Hi-Line
area in Colorado.
Collection of banding area information during the 1972-73
season was hampered by a reduced harvest in the Hi-Line area in Colorado.

Win ter Banding-Northcentral

Colorado

In January 1972, 1,041 Canada geese were banded post-season in northcentral
Colorado.
Size 8 bands were placed on 580 birds that were subjectively
considered large, while 461 small birds were banded with size 7 bands.
These
bandings bring the two year total of birds banded post-season in northcentral
Colorado to 2,034, 1,326 with size 8 and 708 with size 7 bands.
The distribution of second year recoveries from birds banded in January 1971
varied considerably from first year recoveries of that banding cohort.
During the 1971-72 season, the number of recoveries in Alberta and Saskatche~
wan of birds banded with size 8 bands were nearly equal (Table 4). During
the 1972-73 season, reported recoveries on the Canadian prairies of birds
wearing size 8 bands came almost exclusively from Saskatchewan.
In addition,
Alberta was not predominant in the recovery of birds wearing size 7 bands
during the 1972-73 season, as it had been during the 1971-72 season (Table 4).
First year recoveries of birds banded in January 1972 showed an equal distribution of recoveries between the two provinces regardless of band size (Table
5). A similar provincial distribution was noted for first year recoveries of
birds wearing size 8 bands from January 1971 bandings in northcentral Colorado (Table 4).
In general, recoveries of Colorado-banded birds during the 1972-73 season
did not reveal any new potential Hi-Line production areas.

�-57-

Table 1. Banding areas outside Colorado of Canada geese recovered in
northcentral Colorado during the 1972-73 hunting season.
Direct

Indirect

District of Keewatin
Thelon River

o

1

Alberta
Dowling Lake Area
Brooks Area

o

6

1

1

o

1

10

3

Montana
Phillips County
Fergus County
Garfield Coun ty
Sheridan County
Bighorn County
Treasure County

22
0
1
1
2
1

15
1

Wyoming
Goshen County
Sheridan County
Albany County
Fremont County

2
2
1
1

0
0
1
1

Kansas
Phillips County

1

0

New Mexico
Mora County
San Miguel County
Socorro County

0
0
1

2
2

1

o

1

o

Area

Saskatchewan
Moose Jaw Area
Eyebrow Lake
Cypress Hills Area
CENTRAL FLYWAY

4

1
3
0

4

PACIFIC FLYWAY
Idaho
Canyon County
Utah
Dagget County
Weber County

Total

1

o

49

46

�-58-

Table 2. Banding areas outside Colorado of Canada geese recaptured, postseason, in northcentral Colorado, January 1973.
Area
Alberta
Edmonton-Camrose Area
Saskatchewan
Cypress Hills Area

Direct

Indirect

o

1

1

3

o
o

5

1

1

o

1

1

o

3

12

CENTRAL FLYWAY
Montana
Blaine Coun ty
Phillips County
New Mexico
Socorro County
Texas
Randall County

1

PACIFIC FLYWAY
Idaho
Canyon County
Total

Migration Routes and Harvest Patterns
During the 1972-73 hunting season, the percentage of total recoveries of birds
banded post-season in northcentral Colorado increased over 1971-72 levels in
Saskatchewan for both large (size 8) and small (size 7) geese (Table 6).
A slight decline in harvest was noted in Montana and a major decline was
apparent in northcentral Colorado. The recovery distribution in Colorado
during the 1972-73 season indicates a major shift occurred from the northcentral area to southeast Colorado (Table 6). The Colorado harvest distribution confirms that a portion of the Hi-Line population moved south and
southeast, apparently in response to persistent inclement weather conditions
in northcentral Colorado. Recoveries of large birds (size 8 bands) occurred
in non-traditional goose harvest areas along the foothills in southern
Colorado (Pueblo and Las Animas counties) •. Plotting the distribution of
recoveries north of Colorado did not reveal any new possible breeding areas
for Hi-Line geese.

�-59-

Table 3. Banding locations of Canada geese classified as wild-trapped locals
which were recovered or recaptured in northcentral Colorado during the
winter of 1972-73.

Year of Banding
Location

Before
1967

Alberta
Edmonton-Camrose Area
Hay Lakes Area
Dowling Lake Area
Brooks Area
30 mi. S. Bow River
30 mi. SE
Saskatchewan
Cypress Hills Area
N. of Eastend
Vidora Area
Maple Creek Area
Masefie1d Area
Divide Area
Cypress Hills Park

1967

1968

1

1969

1970

1971

1

1
2

1

1972

1
1

1
3
1

1
2

2

3

3
1

18
1
1

1

CENTRAL FLYWAY
Montana
Blaine County
Phillips County
Bighorn County
Treasure County
Wyoming
Goshen County

1
4

2

3
1

1

PACIFIC FLYWAY
Utah
Daggett County

1

�-60-

Table 4. Percentage distribution of recoveries of Canada
geese banded
post-season in northcentral Colorado, January 1971.

Location

Hun ting Season
Band Size

Alberta
Saskatchewan

1971-72 (First Year~
B
7

1972-73 (Second Year)
8
7

13.B (11)

40.0 (6)

2.4 (1)

12.5 (1)

lB. B (15)

6.7 (1)

33.3 (14)

12.5 (1)

7.5 (6)

13.3 (2)

4.B (2)

12.5 (1)

2.5 (2)

6.7 (1)

0.0 (0)

(0)

2.5 (2)

(0)

2.4 (1)

(0)

2.5 (2)

(0)

2.4 (0)

(0)

46.3 (37)

20.0 (3)

35.7 (15)

25.0 (2)

0.0 (0)

13.3 (2)

7.1 (3)

37.5 (3)

0.0 (0)

(0)

4.B (2)

(0)

1.3 (1)

(0)

2.4 (1)

(0)

0.0 (0)

(0)

2.4 (1)

(0)

2.5 (2)

(0)

2.4 (1)

(0)

Montana
South Dakota
Wyoming
Nebraska
Colorado
Northcentral
Southeast
New Mexico
Arizona
Nevada
California

Total Recoveries

BO

15

42

B

Distribution
Coordinated counts throughout the Hi-Line population wintering area indicated
a reduced total population in comparison to the 1971-72 levels (Table 7).
The January count, in 1973, however, was nearly identical to the 1972 total.
Weather conditions, which induced unusual movement as was discussed above,
definitely created some census problems. However, according to the figures
presented, the distribution by state in wintering areas remained about the
same as in previous years.

�-61-

Table 5. Percentage distribution of recoveries of Canada geese banded postseason in northcentral Colorado, January 1972, and recovered during the
1972-73 season.

Area

Band Size 8

Band Size 7

Total

Alberta

24.5 (12)

42.1 (8)

29.4 (20)

Saskatchewan

22.4 (11)

31.6 (6)

25.0 (17)

Manitoba

2.0 (1)

(0)

1.5 (1)

Montana

6.1 (3)

(0)

4.4 (3)

Idaho

2.0 (1)

(0)

1.5 (1)

Wyoming

2.0 (1)

(0)

1.5 (1)

Nebraska

2.0 (1)

(0)

1.5 (1)

Northcentral

24.5 (12)

5.3 (1)

19.1 (13)

Southeast

12.2 (6)

15.8 (3)

13.2 (9)

Other

2.0 (1)

(0)

1.5 (1)

Arizona

0.0 (0)

5.3 (1)

1.5 (1)

Total Recoveries

49

19

Colorado

68

Extreme cold weather resulted in an unusual distribution of geese in northcentral Colorado from mid-December through early January.
The birds moved
in increasing numbers to College Lake in the Fort Collins area, and Va1mont
Reservoir in the Boulder area (Table 8). By January 10, 1973, nearly 80
percent of the geese in northcentral Colorado were located on these two areas.
The extensive movement of geese to Valmont Reservoir resulted in there being
for the first time, more geese in the Longmont-Boulder-Denver
area during
late December and early January than in the Fort Collins-Loveland
area (Table
9). Water releases from the adjacent power plant at Valmont maintains open
water on the reservoir regardless of weather conditions.

�-62-

Table 6.

Percentage distribution of recoveries of Canada geese
banded postseason in northcentral Colorado, 1970-71 and 1971-72.
Hunting
Season
Area

Size 8

Alberta

1972-73
Size 7

13.8 (11)

1972-73
Size 8
Size 7

Size 8

Total
Size 7

40.0 (6)

14.3 (13)

33.3 (9)

14.0 (24)

35.7 (15)

6.7 (1)

27.5 (25)

25.9 (7)

23.4 (40)

19.0 (8)

0.0 (0)

1.1 (1)

0.0 (0)

0.6 (1)

0.0 (0)

13.3 (2)

5.5 (5)

3.7 (1)

6.4 (11)

7.1 (3)

6.7 (1)

0.0 (0)

0.0 (0)

1.2 (2)

2.4 (1)

0.0 (0)

2.2 (2)

0.0 (0)

2.3 (4)

0.0 (0)

0.0 (0)

1.1 (1)

0.0 (0)

0.6 (1)

0.0 (0)

0.0 (0)

2.2 (2)

0.0 (0)

2.3 (4)

0.0 (0)

46.3 (37)
0.0 (0)
0.0 (0)

20.0 (3)
13.3 (2)
0.0 (0)

29.7 (27)
9.9 (9)
1.1 (1)

11.1 (3)
22.2 (6)
0.0 (0)

37.4 (64)
5.3 (9)
0.6 (1)

14.3 (6)
19.0 (8)
0.0 (0)

0.0 (0)

0.0 (0)

2.2 (2)

0.0 (0)

1.2 (2)

0.0 (0)

0.0 (0)

1.1 (1)

3.7 (1)

1.2 (2)

2.4 (1)

0.0 (0)

1.1 (1)

0.0 (0)

0.6 (1)

0.0 (0)

0.0 (0)

1.1 (1)

0.0 (0)

1.8 (3)

0.0 (0)

0.0 (0)

0.0 (0)

0.0 (0)

0.6 (1)

0.0 (0)

0.0 (0)

0.0 (0)

0.0 (0)

0.6 (1)

0.0 (0)

15

91

27

Saskatchewan

18.8 (15)

Manitoba

0.0 (0)

Montana

7.5 (6)

South Dakota

2.5 (2)

Wyoming

2.5 (2)

Idaho

0.0 (0)

Nebraska

2.5 (2)

Colorado
Northcentral
Southeast
Other
New Mexico
Arizona
Nevada
California
Pennsylvania
Unknown
Total

1.3 (1)
0.0 (0)
2.5 (2)
1.3 (1)
1.3 (1)
80

171

42

�-63-

Table 7.

Results of Hi-Line Canada goose population inventories, 1972-73.

Location

November 6

November 22

December 20

January 10
---------

Saskatchewan 1/
Montana

10,130

8,594

1,050

920

Wyoming

4,971

3,000

1,956

1,272

Colorado

17,520

31,863

34,356

33,709

539

859

2,434

2,409

Totals

33,160

44,316

39,796

38.310

Comparative
Totals 1972-73

61,054

62,552

61,236

37.113

New Mexico

!/ No counts in Saskatchewan.

Table 8.
Area

Results of northcentral Colorado goose surveys, 1972-73.
November 6

1/ November 21

December 20

January 10

Ft. Collins-Loveland
Lindenmeir Lake
1,200
Reservoir No. 8
1,300
Reservoir No. 8
Annex
0
Elder Reservoir
0
Reservoir No. 5
0
Reservoir No.6
300
Douglas Reservoir
0
Rocky Ridge Res.
0
Cement Plant Res.
0
Terry Lake
500
College Lake
600
Claymore Lake
0
Sterling Pond
0
Watson Lake
0
Dean Lake
0
Greenwalt (Nelson)
Lake
0
Fossil Creek Res. 1,200
Boyd Lake
30

1,272
0

275
43

0

0
0
21
195
100
18
585
2,789
3,400
1,750
80
1,125
43

72
0
0
0
0
0
0
875
7,270
81
114
137
107

0
0
0
0
0
0
0
720
8,650
43
0
800
0

3
3,000
475

0
25
0

0
20
0

2

-----------------------------------------------------------------------------

�-64-

Table 8.

Results of northcentral Colorado goose surveys, 1972-73 (continued).

Area

November 6 1./ November 21

December 20

January 10

2,000
0
750
0
0
0
0
0
610
0

30
21,118

2,000
0
2,050
40
19
110
0
1
562
1
59
0
13,841

0
13,595

0
360
0
0
600
1

0
322
38
0
200
0

0
0
0
12
50
0

30
10
3,000
180
60
0
0
4,241

0
0
17,000
42
0
57
0
17,659

0
0
18,000
0
0
0
27
18,089

124
81
960
0
257
23
73

30
0
1,561
40
0
0
0

33
0
31
0
309
0
0

5
0
1,523

0
0
1,631

0
247
620

Ft. Collins-Loveland (continued)
Horseshoe Lake
0
Hollister Lake
1,000
New Windsor Reservoir
5,500
Woods (Eaton) Lake
700
Timnath Reservoir
1,300
Cobb Lake
1,600
Black Hollow Reservoir
0
Park Wood Lake
0
Boxelder Reservoir
0
Warren Reservoir
0
Horsetooth Reservoir
Owl Creek Reservoir
0
Sub-total
15,230

0
353
4,729
100
80
330
20
88
530
2

Boulder-Longmont Area
Ish Lake
Terry Lake (Longmont)
McIntosh Lake
Foothills Reservoir
Faivre Ponds
Swede Lake
Lonetree Reservoir
Baller Lake
Boulder Reservoir
Valmont Reservoir
McCall Lakes
Gaynor Lakes
Union Reservoir
Stamp Pond
Sub-total

15
0
0
0
0
0
25
0
0
1,200
0
0
0
0
1,240

Denver Area
Sloans Lake
Standley Lake
Bowles Lake Area
Youngfield Lake
Denver City Park
Gravel Ponds
Washington Park
Tule Lake S. Colo. Blvd.
at Quincy
Various Fields
Sub-total

-----------------------------------------------------------------------------

�-65Table 8.

Results

of northcentral

Area

November

Brighton-Greeley-Ft.

Colorado

goose surveys,

61:./ November

21

1972-73 (cont.).

December

20

10

January

Morgan Area

Barr Lake
Horsecreek Reservoir
Prospect Reservoir
Latham Reservoir
Milton Reservoir
Empire Reservoir
Riverside Reservoir
Jackson Reservoir
S. Platte River
near Jackson Res.
Sub-total

Grand Total

100
0
250
0
0
0
500
100

1,400
2,700
0
6
30
25
450
370

225
0
0
0
0
0
0
0

75
0
0
400
0

100
1,050

0
4,981

1,000
1,225

880
1,405

17,520

31,863

34,356

33,709

0

0
50

1:./ No count in the Denver area.

Hunting

Pressure

and Harvest

An estimated 9,938 hunters harvested 5,270 geese in northcentral Colorado
during the 1972-73 season (Tables 10 and 11). Declines in harvest from the
1972-73 season levels were recorded in all counties (Table 11). The average
seasonal bag per active hunter declined to about one-half a bird, even though
the average days hunted remained comparatively high (Table 10). Nearly 70
percent of the active hunters failed to bag a goose in the permit area during
the 1972-73 season (Table 12). Comparative statistics concerning hunting
pressure and harvest by County for the 1971-72 and 1972-73 hunting seasons
are presented in Table 13.

San Luis Valley
Foreign Band Recoveries
During the 1972-73 goose season, nine birds that were banded outside of the
San Luis Valley were reported recovered in the Valley.
Five of the birds
were banded at Wheatland Reservoir in Albany County, Wyoming (Table 14).
Consistent recoveries of Wheatland Reservoir banded birds in the San Luis
Valley essentially confirm that Wheatland Reservoir, exclusively a molting
area for geese, is being utilized by molt migrants from the San Luis Valley
population.
The other three recoveries during the 1972-73 season were banded
during post-season banding operations in New Mexico, two birds from Soccoro
County in the Rio Grande Valley and one from Mora County.
For the first time
in three years, no birds banded in Hi-Line production areas were recovered in
the San Luis Valley.

�-66-

Table 9.

Winter inventories of the Colorado Hi-Line Canada goose population.

Count Date

Fort Co11insLoveland

Longmont-BoulderDenver

Brighton-Gree1eyFort ~rgan

Total

1967
January 9

9,739

2,883

991

13,613

1968
January 10

12,217

4,029

678

16,924

1968-69
November 20
December 19
January 2 &amp; 13

15,848
20,905
19,693

3,461
4,236
4,874

2,667
1,170
775

21,976
26,311
25,342

1969-70
November 5
November 28
December 23
January 6

8,737
31,350
18,522
30,650

2,255
3,782
5,668
5,060

390
1,374
1,259
1,914

11,382
36,506
25,499
37,624

1970-71
November 4
November 23
December 22
January 6

12,612
29,970
36,034
19,879

1,690
16,710
12,664
15,566

348
1,370
3,055
2,425

14,650
48,050
51,753
37,870

1971-72
November 4
November 23
December 21
January 10

25,699
31,072
31,516
19,117

1,815 !/
9,181
19,525
10,742

6,905
13,525
4,476
3,661

34,419
53,788
55,517
33,520

1972-73
November 6
November 22
December 20
January 10

15,230
21,118
13,841
13,595

1,240 1/
5,764
19,290
18,709

1,050
4,981
1,225
1,405

17,520
31,863
34,356
33,709

1/ Denver area not included.

�-67Table 10.

Hunter activity and success in permit .area.
Average
Days Hunted

Average
Seasonal Bag/Hunter

1,427

6.1

0.48

2,335

1,578

6.5

0.53

1966-67

3,996

2,910

5.1

0.41

1967-68

3,000

2,256

6.7

0.55

1968-69

3,000

2,379

9.2

1.11

1969-70

8,342

6,149

7.4

0.68

1970-71

13,611

11,187

6.9

1.08

1971-72

14,847

12,920

7.3

1.14

1972-73

12,702

9,938

6.1

0.53

Year

Number
Permits Issued

1964-65

1,608

1965-66

Table 11.

Est. Number
Active Hunters

Distribution of harvest, by county, in the permit area.
Boulder

furgan

Adams

Total

Year

Larimer

Weld

1964-65

504

181

1965-66

665

144

29

838

1966-67

764

409

11

1,184

1967-68

944

265

37

1,246

1968-69

1,584

886

161

2,631

1969-70

2,431

1,112

383

146

100

4,172

1970-71

7,486

3,544

620

93

370

12,112

1971-72

7,723

4,484

1,804

464

296

14,771

1972-73

2,855

1,547

413

329

126

5,270

685

�Table 12.

1969-70
Number
Percent

1970-71
Number
Percent

1971-72
Number
Percent

1972-73
Number
Percent

0

4,263

69.3

6,588

58.9

7,006

54.2

6,857

69.0

1

836

13.6

1,620

14.5

2,209

17.1

1,391

14.0

2

452

7.4

1,073

9.6

1,428

11.1

666

6.7

3

238

3.9

564

5.0

768

5.9

298

3.0

4

153

2.5

528

4.7

633

4.9

616

6.2

Season Bag

Total

Estimated distribution of season bag for active hunters in the northcental Colorado permit area.

5

138

2.2

342

3.1

371

2.9

50

0.5

6

69

1.1

472

4.2

505

3.9

60

0.6

6,149

100.0

11,187

100.0

12,920

100.0

9,938

100.0

,
0-

,

00

�-69-

Table 13.
1972-73.

Hunting pressure and harvest, by county, in northcentral Colorado,

Larimer County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

Estimated Number
1971-72

Estimated Number
1972-73

6,760 (+ 285)
41,535 (+3,217)
7,723 (+ 693)

5,067 (± 265)
27,141 (+2,455)
2,855 (+ 403)

6.1

5.4

Percent Change
From 1971-72

0.56
0.105

-25.1
-34.7
-63.0
-11. 5
-50.9
-43.6

5,911 (+ 280)
31,914 (+3,126)
4,484 (+ 514)
5.4
0.76
0.141

4,332 (+ 259)
18,007 (+1,815)
1,547 (+ 290)
4.2
0.36
0.086

-26.7
-43.6
-65.5
-22.2
-52.6
-39.0

2,296 (+ 206)
14,961 (+1,962)
1,804 (+ 353)

1,757 (+ 187)
10,570 (+1,885)
413 (+ 141)

-23.5
-29.4
-77 .1
- 7.7
-69.6
-67.8

1.14
0.186

Weld Count
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

Boulder County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

6.S

o. 79

6.0

0.121

0.24
0.039

1,104 (+ 149)
3,774 (± 738)
464 (+ 166)
3.4
0.42
0.123

924 (+ 143)
3,157 (+ 703)
329 (+ 141)
3.4
0.36
0.104

Morgan County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hqpter Trip

-16.3
-16.4
-29.1
-14.3
-15.5

-------------------------------------------------------------------------------~-

�-70-

Table 13. Hunting pressure and harvest, by county, in northcentral Colorado,
1971-72 (continued).

Adams County

Estimated Number
1971-72

Estimated Number
1972-73

Total Individual Hunters
Total Hun ter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trips

545 (+ 107)
2,288 (+ 681)
296 (+ 152)
4.2
0.54
0.129

462 (+ 108)
2,191 (+ 780)
126 (± 96)
4.7
0.27
0.058

-15.2
- 4.2
-57.9
+11.9
-50.0
-55.0

12,920 (± 189)
94,472 (+4,027)
14,771
850)
7.3
1.14
0.156

9,938 (+ 223)
61,066 (+3,059)
5,270 (+ 516)
6.1
0.53
0.086

-23.1
-35.4
-64.3
-16.4
-53.5
-44.9

Percent Change
From 1971-72

Entire Permit Area
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

(±

Distribution and Harvest
The general distribution of Canada geese during the 1972-73 hunting season
was consistent with previous years. The majority of the geese were located
either along the Rio Grande River or on the MOnte Vista National Wildlife
Refuge (Table 15). The number of geese on the MOnte Vista Refuge increased
as the season progressed. The total number of geese counted in the Valley
during the 1972-73 season varied from 1,000 in October to 1,600 in January
(Table 16).
A total of 400 permits were issued authorizing the taking of one goose during
the 1972-73 special San Luis Valley goose season. According to the harvest
survey, an estimated 311 active hunters harvested 255 geese for an average
season bag of 0.82 (Tables 17 and 18).
During the 1972-73 hunting season, about 47 percent of harvest occurred on
or within two miles of the MOnte Vista National Wildlife Refuge, compared to
60 percent in that area during the 1971-72 season. Another 35 percent of
the harvest was reported taken along the Rio Grande River between Del Norte
and Alamosa.

�-71-

Table 14. Banding areas outside the San Luis Valley of Canada geese
recovered within the San Luis Valley during the 1970-71, 1971-72 and 1972-73
hunting seasons.

Number of Recoveries
1970-71
1971-72
1972-73

Area

Alberta
Dowling Lake

2

o

o

Montana
Phillips

Coun ty

1

o

'0

Garfield

County

o

2

o

1

5

5

o

1

o

2

o

o

1

o

o

So ccoro County

o

1

2

Mora County

o

o

1

o

1

o

Wyoming
Albany

County

Wheatland
Fremont

Reservoir

County

Ocean Lake
Colorado
Northcentral
Texas
Panhandle
New Mexico

Wisconsin
Columbia

Coun ty

�-7:'-

Table 15. Results of Canada goose surveys during the 1972-73 hunting season
in the San Luis Valley.
Area

October 25

November 22

December 18

January 8

565

481

725

1,156

Del Norte to Monte Vista

158

38

30

45

Monte Vista to Alamosa

5

222

81

30

Alamosa to State Line

126

240

226

o

Conejos River

4

o

15

40

San Luis Lakes

119

215

o

o

Smith Reservoir

20

o

2

o

Russell Lakes

9

o

o

o

Monte Vista Nat'l Wildlife
Refuge
Rio Grand River

Area 15 miles South, 2 miles
East of Saguache
Total

Table 16.
Colorado.

Year

350
1,006

1,196

1,079

1,621

Winter inventories of Canada geese in the San Luis Valley of

October

1970-71

Number of GeesJ:.!
November
December

January

1,490

1,050

1,261

1971-72

1,570

913

1,255

1,286

1972-73

1,006

1,196

1,079

1,621

1/ Compared monthly, inventories for the three years generally taken within
fiv~ days of each other.

�-73-

Table 17.

Hunter activity

and success

Year

Number
Permits Issued

Est. Number
Active Hunters

Average
Days Hunted

Average
Seasonal Bag/Hunter

1970-71

200

164

3.4

0.65

L971-72

350

296

4.2

0.60

L972-73

400

311

4.6

0.82

in permit area.

Table 18.

Distribution

of harvest,

by county in the permit

area.

Year

Rio Grande

Alamosa

Saguache

Conejos

Costilla

Total

1970-71

74

31

2

o

o

107

1971-72

121

50

1

5

o

177

1972-73

195

51

2

1

6

255

Only 27 percent of the harvest during the 1972-73 season was reported taken
during the first six weeks of the season.
During the 1970-71 and 1971-72
seasons, 53.5 percent and 59.4 percent of the birds taken were reported
harvested during the same six week time period in.the respective years.
Harvest
hunting

and hunter activity for the entire permit area for each of the three
seasons are presented in Table 19.

Westcentral
Hunter Activity

Colorado

and Harvest

Two hundred and fifty goose permits were issued, authorizing the taking of
one goose during the November 22 to December 17, 1972 hunting season, within
the westcentral goose permit area. An estimated 182 active hunters harvested
58 geese, for an average seasonal bag of 0.32 birds per hunter (Tables 20, 21).
Approximately 41 percent of the birds harvested were taken within five miles
of Highline Reservoir, which is located approximately 20 miles northwest of
Grand Junction.
The results of pre-and post-season aerial inventories in westcentral Colorado
for 1972-73 with comparative 1971-72 totals are presented in Table 22.

�Table 19. Hunting pressure and harvest
area, 1971-72 through 1972-73.

Estimated Number
1970-71

Entire Permit Area

Total

Individual

in the San Luis Valley goose permit

Hunters

Estimated Number
1971-72

164

(± 3)

296

(± 5)

Total Hunter Trips

556

(± 20)

1,236

(± 36)

Total Geese Bagged

107

(± 4)

177

(± 6)

Estimated Number
1972-73

311 (±-

6)

(± 54)

1,440

255 (±.

Average

Hunter Trips/Hunter

3.4

4.2

4.6

Average

Bag/Hunter

0.65

0.60

0.82

Average

Bag/Hun ter Trip

0.19

0.14

0.18

8)

"
,1

Table 20. Hunter activity
permit area.

and success

in the westcentral

Colorado

goose

Il
"

Ij

11

Ii

Number
Permits Issued

Year

!j
Est. Number
Active Hunters

Average
Days Hunted

Average
Seasonal Bag/Hunter

l1
IJ

!

1

1971-72

150

112

2.9

.29

1972-73

250

182

3.7

.32

in the westcentra1

Estimated

Colorado

1972

112

(± 4)

182

Trips

329

(± 20)

579 (±. 30)

Cee se Bagged

33

(± 4)

58

Total Hunter
Total

goose

Number

1971
Hunters

I

·1

Table 21. Hunting pressure and harvest
permit area, 1971 and 1972.

Total Individual

I

(± 5)

(± 5)

Average

Hunter Trips/Hunter

2.9

3.7

Average

Bag/Hunter

0.29

0.32

Average

Bag/Hunter

0.10

0.086

Trips

�-75-

Table 22.

Canada goose inventories,

Area

westcentral

November

Highline

Lake

Colorado

River

Colorado,

1972-73.

21, 1972

Date
January

11, 1973

215

225

107

139

7

45

o

5

B1 ack Canyon

o

204

Delta

o

95

o

470

329

1,183

485

989

Rifle to Debeque Canyon
Grand Junction

to Utah State Line

North Fork River
Hotchkiss
Gunnison

to Gunnison

River

River

to Grand Junction

In fields in Colorado

River Valley

below Grand Junction

Total
Comparative

1971-72

Prepared

&gt;&lt;~&gt;!f.&gt; :/}.;../1 iT' /..:i;',}.---1-Y,,&lt;c; /a//

by

Totals

Michael R. Szymczak 'j ::'
()
Assistant Wildlife Researcher

��October, 1973
-77-

JOB PROGRESS REPORT

State of

~CO~L~O~RAD~~O~

Migratory Bird Investigations

W-88-R-18

Project No.
Work Plan No.
Job Title
Period Covered:
Personnel:

_

2

Job No.
~~7
_
Non-hunting 'Mortality Investigations
~o~f~~C=a=n=a=d=a~q~e~e~s~e~i=n~S~o~u~th~e~a=s=t=e=r~n~C~o~l=o~r~a~d~o
---_

October 3, 1972 through March 31, 1973

W. Adrian, R. Baker, G. Claassen, D.Coven, G. Eyre, R. Keiss,
R. Kitzmiller, S. Porter, R. Rosette, L. Webster and M. Szymczak.

ABSTRACT
Lead shot was found in 26 of 215 soil samples extracted from the basin of
Turk's Pond in Baca County, southeast Colorado. On a percentage basis,
positive samples were more numerous along the south-southeast portion of the
basin where soil was granular and not subject to washing by irrigation water
and resulting siltation. About 48 percent of the lead shot found was size
number two. None of 59 blood samples extracted from normal appearing geese
captured in January and February, 1973 contained any measurable lead.

��-79-

NON-HUNTING MORTALITY INVESTIGATIONS
OF CANADA GEESE IN SOUTHEASTERN COLORADO
Michael R. Szymczak
In late January and early February of 1972, Canada geese of the Short Grass
Prairie population wintering at Turk's Pond were noted to be undergoing
an unexplained mortality. As many as 2,000 geese may have succumbed. Ten
live birds and 15 dead birds were collected at Turk's Pond. The live birds
were in visibly poor condition. Sample birds were sent to the Colorado
State University Diagnostic Laboratory and the Bear River Wildlife Disease
Research Station. No infectious disease organisms were isolated from birds
sent to Bear River. One organism was isolated in at least one of the birds
at CoLorado State University. However, that organism has never been
reported as associated with goose mortality. At least one worn lead shot
was found in five of the eight birds sent to Bear River. In addition, post
mortem findings were generally consistent with diagnoses of lead poisoning.
This study was directed toward confirming the possibility of mortality due
to lead poisoning with additional procedures incorporated to identify other
possible causative organisms or poisons. However, during the winter of
1972-73 no unusual mortality was discernable at Turk's. Therefore, the
majority of the procedures outlined in the project documents we~e not initiated. Only those procedures followed and the concluding results will be
dealt with here.
P. S. OBJEGrIVE
To identify non-hunting mortality factors of Canada geese at Turk's Pond in
southeast Colorado.
SEGMENT OBJECTIVES
1.

Determine the incidence of lead shot in the environment at Turk's Pond
and local vicinity.

2.

Determine if lead poisoning occurs in the waterfowl population at
Turk's Pond.

3.

Determine if other causative agents of disease or poisoning are
responsible for mortality of Canada geese at Turk's Pond.

4.

Determine the magnitude of non-hunting mortality in the Turk's Pond
area.

5.

Analyze data and formulate recommendations for preventative measures
designed to lessen non-hunting mortality of €anada geese at Turk's
Pond.

�-82-

Table 2. Lead shot size of pellets found in soil samples from the basin
area of Turk's Pond, southeast Colorado.

Sam~le Orisins
Below HiSh Water Line
Above Hi~h Water Line
No.
Percent
No.
Percent

No.

6

1

5.3

0

0.0

1

3.4

4

3

15.8

0

0.0

3

10.3

2

9

47.4

5

50.0

14

48.3

BB

4

21.1

3

30.0

7

24.1

No. 4
Buck

2

10.5

1

10.0

3

10.3

No. 0
Buck

0

0.0

1

10.0

1

3.4

Total

19

Shot
Size

Prepared by

10

m;/JE~

Michael R. Szymc~
Asst. Wildlife Researcher

29

Total
Percent

�October,

1973

-83-

JOB PROGRESS

State

of

Project
Work

REPORT

COLORADO
-----------------------No.

W-88-R-18

Plan No.

Migratory

3

Bird

Job No.

Investigations

6

Job Tit Le -'- __ ..;I:;...n_v_e_s.;....t.;....l.
....
·g•.•.
__t..;i_o_n--"o..;f_·
a
_M_a_1_l_a_r_d--.;M~a_n_a-,glo4;..em
__e_n_t-..::U_n_i_t_s
__o....,.f
__E..;..a
__s _te
__r_n_C_o_l_o_t_a_d_o_
Period

Covered:

Personnel:

April

1, 1972 through

March

31, 1973

Michael Szymczak, Clait Braun, Jack Corey, Robert Kitzmiller,
James Jackson, Robin Henry, Delwin Smith, Ed Kautz, Howard Funk
and Richard Hopper.

ABSTRACT
Post-season mallard trapping efforts resulted in the banding of 6,071 birds
in eight study areas during the winter of 1972-73.
Quotas were met or exceeded
in all but one area.
Wintering populations
of mallards apparently
reached a
peak in late November at about 250,000 birds.
This number was much lower than
the peak of over 500,000 mallards counted in late December of the previous
year.
Extreme winter weather conditions
in 1972-73, plus low production,
probably accounted for much of this decline.
Sex ratio counts conducted during
January and February, 1973 produced an overall average of 60.3 percent males
in the Colorado mallard wintering population.
A mallard age ratio of 0.4
immatures per adult in the Colorado harvest (1972-73) was the lowest since the
1961-62 hunting season.
Updating of band recovery data to include the 1964-65
through the 1970-71 hunting seasons showed the continued high survival rates
of male mallards.

��-85-

INVESTIGATION OF MALLARD MANAGEMENT UNITS OF EASTERN COLORADO
Richard M. Hopper

Post-season banding of wintering populations of mallards was continued in
eastern Colorado in 1972-73. Past banding data collected as part of this
job contributed much to the cooperative Central Flyway program which finally
resulted in the acceptance of the High Plains Mallard Management Unit in
1972 by the Bureau of Sport Fisheries and Wildlife. This management unit
concept, as justified by Funk et al. (1971), essentially split the Central
Flyway into two units for the management of mallard populations. However,
banding work is being continued in Colorado to investigate the feasibility
of managing mallard populations or flocks within the State on a IOOre refined
basis. Colorado probably has the best post-season mallard banding data in
the country, largely because birds have been aged since initiation of the
study 1963-64, and we want to continue banding until the cohort of birds
from this first year of banding no longer remains in the population. This
report mostly covers results of banding and survey data collected during
the segment, but it also includes some updating of the overall banding
analysis.

P. S. OBJECTIVE
To develop a harvest formula for eastern Colorado management units.
SEGMENT OBJECTIVES
1.

To analyze, for each of nine eastern Colorado study areas of Wintering
mallards, all data collected from 1963-64 through the 1971-72 hunting
season on the discreteness of populations and publish results in an
appropriate journal.

2.

To determine the feasibility of requesting special hunting regulations
for individual mallard management units within Colorado.

3.

To trap, band, age, and sex samples of mallards during winter in each
management unit of study, based on results of final analysis, of
sufficient size to allow adequate monitoring of each population.
METHODS AND MATERIALS

Cage traps of the Salt Plains type were used to capture most mallards,
although cannon-nets were also employed in the Arkansas Valley. Trapping
began immediately following the end of the duck hunting season on January 14,
1973 and continued through February 22, 1973. Banding was accomplished in

�-86-

nine units of eastern Colorado, including five in the South Platte Valley,
three in the Arkansas Valley, and one in the Bonny Reservoir area. All birds
were aged by use of the wing according to Carney (1964) and Hopper and Funk
(1970).
Periodic aerial censuses were again conducted during the fall and winter to
estimate movements and numbers of birds present in the State.
Sex ratio counts
were conducted from the ground as described in previous segment reports.
A partial update of the banding analysis was accomplished for bandings from
1963-64 through 1969-70, representing recoveries from the 1964-65 through the
1970-71 hunting seasons.
Programs were developed and data from tapes supplied
by the Bird Banding Laboratory were used to obtain print outs of desired
information.
As in past years, age and sex ratios of mallards harvested in Colorado were
obtained through the annual Wing Bee conducted by the Bureau of Sport
Fisheries and Wildlife.
Colorado again participated in gathering hunter
performance data to evaluate the 1972-73 experimental point system duck season.
Methods used in conducting hunter performance surveys are described in detail
by Mikula et ale (1972).

RESULTS AND DISCUSSION
Trapping

and Banding

Banding quotas of six to eight hundred mallards were set for each of eight
study areas. and these were met or exceeded in all but one area (Table 1).
Management Units 10, 11 and 12 (Fig. 1) are listed separately in Table 1, but
for this study are considered one unit.
Cold and snowy weather prevailed
during most of the banding period, which resulted in good trapping conditions
and the ultimate attainment of quotas in most areas.
Nearly 6,100 mallards were banded as a result of trapping efforts in the
eight study areas
(Table 1). Efforts to band about equal numbers of birds by
age and sex were mostly successful.
Adult males were again the easiest to
obtain.
Adult females, normally the most difficult to get of the four age
and sex classes, were banded in good numbers this year, with samples being
even better than those for immature males in many areas.
Immature females
replaced adult females as the group hardest to obtain, with only 988 birds of
the former category being banded during the segment.
Immature birds of both
sexes were especially difficult to obtain in the Fort Collins area (Unit 4).
Severe weather conditions during the winter may have pushed many immature
birds out of Colorado, thus making them unavailable for banding.
Winter Aerial

Surveys

JateS and results of the periodic aerial counts are shown by management unit
i~ Table 2. Mallard numbers appeared to reach a peak in late November when

�-87-

about 250,000 birds were
counted.
This figure was considerably lower than
peak numbers observed during the previous year when over one-half million
birds were counted in late December (Funk 1972). A drastic reduction in
mallard numbers from 1971-72 to 1972-73 occurred during every counting
period, as well as in all management units except the San Luis Valley (15).
This difference between years is no doubt partly explained by extreme
weather conditions during 1972-73 that forced many birds out of the state.
Mild weather, in addition to a good production year, contributed to the above
average mallard populations that existed in Colorado during the fall and
winter of 1971-72.

Winter

Sex Ratio Counts

Mallard sex ratio counts are listed by management unit and area in Table 3.
Nearly 15,000 birds were classified, producing an overall estimate of 60.3
percent males in the wintering sample.
This compares closely with past
years, and continues to indicate a preponderance of males in the wintering
population.
An extensive survey of mallard sex ratios was conducted by all 10 states in
the Central Flyway during the nationwide mid-winter count in early January
of 1973 (Funk 1973). This survey, representing a total sample size of over
61,000 mallards, yielded a weighted average of 61.9 percent males in the
Central Flyway wintering mallard population.
Individual states varied from
52.5 percent males in New Mexico to 78.5 percent males in North Dakota.
Sex ratio surveys conducted in Colorado in recent years, plus the extensive
survey done in the Central Flyway in 1973, adds further evidence to support
the contention that a significant imbalance in the sex ratio in favor of
males exists in wintering populations of mallards in the Central Flyway.
These data also suggest that in Colorado, and probably the remainder of the
Central Flyway states, hunting pressure has not been of sufficient magnitude
to greatly alter sex ratios under drake-oriented regulations experience the
past few years.

Wing Surveys
A mallard age ratio of 0.4 immatures per adult was estimated to be in the
Colorado harvest during the 1972-73 hunting season, based on the Bureau's
wing collection survey (Sorensen et al. 1973). This ratio was the lowest
encountered in Colorado since the 1961-62 hunting season when the Bureau's
Parts Collection Survey came into existence.
However, this was also true
of most other states throughout the country, which reflected the earlier
prediction of relatively poor production on the major breeding grounds and
the beginning of a new drouth period.

�c,.,
••••••
~

es

)

••••••

:it::
~
c:.:s
~

(,;

...•.•...
~"

e
:t:
••••
•••••
~

v
:"i

H

'tj

1

n

o

]

]

J

0

u
x
w

::F

ru,
t'j

~
~
0

!3:

..-1
H

4-1

Q)

t'j

.w

:3

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o

�Table 1.
1972-73.

Numbers and percentages of mallards in the banded sample by location, age and sex, eastern Colorado,

Adult
No.
Banded

No.

Male
Percent

(1) Sterling-Julesburg

749

247

(2) Ft. MOrgan-Sterling

994

(3) Greeley-Ft. MOrgan
(4) Ft. Collins

Managemen t Unit

Immature
Male
Female
Percent
Percent
No.

Female
No.
Percent

No.

33.0

204

27.2

202

27.0

96

12.8

244

24.5

250

25.2

295

29.7

205

20.6

889

242

27.2

278

31. 3

226

25.4

143

16.1

645

200

31.0

266

41.2

95

14.7

84

13.0
I

00

(6) Denver-Greeley

1,112

313

28.1

321

28.9

317

28.5

161

14.5

(9) Bonny Reservoir

618

198

32.0

148

23.9

156

25.2

116

18.8

(10) Lamar-Holly

141

53

37.6

13

9.2

46

32.6

29

20.6

(11) Two Buttes Reservoir

246

87

35.4

88

35.8

24

9.8

47

19.1

(12) Rocky Ford-Lamar

628

241

38.4

110

17.5

172

27.4

105

16.7

(13) Pueblo-Rocky

49

35

71.4

7

14.3

5

10.2

2

4.1

6,071

1,860

30.6

1,685

27.8

1,538

25.3

988

16.3

Totals

Ford

\,C)

I

�Table 2.

Aerial duck counts by date interval and study unit, eastern Colorado, 1972-73.

Managemen t Unit

November 6, 10

(1) Sterling-Julesburg

(4) Ft. Collins
(6) Denver-Greeley
(9) Bonny Reservoir
(10-13) Arkansas Valley
(15) San Luis Valley
Totals

18,560

8,625

2,800

-- }j

25,970

24,800

22,700

31,300

37,220

19,500

53,250

39,000

64,775

4,025

8,700

31,450

11,200

26,625

43,750

30,000

37,500

-- 1./

18,200

22,000

40,500

35,500

25,562 ])

29,044

27,170

35,538

1./

246,269

151,245 if

233,038

175,512

1f October 25, 1972.
Ft. Morgan-Julesburg

I

\D

0
I

1f Not censused.

1/

January 8-10, 12

-- }j

(2) Ft. Morgan-Sterling
(3) Greeley-Ft. Morgan

Number of Ducks Counted
November 21-22, 28
December 11-12, 15, 18

not included.

if Bonny Reservoir not included.

30,800

�-91-

Table 3.
1972-73.

Mallard post-season sex ratio counts by study unit, winter of

Management Unit and Date
Sterling-Julesburg

Male

No. Ducks Counted
Female
Total

Percent
Males

(Unit 1)

1-18-73 Jumbo Reservoir

269

155

424

63.4

2-3-73 South Platte River

304

186

490

62.0

386

214

600

64.3

1-10-73 Boxe1der Reservoir

280

160

440

63.6

2-6-73 Timnath Outlet Ditch

329

205

534

61.6

2-6-73 New Windsor Reservoir

1,477

1,210

2,687

55.0

2-6-73 Woods Lake

80

51

131

61.1

2-7-73 Windsor Area

238

129

367

64.8

1-9-73 Valmont Reservoir

616

405

1,021

60.3

1-11-73 Valmont Reservoir

1,840

1,170

3,010

61.1

1-11-73 Bonny Reservoir

650

350

1,000

65.0

1-15-73 Bonny Reservoir

315

212

527

59.8

1-17-73 Bonny Reservoir

839

672

1,511

55.5

1-18-73 Bonny Reservoir

178

125

303

58.7

1-19-73 Bonny Reservoir

297

226

523

56.8

1-15-73 Two Buttes Reservoir

488

240

728

67.0

1-18-73 Two Buttes Reservoir

351

176

527

66.6

8,937

5,886

14,823

60.3

Greeley-Ft. Morgan (Thlit 3)
1-20-73 South Platte River
Ft. Collins Area (Unit 4)

Denver-Greeley

(Unit 6)

Bonny Reservoir Area (Unit 9)

Arkansas Valley (Unit 11)

Totals

�-94-

Table
4. Relative recovery rate method estimates on survival and mortality
rates of mallards winter-banded in Units 1, 2 and 9 in eastern Colorado from
1963-64 through 1969-70 by age and sex at time of banding. (Data represent
recoveries from the 1964-65 through 1970-71 seasons) (continued).

Unit of Banding
and Age and Sex

No.
B'anded

First
Year

Recovery Rates
Second and
Later Years
Total

Survival
Rate

M:&gt;rtality
Rate

Units 1, 2 and 9 Combined
Adult Male

5,660

.024

.067

.091

.856

.144

Immature Male

4,312

.036

.062

.098

.777

.223

Total

9,972

.029

.065

.094

.819

.181

Adu1 t Female

2,531

.017

.019

.036

.647

.353

Innnature Female

3,434

.019

.024

.043

.603

.397

5,965

.018

.022

.040

.627

.373

Total

�-95-

Table
5. Relative recovery rate method estimates on survival and mortality
rates of mallards wint~r banded in Units 3, 4 and 6 in eastern Colorado from
1963-64 through 1969-70 by age and sex at time of banding. (Data represent
recoveries from the 1964-65 through 1970-71 seasons).

Banded

First
Year

Recovery Rates
Second and
Later Years

Total

Survival
Rate

Mortality
Rate

Adult Male

1,611

.039

.069

.108

.677

.323

Immature Male

1,592

.031·

.067

.098

.781

.219

Total

3,203

.035

.068

.103

.724

.276

Adul t Female

885

.019

.030

.049

.853

.147

1,346

.023

.033

.056

.740

.260

2,231

.022

.032

.054

.811

.189

Adult Male

2,103

.026

.074

.100

.828

.172

Immature Male

1,729

.035

.073

.108

.824

.176

Total

3,832

.030

.074

.104

.822

.178

Adult Female

1,158

.024

.022

.046

.634

.366

Immature Female

1,367

.020

.021

.041

.585

.415

2,525

.022

.021

.043

.529

.471

Adult Male

2,327

.034

.092

.126

.745

.255

Immature Male

1,454

.046

.080

.126

.611

.389

Total

3,781

.038

.087

.125

.674

.326

Adult Female

931

.015

.017

.032

.623

.377

1,066

.024

.030

.054

.570

.430

1,997

.020

.024

.044

.597

.403

Unit of Banding
and Age and Sex

No.

Unit 3

Immature Female
Total
Unit 4

Total
Unit 6

Immature Female
Total

------------------------------------------------------------------------------

�-96-

Table
5. Relative recovery rate method estimates on survival and mortality
rates of mallards winter banded in Units 3, 4 and 6 in eastern COlorado from
1963-64 through 1969-70 by age and sex at time of banding.
(Data represent
recoveries from the 1964-65 through 1970-71 seasons) (continued).

Unit of Banding
and Age and Sex

No.
Banded

First
Year

Recovery Rates
Second and
Later Years

Total

Survival
Rate

Mortality
Rate

Units 32 4 and 6 Combine,d
Adult Male

6,041

.032

.079

.111

.777

.223

Immature

4,775

.037

.073

.110

.798

.202

Total

10,816

.034

.077

.111

.787

.213

Adult Female

2,974

.020

.023

.043

.609

.391

Immature

3,779

.022

.028

.050

.663

.337

6,753

.021

.026

.047

.644

.356

Total

Male

Female

�-97-

Table
6. Relative recovery rate method estimates on survival and mortality
rates of mallards winter banded in Units 10, 11, 12 and 13 in eastern Colorado from 1963-64 through 1969-70 by age and sex at time of banding. (Data
represent recoveries from the 1964-65 through 1970-71 season).

Unit of Banding
and Age and Sex

No.
Banded

First
Year

Recovery Rates
Second and
Later Years

Total

Survival
Rate

Mortality
Rate

Units 10, 11 and 12 Combined
Adult Male

1,429

.021

.061

.082

1.102

-.102

Immature Male

1,162

.025

.038

.063

.742

.276

Total

2,591

.023

.050

.073

.817

.183

Adult Female

796

.011

.025

.036

.602

.398

1,030

.018

.012

.030

.428

.572

1,826

.015

.018

.033

.510

.490

Adult Male

603

.030

.045

.075

.860

.140

Immature Male

327

.034

.058

.092

1.007

-.007

Total

930

.031

.052

.083

.946

.054

Adul t Female

311

.013

.010

.023

.684

.316

Immature Female

253

.024

.024

.048

.411

.589

564

.018

.016

.034

.774

.226

Immature Female
Total
Unit 13

Total

Units 10,11, 12 and 13 Combined
Adult Male

2,032

.024

.056

.080

.996

.004

Immature Male

1,489

.032

.043

.075

.652

.348

Total

3,521

.027

.050

.077

.830

.170

Adult Female

1,107

.012

.021

.033

.867

.133

Immature Female

1,283

.019

.014

.033

.446

.554

2,390

.016

.017

.033

.742

.258

Total

�-98-

LITERATURE CITED

Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification
by means of wing plumage. U. S. Fish and Wildl. Serv., Spec. Sci.
Rept. :Wildl. No. 82, 47 p.
Funk, H. D. 1969. Investigation of mallard management units of eastern
Colorado. Colo. Div. of Wildl., Game Res. Rept., Oct. p. 131-156.
_____ , J. R. Grieb, D. Witt, G. F. Wrakestraw, G. W. Merrill, J. Sands,
T. Kuck, D. Timm, T. Logan, and C. D. Stutzenbaker.
1971. Justification of the Central Flyway High Plains Mallard Management Unit.
Unpubl. Central Flyway Tech. COlDIn.Rept. (Mimeo). 48 p ,
1972. Investigation of mallard management units of eastern Colorado.
Colo. Div. of Wildl., Game Res. Rept. Oct. p. 105-114.
1973. Central Flyway mallard sex ratio survey midwinter count.
Central Flyway Tech. Comm. Rept., (Mimeo). 4 p.
Hopper, R. M., and H. D. Funk. 1970. Reliability of the mallard wing age.determination technique for field use. J. Wildl.Mgmt.
34(2) :333-339.
Kimball, C. F. 1973. Results from the hunter performance survey, 1970-1972,
and the bag check temperature survey, 1972. U. S. Fish and Wildl. Serv.,
Div. of Wildl. Res. Unpublished Rept. (Mimeo). 25 p.
Mikula, E. J., G. F. Martz, and C. L. Bennett, Jr. 1972. Field evaluation
of three types of waterfowl hunting regulations. J. Wildl. Mgmt.
36(2):441-459.
Sorenson, M. F., S. M. Carney, and L. D. Schroeder. 1973. Species, age, and
sex composition
of ducks bagged in the 1972 hunting season in comparison
with prior years. U. S. Fish and Wildl. Servo Office of Migratory Bird
Mgmt. Admin. Rept. 50 p ,

Prepared by ----l/£=:.J.~4;"'Jz::....:.L-:;..!:"~~~=4Z_
.. ---::7~-+~J=----4.?h....L.J.~Mi.;;l&lt;11..'-:L0!:::::- ~_
Richard M. Hopper ~~
Wildlife Researcher

�October,

1973

-99-

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~---------

Project No.
Work plan No.

4

Job Title
April

Bird Investigations
3

Job No.
Trapping

Period Covered:
Personnel:

Migratory

W-88-R-18

and Banding Doves

1, 1972 through March

31, 1973

Jack Randall, Bureau of Sport Fisheries and Wildlife; Clait E.
Braun, Howard D. Funk, J. Edward Kautz, Brett Petersen, Norwin
Smith, Steve Steinert, Mark Stromberg and J. Allen White,
Colorado Division of Wildlife.

ABSTRACT
Efforts initiated in 1964 to trap and band samples of mourning doves (Zenaida
macroura)in Colorado were continued in 1972. Cooperative federal and state
efforts resulted in 4,SSS birds being newly banded.
Of this total, 2,3SS were
immatures, 1,238 were adult males, 961 were adult females and one was an unsexed
adult.
Assigned quotas were accomplished for all areas except extreme eastern
Colorado.
Wing molt data from trapped and harvested immatures suggested four
hatching peaks in 1972. Peak hatching periods were mid- to late May, mid-June,
mid-July and early August.
A sample of 1,200 hunter-harvested
birds, mostly
from eastern Colorado, gave a young-to-old ratio of 1.S:1. Wing molt data
from harvested immatures in this sample indicated that most early hatched young
had migrated from Colorado prior to September 1. Fifty-five band recoveries
were reported in 1972 from doves banded by Division personnel.
Only 23
(41.8 percent) had been banded in 1972. Twenty-three (41.8 percent) were
recovered in Colorado, while 14 (2S.S percent) were from Mexico with the remainder being from scattered locations mostly south and west of Colorado.
Apparent hunting mortality for doves banded in Colorado continues to be less
than 3 percent.

��-101-

MOURNING DOVE TRAPPING AND BANDING
C1ait E. Braun
Intensive efforts to trap and band mourning doves in Colorado initiated in
1964 were continued in 1972 in cooperation with personnel from the U. S.
Fish and Wildlife Service. Data presented in this report are those collected in 1972, the ninth year of this continuing investigation.
P. S. OBJECTIVE
To investigate migration patterns and mortality rates of mourning doves
banded in Colorado by age, sex and area.
SEGMENT OBJECTIVES
1.

To trap and band mourning doves in three selected areas of Colorado for
the purpose of obtaining migration, life history, and annual mortality
data.

2.

To estimate harvest size and hunter success.
METHODS AND MATERIALS

Methods and materials used in 1972 were similar to those described in earlier
reports and summarized by Braun (1970). In addition, less than 100 doves
were trapped in cannon nets in 1972 during trapping operations for bandtailed pigeons (Columba fasciata).
Description of Trapping Sites
Most trapping sites in Eastern Slope areas were similar to those described
earlier (Braun 1970). One new site was utilized in 1972 east of the Rocky
Mountains, in addition to those also trapped in 1971. This site was near
Springfield where trapping was conducted near p1antings of introduced trees
adjacent to grainfields (primarily wheat, Triticum aestivum). West slope
trap sites were essentially those also used in 1971, with the exception of
one site. This site near MOntrose was a livestock feeding area surrounded
mostly by small residential home sites with an abundance of cottonwood trees
(Populus spp.).
RESULTS AND DISCUSSION
Trapping and Banding
Cooperative Federal and State trapping and banding efforts resulted in 4,555
doves being newly banded in 1972. Of this total, 4,100 were banded by Division

�-102-

personnel. Doves were banded at four major sites on the eastern slope, two
sites on the eastern plains and at two major sites west of the Continental
Divide. Additionally, some doves were banded at five other scattered
locations in conjunction with band-tailed pigeon trapping activities. The
banding goal of 4,000 birds was essentially accomplished as over 1,000 were
banded in western Colorado, slightly less than 1,000 in extreme eastern Colorado and over 2,000 were banded along the eastern slope of the Rockies. Sex
and age distribution of birds banded by area are presented in Table 1.
Slightly more immature doves than adults were banded in 1972, a change from
the situation prevailing in most previous years of the study. Production in
1972 was believed to be good to excellent, possibly explaining the higher
percentage of immatures trapped and banded. However, differences in numbers
of adults and immatures trapped is undoubtedly a function of time of season
of trapping effort. Trapping at Vineland, Ft. Collins and Durango was
accomplished primarily in June, while trapping at Ft. MOrgan, MOnte Vista
and Montrose was conducted principally in July and early August. As in
previous years of the study, a higher percentage of males were trapped than
females (27.2 to 21.1 percent). This disparity is probably related to timing
of incubation and brood sharing, as females would normally be exposed to
only one trapping period per day.

Hatching Data
Wing molt data were available for 2,323 of the 2,355 immature doves trapped
and banded in 1972 and from 653 hunter harvested immatures. Estimated
hatching dates (Allen 1963) are presented in Tables 2 and 3. Examination of
data in Table 2 indicates that few mourning doves trapped in Colorado hatched
prior to May 1, with most hatching in mid- to late May and mid-June. While
definite hatching peaks are difficult to ascertain from data in Table 2,
peaks are suggested in mid-May and mid-June. Since trapping ceases in early
to mid-August, progeny of late nests are not represented in Table 2. This
can clearly be seen upon examining Table 3, as 3 peaks of hatching can be
ascertained (late June, mid-July, and late July to early August). These data
from hunter-harvested immature doves are biased toward late hatching juveniles
while trapping data are biased toward early hatching doves. These data also
support the hypo"thesis that progeny of early nests in Colorado are not available during the hunting season in Colorado. It should be remembered that
neither trap or hunter samples may actually represent doves produced in
Colorado. This is especially true for hunter-harvested birds.

Harvest
Efforts were made to collect wings from hunter-harvested doves throughout
Colorado during the 1972 hunting season. Through this effort, 1,200 wings
were examined for age (adult or immature) and primary molt. All but 15 of
these wings were from east slope areas, with the largest samples from Ft.
Carson (756), Pritchett (196), and LaVeta (57). All wings were from September with over 80 percent being from the first two weekends of the month.

�-103-

Table 1. Sex and age distribution of mourning doves banded by area in Colorado, 1972.
Adult Male
No.
Banded Percent

Area

Adul t Female
No.
Banded Percent

Immature
No.
Banded Percent

Total

Eastern Colorado
Ft. Morgan

34

6.8

58

11.6

408

81.6

500

Vineland

128

30.4

164

39.0

129

30.6

421

1

4.4

5

21.7

17

73.9

23

163

17.3

227

24.0

554

58.7

944

Ft. Garland

257

29.9

145

16.8

459

53.3

Monte Vista

101

17.9

84

14.9

380

67.2

565

Denve.;!:/

200

39.9

85

17.0

216

43.1

501

Ft. Collins

150

30.0

169

33.8

181

36.2

500

Ft. Carson

22

64.7

6

17.6

6

17.7

34

LaVeta

3

100.0

o

0.0

o

0.0

3

Subtotal

733

29.8

489

19.8

1,242

50.4

2,464

51

9.4

56

10.4

433

80.2

540

Durango-

247

47.9

159

30.8

110

21.3

516

Unaweep

26

55.3

11

23.4

10

21.3

47

Meeker-Craig

18

41.9

19

44.2

6

13.9

43

Subtotal

342

29.8

245

21.4

559

48.8

1,146

Total All Areas

1,238

27.2

961

21.1

2,355

51.7

Springfield
Subtotal
Eastern Slope

Western Slope
3/
Montrose4/

!/ Total does not include one AHY unknown sex.
~/ Including one bird banded at Evergreen.

1/ Including 9 birds banded at Carbondale.
i/ Including 3 birds banded at Arboles.

�Table 2.

Estimated hatching dates for wild-trapped
Prior to
May 1

Number
Hatching

3

Percent of Total

0.1

immature mourning doves, 1972.

Mal

1-7

44
1.9

8-14

15-21

125

22-28

283

5.4

29-June 4

340

12.2

344

14.6

5-11

June
12-18

19-25

26-Ju1y 2

3-9

Ju1l
10-16

17-23

24-30

326

196

139

135

78

22

0

14.0

8.4

6.0

5.8

3.4

1.0

0

288

14.8

12.4

....
I

Table 3. .Estimated hatching dat.es for hunter-harvested
Prior

Number
Hatching
Percent of
Total

Prior to
May 21

0
0

Mal

22-28

5
0.8

29-June 4

2
0.3

5-11

2
0.3

June
12-18

4
0.6

o
.p-

innnature mourning doves, .1972.

19-25

102

15.6

26-July 2

15
2.3

3-9

48
7.3

Ju1l
10-16 17-23

I

24-30

31-August 6

August
7-13 14-20

21"'2:

109

79

46

111

90

20

20

16.7

12.1

7.0

17.0

13.8

3.1

3.1

�-105-

Immature to adult ratios varied from a high of 4:1 at Monte Vista (N = 35),
2.6:1 at Pritchett (N = 196) to 1.2:1 at Fort Carson (N = 756). For all
wings sampled, the immature to adult ratio was 1.5:1 (654 immatures to 446
adults). Comparable data for an extensive area and large sample size were
not available for previous years. Wing molt data were available for 653 of
the 654 immatures. Calculated hatching dates are presented in Table 3.
Harvest data obtained through use of a mail survey of hunters after the
hunting season indicated that 33,299 hunters harvested 513,898 doves in Colorado in 1972 (H. Riffel, personal communication). These figures are significantly (P&lt;.Ol)
higher than in 1971 when an estimated 22,033 hunters harvested
298,767 doves. Reasons for the large disparity are not know, but must relate
to sampling problems. Reported band recovery rates did not change between 1971
and 1972.
A total of 55 band recoveries were received for doves recovered in 1972 that
had been banded by Division personnel. One of these birds had been initially
banded in 1968, 4 in 1969, 3 in 1970, 24 in 1971, while 23 had been banded in
1972. All but 7 of these were shot recoveries. Of the 55 recoveries, only 9
were from West Slope bandings, with 2 each recovered in Arizona, California.
Mexico and western Colorado, and one from near Las Cruces, New Mexico. Based
on this small sample, it would appear that the majority of doves banded west
of the Continental Divide migrate southwest from Colorado. In contrast, doves
banded in the San Luis Valley (Monte Vista and Ft. Garland) move primarily
south (one recovery each from New Mexico and Texas and 5 from Mexico). However, some doves banded in this area tend to disperse widely as ~ recovery each
was reported from California, Oregon and Illinois. Considering all recoveries,
23 (41.8 percent) were from Colorado, 14 (25.5 percent) were from Mexico,
while 5 were reogvered in Texas, 3 each in California and New Mexico, 2 in
Arizona, and 1 each from E1 Salvador, Nicaragua, Oregon, Illinois and South
Dakota. Apparent hunting mortality continues to be less than 3 percent of all
doves banded in a given year or of a particular age class banded in the same
year.
Recoveries of doves banded outside of Colorado were reported to the Division
of Wildlife for the initial time in 1972. Four doves (all banded as immatures)
banded elsewhere were reported killed in Colorado in 1972. Two each were from
Montana and Nebraska.
LITERATURE CITED
Allen, J. M. 1963. Primary feather molt rate of wild immature doves in
Indiana. Ind. Dept. Conserv., Game Res. Sect. Circ. No.4.
Indianapolis. 4 p.
Braun, C. E. 1970. Mourning dove trapping and banding. Colo. Div. Game,
Fish and Parks, Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. p , 143-149.

Prepared by__

--==L.=.~~

-===.-=Clait E. Braun
Wildlife Researcher

",::~~~~
•....
•

_

��October,

-107-

JOB PROGRESS

REPORT

S ta te of

.....:::.CO::::.;L:::,.O::::;RA:.:;::;D:::.,:O::_

Project No.

W-88-R-18

Work Plan No.

4

Job Title
Period Covered:
Personnel:

..
~ Migratory

April

Bird Investigations

4

Job No.
Band-tailed

1973

Pigeon

Investigations

1, 1972 through March 31, 1973

Spencer Amend and Ken Baer, Bureau of Sport Fisheries and Wildlife;
Jim Cruse, Wayne Paintner and other personnel of the U. S. Forest
Service; John Arther, Lyle Bennett, Clait Braun, Harold Burdick,
Bob Clark, Jack Corey, Howard Funk, J. Edward Kautz, Marie Vendeville
Kautz, Ann Leckler, Richard McDonald, Kris Moser, Brett Petersen,
Charles Roberts, Carl Roy, John Seidel, Norwin Smith, Mark Stromberg,
Charles Taylor, J. Allen White, Wildlife Conservation Officers and
other personnel of the Colorado Division of Wildlife.
ABSTRACT

Investigations of band-tailed pigeons (Columba fasciata) initiated in Colorado
in 1969 in conjunction with the Four Corners States Cooperative Study were
continued in 1972. Major emphasis was placed on ascertaining relative abundance,
trapping and banding samples of pigeons throughout the State, and conducting an
experimental hunting season.
Analysis of all distribution and abundance data
indicated no major shifts in distribution or changes in abundance from 1946 to
1972. Distribution of pigeons was found to be closely correlated with Ponderosa
pine (Pinus ponderosa) and oak (Quercus spp.).
Pigeons were trapped at 23
different locations in 1972 with 7,251 birds (5,981 adults, 285 subadults and
985 immatures) being newly banded.
Over sixteen hundred (1,647) pigeons were
retrapped, including 21 banded outside of Colorado (12 from New Mexico and 9 from
Utah).
During the experimental hunting season, 562 permits were issued with
298 individuals actually hunting.
One hundred and twenty-four hunters were
successful, each harvesting an average of 5.9 pigeons.
Total estimated harvest
(including crippling loss) projected from the questionnaire
survey (92.5 percent
response) was 822 birds, less than one-half of the number harvested in 1971.
Crippling loss approximated 11 percent of the birds shot and retrieved.
Wings
were received from 93 successful hunters.
Immature and subadult pigeons comprised 35.2 percent of the 515 band-tail wings received.
Field checks of 266
(169 adults, 97 immatures) hunter-killed birds indicated that 50.9 percent of the
adults were still involved with nest activities and/or feeding of young.
Recoveries were received from 125 pigeons banded in Colorado.
The majority of these
(85) were recovered in Colorado, with 20 being recovered in New Mexico, 12 in
Mexico, 3 each in Arizona and California, and 1 each in Utah and Washington.
At least 13.6 percent of the birds harvested during the experimental season in
Colorado were banded.
Hatching dates were calculated for 1,169 immature pigeons,
with most hatching between 2 May and 29 August.
Helminth infections continue to
be low (9.5 percent) with females being more frequently parasitized than males.

��-109-

BAND-TAILED PIGEON INVESTIGATIONS
Clait E. Braun

Intensive ecological studies of wild band-tailed pigeons which nest and
reside in Colorado from late April to early November, initiated in 1969,
continued in 1972. Investigations in Colorado represent a portion of the
regionwide Four Corners Cooperative Band-tailed Pigeon Investigation which
was initiated in Arizona in 1967. Data presented in this report are those
collected in 1972, the fourth year of this continuing investigation.
P. S. OBJECTIVE
To investigate life history and ecological facts pertinent to band-tailed
pigeons in Colorado as follows: range; relative numbers; relation between
numbers of birds present by area and such physical factors as weather, food,
vegetative cover, elevation, and salt; breeding and nesting chronology;
migration patterns and chronology; harvest patterns; and mortality rates.

SEGMENT OBJECTIVES
1.

To investigate distribution and relative numbers of band-tailed pigeons in
Colorado by area.

2.

To investigate the correlation between abundance and distribution of
bandtails and physical features of the environment.

3.

To trap, band, age, and sex as many birds as possible in selected areas
of Colorado to investigate migration patterns and chronology of Colorado
bandtails throughout their range.

4.

To conduct an experimental hunting season, gather harvest data, and
evaluate results.
METHODS AND MATERIALS

Most methods and materials used in 1972 were identical to those used in 1969,
1970, and 1971 (Braun 1970; 1971; 1972a). Reporting procedures for field
personnel were standardized in all regions with all monthly reports being sent
to the regional biologists for tabulating and forwarding to Fort Collins.
Trapping methodology in 1972 was that used in 1970 (Braun 1971). Evaluation
of the experimental hunting season and disposition of collected birds were
the same as in 1970 and described by Braun (1971).

�-110-

Description of Trapping Areas
In 1972, pigeons were trapped at 16 locations also utilized in 1969, 1970
and 1971. These locations were near Durango, LaVeta, Monte Vista, Sanborn
Park, Evergreen, Ft. Garland, Unaweep, Beulah, Woodland Park, Longmont,
Estes Park, Stoner, Aspen, Bayfield, Avon, and Manitou Springs, and have been
previously described (Braun 1970; 1971; 1972a). In addition to these sites,
pigeons were trapped near MOntrose, Carbondale, Wetmore, Arboles, Glenhaven,
Pagosa Springs, and Woodmen Valley. These sites were typically grain fields
(Carbondale, Wetmore and Pagosa Springs), livestock feeding sites (Montrose
and Woodmen Valley), grain storage areas (Arboles), or residential areas
where reSidents fed a variety of birds (Glenhaven). As in previous years, all
trap sites were immediately adjacent to readily accessible trees or power
lines used for perches. Trees used were primarily ponderosa pine, Gambel's
oak (Quercus gambelii), Pinyon pine (Pinus edulis), cottonwoods (Populus spp.),
and Engelmann spruce (Picea engelmannii). It was difficult to ascertain
feeding preferences as about equal numbers of pigeons were trapped in fields
where they were feeding on waste barley (Hordeum vulgare) and wheat (Triticum
aestivum). Whole or cracked corn (Zea mays) was frequently used as an attractant and bandtails clearly indicated a preference for this grain.

RESULTS AND DISCUSSION
Distribution and Abundance
Data collection for this objective terminated at the end of the field season
in 1972. All data collected have been analyzed and presented in a technical
paper (Braun 1973). The summary of that paper is as follows.
Band-tailed pigeons are widely distributed throughout the forested mountains
of Colorado, occurring primarily between 5,000 and 9,000 ft in elevation.
They prefer habitats dominated by Ponderosa pine and Gambel's oak, especially
where these vegetative types are interspersed with cultivated fields of small
grains. Nesting habitats utilized vary, with most observations outside of
feeding areas being from the higher limits of Ponderosa pine forests through
the spruce-fir-aspen forests into areas dominated by lodgepole pine and limber
pine. Abundance of pigeons appears related to vegetative type and possibly
parent materials. Bandtails in Colorado are most abundant in areas where
Gambel's oak is a co-dominant and where soils have derived from volcanic and
sedimentary rocks. Mineral springs and salt, while occaSionally used, are
not of major importance for bandtails in Colorado. It is concluded that no
significant changes in distribution and abundance of band-tailed pigeons in
Colorado have occurred since 1946.

Banding
Trapping and banding activities increased over all previous years of the
study. Trapping was conducted at 23 sites including 3 of the 5 1969 trap
sites, 8 of the 13 sites used in 1970, and 13 of the 15 sites trapped in 1971.

�-111-

In all, 7,251 pigeons were newly banded (Table 1). Of the 7,251 birds, 985
(13.6 percent) were immatures and it is apparent from examination of Table 1
that percent of immatures increased throughout the summer. This was especially apparent at LaVeta, Evergreen, Longmont, and Estes Park where trapping
was conducted throughout the summer. The high percentage of immatures trapped
at Woodmen and Evergreen in September may be misleading, as adults that were
previously banded.are not included in the calculations. However, results of
trapping in September were supported by observational data which suggested
that immature pigeons remained at trap sites after adults had dispersed to
new feeding sites or migrated south.
Sex of most adult and subadult pigeons banded in 1972 was determined through
examination of external plumage characters. Of the 5,981 adults, 2,985 were
males (49.9 percent), and 2,996 were females (50.1 percent), while only 84
of the 285 subadults were males (29.5 percent) with the remainder (201) being
females (69.5 percent). These data are similar to those collected in previous
years. The disparity in the sex ratio of subadults is not understood but may
be related to sex differences in rate of plumage molt. It is assumed that sex
ratios are equal at time of fertilization.
Time of day was recorded for all but 24 pigeons captured in 1972. A summary
of these data is presented in Table 2. It is obvious that sex ratios of
birds available for trapping changed during daylight hours. These changes
are undoubtedly due to differences between males and females in timing of
incubation, brooding and feeding. It is also apparent that more than half
of the trapping effort and success occurred after 1000 MDT. This is probably
the major reason why more females than males were trapped. With the exception
of the data for immatures, percentages of birds caught by time period for
adults are remarkably similar to those collected in prior years of the study.
Reasons why juveniles frequent trap sites in higher numbers in early morning
are not known. If time of day that attempts to capture pigeons were made
is examined (Table 2) it is readily apparent that both age and sex ratios
varied independently of time alone.
One thousand, six hundred and forty seven previously banded pigeons were
retrapped in 1972. All but 21 of these had been banded in Colorado, with 277
of this total being recaptured out of the original degree block of banding.
In all, 759 birds were banded prior to 1972. Of this total, 56 were from
1969, 241 were from 1970, and 462 were banded in 1971. Of the 21 out-of-state
bandings retrapped in Colorado, 12 were originally banded in New Mexico, while
9 were originally banded in Utah. In addition, it was learned that 17 pigeons
first banded in Colorado were retrapped in 1972 in New Mexico, while 5 were
retrapped in Utah.

Experimental Hunting Season
1972 was the third year of the 3-year experimental harvest program initiated
in 1970. Hunting was allowed from September 9 through October 1, with a
daily bag limit of 5 birds and a possession limit of 10. During the experimental season all of the state west of Interstate Highway No. 25 was open to

�Table 1.

Area

Durango

Bayfield
Carbondale
LaVeta

Number of band-tailed pigeons banded by area in Colorado, 1972.
Adults 1,/
No.
Banded
Percent

Immatures
No.
Banded
Percent

Totals

May 8-30
June 1-21
July 1-2
August 16-25

361
481
17
34

100.0
97.0
94.4
89.5

0
15
1
4

0.0
3.0
5.6
10.5

361
496
18
38

600+
1,000+
50+
50+

May 9

21

100.0

0

0.0

21

50+

100.0

0

0.0

340

500+
500+
50+
1,000+

Dates

May 11-24

340

Estimated No. of
Birds Using
Trapping Sites

May 11-17
July 18
August 6-24

256
28
409

100.0
90.3
71.3

0
3
165

0.0
9.7
28.7

256
31
574

May 18-24
July 19

223
20

100.0
95.2

0
1

0.0
4.8

223
21

400+
50+

May 20
June 15
July 6-27
August 9
Sept. 5-21

8
16
128
63
57

100.0
100.0
66.3
55.8
38.5

0
0
65
50
91

0.0
0.0
33.7
44.2
61.5

8
16
193
113
148

&lt;50
&lt; 50
200+
200+
200+

Monte Vista May 26
June 26-28
July 5-26
August 5-7

41
101
201
107

100.0
96.2
73.6
79.3

0
4
72
28

0.0
3.8
26.4
20.7

41
105
273
135

100+
200+
400+
300+

168
19
36

100.0
100.0
81.8

0
0
8

0.0
0.0
18.2

168
19
44

300+
50+
100+

Beulah

Evergreen

Woodland
Park

--

May 27
June 2
August 22

I
•....
•....

N
I

-----------------------------------------------------------------------------------------------------------

�Table 1.

Number of band-tailed pigeons banded by area in Colorado, 1972 (continued).
Adults 1/
No.
Banded
Percent

Immatures
No.
Banded
Percent

Totals

Estimated No. of
Birds Using
Trapping Sites

0.0
3.3

47
60

100+
100+

0
2
32

0.0
2.7
54.2

76
74
59

100+
200+
100+

97.5

13

2.5

524

1,000+

Area

Dates

Stoner

May 31
June 23

47
58

100.0
96.7

0
2

Longmont

June 4-21
July 5
August 8

76
72
27

100.0
97.3
45.8

Sanborn Pk. June 12-25

511

Estes Park

June 13
July 7
August 14 &amp; 15

5
32
47

100.0
97.0
52.2

0
1
43

0.0
3.0
47.8

5
33
90

20+
50+
150+

Unaweep

June 14 &amp; 27

267

95.0

14

5.0

281

400+

Glenhaven

June 22 &amp; 24
July 9

124
2

100.0
100.0

0
0

0.0
0.0

124
2

200+
10+

Montrose

June 26
July 11-19

493
369

97.4
85.8

13
61

2.6
14.2

506
430

1,500+
1,000+

Ft. Garland July 7-13

305

95.3

15

4.7

320

500+

Arboles

July 20
August 5

149
67

87.6
74.3

21
25

12.4
25.7

170
92

300+
150+

Avon

August 3

20

95.2

1

4.8

21

50+

-----------------------------------------------------------------------

I

I-'
I-'
w
I

�Table 1.

Area

Aspen
Wetmore
Pagosa
.§prings
Manitou
S~rings
Woodmen

Number of band-tailed

Dates

pigeons banded by area in Colorado, 1972 (continued).

Adults 1/
No.
Banded
Percent

August 9

36

August 16-25

September 3
September 9

Totals

Estimated No. of
Birds Using
Trapping Sites

76.6

11

23.4

47

100+

72.7

86

27.3

315

500+

95

77 .9

27

22.1

122

300+

67

62.6

40

37.4

107

200+

33

31. 7

71

68.3

104

200+

229

August 27

Immatures
No.
Banded
Percent

I

t-'
t-'
.pI

Totals

6,266

86.4

!/
Includes pigeons classified

as subadults.

9'85

13.6

7,251

�Table 2.

Time of day of band-tailed

pigeon trapping, 1972.

Percent of All CaEtures
Adults 1/
Males
Females

Percent of
All Males
Captured.£/

Percent of
All Females
CapturedY

Percent of
All Immat ures
Captured

Percent of
All Adults
Capturedl/

Time MIYr

Percent of
All Success ful
Trap Attempts

0600-0959

34.6

68.0

32.0

51.1

23.1

49.3

36.8

1000-1559

48.7

34.3

65.7

36.8

68.0

34.7

52.7

1600-

16.7

56.6

43.4

12.1

8.9

16.0

10.5
I

!/ Including

subadults.

t-'
t-'

VI
I

2/ Adults and subadults only.

�-116-

hunting. As in previous years, hunting was by free permit obtainable at
7 Division Offices and the headquarters of the Monte Vista National Wildlife
Refuge. No limit was placed on total number of permits to be issued and 562
permits were obtained by hunters. This was an increase of 38 (6.8 percent)
over 1971.
Hunter Questionnaire Survez
The experimental hunting season for bandtails in 1972 closed at sunset on
October 1. On October 2 and 3, letters containing a self-addressed, postagepaid return card were sent to all permittees. Only one (.2 percent)
questionnaire was returned for lack of sufficient address. On October 16,
a followup letter was sent to the 176 permittees (excluding the one permittee
for which an insufficient address was available) who had failed to respond
to the initial letter. Results of the questionnaire survey are presented in
Table 3. Data for the 42 non-respondents were calculated using average values
of the respondents to the follow up survey.
While interest in pigeon hunting increased slightly (6.8 percent) there was a
significant decrease in hunter success (41.6 percent versus 58.4 percent)
and total harvest (822 pigeons versus 1,723) in 1972 when compared to 1971.
As in previous years, a significant number of people who obtained permits
did not hunt (47.0 percent). These people probably obtained a permit in
order to hunt if they saw pigeons while engaged in some other activity. If
pigeons were not observed, they did not hunt. It is doubtful that these individuals can be classified as serious pigeon hunters.
Average number of days hunted per hunter increased slightly in 1972 from 1971
(2.6 to 2.5), while number of pigeons bagged per successful hunter decreased
(7.6 to 5.9).
Percent crippling loss (total number of pigeons reported lost
divided by total harvest) (bagged + crippled and lost) was 11.3 in 1972,
similar to the 10.8 percent reported in 1971. Reported crippling loss is minimal and observations of actual cr~ppling loss indicate that it may represent
as much as 20 percent of the b1rdii actually retrieved.
&gt;

The questionnaire was devised in order to ascertain the proportion of the
harvest that occurred in given time periods. It was found that 44.2 percent
of the total harvest occurred on the opening weekend, 20.6 percent occurred
on the second weekend, 9.4 percent occurred on the third weekend, while only
1.5 percent occurred on the last weekend of the season. Not surprisingly,
only 27.2 percent of the total harvest occurred during the non-weekend portion
of the season (15 days) with 19.0 percent of this occurring during the initial
week of the hunting season. Harvest was not well spaced during the 1972
hunting season, as 83.8 percent of the total harvest occurred during the first
9 days. This suggests that either hunting was poor later in the season or
that hunter interest declined as the season progressed. It is believed that
both factors are involved.
Hunting pressure in 1972 was diffuse, although q areas, Durango (17.3 percent),
Monte Vista (13.7 percent), La Veta (8.1 percent) and Colorado Springs (8.1
percent) received almost 50 percent (47.2) of the total hunting effort. In

�Table 3.

Experimental band-tailed

pigeon season, harvest statistics, 1972. !/
Response
First
Letter

Response
Second
Letter

Sum
First and
Second Letters

Number of Permittees Responding

385 (68.5)

135 (24.0)

520 (92.5)

Number of Permittees Hunting

213 (55.3)

65 (48.1)

278 (53.5)

298 (53.0)

Number of Permittees Not ,Hunting

172 (44.7)

70 (51. 9)

242 (46.5)

264 (47.0)

Number of Successful Hunters

95 (44.6)

22 (33.8)

117 (42.1)

124 (41.6)

Number of Hunter Days

539

171

710

762

Days Hunted Per Hunter

2.5

2.6

2.6

Pro j ec t ed

All (562)
Permittees

2.6
I

Number of Pigeons Bagged

523

156

679

t-'
t-'

729

'-I
I

Pigeons Per Successful Hunter

5.5

7.1

5.8

5.9

Pigeons Per Hunter

2.5

2.4

2.4

2.4

Number of Pigeons Crippled and
Lost

68

19

87

93

Pigeons Crippled and Lost Per
Hunter

.3

.3

.3

.3

Total Harvest (Bagged + Crippled
and Lost)

591

175

766

822

Percent Crippling Loss

11.5

10.9

11.4

11.3

!/
Values in parentheses are percentages.

�-118-

all, 271 hunters reported hunting at 54 locations. On a regional basis, 156
of the 271 hunters (57.6 percent) hunted in southwestern Colorado, 68
(25.1 percent) hunted in southeastern Colorado, 27 (10 percent) hunted in
northeastern Colorado, while only 20 (7.3 percent) hunted in northwestern
Colorado.
Many written connnentswere received from hunters with the majority relating
to a lack of pigeons seen'during the open season. This connnentwas valid,
as in many locations pigeons did leave areas where they had been in mid- to
late sunnner just prior to opening of the season. These movements away from
areas where they can be readily observed appear to be normal and are related
to ripening of berry and acorn c~QPS in more inaccessible locations. It is
also probable that some pigeons initiate southward migration at this time.
Wing Survey
Packets containing 5 w~ng envelo~s with pne days' bag to be placed in each
envelope were issued to each pigeon permit holder in 1972. Envelopes containing 516 wings were rece:f.vedf'rom9~ hunters. Additional data were
received concerning hours hunted (443.8), birds crippled and lost (66),
location of kill and banded birds harvested (70). Major harvest areas were
near Durango (34.9 percent of the harvest), La Veta (18.6 percent), and MOnte
Vista (11.8). All other areas contributed less than 6 percent each of the
total harvest. These data are somewhat different than the distribution of
hunters obtained from the questionnaire survey. Thus, Durango with 17 percent
of the total hunters had 35 percent of the harvest, while the Colorado Springs
area had 8 percent of the hunters and 0 percent of the harvest. This suggests
that hunting at Durango was good in 1972, while hunting was extremely poor
in the Colorado Springs area. Crippling loss calculated from the wing survey
was essentially identical (11.2 percent as compared to 11.3 percent) to that
calculated from the questionnaire survey.
Seventy of the 515 bandtails (excluding one wing from a feral pigeon, Columba
livia) reported harvested on the wing ~urv~y were banded (13.6 percent). This
is the highest percentage recorded in any of the 3 years of the experiment
(6.3 percent in 1970, and 9.3 percent in 1971). Percent of birds harvested
that were banded varied from 36.7 (N = 30) at Longmont, 30.0 (N = 20) at Ft.
Garland, 18.7 (N = 96) at La Veta, 12.8 (N = 180) at Durango, to 8.2 (N = 61)
at Monte Vista. Bands were reported from 5 other locations but sample sizes
of wings received were too small (less than 20) for meaningful comparisons.
With the exception of La Veta and Longmont, none of the banded birds shot were
harvested within one-half mile of a trap site.
It is believed that the 13.6
percent banded birds in the harvest is somewhat higher than the actual percentage of bands in the popUlation. However, it is believed that at least 10
percent of the fall population in Colorado in 1972 was banded.
Comparable data were available from both hunter questionnaires and wing
surveys concerning numbers of birds bagged and those crippled and lost by 83
individual hunters. A reporting difference of 15.0 percent (466 in wing
survey versus 536 in questionnaire survey) was found in number of birds killed
and 5.2 percent (58 versus 61) in number of birds crippled and lost for those

�-119-

hunters who s:entin wings and also responded to the questionnaire. The
difference in birds harvested and retrieved could be related to the fact
that hunters only received 5 wing envelopes each and could thus send in a
maximum of only 25 wings. Several hunters reported killing more than 25
pigeons with one hunter reporting a season harvest of 37 birds. The slight
difference in crippling loss can also be attributed to the number of wing
envelopes each hunter received as the number of pigeons reported lost on wing
envelopes was less than on the questionnaire survey. It is probable that
hunters tend to underestimate or report birds crippled and lost.
All wings received were measured (carpal and all 10 primaries) and classified
as immature, subadult and adult in order to derive age structure of the
harvest. Wings were received from 334 adults (64.8 percent), 6 subadults (1.2
percent), and 175 immatures (34.0 percent). Percent of adults was slightly
lower than in 1970 or 1971 (67.8 and 68.0 percent), while percent immatures
increased markedly from 1970 and 1971 levels (26.6 and 21.4 percent). When
the subadult (birds less than 1 year of age but having molted past P5) age
class is added to the immature class, data for all birds less than 1 year of
age are comparable for all 3 years (32.3 percent in 1970, 32.0 percent in
1971, and 35.2 percent in 1972). Age ratios in the 1972 harvest were somewhat
different to that expected,' based on trapping results in late August and early
September. It was expected that substantially more immatures(in percent)
would be harvested in 1972 than in either 1970 or 1971 as all indications were
that production was excellent in 1972. Reasons why the expected increase did
not occur to a greater extent are not fully understood but may be related to
differential migration of adults and immatures. It is also possible that
production was lower than that predicted from trapping results.
Hunter Field Checks
During the 1972 experimental season, emphasis was again placed on contacting
pigeon hunters in the field in order to examine harvested birds for crop
gland activity. Crops of all adult pigeons checked were classified as being
active (crop gland with curds), stimulated (gland apparent but no curds), or
inactive. In addition, sex by gonadal inspection was obtained for as many
birds as possible. In all, 266 pigeons were checked, with 97 being immatures
(36.5 percent), with the remainder (169) being classified as adults. Of the
adults, 23.0 percent were classified as having active crop glands, while 27.9
percent were classified as being stimulated. Percent of crops showing glandular activity (active + stimulated) was significantly higher in 1972 than in
1971 (50.9 versus 28.7), and was similar to that observed in 1970 (52.3 percent).
This supports field observations that indicated that nesting and/or feeding
of young continued late into August and September.
Data on sex was available for 160 adults. Of this number, 64.4 percent were
males, and 35.6 percent were females. Of the males, 55.3 percent exhibited
some glandular development, while 42.1 percent of the females showed development of the crop gland. These data are somewhat similar to that collected in
previous years. Reasons for these disparities are not fully known, but may
be related to time of day of hunting. However, this would not explain the
differences between males and females in crop activity.

�-120When samples of adults examined by area are compared, it is apparent that
nesting was still occurring in most areas of the state during the hunting
season. These data indicated that 41.7 percent of the 24 birds examined in
the San Luis Valley were still associated with nest activities, this percentage was 51.7 percent in the San Juan Basin (N = 118), 53.3 percent (N = 15)
from Wetmore to La Veta, and 58.3 percent (N = 12) at Longmont.
Mortality
Recoveries of 125 band-tailed pigeons .initially trapped and banded in Colorado were reported in 1972. Five of these were initially banded in 1969, 22
were banded in 1970, 25 were banded in 1971, while 73 were initially banded
in 1972. All but 10 were shot recoveries. Of the 115 shot recoveries, 73
were first year recoveries, 21 were second year recoveries, 19 were third
year recoveries, while only 2 were fourth year reoaveries. Including all
recoveries, 85 were recovered in Colorado, 20 in New Mexico, 12 in Mexico, 3
each in Arizona and California, and 1 each in Utah and Washington.
It would
appear that hunting mortality during migration (primarily in New Mexico) is
substantially higher than originally anticipated (Braun 1972b). Overall,
reported hunting mortality remains low and it is doubtful that hunting is
presently an important cause of lIlortality.
Hunting seasons for bandtails were also conducted in New Mexico (September 224), Utah (September 1-23), and Arizona (Occobe r 14-23) in 1972. One Colorado banded bird was recovered on 26 September in eastern Utah, just west of
where it was banded, while 3 were recovered in Arizona on 14 October (2) and
15 October (1). These 3 birds were south and west of initial banding and
presumed breeding locations. Of the 20 birds banded in Colorado that were
shot in New Mexico 2 were harvested the initial weekend of the season, 4
during the first week, 6 the second weekend, 3 the second week, and 5 during
the third weekend of the season. These data sugges t , provided all banded
birds shot were summer residents of Colorado, that migration from Colorado
had started by September,l with major numbers of birds passing through New
Mexico from September 4 t1l.~ouih15. Thus, 1il portion of the Colorado population, possibly as high as 30 percent, had migrated from the state prior to
opening of the hunting season.
No pattern could be discerned in age class, as only 1 of the 20 birds harvested in New Mexico was an innnature. This bird was harvested on 15 September.
Of the 20 birds harvested, 2 were banded in 1970, 4 in 1971, and 14 in 1972.
Four of the 6 birds shot in New Mexico prior to the season opening in Colorado
were banded in 1972. Origin of birds shot in New Mexico varied with the
majority being from La Veta (6), Ft. Garland (4), Monte Vista-Del Norte (3),
Salida (2), and Rye-Beulah (2). Of the 20 banded birds harvested in New
Mexico, only 1 (Montrose) was banded west of the Continental Divide.
Breeding Phenology
Data on breeding phenology were obtained through crop and gonadal inspection
of birds accidentally lost in trapping operations, and those systematically
collected. Additional data were collected from 217 hunter-killed pigeons
examined from September 8 through October 1. Excluding the 217 hunter-killed

�-121-

birds, 138 birds were available for detailed examination.
Of this number,
117 were adults or subadults (53 males and 64 females), while the remaining
21 were immatures.
The sample included birds taken from 8 May through 1
October, with the following monthly totals: May - 15, June - 51, July - 9,
August - 8, September - 52, and October - 3. The 1972 field season represented the last year of this phase of the study.
All pertinent data have
been analyzed and reported in the Job Final -Report for Work Plan 4, Job 5
(Gutierrez 1973).
Wing Measurement

and Molt

Data

Data concerning length of primaries, rectrices, total body length, carpal
length and bill length were obtained from most pigeons collected in 1972.
Once adequate samples are available, these data will be analyzed to examine
differences between age and sex classes of pigeons.
Prior to dissection of
collected birds, plumage characters of each adult specimen were examined and
recorded.
Each bird was then classified as either male or female.
Upon
dissection, gonads were examined and actual sex of each bird was compared with
the classification
based on plumage appearance.
Comparative data were available for 116 adults and subadults (53 males and 63 females) and 10 immatures.
Only 1 bird was inaccurately
sexed.
This bird appeared to be a male by
plumage, but was actually a female.
This percentage of error (0.8) was the
lowest of the 4 years studied.
Information concerning primary molt was available for 1,169 immatures (974
banded and released, 174 from the hunter wing survey, and 21 collected).
Of
the 1,169 immatures, 587 had not molted primary 1, 283 were" molting PI, 180
were molting P2, 80 were molting P3, 26 were molting P4, 8 were molting P5,
no birds were molting P6 and 7, 1 was molting P8, 1 was molting P9, while
3 were molting PIO. Hatching dates were estrurnated usin&amp; data from White
(1973).
Estimated hatching dates for all immatures for which data are available are presented in Table 4.
As in earlier years (Braun 1970; 1971; 1972a) hatching and fledging of bandtailed pigeons were spread over a wide time period.
It is interesting
to
note that of the :j pigeons hatching prior to April 11, ;L was estimated to
have hatched on 6 December 1971, 1 on 12 January, 1 on 31 January, 1 on 6
February, 1 on 7 March, 1 on 22 March, and 1 on 2 April.
It is exceedingly
questionable
if any pigeons hatched in Colorado prior to 1 May, 1972, as
few, if any bandtails occur in Colorado from early December through midApril (Braun 1973).
Young pigeons fledging after early September are not
represented in Table 4, as they had little chance of being shot or trapped.
It is important to note the steady decrease in number of pigeons hatching
after August 9. These data suggest that most young pigeons fledge by
September 10-15.
Thus, hunting after this period should be of little importance in nestling loss.
Of the 1,169 immatures for which data are available, only 166 (14.2 percent)
would have fledged after 1 September, with only 81 (6.9 percent) fledging
after 10 September.
During the hunting season, 51 percent of the adults har~
vested that were examined were found to have some development of the crop

�-122-

glands. These disparities are difficult to understand unless adults feed
free flying young or most adults that had terminated all young feeding
activity had migrated south. Data supporting either of these ideas are not
available, although imttlaturepigeons have been observed begging and being
fed while sitting on power lines.
Table 4.

Estimated hatching dates for wild band-tailed pigeons, 1972.

Time Period

Number
Hatching

Prior to April 11
April 12-21
April 22-May 1
May 2-11
May 12-21
May 22-31
June 1-10
July 11-20

Percent

TilliePeriod·

Number
Hatching

Percent

7

0.6

June 21-30

155

13.3

9

0.8

July 1-10

113

9.7

7

0.6

July 11-20

125

10.7

44

3.8

July 21-30

47

4.0

88

7.5

July 31-August 9

85

7.3

77

6.6

August 10-19

46

3.9

171

14.6

August 20-29

33

2.8

160

13.7

Aug. 30-Sept. 8

2

0.2

Data collected through the survey indicated that about 65 percent of the
harvest was comprised of adults. This would approximate about 534 of the 822
pigeons harvested. When it is realized that 51 percent (272 birds) of this
total exhibited some crop gland development, and that one parent can successfully rear a squab to flight stage, it is readily apparent that hunting as
it presently Occurs in Colorado has little effect on overall production.
Parasitic Load
Information on helminths was available for 137 band-tailed pigeons (116 adults
and 21 immatures) necropsied.
Thirteen (all adults) of the 137 birds examined
had helminth infections. The infection rate of 9.5 percent is about onehalf of that found in 1971, but similar to the 10.4 and 13.2 percent found in
1970 and 1969, respectively. As in other years, more adult females (15.9
percent) harbored helminths than did males (5.7 percent). Tapeworms were
found in 8 birds, while 6 birds had nematodes (1 bird, an adult female, had
both tapeworms and nematodes). Worms were found in pigeons collected as early
as 8 May and as late as 17 September. No deleterious effects of the observed
parasitism were noted. All worms were preserved for later .identification.

�-123-

LITERATURE CITED
Braun, C. E. 1970. Band-tailed pigeon investigations. Colo. Div. Game,
Fish and Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. p.
151-171.
1971. Band-tailed pigeon investigations. Colo. Div. Game, Fish and
Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. October. p. 209-236.
1972a. Band-tailed pigeon investigations. Colo. Div. Wi1d1.
Res. Rept., Fed. Aid Proj. W-88-R. October. p. 123-142.

Game

1972b. Movements and hunting mortality of Colorado band-tailed
pigeons. Trans. No. Am. Wi1d1. and Nat. Resour. Conf. 37:326-334.
1973. Distribution and habitats of band-tailed pigeons in Colorado.
Proc. Western Assoc. State Game and Fish Comma. 53:In Press.
. /
d
Gut1errez,
R. J. 1973. Band-tailed pigeon investigations:breeding an nesting
chronology studies. Colo. Div. Wi1dl. Game Res. Rept., Fed. Aid Proj.
W-88-R. October. p. In Press.
White, J. A. 1973. Band-tailed pigeon investigations:p1umage studies. Colo.
Div. Wildl. Game Res. Rept., Fed. Aid Proj. W-88-R. October. p. In Press.

Prepared by

_--'&amp;~~~.
=:=-:·~r=::-. ---L.~~--':'-'--=-C1ait E. Braun
Wildlife Researcher

_

��October,

-125-

1973

JOB FINAL REPORT

State of

COLORADO

Project No.

W-88-R-18

Work Plan No.

4

Job Title:

Bird Investigations

Job No.
Band-tailed

Period Covered:
Personnel:

Migratory

April 1,

Pigeon Investigations

5

- Plumage

Studies

1971 through March 31, 1973

Clait E. Braun, Jack Corey, J. Edward Kautz, Brett Petersen,
Larry Webster and J. Allen White.

ABSTRACT
Band-tailed pigeons (Columba fasciata) were studied in captivity to develop
techniques suitable for care, maintenance and breeding; to document progression
of molts and plumages from juvenile to adult age; and to investigate the validity
of using plumage characters in accurately distinguishing among juvenile (HY),
sub-adult (SY), and adult (AHY) age classes and between sexes.
These pigeons
were easily maintained in captivity using techniques suitable for domestic
pigeons (Columba livia) when special attention was given to pen size, wire mesh
size and amount of privacy given to breeding birds.
Immature pigeons began
post-juvenal molt at 47 days (n = 9) and completed this molt at 318 days
(n = 47, range = 284-340).
Only three plumage characters were found useful
for identification of HY and SY band-tailed pigeons.
These were the presence
of juvenal wing coverts, secondaries. and primaries.
While juvenal primaries
were still present, it was found that hatching dates could be accurately
calculated up to a mean of 252 days (n = 84). Presence and stage of secondary
molt could be used to identify SY individuals for more than 340 days (n = 24).
The last greater secondary coverts (over S3-S6) were molted by 260 days of age.
Using coloration of the head and breast, it was found that 96 percent of the
immature pigeons examined (n = 110) could be accurately classified according
to sex by 80 days of age.

��-127-

ACKNOWLEDGMENTS

I would like to thank the members of my graduate committee, Dr. Clait E. Braun,
Dr. Dale Hein, Dr. Ronald A. Ryder, and Dr. Vern E. Youngman for guidance concerning my research and course work at Colorado State University.
To Dr. Dale Hein, my graduate adviser~ for reviewing and editing rough drafts
of this thesis and encouragement throughout my stay at Colorado State University, go special thanks. Without his interest I would never have been able
to complete my education and I would never have been associated with the
Colorado band-tailed pigeon investigation.
Special thanks to Clait E. Braun, Wildlife Researcher of the Colorado Division
of Wildlife, my supervisor, close friend and the most industrious researcher
that I have ever met. Without his support this study would never have been
undertaken.
My close friends, Larry Webster and Jack Corey, Colorado Division of Wildlife,
helped design and construct most of the facilities used in this study and
helped care for the pigeons over the past two years. Howard Funk, Wildlife
Research Leader, Colorado Division of Wildlife, provided administrative
assistance. Finally, thanks to all other Division of Wildlife personnel
who contributed to this project.
I would like to thank the Department of Fishery and Wildlife Biology,
Colorado State University, for faculty assistance and use of facilities;
their support was appreciated.
Funding for this project was obtained through the Accelerated Research Program
of the U. S. Fish and Wildlife Service, administered through Colorado Federal
Aid in Wildlife ~toration
Project W-88-R.
To my wife, Julie, for helping care for pigeons, recording data, typing parts
of this manuscript and most of all for encouragement, without which I could
not have started or completed this work.

��-129-

TABLE OF CONTENTS

........

INTRODUCTION • . • . • •
METHODS AND MATERIALS
RESULTS AND DISCUSSION • .
Care, Maintenance and Breeding
Post-juvenal Molt • .
SUMMARY
LITERATURE CITED •

...........
LIST OF TABLES

Table
1

Sequence and timing of primary and secondary
feather molt • • • • • • • • •
••••
•

.....

LIST OF FIGURES

Figure
1

Overall sequence and timing of post-juvenal molt
in band-tailed pigeons (horizontal bars indicate
ranges, exact days in parentheses) • • • . • • .

2

Sequence and timing of post-juvenal secondary molt
in band-tailed pigeons (horizontal bars indicate
ranges, vertical bars indicate means, sample sizes
in parentheses)
••..••••••••••.•.

�.•130-

LIST OF FIGURlS

('ot\tit\"~&lt;i)

Figure
3

Timing and extent of upper wing covert molt
compared to mean stageoi, primary molt

.....

4

Percentage of ~~-tail.d
pigeons molting upper wing
coverts;compared to mean stage of primary molt • •

5

Timing and extent of breast feather molt compared
to mean stage of primary molt . • • • • • • • •

6

Percenta~e of band-r~~
R~geone molting breast
feathers compareci to~me.n titOilge of primary molt

7

Timing and extent of crown, occiput, nape and
upper back molt compared to mean stage of
primary mol t . . . . .

. . . ,. . . . . . . . .

8

Percentage of band-tailed pigeons molting crown,
occiput, nape and upper back compared to mean
stage of primary molt .••
• • • • . • • • • •

9

Weight of band-tailed pigeons from hatching until
beginning of post-juvenal molt (horizontal bars
indicate ranges.,vertical bars indicate means,
sample sizes in parentheses) . • • • • • • • • .

10

Weights of band-tailed pigeons at mean stages of
post-juvenal primary molt (vertical bars indicate
ranges, horizontal bars indicate means, sample
sizes in parentheses) ••••••••••••••

�-131-

PLUMAGE STUDIES OF BAND-TAILED PIGEONS
J.

Allen White

iNTRODUCTION

The band-tailed pigeon (Col~~
fasciata) is widely distributed in the
Western Hemisphere (Goodwin 1967) from I)orth-western South America through
western North America. Two subspecies occur in the United States. A coastal
race (C. f. monilis) is found fro~ ~aja California through southern British
Coltnnbia.- The interior ~ce (C. "f. fasciata) winters in Central Mexico and
migrates seasonally tOA!:izona:- C'oior~do,'New Mexico and Utah (A.O.U. 1957).
Estimates of annual productioll, survival rates for all age classes, and indices
to population tren&lt;is are n~~ded to
a wildlife species. However,
managers must first be able"to cl~ssifi1 individuals into age and sex categories.
Often it is necessary to accomplish this using only certain parts of an animal,
such as wings collected in mail surveys. Unfortunately, the available literature on methods of ascertaining age and sex of band-tailed pigeons was, for
the most part, i~deq~ate for use ;Lna).l management programs.

maP.,e

Miller and Wagner (1955), Silovsky et ill. (1968), Houston (1963) and Drewien
et al. (1966) discussed use of cloacal and bursal characters for determination
of age and sex of band-tailed pigeons. However, cloacal examination is difficult to learn and too slow for large trapping operations during warm months.
Smith (1968), Houston (1963), and Morse (1950) suggested that color of the
plumage of band-tailed pigeons was an accurate means of sex determination.
Braun (1970, 1971, 1972) verJf1ed their reports and found that an experienced
1:~"lOO percent of adult band-tailed
observer could correctly class:J,.,f~::95
pigeons Ln.t;o sex categories using plumage' coloration alone. Silovsky et a1.
(1968) devised a key for age determination of band-tailed pigeons during
September through December. According to Silovsky, useful characters
include buffy tipping on upper wini coverts and primaries, amount of wear
of the primaries and secondaries, and stage of primary molt. His key,
designed for use during fall and winter, has obvious limitations for use
during banding operations in spring and summer. Also, before a satisfactory
key for determination of age could be constructed, it was essential to first
ascertain the age when juvenal plumage characters are lost, and adult
plumage attained.
Few comments in the literature refer to molt of band-tailed pigeons (Houston
1963, Peeters 1962) and no extensive studies of their molt have been made.
Unpublished observations obtained by H. M. Wight (personal communication)
indicated that hatching dates can be calculated for immature band-tailed
pigeons using primary molt patterns. This technique is routinely used for
mourning doves (Zenaida macroura) (Reeves et a1. 1968). Studies of molt in
other species of pigeons (Levi 1969, Niethammer 1970) indicated that age of
band-tailed pigeons could probably be estimated in a similar manner.

�-132-

This study of molts and plumages in captive band-tailed pigeons and time
required for each change, was undertaken to provide knowledge needed to help
manage this species. The objectives were: (1) develop techniques suitable
for care, maintenance and breeding of band-tailed pigeons in captivity; (2)
document progression of molts and plumages from juvenile to adult age; and
(3) investigate the validity of using plumage characters as aids in accurately
distinguishing among juvenile (HY), sub-adult (SY), and adult (AllY)age
classes and between sexes.

METHODS AND MATERIALS
During August and September, 1971, cannon nets were used to capture 72
immature band-tailed pigeons (Braun 1972), in Colorado for subsequent plumage
studies in an aviary. In addition, four immature pig~ons were supplied from
California by F. John Ward of Pebble Beach. In July and August, 1972, a
second group of 28 immatures was trapped and placed in the aviary. Nine
young birds raised by the captive flock in the aviary were added to the second
group of pigeons. All pigeons were marked with serially numbered yellow,
green, or blue plastic bandettes so that each bird could be identified. All
pigeons used in this study, except those from California, were trapped by
Colorado Division of Wildlife personnel at locations near Evergreen, Longmont,
Fort Garland, Manitou Springs, Monte Vista, Del Norte, La Veta, Woodland Park,
and Glenhaven.
Ten pens were used to ascertain type of facilities best suited for keeping
captive band-tailed pigeons. Six of these were in two banks, three pens per
bank and each pen was 1.8 m high, 1.8 m wide, and 3.7 m deep. Other small
pens were 1.8 m high, 1.8 m wide, and 1.8 m deep and 1.5 m high, 2.4 m wide,
and 1.5 m deep. A redwood shed 2.8 m high, 3.0 m wide and 5.6 m long was
used to house sick birds during cold weather. One pen 2.4 m high, 16.8 m
wide, and 22.9 m long was used for breeding. All pens were provided with
adequate perches of both natural limbs and 5-cm by 5-cm lumber. Wooden
boxes, plastic sheeting, and steel or fibre glass roofing material was
provided for shelter as weather conditions required.
Food, water and grit were supplied ad libitum in several different ways
throughout the study. Water was supplied in l2-liter galvanized pans with
or without electric water warmers, depending upon time of year and ambient
temperatures. Suitable grit was available in all pens.
Different feeds were provided to breeding birds and to those being examined
for molt studies. Breeders were fed a commercial pigeon ration (Purina
Pigeon Checkers) which supplied all necessary nutrients. Grain mixtures fed
to pigeons during plumage investigations were varied in order to learn food
preferences. Among the grains tested were whole and cracked dent corn (Zea
mays), field peas (Pisum sativum), whole wheat (Triticum aestivum), whole
milo (Sorghum bicolor), and whole and rolled oats (Avena sativa). A basic
ration of equal parts of whole corn and whole milo was used between trials
of other grains during the first year and exclusively during 1972-73.
Breeding studies to provide known-age immatures began in late April, 1971.
Nest boxes measuring 30.4 cm long, 30.4 cm wide, and 15.2 em deep were

�-133-

provided. Two were placed in each of the six main pens and 18 in the large
breeding pen. Nesting material consisting of pine (Pinus spp.) needles,
sticks, twigs, and some meadow hay and alfalfa (Medicago sativa) hay were
placed in each box and on the floor of each pen. Where possible, burlap was
placed on the sides of each pen so the birds would be .disturbed as little as
possible. Data collected from this segment of the study included: weight of
young at intervals of 3 to 5 days from hatching until beginning of the postjuvenal molt; and age whenpost-juvenal molt began.
During post-juvenal molt of primaries, each pigeon was examined at least three
times each month, with four examinations being preferred. Data concerning each
bird was tape recorded or listed by an assistant for later transcription onto
individual data sheets. Data recorded during examinations included the
following categories:
1.

Sequence and timing of primary feather molt.

2.

Presence or absence of buffy edging on juvenal primary feathers.

3.

Sequence and timing of secondary feather molt.

4.

Presence or absence of fraying or wear on secondary feathers.

5.

Sequence and timing of molt of rectrices.

6.

Timing of tail covert molt.

7.

Presence or absence of buffy edging on j uvenal tail coverts.

8.

Timing and extent (light, moderate, heavy) of upper wing covert molt.

9.

Timing of alular quill covert molt.

10.

Timing of greater secondary covert molt.

11.

Timing and extent (light, moderate, heavy) of breast feather molt.

12.

Color of crown and breast to ascertain earliest time sex may be
determined using plumage characters.

13.

Timing and extent (light, moderate, heavy) of crown, occiput, nape and
upper back molt.

14.

Absence or estimated percentage of adult crescent present.

15.

Absence or estimated percentage of adult nape color present.

16.

Timing and description of eye color change.

17.

Timing and description of cuticle color change.

18.

Timing and description of bill color change.

19.

Timing and description of foot and leg color change.

�-134.-

20.

Weight.

21.

Descriptive connnents on overall plumage aspect of each pigeon (e g ,,
adult or juvenile in overall appearance or unusual observed characteristics).
s

Analysis of all data was confined to calculation of simple descriptive statistics including mean, standard deviation and range. Where possible, arrays of
data were sunnnarized in tables and figures.

RESULTS AND DISCUSSION
Care, Maintenance and Breeding
M. R. Blankenship is probably the most successful breeder of band-tailed
pigeons in the United States, having over 20 years experience and unfailing
results. His techniques are simple. According to Blankenship (personal communication), the three most important considerations in raising any wild
pigeon are: adequate facilities, Prtlviding a minimum of disturbance, and
patience. Care should be kept to a minimum during a set time period each
day, and should be restricted to {ee4ing, watering, supplying nest materials
and cage cleaning. If the keep,eris present at the same time each day, pigeons
will learn to expect him and remain calm. Normally, the best time is around
noon, but never in late afternoon or early morning. These are the peak activity periods for band-tailed pigeons, and if disturbed they may not return to
the nest. It is also deSirable to wear the same subdued-colored clothing
each time the pen is entered. Pigeons, like most birds, appear to recognize
their caretaker and wearing the same clothing each day causes less disturbance.
Although the caretaker can segregate birds into pairs with some success, they
prefer to choose a mate. Once a pair begins to breed they will produce from
four to seven young each year (Blankenship, personal communication). Blankenship believes that the greatest barrier to.raising band-tailed pigeons is the
average caretaker's urge to' constantly"do something. This invariably leads to
stressed birds that will not breed, broken eggs, or abandoned young. The key
is to do as little as necessary.
Levi (1969) described detailed methods for care, maintenance, and breeding of
domestic pigeons. He included chapters on breeding, diseases and disease control, feeds and feeding, and housing. While all of these subjects should concern persons raiSing band-tailed pigeons, the most important consideration is
adequate housing. The principal criterion in selection and construction of
housing should be to simulate the natural habitat of the species (Levi 1969).
This factor accounts for the main differences between adequate facilities for
domestic pigeons (primarily descendents of colonial cave and cliff nesters)
and those most suitable for band-tailed pigeons (solitary pine-oak woodland
nesters). While domestic pigeons require only a small 10ft with many covered
platforms for nesting, band-tailed pigeons do not respond well to those
circumstances.
Large flypens are ideal for keeping and rearing band-tailed pigeons. M. R.
Blankenship (personal communication) produces band-tailed pigeons for sale,

�-135-

using both large flypens and modified courtyard arrangements as described by
Muller et ale (1971). During my study, success was obtained using a breeding
pen 22.9 m long, 16.8 m wide and 2.4 m high. Smaller pens were best suited
for keeping unmated birds or pigeons that were to be examined at regular intervals. Pens smaller than 1.8 m high, 1.8 m wide, and 3.7 m deep were found
unsuitable for band-tailed pigeons, because of the tendency for the birds to
panic when closely approached.
Another major consideration in pen construction is type and size of netting
used. During this study most pigeon injuries and fatalities were directly
caused through use of 5-cm wire mesh, poultry netting. Five-cm mesh resulted
in cut crops, necks and heads. Small pens, where pigeons do not attain high
speeds in flight, may be covered with' 2.5-cm wire netting without concern for
1nJuries. However, large flypens or breeding pens should be covered with
vinyl-covered wire or nylon poultry netting to avoid injuries and fatalities.
In addition to providing perches, breeding pens should contain two or three
nesting structures for each breeding pair. Suitable structures can be made
from small wooden boxes approximately 30 cm long, 30 cm wide, and 10 to 15 cm
deep. While band-tailed pigeons will build a nest of coarse twigs, it is best
to place a padding of hay or grass in each box. Other nesting materials may
be supplied on the floor of the pen. Band-tailed pigeons prefer twigs and
sticks from 2 to 6 mm in diameter, although pine needles were frequently
selected.
Several other requirements must be met in pen construction. All pens should
provide shelter from storms, protection from extreme winds, shade, and privacy
from outside disturbances. Ventilation and drainage of water from all pens is
also important. During fall and winter, 1971, all small pens used in this
study were covered with plastic sheeting to provide protection and shelter.
As a result, 10 birds became sick and seven eventually died. One dead bird
was sent to the Diagnostic Laboratory at Colorado State University; cause of
death was determined to be peritonitis. Another sick bird was taken to the
Colorado State University Small An;imal Clinic for examination and treatment.
The diagnosis was probable aspergillosis (Aspergillus fumigatus), a fungal
condition caused by dampness and moldy grain. Symptoms were similar in all
other sick pigeons and were characterized by a slow decline in weight following
infection, until the birds became extremely weak and thin. Shortly before
death, breathing became labored and traces of blood became apparent in the
saliva. At death, hemorrhaging in the respiratory tract occurred and the lungs
were filled with saliva and blood.
Post-juvenal Molt
At 47 days of age (n = 9) post-juvenal molt began with loss of the innermost
primary (PI). This molt continued until pigeons were 318 days old (n = 47).
However, one individual required only 284 days to complete post-juvenal molt,
and 24 (22 percent) retained secondary 6 and/or S7 for 340 days or longer.
Band-tailed pigeons over 150 days of age were similar to adults in overall
plumage appearance, although these birds could be distinguished from adults
by examination of primary feathers, secondary feathers and upper wing coverts.
Overall sequence of timing of post-juvenal molt is sho~Yn in Fig. 1.

�DAYS OF AGE
25

JUVENAL

PWMAGE

50

75

(4O)

(70)

100

125

150

175

200.

225

250

275

(130)

~

(215)

I

(35)

(1:~0)

TAIL COVERTS

l:2

ffif-.

350

(340)

(100)

RECTRICES

f::

325

-----j

COMPLETE

SECONDARIES

U

300

(60)

r-----------------------

UPPER WING COVERTS

~

(260)

U

&lt;

~

(75)
AWLAR

QUILL COVERTS

GREATER

SECONDARY

(165)

U

I-'

(90)

COVERTS

(260)

I

(45)

.(165)

~-------------~

BREAST

(45)
CROWN,

I

--I

I

OCCIPUT,

NAPE

(200)

I

Pl- P2 P3 P4 PS - ~ P~ ~

~

pIo

(56 )(74) (95) ;(118)(140)(156)(176)(195)(218:)(239)
MEAN STAGE OF PRIMARY

MOLT

Figure 1. Overall sequence and timing of post -j .•
vens 1 molt. in band-tailed pigeons (horizontal bars indicate ranges,
exact days in parentheses).

W
0\
I

�-137-

Immature band-tailed pigeons molted primaries in sequence from the innermost
primary (PI) to the outermost primary (PlO). In this sequence, a new primary
was molted every 19 to 20 days. Each primary was lost and fully replaced in
22 to 26 days, although some minor growth (1 to 4 mm) occasionally occurred
for up to 40 days. Normally, this sequence was not interrupted and no longterm suspension of molt was observed. However, during cold weather in
November and December, 1972, (3 week period: maximum temp = 540F, minimum =
-180F, weekly means = 29.00F, 2.00F, 8.70F, overall mean = l3.20F) 17 pigeons
displayed a short-term (8 to 13 days) interruption of molt. Five stopped on
P5, 9 on P6, and 3 on P7. None stopped on more than one primary. Cold
weather usually has no effect on molt rate in birds (Payne 1972) but Keast
(1968) found that chilling may stop molt for a short time.
Mean number of days for band-tailed pigeons to attain selected stages of
primary development during post-juvenal molt are shown in Table 1. In this
study, primaries were visually classified as empty (E), quill one-fourth of
its full length (1/4), new feather one-half of its full length (1/2), new
feather three-quarters of its full length (3/4), or a new feather that was
full length (F). Using Table 1 it is possible to estimate hatching dates
for immature band-tailed pigeons.
Juvenal primaries are easily distinguished from adult primaries on the basis
of color and shape. Adult primaries are noticeably darker in color and are
gray.
Juvenal primaries are lighter in color and appear brown. This brown
tinge in juveniles is most noticeable at the edge of the primary feathers and
often appears as a buffy edge at the tip. Nearly all (92 percent) juvenile
pigeons had this characteristic on the inner primaries (PI through P6), while
only 24 percent of the immatures examined displayed this characteristic on the
outer primaries :(P7 through PlO). This edge varies in color from nearly white
to brown. An interesting variation in the color of this edge occurs in males
and less commonly in females. I found that 28 percent of the pigeons examined
had an orange edge on the inner primaries (PI through P4). Of these, only 10
percent were females. Braun (personal communication) had also noticed this
characteristic, but did not relate it to the sex of the individual.
Shape was also useful in distinguishing between adult and juvenal primaries.
In adults, PI through P6 were broad and tended to be truncate at the apex,
while primaries P7 through PIO were rounded at the apex. Juvenal primaries
PI through P6 were narrower and rounded, but P7 through PIO were narrow and
pointed when compared to adult feathers.
Secondary feathers are numbered from the outermost (Sl) to the innermost (SlO).
Timing of secondary feather molt is shown in Fig. 2. While there was obvious
overlap in timing of individual secondary molt, the average sequence appeared
to be Sl, S2, S3, SlO, S9, S5, S8, S6, and S7. Secondaries S6 and S7 were
particularly important, because these were the last juvenal feathers to be
molted. Juvenal secondary feathers were rounded and narrow, while adjacent
adult secondaries were obviously truncate and broader in comparison.
One study (Silovsky et al. 1968) used fraying of secondary feathers to identify
immature (HY) band-tailed p~geons. However, during this study no fraying or
wear of secondaries was observed in immature pigeons. Fraying may not have
occurred because of the conditions of captivity, but it seems likely that
some wear would have been noted if this characteristic was a useful indicator
of age.

�-138Table 1.

Sequence and timing of primary and

Mean Days
of Age

Standard
Deviation

Feather Molt
Stage of
Primary Molt

47

2

IE

2

1 1/4

110

0

3

1 1/2

110

0

4

2E

109

0

3

1 3/4

108

0

3

IF

108

0

5

2 1/4

108

0

5

2 1/2

107

0

7

3E

107

0

5

2 3/4

107

0

7

3 1/4

107

0

6

2F

107

0

8

3 1/2

106

0

7

3 3/4

106

0

9

4E

106

0

8

3F

106

0

9

4 1/4

105

0

9

4 1/2

105

0

11

5E

105

0

9

4 3/4

105

0

10

4F

105

0

10

5 1/4

105

0

11

5 1/2

105

0

59
61
65
68
73
79
81
84
90
95

111
114
119
126
129
131
131
136

Secondaries
Most
Often Molting
0

59

105

Sample
Size

9

51

101

secondary feather molt.

---------------------------------------------------------------------------

�-139-

Table 1. Sequence and timing of primary and secondary feather molt
(continued).

Mean Days
of Age

Standard
Deviation

Feather Molt
Stage of
Primary Molt

Sample
Size

145

11

5 3/4

104

0

149

13

6E

104

1

153

14

6 1/4

104

1

155

12

SF

104

1

158

14

6 1/2

105

1-2

165

14

6 3/4

104

1-2

167

16

7E

103

1-2

168

15

6F

103

1-2

173

17

7 1/4

103

1-2

179

17

7 1/2

103

1-2-3

185

18

7 3/4

102

1-2-3

186

21

8E

102

1-2-3

191

19

7F

102

1-2-3

191

22

8 1/4

99

2-3-4

195

22

8 1/2

98

2-3-4

199

21

8 3/4

97

2-3-4

203

24

8F

95

2-3-4-10

205

25

9E

94

2-3-4-5-10

209

25

9 1/4

93

3-4-5-10

216

25

9 1/2

92

3-4-5-10

223

25

9 3/4

91

4-5-9-10

226

27

10E

89

4-5-9-10

Secondaries
Most
Often Molting

----------------------------------------------------------------------------

�-140-

Table 1. Sequence and timing of primary and secondary feather molt
(continued) •

Feather Molt
Stage of
PriIDilryMolt

Sample
Size

Secondaries
Most
Often Molting

9F

87

4-5-9-10

28

10 1/4

85

4-5-9"-10

236

28

10 1/2

85

4-5-6-9

245

28

10 3/4

84

5-6-8-9

252

29

lOF

84

6-7-8-9

265

59

6-7-8

285

31

6-7-8

305

29

6-7

320

25

6-7

340

24

6-7

Mean Days
of Age

Standard
Deviation

230

28

230

Sequence and timing of molt of rectrices was difficult to delineate because
of accidental loss of these featherlil. As.a result, sequence of molt of
rectrices was not discerned during this study. However, timing of this molt
is illustrated in Fig. 1. Because of ease of accidental loss of rectrices,
they appeared to be of little benefit in determination of age in band-tailed
pigeons.
Timing of molt of tail coverts is shown in Fig. 1. Loss of these feathers
through accident was also common and this characteristic was of little value
in age determination because of unreliability and the early age at which
replacement occurred. However, juvenal tail coverts were distinguishable from
adult tail coverts. Juveniles showed a buffy edge similar to that found on
primaries. Of the juvenile pigeons examined, only 12 percent did not show
discernable buffy edging on the tail coverts.
Timing and extent of upper wing covert molt are presented in Figs. 3 and 4.
This molt extends to a maximum of 260 days of age and juvenal wing coverts
are eaSily distinguished from those of the adult. This character therefore
becomes one of the best indicators of age in band-tailed pigeons. Like primaries and tail coverts, wing coverts of juveniles are clearly buffy-edged,
while adults show uniform blue gray colored covert feathers.

�10

I

I

-~(85)

-:-.1

. I

9

I (84)

I

I

8

1(59)

7

1------+---- -(59)

Ul

6

f.------f------- (59)

~

5

fiI
~

o
z
o
o
w
Ul

I,
,

4

,

3
2
1

I --;
I '

I

I

150

125

175

I

t-"
~
t-"

(85)

I

(9'~)

I

I

(84)

I (941)
1(103)

200

250

225

275

·300

325

350

DAYS OF AGE
Figure 2. Sequence and timing of po s t j uve nal . secondary molt in band-tailed pigeons
(horizontal bars indicate ranges, vertical bars indicate means, sample sizes in
c

parentheses)

.

�..

PRIMAR Y MOLT
PI

p~

P?
-

-

~

~

P4
.L

-.

P5

l='h

07

DO

'[")/"1

.----==~~

I

t-'

+'
N
I

I

40

50

60

70

80

90

"j
-.,..---r-""""-"""'-"""'--r---r---r'--r---,..---r'100 110 120 130 140 150 IE-O 170 180 190200210220230240250260
270

AGE IN DAY~:;

Figure

3.

Timing and extent of upper wing COV,'~l'tmolt compared

to mean stage of primary

molt.

�PRIMAR Y MOLT
.•.... ,

to

'0'1

'0"

'DA

1

80

% 60
40

_ ••

2: 46 5'0

r

,

,.

_

••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••

_

• _

••

____

6~...u~~

I

t-"
.po
w
I

I

AGE IN DAYS

Figure 4. Percentage of band-tailed
stage of primary molt.

pigeons molting upper wing coverts

compared

to mean

�-144-

Timing of alular quill covert molt and greater secondary covert molt is
shown in Fig. 1. Alular quill coverts were molted early in post-juvenal
molt and it was difficult to distinguish juvenal feathers of this region
from those of adults. As a result, &amp;lular quill coverts are not useful in
age determination of band-tailed pigeons. Greater secondary coverts of
juveniles are readily distinguished from those new feathers replacing them
during post-juvenal molt. In addition, these are the last upper wing coverts
to be molted. Those greater secondary coverts adjacent to secondaries S4
through S7 were molted last in all cases. Fifty percent of the pigeons
examined retained one or more greater secondary coverts with typical buffy
edging for 220 days (see Fig. 4). Therefore, greater secondary coverts
appear to be a good character for age determination for this species.
Timing and extent of molt of the breast feathers are shown in Figs. 5 and 6.
Since molt in this region begins while birds are fairly young, it is possible
to distinguish sex of bandtails at an early age (see Braun 1972). At 80 days
of age (P2 molt), all pigeons examined had molted one or more breast feathers
and showed enough adult breast coloration to be accurately sexed. During
this study, I classified 96 percent of the immature pigeons examined correctly as to sex, determined from later adult plumsie, at 80 days age or
less. In most cases it {s.possible to determine sex of an immature bandtailed pigeon as soon as one breast feather is replaced. However, one
must look much closer to distinguish between adult female and tMmature
female breast feathers than between the adult male and immature male.
Early in this study it was thought t~~t the neck crescent and irridescent
nape of band-tailed pigeons might be"key characters in age and sex determination. This was subsequently determined not to be the case. Timing and extent
of this molt are shown in Figs. 7 and 8. Since this molt occurs at an early
stage compared to malt of wing coverts, secondaries and primaries, it was
found not to be as good a measure of age in band-tailed pigeons as were
these other characters. Obviously, pigeons without the crescent or metallic
nape should be classifiec:ias ~tures~
At later stages of post-juvenal
molt, however, it is essellti#'t;hat other characters,specifically
the upper
wing coverts, primaries, and secondaries, be checked for positive age classification. As an indicator of sex, these two characters are less useful than
breast feather color. If sex is in doubt and time permits, cloacal examination (Miller and Wagner 1955) should be used.
Color of the eye, cuticle, bill, foot and leg was found to be of little value
in determining age or sex of band-tailed pigeons. In each of these characters,
the immature color is obviously different from that of the adult. The iris
of juvenile pigeons is brownish-pink to brown. This color gradually changed
to reddish-brown after primary molt began with the blood-red eye of the adult
being attained by 100 days of age in all pigeons examined. The color of the
cuticle or eye ring in immatures was found to be highly variable, with pale
shades of pink, orange, or yellow present but with the overall color being
brown. This color also changed by 100 days to the typical dark pink adult
cuticle in 80 percent of the pigeons observed. All pigeons attained typical
cuticle color by 120 days. Eighty-four percent of the immatures examined
had bills, feet and legs noticeably paler than adults. By 85 days of age
all immatures had pigmentation in these areas identical to that of adults.

�PRIMARY MOLT
PI

P2

P3

P4

I

::)~)

P6

P7

P8

P9

PI0

.~'

I

~

VI
I

~~--~~--~~~~--~~--~--~I'--II--r--T--~~r--r--T-~--~--~~-40

50

60

7"0 80

90

100 110120

130 140 150 160 170 180 1902002102202302402502.60
AGE IN DAYS

Figure

5.

Timing and extent of breast

feather

molt compared

to mean stage of primary

molt.

�I_~

PI
100,

P2

1

80

0/0

P3

PRIMARY
P4

MOLT

P5

P6

P7

P8

P9

PI0

60
40
I.

20/
0'

I-'
~

~~~~
i

40

0'\

/$1//!/1'/1?01'(fjm0'W(111'm/P&amp;4'~~~ 1 .'
50

60

70

80

90

HH) 110 120 130 140 ISO

•

.. 1

i

i.

i

i

,

I

1&amp;0 170 180 190200210220230240250

AGE IN DAYS

Figure 6. Percentage of band-tailed
stage of primary molt.

pige ons molting breast

feathers

compared

to mean

I

�PRIMARY MOLT
PI

P2

I

P3

P4

I

i

P5

I~----~----~----~~----~
P6

~-----

I

~
.po.

....•
I

,

4050607080

1

90

1001l012013014015CL60170180190200210220230240250260270
AGE IN DP.YS

Figure 7. Timing and extent of crown,
stage of primary molt.

occiput,

nape and upper back molt compared

to mean

,

�PRIMA:~Y
100

••

• c..

P3

MOLT
P7

P4 .
~."

~~.

f

1 ~J

J

,

•

80

%

60
40

J
•....

.j::'(Xl

20

o

J
I

40

-a It If / /I { /I / , { , . (/ / ( {/ U If!,
I
.i
I
II

50

60

70

80

90

(/

. /I

1

' (III'

D I{t.! (/11'.1./ I,

'. 1

.

I

I

'."

.

I

,

'1"',

100 110 120 130 140 150 160 170 ISO 190200210

22023-0240250

AGE IN DAYS
",,'

':~.

Figure 8. Percentage of band- tailed pigeor·.s niQlting on crown,
compared lomean stage of primary molt.

occiput,

nape and upper back

�WEIGH r IN GRAMS

I

50

1~0

1~0

200

250

300

350

1

I

48-51

295
----+----(8)
281

44- 47

260

40- 43

248

I

36- 39
~

32 - 35

&lt;t:

a

28 - 31

~
0

24-27

Ui

&gt;&lt;

20-23

0

16-19

202

r---(10)

I

(10)

tal

(9)

(10)

I
•....

+:'-

\0

_I

I

150
1-(10)
i--{10)

12-15

102

.

8-11

0-3

(9)

(10)

&lt;t:

4-7

I

I

j

-(10)
( 13)

59
29

1

(14)

_+-1 -(17)

I

I

1·-

50

100

150

,

,....

200

,

,

I

250

300

350

. ..-.-----T---·-·- --~~--,-~~________,

WEIGHT IN GRAMS
Figure 9. Weight of band-tailed
pi ge ons from hatching until beginning of post-juvenal
molt (horizontal bars indicate r an ge s , vertical bars indicate means, sample sizes in
parentheses)

�25

50

75

D~\.YS OF AGE
125
150
175

100

20.0

225

250

275

400
375
350
345 -to340
v:

-IT34.cr-h335 -;p,340
.:

325

t.

]345
. ..",.~.,

l350

335,

~

-&lt;

0::
0
Z
~
f-i
..,...

300
275

.....
0
~

W

I

305

106)

280 I =.
( 107)

I

I . 1(95)

(93)

(99) .

"
I

1(92) '(84)

I-'

1(8~f

(105)

(109)
225
200
PI
P2
(56) (74)

P3
(95)

P4
P5 P6' P7
P8
P9
( 118) ( :.40 )( 15 6H 176)( 195) (2 18)

MEAN STAG::; OF PRIMARY

&lt;:&gt;
I

250

~

VI

PI0
(2 39 )

MOLT

Figure 10. Weights of band- tailed pigeor.s at mean stages of post- juvenal primary molt
(vertical bars indicate ranges, hor i aont.a.l bars indicate means, sample sizes in parentheses)

�-151-

Weights of summer-hatched immature pigeons followed a typical pattern with
a peak (of fall weights) in October (see Figs. 9 and 10). This may be due
to a combination of factors, with deposition of fat during the premigratory
period being most important. It is also possible that this peak could be
explained through the normal increase of weight that occurs just prior to
the time when the most feathers are molted (Payne 1972) apparently because
of the intense growth of feather papillae during this time. It was noted
that at this time the pigeons were 80 to 140 days 01d and were at the peak
of molt activity (Figs. 1, 3, 5, and 7).

SUMMARY
The results of this study indicate that band-tailed pigeons readily adapt to
captive conditions if special attention is given to cage size, wire mesh
size used in pen construction, and if not disturbed at frequent intervals.
In most cases, except where pigeons must be handled regularly, large pens
with small wire mesh, nylon mesh or vinyl-covered wire mesh netting are
recommended.
Post-juvenal molt in band-tailed pigeons began at 47 days (molt of primary
PI) and continued until 318 days (secondary S6 or 87 fully replaced) with
a peak of molt occurring from 100 to 140 days of age.
Of the plumage characters observed, only breast feather coloration was found
to be a valid indicator of sex in band-tailed pigeons over 80 days of age.
Primaries, secondaries, and upper wing coverts are the only three characters
useful to determine age of these pigeons. Through use of post-juvenal primary molt, accurate estimates of hatching dates are possible and by using
post-juvenal secondary molt rough estimates (BY or SY) can be made up to 340
days of age.
It is hoped that with the knowledge gained through this study managers will
be better able to delineate the popUlation dynamics of the band-tailed
pigeon through estimates of mortality in all age classes and improved
estimates of annual production.

LITERATURE CITED
American Ornithologists' Union. 1957. Checklist of North American Birds.
5th ed. Lord Baltimore Press, Baltimore. 691 pp.
Braun, C. E. 1970. Band-tailed pigeon investigations. Colorado Division
of Game, Fish and Parks. Game Res. Rept., Fed. Aid Proj. W-88-R.
Oct. pp. 151-171.
1971. Band-tailed pigeon investigations. Colorado Division of Game,
Fish and Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct.
pp. 209-236.
1972.
Wildlife.

Band-tailed pigeon investigations. Colorado Division of
Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. pp. 123-141.

�-152-

Drewien, R. C., R. J. Vernimen, S. W. Harris, and C. F. Yocum. 1966.
Spring weights of band-tailed pigeons. J. Wi1d1. Manage. 30(1):190-192.
Goodwin, D. 1967. Pigeons and doves of the world.
London. 446 pp.

British Mus. Nat. Hist.,

Houston, D. B. 1963. A contribution to the ecology of the band-tailed
pigeon, Columba fasciata Say. M.S. Thesis, Univ. Wyoming, Laramie.

74 pp.

Keast, A. 1968. Moult in birds of the Australian dry country relative to
rainfall and breeding. J. Zool. 155:185-200.
Levi, W. M. 1969.
667 pp.

The pigeon.

Levi Pub1. Co., Sumter, South Carolina.

Miller, W. J., and F. H. W~gner. 1955. Sexing immature columbiformes by
cloacal characters. Auk 72(3):279-285.
Morse, W. B. 1950. Observations on the band-tailed pigeon in Oregon.
Western Assoc. State Game and Fish Comm. 30:l02~104.

Proc.

Muller, H. D., D. D. Neill, and W. J. Werner. 1971 Reproduction, raising
and releasing gray partridge. Colo. State Univ. Expt. Sta., Fort
Collins. 16 pp.
Niethannner, G. 1970. Zur mauser der ringeltaube (Columba palumbus).
Ornith. 111:367-377.

J.

Payne, R. B. 1972. Mechanisms and control of molt. pp. 103-155. In
Farner, D. S. and J. R. King. Avian Biology, Vol. II. AcademiC-Press,
New York. 612 p.
Peeters, H. J. 1962. Nuptial behavior of the band-tailed pigeon in the San
Francisco Bay area. Condor 64(6):445-470.
Reeves, H. M., A. D. Geis, and F. C. Kniffen. 1968. Mourning dove capture
and banding. U. S. Fish and Wildl. Serv., Spec. Sci. Rept.--Wildlife
No. 117. 63 pp.
Si1ovsky, G. D., H. M. Wight, L. H. Sisson, T. L. Fox, and S. W. Harris.
1968. Methods of determining age of band-tailed pigeons. J. WildL
Manage. 32(2):421-424.
Smith, W. A. 1968.
Fish and Game.

The band-tailed pigeon of California.
54(1):4-16.

Prepared by __~J~.~A=1=1=e=n_Wh~=1·~te=.1/ ...t-, 1'. ~.
Approved by ~~(;{~~~~__ ~L~'-L~~~~~
Clait E. Braun
Wildlife Researcher

_

_

California

�October, 1973
-153-

JOB FINAL REPORT

State of

COLORADO
----~~~~~----------

Project No.

W-88-R-18

Work Plan No.

4

Job Title
Period Covered:
Personnel:

Migratory Bird Investigations

6
Job No.
Band-tailed Pigeon Investigations:
Breeding and Nesting Chronology Studies

April 1, 1969 through March 31, 1973

Clait E. Braun, Howard D. Funk, Ralph J. Gutierfez, J. Edward
Kautz, Chet McCord, Brett Petersen, Michael Robinson, Mark
Stromberg, Michael Watkins and J. Allen White.

ABSTRACT
The reproductive biology of the band-tailed pigeon (Columba fasciata) was
studied from May through September during 1969-1972 in Colorado, and during
1972 in New Mexico. Gonads, body weight, percent body fat and crop activity
from 619 male and female pigeons were used in the analyses of gonadal cycles
of wild birds. Photoperiodic responses were studied experimentally during
1972-73 to help explain the unusual reproductive patterns in this bird. The
breeding season extends from May through August and two nesting attempts
probably occur during this period. This pigeon normally lays one egg per
clutch, but up to eight percent of the females may lay two eggs per clutch.
Fall breeding was observed in one population of pigeons, indicating that bandtailed pigeons responded to environmental conditions other than photoperiod.
The photoperiodic responses showed that these pigeons do not exhibit a
characteristic refractory period and that testes responded to light periods
as short as 10 hours. Ovarian follicles of unmated females can regress once
they have become enlarged (e.g. greater than 5.5 rom in diameter). The testes
of captive birds held in outdoor aviaries continued gonadal regression from
23 September 1972 until the winter solstice (during December), at which time
they began to increase. Full breeding capability was reached during March, 1973.
Female pigeons housed with these males showed no follicle growth until March.

��-155-

CONTENTS

...

Abstract
Contents
List of Tables
List of Figures • • • • •
Acknowledgments •
Introduction
Materials and Methods
Gonadal Studies
Photoperiod Studies
Results
Gonadal Cycles • • • •
Photoperiod Studies
Body Weight, Body Fat, Crop Activity •
Discussion •• • •
Summary • • • • • •
Literature Cited

.. . . ... .

....

LIST OF TABLES
Table

Page

1

Ovarian condition of female band-tailed pigeons from
Colorado and New Mexico 1969-1972.

2

Correlation coefficients of testes volume and body
weight in male band-tailed pigeons.

LIST OF FIGURES
Figure

Page

1

Geographical subpopulations of band-tailed pigeons
within Colorado and New Mexico.

2

Testes volumes of male band-tailed pigeons from
Colorado in 1969 •. Vertical lines indicate ran~e,
horizontal lines mean, box standard error of mean.
Sample size is given above range line.

3

Testes volumes of male band-tailed pigeons from
Colorado in 1970. Vertical lines indicate range,
horizontal lines mean, box standard error of mean.
Sample size is given above range line.

4

Testes volumes of male band-tailed pigeons from
Colorado in 1971. Vertical lines indicate range,
horizontal lines mean, box standard error of mean.
Sample size is given above range line.

�-156-

LIST OF FIGURES (continued)
Figure

Page

5

Testes volumes of male band-tailed pigeons from
Colorado in 1972. Vertical lines indicate range,
horizontal lines mean, box standard error of mean.
Sample size is given above range line. Dashed line
and box represent testes data from southwestern
New Mexico.

6

The relationship of testes volume, and stage of
spermatogenesis. Range, mean, standard error
of mean, and sample size are shown.

7

Relationship of 30 day photoperiod treatments to
testes development. Beginning volumes are indicated
by "A", ending volumes at same photoperiod as liB"
Mean, range, standard error of mean are shown.
OJ

8

Gonadal cycles and body weights of captive male
band-tailed pigeons. Sample size is four in all
cases. Mean, standard error of mean, and range
are shown.

9

Gonadal cycles and body weights of captive female
band-tailed pigeons. Sample size is four in all
cases. Mean only is shown for follicle diameter.
ACKNOWLEDGMENT S

I would like to thank M. Kaye Giffin for encouragement and help in every
phase of the research and typing of the manuscript. Without her help I
surely would not have completed this work by this,date.
Clait E. Braun, Colorado Division of Wildlife, and Thomas P. Zapatka, New
Mexico Department of Game and Fish, collected all field specimens.
I would like to thank my major professor, J. David Ligon, for his support
of the project and sharing of valuable ideas throughout my studies. My
graduate committee members, Donald W. Duszynski, J. Scott Altenbach, James
R. Gosz, and Clait E. Braun critically reviewed the manuscript and offered
valuable suggestions. Russell Lenth, Department of Mathematics wrote the
computer programs used in this study, Steven Cito helped construct aviaries,
and Phillip W. Day, Medical School, provided ketamine hydrochloride and
technical assistance.
I acknowledge financial Support of the Colorado Division of Wildlife,
Federal Aid in Wildlife Restoration Project W-88-R, and the United States
Fish and Wildlife Service's Accelerated Research Program. Spencer Amend of
the Migratory Bird Population Station, Laurel, Maryland, was instrumental
in obtaining support from the Accelerated Re~earch Program.
I

Ralph J. Gutierrez
Department of Biology
University of New MeXico, Albuquerque
May 1973

�-157-

BAND-TAILED PIGEON INVESTIGATIONS
BREEDING AND NESTING CHRONOLOGY STUDIES
/

Ralph J. Gutierrez

INTRODUCTION
The band-tailed pigeon, a forest-dwelling columbid, consists of two major
populations in western North America a coastai and an interior population.
The interior population reported on here, inhabits Colorado, New Mexico,
Arizona, Utah, Texas and Mexico. This bird is one of several species of
western North America which sometimes exhibits unseasonal and variable
nesting periods (e.i. see Ligon 1971). Active nests have been found in some
portion of the species' range in every month of the year (Swarth 1900,
Thayer 1909, Stephens 1913, Lambl926, Abbott 1927, Vorhies 1928, Neff 1947,
MacGregor and Smith 1955).
Although several studies of the breedf.ngbiology of the band-tailed pigeon
have been undertaken on the Pacific Coast;population (Glover 1953, MacGregor
and Smith 1955, Houston 1963, March and Sadleir 1970, Zeigler 1971) few have
been conducted on the interior population (Fitzhugh 1970). The breeding cycle
of the interior band-tailed pigeon is poorly understood, although many observations of nests have been reported (Neff 1947, Fitzhugh 1970).
This study was undertaken to better understand the seasonal breeding cycle
and breeding potential of band-tailed pigeons in Colorado and New Mexico.
Results of the field studies on nesting and associated literature review have
previously been reported by Braun and Benson (1972) and will not be repeated
in this final report.

METHODS !ND MATERIALS
Gonadal Studies
Band-tailed pigeons were obtained from trapping-banding mortalities, farmer
kills (as a result of crop depredations), hunter kills, and systematic collections. The last occurred primarily during May through August 1972, and provided fewest birds. Gonadal materials were collected from 144 birds in 1969,
135 in 1970, 171 in 1971, and 169 in 1972. May through September is the
period that most band-tailed pigeons are present in Colorado and New Mexico.
All data from 1969, 1970, and 1971 were taken from birds collected in Colorado.
Pigeons from both Colorado and New Mexico were studied in 1972. Age was determined as adult or juvenile by plumage characteristics (Silovsky et al. 1968).
C. E. Braun (unpublished data) classified some subadults in 1970-72. Sex was
determined by inspection of the gonads.
Prior to 1972, all collected specimens were enclosed in plastic bags and
frozen. Later, each bird was thawed, weighed (nearest 0.1 gram) and the

�-158-

gonads measured in situ with vernier calipers to the nearest 0.1 mm. The
gonads were excised and stored in either Bouin's fixative or 10 percent
buffered formalin. All collection procedures were the same in 1972 except
that gonads were measured in situ and excised within 15 minutes of death and
fixed in either Bouin's fixative er FAA.
Crop activity was determined by visual inspection of the inner layer of crop
lining and assigned to one of three categories - active, inactive, or stimulated. This visual characterization followed the classification developed
for coastal populations of band-tailed pigeons (March and Sadleir 1970,
Zeigler 1971). Crops were fixed in Bouin's fixative or 10 percent buffered
formalin. The skin, feet, wings, and head were removed; as much fat as
possible was retained on the body. Crop, gizzard and gut contents were
removed and then the remaining carcass (including bones) was thoroughly ground
following procedures outlined by the Association of.Official Agricultural
Chemists for the determination of body fat percentage (Horwitz 1960). Weight,
fat, and crop activity were used as physical parameters indicative of changes
in breeding activity.
For purposes of statistical analyses, pigeons were separated into geographical
subpopulations within Colorado and New Mexico. Braun (1972) established Colorado subpopulations on the basis of less than 10 percent movement between subpopulations; mountain ranges were used to delineate subpopulations in New
Mexico (Fig. 1).
Volumes of the left testis were calculated using the formula for an ellipsoid,
V = 1/2 LW2 where L = 1/2 length and W = 1/2 width at widest point (Lofts et
al. 1966). Samples of testes of different volumes were processed using standard histological techniques and sectioned at 10 p to observe the relationship
between volume and spermatogenic state. Sections were stained with Harris'
haematoxylin and counter-stained with eosin. Spermatogenic stages were classified according to Lofts et al. (1966).
The diameter of the largest follicle was measured and in most cases the
number of follicles greater than 5.5 mm was recorded. Follicles of unmated
females are said not to exceed 5.5 mm; when pair bonds form rapid growth of
the follicle (up to 20 mm) can occur in a short time (Bartelmez 1912). All
ovaries with follicles less than 2.0 mm were classified as granular. Each
ovary was examined macroscopically for ruptured follicles (Meyer et al. 1947).
Photoperiod Studies
Adult pigeons were trapped using cannon nets near La Veta, Colorado in late
August 1972. These birds were transported to the University of New Mexico
and placed in outdoor aviaries (8' x 8' x 8')•. Sex was determined by laparotomy.
Experiments designed to illustrate the relationship of testes growth to day
length are described with the results for the sake of clarity. In addition,
food type was the experimental variable in one experiment.

�-159-

.'

COLORADO

NEW MEXICO
.-

Fig. 1.
Colorado

.;

Geographical subpopulations
and New Mexico.

.~..•.

of band-tailed

pigeons

within

�-160-

All birds housed in outdoor aviaries were fed a mixture of cracked corn and
wheat; grit and water were provided ad libitum.
Perch space sufficient to
prevent excessive aggressive behavior-was provided.
Roofing covering half
of each enclosure provided shade and weather protection.
Birds in the laboratory were housed in cages (4' x 3' x 2'), with either one
or two birds per cage. Temperature was about 250 C. Each experimental group
received the light intensity provided by four 150 watt light bulbs.
These
birds also were fed wheat and cracked corn (except group A of experiment 2).
Water and granite grit were always available.
Gonadal activity was determined by laparotomy.
The birds were anesthetized
with an intra-muscular injection of ketamine hydrochloride
(Ketaset).
Dosage
varied from 30 milligrams/kilo
in early fall to 100 milligrams/kilo
in winter.
The reason for this change in sensitivity to the drug is not fully understood,
but may be related to seasonal change in the physiological state of the birds.
Ether and methoxyflurane
(Metophane), both inhalent anesthetics, proved
inefficient and dangerous to the birds.
Each bird was weighed and the operation begun within 7-10 minutes of injection.
Feathers were wetted with
alcohol to expose the area of the last rib and top of the leg on the left
side of the body.
A dorsal-ventral incision was made posterior to the last
rib through the skin, fat layer, and first mesentary above the muscles.
Two
blunt probes were used to push the muscle aside and break the mesentary to
expose the abdominal cavity.
A sharp-focus light source was then directed
into the abdominal cavity to illuminate the gonads.
The length and width of
the testes were measured by inserting dividers over the testes (length and
width) and withdrawing these dividers and placing the points on a piece of
paper.
These points were then measured with vernier calipers.
Little or no
error was encountered using this method as opposed to measuring with calipers
in situ. The left testis was measured in each case, as was done with birds
taken in the wild.
The largest follicle of the females' ovaries was measured
in the same manner.
In all photoperiod experiments,
gonads were measured by experimental laparotomy at the beginning and end of each photoperiod treatment.
Statistical
tests were used to determine whether or not change had occurred between
starting and ending volumes as a result of light treatment.
The testes of
the group showing no statistical difference in volume increase (10 hr photoperiod) were examined histologically.
Gonads of band-tailed pigeons on
natural photoperiod were measured about the first of each month.
Student's t distribution was used to test significant differences between
two means.
Analysis of variance was used for significance testing when more
than two variables were involved.
Correlation analysis was used to determine
if a consistent relationship existed between gonad size and body weight,
body fat, or crop activity during the entire summer, early summer or late
summer for all years.
The level of statistical significance used was at
least equal to 0.05 in all tests unless otherwise indicated.
RESULTS
Gonadal

Cycles

Testes volumes of males are shown in Figs. 2-5. Gonadal cycles of males
could not be pooled for all years, because there is evidence that significant

�-161-

900

ISO

55
100

7S0

100

'SO

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'"

1

sso

0

8

M

E
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SOO
u..o

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:&gt;
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W

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'00

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300

na
200

3

~

ISO

100

50

IS

MAY -

31

IS

JUNE

30

IS

JULY

31

IS

.•, - AUGUST

31

3

,
15

30

"SEPTEMBER'

Fig. 2. Testes volumes of male band-tailed pigeons from Colorado in
1969. Vertical lines indicate range, horizontal lines mean, box standard
error of mean.
Sample size is given above range line.

�-162-

14

900

ISO

aoo
6
7S0

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9

710

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so

IS

MAY

31

IS

JUNE

30

IS

JULY

31

15

., AUGUST

31

IS

30

"SEPTEMBER'

Fig. 3 •. Testes volumes of male band-tailed pigeons from Colorado in
1970. Vertical lines indicate range, horizontal lines mean, box
standard error of mean. Sample size is given above range line.

�-163-

90.

6

5

.~o
3
810

no
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6S0

7
600

1

~~o

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&amp;

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+

so

IS

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II

IS

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30

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JULY

31

IS

31

IS

30

.. AUGUST ••SEPTEMBER-

Fig. 4. Testes volumes of male band-tailed pigeons from Colorado in 1971.
Vertical lines indicate range, horizontal lines mean, box standard error
of mean. Sample size is given above range line.

�-164-

'HO

900-

&amp;50

I

o

aoo2
]So

700

I

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600.

...,.

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E
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7
T

I
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31

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-IAAY

JUNE

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+
30

- ••-AUuUST---SEPTEMBER.'

Fig. 5. Testes volumes of male band-tailed pigeons from Colorado
1972.
Vertical lines indicate range, horizontal lines mean, box
standard error of mean.
Sample size is given above range line.
Dashed line and box represent testes data from southwestern New
Mexico.

in

�-165-

differences exist between years (e.g. 1969 vs. 1971). Subadult testes are
significantly smaller than those of adults. They may develop more slowly
as in the wood pigeon (Columba palumbus) (Lofts et al. 1966). However,
subadults were included with adults in the analyses because histological
examination of testes indicated a capability to breed as early as adults.
Also, subadults have been observed to breed as early as April in captivity
(C. E. Braun, personal communication).
The period 16 A~gust-15 October was the only one studied in which a significant decrease in testes size occurred. The decrease occurred in each year
except 1971, when testes remained large until September.
Only one locale-time set of data differed significantly from the others
(Fig. 5). These pigeons were taken in September 1972 from the Gila National
Forest in southwestern New Mexico (area 17 in Fig. 1). Testes from this
group were larger in early September than those from any other area, including birds at the same latitude (350 N) in the Lincoln National Forest (area
16 in Fig. 1).
I checked many areas in New Mexico for acorns (Quercus spp.) in 1972, but
found them only in southwestern New Mexico, within 50 miles of area 17 (Fig.
1). However, an average acorn crop apparently was produced in Colorado (C.
E. Braun, personal communication).
Conditions of ovaries from females collected in Colorado and New Mexico are
listed in Table 1. A one-way analysis of variance showed no significant
differences between 1969 and the other collection years, ,between subadults
and adults, or between time periods. This is not unexpected, since an
enlarged follicle is a temporary, short-term condition. The female pigeon
undergoes a rapid final stage of follicle enlargement when sexually stimulated by the male (Bartelmez 1912). The data do indicate, however, that most
females taken (88 percent) were in breeding condition (follicle diameter
2.1-5.5 mm+) from May through August. Altogether, 19 birds (8.6 percent)
had two or more follicles greater than 5.5 mm. Seven birds contained either
an egg and large follicle or two ovulated follicles. Females taken from the
Gila National Forest in September 1972 showed significantly larger follicles
than those taken at the same time from other areas (Table 1). No birds
from this area had granular ovaries. One pigeon collected on 2 September
1972 had two large ruptured follicles (15.5 and 10.7 mm, after fixation and
storage). Another bird had one large ruptured follicle of 10.1 mm.
Results of the histological examination of testes are shown in Fig. 6. A
significant correlation (r = .654, n = 46; P&lt; 0.01) exists between increase
in testes volume and increasing stage of spermatogenesis. All testes examined
above 181 mm3 volume were in stage 5 (producing sperm). On the basis of this
examination, most subadults were thought to be in breeding condition. Juvenile birds were not considered in the gonadal analyses because statistical
analyses of 23 individuals (X = 11.0 mm3; standard error of the mean = 5.6;
range = 4.5 - 30.4 mm3) indicated they were not capable of breeding.

�Table 1.

Ovarian condition of female band-tailed pigeons from Colorado and New Mexico, 1969-1972.

Diameter of Largest Follicle
&lt;2.0 mm
2.1 to
Granular
5.4 mm
&gt; 5.5 mm
Number of Birds in Size Class
1-15 May

No. of Birds
with 2
Follicles
&gt; 5.5 mm

No. of
Birds With
3 or More
Follicles
&gt; 5.5 mm

No. of Birds
With an Oviductal Egg
or Ruptured
Follicle

0

2

2

0

0

0

2

6

9

1

0

0

1-15 June

1

9

6

2

1

2

16-30 June

9

17

10

3

3

4

1-15 July

4

35

28

4

0

10

16-31 July

0

5

7

0

1

3

1-15 August

1

9

2

1

0

2

16-31 August

3

5

0

0

0

0

1-15 September

21

4 (5)*

(3)

(1)

(2)

(2)

16-30 September

16

0

0

0

0

0

Total

57

97

67

12

7

23

16-31 May

I

t-'

0\
0\
I

*(n) indicates birds which were taken in southwestern New Mexico 1972.

�-167-

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W

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(IJ

en

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300

250

3

200

3

ISO

100

so

6

$ $

I

0

$
3

STAGE

OF

II

s

SPERMATOGENESIS

Fig. 6. The relationship of testes volume and stage of spermatogenesis.
Range, mean standard error of mean, and sample size are shown.

�-168-

Photoperiod

Studies

Results of the photoperiod

experiments

are shown in Figs. 7-9.

Experiment l.--Five males with regressed or regressing testes were subjected
to 14 hr light - 10 hr dark photoperiod from 23 September to 23 October 1972.
These birds showed a highly significant (P&lt;O.Ol)
testicular volume increase.
This experiment was designed to determine the presence or absence of a refractory period in band-tailed pigeons.
The responses obtained suggest that
these pigeons are continuously photoresponsive.
Experiment 2.--Two groups of five males with regressed testes were subjected
to 12.5 hr light - 11.5 hr dark photoperiod from 30 November 1972 to 3 January 1973. One group was fed only acorns and the other group a mixture of
wheat and cracked corn. Band-tailed pigeons appear to be dependent on mast
production (acorns and seeds of pinon pines, Pinus edulis) over much of their
range (Neff 1947, Fitzhugh 1970), although they take a wide variety of vegetable foods (Neff 1947, Salt 1953).
I hoped to obtain information both on
the role of food on reproductive patterns and on the effects of this photoperiodic regime on gonadal development.
Additionally, exposure of the birds
to a 12.5 hr light regime 30 days after experiment 1 might demonstrate a
refractory period in the event the birds in experimen~ 1 had not yet become
refactory.
A two way analysis of variance (light and food) showed a highly significant
(P&lt; 0.01) increase in testes volume of both groups of birds.
No significant
increase could be attributed to the type of food the birds received, nor was
there a significant interaction between the two variables.
Experiment 3.--Four males with regressed testes were subjected to 10 hr light _
14 hr dark photoperiod from 15 January to 15 February 1973. Ten hours of
light caused some increase in testes size, however, the increase was not
statistically significant.
Histological examination of the testes showed
spermatogenesis had begun (stages 2 or 3, see Lofts et ale 1966) in all but
one bird.
Experiment 4.--Eight males and four females were housed in outdoor aviaries
from 23 September to 1 November 1972.
Five of the males and one female were
lost and subsequently were replaced by a single male and female.
These eight
birds (two pairs in each aViary) were maintained from 1 November 1972 to 3
April 1973.
Since band-tailed pigeons normally overwinter in Mexico, specimens
available for gonadal analyses are rare. Accordingly, this experiment was
designed to monitor gonadal change throughout the fall, winter, and spring
in relation to naturally decreaSing and increasing photoperiod.
Apparently, male pigeons continue a slight testicular regression until day
length begins to increase (sometime during the period 1 December _ 4 January).
The testicular increase for this period was significant (P &lt; 0.01).
Testes
volume did not increase significantly between 4 January - 1 March 1973. The
testes again increased significantly in the period 1 March - 3 April 1973.

�-169-

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HOURS OF llC,HT

Fig. 7. Relationship
of 30 day photoperiod treatments
to testes
development.
Begtnnf.ng volumes are indicated by "A", ending volumes
at same photoperiod as "B". Mean, range, standard error of mean
are shown.

�-170-

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Fig. 8. Gonadal cycles and body weights of
captive male bandtailed pigeons.
Sample size is four in all cases.
Mean, standard
error of mean, and range are shown.

�-171-

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Fig. 9. Gonadal cycles and body we~ghts of captive female bandtailed pigeons. Sample size is f4ui in all cases. Mean only is
shown for follicle

diameter.

:~"

�-172-

The ovaries of female pigeons
tember 1972 and 1 March 1973.
and 3 April 1973.

showed no significant growth between 23 SepFollicles grew significantly between 1 March

Experiment 5.--Two females were held in outdoor aviaries from September 1971August 1972 under natural photoperiod to observe ovarian change in the
absence of a male.
Fem~le rock doves (Columba livia) and wood pigeons apparently do not undergo final ovarian development if no male is present
(Bartelmez 1912, Murton and Isaacson 1962).
Ovaries of both females remained
granular (follicles less than 2.0 rom in diameter) until the period 1 April _
1 May 1972, when they increased to 5.0 and 4.2 rom. They reached maximum
follicular diameter in June (6.4 rom, 6.8 rom), then began to decrease gradually
until September, when they were once again classified as granular.
Bartelmez
(1912) suggested that in domestic pigeons a follicle greater than 5.5 rom
would continue to increase until ovulation occurred.
Others have used this
5.5 rom criterion as a measure of pigeon breeding activity (Lofts et al. 1966,
March and Sadleir 1970).

Body Weight,

Body Fat, Crop Activity

Correlation coefficients of body weight, body fat, and crop gland to gonad
size for males are listed in Table 2. These were computed both for the entire
summer and for each half of the summer (early and late).
A significant positive correlation exists between testes volume and body
weight during early summer for all years, and a significant positive entire
summer correlation for 1969 and 1970. The slight indication of a negative
correlation in late summer is not significant because weight changes are
not great (Figs. 8 and 9). A significant correlation between body weight
and gonad size was obtained.
No consistent significant correlations were
observed with respect to body fat or crop activity in males or to body weight,
body fat, or crop activity in females for all years and time periods.
The
relationship of body weight to gonad size is clearly seen in Figs. 8 and 9.

DISCUSSION
Band-tailed pigeons breed in Colorado from at least May through August.
The
season may be even longer, however, no birds were collected during March or
April.
In New Mexico the breeding season is Similar, except that fall
breeding apparently has occurred in southwestern New Mexico.
The length of
the breeding season is not a latitudinal response per se as suggested by
March and Sadleir (1970), but almost certainly is a response to environmental conditions.
Two populations of pigeons sampled in September 1972 at
the same latitude (330 N), but 150 miles apart, document this.
Specimens
from one population were in breeding condition, whereas the others were not.
Acorns were locally abundant in southwestern New Mexico in 1972 and bandtailed pigeons apparently bred in the fall in response to abundan t acorns.
Ligon
(1971) found pinon jays (Gymnorhinus cyanocephalus) breeding in fall
in southwestern New MeXico, presumably in response to an abundance of pin~n
pine seeds.

�"

-173-

Table 2. Correlation coefficients of testes volwne and eody weight in male
band-tailed pigeons.

Year

Entire
Sutmner

Early
Summar

Late
Summer

1969

r = 0.242*

r = 0.294*

r = -0.347 ns

1970

1971

1972

(67)

(57)

r = 0.514*

r = 0.554*

(10)

r =

0.341 ns

(49)

(36)

(13)

r = 0.250 ns

r = 0.54:)*

r = -0.157 ns

(35)

(19)

(16)

r = 0.114 ns
(56)

*

- significant (P&lt;0.05)

ns

- not significant (P&lt;0.05)

r = 0.442(25)

r = -0.092 ns
(31)

(n) - sample size

Nomadic bahavior of band-tailed pigeons in response to mast production has
been suggested by Neff (1947) and Smith (1968);· Braun (1972), however, found
that Colorado pigeons have a high fideliti to a given area. The production of
acorns and pinon seeds in the southwest is not predictable. Since band-tailed
pigeons apparently do not possess a true refractory period they may move until
locating a food supply large enough to support a breeding effort, regardless
of increasing or decreasing daylength. Reproductive success must be considered the ultimate factor in the control of any species' breeding cycle.
Food as well as light probably interact as proximate factors which trigger
breeding in band-tailed pigeons. Nomadism is a reasonable response to local
shortages of food. The great mobility of these pigeons can be considered as
an adaptation which allows breeding in an environment with an Wlpredictable
local food supply. Numerous observations of nomadic behavior in this pigeon
have been made in southwestern New Mexico (J. D. Ligon, personal cotmnWlication).
In this respect, the biology of the band-tailed pigeon is similar to that of
the extinct passenger pigeon (Ectopistes migratorius) (Schorger 1955).
Most of BraWl's (1972) study areas are near agricultural areas (e.g. small
grains, orchards). This modifies the environment in that some predictability

�-l/4-

in food supply is attained.
Response to these locally predictable
supplies may explain the high fidelity he found to these areas.

food

Band-tailed pigeons in Colorado and New Mexico retain sexual competency until
mid-August.
The incubation period is 19 days and nestling life is about 20
days (MacGregor and Smith 1955).
Therefore, the birds could easily lay two
clutches of eggs. Also, this bird will sometimes lay eggs in a nest containing a nestling (C. E. Braun, unpublished data).
If this is a common occurrence,
it would considerably shorten two nesting cycles.
Enlarged ovarian follicles are a general indicator of breeding condition.
March and Sadleir (1970) did not find more than one swollen follicle in any
ovary, suggesting that pigeons from British Columbia lay only one egg per
clutch.
Eight percent of the females I examined had two or more enlarged
follicles suggesting that a small percentage of interior pigeons do lay two
eggs.
I do not believe that band-tailed pigeons lay three eggs, although
specimens containing three or more follicles greater than 7.0 rom were recorded
(see Bartelmez 1912).
Neff (1947) and MacGregor and Smith (1955) also
reported clutches of two eggs from interior populations, although these were
not common.
Band-tailed pigeons may attempt to breed three times in years of favorable
environmental conditions, as suggested by fall breeding in at least one
population.
MacGregor and Smith (1955) observed that a pair of band-tailed
pigeons successfully reared three broods in one year.
Gonadal cycles of interior and British Columbia band-tailed pigeons are similar
(see March and Sadleir 1970).
In Britian, the wood pigeon, stock dove
(ColUmba oenas), and rock dove exhibit similar gonadal cycles (Lofts et ale
1966).
Wood pigeons depend on cereal grains and stock doves on weed seeds
during the breeding season (Lofts et ale 1966).
As in these columbids,
band-tailed pigeons undoubtedly respond to food as a proximate factor.
This
is also the case in many other birds (Lofts and Murton 1968).
Other columbids, e. g. the wood pigeon, stock dove, rock dove, and possibly
the mourning dove (Zenaida macroura), show no refractory period like that
seen in many passerines (Cole 1933, Lofts et ale 1966, Lofts et ale 1967a,
Lofts et ale 1967b, Lofts and Murton 1968).
However, another columbid, the
turtle dove (Streptopelia turtur), does show a refractory period (Lofts et ale
1967c).
The experiment involving light and food indicated that light, rather than the
kind or amount of food (acorn group was fed twice daily), was the determining
factor in testicular recrudescence.
This does not discredit the suggestion
that pigeons respond to abundant food in the wild.
Most psycholigical stimuli
necessary for breeding are miSSing in a controlled laboratory environment.
However, the environmental situation dictates the relative importance of each
factor.
Another point brought out by the experiment involving light and food
type is that gonads responded to light per ~,
rather than to more food intake
as a result of long daylength, contradicting the suggestion of Ward (1972).
The fact that band-tailed pigeons are responsive to photoperiods at least
as short as 10 hours (1 January at 350 N) helps to explain the breeding

�-175-

records from Mexico in December and January (Thayer 1909, Lamb 1926).
As
day1ength is always greater than 10 hours in Mexico, breeding presumably
may be attempted if the proximate factors of food and weather are favorable.
The pigeons kept outdoors on natural photoperiods showed a gradual gonadal
regression from September until December (probably until the winter solstice);
when gonads increased rapidly as daylength increased.
Absence of further
growth during January and February may have been the result of overcrowding
in the aviaries.
In any case, after this situation was relieved on 1 March,
gonadal growth resumed.
These data help to explain the ability of the band-tailed pigeon to attempt
unseasonal breeding or to show extended breeding seasons as a response to
environmental conditions (Neff 1947, MacGregor and Smith 1955, Fitzhugh 1970).

SUMMARY
Band-tailed pigeons breed in Colorado and New Mexico from at least May
through August and may attempt two nesting cycles.
Earlier breeding is a
possibility.
Band-tailed pigeons normally lay one egg per clutch, although
small percentage probably lay two eggs.

a

Photoperiod studies of the band-tailed pigeon indicate that the birds are
stimulated by light periods as short as 10 hours a day and that they do not
exhibit a characteristic refractory period.
Gonadal regression begins in
September with a rapid decrease in testes volume.
Further regression takes
place slowly in October and November.
Significant growth, presumably in
response to increasing daylength was observed by 4 January 1972 in birds kept
out of doors, and full breeding capability was reached in March.
The response to photoperiod stimulation and the probable response to other
environmental cues explain the reports of winter nesting and the unusually
long and varied breeding seasons sometimes recorded for this pigeon.
The band-tailed pigeon apparently responds to proximate environmental cues,
particularly mast production, in parts of its range (southwestern New Mexico)
and fall breeding may occur at least occasionally.
This response is not
latitudinal per se but may be influenced by environmental conditions.
This
pattern of breeding in response to environmental cues is also seen in another
southwestern species, the pinon jay. If conditions are poor in a given area,
these pigeons may show nomadic behavior, not unlike that of the extinct
passenger pigeon until finding food adequate to sustain breeding.
This data help explain the unusual
in the band-tailed pigeons.

reproductive

patterns

previously

observed

�-176-

LITERATURE CITED

Abbott, C. G. 1927. Notes on the nesting of the band-tailed pigeon.
Condor 29:121-123.
Bartelmez, G. W. 1912. The bilaterality of the pigeon's egg: a study in
egg organization from the first growth period of the oocyte to the
beginning of cleavage. J. MOrphol. 23:269-330.
Braun, C. E.
pigeons. 1972. Movements and hunting mortality of Colorado band-tailed
Tr. N. Am. Wild!. Nat. Resour. Conf. 37:326-334.

---, and D. E. Benson. 1972. Band-tailed pigeon investigations, breeding
and nesting chronology studies. Colo. Div. of Wildl. Game Res. Rept.
Fed. Aid Proj. W-88-R. October. p. 159-168.
Cole, L. J. 1933. The relation of light periodicity to the reproductive
cycle, migration and distribution of the mourning dove (Zenaidura
macroura carolensis). Auk 50:284-296.
Fitzhugh, L. 1970. Literature review and bibliography of the band-tailed
pigeon of Arizona, Colorado, New Mexico, Utah. Spec. Rept., Arizona
Game and Fish Dept. 53 p.
Glover, F. A. 1953. A nesting study of the band-tailed pigeon (Columba f.
fasciata) in n0rthwestern California. Calif. Fish and Game 39:397-407.
Horwitz, W. 1960. Official methods of analysis.
Agr. Chemists. Washington, D. C. 832 p.

9th Ed. Assoc. of Off.

Houston, D. B. 1963. A contribution to the ecology of the band-tailed
pigeon, Columba fasciata, Say. M. A. Thesis. Univ. of Wyo., Laramie.
74 p.
Lamb, C. C.

1926.

The Viosca pigeon.

Condor 28:262-263.

Ligon, J. D.
Mexico.

1971. Late summer-autumnal breeding of the pinon jay in New
Condor 73:147-153.

Lofts, B., R. K. Murton, and N. J. Westwood. 1966. Gonadal cycles and the
evolution of breeding seasons in British Columbidae. J. Zool., London
150:249-272.

---,

, and
1967a. Experimental demonstration of a postnuptial refractory period in the turtle dove, Streptopelia turtur.
Ibis 109:352-358.

---,

, and
1967b. Photoresponses of the woodpigeon, Columba
palumbus, in relation to the breeding season. Ibis 109:338-351.

�-177-

Lofts, B., R. K. Murton, and N. J. Westwood. 1967c. Interspecific differences
in photosensitivity between three closely related species of pigeons.
J. Zool., London 151:17-25.
Lofts, B., and R. K. Murton. 1968. Photoperiodic and physiological adaptions regulating avian breeding cycles and their ecological significance.
J. Zool., London 155:327-394.
MacGregor, W. G., and W. M. Smith. 1955. Nesting and production of the
band-tailed pigeon in California. Calif. Fish and Game 41:315-326.
March, G. L., and R.M.F.S. Sadleir •. 1970. Studies on the band-tailed pigeon
(Columba fasciata) in British Columbia. 1. Seasonal changes in gonadal
development and crop gland activity. Can. J. Zool. 48:1353-1357.
Meyer, R. K., C. Kabat, and I. o. Buss. 1947. Early involuntary changes in
the post-ovulatory follicles of the ring-neck pheasant. J. Wildl. Mgmt.
11:43-49.
Murton, R. K., and A. J. Isaacson. 1962. The functional basis of some
behavior in the wood-pigeon (Columba palumb~s). Ibis 104:503-521.
Neff, J. A. 1947. Habits" food, and economic status of the band-tailed
pigeon. N. Am. Fauna 58:1-76.
Salt, G. W. 1953. An ecologic analysis of thr~e California avifaunas.
Condor 55:258-273.
Schorger, A. W.

1955.

The ~assenger pigeon.

Univ. Okla. Press.

Norman. 424 p.

Silovsky, G. D., H. M. Wight, L. H. Sisson, T. L. Fox, and S. W. Harris. 1968.
Methods for determining age in band-tailed pigeons. J. Wildl. Mgmt.
32 :421-424.
Smith, W. A. 1968.
Game 54 :4-16.

The band-tailed

pigeon in California.

Calif. Fish and

Stephens, F.

1913.· Early nesting of the band-tailed pigeon.

Swarth, H. S.

1900.

Avifauna of a 100-acre ranch.

Thayer, J. E.

1909.

Letter to the editor.

Vorhies, C. T. 1928.
Condor 30:253.

Condor 15 :129.

Condor 2 :14-16.

Condor 11:142-143.

Band-tailed pigeon nesting in Arizona in September.

Ward, P. 1972. An appeal to physiologists
Ibis 114 :275.

reporting photoperiod experiments.

Zeigler, D. L. 1971. Crop-milk cycles in band-tailed pigeons and losses
of squabs due to hunting pigeons in September. M. S. Thesis. Oregon
State Univ., Corvallis. 48 p.
Prepared by

/

Ralph J. Gutierrez

Approved by _~t4-=-=V:..:..-.· ~r.==-=-. -+~-c.:.....:=~~
Clai t E. Braim
Wildlife Researcher

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                  <text>-1-

January, 1974

JOB FINAL REPORT

State of __

-=C-=O=LO.;:;,;RA=D::..;O~_

Project No.

.W-4l-R-23

Work Plan No.

1

Bighorn Sheep and Mountain Goat Investigations
Job No.

12

Job Title:

Location and Distribution of Bighorn Sheep Herds in Colorado

Period Covered:

June 1, 1972 through May 31, 1973

Personnel:

George D. Bear and George W. Jones.

ABSTRACT

A final report has now been completed, published and distributed during this segment,
in the Game Research Report format under a special cover and binder to provide usable
sources of information for wildlife managers, under the title "History and Distribution
of Bighorn Sheep in Colorado".
.

Prepared by

r

-_--_-_/._;'_~-~~~&lt;~-!---.-.-~-.--

George D. Bear
Wildlife Researcher

r ~'"

_

, •.. ,-,

��January,

-3-

JOB PROGRESS

State of
Project

1974

REPORT

.:;.CO.:::.;L::;.O::;,;RAD=:...:O:.-.
_
No.

Work Plan No.
Job Title

Bighorn

1

Job No.

Evaluation

Period Covered:
Personnel:

W-4l-R-23

of the Nutritional

Sheep and Mountain

Requirements

Goat Investigations

16
of Bighorn

Sheep

June 1, 1972 through May 31, 1973

Gene G. Schoonveld - Field Collections, bighorn sheep food-habits.
Robert E. Keiss and William J. Adrian - laboratory analyses.

ABSTRACT

This investigation concerns nutrition of bighorn sheep and the role of nutrition
to the overall welfare of this animal species.
These past two segments have centered
primarily on making field collections of forages and bighorn sheep fecal samples from
the Buffalo Peaks, Pikes Peak and Trickle Mountain ranges, determining what plants are
eaten by sheep, and testing laboratory techniques for measuring nutritional quality of
forages and nutrients eliminated in feces.
Food-habits data obtained during this second segment do not show extreme fluctuations
in plants eaten between months as previously reported and is believed to better estimate
relative proportions of forages actually eaten.
Grasses and sedges were predominantly
eaten throughout the year with increased usage of browse species occurring during
fall and early winter.
Only one forb species, Arenaria fendleri was eaten by sheep
in substantial quantities.
Testing laboratory techniques for measuring nutritional quality of forages as well as
analyses of fecal samples for nutrients eliminated has been accomplished.
The results
of utilizing these techniques on forages and fecal samples are also contained in this
report.

��-5-

EVALUATION OF THE
NUTRITIONAL REQUIREMENTS OF BIGHORN SHEEP

•
. !J'

Robert E. Keiss
Gene G. Schoonveld

P. S. OBJECTIVE
To develop methods:, (a) for the evaluation of nutritional requirements
of bighorn sheep under natural conditions, and (b) to measure the adequacy
of selected ranges to meet these requirements.
SEGMENT OBJECTIVES
1.

To estimate the nutritional level o f forage species utilized by bighorn sheep.

2.

To evaluate if the nutritional level of selected bighorn sheep ranges
is adequate to meet nutrient requirements of bighorn sheep at all
seasons of the year.

METHODS AND MATERIALS

Nutritional Quality of Forage
Species Used by Bighorn Sheep
Sample Collections and Food-Habits Analysis
Procedures for collecting forages and bighorn sheep fecal samples and microtechniques used in determining food-habits were previously described
(Schoonveld 1973) .. Fecal samples collected during this·segmentwere also
analyzed for numbers of lungworm larvae. Results of these analyses are
reported under the contract segment, "Spontaneous Diseases in Bighorn Sheep".

Laboratory Methods
General
•

In the early years chemists attempted to define the nutritive quality of
feeds (Thaer 1809; Boussingault 1843; Henneberg and Stohmann 1860) and
their chemical methods were ultimately adopted and developed into the total
digestible nutrient (TDN) system of feed analysis as proposed by Henry and

�-6-

Morrison (1910). The Weendemethod
of partitioning forages is based
upon the TDN system (crude protein, crude fat, crude fiber, and nitrogen free extract) but has several major flaws and assumptions which
are very questionable by today's standards of feed analysis.
Other modifications and improvements have been made by a number of workers
in recent years (Crampton and Maynard 1938; Van Soest 1964; Colburn and
Evans 1967; Van Soest and Wine 1967). Fonnesbeck and Harris (1970a, 1970b,
1971) and Harris (1970) have developed a procedure for examining the cell
wall constituents of all types of feeds which have been adopted by our
laboratory and were used throughout this experiment. The plant I:!ellis
basically separated into two main components, the cell walls and the cell
contents. The cell contents.include the majority of the nutritive matter
that is digested by enzymatic action within the digestive system and subsequently absorbed and utilized by the animal. This includes both partially
nutritive matter and non-nutritive matter. Partially nutritive matter includes cell wall carbohydrates, cellulose and hemicellulose.
These
chemically complex compounds are partially broken down by the action of
bacterial enzymes within the gastro-intestinal tract and the total digestibility of the cellulose and hemicellulose can be combined and reported as
holocellulose as the differences for most animals is not significantly
different to report separately. The non-nutritive matter within the cell
walls includes lignin and acid insoluble ash. These constituents have no
known nutritive value.
The system used for analysis of the nutritive value of feeds used in this
experiment requires the determinations of protein, ether extract, ash, cell
walls, cellulose, hemicellulose, lignin, acid insoluble ash, soluble ash,
and soluble carbohydrate.
Mineral analysis for calcium, phosphorous, magnesium, manganese, zinc,
copper, iron, potassium were made on selected forages as a part of this
study •

.Protein
The micro-Kjeldahl method for the determination of crude protein (total
nitrogen x 6.25) was used. Two hundred fifty milligrams of plant material
was placed in a 100 ml Kjeldahl flask with 1 gm of catalytic mixture and
3 m1 of concentrated H2S04. The material was predigested overnight then
placed upon a micro-Kjeldahl digester and heated until the sample was
completely clear. After cooling, the digested sample was transferred to
a micro-Kjeldahldistillation
chamber. Ten ml of 12 N.NaOH is added
and the sample distilled with steam into a boric acid solution containing
2 drops each of bromocresol green and 2,4, dinitro-phenol indicator
solutions. The boric acid solution was titrated with 0.05715 N. H2S04.

,

.,.-

�-7-

Calculation:
Percent Protein = m1. titrant x 2
Reagents:
1. Catalytic Mixture
100gm Cu~04
2. Boric Acid
3.·Bromocreso1 Green Indicator
4.; 2,4 Dinitro-pheno1 Indicator
,5."12 N. NaOH
~. 0.05715 N. H2S04

250 gm Na2S04

Ether Extract
A one gram (ove.ndry) sample is weighed into a previously tared, fat-free
extraction thimble. iThe thimble was placed in aSaxh1et fat extraction
apparatus and extracted for eight hours with anhydrous petro1etnn ether.'
After the extraction period the thimbles were removed, allowed to dry
overnight, cooled in a desiccator and weighed. The loss in weight of the
sample was calculated as percent ether extract on a: dry weight basis •.

Ash
A one gram (oven dry) sample was weighed into a previously tared crucible
and placed into a cold muffle furnace. The heat was allowed to buildup
slowly to 5500 C artd'held overnight. The following day the heat was shut
off, allowiIlg the~rucibles to cool. After the crucibles were cooled to
approximately 2000'C they were transferred to a desiccator where they were
allowed to cool to room temperature'and weighed. Percent ash was calculated on a dry weight basis. The ash was then dissolved in 2 m1 of concentratedNH03 ov'er a steam bath and quantitatively transferred to a 50
ml vo1tnnetric flask, made to vo1tnne, and the solution kept for mineral
analysis.

Cell Walls
.-;.:

•

19ram (oven dry) .samp1e was added to a 600m1 Berze1ius beaker and 75 m1
of citrate buffer and 1 m1 deca LLn added. .The beakers were placed on a
refluxing apparatus consisting of six individually controlled hot plates
and 500 m1 round bottom water cooled flasks on top of the beakers. The
samples were brought to a boil and allowed to reflux for 30 minutes from

�-8-

the onset of boiling. The beakers were removed, cooled and 25 ml of
pepsin solution added. The beakers were placed in a water bath at 450
C.Since
we had no means to keep the flasks in constant motion as
recommended, we allowed the digestion to proceed for approximately 48
hours with occasional stirring. After the pepsin digestion the liquids
were removed from the beakers and the residue rinsed with hot water,
followed by a thorough acetone washing. After the acetone washing,
100 ml of detergent solution was added and the beakers refluxed for 60
minutes on the refluxing apparatus. The detergent samples are then
poured into a previously tared filter crucible while drawing off the
liquid with suction. The cell walls were then thoroughly washed with
large volumes of hot water, followed by thorough rinsing with acetone.
The filter crucibles were dried at 1050 C overnight, placed in a
desiccator to cool, and weighed.
Calculation:
Percent Cell Walls

Weight of Cell Walls
Weight of Sample

X 100

Reagents:
1.

42 gm
Citric Acid
10 gm
Tween 40
1,000 ml
Water
(Adjust pH to 3.0 ,.nthNH40H)
2. Pepsin Solution
Pepsin
'40 gm
Citric Acid
40 gm
Water
1,000 ml
(Make fresh as needed - 1 gm pepsin and 1 gm
citric acid per 25 ml - which is required
for each sample)
3. Detergent Solution
Citric Acid
42 gm
Dodecyl Sodium
Sulfate
20 gm
Water
1,000 ml
(Adjust to pH 3.0 with NH40H)
4. Decalin (Decahydronaphthalene)
5. Acetone
Citrate Buffer

Hemicellulose
The residue from the cell wall determination was removed from the filter
crucible and placed in the Berzelius beaker, taking great care to remove
all the material. Seventy-five ml of 8 percent H2S04 was added to the
beaker and they were refluxed on the refluxing apparatus for 1 hour.
After refluxing, each sample was washed back into the same filter crucible
from which it was removed, the residue washed in large volumes of hot,
water followed by a thorough acetone rinse. The filter crucibles were
placed in the drying oven overnight, removed to a desiccator until cool
and weighed. Hemicellulose was computed as the loss of weight from the
cell wall residue.

�-9':'

.Calctilation :

Wt. Cell Wall Residue Wt. Extracted·Cell Wall Residue ·X 100
Sample Weight

% Hemicellulose

Reagents:
1.

8% Sulfuric Acid (Specific gravity of 1.052 at 20o C)

Lignin and Acid Insoluble Ash
The residue from the hemicellulose determination is digested with 72%
H2S04 by placing the filter crucible into a beaker containing the acid.
The acid is allowed to soak into the residue from the bottom through the
fritted disc so that the residue is wetted from the bottom up. The residue
was left in the acid for 3 hours after which the acid was filtered off,
followed by a thorough washing with large volumes of hot water. It is
important ..
to remove all traces of acid from the residue and filter crucible.
The residue was then dried overnight, cooled in a desiccator and weighed.
The filter crucibles were then placed in the muffle furnace at 5000 Covernight, removed, cooled in a desiccator and weighed.
Calculation:
% Lignin = Wt.Residue

BeforeAshing":" wt. Residue After Ashing X 100
Sample Weight

% Acid Insoluble Ash

Wt. Residue After Ashing - Wt. Tared Crucible
Sample Weight

Reagents:
1.

72% Sulfuric ACid (Specific gravity of 1.63 at 200 C)

Celluiose, by Ca Lcu'Lat.Lon:
Calculation:
% Cellulose

% GellWalls

- (% Hemicellulose + % Acid Insoluble Ash)

Soluble Ash, by Calculation
Calculation:
% Soluble Ash

% Ash - % Acid Insoluble Ash

Cell Contents, by Calculation
Calculation:
% Cell Contents = 100 -% Cell Walls

XIOO

�-10-

Soluble Carbohydrate, by Calculation
Calculation:

% Soluble Carbohydrate =% Cell Contents - (% Protein + % Soluble
Ash + % Ether Extract)
Summary of Procedures:
The cellular components are chemically separated and evaluated by these
methods as described by Fonnesbeckand Harris (1970a, 1970b, 1971) and
Harris (1970) with a few modifications to better adapt the procedures to
our particular laboratory and needs. The basic data produced by these
procedures areas follows:
% Cell Content
% Protein
% Lipid- Ether Extract
% Soluble Ash
% Soluble Carbohydrate
% Cell Walls
% Hemicellulose ) % Holocellulose
)
.
% Ce11u 1ose
% Acid Insoluble Ash
% Lignin

Mineral Analysis
After dry ashing a one gram sample for percent ash determination, the ash
was dissolved in 2 ml concentrated HN03 and transferred to a 50ml volumetric
flask. This solution was the stock solution for running mineral analysis.
Phosphorous was determined by the molybdophosphoric acid spectrophotometrically
using a B &amp; L Spectronic 20. All other minerals were determined using a
Perkin-Elmer 303 Atomic Absorption Spectrophotometer using standard procedures~
The following minerals were analysed for, in selected samples:
..,.

'

Phosphorous
Calcium
Magnesium
Manganese

Zinc}
Copper
Iron
Pota,ssium

RESULTS AND DISCUSSION
Nutritional Quality of Forages Used by Bighorn Sheep
Food-Habits
Residues of plant species identified in fecal samples were similar to those

�-11-

•
;'

previously reported (Schoonveld 1973) however, estimated relative proportions of major forage classes (grasses and sedges, browse, forbs)
do not show extreme monthly fluctuation (Fig. lA). Food-habits data
obtained during this segment is more closely substantiated by field
observations of what sheep were seen to be using and is believed to
better represent relative proportions of forages used. Improved data
accuracy is probably the result of increased knowledge of sheep using
the range, better field collection techniques, and more experience on
the part of the technician in preparing microscope slides, identifying
plant residues and estimating relative proportions· of plants eaten.
A comparison of major forages used (species consistently identified in
substantial amounts &gt; 5 percent of the diet) among the three study areas
is given in Table lA. Complete lists of all grasses and sedges, trees
and shrubs, and forbs identified in fecal samples collected June 1971
May 1973 are listed in Tables 2A through 4A. Grasses and sedges were
predominantly eaten throughout the year with increased utilization of
browse species occurring during fall and early winter (Fig. lA). Only
one forb species, Arenaria fendleri was used by Pikes Peak and Buffalo
Peaks sheep in substantial amounts (Table lA).

Digestion Coefficients, Total Digestible Nutrients (TDN) and Nutritive Ratios

•

Accurate data concerning relative proportions of individual plant species
comprising sheep diets are essential in determining digestion coefficients,
TDN,· and nutritive ratios of the diet. Previous studies (Zyznar and Urness
1969, Stewart 1967, Schoonveld 1973) suggest substantial bias exists in
use of fecal microanalysis techniques for estimating relative proportions
of plants ingested. Recently, an exacting investigation on validity
of the technique (Dearden 1973) found that differential digestion does influence percent relative proportions of plant residues discernible in feces
and correction factors must be used in making diet composition estimates.
He found relative proportions of grasses and grasslike forages were consistently overestimated while forbs and shrubs were underestimated.
Determining correction factors for all major forages used by sheep would
involve extensive feeding trials, preferably using captive bighorn sheep,
which is not feasible at. this time, nor included in the scope of this
investigation. The determination of digestion coefficients, TDN, nutritive
ratios and digestible energy of sheep diets cannot be done with a reasonable
degree of accuracy and is therefore deleted from this study •

•

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I

/

I
I

I

.

.i:

I
I
I

I
I

-

Iii

I

.•.....•...•
•......
. I

OCT

/'"

'"

"'x'"

I
i

I
I

"

",'
,
,"
,
.
-----, --- ..---a

5 O'

•....

"

,~/

60

'"
/

I

/'

-,

,

II

/

'" '"

I

J

~/ /

..;''

/{"

J

/

,.......;'
..

.

I

/

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U.

I

J

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J

,

••••

,~X

,.

•.. 80

I·

-~

-

NOV

x - PlKfS PEAK
., - 'BUFFALO .
II. - TRICKLE

I
.: ••••••
D

......

DEe

.'

'i'

;

JAN

".

".'"
I

';

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.FEBMAR

,

I

APR

i

MAY

Fig. lA. Percent grasses and grasslike forages identified in diets of bighorn sheep using the Trickle Mountain,
Buffalo Peaks and Pikes Peak ranges, June 1972 - May 197'3•. Estimates of percent relative density were made
using fecal sampl~ micro-techniques.

'"

"

•

..

.J.

.-.,

�-13-

Table lA. Major forage species used by bighorn sheep occupying the Buffalo
Peaks, Pikes Peak and Trickle Mountain ranges June 1971-May 1973. Plant
species were identified and percent relative proportions estimated using
fecal sample micro-techniques •.

Species

Percent of Diet (Range)
Buffalo Peaks
Pikes Peak
Trickle Mtn.

Grasses and Grasslikes
Agropyron spp.

T-7

T-IO

Bouteloua gracilis

T-9 11

T-31./

Bromus spp.

T-ll

Carex spp.

3-54

1-30

T-14
8-92

Danthonia parryi

4-19
T-7

Deschampsia caespitosa

T-17

T-4

Festuca spp.

T-67

T-58

Muh1enbergia spp.

T-6

T-56

Oxyzopsis hymenoides

T-5

T-ll

Paa spp.

2-11

T-15

Sporobo1us cryptandus

T-53

4-15
T-8

Stipa spp.

T-ll

T-5

T-17

T-12

T-2

Artemisia frigida

T-4 !/

T':S1./

T-15

Cercocarpus montanus

T-12 1./

T-7 1.1

T-14

Forbs
Arenaria fend1eri
Shrubs and Trees

Eurotia lanata

T-16
T 1/

Ho1odiscus dumosus

••

T-42

Pinus spp.

T-7

Shepherdia canadensis

T-1~

Salix spp.

T-28

T-35

'I-I 0

Yucca spp.

T-61/

T !/

T-16

T-29

1/
- Indicates sheep are moving and foraging between alpine and sub-a1plne ranges •

.. ",.:--

�-14Table 2A. Grass and grasslike forages identified in bighorn sheep fecal
samples collected June 1971 - May 1973·from the Buffalo Peaks, Pikes Peak
and Trickle Mountain ranges ...

Buffalo Peaks

Pikes Peak

Agropyron spp.

X

X

X

Agrostis variabilis

X

X

X

X

X

Species

Aristida spp.

Trickle Mtn.

Bouteloua gracilis

X

X

X

Bromus spp.

X

X

X

Calamagrostis canadensis

X

X

X

Carex spp.

X

X

X

Carex nigricans

X

X

X

X

X

Cyperaceae spp.
Danthonia parryi

X

X

X

Deschampsia caespitosa

X

X

X

X

X

X

Festuca idahoensis
F. thurberi

X

X

Hordeum jubatum

X

X

Juncus spp.

X

X

X

X

X

Koe1eria cristata
Kobresia be1lardi

X

X

X

Muh1enbergia spp.

X

X

X

Oryzopsishymenoides

X

X

X

X

Phleum alpinum
Poa spp.

X

X

X

Sporobolus cr:¥:ptandus

X

X

X

Stipa spp ,

X

X

X

Trisetum spicatum

X

X

~

�-15-

Table 3A. Forbs identified in bighorn sheep fecal samples collected June
1971 - May 1973 from the Buffalo Peaks, Pikes Peak and Trickle Mountain
ranges.
;&lt;iI'
,

'\

;;

Species

Buffalo Peaks

Pikes Peak

Achillea 1antilosa

x

Allium spp.
Anemone spp.

X

X

Amaranthus spp.

X
X

Arenaria fend1eri

X

X

X

Astragalus spp.

x,

X

X

Calochortus nutta11i

'.

X

Castilleja spp ,.

X

X

Chrysopsis vil10sa

X

X

Cryptantha spp.

X

X

Delphinium spp.

X

Draba spp.

x

Erigeron spp.

X

X

X

Erigeronum spp.

X

X

X

Erysimum aspertim
Euphorbia 1athyris

X
X

X

Geum'turbinatum
Hymenoxys spp ,

X

Lesquerel1a spp.

X

Lupinus spp.

X

X

X

X

X
X

X

Ranunculus spp.
Senecio spp.

X

Sphaeralcea coccinea

X

Townsendia sericea
Trifolium parryi

X
X

Po1emonium spp.
Potentil1a spp.

X
X

Opun tia spp ,
Oxytropis hymenoides

X
X

Mertensia1anceo1ata

•

Trickle Mtn.

X

X

X

X

X

X
X

X

X

�-16-

Table 4A. Shrubs and trees identified in bighorn sheep fecal samples
collected June 1971 - May 1973 from the Buffalo Peaks, Pikes Peak and
Trickle Mountain ranges ..

Species

Buffalo Peaks

Pikes Peak

Trickle Mtn.

Artemisia frigida

x

x

x

A. tridentata

x

x

x

Atriplex canescens

x

x

Berberis repens

x

x

x

x

x

x

Cercocarpus montanus

x

Chrysothamnus spp.
Eurotia lanata

x

x

x

Holodiscus dumosus

x

x

x

Juniperus spp.

x

x

x

Pinus spp.

x

x

x

Potentilla fruiticosa

x

x

Rosa spp.

·x

x

x

Shepherdia canadensis

x

x

x

Salix spp.

x

x

x

Yucca spp.

x

x

x

. ." ....

�-17-

Evaluation of Selected Forages by Chemical Procedures
In order to test the analytical procedures for the evaluation of feeds
and feces by the methods of Fonnesbeck and Harris (1970a, 1970b, 1971)
as well as to become proficient in the use of these procedures an
experiment was designed using a series of forage samples. For this work
grass samples collected at two seasons of the year (Summer-Winter) were
selected for testing. These included grasses of the genus Poa, Bouteloua
Festuca, Muhlenbergia, and KObresia. Eight replicate samples collected
in summer and eight replicate samples collected in winter (with the exception of Bouteloua where no winter collections were available) were analyzed
giving a total of 72 grass samples.
All samples were analyzed to determine the percent cell content (% protein,
% ether extract, % soluble ash, % soluble carbohydrate), percent cell walls
(% insoluble ash, % lignin, % cellulose, % hemicellulose).
The results of
these determinations are shown in Tables 1,2,3,4,5,6,7,8, and 9.
All grass samples were analyzed for mineral content using the methods previously described. All samples were analyzed for calcium, phosphorous,
manganese, magnesium, copper, iron, potassium, and zinc. The results of
these analyses are shown in Tables 10,11,12,13,14,15,16,17, and 18.
Sample numbers assigned to forage samples in the cell content-cell wall
analysis correspond to those assigned for mineral analysis.

Evaluate Nutritional Level of Ranges in
Furnishing Nutrient Requirements of Bighorn Sheep
Range evaluations are dependerit upon comparisons of nutritional data or
"qua1ityU of forages used by sheep on the three study areas. These analyses
are continuing and complete evaluations cannot be done until all samples and
data have been analyzed and comparisons made •

••

�Table 1. Poa (swnmer) - chemical partitioning of the nutritive constituents by the methods of
Fonnesbeck and Harris (1970a, 1970b, 1971).
Cell Content
Soluble
Ether
Extract
Ash

(%)

Percent
of
Total

(%)

(%)

(%)

(%)

2.n
1.59
2.32
3.21
1.89
2.26
2.42
2.92

12.48
14.61
20.16
5.48
15.15
11.19
22.72
20.82

67.98
68·.15
61.63
70.97
68.03
n.57
60.80
61.90

2.41
2.84
2.10
2.96
2.67
2.83
1.98
2.79

5.51
6.33
6.30
6.13
5.94
5.81
5.72
5.63

34.71
29.96
26.19
33.86
31. 89
34.77
24.29
26.01

25.35
29.02
26.77
28.02
27 ~53
29.16
28.81
27.47

2.42

15.33

66.47

2.57

5.92

30.21

27.77

Sample
Number

Percent
of
Total

Protein

(%)

(%)

(%)

1
2
3
4
5
6
7
8

32.02
31.85
38.64
29.03
31.97
27.43
39.20
38.10

11.60
10.80
11.60
15.60
10.80
9.60
10.00
10.40

5.22
4.85
4.56
4.74
4.13
4.38
4.06
3.96

-

33.53

11.30

4.49

x

Cell Walls
HemiLignin Cellulose

Insoluble
Ash

Soluble
Carbohydrate

Cellulose

I

t-'
00
I

Table 2. Poa (Winter) - chemical- partitioning of the nutritive constituents by the methods of
Fonnesbeck and Harris (1970a, 1970b, 1971).
Cell Content
Soluble
Ether
Ash
Extract

Cell Walls
Hemi.Lignin Cellulose

Insoluble
Ash

(%)

(%)

(%)

(%)

12.50
14.31
12.40
13.84

35.93
34.46
33.71
34.43
34.58
33.48
35.73
33.25
34.45

Sample
Number

of
Total

Protein

(%)

(%)

(%)

(%)

Percent
of
Total

9
10
11
12
13
14
15
16

14.64
16.71
18.10
17.65
16.28
18.71
17.22
16.19

4.80
6.60
6.20
6.80
6.40
6.60
6.60
6.80

5.56
2.42
3.10
4.30
3.92
4.17
3.17
2.03

3.90
2.07
2.69
2.52
2.02
2.42
3.23
2.75

.38
5.62
6.11
4.03
3.94
5.52
4.22
4.61

85.36
83.29
81.90
82.35
83.72
81.29
82.78
83.81

2.25
3.15
3.14
2.22
3.66
3.10
3.32
2.52

12.64
12.41
12.68

34.68
31.37
32.65
31.86
31. 76
32.07
31.32
34.86

x

16.94

6.35

3.58

2.70

4.30

83.06

2.92

13.06

32.57

Per cent :

(I.

If

Soluble
Carbohydrate

.-1

.~

13. t:

j

.~

Cellulose

�.,

~

•

"

.~

"

Table 3. Kobresia (summer) - chemical partitioning of the nutritive constituents by the methods of
Fonnesbeck and Harris. (1970a, 1970b, 1971).
CellCorttent
Ether
Soluble
Extract
Ash

Sample
Number

Percent
of
Total

Protein

(%)

(%).

(%)

17
18
19
20
21
22
23
24

31. 76
29.92
31.47
27.86 .
31.51
30.95
37.00
35.99

14.80
10.40
11.20
9.60
10.40
10.80
10.40
10.40

4.34
5.9l
4.70
4.59
4.85
4.52
4.09
4.83

-

32.06

11.00

4.73

x

Soluble
Carbohydrate

Cell Walls
HemiLignin Cellulose

Insoluble
Ash

(%)

Percent
of
Total

(%)

(%)

(%)

(%)

2.84
2.74
3.23
2.96
2.71
3.34
3.33
4.02

9.78
10.87
12.34
10.71
13.55
12.29
19.18
16.74

68.24
70.80
68.53
72.14
68.49
69.05
63.00
64.01

3.19
3.26
2.86
3.42
3.38
3.30
3.20
3.27

9.03
9.53
8.27
7.64
8.64
7.69
8.40
7.55

28.88
30.59
29.76
34.77
31.72
31.28
26.96
29.23

27.14
26.70
27.64
26.31
24.75
26.78
24.80
23.96

3.15

13.18

67.94

3.24

8.34

30.40

26.01

Cellulose

I

I-'

1.0
I

Table 4. Kobresia (winter) - chemical partitioning of the nutritive constituents by the methods of Fonnesbeck
and Harris (1970a, 1970b, 1971).
Cell Content
Soluble
Ether
Ash
Extract

Soluble
Carbohydrate

Cell Walls
HemiLignin Cellulose

Percent
of
Total

Insoluble
Ash

(%)

(%)

(%)

(%)

Sample
Number

Percent
of
Total

Protein

(%)

(%)

(%)

(%)

25
26
27
28
29
30
31
32

23.00
22.68
24.62
22.39
21.06
21.88
22.84
24.51

7.60
6.40
6.80
7..
20
6.20
7.20
7.60
9.00

3.52
3.37
1.9l
3.07
2.39
2.38
2.12
3.06

3.15
3.54
3.75
3.44
3.04
3.11
2.82
3.28

8.73
9.37
12.16
8.68
9.43
9.19
10.30
9.17

77 .00
77.32
75.38
77.61
78.94
78.12
77.16
75.49

3.32
3.36
2.80
2.96
3.62
3.95
3.52
2.96

11.86
12.15
13.11
11.92
12.67
11.79
14.06
14.45

30.28
30.21
29.65
29.79
30.47
29.86
28.86
26.•
91

31.54
30.60
29.82
32.94
32.18
32.52
30.72
31.17

-

22.87

7.25

2.73

3.27

9.63

77.13

3.31

12.75

29.50

31.44

x

Cellulose

�iI

Table 5. Fes tuca (swnmer) -chemical partitioning of the nutritive constituents by the methods of
Fonnesbeck and Harris (1970a, 1970b, 1971).
Cell Content
Ether
Soluble
Extract
Ash

Soluble
Carbohydrate

Cell Walls
HemiLignin Cellulose

Insoluble
Ash

(%)

Percent
of
Total

(%)

(%)

(%)

(%)

4.74
4.27
4.71
3.95
2.25
3.76
3.98
4.14

6.05
7.06
9.35
11.86
14.62
8.07
5.74
10.46

76.09
71.91
65.63
72.56
67.29
69.91
66.04
69.27

9.47
6.48
5.03
8.80
7.37
6.77
5.88
7.68

7.27
8;24
7.00
6.58
7.07
7.83
7.07
7.21

25.20
26.29
25.80
22.38
27.61
28.68
23.67
25.16

34.15
30.90
27.80
34.80
25.24
26.63
29.42
29.22

3.98

9.15

69.84

7.19

7.28

25.60

29.77

Sample
Number

Percent
of
Total

Protein

(%)

(%)

(%)

33
34
35
36
31
38
39
40

23.91
28.09
34.37
27.44
32.71
30.09
33.96
30.73

8.40
11.60
14.40
7.20
10.40
13.20
15.20
11.20

4.72
5.16
5.91
4.43
5.44
5.06
9.04
4.93

x

30.16

11. 45

5.59

Cellulose

, I

-tv
0
I

of the nutritive constituents by the methods of
Table 6. Festuca (winter) - chemical partition:i,.ng
Fonneab eck and Harris (1970a, 1970b, 1971).'

Sample
Nuinber

Percent
of
Total

41
42
43
44
45
46
47
48

19.14
20.98
20.02
22.68
20.97
21.02
20.94
24.42

x

21.27

~

'T'

Cell Con tent
Soluble
Ether
Ash
Extract

(%)

(%)

(%)

Percent
of
Total

3~80
3.40
4.00
5.40
5.60
5.00
6.40
7.60

2.55
2.30
3.11
4.17
3.35
4.48
4.39
5.94

3.92
4;24
3.23
3.11
2.89
3.18
2.18
3.14

8.89
11.04
9.68
10.00
9.13
8.36
7.97
7.74

80.86
79.02
79.98
77 .32
79.03
78.98
79.06
75.58

.8.98
6.01
7.80
7.52
6.63
7~78
5.96
5.86

'5.15

3~79

3.24

9.10

78.73

7.07

Protein

(%)

Soluble
Carbohydrate

Insoluble
Ash

Cell \val1s
HemiLignin Cellulose

(%)

Cellulose

(%)

(%)

.(%)

9.26
9.63
9.30
10.54
9.52
11.03
11.66
12.41

27.89
27.96
27.73
26.68
28.25
27.44
28.99
26.90

34.73
35.42
35.15
32.58
34.63
32.73
32.45
30.41

10.42

27.73

33.51

-

.'

~

",

.

~

)"

�•

l.

'jI

Table 7. Muhlertbergia (summer) - chemical partitioning
Fonnesbeck and Harris (1970a, 1970b, 1971).

Sample
Number

Percent
of
Total

Protein

(%)

Cell Content
Soluble
Ether
Ash
Ex.tract

••

"
of the nutritive

(%)

(%)

Percent
of
Total

4.26
3.99
4.20
3.69
4.08
4.02
3.55
4.00

6.24
6.36
9.68
7.87
6.18
6.75
8.60
8.70

77.•91
70.20
71.23
76.99
75.30
73.44
72.30
72.86

'3.97.

7.55

(%)

Soluble
Carbohydrate

~.

!

constituents by the methods of
Cell ~valls
HemiLignin Cellulose

Insoluble
Ash

(%)

(%)

(%)

(%)

7.06
4.07
4.20
5.53
4.30 .
4.36
4.21
3.26

9.76
8.62
8.95
9.89
9.83
9.81
10.30
9.67

30.77
29.03
29.05
33.40
31.53
32.57
29.93
30.66

30.32
28.48
29.03
28.17
29.64
26.70
27.86
29.27

30.87.

28.68

Cellulose

. ;~,-:
','

49
50
51
52
53
54
55
56

-.
x

22.09
29.80
28.77
23~01
24.70
26.56
27.70
27.14
.'

-.

7.00
13.00
10.40
7.80
10.40
11.00
11.40
11.40

4.59 .:
6.45
4.49
3.65
4.04
4.79
4.15
3.04

..
'

26.22'

I

N

I-' .
I

'.

;

Ta.ble a.Muhlertberg-ia.(W1nteJ:)-chemica.1 partitioni~g
Fonnesbeck and Harris (1970a, 1970b, 1971).
.

Sample
Number

Percent
of
Total

Protein

(%)

Cell Content
Soluble
Ether
Ash
Extract

(%)

64

2.13
4.00
.··2.33
2.40
'1.65'
4.80
5.60'
2'~21·
2~16'
18037
S.2.b -.
1. 81
.6;00 ..
18.2.1
1.-96
.16.~88'.' 5.40
3.04
6.60
20.95

x

18.35

57
.58
···.·.59·

··~··60
61'
6.2
63

16.42
17.04
'18.;76"
"'20~17

5.00

2.16

of i:he'nutritiv~ cortstit:uentsby the methods of
.'
Cell Walls
HemiLignin Cellulose

Insoluble
Ash

(%)

(%)

(%)

(%)

(%)

(%)

Percent
of
Total

3.41
3.56
'3.43
2.91
2.52
2.49
2.37
3.20

6.88
8.75
8~88
9~45
8.49
7.91
7.15
8.11

83.58
82.96
81.24
79.83
81. 63
81. 79
83.12
79.05

6.27
7.09
4.76
4.97
4.44
4.12
4.64
4.09

10.27
9.84
10.34
10.34
11.22
11. 77
10.91
10.42

38.48
35.56
33.31
33.29
34.95
35.17
34.65
34.43

28.56
30.47
32.83
31.23
31.02
30.73
32.96
30.11

8.20

81.65

5.05

10.64

34.98

30.99

2.99

Soluble
Carbohydrate

'.'

Cellulose

�Table 9. Bouteloua (summer) - chemical partitioning of the nutritive constituents by the methods of
.Fontlesbeck&gt;andHarris (1970a, 1970b, 1971).

Sample
Number

Percent.
of
Protein
(%),
Total

Cell Content ..
Ether
Soluble
Extract
Ash

Soluble
Carbohydrate

(%)

(%)

(%)

Percent
of
Total

Cel1 Walls
HemiLignin Cellulose

Insoluble
Ash

(%)

(%)

(%)

(%)

Cellulose

65
66
67
68
69
70
71
72

26.21
33.81·
30.90
26.51
29.79
29.58
30.99
30.73

11.60
17.60
17.20
12.40
14.80 .
16.40
17.20
16.00

3.38
3.26
2.49
2.62
3.14.
3.41
3.07
2.92

5.57
5.80
5.33
5.41
4.32
4.26
5.76
5.63

5.66
7.15
5.88
6.08
7.53
5.51
4.96
6.18

73.79
66.19
69.10
73.49
70.21
70.42
69.01
69.27

8.09
4.98
5.09
9.11
7.46
6.10
6.31
5.74

7.68
7.62
7.75
8.29
7.47
7.83
7.63
7.81

29.42
27.58
29.58
29.25
25.72
29.19
29.17
29.18

28.60
26.01
26.68
26.84
29.56
27.30
25.90
26.54

x

29.82

15.40

3.04

5.26

6.12

70.18

6.61

7.76

28.64

27.18

b.

r:

.;

.£

..r

I

N
N
I

�-23-

i
Table

10.

Poa (summer) - mineral analysis.

.Sample
Number

Ca

P

%

%

1

.37
.34
.34 .
.46
.34
..40
.34
.43

.17
.10
.18
.20
.11
.10
.09
.18

.38

.14

2

3
4
5
6
7
8

ppm
Hn

Mg

ppm
Cu

ppm

K

%

Fe

%

ppm
Zn

1:2.18
1:3.40
1:1.89
1:2.30
1:3.09
1:4.00
1:3.78
1:2.39

32.5
29.0.
29.0
65.0
29.0
42.0
29.0
50.0

.12
.10
.12
.l3
.11
.11
.10
.12

3~0
2.5
1.7
3.4
1.6
1.6
1.6
1.6

25.5
23.5
22.0
27.5
22.0
33.0
22.0
23.5

.94
.66
.80
1.15
.71
.62
.57
.71

20.5
19.0
19.0
23.0
19.0
17.5
19.0
25.0

1:2.71

38.2

.11

2.1

24.9

.77

17.4

ppm
Mn

Mg

prm
Cu

ppm
Fe

K

%

%

ppm
Zn

Ca/p

Table 11.

Poa (winter) - mineral analysis.

Sample
Number

Ca

P
%

Cat

%

9

10
11
12
l3
14
15
16

.49
.48
.55
.44
.46
.58
.55
.39

.06
.04
.03
.05
.06
.09
.06
.06

1: 8.17
1:12.00
1:18.33
1: 8.80
1: 7.67
1: 6.44
1: 9.17
1: 6.50

31.5
32.5
41.0
35.5
31.0
35.5
32.0
35.5

.12
.10
.11
.11
.12
.11
.13
.11

4.2
3.7
3.8
3.2
3.1
4.1
3.4
·3.8

41.5
40.0
44.0
42.5
44.0
41.5
42.0
42.0

.15
.15
.15
.15
.13
.17
.15
.13

12.5
·12.5
10 •.
5
11.0
12.5
11.0
l3.0
14.5

x

.51

.06

1.:8.50

34.3

.11

3.7

42.2

.15

12.2

P

�-24-

Table

12.

Kobresia (summer) - mineral analysis.

Sample
Number

Ca

P

%

&lt;1'

70

Ca/p

17
18
19
20
21
22
23
24

.83
.80
.80
.70
.95
.89
.78
.48

.19
.14
.18
.12
.20
.18
.12
.10

x

.78

.15

ppm
. Mn

Mg
%

ppm
Cu

ppm
Fe

.%

ppm
Zn

1:4.37
1:5.71
1:4.44
1:5.83
1:4.75
1:4.94
1:6.50
1:4.80

70.0
80.0
80.0
80.0
65.0
65.0
65.0
65.0

.17
.16
.15
.11
.18
.16
.14
.16

3.8
2.5
2.5
1.6
3.0
3.0
2.1
2.2

23.5
20.0
22.0
20.0
23.5
27.5
27.5
14.5

.54
.50
.57
.66
.57
.57
.54
.43

34.0
12.5
23.0
25.0
21.0
45.0
25.0
19.0

1:5.20

71.3

.15

2.6

22.3

.55

24.0

K

''r

••

�-25-

~

Table

Festuca (summer) - mineral analysis.

14.

Sample
Number

Ca

P
%

Ca/

p

ppm
Mn

Mg

%

33
34
35
36
37
38
39
40

.87
.89
.44
.39
.37
.38
.87
.36

.16
.16
.19
.12
.16
.18
.16
.17

1:5.44
1:5.56
1:2.32
1:3.25
1:2.31
1:2.11
1:5.44
1:2.12

32.5
39.0
42.0
25.5
35.5
45.0
45.0
39.0

.10
.11
.12
.08
.10
.17
.15
.12

x

.58

.16

1:3.63

37.9

.12

%

ppm
Cu

ppm
Fe

K
%

ppm
Zn

2.1
3.4
3.0
2.1
3.0
3.0
3.0

47.5
70.0
70.0
58.5
36.5
65.5
58.5
51.5

.46
.66
.75
.43
.62
.66
.66
.57

17.0
16.0
12.5
11.0
12.5
25.0
14.5
14.0

2.9

57.3

.60

15.3

3.8

••
Table

15.

Festtica (winter) - mineral analysis.

Sample
Number

Ca

a,

P
%

Cal

p

ppm
Mn

'Hg

%

ppm
Cu

ppm
Fe

K
%

ppm
Zn

41
42
43
44
45
46
47
48

•35
.32
.33
.36
.36
.•41
.37
.48

.04
.07
.04
.09
.07
.05
.04
.10

1:8.75
1:4.57
1:8.25
1:4.00
1:5.14
1:8.20
1:9.25
1:4.80

42.0
32.5
35.5
35.5
45.0
29.0
39.0
60.0

.07
.07
.07
.09
.07
.09
.08
.09

2.5
2.1
3.0
2.5
3.8
2.5
3.4
2.5

36.5
36.5
31.0
36:5
35.0
31.0
33.0
33.0

.15
.17
.13
.17
.13
.21
.13
.15

12.5
12.5
12.5
16.0
11.0
9.5
11.0
9.5.

x

.37

.06

1:6.17

39.8

•08

2.8

34.1 .

.16

11.8

Ie

'c,
\

�-26-

,.~.

~

Table 16.

Muh1enbergia (sunnner)- mineral analysis.

Sample
Number

Ca
%

P
%

Ca/p

49
50
51
52
53
54
55
56

.50
.53
.53
.43
.43
.39
.46
.53

.15
.16
.16
.12
.16
.17
.14
.16

x

.48

.15

Table 17.

Muh1enbergia (winter) - mineral analysis.

Sample
~umber

P
%

Ca

%

lP

ppm
Mn

~~
%

ppm
Cu

57
58
59
60
61
62
63
64

.43
.48
.41
.43
.44
.39
.45
.50

.07
.08
.04
.04
.06
.07
.05
.14

1: 6.14
1: 6.00
1:10.25
1:10.75
1: 7.33
1: 5.57
1: 9.00
1: 3.57

19.5
25.5
22.5
22.5
22.5
19.5
22.5
22.5

.07
.07
.07
.08
.07
.08
.08
.08

2.1
2.1
2.1
2.5
2.1
5.0
2.1
2.1

31.0
46.5
29.5
33.0
26.5
31.0
33.0
29.5

.13
.13
.17
.17
.15
.15
.18
.17

10.0
12.0
11.5
13.5
12.5
12.5
12.5
11.0

x

.44

.07

1: 6.29

22.1

.08

2.5

32.5

.16

11.9

Ca

ppm
Mn

Mg

ppm
Cu

ppm
Fe

K

%

%

ppm
Zn

1:3.33
1:3.31
1:3.31
1:3.58
1:2.69
1:2.29
1:3.29
1:3.31

25.5
39.0
42.0
19.5
35.5
25.5
25.5
25.5

.11
.13
.12
.10
.10
.11
.11
.13

3.4
2.5
2.5
3.0
3.0
3.0
2.1
2.1

45.5
41.5
40.5
42.0
44.0
45.5
38.5
45.5

.31
.66
.62
.35
•57
.62
.57
.71

25.0
30.5
19.5
19.0
19.5 .
21.5
16.0
22.0

1:3.20

29.8

.11

2.7

42.9

.55

21.6

ppm

.K

Fe

10

ppm
Zn

"I

�-27-

,~

\

Table 18.· Boute1oua (summer) - mineral analysis.
Sample
Number

Ca

P
%

Cal

p

ppm
Mn

Mg

%

%

ppm
Cu

ppm
Fe

K
%

ppm
Zn

65
66
67
68
69
70
71
72

.63
.73
.72
.60
.69
.65
.67
.75

.26
.26
.25
~22
.25
.26
.23
.24

1:2.42
1:2.81
1:2.88·
1:3.00
1:2.76
1:2.50
1:2.91
1:3.13

42.0
65.0
80.0
39.0·
55.0
39.0
55.0
60.0

.16
.19
.18
.16
.17
.18
.18
.19

3.0
4.7
2.1
2.5
2.1
1.6
2.1
3.8

65.5
58.5
62.0
58.5
58.5
49.5
50.5
58.5

.85
1.11
1.07
.75
.98
.94
1.11
.98

16.0
21.0
17.5
22.0
21.5
14.5
19.5
19.5

x

.68

.25

1:2.72

54.4

.18

2.7

57.7

.97

18.9

EXperimental Conclusions
The data presented shows the methods of Fonnesbeck and Harris (1970a, 1970b,
1971) to be a realistic way to partition feeds, and the procedures developed
have been adopted by this laboratory for use in nutrition studies.
The results of these analyses show signi:ficantdifferences between the nutritional quality of forages from summer and winter ranges. The mean values of
each group of samples are summarized in Tables 19 and 20 and graphically
shown in Fig. 1.

�Table 19. Mean values of nutrient constituents of forages collected from bighorn sheep rangell in
summer and winter.

Genus

Season

Percent
of
Total

Poa

Summer

Protein

Percent
Cell Content
Soluble
of
Ether
Soluble
Carbohydrate Total
Extract
Ash

Cell Walls
HemiInsoluble
Ash
Lignin Cellulose

Cellulose

(%)

(%)

5.92

30.21

27.77

2.92

13.06

32.57

34.45

3.24

8.34

30.40

20.01

(%)

.(%)

(%)

(%)

(%)

33.53

11.30

4.49

2.42

15.33

66.47

2.57

Winter

16.94

6.35

3.58

2.70

4.30

83.06

Summer

32.06

11.00

4.70

3.15

13.18

68.03

(%)

"-

Kobresia

I

Fe~

Muhlenbergia

Boute1oua

Winter

22.87

7.25

2.73

3.27

9.63

77.13

3.31.

12.75

29.50

31.44 ~

Summer

30.16

11.45

5.59

3.98

9.15

69.84

7.19

7.28

25.60

29.77

Winter

21. 27

5.15

3.79

3.24

9.10

78.73

7.07

10.42

27.73

33.51

Summer

26.22

10.30

4.40

3.97

7.55

73.78

4.62

9.60

30.87

28.68

Winter

18.35

5.00

2.16

2.99

8.20

81.65

5.05

10.64

34.98

30.99

Summer

29.82

15.40

3.00

5.26

6.1~

70.18

6.61

7.76

28.64

27.18

Winter
. .

~

~(

.."

. ..

.

.

,

'"

...

-f

.~

I

�.,

II

...

~

...•

Table 20. Mean values of mineral content from forages collected from bighorn sheep ranges in
summer and winter.

~.

�40

30

20

I

W

o
I

10

Protein

§E'J
Fig. 1.
l\

Summer;

EXtract

Carbohydrate

Ash

Ash

Lignin

Cellulose

Hemicellulose

••

Winter

Comparison of the nutritive constituents of all grass samples collected in the summer and winter.
'I

..,.

4.

-{

~

�.-3'1-

Evaluation of Methods for
Partitioning Chemical Constituents of Feces

1f'

In order to evaluate the nutritive quality of bighorn sheep ranges by
adopting nutritional procedures using the methods of Fonnesbeck and
Harris (1970a, 1970b, 1971), it was necessary to collect bighorn sheep
feces and partition the nutritive constituents. In this phase of the
project a series of bighorn sheep fecal samples were examined for three
reasons. One phase was to test for individual animal variation in
chemical breakdown of the feces. Another purpose was to determine if a
composite sample of ten individual pellet groups would represent the
average values of all the individual pellet groups in the composite.
It was felt that the only real "common denominator" for all three study
areas was the fecal samples as the plant compositions were different,
therefore, it would afford some opportunity to compare the three study
areas.
Ten individual pellet groups were collected at three study areas (Trickle
Mountain, Buffalo Peaks and Pikes Peak) at two seasons of the year
(summer-winter). Each pellet group was dried, ground in a Wiley Mill
and thoroughly mixed. One gram subsamples of each pellet group were
composited and mixed making a total of eleven samples for each study
area at each season of the year for a grand total of 66 samples for the
experiment.
All fecal samples were processed for percent cell content (% protein,
% ether extract, % soluble ash, % soluble carbohydrate) and percent cell
walls (% lignin, % insoluble ash, % holocellulose) by the methods of
Fonnesbeck and Har rd.s (1970a, 1970b, 1971). The results of these determinations are shown in Tables 21, 22, 23, 24, 25, and 26.
The results of the composited samples are shown in Table 27 and the
analysis of the results of the experiment are shown in Fig's. 2, 3, 4,
5, 6, 7, 8, 9, and 10.

'1

�-32-

- r-

50%

-r--

- •..

."

~~

45%

- ,...

40%

..,,

-,...

_L..

)I"-~

,"

~~

_L..

- •...

35%
~~

"':'~

;k

-~

30%

25%

-

-

s

w

Trickle Mtn.

.•..
s

W

Buffalo Peaks··

s W
Pikes Peak

Fig. 2. Mean values of percent cell contents arid standard deviation (.05) of the
results often
fecal samples from three study areas at two seasons of the year.
The value of a composite sample of each is designated byX.

�-33-

15%

14%

- •...
--

12%

,~

-,.....

~I'

-

,j,

,

Jf\

11%

J

K

-,.....

)~

_L...

-"-

_L...

-,.....

10%

r-

- •...

"J'

,,~
_L...

9%
_w

.r [ ·0
s

W

Trickle Mtn.

[ I:J
s

W

Buffalo Peaks

-

I" I ]
s W
Pikes Peak

Fig. 3. Mean values of percent protein and standard deviation (.05) of the
analytical results of 'ten fecal samples from three study areas at two seasons
of the year. The value of a composite sample of each is designated by X.

�-34-

16%

- .•...

- .....

-~

15

)K

14%

,~
J~

-~

-~

;k

13

-~

12%

-)(

11

- .•...
'.

,I,-

10%

_ •...

J

_L...

9

8%

-r-

7

"'7
.,

6%

T

1
s

w

Trickle Mtn.

s

w

Buffalo Peaks

J
1
w
s
Pikes Peak

Fig. 4. Mean values of percent ether extract and standard deviation(. 05) of.the
analytical results of ten fecal samples from three study areas at two seasons of
the year. The value of a composite sample of.each is designated by X.

�-35-

..

12

.~

11

.~

10

.~

9

.

,

-r-

'if
J~

..I8

7

-,~

-r-

6

,It

41'
5

.11
~I'

. -r~.

4

-•...

-'-

.

..or-

,~

_I-

Ii'

)~

3

- •...

-I-

2

~

1

.
s

w

Trickle Mtn.

s

w

Buffalo Peaks

s

w

Pikes Peak

Fig. 5. Mean values of percent soluble ash and standard deviation (.05) of
analytical results of ten fecal samples from three study areas at two seasons
of the year. The value of a composite sample of each is designated by X.

�-36-

-,-

22

·

20

~

-r-

- ....

18

-r-

16

·
-r-

14
12

10

~(

·

;~
.

·

.....

-r-

,~
..• ~

,"

_L..

_ •....

,~

_L..

-I""

8

·

6
..•

,

",
4

~

....

2

s

w

Trickle Mtn.

s

w

Buffalo Peaks

s

w

Pikes Peak

Fig. 6. Mean value of soluble carbohydrate and standard deviation (.05) of
analytical results of ten fecal samples from three study areas at two seasons
of the year. The value of a composite sample of each is designated by X.

�-37-

70

65

-~

60

..
-'~~.

-~~

55

.L...

50

45

-

T I s
.,.

T· .~ I
w

Trickle Mtn .

[71 1
s
w

Buffalo Peaks

[ - [
s

I

W

Pikes Peak

Fig.· 7. Mean values .of cell wall content and. standard deviation (.05) of
analytical results of ten fecal samples from three study areas at two seasons
of the year. The value ofa composite sample of each is designated by X.

�-38-

..

20 .

-

18

16

.~

14

..

12

.

10

.

-•...
(

-r-

~K

X

- •....
8

,~

i"

'If

~

6

~r-

'If

I~

"1\

_ ....

-~

-r-

'Ir
. ,,1\

to

4

2

-~

- •....

.

_ •....

-'-

.~
s

w

Trickle Mtn.

s

w

Buffalo Peaks

s

w

Pikes Peak

Fig. 8. Mean values of acid insoluble ash and standard deviation (.05) of
analytical results of ten fecal samples from three study areas at two seasons
of the year. The value of a composite sample of each is designated by X.

�-39-

.,..
25

"-,..

24

-23 .

I(

-

-r-

I""

"

J

22

.•

.1-

~
"
21

.100

"
;~

20

"

1\

-I'"

19

.-

--

1i1

,~

18

...•..

.-

17

-

16
15

T

[

#a

s

I ...w

1

r sJw J

r s

w

Trickle Mtn.
Buffalo Peaks
Pikes Peak
Fig. 9. Mean values of lignin and standard deviation (.05) of analytical results
of ten fecal samples from three study areas at two seasons of the year. The value
of a composite sample of each is designated by X.

"

�-40-

36

X
34

-~

32

30

.~
J~

--

••••

-: ~

28

; 1\

--

26
24

"I""'

,I'
....

22
20
0

T

[

1
s

W

c T
s

l

[
w

s

w

Buffalo Peaks
Pikes Peak
Trickle Mtn.
Fig. 10. Mean value of holoce11u1ose and standard deviation (.05) of analytical
results of ten fecal samples from three study.areas at two seasons of the year.
The value of a composite sample of each is designated by X.

�,.

x

,

36.72

.4

12.00

14.65

3.75

6.32

«

'"

63.28

12.75

20.25

;~,

30.28

�Table 22. Trickle Mountain (winter) - chemical partitioning of the nutritive constituents in bighorn
sheep feces.

Sample
Number

Percent
of
Total

Protein
(%)

Cell Contents
Ether
Soluble
Extract
Ash
(%)
(%)

11

31.04

10.40

8.46

12

28.58

7.20

13

30.58

14

35.82

Cell Walls
Soluble
Carbohydrate
(%)

Percent
of
Total

Insoluble
Ash
(%)

Lignin
(%)

HoloCellulose
(%)

4.37

7.81

68.96

11.41

19.80

37.75

7.36

5.27

8.75

71.42

10.09

25.13

36.20

10.00

8.07

4.38

8.13

69.42

11.99

20.61

36.82

10.00

6.28

3.13

16.41

64.18

11.83

20.91

31.44

N
I

15

32.14

9.20

7.00

4.17

11.77

67.86

10.11

21.57

36.18

16

34.86

10.40

7.43

4.26

12.77

65.14

11.59

22.93

30.62

17

34.94

10.00

7.70

3.72

13.52

65.06

12.35

23.10

29.61

18

35.40

9.60

7.17

3.11

15.52

64.60

11.76

23.09

29.75

19

35.60

9.20

8.06

4.37

13.97

64.32

10.57

19.44

34.31

20

37.18

8.80

7.44

5.37

15.57

62.82

10.52

24.71

27.59

x

33.61

9.48

7.49

4.22

12.42

65.48

11.22

22.13

33.03

.q

(&gt;:

...•.

I

+"-

..

..{

~

�11

.•

~

~

""

)..

Table 23. Buffalo Peaks (summer) - chemical partitioning of the nutritive constituents in bighorn
sheep feces ~
Cell Contents
Ether
Soluble
Extract
Ash

Ceil Walls
Insoluble
Ash

Lignin

;Ho1oCellulose

(%)

(%)

(%)

Sample
Number

Percent
of
Total

Protein

(%)

(%)

(%)

(%)

Percent
of
Total

21

46.96

11.60

15.83

4.08

15.45

53.04

2.91

19.78

30.35

22

39.39

13.20

15.22

5.57

10.97

60.61

12.30

20.04

28.27

23

55.48

15.60

16.79

7.81

15.28

44.52

1.12

16.04

27.36

...
'24

53.12

14.40

14.02

9.63

lS.07

46.88

4.44

15.71

26.73

Soluble
Carbohydrate

I
.p..
Vol
I

25

46.18

16.00

17.73

3.08

9.37

53.82

7.70

20.41

25.71

26

46.88

10.00

14.08

8.S9

14.21

53.12

5.88

18.31

28.93

27

47.42

11.60

15.18

6.20

14.44

52.58

4.50

18.90

29.18

28

3S.44

10.40

11.95

3.88

9.21

64.56

18.79

17.1S

28.62

29

39.09

10.00

15.14

4.68

9.27

60.91

14.05

16.55

30.31

30

46.22

9.60

12.83

8.76

15.03

53.78

6.55

18.32

28.91

x

45.62

12.24

14.88

6.23

12.83

54.31

7.83

18.12

28.44

�Table 24. Buffalo Peaks (winter) - chemical partitioning of the nutritive constituents in bighorn
sheep feces.

Cell Contents
Ether
Soluble'
Soluble
Extract
Ash
Carbohydrate

Sample
Number

Percent
of
Total

Protein

(%)

(%)

(%)

31

48.28

10.80

11.63

32

49.34

7.60

33

46.78

34

Cell Walls
Insoluble
Ash

Lignin

(%)

Percent
of
Total

(%) ,

(%)

HoloCellulose
(%)

9.97

15.88

51.72

3.66

25.69

22.37

9.85

7.51

24.38

50.66

3.42

25.52

21. 72

11.60

11.97

.9.77

13.44

53.22

14.75.

18.28

20.19

49.70

11.60

13.00

9.31

15.79

50.30

11.27

19.82

19.21

35

46.78

12.00

10.27

10.72

13.79

53.22

11.52

20.33

21.37

36

47.24

11.60

10.65

8.61

16.38

52.76

9.28

20.27

23.21

37

45.54

11.20

9.15

7.22

14.38

54.46

10.41

19.88

24.17

38

45.64

11.20

7.27

9.33

17.84

54.36

10.51

21.66

22.19

39

45.84

10.00

10.19

12.41

13.24

54.16

7.52

20.92

25.72

40

45.58

10.60

11.83

12.65

10.50

54.42

10.43

17.78

26.31

x

47.07

10.82

10.58

9.75

15.56

52.93

9.28

21.02

22.65

I

.1&gt;-'
.l&gt;I

Q

«

-q

.4.

-1'

�~

Table 25.
feces.

..

•

J,

~.

""

Pikes Peak (summer) - chemical partitioning of the nutritive constituents in bighorn sheep

Cell Contents
Ether
Soluble
Extract
Ash

Cell Walls
Insoluble
Ash

Lignin

HoloCellulose

(%)

Percent
of
Total

(%)

(%)

(%)

2.95

11.54

59.54

11.59

22.23

25.72

14.02

4.93

14.97

54.48

7.54

19.63

27.31

9.20

15.23

2.01

9.30

64.26

11.17

24.37

28.72

10.40

15.65

3.72

10.05

60.18

8.36

24.07

27.75

Sample
Number

Percent
of
Total

Protein

(%)

(%)

(%)

41

40.46

10.80

15.17

42

45.52

11.60

43

35.74

44

39.82

Soluble
Carbohydrate

I

I

I'

I

I

.pV1
I

45

48.00

8.40

11.38

3.89

24.33

52.00

11.25

16.12

24.63

46

42.14

10.80

12.46

4.10

14.78

57.86

11.32

19.33

27.21

47

39.34

8.40

15.04

3.17

12.73

60.66

6.48

23.87

30.31

48

37 •.
36

.11.60

14.72

2.43

8.61

62.64

12.02

21.35

29.27

49

41.80

10.00

10.17

2.46

19.17

58.20

8.09

24.95

25.16

50

42.36

11.60

12.54

3.18

15.04

57.64

8.92

24.41

24.31

x

41.25

10.28·

13.64

3.28

14.05

58.75

9.67

22.03

27.04

�Table 26.
feces.

Pikes Peak (winter) -chemical

partitioning of the nutritive constituents in bighorn sheep

Cell Contents
Soluble
Ether
Extract
Ash

Cell Walls
Insoluble
Ash

Lignin

HoloCellulose

(%)

(%)

(%)

Sample
Number

Percent
of
Total

Protein

(%)

(%)

(%)

(%)

Percent
of
Total

51

45.50

11. 20

8.67

8.82

16.81

54.50

6.98

21.80

25.72

52

45.34

11.60

8.32

7.18

18.24

54.66

11.09

20.21

23.36

53

43.86

11. 20

11.20

5.41

16.05

56.14

6.27

25.58

24.29

54

45.20

11. 60 .

10.70

4.69

18.21

54.80

1.57

25.60

27.63

.j::-o

55

48.36

10.80

9.93

5.95

21.68

51. 64'

8.76

21. 63

21.25 .

""

56

46.42

11.20

10.85

4.71

19.66

53.58

8.16

. 21.45

23.96

57

44.34 .

10.40

10.36

5.64

.17.94

55.66

2.65

29.28

23.73

58

47.88

12.00

8.21

7.42

20.25

52.12

2.27

25.44

24.41

59

.43.36

10.40

9.31

4.91

18.74

56.64

11.13

21. 79

23.72

60

42.96

10.00

10.54

2.32

20.10

51.04

4.02

21.30

25.72

x

45.32

11.04

9.81.

5.71

18.77

55.08

6.29

23.41

24.38

'l

Soluble
Carbohydrate

I

~

!

""'l'

..{

1"

I"

I

I

�.II

•

~

..,

'"

_J

Table 27. Chemical partitioning of the nutritive constituents in composited bighorn sheep feces
from three study areas at two seasons of the year.

Cell Content
Composite
Fecal Sample

Trickle
Mountain

Buffalo
Peaks

Pikes Peak

Cell Walls

Protein
(%)

Ether
Extract
(%)

Soluble
Soluble
Ash
Carbohydrate
(%)
(%)

Percent Insoluble
of
Ash
Total
(%)

Lignin

Ho1oCellulose

(%)

(%)

34.37

11.80

14.21

3.25

5.11

65.63

10.85

19.63

35.15

33.47

9.60

7.02

4.95

11.90

66.53

9.36

21.66

35.51

Season

Percent
of
Total

Summer
Winter

I
oj:'-

Summer

43.86

11.20

13.94

5.81

12.91

56.14

7.66

18.38

30.10

Winter

46.24

11.00

11.22

9.33

14.69

53.76

8.47

20.18

25.11

Summer

41.16

10.20

13.00

3.61

14.35

58.84

8.49

22.26

28.09

Winter

44.21

11.20

10.05

5.03

17.93

55.79

7.76

23.65

24.38

--.J
I

�-48-

Digestibility

Experiments

The digestibility of a particular type of feed or combination of feeds
by specific animals can be determined by feeding trials. These feeding
trials are carried out under very carefully controlled conditions where
all feed consumed by the animal is precisely measured and all feces from
the animal is collected.
Chemical partitioning of both the feed being
tested and the feces being eliminated are made and the difference between
those values being consumed and those values being eliminated is the
digestibility of that forage.
.
The "classical" methods employed in ·the conduct of digestion trials are
not applicable to the free ranging big game animals so several modifications
and innovations have been employed in this experiment.
These methods allow
digestion trials to be analyzed on bighorn sheep under natural conditions
on their natural ranges.
It should be emphasized that these digestion trials
are experimental and are not to be interpreted as be·ing without problems and
shortcomings.
This segment of the project was devoted to trial runs on these
digestion evaluation methods and the results are reported here.
The methods employed depend upon two main factors - one being the use of
an indigestible indicator occurring naturally in the forage and the other
is the accuracy of the identification of plant fragments in the bighorn
sheep feces by the Foods Habits Laboratory at Colorado State University.
The "indicator" used in these studies was lignin.
The plant constituent
is relatively undigestible and is easily assayed for by the methods of
Fonnesbeck and Harris (1970a, 1970b, 1971) as discussed previously.
The theory of this procedure then involves the analysis of lignin as
indicator and using the fecal analysis as inference on the food intake of
the animal.
By chemically partitioning the forages making up the diet of
the animal and the chemical partitioning of the feces we can then estimate
digestibility by using the following formula:

Digestibility

100 - (100 % Indicator
% Indicator

in Feed X % Nutrient
in Feces
% Nutrient

in Feces)
in Feed

In order to compute the total digestible nutrients (TDN) value of a feed,
the digestion coefficient of each nutritive constituent is computed by
multiplying the amount of that nutrient in 100 pounds of total feed by the
digestibility for that constituent.
(The digestibility of fat is multiplied
by 2.25 because it has more energy value than the other nutrients).
The nutritive ratio is that ratio between the digestible protein, expressed
as unity, to the sum of digestible ether extract, digestible carbohydrate,
and digestible holocellulose
(cellulose + hemicellulose).
Two separate experiments were designed to test the feasibility of the procedure to estimate digestibility.
Each experiment consisted of collecting
feeds and feces from three study areas (Trickle Mountain, Buffalo Peaks, and

�-49-

Pikes Peak) at three seasons of the year (summer, fall, winter).
In the first experiment forages were grouped into two categories (grass
and browse) where a composite sample from each grass and each browse
species collected were "pooled", thoroughly mixed and analyzed for the
nutrient constituents. Fecal samples from each area at each season of
the year were "pooled" by taking equal quantities of at least ten separate
pellet groups. These "pooled" fecal samples were thoroughly mixed and
one portion sent to the Food Habits Laboratory at Colorado State University
where they were examined for plant composition by microanalysis of plant
fragments, and the other portion was analyzed chemically to partition the
nutritive constituents.
Table 28 shows the laboratory results of the percent grass and percent
browse present in each of the pooled fecal samples.
Tables 29, 30, and 31 show the chemical analysis for the nutritive constituents in the pooled grass, browse and fecal samples from the three
study areas at three seasons of the year.

Table 28. Pooled fecal sample analysis by micro techniques for forage
identification for percent grass and percent browse.
Study Area

Season

Percent Grass

Percent Browse

Trickle Mountain

Summer

84.58

15.42

Fall

27.52

72.48

Winter

17.74

82.26

Summer

89.62

10.38

Fall

35.29

64.71

Winter

95.47

4.54

Summer

50.00

50.00

Fall

90.12

9.88

Winter

81.26

18.74

Buffalo Peaks

Pikes Peak

.'

�Table 29. Chemical partitioning of pooled grass, browse and fecal samples from the Trickle Mountain
Study Area.

Protein

(%)

(%)

(%)

(%)

Percent
of
Total

Grass

28.72

6.20

3.72

3.43

15.37

Browse

40.97

9.80

14.01

4.79

Feces

34.41

8.30

9.85

Grass

31.19

8.20

8.81

Season
Summer

,Fall

Cell Content
Soluble
Soluble
Ether
Carbohydrate
Extract
Ash

Percent
of
Total

Cell Walls
Insoluble
Lignin
Ash

HoloCellulose

(%)

(%)

(%)

71.28

7.47

5.61

58.20

12.37

59.03

6.79

7.37

44.87

7.63

·8.63

65.59

11.38

20.32

33.89

.4.21

9.97

68.81

'9.98

7.45

51.38

I

VI

0
I

Winter

'i

Browse

35.26

8.60

12.73

5.18

8.75

64.74

8.32

8.63

47.79

Feces

38.74

12.40

13.75

8.23

·4.36

61.26

13.76

21.53

25.97

Grass

21.63

.3.40

3.11

8.75

6.37

78.37

.10.21

9.37

58.79

Browse

27.37

6.20

9.13

5.45

· 6.59

72.63

8.11

7.39

57.13

Feces

30.39

6.40

11.79

8.93

3.27

69.61

11.46

26. rt

31.38

{

'&lt;)

«

-(

)1

�•

«

Ii

j

'"

Table 30. Chemical partitioning of pooled gr ass., browse and fecal samples from the Buffalo Peaks
Study Area;
Cellt.Jalls
Insoluble
Lignin
Ash

Holo..,.
Cellulose

Protein

(%)

(%)

(%)

(%)

Percent
of
Total

Grass

39.15

10.40

11.45

7.93

9.37

Browse

35.93

12.20

13.37

5.17

Feces

38.27

13.00

·12.17

8.37

Grass

31.55

8.20

10.35

6.63

6.37

68.45

5.33

7.93

55.28

Browse

33.12

7.60

13.83

5.07

6.72

66.78

5.19

6.37

55.22

Feces

35.23

12.30

11.27

8.68

2.98

64.77

12.63

20.13

32.01 .

Grass

22.44

6.00

4.63

7.44

4.37

77 .56

.8.39

10.37

58.80

Browse

27.70

8.20

7.95

5.82

5.73

72.30

5.72

7.78

58.80

Feces

28.37

9.40

·8.25

7.36

3.,36

71.63

11.76

24.73

35.14

Season
Summer

Cell Content
Soluble
Ether
Soluble
Extract
Ash
Carbohydrate

Percent
of
Total

(%)

(%)

(%)

60.85

4.39

7.76

48.70

5.19

64.07

4.63

8.76

50.68

4.73

61. 73

11.91

19.37

30.45
I

V1

Fall

Winter

t-'
I

�Table 31. Chemical partitioning of pooled grass, browse and fecal samples from the Pikes Peak
Study Area.

Fall

Cell Walls
Insoluble
Lignin
Ash
(%)
(%)

Ho Lo+

Protein

(%)

(%)

(%)

(%)

Percent
of
Total

Grass

34.96

8.80

6.73

7.70

11.73

65.04

5.63

6.48

51.93

Browse

34.61

8.80

12.17

4.32

9.32

65.39

3.31

8.37

53.71

Feces

34.03

9.80

10.36

9.51

4.36

65.97

12.17

19.33

34.47

Grass

31.68

7.80

9.94

4.57

9.37

68.32

5.73

8.37

54.22

Season
Sunnner

Cell Content
Soluble
Soluble
Ether
Extract
Ash
Carbohydrate

Percent
of
Total

Cellulose

(%)

I
lJl

N
I

Winter

.&gt;:(

Browse

33.19

8.00

13.27

5.96

5.96

66.81

4.37

8.79

53.65

Feces

35.09 .

11060

10.81

8.36

4.33

64.91

13.39

22.37

29.15

Grass

24.61

5.00

4.37

7.37

7.87

75.39

6.37

10.73

58.29

Browse

26.44

7.60

8.97

4.93

4.94

73.56

5.78

8.77

59.01

Feces

26.02

7.10

7.23

8.96

2.73

73.98

13.39

28.73

31.86

..

&lt;i

-&lt;

~

�-53-

After the laboratory analyses were completed, the total digestible
nutrient (TDN) and the nutritive ratio (NR) were computed for each
combination of grass-browse and feces from each study area at each
time of the year. Tables 32, 33, 34, 35, 36, 37, 38, 39, and 40
show the computations used in deriving these data.
-..,1

Composite diet values for protein, ether extract, soluble carbohydrate,
holocellulose, and lignin were derived by multiplying each value of the
nutrient as determined by laboratory analysis by the percent of that
feed component in the total diet and then adding -these together. For
example, if grass was analyzed as having 6.20 percent protein and grass
made up 84.58 percent of t.he diet then grass contributed 5.24 percent
protein (6.20 X .8458) and if browse was analyzed as having 9.80 percent protein and made up 15.42 percent of the diet, then browse contributed 1.51 pe-rcent of the total protein (9.80 X .1542). Adding 5.24
percent and 1.51 percent protein gives an adjusted value of 6.75 percent
protein in the composite diet.

�-54Table 32. Trickle Mountain Study Area (summer) - computations "for total
digestible nutrients and nutritive ratio for each combination of grass-browse
and feces.
84.58 % Grass
----Analysis Adjusted

15.42 % Browse
Analysis Adjusted

Composit2
Diet

Feces
Analysis

Protein

6.20

5.24

9.80

1.51

6.75

8.30

Ether Extract

3.72

3.15

14.01

2.16

5.31"

9.85

Sol.

15.37

13.00

12.37 ""

1.91

14.91

8.63

Ho10ce11ulose

58.20

49.23

44.87

6.92

56.15

33.89

Lignin

5.61

4.74

7.37

1.17

5.91

20.32

Carbohydrate

--------_
Protein

100

(100

Ether Ext;rac t

100

(100

Sol. Carbohydrate

100

(100

..

Digestion
Coefficient:

_.

"5~21

""8."3Q"
6.75
9.85
5.31

X

20.32
5.91
20.32

X

~.21

X

20.32
Ho Lo ce l.Lu Los e

100 - (100

5.91
20.32

X

)

64.24

)

46.06

8.63
14.91

)

83.17

56.15
33.89

) =

51.82

Pounds of Nutrient
in 100/'/ Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

6.75

.6424

-4.34

Ether Extract

5.31

.4606

5.50

Sol. Carbohydr~te

14.91

.8317

12.40

Ho10cel1u10se

56.15

.5182

29.10

Total Digestible Nutrients

51.34

Nutritive Ratio =
NR

(2.25)

Digestible
( Digestible)
(Ether Extract) + Carbohydrate

(Digestible Protein)
(5~50)(12.40)(29~1.Q.L
__

0=

(4.34) "
NR =

(Digestible)

+ Ho10ce11u10se

1

10.80

"-----""--

'"'_____
47.00
4.34

=

10.80

1"'-"

�-55Table 33. Trickle Mountain Study Area (fall) - computations for total
digestible nutrients and nutritive ratio for each combination of grass-browse
and feces~
72.48 x Browse
Analysis Adjusted

27.52 % Grass
Analysis Adjusted

Composite
Diet

Feces
An3.1ysis

Protein

8.20

2.26

8.60

6.23

8.49

12.40

Ether Extract

8.81

2.42

12.73

9.23

1l.65

13.75

Sol. Carbohydrate

9.97

2.74

8.75

6.34

9.08

4.36

Holocellulose

51.38

14.14

47.79

34.64

48.78

25.97

Lignin·

7.45

2.05

8.63

6.26

8.31

21.53

Digesti.on
COef[icien~
8.31
21.53

Protein

100

(100

Ether Extract

100

(100

Sol. Carbohydrate

100

HolocelluJ.ose

100 - (100

X

8.31
21.53
(100 - 8 31
21.53

X
X

8,31
21.53

X

12~)
8.49
13.75
11.65
4,36
9.08
25.97

=

43.62

)

=

54.44

)

=

81.46

)

=

79.45

48.78

Pounds of Nutrient
in 10011 Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

8.49

.4362

3.70

Ether Ex t r act;

11.65

.5444

14.27

Sol.Carbohydra re

9.08

.8146

7.39

Holocellulose

48.78

.7945

38.76

Total Di.gestible Nutrients

64.12

(2.25)

Digestible
(Digcst:ible ).-I- (Digestible
(Ether Extract) + Carbohydrate
Holocellulose)
Nutritive Ratio =
NR=

..(Df.gcs t Lb Le

.(14.27)(7.3~~.

(3.70)
16.33

Protein)

76':!J.)~_

60.42
3.70

=

16.33

�-56Table 34. Trickle ~1:ountainStudy Area (winter) - computation for tetal
digestible nutrients and nutritive ratio for each combination of grass-browse
and feces.
17.74 % Grass
Analysis Adjusted

82.26 % Br ows e
Analysis Adjusted

Composite
Diet

Analysis

Fe ce s

Protein

3.40

0.60

6.20

5.10

5.70

6.40

Ether Extract

3.11

0.55

9.13

7.51

8.06

11.79

Sol. Ca rb ohyd r at e

6.37

1.13

6.59

5.42

6.55

3.27

Holoc.ellulose

58.79

10.43

57.13

46'.99

47.42

31.38

9.37

1.66

7.39

6.08

7.74

26.77

TOO

.wlgnln

f"'.

Digestion
.Coefficient

-----Protei.n

100

(100

7.74
26.77

X

6.40
5.70

)

=

67.54

Et.he r Extract

100 - (100

7.74
26.77

X

11. 79
8.06

)

=

57.71

Sol.

100

(100

"1 2{~

Carbohydrate

100 - (100

Ho10cellulose

26.77

3 2Z

X

6.55

)

85.57

)

84.20

31.38

7.74
X

26.77

57.42

Pounds of Nutrient
in 100ft Feed

.Digestion
Coefficient

Digestible
Nutrient

Protein

5.70

.6754

3.85

Ether Extract

8.06

.5771

10.47

Sol.

6.55

.8557

5.60

57.42

.8420

48.35

Total Digestible Nutrients

68.27

Carbohydrate

Holocellulose

(2.25)

.:

Digestible
( Digestible )
Digestible
(Ether Extract) + Carbohydrate + (Holocellulose)
Nutriiive Ratio =

(Digestible Protein)
(l0.47) (5•.
.60.) (48.J.oL,,5)L--_ __6_4_.4.;..:.2
__
=

16.73
/T

(3.85)
}ffi

1

16.73

3.85

�-57Trible 35. Buffalo Peaks Study Area (s~~er) - computations for total
digestible nutrients and nutritive ratio for each combination of grass-browse
and feces.·

-_._-_._---'- _------_
..

10.38 % Browse
Analysis Adjusted

89.62 % Grass
Analysis Adjusted

·~l

Composite
Diet

Feces
Analysis

Protein

10.40

9.26

12.20

1.22

10.48

13.00

Ether Extract

11.45

10.19

13.37

1.34

11.53

12.17

Sol. Carbohydrate

9.37

8.34

5.19

0.52

8.86

4.73

Holocel1u1ose

48.70

43.34

50.68

5.07

48.41

30.45

Lignin

7.76

6.91

8.76

0.88

7.79

19.37

..

-Digestion
Coef Id c i.crrt

,..

7.79
19.37

. Protein

100

(100

Ether Extract

100

(100

Sol. Carbohydrate

100

(100

HoLcce.lIu Los e

100 - (100

7.79
19.37
Z~Z9
19.37
7.79
19.37

-

X

X

X

13.00
10.48

)

50.11

12.17
11.53

)

57.55

~.Z3

)

8.86
30.45
48.41

X

=

)

78.53
74.70

_._--_.Pounds of Nutrient
in 100!,fFeed

. Diges t Lo n
Coefficient

DigestibJ.e

·Protein

10.48

.5011

5.25

Ether Extract

11.53

.5755

14.93

Sol. Carbohydrate

8.86

.•7853

6.96

·48.41

.7470

36.16

Total Digestible Nutrients

63.30

Holocellulosc.

(2.25)

Digestible
( Digestible)
(Ether Extract) + Ca rbohyd rate
.Nutritive Ratio =
NR =
NR

(Digestible)

+ Hol~ccllulose

(Digestible Protein)
(14.93)(6.96)(36.16)
(5.25)
1 .: ._--_._----..:..:....11.08

Nutrient

58.05
5.25

11.08

�-58Table 36. Buffalo Peaks Study Area (fall) - computations for total
digestible nutrients and nutritive ratio for each combination of grassbrowse and feces.
64.71 % Browse
Analysis Adjusted

35.29 % Crass
Analysis Adjusted

Composite
Diet

Feces
Analysis

Protein

8.20

2.89

7.60

4.92

7.81

12.30

Ether Extract

10.35

3.65

13.83 ..

8.95

12.60

11.27

Sol. Carbohydrate

6.37

2.25

6.72

4.35

6.60

2.98

Ho1ocellulose

55.28

19.51·

55.22

35.73

55.24

32.01

Lignin

7.93

2.80

6.37

4.12

6.92

20.13

Digestion
Coeff i.c~ent

Protein

100

(100

6.92
20.13

X

Ether Extract

100

(100

6.92
20.13

X

Sol. Carbohydrate

100

(100

6 92

X

Ho Lo cc Ll.u.Los e

100 - (100

20.13
6.92
20.13

X

12.30
7.81
11.27
12.60
2 98
6.60
32.01
55.24

)

45.85

)

69.25

)

84.48

)

=

80.08

Pounds of Nutrient
in 10011 Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

7.81

.4585

3.58

Ether Extract

12.60

.6925

19.63

Sol. Carbohydrate

6.60

.8448

5.58

Holocellu1ose

55.24

.8008

44.24

Total Digestible Nutrients

73.03

(2.25)

Digestible
( Digestible )
Digestible
(Ether Extract) + Carbohydrate + (Holocellu1ose)
Nutritive Ratio

=

(Di~estible Protein)
(19.63)(5.58)(44.24)

---

NR=

(3.58)
NR ==

I

19.40

69.45
3.58

=

19.40

"-

�-59Table 37. Buffalo Peaks Study Area (winter) - computations for total
digestible nutrients and nutritive ratio for each combination of grassbrowse and feces.

--------95.47 % Grass
Analysis Adjusted

4.54 % Br ows e
Analysis Adjusted

Compo si te

Fe ce s

Diet

Analysis

Protein

6.00

5.73

8.20

0.37

6.10

9.40

Ether Extract.

4.63

4.42

7.95

0.36

4.78

8.25

Sol. Carbohydrate

4.37

4.17

5.73

0.26

4.43

3.36

Holocellulose

58.80

56.14

58.80

2.67

58.81

35.14

Lignin

10.37

9.90

7.78

0.35

10.25

24.73

Digestion
Coef f Lc Lent

Protein

100

(100

Ether Extract

··100

(100·

Sol. Carbohydrate

100

(100

Ho Lo ce Ll.u Los.e

100 - (100

10.25
24.73
10.25
24.73
10.25
24.73
10.25
24.73

9.40
6.10

X

8.25
)
4.78
3~3L-)
4.43

X
X

X

35.14
58.81

36.13

)
=

28.46

-.

68.56

)

75.23

Pounds of Nutrient
in 100ft Feed

Digestion
Coefficient

Digestible
Nutrie.nt

Protein

6.10

.3613

2.20

Ether Extract

·4.78

.2846

3.06

Sol. Carbohydrate

4.43

.6856

3.04

Holocellu10se

58.81

.•7523

44.24

Total Digestible Nutrients

52.54

(2.25)

Digestible
Digestible
..( Digestible ) +
(Ether Extract) + Carbohydrate
. (Holocellulose)

Nutritive Ratio = .: -'----~-'-~----'---=----.:.-......:.------(Digestible Protein)
__ (3.06)(3.04)(44.24)
50.34
== 22.88
NR=
(2.20)
2.20
NR =:

1

22.88

---

�-60Table 38. Pikes Peak Study Area (summer) - computations for total
digestible nutrients and nutritive ratio for each combination of grassbrowse and feces.

---~-.-.--50.00% Grass
Analysis Adjusted

50.00 % Br ows e
Analysis Adjusted

Composite
Diet

Feces
Analysis

Protein

8.80

4.40

8.80

4.40

8.80

9.80

Ether Extract

6.73

3.36

12.17

6.08

9.44

10.36

Sol. Carbohydrate

11.73

5.86

9.32

4.66

10.52

4.36

Ho10-::el1ulose

51.93

25.96

53.71

26.86

52.82

34.47

Lignin

6.48·

3.24 .

8.37

4.18

7.42

19.33

Digestion
Coeffici2i1t

Protein

100

(100

7.42
19.33

X

Ether Extract

100

(100

7.42
19.33

X

Sol. Carbohydrate

100

(100

1.42

- X

Holocel1u10se

100 - (100

19.33
7.42
19.33

X

9.80
8.80
10.36
9.44
ll.36
10.52
34.47
52.82

Pounds of Nutrient
in 100fi Feed

)

=

57.25

)

=

57.87

)

=

84.09

)

=

74.95

Digestion
Coefficient

Digestible
Nutrient

.5725

5.04

.5787

12.29

Protein

8.80

Ether Extract

9.44

Sol. Carbohydrate

10.52

.8409

8.85

Holocellulose

52.82

.7495

39.59

Total Digestibl~ Nutrients

65.77

(2.25)

Digestible
( Digestible ) + (Digestible
)
(Ether Extract;:L+ Carboi1ydrate
rio Loce l.Lu Io s e
Nutritiv~ Ratio

(Digestible Protein)
__

NR =

1

(.!..::1~
..
29) (8.85L(39.5~._.::::
(5.04)
12.05

_ _---60.73
..
5.04

12:05

fJr'-

�-61Table
39. Pikes Peak Study Area (fall) - computations for total
digestible nutrients and nutritive ratio for e~ch combination of grassbrowse and feces.

90.12
Analysis

% Crass
--'--Adjusted

Protein

7.80

7.02

8.00 .

Ether Extract

9.94

8.95

301. Carbohydrate

9.37

Holocellulose
Lignin

9.88 % Browse

Composite
Diet

Feces
Analysis

0.72

7.74

11.60

13.27

1.31

10.26

10.81

8.43

5.96

0.59

9.02

4.33

54.22

48.80

53.65

5.30 .'

54.10

29.15

8.37

7.53

8.79

0.87

8.40

22.37

Analysis

Adjusted

1)i~estion
COGff :i.c~.en
t

8.40
22.37

11.60
7.74

x

Protein

100

(100

Ether

100

(100 _-::-::-8_.
4:-,0
__
X ~~l_)
22.37
10.26
X
4.33)
(100 --'-2~~.!..::j;u,~~9.02

Extract

Sol. Carbohydrate

100

Holocellulose

100 _ (100 _---&gt;&lt;.8 4""'0'---_
x
22.37
0..;:'

)

.29.12 __ )
54.10

43.72

=

60.44
81.98
79.77

=

Pounds of Nuffieht
in 10011. Feed .

'!)igesti.on
Coefficient

D:tgesti.ble
:\utr:ient

Protein

7.74

.4372

3.38

Ether Extract

10.26

Sol. Carbohydrate

9.02

.8198

7.39

Holocellulose

54.10

.7977 .

43.16

(2.25)

.6044

Total Digestible

67.88

Nutrients

====================================================-:===--_ .._-Digestible.
(Ether Extract)
Nutritive

Ratio :::

Digestible)
Carbohydrate

(Dig~stible

-,--~(1:.;:.3.::...::.
95}(7. 39) (43.16)
(3..38)

JI.'R ::::,'

l-.'R -

'(

+

1

19.08
._-------'--

+

Digestible·
(Holocellulose)

Protein)

64.)0
3.38 '

'.';"
19.08

�-62Table 40. Pikes Feak Study Area (winter) - computations for total
digestible nutrients and nutritive rati,ofor each combination of gxaas+
browse and feces.
81.26 % Grass
Analysis Adjusted

18.74 % Browse
Analysis Adjusted

Composite
Diet

Feces
Analysis

Protein

5.00

4.06

7.60

1.42

5.48

7.10

Ether Extract

4.37

3.55

8.97

1.68

5.23

7.23

Sol. Carbohydrate

7.87

6.39

4.94-

0.93

7.32

2.73

Holocellulose

58.29

47.37

59.01

11.06

58.43

31.86

Lignin

10.73

8.72

8.77

1.64

10.36

28.73

Digestion
Coefficient

Protein

100

(100

Ether Extract

100

(100

Sol. Carbohydrate

100

(100

Holocellulose

100 - (100

10.36
28.73
10.36
28.73
10.36
28.73
10.36
28.73

7.10
5.48

X
X
X

X

7.23
5.23
2~73
7.32
·31.86
58.43

53.28

)
)

=

50.15

)

=

86.55

)

80.34

Pounds of Nutrient
in lOOt!-Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

5.48

.5328

.2.92

Ether Extract

5.23

.5015

5.90

Sol. Carbuhydrate

7.32

.8655

6.34

Holocellulose

58.43

.8034

46.94

(2.25)

Total Digestible Nut.rients

62.10

Digestible
(Digestible )
Digestible
(Ether Extract) + Carbohydrate· + (Holocellulos)
Nutritive Ratio NR

=

(Digestible Protein)
(5.90) (6.34) (.~4~6~.
9--:.4L.)
(2.92)

NR =:
1 :20.27
-----------_._-,--

=

59.18
2.92

=

20.27

,0-..

�-63-

The second experiment was conducted using specially compounded diets
based upon the percent food composition found in the feces. The
compounded diet was based upon those genera of plants which were found,
to be greater than 5 percent in the feces. Tables 41, 42 and 43 show
the genera of plants which were collected at the indicated season of
the year, from the specific study areas and then compounded prior to
chemical analysis.
Fecal samples were divided into two parts, one part was sent to the
food habits laboratory for analysis and the other part was kept for
chemical analysis. Table 44 shows the results of these analyses.
Tables 45 through 53 show the computations for total digestible nutrients
and nutritive ratio for each compounded forage sample and fecal sample
collected from each study area at three seasons of the year.

Table 41. Forage sample compositions from Trickle Mountain based upon
fecal analysis.
Season and Genera of Plants

Percent

Summer
Poa

7.0

Bouteloua

18.0

Carex

12.0

Festuca

57.0

Artemisia frlgida

6.0

'Fall
Carex

7.0

Festuca

18.0

Bouteloua

17.0

Artemisia frigida

46.0

Poa

12.0

Winter
Car ex

10.0

Festuca

8.0

Muhlenbergia

40.0

Poa

6.0

Artemisia frigida

20.0

Yucca

16.0

�-64-

Table 42. Forage sample compositions from Buffalo Peaks based upon
fecal analysis.
Season and Genera of Plants

Percent

Summer
Festuca

68.0

Carex

26.0

Poa

6.0

Carex

24.0

Festuca

12.0

.Kobr es La

7.0

Fall

Poa
Salix

..

12.0
45.0

Winter
Carex

6.0

Poa

8.0

Fescue

69.0

Kobresia

9.0

Salix

8.0

�-65-

Table 43. Forage sample compositions from Pikes Peak based upon,
fecal analysis.
Season and Genera of P'lantia

Percent

Summer
Carex

63.0

Festuca

13.0

Poa

7.0

Kobresia

8.0

Pinus

9.0

Carex

52.0

Fall

Festuca

(

( .

8.0

)

Poa

7.0

Stipa

6.0

Salix

27.0

Winter
Agropyron

6.0

Carex

22.0

Fescue

52.0

Kobresia

6.0

Pinus

9.0

Poa

5.0

�Table 44. Chemical analysis of compounded forage samples and fecal samples collected from three study
areas at three seasons of the year.

Cell Content
Percent
of
Total

Protein
(%)

Ether
Extract
(%)

Cell Walls

Soluble
Soluble
Carbohydrate
Ash
(%)
(%)

Percent
of
Total

Insoluble
Ash
(%)

Lignin
(%)

Ho1oCellulose
(%)

Area

Season

Trickle Mtn.

Summer

Forage
Feces

29.72
26.76

9.00
12.20

2.89
4.45

3.73
7.75

14.10
2.36

65.28
73.24

5.82
12.75

6.48
25.98

57.98
34.51

Fall

Forage
Feces

20.85
30.20

8.60
8.20

4.94
6.39

3.31
5.81

6.99
9.80

79.16
69.80

2.65
13.60

11.69
24.49

64.82
31. 71

Winter

Forage
Feces

25.14
34.76

7.40
9.20

4.71
6.94

4.27
6.93

8.76
8.69

74.86
65.24

5.80
12.18

10.78
20.60

58.28
32.46
I

Summer· Forage
Feces

25.50
31.02

9.20
60.20

2.54
4.13

3.73
7.31

10.03
3.38

74.50·
68.98

6.29
15.11

8.02
28.52

60.19
25.35

Fall

Forage
Feces

28.04
43.30

10.00
13.40 .

6.71
8.67

3.69
12.27

7.64
8.69

71.96
56.70

3.89
6.46

11.27
23.11

56.80
27.13

Winter

Forage
Feces

21.79
28.53

4.60
6.53

4.16
5.07

2.15
5.91

10.88
8.55

78.21
71.47

.2.48
11.35

8.28
23.07

67.45
37.05

Summer

Forage
Feces

34.06
36.09

14.20
17.00

4.05
5.22

4.19
9.32

11.62
4.55

65.94
69.91

3.82
8.43

8.39
26.14

53.73
29.34

Fall

Forage
Feces

20.19
32.30

5.80
10.00

5.07
4.61

3.21
6.80

6.11
10.89

79.81
67.70

4.44
10.66

9.88
24.35

73.17
32.69

Winter

Forage
Feces

27.84
40.93

8.60
16.20

4.94
6.97

3.31
10.36

10.99
7.40·

72.16
59.07

2.65
3.87

11.69
28.91

57.82
26.29

Buffalo Peaks

Pikes Peak

"'

¢

A

-4

~

-~

0\
0\
I

�-67Table 45. Trickle Mountain Study Area (summer) - computations for· total
digestible nutrients and nutritive ratio for compounded forage and fecal
samples.
Ana1;ysis .

"'\

Forage

Feces

Protein

9.00

12.20

Ether Extract

2.89

4.45

Sol. Carbohydrate

14.10

2.36

Ho10cel1u10se

57.98

34.51

Lignin

6.48

25.98

Digest:ion
Coeffident

Protein

100

(100

6.48
25.9a

X

12.20
9.00

)

=

66.19

Ether Extract

100 - (100

6~
25.98

X

~lo

!i5

)

=

61.60

)

=

95.83

)

=

85.16

Sol. Carbohydrate

100

Ho 16ceLl.uLose

100 - (100

6.tiS

(100

25.98

•

6.48
25.98

2.89
X

X

2.36
14.10
34.51
57.98

Pounds of Nutrient
in 100/1 Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

9.00

.6619

5.96

Ether Extract

2.89

.6160

4.01

Sol. Carbohydrate

14.10

.9583

13.51

Ho10ce11u10se

57.98

.8516

49.37

Total Digestible Nutrients

72.85

(2.25)

Digestible
,( Digestible ) + (Digestible
)
(Ether Extract)'
Carbohydrate
Holoce11u10se
Nutr.l~ive Ratio

=

(Df.ge s t LbLe

(4.01) (13.51) (49.37)

------------------------~~

NR=
NR;:;

(5.96)

1

11.22

Protein)

=

__

66.89
=---=~_;:;
5.96

11. 22

�-68Table 46. Trickle Mountain Study Area (fall) - computations for total
digestible nutrients and nutritive ratio for compounded forage and fecal
samples.
Analysis

,.c--

Forage

Feces

Protein

8.60

8.20

Ether Extract

4.94

6.39

Sol. Carbohydrate

3.99

9.80

Holocellulose

64.82

31. 71

Lignin

11.69

24.49

Digestion
Coe f f i.cd erit;

Protein

100

(100

-Ether Extract

100

(100

Sol. Carbohydrate

100

(100

Holoce11ulose

100 - (100

11.69
24.49
11.69
24.49
11.62
24.49

]].62
24.49

B.20
8.60

X

6.39
4.94
2.8Q
6.99

X
X
X

3] •Z]
64.82

)

54.49

=

38.27

)
)

-

33.0B
76.65

\
I

Pounds of Nutrient
in 100fi Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

8.60

.5449

4.69

Ether Extract

4.94

.3827

4.25

Sol. Carbohydrate

3.99

.3308

1.32

Holoce11ulose

64.B2

.7665

49.68

Total Digestible Nutrients

59.94

(2.25)

Digestible
Digestible
( Digestfble )
(Ether Extract) + Carbohydrate + (Holocell.ulose)
Nutr itive.Ratio =

(Digestible Protein)

~(4~._25~)~(=1.=3=2~)(~4~9.~6=B~)~
=

NR=

NR -

(4.69)
1

11.78

55.25
4.69

=

11.78
C,..

�-69Table 47. Trickle Mountain Study Area (winter) - computations for total
digestible nutrients and nutritive ratio for compounded forage and fecal
samples.
Analysis

,,-,. ,

Forage

Feces

Protein

7.40

9.20

Ether Extract

4.71

6.94

Sol. Carbohydrate

8.76

-B.69

Holocellulose

58.28

32.46

Lignin

10.78

20.60
Dj.~estion
Coefficient

Protein

100

(100

Ether Extract

100

(100

...,..

10.78
20.60

X

lQ.18

X

20.60

-1..0..78

Sol. Carbohydrate

100

Holocellulose

100 - (100

(100 -

20.60

10.18
20.60

X
X

9.20
7.40

)

34.94

6.21
4.71
8.62
8.76

)

22.89

32.~Q
58.28

)

)

48.09

=

70.85

Pounds of Nutrient
in 1001/ Feed

Digestion
Coefficient

Digestible
.Nutrient

Protein

7.40'

.3494

2.59

Ether Extract

4.71

.2289

2.43

Sol. Carbohydrate

8.76

.4809

4.21

Holocellulose

58.28

•.
7085

41.29

Total Digestible Nutrients

50.52

(2.25)

Digestible
( Digestible)
(Ether R'&lt;:tract)
+ Carbohydrate
Nutr:i:tiveRatio =

(Digestible Protein)
(2.43)(4.21)(41.29)

-

NR=
(2.59 )
NR

(Digestible)

+ Holocellulose

J.

19.50

50.52
2.59

= 19.50

�-70Table 48. Buffalo Peaks Study Area (summer) - computations for t ot'a.l
digestible nutrients and.nutritive ratio for compounded forage and fecal
samples.
Anal;t:sis
Forage

Feces

Protein

9.20

16.20

Ether Extract

2.54

4.13

Sol. Carbohydrate

10.03

3.38

Holocellulose

60.19

25.35 .

Lignin

8.02

28.52

(.1

Digestiorl
Coefficient

100 - (100

Protein

8.02
28.52

X

12.20
9.20

)

=

50.48

X

4.13
2.54

)

=

54.28

3.38

)

=

90.52

)

=

88.16

-y

Ether Extract

100 - (100

Sol. Carbohydrate

100

Ho10cel1ulose

100 - (100

(100

8.02
28.52
8.02
28.52
8.02
28.52

X
X

10.03
25.35
60.19

Pounds of Nutrient
in 100/tFeed

Digestion
Coefficient

Digestible
Nutrient

Protein

9.20

.5048

4.64

Ether Extract

2.54

.5428

3.11

Sol. Carbohydrate

10.03

.9052

9.08

Holocellulose

60.19

.•8816

53.06

(2.25)

Total Digestible Nutrients

Digestible
( Digestible)
(Ether Extract) +. Carbohydrate
Nutritive Ratio

=

(Digestible Protein)
(3.11)(9.~3.06)

=

~m;::

(4.64)
NR

+

1

14.06

69.89 .

Digestible
.
(Holocellu10se)

65.25
4.64

=

-.;:-:-

14.06

�-71Table 49. Buffalo Peaks Study Area (fall) - computations for total
digestible nutrients and nutritive ratio for compounded forage and fecal
samples.
....•

Anal~sis

\

Forage

Feces

Protein

10.00

13.40

Ether Extract

6.71

8.67

Sol. Carbohydrate

7.64

.8.69

Ho10ce11u1ose

56.80

27.13

Lignin

11.27

23.11

Digestion
Coeffic:i,ent

Protein

100

(100

11.27
23.11

X

Ether Extract

100

(100

11.27
23.11

X

Sol. Carbohydrate

100

(100

lIolocellulose

100 - (100

II

z:

23.11

13.40
10.00
816Z
6.71
8 69
.7.64

X

)

=

34.65

)

=

36.98

)

=

44.53

;

11.27
23.11

27.13
56.80

X

76.71

)

Pounds of Nutrient
in 100/1 Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

10.00

.3465

3.47

Ether Extract

6.71

.3698

5.58

Sol. Carbohydrate

7.64

.4453

3.40

Ho10ce11u1ose

56. 80~

.7671

43.57

Total Digestible Nutrients

56.02

(2.25)

Digestible
Digestible
.( Digestible ) +
(Holocellulose)
(Ether Extract) + Carbohydrate
NutriUve

----------~-------

Ratio =

(Digestible Protein)
.(5.58)(7.64)(56180)
= _---.,;5_2
_.
5_5
__
(3.74)
3.74

NR =
NR =

1

15.14

=

15.14

�-72Table 50. Buffalo Peaks Study Area (winter) - computations for-total
digestible nutrients and nutritive ratio for compounded forage and fecal
samples.
Ana1~sis
-,.0--.

Forage

Feces

Protein

4.60

9.00

Ether Extract

4.16

5.07

Sol. Carbohydrate

10.88

8.55

,Ho1oce11u10se

67.45

37.05

Lignin

8.28

23.07

Digestion
Coefficient

Protein

100 - (100

Ether Extract

100 - (100

Sol. Carbohydrate

100

HCJ1oce11u10se

100 - (100

(100

8.28
23.07
8.28
23.07
8.28
23.07
8.28
23.07

6.53
4.60

X

X
X

X

5.~)
4.16
8.55
10.88
37.05
67.45

=

49~05

=

56.26

)

=

71.80

)

=

80.29

)

Pounds of Nutrient
in 100fi Feed

Digestion
Coe.fficient

Digestible
Nutrient

Protein

4.60

.4905

2.25

Ether Extract

4.16

.5626

5.27

Sol. Carbohydrate

10.88

.7180

7.81

Ro1ocellulose

67.45

.8029

54.16

Total Digestible Nutrients

69.49

(2.25)

Digestible
( Digestibl-e ) +
Digestible
(Ether Extract) + Carbohydrate
(Ho1ocellulose)
Nutritive Ratio =

(Digestib1e Protein)
__ ~(5~.=2~7~)(~7~.=81~)~(=54~.~1~6L)=
(2.25)

NE ==

NR '"

1

29.88

67.24
2.25

=

29.88

�-73Table 51. Pikes Peak Study Area (summer) - computations for total
digestible nutrients and nutritive ratio for compounded forage and
fecal samples.·

'

Analzsis

..•.

Forage

Feces

Protein

14.20

17.00

Ether Extract

4.05

5.22

Sol. Carbohydrate

11.62

.4.55

Ho1oce1lulose

53.73

29.34

Lignin

8.39

26.14

Digestion
Co ef f Lc f ent;

Protein

100

Ether Extract

100 - (100

Sol. Carbohydrate

100

Holoce11u1ose

100 - (100

(100

(100

8.39
26.14
8.39
26.14
8,39
26.14

X
X

X

8.39
26.14

17.00
14.20
5.22
4.05
4,55
11.62

)

=

58.63

)
)

-

29.34
X

53.73

61.57

)

87.43
82.47

Pounds of Nutrient
in 100# Feed

Digestion
Coefficient

Dig2stible
:Nutrient

Protein

14.20

.6157

8.74

Ether Extract

4.05

.5863

5.33

Sol. Carbohydrate

11.62

.8743

10.16

Ho1oce11u1ose

53.73

.8247

44.31

Total Digestible Nutrients

68.54

(2.25)

Digestible
( Df.g e.st Ib ke )
Digestible
(Ether Extract) + Carbohydrate + (Ho1ocellulose)

-0-

Nutritiv~ Ratio =
NR=

(Digestible Protein)
(5.33)(10.16)(44.31)
(8.74)

NT?
1
----

6.84

59.80
8.74

6.84

�-74Table 52. Pikes Peak Study Area (fall) - computations for total d~gestible
nutrients and nutritive ratio for compounded forage and fecal samples.
Analysis
Forage

Feces

Protein

5.80

10.00

Ether Extract

5.07

4.61

Sol. Carbohydrate

6.11

10.89

Holoce1lulose

73.17

32.69

Lignin

9.88

24.35

Digestion
Coefficient

Protein

100 - (100

9.88
24.35

X

10.00
5.80

)

=

'57.60

Ether Extract

100 - (100

9.88
24.35

X

4.61
5.07

)

=

63.11

Sol. Carbohydrate

100 - (100

9 88

X

' O. 89

)

=

27.69

Holocel1ulose

100 - (100

X

32.69
73.17

)

=

81.87

24.35
9.88
24.35

6.11

Pounds of ,Nutrient
in 100/1'Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

5.80

.5760

3.34

Ether Extract

5.07

.6311

'7.20

Sol. Carbohydrate

6.11

.2769

1.69

Holocellulose

73.17

.8187

59.90,

Total Digestible Nutrients

72.13

(2.25)

Digestible
Digestible
( Digestible ) +
(Ether Extract) + Carbohydrate
(Holocc11ulose)

------------------

Nutritive Ratio

(Digestible Protein)
_-&gt;( 7. 2Qlli..Ji.ll.ill~.~90.t..J.)~
__

(3.34)
NR

1

20.60

=

68.79
_
------3.34

20.60

�-75Table 53. Pikes Peak Study Area (winter) - computations for total.digestible
nutrients. and nutri.tive ratio for compounded forage and fecal samples.
Analysis
';".

.

"

Forage

Feces

Protein

8.60

16..20

Ether Extract

4.94

6.97

Sol. Carbohydrate

10.99

7.40

Holocellulose

57.82

26.29

Lignin

11.69

28.91

Digestion
Coefficient

Protein

100 - (100

Ether Extract

100 - (100

Sol. Carbohydrate

100

Holocellulose

100 - (100

(100

11.69
28.91
11.69
28.91
ll.62
28.91
11.69
28.91

X
X
X

X

16.20
8.60
6.97
4.94
Z.40
10.99
26.29
57.82

)

=

23.82

)

42.94

)

72.77

)

=

81.6I.

Pounds of Nutrient
in 10011 Feed

Digestion
Coefficient

Digestible
Nutrient

Protein

8.60

.2382

2.05

Ether Extract

4.94

.4294

4.77

Sol. Carbohydrate

10.99

.7277

7.99

Holocellulose

57.82

.8161

47.19

Total Digestible Nutrients

62.00

(2.25)

Digestible
(Digestible ) + ( Digestible'
(Ether Extract) + Carbohydrate
Holocellu1ose)
Nutritive Ratio =

(Digestible Protein)
(4.77) (Z~9)(47 .19)
(2.05)

NR=

1

29.24

------

=

'59.95
-----2.05

29.24

�-76-

Tables 54 and 55 show the results of the two experiments previously
described. The total digestible nutrients and the nutritive ratio
resulting from the two methods are comparable, but a great deal of
additional work must be done before either technique can be used to
evaluate range quality for big game animals.

Table 54. Results of nutritive quality of bighorn sheep ranges by the
technique of compounding forage samples by percent grass and percent
browse.
Buffalo Peaks
NR
TDN

. Pikes .Peak .
NR
TDN

Season

Trickle Mountain
NR
TDN

Sunnner

51.34

1:10.80

63.30

1:11.08

65.77

1:12.05

Fall

64.12

1:16.33

73.03

1:19.40

67.88

1:19.08

Winter

68.27

1:16.73

52.54

1:22.88

62.10

1:20.27

Table 55. Results of nutritive quality of bighorn sheep ranges by the
technique of compounding forage samples by the percent plant composition.
Buffalo Peaks
TDN
NR

.Pikes Peak
NR
TDN

Season

Trickle Mountain
NR
TDN

Sunnner

72.85

1:11.22

69.89

1:14.06

68.54

1: 6.84

Fall

59.94

1:11. 78

56.02

1:15.14

72.13

1:20.60

Winter

50.52

1:19.50

69.49

1:29.88

62.00

1:29.24

-y

�-77-

LITERATURE CITED
Boussingault, Jean-Baptiste. 1843. Economic Rurale Consideree dans
ses Rapports avec la chimie, la Physique et la meteorologie.
Bechet Jeunne. Paris. (In 2 volumes). Transplated by Sir George.
Laws. 1845. Rural economics in its relation with chemistry,
physics and meteorology. D. Appleton and Company, New York.
Colburn, M. W., and J. L. Evans. 1967. Chemical composition of the
cell wall constituent and acid detergent fiber fraction of
forages. J. Dairy Sci. 50:1130.
Crampton, E. W., and L. A. Maynard. 1938. The relation of cellulose
and lignin content to the nutritive value of animal feeds. J.
Nutr. 15:383.
Dearden, B. L. 1973. Analyses of the discernibility of plant species
during digestion. PhD Dissertation, Colo. State Univ., Ft. Collins.
126 pp.
Fonnesbeck, P. V., and L. E.Harris.
1970a. Determination of plant
cell walls in feeds. Proc. Western Sec., Amer. Soc. Animal Sci.
21:153.
~

, and
1970b. Determination of holocellulose, lignin and
silica of plant cell walls. proc. Western Sec., Amer. Soc. Animal
ScL
21:162.

, and
1971. Determing hemicellulose in plant cell walls.
----Proc.
Western Sec., Amer. Soc. Animal Sci. 22:77.
Harris, L. E. 1970. Nutrition techniques for domestic and wild animals.
Vol. 1, An international record system and procedures for analyzing
samples. L. E. Harris, Logan, Utah.
Henneberg, W., and F. Stohmann.
fritterung der Weiderkur.

1860.

Henry, W. A., and F. B. Morrison. 1910.
W. A. Henry, Madison, Wisconsin.

Berundung einer rationellen

Feeds and feeding.

10th ed.

Schoonveld, G. G. 1973. Bighorn Sheep &amp; Mountain Goat Investigations Evaluation of the nutritional requirements of bighorn sheep. Colo.
Div.of Wildl. Game Res. Div., Fed. Aid Proj. W-4l-R-22. Game Res.
Rept. January.
Stewart, D. R. M. 1967. Analysis of plant epidermis in faeces; a technique for studying the food preferences of grazing herbivores.
J. App1. Ecol. 4 (1):83-111.

�-78-

Thaer, A. 1809. Grundsatze der ratione11e Landwirtschaft.
Sec. 275, Die Realschu1buchhand1ung. Berlin.

Vol. 1,

Van Soest, P. J. 1964. Symposium on nutrition and forage and pastures:
New chemical procedures for evaluating forages. J. Animal Sci.,
28: 838.
__~_;

and R. H. Wine. 1967. Use of detergents in the analysis of
feeds. IV. Determination of plant cell-wall constituents. J.
Assoc. Off. Anal. Chem. 50:50.

Zyznar, E., and P. J. Urness. 1969. Qualitative identification of
forage remnants in deer feces" J. Wildl. Mgmt. 33(3) :506-510.

=.•. -=:.~K..
•.

Prepared by _ZJ!·~"t:.!=·
I!!~ ~:"I!!'''''''''LRobert E. Keiss
Wildlife Researcher
=.

~~:...

..

�January,

-79-

JOB PROGRESS

State of

1974

REPORT

COLORADO
--------~~~-------------------

Project No.

W-4l-R-23

Work Plan No.
Job Title

Personnel:

1
Manipulation

Period Covered:

Bighorn

Sheep and Mountain

Job No.
of Vegetation

Goat Investigations

17

on Bighorn

Sheep Ranges

June 1, 1972 through May 31, 1973.

George D. Bear, Dennis Davis, Richard

Greene, William

Adrian,and

Robert Keiss.

ABSTRACT

Study ,areas were treated with nitrogen fertilizer, phosphorous fertilizer and 2,4-D
herbicide.
Work this segment was dire~ted at monitoring the effects of these treatments.
Frequency of occurrence was established for each plant species on the study plots.
Herbage production data indicated there was a significant increase in the forage produced on the Cebolla Creek (subalpine) study area due to nitrogen treatment, but there
appeared to be little difference resulting from the level of nitrogen treatment.
There
was no significant difference due to phosphorous or herbicide trea:bments. Alpine study
plots showed a significant decrease in forage production due to herbicide treatment,
and a significant increase due to phosphorous treatment.
There was not a significant
response to the application of nitrogen fertilizer.
Plants collected from the Cebolla Creek study area showed an increase in protein content
due to nitrogen treatments.
There was no change in the phosphorous or calcium content
due to the various treatments.
Chemical composition of the alpine plants was unchanged
by the various treatments.
Bighorn sheep on the alpine area did not show a preference for the vegetation on the
various plots, whereas, elk on Cebolla Creek showed a marked preference for plots
treated with nitrogen fertilizer.

��-81MANIPULATION OF VEGETATION ON BIGHORN SHEEP RANGES
GeorgeD.

·P.S.

Bear

OBJECTIVE

Improve the herbage yield, vegetative density and vegetative composition
on selected bighorn sheep ranges in Colorado.
SEGMENT OBJECTIVES
1.

Alter the quality and quantity of forage on specific study areas with
varied treatments of chemical fertilizers and herbicides.

2.

Determine the effect of each treatment on the composition, production,
and chemical content of plant species.

3.

Determine grazing preferences of wild bighorns on the area in respect
to each treatment.

METHODS AND MATERIALS
The study area is located about 35 miles south of Gunnison, Colorado. This
bighorn herd winters in two areas: in a low (8,000 feet elevation) bunchgrass and ponderosa pine type, and in a high (11,000-14,000 feet elevation)
alpine type. The following work is being conducted on each winter range.
1. Bighorn preference and response of the vegetation was measured on
plots treated with 2,4-D (herbicide), nitrogen, and phosphorus. Two levels
of treatment (0 and 2 lbs./acre) was evaluated for 2,4-D; four levels of
treatment (0, 30, 60, and 90 lbs./acre) for nitrogen; and two levels of
treatment (0 and 30 lbs./acre) for phosphorus. These were evaluated in all
.possible combinations, ·or a 2x4x2 factorial for a total of 16 treatments,
as outlined in the following table:
Treatment
Number
1
2
3
4
5
6
7

8
9
10
11
12
13
14
15
16

2,4-D
0
0
0
0
0
0
0
0
2
2
2
2
2
2
2

2

Treatment
Nitrogen
0
0
30
30
60
60
90
90
0
0
30
30
60
60
90
90

Phosphorus
0
30
0
30
0
30
0
30
0
30
0
30
0
30
0
30

�-82··

Treatment plots. measuring 50 feet by 100 feet. were established
last segment. Each corner was marked with a metal fence post. These
markers were numbered to assist in identifying plots when bighorns are
grazing on them. Three replications were randomly established on each
study area. Each replication consisted of the 16 treatment plots listed
on the preceding page. Plots in each replication were arranged in a
rectangular area eight plots long and two plots deep. Randomized selection of treatment for each plot was determined by using a table of
random numbers.
2. Frequertcysampling. Transects were established on each plot
to determine changes in vegetative composition which may result from
the various treatments. Four transects were established in each plot.
The transects parallel the base of the plot with approximatelylO-foot
spacing between transects. A total of 15 quadrats are located at 6-foot
intervals along each transect. or a total of 60 quadrats for each treatment plot. Preliminary sampling work completed during the last segment
indicated a quadrat size measuring l6x16 inches should be used on the low
winter range at Cebolla Creek; and a split-sampling scheme should be used
on the alpine area. Two quadrat sizes are used on the alpine: 1 inch by
1 inch for measuring Trifoliumrtanum and Carex sp •• and 8x8 inches for all
other species.
Frequency percentages were derived for all plant species encountered
on the quadrats. To be considered as occurring on a given quadrat, the
center of an individual plant or one-half of its basal area must be inside
the quadrat. An "individual" for bunchgrasses was a clump; for single stem
species, such as wheatgrass, it was individual stems; for cactus it was a
lobe; and for shrubs and forbs it was the basal area of all stems. Obviously,
these data are acceptable for determining changes in frequencies for each
individual species only. and cannot be used to compare one species to another.
Frequency (p) is defined p=m/n. where m is the number of quadrats containing
a given species and n is the total number of quadrats observed in each treatment. These transects were read in June and July. when the vegetation was
growing well, but before the crowns become so large they interfered with
observations.
Yield plots. Quantity of forage produced on the treatment plots
was determined by using a Neal Electronics Herbage meter. The herbage
meter measures an area 24 inches by 12 inches. which is designated as a
yield plot. Preliminary sampling data gathered last segment indicated
40 plots were needed on the alpine area for a statistically valid (p = .10)
sample and 30 plots on the Cebolla Creek area. The 60 plots established
on each treatment for the frequency plots above were also used for the
yield plots. Obviously the 60 plots is considerably greater than the
recommended numbers given above; however. the vegetation will be changing
due to treatment and larger samples will be needed as variation increases.

�-83-

In addition to the herbage meter readings, the vegetation on five
plots on each treatment was clipped and weighed. All green vegetation
on these plots was clipped at the root crown and placed in a labeled
paper sack. These samples were taken to the laboratory, oven dried,
and then weighed to the nearest gram. The weights were used to compute
regression equations for converting herbage meter readings into pounds
of forage per acre. Preliminary sampling last segment indicated at
least S clip-plots were needed on the alpine area and 3 clip-plots on
each treatment on the Cebolla Creek area to obtain statistically sound
results (P = .10).
Wire cages will be placed on the study areas to monitor the effect
of grazing on the area. The cages measuring 3 feet by 3 feet by 1 foot
high will be placed on each area. Vegetation produced inside the cages
(no grazing) will be compared with vegetation produced on areas outside
the cages (grazed areas) to get a general idea concerning the effects
of grazing.
Photo points were established on each treatment. Photos will be
taken each year at the end of the growing season to establish visual
records of changes occurring in the vegetation due to the various treatments.
Chemical analysis. Vegetative samples were analyzed for phosphorus, potassium, and protein content. This work was done at the Fort
Collins Research Center Laboratory. Since the study area is a bighorn.
winter range, the vegetation was sampled at the completion of the growing
season, again in mid-winter, and in late winter. Samples of the most
common plant species were collected from the treated areas.
3. Bighorn preference or selection of treatment plots shall be
determined. Number of bighorns on each treatment plot was recorded at
IS-minute intervals during the observation periods. Sex and age of the
sheep, time of day, general activity of the animals, weather conditions,
and snow depth (if present) was also recorded. Observations were made
with a spotting scope during daylight hours. Sampling periods extended
throughout the time the sheep occupied the winter range. Notations were
also kept on other animals (deer or elk) that use the study plots.
Records were kept on the snow depth on each treatment plot to be
related to bighorn preference for given plots. The fence posts marking
the corners of the plots provided permanent snow depth markers.

RESULTS AND DISCUSSION
Treatment of Study Plots
Ammonium nitrate and phosphate fertilizers were applied to the plots during
the last work segment (September, 1971). Herbicide (2,4-D) applications
were made on the designated plots in June, 1972, when plant growth was just
starting.

�-84-

Monitoring Treated Plots
Frequency Sampling
Percent frequency of occurrence for plant species on the study plots
treated with fertilizers and herbicide are presented in Tables 1 and
2. Data from the three replications (180 sampling plots) are combined
to give a single percentage-figure for each plant species within each
level of treatment. These frequency percentages were derived at the
beginning of the growing season; therefore the effects of the fertilizer
and herbicide treatments are not reflected here and the data serves as
a baseline to evaluate changes that occur in following years. Most common
plants encountered on the Cebolla Creek study area were: Muhlenbergia
montana, Festtica atizortica, Sitartiorthysttix; 'BOutelouagtacilis,ArtemiSia
frigida,Chrysothamrtuspattyi,Chenopodium
sp., andCrucifetae.
The most
common plants on the alpine study area were: Kobresicibellardi,Poa
sp.,
Festuca ovina, Ttisetumspicatum; AttemisiaScoptilotum; Gem turb~tum,
Umbelliferae,Polygonum
bistottoides;Ttifolium
nanum, Potentillaconcinna,
Phlox caespitosa, and Cerastium beeringianum.
Yield Plots
The relationship of herbage meter readings to herbage weights was stronger
for green weights than for oven-dry weights (Table 3). However, these
differences do not appear to be great enough to require taking both green
weights and dry weights for all samples. Since fertilizers generally
stimulate the water intake of plants as growth is increased, it is assumed
dry weights would be a better measure of the actual forage produced. Only
dry weight measurements will be made for this study in the future, thus
eliminating unnecessary labor.
The correlation coefficients were lower for these data than those computed
from the sample-data collected during the last work segment. This can
possibly be accounted for by a series of mishaps that interfered with data
collection. Equipment "breaking down" caused a delay that allowed the
alpine vegetation to start drying out before readings could be made. A
second herbage meter was borrowed to complete the readings and it is unknown
how these readings compare to those made with the original meter. It was
obvious that the borrowed meter was less stable and frequent adjustments
and checks were made to try to keep the machine "zeroed out". Of course,
the original sampling (1971) could have been inadequate. However, the
meter readings are still within acceptable limits of accuracy and the speed
of data gathering is far superior to other techniques.
Herbage production data indicated there was a significant (95 percent level)
increase in the forage produced on the Cebolla Creek study area due to the
nitrogen treatment, but there appeared to be little difference due to the
level of nitrogen treatment.
There was no significant difference due to the application of either phosphorus fertilizer or herbicide. There were very few forbs on this area to
be affected by the herbicide treatment; whereas, forbs were abundant on the
alpine area where herbage production was significantly (99 percent level)
reduced by the application of herbicide (Table 4). The alpine area showed

�-85-

a significant (90 percent level) response to application of phosphorous
fertilizer, but not to the application of nitrogen fertilizer.
Even though there were significant statistical differences in production
data, the differences in pounds of forage produced were probably not
sufficient for management purposes. This was a very dry year for the
study area. It is hoped the vegetation will respond more dramatically
to the fertilizer treatments with increased precipitation in future years.
The Cebolla Creek study area received several rainfalls in late August
and the response of the vegetation on the plots to the additional moisture
was quite noticeable.
Chemical Analysis
Samples of the following plant species were collected from the Cebolla
Creek study plots and analyzed for protein, phosphorus, and calcium;
Artemisia frigida, Muhlenbergia montana, Festuca arizonica, and Bouteloua
gracilis (Tables 5-8). Protein content of Artemisia in the fall sampling
period did not show a change as a result of treatment, however, from mid
to late winter the protein content appeared to be higher in this species
due to nitrogen treatment. Protein content in the three species of grasses
was higher on the nitrogen-treated plots throughout the winter. There
were not any definite changes in the protein levels of these four plant
species tested due to ~he various levels of nitrogen treatment. Also,
there weren't any apparent changes in the phosphorus or calcium levelsdue to the fertilizer and herbicide treatments.
Kobresia bellardi, Poa rupicola, Trifolium nanum, Oreoxis bakeri, and
Geum turbinatum were collected from the alpine study plots for chemical
analysis (Tables 9-13). The latter three species are relatively unavailable in late winter, thus, were collected only during the fall sampling
period. There did not appear to be any change in the chemical content
of these plant species due to fertilizer or herbicide treatments.
Grazing Preference
Bighorn sheep in the alpine did not show a preference for any of the
treated plots. Plots used by the sheep are listed in Table 14; these
plots were all on replication number 1 and happened to be nearest. the
bedding area. On many occasions sheep grazed near the plots without
venturing onto them. Most plots were nearly free of snow throughout the
winter so sheep had ready access to the vegetation.
The Cebolla Creek area was frequented by bighorn sheep, elk, and deer but
sheep were never observed on the study plots. It was difficult to
accurately record deer and elk usage on the plots since they were most
active during the nights. A small band of elk that frequented the locality
of replication number 2 were very selective for plants on the nitrogen
treated plots. Arizona fescue plants not treated with nitrogen were 10-14
inches tall, whereas the fescue plants on the nitrogen-treated plots were
grazed-down to a 4-inch height by March. The boundary-line contrast was

�-86-

very pronounced.
winter.

This area was relatively free of snow throughout the

Prepared by ......:~~~_~J_./-:'
=-._. _L--_~_'/_". _/_~_.
George 'ii. Bear
Wildlife Researcher

_-L--_'

�I

Table 1.

Percent frequency

of occurrence

for plant species on the·study

plots treated with fertilizers

and herbicide,

CX&gt;
-...J

Cebolla Creek, 1972.

I

,"(

Treatment!
Species

0-0-00-30-0

Pounds Per Acrua

Nitrogen

- Phos2horus

- Herbicide

30-0-0 30-30-0 60-0-0 60-30-0 90-0-0 90-30-0 0-0-2 0-30-2 30-0-2 30-30-2 60-0-2 60-30-2 90-0-2 90-30-:

G~

Agro2yron smithii
Bouteloua gracilis
Car'~ sp ,
Danthonia 23rryi
Fes[uca arizonica
Koelcria cristata
}!uhlcnbergia montana

-

-

-

25
12

21
2

17
1

51
8
62

66

38
1
65

-

-

T

69

-

}~uh.lenb(;r~

rlchardsoni
~luhll'nbL'rglatorreyi
Ory;~ol'''
Is hymcrioide.•
Poa secunda
Sitan10n hystrix
Stil~~
Vnid. Grass

T

-

-

3
24

-

3

-

3

-

26

-

1
2
1
1
32

-

2
19
14
2
59
47
53

-

4
2
25
3

-

-

-

-

3
13
7

T

-

13
8

20
8

21
14

15
4

8
17

57
1
70

48
2
45

45
7
57

51
4
64

47
4
80

67
2
70

-

-

-

2

-

-

-

-

1
21
2

-

-

5
2
22
T

-

-

-

2
1

-

7

3
27

22
22

-

-

-

-

-

-

25

-

-

3

-

T
T

13

-

1

-

-

T

7
12
8

2
17
7

69

-7

11

3

20
8

19
9

8
5

-

54
2
77

58
1
49

54
3
65

45
4
69

44
4
62

-

-2

3
1

1

1

-

23

28
-

6
-

3

5

31
2

32

38

-

--

3
6

3
3

7
7

5
5

T

1
1

-

61
T

-

22
T

-

-

-

-

-

-

-

-

-

3

T
T

Forbs
~~sp.
Castilleja sp.
Cheuopod Lum sp.
Cruc:ifcrae
Dral~ sp.
Erigcron sp.
Eriogonum sp.
.E.ri:!1;a rea sp.
GE'ranium sp.
Heuchera sp ,
~'noxys
richardsoni
Liwnula

sp ,

--

-T-

-

T

T

2
T
-

8

5
1

-

T

-

-

1

T

-

-

2
T

2
7

-

1
8

-

-

-

-

T

-

-

1
1

T

T

1
2
1

2

T

1

2
3

-

4
7

-8
T

3

-

2
7

-

4

-9-

7

6

-

-

3
-

-

-

.•.

T

3

T

T

T

T
T

2

1

1

1

-

4

1

-

3
-

3

-

T
T

T
6

2
T
-

-2

-2
-

2
1

-

2

-

-

-

2

T

1

-

T

T
1
T
1
3
--------------------------~------------------------------------------------------------------------------------~---------------------------------------_.

LC'ptodactylon pungens

4

4

-1

3

-

2

T

�I

00
00
I

Table

1.

Percent

frequency of occurrence

Species

Forbs (continued)
Hertensia sp.
PenstcMon teucrioides
Potentilla concinna
Senecio sp.
Sph3crnlcea coccinea
Unid. III
Unid. 112
Unid. 113
Un Ld , 114
Unid. 115
Unid. Sage

for plant species on the study plots treated with fertilizers

30-0-0

Treatmentl Pounds Per Acrel Nitrogen - PhosEhorus - Herbicide
30-0-2
90-30-0
0-0-2 0-30-2
60-30-0
90-0-0
30-30-0
60-0-0

0-0-0

0-30-0

4

--

2
T
1
-

-

T

1
59

4
47

3
65

2
51

7

5

-

-

T

-

-

4

-

-

-

-

1
1

2

-

and herbicide,

7

T
T

-

20

1

T

7
T

T

-

-

-

2

4

1
1

T
T

T

-

-

-

-

-

-

-

72

-

4
69

59

59

13

2

5

8

T

-

T

1
-

-

T

-

17

-

T

-

-

-

59

5
48

3

-

T

-

-

-

-

2

-

Cebolla

30-30-2

-

Creek, 1972 (continued).

60-0-:'Z-60-::3&lt;f-2 90-0-2

-

T
1

2

-2

-

90':"30-

T

3

4

2

-

2
1

-

-

-

-

2

53

1
54

2
49

48

1
55

1
47

3

5

1

4

6

9

5

-

-

'. -

-

T

1
1
1

T

-

-

T

-

2

5

3

-

-

~
Artemisia dracuncu1us
Artemisia frigida
Artemisia tridentata
Chrysothamnus parryi
Chrysothamnus
viscidi.florus
Echinocactus sp ,
Physocarpus sp ,
Rhus trilohata
Ribes cereum

------

T

2

-3

-

-

1
-

-

-

1

-

-

-

-

-

T

5

-

T

T

3

1

-

T

-

-

4
1

-

-

2

-

T

1

-

T

-

T

-

T

�,
,

cc

Table 2.

Percent frequency of occurrence

Species

0-0-0

0-30-0

for plant species on the study plots treated with fertilizers

30-0-0

and herbicides,

&lt;o

Alpine,

1972.

30-30-2

60-0-2

60-30-2

90-0-2

90-30-:

8
12

11
18

2
22

24

T

-

T

T

43
2

19
6
27
3

Pounds Per Acre, Nitrogen - PhosEhorus - Herbicide
Treatment
90-0-0
90-30-0 0-0-2 0-30-2 30-0-2
30-30-0 60-0-0 60-30-0

Grasses
Agr(~
scribneri
4
41
Carl~ spp.
Dcsch ••mosia caespitosa
Festuca ovina
35
Fcst,yca
T
Bel i.c
t o t rlcon
m&lt;&gt;rtonlanum
8
Lu-;'-;;]7J
sEicatum
3
Poa sp.
75
Trj"etum sElcatum
9
Ko h res i a bellardi
34
Un Ld , Grass

sP:--

-

-

1
16

36
3
18

36

T

T

2
6
45
27
47

10

-

6
31

-

-

T

2

13

42

24

-

-

-

8
27

5
22

T

-

23
T

43
1

33
2

35
1

31
3

27
2

43
T

9

6

8
4
86

T

T

-

4
2
79
22
46

8

5
58
21
41

9
3
72
8
37
T

9

-

66
26
47

11
67
34
45

3
61
28
47

7

3

40
2

41
4

36

35

T

r

-

79
6
50
2

11
2
75
20
30
4

T
10
63
32
45
T

3
4
71
21
38
2

8
8
80
19
45
2

2

T

T

3

-

-

3
7
17
1

T
74

2
15
36

18
26

-

-

-

20
17
7

-

-

-

-

9
14

2
38
2
36
3

-

4
49

3
16

67
1
45

-

-

-

-

-

-

67
23
36
4

44
1

T

2

-

3

T

1

-

T

T

19
14

-

T

6
25

1
1
66

6

Forbs
Achillea 1anu1osa
Androsace
s"Etentrionalis
Arcnaria fendleri
Arter.lisia scoEulorum
Caltha leptosepala
Castilleja
nee.Ldr-n t n 1 is
CastJ.ll.lli rhexffo1ia
Castj_Ll eja sp ,
Cer'l2.t
r i um
.h.,:~[in,
..Ianum
~,

if "T..'.!.£ (l)

2

-

2

T

14
14

T

67

12
25

T

-

-

-

-

-5
28

-

J:.uu:.i[!' T ,11:' (2)
ELi~~:u'n .rinn;1ti.secta 13
E.r~.r.t'-D. .0rllll£A
3

-

-

-

-

20
4

10
20

-

8
18

-

33
2

5

3
1
19
T

19
1

-

22
17
1
17
25

T

-

-

3

36
-

2
T

11

11
21

34

54

-

-

-

32
4

4
13

4
33

11
17

6
18

22
18

18
8

28

50
23

7
6

14
12

16
5

22
14

10
18

4
25

10
25

-

-

T

-

T

1
T

10
20
1
17
T

-

40
4

--------------------------------------------------------------------------------------------------------------------------------------~----------------

T

�I

1.0

Table 2. Percent frequency of occurrence for plant species on the study plots treated with fertilizers and herbicide, Alpine, 1972 (continued).

Species

0-0-0

0-30-0

Treatment. Pounds Per Acre. Nitrogen - PhosEhorus - Herbicide
30-0-0 30-30-0 60-0-0 60-30-0 90-0-0 90-30-0 0-0-2 0-30-2 30-0-2 30-30-2

60-0-2

60-30-2 90-0-2

0
I

90-30-2

Forbs (continued)
Eritrichium e10ngatum
7
14
Geurr.turbinatum
HaElopappus pygoanuB
3
Hy~cnoxys rrondif1ora
3
H8rtensia bakeri
6
Pedicular.i.s
scopulorum
Phocelia sericea
Phlo~ caespitosa
39
Polcmont um viscosum
2
~3.9..!1Ilmbistortoides 20
Polvgonum sp.
5
Potentilla concinna
40
Potentilla diversifolia Potentilla rubricaulis 9
Ranunc u l us Lnamo enus
4
Saxifr::lI~af Lagel In ris
-

-

Sax Lf racn

r hombo Ldca

SeduClstenopetalum
Sf Lc ne acnul.Ls

Tha:laspialpestre
Trifolium dasyphylum
Trifolium nanum
Trifoliumparryi
Unid. (1)
Gentiana sp.
Umbe 11iferae
UnLd, Forbs

-

T
14
T
2
39

2
57

-

3
9

-

7
36
2
1

-

-

22

38

54
5
7

34

-

-

3
15
2
4

-2

-

-

39

-

7
13
T

-

2
2

2

T

-

30

-

25
22
33
1
62
2

-

-

26
-

67

-

31
2
49

-

5
20
1
5
1
T
29

-

14
2
30

-

4
T
1
2
3
8
T

28
-

12
19

5
43

3
62

T

3
2

3
8

4
6

3
6

1
7

46

54

38

35

34

29

17
2
41

31
7
41

42
1
24

21

50
3
13

48

6
2

6
5
1

2
2
2
1

5
3
2
2

T

T

T

7

8
9

14
5

18
2

37

37
1
15

14
2
25
2
67

2
25
T
2
7

6
31
4
T
T

1

9
8
2
3

14

T

50
5
43

37

26

33

39

18
10
27

28
3
32

4

7
3
2

-

10
3
12
2
4
4

-

13
3
1
29

-

-

T

3

-

33
-

-

1

7
2

8
1

-

6

2
3

T

14

-

2

-

20

4
17

3
2

18
4
58

26

15
38

T
T

34
-

-

T

19

-

T

-3

9
16

25

T

31
-

3
48

1
44

-

T

29
4
56
1

59

28

8
30
1
2
4

24
2
72
2

9

T

13
12

3
42
1
3
13

2
1
T

34

11
3
4
27

19
8

21

-

57

-

-

T

-

63
2

-

-

-

-

-

12
3
1
49

32

42
-

-

-

23
2

50
-

12

-

61

-

-

T

-

2
16
3
4
25

T

-

T

13
71

38
3
23
2
62

T

T

-

T

-

16
69
-

-

77
T

Shrubs
Salix niva1is

-

-

-

T

2
4
1
2

�-91-

Table 3. Relationship of herbage meter readings to oven-dry and green
weights of forage on meter read plots.

.Treatment
Nitrogen-PhosphorusHerbicide

Cebolla Creek
Corr. Coef
Sample
Dry
Green
Size
Weight
Weight

Alpine
Corr. Coef.
Sample
Dry
Green
Size
Weight
Weight

0- 0-0

37

.70

.70

30

.87

.88

0-30-0

36

.78

.80

34

.83

.91

30- 0-0

36

.70

.87

36

.80

.84

30-30-0

36

.57

.59

34

.67

.65

60-.., 0-0

36

.87

.88

30

.75

.78

60-30-0

36

.68

.68

34

.83

.90

90- 0-0

36

.86

.86

30

.88

.89

90-30-0

36

.81

.80

30

.53

.76

0- 0-2

35

.74

.78

30

.56

.78

0-30-2

35

.57

.50

30

.85

.85

30- 0-2

36

.81

.81

34

.75

.75

30-30-2

36

.85

.85

34

.55

.59

60- 0-2

36

.71

.70

30

.69

.72

60-30-2

36

.72

.80

34

.78

.83

90- 0-2

36

.80

.84

30

.78

.84

90-30-2

36

.83

.88

36

.88

.91

�-92-

Table 4. Mean herbage production (pounds per acre) on plots treated
with nitrogen fertilizer, phosphorus fertilizer, and 2,4-D, 1972.

Treatment };..!
Nitrogen-Phosphorus-Herbicide

Herbage Production
(Pounds Per Acre) 2/
Cebolla
Alpine

0- 0-0

694

1145

0-30-0

627

1663

30- 0-0

794

1884

30-30-0

757

1234

60- 0-0

827

1155

60-30-0

774

1238

90- 0-0

801

1055

90-30-0

699

1456

0- 0-2

577

871

0-30-2

731

689

30- 0-2

857

837

30-30-2

999

841

60-0-2

770

915

60-30-2

908

1024

90- 0-2

797

1023

90-30-2

697

1496

1./
Treatment rate is in pounds per acre.
2/
- Pounds per acre figure for herbage was computed on a dry weight basis.

�Table 5.

Chemical analysis of Artemisia frigida collected on the Cebolla Creek Study Area, 1972-73.
Collection Date and Ana1xsis (%2
Januarx
Calcium
Phosphorus
Protein

Protein

AEril
Phosphorus

Calcium

.94

9.4

.16

.78

.14

.88

10.6

.18

.93

9.4

.15

.78

13.2

.21

.85

.93

9.4

.16

.78

13.2

.16

1.08

.14

.90

10.2

.13

.75

13.0

.18

.82

17.2

.18

.90

10.0

.07

.79

14.4

.25

·95

90- 0-0

15.6

.14

.76

10.2

.16

.85

15.4

.23

.84

90-30-0

16.8

.19

.86

11.4

.19

.81

17.4

.25

.96

Protein

SeEtember
Phosphorus

Calcium

0-0-0

14.8

.16

.81

7.0

.13

0-30-0

15.2

.16

.83

7.4

30-0-0

16.8

.18

.86

30-30-0

14.0

.16

60- 0-0

16.0

60-30-0

Treatment
(N-P-H)

I
\0

w
I

�Table 6 • Chemical analysis of Muh1enbergia montana collected on the Cebolla Creek Study Area, 1972-73.

Treatment
(N-P-H)

Collection Date and Analysis (%)
January
Phosphorus
Calcium
Protein

Protein

SeEtember
Phosphorus

Calcium

Protein

AEril
Phosphorus

Calcium

0-0-0

8.8

.16

.42

4.0

.07

.43

3.0

--

.47

0-30-0

7.6

.19

.51

2.4

.08

.48

3.6

.04

.51

30-0-0

13.2

.17

.41

4.8

.04

.41

4.4

.04

.49

30-30-0

11.2

.18

.40

5.6

.01

.43

4.2

.04

.52

60-0-0

13.2

.16

1.70

5.2

.06

.44

5.6

.05

.55

60-30-0

11.6

.17

.41

6.0

.07

.39

6.0

.05

.58

90-0-0

12.4

.16

.47

5.4

.05

.45

6.4

.05

.58

90-30-0

14.4

.18

.81

6.6

.14

.50

7.0

.07

.62

0-0-2

10.0

.16

1.34

4.0

.07

.55

3.6

.06

.45

0-30-2

8.4

.18

.41

3.4

--

--

3.6

.04

.42

30-0-2

10.0

.18

.50

4.4

--

--

4.4

.04

.47

30-30-2

10.8

.17

.42

5.2

.07

.53

5.8

.05

.49

60-0-2

12.0

.17

.58

5.8

.07

.68

5.2

.05

.58

60-30-2

12.4

.17

.63

6.0

.06

.60

5.0

.05

.56

90-0-2

13.6

.16

.58

5.0

--

--

5.4

.04

.67

90-30-2

14.0

.16

.58

7.4

.08

.73

6.4

.07

.58

I

\0

~
I

�Table.7 . Chemical analysis of Festuca ariZortica collected on the Cebolla Creek Study Area, 1972-73.
Collection Date and Analysis (%)
January
Calcium
Phosphorus
Protein

Protein

.04

.35

3.8

.06

.45

3.4

.07

.32

5.6

.06

.52

.34

4.0

.04

.33

3.6

.04

.49

.14

.40

5.4

.09

.36

4.0

.06

.52

14.0

.13

.37

5.6

.07

.34

4.6

.06

.45

14.4

.14

.37

5.0

.05

.41

5.0

.06

.52

4.6

.05

.42

Protein

SeEtember
Phosphorus

Calcium

0-0-0

7.2

.10

1. 70

3.8

0-30-0

8.4

.16

.41

30-0-0

.14

30-30-0

10.8
11.2

60-0-0
60-30-0

Treatment
(N-P-H)

April
T'hosphorus

Calcium

------

90-0-0

15.2

.15

.50

6.4

.04

.37

90-30-0

14.4

.14

.43

7.6

.10

.48

4.8

.07

.45

0-0-2

9.2

.12

.43

3.6

--

--

3.8

.06

.49

0-30-2

8.4

.14

.38

3.4

.10

.58

4.8

.06

.45

30-0-2

13.6

.12

.47

5.0

.08

.53

4.2

.06

.45

30-30-2

10.4

.18

.41

5.8

.09

.53

4.4

.05

.42

60-0-2

11.2

.13

.38

6.6

.10

.55

5.2

.05

.49

60-30-2

14.4

.14

.44

6.0

.09

.53

4.8

.07

.49

90-0-2

11.6

.14

.38

5.0

.17

.53

5.0

.04

.47

90-30-2

13.2

.14

.44

3.6

.08

.43

4.4

.07

.45

I

\0
VI
I

�Table 10- Chemical analysis of Poa rupicola collected on the Alpine Study Area, 1972-73.
Collection Date and Anal~sis (%)
Januar~
Calcium
Phosphorus
Protein

Protein

April
Phosphorus

Calcium

.49

4.0

.04

.58

.04

.48

4.2

.07

.68

6.2

.03

.55

5.4

'.04

.65

.33

6.8

.05

.44

5.4

.06

.55

.11

.30

6.4

.06

.41

6.2

.04

.65

15.6

.10

.44

6.6

.09

.58

5.6

.05

.62

.10

.34

6.6

.06

.52

6.8

.04

.68

90-30-0

13.6
11.6

.14

.31

6.8

.06

.39

6.4

.07

.82

0-0-2

9.6

.10

.37

5.2

.07

.61

4'.8

.04

.62

0-30-2

10.4

.12

.42

5.2

.07

.50

5.0

.05

.68

30-0-2

10.0

.10

.33

4.6

--

--

4.6

.03

.65

30-30-2

13.2

.11

.30

5.4

.05

.68

5.4

.04

.65

60-0-2

10.8

.09

.35

6.0

.06

.61

5.4

.04

.64

60-30-2

14.0

.10

.35

5.8

.04

.43

5.8

.05

.57

90-0-2

13.2

.10

.31

5.8

.05

.61

6.2

.06

.62

90-30-2

12.4

.11

.30

5.0

.08

.58

6.0

.09

.68

Protein

SeEtember
Phosphorus

Calcium

0-0-0

11.2

.10

.43

4.8

.06

0-30-0

10.8

.10

.81

6.6

30-0-0

10.8

.10

.34

30-30-0

11.6

.11

60-0-0

12.0

60-30-0
90-0-0

Treatment
(N-P-H)

I

\0

00
I

�-99-

Table 11. Chemical analysis of Trifolium nanum collected on the
Alpine Study Area, Sept. 1972.

Treatment
(N-P-H)

Protein

-AnalysiS -(%) Phosphorus

Calcium

0"';0-0

17.2

.14

2.20

0-30-0

17.2

.16

2.34

30- 0-0

16.0

.11

2.34

30-30-0

19.6

.15

2.28

60- 0-0

16.8

.32

2.26

60-30-0

14.4

.15

2.28

90- 0-0

19.6

.12

2.28

.16

2.34

90-30-0

Table 12. Chemical analysis of Oreoxisbakeri
Alpine Study Area, September, 1972.

collected on the

Treatment
(N-P-H)

Protein

Ana1ysis-(%) Phosphorus ,__

Calcium

0- 0-0

10.8

.10

1.65

0-30-0

10.0

.10

1.51

30-CO';'0

11.2

.10

1.70

30-30-0

12.0

.12

1.49

60- 0-0

9.2

.09

1.70

60-30-0

12.0

.13

1.70

90- 0-0

11.2

.13

1.64

90-30-0

14.0

.14

1.50

�-100Table 13. Chemical analysis ofGeum
Alpine Study Area, September, 1972.

turbinatum collected on the

Treatment
(N-P-H)

Protein

Arta1xsis (%)
Phosphorus

Calcium

0- 0-0

8.0

.10

1.44

0-30-0

6.8

.10

1.25

30- 0-0

7.6

.09

1.22

30-30-0

9.2

.09

1.40

60- 0-0

8.0

.09

0.49

60-30-0

8.8

.09

1.50

90- 0-0

10.4

.11

1.40

90-30-0

10.1 .

.08

0.43

Table 14. Number of bighorn sheep, deer and elk observed grazing on the
study plots during the period January-May, 1973.

Plot
Treatment
(N-P-H)

Number of Observations
Cebolla Creek
A1Eine
Bighorn SheeE
Elk Deer
.Bighorn SheeE
Ewes Lambs Rams Total Ewes Lambs Rams Total No. No.

0- 0-0

3

3

0-30-0
30- 0-0

2

30-30-0

3

60- 0-0

5

3

8

1

60-30-0

10

4

14

1
9

90- 0-0
6

90-30-0

6

00- 0-2
...•

6

0-30-2

3

3

30-00-2

4

6

10

30-30-2

4

5

9

6

4

4

60- 0-2
60-30-2
90- 0-2
90-30-2

2

4·

2

�January, 1974

~lOl-

JOB FINAL REPORT

State of

COLORADO

Project No.

W-4l-R-23

Work Plan No.
Job Title

Job No.

18

Control of Lungworm in Bighorn Sheep

Period Covered:
Personnel:

1

Bighorn Sheep and Mountain Goat Investigations

June 1, 1972 through May 31, 1973

Robert Lange, Steven Steinert, Julius Klein, George Bear and Robert Keiss,
Colorado Division of Wildlife; Dr. Charles Hibler, Department of Pathology,
Colorado State University.

ABSTRACT

This portion of the study is covered under contractual agreement with Colorado State
University personnel in a report entitled II Spontaneous Diseases of Bighorn Sheep",
W~4l-R, Job Contract.
No 4l-R funds were expended on this Job (Work Plan 1, Job 18).

:',

.

��-103-

January, 1974

JOB FINAL REPORT

.State of

COLORADO

Project No.

W-41-R-23

Work Plan No.

:;.1

Job Title:
Period Covered:

Bighorn Sheep and Mountain Goat Investigations
--'-_--'-

_

Job No.

Mineral Block Experimentation
June 1, 1971 through May 31, 1973

Personnel: William H.Rutherford

and Pale W. Stahlecker.

ABSTRACT

Publication plans are outlined.

19

��-105-

MINERAL BLOCK EXPERIMENTATION
William H. Rutherford

P. S~ OBJECTIVE
To assess factors involved in use of mineral blocks by bighorn sheep.

RESULTS AND DISCUSSION
A manuscript entitled "Techniques for Supplementing Diet, and Attracting
and Baiting Bighorn Sheeptl is being prepared for publication as a Game
Information Leaflet by the Colorado Division of Wildlife.
The section
of this publication dealing with the use of mineral blocks constitutes
the Job Final Report under the P.S. Objective for Work Plan 1, Job 19,
and thereby satisfies Federal Aid requirements for this study.

��January, 1974

-107-

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~~-------------

Project No.

W-41-R-23

Work Plan No.
Job Title

Job No.

20

Collection of Bighorn Lambs

Period Covered:
Personnel:

1

Bighorn Sheep and Mountain Goat Investigations

June 15, 1972 through September 15, 1972

Robert L. Schmidt, Gene G. Schoonve1d and William H. Rutherford.

ABSTRACT

Surveillance of bighorn sheep herds and collecting of lambs resulted in four lambs
being taken from Pikes Peak, two from Trickle Mountain, and one from Buffalo Peaks,
all taken as specimens for necropsy study to be done on an outside contract basis.

��-109-

COLLECTION OF BIGHORN LAMBS
William H. Rutherford

P. S. OBJECTIVE
To collect bighorn lambs as specimens for necropsy study; such study to
be done on out.s'Lde contract basis.
SEGMENT OBJECTIVES
1.

Monitor bighorn sheep herds to detect clinically ill lambs.

2.

Collect six clinically ill lambs.
METHODS AND MATERIALS

Surveillance of bighorn sheep herds on Pikes Peak, Trickle Mountain and
Buffalo Peaks was started in mid-June and continued into September in an
attempt to document characteristics of lamb behavior and appearance that
would be indicative of the progression of illness. Collection of lambs
was done on foot, using conventional firearms and placing shots for neck
hits only.
RESULTS AND DISCUSSION
Lambs on Trickle Mountain and Buffalo Peaks were never observed to show
any external indications of illness. On Pikes Peak, many lambs were showing a light-colored scruffy pelage, small size, lethargic behavior, and
spasms of coughing by late July and early August.
Collections were initiated on August 15 and terminated on September 13.
During this time, seven lambs were collected. Four were taken from Pikes
Peak on August 15, 17, 18 and 21. Two were taken from Trickle Mountain on
August 29 and 30, and one was taken from Buffalo Peaks on September 13.
The complete descriptions and necropsy reports on these lambs appear in the
report submitted by the contractor, the Department of Pathology, College
of Veterinary Medicine and Biomedical Sciences, Colorado State University.

Prepared by __~~~~~~~~~~~~~~
Rutherf rd
Wildlife Researcher

��-i u-

January, 1974

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~--------------

Project No.

W-4l-R-23

Work Plan No.

1

Job Title

Job No.

CONTRACT

Spontaneous Diseases of Bighorn Sheep

Period Covered:
Personnel:

Bighorn Sheep and Mountain Goat Investigations

July 1, 1972 through June 30, 1973.

Gene Schoonveld, Robert L. Schmidt, William H. Rutherford, Robert E. Keiss of
Colorado Division of Wildlife. C. P. Hibler, G. C. Solomon, L. H. Lauerman,
T.R. Spraker, R. E. Lange, C. J. Metzger, G. Heideman of the Colorado
State University.

ABSTRACT

Seven bighorn lambs were collected during the summer of 1972 and given a complete
postmortem examination to determine the cause of the pneumonia in some herds inColorado. The first four were collected on Pikes Peak. All four had severe pneumonia,
with the ventral lobes of both lungs completely consolidated. These animals had
heavy lungworm infection and Pasteurella spp. were isolated from the upper and lower
respiratory tract. PI-3 virus was isolated from 3 of the 4. Two lambs were collected
from the Saguache, Colorado herd. These were extremely healthy lambs that did not
have pneumonia, a light infection with lungworm and no PI-3 virus. One of the two had
Pasteurella in the upper respiratory tract. The last lamb was collected at Buffalo
Peaks. It had some small areas of pneumonia, but a light lungworm infection. Pasteurella was isolated from both the upper and lower respiratory tract, and PI-3 virus
was isolated.
.
Six foetal and neonatal bighorn lambs were examined for evidence of transplacental
transmission of lungworm and in all six the results indicated that this type of transmission is the usual means by which bighorn sheep obtain their initial infection
with lungworm, at least on Pikes Peak in Colorado. Most of these lambs were born with
heavy burdens of lungworm.
An experiment was initiated using captive sheep to experimentally reproduce this type
of transmission and determine if the ewe was capable of storing the larvae for long
periods. The results, although incomplete at present, indicate the experiment was
successful. The data have not been evaluated at this time.

(Abstract continued on reverse)

�-112-

A program of monitoring bighorn sheep herds for.evidence of their health status
by the nulnber of lungworm larvae in fecal droppings was initiated in the Pikes Peak,
Buffalo,Peaks, Trickle Mountain and Poudre Canyon herds. The results, although
inconclusive, indicate that if the sample size is sufficient, the procedure is a
good criterion for evaluating health status.
Chemotherapy trials to evaluate the efficacy of tramisol and cambendazole for
efficacy 'against the adult lungworm and third stage larvae stor~d in the ewe's
tissues indicate that.the drugs will not kill the adult paras Ltie or the third
stage larvae, but will decrease larval output by 95-98 percent for periods of six
weeks.

�.-113-

SPONTANEOUS DISEASES OF BIGHORN SHEEP
Dr. Charles Hibler

P. S. OBJECTIVE
To determine causes, nature and effects of spontaneous diseases in Colorado
bighorn sheep with particular reference to mortality in lambs, and study
various means of controlling these diseases.
SEGMENT OBJECTIVE NUMBER 1
1.

Obtain 6 clinically ill bighorn lambs from the commission for a detailed
postmortem examination.

Seven Rocky Mountain bighorn sheep were collected for studies on lamb mortality and necropsied by the personnel in the College of Veterinary Medicine
and Biomedical Sciences. Details of these collections, including dates,
sources of the animals, and gross and histopathology findings are given below.

PROCEDURE
Necropsy examinations of animals killed or dead were conducted by a veterinary
pathologist and included gross and histopathologic evaluation of all organs
and tissues as well as collection of appropriate specimens, as indicated, for
nutritional, parasitologic, bacteriologic, virologic, and toxicologic studies.
RESULTS
The results are given separately for each sheep. A tabular summary, Table 1,
precedes the written results. A detailed histologic description was written
for 72BHL-3 and 72BHL-6 and all other lambs were compared to these two lambs.

�Table 1.

Summary of pos tmo rtem resu1t.s froIl1s.evenbighorn .Lambs co.l.Lec
ted in .19.72.

Lamb
Number

Location of
collection
and date

Sex

Condition of
hair coat

Body
weight

Pneumonia

AbdoIl1ina1
fat
LungworIl1s

Pasteurella
isolated

spp.

PI-3
Isolated

72BHL-2

Pike's Peak
8-15-72

Male

Good

5011

Severe

+

+H+

+

72BHL-3

Pike's Peak
8-17-72

Male

Good

5611

Severe

+

+H+

+

72BHL-4

Pike's Peak
8-18-72

Male

Yellow and
rough

4011

Severe

-

+H+

+

+

72BHL-5

Pike's Peak
8-22-72

Male

Yellow and
rough

38/1

Severe

-

+H+

+

+

72BHL-6

Trickle Mt.
8-29-72

FeII1ale Excellent

xe.

FeIl1ale Excellent

57/1

Male

7011

+

I

72BHL-7

Trickle
8-30-72

72BHL-8

Buffalo
Peaks
9-13-72

Excellent

55/1

++

+

-.

None

++

+

+

Two sII1all
areas of
pneumonia

++

+

+

None

-

+

I-'
I-'
.pI

�-115-

I.

PATHOLOGY
72BHL-2

History
This lamb was collected on August 15, 1972 from the Cog Railroad on the
southeast side of the summit of Pikes Peak. The animal was in good physical condition and had a normal hair coat, but was observed to be coughing
at the time of collection.
Postmortem
This animal was a male lamb, weight 50 pounds, shot in the head. Blood was
taken immediately after death. Fifteen minutes after collection, postmortem
examination was initiated.

Respiratory Tract
Nasal Cavity-- The nasal mucosa was inflammed, however, no exudate was present.
The retropharyngeal lymph nodes were enlarged and moist.
Trachea--The trachea was mildly inflammed, but no exudate was present on the
mucosa.
Lungs--The changes in both the left and right lungs were essentially identical.
The ventral one-third of the apical, cardiac and diaphragmatic lobes were
completely consolidated and had a firm, meaty texture. They were dark red in
color, very similar to liver, both in color and consistency. The cut surface
was very granular as the result of many small (~ to 1 mm) nodules, all of
which bulged out from the cut surface. Very little fluid could be expressed
from the cut surfaces. The granular nodules were probably the result of lymphofollicular hyperplasia. Distended lymphatics were obvious in the subpleural
layer of the lungs. The dorsal portion of the lungs were sprinkled with white
to reddish-brown foci measuring 1 to 2 mm in size. The extreme posterior
aspect of the diaphragmatic lobe was very firm and yellowish-white in color.
The involved area was about 2.5 to 3.5 cm in size. This area was examined
in the Baermann funnel and found to contain well over a million first stage
larvae of Protostrongylus, indicating the presence of an extremely large mass
of P. stilesi. !. rushi was not found in any of the bronchioles.
The hilar and mediastinal lymph nodes were enlarged and quite moist.

Digestive System
No gross lesions were present in the digestive system. The rumen was filled
with grass and milk curd was present in the abomasum. The mesenteric lymph

�-116-

nodes were small, white and moist. Formed feces were present in the colon
and rectum. There was a moderate amount of abdominal fat present in the
abdomen.

Musculoskeletal

System

No gross lesions were present.
Central Nervous System
The eyes and brain were grossly normal.
Skin
This lamb had a smooth, tan hair coat.
Cardiovascular

No ectoparasites were found.

System

There was a moderate amount of fat around the coronary band of the heart.
No lesions were present.

Urogenital System
The testicles had both descended.
No lesions were present.

There was a small amount of kidney fat.

Histopathological
Respiratory

Changes

System

Nasal Cavity and Trachea--Same as 72BHL-3 (see 72BHL-3).

Lungs--There were more areas of alveolar and pleural edema in these sections
compared to 72BHL-3. Some of the lymphocytic hyperplasia was lighter staining
and had been replaced by fibrous connective tissue. The main histological
difference from 72BHL-2 was the lungworm nodules. The area was completely
consolidated with Langerhans' giant cells and macrophages.
Lymphocytes,
eosinophils and a few neutrophils were present. Peribronchiolar lymphocytic
hyperplasia was also very prominent. There were three to ten adults in all
the nodules and all adults were dead. The dead adults were usually surrounded
by an amorphous eosinophilic material, macrophages, Langerhans' giant cells,
and eosinophils (small granulomas). Most first stage larvae were alive;
however, a few were dead and had a similar reaction as the adults. Very few
eggs were present. There was an extensive amount of fibrosis in the alveolar
walls. The pleura was thickened in most areas due to infiltration of mononuclear cells and eosinophils.
Mediastinal

and hilar lymph nodes were similar to 72BHL-3.

�-117-

Digestive System
This lamb had several mononuclear cell infiltrations in the liver.
lesions were found in any other organ of the digestive system.

No

Musculoskeletal System
No lesions found.
Cardiovascular System
No lesions found.
Urogenital System
No lesions found.
Endocrine System
No lesions found.

Central Nervous System
No lesions found.

73BHL-3
History
This lamb was collected on August 17, 1972 from Pikes Peak. He was first
observed in a large herd which included several ewes and lambs. When observed,
he appeared to be quite normal and healthy, but would frequently have paroxysms
of coughing and lag behind the herd.
Postmortem
This animal was a male lamb, weight 56 pounds. He had a smooth, tan hair
coat. He was shot in the anterior cervical region. One hour and 10 minutes
after collection tpe postmortem examination was initiated.
Respiratory Tract
Nasal Cavity--The nasal mucosa was slightly inflammed, but no exudate was
p~esent. The retropharyngeal lymph nodes were enlarged and moist (about 1.5
em in diameter).

�-118-

Trachea--The mucosa was slightly inflammed, but no exudate was present.

Lungs--(Right lung); The entire apical lobe, the ventral two-thirds of
the cardiac lobe, and the lower one-third of the diaphragmatic lobe and
lower one-half of the intermediate lobe were reddish-brown, firm and
meaty (texture of liver) and granular in appearance. The granular appearance was due to small (~ to I mm) white foci which was probably lymphofo~licular hyperplasia. A sharp line of demarcation was also present
between the diseased and functional portion of ~he lung. Emphysema was
present on the tips of many of the lobes and between the dark red consolidated lung and the functional dorsal portion of the lung. The dorsal
portion of the lung was sprinkled with white and red foci on a pink background. There were 3 to 4 large, white, firm, circular areas on the
posterior dorsal aspect of the diaphragmatic lobe. The cut surface of
the lung (ventral aspect) had a dry, granular appearance. Only a small
amount of a mucoid material could be expressed from the bronchioles. No
Protostrongylus rushi were found in the large bronchi.
(Left lung); The lower one-half of the apical lobe, lower threefourths of the cardiac and lower one-fourth of the diaphragmatic lobe were
reddish-brown and firm. In all aspects the left lung was identical to the
right.
Mediastinal lymph nodes--These nodes were enlarged and moist.
Digestive System
There was a moderate amount of abdominal fat present in this lamb. The
rumen was filled with grass, and milk curd was present in the abomasum.
Musculoskeletal

System

No lesions were found.

Central Nervous System
No lesions were found in the brain or eyes.
Integument
Hair coat was smooth and tan.
Cardiovascular System
A moderate amount of fat was around the coronary band.
were found.

No gross lesions

�-119-

Urogenital System
Both testicles had descended. There was a small amount of renal fat.
lesions were found in the kidneys.

No

Histopathological Changes

Respiratory System
Nasal Turbinates--Lymphocytes, plasma cells and a few neutrophils were present in substantial numbers in the lamina propria of the nasal cavity. The
lymphatics and small capillaries appear slightly dilated in the lamina propria. A moderate number of mononuclear cells and plasma cells were encircled
around the mucous glands. One mucous gland contained cellular debris. Focal
areas of eosinophilia were present in the lamina propria.
Trachea--The ciliated pseudostriated squamous epithelium of the trachea was
hyperplastic in areas and atrophied in other areas (squamous metaplasia).
Short cilia were present over the hyperplastic epithelium but were absent
over the atrophied epithelium. There were a few neutrophils migrating through
the epithelium of the trachea. There was a substantial number of mononuclear
cells (lymphocytes and macrophages) and plasma cells in the lamina propria
beneath the areas of hyperplasia compared to the cellularity beneath the
areas under the atrophied mucosa. There were several large eosinophilic
amorphous structures located intracellularly in the mucosa. Several mucous
glands in the lamina propria contained exudate consisting of cellular debris
and neutrophils.
'
Lungs--(Bronchi): The mucosa of the bronchi was hyperplastic in most areas.
Cilia was also present on the epithelium. Neutrophils, eosinophils and mononuclear cells were migrating through the epithelium in many areas. A normal
number of goblet cells and mucous glands were also present. The lamina propria was thicker than normal due to an infiltration of mononuclear cells and
plasma cells with a few eosinophils and neutrophils. Many of the bronchi
contained distended blood and lymphatic vessels in their lamina propria.
Nearly all of the bronchi had hyperplastic lymphofollicular nodules. These
nodules contained large and small lymphocytes with very few mitotic figures.
The muscle coats of most bronchi were normal in thickness, however some were
atrophied. The lumen of most bronchi were partly filled with cellular debris,
composed of leukocytes, macrophages, desquamated epithelial cells, mucus, and
first stage larvae of Protostrongylus stilesi. Several vessels near bronchi
had foamy muscular coats. Many eosinophils were associated with these vessels.
The bronchioles and respiratory bronchioles were all partly filled with
cellular debris composed of neutrophils, macrophages, eosinophils, desquamated
epithelial cells, mucus, and edema. First stage larvae of P. stilesi were
also present. Some of the bronchioles were normally constricted but others
remained dilated. Cilia was still present on the epithelium of many bronchioles.

�-120-

The mucosa was hyperplastic with a normal number of goblet cells. Desquamation of epithelial cells was very common and leukocytes were migrating
through the mucosa into the lumen of many bronchioles.
The main cellular
reaction of the bronchioles was a peribronchiolar lymphocytic hyperplasia.
In some areas this was in the form of germinal centers in the lamina propria while in others a complete encircling of the medium-sized bronchioles
occurred. This complete encircling of lymphocyte hyperplasia appeared to
cause a total obstruction of many bronchioles.
There was also a large number of macrophages and a few eosinophils and
neutrophils in the lamina propria. The majority of eosinophils were, however,
outside of the muscular coat and especially around vessels. Many of the
vessels appeared to have cystic areas or holes in their tunica muscularis.
The bronchioles in the ventral aspects of the consolidated lobes often had
severely atrophied muscular layers, or none at all. Many bronchioles in
lungworm nodules in the posterior dorsal portion of the diaphragmatic lobe
were hypertrophied.
Perivascular lymphocytic cuffing occurred around many of the vessels. A
few had foamy muscular layers with eosinophils nearby. Very few vessels in
the consolidated areas were thrombosed.
The alveoli in the ventral aspects of the lobes had several different characteristics. Some lobules appeared normal except for mild peribronchiolar
lymphocytic hyperplasia.
Other lobules contained alveoli which were atelectatic with an accompanying alveolar epithelial hyperplasia.
In these
collapsed lobules many alveolar macrophages were present in the alveolar
spaces. Neutrophils were thinly scattered through these lobules. Many
respiratory bronchioles were packed with neutrophils and there appeared to
be an endobronchiolar spread of pneumonia, i.e. the number of neutrophils
were highest near the respiratory bronchioles and decreased in number as the
distance from bronchioles increased. Also, in less affected lobules, the
alveoli encircling the bronchioles were edematous and filled with macrophages and neutrophils indicating also a peribronchiolar spread of pneumonia.
Only a few first stage Protostrongylus larvae were found in the ventral
aspect of the lobes.
The majority of the P. stilesi were found in nodules in the most posterior
dorsal aspect of the-diaphragmatic lobes. These nodules contained many adults
(all alive), first stage larvae and eggs. The primary cellular reaction was
a mononuclear cellular, plasma cell and eosinophilic cellular infiltration
around the worms, eggs and larvae. Langerhans' giant cells were very common
in some areas. Many lymphocytic germinal centers were present in the tissue
especially around bronchioles and blood vessels and in the subpleura. Many
areas had a hyperplasia of the smooth muscle coat around these bronchioles
and throughout the alveolar tissue. Many bronchioles had an increased number
of goblet cells in the mucosa. Cellular debris, mucus and first stage
Protostrongylus larvae almost filled many of the bronchioles. Very few dead
larvae are present. These dead larvae are usually encircled by an eosinophilic substance, eosinophils, macrophages, ~angerhans' giant cells, and
mononuclear cells.
The pleura was relatively uninvolved over the pneumonic areas of the ventral
aspect of the lungs. The pleura over the lungworm nodules was thickened due
to mononuclear cells, eosinophils and many subpleur~l lymphocytic nodules.

�-121-

Mediastinal, Hilar, Retropharyngeal Lymph Nodes--The cortex was thickened
as a result ·of many active germinal centers. The sinuses were filled with
lymphocytes, erythrocytes and plasma cells. Thus, there was a lymphoid
hyperplasia with a high mitotic index. Also, a reticuloendothelial cell
hyperplasia was present (RE hyperplasia). Few foamy vessels were present
in the hilar nodes.

Reticuloendothelial

System

Thymus--Several foamy vessels were present. The primary change was islets
of eosinophils in the medulla of the thymus.
Spleen--No histologic lesion seen.
Digestive System
Liver--Several areas of lymphocytic infiltration were present. These were
probably due to the protostrongyloid larvae migrating in the liver before
birth.
No other histologic lesions were present in the digestive system.
Cardiovascular System
No lesion seen.

Urogenital System
No lesion seen.

Endocrine System,.
Adrenal and thyroids - normal.
Central Nervous System
Brain and cord - normal.
72BHL-4

History
This lamb was collected on August 18, 1972 about 100 yards off the Pikes Peak
toll road, near the summit. It was small, and had frequent paroxysms of
coughing. The lamb lagged behind the ewe; however, she would wait for it to

�-122-

catch-up.

'Postmortem
This was a male lamb, weight 42 pounds. The hair coat was a dull,
yellowish-tan, ragged in appearance and brittle. The postmortem interval was 40 minutes.

Respiratory System

Nasal Mucosa--The nasal mucosa was slightly inflammed and a thick, tenacious, white exudate was present between the turbinates.

Trachea--The mucosa was slightly inflammed and a thick, white, tenacious
exudate was present in the lumen. About 10 to 20 cc of this exudate poured
out from the bifurcation of the trachea when the rear of the animal was
elevated.

Right Lung--The lungs did not collapse when the thorax was opened. No
excess of fluid was present in the thoracic cavity. There was a sharp line
of demarcation between the lower one-half of the entire lung and the upper
portion. No emphysema was present above this line of demarcation as in
72BHL-3. The lower portion was dark red, meaty and firm in texture. The
cut surface was granular in appearance due to many small, one-half to threequarter nun sized nodules. There was a considerable amount of a mucoid
exudate in the small bronchiolesof
these lungs. The entire apical lobe
was consolidated, the lower three-fourths of the cardiac, about one-half of
the anterior ventral aspect of the diaphragmatic, and the entire intermediate
were completely consolidated. No f.rushi were seen in the bronchi.
Several small adhesions affixed the lower diaphragmatic lobe to the diaphragm.
Three small abscesses were associated with these adhesions.

Left Lung--The left lung was grossly identical to the right.

Mediastinal,
and moist.

Hilar and Retropharyngeal

Cardiovascular

Lymph Nodes--These nodes were enlarged

System

No fat was present around the heart.

No lesions were found in the heart.

Digestive System
No abdominal fat was present. The rumen was filled with vegetation and milk
curd was present in the abomasum.

�-123-

Musculoskeletal System
The muscles appeared to be a little pale, but this may have been due to
the fact that this lamb's heart beat was still strong during removal of
blood from the heart. A very large volume of blood was removed.

Urogenital System
No lesions; no fat around the kidney.

Endocrine System
The adrenal appeared to be congested.

Thyroids appeared normal.

Central Nervous System
No lesions were seen in the eyes or brain.
Histopathological Changes

Respiratory System
Nasal Cavity--The cellular reaction was similar to 72BHL-3 except there
were more inflammatory cells present and exudate was present in the nasal
cavity.
Trachea--The trachea was similar to 72BHL-3.
Lungs--The lungs were similar to 72BHL-3. The most impressive change was
a peribronchiolar lymphocytic hyperplasia encircling medium-sized bronchioles.
The alveoli were very hyperplastic and alveolar macrophages completely filled
the alveolar spaces.
Digestive System
No lesions were seen.
Lymphohemopoietic System
No lesions were seen.
Musculoskeletal System
No lesions were seen.

�-124-

Urogenital 'System
No lesions were seen~

Endocrine.System
No lesions were seen.

Central Nervous System
No lesions were seen.

'72BHL-5

HiStory
This lamb was collected on August 22, 1972. He had frequent paroxyms of
coughing and lagged behind the herd but the ewe waited for him to catch-up.
Postmortem
This was a small male lamb, weighing 38 pounds. He had a dull, yellow,
ragged, brittle hair coat. The body temperature was 104.20 F at the time
of death. The postmortem interval was 60 minutes.
Respiratory

System

Nasal Cavity--The mucosa was slightly inflammed, but no exudate was present.
Trachea--The

tracheal mucosa was slightly inflammed.

Lungs--The lower one-half of both lungs were identically involved. The
typical line of demarcation was present, dividing the more normal dorsal
part from the ventral dark reddish-brown, meaty tissue. Several lungworm
nodules were present in the dorsal posterior area of the diaphragmatic lobe.

Mediastinal, Hi1ar and Retropharyngea1
inf1ammed and moist.

Digestive

Lymph Nodes--These

nodes were enlarged,

System

Liver--There were several white foci (about one-half mm in diameter) just
under the capsule of the liver. The remaining portion of the digestive tract

�.-125-

appeared normaL
The rumen was filled with vegetation and milk curd was
present in the abomasum. No abdominal fat was present.

Musculoskeletal

System

Normal except again the muscle appeared a little pale.

Urogenital System
No lesions were found.

No fat was present around the kidneys.

Central Nervous System
No lesions were found.
Histopathological

Changes

All histologic changes are similar to 72BHL-3.
73BlU.-6

History
This lamb was collected on August 29, 1972 in Saguache, Colorado~ She
appeared to be quite healthy. Her hair coat was smooth and tan, and she
did not cough or lag behind the herd.
Postmortem
This lamb was a female, weight 55 pounds, body temperature 104.60 F (she ran
a short distance before collection). She was in excellent body condition and
had a large amount of subcutaneous body fat. No ectoparasites were found.
The postmortem interval was 15 minutes.

Respiratory System
Nasal Cavity--Lamb 72BlU.-6was shot at the base of the head. The nasal
turbinates and nasopharynx were totally destroyed. The trachea appeared normal.
Lungs--The lungs collapsed normally when opened. They were a soft, salmonpink color. The only lesions were 2 or 3 small, whitish-yellow, firm nodules
(one~half to one cc in diameter) located in the most dorsal posterior aspect
of the diaphragmatic lobes. The mediastinal and hilar nodes were small and
dry. No adhesions or pleural fluid were present. No P. rushi were found.

�-126-

Cardiovascular System
The heart was normal grossly.
the coronary vessels.

There was a large amount of fat around

Digestive System
The rumen was filled with vegetation and there was a small amount of milk
curd in the abomasum. The digestive tract seemed to be larger in this lamb
than the lambs from Pikes Peak. The mesenteric lymph nodes were large,
white and moist (this is normal for these nodes).
Lymphohemopoietic

System

The spleen was small and contracted. There were many red, raised foci
around the peripheral area of the spleen which are normal. A small amount
of fat was present in the bone marrow.
Musculoskeletal

System

No gross lesion.

Urogenital System
No gross lesion.

Fat was present around both kidneys.

Endocrine System
Thyroids and adrenals were grossly normal.

Central Nervous System
Brain and eyes were grossly normal.

Histopathological

Changes

Respiratory System

Nasal Turbinates--A few eosinophils, mononuclear cells and plasma cells
were present in the lamina propria.
Trachea--Qne section showed that the mucosa on the ventral aspect of the
trachea was slightly thicker than the dorsal part. This ventral aspect also
had a few eosinophils, mononuclear and plasma cells in the lamina propria
under the slightly thickened epithelium.

�-127-

Bronchi--There were no lesions in the bronchi.
BronchiolesandRespiratorYBronchioles:
A few bronchioles contained mucus
and cellular debris. Most bronchioleswere
clear. The mucosa was not hyperplastic and the lamina propria had a normal cell population. About 10 to 20
percent of the bronchioles had a peribronchiolar lymphocytic hyperplasia in
the apical and cardiac lobes and ventral aspect of the diaphragmatic lobes.
From 50 to 70 percent of the bronchioles in and near lupgworm nodules also
had this lymphocytic hyperplasia.
These lymphatic nodules were usually
round and were located either in the lamina propria or just outside the
muscle coats of the bronchioles.
Alveoli--There was a diffuse stromal thickening of all slides of all lobes.
There were very few neutrophils, eosinophils and large mononuclear cells in
these areas. The importance of this was not known - possibly some type of
allergic response. There was a minimal amount of perivascular cuffing of
lymphocytes.
The dorsal posterior aspect of both diaphragmatic lobes had small lungworm
nodules. These nodules were characterized by several adults and many eosinophilic staining eggs and first stage larvae. The first stage larvae could
be identified as Protostrongylus larvae by their sharply pointed tail, with
a nick at the base of the tail. The adults were usually at the periphery
of the nodule. Some adults had no cellular reaction around them. It appears
that the adults crawl around and deposit their eggs then leave to another
area. The eggs then hatch into first stage larvae and make their way into
the bronchioles.
The majority of the cellular reaction was associated with
the eggs and larvae and around some of the adults. The total cellular reaction is minimal compared to lungworm nodules of "sick lambs". The cellular
response in this lamb was an epithelial hyperplasia in some areas; many
bronchioles and vessels were encircled by a lymphocytic hyperplasia (germinal
centers). Macrophages, eosinophils, Langerhans' giant cells, and lymphocytes
were associated with these larvae and eggs. Many germinal centers of lymphocytic tissue were subpleural.
Mediastinal Lymph Node--This lymph node contained many active germinal centers
in the cortex. These germinal centers had a mixture of small and large lymphocytes. Mitotic figures were few. TWo parasites were present. There was an
eosinophilic and lymphocytic infiltration around one of the worms. The other
worm had no noticeable cellular reaction. The afferent lymphatics were filled
with lymphocytes and a few eosinophils.
Digestive System
The liver had one small focus which was an area of lymphocytic infiltration.

All Other Systems
All other systems were histologically normal.

�-128-

. 72BHL-7
.·HistorY
This lamb was collected on August 30, 1972 at Saguache, Colorado. She
appeared to be quite healthy and did not cough or lag behind the herd.
Postmortem
72BHL-7 was a female lamb, weight 57 pounds. Her hair coat was in excellent
condition and she had a considerable amount of body fat. The postmortem
interval was 65 minutes.
Respiratory

System

Nasal Cavity--The lamb regurgitated before death and rumen contents were
present in the nasal cavity; however, the cavity appeared to be quite normal.
Trachea--Rumen contents did not extend beyond the larynx. The tracheal
mucosa was grossly normal. No exudate was present. The retrophar:yngeal
lymph nodes were small (about one mm in diameter) and dry, with a white
cortex and slightly brown medulla.

Lungs--Both lungs collapsed normally when the thoracic cavity was opened.
The lungs were a soft salmon-pink with a soft, spongy consistency. There
were three small, firm, whitish-yellow nodules located in the posterior
dorsal aspect of both diaphragmatic lobes. No adhesions were present and
no fluid was present in the thoracic cavity. The pleura seemed normal on
gross examination.

Cardiovascular

System

There was considerable fat around the coronary band of the heart.
lesions were present in the heart or the vessels.
Lymphohemopoietic

No gross

System

The spleen was small and contracted. There were many small, red foci around
the peripheral aspect of the organ. '
Digestive System
The rumen was filled with vegetation.
The abomasum had a small amount of milk
curd. No parasites were found. The entire digestive tract was as large as
72BHL-6. There were three focal hemorrhages in the cecum. Abdominal fat was
present in this animal. The liver was grossly normal.

�-129-

Musculoskeletal

System

No gross lesions.
Urogenital· System
Renal fat was present.
productive tract.

No gross lesions were found in the kidneys or re-

Endocrine·System
Thyroids andadrenals

- normal.

Central Nervous System
The animal had been shot in the brain.
Histopathological

Eyes were normal.
Chartges

The histopathological changes in 72BHL-7 were very similar to 72BHL-6.
Only stromal thickening with infiltration of inflammatory cells was obvious.
Focal areas of pneumonia were present in this lamb. Foamy vessels in the
lung were also present in this animal. Some vessels were in the center of
hemorrhagic areas and eosinophils and mononuclears were surrounding these
vessels.
The only other difference in 72BHL-7 was the presence of 6 parasites, three
alive and three dead, located in a mandibular lymph node. Another parasite
was present in the anterior part of the muzzle. The thymus has focal islets
of eosinophils in the medulla of the gland. One area of eosinophilic infiltration was present in the liver.
72BHL-8

History
This lamb was collected on September 13, 1972 at Buffalo Peaks. The animal
was in excellent physical condition and did not cough or lag behind the herd.
Postmortem
72BHL-8 was a 70 pound male lamb in excellent body condition with a smooth,
tan hair coat and considerable body fat. The postmortem interval was 2 hours.

�-130-

Respiratory System

Nasal Cavity--Nasal turbinates were slightly inflammed, with a slight
excess of mucus.

Trachea--The trachea was reddened with an increase in mucoid material
present on its mucosa.

Lungs-~(Right lung): The lungs collapsed moderately when the thorax was
opened. There was a small, firm, dark red, depressed lobule on the most
ventral aspect of the anterior diaphragmatic lobe. There was a very small
adhesion from this depressed, firm area to the diaphragm. Four to five firm,
yellowish-white, circular nodules were present on the dorsal posterior aspect
of the diaphragmatic lobe. A copious amount of a mucoid exudate was present
at the bifurcation of the trachea and extended downward into both mainstem
bronchi. Exudate (mucoid) was not noticed in the small bronchioles of the
lung.
(Left lung):. The left lung was similar grossly.' There was a firm,
dark red, depressed area located on the ventral aspect of the cardiac lobe.
Four to five circular, yellowish-white nodules were present on the dorsal
posterior diaphragmatic lobe. One small focus of hemorrhage was present
on the dorsal aspect of the cardiac lobe.

Mediastinal and Retropharyngeal Lymph Nodes--The mediastinal and retropharyngeal lymph nodes were slightly enlarged.
Cardiovascular System
There was a moderate amount of fat around the coronary band. No excess of
fluid was present in the pericardial sac. No lesions were seen on the heart.
Lymphohemopoietic

System

No lesions were seen in the spleen. The spleen was small and contracted and
had many small red foci around the periphery of the organ.
Digestive System
No lesions in the liver or intestines were observed. The rumen and abomasum
were filled with vegetation; no milk curd was present in the abomasum. Formed
feces were present in the colon and rectum. Considerable abdominal fat was
present.

Musculoskeletal

System

The right frontal sinus contained an excessive amount of a mucoid exudate

�-131-

which contained white flecks of material (Pasteurella hemolytica and
Streptococcus spp. were cultured from the sinus).

Urogenital SYstem
No lesions were seen.

Renal fat was present.

Endocrine System
Thyroids and adrenals appeared normal.

Central Nervous System
No lesions in the brain or eyes were noted.
Histopathological Changes
Respiratory System
Nasal Turbinates--Some areas of nasal mucosa were hyperplastic and others
were beginning to undergo metaplasia to a more cuboidal type epithelium.
The mucosa seemed normal in other areas. There was an increased number of
mononuclear cells and plasma cells in the lamina propria, but only a few
neutrophils or eosinophils could be found. The capillaries in the lamina
propria appeared slightly dilated.

Trachea--The mucosa was normal to slightly hyperplastic in areas. The
cilia were either absent or small. There were a few neutrophils and mononuclear cells migrating through the mucosa. An increase in mononuclear
cells and plasma cells was present in the lamina propria just below the
mucosa. The capillaries in the lamina propria were engorged. There appeared
to be a normal number of goblet cells in the mucosa.

Lungs--(Bronchi): Most bronchi had a moderate amount of mucus and cellular
debris. The mucosa was slightly hyperplastic with a few neutrophils and
mononuclear cells migrating through the mucosa. A few desquamating epithelial
cells were present. The lamina propria had an increased number of plasma
cells and mononuclear cells (mostly lymphocytes) present just under the
lamina propria. The muscular coat was of normal thickness. The vessels
appeared normal.
(Bronchioles): The primary cellular change in the bronchioles was
again a lymphocytic hyperplasia around the medium-sized bronchioles. Sections
taken more ventrally have a higher percentage of bronchioles involved with
peribronchiolar lymphocytic hyperplasia. The magnitude of this lymphocytic
hyperplasia appears to have completely cut off or obstructed some of the
medium sized bronchioles. The mucosa of these bronchioles was hyperplastic

�-132-

and some of their epithelial cells were desquamating. The brortchioles
are partly filled with a mucoid exudate with cellular debris. Leukocytes were migrating through the mucosa in many places. There was an
increase in cell population in the lamina propria which was made up mostly
of mononuclear cells and plasma cells, with a few neutrophils and very few
eosinophils. The muscular coat around vessels which were encircled by
lymphocytic hyperplasia had atrophied. Areas where there were no lymphocytic hyperplasia, the muscle coat was normal in thickness. Many small
an4 medium-sized vessels were also encircled by lymphocytis.
(Alveoli): In most sections, a similar type of stroma thickening as
in 72BHL-7 was also present in this lamb. Several lobules were atelectatic
(collapsed). These atelectatic lobules were associated with areas in which
nearly all of the bronchioles were severely involved or encircled with this
lymphocytic hyperplasia. These atelectatic areas were characterized by
collapsed alveolar walls and alveolar epithelial cell hyperplasia with a
few large macro phages in the narrow alveolar space. In most areas, alveolar
capillaries were still open and transporting erythrocytes. Very few neutrophils were in these areas; in fact they were difficult to find.
The lungworm nodules are similar to 72BHL-7 except there were more adults
present.
Cardiovascular System
No lesions were seen.

Lymphohemopoietic

System

The spleen appeared normal.
medulla of the gland.

The thymus had islets of eosinophils in the

Digestive System
Liver had several focal areas of a mononuclear cell infiltration.
maining digestive tract had no histologic lesions.
Musculoskeletal

System

No lesions were seen.
Urogenital System
No lesions were seen.

Endocrine and Central Nervous System
No lesions were seen.

The re-

�-133-

II.

PARASITOLOGY

Lamb Collections
During 1972, a total of seyen lambs were collected and examined to determine the cause of the pneumonia responsible for the severe mortality among
some herds in Colorado. Routine parasitology examinations were conducted
on all of these lambs to determine if ectoparasites and gastrointestinal
parasites were a contributing factor.

Ectoparasites
No ectoparasites were found on any of the lambs.
Gastrointestinal

Parasites

A few of the lambs had an extremely low level of infection with a"Marsha11agia
sp. Most of the lambs had a low level of infection with Eimeria spp. The
pattern and level of infection was not significantly different from the results obtained in 1971. For the species of Eimeria found in these animals,
please refer to the 1971 report.

Protostrongylus
The only species of Protostrongylus found in any of the lambs collected was
P. sti1esi (Table 2). As with last year, interpretation of the level of
infection is based upon the number of first stage larvae recovered from the
Baermann funnel. This, of course, furnished information relative to the level
of infection between lambs and between areas. Unfortunately, there is no
accurate method of determining the number of adult parasites.
Table 2. The number of first stage larvae ofP.stilesi
lungs of lambs collected in 1972.

recovered from the

Lamb

Area

Number of
Larvae Recovered

72BHL-2

Pikes Peak

3,062

72BHL-3

Pikes Peak

2,052

72BHL-4

Pikes Peak

4,623

72BHL-5

Pikes Peak

2,866

72BHL-6

Saguache

282

72BHL-7

Saguache

326

72BHL-8

Buffalo Peaks

305

�-134-

The results of this year's collection indicate that a much higher level
of infection with P. stilesi exists among animals on Pikes Peak as compared to Saguache
Buffalo Peaks. Comparisons with 1971 must be interpreted with caution because lambs were collected much earlier this
year on Pikes Peak; thus the female parasites had not yet reached their
full reproductive capacity. See the section on transplacental transmission for an interpretation of the reasons for such a high level of
infection in Pikes Peak.

or

III.

BACTERIOLOGY

Collected Lambs
The procedure for the isolation and identification of bacteria found in
bighorn sheep lambs has been outlined in the 1971-72 annual report. The
same procedure was used this past year. The bacteriology results for lambs
collected during 1972-73 are summarized separately in tabular form.
The most significant findings in the collected lambs were the presence of
Pasteurella hemolytica, Pasteurella multocida and a Neisseria sp. deep down
in the respiratory system of the lambs collected on Pikes Peak. These lambs,
of course, had heavy prenatal lungworm burdens and severe pneumonia. The
lambs collected from Saguache were healthy and had low levels of lungworm.
One of these lambs (72BHL-6) did not have Pasteurella and the other (72BHL-7)
had Pasteurella only in the upper respiratory tract. The last lamb was
collected on Buffalo Peaks and had a slight pneumonia. As expected, P.
hemolytica was distributed throughout the respiratory system.
Meaningful interpretations are difficult because so few control lambs were
examined, but the discovery of heavy lungworm burdens as a result of transplacental transmission of f. stilesi to lambs born on Pikes Peak indicates
that this overwhelming burden of parasites predisposes the lamb to infection
with Pasteurella and Neisseria, both of which are opportunistic organisms.
Either one, or both, are quite capable of causing pneumonia, but more often
than not they are both secondary invaders.

�-135-

(J)

c

SUMMARY
72BHL-2

Right apical lobe

+

Right cardiac 10be
~~+----~--+_--.+_--+--~--+~--_+--_+.--_4'------~--~--+_-----------------_
+
Right diaphragmat i c lobe
+
~_e_f_t_~ardiaclOb~e~
Left diar:&gt;hrai&lt;tllatic
lobe

+-_+
__-4

lobe

+

Left apical lobe

+

Intermediate

~ __~---_+---~--~--4---_+-----~I--_+---+--------------------_
i

i

+

+

I

+

Bronchus - E_~lt'::._~s-=i~d=-e
~-+--_+----+-+--~--+--+--_+-+--+-+--~--+
__--_+--_+--~-----I
+
~r~x
+
+
+

+

Nasal cavity

+
+

Nasophar~x

+

Trachea
Mediastinal

lymph node

Pharyngeal

lymph node

+

Gram + rods

+
+

Urine
Spleen

+

Kidney - right
Thymus
Liv '
Duodenum

+

Jejunum

+

Ileum

+

Cecum

+

Colon

+

+
+
Gram + rod~

+

Pleura - right

+

Abomasl!ID
Pericardial

+

+

Peritoneum

fluid

+

Blood - heart

+

Conjunctiva

+

Pleura - left
Lateral ventricle

+
+

Gram - rods

_

�-136-

...•

III

SUMMARY
72BHL-3

~as

~C)
CII ••••

•••••
;:I &gt;QI~

••0

: m

Po.c

Right apical lobe
Right cardiac lobe
Right diaphragmatic

Intermediate

...•

El

Q,

.

III~

III

III

C)~
tJO
oEl
C)CII
o.c

~
~CII

-..4as

"";:I

""CII

CII

III
III

•• ...•
III

III

Po

CII

l2:

+
+

+
+

+
+

+
+

+

+
+

+
+

+
+
+

;:1&gt;-

••
Q,III

III
Q,

III

•••

C)
C)

Q,

~~

0

III

§
...•

III

'tl

0
Q,

CII.c

•••Q,

~~

en&lt;

•••
•••
•••

III

.c

~
~

CII

Q,

;:I

C)

~0

&gt;.c
Q,
III

~

en

;:I

~
~
-..4
C)

III
I'Q

Q,
III

~
-..4•....

§
-..4

CII

••

~
C)

as III

-..4

.c

~
""
III

CII

0

~

u

•••
CII

CII

C)

as

~g:, 11~
c::

""
0
o &gt;UQ,

•••

0
U

III
CII
CII

110
0

•••
CII
III

•••
CII

••
C)

III

.c

0

•••

~

5

0

12
:5

lobe

Left cardiac lobe
Left diaphragmatic

Q,

~
...•

Q,

IIItJ

. c::

CII

.

C)

0

~
~;:I

UI

d

III
'tl

lobe

lobe

Left apical lobe
Bronchus - right side

+

Larynx
Nasal cavity
Nasopharynx
Trachea
Mediastinal

lymph node

Pharyngeal

lymph node

+
+

+
+

Urine
Spleen

+

Kidnev - riltht
Thymus
Liv~-Duodenum
Je1unum
Ileum
Cecum

+

Colon

+
+
+
+
+

+
+
+
+
+

Gram

+ rods

+

Pleura - right
Peritoneum
Abomasum
Pericardial

+
+
+

fluid

Blood - heart
Conjunctiva

Gram - rods

Pleura - left
Lateral ventricle

+

+

+
(Continued)

�-137-

III

e
QI
~
...•
~
••••
~
QI

\IS.
"0-

-..4

U
0
4.1

....•
::l
S

SUMMARY

72BHL-3
Continued

t\I
•.•.•
co
•.•.•U

QI-..4
~

4.1

::l &gt;-

QI •••.•
4.10

: m

Po&lt;..c::

Diaphragmatic
Bronchus

co
....•
....•
QI
~
::l
QI

4.1
10
00

Po&lt;

e,

.

Ilo
10
111

-..4

~
QI
III
III

-..4

QI

Z

nodule

+

- left 1unR

+

Pharynx
Left bronchial
Bronchus
Abdominal

fluid

RiKht diaphra2matic

nodule

lymph node

Mesenteric
Left kidney
Posterior

nasal cavity

Bronchus -left
Tracheal

diaphragmatic

bifurcation

Bronchus - riRht apical
Bronchus

- left

Lateral ventricle
Bronchus

- distal diaphra2matic

Bronchus - distal riRht diaphra2matic
RiRht frontal sinus exudate
Let.. frontal sinus exudate
Left diaphragmatic

nodule

Rbht

diaphraRmatic

abscess

Rbht

intermediate

lobe

Diaphragmatic

lobe bronchus

Diaphragmatic

med. bronchus

Right cardiac

lobe

Thorax
Retropbar)'DAeal lymph node

+

Ilo
IOU

-..4

qI
..c::

1114.1

Ilo

::l&gt;U •.•.•
uO

oS
uQI
o..c::

4.1
1lo00

~
~
....•
0
Ilo

••• p..

4.1 •••.•

III

::l

U
U

0

~
~

. §

Ilo
10

U

0
....•

&gt;..c::

e,

QI..c::

en&lt;

10

00
4.1

en

~
~
QI

co
.0

4.1
U
111

-..4

4.1
U

4.1
U

:::I

4.1
10
0

~

QI

c:
QI
co
0

-..4

as III
.0 QI
QI c:

III

~
~
QI

-..4

...•
U
111
I:Q

III

Ilo

CD

::l
....•
....•

ci.

~
QI
co

~
QI

~:o ~ .g~

....•

••• 0

u

Ullo

o

&gt;-

•••

0
U

~

~
0
~
:::&gt;

�-138,

I::
Q)

"Cl

....•
U

41 •••.•

•...
~;::I
e
&lt;IS
~
~41

SUMMARY
72BHL-4

;::I &gt;.

;::I

Q)~

41
•...

~ ~

III

~

&lt;IS

&lt;IS

~

t)

•... •...
•..•
0
A...c

Ri.ght apical lobe

•...

&lt;IS
A..

c:&gt;.
en ....•
U
c:&gt;.
III

&lt;IS
....•

•...

41
III
III

....•
Q)

Z

Ill ••.•

;::I&gt;.

u~
UO

oS

UQ)
o.c

•...

c:&gt;.&lt;Il
Q).c
•..• c:&gt;.

•...~

en":

....•

e,

&lt;IS
.c

c,
•...
0

~~
0

c:&gt;.

m
....•
~

•...
""'•...
Q)
c,

en
en

;::I
U
U
0
U
0
&gt;.
.c

c:&gt;.
III

§
....•

en

"Cl

•...

c,

~
~
....•

•...

U
&lt;IS

~

&lt;Il

en

;::I

!Xl

....•

•...

c:&gt;.
III

§

Q)

I::
41
bI)
0

§
....•

•...

•...

&lt;IS

41
•...

41
•...

•...

U
&lt;IS

41
•...

•...

U
&lt;IS III

..0
41

41
I::

..0

~~

Qj

•..•
0

~
•...

u

U

u

~

Ul

....•

0

en

o &gt;.

c:&gt;.

0

41

U
&lt;IS

..0

0
•...

..:
41

~
0
~r::
:J

+

+

+

+

+

Right cardiac lobe

.

bI)
I::

0

Right diaphragmatic

en

&lt;IS

+

lobe

Left cardiac lobe
Left diaphragmatic
Intermediate

lobe

lobe

+

+

+

+

+

+

+

+

+

+

+

+

+

+

Left apical lobe

+

Bronchus - right side

+

+
+

Nasopharynx

+
+

+

+

+

+

Trachea

+

+

+

+

+

Gram + rods

Larynx
and exudate

Nasal cavity

Mediast"inal lymph node
Pharyngeal

lymph node

Urine

+

Spleen
Kidnev - dght

Gram - and + rods

+

ThYJlluS
Liv2"'
Duodenum

+

Je1unum

+

Ileum

+

+

Cecum

+

+

Colon

Gram + rods

+
+
+

Gram + rods

+
+

Gram + rods

+

+

Gram + rods

+

+

Gram + rods

Pleura - dght
Peritoneum
Abomasum
Pericardial

+

fluid

Blood - heart

+

+

+
+

Conjunctiva
Pleura - left
Lateral ventricle
(Continued)

�-139-

III

~
Q)
00
~
....•

t1I

'tl
....•

.

..-t

~u

•...

SUMMARY
72BHL-4
Continued

::l
S
t1I
..-t t1I
..-t&lt;J
Q) •••.•

,.. •...
::l &gt;-

Q)..-t
•..•
0
tD S
t1I Q)
Il&lt;.c:

Diaphra~atic

u
0

t1I
..-t
..-t
Q)

,..::l

Q)

•...
III
t1I

~

.

p.
III

t1I
....•

,..

Q)
III
III

....•

Q)

Z

....•

Ill •..•

::l&gt;-

u..-t
uO
oS
uQ)
o.c:

•...

p.t1I
Q).c:
,..p.
•..•..-t
tIl~

e,
tD

t1I
.c:

,..

P.

tD

::l

u
U

.

P.

,..
.•..
,.. §
....•
P.
,..
Q)

0

tD

§
....•

0

III

&gt;-

::l

-e
....•

.c:

..-t
..-t

~
....•

....•

tD

t1I

•...

:E

U
t1I

tIl

I'Q

~

..-t
0
p.

u

..-t

e,

,..
•...

Q)
•...
t1I tD
.0 Q)
Q) ~

o

p.
III

§

....•

,..

Q)
•...
u
t1I
.0
Q)

~g:, ,..~
o &gt;0

0
..-t

,.. 0

o

()P.

()

III
Q)

~
Q)
00

,..
0

Q)
t1I

,..

Q)
•...
u
t1I
.0
0

,..

Q)

~

50

.e

~
=&gt;

nodule

Bronchus - left lung
Pharynx
Left bronchial
Bronchus
Abdominal

fluid

Right diaphragmatic
Mesenteric

nodule

+

lymph node

Grar - rod

Left kidney
Posterior nasal cavity
Bronchus - left diaphra2matic

+

+

+

+

+

Tracheal bifurcation
Bronchus - ri2ht apical
Bronchus - left
Lateral ventricle
Bronchus - distal diaphralllllatic
Bronchus - distal right diaDhra2matic
Right frontal sinus exudate
Leh

frontal sinus exudate

+

+

Left diaphra2matic

nodule

Ri2ht diaphra~tic

abscess

+

lobe

+

+

Diaphragmatic

lobe bronchus

+

+

+

+

+

Diaphra2matic

med. bronchus

+

+

+

+

+

R12ht intermediate

Right cardiac lobe

+

+

Thorax
Retropharvn2eal

+

lymph node

+

+

Grar + rod

�-142-

&lt;IJ
s::
Q)

&lt;1S

....•
"d

0

SUMMARY
72BHL-6
&lt;1S

M
&lt;1S
M
U
Q) ••..•

,... •...
::s »
Q)M

•..•
0

~ m

p...s::

•...

p.

M
::s

COU
-rl

.

s

Po

&lt;1S

I/)

.-l
.-l
Q)

,...

::s
Q)
•...
&lt;IJ

&lt;1S

p..

&lt;U
....•

,...

Q)
I/)

co
....•
Q)

Z

1/) ••.•

::s»
U.-l
uo
oS
U Q)
o.s::

•...

p.&lt;1S

I/)

&lt;1S

,...Po

en&lt;

&lt;IJ

Po

Q)

Po

::s

,...

o
U

Po

~

0

I/)

0
c,

.-l
»
.s::

0

co

0

U

.-l

Q).s::

•... .-;

s::
....•
,...
.....
,...

Po
.s::

~
:E:

p.
&lt;1S

•...

en

.

eo

.

U

::s
.-l
M

....•
U

&lt;1S
!Xl

p.

I/)

§

0
M

U

&lt;1S

Q)

•...

,...

U

Q)
•...

&lt;U co

&lt;IJ

,...

Q)

,...

•...

,...

ee
0
,...

....•

§

•...

§

....•

-e

....•
....•

&lt;IJ

Q)
s::
Q)

U

.0 Q)
Q) s::

~g:,
,...0
o

»

UPo

&lt;U

.0
Q)

~
,...
0
U

Q)

U

&lt;U

.0

0
,...
Q)

&lt;

~
0
]
s::

::&gt;

Ri,ght apical lobe
Ri,ght cardiac lobe

Gram + rods

R:!&amp;ht diaphra2Dlatic lobe

Gram + rods

Left cardiac lobe
Left d Iaphr agraatLc lobe
Intermediate

lobe

Left apical lobe
Bronchus - right side
Larynx
Nasal cavity

+

Nasopharynx

+

Trachea
Mediastinal

lymph node

Pharyngeal

lymph node

Urine
Spleen

+

+

Kidney - rbht
Thymus
Livc:r

+

Duodenum
Jejunum

+

+

Ileum

+

Cecum

+

+
+

Colon

+

+

+

+

Pleura - right
Peritoneum
Abomasum
Per icardial fluid
Blood - heart
Conjunctiva

+

Pleura - left
Lateral ventricle

(Continued)

�-143-

GI

~
~

as

"t:I

""

•..

SUMMARY
72BHL-6
Continued

GlO

a

GI'"

as
M
M
QI

(11M

""::l
Q)

•..

•.•0

~ m

Il-o..c:

Bronchus

M

M as
M 0
QI""

""::l &gt;-

Diaphra£matic

c:lo

::l

as

...
GI

as

Il-o

e,.

.

0
0

"c:lo
GI

as

""""

QI
GI
GI

""z

Q)

""
::l&gt;OM
00
ot:!
0Q)
o..c:

...c:loas

Q)..c:

""c:lo

",M
CIl&lt;

as

..c:

e,

""

~

M

0

c:lo

~

""

:E

""

GI

'101
""

GI

QI
""e,

::l

0
0
0
0
0
M

&gt;-

..c:

e,

e,
GI
GI

::l

M
M

as
""as
•..
0

CIl

pC!

nodule

- left lung

Pharynx
Left bronchial
Bronchus
Abdominal

+

fluid

Right diaphra2matic

nodule

lymph node

Mesenteric
Left kidney
Posterior

nasal cavitv

Bronchus - left diaphraRmatic
Tracheal

bifurcation

Bronchus - riltht apical
Bronchus - left
Lateral ventricle
Bronchus

- distal diaohra2m8tic

Bronchus - distal ri2ht diaphra2matic
Ri2ht frontal sinus exudate
Leh

frontal sinus exudate

Left df.aohraRmS tic nodule'
Ri2ht diaphra2m8tic

abscess

Ri2ht intermediate

lobe

Diaphra2matic

lobe bronchus

DiaphraRmatic

med. bronchus

Riaht cardiac

lobe

Thorax
Retropharyngeal

lymph node

e,

§

""~

GI

~

~

""""

""""

...

...

Q)

QI

0

0

as

GI

...""

.oQ)

U

Uc:lo

GI
0
M

as

.0

GI
QI

c::
QI
co
0

""QI
as

...""

Q)
0

~!~

.gas

~
0

""
o 0
&gt;-

""QI
&lt;

~
::&gt;

""0
U

�-144-

d

"0
..-4
tJ
0
,..,j

00
d

CII

•..

SUMMARY
72BHL-7

P.

1-0

OJ.(
III
IIltJ
CIS
1-0.
;::j
..-4 ,c
III
CII §
P.
;::j
S
Ill'"
P.
..-4
P.
;::j&gt;"
1-0
tJ
1-0
tJ,..,j
eo
CIS
III
tJ
P.
0
,..,j
,..,j
co
CII
tJO
§
0
III
,..,j
tJ ,..,j eo
~ tJ III ..-4 tJ
,..,j
CII..-4 CII ..-4 oS
tJ
CII
0
"0
CIS
1-0 •••
,..,j ;::j ..-4 ,J:l III
1-0
1-0
o,c
0
CII
::l &gt;..
;::j
,..,j
p.
CII p.eo
&gt;..
CIId
CII,..,j
,c ,..,j 1-0
CII
Ul
•.•0
CII,c
p.
ell
..-4
&lt;Il
..-4 1-oC&gt;&lt;Il
tJ 0
m eo eo
CIS
..-4
~ m Ilo.
CII •.•..-i
1) ~.
Ilo.,c
en&lt;
Z
l::
en
up.
~ M
u

Right

apical

lobe

Right

cardiac

lobe

Right

diaphragmatic

Left

cardiac

Left

d Laphr azmat Lc

.

.

Intermediate

lobe
lobe

•..

...

•.. ~g:,

•..

II)

§

CII
CII

d

00
0

..-4 1-0CII
1-0
eo
CII 1-0
tJ
CII
co
,J:l
tJ
CII eo
~
,J:l
0

•..
~

1-0

0

u

•..
0

CII ~d

1-0

&lt;

~

+
+
lobe

+

lobe

Left

Bronchus

Ul

..-4

P.

p.

•..

apical

Ul

eo

+

lobe

+

+

+

+

- riJ;1;ht side

Gram

- rods

Unknowns

+

Larynx
Nasal

cavity

NasoPharvnx

+

+

Trachea

+

+

Mediastinal

lymph

node

PharvnJ;1;eal lymph

node

Lots

of unknowns

Gram

- and + rods

Gram

+ rods

Gram

- and + rods

Gram

- rods

Urine
Spleen
Kidney

- right

+

Thymus
I

Liv·.~'
Duodenum

+ +
+ +

Jeiunum

+

Ileum

+

Cecum

+

+ +
+

+

+

Colon
Pleura

- right

Peritoneum

+

Abomasum

+

Pericardia1
Blood

fluid

.

- heart

Conjunctiva
Pleura

- left

Lateral

ventricle

+

+

(Continued)

�-145-

III
s::
QI

l'\l

"0

..-I
0
U

~
:::s
a
l'\l
l'\l
~ l'\l
~
~
U
~&lt;II
QI ••.••

SUMMARY

72BHL-7
Continued

I-&lt; U

:::s

&lt;II~

&gt;.

Diaphragmatic
Bronchus

CIl

U~

.....

uO

a

I-&lt;

I-&lt;

o
UQI
o..c

&lt;II

&lt;II
CIl

U
p.l'\l

:::s

••• 0

U

III
l'\l QI
p....c

p..

a

P.

.....
OOU
:::S&gt;.
ooU

l'\l

CIl

l'\l

rn
.....
&lt;II

z

s::
..-II-&lt;

P.

P.

QI..c

I-&lt;p.

u~
tI'l&lt;

00

•••
I-&lt;

00

QI

l'\l

..c

e,

...

~~

0
P.

l'\l

..-Ia
~

:::s

.

U
U

P.

0

III

U

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..c

p.
l'\l
U

tI'l

.

00

.

U

00

:::s

~
~
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U

l'\l
c:Q

p.

§
..-I
"0
..-I

I-&lt;
U
CIl

~0

U

P.
III

§
..-II-&lt;
QI

§
..-II-&lt;
QI

U

U

U
III CIl
,&amp;J QI
QI s::

,&amp;J
QI

~~
I-&lt; 0

o &gt;.

up.

U

III
QI

s::

QI

00
0
I-&lt;
QI
l'\l

I-&lt;
QI

III

U

e
&gt;.

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0
u

U

l'\l
0
I-&lt;
&lt;II

&lt;:

~
0

.e

s::

~

nodule

- left lunlt

PharYnx
Left bronchial
Bronchus
Abdominal

fluid

Ri2ht diaohramatic
Mesenteric

+

nodule

+
+

lvmoh node

+
+

Left kidnev
Posterior

nasal cavity

Bronchus

- left diaphraltDlatic

Tracheal

bifurcation

Bronchus

- riltht apical

Bronchus

- left

Lateral

ventricle

Bronchus

- distal diaphragmatic

Bronchus

- distal right diaphragmatic

+

+

+
+

+
+

+

Riltht frontal sinus exudate
Left frontal

sinus exudate

Left df.anhragma tic nodule
Riltht diaphragmatic

abscess

Ri2ht intermediate

lobe

Diaphragmatic

lobe bronchus

DiaphraltDlatic med. bronchus
Riaht cardiac

lobe

Thorax
Rat ronharvnzeaj, lymph node

+

Gr Pn - rods

�en
s::

-146-

co

Q)
00

"0

~

.
c,

U
0

•..

...-f
;:J

SUMMARY
72BHL-8

e

co

...-f ."
...-f U
Q)~

"";:J

•..
e-,

Q)...-f
•.• 0

~ ~

Il..o!:

co
...-f
...-f
Q)

"";:J

Q)
•...

en

."

Il..

III U
c:l.

en

co

~

""Q)
III
III

~
en •.•
;:J&gt;.
U...-f
U 0

o
e
UQ)

•..

Oo!:

~Q)

c:l.'"
Q)o!:
).jc:l.
•.•...-f

Z

CIl&lt;

s::

co
e,

o!:
).j
0

~
...-f
0
e,

~~
~

.....""

III

en
;:J
U
U
0
U
0
...-f
&gt;.
o!:

c,
co
•...

CIl

""Q)
c,

e,
en
en

;:J
...-f
...-f

~
U

~."

~3
"0
~

c:l.
III

§

§

~
""Q)
•...

~

•..
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U

co en

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Q) s::

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en

0
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U

en
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e,

o 0&gt;.

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U

Po.

U

."

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Q)

~
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U

Q)
00
0

""Q)

co

""
•..
Q)
U

."

.0
0

""Q)

&lt;

5

0

]

s::
::&gt;

Right apical lobe
,

+

Right cardiac lobe
Right diaphra2ll1atic lobe
Left cardiac lobe

+

+

Left d Laphr azmat ic lobe

+

+

Intermediate

lobe

Left apical lobe

+
+

Bronchus - right side
Larynx
Nasal cavity

+

Nasopharynx

+

+

Trachea
Mediastinal

lymph node

Pharyngeal

lymph node

+

Urine
Spleen
Kidn~

+
- right

+

+

T~us
I

Liv2-':

+
+

+

Ileum

+

+

Cecum

+

+

Colon

+

+

Duodenum
Jeiunum

+

Pleura - ri2ht
Peritoneum

+

Abomasum
Pericardial

fluid

Blood - heart

Gram - rods

+

+

Conjunctiva
Pleura - left
Lateral ventricle
(Continued)

�-147-

III

c::

oS

Q)

""

~

.

0

M

SUMMARY

:::l

a

72BHL-8
Continued

."
M ."

M 0
Q) ••.••
"" .j..I

;:I &gt;-.

Q/M
.j..I 0
III
." Q)
p....c::

a

Diaphragmatic

oS
M
M
Q)

"":::l

Q)
.j..I
III
oS
p..

.

p..
III

oS

.....

""Q)
III

III
.....
Q)

z

!E'o
.....
1II.j..1

;:1&gt;-.
C)M
C)O

oa
OQ/

o..c::
.j..I
p..oS
Q)..c::
""p..
.j..IM
00&lt;

nodule

+

Bronchus - left luna
Pharynx
Left bronchial
Bronchus
Abdominal

fluid

Riaht diaDhraamatic
Mesenteric

nodule

+

Lvmoh node

Left kidney
Posterior nasal cavity
Bronchus - left diaphraamatic
Tracheal bifurcation

+

Bronchus - riaht aDical

+
+

Bronchus - left
Lateral ventricle

+

+

Bronchus - distal diaDhra~tic

+

+

Bronchus - distal riaht diaDhragmatic
Ri2ht frontal sinus exudate
Lei.. frontal sinus exudate

+

+

+

+

Left ddaphr aama tic nodule
Rbht

diaDhraamatic

abscess

Rbht

intermediate

lobe

Diaphra~tic

lobe bronchus

Diaphra~atic

med. bronchus

Riaht cardiac lobe
Thorax
RetroDharyngeal

lymph node

Bronchus - dorsal right dia~hragmatic

+
+

~~
.•...
""

c,

0
.j..I

III
oS
..c::

e,
""

III

;:I
0
0

.

p..
III

~
M

0
0

III

0
e,

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;:I

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M
M

~
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0

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oS
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00

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p..

s

~
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~

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M

0

IQ

0

U

p..
III

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""Q)
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0
." III
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0

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o 0
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up..

III
Q)

c::
Q)
co
0

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0

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0

50
~

P

�..;.148-

IV.

VIROLOGY

Lamb Collections
The seven bighorn sheep lambs collected during the summer of 1972 were
examined for possible infection with, or exposure to, 5 different viruses,
including Parainfluenza-3 (PI-3), Infectious Bovine Rhinotracheitis (IBR),
Bovine Viral Diarrhea (BVD), Bluetongue (BT), and the Parvoviruses. The
serological results are presented in tabular form (Table 3).
PI-3 was isolated from 72BHL-2, 72BHL-4, 72BHL-5, and 72BHL-8. PI-3 was
not isolated from 72BHL-3, 72BHL-6 and 72BHL-7. No other virus was isolated
from these animals.
PI-3 is a ubiquitous virus and it is not at all unusual to find a titer in
most animal~ asa result of infection; undoubtedly it does contribute to
the overall picture of pneumonia. However, present evidence indicates that
this organism alone does not induce the serious pneumonia responsible for
such high lamb mortality.
In 1971 and in 1972, some of the lambs were examined for serological evidence
of infection with adenoviruses, many of which are capable of producing respiratory infection. Some of the lambs from both years had rather high titers to
adenovirus, but the significance of this finding has not yet been determined
(see Table 3).

�-149-

Table 3. Serology for evidence of virus infection in
Colorado bighorn sheep.
Lamb
Number

PI-3

IBR

BVD

71BHL- 5

2

0

0

0

128

71BHL- 6

8

0

0

0

16

71BHL- 7

8

0

71BHL-17

2

0

0

0

64

71BHL-18

64

0

0

0

32

71BHL-19

8

0

0

0

128

71BHV-20

4

0

0

0

128

71BHL-21

0

0

0

0

George

8

0

64

512

George

0

ND

32

8

George

2

NA

NA

NA

George

0

0

16

&gt;64

32

George

0

0

16

16

72BHL- 2

16

0

0

0

16

72BHL- 3

0

0

0

0

64

72BHL- 4

4

0

0

0

64

72BHL- 5

4

0

0

0

32

72BHL- 6

0

0

0

0

16

72BHL- 7

0

0

0

0

0

72BHL- 8

4-

0

0

0

0

BT

Parvovirus

8

&gt;64

�-150-

V.

MYCOPLASMA

Lamb Collections
Four of the seven lambs collected in 1972 were examined for evidence of
Mycoplasma infection by the egg-inoculation technique. All four were
nesative. Mycoplasma spp. are capable of producing respiratory disease
in some animals, but this examination was routinely conducted to incriminate or eliminate this as a possibility.

SEGMENT OBJECTIVE NUMBER 2
Perform complete necropsy examination of animals experimentally infected
with bacteria, virus and lungworms.

PROCEDURE
Previously uninfected animals are to be inoculated with bacteria, virus
and lungworms and then examined at postmortem when clinical signs of
infection appear.

RESULTS
Two male lambs obtained from the Denver Zoo were not infected with lungworm,
and did not have a titer to PI-3 virus, but one had Pasteurella in the upper
respiratory tract. In view of the numbers, the research team decided to
challenge these animals with PI-3 to determine if the virus alone, or in
combination with Pasteurella but in the absence of lungworm would induce
pneumonia. Two days after inoculation both animals developed a slight nasal
exudate, but recovered without any noticeable serious effects from PI-3.
Both then developed a substantial titer to this v:i.rus.·
Experimental infections with Protostrongylus were conducted on sheep at the
Little Hills Experiment Station. Following the discovery of transplacental
transmission, the emphasis on experimental infections was shifted to favor
this approach. These results are given partially under Segment Objectives
4 and 8.
No attempts have been made to inoculate lambs with Pasteurella or Neisseria;
both seem to be ubiquitous inhabitants of the respiratory tract and the
logistics involved in checking an animal for the presence or absence of
these bacteria was just recently worked out to the satisfaction of the research team.

SEGMENT OBJECTIVE NUMBER 3
Examine the 7 bighorn sheep for the incidence of infection and level of infection with ectoparasites and endoparasites and evaluate the significance

�-151-

of these parasites on the health and well-being of the animal.

PROCEDURE
Survey of the incidence and level of infection with parasites will be
accomplished by:
a.

Following examination by the pathologist, the sheep listed
under Item 3 above will be examined for any evidence of ectoparasites.

b.

The lungs will be opened, the parasites removed and identified.
To facilitate recovery, the lungs will also be Baermannized.
Assessment of the severity of infection will be done in cooperation with the pathologist.

c.

The parasites in the abomasum, small intestine and large
intestine will be washed out and the total counted by species.
RESULTS

See under Segment Objection Number 1, section II, lamb collections for
parasitology examination.

SEGMENT OBJECTIVE NUMBER 4
Elucidate the bioloKical cycle of Protostrongylus.
PROCEDURE
Elucidate the biological cycle of Protostrongylus. Land snails have been
colonized in the laboratory, and successfully infected with the first stage
larva of Protostrongylus.
Snails with infective larvae will be used to
experimentally infect domestic sheep, bighorn sheep and domestic goats.
Domestic sheep and goats will be purchased locally, and bighorn sheep will
be obtained as lambs from the Little Hills herd and at zoos throughout the
state. These animals will be infected by introducing infected snails orally.
Some of the domestic sheep will be given immunosuppressants to enhance
susceptibility to the parasite.
RESULTS
Following discovery that Protostrongylus was transmitted transplacentally
to the lambs, all efforts on the biological cycle were directed towards
proving this portion of the cycle because present evidence strongly indicates
that transplacental infection predisposes the lambs to the pneumonia that
follows. Thus, elucidation of the biological cycle, and experimental infection of lambs by feeding infected snails would be purely academic.

�-152-

The basic life cycle of lungworms was completed by Ruth Monson in 1971.
She completed the life cycle in mouflon-bighorn crossbred sheep at
Rachelwood Wildlife Research Preserve, New Florence, Pennsylvania. There
are two species of Protostrongylinlungworms
in bighorn sheep, protostrongylus stilesi and Protostrongylus rushi. AdultProtostrorigylus
stilesi are found with the parenchyma of the lung andProtostrOrigylus
·rushi are found within the bronchi and bronchioles.
The adult lungworms mate in the lungs and an embryonated egg is produced
which develops and hatches within the lung. A first stage infective
larvae exits~the egg. It makes its way either actively or by coughing
and ciliate motion in the trachea to the pharynx. The larvae is swallowed
and passes unharmed through the gastrointestinal tract of the sheep. It
passes out of the body in fecal pellets during defecation. Development of
the first stage larvae is halted unless it is able to enter a suitable
intermediate host'and undergo several moults in a process of maturation.
Suitable intermediate hosts are any of several species of land molluscs,
all exceedingly small. After an appropriate period of maturation the
larvae within the foot tissues of the snail are third stage larvae and
are infective to bighorn sheep. No other mammal has been successfully infected with them. The snail is consumed incidentally with forage while the
sheep grazes. The infected snail enters the digestive tract of the final
host. It penetrates the alimentary mucosa and makes its way to the lungs
by way of the hepatic portal system, the liver and the posterior vena cava.
Within the lungs third stage larvae mature to adults, mate and renew the
life cycle. This basic life cycle has recently been supplemented by an
alternate route of migration indicated by recent work. That is, the isolation of third and fourth stage lungworm larvae from fetal and neonatal bighorn lamb tissues.
In transplacental transmission of lungworms the third stage larvae crosses
from the ewe through the placenta and into the fetus at some time in pregnancy.
Larval development in the fetus is halted after transmission and most of them
remain in the liver. After birth larvae complete migration to the lungs where
they mature, mate and renew the life cycle. Thus, a neonatal lamb is infected
with lungworms before it ever hits the ground. This method of infection has
previously been shown with Toxocara canis andAncylostomacariinum
in dogs and
with Neoascaris vitulorum in calves (Sprent 1954; Yutuc 1949).
There have been indications of transplacental transmission of lungworm larvae
for many years. Pillmore (1956a) collected lamb pellet group samples on the
Buffalo Peaks range in Colorado during June and July. One of four pellet
groups contained lungworm larvae on June 16, 1955. A sample collected June
22 was positive. Four samples collected July 26 and two of three samples
collected July 29 were positive for first stage lungworm larvae. Pillmore
(1959b) recalls a lamb that was born in captivity in Denver on June 1, 1940.
It was coughing, had a rough hair coat and breathed with difficulty when
exercised. On July 19 the lamb was found to have 1ungworms. The pen it
occupied was dry and there was minimal chance for presence of intermediate
host land snails. Another example he recounts occurred at Sybille Wildlife
Research Station in Wyoming. Six ewes had been brought to the station after
being trapped near Dubois in January 1956. Five of them produced lambs and
all of these began passing first stage larvae in July. Suitable intermediate

�-153-

hosts were found in the pasture. Another example is the lamb out of a
ewe caught at Glenn Eyrie, Colorado Springs in January, 1958. The lamb
began passing first stage larvae about a month after birth. It had been
a healthy, active lamb but three weeks after first passage of larvae it
died. A necropsy was performed and a very heavy lungworm infection of
the diaphragmatic lobes of the lungs was found. A similar example is a
lamb caught on the Buffalo Peaks range on May 23, 1958 when still too
young to follow its mother. It was penned at the Cheyenne Mountain Zoo
in Colorado Springs where bighorn sheep had not been kept for a number
of years. A lactating goat was used as a foster mother. On June 23 the
lamb began passing first stage lungworm larvae. Pillmore (1959b)"believed
these last two cases in particular indicated en utero infection of the
fetus with lungworms. Pillmore (1961) recall;-another lamb kept at the
Cheyenne Mountain Zoo. It died on July 7, 1959 from pneumonia and in
necropsy first stage Protostrongylin larvae were isolated. All of these
examples are strong indications of prenatal infection but none of them
are conclusive. Attempts to prove the mechanism were unsuccessful.
Pillmore (1959b) suggested early first stage larvae passage in lambs
could come from three sources. First they could reflect coprophagous habits
of lambs who might eat pellets of infected sheep. It should be considered
how many pellets a lamb would have to eat to produce a significant larvae
output in its own feces. Pellets eaten by a lamb would be diluted'by the
contents of its gastrointestinal tract. A lamb would have to eat a lot of
infected pellets to pass many first stage larvae. One of us (Lange), has
sat with lambs while they played, nibbled at vegetation and rested and have
observed very little of this behavior. A second explanation of early first
stage larvae passage is postnatal infection of lambs. While a lamb does
begin almost immediately after birth to nibble vegetation there is little
likelihood that he could consume sufficient numbers of infected snails to
produce the levels of larvae output that we have witnessed. One lamb from
Pikes Peak produced over 10,000 first stage larvae per gram of feces
(Hibler 1972). The third possibility Pillmore suggested is prenatal infection. He thought this mechanism the most likely. A fourth possibility
exists. This is transmammary transmission soon after birth. Third stage
infective larvae would cross from the ewe to the neonatal lamb through her
milk. This mechanism has been observed with Toxocara cati in cats
(Swerczek, Nielsen and Helmboldt 1971), with Ancylostoma-canium in dogs
(Stone and Girardeau 1966), with Toxocara canis in dogs (Stone and Girardeau
1967), with Strongyloides westeri in horses (Lyons, Drudge and Tolliver 1969),
and with Neoascaris vitulorum in cattle (Warren 1969). We have examined
mammary tissues of several post-parturient ewes and found no larvae but we
cannot rule out this possibility.
In May, 1972 a pregnant ewe in late gestation was delivered to the Colorado
State University Wildlife Disease Center by State Division of Wildlife personnel. Third stage lungworm larvae were isolated from liver, lung and
cotyledonary tissues of the fetus and placenta (Hibler, Lange and Metzger
1972). This was the first direct evidence of transplacental transmission of
protostrongylin lungworm larvae. Since then third or fourth stage lungworm
larvae have been recovered from six other fetal or immediately postnatal
lambs (Hibler et al. 1973). The numbers of larvae recovered have varied
from one to 20~-rhis
despite the fact that a fair proportion of the tissues

�-154-

were lost to histopathological, viral and bacteriological sampling. It
is apparent that prenatal infection does occur on the Pikes Peak range,
at least. It is more likely that this mechanism occurs on most Rocky
Mountain bighorn sheep range throughout western North America.
Lungworm larvae consumed during grazing by the ewe can cross the placenta
to infect the fetus. An important question is whether the ewe can accumulate
third stage larvae through the year or must larvae cross the placenta soon
after being consumed by the ewe. That is, does somatic storage of third
stage larvae exist in bighorn sheep? This is an important question. If
ewes can transmit larvae to the fetus only at some point in gestation and
are unable to store larvae they encountered at other times of the year they
will pass far fewer larvae to the fetus. However, if they can store larvae
all year long and then transmit them in a large migration to the fetus during
gestation a large dose will be delivered in a relatively short period. This
mechanism would make range density and availability of winter range vital
because year-round population density would influence the intensity of infection among intermediate and definitive hosts. Any changes to increase
density or decrease seasonal home range would add significantly to the dose
of third stage iarvae available to cross the placenta. Somatic storage has
been observed with the dog ascarid, Toxocara canis. Immature stages of
larvae of this species are stored in the muscles, central nervous system,
kidneys, liver, and other highly vascular organs (Webster 1958). They are
mobilized during the last two weeks of gestation at which time they cross
the placenta to infect the fetus.
PURPOSE
The objective is to elucidate the mechanisms of transplacental transmission.
First, can it be reproduced and second, can we demonstrate somatic storage
of third stage lungworm larvae.

METHODS AND MATERIALS
Two ewes were selected at Little Hills Game Experiment Station near Meeker,
Colorado to test somatic storage of third stage protostrongylin lungworm
larvae. Ewe I received 754 and Ewe II 760 mature third stage larvae (see
Table 4). The intermediate host snails were cultured and infected in the
laboratory at Colorado State University. Larvae were permitted to mature
in the snails, Vallonia pulchella andcyclophorella
for known periods of
time (Table 4). Periodically, infected snails were transported to Little
Hills where they were administered to the test sheep. Infected snails were
placed in number one gelatin capsules and these were secreted in alfalfa
cubes that had been hollowed out. The alfalfa cubes could be fed by hand
to the test sheep in most cases. Administrations were made by Steve
Steinert, Research Assistant at Little Hills, and by the author. Total
dosages to these two ewes were completed before breeding season. The test
was set-up so lambs of these two ewes, to pass first stage larvae during the
following summer, could only do so through the mechanism of somatic storage
of administered third stage larvae and prenatal infection of the fetus with
them.

�-155-

In the second portion of the study two other ewes, Ewe III and Ewe IV,
were selected for infection postbreeding. These ewes were administered
third stage larvae between February and late April (Table 5). Ewe III
received 1643 larvae and Ewe IV received 1489. Each of the larvae matured
in snails for a specific amount of time. Lambs of these two ewes would
demonstrate direct transplacental transmission if somatic storage did not
exist. Lambs of these two ewes could not rule out somatic storage though
if they did pass first stage larvae. Only lambs born to Ewes I and
II could do that. The ram in the herd at Little Hills would serve as a
control because he never had a patent lungworm infection.
There are two questions that will occur to the reader. How do we know that
ours are the only third stage larvae the ewe might encounter and how do we
know no somatically stored larvae from years past in the ewes could cloud
our results? First, it unlikely there is completion of the life cycle
through intermediate hosts on this range. The range at Little Hills is
semiarid and is composed of a pinon-juniper association on rocky hills
with cliffs and sagebrush bottoms. I have surveyed the range at Little
Hills and have found four species of land snails. Two of these species are
suitable intermediate hosts. However, after two years of searching I have
been unable to find live specimens of either of these snails, Pupilla blandi
or Vallonia cyclophorella. I find older, bleached shells. Obviously, there
are live intermediate hosts present but they must be scarce. On every other
bighorn range I have examined, including Pikes Peak, Buffalo Peaks, Ramparts
Range, Trickle Mountain, and the Pecos Wilderness and Red River area in New
Mexico, I have been able to find suitable intermediate hosts. Little Hills
is a dry range and seems to be in a dry cycle. It is not extraordinary to
find an area where the life cycle is not completed. The Arkansas Canyon herd
between Cotapaxi and Royal Gorge was entirely free of lungworms in the 1950's
(Pillmore 1958). Furthermore, the sheep at Little Hills are primarily browsers.
They consume mostly mountain mahogany, serviceberry and bitterbrush, thus not
being exposed often to surface feeding when they might consume a snail. Finally,
we have monitored larvae putput of each of the sheep at Little Hills since June,
1971. This has been done by collecting pellet group samples several times per
month and baermannizing them. One of these sheep is a ram who has no lungworm
infection and presumably no resistance to infection. He has never had a lungworm infection and he has not picked one up at Little Hills. He spends his
time with the other sheep. If infected intermediate host snails were available for consumption by the sheep the ram should pick them ~p and develop a
patent infection. We may be fairly certain, though not conclusively, that
artificial dosages of lungworm larvae are the only ones the sheep have encountered at Little Hills. The second question is whether the lungworms of
the lambs could be traced to somatic storage of larvae acquired before the
sheep were transplanted to Little Hills. The answer is probably not but it
doesn't matter. Ewe I bore a lamb on this range two years ago which passed no
more than a trace of first stage larvae for 18 months (Table 4, Ewe II). If
she had any third stage larvae in somatic storage they should have passed to
her lamb two years ago, and that lamb's larvae output should have reflected
the event. Apparently, she had few third stage larvae to transmit to her
fetus. The second test ewe, Ewe II, used to test somatic storage was the lamb
born to Ewe I in 1971. She was 18 months old at breeding and 24 months old
at parturition. She has been on the Little: Hills: range all her life and it is
unlikely she has been exposed to infection there. In Ewes III and IV we are
not testing somatic storage. Though some few larvae may derive from earlier
somatic storage of third stage larvae it does not affect our objective of
showing transplacental transmission with these two ewes.

�-156-

Table 4. Preconception doses of third stage Protostrongylus ~.
administered to two ewes at Little Hills.

Ewe I (Mom)

larvae

Ewe II (Lamb)

Date

Larvae

Snails

Days in
Snails

7-15-72

15

12

over 40

9-23-72

200

148

46

11-11-72

193

110

39

11-11-72

132

69

48

11~11-72

214

89

45

Larvae

Snails

Days in
Snails

124

95

46

11-11-72

236

149

35

11-11-72

33

31

36

1.1-11-72

367

118

47

760

393

TOTALS

754

428

Table 5. Postconception doses of third stage Protostrongylus ~.
administered to two ewes at Little Hills.

Ewe III ~Boss)
Date

Larvae

Snails

Days in
Snails

2-10-73

68

47

over 61

2-24-73

163

108

53

4-7-73

659

225

46

larvae

Ewe IV (Smok:~~)
Days in
Larvae
Snails
Snails

189

91

46

4-8-73

180

75

47

4-9-73

197

75

48

387

110

49

4-20-73

330

100

50

4-21-73

206

57

51

1489

508

4-19-73

TOTALS

753

1643

225

605

49

�-157-

Plans to.test sematic sterage and transplacental transmissien ef third
stage lungwerm larvae were helped when all feur test ewes bere lambs.
Ewe I bere a ewe lamb between June 19 and 24. Ewe II had a ewe lamb between June 2 and 8. Ewe III had a ram lamb between May 20 and 31 in Ewe
IV had a ewe lamb between May 20 and 31.
The study area enclesed 363 acres ef pasture at Little Hills. It is hilly,
with cliffs and steep hillsides. A pinen-juniper asseciatien is predeminant
with sagebrush flats en the canyen bettems. Annual precipitatien averages
12.5 inches including up to. 18 inches ef censtant snew cever en the ridgeteps and in the canyen bettems during the winter. Temperatures range from
30 degress Fahrenheit belew zero. in the winter to.100 degrees during the
summer. The sheep are en a high plane ef nutritien. They are given
pelleted feed as a supplement during the winter. The state ef these animals
is addressed by the fact that a two.year eld ewe bere a lamb in 1973.
The sheep are Recky Meuntain bighern sheep (Ovis canadensis). While the ram
came frem a Birmingham, Alabama zee the ether animals were transplanted frem
Glenweed Canyen, Celerade in the winter ef 1971. They are tame because they
have been werked with extensively. Each sheep can be distinguished frem the
ethers by hem characteristics, size and persenality.
.
Intermediate hest land snails used were Vallenia cyclepherella and pulchella.
The latter species is the ene used successfully by Mensen (1971) to. complete
the pretestrengylin life cycle in meuflen-bighern hybirds. Land snails fer
initial laberatery cultures were cellected at Glenn Eyrie and at a private
residence in Celerade Springs. Culture, prepagatien and infectien ef intermediate hests was discussed in the 1971 repert.
RESULTS
At this writing (7-6-73) the study is incemplete. One lamb, frem Ewe IV,
passed first stage lungwerm larvae at the rate ef 6 larvae/gram ef dried
feces in a sample cellected 7-3-73. This is censistent with earlier werk
and indicates that lungwerms are just new maturing in these sheep. Signs
that weuld indicate develepment ef respiratery pathelegy have been few to.
date. AlIef the lambs have reugh ceats with the eldest enes leeking the
reughest. One lamb has been ebserved ceughing fer a minute er two. en two.
eccasiens.
This study is centinuing. In the weeks to. ceme we expect to. track the rise
in larvae eutput as lungwerms mature, shewing transplacental transmissien.
Also., frequent ebservatiens are being made to.watch fer signs of respiratory
disease.
.

SEGMENT OBJECTIVE NUMBER 5
Study the epid~ielegy

ef Pretestrengt1us

in Celerade bighorn sheep.

PROCEDURE
Fecal samples from various herds throughout the state will be collected at
random on a menthly basis and the lungworm larval output between herds compared;

�-158-

hopefully this procedure will be an indication of herd health.
RESULTS
During this study the sheep were continually monitored for level of first
stage larvae putput. Using this method, larvae counted in dried, crushed
fecal samples were related in larvae per gram of feces. A legitimate
question is what do larvae output figures tell us? First, larvae output
figures can tell us if a herd is infected. This is not particularly valuable information since all herds are infected. Second, larvae output
figures detect seasonal variation. Forrester and Senger (1964) noted that
high concentrations of larvae in monthly fecal samples collected on Wildhorse
Island, Montana during 1959-60 and 1961-62 occurred during April and May.
They observed that most moisture fell in the area during April, May and
June. Pillmore (1958c) collected fecal pellets at Glenn Eyrie in Colorado
Springs. He observed higher larvae outputs during and after January than
he had seen in November and December. During the past year personnel of
the Colorado State Division of Wildlife have collected 260 pellet groups
on Pikes Peak, Trickle Mountain, Cache la Poudre and Buffalo Peaks ranges.
A cursory glance at these larvae output figures (Table 6) shows differences
between seasons on these ranges. Lower larvae outputs are in August and
September. Higher counts are during winter and spring. There are several
interpretations of seasonal fluctuation. Larvae outputs may reflect depressed immune response to first stage larvae, increased infection with
adult lungworms from range snails, greater concentration of fecal material
during winter months, or activation and infection with somatically stored
larvae. Most of the increase probably reflects a stress-related decline in
the immune response. There is a third piece of information that larvae output figures may furnish. There is some indication they can be used to compare
the state of infestation between herds. Forrester and Senger (1964) felt that
there was an association between the level of infection with lungworms in the
sheep and the larval output when compared on a herd basis. That is to say
that differences in levels of infection in herds are reflected by differences
in larvae output. This appears to be true. Such a comparison may be made
between January and March 1973 figures on Pikes Peak and January and March
figures at East Trickle Mountain (Table 7). The value of such information is
that it may describe the general health of the herd. The herd on Pikes Peak
is unhealthy and declining at present while the herd at Trickle Mountain
seems to be healthy and increasing. The figures seem to reflect this. The
comparisons are interesting but more work must be done to see if larvae output can be used to compare the health state of herds.
Finally there are some things that larval output cannot do. A single pellet
group cannot be counted and interpreted. It may be possible to collect a
series of pellet groups and make a statement as discussed above. Larval
output figures are not repeatable from day to day except at very low levels
of infection. There is so much randomness of larval putput that large numbers
of samples are needed from a herd or repeated samples from an individual
sheep if a trend is to be established.

�-159-

Table

6.

Lungworm larvae output in four Colorado bighorn sheep herds.
Aug. 72

Sept. 72

Nov. 72

Dec. 72

Jan. 73

Mar. 73

Pike's Peak

63

11

530

116

364

253

Buffalo Peaks

19

42

East

8

22

Middle

67

West

56

Area

May 73

561

129

Trickle Mountain
121

117

89

147

9
46

Cache la Poudre
Figures represent first stage lungworm larvae per gram of air-dried feces and are
averages of mUltiple samples.

344
212

�-160-

Table

7 • Lungworm larvae output in four Colorado bighorn sheep herds*.

Aug. 1972

Ave.

Sept. 1972

P I K E ' S

PEA

Nov. 1972

Dec. 1972

Jan. 1973

March 1973

K

1

1

717

38

610

None

None

27

107

7

1471

144

None

1

2489

338

699

292

None

26

77

46

423

133

None

4

487

23

184

767

1

4

149

63

129

251

None

13

173

294

158

142

92

56

44

360

719

183

2

55

153

121

None

160

102

1

24

12

192

449

10

104

138

19

22

231

1

19

None

22

3

1

None

2

None

63

11

364

253

530

116

-----------------------------------------------------------------------------------*Numbers in larvae per gram of air-dried feces.

�-161-

Table 7 • Lungworm larvae output in four Colorado bighorn
sheepherds* (continued).
PEA

BUFFALO

K S

Sept. 1972

Jan. 1973

May 1973

1

None

61

355

92

9

27

298

16

48

77

430

None

1

395

81

8

.17

29

338

7

1

423

1832

None

28

253

249

20

229

69

1857

None

109

54

518

5

1

25

577

37

11

30

428

58

47

100

263

Aug. 1972

72

15
13
Ave.

19

42

129

561

--------------------------------------------------*Numbers in larvae per gram of air-dried feces.

�-162-

Table 7 .• .Lungwo.nnlarvae. Qut:put:in JQuz:Go19radQ

~:lgllQ:rn
~heep herds*
..

. . . . .

TRICKLE
Aug. 1972

Sept. 1972

MOUNTAIN

(continued) •

.

E A S T

Nov. 1972

Jan. 1973

March 1973

May 1973

12

9

21

54

68

644

5

4

183

85

16

30

None

18

63

92

28

49

10

1

329

196

58

137

None

58

211

221

145

32

10

3

12

208

10

67

15

13

61

81

369

121

19

3

69

54

100

181

11

1

36

10

93

193

5

6

38

3

20

1

133

.243

89

18

19

24

None
Ave.

8

22

121

117

89

147

--------------------------------------------~---------------------------------------*Numbers in larvae per gram of air-dried feces.

�-163-

Table 7. Lungworm larvae output in four Colorado bighorn sheep herds*
(cont Imied) •
TRICKLE MOUNTAIN MIDDLE

TRICKLE MOUNTAIN WEST

Aug. 1972

Nov. 1972

Aug. 1972

Sept. 1972

May 1973

178

7

159

16

323

27

None

25

43

1672

163

None

20

68

78

29

10

10

36

299

47

7

159

36

601

38

None

20

2

222

138

10

22

51

44

156

None

65

111

129

24

44

20

136

39

27

36

46

344

14
34
14
22
Ave. 67

9

56

CACHE LA POUDRE
May 1973
130

99

40

125

88

1

123

360

3

337

307

389

478

482

Ave. 212
*

Numbers in larvae per gram of air-dried feces.

�-166-

third-stage larvae in the liver and lungs at birth, for a projected total
of 253, and the other had 147 in the liver and lungs, fora projected total
of 221 (see Segment Objective Number 8). Obviously the Cambendazole did not
kill the infective larvae stored in her tissues.
A young ewe lamb captured on Pikes Peak in late May, 1973 was treated with
Cambendazole to determine if this compound would kill her third stage larvae.
When she died, a total of 57 third and fourth stage lungworm were found in
he~ liver and lungs. Many of the fourth stage were digested; consequently,
there is no way of arriving at the exact level of infection (see Segment
Objective Number 8). While Cambendazole may have reduced the burden of infection, the drug certainly did not eliminate the parasites.
Of the 12 ewes brought into captivity, four gave birth to five lambs, but
the twin lambs from one ewe died, leaving three alive. Fortunately, one
lamb is from a control ewe, one from a Tramisol-treated ewe, and one from a
Cambendazole-treated ewe. The lambs are presently at the age when they
should begin showing signs of infection in the wilds; thus, there is no way
of evaluating the efficacy of treatment for another two to four weeks.
The sheep captured, treated, banded, and released on Pikes Peak (see Segment
Objective Number 18), are currently being monitored, but evaluation must be
postponed until after the lamb die-off on Pikes Peak. This will be in late
September or early October of 1973.

SEGMENT OBJECTIVE NUMBER 8
Examination of bighorn sheep for possible transplacental or transmammary
transmission of Protostrongylus.

PROCEDURE
Recent evidence strongly suggests that infection may be acquired in ways other
than by oral ingestion of infected snails. One effective means of evaluating
this possibility is to collect and examine ewes in late pregnancy. At least
two pregnant ewes should be examined. The placenta and mammary glands from
the ewes and the liver, lungs and lymph nodes from the foetus should be
macerated and run through the Baermann funnel to collect any larvae of Protostrongylus which might be present. A portion of each organ or tissue would
be preserved for histopathology. The foetal kidneys would be needed by the
virologist for tissue culture procedures used in the isolation of viral agents.

RESULTS
A ewe in late pregnancy was collected from Pikes Peak on May 16, 1972 and her
foetal tissues were examined for evidence of transplacental transmission of
Protostrongylus. The results of that examination revealed that transplacental
transmission does indeed occur. These findings are reported in the 1971
annual report. Since one case of transplacental transmission merely suggests

�-167-

that this is the usual means by which lambs receive their initial infection, additional evidence was necessary. Therefore, all pregnant ewes
or lambs dying accidentally were carefully examined for additional instances
of this type of transmission.
During the first week of April, 1973 a pregnant ewe died accidentally during
routine trapping operations on Pikes Peak. Measurement of her foetus indicated that she was within six weeks of lambing. Tissues from the ewe and
the foetus were exhaustively examined in an effort to obtain additional
evidence of transplacental transmission, and find the site of somatic storage
in the adult animal. Tissues from the ewe included lymph nodes, liver, uterus,
cotyledons, muscles from the diaphragm, and all four limbs. Tissues from the
foetus included the liver, lungs and lymph nodes. About two-thirds was used
to recover intact larvae, the remaining one-third was preserved for histopathology study. These tissues were finely minced and placed in the Baermann
funnel for 24 hours. The material collected was preserved in 10 percent
formalin for subsequent study.
A total of 43 third-stage larvae of Protostrongylus sp. was recovered from the
foetal liver; none was recovered from any of the other tissues listed above.
In the second week of April, 1973 a captive ewe died of pasteurellosis.
Measurements of her foetus indicated that she was 2-2~ months from lambing.
A similar examination of this ewe and the foetus was performed using the
Baermann apparatus. One third-stage larva was recovered from the cotyledons;
none were found in any of the other tissues.
On April 29, 1973 a captive ewe aborted a full-term ewe lamb. This ewe had
been ill for several months and the dominant animals in the herd fought her
continuously. Examination of this lamb revealed that third stage larvae of
Protostrongylus were present in the fetal liver and lungs. The liver weighed
116 grams and 45 grams were used to recover third stage larvae. The remainder
was preserved for histopathology. A total of 189 larvae were found in the
liver. Theoretically the liver possessed 472 larvae. The left lung was perfused with 10 percent formalin for histopathology study, the right was
baermannized for the recovery of third-stage larvae. Seven infective larvae
were found in the right diaphragmatic lobe, 17 in the right apical lobe and
22 in the right cardiac lobe making a total of 46 larvae. Theoretically, the
lamb had 92 larvae in the lungs at the time of birth.
On May 21, 1973, a lamb born to a captive ewe died a day after birth and was
examined for evidence of transplacental transmission of Protostronglyus. The
tissues from this lamb were examined by the Baermann funnel and the pepsinhydrochloric acid digestion technique. Seventy-five percent of the lungs were
digested, the remainder was preserved for histopathology examination. A total
of 147 third stage larvae were recovered from the lungs. Theoretically, the
lungs possessed a "total of 196 larvae. Seventy-five percent of the liver was
digested, the remaining 25 percent was preserved for histopathology examination.
A total of 43 third stage larvae was recovered. Theoretically the liver
possessed 57 larvae." The total for this lamb was 189, with a proj ected total
of 253.
The Baermann funnel is extremely effective for separating larval stages from

�-168-

feces 9 cultures, etc., but efficiency decreases when tissues such as lung,
liver or cotyledons are examined.· These tissues readily break down and the
finer particles block the screens, inhibiting passage of the larvae. Considerable difficulty was encountered with tissues from the first sheep,
which may account for the low number recovered, but with the last three
animals the technique was improved by usirig smaller amounts of tissue in
more funnels. The pepsin-hydrochloric acid digestion technique, which is
so effective in removing nematodes from tissues, will digest Protostrorigylus
adults; therefore, we thought that larvae would also digest. Thus, the
material obtained from the first three sheep was not subjected .to the digestion
technique, but after the material from the fourth sheep had been baermannized
it was subjected to digestion. A total of 58 larvae were found by the Baermann
technique, 131 by the digestion technique; consequently, we must assume that
baermannization is a very poor procedure. We must also assume that the number
of larvae recovered from the first three sheep represents a small fraction of
the total present in fetal tissues.
In mid-May, 1973, a lamb (73BHL-lO) born of captive ewe BHE-4 died of an
enteric clostridial infection 2~ to 3 days after birth. As with the preceding
lamb, 75 percent of the lung and liver tissue was examined for larvae, the
remaining 25 percent preserved for histopathology study.
Examination by the Baermann technique revealed that 30 third stage larvae
were present in the lungs and 19 in the liver. By the digestion technique,
using the same tissue, 103 were recovered from the lungs and 14 from the liver.
This is an actual recovery of 166 third stage larvae. Theoretically, the
animal possessed a total of 44 in the liver and 177 in the lungs, or a theoretical total of 221 third stage larvae.
.
In late May, 1973 a newborn female lamb was captured on Pikes Peak and maintained in captivity. Seven days later, on the lamb's eighth day of life,
she died of a clostridial infection of the gastrointestinal tract. Since this
animal was old enough to be infected withProtostrorigylus which would be in
the process of developing from third to fourth stage, and possibly even to
immature adults (fifth stage), she was examined first of all by the Baermann
technique and her tissues were then digested. As with the preceding animals,
75 percent of the liver and 75 percent of her lungs were examined. The remainder preserved for histopathology.
No larvae were found in the liver tissue examined by the Baermann technique9
but five were recovered after digestion. A total of 39 larvae, essentially
equal numbers of late third stage and fourth stages, were recovered from lung
tissue examined by the Baermann technique. The great majority (36) were
recovered from the diaphragmatic lobes. This same lung tissue was then
digested and an additional 13 larvae were recovered from the diaphragmatic
lobes, but none were found in the other lobes. These 13 were all developing
third stage larvae. However, large numbers of partially to almost totally
digested larvae were evident in the digest. There was no satisfactory procedure for counting these pieces, but an estimate of 100 worms was considered
conservative.
A total of 57 third and fourth stages were recovered from this lamb. Five of
these came from the liver, the remaining 52 from the lungs. Undoubtedly, the
great majority had already migrated to the lungl;land began development. The
52 found in the lungs theoretically represents a total of 70 in the lungs;

�-169-

however, since development to the fourth stage had occurred, and many
digested pieces were found, it must be concluded that this number represents
only a fraction of the total. Moreover, the discovery of digested larvae
indicates that the fourth stage, like the adult, is susceptible to digestion.
Thus the accuracy is considerably reduced.
Thus far, evidence of transplacental transmission has been obtained from all
lambs or foetuses examined. However, present techniques indicate that results
obtained with the first three animals was not accurate (see paragraph discussing the Baermann funnel).
When the ewe was collected on Pikes Peak in late May of 1972, some additional
circumstantial evidence in support of transplacental transmission of lungworm
was fortuf.tously obtained. A young lamb, not more than one or two days old,
was captured and reared in captivity. At an estimated age of 35 days, he
began passing first stage lungworm larvae in the feces. Over the next three
weeks his output increased, eventually reaching the extremely high level of
lO,OOO/gram. Concomitant with this heavy infection, the lamb developed a
chronic cough, shallow respiration and tired easily. He was also light in
weight, light in color and had a rough hair coat, most of which was ascribed
to his diet at that time. About the first of August, this lamb had learned to
eat grass and had a free-choice of grass, hay and grain in addition to the
cow's milk he received three times daily. At this time he began to grow and
put on considerable weight. His high larvel output remained unchanged until
about August 15, 1972, when, in the short span of a week, the output dropped
from lO,OOO/gram to lO/gram of feces. Presently, this lamb is an extremely
healthy animal and his larval output is a consistent lO/gram of feces.
Several conclusions can be drawn from studies on transplacental transmission of·
Protostrongylus:
(1) the data, while incomplete, indicates that this is the
usual means by which lambs receive their initial infection with P. stilesi in
herds experiencing high lamb mortality; (2) transmission probably occurs in
late pregnancy; (3) the larvae are stored in the foetal liver until birth, or
shortly before, and then migrate to the lungs; (4) the ewe probably accumulates
these larvae throughout the year and stores them (somatic storage) in some
site presently unknown; and (5) the lamb is born with an overwhelming infection
which probably predisposes it to secondary invasion by bacteria.
Now that we are aware of transplacental transmission of lungworm in bighorn
sheep, what can be said about the cycle from this point on is theoretical, and
will require several years of difficult work to prove, but proof is now within
our grasp. Theoretically, the pregnant ewe obtains third stage (infective)
larvae with snails she aceidentally ingests while grazing. When these larvae
are released in the gastrointestinal tract, they apparently enter her circulatory system, cross the placenta, and accumulate in the foetal liver until
birth. When the lamb is born, infective larvae are already present, ready
to enter the lungs and continue development. In the lung they grow, develop,
and molt through two additional stages to the adult parasite. Adult parasites
reach sexual maturity, mate and begin producing eggs. First stage larvae
appear in the feces 4-6 weeks after birth.
There are a number of reasons why this mode of transmission is extremely important, and all of them are important to the management of the bighorn sheep
herd; consequently, the problem·must be thoroughly studied and all of its
implications evaluated.

�-170-

When the ewe eats snails, and acquires infective larvae, some of these
larvae may be destined to cross the placenta and enter the foetus, and
some may be destined to enter her lungs and mature. However, since she
already has lungworm, she is probably immune to a higher level 6f infection, especially if she is healthy; thus, the majority of the larvae
are destined to cross the placenta and enter the foetus. Presently,
circumstantial evidence indicates that the larvae acquired by a ewe in
the course of grazing throughout the year go into her body tissues and
become dormant. Here they could accumulate, often in great numbers, to
be released during pregnancy year after year, even with no further intake
of larvae. One important management problem which comes immediately to
mind is that lungworm-free herds cannot be established by planting lambs
raised in a snail-free environment.
Another significant implication of transplacental transmission of lungworm is the potential problems which could (and probably do) develop on
poor and overgrazed sheep ranges. All parasites require a number of
factors to insure perpetuation of their own kind; crowding and malnutrition
are possibly the most important. During the past 50-70 years the bighorn
sheep ranges have become smaller and smaller, forcing animals into crowded
situations. When the sheep are crowded together, more snails per unit area
are infected (especially on lambing grounds), furnishing a better chance
for ewes to accumulate greater numbers of infective larvae for their offspring. Crowding promotes overgrazing, and this too means more snails ingested because grasses are more closely cropped. If the bighorn sheep is
on poor, overgrazed range, it is undoubtedly in a marginal or suboptimal
plane of nutrition. Most likely the lambs born of these ewes are in the
same condition. If the ewe gives birth to a weak lamb, and does not furnish adequate milk, the lamb has several strikes against it at birth.
When these factors are combined and the parasites added as a final ingredient,
the result is that most lambs are born predestined to die.
If this is the correct interpretation of the cause of lamb mortality, at
least two avenues of control must be explored: (1) kill the third stage
larvae in the ewe before she can transmit the parasite to the lamb; and/or
(2) kill the third stage larvae in the bedding and lambing grounds, thus
insuring that ewes will not acquire large numbers to pass to the lambs.
Presently (see above under Segment Objective Number 6) treatment of the ewe
is being explored with Tramisol and Cambendazole. This summer and next year,
the feasibility of treating the lambing and bedding grounds will be explored.
SEGMENT OBJECTIVE NUMBER 9
Identify the virus recently isolated from captive bighorn sheep in Wyoming.
This virus was identified as PI-3 and reported in 1971.
SEGMENT OBJECTIVE NUMBER 10
Experimentally inoculate domestic lambs with the virus mentioned above to
determine the effect of the agent on susceptible animals. Since the virus
was PI-3, and the effect on domestic sheep is well-documented, there was no
need to pursue this objective.

�-171-

SEGMENT OBJECTIVE NUMBER 11
Develop and evaluate a vaccine against the virus named above if warranted.
Present results indicate that PI-3 is one of the agents responsible for
fatal pneumonia in lambs, but not the predisposing agent; therefore,
preparation of a vaccine seems unwarranted, moreover, no long-term benefits
can be obtained by inoculating sheep other than captive animals because
PI-3 is essentially an ubiquitous virus.
SEGMENT OBJECTIVE NUMBER 12
Initiate tissue cultures from a healthy lamb. This is a routine step
necessary to isolate viruses and this objective was met in 1971 with the
collection of a pregnant ewe on Pikes Peak. In 1972, other tissue culture
systems were employed.
SEGMENT OBJECTIVE NUMBER 13
Continuation of serology, using the serum samples on hand and procurement
of additional serum samples. This objective was met, and the results are
reported in Segment Objectives Number 1 and 18.
SEGMENT OBJECTIVE NUMBER 14
Obtain fresh tissue and nasal washings from bighorn sheep and attempt viral
isolation. This objective was met and the results are reported in Segment
Objective Number 1.
SEGMENT OBJECTIVE NUMBER 15
Attempt isolation of chlamydial agents. This objective was not met and
unfortunately no attempt was made to do so.
SEGMENT OBJECTIVE NUMBER 16
Microscopic examination of tissue culture cells for evidence of viral infection (cytology). This is a routine objective and is a procedure required
of the virologist to interpret presence or absence of infection.
SEGMENT OBJECTIVE NUMBER 17
Report on Segment Objectives.
SEGMENT OBJECTIVE NUMBER 18
This objective was not written into the plans by the research team but came
about as the result of discoveries made after the plans had developed. The

�-172-

plan was conceived by the Division of Wildlife as a multipurpose, multidisciplinary objective. That objective was to trap, band and release
sheep on Pikes Peak to: (1) determine behaviour and migratory habits;
(2) determine the age structure, (3) determine the presence or absence
of pathogens in the herd; and (4) acquire captive animals for study. A
summary of the results are presented in Tables 8, 9 and 10.
Six of the ewes captured early in the experiment died as a result of age
and stress. All of these animals were old and debilitated. The postmortem
examination revealed that they had a chronic pneumonia of long-standing
duration, accompanied by severe pleural adhesions; thus they apparently
died of a combination of stress, age and pneumonia. Moreover, the first
four animals were to be transported to Little Hills, Colorado and a severe
storm delayed progress for 24 hours, which was an added stress. These
died shortly after their arrival.
Nasal swabs were taken and examined for the presence of Pasteurella in the
upper respiratory tract in as many of the sheep as was possible under the
circumstances. Pasteurella was isolated from only a few of the sheep, but
postmortem examination of those dying revealed that the organism was present. For example, Pasteurella was obtained from the upper respiratory
tract of only one of the first 5 sheep that died, but was obtained from the
lower respiratory tract (lungs) of all of them. Winter conditions on Pikes
Peak are extremely rigorous and our present interpretation is that Pasteurella
cannot survive the cold (as low as -200 F) air in the nasal passage.
As expected, most of the sheep had a titer to PI-3, indicating exposure to
this virus at sometime in their life. The only titers run at present are
the 1:2, which indicates only exposure and does not reveal the strength of
the immunity; the press of other work has delayed these studies, necessitating
they be reported in the 1973 annual report.

�Table 8.

Bighorn ewes - 0 through 26; April 6, 1973.

Treatment

Pasteurella
Isolated From
Nasal Swabs

Titer
PI-3
1:2

Blood
Taken

Place of Capture

Captive or
Released

Tag Color and
Number

7-8 yr.

Tram. *

---

+

Yes

Old Trap Site

Captivet

Red/White 114

73BBE- 5

6-7 yr.

Tram. *

---

+

Yes

Old Trap Site

Captivet

Red/White

115

1-9

73BBE- 6

7-8 yr.

Tram. *

---

+

Yes

Old Trap Site

Captivet

Red/White

116

1-9

13BBE- 7

7-8 yr.

Tram .•*

---

+

Yes

Old Trap Site

Captivet

Red/White

117

1-11

73BBE;" 8

6-8 yr.

Tram. *

P. hemolytica

+

Yes

Old Trap Site

Captivet

Red/White 118

1-11

13BHE- 0

6-7 yr.

Tram. *

+

Yes

Old Trap Site

Captivet

Red/White 110

+

Yes

Sheep Mt.

Captivett

Red/White

Date
cap.

Ewe Number

Age

1-9

73BHE- 4

1...•
9

I

2-13

13BBE- 9

6 yr.

Camb.**

-----

2-13

73BBE-IO

8ye

:Control

Past. ap.

+

Yes

Sheep Mt.

Cap tivet t

Red/White 114

2..;.13 73BHE-l1

6yr

Tram. *

---

+

Yes

Sheep Mt.

Cap tivet t

Red/White 116

....•
....•
W
I

113

2-13

7111HE-12

8-'9 yr.

Tram. *

P. hemolytica

+

Yes

Sheep Mt.

Captivett

Red/White

2-16

73BHE-13

6-.7 yr.

Camb.**

+

Yes

Sheep Mt.

Captivett

Red/White 118

2-16

73BHE-14

4 yr.

Control

+

Yes

Sheep Mt.

Cap tivet t

Red/White 119

2-21

13BBE-lS

1-9 yr.

Camb.**

-------

+

Yes

Sheep Mt.

Captivett

Red/White 1110

2-21

73BUE-16

5-:6 yr.

Tram. *

---

+

Yes

Sheep Mt.

Captivett

Red/White 1111

2-21

73JmE-17

5~yr.

Control

---

+

Yes

Sheep Mt.

Captivett

Red/White 1112

3-1

73BHE-18

6-7 yr.

Camb.**

---

+

Yes

Sheep Mt.

Released

Orange 111

-----------------------------------------------------------------------------------

115

�Table 8 - Bighorn ewes - 0 through 26; April 6, 1973 (continued).

Treatment

Pasteurella
Isolated From
Nasal Swabs

Titer
PI-3
1:2

Blood
Taken

Place of Capture

Captive or
Released

Tag Color and
Number

7-8 yr.

Camb.**

---

+

Yes

Sheep Mt.

Released

Orange 112

73BHE-20

5-6 yr.

Camb.**

+

Yes

Sheep Mt.

Released

Orange 114

3-1

73BHE-21

5-6 yr.

Tram. *

+

Yes

Sheep Mt.

Released

Red/White 113

3-1

73BHE-22

7-8 yr.

Tram.**

-------

+

Yes

Sheep Mt.

Released

Red/White 1f4

3-1

73BHE-23

4 yr

Tram. *

---

+

Yes

Sheep Mt.

Captivet

Red/Whi te 111

3-1

73BHE-24

4 yr.

Camb.**

+

Yes

Sheep Mt.

Captivett

Red/White

3-1

73BHE-25

3 yr.

Control

-----

+

Yes

Sheep Mt.

Captivett

Red/White 1113

4-4

73BHE-26

4 yr.

Dead in net (4-mile location)

---

-------

--------

*
**

Tramisol
Cambendazole
Dead
Alive

Date
Cap.

Ewe Number

Age

3-1

73BHE-19

3-1

+
tt

In

.....I
~
"
I

-------

�Table 9.

Bighorn rams - 1 through 19.

Date
Ca2·

Ram Number

Age

Treatment

1-9

73BHR- 1

5 yr.

1-9

73BHR- 2

5 yr.

1-9

73BHR- 3

5 yr.

1-9

73BHR- 4

7 yr.

Pasteurella
Isolated From
Nasal Swabs

Titer
PI-3
1:2

Blood
Taken

Place of Ca2ture

Captive or
Released

Tag Color and
Number

No Sample

+

Yes

Old Trap Site

Released

Tag 111

----

-

Yes

Old Trap Site

Released

Tag 112

----

+

Yes

Old Trap Site

Captive*

Tag 1/3

----

+

Yes

Old Trap Site

Released

------

----

+

Yes

Old Trap Site

Released

------

1-9

73BHR- 5

9 yr.

-----------

1-9

73BHR- 6

11 yr.

---

----

+

Yes

Old Trap Site

Released

------

1-9

73BHR- 7

8 yr.

---

No Sample

+

Yes

Old Trap Site

Released

------

1-9

73BHR- 8

9 yr.

---

----

+

Yes

Old Trap Site

Released

------

1-11

73BHR- 9

7 yr.

----

+

Yes

Old Trap Site

Released

------

2-1

73BHR-10

5 yr.

-----

----

No
Blood

Lost

Old Trap Site

Released

Red Tape

2-1

73BHR-ll

7 yr

----

+

Yes

Old Trap Site

Released

Red Tape

2-1

73BHR-12

8 yr.

-----

+

No
Blood

---

Old Trap Site

Released

Red Tape

2-1

73BHR-13

9 yr.

---

----

+

Yes

Old Trap Site

Released

Red Tape

2-1

73BHR-14

4 yr.

---

----

+

Yes

Old Trap Site

Net Death

------

---

I
f-I

Red Tape
Sheep Mt.
Released
Yes
---+
-------------------------------------------------------------------------------------------------------------------2-16

73BHR-15

5 yr.

"

\JI
I

�Table 9 - Bighorn ~ams - 1 through 19

(continued).

Date
Cap.

Ram Number

Age

Pasteurella
Isolated From
Nasal Swabs

2-13

73BHR-16

5 yr.

2-13

73BHR-17

3 yr.

2-16

73BHR-18

5 yr.

Cl. perf.
D. Vacc.

3-1

73BHR-19

6 yr.

70 cc.
Cambin.

Treatment

No Sample

Titer
PI-3
1:2

Blood
Taken

Place of Capture

Captive or
Released

Tag Color and
Number

+

Yes

Sheep Mt.

Released

Red Tape

Yes

Sheep Mt.

Released

Red Tape

+

Yes

Sheep Mt.

Captive*

White/Red /17

+

Yes

Sheep Mt.

Released

Red Tape
I
•....

-...J

0\

*

Alive

I

�Table 10.

Bighorn lamb Number 1.

Date
Cap.

Lamb Number

Age

Treatment

Pasteurella
Isolated From
Nasal Swabs

1-9

73BHL-l

8 mo.

10 cc P-S

P. multiocidia

Titer
PI-3

1:4

Blood
Taken

Place of Capture

Yes

Old Trap Site

Captive or
Released

Tag Color and
Number

Captive
Dead

None

I

t-'

"
"
I

�-178-

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Davis, W. B. 1938. Summer activity of mountain sheep on Mt. Washburn,
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�-179-

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1973.

J. Wi1d1. Dis. Assoc.

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Research note; J. Wi1d1.

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Honess, R. F.
Wyoming.

1942. Lungworms of domestic sheep and bighorn sheep in
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1955.
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Land snails and the lungworm of bighorn sheep. Fed. Aid
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___

, and M. Frost. 1942. A Wyoming bighorn sheep study.
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Wyo. Game and

__

~, and K. B. Winter. 1955. Incidence of lungworm parasitism in bighorn sheep. Fed. Aid Projects. Quarterly Reports. Wyo. Game and Fish
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Hunter, G. N., and R. E. Pi11more. 1954. Hunting as a technique in studying
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93-98.

�-180-

Jones, G. W., and C. E. White. 1950. Rocky Mountain bighorn sheep surveys and investigations. Fed. Aid Projects. Quarterly Reports.
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________
, and C. A. Moser. 1953. Census and distribution studies on herds
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Kassai, T. 1958. Larvae of protostrongylins
8(3):224-236.

in snails.

Keyes, M. C. 1965. Pathology of the Northern fur seal.
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Acta Vet.

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Parasites from

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Pneumonia in Rocky Mountain bighorn sheep.

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J. Mammal.

A preliminary study of the bighorn sheep of Yellowstone
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__~

, and G. W. Jones. 1954. Lambing study and predator loss to lambs.
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--------Projects.
Quarterly Reports.

Study of mortality factors. Fed. Aid
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�-181-

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1954b.
19-21.

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�-182-

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�-183-

ADDENDUM

�-184-

31 Bighorn Rams Captured on Pike's Peak
in the Winter of 1973

Date Capt.

Ram Number

Age

Tag Color and
Number

Place of Capture

1-9

73BHR- 1

5 yr.

Red/White III

Old Trap Site

1-9

73BHR- 2

5 yr.

Red/White 112

Old Trap Site

1-9

73BHR- 3

4 yr.

Red/White 113*

Old Trap Site

1-9

73BHR- 4

4 yr.

------

Old Trap Site

1-9

73BHR- 5

9 yr.

------

Old Trap Site

1-9

73BHR- 6

11 yr.

------

Old Trap Site

1-9

73BHR- 7

8 yr.

------

Old Trap Site

1-9

73BHR- 8

9 yr.

------

Old Trap Site

1-9

4 yr.

-----

Old Trap Site

1-9

6 yr.

------

Old Trap Site

1-11

73BHR- 9

7 yr.

------

Old Trap Site

2-1

73BHR-lO

5 yr.

Red Tape

Old Trap Site

2-1

73BHR-11

7 yr.

Red Tape

Old Trap Site

2-1

73BHR-12

8 yr.

Red Tape

Old Trap Site

2-1

73BHR-13

9 yr.

Red Tape

Old Trap Site

2-1

73BHR-14

4 yr.

------

Old Trap Site

2-16

73BHR-15

5 yr.

Red Tape

Sheep Mt.

2-13

73BHR-16

5 yr.

Red Tape

Sheep Mt.

,2.,..13

73BHR-17

.5 yr.

Red Tape

Sheep Mt.

?-16

73BHR-18

3 yr.

Red/White**

Sheep Mt.

3.•
1

73BHR-19

6 yr.

Red Tape

SheeP Mt.

3•••
23

......

4 yr.

Red Tape

Cathedral Park

Treatment

Net death

Net death

---~~

Camb ,

~.~.~~__.__~_w~_.~~_-.
_____~~_____~~__~~___~~~~__~~____---~~---~----_~
__________
~~__

�-185-

31 Bighorn Rams Captured on Pike's Peak in the Winter of 1973 - Continued.

Age

Tag Color and
Number

Place of Capture

3-23

5 yr.

Red Tag

Cathedral Park

4-1

4 yr.

Red/White 115

Cathedral Park

4-1

5 yr.

Red/White 116

Cathedral Park

4-1

4 yr.

Red Tape

Cathedral Park

4-4

4 yr.

Blue 112

Four-Mile

4-4

4 yr.

Blue 113

Four-Mile

4-4

7 yr.

Blue 114

Four-Mile

4-4

4 yr.

Blue 118

Four-Mile

4 yr.

Orange 1111

Gravel Pits

Date Capt.

5-12

*
**

Ram Number

73BHR-20

Sheep taken to Little Hills
Sheep taken to Fort Collins

Treatment

Camb.

�-lS6-

43 Bighorn Ewes Captured on Pike's Peak
in the Winter of 1973

Date
Cap.

Ewe Number

Age

Tag Color and
Number

Place of Cap.

Treatment

Captive or
Released

1-9

73BHE- 4

7-S yr.

Red/White /14

Old Trap Site

Tram.

Capt. - D*

1-9

73BHE- 5

6-7 yr.

Red/White /15

Old Trap Site

Tram.

Capt. - D*

1-9

73BHE- 6

7-S yr.

Red/White /16

Old Trap Site

Tram.

Capt. - D*

1-9

73BHE- 7

7-S yr.

Red/White 117

Old Trap Site

Tram.

Capt. - D*

1-11

73BHE- S

6-S yr.

Red/White /IS

Old Trap Site

Tram.

Capt. - D***

1-11

73BHE- 0

6-7 yr.

Red/White /10

Old Trap Site

Tram.

Capt. - D***

2-1.3

73BHE- 9

6 yr.

Red/White 113

Sheep Mt.

Camb.

Capt.**

2-13

73BHE-10

S yr.

Red/White /14

Sheep Mt.

Control

Capt.**

2-13

73BHE-11

6 yr.

Red/White fl6

Sheep Mt.

Tram.

Capt.**

2-13

73BHE-12

S-9 yr.

Red/White 115

Sheep Mt.

Tram.

Capt.**

2-16

73BHE-13

6-7 yr.

Red/White lIS

Sheep Mt.

Camb.

Capt.**

2-16

73BHE-14

4 yr.

Red/White 119

Sheep Mt.

2-21

73BHE-15

7-9 yr.

Red/White 1110

Sheep Mt.

Camb.

Capt.**

2-21

73BHE-16

5-6 yr.

Red/White 1111

Sheep Mt.

Tram.

Capt.**

2-21

73BHE-17

5-6 yr.

Red/White 1112

Sheep Mt.

3-1

73BHE-18

6-7 yr.

Orange 111

Sheep Mt.

Camb.

Released

3-1

73BHE-19

7-S yr.

Orange 112

Sheep Mt.

Camb .

Released

3-1

73BHE-20

5-6 yr.

Orange 114

Sheep Mt.

Camb.

Released

3-1

73BHE-21

5-6 yr.

Red/White 113

Sheep xe.

Tram.

Released

3-1

73BHE-22

7-8 yr.

Red/White 114

Sheep Mt.

Tram.

Released

3-1

73BHE-23

4 yr.

Red/White 111

Sheep Mt.

Tram.

Capt.**

Capt.**

Capt.**

------------------------------------------------------------------------------------

�-187-

43 Bighorn Ewes Captured on Pike's Peak in the Winter of 1973 - Continued.

Date
Cap.

Ewe Number

Age

Tag Color and
Number

Place of Cap.

3-1

73BHE-24

4 yr.

Red/White fI2

Sheep Mt.

3-1

73BHE-25

3 yr.

Red/White 113

Sheep Mt.

4-1

8 yr.

Orange 115

Cathedral Park

Camb ,

Released

4-1

6 yr.

Orange 116

Cathedral Park

Camb ,

Released

4-4

4 yr.

Orange 113

Four-Mile

Camb ,

Released

4-4

4 yr.

Blue 111

Four-Mile

Camb.

Released

4-4

8 yr.

Blue 112

Four-Mile

Camb.

Released

4-4

5 yr.

Blue 113

Four-Mile

Camb ,

Released

4-4

6 yr.

Blue 114

Four-Mile

Camb ,

Released

4-4

4 yr.

Blue 115

Four-Mile

Camb.

Released

4-4

3 yr.

White 1114

Four-Mile

Tram.

Released

4-4

4 yr.

White 1115

Four-Mile

Tram.

Released

4-4

5 yr.

White 1116

Four-Mile

Tram.

Released

4-4

5 yr.

White 1/17

Four-Mile

Tram.

Released

4-4

3 yr.

Blue 1/6

Four-Mile

Camb.

Released

Treatment
Camb .

Captive or
Released
Capt.**
Capt.**

4-4

73BHE-26

4 yr.

5-12

73BHE-28

6 yr.

Orange 117

Gravel Pits

Camb ,

Released

5-12

73BHE-27

7 yr.

Orange 118

Gravel Pits

Camb.

Released

5-12

73BHE-29

7 yr.

Orange 119

Gravel Pits

Camb.

Released

5-12

73BHE-30

7 yr.

Red/White 117

Gravel Pits

Tram.

Released

5-12

73BHE-31

6 yr.

Red/White 118

Gravel Pits

Tram.

Released

5-12

73BHE-32

10 yr.

Orange 1110

Gravel Pits

Camb ,

Released

*

**
***

Four-Mile

Sheep' taken to Little Hills
Sheep taken to Fort Collins
Died

Net death

�-188-

One Bighorn Lamb Captured on Pike's Peak
in the Winter of 1973

Date

Lamb Number

Age

1-9

73BID..-l

8 mo.

Tag

Place of Cap.
Old Trap Site

Treatment

Captive
Fort Collins

�-189-

Bighorn Ewes Captured on Pike's Peak
Treated and Released

Tag Color and
Number

Place of Capture

6-7 yr.

Orange 111

Sheep Mt.

Camb.

73BHE-19

7-8 yr.

Orange /12

Sheep Mt.

Camb.

3-1

73BHE-20

5-6 yr.

Orange 114

Sheep Mt.

Camb.

3-1

73BHE-2l

5-6 yr.

Red/White

113

Sheep Mt.

Tram.

3-1

73BHE-22

7-8 yr.

Red/White 114

Sheep Mt.

Tram.

4-1

8 yr.

Orange //5

Cathedral Park

Camb.

4-1

6 yr.

Orange 116

Cathedral Park

Camb.

*4-4

4 yr.

Orange 113

Four Mile

Camb.

4-4

4 yr.

Blue /11

Four Mile

Camb.

4-4

8 yr.

Blue /12

Four Mile

Camb.

4-4

5 yr.

Blue //3

Four Mile

Camb.

4-4

6 yr.

Blue 114

Four Mile

Camb ,

4-4

4 yr.

Blue 115

Four Mile

Camb.

4-4

3 yr.

White /114

Four Mile

Tram.

4-4

4 yr.

White 1115

Four Mile

Tram.

4-4

5 yr.

White 1116

Four Mile

Tram.

4-4

5 yr.

White 1117

Four Mile

Tram.

4-4

3 yr.

Blue 116

Four Mile

Camb .

Date Capt.

Ewe Nwnber

Age

3-1

73BHE-18

3-1

Treatment

5-12

73BHE-28

6 yr.

Orange 117

Gravel Pits

Camb.

5-12

73BHE-27

7 yr.

Orange 118

Gravel Pits

Camb.

5-12

73BHE-29

7 yr.

Orange /19

Gravel Pits

Camb.

5-12.

73BHE-30

7 yr.

Red/White 117

Gravel Pits

Tram.

5-12

73BHE-3l

6 yr.

Red/White fl8

Gravel Pits

Tram.

5-12 .

73BHE-32

10 yr.

Orange /110

Gravel Pits

Camb ,

��-191-

Determination of Some Trace Elements in Body Tissues and Blood Serum
of Rocky Mountain Bighorn Sheep by Atomic Absorption Spectrometry (AAS)

Project No. W41R-22

Robert H. Udall and Timothy Jon Raczniak
Department of Pathology,
Colorado State University,
Fort Collins, Colorado

��-193-

INTRODUCTION
Analysis of trace elements in biological substances by atomic absorption
spectrophotometry (AAS) has been a very useful technique in the field of
medicine. The development of methods of analysis have led to an increased
understanding of the roll of trace elements in metabolism as well as the
pathologic conditions resulting from metabolic disorders.
It is the purpose of this preliminary report to present a method suitable
for trace element study in the bighorn sheep. We present data collected
by the technique of AAS and compare it to the literature.
PROCEDURES
Sample Collection
Eight Rocky Mountain bighorn sheep were collected (August-October, 1972)
from various herds in Colorado. All animals were necropsied in the field.
Postmortem examination included the collection of tissues for histopathological
examination and fluids for hematological, bacteriological and virological
analysis. Tissue samples taken for trace element analysis were placed in
polyethylene bags at necropsy, frozen immediately, and kept frozen until
processed in the laboratory. Blood samples were taken by venipuncture,
refrigerated immediately. The blood was then centrifuged at the laboratory
and the serum layer removed and frozen until analyzed.

Sample Preparation
A modification of the method by Adrian (1973) was used for the digestion of
tissue. Subsamp1es of frozen tissue were taken, dried for 48-60 hr at 600
C., cooled, weighted (to the fourth place), and recorded as dry-weight.
The tissue samples were placed in Na1ge narrow mouth bottles (Na1ge-Syborn
Corporation, Rochester, New York), 3 m1 of perch10ric acid (HC104) and 6 m1
of nitric acid (HN03) were added. The acids were delivered via an automatic
buret assembly, increasing speed and accuracy with which the acids cou1d·be
added. The samples were capped and allowed to predigest for 24 hr at 250 C.
The samples were then placed in a water bath at 700 C. for 5 hr. After
further digestion in the water bath the caps were removed from the bottles
and approximately 5 m1 of distilled, de-ionized water was added to each
sample, to expel the excess volatile acid. The samples were then diluted
in 25 ml volumetric flasks and made to volume with distilled, de-ionized
water. The diluted solutions were returned to new Na1ge bottles and capped,
ready for analysis by AAS.
Serum samples were diluted (1:10) with lanthanum chloride in an automatic
diluter. Standa,rd preparation of lanthanum chloride, LaC1 ·7Hz0, was carried
3
out and diluted to make a 1 percent solution (w/v).
Standard solutions of calcium, copper, iron, potassium, magnesium, sodium,
and zinc.were prepared commercially (Aztec Instruments, West Port, Connecticut).
In all preparations spectrographic grade reagents were used; all glassware
was washed in dilute sulfuric acid and rinsed thoroughly in distilled, deionized water.

�-194-

Trace element analysis was carried out on a Perkin-Elmer Atomic Absorption
Spectrophotometer Model 303, equipped with a Perkin-Elmer Recorder Model
165. A prepared standard was aspirated every 25 samples, if there was a
deviation from the baseline, the calibration control was readjusted.
Total phosphorus was determined as a yellow molybdivandophosphoric
spectrophotometrically.

acid

A program for tissue analysis was written and statistical analysis was
carried out on a SCM Marchant Cognito/1016PR calculator.

RESULTS
The mean concentration and deviation of detected elements of various tissues
and blood samples are shown in the Appendix (Tables 5-13).
Table 1 demonstrates copper concentration in the liver, comparing normal and
copper-deficient diets. Copper concentration in the blood of ovine, bovine,
and bighorn sheep is shown in Table 3.
A composition of zinc values is shown in Table 4.
Phosphorus values were determined spectrophotometrically; but a heavy precipitate prevented accurate phosphorus determinations of spleen and liver
tissues.
Chromium, cadmium, selenium, cobalt, and molybdenum analysis were also
carried out on AAS; however, reproducible results were unattainable and
analysis for these trace elements was discontinued.

DISCUSSION
Perhaps the most important observation made during the experiment was that
copper appeared in relatively low amounts in the serum as well as tissue
samples. For humans, dogs, cats, and sheep the range of copper in blood is
given as .5 - 1.5 ug/ml. Levels of copper below .6 ug/ml in the plasma of
sheep are indicative of copper deficiency (Beck 1961).
Copper deficiency is manifested clinically by reproductive failure, due to
fetal death and reabsorption (Dutt 1960; Hall 1969). In addition, copper
deficiency affects many other body systems, showing anemia (hypochromic and
microcytic), hypopigmentation and abnormalities of hair texture, and defective
structure of vascular intima (Sandstead 1970a; 1970b).
Copper is also a component of numerous oxidative enzymes, including cytochrome,
oxidase, tyrosinase, ascorbate oxidase, and uricase.
Underwood (1971) and Everson (1968) have shown abnormalities of myelin formation in copper-deficient lambs and guinea pigs, respectively.
Little is known of the iron requirement of sheep. Lawlor (1965) found that
10 ug/ml of copper in blood of sheep is inadequate. Iron deficiency tends

�-195-

to decrease growth, decrease resistance to infection, and decrease the
ability to protect against infestation of parasites (Richard 1954).
Ott (1966a; 1966b) conducted studies on tissue changes in zinc toxicity
in sheep and cattle. He found with high levels of zinc, subnormal liver
copper concentrations and increased iron concentrations. We found with
high zinc levels a decrease in both copper and iron concentration levels
(Tables 2-3). Sandstead (1969) found with high levels of dietary zinc
(0.5 ppm) there is a high incidence of abortion.
Zinc is also essential for the growth of tissue and plays an important
role in enzyme function (Hurley 1969: Prasad 1969; Parisi 1969).
Chromium, cadmium, selenium, cobalt, and molybdenum analysis proved to
be unsatisfactory using AAS; however, using a more sensitive analytical
technique (gas chromatography) additional studies aimed at detecting
these trace elements in bighorn sheep could be conceived and carried
through.

�Table 1.

Copper concentrations

(ug/G)

* of liver tissue.

Species

Age &amp; treatment

Sheep

New born:norma1

Sheep

No. of animals

Copper
concentration
(ug/G)

Reference

diet·

27

168 (74~430)

Cunningham, 1946

New born:Cu-deficient
diet

29

13 (4-34)

"

Sheep

Mature:norma1

44

599(186-13741

"

Sheep

Mature: Cu-deficient
diet

35

27(7-106)

"

Bighorn Sheep

*dry-weight basis

Pooled

diet

7

15(3-27)

Present study

I
'"""

1.0
0\
I

�Table 2.

Serum iron (ug!m1).

No. of animals

Serum iron
(ug/m1)

Reference

Normal cows.

10

14.6(8.9-25.3)

UnderwoodJ1963

Normal bulls

10

14.5 (9.2-27 .0)

Normal ewes

12

18.2 (10.2-30.4)

Normal rams

12

15 •2 (11.4-19 .1)

7

10.7(2.0-24.5)

Species

Bovine Adults
II

Ovine Adults

Bighorn

Sheep

II

II

....
I

\0

.....•

Present

study

I

�Table 3.

Copper concentration

in the blood of several species.

Species

Age

Copper concentration
(ug/ml)

Reference

Ovine

Mature

Beck,1956

Ovine

Mature

l.Ol±.96 a
.9la

Bovine

Mature

.93a

Bighorn Sheep

Pooled

.O6±.O8

b

Cunningham, 1946

"
Present study

...•
I

\0

00
I

awhole blood
b

serum

�Table 4.

Zinc concentrations of various tissues

*.

(ug/G).

Tissue

Mana

Ratb

PigC

Bighorn Sheepd

Kidney

55
21

23

40

104

28

33
55
15

24
21
30
22

111
80
173
142

54

13

Spleen
Cardiac
Liver
Lung
Skeletal muscle

40

153

I

~

1.0
1.0
I

*dry-weight basis
aTipton and Cook, 1963
bGilbert and Talyor,1956; Mawson and Fischer,1953
c Hoekstra, 1967
d

Present study

�-200-

REFERENCES
1.

Adrian, W.J. in press The Analyst, 1973.

2.

Beck, A.B., Aust. J. Zool., i, 1, 1956.

3.

Beck, A.B., Aust. ~. Agr. Res., 12, 743, 1961.

4.

Cunningham,

5.

Dut t , B. and C.F. Mills, ~.Comp.

6.

Everson, G.J., R.E. Shrader, and 1. Wang, ~. Nutri., 94, 89,1968.

7.

Gilbert, 1.G., and D .M. Taylor, Biochem. Biophys. Acta., ~

8.

Hall, G.A. and J. McChowell,

9.

Hoekstra, W.G., E.C. Faltin, C.W. lin, R.F. Roberts, and R.R.
Grummer, ~. Animal Sci., 26, 1348, 1967.

10.

Hurley, L. S., Am. ~. Clin. Nutri., 22, 1332, 1969.

11.

Lawlor, M.J., W.R. Smith, W.M. Beeson, ~. Animal Sci. 24, 742, 1965.

12.

Mawson, C.A., M.I. Fischer, Biochemtstry, 55, 696, 1953.

13.

Ott, E.A., W.R. Smith, R.B. Harrington, and W.M. Beeson, J. Animal
Sci" ~, 414 and 419, 1966a.

14.

Ott, E.A., W.R. Smith, R.B. Harrington, M. Stod, R.E. Parker j and
W. M. Beeson, ~. Animal Sci. ,25, 432, 1966b.

15.

Parisi, A. and B. Vallee, Am. ~. Clinical Nutri., ~,

16.

Prasad, A.,D. Oberleass, P. Wolp, and J. Rorwitz, ~. Lab. Clin.
Med., 11, 486, 19Ei9.

17.

Richard, R. M., R. Shumard, A. Pope, P.R. Phillips, C.A. Rerrick,
and G. Bohsteadt, ~. Animal Sci., 13, 274 and 674, 1954.

18.

Sandstead, H.R., and Glasser, SS, R., Clin. Res. 17, 549, 1969.

19.

Sandstead, R.R., C. Lanier, G. Shepard, and P. Gillespie,
Am. J. C1in. Nutri., 23, 514, 1970a.

20.

Sandstead, R.R., R.F. Burk, G.R. Booth, W.J. Darby, Med. Clinics
N. Am., ~, 1509, 1970b.

21.

Tipton, r.u, and ~1.J. Cook, RealthPhys.,

22.

Underwood,

1. J., New Zeal. ~. Sci. Tech~ Sect. A Y..., 372,381, 1946.
Path. 70, 120, 1960.

Brit. ~. Nutri., Q, 41, 1969.

1222, 1969.

2., 103, 1963.

E.J. and E. Morgan, Aust. J. Exp. BioI. Med. Sci.

41, 247, 1963.
23.

546, 1956.

Underwood, E.J. Trace Elements in Ruman and Animal Nutrition,
New York and London, Academic Press, 3 rd. Edition, 1971.

�-201-

APPENDIX

��Table 5.

Serum calcium, copper, iron, potassium, magnesium, and zinc (ug/m1).
Calcium

Copper

Iron

Potassium

Magnesium

Zinc

2

14.00

.25

15.00

1070.00

5.90

.90

3

31.50

.10

17.50

1350.00

9.80

1.50

4

9.00

.03

5.00

850.00

4.40

.50

5

6.00

.02

24.50

2150.00

12.60

1.40

6

14.30

.01

2.00

400.00

22.00

.50

7

13:90

.01

8.00

1150.00

4.60

.60

Animal No.

I

N

0

8

15.00

.01

3.00

820.00

4.10

.80

14.8±8.07

.06±.08

10.71±8.46

1112.85±547.54

9.05±6.54

.88±.41

w
I

�Table 6.

Calcium concentration

Animal No.

Kidney

Spleen

Heart

Liver

Lung

Skeletal Muscle

2

.34

.18

.17

.13

.36

.14

3

.42

.22

.18

.16

.31

.17

4

.34

.20

.13

.16

.38

.15

5

.33

.22

.04

.15

.17

.18

6

.29

.20

.14

.13

.33

.08

7

.27

.19

.01

.12

.30

.13

8

in tissue (mg/G).

I

N

0
.po

.47

.23

.18

.17

.50

.15

.35±.07

.20±.01

.12±.06

.14±.01

.33±.09

.14±.03

----------------------------------------------------------------------------------------------------------

I

�Table 6.

Calcium concentration

in tissue (mg/G) (continued).
Abomasum
(Greater Curve)

Jejunum

4.62

.63

.52

1.11

4.44

.36

.80

.78

6.11

1.09

.46

.24

1.03

.88

1.16

4.82

.35

.43

1.43

1.61

5.20

3.09

.35

.57

2

1.27

1.24

4.91

3

1.03

1.21

4

1.03

5
6

Animal No.

Abomasum
(Pylorus)

Thymus

Neck Hair

Rump Hair

I

N

7

1.23

1.26

8

1.29

1.18±.15

.38

1.99

4.87

2.42

.78

4.49

1.38

.12

1.17

1.10±.30

3.97±2.00

3.12±1. 55

.37±.15

.81±.59

0

VI
I

�Table 7.
Animal No.

Copper concentration

in tissue (ug/G).

Kidney

Spleen

Heart

Liver

Lung

2

20.01

2.65

13.66

3.64

8.46

5.82

3

12.44

2.54

15.73

13.46

7.60

4.86

4

4.92

3.50

14.35

15.37

4.65

4.01

5

10.29

4.05

15.74

14.13

8.57

6.86

6

13.40

4.52

18.26

18.27

10.66

12.91

7

13.23

4.34

14.68

27.77

11.13

9.42

10.06

2.55

18.72

16.16

11.83

3.77

12.05±4.55

3.45±.87

15.87±1.93

15.54±7.13

8.98±2.47

6.80±3.31

Skeletal Muscle

I

N

0

0\

8

I

-------------~-------------------------~-----------------------------------------------------------------

�Table 7.

Copper concentration in tissue (ug/G) (continued).
Abomasum
(Greater Curve)

Abomasum
(Pylorus)

Thymus

Jejunum

Neck Hair

Rump Hair

7.61

6.43

15.30

6.80

12.67

3.02

2

3.90

7.05

9.80

.52

.97

9.14

3

4.34

5.10

18.22

.77

8.43

6.20

4

3.71

9.18

47.68

.87

9.24

32.87

5

1.81

3.94

8.81

10.32

17.45

9.61

6

Animal No.

I

N

5.53

5.07

14.32

6.06

13.20

25.23

7

3.17

4.01

13.79

1.97

2.41

4.33

8

4.29±1. 84

5. 81±1. 87

18.27±13.35

3.90±3.83

9.19±5.92

12.91±11.48

0
....•
I

�Table 8.
Animal No.

Iron concentration in tissue (ug/G).
Kidney

Spleen

Heart

Liver

Lung

2

31.42

83.54

40.27

45.51

66.51

13.11

3

31.10

50.23

43.00

23.43

71.25

6.67

4

23.88

37.51

26.18

40.89

62.71

6.82

5

48.69

30.99

26.24

45.09

67.22

7.91

6

54.87

46.85

28.66

24.02

39.99

8.72

7

55.12

49.80

28.51

21.36

101.19

13.85

8

Skeletal Muscle

32.83

31.36

26.37

20.42

68.25

8.17

39.70±12.83

47.18±18.00

31.31±7.16

31.53±11.66

68.23±17.91

9.32±2.93

---------------------------------------------------------------------------------------------------------

I

N

0
00
I

�Table 8.

Iron concentration

in tissue (ug/G) (continued).
Abomasum
(Pylorus)

Thymus

Jejunum

38.26

23.82

53.88

26.80

80.70

10.42

6.58

35.04

28.57

129.26

91.88

49.36

18.06

29.87

18.60

5

91.29

108.37

18.23

24.79

30.80

17.48

6

36.38

29.62

20.66

21.50

34.97

28.84

Neck Hair

Rump Hair

2

95.22

150.25

3

32.97

4

Animal No.

Abomasum
(Greater Curve)

,
,

N

7

72.28

34.89

18.57

30.34

38.78

16.79

8

34.43

107.80

17.93

5.93

12.08

14.96

72.88±35.37

86.2l±42.73

24.77±13.76

18.7l±9.28 33.63±12.42 21.64±6.00

0

\0

�Table 9.

Magnesitnn concentration

in tissue (mg/G).

Animal No.

Kidney

Spleen

Heart

Liver

Lung

2

.83

.72

1.02

.63

.65

1.00

3

•BO

.89

1.09

.66

.63

1.22

4

.75

.87

1.05

.65

.77

1.31

5

.77

.99

.74

.66

.89

1.26

6

.68

.97

.99

.64

.77

1.11

7

1.43

.B3

.83

.64

.60

1.10

8

.75

.88

1.08

.67

.63

1.25

•85±. 25

.B7±.08

.97±.13·

.65±.01

.70±.10

1.17±.11

Skeletal Muscle

I

N
•.....

0

---------------------------------------------------_.------------------------------------------------------

I

�Table

9.

Magnesium concentration

in tissue

(mg/G) (continued).
Abomasum
(Pylorus)

Thymus

.89

.95

.84

1.85

.40

.36

.21

.63

.91

.36

.25

1.67

.31

.99

1.53

5

.30

.62

.89

.75

1.23

1.04

6

.36

.39

1.02

.94

1.05

.96

Neck Hair

Rump Hair

2

.38

.55

3

.24

4

Animal No.

Abomasum
(Greater Curve)

Jejunum

7

.36

.38

1.35·

.85

1.35

2.31

8

.41

.26

.79

.63

1.00

1. 73

.34±.05

.40±.13

.99±.41

.66±.29

1. 01±. 23

1.47±.52

I

N
to-'
to-'
I

�Table 10.

Potassium

concentration

in tissue (mg/G).
Skeletal Muscle

Kidney

Spleen

Heart

Liver

Lung

2

10.70

11.96

4.75

7.20

10.88

9.71

3

10.10

15.10

10.98

6.77

10.68

7.65

4

10.85

11.25

10.21

7.61

11.24

8.02

5

10.43

10.03

7.83

6.91

11.91

10.81

6

8.80

13.18

11.68

6.27

10.96

12.39

7

16.53

13.13

5.87

7.26

10.62

12.74

8

9.78

4.64

10.22

7.08

10.23

11.18

11. 02±2. 52

11. 32±3. 35

8.79±2.67

7.01±.42

10.93±.53

10. 35±1. 99

Animal No.

I

N
J-I

N

------------------------------------------------------------------------------------------------------------

I

�Table 10.

in tissue

(mg/G) (continued).

Neck Hair

Rump Hair

Abomasum
(Greater Curve)

2

5.57

6.43

5.00

3

3.41

6.75

4

4.83

5
6

Animal No.

Potassium

concentration

Abomasum
(Pylorus)

Thymus

Jejunum

.89

8.45

10.23

2.63

2.37

9.96

4.28

2.50

15.18

3.37

18.90

6.57

5.10

5.69

5.32

4.31

18.99

13.29

5.45

7.56

6.97

5.80

19.2;3

5.76

l

I:

,...
N·

7

4.92

5.49

3.53

5.26

19.87

6.83

8

9.03

7.52

2.60

8.11

17.13

6.30

5. 47:!:1.
72

S.99:!:1. 73

S.89:!:4.38

4.30:!:2.37

16. 07:!:4.
78

7.60:!:3.08

W'
I

�Table 11.

Phosphorus concentration in tissue (mg/G).

Animal No.
2

Kidney

Spleen*

Heart

.48

ND

.53

Skeletal Muscle

Liver*

Lung

ND

.55

.39

I

3

.58

ND

.55

ND

.58

.37

4

.59

ND

.60

ND

.48

.46

5

.60

ND

.16

ND

.46

.55

6

.54

ND

.46

ND

.35

.53

7

.34

ND

.56

ND

.36

.48

8

.57

ND

.59

ND

.35

.32

.52±.09

ND

.49±.15

ND

.44±.09

.44±.08

----------------------------------------------------------------------------------------------------------

*Not determined.

N

I-'
~
I

�Table 1l.

Animal No.

Phosphorus concentration in tissue (mg/G) (continued) •

Neck Hair

Rump Hair

Abomasum
(Greater Curve)

Abomasum
(Pylorus)

Thymus

Jejunum

2

.41

.50

.32

.27

.23

.55

3

.45

.55

.42

.16

.57

.48

4

.35

.42

.36

.18

.53

1.50

5

.52

.44

.34

.26

.47

.57

6

.54

.46

.38

.34

.55

.53
I

N

.35

.27

.54

.46

7

.35

.40

8

.55

.54

.40

.25

.46

.42

.45±.08

.47±.05

•36±.03

.24±.06

.47±.1l

.45±.14

•.....
V'I

I

�Table 12.Sodiwa

concentration in tissue (illIG).

An1llal No •.

IClclney

Spleen

Heart

Liver

Luna'

2

8.96

4.27

5.00

4.24

6.89

.68

3

7.38

4:.13

5.21

2.42

6.57

2.19

4

8.32

3.50

4.98

4.28

6.14

1.92'

5

6.95

3.77

1.22

4.20

6.60

2.04

6

5.76

3.81

1.32

2.78

6.96

1.74

7

8.26

4.07

3.04

2.67

6.32

2.07

9.92

1.74

4.15

2.47

8.41

1.19

7.93±1. 36

3.61±.86

3.56±1. 72

3.29±.89

6.84±.74

1.69±.55

Skeletal'!Iu8cle

I

N
,,...
0\

8

---------------------------------------------------------------------------------------------------------

I

�Table 12.

Sodium concentration

Animal No.

Neck Hair

Rtmlp

2

1.08

1.07

3

1.46

4

(mg/G) (continued).
Abomasum
(Greater Curve)

Abomasum
(Pylorus)

Thymus

Jejunum

4.76

7.75

8.98

5.95

1.15

2.75

2.76

6.31

4.00

1.38

1.14

2.50

1.55

7.39

3.34

5

1.39

.91

3.23

3.34

5.13

5.59

6

1. 72

1. 38

4.30

5.23

6.68

4.32

7
8

1.99

Hair

1.16

2.60

6.01

6.78

7.35

.94

3.84

4.87

4.15

5.01

3.54

1.42±. 35

1.52±1.03

3.57±1.04

4.39±2.10

6.61±1.36

4. 87±1. 47

I

N
•....

-...J
I

�Table 13.

Zinc concentration in tissue (ug/G).

Skeletal Muscle

Animal No.

Kidney

Spleen

Heart

Liver

Lung

2

208.24

96.84

82.98

273.09

91.91

182.84

3

87.09

136.69

77.78

238.69

83.91

123.54

4

98.73

92.28

83.40

162.59

73.82

122.85

5

93.35

146.51

84.99

103.72

445.75

175.68

6

65.16

105.23

89.19

200.23

115.54

124.64
I

7

104.74

108.66

72.56

106.81

80.95

130.72

N

t-'

00
I

8

77 .13

92.94

70.42

126.04

106.93

213.71

104.92±47.46

111.30±21.74

80.18±6.85

173.01±66.52

142.68±134.44

153.42±36.89

----------------------------------------------------------------------------------------------------------

!;

�Table 13.

Zinc concentration in tissue (ug/G) (continued).
Abomasum
(Greater Curve)

Abomasum
(Pylorus)

Thymus

Jejunum

159.43

115.70

41.20

115.23

97.37

42.67

27.70

50.53

29.72

94.30

100.23

208.83

31.27

62.98

111.63

5

80.46

87.54

109.40

97.30

102.69

118.91

6

97.33

98.73

94.76

103.23

80.73

105.76

7

110.74

97.28

67.23

86.18

87.27

103.31

8

86.07

73.87

88.31

81.55

96.68

94.51

95.83±11.21

93.39±9.57

110.09±56.68

77.56±34.69

74.54±23.51

97.01±30.75

Animal No.

Neck Hair

Rump Hair

2

92.50

98.73

3

109.43

4

I

N

I-'
\0
I

��-221-

JOB PROGRESS
REPORT

State
Project

of

Colorado
----------~--~~~--No.

W-l.24-R-l

Work Plan No.

1

Raptor Investigations
Job No.

Job Title:

Raptor Populations

Period Covered:

February 1, 1972 through january

Personnel:

1

and Characteristics

Studies

31, 1973

Gerald R. Craig

ABSTRACT
Four inventory methods were investigated
to monitor populations of '·wintering
raptors.
These were: 1) statewide eagle census by fieldmen,
2) aerial
flights
for eagles in northeastern
Colorado,
3) 56 square mile censuses
in northeastern Golo-rado, and 4) ground transects.
The aerial flights
show merit for monitoring population trends and age structure
of eagles,
while the ground transects and area counts should establish
chronology of
. movement and habitat preferences of smaller raptors.
Nesting investigations
were initiated
and total of 59 golden eagle nests and 34 prairie
falcon
eyries were located.
The productivity
of golden eagles averaged 1.6 young
per nest in Northwestern Colorado, .5 young per nest in Northeastern Colorado
and .4 young per nest in Southeastern Colorado.
Productivity
of prairie
falcons averaged 2.9 in Northeastern Colorado and 3.7 in east-central
Colorado.
Jackrabbits
and cottontail
rabbits accounted for the majority (91%) of the l'rey
ite~s present in the golden eagle nests of northeastern
Colorado.
Shooting
accounted for 37%and starvation and collision
each accounted for 8% of the
causes of mortality of 62 raptors turned into the Colorado Division of
Wildlife.
August, September and November were the months of highest mortality.
Forty-four prairie
falcons and six golden eagles were amongthe 87 rapt ors
banded.

��-223-

RAPTOR POPUlATIONS AND CHARACTERISTICS STUDIES
Gerald R. Craig
To determine status of current populations, characteristics and trends
including'distribution, migration patterns and chronology of Bald and Golden
Eagles, Peregrine and Prairie Falcons and Burrowing Owls in or through
Colorado.

SEGMENT OBJECTIVES
1.

Develop and refine inventory methods for Bald,and Golden Eagles,
Peregrine and Prairie Falcons and Burrowing Owls in Colorado.

2.

Locate and categorize major and important habitats and associated
Bald and Golden Eagle, Peregrine and Prairie Falcons and Burrowing
Owl populations in Colorado.

3.

Establish and maintain annual or periodic inventories of Bald and
Golden Eagles, Peregrine and Prairie Falcons and Burrowing Owls in
Colorado from inventory methods selected from the literature.

4.

To monitor mortalities of Bald and Golden Eagles, Peregrine and
Prairie Falcons and Burrowing Owls in Colorado and, where possible,
determine causes.

5.

To initiate and maintain a ten year raptor banding program in support
of Project Statement Objective I.

METHODS AND MA.TERIALS
Inventory Methods
Winter Inventories
Four census techniques were selected for testing to establish their
worth as indicators of population densities, fluctuations and trends of
winter raptor populations present in Colorado. While both golden and
,bald eagles were initially designated for census, attempts were made to
census all raptors observed in a given area. These four techniques were:
(1) counts of eagles observed by WCOls in February and March, (2) aerial
censuses from fixed-wing aircraft in northeastern Colorado,
(3) area
censuses of given sample sites, and (4) linear transects from vehicles
traveling along secondary roads.
All WCOls were requested to locate and count eagles observed in their
districts during the periods of February 14-16 and March 1-3. The purpose
was to locate major concentration areas of eagles and establish the
responsiveness of WCOls to raptor censuses. The request was made in the
form of a field order (Appendix A through C) •

�-224-

The aerial census was established in the manner described by Boeker and
Ray (1971) and Wrakestraw (1972). An area of 3,000 square miles was
designated in northeastern Colorado and 13 north-south transects were
randomly selected so that each transect was at least two miles away from
its neighbor (Fig. 1). The transects represented a total of 600 linear miles.
The transects were then flown in a Cessna 185 and all raptors observed
within ~ mile of either side of the aircraft were mapped and recorded
(Appendix D). Thus, the transects accounted for 300 square miles, or a
10% sample of the study area. Upon completion of this count, the aircraft
would then be flown along the South Platte River b.etween Fort Morgan
and Greeley and all eagles sighted along the river were counted. Aerial
censuses were flown in December of 1972 and January and February of 1973.
Incidental eagle sightings were also recorded on aerial censuses for other
wildlife such as antelope, deer, elk and waterfowl and will be compared
for compatability.
One 56 square mile census area and technique that was established by
Dr. R. A. Ryder for International Biological Programme studies on the
Pawnee National Grasslands (Ryder, 1969) was again censused to provide
long term information on the densities of wintering raptors.
In addition,
two other 56 square mile census areas were delineated in agricultural
areas to compare raptor densities and composition.
To accomplish these
censuses, each area was traversed slowly in a vehicle and all raptors
that were Sighted within one mile of the vehicle were identified and
pertinent information such as location and activity was recorded on an
observation card (Appendix E).
Several localities were investigated for potential as linear transects
to establish trends and changes in species composition of raptors and
a census form was developed (Appendix F). Bait stations in the Meeker.
area were established and will be investigated for potential as counting,
trapping and marking stations.
Unpublished data from transects in
southeastern Colorado has been made available for integration into this
study by William A. Anderson of Otero Junior College.

Nesting

Investigations

Nesting investigations were commenced in northwestern Colorado.
In general,
likely areas were surveyed using four-wheel drive vehicles, trail bikes
and foot. All active and inactive sites were mapped and pertinent data
about each site was recorded on forms like that in Appendix G. Similar
work was done by Olendorff in northeastern Colorado under Amendment I
to Job 1, and WilliamA.
Andersen provided data for nest sites in the
southeastern portion of the state.
In addition, all field personnel were requested to complete and submit
nest report forms (Appendix H) for each active raptor nest about which
they had knowledge.
These reports, as well as information obtained from
other sources were investigated where practical.

�-225HABITATS
Information about habitat type, disturbance factors and site descriptions
were recorded on all nest sites that were investigated.
Major vegetation
types and response to disturbances were also noted for raptors observed on
winter inventories.

MORTALITY
Mortality report forms (Appendix I) were distributed to all WCO's in an
attempt to obtain information on dead raptors.
Where feasible, carcasses
were collected·for analysis as to cause of death.
Also, injured raptors that were discovered by the Division (Appendix J),
the public, or other personnel were inspected for the cause of incapacitation
and the birds were turned over to a group that was authorized to care for
and rehabilitate them. For the purposes of analysis, all sick, injured or
incapacitated raptors that were received were considered as mortalities
since, had they not been discovered, they would have died.

BANDING
Federal and state bird banding permits were obtained.
Nestlings were
banded when the investigator visited nest sites to obtain information
on prey items and productivity.
Several trapping blinds were established
in the foothills west of Denver to trap and band adult raptors moving
through on migration.
Raptors were trapped and banded from the vehicle
during winter inventories.
Several Swedish goshawk traps were constructed,
but were not utilized in 1972.

RESULTS AND DISCUSSION
Inventory
WCO Eagle Concentration

Methods

Census

Four hundred and sixty-six eagles were counted by fifty-seven field men
from February 12-18 and three hundred and thirty-three eagles were
counted by fifty-four persons from March 1-6.

Table

1 -- Summary

of the Preliminary

Type

Eagle Concentration

Counts,

1972.

Bald

Golden

Adult
Immature
Unknown

150
17
64

153
39

14

29

SUBTOTAL

231

206

29

Unknown

Count Period: February 12-18
No. of Respondents: 57

TOTAL:

466

�-226Table 1 --Summary
(Cont'd.)

of the Preliminary Eagle Concentration. Count s ,..1972.
.

Type

.

. . :.:

......

Bald

Golden

Adult
Immature
Unknown

101
32

137

o

26
21·

SUBTOTAL

133

184

-

,

,

. ......

UIlknOwn

Count Period: March 1-6
No. of Respondents: 54

TOTAL:

. -15·

333

The observation forms also provided space for the observer to record the
behavior and activity of eagles observed. Although several were seen
pursuing quarry, no eagles were observed catching prey. A total of 59 eagles
were observed feeding on carrion (see Table 2) and in only 10 of 24 instances
solitary eagles were seen on carrion. This indicates that both species are
gregarious and will share their meals.
,-..&gt;

Table 2 -- Eagles Observed Feeding on Carrion February 12-18 and March 1-6,
1972.
SEecies of Eagle
Golden
Bald
1
7
1
1

2

2

Total Number of
Eagles ·Per Carcass
1
7
1
3

2
1

1

1

1

7

7

1

1

3

3

2

2
2

2
5
2

5

3
3

3
3

2

2
1
.2
1
1

1
.1

4
TOTALS:
41

18

2
1
2
1
1
1
1
1- ..

59

Kind of Carcass
Cow
Cow
Calf
Elk
Elk
Elk
Elk
Deer
Deer
Sheep
Sheep
Sheep
Coyote
Coyote
Jackrabbit
Jackrabbit
Jackrabbit
Jackrabbit
Rabbit (Unk. Spp.)
Rabbit (Unk, Spp.)
Rabbit (Unk. Spp.)
Rabbit (Unk. Spp.)
Rabbit (Unk. Spp ,)
Unknown
24

G

-Ii.

�-227-

As was expected, the majority of the bald eagles were concentrated along the
major rivers and .large reservoirs. The accessibility of fish and waterfowl
is possibly the cause for this behavior. The following appear to be the
major areas of concentration although birds were observed elsewhere.
San IllisValley

- 61 observed in the February count below Monte
Vista on the Rio Grande.
- 30 near Riverside, Jackson and Empire Reservoir
complex.
25 scattered along the river from Glenwood Springs
to the Utah line.
24 scattered.along the river from Meeker to
Rangely.

South Platte River
Colorado River Drainage
White River Drainage.

(See Map Showing Bald Eagle Concentrations, Figure 112)
Due to their nature, golden eagles were scattered throughout the state and
no large concentrations were noted. Observations confirmed earlier reports
of major wintering populations. in several areas. The number of golden eagles
observad in North Park was surprising as relatively few birds have been
reported wintering in this area. It is believed that substantial wintering
populations occur on the eastern plains near Wyoming in the northeast and
throughout the southeast quarter of the state.
(See Map Showing Golden Eagle Concentrations, Figure 1/:3)
As expected, the results cannot be used as any indication of actual numbers
of wintering eagles present in Colorado. The technique does show merit
as method of obtaining an index of relative abundance, species composition
and possibly age ratio.' The reports also give insight into activities,
behavior, habitat preferences and prey preferences of eagles.
Aerial Golden Eagle Censuses
Three aerial flights were made in northeastern Colorado, on December 19, 1972,
January 24, 1973 and February 16, 1973 to count golden eagles on a 3,000
square mile study area (Fig. 1). Table 3 summarizes the results of these flights.
Table 3 -- Results of Aerial Flights for Golden Eagles in NE Colorado,
1972-73.
Date

Golden Eagles
Ad. Juv. Unk.

Dec. 19
Jan. 241/
Feb. 16-

22
16
18

6
8
7

4
0
0

Total Area
Estimate

Eagles per
100 Sq. Miles

320
240
250

10.6
8.0
8.3

1./ While this flight was made in project year 2, it is included here for
continuity.

�-228-

In the future, the number of flights will probably be reduced to one in
January. Results from one year to the next will be compared for trend
data. Results from these flights will also be provided to the U.S.
Bureau of Sport Fisheries and Wildlife to enable them to compare these
flights to those made in northwestern Colorado, Wyoming, Utah, Nevada,
New Mexico and Idaho. Another aerial flight should be designated in
east-central or southeastern Colorado for 1974.
Aerial Bald Eagle Census
After the aerial golden eagle census was completed, the investigators would
fly above the South Platte River between Fort Morgan and Greeley, Colorado
and count those bald eagles present along the river. Table 4 provides the
results of these flights.
Table 4 -- Results of Aerial Flights for Bald Eagles in NE Colorado,
1972-73.
Date

Ad.

Juv.

Unk.

Total

% Juv.

Dec~ 19
Jan. 24
Feb. 26

23
18
47

15
13
19

0
0
0

38
31
66

39
42
29

,~

This is a direct count of all eagles seen along the riverbottom. Since
bald eagles tend to concentrate along watercourses where waterfowl are
in abundance, an exact count is feasible.
The adult to juvenile ratio is important since it is an indicator of the
viability of this particular population. Considering that juveniles don't
attain their adult plumage until they are four to five years of age and
that the potential lifespan of a bald eagle may be thirty to forty years,
a population containing thirty to forty percent juveniles would indicate
a high reproductive success.
With a little practice, it is a simple matter to distinguish immature bald
eagles from adult golden. eagles. The former have more mottling and are
not as uniform in color as the golden eagles. The lighter head and neck
feathers of the golden eagles also stand out.
This method shows the greatest potential for obtaining an accurate census
of bald eagles. With their predilection for open water, it is a simple
matter of flying the major watercourses to obtain a very close estimate
of the actual wintering population present.
lJ

Area Censuses in Northeastern Colorado
Three 56 square mile census areas on grassland, dryland farming and irrigated
farming vegetative types were censused at two week intervals from December 17,
1972 through March 10, 1973 (Figure 4, 5, 6 &amp; 7). The counts of February and.

�-229-

•

March 1973 are included here for continuity of data. A total of twenty counts
were conducted. The total number of birds seen on each area was similar but
total biomass was twice as great for the short grass prairie as the wheat
farming and five times greater for the irrigated farmland.
Of the eight raptor species observed, the red-tailed hawks and American
kestrels were peculiar only to .the irrigated site, while ferruginous hawks
(72% of those observed) and golden eagles (75% of those observed) were
present on the short grass prairie site (Table 5) •..Rough-legged hawks
and golden eagles accounted for more than 7fJ% of the total raptor biomass
on each study area (Table 6).

�-230-

Table 5 -- Total Numbers of Each Species of Raptor Counted on Three 56 Sq.
Mile Stud~ Areas During Winter 1972-73.
Pawnee

Site
Briggsdale

Ault

Total

Golden eagle

48

14

2

64

Ferruginous

13

1

4

18

Rough-legged hawk

33

45

30

108

Red-tailed hawk

0

0

5

5

Unidentified Buteo

1

5

9

15

Marsh hawk

4

4

6

14

American kestrel

0

0

23

23

Prairie falcon

6

7

7

20

Merlin

1

2

3

6

TOTAL

106

78

89

273

S:eecies

Table 6 -- Percent TOtal Biomass of Rough-legged Hawks and Golden Eagles on
~hree 56 Square Mile Study Areas During Winter 1972-73.

Site

Rough-legged H'lwk
Biomass
% Total
No.
(kg.)
Biomass

Golden Fagle
Biomass
% Total
(kg.)
Biomass

No.

Pawnee

33

33.0

12.6

48

206.4

75.0

Briggsdale

·45

45.0

39.3

14

60.2

52.5

Ault

30

30.0

54.9

2

18.6

15.8

Linear Transects
Several routes were investigated in the San Luis Valley for potential
andaithough one route.was not settled upon, at least one transect will
be selected. The information will be recorded on forms similar to that
in Appendix F.
Two transect lines were experimentally followed on the praari,e east and
north of Denver. However, these routes will have to be followed-for
several more years before sufficient data will be gathered to permit
comparisons.

Nesting Investigations
Northwestern Colorado
The prime areas of investigation in northwestern Colorado were: Rio Blanco,
Moffat and the western portion of Routt counties. The topography of this

�-231-

region provides numerous escarpments, river canyons, and erosional reminants
that are ideal nest sites for golden eagles, red-tailed hawks and to a
lesser extent ferruginous hawks and Swainson's hawks.
A total of 38 golden eagle eyries were located. The reproductive success for
these nests averaged 1.6 young per nestwhich is comparable to other
localities. Eight of the nests were visited to band the young and note
the prey items. Jackrabbits and cottontail rabbits were the predominant
prey items accounting for 91% of the prey items present.

Table 7 -- Prey Items Present in Golden Eagle Nest in NW ·Colorado.
Prey Item
Number
% of Total
Lagomorphs (unk. spp.)
86
76.8
Jackrabbit

(sp.?)

10

8.9

Cottontail Rabbit (sp .?)

6

5.3

Black-bi11edMagpie
Mule Deer (Fawn)·!.!

2
2

1.8
1.8

Marmot

1

0.9

Sage Grouse

1

0.9

Red-headed Duck

1

0.9

Fish (sp.?)

1

0.9

Snake (sp,?)

1

0.9

Bird (sp.?)

1

0.9

112

100.0

TOTAL

11

Day old fawns, probably still-born, were present in two nests located
in deer habitat. (pinyon-juniper)

Listed in Table 8 are data regarding other raptor nests that were located
in the region.
Table 8

Other Raptors Nesting inNW

Species

~

Colorado.

Nests

Young

Young/~est

Ferruginous hawk

2

6

3.0

Swinson's

hawk

2

4

2.0

Red-tailed hawk

28

57

2.0

�-232-

Northeastern

Colorado

Information discussed here is covered in greater detail in the 'report
submitted as part of the requirements of Job 1, Amendment I.

Table 9

Breeding Densities and Nesting Success of Raptors in NE Colorado.
No. of Pairs

Species

Pairs/IOO Sq. Mi.

Successful Pairs

Golden Eagle

12

1.20

5 (41.6%)

Prairie Falcon

14

1.40

11 (78.6%)

Ferruginous Hawk

26

2.60

16(16.6%)

Red-tailed Hawk

8

0.80

?!j

Swainson's Hawk

68

6.80

38(56.8%)

Great Horned Owl

30.5

3.05

8(29.1%)

All Species

158.5

15.85

78(53.5%)

!/

Productivity was not determined.
,-t:)

Table 10

Productivity of Raptors in NE Colorado.

Species

Young
Produced

Young/Pair
w/Known Production

Young Produced/
100 Sq. Miles

Golden Eagle

6

0.50

0.60

Prairie Falcon

41

2.93

4.10

Ferruginous Hawk
Red-tai led HawJ;./

48

1.85

4.80

(8)

(1.00)

(0.80)

Swains on's Hawk

80

1.19

8.00

Great Horned Owl

18

0.65

0.69

All Species

201

1.37

20.10

!/

Estimated productivity.

East Central Colorado
This region was surveyed only to locate pra~r~e falcon eyries and attempt to
establich productivity. A total of 20 eyries were located and the data is
summarized in Table 110

�-233-

Table

11 -- Productivity

Falcons

in East-Central

Site

Clutch Size

No. Produced

1
2

5
5
5
5
5

4

0

3

••

of Prairie

4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20

1/

Colorado.
No. Fledged

2

(undet ermined)
(undetermined)
(unde"termined)
(undetermined)
(undetermined)

4

(undetermined)

but not breeding
(undetermined)
(undetermined)
(undetermined)

(undet ermined)
(undetermined)
(undetermined)

4

4
(undetermined)

Inactive

5
Pair present,

3
3
5
(4)})

4
4
5
5
1
4
2
4
5

(4)·Y
(5)1/
(5)·Y

(l)·Y

(4)))
(2)1/

(4)])
(5)!/

0
4
(undetermined)

5
1
4
2
4
5

Estimated from later visit after young had hatched.
minimum figure.

Average clutch size -- 4.17.
Number of young produced per pair
Number of young produced -- probably

Southeastern

This represent

a

3.71.
quite close to 3.71 figure.

Colorado

The area of this study consists of the Comanche National Grasslands and in
the vicinity of LaJunta, Colorado and northward.
The following informati7on
was provided by William C. Andersen of Otero Junior College.

Table

12

Productivity

SEecies
Golden

cj

Eagle

in SE Colorado.

Nests

Young Produced

9

4

0.44

Young/nest

Ferruginous

Hawk

19

41

2.19

Red-tailed

Hawk

11

12

1.09

Swainson's

Hawk

113

157

1.85

Kestrel

3

7

2.33

Kite

3

2

0.66

Great. Horned Owl

36

43

1.19

White-necked

15

44

2.93

American

Mississippi
~

of RaEtors

Raven

�-234-

--

Table 13
Swmnary of Dead arid Injured Raptors Turned into the Division
During Jan. - Dec. 1972.
.u
Il-4

.u

III

0

.....,
&lt;IJ

~

0
.r-!

.u

::l

bO

~
...•

0
0

co

.r-!
.....•
.....•

0

u

&gt;

.u
"'"
&lt;IJ
.....•

S,eecies

tI.l

r::l

-e

Golden Eagle

2

2

1

.u

0
.u

::l

-'

Bald Eagle

,!J
0

0
'r-!

III
"'"

0

-

~

.u

III

0
"'"

III

.bO
&lt;IJ
.....•

M

~
&lt;IJ
~

.....•.

E-I

u

0
...•
co
...•

.....•

tI.l

u.

0

M
H

III

f'"'

'r-!
"'"

~
~

0

.c

:&gt;.
.....•
.....•

~
~

...•.
co
0

...•

.....•
0

3

~
~
~
~

.....•
III
.u

::&gt;

H

2

10

3

Goshawk

2

Marsh Hawk

1

0

3
1

3
3

4

1

2

5

4

9

Swainson's Hawk

1

1

2

Rough-legged

1

Ferruginous

Hawk

1

-

Red-tailed Hawk

-

Hawk

1
1

Pigeon Hawk

1

American

1

2

·1

5

4
3

3

10

1

2

8

1

1

Owl

....

Screech Owl
TOTAL

5

2

1

Great Horned Owl

Short-eared

1

3

Kestrel

Barn Owl'

0

1

Peregrine Falcon
Prairie Falcon

-e

2

2

23

5

5

1

1

1

1

1

19

62

\...._~

Of a total of 62 raptors that were turned into the Division, 37% were shot,
8% were starving, 8% had collided with some object, 3% were electrocuted,
3% collided with automobiles and 30% were injured or died of undetermined
causes. All of the bald eagles (3 individuals) were shot, the majority
(4%) of the prairie falcons suffered injuries from collisions with objects.

�-235Figure 8 -- Breakdown by Month of Dead or Injured Raptors Reported to the
Division.
No. of Raptors

Jan

Feb

Mar

Apr

May

Jun

Ju1 Aug

Sep

Oct

Nov

Dec

108-

6-

420-

This graph roughly shows the seasons of highest mortality, assuming that
·al1 other factors, such as percentage of persons reporting injuredraptors,
remains constant. Generally, late winter is the time of least mortality,
most the raptors have already encountered various factors that adversely
affect them. Mortality drops sharply when the adults enter the breeding
season in the spring. Mortality increases as juveniles fledge and encounter
problems, and reaches its peak in the fall when migration commences and
naive juveniles move into new territories.
Unfortunately, the mortality factors, such as shooting and weather. do not
remain constant. The number of gunners in the field increases greatly in
the fall, thus more raptors are shot. In the fall, the raptors encounter
harsher weather conditions and survival is difficult. Ideally, mortality
should drop late in the winter when raptor acclimate to the harsh surroundings
and become wary of humans.

BANDING
No special point was made to attempt to band raptors specifically for that
purpose in 1972. Those raptors that were encountered in the course of study
that were available were banded. In the future, when more intensive work,
such as nest site viSitations, is undertaken, more nestings will be encountered
and banded. Table 14 lists those .raptors banded in the course of study.
Table 14

Raptors Banded by'the Division in 1972.

Species

;j

Number Banded

Prairie· Falcon
Red-tailed Hawk
Goshawk
American Kestrel
Golden Eagle
Swainson's Hawk
Coope r 's Hawk
Ferruginous Hawk
Great Horned Owl
Rough-legged Hawk

44
14
9
7

TOTAL

87

6
3

1
1
I

1

�-236-

LITERATURE CITED

Ryder, R.A. 1969. Diurnal Raptors on the Pawnee Site. U.S. IntI.
BioI. Prof., Grassland Biome Tech. Rep. 26, l6p.
Boeker, E.L. and T.D. Ray 1971. Golden Eagle Population, Studies in the
Southeast. Condou, 73, (no 4) 463-467.
Wrakestraw, G. F. 1972. 1972 Wyoming Bald and Golden Eagle Survey.
Completion Report, Game Bird Survey Proj. No. W-50-R-2l.
Title: Establish Number of Bald and Golden Eagles. Proj.
Objective No.7, Job No. 31.

Job

a

15

�'o.l.

..t'

~

z.

.,;.

~.

Figure 1 -- Aerial Eagle Census of NE Colorado

._'"!W!r..:.....

5

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�-240Figure

4 -- 56 Sq. Mile Census Areas

~

.

.. ---------

~.

CHEYEl'!NE

Wyoming

--------------------

Neb!a~k:.

ffiIJ
FORT CCLLINS
~

STUDY· AREAS

'n
a

II

.~

Prairie

Dryland

.Irrigated

, 16 miles,

_

�-241Figure 5 -- Pawnee 56 Sq. Mile Census Area.

-f

· J~
~

(j
.····

.·

~
.. ~

'.f

~

!

.~

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J~
I

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tart

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KEY
~

r

Census Route
~

Standing Water

~.

Intern)i t tent Water

t mile
•

-

-

• Q

�-2426 -.:. Ault 56 Sq. Mile Census Area.

Figure

.

,

....

Fini:. h
.. ~.

IT

~~

";

4l11IP

,
..•

4I~
iIr

~

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f
:;,

1 mile

,,!)a.

�-243Figure

7 -- Briggsdale

56 Sq. Mile Census Area.

,.,
....,.~.

Sh.rt

.

Fi11 sh
'I··
"

Ql.
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,

IT

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a

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r

Hater

1 roDe-4

�-244-

APPENDIX A
Use separate

form for each eagle or group of eagles.
MLD

Date

&amp; GOLDEN EAGLE OBSERVATION FORM
Time

--------------------

Species

and No. P~esent:

Bald

_

lJo. of Adults

_

No. of Unknown
or Activity

Location
drainage

(Direction in miies
or i~poundllient)

Weather

Condition

Other

_

No. of Juv.
_

Behavior

Eab Lt.at;

Golden

---------

Unknown
Age:

_

_

from ne3.rest tOvm or other

(Hind, Cloud Cover,

landmark,

PPT)

.-----------------------------------~-~~. ----

Not es.

Comments

--'

_

---- ---------------------

Observed

by

_

i!

�,..245-

.

APPENDIXB

KEYTO THE FIELD IDENTIFICATIONOF BALDANDGOLDEN
EAGLES
The Bald and .Golden Eagles are among the largest birds of prey of North America.
The wingspan of the Bald Eagle approaches seven feet while that of the
Golden Eagle is nearlrsix and one half feet.
The only North American bird of
prey larger than the eagle is the California
Condor whLch is restricted
to
Southern California.
BALDEAGLE
Adult

Haliaeetus

leucocephalus

-- Body and wi.ngs completely brownish black.
pletely~.
Bill,
eye and feet are bright

Head and tail
yellow.

are com-

Emmat.ure -~ Body, wings, head and tail are completely br own, but not as da r k
as the adult.
Attainment of Lie adult plumage is gradual with the head
and tail becoming white at four or five years of age.
Eye and cere*
are pale ye l l.owi.sh grey; bill
is br own; legs are pale yellow.
Field

Ch~:!:~cte~§.-- II!lI'Jature Btlld Eagles may be confused with the adult
Golden Eagle, but can be distinguished
by the larger 1!.eac!~heavier
andunfeather.ed
vellmv legf:..

GOLDEN
EAGLE

Aguila

bill,

chrysaetos

Ad~lt;. -- Crown and nape feathers
edged. w:tth golden buff (which gives the head
The rest of the body, tail and wings are dark
a "Golden ·appearance).
brown. May"be a slight \"lh_itish' patch on the und erwi.ng at the "wr Lst "
joint.
Legs are feathered
down to the toes.
Cere* and feet yellow.
ll

Immature -- The cr ovn and nape are darker than the adult (doe sn ' t appear as
"go Iden"},
Tail feathers
are basally whi.t e ''lith 2/3 the terminal
portion blackish.
Both on the top and under the wing is a conspicuous
white patch formed by the whLt e bases of the inner primaries
and secondaries ~ Eye is darker br-own than the adult;
cere* and feet are yellow.
Leg is feathered
down to the toes.
Field

Characte~..§. _.. Golden Eagles can be distinguished
from either
adult or
immature Bald Eagles by the feathered
legs down to the toes, but thi s
characteristic
is usually ev Lderrt only at close r ange , Immat.ur e Golden
Eagles can be distinguished
from either
adu Lt; 0: Lmmatur e Bald Eagles
by the large areas of white at the baG€.:. of the tG.:i.l and "wrist"
joint
of the wing in flight.

~~The fleshy,
the nostrils

membranous covering
open.

at

the base

01:

t.he upper

beak through

whLch

�-246-

APPENDIX B (Continued)

KEY TO NORTH AMERICAN EAGLES
1.

2.

a.·

Basal 1/3 of tail \\Thite;large patch at "wrist"
of wing • . • • •
• IMMATURE GOLDEN EAGLE

b.

Tail all white or all dark; white patch small .

n.

Tail and head all white; bright yellow legs and
bill

3.

. . . .

. . . . . . . .

• • 2

ADULT BALD EAGLE

h.

Tail and head not white • • '.

a.

Completely brown eagle with "Golden" head and
small ~.Jhitishpatch at wri~t on underwfng ; legs
feathered . . . . . . . . . . . . • •

ADULT GOLDEN EAGLE

Completely brown eagle with !!£ white; light
yellow legs are feathered • • • • • •

IMHATURE BALD EAGLE

b.

• . 3

If

�-247-

APPENDIX C

HA.BITATS PREFERRED' BY EA.GLES

Bald Eagles:

Bald Eagles arrive in Colorado during the fall migration and
soine remain throughout the winter, departing in early Spring.
Most of the Bald Eagles inColcrado belong to northern
nestidng populations.
They are often gregarious and from two
to nearly a hundred may congregate near open water such as
large river drainages and large reservoirs. A. diet of fish
is preferred and winter-killed fish will attract them. They
may also frequent areas where waterfowl concentrate and feed
on the dead or crippled ducks and tg eese , They can be seen
standing on. the ice near open water or perched in bare trees
and snags surrounding the reservoirs.
Bald Eagles have been reported on the eastern pra Lr'Les with
up to half a dozen birds in a solitary tree. In these cases,
tl:J.eyprobab
ly feed on carrion and jackrabbits. Large Iivestock die-offs will also concentrate both Bald and Golden
Eagles.

Golden Eagles: In Colorado, Golden Eagles are more numerous than Bald Eagles
and may be found throughout the state from above timberline
tbthe prairies. They are especially abundant on the prairies
and open mountain parks in the winter. Golden Eagles do inhabit forested areas, but they are not abundant and they are
difficult to observe, so such areas generally should not be
included in winter census activities.
Golden. Eagles take more live prey than Balds but will feed
on Carrion when.the occasion presents itself, so they may
also be found in:'the company of Bald Eagles. Because of this,
care nrustbe' taken to avoid confusing adult Golden with immature
Bald Eagles.

�-248Appendix
AERIA.L EAGLE
Date:

--

_

Locaticn:

D
CC:SUS

Flight

A.M.

Time:

to
-----------_
P.M.
to

_

Crew:
WE.~THER:

Start

--

Temp.

Mid.

--

Temp.

End

--

Temp.

_

Wind
_

_

Visibility

Cloud

Cover

Wind

_

Visibility

Cloud

Cover

Wind

_

Visibility

Cloud

Cover

_

----

----

Counting

c~nditions:

Trans.j

f&gt;.':"I.D r~\G~_S
_-:::G...::.G=.:i[)~!c::.::r---=-·:..\:...:G;.=L.::..i:'_;_-:_Other
Coyotes &amp;
Ur:ident.
Buteo
J
?
Adult
I~;::.
I
Eagles
Hawks
Ar::..: 1t
....::.;:!:
·~-+----.:.._+-=:::..::...::'__-+-_=:::;.:.._+---..::==~f__-==.::..:.:.:==--_+_-=R=a~=.;:.t.::..or:..:s~--=.A:.:.n.:..::t:.::::e-=-l
~o;J::....:e=--

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!

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i

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i

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G. - GrOU;1c!

F .1' .

o.e

j

_----1

- F'enc e Post
- Outcropping

__

-

~-. __ .: '\
--1...L_

T.
Tree
O. - Ot he'r

/J

�-249-

Appendix E

OBSERVATION CARD

Band No.

Species

Sex

No. present

Date

Location

f1f'

Weather
Behaviour
Observed or collected by
Other notes

J

Vegetation

Age
Time

�.
..
Appendix F

DATE:

-250-

OBSERVERS:

TIME START:

MILE.o\GE START:

TIME

I1ILFAGE

&amp;'W:

END:

WEATHER:

TINE

-

MILFAGE

l

SPECTES

I

I

ACTIVITY

• u~

,-

I

I

I

,

I

j

I

I

I
l

I

I

I

i

..

I

I
I

i

I

"

j

I

I
i

I

[

I

i

!

I
i

I

I

!

I

l

I

1

!

I

,

!
I

I
I

I

I
--, !

f

I

I

l==~1--~---~--'-'~:~------r-----~------.~--------~

--

I

i

!

j

I

I

I

I

I

!
i

i

I
I

I

I

i
I

I

I
I

I
I

I

i

I

..

I

~

!

r=

:

!_

i
.---.L--.-.
---'-----,_L

LI

, ,----+------,-

,

I
~I'-----

,----

I
..;..------+------------'------I

'

(

\.

�Appendix G
Date

NEST DATA

,J

Species

_

Time

_

Landowner

Location: T

N or S;

R,

_

W; Sec.

County

I.

-251-

_ ___

,,"--_1/4

Additional Details

Supporting Structor Information
CLIFF OR OUTCROPPING
Pothole: width
Ledge:
width~

height

_

_

len~th

_

_

1ength,

-----------

Over'hang
: width,
Other:

~

_

depth

_

ht to overhang

_
_

Direction of Avis

--------------------------

~eol"gy:_ _:_----------------------------------------------TREE
~:I)ecies:

------------------------

rio _ of trees in vicinity

height

DBH~

_

_

c~m":R STRUCTURE (describe)

,.--

_

I1c;tails
of nest
Ht. above ground:..--.

Ht. to top of structure

Exposure (directLou)

_

Size:

_

Nest Materials

----~-------------------------~-----------------

II.

Habitat nata
Topography

-:__~

~

-------

Dominant Vegetation~

_

GeologicFormations

~

~

Drainages/Impoundments: Direction:..--.------------ Distance~

_

Soil types~----------------~---~-------------------------Land Use-----:~-

_

IiI.,Behavior
Adult s Activity;

Male

----~--------------------------------

Female
Young Activity:

------------------------------------

�-252-

IV.

Prey Items
Species

No. of Individuals

Portion Remaining

Unidentified item I.D. No.
V.

_

Eggs/Young
No. Present
Band No.

Age~

_

-------------------------_ Sex

_

o

VI.

Nest Failure
No. of unhatched eggs
No. of dead young,

-------

_

No. of destroyed eggs_·
Age.

Cause of Destroyed eggs~

_
~

Cause of Dead Young,

_
~

~

Remains and Shell I.D. No.
VIi.

Disturbance Factors
type
1oad/Highway
type

TC'lil/Track
h::: b itation:
Other Human Activities
VII.

_

_

~~--------------

_
_

Degree Use.~

_

Dist.&amp; Direct.

Degree Use.

_

Dist.&amp; Direct.

Dist. &amp; Direct •
~

------------_
_

Additional Comments:

---_.------

- .-. - - -------_.

--.---------------------------------

�-253Date:

_

Property Owner Data:
Name:

------ Phone:

_

Mailing
Address:

_

Location of Residence:

_

Attitude toward raptors and willingness

to protect raptors nesting on

his property:

-----------

Att Lt.ude toward pub lic access :

--..,.-

-------------

Description (sketch map) of land he holds in the vicinity of the nest
site(s).

List of rapt or nests on his property
Species

Location

Were the following left with the landowner?

,=1

Division Identification Card

Yes

'Violation Report Card

Yes 0

NoP
No 0

�· --254Appendix H

BIRD 010· PREY NEST· REPORT

Species

~-- __ --

Observer

_

Discovery Date

--------

_

r

------------------------------------------------

Phone number

and where may be contacted

--------------------------------

Exact location of nest (legal description by township, range, .section and 1/4
s ec t Lon , a Lso give :lirection from a known landmark.
\.Jheneverpossible, include
map show i ng access):

Property owne r (if known)
Description of surrounding habitat

Description of nest (supporting structure, aspect, height above ground)

Activity at nest (building, incubating, etc)

~~

~~

__~

_

Reffiarks

------------------------~----------~----------------

Mail

form to:
Gerald R. Craig
.Colorado Gawe, Fish &amp; Parks Division
6060 Broadway
Denver, CO
80216

[!

�-255.'

Appendix I
Colorado Division of
Wildlife

.~.

RAPTOR MORTALITY REPORT
1.

Date of discovery

2. Approximate date when mortality occurred
3. Location (county, nearest post office, township, name of electric utility and
pole number)

4. Species, age, sex (if known) of each bird found
.5.

Probable cause of mortality (electrocution, gunshot, etc.)

6.

Vegetation (forest, grassland, etc.) and terrairt (hilly~ flat, etc.)

7.

Check pole type from those below

N6 crDssarm
----~

. 8.

No crossarm,
but transformer
present

___

Crossarm(s)

Disposition of carcass (left, picked up, etc.)

9. '.Name. address, phone numbe.r and agency of person making report

Send Report to: Hildli fe ~!anagemcnt
6060 !!roaJway
Denver, CO 80216

Other
---(describe on
back) .

�-256-

Appendix J

REPORT OF INJURED RAPTORS

The Colorado Division of Hi1dlife requires' this form to be completed upon the
acquisition

of each raptor that is accepted.

This form should be mailed 19ith-

;

in 48 hours to:
Gerald R. Craig, Raptor Biologist
Colorado Division of Wildlife
6060 Broadway
Denver, CO

80216

(Phone:

825-1192)

Species of raptor:
Information on party that first found the r.apt6r
Phone

Name:
Address:
(Street)
Locality where the raptorwas

(City)

(Zip)

(State)

discovered:

Date of Discovery:
Details of injury:

Information

on party that brought the rapt or in (if you know the individual,

just provide his name):
Name:
Address:
(Street)
Raptor received by:

(City) .
________________________________ ~

(State)
~Date.~

(Zip)

--

----

(f

�-257-

FIGURES
111

Map of Aerial Eagle Census of NE Colorado.

#2

Map of Bald Eagle Concentrations

#3

Map of Golden Eagle Concentrations

114

Map of 56 Sq. Mile Census Areas.

115

Pawnee 56 Sq. Mile Census Area.

llq-

Au1t 56 Sq. Mile Census Area.

#7

Briggsdale 56 Sq. Mile Census Area.

118

Breaksown by Month of Dead or Injured Raptors Reported to the Division.

TABLES

t

J

1fl

Summary of the Preliminary Eagle Concentration Counts, 1972.

1foZ

Eagles Observed Feeding on Carrion February 12-18 and March 1-6, 1972.

113

Results of Aerial Flights for Golden Eagles in NE Colorado 1972-73.

114

Results of Aerial Flights for Bald Eagles in NE Colorado 1972-73.

115

Total Number of Each Species of Rap t or- Counted on the Study Areas
During Winter 1972-73.

116

Percent Total Biomass of the Rough-legged Hawk and Golden Eagle on the
Study Areas During Winter 1972-73.

1F?

Prey Items Present in Golden Eagle Nests in NW Colorado.

118

Other Raptors Nesting in NW Colorado.

119

B'reedLng Densities and Nesting Success of Raptors in NE Colorado.

1110

Productivity of Raptors in NE Colorado.

1111

Productivity of Prairie Falcons in East-Central Colorado.

1112

Productivity of Raptors in SE Colorado.

ns
..

.,

1114

Summary of Dead and Injured RaptorsTurned
During Jan. - Dec. 1972.
Raptors Banded by the Division in 1972 •

into the Division

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Jon

State
Project

COLOl&lt;..'1 i":0.

01:

_

Game Bird Survey

1

\&lt;Jork Plan No.

Job ;-;0.-0.

.1::..:.8

Pheasant Roadside Cover Evaluation ~t_u_d~y

Period Covered:
Personnel:

PROGRESS m:PORT

W-37-R-27

No.

Job Title

April 1974

.

_

_

April 1, 1973 to March 31, 1974

Ted Behlke, John Klein, Mike Loftsgard and Warren Snyder.

ABSTRACT
Pheasant nesting utilization of seeded roadside plots dropped from 0.94 nests
per acre in 1972 to 0.54 nests per acre in 1973. Unfarmed nat.ural, roadsides
retained slightly higher nesting use than in previous years. Farmed controls
received decreased use in comparisons with 1972. Decreased nest occurrence
was possibly attributed to (1) heavy winter snow pack which flattened all
residual cover, (2) decreased hen pheasant density, or (3) late spring stubble
plowing which prompted hens to move into green wheat to renest. Approximately
21 percent of the-nests hatched successfully in 1973, whereas about 58 percent
were predator destroyed. These figures are similar to those obtained in
previous 'years. During the four years'of study, one successful nest was
found per 2.42 miles of seeded roadside, per 3.54 miles of unfarmed roadside,
and per 12.23 miles of farmed roadside. Currently it is estimated that
unfarmed and farmed roadside yields only five to six percent of the total
pheasant production in the tablelands of northeast Colorado. Unfarmed roadside occupies 38 percent and farmed roadside occupies 62 percent of the total
roadside -in the region. If all roadsides were seeded, total pheasant production Would be increased by about eleven percent. There was no evidence
that hens selected roadsides for nesting in relation to adjacent field crop
types. Vegetative measurements in May, 1973 were markedly shorter and less
dense than those taken in previous years, however, by late June roadside
vegetation had made rapid growth and achieved normal height and density.

��-3-

PHEASANT ROADSIDE COVER EVALUATION STUDY
Warren D. Snyder
P. S. OBJECTIVE

To compare pheasant production under natural roadside conditions with the
following cover types to be established along roadsides: (1) grass, and (2)
grass-legume mixtures.

SEGMENT OBJECTIVES
1.

To measure pheasant production in test and control plots.

2.

To measure life-form characteristics of roadside cover.

METHODS AND MATERIALS
Reference is made to Snyder (1969, 1970, 1971 and 1972) concerning methods
and materials used in this study. A minor supplement to these is added as
follows:
In addition to height and density measurements of vegetation, ocular estimates
of litter were obtained during the first (May) sample period. One sample of
litter density was obtained each time a vegetation measurement was obtained.
Estimates were calibrated on a scale of one to three for sparse to dense
litter cover, repectively.

RESULTS AND DISCUSSION
Measurement of Pheasant Production
Factors Influencing 1973 Roadside Pheasant Production
Pheasant use of roadsides for nesting declined markedly in 1973 compared to
the three previous years of study. Tom Lines, Wildlife Conservation Officer,
(pers. comm.) reported that spring 1973 crowing indices did not provide evidence of significant pheasant population decline following prolonged cold
and snow conditions in November and December 1972. However, persistent
heavy snows did flatten nearly all residual vegetation in the roadsides so
that most offered marginal to poor nesting habitat in May.

�-4-

Above normal precipitation in March and April 1973 slowed vegetative growth
and also prevented plowing of wheat stubble during the usual April to early
May period. Instead, plowing was not initiated until mid-May and progressed
into early June on many farms before completion. If pheasants started
nesting in late April or early Mayas usual, then many nests undoubtedly were
destroyed in the vast stubble fields. It is probable that some hens were also
killed on the nests in instances where incubation was underway.
Warming temperatures in May resulted in rapid wheat growth which advanced far
ahead of roadside vegetation. Potentially, then, hens moved directly from
plowed stubble fields to green wheat to renest.
Pheasant brood count indices, numerous hunting season contacts, Fleming Check
Station data, and reports following the season indicate a greatly reduced
fall population level existed in 1973 compared to 1972. Therefore, the
reduced nesting density observed during nest searches apparently was indicative of generally reduced production. Pheasants may not have bypassed
roadsides for nesting in 1973. Instead it is possible there were fewer hens
surviving either following spring plowing or early winter snows.
Nesting Rate of Pheasants per Cover Type in Roadsides
In 1972, 58 nests were found in approximately 86 acres of roadside. In 1973
only 38 nests were found in approximately 80 acres of roadside. This represents a nest density reduction of about 30 percent from one year to the next
which corresponds to a 30 percent reduction in birds per mile observed in
August brood counts. In seeded roadsides densities dropped from 0.94 nests
per acre in 1972 to 0.54 in 1973 (Table 1). In farmed roadsides they declined
from 0.44 nests per acre to 0.19 nests per acre. Natural unfarmed roadsides
however, showed an increase in use from 0.42 nests per acre in 1972 to 0.65
in 1973. This latter figure exceeded nest densities found in seeded roadsides
for the first time during the four year period of study. Nests per acre
figures for successive years, 1970 through 1973 are presented in Table 1.
As observed in Table 1, nest densities in seeded plots increased slightly each
year until 1973 when they dropped markedly. Unfarmed roadsides were more
stable in use at a lower level. Nest densities in farmed roadsides were quite
low each year except in 1972 when nesting use approximated that in unfarmed
controls.
In 1973 seeded roadsides contained significantly more nesting attempts than
farmed controls (Chi-square = 4.05* 1 d. f.), but there was no significant
difference between seeded and unfarmed roadside utilization (Chi-square =
1.63, 1 d. f.). Unfarmed plots contained significantly more nests per acre
than farmed controls (Chi-square = 4.93*, 1 d. f.).
Pooled four-year nesting data, presented in Table 2 are sub-tended by chisquare values of comparisons among cover types. Seeded plots contained 0.80
nests per acre which was greater with high significance than either of the
controls. Unfarmed controls averaged 0.52 nests per acre which was significantly greater than densities of 0.23 nests per acre found in farmed
controls.

�-5-

Table 1. Nesting attempts and successful nests found along roadsides
during successive years of the four year evaluation. !/

Cover Type

Number
of Plots

Total
Acres

Nesting
Attempts

Nests/
Acre

Success.
Nests

Success. Nests
per Acre

0.15

1970
Seeded

48

26.63

22

0.83

4

Un farmed

23

11.67

5

0.43

0

Farmed

28

15.54

2

0.13

0

1971
Seeded

67

35.94

32

0.89

7

0.19

Unfarmed

38

19.88

11

0.55

5

0.25

Farmed

38

20.38

2

0.10

0

1972
Seeded

76

41.52

39

0.94

9

0.22

Unfarmed

42

21.30

9

0.42

4

0.19

Farmed

42

22.94

10

0.44

3

0.13

1973
Seeded

71

38.76

21

0.54

7

0.18

Unfarmed

39

20.12

13

0.65

1

0.05

Farmed

39

21.29

4

0.19

0

1/
-For

four year average see Table 2.

�-6-

Table 2. Nesting attempts and successful nests found along roadsides
during four years of evaluation, 1970-1973.
Number
of Plots

Total
Acres

Nesting
Attempts

Nests/
Acre

Success.
Nests

Success. Nests
per Acre

Seeded

262

142.85

114

0.80

27

0.19

Unfarmed
Control

142

72.96

38

0.52

10

0.14

Farmed
Control

147

80.16

18

0.23

3

0.04

295.92

170

Cover Type

Total

40

Chi-square Values

Comparison

Total Nesting
Attempts

Successful
Nesting Attempts

Seeded vs. Unfarmed Control

12.80** !/

3.06

Seeded vs. Farmed Control

29.50**

8.72*

Unfarmed vs. Farmed Control

17.43**

5.66*

1/

- One asterisk denotes significance, two denote high significance with 1 degree
of freedom.

Production Rate per Cover Type in Roadsides
More nesting attempts per acre were found in seeded plots than in natural
unfarmed controls, but nesting success was apparently lower in seeded plots
resulting in no significant difference in production between the two groups
(Table 2). However, both seeded and unfarmed controls contained significantly higher successful nest densities than were found in farmed roadsides.
Over the four-year period less than one-fourth of the nests hatched successfully so that the production rate per acre was very low (Tables 1 and 2). In
seeded plots, one successful nest was found per 2.42 miles of roadside (0.19
nests per acre). In natural roadsides one successful nest was recorded per

�-7-

3.54 miles (0.14 nests per acre), and in farmed controls one hatched nest
per 12.23 miles was observed (0.04 nests per acre). Both seeded and natural
roadsides produced significantly more young than farmed roadsides, but there
was no significant difference between seeded and natural plots (Table 2).
The low production rate in roadsides reflects a fairly high rate of predation
(Table 3) by a variety of predators. However, this rate of nesting success
is proximal to that found along roadsides in other states as illustrated
in Table 4. The narrow strips of cover lend themselves well to predators,
especially skunks, which have been observed in early mornings conducting their
own nest searches along roadsides. Apparently, few nests escape predation in
early spring when cover is sparse and little other cover exists elsewhere.
Higher nesting success was observed in mid-summer along roadsides after cover
on and off the plots developed, rodent populations increased, and predation
became less a factor.

Table 3. Fate of pheasant nests found in roadside study plots during the
sununers of 1970, 1971, 1972 and 1973.
1970

1971

1972

1973

Total
No.

Nest Fate

No.

%

No.

%

No.

%

No.

%

Successful

4

13.8

12

26.7

16

27.6

8

21.1

40

23.5

Predator
Destroyed

16

55.2

22

48.9

33

56.9

22

57.9

93

54.7

Abandoned

4

13.8

4

8.9

5

8.6

6

15.8

19

11.2

Unknown

5

17.2

7

15.5

4

6.9

2

5.2

18

10.6

%

Pheasant Production From Roadsides Projected on a Square Mile Basis
Current Pheasant Production from Roadsides in Northeast Colorado:--Accurate pheasant population information is not available for northeast Colorado.
Therefore, rough calculations based on (1) harvest-juvenile mortality data,
and (2) spring density data, are presented in Appendix A. These calculations
provide estimates of 88 to over 100 young hatched per square mile in the
tablelands of Phillips and Sedgwick counties. Or there were at least eleven
successful nests per section since the average successful nest in roadsides
produced about 8 young.

�-8-

Table 4. Percentage of pheasant nests terminated successfully in various
study sites.
State

Reference

Roadsides

Small Grain

All Covers

South Dakota

Trautman (1965)

22.2

43.7

23.9

Nebraska

Linder et al. (1960)

19.3

24.8

15.1

Pennsylvania

Hartman (1971)

16.7 !!

33.3

Iowa

Klonglan (1955)

29.0

18.8

17.3

Colorado

Lyon (1950)

34.4 !!

48.4

24.0

Illinois

Joselyn (1970)

26-29

Colorado

This Study

23.5

1/ Permanent cover including roadsides.
A systematic sample of 528 miles of roadside in 22 townships shows that 38
percent of the roadsides were un farmed , whereas about 62 percent were farmed
to the road shoulder in 1973. Calculations presented in Table 5 show that
only 0.63 successful nests could be expected from natural (unfarmed) and
farmed roadsides per section per year. If eleven nests per section is near
accurate, then only five to six percent of the current production comes from
roadsides. In terms of young, five would be hatched.
Based on calculations in Appendix A, only 0.75 cocks per section are contributed to the hunter's bag per section from roadsides. Hunter harvest
surveys indicate about 19 cocks per section are bagged in the region of the
roadside study. In other words, roadsides supply a very small percent of
the total harvest in northeast Colorado at present.
Production Assessment if all Roadsides Were Farmed
If all roadsides per section were farmed, and in some localities they are,
then we can expect only 0.33 successful nest per section (Table 5). On the
average, 2.4 chicks would be produced, yielding about 0.36 cocks per section
to the hunter's bag.
It should be remembered that the average farmed roadside is approximately
four feet wide and contains 0.56 acres of weeds or grassy cover per linear
mile.

�-9-

Table 5~ Pheasant production in roadsides in northeast Colorado based on
projections from study results.

Roadside
Status
,

Percent

·PerSeatiOnStattis
Linear
Total
Miles
Acres 1/

Successful
Nests

Number
Young ?:/

Harvested
Cocks 3/

Yield Under Existing Northeast Colorado Conditions
Farmed

62

2.48

5.42 1/

0.20

Unfarmed

38

1.52

3.14

0.43

Total

100

4.00

8.56

0.63

5.0

0.75

2.4

0.36

9.0

1.35

13.2

2.00

Yield in Farmed Roadsides Eer Section
Farmed

100

4.00

8.74 1/

0.33

Yield in Unfarmed Roadsides Eer Section
Unfarmed

100

4.00

8.26

1.13

Yield in Seeded Roadsides Eer Section
Seeded

100

4.00

8.74

1.65

1/
- Farmed roadside includes farmed and road shoulder acreage equivalent to that
of seeded plots used in study. The road shoulder portion would total 2.24
acres per section.

2/
- Based on 8 young per successful nest as observed during study nest searches.

3/
- Based on 40 percent natural mortality to harvest and 50 percent cock harvest
as illustrated in Appendix A.

�-10-

Production Assessment if all Roadsides Were Unfarmed:--If all roadsides
per section were unfarmed, then production would increase to about 1.13
successful nests, or 9 young per section. Based on 40 percent mortality to
hunting season and 50 percent cock harvest, 1.35 roosters per square mile
would be added to the hunter's bag from roadsides. Data in Table 5 show
that conversion of roadside from unfarmed natural state to farmed status
reduces cock harvest by about a bird per section per year. Unfarmed roadsides produce about four times as many young per year as do farmed roadsides.
Production Assessment if all Roadsides Were Seeded:--Seeding of all
roadsides per section would result in 1.65 successful nests per year or
production of 13.2 young. Two cocks per section would be added to the hunter's
bag per year (Table 5). An approximate 11 percent increase in production
would result from seeding 1.36 percent of the total land area. Seeded roadsides are more than five times more productive than farmed roadsides, but as
previously stated, production in seeded roadsides was not significantly
greater than in unfarmed roadsides (Table 2). Additional cover provided by
seeded or natural roadsides for brood rearing and winter cover are other
assets to pheasants for which there is no ready measurement.

Pheasant Production in Small Grains Compared with that in Roadsides
Cover mapping in the 1960's provided information showing that 39.6 percent
of 642,000 acres in northeast Colorado was in small grains, primarily green
wheat (Snyder 1970). The average section contained about 250 acres of wheat
and a slightly greater amount of summer fallow to be planted in fall. Corn,
sorghums, millets, beets, and beans were not considered as nesting cover.
Pastures, which comprised about 10 percent of the land area, were sodded
shortgrass for the most part, and of little value to pheasants. Linder et
ale (1960) found that pastures in southcentral Nebraska only occasionally
produced pheasants. Pasture cover is less abundant in the tablelands of
northeast Colorado.
Alfalfa occupied less than two percent of the land, but probably contributed
a small fraction of the total pheasant production. Most studies show
alfalfa as highly attractive for nesting, but due to haying, pheasant production is almost nil.
Vacant farmlots, draws and low areas occupied about one percent of the land
and undoubtedly provided fair to good nesting cover in many instances.
Assuming nesting success on these was the same as on unfarmed roadsides, they
contributed 0.88 successful nests per section.
Based on these data, small grains contribute most of the successful nests in
the tablelands of northeast Colorado. Assuming 9 nests per 250 acres, the
nest density is low, one nest per 28 acres. However, the vast acreage, in
comparison with roadsides, and the lower probability of predation apparently
establishes small grains as the predominant producer of pheasants in eastern
Colorado. For the most part, there is little other cover available for their
use in nesting. Meanwhile, the transition to less roadside and more irrigated row crops continues. It is fortunate for the pheasant, that much of
eastern Colorado will remain dryland and in wheat, due to lack of irrigation
water.

�-11-

Reference is made to Table 6 which illustrates that small grains usually
have low nest densities compared with roadsides.
But most studies show
lower predation rates in small grains and much higher percentage nesting
success.

Table 6. Comparisons
small grain fields.

of pheasant

(Harvard)

and

Reference

Pheasants
per Square
Mile

Pheasant
Roadsides

Linder et al.

29

1.67

0.15

37

1.91

0.20

147

1. 87-2.82

0.14-0.57

Location

Nebraska,

density and nest density of roadsides

Nests per Acre
Small Grain

(1960)

Nebraska,
Combined

Data

Baxter &amp; Wolfe
(1973)

South Dakota

Trautman

(1960)

Illinois

Joselyn

(1970)

Pennsylvania
Colorado,

Hartmann

(1971)

This Study

of Nesting

40-120

to Adjacent

0.10
0.04

25-30

Nest Association
Relationship

2.6 Seeded
1.2 Unmanaged

0.8 Seeded
0.5 Un farmed
0.2 Farmed

0.03

with Cover Trees
Field Cover Type
t

. ... -

_

There was no evidence during the four years of st:1Jdythat pheasants selecteo
nests in any of the roadsides in relation to adjac;.entfield typeel. Ind1v:t,
..•
dual comparisions for seeded, fa~d,
unfarmed and combined data, tot31e4
for the four years, revealed that ohset;'vedoccurrences app:ro~:t.matedtho
expected.
In no instance did Chi-square values app:r;oachlevels of s:lgn:U1~
cance in any 0 f the comparisons.
Wheat stubble J p:resent ,in sPl"inS,wa@ soo.n
plowed under, leAVing fallow througll the SUll111ler.Determinations CQW,d. not be
made as to whether most nests were established while adjacent stybble existed
or after it was plowed.

�-12-

Table 7. Utilization of vegetation within roadside cover types by nesting
pheasants during the summers of 1970 through 1973.
Number of Nests
Plot and Cover Type

Combined
Total

1970

1971

1972

1973

4
1
1
0
0
1
0
2
0
1
0
11
1
0

9
1
1
3
4
6
2
1
2
0
2
1
0
0

8
6
0
6
1
4
2
1
1
1
2
0
3
4

8~
6
4
0
1
0
1
0
0
0
0
0
1
0

22

32

39

21

114

0
0
0
1
2
2
0
0
0

r
3
2
0
1
2
2
0
0

1
1
2
1
1
2
1
1
0

0
0
3
2
1
3
0
3
1

1
4
7
4

5

11

9

13

38

Native Grass
D. Brome and Residual
Green Weeds and Residual
Straight Downy Brome
Green Weeds
D. Brome and Green Weeds
Green Weeds, Residual and
D. Brome

0
0
2
0
0
0

0

1
0
0
0
2
1

2
2

0
0
0
0

1
0
1
3
2
2

0

0

1

0

1

Sub-total

2

2

10

4

18

Seeded Roadside
Seeded Grass
Grass-Alfalfa
Seeded Alfalfa
Grass - D. Brome
Alfalfa - D. Brome
Straight Downy Brome
Grass and Residual Weeds
Grass, Alfalfa and Weeds
Grass, Alfalfa and D. Brome
Alfalfa, Weeds and D. Brome
Green Weeds
Weeds and Residual
Weeds, Residual and D. Brome
Grass, G. Weeds and Residual
Sub-total

29
14
6
9
6
11

5
4
3
2
4
12

5
4

Unfarmed Roadside (C2)
Native Grass
N. Grass and Downy Brome
Straight Downy Brome
Downy Brome and Residual
Green Weeds
G. Weeds and Residual
Straight Residual Cover
D. Brome, Weeds and Residual
N. Grass and G. Weeds
Sub-total

5
9
3
4
1

Farmed Roadside (Cl)
2

3
3
4

3

�-13-

Proximal Cover Association
In seeded roadsides, proportionally increased use of seeded grass and alfalfa
for immediate nesting cover was noted over previous years of study (Table 7).
Downy brome received high use in the unfarmed, natural roadsides. The
combined four year total shown in the table indicates that a wide variety of
covers and cover combinations were used by pheasants. Direct comparisons of
use and vegetative type occurrence cannot be made to show preference.

Life Form Characteristics of Roadside Cover
Vegetative Measurements
Measurements of height and density were taken in early May and again in
late June, 1973 as in prevtous years. Twelve plots, each, of seeded,
unfarmed and farmed roadside were sampled. In the farmed sample, only plots
containing green wheat were measured. In May, both height and density
measurements were markedly lower than in previous years (Table 8). This
was primarily due to lodging of residual cover under heavy snow during the
previous winter followed by a cold spring which delayed new growth. By
mid-June, warming temperatures and ample moisture had stimulated growth so
that evidence of reduced cover no longer existed, as shown in 'Table 8.

Table 8. Comparisons of height and density for 1971, 1972 and 1973 on
roadside study plots. );/

Plot Type

1971

May Measurement
1972
1973

1971

June Measurement
1972
1973

Height
Seeded

8.71

l3.l8

6.37

16.31

19.00

21.28

Unfarmed

6.08

8.07

4.72

13.06

15.10

15.05

Farmed

14.18

17.33

6.18

26.72

25.53

26.56

Density
Seeded

1.65

2.14

1.14

2.23

2.51

2.36

Unfarmed

1.35

1.71

1.l3

2.01

2.50

2.13

Farmed

1.66

1.95

1.17

2.19

2.43

1.84

�-14-

Ground Cover Measurement
Residual ground cover is considered an important factor in pheasant nest
establishment. The bowls of pheasant nests are composed of litter and it
also serves in breaking up the outline of the nest and the hen. Usually,
the hen's natural color closely approximates the color of dead vegetation.
Therefore, in May 1973 a sample of litter density was obtained. Based on a
sparse to dense litter index of 1 to 3, seeded roadside yielded an index of
2.36. Farmed roadside had a much lower index of 1.84 compared to a medium
index of 2.13 for unfarmed natural roadside. Although. the actual importance
of this variable in nest selection is unknown, seeded and unfarmed roadside
offered much better litter cover for pheasant use. Green wheat planted in
roadsides offers almost no residual litter for use in nesting.

LITERATURE CITED
Baxter, W. L., and C. W. Wolfe. 1973. Life history and ecology of the ringnecked pheasant in Nebraska. Nebr. Game and Parks Comm. Tech. Pub.
83 p.
Hartmann, F. E., and D. E. Sheffer. 1971. Population dynamics and hunter
harvest of ring-necked pheasant populations in Pennsylvania's primary
range. Northeast Fish and Wildl. Conf. 28. 39 p.
Joselyn, G. B. 1970. Management of roadside cover for nesting pheasants.
Job Compl. Rpt. Ill. Proj. 10-66-R-8, Job No. 3. 41 p.
Klonglan, E. D.
Ia. , 1954.

1955. Pheasant nesting and production in Winnebago County,
Proc. Ia. Acad. Sci. 62:626-637.

Linder, R. L., D. L. Lyon, and C. P. Agee. 1960. An analysis of pheasant
nesting in southcentral Nebr. Trans. N. Am. Wildl. Conf. 25:214-230.
Lyon, L. J. 1950. Pheasant nesting success on agricultural land in Colo.
Colo. Coop. Wildl. Res. Unit. Quart. Rpt. 4(1):1-19.
Snyder, W. D. 1970. Pheasant hen harvest investigation - final rpt.
Div. of Game, Fish and Parks. Game Res. Rpt. April. p. 3-81.

Colo.

Trautman, C. G. 1960. Evaluation of pheasant nesting habitat in eastern
South Dakota. Trans. N. Am. Wild1. Conf. 25:202-213.
1965. Evaluation of pheasant nesting habitat in eastern South
Dakota. S. Dak. Dept. of Game, Fish and Parks. F. A. Comp1. Rept.
W-75-R.

Prepared by

1fa~s~
Wildlife Researcher

�-15-

APPENDIX A
Part 1. Projected pheasant production per square mile in Phillips County
based on hunter harvest survey data.

Variable Per Square Mile
Harvest on 600 square miles averaged over a
14 year period (1955-1968)

18.9 cocks

Seventy percent of cocks are young (1966-1970)

13.2 young cocks

Assuming a 50 percent harvest, then 26.4 young
cocks were present at start of hunting season
or the total young approximated

52.8 young birds

Numerous references show an average 35 percent
mortality from hatch to 6-10 weeks of age.
Assuming an additional 5 percent mortality to
hunting season, then 52.8 young equals 60
percent of the original hatch or

88 young hatched

Nest searches show mean clutch size approximated
8 eggs per nest, or successful nests numbered

11 successful nests

Part 2. Projected pheasant production per square mile in Phillips County
based on breeding population density.

Variable Per Square Mile
Mean pheasant crowing index equaled 49.75 calls
per station (1963-1968) or there were about
(Assumes an approximate 1 mile hearing distance)

12.4 cocks

Hens per cock equaled 2.34 (1963-1968), or

29.0 hens

Wide variation in percentage nesting success
is cited in the literature, so a conservative
50 percent success rate is used here. Assuming
50 percent of hens nest successfully, then there
were

14.5 nests

14.5 nests times 8.eggs per nest equals

116.0 young birds

��April 1974
-17-

JOB PROGRESS REPORT

.State of

~C:::O::.;:LO::::..::RAD::.::::..::.O
_

Project No.

W-37-R-27

Work Plan No.

3

Job Title

Game Bird Survey
Job No.

B_a

_

Effects of Sagebrush Control,on Distribution and Abundance of Sage Grouse

Period Covered:

April 1, 1973 to March 31, 1974

Personnel: Fred Giese and Robert Streeter, U. S. Fish and Wildlife Service;
·Thomas Beck, Clait Braun, Jack Corey, Courtney Crawford, John Ellenberger,
Heather Flanagan, Howard Funk, Jack Gustafson, Richard Hoffman, Russell
Kozacek, Ronald Oakleaf,Steven Porter, Wayne Russell, Michael Szymczak,
John Wagner, Richard Wenger, Colorado Division of Wildlife.
ABSTRACT
In all, 321 sage grouse (Centrocercus urophasianus) were banded during this
segment, 134 on strutting grounds, 154 on nearby areas of concentration, .and
33 on wintering areas. Eight active strutting grounds were located within or
adjacent to the study area, three of which were new or relocated grounds. The
peak male count was 257 cocks. Two strutting grounds accounted for 54.5 percent (140) of the cocks observed. Twelve nests were located in 280 acres of
nest search plots, none in the block sprayed areas. Of the 71 adult females
observed during the summer of 1973, 32.4 percent (23) had broods. Mean brood
size was 4.05 chicks per hen and the young per adult female ratio was 1.15:1.
No significant differences existed between relative frequencies of grasses,
forbs, and shrubs in sprayedvs. unsprayed plots. Sagebrush densities by
age class in sprayed and control blocks indicate the range is in a rejuvenative stage. Thirty-one percent (127) of all sage grouse sighted during the
summer of 1973 were in treated areas. Three hundred sixty-six hunters checked
193 sage grouse through check stations on the opening weekend of a three day
season. W.inter distribution was plotted on the basis of 130 flock sightings,
amounting to 2,755 birds. Six high use areas comprisingless·than
27 square
miles of the sagebrush lands in North Pal:k were evident. Strong segregation
by sexes is evident in sage grouse flocks. Of the 118 flocks classified, 74
percent (87) were composed of at least 90 percent birds of like sex , Mean
size of male flocks was 11 birds; that of female flocks 32 birds. Flock size
was less than 50 birds in 92 percent of all observations. No differences
could be detected in vegetal characteristics of winter sites used by male and
female flocks. Sixty-four percent (N = 87) of the winter use sites had slopes
less than 5 percent while only 2 percent of the areas had slopes greater than
15 percent. Seventy-one percent (N = 86) of the winter flock locations were
on south, southwest, or west facing slopes. Winter sightings of 71 banded
birds provided mean distance traveled for males of 6.7 miles (N = 51) and for
females of 11.9 miles (N = 5). Sex ratio of 2,356,birds observed during the
winter of 1973-74 was 61 males:lOO females.
.

�-18RECOMMENDATIONS

1.

Nest searches

on permanent

2.

Nest searches should be conducted in suitable habitat in areas not
included in the original study area with the objective of locating
areas of high nesting density.

3.

Vegetation

4.

Brood counts should be made throughout North Park with emphasis
locating females banded in the Lake John Area.

5.

All areas of sagebrush
acreage determined.

transects

5 acre plots should be. discontinued.

should be discontinued.

disturbance

on

in North Park should be mapped and

�~19-

EFFECTS OF SAGEBRUSH CONTROL ON
DISTRIBUTION AND ABUNDANCE OF SAGE GROUSE
Thomas D. I. Beck
Sage grouse, the largest North American member of the Tetraonidae, inhabit
areas dominated by sagebrush (Artemisia spp.) and sub-dominant grass types
with interspersed wild hay meadows. As a result of agricultural demands,
overgrazing, and encroachment of man-made developments, there has been a
continued and rapid destruction of sage grouse habitat. In the face of
such changing land use patterns sage grouse have continued to be highly
specialized in their habitat requirements. They have not adjusted and it is
doubtful if they can adjust their life processes to fit a land use pattern
which eliminates or disturbs large tracts of lands dominated by sagebrush.
Compounding this habitat specialization problem sage grouse are one of
relatively few animals which depend upon a climax vegetation stage. Climax
vegetation types are hazardous food sources due to their slow recovery after
natural or man-caused catastrophies.
Due to pressures to alter sagebrush r?Ugelands, the Colorado Division of
Game, Fish and Parks (now Colorado Division of Wildlife) in cooperation with
the Bureau of Land Management, initiated a study in North Park, Colorado to
document the effects of 2,4-D spraying of sagebrush on sage grouse abundance
and distribution. At the time of the study's initiation in 1963, 2,4-D
spraying to reduce or eliminate sagebrush was a popular tool of land management agencies.
Pre-treatment studies were accomplished in 1963-65 with study patterns for
spraying designated and actual spraying accomplished in June, 1965 (Gill
1965). nvO years of study on the immediate effects of spraying sagebrush
followed, being completed in 1967 (Carr 1967). During the summers of 1968
and 1969, studies were conducted on distribtuion of females in relation to
the sprayed areas, by use of radio telemetry (Poley 1969, May 1970).
During this research segment, the third and final segment of the North Park
research project was initiated. Emphasis during this study will be directed
toward the winter period on the assumption that if sagebrush eradication
programs did affect North Park sage grouse populations, any such changes
should be manifested during a time of limited habitat - winter.
The question investigated concerned winter distribution of sage grouse and
characteristics of winter use areas. In addition, most of the parameters
investigated by earlier researchers are being monitored to provide longterm comparisons.
P. S. OBJECTIVE
To determine the effects of 2,4-D spraying of sagebrush on (1) sage grouse
abundance, (2) sage grouse distribution, and (3) vegetative composition and
density.

�-20-

SEGMENT OBJECTIVES
1.

To investigate seasonal numbers of sage grouse present by age and sex
throughout the year.

2.

To investigate the seasonal distribution of sage grouse throughout the
year.

3.

To investigate the current correlation between numbers and distribution
of sage grouse in relation to vegetative and habitat composition, patterns
and characteristics, and initiate comparisons of current results with
those obtained in previous segments.

4.

To prepare the required annual job progress report.

In addition, attempts were made to (1) characterize winter flocks by sex
composition and size of flock, (2) detect shifts in distribution during the
winter, and (3) locate banded birds and calculate movements.

REVIEW OF LITERATURE
Intensive studies of sage grouse throughout the west have provided extensive
data on most phases of the life history of this bird. MOst research has
been devoted to the requirements for successful population maintenance or
to behavioral studies.
Food habits studies have been conducted in most states, with the seasonal
food requirements of sage grouse being well documented. Sagebrush is the
major food item throughout the year except for the summer months (Griner 1939,
Wallestad et al. 1973). Throughout the summer months, succulent forbs were
the main food item for adults (Patterson 1952, Martin 1970, Wallestad et al.
1973), with juvenile birds utilizing succulent forbs and insects (Klebenow
and Gray 1968, Peterson 1970). The winter diet of sage grouse is almost
100 percent sagebrush (Bean 1941, Patterson 1952, Dalke et al. 1963). All
food studies have clearly shown that sage grouse are dependent on sagebrush
for food.
Most researchers have characterized strutting sites as being an open area
surrounded by sagebrush (Scott 1942, Patterson 1952, Dalke et al. 1963,
Gill 1965). Dalke et al. (1963) suggested that a possible management tool
is the clearing of ~ - ~ acre areas in dense stands of sagebrush to provide
areas for males to strut and open up adjacent lands for nesting. This
reference to nesting implies that nesting distribution is dependent upon
strutting ground locations. Gill (1965), Martin (1970) and Wallestad and
Pyrah (1973) reported over 80 percent of the sage grouse nests located in
their study areas were within two miles of a strutting ground. However,
Autenrieth (1970), studying a migratory population on the Snake River plain
in Idaho reported that a four mile radius was needed to include 80 percent
of the nests located. In studying two sedentary populations, Autenrieth
(1969) found that a five mile radius was needed to include 80 percent of all
nests located in the Little Lost River drainage and a three mile radius

�-21-

was needed to include 80 percent of all nests found in the Shoshone Basin.
Canopy coverage of the three areas was similar (24, 30 and 29 percent,
respectively).
Distance from water appeared to have no relation to distances
of nests from strutting grounds.
Nearly all nests located have been under sagebrush, although nests have been
found under various shrub species and in open areas (Patterson 1952, Gill
1965, Klebenow 1969, Autenrieth 1970).
Stands of 20 to 30 percent canopy
coverage are most often selected for nesting (Patterson 1952, Klebenow 1969,
Autenrieth 1970), and thus, areas where sagebrush has been eradicated provide
no nesting habitat (Klebenow 1970).
Brood rearing habitat studies have shown that meadows provide needed forbs
and insects for chicks in most areas (Griner 1939, Gill 1965, Autenrieth 1970,
Oakleaf 1971, Wal1estad 1971). Klebenow (1969) detected altitudinal movements of sage grouse broods as they moved to higher elevations when succulent
forbs on lower ranges dried out. Autenrieth (1969) points out that in areas
of high precipitation, sagebrush eradication programs can benefit grouse
populations by opening up dense stands of sagebrush and allowing forbs to
become established.
He has documented such an occurrence on Edie Creek
Bench in southern Idaho. The annual precipitation in this area is about 18
inches.
Klebenow (1970) also found that some sprayed ranges were providing
excellent brood habitat but only after several years had passed since
spraying.
During the interim years the sprayed areas received little sage
grouse use.
Little scientific effort has been directed toward the winter ecology of sage
grouse.
Numerous
references have referred to winter distribution of sage
grouse, but little or no quantitative data has been presented.
In areas of
greatly varying elevations, winter distribution and movements of sage grouse
are dependent upon snow depth, with birds moving to lower elevations as snow
covers the sagebrush on higher ranges (Griner 1939, Patterson 1952, Dalke
et a1. 1963).
Eng and Sch1adweiler (1972), studying five radio telemetered
hens, found little movement during the winter, with birds primarily using
areas where canopy coverage exceeded 20 percent.
Their study area in central
Montana does not have elevational variation as great as the areas studied
by Griner, Dalke and Patterson.
Eng and Schladweiler also reported that
wintering sage grouse preferred areas of low relief and avoided dense stands
of sagebrush on steep slopes.
No studies on sage grouse have been devoted to winter flock composition.
However, Patterson (1952) referred to partial segregation of sexes in winter
flocks in Wyoming, "There was no complete segregation of sexes or age groups
in any of the large concentration areas, although flocks composed predominately of males or females were commonly seen within the scope of the concentration areas."
He p.rovided no quantitative data indicating the degree of
segregation.
Patterson further states that "there was a pronounced tendency
for adult birds to segregate themselves according to sex on their wintering
grounds."
Patterson noted that the prevalent social unit was the flock, and
flock sizes generally varied between 10-100 birds, with occasional flocks of
100-200 birds.

�-22-

Autenrieth (1969), working in Owyhee County, Idaho, noted that "it is not
uncommon for adult cocks to winter in groups away from hens and juveniles."
Flock sizes for 56 flocks in Owyhee County varied between 2-110 during the
mild winter of 1969-70, whereas during the severe winter of 1968-69 several
groups of 500+ birds were seen. No distinction was made as to whether these
500+ bird sightings were one flock or numerous flocks in close proximity
(Autenrieth 1970).
Scientific references to all grouse species during the winter are limited.
Weeden (1964) documented sexual segregation of Alaskan'rock ptarmigan (Lagopus
mutus) during the winter. Segregation occurred by habitat types with males
utilizing wind blown areas near their breeding territories, while females
moved great distances to areas with abundant willow (Salix spp.) stands.
Braun and Schmidt (1971) reported similar results from research on whitetailed ptarmigan (Lagopus leucurus) in Colorado. Partial segregation of
sexes in winter occurred, with males utilizing high windswept areas and
females utilizing lower areas near tree line, where tall, dense willow stands
provided food. Snow quality was of extreme importance to white-tailed
pta:nnigan and bird~ Vi'e.~e.
~lQt;.e.(1
t;.Q
W.Q'le. ~
t;Q. one mile frorafeedmg areas in
search 'of snow suitable for roosting.
Seiskari (1962), in comparing winter habitats of capercaillie (Tetrao
urogallus) and black grouse (Lyrurus tetrix) in Finland, observed partial
segregation of sexes in wintering flocks of both species. Segregation was
stronger in capercaillie, a sp~cies in which females and males utilize
different winter habitats. Both sexes of black grouse utilized similar
habitat.
Koskimies (1957), in describing flocking behavior of capercaillie and black
grouse, postulated that capercaillie male flocks were probably composed of
males utilizing the same display area, owing to the sedentary nature of males.
Black grouse winter flocks show some sexual segregation, whereas capercaillie
flocks are distinctly unisexual (75 to 85 percent of one sex in a flock).
Strongest segregation occurred in flocks composed of adult males, suggesting
that female flocks may be composed of family aggregations, including subadult males.
Ellison (1973) found little segregation by sex of spruce grouse (Canachities
canadensis) in winter flocks. In general, females and juveniles of both
sexes were found in association, while most adult males were found with
females or as singles.
In most references, flocking is generally attributed to being a response to
unfavorable weather conditions. Koskimies (1957) presents evidence to suggest
that temperature is a factor in flocking intensity, with lower temperatures
resulting in more intense flocking. Braun and Schmidt (1971) noted that
birds began to appear on known wintering areas following intense snow storms.
Flocking is not as pronounced in capercaillie (Koskimies 1957), spruce grouse
(Ellison 1973), and ruffed grouse (Bonasa umbellus) (Gullion 1970) as in

�-23-

White-tailed ptarmigan (Hoffman 1973), rock ptarmigan (Weeden 1964) and black
grouse (Koskimies 1957), based on frequency of sightings of single birds,
nor is flock size as large in the capercaillie, spruce grouse, and ruffed
grouse as in the other grouse species studied. Flock sizes of capercaillie
and black grouse were greater in areas of large uniform habitat blocks
(Koskimies 1957).
Of all investigations on winter flocking of grouse mentioned in this review,
only Ellison (1973) used an arbitrary distance (18 meters) of association
between birds as a basis for determining flock size. All others based flock
size on the decision of the observer as to Which birds seen were in association.
METHODS AND MATERIALS
Capturing and Banding
Capturing and banding of sage grouse in the Lake John area was initiated on
26 March 1973 and continued until 1 May 1973 when capture activities were
switched to strutting grounds. Trapping continued on strutting grounds
until 30 May 1973. Capture operations were conducted sporadically throughout
the period 7 January 1974 to 13 March 1974. More intensive capture operations
were conducted after 13 March when birds were located on strutting grounds.
Birds were located by spotlighting and were then netted While on roosting
areas as described by Gill (1965). An adaptation to this procedure was
developed during the spring of 1973. Instead of using a motor vehicle
electrical system for the light source, an automobile battery (12 volt)
fastened to a pack frame was used. This allowed greater mobility for
personnel and enabled workers to search areas inaccessible to motor vehicles.
Best results were obtained when using an aircraft landing light (250 watt
bulb) for spotlighting, a long handled (3 meter) net for capturing, and a
tape recording of snowmobile noises to mask sounds of the approach. In 1973,
3.8 centimeter mesh cotton netting was used, with a hoop diameter of 100 cm.
In 1974, 1.25 centimeter mesh nylon netting was used in efforts to reduce
injuries to primary feathers. In addition, hoop diameter was reduced to 70
cm and lighter, aluminum tubing was used in net construction to provide
greater maneuverability. Captured birds were placed in burlap bags for
holding until they could be banded.
All captured birds were banded with aluminum, serially-numbered bands.
Males were banded with size 16 bands, and females with size 14 bands. Colored
plastic bandettes were placed on all but 55 of the birds. Any bird recaptured
during 1974 which did not have a plastic bandette was banded with the color
which was coded to the initial capture site. Each capture area was assigned
a specific color code. Birds caught on strutting grounds were color banded
on their left leg while birds caught in all other areas were color banded
on their right leg. Age, sex, weight, date, and location of capture were
recorded for each bird. Age and sex of all birds handled were determined by
techniques described by June (1967). In addition, blood smears for parasitological examination were taken from most birds captured during 1974.

�-24-

Strutting Ground Counts
Known strutting grounds in the area were censused regularly from 23 April
1973 until most strutting activity had ceased, near the middle of May.
During this period three new or relocated grounds were found and were censused
regularly thereafter. These grounds were located from the air during two
mornings of aerial search in early May. The primary study area was searched
by flying east-west transects at quarter mile intervals. All grounds censused
were approached in a slow moving vehicle and observers attempted to obtain
accurate counts of the numbers of birds present by sex class. Frequently,
observers had to approach grounds on foot due to deep snow or impassable
roads. All counts were made within one half hour of sunrise. - In 1974, birds
were first observed on strutting grounds on 13 March, 10-14 days earlier than
normal, probably due to the mild winter and early spring. Data collected
include high count of males and females, number of banded birds, time of
strutting, and adult male:sub-adult male ratios. Aging of males on strutting
grounds is based on the shape and length of rectrices (Lumsden 1968). This
method is somewhat questionable (Dalke et al. 1963) and all captured males
will be examined to learn if aging by rectrices corresponds to the accepted
technique of aging by appearance of the outer primaries.
Nest Search
Systematic nest searches were initiated on June 20 on five acre plots
originally established and marked by Carr (1967). The areas were searched
by two and three man teams on foot. A distance of ten feet was maintained
between the searchers. All nests of the year, old nests, and shell fragments
found were recorded. Plots are unevenly distributed throughout the large
block spray area, the small block spray area, the strip sprayed area, and two
control areas.
Brood Counts
Brood counts were conducted on a regular basis throughout July and the first
three weeks of August. Routes were usually run during the three hour period
immediately following sunrise and immediately preceding sunset. Records of
brood sightings were also recorded during mid-day. Routes along the river
bottoms were covered from horseback while routes through the sagebrush were
covered on foot or in a vehicle. Veh~cle speed was maintained between 5-10
mph. The intent of the brood counts WqS to achieve maximum number of sig~tings,
therefore no attempt was made to record mileage or always run the same routes.
All sightings of sage grouse were recorded, as Well as observations of predator;yanimals.

Vesetation Transects
Vegetation transects were run on plots established by G;L11(196.5). All
transects were run dUring early August after the grasses had headed out.
Measurements were taken to provide information on percent composition and
shrub density. A detailed description of the methods used was presented by
Gill (l,965) ,
, .'

�-25-

Summer Movements
Data on summer movements of sage grouse from s~rutting grounds and nearby
concentration areas were collected from observations of color marked birds
and from band recoveries obtained during the hunting season (early September).
All locations of the sightings or kills were plotted on a map (D.S.G.S. 7.5
minute) and the airline distance from the banding site was measured.
Distances reported are not necessarily the total distance traveled by the bird,
only the minimum distance traveled.

Summer Distribution

and Abundance

All sage grouse observed were counted, classified by sex when possible and
the location and vegetative type were recorded.
Special emphasis was placed
on categorizing as treated or untreated areas where grouse were observed.
This was frequently difficult as the strip sprayed areas had a poor initial
kill of sagebrush and boundaries were often difficult to discern.

Harvest

Data

Check stations were operated at Walden and Cowdrey on September 8 and 9,
the first two days of the three day season.
Grouse wings were collected
from hunters for aging and sexing, using the technique described by Crunden
(1963).
Bands were collected and hunters questioned as to the location
where banded birds were shot. This information was used to calculate age
and sex composition of the harvest, movements of banded birds, peak of hatch,
and fall production estimates.

Delineation

of Winter

Distribution

Distribution of wintering sage grouse in North Park was determined by
searching all sagebrush lands in the park with all probable areas used being
searched at least once. An area was considered a probable use area if any
sagebrush was visible above the snow. Areas inaccessible by four wheel drive
vehicle were searched from snowmobile or snowshoes.
Aerial search of North
Park along random transects was conducted on 21-22 February and 17 March.
Data concerning grouse flocks observed from the air were used in mapping
distribution but were disregarded when calculating flock sizes due to
inaccuracies involved in counting flying grouse.
Written records were made
on both grouse observations and indications of grouse use (droppings, tracks,
roosts).
Observations reported by reliable observers were followed up by
intensive
searches of reported locations within one day of the initial
sighting.
The winter distribution work covers the period 8 January to 8
March, 1974. March 8 was selected as the ending date as the Lake John study
area was approximately
90 percent snow free at this time.

Characterization

of Winter

Flocks

A flock was considered to be any group (2 or more) of birds in close·'p-roxdmf.t y
to each other.
The decision as to flock size was made by the observer as

�-26-

flocks often ranged over distances from 10-200 meters, thus inhibiting the
use of standard distances between birds as criterion for flock size. The
total number of birds in a flock was recorded, and when possible, flocks were
classified as to sex composition. For example~ a flock comprised of greater
than 50 percent females was referred to as a female flock, and vice versa.
Sightings of single birds were also recorded.
Sage grouse are sexually dimorphic, thus enabling one to sex birds visually
at a distance. Male sage grouse in breeding plumage are best characterized
by contrasting color zones of black and white from the base of the lower
mandible to the abdomen. The head is dark in appearance with a mixture of
gray and black feathers. The black and white speckled area at the base of
the lower mandible is separated from the black throat by a narrow white
collar. This collar runs from the eye down under the head and is visible
from the front and side. Lines of demarcation are distinct on the collar;
The black feathers of the throat are white tipped, but contrast sharply with
the solid white breast feathers. The feathers on the lower breast are white
with black tips. The black portion of these feathers becomes greater as
one progresses to the abdomen. The abdomen appears solid black. Males
have long (10-15 mm) filoplumes extending from the side of the neck which
are readily visible, especially on windy days. Males are much larger in
size, averaging twice as large and heavy as females. Adult sage grouse undergo a complete post-nuptial molt which is completed by late September and
juvenile birds acquire their first adult or winter plumage by late October.
Thus, all males were in breeding plumage during the study period (Patterson
1952, Bent 1932).
Female sage grouse are smaller and more drab colored than males. The abdomen
is black but not as solid as in males, being splotched with gray and white.
The breast is white, gray, and tan, giving a splotched appearance with no
discrete color patterns visible. There is no white neck collar nor any filoplumes on females. The area at the base of their lower mandible is more white
in females than it is in males. The back and sides of females are lighter
in color, having less black than males. Using these criteria, over 85 percent
(2,356 of 2,755) of all birds observed were classified as to sex.
Characteristics of Winter Use Sites
The location of each flock sighting was plotted and recorded to the nearest
quarter section. The following physical and vegetal characteristics of
each site were recorded: aspect, slope, snow depth and condition, sagebrush
density, average crown breadth, and height of sagebrush above snow. Weather
conditions such as temperature, cloud cover, precipitation, and wind direction
were also recorded for each sighting.
Winter Movements
Sightings of banded birds were recorded, as well as the number of definitely
unbanded birds by sex classes. MOvement data were obtained from observations
of birds banded with colored bandettes coded to banding sites. The location
of each sighting was plotted on a map and the airline distance from the
banding site measured. Distances reported are measures of minimum distances
travelled. Directions of movements were determined from banding location to
observation sites by plotting the two locations on a map.

�-27-

Aerial

Search

Two aerial censuses were made, one each in February and March.
These flights
were along 12 random transects running east-west across North Park. Two
observers were present on each flight and were responsible for counting all
grouse observed in a one eighth mile strip of ground on either side of the
plane.
This resulted in each transect being one quarter mile wide.
In this
way, approximately 9 percent of the park was covered.
The location and size
of each flock were recorded On cassette tapes. The purpose of these flights
was to examine the feasibility of random aerial census for estimating sage
grouse populations just prior to the breeding season.

Analysis

of Data from Previous

Years

Comparisons were made of weather data, strutting ground counts, brood counts
and harvest data. Various statistics were graphed in an attempt to find
visible correlations.
Sage grouse management in Colorado was established on
the premise that harvest data are a reflection of strutting ground counts
and production estimates.
These comparisons serve as a test for the accuracy
of the management plan.

DESCRIPTION

OF STUDY AREA

North Park (Jackson County) was chosen for study due to a proposed land
management plan by the Bureau of Land Management (BLM) which would result in
every acre of BLM controlled sagebrush lands in North Park being sprayed.
In addition, North Park supported the highest population of sage grouse in
Colorado (Rogers 1964:47).
Pre-treatment investigations outlined an intensive study area in the northwest portion of North Park. This area, in the Lake John vicinity, encompasses
approximately 60 square miles.
MOst of the land in the study area is under
the control of the BLM. The remainder of the intensive study area, primarily
meadowland along streams is under private control.
The area is bounded on
the north by Independence Mountain, on the east by the North Platte River,
on the south by the North Fork of the North Platte, and on the west by
Sheep Mountain and Boettcher Ridge. There are two major vegetation types in
the area: sagebrush-grassland,
and wild hay-meadowland.
The meadowland is
restricted to the water courses, with two major drainages in the area, the
North Platte River on the east side and the Lake John-Lake Creek drainage on
the west.
The sagebrush-grassland
type is found in the hilly terrain between
the two major stream systems.
A more detailed description is presented by
Gill (1965). Areas adjacent to the described study area are considered as
part of the study unit for the winter distribution work.
These areas lie to
the east and south of the formal study area and are necessarily included due
to suspected limited wintering habitat within the Lake John study area and
the mobility of sage grouse flocks.

�-28-

RESULTS AND DISCUSSION
Capturing and Banding
In all, 288 birds were banded during the spring of 1973, 134 on strutting
grounds and 154 on nearby areas of concentration. In addition, 33 birds
were caught during the period 8 January-31 March 1974. This brings the
total number of birds banded in this segment to 321, 230 more than the previous high number banded in any segment (91 in 1967). Data on sex, age
and specific locations of the spring caught birds are presented in Table 1.
Twenty-two birds were recaptured during the spring period and none during
the winter. One-half of the birds recaptured were caught within seven days
of initial capture and were in the same general area. Four birds were killed
during capture operations for a capture mortality rate of 1.2 percent.
During spring operations, 0.83 hours of field time were required to capture
and band one bird, whereas during the winter this figure rose to 3.8 hours
per bird. Winter trapping was not successful for two reasons. First,
birds were difficult to locate in areas accessible to four wheel drive
vehicles. Second, birds generally roosted in a small area which meant only
one or two birds could be netted before the flock flushed. Roosting in
restricted areas is contrary to the situation found during all other seasons
when birds are usually spread out over an area with seldom more than 3-5
birds in close proximity.
No lab work has been completed on the blood smears taken from birds caught
in 1974. Analysis of this portion of the work will be presented in the
final report.
Strutting Ground Counts
Eight active grounds were located within or immediately adjacent to the
primary study area. The total male peak for these eight grounds in 1973 was
257 cocks. Incomplete counts were made in 1972, so no comparison can be
made, but the 1973 count was less than the 1971 count of 326 cocks. The five
year average for the years preceding 1973 (excluding 1972) is 291 cocks. It
is possible that the difference may be due to natural fluctuations or human
error rather than a major population decline. Possibly, counts were affected
by weather conditions as the preceding winter (1972-73) was one of heavy snow
cover. Also, storms frequently occurred throughout April, 1973. Active
strutting was either delayed and/or not detected until late in the normal
strutting season. Two strutting grounds accounted for 54.5 percent (140)
of the cocks observed.
The peak periods for males and females on these strutting grounds, Alkali Lake
and Boettcher Junction, are plotted in Figs. 1 and 2, respectively. Table 2
gives the results of strutting ground counts on the eight active grounds for
the spring of 1973. The Monahan Draw strutting ground, which is in the large
block spray area, only had 11 cocks present this spring. This ground had
maintained high numbers of birds in the years since the spraying (64 in 1966,
74 in 1967, 53 in 1968, 80 in 1969, 57 in 1970, 81 in 1971), and the sudden
drop in 1972 (10), and 1973 (11) is at present unexplainable. Counts on the
grounds as of 31 March 1974 were approaching 50 percent of the peak male

�Age and sex classes of sage grouse banded at specific locations, spring 1973.

Table l.

Color Marking

Area

Unknown Age
F
M

Ad M

SA M

Ad F

SA F

Total

Y on B

RL

3

1

2

2

5

9

22

Y on R

RL

0

1

6

9

13

6

35

Jackson Co. 7 (2-5 mi S of L. John)

Ron

Y

RL

0

1

11

5

5

3

25

Jackson Co. 7 (0-2 mi S of L. John)

B on Y

RL

0

0

1

1

3

1

6

Jackson Co. 7 (N of L. John)

Y

RL

0

0

5

2

4

1

12

Jackson Co. 6 (E of Michigan River)

G

RL

0

0

2

0

0

0

2

Case Flats

R

RL

1

8

2

3

14

10

38

Jackson Co. 12

Y on G

RL

0

3

3

0

2

1

9

Jackson Co. 6 (W of Michigan River)

B

RL

0

0

1

0

0

0

1

E of Walden

G on B

RL

1

0

2

0

0

1

4

Boettcher Jct. S. G.

Y

LL

0

0

15

8

5

1

29

Alkali Lake S. G.

R on Y

LL

4

0

24

20

2

2

52

Walden S. G.

Y on B

LL

0

0

8

7

0

2

17

Monahan Draw S. G.

B on G

LL

0

0

2

0

0

0

2

Riley S. G.

Ron

LL

0

0

1

4

0

0

5

New Cowdrey S. G.

B

LL

0

0

2

7

0

0

9

Y on R

LL

0

0

3

12

1

4

20

9

14

90

80

54

41

288

Jackson County 33
Jackson Co. 7 (5-7 mi Sof

,

}

Wattenburg #2 S. G.

Total

L. John)

B

I

N
\0
I

�-30-

••
••
••
••
••
••
••
•••
••
••
••
.-

150

140
130
120
110
100
90
80

70
Q

Z

60

::)

o

50

~

C!)

z

o
30

10

DAY E (mo./day)

4/24 4/27 4/28 4/305/2

I

MALES

Fi 9 .1.

A 1k ~ ,Ii

L a k eSt

rut

tin

5/5

5/8 5/13 5/14

!

FEMALES

9

G r 0 u n d, 19 73.

5/17

�Q

Z
~

o

t.:

C)

z

I

o

W

•.....
I

l:1li:
UoI

&amp;G

I

no
grouse

~

z
DATE

4/25/13

Fig.a.

4/26/13

Boettcher

!!FEMALES

4/28/13

4/30/13

Lake

Jet.

Strutting

.MAL

ES

5/2/13

5/4/13

Ground.

5/1/13

5/16/13

�-34-

and unsprayed strips were indistinguishable. Therefore, in most of the strip
sprayed areas there was little change in gross habitat features and one
wouldn't suspect nesting incidence to be substantially altered.
The Lake John area appears to contain poor nesting habitat, as evidence by
low numbers of nests and broods located. However, the Walden Reservoir area
two miles east of the study area has possibly the best nesting habitat in
North Park. Two miles is not a great distance for hens to travel in search
of ~ nesting site (Autenrieth 1970). During the upcoming segment, extensive
efforts will be made to survey areas near Walden Reservoir and other apparently suitable nesting locations throughout North Park in search of females
banded in the Lake John area. This work will test the hypothesis that sage
grouse disperse throughout North Park during the summer as well as in winter
(see section on Winter Movements).
Brood Counts
Three hundred and sixty-seven sage grouse were seen in the study area from
11 June through 23 August 1973. Of these, 154 were adult males, 55 were
unclassified, with the remainder being hens and broods. A total of 71 adult females.were observed with 32.4 percent (23) of these having broods. Average brood
size was determined for hens where complete counts of broods were made. The
average brood size was 4.05 chicks per hen for 19 broods. The young per
adult female ratio was 1.15:1. No large decrease in brood size was evident
as the summer progressed nor were any hens seen with more than six chicks.
This is contrary to the general pattern of hens "adopting" chicks from
several broods in late summer as reported by Patterson (1952). A partial
explanation may be in the delayed hatch in 1973 as indicated from juvenile
wing molt (see Harvest Data). Previous researchers in North Park (Gill 1965,
Carr 1967, Poley 1969, May 1970) all reported concentrations of broods along
stream bottoms during the summer. However, of the 23 broods found in 1973
only one was along a stream bottom. Well distributed rains throughout the
summer resulted in apparent abundance of succulent forbs in sagebrush areas
and may be the reason that concentrations of grouse were not observed in
meadows.
Vegetation Transects
Relative frequencies of forbs, grasses, and shrubs for 1973 are presented
in Table 4. There were no significant differences (P &lt; .05) in relative
frequencies of shrubs, forbs, and grasses between sprayed and unsprayed areas.
Comparisons of relative frequencies with results of previous workers are of
little value due to varying weather conditions and differing times of the year
when measurements were taken. Density of sagebrush by age classes was determined on each plot and is presented in Table 5. Comparisons of sagebrush
densities for 1973 can be compared to densities recorded in 1964 and 1966 as
abundance of sagebrush is not dependent upon weather conditions for that year.
Spraying of sagebrush with 2,4-D was conducted during June, 1965, so densities for 1964 and 1966 represent the pre- and post-spray years (Table 6).

�-35Table 4. Relative frequencies by major plant groupings, 1973. Values
are means from 4 sprayed plots and 3 unsprayed plots in each sage type. ~/
Growth
Form

Forbs

Grass

Sage

Total Shrubs
Including Sage

Al Sprayed

35.3

43.0

9.6

21.6

Al Unsprayed

42.4

32.3

12.9

24.9

A2 Sprayed

31.0

46.8

9.3

22.1

A2 Unsprayed

36.1

33.9

14.7

29.8

A3 Sprayed

19.2

53.3

14.5

27.4

A3 Unsprayed

23.4

51.6

18.2

24.9

~/ All values are in percent.

Table 5.

Densities of Artemisia tridentata.
Al
Sprayed Unsprayed

Seedling-Young:
Strip
Block
x

Mature-Decadent:
Strip
Block

x

1.6
8.0
4.8
56.0
11.6
33.8

A2
Sprayed Unsprayed

1.2

22.4
8.1
15.2

55.0

14.5
11.2
12.9

5.5

11.8
l3.8
12.8

1.7

31.9
23.9
27.9

85.7

27.9
34.0
31.0

12.6

40.2
42.1
41.1

A3
Sprayed Unsprayed

4.1

43.8
38.2
41.0

25.9

32.6

22.2
19.0
20.6

47.6

6.6

19.8
12.9
16.3

12.7

8.1

47.8
34.0
50.9

30.8

74.6

29.0
25.6
27.3

53.7

Dead:
Strip
Block

x
Seedling-Young:
Strip
Block

x
Mature-Decadent:
Strip
Block

x

8.5
13.1
10.8
4.8
24.6
14.7

72.3
33.5
52.9

Dead:
Strip
Block
x

22.8
41.8
32.4

23.2
17.3

20.4
21.8

15.6

�-36-

Table 6. Densities of sagebrush by age classes and growth forms, 1964,
1966, 1973.

Growth
Form

Type of
Plot

Al

Sprayed

AI

Unsprayed

A2

Sprayed

A2

Unsprayed

A3

Sprayed.

A3

Unsprayed

Densit~ of Sagebrush (Plants/IOO ft~
1964i7
196627
1973
Seedling- MatureSeedling- MatureSeedling- MatureYoung
Decadent
Young
Decadent
Young
Decadent

1.0

6.0

76.0

50.0

28.0

22.0

0.1

10.0

4.8

33.8

0.0

44.0

1.2

55.0

3.0

11.0

15.0

13.0

0.8

29.0

4.0

32.6

43.0

15.0

41.0

20.6

137.0

68.0

26.0

47.6

1./ From Gill 1965.
J:../ From Carr 1967.
Table 6 may at first glance, pose more questions than provide answers in
regard to changes in sagebrush densities. One should first examine the data
for each year for relationships rather than just for numbers. The control
year, 1964 requires little explanation. From the 1966 data, collected one
year after spraying, it is apparent that sagebrush densities in sprayed plots
were lower within each growth form. Thus, it would appear that spraying
substantially reduced the amount of sagebrush. Two important relationships
are apparent in the 1973 data. First, in all growth forms, the density of
seedling and young plants in sprayed plots exceeded the density in the
unsprayed plots. This indicates the sagebrush is re-invading the sprayed
areas. Second, densities of mature and decadent sagebrush in sprayed plots
are still less than those found in unsprayed plots, thus sprayed areas
have not regenerated to the stage they were eight years ago. Based on the
1973 data, it is apparent that sprayed areas are still in the recuperative
stage following the spraying in 1965. The major discrepancy in Table 6 is
found in the unsprayed plots in the A-3 growth form. In both 1964 and 1966
density of seedlings and young plants greatly exceeded density of mature
plants, yet in 1973 the reverse was true. One possible, and probable,
explanation is observer differences in classifying plants as either young or
mature. In all plots total shrub occurrence is greater than sage occurrence,
primarily due to the high incidence of snakeweed (Gutierrezia) found in the
area. No mention of this species was made by previous researchers.

�-37-

Summer Movements and Distribution
Movements were determined from 20 band recoveries obtained during the
hunting season and from 6 observations of color banded birds during the
summer. Of the 20 birds killed during the season, 9 were banded on spring
concentration areas and 11 were banded on strutting grounds. The birds
banded on strutting grounds (1 female and 10 males) moved an average distance. of 3.1 miles, with a range of 0.5 to 5.5 miles. The nine birds
banded on spring concentration areas (6 females and 3 males) moved an
average of 10.4 miles with a range of 1.5 to 20 miles. Three of these
birds (two females and
male) moved 17, 20 and 17.5 miles, respectively. Of
the six marked birds observed in mid-summer, five were males banded on
strutting grounds. These birds had moved an average of 2.6 miles, with a
range of 1.2 to 7.2 miles. One unclassified bird which had been banded on
a spring concentration area was seen 5.5 miles from this area in late August.

a

Average distance traveled for females (N = 7) was 9.5 miles, with a range of
2 to 20 miles, whereas the average distance traveled by males (N = 13) was
4.7 miles, with a range of 0.5 to 17.5 miles. The long female movements
were to the south. Sample sizes were too small to detect any pattern in
directional movements.
Distribution information is based on the sighting of 405 birds in the Lake
John area from 11 June to 30 September 1973. Thirty-one percent (127) of
all birds sighted were in treated areas. Most of these birds were seen in
crested wheat (Agropyron spicatum) and rye (Elymuss spp.) fields. No birds
were seen in the large block spray area. According to observation data, no
grouse fed on the wheatgrass or rye. There is a substantial stand of young
sagebrush in these fields, which were disked and seeded in 1961. The large
number of sightings in the crested wheatgrass field may be biased as the
field is along a major road through the study area and birds are more visible
in grass cover. Of the 127 grouse seen in treated areas, 100 were adult
males. From these observations, it is apparent that treated areas are being
utilized by sage grouse, primarily bachelor flocks of males.
Harvest Data
Results of the 1973 check station operations are presented in Table 7 as
part of the 18 year summary. The age ratio in the harvest was 91 juveniles:
100 adults. Sex ratios were determined for each age class, with a ratio of
of 69 males: 100 females for juveniles and 44 males: 100 females for adults.
The 17 year pooled data ratios are 78 males:lOO females for juveniles, and
55 males: 100 females for adults.
All juvenile wings were aged in order to estimate the peak of hatch (Eng 1955).
Data from 60 wings indicated that 75 percent of the young hatched between 10
June and 30 June. The peak week was June 17-23 (35 percent), roughly eight
weeks after the peak of mating activity. This is considerably longer than
reported by others (Patterson 1952, Griner 1939), but may be explained by
adverse weather conditions during the spring of 1973. The young to adult
female ratio in the harvest varied little from the ratio obtained from brood
counts, 1.3:1 and 1.15:1, respectively.

�Table 7.

Year

North Park sage grouse hunter checks, 1955-1973.·Y

Bag
Limit

2
1955
1956
3
2
1957
1958
2
1959
3
1960
1961
3
1962
2
1963
2
2
1964
1965 2:../ 2
2
1966
1967
2
2
1968
2
1969
2
1970
2
1971
2
1972
2
1973

Hunters
Checked

Hours
Hunted

Adult
Cocks

221
349
198
285
257

641
1,251
728
863
1,003

47
119
45
51
72

193
551
592
217
150
306
300
546
662
564
357
452
366

612
2,001
2,460
624
626
1,227
1,177
2,604
2,936
2,617
1,802
2,377
1,599

Adult
Hens

Percent
Adults

41
46
132
48
70
58
71
45
151
67
No checks made
66
91
69
12
44
38
42
62
150
25
81
59
27
30
49
31
116
56
50
127
67
42
135
80
180
70
79
58
55
89
28
51
67
67
50
69
52
24
54

Birds
Hours
Unclassified
Per
Hunted
(Dressed) Total
Birds
Birds Hunter Per Bird

Juvenile
Cocks

Juvenile
Hens

Percent
Juvenile

48
122
36
48
40

85
150
46
102
68

59
52
42
55
33

57
129
25
150
44

283
652
222
422
375

1.28
1.67
1.12
1.48
1.46

2.27
1. 92
3.28
2.05
2.67

37
34
113
28
26
45
42
156
37
53
54
40
29

35
29
181
45
26
71
46
141
74
66
39
18
42

31
56
58
41
51
44
33
58
30
45
49
33
48

39
393
- 3/

268
506
506
179
116
263
267
512
569
266
264
253
193

1.39
0.91
1.03
0.82
0.77
0.86
0.89
0.94
0.86
0.47
0.74
0.56
0.53

2.28
3.95
4.86 I
3.49 w
00
5.40 I
4.67
4.41
5.09
5.16
9.84
6.83
9.40
8.28

l/ Based on Walden and Cowdrey check station data during opening weekends.
2:../ A one day season; hence only a one day check.

1/ Composition of kill projected from wing samples; hence no unclassified birds.

- 3/
- 3/
- 3/
- 3/

-]j
199

76
76
44

�-39-

Delineation

of Winter Distribution

One hundred thirty sage grouse flocks were sighted, amounting to 2,755
birds.
The winter period was divided into four two-week periods.
Number
of flocks and total birds seen during each period are presented in Table
8. No explanation is presently available for the pattern of sightings.
Weather data alone (average high and low temperatures and precipitation for
the period) do not provide evidence that climatic factors affected frequency
of sightings nor flock size. As can be seen from Table 9, the winter of
1973-74 was warmer than the winter of 1972-73 in North Park. The location
of each flock was plotted by period observed to detect obvious shifts in
areas utilized during the winter.
No significant changes were apparent.
In general, sage grouse could be found throughout the winter anywhere that
sagebrush was available above the existing snow cover.
It appeared that
flocks were very mobile, showing little fidelity to a certain area. However,
six areas of heavy use were evident from the plotted locations of 130 flock
sightings.
Sixty-eight percent (N = 130) of the flocks, and 71 percent of
the birds observed were seen in these six areas (Table 10). These six areas
comprise less than 27 square miles of the 611 square miles of sagebrush lands
in North Park (Rogers 1964). Undoubtedly, acreage of sagebrush in North Park
has been reduced due to sagebrush control activities since Roger's work in
the early 1960's.
The exact acreage of disturbed sagebrush lands in North
Park is presently unknown.
This will be investigated during the 1974 field
period.
One estimate is that less than 15 percent of the 611 square miles of
sagebrush lands have been altered.
Approximately 75 percent of the sagebrush lands provided possible habitat for sage grouse in the winter of 197374. Most areas where snow completely covered the sagebrush were in the
southern part of North Park, where elevations range 500-600 feet higher than
in the northern part. The indicated heavy usage of these 27 square miles of
sagebrush lands shows selectivity by grouse rather than random distribution
throughout the available sagebrush lands.
Birds could not always be found in preferred areas, indicating movement into
and out of the area.
Flock sizes in these areas varied and since birds are
not individually marked, it was impossible to tell if a particular flock was
habitually in an area.
It was also impossible to judge stability in numbers
of a flock. There were no significant differences in the vegetal and physical
characteristics measured between the six heavy use areas and the remaining
areas of sightings.
At present, data are not available to show obvious
reasons for birds selecting these areas.
These six areas were not searched more frequently than otheI less productive
areas, thus eliminating the bias of heavy use due to a high frequency of
observation time.
Searches for sage grouse were conducted during 60 days
during the winter period.
At least one or more of the six heavy use areas
was searched on 34 of the total days. Areas that were subsequently determined to be used lightly or not at all were covered on at least 25 days
during the period.

�Table 8.

Observations of sage grouse flocks, North Park, winter 1974.

Time Period

Hours
of
Search

Total
Flocks

Flocks
Observed/
Hour

Total
Birds

Birds
Observed/
Hour

70

18

0.26

283

4.0

17

191

1

52

-

40

121.5

45

0.37

826

6.8

23

305

17

427

5

94

Male Flocks
No. of
Total
Flocks
Birds

Female Flocks
No. of
Total
Flocks
Birds

Unclassified
No. of
Total
Flocks
Birds 1/

January 8January 21
January 22 February 7

I

February 8 -

.p-

o
I

107

21 ']j

0.20

576

5.4

10

206

7

290

3

80

March 8

110

46

0.42

1,070

9.7

17

192

25

693

4

185

Total

408.5

130

0.32

2,755

6.7

67

894

50

1,462

12

399

February 21
February 22-

1/ One flock of 50 percent males and females.
~/ Includes unclassified birds from male and female flocks.

�1/
2/
Table 9. Temperature- and precipitation- data for winter months, North Park.
November
~:ean Daily Me an
Nax.
Hin. Daily

Pp t ,

Winter 1973-74

37.2

17.4

27.3

Winter 1972-73

33.3

9.2

~~x.

December
Daily Mean
Hin. Daily

Mean
Ppt. Max.

January
Daily ~~an
Hin. Daily

1.91

31.4

6.6

19.0

0.69

25.4

2.5

21.3

0.31

24.5

3.0

13.8

1.51

21.7

-4.3

28.2

0.48

17.0

0.57

Hean

Ppt.

Mean
Max.

February
Daily Mean
Hin. Daily

Ppt.

Mean
Max.

March
Daily Mean
Min. Daily Ppt.

14.0

0.99

29.2

2.8

16.0

0.18

41.6

20.6

31.1

0.94

8.7

0.25

25.0

-5.6

9.7

0.03

33.8

8.1

21.0

0.54

12.9

0.53

17.6

0.42

22.6

0.39

"Average"
Winterl/

I

.;-

•...
I

11 In degrees Fahrenheit.

!:. / In inches.
.~/ Averages for 1961-67 (temperature) and 1961-70 (precipitation).

�-42-

Only two of the six heavy use areas are in the original intensive study
area. One, the California Gulch area (No.4, Table 10) is the only area in
the Lake John study area where wintering sage grouse were located in the
winters of 1963-64, 1964-65 and 1965-66 (Gill 1965, Carr 1967). Both of
these researchers reported concentrations of birds along Colorado State
Highway 125 north of Walden. This area corresponds to winter use area
number two in Table 10. Birds banded on Lake John complex strutting grounds
were observed in each of the six high use areas.
Characterization of Winter Flocks
Sage grouse in North Park are gregarious, being found in flocks most of
the time. Only nine single birds (seven males, two females) were seen in
the winter of 1973-74 out of a total of 2,755 birds observed. Strong segregation by sexes is evident in sage grouse flocks. Of the 118 flocks classified, 74 percent (87) were composed of at least 90 percent birds of like
sex (Table 11). Male flocks were smaller than female flocks, averaging 11
birds, compared to 32 females per flock. Flock size was less than 50 birds
in 92 percent of all cases. The strong segregation of sexes and differing
flock sizes suggests the possibility of differential niche selection within
the sagebrush vegetation type. This question will be further investigated
in the next segment, as present data indicate no apparent differences in
characteristics of areas used by male and female flocks.
Characterization of Winter Use Sites
The vegetal characteristics for 68 flock locations are presented in Table
12. Measurements were not taken at all flock locations as this required
flushing the birds which would prevent trapping during the night. In these
situations, strong winds and snowfall would frequently change characteristics
of the site before the area could be revisited the following day. Measurements were only taken at sites where the sex classification of the flock
could be determined, thus eliminating 13 flock locations. Statistical
analysis of this data has not been completed. The only difference appears
to be that female flocks may be utilizing denser stands of sagebrush than
male flocks.
Slope appears to be a significant factor in selection of a use site by sage
grouse. The average percent slope for areas used by female flocks was 4.9,
and for male flocks was 6.9. Sixty-four percent (N = 87) of the areas had
slopes less than 5 percent, while only 2 percent of the areas had slopes
greater than 15 percent. Many areas of North Park have abundant sagebrush
available during the winter on slopes greater than 15 percent, but few birds
were ever seen in these areas.
Eng and Schladweiler (1972) noted that sage grouse winteriIlg in the Winnet
area in central Montana utilized nearly level terrain, avoiding areas of
steep slopes even though such areas had dense stands of sagebrush.
Aspect also appears to be a significant factor in site selection. Seventyone percent (N = 86) of the flock locations were on south, southwest, or
west faciIlg slopes. The prevailing wind in North Park is from the southwest
and strong wiIlds were present most days during the study period.

�Table 10.

Areas of high use by wintering sage grouse, North Park.

Description

Area

Approximate
Square Miles

No. of Flocks
Observed

Total Birds
Observed

1

NE 1/4 Sect. 13, E 1/2 Sect. 12; R80W, T10N
Sect. 18, W 1/2, SE 1/4 Sect. 7; R79W, T10N

2.5

14

114

2

SE 1/4 Sect. 19, S 1/2 Sect. 20, SW 1/4 Sect. 27,
S 1/2, NW 1/4 Sect. 28, Sect. 29, E 1/2 Sect. 32,
Sect. 33, Sect. 34; R79W, T10N
N 1/2 Sect. 3, N 1/2 Sect. 4, NE 1/4 Sect. 5, R79W

6.5

16

214

19

223

p

T9N
I

3

SE 1/4 Sect. 1, E 1/2 Sect. 12, NE 1/4 Sect. 13; R81W, T9N
SW 1/4 Sect. 7, NW 1/4, S 1/2 Sect. 18, N 1/2 Sect. 19;
R80W, T9N

2.5

4

SE 1/4 Sect. 7, S 1/2 Sect. 8, N 1/2, SE 1/4, Sect. 17,
W 1/2 Sect. 16, NE 1/4 Sect. 20, Sect. 21; R80W, T10N

3.25

8

179

5

E 1/2 Sect. 7, SW 1/4 Sect. 8, SW 1/4 Sect. 16, Sect. 17,
E 1/2 Sect. 18, E 1/2 Sect. 19, Sect. 20, Sect. 21,
SW 1/4 Sect. 22, W 1/2 Sect. 27, Sect. 28, Sect. 29,
NE 1/4 Sect. 30, NE 1/4 Sect. 33; R78W, T9N

8.25

19

866

6

S 1/2 Sect. 31, S 1/2 Sect. 32: R78W, T8N
W 1/2, NE 1/4 Sect. 5, Sect. 6, N 1/2 Sect. 7, NW 1/4
Sect. 8, R78W, T7N

3.5

13

371

26.5

89

1,967

Total f
}

.j::W
I

�-44-

Table II.

Sex composition of winter flocks of sage grouse, North Park. 1/

Size of Flock

51-59

60-69

Percent in Flock
70-79
80-89

90-100

Total

Males
0-9

1

2

1

29

33

10-19

1

3

21

25

20-29

1

2

3

6

1

3

6

54

67

3

2

11

16

10-19

1

2

12

15

20-29

1

2

2

5

30-39

1

1

2

5

40-49

3

3

75-99

1

1

50-74

1

1

Sub-total

1

2

4

Females
0-9

&gt;100
Sub-total

1

1

2

2

5

6

2

9

33

50

117

Total

1

8

6

15

87

Percent of all
Flocks

1

7

5

13

74

1/ Excluding one flock of 50:50 males and females.

�-45-

Table 12.

Vegetal characteristics of winter use sites.

N

Mean Sagebrush
Density
(Plants/O.Ol Acre)

Mean Height of
Sagebrush Above
Snow (em)

Mean of Average
Crown Breadth of
Sagebrush (cm)

M

16

50.8

25.1

47.6

0-9

F

9

67.7

26.0

50.1

10-19

M

15

42.6

18.9 .

48.9

10-19

F

7

79.4

19.4

42.6

20-29

M

5

47.2

27.2

56.6

20-29

F

5

55.2

20.4

55.2

30-39

F

4

71.5

20.8

38.6

40-49

F

2

61.5

25.5

67.7

50-74

M

2

60.0

12.5

31.4

75-99

F

1

35.0

20.0

56.0

&gt;100

F

2

59.5

21.0

59.9

x for male flocks N = 38

47.6

22.3

48.4

x for female flocks N = 30

66.8

22.3

51.0

No. in
Flock

Main Sex
of Flock

0-9

Winter Movements
A total of 71 banded birds were seen in the winter of 1973-74, of which only
five were females. Of the 66 banded males sighted, 51 were banded with
colored bandettes. Average distance traveled from banding sites for males
was 6.7 miles (N = 51), and for females 11.9 miles (N = 5). The small number
of female sightings prevents valid comparisons with males. Eighty-four
percent(N = 51) of the male movements were in the following four directions:
north, northeast, east, southeast. This corresponds strongly with the
directions of the main winter use areas from banding sites. The longest movement was 21.8 miles by a male trapped on a strutting ground in May, 1973.
There are several possibilities as to why so few banded females were~bserved.
One is that females have a higher mortality rate than males, thus red~ing
the number of banded females in the winter population. Another possibility
is that there are many more females in the population than males, thus

�-46-

resulting in more unbanded females in relation to banded females than is
the case for males. A third possibility is that major areas of winter use
by banded females were not located. Observational data from the winter of
1973-74 support the second possibility.
Of the 2,755 birds observed, 2,356 were classified as to sex. The sex ratio
of these 2,356 birds was 61 males:lOO females (894:1,462). This represents
a significant deviation from a 1:1 sex ratio and is distorted in favor of
females. This ratio is very similar to that (55 males:lOO females) derived
from pooled harvest data for adults from 1955-1973. Of the 1,462 females
observed, 886 were definitely unbanded, while only 5 were banded. The maximum number of banded females in the population was 102 (number banded in
spring minus known hunting mortality). Of the 894 males observed, 462 were
definitely unbanded and 66 were banded. The maximum number of banded males
in the population was 166. The much lower ratio of banded to unbanded birds
for females supports the observed distorted sex ratio. Further evidence
which supports the observed ratio is the sex ratio of juvenile birds in the
harvest. The sex ratio determined from check stations over the period 19551973 is 78 males:lOO females.
Efforts will be made during the next research segment to document seasonal
movements of females and extensive searches for female flocks will be
conducted.

Aerial Search
Aerial censuses were flown along random transect lines in North Park on
February 21-22 and March 17, 1974. The flights in February were conducted
under ideal observing conditions, bright sun and fresh snow cover. The
southern half of North Park was covered on 21 February. The search was
halted by a low cloud mass moving in from the north. The north half of
North Park was covered the following day. Five flocks totaling 108 birds
were counted. An extrapolation of this figure (108 birds, 9 percent of the
area) would result in a sage grouse population of 1,200 birds. This is
suspected to be a low estimate. It is important to note that none of the
transects included any of the six high use areas. Two transects passed
along the boundaries of two wintering areas. The possibility of using an
orderly random census method to include these areas will be investigated
next winter. The second flight, on 17 March resulted in few observations, 3
flocks and 27 birds, due to poor visibility caused by patchiness of snow
cover. The Lake John study area was 90 percent bare ground on 8 March and
observation of grouse flocks in this area would have been most difficult.
The plotted sightings of grouse flocks were used in outlining winter
distribution.
Analysis of Data from Previous Years
Graphs were made in attempts to correlate various weather characteristics,
strutting ground counts, brood counts, and harvest in a multitude (greater
than 60) of combinations. However, no visible correlations existed.

�-47-

Regression values will be calculated for two possible correlations which
compare percent of juveniles in harvest to the number of days where minimum temperatures were below 00 C during the months of May and June. Results
of all correlations will be presented in the final report.

LITERATURE CITED
Autenrieth, R. E. 1969. Sage grouse investigations.
Rept. Idaho Fish and Game Dept. 25 p. Mimeo.

Third Annual Prog.

1970. Sage grouse investigations. Fourth Annual Prog. Rept.
Idaho Fish and Game Dept. 21 p. Mimeo.
Bean, R. 1941. Life history studies of the sage grouse (Centrocercus
urophasianus) in Clark County, Idaho. M. S. Thesis, Utah State Agric.
Coll., Logan.
Bent, A. C. 1932. Life histories of North American gallinaceous birds.
Smithsonian Inst., U. S. Nat. Mus. Bull. 162. 490 p.
Braun, C. E., and R. K. Schmidt, Jr. 1971. Effects of snow apd wind on
wintering populations of white-tailed ptarmigan in Colorado. In
Haugen, A. o. (Ed.). 1971. Proc. Snow and Ice Symposium, Iowa State
University, Ames. 280 p.
Carr, H. D. 1967. Effects of sagebrush control on abundance, distribution,
and movements of sage grouse. Colorado Game, Fish and Parks Dept.,
Game Res. Rept., Proj. W-37-R-20. 106 p.
Crunden, C. W. 1963. Age and sex of sage grouse from wings.
Mgmt. 27(4):846-849.

J. Wildl.

Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F.
Schlatterer. 1963. Ecology, productivity and management of sage grouse
in Idaho. J. Wildl. Mgmt. 27(4):811-841.
Ellison, L. N. 1973. Seasonal social organization and movements of spruce
grouse. Condor 75(4):375-385.
Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios
from wings. J. Wildl. Mgmt. 19(2):267-272.
_____ , and P. Schladweiler. 1972. Sage grouse winter movements and habitat
use in central MOntana. J. Wildl. Mgmt. 36(1):141-146.
Gill, R. B. 1965. Effects of sagebrush control on distribution and movements of sage grouse. Colo. Game, Fish and Parks Dept., Game Res.
Rept., Part 3. W-37-R-17. 185 p.
Griner, L. A. 1939. A study of the sage grouse (Centrocercus urophasianus),
with special reference to life history, habitat requirments, and numbers
and distribution. M. S. Thesis. Utah State Agric. ColI. 83 p.

�-4B-

Gullion, G. W. 1970. The ruffed grouse in northern Minnesota. Univ.
Minnesota. Forest Wildl. Relations Proj. 37 p. Multilith.
Hoffman, R. W. 1973. Migration of white-tailed ptarmigan to and from a
major wintering area. Colorado Div. of Wildlife, Job Prog. Rept.
W-37-R-26. 30 p.
June, J. W. 1967. Sage, blue and ruffed grouse sexing and aging characteristics. Wyoming Game and Fish Comm. 59 p.
Klebenow, D. A. 1969. Sage grouse nesting and brood habitat in Idaho.
J. Wildl. Mgmt. 33(3):649-661.
1970. Sage grouse versus sagebrush control in Idaho.
23(6): 396-400.
-----, and G. M. Gray. 196B.
Range Mgmt. 21(2):BO-B3.

J. Range Mgmt.

Food habits of juvenile sage grouse.

J.

Koskimies, J. 1957. Flocking behavior in capercaillie, Tetrao urogallus
(1.), and blackgame, Lyrurus tetrix (L). Finnish Papers on Game
Research No. lB. 32 p.
Lumsden, H. G. 196B. The displays of the sage grouse. Res. Rept. (Wildlife)
No. B3. Ontario Dept. of Lands and Forests. 94 p.
Martin, N. S. 1970. Sagebrush control related to habitat and sage grouse
occurrence. J. Wildl. Mgmt. 34(2):313-320.
May, T. A. 1970. Effects of sagebrush control on distribution and abundance
of sage grouse. Colo. Game, Fish and Parks Div. Job Prog. Rept. Proj.
W-37-R-23, Work Plan 3, Job Ba. 23 p.
Oakleaf, R. J. 1971. The relationship of sage grouse to upland meadows in
Nevada. Nevada Dept. of Fish and Game. Job Final Report W-4B-2. 64 p.
Patterson, R. L. 1952.
Denver. 341 p.

The sage grouse in Wyoming.

Sage Books, Inc.

Peterson, J. G. 1970. The food habits and summer distribution of juvenile
sage grouse in central Montana. J. Wildl. Mgmt. 34(1):147-155.
Poley, B. E. 1969. Effects of sagebrush control on distribution and abundance of sage grouse. Colo. Game, Fish and Parks Div. Job Compl. Rept.
Proj. W-37-R-22, Work Plan 3; Job Ba. 31 p.
Rogers, G. E. 1964. Sage grouse investigations in Colorado.
Fish and Parks Dept. Tech. Pub. 16. 132 p.

Colo. Game,

�-49-

Scott, J. W.
498.

1942.

Mating behavior of the sage grouse.

Auk 59(10):477-

Seiskari, P. 1962. On the winter ecology of the capercai11ie, Tetrao
uroga11us, and the black grouse, Lyrurus tetrix, in Finland. Finnish
Papers on Game Research No. 22. 119 p.
Walles tad , R. O. 1971. Summer movements and habitat use by sage grouse
broods in central Montana. J. Wi1d1. Mgmt. 35(1):129-136.
_____ , and D. B. Pyrah. 1973. Movements and nesting requirements of sage
grouse hens in central Montana. J. Wi1d1. Mgmt. (In Press).
_____
, J. G. Peterson, and R. L. Eng.

grouse in central Montana.

1973. Food habits of adult sage
J. Wi1dl. Mgmt. (In Press).

Weeden, R. B. 1964. Spatial separation of sexes in rock and willow ptarmigan
in winter. Auk 81(4):534-541.

~d~·
J~'...........;d;.."..-;;.~-----=-~_

Prepared By_J=--:;;~~·
Thomas D. I. Beck
Graduate Research Assistant

��April

1974

-51-

JOB

State of
Project

FINAL REPORT

CO::..::..:LO=RAD=~O
_
W-37-R-27

No.

Work Plan No.

Game Bird Survey
Job Nc.

8

Job Title

Inventory

Period

April 1, 1959 to March

Covered:

Personnel:

of Lesser Prairie

2

_

Chickens

31, 1974.

Charles H. Gibson and Elmer Miller, U. S. Forest Service, Glen Eyre,
Lloyd Hazzard, Daniel Potts, Walter Schuett, Warren Snyder and
Donald Hoffman, Colorado Division of Wildlife.

ABSTRAcr
A basic inventory of lesser prairie chickens (Tympanuchus pallidicinctus)
was conducted during the four year period from 1959 through 1962 in southeastern Colorado.
A re-inventory was completed during the two year period
from 1972 through 1973, approximately 10 years after the basic inventory
on this species.
During the spring of 1962, 130 lesser prairie chickens
were counted on 13 active grounds.
Included in this total were 104 cocks,
one hen and 25 unclassified as to sex. Total numbers of lesser prairie
chickens counted in the spring of 1973 on 10 active booming grounds was 129,
and included 60 cocks, 5 hens, and 64 unclassified as to sex. In occupied ~
habitat southeast of Holly, Colorado in Prowers County, two booming grounds
were active in 1962 and 1972. Only the one ground nearest the state line
was being used in 1973. Populations had increased from a total of 14 on
two grounds in 1962 to 28 on the one ground in 1973, indicating redistribution
had occurred.
Extensive disking to reduce sand sagebrush and increase
grasses in 1962 on privately owned grazing lands resulted in no apparent
changes in habitat by 1973. In two areas southwest and southeast of Campo,
Colorado in Baca County, three booming grounds were located during the period
1959 through 1962. All three grounds, however, had disappeared by 1972.
No major habitat changes or land use practices were noted in comparing 1973
with 1962 but an isolation factor may have resulted in their disappearance.
In occupied habitat south of the towns of Vilas and Walsh in Baca County, 69
lesser prairie chickens were counted on six active booming grounds in 1962
compared with 72 lessers counted on seven active booming grounds in 1973.
Populations have remained relatively stable, even though two grounds on private
land disappeared with large scale gravel mining operations, and two grounds
apparently moved for unknown reasons.
Five new grounds were established since
1962 to stabilize this population.
In the area south of the Cimarron River
in Baca County, three active booming grounds with a total of 31 birds were
counted in 1962, compared with a total of 29 birds counted on two active
booming grounds in 1973.
Populations have remained relatively stable, even
though some redistribution has occurred.
Fewer resident farm families, additional clearing of sand sagebrush habitat for farmground, and installation of
several circular sprinkler systems were changes noted from 1962 to 1973.

�-52RECOMM:ENDATIONS

Continued spring counts of lesser prairie chickens on booming grounds are
reconunended for this species, which has been classified as endangered in
Colorado. Occupied habitats in Colorado are on the northwestern fringe of
their historical range, so quality of habitat is 'lower than found in Oklahoma and Kansas. For this reason, booming grounds in Colorado are often not
as permanent as those in higher quality range. Annual counts are reconunended
since population shifts and changes are more easily determined than use of
counts every five or 10 years. Only through periodic inventories can the
future welfare of lesser prairie chickens be revealed.
Future inventories should, however, be accomplished by Southeast Region
personnel of the Division of Wildlife. Cooperative counts with personnel
of the U. S. Forest Service for all booming grounds located on the Comanche
National Grasslands are reconunended. Six inventory booklets listing all
past counts and locations have been prepared for individuals involved and
Wildlife Conservation Officers in the Springfield, Colorado and Lamar, Colorado districts have been orientated in ground locations and counting
procedures.

�-53-

INVENTORY

OF LESSER PRAIRIE CHICKENS
Donald M. Hoffman

P. S. OBJECTIVE
To determine

the population

status of lesser prairie

chickens

in Colorado.

METHODS AND MATERIALS
Early morning listening checks were made along roads and trails to locate
booming grounds during the periods April through June, 1959 through 1962,
and 1972 through 1973. Reports of past sightings were used along with
surveys in preferred habitat types to locate populations.
Most of the potential lesser prairie chicken habitat in Baca, Bent, Las Animas and Prowers
counties were searched during the years of the study. Limited amounts of
time were spent in Cheyenne, Kiowa, and Lincoln counties searching for
populations without success.
Potential lesser prairie chicken habitat along
the Big Sandy Creek drainage from Chivington in Kiowa County to south of
Hugo in Lincoln County was worked very little because of its inaccessibility.
Numbers of males and females using booming grounds were determined through
early morning counts.
Flushing counts were secured on both accessible and
inaccessible
grounds.
A partial inventory of known populations in southern
Baca County was made by Warren D. Snyder in 1963.
Persons living and working within lesser prairie chicken
viewed to secure leads on locations of new flocks.

ranges were inter-

Maps of locations of booming grounds were drawn for future reference
inventory booklets were prepared to assist in future inventories.

and

RESULTS AND DISCUSSION
Table 1 lists highest numbers of lesser prairie chickens counted by individual booming ground and by year during the years 1959-1962 and 1972-73.
Data for 1963 when only a partial count was secured is also included in
Table 1. Grounds were numbered consecutively by date located, and individual grounds were further identified by land ownership in naming the grounds.
Where more than one ground was located on an individual landowner's place,
these were designated by the letters b, c, d, etc., in the order located.
The map locations of all known booming grounds (active and abandoned) are
shown in Figs. 1, 2, and 3.
Total numbers of birds counted in 1973 on 10 active booming grounds were
approximately equal to the number counted in 1962 on 13 active grounds (129
in 1973 compared with 130 in 1962, Table 1). Only five of the 13 active
grounds being used in 1962 were still active in 1973. Of the 130 birds
counted in 1962, 104 were cocks, one was a hen and 25 were unclassified as
to sex (Table 1). Of the 129 birds counted in 1973, 60 were cocks, five
were hens, and 64 were unclassified as to sex (Table 1).

�Table 1.

Summary of highest lesser prairie chicken booming ground counts, 1959-1973.

Booming Ground
Number

I

...;:t
II)

I

Name

Year
1959
Cocks Unclass.
Hens
Total

1 C. Low (a)
4
2 T. Sta1ford (a) 0
3 Dye, Glover (a) 2
4 T. Sta1ford (b)
5 T. Stalford (c)
6 Hanes (a)
7 Perkins,
Tanner (a)
8 C. Low (b)
9 Hanes (b)
10 Dye, Glover (b)
11 T. Sta1ford (d)
12 Lowder (a)
13 Dye, Glover (c)
14 Schnauffer (a)
15 Chick, Dow (a)
16 Shell Oil (a)
17 Dye, Glover (d)
18 Dye, Glover (e)
19 Shell Oil (b)
20 Dye, Glover (f)
21 Schnauffer (b)
22 T. Sta1ford (e)
Totals

2
0
2

6 4

0
8
0

8

6
8
4

18

1/ Counts by W. D. Snyder (1963).

1960
Cocks Unc1ass.
Hens
Total

1961
Cocks lhc1ass.
Hens
Total

1962
Cocks Unc1ass.
Total
Hens

1963.1.
Cocks Unc1ass.
Total
Hens

1972
Cocks Unc1ass.
Hens
Total

1973
Cocks Unc1ass.
Total
Hens

7
10
6
7
2
6

2
0
5
0
0
0

0
2
0
0
0
0

9
12
11
7
2
6

6 4
0
7 0
2
12 0
0
Abandoned
5 0
8
0 0
13

10
9
12

5 1
0
10 0
3
12 0
3
Abandoned
13 0
3
12 0
8

Not counted
Not counted
20 2
0
22
Abandoned
Not counted
19 0
7
26

0 0
8
Abandoned
11 1
0
Abandoned
0 0
23
Abandoned

Abandoned
Abandoned
12 0
1
Abandoned
25
0 0
Abandoned

1

0

0

1

7
5
0
4
2
6
0
2

8
6
7
4
2
10
4
2

5 0
0
5
Not counted
Abandoned
4 0
0
4
Not counted
17 1
0
18
0 1
15
16
Abandoned
0 0
13
13
0 0
7
7
11 0
11
0
0 0
3
3

Abandoned
0 0
20
Abandoned
Abandoned
Abandoned
10 0
5
15 1
0
Abandoned
Abandoned
Abandoned
Abandoned
5 0
0
4 0
3
4 0
0

76 4

49

39

7

2

48

56

1
0
0
0
0
1
0
0

6

0
1
7
0
0
3
4
0

13
13

38 100

4 0
2
7 0
1
Abandoned
5 0
1
2 0
0
13 0
3
7 0
0
Abandoned
9 0
1
5 0
0

104 1

25

6
13
15
16
20
6
8
6
2
16
7
10
5

130

45

125

2

59

8
12
23

20

15
16

5
7
4

110

13
25

Abandoned
12 1
15
Abandoned
Abandoned
Abandoned
12 0
2
13 3
0
Abandonecj
Abandoned
Abandoned
Abandoned
2 0
6
7
0 0
6 1
3
0 0
4
1
3 0

8
7
10
4
4

64

129

60

5

28

14
16

�-55Populations and Habitat Changes by Area-1962 to 1973
(1) Area Southeast of Holly, Colorado in Prowers County (Fig. 1).
Although two grounds (Table 1 and Fig. 1, numbers 1 and 8) were active in 1962
and 1972 only the one ground (number 8) nearest the state line was being used
in 1973. Populations had increased from a total of 14 on the two grounds in
1962 to 28 on the one ground in 1973 (Table 1). Several large tracts, including the booming ground sites and 'surrounding areas were known to'have been
disked in the spring of 1962 to reduce sand sagebrush and increase grasses
on privately owned grazing lands. The sand sagebrush had recovered by 1972
with little evidence noted, except for scattered dead sand sagebrush stems.
A large area to the west of the booming grounds was aerial sprayed with a
herbicide in the spring of 1973 also, to reduce the amount of sand sagebrush and increase the grasses.
(2) Areas Southwest and Southeast of Campo, Colorado in Baca County (Fig. 2).
During the early 1960's, small isolated populations of lesser prairie
chickens were found approximately five miles southwest of Campo, Colorado
(Table 1, Fig. 2, ground number 7), and four to six miles southeast of Campo
(Table 1, Fig. 2, ground numbers 14 and 15). Both of these isolated populations,
however, had disappeared by 1972. All other resident populations in Colorado
are in close proximity with populations in either Kansas or Oklahoma so it is
possible that the isolation factor may have resulted in their eventual disappearance.
No major apparent changes in habitat was found for either area in comparing
1973 with 1962 and land use practices appeared to be similar
(3) Area South of Vilas and Walsh, Colorado in Baca County (Fig. 3).
During 1962, a total of 69 lesser prairie chickens were counted on six ,active
booming grounds in this area (Table 1, Fig. 3, numbers 3, 6, 10, 12, 13 and
16) compared with 72 lessers counted on seven activing booming grounds in 1973
(Table 1, Fig. 3, numbers 3, 12, 13, 18, 19, 20, and 21). Populations have
remained relatively stable even though two grounds on private land (Table 1,
Fig. 3, numbers 6 and 9) have disappeared due to gravel mining operations,
and two grounds on Federal lands (Fig. 3, numbers 16 and 17) have moved for
unknown reasons. One ground (Fig. 3 number 17 was discovered in 1963, but
was abandoned by 1972). There is little or no apparent change in habitat
orland use patterns in this area except as noted above. This area is pri~
marily grazing lands within the Comanche National Grasslands, and appears
to have the best potential for long term survival of the species in Colorado.
(4) Area South of Cimarron River in Baca County (Fig. 3).
Three booming grounds in the area south of the Cimarron River were active
in 1962 (Table 1, Fig. 3, numbers 2, 5 and 11) with a total of 31 birds, and
two (Table 1, Fig. 3, ground numbers 5 and 22) were being used in 1973 with
a total of 29 birds, so populations appear to have remained relatively stable,
even though some redistribution has occurred. One ground (Fig. 3, number 4)
was discovered in 1960, but was abandoned in 1961. There were only two farm

�R,.4IW.

\
T. 23 S.

23

19

24

..;
I

26

~

25

I

--

2

II

o

.

27

26

2~

•
o

21

H

26

25

30

"II

29

•

=~:~i
R ·~~~~[JJ~f~l=~tJi5:T~T~:;~~",3.-= -1.

U.-

,.

28

I

~..=....--~

II
II
II

•

6

I)

V1
0\
I

5

/'
.

II

II

.2

7

0

0

/

T

10

"

12

I
I
I
I
I
I

,

r

"

'I
I
I
I

'IL

_IF

Q

I
I
I
I

.

t

Fig. 1.
County.

Location and reference number (Table 1) of lesser prairie
Active grounds (in 1973) are indicated in solid color.

chicken booming

grounds

in Prowers

�r~-~-~

KEY

I

N~

rl ---~''777777'7777777777'77'!".

T\ J

----"
27

i

I
~~

II

_,
•

-

II

26

/

30

25\

2.

29

\

II

'

II
II

~~~

~~~~~J'

,,"

,,"

"
W

TOT

.,

I

"

-

""
::

27

II

II
II

~
:~~~~~~"
bII

II

33"

."

::

34

I

1

"

\"

- ~,

1,"

::C
"9'-.

T.34S
I
VI

~
I

En

36

~·\'·b":~.
~
:~:~'.,

.e
-. '; '::.&gt;:;i.,.:

:':'~-~&gt;::::it::.

1t j

10

E~~

Fig. 2. Locations and reference numbers (Table 1) of lesser prairie chicken booming grounds
central Baca County.
All grounds shown on this map were classified as abandoned (in 1973).

-,37(;

in south-

�N

I

II

IIH -

, I

"

25

![~.

~

'.H"

Ilwz ~l-IrrT I " '\

q,
~::~,

30

Ii

29

~

I

2.

~27

~

~

ii

i 2.

\:o"~

25:1

t

29

r

2.

27

2.

N

,

I~

.~ ~"

II

L

••

:."'1

III
i.1

SHEET:iI :::'

",~~
"',,

z
__

SHEET

51

I

32

[]

I

33

34

fl

35

II

36

2

1

I
1
1

'I
I

~
121

I

2.

25

I

t

I

I'

II
3.

II

31

I

32

I I

33

I

3'

I

3S"':- 41

301

I"

'~i

I

"r

J"

,22 ,
13

R.44W.

I

n: •.••
~ vv

"'

Colo.'

w

I

ra

I

'7

I

I.

I

'5

I

11

"
"

1

'2

I

,.

:35J
.-

R.42W.

Fig. 3. Locations and reference numbers (Table 1) of lesser prairie chicken booming grounds in ~outheastern Baca County. Active grounds (in 1973) are indicated in solid color.

VI
00
I

�-59-

families residing in the area in 1972 and 1973, compared with four families
in 1962, similar to trends in many farming areas of Colorado and elsewhere.
Several circular sprinkler systems have been installed and are in use in this
area. In addition, two large tracts near the booming ground locations of
approximately one square mile each have been cleared of sand sagebrush and
are now being farmed. These are farther west than previously farmed areas.
The remaining sand sagebrush-grassland type appears similar to that of the
early 1960's.
SUMMARY AND CONCLUSIONS
Basic inventories of lesser prairie chickens conducted during the period
1959 through 1962, and 1972 through 1973 indicated overall populations in
Colorado have remained relatively stable during the period. Isolated populations southwest and southeast of Campo, Colorado in Baca County have disappeared, possibly due to an isolation factor. Overall status remained similar
due to establishment of new grounds and redistribution of established populations.
A total of 130 lesser prairie chickens was counted on 13 active booming
grounds in 1962, compared with 129 on 10 active booming grounds in 1973. The
total popUlation of lesser prairie chickens in Colorado is probably less than
250 birds based upon these inventory results.
Annual counts to be accomplished by Southeast Region personnel of the Division
of Wildlife are recommended since population shifts and changes are more
easily determined than using periodic counts conducted every five or 10 years.
The Colorado Division of Wildlife hosted the Tenth Conference of the Prairie
Grouse Technical Council on September 5-7, 1973 in Lamar, Colorado. Fiftytwo people from various states with an interest in prairie grouse research
and management attended. A proceeding of 22 pages was compiled and distributed to those attending.
PUBLICATIONS RESULTING FROM STUDY
Hoffman, D. M.
28-30.

1960.

Prairie chicken comeback.

Colorado Outdoors 9(3):

1963. The lesser prairie chicken in Colorado.
27(4):726-732.

J. Wildl. Manage.

1965. Prospects for the extension of lesser pra~r~e chicken range
in Colorado. Paper presented at Sixth Conference, Prairie Grouse Tech.
Council, Warroad, Minnesota. September 14-16. 3 p. Mimeo.
1973. Changes in populations and habitats of lesser prairie chickens
in Colorado, 1962 to 1973. Paper presented at Tenth Conference, Prairie
Grouse Tech. Council, Lamar, Colo. September 5-7.7
p. Mimeo.
(Compiler). 1973. Proceedings Tenth Conference Prairie Grouse_Tech.
Council, Lamar, Colorado. Colo. Division of Wildlife, Denver. Sept.
5-7. 22 p.

�-60-

LITERATURE CITED
Hoffman, D. M. 1959. Habitat requirements of lesser prairie chickens. Fed.
Aid Job CampI. Rept. Proj. W-37-R-13. Colo. Dept. of Game and Fish,
Denver. p. 9-31.
1961. Census of lesser pra~r~e chickens. Fed. Aid Job CampI. Rept.
Proj. W-37-R-14. Colo. Dept. of Game and Fish, Denver. p. 195-201.
1962. Census of lesser prairie chickens. Fed. Aid Job Camp!. Rept.,
Proj. W-37-R-15. Colo. Dept. of Game and Fish, Denver. p. 93-99.
1963. Census of lesser prairie chickens. Fed. Aid Job CampI. Rept.,
Proj. W-37-R-16. Colo. Dept. of Game, Fish and Parks, Denver. p. 145149.
1973. Inventory of lesser pra~r~e chickens. Fed. Aid Job Prog.
Rept., Proj. W-37-R-26. Colo. Division of Wildlife, Denver. (In Press).

Prepared by

i~~~

Donald M. Hoff
Wildlife Researcher

�-61 ....
April, 1974
JOB PROGRESS REPORT

State of

COLORADO
--------~~---~----W-37-R-27

Project No.

Game Bird Survey

--------------------~

Work Plan No.

10

Job No.

2

----~--------------------------

Job Title

Experimental Breeding of Hungarian Partridge

Period Covered:

April 1, 1973 to March 31, 1974.

Personnel:

Many individuals have contributed to the completion of this study.
A complete listing is found in Hoffman et a1. (1974b).

ABSTR.,.\CT
.
Publications and a manuscript awaiting final editing covering results of
this study include:
Hoffman, D. M. 1973. Reproductive performance in the gray partridge.
Ph.D. Dissertation. Dept. of Animal Science,Colorado State University,
Fort Gollins. 89 p.
_____ , C; L. Quarles,L. A. Webster, and L. R. Crooks. 1974a. Raising
gray (Hungarian) partridge. Game Bird Breeders Gazette 23 (2):30-35.
1974b. Experimental propagation of
----, and
gray partridges. Special Report. Colo. Division of Wildlife, Denver.
(awaiting final editing and publication).

��-63-

EXPERIMENTAL BREEDING OF HUNGARIAN PARTRIDGES
Donald M. Hoffman

P. S. OBJECTIVE
To develop gamebird farm production techniques for Hungarian or gray
partridges (Perdixperdix perdix).

SEGMENT OBJECTIVE
To compile data, summarize, and prepare final reports.

RESULTS AND DISCUSSION
The objective of this study has been completed and three manuscripts have
been written. Two of these manuscripts including an article submitted to
the Game Bird Breeders Gazette and a Ph.D. dissertation have been either
published or accepted. A third manuscript to be published in the Colorado
Division of Wildlife's Special Report series was completed and submitted
to the Small Game Section Leader on November 29, 1973. Final editing and
publication will be accomplished during 1974.

,('
Prepared by

..

(k~~&lt;rrJ~~
Donald M~f~
Wildlife Researcher

..

��April 1974

-65-

Jon l'ROGP-ESSREPORT

State of

.

C::..'O::.:LOf
...=";A...:;T::...;) 0=---'-

W-37-R-27

Project No.
Work Plan No.
Job Title

_

Game Bird Survey

~~----~-----------

17

Job No.

2

Continued Inventory of Selected Ptarmigan Populations

Period Covered:

April 1, 1973 to October 20, 1973

Personnel: Clait E. Braun, Richard W. Hoffman, Tom Lytle and Charles Wagner.

ABSTRACT
Investigations of white-tailed ptarmigan (Lagopus leucurus) populations in
Colorado initiated in 1965 were continued in 1973 with major emphasis in
ascertaining popUlation levels and relationships of nesting success and
production to climatic conditions. Breeding densities decreased on all areas
studied except Crown Point where they increased. Reasons&gt; for the observed
decreases at Mt. Evans and Independence Pass were related to heavy htinter
harvest in 1972. The continued decrease at Rocky Mountain National Park
was related to lower overwinter hen survival and poor recruitment of sub,...
adults into the breeding. popuLatLon , Nesting failed at Crown Point in 1973
and success was low (about 30 percent) on all areas studied •. The observed
low nesting success was possibly related to age of females and late nesting
due to cooler temperatures and late lying snow cover in 1973. Hunter interest
was low in 1973 and pressure was almost non-existent at the two study areas
open to hunting.

��-67-

CONTINUED INVENTORY OF SELECTED PTARMIGAN POPULATIONS
Clait E. Braun
Knowledge concerning population levels of white-tailed ptarmigan and factors
influencing observed densities is essential for proper management. This
project was undertaken to examine long term population trends and effects of
climatic conditions on white-tailed ptarmigan in Colorado. The initial 5
years of research on this grouse have been previously reported by Braun and
Rogers (1971). This report presents data collected during the fourth year
of the last 5-year segment. Data collected during the first three years of
the second 5-year segment were presented by Braun (1971, 1972, 1973).
P. S. OBJECTIVES
To test the hypotheses that (1) populations of white-tailed ptarmigan in
Colorado are not cyclic, (2) production and fall population densities of
this species in Colorado are related to temperature and moisture conditions
during June and early July of the same year, and (3) harvest of over 50
percent of the fall population adversely affects breeding densities the
following spring.
SEGMENT OBJECTIVES
1.

To estimate breeding population levels of ptarmigan in 4 study areas (Mt.
Evans, Crown Point, Independence Pass, and Rocky Mountain National Park).

2.

To estimate nesting success and production in the above areas.

3.

To obtain weather data from a representative alpine area for use in
determining correlations between spring weather conditions and fall
ptarmigan populations.

4.

Manipulate hunting seasons at Mt. Evans through closures, reduced length,
late timing, etc., in order to affect breeding densities.

5.

To compile data and prepare progress report.
METHODS AND MATERIALS

Techniques used were essentially those developed under Work Plan 17, Job 1
and reported in detail by Braun and Rogers (1971), and updated by Braun
(1971). In 1973 green bandettes with black numerals were used to mark all
newly banded birds and those whose older bandettes had become too worn for
individual recognition at Crown Point and Independence Pass. Yellow bandettes
were used in Rocky MOuntain National Park, while white, green, yellow and
blue bandettes were used at Mt. Evans.

�-68-

Weather data were obtained from the Institute of Arctic and Alpine Research
(INSTAAR), University of Colorado, for reasons previously explained (Braun
1971). Hunting statistics were collected through return of bands from
successful hunters (Independence Pass and Crown Point).
Description of Area
Areas intensively investigated have been described in detail by Braun and
Rogers (1971) and have been presented by Braun (1971).
RESULTS AND DISCUSSION
Breeding Densities
Censuses of breeding birds were initiated on May 5 in Rocky MOuntain National
Park, with several pairs being observed along with lone males, and males with
more than one female. Breeding surveys were initiated on May 11 at Mt. Evans,
May 12 at Crown Point, and May 31 at Independence Pass. Most pairing in 1973
appeared to have occurred between May 1 and 15 except at Independence Pass
where it appeared to have occurred in mid- to late May. While timing of
breeding events was similar to that observed in most recent years except 1971,
fidelity to breeding territories was sporadic due to frequent storms in May.
Densities observed are presented in Table 1.
Table 1.

White-tailed ptarmigan breeding densities, all areas, 1973.

No. of
Breeding
Pairs

Unmated
Males

Total
Breeding
Population

Birds per
Square
Mile

1.25

7

4

18

14.4

Toll Memorial

.19

3

3

9

47.4

Fall River Pass

.70

6

4

16

22.9

Total Rocky Mountain National
Park

2.14

16

11

43

20.1

Crown Point

1.93

3

3

9

4.7

Mt. Evans

1.54

11

3

25

16.2

Independence Pass

1.12

8

3

19

17.0

Study Area - S.
1ze-1/
Rocky Mountain National Park
Tombstone Ridge-Sundance Mtn.

1/ In square miles.

�-69-

Breeding densities in 1973 decreased o.nall study areas except Cro.wn Po.int
f rom levels obse rve'd in 1972 as illustrated in Table 2. The significant
increase at Cro.wn Po.int (fro.m1.0 to.4.7 birds per square mile) was expected
earlier (1972), after the co.mp1ete remo.va1 o.f all breeding birds in 1971.
It wo.uld appear that it takes at least two. years fo.ra po.pulatio.no.f ptarmigan
to.reco.ver fo.11o.wingmajo.r lo.sses. This waS indicated fro.m earlier wo.rk at
Mt. Evans (Braun 1971, 1972). Po.pulatio.nsat Mt. Evans o.n1y decreased abo.ut
3 birds per square mile fo.11o.wingthe o.verharvest (54 percent o.f the fall
po.pu1atio.n)during the 1972 hunting seaso.n. Ho.wever, o.ver o.ne-ha1f (15 o.f
25) o.f the breeding birds at Mt. Evans in 1973 were subadu1ts, thus reflecting the heavy harvest of adults the preceding fall. At Independence Pass
appro.ximate1y 60 percent (61.1) o.f the fall po.pu1atio.nwas harvested in 1972.
While this o.verharvest was no.t reflected in a majo.r decline in breeding
po.pulatio.nin 1973 (a decrease o.fo.n1y 1.7 birds per square mile), it was
reflected in the age co.mpo.sitio.n
o.f the breeding po.pu1atio.n. Only 3 o.f 19
birds were o.lder than 1 year. In co.ntrast, 35 o.f 43 breeding birds at Ro.cky
Mo.untain Natio.na1 Park were o.lder than 1 year in 1973. The decrease o.bserved
at R.M.N.P. in 1973 was the fo.urth year o.f the decline first o.bserved in 1970
(Table 2). This po.pu1atio.nhas no.w decreased by o.ne-third as the decline is
the result o.f increased hen mortality and decreased recruitment of yo.ung
birds into. the spring breeding po.pu1atio.n. With the exceptio.n o.f1970, the
breeding po.pu1ation at Independence Pass has remained remarkably stable since
1966. Apparently, little affects the po.pulatio.nat this site except severe
hunting pressure fo.r at least 2 successive years (Braun 1971).
Table 2.

White-tailed ptarmigan breeding densities 1966-73.

Study Area

1966

1967

Birds Per Sguare Mile
1968
1971
1969
1970

1972

1973

Ro.cky Mo.untain
Natio.na1 Park

29.0

25.2

29.4

30.8

24.8

23.4

22.4

20.1

Cro.wn Po.int

14.5

21.2

18.1

14.5

10.4

5.7

1.0

4.7

Mt. Evans

7.8

7.1

7.1

5.8

5.2

11.0

19.5

16.2

Independence Pass

18.7

18.7

18.7

17.9

10.7

21.4

18.7

17.0

Nesting Success and Pro.ductio.n
Laying was initiated o.nmo.st areas by June 10, with incubatio.n underway as
early as June 20. One nest was lo.cated in 1973, fo.und o.n June 20 with 5 eggs.
This nest was unsuccessful as it was destro.yed by a weasel o.n June 22. The
earliest calculated hatching date was July 23 and the latest was August 4.

�-70-

Calculated peak of hatch was between July 23-30, about one week later than
that observed in most years. Nesting success in 1973 was poor, possibly the
lowest of the 8 years of study. Nesting at Crown Point failed in 1973, as
no successful hens were located. In all, only 10 successful hens were located
on the study areas, whereas 12 unsuccessful hens were observed. This is
misleading, as unsuccessful females do not respond to tape recorded calls
and have low fidelity to breeding areas, whereas successful hens do. Estimated nesting success on all areas was between 25 and 30 percent in 1973.
This could be attributed partly to inexperience of hens (most were subadults)
and delayed nesting due to late-lying snow cover. In a larger survey in
1973, R. Hoffman (personal communication) located 22 successful and 12
unsuccessful females. His data also suggest that nesting success was lower
in 1973 than in 1972. In 1972 in the same area, he located 37 successful
hens and 8 without broods. His data suggests that nesting success in 1973
decreased by about 40 percent (40.5) from levels estimated in 1972 (about
70-75 percent).
Initial brood size in 1973 upon hatching was about 4 chicks. By September 1
this number had decreased to about 3 chicks per succ~ssful female (Table 3).
These data are different from all years, except 1969 when average brood size
was one chick less than normal on September 1. Normal brood size to September
1 is 4 chicks per successful hen.

Table 3.

Month

Number of broods and average brood size, all areas, 1973.l1
Number of
Broods Observed

Average Number of
Chicks per Brood

July

0 (0)];.1

0.0

August

10 (22 )?:..I

3.4 (3.3)];.1

September

2

(O)?:..I

(0)];.1

3.0 (O)?:..I

II Only distinct broods are included.

21 Data from R. Hoffman gathered in the Guanella Pass-Loveland Pass area
(personal communication).
Fall Densities
Estimates of densities of white-tailed ptarmigan on September 1, while useful in illustrating population gain through production, are difficult to
accurately derive due to a number of variables. Estimated fall densities
for each area studied are presented in Table 4. These densities were calculated following the three basic assumptions discussed in detail by Braun
and Rogers (1971).

�Table 4.

Estimated fall densities of white-tailed ptarmigan, 1973.
Total
Population
on
September 1

Estimated
Percent
Nesting
Success

Average
Brood Size
on
September 1

Total
Production

Total
Breeding
Population

Rocky Mountain
National Park

30

3.2

16

43

55

Crown Point

0

0.0

0

9

8

4.1

0.0

Mt. Evans

30

4.0

16

25

39

25.3

41.0

Independence Pass

30

3.7

11

19

28

25.0

39.3

Area

Birds Per
Square Mile

Percent Gain

25.7

29.1

-..JI

•...•
I

�-72-

In 1973 all alpine areas of the state were open to ptarmigan hunting from
September 8 through October 8, except for the area lying within ~ mile of
the Mt. Evans highway where the season was closed. As in recent years, no
ptarmigan hunting was allowed during any of the early big game seasons.
Bag and possession limits for ptarmigan were 3 and 6.
With the closure of the area along the Mt. Evans highway, no check station
was operated in 1973 and no reports of hunting were received. No bands
were returned from Crown Point for the second consecutive year, reflecting
both low hunting pressure and the few birds available. Reports from
Wildlife Conservation Officers Tom Lytle at Leadville and Al Whitaker of
Aspen indicated no hunters at Independence Pass in 1973. Only one band
was received from this site and the reporting hunter failed to respond to
an inquiry about his hunting success. All observations indicate that few
hunters were afield for ptarmigan in 1973. Reasons for this are not known.
Relationships of Weather:Nesting Success and Production
Weather data obtained from the 12,300 ft level of Niwot Ridge, through the
courtesy of INSTAAR are not yet available for 1973. These data are normally
available about June 1 of the year following collection. These data will be
obtained once available, and detailed analysis and statistical correlation
will be completed for the final report in 1975-76. In general, weather conditions in 1973 appeared to be cooler and possibly wetter than in 1972, with
the result that much of the study areas remained snow covered into mid-June.
LITERATURE CITED
Braun, C. E. 1971. Continued inventory of selected ptarmigan populations.
Colo. Div. of Game, Fish and Parks, Game Res. Rept. Fed. Aid Proj.
W-37-R. April. p. 91-106.
1972. Continued inventory of selected ptarmigan popul.at Lons , Co l.o ,
Div. of Wildl. Game Res. Rept. Fed. Aid Proj. W-37-R. April. p. 91-98.
1973. Continued inventory of selected ptarmigan populations. Colo.
Div. of Wild 1. Game Res. Rept. Fed. Aid Proj. W-37-R. April. In Press.
and G. E. Rogers. 1971. The white-tailed ptarmigan in Colo.
---,Game,
Fish and Parks. Tech. Publ. No. 27. 80 p.

Prepared by __

J-6~/~r.a:'~1-~,;~7_-~(~;;....).::..;2d.ivW~_'
Clait E. Braun
Wildlife Researcher

_

Colo.

�April 1974

-73-

JOB PROGRESS REPORT

State of

C;;..;O:;.,;;LO=RAD=..;;;.O
_

Project No.

W-37-R-27

Work Plan No.

17

Game Bird Survey
Job No.

----3------------------~~

Job Title Experimental Removal of a Breeding Population of White-tailed Ptarmigan
Period Covered:
Personnel:

April 1, 1973 to October 30, 1973.

Clait E. Braun and RichardW.

Hoffman

ABSTRACT
A minimum of 9 white-tailed ptarmigan (Lagopus leucurus) occupied the Crown
Point study area in 1973. This total was comprised of3 pairs and 3 urunated
males. All females and at least 1 male were .subadult s, with the other 4
unmarked males also classed as subadults based on their behavior. The.l
subadult male marked on the area in 1972 returned in 1973. No broods were
observed in 1973, and most, if not all birds left the area by early August.
All areas used by ptarmigan had been used in prior years of the study.

��-75-

EXPERIMENTAL REMOVAL OF A
BREEDING POPULATION OF WHITE-TAILED PTARMIGAN
Clait E. Braun
Management of wild animals is dependent upon knowledge about population
levels, reproductive capacity and success, survival and mortality rates,
and habitat requirements. This study to investigate habitat preferences and
pioneering ability of white-tailed ptarmigan was initiated in 1971 after
five years of previous study of the Crown Point population (Bratm and
Rogers 1971, Bratm 1971). This report presents data collected during the
third year of a 4 year study. Data collected during the first 2 years of
the study were presented by Bratm (1972, 1973).
P. S. OBJECTIVE
To test the hypotheses that (1) ptarmigan select sites for breeding territories which contain readily available bushes of Salix spp. (&gt; 10 cm in
height), and snowfree locations, (2) immature birds initially pioneer new or
empty habitats, and (3) number of occupied territories is affected by age
class of male ptarmigan.
SEGMENT OBJECTIVES
1.

To map and compare number and characteristics of all occupied breeding
territories at Crown Point.

2.

To compile data and prepare progress report.
METHODS AND MATERIALS

Ptarmigan were located during the breeding season through use of taperecorded male challenge calls as described by Bratm et al. (1973). Techniques used in capturing, observing, banding, etc. were essentially those
described in detail by Braun and Rogers (1971) and updated by Braun (1972).
Description of Study Area
This area was described in detail by Bratm and Rogers (1971) and will be
redescribed in the final report. Extent of the area investigated was shown
in the initial report (Braun 1972).
RESULTS AND DISCUSSION
Population Status
The study area was systematically surveyed 3 times during the breeding season
and twice during the late summer brood period. A total of 13 different birds

�-76-

were observed of which 8 were newly banded. Six males occupied territories
in 1973 of which 1 was banded as a subadu1t in 1972, 1 was banded as a subadult in 1973, and 4 were unmarked males. These males were extremely wild
and were thought to be subadults based on behavior and reaction to the taperecorded male challenge call. Three of these males were unmated. Seven
females were banded on the study area, and 2 additional unmarked hens were
observed in early May. Only 3 hens became paired and remained on the study
area. All other females disappeared by late May and are presumed to have
moved to the Mummy Range in search of suitable males or territories.
During the breeding period only 3 pairs and 3 unmated birds occurred on the
Crown Point study area. This density of 4.7 birds per square mile is up
significantly from 1972 (1.0 birds per square mile) but still below the 5.7
birds per square mile that were present in 1971 when the removal experiment
was initiated. Areas occupied in 1973 are shown in Fig. 1.
No broods were located at Crown Point in 1973, and it is assumed that all
hens were unsuccessful. Observations indicated that most, if not all birds
had left the area by early August.
Description of Breeding Territories
All areas used by ptarmigan in 1973 had been used in one or more prior years
and had been previously described (Braun 1971). All areas occupied in 1972,
1973, and 1974 will be vegetatively described in 1974.

LITERATURE CITED
Braun, C. E. 1971. Habitat requirements of Colorado white-tailed ptarmigan.
Proc. Western Assoc. State Game and Fish Comms. 51:284-292.
1972. Experimental removal of a breeding population of white-tailed
ptarmigan. Colo. Div. Wildl., Game Res. Rept. Fed. Aid Proj. W-37-R.
April. p , 99-109.
1973. Experimental removal of a breeding population of white-tailed
ptarmigan. Colo. Div. Wi1d1., Game Res. Rept. Fed. Aid Proj. W-37-R.
April. In Press.
Braun, C. E.,and G. E. Rogers. 1971. The white-tailed ptarmigan in Colorado. Colo. Div. Game, Fish and Parks. Tech. Publ. No. 27. 80 p.
, R. K. Schmidt, Jr., and G. E. Rogers. 1973. Census of Colorado
-----white-tailed
ptarmigan with tape-recorded calls. J. Wildl. Mgmt.
37(1) :90-93.

Prepared by _--!:a~d.-::=:'.L..· -=Z:..;..
.. -J&lt;..~==-:...;-'---C1a.it E. Bra.un
Wildlife Researcher

�-77-

"~....
0

6'00

~.---.,

I

.
•

/

~

__

-'.

--~.

-

--.:

-

I"
I
I
'1,

, -,
J
I

XII14S,"'"

10600

••.

....

"'-.

--

-""'-_.

,

I

..

-~--.---

-~-

"

/'

&lt;&gt;:&gt;,
;;:.
~. ".

."

----

~----.

CROWN

AREA

COLORADO

N

f

POINT

-

Study Area Boundary

.,~'

Fig. -1.

Crown Point study area

(X's mark occupied breeding

areas,

1973).

��April 1974

-79-

JOB FINAL REPORT

State of

COLORADO
----------~~~~--------~

Project

No.

W-37-R-27

Game Bird Survey

17

Job Title:

4
: Job No.
C~h-a-r-a-c-t~e~r~i-s~t~i-c-s--an~a
Migration ot Wintering
P_o~p_u_l_a_t_i_o_n_s_·_o~f
__C_o_l_o_r_a_d_o
__W_h_l_Ot_e_-_t_a_l_Ol_e
__d__P_t_a_rm
__i_g_an

Period

April 1, 1972 through March 31, 1974.

Work Plan No.

Covered:

Personnel:

_

C. Braun, H. Funk, P. Gilbert, R. Hoffman, D. Luce, R. Oakleaf, and
C. Wagner, Division of Wildlife; D. Hein, P. Lehner, S. Martin and
R. Ryder, Colorado State University; T. May, University of Colorado,
and J. Kitzmiller.

ABSTRACT
Studies of the characteristics
and migration of wintering populations of
white-tailed ptarmigan (Lagopus leucurus) were conducted in north-central
Colorado from April 1972 through April 1974 primarily at Guanella Pass.
In
all, 293 km2 of alpine .habl t at were selected for studying movements from and
to the wintering area.
Guanella Pass was utilized principally by females (80 percent of 799 birds
identified) and primarily adults (65 percent).
Numbers of subadults varied
with year depending upon production success.
The 172 winter flocks observed
ranged from 2 to over 80 birds per flock, but most contained 2 to 25. Females
typically congregated in larger flocks (20-30. birds) than males « 15 birds);
probably the result of differences in winter habitat preference as partial
segregation of sexes occurred.
Apparently, winter flocks were not composed ot
family units nor were the same group of individuals associated together
throughout the winter.
Sixty percent of the ptarmigan banded at Guanella Pass
and relocated on breeding or summering sites returned in subsequent winters.
Adults exhibited the greatest affinity for the wintering grounds with 73
percent returning, while only 51 percent of the subadults returned.
Willow
(Salix spp.) was the most important vegetation found on wintering areas.
Essentially all sites on the winter range were utilized, but some preferences
were noted.
Major factors affecting distribution were availability of willow
and snow quality.
Eight chick movements were documented to wintering or breeding sites.
Hunter recoveries indicated chicks were relatively sedentary prior
to brood dispersal.
Females banded as chicks traveled greater distances than
males.
Males showed high fidelity to natal areas.
Ninety-nine and 45 movementswere
documented from and to winter use sites, respectively.
Subadults
were more mobile than adults, with subadult females traversing the greatest
distances.
Female grouse were consistently more mobile than males.
Average
movements of males were generally restricted to less than 2.5 km for adults
and 4.0 km for subadults, while movements of females frequently exceeded
these distances.
The longest movement was 22.7 km for a subadult female.
Seventy-four percent of the movements from Guanella Pass were in westerly
directions with 28 percent to the northwest.

�-80-

ACKNOWLEDGEMENTS
-;

I wish to thank Dr. Ronald A. Ryder, Professor, Department of Fishery and
Wildlife Biology, Colorado State University, for serving as my graduate
advisor, for providing advice and assistance throughout my graduate training,
. and for reviewing and editing this thesis.
Thanks are expressed to Dr.
Stephen Martin, Assistant Professor, Department of Zoology, Colorado State
University,
for serving as a memberof my graduate committee and for reading
this manuscript.
Special thanks are extended to Dr. Clait E. Braun, Wildlife Researcher,
Colorado Division of Wildlife, who.was instrumental in initiating
this project
and assisted with fieldWOrk throughout the study.
Dr. :Braun as a memberof
mycommittee, contributed many helpful suggestions, provided additional data,
and critically
evaluated the thesis.
His understanding and encouragement -are
greatly appreciated.
..

.'

I also thank the Colorado Division of Wildlife through Federal Aid in Wildlife
. Restoration Project W-37-Rfor providing equipment, financial support, and use
of their facilities
during this project.
Howard Funk, Project Leader,Colorado
Division of Wildlife, provided administrative
assistance and reviewed this
manuscript.
Myappreciation is extended to all other Division of Wildlife
personnel who contributed to this proj ect.
in field
to Ronald Oakleaf and James Kitzmiller .for assisting
work, ".and to D:r.Dale Hein , . Associate Professor, .Department of Fishery and
Wildlife Biology, Colorado State Uni:versity, for providing helpful suggestions.
Thanks are extended to'rerry
A. May for aiding in fieldwork
and providing
suggestions concerning my research, arid to Dr. PhillipN . Lehner, Assistant
'Pro fes sor , Department of Zoology, for evaluating the thesis.
I am grateful

:".

:

",

Finally, tomy wife, Barbara, I wish to express my deepest appreciation
her support and. patience, .without which I could not have completed this
study.

for

�-81-

LIST OF FIGURES
Figure
1

Montezuma-Georgetown
Distribution
Colorado

86

study area

of alpine tundra and ptarmigan

in

3

Guanella Pass study area

4

Percentage of different age and sex classes of
white-tailed ptarmigan returning to Guanella
Pass •

5

6

87

89

100

Distances traveled by white -tai.Ied ptarmigan
from wintering to breeding or summering a re a s .' .

. 105

Directions and distances recorded for movements of white-tailed ptarmigan from Guanella
Pass to breeding or summering sites

• 107

�-82-

LIST OF TAB LES

Table
1

Arrival
and departure
Guanella Pass
.

dates

of pta rrn ig an at

2

Sex and age cornpo s it ion of white -tailed
pta rrn igan wintering at Guanella Pass
.

3

Nurnb e r and frequency of encounters
with lone
birds and flocks of white -tailed pta rrn igan,
Guanella Pass,
1966-1974

4

5

6

7

91

92

94

Yearly trends in flock size of white-tailed
pta rrntgan at Guanella Pass
.

• 94

Monthly trends in flock size of white -t ai Led
pta rrn igan at Guanella Pass
.

• 95

Com puted chi square values for corrrpa r is ona
between nurnb e r s of white -rai Ied' pta rrn igan of
different
age and sex classes
returning to
Guanella Pas s .

• 99

Contrasting
statistical
analyses for distances
rn oved by different age and sex classes
of
white -tailed pt a rrn iga n .

·106

�-83-

TABLE OF CONTENTS

INTRODUCTION

• 85

DESCRIP TION OF STUDY AREAS

85

METHODS AND MATERIALS.

88

RESULTS AND DISCUSSION

• 90

Winter Investigations
Arrival and Departure.
Sex and Age Composition.
Flocking Behavior
Distribution.
Affinity for Wintering Sites

90
90
• 91
• 93
97

.

Movements
Movements of Chicks to Wintering and
Breeding Areas
•
Movements from Wintering to Breeding
or Summering Sites.
Movements from Breeding or Summering
to Wintering Sites
SUMMARY.
LITERATURE

• 98
• 102
· 102
.104
• 109
• 110

CITED

•113

��-85-

INTRODUCTION

White-tailed ptarmigan (Lagopus leucurus), the only species of Lagopus
confined to North America, occupy alpine habitats from south-central Alaska,
the Yukon Territory south through the mountains of western Canada and United
States to northern New Mexico (Aldrich 1963). The range of the southern
white-tailed ptarmigan (Lagopus leucurus altipetens) in Colorado correlates
with the distribution of alpine tundra from 3353 to over 4267 m in elevation,
an area of approximately 9713 km2 (Braun and Rogers 1971).
Seasonal movements of white-tailed ptarmigan have been described in general
terms by several authors.
Packard (1945), Racey (1948), Bailey (1927),
Bent (1932), and Gabrielson and Lincoln (1959) describe the species as
spending the summer above timberline and descending to or below treeline in
winter.
Choate (1963), suggested some flocks remain above timberline during
the winter, while others move below timberline.
Braun (1969) documented the
movements of white-tailed ptarmigan in Colorado throughout the year.
His
studies indicated that males are relatively sedentary, usually moving less
than 2 km in a year, while females are highly mobile with some birds moving
16 to 24 km from wintering to breeding areas.
in Colorado (Braun 1969, Haskins
extensive data on
general ecology, population dynamics, movements, habitat, seasonal foods,
behavior and parasites.
In order to further understand seasonal movements
of ptarmigan in Colorado, this study was initiated in 1972. The primary
objective was to document movements by age and sex class from and to a
major wintering area in order to ascertain the importance of such areas to
surrounding breeding sites and to determine the directions and distances
involved in annual migrations.
In addition, attempts were made to: (1)
document departure and arrival dates to and from wintering areas, (2) record
and analyze winter flock sizes, (3) describe the distribution of birds on a
wintering area, (4) ascertain the affinity of birds for individual wintering
sites, and (5) individually identify chicks on summering areas in order to
document their movements to wintering areas.
Intensive

studies of white-tailed

ptarmigan

1969, Schmidt 1969, and May 1970, 1974) have provided

DESCRIPTION

OF STUDY AREAS

Approximately 293 km2 of alpine habitat in the Front Range of the Rocky
Mountains in Colorado were selected for studying movements of ptarmigan to
and from a major wintering area. This area, hereafter referred to as the
Montezuma-Georgetown
study area (Fig. 1), centers around Guanella Pass,
which was chosen as the primary winter study area principally because of
the relative abundance of wintering ptarmigan, accessibility,
and availability of information on this area from prior studies (Schmidt 1969, May
1970, Braun and Schmidt 1971). In addition, limited winter investigations
were conducted near Waldorf, Naylor Lake, Steven's Gulch, and Horseshoe
Basin where there were small numbers of ptarmigan.
Fig. 2 depicts the
location of study areas in relation to alpine tundra in Colorado.

�-86-

.

.

r-I

rx..

eo
•..-l

�• Cre.,

• ,e,t ,elll ••

~

~
~
,

-'.

• O ••••.• r
••TrV •••
CU'.ILL •••••
NAnOfil

LA'"

I
• G'.lIIt

00

,h"'ti ••

"-J

I

~

o

.c•.••
_ ••..•••

•

b

t

"MI ••10n,O"""

.,.

(;P

.'" .....
Fig.

2.

Distribution
and Rogers

of alpine
1971).

tundra

and ptarmigan

in Colorado

(after

Braun

�-88-

Weimer and Haun (1960) have described the geology of the Front Range in
Colorado while Judson (1965) and Marr (1961) have discussed the climatic
features of the area. Marr (1961) also characterized the vegetation of the
alpine tundra in the Front Range of Colorado.
Descriptions of habitats
seasonally occupied by ptarmigan in Colorado were presented by Braun (1971).
Much of the above information concerning the Guanella Pass area has been
summarized by Braun and Schmidt (1971) and only limited discussion follows.
Guanella Pass lies 13 km south of Georgetown, Colorado, in the Pike and
Arapaho National Forests.
Area investigated is in T5S, R74W in parts of
Clear Creek and Park counties at 1050 37'W long., and 390 35'N lat. Total
area investigated was 9 km2 (Fig. 3). The four additional wintering sites
briefly investigated were smaller, varying in size from 2 to 4 km2.
Guanella Pass is primarily dominated by Salix spp. Higher, in Krummholz
areas, Salix is codominant with Picea engelmannii.
The presence and abundance of Salix spp. were the most important factors determining where
ptarmigan wintered on all areas studied.
The topography is rolling, with
slopes ranging from 5 to 30 percent and elevations from 3475 to 3655 m above
sea level.
Snow depths vary considerably within the study area, however,
much of the area is unprotected from the prevailing westerly winds with
Salix never completely snow covered.
The larger Montezuma-Georgetown
study encompassed approximately 293 km2
but only 65 to 78 km2 were considered suitable breeding and summering sites.
Alpine areas searched included portions of T4S, R75 and 76W; T5S, R73, 74,
75 and 76W; and T6S, R73, 74, 75 and 76W in Clear Creek, Park, and Summit
counties.
The topography is irregular, varying from precipitous slopes and
sharp peaks to gently rolling expanses of hundreds of hectares.
Elevations
range from over 4267 to 3353 m. About 99 percent of the area studied is
public land administered by the U. S. Department of Agriculture, Forest
Service.
METHODS

AND MATERIALS

Winter groups of ptarmigan were located by concentrated searching on foot
with periodic stops to scan surrounding areas for tracks, snow roosts, and/
or birds themselves (Braun and Schmidt 1971).
During the breeding and brood
seasons, ptarmigan were located through use of tape recorded calls (Braun et
al. 1973), while summer flocks of males and unsuccessful females were found
by systematically searching rocky areas along ridgetops.
Once birds were
located, they were first observed through 7 X 50 binoculars to determine if
they were banded.
Also, efforts were made to ascertain sex of adults and
count the birds present.
Attempts were made to determine band color and
number of those birds already banded.
Some unbanded ptarmigan and all birds
with unreadable bandettes were pursued until caught or flushed and not
relocated.
Each observation was recorded at the time on standardized observation cards (Braun 1969). All locations were subsequently plotted on 7.5
minute U. S. Geological Survey topographic maps.
Birds were captured with a 5 or 7 m long telescoping noose pole as described
by Zwickel and Bendell (1967) for blue grouse (Dendragapus obscurus) and

�',~

'.,--- --'
,

/
Ii'
;-

~,\,.I•f•t..nkf
...•.
J-"':-

/
/'

&lt;,

/'

~

-,

-. r-:

,Pj{j-.........

G\Janelia

p~

s

/'
--"J
&lt;;

V/
---

r-"

T(.p
I

00

\0
I

The
Sawtooth
'~

" GUANELLA

r

t --

PASS

COLORADO
Siudy

Area Bo un d r y

SCAl:E

IN

o

==:J

~~;\
'::9JO

/'

t::1
:d

-

f!~

-1-

~. ?l~~

-@' ",
co
-'---;---------,
j

APPROXJWAT£80UWOA'RY

&lt;,

~-"

Fig. 3.

__

._.

'

\':\

Guanella Pass study area.

KlL(H/.[rCi{$
1/2

I

~

AREA

_

.,.~

�-90-

reported by Braun and Rogers (1971) for white-tailed ptarmigan.
Upon
capture, birds were banded, and classified to age and sex following methods
described by Braun and Rogers (1971).
Distribution of ptarmigan on the wintering area was determined by periodically traversing the entire area and through reports of sightings from
reliable observers.
Sites were classified as being utilized if tracks,
roosts, droppings, or ptarmigan were observed.
Data were available on the
location of wintering flocks at Guanella Pass since 1966 (files of C. E.
Braun, Colorado Division of Wildlife).
Although all sites on the wintering
area were not equally searched, it is believed that during the 7 years for
which data were available, all sites being utilized by ptarmigan were noted.
Arrival and departure dates to and from the wintering area were documented
by frequently searching the study area in October and April.
Throughout the
winter, efforts were concentrated on identifying as many birds as possible.
Surrounding breeding and summering areas were searched from late April until
mid-September to locate birds banded on wintering sites. During the summer,
hens and chicks were banded to document their movements to winter sites.
Movements were plotted on 7.5 minute (scale 1:24,000) U.S.G.S. topographic
maps.
Straight lines were drawn from initial banding sites to recovery
locations, thus, all movements represent minimum distances traveled.
Directions in which movements occurred were determined from wintering areas to
breeding or summering sites where birds were observed.
All information
about each movement and bird involved was recorded on a standardized form.
Hunter bags were checked on weekends during 1972 and 1973 primarily to obtain
data on banded birds harvested.
Data prior to 1972 were available from
hunter recoveries, collections, and observations, courtesy of C. E. Braun.

Arrival

RESULTS

AND DISCUSSION

Winter

Investigations

and Departure

Winter was arbitrarily designated as late October, when ptarmigan became
entirely white, through mid- to late April when the prenuptial molt of males
was initiated.
During these 6 months, ptarmigan were rarely observed away
from winter use sites.
As previosuly reported (Braun and Schmidt 1971),
arrival and departure to and from wintering areas were greatly influenced
by climatic conditions.
Prolonged mild weather during the fall of 1972
delayed the arrival of birds to Guanella Pass until early November, while
extended winter conditions the following spring delayed departure until early
May. However, in most years birds were on the wintering area by late October
and remained within the area until late April when they departed for breeding
areas. Arrival and departure dates are shown in Table 1.

�-91-

Table 1.

Arrival and departure dates of ptarmigan at Guane11a Pass.

Winter

Arrival Date

Departure Date

1971-72

23-26 October

19-22 April

1972-73

3-5 November

3-10 May

1973-74

20-26 October

27-30 April

Sex and Age Composition
A total of 799 ptarmigan were identified on the Guane11a Pass study area from
1966 through 1974. These included observations and recaptures of birds
banded in previous winters, newly banded birds, and collections for scientific
study. Significantly (P&lt;O.OOl)
more females than males were identified each
year (Table 2). However, no significant difference (p&gt; 0.50) was evident in
the proportion of males and females occupying the area between different
winters. Nearly 80 percent of the estimated 200-300 birds annually utilizing
this area were females, while only 20 percent were males. Apparently,
suitable wintering grounds for hens are limited; consequently, considerable
numbers of females are attracted to the few suitable sites.
A significant difference (P&lt;O.005) was detected in the number of adults and
juveniles annually utilizing the wintering area; consistently more adults
than juveniles were present each winter. Numbers of juveniles appeared correlated with nesting success in the previous summer. Thus, a summer of good
production resulted in a high percent of juveniles during the following winter.
In the summer of 1972, 37 (82.2 percent) of 45 females encountered on breeding
areas surrounding Guane11a Pass were accompanied by chicks, while in 1973
only 19 (54.2 percent) of 35 females located were with broods. The estimated
peak of hatch occurred approximately 3 weeks later in 1973 (July 28) than in
1972 (July 5). Similar indications of good nesting success in 1972 and poor
success in 1973 were reported by Braun (1973, 1974) for other areas of the
state, including the Mt. Evans area which lies approximately 6 km east of
Guane11a Pass. Juveniles comprised 42.3 and 27.2 percent of the birds identified at Guane11a Pass during the 1972-73 and 1973-74 winters, respectively
(Table 2). These data support the hypothesis that more juveniles (percent of
total birds) occur on the wintering area following summers of good production
than after summers of poor production.
Some error in sex determination is possible in winter as sex characters are
not visible. However, through measurements of certain wing characteristics
(Braun and Rogers 1971) over 90 percent of the birds can be accurately sexed.
During this study, of 115 birds identified on the wintering area and relocated
in spring or summer when the sex could be determined by plumage differences
alone, only 3 (2.6 percent) had been incorrectly sexed when initially banded

�Table

2.

Sex and age composition

of white-tailed

Winter

N

b

wintering

b

at Guanella

Pass.
=
Males

Females

Juveniles

Adults
a

ptarmigan

b

b

Percent

N

Percent

N

Percent

N

Percent

b

N

1967-68

66

54.5

36

45.5

30

72.7

48

27.3

18

1968-69

43

70.0

30

30.2

13

81.4

35

18.6

8

1969-70

III

78.4

87

21.6

24

78.4

87

21.6

24

1970-71

125

61.6

77

38.4

48

80.8

101

19.2

24

1971-72

142

56.3

80

43.7

62

78.9

112

21.1

30

1972-73

153

57.7

88

42.3

65

77.4

118

22.6

35

1973-74

125

72.8

91

27.2

34

84.0

105

16.0

20

Total
Mean

765

I

I

b

Total

winter

of 1966-67

number

of birds

was excluded
identified.

21.0

79.0

35.5

due to small

sample

159

606

276

489
64.5

aThe

\0
N

size

(34 birds

identified).

�-93-

on the wintering area. Some error also exists in separating age classes.
Age determination is based on the presence (subadult) or absence (adult)
of pigmentation on the ninth primary.
Developing and using this technique,
Braun and Rogers (1971) reported that 98 percent of the ptarmigan handled
were correctly placed in one of the two age classes.
Considering the small
error in aging and sexing techniques, it is unlikely that considerable bias
occurs in Table 2.

Flocking

Behavior

Like many grouse, ptarmigan are gregarious and generally associate in flocks
throughout the year.
Flocks are temporarily fragmented from late April until
mid-July due to territorial activities accompanying reproduction.
Flocking
tendencies are most evident from late October until late April when ptarmigan
are concentrated on winter use sites. During the winters of 1966-67 through
1973-74, 172 individual flocks totaling over 3,165 ptarmigan were observed
on the Guanella Pass study area.
Following Koskimies (1957), two or more grouse were considered a flock.
Ellison (1973) in studying seasonal flock size of spruce grouse (Canachities
canadensis) arbitrarily chose 18 m as the distance two birds had to be within
each other to be considered a flock. He based this distance on visibility
in the forest.
In this study, no specific distance was designated because
of greater visibility in alpine areas and the tendency for ptarmigan to congregate in large, loose flocks with individuals scattered over an area of 50 m2
or more.
The decision as to whether a group of birds comprised a flock was
decided by the observer.
A bias in estimating flock size resulted when all
birds present could not be counted.
This bias was most serious when considerable numbers
30) of birds were dispersed over a large area such that all
could not be seen, or when the birds flushed before all individuals could be
counted.
In such instances, only minimum estimates of numbers of individuals
in flocks were obtained.
Flock sizes recorded during the winters of 1966-67,
1967-68, and 1968-69 were excluded from the comparison of flock sizes between
winters because of small sample sizes « 15 flocks observed per winter).
Since only 2 to 8 flocks per month were observed in anyone
year, sampie sizes
were not large enough to compare differences in monthly flock sizes between
or within years.
Consequently, monthly analysis of flock size was based on
pooled data which included flock size information collected during all winters
from 1966 through 1974.

o-

Winter flock size varied from less than 5 to over 80 birds, but 68.6 percent
of the flocks observed contained between 2 and 25 individuals.
Of 185 grouse
encounters, only 13 (7.1 percent) were of lone birds (Table 3), indicating
the social nature of ptarmigan during the winter.
Mean flock size was not
significantly different (p&gt; 0.05) between winters for the 5 winters of 196974 (Table 4). The 135 flocks observed during these 5 winters averaged 15.5
birds per flock. No signficant difference (P&gt; 0.05) was noted in mean flock
size for each of the 7 winter months, as average flock size varied from 16.0
to 23.8 birds per flock (Table 5). However, flocks tended to be smaller in
October and April, 16.0 and 16.2 birds per flock, respectively.
Both months

�-96-

represent transition periods when birds are either arrlvlng on (October), or
departing from (April) the wintering area. Since birds do not arrive or
depart simultaneously, probably fewer birds were present at these times to
form flo cks •
Arrival of ptarmigan on wintering areas and formation of flocks were correlated with unfavorable weather.
Braun and Schmidt (1971) noted that following
intense snow storms in October, birds appeared along the periphery of known
winter use sites.
A similar relationship between winter flock formation and
adverse weather has been reported for rock ptarmigan (Lagopus mutus) (Watson
1965), red grouse (Lagopus lagopus scoticus) (Watson and Jenkins 1964), black
grouse (Lyrurus tetrix), and capercaillie (Tetrao urogallus) (Koskimies 1957).
Size of winter flocks of white-tailed ptarmigan at Guanella Pass were similar
to that reported for rock ptarmigan (Weeden 1964), willow ptarmigan (Lagppus
lagopus) (Irving et al. 1967), and black grouse (Koskimies 1957), whereas
spruce grouse (Ellison 1973), ruffed grouse (Bonasa umbellus) (Gullion 1970),
and capercaillie (Koskimies 1957) generally form smaller flocks.
Encounters
with lone ptarmigan were also less than reported for ruffed grouse, spruce
grouse and capercaillie.
Percentage of grouse encounters during winter that
were with lone birds was 58 percent for spruce grouse (Ellison 1973),70
percent for ruffed grouse (Gullion 1970), and 30 percent for capercaillie
(Koskimies 1957).
Koskimies (1957) suggested increased population numbers and good nesting
success led to larger flocks, and to a lesser degree, more flocks. Considering the failure to detect a significant difference between years in mean
size of winter flocks observed at Guanella Pass from 1969 through 1974, even
though nesting success varied from year to year, there was no indication
that nesting success influenced winter flock size. However, following the
good production of 1972, 2.37 flocks were observed per day on the wintering
area, while in the winter of 1973-74 following a poor production year, only
1.29 flocks were located per day. These data suggest a possible relationship
between nesting success and number of flocks utilizing the wintering area.
Assuming more birds are present on the wintering area following a good production year, they apparently form more flocks instead of gathering into
larger flocks.
However, this observation is based on only two years of data
which may not be representative.
Mild weather during the winter period did not cause disintegration of winter
flocks as reported by Koskimies (1957) for black grouse and capercaillie.
On the contrary, large flocks appeared to divide into smaller units during
inclement weather.
Apparently, winter flocks were not composed of family
units.
Ptarmigan broods disperse in late September or early October (Braun
1969) prior to arrival on wintering areas. In no instances were females or
any of their chicks that were banded together during the sunnner relocated
in the same winter flock. In addition, considerable interchange of members
occurred between flocks, as the same group of individuals were seldom observed
together on consecutive visits to the wintering area. Although exact numbers
cannot be Cited, many birds banded at Guanella Pass were relocated several
times throughout the winter associated with flocks of various sizes comprised
of different individuals at different locations.
No noticeable aggressive

�-97-

behavior was exhibited towards new members of a flock. A similar observation was reported for willow ptarmigan in Alaska (Irving et al. 1967).
Partial segregation of sexes of white-tailed ptarmigan occur on wintering
areas (Braun and Schmidt 1971). Flocks of mostly males were seldom observed
at Guanella Pass during this study because most banding efforts were concentrated on areas utilized principally by females.
Thus, minimum data were
collected for determining differences in size of winter flocks of males and
females.
Since sex of ptarmigan cannot be distinguished by visual characteristics during the winter, it was impossible to ascertain sex composition of
a flock unless most birds could be captured or individually identified through
observation of bands.
Flocks of primarily females observed at Guanella Pass
varied considerably in size (2-80 birds per flock) but usually contained 20
to 30 birds.
Seven flocks of males were observed on the Horseshoe Basin
wintering area, with the largest flock being 14 birds.
Only 4 flocks of
primarily males were encountered at Guanella Pass during this study, with
the largest flock consisting of 9 individuals.
It should also be noted that
of 13 lone birds found, 8 (61.5 percent) were males, 3 (23 percent) were
females, and 2 (15.5 percent) were of unknown sex. These data suggest that
males do not gather in large flocks typical of wintering females.
Similar
results were reported by Braun and Schmidt (1971), who observed that flocks
of males were usually small « 15 birds), while females congregated in
larger flocks.
Differences in flock sizes of males and females may possibly
be related to differences in the winter habitat utilized.
Females typically
winter in areas where dense, tall stands of willow occur; thus, food supplies
are abundant and they can afford to congregate in large groups. Conversely,
males winter at higher elevations where willow is sparsely distributed and
food availability is primarily dependent upon wind action.
Such areas are
limited in size and scattered in distribution with each site possibly capable
of supporting only a small group of birds.

Distribution
Locations of 172 winter flocks, plus 13 lone birds observed at Guanella Pass
since 1966 were plotted on both a monthly and yearly basis in order to
determine if there were changes in distribution between months or years.
Examination of these data indicated that: (1) essentially all sites on the
wintering area were utilized, (2) some sites were highly preferred to others,
(3) there were no apparent differences in distribution of birds on the
wintering area between years, and (4) changes in monthly distribution occurred.
Since the buds and twigs of willow provide the bulk of the food eaten by
ptarmigan during the winter (May and Braun 1973), flocks were seldom found
where willow was absent.
Changes in monthly distribution appeared related to
changes in the availability of willow as influenced by the amount of snow
present (Braun and Schmidt 1971). Sites used by ptarmigan early in the winte.r
were devoid of birds later, primarily because increasing snow depths covered
the willow. The birds moved to other sites where tall stands of willow occurred
and snow rarely covered all the bushes.
As spring approached, the snow
settled, exposing unused buds, and birds were again observed on areas previously
abandoned.

�-100-

o

0
I

PERCENT RETURNING
CD
(Jl
m ~
(JI
~
N

o

o

o

o

0

I

I

I

I

I

o

--

TOTAL

I~

FEMALES

l~

I

I

I~
- IOJ-

ADULTS

-

l~

I~

ADULT
FEMALES
SUBADULT
FEMALES
ADULT
MALES
SUBADULT
MALES

I

I

o
o

-

MALES

SUBADULTS

o

w
o

g

-

I~

I§

-

]~

Fig. 4. Percentage of different age and sex classes of white-tailed
ptarmigan returning to Guanella Pass (excluding known losses).
Sample
sizes are given in paren theses.

�-101-

Subadults suffer a higher mortality than adults (Braun 1969); consequently,
few are available to return.
In addition, subadults have no prior attachment to breeding sites and upon leaving wintering areas in spring they
frequently travel considerable distances in search of vacant territories
where they can establish.
Because territorial males generally winter adjacent
to their territories (Braun and Rogers 1971), only subadult males becoming
established on territories in the vicinity of Guanella Pass should return.
Those becoming established farther away probably winter adjacent to their
territories and thus, do not return.
Documented movements of subadult males
indicate they frequently move over 3 km to a breeding site. Movements of
adult males to breeding areas seldom exceed this distance.
Thus, because
most adult males banded at Guanella Pass were established on territories in
the viCinity, they returned year after year. Only subadult males becoming
successfully established on territories within 3 km of Guanella Pass returned
in succeeding winters.
All but one failing to return were located over 4 km
from Guanella Pass and probably wintered elsewhere.
The situation with females is similar to that of males except females
returned from greater distances.
While most s.ubadult males moving over 4 km
away from Guanella Pass failed to return, subadult females returned from at
least 10 km away. This supports the hypothesis that areas capable of supporting females during the winter period are limited.
Of the 15 subadult
females banded at Guanella Pass that were located on breeding or sunnnering
areas and that failed to return, 10 (67 percent) were located over 8 km from
Guanella Pass. Only 5 (29 percent) of 17 returning subadult females located
on breeding or summering areas moved greater than 8 km. This suggests that
most subadult females traveling beyond a certain distance (8 km) in search
of unmated territorial males generally do not return.
Undoubtedly, some die,
while others probably winter closer to their territories.
Since movements
of adult females to breeding sites rarely exceeded 8 km, they showed a greater
tendency to return.
Sufficient time was not available to intensively investigate other winter
use sites; thus, limited evidence was obtained to support the hypothesis that
some birds (primarily subadults) initially banded at Guanella Pass were
utilizing other sites during the winter period.
Of 40 birds identified on
other winter use sites during 6 field days, 4 (10 percent) were initially
banded at Guanella Pass. All were banded as subadults with 2 males being
located at Naylor Lake, 1 female at Waldorf, while 1 female was observed at
Steven's Gulch. Although the sample is small, it does indicate that ptarmigan
banded as subadults are more likely to disperse from wintering areas where
they were initially banded and subsequently winter elsewhere.
No differences were apparent in the proportion of adult males (75 percent) and
females (73 percent) returning to Guanella Pass.
Since wintering sites utilized by males were not searched as thoroughly as areas used by females the
proportion of males returning may possibly be underestimated.
Braun (1969)
reported a 25 percent annual turnover of adult males in an unhunted population of ptarmigan.
This value is identical to the percent of adult males
(25 percent) failing LO return to the wintering site.
For the same unhunted
population, Braun reported a 30 percent annual turnover for adult females.

�-102-

This percentage is almost identical to the proportion of adult females
(27 percent) failing to return to Guanella Pass. Since adults show a high
fidelity to wintering sites, it can be assumed that those not returning
were probably dead, consequently, percent of adults returning approximates
annual survival rates for this population. No similar assumption can be
made for subadults because non-returning subadults are not necessarily lost
from the population.
MOVEMENTS
Movements of Chicks to Wintering and Breeding Areas
One hundred and twenty six chicks were individually marked on the MontezumaGeorgetown study area during the summers of 1972 (75) and 1973 (51). Chicks
were banded from less than 1 to over 18 km from Guanella Pass. The sample
included 47 males, 47 females, and 32 birds of unknown sex. In order to
estimate percentage of birds initially banded as chicks that were recovered
on wintering and breeding sites, 16 birds (7 males, 5 females, and 4 of
unknown sex) were excluded from the sample since they were harvested before
having an opportunity to move to a wintering area. All were shot within 2 km
of their initial banding site, indicating that broods are relatively sedentary
during summer and early fall prior to brood dispersal in late September and
October. Braun (1969) similarly noted that females and their broods remained
in the same general area throughout the summer.
Of 99 subadult ptarmigan identified at Guanella Pass during the winters of
1972-74, only 2 (both females) were initially banded as chicks during this
study. Additionally, none of the 12 subadults identified on other winter use
sites was banded as a chick. Based on 110 chicks (excluding losses) banded
on the Montezuma-Georgetown study area, movements to wintering areas were
documented for only 1.8 percent (2 of 110) of the birds. Both individuals
identified at Guanella Pass were banded on brooding areas in 1973 and recovered the following winter (1973-74). Distances and directions traveled from
original banding location to Guanella Pass were 10.7 km east-northeast and
9.9 km northeast.
An additional 2 birds (one male and one female) banded as chicks on Mt. Evans
in 1971 were known to have moved 5.6 km west-northwest and 6.4 km west, respectively, to Guanella Pass. Studies by Braun (1969) indicate that following
brood dispersal chicks may wander several kilometers from their natal areas
in a few days. Probably, the two chicks banded at Mt. Evans in October 1971
hatched elsewhere; thus, their movements to wintering sites were not comparable with the movements of the two chicks banded and recovered in this study.
Both chicks banded at Mt. Evans in 1971 were not relocated at Guanella Pass
until the winter of 1972-73. Where they spent the winter of 1971-72 is unknown.
Only four (all males) of the 70 chicks (excluding 5 losses) marked on the
study area in 1972 were relocated on breeding sites in 1973. Distances
traveled from their initial summer banding location to a territory were 0.9
km, 0.7 km, 0.7 km, and 1.2 km. Braun (1969) reported that males banded as

�-103-

chicks were not observed during the next breeding season more than 1.0 km
from where originally banded (N = 26). Thus, males show high fidelity to
natal areas. Probably some males banded as chicks which are unsuccessful in
competing for territories near their natal areas search elsewhere for unoccupied areas where they can become established. Others may remain near natal
areas as non-territorial males occupying areas on the periphery of established
territories.
Since no females banded as chicks were relocated on territories, any statements about their movements to breeding sites are speculative. Braun (1969)
found that females banded as chicks traveled considerable distances from their
natal areas. Of the two females banded as chicks that Braun relocated in
subsequent years, one was found as a transient 9.6 km from where originally
banded, while the other was established on a territory 19.3 km from the initial
banding site.
The high mortality rate of ptarmigan during their first year of life is undoubtedly a major factor responsible for the low reobservation rate of banded
chicks. Nearly 70 percent of the chicks hatched do not survive their first
year (Braun and Rogers 1971). Additional factors possibly contribute to the
low rate of reobservations of marked chicks. Most juvenile males are unsuccenssful in competing for territories and consequently, cannot be located
using tape recorded calls. Obviously, chicks are moving to Guanella Pass as
indicated by the 99 subadults identified during this study. This suggests
that chicks hatched outside the study area were moving to the wintering area.
It is also possible that many broods within the study area went undetected.
Thus, the banded sample may only represent a small proportion of the chicks
available to move to wintering sites. Finally, and probably most important,
insufficient numbers of chicks were banded to offset their high mortality rate.
Increased surveys in other wintering sites, extending the area searched
during the breeding and brood rearing periods, and larger samples of banded
chicks may be necessary if chick movements are to be documented.
Because findings from the present study are based on small samples, movements
of white-tailed ptarmigan chicks are still basically unknown. Present evi~
dence indicates that females accompanied by chicks remain within a limited
area during summer and early fall. Documented mortalities of ptarmigan
suggests that most losses occurring in the fall are young of the year (Braun
and Rogers 1971). This mortality can be attributed to increased mobility of
juveniles following brood breakup. Bendell and Elliott (1967), Gullion and
Marshall (1968), Zwickel et al. (1968), and Robel et al. (1970) all reported
similar increased movements of juvenile grouse in fall, and interpreted this
increased mobility as dispersal. Possibly, competition for space on wintering
areas results in some juveniles becoming established, while others continue
their nomadic life and wander between wintering sites. Inexperienced females
(young of the year) travel greater distances than inexperienced males to
winter use sites. It is possible that wintering areas suitable for females
are limited, forcing females to move longer distances. Upon departing from
wintering areas in spring, most juvenile males return to their natal areas
to compete for breeding territory. Juvenile fe1i1alesmove greater distances
than juvenile males to breeding sites, and do not return to their natal
areas.

�-104-

Movements From Wintering to Breeding or Summering Sites
Prior to 1972, 351 different ptarmigan were marked during the winter at
Guanella Pass. Originally, hunter recoveries were expected to provide
enough data to document movements of white-tailed ptarmigan from wintering
areas. However, due to the inaccessibility of ptarmigan habitat and lack of
hunter participation, only 11 band recoveries were obtained from 1967 through
1971. As a result, the present study was initiated in 1972. From 1972 to
1974, 278 different ptarmigan were individually identified at Guanella Pass.
An additional 40 birds were marked on other winter use sites in the MontezumaGeorgetown study area, including 12 at Naylor Lake, 10 at Waldorf, 12 at
Horseshoe Basin, and 6 at Steven's Gulch.
Ninety nine birds were known to move from wintering to breeding or summering
sites; 28 adult females, 39 subadult females, 14 adult males, and 18 subadult
males. This sample includes eight of 11 movements recorded prior to 1972.
Three of the 11 movements were excluded because of incomplete records. Ten
movements were obtained from hunter recoveries during the 1972 (4) and 1973
(6) grouse seasons. The remaining 81 movements were obtained by locating
banded ptarmigan during the spring and summer through the use of tape recorded
calls (Braun et ale 1973). Forty six and 35 movements were documented by
this method in 1972 and 1973, respectively. Compared to the number of birds
identified on wintering areas, few were actually relocated on breeding or
summering sites. Undoubtedly, some birds died or emigrated from the area,
but the difficulty of thoroughly searching all breeding and summering sites
within the study area was the primary limitation. Probably many banded birds
went undetected, especially marked grouse occupying territories outside the
Montezuma-Georgetown study area.
Most ptarmigan (90) for which movements were documented were initially banded
at Guanella Pass, however, 3 were banded at Mt. Evans, 3 at Horseshoe Basin,
2 at Waldorf, and 1 at Steven's Gulch. Distances traveled by birds banded on
all winter use sites were similar. All data were analyzed to identify degree
of variation in movements between years. Differences were tested between
movements from wintering to breeding areas and wintering to summering areas.
No statistically significant differences (P &gt;0.05) were evident in comparisons between years for grouse of the same age and sex. Consequently, data
for all years were combined.
Differences between each age and sex class in distances traveled were tested
through standard statistical analyses. These data are given in Table 7, and
graphically presented in Fig. 5. Differences between means were tested by
student "t" tests for adult females vs. subadult females, adult males vs.
subadult males, adult females vs. adult males, subadult females vs. subadult
males, adult females vs. subadult males, and subadult females vs. adult males.
Results of these tests indicate that (1) females of both age classes moved
significantly greater distances (P&lt;O.Ol) than males, (2) no significant
difference (P&gt; 0.05) was detected in distances traveled between adult and
subadult females, and (3) no significant difference (P&gt; 0.05) was evident in
distances traveled between the age classes of males.

�-105-

30-

25-

-

en

0::
W

~ 20~

o
-l

~

z 15•••••

z

w
~ lOw

&gt;
o
::E

..,,..

-H-t-(39)
(28) -+-t-t-

5-

-I'"

-++-+-(18)

--

Fig. 5. Distances traveled by white-tailed ptarmigan from wintering to
breeding or summering areas. Vertical line = range, horizontal line =
mean, vertical bar = standard deviation.
Sample sizes are given in
parentheses.

�-106-

On the average, subadults moved greater distances than adults of the same
sex. The failure to detect a significant difference in distances traveled
between age classes of the same sex was apparently the result of small
samples for adults and because several adults included in these samples made
unusually long movements. For example, the sample of adult males was small
(14 birds) and included two birds moving 7.9 and 5.9 km to breeding sites.
The other 12 adult males each moved less than 2.5 km. In contrast, 9 of 18
subadult males moved over 4 km with the longest movement exceeding 10 km.
Similar findings were noted for females. While most adult hens (19) did not
venture beyond 8 km from the wintering area, 5 moved over 10 km. Movements over
10 km were not uncommon for subadu1t females, as 15 of 3~ movements exceeded
this distance. Thus, although some adults and subadults made extensive moves,
subadults of both sexes moved further than their adult counterparts.

Table 7. Contrasting statistical analyses for distances moved by different
age and sex classes of white-tailed ptarmigan.

Males

Adult
Females

Males

Subadu1t
Females

Distances Traveled
Mean (km)

2.3

6.3

3.8

8.5

Range (km)

.6-7.9

1.6-15.7

.2-10.8

.8-22.7

Standard deviation

2.1

4.0

2.9

6.0

Sample size

14

28

18

39

Birds departing from Guanella Pass showed a striking tendency to move westerly,
especially northwest (Fig. 6). This tendency was mainly noted for females of
both age classes and subadult males. Adult males did not prefer to move in
one direction more than another. The longest movements were also in the
westerly directions, being longest towards the northwest (Fig. 6). Of 90 total
movements documented from Guanella Pass, 67 (74.4 percent) were ~rom southsouthwest to north-northwest with 25 (27.8 percent) to the northwest. The
average distance traveled by grouse moving northwest was 10.1 km. This distance was significantly greater than the mean distance traveled in any other
direction. Additionally, more subadults (19) than adults (6) moved northwest.
No birds were recorded moving east and apparently only 4 birds moved in any
other easterly direction. The longest movements in easterly directions were

�-107-

..

Q
..,CD
ZI)(

'-"

Fig. 6. Directions and distances recorded for movements of white-tailed
ptarmigan from Guanella Pass to breeding or summering sites. Each concentric
circle represent one kilometer. The number of birds moving in each direction
(N) and the mean distance traveled (X) are shown in parentheses. Movements
are plotted along the nearest directional heading.

�-108-

by males (2 adults and 2 subadults), while females of both age classes
exhibited minimal movements in these directions.
The 4 males were recover.ed
on the Mt. Evans study area which lies about 6 km east of Guanella Pass.
Ptarmigan investigations have been conducted at Mt. Evans since 1966, and
no females banded at Guanella Pass have been recovered there.
Reasons for the observed differences in movement patterns between age classes
of the same sex and between sex classes are not definitely known. However,
using data collected during this and prior studies (Braun 1969, Braun and
Schmidt 1971, Braun and Rogers 1971, and Schmidt 1969), the following explanation is presented.
Nearly 80 percent of the 200-300 birds annually utilizing Guanella Pass are
females and sites utilized by females during the winter are generally farther
from breeding areas than locations where males winter.
In addition, although
wintering females were located in three other sites within the MontezumaGeorgetown study area the only major concentration was at Guanella Pass.
At breeding densities of 4 females per km2, the 160 to 240 females wintering
at Guanella Pass must disperse into 40 to 60 km2 of surrounding breeding
habitat.
It would appear then that suitable wintering grounds for females
are limited, consequently, females are attracted from considerable distances
to suitable sites.
Since adult ptarmigan have great fidelity to breeding and
wintering sites used in prior years, it follows that movements from wintering
to breeding areas must be similar to those from breeding and/or summering
sites to wintering areas.
Subadult females have no prior attachment to
breeding areas and must search for territories occupied by unmated males.
Some subadult females are able to establish themselves close to the area where
they initially wintered, while others must travel considerable distances to
locate unmated territorial males. However, this factor alone cannot account
for the almost complete failure to relocate females that were banded as chicks
on brood areas.
Most adult males winter adjacent to breeding areas and only have to move short
distances to these areas in the spring.
Subadult males must compete with
established adults for space on their natal areas. Only a few are successful
in becoming established in their first year, while most remain on peripheral
areas as non-territorial males.
Undoubtedly, some subadult males unsuccessful
in competing for space on their natal areas search elsewhere for unoccupied
territories.
These birds probably account for the exceptionally long movements recorded for some subadult males.
Most alpine habitat and consequently, most breeding habitat occurs west to
northwest of Guanella Pass (Fig. 1). Birds could travel about 32 km northwest across continuous alpine habitat, while the maximum distance they could
travel in any other direction without encountering extensive areas of unsuitable habitat, mainly forest, was 18 km. Considering only the easterly directions, the maximum distance ptarmigan could venture from Guanella Pass was
14 km. The greater probability of finding vacant territories in the direction
where most breeding habitat occurs may explain why subadults preferred to
move northwest.

�-109-

As most adult males banded at Guane11a Pass occupied territories immediately surrounding the wintering grounds, it is not unusual that they showed
no significant movements in anyone direction. At present, no evidence is
available to explain why females avoided moving longer distances in easterly
directions. From studies conducted at Mt. Evans, there is no indication that
lack of vacant territories or quality of the breeding habitat controlled
easterly movements of females. Perhaps females breeding on areas east of
Guane11a Pass winter elsewhere. While evidence to support this hypothesis is
not presently available, it is believed that many of these females winter in
a large willow basin approximately 6 km south of Mt. Evans. Unfortunately,
this basin was inaccessible during the winter, and consequently, could not
be investigated.
Movements from Breeding or Summering to Wintering Sites
Although the major emphasis of this study was to document movements from
winter use sites, 45 movements were documented to Guane11a Pass. This sample
included 16 adult females, 17 subadu1t females, 6 adult males, and 6 subadu1t
males. Mean distances traveled by these birds to Guane11a Pass were 6.8, 6.5,
1.4, and 3.1 km, respectively. In all cases, movements to winter use sites
were shorter than movements from these sites, but differences were not statistically significant (P&gt; 0.05). From.data presented earlier it is apparent
that most females moving over 10 km and males moving over 4 km failed to
return to the wintering site where they were banded. This can be partially
attributed to mortality during the spring and summer, but it was noted that
some birds (primarily subadu1ts) traveling long distances from the wintering
site where they were initially banded utilized other areas closer to their
territories in subsequent winters. This may explain why return movements
were shorter.
Almost identical patterns were noted for movements of ptarmigan to wintering
areas as described for movements from these sites. Since the majority of
adult males banded at Guane11a Pass were established as breeding birds in the
immediate vicinity, they showed the shortest movements to the wintering area.
Experienced (adults) and inexperienced (subadu1ts) females moved longer distances than males to Guane11a Pass. Differences in wintering habitat previously described were probably major factors affecting hen movements to
winter use sites. As expected,birds moved towards Guane11a Pass mainly from
breeding and summering sites west to northwest of the wintering area.
However, returning movements were not as prevalent from areas to the northwest as were movements in this direction. While 25 birds moved northwest
from Guane11a Pass, only 12 were known to return. Considering that mainly
subadu1ts moved northwest and the extensive distances they traveled in this
direction, it is probable that many of these birds wintered elsewhere. Even
so, the longest returning movements ( &gt; 15 km) were by subadult females
originally moving northwest from Guane11a Pass.
Movement patterns noted in this study are similar to those reported by Braun
(1969). Similar findings relating to the greater mobility of females have
also been preported for rock (Weeden 1964) and willow ptarmigan (Irving et a1.
1967). Both studies found that most males winter near their breeding grounds,

�-110-

while females frequently moved long distances to or below treeline to winter
ranges. As suggested by Weeden (1964) migration patterns and segregation
of sexes of rock and willow ptarmigan during winter appear to be closely
related. This relationship is also true for white-tailed ptarmigan. No
evidence is available to explain why sexes need to live separately during
the winter. Perhaps this is an adaptation to allow for more efficient utilization of winter food supplies.
The greater mobility of females is not unique to ptarmigan as demonstrated
from studies of other species of tetraonids including: sharp-tailed grouse
(Pedioecetes phasianellus)(Hillman and Jackson 1973); ruffed grouse (Hale
and Dorney 1963, and Gullion 1970); greater prairie chicken (Typanuchus cupido
pinnatus) (Hamerstrom and Hamerstrom 1973); black grouse, and capercaillie
(Koskimies 1957). Compared to many species of grouse, white-tailed ptarmigan
do not move unusually long distances. When compared to documented movements
of rock (Weeden 1964) and willow ptarmigan (Irving et al. 1967), movements
of white-tailed ptarmigan are relatively short. Thus, ptarmigan are migratory with white-tailed ptarmigan exhibiting the shortest annual migrations
documented to date.

SUMMARY
Intensive studies of characteristics and migration of wintering populations
of white-tailed ptarmigan were conducted at Guanella Pass and surrounding
alpine areas in north-central Colorado. Data were collected from April 1972
to April 1974.
Arrival and departure from and to winter use sites were greatly influenced
by climatic conditions. However, in most years, ptarmigan were on wintering
areas by late October and remained on these areas until late April when they
departed for their breeding grounds.
Seven hundred and ninety nine ptarmigan were identified at Guanella Pass.
Nearly 80 percent of the birds handled each year were females. Consistently
more adults than subadults utilized the area; however, the age composition
varied considerably between winters. Proportionally, more subadults were
identified on the wintering area following summers of good production (1972)
than poor production (1973).
One hundred and seventy two flocks were observed at Guanella Pass. Flock
sizes ranged from 2 to over 80 birds, but 69 percent of the flocks encountered
contained 2 to 25 members. Mean yearly flock sizes ranged from 12 to 21 birds,
but differences were not significant. Flocks tended to be smaller in October
and April (about 16 birds per flock); however, mean monthly flock sizes were
not significantly different. Flocks were not composed of family units nor
were the same individuals associated together in the same flock throughout
the winter. Following a summer of good production, ptarmigan apparently
formed more winter flocks instead of gathering into larger groups.
Partial segregation of sexes was observed, with males preferring higher, more
windswept ridges than females. Flocks of males were usually small « 15 birds)
while females congregated in flocks of larger size (20-30 birds).

�-111-

Sixty percent of the ptarmigan banded at Guanella Pass and relocated on
breeding or summering sites returned in subsequent winters.
Seventy three
percent of the adults returned, while only 51 percent of the subadults
returned.
Differences were due to greater mobility and higher mortality of
subadults.
Most adults failing to return were probably dead, while some
subadults wintered elsewhere in winters following their banding at Guanella
Pass.
Presence and abundance of willow were the most important factors determining
where ptarmigan wintered.
Essentially all sites on the Guanella Pass wintering area were utilized, but some sites were highly preferred to others.
Yearly differences in distribution were not evident.
Monthly differences in
distribution resulting from changes in availability of willow and snow quality
were detected.
Human disturbance had some influence upon the distribution of
birds at Guanella Pass.
Females and their broods moved relatively short distances in summer and early
fall prior to brood breakup.
Of 126 chicks banded during this study, only
2 were located on winter use sites.
Four other chicks were located on breeding
areas .the year following banding.
This low reobservation rate was primarily
the results of high mortality of chicks.
Females banded as chicks traveled
farther than males.
Males showed a high fidelity to natal areas, with no
males banded as chicks relocated on breeding sites more than 1.2 km from
their initial banding location.
Ninety nine movements were documented from wintering to breeding or summering
sites.
On the average, subadu1ts moved greater distances than adults of the
same sex. Females were consistently more mobile than males.
Females wintered
farther from breeding sites; possibly because wintering areas suitable for
females were limited.
Adult and subadult males were relatively sedentary,
usually moving less than 2.5 to 4.0 km, respectively.
Average movements of
females were 6.3 km for adults and 8.5 km for subadults, with the longest
movement being 22.7 km for a subadu1t female. Birds departing from Guanella
Pass showed a striking tendency to move in the westerly
directions, especially
northwest.
This tendency was probably related to the large amount of alpine
habitat located west to northwest of the wintering area.
Forty five movements were documented to Guane11a Pass.
Movements to winter
use sites were shorter than movements from these sites; however, movement
patterns were similar.
Most birds moved towards Guane11a Pass from breeding
or summering sites west to northwest of the wintering area.

�-113-

LITERATURE CITED

Aldrich, J. W. 1963. Geographic orientation of American
onidae. J. Wildl. Manage. 27(4):529-545.
Bailey, A. M. 1927. Notes on the birds of southeastern
Auk 44(2): 184-2 05.

Tetra-

Alaska.

Bendell. J. F •• and P. W. Elliott.
1967. Behavior and the regulation of numbers in blue grouse.
Canadian Wildl. Ser v . Rept.
Ser. 4. 76pp.
Bent, A. C.
birds.

1932. Life histories of North American
U. S. Nat. Mus. Bull. 162. 490pp.

Braun. ·C. E. 1969. Population dynamics, habitat,
of white-tailed ptarmigan in- Color ado . Ph.D.
rado State Univ., Fort Collins.
189pp.

gallinaceous

and movements
Thesis.
Colo-

1972. Habitat requirements of Colorado white -tailed
ptarmigan.
Proc. Western Assoc. State Game and Fish Commissioners 51:284-292.
1973. Continued inventory of. selected ptarmigan populations.
Colorado D'iv, Wildl. Game Res. Rept., Fed. Aid P'r oj .
W-37-R. April.
(In press)
1974. Continued inventory of selected ptarmigan populations.
Colorado Di.v, Wildl. Game Res. Rept., Fed. Aid Proj.
W-37-R. April. (In press)
___

-.
and G. E. Rogers.
1971. The white-tailed ptarmigan in
Colorado.
Colorado D'iv, Game, Fish and Parks Tech. Publ.
27. 80pp.
and R. K. Schmidt, Jr.
1971. Effects of snow and wind
on wintering populations of white-tailed ptarmigan in Colorado.
Pp.238-250.
In A. O. Haugen, ed. Proceedings snow and ice
symposium.
Iowa Cooperative Wildl. Res. Unit, Iowa State
Univ .• Ames. 280pp.

�-114-

Braun,

C. E., R. K. Schmidt, and G. E. Rogers.
1973. Census of
Colorado white -tailed ptarmigan with tape recorded calls.
J.
Wildl. Manage. 37( 1):90-93.

Choate, T. S. 1963. Ecology and population dynamics of whitetailed ptarmigan (Lagopus leucurus) in Glacier National Park,
Montana. Ph.D. Thesis.
Univ. Montana, Missoula.
205pp.
Ellison, L. N. 1973. Seasonal social organization
of spruce grouse.
Condor 75(4):375-385.

and movements

Gabrielson, r. N., and F. C. Lincoln.
1959. The birds of Alaska.
The Stackpole Co. and Wildl. Manage. Inst., Harrisburg,
Fa.
and Washington, D. C. 922pp.
Gullion, G. W. 1970.
Univ. Minnesota,
(m.ultilith) .
____

The ruffed grouse in Northern Minnesota.
Forest Wildl. Relations Proj.
37pp.

, and W. H. Marshall.
1968. Survival of ruffed grouse in
a boreal forest.
The Living Bird 7:117-167.

Hale, J. B., and R. S. Dorney.
1963. Seasonal movements of
ruffed grouse in Wisconsin.
J. Wildl. Manage. 27(4):648-656.
Hamerstrom,
F., and F. Hamerstrom.
1973. The prairie chicken
in Wisconsin.
Wisconsin Dept. Nat. Res. Tech. Bull. 64.
52pp.
Haskins, A. G. 1969. Endoparasites of white-tailed ptarmigan
(Lagopus leucurus) from Colorado. M.S. Thesis.
Colorado
State Univ., Fort Collins.
147pp.
Hillman, C. N., and W. W. Jackson.
1973.· The sharp-tailed grouse
in South Dakota. South Dakota Dept. Game, Fish and Parks
Tech. Pub!. 3. 64pp.
Irving, L., G. C. West, L. J. Peyton, and S. Paneak.
1967. Migralion of willow ptarmigan in arctic Alaska. Arctic 20(2 ):77-85.
Judson, A. 1965. The weather and climate of a high mountain pass
in the Colorado Rockies. U. S. Forest Servo Res. Paper
RM-16. 28pp.

�-115-

Koskimies, J. 1957. Flocking behavior in capercaillie,
Tetrao
urogallus (L.), and blackgame, Lyrurus tetrix (L.).
Finnish
Papers on Game Res. 18:1-32.
Marr,

J. W. 1961. Ecosystems of the east slope of the Front Range
in Colorado. Univ. Colorado Studies. Sere BioI. 8. 134pp.

May, T. A. 1970. Seasonal foods of white-tailed ptarmigan in
Colorado.
M.S. Thesis.
Colorado State Univ., Fort Collins.
55pp.
1974. Bioenergetics of white-tailed ptarmigan in Colorado. Ph.D. Thesis.
Univ. Colorado, Boulder.
(In prep.)
, and C. E. Braun.
1973. Seasonal foods of adult white-----.tailed ptarmigan in Colorado. J. Wildi. Manage. 36(4}:11801186.
Packard, F. M. 1945. The birds of Rocky Mountain National Park,
Colorado. Auk 62(3 ):371-394.
Racey

K. 1948. Birds of the Alta Lake region,
Auk 65(4}:383-401.
I

British

Columbia.

Robel, R. J., J. N. Briggs, J. J. Cebula, N. J. Silvy, C. E. Viers,
and P. G•.Watt. 1970. Greater prairie chicken range, movements and habitat usage in Kansas.
J. Wildl. Manage. 34(2}:
286-306.
Schmidt, R. K., Jr.
1969. Behavior of white-tailed ptarmigan in
Colorado. M.S. Thesis.
Colorado State Univ., Fort Collins.
174pp.
Simpson, G. G., A. Roe, andR. C. Lewontin. 1960. Quantitative
zoology. Revised Ed., Harcourt, Brace and Co . , New York.
440pp.
Watson, A. 1965. A population study of ptarmigan
in Scotland. J. Anim. Ecoi. 34:135-172.
____

(Lagopus mutus)

"_, and D. Jenkins.
1964. Notes on the behavior
grouse.
British Birds 57(4):137-170.

Weeden, R. B.
ptarmigan

of red

1964. Spatial separation of sexes in rock and willow
in winter.
Auk 81(4):534-541.

�-116-

Weimer,
R. J., and J. D. Haunt eds.
1960.
Guide.to the geology of
Colorado.
Ge o l , Soc. Ame r, , Rocky Mountain Assoc.
Geol.,
ColoradoSci.
Soc., Denver.
310pp.
Zwickel,
F.,
grouse.

and

J. F. Bendel!.

J. Wildl.

Manage.

1967.
A snare
31(1}:202-204.

for capturing

blue

, 1. O. Buss, and J. H. Brigham.
1968.
Autumn move---ments of blue grouse and their relevance
to populations
and
J. Wildl.

management.

Prepared
.

L«~

by
I

Richard

Manage.

dfItiFffL-

W.'

offman

32(3):456-468.

�-117-

April 1974

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~-----------

Project No.

W-37-R-27

Work Plan No.

Game Bird Survey

20

Job No.
1
Investigation of Population
Status of Bobwhite Quail in Eastern Colorado

----------~~-------------------

Job Title

Period Covered: April 1, 1973 to March 31, 1974
Personnel:

Donald Hoffman, Marvin Gardner, Earl Downer, Mike Loftsgard,
Ted Behlke and Warren Snyder.

ABSTRACT
Summer whistling counts on the Tamarack Study Area and elsewhere along the
South Platte and Republican River drainages generally dropped from 1972
levels. Change in ratio projections indicated that bobwhite quail suffered
an approxiinate 60 percent natural mortality from fall banding, 1972 to February
1973 on the Tamarack. This loss, in combination with harvest removal of 26.7
percent accounted for an 86.8 percent late fall and winter lIlOrtality rate.
Only six percent of the original 2,319 quail were lost through the subsequent
spring and summer of 1973, bringing the annual fall to fall mortality rate to
93.7 percent on the Tamarack. Severe cold and prolonged snow through the 1972
hunting season was responsible for the except tonal.Lyhigh winter loss. A
major flood inundated the South Platte VClll,~y
and the study area for several
weeks during the spring of 1973. Subsequent vegetative growth and weather
conditions permitted nesting which was delayed by about two weeks in comparison with the previous year. The 1973 fall population estimate of 722 quail
was 254 percent greater than the late winter, 1973 population. In comparison, the 1972over-sunnner gain was 235 percent above the late winter base.
Hunters removed.about 31.7 percent of the quail, including crippling loss,
during the 33 day season in 1973. Six to ten inches of snow cover and extreme
cold starting December 18, 1973 and lasting for almost a month, caused high
quail loss similar to that of the previous winter. Winter, 1974 trapping
efforts were near futile. It is estimated that only about 100 quail survived
the winter to reproduce in 1974.

�-118-

RECOMMENDATIONS
1.

Whistling count indices should be continued on the Tamarack to determine
their relationship to preceding late winter and subsequent fall population and harvest levels. It is recommended that Wildlife Conservation
Officers continue inventory counts on routes established elsewhere in
eastern Colorado.

2~

Quail hunting seasons should cOIllIIlence
in early November and run
continuously to termination by early to mid-December at the latest.
A split or delayed second season for bobwhite is not recommended.

3.

Phragmites stand locations along the Tamarack should be plotted. The
merits and problems connected with uSe of this winter cover for all
wildlife should be weighed •. Transplants of phragmites into new locations
would potentially follow.

�-119-

INVESTIGATION OF POPULATION
STATUS OF BOBWHITE QUAIL IN EASTERN COLORADO
Warren D. Snyder

P. S. OBJECTIVE
To investigate (1) the distribution and relative density of bobwhite quail
in eastern Colorado, and (2) population structure and level, and rate of
harvest, and identify key factors that limit bobwhite on the Tamarack Management Area.

SEGMENT OBJECTIVES
1.

To investigate the distribution and density of bobwhite in eastern
Colorado.

2.

To investigate population structure and level, and rate of harvest of
bobwhite quail on the Tamarack Management Area.

3.

To identify key factors limiting bobwhite on the Tamarack Management
Area.

METHODS AND MATERIALS
Methods and materials used in this investigation were previously listed by
Snyder (1972, 1973) in previous segment reports.
A brief clarification is presented here as to analytical procedures used
in the section on Population Structure, Level and Rate of Harvest. To
increase sample sizes, these data utilize fall retrapped adults to supplement fall banded samples. For example in the fall of 1972, 802 quail were
banded and 34 previously banded adults were retrapped. The combined total,
836 quail, were used as the fall sample on which to base population level
estimates, and other variables. However, subsequent recoveries of other
previously banded quail, not retrapped in the fall, were not utilized in
these data analyses.
RESULTS AND DISCUSSION
Bobwhite Distribution and Density in Eastern Colorado
South Platte River
Count summaries are provided in Tables 1 and 3. With the exception of
Atwood-Merino and Sterling-Iliff routes, the counts were generally down.
The greatest decline was noted in the Brush-Fort Morgan area. Eleven counts
on the Tamarack indicated about half ~s many calling males as in 1972. Note
the overall summary at the bottom of Table 1.

�-120-

Table 1. A comparison of 1971, 1972 and 1973 bobwhite quail whistling
counts completed along the south Platte River.

Number
of
Counts

Number
of
Stops

2

Natural
Tally

Mean

20
20
40

78
65
93

3.90
3.25
2.33

2
4'
6

20
45
64

67
202
135

3.35
4.49
2.10

1971
1972
1973

6
7
5

75
79
57

219
312
109

2.92
3.95
1.91

Sterling-Iliff

.1972
1973

2
2

19
20

63
73

3.33
3.65

Atwood-Merino

1971
1972
1973

2
2
3

20
20
30

108
182
246

5.40
9.16
8.20

Brush-Fort Morgan

1971
1972
1973

2
2
3

20
20
30

77
163
60

3.85
8.15
2.00

Goodrich-Orchard

1971
1972
1973

2
2
1

20
23
11

23
32
7

1.15
1.39
0.64

South Platte River
Average for all
counts

1971
1972
1973

175
226
252

572
1,019
723

3.27
4.51
2.87

Route

Year

Stateline-Ovid

1971
1972
1973

2
4

East Tamarack

1971
1972
1973

West Tamarack

r

�-121-

Arkansas River
The primary bobwhite areas along the river from Las Animas to Rocky Ford
show a decline in bobwhite numbers compared with 1972 (Table 2). The
lower Purgatoire, which contains a high concentration of bobwhite was also
down in population levels. The overall average along the Arkansas drainage
was 3.03 calling males per stop in 1972, compared to 2.40 calling males per
stop in 1973.
Cimarron Drainage
Table 2 indicates a considerable decline in the 1973 population compared
with previous years. Since the Baca County route runs through sand sage
rangelands, quail are heard calling in all directions rather than in a semicircle as along the rivers. Less vegetation and less noise interference
also permit location of calling males at greater distances. Therefore, the
BacaCounty counts cannot be directly compared with other counts.
Republican Drainage
Results from past years (Tables 2 and 3) indicate the Republican drainages
generally possess low densities of bobwhite. The count on the Arikaree was
down considerably from the previous year.
Whistling Counts and Subsequent Fall
Population Trends on the Tamarack Study Area
Do summer whistling counts foretell fall population levels? Preliminary
Tamarack data presented in Table 4 indicate that whistling counts can be
used to predict population changes. In 1972, when the call index was up by
about a third, fall population levels and harvest also increased by about
that amount. In 1973, the index, the fall population level, and harvest
decreased, but not by quite the same percentages. Additional years of evaluation plus complete review of previous studies on the subject are planned for
the future.
Population Structure, Level and
Rate of Harvest on the Tamarack Study Area
Past Segment Band Recoveries and Population Estimates
First Segment Band Recoveries.--In the fall of 1971, 531 bobwhite were
banded. Only two recoveries from this original sample were obtained during
the fall and winter samples in 1973-74. Four bands were recovered during the
same period from 155 birds banded in the late winter of 1972. These data are
considered inadequate for use in revision of previously established population estimates. They do illustrate the short life span of bobwhite and
will be utilized later in the derivation of mortality and survival rates.
Second Segment Band Recoveries.--The initial projection of 2,3l9 quail alive
in October, 1972 will be retained as the best estimate of population level

�-122-

Table 2. A summary of bobwhite whistling counts completed in southeast
Colorado and the Republican River Drainage in 1971, 1972 and 1973.

Year

Nwnber
of
Counts

Number
of
Stops

Natural
Tally

Mean

E. Prowers County

1971
1972
1973

1
3
2

10
30
26

8
33
45

0.8
1.1
1.73

Purgatoire

1971
1972
1973

1
2
3

10
24
40

52
194
158

5.2
8.1
3.95

Las Animas-La Junta

1972
1973

2
4

27
47

152
128

5.6
2.72

La Junta-Rocky Ford

1972
1973

2
3

23
37

76
87

3.3
2.35

Rocky FordManzanola

1972
1973

2
2

18
17

8
12

0.4
0.71

Fowler-Boone

1972
1973

3
1

34
14

9
4

0.3
0.29

1971
1972
1973

1
2
2

10
28
25

38
145
49

3.8
5.2
1.96

Arikaree

1972
1973

1
2

11
20

31
8

2.8
0.40

South Fork-Bonny

1971
1972
1973

3
30
2
27
No counts made

21
12

0.7
0.4

Location and Route

Arkansas River Valley

Cimarron River Valley
S. E. Baca County

Republican River Drainage

�-123-

Table 3. A comparison of high counts obtained on bobwhite quail census
routes in eastern Colorado during 1971, 1972 and 1973.

Route

1971

Year
1972

1973

South Platte Drainage
Stateline-Ovid

5.3

3.4

4.0

East Tamarack

4.0

6.1

3.1

West Tamarack

4.1

8.2

2.2

4.3

4.6

Sterling-Iliff
Atwood-Merino

6.7

9.1

9.5

Brush-Fort Morgan

3.9

8.3

2.3

Goodrich-Orchard

1.4

2.3

0.6

2.8

0.6

Republican Drainage
Arikaree

Arkansas Drainage
E. Prowers County

0.8

1.8

1.8

Purgatoire

5.2

9.4

4.5

Las Animas-La Junta

6.1

3.7

La Junta-Rocky Ford

3.5

2.5

Rocky Ford-Manzanola

0.9

0.7

Fowler-Boone

0.4

0.3

5.5

2.1

Cimarron Drainage
S. E. Baca County

3.8

�-126Table 5. Derivation of natural mortality of bobwhite during the fall and
winter of 1972-73.
West
Tamarack

East
Tamarack

Total

Fall, 1972 banded quail remalnlng
(corrected for emigration)

483

306

789

Projected 1972 harvest removal
(corrected for crippling loss)

138

73

211

Fall, 1972 quail subject to natural
mortality

345

233

578

Quail banded or retrapped in
February, 1973

63

15

78

From Fall, 1972 banded sample
From February, 1973 sample

11
10

4
2

15
12

Fall, 1972 banded alive at end of
winter, 1973 1./

69

29

98

Fall-winter natural mortality of
banded sample

276

204

480

Projected 1972 fall population estimate

1,521

798

2,319

Fall-winter natural mortality of total
population 2:./

869

532

1,401

57.13

66.67

60.04

444

190

634

28.57

23.86

26.74

Item

Recoveries obtained during fall
and winter, 1973-74:

Percent natural mortality, fall and
winter
Total 1972 season harvest and crippling
loss
Percent harvest mortality
Total mortality, natural + harvest
Percent mortality, natural + harvest
Total bobwhite alive mid-February, 1973

1,313

722

2,035

85.70

90.53

86.78

208

76

284

l/ The ratio of fall to winter band recoveries based on the total winter banded
sample was used to project the number of fall-banded quail alive in late
winter, 1973.

£/ Based on a 33.77 percent fall-banded sample as shown in the 1972 harvest
recovery of 548 quail.

�-127-

Table 6. Natural mortality of bobwhite through the
1973 on the Tamarack Study Area.

Item

spring and summer of

Males

Females

Total

Bobwhite alive February, 1973 11

154

130

284

Adult bobwhite projected alive October, 1973 11

82

64

146

56.2

43.8

72

66

Proportion of mortality between sexes

52.2%

47.8%

Percent mortality, Spring through Summer
based on February population level

46.7

50.8

48.6

Percent mortality, Spring through Summer
based on 1972 Fall population level

5.7

6.2

5.9

Percent composition of survivors
Natural mortality--Spring

through Summer, 1973 11

Annual mortality--Fall, 1972 to Fall, 1973
Percent annual mortality, Fall, 1972 to
Fall, 1973

1,180

93.5

993

138

2,173

93.9

1lBased on sex ratio data from Table 19 in Snyder, 1973.
1lBased on 1973 fall population estimate (Table 8) and age and se~ ratios
(Tables 14 and 15).
liThe difference between line 1 and line 2 above.

93.7

�-128-

Table 7. An estimate of mortality and survival of bobwhite quail from 1971
through 1973 based on the relative recovery rate method. !!

Period Banded

Number Banded

Number of Recoveries
Each Year After Banding
1
2

Fatl, 1971

531

119

34

Fall, 1972

802

175

13

Fall, 1973

300

83

Recovery Rate
Each Year After Banding
1
2

0.0640
0.2182

0.0162

0.2767

0.4949
Percent

0.0802
Percent

First Year Survival Rate

29.33

Mortality Rate

70.67

Second Year Survival Rate

5.85

Mortality Rate

94.15

Average Survival Rate

16.21

Mortality Rate

83.79

"};/ Based on procedures from Funk, 1966.

Fall Banding, 1973
During the fall of 1973, 197 bobwhite were banded on the West Tamarack and
103 quail were banded on the East Tamarack. Nine retrapped adults brought
the total sample to 309. CIR data in Table 8 indicate that 41 percent of
the quail were marked. High river levels restricted much of the trapping
to the south side of the Tamarack. Harvest also was restricted in the same
way to some degree. If neither banding or harvest was randomly distributed,
then a higher proportion of banded quail were taken and the actual population
level was higher than that projected in Table 8. It is not believed that
the bias was too great, and for the most part, it was confined to the West
Tamarack. Future samples will be too small to permit correction.
Fall :t-k&gt;vemen
t
Correction for movements and loss of banded quail from the study area prior
to harvest were calculated by the same method as in previous years (Snyder
1973). Most coveys were quite stable, with only one known covey moving a
long distance upstream to near Sterling. Correction for loss of banded quail
is applied in Table 8.

�-129-

Table 8. A summary of Fall, 1973 bobwhite banding and harvest information
from the Tamarack Management Area.

West
Tamarack

East
Tamarack

Total
Area

197

103

300

3

9

203

106

309

13

3

16

190

103

293

58

23

81

2

o

2

60

23

83

Crippling loss (projected) total quail

17

8

25

Known harves t

139

63

202

Projected total harvest

161

70

227

Percent crippling loss

15.83

11.11

12.38

Projected total band recoveries

67

26

93

Percent banded

43.16

36.51

41.09

Percent harvest (projected total)

35.26

25.Z4

31. 74

Item

Number of Fall, 1973 banded qu~i1
Number of retrapped adults
Total marked
Marked lost to emigration
Marked remaining

Band recoveries--Fa11,

1973 banded

Retrapped adults
Total band recoveries

Fall population estimate

440

Confidence limits

± 85.22

± 97.28

± 113.96

Confidence limits with
finite correction

± 67.80

± 84.28

±

+

282

=

722

94.38

�-130-

Fall Population Estimate
The fall population in 1973 was much lower than that in preceding years
(Tables 8 and 9). Marked and recapture samples were not sufficient to
prevent errors greater than 10 percent, with 95 percent accuracy as
recommended by Robson and Regier (1964). Due to high winter mortality, this
level of sampling will not be achieved. Confidence limits for the fall
population are presented in Table 8.

Table 9.
1973.

Bobwhite fall popUlation and harvest statistics for 1971 through

Item

Fall population estimate

197i

1972

1973

1,745

2,319

722

Season length (days)

20.5

31.5

32.5

Known harvest

411

548

202

Projected total harvest mortality !/

472

634

227

Percent harvest mortality y

27.0

26.7

31.7

!/ Includes projected crippling loss.
Harvest 1973
The 1973 quail season in Small Game Management Unit #1 began at noon on
Saturday, November 17 and ended on the 25th. An extended season along the
South Platte River opened on December 8 and terminated at the end of the
month. Quail and pheasant hunting was thus permitted for 32.5 days in
1973, compared to 31.5 days in 1972, and 20.5 days in 1971.
Hunters removed 202 bobwhite from the Tamarack during the 1973 season.
Projected crippling loss increased the mortality to 227 birds (Table 8).
This harvest was well below that of either of the two preceding years
(Table 9).
Hunting Pressure and Hunting Success
Hunting success of bobwhite was much lower in 1973 than in either 1971 or
1972. During the first 8 days of the 1973 season, 5.8 hunters checked into
the Tamarack per each quail taken from the area (Table 10). In contrast,
in 1971 and 1972, 2.47 and 2.63 hunters per bagged quail, respectively,
were checked. Hunter success declined with season progression each year, but
again, the most dramatic decline occurred in 1973 (Table 10).

�-131-

Table 10. Hunter success ratios on bobwhite quail with the progression of
hunting seasons in 1971, 1972 and 1973.
Interval

Number of Hunters!.!

Number of Quail

Hunters/Quail

1971 Season
First 8 days

638

258

2.47

Second 8 days

363

100

3.63

Last 5 days

258

53

4.87

1972 Season
First 8 days

606

230

2.63

Second 8 days

386

167

2.31

Third 8 days

369

100

3.69

Fourth 8 days

392

36

10.89

1973 Season
First 8 days

823

142

5.80

Second 8 days

339

36

9.42

Third 8 days

161

10

16.10

Fourth 9 days

303

13

23.30

!/ Includes all hunters using the Tamarack.

During 1972 and 1973 hunters
during the latter part of the season were primarily duck hunters.

Hunting pressure was nearly the same during the early parts of the 1971 and
1972 seasons, whereas increased pressure was recorded during the first eight
days of 1973 (Table 10). This pressure dropped off more rapidly than in
previous years, undoubtedly due to poor hunting success on most species
(Table 11). Increased hunting pressure, primarily attributed to duck
hunters, was noted during the holiday period of late December.

�Table 11. Harvest of small game per day on the Tamarack Management Area during the 1973 quail and pheasant
season. y
Bobwhite Quail
Date

Pheasants

Duck;!:..! Rabbits

Squirrels

Adult
Male

Adult
Female

Imm.
Male

Female

Unclassified

Total

5
2
0
2
0
0
1
0
3

18
8
4
8
6
1
10
7
3

8
17
4
3
4
3
6
6
2

3
0
1
0
0
0
0
0
O·

40
28
9
14
10
4
22
14
12

13

65

53

4

153

1
1
0
0
0
1
0
0
0
0
0
0
0

4
1
0
1
0
1
0
0
0
3
0
0
2

3
3
0
2
0
2
1
1
1
0
0
0
2

1

11
5
0
4
0
4
1
2
1
3
0
0
4

Imm.

First Season
11-17
11-18
11-19
11-20
11-21
11-22
11-23
11-24
11-25

Subtotal

:

0
1
0

9
9
7
3
3
1
1
3
5

45
37
15
50
15
4
11
23
7

8
4
4
0
8
0
5
4
1

0
0
0
1
0

6
1
0
1
0
0
5
1
4

41

207

34

2

18

o

Extended Season
12-8
12-9
12-10
12-11
12-12
12-13
12-14
12-15
12-16
12-17
12-18
12-19
12-20

5
2
0
2
1
1
0
0
1
0
0
0
1

10
5
2
1
3
3
0
3
0
0
22
27
7

2
2
0
0
0
0
0
2
0
0
0
0
0

0
0
0'
0
0
0
0
1
0
0
0
0
0

2
0
0
1
0
0
0
1
0
0
0
0
0

0
0
0
'0
0
0
0
0
0
0
0
0

------------------------------------------------------------------------------------------------------------------

I
•...•

w

N
I

�Table 11. Harvest of small game per day on the Tamarack Management Area during the 1973 quail and pheasant
season (continued). 1./

Date

Pheasants

Ducks-2/

Rabbits

Squirrels

Adult
Male

Adult
Female

Imm.
Male

Bobwhite guail
Imm.
Female
Unclassified

Total

Extended Season (cont.)
12-21
12-22
12-23
12-24
12-25
12-26
12-27
12-28
12-29
12-30
12-31

0
0
1
0
0
0
4
0
3
0
0

9
30
15
2
0
15
8
12
23
21
16

0
5
2
1
0
0
2
0
11
0
2

0
0
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
1
1
1
0
0

0
0
0
0
0
0
1
0
1
0
0

0
0
1
0
0
0
1
0
1
0
1

0
1
1
0
0
0
1
0
1
0
1

0
0
0
0
0
0
0
0
0
0
0

0
1
2
0
0
0
4
1
4
0
2

Subtotal

21

234

29

1

7

5

16

20

1

49

Total

62

441

63

3

25

18

81

73

5

202

1/ Seasons for waterfowl, rabbits and squirrels extend beyond dates listed here.
'!:../ Includes 393 mallards and 48 others of several species.

I

~

w
w
I

�-134-

Hunter

Use and Success by Hunting Area

Utilization of hunting areas is indicated in Table 12. Also included are
hunter hours, quail banded, total quail harvested and banded quail harvest
per area, and per east and west sections of the Tamarack.
Assuming six
square miles of hunting area (Snyder 1972), each square mile of the 'I'amanac
k
river bottom was hunted by about 270 hunters.
Each square mile sustained
about 909 man hours of hunting pressure during the quail season.
Categorizing this hunting pressure per species or even between upland game and waterfowl cannot be made.
Many additional hunter days and man hours of use were
accumulated during dove, deer, duck and rabbit seasons which extended
through various parts of the fall and winter.
Harvest

of Bobwhite

in Relation

to That of Other Species

Species harvest by day of season is illustrated in Table 11. Harvest of
quail is broken down to age and sex. Table 13 compares the harvest of small
game for years 1971, 1972 and 1973. Harvest of all species declined in 1973
from 1972 levels.
Only 1973 duck and rabbit harvests exceeded those in 1971.

Winter Banding 1974
Most of February, 1974 was utilized in quail trapping efforts.
Donald M.
Hoffman assisted for two weeks.
Despite these tenacious efforts, only twelve
bobwhite were banded, and six were retrapped at various locations on the
Tamarack.
Efforts to locate quail by tracking in snow in combination with
use of bird dogs was productive.
However, these efforts were restricted due
to lack of snow cover through most of the month.
Searches that were completed
indicated that large areas of the Tamarack were completely void of bobwhite,
and the few coveys that were located by baiting or searches were all quite
small.
Only about 40 quail were found through trapping, searches and contacts, but it is possible that two to three times that number of quail are
present.
The small trapped sample will not provide an adequate base for use in CIR
projections from subsequent band returns.
A map is enclosed (Fig. 1)
showing known covey locations, but it must be accepted that accuracy in
measurement of population variables is directly related to population size.
The smaller the population, the less accurately we can measure it.

Age Ratios-Fall

to Winter,

1973-74

Trap bias toward young was again evident in the fall banded sample as in
previous years (Table 14). The percentage of adults taken during the November hunting season (20.3 percent) was considered to benost
closely approximate
the proportion of adults in the fall population.
Disproportionately
higher
juvenile mortality may have increased the proportion of adults in the subsequent harvest and winter banding samples.
In 1972, approximately 16.3
percent of the population were adults and in 1971 the percentage adults
was 19.2

�-135Table 12. A summary by hunting area of fall banded bobwhite, hunter use,
total harvest and banded harvest recoveries on the Tamarack Management Area
in 1973.

Area

Number Quail
Banded

Number of
Hunters y

Hunter
Hours }j

Total Quail
Harvest '!;.I

Banded Quail
Harvest 11

9

4
3

West Tamarack
1
2

4

3
4
5

16

6
7

16

53
50
46
47
27
48
41

8

2

33

9

2

6

o
2

o

146
114
69
79
66

13

o

14
15
18
19
20
21
22
23
24
25

5

46
39
53
57
50
54

o

33

17

97

9

55

11
27
18

30

48
47
59
43
39
30

128
102
64
119
106
175
174
206
179
128
414
222
256
194
230
211
148
188

197

1,095

3,718

10

6

11

18

12

Subtotal

o

8

o

5
3

o

o
o
o

o
6

1

2
2

o

4

1
1
6

o
8
8
3
2
4

12
20
12
3

14
10

o
o
o

o

o
6

11
8
1
7

6

1

o

11

2

139

57

o

o

East Tamarack
Braddock Pasture 34 ~I
1

14

2
3
4
5
6
7
8

6
1
2

9

13
6
2

25

o
o

68
58
49
35
45
50
54
36
30
45

182
203
151
120
140
184
187

4

3

7
11

1

2

15
12

III
III

7
2

o
o
8
6
5
1

o

172
172

o

55

3

o
o

Subtotal

103

525

1,733

63

23

Total

300

1,620

5,451

202

80

10
11

II Includes only known hunters and hours reported through the check station.

21 Includes only known harvest.

Some hunting probably went unrecorded.
Exact location unknown.
~I Braddock pasture and Areas 16 and 17 West are closed to public hunting.

11 Approximates the location where quail were shot.

�-136-

Table 13. A comparison of small game harvest during the 1971, 1972 and
1973 quail and pheasant seasons.

Year

Length of
Season

Quail

1971

20.5 days

411

1972

First 20.5 days

1972

Pheasants

Duck

Rabbits

Squirrels

87

196

30

9

480

130

555

126

4

31.5 days

548

143

704

156

4

1973

First 20.5 days

184

53

283

40

3

1973

32.5 days

202

62

441

63

3

Sex Ratios
Table 15 illustrates the greater proportional mortality among adult hens
than among adult cocks. As in 1972, this differential mortality occurred
during the summer. Some difficulties are encountered in sexing young juveniles during fall banding. Therefore, the sex ratios of fall trapped juveniles are not considered highly accurate.

Factors Limiting Bobwhite on the Tamarack
Fall and Winter Mortality
Available evidence indicates that two successive winters with prolonged snow
cover accompanied by extreme cold have nearly wiped out the Tamarack quail
population. These weather conditions came in conjunction with the 1972
hunting season, so hunting stress probably added to the natural mortality.
Rosene (1969) stated that bobwhite are quite susceptible to cold and exposure
when their feathers become wet. Short days and long nights in winter limit
the hours of feeding. Coveys must be united to provide communal warmth over
night in below zero weather. Hunting, especially under snow conditions and
in late afternoon could greatly heighten stress by causing wet feathers and
by preventing feeding and covey reorganization. Available evidence indicates
hunting removed about 634 quail in 1972, while 1,401 birds were lost to
unknown causes. The drastic decline in hunter success by the end of the 1972
season (Table 10) provided evidence that much of this mortality had occurred
prior to the end of the hunting season. Pre-baiting and covey searches in
January, 1973 confirmed this.

�TAMA RACK MANAGEMENT

AREA

'fl £$1

AREAS

I
•.....
w

~
IV

/
EAST

AREAS

SCALE

o
LEGEND
1

l&gt;iY!GRASSLAND

Fig. 1.

1 Mi.

()

Covey Location - 1973

~

Covey Location - 1974

HUNTING AREA

~UNFARMED
RIVER BOTTOM ~
ALFALFA
~
AND RIVER CHANNELS
L.:.-J

DHAY

Yo

MEADOW

[:~~:~

ROW CROPS
PLANTING

Bobwhite coveys located during late winter trapping efforts in 1973 and 1974.

I

�-138-

Table 14. Bobwhite age ratio information obtained during the fall and
winter of 1973-74.

Sample

Number
Adults

Number
Young

Total

Percent
Adults

Adul t:Yearling
Ratio

Fall Banded

31

278

309

10.03

1:8.97

Banded

4

62

66

6.06

Unbanded

26

56

82

31.71

Total

30

118

148

20.27

Banded

3

l3

16

18.75

Unbanded

8

23

31

25.80

Total

11

36

47

23.40

Banded

7

75

82

8.54

Unbanded

34

79

113

30.09

Total

41

154

195

21.03

1:3.76

5

13

18

27.78

1:2.60

November Season

1:3.93

December Season

1:3.27

Combined Season Data

Winter Banded

In 1973, snow cover began about December 18, after the main hunting stress
had passed. Snows, deeper than in the previous year (8 to 10 inches), persisted until mid-January. Two small coveys were observed flying an eighth
of a mile or more out to exposed roadways to feed, indicating a stress for
food. The few small coveys that survived the winter were resident in the
heaviest available cover. Snow berry (Symphoricarpos sp.) and wild rose
(Rosa sp.) generally did not provide adequate cover in the deep snow. Most
willow (Salix sp.) clumps were too sparse and open. Cordgrass (Spartina
pectinata), other grasses and most forbs were flattened under the snow. The
tall dense stands of Phragmites grass provided the key to survival for most
rema1n1ng coveys. Pheasants also concentrated in these locations. Many of
the Phragmites stands, which did not contain quail, also did not contain
pheasants. Lack of food and adequate adjacent cover were probably factors
limiting use of these cover patches. The presence of pheasants probably
increased food stress to bobwhite.

�-139-

Table 15. Sex ratios of adult and juvenile
and winter trapping and harvest, 1973-74.

bobwhites

obtained

during fall

Adult
Males

Adult
Females

Immature

Sample

Males!1

Immature
Females

Fall trapped

21

10

126

146

Percent by age class

67.7

32.3

46.3

53.7

Hunting

25

18

81

73

Percent by age class

58.1

41.9

52.6

47.4

Winter trapped

4

1

6

7

Percent by age class

80.0

20.0

46.2

53.8

Total

50

29

21.3

226

Mean percent by age class

63.3

36.7

48.5

51.5

determined

during this

Season

(total birds)

II Sex of immatures
sample period.

could not always be accurately

The part which predators played in decimating quail populations under snow
stress conditions remains unknown.
However, it undoubtedly played an important part, since the quail were forced to expose themselves when feeding and
could not readily find escape cover.
Goshawks, great horned owls and coyotes
were among the most active predators along the river during snow stress
periods.

Spring and Summer Mortality

and Reproduction

Whistling counts conducted along the South Platte and Tamarack indicated population declines occurred in 1973 based on 1972 levels (Table 4). A second
variable, spring flooding in 1973 was also a factor to consider in quail
survival and reproduction.
Present evidence indicates the snow stress, not
flooding, decimated the population.
The flood delayed nesting, but a significant reduction in nesting and reproduction is not evident.
Flooding
occurred early enough in spring to permit abundant
vegetative growth and midto late summer quail reproduction.
Projected Period and Peak of Hatch.--Fig. 2 illustrates the period and general
peak of quail hatch on a sample of 288 birds measured during fall trapp~g.

�-139-

Table 15. Sex ratios of adult and juvenile
and winter trapping and harvest, 1973-74.

bobwhites

obtained

during fall

Adult
Males

Adult
Females

Immature

Sample

Males!1

Immature
Females

Fall trapped

21

10

126

146

Percent by age class

67.7

32.3

46.3

53.7

Hunting

25

18

81

73

Percent by age class

58.1

41.9

52.6

47.4

Winter trapped

4

1

6

7

Percent by age class

80.0

20.0

46.2

53.8

Total

50

29

21.3

226

Mean percent by age class

63.3

36.7

48.5

51.5

determined

during this

Season

(total birds)

II Sex of immatures
sample period.

could not always be accurately

The part which predators played in decimating quail populations under snow
stress conditions remains unknown.
However, it undoubtedly played an important part, since the quail were forced to expose themselves when feeding and
could not readily find escape cover.
Goshawks, great horned owls and coyotes
were among the most active predators along the river during snow stress
periods.

Spring and Summer Mortality

and Reproduction

Whistling counts conducted along the South Platte and Tamarack indicated population declines occurred in 1973 based on 1972 levels (Table 4). A second
variable, spring flooding in 1973 was also a factor to consider in quail
survival and reproduction.
Present evidence indicates the snow stress, not
flooding, decimated the population.
The flood delayed nesting, but a significant reduction in nesting and reproduction is not evident.
Flooding
occurred early enough in spring to permit abundant
vegetative growth and midto late summer quail reproduction.
Projected Period and Peak of Hatch.--Fig. 2 illustrates the period and general
peak of quail hatch on a sample of 288 birds measured during fall trapp~g.

�-140-

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~
Il.
5

\
\\

[hI

..."

,/
v ..•.

"

I
r-_
-1
I

I

4

Sample Size

I
I

3

N = 768 in 1972

f

I

N = 288 in 1973

/

2
I
I

1

,

I

I

o
18

25

May

2

9

17

June

24

3

11

19

27

July

4

12

20

August

28

7

14

Sept.

Weekly Mid-points

Fig. 2. Period and peak of bobwhite quail hatch projected from prima~y
feather growth for 1972 and 1973 on a percent of total basis.

�-141-

Wings were measured and age data projected using a method described by
Rosene (1969). Very little hatching occurred prior to mid-June, and most
occurred after the first week in July. A second high peak occurred during
the second week of August. Newly hatched broods were present even as late
as the second week of September illustrating the tenacious efforts of the
hens to bring off a brood.
Most of the hatch in 1972 came about two weeks
ahead of that in 1973 (Fig. 2).
Nesting Success.--The February, 1973 base population of 284 quail gained in
number by 254 percent to a fall level of 722 birds.
In the preceding year,
the gain from the February base of 988 quail was 235 percent.
This indicates
that the 1973 population made a better gain percentage-wise than in the
previous year, despite the spring flood.
Since the population level was much
lower in 1973, this probably helped in reducing mort.ality over summer and in
making available more optimum nesting sites.
The percentage of adults in fall was 19.2, 16.3, and 20.3 for 1971, 1972
and 1973, respectively.
The higher percentage adults in 1973 might indicate
lower nesting success that year. However, it might also indicate a higher
rate of survival among adults.
Previously, presented data also indicate a
higher percent adult survival in 1973 than in 1972 from late winter to early
fall.
Vegetation

Sampling

Late summer inventories of vegetation were again obtained along the Tamarack
in 1973.
Samples were collected in west areas 2-3, 6-7, and 21-25, and in
east areas 4-6 and the Braddock pasture.
Results are illustrated in Tables
16, 17 and 18. Notable differences from the previous year included increases
in bare ground, curly dock and cockle burr due to flooding.
Sweet clover
which was abundant in 1972 was near non-existent in 1973. Ragweeds continued
among the dominant forbs. Cordgrass, witchgrass, barnyard grass and Canada
wild rye remained among the most abundant grasses.
Cover density averaged
slightly less in 1973 and height averaged slightly greater than in 1972.
However, two of the locations were not sampled the previous year, so direct
year to year comparisons are questionable.
Michael Loftsgard, a Colorado State University Wildlife Management Student
conducted food habits analyses on quail crops collected through the November,
1973 hunting season, and on a limited sample obtained during the 1972 hunting
season.
His information, presented in Table 19, provides a comparison of
fall food consumption in relation to plant species occurrence along the study
area. Wassink's (1972) information is also included.
Ragweeds (Ambrosia spp.),
knotweed and other Polygonum spp. were among the most important fall foods.
Sunflower (Helianthus spp.), milkweed (Asclepias spp.), and sweet clover
(Melilotus spp ,) were also commonly utilized.
Field penny cress (Thlaspia
arvense) and other mustards mature early in spring and summer, so may be more
important as food earlier in the year.

�Table 16. Percentage occurrence of vegetation, litter and bare ground on several sites within the Tamarack Management Area in late summer, 1973.

Cover Type

2-3 West
Sum
Percent

6-7 West
Sum
Percent

Location
21-25 West
4-6 East
Sum
Percent
Sum
Percent

Braddock
Sum
Percent

Grass

436

43.9

334

39.2

656

54.6

407

40.1

322

33.0

2,155

42.8

Brush and vines

312

31.4

94

11.0

147

12.2

367

36.2

244

25.0

1,164

23.1

Forbs

141

14.2

218

25.6

262

21.8

110

10.8

184

18.9

915

18.2

Dead vegetation

12

1.2

9

1.1

5

0.4

3

0.3

12

1.2

41

0.8

Combined
Sum
Percent

I

Litter

31

3.1

27

3.2

15

1.3

15

1.5

3

0.3

91

1.8

I-'
.po
N
I

Bare ground

61

Total Occurrences

993

6.2

169
851

19.9

116
1,201

9.7

112
1,014

11.1

210
975

21.6

668
5,034

13.3

�-143-

Table 17. Height and density measurements obtained at several locations
along the Tamarck Management Area in late summer, 1973.

Location

No. of Measurements

Mean Height
in Inches

. 1/
Mean Densl.ty-

2-3 West

210

28.82

1.62

6-7 West

210

18.87

1.96

18-19 West

210

25.43

2.47

4-6 East

210

15.80

1.98

Braddock Pasture

210

21.06

2.13

18.33

1.69

Average

1/ Based on an ocular estimate scale of 1, 2, or 3, respectively, representing
open, medium and dense cover.
LITERATURE CITED
Funk, H. D. 1966. Review of duck literature relating to population dynamics,
and banding analysis techniques and findings. Colo. Dept. Game, Fish
and Parks. Game Res. Rept. Oct. p. 77-108.
Loftsgard, M. 1974. Fall food habits .of bobwhite on the Tamarack 11anagement.
Area. Colo. State Univ., Dept. of Fishery and Wild1. Biology. Mimeo Rept.
Robson, D. S., and H. A. Regier. 1964. Sample sized in Peterson mark-recapture
experiments. Trans. Amer. Fish. Society 93(3):215-226.
Rosene, W. 1969. The bobwhite quail.
New Jersey. 418 p.

Rutgers Dniv. Press, New Brunswick,

Snyder, W. D. 1972. Population status of bobwhite quail in eastern Colorado.
Colo. Div. of Game, Fish and Parks. Game Res. Rept. April. p. 121-150.
1973. Population status of bobwhite quail in eastern Colorado.
Div. of Wildlife. Game Res. Rept. April. (In Press).

Colo.

Wassink, J. L. 1972. Fall food habits of bobwhite in eastern Colorado.
State Dniv., Dept. of Fishery and Wildl. Biology. Mimeo Rept.

Colo.

Prepared bY

~~~~~~~~~~~

~~~~~~~~~

wa;renrJ.SrlYde~

Wildlife Researcher

)_

�Table 18.

Forb occurrence on five sites along the Tamarack Management Area in late summer, 1973.!I

Genera or Species

2-3
West

6-7
West

21-25
West

4-6
East

Braddock
Pasture

Total

Percent
Occurrence

Per. Ragweed
Lambs quarter
Kochia
Horseweed spp.
Sunflower
Annual ragweed
Wild lettuce
Milk purslane
Purslane
Curly dock
Ground cherry
Poison hemlock
Dandelion
Gumweed
Crotons
Snow-on-the-mountain
Pigweed spp.
Milkweed
Indian hemp
Mustard spp.
Licorice
Vervain spp,
Goldenrod
Mint
Plantain
Prickly pear
Cocklebur
Canadian thistle
Clammy weed

13
1
3
0
3
1
0
0
0
14
5
8
0
0
0
0
0
21
34
0
1
0
1
0
0
2
2
7
3

10
2
5
0
4
16
1
2
0
40
5
6
0
0
0
3
32
4
14
2
1
0
0
0
0
1
25
0
4

29
0
0
1
14
4
0
1
0
88
6
6
0
0
0
5
3
49
0
0
9
0
0
2
0
3
26
0
5

26
0
0
0
3
4
0
5
0
1
4
12
2
1
0
1
7
8
16
0
1
0
0
0
0
7
5
0
2

15
0
0
0
0
0
0
5
19
14
0
3
7
0
1
1
4
0
1
0
18
9
0
0
1
1
34
0
8

93
3
8
1
24
25
1
13
19
157
20
35
9
1
1
10
46
82
65
2
30
9
1
2
1
14
92
7
22

10.16
0.33
0.87
0.11
2.62
2.73
0.11
1.42
2.08
17.16
2.19
3.83
0.98
0.11
0.11
1.09
5.03
8.96
7.10
0.22
3.28
0.98
0.11
0.22
0.11
1.53
10.05
0.77
2.40

-----------------------------------------------------------------------------------------------------------

I
•.....

.p.
.p.
I

�Table 18.

Forb occurrence on five sites along the Tamarack Management Area in late summer, 1973 (cont.).l/

Genera or Species

2-3
West

6-7
West

21-25
West

4-6
East

Braddock
Pasture

Total

Percent
Occurrence

0
13
0
8
1
0
0
0
0

18
5
1
14
0
2
1
0
0

0
5
0
4
2
0
0
0
0

0
0
0
1
2
2
0
0
0

2
7
4
9
6
0
0
1
14

20
30
5
36
11
4
1
1
14

2.19
3.28
0.55
3.93
1.20
0.44
0.11
0.11
1.53

-Knotweed
Smartweed
Uniden t·ified compos itae
Tickseed
Snakeweed
Sedges
Buffalo burr
Puncture vine
Unidentified forb

I
•....
.po

V1
I

Summation

141

218

262

110

184

915

1/ Species present in 1972 but not in 1973 - clover, four o'clock, mullein, horsetail, Russian thistle,
tall evening primrose, prostrate primrose,

butterfly guara, goatsbeard, gi1ia.

�-146-

Table 19. A summary of principal foods utilized by bobwhite quail during late
fall periods in comparison with occurrences of some of these species in late
summer vegetative samples. !I

Species

Presence in CroEs
Percent Occurrence
Percent b~ Volume
1973
1972
1971
1973
1972
1971

Annual ragweed

45.4

12.8

Perren~a1 ragweed

27.9

12.8

Milkweed

32.6

10.3

Halberd-leaved
orache

19.8

46.2

Lambs quarter

3.5

10.3

Pigweed

14.0

7.7

Clammy weed

45.4

0.0

Smart weed

3.5

7.7

Knot weed

31.4

Sunflower

68

14.9

0.2

2.8

4.3

30

4.0

1.3

38

0.3

0.9

0.5

0.0

T

T

3.2

0.0

2

T

0.1

10.3

80

6.8

52.3

41.0

58

Kochia

18.6

15.1

Curly dock

36.1

5

Percent Occurr.
in Veg. SamEles
1973
1972
2.7

4.3

10.2

18.0

1

9.0

1.7

1

Y
0.3

7.1

5.0

0.4

2.4

0.1

T

3.3

0.9

0.2

20

2.2

0.1

7.0

2.8

4

2.6

5.6

20

T

0.2

1

0.9

5.7

33.3

44

0.8

1.2

T

17.2

1.7

Snow-on-the-mountain 16.3

2.6

22

0.9

0.1

T

1.1

0.3

Texas croton

18.6

0.0

0.7

0.0

0.1

1.1

Spurge

1.1

0.0

T

0.0

2/

Milk purslane

2.3

0.0

T

0.0

1.4

4.1

Ground cherry

23.3

2.6

0.3

0.1

2.2

2.1

Verbena spp.

17.4

2.6

2

T

0.1

T

1.0

1.6

Sweet clover

4.6

23.1

14

0.1

1.2

T

0.0

10.1

Russian thistle

7.0

2.6

T

0.1

0.0

0.2

Motherwort

2.3

0.0

T

0.0

0.2

0.4

0.0

1.0

0.0

0.0

0.0

Mint sage

9.3

16

T

-------~-----------------------------------------------------------------------

�-147Table 19. A summary of principal foods utilized by bobwhite quail during late
fall periods in comparison with occurrences of some of these species in late
summer vegetative samples. 1/

Species

Presence
Percent Occurrence
1973
1972
1971

Field penny cress

7.0

61.5

Crownbeard

12.8

0.0

Night shade

3.5

0.0

Mercury

3.5

0.0

Poison ivy

5.8

7.7

Grape

12.8

7.7

Witchgrass

19.8

0.0

Switch grass

16.3

0.0

Foxtail

2.3

0.0

Dropseed

11.6

Proso millet

58

in Crops
Percent by Volume
1973
1972
1971

T

22.8

0.6

0·0

0.9

0.0

T

0.0

4.0

1.1

0.8

0.7

T

0.0

T

0.0

T

0.0

0.0

2.3

0.0

7.0

0.0

13.5

0.0

Corn

16.3

28.2

21.7

53.7

Insect matter

54.7

7.7

9.4

0.4

Vegetative

29.1

10.3

0.1

0.1

matter

2

4

4

14

48

5

Percent Occurr.
in Veg. Samples
1973
1972

0.2

1.0

Not identified
T

0.0

0.0

Not identified
T

T

32

4

1/ Crop analysis
Wassink

data for 1972 and 1973 by Loftsgard (1974). Data from 1971 by
(1972). Crop sample size for 1972 was considerably below that for 1973.

~ Orache and lambs quarter were not identified
sampling.
Croton and spurge were also combined

separately in vegetative
in sampling.

11 Poison ivy and grape are both common, but were not listed by percent
occurrence.
~ Grasses were not separated by species in vegetative sampling.
are common to abundant in occurrence along the Tamarack.

All listed

~/ Proso millet was planted in food plots on the east Tamarack.
Corn was used
in pre-season trapping and is also available adjacent to the river bottom in a
few locations each year.

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                  <text>-1-

July, 1974
JOB PROGRESS REPORT

State of
Project No.

COLORADO

4

\-lork Plan No.

Job Title

Job No.

1a

_

----~.----------~------.---------

Inventory of Range Manipulation Projects in Colorado

Period Covered:
Per sonnal.:

Game Range !~ves_t_i~g~a~t_i_o_n_s
~

---W-101-R-16

April 1, 1973 through March 31, 1974.

Roland C. Kufe1d and Regional Game Biologists.

ABSTf\ACT
An inventory was made of all range vegetation-modification projects completed
during 1971, in the western half of Colorado, on lands administered by the
U. S. Forest Service and Bureau of Land Management. Acreages treated were:
Forest Service - 3,690 and Bureau of Land Management - O.

��-3-

INVENTORY OF RANGE MANIPULATION PROJECTS IN COLORADO
Roland C. Kufeld

P. S. OBJECTIVE
To collect information which describes proposed and completed range vegetation modification projects that are located in Colorado, and their effects
on the range, wildlife and livestock; and to provide desired IBM listings of
these data to cooperating agencies upon their request.
SEGMENT OBJECTIVES
1.

To collect information which describes proposed and completed range
vegetation modification projects that are located in Colorado and their
effects on the range, wildlife and livestock.

2.

To compile, codify, process and analyze inventory data.

3.

To provide desired IBM listings of inventory data to cooperating agencies
upon their request.
METHODS AND MATERIALS

An inventory was made of all range vegetation modification projects completed
through 1971, in Colorado, west of Interstate Highway 25 on lands administered
by the U. S. Forest Service and Bureau of Land Management (Kufe1d 1968, 1970,
1971 and 1973). Kufeld (1968 and 1970) also covered projects completed on the
Southern Ute and Ute Mountain Indian Reservations through 1969. Indian
Reservation projects were dropped from the inventory after 1969. This report
concerns vegetation modification projects completed on U. S. Forest Service
and U. S. Bureau of Land Management lands west of Interstate Highway 25
during 1972.
Data were collected using procedures outlined in Colorado Division of Wildlife
Administrative Directive No. 204 entitled "Range Vegetation Modification
Projects", and by Kufeld (1968 and 1970). Additional procedures initiated
this year and which will continue to be used during future post-treatment
evaluations are described by Greg L. Schenbeck and included in this report.
Schenbeck's report also includes helpful suggestions which will increase the
efficiency of those involved in making post treatment evaluations.
Inventory data were transferred from original data sheets to Mark Page Reader
Forms, and then to IBM cards.
Procedures outlined in Administrative Directive No. 204, call for evaluations
to be made on each vegetation modification project at the end of the 2nd, 5th,

�-4-

and 10th years following treatment. During 1972, 2 year post-treatment
evaluations were made on 15 vegetation modification projects that were
completed in 1970, and data recorded on a form described by Kufeld (1971).
Two requests for range vegetation modification project inventory information were processed during the 15th segment. These were for the U. S.
Forest Service Rocky Mountain Regional Office and the Natural Resources
Defense Council, Inc.

RESULTS AND DISCUSSION
Acreages treated during 1972, on all lands administered by the U. S. Forest
Service west of Interstate Highway 25, are shown by vegetation type and kinds
of treatment in Table 1. These acreages are shown by individual National
Forests in Table 2. No vegetation modification projects were completed on
Bureau of Land Management lands in Colorado during 1972.
A total of 3,690 acres were treated on National Forest lands in 1972. To
date (through Dec. 31, 1972)a total of 582,072 acres have been treated in
Colorado west of I~25 on lands administered by the U. S. Forest Service and
Bureau of Land Management. Of this total 242,565 acres were on Forest Service
and 339,507 acres were on Bureau of Land Management lands.
Since 1972 was only the third year for 2 year post-treatment evaluations of
completed vegetation modification projects, and since no 5 year evaluations
have yet been made, data from post-treatment evaluations of 1968, 1969 and
1970 projects will be kept on file and covered in a future report when enough
2 and 5 year post-treatment evaluations have been made to provide a basis for
a meaningful data analysis.
LITERATURE CITED
Kufeld, Roland C. 1968. Inventory of range manipulation projects in Colorado.
Colo. Div. Game, Fish and Parks. Game Res. Rept. P. R. Proj. W-lOl-R-lO
WP 4, Jla, July, Part 1. p. 1-121.
1970. Inventory of range manipulation projects in Colorado. .Colo.
Div. Game, Fish and Parks. Game Res. Rept. P.R. Proj. W-lOl-R-12, WP 4,
Jla, July, Part 1. p. 59-94.
1971. Inventory of range manipulation projects in Colorado. Colo.
Div. Game, Fish and Parks. Game Res. Rept. P.R. Proj. W-lOl-R-13, WP 4,
Jla, July, Part 1. p. 1-15.
1973. Inventory of range manipulation projects in Colorado. Colo.
Div. Wildl., Game Res. Rept. P. R. Proj. W-lOl-R-15, WP 4, Jla, July,
Part 1. p , 1-8.
Prepared bY~C/~
Roland C. Kufeld
Wildlife Researcher

�-5-

Table 1. Acreages of range land treated during 1972, in Colorado by the
U. S. Forest Service.
Vegetation
Type

Grass

Sagebrush

Browse

!/

Conifer

Abandoned Land

All Veg. Types

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as .
Part of Treatment

Spray

1

500

500

1

500

500

3

1,370

0

3

1,370

0

Seed Only

1

20

20

Chain 2:../

8

1,690

1,145

9

1,710

1,165

1

10

10

1

10

10

1

100

100

1

100

100

15

3,690

1,775

Spray

Seed Only

Seed Only

GRAND TOTAL

!/

Except for one 20 acre project all vegetation modification projects conducted
during 1972 in the "browse" vegetation type were in the "oakbrush" subtype.

2/
- Chaining includes chaining, cabling, railing and bulldozing.

�-6-

Table 2. Acreages of range land treated during 1972, within each
National Forest in Colorado.
National
Forest

Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seed as
Part of Treatment

Grand Mesa

Sagebrush

Spray

1

150

0

Gunnison

Sagebrush

Spray

2

1,220

0

Rio Grande

Browse

Seed Only

1

20

20

San Juan

Grass

Spray

1

500

500

8

1,690

1,145

Total all
Forests

!/

J:./

Browse !/

Chain

Conifer

Seed Only

1

10

10

Abandoned
Land

Seed Only

1

100

100

Total all
Veg. Types

Total all
Treatments

15

3,690

1,775

!l
Except for one 20 acre project on the Rio Grande National Forest all vegetation modification projects conducted during 1972 in the "browse" vegetation
type were in the "oakbrush" subtype.

J:./
Chaining includes chaining, cabling, railing and bulldozing.

�-7-

ADDITIONAL PROCEDURES FOR EVALUATION
of
RANGE TYPE CONVERSION PROJECTS
Greg L. Schenbeck

Introduction
A uniform procedure for evaluating range-type conversion projects must be
established to insure acceptable accuracy in the findings. Basically, the
evaluation is composed of the following 3 steps:
1.

Completion of a #4A or #4B form. (See A System for Inventory,
Evaluation and Exchange of Information on Range Type Conversion
Projects) •

2.

Making a photographic comparison between the treated and adjacent
untreated area and between successive pictures of the treated
area at various time intervals.

3.

Completion of a lOO-point paced transect to determine range
condition and trend. (See Step 5 - Determine Range Condition
and Trend).

The procedures to follow for each step will be described in entirety.

Procedures
To complete the #4A or #4B form the agency responsible for the project must
be consulted. Prior to the time of the evaluation, the agency is contacted,
and a representative of the agency is invited to accompany you. If a
representative can accompany you, make sure to inform him that you will be
at the site for several hours; he may wish to take another vehicle. If it
is not possible for them to send a representative into the field with you,
an appointment should be made to come to their office to get the necessary
information. It is helpful to get a copy of their surveyor planning report
on the project. It is also advisable to acquire the treatment dates of all
proposed projects early in the summer. After you get the dates, schedule
your inspections so you can evaluate the areas before the treatment is started.
On the back of the #4A and #4B forms there is a series of questions pertaining
to wildlife use of the area. This information can be acquired from a representative of the agency or from the W.C.O. whose district includes the project. Problems will usually arise when trying to schedule appointments with
the W.C.O.; do your best.

�-8A System of Evaluation of Range Type Conversion Projects
Page 2

Make sure that you include on the forms both the code numbers and verbal
description in the blanks. By including the verbal descriptions, it is
not necessary to return to the revegetation manual each time the forms
are examined. Also, at the request of the Regional Manager, C. E. Till,
write "pretreatment evaluation" at the top of thef/4A forms and "posttreatment evaluation" with the number of the years since the treatment
at the top of the #4B forms.
Any extra information that would help to evaluate the range modification
projects should be included in the remarks sections of the forms.
When in the office, a written evaluation of each project should be made
using the available information. Be careful which data is compared because in a few cases the transects may have been run in different areas
of the project.
It is necessary that a uniform procedure be followed in order that a photograph be taken of an identical area in the treated portion. First, a
cross-section of the area is chosen to represent the treated area. This
cross-section will be the route of the 100-point paced transect and is also
included in the photograph. Drive a post (a 6'6" CF&amp;I fence post) approximately 30 feet inward from the beginning of the cross-section. Measure 30
feet outward from this stake, and from this point, record the bearing of
the driven stake from where you're standing. At this point, drive in an
orange-topped post, and from some nearby, noticeable landmark, record the
bearing and distance from that landmark to the orange post. The PhotoPoint Reference Form lists the data which needs to be determined and recorded. At the bottom of the form draw a sketch of how to get to the
photopoint from some main roadway. A forester's compass, Jacob staff,
and 100-foot tape are required to gather the necessary data. When pictures
are taken, the camera is rested on the top of the orange-topped post, and
a chalk board is placed on the other post. The date, project number or name,
and the forest or BLM district are placed on the board. The photographer
places the chalk board in the center of the frame, focuses on the board,
and snaps the picture.
Photographs of the untreated areas are not taken in identical areas year
after year. Therefore, a photo-point is not established. It is suggested
that the photographer walk about 50 yards into the untreated area to take
the picture. The chalkboard with the necessary data on it can be placed
against a rock or trunk, etc.
When returning to photo-points already established, follow the sketch map
on the photo-point form. Find the orange-topped post and sight through the
opposite post; this is the route of the transect. A landmark that is in
line with the 2 posts is chosen, and this landmark is used to help keep you
on the transect course.

�-9-

A System of Evaluation of Range Type Conversion Projects
Page 3

When evaluating a proposed project, a transect needs to be completed only
in the proposed site. If an area is proposed for treatment, a transect
is run only in the area to be treated. However, if the area has been
completed years previous, transects should be run in both the treated and
untreated areas. There will be times when you cannot complete a transect
in untreated areas because of thick vegetation. Other situations will
occur that will require you to make your own adjustments in the procedures.
Transects in untreated areas cannot always be run in straight lines. For
example, large oak or P-J trees will have to be circumvented.
At each hit, the item found is dot-tallied. Chained oak. sage or pinyonjuniper plants are counted as litter if the plants are dying or dead.
Also, the nearest browse and grass species within a 1800 arc in front of
the hit and within 5 steps are identified, recorded, and dot-tallied. It
has been recommended that the initial move be 15 steps and the consecutive
moves be 5 steps each.
At each tenth hit, a pellet-group count is made. To be counted, a group
must contain at least 5 pellets, and at least one-half of the group must be
in the plot. All groups are counted, regardless of age. The number of
browse stems in the plot are dot-tallied according to age class. Step 5
of the Reg. 2 U. S. Forest Service Range Analysis Handbook for big game
"Determine Range Condition and Trend" gives instruction and advice in completing the transect. In addition, the number of living pinyon-juniper
plants per plot is determined and recorded.
While at the project site, plant species which are not in the plant
collection should be taken and prepared for the collection.

Note:

Staple all forms in file to each copy.

4A, 4B, Transect, Photo-point.

�-10PACED TRANSECT RECORD - BROWSE RANGE CONDITION
Project Area

_

Forest or BLM District
Examiners

Date

--------------------------------------------------------------------------------------------------------------------------------------------------Type Identification
No. (from map)
point, direction of travel, interval between samples)

GROUND COVER DATA
Item

Dot tally
transect
hits

Bare ground
and era!': na1le
Rock
Moss
Litter
Grasses
Forbs
Browse
Browse

No.
of
Hits

Ground
cover
%

PELLET GROUP COUNTS
Plot

D

-.-No.

E

1

t

1
2

i

,
I

,

3

1! C

S

Other
i

II

I

I

I

I

,

I

I

!

;

i

4

I

i

I

;

5
6

I

I

7
i

8
Grass

9

10
Totals

J
I

Per Acre
Factor

I
!

15

13

12

13

Days/A.

Totals

BROWSE CONDITION &amp; DENSITY/ACRE

I

Age Class
y

1
2
3
4
5
6

7
B

9

10

M

D

P-J

Total

�-11-

PHOTO-POINT REFERENCE

Project
Agency
CPS
CBS

_
_

Camera Point Stake (Orange-Topped)
Chalk Board Stake

CPS From:

_
Bearing

_

Distance

-------------~--------

CBS From CPS
Bearing
Distance

_
_

Camera Height (Top of CPS)

_

Distance From Lower Edge of Chalk Board to Ground

Sketch

-----------------------------

��-13-

July, 1974

Jon PROGRESS REPORT

State 0 f

COLORADO
--------~~~~-----------

Project No.

W-lOl-R-16

Work Plan No.

4

Game Range Investigations
Job No.

2

Job Title

Experimental Improvement of
Oakbrush on Deer and Elk Winter Ranges - Beaver Creek

Period Covered:

April 1, 1973 through March 3l~ 1974.

Personnel:

Roland C. Kufeld.

ABSTRACT
A contract was entered into with Colorado State University Statistics
Department to computerize and analyze pre- and post-treatment oak study
data. The analysis was not completed during this project segment so results will be presented in a later report.
A "Research Paper" by Roland C. Kufeld, O. C. Wallmo and Charles Feddema
summarizing 99 mule deer food habits studies, and ranking plants eaten by
relative value was published by the U.S. Forest Service, Rocky Mountain
Forest and Range Experiment Station (Res. Pap. RM-lll, 31 p.).

_

��-15-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH ON
DEER AND ELK WINTER RANGES - BEAVER CREEK
Roland C. Kufe1d

P. S. OBJECTIVE
To determine if deer and elk forage production and game use can be increased on overage Gambe1 oakbrush winter game ranges by spraying with
2,4,5-TP to induce sprouting.
SEGMENT OBJECTIVES
1.

To summarize and analyze pre- and post-treatment vegetation measurement
data.

2.

To write a report of the findings for publication in a scientific journal.

RESULTS AND DISCUSSION
A contract was entered into with Colorado State University Statistics Department to computerize and analyze pre- and post-treatment data from both the
Beaver Creek and Hightower MOuntain oak study areas. The analysis was not
completed during this segment so the results will be presented in a later
report.
A publication entitled "Foods of the Rocky Mountain Mule Deer" by Roland C.
Kufeld, O. C. Wallmo and Charles Feddema was published by the U. S. Forest
Service, Rocky Mountain Forest and Range Experiment Station (Research Paper
RM-lll, 31 p.). This was based on a literature review of mule deer food
habits completed during the 15th project segment. Ninety-nine food habits
studies were combined to determine plants normally eaten by the Rocky Mountain
mule deer (Odocoileus hemionus hemionus) and the relative value from a
manager's viewpoint based on the response deer have exhibited toward them.
Plant species are classified as heavily, moderately or lightly eaten on the
basis of their contribution to the diet in food habits studies where they
were recorded. A total of 202 shrubs and trees, 484 forbs, 84 grasses,
sedges and rushes, and 18 lower plants are listed as deer forage and
categorized according to relative value.

C, /(@

Prepared by ~
Roland C. Kufeld
Wildlife Researcher

��-17-

July~ 1974
JOB PROGRESS RErORT

Stat e 0 f

C::.:(::.;JL::.:O::.;R:=U.::ill:..,:O=-----''----

Project No.

W-lOl-R-16

Game Range Investigations

4

Job No._._--''---=-3--------~
__ --Experimental Improvement of
Job Ti t l,e_....:O~a::k~b~ru~~sh~~o!!n~D-=e:::e~r..1~-=E:=l:.!::k~a:=n~d~C~a!:.!t:..!t:.:l~e~Ra=n:!
ogh~t~o.!!w.=e:.=.r--=:-'M~o""un~t~a.

Work Plan No.

Period Covered:
Personnel:

April 1, 1973 through March 3l~ 1974.

Roland C. Kuf eld , Matthias Kn.Lesel.,John Arthur~ John Eer rdauo.l,o
,
Galen Burrell.

ABSTRACT
Two year post-treatment vegetation measurements were made in 600 permanent
meter square plots on the Hightower Mountain oak study area. Data analysis
is underway and results will be presented in a future report.
Cattle preference for burned oakbrush areas 2 years after treatment averaged
about 6 times that exhibited for areas treated by spraying and chaining.
Cattle use increased 18.7 cow days per acre on two 30 acre burned areas.
This was sIgnf.f Icant; of the 5 percent confidence level. Cattle use on two
30 acre sprayed areas increased 3.3 cow days per acre. A valid average
cattle use estimate could not be computed for the chained areas because of
the influence of an adjacent preferred burned area to one chaining~ and physical
barriers, which kept cattle from using another chaining. Because of these
factors it could not be concluded that chained areas were less preferred than
sprayed areas~ although. data indicated chainings were not more preferred.
A "Research Paper" by Roland C. Kufeld~ o. C. Wallmo and Charles Feddema
summarizing 99 mule deer food habits studies~ and ranking plants eaten by
relative value was published by the U. S. Forest Service, Rocky Mountain
Forest and Range Experiment Station (Res. Pap. RM-lll~ 31 p.).

��-19-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH
ON DEER, ELK AND CATTLE RANGES - HIGHTOWER MOUNTAIN
Roland C. Kufe1d

P. S. OBJECTIVE
To determine the extent to which deer, elk and cattle forage production
and game use can be increased and maintained by chaining, spraying and
controlled burning on overage Gambe1 oak winter game ranges.
SEGMENT OBJECTIVE
1.

To determine the extent of vegetative composition and forage production
changes, and the extent of deer and elk use changes which have resulted
from implementation of each habitat improvement method tested. Determine
if cattle exhibit a preference toward certain habitat improvement units.

METHODS AND MATERIALS
Post-treatment Vegetation Measurements
Two year post-treatment vegetation measurements were made in 600 permanent
meter square plots on the Hightower Mountain oak study area using the same
procedures employed during pre-treatment evaluations (Kufeld 1971). The
measurement period was July 9, 1973 through August 16, 1973. Prior to beginning vegetation measurements observers were trained in the use of vegetation measurement technique, and in identification of all plants on the
study area.
Determination of Nutrient Quality Changes
Samples of plant species designated by Kufe1d (1971) as "moisture indicator
plants" and used to reflect changes in vegetation moisture content during the
measurement period were collected in Units 5, 6, 7 and 8 between July 24, and
July 27, 1973, inc. Samples consisted of approximately 100 grams green weight
of each species collected throughout each habitat improvement unit. In units
5 and 7 samples were collected from those areas where it was certain burning
and spraying had been severe.
Samples will be compared with pre-treatment samples of the same species
collected in unit 6, a control area, to determine changes in nutrient quality
of forage resulting from chaining, burning and spraying. Standard laboratory
procedures will be used to give an estimate of digestible dry matter, digestible

�-20-

protein~ digestible energy and trace mineral values. These procedures
will involve the use of fistulated goats to arrive at estimates of digestibility. Laboratory analysis has not been completed at the end of the 16th
project segment, so results will be presented in a later report.

Photo Point Photographs
Post-treatment photos were taken at permanent photo points established in
1970, prior to treatment. These show effects of spraying~ burning and
chaining during the first summer following treatment. A black and white
photo, and a 35 mm color slide was taken at each photo point with the same
cameras used to take pre-treatment photos.

Post-treatment Deer, Elk and Cattle Use Measurements
Accumulated deer and elk pellet groups were removed from all pellet plots
on September 3, 1973. Groups deposited during the winter of 1973-74 will
be counted in May, 1974.
When plots in units 1, 5~ 6~ 7, 8 and a portion of 3 were cleared a record
was kept of the number of cow chips in each plot in order to provide an
index to cattle use. Cow chip counts in units 2, 4 and the remainder of 3
were delayed until October 3~ because these units are in a different cattle
allotment~ and cattle were still grazing until the end of September. Only
those chips which could be identified as having been deposited during the
summer of 1973, were counted.

RESULTS AND DISCUSSION

Post-treatment Vegetation Measurements
Data have been summarized and a contract has been entered into with Colorado
State University Statistics Department to computerize and analyze pre- and
post-treatment data from both the Hightower Mountain and Beaver Creek oak
study areas. The analysis was not completed during this segment, so the
results will be contained in a later report.

Post-treatment Cattle Use Measurements
An analysis of 1970 and 1973 cattle use data to determine cattle preference
two years after treatment for areas burned, sprayed and chained is presented
in Table 1.
Control Areas
Data show cattle use increases in both units 4 and 6~ although neither increase
is Significant at the 5 percent confidence level. Much of the increase in

�-21-

unit 6 is believed due to movement of cattle around unit 7 and between
units 5 and 7 to which cattle were attracted.
Unit 6 lies directly between
units 5 and 7. Thus, there was very little, if any, actual change in
cattle use on the control areas.

Burned Areas
The most preferred treatment by cattle two years after treatment was
definitely burning.
Units 1 and 7 showed the greatest increases in cattle
use after treatment, and these were significant at the 5 percent confidence
level. Cattle use in units 1 and 7 increased 15.2 and 22.1 cow days per
acre respectively, and averaged 18.7. Cattle preferences for these units
were also obvious from field observations.
Preference for the burned areas
was approximately 6 times greater than for the sprayed or chained areas.

Sprayed Areas
One spray area (unit 5) showed a substantial increase in cattle use which
was significant at the 5 percent confidence level as a result of spraying.
The other (unit 2) showed a slight decrease~ although not significant at
the 5 percent level. According to the data, which show an average increase
of 3.3 cow days use per acre in the sprayed areas, spraying would appear to
be the second most preferred treatment by cattle.
However, due to circumstances
described in the following discussion on chaining it is not valid to conclude
that cattle preferred the sprayed areas more than the chained areas at the
end of the second year following treatment.

Chained

Areas

No use was recorded in unit 3 in 1973, probably due to the fact that it is
on the south side of Buzzard Creek and high water during 1973 kept Buzzard
allotment cattle on the north side. Also, unit 3 is difficult for both
Buzzard and Porter allotment cattle to find. The almost impenetrable oak
thicket in unit 3 was opened substantially by chaining, and it is felt
cattle use would have occurred in 1973, if cattle had been able to get to
the area.
Cattle use in unit 8 dropped from 18.3 cow days use per acre in 1970 to 8.7
in 1973. Although this drop was not significant at the 5 percent confidence
level it appeared that cattle were simply drawn off unit 8 into the more
preferred burned area of adjacent unit 7. This occurred even though conditions in unit 8 were improved for cattle by chaining.
Because of the statistical insignificance of the drop in unit 8, and inability
of cattle to find and get into unit 3, it is not valid to conclude that
chained areas were any less preferred by cattle than sprayed areas.
Had it
not been for the existence of the highly preferred burned areas cattle would
have probably exhibited a greater preference for areas treated by spraying
and chaining.

�Table 1. Cattle use of burned, sprayed, chained and control areas before (1970) and two years after
(1973) treatment.

!/

Cow Days Use
Per Acre During
1970 Season 1:./

Cow Days Use
Per Acre During
1973 Season 1/

Change in Cow
Days Use Per Acre
From 1970 to 1973

Significance
of Change
at d... .05 level

Unit

Treatment

4

Control

0.0

3.0

+ 3.0

No

6

Control

1.3

6.5

No

4 &amp; 6

Control

0.7

4.8

+ 5.2
+ 4.1

1

Burn

4.4

19.6

7

Burn

9.6

31. 7

1 &amp; 7

Burn

7.0

25.7

+ 15.2
+ 22.1
+ 18.7

2

Spray

8.7

5

Spray

2 &amp; 5

No
Yes
Yes
Yes

I

N
N
I

7.4

-

1.3

No

3.0

10.9

Yes

Spray

5.9

9.2

+ 7.9
+ 3.3

3

Chain

0.0

0.0

0.0

No

8

Chain

18.3

8.7

- 9.6

No

Chain

--=-r;J

3 &amp; 8

l/Burned,

No

sprayed and chained areas were also seeded.

~/Cow defecation

rate is 11.5 chips per day as determined by Ju1ander

(1955).

3/
- Post-treatment data for all units have been adjusted for differential cattle stocking rates which existed
during 1970 and 1973. Data have also been adjusted for differences in stocking rates between two cattle
allotments which comprise the study area. Units 1, 5, 6, 7, 8 and a portion of 3 are in Buzzard allotment,
and 2, 4 and the remainder of 3 are in Porter allotment.
i/Data for chained areas were not averaged. High creek water and difficulty for cattle in finding Unit 3
caused the absence of cattle use in unit 3 during 1973, even though the area had been substantially improved
for cattle by chaining.

�-23-

LITERATURE REVIEW
A publication entitled "Foods of the Rocky Mountain Mule Deer" by Roland
C. Kufe1d, O. C. Wa11mo and Charles Feddema was published by the U. S.
Forest Service, Rocky Mountain Forest and Range Experiment Station (Research
Paper RM-lll, 31 p.). This was based on a literature review of mule deer
food habits completed during the 15th project segment. Ninety-nine food
habits studies were combined to determine plants normally eaten by the
Rocky Mountain mule deer (Odocoileus hemionus hemionus) and the relative
value from a manager's viewpoint based on the response deer have exhibited
toward them. Plant species are classified as heavily, moderately or lightly
eaten on the basis of their contribution to the diet in food habits studies
where they were recorded. A total of 202 shrubs and trees, 484 forbs, 84
grasses, sedges and rushes, and 18 lower plants are listed as deer forage
and categorized according to relative value.
LITERATURE CITED
Julander, Odell. 1955. Determining grazing use by cow chip counts.
Range Manage. 8(4) :182.

J.

Kufeld, Roland C. 1971. Experimental improvement of oakbrush on deer,
elk and cattle ranges - Hightower Mountain. Colo. Div. of Game,
Fish and Parks. P-R Project W~101-R-13, WP-4, J-3, Game Research
Rept. July. Part One. pp. 23-86.

Prepared by

~CI%/rhJ
Roland C. Kufeld
Wildlife Researcher

��-25-

July, 1974

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~--~--~---

Project No.

W-101-R-16

·Work Plan No.

5

Game Range Investigations

Job Title

Bighorn Sheep Range Inventory

Period Covered:

April 1, 1973 .toMarch 31, 1974

Personnel:

Harold R. Shepherd, Roger Bradshaw, Denis Davis, JamesL.
Shepherd son , Alan Titche, and John Weins.

ABSTRACT

A range inventory of the Buffalo Peaks Bighorn Sheep study Area, near
Fairplay, Colorado was completed. The resulting data were compiled and
summarized by transect and vegetation type, and a map was prepared showing the distribution, extent and composition of vegetation and the loca~
tion and extent of seasonal ranges in the study area.

��-27-

BIGHORN SHEEP RANGE INVENTORY
Harold R. Shepherd

P. S. OBJECTIVE
To inventory, contrast and evaluate certain range attributes of a productive
and an unproductive bighorn sheep habitat.
SEGMENT OBJECTIVES
1.

Determine the distribution and extent of vegetation types in relation
to physical features of the habitat on ranges with good (Trickle Mountain)
and poor (Buffalo Peaks) bighorn sheep production.

2.

Estimate percentages plant species composition, frequency, crown cover,
litter, rock and bare ground on ranges with good (Trickle Mountain)
and poor (Buffalo Peaks) bighorn sheep production.

3.

Organize, compile and summarize data by vegetation type.
DESCRIPTION OF AREA

The study area is located on East and West Buffalo Peaks on the boundary
between Park and Lake counties, Colorado in South Park, and it is 16 miles
south and 7 miles west of the town of Fairplay (Fig. 1). The county boundary
is also a common boundary between the Pike and San Isabel National Forests.
Generally, the study area is delimited by timberline at about 11,800 feet
around the two peaks, and it extends upward to the top of West Buffalo Peak
at an elevation of 13,326 feet.
METHODS AND MATERIALS
Establishing Study Area Boundaries
It is the consensus of Division personnel and others who have studied and
observed the Buffalo Peaks bighorn sheep that in recent years the herd has,
for the most part, restricted itself to that portion of the Buffalo Peaks
area above timberline. Consequently, for the purposes of this study, timber
line or a few hundred feet below timberline on the extreme eastern edge of the
area marks the boundary of the study area.
Preparation of Photo Mosaics
A photo mosaic of the study area was prepared from 9X9-inch color aerial
photographs, and the study area boundary was delineated on it in red ink.
Photographic copies of the master mosaic were made, and from one of these was
prepared a cloth-backed folding mosaic map used in the field during transecting
activities.

�:(

I

W

o
r

BUFFALQ
n:CHORN SHEEP

PEAKS
STUDY

Po",

Counties,

ana Chaffee

Son 'socer and P,ke Nat!
t)t'o!oqrCOhed

for

Colo. Division

AREA
Colo
Forests

N

J

of W'k1lrte

By
i&lt;"(~n

and

"r8[&lt;III)(j

Ly

J~g0~1'Jr;

I~S$OC.

H.R
Sco!t~

Denver.

Colo., Sept 7. 1971

Shepherd 8, J.L
I 15840

Shepherdson

Fig. 2. Mosaic showing black and white and color aerial photograph
Bighorn Sheep Study Area.

coverage

available

of Buffalo Peaks

�-31-

Preparations

for Transecting

A field crew of four game management students reported for work June 4.
The first seven days of the field season the crew was trained in plant
identification, using the field herbarium of mounted plants and color
slides of plants previously collected in the study areas. June 13th,
the crew left Fort Collins for Buffalo Peaks, and the next four days were
devoted to clearing snowdrifts and down timber from the road up Lynch
Creek to a permanent camp site just below timberline in the cirque between east and west Buffalo Peaks. A condition of permission from the
Forest Service to conduct the range inventory and to camp within the
cirque was that we would establish locked gates blocking the lower end
of the road up Lynch Creek to the public and that we would erect signs
declaring that the country behind the signs was a bighorn sheep study
area and off limits to the general public.
This was done. A temporary
camp was established at the old saw mill site near the junction of the
Lynch Creek and Rough and Tumbling Creek roads.
Actual transecting was begun east of the Buffalo Peaks cirque on June 17
and was completed September 6.
Before and during the transecting of a vegetation type, its boundary as
previously delineated under stereo was checked against on-the-ground
conditions and any desired modifications made on the photos.
In only
a few instances were changes made, and these were slight.

Transecting

Vegetation

Types

Because of the extremely rocky and precipitous terrain and absence of
roads, vehicular travel within the study area was impossible.
In fact,
only the perimeter of the area could be reached by automobile and this
in only five places, three on the north and two on the south sides of the
area.
Consequently, the field crew was transported from base camp in the
Buffalo Peaks cirque to the four other avenues of approach.
From these
the entire area was covered on foot. The long driving time required from
base camp to the two southern approaches to the area, made the use of these
approaches less economical of time for all except the transecting of nearby
areas than the base-camp approach.
Consequently, most of the area was
reached on foot from the base camp.
Each crew member carried a back pack
on a pack frame in which he transported his share of the equipment load
besides items of personal gear. The transecting equipment and supplies
included two Daubenmire plot frames, two 300-foot steel tapes and reels,
four steel stakes, three clipboards, two hand levels, three compasses,
two ball peen hammers, a chalk board, aerial photos and a folding aerial
mosaic map of the study area, pencils, sharpie pens, Kohinoor pens and
ink, a small metric scale, a protractor, several plant manuals, a plant
press with blotters and folders, hand lenses, a plant collecting trowel,
three field glasses, a 35 rom reflex camera and film, and a supply of data
forms.
In addition to transecting equipment, each man carried such personal gear as a down jacket, rain poncho, thermal underwear, camera equipment, emergency first aid and survival items and his lunch.

�-32-

The Wildlife Researcher as crew leader, led the crews to sampling sites
located, as previously described, on aerial photos.
The location of each
sample site was determined approximately by reference to features of the
terrain recognizable on the ground and in the photos.
These were such
features as rock outcrops, individual or clumps of trees, patches of
willows, different kinds and colors of rock and soil. gullies. streams,
rock garlands, rock streams, rock stripes, rock polygons and circles
comprising patterned ground, etc.
Each sample site became one end of a 300-foot transect placed parallel
to the contour.
Each end was marked with a rock cairn.
Steel stakes
were driven at either end and a 300-foot steel tape stretched between them.
Each sample site was photographed by the crew leader along the transect
from one of its ends, using a 35 rom Exakta camera and Kodacolor film. A
small slate on which was written with white chalk the transect's number
and the type number and date was included in the picture foreground.
Two men comprised a transect crew. At the outset. each took turns reading
and recording transect data so that each would be familiar with the procedures and the vegetation.
Later, as differences in skills and aptitudes
became evident, certain individuals were designated readers and others
recorders.
But throughout the investigation, members of a team collaborated
with and double-checked one another's plant Ldent.Lf Lca t Lon, canopy-coverage
estimates and physiognomic descriptions.
The crew leader attemped to so organize and direct the order of transecting
that the two crews, each reading a different transect. would be near enough
that he could supervise each.
Transects were placed along the contour so that they could be kept within
the same general soil type and, as a consequence, changes in vegetation
composition would be kept to a minimum.
Vegetation was sampled systematically
along the transect at five-foot intervals with a Daubenmire canopy-coverage
frame. When transect data from the Trickle Mountain study area had been
tested the previous summer with species-area curves it had been found that
40 plots per transect did not yield acceptable accuracy in some types but
that 60 plots were sufficient in most cases.
Similar results were found
for selected Buffalo Peaks data, so 60 plots were read for each Buffalo
Peaks transect.
The Daubenmire (1959) canopy-coverage
plot frame (Fig. 3) used for the Trickle
Mountain inventory was used for the Buffalo Peaks inventory.
Its modifications
and use were described (Shepherd 1973) in the previous report.
Upon establishment of a transect, the crew leader marked its location within
the vegetation type on the aerial photograph.
Using a rapidograph pen, a
short, red line to scale was marked on the photograph to indicate the pOSition,
length and bearing of the transect.
A numeral indicating the number of the
transect in its type was placed at one end of the mark designating the transect.

�-33-

�-34-

Using the system described by Daubenmire (1959), the two men of a transecting team esti.mated and recorded the canopy-coverage per species and
the ground-surface data within each of 60 plots of a transect.
These
data were recorded on prepared forms (Fig. 4). When all of the plots
along a transect had been read, physiognomic data for that transect and
its stand in the immediate vicinity were recorded on a prepared form
(Fig. 5). The Braun-Blanquet system of vegetation analysis and description (Kuchler 1955) was used to record physiognomic data for each species.
The two men of a team collaborated in assigning the physiognomic descriptions.
Consequently, they compared each other's estimates of sociability
and coverage, thus increasing the uniformity and reliability of their
estimates.
Transecting was completed September 6. The crew was brought to the Research
Center where they began compiling and summarizing the summer's collection
of data from 12,060 plots and 201 transects.

Data Compilation

and Summarization

A form was prepared for each transect, summarizing the data for all plots
along the transect (Fig. 6). To the original physiognomic data form was
stapled a 3X5-inch color photograph, showing the transect's location, the
vegetation it sampled and the characteristics of the ground surface and
topography of the site.
Data for all the transects within a type were displayed together in an
abbreviated, simplified manner on a form prepared for the purpose so that
comparisons could more readily be made between types. Using this information together with physiognomic and photographic data, similar types
were combined to form larger areas of vegetation, and these were given names
descriptive of the three or four dominant plant species characterizing them.
All of this information was assembled into large loose-leaf binders to provide an organized file of data with which to prepare a final report for
publication of the results of the combined range inventories of Trickle
Mountain and Buffalo Peaks study areas.

RESULTS AND DISCUSSION
The results of the comprehensive range inventory for both the Trickle Mountain
and Buffalo Peaks bighorn study areas will be presented as a final report in
the next project segment.
Since the data collected, although in summarized
form, is too voluminous to justify inclusion in a Progress Report, only a
vegetation type map of the Buffalo Peaks study area (Fig. 7) and detailed data
(Fig.8) for one of its plant associations comprising the lambing range on East
Buffalo Peak are shown here.
The seasonal ranges on the Buffalo Peaks Bighorn Study Area (Fig. 7) are comprised of 197 acres of summer range, 261 acres of winter range and 44 acres
of lambing range. Other range included within the boundary of the study area
is used much less intensively or seasonally by sheep and is termed "other
range".
It comprises 2106 acres.
Total acreage of the study area is 2608
acres.

�-35CANOPY-COVERAGE-PLOT

DATA FORM
Sheet No.

Stand No.

':1.7-/

Stand Location:
Aspect &amp; Slope

Veg. Type

PeQXS

Study Area Name Bv{(%

Date
Town.

7/g,77.J
r I
~-

335°,

Slope

_

No. (from map) 27(Summer

RAn~J
Examiners Veins. ];tc..he

/s- ~

Photo No.
Range
Soil:

:/,0

&amp;

~

Section~

_
Series.

_

Class.

_

j

Physiognomy
Transect:

,

&amp; Structure

No •

[lev. ~

_

...:...J

Location

J ').J 'l60

"

.,
&amp;

3?S
Bearing,..-::::....:.):::...::
0

.,

11

_

e:

Plot NO"_--L __
Canopy Coverage

I

Species
Class
No.

PAS£.
A ROB

/

:J.

Class
Hidpoint %
~.5
!
t
/5.0
/5. 0

TRNA

:1

ARAR

I
J

1.5

{

~.5

8'

40.0

PoA
FEOvB

Totals
Plot No.

'1..

Ground-surface
&amp; Vegetation
Point Estimates
Vascular
Hoss
I Rock Litter
Bare
Plant %
Lichens %
%
%
Ground '7,

I

75

:z:;

s:

2.
Ground-surface
&amp; Vegetation
Point Estim2tes
Vascular
:loss
Roc Litter
Class
I Bare
Plant %
%
Lichens
%
%
%
~1idpoint % Ground

Canopy Coverage
Species

Class
No.

I

PASf
ARa/3

I

2.S

z

/5.0

I

TRNA

)

~.S

t

ORAL

I

~ ..)

ERst
}&lt;'OOE

I

POA

I

~.5"
:l.S

8

30.0

Total~
Fig.

I

i
SO
'-.5

:2..S

4. Canop,y-coverage data field form and sample plot data.

,"-s-

�-36VEGETATION-TYPEDATARECORD
Sheet
Study

Hza.ks

2. 7

Type Ho.
Stand

Buffa.lo

Uame

AYe:)..

Aspect

Transect

Stand

slop(/!

-1-0

/181&gt;

2. 7 - I

No.

3;, 5'0

and Slope

Location

Cs u m mar:

No.

,..a.ngcZ)

-~---

'I'r'ans ec t No. __ i.

2 D

W(/!~f ELL f-fa/o

_

--------

Pea.k

------------------------------------------------------------------Transect:

Bearing

Aerial

Photo

Soil

&amp; Ground

crr cle s

No.

By

Floristic
Composition
(S..B.ecies)

S" I&gt;

Length

IS- - 6

Surface

o",d

Recorded

'33

rro e ]:

Wttins

Stand

.&lt;icvndy

Photo
a.Vld

lao-WI

360 .f-l.

No. Plots

No. TY- '27 TR-lDate
pafferYlfZcI

qroUHd

60
7-16-- 73
I&lt;J

rock::

.:s1y-tra.Wlsal1a

Physiognomy
and
Structure

]j'f-che
Cover
and
Sociability

Remarks

-----

PA s e:

IUp

-t.3

GtZl..l"'"

+urb/n(X

TRNA

HIp

t. 3

Sh(Z(Z p

sea l- Dee unrrzc/

AROB

Hip

1.3

-F'yt?ou.en t lv-

ARA~

!-lIp

+.3

Glp

« z.

FEOV13

G/ p

+.2

ORAL

Hip

1.3

I

HI,..

+.1

KOBE.

/.2-

PORU 2-

G/p
J-I /.£

PHCO .3
GETU

H te
H te

Moss

LIp

SARH

Hlr
Gly

+.1

Hlr

POA

ERS

SP.

AGSC

3

+,3

-+.3
.Q.3
f.3
f.2

PRAN 2

HI,..

+.2
f. I

TR..S P

GJr

+,2-

MEVI

Hly

T'/

ER.N/3

Hlr

[)~AV

Hlr-

+./
+. I

TARAXACUM

HI"..

ARFE

ANSEG

Hly

Mea,A

Hlr

~CL

HIJ-

M ~

HeNI

pav I

Fig.

4-

HII'
HIp

5. Physiognomic

-fGlWl ;J..Y"a.:vza.

-to I
+. I
T. I

+. I
+. '3
+.2...
data field form and sample transect dataG

-------

�-37TRANSECT SUMMARY
Page 1
Date 7-31-73
Photo No. 15-8
Study Area Name Buffalo Peaks - Lambing Range
Stand No.
Transect No. 4
Veg. Type ...,:4..:,.9_-=-:-:-......- __
----,,....,..
_
~-----Aspect &amp; Slope.-=1~4~0
~
_ Soil &amp; Ground Surface
Physiognomy &amp; Structure

Ground-Surface &amp; Vegetation Point Estimates
Moss
Bare %
Litter%
Rock%
Lichen %
Ground
Total

,

') &lt;;.()

20.83

Ave.

,

Q&lt;;;f\

f\

1 Q')e;

17.92

30.83

0

30.42
_

_

iotal
Total
Times
Canopy
Coverar::eOccurred

Species

Vascular
Plant %

l()7e;

Total No. Canopy-Coverage Entries __~3~3~8
Average No. of Species/Plot ~5..:.
•..:.6~3

_

No. Plots ---=--=---60

AVArap.:e Average
Fr-equency Canopy
~average
~

Co:;;pos-

;C'" ition 1&gt;

GRAMINOIDS
10

1f,e; n

?f,

I. '1

Kabresia be11ardi

790.0

58

96.67

13.17

24.85

Paa spp.

152.5

46

76.67

2.54

4.79

1107.5

130

216.67

18.46

34.83

Trifolium nanum

905.0

'l2

R6 67

s .na

28.45

Oreoxis a1pina

507.5

59

98.33

8.46

15.96

Potenti11a

147.5

24

40.00

2.46

4.64

Geum turbinatum

170.0

14

23.33

2.83

5.34

Trifolium dasyphy1lum

20.0

3

5.00

.33

.62

Arenaria abtusiloba

120.0

10

16.67

2.00

3.77

Phlox caespitosa

62.5

15

25.00

1.04

1.96

Saxifraga rhomboidea

10.0

4

6.67

.17

.32

Paronychia

22.5

4

6.67

.38

.72

2.5

1

1.67

.04

.08

e10ngatum

17.5

2

3.33

.29

.55

_.HgIL1.Q.P1!P..IlliS
pygmaeus

25,0

5

R 11

.42

.79._

2010,0

101

11?1.h7

11 'l()

63.21

Moss

f,?

e;

, &lt;;;

I.

1 Of,

Total

3180.0

338

53.00

99.99

Fig. 6.

Sample transect summary form and data.

~('rihnf&gt;ri

A

Sub-Total

')

'1 '1

7&lt;;.

e;

FORBS

Arenaria

rubricau1is

sessilif10ra

fendleri

Eritrichium

Sub-Total

')&lt;;;

f\f\

563.34

t

,

f\

�VEGET A TlON
BUFFALO

PEAKS

AND

SEASONAL

BIGHORN

SHEEP

RANGES
STUDY AREA

J
.:1

I

W
00
I

~~~~;~fGETU·Dl:o(·C.ufX)

FOlliS

LEGEND

CJ
t£I]
IIJI

A,ropyron

UOMU
CUEX
OECA
FIOV
FETH
KOIf

'ramus

',p.

Cor ••.•

pp.

I'OA
OTHER

RANGE

.cribn.,;

D'J(hornp.io &lt;••• pilol.
Fo.II,I(O

•.••ina

fulu(o
Kobt..&gt;a

Ih",,",;

'0••

SUMMER

LAMBING

Fig. 7.

RANGE

Nj

RANGE

IANGE

Scale

"CLio

Achilln

AN'''3

"""11".';'

CACO

Arnico ,or4If,lio
C. •••
min.
(o,41f,lI.

.•••
eo
CAU

bell.,eli

pp.

BROWSE

WINTER

~

ADS(

CI'A2
DO",
IPAN
GETU
HEVI
MEel

C.llha
Ci,.jym

,.",,1, ••

"rYU.tie

,." ••••••
10
,.lIielvm

00",011,.,,,
Epilobil"r.

,ulchallum

.",".'I~

h"binot"",
H,t.,oth,ca
•••
1110
••
0.1,1191

"'-,I,n,h"

CH'AII

ChrY50lhamftU5

ORoe

O.yo.

oclopelola

ORAL

Or.oxil

olpino

'IAR

Pinv.

o.illolo

~D12

Pol,ntillo

div.r,lf,lia

,IEN

Picoo

.ft,elmoftfti

Pol ••.•
lillo

lARAX
rlOA
TINA

T.".o(lIm

"'"

porry;

hUlico50

'OTlt
liMO

Ribft

moftlig."um

SAANA

Soli.

o"glorum

Populul

tte/nllloid

••

oftliplalla

SAGLG

Soli.

glauco

glob.e5Uft5

SA,LM

Soli.

plofti'olio

moftica

1:IS140

Vegetation-type map of Buffalo Peaks Bighorn Sheep Study Area.

pony;

U,.•.

cilia,a

.,p.

"".,.I1",,"

hi f.B",,,, d••
Trifoli"",
T,if.'iull'l

,.".,1

VACAI

V.I••iono

,o,i'ol.

lOCI(

(h\oJd.,fi ••••1.1,", Ser ••• CIIH)

�.•
39 .•
ASSOCIATION

SUMMARY
Page 1

35
Name
Association
Types.
4~9~(6~t~r~a~n~s~e~c~t~s)~

----------------

Seasonal

TRNA-KOBE

------------------------------------- _

Range_'__ -=Lam~b~i~n~g~Ra~n~ge~
Ground-Surface
Bare
Ground %

Rock %

_

&amp; Vegetation Point Estimates
Moss
Lichen %

Litter %

Vascular
Plant %

Total

4125

9600

15,000

275

7000

Ave.

11.46

26.67

41.67

0.76

19.44

Entries

2_O_O_6

_

5_._5_7

_ No. P 1ots.

360

Total No. Canopy-Coverage
Average No. of Species/P1ot.

Species

Total
Total
Times
Canopy
Coverage Occurred

_

Average
Frequency
%

Average
Canopy
Coverage

%

Composition %

GRAMINOIDS
Kobresia be11ardi

3652.5

287

79.72

10.14

20.86

Poa spp ,

1268.0

283

78.61

3.52

7.24

440.0

101

28.06

1.22

2.51

Festuca ovina

447.5

55

15.28

1.24

2.55

Carex spp.

292.5

28

7.78

0.81

1.67

Trisetum spicatum

197.5

54

15.00

0.55

1.13

He1ictotricon

17 .5

7

1.94

0.05

0.10

6315.5

815

226.39

17.53

36.06

291

80.83

11.19

23.02

Oreoxis alpina

4027.5
2905.0

292

81.11

8.07

16.60

Geum turbinatum

2155.0

120

33.33

5.99

12.32

Saxifraga rhomboidea

72.5

29

8.06

0.20

0.41

Potentilla

377 .5

86

23.89

1.05

2.16

Agropyron

scribneri

mortoniana

Subtotal

FORBS
Trifolium nanum

rubricaulis

Arenaria

obtusi1oba

315.0

44

12.22

0.88

1.81

Trifolium

dasyphy11um

132.5

28

7.78

0.37

0.76

262.5

50

13.89

0.73

1.50

Draba aurea

2.5

1

0.28

0.01

0.02

Primu1a an~ustifo1ia

12.5

4

1.11

0.03

0.06

Phlox caespitosa

!rotal

Fig. 8.

Association

summary form and sample canopy-coverage

data.

�-40ASSOCIATION SUMMARY

Page 2

Associat1on~-

_
Total
Canopy
Coverage

Total
Times
Occurred

C1aytonia megarrhiza
Bessya a1pina

20.0
7.5

3
3

0.83
0.83 -

0.06
0.02

0.12
0.04

Hap10pappus pygmaeus

50.0

15

4.17

0.14

0.29

Ste11aria longipes 1aeta

37.5

15

4.17

0.10

0.21

Artemisia spp.

30.0

12

3.33

0.08

0.16

Swertia radiata

2.5

1

0.28

0.01

0.02

Saxifraga f1age11aris

7.5

3

0.83

0.02

0.04

Cirsium pa11idum

7.5

3

0.83

0.02

0.04

Taraxacum spp.
Androsacea septentrionalis glandu10sa

32.5
2.5

8
1

2.22

0.18

0.28

0.09
0.01

0.02

Potentilla diversifo1ia-

2.5

1

0.28

0.01

0.02

Silene acaulis

1

Cerastium spp.

2.5
22.5

9

0.28
2.50 .

0.01
0.06

0.02
0.12'

Senecio spp.

2.5

1

0.28

0.01

0.02

Erysimum niva1e

2.5

1

0.28

0.01

0.02

Chionophila jamesia

7.5

3

0.83

0.02

0.04

Potentilla diversifo1ia

2.5

1

0.28

0.01

0.02

Paronychia sessi1if1ora

127.5

21

182.5

43

5.83
11.94

0.35

Arenaria fend1eri

0.51

0.72
1.05

Eritrichium eLongatum

17.5

2

0.56

0.05

0.10

Eriogonum piperi

5.0

2

0.56

0.01

0.02

10835.0

1094

303.89

30.10

61.91

355.0

97

26.94

0.99

2.04

355.0

97

26.94

0.99

2.04

17505.5

2006

557.22

48.62

100.01

Species

Subtotal

Average
Average ComposFrequency Canopy % ition
%
Coverage
%

MOSS-FERNS-LICHENS
Moss
Subtotal

Total
Fig. 8.

Association SUmmary form and sample canopy-coverage data (continued).

�-41-

BIBLIOGRAPHY
and
LITERATURE CITED
American Society of Photogrammetry. The manual of photogrammetry.
Soc. Photogrammetry. Menasha, Wis. 876 p.

Amer.

Anderson, A. E., D. E. Medin, and D. C. Bowden. 1971. Mule deer fecal
group counts related to site factors on winter range. (Unpublished
manuscript).
Daubenmire, R. 1959. A canopy-coverage method of vegetational analysis.
Northwest Science. 33(1):43-64.
1970. Steppe vegetation of Washington.
Tech. Bull. 62. 131 p.

Wash. Agr. Exp. Sta.

Drew, W. B. 1944. Studies on the point-quadrat method of botanical
analysis of mixed pasture vegetation. J. Agr. Res. 69:289-297.
Fayle, D. C. F. 1959. The point contact method as a three-dimensional
measure of ground vegetation. The Forestry Chronicle. 35(2):135-141.
Harrington, H. D., and L. W. Durrell.
Sage Books. 203 p.

1957.

How to identify plants.

Kuchler, A. W. 1955. A comprehensive method of mapping vegetation.
Annals Ass. of Amer. Geog. 65:404-415.
Loveless, C. M. 1967. Ecological characteristics of a mule deer winter
range. Colo. Game, Fish and Parks Dept., Tech. Pub. No. 20. 124 p.
Shepherd, H. R. 1973. Bighorn sheep range inventory, pp. 25-39. In
Game Research Report, July, 1973. Part 1. Colo. Div. Wildlife.
U. S. Forest Service. 1969. Forester's guide to aerial photo interpretation,
Agr. Handbook No. 308. 40 p.
U. S. Forest Service.

Prepared by

B.P.I. workbook.

~~£4~

;dId R. Shepherd
Wildlife Researcher

~

Washington, D.C.

47 p.

��-43July, 1974
JOB PROGRESS REPORT

State of --------~~~~~--~-COLORADO
Project No.

W-101-R-16

Work Plan No.

7

Job Title Reprint Book:

Game Range Investigations
Job No.

1

~~----~-----------------------

Winter Guide to Native Shrubs of the Central Rocky Mountains

P~riod Covered: July 1, 1973 to March 31, 1974
Personnel:

William T. McKean and Paul H. Neil.

ABSTRACT
No work was accomplished on this job during the nine months period except
that in a conference with Mr. Charles Hje1te, a decision was made to reproduce illustrations by photographing pages of the first edition book
wherever they are of satisfactory quality. Some illustrations will need
to be done over from new specimens or from herbarium specimens.

��-45-

REPRINT BOOK: WINTER GUIDE TO NATIVE
SHRUBS OF THE CENTRAL ROCKY MOUNTAINS
William T. McKean and Paul H. Neil

P. S. OBJECTIVE
To revise and reprint the book: Winter Guide to Native Shrubs of the
Central Rocky Mountains.
SEGMENT OBJECTIVE
To refurbish art work on book illustration pages.
METHODS AND MATERIALS
The original paste-up sheets for each page of illustrations were salvaged
and stored after the first edition was printed. Labels have since then
loosened and need resticking. Black and white prints need retouching or
replacement. All the beaver board sheets on which materials are mounted
need cleaning up or replacement. This job will be worked upon by a student
assistant, under supervision, at odd times during the segment 1973-74.

RESULTS AND DISCUSSION
The only work accomplished was a conference with Division Publications
Chief, Charles Hjelte. Mr. Hjelte examined the paste-up sheets and advised
that most of the art work be done by directly photographing the book pages
rather than trying to refurbish the sheets. In cases where the photo prints
(half-tones) are of too poor quality they should be rephotographed from new
specimens.

2l?~Ie: a~i

Prepared by 1Kt-!{~~~.I&lt;_:;rc
William T. McKean
Wildlife Researcher

��-47July 1974
JOB PROGRESS REPORT

Stat e 0 f .

C=-o::...:L::.:o:..:R.:=:\D::::.-~O:...__'_
_

Project No.

W-101-R-16

Work Plan No.

6

GaIne Range Investigations
Job No.

1
----~--~----------~~------~

Job Title

Game Management Unit· Inventory

Period Covered:

April 1, 1973 to March 31,1974

Personnel:

Bertram D. Baker, William T. McKean, and Paul H. Neil.

ABSTRACT
Broad descriptive and wildlife speciesmanagemeIlt information were compiled
separately for Wildlife ManageinentUnits3l
·(Roan Creek) and 32 (Parachute
Creek). Subjects discussed included: Unit Description (Boundaries, Size,
Physical Features and Climate); Landownership Status; Land Use Status;
Human Populations (by County); Wildlife Species Checklists; Hunting Pressure,
Harvests and Seasons; Narrative and Map Descriptions of Game Species Distribution and Abundance; Introductions. of Game Species;·Census Areas and Routes;
Habitat Restoration Projects; Management Problems Checklist; and Pertinent
Research References by Game Species and Author •

•'i. •.••

��-49-

GAME MANAGEMENT UNIT INVENTORY
Bertram D. Baker~ William T. McKean and Paul H. Neil

For the second time, progress on this job was delayed. Bertram D. Baker
was transferred to Project W-38-R-28 (wp 18, Job 1) after working a little
over one month on the 1973-74 segment of this project. On July 1, Paul
H. Neil joined the project as Conservation Aide. He has served continuously
since that time.

P. S. OBJECTIVE
To compile broad descriptive and wildlife species management information
by wildlife management unit.
SEGMENT OBJECTIVES
1.

Describe Wildlife Management Units by name, number, boundaries, size,
physical features, and climate.

2.

Inventory by unit the cultural and physical characteristics including
landownership status, land use status, and human population.

3.

Inventory by unit wildlife characteristics including species lists,
distribution and abundance, harvest data, introductions and/or past
species records, census areas and routes, and research and/or management studies bibliographies.

4.

Inventory habitat (range) characteristics of vegetation types~ vegetation condition and trend, restoration projects, and transect records,
all from existing sources.

5.

Inventory present and future management problems by unit.

6.

Test application of any item of information compiled or recorded in steps
1-5 above in pilot computerized map project now in operation.

METHODS AND MATERIALS
Five steps were necessary to produce the desired information about each
Wildlife Unit.
1.

The priorities of units were determined, based upon recommendations of
field and administrative personnel.

2.

Working with sources in Fort Collins and Denver~ as much data as possible
concerning each item were assembled and condensed (See APPENDICES A and B).

�-50-

3.

First-draft copies of descriptive material were submitted to Northwest Regional personnel and to knowledgeable Denver office persons
for review and correction.

4.

Group meetings were arranged with field personnel and others to make
pencilled corrections on 1/2"/mile species distribution maps and to
make refinements to the narrative.

5.

As of this writing the above mentioned meetings have not been held
but will be within the first few weeks of the next segment. After
said meetings are held, final revisions to narratives and maps will
be made and all information gathered into file binder folders, except
for the 1/2"/mile maps which will be reproduced by hand and bound in
separate folders. Copies of all this material will be distributed
to the Northwest Regional office, the Denver office (Game Management
Section) and to the Federal Aid Section.

RESULTS AND DISCUSSION
Current status of the work is stated in step 5 above. For the work done
to date on Wildlife Management Units 31 and 32 see APPENDICES A and B.
These narratives and maps have not yet been corrected by field personnel
and others.
No information was obtained concerning habitat (range) characteristics of
vegetative types (objective 4). The "Land Use Status" tabulation is
essentially a listing by type. It lacks figures on densities, species
composition, condition and trend, etc.
Data gathered under this project should be useful in the pilot computerized
map project now in operation by the Division Planning Section (objective 6),
but this has not been realized as yet.
An extensive bibliography will be included in all final copies of this work,
but in the interest of saving paper and time, is not appended to this job
progress report.

/' r;'?l,'

e77-

,1")/].".,

~. .!./

Prepared by '?t'~.c.a-~-~,-,/I- / //c&amp;~~/
William T. McKean
Wildlife Researcher

.

�-51-

APPENDIX

HILDL IrE

A

~f!-.NAGE·!E~;'Im;rT 11 (ROAN CREEK)

Rio Blanco, Garfield, and Mesa Counties, Colorado

Information on Unit Description,
Population;

Landownership,

Hildlife Species Checklists;

Narrative and Hap Descriptions

Checklist; References.

Compiled by:

To:

HarYests and Seasons;

of Distribution

Census Areas; Habitat Restoration

Land Use, Human

and Abundance;

Projects; Hanagement

March 1974.

l-l.T. l-1cKeanand P. H. Neil

Problems

��-53TABLE OF CONTENTS

Item
UNIT DESCRIPTION
LANDOWNERSHIP

(Boundaries,

size, physical features and climate)----_

59

LAND USE STATUS ------------

_

60

HUMAN POPULATION

_

61

GAME SPECIES

STATUS -------------

55

(Garfield County, 1960-1970 and projections)

(Big game mammals,

small game mammals and game birds)

OTHER MAMMALIAN SPECIES (Furbearers,
statute) ----------------------

nongame mammals defined by

62
_

64

_

65

(Nongame birds, raptores)

_

66

DISTRIBUTION MlD ABUNDANCE OF BIG GAME MAM}fALS (Black bear, elk,
mountain lion, mule deer) ----------.

_

73

NO~GAME ?~ALS

(Not defined by statute) -~---

.OTHER AVIAN SPECIES

DISTRIBUTION

AND ABUNDANCE

OF WILD HORSE o ------

DISTRIBUTION AND ABUNDANCE OF SHALL GAHE MM1MALS
snowshoe hare, red (pine) squirrel) -----

_
(Cottontail rabbit,

_

78

79

DISTRIBUTION MlD ABUNDANCE OF SMALL GAME BIRDS (Grouse, chukar, ringnecked pheasant, wild turkey, band-tailed pigeon, mourning dove,
Gambel's quail, ll1aterfm"l) ---------_

82

DEER HARVEST,

93

SEASONS, AND HUNTING PRESSURE

ARCHERY DEER HARVEST AND SEASONS

(1956-1972)

(1956-1972)

ELK HARVEST,SEASONS,

AND HUNTING PRESSURE

ARCHERY

SEASONS, AND HUNTING PRESSURE

ELK HARVEST,

BLACK BEAR HARVEST ~~

SEASONS

_

(1956-1972)

_

95

_

98

(1956-1972)

100

(1955-1972)

102

MOUNTAIN LION HARVEST AND SEASONS 1965-1972) ----------------------__

104

S}~L GAME HARVEST AND HUNTING PRESSURE
~, 1968-1972) ----------------------

(Small game management

Unit
~____

105

SMALL GAME HARVEST AND HUNTING PRESSURE
17 1968-1972) --------------------

(Small game management

Unit
:::::

-'

DUCK AND GOOSE IllffiVEST
AND HUNTING PRESSURE (Garfield County,
1954-1972) -------------------------------------------

_

106

107

�-54-

TABLE OF CONTENTS (Continued)

Item

Page

INTRODUCTIONS OF GAME SPECIES------------------------------------------

108

CENSUS AREAS AND ROUTES ----~------------------------------------------

110

HABITAT RESTORATION PROJECTS ------------------------------------------

112

MANAGEMENT PROBLEMS CHECKLIST -----------------------------------------

113

�-55-

DESCRIPTION - \.JILDLIFENANI\GEHENT UNIT 31 (ROAN CREEK) 1/

Boundaries l./.--"That portion of Garfield and Rio Blanco counties within the Roan Creek drainage north of the Colorado River, that portion of
Mesa County north and west of the Colorado River which drains into the
Colorado River upstream from the town of Palisade."

Size.--Unit 31 was determined to be 697.2 square miles or 446,240 acres
in size.

A planimeter was used to obtain the acreage from a 1/2" = 1 mile

scale Bureau of Land Management base map, 1965.
Physical Features.--Sedimentary

rocks and alluvium within Unit 31 are

primarily or Tertiary origin.

These occur \"ithin the Green River and
o

Wasatch Formations.

The Green River Formation caps the Roan Plateau

which is bordered by the Roan Cliffs.

That part of the Roan Cliffs

occurring in Unit 31 extends from a point approximately four miles wes t
of the town of Grand Valley on the east to a point approximately one and
one-half miles northeast of Palisade, near Mount Lincoln.
River Formation is best knmm

The Green

for its oil shale deposits which have great

economic potential for oil production.

They are thickest in the Parachute

Creek Member, attaining a maximum thickness of about 3,500 feet.
Creek Member likewise has extensive oil shale deposits.

The Roan

These rocks are

relatively resistant to erosion and generally underlie broad plateaus or
mesas that terminate in sheer cliffs (U. S. Dept. of Agriculture 1965).

1/
- Legally termed "Big Game Management Unit" (see footnote 2/ below) but
tentatively called "Wildlife Management Unit" here to incl~de broader aspects
of anLma I life present.

J:...! Colorado \.Jild1ifeDivision Laws and Regu.La t Lons Hdbk ., 1970 (p, 7, Chap.
'2- Big Game).

�-56-

Beneath the Green River Formation lies the Wasatch, a relatively erodible
rock containing large amounts of soluble salts.

From Grand Valley to Cameo,

and up Roan Creek for many miles, alluvium is from the Wasatch and Green
River Formations.
common also.

Shale fragments, stones, and gravel and cobbles are

Varying concentrations of salt and degrees of alkalinity

exist in the alluvium depending upon the influence of the Wasatch Formation.

Terrain varies (traversing from south to north) from the gently sloping
Grand Valley bottoms to deeply eroded, rounded foothills and alluvial fans.

The foothills abruptly terminate at talus slopes beneath tremendous

cliffs rising 2,000,to 3,000 feet in one half to one mile horizontal
distance.

These cliffs constitute the border of the Roan Plateau, whf.ch

in Wildlife Unit 31, extends 18 to 36 miles northward.

It is a high

rolling tableland, which is steeply dissected by Roan Creek and more
lightly dissected by a multitude of tributaries, including Clear Creek,
Brush Creek, Carr-Creek, Kimball Creek, Dry Fork Creek, and Jerry Creek.
It constitutes the divide between Piceance Creek to the north and Roan
Creek drainages.

\.JildlifeUnit 31 elevations range from 4,}50 feet on

the Colorado River between Cameo and Palisade upward to 9,035 feet at the
head of Carr Creek~

In addition to the above mentioned principal drainages in Wildlife Unit
31, there are Conn Creek and Logan Wash on the east, while in the southwest section are Coon Hollow, Sulphur, Horseshoe, Jackson, Roberts,
Ashbury, and Jerry creeks.

These are all quite short, intermittent and

steep except fo~ J~rry Cr~ek.

�-57-

Soils data for Unit 31 can be obtained by consulting U. S. Soil Conservation Service Work Unit personnel located at Glenwood Springs and at
Grand Junction.

Descriptions and a map of broad soil classification

can be found also in the previously cited report of the U. S. Department
of Agriculture

(1965).

C1imate.--The climate of Unit 31 is typical of Colorado's western slope,
with wide extremes resulting from large variations in topography.

Mean

annual precipitation averages 11. 05 inches (41 years) at Rifle, the most
nearly representative, long-term National Weather Service station.
cipitation has ranged from 7.41-22.2 inches (1956-1972).

Pre-

Precipitation

on Roan Plateau at 8,000 to 9,000 feet is approximately 25 inches annually.
Cloud burst storms of high intensity and short duration occur over
localized areas and result from summer convective conditions.

Seasonal

distribution of precipitation is fairly uniform, higher elevations receiving a greater.proportion

in winter in the form of snowfall.

National Weather Service records at Rifle show a mean annual temperature
of 47.5 degrees, ranging from 45.5 to 47.9 over the past 40 years.
free days averaged 121, ranging from 56 to 166 (1956-1972 incl).

FrostGrowing

season length on Roan Plateau is unrecorded but is believed to range from
30 to 60 days.

Mean monthly evaporation in inches, measured from a sunken "Colorado"
pan at Grand Junction 1948-1960 was: March - 5.41; April - 9.07; }1ay 12.83; June - 17.31; July - 17.43; Aug. - 14.64; Sept. - 11.59; Oct.7.32; Nov. - 3.25.

Preceding climatic information was obtained, and in some cases interpreted

�-58-

from reports of the V. S. Department of Agriculture (1965, 1966) and

V. S. Department of Commerce (continuing) - National Weather Service.

Literature Cited
U. S. Department of Agriculture.

1965.

Water and related land resources,

Colorado River Basin in Colorado.

Coop. Study by Colo. Water Conserv.

Board and V.S.D.A. Econ. Res. Service, Forest Service, and Soil
Conserv. Service.

1966.
Colorado.

~~y.

Denver.

183 pp.

(processed).

Water and related land resources, White River Basin in
Coop. Study by Colo. Water Conserv. Board and U.S.D.A.

Econ. Res. Service, Forest Service, and Soil Conserv. Service.
November.

Denver.

92 pp.

V. S. Department of Commerce.
Colorado.

(processed).

(Continuing).

M~nthly and Annual Summaries.

Fed. Bldg., Asheville, N. C.

Climatological data,
National Climatic Center,

28801.

W. T. McKean
January 1974

�-59-

LANDOWNERSHIP STATUS - HILDLIFE !:-'Ii\NAGEMENT
UNIT 31

Bureau of Land Management

265,093

Acres

Private lands

187,320

Acres

Municipal and County lands

60

Acres

Other Federal lands including National
Park Service, Forest Service, Indian
and Military Reservations, and Bur.
Sport Fisheries and Hi1d1ife

o Acres

Colorado Division of Hi1d1ife lands

o

State Land Board Administration

Acres

----o Acres

lands

452,473

Total

Acres *

(706.9 sq. miles)

* Total land acreage differs from totalo in Unit ~~Description
Section
__ ~----~~
(446,240 acres or 697.2 sq. miles). Bureau of Land Management lands,
private lands, and a total unit acreage were derived from p1animetering
on a 1/2" = 1 mile B.L.M. planning unit map (1965).

P. H. Neil
December 1973

�-60-

LAND USE AND VEGETATIVE COVER STATUS - WILDLIFE HANAGEMENT UNIT 31

Irrigated cropland

8,324 Acres

Grassland with half-shrub mixtures

6,115 Acres

Grassland with woodland mixtures

2,391 Acres

Sagebrush

20,542 Acres

Brush (desert and mountain)

143,721 Acres

Woodland (pinon-juniper and oakbrush)

154,853 Acres

Commercial timber to include aspen

47,299 Acres

Riverbottom vegetation

l/

353 Acres

Urban !:../
Miscellaneous

130 Acres

1/

63,086 Acres

Total

446,814 Acres
(698.1 sq. miles)

Cover type categories were derived from 1/2" = 1 mile S.C.S.Land Use
and Cover Type Maps, 1953, 1954, 1955, for Rio Blanco, Garfield and
Mesa counties. A planimeter and grid were used to derive and compute
acreage. Total land acreage differs from total shown in Unit Description
Section (446,240 Acres or 697.2 sq. miles).

1/
- Includes only woods or brush-woods mix along the Colorado River as planimetered from U.S.G.S. 1:24,000 topographic maps (Palisade, Cameo, DeBeque
and Grand Valley quadrangles).

2/

- Urban - Those areas used for residential and municipal purposes as
illustrated on the 1/2" = 1 mile B.L.M. planning unit base map (1965) for
the t own of DeBeque.

3/
- Miscellaneous - Interpreted to consist mainly of bare ground with sparse,
scattered vegetation of various types.
P. H. Neil
December 1973

�-61-

HUMAN POPULATION - WILDLIFE HANAGEMENT UNIT 31

Specific human population data for Unit 31 are impossible to obtain
because U. S. Census Bureau records are on a county-wide basis only.
The unit is split between Garfield and Mesa counties, with only small
segments of each.

This analyses differs from the Bureau of the Census source in that they
consider cities and towns of less than 2,500 population as being rural
in character.

Here, all incorporated cities and towns are considered

as being urban regardless of size, with remaining populations being
rural.

DeBeque, an incorporated city, had a population of 172 in 1960

which decreased to 155 by 1970.

The remaining rural population is roughly

estimated as ranging between 100 and 200 persons, based upon Soil Conservation Service national estimates of 3.2 persons per farm or ranch.
This does not include new residents associated with oil shale mining,
of whom there are known to be a few at this writing.

William T. McKean
January 1974

�-62GAHE SPECIES - HILDLIFE HANAGEHENT UNIT 31
Big game mammals 1:../
Black bear (Ursus americanus) Uncommon.
Elk (Cervus canadens~~) Common.
Mountain lion (Felis concolor) Uncommon to common.
Mule deer (Odocoileus hcmionus) Common.
Small game mammals !/
Cottontail rabbit (Svlvilagus audubonii; S. nuttallii) Common.
Pine (red) squirrel (Tamiasciurus hudsonicus) Common.
Snowshoe hare (Lepus americanus) Common.
Game birds 'l:./
Migratory waterfowl and shorebirds
Great Basin Canada goose (Branta canadensis moffitti) Common yearlong
resident.
Black brant (Branta nigricans) 1/ Possible rare migrant.
Hhite-fronted goose (Branta albifrans frontalis) 3/ Possible uncommon migrant.
Snow goose (Chen caerulescens caerulescens) 3/, 4/, Possible rare migrant.
Mallard (Anas-piatyrhynchos platyrhynchos) C~mmo~ resident.
Gadwall (Anas strepera) Common spring and fall migrant.

1/
- Nomenclature according to Lechleitner, R. R. 1969.
Colorado. Pruett Publishing Co., Boulder. 254 pp.

Hild mammals of

~/Nomenclature from Bailey, A. M., and R. J. Niedrach. 1967. Pictorial checklist of Colorado birds. Denver Mus. Nat. Rist. 168 pp. Information on occurrence
and status adapted from the above reference and Cringan, A. T., and L. Carlson.
1973. Hildlife in the Piceance Creek Basin, In: An environmental reconnaissance
of the Piceance Basin, Rio Blanco and Garfiel~counties, Colorado. A report on
the completion of Part 1, Phase One of the environmental inventory, analysis and
impact study portion of the Regional Oil Shale Study being done for the State of
Colorado by the Thorne Ecological Institute, Boulder, Colorado, 144 pp. Additional information on occurrence, in employing the term "possible", is adapted
from the foregoing references and Davis, H. A. 1969. Birds in Hestern Colorado.
Colo. Field Ornithologists. 61 pp. Hhere adjective "possible" is absent, actual
sightings have been reported verbally by anyone or more Division personnel
Glenn E. Rogers, Claude E. '~ite, George E. Steele, Kenneth C. Dillinger, or
qualified by additional footnotes that follow.

�-63-

Game birds (continued)
Pintail (Anas acuta) Common spring and fall migrant.
American green-winged teal (Anas crecca carolinensis) Common migrant
and uncommon yearlong resident 4/.
B'l.ue-wLnged
teal (Anas discors di~cors) Common migrant.
Cinnamon teal (Anas cyanoptera septentrionalium) Co~~on migrant.
American wigeon (Anas americana) 4/ Common migrant and rare w Lnter resident.
Northern shoveler (Anas clypeata)-~/ Common migrant and uncommon summer
resident.
Wood duck (Aix sponsa) 1/ Possible rare migrant.
Redhead (Aythya americana) Common migrant.
Ring-necked duck (Aythya collaris) Common migrant.
Canvasback (Aythya valisineria) uncommon to rare migrant.
Greater scaup (Aythya marila nearctica) 3/ Rare migrant.
Lesser scaup (Aythya affinis) Common migrant.
Common goldeneye (Bucephala clangula americana) Common migrant and
winter resident.
Barrow's goldeneye (Bucephala islandica) 3/ Rare winter visitor.
Buffelhead (Bucephala albeola) Uncommon spring and fall migrant and
rare winter resident.
Ruddy duck (Oxvura jamaicensis rubida) Common migrant and occasional
summer resident.
Hooded merganser (Lophodytes cucullatus) Rare winter visitor on river.
Common merganser (Hergua merganser americanus) Common ",inter resident.
Red-breasted merganser (Mergus serrator serrator) Uncommon winter
resident.
American coot (Fulica americana americana) Common migrant and occasional
summer resident.
Common Wilson's snipe (Capella gallinago delicata) Common migrant and
rare winter 'resident.

Upland game birds
Blue grouse (Dandragapus obscurus obscurus) Common.
Sage grouse (Centrocercusurophasianus urophasianus) Uncommon to common.
Ring-necked pheasant (Phasianus colchicus) Uncowmon.
Chukar (Alectoris chukar) 4/ Common.
Band-tailed pigeon (Columba fasciata fasciata) Uncommon summer migrant.
Mourning dove (Zenaida macroura margine11a) 4/ Common summer resident.
Gambel's quail (Lophortyx gambelii sanus) Unknown.
Wild turkey (Heleagris gallopavo merriami) Uncommon.

3/

- Unverified in hunters' bag checks but possible rare migrant and legal
game 1971-72.
~/Changes in nomenclature follow the thirty-second supplement to the
American Ornithologists Union check-list of North American birds published
in Auk 90:411-419, Apr{1, 1973.
W. T. McKean and P. H. Neil
December 1973

�-64OTHER MAMMALIAN SPECIES 1./ - WILDLIFE HANAGEMENT UNIT 31
Furbearers ]j
Beaver (Castor canadensis) Common.
Mink (Mustela vison) Uncertain.
Muskrat (Ondatra zibethicus) Uncommon.
Ringtail (Bassariscus astutus) Uncommon.
Weasels (Hustela erminea; H. frenata) H. erminea Uncertain; M. frenata
Uncommon.

Nongame mammals ~/
Coyote (Canis latrans) Common.
Red fox (Vulpes fulva) Uncommon
Gray fox (Urocyon cinereoargenteus) Common.
Raccoon (Procvon lotor) Uncommon to common.
American badger (Taxidea ta~~s) Common to uncommon.
Spotted skunk (Sp LLoz a.Le putorLus) Uncommon.
Striped skunk (~ephitis mephitis) Common.
Bobcat (wildcat) (Lynx rufus) Common.
White-tailed jack rabbit (Lepus townsendii) COID~on.
Black-tailed jack rabbit (Lepus californicus) Uncommon.
Yellow-bellied marmot (Marmota flaviventris) Common.
White-tailed prairie dog (~~
leucurus) Uncommon to common.
Richardson's ground squirrel (Spermophilus richardsonii) Common.
Thirteen-lined ground squirrel (Suermophilu~ tridecemlineatus) Common.
Rock squirrel (Spermophilus variegatus) Common.
Golden-mantled ground squirrel (Spermophilus lateralis) Common.
White~tailed antelope squirrel (Affimospermophilusleucurus) Uncommon.
Least chipmunk (Eutamias minimus) Common.
Colorado chipmunk (Eutamias quadrivittatus) Common
Uinta chipmunk (Eutamias umbrinus) Uncommon to uncertain.
]j
These species, grouped separately as "furbearers" and "Nongame mammals"
and outside of "game" categories, follow Chapter 62, Colo. Rev. Statutes
1963 As Amended, in Colo. Game, Fish and Parks Div. Laws and Regulations
Hdbk., 1970 (Art. 1, Item 3, Definitions, p. 3).
2/
- Nomenclature according to Lechleitner, R. R. 1969. Wild mammals of
Colorado. Pruett Publishing Co., Boulder, Colorado. 254 pp. and Armstrong,
D. H. 1972. Distribution of mammals in Colorado. Monograph of the
Museum of Natural History, the Univ. of Kansas, Number 3, 1972. 415 pp.

W. T. McKean and P. H. Neil
December 1973

�-65NONGi\:-lE MAHMALS NOT DEFINED
BY STATUTE - It.1ILDLIFE
!l..ANAGE-1ENT
[:-lIT31 1./

Water shrew (Sorex oalustri~) Common.
Vagrant shrew (Sorex vagrans) Common.
Merriams shr-ew (Sorex mer r i amf ) 2/ Rare.
Masked shrew (Sorex cinereus) Common.
Townsend's big-eared bat (Plecotus t owns end i.i.) Common.
Pallid bat (Antrozous Dallidus) 2/ Rare.
Spotted bat (Euderma ~aculatu~) 2/ Uncertain - rare.
Silver-haired bat (Lasionvcteris-noctivagans) Common.
Hoary bat (Lasiurus cinereus) Cncoll'.Inon
- common.
Big brown bat (Eptesicus fuscus) C011'~on.
Western pipistrelle (Pipistrellus hesperus) Common.
Long-legged myotis Wvotis volans) Uncoramon.
California myotis (Hyot Ls californicus) Conmon .
Small-footed myotis (~;yotisleibii) Cornmon,
Fringed myotis (~yotis thysanodes) 2/ Uncertain - rare.
Long-eared myotis C-ivotis evo t i s) r;certain.
Little brown myotis C1~'otis Luc Lf'uzus ) Uncertain.
Brazilian free-tailed bat (Tadarida braziliensis) 1.1 Uncertain - rare.
Northern pocket gopher (Thomomvs talooides) Common.
Ord's kangaroo rat (Dipodomvs ordii) Uncor:.non.
Apache pocket mouse (P~rogn~thus aDache) t'ncommon.
Western harvest mouse (Reithrodontonvs megalotis) Cornmon.
Canyon mouse (Peronvscus crinitus)Col'1mon.
Deer mouse (Peromyscus naniculatus) Common.
Pinon mouse (Peromvscus truei) Co~~on.
Northern grasshopper mouse (Onvchomvs leucogaster) Uncommon - Common.
Desert wood rat (Neot oma Lep Lda ) COTI'JTIon.
Bushy-tailed Hood rat (~eotoma cinerea) Common.
Capper's red-backed vole (Clethriono~vs gapoeri) Common.
Meadmv vole C-Ucrotus penn svl.vanLcus ) Uncommon ,
Montane vole (:!icrotus non t anus ) Unc e.rt aLn ,
Long-tailed vole C'~icrotus longicaudus) Common.
Sagebrush vole (Lagurus curtatus) Uncertain.
House mouse (Nus musculus) Uncoll'.Inon.
Western jumping mouse (Zapus princeps) Cornmon.
Porcupine

(Erethizon dorsatum) Common.

1/
- Nomenclature from Lechleitner, R. R. 1969. Wild mammals of Colorado. Pruett
Publishing Co., Boulder. 254 pp. Infotmution on occurrence and status from
the above reference and: Cringan, A. T., and L. Carlson. 1973. Wildlife in the
Piceance Creek Basin, In: an environmental reconnaissance of the Piceance Basin,
Rio Blanco and Garfield counties, Colorado. A report on the completion of Part
1, Phease One of the environmental inventory, analysis and impact study portion
of the Regional Oil Shale Study beinp, done for the State of Colorado by the
Thorne Ecological Institute, Boulder, Colorado, 144 pp.
2/
- Occurrence listed by the Colo. Div. of \.JildlHe as extremely unusual - very
few documented records within the past decade.
(1972 Status Evaluation for
Selected Colorado Species) appended to: 1973 Wildlife Operations Work Plan,
Field Order No.4 - 1973.

�-66OTHER AVIAN SPECIES 1./ - WILDLIFE HANAGEMENT UNIT 31

Nongame birds ~I
Common loon (Gavia immer) Possible rare migrant.
Horned grebe (Podiceps auritus cornutus) Common migrant.
Eared grebe (Podiceps nigricollis californicus) Possible common migrant 11.
Western grebe (Aechmophorus occidentalis) Common migrant.
Pied-billed grebe (Podilymbus podiceps podiceps) Common migrant and
.summer resident.
Double-crested cormorant (Phalacrocorax auritus auritus) Possible rare
migrant.
Great blue heron (Ardea herodias treganzai) Common summer resident ~/, II.
Snowy egret (Egretta thula breHsteri) Possible uncommon summer resident 1/.
Black-crowned night heron (Xvcticorax nycticorax hoactli) Possible common
summer resident.
Least bittern (Ixobrvchus exilis exilis) Possible rare summer migrant.
American bittern (Botaurus lentiginosus) Possible rare summer migrant.
White-faced ibis (Plegadis chihi) Possible rare migrant.
Whistling swan (Olor columbianus) Possible uncommon migrant.
Sharp-tailed grouse (Pedioecetes phasianellus columbianus) Possible
resident 4/.
Sandhill cr;ne (Grus canadensis canadensis; ~. £. tabida) Possible regular
migrant.
Virginia rail (Rallus limicola limicola) Uncommon summer resident.
Sora (Porzana carolina) Uncommon summer resident.
Semipalmated plover (Charadrius semipalmatus) Possible rare migrant.
Killdeer (Charadius vociferus vociferus) Common summer resident ~/, II
rare winter resident.

11
- These species, grouped separately as "Nongame birds" and "Raptores" and outside of "game" categories, follow Chapter 62, Colo. Rev. Statutes 1962 As Amended,
in Colo. Game, Fish and Parks Div. Laws and Regulations Hdbk., 1970. (Art. 1,
item 3, Definitions, p. 327).
~/Nomenclature from Bailey, A. M., and R. J. Niedrach. 1967. Pictorial checklist of Colorado birds. Denver Mus. Nat. Hist. 168 pp. Information on occurrence
and status adapted from the above reference and Cringan, A. T., and L. Carlson.
1973. Wildlife in the Piceance Creek Basin, In: An environmental reconnaissance
of the Piceance Basin,Rio Blanco and Garfield-counties, Colorado. A report on
the completion of Part 1, Phase One of the environmental inventory, analysis and
impact study portion of the Regional Oil Shale Study being done for the State
of Colorado by the Thorne Ecological Institute, Boulder, Colorado, 144 pp.
Additional information on occurrence, in employing the term "possible", is
adapted from the foregoing references and Davis, W. A. 1969. Birds in western
Colorado. Colo. Field Ornithologists. 61 pp. Where adjective "possible" is
absent, actual sightings have been reported verbally by anyone or more Division
personnel Glenn E. Rogers, Claude E. t~ite, George E. Steele, Kenneth C.
Dillinger, or qualified by additional footnotes that follow.

�-67Nongame

birds

(continued)

Mountain plover (Charadrius montanus) Possible rare migrant 1/.
Black-bellied plover (Pluvialis sauatarola) Possible uncommon migrant 3/.
Long-billed curlew (Numenius americanus 8rnericanus) Possible rare migrant.
Spotted sandpiper (Arctitis macularia) Uncommon summer resident.
Solitary sandpiper (Tringa solitaria cinnamomea) Possible common migrant
and occasional summer resident.
Willet (Catontrophorus seminalmatus inornatus) Unco~~on resident.
Greater yelloulegs (Tringa melanoleuca) Possible cormnon migrant 1./.
Lesser yellowlegs (Tringa flavipes) Possible uncommon migrant 1/.
Red know (Calidris canutus rufa) Possible rare migrant.
Pectoral sandpiper (Calidris melanotos) Possible rare migrant 3/.
Baird's sandpiper (Calidris bairdii) Possible cormnon migrant ]/.
Least sandpiper (Calidris minutilla) Possible cormnon migrant 1/.
Long-billed dowitcher (Limnodromus scolopaceus) Uncommon migrant.
Stil t sandpiper (?1icropalma himantoDus) Possible rare migrant.
Semipalmated sandpiper (Calidrispusilla)
Possible rare migrant 11.
Western sandpiper (Calidris mauri) Possible uncommon migrant 1/.
Marbled godwit (Limosa fedoa) Possible rare spring migrant.
Sanderling (Calidris alba) Possible rare migrant 11.
American avocet (Recurvirostra americana) Rare migrant and resident~
Black-necked stilt (Himantonus mexicanus) Possible rare migrant.
Wilson's phalarope (Steganopus tricolor)- Possible common migrant and uncommon summer resident.
Northern phalarope (LobiDes lobatus) Possible unCO~1Uon migrant.
Pomarine jaeger (Stercorarius pomarinus) Possible rare migrant.
Herring gull (Larus argentatus smithsonianus) Possible uncommon migrant.
California gull (Larus californicus) Possible rare migrant.
Ring-billed gull (Larus delm"arensis) Possible uncommon migrant.
Franklin's gull (Larus pipixcan) Possible uncommon migrant.
Bonaparte's gull (Larus philadelphia) Possible r~re migrant.
Sabine's gull (Xema sabini sabini) Possible rare migrant.
Forster's tern (St;rna forsteri) Possible rare migrant.
Common tern (Sterna hirundo hirundo) Possible rare migrant.
Least tern (Sterna albafrons~~ssos)
Possible rare migrant.
Black tern (Chlidonias niger surinamensis) Possible rare migrant.
Rock dove (Columba livia) Possible cormnon resident.
White~winged dove (Z~da
asiatica mearnsi) Possible rare migrant.
Yellow-billed cuckoo (COCCYZUS americanus americanus) Possible uncommon
summer resident.
Poor-will (Phalaenoptilus nuttallii nuttallii) Common surmner resident ii, 6/.

3/
- Changes in nomenclature follow the Thirty-second Supplement to the American
Ornithologists Union Checklist of North American Birds published in Auk 90(2):
411-419. April, 1973.

4/
- Technically a game bird, but comparative rarity makes occurrence in Unit 31
questionable.
Species is cited here to avoid its complete omission, since
chances appear good enough for it to occur in the unit sometime.

�-68-

Nongame birds (continued)
Cornman nighthawk (Chordeiles minor hesperis; f. ~. howelli) Cornmon
summer resident 6/.
White-throated swift (Aeronautes saxata1is sc1ateri) Common summer
resident E.../,
Black-chinned hummingbird (Archilochus alexandri) Common summer
resident 7/.
Broad-tailed-hummingbird (Se1asphorus p1atycercus p1atycercus) Common
summer resident 5/, 6/, 7/.
Rufous hummingbird (Sel;sph~rus rufus) Common late summer migrant 7/.
Calliope hummingbird (Ste11ula calliope) Possible rare migrant andsummer resident.
Rivo1i's hummingbird (Eugenes fu1gens aureoviridis) Possible rare summer
visitor.
Belted kingfisher (Megacery1e a1cyon a1cyon) Possible common resident.
Common flicker (Co1aotes auratus col1aris) Common resident 3/, 5/, E.../,I/,
(C. a. 1uteus) Possible rare migrant 3/.
Lewis' woodpecker (Asvndesmus Lew i s) Co~on summer resident 7/.
Yellow-bellied sapsucker (Sphyrapicus varius nucha1iae) Comm~n summer 2/,
6/, 7/ and possible occasional winter resident.
Williamson's sapsucker (Sphyraoicus thyroideus nata1iae) Possible common
summer resident.
Hairy woodpecker (Dendrocopos vi110sus monticola) Common resident 7/.
Do~~y woodpecker (DendrocoDos pubescens leucurus) Common resident 5/, 6/, I/.
Northern three-toed woodpecker (Picoides tridacty1us dorsalis) Possible
rare migrant.
Eastern kingbird (Tyrannus tyrannus) Uncommon summer resident II.
Western kingbird (Tyrannus verticalis) Common summer resident 7/.
Cassin's kingbird (Tvrannus vociferans vociferans) Possible uncommon
surnrnerresident.
Ash-throated flycatcher (Myiarchus cinerascens cinerascens) Common summer
resident 7"/.
Say's phoebe-(Savornis saya saya) Common summer II and possible rare
winter resident.
Wil1m" flycatcher (Empidonax traillii) Common summer resident II.
Hammond's flycatcher (Empidonax hanunondii) Possible migrant and uncommon
summer resident.
Dusky flycatcher (Empidonax oberho1seri) Summer resident 7/.
Gray flycatcher (Empidonax vrrLght Ld ) Possible summer resident.
Western flycatcher (Empidonax difficilis hellmayri) Common summer resident I/.

u.

5/
- Sight record given in unpublished checklist of birds of Naval Oil Shale
Reserve, 1969-70, by L. :tof. Stevens, (Specific for Wildlife Management Unit 32).
~/Sight record taken from Cringan, A. T. 1973. Annotated list of birds known to
occur in northwestern Colorado. 17 pp. In: The Colony environmental study,
Parachute Creek, Garfield County, Colorado. Prepared by Thorne Ecological Institute, Boulder, Colorado. August, 1973. Chapt. VII. Part II. Vol. 2, pp. 17-33,
(Specific for Wildlife H.anagement Unit 32).
I/Sight record taken from Smith, A. G. 1973. Avian environmental inventory and
impact study for Colony Development Operation in Garfield County, Colorado. Part
I. Environmental inventory by Thorne Ecological Institute for Colony Development
Operation, Atlantic-Richfield Company, Operator, October, 1973. (processed),
(Specific for Hild1ife Management Unit 32).

�-69Nongame birds (continued)
Western wood peewee (Contopus ~ordidulus veliei) Common summer resident 1..1.
Olive-sided flycatcher Uiutt:111ornis boren Lis) Common summer resident JJ.
Ho~ned lark (Ercmophila alocstris lcucolacma) Common residcnt 1../,2...1.
Violet-green swallow (Trachvcincta thalassina leoida) Common summer
resident 51, 6/, 7/.
Tree swallow-(Irfdoorocne bicolor) Common migrant and summer resident !!.../, 1/.
Bank swallow (Riparia rioaria riparia) Possible uncommon migrant and
summer resident.
Rough-winged swallow (Stel~idootervx ruficollis serripennis) Uncommon
migrant and S~Th~er resident 2/.
Barn swallmv (Hirundo rustica ervthro&lt;:&gt;aster)Cormnon summer resident 2...1,1/.
Cliff swallow (Pctrochelidon pvrrhonota oyrrhonota) Common summer
resident 5/, 6/.
Purple marti; (Pro£ne subis subis) Possible rare summer migrant.
Gray jay (Perisoreus canadensis c2oitalis) Uncommon resident !!.../, 2/.
Steller's jay (Cv ano c i t t a stelleri r-acr'o Looha) Common resident 'i/, 2.../,l/.
Scrub jay (Ache Locoma coeruleseens woodhouse i L) Common resident :i/, 2../, 2/.
Black-billed magpie (Pica pica hudsonia) Common resident 5/,6/, 7/.
Common raven (C;~vus ~x~uatus)
Coenon resident 5/, 6/, 7/. Common crow (Corvus brachvrhvnchos brachvrhvnchos) Possible uncommon resident.
Pinon jay (Gymnorhinus cyanoce~halus) Common SUmr:1er1/,2.../,1/ and winter
resident 7/.
Clark's nutcracker (Nucifra£a columbiana) Common resident 5/,6/, 7/.
Black-capped chickadee (Farus articaDillus garrinus) Co~on reside;t 2/, !!.../, 2/.
Mountain chickadee (Parus gaI!lbeli;:;ar::beli)
COr;'J!.Ol1
resident 5/, 6/, 7/.
Plain titmouse (Parus
inornatus ri0;;:rr Common resident 77. Bushtit (Fsaltrinarus minimus p Lumb eus ) Possible common resident 1/.
White-breasted nuthatch (Sitta carolinensis nelsoni) Uncor;'~onresident 1/.
Red-breasted nuthatch (Sitta canadensis) Rare r2sident ~/.
Pygmy nuthatch (Sitta pv~maea melanotis) Possible uncommon resident.
Brown creeper (Certhia familiaris montana) Uncommon resident and common
migrant 7/.
Dipper (Cin~lus nexicanus unicolor) Common resident 5/,6/, 7/.
House wr en (I'r oz lodv t es aedon pa r kraan i.L) Common summer resident 2/, §.../, 2/.
Bewick's wren (Thrv~ma;es beV7i~kii ercnonhilus) Possible common summer
resident and rare winter resident.
Long-billed marsh wren (Telmatodvtes oalustris plesius) Possible rare
winter resident.
Canyon wren (Saloinctes mexicanus consnersus) Possible uncommon summer resident.
Rock wren (Saloinctes obsoletus obsoletus) Co~~on s~er
!!.../, and possible
rare winter resident.
Mockingbird (~!inusoolvglottos leuconterus) Possible uncommon summer resident.
Gray catbird (Dume t e Ll.a carolinensis) Rare summer resident 1/, ]j.
Sage thrasher (Orcoscootes T!1ontanus)Possible common su".rnerresident.
American robin (Turdus mi£ratorius nropinquus) Common summer and winter
resident 1/, 21, J:...I, 21.
Hermit thrush (Catharus guttatus audubonii) Co:nIDonsummer resident 11,1/,1/.
Swaf.nson s thrush (Ca t ha rus ustulatus a Lmae ) Uncommon migrant 7/.
Veery (Catharus fucescens salicicola) Co~~on migrant and summer
resident 3/, 7/.
Western bluebird-(Sialia mcxicana bairdi) Common migrant and uncommon summer
residen t 7/.
Mountain bluebird (Sf.a Lta cu rrucoides) Common mi gran t and sumner resident
5/, 6/, 7/ and pos s i blc occas i onn l w i n t er resident.
Townsend' s ~oli t aIrc (Hyades tes to\V11s('ndi
t ownscndL) Uncommon rcs Ldcnt;
t

2/, §.../,

u.

�Nongame birds (continued)

-70-

Blue-gray gnatcatcher (Polioptfla caerulea amoenissima) Common summer
resident 7/.
Golden-crown;d kinglet (Regulus satrapa amoenus) Possible uncommon
migrant and rare summer resident.
Ruby-c rowned kinglet (Regulus calendula cineraceus) Common migrant 6/, ]J.
Bohemian waxwing (BoDbycilla garrulus pallidiceps) Possible irregul~r
winter migrant 3/.
Cedar waxwing (Zombvcilla cedrorum) Uncommon·resident 7/.
Northern shrike (Lanius excubitor invictus) Possible common winter resident.
Loggerhead shrike (Lanius ludovicianus excubitorides) Possible uncommon
summer and co~on winter resident.
Starling (Sturnus vulgaris vul£aris) Common resident 6/, 7/.
Gray vireo (Vireo vicinior) Uncommon summer resident 7/. Solitary vireo (Vireo solitarius ulumbeus) Cormnon SU!i1Iller
resident 1./.
~ed-eyed vireo (Vireo olivaceus) Possible rare summ~r resident.
Warbling vireo (Vireo £ilvussHainsonii)
COIT'.r1on
summer resident 7/.
Tennessee warbler (Ver::1ivorauere2rina) Possible rare but regular migrant.
Orange-crowned warbler (Vermivora celata orestera) Uncommon migrant and
summer resident 6/.
Nashville warbler (Verraf.vor
a ruficauilla ric.g~·,ayi)
Possible rare migrant.
Virginia's warbler (Vermivora vir£iniae) COIT~on summer resident 7/.
Yellow warbler (Dendroica uetechia aestiva) Co~mon summer reside;t 5/,6/,2/.
Ye Ll.ow+rumped wa rb Ler (Dendroica coronata rnemorabLl.Ls) Common summer
resident 3/, 6/, 7/; (D..c. coronata) Possible common migrant 3/.
'Black-throat;d gray ;arbler (Dencroica ni.z
r esceris) Commo; s~er
resident 2/.
Townsend's warbler (Dendroica t ownseridi.) Uncommon fall migrant 7/.
~~cGillivray's warbler (Oporornis tolmiei monticola) Common summer resident

5/, 7/.
Common yellowthroat (Geothlvuis trichas occidentalis; G. t. campicola)
Uncommon summer resident 3/, 6/, 7/.
- Yellow-breasted' chat (Icteri~ virens-auricollis) Possible common summer
resident.
Wilson's warbler (Wilsonia pusilla pileolata) COI1L.T'[)on
migrant and SUIl'.mer
resident.
American redstart (Setouhaga ruticilla tricolora) Possible rare migrant.
House sparrow (Passer do~esticus domesticus) Conmon resident 6/, 7/.
Bobolink (Dolichonyx oryzivorus) Possible rare summer migrant~
Western meadowlark (Sturnella ne£lecta neglecta) Common summer ~/, 2/ and
possible uncommon ylinter resident.
Yellow-headed blackbird (Xanthocephalus xanthoceuhalus) Common summer
resident 6/, 7/.
Red-wf.nged blackbird (Agelaius nhoeniceus fortis) Common resident 6/, 7/.
Northern oriole (Icterus galbula bullockii) Co~~on summer resident-3/,-6/, 7/.
Rusty blackbird (~uuha9:us carolinus carolinus) Possible rare winter-migrant~
Brewer's blackbird (EunnaQ:us cvanoceDhalus) Cor.unonresident 6/, 7/.
Brown=he aded cowbird C'~olothrus a t er artemisiae) Common summar r;sident 6/, 7/.
Western tanager (Pfranz a lucovici-;;;) Common migrant and summer resident-7/.Scarlet tanager (Piranga olivacea) Possible rare migrant.
Hepatic tanager (Piranga flava dextra) Extremely rare straggler 1./. (Record
observation).
Black-headed grosbeak (Pheucticus melanocephalus melanocephalus) COll1.T'[)on
summer resident 5/.
Blue grosbe.:lk(Cuir-;c&lt;1c;JC'ruleainte&gt;rfusa) Possible uncommon summer resident.
Lazuli bunting ~~-ina
&lt;1::10(';1;1)
Common summer rcs t derrt Jj.
Evening grosbeak (!lcsperiphona vcspertina brooksi) Common winter
resident ~/. Jj.

�-71-

Nongame birds (continued)
Cassin's finch (Carnodacus cassinii) Possible common resident.
House finch (CarV()ciaCliS
mChicanus frontalis) Common swnmer 2/, 7/ and
possible uncommon winter resident.
Pine grosbeak (Finicola enucleator montana) Possible uncommon resident.
Gray-crowned rosy finch (Leucosticte tephrocotis tenhrocotis; L. t.
littoral is) Possible connon winter resident.
Black rosy finch (Leucosticte atrata) Possible common winter migrant.
Broym-capped rosy finch (Leucosticte australis) Possible common winter
migrant.
Common redpoll (Acanthis fla~ea flammea) Possible rare winter migrant.
Pine siskin (Sui~~s pinus ninus) COffi8onresident 5/.
American goldfinch (S~inus tristus tristus; ~. ~.-pallidus) Possible
common summer and unco~~on winter resident.
Lesser goldfinch (Suinus psaltria psaltria) Possible uncommon summer
and rare winter resident.
Red crossbill (Lox La curvirostra) Possible rare, irregular resident.
White-winged crossbill (Loxia leucoptera leucontera) Possible rare
winter migrant.
Green-tailed towhee (Chlorura chlorura) COillIDon
summer resident 5/, 6/, 7/.
Rufous-sided towhee (PiDilo ervthroDhthalnus montanus) Common s~mer and
rare winter resident 6/, 7/.
Lark bunting (Calamosoiz; melanocorvs) Possible uncommon summer resident.
Savannah spar row (Pas sercuLus s andw i chensLs nevadensis; P. s , anthinus)
Common migrant and sumner resident 6/, 7/.
- Grasshopper sparrow (~odramus
savann;rum-Derpallidus) Uncommon summer
resident 5/, 7/.
Vesper sparrow (Pooecetes gr2~ineus confinis) COIT~onmigrant and summer
resident 5/, il.
Lark sparrow-(Ch~ndestes grammacus stri~atus) Possible common migrant and
summer resident.
Black-throated sparrow (Amphisniza bilineata deserticola) COITmon summer
resident 7/.
Sage sparrow-(A~Dhisniza belli nevadensis) Common summer resident 7/.
Dark-eyed junco (Junco hvemalis a~keni) Possible rare winter migrant 3/;
(.:!. h. hyemali;-; J. h. c i sraon t anus ) Rare w i.nt.e
r resident 1./, 7); (J. h. oreganus) Co~~on winter resident 3/, 7/; (J. h. o. var. mearnsi)
Common winter resident 1/, ]j.
-Gray-headed junco (Junco caniceps caniceDs) Common suw~er ~/ and winter
resident 6/, 7/.
Tree sparrow-(Sryizella arborea ochracea) Possible unCOIT~on winter migrant.
Clay-colored sparro\y (Soizella nallida) Common migrant 7/.
Chipping sparrow (Spizella passerina boreoDhila) COITmon-summer resident

6/, 7/.
Bre;er';- sp arrow (Sp i zeLla bre,.;reri
br ewerL) Common summe r resident 7/.
Harris' spar row (Zonotrichia oue rul.a) Possible uncommon migrant and rare
winter resident.
~fuite-crowned sparrow (Zonotrichia leucronhrvs) Common resident ~/, ~/.
White-throated sparrow (Zonotricbi&lt;1 albicollis) Rare migrant 7/.
Fox sparrow (Passerell;].iliac~ scnist:lcea) Kare summer resident 5/.
Lincoln's s parrow (~Ielospiza lincolnii a l t i.co
La) Common migrant and
summer resident l/.
Song sparrow C·le1 OS!)} za rneLodLa) Corrunonsummer l/ and possible uncommon
winter resident ..
Lapland longspurc (C&lt;llcari~_ lapponictls a lasccnsis) Possible r are winter
migrant.

�-72-

Raptores J:../

Turkey vulture (Cathartes aura meridionalis) Common summer 1/, ~/, 1/
and possible rare winter resident.
Goshawk (Accipiter gentilis atricapillus) Uncommon resident ~/, 2/.
Sharp-shinned hawk (Accipiter striatus velox) Rare summer and common
winter resident 7/.
Cooper's hawk (Accipiter cooperii) Uncommon summer 1/ and common winter
resident 7/.
Red-tailed h;wk (Buteo jamaicensis calurus) Common resident 5/, 6/, 7/.
Swainson's hawk (Buteo swaf.nsom ) Uncommon summer 1/, 1/, and possible
rare winter resident.
Rough-legged hawk (Buteo lagopus ~. johannis) Rare summer and uncommon
winter resident or migrant.
Ferruginous hawk (Buteo regalis) Rare summer and common winter resident 1/.
Golden eagle (AGuila chrvsaetos canadensis) Common resident 6/, 7/, 8/.
Bald eagle (Haliaeetus leucoceohalus alascanus) Common winte-r resident 6/,

ij,Y·

-

Marsh hawk (Circus cyaneus hudsonius) Common summer 1/, ~/, 1/, and uncommon winter resident 7/.
Osprey (Pandion haliaetus carolinensis) P~re migrant.
Prairie falcon (Falco mexicanus) Uncommon resident 1/.
Peregrine falcon (Falco peregrinus anatum) Rare migrant, endangered.
Pigeon hawk (Falco columbarius) Rare winter migrant 7/.
Sparrow hawk (Falco sparverius sparverius) Common su;IDer1/, ~/, 2/ and
possible uncommon winter resident.
Screech ow'l (Ot us asio) Uncommon resident.
Flammulated owl (Otus flammeolus flammeolus) Possible rare summer resident.
Great horned owl (Bubo virginianus) Common resident 5/, 6/, 7/.
Pygmy owl (Glaucid~gnoma
californicum) Possible rare resident.
Burrowing owl (Speotyto cunicularia hypugaea) Common summer 1/ and possible
rare winter resident.
Long-eared owl (Asio otus wilsonianus) Uncommon resident 1/.
Short-eared owl (Asio flammeus flaIT~eus)Possible uncommon winter migrant.
Saw-whet owl (Aeg~s
acadicus acadicus) Uncommon resident 1/.

8/
- Golden and bald eagle specifically excluded from statutes defining "Raptore"
as cited in footnote l/ but herein listed to avoid omission

P. H. Neil
December 1973

�-73-

DISTRIBUTION AND ABUNDANCE OF BIG GAME MAMMALS
WILDLIFE MANAGMENT UNIT 31 (ROAN CREEK)

BLACK BEAR
Black bears occupy approximately the areas between 6,000 and 9,000 feet
elevation in Unit 31, although on rare occasions they may wander over almost
the entire unit.

Their yearlong habitat is largely within the timbered or

brush-covered lands above the Roan Cliffs, where fair combinations of food
are available, such as fruit-bearing shrubs, small mammals, and carrion from
livestock and big game herds.

This is a comparatively small area within Unit

31, but fairly well iso~ated because of the abrupt topography inherent in
the Roan Cliffs (see accompanying distribution maps).

Unit 31 isolation may

not be as great as in Unit 32 because of its many long drainages and intervening narrow ridges.

There are at least jeep type roads, or better, within

or upon almost all drainages and ridges.

Minnich (1973) calculated a bears per square mile figure for a large area
extending from Rifle to the Utah line north of the Colorado River and south
of 'the Roan Plateau summit.

Applying his ratio to Unit 31 produces an

estimated 25 bears at 0.36 bears per square mile.

Estimated annual black bear harvest for the 18 year period 1955-1972,
inclusive, was 5.0 head, based upon hunter report card surveys.

Annual kills

ranged from 0-21; there were only four years out of eighteen when no bears
were reported taken.
complaints.

Bears have occasionally been removed because of damage

In 1972 and 1973 no bears were recorded as being removed (Myers

pers. comm. 1974).

W.C.O. Finch reported in a 1972 "Bear Census", three

bears known, seven estimated, four minimum, and 15 maximum.

�-74-

ELK

Summer range for elk in Unit 31 lies approximately within the range 8,0009,000 feet, generally being the highest, most isolated areas along the north
and northwest portion of the Unit.
"Overall Distribution"

Minnich (1973) delineated separately an

(meaning post-calving and fawning) and a winter distri-

bution; these were from fieldmen personal contacts.
adapted from the above.

Accompanying maps are

Herds which summer on aforementioned ranges are also

believed to interchange freely across high ground areas of Unit 22 on the
north, Unit 21 on the northwest, and Unit 30 on the west.

Winter range for elk in Unit 31 decreases, on the average, to only a fraction
of summer range.

Minnich (1973) maps (distribution maps prepared by Minnich

1973) indicate a narrow band about a mile wide and fourteen miles long across
the northeast portion of the Unit.

Also depicted are three small sites of

about one to five square miles in size, largely on southwest exposures, as
follows (see accompanying maps): junction of Willow and Clear Creeks; Brush
Creek at first Standard Parallel south and below Skinner Ridge; Roan Creek
inT6S,

RlOOW.

No aerial trend counts have been established for elk in Unit 31, however,
occasional observations of elk have been made during spring deer aerial trend
counts.

One such count was eight head in 1973 on Brush Creek.

Unit 31

yielded an estimated annual harvest of seven elk for the period 1959-1972
omitting three years that were closed (1962-1964).
26.

See Harvest Data presented in this report.

Harvest ranged from 2 to

Minnich (1973) com-

puterized maps and data indicate an "Overall" density of 0.5 elk per square
mile with an estimated population of 100 head.

Mean winter population,

derived from the same source was 62 head(l. 0 per mile2).

�-75-

MOUNTAIN LIONS
Numerous records of past and recent occurrences of mountain lions in Unit 31
are available.

From these reports the accompanying maps were drawn.

Virtually the entire unit is considered usable range with the possible exception of culti~ated lands on Roan Creek and its tributaries.

DeBeque Canyon

likewise is probably unsuitable any longer for mountain lions.

The better

yearlong habitat is the broken terrain associated with the Roan Cliffs and
Book Cliffs, but the high mobility and secretiveness of the lion makes its
undetected occurrence in almost any portion of the unit possible (see
accompanying maps).

Dixon (1967), and Armstrong

(1972) make specific citations

concerning lion activity within the Roan Creek drainage.

Minnich (1973) calculated a lions per square mile figure for a.large area
extending from Rifle to the Utah line north of the Colorado River and south
of the sunnnit of the Roan Plateau, but also including all of Douglas Creek
(Unit 21).

Based on his ratios, and assuming that Roan Creek is typical, a

lion population of 27 lions at 0.38 lions per square mile results for Unit 31.

Hunter reports of lions killed during the period 1969 to 1972 by year are as
follows: 1969 - 3; 1970 - 5; 1971 - 13; 1972 - 4.

There could be a few

unreported kills by landowners and others under damage control laws or otherwise.

W.C.O. Finch reports in a 1972 "Mountain Lion Census" four lions

known, 10 minimum, and 20 maximum, with the estimated 1971 population up over
the past five year average.

�-76-

MULE DEER
Mule deer summer range encompasses

the entire unit, except for the Roan

Cliffs proper and low-lying areas of human intrusion along the south border
and in cultivated

lands along Roan

Creek and its tributaries

(see accompany-

ing maps from Minnich 1973).

The upper winter range line in Unit 31 is delineated on the east side by
the Roan Cliffs.

On the west side a line along Cow Ridge north of Dry Fork

and a line along Horse Ridge south of Dry Fork would approximately
upper winter range line (see accompanying

mar~ the

winter range maps).

An area of normal or average winter range appears to occupy approximately
lower one-third of the unit, acc.ording to Minnich 1973 maps.
the main Roan Creek bottoms.
range are identified

the

It also includes

Five small islands within the normal winter

as "concentration"

areas of 2 to 5 square miles in size.

A lower winter range line coincides with U. S. 6 &amp; 24 on the southeast and
on the southwest by the Book Cliffs.

Spring meadow counts by helicopter

from 1965-1973 averaged 2,035 head for

closely similar areas and time spent.
to 2,964 (1973).

Minnich's

winter population

of 2,550 head.

Th~ counts ranged from 1,119 (1965)

(1973) computerized

data for 1972 indicated a

Mean winter density was 24.8 deer per

square mile.

On the basis of total harvest

from 1959-1968, Unit 31 ranked third statewide.

Total kill for the ten year span was 39,061, or'an approximate
harvest of 3,906 animals.

mean annual

As elsewhere in Colorado, deer harvest has declined

in the late 1960's and early 70's.

Annual harvest for recent years was:

1969 - 2,912; 1970 - 1,161; 1971 - 786; 1972 - 1,320.

�-77-

Literature

Armstrong,

D. M.

1972.

Cited

Distribution of mammals

Kansas Printing Service, Lawrence.

Dixon, K. R.

1967.

pp. 141-164.
and Parks.

Minnich, D. W.

of Colorado.

Univ. of

415 pp.

Evaluation of effects of mountain lion predation,
In Game Res. Rep., July, Part II.

Colo. Div. Game, Fish

pp. 73-310.

1973.

Statewide mammal

and bird distribution

maps and

data compiled from W. C.O., Area Supervisor, and Regional Director input.
Located at Denver Office, Colo. Div. of Wildlife.

W. T. McKean
January 1974

(Unpublished).

�-78-

DISTRIBUTION AND ABUNDJU~CE OF WILD HORSE
WILDLIFE HANAGEMENT UNIT 31

Little information is available regarding wild horses in Unit 31, how~
ever, W.C.O. Dillinger

(1973 pers. comm.) reported that approximately

50-60 head are known to occur almost year-round in the southwest quarter
of the unit.

The range that these horses use encompasses approximately

140 square miles and consists mainly of pinon-juniper

type vegetation,

but includes irrigated meadows, riparian areas, and native grass areas.
The northern boundary of their range, according to W.C.O. Dillinger,
appears to be the.South Shale Ridge, while the east and west boundaries
are the main Roan Creek area and the Book Cliffs respectively.
Colorado River comprises the south boundary.

The

It is believed that some

movement in and out of the unit may occasionally

take place.

See

accompanying map.
Feral or wild horses presently do not have legal status under Colorado
laws.

Division Northwest Regional Administrators

have, nevertheless,

expressed concern about possible effects that the increasing numbers of
horses ~ight have on Unit 31 range resources.

In light of controversies

that wild horses have caused e1sewher~ in the West, both the BLM and
Division very probably are in for future difficult decisions in wild
horse management here and at other locations in western Colorado.

P. H. Neil
March 1974

�-79A~~ ABU~~ANCE OF SMALL GAME ~AMMALS

DISTRIBUTION

WILDLIFE HANAGEMENT UNIT 31 (ROAN CREEK)

COTTONTAIL RABBITS
Every square mile of Unit 31 is considered capable of supporting at
least some cottontail rabbits yearlong.
(1969) and Armstrong

According

to Lechleitner

(1972), both the desert cottontail and Nut taLl 's

cottontail should occur, the desert cottontail being common at the
lower elevations

(5,000-7,000) and the Nuttall's cottontail common on

the plateau (7,000-9,000).
No specific studies have been conducted regarding cottontail rabbit
density anywhere in western Colorado, however, Shepherd

(1965a) con-

cluded, after a literature survey, that a possible high-cycle density
of 150-200 animals per square mile and a low-cycle density of 15-20
animals were reasonable
applicable

for much of Colorado.

This estimate should be

to Unit 31.

For abundance, as reflected in harvest data, see tables on Hunter Harvest
'for Small Game Management Unit 8, following.
SNOWSHOE HARES
The snowshoe hare is believed to occur commonly in those areas of the
Roan ,Plateau occupied by Douglas-fir

and Douglas-fir-aspen

mix.

Stevens

(1970) reported this species on the Naval Oil Shale Reserve which includes
a large area of the plateau in the eastern one-half of Unit 32.

It is

reasonable to expect their occurrence in areas of Unit 31 having similar
topography and vegetative

type.

Game distribution maps prepared by

�-80-

Minnich

(1973) and Shepherd

present on the plateau

Minnich

top portions of Unit 31.

(1973) calculates

estimates

(1965b) indicate that snowshoe hares are

a population

of field personnel,

of 1,000 hares, based upon

with an average density of 2.7 hares per

square mile in a large portion of Unit 31.

Harvest data, as an indicator of abundance,
year period 1968-1972,

shows that for the five

in Small Game Management

Unit 8, an. average

annual harvest of 491 snowshoes by 109 hunters occurred.
probably represent

This would

the total kill in Wildlife Units 21, 22, 30, 31, and

32 across the Roan Plateau from the Utah line to Highway 13 north of
Rifle.

RED (PINE) SQUIRREL
Lechleitner
squirrel

(1969), Armstrong

should.be

(1972) and Cary (1911) suggested

fairly common in the coniferous

that this

forests, principally

Douglas-fir,

on the higher mesas and plateaus

in Unit 31.

Neil (pers. comm. 1973) reported several observations

this species in small groves of Douglas-fir
Roan Plateau.

Stevens

(1970) reported

the Naval Oil Shale Reserve
vegetative

of

along the divide road on the

their OCcurrence

(Unit 32) at comparable

on portions of

elevations

and

types.

No data on abundance
reasonably

in Garfield County such as

or hunter harvest data are available.

stable populations

exist in the Douglas-fir

Presumably

types.

�-81-

Literature Cited

Armstrong, D. M.

1972.

Distribution

of mammals in Colorado.

of Kansas Printing Service, Lawrence.

Cary, M.

1911.

Univ.

415 pp.

A biological survey of Colorado.

In North American

Fauna No. 33,U.

S. Dept. Agri., Bur. of Bio1. Sur., Wash., Gov't.

Printing Office.

256 pp.

Lech1eitner,

R. R.

Boulder.

1969.

Wild Mammals of Colorado.

Pruett Pub. Co.,

254 pp.

Minnich, D. w.

1973.

Statewi.de mammal and bird distribution maps and

data compiled from W.C.O., Area Supervisor, and Regional Director
input.

Located at Denver office, Colo. Div. of Wildlife.

Un-

published.

Shepherd, H~ R. .1965a. Colorado long range management
species, 1965-1975, for cottontail rabbit.
Parks Dept. rep. 12 pp.

1965b.
1965-1975,
rep. 11 pp.

plans for game

Colo. Game, Fish and

(Mimeo).

Colorado long range management
for snowshoe hares.

plans for game species,

Colo. Game, Fish and Parks Dept.

(Mimeo).

W. T. McKean
February 1974

�-82DISTRIBUTION AND ABUNDANCE OF S}~L GAME BIRDS
WILDLIFE ~ANAGEMENT UNIT 31 (ROAN CREEK)

BLUE GROUSE
The distribution

maps of Rogers

(1965, 1968) and Minnich

(1973) in-

dicated that the forested and brush covered portions of the Roan
Plateau above 7,000 feet within Unit 31 are available habitat for blue
grouse.

Smith (1973) reported them at comparable

tative types in Unit 32 adjoining
Little information

to the east.

elevations and vege-

See accompanying

maps.

is available regardirtg the true abundance of blue

grouse in Unit 31, however, Minnich

(1973) estimated. a density of 4.9

birds per square mile of blue grouse habitat in this area which results
in an estimated population of 1,513 birds for Unit 31.

Harvest of blue grouse in Unit 31 can be estimated from the Small Game
Harvest and Hunting Pressure tables included in this report.

SAGE GROUSE
Rogers

(1964) indicated several areas within Unit 31 where light popu-

lations of sage grouse are present.

He reported that during the summer

months these birds range from the head of Parachute Creek on the Roan
Plateau west to Douglas Pass near the.Utahline.
vided by Minnich

Distribution

(1973) indicated that sage grouse maybe

out the sagebrush areas on the plateau in most of Unit 31.
accompanying

maps pro-

present throughSee

maps.

Winter range for these birds is not known within the unit, but they are
believed
1964) •

to drop off the plateau onto the lower sagebrush mesas

(Rogers

�-83-

Rogers

(1964) suggested

that fall densities of sage grouse range from

1 to 10 birds per square mile in occupied range in Unit 31.

Minnich

(1973) supports this estimate with an average of 5.25 birds per square
mile, which results in an estimated population of 1,650 sage grouse
for the unit.

Harvest of sage grouse in Unit 31 can be estimated from the Small Game
Harvest and Hunting Pressure tables included in this report.

Between 1952 and 1963 (inclusive), 821 chukars were released within the
boundaries

of unit 31 (see Game Species Introductions)

Canyon, Coon Hollow, Horseshoe

with DeBeque

Gulch, Coal Creek (Cameo area) and Roan

Creek being the main areas of release

(Rogers and Jones 1963; Sandfort

.1956, 1957; Sandfort and Rogers 1960; Woodw'ard and Rogers 1961; Wood,.,ard
et al. 1958).

Minnich

(1973) estimated the present population

of chukar

to be. 982 with an overall density of 7.63 birds per square mile in
habitable range.

Based on .this estimate, the population

increased within the last decade.

has slightly

�-84-

Trend count information
information
lation.

for Unit 31 is lacking and the only comparable

is from Unit 32, which indicates a downward

Unit 32 counts for the period 1964-1973

trend in popu-

ranged from a peak

of 60 in 1964 to 16 in 1972 with no birds being observed
1967, 1968, and 1969.

A summary of Colorado chukar counts from 1958

to 1971 revealed a definite downward
population.

in 1965,

trend in the statewide chukar

Counts ranged from 2,218 in 1963 to 178 in 1971 (Tully

1974 pers. comm.).

Data was not available

for 1972.

Harvest

for chukar is available

in the Small Game Harvest

information

table in this report.
RING-NECKED

PHEASANT

Swope (1965) outlined pheasant range along the Colorado riverbottom
and agricultural

areas extending

from the eastern boundary of Unit 31

to the mouth of Roan Creek and from the mouth of Ashbury Creek to the
western boundary

of the unit.

He also classified

this as poor quality

habitat with a spring breeding population

index of 0-15 based on spring

crow counts and winter sex ratio counts.

Minnich

smaller distribution

pattern located in the riverbottom

areas at the mouth of Roan Creek (see accompanying
density

for this area was estimated

sulting in an estimated

population

Recent records of ring-necked
W.C.O. Dillinger
been observed
population

(1973) showed a much

maps).

Population

at 4.9 birds per square mile reof 50 pheasants

pheasant

(Minnich 1973).

trend counts are lacking, however,

(1973 pers. comm.) reported

that very few pheasants

in the area in recent years and believes

is very. small.

and agricultural

that the local

have

�-85-

WILD TURKEY
Relatively

few wild turkeys are reported

to exist in Unit 31 which

offers little good quality turkey habitat.
occupied

Ho'ffman (1965) showed

turkey range on the plateau near the headwaters

of Roan

Creek and Minnich

(1973) indicated

that there is a small population

of approximately

5 birds in the upper Jerry Creek area with a popu-

lation density of .86 turkeys per square mile and another small
population

of approximately

with a population

density of 2.3 birds per square.

(1973 pers. comm.) reported
while patrolling

20 birds in upper South Dry Fork Creek
W.C.O. Dillinger

that only a few turkeys have been observed

the unit.

Trend count information

for turkeys in Unit 31 is lacking.
Q

to Tully (1974 pers. comm), Unit 31 is generally

According

poor turkey country

and the few turkeys that may be present do not make yearly trend counts
feasible.

A search of turkey harvest data for the years 1959-1969

revealed

no turkeys have been taken from Unit 31 for that period.
information

revealed

turkeys were harvested

that from 1969-1973

an estimated

from Small Game Management

that

Additional

total of three

Units 8 and 17, which

include Unit 31, however, Tully (1974 pers. comm.) stated that these
birds probably did not come from within the boundaries

BAND-TAILED
Braun (1974 pers. comm.) reported
is mainly confined

of Unit 31.

PIGEONS
that band-tailed

pigeon distribution

to the squtheast one-half of Unit 31 and that this

,

area is the northwest

fringe of occupied pigeon range in Colorado.

�-86-

Being migratory by nature, these birds usually arrive in Colorado about
April and depart around October (Braun 1970).

The availability and crop

success of Gambel's oak and various ripening berries within the unit
undoubtedly

influence the distribution of these birds.

Little data are available regarding the abundance of band-tailed pigeons
in Unit 31, however, Braun (1974 pers. corom.) suggested that the population is few and scattered during the breeding season (perhaps less than
one bird per square mile or less than 100 birds on the habitable range)
but may increase during migration periods, particularly in areas of oak.
Unit 31 is considered marginal habitat for band-tailed pigeons largely
because of the lack of ponderosa pine in association with Gambel's oak
(Braun 1974 pers. corom.). Hencefortb, it is believed that band-tailed
pigeons are generally uncommon to this area and that their population
densities fluctuate from year to year in proportion with availability
of food resources

(berries, acorns, new conifer buds, and insect larvae)

and during migration periods.
MOURNING DOVE
Minnich

(1973) indicated that the overall current distribution of

mourning doves included all of Unit 31.

Grieb (1965) indicated that only

the lower elevations of Unit 31 are included in the breeding range of
mourning doves.

Braun (1974 pers. comm.) also suggested that doves are

more common below 8,000 feet along the major drainages and the Colorado
River bottomlands,
accompanying maps.

particularly during migration periods.

See

�-87Mourning doves are common during spring, summer and fall in Colorado
and, according to Bailey and Niedrach (1965), a few may rerr~in over
the winter months in sheltered areas at lower elevations of Unit 31.

Little is known of the actual abundance of mourning doves in Unit 31,
however, Minnich (1973) estimated a density of 5.4 doves per square
mile which results in a population of approximately
doves for the entire unit.

3,765 mourning

It is believed that the local population

density varies considerably from year to year.

Mourning dove harvest for Small Game Management Units 8 and 17, whf.ch
include Unit 31, is illustrated in the Small Game Harvest Table in
another section of this report.

GAMBEL'S QUAIL
Sandfort (1965b) indicated that the distribution of Gambel's quail was
mainly along th~ Colorado River and the lower elevations of Unit 31.
Minnich (1973) indicated that a small population of approximately

15

birds exist in upper Roan Creek near its confluence with Brush Creek
and a larger population of approximately 200 birds exist in the low
lands between DeBeque and the Coon Hollow area.

See accompanying maps.

Trend counts from 1952 and harvest data from 1955 to 1964 revealed that
some birds were observed and harvested in habitable range in Unit 31.
Average ,estimated harvest for Small Game Management Units 8 and 17,
which include Unit 31, was 46 and 1,806 birds respectively for the years
1968-1972 (Funk and Tully 1972, also see Small Game Harvest table).

�-88-

WATERFOWL
Breeding populations:

The mallard and the green-winged

teal are the

most common summer resident ducks in Unit 31, particularly

in those

areas north of the Colorado River (Szymczak pers. comm. 1974).
Mallards have been observed nesting along Parachute Creek and its
tributaries

in Unit 32 to the east (Neil 1972) and since Roan Creek

is a similar but larger drainage offering more waterfowl
habitat,

breeding

it can logically be assumed that mallards and other species

nest here also.
green-winged

Grieb (1965) indicated that the breeding range of the

teal extends over all of the unit and that the mallard

remains the most 'common nesting species in the state.

The Colorado

River and adjacent

canals and ponds,

including

irrigated farmlands with associated

those in the Roan Creek drainage, provide food and cover for

many migratory

species of waterfowl.

According

to Szymczak

comm.) small numbers of Canada geese are beginning

(1974 pers.

to pioneer up the

Colorado River from the Grand Junction area to nest in the spring.
Canada geese were observed along the Colorado River near DeBeque in
May, 1973 by members of the Audubon Society of Western Colorado,
(Dismant 1974).

Other species of waterfowl

Inc.

observed by this club along

the Colorado River and the lower portions of Roan Creek during the
months of March, April and May, 1973, included the following:
eye, common merganser,

mallards,

tail, gadwall, green-winged
western

cinnamon teal, blue-winged

teal, wigeon, ring-necked

grebes, and Canada geese (Dismant 1974).

redhead, and lesser scaup.

teal, pin-

ducks, shovelers

Other species believed

to be present, but not listed by the bird club, include:
common goldeneye,

Golden-

American coot,

�-89-

Little information

is available regarding

spring breeding population

estimates for this area, however, based on somewhat comparable

infor-

mation from the Piceance Creek area (Jobman 1967), an estimated spring
population of waterfowl
Roan Creek-Colorado

Wintering

ranging from 1,000-1,500

may be expected in the

River wetlands.

populations:

Wintering

areas for waterfowl

in Unit 31 con-

sist mainly of available open waters along the Colorado River and Roan
Creek and possibly some of its tributaries.
mergansers,

and mallards were personally

Goldeneyes,

common

observed in December,

1971 on

the Colorado River all along the southern boundary of Unit 31.
mallard is the principal wintering

The

duck species while the majority

of

the geese winter in the Grand Junction area (Szymczak 1974 pers. comm.).
The duck population along the Colorado River between SUt
which includes approximately

35 miles of riverbottom

and Palisade,

in Unit 31, for the

three year period 1971-1973, based upon aerial trend counts in January
by regional personnel,
1,960.

has averaged 1,483 birds ranging from 980 to

Geese averaged 117 birds ranging from 32 to 179 (unpublished

information

- Colo. Div. of Wi1d1.) •

. Literature
Bailey, A. M., and R. J. Niedrach.
of .Nat. Rist.

Braun, C. E.

1970.

Cited

1965.

Birds of Colorado.

Denver Mus.

2 Vol., 895 pp.

Band-tailed

pigeon investigations,

pp . 151-171.

Game Res. Rep., Oct., Colo. Div. Game, Fish and Parks.

171 pp.

In

�-90-

Dismant, S. V.

1974. Palisade, Colorado.

Correspondence

received

containing results of spring bird counts conducted by Audubon
Society of Western Colorado, Inc.

Funk, H. D., and R. J. Tully.
survey - 1971 and 1972.
tables.

2 pp. Typewritten.

Colorado small game hunter harvest
Colo. Div. of Wildlife.

Misc. loose

(Unpublished).

1965. Colorado long range game species management plans,

Grieb, J. R.

1965-1975, for migratory birds.

Colo. Dept. Game, Fish and Parks.

36 pp.
1967. Waterfowl production and management recommendations

Jobman, W. G.

for state-owned lands near Little Hills Experiment Station, Meeker,
Colorado.
38 pp.

CSU term paper for FH 495 bx - Spec. Studies in Wildlife.

(typewritten).

Minnich, D. W.

'1973. Statewide mammal and bird distribution maps and

data compiled from W.C.O., Area Supervisor, and Regional Director
input.

Neil, P •.H.

Located at Denver office, Colo. Div. of vJildlife.

1972.

(unpublished).

Inventory of beaver ponds, East Middle Fork, Parachute

Creek, Aug. 17, 1972.

In Colony environmental study, Parachute

Creek, Garfield County, Colorado.

Prepared for Colony Development

Operation by Thorne Ecological Institute, Boulder, Colorado.
Aug. 1973.

Part III.

Rogers, G. E., and H. Jones.

1963. Chukar planting record.

March.

Original release record located in Sandfort's files at Fort Collins
Research Center.

�-91-

Rogers, G. E.

1964.

Pub. No. 16.

1965.

Sage grouse investigations in Colorado.

Colo. Dept. Game, Fish and Parks.

Tech.

132 pp.

Colorado long range game species management plans,

1965-1975, for blue grouse, sage grouse, sharp-tailed grouse,
ptarmigan.

1968.

Colo. Dept. Game, Fish and Parks.

The blue grouse in Colorado.

Colo. Dept. Game, Fish and Parks.
Sandfort, W. W.

1956.

16 pp.

Tech. Pub1. No. 21.

63 pp.

Chukar partridge release records - Northwest

Region and adjoining areas, Dec. 9, 1951 - Jan. 9, 1956.

Memo

to Commf.ssLon , Fed. Aid Proj. l-l-37-R. 2 pp.
1957.

Chukar partridge release records - Northwest Region,

}1arch 20-26, 1957.

Memo to Commission.

Fed. Aid Proj. W-37-R.

2 pp.
1965a.

Colorado long range game species management plans,

1965-1975, for chukar partridge.

Colo. Dept. Game, Fish and Parks.

18 pp.
1965b.

Colorado long range game species management plans,

1965-1975, for Gambe1's quail.

Colo. Dept. Game, Fish and Parks.

9 pp.

---

, and G. Rogers.

1960.

Chukar planting record.

March.

Original

release record located in Sandfort's files at Fort Collins Research
Center.

�-92-

Smith, A. G.

1973.

Avian Environmental Features at the Colony Plant

Site, Garfield County, Colorado. Part 1. Environmental
by Thorne Ecological
Development

Institute, Boulder, Colorado, for Colony

Operation, Atlantic Richfield Company, Operator,

October, 1973.

Swope, H. M.

1965.

(Processed).

Colorado long range game species management

1965-1975, for ring-necked pheasant.
Parks.

Inventory

plans,

Colo. Dept. Game, Fish and

54 pp.

Woodward, W., M. Lowry, loT.Sandfort, and G. Rogers.
planting record.

April.

1958.

Chukar

Original release record located in

Sandfort's files at Fort Collins Research Center.
______ :,and G. Rogers.

1961.

Chukar planting record.

March.

Original release record located in Sandfort's files at Fort
Collins Re~earch Center.

P. H. Neil
March 1974

�-93DEER HARVESTt SEASONS AND HUNTING PRESSURE
WILDLIFE NANAGD1ENT UNIT 31
(From Colorado Big Game Harvest 1956-1972)

YEAR

HUt-."TING
PRESSURE 1/ BUCKS

1956

2112

1957

HARVEST
HUNT AND SEASON

DOES

FAWNS

TOTAL

1634

913

213

2760

2ES

10/15 - 10/31

2678

1737

1195

294

3226

2DM
2DM
2DM

.
2/
10/1 - 10/14 2/
10/1 - 12/31 10/15 - 11/17

1958

4006

1730,

995

328

3113

1DM
IDM

10/15 - 11/2
11/3 - 11/30

1959

3770

1795

1112

353

3260

1DH
1DM

10/17 - 11/3 2/
11/21 - 12/6 -

1960

1427

1023

·580

115

1718

1DM

10/17 - 11/13

1961

2680

2194

1764

599

4557

3ES
3ES

10/21 - 11/8
11/18 - 12/17 !/

1962

5162

4086

4003

1215

9304

3ES
3ES

10/20 - 11/4
11/5 - 12/31

1963

2929

1422

1374

494

3290

3DA
1DM

8/17 - 9/2
10/19 - 11/7

1964

2587

1471

1020

262

2753

1DM

10/17 - 11/5

1965

2758

1376

1095

365

2836

1m!

10/16 - 11/5

1966

2699

1574

1359

396

3329

2ES

10/15 - 10/24

1967

3053

2256

1710

434

4400

2ES

10/21 - 11/9

1968

3064

1936

1346

332

3614

ES
2ES

10/19 - 10/21
10/22 - 11/7

1969

2427

1501

1130

281

2912

ES
2ES

10/18 - 10/24
10/25 - 11/6

1970

1511

725

348

88

1161

ES
ES
ES

10/17 - 10/21 !~
10/17 - 11/6 !
12/5 - 12/6!/)
12/12 _ 12/13)3?0 per12/19 _ 12/20)m~ts only

1971

1532

786

786

1DB

10/30 - 11/11

1972

1857

942

1320

1DB
ES
ES

10/14 - 10/23
10/14 - 10/23 2/
12/2 - 12/10 -

(Footnotes on following page).

339

39

t~

�-94-

DEER HARVEST, SRASONS AND HUNTING PRESSURE (Continued)
WILDLIFE MANAGEMENT UNIT 31

1/
- Based upon combined total resident and non-resident license sales
and expressed in number of licenses. '

2/
- In part of Unit 31.

Note:

Hunt symbols explained as follows:

ES

=

Either sex, one deer (one license).

2ES

=

Either sex, two deer (one license).

3ES

Either sex, three deer (two licenses plus 3rd deer coupon
on 2nd license).

IDB

=

One deer, buck only (one license).

IDM

=

One deer multiple, either sex (2 licenses and 2 deer per
individual) •

2DM

=

Two deer multiple, either sex (2 licenses and 4 deer per
individual).

3DA

=

Three d~er, one must be antlerless
coupon on 2nd license).

(2 licenses plus 3rd deer

Note: The term IDM evolved into 2D,HC or two deer, hunter's choice
(2:licenses and 2 deer per individual) and is synonymous.

P.H. Neil
December 1973

�-95ARCHERY DEER HARVEST A,.~DSEASONS
WILDLIFE MANAGErtENT UNIT 31
(From Colorado Big Game Harvest 1956-1972 )
YEAR !/

BUCKS

DOES

FAWNS

TOTAL ~/

HUNT AND SEASON

1956 1/

ES
2ES

10/1
10/15

-

10/14
10/31

1957

2DM
2DM
2DM

10/1
10/1
10/15

-

-

10/14, In Part
10/31, In Part
11-17

1958

0

0

0

0

ES
1DM
1DM

9/1
10/15
11/3

-

9/30
11/2
11/30

1959

0

0

0

0

ES
1DM
1DM

9/1
10/17
11/21

-

9/30
11/3
12/6, In Part

1960

0

0

0

0

ES
1DM
1DH

9/10
10/17
12/2

-

9/30
11/13
12/18

1961

2

2

0

4

3ES
3ES
3ES
3ES

8/26
10/21
11/11
11/8

-

9/10
11/8
11/26
12/17, In Part

1962

4

12

5

21

ES
3ES
3ES

8/25
10/20
11/5

-

9/23
11/4
12/31

1963

7

6

0

13

ES
3DA
1DM

8/17
8/17
10/19

-

9/8
9/2
11/7

1964

10

8

0

18

ES
1DM

8/15
10/17

.-

9/13
11/5

1965

8

17

0

25

ES
1DM

8/21
10/16

-

9/12
11/5

1966

16

28

·5

49

ES
2ES

8/20
10/15

-

9/18
10/24

�11 Free permits issued to holders of regular big game hunting licenses by application
;nly. Years 1957-1960, no issue of special permits; archery hunting allowed to
holders of valid deer licenses. Years 1961-1972, separate archery license regulations in effect.
11 Inconclusive kill data are omitted for ye~rs 1956 and 1957.

1/ For years 1956-1967 no hunter pressure data available.
il 203 total resident and non-resident

archery licenses issued for Unit.

51 631 Total hunts for 1969 archery season in Unit 31; total does not reflect
number of hunters.
il 130 resident and non-resident

archery hunters.

71 188 hunters based on total number of hunts; 150 total resident and non-resident
archery licenses issued for Unit.
81 179 total resident and non-resident

archery licenses issued for Unit •.

�-97Ar.chery Deer Harvest and Seasons -Wildlife

Note:
ES
2ES
3ES
lDB
lDM
2DM
3DA

Managment Unit 31 (cont~nued).

Hunt symbols explained as follows:

=

•••

=
=
=
=

=

Either sex, one deer (one license).
Either sex, two deer (one license).
Either sex, three deer (2 licenses plus 3rd deer coupon on 2nd license).
One deer, buck only (one license).
One deer multiple, either sex (2 licenses and 2 deer per individual).
Two deer multiple, either sex (2 licenses and 4 deer per individual).
Three deer, one must be antlerless (2 licenses plus 3rd deer coupon
on 2nd license).

Note: The term lDM evolved into 2D, He or two deer, hunters choice (2 licenses and
2 deer per individual) and is synonymous.

P. H. Neil

December 1973

�-98ELK HARVEST, SEASONS AND HUNTING PRESSURE
WILDLIFE MANAGEMEl'-li UNIT 31
(From Colorado Big Game Harvest 1956-1972)

YEAR

HUNTING
PRESSURE Jj BULLS

COWS

CALVES

TOTAL

HUNT AND SEASON

1956

CLOSED

1957

CLOSED

1958

CLOSED

1959

2

2

0

0

2

AO

10/17 - 11/3

1960

12

3

0

0

3

AO

10/17 - 11/6

1961

14

10

0

0

10

AO

10/21 - 11/8

1962

CLOSED

1963

CLOSED

1964

CLOSED

1965

9

2

0

0

2

25 B + 25 C '!:../

10/16 - 11/5

1966

5

2

2

0

2

25 B+

25 C 1/

10/15 - 11/3

1967

2

2

0

0

2

30 B + 30 C i/

10/21 - 11/9

1968

19

2

2

0

2

30 B+

1969

14

3

0

0

3

30 B + 30 C i/

1970

56

21

5

0

26

AO + 30 C

1971

79

16

2

0

18

AO + 20 C

1972

105

7

1

0

8

AO + 30 C

30 C i/

}../
}../

E./

10/19

11/7

10/18 - 11/6
10/17 - 11/6
10/16 - 10/25
10/28 - 11/12

1/
- Based upon combined resident and non-resident license sales and expressed in
number of licenses. During the period 1956-1970, when elk season coincided
with deer season, the hunting pressure statistics were very probably conservative. Despite difficulties verifying that assumption, unknown substantial
numbers of deer hunters may also have carried elk licenses hoping for a bull
elk under antlered only regulations. This latter happens regardless of low
elk populations. Thus, because of low success and tendency (above non-reporting
statewide figures used in projections) for non-successful elk license holders
not to report by card, elk present in Unit 31 probably received extra pressure
over that indicated by card return projection on years of antlered only seasons.
Discrepancies between hunting pressure and total allowable specified permits
for the years 1965-1969 are probably due to projection error tendencies that
are inherent for all units having small numbers of permits.
(Footnotes continued on following page).

�-99-

ELK HARVEST, SEASONS AND HUNTING PRESSURE (Continued)
WILDLIFE MANAGEMENT UNIT 31

2/
- Units 21, 22, 30, 31, 32 combined.

3/

- Units 31, 32, 33, 42 and part of Unit 41 combined.

!!./
Units 21, 22, 31, and 32 combined.
5/
- Units 22, 31, and 32 combined.

~/

Units 22, 31, 32 and part of Unit 21 combined.

Note:

Hunt symbols explained as follows:

AO

= Antlered only.

B + C

Bull (antlered) and cow (antlerless) permits only (specified).

AO + C = Antlered .only plus specified number of cow permits.

P. H. Neil
December 1973

�-100ARCHERY ELK HARVEST2 SEASONS AND HUNTING PRESSURE
WILDLIFE r-Wl'AGEMENTUNIT 31
(From Colorado
Big Game Harvest
1956-1972)
HUNTING 1/
PRESSURE -

YEAR

BULLS

COWS

CALVES

TOTAL

1956

HUNT AND SEASON
Closed

1957

Closed

1958

Closed

1959

No Data

AO

10/17

- 11/3

1960

No Data

AO

10/17

- 11/6

1961

No Data

AO

10/21

- 11/8

1962

Closed

1963

Closed

1964

Closed

1965

No Data

25B + 25CY

10/16

- 11/5

1966

No Data

25B + 25C1./

10/15

- 11/3

1967

No Data

ES
30B + 30C!!)

9/2
10/21

- 9/17
- 11/9

1968
1969
1970
1971

No Data

o

0

0

0

30B + 30C!!../

10/19

- 11/7

No Data

0

0

0

0

30B + 3004/

10/18

- 11/6

No Data

0

0

0

0

AO + 3005/

10/17

- 11/6

No Data

0

0

0

0

AO + 20C~/

10/16

- 10/25

0

0

0

0

AO
30C§.../
AO +

8/19
10/28

- 8/31
- 11/12

total

resident

and expressed

in

1972

2

!/ Based
number

upon combined
of licenses.

'lj Units

21,

22,

30,

31,

and

1/ Units

31,

32,

33,

42,

and part

~j Units

21,

22,

31,

and

32 comb ined •

~j Units

22,

31,

and

32 comb in ed •

§.../ Units

22,

31,

32,

and part

and non-resident

32 combined.
of 41 combined.

of 21 combined.

license

sales

�-101-

Archery Elk Harvest, Seasons and Hunting Pressure, Wildlife Management Unit 31 (cont., },
Note:
AO
B + C
AO + C

Hunt symbols are explained as follows:

=
=

=

Antlered only.
Bull (antlered) and cow (antlerless) permits only (specified).
Antlered only plus specified number of cow permits.

P. H. Neil
December 1973

�-102BLACK BEAR HARVEST AND SEASONS
WILDLIFE
t-1A.NAGENENT UNIT 31

(From Colorado Big Game Harvest 1955-1972)

!/

Jj, 1.1

YEAR

HARVEST

1955

3R

8/15 - 10/1 and regular deer season.

1956

5R

5/15 - 8/15 and the regular deer season.

1957

6R

4/1 - 11/15 and the pre and regular deer seasons.

1958

3R+4S

4/1 - 11/15 and the regular, and post deer seasons.

+959

o

No data on special season; any regular and post deer
and/or elk season.

1960

4R

4/1 - 9/15 and the regular deer and/or elk seasons
ending 11/6/60.

1961

o

4/1 - 9/15 and the regular and post deer and/or elk
seasons ending 11/8/61.

1962

6R

4/1 - 9/15 and the regular deer season. ending 11/4/62.

1963

6S

4/1 - 9/15 and the pre and regular deer seasons ending
11/7/63.

1964

1R

4/1 - 9/15 and the regular deer season ending 11/5/64.

1965

o

4/1 - 9/30 and the regular deer and/or elk seasons
ending 11/5/65.

1966

21S

4/1 - 9/30 and the regular deer and/or elk seasons
ending 11/3/66.

1967

7

Special season probably 4/1 - 9/30 and regular deer
and/or elk seasons ending 11/9/67.

1968

o

4/1 - 9/30 and the regular deer and/or elk season
ending 11/7/68.

1969

13

4/1 - 6/30 and the regular deer and/or elk season
ending 11/6/69.

1970

8

4/1 - 6/30 and the regular deer and/or elk season
ending 11/6/70.

1971

2

4/1 - 6/30 and the regular deer and/or elk seasons
ending 11/11/71.

1972

1

4/1 - 6/30 and the regular deer and/or elk seasons
ending 11/12/72.

HUNT AND SEASON

(Footnotes on following page).

�-103-

BLACK BEAR HARVEST AlID SEASONS (Continued).
WILDLIFE HANAGEMENT UNIT 31

1/
- R = Regular big game (Deer and/or Elk licenses; S = Bear license for
Special Spring, Summer, or Spring-Summer seasons; Also see annual regulations
for archery hunting.

1:../
From 1955-1959, 1 bear per special bear license and/or 1 bear per bear
coupon on either or both deer and elk license; from 1960-1965, same as
1955-1959, except special bear license variously invalid after September 15
or 30, when bear coupon on either or both deer and elk license covered bag
and possession limits; 1966, same as 1960-1965, except one bear, hunter's
choice, per person per calendar year; from 1967 to present, bear coupon removed from regular deer and elk licenses; thus, one bear, hunter's choice,
per special bear or sportsman's license per person per calendar year only
during special bear and regular deer and/or elk seasons.

3/

- Dogs permitted except when bear season~ concurrent with deer and/or elk
seasons.

P. H. Neil
December 1973

�-104-

MOUNTAIN LION Ilt\RVESTAND SEA.SONS
WILDL IrE m·u'~AGnlENTtJNIT 31

YEAR}j

HARVEST

1965

IF 2:./
(Garfield Co.)

10/16/65 - 3/31/66 West Slope; 10/23/65 - 3/31/66
East Slope, 1 either sex.

1966

5M·, 2F 2/
(Garfield Co ,)
5M 2:./
(Mesa Co.)

1/1/66 - 12/31/66 in all counties ",'estof the
Continental Divide and in Jackson, .Conejos, Alamosa,
Hinera1, Saguache, Rio Grande, Costilla, Archuleta,
Hinsdale, and San Juan counties; 3 either sex.

HUNT AND SE.'\SON

10/22/66 - 2/28/67 in all counties east of the
Continental Divide except those specifically listed
above; 3 either sex.
1967

No Data

9/1/67 - 3/31/68 stat ewfde ; 9/1/67 - 5/31/68 west
of State Hwy. #13 and north of U.S. H"~. #6; 1 either
sex.

1968

1M

9/1/68 - 3/31/69 st.at.ewfde
; 1 either sex.

1969

2M; IF

9/1/69 - 3/31/70 statewide; 1 either sex.

1970

4M; IF

9/1/70 - 12/31/70 designated areas, and regular
deer and elk seasons statewide; 1 either sex.

1971

6M; 7F

3/1/71 - 4/30/71,
1 male only, designated areas;
9/1/71 - 10/12/71 and 11/12/71 - 12/31/71, 1 either
sex, designated areas including Unit 31 •

1972

2M;2F

.2/1/72 - 4/30/72, 1 male only, designated areas;
regular deer and elk seasons, statewide, 1 either
sex, 11/13/72 - 12/31/72, 1 either sex, designated
areas including Unit 31.

o

!/Bounty of $50 per lion paid within period May 7, 1929 - Harch 12, 1965;
protected by statute as of July 1, 1965 when mountain lion license was established.

J:./
Credited to "1965-1966" and "1966-1967" for Garfield and Mesa counties as
reported by Dixon, K. R. 1967. Evaluation of effects of mountain lion predation, p , 147; In Game Res. Rep., July - Part II. Colo. Div. Game, Fish and
Parks. p. 73-310. No data available in reports from section of Game Management for 1965-1967. Dixon's information is probably conservative in that
the damage law allows for lax reporting of kills of marauders by stockmen.
For years 1968-1972 data are from Colorado Big Game Harvest Surveys.
P. H. Neil
December 1973

�SMALLGAMEHARVESTANDHUNTINGPRESSURE
SMM.LGAMENANAGENENT
UNIT 8, 1968-1972

Y

1968

1969

1970

HARVEST
1971

1972

Average

1,509

24,813

11,606

9,276

14,148

13,51l.

14,670

244

109

134

78

of!./ 1,403~/

838

491

124

442

246

1,172

73

368

78

815

501

258

267

537

311

603

767

724

597

1,134

765

-

-

-

106

-

-

-

-

-

169

122

453

151

223

-

237

76

606

247

358

-

322

27

-

61

-

-

44

31

-

153

-

-

92 ~

586

1,040

127

246

446

489

-

5,828

2,412

2,091

4,373

3,676

1968

1969

NUHBEROF HUNTERS
1972
1971
1970

Average

1,984

1,328

1,319

1,411

1,502

Snowshoe hare

101

30

73

95

E1ue grouse

298

154

213

Sage grouse

190

301

- ]j

Chukar
Gambe1's quail

SPECIES
Cottontail

rabbit

Sharp-tailed

grouse

Pheasant
Band~tailed
}louming

Y

0
VI

313

621

129

250

256

314

-

276

290

242

447

314

pigeon

dove

11 The area of that portion of ~.Ji1dlife Unit 31 which falls within Small Game rIanagement Unit 8 is approximately
362
;quare miles.
This is approximately
7.2 percent
(7.2%) of Unit 8's total area (362 -; 4,970 sq mi) as p1animetered from
Colorado topographic
map 1: 500,000.
The other portion of Wildlife
Unit 31 falls within Small Game Unit 17. The area
of this portion
is approximately
335 square miles.
This is approximately
15.5 percent
(15.5%) of Unit 17's total area
(335 f 2,286 sq mi).

2:..1 Hean for years

11 ~o season,
il Possible

shown.

or no data.
sample size

error.
W. T. McKean
January 1974

I

�SMALL CAME HARVEST AND HUNTING PRESSURE
SHALL GANE MANAGEMENT UNIT 17,__1968-1972

1968

1969

NUMBER OF HUNTERS
1970
1971

1972

Average'l:.l 1968

1,763

1,549

2,340

2,037

2,967

2,131

Snowshoe hare

156

158

106

55

81

Blue grouse

83

106

61

183

Sage grouse

69

55

36

79

98

-Chukar

496

Gambe1 's quail

365

SPECIES
Cottontail

rabbit

Sharp-tailed

grouse

JJ
1969

1970

HARVEST
1971

1972

AveraBe

15,998

11,329

19,458

22,470

27,056

19,262

111

902

350

334

526

880

598

475

182

872

218

88

404

1,629

644

100

.286

109

144

87

58

172

577

233

-

58

211

112

178

38

-

77

253

137

A17

342

239

-

374

1,913

806

569

418

-

905

430

420

359

509

419

-1,161

2,222

2,014

1,576

2,573

1,900
I

Pheasant
Band-tailed
Mourning

1,19l

1,242

1,163

-

-

-

402

pigeon

dove

1,132

1,845

-

12

10

531

573

932

1,315

3,224

1,932

2,787

2,713

4,128

2,960

1J.

-

-

-

117

12

64

606

-

3,561

7,539

5,378

9,994

6,640

for years shown.

1/ No season,

0
0\
I

1/ The area of that port ion of Wildlife Unit 31 which falls within Small Game Management Unit 17 is approximately 335
;quare miles.
This is approximately fifteen percent
(15.5%) of Unit 17's total area (335 ~ 2,286 sq mi) as p1animetered
from Colorado topographic map 1:500,000.
The other portion of Wildlife Unit 31 falls within Small Game Management Unit
8. The area of this portion is approximately 362 square miles.
This is approximately seven percent (7.2%) of Unit 8's
total area (362 ~ 4,970 sq mi). Data from both Small Game Units 8 and 17 are included herein.

1/ Mean

~

or no data.
W. T. McKean
January 1974

�-107DUCK AND GOOSE HARVEST AND HUNTING PRESSURE GARFIELD COUNTYz 1954 - 1972 1/
(From Colorado Waterfowl Kill Surveys)

YEAR

DUCKS
EST. NO. HUNTERS
EST. HARVEST

1954

No Data

2,284

1955

No Data

4,231

1956

No Data

1,764

1957

No Data

1,909

1958

372

1,442

1959

541

1,721

1960

112

1,194

1961

291

674

1962

260

414

1963

277

1,504

1964

291

1,404

1965

362

1966

50 11

1967

184

406

1968

265

1,348

1969

312

2,901

1970

403

1971
1972

GEESE '.!:./
EST. NO. HUNTERS EST. HARVEST

No Data

19

35

34

60

57

1,076

32

No Data

1,945

16

No Data

2,795

25

No Data

538

4,923

4
75 Permits

1
(2)

302

1,219

No Data
250 Permits

1
(l)

!/Waterfowl kill data are available only by county. Riverbottom lands in
Wildlife Unit 31 resemble those in Unit 32 (Garfield County) more closely
than they do Unit 30 (~!esaCounty). Accordingly, for harvest approximation
purposes Garfield County dat a are show-n. For purposes of making comparisons
these figures could readily be converted to birds harvested per mile of
riverbottom.
~/Accuracy of data questionable due to sampling difficulties
with very small populations. geese and goose hunters alike.
l'obvious

associated

inaccuracy.

W. T. HcKean
January 197/•

�GAME SPECIES INTRODUCTIONS - WILDLIFE MANAGEMENT UNIT 31
Species

Date

Hales

Females

Total

Release Area

Chukar

12/22/52

3

3

6

Coon Hollow-l mi. so. and 1-1/2
mi. west; of DeBeque on Ed Finley
ranch.

Escalante - '''ild- Sandfort 's Files
trapped
(Ft. Collins Res.Cen.)

24

20

46

DeBeque Canyon-directly across
river from cable car, 3.3 miles
above jct. of Colo.65 and U.S. 6
&amp; 24 on first bench above river.

Escalante - wildtrapped

Sandfort 's Files
(Ft. Collins Res.Cen.)

12/8/55

(2) Un-

classified

Source of Stock

References

1/9/56

11

19

30

DeBeque Canyon-in small draw on
Escalante - wildwest side of Colo. River about ~
trapped.
mi. above cable crossing river; 4.2
mi. above jct. of Colo.65 and U.S.
6&amp;24.

Sandfort's Files
(Ft. Collins Res.Cen.)

3/22/56

25

25

50

Same as 1/9/56.

Little Hills

Sandfort 's Files
(Ft. Collins Res.Cen.)

3/23/57

50

50

100

DeBeque Canyon-at mouth of Jackson Gulch,approximately 5.0 mi.
above DePlato on w , side of Colo.
River.

Little Hills

Sandfort's Files
,
(Ft. Collins Res.Cen.)t;
0)
I

4/3/58

22

50

72

Near mouth of first canyon (probably Horseshoe Gulch) upstream
from Akin.

Little Hills

Sandfort's Files
(Ft. Collins Res.Cen.)

3/23/60

75

75

150

Coal Creek (Cameo area) about 2.8
mi. up Coak Cr. from canal; just
west of Colorado River.

Little Hills

Sandfort's Files
(Ft. Collins Res.Cen.)

3/17/61

75

75

150

Coal Creek (Cameo area) about 3.0
mi. west of canal road.

Little Hills

Sandfort's Files
(Ft. Collins Res.Cen.)

3/30/63

65

52

117

Roan Creek-14 mi. w. of DeBeque on Little Hills
Deer Pk. Rd. near s. side of large
reservoir.

Sandfort's Files
(Ft. Collins Res.Cen.)

3/30/63

50

50

100

Roan Creek - 1.2 mi. no. of DeBeque on Roan Creek Road.

Sandfort's Files
(Ft. Collins Res.Cen.)

(Literature Cited on folloving page).

Little Hills

�-109-

GAME SPECIES INTRODUCTIONS (Continued)
WILDLIFE MANAGEMENT UNIT 31

Literature Cited

Rogers, G., and H. Jones. 1963. Chukar planting records. March.
Original release records located in Sandfort's files at Fort
Collins Research Center.
Sandfort, W. W. 1956. Chukar partridge release records - Northwest
Region and adjoining areas, December 9, 1951- January 9, 1956.
Hemo to Commission.
Fed. Aid Proj. 'ol-37-R. 2 pp ,
1957. Chukar partridge release records - Northwest Region,
March 20-26, 1957. Memo to Commission. Fed. Aid Proj. W-37-R.
2 pp.
, and G. Rogers. 1960. Chukar planting record. March. Original
---release
record located in Sandfort's files at Fort Collins Research
Center.
Woodward, W., and G. Rogers. 1961. Chukar planting record. March.
Original release record located in Sandfort's files at Fort Collins
Research Center •
• M. Lowry., W. Sandfort, and G. Rogers. 1958. Chukar planting
---record.
April. Original release record located in Sandfort' s files
at Fort Collins Research Center.

P. H. Neil
January 1974

�-110-

CENSUS AREAS A~~ ROUTES - WILDLIFE MANAGB1ENT UNIT 31

MULE DEER
An annual spring trend count by helicopter has been conducted in Roan
Creek and the majority of its tributaries from 1965-1973.
aircraft were used previous to 1965.

Fixed-wing

Procedural details are available

in the Northwest Regional Office; summaries of results may be found
there and in the Denver office of the Game .Manager. These summaries
do not necessarily

show animal numbers by Wildlife Management Unit, but

more commonly by drainage.
ELK
Because of the relative low density of elk in Unit 31, specific winter
aerial trend counts have not been established for the unit.

Occasional

observations of elk and their locations have been recorded during deer
aerial trend counts.

Any available information regarding the results

of these observations

are on file in the Northwest Regional Office.

BLACK BEAR
Incidental to deer aerial trend counts, black bear are tallied in Units
22, 31, 32 and others.

In addition, an annual estimate, by WCO district,

of bear populations was begun in 1972.

This estimate also included har-

vest by trappers and others, and management objectives for the species.

CHUKAR
We have no records of established chukar production counts for Unit 31,
however, several summer counts and population estimates were made in
De Beque Canyon and Coal Creek (Cameo area) which were both release areas

�-111-

for introduced chukar (see Game Species Introductions).

Results of

these counts may be found in Sandfort's files on chukars at the Fort
Collins Research Center.
WATERFOWL
Wintering populations of ducks and geese in Unit 31 have been
censused annually from fixed-wing aircraft at least in 1971,
1972 and 1973, possibly earlier, by Regional Management personnel.
Areas covered were on the Colorado River from Silt to Palisade.
Summarizing tables do not break down specific river section populations.
Field notes of observers may do so -- see Northwest Regional personnel.

P. H. Neil
March 1974

�HABITAT RESTORATION PROJECTS - WILDLIFE MANAGEMENT UNI~ 31 1/
Date
9/11-12/59

Location

Vegetative Type

Treatment

No. Acres
Treated

Purpose of
Project

Agency

T9S R99W Sec. 10,11,
15

Sagebrush

Plow and seed with
Agropyron cristatum,
Me1ilotus officina1is

510

Brush removal
and revegetation

B.L.M.

10/11/59

Ashbury Creek
T9S R99W Sec. 8,9,
18

Sagebrush

Plow and seed with
Agropyron cristatum,
Melilotus officinalis

156

Brush removal
and revegetation

B.L.M.

12/3-15/59

T9S R98W Se~. 11-13

Sagebrush

Plow and seed with
Agropyron cristatum,
Melilotus officinalis

211

Brush removal
and revegetation

B.L.M.

10/8-11/6/61

Kimball Creek
T7S R99W Sec. 17

Gambel's oak
&amp; Serviceberry

Burn and seed with
Agropyron cristatum,
Melilotus officinalis

358

Range rehabilitation

B.L.M.

1967

Pine Gulch
T8S R99 W Sec.19,20,30
T8S RlOOW Sec. 25, 26,
34, 35, 36

Sagebrush

Chain (double)

Modify range
from sage
class

B.L.M.

1968

Brush Mountain
T5S moov Sec. 1, 2,.
3, 4, 10, II, 12

Sagebrush

Brush cutting with rotary
cutter

Soil and watershed modification

B.L.M.

1,254

531

1/
If details are desired on any of these projects, make requests directly to Mr. Ron Kufe1d, Colorado Division of
Wildlife Research Center, Fort Collins, Colorado 80521.
P.H. Neil and W. T. McKean
January 1974

,
•...
•...
,

N

�-113-

MANAGEMENT PROBLEMS CHECKLIST - WILDLIFE MANAGEMENT UNIT 31

1.

Deer-auto collisions on U. S. Highway 1-70.
Though not as serious as in Unit 32, this problem exists and will
become more acute when Interstate 1-70 is completed to Plateau
Creek as planned.

For a detailed discussion of this problem, in-

cluding projected numbers of collisions, see: Colo. Div. of Wildlife. 1973.

Fish and wildlife analysis of the Interstate 70

highway corridor Rifle to Plateau Creek. Draft report. July. 126 pp.
Persons assisting in this report were: John Torres, Jerry Craig,
George Kidd, Dale Reed, Eddie Kochman, Maurice Potter, and Northwest
Regional personnel.

Copies of this report may be obtained in Denver

and Northwest Regional offices.

With growth of the oil shale in-

dustry, particularly at the Logan Wash in-situ plant, deer-auto
collisions will increase along lower Roan Creek from November through
April each year.

2.

Winter range losses.
A proposed alternate route for U. S. Highway 1-70 (alternates 2-C)
woul~ be constructed north of the Denver and Rio Grande Western Railroad from a point near the mouth of Horseshoe Canyon north eastward
to a point about two miles north of DeBeque,thence

south eas tward to

a point of juncture with old U. S. 6-24 approximately five miles east
of DeBeque.

This loop would be very destructive since it would bisect

a major deer winter range, cutting off access to Colorado River water,
and intercept migration south to Unit 42 even more than does existing
U.S. 6-24.

For a more detailed account of the impact of 1-70 see the

�-114-

report cited above.

Stress to wildli.fe, especially big game, from the oil shale industry
with its many ancillary developments is deemed to be a serious future
management problem in Unit 31.

A more detailed analysis of proposed

oil shale development and possible effects on the environment can
be found in: Thorne Ecological Institute (1973), Tweedy (1971) and
U. S. Dept. of Interior (1973).

3.

Deer damage to farm crops.
In general, the problem of deer damage to farm crops in Unit 31 is
relatively small in comparison to some other units.

According to

Myers (1973 pers. comm.), claims for 1971-72 totaled $1,770.70 for
damage to alfalfa and grasses in Units 31 and 32 combined.
for 1972-73 totaled only $130.00.

Claims

Decreased production of alfalfa

and spring grasses is alleged because of early grazing by deer.
Some damage to stacked alfalfa has been fairly common in winter, as
well as to the few scattered orchards in this area.

Literature Cited

Thorne Ecological Institute.

1973.

The Colony environmental study,

Parachute Creek, Garfield County, Colorado.

Tweedy, J. B., et ale

1971.

August.

Boulder.

3 vol.

Report on economics of environmental pro-

tection for a Federal oil shale leasing program.

A Special Committee

of the Governor's Oil Shale Advisory Committee (for Colorado).
January.

204 pp.

U. S. Department of Interior.

1973.

Final environmental statement for the

prototype oil shale leaSing program.

Washington, D. C.

6 vol.

�-115-

APPENDIX

H

\\TILDLIFEXANAGMENT UNIT 32 (PARACHUTE CREEK)
Garfield County, Colorado

Information on Unit Description, Landownership, Land Use, Human
Population; Wildlife Species Checklists; Harvests and Seasons;
Narrative and Hap Descriptions of Distribution and Abundarice ;
Census Areas; Habitat Restoration Projects; Management Problems
Checklist; References.

Compiled by:

To:

October 1973.

H. T. HcKean and P. H. Neil

�-116-

TABLE OF CONTENTS
Item
UNIT DESCRIPTION
LANDOWNERSHIP

(Boundaries, size,physica1

features and climate)----- ...•
118,

STATUS-----------------

---122

LAND USESTATUS--------------.HUMAN POPULATION

---------123

(Garfield County, 1960-1970 and projections)-----------124

?AME SPECIES (Big game mammals, small game mammals and game birds)------127
OTHER ~~Lhl~
SPECIES (Furbearers, nongame mammals defined
by statute)-- ....
-----------------------129
NONGAME MAMMALS

(Not defined by statute)-----------------

130

OTHER AVIAN SPECIES (Nongame birds, raptores)--------------

131

DISTRIBUTION AND ABUNDANCE OF BIG GAME MAMMALS (Black bear, elk,
mountain lion, mule deer)----------------------

138

DISTRIBUTION AND ABUNDANCE OF SMALL GAME MAMMALS (Cottontail rabbit,
snowshoe hare, red (pine) squirrel)---------------

143

DISTRIBUTION AND ABUNDANCE OF SMALL GAME BIRDS (Grouse, chukar,
ring-necked pheasant, wild turkey, band-tailed pigeon,
.
mourning dove, Gambe1's quail, waterfow1)--------~-------------146
DEER HARVEST, SEASONS, AND HUNTING PRESSURE

(1956-1972)-----------

_ 155

ARCHERY DEER HARVEST AND SEASONS (1956-1972)----------

_ 157

ELK HARVEST, SEASONS, AND HUNTING PRESSURE

_ 159

(1956-1972)----------

ARCHERY ELK HARVEST, SEASONS, AND HUNTING PRESSURE (1956-1972)---

161

BLACK BEAR HARVEST AND SEASONS (1955-1972)---------

163

MOUNTAIN LION llARVEST
AND SEASONS (6)
19 5-1972
.

_ 165

SMALL GAME HARVEST AND HUNTING PRESSURE (Small game management
Unit 8, 1968-1972)------------

_

DUCK AND GOOSE HARVEST AND HUNTING PRESSURE
1954-1972)---------------INTRODUCTIONS

(Garfield County,

OF GAME SPECIES----------------------

~---------

166

167
168

•

�-117-

~

OF CONTENTS (Continued)

Item

Page

CENSUS AREAS AND ROUTES------------------------------------------------

169

HABI~T

RESTORATION PROJECTS -------~----------------------------------

171

MANAGEMENT PROBLEMS CHECKLIST -----------------------------------------

172

�-118-

DESCRIPTION - WILDLIFE MANAGEHENT UNIT 32 (PARACHUTE CREEK) 1/

Boundaries l:..l.--"That
portion of Garfield County uithin the Parachute
Creek drainage and in the Government Creek drainage west of State
Highway 13, and north and west of the Colorado River."

Size.--300.7 square miles or 192)416 acres.

Planimetered from a Bureau

of Land Management base map, 1969, 1/2" = 1 mile.

Physical Features.--Sedimentary

rocks and alluvium within Unit 32 are

primarily of Tertiary origin.

These occur within the Green River and

Wasatch Formations.

The Green River Formation caps the Roan Plateau

which is bordered by the Roan Cliffs.

That part of the Roan Cliffs

occurring in Unit 32 extends from Rifle on the east to a point approximately four miles west of the town of Grand Valley.

The Green River

Formation is best knoml for its oil shale deposits which have great
economic potential for oil production.

They are thickest in the

Parachute Creek Member, attaining a maximum thickness of about 3,500
feet.

These rocks are relatively resistant to erosion and generally

underlie broad plateaus or mesas that terminate in sheer cliffs (U. S.
Dept. of Agriculture

1965).

Beneath the Green River Formation lies the

Wasatch, a relatively erodible rock containing large amounts of soluble
salts.

From Rifle to Grand Valley alluvium is from the l-lasatchand

Green River Formations.
are common also.

Shale fragments, stones, and gravel and cobbles

Varying concentrations of salt and degrees of alkalinity

..
!.lLegally termed "Big Game Management Unit" (see footnote Jj below) but
tentatively called "~nldlife Management Unit" here to include broader
aspects of animal life present.
2/
- Colo. Div. Wildlife Laws and Regulations Hdbk., 1972 (p , 7, Chap. 2 _
Bi P, Game.

�-119-

exist in the alluvium depending upon the influence of the \.JasatchFormation.

Patches of white salt are common in the exposed Wasatch between

Rifle and De Beque .

.Terrain varies (traversing from south to north) from the gently sloping
Grand Valley bottoms to deeply eroded, rounded foothills and alluvial
fans.

The foothills abruptly terminate at talus slopes beneath tremendous

cliffs rising 2,000 to 3,000 feet in one half to one Nile horizontal
distance.

These cliffs constitute the border of the Roan Plateau, which

extends six to ten miles northward.

It is a high rolling tableland, which

is steeply dissected by Parachute Creek and more lightly dissected by a
multitude of tributaries.

It constitutes the divide between Piceance Creek

and Parachute Creek drainages.

Unit elevations range from 4,996 feet on the Colorado River near Una upward to 9,286 feet at the head of East Parachute Creek above Anvil Points.

Other than Parachute Creek, the only drainages of consequence are:
Government Creek and Hubbard Creek on the east.

Yellow Slide Creek,

Balzac Creek, Cottonwood Creek, Hayes Creek, and Kelley Creek are all intermittent, ·steep, southwar d flowing draLnageways of short mileage.

Soils data for Unit 32 can be obtained by consulting U. S. Soil Conservation Service Work Unit personnel located at Glenwood Springs.

Descriptions

and a map of broad soil classifications can be found in a previously cited
report of the U. S. Department of Agriculture (1965).

Climate.--The climate of Unit 32 is typical of Colorado's western slope,
with wide extremes resulting from large variations in topography.

Mean

�-120-

annual precipitation

averages 11. 05 inches (41 years) at Rifle, the only

long-term National Weather Service station.
from 7.41-22.2 inches (1956-1972).

Precipitation has ranged

Precipitation on Roan Plateau at

8,000 to 9,000 feet is approximately 25 inches annually.

Cloud burst

storms of high intensity and short duration occur over localized areas
and result from summer convective conditions.

Seasonal distribution of

precipitation is fairly uniform, higher elevations receiving a greater
proportion in winter in the form of snowfall.

National Weather Service records at Rifle show a mean annual temperature
of 47.5 degrees, ranging from 45.5 to 47.9 over the past 40 years.
Frost-free days averaged 121, ranging from 56 to 166 (1956-1972 incl.).
Growing season length on Roan Plateau.is unrecorded but is believed to
range from 30 to 60 days.

Mean monthly evaporation in inches, measured from a sunken "Colorado"
pan at Grand Junction 1948-1960 was: March - 5.41; April - 9.07; May 12.83; June - 17.31; July - 17.43; Aug. - 14.64; Sept. - 11.59; Oct.7.32; Nov. - 3.25.

Preceding climatic information was obtained, and in some cases interpreted,
from reports of the U. S. Department of Agriculture

(1965, 1966) and U. S.

Department of Commerce (continuing) - National Weather Service.

Literature Cited
U. S. Department of Agriculture.

1965.

Water and related land resources,

Colorado River Basin in Colorado.

Coop. Study by Colo. Water Conserve

Board and U.S.D.A. Econ. Res. Bervf'cejFo res t Service, and Soil Conserve
Service.

May.

Denver.

183 pp. (processed).

�-121-

U. S. Department of Agriculture.

1966.

White River Basin in Colorado.

Water and related land resources,

Coop. Study by Colo. Water Conserv.

Board and U.S.D.A. Econ. Res. Service, Forest Service, and Soil
Conserv. Service.

November.

U. S. Department of Commerce.
Colorado.

Denver.

92 pp. (processed).

(Continuing).

Monthly and Annual Summaries.

Fed. Bldg., Asheville, N.C.

Climatological data,
National Climatic Center,

28801.

W. T. McKean
July 1973

�-122-

LANDOWNERSHIP

STATUS - lHLDLIFE MANAGE11ENT UNIT· 32
Acres

Bureau of Land Management
Outside Naval Oil Shale Reserve boundary
Within Naval Oil Shale Reserve boundary

25,989
53,814
Total

Private lands
Outside Naval Oil Shale Reserve boundary
Within Naval Oil Shale Reserve boundary

79,803

106,949
4,255
Total

Municipal and County lands

111,204
36

Other Federal lands including National Park
Service, Forest Service, Indian and
Military Reservations and Bur. Sport
Fisheries and Wildlife

o

Colorado Division of Wildlife lands

1

State Land Board·Administration

o

lands

Total

191,044 *

*Total land acreage differs from total in Unit Descriptio~ Section (192,416
acres or 300.7 square miles). Bureau of Land Management lands and Naval
Oil Shale lands and 8. total unit acreage wer e derived from p1animetering
on 1" = 1 mile or 1/2" = 1 mile B.L.M. planning unit base maps (1965 &amp; 1969).

W. T. McKean and Paul H. Neil
July 1973

�-123LAND USE AND VEGETATIVE COVER
STATUS - WILDLIFE HANAGEMENT UNIT 32

Irrigated cropland

4,922 Acres

Grassland with half-shrub mixtures

4,747 Acres

Grassland with woodland mixtures

1,326 Acres

Sagebrush

28,899 Acres

Brush (desert and mountain)

53,015 Acres

Woodland (Pinon-juniper and oakbrush)

17,621 Acres

Commercial timber to include aspen

51,676 Acres

River bottom vegetation!/

678 Acres

Urban];'/

411 Acres

Mineral 1/

2,130 Acres

Nisce1laneous

!:)

24,164 Acres

Total

189,589 Acres
(296.5 sq. miles)

Cover type categories were derived from 1/2" = 1 mile S.C.S. Land Use
and Cover Type Maps, 1953, for Garfie14 County. A planimeter and grid
were used to derive and compute acreage. Total land acreage differs
from total shown in Unit Description Section (192,416 or 300.7 sq. miles).

1/

- Includes only woods and brush-woods mix along the Colorado River as planimetered from USGS 1:24,000 topographic maps (Rifle, Rulison, and Grand
Valley Quadrangles).
2/
- Urban - Those areas used for residential and municipal purposes as
illustrated on the 1/2" = 1 mile S.C.S. Cover Type and Land Use Haps for
the towns of Grand Valley and Rifle.
3/
- Mineral - Those areas where mining facilities exist for the purpose of
extracting oil shale or other minerals. This does not reflect total land
ownership.

4/

- Miscellaneous - Interpreted to consist mainly of bare ground with sparse,
scattered vegetation of various types.
Paul H. Neil
July 1973

�-124-

HUMAN POPULATION - WILDLIFE MANAGEMENT UNIT 32

Specific human population data for the unit are impossible to obtain
because U.S. Census Bureau records are on a County-wide basis only and
because the east boundary splits the to\VDof Rifle.

Instead, urban and

rural lands data and trends for 1960 and 1970 are presented in total for
Garfield County.

All tOvms, regardless of size, are here considered

urban with remaining population being rural.

By comparing 1970 figures with those of 1960 in Table 1, it is apparent
that rural populations gained three times that of municipalities.

Grand

Valley and Rifle, the only towns in Unit 32, experienced markedly smaller
gains than did towns in the east and south half of the County.

Garfield County contains 3,000 square miles and 4.95 people per square
mile (1970) including municipalities.

Applying this ratio to Unit 32

(300 sq. mi.) gives approximately 1,486 people.

Since approximately one-

half the population of Rifle reside west of the unit boundary (Hf.ghway
a figure of 1,075 results.

13),

This, p Lus 270 Grand Valley residents, gives

an urban population figure of 1,345 and 141 rural residents.

Additional analysis by the Soil Conservation Service of the population of
a portion of Garfield County encompassing the Colorado River "sub-basin"
known as Rifle-West Divide are presented on page 6 of the following report:
U. S. Department of Agriculture.

1965.

Water and related

land resources Colorado River Basin in Colorado.

Coop. Study

Rep. of Colorado Water Conserv. Board and U.S.D.A. Econ. Res.
Service, Forest Service, and Soil Conserv. Service.
Denver.

183 pp. (processed).

May.

�-125-

The Colorado State Planning Office (pers. comm. 1973) provided the
following advance projections

for Garfield County:

1970 - 14,821,

1975 - 17,018, 1980 - 19,277.

w. T. McKean
"July 1973

�Table 1.
32 ]:./.

Populations

and trends of Garfield County and municipalities

County Population

City POEu1ation

Rural POEu1ation

%

Garfield

involved, in part, with Unit

1960

1970

Change

12,017

14,821

+ 18.9

l/

%

1960

1970

Change

2,135

2,150

+ 0.6

Grd. Valley

245

270

+ 9.2

Silt

384

434

+ 11.5

New Castle

447

499

+ 10.4

Glenwood Sp.3,637

4,106

+ 11.4

726

+ 15.2

8,185

+ 10.2

Rifle

Garbonda1e

612

1970

Change

-

-

-

-

-

4,557

6,636

+ 30.2

e

Total

7 ,460

1/
- Sources are the U.S. Bur. of the Census; Colorado Yearbook, 1959-61; Colorado State Planning Office
(pers. comm. 1973).
2/
- County rural population
above (urban population).

x

1960

is based upon total county population less total of towns and cities listed

I
•....

N
0\
I

�-127-

GANE SPECIES - vlILDLIFE MANAGE.':1ENT
UNIT 32

Big game mammals 11
Black bear (Ursus americanus) Uncommon
Elk (Cervus canadensis) Common.
.
Mountain lion (Felis concalor) Uncommon.
Mule deer (Odocill;;s hemi-onus) Common.
Small game mammals 11
Cottontail rabbit (Sylvilagus audubonii; S. nuttallii) Common.
Pine (red) squirrel (Tamiasciurus hudsoni~us) Common.
Snowshoe.hare (Lepus americanus) Common.

Game birds 1./
Migratory waterfowl and shorebirds
Great Basin Canada goose (Branta canadensis moffitti) Common yearlong
resident.
Black brant (Branta nigricans) 11 Possible rare migrant.
White-fronted goose (Branta a1bifrons frontalis) 31 Possible rare migrant.
Snow goosc(Chen caeru~~s
caeru1escens) 11, ~/: Possible rare migrant.
Mallard (Anas platyrhynchs&gt;s platvrhvnchos) Common resident.
Cadwa Ll, (Anas strepe.ra) Common spring and fall migrant.

11

-"Nomenclature according to Lechleitner, R. R. 1969.
Colorado. Pruett Publishing Co., Boulder. 254 pp.

Wild mammals of

1./Nomenclature from Bailey, A. M., and R. J. Niedrach. 1967. Pictorial checklist of Colorado birds. Denver Mus. Nat. Hist. 168 pp. Information on occurrence
and status adapted from the above reference and Cringan, A. T., and L. Carlson.
1973. Wildlife in the Piceance Creek Basin, In: An environmental reconnaissance
of the Piceance Basin, Rio Blanco and Garfield counties, Colorado. A report on
the completion of Part 1, Phase One of the environmental inventory, analysis and
impact study portion of the Regional Oil Shale Study being done for the State of
Colorado by the Thorne Ecological Institute, Boulder, Colorado, 144 pp. Additional information on occurrence, in employing the term "possible", is adapted
from the foregoing references and Davis, W. A. 1969. Birds in western Colorado.
Colo. Field Ornithologists. 61 pp. Where adjective "possible" is absent, actual
sightings have been reported verbally by anyone or more Division personnel
Glenn E. Rogers, Claude E. \~lite, George E. Steele, Furman W. Dunham, Don Crane,
or qualified by additional footnotes that follow.

�-128-

Game birds (continued)

Pintail (Anas acuta) Common spring and fall migrant.
American green-winged teal (Anas crecca carolinensis) Common migrant.
and uncommon yearlong resident if.
Blue-winged teal (Anas discors discors) Common migrant.
Cinnamon teal (Ana;-CYanoptera sententrionalium) Common migrant.
American wigeon (Anas americana) il Common migrant and rare winter resident.
Northern shoveler (k1as clvpeata) il Common migrant and uncommon summer
resident.
~~ood duck (Aix sponsa) 1/ Possible rare migrant.
Redhead (Aythya americana) Common migrant.
Ring-necked duck (Aythya collaris) Common migrant.
Canvasback (Aythya valisineria) Uncommon to rare migrant.
Greater scaup (Aythya marila nearctica) 11 Rare migrant.
Lesser scaup (Aythya affi.nis) Common migrant.
Common goldeneye (Bucephala clangula americana) Common migrant and
winter resident.
Barrow's goldeneye (Bucephala islandica) 11 Rare winter visitor.
Bufflehead (Bucephala albeola) Uncommon spring and fall migrant and
rare winter resident.
Ruddy duck (Oxyura jamaicensis rubida) Common migrant and occasional
summer resident.
Hooded merganser (Lophodytes cucullatus) Rare winter visitor on river.
Common merganser (Mergus merganser americanus) Common winter resident.
Red-breasted merganser (Mergus serratorserrator) Uncommon winter
resident.
American coot (Fulica americana americana) Common migrant and occasional
summer resident .
.COIDre0nWilson's snipe (Capella gallinago delicata) Common migrant and
rare winter resident.

Upland game birds
Blue grouse (Dendragapus obscurus obscurus) Common.
Sage grouse (Centrocercus urophasianus urophasianus) Uncommon to common.
Ring-necked pheasant (Fhasianus colchicus) Uncommon.
Chukar (Alectoris chukar) 41 Common.
Band-tailed pigeon (Columb~ fasciata fasciata) Uncommon summer migrant.
Mourning dove (Zenafda macroura marginella) il Common summer resident.
Gambel's quail (Lophortyx gambeHi sanus) Unknown.
Wild turkey (Mel~i~llopavo
merriami) Uncertain.

11 Unverified

in hunters' bag checks but possible rare migrant and legal

game 1971-72.
i!.IChangesin nomenclature follow the thirty-second supplement to the
American Ornithologists Union check-list of North American birds published
in Auk 90:411-419, April, 1973.
W. T. McKean &amp; P. H. Neil
July 1973

�-129-

OTHER MAHMALIAN

SPECIES 1.1 - WILDLIFE MANAGE1ENT UNIT 32

Furbearers J:./
Beaver (Castor canadensis) Common.
Mink (Hustela vison) Uncertain.
Muskrat (Ondatra zihethicus) Uncommon.
Ringtail (Bassariscus astutus) Uncommon.
Weasels (Mustela erminea; M. frenata) ~. erminea Uncertain; M. frenata
Uncommon.
Norigamemammals !I
Coyote (Canis latrans) Common.
Red fox (Vulpes fulva) Uncommon.
Gray fox (Urocyon cinereoargenteus) Uncommon.
Raccoon (Procvon lotor) Uncommon.
American badger (Taxidea taxus) Common to uncommon.
Spotted skunk (Spilogale putorius) Uncommon.
Striped skunk O'lephitis mephitis) Common.
Bobcat (wildcat) (Lvnx rufus) Common.
White-tailed jack rabbit (Lepus townsendii) Common.
Black-tailed jack rabbit (Lepus californicus) Uncommon.
Yellow-bellied marmot (Marmota flaviventris) Common.
White-tailed prairie dog (Cynomys leucurus) Uncommon.
Richardson's ground squirrel (Spermophilus richardsonii) Common.
Thirteen-lined ground squirrel (Spermophilus tridecemlineatus) Common.
Rock squirre~ (Spermophilus variegatus) Common.
Golden-mantled ground squirrel (Spermophilus lateralis) Common.
White-tailed antelope squirrel (Ammosuermophilus leucurus) Uncommon.
Least chipmunk (Eutamias minimus) Common.
Colorado chipmunk (Eutamias guadrivittatus) Common.
Uinta chipmunk (Eutamias umbrinus) Uncommon to uncertain.

11
- These species, grouped separately as "Furbearers" and "Nongame mammals"
and outside of "game" categories, follow Chapter 62, Colo. Rev. Statutes
1963 As Amended, in Colo. Game, Fish and Parks Div. Laws and Regulations
Hdbk., 1970 (Art. 1, Item 3, Definitions, p. 3).

21

- Nomenclature according to Lechleitner, R. R. 1969.
Wild mammals of
Colorado. Pruett Publishing Co., Boulder, Colo. 254 pp. and Armstrong,
D. M. 1972.
Distribution of mammals in Colorado. Monograph of the
Museum of Natural History, the Univ. of Kansas, Number 3, 1972. 415 pp.

W. T. McKean
July 1973

�-130-

NONGAME YI.l\}1HALS
NOT DEFINED
BY STATUTE - WILDLIFE J"Y,NAGEMENTUNIT 32 1:./

Water shrew (Sorex .£..~.ll1stris)
Common.
Vagrant shrew (Sorex vaz rans ) Common.
Merriams shrew (Sorex nerriami) 2/ Rare.
Has ked shrew (So~-inereus)
Co~on.
Townsend's big-eared bat (Plecotus to,vnsendii) Common.
Pallid bat (Antrozous pallidus) l/ Rare.
Spotted bat (Euderma maculatum) 1/ Uncertain - rare.
Silver-haired bat (Lasionycteris noctivagans) Common.
Hoary bat (Lasiurus cine~eus) Uncommon ~ common.
Eig brown bat (Eptesicus fuscus) Common.
Western pipistrelle (Pipistrellus hesperus) Common.
Long-legged ;nyotis (Hyotis volans) Uncommon.
California myotis (Hvotis .californicus) Cornmon.
Small-footed myo t Ls (Hyot Ls leibii) Common.
Fringed myotis (Myotis thysanodes) l/ Uncertain - rare
Long-eared myotis (Hyotis evotis) Uncertain.
Little brown myotis (:-~votis
LucLfugus ) Uncertain.
Brazilian free-tailed bat (Tadarida braziliensis) 2/ Uncertain - rare.
Northern pocket gopher (Thomomys talpoides) Common~
Ord's kangaroo rat (Dipodomys ordii) Uncommon.
Apache pocket mouse (Perognathus apache) Uneommon.
Western harvest mouse (Reithrodontomys megalotis) Uncommon.
Canyon mouse (Peromyscus crinitus) Common.
Deer mouse (Peromyscus maniculatus) Common.
Pinon mouse (Peromyscus truei) Common.
Bushy-tailed wood rat (Neotoma cinerea) Common.
Gapper's red-backed vole (Clethrionomys gapperi) Common.
Meadow vole (Hicrotu~ pennsylvanicus) Uncommon.
Montane vole (Hicrotus I:1ontanus)Uncertain.
Long-tailed vole (Hicrotus longicaudus) Common
Sagebrush vole (Lagurus curtatus) Uncertain.
House mouse (Hus musculus) Uncertain.
Western jumping mouse (Zanus princeps) Common.
Porcupine (Erethizon dorsatum) Cowmon.
l/Nomenclature from Lechleitner, R. R. 1969. Wild mammals of Colorado. Pruett.
Publishing Co., Boulder. 254 pp. Information on occurrence and status from
the above reference and: Cringan, A. T., and L. Carlson. 1973. Wildlife in the
Piceance Creek Basin, In: an environmental reconnaissance of the Piceance Basin,
Rio Blanco and Garfield counties, Colorado. A report on the completion of Part
1, Phase One of the envirornnental inventory, analysis and impact study portion
of the Regional Oil Shale Study being done for the State of Colorado by the
Thorne Ecological Institute, Boulder, Colorado, 144 pp.
~/Occurrence listed by the Colo. Div. of Wildlife as extremely unusual - very
few documented records within the past decade. (1972 Status Evaluation for
Selected Colorado Species) appended to: 1973 Wildlife Operations Work Plan,
Field Order No. 4 - 1973.

�-131-

OTHER AVIAN SPECIES 1./ - WILDLIFE l-1ANAGEMENTUNIT 32

Nongame birds J:...I
Common loon (Gavia immer) Possible rare migrant.
Horned grebe (Podlceps auritus cornutus) Possible rare migrant.
Eared grebe (Podiceps nigricollis californicus) Possible cornmon migrant 11.
Western grebe (Aechmophorus occidentalis) Possible rare migrant.
Pied-billed grebe (Podilymbus podiccns podiceps) Possible uncommon migrant
and rare summer resident.
Double-crested cormorant (Phalacrocorax auritus auritus) Possible rare
migrant.
Great blue heron (Ardea herodias treganzai) Common summer resident ~/, II.
Snowy egret (Egretta !hula brewsteri) Possible uncommon summer resident 11.
Black-crowned night heron (Nycticorax nycticorax hoactli) Possible common
summer resident.
Least bittern (Ixobrvchus exilis exilis) Possible rare summer migrant.
American bittern (Bot~urus lenti?inosus) Possible rare summer migrant.
White-faced ibis (Plegadis chihi) Possible rare migrant.
Whistling swan (Olor columbianus) Possible uncommon migrant ..
Sharp-tailed gro~(Pedioecetes
phasianellus columbianus) Possible
resident 4/.
Sandhill crane (Grus canadensis canadensis; ~. £. tabida) Possible regular
migrant.
0
Vi~ginia rail (Rallus limicola limicola) Possible uncommon summer resident.
Sora (Porzana carolina) Possible uncommon summer resident.
Semipalmated plover (Charadrius semipalmatus) Possible rare migrant.
Killdeer (Charadius vociferus vociferus) Common summer resident 6/, 71 and
rare winter resident.
-

_11These species, grouped separately as IlNongame birdsll and IlRaptores" and outside
of "game" categories, follow Chapter 62, Colo. Rev. Statutes 1963 As Amended, in
Colo. Game, Fish and Parks Div. Laws and Regulations Hdbk., 1970. (Art. 1, item
3, Definitions, p. 327).
J:...I

Nomenclature from Bailey, A. M., and R. J. Niedrach. 1967. Pictorial checklist of Colorado birds. Denver Mus. Nat. Hist. 168 pp. Information on occurrence
and status adapted from the above reference and Cringan, A. T., and L. Carlson.
1973. Wildlife in the Piceance Creek Basin, In: An environmental reconnaissance
of the Piceance Basin, Rio Blanco and Garfield counties, Colorado. A report on
the completion of Part 1, Phase One of the environmental inventory, analysis and
impact study portion of the Regional Oil Shale Study being done for the State of
Colorado by the Thorne Ecological Institute, Boulder, Colorado, 144 pp. Additional information on occurrence, in employing the term "possible", is adapted
from the foregoing references and Davis, W. A. 1969. Birds in western Colorado.
Colo. Field Ornithologists. 61 pp. Where adjective "possible" is absent, actual
sightings have been reported verbally by anyone or more Division personnel
Glenn E. Rogers, Claude E. White, George E. Steele, Furman W. Dunham, Don Crane,
or qualified by additional footnotes that follow.

�-132Nongame birds (continued)
Mountain plover (Charadrius montanus) Possible rare migrant 3/.
Black-bellied plover (Pluvi~alis s(Juatarola) Possible uncommon migrant 3/.
Long-billed curlew (Numenius americanus americanus) Possible rare migr;nt.
Spotted sandpiper (Arctitis macularia) Possible common summer resident.
Solitary sandpiper (Tringa solitaria cinnamomea) Possible common migrant
and occasional summer res Lden t •
Willet (Catoptrophorus semiDalmatus inornatus) Possible rare migrant.
Greater ye1lowlegs (Tringa r.lelanoleuca)Possible common migrant 3/.
Lesser yellow1egs (Tringa flavi~es) Possible uncommon migrant 3/~
Red knot (Calidris canutus rufa) Possible rare migrant.
Pectoral sandpiper (Ca1idris melanotos) Possible rare migrant 1/.
Baird's sandpiper (Calidris bairdii) Possible common migrant 1/.
Least sandpiper (Calidris minutilla) Possible common migrant 1/.
Long-billed dowitcher (Limnodromus scolopaceus) Possible uncommon migrant.
Stilt sandpiper (f.1icroDalmahimantopus) Possible rare migrant.
Semipalmated sandpiper (Calidris pusilla) Possible rare migrant 1/.
Western sandpiper (Calidris mauri) Possible uncommon migrant 1/.
Marbled godwit (Limosa fedoa) Possible rare spring migrant.
Sanderling (Ca1idris alba) Possible rare migrant 1/.
American avocet (Recurvirostra a~ericana) Possible rare migrant.
Black-necked stilt (Eimantopus mexicanus) Possible rare migrant.
Wilson's phalarope (Steganopus tricolor) Possible common migrant and uncommon summer resident.
Northern phalarope (Lobipes lobatus) Possible uncommon migrant.
Pomarine jaeger (Stercorarius pomarinus) Possible rare migrant.
Herring gull (Larus argentatus smithsonianus) Possible uncommon migrant.
California gull (Larus ca1ifornicus) Possible rare migrant.
Ring-billed gull (Larus dela~varensis) Possible uncommon migrant.
Franklin's gull (Larus pipixcan) Possible uncommon migrant.
Bonaparte's gull (Larus philadelphia) Possible rare migrant.
Sabine's gull (Xema sabini sabini) Possible rare migrant.
Forster's tern (Sterna forsteri) Possible rare migrant.
Common tern (Sterna hirundo hirundo) Possible rare migrant •
.Least tern (Sterna albafrons atha1assos) Possible rare migrant.
Black tern (Ch1idonias niger surinamensis) Possible rare migrant.
Rock dove (Columba livia) Possible common resident.
White-winged dove (Zenaida asiatica mearnsi) Possible rare migrant.
Yellow-billed cuckoo (Coccyzus americanus americanus) Possible uncommon
summer residen t ,
Poor-will (Phalaenoptilus nutta1lii nutta11ii) Common summer resident 2/, ~.

l/Changes in nomenclature follow the Thirty-second Supplement to the American
Ornithologists Union Checklist of North American Birds published in Auk 90(2):
411-419. April, 1973.

4/
- Technically a game bird, but comparative rarity makes occurrence in Unit 32
questionable. Species is cited here to avoid its complete omission, since
chances appear good enough for it to Occur in the unit sometime.

�-133-

Nongame birds (continued)
Common nighthawk (Chordeiles minor hesperis; ~. m. howelli) Common
summer resident 6/.
White-throated swift (Aeronautes saxata1is sclateri) Common summer
resident 6/, 7/.
Black-chinned hu;mingbird (Archilochus alexandri) Common summer
resident 7/.
Broad-tailed-h~~ingbird
(Selasphorus platycercus platycercus) Common
summer resident 5/,6/, 7/.
Rufous hummingbird (Sel;sphorus rufus) Common late summer migrant Jj.
Calliope hUTI1.'11ingbird
(Stellula calliope) Possible rare migrant and
summer resident.
Rivoli's hummingbird (Eugenes fulgens aureoviridis) Possible rare summer
visitor.
Belted kingfisher (Hegaceryle alcyon alcvon) Possible connnon resident.
Common flicker (Colaptes auratus collaris) Common resident 1../, ~../, §../,Il .
(C. a. luteus) Possible rare migrant 3/.
Le,vi;';:;oodpecker(Asyndesmus Lewi.s ) Co~on summer resident Jj.
Yellow-bellied sapsucker (Sphyrapicus varius nuchaliae) Connnon summer 2/,
6/, 7/ and possible occasional winter resident.
William;on's sapsucker (Sphyrapicus thyroideus natalia€!) Possible common
summer resident.
Hairy woodpecker (Dendrocopos vi110sus montico1a) Common resident I/·
Downy woodpecker (Dendrocopos pubescens leucurus) Common resident 5/, §../,7/.
Northern three-toed woodpecker (Picoides tridactylu8 dorsalis) Possible
rare migrant.
Eastern kingbird (Tyrannus tyrannus) Uncommon sumnler resident 7/.
Western kingbird (Tyrannus verticalis) Common summer resident I/·
Cassin's kingbird (Tvrannus vociferans vociferans) Possible unconnnon
summer resident.
Ash-throated flycatcher (Hyiarchus cinerascens cinerascens) Common summer
resident 7/.
Say's phoebe-(Sayornis saya saya) Common summer I/ and possible rare
winter resident.
Willow flycatcher (Empidonax trail1ii) Common summer resident I/·
Hammond's flycatcher (Empidonax hammondii) Possible migrant and uncommon
summer resident.
Dusky flycatcher (Empidonax oberho1seri) Summer resident 7/.
Gray flycatcher (Empidonax wrightii) Possible summer resident.
Western flycatcher (Empidonax difficilis hellmavri) Common summer resident I/.

51

- Sight record given in unpublished checklist of birds of Naval Oil Shale
Reserve, 1969-70, by L. M. Stevens.
§../Sightrecord taken from Cringan, A. T. 1973. Annotated list of birds knmm to
occur in northwestern Colorado. 17 pp. In: The Colony environmental study, Parachute Creek, Garfield County, Colorado. Prepared by Thorne Ecological Institute,
Boulder, Colorado. August, 1973. Chapt. VII. Part II. Vol. 2, pp. 17-33.
I/Sight record taken from Smith, A. G. 1973. Avian envirorunental inventory and
impact study for Colony Development Operation in Garfield County, Colorado. Part 1.
Environmental inventory by Thorne Ecological Institute for Colony Development Operation, Atlantic-Richfield Company, Operator, October, 1973. (processed).

�-134Nongame

birds

(continued)

\-lesternwood peewee (Contopus sordidulus veliei) Common summer resident 5/.
Olive-sided flycatcher (Nuttallornis borealts) Cornmon summer resident 7/-:
Horned lark (Eremophila alpestris leucolaema) Common resident 2/, §j. Violet-green swallow (Trachycineta thalassina lepida) Common summer
resident 5/, 6/, 7/.
Tree swa Ll.ow- (Irfdop"I="ocnebicolor) Common migrant and summer resident i/, Jj.
Bank swallow (Riparia riparia riparia) Possible uncommon migrant and
summer resident.
Rough-winged swallow (Stelgidopter\~ ruficollis serriDennis) Uncommon
migrant and s~~er
resident 7/.
Barn swa l.l.ow(Hirundo rustica ervthro9;aster) Common summer resident i/, Jj.
Cliff swallow (Petrochelidon pyrrhonota pyrrhonota) Common summer
resident 5/, 6/.
Purple marti~ (P~ogne subis subis) Possible rare summer migrant.
Gray jay (Perisoreus canadensis capitalis) Uncommon resident iI, il.
Steller's jay (Cya~ocitta stelleri macroloDha) Common resident 5/,6/,
7/.
Scrub jay (ApheLo coma coerulescens vlOodhouseii) Common resident-Sf ,-6/
Black-billed rnagpi e (Pica ~
huds on i.a) COITcDlon
resident 2/, il ,-il.Common raven (Corvus corax sinuatus) Common resident 51, 6/, 7/.
Common crow (Corvus brnhYrhvnchos
brachyrhvnchos)
Po"Ssible u~common resident.
Pinon jay (Cvmno rh tnus cyanoceohalus) Common summer 1/, iI, il and winter
resident 7/.
Clark's nutc"I="acker(Nucifraga coltL.':lbiana)
Common resident 5/, 6/, 7/.
Black-capped chickadee (Parus articapillus garrinus) Comrno~ re"Side~t 2/, i/, Jj.
Mountain chickadee (Parus gambeli gambeli) Common resident 5/, 6/, 7/.
Plain titmouse (Paros inornatus rid~Hayi) Common resident
Bushtit (Psaltriparus minimus p Lumbeus ) Possible common re"Sident 3/.
~~ite-breasted
nuthatch (Sitta-carolinensis
nelsoni) Uncommon reSident i/.
Red-breasted nuthatch (Sitta canadensis) Rare resident 5/.
Pygmy nuthatch '(Sitta pygmaea melanotis) Possible unco~on
resident.
Brown creeper (Certhia familiaris montana) Uncommon resident and common
migrant 7/.
Dipper (Cinclus mexicanus unicolor) Common resident 2/, iI, Jj.
House wre n (Troglodytes aedon par kmaru.L) COTTU:lon
summer resident 'i/, ii, il.
Bew Lck s wren (Thryomanes EeVlickii,;.
_eremoDhilus) Possible common summer
resident and rare winter resident.
Long-billed marsh wren (Telmatodytes Dalustris plesius) Possible rare
winter-resident.
Canyon wren (Salpinctes mexicanus conspersus) Possible uncommon summer resident.
Rock wren (Salpinctes obsoletus .obsoletus) Common summer iI, and possible
rare winter resident.
Hockingbird
(Nimus ~lottos
leucopterus) Possible uncommon summer resident.
Gray catbird (Dumetella carolil£nsis) Rare summer resident 3/, 7/.
Sage thrasher (Oreoscoptes montanus) Possible common summer-resident.
Americari robin (Turdus migratorius propinquus) Common summer and winter
resident 3/, 51, 6/, 7/.
Hermit thrush (C;tha"I="us
guttatus audubonii) Common summer resident 1/, 'i..l, II.
Swainson's thrush (Catharus ustu13tus almae) Uncommon migrant 7/.
Veery (Catharus fucescenssalicicola)
Common migrant and summer
resident ]}: J].
Western bluebi.rd (Sialia mexicana bairdi) Common migrant and uncommon summer
resident 7/.
Mountain blu;'bird (Sialia currucoides) Common migrant and SUmmer resident
51, 6/, 71 and possible occasional winter resident.
Townsend's "S"olitaire (Hyad e.st es tmmsendi
townsendi) Uncommon resident

,2/.

77. -

i

'i/. iI, 7/.

.

-

�-135-

Nongame birds (continued)
Blue-gray gnatcatcher (Polioptila caerulea amoenissima) Common summer
resident 7/.
Golden-crowned kinglet (Regulus satrapa amoenus) Possible uncommon
migrant and rare SQmmer resident.
Ruby-crowned kinglet (Regulus calendula cineraceus) Common migrant ~/, 7/.
Bohemian waxwing (Bombycilla garrulus pallidiceps) Possible irregular
winter migrant 3/.
Cedar'vaxwing (Bombvcilla cedrorum) Uncommon resident 7/.
Northern shrike (Lanius excubitor invictus) Possible c;mmon winter resident.
Loggerhead shrike (Lanius ludovicianus excubitorides) Possible uncommon
summer and common \vinter resident.
Starling (Sturnus vulgaris vulgaris) Common resident 6/, 7/.
Gray vireo (Vireo vicinior) Uncommon summer resident 71. Solitary vireo (Vireo solitarius plumbeus) Common sum;er resident 2/.
Red-eyed vireo (Vireo olivaceus) Possible rare summer resident.
Warbling vireo (Vireo gih'Us swa'i.ns on i i.) Common summer resident 2/·
Tennessee warbLer (Vermivora peregrina) Possible rare but reguLar'migrant.
Orange-crowned warbler (Vermivora celata orestera) Uncommon migrant and
summer resident ~/.
Nashville warbler (Vermivora rufic.apilla ridgHayi) Possible rare migrant.
Virginia's warbler (Vermivora virginiac) COI"1Ilon
summer resident 7/.
Yellow warbler (Dendroica petechia aest Lva) Common summer reside;-t '2/' §j, JJ.
Ye l.Low+rumpedwarb Ler (Dendroica coronata memorabilis) Common summer
resident 3/, 6/, 7/; (D. c. coronata) Possible common migrant 3/.
Black-throated gray warbler (Dendroica nigrescens) Common summer resident 1/·
Townsend's warbler (Dendroica t ownsend i ) Uncommon fall migrant 7/.
MacGillivray's warbler (Oporornis tolmiei monticola) Co~.mon su~er resident

}.j,JJ.

Common yellowthroat (Geothlypis trichas occidentalis; Q. ~. campicola)
Uncommon summer resident 3/, 6/, 7/.
Yellow-breasted chat (Icteri~ virens-auricollis) Possible common summer
resident.
Wilson's warbler (Wilsonia pusilla pileolata) Common migrant and summer
resident.
American redstart (Setophaga ruticilla tricolor a) Possible rare migrant.
House sparrow (Passer domesticus domesticus) Common resident 6/, 7/.
Bobolink (Dolichonyx oryzivorus) Possible rare summer migrant~
Western meadowlark (Sturnella neglecta neglecta) Common summer ~/, II and
possible uncommon wi n t er resident.
Yellow-headed blackbird (Xanthocephalus xanthocephalus) Common summer
resident 6/, 7/.
Red-wt.nged blackbird (Agelaius phoeniceus. fortis) Common resident ~/, II.
Northern oriole (Icterus galhula bullockii) Common summer resident 1/, ~/, 1/·
Rusty blackbird (Euphagus carolinus caroJ.inus) Possible rare winter migrant.
Brewer's blackbird (Euphagus cyanocephalus) Common resident ~/, 1/·
Bro"tvn-headedcowbird (Molothrus ater artemisiae) Common summer resident 6/, 7/.
Western tanager (Piranga ludovic~)
Co~~on migrant and summer resident-II.Scarlet tanager (Piranga olivacea) Possible rare migrant.
Hepatic tanager (Piranga flava dextra) Extremely rare straggler II. (Record
observation).
Black-headed grosbeak (Pheucticus melanocephalus melanocephalus) Common
summer resident 5/.
Blue grosbeak (Guiraca cacrulea interfusa) Possible uncommon summer resident.
Lazuli bunting (Passerina amocna) Common summer resident 1/·
Evening grosbeak (llesperiphonavespertina hrooksi) Common winter
resident ~/, 1/.

�-136-

Nongame birds (continued)
Cassin's finch (Carpodacus cassinii) .Possible common resident.
House finch (Carnodacus mexicanus frontalis) Common summer 5/, lj and
possible uncommon winter resident.
Pine grosbeak (Pinicola enucleator montana) Possible uncommon resident.
Gray-crowned rosy finch (Leucosticte tephrocotis tephrocotis; L • ..!:..
littoral is) Possible common winter resident.
Black rosy finch (Leucosticte atrata) Possible common winter migrant~
Brown-capped rosy finch (Leucosticte australis) Possible common winter
migrant.
Common redpoll (Acanthis flammea flammea) Possible rare winter migrant.
Pine siskin (Spinus Dinus pinus) Common resident 2/'
American goldfinch (~inus tristus tristus; ~. ~. pallidus) Possible
common summer and unco~~on winter resident.
Lesser goldfinch (Spinus psaltria psaltria) Possible uncommon summer
and rare winter resident.
Red crossbill (Loxia curvirostra) Possible rare, irregular resident •.
White-\vinged crossbill (Loxia leucoptera leucoptera) Possible rare
winter migrant.
Green-tailed towhee (Chlorura chlorura) Common smnmer resident 5/, 6/, 7/.
Rufous-sided towhee (PiDilo erythrophthalmus montanus) Common s~mer and
rare winter resident 6/, 7/.
Lark bunting (Calamospiza melanocorys) Possible uncommon summer resident.
Savannah sparrow (Passerculus sandwichensis nevadensis; P. s. anthinus)
Common migrant and summer resident 6/, 7/.
- Grasshopper sparrow (Ammodramus savann-;rrmn-perpallidus)Uncommon summer
resident 5/, 7/.
Vesper sparrow (Pooecetes gramineus confinis) Common migrant and summer
resident 5/, 7/.
Lark sparrow-(Chondestes grammacus strigatus) Possible common migrant and
summer resLdent ,
Black-throated sparrow (Amphispiza bilineata deserticola) Common summer
resident 7/.
Sage sparrow-(Amphispiza belli nevadensis) Common summer resident 2/.
Dark-eyed junco (Junco hyemalis aikeni) Possible rare winter migrant 1/;
(~. h. hyemalis; ~. ~. cismontanus) Rare winter resident 1/, 2/;
(J. h. oreganus ) Common winter resident 3/, 7/; (J. h. o. var , mearnsi)
Common winter resident 3/, 7/.
- - Gray-headed junco (Junco c~nic;ps caniceps) Common summer 2/ and winter
resident 6/, 7/.
Tree sparrow-(Spizellaarborea
ochracea) Possible uncommon winter migrant.
Clay-colored sparrow (Spizella pallida) Common migrant I/.
Chipping sparrow (Spizella passerina boreophila) Common summer resident

6/, 7/.

Brewer's sparrow (Spizella bre\veri breweri) Common summer resident 7/.
Harris' sparrow (Zonotrichia querula) Possible uncommon migrant and rare
winter resident.
White-crowned sparrow (Zonotrichia leucroDhrvs) Common resident 5/, ~/.
White-throated sparrow (Zonotrichia albicollis) Rare migrant 7/.Fox sparrow (Passerella iliaca schistacea) Rare summer resident 5/.
Lincoln's sparrow C·1elospiza lincolnii alticola) Common migrant ~d
summer resident 7/.
Song sparrow (Helospiza melodia) Corrnnonsummer 2/ and possible uncommon
winter resident.
Lapland longspure (Calcarius lapponicus alascensis) Possible rare winter
migrant.

�-137Raptores '!:./
Turkey vulture (Cathartes aura meridionalis) Common summer 2/, 2..1, J)
and possible rare winter resident.
Goshawk (Accipiter p;catl lis atricapi Ll.us) Uncommon resident 2..1, J).
Sharp-shinned hawk (.Accipiter striatus ve l.ox). Rare summer and common
winter resident 7/.
Cooper's haHk (Accipiter cooperii) Uncommon summer II and common winter
resident 7/.
Red-tailed h;wk (Buteo l£maicensis calurus) Common resident l/, 2../, ZI.
Sw.ainson's hawk ~teo
swa LnsonL) Uncommon summer l/,
Z/. and possible
rare winter resident.
Rough-legged hawk (Buteo lagopus ~. johannis) Possible rare summer and
uncommon ,,,interresident or migrant.
Ferruginous hat....
k (Buteo_regalis) Rare summer and common winter resident ZI.
Golden eagle (Aquila chrvsaetos canadensis) Common resident 6/, 7/, 8/ •
.Bald eagle (Haliaeetus lcucocephalus a.l ascanus ) Common winte~ re;ide;-t 2..1,

Z/,

§./.

Marsh hawk (Circus cyaneus hudsonius) Common summer 5/,6/,
7/, and uncommon wi.nt er resident 7/.
Osprey (Pandion haliaetus ~arolinensis) Possible rare migrant.
Prairie falcon (Falco mexicanus) Rare resident 7/.
Peregrine falcon (Falco peregrinus anatum) Possible rare migrant, endangered.
Pigeon hawk (Falco columbarius) Rare winter migrant 7/.
Sparrow hawk (Falco sparverius suarverius) Common su~er l/, 2..1, Z/ and
possible uncommon winter resident.
Screech owl (Otus asio) Possible uncommon resident.
Flammulated owl (Otus flarr~eolus fla~meolus) Possible rare summer resident.
Great horned owl (Bubo virginianus) Common resident 5/,6/,
7/.
Pygmy owl (Glaucid~gnoma
californicum) Possible r~re ~esident.
Burrowf.ng owl (Speo tyt o cunicularia hvpugaea) Common summer 5/ and possible
rare winter resident.
Long-eared owl (Asio otus wilsonianus) Uncommon resident l/.
Short-eared 0\,,1 (Asio flammeus flammeus) Possible uncommon wi.nt er migrant.
Saw-whet owl (Aeg~s
acadicus acadicus) Uncorr~on resident Z/.

~I
Golden and bald eagle specifically excluded from statutes defining "Raptore"
as cited in footnote l/ but herein listed to avoid omission.

P. H. Neil
August 1973

�-138-

DISTRIBUTION
AND ABUNDi\.r..;rCE OF BIG r,Al'1E HANMALS
WILDLIFE
NANAGEI-1ENT UNIT 32 (PARACHUTE
CREEK)

BLACK BEAR
Black bears occupy approximately the area between 6,000 and 9,000 feet
elevation in Unit 32, though on rare occasions they may wander over almost
the entire unit.

Their yearlong habitat is largely within the·timbered

or brush-covered lands above the Roan Cliffs where abundant combinations
of food are available, such as fruit-bearing shrubs, small mammals, and
carrion from livestock and big game herds.

This is a comparatively small

area, but well isolated beca.use of the tremendously abrupt topography
surrounding it (see accompanying distribution maps).

Estimated annual black bear kill for the period 1950-1972 inclusive was

3.4 head, based upon hunter report card surveys.

Annual kills ranged from

0-15; there we re eight years out of 22 when no bears were reported taken.
Bears have frequently been removed because of damage complaints, principally from woolgrowers.

In 1972 two bears were so removed (Myers pers.

corom. 1973).

ELK
Su~ner range for elk in Unit 32 lies approximately within the range 7,0009,000 feet, generally being the highest most isolated areas along the north
boundary of the Unit.

Potentially the entire Roan Plateau at this eleva-

tional range could be summer range, but actually only headwater areas as
follows have been consistently occupied: Spring Gulch eastward to Trapper
Creek and the north fork of West Fork Parachute Creek.

Herds that summer

on aforementioned ranges are also believed to interchange freely across
high ground areas of Unit 22 adjoining on the north (see accompanying maps) •

't,....
.,...
\

�-139-

Winter range for elk in Unit 32 decreases in size on the average, to only
a fraction of summer range.

In 1972-73, a severe winter, ground observa-

tion revealed signs of one small elk herd in East Parachute Creek between
the confluences of Ben Good Creek and Bull Gulch (see accompanying map).
Another group were reported (rancher) wintering in Gran1ee and Helm gulches
east of main Parachute Creek (Neil pers. comrn. 1973).

The principal winter

ranges, on average years, are on the Roan Plateau at the aforementioned
Spring Gulch-Story Gulch headwaters, and in the extreme northwest portion
of West Fork Parachute drainage.
While elk winter range area may decrease in size from summer range, the elk
population apparently increases due to migration from Units 23 and 33.
This fact has not been accurately assessed.

Denney (1965) indicated that

hunter harvest on the Roan Plateau was less than 100 animals annually.
Unit 32 yielded an estimated harvest of 26.4 elk annually for the ten seasons between 1959 and 1971 (Harvest Data) presented elsewhere in this report.
Four years were closed to elk hunting in the unit - 1958, 1962, 1963, and
1964.

This is a substantial harvest considering that Units 21, 22, 31 and

32 together yielded an estimated harvest of only 49.7 elk annually for the
same period.

Minnich (1973) indicates an "Overall Current" 1972 population

of approximately 150 head with densities ranging from 1.4 to 1.5 elk per
.square mile.

Comparable winter populations calculated were 170 head and

densities ranging from 1.1 to 914 elk per square mile depending on the site
within Unit 32.

MOUNTAIN LION
The literature available makes no specific mention of mountain lions occurring
within the specific environs of Unit 32, however, numerous occurrences are

�-140-

cited from areas to the immediate east (Unit 33) and west (Unit 31)
(Armstrong 1972).

Neil (pers. comma 1973) reported seeing a lion feeding

upon a mature doe mule deer freshly killed approximately
of the confluence

of West Parachute Creek and Light Gulch in April 1972.

Two other separate, but undetailed,

reports of lion occurrences

by members of the staff of Thorne Ecological
Colony Development
resembles

rence possible
Dixon

Operations.

Obviously,

that o~ black bear - predominantly

.the high mobility

1/2 mile north

and secretiveness

were made

Institute while working for

the better yearlong lion habitat
that above the Roan Cliffs, but

of the lion makes its undetected

in almost any portion of the unit (see accompanying

(1967) mentions

occur-

maps).

lion kill records from Garfield County in the Roan

Cliffs west of Grand Valley.

Hunter reports of lions killed during the period 1968-1972 by year are as
follows:

1968-0; 1969-0; 1970-0; 1971-0; 1972-1.

was reported taken in five years.

Thus, only one animal

No data are available for the 3-year

period 1965-1967 because no reports were required of hunters.
mountain

lions were not considered

game animals.

Before 1965

It is highly probable

that lions were being taken here by trappers, guides, livestock men, or
others and were not being reported.

MULE DEER
Over one-half of Unit 32 is unavailable
high elevation and deep snow.

Roan Plateau elevations range fromapproxi-

mately 7,500 upward to 9,200 feet.
is clearly delineated

to mule deer in winter because of

The upper winter range line in Unit 32

by the Roan Cliffs (see accompanying

maps from Minnich

�-141-

1973).

The normal or average winter range line occurs some 1/2 to four

miles below the Roan Cliff base.

A normal winter range zone of one to

three miles width follows the borders of the Unit beginning on the northeast corner (approximately T.5S, R.93W, 8.5).

This peripheral zone ends

near Grand Valley; here it turns northward up Parachute Creek and extends
in an increasingly narrow band between the cliff bases to the confluences
of East, Middle and West Parachute Creek, thence up each of these forks
approximately three miles.

The average upper winter range line varies

between 6,000 and 7,000 feet.

A lower winter range line for the Unit

coincides with State Highway 13 on the east, U. S. Highway 6 &amp; 24 on the
south and Parachute Creek bottoms on the west.

Summer deer range largely coincides with the Roan Plateau in Unit 32, witll
additional areas of upper Parachute Creek and its tributaries.

The afore-

mentioned Roan Cliffs constitute a natural boundary between upper winter
range and lower sUmmer range.

Logically, a scattered few deer subsist

yearloilg on river and creek bottom lands and adjoining cultivated fields.
Summertime interchange between Units 31 to the west and 22 to the north are
unverified but believed common.

Abundance of deer by year from one aerial trend count is available.

This

trend count (Rifle-Grand Valley) has had a seven-year (1966-1972) mean of
975 animals.

Annual census figures are available in the Denver and Regional

Game }1anagement offices.

Minnich (1973) calculated a "normal" winter range deer density of 54.2 deer
per square mile for Parachute Creek drainage and a much smaller 9.0 deer

�-142-

per square mile for the remaining south and east wintering areas of Unit
32.

An overall density of 20.4 deer per square mile was indicated.

These

figures were based upon estimates supplied by fie1dmen.

On the basis of total harvest for years 1959-1968 inclusive, Unit 32
ranked 6th statewide.

Total kill for that 10-year span was 32,677, or an

approximate average of nearly 3,268 animals.

As elsewhere in Colorado, deer

numbers declined in the late 60's and early 70's.

Kill by year for recent

years was: 1969-592; 1970-898; 1971-469; 1972-1,436.

Restrictive hunting

regulations are believed to have accounted for some of the drop in kill,
particularly in 1971.

Nonetheless, it is evident that Unit 32 has sustained

at least an above average deer population for many years.

Literature Cited

Armstrong, D. M.

1972.

Distribution of mammals in Colorado.

Univ. of

Kansas Printing Service, Lawr ence , 415.pp.

Denney, R. N.

1965.

Colorado long range species management plans,

1965-1975, for elk.

Dixon, K. R.

1967.

pp. 141-164.

61 pp.

Evaluation of effects of mountain lion predation,
In Game Res. Rep., July-Part II.

Fish and Parks.
Minnich, D. v].

Colo. Dept. Game, Fish and Parks.

1973.

Colo. Div. Game,

pp. 73-310.

Statewide mammal and bird distribution maps and

data compiled from WCO, Area Supervisor, and Regional Director input.
Located at Denver office, Colo. Div. of Wildlife. (unpublished)

W. T. McKean
July 1973

�-143-

DISTRIBUTION AND ABUNDANCE OF SMALL GAHE MAMMALS
WILDLIFE HANAGEMENT UNIT 32 (PAPV\CHUTECREEK)

COTTONTAIL RABBIT
Every square mile of Unit 32 is considered capable of supporting at least
some cottontail rabbits yearlong.

According to Lechleitner (1969) and

Armstrong (1972), both the desert cottontail and Nuttall's cottontail should
occur, the desert cottontail being common at the lower elevations (5,0007,000 feet) and the Nuttall's cottontail common on the plateau (7,000-9,000
feet).
No specific studies have been conducted regarding cottontail rabbit density
anywhe re in wes t ern Colorado, however , Shepherd (1965a) concluded after a
literature survey, that a possible high-cycle density of 150-200 animals per
square mile and a low-cycle density of 15-20 animals per square mile wez e
reasonable for much of Colorado.
32.

For abundance"as

This estimate should be applicable to Unit

reflected in kill data, see tables on Hunter Harvest

for Small Game Management Unit 8, following.
SNOWSHOE HA..~E

The sno\olshoehare is believed to occur commonly in those areas of the Roan
Plateau occupied by Douglas-fir and Douglas-fir-aspen mix.

Stevens (1970)

reported this species on the Naval Oil Shale Reserve which includes a large
area of the plateau in the eastern one-half of Unit 32.

It is reasonable

to expect their occurrence in areas of similar topography and vegetative
type in the western one-half of the unit.

Game distribution maps prepared

by Minnich (1973) and Shepherd (1965b) indicated that snowshoe hares are
welI represented on the majority of the plateau top areas within Unit 32.

�-144-

Kill data~ as an indicator of abundance~ shows that for the five year
period 1968-l972~ in Small Game }mnagement Unit 8~ an average annual harvest of 491 snowshoes by 109 hunters occurred.

This would probably re-

present the total kill in VIildlife Units 21, 22, 30, and 32 across the Roan
Plateau from the Utah line to Highway 13 north of Rifle.

Minnich (1973) calculates a population of approximately 660 hares~ based
upon estimates of field personnel , with an average density of 10.2 hares
per square mile, ranging from 0.8 to 27.5 for various small segments of their
range.

RED (PINE) SQUIR..l{EL
Limited information is available regarding the distribution of this species
within Unit 32.

According to Lechleitner (1969) and Armstrong (1972), they

should be common in the coniferous areas on the higher plateaus and mesas
throughout Garfield.County.

Neil (pers. comm. 1973) reported several obser-

vations of this species along Trappers Creek and in small groves of Douglasfir along the divide road on the plateau.

Stevens (1970) also reported

their occurrence on portions of the Naval Oil Shale Reserve.

No data on abundance are available, but it·is believed that a reasonably
stable population exists within coniferous types.

Literature Cited
Armstrong, D. M.

Distribution of mammals in Colorado.

Printing Service, Lawrence.

Lechleitner, R. R.
Boulder.

1969.

254 pp.

Univ. of Kansas

415 pp.

Wild mammals of Colorado.

Pruett Pub. Co.,

�-145-

Minnich, D. w.

1973.

Statewide mammal and bird distribution maps and

data compiled from W.C.O., Area Supervisor, and Regional Director
input.

Located at Denver office, Colo. Div. of Wildlife.

Un-

published.

Shepherd, H. R.

1965a.

Colorado long range management plans for game

species, 1965-1975, for cottontail rabbit.
Parks Dept. rep.

Shepherd, H. R.

1965b.

12 pp. (Himeo)•

Colorado long range management plans for game

species, 1965-1975, for snowshoe hares.
Dept. rep.

11 pp.

Colo. Game, Fish and

Colo. Game, Fish and Parks

(Mimeo).

P. H. Neil
September 1973

�-146-

DISTRIBUTION A}..TJ) ABUNDANCE OF Sr"'ALLGA11E BIRDS
\HLDL IFE HPv'lAGB-fENT
UNIT 32 (PARACHUTE CEEEK)

BLUE GROUSE
The distribution maps provided by Rogers (1965a, 1968) and Minnich (1973)
indicated that the majority of the Roan Plateau area within Unit 32 is
available habitat for blue grouse.

Current reliable reports (Smith 1973)

confirm their presence on the plateau near the headwaters of Trappers
Creek, East Fork and East Middle Fork of Parachute Creek and in Douglasfir and aspen types scattered along the divide road in the northern part
of the unit.

The true abundance of blue grouse in Unit 32 is not known, however, data
provided by Minnich (1973) estimates ~lue grouse density at 4.9 birds per
square mile which results in an estimated population of 908 blue grouse
in the habitable range in Unit 32.

Smith (1973) reported observing

numerous blue grouse in the unit during bird census wo rk for Colony
Development Operation

(an oil shale company).

He worked on a two week

schedule each month from February through September, 1973.

In a study done

in west slope areas comparable to Unit 32, Rogers (1968) indicated a range
of 10.3-38.7 road miles per bird.

SAGE GROUSE
Rogers (1964) listed several areas within Unit 32 where light populations
of sage grouse are present.

He reported that during summer months these

birds range from the head of Parachute Creek on the plateau west to Douglas
Pass near the Utah line.

Cringan (pers. comm. 1973) reported observing a

brood on the plateau between East Middle Fork and East Fork of Parachute
Creek during the early summer of 1971 which confirms their occasional
presence in that portion of t.heunit.

Minnich (1973) maps current distri-

�-147-

bution of Roan Plateau sage grouse including four areas within Unit 32.
Winter range for these birds is not known, but they are believed to drop
off the plateau onto the lower sagebrush mesas.

Birds have been observed

on Hubbard Mesa just northwest of the town of Rifle (Rogers 1964).

According to Rogers (1964) fall densities of sage grouse range from 1 to
10 birds per square mile in occupied areas of Unit 32, and more recent
information from l1innich (1973) estimates a density of 5.8 birds per square
mile of occupied range which yields an estimated population of 280 sage
grouse for the unit.

Smith (1973), in work cited above, reported no sage

grouse seen - probably a significant negative result.
CHUKAR
Distribution maps prepared by Sandfort (1965a) and Minnich (1973) indicated
that chukars occur at the lower elevations of Parachute Creek and in the
sage flats along Highway 6 and 24 between Rifle and Grand Valley (see
accompanying maps).
Neil (pers. comm. 1973) reported observing chukars in both of these locations
during the early summer months of 1972 and 1973 and Stevens (1970) reported
their occurrence on portions of the Naval Oil Shale Reserve, presumably in
about the same locations.

Between 1940 and 1965, 525 chukars ~ere released (4 releases - see Game Species
Introductions) near Garden Gulch in main Parachute Creek and on the Mahaffey
Ranch near Grand Valley (Carhart 1942, Jones and Rogers 1964, Sandfort 1956,
1957) and it is believed that since that time the population has declined to
approximately 250 in the two above mentioned areas of Unit 32 (Hinnich 1973).
Trend counts for the period of 1964 to 1972 also indicated a downward trend in
population.

Counts ranged from a peak of 60 in 1964 to 16 in 1972 with no

birds being observed in 1965, 1967, 1968, and 1969.
f.n !lenver and

Nor t hwe s t Game Hanaaemerrt offices.

Records may be found

�-148-

RING-NECK.ED PHEASANT
According to Ninnich (l973) a small population of·these birds range from
Rifle to the west boundary of Unit 32 along the Colorado -River and up
Parachute Creek to the confluence of East Fork being mainly confined to
the river bottom and agricultural areas (see accompanying maps).

Swope

(1~65) show~d a similar distribution and classified this as poor quality
habitat with a spring breeding population index of 0-15 based on spring
crow
counts and ~,inter sex ratio counts.
,

He have no recent records of

ring-necked pheasant trend counts to comment on their present abundance.

WILD

TlJRKEY

Hoffman (1965) showed occupied turkey range southwest of Grand Valley on
the south side of the Colorado River and to the east of Rifle on both
sides of the Colorado River,
1971).

as a result of introductions in 1961 (Burget

Recent information from local residents revealed that the wild

turkey used to be-moderately abundant around the lower Parachute Creek
drainage (Neil pers. comm. 1973), however, we have no recent records to
confirm that the ~,ild turkey is still present in the area.

It is believed that if turkeys are present in the immediate adjacent ranges
mentioned above, some drift into Unit 32 probably occurs but their numbers
are few.

A search of turkey harvest data for the years 1959-1970 revealed

that in only two years were any turkeys reported taken from Unit 32 - 1969
(2 birds) and 1965 (4 birds).

Adjoining Units 33 and 42 to the east and

south, produced a combined turkey kill averaging 28 birds during the same
period.

None were reported taken from Units 31 or 22 to the west and north,

respeclivf'ly.

�-149-

BAND-TAILED PIGEON
Braun (1970) indicated that band-tailed pigeon distribution extends
throughout Unit 32.

Being a migratory species, they usually arrive in

Colorado around April and according to Braun (1970) the timing of fall
migration is apparently related to timing of cold fronts in October.

The

availability and crop success of Gambel's oak and various ripening berries
within the unit undoubtedly influence the distribution of these birds.
The cultivated croplands, livestock feeding areas, and the few scattered
orchards in the lower Parachute Creek drainage provide a source of food
for band-tailed pigeons, especially during the breeding season (Braun 1970).
Stevens (1970) reported their occurrence on portions of the Naval Oil
o

Shale Reserve and recent reliable reports (Smith 1973) confirm their presence in the Parachute Creek area.

Little data are available on the abundance of band-tailed pigeons in Unit
32, hmV'ever,.Braun (pers . comm, 1973) suggested that they are probably uncorr~on in the area largely because of the lack of ponderosa pine in association with the Gambel's oak.

It is reasonable to expect that their densities

fluctuate in proportion with availability of food resources (berries, acorns,
new conifer buds, and insect larvae) and during migration periods.

MOURNING DOVE
Grieb (1965) indicated that only the lower elevations of Unit 32 are ineluded in the breeding range of mourning doves.

Braun (pers. counn. 1973)

also suggested that they are more counnon below 8,500 feet along the major
drainages and the Colorado River bottom, especially during migration periods.

�-150Stevens (1970) reported their occurrence on portions of the Naval Oil Shale
Reserve and current reliable reports confirm their presence in the Parachute
Creek area.

Migratory in nature, mourning

summer, and fall in Colorado.

doves are common during spring,

According to Bailey and Niedrach (1965) a

few may remain over the winter months in sheltered areas at lower elevations
in Colorado which, during mild winters, may apply to the lower elevations
of Unit 32.
Little is known of the actual abundance of mourning doves in Unit 32, however, according to Davis (1969) and Braun (1973 pers. comm.) they should
be very common during the summer months.

Minnich (1973) estimated a popu-

lation density of 3.8 doves per square mile in the main Parachute Creek
Canyon below the plateau, which yields an estimated population of 100 birds.
Data are lacking for population estimates on the plateau in Unit 32, but as
Braun (1973 pers. corom.) has suggested, fewer birds can be expected at the
higher elevations.

It is believed that the local population density within

the unit varies considerably from year to year.
GAMBEL 's QUAIL
Sandfort (1965b) indicated that the distribution of Gambel's quail is similar
to that of chukar in Unit 32.

Trend counts from 1952 and harvest data from

1955 to 1964 revealed that some birds were observed and harvested in habitable
range of Unit 32.
We have no recent reports of Gambel's quail within the unit and it is believed
that the population has declined to a point barely discernable.

Average esti-

mated harvest for Small Game Management Unit 8, which includes Big Game Unit
32, was only 61 birds for the years 1968-1970 (Funk and Tully 1971, 1972).
V!ATERFOWL
Breeding populations:

The mallard and grecn-wfnged teal are undoubtedly

the most common summer resident ducks in Unit 32, especially in those areas

�-151-

north of the Colorado River (Szymczak pers. corrm. 1973).

Grieb (1965)

indicated that the breeding range for green-winged teal extends over all
of the unit and that the mallard remains the most common nesting species
in the state.

Mallards have been observed nesting along Parachute Creek

and its major tributaries.

Small numbers of ducks also nest in isolated

beaver ponds scattered throughout the unit (Neil 1972).

The Colorado

River and adjacent irrigated farmlands \-lithassociated canals and ponds
provide food and cover for many migrating species ofwaterfo"Tl to include
gadwall, blue-winged teal, wigeon, redheads, lesser scaup, common mergansers,
American coots, and common (Wilson's) snipe.

Szymczak (pers. corom. 1973)

added that small numbers of Canada geese are beginning to pioneer up the
Colorado River from the Grand Junction area to nest in the spring.

Jobman (1967) in a duck production study on irrigated meadows and creek
bottoms of Piceance and Dry Fork Creeks, estimated a peak summer production
of approximately 17 birds per lineal mile.

Assuming that the Parachute

Creek-Colorado River wetlands are somewhat comparable to Piceance Creek a
projected population ranging between 900 and 1,000 birds results.

This seems

reasonable.
Wintering populations:

The only known areas where waterfowl, including geese,

winter within Unit 32 are in open waters along the Colorado River and
Parachute Creek including small ponds.

The principal wintering duck species

is the mallard while the majority of the geese wi.nter in the Grand Junction
area (Szymczak pers. comm. 1973).

The duck population along the Colorado river between Silt and Palisade, which
includes approximately 20 miles of river bottom in Unit 32, for the three

�-152-

year period 1971-1973, based upon aerial trend counts in January by
regional personnel, has averaged 1,483 birds ranging from 980 to 1,960.
Geese averaged 117 birds ranging from 32 to 179 (unpublished information
Colo. Div. of Wildl.).

Literature Cited

Bailey, A. M., and R. J. Niedrach.
of Nat. Hist.

Braun, C. E.

1965.

Birds of Colorado.

2 Vol., 895 pp.

1970.

Band-tailed pigeon investigations, pp , 151-171.

Game Res. Rep., Oct., Colo. Div. Game, Fish and Parks.
Burget, M. L.

1961.

Wild turkey development.

Rep., Oct., Colo. Dept. Game and Fish.

Carhart, A. H.

1942.

Commission.

Davis, W. A.

Denver Mus.

171 pp.

In Quart.
Prog.
.

pp. 33~43.

Chukar-pheasant experimental plantings.

Fed. Aid Proj. W-17~R~3.

1969.

p , 33.

In

Memo. to

5 pp.

Birds of western Colorado.

Colo. Field Ornithologist.

61 pp.

Funk, H. D~, and R. J. Tully.
1971 and 1972.

Grieb, J. R.

Colorado small game hunter harvest survey-

Colo. Div. l.Jildlife.·Misc. loose tables.

1965.

(unpublished).

Colorado long range game species management plans,

1965-1975, for migratory birds.

Colo. Dept. Game, Fish and Parks.

36 pp.

Hoffman, D. M.

1965.

Colorado long range game species management plans,

1965-1975, for wild turkey.

Colo. Dept. Game, Fish and Parks.

11 pp.

�-153-

Jobman, W. G.

1967.

Haterfow1 production and management recommendations

for state-owned lands near Little Hills Experiment Station, Meeker,
Colorado.

CSU term paper for FW 495 bx-Spec. Studies in Wildlife.

38 pp. (typewritten).

Jones, H., and G. Rogers.

1964.

Chukar planting record.

April.

Original

release record located in Sandfort's files at Fort Collins Research Center.

Minnich, D. W.

1973.

Statewide mammal and bird distribution maps and

data compiled from WCO, Area Supervisor, and Regional Director input.
Located at Denver Office, Colo. Div. of Wildlife.

Neil, P. H.

1972.

Inventory of beaver ponds, East Middle Fork, Parachute

Creek, Aug. 17, 1972.

In Colony environmental study, Parachute Creek,

Garfield County, Colorado.

Prepared for Colony Development Operation

by Thorne Ecological Institute, Boulder, Colorado.

Rogers, G. E.
16.

1964.

August, 1973.

Sage grouse investigations in Colorado.

Colo. Dept. Game, Fish and Parks.

1965.

(unpublished).

Part III.

Tech. Pub. No.

132 pp.

Colorado long range game species management plans, 1965-

1975, for blue grouse, sage grouse, sharp-tailed grouse, ptarmigan.
Dept. Game, Fish and Parks.

--- •. 1968.

16 pp.

The blue grouse in Colorado.

Dept. Game, Fish and Parks.

Sandfort, W. W.

1956.

2 pp~

Tech. Pub. No. 21.

Colo.

63 pp.

Chukar partridge release records-Northwest

adjoining areas, Dec. 9-1951-Jan. 9, 1956.
Proj. W-37-R.

Colo.

Region and

Memo. to Commission.

Fed. Aid

�-154-

1957.

Trapping and transplanting chukar partridges, pp. 31-38.

In Quart. Prog. Rep ,, July, Colo. Dept. Game and Fish.

1965a.

119 pp.

Colorado long range game species management plans,

1965-1975, for chukar partridge.

Colo. Dept. Game, Fish and Parks.

18 pp.

1965b.

Colorado long range game species management plans, 1965-

1975, for Gambel's quail.

Smith, A. G.

1973.

Colo. Dept. Game, Fish and Parks.

9 pp.

Avian environmental inventory and impact study for

Colony Development Operation in Garfield County, Colorado.

Part I.

Environmental Inventory by Thorne Ecological Institute, Boulder,
Colorado, for Colony Development Operation, Atlantic-Richfield Company,
Operator, October, 1973.

Stevens, L. M.

1970.

Checklist of birds and mammals of the Naval Oil

Shale Reserve, 1969-70.

Swope, H. M.

1965.

(processed).

4 pp.

(unpublished).

Colorado long range game species management plans,

1965-1975, for ring-necked pheasant.

Colo. Dept. Game, Fish and Parks.

54 pp.

w. T. McKean and P. H. Neil
August 1973.

�-155HUNTING PRESSURE
DEER HARVEST, SEASONS At-.rn
WILDLIFE MANAG8-ml'o'TUNIT 32
(From Colorado

YEAR

lf0NTING
PRESSURE 1...1

Big Game Harvest 1956-1972)

BUCKS

HARVEST
DOES
FAlmS

TOTAL

HUNT AND SEASON

1956

1983

1494

1028

248

2770

2ES

10/15 - 10/31

1957

2408

1374

1172

313

2859

2DM
2DM
2DM

10/1 - 10/14
10/1 - 12/31 10/15 - 11/17

1958

3694

1424

1001

320

2745

1DM
1DM

10/15 - 11/2
11/3 - 11/30

1959

3734

1652

1057

385

3094

1DM
1DM

10/17 - 11/3
11/21 - 12/6 2/

1960

1443

1031

590

211

1832

1DM

10/17 - 11/13

1961

2158

1659

1270

528

3457

3ES
1DM

10/21 - 11/8
11/18 - 12/3 ]j

1962

4532

3546

3079

971

7596

3ES
3ES

10/20 - 11/4
11/5 - 12/31

1963

1985

877

838

418

2133

1DM

10/19 - 11/7

1964

2362

1161

1050

317

2528

1DM

10/17

1965

2615

1032

1107

344

2483

2ES

10/16 - 11/5

1966

2856

1364

1299

516

3179

2ES

10/15 - 10/24

1967

2793

1795

1519

414

3728

2ES

10/21 - 11/9

1968

2320

1246

1104

297

2647

ES
2ES

10/19 - 10/21
10/22 - 11/7

1969

1108

374

141

77

592

ES

10/18 - 11/6

1970

1521

626

247

25

898

ES

10/17 - 11/6

1971

864

469

469

IDB

10/30 - l1/11

1972

1918

908

1436

ES
ES

10/14 - 10/23
12/2 - 12/10

445

1./

83

Based upon combined total resident and non-resident
expressed in number of licenses.

2/
- In part of Unit 32.
(footnotes continued

on next page)

%~

11/5

license sales and

�-156-

DEER HARVEST, SEASONS AND HUNTING PRESSURE

(Continued).

Note:

Hunt symbols explained as follows:

ES
2ES
3ES
lDB
lDM
2DM
3DA

= Either sex, one deer (one license).
= Either sex, two deer (one license)
= Either sex, three deer (2 licenses plus 3rd deer coupon on 2nd license).
= One deer, buck only (one license).
= One deer multiple, either sex (2 licenses and 2 deer per individual).
= Two deer multiple, either sex. (2 licenses and 4 deer per individual).
= Three deer, one must be antlerless (2 licenses plus 3rd deer coupon
on 2nd license).

Note:

The term lDM.evolved into 2D, HC or two deer, hunter's choice (2'
licenses and 2 deer per individual) and is synonymous.

Paul H. Neil
September 1973

�-157-

ARCHERY DEER HARVEST AND SEASONS
lHLDLIFE MANAGE!'1ENTUNIT 32
(From Colorado Big Game Harvest 1956-1972)

YEAR 1./

DOES

BUCKS

FAHNS

TOTAL J:./

HUNT MTD SEASON
10/1 - 10/14
10/15 - 10/31

1956 ]./

ES
2ES

1957

2DM 10/1 - 10/14, In Part
2D}1 10/15 - 11/17

1958

0

2

0

2

9/1 - 9/30
ES
1DM 10/15 - 11/2
1D}! 11/3 - 11/30

1959

0

2

5

7

9/15 - 9/30
ES
lDM 10/17- 11/3
1D}! 11/21 - 12/6, In Part

1960

3

2

0

5

9/10 - 9/30
ES
lD}! 10/17 - 11/13

1961

3

0

0

3

3DA
3DA
1DM
1DM

8/26 - 9/10
10/21 - 11/8
11/11 - 11/26
11/18 - 12/3, In Part
8/25 - 9/23
10/20 - 11/4
11/5 - 12/31

'"

1962

4

2

0

6

ES
3ES
3ES

1963

1

0

0

1

8/17 - 9/8
ES
1D}! 10/19 - 11/7

1964

4

0

0

4

ES
1DH

8/15 - 9/13
10/17 - 11/5

1965

10

12

6

28

ES
2ES

8/21 - 9/12
10/16 - 11/5

1966

28

~4

6

58

ES

2ES

8/20 - 9/18
10/15 - 10/24

1967

2~

10

5

37

ES
2ES

8/19 - 9/17
10/21 - 11/9

1968 i/

10

14

4

28

ES
ES
2ES

all.7

&gt;'

"'!

9/15

10/19 - 10/21
10/22 - 11/7

�-158ARCHERY DEER HARVEST AND SEASONS (Continued) •
YEAR 1./

BUCKS

DOES

FAWNS

1969 ~/

23

0

0

23

lDB
ES

8/16 - 9/14
10/18 - 11/6

1970 fl./

5

0

0

5

lDB
lDB
ES

8/15 - 8/31
9/1 - 9/20
10/17 - 11/6

1971 ]j

33

0

0

33

1DB
lDB

8/21 - 9/6
10/30 - 11/11

1972 ~/

37

0

0

37

1DB
ES
ES
ES

8/19 - 8/31
9/1 - 9/20
10/14 ..,10/23
12/2 - 12/10

TOTAL ~/

HUNT M"D

SEASON

1/
Free permits issued to holders of regular big game hunting licenses by application only. Years 1957-1960, no issue of special permits; archery hunting
'allow'ed to holders of valid deer licenses. Years 1961-1972, separate archery
license regulations in effect.
~/Inconc1usive

kill data are omitted for years 1956 and 1957.

3/
- For years 1956-1967 no hunter pressure data available.

i/109 total resident and non~resident

archery licenses issued for unit.

i/517

total hunts for 1969 archery season in Unit 32; total does not reflect
number of hunters.

fl./114total resident and non-resident archery licenses issued for unit.
l/163 hunters based on total number of hunts; 126 total resident and non-resident
archery licenses issued for unit.
~/16l total resident and non-resident
Note:
ES
2ES
3ES
lDB
lDH
2Dr1
3DA

licenses issued for unit.

Hunt symbols are explained as follows:

=
=
=
=
=

=

Either sex, one deer (one license).
Either sex, two deer (one license).
Either sex, three deer (2 licenses plus 3rd deer coupon on 2nd license).
One deer, buck only (one license).
One deer multiple, either sex (2 Hcenses and 2 deer per individual).
Two deer multiple, either sex (2 licenses and 4 deer per individual).
Three deer, one must be antlerless (2 licenses plus'3rd deer coupon on
2nd license.

Note: The term lDr1 evolved into 2D, He or two dee r , hunter's choice (2 licenses
and 2 deer per individual) and is synonymous.
P. H. Neil
October 1973

�-159ELK HARVEST, SEASONS AND HUNTING PRESSURE
WILDLIFE MANAGEMENT mUT 32
(From Colorado Big Game Harvest 1956-1972)

YEAR

HUNTING
PRESSURE l/BULLS

COIolS

CALVES

TOTAL

HUNT AND SEASON

1956

94

21

12

2

35

AO + 100 ESP 'lj

10/15 - 10/31

1957

110

6

16

0

22

AO + 100 ESP 'l:./

10/15

10/31

CLOSED

1958
1959

49

·14

0

0

14

AO

10/17 - 11/3

1960

44

12

0

0

12

AO

10/17 - 11/6

1961

32

17

0

0

17

AO

10/21 - 11/8

1962

CLOSED

1963

CLOSED

1964

CLOSED

1965

45

12

4

0

16

25 B + 25 C ~/

10/16 - 11/5

1966

32

'10

9

4

23

25 B + 25 C 2/

10/15 - 11/3

1967

36

14

13

'l

30

30 B + 30

C !!./

.J

10/21 - 11/9

1968

78

15

13

2

30

30 B + 30 C !!./

10/19 - 11/7

1969

24

11

3

0

14

30 B + 30 C !i./

10/18 - 11/6

1970

94

40

2.

0

42

AO + 30 C }J

10/17 - 11/6

1971

172

61

5

0

66

AO + 20 C ~j

10/16 - 10/25

1972

169

31

8

1

40

AO + 30 C E./

10/28 - 11/12

1/

- Based upon combined resident and non-resident license sales and expressed in
number of licenses. During the period 1956-1970, when elk season coincided with
deer season, the hunting pressure statistics were very probably conservative.
Despite difficulties verifying that assumption, unkno~~ substantial numbers of
deer hunters may also have carried elk licenses hoping for a bull elk under
antlered only regulations. This latter happens regardless of low elk populations.
Thus, because of low success and tendency (above non reporting statewide figures
used in projections) for non-successful elk license holders not to report by card,
elk present in Unit 32 probably received extra pressure over that indicated by
card return projection years of antlered only seasons. Discrepancies between
(footnotes continued on next page)

�-160-

ELK HARVEST, SEASONS A.fID HUNTING PRESSURE

(Continued).

!/ continued
hunting pressure and total allowable specified permits in 1968 are probably
due to projection error tendencies that are inherent for all units having
small numbers of permits.
2/ .
- Un~ts 11, 22, and 32 combined.
1/Units 21, 22, 30, 31 and 32 combined.
~/Units 21, 22, 31, and 32 combined.
~/Units 22, 31, and 32 combined.
~/Units 22, 31, 32, and part of 21 combined.
Note:

Hunt symbols are explained as follows:

AO
. = Antlered only.
AO + ESP = Antlered only, plus either sex permits.
B+C
= Bull (antlered) and COy7 (antlerless) permits only (specified).
AO + C
= Antlered only plus specified number of cow permits.

P. H. Neil
October 1973

�-161-

ELK HARVEST, SEASONS AND HUNTING PRESSURE
WILDLIFE HANAGEMENT UNIT 32
(From Colorado Big Game Harvest 1956-1972)

ARCHERY

YEAR

lillNTING
PRESSURE );/ BULLS

COWS

CALVES

TOTAL

HUNT AND SEASON

1956

----

NO DATA

-

AO + 100 ESP JJ

1957

----

NO DATA

-

AO + 100 ESP '1:../
10/15 - 10/31

10/15 - 10/31

CLOSED

1958

----------

1959
1960
1961

NO DATA

- .- -

AO

10/17 - 11/3

NO DATA

-

AO

10/17 - 11/6

NO DATA

-

AO

10/21 - 11/8

1962

CLOSED

1963

CLOSED

1964

CLOSED

-------

NO DATA

-

NO DATA

-

1967

----

NO DATA

-

1968

3

0

0

25 B + 25 C 1/

10/16 - 11/5

25 B + 25 C 1/

10/15 - 11/3

ES
30 B + 30 C i/

9/2 - 9/17
10/21 - 11/9

30 B + 30 C 4/

10/19 - 11/7

30 B + 30 C if

10/18 - 11/6

1970

AO + 30 C 2./

10/17 - 11/6

1971

AO + 20 C 2./

10/16 - 10/25

AO
AO + 30 C 6/

8/19 - 8/31
10/28 - 11/12

1965
1966

----

1969

7

1972

NO DATA

0

- 0

0

- - -

0

0

0

!/Based upon combined total resident and non-resident
in number of licenses.
'1:../Units
11, 22, and 32 combined.
l/Units

21, 22, 30, 31, and 32 combined.

i/Units

21, 22, 31, and 32 combined.

(footnotes

continued

on next page)

license

sales and expressed

�-162-

ARCHERY ~LK tU"RVEST,

SEASONS AND HUNTING

PRESSURE

(Continued).

5/
- Units 22, 31, and 32 combined.
6/
- Units 22, 31, 32, and part of 21 combined.

Note:

Hunt symbols are explained as follows:

AO

AO + ESP
ES
B + C
AO + C

= Antlered only.
= Antlered only plus either sex permit.
One elk, either sex.
= Bull (antlered) plus cow (ant1erless) permits only (specified).
= Antlered only plus specified number of cow permits.

P. H. Neil
October 1973

�-163BLACK BEAR HARVEST AND SEASONS

(From Colorado

- WILDL IrE

Big Game Harvest

!/

HANAGEl'lENT UNIT

32

1955-1972)

2:./, }j

YEAR

HARVEST

1955

3R

8/15 - 10/1 and the regular

1956

8R

5/15 - 8/15 and the regular deer and/or elk season.

1957

3R

4/1 - 11/15 and any pre, regular,
elk season.

1958

7R + 5S

4/1 - 11/15 and the regular and post deer seasons.

1959

o

1960

3R

4/1 - 9/15 and the regular deer and/or elk season.

1961

3R

4/1 - 9/15 and the regular deer and/or

1962

3R

4/1 - 9/15 and the regular and post deer seasons.

1963

4S

4/1 - 9/15 an'a the regular deer season ending 11/7/63.

1964

2S

4/1 - 9/15 and the regular deer season ending 11/5/64.

1965

o

1966

o

1967

15

1968

o

1969

o

1970

o

1971

o

1972

2

HUNT Al\TJ) SEASON

deer and/or elk season.

and post deer and/or

No data on special season; any regular and post deer
and/or elk season.

elk season.

4/1 - 9/30 and the regular deer and/or
ending 11/5/65.

elk seasons

4/1 - 9/30 and the regular deer and/or
ending 1113/66.

elk seasons

Special season probably 4/1 - 9/30 and the regular deer
and/or elk seasons ending 11/9/67.
4/1 - 9/30 and the regular deer and/or
ending 11/6/68.

elk seasons

4/1 - 6/30 and the regular deer and/or elk seasons
ending 11/6/69.
4/1 - 6/30 and the regular deer and/or
ending 11/6/70.

elk seasons

4/1 - 6/30 and the regular
ending 11/11/71.

deer and/or elk seasons

4/1 - 6/30 and the regular
ending 11/12/72.

deer and/or elk seasons

1/
- R=Regular big game (Deer and/or Elk) licenses; S=Bear license for Special
Spring, Sununer, or Spring-Summer seasons; Also see annual regulations for
archery hunting.
(footnotes

continued

on next page).

�-164-

BLACK BEAR HARVEST AND SEASONS - WILDLIFE MANAGEMENT UNIT 32 (Continued).

21
- From 1955-1959, 1 bear per special bear license and/or 1 bear per bear
coupon on either or both deer and elk license; from 1960-1965, same as
1955-1959, except special bear license variously invalid after September
15 or 30, when bear coupon on either or both deer and elk license covered
bag and possession limits; 1966, same as 1960-1965, except one bear,
hunter's choice, per person per calendar year; from 1967 to present, bear
coupon removed from regular deer and elk licenses; thus, 1 bear, hunter's
choice, per special bear or sportsman's license per person per calendar
year only during special bear and regular deer and/or elk seasons.
llDogs permitted except when bear seasons concurrent with deer and lor elk
seasons.

P. H. Neil
October 1973

�-165MOUNTAIN LION HARVEST AND SEASONS - HILDLIFE HANAGEHENT UNIT 32
YEAR !/

HARVEST

1965

1 F '1:./

1966

5 M·, 2F

HUNT AND SEASON
10/16/65 - 3/31/66 West Slope; 10/23/65 3/31/66 East Slope, 1 either sex.
'1:../

1/1/66 - 12/31/66 in all counties we s t of
the Continental Divide and in Jackson, Conejos,
Alamosa, Mineral, Saguache, Rio Grande, Costilla,
Archuleta, Hinsdale and San Juan counties, 3
either sex.
10/22/66 - 2/28/67 in all counties east of the
Continental Divide except those specifically
listed above; 3 either sex.

1967

No Data

9/1/67 - 3/31/68 statewide; 9/1/67 - 5/31/68
west of State Hwy. #13 and north of U. S. Hwy.
116; 1 either sex.

1968

o

9/1/68 - 3/31/69 statewide; 1 either sex.

1969

o

9/1/69 - 3/31/70 statewide; 1 either sex.

1970

o

9/1/70
12/31/70 designated areas, and regular
deer and elk seasons statewide; 1 either sex.

1971

o

3/1/71 - 4/30/71, 1 male only, designated areas;
9/1/71 - 10/12/71 and 11/12/71 - 12/31/71, 1
either sex, designated areas.

1972

1

2/1/72 - 4/30/72, 1 male only, designated areas;
regular deer and elk seasons, statewide, 1
either sex; 11/13/72 - 12/31/72, 1 either sex,
designated areas including Unit 32.

o

!l Bounty of $50 per lion paid within period May 7~ 1929 - Harch 12, 1965;
protected by statute as of July 1, 1965 when mountain lion license was
established.
to "1965-1966" and "1966-1967" for Garfield County as reported by
'1:../Credited
Dixon, K. R. 1967. Evaluation of effects of mountain lion predation, p. 147;
In Game Res. Rep., July - Part II. Colo. Div. Game, Fish and Parks. pp. 73-310.
No data available in reports from section of Game }1anagement for 1965-1967.
Dixonts information is probably conservative in that the damage law allows
for lax reporting of ki11sof marauders by stockmen. For years 1968-1972 data
are from Colorado Big-Game Harvest Surveys.

P. H. Neil
October 1973

�SMALL GAME HARVEST AND HUNTING PRESSURE
SMALL GAl'1E HANAGEMENT UNIT 8, 1968-1972 1/

SPECIES

NUMBER OF HUNTERS
1970
1971 1972

1970

HARVEST
1971

1972

Average

24,813 11,606 9,276

14,148

--

14,961

1968

1969

1,984

1,328

Snowshoe hare

101

30

73

95

--

75

134

78

Blue grouse

298

154

213

124

442

246

1,172

73

368

78

815

501

Sage grouse

190

301

258

267

537

311

603

767

724

597

1,134

765

Chukar

122

453

151

223

--

237

76

606

247

358

--

322

Gambel 's quail

27

--

61

--

--

--

31

--

153

Pheasant

313

621

129

250

256

314

586

1,040

127

246

446

489

Cottontail

Band-tailed

rabbit

pigeon

Mourning dove

---

,1,319 1,411

--1/

Average 2/ 1968
1,510

--

--

--

--

--

2"76

290

242

447

314

---

1969

o !!./
1,403!:'/--

--

--

--

--

5,828 2,412

2,091

4,373

3,676

--

1/
- See Chap. 3 - Colo. Game,; Fish and Parks Division Laws and Regulations Handbook; Wildlife Management Unit
32 is wholly contained within Small Game Management Unit 8 and approximates six percent (6%) of Unit 8's
total area (300 sq. mi. ~ 4,970 sq. mi.) as p1animetered from Colorado topographic maps 1:500,000 scale.
Data from Colorado Small Game Harvest Surveys 1968-1972.
2/
- Hean for years data is shown.
l/No data.
!/Possib1e

sa~ple size error.
W. T. McKean
October 1973

I

I-'
0'\
0'\
I

�-167DUCK AND GOOSE HARVEST AND HtTNTI~G PRESSURE GARFIELD COUNTY, 1954 - 1972

1./

(From Colorado I-Jaterfo"loll
Kill Surveys)
GEESE '!:.../
EST. NO. HmiTERS

EST. HARVEST

No Data

19

35

34

60

57

1,076

32

No Data

1,945

16

No Data

2,795

25

No Data

538

4,923

4
75 Permits

1
(2)

302

1,219

250 Permits

(1)

YEAR

DUCKS
EST. NO. HL"1HERS

EST. HlI.EVEST

1954

No Data

2,284

1955

No Data

4,231

1956

No Data

1,764

1957

No Data

1,909

1958

372

1,442

1959

541

1,721

1960

112

1,194

1961

291

674

1962

260

414

1963

277

1,504

1964

291

1,404

1965

362

1966

50

1967

184

406

1968

265

1,348

1969

312

2,901

1970

403

1971
1972

1!

II

- The closest comparable area to Wildlife Management Unit 32 for which data are
available is Garfield County •. Excluding. National Forestlands,
Garfield County
contains 2,194 square miles. Unit 32 contains 300.7 square miles or 13.7 percent of the area concerned. For purposes of making comparisons these figures
could readily be converted to birds harvested per mile of riverbottom.
'!:.../Accuracy of data questionable due to sampling difficulties associated with
very small populations, geese and goose hunters alike.
110bvious inaccuracy.
,~. T. McKean
October 1973

�GAME SPECIES INTRODUCTIONS

- WILDLIFE MANAGEMENT UNIT 32 .
Release Area

Species

Date

Males

Females

Young

Total

Source of Stock

References

Chukar

9/5/41

?

?

?

50

Upper Mahaffey Ranch, Grand
Valley, Colorado.

Game Farm

FA Report 4-42

6/28/55

55

170

0

225

Parachute Creek near mouth of
Garden Gulch on Floyd Spangler
Ranch.

Game Farm

Sandfort's Files,
(Ft. Collins
Research Center)

3/22/57

50

50

0

100

Parachute Creek, 8.4 miles above
jct. of U.S. 6 &amp; 24, in Grand
Valley on slope west of Benson's
house.

Little Hills

Sandfort's Files,
(Ft. Collins
Research Center)

4/6/64

75

75

0

150

Mahaffey Ranch, 12.4 miles west
of Rifle.

Little Hills

Sandfort's Files,
(Ft. Collins
Research Center)

------------------------~--------------------------------------------------------------------------------------------8/27/41

Ring-

?

?

?

50

necked

Hahnewald's Farm, Grand
Valley, Colorado.

Otto

Game Farm

FA Repor t 4-42

....
~
I

00

pheasant

I

9/5/41

?

?

?

50

Mahaffey Ranch, Grand Valley,
Colorado.

Game Farm

FA Report 4-42

Literature Cited
Carhart. A. H.

1942.

Chukar-pheasant

experimental p Lant.Lngs ,

Jones, H., andG. Rogers. 1964. Chukar planting record.
files at Fort Collins Research Center.

Memo. to C,ommission. Fed. Aid Proj. l-l-17-R-3. 5 pp.

April.

Sandfort, W. W. 1956. Chukar partridge release records-Northwest
1956. l1emo. to Commission.
Fed. Aid Proj. W-37-R. 2 pp.

Original release record located in Sandfort's
Region and adjoining areas, Dec. 9-195l-Jan. 9,

1957. Trapping and transplanting chukar partridges, pp. 31-38.
Game and Fish. 119 pp.

In Quart. Prog.Rep.

July, Colo. Dept.
P. H. Neil
September 1973

�-169CENSUS AREAS AND ROUTES - WILDLIFE MANAGfl1ENT UNIT 32

MULE DEER
A spring trend count by fixed-wing airplane has been made from Rifle to
Grand Valley in Unit 32 annually since 1967 at least.
used in 1971, 1972 and 1973.

Helicopters

were

Procedural details are available in the

Northwest Regional office; summaries of results may be found there and
in the Denver office of the Game Manager.
necessarily

These summaries do not

show animal numbers by game management

unit, but more

commonly by drainage.

ELK
A winter aerial trend count for elk has been conducted ·by Northwest
Regional personnel in adjoining portions of Unit 22.

The trend area

also includes upper elevations of the Roan Plateau in Unit 32 from
headwaters of 'vest Parachute Creek to Davis Gulch, appx·oximate1y.

No

information has been supplied on dates and count results, but it is
assumed that those data are on file in the Northwest Regional office.
BLACK BEAR
Incidental to deer and elk trend counts black bear are tallied in Units
22, 32, 31 and others.

In addition, an annual estimate, by WCO district,

of bear populations was begun in 1972.

This estimate included also kill

by trappers and others, and management

objectives

for the species.

CHUKARS
A production count along Parachute Creek was conducted annually from 19571967. and from 1968-1972, with 1970 and 1971 missing

(Sandfort 1950-~952).

�-170-

Details of technique, map of area counted and results may be found in
the Northwest Regional office, in the Denver office, and in Sandfort's
files on chukars at the Fort Collins Research Center.

WATERFOWL
Wintering populations of ducks and geese in Unit 32 have been censused
annually from fixed-wing aircraft at least in 1971, 1972 and 1973,
possibly earlier, by Regional Hanagement personnel.

Areas covered were

on the Colorado River from Silt to Palisade.

Summarizing tables do not

break down specific river section populations.

Field notes of observers

may do so --:-see Northwest Regional personnel.

Literature Cited
Sandfort, W. \-1.

1950 through 1952.

Game bird surveys.

Rep., Colo. Dept. Game, Fish and Parks.

In Fed. Aid Quart. .

Pages variable by year.

(processed).

William T. McKean
November 1973

�HABITAT RESTORATION PROJECTS - WILDLIFE MANAGEMENT UNIT 32 1/

VEGETATIVE
TYPE

LOCATION

DATE

TREATHENT

NO. ACRES
TREATED

PURPOSE OF
TREATMENT

AGENCY

4/10 - 12/44

E. Parachute Cr.
T5S R94W Sec. 35

Sagebrush

Seeded with Agropyron
cristatum, Helilotus
officinalis, Poa
pratensis.

20

Revegetation

BLM

8/1/58 - 6/30/59

Middle Parachute Cr.
T5S R94H Sec. 7,18
T5S R95W Sec.12,13

Burned area
Sagebrush

Burn and reseed with
Agropyron cristatum,
Melilotus officinalis.

155

Res eed burned
area

BLM

6/18 - 22/62

E. Parachute Cr.
T6S R95W Sec.4-6

Sagebrush,
Oak

Aerial spray with 2
lbs. 2,4-D in 2.5
gal. of diesel.

800

Brush control

BLH

"
I

10/1 - 29/64

E. Parachute Cr.
T6S R94W Sec. 6,7

Sagebrush

Brush cutting with
a brush beater.

80

Brush control

BU1

1967

Parachute Creek
T5S R94H Sec. 2.3,

Sagebrush

Brush cutting with
rotary brush cutter

813

Soil and watershed modification

BLM

11,8,9,10,19,30

T5S
24

R95W

I
•...•
•...•

Sec.14,23,

1/
If details are desired on any of these projects, make requests directly to Mr. Ron Kufeld~ Colorado Division of
Wildlife Research Center, Fort Collins, Colorado 80521.

P. H. Neil and W. T. McKean
January 1974

�-172-

MANAGEMENT PROBLEMS CHECKLIST - WILDLIFE HANAGEMENT UNIT 32

A.

Present.
1.

Deer-auto collisions, State Highway 13 and U. S. Highway 6 and 24.
The location of these Highways are such that they intercept travel
routes and wintering areas of large numbers of migratory deer.
This situation creates a continuing potential for high incidence
of collisions between motor vehicles and animals.

The results of the initial phases of studies on deer-highway
crossing safety measures revealed that a 26 mile section of Highway
6 and 24 between Rifle and Grand Valley had the highest incidence
of deer road-kills, based on information collected at the time
from WCOI s throughout the state (Gordon 1969).

This particular

section of Highway 6 and 24 was selected for installation of van
de Ree mirrors as an experimental safety measure to reduce the
number of road-kills, however, Gordon (1969) concluded that the
mirrors did not significantly reduce deer-highway mortality in
this area.

This study did reveal that the great majority of road-

killed deer in this vicinity were killed in the months November to
March when the animals were concentrated on their winter range.

No information is available regarding road-kills on State Highway
13 immediately north of Rifle, however, it is believed that the
problem of deer losses here and on Highway 6 and 24 will be of
greater magnitude with the proposed oil shale development in
Parachute Creek and on those BLM lease sites located in the
Piceance Creek drainage to the north of Unit 32.

�-173-

2.

Deer damage to farm crops.
In general, the problem of deer damage to farm crops in Unit 32
is relatively small in comparison to some other units.

According

to Myers (1973 pers.connn.), claims for 1971-72 totaled $1,770.70
for damage to alfalfa and hay in Units. 31 and 32 combined.
for 1972-73 have only totaled $130.00 so far.

Claims

Decreased production

of alfalfa and spring grasses is alleged because of early grazing
by deer .

Some damage to stacked alfalfa has b een fairly common in

winter, as well as to the few scattered orchards in this area.
B.

Future.
1.

Oil shale development.
The proposed future oil shale development in Pa rachut e- Creek and
perhaps the Anvil Points area will undoubtedly create a multitude
of management problems, not only for all species of wildl ife but
all componen t s of the environment as well.

The increased human

and vehicular activity, to include residential and commercial
developments, associated with oil shale development will play a
major role in reducing the carrying capacity of the existing mule
d~erwinter

range, particularly in the Parachute Creek drainage

area, and result in loss of habitat for many small game and nongame species of mammals and birds.

A more detailed analysis of proposed oil shale development and
possible effects on the environment can be found by consulting
Thorne Ecological Institute (1973), Tweedy (1971) and U. S. Dept.
of Interior (1973), which are cited in greater detail in the Reference section at the end of this discussion.

�-174-

2.

Extension of Interstate 70 through Rifle and Grand Valley.
Interstate 70 will intercept mule deer winter range and undoubtedly
interrupt several migration routes, resulting in higher winter
losses and still higher numbers of deer-auto accidents.

A more

detailed analysis of this particular problem can be found by consuIting Colo. Div. Wild!. (1973) regarding the environmental
analysis and probable effects on all species of wildlife of the
proposed extension of Interstate 70 from Rifle to Plateau Creek.

Literature Cited
.

.

Colorado Division of Wildlife.

1973.

Fish and wildlife analysis of the

Int;erstate 70 Highway corridor, Rifle to Plateau Creek.
July.

126 pp ,

Gordon, D. F.
pp. 5-23.

1969.

Evaluation of deer-highway crossing safety measures.

Final Report.

In Quart. Res. Rep, , July-Part 1.

Div. Game, Fish and Parks.

Thorne Ecological Institute.

Tweedy, J. B., et al.

1971.

Colo.

140 pp.

1973.

The Colony environmental study, Para-

chute Creek, Garfield County, Colorado.

t Lon

Draft report.

August.

Boulder.

3 vol.

Report on economics of environmental protec-

for a Federal oil shale leasing program.

A Special Committee of

the Governor's Oil Shale Advisory Committee (for Colorado).

January.

204 pp.

U. S. Dept. of Interior.

1973.

Final environmental statement for the pro-

.totype oil shale leasing program.

Washington, D. C.

6 vol.

P. H. Neil and W. T. McKean
November, 1973

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                  <text>-175-

July, 1974

JOR PROGRESS REPORT

State

of ______

(:(_)L_O!~~DO.

_

Deer..:..Elk
Investigations
Proj ect No. __ . W-38-R.:.:.-.-.:::,2.:::,8
_
llC

Wor.k Plan No.

3

Job Title

Evaluation of Experimental Elk Management Procedures

Period

April 1, 1973 through March 31, 1974

Personnel:

Covered:

Raymond J. Boyd and James F. Lipscomb.

ABSTR\CT
No work was accomplished on this job the past segment as all objectives and
procedures were incorporated into a new job(WP 18, J 1, this project) with
four rese&lt;:trchersacting as a Task Force to examine the entire deer and elk
management system in Colorado. This examination of the entire state was to
generally follow the objectives and procedures of this job, but on a statewide scope and both for deer and elk •

/")

.

~/J~~Q~lL.IrOiiII://(~
': £~.~.

Prepared by ~l-l....::~~·
...•-+=..//-=.I =-'
'Ra~#~
Asst. Wildlife Researcher

��-177-

July, 1974

JOB FINAL REPORT

State of
Project

,.-:.C:.,:O:..:;L:,:O:.:.RAD=.:=..O
_
No.

Work Plan No.

W-38-R-28

Deer-Elk

Investigations

14

Job No.

1

Job Tit1e:

Mi
__d_d_1_e_P_a_rk_D_e_e_r_S_t_u_d:.:.y_-~P-o~p-ul_a_t_i_o_n
__D_,i_s_t_r_i_b_u_t_i_on

Period

April 1, 1973 through March

Covered:

Personnel:

_

31, 1974.

W. M. Brandes, G. L. Brown, L.R. Carpenter, R. D. Clippinger,
M. C. Coghill, J. Cooney, J. J. Dorrance, P. F. Gilbert, R. B.
Gill, D. Roart, S. Rom, J. J. Klein, Jr., D. Luce, P; K. Mason,
D. Merriam, K. Moser, L. A.Roper,
R. L. Schmidt, B. C. Sigler,
W. Travnicek, M. Ward, J. B. Weir, J. L. Wolfe, and W. B. Woodward.
ABSTRACT

A draft of a manuscript entitled ''Mule Deer Movem.ents and Distribution
in
Middle Park Colorado" has been completed and is now in the editing stages.
ManUscript will be submitted to an appropriate journal for publication.

PreparedbYJ~
(/ ~
, , Laren A. Roper
Asst. Wildlife Researcher

��-179July, 1974
JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~-----------

Project No.

W-38-R-28

Work Plan No.

14

Deer-Elk Investigations
Job No.

~4~

Job Title Middle Park Deer Study - Physical Characteristics and Food Habits
Period Covered:
Personnel:

April 1, 1973ihroughMarch

31, 1974

L. A. Roper

ABSTRACT
No work was accomplished on the Physical Characteristics portion of this
job during the current segment. The Physical Characteristics portion has
been amended for one more year to allow time for reporting the data in a
Game Information Leaflet. Originally, these data were intended for publication in the Division of Wildlife _~E~cial Report Series. But subsequent
analyses of portions of the data indicated most of the data were too sparse
to permit extrapolation to Middle Park deer populations. Hence, only
pertinent descriptive physical measurements will be reported in the Game
Information Leaflet.
The food habits portion of Job 4 were presented in full in the 1972-73
Segment (W~38-R-27, WP 14, Job 4).

Prepared by

cGvaeC.i:r&amp;!
.Laren A. Roper.
.
Asst. Wildlife Researcher

_

��-181July, 1974
JOB FINAL REPORT

State of

C;:.O;:,;L::.;O;:,;RA=D::.;:O:.._

Project No.

W-38-R-28

Work Plan No.

14

Job Title:

Job No.

5

Middle Park Deer Study - Experimental Range Fertilization

Period Covered:
Personnel:

Deer-Elk Investigations

Len H

April 1, 1973 through March 31, 1974.
e .

Carpenter, R. Bruce Gill, Donald Reichert, and 0. C. Wallmo.

ABSTRACT
A dissertation (Carpenter 1974) is being prepared based on info~tion
and
date obtained from this study and is in the final editing stages. This
thesis, copies of which will be on file at the Colorado Division of Wildlife
Research Center Library and at Colorado State University Library, becomes the
.final report for this job.

Prepared by --=1:,;.:::. ...:.4':';1.-:::...-_11_·, _C:..;:._4._'1r1-&lt;~_1 '_~_~_' '_,
Len H. Carpenter
Student Assistant IV

__

��-183July, 1974

JOB PROGRESS REPORT

State 0 f

COLORADO
--~--~~~~~---------

Project No.

W-38-R-28

Work Plan No.

14

Deer-Elk Investigations
Job No.

5_a
__~

Job Title

Middle Park Deer Study - Range Fertilization

Period Covered:

April 1, 1973 to April 1, 1974

Personnel:

R. Bruce Gill, O. C. Wallmo, Don W. Reichert, Wayne L. Regelin,
and Len H. Carpenter.

ABSTRACf
A study area on native sagebrush range in western Grand County, Colorado
was selected and fenced to investigat-e the response of mule deer forage
species to combined fertilizer-herbicide treatments applied at various
dates during the growing season. A randomized block design with six
herbicide application dates and six controls was replicated three times
Pelleted ammonium nitrate fertilizer (33.5% N) at the rate of 60 pounds
elemental N per acre was applied to each tr~atment plot at the start of
the experiment.· Beginning with April 4, and at two week intervals until
June 13, six, i~4-D herbicide applications at the rate of 2 pounds acid
equivalent per acre were applied to appropriate plots. Aerial cover
measurements of selected plant species were made. on each treatment plot
during August by the rated microplot technique.· Covariance and graphic
analyses of the data were made. April 17 appeared to be the most
promising herbicide application date for maximizing beneficial responses
of important mule deer food species.

_

��-185-

MIDDLE PARK DEER STUDY
EXPERIMENTAL RANGE FERTILIZATION
Len H. Carpenter

This report presents the design, methods, and treatment data gathered
for the first year of this study.

P. S. OBJECTIVE
To test the effects of fertilizer on deer forage and deer feeding responses
to fertilization of critical winter range areas.
SEGMENT OBJECTIVE
Investigate the response of deer forage species to combined fertilizer
herbicide treatments applied at different periods during the growing season.
METHODS AND MATERIALS
A study area on native sagebrush range near Kremmling, Colorado was chosen
for this study. A study site was chosen where there were sufficient forbs
to permit measurement of changes in density attributable to 2,4-D treatments. The experimental design chosen was a randomized block with six
treatments (herbicide application dates) where each treatment plot was
split and one-half was treated while the other half was untreated and
served as a control. The location of the control plots (either east, west,
north or south of the treatment plots) was randomly chosen. This design
resulted in 12 plots in each replicate. This layout was re-randomized and
replicated three times at the study site (Fig. 1). Each treatment unit was
50 X 100 feet with each split-plot unit being 50 X 50 feet (Fig. 2). This
resulted in a block of land 300 X 300 feet wi th each replicate being 100
X 300 feet.
All split-plot units that were treated with herbicide also received an
application of pelleted ammonium nitrate fertilizer (33.5% N) at the rate
of 60 pounds elemental nitrogen per acre. The fertilizer was applied on
April 2 and 3 by a "whirlybird" hand fertilizer. Herbicide was applied
on April 2, April 17, May 2, May 16, May 31, and June 13; that is,
approximately 15 days apart. At each treatment date, 2,4-D herbicide was
applied at a rate of 2 pounds acid equivalent per acre with a back-pack
sprayer modified with a 5-foot boom containing four nozzles. The 2,4-D
consisted of 72.8% propylene glycol butyl ether esters. The herbicide
was applied with water as a carrier at a rate of 5 gallons per acre. A
small amount of wetting agent was also used to increase effectiveness of
the spray. The entire study site was enclosed with a three-strand barbed
wire fence to prevent livestock use.

�I

I

June

I

I

13

Control

Control

I

4

17

I
I

I

I

I

I
I
I

May

April

16

4

I
I
I
I

May

r

Control

31

I
I

April
Control

17

I

I

I

I

I
I
I

Control

I

I
I

lY

May
Control

Control

16

:I
I
I

Control

I

I

I

Control

I
I
I
I

Rep 2

I

I

I
I
I

r

I
I

Control

I

I

I

13

2

I
I

I

I,

June

April

I
I

I
I
I

April

I

May

I

I

Control

:I

Rep 1
May

May

I
I

MIl,.

Control

1-------

16

31

2

April
Control

Fig. 1.

May

--.-. --- -------

------

April
4

May
Control

13

Control

Control

I

I

2

I

June

,...
00
0-

Control

31

~E

Control

17

Physical layout and randomized location of the six herbicide
treatment dates and controls for the three replications.
Solid lines represent treatment boundaries--dotted lines
represent split-plot boundaries.

Rep 3

�I

T

,
I
I
I
I
I
I

100'

I
I
I
I

1
~

....,
I

I
I

i

:

SO'~

r--Fig. 2.

I

~
0;

300'

Individual replicate and treatment dimensions.

-J

�-188-

There were approximately 12.5 inches of crusted snow covering 75 percent
of the ground on the treatment plots at the time of fertilizer application.
The ground remained frozen during fertilizer applications, and limited
snow melt occurred during the warm period of the day. Weather, moisture
and plant phenology at each herbicide application date are presented
(Table 1).
Nitrogen fertilizer was used along with herbicide in order to obtain information relatable to other studies being conducted on similar vegetation in the immediate area (Carpenter 1970, 1971, 1972). In this work
nitrogen and 2,4-D resulted in a reduction in sagebrush (Artemisia
tridentata tridentata), an increase in grasses, and a general increase
in total plant biomass, but also a general decrease in forbs. The herbicide was applied early in June at the optimum state of susceptibility of
big sagebrush. It was hypothesized that earlier treatments would cause
some reduction of the sagebrush plants but would have minimal effects on
the associated forbs.

Vegetation Measurements
The rated microplot method as described by Morris (1971) was employed to
measure treatment effects, using a 1 X 2-inch rectangular plot. All
measurements were made in August approximately two months after final
herbicide treatment date. Aerial cover was recorded for selected plant
species of special interest and for plant groups, in the categories
"other grasses", "other forbs", and "other shrubs", for species of very
low frequency or of little importance as deer forage. Cover was rated
to the nearest one-tenth (1/10 = score of 1, 10/10 = score of 10) of the
area occupied in the microplot. Two hundred microplots were systematically
located in each of 36-50 X 50 feet treatment plots and controls. Cover
ratings were then added together for the 200 separate microplots for each
plant species or group to obtain a measurement value for that treatment
or control plot. While this total rating may not directly relate to cover,
the rated plot method will give a precise estimate of plant cover (Morris
1971) .
Cover information was obtained separately on both dead and live cover for
big sagebrush, green rabbitbrush (Chrysothamnus.viscidiflorus), snowberry
(Symphoricarpos oreophilus), Vasey rabbitbrush (Chrysothamnus vaseyi),
other shrubs and total shrubs. Forbs recorded by species included buckwheat (Eriogonum umbellatum), bluebell (Mertensia lanceolata), penstemons
(Penstemon caespitosus and !.. watsoni), and phlox (Phlox multiflora andP.
bryoides). Grass recorded by species included western wheat grass (Agropyron
smithii), bluebunch wheatgrass (A. spicatum)-, muttongrass (Poa fendleriana),
bottle brush squirreltail grass (Sitanion hystrix), and pine needlegrass
(Stipa pinetorum).
The data obtained for big sagebrush (live cover only), buckwheat, both penstemon species, western wheatgrass, muttongrass, pine needlegrass, and total
shrubs, forbs and grasses were analyzed by covariance analysis. Lack of
uniform data for each treatment date prevented analysis for other species.

�-189-

Table 1. Weather, moisture and plant phenology at each herbicide
application date.

Date

Environmental
Conditions

Selected
Plant Phenology

April 4

Air temperature 390 F. Wind
less than 5 mph. Average
snow depth 12.5 inches. 8085 percent snow ground cover.

Artemisia tridentata stem
elongation beginning.

April 17

Air temperature 450 F. Wind
5-7 mph. Snow in patches
4-6 inches deep. Soil
very wet and muddy.

Chrysothamnus viscidiflorus
beginning to leaf bud.

May 2

Air temperature 450 F. Wind
5-20 mph. Soil moisture
good.

Phlox bryoides beginning
bloom. Penstemon
caespitosus showing vegetative growth. Stipa
pinetorum starting green
growth.

May 16

Air temperature 600 F. Wind
5-15 mph. Top 2 inches soil
dry.

Phlox bryoides ending bloom.
Penstemon caespitosus much
vegetative growth. Koeleria
cristata heads forming.
Poa fendleriana green growth
at base.

May 31

Air temperature 60 F. Wind
less than 5 mph. Soil
surface dry.

Mertensia lanceolata full
bloom. Potentilla concinna
beginning bloom. Eriogonum
umbellatum near bloom.
Koeleria cristata full head.
Poa fendleriana full vege~
tative growth.

June 13

Air temperature 700 F. Wind
less than 5 mph. Soil
surface dry.

Mertensia lanceolata postfloral. Phlox multiflora
blooming. Eriogonum
umbellatum blooming. Poa
fendleriana full head.

o

�-190:--

Each of the 18 control plots served as the independent value while each
of the 18 treatment plots were the dependent values. Covariance analysis
allowed for more precise measurement of true treatment effects by adjusting each treatment cover mean by its own control~ This procedure
decreased the effects of heterogeneous vegetation. Resulting adjusted
means were used for graphic analysis of vegetation response by treatment dates. For some species percent kill was obtained from the unadjusted live and .dead cover measurements.
.

DESCRIPTION OF AREA
The study area, on public land administered by the Bureau.of Land Management, U.S.D.!., is located 5 miles northeast of Kremmling, Grand County,
Colorado (T 2 N, R 80 W, SE 1/4 NW 1/4, Sec. 22).

RESULTS AND DISCUSSION
The percentage kill of total forbs and total shrubs (calculcated from
measurements of live and dead cover) increased from the April 4 herbicide
treatment to the May 31 treatment and decreased thereafter· (June 13 treat'ment) (Fig. 3). The same trend was true for big sagebrush except that
there was little difference between May 31 and June 13 treatments. The
percent kill estimates were not subjected to statistical analysis, because
the analysis was designed to interpret changes in aerial cover over years
and in most cases, mortality of plants can be recorded only as an immediate
effect.
Aerial cover of live forbs, measured in August, was greatest on plots
treated April 4 and least on plots treated May 31 (Fig. 4). Plots treated
June 13 had more forb cover in August than those treated May 2, May 16, or
May 31. Most of the increase was attributable to Pertstemoncaespitosus
(Fig. 5), though the same trend was apparent irtEriogortumumbellatum.
Apparently their peak of susceptibility to 2,4-D is around June 1. Peak
susceptibility of !. watsoni was around May 15.
Aerial cover of shrubs showed the same trend, with peak susceptibility
around May 31. While there was little difference in cover reduction of
big sagebrush on plots treated May 31 and June 13 (Fig. 3), there was
slightly more live sagebrush cover on June 13 plots than.on May 31 plots
(Fig. 5). Even with this diminished effect on June 13, the date of herbicide application did have a significant (p = .05) negative linear effect
on aerial cover of big sagebrush, buckwheat, both penstemons, total shrubs,
and total forbs. Adjusted means for microplot cover by treatment date for
all the various plant variables are given (Table 2).
In the case of grasses there was not a clear pattern of response to date
of treatment (Fig. 4). However, grasses are not directly affected by
2,4-D. They would be expected to respond to reduced competition from other
plants and fertilization, and this response should be measurable in future
years.

�100
Artr

80
Total
shrubs

'""""'

~

w

~

U
H

o
,....l

60

p,..

o

Total
forbs

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.~
.....,
,....l
,....l
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....
I.

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p,..

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/

/

;',/
;'
;'
,/

;'

;'
;'

/

April
4

Fig. 3.

April

17

May
16
2
DATE OF TREATMENT
May

May
31

June
13

Percent kill of big sagebrush, total shrubs, and total forbs on plots treated
with 2,4-0 herbicide at six dates in spring.

�280

I

•,
\
\
\ ..A

240

\0

L

\'d
\ ~
\
\

\
\

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\

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L

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,
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Ul

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40

Fig.

4.

August cover

April

April

4

17

of f orb s , grasses,

May

May

June

2
16
DATE OF TREATMENT

31

13

May

and shrubs

on plots

treated

with

2,4-0

herbicide

at six

dates

in spring.

�200

Peca - Penstemoncaespitosus

Peca
160

Artr = Artemisia tridentata
Erum = Eriogonum umbe11atum

~

Pewa a Penstemon watsoni

§
H

en

...,
::;l

120

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~
~
~

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...... ..•....._

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....-

-......•
... --------

.

May
31

June
13

.....
.....
-....
..•.
_

April
4

April
17

May
2

I

May
16

_---

DATE OF TREATMENT
Fig. 5.

August cover of four plant species on plots treated with 2,4-D
herbicide at six dates in spring.

�-194-

Table 2. August cover of selected plant species and groups on plots
treated with 2,4-D herbicide at various dates in spring. Values are
means of micro plot -cover-estimates as adjusted by covariance-analysis.Dates
.Adjusted Means
Max
1
15

Species or
Plan t Groups

1

A:eril
15

Big sagebrush

108

40

21

Total shrubs

169

93

Buckwheat

29

Penstemon caespitosus

1

Jl.IDe
15

30

10

14

71

55

25

46

19

7

8

4

7

181

89

27

23

11

68

Penstemon watsoni

14

24

9

0

4

7

Total forbs

270

162

58

55

24

103

Western wheatgrass

13

11

16

13

14

14

Muttongrass

68

90

95

64

77

68

Pine needlegrass

11

9

18

11

13

11

Total grass

91

123

143

96

116

94

Figure 3 suggests two reasonable alternatives for timing the application
of 2,4-D if the desire is to achieve a reduction in sagebrush while preserving forbs. Application of2,4-D on April 4 achieved a 26 percent kill
of sagebrush and no forb mortality. The April 17 treatment killed 63 percent of the sagebrush and only 17 percent of the forbs. The snowcover on
April 4 and April 17 (Table 1) probably had some influence on the effectiveness
of 2,4-D on forbs. In other years forbs might be more or less advanced
phenologically and slightly different results would occur if herbicide
were applied on those dates. However, managers can use phenological development as a guide for spring applications just as they now do for summer
application.
Finally, it is emphasized that the results reported here are from the first
growing season after application. Ultimate interpretations will be contingent upon subsequent measurements.

�-195-

. LITERATURE CITED

Carpenter, L. H. 1970. Middle Park deer study - range fe~ti1ization.
Colo. Div. Game, Fish and p'arks. Game Res. Div., Fed. Aid Proj.
W-38-R-24. Game Res. Rep ,, July, Part 3. pp , 369-393.
Carpenter, L. H. 1971. Middle Park deer study - range fertilization.
Colo. Div. Game, Fish and Parks. Game Res. Div., Fed. Aid Proj.
W-38-R-25. Game Res. Rept., July, Part 3. pp. 225-253.
Carpenter, L. H. 1972. Middle Park deer study - range fertilization.
Colo. Div. Wildlife. Game Res. Div., Fed. Aid Proj. W-38-R-26.
Game Res. Rept., July, Part 2~ pp. 211-251.
Morris, M. J. 1971. Estimating understory plant cover with rated
microp1ots. Final Report FS-RM 1702. Rocky Mountain Forest and
Range Experiment Station, Fort Collins, Colo. 29 p.

��-197July, 1974,
JOB PROGRESS REPORT

State of

C::.;O:,:LO=M::.;ffi:::..;:.O-----

Project No.

W-38-R-28

Work P1ailNo.

14

Deer-Elk Investigations
Job No._~-8--_~----~--~------

Job Title

Middle Park Deer Study - Experimental Harvest Regulations

Period Covered:

April 1, 1973 rhrough Harch 31, 1974

Personnel:

W. J. Adrian, G. L.Brown, L. M. Brown, L. Brevard, R. D. Clippinger,
G. Demarest, F. G. Dienes, R. E. Keiss, J.F. Lipscomb, J. A. }1'.aas,
D. Mueller, D. Luce, J. M. Neisus, H. Riffel, L. A. Roper, G. G.
Schoonve1d, J. A. Tulloch, D. Walsworth, and J. L. Wolfe.

ABSTRACT
The January 1973 deer population estimate for deer in Midd1e.Park was
9,670 ± 2,566 deer. Winter losses were estimated at 6.4 percent or 647
deer. Productivity, based on pre-season deer classifications amounted to
86.9 ± 11.8 fawns per 100 adult females. There was no significant difference between October, December and January sex and age classifications
even though a harvest of 775 deer was estimated after the October count
,and a 966 harvest was estimated following the December count. The mean
estimate of bucks per 100 does for each count was 55.4, 58.8 and 62.7.
Fawns for each classification. were estimated at 86.9, 88;1 and 89.4 per
100 does. The total harvest estimate for all hunting seasons was 1,771 deer
or 41 percent below the harvest objective of 3,000 deer. Failure to
distribute all post season deer permits was primarily responsible for the
low harvest. Hunter success was an estimated 22.2 percent for the regular
deer season and 58.1 percent for the post season. AD. estimated 83 percent
of the'permitted hunters hunted in the regular deer season and 77 percent'
hunted .in the post deer hunt.
'

��-199-

MIDDLE PARK DEER STUDY - EXPERIMENTAL HARVEST REGULATIONS
Laren A. Roper

P. S. OBJECTIVE
To adjust a mule deer population to a specified density on the winter
range.
SEGMENT OBJECTIVES
1.

Estimate.deer density, structure, productivity and winter mortality
in Middle Park.

2.

Recommend season regulations and publicize the procedures for hunters
to obtain permits and participate in the experimental hunt.

3.

Measure and estimate the harvest by sex and age class and the success
of hunters.

4.

Determine if the hunting public wilL participate on a continuing basis
to achieve adequate harvests utilizing specified permits in post season
hunts.

5.

Determine if specified permits are capable of achieving predetermined
harvest goals.
METHODS AND MATERIALS

Estimate Deer Density, Structure, Productivity and Winter Mortality
Methods have been reported by Roper (1973, Gill (1969 and 1971).

Recommending and Publicizing Season Recommendations
Methods have been reported by Roper (1973).
Estimate the Harvest and Success of Hunters
Methods have been reported by Roper (1973).
1.

Exceptions are:

A 50 percent sample of permitted hunters was obtained for an estimate
of the regular season harvest and 33 percent of the post season hunters
were surveyed. The statewide game management survey was used for estimating the harvest for archers and high country buck hunters. A 50 percent random survey of all archers in the State was used and a 100 percent
survey for high country buck hunters was attempted.

�-200-

2.

The self-service drop boxes were eliminated as were the report cards
attached to each permit.

3.

The surveys were incorporated into the regular big game survey system
utilized by the Game Management Section. This system uses automatic
data processing where non-responding hunters receive follow-up letters
at two week intervals until the percent return approaches or exceeds
90 percent.

Determine if Hunters Will Participate
See Roper (1973).

Determine if Specified Permits are
Capable of Achieving Predetermined Harvest Goals
The deer population life equation was used to assess the success of the
hunting season as determined by the random surveys of 50 and 33 percent
of the permitted hunters in the October and December hunts to obtain harvest estimates.

RESULTS AND DISCUSSION
Density Estimates
The 1973-74 winter population estimate of deer for Middle Park was 9,670
± 2,566 (P&lt;.lO) deer or 16.5 ± 4.4 (P&lt;.10) deer per square mile of winter
range. Table 1 is a summary of deer counted per quadrat and Table 2 is a
list of population estimates by each deer herd sub-unit. Table 3 is the
life equation data for deer in Middle Park from 1967 to present and Table
4 is a more detailed life equation for deer from January 1973 through
January 1974.

Structure (Sex and Age Composition)
Pre-season Counts
On October 10, 11 and 12, 760 deer were classified by one observer using a
helicopter. A fresh snow of 3-4 inches throughout the Muddy Creek area provided excellent count conditions. The_Blue River area had considerable bare
ground and occasional winds resulted in fair count conditions. In the
Troublesome Creek and Williams Fork River areas the count was hampered by
fog and lightly falling snows. Conditions for counting were poor.
A total of 96 (12.6 percent) mature bucks, 78 (10.2 percent) young bucks,
314 (41.3 percent) does and 273 (35.9 percent) fawns were-classified.
Resulting sex and age ratios (Table 5) were: 55.4 ± 8.61 bUGks:100 does:86.94
± 11.84 fawns (P&lt;.10) (Riney 1956).

�Table 1.
1974.

Numbers of deer counted per quadrat within eight sampling strata, Middle Park, Colorado,

Muddi: Creek
Low
High
Density
Density

£=Total
y
yh

Blue River
Low
High
Density
Density

Troublesome Cr.
High
Density

Granbi:
Low
Density

0

0

14

0

12

3

15

0

0

4

0

4

0

0

41

0

0

0

36

3

60

0

0

20

0

0

65

90

17

0

54

0

0

0

128

87

15

0

30

0

0

0

0

84

0

0
13
25
125
52
54
66
32
0
0
0
0
0
3
0

0
0
0
1
7
0

12
48
50
79
65
2

0
91
0

14
41
1
16

0

370
18.5

12
1.0

499
41.6

275
30.6

296
29.6

20
2.9

= 16.4552
yst = 2.6316

N yst

Williams Fk. River
Low
High
Density
Density

C.L. = 16.46 ± 4.37
= 9670 estimated deer
C.L. = 9670 ± 2566

104
20.8

3
.6

I

N

....
0
I

�Table 2.

Deer population estimates by sub-units in Middle Park, 1968-1974.

Year

Muddy
Creek

Blue
River

Williams Fork
River

1968
1969
1970
1971
1972
1973
1974

4,101
2,994
1,429
794
1,002
2,856
2,272

4,074
3,531
3,291
3,335
2,259
4,186
4,663

843
774
271
569
742
556
622

Table 3.

Troublesome
Creek

Granby

Total

1,622
1,679
1,910
889
1,404
2,340
1,897

No Count
134
306
137
22
149
216

10,640
9,112
7,207
5,724
5,429
10,987
9,670

Wounding
Loss (20%
Harvest)

Post-hunt POE'
Ca1cu- Census
1ated
Est.

Lif.e equation for deer in Middle Park, winter 1967-68 through 1973-74.
I

Year

Jan. Pop.
Est.

1967-68
1968-69
1969-70
1970-71
1971·72
1972-73
1973-74

10,640
9,112
7,205
5,719
5,429
10,087
9,670

Winter Loss
No.
%

Other
Known
Losses

PreFawn
Pop.

Fawn
ReEroduction
%
No.

Pre
Hunt
Pop.

Harvest
No.
%

-

42.65
53.48
45.35
47.36
52.22
50.00

3,810
4,531
2,753
2,477
3,675
4,621

12,744
13,004
8,823
7,707
10,713
13,870

23
51
28
7
3
13

2,909
4,503
2,436
516
356
1,771

582
901
487
103
71
354

9,253
7,600
5,900
7,088
10,541
11,745

9,112
7,206
5,730
5,429
10,987
9,670

*

*

*

*

*

*

*

*

16.0
7.0
15.8
8.6
.7
6.4

1,706
639
1,135
489
50
647

101
192

8,934
8,473
6,070
5,230
7,038
9,249

*

*

*

*

*Data not available this segment.

N

0

N
I

�Table 4.

Life equation for deer in Middle Park from January 1973 - January 1974.

Bucks

Does

Buck
Fawns

1973 Winter Pop.

26.0%
2,625

40.7%
4,100

18.3%
1,849

15.0%
1,513

100.0%
10,087

Winter Mortality
(6.4%)

43.5%
281

8.7%
56

26.3%
170

21.5%
139

100.0%
646

Accidental
Deaths

26.1%
50

40.6%
78

18.2%
35

15.1%
29

100.0%
192

Spring Pop.

24.8%
2,294

42.9%
3,966

17.8%
1,644

14.5%
1,345

9.249

Female
Fawns

Total

Percent
Yearlings
Bucks
Does

I

N

Pre-fawning

Pop.

42.6%
3,938

w

57.4%
5,311

Fawn Crop
(.93 Fawns/Doe)
Pre-hunt Pop.

·0
I

39.9
55.0%
2,542

45.0%
2,079

4,621

28.4%
3,938

38.3%
5,311

18.3%
2,542

15.0%
2,079

100.0%
13,870

Harvest (Archery, Regular 59.6%
and Post-Season Totals)
1,056

34.4%
610

3.4%
61

2.6%
44

100.0%
1,771

Wounding Loss
(20% of Harvest)

211

122

12

9

354

Post-season
(Calculated)

22.8%
2,671

39.0%
4,579

21.0%
2,469

17.2%
2,026

100.0%
11,745

Post-season
(Actual Est.)

24.9%
2,408

39.7%
3,839

20.5%*
1,985

14.9%
1,438

100.0%
9,670

* Fawn harvest = 35.4% of total kill.
Buck fawns comprised 58.0% of total fawn harvest.

24.5

�-204-

Table 5.

Pre-season deer classification

Year

Bucks

Does

Fawns

Total

Bucks:
100 Does

Fawns:
100 Does

1969

195

451

419

1,065

43

93

1970

152

361

258

771

42

72

1971

20

41

30

91

49

71

1972

73

102

89

264

72

88

1973

174

314

273

761

55

87

Post-OctoberDeer

comparisons from 1969-1973.

Season Counts
,

Helicopter classifications by one observer on December 10, 11 and 12, 1973,
resulted in classifying 1,664 deer. Classifications were 244 (14.7 percent)
mature bucks, 152 (9.1 percent) young bucks, 674 (40.5 percent) does, and
594 (35.7 percent) fawns. Count conditions were .good ·to excellent and
ratios were: 58.7 ± 6.1 bucks:100 does:88.1 ± 8.2 fawns (P&lt;.10), (Table 6).

Table 6. Post-season classification of deer in Middle Park from 1968-1973
in November and December.

Year

Type of
Count

Bucks

Does

Fawns

Total

Bucks

100 Females

Fawns

1968

Ground

147

396

363

906

37

100

92

1969

Ground

319

582

505

1,406

55

100

87

1970

Ground

104

231

178

513

45

100

77

1971

Ground

356

783

321

1,460

45

100

41

1972

Ground

818

1,281

1,047

3,146

64

100

82

1973

Aerial

396

674

594

1,664

59

100

88

�-205-

Post-December Deer Season Counts
Following the controlled quota post deer season sex and age classifications
were made using a helicopter after the deer census. All classifications
were made on January 13, 1974. Conditions were excellent with snow background and little wind.
Of 1,266 deer classified, 160 (12.6 percent) were mature bucks, 155 (12.2
percent) young bucks, 502 (39.7 percent) does, and 449 (35.5 percent) fawns.
Ratios were: 62.7 ± 7.4 bucks:lOO does:89.4 ± 9.56 fawns (P&lt;.lO), (Table 7).

Table 7. January classifications of deer in Middle Park from 1969-1974.
The 1974 classification follows a post-hunting season.

Year

Type of
Count

Bucks

Does

Fawns

Total

Bucks

100 Females

Fawns

1969

Aerial

316

558

501

1,375

57

100

90

1970

Aerial

261

490

418

1,169

53

100

85

1971

Aerial

211

471

363

1,045

45

100

77

1972

Aerial

1973

Aerial

362

693

540

1,595

52

100

78

1974

Aerial

315

502

449

1,266

63

100

89

Comparisons Between October, December and January Classifications
Chi square tests (Snedecor and Cochran 1968) for bucks and fawns per lOO
2
does between the October and December counts were non-significant (X = .17,
.006 respectively). Similar non-significance occurred betwee~ October and
January classifications of bucks and fawns per 100 females (X = .69, .14
respectively) •
Table 8 is a summary of the range of variation as shown by the 90 percent
confidence limits for the buck and fawn ratios per 100 does.
Apparently deer behavior and distribution of the sexes are masking the effect
of hunting on the overall sex and age structure classifications of deer
throughout Middle Park.

�-206-

Table 8. Confidence limits (p &lt; .10) for buck and fawn ratios per 100
female mule deer in Middle Park observed during helicopter surveys in
October, December and January, 1973 and 1974.
Date

Bucks : 100 Females

Fawns

October 1973

46.8 ~ 55.4 ~ 64.0

75.1 S. 86.9 ~98.7

December 1973

52.7 ~ 58.8 'S: 64.9

79.9 s, 88.1 ~96.3

January 1974

55.3 ~ 62.7 &lt; 70.1

79.8 ~89.4

100 Females

~99.0

Productivity Estimates
Pre-season fawn:doe estimates prior to the 1973 deer season were 86.9 ±
11.8 fawns per 100 does. Productivity estimates obtained from JanuaryMay 1973 were 162 fawns per 100 mature doe deer (Roper 1973).
Mortality Estimates
Over-winter mortality for the 1972-73 winter was 6.4 percent or 647 deer.

Recommending and Publicizing Season Recommendations
In the 1973 deer hunting seasons Wildlife Conservation Officers reported
nearly complete compliance of hunters in having permits while hunting in
Middle Park. To accomplish this all roads entering the Middle Park area
were signed similar to 1972 (Roper 1973) and a news release in OUTDOOR
LIFE, October 1973, was printed along with newspaper articles and a publication in COLORADO OUTDOORS magazine (Roper 1973).
Estimate the Harvest by Sex
and Age Class and Success of Hunters
Harvest and Hunter Success Estimates
Sampling of hunters involved four different hunts. All hunters were required to have special permits to ~unt in Middle Park in the archery, regular
deer and post deer seasons. A. portion of Middle Park was open to high
country buck hunting by special permit only in the Ten-Mile mountain range.
Sampling was from finite populations of known permittees for each season
except archery. A total estimate of 1,771 deer were harvested in all seasons
(Table 9).

�-207-

Table 9.

Harvest of mule deer in Middle Park in 1973.

Type of
Hunt

Bucks

Does

Buck
Fawns

Doe
Fawns

Total

Percent

Archery

21

0

1

2

24

1.4

High Country
Buck

6

--

6

.3

Regular Deer

436

284

25

30

775

43.8

Post-Season
Deer

593

326

35

12

966

54.5

1,056

610

61

44

1,771

100.0

34.5

3.4

2.5

Total
Percent

59.6

100.0

During the archery season 545 permits were distributed. Hunters were not
limited to Middle Park and 205 hunters had 11.7 percent success in harvesting 24 deer.
During the regular deer season (October 13-23, 1973) hunters were limited
to hunting in Middle Park. Antlered-only permits were unlimited in number
while either-sex permits were limited to 2,000. A total of 4,200 permits
were issued of which ~,277 were antlered-only and 1,923 were either-sex.
An approximate 50 percent random sample of permit holders was used to
estimate the harvest. Of these the real sample size was 1,968 or 46.8
percent of the total permits.
The projected number of hunters was 3,485 (83.0 percent participation).
The success of all hunters was 22.2 percent. Either-sex permit holders
had a 41.3 percent success ratio and antlered-only hunters had 16.8 percent
success. Nonresident antlered-only hunters reported a 37.8 percent success
while 35.0 percent of the either-sex non-resident hunters harvested a deer
(Table 11).
The total regular season harvest was 775 ± 80 (P&lt; .05) deer (Table 10).
Three thousand either-sex post-season permits were available for 3,419
eligible hunters from December 15-23. We issued 2,157 permits and 77.1
percent (1,663) of the permit holders actually hunted in the post season.
Usable returns from a 33 percent random sample amounted to 31.2 percent of
the total hunters with permits.
The estimated harvest was 966 ± 93 (P&lt;.05)
cent (Table 10).

deer for a success of 58.1 per-

�-208-

Table 10. Comparison of the number of permits, hunters, success ratio
and harvest estimates in the October 13-23 and December 15-23, 1973
deer seasons in Middle Park, Colorado.
Season Dates
October 13-23
December 15-23
Total Number Permits Issued
Antlered Only Permits
Either Sex Permits

4,200
2,277
1,923

2,157

Total Projected Hunters

3,485

1,663

Percent Hunters Participating

83.0

77.1

Success Ratio
Antlered Only Permits
Either Sex Permits
All Permits

16.8
41.3
22.6

58.1

775 ± 80

966 ± 93

Harvest Estimate*

o
2,157

*P &lt; .05.
Sex and Age Structure of the Harvest
A comparison of the bucks, does and buck and doe fawns in the regular and
post deer season is tabulated in Table 11.
An incisor tooth was collected from a sample of deer from the harvest in
the regular and post seasons and will be sectioned and aged this coming
segment.

Table 11. A comparison of the harvest of bucks, does and buck and doe
fawns in the October 13-23 and December 15-23 deer seasons in Middle Park,
1973.
Season
Dates

Kind of
Permits

Bucks

Does

Buck
Fawns

Doe
Fawns

Total

October 13-23

2,277 Antlered Only
1,923 Either Sex

56.3%
436

36.6%
284

3.2%
25

3.9%
30

100.0%
775

December 15-23

2,157 Either Sex

61.5%
593

33.7%
326

3.6%
35

1.2%
12

100.0%
966

�-209-

Determine if Hunters Will Participate on a Continuing Basis
In 1972, 4,317 permits were issued and 3,336 (77.3 percent) hunters hunted.
In 1973, 4,200 permits were issued and 3,485 (83.0 percent) hunters participated.
We apparently haven't lost participation to date and some indications are
that as a result of the post season success ratio and satisfied hunters
that participated, the Middle Park experimental harvest season might gain
in popularity in 1974. This may be especially true if other areas in
northwestern Colorado provide more restrictive deer hunting opportunities
than in past years.

Determine if Specified Permits Help Obtain Harvest Goals
Based on the life equation, the pre-hunt population was 13,870 deer. Our
goal was to harvest 3,000 animals. Primarily because all of the post season
permits were not distributed the harvest was only 1,771 or 41 percent below
the goal. Plans are to allow any hunter to have the opportunity to apply
for a post-season permit after 10 days of preference to Middle Park permit
holders who were unsuccessful in the regular deer season. This should help
us come closer to harvesting deer within 10 percent of next year's predetermined harvest goal.

LITERATURE CITED
Gill, R. B. 1969. Middle Park Deer Study - population density and structure.
Colo. Game, Fish and Parks, Game Res. sec., Fed. Aid Proj. W-38-R.
Game Res. Re~., July, Part 1, pp. 105-122.
Gill, R. B. 1969. Middle Park Deer Study - productivity and mortality.
Colo. Game, Fish and Parks. Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rept., July, Part 1, pp. 123-140.
Gill, R. B. 1971. Middle Park Deer Study - population density and structure.
Colo. Game, Fish and Parks. Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rept., July, Part 2, pp. 170-188.
Riney, Thane. 1956. Differences in proportion of fawns to hinds in red
deer (Cervus elaphus) from several New Zealand environments. Nature.
177 :488-489 •
Roper, L. A. 1973. Middle Park Deer Study - experimental harvest regulations.
Colo. Div. Game, Fish and Parks. Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rept., July. (In press).
Sne decor, G • W " an d W ••G Cochran ,.1968
State Univ. Press, Ames. 593 p.

Statistical methods.

()..L&lt;.J..:;(~)=--_

Prepared by ....:...-L.jJ.d::.!t'W:.£l...l.~Z
Laren A. R~
Asst. Wildlife Researcher

The Iowa

��-211-

July, 1974

JOB PROGRESS REPORT

State of

COLOP~O
--~------~~~~~---------------

Project No.
Work Plan No.

W-38-R-28

Deer-Elk Investigations
Job No.~

14

9

~

_

Job Title Middle Park Cooperative Deer Study - Deer Habitat Evaluation
Period Covered: April 1,1973-

March 31,1974

Personnel: R. B. Gill, D. L. Baker, O. C.Wallmo, D. W. Reichert, L. A. Roper,
F. Deines, D. Wallsworth, G. Demerest, W. L. Regel in , L. H.
Carpenter, R. M. Bartmann and S. F. Steinert.

ABSTRACT
During the 1973-74segment a physical plant was constructed t.o facilitate
experiments designed to estimate the nutritional requirements of Middle
Park mule deer in winter. An initial study was designed and conducted
estimating the energy requirements of fawns in winter. Results of the
study are incompletely analyzed, but initial summaries are presented.

��-213-

MIDDLE PARK COOPERATIVE DEER STUDY DEER HABITAT EVALUATION
R. B. Gill and Dan L. Baker

P. S. OBJECTIVE
Establish practical procedures for managers to use in evaluating the
current and time-relative condition of deer ranges, including quantification of carrying capacity of "crucial ranges" in Middle Park, developing
the methodology for general use on game ranges in Colorado.
SEGMENT OBJECTIVES
1.

2.

Quantification of carrying capacity:
a)

Determine the number of deer (if any) in normal mid-winter nutritional state that can be carried through a 2-month late-winter
period on low elevation sagebrush winter range so as to retain
the deer in condition for satisfactory reproduction, exp'ressing
the results in animal units per land-area units.

b)

Determine the same for previously well-nourished deer (for information on advisability of directing attention to improving
early-winter deer use areas).

Game range evaluation and surveillance:
a)

Design a first-stage sampling procedure that (a) will permit
expression of quantities of deer forage of various quality and
the net quality of existing deer forage for range areas of
specific interest, that (b) can be integrated into the ground~
air~sate1lite
sampling procedure, and that (c) is feasible
for use by resource managers.

b)

Design the second-stage aerial photographic sample and signature
reference plots to integrate with above 1st stage sample units
a sample that will permit time-relative and place-relative
definition of important vegetation parameters.

METHODS AND MATERIALS

Physical Plant
The period 1973-74 was one of developing facilities to implement objectives
1a and 1b of the Habitat Evaluation Study. The Junction Butte property of
the Radium Wildlife Management Area was chosen as the site for these
facilities because of its proximity to Kremmling, Colorado (2 miles south

�-214-

of Kremmling), because some facilities already existed there, and because it is fairly typical of Middle Park's "crucial deer winter ranges".
The general layout of the facilities constructed in the past segment is
diagramed in Figure 1. Two pastures were constructed, one approximately
10 acres (4 hectares) and the other approximately 6 acres (2.4 hectares).
The larger pasture was built for studies of the energy budgets of unrestrained, tame deer in winter. The smaller pasture was constructed
to quantify aspects of cover and deer behavior that are important in
minimizing energy losses. A series of 18 individual deer isolation
pens were erected adjacent to an eXisting deer holding pen (Figs. 2 and
3). These pens were designed to measure the winter nutritional requirements of mule deer under conditions where diets could be controlled and
activity minimized. A weighing and handling facility was constructed
as a functional part of the isolation pen complex so that periodic weights
and blood samples could be collected with minimal disturbance to the deer
(Fig. 3).

Development of a Tame Deer Herd
Acquisition of Fawns
Twenty-six mule deer fawns were hand-reared during the period June 1 _
September 1, 1973 for the purposes of (1) investigating the nutritional
requirements and energy budgets of mule deer in winter, and (2) development of a nucleus tame deer herd for use in future research.
Mule deer fawns were obtained from 3 sources. Twenty-eight were taken
from captive wild does trapped the previous winter in Middle Park and held
until parturition. Sixteen were obtained from semi-tame does held at the
Foothills Deer Pens, Fort Collins, Colorado. Thirteen were orphans
collected from various areas in Colorado by Division of Wildlife Conservation Officers.
Newborn fawns initially were reared at Fraser Experimental Forest. Later,
because of continued, inclement weather in June and early July and resultant
high fawn mortality, fawn-rearing operations were moved to the Foothills Deer
Pens in Fort Collins. Fawns remained at this location until January 3, 1974,
when they were transported to the Junction Butte Deer Research Center near
Kremmling.

Fawn Rearing and Mortality
Fawns were taken from the dam approximately 12-24 hours after birth. Fawns
were bottle-fed and handled according to procedures outlined by Reichert
(1972). Out of an initial total of 57 fawns, 26 were reared to 6 months of
age. This represented a 54 percent mortality over the entire rearing period.
The major part of this loss (70 percent) was indirectly attributed to pneumonia
as a result of exposure to freezing rain, snow, wind, and sub-normal temperatures while the fawns were being held at Fraser Experimental Forest. Fawns
that died during this period were 2-3 weeks old. After they were moved to
Fort Collins, 6 fawns died of unknown causes associated with intestinal upsets.

�-215-

N

Fig. 1. Gsner-a L Layout of facilities
Research Center, 1973-74.

constrncted

at Junction Butte Deer

�-216Ci&gt;

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measure nutritional requirements of mule deero

�-217-

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pens and weigh b~lllding facilities

�-218-

Growth Rates and Training
Fawn growth rates were measured by taking weights weekly from 11 males
and 9 females. Up to 4 weeks old, all fawns were weighed by placing them
in a wooden transporting box, then weighing on a balance beam scale to the
nearest 1/10 pound (45.4 g). At 4 weeks of age fawns were trained to stand
on a platform scale located beneath the floor of a wooden handling chute.
They quickly accepted this procedure and would walk into the weigh chute
with little or no forcing by the handler.
Mean daily gain during the first 6 months was .37 pounds (.17 Kg) for males
and .31 pounds (.14 Kg) for females. Daily gain for both sexes combined
averaged .35 pounds (.16 Kg). This is less than the .53 pounds (.24 Kg)
daily. gain reported by Murphy and Coates (1966) for Missouri white-tailed
deer fawns and the .44 pounds (.20 Kg) per day reported by Cowan and Woods
(1955)for Columbian black-tailed deer over a similar period. Robinette
et al. (1973) reported a daily gain of .48 pounds (.22 Kg) for males and
.46 pounds (.21 Kg) for female mule deer fawns in Colorado. However, these
data are reported for fawns which were left with the dam. Fawns in our
study were bottle-fed from 24 hours after birth until weaning (approximately
84 days) at which time they weighed an average of 30.2 pounds (13.76 Kg).
Males were castrated at 3 months with an elastrator. The sex of fawns at
6 months for 26 survivors was 14 males and 12 females compared to the
original 57 fawns of which 34 were males and 23 were females.
Since these fawns were acquired to measure maintenance energy requirements
of mule deer under winter conditions, special training was required to
obtain data on energy losses in urine and feces. Attempts were made to
train fawns to accept attached urine-feces collection bags. While a
functional urine-feces collection bag was being developed, fawns were
trained to wear a light weight harness an an elastic training bag. A bag
to collect uncontaminated samples of both urine and feces was not perfected
during this segment. But, a partially successful feces collection bag was
designed and tested on male fawns. This bag was constructed from a small
gauge nylon mesh with a drawstring at the bottom to allow for daily feces
collection without removing the bag. The feces bag was attached directly
to the rump of the deer with winter viscosity branding cement.
Pre-trial testing indicated that male fawns would carry the bag for at
least 24 hours. However, during actual trial conditions, fawns failed to
carry the bags for more than 3 hours at a time. The special low-temperature
branding cement held the bag in position adequately, but fawns were able to
grasp the bags orally and remove them. Because fecal collection bags were
not usable to estimate both urinary and fecal energy losses, complete
balance trials to determine digestible and metabolizable energy of feeds
will be conducted during the next segment in Fort Collins.
Energy Requirements of Fawns in Winter
Objectives and Experimental Design
This phase of the study was conducted in conjunction with objectives and
manpower availability of Work Plan 14, Job 6 and is really an unexpected

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bonus for Work Plan 14, Job 9 as it puts us roughly one year ahead of
our programmed schedule.
The primary objective of this study was to estimate metabolizable energy
requirements of captive mule deer fawns for winter maintenance. Subsidiary
objectives were: (1) to determine minimum dietary energy requirements
(grams of metabolizable energy/kilogram of body weight .75) for winter
survival of mule deer fawns; (2) to determine length of survival (days)
on a given level of energy intake; (3) to determine dietary energy
deficiencies by measuring blood metabolites; and (4) to measure and relate weather parameters (ambient temperature, relative humidity, and wind
speed and direction) to energy balance.
To accomplish these objectives, 18 hand-reared fawns (10 males and 8 females)
were selected. The deer were transported from the Foothills Deer Pens in
Fort Collins to the Junction Butte Deer Research Center near Kremmling on
January 2, 1974. Energy experiments began January 4, and ended March 28,
1974.
Fawns were acclimated to the Kremmling environment 10 days prior to initiation
of actual feeding trials. Each fawn was assigned to an individual isolation
pen at random in order to minimize pen effects on treatment results. Each
isolation pen measured 10 x 30 feet (3.04 x 9.12 m) and consisted of woven
wire sides and dirt floors denuded of vegetation. A 10 x 10-foot (3.04 x
3.04 m) wooden roof partially covered each pen. Then the 18 fawns were
grouped into trios on the basis of metabolic weight so that each trio
contained. fawns of approximately equal weights. Next the six sets of
trios were ranked from lightest to heaviest. Thus an experimental design
was framed to accommodate 3 treatments (energy levels of feeds) with 6
replicates deer per treatment, so that weight variation within treatments
was similar for all 3 treatments (Fig. 4). All deer were fed a pelleted
ration (Table 1) which they had been receiving ad libitum 4 months prior
to the start of the experiment.
Treatments were assigned to each of the 3 groups of 6 deer as follows:
treatment A deer were offered more food than they could consume in a 24
hour period (food offered ad libitum), treatment B deer received 75 percent
of the ad libitum removal rates of treatment A deer, and treatment C deer
received 50 percent of ad libitum intake (Table 2). Treatment A deer removed an average of 2.601pounds (1.22 Kg) of food per day for a 2-week pretrial period, but were offered 4.56 pounds (2.08 Kg) per day during the
experiment to guarantee sufficient food for ad libitum removal. Treatments
Band C deer were offered 2.01 and 1.34 pounds per day (.915 and .610 Kg),
respectively. Expressed as a function of metabolic weight, treatments A,
B, and C deer received 134.93, 59.36, and 39.57 grams of dry matter/
kilogram of body weight .75, respectively. Correlations were sought between treatments and blood metabolite levels, weight changes over time,
and weather variables. Snow or water was provided each deer ad libitum.

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14
12

10

8
6

Treatment
A

4
w

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u

t
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2

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B

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C

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o

1

2

J

4

5
WEEKS

6

7

8

9

10

Fige 4. Average weekly weight changes (percent) of fawns on three different
levels of dietary ener-gy, Biddle Park, Colorado, Winter 1974.

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Table 1.

Composition of pelleted feed.

Ingredient

Percent

Barley, pulverized

10.0

Corn, pulverized

30.0

Milo, pulverized

5.0

Oats, pulverized

7.5

Wheat Middlings

(Mill Feed)

6.5

Beet Pulp, shredded

2.5

Brewers Grain

35.0

Dicalcium Phosphate

1.0

Molasses, cane

2.5

Vitamin A, D, E. Premix

.002

Trace Mineral Package (Mg, Zn, I, Fe, Co, Cu, Ca)

.005
100.00

Table 2. Experimental design framework for study of maintenance energy
requirements of mule deer fawns in winter.

Rep. 1

Treatment A
Lightest Deer

Treatment B
Lightest Deer

Treatment C
Lightest Deer

Heaviest Deer

Heaviest Deer

Heaviest Deer

Rep. 2
Rep. 3
Rep. 4
Rep. 5
Rep. 6

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Metabolizable Energy Intake
Food removal and orts were weighed daily from January 4 - March 28, 1974
to estimate gross energy intake. Because fecal collection bags were not
successful~ complete energy balance trials to estimate metabolizable
energy intake were scheduled at the Foothills Deer Pens in Fort Collins
at the conclusion of the winter trials in Middle Park. Mean percent in
vivo digestibility will be determined for 4 fawns selected at random --from each treatment. Each fawn will be placed in a 10 x 10 foot (3.04
x 3.04 m) metabolism cage for a 7 day collection trial. Total urine and
feces will be collected. Samples of feed and feces will be analyzed for
dry matter, crude protein ether extract, gross energy, ash, cell wall
constituents, acid detergent fiber, and lignin. Urine will be analyzed
for gross energy and nitrogen. Energy lost in methane production will
be predicted from equations in Blaxter and Clapperton (1965).
Body Weight Changes
All fawns were weighed at the beginning of the feeding trials and at
weekly intervals thereafter. Trained fawns walked through a specially
designed holding chute consisting of a platform scale situated beneath
a wooden floor. Body weights were recorded to the nearest 1/4 pound
(.11 Kg).

Blood Metabolites
Blood samples were taken from fawns at weekly intervals for additional
information on energy balance. A minimum of 12 fawns (4 from each treatment) were sampled each week. Fawns walked into the weigh chute, were
weighed, and then proceeded into a padded squeeze chute designed for
taking blood samples without injury to deer or handlers. A 6~inch diameter (15.24 cm) hole was cut in the squeeze chute exit door to allow a
deer's head and neck to protrude. The hole and exit door were padded
with 4 inches (10.16 cm) of foam rubber. Both sides of the squeeze
chute were horizontally hinged, one side at the top, and the other at the
bottom, so that firm even pressure could be applied to immobilize the deer.
Each side was padded with 6 inches (15.24 cm) of foam rubber so the sides
could conform to deer body contours. Then, one handler squeezed the deer's
body, another grasped its head and positioned it for blood sampling, while
a third collected the blood.
The entire process of weighing and blood letting took approximately 90
seconds per deer. Blood sampling alone took 20-30 seconds from the time
a deer was squeezed until blood was collected. Ideally, 10 milliliters
of blood were taken from the jugular vein of each deer. Samples were taken
with a 12 milliliter syringe and a 1 inch, 18 gauge needle. Blood samples
were placed in heparinized tubes and gently rocked to mix heparin and blood
to prevent coagulation. Whole blood was kept chilled and centrifuged within 2 hours of taking the first sample. Blood plasma was pipetted and frozen
immediately for later analysis.

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Plasma samples will be analyzed for non-esterified fatty acid levels
(NEFA), blood urea nitrogen (BUN), glucose, and total protein. Factors
other than nutritional stress may affect NEFA and glucose 1evels~ such
as excitability of the deer at the time of sampling. Since our deer
were unavoidably excited, plasma will also be analyzed for abnormal
levels of lactic acid and epinephrine.
Weight changes, food intake, and blood metabolite values will be correlated with ambient temperature, wind speed and direciton, and relative
humidity which were recorded continuously at the Junction Butte Deer
Research Center during the feeding trials.

Game Range Evaluation and Surveillance
Accomplishment of these objectives (2a and 2b) was contingent upon contractual support funding from the Bureau of Land Management (BLM) and the
National Aeronautics and Space Administration (NASA). Unfortunately
neither proposal was funded and, consequently, the objectives were not meto
RESULTS AND DISCUSSION
Physical Plant
Construction of the Habitat Evaluation Study facilities (the Junction Butte
Deer Research Center) was only partially completed in the 1973-74 segment
Five gates remain to be constructed and numerous small gaps at the bottom
of pasture fences need to be sealed before these structures are "deerproof". Also poultry mesh aprons should be attached to the bottoms of the
pasture and isolation pen periphery fences to preclude entry by dogs and
coyotes. An observation tower overlooking the 10 acre pasture is scheduled
for construction in 1974-75 segment. Within the isolation pen complex, the
lab and storage building needs to be erected and the loading chute runway
fence awaits completion. Most materials for construction are already stockpiled, so that only manpower and time are necessary to complete the construction of facilities. Long range goals include the installation of a
water system and an electric power system.
0

Additional fawns will be reared in 1974-75 and, barring catastrophic mortality, we anticipate adding 30-40 fawns to our tame deer herd. These fawns
will be used to estimate energy requirements and expenditures under both
controlled pen and free-ranging conditions in the coming winter.
Energy Requirements
Metabolizable Energy Intake
Mean daily intake for the 6 fawns in the ad libitum treatment (treatmemt A)

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was 1.086 ± .0~8 Kg (P&lt;.05) for the 10 week feeding trial. Expressed
in terms of metabolic size this amounted to 70.03 ± 2.45 g DM/ Kg BW .75.
Treatment Band C fawns consumed .910 ± .06 and .610 ± .00 Kg per day or
59.95 ± 0.40 and 42.,75 ± 0.00 g DM/ Kg BW .75, respectively. When daily
intake is expressed as a percentage of body weight then treatment A fawns
constnned an equivalent of 2.85 percent of their body weight per day,
treatment B deer - 2.44 percent, and treatment C deer - 1.76 percent.
Samples of the pelleted ration were collected and will be analyzed for
gross energy content to convert dry matter intake to gross energy intake.
Digestion and energy balance trials now in progress will provide information to convert gross energy intake into digestible and metabolizable
energy intake. These data will then be used to predict apparent metabolizable energy requirements for maintenance from regressions of average
daily metabolizable energy intake on average daily weight gains or losses.
The X axis intercept (energy intake) represents the predicted energy intake
at energy equilibrium or maintenance.
Body Weight Changes
Fawns on all treatments lost weight during the first two weeks of trials
(Fig. 4). After 2 weeks treatment A and B fawns gained weight. Treatment
B fawns lost more weight during the initial 2 weeks and gained more slowly
than treatment A fawns. Treatment C fawns lost weight through 8 weeks and
then apparently gained over the final 2 weeks. Initial conclusions are
that treatment C dietary energy levels were below maintenance levels for
growing fawns subjected to the environmental extremes of the 1973-74 Middle
Park winter.

Blood Metabolites
Blood plasma samples have not been analyzed, but will be during the coming
segment (1974-75).

LITERATURE CITED
Blaxter, Ko L., and J. L. Clapperton. 1965. Prediction of the amount of
methane produced by ruminants. Brit. J. Nutr. 19(4):511-522.
Cowan, I. McT., and A. J. Wood. 1955. The growth rate of the black-tailed
deer (Odocoileus hemionus columbianus). J. Wildl. Manage. 19(3):3313360
Murphy, D. A., and J. A. Coates. 1966. Effects of dietary protein on deer.
Trans. No Amer. Wildl. Conf. 31:129-138.

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Reichert, D. W. 1972. Rearing and training deer for food habits
studies. U.S.D.A. Forest Service. Rocky Mtn. For. &amp; Range
Exp. Sta. RM Note 208. 7p.
Robinette, W. L., C. H. Baer, R. E. Pf.Llmo're , and C. E. Knittle
1973.
Effects of nutritional change on captive mule deer. J. Wildl.
Manage. 37(3):312-326.
0

Prepared by
R. Bruce Gill
Wildlife Researcher

��-227-

July, 1974

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-28

Work Plan No. .....;..._1;;;.4~...,....,.,~,..".-____
Job No. ~-=10::-. ,,-'o
'-;"
Middle Park Cooperative Deer Study
Job Title
Junction Butte Wildlife Habitat Improvement Project
Period Covered:

April 1, 1973 to March 31, 1974

Personnel:

Regelin, O. C. Wallmo, R. A. Schmidt

W.L.

_

.ABSTRACT
Snowfences produced drifts averaging 28.7 inches in depth and reduced snow
.depth to an average of 15 inches in protected shrub stands. Neither soil
temperature nor moisture content differed significantly (P &lt; .05) between
snowdrift-covered and control quadrants. CAG of shrubs has not been
influenced by altering normal snow distribution.
All Purshia tridentata
seedlings planted in .October 1971 were dead by the summer of 1972 •. Thirty
percent of the seedlings planted:l.n May 1973 were still alive 2 months
later. Seeds ofChrysothamnusviscidiflorus,
t. nauseosus, and Purshia
tridentata planted. in October 1971 have failed to establish any plants.
Good. stands of Agropyron.desertorum and Melilotus officinalis·have been
established on all seedbeds. Carbon black applied to the snow surface
was effective in producingmid~inter
snow melt at temperatures well
below freezing.

��-229-

MIDDLE PARK DEER STUDY JUNCTION BUTTE WILDLIFE HABITAT IMPROVEMENT PROJECT
Wayne L. Regelin

P. S. OBJECTIVE
To increase availability of forage on critical winter ranges by relocating
snowdrifts and determine if changes in vegetative composition occur due
to altering snow depth.

SEGMENT OBJECTIVES
1.

Measure snow depth and density and configuration of snowdrifts in
relation to presence or absence of snowfences and in relation to
shrub stands protected and unprotected from the accumulation of
snowdrifts.

2.

Measure occupation by deer of shrub stands protected and unprotected from the accumulation of snowdrifts.

3.

Measure the relative use of browse by deer in shrub stands protected
and unprotected from snowdrifts.

4.

Measure the response of native vegetation (composition and density)
in relation to increased or decreased snow depth.

5.

Evaluate the success of forage species planted on sites of created
snowdrifts and on control areas not covered with drifts.

6.

Measure soil moisture with relation to increased or decreased snow
accumulation.

7.

Test the effectiveness of lamp black to accelerate snow melt in
shrub stands protected by snowfences or natural windbreaks, and
determine if changes occur in density and/or crusting characteristics
of the snow due to treatment with carbon black.

METHODS AND MATERIALS
Prior to the current segment, this work was supported entirely by State
funds; therefore, methodology has not been reported in the Federal Aid
in Wildlife Restoration reporting system. Procedures used to meet all
segment objectives, except number 7, are explained in Study Plan No.8
of Research Work Unit FS-RM-1709, Forest Service, entitled "Improvement

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of Deer Winter Range by Manipulating Snow Cover in Middle Park, Colorado"
(Appendix A). Segment objective number 7 was initiated in 1974. Study
Plan No.8, Amendment 1, explains the procedures used to meet this objective (Appendix B). Briefly, the snowdrift manipulation study consists
of nine sites on the Junction Butte property of the Radium Wildlife
Management Area. Five sites have snowfence to relocate snowdrifts formerly covering dense browse stands; each of these sites is associated
with a paired control site. Four sites have snowfence whose primary
purpose is to test the effectiveness and economy of different fence
structural designs. Appendix C includes sections of the Second Annual
Progress Report dealing with the different structural designs of snowfences.

RESULTS AND DISCUSSION
Weather and Snow Measurements
The anemometer and drift gauge were installed for the 1973-74 winter on
December 24, 1973 at the termination of the post-season deer hunt.
Biweekly measurements of snow accumulation behind the snowfences and
on adjacent controls began December 23, 1973 and continued until April 20,
1974. The first major snowfall of the winter occurred on December 28 and
29 when approximately 12 inches of snow fell. This storm was accompanied
by wind in excess of 40 mph and produced the only significant drifting
event during the winter. During January and February, only small amounts
of snow fell, none accompanied by high winds. Windspeeds did exceed 50
mph several times during this period, but only after the snow had crusted,
resulting in little snow being drifted.
The maximum average snow depth behind the five snowfences on the south
side of Junction Butte was 28.7 inches. The deepest drift measured at
any point was 56 inches. Average peak snow depth in the shrub stands
protected by the snowfences reached 15 inches, which is safely below
the limit of trafficabi1ity for deer (Gilbert et a1. 1970). In the
unprotected control shrub stands, average peak depths were 28 inches.
Relative to control areas, the snowfences reduced snow depth an average
of 12.8 inches in the protected shrub stands. Total snowfall during
the winter was approximately 60 inches, but only 12 inches of this total
was subjected to winds strong enough to transport snow. Table 1 shows
the peak average snow depth in each quadrant as measured on February 5,
1974.
Deer Occupancy of Snowfence Sites
Table 2 summarizes the track count data on deer occupancy of areas behind
each snowfence and the adjacent controls. No track crossings were observed on shrub stands where the snowfences reduced drift accumulation
until February 16, 1974, even though many deer were sited on the mountain
prior to this date. The south slopes remained essentially free of snow
throughout the winter except for short periods after light ~nowfa11.

�-231-

Incidents of track crossings were much higher in the areas where normal
drifting was prevented by the snowfences than in adjacent control areas.
While the sum of track counts is impressive, variability among sites
and dates prohibits any inference of statistically significant differences.

Soil Temperature and Moisture
The average soil temperature in each quadrant of sites 2, 3, and 4, at
3-inch depth, is shown in Table 3. Soil temperature did not differ significantly (P &lt; .05) between treatment and control quadrants at any date.
On sites 1 and 5, where seedbeds were established to study revegetation
methods, soil temperatures did not differ significantly (P &lt; .05) between
beds covered with snowdrifts and control beds or between cleared and
furrowed seedbeds at any of the dates measured (Tables 4 and 5).
Soil moisture content has been measured to a depth of 10 inches in each
quadrant of sites 2, 3, and 4 for the last 3 years during spring and
summer. No significant differences (P &lt; .05) in soil moisture content
between treatment and control quadrants have ever been measured, even
though snow depth on these quadrants has differed greatly. To determine
if the water available in the.snowdrifts is stored below 10-inch,depths,
Bouyoucous soil moisture blocks were installed at depths of 16, 22, 34,
and 46 inches in September 1973. Blocks were placed at three locations
within each quadrant at each depth on sites 2, 3, and 4.
Beginning in the spring of 1974, soil moisture content will be measured
at these depths. Tables 6 and 7 show the average soil moisture content
in the quadrants of sites 2, 3, and 4 and the seedbeds of site 1 during
1973.
Growth of Native Plants
Current annual growth (CAG) on the tagged branches of the five shrub
species was measured in September 1973. Table 8 summarizes the CAG
of each species by quadrant for the 3 years data have been collected.
The created snowdrifts which cover the A quadrants have had no effect
on CAG of any species when compared to shrub growth in the control
quadrants (C). Also, reducing snow depth in B quadrants with the
snowfences has not reduced the CAG of these shrub species when compared
to CAG in D quadrants where normal drifting occurred.
Data on species composition and density of native plants in each quadrant
were collected for the third year during July 1973. Analysis of these
data will not be done until the final year's data have been collected,
so covariance analysis can be utilized to determine if altering the
normal snow depth influences species composition and density.

�-232-

Establishment

of Planted Species

Bitterbrush (Purshia tridentata) seedlings were first planted in October
1971 in dry soil and each was initially watered. All seedlings were dead
by the summer of 1972. A second planting was done in moist soil on May 1,
1973 and not watered. Other planting procedures were the same, but native
plants which had invaded the seedbeds since clearing in 1971 were not
disturbed.
Survival of the seedlings was observed on July 30, 1973; results are summarized in Table 9. Average survival 2 months after planting was 30 percent on both sites 1 and 5, even though site 5 has much shallower soil.
No difference in survival was observed between snowdrift-covered and
control seedbeds, but seedbeds cleared of all vegetation in 1971 had
35 percent more live seedlings than furrowed seedbeds. Seedbeds fertilized in 1971 had a 15 percent lower survival rate than unfertilized
seedbeds, possibly due to increased competition for moisture with native
species in the fertilized seedbeds.
Almost no plants were established from the plantings of green rabbitbrush
(Chrysothamnus viscidiflorus), rubber rabbitbrush (£. nauseosus), and
bitterbrush seeds. An occasional plant of these species is encountered
in the seedbeds, but the plantings of these species must be termed a
failure.
Good stands of crested wheatgrass (Agropyron desertorum) have been established in all seedbeds. Linear ground cover per row was greater in 1973
than in 1972 on all seedbed types (Table 10). Average plant height in
1973 was 11.6 inches compared to 3.5 inches in 1972. Nearly all plants
produced seed heads. The best establishment and growth occurred on
cleared seedbeds. Seedbeds covered with snowdrifts produced less linear
cover per row than control seedbeds, possibly due to winter kill under
the snowdrifts or the delay in the start of the growing season.
Yellow sweetclover (Melilotus officinalis) plants were established in all
seedbeds the second year after planting, but establishment in cleared
seedbeds was much greater than any other seedbed type. The number of
plants per row decreased in all types of seedbeds from 1972 to 1973
(Table 11). Even though there were fewer individual clover plants per
row, the linear ground cover increased due to greater plant height.
Average plant height in 1973 was 12.1 inches compared to 2.6 inches in
1972. Several plants were over 36 inches in height. Nearly all the
plants produced large quantities of seeds, which are a potential source
of reseeding.
The 1974 measurements will indicate if this occurs.
Native revegetation of the seedbeds which were cleared of all plants in
1971 is occurring more rapidly in seedbeds covered by created drifts
than on the control seedbeds. Two years after clearing, the seedbeds
covered with snowdrifts have over twice the number of native plants
reestablished per unit area as the control seedbeds (Table 12).

�-233-

Experimental

Snow Melt

Seven sites, each consisting of two paired 10-meter-square plots, were
located on Junction Butte in areas protected from wind-blown snow
accumulation.
Three sites have a southerly aspect and four northerly
aspects; all are on nearly level terrain. Carbon black was applied
to one plot in each site at the rate of 336 kg/ha (330 lb/acre) using
a back-pack fertilizer spreader. First application of carbon black
was made January 30, 1974; two subsequent applications at the same
rate were made on February 4 and 9, 1974. Snow depth in each treatment
and control plot was measured daily after the first application of
carbon black. Within each plot, 13 snow ablation stakes, marked in
10-em increments, were systematically located to facilitate snow depth
measurements.
Snow density was determined at five locations within
each plot using a MOntrose snow sampler. Snow density was determined
four times during the study, immediately before the initial application,
before the two subsequent applications, and at the termination of the
experiment.
Hardness of snow crust was measured at these same four
times using a Ram Penetrometer.
Details of the methods employed are
given in Appendix B.
The snow melt rate was accelerated on all treated plots. The ef{ect of
the carbon black was more pronounced on the three sites with southerly
aspect where the average snow depth was reduced from 42.9 cm to 3.6 cm
compared to control plots where snow depth decreased naturally from
40.6 to 37.6 cm. Average snow depth on the four treatment plots with
a northerly aspect was reduced from 53.3 cm to 27.4 cm, while no
decrease occurred on control plots. Figures 1 and 2 show the average
melt rates on the two different aspects. A light snowfall, 2 days
after the first application, covered the carbon black with 2 to 3 cm
of snow. No melt occurred on the plots with northerly aspects and' only
slight melt on plots with southerly aspects for th~ next 2 days. This
indicates that a light snow cover greatly reduces the effectiveness of
carbon black in producing snow melt. Carbon black was applied again
on February 4, 1974, but 7 to 8 cm of snow fell that evening, covering
the new application.
The third application of carbon black was made
on February 9, 1974. Clear, cold weather prevailed for several days
after this application resulting in rapid melt on all sites. The
quantity of carbon black on the snow surface was increased 3-fold
shortly after this melt period began because the two prior applications
of carbon black were again on the snow surface. By February 11, 1974,
48 hours after the last application of carbon black, the three treatment plots with southerly aspects had melted to bare ground on 30 percent
of their area. On February 13, 1974 over 60 percent of the area in these
plots had bare ground showing.
At the termination of the study, 16 days after the first carbon black
application, all snow on the treatment plots had melted to bare ground,
except in small patches shaded by tall shrubs. The plots w~th northerly
aspects retained more snow at the termination of the experiment, but
small areas within the plots did melt to bare ground.

�-234-

The average temperature during daylight hours during the entire experiment
was -6.7 C. The highest temperature was +5.4 C. Table 13 shows the
daily temperature fluctuation and mean temperature during daylight hours.
Snow did melt on the treatment plots when the air temperature was below
freezing. Free standing water near edges of the plots was observed when
air temperature was -16 C.
Density of snow did not differ significantly (P &lt; .05) between treatment
and control plots. Covariance analysis was used to test differences
between plots with means adjusted for pretreatment values. Adjusted
treatment means were slightly higher than control plot means prior to
both the second and third applications of carbon black, but the differences were not significant. The average density of snow on all sites
was approximately 30 percent.
Hardness of snow crust varied greatly within treatment and control plots.
The Ram Penetrometer was not powerful enough to break the crust in places,
but would fall through the snow to the ground without applying any force
only a few feet away. These extremely variable crust conditions did not
permit quantitative analysis of data on crusting characteristics of the
snow. From general observation, the carbon black treatment did not
appear to alter the hardness of snow crust until rapid melting occurred
and snow depth was shallow. At this time, the snow was very wet and
slushy and at night an ice crust would form on the surface of such snow.
These results indicate that application of carbon black can produce
mid-winter snow melt at subfreezing temperatures. A light snow cover
greatly reduces the effectiveness of the carbon black. The carbon
black produced significant snow melt on both southerly and northerly
exposures, but the results on southerly slopes were more pronounced.
There was evidence of grazing use and bedding by deer on some melted
plots, but the study was not designed to obtain quantitative information
on the use of melted areas.

LITERATURE CITED
Gilbert, P. F.,O. C. Wallmo, and R. B. Gill. 1970. Effect of snow depth
on mule deer in Middle Park, Colorado. J. Wildl. Manage. 34(1):15-22.

�-235-

Table 1. Peak average snow depth of five snowfence sites on the south
side of Junction Butte during the 1973-74 winter.

Quadrant

28.7

Snow depth in inches

1/

B

C

D

15.0

9.7

27.8

A - New snowdrift created by snowfence.
B - Shrub stand with drift removed by snowfence.
C
Control area, normally no drift.
D
Control area, shrub stand normally drifted.

Table 2.
Butte.

Summary of track crossings at the nine study sites on Junction

Snowfence Areas

Location

Created
drift

Feb. 16, 1974

S

0

Feb. 16

N

Feb. 26

Reduced
drift

Control Areas
No drift

Drift

9

9

0

1

110

0

0

S

0

0

O·

0

Feb. 26

N

3

57

0

0

March 11

S

0

0

0

0

March 11

N

0

0

0

0

March 24

S

0

9

24

3

March 24

N

0

0

0

0

4

185

33

3

Date

Total for Winter

�-236-

Table 3. Average soil temperature at 3-inch depth in each quadrant of
sites 2, 3, and 4.

Date
Quadrant

12 April

3 May

16 May

25 June

17 August.

A

30.7

37.8

69.4

61.4

71.2

B

30.6

32.2

56.3

51.6

56.6

C

31.4

35.5

59.2

57.5

65.5

D

30.5

32.2

52.9

50.2

55.6

Table 4.

Average soil temperature at 3-inch depth in seedbeds on site 1.

Date
12 April

3 May

16 May

25 June

17 August

All beds covered by drift

30.9

34.9

70.4

57.0

73.2

All control beds

30.6

41.8

68.2

55.9

76.8

Cleared beds

30.5

38.3

71.3

56.6

75.9

Furrowed beds

31.0

38.4

67.3

56.2

74.1

�-237-

Table 5.

Average soil temperature at 3-inch depth in seedbeds on site 5.

Date
12 April

3 May

16 May

25 June

17 August

All beds covered by drift

40.1

45.8

75.4

64.2

77 .2

All control beds

40.9

42.2

74.4

60.8

70.8

Cleared beds

40.4

43.2

74.4

62.0

75.4

Furrowed beds

40.6

44.8

75.4

63.0

72.5

Table 6. Average percent soil moisture in each quadrant of sites 2, 3,
and 4. The percentage is the average of three determinations at 0-2,
4-6, and 8-10 inch depths, at two sampling points within each quadrant.

Date
Quadrant

17 May

25 June

18 July

16 August

A

15.8

8.1

22.5

9.4

B

22.4

13.3

23.1

8.4

C

18.3

10.4

24.7

8.9

D

29.6

19.7

27.0

15.8

�-238-

Table 7.

Average percent soil moisture in site 1 seedbeds.

17 May

25 June

18 July

16 August

All beds covered by drift

17.1

9.1

23.7

8.1

All control beds

16.3

10.5

23.0

8.7

Cleared beds

17.1

10.9

23.2

9.1

Furrowed beds

16.3

8.7

23.5

7.7

�1/

Species Code:
AIDa1 - Ame1anchier

a1nifo1ia

Syor - Symphoricarpos

oreophi1us

Rice - Ribes cereum
Prvi - Prunus virginianus
Putr - Purshia tridentata

�-240-

Table 9. Survival of bitterbrush seedlings in various types of seedbeds
2 months after planting.
Twenty seedlings were planted in each seedbed.

Percent of seedlings surviving
Seedbed TyPe

Site 1

Site 5

Avg. on both sites

Seedbeds covered by snowdrift

37.5

20.0

28.8

Seedbe~s in control

23.7

39.0

31.4

All fertilized

28.8

16.2

22.5

32.5

42.5

37.5

All cleared seedbeds

51.0

44.0

47.5

All furrowed seedbeds

10.0

15.0

12.5

seedbeds

All unfertilized

seedbeds

Table 10. Summary of establishment of Agropyron desertorum
types of seedbeds 1 and 2 years after planting.

in the various

Total feet of linear ground
cover per 30-foot row,l/
Seedbed Type

1971

1972

Seedbeds covered by created drift

9.91

12.92

Seedbeds in control area

9.70

15.75

All fertilized

10.66

16.67

8.95

12.00

All cleared seedbeds

14.48

18.36

All furrowed seedbeds

5.12

10.32

All unfertilized

1/

seedbeds
seedbeds

Figures are average of seedbed from both planting sites.

�-241Table 11. Summary of establishment of Melilotus
types of seedbeds 1 and 2 years after planting.

officinalis

Avg. number of
plants/30-ft row

in the various

Total feet of linear
ground cover/30-ft row

Seedbed Type

1972

1973

1972

1973

Seedbeds covered by created drifts

40.9

15.25

6.83

6.61

Seedbeds in control area

44.6

16.85

7.60

8.62

All fertilized

33.7

10.70

5.97

6.68

50.5

21.4

8.46

8.56

All cleared seedbeds

57.6

26.5

12.04

13.55

All furrowed seedbeds

27.8

5.6

2.04

2.22

All unfertilized

seedbeds
seedbeds

Table 12. Average number of native plants per square-foot plot on cleared
seedbeds 1 and 2 years after clearing.
The average is obtained from 90
1 X l-ft plots per seedbed.

Average number plants per square foot
1972

1973

All drift covered seedbeds

3.42

6.95

All control seedbeds

1.58

3.30

Drift covered seedbeds, site 1

2.84

6.57

Control seedbeds, site 1

2.17

4.53

Drift covered seedbeds, site 5

4.00

7.33

Control seedbeds, site 5

1.00

2.07

�-242-

Table 13. Daily temperature in C during daylight hours on Junction Butte
during snow melt study.

Date

Mean
Temperature

High
Temperature

Low
Temperature

"1-31-74

- 5.5

-0.6

-12.2

2-1

- 5.2

-3.9

- 8.3

2-2

- 9.5

-5.6

-13.3

2-3

- 8.6

-1.7

-16.1

2-4

- 7.0

-1.1

-15.0

2-5

-10.6

-6.1

-12.2

2-6

- 9.9

-3.3

-17.8

2-7

-14.1

-8.9

-20.6

2-8

-15.2

-8.3

-25.0

2-9

-10.9

-3.3

-22.2

2-10

-10.4

-1.7

-19.4

2-11

- 8.6

-2.2

-19.4

2-12

- 4.6

-0.6

-11.6

2-13

- 5.4

+5.4

-15.6

2-14

- 7.1

+3.6

-19.4

2-15

- 8.1

0

-16.7

�45
42

ontro1 plots

39
36
33
30

plots
27
snow
depth
em.

24

,

21

N
.p-.
W
I

18
15
12
9
6
3
0
0'\

N
I
.-I

0
C"')
I
.-I

Fig. 1.

.-I
C"')

I

.-I

Date
.-I

I

N

N

I

N

C"')

I

N

-:T
I
N

Ifl

\0

l"-

I
N

I
N

N

I

.-I
.-I

N
.-I

.-I

-:T
.-I

Ifl

0'\

o
.-I

C"')

00

I

I
N

I

I

I

I

I

I

N

N

N

Average melt rate on three sites with southerly exposure.

N

N

N

.-I
N

\0
.-I
I
N

�54
51
48
45
42
39
36

snow
depth
em.

J

~

Treatment plots

33
30

27

~

"

I

N

+:+:-

'-

I

I

24
21
18
15
12

0\
N

~

o
M
,...fI

Fig. 2.

,...f
M

,...fI

,...f
I

N

N

I

N

M

I

N

..;t
I

N

LI"\

\0

I
N

I
N

,....
I
N

00
I

N

0\
I
N

~

I

N

,...f
,...f

N

,...f

M

,...f

..;t

I
N

I

I

I

Average melt rate on four sites with northerly exposure.

N

N

,...f

N

LI"\

,...f
I

N

~

I

N

�-245-

APPENDIX A

��-247-

Wildlife Habitat - Central Rockies
FS-R.'1-l80l

STUDY PLA.~ 8

Improvement
by Manipulating

of Deer Winter Range

Snow Cover in Hiddle Park, Colorado

Rocky Mountain Forest and Range Experiment

Station

Fort Collins, Colorado

PREPARED

BY:

DATE;).!/"
, ;I 7 ZDATE

a Ii /72 .
/'

APPROVED

BY:

DATE

-3/' /72-

��-249-

The Problem
Availability of winter forage is an important factor limiti~g
deer population in much of Colorado and other areas in the central
Rocky Mountains. On most winter ranges snow, more than any other
factor, determines the availability of forages for deer CWallmo and
Gill 1970). Southerly slopes and windswept ridges commonly are
severely overgrazed while dense stands of shrubs remain inaccessible
under adjacent snowdrifts. Relocation of such snowdrifts could make
additional forage available for deer use. The necessary theory and
techniques for manipulating snowdrifts 1n accordance with watershed
objectives have been developed by Forest Service scientists (Tabler
1968, Schmidt 1970). The feasibility of applying these techniques
to the improvement of wildlife. habitat will be tested in this project. This study is conducted under the Research Work Unit assigned
Problem Number 1, Evaluation of Factors on Sagebrush Ranges that
Determine Quality of Wildlife Habitat.
Middle Park is a mountain valley in central Colorado, predominated by big sagebrush and enclosed by high mountains which trap
all deer within the Park during the winter. The study. area is on
Junction Butte, a small isolated mountain in the lowest winter range
in Middle Park. The southerly slopes on Junction Butte are heavily
occupied by deer in winter. In general, the heaviest stands of browse,
principally Amelanchier alnifolia, Symphoricarposoreophilus;
Prunus
virginianus, Ribes cereum, and Purshia tridentata occur where snowdrifts prevent their use by deer. These browse stands offer areas
where the relocation of snowdrifts to alleviate forage shortages can
be evaluated.
Literature Review
Snowfences and vegetative windbreaks have been used for many
years to control blowing snow on agricultural lands and along highways.
Recently watershed managers and researchers concerned with water yield,
both quantity and timing, have been studying the effects of snowfences
upon snow accumulation and storage. Tabler (1968) discusses fence
designs and other physical factors necessary to induce snow accumulation.
The amount of snow caught by fences depends primarily upon the size of
the contributing areas and the amount of snowfall (Berndt 1964). Other
important criteria for maximizing snow accumulation are fence location,
fence heights, and fence density. Schmidt (1970) reports on locating
snowfences in mountainous t~rrain. He describes the effects of
surrounding terrain on the type of snowdrift produced. Greb et al.
(1965) studied the effects of fence densities upon the depth and length
of snowdrifts. Fences, 50 feet long and 48 inches high, with 37, 58,
69, 79, and 85 percent air porosity were tested. The largest snow
deposit, in terms of volume, was found leeward of the fence with 79 percent air porosity. The fence with 58 percent air porosity produced the
largest deposit in terms of depth. Generally, the greater the air
porosity (up to 80 percent) of the fence, the longer and more evenly

�-250-

distributed the drift. Standard slat snowfence (58 percent air porosity) produced a drift approximately 40 feet long with maximum depth
of 3 feet (Greb et al. 1965). However, Berndt (1964) produced drifts
4.5 feet deep and 106 feet long with a standard slat snowfence. The
4.5 foot height was equal to the fence height. The drift height
seldom exceeds the fence height. Swank and Booth (1970) .produced
drifts 30 to 100 feet long and 5 feet deep with standard slat snowfences; drift length depended upon surrounding topography. Fences
should be mounted 6 to 12 inches above the ground to clear most low
growing vegetation. This minimizes upwind snow accumulation which
may damage the fence structurally or reduce efficiency (Swank and
Booth 1970).
Properly located snowfences are efficient in catching and storing
snow. Greb and Black (1971) calculated that drift producing storms
deposited 63 percent of the total snow and 71 percent of the total
precipitation per winter season, even though only 39 percent of the
snowfall events were accompanied by sufficient wind velocity to cause
drifting. Tabler (1971) increased peak snowpack-water equivalent on
a Ill-acre watershed by 70 percent with a single fence 12 1/2 feet high
and 1,300 feet long.
Increasing snow cover can affect soil moisture, and soil temperature,
which in turn affects the vegetational community beneath the snowdrift.
Willis and Haas (1970) report that snow cover reduces frost depth in
soil and with a snow cover of 3.5 to 4 feet, the soil will not freeze.
Cronin (1971) found that soil seldom freezes when the snow cover is 18
inches or more. Soil frost is important because snowpack runoff is
much greater when the soil is frozen (Willis et al. 1961). Also, soil
does not freeze nearly as quickly in the fall when it is wet. Induced
snowdrifts may be able to keep the soil from f rcez.Lng at all. This
would increase water absorption by the soil in the spring and the warmer
soil temperature may aid early spring plant growth. Cronin (1971)
studied larkspur growth under snowdrifts. He states that snowdrifts
may reduce the growing season, but beneath drifts, soil moisture is
abundant and soil temperature remains mild throughout the year. The
snow insulates both the soil and plants from harsh winter temperatures
which allows larkspur buds to grow and germination to occur beneath the
snow. Mathews and Conrad (1968) report that tall bluebell (Mertensia
arizonica) begins stem elongation beneath snow before melt starts,.but
heavy snowpack at higher elevations can delay growth for two weeks.
Greb (1970) studied grass production leeward of a 25 percent air
porosity snowfence near Akron, Colorado. He found the snowfence increased soil-water storage and production of Russian wildrye. Production was increased by 440 pounds per acre when compared with corresponding plots without a snowfence.
Collins (1970) is studying the effects of artificially increasing
snowpack upon native vegetation productivity in the Bridger Mountains
in Montana. Collins (1970) found that snowpack 4 and 9 feet deep delayed peak productivity of the vegetation.· Total production of all
vegetation was slightly less than control areas with only normal snow
depth; however, the normal snow accumulation in this area is approximately.

�-251-

1 1/2 to 2 feet.

Certain species, Agropyron subsecundum, Danthonia
intermedia, and Lupinus argenteus, did respond favorably to the increased snowpack by increasing production.

Planting

Forage Species Beneath

Snowdrifts

Numerous attempts have been made to improve wildlife habitat by
artificial revegetation on western rangelands.
Holmgren and Basile
(1959) recommend several procedures necessary for successful establishment of forage plantations.
Three common causes of failure are inadequate moisture during early growth, excessive grazing pressure before plants are established, and planting species not adaptable to the
sites. Establishing forage plantations beneath snowdrifts created by
snowfences should remedy two of these problems.
The snowdrift will
provide protection from grazing by deer and a source of moisture
during the early growing season to help establish the plants.
Bitterbrush (Purshia tridentata) has received more study than
any other species for forage revegetation, probably because bitterbrush
has long been considered a highly nutritious and preferred deer forage
plant and it has a wide range of site adaptability (Hubbard 1964).
Hubbard (1959, 1964) and Holmgren (1954, 1956) have done extensive
studies on planting bitterbrush seeds to improve big game ranges.
These publications discuss seed collection, site preparation and planting procedure, planting dates, protection of plantations, and management
for sustained production.
Hubbard (1964) recommends a planting depth of
1 to 1 1/2 inches. Ferguson (1962) had greater success in establishing
seedlings when 15 to 20 seeds were planted in a small mound of soil than
with only a single seed per mound.
Competition from grasses has a great
effect upon seedling survival.
Ferguson and Medin (1970) reported over
90 percent survival of seedlings when all vegetation had been removed
\ from the planting area before seeding, and zero percent survival where
no attempt was made to remove competition.
Sanderson et al. (1963)
found that grass competition reduced the amount of growth even for older
established bitterbrush plants.
Ferguson and Medin (1970) studied transplanting nursery grown bitterbrush seedlings in Idaho.
Best results were obtained by transplanting in
March before winter dormancy was broken and on scalped seed beds.
Exposing
bare roots to air for extended periods of time also reduced survival.
Highly fertile soil is not necessary for good bitterbrush survival
or growth.
In fact, higher survival and better growth of transplanted
seedlings was found on unfe~tilized plots than on fertilized plots
(Klemmedson and Ferguson 1969).
Many other species of shrubs have been planted to improve big
game winter ranges, but few studies have been conducted to determine
proper planting procedures and adaptability to range conditions.
Rabbitbrush (Chtysothamnus spp.) occurs frequently on western ranges
but has received very little consideration as a deer forage species or
for range rehabilitation and is often considered undesirable.
However,
Plummer (1968) states that many types of rabbitbrush are sought by all

�-252-

grazing animals and its high nutritional quality remains well balanced
throughout the winter. Two species of rabbitbrush(ChrYsothamnus
nauseosus and £. viscidiflorus) were chosen for planting in this study
because they both occur frequently in Middle Park, both grow rapidly
to considerable height, and personal observations indicate they are
heavily used on deer winter ranges.
Standard crested wheatgrass (Agropyron desertorum) is recommended
by Holmgren and Basile (1959) for range rehabilitation because it is
adaptable to the environmental conditions of many deer winter ranges
in the central Rocky Mountains and is desirable for winter and spring
use by deer. Crested wheatgrass exhibits one other advantage for
.
planting on winter game ranges; it begins growth very early in the
spring, often under snow (Knipe 1969), providing deer with new growth
during a critical period. Plummer (1968) rates establishment and
palatability of spring growth of crestedwheatgrass
as "very good" on
big sagebrush ranges, but presents no date to substantitate these ratings.
Yellow sweet clover (Melilotus officinalis) is a biennial forb
which grows well on disturbed sites and is considered palatable forage
for deer (Plummer 1968). Goom (1964) successfully established yellow
sweet clover on dry western ranges. He found it to be highly productive
and very compatible with crested wheatgrass.

Objectives
Removing snowdrifts from browse stands on deer winter range may
increase the amount of available deer habitat and also influence changes
in the plant community present on these areas. Newly created snowdrifts
have the potential of encouraging new forage supplies. The intention
of this study is to manipulate snowdrifts to increase available winter
forage and determine if changes occur in the plant communities or areas
where the snow cover has been altered.
The objectives are:
1.

To determine if changes occur in species composition and
density that are attributable to the elimination or creation
of snowdrifts.

2.

To determine if the growth of shrubs is influenced by the
"elimination or creation of snowdrifts.

3.

To evaluate the feasibility of establishing forage stands
beneath the created drift;; and on areas not covered by a
snowdrift.

4.

To determine if nitrogen fertilization aids in the establishment of such forage stands.

�-253-

5.

To determine the response of deer to the shrub stands
which'are freed of snow.

6.

To determine if soil moisture and temperature are influenced
by the amount of snow cover.
Methods

Sites and Snowfences
Twelve potential snowfence sit~s were located on Junction Butte
in March 1971. Aerial photographs and ground surveys were made of
these 12 sites, and from these data, five sites were selected for construction of snowfences. Four of these sites offer optimum snow storage
and adequate shrub stands. The fifth snowfenceis located on a steep,
sparsely vegetated hillside; only the forage restoration portion of the
study will be conducted on this site. Each of the fences is located so
its effects can be compared with a nearby unfenced area. Required fence
height for each site was calculated from: (a) topographic profiles of
the site, (b) weather station precipitation records from Kremmling and
Green Mountain Reservoir, and (c) estimated snow contribution zones.
Three of the sites require4-foot-high
and two require 8-foot-high fences
in accordance with snow storage characteristics of the site. The 4-foot
fences are 120, 140, and 160 feet long; both 8-foot fences are 140 feet
in length. The fences were constructed of standard slat snowfence (58
percent air porosity) suspended by 5/16-inch wire rope from wooden
posts spaced 20 feet apart. Alternate posts and end posts are braced
both down and upwind.
Three snowfence sites and controls will be used to evaluate the
treatment effects (drift or no drift) on native vegetation, sites 2, 3,
and 4 (see Appendix 1 for location). Each of these sites consists of
four separate areas or quadrants of nearly equal size: (a) area of
newly created drift, (b) area of original drift removed by fence, (c)
control area for quadrant A, without drifts, and (d) control area for
quadrant B with a drift (Appendix II). Dimensions of the quadrants
were determined by the length of the fence and estimated width of the preexisting drift. Quadrants are 65 X 120 feet at site 2 and 70 X 140 feet
at sites 3 and 4. Sites 3 and 4 are the 8-foot-high fences.
Measurements of Native Vegetation

Composition and Density
In order to determine changes in species composition and density,
40 permanent plots, 1 X 2 feet square, will be established in each quadrant
of the three sites. The plots are systematically distributed: eight transects placed 13 feet apart on site 2, and 15 feet apart on sites 3 and 4,
with five plots equally spaced along each transect. Spacing between plots
varies with the width of the estimated drift area: 10 feet apart in quadrants

�-254-

A and C, and 13 feet apart on quadrants B and D on site 1; and 12 feet
apart on quadrants A and C, and 15 feet apart on quadrants B and D on
sites 3 and 4 (Appendix 3). The distance from the outside transects
to the edge of the quadrant (fence ends) is 14.5 feet on site 2 and
17.5 feet on sites 3 and 4. Each plot is marked with two metal stakes
set in the ground. This design provides 480 total plots with 120 plots
in each quadrant.
.
Measurements on these plots will be made each year in late August
after most growth has occurred. The number of plants of each forb and
shrub species rooted within the plot will be recorded. Also, the
height of all shrubs rooted within a plot will be measured and crown
diameter of Artemisia plants recorded. Diameter will be measured to
the nearest centimeter on lines perpendicular and parallel to the plot.
The density of grasses is difficult to determine due to their growth
form. The numerous stems produced by bunch- and sod-forming grasses
make counting of individual stems nearly impossible. Some workers have
counted clumps of grass to determine density, but a clump is difficult
to define. To get a measure of change in density or species composition
of grasses, the 1 X2-foot plot was subdivided into 50 small plots 2.4
inches square. The number of squares in which each grass species occurs
will be counted. The density of grasses cannot be determined by this
method because the number of stems or separate plants was not counted.
But, any increase or decrease in the number of squares containing a grass
species can be determined.

Analysis of Density Data
Changes in plant density will be compared by means of a multivariate
analysis of variance with covariance, using one year pretreatment and 3
years posttreatment data. The more common shrub and forb species will
be considered ·individually. The same type of analysis will be done for
all grass species combined, but a separate program will be used because
of the different magnitude of numbers. This analysis will determine the
average density of each forb and shrub species for each quadrant and the
overall average of the three replicate quadrants for each year the measurements are taken. Comparison will be made between pretreatment and each
posttreatment year and between the control and treatment quadrants to
determine if changes in density are occurring and if the changes are due
to treatments.

Growth and Use on Native Shrubs
Five of the most common species of shrubs occurring on the study
sites, Amelanchier alnifolia,Symphoricatpos
oreophilus,Purshiatridentata,
Ribes cereum and Prurtus virginianus, will be measured each spring and fall
to determine if growth and deer use of these species is influenced by the
amount of snow cover. Fifteen plants of Amelanchier andSymphoricarpos
will be measured in each quadrants on sites 2, 3 and 4, except in three
cases, quadrant C sites 2 and 4, and quadrant A site 3 did not contain 15

�-255-

Ame1anchier plants; all Ame1anchier plants on these quadrants will be
measured (see Table 1 for numbers). Plants will be selected in the
following manner: 15 permanent density plots were randomly chosen from
the 40 in each quadrant.· For each· apecLes, the plant closest to each
of these points is selected for measurement. The three other species
to be measured occurred infrequently on the study area, and all plants
of these species were measured wherever they were found on a quadrant.
On each plant, two live branches less than 5 feet above the ground
will be tagged on 2-year-01d stems which had at least two current-annua1growth leaders, except that only one branch per plant was tagged on
Symphoricarpos oreophi1us plants and on very small plants of the other
species. The measurement at time of establishment (sunnner1970) resulted
in sample sizes of plants and branches as shown in Table 1. Plants are
the first number in the table and the number of branches is in parenthesis.
In September, after most growth has occurred, all current-annualgrowth on eaCh tagged branch will be measured. The length of each leader
or twig will be measured from the terminal end of the newest bud scale
scar to the end of the terminal bud to the nearest centimeter. Each twig
or leader measured will be marked with a dab of orange paint on the bud
scale scar. In April, the tagged branches will be examined to obtain a
record of use. No measurements will be taken; use on each branch will be
recorded as present or absent. This will show the percentage of branches
upon which some degree of use occurred. Each September, the new current
annual growth on the tagged branches will be measured to see if shrub
growth is responding to the treatment.
Analysis of Shrub Growth Data
Comparisons of the average amount of current annual growth per
leader will be made between pretreatment measurements and each year's
posttreatment measurements for each species on each quadrant and quadrant
type. The same plants of each species will be measured each year so the
sample size will not change within a quadrant, but comparisons between
different quadrant types will have unequal sample sizes. An analysis of
variance will be used to determine the average current annual growth per
leader for each species measured and compare treatment and control areas.

Forage Restoration
Two sites, 1 and 5, will be used for forage restoration studies.
Seed beds, each 30 X 30 feet in size, have been established behind the
fence and on the adjacent control area (see Appendix 1). The lower site
(1), with four seed beds behind the fence and four on the control, is
located on a relatively level area with deep soil. The upper site (5)
has 12 seed beds, eight behind the fence and four in the control area.
Site 5 is located on a steep hillside with shallow, rocky soil. In the
sunnnerof 1970, alternate seed beds on each site were: (a) cleared of
all vegetation to eliminate competition, and (b) furrowed every 2.4 feet
to provide loose soil for a seed bed, but not cleared of competing native
vegetation (see Appendix IV).

�TABLE 1.

Site

Site

2

Site

3

Quadrant

Quadrant

Quadrant

4

Species

A

B

C

D

A

B

C

D

A

B

C

D

Ama1

15 (23)

15 (30)

7(11)

15 (28)

14 (23)

15 (27)

15(15)

15 (26)

15(23)

15(29)

12 (22)

15 (30)

Syor
Prvi

15 (15) ,

15(15)

I

15(15)

15(15)

15(15)

Rice

2 ( 4)

Putr

1 ( 2)

5(10)

15(15) .

15(15)

15(15)

15(15)

15(15)

15 (15)

I

2 ( 2)

2 ( 3)

5(10)

15(15)

I'.J
VI
0\

1 ( 2)
6(11)

1 ( 2)

5 (10)

1 ( 2)

5 ( 9)

1 ( 2)

6(11)

4 ( 8)

�-257-

Five species, Purshia tridentata, Chrysothamrtus nauseosus,
Chrysothamnus viscidif10rus,Agropyron
desertorum, and Me1i10tus
officina1e, were planted in each seed bed in rows 2.4 feet apart.
The planting was done in October 1970. Two rows of each species
were planted per bed except Purshia which has two rows of seedlings
and two rows of seeds planted per bed. ThePurshia seedings, about
6 inches tall, were grown at the U. S. Forest Service Mt. Sopris Nursery
near Carbondale, Colorado, from seeds collected near Maybell, Colorado.
Each seedling was placed by hand into a hole about 6 inches deep. The
soil was placed firmly around the roots and about 1 quart of water was
added to the soil immediately. The seedlings were watered again 3 days
later. Twenty seedlings were transplanted in each row, about 18 inches
apart. Purshia seeds were obtained from the same source as the seedlings.
The seeds were planted 18 inches apart in the rows. A small mound of
loose soil was formed and 12 seeds placed about 3/4-inch into the mound.
The Chrysothamrtus seeds were collected from mature plants near the study
area. Approximately 15 seeds were planted every 6 inches along the row
and covered with about 1 inch of soil.
;

Agropyron desertorum and Me1i10tus officina1e seeds were obtained
commercially. Planting was accomplished by spreading the seeds along
the rows at a rate of about 20 seeds per inch. The seeds were covered
by raking soil over the row. Half of the seed beds on each site were
fertilized in November with ammonium nitrate at the rate of 120 pounds
per acre of actual nitrogen.
The six kinds of plants, each duplicated, resulted in 12 rows per
seed bed which were planted in the order indicated in Appendix V.
The success of p1antings in the various seed bed treatments will be
evaluated by the following methods:
1.

Number of surviving Purshia in each seed bed.

2.

Height of all surviving Purshia seeding transplants will be
measured to the nearest centimeter. Measurements will be from
the ground at the base of the plant to the highest point of the
plant in its natural growth form (the plant will not be
straightened to increase height). These measurements will be
made each spring and fall.

3.

The number of Purshia plants which emerge from each seed mound
will be counted in the spring of 1972. The number of successful
mounds will be counted in the fall of 1972 and the height measured
as above. A successful mound is one which has at least one surviving Purshia plant. Height measurements will be repeated each
spring and fall to get an estimate of plant growth.

4.

The number of Chrysothamnus· seedlings established per row of
each seed bed will be counted and the height measured each fall
and spring, as above.

�-258-

5.

Counts of the number of stems per row of Agropyron and
Melilotus plants will be made each spring and fall. The
average height of these plants will be determined by
measuring the height of the nearest plant to every I-foot
mark along the row.

Germination rate of the Purshia and Chrysothamnus seeds used for
planting will be determined using the procedure outlined by Association
of Official Seed Analysts (1965). Two hundred seeds of each species
will be tested. Dormancy of Purshia seeds will be broken by soaking
the seeds in a 3 percent solution of thiourea for 5 minutes.
Measurement of Deer Occupancy
Use by deer of the areas behind each fence and adjacent control
will be estimated on all sites except number 5. The number of deer
tracks crossing three line transects in each quadrant will be counted
biweekly from January to March. On each site, three transects, perpendicular to the fence, run continuously through quadrants A and B. Three
other transects run continuously through quadrants C and D. The location
and length of all transects is shown in Appendix 6.
The sum of tracks crossing on the three transects in each quadrant
at each 2-week sampling period will be the basic sampling units used to
compare deer use on drift and nondrift areas. These measurements cannot
be used to estimate the number of deer using these areas. Rather, they
will indicate the amount of time spent on the area by one or several deer.
A deer which simply moves across the area without stopping to graze would
be counted only once on each transect; but a deer which stays on the area
to eat will very likely move around and cross the transect lines many
times, and thus represent several track crossings.
A hypothetical sample of the expected data is shown below:
Site 1

Fence
Drift
Date

Control

Nondrift

Nondrift

Drift

Sum of Track Crossings on all Transects

Jan. 13-27

1

3

6

0

Jan. 28 - Feb. 10

0

18

12

0

Feb. 11-24

3

24

27

3

Feb. 25 - March 9

6

17

15

8

0

32

27

6

c

March 10-24

Sites 2, 3, and 4 will provide similar data.

�-259-

An analysis of variance will be used to determine if a significant difference in the number of track crossings occur. Major components of the analysis will be treatments (fence or\ no fence), sites
(4), dates (5), and type (drift or no drift).
Deer occupancy will also be recorded by a time lapse movie camera
which takes a photo every 7 minutes during daylight hours. The camera
will be positioned to simultaneously photograph all quadrants on sites
3 and 4. Approximately 6,000 frames will be exposed from January
through March. Each frame will be considered one sampling unit. All
deer in each quadrant will be counted, and the quadrants on each of the
two sites summed for analysis. An analysis of variance will be done to
determine if a significant number of deer spend more time in one quadrant
than another.
Snow depth on each quadrant will be measured in conjunction with
track counts. A snow depth course has been established at the center of
and perpendicular to each fence and control line. Along these lines,
snow depth will be measured at various intervals depending upon the topographic profiles of the land. A steel post marked in I-foot increments
is set at each measurement point. Track counts in each quadrant may be
related to the snow depth in each quadrant. A simple linear regression
will be run to correlate average snow depth with track counts in each
quadrant at each measurement date.
Measurements of Soil Moisture and Temperature
Soil temperature at 3-inch depth will be measured on all sites to
determine if snow cover affects soil temperature during the growing
season. Sites 2, 3, and 4 each have three thermistors located in each
quadrant (see Appendix 3 for location). Four seed beds in each planting
site have three thermistors each (see Appendix 5 for location). Soil
temperatures at these permanent locations will be measured every two
weeks from April to October. The mean temperatures in each quadrant or
seed bed will be compared to determine if differences occur. In the
spring after maximum snow accumulation, the exact snow depth at each soil
temperature measurement station will be measured. Soil temperature beneath the snowpack will be taken at this time and again as soon as all
snow has melted. A simple linear regression will be run to correlate snow
depth with soil temperature at these times. The thermistors will be
located in relation to anticipated snow profile, so the snow depth would
vary above each measurement point.
Soil moisture from surface to 8-inch depth will be measured at one
central point in each quadrant of sites 2, 3, and 4 monthly from May 1
to September 1. Monthly measurements will also be taken from each of the
eight seed beds on the lower planting site. No soil moisture measurements
will be taken on site 5 because of the extremely rocky conditions prevalent
on this area. The gravimetric method of determining soil moisture will
be used, so the exact measurement point will be selected to provide an
area of deep soil and no rocks if possible.
Soil moisture will be compared between quadrants and related to the
average snow depth of each quadrant and snow depth above the measurement

�-260-

point. Average snow depth will be determined by extensively surveying
each site in the spring after maximum snow accumulation.
Plan t Phenology
Observations on the phenology of the 10 most common species of plants
will be made biweekly during the growing season in each quadrant on sites
2, 3, and 4 to note if differences in plant development occur due to the
amount of snow cover. Twenty plants of each species will be examined in
each quadrant at each observation period. Observations will be made along
the three transects established previously for track count information
(see Appendix 6). Along each transect line, six points will be established
on the outside lines and eight points along the center line. At each of
these points, one plant of each species near the point will be examined to
determine its developmental stage. The same plants will not be examined
at each observation period (although it may happen with some species), but
by examining 20 plants each observation period, the time sequence of phasic
development will be established for these 10 species in each quadrant.
Weather Information
A separate study conducted by Proj~ct 1601 (Snow and Avalanche Control)
of the Rocky Mountain Forest and Range Experiment Station will evaluate the
design and location of the snowfence structures in relation to snow accumulation. Information on wind speed, wind direction, and temperature will be
recorded continuously from two sites on Junction Butte. A recording gauge
for blowing snow (Tabler 1971) will measure the time and extent of drifting
events (see Appendix for locations of weather instruments). Precipitation
records will be obtained from weather stations located at Kremmling and
Green Mountain Reservoir. Details of this study are reported by Schmidt
(1971) •
Presentation of Results
Sufficient data for three major publications should be available by
1974. Two publications will be on the effects of snow cover on native vegetation and on planting forage plants beneath snowdrifts. The Journal of
Wildlife Management or Journal of Range Management should be suitable outlets. Another publication, with Research Wor~ Unit 1601, will report on the
effectiveness of collecting snow with snowfences on sagebrush lands.
An annual progress report and preliminary data will be furnished to the
Colorado Division of Wildlife each May.
Personnel Assignments
The following personnel will be involved with this study: o. C. Wa1lmo,
Project Leader 1801. Provides supervision of all phases of work. Estimated
time required, 6 weeks per year. Wayne L. Regel in , Biological Technician. .
Principal investigator response for all budgeting, planning~ data collection,
data analysis, and publications. Estimated time requirements, 50 weeks per
year. Donald W. Reichert, Range Technician. Will help with data collection.
Estimated time required, 3 weeks per year.

�-261-

Literature Review

Association of Official Seed Analysts. 1965. Rules for testing
seeds. Proc. Assoc. Off. Seed Anal. 54 (2). 112 p ,
Berndt, N. W. 1964. Inducing snow accumulation on mountain
grassland watersheds. J. Soil and Water Conserve 19: 196-198.
Collins, D. D. 1971. The Bridger Site, 1970 Progress Report.
Tech. Rep. No. 84. Grassl. Biome, U. S. Int. BioI. Program,
Fort Collins, Colo. 40 p.
Cronin, E. H. 1971. Tall larkspur: Some reasons for its continued
pre-eminence as a poisonous plant. J. Range Manage. 24(4): 258-263.
Ferguson, R. B. 1962. Growth of single bitterbrush plants vs.
multiple groups established by direct seeding. U. S. Forest Serv.,
Intermt. Forest and Range Exp. Sta. Res. Note 90, 2 p.
Ferguson, R. B. and D. E. Medin. 1970. Rehabilitation and management
of deer winter range. Job 1. Site potential studies. Job 2.
Bitterbrush seedling survival studies. Job Completion Rep.' WIIIR15, Idaho Fish and Game Dep., Boise, Idaho.
Greb, B. W. 1970. Cool season grass production leeward of a 28%
wood-slat snowfence. Colo. Agr. Exp. Sta. Prog. Rep. No. 70-24,
3 p.

Greb, B. W., R. H. Mickelson, and G. O. Hinge. 1965. Conservation
research at the Central Great Plains Field Station, Northern Plains
Branch, Soil and Conserve Res., Div. Agr. Res. Serv., USDA Misc.
Leafl., 16 p .
Greb, B. W. and A. L. Black. 1971. Vegetative barriers and artificial
fences for managing snow in the central and northern plains. p. 96111. In Snow and ice in relation to wildlife and recreation symposium.
Iowa State Univ., Ames, Iowa, 280 p.
Goom, F. B. 1964. A comparison of two sweetclover strains and Lodak
alfalfa alone and in mixtures with crested wheatgrass for range
and dryland seeding. J. Range Manage. 17(1): 19-23.
Holmgren, R. C. 1954. A comparison of browse species for big-game
winter ranges in southwestern Idaho. Intermt. Forest and Range
Exp. Sta. Res. Pap. 33, 12 p.
Holmgren, R. C. 1956. Competition between annuals and young bitterbrush (Purshia tridentata) in Idaho. Ecology 37: 370-377.
Holmgren, R. C. and J. U. Basile. 1959. Improving southern Idaho
deer winter ranges by artificial revegetation. Wi1d1. Bull.
No.3, Idaho Dep. of Fish and Game, 61 p.

�-262-

Hubbard, R. L., E. C. Nord, and L. L. Brown. 1959. Bitterbrush
reseeding •••
a tool for the game range manager. U. S. Forest
Serv., Pac. Southwest Forest and Range Exp. Sta. Misc. Pap. 39,
14 p.
Hubbard, R. L. 1964. A guide to bitterbrush seeding in California.
U. S. Forest Serv., Pac. Southwest Forest and Range Exp. Sta.
and Calif. Dep. Fish and Game Resour. Agency, 30 p.
Klemmedson, J. o. and R. B. Ferguson. 1969. Response of bitterbrush
seedlings to nitrogen and moisture on a granitic soil. Soil
Sci. Soc. Amer. Proc. 33 (6): 962-966.
Knipe, O. D. 1969. Establishment and survival of several grasses
in the sagebrush type, west-central N~w Mexico. U. S. Forest
Serv., Rocky Mt. Forest and Range Exp. Sta. Res. Note RM-149, 2 p.
Mathews, V. B. and P. W. Conrad. 1968. An ecological life history
of tall bluebell (Mertensia arizonica var. Leonardi) in Utah.
Ecology 49(6): 1113-1123.
Plummer, A. P. 1968. Restoring big-game range in Utah.
Fish and Game Publ. No. 68-3, 183 p ,

Utah Div.

Sanderson, H. R., R. L. Hubbard, and D. W. Seegrist. 1963. Effects
of grass competition on bitterbrush: second-year report. U. S.
Forest Serv., Pac. Southwest Forest and Range Exp. Sta. Res.
Note PSW-26, 7 p.
Schmidt, R. A. 1970. Locating snowfencesin mountainous terrain.
p. 220-225. In Snow removal and ice control research. Highw.
Res. Bd. Spec-.-Rep. 115, 282 p. Dartmouth ColI., Hanov~r, N. H.
Swank, G. W. and R. W. Booth. 1970. Snowfencing to redistribute
snow accumulation. J. Soil and Water Conserve 25: 197-198.
Tabler, R. D. 1968. Physical and economic design criteria for
induced snow accumulation projects. Water Resour. Res. 4(3):
513-519.
Tabler, R. D. 1971. Design of a watershed snowfence system and
first year snow accumulation. Proc. 39th Annu. West. Snow Conf.,
Billings, Mon t•
Tabler, R. D. 1971. A recording gage for blOWing snow. U. S. Forest
Serv., Rocky Mt. Forest and Range Exp. Sta. Res. Note RM-193, 7 p.
Wallmo, O. C., and R. B. Gill. 1971. Snow, winter distribution, and
population dynamics of mule deer in the Central Rocky Mountains.
p. 1-15. In Snow and ice in relation to wildlife and recreation
symposium.--Iowa State Univ., Ames, Iowa. 280 p.
Willis, W.O., and H. J. Haas. 1970. Snow and snowmelt management will
level benches, small grain stubble, and windbreaks. p. 86-95. In
Snow and ice in relation to wildlife and recreation symposium. Iowa
State Univ., Ames, Iowa, 280 p.

�-263-

Section 21

/

(j

Legend:
. Contour Interval
160 feet
Loca ti on: T 1 a, R 60 ~j, S 21,22,27,20

+Section
corner
--Sectlon lin~s
~ • Tri.angu1ation station
~Aban~ne"d
mine
-

-

-

-

-

-

-

-.:,.

-

•••.•-:

__ 'oJ __

.::;

&gt;

•

'~8::x 3~~~drift ceter
~ Si~es 1, 3, 4, 5 Pence a~d
'\.
"\'C'ite'"
..
'."
,-,''''17
~.,c,.
...J.
'-..••_,..•••
'-"',='--C':'
-~~t~~:: c~ntrJ1 o~lJ·
• "An6moma

v __

'.I

�-264-

Fence,

Control
C
D

Fence:

Oontrol

o

A. Quadrant

A-

11e,-;sno,\'1drift created

by snovrf enc e

B Quadra.nt 13- Area where s:::.'.):·:i=ift rc:.:.c·.c-.:'~ -:

o ~uadrant

0- Control

. D Quad:=ant D- Contrel

Appendix

II'

area

without

sno-;-;drift

Area wi th sno~';drift

snowfence

�I.,'Jyuutof density

•

•
•

•15
•

)G'

e

•

15

•

•
15·

• •
•

•

I.luadrant D

•

•

x

plots for Sites 3 and 4

• •5
• •4

•
•

Plot No.
5

• • •3

• • 15 •
•
x
4 •
• • • • • • 15 •
Quadra.nt B
3 •
• • • • • • 15 •

4

4

-

.;

- .

•

•

It

2

(

•

,

• •.15 • • • • • x • 2
.
•
•
• • •
• •
• 1
15
._5
• • 12 • • • • •

• • 12
• • • •.
•
Quadrant 0
x
• •• 12 • • x•• • • • 3
• • • • • • • •.. 2
,
~ 12
• • 10 • • • • x • •:,·1

•

•

1
5

•

•

•

•

•

•

•
••
•

•

• •
Quadrant A
• • •
3 •
2 •
• • ",
1

•

•

•

x~

,

t·

• •

• •
• •

I

•
15

x

•

•

•

• 12 •

•

•

•

•

•

•

•

,

•

•10 •

• x
•

•

,.

•

15
...:---

x • Location' of thermistors

Appendix III

I

•
12
12

,

12

12
15
12
12
15
15
15
15
15
12
12 19·
t17.~5 15
15
5. 15
6
4
7 8
2
5
3
4
5
0
7
8 Transe ,ts 1
2
3
1
,
Fence Line 140 ft.
Control line 120 ft.
I

I

N
0\
VI

•

•
f

�Layout

of density

plpto on oite 2

plot No.

• • • •
• • D•
• Quadrant
• • • •
• • • •

•
15ft. Distance between plot3
4 •
• • plot:.;Uo.
15
5 •
3 •
•• • • x • • •
•
• • •
x
13
15
4 •
2 •
• • • • • 13 •
• ;~~uad1'an·~B
• • •
15
3 •
• • • • • • •
1.
•
• • • • • 15__•
13
2 •.
• • • • • • •
5'
•
13
• • • • • 15 • 1 • •
•
• • • '.
---13 ._--4'
•
• • •
5 •
•
•
•
•
•
•
•
15
1Q
3 •
•
• ,
•
•
•
• • It. • • • •
• Quadrant
•
~U2.&lt;1r8nt C
10
15
5 •

•

•

x

~

x

x

x

-f

t

.

.

t

x

x

4

x

2 •

•

•

•

•

~

•

•

• •
DiG t~;
nee

. 15

, 1:;)

1

2

• •

o

J

15,

15

15,

4

)

t

•

iJ

I

x

•

•

1

o on :~J.'fJl 1ino 1"-5 ft.
Oontro1 ;~rco. li3 n~)(r..,1-;~ ~~ooyazd o
north i'10;:-d; of ~rn()lifcnco
:::,1'0(-1

•

13

cont.

,

•

•

•

•

• x

10

•lQ •

•

,.

-,..

7.

L

J

]'(;:;'lOU

LLne

x • Location of thermistors

Ap,en41x III

•

. . . . . . . .
•

2 •

•

•

DiG t,~_l1ee be t'-;:f-0l1 "tft-ns ef.ts l,:!
15
lS.i?:G.
L 15 ( 1;.&gt; _•.._I.?__
• }~-+__
,.2_L :'__J_._.....!__ :1:.?--L7
aTrahGt3e lis 1
2
:5
tt
5
6
7 U

bo tn.e en t r.: risco ts
I

3

15

x

1

I

N
0'1
0'1
I

l::-~O 1'-1;.

-__

-.

�-267-

Layout of forage restoration sites

(Sites 1 and 5)

Site 1 (Lower site)

.

30 ft
C "4

F 4

P 3
Pert.

It

Pert.

"

c 3

30 f

Control

P 2 ~

II
7t

l&lt;

5 feet " between beds

----------

Il

/I.

I

140 feet

C 1
F 1 It
~
Fert. Pert.
5 feet between beds
C 2

Fence

140 feet

.

30 ft

.c

F 5
Fert.

C 6

30ft

C

4

c 2

F 3
&amp;

/

Fer't.

)IF 1~
Pert.
~

II,

"

.I(

F 6

C 5

F 4

Fert.

C
1-

5 ft.

3

&gt;&lt;

C 1
l!"'ert. iert.1l
F 2

1{

5 feet between beds

'I

---~I·============~
Control 70 feet
Fence 160 feet
L~gend:
C- Cleared Seedbed
F - Furrowed seed beds
Fert.-Fert1lized
,X

plot

L'ocat10n of ther.n1stors

Append1x

IV

�-268-

Track count transects

Site 2

Transects

Transects

123

1

2

65ft.

65ft.
QuadlQ..nt:a

-I'-

-

65ft.
Quad ant C

25

J--

30

30

-

5
55
ti5
Control line 110 feet

Appendix VIcont.

3

-!---

Quad ant.Ai

25
--I

•

20

40

40

20
60
100
Fence line 120 feet

20

�-269-

Track count transects

Sites 1, 3j and 4

Transect

Transect
1

2

1

3

2

Quad ant D

Quadr nt B

68ft.

68ft.

-- -""-

-

68ft.

68ft ..

Quadra t A

Quad ant C .

-

20ft

40ft.

20ft.

3

40ft.
60ft.

20ft

-

100ft.

•

N

)S

40ft.

40ft.

·30ft.
I

30ft.
Fence

Control Line 120 feet

Appendix VI

30ft.

110ft.
140 feet long

�-270-

Row Sequence in all seedbeds
12
&amp;",

11
~

'

...

10

-,

9

8

6,

7

5

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ROrl Numbers

1 &amp; 7· Helo1i tus officinalis
2 &amp; &lt;3 Pursl1ia tridentata seeds
3 &amp; 9 Chrysothamnus visciciflorus
4 &amp; 10 Agropyron desertor~
5 &amp; 11 Purshia tridentata seedlings
6 &amp; 12 Chrysothamnus nauseosus
)

AppendL.;:v

S 22'E

�-271-

APPENDIX

B

��-273-

Wildlife Habitat--Central Rockies
rS-lU-l-lSQl

.Study Plan S
Amendment No. 1
The Use of Carbon Black to Produce Mid~Winter Snow Melt
on Deer Winter Range in Colorado

Rocky Mountain Forest and Range Experiment Station
Fort Collins, Colorado
in cooperation with
·Division of Wildlife
Colorado Department of Natural Resources

4210

��-275-

STUDY PLAN 8
Amendment No. 1
The Use of Carbon Black to Produce Mid-Winter Snow Melt
on Deer Winter Range in Colorado
by
Wayne L. Regelin

The Problem
Study Plan 8 addresses the problem of strategic placement of snowfences on deer winter range to substantially reduce the amount of windblown snow which accumulates in shrub stands to permit their use by
deer (Regelin 1972). However, snow which falls directly upon the shrub
stands can accumulate to depths which may prohibit deer occupancy of
these areas. The feasibility of using carbon black to produce mid-winter
snowmelt in shrub stands protected by snowfences or natural windbreaks
will be tested in this amendment to Study Plan 8.

Literature Review
Artificial darkening of the snow surface reduces its reflective
power, resulting in an increase of absorbed short-wave radiation which
increases the rate of snow melt. Procedures based on this principle
have been used for many years in Russia and Japan to accelerate snow
melt on airport landing strips and to increase runoff from glaciers.
Slaughter (1966) cites numerous examples of such practices.
Numerous substances, including soil, coal dust, cinders, sand, and
salt, have been used to darken the surface of snow. Arnold (1960)
studied seven types of materials in Canada. The most effective material
was fine cinders (0.2 to 2.0 mm particles), applied at the rate of 1,000
2
g/m , which increased total ablation by 5 times over control plots.
Eight surface additives were tested near Fraser, Colorado, during the
spring of 1967 (Megahan 1968). Carbon black treatments were much more
effective than other black materials. Melt rates on treated snow exceeded the melt rate on untreated snow by 2.5 times.
Storik (1960) studied the effect of particle size of surface additives
upon melt rate. He discovered that the radiation absorptivity of the
material depended upon degree of subdivision of the material as much as
upon color. He concluded it is more practical to use small quantities
of highly dispersible substances rather than large quantities of substances with low dispersity. Williams and Gold (1963), working in Canada,
reached the same conclusion and indicate "dusts" with average particle
size between 0.1 and 2.5 mm are practical. Megahan (1968) discovered
that the melt rate tends to increase as mean particle size decreases and
tends to approach a limit as the rate of application increases. The most
effective melt rate was obtained with 3.2 g/ft2 (300 lb/acre) of an aqueous
suspension of carbon black.

�-276-

To date, most snow melt experiments have been done in spring and
early summer when temperatures are normally above freezing, but Arnold
(1960) found accelerated melting still occurred at temperatures below
-100 C.
Taketa and Murakami (1956) report accelerated melt at temperatures below 00 C, even when surface additives were covered by 20 cm
of new snow. However, new snowfall covering the surface additive does
reduce its effectiveness. Azuma (1956) found a 55 percent loss in radiation absorption due to 1 cm of new snow c~ver. Arnold (1960) reports
radiation can penetrate up to 15 cm of snow. Megahan (1968) reports
effective accelerated melt on plots covered with 2 inches of new snow
overlying the dusted surface.
In Middle Park, Colorado (Gilbert 1959), 18 inches of snow was melted
in 72 hours in early spring by spreading hardwood charcoal on the snow
surface. Megahan (1968) found that applying carbon black at the rate of
300 1b/acre increased the melt rate by 2.8 times over untreated plots on
the first day after application, but the melt rate diminished thereafter.
Shu1'ts (1963) found the effectiveness of all surface darkening agents
was short-1ived,lasting
only 6 or 7 days.

Objectives
The objectives of this study are:
1. To determine the effectiveness of carbon black in causing midwinter snow melt on deer winter range.
2. To determine if changes occur in density and/or crusting characteristics of the snow due to treatment with carbon black.

Methods
Seven experimental snow melt sites will be selected on Junction Butte,
Grand County, Colorado (see Study Plan 8). All sites will be protected
from wind-blown snow accumulation either by snowfences or natural windbreaks provided by dense stands of· serviceberry and other shrubs. Each
of the seven sites consists of a treatment plot paired with an adjacent
control plot. Plots will be paired on the basis of similar aspect,
slope, and vegetative cover. Each plot is 10 msquare with 3m separating
the paired plots. All sites will be located on nearly level terrain, with
four sites on a slight northerly aspect and three sites on a southerly
aspect. General slope and aspect will be measured with compass and level.
Carbon black {Phi1b1ackN550}!l will be applied using a "whirlybird"
backpack fertilizer spreader at the rate of 336 Kg/ha (3.2 g/ft2). The
first application of carbon black will be in late January, 1974. Subsequent applications will be made if new snowfall covers the carbon black.

!/ Purchased from Philips Petroleum Co.

Trade names and company names are
used for the benefit of the reader and do not imply endorsement or preferential treatment by the U. S. Department of Agriculture.

�-277-

Thirteen snow ablation measurement stakes will be systematically
located within each treatment and control plot. The stakes will be
marked at 3-inch intervals so snow depth can be measured without walking
through the plots. Appendix 1 illustrates the placement of the stakes.
Snow depth at each stake will be recorded daily, during late afternoon,
throughout the duration of the study.
Snow density will be determined using a Montrose Snow Sampler and the
hardness of snow crust by using a ram penetrometer. These measurements
will be made at five locations around the periphery of each treatment
and control plot (Appendix 1). Snow density and ram resistance will be
measured on all plots before the first treatment is applied, before any
follow-up applications, and at the termination of the study.
Air temperature, wind speed, and wind direction will be recorded
continuously throughout the study by an anemometer and thermometer located near the plots. Notes on cloud cover and snowfall will be made
daily.

Analysis and Interpretation of Results
Data on snow melt rate will be analyzed by analysis of variance to
determine if treatment effect (carbon black) increases snow melt and if
daily variations in melt rate occur. The hypothesis will be that treat-,
ment has no influence upon melt rate. The numbers used in the analysis
will be the mean snow depths on plots obtained from the 13 ablation
stakes. An example of the analysis of variance table is shown below.
Source of Variation

Degrees of Freedom

Treatments (T)

1

Replications

6

(R)

T X R

6

Dates (D)

15

T X D

15

DXR

90

T X D X R

90

Total

223

The error term to test treatment effect will be the T X R,interaction
and the error term to test date and date X treatment effect will be the
T X D X R interaction.
The snow melt data will also be presented graphically to illustrate
melt rate differences between treatments and controls as influenced by
aspect and temperature. Data on snow density and ram resistance will be
handled by a similar analysis of variance.

�-278-

Application of Results
If the results are promising, a management-scale pilot test will be
recommended to the contractor, Colorado Division of Wildlife, for further
funding and extension of the study.

Publication
Results will be published in one technical journal article and one
popular journal such as Colorado Outdoors.

Scheduling
Layout of plots - Fall, 1973
Application of treatments - January-February, 1974
Measurements - January-March, 1974
Completion date - June, 1975. Field work will be completed in
Spring, 1974, analysis of data and preparation of final
manuscripts will be completed by June, 1975.

Responsibility
W. L. Regelin, investigation leader
D. W. Reichert, field assistance
O. C. Wallmo, consultation and advice

Estimated Costs
Man-days
Field work

20

Compiling and analyzing data

6

~

Computer services

$ 10

Equipment and supplies

$120

Mileage and per diem

$240

11 Costs are covered by Cooperative Agreement No. 16-216 with Colorado
Division of Wildlife.

�-279-

Literature Cited

Arnold" K. C. 1960. An investigation into methods of accelerating the
melting of ice and snow by artificial dusts. In Geology of the
Artie. Proc ,, 1st Int.Symp. on Artie Geol., Vol. 2. Univ. Toronto
Press, Toronto, Ontario.
Azuma, S. 1956. Micrometeorological observations on natural and artificially blackened snow surfaces. Science Report. Saikyo Univ.
(Japan). Nat. Sci. and Living Sci., Vol. 2. SIP 16051.
Gilbert, P. F. 1959. Experimental snow melting on key deer winter
ranges. Colo. Dep. Game and Fish, P-R Proj. Rep. W-38-R-12, 4 p.
Multilith.
Megahan, W. F. 1968. Increasing snow melt rate by surface additives.
Ph.D. Diss., Colo. State Univ., Fort Collins, Colo. 175 p.
Regelin, W. L. 1972. Junction Butte wildlife habitat improvement
project. 2nd Annu. Progr. Rep., Rocky Mt. For and Range Exp. Stn.,
31 p , Mimeo.
Shul'ts, V. L. 1963. On the question of the expediency of artificial
intensification of the melting of snow in the mountains of Central
Asia. Soviet Hydrology: Selected Papers, Vol. 2, No.3, p. 275-278.
Slaughter, C. W. 1966. Snow albedo modification, a review of literature.
CRREL. TR 217, 25 p.
Storik, I. L. 1960. Utilization of certain substances for acceleration
of the melting of ice. Transl. by V. N. Pavloff. Natl. Res. Counc.
of Canada, Tech. Trans!. 1067, 13 p.
Taketa, M., and M.Murakami.
1956. Jinko uysetsu ni tsuite (On artificial
snow melting). Seppyo, Vol. 17, No.3, p. 21-27. Sip 15660.
Williams, G. P., and L. W. Gold. 1963. The use of dust to advance the
breakup of ice on lakes and rivers. Natl. Res. Counc. of Canada,
Div. of Build. Res., Tech. Pap. 165 (NRC 7725), 60 p.

�-280Treatment

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. Appendix 1.

Layout of snow melt site.

prats (indicated
and Ramresistance

by dots).

Location of snow ablation

X indicates

measurements.

sample points

szakes wi~hin

for snow density

�-281-

APPENDIX

C

��-283-

Second Annual Progress Report
for

Junction Butte Wildlife Habitat Improvement Project
May 1, 1972 to April 30, 1973

in conformance with
Cooperative Agreement No. 16-216

between
Division of Wildlife
Colorado Department of Natural Resources

md
Rocky Mountain Forest md

Rmge

Experiment Station

USDA Forest Service

��-285-

Accomplishments

Fence Designs
During the summer 1972, eight snowfences of four different economical designs were constructed on the north side of Junction Butte. All
four fence designs were constructed of standard slat snowfence suspended
by a single strand of No. 9 wire from steel fence posts. All fences are
4 feet high and vary in length from 90 to 100 feet depending upon post
spacing. Figure 1 illustrates the four fence designs. Cost of the four
designs varied from $1.16 to $ .99 per linear foot including labor costs,
and from $ .79 to $ .64 per linear foot excluding labor costs. This is
much less than the long-life fences constructed in 1971 which cost $7.30
per linear foot for 8-ft. high fences and $4.53 per linear foot for 4-ft.
high fences. The fences constructed in 1971 required a contractor with a
backhoe, while the fences constructed in 1972 were built by research personnel with hand tools in a few days. Table 1 shows the cost breakdown
for all fences. The new fences were placed in series of two fences (one
behind the other) because a single fence would have had to be 8-ft. high
to keep snow out of the shrub stands (Fig. 2). However, construction of
fences higher than 4 feet is very expensive and difficult. Fences in
series also provide more area protected by created drifts in which to
establish new forage stands.
All four designs were successful in trapping and storing snow. Drifts
up to 50 inches in depth were produced behind the 4-ft. fences and snow
depth in the shrub stands was reduced to an average depth of 13 inches of
light fluffy snow.
I would recommend design No. 2 (IO-foot post spacing, all posts braced
upwind) for future fence construction. The 10-foot post spacing provides
more support for keeping the slat snowfence suspended 6 to 8 inches above
the ground. This is necessary for proper drift placement. Downwind
bracing does not appear to be necessary unless the fences are located
where wind direction is highly variable. All designs withstood winds up
to 80 m.p.h. without any damage. No summer maintenance appears to be
necessary. It appears to me that these fences will last for several years
without any maintenance.

�-286-

Table 1.

Summary of snowfencecost.

Design 1.
Fence 4 feet high, 100 feet long. Slat snowfence suspended from #9 wire
attached to steel fence posts spaced 10 feet apart. All posts braced
upwind, alternate posts braced doWnwind with #9 wire attached to angle
iron braces.
Item

Cost perlOOft~

Steel post, 8 ft. long

Fence

$ 12.70

Wire, #9 black

19.00

Braces, angle iron

21.30

Slat snowfence

27.00

Labor (9 hrs. at $4.00/hr.)

36.00
Total

$116.00

$1.16 per linear foot - 79¢ per linear ft. excluding labor

Design 2.
Fence 4 feet high, 100 feet long. Construction same as design 3 except
no downwind braces. Alternate posts braced upwind.
Item

Cost per 100 ft~ Fence

Steel post, 8 ft. long

$ 12.70

Wire, #9 black

13.00

Braces, angle iron

14.20

Slat snowfence

27.00

Labor (9 hrs. at $4.00/hr.)

36~00
Total

$102.90

$1.03 per linear foot - 67 c per linear ft. excluding labor

�-287-

Table 1.

Summary of snowfence cost (continued).

Design 3.
Fence 4 feet high, 90 feet long. Slat snowfence suspended from 119 wire
attached to steel posts spaced 15 feet apart. All posts braced upwind,
alternate posts braced downwind.
Item

Cost per 90 ft. Felice

Steel post, 8 ft. long

$

8.10

Wire, 119 black

18.00

Braces, angle iron

15.60

Slat snowfence

27.00

Labor (8 hrs. at $4.00/hr.)

32~00
Total

$100.70

$1.11 per linear foot - 76¢ per linear ft. excluding labor

Design 4.
Fence 4 feet high, 90 feet long. Construction same as Design 5 except
no downwind bracing. Alternate posts braced upwind.
Item

Cost per 90ft~

Steel post, 8 ft. long

$

8.10

Wire, 119 black

11.00

Braces, angle iron

11.40

Slat snowfence

27.00

Labor (8 hrs. at $4.00/hr.)

32.00
Total

$ 89.50

99¢ per linear foot - 64¢ per linear ft. excluding labor

Felice

�-288-

Table 1.

Summary of snowfence cost (continued).

Equipment required to construct Designs 1, 2, 3, and 4.
1.

Post pounder

2.

Fence stretcher

3.

Fencing pliers

Design·5.
Fence 8 feet high,l40 feet long. Slat snowfence suspended from
5116-inch cable attached to wooden posts spaced 20 feet apart.
Braced upwind and downwind by 5/l6-inch cable.

Item

Cost per Fertce (140ft)

Wood poles, 6" top diameter

$

59.00

Slat snowfence

94.50

Wire, 119 black

5.50

L\Dllber,2" X 4" X 8'

8.00

Cable clamps

13.00

Cable, 5/16" diameter

72.50

Braces

20.00

Labor (contracted to Lickteig)

750.00
Total

$7.30 per linear foot of' fence

$1,023.00

�-289-

Table 1.

Summary of snowfence cost (continued) •

Design 6.
Fence 4 feet high, 140 feet long. Construction same as Design 1 except
for fence height and no downwind braces.
Item

Cost Eer Fence .(140·ft)

Wood poles, 6" top diameter

$

44.10

Slat snowfence

40.50

Wire, 119 black

5.40

Lumber, 2" X 4" X 4'

4.00

Cable clamps

13.00

Cable, 5/16" diameter

35.00

Labor (contracted to Lickteig)

492~00
Total

$

634.00

$4~53 per linear foot

Equipment required to construct Designs 5 and 6.
1.

Backhoe with post hole auger

2.

Fence pliers

3.

Fence stretcher

4.

Post tamper

5.

Wrenches

6.

Sledge hammer

..

--~~

�-290-

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- -

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wind
direction

Figure 1.

f.

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Ten-foot post spacing; alternate bracing of up-wind
no braces down-wind.
Fence is 100 feet in length~

"

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....

Ten-foot post spacing, braces on all up-wind posts and
alternate down-wind posts. Fence is 100 feet in length.

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�-291-

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no braces. down-wind.
Fence is 90 feet in length.
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alternate down-wind posts. Fence is 90 feet in length.

Design 4.

"

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/

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dots indicate

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�-292-

FENCE

70 feet

SHRUBS

300 yards

SHRUBS

Figure

2. Layout of snowfences on north side of Junction Butte.

�-293-

July, 1974

JOB PROGRESS REPORT

State of

CO=.;:U)=R=AD..;;:.O
,-.

Project No.

W-38-R-28

Work Plan No •.
Job Title

_

Job No._~la~

Monitor Potentially Critical Deer-Vehicle

Period Covered:
Personnel:

1_5

Deer-Elk Investigations
_
Accident Areas Statewide

April 1, 1973 through March 31, 1974.

Albert E. Byrne, Larry L. Green, William R. Heicher, David C.
Hoart, Richard F. McDonald (and all Wildlife Conservation
Officers in his area), J. Kris Moser, John W. Seidel, James R.
Adams, Thomas M. Pojar, Dale F. Reed, and Thomas N. Woodard.

ABSTRACT
The number and location of deer-vehicle accidents were monitored in five
general areas of Colorado. Recommendations for installation of devices
to reduce deer-vehicle accidents were submitted to the Division of Highways when-warranted.

�-294...,

RECOMMENDATIONS
Recommendations were developed from this study and made to the Colorado
Division of Highways for erection of the eight-foot fences and attendant
structures referred .toin this report.

�-295-

MONITOR POTENTIALLY CRITICAL
DEER-VEHICLE ACCIDENT AREAS STATEWIDE
Thomas N. Woodard

P. S. OBJECTIVE
To establish statewide, the location and number of deer-vehicle accidents
in areas appropriate to the evaluation of devices.
SEGMENT OBJECTIVES
1.

Examine potentially critical deer-vehicle accident areas reported
by management and as determined by Job 1.

2.

As appropriate, record all deer found killed in selected highway area.

3.

As appropriate, estimate deer densities in each selected highway area.

4.

Measure deer activity along the roadside or in the median of each
selected highway area when feasible.

5.

Summarize the Highway Department's traffic volume data for each
selected highway area as needed.

METHODS AND MATERIALS
Methods and materials have been described by Pojar (1972).
The locations and numbers of deer and elk-vehicle accidents during 1972
and 1973 in the Durango area were plotted on U.S.G.S quadrangle maps
(scale 1:24,000).

DESCRIPTION OF AREAS
Highway 6-24 (1-70) Rifle West
See Pojar (1972). Preliminary planning and design of the interstate
highway which will replace Highway 6-24 is completed but construction has
not begun. A total of 8 underpasses and 1 overpass with associated 8-foot
(2.44-m) fencing has been incorporated in the original highway design.
Highway 1-70 Eagle East
Pojar (1972) and Woodard (1973) described this area. A 4.75 mile (7.65 km)
8-foot fence was completed along the north right-of-way during September, 1973.

�-296-

Highway 1-70 Avon-Wolcott
Pojar (1972) and Woodard (1973) described this area.

Durango Area
Pojar (1972) described this area.

Highway 82 - Glenwood-Basalt
Myers (1969) described this area.

RESULTS AND DISCUSSION
Highway 6-24 (1-70) Rifle West
A total of 28 vehicle-killed deer were documented on 18 miles of highway
in the Rifle West study area; a reduction of 69.56 percent from the
.preceeding segment. Seventy five percent of the deer were killed from
October 1, 1973 to February 28, 1974.

Highway (1-70) Eagle East
A total of 60 vehicle-killed deer were documented on 8.5 miles of highway
in the Eagle East study area. Deer kill was reduced on the portions of
highway opposite the 4.8 mile barrier fence (Reed 1974).
Highway 1-70Avon~Wolcott
A total of 75 vehicle-killed deer were documented on 14 miles of highway
in the Avon-Wolcott study area; a reduction. of 42.75 percent from the
preceeding segment. Deer kill continued to be reduced on the portions of
highway opposite two sections of 8-foot fence (Reed 1974).
..
Deer kill decreased along the section of highway from the Eagle River
Bridge, 2 miles east of Wolcott, to approximately 1.5 miles east of that
point •. Thirteen deer were killed in this area. from January through
March in 1973 and 1 was killed in 1974 during the same period. Woodard
(1973) described a 14 x l4-foot (4.27 x 4.27-m) equipment underpass in
this area which could be used with 8-foot fencing as a deer underpass.
However, the decrease in kill during this segment.does not indicate
the need for barrier fencing.

�.•
297-

Durango Area
One hundred and thirty-eight deer and 41 elk were documented killed on
U.S. Highway 550 from Durango to the north approximately 15 miles (24.15 km)
during 1972 and 1973. Two concentration areas were delineated. Fiftyeight deer and elk were killed from the Durango city limits to the north
1.5 miles (2.41 km). Twenty-seven deer and elk were killed on a stretch
of highway beginning 1 mile north of Hermosa to the north 1 mile (0.62 km).
Three and one half miles (5.65 km) of 8~foot fencing on both sides of the
highway have been recommended to be included in a highway construction
project from Rockwood south. The fencing would tie into 2 underpass
structures included in initial construction plans.
One hundred deer and 7 elk were documented killed on U.S. Highway 160
between Durango and Mancos during 1972 and 1973. Concentration areas
were not evident. The deer underpass with associated 8-foot fencing
between Cherry Creek and Mancos was completed and evaluation begun on
April 1, 1974.
Two hundred and forty-five deer and 23 elk were documented killed on U.S.
Highway 160 between Durango and Pagosa Springs during 1972 and 1973. Two
concentration areas were delineated. Twenty-eight deer were killed from
0.4 miles (0.25 Ian) west to 0.65 miles (1.05 km) east of the intersection
of U.S. Highways 160 and 550 south of Durango. Nineteen deer and elk
were killed on a 1 mile stretch of highway east of the intersection of
U.S. Highway 160 and Colorado Highway 151. A deer overpass could be
utilized in this area since the highway bisects a hill near the center
of the kill area resulting in natural approaches from both sides.
Highway 82 - Glenwood-Basalt
Sixty-nine vehicle-killed deer were documented on 18 miles of highway
in the Glenwood-Basalt study area; a reduction of 58.18 percent from the
preceeding segment. Seventy percent were killed during February and
March. March, with 27 deer, had the highest monthly kill.
The highway construction project which includes 8-foot fencing in the
Carbondale East area (Pojar 1972), 9 miles southeast of Glenwood, will
be completed during the next segment. A 1.1 mile 8-foot fence in the
Diamond-S area, 7 miles southeast of Glenwood Springs was reported by
Reed (1974).

�-298-

LITERATURE CITED
Myers, Gary T. 1969. An investigation of deer-auto accidents.
Colo.
Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. W-38-R-23.
Game Res. Rep., July, Part 2. pp. 147-178.
Pojar, Thomas M. 1972. Monitor potentially critical deer-vehicle
accident areas statewide. Colo. Div. Wildl. Game Res. Div. Fed.
Aid Proj. W-38-R-26. Game Res. Rep., July, Part 3. pp. 305-310.
Reed, Dale F. 1974. An evaluation of deer-proof fence length required
to prevent deer movements on or across high speed highways. Colo.
Div. Wildl. Game Res. Div. Fed. Aid Proj. W-38~R-28. Game Res.
Rep. In Progress.
Woodard, Thomas N. 1973. An evaluation of deer-proof fence length
required to prevent deer movements on or across high speed highways.
Colo. Div. Wildl. Game Res. Div. Fed. Aid Proj. W-38-R-27. Game
Res. Rep., July, Part 3. pp.

~

Prepared by

Ab'V'~s 1/ {'JJMe?~

Thomas N. Woodard
.
Wildlife Researcher Candidate

�-299-

July, 1974

JOB FINAL REPORT

State of
Project

....:C:..:o:.::L:..:O~R:..:A.:::.D:::..O
_
W-:-38-R-28

No.

Work Plan No.

15

Deer-Elk

Investigations

Job No.

6

Job Title:

D_e_e_r
__U_n_d_e_r~p_a_s_s
__Ev_a_l_u_a
__ti_o_n

Period

April 1, 1973 through March

Covered:

Personnel:

Thomas N. Woodard,

_
31, 1974.

Thomas M. Pojar,

and Dale F. Reed.

ABSTRACT
The first draft of a manuscript on the behavioral response of mule deer
to the Vail deer underpass has been completed and is in various stages
of editing.
The manuscript will be submitted to the Journal of Wildlife
Management.

��-301-

DEER UNDERPASS

EVALUATION

Dale F. Reed

P. S. OBJECTIVE
Determine if deer migrating from winter range on one side of Interstate
70 to summer range on the opposite side utilize an underpass constructed
in the area.

METHODS AND MATERIALS
See manuscript in preparation described below.

RESULTS AND DISCUSSION
Manuscript in Preparation

Reed, D. F., T. N. Woodard, and T. M. Pojar. Behavioral response of
mule deer to an underpass. J. Wildlife Mgmt.

C:-.

Prepared by

..; . ~&lt; -//1
/ .&gt; ~l /
~ Ai Q.
t:l.A.e&lt;

i

ale F. Reed
Wildlife Researcher

��-303July,

1974

.JOB PROGRESS REPORT

State

of

Project

--.,;OO=ID=RADO::.:=

No.

w-~38_-_R:_-_2_8 ___.:: Deer-Elk. Investigations

Work Plan No. __
Job T1.t1e

..;;;1;;:;5

~:

.rob No.__ 6a

_

Eva1uation of Deer Underpasses
-------------------------~-------------------------------

Period Covered:
Personnel:

_

April ,1, 1973 through March 31, 1974.

ThomasN. Woodard,..ThomasM. Pojar,.. and Dale F. Reed.

ABSTRAC'r
An arch deer underpass with associated 8-foot {2.44-m} fences along the
highway right-of-way was completed west of Durango toward the end of this
segment. De~r use of the underpass will be monitored during the next
segment.

��-305.•

EVALUATION OF DEER UNDERPASSES
Dale F. Reed
Mule deer response to an underpass 3.05 m (10 feet) by 3.05 m and 30.48 m
(100 feet) long under Interstate 70 has been reported (Reed, 1971). This
study purports to evaluate underpasses of different sizes and design.

P. S. OBJECTIVE
Determine the use of underpasses by deer in critical highway deer kill
areas and improve the effectiveness of these structures.

SEGMENT OBJECTIVE
Measure deer use of underpasses.
MATERIALS AND METHODS
Construction of a 20-foot wide and 10-foot high structural plate pipe
arch deer underPass was completed by the Colorado Division of Highways.
The 8-foot (2.44-m) fence was completed to the underpass toward the end
of this segment.

RESULTS AND DISCUSSION
The initial effort in monitoring deer use of the structure was coordinated
with management personnel.

LITERATURE CITED
Reed, Dale. F. 1971. Deer underpass evaluation. p. 341-351. In Game
Research Report, July Part 3. Colo. Game, Fish and Parks Div. Federal
Aid.

Prepared by _C
__
"_~"_·
....•
· .-.:~/~
•.••
/):./I::V...s.."' __ (_~I-/'-;"'::";~"'-o&gt;"'...J"·!I;..;.:~"'::"l";z:.(~i_) __
/kG'L &gt;'s'
b'-JI&lt; ""'.~
Dale F. Reed
"
Wildlife Researcher

--,-

��-307July, 1974
JOB PROGRESS REPORT

State of

COLORADO

-------------------------------------

Project No.
Work Plan No.

W-38-R--28

Deer-Elk Investigations

15

Job No.

7

--------------------~----------------------

Job Title Effects of Highway Lighting on Number of Deer Killed by Vehicles
Period Covered:

April I, 1973 through March 31, 1974.

Personnel: Claudia A. Doose, Larry L. Green, John W. Seidel, Allen F.
Whittaker, Dale F. Reed, and Thomas N. Woodard.

ABSTRACT
The effect of highway lighting on the rate of deer-vehicle accidents was
evaluated. Estimated deer crossings per kill increased by 86.8 percent
with the lights on compared to.lights off. Th~re was no significant
difference in the crossings per kill ratios (X = 0.60, P&gt;0.25).
The
amount of light available at 3 known kill locations during lights on
nights ranged from 0.32 to 3.85 foot-candles. There was no significant
difference (P&gt;0.50) in southbound mean speeds or in northbound mean
speeds (P&gt;0.40) with lights on or off.

��-309-

EFFECTS OF HIGHWAY LIGHTING
ON NUMBER OF DEER KILLED BY VEHICLES
Thomas N. Woodard
P. S. OBJECTIVE
Determine if highway lighting affects the rate of deer-vehicle accidents
on a portion of Highway 82.
SEGMENT OBJECTIVES
1.

Record the number of deer killed by vehicles at each end of, and
within a 3/4 mile lighted portion of Colorado 82.

2.

Estimate deer densities adjacent to the portion of highway affected
by the lighting.

3.

Compare pre-treatment kill and count data to post-~reatment kill
and count data.

4.

Gather additional information which may be needed to understand why
deer-vehicle accident rates are or are not affected by lighting.
METHODS AND MATERIALS

Thirteen 37,000 lumen (1 lumen=0.093 fc/sq.m), 700 watt, clear mercury
vapor lamps mounted on 10-foot (3.05-m) arms at the top of 40-foot (12.2-m)
metal poles were used to light the highway. Nine lamps, spaced at
approximately equal intervals, illuminated a 0.3 mile (0.48 km) section
of highway (Fig. 1). Two lamps installed at each end and spaced at 400foot (122-m) and 600-foot (183-m) intervals provided transition lighting.
The effect of lighting was evaluated from January 1, 1974, to March 26,
1974. The lights were alternately turned on and off for one week periods.
Six week periods each of lights on and off data were collected.
The location of each deer vehicle accident was documented to the nearest
0.1 mile (.161 km) from 0.25 mile (0.40 km) markers established along the
highway (Reed 1969).
The number of deer that crossed the highway was estimated by making
nightly spotlight counts (Reed 1969) and track counts in the median
strip when snow conditions permitted. One crossing was arbitrarily
added for every deer killed on the highway.

�-310-

Vehicl~ speeds were recorded between 7:00 and 11:00 PM during 35 nights
with an automatic vehicle speed recorder. Eighty speeds were randomly
selected from each nights sample for tabulations.
The amount of light (foot-candles) available at each known kill location
during lights on nights was measured with a light meter (Gassen Luma-Pro).
DESCRIPTION OF AREA
The study area is located 3 miles (4.83 km) south of Glenwood Springs on
Colorado State Highway 82. This 0.7 mile (1.13 km) segment of highway
has 4 lanes and a posted speed limit of 55 miles per hour. Adjacent
deer winter range is pinyon-juniper covered slopes and sagebrush flats
to the east and farmland and sagebrush flats to the west.
RESULTS AND DISCUSSION
Estimated crossings per kill increased by 86.8 percent with the lights
on compared to lights off (Table 1). There was no significant difference
in the ratios (x2=0.60, P &gt;0.25).
The amount of light available at three known kill locations during lights
on nights ranged from 0.32 to 3.85 foot-candles.
Mean vehicle speeds with lights on were 49.54 southbound and 49.22
northbound. Mean speeds with lights off were 49.33 southbound and 49.56
northbound. There was no significant difference (P&gt; 0.50) in southbound
mean speeds or in northbound mean speeds (P &gt;0.40) with lights on or off.
Table 1. Estimated deer crossings and total kill in the Highway 82
lighting study area from January 1, 1974, to March 26, 1974.
Lights Off
Lights On
Estimated Crossings

609

506

Total Kill

9

4

Crossings/Kill

67.7

126.5

�-311-

LITERATURE
CITED
Reed, Dale F. 1969.
Techniques fordeterinining
potentially
critical
deer highway crossings.
Game Informat;:ion Leaflet No. 73.
Colo.
Dept .. Natural Resources , Div. of Game, Fish and Parks.
3p.

.
Prepared by

,,.,....-:;.1&amp;. ()l{j/!Af( .~

l' ct-" o-e,&amp;(,-.,.+/
! ..

//

;,,-/,

Thomas N. Woodard
Wildlife
Researcher

Candidate

�I

..-I

N
C"')
I

Fig. 1.

Highway lighting on Colorado Highway 82 south of Glenwood Springs (Photo by Dale F. Reed).

�-313-

July,

1974

JOBPROGRESS
REPORT

State of

COLORADO
--------~~~~-----------

Project

No.

W-38-R-28

:

Deer-Elk Investigations

------------------------10
: Job No.
15
WorkPlan No.
--------------~-------------------------------------An.Evaluation of 8-Foot Fence Length Re~red
Job Title

to Prevent Movementson or Across High Speedtlighways

Period Covered: April 1, 1973 through March 31, 1974.
Personnel:

ThomasN. Woodard,ThomasM. Pojar, William R. Beicher, David
C. Boart, Larry L. Green, Claudia A. Doose, and Dale F. Reed.

ABSTRACf
Annual or seasonal deer kill caused by vehicles was reduced by 61, 82, 63,
and 73 percent along 8-foot (2.44-m) fence lengths of 1.1, 2.25, 2.25, and
4.8 mile&amp; respectively.
One hundred fourteen, 13, and 33 deer escaped the
highway rights-of-way adjacent to the two 2.25 and the 4.8 mile long8-foot
fences through 17 one-way deer gates respectively.
Does (N=8)and bucks
(N=3)movedlaterally
or parallel to the 8-foot fence for mean distances
of 0.37 and 0.58 miles, respectively.

���-315-

AN EVALUATION OF 8-FOOT
FENCE LENGTH REQUIRED TO PREVENT DEER
MOVEMENTS ON OR ACROSS HIGH SPEED HIGHWAYS
Dale F. Reed

P. S. OBJECTIVE
To evaluate length of 8-foot (2.44-m) fence in relation to deer movement
on or across high speed highways.
SEGMENT OBJECTIVES
1.

install 8-foot fence along appropriate sections of highway as determined by Job 1a.

2.

Evaluate the effectiveness of various lengths of 8-foot fence in
affecting deer movements.
METHODS AND MATERIALS

Methods and materials have been described by Pojar (1972) and Reed et a1.
(1974a) •
DESCRIPTION OF AREA
Highway 82 - Diamond S
The Highway 82 study area has been described by Pojar (1972).

Interstate 70 - Avon, Edwards, and Vail
The Interstate 70 - Avon and Edwards study areas have been described by
Woodard (1973) and the Vail study area by Reed (1971) and Reed et a1.
(1974a) •
Interstate 70 - Eagle
The Eagle study area consists of a segment of the Interstate from near the
Eagle Interchange east 4.8 miles. The Interstate highway was opened to
traffic October 5, 1972. An 8-foot (2.44~) fence was completed along the
north side of the highway right-of-way October" 5, 1973. Alfalfa fields are
prevalent south of the highway. Pinyon-juniper and some big sagebrush
occurs on the north.

�-316-

RESULTS AND DISCUSSION
Highway 82 ~ Diamond S
Deer concentrate in the crested wheatgrass fields northeast of the
highway in late winter and early spring (Table 1). The mean number of
deer crossings during March - May periods in 1971 before the 8-foot
fence was installed was 11.7 CN=32, where N=number of 24 hour periods)
per day. After the fence was installed the means were 2.1 (N=38) in
1972 and 5.5 CN=34) in 1973. This represents an 82 and 53 percent
reduction in crossings in 1972 and 1973 respectively.
The mean deer kill per year before the 8-foot fence was installed
(1968-1971) was 7.75. Three deer were killed in 1972 and three in 1973
(after 8-foot fence was installed); a mean reduction of 61 percent.

Table 1. The March-May mean number of deer observed on spotlight
counts between quarter-mile section markers 25 to 30 for 1968 through
1973 on Highway 82 (n=number of counts).
Month

1968

1969

Mean Count
1970
1971

1972

1973

March

134.8
Cn=4)

151.2
(n=4)

104.5
(n=4)

66.8
(n=4)

102.2
(n=4)

137.4
(n=5)

April

73.0
(n=4)

34.0
(n=5)

56.0
Cn=5)

51.4
(n=5)

4.5
(n=4)

47.0
Cn=4)

May

6.0
Cn=5)

0.0
(n=L)

1.5
(n=2)

0.0
(n=2)

66.2
(n=13)

77 .5
(n=10)

63.7
(n=l1)

47.6

53.4

(n=Ll.)

(n=B)

97.2
(n=9)

MEAN
TOTAL

Interstate 70 - Avon
The mean number of deer observed on spotlight counts during the
August - December periods was 15.5 Cn=17) and 13.4 Cn=2l) in 1971 and
1972 respectively, and 10.8 (n=Ll.) in 1973 for October - December. Deer
crossings were not monitored in 1973. However, 114 deer made passages to
the north through six one~ay deer gates lQcated in the 8-foot fence CReed et a1.
1974b) ~ Twenty eight deer were killed from October 5, 1971, to October 5,
1972, before the fence was installed. Five were killed from October 5,
1972, to October 5, 1973, after the fence was installed; a reduction of
82 percent.
A deer guard in the Avon 8-foot fence was monitored for deer use. Deer use
of this guard plus deer use and response to another guard was reported previously (Reed et al. 1974a).

�-317-

Interstate 70 - Edwards
Twenty seven deer were killed on this segment of highway before the
fence was installed from October 5, 1971, to October 5, 1972. Ten were
killed after the fence from October 5, 1972, to October 5, 1973; a
reduction of 63 percent. Three areas in the fence were in need
of repair and maintenance and were not considered "deer proof".
Thirteen passages occurred through four of seven one~ay deer gates
located in the 8-foot fence.

Interstate 70 - Vail
No additional data was collected in this study area in regard to fence
length.

Interstate 70 - Eagle
The mean number of deer observed on spotlight counts.during October March 1972-73 was 80.5 (n=25). One hundred forty six deer were· killed
from October 5, 1972, to March 31, 1973, before the fence was installed.
Thirty nine were killed from October 5, 1973, to March 31, 1974, after
the fence was installed; a reduction of 73 percent. An estimated
90-100 deer, including several collared animals (Fig. 1), came across
the highway from the south and either 1.) passed through the one-way
deer gates or an opening in the fence, 2.) went around the ends of the
fence, 3.) went back to the south to or across the Eagle River, or 4.)
were killed on the highway. Thirty three deer made passages to the
north through seven of ten one-way deer gates located in the 8-foot
fence.
According to 39 observations of collared deer north of the 8-foot
fence, does (N=8) and bucks (N=3) have moved laterally or parallel to
the fence for mean distances of 0.37 and 0.58 miles, respectively.
Buck No. 64 moved laterally or parallel to the fence for the greatest
distances (Fig. 2).

LITERATURE CITED
Pojar, Thomas M. 1972. An evaluation of deer-proof fence length required
to prevent deer movements on or across high speed highways. Colo.
Div. Wi1d1. Game Res. Div. Fed. Aid. Proj. W-38-R-26. Game Res.
Rep., July, Part 3. pp. 305-310.
Reed, Dale F. 1971. Deer underpass evaluation. Colo. Div. Wi1d1.
Game Res. Div. Fed. Aid. Proj. W-38-R-25. Game Res. Rep. July,
Part 3. pp. 341-351.
Reed, D. F., T. M. Pojar, and T. N. Woodard. 1974a. Mule deer responses
to deer guards. J. Range Manage. 27(2):111-113.

�-318-

Reed,

D. F., T. M. Pojar, and T. N. Woodard. 1974b. Use of one-way
gates by mule deer. J. Wildl. Manage. 38(1):9-15.

Woodard, Thomas N. 1973. An evaluation of deer-proof fence length
required to prevent deer movements on or across high speed
highways. Colo. Div. Wildl. Game Res. Div. Fed. Aid. Proj.
W-38-R-27. Game Res. Rep., July, Part 3. pp.

Prepared

b1:~d/di;i)
ale F. Reed
Wildlife Researcher

�-319-

Fig. 1. Collared doe No. 50 came across Interstate 70 from the unfenced
south side previous to January 15, 1974. The animal was identified on
the north side of the 8-foot fence on three subsequent occasions, having
apparently passed through one of the one~ay gates (Photo by Larry Green).

�-320-

-N10 DEC72
22 NOV73

.75
10DEC 73

~w-

-E- .

I.~
2 JAN 74

eJAN.74

I--

O;.,o,,;;;;.9.........-_~ &lt;i&gt;.

13JAN74

@

1.30

SCALE: III • 0.5 MILE

x •ORIGINAL TRAP SITE

,fig. 2. A diagram illustrating
the apparent movements of buck No. 64
north of the Eagle 8-footfence'acccrding
to seven dated sightings
(2-8).
The animal was collared at point X December 10, 1972.

�-321-

July, 1974

JOB PROGRESS REPORT

State of

COLORADO
------------~~~~---------------Job No.
1~1~
~---Effects of a Simulated 8-Foot Fence Angle
in Diverting Deer from Their Established Direction of Movement

Work Plan No.
Job Title

Period Covered:
Personnel:

Deer-Elk Investigations

W-38-R-28

Project No.

15

April 1, 1973 through March 31, 1974

Thomas M. Pojar, Dale F. Reed, and Thomas N. Woodard.

ABSTRACT
Time was measured and observations made on mule deer (Odocoi1eus hemionus)
response to five simulated fence angles in 1972 and three simulated fence
angles in 1973. Mean time needed for deer to successfully travel the 27.42
meter length of 900 fence angle was significantly greater (P&lt;O.OOl) than
the mean control time (00 angle). The number of deer that failed to move
through the zone during the 900 .fence angle measurements was not significantly
greater (X2 = 0.202, P&gt;0.50) than the number during control.

��-323-

EFFECTS OF A SIMULATED 8-FOOT FENCE ANGLE
IN DIVERTING DEER FROM THEIR ESTABLISHED DIRECTION OF MOVEMENT
Thomas N. Woodard
P. S. OBJECTIVE
Evaluate the effectiveness of three angles of a simulated 8-foot fence
to divert deer from their established direction of movement.
SEGMENT OBJECTIVES
1.

Install a movable 8-foot section of 4-inch square netting adjacent to and
across heavily traveled deer trails and concentration points.

2.

Test the effectiveness of three angles under field conditions.
METHODS AND MATERIALS
Methods and materials have been described by Reed (1973).
Observations were made during 21 days in 1973 from June 12 to July 9.
600 angle was omitted in 1973.

The

DESCRIPTION OF AREA
The study area has been described by Reed (1973).
RESULTS AND DISCUSSION
Time was measured on deer in the simulated fence angle area on 55 occasions
(Table 1). Twenty-nine times were recorded during control measurements.
Mean time needed for deer to successfully travel the 27.42 meter length of
900 fence angle was significantly greater (P &lt;0.001) than mean control
time. The number of deer that failed to move through the zon during the
900 fence angle measurements was not significantly greater (X -0.202,
P &gt;0.50) than the number during control; thus the simulated fence angle
did not preclude deer from their apparent direction of movement.

z

Statistical tests were not applied to the other angle variables because of
insufficient sample size.

�-324-

Table 1. The mean times (seconds) of successes or failures of deer moving
through the zone with either the simulated fence angle removed (control)
or installed (variables: 45°,60°, or 90°). Successful animals moved along
the permanent 8-foot (2.44-m) fence or along the simulated fence. Failure
animals moved into the zone, turned around, the left in the direction that
they had come.
Angles

Successful

Failure

Percent Successful

0°

1/
75.8 (n=19) -" 85.6 (n=lO)

66

45°

168.2 (n=3)

60

60°
90°

0
173.6 (n=9)

52.0 (n=Z)
0
78.3 (n=6)

120°

0

121.3 (n=4)

135°

73.0 (n=L)

117.0 (n=L)

60

50

Number of observations (n).
When deer approached the simulated fence angles of 60° and 45° from the
opposite direction (east), they encountered converse angles of 120° and 135°.

Prepared

by

--z1.;:~;yti. w1;;,da!~),'-.,;.JLv,f
Wildlife Researcher Candidate

�-325July,

1974

JOB r~!OGi-:ESS P.h"1'ORT

State of

-----------------------------

Project: No.

W-38-R-28

Work Plan No.
Job Title

16
:
------~~-------------

Deer-Elk Investigations
.!oh lh.

1
-----~=----------------------

Piceance Deer Study - Population Distribution

Period Covered: April 1, 1973 through March 31, 1974.
Personnel:

R. M. Hartmann, S. F. Steinert,
and M. J. Loftsgard.

J. J. Klein, Jr.,

H. Reiswig,

Deer were trapped and marked from December18, 1973 through March 8, 1974.
Five hundred forty-one deer were marked of which 496 were both eartagged
and neckbanded, 44 eartagged only and .1 neckbanded only. A mean 6.7 deer
per day were marked. Trapping was completed in GameManagementUnit 22 and
operations were movedfarther up the White River into Unit 23 where marked
deer quotas were nearly reached in two more areas.
The composition of the
total catch was_108male fawns, 86 female fawns, 56 mature males, 290
mature females and 1 unknownfemale. Sightings of 346 deer were reported
during the year but only 74 occurred during the time deer were considered
to be. on or near the sUIIIDer
range. Eighty-one marked deer were recovered
but at least 33 of these died during winter or spring, 1973, on the winter
range. Three aerial deer dist-:ribution surveys were flown during winter.
The survey flown in February was based on accumulated tracks and showed
the deer to be somewhatconcentrated because of deep snow.

��-327-

PICEANCE DEER STUDY - POPULATION DISTRIBUTION
Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.

SEGMENT OBJECTIVES
1.

To delineate deer sub-population boundaries and concentration areas
on the Piceance winter range.
a.

Define the winter range boundaries of the Piceance deer population
under mild, moderate, and severe winter conditions.

b.

Define deer SUb-population boundaries within the Piceance winter
range.

c.

Delineate major deer concentration areas within the Piceance winter
range.
METHODS AND MATERIALS

See Bartmann (1972).

RESULTS AND DISCUSSION

Deer Trapping
Deer trapping began December 18, 1973 and terminated March 8, 1974. Snow
cover conditions were similar to the previous year. Mean snow depths averaged
near 15 to 18 inches most of the winter and snowmobiles were required to set
and check traps almost the entire winter. Temperatures, however, averaged
slightly higher than last year and deer condition remained good through
most of the winter except that some fawns began looking "rough" around late
February. The number of traps used was similar to last year, 30 to 40.
Trapping success was the highest thus far, averaging 6.7 deer per day compared to 5.9 and 5.7 deer per day the past two years. Trapping was completed
in Game Management Unit 22 and operations were moved farther up the White
River into Unit 23 where quotas were nearly reached in two more areas. This
leaves about 560 deer to be banded in six areas.

�-328-

Five hundred forty-one deer were marked during the 1973-74 winter, of
which 496 were both eartagged and neckbanded, 44 eartagged only, and 1
neckbanded only (Table 1). The sex and age composition was 108 male
fawns, 86 female fawns, 56 mature males, 290 mature females, and 1 unknown female. Fawns comprised 36 percent of the total compared to 26
percent in 1972-73 and 34 percent in 1971-72. The percent males was 10
compared to 17 and 18 percent the previous two years. Mature females
comprised 54 percent of the total compared to 57 and 48 percent the previous two years.
It was noted that during the last 16 days of trapping, 45 deer were caught
and only 2, or 4 percent, were fawns. During the preceding 20 days in
February, 57 of 160 deer, or 36 percent, were fawns. The reason for this
discrepancy is not known at this time.

Marked Deer Sightings and Recoveries
Sightings of 346 ~Erked deer were recorded from March 16, 1973 through
February 28, 1974 (Table 2). Another four sightings made before the
earlier date were also reported during this period and are included.
Most sightings, 213, were recorded during spring prior to June 1, 1973.
During this interval, deer were present on meadows and other areas where
large numbers could be easily observed and, with several exceptions,
sightings were recorded only if the number on the neckband was read.
Another 74 sightings were recorded from June through October, 1973.
During this period most deer were on or near summer range. The remaining
63 sightings occurred during the winter months.
Previously, observations from May through September were considered summer
sightings and those from October through April winter sightings. However,
in 1973 most deer were still on the winter range in May. The October
hunting season fell during mid-month with mild weather and deer were mostly
near the higher summer range elevations at that time.
Recoveries of 81 marked deer were reported from April, 1973 through February,
1974 (Table 3). At least 33 of these died during the winter and spring of
1973, many soon after they were marked. Ten of the 33 died after getting
caught in fences.

Deer Distribution
The 1973-74 winter was similar to the 1972-73 winter with respect to snow
cover. Snow began accumulating in December and depths eventually ranged
from 12 to 30 inches and averaged around 15 to 18 inches most of the winter.
South slopes, except in a few areas in the southwest part of the winter.
range, remained snow-covered into early February. The mean temperature
from December through Februarz was nearly 50 F warmer than for the same
period in 1972-73 and about 4 F colder than in 1971-72.

�-329-

Three aerial distribution surveys were accomplished. The first was
made December 19, 20, and 21, 1973. Snow depths ranged from 8 to 15
inches and averaged from 10 to 12 inches. Greatest depths occurred
in the eastern and southern portions of the winter range.
Deer were fairly well scattered, this being the case for the first
survey the past two years (Fig. 1). Only the highest elevations in
the south and southwest part of Unit 22 and the higher areas of the
Piceance triangle were devoid of deer tracks. Numerous areas of little
or no deer use were observed throughout the winter range but distribution was still fluid and many of these areas contained deer a few
days or weeks later. The few areas of heavy use mostly appeared in the
same general areas as in the past.
From six to eight inches of snow fell the last week of December increasing
the average depth to 18 to 20 inches. The second survey was flown on
December 31, 1973 and January 2 and 3, 1974. The deer were still scattered
but fewer areas of little or no use were found at the lower elevations,
particularly on the more extensive slopes (Fig. 2). A few new areas of
heavy use were noted but there was no large increase in the to.tal area
of such use. In a few places the deer were starting to use trails and
were already avoiding the larger treeless expanses.
As in the past two years, little snow fell after mid-January. Therefore
an effort was again made to estimate deer distribution based on accumulated
tracks on February 12 and 13, 1974. For several weeks prior to the survey
snow had been slowly melting and the steeper south slopes were beginning
to bare. The melting process made fresh tracks impossible to distinguish.
The deer were also using trails instead of moving separately as they did
earlier in the winter. During the survey, snow depths over much of the
occupied winter range averaged near 15 inches with greater depths, up to
two feet, on north slopes.
The upper limits of deer use had changed noticeably in most areas (Fig. 3).
Heaviest usewas found in the eastern portion of the unit although there
were still some animals present on north slopes. Deer use appeared
considerably lighter in the western portion of the unit and many of the
tracks were believed old. Thus, the distribution limits, at least in this
area, probably exaggerate to some extent the actual distribution. The
avoidance by deer of treeless areas was especially evident during this
survey.
LITERATURE CITED
Bartmann, R. M. 1972. Piceance deer study - population distribution.
p. 315-337. In Game Research Report. Colo. Div. Wildl., Denver.
3(Part 3):253-377.

.....
~~
Prepared by £A2.~1~,;Z
Richart M. Bartmann
Wildlife Researcher

�Table l. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued) •

-

Date
(1973)

Sex

Age

Eartag
no.

Neckbnnd
no.

Twp.

12-23
12-23
12-23
12-24
12-24
12-24
12-25
12-25
12-25
12-25
12-26
12-26
12-26
12-26

Female
Female
Male
Male
Male
Male
Female
Female
Female
Female
Male
Female
Male
Female

Mature
Mature
Fawn
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn

L-2342
L-2343
L-2344
L-2345
L-2346
L-2347
L-2348
L-2349
L-2350
L-235l
L-2352
L-2353
L-2354
L-2355

31
59
66
68
98
92
58
63
69
62
48
71
60
93

2S
2S
lS
lS
2S
2S
lS
lS
IS
IS
IS
2S
2S
2S

Summary:

Location
Range
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

Sec.

Remarks

3
3

34
36
3
3
16
9
15
36
28
12
1
3

21 Male Fawns ; ·16 Female Fawu~; .'8 :Ma.tu~~Males_;'31 Mature Females
Area 4 - Green with White Stripe·Neckband·

12-21
12-21
12-22
12-23
12,...23
12-23
12-23
12-23
12-23
12-23

Female
Female
Female
Male
Female
Female
Female
Female
Female
Female

Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature

L-220l
L-2202
L-2203
L-2204
L-2205
L-2206
L-2207
L-2208
L-2209
L-22l1

65
86
62
81
63
70
.67'
69
66
60

lN
lN
lN
lN
lN
IN
lN
IN
lN
IN

96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

18
18
17
17
17
17
17
7
7
7

------~------~----------------------------.-------------------~~----------------------'~----------

I

w
w

I'.)
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued) •
Date
(1973)

Sex

Age

12-23
12-23
12-24
12-24
12~25
12-25
12-25
12-25
12-25
12':"26
12-26
12-26
12-26
12-26
12-26
12-26
12-27
12-27
12-28
12-28
12-28
12-28
12-28
12':"28
12-29
12-29
12-29
12-29
12-29
12-29
12-29

Female
Male
Female
Male
Male
Female
Female
Male
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Male
Female
Male
Female
Female

FaWn
Mature
Mature
Fawn
Fawn
Fawn
Fawn
Fawn
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Fawn
Fawn
Mature

Eartag
no.
L-2210
L-2212
L-2213
L-2214
L-2215
L"""2261
L-2262
L-2263
L-2264
L-2265
L"""2266
L-2267
L-2268
L-2269
L-2270
L-2271
L-2272
L-2273
L-2274
L-2275
L-2276
L-2277
L-2278
L-2279
L-2280
L-228l
L-2282
L-2283
L..;,,2284
L-2285
L-2286

Neckband
no.

Twp.

74
82
87
126
127
165
61
79
68
110
90
178
189
132
72
176
175
174
114
115
None
109
154
179
156
193
94
154
93
158
162

1N
1N
1N
1N
1N
1N
1N
1N
1N
1N
IN
1N
1N
IN
IN
1N
IN
IN
1N
1N
IN
1N
1N
1N
1N
1N
1N
1N
1N
1N
1N

Location
Range

Sec.

96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

7
7
7
18
18
17
17
26
26
7
7
18
16
16
16
26
7
17
17
17
16
16
16
22
18
7
7
7
17
17
20

Remarks

I

May be wrong band no.

May be wrong band no.

May be wrong band no.

May be wrong band no.

--------------------------~--,--------~~------------------------------------------------------------------------

w
w
w
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
Date
(1974)

Sex

1-12
1-12
1-12
1-12
1-12
1-12
1-12
1-12
1-13
1-13
1-13
1-13
1-13
1-13
1-13
1-14
1-14
1-14
1-14
1-14
1-15
1-15
1-15
1-15
1-15
1-15
1-15
1-15
1...
15
1-16
1-16
1-16
1-16

Female
Female
Female
Male
Female
Female
Male
Female
Male
Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Male
Female
Female
Male
Female
Male
Male
Male
Male

Age

Eartag
no.

Neckband
no.

Twp.

Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn

L-2435
L-2436
L-2437
L-2438
L-2439
L-2440
L-2441
L-2442
L-2443
L-2444·
L-2445
L-2446
L-2447
L-2448
L-2449
L-2450
L-2451
L-2452
L-2453
L-2454
L-2455
L-2456
L-2457
L-2458
L-2459
L-2460
L-2461
L-2462
L-2463
L-2465
L-2466
L-2467
L-2468

147
145
119
None
135
None
207
163
176
100
2];5
None
221
220
216
219
216
223
218
227
143
226
222
None
None
None
230
None
None
None
205
208
None

18
1N
18
18
1S
18
18
18
18
18
18
18
1N
1N
18
18
1N
18
18
18
18
18
18
1N
18
18
18
18
18
18
18
1S
18

Location
Range
95W
95W
95W
95W
95W
95W
95W
94W
96W
96W
96W
96W
95W
95W
95W
95W
95W
94W
94W
94W
95W
95W
95W
95W
94W
94W
94W
94W
94W
95W
95W
95W
95W

Sec.

Remarks

34
34
2
2
2
2
2

17
1
1
6
6

34
34
2

2
34
17
17
17
2
2
2

34
17
17
17
17
17
6
2

2
3

-----------------------------------------------------------------------------------------------------------

I

W
W
0\
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
~

Date
(1974)

Sex

Age

Eartag
no.

Neckband
no.

l'wp.

1-16
1-16
1-16
1-16
1-16
1-16
1-16
1-16
1-16
1-17
1-17
1-17
1-17
1-17
1-17
1-17
1-17
1-17
1-17
1-17
1-17
1-18
1-18
1-18
1-18
1-18
1-19
1-19
1-19
1-19
1-19

Female
Male
Female
Female
Male
Male
Female
Male
Female
Female
Male
Male
Male
Female
Female
Female
Female
Male
Female
Female
Male
Male
Female
Male
Female
Female
Female
Male
Male
Female
Male

Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn

L-2469
·L-2470
L-2471
L-2472
L-2473
L-2474
L-2475
L-2476
L-2477
L-2479
L-2481
L-2482
L-2485
L-2483
L-2484
L-2486
L-2487
L-2488
L-2489
L-2490
L-2491
L-2492
L-2493
L-2494
L-2495
L-2496
L-2497
L-2498
L-2499
L-2500
L-2356

233
169
229
224
202
None
228
None
234
None
None
None
None
210
None
None
225
None
199
232
None
172
211
97
204
201
200
170
168
206
128

IS
IS
IS
IS
IS
18
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IN
IN
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS

e

Location
Range

Sec.

94W
94W
94W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
94W
94W
94W
94W
94W
94W
94W
95W
94W
94W
94W
94W

17
17
17
17
17
17
17
17
17
17
17
17
17
2
2
2
2
34
34
7
7
7
7
7
17
17
6
7
7
7
7

Remarks

-----------------------------------------------------------------------------------------------------------

I

w
w
-....I
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).

(£~14)

Sex

1-19
1-19
1-19
1-20
1-20
1-20
1-20
1-20
1-20
1-20
1-20
1-20
1-21
1-22
1-23
1-23
1-23
.1-24
1-25
1-25
1-25
1-26
1-26
1-26
1-27
1-27
1-27
1-28
1-28
1-28
1-29

Female
Female
Female
Female
Female
Male
Male
Male
Female
Male
Male
Female
Female
Female
Male
Male
Female
Female
Male
Female
Male
Female
Male
Male
Female
Female
Female
Male
Female
Female
Male

Age

Eartag
no.

Neckband
no.

Twp.

Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn

L-2357
L-2358
L-2359
L-2360
L-2361
L-2362
L-2363
L-2364
L-2365
L-2366
L-2367
L-2368
L-2369
L-2370
None
L-2371
L-2372
L-2373
L-2374
L-2375
L-2501
L-2502
L-2503
L-2504
L-2505
L-2506
L-2507
L-2508
L-2509
L-2510
L-2511

213
214
209
101
231
96
105
212
129
144
95
186
240
244
203
241
236
None
None
248
238
249
149
None
245
246
247
410
None
253
243

1S
1S
18
1S
18
1S
18
18
18
18
1S
18
18
18
1N
18
18
18
18
18
18
1N
18
18
1N
18
1S
1N
1S
18
1N

Location
Range

Sec.

94W
94W
94W
95W
94W
94W
94W
94W
94W
94W
94W
94W
94W
94W
95W
94W
94W
94W
94W
94W
94W
95W
94W
94W
95W
94W
94W
95W
94W
94W
95W

7
17
29
34
7
7
7
7
17
17
17
17
7
17
34
17
17
17
17
17
17
34
17
17
34
17
17
34
17
17
34

Remarks

---------------------------------------------------------------------------------------------------------

I

w
w
ex&gt;
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).

-

Date

·r .•

Eartag
no.

(1974)

Sex

Age

1-29
1-29
1-30
1-30
1-30
1-30
1-30
2-7
2-8
2-8
2-9
2-17
2-17

Female
Female
Female
Female
Male
Female
Male
Male
Female
Female
Female
Female
Female

Mature
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature

Summary:

45 Male Fawns;

L-2512
L-2513
L-2518
L...;,2519
L-2520
L-2521
L-2522
L-2614
L-2615
L-2616
L-2617
L-2672
L-2673
44 Female Fawns;

Ne ckb and
no.

Twp.

251
None
252
255
242
None
80
237
254
None
250
239
106

IN
15
IN
IN
IS
15
IS
IN
IN
IN
IN
IN
IN

15 Mature Males;

Location
Range

Sec.

95W
94W
95W
95W
94W
94W
94W
97W
97W
97W
97W
97W
97W

34
17
34
34
17
17
17
36
36
36
36
36
36

Remarks

108 Mature Females

Area 8 - White and Orange Block Neckband
1-29
1-29
1-29
1-30
1-30
1-30
1-30
1-30
1-30
1-30
1-31

Male
Female
Female
Male
Male
Male
Female
Male
Male
Female
Male

Fawn
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Fawn
Fawn

L-2514
L-2515
L-2517
L-2523
L-2524
L-2525
L-2528
L-2529
L-2526
L-2527
L-2531

94
'25
31
None
82
None
27
107
111
24
None

IS
IS
15
IS
IS
IS
15
IS
IS
IS
15

93W
94W
94W
93W
93W
94W
94W
94W
94W
94W
93W

5
14
14
5
8
14
14
14
34
34
5

-------------------------------------------------------------------------------------------------------------

I

w
w

\0
I

�Table 1. Record of deer trapped and marked on the winter ra~ge in Game Management Units 22, 23 and 11,
winter 197~-74 (continued).
Date
(1974)

Sex

1-31
1-31
1-31
1-31
1-31
1-31
1-31
1-31
1-31
1-31
1-31
1-31
1-31
1-31
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-1
2-2
2-2
2-2
2-2
2-2

Female
Female
Female
Female
Female
Female
Male
Male
Female
Female
Female
Male
Female
Female
Male
Male
Female
Female
Male
Female
Female
Female
Male
Male
Female
Female
Female
Female
Male
Male

Age

Eartag
no.

Neckband
no.

Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature

L-2532
L-2533
L-2534
1..-2535
L-2536
L-2537
L-2541
L-2540
L-2539
L-2530
L-2542
L-2538
L"""2543
L-2558
L-2559
L-2560
L-2561
L-2562
L:-2563
L-2564
.L-2571
L-2565
L-2566
L-2567
L-2568
L-2569
L-2570
L-2544
L-2545
L-2546

127
26
30
28
123
47
65
109
29
56
57
112
32
58
100
102
14
15
101
19
59
60
104
97
21
22
18
2
103
105

Twp.

IS
IS
IS
IS

1~
IS
IS
IS
IS
IN
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS

Location
Range

Sec.

93W
93W
93W
·94W
94W
94W
94W
94W
94W
93W
94W
94W
94W
94W
93W
93W
94W
94W
94W
94W
94W
94W
94W
94W
94W
93W
93W
94W
94W
94W

8
8
8
35
35
36
36
34
22
32
22
14
14
14
8
8
35
35
36
34
22
14
14
14
14
8
8
35
34
14

Remarks

I

IN

~

0
I

Found dead 4-25-74

------------------------------------------------------------------------------------------~-----------------

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
Date
(1974)

--

2-2
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-5

Sex
Female
Female
Female
Female
Female
Male
Female
Female
Male
Male
Female
Female
Male
Male
Male
Female
Female
Female
Female
Male
Female
Female
Female
Male
Female
Female
Female
Female
Female
Male

Age

Eartag
no.

Neckband
no.

Fawn
Mature
Mature
F~wn
Mature
Mature
Fawn
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn

L-2547
L-2548
L-2549
L-2550
L-255l
L-2552
L-2553
L-2554
L-2555
L-2556
L-2557
L-2587
L-2588
L-2589
L-2590
L-259l
L-2593
L-2594
L-2592
L-2595
L-2596
L-2597
L-2599
L-2598
L-2572
L-2573
L-2574
L-2575
L-2576
L-2577

1
55
13
20
10
90
None
3
92
95
8
51
106
108
96
23
None
4
5
87
16
7
None
None
52
None
17
50
11

63

Twp.
IS
IS
IS
IS
IS
IS
IN
IN
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IN
IS
IS
IS
IS

Location
Range
94W
94W
94W
94W
94W
94W
93W
93W
93W
93W
93W
93W
93W
93W
93W
94W
94W
94W
94W
94W
94W
94W
93W
93W
93W
93W
94W
94W
94W
94W

Sec.

Remarks

14
35
35
36
14
14
32
32
8
8
8
8
8
8
8
35
36
36
34
22
14
14
8
8

8
31
35
35
36
34

----------------------------------------------------------------------------------------------------------

I

w
.j::-o
t-'
I

�Table l. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued) •
Date
(1974)
2-5
2-5
2-5
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-6
2-7
2-7
2-7
2-7
2-7
2-7
2-7
2-7
2-7
2-7
2-8
2-8
2-8
2-8
2-8
2-8
2-8
2-9

Sex

Age

Eartag
no.

Ne ckband
no.

Twp.

Female
Male
Male
Male
Male
Female
Female
Female
Male
Male
Male
Male
Female
Female
Female
Female
Male
Male
Female
Male
Male
Male
Male
Female
Male
Female
Female
Female
Female
Male

Mature
Fawn
Fawn
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Mature
Matur:e
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn

L~2578
L-2579
L-2580
L-2581
L-2582
L-2583
L-2584
L-2585
L-2600
L-2601
L-2602
L-2603
L-2604
L-2605
L-2606
L-2607
L-2608
L-2609
L-2610
L-2611
L-2612
L-2613
L-2628
L-2629
L-2630
L-2631
L-2632
L-2633
L-2634
L-2635

54
None
88
85
86
36
43
53
79
93
113
None
None
48
34
38
115
124
12
119
83
80
71
139
68
62
46
40
37
110

IS
IS
18
IS
IS
18
IS
18
IN
IN
18
18
18
IS
18
IS
IS
IN
IN
18
IS
IS
IS
18
IN
IN
18
18
IS
18

Location
Range

Sec.

94W
94W
94W
93W
93W
93W
93W
93W
93W
93W
94W
94W
94W
94W
94W
94W
94W
93W
93W
93W
93W
93W
94W
94W
93W
93W
93W
93W
93W
94W

22
14
5
8
8
8
8
8
31
31
35
14
35
36
22
14
14
31
31
5
5
8
14
14
31
31
8
8
8
35

Remarks

--------~--------------------------------------------------------------------------------------------------

I

w
+'-

N
I

�Table l. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
Date
(1974)

--

2-9
2-9
2-9
2-9
2-9
2-9
2-9
2-9
2-10
2-10
2-10
2-10
2-10
2-10
2-11
2-11
2-11
2-11
2-12
2-12
2-12
2-13
2-13
2-14
2-14
2-14
2-14
2-14
2-15
2-15
2-16

Sex
Female
Female
Female
Male
Male
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Male
Female
Female
Female
Male
Female
Female
Female
Male
Male
Female
Female
Male
Female
Female

Age

Eartag
no.

Neckband
no.

Twp.

Fawn
Mature
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature

L-2636
L-2637
L-2638
L-2639
L-2640
L-2641
L-2642
L-2643
L-2644
L-2645
L-2646
L-2647
L-2648
L-2649
L-2650
L-2651
L-2652
L-2653
L-2654
L-2670
L-2671
L-2655
L-2656
L-2657
L-2658
L-2659
L-2660
L-2661
L-2662
L-2663
L-2665

33
45
49
64
67
145
148
141
42
151
39
181
6
129
128
152
72
35
135
44
117
136
140
127
126
118
41
134
116
138
133

IS
IS
IS
IS
IS
IS
IN
IN
IS
IS
IN
IN
IN
IN
IS
IS
IN
IN
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS
IS

.Location
Range

Sec.

94W
94W
94W
94W
94W
94W
93W
93W
94W
94W
94W
93W
93W
93W
94W
94W
93W
93W
94W
94W
94W
94w
93W
94w
94W
94W
93W
93W
94W
93W
94W

36
36
36
34
34
14
31
31
35
34
36
31
31
31
14
14
31
31
36
14
14
36
26
36
36
36
26
26
36
26
36

Remarks

--------------------------------------------------------------------~------~-------------------------------

I

w
~
w
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
Location
Range

Sec.

IS
IS
15

94W
94W
93W

36
36
36

5

15

93W

36

146
74
130
154
137
147
144
143
77
122
149
153
150
69
76
121
73
132
120
66
78
70
131

15
IS
IN
15
15
15
15
15
15
15
15
15
15
IS
15
15
15
15
IS
15
15
15
15

94W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W

36
36
19
26
26
36
36
26
36
36
36
36
36
26
26
36
36
26
36
36
36
36
36

Date
(1974)

Sex

Age

Eartag
no.

Neckband
no.

Twp.

2-17
2-17
2-17

Female
Male
Male

Mature
Fawn
Fawn

L-2666
L-2667
L-2694

142
None
23

2-17

Female

Unknown

L-2695

2-18
2-18
2-18
2-20
2-20
2-20
2-20
2-21
2-21
2-21
2-21
2-21
2-22
2-23
2-23
2-23
2-26
2-26
2-28
2-28
3-5
3-6
3-8

Fetnale
Male
Female
Female
Female
Female
Female
Female
Male
Male
Female
Female
Female
Female
Female
M...ale
Female
Female
Female
Female
Female
Female
Female

Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Hature
Mature
Hature
Hature
Mature
Mature
Mature
Mature
Mature

L-2668
L-2669
L-2679
L-2706
L-2707
L-2709
L-2708
L-2680
L-2681
L-2682
L-2683
L-2715
L-2717
L-2726
L-2727
L-2728
L-2744
L-,z745
L-2758
L-2759
L-2773
L-2775
L-2778

5unnnary: 35 Male Fawns;

31 Female Fawns;

21 Mature Males;

81 Mature Females;

Remarks

Has a yellow and blue
neckband.
Has a yellow and blue
neckband.
I

w
.p.pI

1 UnknoWn Female

----------------~---------------------------------------------------~----------------------~----------------

�Table 1- Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
Date
(1974)

Sex

Age

Eartag
no.

Neckband
no.

Twp.

Location
Range

Sec.

Remarks

Area 9 - Yellow and Blue Block Neckband
2-13
2-13
2-14
2-14
2-15
2-15
2-15
2-15
2-15
2-15
2-16
2-16
2-16
2-16
2-17
2-17
2-17
2-17
2-17
2-18
2-18
2-18
2-18
2-18
2-18
2-19
2-19
2-19
2-19

Female
Female
Hale
Male
Female
Hale
Female
Female
Female
Female
Female
Male
Female
Female
Male
Female
Female
Female
Male
Female
Female
Male
Female
Female
Female
Female
Female
Female
Female

Mature
Hature
Fawn
Fawn
Mature
Fawn
Ha tur e

Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature

...;.---------------~--------.--..~

L-26l9
L-2620
L-2621
L-2622
L-2623
L-2624
L-2625
L-2626
L-2627
L-2664
L-2684
L-2685
L-2686
L-2687
L-2688
L-2689
L-2690
L-2691
L-2693
L-2674
L-2675
L-2676
L-2677
L-2678
L-2696
L-2697
L-'2698
L-2699
L-2700

14
12
18
None
6
19
4
3
1
2
10
25
11
16
24
None
8
15
26
45
7
35
42
43
13
39
46
37
40

IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
1N
IN
1N
1N
1N
1N
1N
1N
1N
1:N
1N
1N
1N
1N
1N
1N

94H
94H
94H
94H
94W
94H
94W
94H
94W
94W
94W
94W
94H
94H
94W
94W
94W
94W
94W
94W
94W
94W
94H
94H
94H
94W
94W
93W
93W

28
28
20
20
28
20
20
20
29
18
20
28
28
28
28
20
29
29
18
20
20
28
28
29
18
18
20
19
19

I

w
+:U1
I

Found dead 3-12-74

-----------------------------------------------------------------------------

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
Date
(1974 )
2-19
2-19
2-20
2-20
2-20
2-20
2-20
2-21
2-21
2-21
2-22
2-22
2-22
2-22
2-22
2-23
2-23
2-23
2-23
2-23
2-23
2-24
2-24
2-24
2-24
2-24
2..;.25
2-25
2-25
2-25
2.,...25

Sex

Age

Eartag
no.

Ne ckb and
no.

Female
Female
Female
Female
Male
Female
Female
Female
Female
}fale
Female
Female
Hale
Femal,.e
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
l1ale
Female
Female
Male
Female

Mature
Mature
Fawn
Fawn
Mature
}fature
Fawn
Mature
Mature
Nature
Mature
}fature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Ha t ur e
Mature
Mature
Mature
Mature
.Mature

L-270l
L-2702
L-2703
L-2704
L-2705
L-2710
L-2711
L-2713
L-2714
L-2716
L-2718
L-2719
L-2720
L-272l
L-2722
L-2723
L-2724
L-2725
L-2729
L-2730
L-273l
L-2732
L-2733
L-2734
L-2735
L-2736
L-2737
L-2738
L-2739
L-2740
L-274l

41
48
None
50
29
44
38
47
49
22
78
74
34
72
73
75
85
84
83
82
81
77
71
80
76
31
28
79
105
32
90

Twp.
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN

IN
IN
IN
IN
IN
IN
IN
IN
IN

Location
Range
93W
93W
94W
94W
94W
93W
94W
94W
93W
93W
93W
93W
93W
94H
94W
94W
94H
94W
93W
93W
93\oJ'
93W
93W
93W
93W
94W
94W
93ToJ'
93'(0]

93W
93W

Sec.

Remarks

19
19
20
29
29
19
18
20
20
19
19
19
19
20
29
28
28
28
19
19
19
19
19
19
19
18
18
19
19
19
19

-----------------------------------------------------------------------------------------------------------

I

w

~

'"
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
Date
(1974)

Sex

2-25
2-25
2..•..
26
2-26
2-26
2-26
2-26
2-26
2-27
2-27
2-27
2-27
2-27
2-27
2-28
2-28
2-28
2-28
3-1
3-1
3-1
3-1
3-2
3-4
3-4
3-4
3-4
3-5
3-6
3-6
3-8

Male
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Male
Male
Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female

Age
Hature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
M..ature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
l'1ature
Mature

Eartag
no.
L-2742
L-2743
L-2746
L-2747
L-2748
L-2749
L-2750
L-275l
L-2752
L-2753
L-2754
L-2755
L-2756
L-2757
L-2760
L-276l
L-2762
L-2765
L-2763
L-2764
L-2766
L-2767
L-2768
L-'-2769
L-2770
L-277l
L-2772
L-2774
L-2776
L-2777
L-2779

Neckband
no.

Twp.

33
101
92
97
94
93
98
21
104
95
88
87
103
30
68
61
54
52
55
63
102
57
100
60
59
58
20
53
56
,51
62

IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
iN
IN
IN
IN
IN
IN

Location
Range
93W
94~.J

93W
93W
93W.
93W
93W
94W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W
93W

Sec.

Remarks

19
18
20
19
19
19
19
18
20
19
19
19
19
19
20
19
19
19
19
19
19
18
19
19
19
19
18
19
19
19
19

-------------~-------------------------~---~---------------------------------------------~---------------

I
\,.0.)

.p-

-..J
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 22, 23 and 11,
winter 1973-74 (continued).
Date
(1974)

Sex

Age

Eartag
no.

Neckband
no.

Twp.

3-8
3-8
3-8

Female
Male
Female

Mature
Fawn
Mature

L-2788
L-2789
L-2787

69
70
64

IN
IN
IN

Summary:

7 Male Fawns;

Grand
Summary: 108 Male Fawns;

5 Female Fawns;

12 Mature Males;

Location
Range

Sec.

93W
93W
93W

Remarks

19
19
18

70 Mature Females

I

86 Female FaWnS;

56 Mature Males;

290 Mature Females;

1 Unknown Female

W
.j::--

co
I

Blue and White Neckbands
2-11-73
2-11-73

Female
Female

Mature
Mature

Corrections to 1973 Trapping Records
(Corrections are Underlined)
L-2084
L-2085

27

28

2S
2S

97W
97W

3
3

May be neckband No. 24
May be neckband No.3

IS
IS

95W
94W

6
17

Green &amp; white neckband
Blue &amp; white neckband

.Recaptures
1-14-74
1-21-74

Male
Female

Mature
Mature

L-1681
L-2110

75
25

�-349-

Table 2. 8ightings of deer marked on the winter range in Game Management
Unit 22, March 16, 1973 through February 28,1974 plus several earlier
sightings reported after March 15, 1973.

Ne ckband

Date
(1973)

Color

Number

Twp.

2-01
2-11
2-11
2-11
3-16
3-16
3-16
3-16
3-16
3-16
3-16
3-16
3-16
3-16
3-17
3-23
3-24
3-24
3-24
4-09
4-09
4-11
4-11
4-11
4-11
4-11
4-12
4-12
4-14
4-14
4~14
4-14
4-14
4-14
4-14
4-14
4-14
4-14
4-14
4-15
4-15
4-15
4-15

White
Pink
Pink
Pink
Blue and white
Red w/white stripe
Red w/white stripe
Blue and white
Blue and white
Blue and white
Blue and white
Blue and white
Red w/white stripe
Yellow wIred stripe
White
White
White
White
White
Blue
Blue
Pink
Pink
Pink
Blue and white
White
Yellow wIred stripe
Pink
White
White
White
Blue and white
Redw/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Pink
Blue and white

55
59
89
75
3
128
107
8
34
1
105
90
109
1
124
87
160
136
72
19
79
59
89
75
103
156
11
45
62

18
38
38
38
28
28
28
28
28
28
28
28
28
18
18
28
18
28
18
2N
2N
28
28
28
18
1N
1N
28
18
18
18
28
28
28
28
28
28
28
28
28
28
28
18

?
?

34
75
107
65
57
43
94
69
55
29
34
27

Location
Range
8ec.

98W
98W
98W
98W
97W
97W
97W
97W
97W
97W
97W
97W
97W
96W
97W
97W
97W
98W
98W
98W
98W
96W
96W
96W
97W
96W
96W
98W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W

2
5
5
5
16
16
16
9
9
4
4
4
4
4
32
9
31
1
33
3
3
32
32
32
21
9
26
24
28
28
28
5
5
16
16
16
16
21
22
36
36
36
21

---------------------------------------------------------------------------------

�-350-

Table 2. Sightings of deer marked on the winter range in Game Management
Unit 22, March 16, 1973 through February 28, 1974 plus several earlier
sightings reported after March 14, 1973 (continued).

Date
(1973)

Color

Number

Twp.

Location
Range

4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-15
4-16
4-16
4-16
4-16
4-16
4-16
4-16
4-16
4-16
4~16
4-16
4-16
4-16
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17
4-17

Blue and white
Blue and white
Blue and white
White,
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Yellow wIred stripe
Pink
White
White
White
White
White
White
White
White
White
White
White
White
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Pink
Red w/white stripe
Pink
Pink
Pink
Pink
Pink
Pink
Pink
Pink

53
55
21
84
126
29
19
16
37
110
48
106
58
33
117
7
27
6
84

IS
IS
IS
IS
2S
2S
2S
2S
2S
3S
3S
3S
3S
3S
3S
3S
3S
IS
2S
IN
IN
IN
IS
IS
2S
2S
2S
2S
2S
25
25
25
25
25
25
25
25
25
25
25
2S
2S
25
25
25
25
25

97W
97W
97W
97W
97W
97W
97W
97W
97W
96W
96W
96W
96W
96W
96W
95W
95W
96W
97W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
97W
97W
97W
97W
97W
97W
96W
97W
97W
96W
96W
97W
97W
98W
97W
97W

Neckband

?
?

188
187
210
?
?
?
?
?
?
?

129
126
32
107
47
29
89
57
55
72
32
45
88
52
7
30

Sec.

28
28
28
28
16
16
26
29
36
4
3
3
3
3
11
9
9
13
16
36
36
36
20
29
29
29
29
29
29
29
29
16
16
36
16
36
36

31
36
36
21
21
19
19
24
29
29

-----------------------------------------------------------------------------------

�-351-

Table 2. Sightings of deer marked on the winter range in Game Management
Unit 22, March 16, 1973 through February 28, 1974 plus several earlier
sightings reported after March 15, 1973 (continued).

Date
(1973)

Color

4-17
4-18
4-21
4-21
4-21
4-22
4-22
4-22
4-22
4-22
4-23
4-23
4-23
4-23
4-23
4-23
4-23
4-23
4-23
4-23
4-24
4-24
4-24
4-24
4-25
4-26
4-26
4-26
4-26
4-26
4-26
4-26
4-26
4-26
4-27
4-27
4-27
4-27
4-27
4-27
4-27
4-27
4-27
4-27

White
Blue
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Pink
Pink
Pink
Pink
Pink
Blue and white
Pink
Pink
Pink
Pink
Pink
Pink
Pink
Pink
Green w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
White
White
White
White
White
White
White
White
Blue
Blue
Blue and white
Blue and white
Blue and white
Blue and white
Blue and white
Blue and white
Blue
White
Blue
Blue

Neckb and

Number

Twp.

34

2S
2N
IS
IS
IS
2S
2S
2S
3S
2S
IS
2S
2S
3S
3S
3S
3S
3S
3S
IN
IN
IN
IN
IN
2S
IS
IS
2S
IS
IS
2N
2N
2N
2N
IS
IS
IS
IS
IS
IS
2N
2N
2N
2N

?

10
36
24
23
63
36
41
26
28
44
60
90
66
94
73
64
40
?

75
31
2
?

181
160
115
175
115
239
29
1
107
?

43
68
22
5
84
36
104
48
39
77

Location
Range
Sec.

97W
94W
96W
96W
96W
97W
97W
97W
98W
97W
97W
97W
97W
96W
96W
96W
96W
96W
96W
95W
96W
96W
96W
96W
98W
97W
98W
98W
98W
98W
98W
98W
99W
99W
97W
97W
97W
97W
97W
97W
98W
97W
98W
98W

20
31
10
10
10
29
32
29
1
31
31
27
27
3
3
3
3
3
3
35
36
27
16
27
10
32
29
19
29
21
11
3
9
4
11
11
11
11
21
21
6
34
6
6

---------------------------------------------------------------------------------

�-352-

Table 2. Sightings of deer marked on the winter range in Game Management
Unit 22, March 16, 1973 through February 28, 1974 plus several earlier
sightings reported after March 15, 1973 (continued).

Neckband

Date
(1973)

4-27
4-27
4-27
4-27
4-27
4-27
4-27
4-27
4-27
4-28
4-28
4-28
4-28
4-28
4-28
4-29
4-29
4-29
4-29
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-01
5-02
5-02
5-02
5-02
5-02
5-02
5-02
5-02
5-02

L~

Color

Blue
Blue
Blue and white
B1\1eand white
Blue and white
Blue and white
Blue and white
Blue and white
Blue and white
Pink
White
Blue and white
Blue and white
Blue and white
Yellow wIred stripe
Blue and white
Blue
Blue
Blue
White
Pink
Red w/~hite stripe
(;
Pink
Red w/white stripe
Red w/white stripe
Red w/white stripe
Pink
Pink
Pink
Pink
Pink
Pink
Red w/white stripe
Red w/white stripe
Pink
Pink
Red w/white stripe
Red w/white stripe
Blue and white
Red w/white stripe
White
White
White
Blue and white
Blue and white

',,~

"

Number

Twp.

40
42
104
132
54
131
59
16
65
8

2N
2N
IS
IS
IS
IS
IS
IS
IS
2S
2S
IS
IS
2S
IS
2S
2N
2N
IN
IS
2S
2S
3S
3S
3S
3S
3S
2S
2S
2S
2S
2S
2S
2S
2S
2S
2S
2S
2S
2S
IS
IN
IN
2S
2S

?
?
?
?

6
66
41
103
96
148
3
123
18
101
125
66
39
3
4
72
15
42
130
12
24
57
128
43
3
70
160
89
?

95
47

Location
Range
Sec.

98W
98W
97W
97W
97W
97W
97W
97W
97W
97W
97W
96W
96W
97W
96W
96W
98W
98W
99W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
96W

5
5
21
11
11
11
11
15
28
29
29
33
32
1
12
9
9
5
2
21
35
35
2
2
2
2
2
35
35
35
35
35
36
36
36
36
16
16
16
16
32
21
21
11
9

---------------------------------------------------------------------------------

�-353-

Table 2. Sightings of deer marked on the winter range in Game Management
Unit 22, March 16, 1973 through February 28, 1974 plus several earlier
sightings reported after March 15, 1973 (continued).

Neckband

Date
(1973)

€olor

5-02
5-02
5-03
5-03
5-04
5-04
5-04
5-04
5-04
5-04
5-04
5-04
5-04
5-04
5-04
5-04
5-05
5-05
5-05
5-07
5-09
5-09
5-09
5-11
5-11
5-12
5-13
5-13
5-15
5-17
5-23
5-30
5-30
5-?
6-05
6-21
6-26
6-26
6-30
7-03
7-09
7-?
7-?
7-?
7-?
8-07

White
Blue and white
Yellow wIred stripe
Blue and white
Pink
Red w/white stripe
Pink
Pink
Pink
Pink
Pink
Pink
Pink
Red w/white stripe
Red w/white stripe
No neckband
Yellow wIred stripe
Green w/white stripe
Greenw/white stripe
Green w/white stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Red w/white stripe
Red w/white stripe
Blue and white
Green w/white stripe
Green w/white stripe
Blue and white
Blue and white
Pink
Green w/white stripe
Pink
Pink
Red w/white stripe
White
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Blue and white
Blue and white
Blue and white
Red w/white stripe
Red w/white stripe
Red w/white stripe

Number

Twp.

62
12
?
77
60
52
37
10
54
46
11
31
44
124
22
Eartag L-1956
6
49
36
25
20
18
39
111
86
?
?
?
85
?
?
?
?
94
?
?
33
92
?
?
?
?
?
?
?
?

lS
lS
2S
lS
2S
2S
2S
3S
3S
3S
3S
3S
3S
2S
2S
2S
1N
lS
lS
1N
lS
lS
lS
2S
2S
4S
1N
1N
4S
3S
2S
3S
4S
3S
2S
2S
2S
2S
2S
2S
9N
2S
2S

IS
lS
2S

Location
Range

Sec.

97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
95W
96W
96W
96W
96W
96W
96W
95W
95W
96W
96W
95W
98W
97W
97W
96W
96W
96W
90W
90W
94W
94W
94W
94W
89W
94W
94W
93W
93W
94W

32
15
26
15
27
36
36
8
8
8
8
8
3
26
25
25
31
14
14
26
10
10
10
32
32
6
36
32
2
11
29
36
4
5
23
23
23
26
23
3
6
11
11
17
17
23

---------------------------------------------------------------------------------.
.

�-3S4-

Table 2. 8ightings of deer marked on the winter range in Game Management
Unit 22, March 16, 1973 through February 28, 1974 plus several earlier
sightings reported after March 15, 1973 (continued).

Date
(1973)

Color

Number

Twp.

8-07
8-19
8-22
8-26
8-26
8-?
9-18
9-24
9-24
9-24
9-24
9-24
9-24
9-24
9-24
9-27
9-28
9-29
9-29
10-5
10-11
10-11
10-12
10-12
10-12
10-13
10-13
10-13
10-13
10-13
10-13
10-13
10-13
10-14
10-14
10-14
10-14
10-14
10-15
10-15
10-15
10-15
10-15
10-lS
10-16
10-16
10-17
10-17

Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Pink
Yellow wIred stripe
Pink
White
Blue
Blue
Blue
White
Pink
Pink
White
Blue and white
Blue and white
Blue and white
Blue
White
Blue
Blue and white
White
Blue
White
Pink
Pink
Pink
Red w/white stripe
White
Blue
Blue
Blue
White
White
Blue
Red w/white stripe
White
Pink
Blue
White
Blue
White
White
Red w/white stripe
White
Pink
Blue and white

?
?
?
123
?
?
2
?
?
?
?
?
?
36
27
?
?
?
?
?
?
?
?
?
?
?
?
52
?
?
?
?
?
?
?
?
?
60
?
?
?
?
?
223
?
?
?
?

28
S8
58
58
58
58
18
S8
48
48
48
S8
48
68
68
IN
IN
18
48
2N
3N
48
58
18
18
38
38
58
S8
58
5N
5N
28
28
28
2N
48
18
38
2N
2N
2N
38
38
28
2N
38

Neckband

Location
Range

8ec.

94W
94W
94W
95W
95W
97W
90W
97W
99W
99W
99W
97W
99W
97W
97W
92W
92W
93W
99W
86W
93W
9SW
96W
91W
91W
94W
94W
97W
97W
97W
91W
91W
94W
99W
99W
95W
96W
99W
96W
94W
94W
94W
97W
97W
94W
99W
96W
90W

23
31
21
14
14
23
9
17
19
19
19
17
16
5
36
3
23
36
15
4
28
33
26
18
18
2S
25
1
1
1
34
34
4
3
3
16
.9
32
9
4
3
17
14
27
18
31
36
9

IS
--------------------------~-------------------------------------------------------

�-355-

Table 2. Sightings of deer.marked on the winter range in Game Management
Unit 22, March 16, 1973 through February 28, 1974 plus several earlier
sightings reported after March 15, 1973 (continued).

Date
(1973)

Color

Ne ckband

10-17
10-18
10-20
10-20
10-21
10-24
10-24
10-25
10-25
10-29
10-30
10-?
10-?
10-?
12-12
12-12
12-12
12-12
12-12
12-12
12-13
12-13
12-19
12-20
12-20
12-21
12-21
12-21 .
12-21
12-21
12-28
12-28
12-28
12-28
12-28
12-28
12-31
1-10-74
1-10-74
1-10-74
1-10-74
1-10-74
1-10-74
1-11-74
1-11-74
1-11-74
1-11-74

White
Pink
Blue
Red w/white stripe
White
Red w/white stripe
Blue
Pink
Red w/white stripe
White
Red w/white stripe
Red w/white stripe
Red w/white stripe
Blue
White
White
White
White
Yellow wIred stripe
Yellow wIred stripe
Green w/white stripe
Yellow wIred stripe
Blue and white
White
White
White
White
White
Blue and white
Blue and white
Blue
Blue
White
White
Red w/white stripe
Pink
White
Blue and white
Blue and white
Blue and white
Blue and white
Blue and white
Red w/white stripe
Yellow wIred stripe
Yellow wIred stripe
Blue and white
Blue and white

Number

Twp.

Location
R~nge

Sec.

?
?
?
?
?
?
?

2S
IS
IS
2S
2S
IS
IS
4S
4S
4N
4S
3S
4S
IN
IS
IS
2N
2N
IN
IN
IS
IS
2S
IS
IS
3S
3S
3S
2S
2S
2N
2N
IN
2N
3S
3S
IN
IS
IS
2S
2S
2S
2S
IS
IS
2S
IS

99W
96W
98W
92W
97W
90W
90W
96W
96W
89W
94W
97W
97W
93W
97W
97W
98W
98W
96W
96W
95W
95W
96W
99W
99W
99W
99W
99W
96W
96W
99W
99W
98W
98W
94W
97W
96W
96W
97W
97W
97W
97W
97W
96W
96W
97W
97W

34
5
9
2
31
9
9
5
21
16
1
13
23
20
9
9
10
10
36
36
6
6
2
19
19
3
3
3
1
2
16
14
35
16
8
2
7
20
35
3
3
3
15
29
29
3
34

3

?
?
?
?
?
?
157
47
14
?
2
4
?
?
104
?
?
?
?
?
44
88
112
?
?
?
?
72
?
?
?
?
?
?
?
?
?
?
?

---------------------------------------------------------------------------------

�-356-

Table 2. Sightings of deer marked on the winter range in Game Management
Unit 22, March 16, 1973 through February 28, 1974 plus several earlier
sightings reported after March 15, 1973 (continued).

Date
(1974)

Color

1-11
1-11
1-11
1-14
1-20
1-21
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-22
1-24
1-24
1-25
1-25
1-25
2-05
2-07
2-10
2-11
2-15

Blue and white
Red w/white stripe
Red w/white stripe
Green w/white stripe
Yellow wIred stripe
Blue and white
Blue and white
Blue and white
Blue and white
Blue and white
Blue and white
Blue and white
Redw/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Pink
Pink
Blue
White
Blue and white
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Blue and white
Blue and white
Yellow wired stripe

Neckband
Number

Twp.

Location
Range

Sec.

?
?
?

IS
2S
2S
IS
IS
IS
IS
IS
IS
2S
2S
2S
2S
2S
2S
3S .
3S
2S
3S
3S
2N
2N
IN
IN
IN
IN
IN
IS
IS
IS

97W
97W
97W
95W
96W
94W
97W
97W
97W
97W
97W
97W
97W
97W
97W
96W
95W
97W
97W
97W
98W
99W
97W
96W
97W
97W
97W
97W
96W
96W

34
10
10
6
14
17
12
12
12
3
3
3
10
10
11
11
2
24
2
2
8
36
28
30
25
36
36
14
3
11

75
?

25
?
?
?
?
?
?
?
?
?
?
?
?

57
?
?
?
?
?

?

74
72
66
?

26

�-357-

Table 3. Recoveries of deer marked on the winter range in Game Management
Unit 22, April, 1973 through February, 1974.

Date of
Recovery
(1973)
4-02
4-11
4-11
4-24
4-26
4-26
4-29
4- ?
5-02
5-04
5-09
5-09
5-11
5- ?
5- ?
5- ?
5- ?
6- ?

7- ?
8-18
8- ?

10-12
10-13
10-13
10-13
10-13
10-13
10-13
10-13
10-13
10-13
10-14
10-14
10-14
10-14
10-14
10-14
10-14
10-14
10-15
10-15
10-15
10-15

Neckband
Color

No.

116
Red w/white stripe
126
White
28
Pink
130
White
86
Blue and white
13
Yellow wIred stripe
40
Blue and whi te
62
Pink
40
Red w/white stripe
27
Blue and white
81
Blue
238
White
25
Red w/white stripe
110
Red w/white stripe
No neckband - eartag L-2013
8
Blue and white
189
White
32
Blue and white
51
Greenw/white stripe
?
White
7
Blue and white
114
Blue
130
Red w/white stripe
122
Red w/white stripe
No neckband - eartag L-2046
166
White
46
Green w/white stripe
50
Green w/white stripe
7
Yellow wIred stripe
45
Red w/white stripe
106
Blue
169
White
104
Blue
180
White
No neckband - eartag L-2009
106
Red w/white stripe
L-2022
No neckband - ear tag
11
Red w/white stripe
99
White
No neckband - eartag L-2312
31
Pink
13
Green w/white stripe
24
White

Twp.

Location
Range Sec.

3S
IS
2S
2S
2S
IS
IS
2S
2S
IS

96W
97W
97W
98W
97W
96W
97W
97W
97W
97W

3

32
30
11
4

14
11
29
25
21

?

?

?

IS
3S
3S
3S
2S
2S
IS
IS
5S
IS

98W
95W
96W
96W
97W
99W
96W
96W
94W
97W

16
9
4
5

4

25
29
3

27
15

?

?

?

5S
5S
5S
IS
IS
IS
IS
3S

94W
94W
94W
99W
96W
96W
92W
91W

26
18
26
16
11
11
12
22

?

?

?

3S
2N
IS
5S
3S
3S
IS
3S
IS
5S
2N
4S

98W
92W
98W
94W
95W
94W
94W
96W
96W
97W
92W
95W

12

Remarks

Died soon after marking
Died in fence
Died soon after marking
Died in fence
Died soon after marking
Died soon after marking
Died soon after marking
Died soon after marking
Died soon after marking
Died soon after marking
Only head found
Died soon after marking
Died soon after marking
Road-kill
Died soon after marking
Died soon after marking
Died in fence
Died soon after marking
Found dead
Found dead
Died soon after marking

Found dead
Found dead

4

2

Found dead

17
17
1

17
27
27

Died soon after marking
Died soon after marking

3

16
36

Found dead

----------------------------------------------------------------------------------

�-358-

Table 3. Recoveries of deer marked on the winter range in .Game Management
Unit 22, April, 1973 through February, 1974 (continued).
Date of
.Recovery
(1973)

Color

10-16
10-16
10-16
10-17
10-17
10-17
10-17
10-17
10-18
10-18
10-18
10-19
10-19
10-19
10-19
10-20
10-20
10-20
10-20
10-21
10-21
10-21
10-22
10-31
10- ?
10- ?
10- ?
10- ?
10- ?
10- ?
10- ?
10- ?
10- ?
10- ?
10- ?
10- ?
11- ?
2-26-74

Pink
Green w/white stripe
Blue
Blue
Pink
Blue
Blue and white
Pink
No neckband - eartag
Pink
No ne ckband - eartag
White
Yellow wired stripe
White
Green w/white stripe
Blue
Green w/white stripe
Red w/white stripe
Red w/white stripe
White
White
White
Green w/white stripe
Red w/white stripe
No neckband - eartag
Blue and white
White
Pink
White
Blue and white
Blue and white
Red w/white stripe
Blue
Green w/white stripe
Blue
Blue
Blue
Red w/white stripe

Neckband
No.

Twp.

77
44
114
108
10
5
23
12
L-2065
65
L-2044
28
29
81
17
132
30
112
128
226
148
7
55
85
L-2010
11
134
80
77
47
77
125

48
18
2N
1N
58
38
28

Location
Range 8ec.

94W
95W
98W
99W
96W
96W
93W

31
32
12
34
36
36
28

?

?

?

18

96W

31

?

?

?

18
1N
28
1N
18
1N
1N
38
78
28
38
2N
18
2N
48
48
48
48
2N
18

96W
96W
·95W
97W
95W
lOOW
96W
95W
97W
98W
98W
97W
96W
91W
97W
97W
97W
95W
93W
95W

31
16
28
8
32
16
18
2
35
7
26
1
13
12
3
3
3
20
24
36

?

?

?

48

97W

23

?

?

?

?

52
50
95
90
51

18
2N
5N
2N
28

92W
98W
91W
98W
97W

10
5
34
5
24

Remarks

Found dead
Found dead

Died soon after marking
Died soon after marking
Died soon after marking

Died soon after marking

Found dead
Road-kill
Found dead

�GAME

MANAGEMENT UNIT

22

-PICEANCE-

~

•....

..••.

-.

r
\ ---,-~":".:. '.'

I

W
VI
\0

I
\
q

I

i,

\

~\
'l

Deer Track Density

1
"

\

•

I "--..r
I

\...

r=-----] Little
r:::::::::;:::::::::::l

or none

::::::::::.:.:.:.:.;.;
Moderate
" ... ····,·0

I

I Heavy

i/

/'

/'

v.

.Q

I

I

•

::'

! :.~
..

,./

••

t·::

/

) //',--,/j

I~/?

II,
,/

--

'

.

'( ( _

_-\.-

•••••••••• I

...•

I,

"Fig.

1.

Deer distribution

on the Piceance winter range, December 19, 20 and 21, 1973.

�GAME

MANAGEMENT UNIT

22

-PICEANCE-

~

•....

.•..

r

L
I
\

I

W
C'\

o
I

n
~I
I

\~\
"1

l

"\
\

Deer Track Density

''\

[_..
••

(

•• 0_ ••••••••

-] Little or none
,

Heavy

Fig. 2.

Deer distribution

on the Piceance winter range, December 31, 1973 and January 2 and 3. 1974.

�GAME

MANAGEMENT UNIT

22

-PICEANCE-

~

..

,,,

.•..

I

W
0\

•...
I

Fig. 3.

Deer distribution on the Piceance winter range, February 12 and 13, 1974.

��-363July, 1974
JOB PROGRESS PYPORT

State

COWRADO

0f

Proj cc.tNo.

~38-R-28

Work Plan No.

16

:

Deer-Elk Investigations
Job No.

2

--------

Job Title

Piceance Deer Study - Population Density and Structure

Period Covered:

September 1, 1973 through January 31, 1974.

Personnel:

R. M. Bartmann, S. F. Steinert, J. J. Klein, Jr., and C.E.

White.

ABSTRACT
The 1973-74 winter deer count was made January 20-25. Eight hundred eightynine deer were counted on 120 1/4-square mile (l/2-mile by 1/2~mile) quadrats
for a mean 8.2 deer per quadrat or 32.8 deer per square mile. The projected
total deer population on the Piceance winter range was 21,812 ± 4,256 deer at
the 90 percent confidence level. Visibility of quadrat markers painted with
non~reflective orange paint was poor except in bright sunlight, so all such
markers will have to be repainted with fluorescent paint. Post-season
classifications on the Piceance winter range extended from December 11, 1973
through January 10, 1974 because of weather delays. The 1,597 deer classified,
which included 244 from Northwest Region .counts, yielded abuck:doe:fawn
ratio
of 22:100:63. Resulting composition of the total winter population was 2,596
adult males, 11,778 adult females and 7,438 faWns. A revised estimate for the
1972-73 winter population was calculated by working the population equation
backwards using the 1973-74 winter population as a base. The revised estimate
was 29,898 deer, which was 10,214 animals greater than originally figured.

��-365-

PICEANCE DEER STUDY - POPULATION DENSITY AND STRUCTURE
Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.
SEGMENT OBJECTIVES
1.

To establish mule deer population density and sex and age structure
estimate techniques for the Piceance winter range.
a.

Estimate the size of the total wintering deer population in Game
Management Unit 22.

b.

Estimate the sex and age structure of the wintering deer population
in Game Management Unit 22.

METHODS AND MATERIALS
Deer Density
A preliminary evaluation of quadrat size and sample size was presented in
the previous segment report (Bartmann 1973). Based on this work, a 1/4square mile quadrat (1/2-mile by 1/2-mile) was selected and the original
60 quadrats were increased to 120, all randomly selected on the winter
range (Fig. 1). The corners of all quadrats were marked with 1 x 2-foot
pieces of lIB-inch tempered masonite placed in tree tops whenever possible,
on tall shrubs, or on fence posts. About one-third of the markers were
painted with a fluorescent red-orange paint, while a commercially available
bright orange, non-reflective paint was used on the remainder.
Deer were counted as soon as winter deer distribution conformed fairly close
with pre-determined limits. Two observers were used; one to count and the
other to count and navigate. Aerial photos of each plot were carried in the
helicopter to help insure staying within plot bounds.
The perimeter of a plot was flown first to locate corners and help orient
the observers with the plot boundaries. Deer on or near the edge were
classed as either in or out of the plot at that time. The pattern flown
over the plot interior varied according to terrain and cover features,
which were used to insure complete coverage and help keep track of deer
already counted.

�4'.:--- ;~,---

~\«:~.
a3/..I.).! .
,~7/9
!1/~~,,,

".

I

r
L.__ ~

f. ~

22

-PICEANCE-

~~

"...

..

•

....,.~

•

MMhr

•

'.,

.

.
"It _.

Fig. 1.

MANAGEMENT UNIT

!

\ .-/_.-/, ~/'-'. .~ Ii...JI /
I ,'''- ---

\

GAME

"

,.

_ , ~.

-

Locations of !-square mile deer census quadrats on the Piceance winter range.

J

W
00J

�-367-

Population Structure
The composition of the winter deer population is estimated from postseason sex and age classifications made from a helicopter. Northwest
Region personnel also conduct such classifications in the southern part
of Game Management Unit 22 and work is coordinated with them to avoid
duplication of effort. Both sets of data are later combined to estimate
the composition for the entire winter range. A minimum goal of 2,000
animals classified is set for the entire winter range with any ground
counts being additional. Animals are classified as bucks) does, fawns,
or unknown. Check station data are used to estimate the fawn sex ratio
and percent yearling females.
RESULTS AND DISCUSSION

Population Density
The deer count was started on January 11, 1974 but was aborted after 20
quadrats were completed due to logistical problems. The count was resumed
January 20 and completed January 25. The 20 quadrats were recounted as
deer distribution had changed during the interim.
Nine hundred eighty-nine deer were counted on 120 1/4-square mile quadrats
for a mean 8.2 deer per quadrat with a range of 0 to 48 (Table 1). This
projects to 32.8 deer per square mile or 21,812 ± 4,256 deer on the Piceance
winter range at the 90 percent confidence level.
The interval estimate is wider than desired but is still within ± 20 percent of the mean. It is presumed that the change in deer distribution
between the time of the first count attempt and the actual count produced
increased variability. Deer were forced out of the higher wintering areas
and a greater than expected number of plots (43) had no deer. This is
partially demonstrated by comparison of the duplicate counts on the 20
plots. During the first count, only one out of 20 plots contained no deer.
During the repeat count about one and one-half weeks later, six plots contained no deer with all but one of these situated near the upper winter range
limit.
The pilot, Mr. Ben Morris, did an exceptionally good job, particularly in
keeping himself oriented once he was shown the plot boundaries. This took
a large part of the job away from the "navigator" allowing him to concentrate
more on looking for deer.
Notes were also made on visibility of corner markers. Some markers need to
be repositioned and all markers painted with non-reflective paint will have
to be repainted with fluorescent paint. The regular orange paint was
satisfactory in bright sunlight but hard to see in the.shadows.

�-368-

Table 1.
Number of deer counted on quadrats on the Piceance
in Game Management Unit 22 during January, 1974.

winter

range

Quad.

Deer

Quad.

Deer

Quad.

Deer

Quad.

Deer

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30

11
7
24
6
11
4
19
0
0
0
0
6
0
7
3
16
8
8
0
17
10
32
16
41
31
13
3
0
0
5

31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60

3
0
26
28
0
26
15
14
0
0
6
9
1
4
19
11
21
0
1
0
0
0
2
0
8
0
0
0
0
0

61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90

0
0
28
5
25
48
17
26
6
2
8
9
3
8
0
0
5
0
0
6
4
6
0
10
0
27
0
1
28
19

91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120

5
3
0
15
6
21
4
10
0
0
0
11
1
18
18
6
0
1
2
0
16
22
34
0
0
0
6
7
0
0

2

n

=

120

s

ix

=

989

s

=

10.25

s

=

0.94

-

x

i.x2
(ix)2

=
=

8.2

= 105.09

x

20,657

= 978,121

C.

v. =

90% Confidence

.

Interval

8.2 ± (1.658)(.94) = 8.2 ± 1.6

Total

POEulation

Estimate

125%
21,812 ± 4,256

�-369-

Population Structure
Aerial post-season classifications in the southern part of the Piceance
winter range were made by Mr. Claude White, Northwest Regional Biologist,
on December 11, 1973. Classifications over the rest of the winter range
were extended over about a one month period due to weather delays. Resultant counting dates were December 12, 13, 28 and 29; 1973 and January
10, 1974. Ground counts were attempted but not enough deer could be found
to warrant continuation.
The 1,597 deer classified included 244 from Northwest Region counts (Table
2). The buck:doe:fawn ratio was 22:100:63 which is 6 bucks and 11 fawns:
100 does less than in 1972.

Table 2. Sununaryof post-season deer classifications on the Piceance winter
range in Game Management Unit 22 during winter, 1973-74.
Number of Deer
Bucks
Does
Fawns

Buck: Doe
Ratio

Fawn:Doe
Ratio

Research

166

721

466

23:100

65:100

Northwest Region

25

139

80

18:100

58:100

Combined Total

191

860

546

22:100

63:100

Count

Check station data for Game Management Unit 22 show that of 70 fawns, 47
percent were males and 53 percent females. Of 194 females older than fawns,
10 percent were classed as yearlings by tooth replacement.
A summary of the 1973-74 deer population density and structure data follow:
Total Population Estimate

21,812 ± 4,256

Total Adult Males

2,596

Total Adult Females

11,778

Yearling Females
Mature Females

1,178
10,600

Total Fawns
Male Fawns
Female Fawns

7,438
3,496
3,942

�-370-

Revision of 1972-73 Population Estimate
It was previously expressed that a greater than "normal" number of animals
were probably missed during last year's census because of the experimental
nature of the count (Bartmann 1973). This resulted in an underestimate of
the total population. An effort to estimate the magnitude of that error
is made here by working the population equation backwards. The projected
1973 harvest estimate for Game Management Unit 22 is based on check station
data a-sthe Game Management Section's random survey data are not yet available.
Total bucks and does during winter, 1973-74

= 14,374

Total 1973 buck and doe harvest including wounding loss

+ 3,634

1973 winter and highway mortality (bucks, does and fawns)

+ 11,890

Total revised 1972-73 winter population estimate

= 29,898

The original 1972-73 winter population estimate was 19.684 deer. The revised estimate is 10,214 animals greater, indicating about one-third of the
deer were missed during the count. The accuracy of the revised estimate is
still unknown, but is thought to be closer to the actual population figure
than the original estimate.

LITERATURE CITED
Bartmann, R. M. 1973. Piceance deer study - population density and
structure. In Game Research Report. Colo. Div. Wildl., Denver.
July. (In press).

Prepared by

~
-- ,'.
/ /;/ .';~/~?
~//~
_7'-'-(.
,"-..&lt;,.~-.--.. _
L

i

(

.""

/

/

C

Richard M:" Bartmann
Wildlife Researcher

�-371-

Ju1y~ 1974

JOR PROGRESS REPORT

State of _~~_CO~LO=RADO=::.::..._
Project No.
WorkPlan No.

Job Title

~__
38-R_-_2_8

:

Deer-Elk Investigations

16
: Job No.
3
-----------------~----------Piceance Deer Study - Productivity

and Mortality

Period Covered: April L, 1973 through l'f..arch3l~ 1974•
. Personnel: R. M. Hartmann, S. F. Ste~ert,
J. J. Kl.edn
, Jr.,
Stone, D. Talcott, J. F. Corey, and C. E. White.

L. Crooks, R.

ABSTRACT

Combinedpre-season ·c1assification data from the upper White River and Roan
Plateau showed72 fawns:lOOdoes. Elimination of essentially non-productive
.yearl:ingdoesfrom the doe population produced a fawn:mother ratio of 80:100.
An estimate of the total deer kill from the GameManagementSections's random
survey for the regular season is not yet available.
Six regular and one
temporary check stations tallied 1,264 deer from Unit 22 during the regular
season. The temporary station at Douglas Pass checked onl.y 10 deer from Unit
22, so this station will not be operated in the future.
The estimated archery
harvest in Unit 22 was 43 deer. Searches of 40 l/8-souare mile (1/4-mile by
l/2-mile) quadrats yielded 88 dead deer for·a mean 2.2 deer per plot, 17.6
dead deer per square mile, or 11,704 total dead deer on the winter range.
This is nearly 40 percent of the total revised 1972-73 winter population of
29,898 deer.
Compositionof the projected dead deer population was 3~724 .
male fawns, 4,722 female fawns, 465 mature males and 2,793 mature females.

��-373-

PICEANCE DEER STUDY - PRODUCTIVITY AND MORTALITY
Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.
SEGMENT OBJECTIVES
1.

To estimate increments and losses to the Piceance deer population to
allow evaluation of density estimate techniques.
a.
b.
c.

Estimate productivity rates.
Estimate hunter harvest rates.
Estimate winter mortality rates.
METHODS AND MATERIALS

Pre-Natal Productivity
See Bartmann (1972).
Pre-Season Classifications
Fawn:doe classifications were made from a helicopter on summer range areas
within the White River drainage during late September, 1973. Results of
similar classifications made on the Roan Plateau by Northwest Region biologists
were included with those from the upper White River.
Hunter Harvest
An estimate of the total deer harvest during both the archery and rifle
seasons is obtained from random surveys by the Game Management Section.
Information on sex and age composition of the rifle harvest is obtained from
permanent Division check stations located throughout the State and, in 1973,
from a temporary station at Douglas Pass.

Winter Mortality
An estimate of winter mortality was made from searches of 40 lIB-square mile
quadrats (l/4-mile by l/2-mile). Mortality quadrats were randomly selected
from the original 60 1/4-square mile census quadrats (Fig. 1). Each selected

�-374-

census quadrat was divided into north and south or east and west halves,
depending on which division provided the greatest elevational change.
One of the halves was then randomly selected for mortality searches.
Forty quadrats were used as a preliminary sample to identify problems and
to determine how many quadrats could be searched within the six-week period
allotted to this work. Each quadrat was sub-divided lengthwise into four
equal strips about 330-feet wide by ll2-mile long. Strips were searched
by a five to six-man crew spaced across the width of each strip. The two
outside crew members carried a photo of each plot, compass, and surveyor's
flagging to help keep on course and to insure complete coverage of each
quadrat.
Each deer judged as having died during the current season was classed as
to sex and age when possible. An incisor was collected from deer estimated
as older than a yearling for later aging by dental cementum. A deer was
counted only if the pelvic girdle was present (Gill 1969). The location of
each carcass was marked on a photo to aid in separating current from older
mortalities in later years. Road-killed animals were not included as these
were tallied separately. All deer found incidental to the mortality searches
were also recorded.

Highway Mortalities
Semi-regular checks were made of major roads in and around Game Management
Unit 22 for road-killed deer during March and April. At other times, roadkills were recorded whenever they were encountered during the course of
other activities. No estimate of the total highway kill is attempted.
Instead, the tally is considered a minimum count.

RESULTS AND DISCUSSION

Pre-Natal Productivity
Pre-natal productivity data are largely collected from does killed along
highways during Marchand April. Because of new reporting deadlines, the
spring 1974 data will necessarily be included in the Segment 29 report.
Pre-Season Classifications
The 1973 pre-season classifications in the upper White River area were made
September 26-29. Classifications were made on the Roan Plateau by Mr. Claude
White, Northwest Region biologist, September 24 and 25. The 1,785 total deer
classified, which included 545 deer from the Roan Plateau, consisted of 634
fawns, 883 does and 282 bucks (Table 1). The resultant fawn:doe ratio was
72:100. Previous data are only available for the Roan Plateau and only for
the procedingtwo years. Ratios for 1971 and 1972 were 78 and 79 fawns:100
does, respectively. The 1973 ratio for the same area was 70 fawns:100 does.

�GAME

MANAGEMENT UNIT

22

-PICEANCE-

~~

•....

....•..,

)•••,

.,...,.....- ..;", ..•.

I

.\--'----""'~-"--.,..'••••

~.

l(~"'-.~

W
•....•

+

••
"

I~~~)h

r ///'0
I

t

~j

I~

\'-..

I

.0

./ . 1ft.

\~c.

Fig. 1. Looations of l/8-square mile deer winter mortality quadrats on the Pioeanoe winter range.

VI
I

�-376-

Table 1. Results of pre-season deer classifications in the White River
and Roan Plateau areas during fall, 1973.

Bucks

Does

Fawns

Total

Buck:Doe
Ratio

Fawn:Doe
Ratio

Fawn:Mother
Ratio!!

Strawberry CreekSulphur Creek

15

84

38

137

18:100

45:100

50:100

Nine Mile-Coal Creek

56

139

97

292

40:100

70:100

78:100

Oak Ridge

33

105

47

171

31:100

45:100

49:100

Miller CreekFlag Creek

70

252

228

550

28:100

90:100

100:100

Piceance Triangle

9

41

40

90

22:100

98:100

108:100

Naval Oil Shale !!

79

208

158

445

38:100

76:100

84:100

Square S-Cathedral !! 20

54

26

100

37:100

48:100

54:100

282

883

634

1,785

32:100

72:100

80:100

Area

Totals

JJ

Fawn:Mother Ratio = Fawn:Doe ratio recalculated after removing the estimated
number of non-producing yearlings from the doe category.

!!
Data are from counts made by Northwest Region personnel.

A second ratio estimate was calculated based on producing does only. Check
station data provided the estimate of the proportion of essentially nonproducing yearling does (10 percent) which were then removed from the total
doe population. The resulting fawn:mother ratio (Anderson 1965) was 80:100.
Similar treatment of the 1972 Roan Plateau data, when 28 percent yearlings
were estimated in the harvest, showed 110 fawns:lOO mothers. This suggests
an even lower productivity in 1973 relative to the previous year than was
indicated by the fawns:lOO total does r~tio. Estimated pre-natal productivity,
based on data from road-killed does, was identical in 1972 and 1973 (145
fetuses:lOO does). This suggests that most of the mortality implied by the
above data probably occurred during the interval from near parturition to
late September when pre-season classifications were made.

�-377-

Hunter Harvest
A statewide II-day deer season was set for October 13-23, 1973. In Game
Management Units 21, 22, 23 and 24 in the White River drainage, the first
six days were open to one deer, antlered only, followed by five days of
one deer, either sex. In Unit 11, part of which is in the White River
drainage, the entire season was open to one deer, either sex.
The total estimated deer harvest from the Game Management Section's random survey will not be available until late spring, 1974. A preliminary
estimate based on projection of check station data for Unit 22 is 3,094
deer. The random survey for the archery season is complete, however, and
shows only 43 deer killed in Unit 22 (Table 2).

Table 2. The 1973 archery deer harvest report for four upper White River
Game Management Units.

Unit

Bucks

Does

Fawns

11

6

8

3

22

22

18

3

23

22

3

3

24

3

o

6

Twelve hundred sixty-four deer from Unit 22 were tallied at check stations
throughout the State (Table 3). The temporary station at Douglas Pass
(instead of Dinosaur as originally planned) checked only 10 deer from Unit
22 (Table 4). Therefore, this station will not be operated in 1974.

Table 3. Sex and age composition of the 1973 deer harvest in four upper
White River Game Management Units inspected at six regular and one research
check station.
Unit

Fawns

Bucks
Yrlgs.

2+

Unk.

Fawns

Yrlgs.

Does
2+

Unk.

11

140

68

403

276

103

49

510

332

22

33

79

598

222

37

20

174

101

23

14

28

212

124

9

4

37

49

24

1

16

73

37

9

3

15

19

�-378-

Table 4. Summary of deer checked at the Douglas Pass check station
during the first nine days of the 1973 deer season.
Unit

Bucks

Does

Fawns

Unknown

Total

.10

20

0

0

5

25

11

7

2

0

4

13

21

162

20

7

14

203

22

8

0

0

2

10

23

0

1

0

0

1

30

52

36

11

9

108

31

4

1

0

0

5

44

1

0

0

0

1

61

2

0

0

0

2

Total

256

60

18

34

368

Winter Mortality
Dead deer searches were made from May 2 through June 6, 1973. Eighty-eight
dead deer were found that were judged losses of the current year (Table 5).
The estimated mean of 2.2 deer per plot projects to 17.6 dead deer per
square mile or 11,704 total dead deer on the winter range in Game Management
Unit 22.
The sex and age breakdown of the total estimated winter loss, with unknowns
apportioned among the various classes, was 3,724 male fawns, 4,722 female
fawns, 465 mature males and 2,793 mature femal~s. An additional 49 dead
deer were found incidental to the mortality survey which included 13 male
fawns, 14 female fawns, 2 unknown fawns, 1 mature male, 18 mature females,
and 1 unknown adult.
The mortality estimate includes an unknown proportion of crippling loss and
illegal kill from the previous hunting season but no attempt was made to
allow for these animals as the magnitude and composition of the loss is unknown. Too, further complication would arise as only part of the loss occurs
on the winter range and this proportion can change yearly depending on the
timing of the hunting season together with weather and other conditions.
Another consideration is that some of the winter related mortality occurring
later in the spring is outside the bounds of the winter range area sampled.

�-379-

Table 5. Results of dead deer searches on 40 1/4-mi1e by 1/2-mi1e
quadrats on the Piceance winter range during spring, 1973.
Quadrat
Number

2
5
6
7
8
9
11
12
13
14
15
16
17
21
22
23
24
26
27
28
29
33
34
35
37
38
39
44
45
46
47
50
52
53
54
55
56
58
59
60
Totals

Male

Fawns
Female

Unknown

Male

Mature
Female

Unknown

Unknown

None
3

1

2

None
1
1
1
1
2

1
None
None
2
1

1
1

1

2
1

None
2

1

1
2

1
1

1
1

3

2
None
1

1

1

1

None
1
None
1
2

1

None
1
1
5
3
1

1
1

1
1

2
2
1
2

2
2
1
1
None

3
2

5
1

1
2
None

23

29

10

3

19

2

2

�-380-

A revised winter population estimate was calculated for 1972-73 by
working the population equation backwards from the 1973-74 winter estimate (see report on Population Density and Structure elsewhere in this
Game Research Report). If the revised estimate of 29,898 deer is considered more appropriate, nearly 40 percent of the total population died
during or immediately following that winter.

LITERATURE CITED
Anderson, A. E. 1965. Population density and structure. P. 47-74. In
Game Research Report. Colo. Game, Fish and Parks Dept., Denver.
l(Part 1):1-246.
Bartmann, R. M. 1972. Piceance deer study - productivity and mortality.
P. 345-350. In Game Research Report. Colo. Div. Wi1d1., Denver.
3(Part 3):253-377.
Gill, R. B. 1969. Middle Park deer study - population productivity and
mortality. P. 123-140. In Game Research Report. Colo. Game, Fish
and Parks Div., Denver. 3{Part 1):1-140.

Prepared

�-381-

July, 1974

JOB PROGRESS REPORT

State of

......::CO:;=.=LO.::,:RAD:;:.:::.;:O:..._

Proj ect No.

W_-_3.,;.8-_R_-_2_8 _

Job No.

17

Work Plan No.

Deer-Elk Investigations

Job Title

Systems Modeling Big Game Populations

Period Covered:

April 1, 1973 through March 31, 1974

Personnel:

1__ ~

_

James F. Lipscomb

ABSTRACT
Efforts during this segment were concentrated on generalizing and standardizing
modeling procedures to save time and insure optimum results as .addLtdonaf big
game herds are modeled. This was judged to be essential for the successful
completion of Work Plan 18 of this project.
The entire data analysis portion of the modeling procedure was structured
as a nonlinear programming problem. Efforts are currently being directed
toward testing various solution algorithms to determine which are the most
dependable and most efficient. A detailed description and explanation of
the analysis method chosen will be presented at the meeting of the Western
Association of State Game and Fish Commissioners in July 1974, and will become
a part of the published transactions.
This job is currently about six months behind schedule due to my past and
present assignments to the Colorado Division of Wildlife's planning project.

Researcher

��-383-

July, 1974

JOB PROGRESS REPORT

COLORA:..:;:D::..;O=----

Proj ect No.

W-38-R-28

Work Pinn l~o.

18

_

Deer-Elk Investigations
Job No.

---

Job Title

Deer and Elk Management Study

Period Covered:

April 1, 1973 to March 31, 1974

Personnel:

1

~----------~---------------

Bertram D. Baker, Raymond J. Boyd, James -P, Lipscomb and Thomas
M. Pojar.

The State was divided into data analysis units, with separate units for deer
and elk, according to predetermined criteria. Organization and evaluation
of data with potential for management purposes was begun. A hunter destinationroute survey was initiated and conducted at selected deer and elk check stations
during 1973 seasons. Data gathered indicated that closure of two deer check
stations and opening a new station near Penrose would check over 90 percent
of the deer normally checked and would eliminate double checking of hunters.
Closure of one elk station and moving another to the east side of LaVeta Pass
would greatly Increase efficiency and eliminate double checking.

. .....

'",.

-:

��-385-

DEER AND ELK MANAGEMENT STUDY
Bertram D. Baker
Raymond J. Boyd
James F. Lipscomb
Thomas M. Pojar

There has always been a need to quantify management objectives for wild
populations. The difficulty of obtaining reliable information upon which
to base these objectives has long been realized by wildlife managers. However, methods for estimating various population parameters are available
and methods of analysis are currently being tested (Work Plan llC, Job 3;
and Work Plan 17, Job 1) using the computer population simulation approach.
If feasible, this approach will be used to incorporate any presently available or potentially available information into a data analysis system for
each data analysis unit.
P. S. OBJECTIVE
Devise and test a statewide deer and elk management system.

SEGMENT OBJECTIVES
1.

Divide the state into a system of data analysis units (DAD's) that are
deemed practical for gathering and analyzing population data.

2.

Based upon currently available information, estimate population parameters.

3.

Identify data that are most useful for improving estimates of population
parameters and recommend procedures for collecting such data by DAD.
METHODS AND MATERIALS

The four regional biologists were asked to delineate on a state map the data
analysis units for deer and elk in their respective regions. The criteria to
identify a data analysis unit (DAU) are:
a.

The area enclosed will encompass the year-round range of the population.

b.

The data collected are characteristic of the population inhabiting
the DAD (i.e., the animals counted in the DAD are the same animals
that are hunted in the DAD).

c.

The DAU boundaries should follow existing game management unit
boundaries wherever possible but can encompass more than one game
management unit.

�-386-

Tagging recoveries of both deer and elk were used to help verify DAU
boundaries.
A list of potentially available population data was made (APPENDIX A)
with possible sources of this data being: Planning Section files, Regional
files, Game Management files in Denver, Game Research project files, U. S.
Forest Service and Bureau of Land Management files. A filing system was
set up so that existing data can be accumulated for each DAU.
Att"empts will be made to identify those data that are most useful for improving population parameter estimates using sensitivity analysis from the
population model developed under Work Plan 17, Job 1.

RESULTS AND DISCUSSION
The four regional biologists drew the boundaries of DAU's for both deer and
elk on 1:500,000 scale maps for their respective regions according to the
criteria mentioned above. Any boundary discrepancies of DAU's that involved
two or more regions was resolved by the regional biologists involved. This
is to be considered a preliminary effort to divide the State into realistic
data analysis units. Consequently, the boundaries will be changed in the
future if evidence is discovered that indicates the initial DAU does not meet
specified criteria.
Deer and elk banding records were updated and summarized as in APPENDIX B
on a statewide basis. Banding recovery data, although meager in most cases,
was used to indicate validity of DAUboundaries.
The accumulation and tabulation of available population data is nearly complete. A filing system where data are accumulated by DAU has been established.
Mean 10-year harvest by DAU was calculated and the DAU's were ranked accordingly
(Tables 1 and 2; Figures 1 and 2). The data accumulated by DAU will be used
in a model developed under WP 17, J 1 to estimate population parameters.
Segment objectives 2 and 3 have not been attained in this segment because of
a delay in progress of WP 17, J 1 due to reassignment of the principal investigator.
Sex and age of harvested animals is presently obtained from check stations.
A review of the effectiveness and efficiency of the current check station
system resulted in an effort to locate check stations to accomplish the following:
(1) eliminate potential double check of hunters, (2) obtain reasonable samples
from areas that historically yield significant proportions of the statewide
harvest, and (3) reduce oversampling where it occurs. To accomplish these
objectives some of the present check stations would need to be abandoned or
relocated. The results of a hunter destination-route survey conducted during
1973 seasons at stations considered for relocation or abandonment are presented
in Tables 3 and 4. Based on the survey and projections of how many animals
may be checked at various sites~ it was decided that 1974 check stations would
be located at Idaho Springs, Canon City, and Towaoc during deer season and
at Idaho spring~narCh,
and LaVeta :ass during elk season.
Prepared by

It-&amp;'~-n£j}/k
Raymond J. Bo'§~'
Asst. Wildlife Researcher

�-387-

Table 1.

Deer DAU's - ranked

in order of mean 10-year kill.

DAU

Rank

Mean 10-year
Kill

D-7

1

19,337

White River (S 1/3 of 11 (60% of Ki11)l/, 12, 13,
22, 23, 24, 25, 26, 31, 32, 33, 34)

D-12

2

6,119

Grand Mesa (41, 411, 42, 52, 521)

D-6

3

4,558

Rangely (10, N 2/3 of 11 (40% of Ki11)l/, 21)

D-19

4

4,133

Uncompahgre

D-30

5

3,927

San Juan (75, 751

D-2

6

3,483

Bear's Ears (4, 441, 5, 14)

D-24

7

3,462

Groundhog (70,71)

D-9

8

3,236

Middle Park (18, 28, 27, 37)

D-8

9

3,184

State Bridge (15, 35, 36)

D-16

10

2,943

Cripple Creek (49, 57, 58, 581, 59)

D-25

11

2,666

Powderhorn Creek (65, 66, 67)

D-20

12

2,593

Crawford (53, 63, 64)

D-14

13

2,556

Red Tables (44, 45)

D-13

14

2,283

Maroon Bells (43, 47)

D-29

15

2,201

Mesa Verde (72, 73, 74)

D-18

16

2,190

Glade Park (40)

D-27

17

2,154

Westcliffe (69, 82, 84, 86)

D-33

18

1,793

Great Divide (3)

D-4

19

1,693

Red Feather (7, 8, 9, 19)

D-ll

20

1,225

Book Cliffs (30)

D-1

21

1,202

Little Snake (1, 2)

D-32

22

1,169

Trinidad (83, 85, 112)

D-17

23

1,127

Bailey (39, 391, 46, 461, 50, 501, 51, 511)

DAU Name and Units Involved

(61, 62)
77, 78)

--------------------------------------------------------

�-388-

Table 1.

Deer DAU's - ranked in order of mean 10-year kill (continued).

DAU

Rank

Mean 10-year
Kill

D-3

24

1,084

North Park (6, 16, 161, 17)

D-31

·25

1,066

Rio Grande (76, 79, 80, 81)

D-21

26

1,023

West Elk (54)

D-lO

27

1,013

Bould er (20, 29, 38, 381)

D-22

28

893

Taylor Park (55, 551)

D-15

29

667

Cottonwood Creek (48, 56)

D-26

30

564

Saguache (68)

D-5

31

494

South Platte (87, 88, 89, 90, 91, 92, 93, 94,
95, 96, 99, 100, 101, 104, 105)

D-28

32

491

Arkansas Valley (97, 98, 102, 103, 106, 107,
108, 109, 110, Ill, 113, 114)

D-23

33

422

LaSal (60)

DAU Name and Units Involved

~/Estimate of NW Reg. Biologists, White and Blumberg.

�-389-

Table 2. .E1k DAU' s - ranked in order of mean 10-year kill.

DAU

Rank

Mean 10-year
Kill

E-6

1

2,934

White River (units 12, 13, 23, 24, 25, 26, 33, 34)

E-31

2

1,958

San Juan (units 75, 751, 77, 78)

E-32

3

1,422

Rio Grande (units 76, 79, 80, 81)

E-21

4

1,047

Gunnison River (units 63, 64, 53, 54, 55, 551)

E-2

5

841

Bear's Ears (units 4, 5, 441, 14)

E-25

6

825

Lake Fork (units 65, 66, 67)

E-14

7

741

Grand Mesa (units 41, 411, 42, 52, 521)

E-30

8

597

Hermosa (unit 74)

E-24

9

528

Disappointment Creek (units 70, 71)

E-16

10

510

Frying Pan (units 44, 45, ~47)

E-15

11

347

Avalanche Creek (units 43, %47)

E-3

12

342

North Park (units 6, 7, 16, 161, 17)

E-7

13

284

Gore Pass (units 15, 27)

E-13

14

273

Williams Fork (units 28, 37)

E-8

15

265

Troublesome (unit 18)

E-26

16

260

Saguache (unit 68)

E-9

17

251

St. Vrain (units 20, 29, 38, 381)

E-18

18

243

Kenosha Pass (units 50, 501, 51, 511, 46, 461,
39, 391)

E-29

19

186

Mesa Verde (units 72, 73)

E-4

20

164

Poudre River (units 8, 19)

E-33

21

148

Trinchera (units 83, 85)

E-17

22

147

Collegiate Range (units 48, 56)

E-5

23

126

Strawberry (unit 11)

DAU Name and Units Involved

-----------------------------------------------------------

�-390-

Table 2.

Elk DAU's

- ranked in order of mean 10-year kill (continued) .

Mean 10-year
Kill

DAU

Rank

E-20

24

121

Uncompahgre

E-l

25

102

Cold Springs (unit 2)

E-22

26

89

Buffalo Peaks (units 49, 57, 58)

E-12

27

70

Piney (units 35, 36)

E-28

28

42

Grape Creek (units 69, 84)

E-ll

29

41

Roan Bluffs (units 31, 32)

E-27

30

37

Sangre de Cristo (units 82, 86)

E-19

31

35

Glade Park (unit 40)

E-23

32

26

II-Mile (units 581, 59)

E-lO

33

19

Yellow Creek (unit 22)

DAU Name and Units Involved
(units 61, 62)

�I

W
\0

,....
I

,_ .. 1
•
;"
•••
v." "" ••••, •.••••" ••••
to

Fig. 1. Deer Data Analysis Units (DAU's) with the top 10 DAU's in mean harvest (1963-1972) shown as
cross-hatched areas. These 10 DAU's account for 62.54 percent of the statewide deer harvest.

�-397APPENDIX B
BIG GAME TAGGING AND BANDING SUMMARY
(Northwest Region)

Trap Site Name - Big Game Management Unit 22 - Piceance
Location{s):

See accompanying maps
Name

Ear Tag Numbers

Neckband Color and Numbers

1

Boise-Barcus

L1697 - L1698*
L1700
Ll714 - Ll717
Ll719
Ll722 - Ll723
Ll725 - L1747*
L1749 - L1752
L1754 - L1757
L1760
L1764 - L1765

Blue, 58 - 59, 73, 76, 78,
85, 92 - 93, 95 - 96,
101, 103, 105, 107, 109,
112 - 115, 118 - 139

2

Barcus-Ryan

Ll718
Ll720 - Ll721
Ll724
L1753
L1758 - L1759
L1761 - L1762
L1766 - Ll775
L1807 - L1808
L1814
L1818
L1823 - L1825
L1833 - L1836

White, 207, 210 - 232,
235 - 236, 238 - 240

3

Ryan-Stuart

L1763
Ll776 - L1806
L1809 - L1813
L1815 - L1817
L1819 - L1822
L1826 - L1832
L1837 ...;
L1883**
L1888 - L1891**
L1898 - L1899**
L1940 - L1942**
L1946**
L1953**
L1963**
L1968**

Pink, 3 - 4, 6 -11,
13 - 97, 99 - 100
(Also, 2 No. 62's)

Area Number

*L1699 and L1748 are missing.
**See Area 7 for L1884-L1887, L1892-L1897, L1900-L1939, L1943-L1945, L1947-L1952,
L1954-L1962, L1964-L1967, and L1969-Forward.

�-398BIG GAME TAGGING AND BANDING SL~Y

Location(s):

- Northwest Region, Piceance (continued)

See accompanying maps

Area Number

Name

Ear Tag Numbers

Neckband Color and Numbers

5

Dry ForkGreasewood

L2182
L2184
L2186
L2188
L2301 - L2331

Yellow with Red Stripe,
1 - 8, 10 - 26, 28 - 29,
32, 34, 36, 38, 39

6

GreasewoodDudley Bluffs

L2035 - L2037
L2048
L2068 - L2076*
L2081 - L2099*
L2101 - L2143*
L2145 - L2181
L2183
L2185
L2187
L2189 - L2200

Blue and White, 1 - 2,
4 - 8, 10 - 26, 29 - 60,
62 - 80, 82 - 88, 90,
92 - 97, 100 - 107
(Also, 2 No. 23's and
24's)

7

Dudley B1uffsFourteenmi1e

L1884 - L1887
L1892 - L1897
L1900 - L1939
L1943 - L1945
L1947 - L1952
L1954 - L1962
L1964 - L1967
L1969 - L2034
L2038 - L2047
L2049 - L2067
L2078 - L2080

Red with White Stripe,
1 - 60, 63 - 130
(Also, 2 No. 3's)

Period of Operation:
Total Deer Marked:

December 13, 1972 - March 9, 1973
517

*L2077, L2100 and L2144 are missing.

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                  <text>-399July, 1974

FINAL REPORT

State of
Project No.

COLORADO
W-38-R-28

Deer-Elk Investigations

Work Plan No ._------:.1-=..4
Job Title

_

Job No.

6

--------------

t4iddle Park Cooperative Deer Study - Physiology and Prevention
of Deer Starvation

Period Covered:
Personnel:

June 1969 through June 1974

Julius G. Nagy, Principal Investigator
Dan L. Baker, James A. Bailey, David S. de Calesta,
David E. Reeder and Gene G. Schoonveld, Investigators.
ABSTRACT

Various problems associated with mule deer (Odocoileus hemionus) starvation and refeeding were investigated. It was found that deer and deer
rumen microorganisms can digest good quality alfalfa hay without adverse
effects. Poor quality alfalfa hay, even in ground and pelleted form can
cause severe and occasionally fatal digestive disturbances. Domestic sheep
and goats fed the same alfalfa rations as deer did not develop digestive
problems. Relative to body size deer have a smaller rumen, omasum and abomasum than sheep and goats.
Deer can be starved, depending on their previous physical condition, for
4-8 weeks without significantly affecting rumen microbial digestion. Starved
deer can be refed successfully. Refeeding did not result in toxicity, liver
malfunction or large scale mortality. After ten days of refeeding, previously starved deer showed similar blood energy metabolites as non-starved
animals. Limited field studies with a pelleted concentrate ration indicate
that deer, under the conditions tested, will use this concentrate as a supplement to their natural diet.

�-400-

RECOMr~ENDATI ONS

1. Periodically it might be necessary to feed starving deer a supplemental

emergency ration. Before attempting to execute such a program each case
must be examined carefully from biological viewpoints. Detailed plans
for periodic emergency feeding should be established well ahead of the
stress period. Haphazard feeding of starved animals often results in high
ex~enses, and dead animals which result in a shattered faith on the
part of the public in the knowledge of game managers.
2. Good quality alfalfa hay should be used only in conjunction with some
high energy concentrate as an emergency supplemental feed. Hay when compared to concentrates is slowly digested and consequently does not give
as much energy to the animal per unit time. Hay is also bulky~ difficult
to distribute and much of it is wasted.
3. For emergency feeding programs, a high energy, easily digestible feed
should be used. Depending on the amount of natural vegetation available
at the time of feeding approximately 500-1,000 grams of concentrate should
be calculated per animal per day.
4. Possibly several small feeding stations would be more effective in terms
of feeding deer than a few large ones.

�-401-

GENERAL INTRODUCTION

Research summarized in this Final Report was conducted by Colorado State
University through contractual arrangement with the Colorado Division of
Wildlife.

Basic information was deficient concerning the digestive physiology

of mule deer (Odocoileus hemionus) especially during periods of nutritional
deprivation.

With such knowledge, the role of supplemental winter feeding

programs in deer management programs could be evaluated more objectively.
Over the past few decades controversy has raged over whether wild deer
can or cannot be fed "supplemental" diets to reserve terminal starvation
process.

Included in the controversy were such questions as:

what should

be fed?; when should feeding commence?; what happens to native forages in
the vicinity of supplemental feed sites?; is supplemental feeding "practical"?;
etc. This study originally attempted to address all of these questions.
The approach was first to investigate the physiological aspects of supplemental
feeding, through controlled experiments with penned deer.

Ultimately, the

knowledge gained from the controlled experiments was to be tested in field
trials with wild, free-ranging mule deer.
The original objectives of this study were to:
1. measure basic metabolic rate of mule deer
2. determine the critical ambient temperature where deer must increase
their rate of metabolism to compensate for heat losses due to cold
stress
3. determine changes in mule deer physiology during the process of
starvation
4. determine the "critical" point of starvation beyond which deer can not
recover.

�-4025.

investigate methods of aiding deer recovery during starvation before
the "critical" point is passed

6. evaluate the biological and economical feasibility of preventing starvation
in wild deer populations
Two of these objectives, 1 and 2, were not investigated because of recent
work by Silver et al. (1971).
For the sake of clarity, this Final Report is divided into three parts,
each containing an introduction, methods, and results and discussion section.
Part I concerns itself with Deer Feeding Experiments, Part II with Physiology
of Deer Starvation, and Part III with Supplemental Feeding of Wild Deer.
ACKNOWLEDGMENTS
This study (Work Plan 14 Job 6) was one job of several conducted under the
auspices of the Middle Park Cooperative Deer Ecology Study.

Colorado State

University through the Department of Fishery and Wildlife Biology assumed major
responsibility for design, conduct and analysis of this phase of the overall
Middle Park deer program.

We wish to acknowledge and express appreciation for

assistance received from the Colorado Division of Wildlife and the Rocky Mountain
Forest and Range Experiment Station, USDA, Forest Service.

Specifically, we

thank O. C. Wallmo, D. W. Reichert, W. L. Regelin, R. B. Gill, H. M. Swope,
W. Aldridge, T. Barber, S. Kerr, J. Olson, W. Paintner, and C. Schwartz for
their guidance, criticism, and physical effort toward the success of this
endeavor.
P. S. OBJECTIVE
To ascertain what takes place in deer physiology when starvation occurs,
and investigate the economic and biologic feasibility of preventing starvation.

�-403PART I
DEER FEEDING EXPERIMENTS
Chapter 1
General Description of Study Areas, Facilities and Animals
Feeding, food restriction and refeeding trials were conducted at the
Deer Research Facilities of Colorado State University Foothills Campus (Fig. 1).
Field trials were conducted in Middle Park, Colorado at the Colorado Division
of Wildlife Junction Butte property.

Laboratory work was conducted at the

Department of Animal Science, and the Department of Physiology and Biophysics,
Colorado State University.

At the CSU Foothills Campus fifteen isolation

pens (6.1 x 1.8 x 2.4 m high) connected to weighing and restraining chutes
were constructed.

Pens were adjacent and were separated by double strand V-mesh

wire; ends were constructed of horizontal boards spaced 3-5 cm.

Covered

feeders providing for one feed pail and one water pail were installed in each
pen.

One end was roofed for 2 m and opened into a common alleyway leading to

weighing and restraining chutes (Fig. 2).

The alleyway was constructed of

horizontal, unspaced lumber, 2.4 m high.

Weighing and restraining chutes were

1.1 x 0.33 x 1.2 m high and constructed of plywood.

Deer were forced from the

alleyway into weighing or restraining chutes by a researcher carrying a crowding
shield.

Overhead doors were lowered to hold an animal in the weighing chute

or to force it into the restraining chute.

The weighing chute floor was the

base of a platform scale where deer weights could be read to the nearest 1/4 kg.
Deer were held by the neck in the restraining chute by clamping a wooden collar
after the animal had passed its head through a 20 cm diameter opening cut in the
chute end (Fig. 2).
was 62 cm.

Distance from the center of this opening to the floor

This height was critical in order to pass a stomach tube down the

esophagus to sample rumen contents.

When the hole was higher, the tube

invariably entered the animal's trachea.

�v-,

I

SHED

o Wl:iGHti\G

AND HANDLING

b WORKING

SPACE

CHUTE'

)

J7~::::::==========:::=:~':rttl~5~To~~
.po

a

~
I

,,,GO'~L~~~YI~OP~N/
HI
"

';)

I~

,___

.

1 ILAB IFl

II........!

~

I C.J I

1-'

,

.•...

f

I / HOUSE
,I
I TRA!LER

GROUP
SOLATiON

'--J
o

.oo

C=~~==~==~==x====~----~
:;0

::;CALE

I N FeE;"

Figure 1. Physical layout of deer holding facilities.

-,

iSOLATION

DI~CfG~dcON
$t1=.LI ~R

STALLS

�il

-

----I

-405-

I

.01

I.

';,

.

~

'.

,.._;~.

I ~~\"'::::'
\
~

\
\

\

,

\

~I

!I. '. I \ \ '.\ ~ \ .C:~--=--=--=-=-. \\;'.'

~-=:-----=-

!'/

!I/,/,

.I

�-406-

Animals used in this study were captured from wild populations of deer
at the·Colorado Division of Wildlife Little Hills Experiment Station or in
Middle Park, Colorado and transported to Fort Collins.
hand raised, tame animals also were used.

In later trials,

�-407Chapter 2
Effects of Prolonged Malnutrition and Alfalfa Hay
Refeeding on Mule Deer
Introduction
These experiments were conducted to study the effects of prolonged submaintenance food intake followed by sudden changes from this diet to
alfalfa hay.
Methods and Materials
Facilities
Experiments were conducted at the Deer Research Facility of Colorado
State University.
Experimental Deer
Five adult mule deer does and four fawns (1 male, 3 female) were used.
Adults were trapped in the wild and held in captivity for approximately 1
year pretrial.

One adult doe died during the 6th week of the trial, and

another during the 12th week; both losses were attributed to handling.

Fawns

in the experiment were born in captivity, raised by their does and were
approximately 8-9 months old when the experiment began.

On January 1, 1970

the adult does and male fawn were confined to individual isolation pens and
began a 6-week pretrial acclimation period.

The three female fawns were

confined in isolation pens on 25 February.

These fawns adjusted to confinement

rapidly and were allowed only a 3-week pretrial acclimation period.

All

experimental deer appeared to be eating normally and were judged in excellent
physical condition when trials started.
Trial Diets and Feeding
Prior to confinement all animals had access to concentrate and alfalfa
hay ad. libitum.

Concentrate consisted of 50 percent barley, 20 percent corn,

20 percent milo and 10 percent bran.

During pretrial confinement deer

�-408continued to receive concentrate ration ad libitum.

Daily food intake was

measured during pretrial confinement, during the trial, and upon refeeding
by offering each animal an excess quantity of ration and weighing back
orts daily.

Both feed and water were offered in No. 10 galvanized pails

and changed or added to daily.

Water consumption was not measured.

A compounded, poor-quality ration (Diet III) fed during the trial
consisted of 15 percent wheat straw, 75 percent cotton seed hulls and 5
percent each of wheat bran and a 32-percent protein supplement.

Components

were ground in a hammer mill until they passed through a 3 mm screen and after
adding 1.5 percent molasses and steam, pellets measuring 7 x 12 mm were
produced.

Dehydrated alfalfa was fed in 4 cm square cubes ranging in length

from 2-5 cm.

The compounded ration and alfalfa cubes were analyzed for dry

matter, crude protein, ether extract and ash (Association of Official Agricultural
Chemists 1965), cell wall constituents (Van Soest and Wine 1967) and acid
detergent fiber (ADF) and lignin (Van Soest 1963) (Table 1).
Weight Change
Food and water were removed from each deer 4 hours prior to weighing.
Deer were weighed initially at the start of the trial, periodically during
and at the end of the trial, and again after 2 weeks of refeeding.

Weight

gain or loss was recorded to the 'nearest 1/4 kg.
Rumen contents were sampled by passing a 17 mm outside diameter (9 mm
inside diameter) stomach tube down the esophagus into the rumen.

A plastic

pipe (1 x 12 inches long) served as a speculum and facilitated insertion and
manipulation of the stomach tube.

Negative pressure was supplied by a portable

vacuum pump attached to the stomach tube through a 1 liter or larger vacuum
flask.

With some deer, obtaining 50 ml of rumen contents required considerable

effort.

With others, rumen contents would siphon and excess of 2 1iters was

obtained.

Initial amounts of rumen content samples were discarded to minimize

�-409saliva contamination.

The pH of rumen contents was immediately measured using

a Fisher Accumet Model 210 pH meter calibrated against standard buffer prior
to use.

Rumen fluid was separated from solid material by filtering the

contents through several layers of cheesecloth.

A sample was prepared for

protozoan counts by pipetting 1 ml of fluid into 9 ml of 10 percent formalin.
Another 10 ml of fluid was acidified with 2 ml of 25 percent meta-phosphoric
acid for VFA determination.

Both samples were frozen at -15C for laboratory

analysis.
Ciliate protozoa within each sample were counted using a Spencer AO Brightline
hemacytometer filling the chamber four times (Warner 1962b).

A 1:10 dilution and

100X magnification using dark field illumination was most satisfactory for
counting protozoa.

Mean number of protozoa counted in the four chambers were

used in calculating protozoa concentration per m1 of rumen fluid.
Concentrations of VFA in rumen fluid were measured by gas chromatography.
A Varian Aerograph, Model 1740 chromatograph with hydrogen flame detector
connected to a Model 20 recorder and Model 477 disc integrator with digital
printer was used.

Samples were injected in duplicate.

Nitrogen carrier gas

flow was 60 m1lminute, hydrogen flow 25 m1lminute, attenuation 4, and column
temperqture 175C.

Column used was 5 feet by 1/8 inch packed with 20 percent

Tween 80 plus 2 percent phosphoric acid (85 percent) on W 60-80 mesh.
Concentrations of VFA in rumen fluid were determined by comparing each sample
against standards using the area under the curve method.
Blood samples were aspirated from the jugular vein using a 20 ml syringe
and 18 ga x 1 inch needle.

Blood was allowed to clot, after which the serum

was withdrawn and frozen for laboratory analysis.

Care was taken to avoid

withdrawing any formed elements which would bias laboratory analysis.

Serum

was analyzed for total protein and urea by the Veterinary Hospital, Diagnostic

�Table 1.
cubes.

Chemical composition

(oven dry basis) of the compounded ration and dehydrated alfalfa hay

Chemical Analysis
Diet

Crude
Protein

Ether
Extract

(Percent)

Moisture

Ash

Cell Wall
Constituents

I
..j:::.

ADFb

Lignin

Compounded rgtion
(Diet III)

5.5

4.9

8.5

5.2

74.3

52.8

15.1

Dehydrated alfalfa cubes

15.9

4.0

7.2

7.6

53.0

38.6

6.8

aComposition -- 75 percent cotton hull seeds, 15 percent wheat straw and 5 percent each of wheat
bran and 32 percent protein supplement.
bAcid-detergent

fiber.

-'
0
I

�-411Laboratory, CSU.

Ureau concentrations were measured by a Urograph (Warner-

Chilcott Laboratories, General Diagnostic Div.).

This procedure is chemically

identical to the Conway microdiffusion method (Conway 1958) but uses
chromatography paper bonded with controlled amounts of reagents to measure
urea nitrogen contained in the sample.

Serum protein was measured using a

refractometer (American Optical Co.).

This procedure is a colorimetric

test that compares color of the sample after adding reagents to color of
standards with known protein content.
Data Analysis
Many of the parameters measured were not evaluated statistically because
of the small numbers of observations and animals in the experiment.

Where

appropriate, a least-squares analysis of variance was used to evaluate sample
differences.

Many physiological changes monitored during the experiment and

upon refeeding were sudden and of such magnitude they were subjectively
considered significant.
Results and Discussion
General Physical Condition
Experimental deer were shaggy and had not lost their winter coats by the
end of the 90-day experiment.
in full summer coats.

In contrast, deer not on trial were sleek and

Antler growth of the one buck used was limited to l-inch

buttons at trial's end and little additional growth occurred therafter during
the rffoaining growth period.

By comparison a similar-aged buck, not used in

the experiment, had branched antlers when the trial ended.

Feeding alfalfa

had a more diverse effect upon experimental deer, especially fawns, than did
the poor-quality. compounded diet.

Within two weeks after feeding alfalfa

began, one fawn died and apparent physical condition of two others declined
markedly,

�-412-

Cone.. ~

_

Diet III

2500

2000

~ 1500
III

~
eo

o

"'-0(4)

500

x-x Adults
0-0 Fawns

o

pt

1

2

3

4

5

6

7

8

9

10

11 12

I 3 14

15

Weeks
Fig.

3.

Mean daily food intake of concentrate
(pt = pretrial),
Diet III
and dehydrated alfalfa cubes by adult and fawn mule deer.

Figures in parentheses indicate numbers of animals remaining
on trial.

�-413-

1--.-------

Diet

-&gt;

------~Alf.

III

I
I

3

I

I

2
X~

:

I

~~

:g
~
l-&lt;

'0

i

Q)

P.

I
I

x~

x

~ (-) 1

~
III

~ J

..s::::

o (-)2
j:;

b.o)
....•
Q)

~

o

o

1

2

3

4

0

5

6

7

8

9

}(--X

Addtsl

\

0--0

F'awn s]

12.6

10

11

~4)

12 13 14 15

Week~

Fig. 4 Mean cumulative we iqht changes(percent of initial body weight)
of adult and fawn mule deer. Figures in parentheses indicate
numbers of animals remaining on trial.

�-414Rumino-reticular

Contents

Protozoa concentrations within rumen fluid of deer decreased from pretrial
levels during the experiment and protozoa were either completely eliminated
or greatly reduced when fed dehydrated alfalfa cubes (Fig. 5).
concentrations

Protozoa

in rumen fluid obtained from mule deer during this experiment

were lower than numbers reported by Pearson (1969) in fluid of mule deer fed
compounded diets (570-590 x 103/ml) but higher than protozoa numbers he found
in wild mule deer collected during winter (160 x 103/ml).

Factors responsible

for eliminating or greatly reducing protozoa in rumen contents of deer fed
dehydrated alfalfa cubes are unknown.

One explanation may be that protozoa

in deer rumens are intolerant of high pH.

Protozoa were absent in rumen fluid

having pH from 7.1-7.3 and detectable concentrations were near zero in rumen
fluid having a pH of 6.9-7.0.
Rumen-fluid pH readings from deer on Diet III and on alfalfa during Experiment
(Table 2) were higher than values reported by other researchers with the
exception of Ullrey et~.

(1964).

High pH levels obtained did not appear

to be related to saliva contamination because pH readings in rumen fluid
samples in excess of 1 liter were as hi'ghor higher as readings in samples of
only 50-100 ml.
Significantly lower (p &lt; 0.05) rumen fluid pH levels were observed as
feeding Diet III progressed over the 13 weeks; a trend especially apparent
in fawns (Table 2). After feeding alfalfa cubes. rumen fluid pH increased
(Table 2) becoming alkaline in five of seven deer.

These data suggest rumen

fluid pH levels in deer decline as intake and rate of food passage increase
or possibly while a change occurs in the microflora and fauna due to diet
change.

�-415-

Cone. 4--------2300
600

Diet III

~

AI£.

I
I

I

I
I

I

I

500

X

X

~x~

I

(3j-x

x-:--tI

;:;

S
J.&lt;

I
I

400

C1.l

p..

o

I

0
•....•

~

!

('t)

~

.

0

s::
s::

300

o~%

0
...•
~
'"
J.&lt;

~C1.l
u

s::

200

0

U

100

x-x Adults
0--0

Fawns
( 3)

L-~-L~

o

pt

1

__~~~~
2

3

4

__L-~~~~~~~~~(4)
5

6

7

8

9

10 11 12

3 14

Fig. 5. Protozoa concentrations in rumino-reticular fluid of adult
and fawn mule deer fed concentrate (pt=pretrial) Diet III
and dehydrated alfalfa cubes. Figures in parentheses indicate numbers of animals remaining on trial.

15

�Table 2. Average rumino-reticular
mule deer.

fluid pH, VFA concentration (mM/l) and molar proportions of acetic:propionic

f Numbers in parentheses

mear'l range)

Adults

Week of

acids (C :C ) in adult and fawn
2 3

Fawns

Trial
N
pt

pH

VFA Conc.

VFA C2:C

3

N

pH

VFA Cone,

VFA C2:C3

4

6.4 (6.3-6.5)

39 (31-60)

3.4(2.7-3.8)

a

- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -.- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2

5

6.7(6.4-6.9)

66 (44-101)

4.5 (3.7-5.1)

4

6.8 (6.5-6.9)

62 (45-74)

5.5 (5.4-5.7)

4

5

6.7(6.5-6.9)

74 (55-87)

5.1(4.0-6.2)

4

6.6 (6.6-6.7)

70 (50-79)

5.2 (5.1-5.4)

4

6.6 (6.3-6.9)

73 (52-84)

5.4 (5.0-5.8)

I

6
8

4

6.6 (6.5-6.9)

69(64-74)

5.1 (4.7-5.4)

4

6.3 (6. 1-6.6)

58 (49-75)

5.4 (5.1-5.7)

10

4

6.6 (6.3-7.1)

69 (61-81)

4.9 (4.3-5.9)

4

6.2 (6.2-6.4)

67 (57-76)

5.6(5.3-6.0)

12

3

6.7(6.3-7.2)

71 (64-75)

5.2 (4.6-6.4)

3

6.2 (6.2)

71 (67-75)

5.8 (5.4-6.4)

13

3

6.7(6.1-7.2)

64 (50-76)

4.8 (4.6-5.5)

4

6.2 (6.0-6.4)

71 (53-82)

5.8 (5.5-6.4)

94 (91-103)

3.8 (3.5-4.3)

.

------------------------------------------------------------------------------b
15
3
7.1 (6.8-7.3)
103 (98-111)

a
Pre-trial,

fed concentrate diet ad libitum.

b
Fed dehydrated alfalfa cubes ad libitum.

3. 7 (3. 5- 3. 8)

4

7.0(6.8-7.1)

~
•.....•
0'1
I

�-417Rumen-fluid VFA concentrations and molar proportions of acetic:propionic
acids differed significantly

(p &lt; 0.05) among weeks uurlng the 90-day

feeding of Diet III (Table 2); however no consistent trends were evident
for either parameter.

Total VFA concentrations

in rumen fluid of deer fed

Diet III were similar to values (58-88 mM/l) reported by Short (1963) in rumen
fluid of white-tailed deer fed white cedar (Thuja occidental is) and big-tooth
aspen (Populus grandidentata).

Values measured in rumen fluid of deer fed

Diet III or alfalfa cubes were considerably lower than mean VFA concentrations
(125 mM/l) in rumen fluid of mule deer on natural browse (Nagy et~.
Molar proportions of acetic:propionic

1967).

acids in rumen fluid of deer fed Diet

III were higher than values reported by other researchers and are an indication
of the diet's poor quality.

Proportions of acetic:propionic

acids in rumen

fluid of deer fed dehydrated alfalfa cubes are similar to values reported
by Short for deer fed high quality diets.
concentration

It is interesting to note that VFA

increased on alfalfa hay diet.

lower in ADF and lignin than Diet III.

This diet was higher in protein,

One must recognize the fact that

VFA levels in the rumen are not only related to the rate of microbial activity
but also to the rate of VFA absorption through the rumen wall.
Serum Protein and Urea
No statistically significant trends in serum protein or in serum urea
levels were evident during the 13 weeks deer were fed Diet III. (Table 3).
Levels of both parameters were similar to values reported by other researchers.
No relationship between serum protein levels and general condition of deer
fed Diet III or alfalfa cubes was noted in this study (Table 3).

�Table

3.

Average

serum

urea

(mg/IOO

ml)

and protein

(observed ranges in parentheses)~

ml) in adult and fawn mule deer.

Adults

Week of
Trial

pt

(g/IOO

N

Serum

Urea

Fawns
Se rurn Protein

N

Serum

3

21.6 (15-35)

Urea

Serum

Protein

a

2

- - - .. - - -

3

6.6 (5-10)

-------8.4: (8.3-8.5)

3

-------21. 6 (20..,30)

7.9

(7.2-8.

5)

- - - - -- - - 6.6(6.1-7.2)

4
6
3
8

3

20

(15-25)

7.1 (6.6-8.2)
I

21. 6 (20-25)

+&gt;0
•....•

7.5 (7.0-8.5)

co
I

10

,

••
-r

21. 2 (20-:25)

6.5 (6.0-7.0)

--------

4

21. 2 (20-25)

6.4

8.1 (8.0-8.5)

4

12
13

3

ISb

21. 6 (20-25)

- - - - - - -

.3

aprc-trial,

30.0(25-35)
fed concentrate

bFcd dehydrrlted

alfalfa

7.1.(6.4-7.8)

diet ad libitum.

cubes ad libitum.

-------42.5

(30-60)

(6.0-7.0)

- - - - - - - 6.7 (5.3-8.3)

�-419Serum urea levels however, increased upon refeeding with alfalfa (Table
3).

Higher serum levels during refeeding may be related to two factors:

(1) increased deamination of diet protein by microorganisms in the rumen
and more ammonia absorbed into the blood, or (2) increased catabolism of
body protein.

Since higher blood urea levels were not associated with

rapid weight loss in three adults and one fawn, the increased blood urea
levels in these deer were most likely due to increased percent of crude
protein in alfalfa.

Three fawns that had significantly higher blood urea

levels than other deer also experienced rapid weight loss.

Exceptionally

high blood urea levels in these fawns were most likely due to body protein
catabolism but may also have been influenced by liver and/or kidney malfunction
as hypothesized by Keiss (1965).

��-421Chapter 3
Digestibility and Rate 0f Passage Trials
Introduction
In the previous experiment (Chapter 2) we found that deer under pen
conditions were able to survive and maintain their body weight, with only
minor changes when fed on a diet which is low in protein, high in ADF and
consequently of marginal nutritional value.

Protein is nutritionally

important and possibly low during winter in native forages available to
deer.

Protein supplements are, however, expensive to feed and one should

know the required levels of protein necessary in supplemental deer feeds to
help deer through starvation periods.

In this trial we investigated the

influence of different protein levels on dry matter digestion and rate of
passage through the deer digestive system.
Methods and Materials
Facilities
Four digestion cages 1.5 x 0.9 x 1.2 m high with 3/4-inch mesh flat
expanded metal floors were constructed.

Cages were built of 2.5 x 10.2 cm

boards with attached covered feeders and placed out of doors.

The top and

two sides were covered with canvas which afforded the deer some protection
from weather.

Window screen was placed beneath each cage to funnel feces

into collection pans beside each cage.

Urine was not collected.

Experimental Deer
Four tame mule deer does aged 1.5-7 years began this experiment, however,
one deer was injured and replaced during the second trial by a 9-10 month
old semi-tame female fawn.

To gradually condition deer to confinement,

they were placed individually in pens for 2 weeks prior to being placed in
digestion cages.

All deer rapidly became accustomed to close confinement

and presented only minor problems during the 10-13 day feeding trials.

�-422Trial Diets and Feeding
Compositions and proximate chemical analyses of the two diets are
given in Table 4.

Diet III was the ration fed during the previous trial

(Chapter 2) and Diet I was basically the same ration with added protein.
Both diets were fed in pelleted form and offered to each deer according
to 2 x 2 cross-over design.

Methods of producing feed, feeding and

chemical analysis were described in Chapter 2. Deer were accustomed to
the test diet for at least 8-12 days in isolation pens and 3-5 days in
digestion cages before daily food intake and fecal production were
measured.
Dry Matter Digestibility
Dry matter digestion of Diets I and III was determined by procedures
outlined by Maynard and Loosli (1962).

Feces produced by each deer were

collected at least 2-3 times daily for 5 days, placed in paper bags,
labeled and dried for 1-2 weeks at 60C. Apparent dry matter digestion
(percent) was calculated between difference of total intake and feces
produced on a dry-weight basis (Table 5).
Rate of Passage
Rate of passage of each diet was measured during the same 5 days in which
feces collections were made for determining diet digestibility.

General

procedures used were those described by Balch and Campling (1965).
Approximately 5 percent of one meal was stained with crystal violet.

To

minimize secondary staining of other food in the digestive system, the stained
diet was thoroughly rinsed with tap water and dried prior to feeding.

Time

required for stained food to pass through the deer digestive systems was
determined by visually identifying numbers of stained particles in samples
of feces collected at intervals post-feeding.

These data are presented as the

time interval between when 5 percent and 95 percent of all identified stained
particles had passed the digestive tract (Table 5).

�Table 4. Compositions and proximate chemical analyses of the two compounded diets fed in
digestibility and rate of passage trials.

Diet

Composition (percent)
32%
Protein
Cotton
Seed
Wheat
SuppleHulls
ment
Straw

Wheat
Bran

Crude
Protein

Analysis (percent, oven-dry basis)
Cell
Ether
Wall
MoisExContract
ture
Ash stituents ADF2

Lignin

I

+&gt;0
N

W
I

Diet III

75

5

15

5

5.5

4.9

8.5

5.2

74.3

52.8

15. 1

Diet I

70

30

--

--

12. 1

2.5

8.8

6.1

73.1

46.7

15.0

aAcid-detergent

fiber.

�-424Dry Matter Digestibility
Dry matter digestion of Diets I and III was determined by procedures
outlined by Maynard and Loosli (1962).

Feces produced by each deer were

collected at least 2-3 times daily for 5 days, placed in paper bags,
labeled and dried for 1-2 weeks at 60C.

Apparent dry matter digestion

(percent) was calculated between difference of total intake and feces
produced on a dry-weight basis (Table 5).
Rate of Passage
Rate of passage of each diet was measured during the same 5 days
in which feces collections were made for determining diet digestibility.
General procedures used were those described by Balch and Campling (1965).
Approximately 5 percent of one meal was stained with crystal violet.

To

minimize secondary staining of other food in the digestive system, the
stained diet was thoroughly rinsed with tap water and dried prior to feeding.
Time required for stained food to pass through the deer digestive systems
was determined by visually identifying numbers of stained particles in
samples of feces colleced at intervals post-feeding.

These data are

presented as the time interval between 'when 5 percent and 95 percent of all
identified stained particles had passed the digestive tract (Table 5).
Results and Discussion
Data on food consumption, food digestibility and rate of passage
show more variation among deer than between Diets I and III (Table 5).
Comparisons between diets are best made for the three deer completing both
tests.

�-425Daily food intake, apparent dry matter digestion, and rate of passage
were significantly greater for Diet I compared with Diet III for the three deer.
These differences are attibuted to added protein in Diet I as other measured
constituents of the diets were nearly equal.
Mean dry matter digestion coefficients for Diets I and III were
less than dry matter digestion coefficients of 46.1-53.4 reported by
Short (1966) for a compounded diet containing 41 percent crude fiber
and 20.2 percent crude protein.

Diets containing similar levels of protein

with less crude fiber had significatnly higher digestion coefficients and
Short conlucded there was an inverse relationship between diet crude-fiber
content and dry matter digestibi1ity.

Data from this experiment suggest

there was also a relationship between diet protein and dry matter digestion.
The lower level of protein in Diet III may have limited the activity of
microbial populations in the rumen.

Diet protein is a source of nitrogen

necessary for synthesis of microbila protein.

By adding protein to the

ration, as in Diet I, additional nitrogen was available and a greater population
of microbes may have been present for food digestion. Increased food
digestion enabled deer to pass food through their digestive systems at a
faster rate

which permitted intake of more food as observed with Diet I

vs. Diet III. Results of this experiment as well as the results of the
previous experiment (Chapter 2) suggest that 5.5 percent crude protein
in the diet is marginal and under normal circumstances, deer need higher
protein levels.

However, deer, at least under pen conditions, can survive

for long periods of time (90 days) on a diet containing 5.5 percent protein.

�Table 5. Mean daily food consumption, apparent dry matter digestibility and rate of passage of Diets I and III
fed to five mule deer.

Consumption
(grams)

Digestibility
(percent)

Rate of Passage,
5-95 Percent Time
(Hrs)

Animal No.

Diet I

Diet III

Diet I

Diet III

Diet I

Diet III

201

910

632

49.6

47.1

19.0

54.0

202

1197

1042

42.6

40.1

42.5

57.5

203

1445

1134

43.8

38.4

45.5

48.0

204

1134

--*

41.8

I

~

N

0)
I

243

1412

36.5
36.8

34.5

--------------------------------------------------------------------------------------------------------------Mean

1172

1055

44.4

*Animal injured during trial, substituted with No. 243.

40.6

35.9

51.0

�-427-

Chapter 4
Alfalfa Hay Feeding Experiments Using Deer,
Domestic Sheep and Goats
Introduction
There are numerous reports in the literature (Doman and Rasmusen,
1944; Smith, 1952; Carhart, 1943) which suggest that feeding alfalfa hay
to starved deer might be detrimental to the animal.

During our feeding

experiments (Chapter 2), deer, especially fawns, decreased their food
intake when fed good quality dehydrated alfalfa cubes and fawns lost 12-19
percent of their pretrial weight within two weeks (Fig. 4).
experiments (Nagy, et ~.,

Previous

1967) suggest that deer rumen microorganisms

are able to digest alfalfa hay in vitro as well as microorganisms

from

the rumen of cattle and sheep.
In the following experiments we wanted to investigate the abilities
of deer, domestic sheep and goats to digest poor quality alfalfa hay fed
in two physical forms.
12-day trial.

Alfalfa hay was fed first in pelleted form during a

In a second, 14-day trial the same quality alfalfa was

fed in cubed form.
Methods and Materials
Facilities
Sheep, goats and deer were individually confined in pens and digestion
cages.

Goats were confined to digestion cages due to insufficient numbers

of isolation pens.
Experimental Animals
Eight mule deer does (4 adults and 4 yearlings), four adult domestic
sheep and four adult domestic goats were used for each trial.

Two deer

(1 adult and 1 yearling) experienced severe digestive disorders and were
sacrificed when near death after eating pelleted alfalfa,

These animals

�428-

were replaced with similar-aged deer for the second trial.

Deer used

during each trial were born in captivity, raised by their does but were
not tame.

For approximately 9 months pretrial and during 2 weeks between

trials, deer were fed ad libitum concentrate and alfalfa hay.

At the

start of each trial deer were judged in excellent physical condition;
weight of adults was 55-70 kg and yearlings 53-61 kg.
Sheep and goats were crossbred adult ewes and nannies.

Sheep were

purchased 6 weeks prior to the trials, however goats had been used extensively in previous experiments.

During 6 weeks pretrial and 2 weeks

between trials, sheep and goats received ad libitum concentrate and alfalfa
hay and were also judged in excellent physical condition at the start of
each trial, weighing 53-66 kg and 25-48 kg respectively.
Trial Diets and Feeding
Alfalfa, cured in the field until mid-August, was used as the test
diet.

At cutting, most leaves had dried and fallen, leaving partially

green stems.

Stems were chopped, dehydrated and compressed into cubes

4 x 4 cm by 2-6 cm long. A quantity of these cubes was ground in a hammer
mill,

passed through a 3 mm screen and compressed into 6 x 12 mm pellets.

The alfalfa pellets produced were offered abruptly and fed ~

libjtym during

trial 1. During trial 2, unaltered alfalfa cubes were offered gradually
with decreasing amounts of concentrate for 4 days and then fed gg libitym
for 14 days.

The method used to measure daily food intake and procedures

for chemical analysis of the test diet (Table 6 ) are described and
referenced in Chapter 2.
was not measured.

Water was offered QQ libitum and daily consumption

�-429-

Table 6.

Chemical corriy o s it ion (oven-dry
in both pelletcd and cube forms.
Constituents

basis)

of alfalfa

Analysis

1.1

Ash

6.9

Crude protein

7.9

Ether

3.2

A cid-detergent-fiber

(ADF)

fed

(percent)

.Mo i st iir e

extract

stems

44.2

Lignin

9.3

Cell wall constituents

56. 1

Deer Necropsy
Two deer suffered severe digestive upsets after eating pelleted
alfalfa, would not respond to refeeding with concentrate,
sacrificed 'when near death.

and were

Blood samples were aspirated from the

jugular, placed in sterile vacuo-tubes containing EDTA and taken within
30 minutes to the Diagnostic Laboratory, Veterinary Hospital, CSU, for
analysis.

Ligations were made above the cardia and at the beginning

of the colon and the stomachs and small intestine with contents removed
and taken to the Diagnostic Laboratory, College of Veterinary Medicine,
CSU, for necropsy.

Four deer that evidenced similar digestive problems

after eating cubed alfalfa were also ~acrificed.
dimensions of the rumen-reticulum,

Weight, volume and

omasum; and abomasum were measured and

samples of contents taken from these organs.

All sheep and goats used

during the experiment and two deer that did not indicate digestive problems
were also sacrificed at the end of trial 2 and the same stomach measurements
made and silmples of contents taken.

�-430-

Stomach Measurements
Volumes of the rumen-reticulum, omasum and abomasum with contents
were measured by water displacement.

After all gas had been expelled

from the rumen, it was immersed in a container partially filled with
water and the displaced volume measured.

Volumes of the omasum and

abomasum, with contents, were measured by immersing each organ in a
partially filled graduated beaker and observing displaced volume.
Length, width and circumference of each omasum and abomasum were
measured to the nearest 3 mm with a flexible plastic tape.

Length

measurements were made along the center line of each organ by bending the
tape to conform with the organ's natural curvature.

Width and circumference

measurements reported are the greatest values obtained.

Once the abomasum

was opened, the pyloric sphincter was cut, laid flat and the inside
diameter measured to the nearest mm.
Weights of the rumen-reticulum, omasum and abomasum with contents
were measured after all fat had been removed from each organ.

After contents

had been removed and the tissue washed and blotted dry, each organ was weighed
empty.

The weight of contents within each organ was calculated by difference.

Histological Sections
Abomasal tissue was obtained for histological examination from
a sheep, a goat, a normal deer, and a deer that experienced an abomasal
disorder.

Samples were taken laterally through the fundus, in the pylorus

region 4 cm forward of the pyloric sphincter, and through the pyloric
sphincter.

Tissues were immediately fixed in 10 percent formalin and 2

percent sodium acetate and prepared by the Research Laboratory, Department
of Anatomy, CSU for interpretation.

Tissues were processed in graded strengths

of ethyl alcohol, cedarwood oil, xylene and embedded in paraplast.

Blocks

�-431-

were sectioned at 4 microns on rotary microtome, stained with hematoxylin
and eosin and mounted on microscope slides with paramount.

Photographs

and histological interpretations of abomasal tissues were made by Dr.
G. C. Soloman, Department of Anatomy.
Digestive r~easurements
The pH of rumino-reticular, omasal and abomasal contents of all
sacrifi ced animals was measured using a Fisher Accumet Model 210 pH
meter calibrated prior to use against standard buffer.

All readings

were made within 30 minutes after death.
Food contents within each stomach of all sacrificed animals were
samples to quantify degrees of food digestion.

Stomach contents were

thoroughly mixed and an approximately 100g sample removed, placed in a
small jar, and fixed in 10 percent formalin for laboratory analysis.

Food

particules in each sample were separated by size by gently washing the
sample through soil sieves of 4, 2, 1 and 0.42 mm.

Fractions separated by

each sieve were backwashed into containers, filtered through pre-weighed
Whatman FG/A glass filter paper, dried for 24 hours at 60C and weighed.
Filtrate containing food particles less than 0.42 mm was collected and its
volume measured.

After thorough mixing, three 100 ml subsamples of filtrate

were withdrawn, filtered and dried using the above methods.

Dry weights

for each of the four separated fractions were obtained after subtracting
filter paper weights.

Average dry weight of food particles less than

0.42 mm contained in the three 100 ml sub-samples of filtrate was related to the total filtrate volume and used to calculate weight of this
fraction.

Percent of each fraction was determined from total dry weight

of all five fractions.

�-432-

Data Analysis
Changes in food intake and among stomach measurements of deer,
sheep, and goats data were not analyzed statistically.

Data on food

particle sizes in stomachs of sheep, goats, deer with normal digestion
and deer with digestive upsets were analyzed using Duncan's New Multiple
Range Test (Li 1968:270).
Results and Discussion
Physical Condition and Food Intake
Six of eight deer fed pelleted alfalfa stems indicated varying degrees
of digestive problems and five of eight deer fed alfalfa cubes evidenced
similar digestive upsets,

Two deer and all sheep and goats fed both physical

forms of alfalfa showed no symptoms of digestive problems, remained in good
physical condition and maintained or increased daily food intake (Figs. 6. 7).
Digestive upsets in deer were indicated by an abrupt decline in food
consumption, usually with no food being eaten following 3 days of digestive
upset.

(Tables 7, 8).

At the onset of digestive problems, feces were sour

smelling and contai~d blood; defecatio~ also ceased within 3 days.

Other

visual symptoms of digestive problems were a dry nose, glassy~vacant
stare of the eyes, ruffled hair and a disinclination to move or get up
when approached.

If the condition persisted for 5 days or longer, the

animal was inclined to lie with head and neck outstretched on the ground.
While in this position small amounts of rumen fluid ran from the nostrils
or mouth,

When sacrificed. rumen fluid gushed from the mouths of these deer.

�· -433-

3500

3000

-

2500

(JJ

S

rd

H

2000

co
Q)

~
rd
....,
.....~

1500

1000

500

Fig.

6.

Mean daily food intakes of mule deer and domestic sheep
and goats fed pelleted alfalfa stems.
Figures in parentheses
indicate numbers of animals remaining on trial.

�-434-

3500

3000

2500
tIl

S

~2000
ee
Q)

~ 1500
~

500 -

o

1

2

3

4

5

6

7

8

9

10 11 12 13 14

Days

Fig.

7.

Mean daily food intakes of mule deer and domestic sheep and
goats fed cubed alfalfa stems.
Figures in parentheses indicate numbers of animals remaining on trial.

�-435-

TABLE 7.

Daily food intake of pelleted alfalfa stems by four adult
and four yearling mule deer does.

Day
of
Trial

Inta ke

(grams)

Adults

Fawns

1388

1759

2000

1988

1796

1168

1722

1652

2

1310

600

592

546

1346

1008

946

1016

3

1076

588

182

162

1254

914

700

72

4

1066

1350

0

0

910

1000

0

0

5

614

670

0

0

894

810

0

0

6

780

30

0

0

1280

1020

0

0

7

1200

0

0

0

846

1152

0

0

8

890

0

Oa

0

648

890

0

0

9

750

0

230

514

1086

0

0

10

206

0

280

504

1376

0

Oa

11

0

0

656

582

1310

0

12

Ob

Ob

690

613

1145

Ob

a

Did not respond to refeeding; sacrificed when very near death.

bResponded with refeeding of concentrate diet.

�-436-

TABLE 8.

Daily food intake of cubed alfalfa stems by four adult and
four yearling mule deer does.

Day

Intake (grams)

of
Trial

Adults

Fawns

700

1190

1506

1182

l300

1106

1108

790

2

376

856

974

510

1214

776

230

724

3

80

1018

930

178

860

1216

0

664

4·

377

922

1220

0

1200

860

0

466

5

758

1060

1161

0

1184

1276

0

688

6

772

1638

1404

0

1482

l366

0

832

7

120

1100

676

Od

800

456

0

670

8

0

1120

0

900

1120

0

390

9

1160

950

0

1076

882

0

790

10

600

588

0

1200

944

0

868

11

172

350

0

1068

1180

0

690

12

100

0

a

1146

1042

OC

974

13

14

0

0

14

Oa

ad

OC

1432
1344b

1362
1300b

1182
884

aResponded to refeeding of concentrate diet, however remained in
poor physical condition 6 months after trial
bSacrificed--normal digestion.
cSacrificed--omasum impacted.
dSacrificed--abomasum upset.

�-437-

Animal Necropsy
Physical condition of two deer declined so rapidly after ingesting
pelleted alfalfa they were near death within 10 days and were sacrificed.
Cursory examination during necropsy revealed no external or internal
injuries or malformations.
body fat.

Both deer were in good flesh and had considerable

Pathologist reports on digestive tracts listed the following

conditions for both deer: (1) no lesions or abnormalities were present
in the rumen or omasum; (2) multiple ulcers, inflammation and hemorrhage
were widespread in the abomasum with some necrosis, and (3) hemorrhagic
ulcers were prevalent around the pyloric sphincter and hemorrhagic enteritis
occurred throughout the small intestine.

Bacterial cultures prepared from

fluid taken from within the tissue lining of the abomasum were positive for
Escherichia coli and Clostridium perfringens.

Blood of one deer was

analyzed; results were: packed'cell volume 71 percent, red blood cells
24.8 x 106/mm3, white blood cells 2.0 x 103/mm3 , serum protein 9.1 g/lOO ml,
and serum urea 85 mg/100 ml.
Four of five deer that suffered digestive problems after ingesting
cubed alfalfa were also sacrificed.

Necropsy of two of these deer revealed

similar conditions within the abomasum and small intestine as in deer that
experienced digestive problems after eating pelleted alfalfa.

However, the

other two deer had greatly distended omasums containing solidly impacted
food particles.

Such a large amount of food was impacted into these organs

that blood capillaries within the organ muscle tissues ruptured.

Removing

irn:-&gt;acted
food from these organs revealed that some primary omasal leaves had
torn and were necrotic.

Inflammation, ulcers and hemorrhaging were also

observed within the abomasums and small intestines of these deer.
Omasal impaction has been reported in domestic ruminants but is difficult
to diagnose and has been observed only during necropsy.

Specific cases of

�-438-

impacted omasums in dairy cows have been reported by Hughes and Cartwright
(1962) and Blampied et~.
during this~udy

(1964).

Symptoms of impacted omasums in deer

were similar to those described for domestic ruminants

(Blood &amp; Henderson 1968: 107) .
Necropsy of sheep, goats and the two deer that did not show symptons
of digestive problems revealed no ulceration or inflammation had occurred
within the animals digestive tracts. Digestive organs in these animals
appeared to be functioning normally.
Histological Sections
Histological sections of abomasal tissue from a sheep, a goat,
and a deer with normal digestion indicated no qualitative structural
differences among the three species.

Examination of abomasal tissue

from a deer experiencing digestive problems revealed extensive abrasion of
the mucosal

lining, blood capillaries within the muscle tissue were

dilated, and abnormally high concentrations of lymphocytes were present.
With abrasion of the protective abomasal mucosal lining, underlying blood
capillaries were closer to the lum~n of the abomasum and more susceptible to
injury.

It is possible that dilation of blood capillaries was caused by

infection or a bacterial toxin.

The abnormally high concentration of

lymphocytes in the tissue indicates infection and a possible bacterial invasion.
Anatomical t1easurements
Measurements of the rumen-reticulums, omasums, and abomasums of
sheep, goats and deer show obvious size differences among the three species.
Relative to body size, weights of rumino-reticular contents account for about
10 percent of body weight in deer compared with 17-20 percent for sheep and
goats (Table 9). The omasum and abomasum are also smaller in deer compared

�-439-

with these organs in sheep or goats (Tables 10, 11).

Figure 11 shows

relative sizes of omasums and abomasums from a sheep, a goat, a deer
with normal digestion and a distended omasum of a deer due to food impaction.

Differences in relative organ sizes can be seen readily in this

figure. Degree of distension was indicated by the differences

between

weights, volumes, and circumferences of impacted omasums and normal omasums
(Table 10).

Abomasums from deer with abomasum upsets also indicated these

organs were larger than normal deer abomasums (Table 11).

Mean body wc i glrt s and weight, volume, percent dry matter
and pH of rum.ino- reticular
contents of mule deer and do~e stic sheep and goats fed cubed alfalfa stems (number of am-

Table 9.

mals in parentheses).
Contents

Rum.ino- reticular

(1)

Dry
Matter
(percent)

pH

37.2

19.0
18.5

13.3
7.6

13: 3
14.6

6.4
6. 6

54.5
60.8
59.0

10.6
12.8
10.4

9.9
12.9
7. 1

14. 1
2.7
6. 1 .

6.2
7. I
7..1

to sacrifice;

rumen

Mean
Body
Wt. (kg)

Weight
(percent
body wt.)

Vol.

Sheep (4)

51.7

Go at s (4)

Deer
no r mal d i ge stion (2)
a
o rri a s u i'n impa ctec1 (2)
abom.asum. up s et" (2)

Animal

a Deer

did not eat for

tents very fluid.

3-8 days

prior

con-

�-440-

Table

10.

Mean size 'of the omasums and weight, percent dry matter
and pH of omasal contents of mule deer and dome stic sheep
and goats fed cubed alfalfa stems (number s of animals in
parentheses).

Omasum Size
(with contents)

Vol.
(ml)

Sheep (4)
Goats (4)
Deer
. normal digestion (2)
omasum impa ct ed" (2)
abomasum up s e t+ (2)

Animal

ci-.

Omasal

( g .)

Dry
Matter
(percent)

pH

125
101

757
530

17.5
19.8

7.2
6.2

50
109
49

314
980
107

20.6
18.6
13.0

6. I
7.0
6.8

W.

Wt.

(mm)

L.
(mm)

(mm)

861
546

271
297

189
155

239
1132
126

154
350
138

88
205
100

o

aDeer did not eat for 3-8 days prior

Contents

to sacrifice.

Digestive Measurements
Percent dry matter of rumino-reticular,

omasal and abomasal contents

of deer with normal digestion, sheep and goats all fed alfalfa cubes was
similar (Tables 9-11).

Percent dry matter in rumino-reticular

and omasal

contents of deer with digestive upsets was less than levels in these organs
of normal deer.

Dry matter levels in abomasal contents of deer with digestive

upsets were, however , higher than levels in normal deer, sheep or goats.
The pH of omasal contents of deer with impacted omasums and sheep
with normal digestion were neutral or alkaline; however acidic pH readings
were observed in omasal contents of goats and deer with normal digestion
(Table 10).

Abomasal contents pH in deer with digestive upsets was higher

than pH readings in animals with normal digestion (Table 11) indicating
cessation of normal acidic digestion.

�Table 11. Mean size of the abomasums and weight, percent dry matter,
and pH of abomasal contents
of mule deer and domestic sheep and goats fed cubed alfalfa stems (numbers of animals in
parentheses).
Abomasum

Abomasal

Size (w.fh contents)
Inside Dia.
Pyloric
Sphincter
(rom)

Contents

I

Wt.
(g)

Dry
Matter
(percent)

pH

+&gt;0
+&gt;0

.....•

Vol.
(rol)

Cir.
(rnm)

(rom)

W.
(rnm]

Sheep (4)
Goats (4)

1570
843

280
260

437
338

134
121

61
52

1134
759

12.7
13.3

3. 5
3.4

Deer
normal digestion (2)
omasum impacteda (2)
aborna sum upseta (2)

448
215
661

206
174
222

.267
244
277

82
71
95

48
44
52

521
212
633

12.4
15.7
17.0

5.8
6.5

Animal

aDeer did not eat for 3-8 days prior

L.

to sacrifice.

I

3. 3

�-442-

Size of food particles in rumino-reticular contents of sacrificed animals
would be affected by time between ingestion and sacrifice for each animal,
a variable that was not controlled in this experiment.

Percent distribution

of particle sizes by weight in rumino-reticular, omasal and abomasal
contents did not vary significantly among normal deer, sheep or goats(Table 12).
Distribution of particle sizes in the stomachs of deer with digestive
problems differed significantly from distribution in stomachs of normal deer
(Table 12).

There were fewer large particles in rumino-reticular contents

of deer with digestive problems than in contents of normal deer.

This was

expected since deer with digestive problems did not eat for 3-10 days prior
to sacrifice.
Impacted deer omasums contained a higher proportion of food particles
larger than 1 mm than did normal deer omcsums; abomasums of deer with either
impacted omasums or upset omasums contained more particles larger than 0.4
mm than did abomasums of normal deer (Table 12).

These data suggest a failure

of mechanisms regulating passage of large particles from the rumen-reticulum of
some deer fed alfalfa.

It is not known if this failure was the cause or the

effect of the digestive disorders observed.

�Table

12.

Distribution

of Cood particle

size.

mule deer lcd cubed a.l!alfa. stems

(percent

by weight)

in rumino-reticular,

omasal

and abomasal

cont ent s o{ domestic

Rumen •.r e t ic u'Iu rru Food Particle
(percent)

Orn r surnr

Siz.c

Food Particle
{pe r c e nt]

Abomasum:

Size

&lt;0.4
nun

0

2.5
1.4

15.0
5.80

36.7
37.8

40.0
55.-1

0
0
2.5

1.4
5.4
6.6

8.0
16.2
18.2d

31.5
38.2
35.9

59. d
40.3c
36.9°

&lt;0.4
mm

&gt;4.0

2-4
mm

1-2

0.4-1

rom

mm

mm

&lt;0.4
mm

&gt;4.0
mm

9.4
9.7

8. I
5.9c

24.6
18.6

30.5
32.6

27.4
33.2

0
0

2.2c
1.3c

12.9c
5. Ice

39.6
38.0

45.4
55.7d

·0

$C.cep (4)
Goat s (4)
"
Deer
normal digestion
(2)
omasum
impacteda
(2)
ab or-ia s urn upset" (2)

11.4d
2.7e

9 •.2
13.5b
9.5

20.3
29.5
26.2

2? 3
37: 1
31. 4

29.8
17.229.2

0
0
0

2.0c
11.5b
4.4b

11.0e
24.0b
15.4cd

35.8
39.4
36.1

51. 3
25.0°
44.2

eMean.b

Multiple

Range

TeBt.

di{{er

.ig"i!icantly

2-4

to .acrince.

(p &lt; 0.01)

{rom mean.c,

and mcansd

di.!Cer signilicantly

(p

&lt; 0.05)

Si:te

nun

0.42-1
mm

did not eat for 3-10 .days prior

Food Particle
(percent)
0.4-1

1-2
mm

b, c.d.

and

1-2
mm

2-4
mm

aDeer

goat.

mm

&gt;4.0
mm

3.70

.heep.

and sacrificed.

e

{~om means

according

to Duncan'.

Ne'f

i

..,.
..,.
I

W
I

�-444Inability of deer to digest alfalfa stems fed during this experiment
appears to be related to stomach size and morphology.

Deer have small

stomachs, especially the omasum and abomasum, that are not adapted for
digestion of bulky, high-roughage diets.

The structure of a deer's rumen,

reticulum and omasum is similar to the same organs in other ruminants with
selective food habits.

Hofmann (1968) noted ruminants with selective food

habits have a small rumen with weak pillars, the reticulum was comparatively
large and contained shallow crests, and the omasum was "extremely small,
kidney shaped," and contained few laminae.

Sizes and structures of the

rumens, reticulums and omasums in sheep and goats were, however, similar
to these organs in ruminants with non-selective food habits that are adapted
for digestion of high-roughage diets.

Hofmann (1968) noted the rumens in

"roughage eaters" were very large with powerful pillars, the reticulums
relatively small and contained deep crests, and the omasums were large,
spherical and contained numerous laminae.

To compensate for their small

stomachs ruminants with selective food habits require forage that is more
easily digested.

Rate of food passage may also be faster in animals with

small stomachs than in ruminants with non-selective food habits.

During

this experiment deer that experienced digestive problems may have been
unable to retain particles of coarse alfalfa stems in their rumens long
enough to permit adequate digestion.

Incomplete food digestion resulted

in large, abrasive food particles being passed through deer digestive tracts
that resulted in physical damage to the abomasum and small intestine mucosa
(Fig. 8).

�GOAT

SHEEP

DEER
NORMAL

DEER
OMASUM

IMPACTED

0

6
12

~j;I('~;:;;~~
'"' I "
r.:

I,',

'I

"

'I' .\
-.
~;tl:'
\ :")".'"/."')~,J'"
fl

I'

.'

.

\'\'

','

'\~\'{/ / i:

18

bi :&gt;,::~.

24

I
h~")
l'j"'i'6

(J)

a:: 30
w
I-

w 36
~
I-

1

' .• 'i

(

~'

~

"
"

./. ;

I
..j::&gt;

"I .

~

d:\

f.il~1,
(I,,\..'
-..:;

(J"I

.

42

z'

w 48
u

54
60
66

72
FIGURE

8. Relative sizes of goat, sheep and deer omasums and abomasums (after a photo by Nagy)

I

�-446-

Table

13.

Chemical composition
mountain hay.

(oven-dry

basis)

Constituents

of pelleted

Analysis

(percent)

Moisture

13.4

Ash

5.8

Crude protein

12. 1

Ether

. 5.8

extract

Acid-detergent-fiber

(ADF)

43.6

Lignin

4.8

Cell wall constituents

58.0

TABLE 14.

native

Daily food intake of five mule deer fed pe11eted native mountain hay.

Day of
Trial

Intake(grams)

1

1210

1420

1100

1130

2

1194

1620

1256 .

780

3

1332

1182

792

n04

688

4

1370

730

402

88

486

5

1336

994

914

0

956

6

1436

974

750

0

976

7

1690

1214

1020

0

1258

8

1736

1190

824

82

1124

9

1428

1322

884

48

1276

10

1860

1308

936

270

1048

11

1344

1116

582

590

794

12

1460

1138

966

1286

690

13
]4

1060
1566

1010
1280

788
768

1234
1260

1084
1050

1060
,

1240

�-447-

Results and Discussion
One deer evidenced digestive problems but recovered within 4 days and
had a normal food intake w~en the trial ended (Table 14).

The only symptoms

of digestive problems evidenced in this deer were an abrupt decline in food
intake and a dry nose.

Daily food intakes of four deer (Table 14) not

evidencing digestive problems were approximately the same as intakes of two
deer that did not experience digestive problems when fed alfalfa (Tables 7,8).
Deer fed native hay did not experience digestive problems to the
same degree as deer fed pelleted alfalfa during Experiment 3.

Reasons

why deer were able to digest pel1eted native hay better than pelleted
alfalfa are unknown, but may be related to structural differences due
to lignification.

Alfalfa contained more lignin (9.3 percent) than did

native hay (4.8 percent).

Higher lignification in alfalfa may be related

to structural characteristics that resulted in alfalfa being less
digestible and more abrasive to the digestive tract than native hay.
Digestive problems observed however cannot be entirely the result of
a high percent of lignin in alfalfa.

Feeding trials of mule deer fed

browse diets of mounta in mahogany (Cercocarpus montanus), bitterbrush
(Purshia tridentata) and big sagebrush (Artemisia tridentata) containing
22, 26 and 30 percent lignin respectively (Short and Remmenga 1965) did not
result in digestive problems.

��-449Chapter 5
Alfalfa and Native Hay Feeding Experiment
Introduction
During the previously described feeding trials it became clear that
deer can experience digestive problems when fed poor quality alfalfa hay.
It has been observed that an abrupt change in dietfrom a concentrate to
alfalfa hay can cause digestive problems in deer as evidenced by decreased
food intake for several days (Chapter 2; Nagy et~.

unpublished data).

When deer were fed native hay (Chapter 5) they did not experience severe
digestive problems.

These results suggest that deer may be responding

differently to various amounts of alfalfa or native hay in their diets.
Investigations to determine what amounts of alfalfa or native hay contained
in a deep ration that would result in digestive problems were therefore
conducted.
Methods and Materials
Facil ities
Deer were confined in individual pens through the trial.
Experimental Animals
Eleven adult mule deer (5 castrated males and 6 females) weighing 60-75
kg and judged to be in good health were used.

All animals had been raised

in captivity and were fed ad libitum a high quality ration including alfalfa
hay for 3 months before the experiment.
friaLDiets

and Feeding

Five rations (Table 15) including the norma lly fed deer ration(control)
were ground and pelleted.
hay.

The control ration was concentrate without

For the four test rations alfalfa and native hay were each substituted

to provide 37 and 75 percent of the diet.

Alfalfa hay used was a leafy.

third cutting and the native hay was also of good quality. containing
mostly tirnothy(~hleun:!.
pratense) and sedge(Carex spp.). All five rations con-

�-450-

tained similar levels of protein (Table 16).

Except for lignin content,

the diet containing 37 percent alfalfa was similar to that containing 37
percent native hay and the,diet containing 75 percent alfalfa was similar
·to that containing 75 percent native hay (Table 16).

Table 15. Compositions of five pelleted
and native hay feeding trials.

rations used in the alfalfa hay

ComEosition (Percent Air-Dry v7eight)
Corn
Protein SUPP'.!..1
Native Hay

Ration

Barley

Control

60

20

2.0

0

0

N 37

38

12.5

12.5

37

0

N 75

15

5

'5

75

O·

A 37

38

12.5

12.5

0

37

A 75

15

0:

5

0

75

oJ

Alfalfa

1/
- 32 percent protein supplement.
Table 16. Chemical composition
native hay feeding trials.

of five rations used in the alfalfa hay and

Hay

�-451-

Deer were divided into two groups with six receiving the diet containing
alfalfa and five receiving the diet containing native hay. A 3 x 3 Latin
Square design was used in the experiment, the-rations being randomized among 3
groups of deer in 3 consecutive 7-day periods (Tables 17 and 18).
sequence, the diet was changed after each 7-day period.
by each deer was measured daily.

In each

Food consumption

Throughout the experiment water was

available ad libitum and intake not measured.
Data Analysis
Average daily food intake of the last five days of each 7-day feeding
period was analyzed for each ration using the procedure outlined by Dixon
and Massey (1969:310).

Comparison between the hays was made also using

Dixon and Massey (1969:175).
Results and Discussion
None of the deer on either alfalfa or the native hay containing
diets showed ~ny symptoms of digestive problems other than minor adjustments
to the abrupt changes in diet.

Average daily food intake of deer in both

the alfalfa and native-hay series showed no significant (P&gt; 0.10) effects
due to periods(columns), animals (rows), or to percent of hay in the ration
(treatments) (Table 19).

However, sample sizes are small, variation among

deer is large and treatment effects are confounded with the sequence in
which rations were fed.

Further, the short feeding period (7 days) may

not have allowed sufficient adjustment to the rations.

�-452-

Table 17. Average daily food intake (grams) of six deer fed alfalfa-hay
rations during 7-day periods !/. Rations control (0), A 37 and A 75 contained
0, 37, and 75 percent alfalfa, 'respectively.
Deer
Number

Period
I

II

III

o

A 37

A 75

5

392

,1078

a54

6

913

442

1253

Mean

652

760

1053

37

A 75

o

1

1466

260

1078

2

1945

,'226

1522

Mean

1706

243

1300

A 75

o

A 37

3

590

1103

861

10

1102

1028

,'962

Hean

846

1065

911

A

1/

o (control), overall mean intake = 1006 g.
A 37, overall mean intake = 1126 g.
A 75, overall mean intake = 714 g.

�-453-

Table 18. Average daily food intake (grams) of five deer fed native-hay
rations during 7-day periods !/. Rations control (0), N 37 and N 75 contained 0, 37 and 75 percent native hay, respectively.
Dee r

NUillber

Period
I

II

III

0

N 37

N 75

972

1851

1948

N 37

N 75

o·

7

298

1182

761

8

·1931

1526

947

Mean

1114

1354

854

N 75

0

N 37

--

138

1542

11

4

1450

9

1371

186

1115

Mean

1410

162

1328

-.

1/
-- 0 (control), overall mean intake = 601 g.
N 37, overall mean
75&gt;

intake

1347 g.

overall mean intake

1495 g.

,

�-454Table 19. Analyses of variance of data from the alfalfa and native-hay
feeding trial.
Source

df

Mean Square

Alfalfa Hay Rations
Total

8

Co1unms (Periods)

2

151361.8

0.44

Rows (animals)

2

51357.4

0.15

Treatments (% Hay)

2

134469.4

0.40

Residual

2

340280.4
Native-Hay Rations

Total

8

Colunms (Periods)

2

55598.1

O.lq

Rows (animals)

2

321018.11

0.57

Treatments (% Hay)

2.

217153.45

0.38

Resi~ua1

2

566183.4

1/
- F ratios for coluons, rows, and treatments are to be conlpared with
F.90 (2,2) = 9.00.

Throughout the experiment (Tables 17 and 18) deer consumed more (P 0.10) of the
rations containing native hay than of the rations containing alfalfa.

This dif-

ference in consumption was greater for the rations containing 75 percent hay than
for the rations containing 37 percent hay.

Greater consumption of native hay may

be related to the higher concentrations of lignin in the alfalfa-hay rations(Table
Results of this experiment are by no means clearcut.

They suggest, however,

that deer have difficulty with higher levels of alfalfa hay in the diet even when
the hay is of good quality.

This does not occur with native hay.

Results also

suggest that a gradual increase of hay levels is more desirable when feeding deer
with diets containing hay than an abrupt change from low to high levels of hay.

16).

�-455-

Chapter 6
Rate of Food Passage Through the Digestive
Tract of Deer and Sheep
Introduction
Food intake is related to the rate of passage of the ingesta and rate
of passage can be expressed by the retention time, the time the ingesta remains
in the digestive tract.

Rentention time would depend on rate of digestion,

more digestible food, e.g., low fiber diet would pass through the tract faster,
and on physical form of the diet e.g., a diet which is of small particle size will
pass through faster.

Rentention time is also characteristic for the particular

species of animals.
In earlier experiments deer experienced digestive problems on diets containing high proportions of alfalfa hay fiber.

We hypothesized that sheep, as

the fiber (alfalfa hay) content of the diet increases will eat more, rate of
passage through the digestive tract will increase and retention time will
decrease.

The opposite would be true in case of deer.
Methods and Materials

Fac ilities
Deer and sheep were kept in individual pens describe9 previously
(Chapter 1). During the experiment animals were placed in 3 x 4 x 6
foot metabolic cages designed to collect fecal material.
Ex.~e!,_imen.!.a
1 An ima1s
Six yearling mule deer raised in captivity by their dames and six
y ..
:d,'llng

crossbred sheep were used in the study. Sheep were purchased 6

weeks prior to the beginning of the trial and received the same diet as
the deer, ad libitum concentrate and alfalfa hay.

�-456-

Design
Four pe11eted diets were formulated

(Table 20) with 0, 15, 35 and 50

percent alfalfa hay mixed in the ration.

Table 20.

Chemical analysis of four diets fed during rate of passage

trials.

..

Chemical Analysis
Crude
Ether
Fiber
Extract

NEF

Diet

Percent
Alfalfaa

Protein

1

0

13.8

2.4

18.6

52.9

II

15

13.8

2.3

20.1

51.0

III

35

13.4

2.1

22.8

47.7

IV

50

13.5

2.0

24.9

45.3

a

The remalnlng portions of the compounded rations consisted of 40 percent
barley, 30 percent cottonseed hulls, 20 percent of 32 percent protein
supplement and 10 percent corn.
b

Nitrogen free extract.

b

�-457-

Animals were placed in the metabolic cages and were allowed to adjust
for ten days to cage conditions and to the diet (basically similar
to their ad libitum diet) which contained no alfalfa (control).

After ten

days, when food and water intake appeared stable the animals were given
a 0.5 gm portion of food containing 2 to 5 microcuries of chromium - 51
(51 Cr C13).

Fecal collection began six hours post-spike and continued at

two hour intervals to twenty hours when the interval was changed to four
hours.

Four hour collections were maintained until 64 hours post-spike.

Eight hour collections were made from 64 hours until 96 hours when trials
were concluded.

Subsequent trials followed this scheme with dietary pro-

portions of alfalfa increasing from 0 to 15, 35 and 50 percent alfalfa in
the pelleted ration.
alfalfa trial.

Control diet was fed again after the 50 percent

Prior to initiation of each trial, animals were conditioned

for ten days to the diet.

Food and water were given ad libitum.

Urine samples

collected from two animals contained no chromium -51.
o

After collection, fecal samples were placed in a dehydrator (163 F) for
24 to 48 hours then weighed to the nearest 0.1 gram.

Samples were

homogenized and slurred with 0.5 N HCl for comparison to liquid standards
of similar density and configuration and assayed with 5 x 5 inch No. I
crystal attached to a single channel analyzor.

Duration of counts was such

that the standard deviation of the difference between the sample ~nd the
background did not exceed 5 percent of the net sample count rate.
Net counts per minute were converted to microcuries per gram of dry
sample and plotted as a net cumulative excretion curve.

The mean retention

time was calculated using the 5-95 percent elimination method of Castle
(1956) ,

�Table

21

Daily feed consu~ption, dry matter digestion and ~ean retention
fed 0, 15, 35, and 50 percent alfalfa diets.

(n=6)
DEER
Daily Fe~d Consump t.ion (g)
Appa ren t Dry I,ratterDigestion
}fean Retention Time (hrs)
(n=6)
D3ily Feed Cor.sumption (g)
App ar en t Dry Matter Digestion
Hean Retention Time (hrs)

(%)

time for deer and sheep

Trial III
35% Alfalfa
~'lean
SE

Trial IV
50% Alfalfa
}~eon
SE

991

2.3

944'
147
1.8
64.,4
1.3
21.3

73
1.3
1.9

1660
66.8
20.6

Trial I
0% Alfalfa
SE
Mean

Trial II
15;~ ~lfa1fa
SE
Bean

698
121
70.8
0.8
1.3
23.6 '

933
71. 7
23.5

135
0.9
2.0

67.4
2' '

_ ...•

LO

1277
69.3
32.5

1453
72·.0
29.7

84'
1493
68.6
0.8
2.2 . 25.4

135

Trial V
0% Alfalfa
}!ean

c::'C'

w •.••

672

72

--

-2.1

23.1

SHEEP

(%)

114

1.0
2.1

84
1.1
0.7

1413
30.2

83

1.4

I

.+:&gt;
(J'1

00
I

�-459-

Derived mean retentin times and food intakes are given in Fig.
21 respectively.

9 and Table

Included in these tables are results of a fifth trial during

which deer and sheep were fed a diet containing no alfalfa hay. These trials
indicated that, increased amounts of hay in the diets did not influence intakes
and mean retention times which were characteristic for the two species at the
beginning of the trials.

Comparing mean retention times to percent alfalfa

hay in the diet showed that sheep had a statistically significant (P&lt;O.Ol)
decrease in the mean retention time of marked particles with an increase of hay
in the diet (Fig. 9).

In contrast, deer showed a very slight, however sig-

nificant (P&lt;0.05) decline in mean retention times associated with increase of
hay in diet.

Mean retention times of deer were significantly different from

those of sheep for diets with 0 percent(P&lt;O.Ol) and 15 percent (P&lt;0.05)alfalfa,
but not for diets with 3 percent (P&lt;0.25) or 50 percent(P&lt;0.05) alfalfa. Sheep
exhibited a significant (P&lt;0.05)increase of food intake with increased amounts
of hay in the diet.
Food retention time in the digestive tract, rate of passage through the
digestive tract and food intake are closely related.

As the fiber content of

ground and pelleted diets increases, retention time of food in the digestive
tract of many ruminants may decrease with a corresponding increase in food
intake.

This means that per unit time there is more food passing through

the digestive tract of animals receiving a poor quality diet. For this reason
the animal's energy supply is somewhat compensated.
this trial with sheep.

This is well documented in

Deer, however, were unable to increase their food

intake on these diets; consequently, as fiber content of the diet went up they
must have received less and less energy per unit time.

In our previous ex-

periments (Chapter 4) it was shown that there are important differences in the
rations of rumen, omasal and abomasal size to body size between deer and sheep.
It is possible that deer having relatively smaller rumen, omasum and abomasum
than sheep are unable to increase rate of passage of certain high fiber foods.

�-460-

(/,

1-

::J

o
s:

~
w
:::?:
I-

39
?-8
37
36
35
34
33
32 ::31

Z

30 29

I-

28

Z

26

o
w

I-

w

0::
Z
c:l:

w
:::?:

2"1 -

25
24.
23

DE_E_R_
--

_

22

:~['
19
18

o

35

15

PERCENT

ALFALFA

50

HAY

IBOO
1700
1600 -

SHEEP

1500 1400
(/I

E
o

1200

1-

0&gt;

1100-

W

1000

&lt;l:
I-

900

Z

BOO I""

~

700

DEER_---

---

__

-------

---

»&gt;

C--

600
500 400
300
200L-------------------~---

o

15

--------------~~--------------~
35

PERCENT

ALFALFA

50

HAY

Fig. 9 - Mean retention time and food intake changes of sheep and deer on
diets with varying percentages of alfalfa hay_

�-461-

Discussion
During the previous experiments (Part I) we tried to gain some basic
information on the digestive ability of mule deer on different concentrate rations, on different hay rations and we compared certain characteristics of deer
with that of sheep and goats.

We have shwon that deer rumen microorganisms can

digest alfalfa hay and deer can utilize the end products of this fermentation
without ill effects.

Deer not in starved condition and given good quality

(leafy) alfalfa hay may select between the leafy and stemy parts and survive
on leafy alfalfa hay since it is digestible and nutritious to other ruminants.
An abrupt change to a diet containing high proportion of hay might upset the
digestive processes of deer.
We have shown, however, that deer are much more susceptible to digestive
upsets (often fatal) if the hay is of poor quality than are sheep or goats.
Sheep, when increasing amounts of hay are offered in the diet, are able
to increase their food intake to compensate for the poorer quality diet.
Deer were unable to do so. Differences in the digestive organ sizes
between deer, sheep and goats might be the prime reason for this.

In the

wild, deer feed on alfalfa hay when they come down to haystacks, and in
some supplemental feeding programs alfalfa hay is the principle supplement.
Sudden change to this type of a high-fiber diet might further excelerate an
already deteriorating physiological state.

Deer might then develop serious

digestive upsets and complications such as impaction of the omasum and/or
in;ection of the abomasum, and consequently die.

��-463-

PART II
PHYSIOLOGY OF DEER STARVATION
General Introduction to Part II
In the past most attempts to feed starving deer occurred after
some deer had begun to die and often were the results of pressures applied
by lay groups to save the remaining deer (Severinghaus, 1963; Keiss and
Smith) 1966; Giles and McKinney 1968).

Generally, such. attempts were

hastily conceived and conducted with little regard to how or if
supplemental feeds .met qualitative and quantitative nutritional needs of
starving deer.

In nearly all cases, deer died after days or weeks of

continual feeding (Carhart 1943, Doman and Rasmussen, 1944, Hesselton
1964-65).

These results have led most observers to conclude that starving

deer cannot be fed to prevent or minimize winter losses (Carhart, 1943;
Doman and Rasmussen, 1944, Cheatum, 1956, Haynes, 1967, Giles &amp; ~1cKinney,
1968; Kelsey, 1973).
In this part of the overall study we made an effort to test
various hypotheses regarding the failure of deer to respond to emergency
feeding. We tried through controlled experiments to imitate starvation
conditions to find out what physiological changes occur in deer during
starvation, and we tried to investigate ways to prevent or minimize
starvation losses.

�-464Chapter 1
Effects of Starvation on Deer Rumen Bacteria
Introduction
In this series of experiments we investigated the possibility that
deer, during prolonged starvation may lose their functional rumen microbial
population through various means.

Chief among our hypotheses were that:

1) microorganisms may die due to starvation or 2) become inactive and unable
to digest food when refed.
Methods and Materials
Facilities
Facilities used during these experiments were described in Part I.
Experimental Animals and Design of the Experiments
A pilot study was initiated on September 25, 1970, investigating the
hypothesis that deer rumen microorganisms die or lose digestive function
when deer starve.

Five adult mule deer were sacrificed one each after 0,

7, 14, 21 and 28 days of starvation in 1.8 x 6.1 m isolation pens at
the Colorado State University Foothills Campus.
Counts of viable bacteria and dry matter digestion (DMD) of substrates
in vitro by bacteria from rumen fluid of starved deer were compared
between the deer sacrificed at

° days starvation and deer sacrificed after

7, 14, 21 and 28 days of starvation.
There appeared to be differences of counts of viable bacteria and
DMD between starved and non-starved deer in the pilot study.

A second trial

was begun on September 23, 1971, with larger sample sizes and concurrent
investigation of how deer maintain large numbers of viable, functioning

�-465rumen bacteria during starvation.

It also was desired to measure effect

of starvation upon ability by deer to digest various components of feeds.
This was accomplished by estimating digestion of cell wall constituents
(cellulose, hemicellulose and lignin) (CWCO) and cell contents (soluble
carbohydrates, fats and proteins) (CCO) of substrates by rumen bacteria.
Techniques associated with analysis of cell wall constituents are regarded
as more accurately representing the cellulose, hemicellulose and lignin
fractions of feeds than those of the Weende analysis crude fiber fraction.
Deer were starved to death in pens with 3 different floor types (Table 22).
One non-starved deer was sacrificed at 20 days 'after initiation of starvation
of experimental deer, another at 30 days and a third at 40 days.
Comparisons of counts of viable bacteria and of related in vitro
digestive abilities of rumen fluid from fed and starved deer were used to test
the hypothesis that rumen bacteria in starved deer die or lose the ability
to digest substrates.

Comparing these parameters among groups I, II and

III deer (Table 22) provides insight into mechanisms deer might use to
preserve viable rumen bacteria during starvation.

If group I deer exhibit

higher counts of viable bacteria and higher in vitro digestion than deer
in groups II and III, reinfection by rumen bacteria from eating infected soil
or feces is suggested as a mechanism for preserving viable rumen bacteria
during starvation.

If these parameters for groups I and II are similar and

higher than those for group III deer, eating soil and f~ces might sustain
rumen bacteria during starvation.

If these parameters for groups I and II

are similar and higher than those for group III deer, eating soil and feces
might sustain rumen bacteria during starvation by providing a nutrient source.
If no discernible differences in bacteria-population

parameters exist among

groups J, II and III deer, other mechanisms must account for maintenance of
high counts of viable, functioning rumen bacteria in starved deer.

�-466-

Table 22.

Starvation regimes, fall 1971 trial.

Group

n

I

4

2-31m x 8m pens, dirt floor

Water, soil, feces

II

2

2-6.1m x 1.6m pens, concrete
floors

Water, sterilized
soil

III

3

3-6.1m x 106m pens, concete
floors

Water

IV1

3

1-40m x 40m pen, dirt floor

Water and feed

Holding Facil ity

lNon-starved, control deer.

Potential Sources
of Exogenous Bacteria

�-467Collection of Rumen Fluid
The peritoneal cavity was opened immediately after death of a deer.
The rumen was removed and its contents thoroughly mixed.

A 300 ml sample

of these contents was withdrawn and placed into a pre-warmed thermos.

These

contents were strained through double-layered cheesecloth and used in
culturing and in vitro digestion determinations.
recorded.

Rumen contents' pH was

Volume of the remaining rumen contents was measured.

Counts of Viable Bacteria
Hungate's (1966:23-29) techniques for culturing rumen bacteria generally
were followed.

A broad-spectrum-agar medium allowing growth of cellulolytic,

proteolytic and amylotic bacteria was used.

Nine ml of autoclaved culture

medium were anaerobically pipetted into autoclaved 15 x 180 mm glass tubes.
These tubes were stoppered and immersed in water at 55 C.
Eight successive 1:10 dilutions were made of raw, strained rumen fluid
with sterile, isotonic diluting fluid, progressing from pure rumen fluid to
8
1:10 parts rumen fluid:diluting fluid. Three 1 ml aliquots from each
dilution were anaerobically inoculated into three of the prepared culture
medium tubes.

Inoculated tubes were gassed with CO2, stoppered immediately

and rolled horizontally under a stream of tap water at approximately 5-10 C
until the medium gelled, forming a translucent film of uniform thickness
about the inner surface of the tube4

Rolled tubes were incubated in an

oven at 38 C for 48 hours, after which visible bacteria colonies embedded
in the medium were counted for each tube. Multiplying mean number of
bacteria colonies times the dilution factor of that dilution produced an
average of 30-300 visible bacteria colonies.

This yielded an estimate of

number of viable bacteria per ml raw rumen fluid.

Multiplying this number

by volume of rumen contents gave an estimate of number of viable bacteria
in the rumen.

�-468-

Three control test tubes containing culture medium were treated
similarly and incubated to serve as a check against bacterial contamination
during culturing.
Digestive Ability of Rumen Bacteria
Percent DMD was determined using two-stage techniques of Tilley and
Terry (1963) as modified by Pearson (1970).

Percent DMD was estimated

by the equation:
- CTR x 100
Percent DMD = DM - DMDR
DM
DM = Dry Matter
DMDR = Dry Matter Digestion Residue
CTR = Control Tube Residue
Percent CWCD was determined by combining techniques of Van Soest and
Wine (1967) with those of Pearson (1970).

Three 0.25 g oven-dried samples

of each substrate were subjected to Tilley and Terry two-stage digestion.
Tubes containing digested substrates were then centrifuged at 50,000 rpm
for 10 minutes.

Centrifugate from each tube (DMDR) was then subjected

to neutral detergent digestion (Van Soest and Wine, 1967).

Percent CWCD

was estimated by the equation:
Percent CWCD = CWC - CWCR - CTR x 100
CWC
CWC = Cell Wall Constituents
CWCR = Cell Wall Constituents Residue
Percent CCD was estimated by combining DMD and CWCD results in
the equation:
Percent CCD = (OM - CWC) - (DMDR - CWCR)
OM - CWC

x 100

Pilot Study
Counts of Viable Bacteria
Counts of viable bacteria per m1 of rumen fluid from starved deer
declined 1-2 orders of magnitude from those of the non-starved control

�-469Table 23.

Diets used in study.

Chemical Analysis
Crude
Protein

Fat

CWC1

NFE2

Diet I

18.0%

3.7%

26.1%

58.4%

Diet II

15.0%

2.0%

42.0%

51.0%

Diet III

14.0%

4.0%

23.2%

57.1%

Diet IV

3.8%

1.3%

52.0%

39.3%

Alfalfa

15.0%

1.7%

33.5%

38.3%

lCWC = cell wall contents
2NFE = nitrogen free extract

�-470-

deer (Table 24).

However, starved deer had over one billion viable

bacteria (estimated) in their rumens when sacrificed.
Digestive Ability of Rumen Bacteria
DMD of alfalfa by bacteria in rumen fluid of starved deer did not
decline from that of the control deer.

DMD of Diet I by bacteria in rumen

fluid of starved deer appeared to decline progressively as length of
starvation increased (Table 24).
Rumen Contents Volume
Rumen contents volume of the non-starved deer was 5535 ml, mean volume
for starved deer was 2089 ml (Table 24).
Fall 1971 Trial
Starved deer began to die after 16 days starvation; by 47 days the
last deer had died.
Counts of Viable Bacteria
Counts of viable bacteria per ml rumen fluid from Group I starved
deer (allowed to eat soil and feces, presumably infected with rumen bacteria,
during starvation) were not significantly different (P &gt; 0.1) from those
of non-starved Group IV deer.

Total number (estimated) of viable bacteria

in rumens of Group I deer was significantly lower (P ~ 0.001) than that of
Group IV deer (Table 25).
Counts of viable bacteria from rumen fluid of deer in Groups II
(allowed to eat autoclaved soil and feces during starvation) and III
(starved without any source of solid matter intake) were lower than those
of Group I deer (Table 25).

Unfortunately, counts of viable bacteria for

two of three Group III deer were biased by contamination.

Resulting small

sample sizes of deer in Group II (2) and Group III (1) prohibited tests of

�Tab 1e 24.

Deer ru.oen fi u .j parameters.

pil ot study 1970.

Counts of
Viable Bacteria
Per ml
Tota 1 number
rumen fl uid
~er rumen

In Vitro
Digestion of Dry
Matter {~ercentl
Alfalfa

Diet I

Rumen Contents
Volume (ml)

1.2xlOll

69

76

5535

65

71

3050

4.1xl05

1.3xl010
1.6xl09

--

--

3840

1

--

--

74

63

2210

1

4.9xl05

9.8xl09

66

58

2000

Days
Starved

Number
Deer

01

1

7

1

2.1 xl 07
4.4xl06

14

1

21
28

~
'-J

.....
I

lNon-starved control deer

�Table 25.

Group
Number

Deer rumen fluid counts of viable bacteria, volume and pH, fall 1971 trial.

n
-

Days
Starved

Counts of Viable Bacteria
Per ml rumen
Total number
6
fluid (xl0 )
per rumen (108)

Rumen Contents
Volume (ml)
~H

X

SD

X

SD

X

SD

X

SD

4

20,22
22,29

37.4

39.6

534.81

500.9

1605'

430.6

7.' ,

0.3

II

2

16,22

2.9

0.9

28.3

21. a

1251

915.0

7.2

0.3

III2

1
3

47
31,37
47

0.9

--

36.0
1936

1534.7

7.2

O. 1

3

0

34.6 '

IV3

7.4

2066.81

I

404.1

59801

237.3

6."

lDifferences between means significantly different (P&lt;0.005).
2Culture counts of two of three deer contaminated,
for all deer in this group.
3Non-starved control deer.

but rumen volume and pH measurements

collected

O. 1

+=&gt;
.....•
N
I

�-473differences of viable bacteria among Group I, II and III deer.

All

starved deer died with over one billion (estimated) viable bacteria in
their rumens.
Digestive Ability of Rumen Bacteria
Differences in alfalfa CWCD among Groups I, II and III deer were
statistically significant (P ~ 0.1).

There were not other statistically

significant differences (P &gt; 0.1) in DMD, CCD or CWCD between Group I and
IV deer, nor among Group I, II and III deer for any of the three diets
(Table 26).
Lack of significant differences (P &gt; 0.1) between digestive abilities
of rumen bacteria from starved Group I deer and non-starved deer and among
groups of starved deer (excepting alfalfa CWCD) prompted examination of
the relationship between length of starvation and digestive ability of rumen
fluid bacteria (Fig. 10).

Regression of DMD, CCD and CWCD of all diets on

length of starvation produced slightly negative slopes, but 90 percent
confidence intervals of these Slopes included zero (Table 27).

Thus, DMD,

CCD and CWCD of diets by bacteria in deer rumen fluid did not decline
significantly as length of starvation increased.
In Vitro Digestion of Different Diets
Differences between alfalfa and Diet II DMD, CCD and CWCD were not
statistically significant (P &gt; 0.1) whether determined within Groups I,
II, III and IV or for all deer.

DMD, CCD and CWCD of Diet I were

significantly higher (P ~ 0.01) than those of alfalfa and Diet II when
determined within deer groups and for all deer.

Higher digestion coefficients

for Diet I probably reflect its lower concentrations of cell wall
constituents (Table 23).

�Table 26. ~

vitro digestion of three diets by rumen fluid from Group I-IV deer, fall 1971 trial.

�-475-

CELL

WALL

70

/

P

/

60 - --

--

50
40 L'r--=

--=--~-

m--

__

-

!!II-

-

-

-Eli--..g/

G-----&lt;1

20

/

~

~
&lt;,

Die t I
Alfalfa

-;

'(,-----

--

--"

5

10

/

~

~DietII

Control

/

"-,,-

&lt;,

---:----....
-IE

/

&lt;,

''0-

30

/

-it----II

20

15

CELL

25

30

C!

45

40

35

CONTENTS

100

-------~.

90

-------

--------81_
-

-ir-'tr/rt-~----~----II!-----=

-"'- ~
- --

-------~

80

------8---0-'

\

-

z;

o 70 f-t

&gt;-1

~

::

J

IS-----l'l!

Diet

O-------('j

Alfalfa

!::s.

Control

!'J

...--Li

11

~

\

//

I

I:::. Diet IT

5

10

15

20
DRY

25

30

40

35

45

MATTER

l

---.--70
60
50
40

8!l-----J!

Diet I

0----0

Alfalfa

6---6

Diet

,,

-,

!::s.
,

,,

/'

'",/

IT

L
Control

5

_10

15

20
DAYS

25

30

35

40

45

STARVED

Fig. 10. Effect
of~tarvation
on in vitro digestion
of three
by deer r urn e n fluid.
Cont1"ol deer were fed ad Libi tu rn .

diets

�Table 27. Slopes and associated 90 percent conf1dence intervals for regression of in vitro DMD, CCD
and CWCD of three diets on length of starvation.

Slope 2

DMD
-0.24

Confidence
interval

+0.32

Alfalfa
CWCD
_.-

DMD

Diet I
CWCD

-0.09

-0.13

-0.14

Diet II
DMD
CWCD
----0.05
-0.09

+0.38

+0.44

+0.19

+0.24

--

-0.18

CCD
--0.32

+0.32

+0.54

CCD

--

+0.53

CCD
0.07
+0.26

lDMD = dry matter digestion; CWCD = cell wall constituents digestion; CCD = cell contents
digestion.
2Change in percent digestion per day of starvation.

..,.
I

'-J
O'l
I

�-477-

Rumen Contents Volume
Mean volume of rumen contents for Group IV deer was significantly
higher (P 20.01)

than that for Group I deer.

This could possibly mean

that during starvation deer reduce their water intake.

Differences of

mean pH among Group I, II and III were not significant (P &gt; 0.1) (Table 4).
Chapter 2
Discussion-Effect of Starvation
on Rumen Bacteria
Counts of Viable Bacteria
Deer starved in the pilot study in Chapter I retained fewer (1-2 orders
of magnitude) viable bacteria per ml rumen fluid than the non-starved
control deer.

In the fall 1971 trial there was no significant difference

in numbers of viable bacteria per ml rumen fluid between starved and
non-starved deer.

The 11 starved deer from 1970 and 1971 trials retained

.9-37 million viable bacteria per ml rumen fluid at death.

These numbers

were well within ranges (2 million-23 billion) of number of viable bacteria
cultured from rumen fluid of non-starved domestic ruminants (Hungate 1966:
34-35).

Volumes of rumen contents for deer starved in the pilot study

and fall 1971 trial were reduced from one-half to one-fifth of control
deer.

Resultant estimate of total number of viable rumen bacteria from

starved deer was significantly lower than that for non-starved deer.
These results agree with pertinent literature indicating that number
of viable bacteria declines during starvation.

However, there is no

evidence demonstrating total loss of viable rumen bacteria during starvation.
Rather, deer in this study retained over one billion viable rumen bacteria
during 7-47 days of starvation.

�-478Indirect evidence that viable bacteria are not eliminated from rumens
of starved deer was provided by results from spring trials during 1971,
1972 and 1973.

Of 41 deer starved 10-64 days, 39 were successfully refed,

indicating that, at the end of starvation, deer retained numbers of viable
bacteria sufficient to provide metabolites, upon refeeding, required for
sustenance.

Ozoga and Verme (1970) reported successful refeeding of deer

starved for six weeks; this success required survival of rumen bacteria
during starvation.
Unfortunately, culture techniques utilized in this study did not
permit specific estimates to be made of numbers of viable proteolytic,
amylolytic or cellulolytic bacteria existing in rumens of starved deer.
Likewise, species composition of bacteria cultured from deer rumen
fluid was not determined.

Change in composition of bacterial species

.occurring during starvation was likely, since mean pH of rumen fluid from
starved deer (7.2) was significantly higher than that from non-starved
deer (6.1). Rise in pH reflects a reduction in production of volatile
fatty acids, an expected result of starvation.
Digestive Ability of Rumen Bacteria
DMD, CWCD and CCD of diets by bacteria in rumen fluid from deer
starved 16-47 days were not significantly less than those for non-starved
deer.

These results do not contradict results from starvation of domestic

ruminants.

Suppression of VFA production cited by Coop (1949) and Quinn

(1943) lasted 3-10 days but was associated with voluntary restriction of
food intake during the first few days of refeeding.
expected to be lower with restricted intake.

VFA levels would be

Slow fermentation rates

occurring upon resumption of feeding cited by Quinn (1951) are not comparable
with data from this study, which measured absolute digestion rather than
rates of digestion.

Meiske et £1. (1950) reported severely restricted

�-479-

cellulose digestion in vitro associated with starvation but digestion
trend lasted for only 24 hours.

Additional 24 hours of digestion

allowed in this study may have permitted cellulolytic bacteria to
undergo logarithmic growth, increasing their numbers sufficiently to
permit maximum digestion of cellulose.
Above cited ability of deer starved 10-64 days to survive refeeding
implies that starved deer maintained sufficient numbers of viable
proteolytic, amylolytic and cellulolytic bacteria to provide required
metabolites upon resumption of feeding.

These results demonstrate:

1) that rumen bacteria retain digestive abilities during starvation of
deer; and, 2) that this ability includes digestion of different feed
components.

This latter statement is important, for it provides evidence

that starving deer retain adequate numbers of rumen bacteria capable of
digesting major substrate categories (i.e., cell wall constituents and
cell contents) and producing quantities of essential metabolites required
for sustenance.
In this study, reduced number and digestive ability of viable rumen
bacteria from starved deer were not significantly lower biologically nor
(in the case of digestive ability) statistically than those from non-starved
deer.

It is doubtful that wild deer experience total starvation (complete

abstinence from food) as long as some of the deer in this study.

These

results suggest that during starvation wild deer retain qualities and
quantities of rumen bacteria at refeeding sufficient to provide them with
required metabolites to sustain life.
Retention of Viable Rumen Bacteria During Starvation
It is argued that counts of viable bacteria from Group II and III deer
are not biologically different from those of Group I deer, based on

�-480-

lack of differences (excepting digestion of alfalfa cell wall) among
digestive abilities of rumen bacteria from these groups to digest three
diets (Table 26).
Group III deer were unable to reinfect their rumens with live bacteria
and had no access to organic matter other than their hair, sloughed rumen
epithelium and soil, apparently retained in the rumen, during starvation.
Thus, starving deer may not depend upon reinfection to maintain rumen
bacterial populations.

It had been assumed that bacteria were removed from

rumens of starving deer, with passage of ingested water, to the lower tract.
Rumen fluid volume of starved deer was less than half that of non-starved
deer, probably reflecting a sharply-reduced water intake during starvation.
Since starved deer maintained large numbers of viable rumen bacteria in this
trial, it is postulated that most of the ingested water was absorbed through
the rumen wall with little passing to the lower tract.

Thus, bacteria were

not lost from the rumen by passage, suspended in water.

Church (1969) cited

studies indi.cating that rumen motility decreases in rate and strength in
fasted sheep.

Perhaps rumen motility in fasted deer ceases entirely, curtailing

passage of water from the rumen.
Wild ruminants host species of rumen microorganisms similar to those
of domestic ruminants (Pearson 1965, 1969, McBee 1969).

Of microbial

genera found in domestic ruminants, only one, Clostridium, sporulates under
adverse conditions such as starvation.

Thus, deer may contain only one

genus of sporulating bacteria in their rumens.

However, rumen bacteria may

survive starvation without sporulating.
Lamanna and Mallette

(1959:617) and Ensign (1970) suggested that

microorganisms resistant to starvation should have evolved.

Barnes and

Burton (1970) demonstrated that counts of viable cecal bacteria from
thirteen-lined ground squirrels (Citellus tridecemlineatus) at 6 and 42

�-481days of hibernation were of the same magnitude as those from non-hibernating squirrels, indicating that cecal bacteria are capable of surviving
cold and starvation stresses.

Ensign (1970) demonstrated 65 percent

viability of Arthrobacter cyrstal10poietes, a non-sporulating soil
bacteria, at 60 days of starvation.
Strange et~.

(1961), Dawes and Ribbons (1965) and Sierra and Gibbons

(1962) demonstrated that starving bacteria utilized endogenous carbohydrates
and proteins.

When stored nutrients are depleted, starving bacteria may

remain viable by metabolizing products from lysed bacteria (Postgate 1967).
Harrison (1960) postulated that this cryptic existence might provide minimal
nutritional requirements for starving bacteria in the transition to a
stationary, viable state devoid of exogenous nutrients.

Finally,.

"cells which survive for a sufficient time might be able to adjust to
greater resistance through loss of the characters of logarithmic phase cells
and acquisition of characteristics of maximum stationary phase cells."
(Harrison 1960:427).

Ensign (1970) and Burleigh and Davis (1967) demonstrated

that bacteria drastically reduce endogenous respiration within 1-2 days of
starvation, significantly reducing rate of autodegradation.

Possibly, rumen

bacteria in starving deer may use all available nutrients in the rumen and
then revert to a state of minimal metabolic activity, remaining viable until
deer eat again.
Digestion of Different Diets by Rumen Bacteria
Rumen bacteria from starved deer exhibited significantly higher ~
digestion of Diet I than of any other diets.

vitro

The greater the digestion of a

feed, the greater the amounts of metabolites provided to the host organism.
Feeding Diet I to deer starved 10-64 days produced no mortality nor digestive
upsets.

Diet I is thus a logical candidate for feeding starving wild deer.

��-483-

Chapter 3
Physiology of Deer Starvation and Refeeding
Starved Deer
Introduction
In this chapter attempts were made to simulate the physiological
changes which take place in deer during starvation.

These changes were

monitored, and deer at different stages of starvation were refed.
Methods and Materials
Facil ities
Facilities for conducting starvation and refeeding trials, including
isolation and multiple-animal pens and apparatuses for weighing deer
and collecting rumen and blood samples were described in Part I. Six
8 x 31 m pens and two 31 x 31 m pens, located at the Colorado Division of
Wildlife Regional Research Center in Fort Collins, Colorado also housed
starved and refed deer used in this study.
A laboratory suitably equipped for conducting digestion trials and
bacterial culturing was provided by the Department of Animal Sciences,
Colorado State University.

A second laboratory, wherein analysis of blood

parameters was performed, was provided by the Department of Physiology
and Biophysics, Colorado State University.
Design
A pilot study, initiated March 26, 1971, investigated the feasibility
of starving and refeeding deer.

Preliminary information sought was:

1)

definition of a point during deer starvation beyond which feeding fails to
prevent deat~; 2) ability of deer to regulate intake during refeeding; and
3) survival of starved deer when refed.

�-484-

Twenty-one adult and yearling deer were transported from the Little
Hills Research Station, Colorado Division of Wildlife, located near Meeker,
Colorado and placed in two 12.3 x 18.5 m pens at the Colorado Division of
Wildlife research facility, Fort Collins, Colorado.

These deer were fed

Diet I (Table 2) and alfalfa hay ad libitum for 2 weeks while deer
acclimated to the pens.
Beginning March 26, 1971 these deer were starved.

At 10, 15 and 22

days of starvation respectively, three deer, selected randomly, were
separated from the herd, placed in 1.8 x 6.1 m isolation pens and refed ad
libitum Diet II (Table 23), which was formulated to simulate levels of crude
protein, crude fiber, and nitrogen free extract (NFE) expected in native
forage.

All deer dying during the trial were necropsied.

Feed intake of

refed deer was recorded for 12-18 days.
The pilot study indicated that the starving and refeeding of deer could
be studied under penned conditions.

Pilot study results indicated that

3 weeks of starvation might approximate a point beyond which feeding might
not save starving deer.
A second study was initiated in late winter 1972 to test a number of
hypotheses.

The first (Hl) was that fawns would succumb more quickly than

does to starvation and exhibit higher refeeding mortality.
comparing

This was tested by

mortality rates of fawns and does for the two periods.

The second

(H2) hypothesis was that deer fed before 20 days starvation would exhibit
lower refeeding mortality than deer refed after being starved longer than
20 days.

Again, comparing mortality rates for the two periods sufficed to

test the hypothesis.

A third hypothesis (H3), that ammonia and/or urea
toxicity results from feeding diets of high protein content, to starved deer,

was tested by feeding two diets differing in protein content to starved deer

�-485(Diets III and IV: Table 23).

Higher refeeding mortality and blood urea

nitrogen levels among deer fed Diet III would support the hypothesis.
A fourth hypothesis (H4) was that starvation induced liver necrosis or
malfunction, resulting in fatal metabolic disruptions.

Elevated blood levels

of glutamic oxa1acetic transaminase (GOT) and ornithine carbamyl transferase (OCT)
provide positive identification of liver necrosis; blood levels of these
enzymes were monitored in starved and refed deer to determine whether or not
liver malfunctions resulted from refeeding.
An additional reason for feeding starving deer two diets was to compare
refeeding performance (weight gain, mortality, intake) of starved deer on
2 diets of different qualities.
On February 29, 1972 jugular blood samples and body weights were obtained
from 13 doe and 13 fawn mule deer.
m pen and starved.

These deer were then placed in a 31 x 31

Water consumption was ad libitum.

According the original

sampling design, two does and two fawns were to be refed after la, 17, 24, 31,
and 38 days of starvation, assuming that several deer would die before they
could be refed.

At each refeeding date, one fawn and one die were to be fed

Diet III and the remaining doe and fawn were to be fed Diet IV.
By the 24th day of starvation, none had died subsequent to refeeding
and remaining starving deer appeared strong and in no immediate danger of
dying.

One of the objectives of the trial was to present deer of nutritional

states ranging from good (starved only ten days) to extremely poor (starved
almost to death) to refeeding.

It was anticipated that under the original

design, considering the apparently good condition of remaining starved deer,
deer in extremely poor condition might not be produced prior to refeeding.
Therefore, after the 24th day of starvation deer were not refed until they had

�-486-

lost approximately 30 percent of pre-starvation body weight, a figure
several investigators stated to be indicative of deer near death (Davenport
1939, Doman and Rasmussen 1944).
Results of the 1972 spring trial failed to substantiate hypotheses
linking refeeding failure to development of toxicity or liver malfunction
subsequent to refeeding.

Failure of hypotheses tested in this study to clarify

the issue of refeeding starving deer resulted in a final set of hypotheses,
tested in winter, 1973.

One possibility is that starving wild deer may not

have received enough supplemental feed when fed in the wild to alleviate
negative energy balance.

Accordingly, final hypotheses tested revolved

about investigation of deer energy metabolism during starvation and refeeding.
The first hypothesis (H5) tested was that nutritional plane of deer prior to
starvation influences their ability to survive starvation and to successfully
adjust to refeeding.

This was tested by comparing mortality during starvation

and refeeding between deer fed ad libitum for 6 weeks prior to starvation and
deer of severely restricted intake during the same 6 week period.

The second

hypothesis tested (H6) was that deer dying during 1972 and 1972 spring starvation
trials depleted their energy reserves, entered negative energy balance
and were unable to maintain homeostasis.

This was tested by comparing blood

levels of energy-metabolites between deer dying of starvation and deer
surviving starvation.
Intakes of 13 deer placed in isolation pens and fed Diet III ~
were recorded from January 4, 1973 to January 13, 1973.

libitum

Five deer were

selected as control deer, remaining deer were matched to control deer on the
basis of mean feed consumption and body weight.

A six week conditioning

period was conducted from January 16 to March 1, 1973.

Experimental deer

�-487-

received one-fourth the previous day's ad libitum intake of their matched
control deer prior to total starvation.

This conditioning period was designed

to produce two groups of deer: control, representing deer on a high
nutritional plane; and experimental, representing deer in a semi-starved
condition.

All deer were starved at termination of conditioning until any three

deer died, whereupon remaining deer were refed Diet III ad libitum for 10
days.

Jugular blood samples and body weights were obtained at initiation of

conditioning, at 3 weeks of conditioning, at initation of starvation, at
initiation of refeeding and 10 days post refeeding.
recorded during refeeding.

Daily feed intake was

Body weight and a jugular blood sample were taken

from dying deer when possible.
Sampling Procedures
Body weight.

Weighing apparatus was described in Part I. Deer were

forced through an alleyway into a darkened chute narrowing to two 1.1 x 0.33
x 1.2 m plywood boxes.
scale.

The first box was mounted directly over a platform

Closure of doors on both ends of this box forced deer to stand upon

the scale, whereupon their weight was recorded.
Blood samples.

Deer were forced from the weighing box into the second

box fitted for immobilizing the head and neck.

Immobilization was accomplished

by pinching a deer's neck between two wooden, fur-cushioned clamps after
the animal had passed its head through a 20 cm diamater opening.

The clamps

were attached to the box by hinges, permitting the neck to be stretched
sufficiently for easy withdrawal of blood (Fig. 2).
Forty ml jugular blood samples were withdrawn with a 50 ml syringe.
Approximately 0.05 ml blood was drawn from the syringe tip into a heparinized
capillary tube.

Remaining blood was placed immediately into a 50 ml centrifuge

tube containing sodium-fluorine-heparin.

The tube was capped, everted gently

several times to mix blood and heparin and kept in an ice bath until taken

�-438-

to the laboratory.

Forty ml blood samples were centrifuged for 5

minutes at 3000 rpm and resulting plasma frozen.

Capillary tubes containing

heparinized blood were centrifuged for 5 minutes at 3000 rpm.
Blood Parameters Examined
Toxicity indicators.

Plasma urea nitrogen levels were determined from

blood samples collected during 1972 and 1973 spring trials.

Techniques

and reagents used are described in Sigma Technical Bulletin No. 535 (Sigma
Chemical Co. 1971).

Evaluation of plasma ammonia levels was desired as an

index of ammonia toxicity.

However, variations of time interval between

drawing and freezing blood (1-4 hours) among deer resulted in unequally
biased blood ammonia levels (Tietz 1970:665), rendering valid comparisons
of blood ammonia levels among deer impossible.
Liver malfunction.

Plasma levels of two liver enzymes (OCT and GOT)

were determined from 11 deer in the spring trial of 1972.

Deer selected

for sampling were two non-starved deer, four starved deer, four refed deer
and the one deer dying during refeeding.

Techniques followed and reagents

used in determining plasma levels of the two enzymes were described in Sigma
Technical Bulletin Nos. 108 (OCT) and 55-UV (GOT) (Sigma Chemical Co. 1971-1972).
Energy metabolism.

Levels of plasma glucose, urea nitrogen and free

fatty acids were determined from deer in spring trials, 1972 and 1973.
Duplicate plasma urea nitrogen (PUN) levels for each blood sample
were determined according to procedures outlined above.
Duplicate plasma free fatty acids (FFA) levels for each blood sample
were determined according to techniques of Mosinger (1965) and Dole and
Meinertz (1960).
Duplicate plasma glucose levels for each blood sample were determined
using a Beckman glucose autoanalyzer.

�-489-

Other parameters.

Hematocrits of deer from spring trials in 1972 and

1973 were determined by placing centrifuged capillary tubes containing
heparinized blood in an International Micro-capillary Reader (International
Equipment Co.) and reading hematocrit from a sliding scale.

Two drops of

plasma from each centrifuged capillary tube were placed onto a Goldbert TIC
Refractometer (American Optical Co.) and optical density recorded.

Plasma

total protein levels were obtained from conversion tables based on optical
density.

Hematocrit and total protein values were obtained in hopes of

making fortuitous comparisons with other blood parameters and with deer
performance during starvation and refeeding that might provide further insight
into the area of investigation.
Parameters from free-ranging deer.

As part of the overall Middle Park

Deer Study the Colorado Division of Wildlife was collecting data on a
number of mule deer physiological characteristics during winter.

Heparinized

plasma samples were obtained from 12 wild does on winter range in Middle Park,
Colorado between March 7 and April 21, 1972.

Plasma glucose, FFA and PUN

levels of these deer were determined according to procedures outlined above.
Results
Pilot Study
Mortality DUring Starvation,
Deer began to die after 6 days of starvation; by 28 days of starvation
all deer that had been refed or had died (Table 28). All deer refed after 10
and 15 days of starvation survived refeeding as did two deer starved 22
days.

One deer, starved 22 days, died the third day of refeeding.

animal was extremely weak when refed.

This

During refeeding it lay on the ground

with head and neck outstretched, arising only to eat.

�-490Intake During Refeeding
Daily intake during refeeding did not differ significantly (P &gt; 0.1)
among deer refed after 10, 15 and 22 days of starvation.

Mean daily intake

by refed deer began at approximately 525 g, remained at this level for a
second day, doubled by the third day and levelled off at about tripled the
first day's intake by the seventh day.
Spring 1972 Trial
Mortality During Starvation
Deer began to die after 33 days' starvation; by 64 days all deer
had been refed or had died (Table 29).
Comparison of Refeeding Performances of Does and Fawns
Weight changes during starvation and refeeding.

Starved fawns lost weight

at approximately double the rate of starved does (Fig. 11); this difference
was statistically significant (P ~ 0.01).

No significant differences

(P &gt; 0.1) existed between rates of weight gain by does and fawns during
refeeding.
Starvation mortality.

Four fawns died during starvation; one before it

could be refed (starved 33 days) and three (starved 33, 34, 34 days) within
hours of refeeding.

The latter three fawns were considered dying when refed,

based upon their extremely weakened condition.

None was able to walk without

falling; all had to be carried to isolation pens for refeeding and struggled
only feebly.

Mean loss of body weight by these four fawns was 29.8 percent.

One doe died after 54 days' starvation, losing 37 percent of its body weight
(Table 29).

Fawn mortality during starvation (30.8 percent) was significantly

(P ~ 0.1) higher than that of does (7.7 percent).
Refeeding mortality.
refeeding.

Of 12 starved and refed does, only one died during

This doe had been starved 56 days and lost 31.9 percent of its

�-491Table 28.

Fate of starved and refed deer, spring 1971 pilot study.
Days
Starved

Deer

Final Disposition

Adult female

6

Died sixth day of starvation.
malnourished

Appeared

Adult female

10

Died tenth day of starvation.
slightly malnourished.

Appeared

Adult females (3)

10

Refed Diet II. SSR1.

Yearling male

11

Died 11th day of starvation.
malnourished.

Appeared

Adult female

12

Died 12th day of starvation.
malnourished.

Appeared

Yearling male

12

Malnourished.

Adult females (2)

13

Died 12th day of starvation.
Died 13th day of starvation.

Malnourished.

Year1ing ma 1e

14

Died 14th day of starvation.

Malnourished.

Adult female

15

Refed Diet II. SSR.

Adult males (2)

15

Adult female

16

Refed Diet II. SSR.
Died 16th day of starvation.
malnourished.

Adult females (3)

22

Yearling male

23

Adult f ema1e

27

Died 27th day of starvation.
malnourished.

Grossly

Adult male

28

Died 28th day of starvation.
malnourished.

Grossly

lSSR

Survived starvation and refeeding.

Grossly

Refed Diet II. One died after two days
on feed; grossly malnourished. Others
SSR, and appeared grossly malnourished.
Died 23rd day of starvation. Grossly
malnourished.

�-492Tab1 e 29.

Deer 1

Fate of starved and refed deer, spring

Days
Starved

Diet
Refed

1972 trial.

Bod1 Weight Changes
During
During
Starvation
Refeeding
---3
-19.3
+ 3.03

Final o isposition
SSR4

MF2

10

III

FF5

10

IV

- 9.7

+ 10.1

SSR

10

III

-10.4

+

0.8

SSR

IV

- 7.4

-

1.4

SSR

Doe
Doe

10

FF

17

III

-19.8

+ 11.4

SSR

17

IV

-14.6

+

2.6

SSR

III

- 7.5

+

2.0

SSR

17

IV

-24.6

+ 17.0

SSR

24

III

-24.0

+ 4.0

SSR

-

5.6

SSR

FF
Doe
Doe
MF

17

FF

24

IV

-11.9

Doe

24

III

- 7.1

0.0

SSR

24

IV

-14.9

- 13.6

SSR

33

III

-27.4

33

IV

-19.8

33

None

-29.5

III

-15.3

+

2.6

SSR

-

1.6

SSR

Doe
MF
FF
MF
Doe

33

Died 3.5 hours
after being refed
+ 4.1

SSR
Died before
could be refed

Doe

33

IV

-16.4

MF

34

IV

-33.9

Died within three
hours of refeeding

III

-28.1

Died within five
hours of feeding

35

III

-30.0

+

7.5

SSR

35

IV

-28.6

+

5.3

SSR

IV

-29.7

+

5.4

SSR

FF
MF
MF
Doe

34

36

�-493-

Tabl e 29. Continued
Bod~ Weight Changes
During
During
Starvation
Refeeding

Days
Starved

Diet
Refed

Doe

54

none

-37.0

Doe

55

III

-26.0

Doe

56

IV

-36.3

Doe

64

IV

-24.2

Deerl

Died before
could be refed
+

+

2MF = male fawn.
3percent body weight.
4SSR = survived starvation and refeeding.
FF = female fawn.

6.0

SSR
Died 5th day of
refeeding

lOne animal per entry.

5

Final Dis~osition

6.0

SSR

�-494-

0

f:j.

Does

••

Does

0

Fawns

e

Fawns

dying

t r i aI

during

dying

during

trial

o

f:j.

Do e s- .

o

o

Fawns:

Y = -0.81X
.z , 0.75
f:j.

o

35

~------~------~

-L

o

30

10

20

~

~

40

50

60

DAYS STARVED
Fig.
11. Loss of pre - starvation
bodyweight
during 0-64 days starvation,
Spring
1972.

by doe s and fa wns

�-495body weight.

Diet IV (high fiber, low protein) was refed.

The doe ate 510 9

the first day, 216 9 the second day, 590 9 the third day and 335 9 the fourth
day.

Her intake did not exhibit the sharp rise exhibited by other refed

deer after the second day of refeeding.
survived.

All nine starved and refed fawns

Difference in refeeding mortality rates between does (8.3 percent)

and fawns (0.0 percent) was not significant (P &gt; 0.1).
Effect of Length of Starvation Upon Refeeding Success
Four fawns and four does starved less than 20 days survived refeeding.
Of 5 fawns and 8 does starved over 20 days only one (a doe) died during
refeeding.

Difference of refeeding mortality between deer starved less

than, vs greater than, 20 days was not statistically significant (P &gt; 0.1).
Of 30 de~r s~arved and refed in the pilot study and in the spring 1971
trial only 2 deer died during refeeding.

These results indicate that: 1)

refeeding success is unrelated to length of starvation itself; and 2) little
mortality occurs due to refeeding.
Causes of Refeeding Mortality
Tests of hypotheses concerning refeeding mortality were negated by
failure to induce significant refeeding mortality.

However a brief

presentation of pertinent results is merited.
Urea toxicity.

Mean PUN levels on the tenth day of refeeding for deer

refed Diet III (high protein, low fiber) (Y = 17.6 mg percent for does; 13.9
mg percent for fawns) were significantly higher (P&lt; 0.1) than those for deer
refed Diet IV (high fiber, low protein) (Y = 10.9 mg percent for does; 13.1
percent for fawns).

However, these levels were not significantly different

(P ~ 0.1) from those of non-starved deer.

PUN for the doe dying on the fifth

day of refeeding was 26 mg percent. a value considered not toxic.

�-496Mean PUN level on the tenth day of refeeding for does refed Diet III
was identical to that for non-starved does.

Mean PUN levels on the tenth

day of refeeding for does refed Diet IV and for fawns refed both diets were
Significantly lower (P ~ 0.1) than those fur non-starved deer.
Liver malfunction. Plasma OCT and GOT levels for starved and refed deer
were well below levels indicative of liver damage (Table 30).

Plasma

OCT and GOT levls for the doe dying 4 days after refeeding began were
well below levels indicative of liver damage.
Effects of Feed Refed Upon Refeeding Performance
Weight gain.

Percent pre-refeeding weight gain on the tenth day of refeeding

was compared between deer refed Diet III and deer refed Diet IV. Separate
comparisons of weight gain related

to refeeding diet were made for fawns

and does to avoid age-class bias.

In six of seven such comparisons, deer

refed Diet III exhibited non-significantly higher (P &gt; 0.1) percent prerefeeding weight gain than deer refed Diet IV.
Intake during feeding.

Deer co~sumed a daily average of 134.1 g, or 8.7

percent, less of Diet IV than of Diet III during refeeding.

This difference

was highly significant (P ~ 0.001).

Regardless of diet refed, doe and fawn

intakes exhibited large variation.

First day intake ranged from 1488 g

(doe refed Diet III) to 27 g (doe refed Diet III).

For some deer, subsequent

intakes never matched that of first day; for others intake during refeeding
remained at first day's level.

The intake pattern displayed by deer in the

pilot study (intake on the first day was duubled by the third day and tripled
by the seventh day) was not observed in 1972.
Spring 1973 Trial
Conditioning Period
Experimental deer experiencing negative energy balance during conditioning would mobilize stored fat and muscle protein for conversion in the liver
to such primary energy sources as glucose and ketone bodies.

This phenomenon

�-497-

Table 30.

Deer plasma enzyme levels. spring 1972 trial.

Deer
Control
5C3

T5

25.0

5C

T

T

:iC

T

1.0

6C4

T

0.5

6C

T

12.5

6C

T

2.5

Fawn, 17 days

T

9.0

Doe. 10 days

T

Doe. 64 days

T

6.5

Fawn, 10 days

T

T

Fawn, 10 days

T

15.5

Fawn, 10 days

T

18.5

day

T

7.5

Doe, 3 days

T

1.0

Doe, 4 days6

T

5.0

Doe, 10 days

T

7.5

Starved

Refed

Doe,

10rnithine carbamyl transferase, Sigma unit levels indicative of
liver damage ~ 1000 Sigma units/ml.
2Glutamine oxalacetic transaminase, Sigma unit levels indicative
of liver damage ~ 28-50 Sigma units/ml.
3 4

, Non-starved control deer sampled throughout trial.

5T = Trace.
6Deer died on fourth day of refeeding.

�-498-

would be indicated by losses of body weight and increased blood levels of
free fatty acids and urea.
Weight change.

Experimental deer lost an average of 12.0 percent body

weight during conditioning, whereas control deer gained an average of 3.7
percent (Table 13). Weight change difference between control and experimental
deer was highly significant (P ~ 0.01).
Plasma energy-metabolite levels.

Mean PUN level for experimental deer

at the third week of conditioning was significantly lower (P ~ 0.05) than
when conditioning began (Table 31).

This was contrary to expectations pre-

dicting an increase in PUN level as a reflection of catabolism of muscle
protein.

At the end of conditioning, mean PUN level for experimental deer had

risen significantly (P ~ 0.1) above third week level but remained significantly
(P ~ 0.05) below pre-conditioning levels.

PUN level for control deer declined

throughout conditioning; values at the end of conditioning were significantly
(P ~ 0.05) lower than at pre-conditioning and at the third week of
conditioning.
Inexplicably, mean PUN levels prior to conditioning were significantly
different (P ~ 0.05) between control and experimental deer.

By the end of

conditioning, mean PUN level for experimental deer was significantly higher
(P :5_0.05) than that for control deer.
During conditioning mean level of plasma FFA for experimental deer
increased steadily and significantly (P ~ 0.05), whereas that for control
deer did not change (Table 32). At the end of conditioning, mean level of
plasma FFA for experimental deer was significantly higher (P ~ 0.1) than
that for control deer.
Mean level of plasma glucose for control and experimental deer exhibited
large, non-significant (P &gt; 0.1) variations during conditioning (Table 33).
The significant (P &lt; 0.1) difference between control and experimental deer

�-499Table 31. Means and standard deviations of plasma urea nitrogen
concentrationsa, spring 1973 trial.
Control Deer
SD
n
-

X

Ex~erimental Deer
n
SD

Trial Phase

X

Begin
b
Cond ition ing

21.4h,i

,3.6

4

25.1h,i,k

3.6

8

3d Week of c
Conditioning

20.4k

3.7

5

18.4i,j

1.6

8

Begin d
Starvation

16.4h,i,k

3.0

5

20.5h,k,j

4.3

7

Begin
Refeeding e

16.7

8.3

5

22.6

10.9

6

Ten Days
Post-feeding f

17.1

9.9

4

17.2

5.9

5

aMg urea nitrogen per 100ml plasma.
bBeginning of conditioning period, control deer fed ad libitum,
experimental deer' fed one-quarter of control deer intake.
cTwenty-third day of conditioning.
dForty-fourth day of conditioning, beginning of food deprivation,
all deer.
eTwenty-third day of food deprivation, beginning of refeeding,
all deer.
fTenth day of refeeding.
hDifferences between means in rows significant at P ~ 0.05.
i,kOifferences between means in columns significant at P &lt; 0.05.
jOifferer,ces between means in columns significant at P &lt; 0.1.

�-500Table 32. MeaRs and standard deviations of plasma free fatty acid
concentrations, spring 1973 trial.

Control Deer
Trial Phase
Begin
b
Conditioning
3d Week of c
Conditioning
Begin
Starvationd
Begin e
Refeeding
Ten Days
Post-feeding f

X

SD

0.29i

Experimental Deer

-

n

X

SD

n
-

0.08

4

0.25i,j,k

0.03

8

0.26k

0.05

5

0.33i,n

0.10

8

0.30h,j

0.13

5

0.43h,j,n,1

0.20

7

0.7oiij,k 0.28

5

0.73k,1,m

0.20

6

0.271

4

0.20m

0.06

5

0.15

aMicro-equiva1ents of free fatty acids per m1 plasma.
b-fSee Table 8.
hDifferences between means in rows significant at P ~ 0.1
i-mDifferences between means in columns significant at P ~ 0.05.
nDifferences between means in columns significant at P &lt; 0.1.

�-501-

TABLE 33. Neans and standard deviations of plasma glucose concentrationsa,
spring 1973 trial.

Control
Trial

Phase

Deer

X

Experimental

Deer

SD

n

X

SD

n

28.4

5

131. sj

20.0

8·

Begin
b
Cond it ion irig

114. 3

d
3 Week of
c
Conditioning

129.1

17.5

5

112. 8

19.9

.8

Begin
d
Starvation

101. S

38.9

5

119.3

54.9

8

Begin
'e
Refeeding

132.8

46.2

5

108.9

26.4

6

Ten Days f
P'os t fe e dtng

138.5

40.9

4

113.4

33.9

5

significant

g

g

j

e

a

Mg glucose

b-f

per

lOOml plasma.

.
See Table 8.

gDifferences

between means

in rows

jDifferences

between means

in columns

at P:::. O. 1.

significant

at P~O. 05.

�-502-

Table
trial.

34. Hematocrit

means and standard

deviations,

Control Deer
Trial

Phase

spring

Experimental

X

SD

n

X

52.6

2.9

5

54.6

Begin
d
Starvation

51. 0

3.7

5

52.3 i

Begin
e
Refeeding

50.2

3.0

5

50.8

0.7

2

43.8

1973

Deer
SD

n

2.5

7

2.5

7

5. 1

6

4.9

5

Begin
b
Conditioning
d
3 Wee'k of
c
Conditioning

Ten Days
Post-feeding

s-r
i

f

i

45.5

i

i

j

j

See Table 8.

Differences

jDifferences

'

between means

in columns

significant

at P.sO. 1.

between means

in columns

significant

'at P=:;.O.
05.

�-503mean level of plasma glucose at the third week of conditioning was judged
not biologically significant.
Mean hematocrit did not differ significantly (P ~ 0.1) between control
and experimental deer during conditioning (Table 34).

Decl"ine in experimental

deer hematocrit from the third to the sixth week of conditioning was
significant (P &lt; 0.1).
Mean level of plasma total protein did not differ significantly (P &gt; 0.1)
between control and experimental deer at the end of conditioning (Table 35).
The significant difference (P &lt; 0.1) between control and experimental deer
mean level of plasma total protein at the third week of conditioning is
inexplicable and judged not biologically significant.
Pre-starvation Nutritional Plane and Starvation Performance
Mortality.

Of eight experimental deer starved, three died; one at

11, one at 12, and one at 23 days of starvation.

Difference between

mortality of experimental deer (37.8 percent) and control deer (none) was
significant (P 2 0.1).
Weight loss.

Percent weight loss during starvation for control deer

was not significantly different (P &gt; '0.1) from that for experimental deer
(Table 3~.

Percent weight loss during entire trial for experimental deer

was significantly higher (P &lt; 0.01) than that for control deer (Table 36).
Percent weight losses exhibited during conditioning and starvation
between experimental deer surviving and experimental deer succumbing to
starvation were not significantly different (P &gt; 0.1) (Table 36).
Experimental deer lost weight at a significantly higher rate (P 20.01)
during starvation than during conditioning (Table 36).
Levels of plasma energy metabolites.

A blood sample was collected,

immediately prior to death, from the experimental deer dying on the 12th
day of starvation.

Similar blood samples were not obtained from the other

two experimental deer dying during starvation.

�-504-

Ta.ble 35. Means and standard deviations of total protein concentrationa,
spring 1973 trial.

Control Deer

Experimenta 1 Deer

Trial Phase

X

SO

n

X

SO

n

Begin
Conditioningb

7.6

0.5

4

8.01

0.5

3d Week of
ConditioningC

8

7.6g

0.4

5

8.0g,k

0.5

7

Begin
d
Starvation

7.6

0.6

5

7.7j

0.6

7

7.5

0.5

5

7.3j,k.l

0.4

6

8.0h

0.4

2

7.0h

0.4

5

Begin
Refeedinge
Ten Oaysf
Post-feeding

aG total protein per 100ml whole blood.
b-fSee Table 8.
gOifferences between means in rows significant at P ~ 0.1.
hOiffercnces between means in rows significant at P ~ 0.05.
jOifferences between means in columns significant at P ~ 0.01.
k,lOifferences between means in columns significant at P &lt; 0.05.

�Table 36.

Body weight changes, spring 1973 trial.

Percent Weight
Loss During
Starvation
SO
X

Percent Weight
Loss,
Entire Trial
SO
X

Percent Wei ght
Gain During
Refeeding
SO
X

-14.Sh

5.6

-13.4j

7.5

+5.8

6.5

5..2

+6.7

3.7

Deer

n

. Control

5

Percent Weight
Loss During
Conditioning
SO
X
+3.7h,j
3.0

Experimental
deer surviving
starvation

5

-11.99,j

5.1

-16.49

3.3

-25.9j .

Experimenta 1
dying durin9
starvation

3

-12.29,j

1.4

-15.8g

2.7

-26.7

90ifferences

between means in rows significant

(P ~ 0.1).

hOifferences between means in columns significant

(P ~ 0.05).

jDifferences

(P ~ 0.01).

between means in columns significant

j

2.5

0,
0
U1
I

�-506Mean PUN level for experimental deer surviving starvation was not
significantly different (P &gt; 0.1) from that for control deer.

PUN

level for the experimental deer dying during starvation (40.0 mg percent)
was outside the 95 percent confidence intervals of mean PUN levels for
control deer and experimental deer surviving starvation and judged
significantly different (P-&lt; 0.05) from both.
Mean levels of plasma FFA for control deer and experimental deer
surviving starvation were not significantly different (P &gt; 0.1)

Plasma FFA

level of the experimental deer dying during starvation (0.22 micro-equivalents/ml) was outside 95 percent confidence intervals of mean levels of
plasma FFA for control deer and experimental deer surviving starvation and
judged significantly different (P &lt; 0.05) from both.
Mean levels of plasma glucose for control deer and experimental deer
surviving starvation were not significantly different (P &gt; 0.1)

Plasma

glucose level for the experimental deer dying during starvation (24 mg
percent) was outside 95 percent confidence intervals of mean levels of
plasma glucose for control deer and experimental deer surviving starvation
and judged significantly different (~20.05) from both.

Mean glucose

level for experimental deer surviving starvation was significantly lower
(P 2 0.05) than at beginning of conditioning.
Mean hematocrit for control deer was not significantly different
(P &gt; 0.1) from that for experimental deer surviving starvation.

Hematocrit

for the deer dying during starvation (48) was within 90 percent confidence
intervals of means for other deer and judged not significantly different
(P &gt; 0.1) from them.
Mean levels of plasma total protein for control deer and experimental
deer surviving starvation were not significantly different (P &gt; 0.1).

Leve1

�-507of plasma total protein for the experimental deer dying during starvation
(6.1 g percent) was outside 95 percent confidence intervals of mean levels
of plasma total protein for other deer and judged significantly different
(P 2 0.05) from them.
Pre-starvation Nutritional Plane and Refeeding Performance
Mortality.

Neither control or experimental deer experienced refeeding

mortality.
Weight gain.

Percent weight gain during refeeding for control deer was not

significantly different (P &gt; 0.1) from that for experimental deer
(Table 36).
Levels of plasma energy-metalolites.

Mean PUN levels on the tenth day of

refeeding for experimental deer and control deer were not significantly
different (P &gt; 0.1) from respective levels at beginning of refeeding.
Mean levels of plasma FFA on the tenth day of refeeding for experimental
deer and control deer were not significantly different (P &gt; 0.1).

These

levels were significantly lower (P 2 0.05) than respective levels at beginning
of refeeding.
Mean levels of plasma glucose for control deer and experimental deer
were not significantly different (P &gt; 0.1) from each other on the tenth
day of refeeding nor from respective values at beginning of refeeding.
Mean hematocrits on the tenth day of refeeding for control deer and
experimental deer were not significantly different (P &gt; 0.1).
these values were significantly lower (P 20.1

However,

within control deer; P 20.05

within experimental deer) than respective values at beginning of refeeding.
Mean level of plasma total protein on the tenth day of refeeding for
control deer was significantly higher (P 2 0.05) than that for experimental
deer.

These values were not significantly different (P &gt; 0.1) from respective

values at beginning of refeeding.

�-508Intake During Refeeding
Experimental deer ate significantly more (P &lt; 0.005) during refeeding
than controls.

Intakes of experimental and control deer approximately

doubled by the third day of refeeding.

Experimental deer tripled initial

intake during 6-8 days of refeeding; control deer intake appeared to
increase at a slower rate and never tripled that of the first day.
Blood Energy-Metabolites From Free-Ranging Deer
Levels of blood energy metabolites for free-ranging deer are presented
in Table 37 and discussed in relation to levels for penned deer in 1972
and 1973 spring trials under appropriate discussion sections.
Blood Energy-Metabolites From Spring 1972 Trial Deer
Starvation
PUN.

Mean PUN levels for does and fawns differed significantly (P &lt; 0.025)

only at 33-36 days of starvation (Table 38).

Three of six fawns starved

33-36 days died; mean PUN for these deer (42.3 mg percent) was significantly
higher (P ~ 0.05) than that for surviving fawns (26.5 mg percent).

Mean

PUN level for fawns starved 33-36 days was significantly higher (P &lt; 0.001)
than that for non-starved fawns.
PUN level for the doe dying during refeeding was within the 90 percent
confidence interval of mean PUN level for two does starved 55-64 days and
judged not significantly different (P &gt; 0.1) from them.
FFA. Mean levels of plasma FFA for non-starved does and fawns were not
significantly different (P &gt; 0.1).

During starvation, mean levels of

plasma FFA for does were significantly higher (P ~ 0.05) than those for
fawns (Table 39). Mean levels of plasma FFA for fawns starved 33-36 days
was significantly higher (P ~_0.1) than that for non-starved fawns.

Mean

FFA levels for remaining does and fawns starved 10-64 days were significantly
higher (P ~ 0.05) than those for non-starved deer of respective age
classes.

�Table 37. Comparisons of mean values of physiological parameters among adult does during winterearly spring.
Michigan W_T1 Does
19682

19723

New York4
W-T Doe

1972

1973

Free
Rangi ng
Does

126.5

125.8

106.8

97-123

100-168

72-111

17.7

23.9

8.2

25.6

11-22

15.7 -28.4

Plasma FFA7

0.24

0.27

0.28

Hematocrit

52.1

--

--

51-54

58.7

50-51.5

--

Spri_ni)Trial Does
-

Plasma 5
Glucose
Blood6
Urea
Nitrogen

Tota ,8
Protein

--

7.6

--

6.7-7.3

5.7-5.9

Body weight (kg)

63.9

59.4

61. 2

50.6-56.3

55.0

lW_T = white-tailed deer.

7Micro-equ ival ents/ml .

2Data from Ullrey et al. 1968.

8G percent.

30ata from Seal et al. 1972.
40ata from Maynard et al. 1935.
5

Mg percent.

6Mg percent.

I
&lt;..T1

0
\0
I

�-510-

Table 38. Means and standard deviations of plasma urea nitrogen concentrationsa
from deer starved 0-64 days, spring 1972 trial.

Days
Starved

Does

X

SD

17 .7

. Fawns

X

SD

n

3.0

n
13

17 .2

3.0

13

22.3

6.7

2

17 .0

0.0

2

15.0

1.4

"2

20.5

4.9

2

24

17.5

1.4

2

20.5

0.7

2

33-36

18.0b

5.2

3

34.4b

11.0

6

20.5

2.8

3

0
10
17

55-64

aMg urea nitrogen per 100ml plasma.
bDifferences between means significant at P ~ 0.025.

�-511-

Table 39 .. Means and" standard deviationsbof plasma free fatty acid concentrat tons''
from deer starved 0-64 days, spring 1972 trial.

Fawns

Does
Days
Starved

X

SO

n

X

SD

n

0.24c

0.04

13

0.26c,d

0.04

13

0.24

2

2

0.54 "

2

0.34c
0.39c

0.08

17

0.53c
1.32c

0.01

2

24

0.65c

0.16

2

0.37c

0.16

2

0.6lc
c
0.69

0.42

3

0.29d

0.06

6

0.61

3

0
10

33-36
55-64

aMicro-equivalents of free fatty acids per m1 plasma.
bAccumu1ated comparisons of doe and fawn mean values during starvation
significant at P 2 0.05.
cMeans of non-starved deer significantly lower (P &lt; 0.025) than those
of all starved deer.
dDifferences between means significant at P &lt; 0.1.

�-512Plasma FFA level for the doe dying during refeeding was outside
the 95 percent confidence interval of mean plasma FFA for two does at
55-64 days of starvation and judged significantly different (p &lt; 0.05) from
them.
Glucose.

Mean level of plasma glucose for does starved 55-64 days

was significantly lower (P ~ 0.05) than that for non-starved does.

Mean

levels of plasma glucose for starved fawns and other starved does were
not significantly different (P &gt; 0.1) from each other nor from those
of respective age-class non-starved deer (Table 40).

Mean levels of plasma

glucose for three fawns starved 33-36 days and dying (32.7 mg percent)
was significantly lower (P ~ 0.025) than that for fawns surviving 33-36
days of starvation (113.8 mg percent).
Plasma glucose level for the doe dying during refeeding was outside
the 95 percent confidence interval of mean level of plasma glucose for
two does starved 55-64 days and judged significantly different (P &lt; 0.05)
from them.
Hematocrit.

Mean hematocrits for fawns and does were not signifi-

cantly different (P &gt; 0.1) (Table 41).

Mean hematocrits for does and fawns

starved ten days were significantly higher (P ~ 0.1) than those for respective age-class non starved deer.

Mean hematocrit for does starved

55-64 days was significantly lower (P ~. 0.01) than that for non-starved
does.
Hematocrit for the doe dying during refeeding was within the 90 percent
confidence interval of mean hematocrit for two does at 55-64 days
of starvation and judged not significantly different (P &gt; 0.1) from them.

�-513-

Table 40. Means and standard deviations of plasma glucose concentrationsa
from deer starved 0-64 days, spring 1972 trial.

Fawns
SO

n

145.1

35.5

13

2

183.3

22.9

2

. 30.1

2

106.0

22.9

2

118.8

36A

2

145.5

2.8

2

33-36

101.5

28.2

3

73.3

49.9

6

55-64

89.7b

41.9

3

v

Does
SO

n

X

0

126.5b

28.9

13

10

162.5

48.8

17

97.8

24

Deer
Starved

1\

a Mg glucose per 100 ml plasma.
bOifferences between means significant at P .:.0.01.

�-514-

Table 41. Hematocrit means and standard deviations from deer starved
0-64 days, spring 1972 trial.

0

52.1a,b,c

Does
SO
2.8

n
13

10

56.5a

0.7

17

55.0

24
33-36

54.0
53.5b

55-64

46.3c

Days
Starved

a,bOifferences

X

50.3a

Fawns
SO
4.2

13

2

56.0a

1.4

2

1.4

2

53.5

5.0

2

4.2

2

53.5

3.5

2

0.7

2

50.3

6.1

6

3.1

3

X

between means significant at P ~ 0.1.

CDifferences between means signifi~ant at P &lt; 0.01.

n

�-515Refeeding
Effect of diet refed on level of blood energy-metabo"'ites. Levels of
blood energy-metabolites on the tenth day of refeeding were compared
respectively between:

1) fawns; and 2) does refed Diet III and Diet IV

for each starvation period.

There were no significant differences (P &gt; 0.1)

in levels of blood energy-metabolites between deer refed Diet III and deer
refed Diet IV.
Effect of length of starvation on level of blood energy-metabolites for
refed deer.

Assuming levels of blood energy-metabo1ies were not affected

by diet refed, values of each blood parameter on the tenth day of refeeding
were regressed on length of starvation prior to refeeding.

This was done

to determine whether or not length of starvation prior to refeeding affected
levels of blood energy-metabolites on the tenth day of refeeding.

Hematocrit

of does on the tenth day of refeeding declined significantly (P ~ 0.05) as
length of starvation prior to refeeding increased (Table 42).

Ninety percent

confidence intervals of slopes of other blood parameters from fawns and does
included zero, indicating that length of starvation prior to refeeding had no
influence on levels of blood parameters on the tenth day of refeeding
(excepting doe hematocrit).
Comparison of levels of blood energy-metabolites between the tenth day of
refeeding and at beginning of starvation.

Fawn mean PUN level on the tenth

day of refeeding was not significantly different (P &gt; 0.1) from prestarvation level.
All levels of blood energy-metabolites on the tenth day of refeeding for
does were significantly lower than pre-starvation levels (P &lt; 0.1 PUN;
P &lt; 0.01 plasma glucose; P ~ 0.05, plasma FFA).

�Table 42. Slopes and associated 90 percent confidence intervals for regression of mean level of blood
parameters (at the tenth day of refeeding) on length of starvation prior to refeeding.

Slopec
Confidence
1evel

PUNa

FFAb

Does
Glucose

0.05

0.00

-0.12

Ht!matocrit

PUNa

-0.19d

0.00

FFAb

Fawns
Glucose

Hematocrit

0.00

0.61

-0.24

I
01

--'

+ 0.21

+0.00

+0.75

+0.15

apUN = plasma urea nitrogen.
bFFA = free fatty acid.
CChange in mean blood parameter value per day of starvation.
dSlopesignificantly

different from zero (P ~ 0.05).

+0.42

+0.01

0'1
I

+2.51

+0.32

�-517Levels of plasma energy-metabolites

for the doe dying during refeeding.

A blood sample was obtained from this doe immediately prior to death,
on the fourth day of refeeding.
was 0.2 micro-equivalentsjml;
hematocrit was 43.

PUN was 26 mg percent; plasma FFA

plasma glucose was 36 mg percent and

��-519Chapter 4
Discussion - Deer Nutritional Physiology
Metabolic Disfunction
Toxicity Occurring During Refeeding
Urea toxicity.

None of the diets for the two starving deer in this study

resulted in observable urea toxicity.

PUN for the 1972 trial doe dying

during refeeding (26 mg percent) was only slightly higher than before it
was refed (23.5 mg percent).

Neither this deer nor the doe dying during

refeeding in the pilot study exhibited typical urea-ammonia toxicity
symptoms (dyspnea, incoordination, tetany and tetanic spasms, slobbering
and severe muscular twitches) (Church 1969:298).
Acute indigestion.

Dahlberg and Guettinger (1956) speculated that

starving deer may have died after refeeding as a result of overeating.
Overeating toxicity is caused, primarily, by ingesting "...
normal amounts of readily fermentable carbohydrates..

larger than

resulting in

abnormal and toxic fermentation products ... " (Church 1969:293).
Clinical symptoms include loss of appetite, dehydration (evidenced by
increased hematocrit), high blood glucose, diarrhea and ataxia (motor
incoordination) (Church 1969:294).
None of the diets fed to starving deer in this study resulted in
acute indigestion.

During spring 1971 and 1973 trials, deer voluntarily

restricted food intake the first 2 days of refeeding following starvation.
The one doe dying during refeeding in the 1971 spring pilot study
similarly restricted its intake the first two days of refeeding.

This

deer did not exhibit ataxia, diarrhea, loss of appetite, high blood
glucose or dehydration (based on hematocrit, which declined rather than
increased) and was judged not to have died from acute indigestion.

�-520Mortality.

Of 4 starved deer refed, only 2 died during refeeding.

Other starving deer presumably were in condition comparable to that of
the deer dying during refeeding.

Development of toxicity upon refeeding

of 40 deer should have included more than two deer if it were a genera11yoccurring phenomenon.
Refeeding starved deer Diets II, III and IV did not result in
observable toxicity nor in mortality attributable to toxicity nor in
appreciable mortality.
Liver Malfunction
Blood levels of OCT and GOT were well below those considered indicative
of liver damage/malfunction.

These results, coupled with low refeeding

mortality, suggest that liver malfunction, occurring during refeeding, is
an important mortality agent of starved and refed deer.
Nutrition
Qualitative Aspect - Feeds Fed to Starving Deer
Starving deer refed Diet IV exhibited blood parameter values similar
to those of starved deer refed Diet III, indicating that deer were able to
produce metabolites required for sustenance from both diets.

The non-

significantly higher weight gain exhibited by deer refed Diet III is probably
a reflection of the significantly higher consumption of Diet III during
refeeding.

However, deer refed Diets II and IV exhibited 10.5 percent

refeeding mortality (2 deer died of 19 refed), whereas deer refed Diet III
exhibited no mortality.

This difference was significant (P ~ 0.1).

of death in these two deer is not understood.

Cause

Blood samples were collected

from the deer dying during the 1972 spring trial but not from the deer
dying during the 1971 spring trial.

Of all comparisons of blood parameters

between this deer dying during refeeding and deer surviving refeeding, only

�-521-

plasma glucose levels were different.

Plasma glucose for the-deer dying

during refeeding (36 mg percent) was less than half that for deer surviving
refeeding but similar to that for deer (fawns and dow) dying during
starvation.

Apparently, gluconeogenesis was impaired in the deer dying

during refeeding and may have been the cause of death.

However, blood

levels of liver enzymes from this deer were not indicative of liver
malfunction.
Quantitative Aspect - Energy Intake of Starving and Refed Deer
Spring 1973 trial.

Deer of the two starving nutritional strata were produced

during conditioning.

Control deer lost no weight and exhibited no elevations

of PUN or plasma FFA.

Experimental deer exhibited significant weight losses

and plasma FFA-elevations during conditioning, indicating that fat stores
were mobilized to offset inadequate exogenous energy intake.
deer PUN did not increase during conditioning.

Experimental

This result may be inter-

preted as indicating that catabolism of muscle protein was not occurring in
experimental deer during conditioning.

However, Young and Scrimshaw (1971)

stated that catabolism of muscle protein occurs throughout periods of
reduced food intake.

It is more likely that catabolism of muscle protein

was occurring duri~g conditioning and that the kidneys were clearing urea
from the blood at a rate sufficient to prevent blood urea level from rising.
As expected, the first three deer to die during starvation were
experimental deer.

Weight losses and levels of blood energy-metabolites

for these deer were not significantly different from those for other
~xperimental deer at beginning of starvation.

Weight losses of these deer

during starvation were similar to those of experimental deer surviving
starvation.

However, levels of blood energy-metabolites obtained from one

deer dying during starvation strongly suggest that it succumbed to inadequate

�-522energy metabolism.

PUN level was significantly higher and plasma FFA,

glucose and total protein levels were significantly lower, than those for
surviving experimental deer and control deer.

Apparently this deer depleted

its fat reserves (indicated by high PUN) to provide an endogenous supply of
energy metabolites.

Catabolism of this muscle protein did not provide the

degree of gluconeogenesis required to maintain energy balance (indicated by
low levels of plasma glucose and plasma total protein) and the deer died.
Extremely low plasma glucose level (24 mg percent) of the deer dying during
starvation suggests that hypoglycemia was the primary mortality agent.
Surviving starved deer had levels of plasma glucose greater than 73 mg
percent.
Experimental deer lost weight at a significantly higher rate during
starvation than during conditioning.

This was expected, for deer were

obtaining part of their energy from feed during conditioning and catabolizing
fat at a slower rate (mean plasma FFA level of experimental deer during
starvation was significantly higher than during conditioning than during
starvation.

Mean weight loss of deer dying during starvation (27 percent)

was similar to the 30 percent figure considered fatal (Davenport 1939,
Doman and Rasmussen 1944).
Control deer and experimental deer surviving starvation exhibited
similar levels of blood parameters (excepting total plasma protein which
is inexplicable), weight changes and lack of mortality at 24 days of
starvation and at 10 days of refeeding.

Mean levels of plasma glucose,

FFA, PUN and total proteins on the tenth day of refeeding for both groups
of deer were similar to those at beginning of conditioning.

Mean hemat-

ocrits, however, were significantly lower than during conditioning, starvation
and those of non-starved deer in the spring 1972 trial, indicating that
an inexplicable hyper-hydration occurred in spring 1973 trial deer.

�-523These results indicate that:

1) deer starved to the point of death

can be refed successfully (assuming experimental deer surviving starvation
were physiologically equivalent to experimental deer dying during starvation);
2) experimental deer surviving starvation were catabolizing stored fat during
starvation and metabolically equivalent to control deer during starvation
and refeeding; and 3) deer dying during starvation failed to maintain levels
of blood energy-metabolites required for sustenance.
Spring 1972 trial. As in the 1973 trial, starving does lost weight and
exhibited increasing plasma FFA levels and constant PUN levels.

Plasma

glucose levels remained constant until 55-64 days of starvation when they
declined to values significantly lower than before starvation.

Weight

loss was linearly related to length of starvation, whereas plasma FFA
levels appeared to increase rapidly, peak at 17 days of starvation and
drop to a constant level as length of starvation increased.

Fawns also

lost weight linearly but at a significantly higher rate than does.

Fawn

plasma FFA levels were significantly lower than those of does during
starvation.

PUN levels for starved fawns remained at pre-starvation level

until 33-36 days of starvation whereupon they increased sharply and were
significantly higher than those for does starved the same length of time.
Plasma glucose levels of starved fawns likewise remained at pre-starvation
levels until 33-36 days of starvation whereupon they dropped significantly.
Differences in levels of blood energy-metabolites between fawns
dying and fawns surviving 33-36 days of starvation were of particular
interest.

Dying fawns exhibited significantly higher PUN, lower plasma

glucose and non-significantly lower FFA and higher weight loss than surviving
fawsn.

Mean weight loss of fawns dying during starvation was 30 percent,

identical to that considered fatal to deer in winter (Davenport 1939,
Doman and Rasmussen 1944).

Mean level of plasma glucose for starved fawns

dying (32.6 mg percent) was similar to those for the doe dying of starvation

�-524during the spring 1973 trial (24 mg percent) and the doe dying during
refeeding in the spring 1972 trial (36 mg percent).
These results suggest that fawns and does catabolized stored fat during
starvation (indicated by weight losses and increased plasma FFA).

Fawns

increased the rate of catabolism of muscle protein when plasma FFA levels
dropped, indicating that stored fat reserves were depleted.

Does and

fawns surviving starvation maintained plasma glucose at or above levels
required to sustain life.

Dying fawns apparently exhausted fat reserves

(indicated by low plasma FFA) and subsequent increased protein catabolism
(indicated by high PUN levels) was not sufficient to maintain plasma glucose
at levels required to sustain life.
Higher rate of weight loss during starvation for fawns suggests that
they catabolized stored fat and muscle protein at higher rates than does.
Lower mean level of plasma FFA for fawns during starvation indicates that
they extracted FFA from blood at a higher rate or mobilized FFA at a slower
rate than does.

Given the more rapid depletion of fat stores by fawns,

it is reasonable to expect that fawns would succumb to starvation before
does, as they did.
Levels of blood energy-metabolites for fawns on the tenth day of
refeeding were similar to non-starved levels, indicating that fawns, as
did does in the 1973 spring trial, normalized their metabolism by the
tenth day of refeeding.

Mean levels of PUN, plasma glucose and hematocrit

for does on the tenth day of refeeding were significantly lower that prestarvation levels.

Depressed doe mean hematocrit (48.1) was not as low

as that of spring 1973 trial does at 10 days of refeeding (44.5), indicating
a lessened degree of hydration.

Lower mean levels of PUN and plasma

glucose for does are inexplicable and may not be biologically significant
as no refeeding mortality occurred (excepting the doe, discussed above,
dying during refeeding with high PUN, low plasma FFA and extremely low

�-525(36 mg percent) plasma glucose.
These results indicate that: 1) starving deer can be refed successfully;
2) does and fawns are not metabolically equivalent during starvation and
refeeding and 3) deer dying during starvation succumbed to hypoglycemia.
Non-starved deer.

Levels of blood energy-metabolites and body weights for

does from spring 1972 and 1973 trials were similar between years, to those
for does in other winter-early spring studies and to those for free-ranging
wild does sampled in spring 1972 (Table 37).

The only apparent disagreement

among these values is the lower PUN exhibited by free-ranging does.

As a

result of this general agreement, physiological parameters for does during
spring 1972 and 1973 trials were regarded as being representative of does
in good nutritional condition in winter-early spring.

No such comparisons

could be made for fawns; values of 1972 spring trial fawns were assumed to be
representative of fawns in good nutritional condition in winter.
Starved/malnourished deer.

Levels of blood energy-metabolites and body weight

losses for does from spring 1972 and 1973 trials were similar between years
and to those of does in other winter-early spring studies (Table 43).

Data

of Teeri et al. (1958), indicating that fawns malnourished 2-3 months during
winter and losing 19-23 percent body weight were hypoglycemic (43-60 mg percent
blood glucose) within 2 days of death agree with findings in this study.

Three

of four mule deer fawns subjected to a "poor-quality" diet for 2 weeks during
winter exhibited weight loss and elevated mean PUN (from 21.6 mg percent
pre-trial to 42.5 mg percent post-trial) (Schoonveld 1971).

These data

indicate that weight losses and levels of blood energy metabolites for deer
from spring 1972 and 1973 trials were valid and representative of starving
and malnourished deer.

�-526-

Table 43. Comparison of mean values of physiological parameters among adult
does during winter-early spring.

Glucose

118.8

Sl2ring Trial Does
1973
Michigan W-T4 Does
Contro 1&lt;Exl2erimenta13 19689
197210
132.8
112.8-119.3
34-102
100-140

Blood urea5
nitrogen

17 .5

16.6

FFA

0.65

0.70

Hematocrit

54.0

19721

')

18.4-20.5

6.1-16.2

50.2

50.8-52.3

55.0-57.0 43.5-51.0

7.5

7.7-8.0

7.0-7.6

5.3-5.4

14.8

11.9-16.4

8.6-20.6

24.0

16.0

Pla~ma

Tota 1 7
Plasma
protein
Weig§t
Loss

11.0

lDeer starved 24 days.
2Deer starved 23 days.
3Deer malnourished (intake restricted to 1/4 normal); first number =
deer malnourished 3 weeks, second number = deer malnourished 6 weeks.
4W_T = white-tailed deer.
5Mg percent.
6Micro-eQuivalents/ml.
7G percent.
8percent body weight loss.
9Data from Ullrey et al. 1968.
100ata from Seal et al. 1972

�-527Effect of Length of Starvation on Refeeding Success
The hypothesis that deer cannot be successfully refed after 20 days
of starvation, empirically based on results of the spring 1971 pilot
study, was refuted by results of spring 1972 and 1973 trials, wherein 23
of 24 deer starved longer than 20 days were successfully refed.
Management Implications
Results of three years' refeeding trials, indicating that starving
deer can be refed successfully, contradict field findings.

Three factors

are considered responsible for this discrepancy.
Type of feed.

Although starved deer performed similarly when refed different

diets in this study and exhibited negligible mortality, they were not challenged
by diets containing alfalfa hay as were deer dying during refeeding in the field.
In Part I of this final report we indicated that mule deer fed ground or cubed
"poor quality" alfalfa develop fatal digestive upsets, omasa1 impactions and
hemorrhagic ulcerations of the abomasum.

It is possible that structural

characteristics of lignin in alfalfa may have been responsible for the inability
of deer to assimilate alfalfa hay.
Energy.

Literature concerning refeeding of deer in the field indicated that

starving deer did not receive enough food to satisfy daily energy requirements.
Preponderance of fawns among deer dying during field refeeding and findings of
this study wherein fawns succumbed to starvation before adults suggests that
deer refed in the field were dying from lack of adequate energy intake.
Environment.

Deer in this study were not subjected to the rigorous climatological

conditions typical of winter range.

Stresses imposed by colder temperatures and

exertion of moving through deep snow may increase refeeding mortality to levels
above that associated with qualitative and quantitatve aspects of deer refed.

�-528-

Probably a combination of these factors contributed to the disparity
between refeeding mortality observed in the field and in this study.
It is suggested that if starving deer are refed ad libitum a high quality
diet, such as Diet I, without alfalfa, there will be little refeeding mortality.
However, results of this study can not be applied to the problem of deer
starving in the wild.

Rather, additional refeeding experiments must be

conducted with deer on winter range.

Furthermore, it should be recognized

that winter feeding should not and cannot be conducted until the philosophy
and logistics of winter feeding are fully explored and delineated.

�-529-

Chapter 5
Conclusions
1.

During malnutrition and' starvation deer retain sufficient quantities

and variety of rumen bacteria to produce metabolites required for
sustenance upon refeeding.
2.

Starved deer refed diets of high protein (15-18 percent) and high

NFE (51-58 percent) concentrations do not become toxic or experience
metabolic upsets.
3.

Successful refeeding of starved deer is dependent upon physiological

condition of the deer rather than absolute length of time starved ..
4.

During malnutrition and starvation deer catabolize stored fat and

muscle protein to provide the body with required energy-metabolites.
When stored fat reserves are depleted. muscle protein catabolism increases.
Catabolism of muscle protein cannot provide adequate amounts of energy·
precursors; deer reduced to this state soon die. probably due to hypoglycemia.
Fawns succumb to starvation before does. probably due to lower fat stores
and faster catabolism of these fats. resulting in earlier depletion of fat
stores. increased muscle protein catabolism and hypoglycemia.
5. Mortality of refed deer in the wild probably resulted from various
combinations of: a) inadequate energy intake; b) poor quality feeds used
producing on1asal compactions or disruptions of digestive processes and
infections of digestive organs; and c) increased severity of environmental
stresses.
6.

Refeeding survival of starved deer is dependent upon quantity of feed

ingested as wel l as quality.

Certain feeds. such as "poor-quality"

alfalfa.

will cause digestive upsets. however. and should be identified and excluded
from feeds fed starving deer.

If starving deer receive all they can eat of a

high-quality diet, they will probably survive.

��-531PART III
SUPPLEMENTAL FEEDING OF WILD DEER
Chapter 1
Feeding Experiments, Winter 1973 - 1974
Introduction
Studies of penned deer under controlled experimental conditions were
conducted to determine a desirable composition of pelleted feed for nutritionally
stressed deer in winter (Nagy 1970,1971,1972,

1973).

When this pelleted feed

was offered in the field to wild deer in 1972-73 winter, feeders were visited,
but wild deer did not remove significant amounts of the ration.

But, at the

same time, it was noted that wild deer were attracted in considerable numbers
to trap sites baited with alfalfa hay. This suggested that wild deer could be
attracted to feeder sites until they discovered the pelleted ration by baiting
them in with alfalfa hay.

In 1973-1974 winter, a study was designed to

test the hypothesis that alfalfa hay would decrease discovery time and increase
consumption of pelleted feed when compared with unbaited sites.
But simply getting wild deer to use'feeders may not contribute significantly
to the winter maintenance of a deer population in severe winters.

Dean (1972)

reported that domination of feed sites by aggressive adults precluded the
use of feed by sub-dominant deer (especially fawns).

Since fawns are presumably

under greater nutritional stress in winter than adult deer they would benefit
most from supplemental feeds.

If Dean's contentions are true, then feeding

methodology would have to be developed to circumvent this apparent behavioral
"bottleneck".

Unfortunately, Dean (1972) offered little quantification for

his theory of feeder dominance by aggressive adults.

Hence, we tried to

quantify dominance interactions of deer at a feed site.

�-532-

Methods and Materials
Object ive I
Pelleted feed (Table 44) was offered in la-quart galvanized buckets
to permit accurate weighing.

At each feeding location, two buckets of food were

placed in wooden boxes or "feeders" (Figure 12). This experiment was designed
to compare two treatments:

pelleted feed alone and pelleted feed with alfalfa

hay distributed in a radius of about 3.0-4.5 m around the feed box.

Each

treatment was replicated four times for a total of eight feeder locations
(Figure 13).

Feeders were placed in areas known to be frequented by deer.

(Elliot Creek, Grand County, Colorado, Sect. 27, 28, 33, 34, TIN, R80W, 6pm).
Deer activity, vegetation and topography in Elliot Creek are similar
along a 3.2 km length from section corner 29/28 to 27/26, so feeder locations
32/33
34/35
were situated at approximately 0.5 km intervals. At each location, feeders
were placed at locations where deer crossed Elliot Creek as evidence from tracks.
Feeder locations were numbered from 1 to 8, beginning at the west end of the
line.

Treatments were assigned to locations alternately after a random start

at location number one.

Thus, location numbers 1, 3, 5, and 7 were unbaited

and numbers 2, 4, 6 and 8 were baited.
The amount of pelleted feed removed at each location was measured at
the end of each week.

Measurements were taken once a week to minimize dis-

turbance of deer at feed sites.

Initially, each location was supplied with

16.4 kg of the pelleted ration.

Each week, after measuring amounts of feed

removed, each feeder was resupplied with pelleted deer ration so that the
total feed offered at each location (exclusive of alfalfa) equaled 16.4 kg
In addition to feed removal, notes were kept on relative abundance of deer
tracks and sign (abundant, moderate, little or none), approximate discovery
time of the feeder and numbers of deer observed at or near feeders.

�-533-

TABLE 44.

Composition of pelleted feed, used in experimental deer
experiment in Middle Park, Colorado, winter 1973-74.

Ingredient

Percent

Barley, pulverized

10.0

Corn, pulverized

30.0

Milo, pulverized

5.0

Oats, pulverized

7.5

Wheat Middlings (Mill Feed)

6.5

Bea t Pulp, shredded

2.5

Brewers Grain

35.0

Dicalcium Phosphate

1.0

Molasses, cane

2.5

Vitamin A, 0, E Premix

.002

Trace Mineral (P, K, Ca, P)

.005

TOTAL

100.00

�=:':':':
,(//~~--.--~---===~
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i iI~=='cC&gt;~1
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,./

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Figure 12.

Diagram of feeders used in the Middle Park deer feeding experiments,

,
.

\

---'

II(~(
-

" Vv\
0"'/

.

~

,

;

{/;(I

i; .. (. .(__

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iI· /: I

I

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winter 1973-74.

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�-535-

...

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Fig. 13. Location of feeder sites of Middle Park deer winter feeding
study, 1973-74.

�-S36-

Time of discovery was based on the presence of deer tracks close
enough to the feeder so that deer could have removed feed. coupled with
removal of measurable amounts of food.

Differences in feed removal from

baited vs. unbaited sites were analyzed by an unpaired test (Dixon and
Massey 1969).
Objective II
One simple feeder was placed in an area known to be frequented by
deer (Fig. 16).

This feeding station was used to attempt to quantify and

qualify aggressive behavior of deer at feeders to determine if such behavior
restricts the use of feeders by sub-domi'nant deer (Table 4S).

Observations

were made from a distance of approximately O.S km. using 8 x binoculars and
a zoom lS-60x spotting scope.

Forty-three, 60 minute observation periods

were made alternating between morning and evening observations.

In addition

to daily observations, weekly measurements were made to determine the amount
of feed removed.
Measurement #1:

Record the number of dominant interactions that occured

at the feeder by sex, age group.
Dominant
Doe
Doe
Doe
Buck
Buck
Buck
Fawn
Fawn
Fawn
Measurement #2:

Possibilities were:
Subdominant
Doe
Buck
Fawn
Buck
Doe
Fawn
Fawn
Doe
Buck

Record amount of feeding time (minutes) spent at the

feeder by each sex and age group.
Measurement #3:

Record the amount of feed removed (kg) once each week.

�-537Data form·to be used in recording

TABLE 45.

deer behavior near feeding stat1ons~

BEHAVIOR BAIT SITE
(BBS)
Date:
Observer:

Time

Maximum No. of Deer
at Feeder Site:

}~ximum No. pf Tagged Deer
at Feeder Site:
Tag!\o.'s:

A I:: Adult
B .;Buck
D ~ Doe

B

End:

F' :: Fawn

Dominance Surr:mary

DL ::Large deer
DS = Small deer

Sub=Dorrdnan t

Dominant
A

Begin:

F

D

n

A

..

II

I

m.

F

I

I

I
-

-;

:

I,

I

I

I
I

f

I"

.

.

I

Feeding Hinutes
A

B

D

F.

Total

Reoarks:

�-538-

Measurement #4:

In the past two years (1972 - 1973 and 1973 - 1974)

deer have been trapped and individually marked with conspicuous collars in the
vicinity of this feeder site.

The number of marked deer still present will

be determined from intensive reconnaissance of the general area.

The ratio

of marked to unmarked deer using the feeder will be used as a basis for
estimating total use, as follows:
(a) Total deer use (D) _ all deer recorded at feeder (d)
Total marked deer(M) - marked deer recorded at feeder (m)
Md
(b) D -m

Measurements 1, 2, and 3 were used to estimate amounts of feed removed by
each sex and age class, a5suming equal removal per unit time by the three
classes recognized.
ObJective III
The following factors were considered in estimating the cost of emergency feeding of wild deer:
1. Calculate the amount of feed required to feed a specified number of
deer:
a. Decide what percent of the-population is to be fed (50% 5000
deer, 25% 2,500 etc.)
b. Decide how many kolograms of pelleted feed will be fed per deer
per day (ad libitum to all deer or feed 50% of ad libitum consumpti on and depend on the rema inder to be supplTed by ava ilab 1e
forage.
2. From experience in objectives #1 and #2 and from previous deer distribution studies decide speculatively on the number of locations needed to
feed the specified number of deer.
3. Estimate numbers and kinds of feeders needed to feed the specified

�-539-

number of deer the specified amount.
4. Set an arbitrary length of feeding period based on expected winter
nutrional stress period for Middle Park deer.
5. Estimate man-power, equipment, feed and delivery cost to achieve the
objectives.
Results and Discussion
Objective I
From the period of February 17 to March 29, 1974, a combined total of
560.22 kg. of pelleted feed was removed from both baited and unbaited feed
bunkers (Table 46).

A total of 259.79 kg. was removed from the unbaited

sites while 300.43 kg. was taken from the baited sites.

There was no signi-

ficant (P&lt;0.05) difference in the amount of feed removed from sites baited
with alfalfa and those left unbaited.

The time required for deer to discover

and use the feed bunkers varied from two to four days with no apparent preference for baited vs. unbaited sites.

However, due to the weekly visitation

schedule coupled with the difficulty in determing fresh tracks, deer discovery
of the feeders could only be approximated.
No quantitative measurement was made to determine the number of deer
using the feeders, however, ten to twenty-five deer were observed at or near
the feed sites each week when the feed buckets were weighed and refilled.
Summary
Although there was no significant difference in feed removal or initial
discovery time between baited and unbaited sites in this experiment, it is
possible that the 0.5 km distance between baited and unbaited feeders may
not have been of sufficient magnitude to independently test baited feed sites

�-540versus

unbaited sites.

However, for the purpose of this study an effort

was made to locate all feed bunkers in an area as similar as possible in order
to minimize the effect of location on deer use.
of 0.5 km between feeding sites.

TABLE 46.

Total amount of pelleted feed removed (kg) from baited and unbaited
feed bunkers from February 17 to March 29, 1974.

Date

2-17-74
2-24-74
3-1-74
3-8-74
3-15-74
3-22-74
3-29-74

TOTAL
Weekly Mean

This necessitated a maximum

Pelleted Feed Removed (kg)
Unbaited
Baited
12.95

8.63

49.77

50.45

22.09

54.09

47.04

65.45

14.09

26.36

48.40

46.36

65.45

49.09

259.79

300.43

37.11

42.91

Objective II
Measurement #1. A single feed bunker was filled with 16.4 kg of pelleted
feed and baited with approximately 1/2 bale of alfalfa hay and 34.0 kg of
apple pulp.

An effort was made to attract as many deer as possible to this feed

site in order to attempt to quantify and qualify aggressive behavior of deer at
feeders and to determine if such behavior restricts the use of feeders by sub-

�-541dominant deer.

For the purpose of this experiment, as aggressive action was

intrepreted as any pugnacious action displayed by one deer over another.
From February 17 to March 29, 1974, a total of 38 dominant interactions were
recorded for 43 hours of observation.

Due to observation distance and varible

weather conditions which made sex and age classification difficult, deer observed at the feed site were recorded only as adults or subadults.

Table

(47) shows the total number of dominance interactions observed between adults
and subadults.
Sixty-eight percent of the aggressive activity at the feed bunker was
observed between adults, while thirty-two percent of the encounters Occurred
between adults and subadults.

Dominance appeared to take several forms.

In

some cases lowered ear tips and extended head toward another animal was
sufficient to force the subordinate deer away.

Other interactions were more

forceful and resulted in a series of lunges or strikes with the forefeet toward
the opponent.

The intimidated deer usually retreated 5-10 meters while the

dominant deer returned to the feed bunker.
Measurement #2.

Deer spent approximately 17 hours at the feed bunker.

When more than four deer were at the feed bunker together it became difficult
to record actual feeding minutes for each individual deer.

Therefore, deer

were recorded as feeding as long as they remained within 3 meters of the
feeder.

This included feeding on the pelleted ration placed in buckets

within the feeder as well as the alfalfa and apple pulp attractants located
in a 3 meter radius around the bunker.

Table (48) shows the total feeding

minutes recorded for adults and subadults during this study.

Adults were

observed at the feed bunker a total of 826 minutes which accounted for 81
percent of the time.

Subadults spent 193 minutes at the feeder or 19 percent

�-542-

TABLE 47.

Total number and percent of aggressive dominant interactions recorded between adult and subadult deer.

Age Class

Number of Dominant Interactions

Percent

Adult

Adult

26

68.42

Adult

Subadult

12

31.58

38

100

TOTAL

TABLE 48.

Total minutes and percent of time spent at the feed bunker by
adult and subadult deer.

Age Class

Minutes observed at feed bunker

Percent

Adults

826.0

81.06

Subadult

193.0

18.94

TOTAL

1019.0

100

of the time.

No nocturnal observations were attempted although occassionally

deer were still at the feed bunker when darkness dictated termination of the
observation period.

Time spent at the feed bunker was recorded as long as

deer were present and could be classified as adults or subadults.
Measurement #3. Approximately 147.6 kg of pelleted feed was removed
from the single feed bunker during this experiment (40 days).

Assuming an

equal removal rate for adults and subadults approximately 119.56 kg of pelleted feed was removed by adults while a total of 28.04 kg was removed by

�-543-

subadults.

These calculations are based on the percent time spent at the feeder

by adults and subadults.
Measurement #4. A total of 48 deer were trapped and individually marked
with conspicuous collars in the vicinity of the feed site.

Eight of these

marked deer were observed at the feed bunker on different occasions during the
study period.

By knowing the total number of marked deer as well as the total

number of marked deer observed and the overall total of deer at the feeder an
estimate can be made on the total number of deer using the feeder over the 40 day
period.

This value is calculated using the formula D=Md where M{total marked
m

deer)=48; m{total marked deer observed feeder)=8; d{total deer observed at the
feeder)=84.

Therefore, an estimated 504 deer used the feed bunker during this

experiment.
Weather Data at Behavior Study Site
o

Mean low temperature during the study period was -11.1 C while the mean
o

high was +1.11 C. Thelast week in February was the coldest period with a
o

0

weekly low mean temperature of -25 C and mean weekly high of -11.1 C. Maximum snow accumulation during this period was approximately 38 cm. which occurred
during the last week of February.

From March 5 until the termination of the

experiment no significant amounts of snow were recorded.
Deer activity and feed removal observed at the feeder remained relatively
constant throughout the study period.

Snow accumulation decreased from about

38 cm. at the beginning of the study to approximately 0 cm. by the end.

Through-

out the study deer continued to be observed at the feeder even though natural
vegetation appeared to be available.
Summary
Lack of adequate controls in number, sex and age of deer using the feeder,

�-544-

as well as weather variability, animal identification, and the subjective
interpretation of animal behavior make it difficult to draw any definate
conclusions from this experiment regarding the effect of aggressive behavior
on subdominant deer use of feed bunkers or numbers and kinds of deer visiting
a feed site.

The data in this study appear to indicate that aggressive action

does exist between deer in this type of feeding situation however, it is
difficult to determine the extent of this dominant behavior in precluding
the use of feeders by subdominant deer.

The values obtained for feeding minutes,

number of dominant interactions and total number of deer using the feeder
were calculated based on several assumptions: 1) deer could be identified as
either adult or subadult; 2) all deer within 3 meters of the feed bunkers
were considered eating; 3) marked deer visited the feed site at the same rate
as unmarked deer.

��-546-

Chapter 2
Estimating the Cost of Emergency Winter Feeding
of Mule Deer
In estimating the cost of emergency winter feeding of mule deer in Middle
Park a hypothetical situation was created.

In this theoretical case an approx-

imated value was calculated for pelleted feed, manhours, equipment, material and
delivery cost required to feed a specified number of deer for a predetermined
period of time.

Based on these estimates a cost per deer value was calculated

as well as an overall total cost for the entire feeding program (Table 49).
Cost estimates will vary depending on the level of manpower used to carry out
and supervise the feeding program.

Manpower cost will also vary depending on

if a particular emergency feeding operation is considered a special function
or if it is included in the daily work schedule of the personnel involved.
For the purpose of this experiment, manpower cost was considered as a special
function and estimated on an hourly basis.

In this example wildlife conser-

vation officers supervised the emergency feeding program at an estimated hourly
rate of $5.00 an hour.
In this hypothetical case, it was arbitrarily decided to make an effort to
feed 2500 deer or one-quarter of the estimated winter population of deer in
Middle Park.

Since a severe winter in Middle Park consisting of low temper-

atures (-17.8° C to -1.11° C), wind and deep crusted snow ( 38 cm) may nutritionally stress deer a minimum of three weeks, it was recommended that the
feeding program be calculated for a 21 day period (Gill-personal communique).
The length of artificial feeding may vary from year to year depending on the
severity of the winter, available manpower and the economic situation.

�-547TABLE 49.

Estimated cost of manpower, feed, equipment and material
necessary to maintain 2500 deer for 21 days in an emergency
winter feeding program.

Material or Service

Manhours

Manhour cost @$5.00/Hr.

Commercial Pelleted
Feed (75 ton)

Material Cost
$12,612.00

144 Feed Bunkers
(22.68 kg. capacity)

144

$720.00

2,160.00

Transportation cost
(Ft. Co11ins to
Kremmling)

80

400.00

75.00

2816

14,080.00

Feed Dispersal
a) gasoline

704.00

b) food &amp; lodging
Subtotal

3,080.00
3040

15,200.00

TOTAL

18,631.00

$33,831.00

Previous data indicates that penned fawns weighing approximately 45 kg
require 1.0-1.3 kg per day of a high energy-high protein ration to maintain
body weight (Gill and Baker' 1974).

The cost of this pelleted ration (Table 49)

is approximately $168.00 per ton.

Therefore, in order to maintain 2500 deer

for 21 days it would require a minimum of 75 tons of feed at a cost of $12,612.60.
In order to adequately distribute feed to deer and minimize spillage and
waste, feed bunkers would be needed.

To feed 2500 deer, it would require ap-

proximtely 144 of the simple wooden feeders (22.68 kg capacity) used in Part I
of this study (Fig. 15).

The minimum material and labor cost for these feeders

would be approximately $20.00 per feeder or a total of $2,880.00.

This type of

feeder would require daily refilling during the feeding period which would in-

�-548-

crease manpower and equipment cost.

Fewer and more elaborate self-feeders could

be used in order to minimize maintenance cost, however these would be more expensive to build and logistically more difficult to situate.

Also in order to

reduce aggressive interaction by dominant animals, several smaller feeders
would seem more reasonable than fewer large feeders.
Further cost would be accumulated in transporting this large quanity of
feed and feeders to a central location such as Kremmling, Colorado.

Here the

feed and feeders would have to be stored temporarily until they could be distributed to the critical concentrations of starving deer.

The minimum estimated

cost for this phase of the feeding program would be approximately $475.00.
The process of actually distributing the feed to the deer and daily visits
to each feeder location for replenishing the feed supply would require the major
emphasis in planning, organizing and implementation of manpower and equipment.
The highest cost of the entire feeding program would be incurred at this level
of operation due to manpower, equipment and per diem expenses.
In this hypothetical situation, four locations were selected in Middle
Park for deer feeding sites.

Each location was to be maintained by four wild-

life conservation officers.

Each crew was to have access to one snow cat and

two snowmobiles with which to transport feed and feeders for the 21 day feeding
operation.

The cost associated with this phase of the feeding program includes

1) gasoline and maintenance for pickups, snowcats and snowmobiles; 2) manhours
for 16 wildlife conservation officers (WCO's) for 21 days; 3) food and lodging
for 14 weo's.

The total estimated cost for this phase would be a minimum of

$17,864.00 (Table 49).
Therefore, total expenditure for the entire feeding program from the time
the feed is bought, to the time it reaches the deer would amount to approximately

�-549-

$33,831.00.

Hence, to feed 2500 deer for 21 days would require a minimum cost

of $13.53 per deer.
Summary
Each emergency winter deer feeding situation is different and the associated cost will fluctuate depending on the number of deer to be fed, duration of the feeding period, feed cost and level of manpower and equipment
utilized in implementing the feeding program.

It should be emphasized that the

cost value computed in this example is based on a hypothetical case and not
on a actual feeding operation.

�-550LITERATURE CITED
Association of Official Agricultural Chemists. 1965. Official methods of
of analysis. lOth Ed. Wash., D.C. 957 pp.
Balch, C. C. and R. C. Campling. 1965. Rate of passage of digesta through
the ruminant digestive tract. In Physiology of Digestion in the Ruminant,
pp. 108-123. R. W. Dougherty (Ed.). Butterworths, Wash., 480 pp.
Barnes, Ella M. and Glenna C. Burton. 1970. The effects of hibernation on the
caecal flora of the thirteen-lined ground squirrel (Citellus tridecemlineatus). J. Appl. Bact. 33:505-514.
Blampied, P. H., D. A. LeCheminant, and I. M. Howey. 1964. A case of bovine
omasal impaction. The Vet. Record. 76(19):533-534.
Blood, D. C. and J. A. Henderson. 1968.
Wilkins Co., Baltimore, Md. 924 pp.

Veterinary medicine.

Williams and

Burleigh, I. G. and E. A. Davis. 1967. Studies on the endogenous metabolism
and senescence of starved Sarcina lutea. Biochem. J. 102:236-250.
Carhart, A. H. 1943. Fallacies in winter feeding of deer.
Wildl. Conf. 8:333-338.

Trans. N. Am.

Castle, E. J. 1956. The rate of passage of food stuffs through the alimentory
tract of the goat. 1. Studies of adult animals fed on hay and concentrates.
Brit. J. Nutr. 10(1):15-23.
Cheatum, E. L. 1956.

Too many deer?

Conservationist. 10(6):2-4.

Church, D. C. (Ed.). 1969. Digestive physiology and nutrition of ruminants.
Vol. 1. Oregon State University ,Book Stores Inc. Corvallis. 306 pp.
Conway, E. J. 1958. Microdiffusion analysis and volumetric error. (Fourth
Ed ition). Macmillan Co., N.Y. 465 pp.
Coop, I. E. 1949. The effect of starvation and of feeding after starvation,
on the metabolic activity in the rumen. New Zealand J. Sci. and Tech.
31(A):1-12.
Cowan, R. L. 1962. Studies on antler growth and nutrition of white-tailed
deer. In Proc. First Nat. White-tailed Deer Disease Symposium. Univ.
of Georgia, Athens. pp. 54-60.
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Wisconsin Cons. Dept. Tech. Wildl. Bull. 14. 279 pp.
Davenport, L. A. 1939. Results of deer feeding experiments at Cusino, Michigan.
Trans. N. Amer. Wildl. Conf. 4:268-274.
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of Escherichia coli. Biochem. J. 95:332-343.

�-551Dean, R. 1972. Deer ration research.
Div., Fed. Aid. Prog. W-70-R-2.
p.6.

Oregon State Game Commission, Research
Game Res. Rep. July, Study 5, Job 1,

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6(1):46-47.

1968.

Feeding deer to death.

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Nat. Wildl.

Game Res. Div.

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19 (5):10-13.

Food is the key to deer management.

Idaho Wildl. Rev.

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533 pp.

A case of distension of the omasum.

The rumen and its microbes.

Academic Press, New York.

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.
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April 15 (2):1-8.
Kelsey, P. M.

1973.

1966.

Can we feed deer?

Winter deer feeding.

Colorado Outdoors.

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The Will iams

March-

�-552Li, J. C. R. 1968. Statistical inference.
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N.Y. 533 pp.

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1962. Animal Nutrition.

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McGraw-Hill Co.,

Rumina1 microorganisms

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.
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App1. Micro.

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�-553-

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.
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~1. B. Saunders

�-554-

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. Church, D. C. (Ed.). 1972. Digestive physiology and nutrition of ruminants. Vol. III. Oregon State University Book Stores Inc. Corvallis.
350 pp.
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1971.

J. Ger.. Mi-

The physiology of starvation.

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                  <text>October 1974

-1-

.JOB :r'ROGRESS REPORT

State of

---

Project No.
\'brk Pl.an No.

Job Title

COLORADO
W-88-R-19

Migratory Bird Investigations

1

Job 1\0.

1

_

Waterfowl Production Surveys

Period Covered: April 24, 1973 to June 30, 1973
Personnel: C. Bryant and Staff, MOnte Vista National Wildlife Refuge; W.
Wilson, Brown's Park National Wildlife Refuge; F. N. Folks, C. Jensen and
A. Regenthal, Utah State Division of Wildlife Resources; R. Blumberg, R. Clark,
J. Corey, R. Desilet, W. Dolezal, R. Hopper, J. Lotentzson, R. Lowry, F. Rinella,
W. Russell, K. Wagner, M. Zgainer, and M. Szymczak, Colorado Division of
Wildlife.
AIlSTIJ\CT

Water conditions were considered good in North Park, and the Cache La Poudre
and South Platte River Valleys, but POOi[' in the San Luis Valley. The total
estimated number of duck breeding pairs increased to 56,539, a level approximately 19 percent above the 1972 population and nearly equal to the long
term average. Major increases in the duck breeding popu.lation were noted in
North Park and the Cache la Poudreand Yampa Valleys. The number of redheads
in the breeding population increased markedly. The mallard population
remained numerically stable. The post-nesting season population of Canada
geese in Moffat County was about 10 percent above the 1972 level, even
though production was down approximately 7 percent. Gosling production in
north-central Colorado was about 31 percent above the 1972 level but the
overall population remained stable.

��-3-

WATERFOWL PRODUCTION SURVEYS
Michael R. Szymczak
P. S. OBJECTIVE
To formulate waterfowl harvest regulations for Colorado.

SEGMENT OBJECTIVES
To estimate the number of duck and goose breeding pairs, by species, in
each of Colorado's major waterfowl breeding areas.
METHODS AND MATERIALS
Present duck breeding pair and production surveys consist of a breeding pair
inventory of only major production areas.
The 1973 duck breeding pair surveys were conducted during the period of May
14 to June 30. Surveys in North Park and the Cache 1a Poudre and South
Platte Valleys were conducted exclusively from the air. Ground counts were
made in the Yampa Valley and Brown's Park. In the San Luis Valley intensive
aerial counts were adjusted for visibility by air-ground comparison studies.
Pair estimates for the Monte Vista National Wildlife Refuge in the San Luis
Valley were obtained from'nesting transects. All survey methods and sample
areas for ducks remained the same as in previous years.
Surveys of Canada goose production were conducted within the period of
April 24 to June 16. Production estimates for Moffat County were made from
as complete a count as possible of hatched or active nests and brood size.
Population estimates in north-central Colorado were obtained from counts of
goslings and adults conducted from the ground during the period in which
the birds were flightless. The west-central Colorado survey was conducted,
however, poor results were obtained and are therefore not presented in this
report.
All flying was accomplished with a Cessna 185 aircraft. Areas sampled by
section or block were flown with one observer, while two observers were
used in sampling by transect.
On the basis of these studies, 'a report is submitted to the Bureau of Sport
Fisheries and Wildlife, which constitutes Colorado's part in the annual
continental cooperative breeding ground survey.

RESULTS AND DISCUSSION
Water conditions in Colorado's major waterfowl breeding areas were found to
be quite variable throughout the state during the survey period. In spite
of a record snowpack in mountain areas surrounding the San Luis Valley,

�-4-

the Valley proper was as dryas

in 1972 in most areas.

A late spring in southern Colorado retarded both the runoff from the surrounding mountains and irrigation activities in the Valley. Therefore,
many of the ditches and ponds resulting from irrigation water were dry.
Because of the expected late water, prospects were excellent for brood
survival. Heavy winter snows produced excellent water condition for birds
breeding in North Park. In the Cache la Poudre and South Platte River
Valleys, late spring storms filled most marshes and drainage basins, but
retarded the irrigation season. High water on rivers in northwestern
Colorado resulted in some flooding loss of Canada goose nests, particularly
on the Yampa River.
The number of duck breeding pairs in 1973 increased 19 percent above the
1972 level, essentially equalling the long term average (Table 1). All areas
with the exception of Brown's Park recorded increases over the 1972 level.
The major increase in terms of numbers was noted in North Park. However,
neither North Park nor the other major Colorado duck breeding area, the San
Luis Valley, reached the long term average level in numbers of breeding
pairs.
Table 1. Summary of Colorado's duck breeding ground popUlation estimates
in selected areas, 1973.

Total Est. Breeding Pairs
Long Term
1973
1972
Averagell

Percent Change
From
From Long
1972
Term Average

San Luis Valley

24,942

23,509

27,335

+ 6.1

- 8.8

North Park 2:./

14,255

8,922

16,176

+59.8

-11.9

South Platte Valley

7,917

7,019

6,062

+12.8

+30.6

Cache la Poudre Valley

5,852

4,630

3,107

+26.4

+88.3

Yampa Valley

2,571

1,857

2,814

+38.4

- 8.6

Brown's Park

632

1,339

1,045

-52.8

-39.5

56,169

47,276

56,539

+18.8

- 0.7

Area

Total

II San Luis Valley and North Park averages are based on results of 1964
through 1972 and 1968 through 1972 surveys, respectively, because of changes
in survey methods utilized prior to those dates. Figures for other areas are
19 year averages.
~I Aerial counts corrected by species from visibility ratios obtained in the
San Luis Valley.

�-5-

Increases in numbers of three species, the redhead, gadwall and pintail, were
responsible for the majority of the increase in Colorado duck breeding numbers
in 1973 over the 1972 level (Table 2). The redhead, which normally comprises
about 4 percent of the Colorado breeding populations made up about 11 percent
of the 1973 population. The mallard continues to be the major breeding
species in the state, making up about 40 percent of the 1973 population.
Table 2.

Species composition of Colorado's 1973 duck breeding population.
Number of Breeding Pairs
1954-721/
1973
1972
Average-

Percent SEecies ComEosition
1954-72
1972
Average
1973

Mallard

22,608

22,504

27,825

40.2

47.6

56.2

Blue-:winged and
Cinnamon Teal

6,374

6,532

4,852

11.3

13.8

9.8

Gadwall

7,400

4,666

5,129

13.2

9.9

10.4

Pintail

5,794

3,073

3,401

10.3

6.5

6.9

Green-winged Teal

1,161

2,094

2,283

2.1

4.4

4.6

Shoveler

4,556

4,152

2,183

8.1

8.8

4.4

Redhead

6,416

1,802

1,812

11.4

3.8

3.7

789

1,331

878

1.4

2.8

1.8

Other Divers

1,071

1,122

1,141

1.9

2.4

2.3

Total

56,169

47,276

49,504

Species

American Widgeon

1../ Species composition computed from data from all areas for the 19 year period
regardless of changes in survey methods.

The post-nesting season population of Canada geese along river systems in
northwest Colorado was estimated to be approximately 1,500 in 1973 (Table 3).
The total number of geese increased about 10 percent above the 1972 level,
but estimated gosling production was off about 7 percent (Tables 4 and 5).
High water along the Yampa River created major problems in estimating production. The results of the 1973 Canada goose production census in northcentral Colorado are presented by individual area in Table 6. Gosling production Lncreased to 1,450 birds, about 31 percent above the 1972 level and
28 percent above the average (Table 7). All five trend areas showed increases
°jn gosling production. The total population remained essentially stable
(Table 8).

�-6-

.Table 3. Number of Canada geese observed and estimated production, Moffat
County, Colorado, 1973.

Nesting
Pairs

Non-nesting
Birds

Total
Adults

Estimated No.
Gos1ingsl}

Total
Birds

7
10
5

257
88
116

271
108
126

18
37
12

299
145
138

461

505

67~/

582~/

40
29

206
83

286
141

151
105

437
246

Subtotal

69

289

431

256

683

Little Snake River

28

120

176

99

275

Grand Total

119

870

1,112

422

1,534

Area
Yampa River
Craig to Juniper Spgs.
Juniper to Cross Mtn.
Lily Park

222:./

Subtotal
Green River
Brown's Park
Dinosaur Nab'l Monum.1/

1:./ Calculated using average brood size and number of successful nests.
~/ Figures are minimal; extremely difficult to determine number of nesting pairs
and gosling production because of high water during and prior to survey.

1/ Area first surveyed in 1970; data supplied by F. Neil Folks, Utah State
Division of Wildlife Resources.

Table 4.

Total Canada geese observed, Moffat County, Colorado 1973.

Area
Yampa River

Number Geese Counted
1956-72
1973
1972
Average

Percent·Change
From 1956-1972
From 1972
Average

582

513

415

+ 13.5

+ 40.2

Green River
Brown's Park
Dinosaur Nat'l
Monum. Jj

437

264

139

+ 65.5

+214.4

246

300

363

- 18.0

- 32.2

Little Snake River~/

275

320

246

- 14.1

+11.8

1,534

1,397

1,163

+

+ 31.9

Total

!/ Area first surveyed in 1970
~/ Not included in survey until 1962.

9.8

�-7-

Table 5.
1972.

Estimated number of Canada goose goslings, Moffat County~ Colorado,

Number of Goslings
1956-72
1973
1972
Average

Area
Yampa River
Green River
Brown's Park
Dinosaur Nat'1
Monum. 1/
Little Snake River '!:../
Total

Percent Change
From 1956-1972
From 1972
Average

67

117

141

- 42.7

- 52.5

151

139

54

+

8.6

+179.6

105

136

121

- 32.8

- 13.2

99

61

78

+ 62.3

+ 26.9

422

453

394

6.8

+

7.1

1/ Area first surveyed in 1970.
'!:../

Not included in survey until 1962.

Fall Flight
The prospect for excellent late water conditions in the San l,uis Valley along
with the good water conditions in the other Illajorbreeding areas resulted in
excellent duck production and brood survival. Therefore, even though the
duck breeding population is about vaver-age , the fall flight should be much
better than average. The Canada goose flight from both northwest and northcentral Colorado will be very similar to 1972 •

.-vYJ ._ J , I} -7fJ

J}

/J

Prepared by__ -L.
1..:/ ~~,u.../=--\.~....:::;0=-_/.::.u'_
. ........,---&lt;~~~.
~:t:~~..cJ:.~&lt;"",0~,-:!...-1b.s......
__
Michael R. Szymczak.IO
(;
Assistant Wildlife Researcher

�Table 6.

Results of the north-central Colorado goose census, June 15, 1973 (continued).

Production
Area

Number
Broods

Total
Number Goslings

Total Number
Adults &amp; Yearlings

Total No.
of Birds

2
0
4
0
3

12
0
13
0
7
45
33
73
5
50
50
4
16
23
24
14
0
11

47
15
29
164
150
37
38
30
2
70
105
13
54
34
32
41
2
5

59
15
42
164
157
82
71
103
7
120
155
17
70
57
56
55
2
16

380

868

1,248

1,450

2,579

4,029

Denver

Total
Grand Total

Water Area
South Colorado Blvd. &amp;
Quincy
Blackmer Lake
Reservoir No. 3
Denver City park.
Sloans Lake
Federal Center Pond
C1aifie1d Reservoir
Gallup Reservoir
Columbine Country Club
Bowles Lake
Kings Pond
Lower Tu1e Lake
Marston Reservoir
Grant B Reservoir
Patrick Lake
Pinehurst Country Club
Kendrick Lake
Mohn Estate

8
14
1

2
4
5
5
3
0
2

I

~

0
I

�-11-

Table 7. Total number of Canada goose goslings produced in north-central
Colorado production trend areas, 1973.

Area

Number of Goslings
1969-72
1973
1972
Average

Percent Change
From 1969-72
From 1972
Average

Wellington

347

219

269

+58.4

+28.9

Fort Co11ins

347

3181:/

267

+ 9.1

+30.0

Loveland

106

74

75

+43.2

+41.3

Boulder

270

199

238

+35.7

+13.4

Denver

380

294

280

+29.2

+35.7

Total

1,450

1,104

1,129

+31.3

+28.4

!/ Includes 23 birds raised at Ft. Collins Wildlife Research Station.

Table 8. Total number of Canada geese observed in north-central Colorado
production trend areas, 1973.

Area

Number of Geese
1969-72
1973
1972
Average

Percent Change
From 1969-72
From 1972
Average

Wellington

828

731

853

+13.3

- 2.9

Fort Collins

843

868

747

- 2.9

+12.8

Loveland

283

149

177

+89.9

+59.9

Boulder

827

679

717

+21.8

+15.3

Denver

1,248

1,530

1,387

-18.4

-10.0

Total

4,029

3,957

3,881

+ 1.8

+ 3.8

��October, 1974

-l3-

J013 PROGRESS

j{EPOKT

St at e of

Pro j ect No.
i.JorkPlan No.
J~b Title

Migratory Bird Investigations

W-88-R-19

Job No.

1

Trapping and Banding Ducks and Geese

2

--~------------------

Period Covered; April 1, 1973 to March 31, 1974
Personnel~ C. Bryant and staff of Alamosa and Monte Vista National Wildlife
refuges, C. Braun, J. Carsella, J. Corey, D. Coven, H. Cox, C. Crawford, M.
DePra, R. Desilet, K. Dillinger, G. Eyre, B. Goetz, J. Gray, J. Gustafson,
T. Henry, J. Hicks, R. Holder, J. Jackson, D. Kevin, R. Kitzmiller, J~Lewis,
J. Lorentzson, R. Mason, K. Miller,. J. Miller, W. Mink, G. Nugent, R. Oakleaf,
B. Peterson, H. Schultz, B. Smith, G. Smith, S. Steinert, M. Sudheimer, M.
Szymczak, K. Wagner and R. Hopper.
ABSTRACT
Trapping activities in Segment 19 resulted in the banding of 14,176 ducks of
11 species at eight locations in Colorado. The mallard constituted the major
species in the banded sample, contributing 9,616, or about 68 percent of the
total. Pre-season banding in the high country yielded 9,336 ducks, including
2,982 in North Park, 2,039 in South Park, and 4,315 in the San Luis Valley.
¥~llards and pintails provided the bulk of the sample in these three areas.
Post-season banding resulted in a banded sample of 6,743 mallards, including
5,643 in eastern Colorado and 1,100 in west-central Colorado. Activities in
the west-central portion of the state represent a new banding study in an area
supporting a significant population of wintering mallards.
Goose trapping operations yielded a banded sample of 1,392 Canada geese in
Segment 19. Goslings, hand-reared or captured in the wild and transplanted in
various locations during the summer, accounted for 377 of this total. The
remainder were banded post-season in the Arkansas Valley (337), Cache la Poudre
Valley (270) and South Platte Valley (408).

��-15-

TRAPPING AND BANDING DUCKS AND GEESE
Richard M. Hopper

This report summarizes all waterfowl banding activities conducted under
Federal Aid Project W-88-R-19 for the segment year April 1, 1973 to March
31, 1974. Band recovery data are analyzed as part of another job (Work
Plan 1, Job 3) and thus, will not be included in this report. This report
simply presents a listing of numbers of ducks and geese banded by species
and location during the segment.

P. S. OBJECTIVE
To formulate waterfowl harvest regulations for Colorado.
SEGMENT OBJECTIVES
1.

To provide adequate samples of banded ducks and geese in Colorado's major
breeding and harvest areas for the purpose of accumulating migration and
annual mortality data.

2.

To report species and numbers of mallards winter banded in the South
Platte Valley, Ark?nsas Valley, and Bonny Reservoir as part of Work Plan
3, Job 6~

3.

To report transplant geese banded underWork

Plan 2, Job 2.

METHODS AND MATERIALS
Trapping and banding operations in Segment 19 continued the same as in recent
years with three major phases: (1) mid-summer banding of Canada goose transplants, (2) late summer (pre-season) duck banding, and (3) winter (post+ "
season) duck and goose banding. Goslings for transplanting were hand-reared
and alSo obtained from the wild by drive-trapping prior to the time the birds
were capable of flight. Pre-season duck banding was conducted in August and
September using baited Salt Plains traps (cage-type) to capture birds in North
Park, South Park, and the San Luis Valley. Night-lighting was again employed
experimentally to catch ducks in North Park during the pre-season period.
This technique is best described by Cummings and Hewitt (1964) and Drewien et
al. (1967). Post-season banding (January-February) utilized baited Salt Plains
traps and cannon-nets for duck trapping, and baited cannon-nets for goose
trapping.

�-16-

Ducks
The banded sample in Segment 19 consisted of 14,176 ducks of 11 species
(Table 1). An additional banding location (West Slope) was added to the
seven locations of recent years, with rhe South Platte Valley agafn yielding
the largest sample (3,350). North Park (2,982), the San Luis Valley (2,412)
and South Park (2,039) were next in sample size.
As normal, the mallard dominated as the species most abundant in the banded
sample, contributing 9,616, or about 68 percent of the total. Pintails and
teal were the only other species that added significantly to the banded sample,
with 3,137 and 1,075, respectively.
High Country Study
Objectives of the high country study of duck populations are given by Hopper
(1972). Species composition by age and sex for 9,336 ducks banded in the
three high country areas during 1973 is shown in Tables 2-5. Bandings in
the San Luis Valley were accomplished by both State and Federal personnel
(Table 4). The Bureau of Sport Fisheries and Wildlife is cooperating in
this study by banding a portion of the Valley quota on the Monte Vista and
Alamosa National Wildlife Refuges.
Mallards and pintails continued to supply most of the banded samples in all
three high country areas (Tables2-4).
They contributed 3,683 and 3,728
birds, respectively, for all three areas combined (Table 5). Generally poor
samples of the other species were obtained, with the possible exception of
green-winged teals (748), blue-winged and cinnamon teals (618), and redheads
(401).

Winter Banding
Post-season banding of terminal wintering populations of mallards was continued in eastern Colorado for the eleventh consecutive year under Work Plan
3, Job 6 (Investigation of Mallard Management Units of Eastern Colorado).
Quotas were easily attained by age and sex class .in all areas except the
the Arkansas Valley. The total banded sample amounted to 5,643 mallards for
the post-season period of Segment 19 in eastern Colorado (Table 1).
An additional 1,100 mallards were banded post-season on the Western Slope of
Colorado during Segment 19 (Table 1). This activity was the initiation of a
new banding program for Colorado. A study of at least five years duration is
planned in the Colorado-Gunnison-Uncompahgre River Complex of west-central
Colorado to investigate (1) population status, (2) breeding area derivation,
(3) harvest distribution, and (4) mortality rates of the mallard population
wintering in this area. A minimum population of 10,000 mallards winter here
and until now little has been done to assess their welfare.

�1/

August-September.

2/

- January-Feburary.

1/

Colorado-Gunnison-Uncompahgre

River Complex.

�-18Table 2. Age and sex composition of'duckS banded pre-season in North
Park, 1973.
Number of Ducks Banded
Age and Sex
Species

AM

1M

AF

IF

Total

Mallard

315

320

254

260

1,149

Gadwall

7

2

5

7

21

American Widgeon

5

3

1

3

12

Green-winged Teal

34

15

12

4

65

Blue-winged and Cinnamon Teal

13

12

6

10

41

Pintail

791

202

462

206

1,661 '

Redhead

7

1

23

0

31

Canvasback

1

0

0

0

1

Ring-necked Duck

1

0

0

0

1

1,174

555

763

490

2,982

Total

Table 3. Age and sex composition of ducks banded pre-season in South
Park, 1973.
Number of Ducks Banded
Age and Sex
Species

1M

AF

IF

Total

Mallard

250

190

192

113

745

Gadwall

0

0

1

0

1

American Widgeon

2

2

1

1

6

Green-winged Teal

281

123

59

100

563

Blue-winged and Cinnamon Teal

29

43

14

38

124

Pintail

186

123

147

109

565

Redhead

4

1

28

0,

33

Canvasback

1

0

0

0

1

Hooded Merganser

0

1

0

0

1

753

483

442

361

'2,039

Total

�-19-

Table 4. Age and sex composition of ducks banded pre-season in the San
Luis Valley, 1973 l/.
Number of Ducks Banded
Age and Sex
IF
IM
AF

Species

Total

Mallard

444

458

421

466

1,789

Gadwall

1

45

6

58

110

American Widgeon

a

2

1

a

3

Green-winged Teal

64

26

8

22

120

Blue-winged and Cinnamon Teal

176

108

62

107

453

Shoveler

a

a

0

1

1

Pintail

375

360

429

338

1,502

Redhead

105

49

129

54

337

Total

1,165

1,048

1,056

1,046

4,315

II

- Includes ducks banded by Federal personnel on Alamosa and Monte Vista
National Wildlife Refuges as part of high-country duck study.

Table 5. Age and sex composition of ducks banded pre-season in the three
high-country areas combined, 1973 !/.

Species

AM

Number of Ducks Banded
Age and Sex
AF
IF
IM

Total

Mallard

1,009

968

867

839

3,683

Gadwall

8

47

12

65

132

American Widgeon

7

7

3

4

21

379·
218

164
163

79
82

126
155

748
618

Shoveler

a

a

a

1

1

Pintail

1,352

685

1,038

653

3,728

Redhead

116

51

180

54

401

Canvasback

2

a

0

2

Ring-necked Duck

1

a
a

0

a

1

Hooded Mergariser

0

1

0

0

1

3,092

2,086

2,261

1,897

9,336

Green-wiIlgedTeal
B1ue-win~ed and Cinnamon Teal

;,~

Total

!/Inc1udesducks banded by Federal personnel on Alamosa ~nd Monte Vista National
Wildlife Refuges Ln. the San Luis Valley as part of the h1gh-country duck study.
~"~.

�-20-

Good age and sex ratios were obtained in the sample of 1,100 maliards
banded in west-central Colorado, with nearly equal total numbers for the
two banding locations (Table 6). Some suitable trapping sites have now
been found in the Grand Junction and Delta areas, but efforts in future
segments will be to include the Montrose area in the banded sample.

Table 6. Age and sex composition of mallards banded post-season
in westcentral Colorado, 1973-74.

AM

Number Banded
Age and Sex
IM
AF
IF

Grand Junction Mack - Palisade

167

113

141

82

503

Delta

140

161

134

162

597

Total

307

274

275

244

1,100

Location

Total

Geese
Nearly 1,400 Canada geese were banded during Segment 19 (Table 7). Summer
transplant geese contributed 377 birds to the banded sample. These goslings
were banded and released just prior to flight stage at Walden Lake in North
Park (100), Antero Reservoir in South Park (80), and Prewitt Reservoir (96)
and Jumbo Reservoir Annex (101) in the South Platte Valley. The sex compos ition of transplant geese was somewhat unequal at each release site, but
essentially equal numbers of males and females were the case for all areas
combined (Table 8). Work Plan 2, Job 2 discusses the transplant program in
more detail.
Post-season trapping resulted in the banding of 1,015 Canada geese in the
Arkansas Valley (337), Cache la Poudre Valley (270), and South Platte Valley
(408) (Table 7). Samples were not as large as desired in any of the above
three areas, but a considerable improvement was experienced over last segment
in the Cache la Poudre and South Platte valleys.

�~21-

Table 7.
1973-74.

Number of Canada geese banded by location and period of year,

Number Banded
Summer
Post-season 1./
Transplants 1/

Location

Total

Arkansas Valley

0

337

337

Cache la Poudre Valley

0

270

270

North Park

100

0

100

South Park

80

0

80

South Platte Valley

197

408

605

Total

377

1,015

1,392

1/
- June - July.
2/
- January - February.

Table 8.

Sex composition of goslings banded for summer transplants, 1973-74.
Number Banded
Sex
Male

Female

Total

North Park

59

41

100

South Park

46

34

80

South Platte Valley

86

111

197

Total

191

186

377

Release Site

The age composition of the banded samples of geese appeared to be distorted
in favor of adults at each banding location, while sex ratios seemed nearly
equal (Table 9). Additional information regarding post~season banding
activities can be found in the Job Progress Report for Work Plan 2, Jobs 5
and 6.

�-22-

Table 9. Age and sex composition of Canada geese banded post-seasol} in
eastern Colorado, 1973-74.

Number of Geese Banded
Age and Sex

Location

AM

1M

AF

IF

UF

Total

Cache la Poudre and South
Platte Valleys

270

51

7

5

345

678

Arkansas Valley

105

77

100

55

o

337

Total

375

128

107

60

345

1,015

LITERATURE CITED
Cummings, G. E., and O. H. Hewitt. 1964.
birds at night with light and sound.

Capturing waterfowl and marsh
J. Wildl. Manage. 28(1):120-126.

Drewien, R. C., H. M. Reeves, P. F. Springer, and T. L. Kuck. 1967.
unit for capturing waterfowl and upland game by night-lighting.
Wildl. Manage. 31(4): 778-783.
.
.
Hopper, R. M. 1972. Trapping and banding ducks and geese.
Wildl., Fed. Aid Game Res. Rept., Oct. pp. 13-21.

..;.

"J

Prepared by

.

-.

"

·1:~,c14J~/'&amp;25/"&gt;1
, Ri.chard M. Hoppe r« /
Wildlife Researcher

Backpack
J.

Colo. Div.

�October 1974

-23-

JOB PROGRESS
REPORT

State
Project

COLORADO
------~~~~~---------

of

No.

W
__
-_88_-_R_-_l_9

~

1

_

Work Plan No.
Job Title~
Period

Analysis

Covered:

Personnel:

April

Migratory

Bird Investigations

of Waterfowl Banding Data
~------~--------------------------1, 1973 to March 31, 1974

Michael Szymczak.

ABSTRACT
A decision was made to delay analysis of recoveries
of ducks banded preseason in North Park, South Park and the San Luis Valley until at least four
complete years of recovery data are available.
The project has beenre-written
to allow for an evaluation
and adjustment of annual banding quotas for each
area by species,
sex, and age.

��-25-

ANALYSIS OF WATERFOWL BANDING DATA
Michael R. Szymczak
P. S. OBJECTIVE
To formulate waterfowl harvest regulations for Colorado.

SEGMENT OBJECTIVES
To investigate migration, mortality, recovery distribution and relationships
among populations of ducks banded in Colorado's high country, specifically
in North Park, South Park, and the San Luis Valley.
METHODS AND MATERIALS
Recovery cards from bandings in high mountain areas and associated computer
printouts received from the Bird Banding Laboratory were filed. No other
work was accomplished under this job.

RESULTS AND DISCUSSION
An evaluation of banding programs in general and resulting analysis led to
the decision to delay of recoveries of birds banded in North Park, South Park
and the San Luis Valley until at least four complete years of recovery data
are available.

��October 1974
-27-

JOB PROGRESS 1I.EPORT

State

COLORADO

of

Projec.t:No.

W-88-R-19

Hork Plan No.

1

Job Title

Job No.

Bird Investigations
~1~2~

S_a_n
__L_u
__i_s--v_a_l_l_e~y--C_o_o~p_e_r_a
__
t~i_v~e--Ma--l~l_a_r_d
__I_n_v_e_s_t
__i~ga
__t_i_on

Pe.riod Covered:
Personnel:

Migratory

_
~ __

April 1, 1973 to March 31, 1974

Richard M. Hopper

ABSTRACT

The final report covering this job will be submitted to Wildlife Monographs
in Mayor June of 1974 for possible publication.
All work except typing
of the final manuscript was completed prior to the end of Segment 19.

��-29-

SAN LUIS VALLEY COOPERATIVE MALLARD INVESTIGATION
Richard M. Hopper
P. S. Objective
1.

To develop a harvest formula for the San Luis Valley mallard population.

2.

To determine hunter reaction to various types of hunting regulations.
SEGMENT OBJECTIVES

Evaluate results of the experimental hunting seasons since 1963 and prepare
final reports covering all phases of the investigation in cooperation with
the Bureau of Sport Fisheries and Wildlife.
METHODS AND MATERIALS
Compile and analyze all data in cooperation with the Bureau of Sport Fisheries
and Wildlife. Prepare final report and publish as Division Technical Bulletin
or Journal of Wildlife Management Monograph.
RESULTS AND DISCUSSION
The final report for this job is in the last stages of preparation and will
be submitted to Wildlife Monographs in Mayor June of 1974 for possible
publication.
As of March 31, 1974 the manuscript had received final editing by Division
editor Dr. Lee E. Yeager, and the four authors (R. M. Hopper, A. D. Geis,
J. R. Greib, and L. Nelson, Jr.). Typing of the final copy will be completed
by mid-April, 1974. The manuscript will then be sent to Washington, D. C.
where it will be reviewed by the Bureau of Sport Fisheries and Wildlife prior
to submission to Wildlife Monographs. The title select~d for this publication
is "Experimental Duck Hunting Seasons - San Luis Valley, Colorado, 1963-1970."

Prepared by ----JI/:...,.i.:·.•..•••
",&lt;=t;4-t.•..•
·./u.,.o::J:ol-.· -'?&lt;-----"~y
__ /~£~!~;.~, ·i;:...
•.·l-I
Richard M. Hopper
v'/'
Wildlife Researcher

_

��October

-31-

JOB PROGRESS

St ate

REPORT

-'-..:C..:O.:.;L:,::O.:.;R:.=.AD.::.O---:----,--

0f

Project

1974

Migratory

W-88-R-19

No.

Bird Investigations

1
Job NO.-'~1~4__ ---- __ ----~~--Determination
of Methods for Developing and Managing
Job Tit 1e __ w~a:::...t::...e::..:r::..:f_o::...w....:l:::...·....:H=a=b:..:i:.:t=a:.:t::...-..:.Annn===o
. .:.N:.:i:.:t:.:r:.:a:.:t:.:e;.....::.P-=o-=t:.:.:h:;::o=l:.
_.=B=l::::a=s.;::.t=i.:::n~ge-;::.S=t.;::.
__

Wark Plan No.

Period

Covered:

Personnel:

April

Richard

1, 1973-March

31, 1974

M. Hopper

ABSTRACT·

.Field work was re-initiated
in Segment 19 to continue the "life expectancy"
phase of.the pothole blasting study.
This is the only portion of the overall
investigation
remaining .to be completed. The 76 potholes on the study area
had an average depth of 32.0 inches in 1973, yielding a loss of lS.3 percent
from 1970 to 1973 and a 26.1 percent overall reduction from 1968 to 1973.
The lSO-lb potholes showed t he greatest loss of depth during five years of
life (30.S percent), while 2S-lb potholes showed the least (22.2 percent).
All sizes of potholes, regardless of block occupied, were invaded by emergent
vegetation.
The average proportion of surface area grown up to emergents
was greatest for 2S-lb potholes (S4.S percent) and least for 7S-lb potholes
.(36.1 percent).
A notable reduction was observed in the number of 2S-, SO- and 7S-lb potholes visited by ducks in 1973 compared to the 1968-1970 period, while an
increase was recorded in the number of lSQ-lb potholes used. The proportion
of total duck visits received through the years by lSO-lb potholes has
gradually increased (32.6-S8.9 percent), while that for the other three
sizes has generally decreased. .

��-33-

AMMONIUM NITRATE POTHOLE BLASTING STUDY
Richard M. Hopper

Field work was again initiated in Segment 19 (1973-74) on the pothole
blasting study following two years. of inactivity (1971-72, 1972-73). This
study was first statted in 1967, with all phases of the objective completed
and results' reported except for the "life expectancy" portion (Hopper 1972).
The life expectancy of the 84 potholes blasted in 1967 is an important part
of the overall study and the subject of this report deals with that determination. Obviously, a given pothole may exist physically for many years,
but because of loss of depth,encroachment of emergents, or for some other
reason, this pothole will lose its attractiveness to waterfowl after a
period of time. Its usefulness as waterfowl habitat has ended or been
greatly curtailed and the waterfowl manager can conclude that the pothole
has reached its life expectancy. Thus, in this study, duck use observations
are playing a major role in the evaluation of life expectancy, or what
might be better termed "useful life" of the potholes.
It might be appropriate here to briefly revie.w the results of the waterfowl
use and cost relationships among the four charge sizes of annnonium nitratefuel oil mixtures (AN-FO) as presented previously (Hopper 1972). It was
concluded in this phase of the study that 75-lb charges of AN-FO yielded
the most efficient potholes in terms of (1) lowest average cost per 100
square feet of surface area created and per duck visit received, and (2)
highest average number of duck visits per 100 square feet of surface area.
The l50-lbpotholeswere
next:most efficient, and over an extended period
of time may prove to have a longer useful life than those produced by the
other charge sizes.
P. S. OBJECTIVE
To evaluate various size potholes blasted with ammonium nitrate in terms
of (a) life expectancy, (b) plant succession, (c) soil and water characters,
(d) waterfowl use and hunting potential, and (e) cost.
SEGMENT OBJECTIVES
1.

Obtain pothole measurements.

2.

Observe and record duck use of potholes by species during the spring
period.

�-34-

METHODS
ANDMATERIALS
Methods and materials employed during Segment 19 remained the same as those
used in Segments 13 and 1'4 (Hopper 1968, 1969) and will not be repeated
here.
An additional method was used to evaluate pothole life expectancy.
This involved Visiting the potholes during the March-April period and estimating the proportion of the surface area of each pothole that was grown up
to emergent vegetation.
Twoestimates were made for each pothole during
this period, with the second estimate being made without prior reference
to the first.
An average figure was then obtained for each pothole and
comparisons were made by pothole size and amount of duck use received.
RES
UL
TS ANDDIseus SION
Pothole Measurements
Depth
Tables 1-3 list pothole depths in March-April 1973 by block (Fig. 1) and
charge size, as well as comparisons with measurements t aken in March 1968
and March-April 1970. Measurements in 1973 showed a continued loss of
depth with age for all potholes in the three bLocks,. Potholes of the 50-lb
charge size continued to maintain the greatest depth of the four charge
sizes in BlocksB and C, but for the first
time slipped slightly below the
average depth of 25-lb potholes in Block A. Average depths of l50..,.lb
pothdlesinBlock
A (27.6 inches) and 75-lb potholes in Block B (28.1 inches)
were considerably less than.those'for
the other sizes in their respective
blocks.
'
The greatest loss of depth occurred for 150-lb potholes in Block A between
1970 and 1973 (28.9 percent) and between 1968 and 1973 (35.2 percent).
The
least loss occurred for 25-lb potholes in Block e between 1970 and 1973
(4.7' percent), while the 75-lb potholes in Block e had the lowest overall
10s80f depth for the 1968-1973 period (15.8 percent).
Average depths and percent loss of depth by year and charge size are
presented in Table 4 for all blocks combined. The 76 potholes on the study
area yielded an average depth of 32.0 inches in 19.73, reSUlting in a loss of
15.3 percent from 1970 to 1973 and a 26.1 percent overall reduction from
1968 to 1973., After five years of life,
the 50-lb potholes had the greatest
average depth (34.4 inches) of the four charge sizes, with the 150-lb size
being the shallowest (29.9 inches).
The 150-lb potholes also showed the
most loss of depth during their life (30.5 percent).
The smallest potholes
(25:..1b) sustained the smallest overall loss of depth at 22.2 percent.

�-35-

Table 1. Comparison of pothole depth measurements in March, 1968, MarchApril 1970, and March-April 1973 by charge size in Block A.
Percent Loss of DeEth
From
From
From
1968-1973
1970-1973
1968-1970

Charge
Size 1/
(Lbs)

Average DeEth ~Inches)
1973
1968
1970

25

45.4

40.3

33.7

11.2

16.4

25.8

50

45.7

39.3

32.8

14.0

16.5

28.2

75

43.0

37.8

31.2

12.1

17.5

27.4

150

42.6

38.8

27.6

8.9

28.9

35.2

Total

.44.2

39.1

31.3

11.5

19.9

29.2

1/ Seven potholes of each charge size.
Table 2. Comparison of pothole depth measurements in March 1968, MarchApril 1970, and March-April 1973 by charge size in Block B.
Percent Loss of DeEth
.From
From
From
1968-1973
1968-1970
1970-1973

Charge
Size 1/
(Lbs)

Average DeEth (Inches).
1973
1968
1970

25

40.3

36.8

31.7

8.7

13.8

21.3

50

47.9

40.4

34.9

15.6

13.6

27.1

75

39.9

34.1

28.1

14.5

17.6

29.6

150

43.2

36.7

31.0

15.0

15.5

28.2

Total

42.8

37.0

31.4

13.6

15.1

26.6

!/ Seven potholes of each charge size.

�N

o

12m
Scale;&gt; :

1
o TS"
... 0;,?S'.

···0···

··0
...

• . '.

,fo

o ... 0

0··

0

..2.J"B

·0/~o

~0·.o ..·0,
..
.
··0..
.0.· 0 ..... (0
0'
.... (0.

o

0

0····0.J..o ..
..
O,s···(}0

.. 0
I

0

0

C:) (0

-.'..

W
.0\

.0

I

S"o

0

.,.
·0·
0
0 -.
0
~'\ ..',0
. . 0
.\:J
-. 0
0··

B~n('h

o®.
100 ... 0
.~
0
o

Area

o(~ .

o

C

·02S·

o

o

Fig. L Pothole blast:i.ng study area, showing
the position and size e&gt;fphotholes by study area ,

0°

o

�-37-

Table 3. Comparison of pothole depth measurements in March 1968, MarchApril 1970, and March-April 1973 by charge size in Block C.
Percent Loss of DeEth
From
From
From
1968-1973
1968,..1970 1970-1973

Charge
Size 1:./
(Lbs)

Average DeEth (Inches)
1973
1970
1968

25

39.5

33.8

32.2

14.4

4.7

18.5

50

47.3

40.0

35.5

15.4

11.2

24.9

75

39.8

·36.2

33.5

9.0

7.4

15.8

150

43.7

36.7

32.6

16.0

11.2

25.4

Total

42.9

36.9

33.7

14.0

8.7

21.4

II A total of 20 potholes, including 6 25-1b, 7 50-1b, 4 75-1b,

and 3 ISO-lb.

Table 4. Comparison of pothole depth measurements in March 1968, MarchApri11970, and March-April 1973 by charge size and for a11b1ocks combined.
Percent Loss of DeEth
From
From
From
1968-1973
1970-1973
1968-1970

Charge
Size 1/
(Lbs)

Average Depth (Inches)
1973
1970
1968

25

41.8

37.2

32.5

11.0

12.6

22.2

50

47.0

39.9

34.4

15.1

13.8

26.8

75

41.0

36.0

30.5

12.2

15.3

25.6

150

43.0

37.6

29.9

12.6

20.5

30.5

Total

43.3

37.8

32.0

12.7

15.3

26.1

1/ A total of 76 potholes, including 20-25 1b, 21-50 1b, 18-75 1b,
17-150 lb.

and

�-38-

Invasion by Emergent

Vegetation

It was thought that another indication of pothole longevity might be the
degree to which the potholes were invaded by emergent vegetation, thus
reflecting their usefulness and attractiveness to waterfowl as discussed
earlier.
Tables 5-7 present the estimated proportion of pothole surfaces
grown up to emergent vegetation by block and charge size. These data
show that no pothole. regardless of size or block occupied, was immune
from invasion by emergents.
All sizes of potholes in each block also
exhibited a wide range in the amount of emergent vegetation present.
Emergent vegetation consisted of cattail (Typha latifolia) and bulrush (Scirpus
spp.).
.
The average percent of surface area grown up to emergents was least for
50-lb potholes in Block A (28.9 percent) and greatest for 25-lb potholes in
Block C. Potholes in Block A, as a group, seemed to have the least affinity
for emergent growth, with an average of 36.8 percent of their surfaces grown
up to cattails or bulrush, followed by 43.9 percent for potholes in Block
Band 50.0 percent for those in Block C.
For all three blocks' combined, the average proportion of surface area
occupied by emergents was greatest for 25-lb potholes (54.5 percent) and
least for 75-lb potholes (36.1 percent) (Table 8). Potholes of the 50- and
l50-lb charge sizes showed percentages only slightly higher than that for
the 75-lb size, 40.1 and 42.0 percent, respectively.
Thus, with the possible
exception of 25-lb potholes, there appeared to ,be little difference among
the pothole sizes in regard to percent of surface area encroached upon by
emergent vegetation.
However, the amount of surface area that remains open in a pothole following
invasion by emergents may be more important than the proportion of the
surface area grown up to emergents in terms of usefulness or attractiveness
.,
to waterfowl.
For example, a l50-lb pothole with 75 percent of its surface
area grown up to emergents would still maintain a larger surface area of
open water than a 25-lb pothole with only 25 percent its surface area,grown
up to emergents.
Waterfowl use in relation to emergent invasion of potholes
will be discussed in the next section.

Waterfowl

Use

Duck use of the pothole study area was recorded during four observation
periods totaling 67.5 hours in 1973, (1) March 26-28, (2) April 16-20, (3)
May 1-4, and (4) May 16-18. Mallards accounted for over 95 percent of the
total use. Only 35 of 79 potholes (44.3 percent) were .use d one or more times
during these observation periods (Table 9) . This compares to an average of
58, or 70.7 percent, of 82 potholes used at least once during the three year
period of 1968-1970.
A considerable reduction was noted in the number of 25-,
50-, and 75-lb potholes visited by ducks in 1973, as compared to the 19681970 period, while an increase was observed in the number of l50,,:lbpotholes
used.
The greatest reduction occurred for the 25-lbpotholes,
followed by
the 50-lb potholes, 1. e., pothole size was inversely related to the amount
of decrease in the number of potholes visited at least once.

�-39-

Table 5. Proportion of pothole
vegetation,
March-:-April 1973.

Charge Size
(Lbs) .

surfaces

in Block A grown up to emergent

Percent of Surface Area Grown Up To Emergents
Range
Average

25

5.0 - 85.0

37.1

50

5.0

85.0

28.9

75

5.0

95.0

42.8

150

7.5 - 75.0

38.8

5.0 - 95.0

36.8

Total

Table 6 . Proportion of
vegetation,
March-April

pothole

surfaces

in. Block B grown up to emergent

.Charge Size
(Lbs) .

Percent of Surface Area GrownUp To .Emergents
Range
Average

1973.

·25

5.0 - 85.0

58.2

50

S.O - 95.0

48.6

75

5.0

75.0

31.4

150

5.0

85.0

37.5

5.0 - 95.0

43.9

Total

�-40-

Table 7. Proportion of pothole
vegetation,
March-April 1973.

Charge Size
(Lbs)

surfaces

in Block C grown up to .eme rgen t

Percent of Surface Area Grown Up To Emergents
Range
Average

25

20.0 - 100.0

68.2

50

20.0 - 80.0

42.8

75

12.5 - 90.0

33.8

150

7.5

80.0

55~0

7.5 - 100.0

50.0

Total

Table 8. Proportion of pothole
grown up to emergent vegetation,
Charge Size
(Lbs )

surfaces in all three
March-April 1973.

blocks combined

Percent of Surface Area Grown Up To Emergents
Range
Average

25

5.0

50

5.0 - 95.0

40.1 .

75

5.0 - 95.0

36.1

150

5.0 .•.85.0

42.0

5.0 - 100.0

43.4

Total

100.0

54.5

�-41-

Table 9. Number and percent of potholes used by ducks in 1973, compared to
the 1968-1970 period.
1973

1968-l97aY
Average Number
Used

Charge Size
(Lbs)

Number of
PothoLes

Number
Used

Percent

25

21

4

19.0

10.7

51.0

50

21

7

33.3

13.7

65.2

75

20 Y

11

55.0

17.3

82.4

150

17 '1:../

13

76.5

16.3

85.8

79

35

44.3

58.0

70.7

Total

Percent

J:./ From Hopper (1972).
'1:../ Twenty-one 75-lb potholes and 19 l50-lb potholes constituted the sample
during the 1968-1970 period.

The reduction in number of potholes utilized in 1973 could be a reflecti-on
of decreased attractiveness to ducks since the 1968-1970 period. However,
one must be cautious in comparing duck -use between years because of such
variables as duck density from year to year in a locality and annual
differences in dates of peak populations in relation to designated observation periods.
The number of duck visits received by size of pothole and year is compared
in Table 10 for all blocks combined. Total number of duck visits is not
comparable among years because of annual differences in hours of observation.
The main feature in Table 10 is that through the years the proportion of
total duck visits received by l50~lb potholes has gradually increased
(32.6-58.9 percent), while the proportion received by the other three sizes
have generally decreased.
Duck use of the various sizes of potholes during the four observation
periods in 1973 is shown by block in Tables 11-13 and for all blocks combined in Table 14. Block A was the most frequently used of the three blocks,
yielding 5.44 duck visits/hour compared to 2.75 and 1.15 duck visits/hour
for Blocks Band C, respectively. The peak.of duck use occurred during the
April 16-20 period, no doubt reflecting the early activities of the mallard.

�Table 10~

Charge Size
(Lbs)
25
50
75
150
Total

Comparison of number and percent of duck visits by charge size and year for all blocks combined.
1968
Number of
Duck Visits
Percent
54

11.0

80

1969
Number of
Duck Visits
Percent

1970
Number of
Duck Visits
Percent

1973
Number of
Duck Visits
Percent

8

4.4

45

8.9

16

7.7

16.3

28

15.6

48

9.5

21

10.2

197

40.1

60

33.3

148

29.4

48

23.2

160

32.6

84

46.7

262

52.1

122

58.9

491

100.0

180

100.0

503

99.9

207

99.9

I

~
I'.)
I

�-43-

based upon
Table 11. Duck use of potholes in Block A by charge size,
watching from observation points, 1973.
Average Number Duck Visits/Hour
Charge Size (Lbs2I7
Total
150
75
50

Number Hours
Observed

25

3/26 - 3/28

4.00

O.~/

1.00

1.50

2.92

5.4211

4/16 - 4/20

8.50

0.82

0.47

2.12

6.86

10.27

5/1 - 5/4

5.25

0.38

0.0

0.0

0.67

1.05

5/16 - 5/18

5.50

0.18

0.0

0.54

1.48

2.20

Total

23.25

0.43

0.34

1.16

3.51

5.44

Date

l./ Seven potholes of each charge size except for the l50-lb s Lz e which had
only six potholes.
~/ Combined duck use for all seven potholes in the row.

1/ Combined duck use for all potholes in the block.
Table 12. Duck use of potholes in Block B by charge size, based upon
watching from observation points, 1973.
Average Number Duck Visits/Hour
Charge Size (Lbs)17
Total
150
75
50

Number Hours
Observed

25

3/26 - 3/28

4.00

0.5~/

0.50

1.00

2.75

4.7511

4/16 - 4/20

8.00

0.50

0.25

0.50

3.50

4.75

5/1 - 5/4

10.50

0.0

0.10

0.86

0.76

1.72

5/16 - 5/18

5.50

0.0

0.0

0.0

0.36

0.36

Total

28.00

0.21

0.18

0.61

1.75

2.75

Date

l./ Seven potholes of each charge size.
~/ Combined duck use for all seven potholes in the row.

1/ Combined duck use for all potholes in the block.

�-44Table 13. Duck use of potholes in Block C by charge size, based upon
watching from observation points, 1973.

Average Number Duck Visits/Hour
Charge Size (Lbs) 17
50
75
150
Total

Date

Number Hours
Observed

25

4/16 - 4/20

5.50

O. rJ:-/

1.09

0.0

0.95

2.041/

5/1 - 5/4

5.25

0.0

0.0

0.0

0.0

0.0

5/16 - 5/18

5.50

0.0

0.36

1.02

0.0

1.38

Total

16.25

0.0

0.49

0.34

0.32

1.15

!/ Seven potholes of each charge size except for the 75-lb size which had only
five potholes and the l50-lb size with only four potholes.
~/ Combined duck use for all seven potholes in the row.

1/ Combined duck use for all potholes in the block.

Table 14. Duck use of potholes in Blocks A,
B, and C combined by charge size,
based upon watching from observation points,
1973.

Average Number Duck Visits/Hour
Charge Size (Lbs)17
50
75
150
Total

Number Hours
Observed

25

3/26 - 3/28

8.0

0.5rJ:-/

1.50

2.50

5.67

10.173/

4/16 - 4/20

22.0

1.32

1.81

2.62

11.31

17.06

5/1 - 5/4

21.0

0.38

0.10

0.86

1.43

2.77

16.5

0.18

0.36

1.56

1.84

3.94

67.5

0.64

1.01

2.11

5.58

9.34

Date

5/16 - 5/18
Total

!/ Twenty-one potholes of each charge size except for the 75-lb size which had
only 19 potholes and the l50-lb size with only 17 potholes.
~/ Combined duck use for all 21 potholes of the same charge size in the three
blocks.

2/ Combined duck use for all potholes in the three blocks.

�"':45-

The l50-lb pothole received
the most duck visits/hour
in Blocks A and B
(3.51 and 1. 75), but 50-lb potholes predominated in Block C (0.49).
The
75-lb potholes received the next highest use in all three blocks.
For all
blocks combined, average riuniber of duckvisitsihour
was directly proportional
to charge size (Table 14).
Overall duck use (duck ~isits/hour) .and that for 50;...and l50-lb charge sizes
were greater in 1973' that in any of the other three years of study (1968-:
1970) (Table 15). Tnaddition,150-lb
potholes far outstripped the other
. charge sizes in duck visits/hour during 1970 cind1973. These data indicate
'. that the pothole study area as a whole may be maintaining. its attractiveness
to waterfowl, but with a possible shift of use to150-lbpotholes.However,
as noted earlier,
caution must be exercised in making comparisons amongyears
with regard to these data.
.

')
..
Table 15.

Duck use by charge size

and year for all

.

'.

1/

blocks combined.-

.Average' NumberDuck Visits/Hour
1970
1969

Charge Size
(Lbs)

1968

25

0.62 .

0.26

0.73.

0.64

50

0.92

0.90

0.78

1.01

75

2.26

1.92

2.41

2.11

150

2.03

2.97

4.71

5.58

5.83

6.05

8.63

9.34

Total

1973

1/ Data for the years 1968-1970 from Hopper (1972).

Information regarding relationships
between duck use and proportion of pothole surfaces grown up to emergents are as yet inconclusive.
Another year
of data will be helpful.
However, there was some indication that ducks
avoided potholes that had 76-100 percent of their surface areas grown up to
emergents.
Only three of 200 total duck visits occurred on the 18 potholes
that fell in this category.
In contrast,
36 potholes with 0-25 percent of
their surfaces grown up to emergents received 123 duck visits.

�-46-

LITERATURE

CITED

Hopper, R. M•. 1968.
Determination
of methods for developing and managing
waterfowl habitat-ammonium nitrate
pothole blasting
study.
Colo. Game,
Fish and Parks. Div., Fed. Aiq.Game Res. Rpt , Oct.
p. 51-64.'
.

.

Hopper, R. M.1969.
Determination
of methods for developing and managing
'waterfowl habitat-ammonium nitrate
pothole blasting
study.
Colo. Game,
Fish and Pazks Div., Fed , Aid Game Res. Rpt.
Oct.
p , 73-85.
Hopper, R.M.
.
J.Wildl.

Prepared

1972.
Waterfowl use in relation
Manage. 36(2):459-468.

to size

and cost

of potholes.

by _.o.I,t;+' ~·,,",1.C'"'RA:"':i~=~·~..:lIia.r::"d-/c..r""":::"';;;"H~;,L.?!f9;..J~"-=e::;(""?~Mo:::.
~
•..
--'_
Wildlife

Researcher

�October

-47-

JOB PROGRESS

State

0f

Project

1974

REPORT

..::::C;:.O:.::.LO::.:l:...:~..i:...:\.D::...:O=__
_
No.

Work Plan No.

Migratory

W-88-R-19

Job No.

2

Job Title

Experimental Studies on Improving
The Status of Canada Goose Populations

Period

April

Covered:

Bird Investigations
2

_

6, 1973 to March 17, 1974.

Personnel:
J. Corey, M. DePra, W. Dolezal, J. Hicks, R. Hopper, T. Lines,
J. Lorentzson, W. Olmstead, S. Porter, K. Wagner, J. Wagner, L. Webster,
and M. Szymczak.

ABSTRACT
There were 3 successful nesting attempts at the Jumbo Reservoir Annex in
1973.
An additional 101 goslings were released on Jumbo Reservoir Annex.
One nest was established near Prewitt Reservoir in 1973. A total of 96
goslings were released on Prewitt Reservoir in 1973. Approximately
60 geese
were observed on Lake John Annex in North Park on April 6, 1973. A total of
105 adult and subadult geese and 16 goslings were observed on surveyed areas
in North Park on June 18; 1973. Unusually late spring breakup of the ice
and snow pack probably hampered nesting in North Park in 1973.
Seven known
nests were established in North Park in 1973, 4 hatched, producing an estimated total of 20 goslings.
Six birds which had been released in North
Park during previous years were recaptured during their moult at Wheatland
Reservoir, near Wheatland Wyoming.
One-hundred goslings were released on
Walden Reservoir in North Park in July 1973.
Eighty goslings were released
in July on Antero Reservoir in South Park.

��-49-

EXPERIMENTAL STUDIES ON IMPROVING
THE STATUS OF CANADA GOOSE POPULATIONS
Michael

R. Szymczak

P. S. OBJECTIVE
To investigate the success of attempts to establish breeding populations of
Canada geese in suitable habitat where they do not currently exist in Colorado.
SEGMENT OBJECTIVES
1.

To prepare a technical bulletin on the techniques
breeding population of Canada geese.

of establishing

a

2.

To expand the breeding flocks of Canada geese in the South Platte Valley
and North Park and continue efforts to establish a breeding flock of
Canada geese in South Park.
METHODS AND MATERIALS

Canada goose goslings for transplant purposes were obtained from three different sources: (1) goslings from the metropolitan Denver area; (2) goslings and
eggs from the Fort Collins area, and goslings from Faivre Ponds in the Boulder
area. All eggs were transported to the Fort Collins Wildlife Research Station
for incubation and subsequent raising.
All goslings were transported to the
Research Station, held for variable amounts of time, then transported to the
various release sites.
Birds were released on Walden Reservoir, Prewitt
Reservoir, Jumbo Reservoir Annex and Antero Reservoir.
Because of change in reporting
recoveries were analyzed.

deadlines,

no 1973-74 hunting

season band

RESULTS AND DISCUSSION
Preparation

of Technical

Bulletin

The first draft of the technical bulletin
restoration proj ect is nearly complete.

outlinifg

the Foothills

South Platte Valley-Jumbo

Reservoir

Annex

area goose

Approximately 100 Canada geese were observed in the Jumbo Reservoir area in
early spring 1973 (Szymczak 1973).
There were 3 reported successful nests
on the Annex, with brood sizes, of 1, 3 and 5. A total of 9 nests were established on ponds located on private land in the Jumbo Reservoir area.
On July 19, 1973, 42 male and 59 female goslings were released on Jumbo
Reservoir Annex, bringing the total number of geese released there in 3 years
to 303. Generally, the birds remained in the vicinity of the Annex throughout the summer and into the early fall.

�-50-

South Platte Valley-Prewitt

Reservoir

In July 1972, 70 Canada goose goslings were released on Prewitt Reservoir,
located near the South Platte River in Washington County, in northeast Colorado.
In the spring 1973, a small flock of birds were present periodically
and 1 nest was reported near Prewitt.
The nest was established in a marsh
area located below the dam. The nest was apparently established by a pair
of birds that had either pioneered into the area, or were members of the 19
birds released there in 1970.
On June 25, 1973, 39 male and 41 female goslings were released at Prewitt.
An additional 5 males and 11 females were released at the same location on
July 19, 1973. The birds remained on the reservoir until early fall when they
moved to nearby South Platte River.
North Park
By August 1972, a total of 831 Canada goose goslings had been released during
a 4 year period at 2 locations in North Park. These birds had established
the lower Colorado River bordering California and Arizona and the Imperial
Valley of California as their winter area (Szymczak 1973). Some of the
released birds were using Wheatland Reservoir in Albany County, Wyoming as a
molting area (Szymczak 1973). Nesting attempts began in North Park in 1971
and increased in 1972, but comparatively large numbers of birds had not
returned to North Park (Szymczak 1973).
Spring Returns
On March 28, 1973, 25 geese were observed at Lake John Annex.
On April 6,
1973 approximately 60 geese were observed on the Annex. About 15 birds were
grouped and feeding on grain placed in the artifical feeder .. Mbstof
the
other birds were in pairs and were feeding in the pasture to the south of the
Annex.
Some pairs were closely associated with nesting structures.
At least
2 pairs were observed copulating.
Snow and ice cover was nearly continuous
throughout the area except for a small area of open water on the very southern
part of the Annex and about a 30 acre area of the pasture located directly
south and adjacent to the open water.
No potential natural nest sites were
free of snow.
On April 27, 1973, 30 to 31 geese were observed for a period of time on Lake
John Annex.
Most birds were in pairs.
One nest was observed on an elevated
structure on the southeast corner of the pond. Also on April 27, 1973, 9
geese were observed on the extreme southeast corner of Walden Reservoir, the
only part of the reservoir in which the water was open.
Summer Res iden ts
On May 23 and 24, 1972 a complete survey of all major water areas, with the
exception of the river's systems, was conducted.
The survey was duplicated
June 18, 1973. The number of adult and subadult geese observed on those
surveys by area are presented in Table 1.

�-51-

Table l. Number of full grown Canada geese observed
surveys in North Park, Colorado.

during late spring

No. in Pairs

1972
No. in Groups

Lake John. Annex

4

33

16

59

Walden Reservoir

10

10

0

15

MacFarlane

2

0

0

10

0

0

0

5

6

0

0

0

22

43

16

89

Location

Reservoir

Pole Mountain
Boettcher

Reservoir

Lake

Total

1973
No. in Pairs 1/ No. in Groups

11 Count taken during molt, grouping of birds tends to mask observation

of

pairs.

One hundred five full grown birds were observed during the 1973 survey,
compared to 65 in 1972
(Table 1). IIi 1971 the largest group observed during
that period was 50 (Szymczak 1972).
Nesting and Production
Thorough nest searches have not been conducted in North Park, mainly because
of the disturbance factor.
Information on nesting and production has been
gathered through long distance observation, chance encounters and counts
of broods and nests during the late spring surveys discussed above.
During 1973, nesting attempts were hampered by an unusually late spring.
There is a possibility that many of the potential nesting birds either did
not attempt to nest or were unsuccessful because of inclement weather.
Known
production information for 1973 is summarized along with previous years data
in Table 2.
Molting
Birds released in North Park continue to be recaptured on Wheatland Reservoir
in Wyoming during banding operations.
In June, 1973 6 birds were captured
at.Wheat1and.
Only one of the 6 had not been captured there before, and
one bird was·recaptured for the fourth consecutive year.

�-52-

Table 2.

Known production

data for Canada geese in North Park 1971-1973.

No. of
Nests Established
1971 1972 1973

Area

Lake John Annex

1

3

6

1

2

4

Boettcher

0

2

0

0

2

0

Arapahoe National
Wildlife Refuge

0

1

1

0

1

1

Total

1

6

7

1

5

5

l/ Actual
goslings

Lake

number unknown,
produced.

New Releases,Summer

Estimated No.
Goslings Produced
1971 1972 1973

No. Hatched
1971 1972 1973

7

16

0

11

0

0

-1/

- 1/

23

20

average brood size used for estimates

Observations

of total

and Early Spring Returns.

On July 9, 1973, 59 males and 41 females were released on Walden Reservoir,
bringing the total number of birds released in North Park to 931. Periodic
observations indicated that the birds remained on Walden Reservoir throughout
the summer and into early fall. Approximately 30 birds were observed on
Walden in late November.
On March 17, 1974, 3 geese were observed flying over Walden Reservoir.
Annual, first early spring observations for Canada geese are summarized
Table 3.

Table 3. First spring observations
and location.

Date

in

for Canada geese in North Park by year

Number Observed

Location

Observed

April 11, 1970

11

Lake John Annex

April

10, 1971

10

Illinois River, west of
Walden

March

12, 1972

16

North Platte River, near
Delaney Buttes

March 28, 1973

25

Lake John Annex

March 17, 1974

3

Walden

Reservoir

�-53-

South Park-Antero

Reservoir

On July 2, 1973, 46 male and 34 female Canada goose goslings were released
on Antero Reservoir in Park County.
Seventy-six birds were released at
that same location in July 1972.
Restoration efforts in South Park are designed to supplement a small nesting
population which is present on Antero (Szymczak 1973).
In the spring of
1973 birds did return to Antero and successfully reproduce.
It is assumed
that those nesting birds were members of the remnant population.

LITERATURE

CITED

Szymczak, M. R. 1972.
Experimental studies on improving status of Canada
goose populations.
Colo. Div. of Wildl., Game Res. Rept., Fed. Aid
Proj. W-88....:R.October.
p. 39-49.
1973.
Experimental studies on improving status of Canada goose
populations.
Colo. Div. of Wildl., Game Res. Rept., Fed. Aid Proj.
W-88-R.
October.
In Press.

Prepared

J!i'~i

by . ·~/-!····1/tJlJ:J:~'·Co·:=::·~. _.-&lt;Jf?t:..... !..!.•.••.•..
/..:l.~ A~.f-· sx:-~
~.~:2..Michael R. ~~
Assistant Wildlife Researcher
/1

��October 1974
-55-

JOB FINAL REPORT

,State of

---.;C:.;;;O..;;;LO=ru:.;;;AD;:;.-O~
_'___

Project No.

W-88-R-19

Work Plan No.
Job Title

Job NO._, __

~~5~

- __ ~~_

Arkansas Valley Canada Goose Flock Management Studies

Period Covered:
Personnel:

2

,Migratory Bird Investigations

April 1, 1966 through March 31, 1973

Ann Leckler, D. Bogart, J. Carsel1a, V. Clark, G. Eyre, R. Forbes,
B. Goetze, L. Green, J. Grieb, A. Heins, D. Hopper, L. Johnson,
R. Kitzmiller, C. Leonard,D. Minnich, H. Moorhead, G. Nugent,
P. Olson, D. Potts, R. Rosette, F. Roth, W. Russell, W. Rutherford,
R. Velarde, B. Widhalm, and M. Szymczak.
ABSTR.~CT

The earliest arrival date of Canada geese (Branta canadensis) in southeast
Colorado during the 1966 through 1971 period was November 3, 1967 with the
latest being December 10, 1962. During the last 10 years there has
been a general increase in the number of birds staying at Turk's Pond during
the winter, with a decline in the number of birds staying at Two Buttes
Reservoir and at the Eads Lakes. The number of birds staying at Meredith and
John Martin Reservoirs have remained fairly stable. In January 1974 the total
short grass prairie population numbered 149,270 birds, approximately eight
percent below the population size objective for that year. Special regulations
designed to retard movement of geese out of southeast Colorado into the Texas
Panhandle had no effect through 1968. Various one-half day hunting regulations,
first initiated in 1969 definitely were effective in retaining geese in southeast Colorado Under most conditions. Feeding flight activity during various
half day regulations is discussed. During the seven year period of this study
the percent of young in the harvest on firing lines varied from 59 percent in
1971-72 to 7.5 percent in 1972-73. On decoy hunting areas the portion of
young in 'the harvest was on the average, 19 percent higher than on firing line
areas. During the 1970-71 season hunters on the Two Buttes Management Area
harvested 1,733 geese in 3,170 hunter days for an average of 0.55; the latter
figure a Two Buttes Management Area record. Seventy-eight percent of the
geese harvested on the Two Buttes Management Area were taken on the south side
of the reservoir during the 1966-67 through 1968-69 periods. During the 196970 season a record 512 geese were harvested in 1,361 hunter days, for an
average bag per hunter day of 0.38. Harvest in all of southeast Colorado reached
a record 30,329 during the 1970-71 season. Baca County was the major county
of harvest in five of the seven seasons from 1966-67 through 1972-73. Males
composed 50.3 percent of the 5,748 birds trapped during post-season banding
operations from 1966-67 through 1972-73. On the basis of weight loss during
the season, geese mainta'ined better body condition during the various halfday hunting regulations than during season long full day hunting regulations.
Beginning in 1966, Saskatchewan replaced Alberta as the major recovery locations of Canada geese banded in southeast Colorado and taken on the migrational
staging area. Harvest on the migrational staging area in Alberta and Saskatchewan of birds banded post-season in southeast Colorado has accounted for an
average of 39 percent of the annual band recoveries since the 1966-67 hunting

�-56-

ABSTRACT
(Continued)

season compared to 44 percent prior to 1966-67. Harvest in Colorado has
accounted for an average of 33 percent of the annual band recoveries since
1966-67 compared to 29 percent prior to that period.
Mortality rates for
geese banded post-season in southeast Colorado calculated by the composite
dynamic method are .257, .263, and.260 for adults, juveniles and all age
classes combined, respectively.

RECOMMENDATIONS
1.

At least two inventories of Canada geese in southeast Colorado should. be
conducted each year.
These surveys, one in early December and one in
early January should be coordinated with other agencies in Nebraska,
Texas and NewMexico so that as complete a count as possible of the
short grass prairie popuLatLon will be obtained.

2.

Experimentation with management practices and season regulations should
be continued in attempts to obtain an acceptable distribution
of geese
both within southeast. Colorado and the entire short grass prairie wintering range.

3.

In order to enhance the possibility
of successful public hunting at Two
Buttes Reservoir, ari attempt should be made to gain control of the land
adjacent to the south side of the reservoir.

4.

A quota of Canada geese should be banded post-season in southeaSt Colorado.
Currently, the quota shoul.d iba 1,000 birds of any age or sex
banded at any location,
or number of locations.
The age' of. each bird
should be determined by the notched tail feather method, and the sex
determined by cloacal examination.
Bands should be placed only on birds
of known age and sex.
Continued banding is essential
to monitor the
mortality and harvest distribution
of the population.
Annualanalysis
of recoveries should be a: cooperative effort between the management staff
responsible for the banding and the migratory bird research staff.

5.

A representative
of the management staff of the Colorado Division o f
Wildlife should function as a memberof the Short Grass Prairie Canada
Goose Subcomridttee of the Central Flyway Technical Commi.t
t ee ,
The
representative
should be responsible forgathering
allpertinentinformation concerning the Colorado segment of the short grass prairie goose
population and presenting the information to other members of the sub~
comridttee. The representative
will participate
in all subcommittee
discussions and decisions concerning any aspect of the short grass prairie
Canada goose flock.

�-57-

ARKANSAS

VALLEY GOOSE FLOCK MANAGEMENT
Michael

STUDIES

R. Szymczak

Arkansas Valley goose flock management studies evolved from earlier investigative programs conducted and reported by Grieb (1970) and Rutherford (1970).
This study was designed to carryon
various management programs initiated
during the two previous studies.
The basic objective was to collect information which would enable an annual evaluation of the status of the Arkansas
Valley segment of the short-grass prairie Canada goose population.
In addition,
during the course of the study, new approaches to the management of Canada
geese in the Arkansas Valley were evaluated.
The termination of this project
in no way indicates that Colorado will discontinue the collection of vital
management information.
The collection of these data will continue as a
function of the management staff of the Colorado Division of Wildlife.

P. S. OBJECTIVE
To collect management
goose population.

information

on the wintering

short-grass

prairie

Canada

METHODS AND MATERIALS
Periodic aerial counts were conducted in the Arkansas Valley throughout the
study. At least two inventories each year were coordinated with other agencies
resulting in essentially simultaneous counts throughout the short-grass
prairie wintering range in Nebraska, Colorado, Oklahoma, Texas and New Mexico.
Tail fans were collected annually at check stations at the Two Buttes and
Lamar-Eads Management areas in order to obtain some information on the age
composition of the flock. Occasionally tail fans were obtained through hunter
bag checks at other areas throughout the Arkansas Valley.
The age of each
Canada goose banded was recorded.
Family group counts were conducted during
the first year of the study.
Records concerning hunting pressure and harvest were collected at the Two
Buttes and Lamar-Eads Management areas. Estimates of harvest for all of
southeast Colorado were obtained from the annual small game hunter harvest
survey.
During a portion of the study, Canada goose feeding flights were observed at
various locations throughout southeast Colorado.
Information on time and size
of flights and the general behavior of the birds was recorded.
Canada geese were trapped and banded in southeast Colorado-post season throughout the length of the study. The sex, age and weight of each goose banded
was determined.
Morphological measurements of geese captured were taken during
two post-season trapping periods.

�-58-

RESULTS AND DISCUSSION
According to Grieb (1970) the time of arrival of geese on the wintering
ground in southeast Colorado is dependent on a major storm appearing on the
staging area in Alberta and Saskatchewan. In response to the storm, the
birds generally move south en masse to their wintering areas. From 1960
through 1971 the earliest arrival date in southeast Colorado was November 3,
and the latest December 10 (Table 1).

Table 1.

Arrival dates of Canada geese in southeast Colorado, 1960-71. 1/

Year

Date of Arrival

1960

November 8

1961

November 5

1962

December 10

1963

November 22

1964

November 20

1965

November 12

1966

November 10

1967

November 3

1968

November 17

1969

November 16

1970

November 17

1971

November 10

1/ Dates for 1960 through 1965 are from Grieb 1970.
In the fall of 1970 the chronology of arrival of geese on the wintering
ground in southeast Colorado was definitely atypical. Some birds began
arriving as early as October 15. About October 30 a substantial movement
into southeast Colorado occurred, resulting in a total of about 25,000 on
November 9. About November 17 the major portion of the migrant birds moved
into the area with the aerial survey on November 23 recording approximately
90,000 birds. Apparently the early season movement south by a portion of the
population was the result of an early cold period on the Alberta-Saskatchewan

�-59-

staging areas. Normally, the first cold period on the staging area moves
birds from the lakes and reservoirs to the rivers (A. Dzubin, personal
communication).
Distribution of the Wintering Population
Rutherford (1970) discussed the combined influence of weather, water, food
conditions and to some degree, hunting pressure on the distribution of the
wintering population of Canada geese in southeast Colorado. From the 1966
through 1972 period, no fewer than three and no more than seven aerial flights
were conducted to determine the population size and distribution of these
birds.
In order to obtain an index that is comparative between areas an average
number of birds present throughout a particular season, using all aerial
counts, was calculated for an area and presented as a percent of the total
birds in southeast Colorado (Table 2). This method results in an average look
at anyone season.
Over the ten year period each area underwent some drastic fluctuations, probably in response to variations in conditions discussed by Rutherford (1970).
On a long term basis, as partially indicated by five year average presented
in Table 2, there is a general increase in the percent of birds at Turk's
Pond and declines at the Eads Lake and Two Buttes. Both the Meredith area
and John Martin seems to have remained fairly stable.
During the seven year period of this study, various changes in regulations
were instigated in attempts to alter the distribution of geese in southeast
Colorado. In 1966, all areas south of the Arkansas River Valley proper had
a later opening hunting date than the remainder of the Arkansas Valley. The
same regulation had been in effect in 1965 (Rutherford 1966). In 1967, 1968
and 1969 the Two Buttes Reservoir area only, was subject to the delay. In
addition, in 1967 and 1968, the north and south sides of the reservoir were
open and closed to hunting alternately, at weekly intervals. Regulations
during the 1965 and 1966 seasons were designed primarily to encourage birds
to remain in southeast Colorado in general and in Baca County specifically~
for the entire hunting season. Regulations in 1967, 1968 and 1969 were
designed not only to achieve the above goal but also to maintain birds at
Two Buttes Reservoir because most of them had been gathering and remaining at
Turk's Pond. The effects of these types of regulations on distribution in
southeast Colorado can only be described as erratic. Regulations at Two Buttes
seemed to achieve the objective in 1967, but not in 1968 or 1969 (Tables 2 and
3) •

In 1972 a full day season was in effect in the Two Buttes-Turk's area while
only half-day hunting was allowed in the Arkansas Valley proper. This regulation was designed to maintain more birds in the more northerly Meredith,
John Martin, Eads Lakes areas, and hopefully, delaying mass movement to the
Two Buttes-Turk's area at least until later in the year; an objective which
was not met.
In the final analysis it seems that no regulation applied to date, designed
to influence the distribution of geese within southeast Colorado has been
conSistently effective.

�-60-

Table 2. Percentage distribution of geese located on selected wintering
areas in southeast Colorado, pooling all census results by year.

1/
Year--

Meredith

John Martin

Area
Eads Lakes

Two Buttes

Turk's

1963

9.0

14.3

31.5

41.1

1.5

1964

0.7

2.7

47.4

31.5

17.6

1965

5.6

11.2

20.3

30.1

24.0

1966

11.4

19.0

32.1

7.3

18.4

1967

6.9

16.1

24.0

33.0

16.4

1968

4.5

16.2

28.5

15.7

32.4

1969

6.2

12.2

35.3

16.1

24.6

1970

10.7

18.2

16.2

32.9

20.6

1971

5.3

12.5

22.2

18.8

35.3

1972

3.3

15.7

13.2

24.6

42.5

Totals
1963-67

6.7

13.1

27.8

28.6

17.6

Totals
1968-72

6.4

14.4

23.4

21.6

30.2

1/ Years 1963 through 1965 from Rutherford 1970.

�Table

3,

Year

Ae r Lal,

Date

counts

Meredith
Reserv.

0f

Canada geese,

southeast

Meredith

Lake Area

Holbrook
Reserv.

Dye
Reserv.

1966··67

11/15
12/1
1/2

4,150
6,700
1,475

0
400
200

0
350
175

1967-68

11/17
11/28
121'1
1/1l!il

1,850
5,275
2,550
1,400

0
0
165
0

0
50
500
450

11/20
12/5
12/17
1/2
1/16

3,810
1,075
600
600
890

0
0
IS
0
0

1969-70

11/21
12/9
12/19
1/7
1/16

7,600
3,900
1,075
2,650
845

0
100
400
600
1,200

1970-71

11n3
12/11
12/23
11M

9,700
9,350
10,7201/
4,695-

0
0
150
0

1971-72

IDU1l3
lIf9
11/22
12/6
12/13
12/21
1

3,400
13,000
1,300
5,200
3,000
250
1,500

1968-69

't..

1972-73

lUlllO
1l/28
Ub12
l/Z

3,500
1,100
500
200

1/ Est Lmat.eefrom ground.

0
0
30
0
200

Others

Colorado,

1966-1972.
John

Blue Lake Area

Ho r se c re ek

Blue

Rese rv .

Lake

5,000
1,200
0

Other

1,800
900
0

John
Mart in
Re s er v.

8,300
5,000
3,500

".'e rh oe f f
Pond

0
25
2,000

Martin
I

Ead s Lakes Ar~a

Area

S

Ot he r

Sweetwater
Reserv.

Nee
Noshe

Upper
Queens

flamingo

Swede
Lake

Other

Two
Buttes

2,200
1,500
275

3,300
75
1,0

13,300
9,000
9,125

I, lOO
25
2,000

0
0
0
150

0
900
1,800
3,200

15,500
9,000
1,800
3,000

450
950

Specific

-

0
4,500
0

0,
12

n

J.50

0
900
40
1,375
250 i,iS5
o 3,500

150
0
1,500
500
3,200

80
175
450
0
100 1,100
0
700
125
120

Turk IS
Pond

Bonny

nese rv.

Other

Totals
--

8,000
500
130

2,500
6,800
12,500

0
150
305

49,650
37,137
31,835

13,500
15,500
15,000
11
6, 80CP

5,000,
5,000
8,8001/
l1,50CP

0
0
0
32

44,500
55,580
41,500
42,682

11,500
600
500
600
11,000

9,500
10,000
15,000
11,000
4,500

0
108
1,777
631
19

45,000
24,833
26,592
24,5::n
33,366

0
0
5
394
25

56,846
60,700
55,980
63,444
64,759

0
25
450
0

700
0
0
0

1,100
'1,790
300
0

5,500
15,675
6,000
2,500

0
0
0
0
12

450
0
0
0
0

350
400
300
0
0

100 12,300
0 4,000
0
700
0 3,500
0 4,500

185
3,700
0
0
0

6,400
4,500
5,000
7,000
9,000

0
0
20
0
0

5,000
1,700
0
0
4

2,200
4,000
0
0
0

4,000
400
16,000
12,000

6,800
1,900
0
0
55

3,500
7,54';
19,OOO 4,600
12,000
6,500
9,000
4,000
In,OOO
11,000

0
1,900
400
0
0

201
1,200
1,000
0
0

0
0
50
0
0

12,000
9,000
4.,500
6,000
17,OnO

7,000
12,000
13,500
27,800
14,000

1,000
1,000
500
1,000
0

500
0
0
0

0
0
9

29,00011
24,60CP
20,400
32,200

12,0001,
21,20CP'
20,000
13,400

I,OOO!!
0
1,7001/
1,00CP

89,400
90,550
80,249
62,720

600
3,800
600
0
0

100
6,000
0
7,500
7,000
300
0

2,000
5,600
8,000
15,000
22 ,SOD
2,,200
9,500

11,000
14,000
16,400
6,700
18,500
25,000
6,000

8,300
16,000
17, 300
32,000
25,000
34,000
51,000

600
0
0
600
0
0
300

40,000
80,200
65,050
85,500
85,000
87,750
76,400

400
0
0
0

3,900
13,100
900
5,000

1,300
11,500
10,000
13,000

10,900
10,000
18,000
23,000

0
300
0
0

24,200
47,500
32,650
41,300

°
0
0
300
0
0

a

4,430
350
23,700
0
10,000
14,750 / 1,100
9,OOol0

2,/00

220
0
120
300

5,300
11,600
3,800
600

330
400
5,700
1,500

7,300
9,000
1,700
0

0
0
3,0001,400
100
0
25
0

0
0
1,400
3,000
0
0
0

0
15,000
12,000
6,000
5,000
12,000
0

10,600
2,700
50
i , ';00
4,000
14,000
8,100

0
4,100
8,000
8,000
0
0
0

3,400
0
0
0
0
0
0

2,000
11,000
3,000
0

2,200
200
700
100

-

,

-

'",

�-62-

Size of the Wintering

Population

Total Population
Grieb (1970) stated that a basic management objective for the short grass
prairie Canada goose population should be an increase in numbers at the rate
of 5 percent per year as long as restraints on the population are not warranted.
When this study was initiated in 1966 the population was estimated
to be about 110,000 birds (Table 4). According to Grieb's suggested rate of
increase, the population should have numbered 155,000 by January, 1973. unfortunately, counts of short grass prairie Canada geese in January, 1973 were
not reliable.
The January, 1974 count, taken after the termination of data
collection under this project, totaled 149,270, or about 8 percent below the
projected population figure of 162,000 for January, 1974.

Table 4. Post-hunting season status of Short Grass Prairie Canada goose
populations, 1960-73, data generally from regular mid-winter inventories.

Year

Number of Birds

Year

Number of Birds

1960

77,709

1967

111,452

1961

103,355

1968

127,903

1962

80,133

1969

112,399

1963

93,940

1970

147,414

1964

81,221

1971

152,734

1965

103,435

1972

134,500

1966

110,485 1./

1973

257,500 ~/

11 Inven tory of February 15, 1966 substituted
inventory.

Y Unreliable

for unsatisfactory

January

coun t.

A definite problem in obtaining a good reliable count has arisen in recent
years.
Birds from the short grass prairie nesting popUlation are beginning
to winter in the Hi-Line population wintering range in north-central Colorado
and southeastern Wyoming (Szymczak In press).
To date, the portion of the
popUlation wintering in the Hi-Line area has not been isolated in terms of
numbers, and therefore total short grass breeding population numbers have not
been obtained.
Numbers of short grass birds Wintering in the Hi-Line area
seem to have been erratic from year to year, but have probably not exceeded
10,000 birds.

�-63-

Southeast Colorado Population
Earlier in this report a discussion concerning the distribution of Canada
geese in southeast Colorado touched on the implementation of various regulations designed to maintain a huntable population of Canada geese in southeast
Colorado throughout the hunting season. For five years, beginning in 1964,
delayed opening type regulations designed to allow geese to establish feeding
patterns prior to the opening of the hunting season were in effect. In 1967
and 1968 the season opening date on the Two Buttes Wildlife area was delayed
even further. In 1968, an evaluation of the five years of expe riment ation
indicated that the regulations had not been effective in retarding the movement of geese into the Texas Panhandle, as was the original objective. Colorado was retaining a decreasing percent of an increasing short grass prairie
popuiation (Fig. 1).
In 1969, Colorado initiated half-day hunting regulations in order to meet the
wintering grounds numbers objective of at least 40,000 birds throughout the
hunting season in southeast Col.orado (Rutherford 1970). The regulations from
1969 through 1972 were as follows:
1969 - November 22, 1969 through January 15, 1970; one-half hour before sunrise to noon.
1970 - October 31, 1970 through December 11, 1970; one-half hour before sunrise to noon.
December 12,1970
rise to sunset.

through January 17,1971;

one-half hour before sun-

1971.- November 1, 1971 through December 10, 1971; one-half hour before sunrise to noon.
December 11, 1971 through January 16, 1972; one-half hour before sunrise to sunset.
1972 - For all areas except Baca County (Two Buttes-Turk's Area) October 28,
1972 through December 1, 1972; one-half hour before sunrise to noon.
December 2, 1972 through January 14, 1973; one-half hour before sunrise to sunset.
Baca County - October 28, 1972 through January 14, 1973; one-half hour before
sunrise to sunset.
The popUlation seemed to respond immediately to the regulation. In 1969 counts
in southeast Colorado totaled over 55,000 in late November and remained high
for the entire winter (Table 3). In 1970, inventories in southeast Colorado
averaged nearly 81,000 birds for the entire season. In 1971, Colorado maintained the highest sustained percentage of the population having 80,000 birds
in the state in early November, and a record 76,000 or about 57 percent of
the total population duritig the January 1972 inventory (Tables 3 and 5, Fig. 2).

�150,000
JANUARY INVENTORY

125,000

&lt;,
100,000
~
~
~
(.;
CI.l

~

0
p::

~

.....,

~
75,000

I

0\
.j::'-

J

I

~.

50,000

'"

...

/

'"

""

/

..•. ..•

.•.. ,Colorado Population
..•..•.-

--

--- -

/

---- - - - - - •...••.

/

, ...•

...•..'" ,-

/

..•.•

/

'"

.•..•

....

....
&lt;,
&lt;,

25,000

1961

1962

.------------------,------------------,~----------------,_----------------~~----------------_r------------------r----------

1963

1964

1965

1966

1967

1968

1969

YEAR
Fig. 1. Comparison of the Colorado and Texas Panhandle segments of the Short Grass Prairie Canada Goose
Populations, January 1961 through January 1969.

�-65-

JANUARY INVENTORY

150,000

125,000

~
~
~

.00

100,000

C-'
~

o
~
~
~
~

75,000

I

/

50,000
Colorado Population

••...

/

I

I

/
/

25,000

1967

1968

1969

1970

1971

1972

YEAR
Fig. 2. Comparison of southeast Colorado's population with total
Short Grass Prairie Population according to the coordinated
January inventory.

�-66-

However, Colorado populations during the 1972 hunting season fell to prehalf day hunting season levels with flocks generally equally reduced in all
areas. Still the large concentration of birds during the 1972 season was
in the Two Buttes-Turk' s Pond area, the area which was subject to full day
hunting.

Table 5.
1948-74.

January inventory of Canada geese, Arkansas Valley, Colorado,

Year

Goose Count

Year

Goose Count

Year

Goose Count

1948

4,798

1957

24,617

1966

38,635 '1:../

1949

12,286

1958

35,894

1967

29,835

1950

13,170

1959

44,660

1968

42,682

1951

19,320

1960

37,394

1969

29,201

1952

30,463

1961

31,360

1970

63,444

1953

20,236

1962

40,250 1/

1971

62,720

1954

20,280

1963

35,889

1972

76,400

1955

25,110

1964

33,750

1973

41,300

1956

24,212

1965

37,693

1974

27,400

1/ Inventory of February 7, 1962 substituted for January, 1962 inventory.

l/ Inventory of February 15, 1966 substituted for January, 1966 inventory.
Four years of experimentation with half-day hunting regulations have shown
that birds can be retained in southeast Colorado under most conditions using
the half-day hunting technique. Apparently under existing conditions, the
Two Buttes-Turk's complex in Baca County is the critical area. Therefore,
if comparatively large numbers of birds are to remain in southeast Colorado
throughout the season, more than likely the majority of those birds will be
in Baca County.
Effects of Half-day Hunting Regulations on Flight Patterns
During the 1969-70 and 1970-71 hunting seasons the feeding flight patterns
of Canada geese were observed in order to document the possible influence of
the regulation on flight patterns. As stated before, during the 1969-70

�-67-

season hunting was allowed only from sunrise until 12:00 o'clock noon.
In
1970-71, half-day hunting was in effect only through December 11, 1970,
followed by sunrise to sunset hunting for the remainder of the season.
Because of variations in ecological factors influencing patterns as discussed
by Rutherford (1970), each season was considered separately.
The following
are general conclusions resulting from the observations.

Season - 1969-70
1.

Prior to the beginning of the hunting season geese made feeding flights
twice each day, one in the morning and a second in the afternoon in spite
of approaching full moon conditions.

2.

In the Eads Lakes, John Martin, Meredith Lake area birds began feeding
exclusively in the late afternoon and through the night on November 23,
the day that the season opened and the height of the full moon period.
In the Two Buttes area birds continued to fly twice each day after November 23, but afternoon flights gradually began later in the day and night
feeding was common during the November 22 through November 28 period.
Hunting pressure was heavy in the Eads Lakes area and light in the Two
But tes area.

3.

In general, the cessation of morning flights was closely correlated to
light of the moon phases during the hunting season in the months of
November and December.
Rutherford (1970) found that geese in southeast
Colorado fed exclusively at night for about 10 days each month during the
full moon phase.

4.

Geese subject to firing line hunting around reservoirs showed a more
definite response to light-of-moon phases by discontinuing morning flights,
than under non-firing line situations.

5.

Inclement weather stimulated morning flights even during light of the
moon periods.
Cold weather retarded morning flights even during dark of
the moon pe riods -.

6.

There were definite indications toward the end of the season, that geese
responded to morning hunting pressure by delaying feeding flights tintil
afternoon.

Season - 1970-71
1.

Generally, two a day flights occurred from the time geese arrived in
southeast Colorado throughout the entire half day hunting portion of the
season, with the November full moon having very little effect on the
flight patterns.

2.

For about a nine day period December 11 through 19 with the height of the
full moon period on December 12, essentially no morning flights occurred.

�-68-

3.

Morning flights resumed on December 20, but were not of significant
size for the remainder of the season.

4.

Afternoon flights occurred from December 12 through the remainder of the
season.
Full moon periods did not stimulate exclusive night feedings.
However, on some days birds would leave reservoirs in late afternoon and
not return until after dark.

5.

Again,

as during

the 1969-70 season cold weather

retarded morning

flights.

Age Composition
Grieb (1970) and Rutherford (1970) both discussed the value of obtaining
population age-ratio information on migration and wintering areas as an
indirect indication of production.
Each agreed that age ratios obtained using
birds harvested by hunters tended to over estimate the percentage of young in
the population because young are taken at a faster rate than adults, as
pointed out by Rutherford (1968).
Grieb (1970) discussed the wide variety of opinion relative to the validity
of cannon-net trapping in obtaining Canada goose age ratios. He concluded
that age-ratios obtained from cannon-net trapping in southeast Colorado
"could well be representative of the age ratio of birds in the vicinity of
the trap site", and therefore, proper sampling with the cannon-net would provide reliable estimates.
Trapping techniques during this study followed no specific sampling technique.
Other factors such as the mere presence and accessibility of a trap site and
its physiographic features were of prime importance.
Many times, catches
were made at the same site throughout the trapping season.
Therefore, for
the population to be properly sampled, the geese had to be distributed somewhat randomly.
In analysis of age data collected during the period of this study the assumptions relative to obtaining reliable estimates were not made.
Age data collected were considered only as a possilbe gross indicator of production and no
management decisions were made based on age-ratio data.
Age ratio information for both hunter harvested birds and trapped birds for
this study are presented in detail in the preceding annual progress reports
for this job. For the purpose of this report, annual summaries are presented
in Table 6.
Rutherford (1970) found that through 1967-68 the percent of young in the sample
harvested by hunters was higher than the percent in the trapped sample in nine
out of eleven years.
From the 1968-69 season through the 1972-73 season, only
one season, 1972-73, -showed a lower percent of immatures in the harvested
birds as compared to the trapped birds (Table 6). The harvested sample for
1972-73 numbered only 67 birds.

�Table 6.

Age composition of the short grass prairie Canada goose population in southeastern Colorado.

Banding
Year

No.

Adult
Percent

Young
No.
Percent

1950-51
1951-52
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69
1969-70
1970-71
1971-72
1972-73

344
649
346
214
269
1,171
671
607
360
789

53.3
50.9
66.4
42.2
60.0
62.3
77.7
44.7
57.7
80.6

300
627
175
293
179
710
192
750
264
189

46. 7
49.1
33.6
57.8
40.0
37.7
22.3
55.3
42.3
19.4

438
700
928
379
549
703
496

59.3
70.6
83.2
61.1
64.1
67.2
87.9

300
292
187
241
307
343
68

40.7
29.4
16.8
38.9
35.9
32.8
12.1

Check Station
Firing Line
Decoy
Adult
Young
Adult
Young
No.
Percent
No.
Percent
No.
Percent
No.
Percent

356

46.8

404

53.2

929
377
612
527
204
377
442
571
217
245
136
110
672
867
62

47.9 1,010
36.5
655
45.5
732
55.7
419
48.9
213
52.1
346
63.1
259
340
62.7"
45.3
262
59.9
164
61.8
84
55.3
89
40.9
972
62.4
522
92.5
5

52.1
63.5
54.5
44.3
51.1
47.9
36.9
37.3
54.7
40.1
38.2
44.7
59.1
37.6
7.5

t

0'

\0
t

79
143
77
128
102

51.3
35.9
27.4
33.7
71.3

75
255
204
252
41

48.7
64.1
72.6
66.3
28.7

�-70-

The establishment of a decoy hunting only, area in southeast Colorado in
1968 provided an opportunity to compare the age-ratio of birds harvested
over decoys with those harvested on the Two Buttes firing line. For the five
year period the percent of young in the decoy harvest exceeded the young in
the firing line harvest by an average of 19 percent. The differences ranged t
from about 11 to 29 percent. The percent of young in the banded sample from
1966-67 through 1972-73 was between 30 and 40 percent for five of the seven
years (Table 6).
In 1968-69 and 1972-73 the percent young was comparatively
very low, 16.8 and 12.1 percent, respectively. According to post-season
population counts, the low number of young in 1968-69 apparently did indicate
a reduced population (Table 4). But no such direct correlation was present
in 1972-73, nor had it been in 1964-65 when the percent of young in the banded
sample fell below 20 percent.

Hun ting Pressure and Harves t
Two Buttes Management Area
Estimates of hunting pressure and harvest on the Two Buttes area were collected
annually through the 1972-73 hunting season. Collection of information at
Two Buttes began in 1957. During the 1958-59 hunting season, a record 1,947
geese were harvested by individuals hunting a record 7,343 days, for a .27
bird per hunter day average.
The most successful hunting year on record at Two Buttes was the 1970-71 season
when 1,398 hunters hunted a total of 3,170 days and bagged 1,733 geese for an
average bag per hunter day of 0.55 (Table 7). Regulations during the 1970-71
season permitted sunrise until noon hunting from October 31 until December 11,
1970, followed by sunrise to sunset hunting until January 17, 1971. The average bag per hunter day during half-day hunting was 0.66, compared to 0.46
during full day hunting. Counts at Two Buttes during the 1970-71 season indicated the population present varied from 20,000 to 32,000 (Table 3). During
the 1971-72 season the regulations were essentially the same, the population
was usually fairly substantial, but did fluctuate, but hunting success on
the management area declined primarily because of the loss of the land on the
south side of the reservoir to public hunting.
Rutherford (1970) pointed out that maintaining optimum numbers of geese at
Two Buttes Reservoir was essential in order to provide the greatest amount
of public recreation at Two Buttes Management· area. During a six year period
beginning in 1963, the harvest on the Two Buttes Management area was recorded
by pit number, and whether it occurred on the north or south side of the reservoir. From the 1963-64 through the 1965-66 seasons, 60 percent of the geese
harvested on the management area were taken on the south side (Rutherford 1964,
1965, 1966).
From the 1966-67 through the 1968-69 period, 78 percent of the
birds were harvested on the south side. In addition to the distribution of
harvest information, observations of goose feeding flights have indicated a
general increase in flights south, both leaving and returning to Two Buttes.
Daily exchanges of geese between Two Buttes and Turk's Pond, which is located
southeast of Two Buttes Reservoir, were mentioned by Rutherford (1970).
The

�-71-

loss of the south side of Two Buttes Reservoir for public hunting was extremely
detrimental to public goose hunting at Two Butt.es. Not only must optimmn
numbers of birds be maintained at Two Buttes, but given current prevailing
flight pat terns, the south side of the reservoir must be available for public
use in order to provide the greatest amount of public recreation at the Two
Buttes Management area.
Table 7. Goose harves t, hunting pressure, and hunter success, Two Buttes
Managemen tAre a.

Year

No. of
Individual
Hunters

No. of
Hunter
Days

No. of
Geese
Bagged

1966-67

1,257

2,413

1967-68

840

1968-69

Ave. Bag
per
Hunter

Ave. Bag
per
Hunter Day

493

0.36

0.20

1,554

433

0.52

0.28

792

1,309

243

0.31

0.19

1969-70

659

1,182

248

0.38

0.21

1970-71

1,398

3,170

1,733

1.24

0.55

1971-72

830

134

0.16

1972-73

1,003

90

0.09

Lamar-Eads Management Area
Prior to the 1968-69 goose hunting season, the Lamar-Eads Management area was
established near the Eads Lakes complex north of Lamar. Regulations on the
management area permitted decoy hunting only.
A record of hunting pressure and success was maintained on the management area
through the 1972-73 hunting season. During that five year period, hunting
success was highest during the 1969-70 season when 512 geese were harvested by
703 hunters at a rate of 0.38 geese per hunter day (Table 8).
Southeast Colorado
An estimate of hunting pressure and harvest for all of southeast Colorado was
obtained annually from the regular state-wide small game random harvest survey.
Both Grieb (1970) and Rutherford (1970) mentioned problems in obtaining
harvest estimates through this type of a survey. Rutherford (1970) indicated
the survey probably provided a good estimate of the numbers of hunters participating, but cited Grieb's (1970) comment that the harvest estimates were probably

�-72-

inflated.
In final analysis, Grieb (1970) surmised that since the statewide
survey was conducted in the same manner each year it was of year to year trend
value.

Table 8. Goose harvest,
Man agemen t Are a.

hunting

pressure,

and hunter success,

Lamar-Eads

Year

No. of
Individual
Hunters

No. of
Hunter
Days

No. of
Geese
Bagged

1968-69

598

1,062

160

1969-70

703

1,361

512

0.73

0.38

1970-71

705

1,128

304

0.43

0.27

1971-72

1,378

439

0.32

1972-73

814

88

0.11

Ave. Bag
per
Hunter

Ave. Bag
per
Hunter Day

0.15

Goose harvest in southeast Colorado fluctuated considerably during the period
of this study.
The lowest harvest occurred in 1969 when the fewest number of
hunters recorded since the survey began in 1954 harvested an estimated 10,597
birds (Table 9). Reduced hunting pressure in 1969 was probably a result of
the negative reaction of goose hunters to the morning hunting only, season.
According to aerial counts between 55,000 and 65,000 geese were in southeast
Colorado throughout the 1969-70 season.
Harvest in the 1970 season reached a record 30,329 (Table 9). The 1971 season
harvest was nearly as high, totaling 29,114. Regulations during both of
these seasons were identical, with morning hunting only, during the first full
week in December, followed by full day hunting for the remainder of the season.
The population in southeast Colorado during both .those years varied from 65,000
to 90,000 throughout most of the hunting period.
Estimated harvest during the 1972-73 season was less than one-half that recorded
for the previous season.
The harvest paralleled the decline in population
numbers in southeast Colorado (Table 3).
Baca County was the dominant harvest area for five out of the seven seasons
from 1966 through 1972 (Table 10). During the 1969-70 season, the half-day
hunting regulation discouraged hunter participation, particularly in remote
areas such as Baca County.
During the 1972-73 season, weather conditions in
Baca County made hunting areas virtually inaccessible.
Percent of the total
Arkansas Valley goose population present in Baca County (Two Buttes-Turk's area)
during those two years was 41 percent in 1969-70 and a record 67 percent in
1972-73 (Table 2).

�-73-

Table 9.

.:»
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69
1969-70 2/
1970-71 3/
1971-72 4/
1972-73

""fJ

Goose hunting season statistics, 1954-72.

Dates of Season

Estimated
Goose Hunters

Southeast Colorado
Average
Season Bag

Estimated
Kill

11/1 - 12/30
11/1 - 12/30
11/9 -1/7
11/2 - 11/31
11/17 - 1/15
10/26 - 1/8
10/26 - 1/8
11/10 - 1/8
10/31 - 1/13
11/2 - 1/15
11/2 - 1/15
11/2 - 1/15
11/19 - 1/15
11/18 - 1/14
11/16 - 1/15
11/22 - 1/15
10/31 - 1/17
11/1 - 1/16
10/28 - 1/14

7,071
9,054
9,833
9,113
10,082
8,888
9,838
7,577
6,021
6,668
8,016
6,313
9,357
6,975
6,668
5,327
7,875
9,468
6,573

1.04
1.54
1.05
1.39
1.51
1.61
1.39
1.68
1.58
2.17
2.30
1.52
2.59
2.23
1.66
1.99
3.85
3.07
2.08

7,372
13,904
10,276
12,656
15,205
14,309
13,629
11,724
9,495
14,444
18,474
9,613
24,269
15,558
11,046
10,597
30,329
29,114
13,680

!I Data for 1954-55 through 1965-66 from Grieb 1970.
~/ Morning hunting only throughout the season.

1/ Morning hunting only through December 11, full day hunting for the
remainder of the season.

i/ Morning hunting only through December 10, full day hunting for the
remainder of the season.

1/ Morning hunting only through December 2, full day hunting for the
remainder of the season.

Since 1954 Baca County has been the major harvest area in southeast Colorado
for 13 out of 19 years (Rutherford 1970). During the four seasons from 1961
through 1964, and again in 1969, Kiowa County was the major harvest area
(Rutherford 1970). In 1972-73 Bent County was the major harvest area.

�Table 10.

Goose harvest by County,

County and Reservoirs

southeast

1966-67
Percent
Number

Colorado,

1966-1972.

1967-68
Number
Percent

1968-69
Percent
Number

1969-70
Percent
Number

1970-71
Percent
Number

1971-72
Number
Percent

1972-73
Percent
Number

--Baca
Two Buttes

and Turk's

7,930

32.8

7,144

46.0

4,613

41.8

2,070

19.5

13,055

43.0

13,264

45.6

2,488

11.9

4,836

19.9

3,745

24.1

1,802

16.3

1,946

18.4

2,229

7.3

2,620

9.0

3,630

17.4

Bent
John Martin,
Horsecreek

Blue and

Crowley
751

3.1

1,017

6.5

523

4.7

973

9.2

3,025

10.0

2,305

7.9

784

3.8

30

0.1

0

0.0

0

0.0

71

0.7

0

0.0

52

0.2

75

0.4

6,758

27.8

2,427

15.6

1,318

11. 9

3,786

35.7

7,005

23.1

6,039

20.7

1,567

7.5

120

0.5

23

0.1

252

2.3

88

0.8

120

0.4

880

3.0

298

1.4

1,231

5.1

139

0.9

426

3.9

531

5.0

557

1.8

734

2.5

2,127

10.2

2,463

10.1

901

5.8

1,957

17.7

884

8.3

3,980

13.1

2.686

9.2

2,077

9.9

Pueblo

150

0.5

162

1.0

155

1.4

248

2.3

358

1.2

534

1.8

634

3.0

Total

24,269

Meredith
~fano

Kiowa
Eads and Blue
Las Animas
Otero
Horsecreek,
Holbrook

Dyes and

Two Buttes

and Eads

Pr~

15,558

11,046

10,597

30,329

29,114

13,680

,
~
",

�-75-

Banding Investigations
Sex Composition
The sex of each goose trapped and banded in southeast Colorado since the
winter of 1957-58 has been determined by cloacal examination. The importance
of determining the sex of a bird at time of banding has been questioned by
some investigators. Imber (1968) identified some Canada goose populations
in which mortality rate of males were higher than females. Grieb (1970)
found that males banded at Two Buttes had a four percent higher mortality
rate than females. Grieb (1970) did not consider the difference indicative
of any problem in the popUlation.
From the winter of 1966-67 through the 1972-73 trapping season the sex of
5,748 geese trapped in southeast Colorado was determined. Males dominated
the sample in three of the seven years, including 1971-72 and 1972-73 (Table
11). Over the entire period, the sample consisted of 50.3 percent males.
Assuming that the trapped sample is an indicator of the sex composition of
the population, greater mortality of males has not created an unbalanced
sex ratio.
Table 11. Sex ratios of Canada geese trapped in southeast Colorado 1966-67
through 1972-73.

Males

Females

Year

Number

Percent

Number

1966-67

345

49.2

356

50.8

1967-68

403

49.4

412

50.6

1968-69

572

50.1

571

47.9

1969-70

293

47.3

327

52.7

1970-71

416

48.0

451

52.0

1971-72

569

54.8

469

45.2

1972-73

292

51.8

272

48.2

Total

2,890

50.3

2,858

49.7

Percent

�-76Table 12. Comparison of weights, in pounds, of geese from trap and check
station samples, southeast Colorado, 1966-67 through 1972-73.

Year

Source

No.

Adults
Ave. Wt.

1966-67

Check Station
Two Buttes

217

105

Immatures
No. Ave. Wt.

No.

Total
Ave. Wt.

5.67

262

5.10

479

5.36

311

5.47
6.08

68
217

4.85
5.48

173
528

5.23
5.83

Check Station
Two Buttes

255

5.41

170

5.23

425

5.34

Trapping
Baca County
Northern Areas

595
101

5.36
5.70

220
70

5.15
5.28

815
171

5.31
5.63

Check Station
Two Buttes
Lamar-Eads

134
78

5.32
5.46

79
74

5.00
5.51

213
152

5.20
5.48

Trapping
Baca County
Northern Areas

616
315

4.90
5.41

135
55

4.89
5.39

751
370

4.90
5.41

Check Station
Two Buttes
Lamar-Eads

110
142

5.56
5.73

85
248

5.25
5.41

195
390

5.43
5.53

Trapping
Baca County

379

6.03

241

5.74

620

5.92

Check Station
Two Buttes
Lamar-Eads

673
78

5.47
5.68

949
210

4.92
4.99

1,622
288

5.15
5.18

Trapping
Baca County
Northern Areas

332
215

5.08
6.30

192
128

4.59
5.63

524
343

4.90
6.05

Check Station
Two Buttes
Lamar-Eads

126
126

5.49
5.50

79
254

5.12
4.78

205
380

5.35
5.02

Trapping
Baca County

695

5.49

343

5.37

1,038

5.45

Trapping
Baca County

496

5.72

68

5.53

564

5.70

Trapping
Baca County 11 2
Northern Areas _I
1967-68

1968-69

1969-70

1970-71

1971-72

1972-73

II
- Includes all birds trapped at Two Buttes Reservoir or Turk's Pond
l1Inc1udes all birds trapped at the Eads Lakes, John Martin Reservoir or
Verhoeff's Pond; all located in the Arkansas Valley proper.

�-77-

Weights
Weights of Canada geese harvested on the Two Buttes and Lamar-Eads Management
areas were recorded during this study during all years except 1972-73.
Weights of all birds captured during post-season banding operations were also
recorded.
Weight information collected throughout this study was used primarily as a
possible indicator of body condition by comparing average weights of geese
by periods.
Assuming that annual and seasonal distribution of various subspecies remained fairly constant and percentage composition of the population
by subspecies did not change drastically from year to year, these comparisons
are valid.
Birds trapped and/or banded in the northern areas, Eads Lakes, John Martin,
Verhoeff's, were consistently heavier than those from Baca County areas, Two
Buttes and Turk's (Table 12). The only exception occurred in 1971-72 when
immatures taken on the Lamar-Eads Management area were, on the average,
smaller than those taken on the Two Buttes Management area. Adults from the
two areas in that same year averaged nearly the same weight (Table 12).
Disregarding the weights of trapped birds during the 1970-71 season, it seems
that the difference in weights of birds from the two areas decreased during
the 1969-70 through 1971-72 period, a period in which one-half day hunting
seasons were in effect throughout southeast Colorado.
During the 1970-71
trapping season the birds trapped in the northern area at Verhoeff's Pond
were considered on the average morphologically
larger than most birds in
southeast Colorado.
Converging average weights for the two areas during the
various one-half day season regulations indicates that birds in northern
areas either lost weight or those in Baca County gained, or a combination of
those two occurrences.
Information from northern areas is insufficient to
compare the two three-year periods, 1966-67 through 1968-69, with 1969-70
through 1971-72. But birds trapped in Baca County definitely, on the average,
were heavier during the 1969-70 through 1971-72 period, whereas those harvested were not (Table 13). Assuming the harvested and trapped birds were taken
from the same population, the average weight of the birds increased from the
harvest period to the trap period during the various half-day season regulations.
A weight decrease between the time the birds are harvested and the
time they are banded was always the rule, until half-day regulations began
in 1969-70. Unfortunately, no information is available for Baca County for
1972-73 when full day seasons were re-initiated.
Apparently in terms of
body condition the half-day regulations have definitely been of benefit to
geese in southeast Colorado.
Table 13. Average weights of Canada geese taken on Two Buttes Management area
or trapped in Baca County during the 1966-67 through 1968-69 period, compared
with those harvested or trapped in the same area during the 1969-70 through
1971-72 period.
Harvested

Trapped

Period

Adult

Immature

Adult

Immature

1966-69

5.48

5.13

5.15

5.02

1969-72

5.48

4.96

5.54

5.21

�-78-

Distribution

of Harvest

Rutherford (1970) summarized the distribution of band recoveries of Canada
geese banded in southeast Colorado and reported taken during the 1951-52
through 1966-67 seasons.
During that period, roughly 44 percent of the birds
reported recovered were harvested on the staging area, as defined by Grieb
(1970), of Alberta and Saskatchewan.
Grieb (1970) stated that two thirds
of the harvest on the staging area of birds banded in southeast Colorado
occurred in Alberta.
Grieb's analysis of band recoveries through the 1964-65 season resulted in
splitting the short grass prairie population into two segments, (1) an eastern
segment which are coastal nesters breeding along the southern coast of
Victoria Island and the adjacent Arctic Coast of the District of Mackenzie,
staging primarily in western Saskatchewan and wintering in the Waggoner Ranch
area near Vernon, Texas; and (2) a western segment nesting predominantly in
the forests along the Mackenzie drainage, staging primarily in Alberta and
wintering primarily in southeast Colorado and the Buffalo Lakes area of the
Texas Panhandle.
On the staging area two thirds of the western segment was
located in Alberta, the same distribution as indicated by the band recoveries.
Beginning in 1966 a shift occurred on the short grass staging area in the
recovery distribution of geese banded in southeast Colorado.
In four of the
recovery years from 1966-67 through the 1972-73 seasons, more banded birds
were recovered in Saskatchewan than in Alberta below 53 degrees latitude
(Table 14). Assuming band reporting rates are the same for the two Provinces,
either a population shift has occurred or harvest rates have changed considerably.
One possible explanation is the merging of the eastern and western
segments on the wintering area, allowing eastern as well as western segments
the opportunity to be banded in southeast Colorado.
The status of the eastern
group should probably be explored further.
The-percent of total recoveries on the staging area has declined somewhat.
Since the 1966-67 season, recoveries on the staging area have averaged about
39 percent compared to the 44 percent prior to that period.
The recoveries
in Colorado have increased to an average of about 33 percent during the 196667 through 1972-73 period, compared to 29 percent during the earlier period.
Increased harvest in Colorado under various half day regulations in 1970-71
and 1971-72 are primarily responsible for the shift in recovery distribution
(Tables 9, 14).
The take in other harvest areas during the 1966-67 through 1972-73 period
remained similar to that reported by Rutherford (1970), with Nebraska harvesting about 7 percent and Texas about 8 percent of southeast Colorado
banded birds.

Mortality

Estimates

Rutherford (1970) reviewed the mortality rates of Canada geese banded postseason in southeast Colorado through the 1966-67 recovery year.
Using the
composite dynamic analysis method he found mortality rates of adults and

�-79-

juveniles (birds less than one year old at time of banding) to be .259 and
.271, respectively; and the rate for all age classes combined to be .264.,
The same analysis using recoveries through the 1971-72 season results in
rates of .257, .263, and .260 for adults, juveniles and all age classes,
respectively (Tables 15, 16 and 17).
Average mortality rates for Canada geese banded post-season in southeast
Colorado were also calculated by the relative recovery rate method. Mortality rates for adults, juvenile, and all age classes through the 1971-72
recovery year are .255, .251, and .256, respectively (Tables 18, 19 and 20).
These mortality rates have allowed the population to grow somewhat consistently.

LITERATURE CITED
Grieb, J. R.
Monogr.

1970. The shortgrass prairie Canada goose population.
No. 22. 49 p.

Imber, M. J. 1968.
32 (4):905-920.

Sex ratios in Canada goose populations.

Wildl.

J. Wildl. Mgmt.

Rutherford,W. H. 1964. Investigation of the Arkansas Valley wintering
goose flock. Colo. Game, Fish and Parks Dept., Game Res. Rept., Fed.
Aid Proj. W-88-R. Oct. p. 27-46.
1965. Investigation of the Arkansas Valley wintering goose flock.
Colo. Game, Fish and Parks Dept., Game Res. Rept., Fed. Aid Proj. W-88-R.
p , 87-104.
1966. Investigation of the Arkansas Valley wintering goose flock.
Colo. Game, Fish and Parks Dept., Game Res. Rept., Fed. Aid Proj.
W-88-R. p. 53-67.
1970. The Canada geese of southeastern Colorado.
Fish and Parks. Tech. Publ. No. 26.

Prepared by _-':"&gt;-J:/1r...-~1"",,~.&amp;L· .~?~~. _?....::.,KI......-....·:::=;·:oq.q
j~~~~~J!L'7
__
Michael R. szymcz~n.J
Assistant Wildlife Researcher

Colo. Div. Game,

��Table 14.

Percentages of total band recoveries,

Five Year Averages
1951-55
1956-60
1961-65

Area

Central F1~aI

Pacific Flyway
F1~aI

Mexico

Total Number
of Recoveries

1966-67

1967-68

7.5
0.7

7.5

bandings,

by area and year of recovery,

all bandings

(continued).

1971-72

1972-73

Total
Number
Recoveries

2.5
3.0

2.4
1.0

5.5

3.3

7.9
0.8
3.2
11.9

127
54
21
202

3.9
1.7
0.6
6.2

1.0

1.4

1.6

57

1.8

RecoverI Year
1968-69
1969-70
1970-71

Percentage
of Total
Recoveries

(cont.)

Texas
Panhandle
Waggoner Ranch
Gulf Coast
Total

MississiEEi

Arkansas Valley post-season

2.3
2.0
0.8
5.1

2.8
2.2
0.2
5.2

2.4

2.2

4.6
1.5
0.5
5.5

8.2

0.8

2.2
9.7

5.0
0.7
1.4
7.1

0.7

1.4

6.7
1.0
1.0
8·7

0.1

0.7

0.7

0.1

748

I

00
.....

677

770

134

134

141

105

198

211

126

3

0.1

1

0.03

3,244

I

�Table 15.

Composite

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69
1969-70
1970-71
1971-72

344
650
900
0
0
0
0
347
217
250
306
334
369
335
780
0
389
646
905
358
532
694

8,356

dynamic analysis of band recoveries

from Canada geese banded as adults, Arkansas

Valley, Colorado,

1

2

3

4

5

6

Recoveries
7
8

45
71
85

23
37
55

19
24
50

10
23
31

7
21
25

12
6
20

5
11
9

3
6
12

3
9
10

2
7
6

2
7
2

0
4
4

X
X
X

X
X
X

X
X
X

X
X
X

X
X
X

X
X
X

X
X
X

X

X
X

X

X

X
X
X

X

X

X

X

X

X

X

X

X

33
21
27
29
26
35
24
55

27
9
16
13
18
21
12
40

17
7
6
18
12
14
15
20

8
4
12
12
5
4
10
22

5
4
8
4
13
11
4
12

8
2
4
7
3
5
4
13

2
1
6
3
2
2
11
9

2
1
3
2
2
5
3
5

2
0
2
3
4
2
1

X

X

X

X

X

X

X

29
35
22
21
23
23

26
17
39
21
14

10
29
42
6

9
25
15

8
10

6

604

388

289

226

132

90

61

44

36

23

15

10

6

3

2

2

0

0

0

Banded Birds
Eligible

8,356

7,662

7,130

6,772

5,867

5,221

4,832

4,832

4,052

3,717

3,348

3,014

2,708

2,458

2,241

1,894

1,894

1,894

1,894

Recoveries per
1,000 Banded

72.3

\50.6

40.5

33.4

22.5

17.2

12.6

9.1

8.9
214.2

6.2

4.5

3.3

2.2

1.2

1.1 . _ Q.O
0.9
286.5

0.0/

0.0

28.3
826.5
.259

19.4

0.0/

0.0

Total
Recoveries

Alive Going
into .Period
Mortality

Rate

286.5 \214.2
.252

163.6

123.1

89.7

67.2

X

bl: Year Fo11owins Bandins
9
10
11
12

1951-1972.

X
X

3
1
1
2
1
0

13

14

15

16

1
2
2

1
0
0

0
1
1

1
0
1

X

X
X
X

X

X

X

X
X
X

X

X

X
X
X

0
0

X

X

X

X

1
0
1
1
1

0
0
0
2

1
0
0

2
0

0
0

X

X

17

18
0
0
0

0
0

19
0
0
0
0
0

0
0
0
0

I

00

."
I

50.0

37.4

1: =.
13.2

8.7

5.4

3.2

1.1
2.0
1: = 1,113.0
.257

0.0

�Table 16.

Composite

dynamic

Number
Banded

Year
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964"-65
1965-66
1966-67
1967-68
1968-69
1969-70
1970-71
1971-72

analysis

of band recoveries

from Canada geese banded as juveniles,

Recoveries

Arkansas

bI Year Following

Valley, Colorado,

1951-1972.

Banding

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

31
58
52

25
31
25

11
33
25

11
20
24

4
16
20

5
8
11

4
7
11

7
6
9

1
4
2

1
4
4

0
0
4

0
5
1

0
2
3

1
1
1

1
0
1

0
0
0

0
1
2

1
0
0

X
X
X
X

X
X
X
X

X
X
X

X
X
X

X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X

X
X

X
X

X

0
0

0

X

X
X
X
X

X
X

X

X
X
X
X

X
X

X

X
X
X
X

0
0
0
0

21
25
16
15
10
40
16
15

16
16
10
5
12
24
11
7

6
16
3
4
10
9
11
8

4
8
14
7
0
14
13
1

4
7
8
8
1
6
5
0

1
3
0
7
1
2
2
3

4

0
0
3
1
2
8
1

0
1
1
:1
1
5
5
1

3
1
0
1
1
2
0

1
2
0
2
0
1

0
1
1
2
0

0
0
0

0
0

X

300
628
578
0
0
0
0
176
298
167
248
123
399
251
198
0
285
270
184
241
301
337

X

X

X

X

X

X

18
16
9
18
19
15

10
9
11
9
8

5
8
11
3

12
4
3

9
6

7

4,984

394

229

163

135

94

50

X
X

X
X

0
1
0
0

0
0

0

I

00
IN
I

--

Total
Recoveries
Banded Birds
Eligible
Recoveries per
1,000 Banded
Alive Going
into Period
Mortality

Rate

41

38

15

15

8

7

5

3

2

0

3

1

0

4,984 4,647 4,346 4,105 3,921 3,651 3,366 3,366 3,168 2,917 2,518 2,395 2,147 1,980 1,682 1,506 1,506 1,506 1,506
79.1 \49.3

37.5

32.9

24.0

283.6 \204.5

155.2 117.7

84.8

.279

12.2· 11.3
4.7
204.5
60.8 _41._1.~A.9
23.6
795.0
.257
13.7

5.1

3.2

2.9

2.3

1.5

18.9

13.8

10.6

7.7

5.4

1.2
0.0
I:= 283.6
2.7
3.9
l:.,1,078.6
.263

2.0

0.7/

0.0

2.7

0.7/

0.0

�Table 17.

Composite dynamic analysis of band recoveries

Year
Banded

Number
Banded

1950-51
1951-52
1952-53
1953-54
1954-55
1955-56
1956-57
1957-58
1958-59
1959-60
1960-61
1961-62
1962-63
1963-64
1964-65
1965-66
1966-67
1967-68
1968-69
1969-70
1970-'-71
1971-72

from Canada geese banded in all age classes, Arkansas Valley, Colorado, 1951-1972.
Recoveries bX Year Following Banding
11
10
7
8
9

12

13

14

15

16

17

18

19

2
7
6

0
9
5

1
4
5

2
1
1

1
1
2

1
0
1

0
1
2

1
0
0

0
0
0

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X

X
X

X

4
3
1
4
1
1

1
1
2
3
1

0
1
0
2

1
0
0

2
0

0

1

2

3

4

5

6

644
1,278
1,478
0
0
0
0
523
515
417
554
457
,768
586
978
0
674
922
1,089
599
836
1,032

76
129
137

48
68
80

30
57
75

21
43
55

11
37
45

17
14
31

9
18
20

10
12
21

4
13
12

3
11
10

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X

54
46
43
44
36
75
40
70

43
25
26
18
30
45
23
47

23
23
9
22
22
23
26
28

12
12
26
19
5
18
23
23

9
11
16
12
14
17
9
12

9
5
4
14
4
7
6
15

6
1
6
6
3
4
19
10

2
2
4
4
3
10
8
6

5
1
2
4
5
4
1

X

X

X

X

X

X

X

47
51
31
39
42
38

36
26
50
30
22

15
37
53
9

21
29
18

17
16

13

13,350

998

617

452

325

226

139

102

82

51

38

23

17

11

6

4

2

3

1

0

Banded Birds
Eligible

13,350

12,318

11,482

10,883

9,794

8,872

8,198

8,198

7,220

6,634

5,866

5,409

4,855

4,438

3,923

3,400

3,400

3,400

3,400

Recoveries per
1,000 banded

74.8

,50.1

19.4

.~._9_

23.1

15.7

12.5

10.0

7.1
5.7
206.8

3.9

3.1

2.3

1.2
1.0
E - 281.6

0.6

0.9

0.31

0.0

Alive Going
into Period

281.6

"206.8

156.7

117.3

87.4

64.3

48.6

36.1

26.1
19.0
801.4

13.3

9.4

6.3

4.0
2.8
E = 1,083.0
.260

1.8

1.2

0.31

0.0

I

00

+I

--

Total
Recoveries

Mortality Rate

.266

.258

�-85-

Table 18. Relative recovery rate analysis of band recoveries from Canada
geese banded as adults, southeast Colorado, 1951-72.
Year
Banded

Number
Banded

1950-51

344

CD 1/

89

1951-52

650

229

158

.3523

1952-53

900

313

CD

.3478

1957-58

347

CD

78

1958-59

217

50

29

1959-60

250

86

1960-61

306

1961-62

Number of Recoveries
1-n
2-n

Recovery Rates
1-n
2-n

Survival
Rate

.2587

.734

.2431

.699

.2248

.976

.2304

.1336

.393

59

.3400

.2360

.752

96

67

.3137

.2190

.841

334

87

61

.2605

.1826

.681

1962-63

369

99

64

.2683

.1734

.692

1963-64

335

84

60

.2507

.1791

.794

1964-65

780

176

CD

.2256

1966-67

389

CD

59

.1517

.845

1967-68

646

116

81

.1796

.1254

.962

1968-69

905

118

96

.1304

.1061

.787

1969-70

356

48

27

.1348

.0758

1.091

1970-71

532

37

14

.0695

.0263

.795

1971-72

694

23

Total

8,354

.0331
3.1367

Average Survival Rate

.745

Average Mortality Rate .255
1/
- Cannot use

2.3356

Mortality
Rate

�-86-

Table 19. Relative recovery rate analysis of band recoveries from Canada
geese banded as juveniles, southeast Colorado, 1951-72.
Year
Banded

Number
Banded

1950-51

300

CU 1/

72

1951-52

628

196

138

.3121

1952-53

578

195

CU

.3374

1957-58

176

CU

39

1958-59

298

81

1959-60

167

1960-61

Number of Recoveries
1-n
2-n

Survival
Rate

Mortality
Rate

.2400

.769

.231

.2197

.651

.349

--

.2216

.815

.185

56

.2718

.1879

.592

.408

53

37

.3174

.2216

.981

.019

248

.56

41

.2258

.1653

.•550

.450

1961-62

123

37

27

.3008

.2195

.834

.166

1962-63

399

105

65

.2632

.1629

.576

.424

1963-64

251

71

55

.2829

.2191

1.205

0.000

1964-65

198

36

CU

.1818

1966-67

285

CU

43

.1509

.947

.053

1967-68

270

43

27

.1593

.1000

.541

.459

1968-69

184

34

25

.1849

.1359

1.088

0.000

1969-70

241

30

12

.1249

.0498

.555

.445

1970-71

301

27

8

.0897

.0266

.598

.402

1971-72

337

15

Total

4,984

Recove!:Z Rates
1-n
2-n

.0445
3.·09·65 2.3208

Average Survival Rate

.749

Average Mortality Rate .251
1/

Cannot use

�-87-

Table 20. Relative recovery rate analysis of band recoveries from Canada
geese banded in all age classes, southeast Colorado, 1951-72.
Year
Banded

Number
Banded

1950-51

644

1951-52

Number of Recoveries
1-n
2-n

Recovery Rates
1-n
2-n

Survival
Rate

Mortality
Rate

.2500

.752

.248

.2316

.674

.326

.2199

.858

.142

CU 1:/

161

1,278

425

296

.3326

1952-53

1,478

508

CU

.3437

1957-58

523

CU

115

1958-59

515

132

86

.2563

.1670

.501

.499

1959-60

417

139

96

.3333

.2302

.829

.171

1960-61

554

151

107

.2776

.1967

.725

.275

1961-62

457

124

88

.2713

.1967

.752··

.248

1962-63

768

201

126

.2617

.1641

.620

.380

1963-64

586

155

115

.2645

.1962

.910

.·090

1964-65

978

211

CU

.2157

1966-67

674

CU

99

.1469

.857

.143

1967-68

922

158

107

.1714

.1161

.854

.146

1968-69

1,089

148

117

.1359

.1074

.825

.175

1969-70

599

78

39

.1302

.0651

.850

.150

1970-71

836

64

22

.0766

.0263

.715

.285

1971-72

1,032

38

Total

13,350

.0368

3.1109
Average Survival Rate

.744

Average Mortality Rate .256
1/
.
- Cannot use

2.3142

��October 1974

-89-

JOJ~ PROGRESS REPORT

State of

COLORADO

Proj ect No.

W-88-R-19

"lork Plan No.
Job Title

Migratory Bird Investigations
Job No.---~--=-6

2

Studies of Canada Goose Populations in Colorado Transplant Areas

Period Covered:

April 1, 1973 to March 31, 1974

Personnel:
C. Bryant and Staff, MOnte Vista National Wildlife Refuge; M.
Babler, D. Benson, C. Brown, G. Claassen, R. Clark, C. Crawford, M. DePra,
J. Froth iIigham , H. Funk, B. Goetze, J. Gumber, T. Henry, J. Hicks, J. Hobbs,
R. Hopper, C. Leonard, J. Lorentzson, T~ Lynch, F. Marcoux, M. McLain, K.
Miller, D. Owens, S. Palm, J. Pogorelz, C. Roberts, L. Rottman, W. Russell,
G. Saville, L. Searle, H. Spear, E. Wagner, and M. Szymczak, Colorado
Division of Wildlife.
ABSTRACT
According to tail fans, 1,200, or 12 percent of the total harvest of Canada
geese in north-central Colorado was composed of "small" Canada geese. A
coordina~ed count throughout the entire Hi-Line population wintering range
totaled a record high 74,305 on December 13, 1973. Only 28,487 Canada geese
were recorded in Hi-LiIie winter areas on January 9, 1974. The reduction in
goose numbers iIithe Hi-Line area between the two counts was attributed
primarily to emigration. Within Colorado, counts indicated there were
52,274 and 23,465 geese, respectively, on December 13, 1973 and January 9,
1974 iIithe Hi:-Line area. An estimated 11,048 hunters harvested 10,169
Canada geese iIinorth-central Colorado duriIig the 1973 hunting season. The
season bag restriction of six geese was not iIieffect during the 1973 season,
yet only 1.3 percent of the hunters reportedbaggiIig more than 6 geese,
accountiIig for only six percent. of the total harvest. According to periodic
inventories the San Luis Valley wintering Canada goose population size in
1973-74 was only 60 percent of the previous year's total. An estimated 233
hunters harvested 112 geese during the special season in the San Luis Valley.
In west-central Colorado 254 hunters in Mesa County bagged 75 Canada geese
and in Garfield County 46 hunters bagged 9 geese. A total of 965 Canada
geese were recorded on January 11, 1974 during an inventory of waterfowl
in west-central Colorado.

��-91-

STUDIES OF CANADA GOOSE POPULATIONS IN COLORADO TRANSPLANT AREAS
Michael R. Szymczak
P. S. OBJECTIVE
To investigate the status of resident and migrant Canada goose flocks and
their interrelationships in areas in which populations have been established
through transplant programs in Colorado.
SEGMENT OBJECTIVES
La , To determine breeding areas of Canada geese wintering in north-central
Colorado.
lb.

To examine migration routes and/or harvest patterns and distribution
of Canada geese wintering in north-central Colorado arid the San Luis
Valley.

2.

To estimate hunting pressure on, and hunter harvest of, Canada geese in
north-Central Colorado, the San Luis Valley and west-central Colorado.

3.

To estimate annual mortality of Canada geese wintering in north-central
Colorado.

4.

To make recommendations for continuing Canada goose hunting seasons in
west-central Colorado, north-central Colorado and the San Luis Valley.
METHODS AND MATERIALS

Because of a change in Federal Aid report deadlines, band recoveries were
not analyzed prior to the completion of this report. Therefore, analysis of
information for achieving segment objectives 1, 3 and 4 is not complete.
Tail fans collected through the Bureau of Sport Fisheries and Wildlife
Waterfowl Parts Collection Survey were examined.
Canada geese in north-central Colorado were counted on December 13, 1973
and January 9, 1974. In the San Luis Valley geese were counted on October 24,
November 27, December 19, 1973, and January 16, 1974.
A mechanica~ly random $ample of hun.t~rs issued permits to hun~ geese in th~
nOJ;"th-cenc}:"al
Colo}:"ado"Special P~rmit" goose hunting area we}:"~lMile.q
questionnaires inquiring about thei}:"hun.ting activity and §t,lcce.!i!~.
All of
the hunters hcl.dtng spec Lal, permits to hunt· geese. in the.San Lt,li§ValleY
and west-Ce.ntra+ Colorado were mailed similar questionna1re.s. All data
recieve.dwe.re tabulated and st atLst Lca.LLyana Lyzed ,

�-92-

RESULTS
ANDDISCUSSION
North-central Colorado - Breeding
Range of the Hi-Line Wintering Population
Tail Fans
Measurements of tail fans collected through the "Parts Collection Survey"
from geese ba.gged.in north-central
Colorado during the 1973 season indicate
that 17 percent of the birds bagged were "small" Canada geese, ..possibly
members of the Arctic nesting Short Grass Prairie Population.
According to
.. tail fans collected,
small geese have made up 11 percent, 17 percent, and
in north--central Colorado in the years 1970, 1971,
8percerit of the·harvest
and 1972, respectively
(Table 1) • However, pooling Counties together without weighting factors is not consi.deredvalid,
as the number of tail fans
collected
b)TCountyare generally not proportional to the ha~est.
If.
sampl,e sizesareadequate,the
distribution
of harvest classified
as "large"
or "small" can be compared on a county basis.
Since 1970, the harvest in
Weld County has consistently
been composed of a higher percent of small
geese than the harvest in Larimer or Boulder counties (Taple 1).
.

..

Tail fan sample size for the 1973 season seems to be adequate for Larimer,
:Soulder,and possibly Weld count.Les, Applying the percentages presented in
Table 1 to theharyest
information which appears in Table 6 results in an
estimatedharvestof·343
small birds in Larimer County, 187 in Boulder
County, and 671 in Weld County.
PopulationDis

tribtition

Coordinated inventories
throughout the Hi-Line population's
wintering area
were reduced to 2 in 1973-74. A record high number of 74,305 birds was
recorded during the December 13,1974coui:tt,
and a record low of 28,487 was
. recorded on the January 9, 1974 inventory (Table 2). About 70 percent of
the·populationwasfound
in Colorado in mid-December compared to 82 percent
.in January • It is difficult
.t o explain the drastic
fluctuation
in Hi-Line
population.
The spring· of 1973 was aipoo r production year for Hi-Line
Canada geese.
The most logical explanation woul.d be that a larger than
usual portion of the December population was composed of Short Grass Canada
geese.
Muchmore information than is currently available is needed to
analyze the situation.
Within Colorado, approximately 60 percent of the geese were located in the
Fort Collins-Loveland area during both inventories
(Table 3). An unusually
large number of birds (10,380) were located in the Brighton~Greeley-Fort
MOrganarea during t.hevmfd-Decemharinventory.
Only in late November1971
have more than 10,000 geese been counted in the Brighton-Greeley-Fort
. Morgan area (Table 4).
Over 8,000 of the birds counted during the 1973 midDecember inventory were located on Empire or Riverside· Reservoir (Table 3).
North-central
Colorado experienced a cold weather period in late December
and early January, which undoubtedly had a major effect on the movementof
geese out of the area.

�Table 1.
Classification
of Canada goose t.ailfans
collected
through the Bureau of Sport Fisheries
and Wildlife's
Parts Collection
Survey,. from birds harvested in no rt.h=cen t ra.l, CoLorado , 1970-1973 hunt.Lng seasons.

County of Harvest

1970
Percent
Percent
. Small
Large

1971
Percent
Percent
Small
Large

Year
1972
Percent
Percent
SmAll
Large

8(2)

Larimer

92 (36)-!./

8(3)

91(41)

9 (4)

92(22)

Boulder

100(1)

-

86(18)

14 (3)

100(10)

-

Arapahoe

-

-

-

-

-

.

100 (2) .

Adams

100(2)

-

100(4)

-

100(1)

Weld

82(18)

18(4)

59 (13)

41(9)

67(2)

Morgan

-

-

-

-

-

Total

89(57)

11(7)

83(78)

17(16)

92(35)

1:./ Sample size

in parentheses.

.

1973
Percent
Percent
Small
Large

Total
Percent
Percent
Small
Large

. 93(28) ..

7 (2)

92 (127)

8(11)·

84(42)

16(8)

87 (71)

13(11)

_.

-

100(2)

.1

\0

..,.

-

100(7)

-

33(1)

79 (11).

21(3)

72(44)

28(17)

-

-

100(4)

-

100(4)

83(81)

17(17)

8(3)

\.oJ
1

�-94-

Table 2.

Results of Hi-Line Canada goose population inventories, 1973-74.

Location

December 13

January 9

Montana

7,618

1,285

Wyoming

13,144

1,833

Colorado

52,274

23,465

New Mexico

1,269

1,904

Totals

74,305

28,487

Table 3.

Results of north-central Colorado goose surveys, 1973-74.

Area

December 13

January 9

925
50
20

3,250
23

Fort Collins-Loveland
Lindenmeier Lake
Reservoir No. 8
Reservoir No. 8 Annex
Reservoir No. 5
Reservoir No. 6
Bureau of Standards
Douglas Reservoir
Reservoir No. 15
Boxe1der Reservoir (Boxeye)
Indian Creek Reservoir
Terry Lake
Watson Lake
Claymore Lake
Sterling Ponds
Dean Acres
College Lake
Parkwood Lake
Greenwalt Reservoir
Fossil Creek Reservoir
Boyd Lake
Hollister Lake (Windsor Reservoir)
New Windsor Reservoir
Timnath Reservoir
Woods Lake
Cobb Lake
Subtotal

6

108
21
30
100
1,000
153
2,430
607
2,200
448
85
8,144
4
30
5,700
1,600
175
7,600
97
25

o

o
13

o
220

o
o
o
275
1,800

o
26
33
1,419

o

o

o

225
475
ll5
5,000
960
500
520

31,558

14,854

--------------------7-----------------------------------------------

_

�-95Table 3.

Results

of north-central

Location

Boulder-Longmont

Colorado

goose surveys,

December

13

1973-74.
January

9

Area

o
o
o
o

Terry Lake
Crystal Lake
Left Hand Valley Reservoir
Culver Reservoir
McCall Reservoir (Stamp)
Baller Lake
Faivre Ponds
Boulder Reservoir
Baseline Reservoir
Ish Lake
Swede Lake
Valmont Reservoir

1,225
80
20
71
336
35
90
200
65
1

o
o
o

35
6,100

.4,640

Subtotal

8,258

4,765

105
900
187

1,500
75

125

o

o

o

Denver Area
Marston Reservoir
Bowles Lake Area
Sloans Lake
Denver City Park
Standley Lake
Main Reservoir
KettringLake
Green Gables Country Club
Norgren Reservoir
Lowry Air Base
In Field Along Belleview Avenue

o
150

o

5

225
95

54
222

a
a

o
o

57
80

460

a

2,078

2,037

Barr Lake
Horsecreek Reservoir
Milton Reservoir
Latham Reservoir
Seeley Reservoir
Riverside Reservoir
Empire Reservoir
Jackson Reservoir
Orchard Area-South Platte River

630
550
25
15
625
4,850
3,650
35

1

o

108
1,700

Subtotal

10,380

1,809

Grand Total

52,274

23,465

Subtotal

Brighton-Greeley.,.Fort Morgan Area

a
a
a
a

a
a

�-96-

Table 4.

Winter inventories of the Colorado Hi-Line Canada goose population.
Fort Co11insLoveland

Longmont-Bou1derDenver

9,739

2,883

991

13,613

12,217

4,029

678

16,924

1968-69
November 20
December 19
January 2 &amp; 13

15,848
20,905
19,693

3,461
4,236
4,874

2,667
1,170
775

21,976
26,311
25,342

1969-70
November 5
November 28
December 23
January 6

8,737
31,350
18,522·
30,650

2,255
3,782
5,668
5,060

390
1,374
. 1,259
1,914

11,382
36,506
·25,499
37,624

1970-71
November 4
November 23
December 22
January 6

12,612
29,970
36,034
19,879

1,690
16,710
12,664
15,566 .

·348
1,370
3,055
2,425

14,650
48,050
51,753
37,870.

1971-72
November 4
November 23
December 21
January 10

25,699
31,072
31,516
19,117

1,815 };./
9,181
19,525
10,742

6,905
13,525
4,476
3,661

34,419
53,788
55,517
33,520

1972-73
November 6
November 22
December 20
January 10

15,230
21,118
13,841
13,595

1,240 };./
5,764
19,290
18,709

1,050
4,981
1,225
1,405

17,520
31,863
34,356
33,709

1973-74
December 13
January 9

.31,558
14,854

10,336
6,802

10,380
1,809

52,274
23,465

Count Date
1967
January 9
1968
January 10

};./
Denver area not included.

Brighton-Gree1eyFort Morgan

Total

�-97-

Hunting Pressure and Harvest
An estimated 11,048 hunters harvested 10,169 geese in north-central Colorado
during the 1973 hunting season (Tables 5 and 6). Harvest increased consid-'
erably over 1972 levels in all counties .and reached record levels in furgan
County (Table 6). Comparative statistics concerning hunting pressure and
harvest by County for 1972 ,and 1973 l:Iuntingseasons are presented in Table 7.

Table 5.

Hunter activity and success in permit area.

Year

Number
Permits Issued

Est. Number
Active Hunters

Average
Days Hunted

Average
Seasonal Bag/Hunter

1964-65

1,608

1,427

6.1

0.48

1965-66

2,335

1,578

6.5

0.53

1966-67

3,996

2,910

5.1

0.41 '

1967-68

3,000

2,256

6.7

0.55

1968-69

3,000

2,379

9.2

1.11

1969-70

8,342

6,149

7.4

0.68

1970...,71

13,611

11,187

6.9

1.08

1971-72

14,847

12,920 '

7.3

1.14

1972

12,702

9,938

6.1

0.53

1973

13,253

11,048

6.1

0.92

The six bird seasonal bag restrictions for the north-central Colorado permit
area was lifted for the 1973 season. Only 1. 3 percent of the hunters
reported bagging more than 6 geese during the 1973 season (Table 8). If
these hunters would have been restricted to bagging only 6 birds, the estimated harvest for the north-central Colorado permit area would have been
9,587,6 percent less than the actual harvest.

�-98-

Table 6.

Distribution

of harvest,

Larimer

Weld

1964-65

504

181

1965-66

665

144

29

838

1966-67

764

409

11

1,184

1967-68

944

265

37

1,246

1968-69

1,584

886

161

2,631

1969-70

2,431

1,112

383

146

100

4,172

1970-71

7,486

3,544

620

93

370

12,112

1971-72

7,723

4,484

1,804

464

296

14,771

1972

2,855

1,547

413

329

126

5,270

1973

4,895

3,195

1,166

590

253

70 10,169

Hunting pressure

and harvest,

Year

Table 7.

by county, in the permit area.

Boulder

Morgan

Adams

Other

Total

685

Estimated

by county, north-central

Number

Estimated

Number

Colorado.

County

1972

1973

Percent Change
From 1972

Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

5,067 (± 265)
27,141 (+2,455)
2,855
403)
5.4
0.56
0.105

5,232 (+ 271)
29,854 (+2,556)
4,895
706)
5.7
0.94
0.164

+,3.3
+ 10.0
+ 71.5
+ 5.6
+ 67.9
+ 56.2

4,332 (+ 259)
18,007 (+i,815)
1,547 (+ 290)
.4.2
0.36
0.086

4,607 (+ 266)
19,741 (+1,814)
3,195
503)
4.3
0.69
0.161

+ 6.3
+ 9.6
+106.5
+ 2.4
+ 91.7
+ 87.2

Larimer

(±:

(±

Weld Count
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

(±:

----------------------------------------------------------------------------

�-99Table 7. Hunting pressure and harvest, by county, north-central Colorado (cont.).

Boulder County

Estimated Number
1972

Estimated Number
1973

Percent Change
From 1972

Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
.Ave. Bag/Hunter
Ave. Bag/Hunter Trip

1,757 (+ 187)
10,570 (±:1,885)
413 (+ 141)
6.0.
0.24
0.039

1,868 (±192)
12,430 (±1,85l)
1,166 (± 274)
6.7
0.62
0.094

+ 6.3

924 (+ 143)
3,157
'703)
329 (+ 141)
3.4
0.36
0.104

1,011 (+ 147)
3,363 (+ 684)
590 (± 197)
3.3
0.58
0.175

+

462 (± 108)
2,191 (± 780)
96)
126 (±
4.7
0.27
0.058

499 (+ 111)
1,975 (+ 712)
253 (± 152)
4.0
0.41
0.128

+

+ 17.6
+182.3
+ 11.7
+158.3
+141.0

Morgan County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

(±

9.4

+ 6.5
+ 79.3
2.9
+ 61.1
+ 68.3

Adams County
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trips

8.0
9.9
+100.8
- 14.9
+ 51.9
+120.7

Arapahoe and Jefferson Counties
21
77
70
3.7
3.33
0.909

Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave. Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip
Entire Permit Area
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Ave..Hunter Trips/Hunter
Ave. Bag/Hunter
Ave. Bag/Hunter Trip

9,938 (+ 223)
61,066 (±3,059)
5,270 (± 516)
6.1
0.53
0.086

11,048 (+ 207)
67,440 (+3,271)
10,169 (+ 936)
6.1
0.92
0.151

+ 11.2
+ 10.4
+ 93.0
+ 73.6
+ 75.6

�Table 8.

Season Bag
0
1
2
3
4
5
6

Estimated distribution of season bag for active hunters in the north-central Colorado permit area ,

Number

1969-70
Percent

Number

1970-71
Percent

Number

1971-72
Percent

Number

1972
Percent

.Number

1973
Percent

4,263

69.3

6,588

58.9

7,006

54.2

6,857

69.0

6,817

61.7

836

13.6

1,620

14.5

2,209

17.1

1,391

14.0

1,823

16.5

7.4

1,073

9.6

1,428

11.1

666

6.7.,

1,061

9.6

3.9

564

5.0

768

5.9

298

3.0

552

5.0

2.5

528

4.7

633

4.9

616

6.2

287

2.6

138

2.2

342

3.1

371

2.9

50

0.5

210

1.9

69

1.1

472

4.2

505

3.9

60

0.6

155

1.4

55

0.5

22

0.2

22

0.2

44

0.4

11,048

100.0

452
238
153

8
9
10+

6,149

100.0

11,187

100.0

12,920

·100.0

0
0
I

7

Total

I
f-l

9,938

100.0

�-101-

.San Luis Valley·
Distribution
Inventories of Canada·geese in the San L~is Valley through the fall and
winter of 1973~74 indicated a r-educed popul.atiLontf'I'ab.Le 9). Undoubtedly,
a number of birds .were .mtssed during the OCtober census.
The number of
birds recorded during the November, December and January counts were on the
average, only 62 percent of the previous year's totals.

Table 9•. Winterinveritories

of .Canada geese in the San .Lud.s Valley of Colorado .

.... 1/

Year

October

Numberof GeeseDecember
November

January

1,490

1,050

1,261

1970-71
1971-72

1,570

913

1,255

1,286

1972~73

1,006

1,196

1,079

1,621

1973-74

289

736

774

905

for the 4 years

generally

11 Comparedmonthly inventories

taken within

5 days .

of each other.

The distribution
of geese within the San Luis Valley during the fall. and
winter of 1973-:74wasquite
simila.r to previous years with the majority of
the birds located either on the Monte Vista National Wildlife Refuge or on
the Rio Grande River (Table 10).
Harvest
A total of 300 permits were issued authorizing the taking of one goose during
the 1973 special San Luis Valley goose season.
An estimated 233 active.
hunters harvested 112 geese for a success ratio of 0.48, the lowest success
ratio recorded since. permits were first issued in 1970 (Tables 11 and 12).
Harvest during the 1973 season declined 56 percent
level (Table 13). The harvest was down 68 percent
22 percent in Alamosa County.

from the 1972-73 season
in Rio Grande County and

Forty-four percent of the geese taken in the San Luis Valley were reportedly
harvested on or within 2 miles of the Monte Vista National Wildlife Refuge.
About· 30 percent of the harvest occurred along the Rio Grande River between
Monte Vista and Ahunosa.

�-102Table 10. Results of Canada goose surveys during the 1973-74 hunting
in the San Luis Valley.

Area

October

24

November

27

December

19

season

January 16

Monte Vista National
Wildlife

Refuge

133

112

325

890

South Fork to Del Norte

0

0

27

0

Del Norte to Monte Vista

0

38

130

15

122

172

5

0

0

173

228

0

0

104

55

0

San Luis Lakes Area

34

80

0

0

Russell

0

0

4

0

Smith Reservoir

0

40

0

0

Sanchez Reservoir

0

17

0

0

289

736

774

905

Rio Grand River

Monte Vista to Alamosa
Alamosa

to State Line

Conejos River
McIntire
Area

and Sego Springs

Lakes

Total

Table 11.

Hunter activity

Years

Number
Permits
Issued

Est. Number
Active
Hunters

Average
Days Hunted

Average
Seasonal
Bag/Hunter

1970-71

200

164

3.4

0.65

1971-72

350

296

4.2

0.60

1972-73

400

311

4.6

0.82

1973-74

300

233

4.2

0.48

and success in the San Luis Valley goose permit area."

�-103-

Table 12. Distribution
permit area.

of harvest,

by county

in the San Luis Valley goose

Year

Rio Grande

Alamosa

Saguache

Conejos

Costilla

Total

1970-71

74

31

2

0

0

107

1971-72

121

50

1

5

0

177

1972-73

195

51

2

1

6

255

1973

62

40

3

1

6

112

Table 13. Hunting pressure and harvest
area 1972-73 and 1973 hunting seasons.

Estimated

in the San Luis Valley goose permit

Number

1972

Total Individual

311 (±

Hunters

Number

Percent Change
From 1972

5)

25.1

974 (± 65)

- 32.4

112 (±

- 56.1

Estimated

1973

6)

Total Hunter Trips

1,440 (± 54)

Total Geese Bagged

255 (±

8)

233

(±

.Average Hunter Trips/Hunter

4.6

4.2

Average

0.82

0.48

0.18

0.11

Bag/Hunter

Average Bag/Hunter

Trip

West-Central
Hunter Activity

6)

Colorado

and Harvest

For the 1973 west-central Colorado goose season, 300 permits authorizing the
taking of 1 goose were issued for Mesa County, and 66 permits with the same
bag limit were issued for Garfield County.
In Mesa County 254 active hunters bagged a total of 75 geese for an average
seasonal bag of 0.30 birds per hunter (Tables 14 and 15). In Garfield County
46 of the 66 hunters were active, bagging nine geese for a 0.20 bird per
hunter bag (Tables 14 and 15).

�-104-

Table
area.

14 .. Hunter activity

and success

in west-central

Colorado goose permit

Average
Days Hunted

Average
Seasonal Bag/Hunter

Year

Number
Permits Issued

1971

150

112

1972

250

182

. Mesa Co.

300

254

3.9

.30

Garfield
Co.

66

46

3.7

.20

Entire
Area

366

300

3.8

.28

Est.
Active

Number
Hunters

2~9

.29
.32

1973

Table 15. Distribtit
permit area.

ion of harvest. by year in west-central

Year

Mesa County

CoLorado goose

Garfield

County

Total

1971

33

1972

58

1973

75

9

Hunting activity
and harvest information for the entire west-central
rado permit area for 1971, 1972, and 1973 is presented in Table 16.
The results
of a Canada goose inventory
in Table 17.

by7/ZJ:vL.Jl7? ....~
rv

Prepared

in west-central

.~'

..

Michael R. Szymc~ ~
Assistant Wildlife Researcher

"

84

Colo-

Colorado is presented

�-105Table 16. HUnting pressure and harvest in the west-central Colorado goose
permit area, 1971, 1972 and 1973.
1971

1972

1973

Total Individual Hunters

112 (+ 4)

182 (± 5)

300 (+ 6)

Total Hunter Trips

329 (+20)

579 (+30)

1,152 (+51)

Total Geese Bagged

33 (± 4)

58 (+ 5)

84 (+ 7)

Average Hunter Trips/Hunter

2.9

3.7

3.8

Average Bag/Hunter

0.29

0.32

0.28

Average Bag/Hunter Trip

0.10

0.100

0.073

Table 17.

Canada goose inventory. west-central Colorado, January 11, 1974
Number of Geese

Area
Highline Lake

65

In Field North of Loma

175

Colorado River
Silt to DeBeque Canyon

249

Palisade to Grand Junction

12

Grand Junction to Horsethief Canyon

6

Hor set.h Iaf .Canyon to Utah State Line

131

North Fork of Gunnison
Hotchkiss to North Fork COnfluence

10

Gunnison River
Black Canyon

25

Confluence with North Fork to Delta

79

Delta to Grand Junction

206

Sweitzer Lake

7

Total

965

Comparative Total January 1973

989

1972

1,183

1971

879

��-107-

JOB PROGRESS

State of ~

C=.;O;;.;;LO=RAD=..;;O
_

Project

W-88-R-19

No.

Work Plan No.
Job Title
Period

Covered:

October

1974

REPORT

}ligratory Bird Investigations
Job No

2

0

7

_

Non-Hunting Mortality Investigations
of Canada Geese in Southeastern· Colorado
October

27, 1973 to March 31, 1974

T. Spraker, Colorado State University; W. Adrain, J. Carsella,
Personnel:
G. Eyre, J. Gustafson, J. Jackson, D. Kenvin, R. Oakleaf, B. Petersen,
R. Rosette, and M. Szymczak, Colorado. Division of Wildlife.

ABS~RACT
Measurement of the comparative incidence of surface lead shot in agricultural
fields surrounding Turk's Pond indicated that fewer lead shot were present in
fields north and northwest of the pond than in other areas.
Greatest lead
shot accumulations were located ina portion of an alfalfa field north of
the pond which had not been plowed fora number of years.
Accumulations in
annual croplands were greatest to the east and southeast of the pond. Lead
s1;lotpellets were found in a predictable pattern in reference to the firing
line, with relative densities increasing slightly for 600 feet in front of
the firing line and decreasing beyond.
Densities remained fairly stable for
400 feet behind the firing line before decreasing.
Post-mortem examinations
of Canada geese collected at Turk's Pond indicated that 52.9 percent of the
birds had an impacted proventricali, 64.5 percent had lead shot fragments in
their ventriculus and 80 percent had either one of the conditions, or both.
Lead concentration levels from liver, kidney, muscle and bone tissues from
90 Canada geese are presented.
Sixty-three of 221 gizzards sampled from
1,522 dead geese picked up around Turk's Pond and judged to have died
shortly after the hunting season had at least one lead pellet fragment in
their ventriculi.
Sixty-one of the 63 gizzards contained fragments from
only one lead pellet.
According to chemical analysis of tissues, only 76.9
percent of those birds suffering from lead poisoning had lead shot fragments
in their gizzards.
Therefore, 565, or 37.1 percent of the 1,522 were actual
or potential lead shot victims.
Ninety-three percent, or 343 of 368 geese
removed from the Turk's Pond area which were Judged not to be victims of
gunshot wounds' and definitely died after the hunting season were considered
definite victims of lead poisoning.
A total of 908 Canada geese removed
from Turk's Pond area were considered victims or possible victims of lead
poisoning.

��-109-

NON-HUNTING
MORTALITY
INVESTIGATIONS
OF CANADA
GEESEIN SOUTHEASTERN
COLORADO
Michael R.Szymczak

The possibility
of some mortality
factor other than actual hunting or
crippling loss affecting the Canada goose population at Turk's Pond was first
evident in late January and early February, 1972.
Canada geese collected
at
Turk's Pond at that time exhibited definite
signs of lead poisoning.
In
the fall of 1972~ th.e basin of Turk's Pond was sampled to determine the
general occurrence of lead shot.
The results
of the basin sampling were
documented by Szymczak (In Press) •. Duri~gthe winter of 1972-73 no unusual
mortality was discernible
at Turk's.
However, a die:-off was again experienced in the winter of 1973"':74 with the bulk of this report devoted to investigational
techniques utilized
and results
of the study during this period.
P. S. OBJECTIVE
To identify non-hunting
southeast Colorado.

mortality

factors

of Canada geese at Turk's

Pond in

1.

Determine the incidence
and local vicinity.

of lead shot in the environment at Turk's

2.

Determine if
Pond.

3.

Determine if other causative agents. of disease or poisoning are responsible for mortality of Canada geese at Turk's Pond.
.

4.

Determine the magnitude of non:-hunting mortality

5.

Analyze data and formulate r ecommendatLons for preventative
me.asures
designed to lessen non-hunting mortality of Canada geese at Turk's Pond."

lead poisoning

occurs in the waterfowl

population

in the Turk's

Pond

at Turk's

Pond area .

METHODS
ANDMATERIALS
Agricultural
fields located directly
adj acent to Tllrk' sPond were sampled to
. determine the general incidence of lead shot on the surface of the ground in
respect to the pond and the firing line.
All fields were sampled with the
exception of the picked cornfield located directly
east of the pond and the
picked cornfield that was located outside the firing line, southeast of the
pond (Fig. 1).
The two afore mentioned cornfields
were not sampled because
(1) the cornfield to the southeast was disked before the sampling could take
place, and (2) sampling in f LeLds (Nos. 5, 6, 7 and 8) adjacent to the cornfield located directly
east of the pond indicated that the incidence of lead·
shot in the field· would be minimal.

�x
X
l\

N

'Y.

,

,

'f

wl-\SAT

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WHEAT

WHEAT
/.

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,
,,;

.,

., /lLt=ALFA

WHEAT

.\.W

f?\

,~.

I ALI='ALrA

'to

&lt;/
'9"~

:
&lt;, &lt;:

I

coRN

'"

""

r- - - -

WHEAT·

-r-=-+ .,- -

)C

~
~

COR III

CORN

I·.~

______
I P,·e, I.!.
I

. _I

No, rID

.I

C]

BO,-\/lJo..,.y L'-" •..,.' +0 r- S'\.""f'I~ P•.•.
I~s
Bo"l\dC&gt;,.('y L;",,, ~o.,.. C •..•p Cl).'l~c.
\.v d-I, ,'" S"" "'pit. F='idd.s
/J1A",be.v- 0+ T""o.nHc...t.:.
SQ,~f/ •. (',dcts

G~n •.•..
o..\ Oi•.td'e&gt;&gt;\

i

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I

X

J

x

X

y;

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WHEAT

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~

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.\:)(

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o

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CORN
WASTELANO

I

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1
:f

~ -

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MILO

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WIH.AT

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x -.
%

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WAST/; LANO

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I

x

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&gt;()(

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K

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l(

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lI'

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&gt;(

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WHGAT

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)(

WASTELANQ

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I i nc.h -:: .....•.S&gt;Oc.-Ft

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)(

Fig. 1.
Turk is
field
indicated.

Pond and vicinity

Fith

sample fields,

general

crop patterns

and number of transects

by

�-111-

Plots were located along a total of 50 transects in seven fields (Fig. 1).
Transects were parallel in four of eight fields and were generally spaced
at 150 foot intervals.
Transect starting points in the three fields north
of the porid (Field Nos. 2, 3 and 4) were equidistant apart.
Transects .Ln
these north fields radiated out at compass settings designed to space the
transects throughout the triangular shaped fields.
Transects in the field
located in the southeast corner of Fig. 1 (Field No.6) had a common starting
point and radiated out at pre-determined compass settings.
All transects
were generally perpendicular to the firing line. All transect starting points
were positioned along field boundaries which were nearest to and generally
paralleled the water's edge.
Plots were one-half meter square (0.5 m sq) in size and located at the origin
of the transect and at 100 foot intervals along the transect.
Transects were
variable in length, but each extended at least 700 feet beyond the firing line.
A few transects in Field No. 6 were terminated at the edge of a wasteland
area less than 700 feet outside the firing line (Fig. 1).
Non-rooted vegetative material, such as corn leaves and stalks, located
within the plots was removed carefully so that lead shot suspended in the
material would fall within the plot. Loose dirt and duff-like surface
material·was moved within the plots in order to expose lead. Lead shot
found were placed in an envelope on which the field, transect and plot
number were recorded.
Notes were taken concerning the "habitat tyrpe" in
which transects occurred and at what point the transects crossed the firing
line.
In.analysis, the numbers of lead pellets were recorded by field, transect
andplot,and
averaged by distances using the firing line as a reference
point. .The shot size of each lead pellet found was also recorded.
No
attempt was made to stratify fields or plots within fields by "habitat type".
All fields used for analysis had definite geographic or habitat type boundaries with the exception of thealfalfa-wheat
fields (Field Nos. 3 and 4)
located directly north of the pond (Fig. 1). In the latter case the field
division was based on the relative number of shot found on the three transects in the east portion of the field (Field No.4),
compared to what was
found on the three transects in the west portion (Field No.3),
thus indicating the influence of the large number of pits in Field No.4.
Samples of dead and live Canada geese having no external visible signs of
gunshot wounds were collected on February 5, February 11, February 21 and
March 5, 1974. An attempt was made to capture at least .ten live birds on
each collection date.
Live birds were captured by hand or with the use of
a net.
Selection of live birds was based almost solely on ease of capture.
No specific sampling scheme was used in the collection of dead birds.
On
each sampling date an attempt was made to collect only birds that had died
a few days prior to that date. The "time of death" judgement was purely
subjective in nature, based primarHyori
the condition of the eye of the
bird and/or the general dryness of the carcass.
On March 5, 1974 an attempt
was made to capture "normal" appearing flightless birds.

�-112-

Birds were transported on the day of collection to the Fort Collins Wildlife·
Research Center and placed in isolation.
Live birds were sacrificed and
necropsied on February 6, 13, and 22, and March 6, 1974. Post mortem examinations were performed either at the C~lorado Sta:te University School of
Veterinary Medicine or at the Wildlife Research Center.
A sample of blood was taken from each live bird at the time of sacrifice.
A subjective judgement as to the general body condition and the status of
body fat was recorded.
Impaction of the proventricular and esophageal
areas, a sign of lead poisoning described by Jordan and Bellrose (1951) for
mallards and by Cook and Trainer (1966) for Canada geese, was noted. Any
gross pathological conditions or other abnormalities were recorded.
A
portion of the liver, a kidney, breast muscle t Lssue and the radius bone
were .collected from each goose to determine lead content· of those tissues.
In addition, the brain, remainder of the liver, a kidney, a section of
breast muscle were taken from every live bird collected and sections of the
heart, the thyroids, spleen, sciatic and vagal nerves, portions of the .intestine, sections of the ventriculus and proventriculus and lung were taken from
most birds sacrificed.
All the above tissues were collected for histopathological examination.
In general terms, the contents of the esophagus, proventriculus and ventriculus were recorded, including lead shot. All shot
found in the ventriculus was collected and labeled as to bird number.
The same routine, with the exception of the collection of tissues for
histopathological
work was followed for birds which were dead at the time of
collection or died between the time of collection and the time of necropsy.
Each tissue collected for analysis of lead content, with the exception of
blood, was oven dryed at 80 degrees centigrade for at least. 72 hours, weighed
and digested in 2 ml HCL04 and 3 ml HN03 in a hot water bath for 6 hours.
Samples were then. diluted to 50 ml and analyzed by atomic absorption spectrophotometry with the deuterium arc background correction.
On January 28 and 29, 1974, 1,092 Canada goose carcasses were removed from
the Turk's Pond area. A random sample of 221 ventriculi (gizzards) were
obtained from the 1,092 Canada goose carcasses were removed from the Turk's
Pond area. A random sample of 221 ventricula (gizzards) were obtained from
the 1,092 carcasses.
These gizzards were examined and the presence of lead
shot reco rded , On February 20 and 21, 1974, 430 carcasses of birds judged
to have died during or shortly after the goose season were removed from
Turk's Pond.
From February 5, 1974 through April 2, 1974 a total 371 geese that were not
considered victims of gunshot wounds, including those collected for examination, were removed from Turk's Pond.
RESULTS AND DISCUSSION
Distribution

of Lead Shot

The sampling technique was designed only to document the general relative
density of surface lead in respect to the firing line. The method resulted
in sampling rates which were fairly uniform, by area, in fields that were

�-113-

rectangular in shape (Field Nos. 1, 5, 7 and 8), but in fields that were
triangular in shape (Field Nos. 2, 3, 4 and 6) the rate decreased continuously as one moved out the transect from the point of origin.
In analysis,
samples were not weighted by surface area and are basically not comparable.
However, the resulting information does provide a general picture of the
lead shot distribution.
On a comparative basis, very few lead pellets were found in fields numbers
one and two located west and northwest of the pond (Table 1 and Fig. 1).
Although no quantitative data were collected, there were indications that
very few birds leaving the pond flew west or northwest.
There apparently
were some flights that returned to the pond from the west (J. Randall, pers.
comm.) .
Available surface lead in fields three and four was influenced by the presence of a perennial crop, alfalfa, between the firing line and the pond
(Fig. 1). The alfalfa field had not been plowed on an annual basis and
therefore, lead pellets on the surface had accumulated, for more than one
hunting season.
The decision to divide the field north of the pond into
two separate fields for analysis was based on the comparatively heavy accumulation of lead pellets in the eastern portion of the field, compared to the
western portion.
These heavy accumulations were no doubt the result, in
part, of the large number of goose hunting pits and potential hunters in the
eastern area. A total of 25 pits from which hunters could distribute lead
pellets in the area were located.
Hunters from only seven pits were in a
position to distribute lead pellets in the western portion of the north
field. In addition, there were apparently substantially more birds which
left the pond flying north-northeast than in a more westerly direction.
According to observers, the major flights of geese leaving and returning to
the pond throughout the hunting season were generally over fields to the
east and southeast.
The presence of lead in fields five and six, and to a
lesser extent, seven, indicate a comparatively heavy amount of gunning
pressure in that area (Table 1).
'
Field ,numbers five through eight were all planted to annual crops and therefore,plowed
sometime after the 1972-73 hunting season, but before the 197374 season. Therefore, most lead on the surface had accumulated during the
1973-74 season (Table 1). These fields, grouped together, provide a picture
of what might be expected, in terms of lead distribution, and in part,
accumulation from one hUnting season's activity around the pond under current
conditions (Table 2). Some obvious changes in relative density of pellets
occur as one moves away from the firing line in either direction (Table 2).
Data from all fields are grouped together in Table 3 to provide an "average"
look at the total area.
It is obvious that many variables were disregarded in the analysis of sur face
lead distribution.
However, data gathering techniques and subsequent
analysis, to date, have shown that lead pellets were available in varying
densities on the surface' of the ground and they were distributed in a some- ~'
what predictable fashion.

�-114;..

Table L Average number of lead pellets found in plots
at 100 foot intervals
along transects in fields sampled around Turk' s Pond.

Approximate
Distance Relative
to the Firing Line

1(10}1/

1,500
1,400
1,300
1,200
1,100
1,000
900
800
700
600
500
400
300
200
100

0.0
0.1
0.0
0.0
0.2
0.1
0.1
0.4
0.0
0.3
0.5
0.5
0.4
0.8
0.1

Firing

'"0

s::
0

P-&lt;
'"0
H

~

0

E-;

Line

100
200
300
400
500
600
700
800
900
1,000
1,100

1/ Number

Firing Line

2(2)

0.0
0.0
0.5
0.0
0.0
0.0
0.5
0.0
0.0
0.0
0.0
0.0

3(3)

4 (3)

5(10)

6(8)

7 (7)

8(8)

0.3
1.3
1.0
1.3
1.3
1.7
1.3
1.3
1.0
0.7
0.3
1.3

0.0
0.0
0.3
0.7
1.7
1.3
2.0
2.3
3.0
3.0
1.3
3.0
4.7
2.3
2.7

0.2
0.2
0.7
0.4
0.3
1.3
1.1
1.2
1.7
2.4
2.4
2.6
1.8
2.4
1.9

0.4
0.3
1.1
0.8
1.6
1.4
2.3
2.5
2.9
3.4
1.9
1.9
1.9

0.1
0.6
0.1
0.6
1.1
1.1
1.6
1.1 .
1.9
1.1
1.4
0.9
i.6

0.1
0.1
0.3
0.3
0.1
0.6
0.6
2.3
1.4
1.9
0.9
1.1
2.6

1.3
0.9·
1.5 . ·1.3
1.6
0.6
1.1
0.9
1.1
0.4
0.7
0.6
0.3
0.3
0.5
0.6
1.3
0.5

0.4
0.1
0.1
0.7
0.6
0.3
0.7
0.6
0.4

-------0.6
0.3
0.6
0.5
0.5
0.3
0.4
0.4

of transects •.

- - - -.- - - - - - - 1.0 . 0.0
1.3
0.5
0.0
0.7
0.0
0.7
0.0
0.3
0.0
0.0
0.0
0.3

1.0
1.7
2.0
1.3
1.3
1.7
0.0
1.7
1.3
··0.3
0.3

2.0
1.2
1.6
1.4
1.3
1.2
0.6
0.6
0.2

�-115-

Table 2. Number of lead shot found in plots at 100 ft intervals along
transects in fields to the south and east of Turk's Pond.

Number of
Shot Observed
12

100 ft

1300 ft

Averages by Interval
200 ft
300 ft
400 ft

500 ft

0.375

11

0.344

15

0.469

0.406
0.531
0.656
25

0.781

33

1.031

35

1.094

50

1.563

0.906
1.229
1.328

2.125

68

2.·172

1\

71

1.319
1.750

500 ft

2.219

2.219
2.313

74

1.969
52

1.625

53

1.656

1.938
1.867
1.719
1.766

60
Firing Line
39

100 ft
- - - - --100 ft

1.875
- ~ - - - - - - - - - - - - - - - - 1.219
1.141

34

1.063

34

1.063

34

1.063

1.115.
1.102
1.063
1.063

28

500 ft

0.903 '

0.903
0.820
0.733

22

0.692
0.536

15

0.554
16
.16

·900 ft

�-116-

�-117-

MOrtality-Lead Poisoning
Necropsy-Gross

Observations

A total of 107 Canada geese were collected at Turk's Pond for examination.
Twenty-seven birds were sacrificed
at the time of necropsy (Table 4).
The
remainder of the birds were either dead at the time of collectioIl or were
collected alive but died prior to necropsy.

Table 4.

Schedule of bird collections

at Turk's

Pond, southeast

Colorado.

Live

Numberof Birds Collected
Dead

Total

February 5, 1974.

3

2#

26

February 11,

1974

4

26

30

February 21, 1974.

l3

18

31

March 5, 1974

7

13Y'

20

Date

};./ Total examination not accomplished on all

birds.

To date, only the number of birds with impacted proventriculi
and/or lead
. shot fragments in their ventriculi
have been tabulated.
The results indicate that 52.9 percent of the birds examined had impacted proventriculi
. (Table 5), 64.5 percent had lead shot fragments in their ventriculi
(Table
6), and 80 percent had either one of the conditions
or both (Table 7).

Table 5. The incidence of impacted proventriculi
in Canada geese examined'
that were collected at Turk's Pond, southeast Colorado.

Date Collected

NumberExamined

Numl:&gt;er
Positive

18

11

February 5, 1974

Percent 'Positive

61.1
t:

February 11, 1974

30

11

February 21, 1974

31

21

85

45

36.7

March 5, 1974

Total

-,

,52.9 .

�-118-

Table 6. The incidence of lead shot fragments in the ventriculi of Canada
geese examined that were collected at Turk's Pond, southeast Colorado.

Date Collected

Number Examined

Number Positive

Percent Positive

February

5, 1974

26

11

42.3

February

11, 1974

30

24

80.0

February

21, 1974

31

21

67.7

March 5, 1974

6

4

66.7

Total

93

60

64.5

Table 7. The incidence of impacted proventriculi and/or lead shot fragments
in the ventriculi of Canada geese examined that were collected at Turk's Pond,
southeast Colorado.

Date Collected

Number Examined

Number Positive

Percent

Positive

February

5, 1974

18

14

77.8

February

11, 1974

30

25

83.3

February

21, 1974

31

25

80.6

March 5, 1974

6

4

66.7

Total

85

68

80.0

Lead Content of Organs and Tissues
Lead levels of organs and tissues taken from 90 Canada geese collected at
Turk's Pond and from ten pen reared control birds are presented in Table 8.
In controlled studies Cook and Trainer (1966) found that the level of lead
in the liver was related to the quantity of lead the bird received, as well
as the length of time the birds retained the pellet. At the time of death,
liver tissues from birds experimentally dosed with from five to 100 No. 4
lead pellets contained from five to 32 ug/g (or ppm) lead. Livers froInsuspected lead poisoned birds collected at Turk's Pond have generally greater
lead concentrations.
Complete analysis of these lead concentration data is
in progress.

�-119-

Table 8. Lead levels, in ppm, in tissues·from Canada geese col.Lect ed at
Turk's Po.nd,so.utheastCo.1orado.,
and from birds raised.artificia11y for
control purposes.
Goose
Sample No.
1
3
5
9
11

13
15
17
19
21

23
25
27
29
31A
33A
34A
35A
30
31
32
33
34
35
36
37
38
39
40
41

Liver

Kidney

40

59

&lt;2
&lt;2

30
148
154

5
149

44

109
148

52
53
54

8
8

28
34
63

296
373
&lt;6
154
76
51
74

73
216

81 .

8

55.

119
84
69
125
160
69
115

43

50
51

38
39
30

99

53

69

49

7
18
3

169
101

124

129

165
160
173
123

157
118 .
73
284
151
97
.103
13
90 .

94

6
9
5

7
4
5

242
95

5
4

138
140
187

14
5
4
4

1784
231

20
83
103
12
35
51
38
36
70
36
43
41

4

1

2

137
101

Remarks

2

119
150
132
&lt;4
93
82
88
118
52
76
77
136
90

Bone

&lt;2.
42

44

42

45
46
47
48

1:./

Tissues
Muscle

115
187

3
6

96

7

62
11

Internal hemorrhage from
lead shot wourid

64

48
29

-------------------------------------------------------------------.----------~

�-120Table 8. Lead levels, in ppm, in tissues from Canada geese collected at
Turk's Pond, southeast Colorado, and from birds raised artificially for
control purposes. (continued).
Goose
Sample No.

Liver

Kidney

55
56
.57
58
59
60
61
63
64
65
66
67

138
82
80
104
76
104
247
87
126
57
105
8

148
107

68
69
70
71
72
73
74
75
76
77
78
79
80
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96

97
80
125
100
130
50
54
75
52
93
79
160
170
51
53
67
122

120
56
104
629
56
88
39
144
6

Tissues
Muscle

Bone

8
6
1,738
14
38

35
238
31
49

5
4
3
9
3
2

76
28
29
60
26
106
13

76
182
44
94
108
116
283
106
62
3

108
73
143
114
128
77
77
66
48
93
128
196
151
105
67
114
109
68
86
199
245
151
87
295
461
76
40
5

7
7
4
4
4
4
5
10
15
7
12
8
19
5
5
3
2,137
4
7
8
2
5
2
7
9
6
6
3

59
34
31
47
37
446
31
35
59
44
33
37
131
29
87
59
60
108
39
76
43
74
79
13

97
98
99
100

5
79
81
3

6
104
110
12

2
2
6
2

26
40
32
14

101

2

3

2

2

Remarks

Collected as a normal
appearing cripple

38
37
48

Collected as a normal
appearing cripple

Collected as a normal
appearing cripple
Collected as a normal
appearing cripple

------------------------------------------------------------------------------

�-121"'-

Table 8. Lea~ levels, in ppm, in tissues from Canada geese collected at
.Turk's Pond, southeast Colorado, and from birds raised artificially for
control purposes (continued).

Goose
Sample No.

Liver

Kidney

102
103
.Cl
C2
C3
C4
C5
C6
C7
C8
C9
ClO

10
66
12
4
2
2
4
20
4
2
2
2

69
48
11
5
4
2
2
11

Tissues
Muscle

6
5

1./ Less than signs «)

Remarks

Bone

2
6
2
5
4
4
3
3
3
2
2
2

17
23
·6
10
8
8

9

Control
Control
Control
Control
Control
Control
Control
Control
Control
Control

are a function of weight of the sample.

MOrtality~Other

Causative

Agents

To date, no infectious disease or other causative agents of disease or poisoning, other than lead poisoning, have been identified in the 1973-74 samples.

Magnitude

of Non-hunting

MOrtality-Preliminary

Results

Gizzards from 221 geese sampled from a population of 1,092 dead birds picked
up around Turk's Pond shortly after the hunting season were examined for the
presence of lead shot fragnients. The majority of these birds died during the
hunting season.· A total of 63 gizzards, or 28.5 percent of the sample,
containeq lead shot.
Sixty-one of the gizzards contained a fragment
from
only one lead shot; one gizzard contained fragments from three lead shot; and
.one gizzard fragments from four lead shot. All pellets from 61 of 63 gizzards
exhibited
considerable amount of erosive wear.
These pellets, both in
quantity of occurrence and wear, resembled those found in gizzards from geese
that were definite victims of lead poisoning.
It was concluded that regardless of the actual cause of death, 311 of the 1,092 birds, or 28.5 percent,
were potential lead poisoning victims because of the presence of lead shot
in their gizzards.
In addition, by tabulating the incidence of lead shot in
the gizzards of those birds that were definite victims of lead poisoning,
according to chemical analysis, we found that only 76.9 percent of those
birds suffering from lead pOisoning had lead shot fragments in their gizzard.
Therefore, an additional 94 of the 1,092 birds may have died of lead poisoning, bringing the potential total deaths caused by lead poisoning to 405 or
37.1 percent of the sample.

a

�-122-

An additional 430 carcasses of birds judged to have died during or shortly
after the goose hunting season at Turk's Pond were removed in late February.
These carcasses were not picked up earlier for various reasons.
It was
concluded that lead poisoning victims in this group occurred at the same rate,
.371, as they occurred in the population of 1,092 birds removed from the.
pond earlier.
Therefore, 160 more birds were categorized as in-season lead
pOisoning victims bringing the .total to 565.
.
A total of 371 live and dead. birds judged not to be victims of gunshot
wounds were removed from Turk's Pond from February 5, 1974 through April 2,
1974 (Table 9). Birds for postmortem
examination and tissue analysis were
selected from this group. Three of the birds collected were selected as
·normal appearing wing cripples and should not be considered as part of the
sample of victims of non-hunting mortality .. Analysis of lead levels in
tissues indicate that approximately 93 percent, or 343 of the 368 birds
removed that were categorized as non-gunshot victims, were victims of lead
poisoning.

Table 9. Sequence of removal of carcasses
be victims of gunshot wounds.

Date

Total
Number
Birds
Removed

Number
Retained for
Examination

of Canada geese judged not to

Remarks

February

5, 1974

26

26

Only birds for examination

removed

February

11, 1974

30

30

Only birds for examination

removed

February

21, 1974

163

18 1/

All carcasses

removed

March 5, 1974

104

13 2/

All carcasses

removed

March

13, 1974

37

0

All carcasses

removed

April

2, 1974

11

0

All .carcasses removed

l/ Includes

one bird selected as a normal appearing

wing cripple.

~

two birds selected as normal appearing

wing cripple.

Includes

Although some mortality continues to occur at Turk's Pond, the rate of death
has slowed considerably.
From March 13 to April 2, 1974, only 11 birds
succumbed for a death rate of about .6 birds/day.
During the previous period,
March 5 to March 13, 1974.birds were dying at the rate of 5.4/day.
Reduction
in the number o f birds using the Turk's Pond area was probably the major
factor in the decline in the observed mortality rate.

�-123-

To summarize, the death of 565 Canada geese at Turk's Pond during or shortly
after the hunting season was attributed to lead poisoning.
An additional
343 birds died of lead poisoning at Turk's prior to April 2, 1974 bringing
total lead poisoning mortalities to 908.
Conclusions
Because of the magnitude of the problem at Turk's Pond and the desire to
implement measures to decrease non-hunting mortality as soon as possible,
some preliminary conclusions are in order.
1.

Of 1,890 carcasses of Canada geese removed from Turk's Pond during the
winter of 1973-74,908
or 48 percent were estimated to be victims or
possible victims of lead poisoning.

2.

Birds are ingesting lead pellets which are distributed on agricultural
lands around Turk's Pond, primarily by hunters shooting from the firing
line.

3.

The ingestion of lead pellets by geese begins prior to the conclusion
of the hunting season.

4.

There is evidence
may be sufficient
Turk's Pond.

to suggest that the ingestion of a single lead pellet
~o cause lead poisoning in geese associated with

Sa. Normal practices associated with annual agricultural crops, such as
plowing and diskingare
currently effective in removing most lead
pellets from the surface of the ground.
However, because of the timing
of these· practices lead is not eliminated during the critical period of
goose habitation.
b. Maintenance of perennial crops such as alfalfa allows accumulation
surface lead for a number of years.
6.

The number of lead pellets distributed in anyone
the number and direction of shots fired.

7.

Lead pellets are distributed
fir:i.ngline.

in a predictable

LITERATURE

of

area is a function

pattern

in respect

of

to the

CITED

Cook, R. S., and D. O. Trainer.
1966. Experimental
Canada geese. J. Wildl. Mgmt. 30(1) :1-8.

lead poisoning

of

Jordan, J. S., and F. C. Be11rose.
1951. Lead poisoning in wild waterfowl.
Ill. Nat. Hist. Survey Div. BioI. Notes No. 26. 27 p.

Prepared

bY?1i~ilg?.j/~
.

Michael R. szym~O
AssistanFWild1ife

&amp;-,7

Researcher

��-125-

October 1974

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~-----------

Project No.

W-88-R-19

Work Plan No.

2

Job Title

Job No.

8

~

~

Investigation of Canada Geese in the San Luis Valley

Period Covered:
Personnel:

Migratory Bird Investigations

April 1, 1973 to March 31, 1974

H. Burdick, B. Clark, R. Desi1et, H. Funk, G. Hinshaw, D. Lowry,
J. Rauch, G. Saville, W. Schuett, M. Szymczak, R. Velarde, R.
Weldon, B. Widhalm, M. Zgainer of Colorado Division of Wildlife;
C. Bryant, J. Sarvis, S. Cornelius of Monte Vista National Wildlife
Refuge; B. Darnell, D. Ziegler of Alamosa National Wildlife Refuge;
R.Tragstaad of Bureau of Land Management.
ABSTRACT

This report presents the results of the first spring-summer season of a field
study concerning the "size, distribution and productivity of the Canada goose
population in the San Luis Valley. Habitat occupied by nesting geese was identified by aerial and ground inspection of all water areas in the valley. Riverbottoms with woody vegetation, ponds, sloughs and islands were preferred, as
were permanent lakes and ponds with Lal.ands : ox emergent:;vegetatioR'" Aerial
counts and ground counts on the same areas were compared to ascertain the
usefulness of count Lng and estimating numbers of geese from a fixed-wing aircraft. Air/ground ratios were consistent from area to area for certain counts
made in open habitat, during mid-day. Most air/ground ratios were inconsistent,
in heavily vegetated riverbottom areas, and could not be used for population
estimates for the entire study area. Aerial brood counts of downy goslings
were not successful. Seventy-three nests were located. Forty-four nests were
on natural sites averaging 18 feet-l inch (5.51 meters) from open water and 15
inches (0.38 meters) above water. Twenty-nine nestswere on artificial nesting
structures either near or in open water. Peak of egg laying occurred from 29
March to 15 April, and peak of hatching occurred from 2 May to 14 May. Mean
number of eggs was 5.05 for 40 nests; 4.82 for 22 successful nests and 5.33 for
18 unsuccessful nests. Hatching success and mean number of eggs hatched was 30
percent and 1.50 eggs for all nests, and 84.6 percent and 4.08 eggs for successful nests. Gosling mortality was at least 10.9 percent on nine different,
isolated areas from date of hatch to flight stage.

�-126-

RECOMMENDATIONS
It is felt that a helicopter instead of a faster moving, fixed~wing
aircraft would allow more reliable aerial counts of geese, especially
along the heavily vegetated riverbottom areas. Counts should be conducted at various daylight hours, depending on weather and on movements
and locations of geese at different times of day during the changing
season.

�-127-

INVESTIGATION OF CANADA GEESE IN THE SAN LUIS VALLEY
Dennis G. Lowry
The resident Canada goose population of the San Luis Valley developed
from restoration efforts by federal and state personnel which began in
1953 and 1955, respectively. Few data have been collected on the size,
distribution and productivity of this expanding population; information
that is basic to satisfactory management of any wildlife population.
This study was designed to provide such knowledge, and this report presents the results of the first spring-summer season of the field study.

P. S. OBJECTIVE
To investigate the size, distribution and productivity of the Canada goose
population in the San Luis Valley during the spring and summer period and
develop a management plan.

SEGMENT OBJECTIVES
1.

To locate, examine, and map the distribution of occupied and potential
Canada goose nesting habitat.

2.

To develop a technique for estimating the annual size and distribution
of the breeding population of Canada geese in the San Luis Valley.

3.

To develop a technique for estimating the annual productivity of
Canada geese in the San Luis Valley.

4.

To compile data, analyze results and prepare a progress report.
METHODS AND MATERIALS

Delineation of Nesting Habitat
Aerial inspection on 20 and 24 April, and ground inspection from early April
to late August allowed the determination and recording of occupied goose
nesting habitat. The Rio Grande River below 8,200 feet (2197 meters) and
all other water areas below 8,100 feet (2469 meters) were searched. Aerial
inspection was from a Cessna 180 aircraft, with the investigator and the
pilot both scanning for geese in and near water, from elevations of 100 to
150 feet (30 to 46 meters) above the ground. Ground inspection was conducted primarily on foot and occasionally from a 4-wheel drive pickup.
Seven-power binoculars and a 20-power spotting scope were used to facilitate
the ground search.

�-128-

Aerial-Ground Counting Techniques
Numbers of adults and subadults found in pairs, singles and groups, plus
numbers of broods observed from a fixed-wing aircraft on selected sample
areas were recorded. Aerial counts were compared with closely-timed
ground counts on the same selected areas in order to evaluate the use of
aerial counts.
Aerial counts were made from a Cessna 180 aircraft at elevations from 100
to 150 feet (30 to 46 meters) above the ground and at speeds from 75 to
100 miles per hour (121 to 161 kilometers per hour). Two observers counted,
scanning the area below both sides of the aircraft and up to .25 mile (402
meters) away. Transects were, (1) systematically counted by making two
passes over each 1 mile-wide area, such that a .5 mile strip would be scanned
on each pass, and making additional passes over areas where sightings were
uncertain; or (2) by spotting geese and then counting, without a predetermined flight pattern.
Ground counts on corresponding areas counted from the air were usually made
by the investigator before or after the flight. However, comparisons of
counts by different teams of ground and aerial observers were tried. Most
corresponding ground and air counts were made within 4 hours of each other
to minimize the chance of geese entering or leaving the area. Air and
ground counts were made during mid-day, a period when geese were flying
the least. Ratios of air-ground counts were tested for consistency using
Chi-square tests.
Breeding pair counts were attempted from the ground but were incomplete due
to the investigator's unfamiliarity with the area early in the season.
Aerial counts of breeding pairs were not attempted for the same reason and
also because of bad weather.

Nesting, Productivity and Survival
Nests were located and examined, and broods were periodically counted to
estimate productivity and mortality. Egg laying dates were estimated by
back-dating from hatching dates, assuming 28 days to be the average incubation period and 1.5 days to be the average time required to lay one egg.
Nests with at least one hatched egg were considered successful. Gosling
mortality was estimated from periodic ground counts on nine different,
isolated areas (Fig. 1) from late-May to late-July.
RESULTS AND DISCUSSION
Delineation of Nesting Habitat
Habitats used for nesting included river ,lake and pond areas. Riverbot toms
with small ponds, sloughs and islands surrounded by willow and cottonwood
were preferred river areas. Permanent lakes and ponds with islands and/or
emergent vegetation were favored open water areas. Areas currently occupied
by nesting Canada geese are indicated in Figure 1.
Nesting is possible in and near all water areas of the San Luis Valley, but
is a matter of speculation. Possibly, additional data from the 1974 nesting
season will help define the most probable potential nesting·areas.

�-129-

N

R~s.5e/l

Lakes

®

LEGEN 0

COh-f: •.•••
eJ
To Have.Nest: "'!J
/=J"eqs

be e.s e

R"eClS

Reporte.d

To /tCl"e. Nest;"J
Gee.s e

x

AreQ~

Ob.s,rv.el

For Go.sl:"'!I
S~\"v,'Vq'

SeRLE.

o

.

IN

5

MILES
1.0

15

•

Mana!\sq

Fig. 1.

Areas occupied by Canada geese in the San Luis Valley, 1973.

�-130-

Aerial-Ground Counting Techniques
Results of air-ground comparison counts of partially- and fully-feathered
geese showed that all air/ground ratios were inconsistent and could not
be used for population estimates for the entire study area.
Chi-square tests were run on selected air-ground counts from heavily vegetated riverbottom areas and from open areas with little or no tall woody
vegetation. Only the open area counts of Table 1, Lines 2, 3, 4, 5, 16,
18 were found to have consistent air/ground ratios from area to area at
the 0.025 significance level. These particular counts were ones which:
(1) the investigator believed no geese had left or entered the area in
the time between corresponding air and ground counts, and (2) the ground
count was considered complete and accurate. The number of geese seen from
the air in these open areas averaged 61 percent of the ground count.
Selected riverbottom counts, using some criteria, were not consistent.
Error in both air and ground counts was probably high. Tall, dense vegetation and high eircraft speeds made aerial counts inaccurate. Ground
counts were difficult due to the above mentioned vegetation and the
secretive nature 'of geese, especially during their flightless molting and
brood-rearing periods from mid-May through July.
Brood counts of downy goslings from the air were not successful; the young
were even less obvious than the adults. Brood counts of downy young are
not included in Table 1. However, partially-to fully-feathered goslings
were recognized, and are included in the June, July and August counts
(Table 1).
Breeding pair counts from the ground were unreliable because, as previously
mentioned under Methods and Materials, the investigator was not familiar
with the area early in the season. Aerial counts of pairs were not available for comparison, as explained previously. However, the investigator
plans on reliable ground and aerial breeding pair counts for the 1974
season, because of last season's knowledge concerning the nesting areas.

Nesting, Productivity and Survival
Data were collected from various areas in the San Luis Valley that nesting
Canada geese were found to inhabit (Fig. 1). The investigator's data do
not include data from the Monte Vista National Wildlife Refuge; however, those
data are labeled and presented separately when comparable.
Nesting Sites
Seventy-three nests were found and studied, with 44 nests on natural sites
and 29 nests on man-made nesting structures (Table 2). TWenty-five nests
were built in bulrush or cattails on either muskrat houses or matted vegetation, located above either water or moist ground. Fourteen nests were
built on islands on bare groUnd using greasewood, driftwood or willow as
cover. TWenty-five nests were found on elevated artificial structures with
straw or hay being the nesting base. Three nests were built on artificial

�Table 1- Comparison of aerial and ground counts of total numbers of partia11y- and fully-feathered geese
in the San Luis Valley, 1973.

Line

Aerial Counts
Area 1./
Date

Count

Air/Ground
Pass '];./ (Percent)

Date

Ground Counts
Area !./

Count

Habitat
Type 11

1

30 May

RL

12

1FA

17

30 May

RL

70 !!./

0

2

30 May

RL

56

2FA

80

30 May

RL

70 !!../

0

3

30 May

SL

27

1FA

63

31 May

SL

43

0

4

30 May

DL

3

1FA

60

29 May

DL

5

0

5

18 Jun

GP

36

1FA

35

18 Jun

GP

102 '2/

0

18 Jun

22

86 'i/

V
V

6

18 Jun

22

14

1FA

7

18 Jun

23

14

1FA

18 Jun

23

8

18 Jun

26

22

1FA

18 Jun

26

o '2/
o '}../

18 Jun

27

25

1FA

18 Jun

27

o '2./

V

9
10

18 Jun

29

31

1FA

18 Jun

29

o '2./

V

11

18 Jun

30

2

1FA

18 Jun

30

0

V

12

18 Jun

31

6

1FA

120

18 Jun

31

5 '2/

V

13

9 Ju1

22

39

1FA

33

9 Ju1

22

119

V

14

9 Ju1

AL

25

1FA

89

3 Ju1

AL

28

0

15

13 Aug

113

22

1A

110

13 Aug

113

20

0

16

13 Aug

113

20

1F

100

13 Aug

113

20

0

17

13 Aug

22

19

1FA

16

13 Aug

22

120

V

V

-------------------------------------------------------------------------------------------------------------

I
I-'
Vol
I-'
I

�Table 1. Comparison of aerial and ground counts of total numbers of partially- and fully-feathered
in the San Luis Valley, 1973 (continued).

geese

Date

Aerial Counts
Area·l]

Count

Pass '27

Air/Ground
(Percent)

Date

Ground Counts
Area 17

Count;

Habitat
Type 31

18

13 Aug

GP

49

lFA

65

13 Aug

GP

75

192/

0

15 Aug

III

26

lFA

39

15 Aug

111

66

V

202/

15 Aug

39

lA

42

15 Aug

21 2..1

V

37

2A

40

15 Aug

112
112

93

15 Aug

112
112

93

V

22 ~/

15 Aug

112

47

IF

51

15 Aug

23 2..1

112

93

V

15 Aug

112

31

2F

33

15 Aug

24 2./

112

93

V

15 Aug

113

25

lFA

357

15 Aug

113

7

0

25 2..1

15 Aug

GP

86

lFA

115

15 Aug

GP

75

0

26 ~/

15 Aug

GP

84

2FA

112

15 Aug

GP

75

0

27 &amp;/

22 Aug

34

118

lFA

78

22 Aug

34

151

V

28

22 Aug

22

10

lFA

63

22 Aug

22

16

V

Ll.ne

11

.

.

- RL = Russell Lakes; SL = San Luis Lake; DL = Dry Lakes; AL = Adams Lake; GP= Goose Pen Area at Monte
Vista National Wildlife Refuge; All Numbers = Various Rio Grande riverbottom areas between Monte Vista
and Alamosa.

21
- Number indicates first or second pass with aircraft; Letter indicates count by fore or aft observer or
by both.

llV = heavily vegetated

riverbottom; 0 = open habitat.

i/Ground count of 30 May was estimated; all others were actual counts.

51

..

- Ground counts of lines 5 - 10 and 12 were not by same observer who made aerial counts; other counts were
by same observer.
i/For the aerial counts in lines 19 - 27, geese were spotted and then counted without a predetermined
pattern; all other aerial counts were made by flying predetermined transects.

flight

,...
I

w
!-..)
I

�-133-

Table 2.

Nesting sites of rknown..
nests in the San Luis Valley, 1973. 1..!
Description· of
Nest Base

Number of
Nests

Natural

Vegetation Above of Near Water

25

Natural

Bare Ground of Island

14

Natural

Haystack

2

Natural

Gravel Beach

1

Natural

Duck-hunting Blind

1

Natural

Bare Ground Under Log

1

Type of
Nest SiteJ:...!

44

Sub-total

Artificial

Hay or Straw, Elevated Above
Land

17

Hay or Straw, Elevated Above
Water

8

Artificial

Wooden Floating Structure

3

Artificial

Beached, Wooden Floating
Structure

1

Artificial

Sub-total

29

Total

73

1/
-Data do not include sites of nests at Monte Vista National Wildlife
Refuge, 1973.
J:...!
Artificial sites are considered only those structures specifically constructed for goose nesting.

�-134-

floating structures with straw or emergents used as nesting material.
Haystacks, a gravel beach, a fallen log, a beached floating structure
and a duck hunting blind provided sites for the remaining six nests.
Natural nests averaged 18 feet-l inch (5.51 meters) away from open water,
and ranged from 1 foot-6 inches to 55 feet-lO inches (0.46 to 17.02 meters)
away. Twenty-five of these 44 nests were 14 feet (4.27 meters) or less
away from open water.
Eggs in natural nests averaged 15 inches (0.38 meter) above water, and
ranged from 2 to 64 inches (0.05 to 1.63 meters) above water. Half of
these nests held eggs 12 inches (0.31 meter) or less above water.
Occupied elevated, man-made structures were located either in open water
or up to 25 feet (7.62 meters) away from open water. Those on land accounted
for more than two-thirds of the occupied sites and averaged 7 feet-lO inches
(2.39 meters) awa~ from open water.
Eggs in the 17 artificial, standing structures on land averaged 6 feet-8
inches (2.03 meters) above ground, and ranged from 10 feet to 5 feet-4 inches
(3.05 to 1.63 meters) above ground. More than four-fifths of these nests
held eggs between 5 and 7 feet (1.52 and 2.13 meters) above ground. Eggs in
the eight artificial, standing structures in water averaged 4 feet-lO inches
(1.47 meters) above water, and ranged from 4 feet-l inch to 6 feet-2 inches
(1.25 to 1.88 meters) above water.
Nesting Chronology
Egg laying began on 24 March and continued through 22 May, with 29 March
through 15 April the peak egg-laying period. Hatching began on 29 April
and continued through 20 June, with 2 May through 14 May the peak hatching
period.

Clutch Sizes
Although 73 nests were studied, the clutch sizes for only 40 nests were
determined. The remaining nests were either hatched or destroyed before
the clutch size could be accurately determined. Based on these 40 nests,
the mean clutch size was 5.05 eggs and the mode was 5 eggs.
The mean clutch size of the 22 successful nests was 4.82 eggs and the mode
was 5 eggs. In the 18 unsuccessful nests, the mean clutch size was 5.33
eggs and the mode was 6 eggs. Mean clutch size in 29 nests at the Monte
Vista National Wildlife Refuge was 4.52 eggs (J. Sarvis, unpublished data).
Nesting Fate
All 73 nests studied were classified as to fate since destroyed egg remains
differ in appearance from hatched eggs. Of these 73 nests, 27 (36.9%) were
successful and 46 (63.1%) were failures. Success rates varied from area to

�-135-

area (Table 3). Thirty';"nineof the nests that failed were apparently
destroyed by avian or mammalian predators and the remaining seven were
deserted.
Of the 79 nests studied at Monte Vista National Wildlife Refuge (Table
4), 52 (65.8%) were successful, 25 (31.6%) were failures and 2 (2.6%)
were unknown as to fate. Seventeen of the nests that failed were destroyed, nine by mammalian predators and eight by avian predators. Of
the remaining nests that failed, four were abandoned, two were flooded
and two were eventually deserted due to chilled or addled eggs (J. Sarvis,
unpublished data).

Hatching Success
Using the data from the 40 nests in which clutch sizes were determined,
the percent of eggs hatching in all 73 nests studied was estimated at 30
percent, with a mean hatch of 1.50 eggs per nest. The 27 successful nests
had an estimated 84.6 percent of the eggs hatch, with a mean hatch of 4.08
eggs per nest. Percent of eggs hatching in all 79 nests studied at Monte
Vista National Wildlife Refuge was estimated at 52 percent, with a mean
hatch of 2.34 eggs per nest (J. Sarvis, unpublished data).
Gosling Mortality
On nine different, isolated areas (Fig. 1) at least 129 goslings were
produced and 115 goslings remained when the birds reached flight stage.
Gosling mortality in these areas was estimated to be at least 10.9 percent.

f,

Prepared by

,~,

.

=-:-0. __ &gt;

' '"&gt;

,&gt;

'- __ '}.~----:---

'~

-

Dennis G. Lowry
Graduate Student

&gt;

�-l36Table 3.

Nesting fate of known nests in the San Luis Valley, 1973. !I

Area and Type of
Nest Structure Y

Number
of Nests

.Successful Nests
Number
Percent

Rio Grande Management .Area
Artificial

20

0

0.0

Natural

3

1

33.3

Total

23

1

4.3

1

50.0

Natural

2
22

14

63.6

Total

24

15

62.5

Artificial

7

2

28.6

Natural

19

9

47.4

Total

26

11

42.3

Artificial

29

3

10.3

Natural

44

24

54.5

Total

73

27

36.9

Russell Lakes
Artificial

Other Areas

Total Area

1/
- Data do not include fate of nesting geese at the Monte Vista National
Wildlife Refuge, 1973. These data are presented in Table 4.
2/
- Artificial sites are considered only those structures specifically constructed for goose nesting.

Table 4. Nesting fate of known nests at Monte Vista National Wildlife
Refuge, 1973 (J. Sarvis, unpublished data).
Type of
Nest Structure!/

Number
of Nests

Successful Nests
Number
Percent

·Artificial

28

21

75.0

Natural

51

31

60.8

Total

79

52

65.8

1/
- Artificial sites are considered only those structures specifically constructed for goose nesting.

�October 1974

-137-

JOB PROGRESS
REPORT

State

of

Project

.::.CO::..LO=RAD=:.;O~
__ ...,..._,-No.

Job Title

Migratory Bird Investigations

W-88-R-19

InvestigatHmof

.

Period

Covered: . April
.

..

..

'"

Units of Eastern

Mallard
.

.:

...

'

-,'

"

-

....

-

.,',

",

:....

-'

1, 1973 through March 31, 1974
..

'.

,"

,"

,"

,"",

":-,.',

".:-,

Colorado

-:.

::.

Personnel:
C. Braun, J.Carsella,
R. Clark~J.
Corey , D.Coven, C. Crawford,
M. DePra, G.East,G.
Eyre,R.Forbes~H.FUn.k,B.Goetze,
.J. Gustafson,
.
A. Hemmert, T. Henry, J. Hicks, J.Jackson,
D.Kenvin, jr.,
R. Kitzmiller,
C. Leonard, J~ Lewis, G.Lorentzson,
R.Mason, R. Oakleaf, P.Olson,B.
Peterson, W. Russell, G. Saville,
II.. Schul.tz., G. Smith, Jr.,
M. Szymczak,
aridR.Hoppet.
ABSTRACT
Poat.-aeason mallard trapping efforts yielded 5, 614 banded birds in nf.ne study
units during the winter of 1973;.,.74.; Banding quotas of 600:-800ma.1lards were
met or exceeded in all but:, 'the three 001tsofsolltheastern
Colorado. . Mallard
numbers appeared tobegenerally
lower ill easternC()loradoduring
1973';':'74,·
as compared to the previous year •. Peak numbers occurred between mid~N6vember
and mid-December., A firial analysis of all band recovery data relative
to '
this study will be iriitiatedin
the next Segment.

��-139-

INVESTIGATION OF MALLARD MANAGEMENT
UNITS OF EASTERN COLORADO
RichardM.

Hopper

Periodic aerial surveys and post-season banding of wintering populations of
mallards were continued in eastern Colorado in 1972-73. Past contributions
of this study as part of a cooperative venture with other Central Flyway
States were reviewed in the previous progress report (Hopper 1973). The
present status of this study as it relates to Colorado was also discussed
in this same report. The present report for the most part, covers results
of banding and survey data collected during the Segment.
P. S. OBJECTIVE
To develop a harvest formula for eastern Colorado management units.
SEGMENT OBJECTIVES
1.

To analyze, for each of nine eastern Colorado study areas of wintering
mallards, all data collected from 1962-63 through the 1972-73 hunting
season on the discreteness of populations and publish results in an
appropriate journal.

2.

To determine the feasibility of requesting special hunting regulations
for individual mallard management units within Colorado.

3.

To trap,band, age, and sex samples of mallards during winter in each
management unit of study, based on results of final analysis, of
sufficient size to allow adequate monitoring of each population.
METHODS AND MATERIAL S

Cage traps of the Salt Plains type were employed to capture most mallards,
although cannon,....nets
were also used in the Arkansas Valley. Trapping operations began inunediately following the end of the duck hunting season on
January 16, 1974 and continued through February 15, 1974. Banding was
accomplished in nine units of eastern Colorado, including five in the South
Platte Valley, three in the Arkansas Valley, and one at Bonny Reservoir. All
birds were aged by use of the wing according to Carney (1964) and Hopper and
Funk (1970).
.
Periodic aerial censuses were again conducted during the fall and winter to
estimate movements and numbers of birds present in eastern Colorado. Sex
ratio counts were not conducted during the 1973-74 period.

�-140-

RESULTS AND DISCUSSION
Trapping and Banding

Banding quotas of 600-800 mallards were established for each on nine waterfowl management units (Fig. 1), and these were met or exceeded in all but
three units (Table 1). Banding activities were highly successful at Bonny
Reservoir and in the five units of northeastern and northcentral Colorado,
despite relatively mild weather during the trapping period. Mild weather
has always created problems in attaining banding quotas in southeastern
Colorado, and this year was no exception.
Trapping efforts resulted in the banding of 5,614 mallards in the nine study
units (Table 1). Attempts to band equal numbers of birds by age and sex
were mostly successful. Overall samples by age and sex were more equal than
those obtained during the previous year. Females were again the most difficult sex to obtain, with success by age varying from area to area.
Winter Aerial Surveys
Dates and results of the periodic aerial counts are shown by management unit
in Table 2. These counts were made rather inconsistently, with the midwinter survey (January) being the only one that included counts of all areas
at approximately the same time. A high of 222,407 ducks were counted during
the mid-December period, but the mid-November count may have been higher if
all areas had been surveyed at that time. It is difficult to compare survey
results between years, but it appeared that numbers of ducks were generally
lower in 1973-74 than in the previous year. The January Inventory of 1974
(160,665) was considerably below that of 1973 (233,038). Variables such as
weather and flying conditions make it hard to interpret results between years.
Analysis of Band Recovery Data
No analysis of band recovery data was conducted during Segment 19. It now
appears that the cohort of birds from the first year of banding (1963-64) is
nearly eliminated from the population. Thus, a final analysis will be initiated next Segment to include recovery data from the 1964-65 through the 197374 hunting seasons. Existing programs will be utilized in the analysis,
along with new ones developed for more recent methods of analysis. A program
for the Seber Method (Seber 1970) of estimating survival will be available
from the Bureau of Sport Fisheries and Wildlife by the summer of 1974. This
analysis method is very promising and will play a major role in the analysis
of the band recovery data in question.

�WATERFOWL
\\

(;r 1'- R-\j- J

Y

0

~., ,

!\;

MANAGEMENT

UNITS

"

F

B

R

1\

c,

K

A

G

Waterfowl managem~nt Units.

,\

,&lt;

I

"

HUM

A

�'Table 1 • Numbe.rsoaaa percentages of mallards in the banded sample by location, age and sex, eastern Colorado,
.1973-74.
Immature

Adult
Male
Number
Percent

Female
Number
Percent

18.6

257

27.1

262

27.7

212

26.6

205

25.8

173

21.7

24.8

146

18.4

209

26.4

241

30.4

153

25.7

112

18.8

156

26.2

174

29.2

797

200

25.1

204

25.6

228

28.6

165

20.7

869

263

30.3

235

27.0

224

25.8

147

16.9

(11) Two Butt:es~e'n'0f.r 498

154

30.9

214

43.0

47

9.4

83

16.7

(12) Rocky Ford-Lamar

38

20

52.6

6

15.8

8

21.0

4

10.5

(13) Pueb1o-RoekyFord

281

124

44.1

91

32.4

37

13.2

29

10.3

5,614

1,569

27.9

1,396

24.9

1,371

24.4

1,278

22.8

Male
Percent
Number

Female
Number
Percent

947

252

26.6

176

(2) Ft_M::&gt;rg••~Ste;r:ung :796

206

25.9

(3) Gree1 ey-Ft:,.
l(o:rgan

79.3

197

(4) Ft",Co1JJlns

595

(6) Denver-Greel~y
(9) Bonny Rese.nroir

Number
Managemen t Unil:
(1) Sterling-Julesburg

.Banded

I

I-'
.pN

Totals

I

�-143-

Table 2. Aerial duck counts by date interval and study unit, eastern
Colorado, 1973-74.

Management Unit

November
5-6

(1) Sterling-Julesburg

Number of Ducks Counted
November
December
January
16-17
10, 19-20
7, 9, 11, 13
35,000

14,450

6,175

(2) Ft. Morgan-Sterling

31,500

28,000

29,500

30,050

(3) Gree1ey-Ft. Morgan

35,950

42,000

31,000

31,150

(4) Ft. Collins

11,400

60,300

9,550

27,900

(6) Denver-Greeley

13,250

1,000

23,000

9,375

(9) Bonny Reservoir

52,000

10,000

(10-13) Arkansas Valley

56,750

17,400

6,157

28,615

222,407

160,665

(15) San Luis Valley
Totals

8,68(}~J

92,100

1/ Coun t made on November 27.

174,980

�-144-

LITERATURE

CITED

Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification by means of wing plumage. U. S. Fish and Wildl. Serv., Spec.
Sci.Rept. :Wildl. No. 82. 47 p.
Hopper, R. M.
Colorado.

1973. Investigation of mallard management units of eastern
Colo. Div , of Wildl., Game Res. Rept. Oct. In Press.

Hopper, R. M., and H. D. Funk. 1970. Reliability of the mallard wing agedeterminat ion technique for field use. J. Wildl. Manage. 34 (2) :333- 339.
Seber, G.A.F.1970.
Estimating time-specific survival and reporting rates
for adult birds from band returns. Biometrika 57(2):313-318.

Prepared bY_-'~4/r.?"":'::'/-::W1:h&amp;;...jo4ol·
~.......:.o
Jg.;,/,-7 .L.2:
.•Z~-_0.L.
'J6",-"G~3.M?_U""-J«,-,--"-__
Richard M. Hopper {}
Wildlife Researcher

�-145-

JOB PROGRESS

Stat e 0 f __ --'-_....:C:..:O~J.:..:.O.:;R=.:A:.::D.:::O
Project

No.

Work Plan No•

W-88-R~19
4

October 1974

I~EPORT

__.,.--

Migratory Bird ,Investigations
Job No.

.Iob Title

Trapping and Banding, Doves

Period Covered:

April 1, 1973 through March 31, 1974

3

~

Pe:rsonne1: Jack Randall, Bureau of Sport Fisheries and Wildlife; W. John
Arthur, Clait E. Braun, Jack Corey, HowardD. Funk, James o. Goodyear, J.
Edward Kautz, Marie Vendevil1eKautz,
Charles W. Loeffler,
Ronald Oakleaf,
Brett Petersen, Brent Smith; Darryl J. Todd, Jack Vayhinger, and Kenneth
C. Wagner, Colorado Division of Wildlife.
ABSTRACT
Efforts initiated
in, 1964 to trap and band samples of mourning doves
(Zenaida macroura) in three geographic areas of Colorado were -continued in
1973. Cooperative Federal and State efforts resulted in 4,399 birds being
newly banded. Of this total;
1,822 were innnatures, 1,341 were adult males,
1,148 were adult females, while age and sex were not determined for 88 birds.
Assigned quotas were accomplished for all areas except extreme eastern Colorado ~ Wing molt data from trapped and harvested immatures suggested that
nesting was later; in 1973 than in 1972, with hatching peaks in late May,
late June, late July, and late August. A sample of 2,960 hunter-harvested
birds, all from east of the Rocky Mountains, gave a young-to-oldratio
of
1: 1. Wing molt data from harvestedimmatures
in this, sample indicated that
most early-hatched young had migrated from Colorado prior to September l.
Ff.f ty=one iband recoveries
from doves banded by Division personnel were
reported to date in the 1973 recovery year.
Twenty~nine (56.9 percent) had
been banded in 1973. Of the recoveries, 27 (53. o percent) were recovered
in CoLo rado , while 8 (15.7 percent) were from Texas, with the remainder
being from scattered locations mostly south and west of Colorado.
Apparent
hunting mortality for doves banded in Colorado continues to be less than 3
percent.

_

�-146RECOMMENDATIONS
1.

It is recommendedthat the field portion of this study be extended
through 1974 in order to contribute to the overall Central Management
Unit Dove banding project.

2.

It is recommended that this project be held inactive in 1975 and 1976
with the final· report: to be written once most recoveries have been
received, probably in 1977 or 1978.

�-147-

MOURNING DOVE TRAPPING AND BANDING
C1ait E. Braun

Intensive efforts to trap and band mourning doves in Colorado initiated in
1964 were continued in 1973 in cooperation with personnel from theU. S.
Fish and Wildlife Service. Data presented in this report are those collected in 1973, the tenth year of this continuing investigation.
P. S. OBJECTIVE
To investigate migration patterns and mortality rates of mourning doves
banded in Colorado by age, sex, and area.

SEGMENT OBJECTIVES
1.

To trap and band mourning doves in three selected areas of Colorado for
the purpose of obtaining migration, life history, and annual mortality
data.

2.

To estimate harvest size and hunter success.
METHODS AND MATERIALS

Methods and materials used in 1973 were similar to those described in earlier
reports and summarized by Braun (1970).
Description of Trapping Sites
Most trapping sites in Eastern Slope areas were identical or similar to those
described earlier (Braun 1970). One new site was utilized in 1973 east of
the Continental Divide, in addition to those also trapped in 1972. This site
was near Salida where trapping was conducted along edges of fields planted
to wheat (Triticum aestivum) and alfalfa (Medicago sativa). West slope trap
sites were essentially those also used in 1972.
RESULTS AND DISCUSSION
Trapping and Banding
Cooperative Federal and State trapping and banding efforts resulted in 4,399
doves being newly banded in 1973. Doves were banded at five major sites on
the eastern slope, two sites on the eastern plains and at two major sites

�-148-

west of the Continental Divide. The banding goal of 4,000 birds was essentially accomplished as 1,000 were banded in western Colorado, somewhat less
than 1,000 in extreme eastern Colorado and over 2,000 ~ere banded along the
eastern slope of the Rockies. Sex and age distribution of birds banded by
area are presented in Table 1.
Slightly fewer inunature doves than adults were banded in 1973, a change from
the situation prevailing in 1972, but similar to most previous years of the
study. Production in 1973 was believed to be only fair until late sUIlUller,
possibly explaining the lower percentage of immatures trapped and banded.
However, differences in numbers of adults and immatures trapped is undoubtedly
a function of time of season of trapping effort. Trapping at Durango, Montrose,
Salida and Monte Vista was accomplished primarily in June and early July,
while trapping at Ft. Morgan and Denver was conducted principally in late
July and early August. As in previous years of the study, a higher percentage
of males were trapped than females (31.1 to 26.6 percent). This disparity is
probably related to timing of incubation and brood sharing, as females would
normally be exposed to only one trapping period per day.
During the 1973 trapping period, 83 mourning doves were recaptured that had
been banded in previous years. Eighty-one returned to the area where they
were initially banded, while one moved from Ft. Garland to Montrose and one
was banded out of Colorado (location still unknown). Of the eighty-two doves
banded in Colorado, 2 were banded in 1968, 5 were banded in 1970, 17 were
banded in 1971, while 58 were banded in 1972.

Hatching Data
Wing molt data were available for 1,815 of the 1,822 immature doves trapped
and banded in 1973, and from 1,509 hunter harvested immatures. Estimated
hatching dates (Allen 1963) are presented in Tables 2 and 3. Examination
of data in Table 2 indicates that few mourning doves trapped in Colorado hatched
prior to May 1, with most hatching in late May and mid- to late June. While
definite hatching peaks are difficult to ascertain from data in Table 2,
peaks are suggested in late May and late June. Timing of hatching based on
trapped samples in 1973 was later than in 1972 (Braun 1973). Since trapping
ceases in early to mid-August, progeny of late nests are not represented in
Table 2. This can clearly be seen upon examining Table 3, as two peaks can
be ascertained (late July and late August). When hatching dates of hunter
harvested doves in 1972 and 1973 are compared, it is obvious that nesting
continued later in 1973 that 1972 (Braun 1973). Data in Table 3 suggest
that early hatched doves have migrated from Colorado prior to 1 September.
It is apparent that neither trapping or hunting data alone accurately
represent actual hatching dates as one method (trapping) fails to show late
nesting, while the other (hunter harvest in Colorado) misrepresents production early in the nesting period. It should be remembered that neither
trap or hunter samples may actually represent doves produced in Colorado.
Harvest
Efforts were again made to collect wings from hunter-harvested doves throughout Colorado during the 1973 hunting season. Through this effort 2,960

�-149-

wings were classified as to age (adult or immature) and primary molt. All
wings were from eastern and east slope areas of Colorado with the largest
samples from Ft. Carson (2,743) and the Pritchett area (189). All wings
were from September to early October with about one-third (32.3 percent)
being taken after 21 September. Immature to adult ratios varied from a
high of 2.5:1 at Pritchet (N = 189), and Louviers (N = 28) to 1:1 at Ft.
Carson (N= 2,743). At Ft. Carson the ratio of immatures to adults in the
harvest started off low (.8:1), decreased during the second and third week
of the season (.6:1 and .5:1), and then increased late in the season to 2.4:1
and 2.6:1.
Harvest data obtained through use of a mail survey of hunters after the 1973
hunting season are not presently available. Final results of this survey
are normally not ready for distribution until 30 June of the year following
the season (H. Riffel, personal communication).
To date, recoveries of 51 mourning doves banded by Division personnel have
been reported in the 1973 recovery year (June 1, 1973 to May 31, 1974). An
additional 9 recoveries from the 1972 recovery year have been received since
preparation of the 1972 progress report. All but two of these 9 late
recoveries were from outside of Colorado, with 5 being from Mexico and one
each from Texas and Arizona. Six were first year recoveries (banded in
1972), while two were banded in 1971 and one was initially banded in 1969.
Eight of the nine were from bandings east of the Continental Divide, while
only one was banded in western Colorado.
Of the 51 recoveries reported in the 1973 recovery year, one was banded in
1968, two in 1969, three in 1970, 9 in 1971, 7 in 1972, while 29 were first
year recoveries (banded in 1973). All but 4 were shot recoveries. Eleven
of the 51 recoveries were banded west of the Continental Divide, with four
being recovered in western New Mexico, three in Mexico and one each in Utah,
california, western Colorado and eastern Colorado. These data are similar
to those gathered in previous years and suggest that doves banded west of
the Continental Divide have little affinity for areas east of the
Continental Divide. Considering all recoveries, 27 (52.9 percent) were from
Colorado, 8 (15.7 percent) were from Texas,S each (9.8 percent) were taken
in New Mexico and Mexico, with 2 recovered in Utah and one each in Minnesota,
Kansas, California and Arizona. Examining first year recoveries only, 10
of 29 were banded as immatures (34.5 percent). This is less than the proportion of immatures banded in 1973 (42.2 percent), suggesting that adults
and juveniles do not have differential vulnerability to hunting pressure,
but do migrate at different times. All present indications are that hunting
is not a serious mortality factor of doves banded in Colorado, as recovery
rates are less than 3 percent of all doves banded in a given year, or of a
particular age class banded in the same year.
Six doves banded in other states were reported recovered in Colorado in 1973.
Three were banded in South Dakota, while one each was banded in Texas,
Wyoming, and Montana. Four of the six doves were banded as immatures.

�-150-

Table 1. Sex and age distribution of mourning doves banded by area in
Colorado, 1973.

Area

Adult Male
No.
Banded Percent

Adult Female
~.
Banded Percent

Immature
No.
Banded Percent

Total

Eastern Colorado
Ft. Morgan

60

12.0

130

26.0

310

62.0

500

Vineland

64

33.3

35

18.2

93

48.5

192

Subtotal

124

17.9

165

23.8

403

58.3

78o!-'

Denver

95

10.3

87

9.5

738

80.2

920

Monte Vista

217

37.8

192

33.4

165

28.8

574

Ft. Garland

160

32.0

134

26.8

206

41.2

500

Ft. Collins Area

203

40.6

118

23.6

179

35.8

500

Salida

70

56.0

45

36.0

10

8.0

125

Subtotal

745

28.4

576

22.0

1,298

49.6

2,619

Durango

284

56.8

210

42.0

6

1.2

500

Montrose

188

37.6

197

39.4

115

23.0

500

Subtotal

472

47.2

407

40.7

121

12.1

1,000

1,341

31.1

1,148

26.6

1,822

42.2

4,399~/

Eastern Slope

Western Colorado

Total All Areas

~/ Including 88 unknown age and sex mourning doves banded at Vineland.

. '.'

�Table 2.

Estimated hatching dates for wild-trapped

immature mourning doves, 1973.

June
5"""11 12-18

19-25

26-Ju1y 2

3-9

Ju1l
10-16

17-23

260

220

239

293

237

77

0

0

14.3

12.1

13.1

16.1

13.0

4.2

0.0

0.0

Prior to
May 1

1-7

8-14

Ma
15-21

22-28

29-June 4

Number hatching

4

14

53

126

292

Percent of total

0.2

0.7

2.9

6.9

16.0

Table 3.

Estimated hatching dates for hunter-harvested

immature mourning doves, 1973.

...•
...•
I

V1

Prior to
June 12

June
12-18 19-25

26-Ju1y 2

3-9

Ju1
10-16

17-23

24-30

31-Aug. 6

I

August
7-13 14-20

21-27

SeEt.
28-Sept. 3

Number hatching

7

50

90

28

131

163

102

226

206

137

118

194

57

Percent of total

0.5

3.3

6.0

1.9

8.7

10.8

6.8

15.0

13.7

9.1

7.8

12.9

3.8

�-152-

LITERATURE CITED
Allen, J. M. 1963. Primary feather molt rate of wild immature doves in
Indiana. Ind. Dept. Conserv., Game Res. Sect. Circ. No.4.
Indianapo1is. 4 p.
Braun, C. E. 1970. Mourning dove trapping and banding. Colo. Div. Game,
. Fish and Parks, Game Res. Rept., Fed. Aid Proj. W-88-R. Oct.
p. 143-149.
1973. Mourning dove trapping and banding.
Game Res. Rept., Fed. Aid Proj. W-88-R. Oct.

Prepared

Colo. Div. Wi1dl.,
In Press.

by_---:::~tIu~~/.~I~.__f_L~~~--C1ait E. Braun
Wildlife Researcher

�-153-

October 1974

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-88-R-19

\~ork Plan No.

4

Job Title
Period Covered:

Migratory Bird Investigations
Job No.

4

~

~--~-

Band-tailed Pigeon Investigations
April 1, 1973 through March 31, 1974

Personnel: Clait Braun, Ron Desilet, Richard Fentzlaff, Howard Funk, J. Edward
Kautz, Marie Vendeville Kautz; Harold Lanning, Ann Leckler, Richard McDonald,
Kris Moser, Ron Oakleaf, Robert Rouse, Brent Smith, Darryl Todd, Willie
Travnicek,Ron Velarde, Ken Wagner, Pat Waters, J. Allen White, Mike Zgainer,
Wildlife Conservation Officers and other personnel of the Colorado Division of
Wildlife.
ABSTR..I\Cr
Investigations of band-tailed pigeons (Columba fasciata) initiated in Colorado
in conjunction with the Four Corners States Cooperative Study were·continued
in 1973. Major emphasis was placed on banding and continuation of analyses
of hunting statistics. Pigeons were trapped at 21 different locations in·1973
with 3,559 birds (2,812 adults, 177 subadu.l.t s, and 570 immatures) being newly
banded. Over one thousand (1,017) pigeons were retrapped, including 7 banded
outside of Colorado (6 'from New Mexico and 1 from Utah). During the hunting
season, 455 permits were issued, with 212 individuals actually hunting. Only
60 hunters were successful, each harvesting an average of 5.5 pigeons. Total
estimated harvest (including crippling loss) projected from the questionnaire
survey (92.1 percent response) was 366 birds, less than one-half of the number
harvested in 1972. Crippling loss .approxfmated 10 percent of the birds shot
and retrieved. Wings were received from 44 successful hunters. Immature and
subadult pigeons comprised 22.9 percent of the 236 baridtailwings recieved.
Field checks of 122 (95 adults, 27 innnatures) hunter-killed birds indicated
that 63.2 percent of the adults were still involved with nest activities and/
or feeding of young. Recoveries were received from 92 pigeons banded in Colorado. Most (67) were recovered in Colorado, with 17 being recovered in New
Mexico, 4 in Arizona, 3 in Mexico and 1 in California. At least 18.2 percent
of the birds harvested during the hunting season in Colorado were banded. one
pigeon banded in Arizona was harvested in Colorado during the 1973 season.
Hatching dates were calculated for 623 pigeons with lIDst hatching between 12
May and 19 August. Helminth infections continue to be low (13.9 percent),
with females being more frequently parasitized than males.

�-154RECOMMENDATIONS

1.

It is recommended that this project be extended for completion of banding
objectives through 1974 and for analyses of data and publication of
results through 1975.

2.

It is recommended
be continued.

that the hunting

season, permits and harvest

surveys

�-155-

BAND-TAILED PIGEON INVESTIGATIONS
C1ait E. Braun
Intensive ecological studies of wild band-tailed pigeons which nest and
reside in Colorado from late April to early November, initiated in 1969,
continued in 1973. Investigations in Colorado represent a portion of ~he
regionwide Four Corners Cooperative Band-tailed Pigeon Investigation which
was initiated in Arizona in 1967. Data presented in this report are those
collected in 1973, the fifth year of this investigation.

P. S. OBJECTIVE
To investigate life history and ecological facts pertinent to band-tailed
pigeons in Colorado as follows: range; relative numbers; relation between
numbers of birds present by area and such physical factors as weather, food,
vegetative cover, elevation, and salt; breeding and nesting chronology;
migration patterns and chronology; harvest patterns; and mortality rates.
SEGMENT OBJECTIVES
1.

To trap, band, age, and sex as many birds as possible in selected areas
of Colorado to investigate migration patterns and chronology of Colorado
bandtai1s throughout their range.

2.

To conduct a hunting season, gather harvest data, and evaluate results.

3.

Compile and initiate analyses of all data collected in Colorado.
ate in preparing final report for Four Corners States.

Cooper-

METHODS AND MATERIALS
Methods and materials used in 1973 were identical to those in previous years
of the study (Braun 1970, 1971, 1972a, 1973a). Written reports of pigeons
were not solicited from field personnel although communications were encouraged for ease in locating groups of pigeons in trappab1e locations. Trapping
methodology in 1973 was that used in previous years, with most birds being
captured through use of cannon nets. Evaluation of the hunting season and
disposition of collected birds were the same as in previous years.
Description of Trapping Areas
In 1973 pigeons were trapped at 21 locations utilized in previous years.
These locations were near Del Norte, LaVeta, Salida, Wetmore, Durango, Monte
Vista, Avon-Minturn, Stoner, Evergreen, Estes Park, Montrose, Arboles, Longmont,
Uniweep Canyon, Aspen, Sanborn Park, Ft. Garland, Woodland Park, Buford,

�-156-

Carbondale and Woodmen, and have been previously described (Braun 1970, 1971,
1972a, 1973a). In addition to these sites, pigeons were trapped near Molina,
Gypsum, Hesperus, and Chipita Park. These sites were grain fields (Molina),
livestock feeding areas (Gypsum and Hesperus), and residential areas (Chipeta
Park) where residents fed a variety of birds. As in previous year, all trap
sites were immediately adjacent to readily accessible trees or power lines
used for perches. Trees used were primarily ponderosa pine, Gambel's oak
(Quercus gambelii), Pinyon pine (Pinus edulis), cottonwoods (Populus spp.),
and.Engelmann spruce (Picea engelmannii). It was difficult to ascertain
feeding preferences, as about equal numbers of pigeons were trapped in fields
where they were feeding on waste barley (Hordeum vulgare), and wheat (Triticum
aestivum). Whole or cracked corn (Zea mays) was frequently used as an attractant and bandtails clearly indicated a preference for this grain.
RESULTS AND DISCUSSION
Banding
Trapping and banding activities decreased from 1972 levels, but numbers of
birds banded (3,559) in 1973 were still third highest of the five years of
study. The reduced number of birds banded in 1973 was the result of decreased
emphasis on banding large numbers of birds at all sites. Emphasis instead,
was placed on banding samples of pigeons (usually less than 200 birds per
site) in as many locations as feasible. As a result, more than 200 birds
were banded at only four of the 25 trapping locations. Efforts were continued
to maintain banding at as many of the sites used in previous years as
feasible. Thus, trapping was successful at 4 of the 5 sites used in 1969,
10 of the 13 sites used in 1970, 12 of the 15 sites trapped in 1971, and 18
of the 23 locations used in 1972. In all, 3,559 pigeons were newly banded
(Table 1). Of this number, 570 (16.0 percent) were immatures. Percent
immatures trapped in 1973 was similar to that reported in 1970 and 1972
(16.6 and 13.6 percent, respectively), but lower than the 23.6 percent found
in 1969, and over twice as high as that found in 1971 (7.2 percent). Upon
examination of data in Table 1 it is appraent that percent immatures in trap
samples increased throughout the summer. The initial immature trapped was
on 4 July (Montrose) about 15-20 days later than in previous years. This
suggets that nesting was later in 1973 than in some years, a hypothesis
supported by field observational data of breeding activities.
Sex of adult and subadult pigeons was determined through examination of
external plumage characters. Of the 2,812 adults, 1,343 were males (47.8
percent), and 1,469 were females (52.2 percent), while only 68 of the 177
subadults were males (38.4 percent), with the remainder (109) classified as
females (61.6 percent). These data are nearly identical to those collected
since 1970. The disparity in sex ratios of adults is probably related to
time of day trapping was conducted. The variance in percent of subadult
males and females trapped compared to percent adult males and females
captured (difference of 9.4 percent) is not understood. It may be related
to differences in molting rates between males and females, differential
mortality or observer error.

�Table 1.

Number of band-tailed pigeons banded by area in Colorado, 1973.

Area

Date(s)

Adults];..!
Percent
No. Banded

Innnatures
No. Banded
Percent

Totals

Estimated No. of
Birds Using
Trapping Sites

Molina

May 19

69

100.0

0

0.0

69

lS0

Gypsum

May 21

86

100.0

0

0.0

86

lS0

Del Norte

May 23-26
October 18

238
7

100.0
S3.8

0
6

0.0
46.2

238
13

400+
SO

LaVeta

May 24-27
August 21

219
3S

100.0
76.1

0
11

0.0
23.9

219
46

SOO+
100+

Salida

May 2S-26
June 19-30
July 20-21

10S
17
11

100.0
100.0
91.7

0
0
1

0.0
0.0
8.3

10S
17
12

lS0
SO
2S

Wetmore

June 6-7

122

100.0

0

0.0

122

2S0

Hesperus

June 6

19

100.0

0

0.0

19

SO

New MexicoV

June 7-13

22

100.0

0

0.0

22

Durango

June 8-20

446

100.0

0

0.0

446

SOO+

Monte Vista

June 11
July 10-14
August 10-29

6
32
177

100.0
88.9
42.3

0
4
241

0.0
11.1
S7.7

6
36
418

2S
SO
SOO+

Chipita Park

June 11
July 21

70
SO

100.0
6S.8

0
26

0.0
34.2

70
76

lS0
lS0

Avon

June 13

123

100.0

0

0.0

123

200+

,

•....
VI

---------------------------------------------------------------------------------------------------------------

~,

�Table 1.

Number of band-tailed pigeons banded by area in Colorado, 1973 (continued).

Area

Date(s)

Adu1ts1/
No. Banded
Percent

Immatures
No. Banded
Percent

Totals

Estimated No. of
Birds Using
Trapping Sites

Stoner

June 19

88

100.0

0

0.0

88

100+

Evergreen

June 20
August 10-22
September 3-28

57
18
33

100.0
52.9
75.0

0
16
11

0.0
47.1
25.0

57
343/
44-

100
75
75

Estes Park

June 21
August 10

51
16

100.0
88.9

0
2

0.0
11.1

51
18

100
50

Montrose

I

~

VI

June 22-28
July 4

24
64

100.0
92.8

0
5

0.0
7.2

24
69

100
100

Arboles

June 23

64

100.0

0

0.0

64

100+

Longmont

June 24
July 10
August 6

14
56
31

100.0
100.0
27.7

0
0
81

0;0
0.0
72.3

14
56
112

100
100
200+

Uniweep Canyon July 5

24

100.0

0

0.0

24

50

Aspen

July 6-7

112

98.2

2

1.8

114

150

Sanborn Park

July 12-13

127

92.0

11

8.0

138

200

Ft. Garland

July 15-31
August 1

131
0

78.0
0.0

37
1

22.0
100.0

168
1

200
200

Woodland Park

July 22
August 18

49
10

56.3
32.3

38
21

43.7
67.7

87
31

150
150

---~------------------.---------------------------------------------------:--------------------------------------

00
I

�Table 1.

Number of band-tailed

pigeons banded by area in Colorado, 1973 (continued).

Totals

Estimated No. of
Birds Us:ing
Trapping Sites

23.5

183

200+

13

36.1

36

100

100.0

0

0.0

3

25

84.0

570

16.0

3,559

1/
AdultsPercent
No. Banded

ImmatureS
Percent
No. Banded

Area

Date(s)

Buford

July 23-25

140

76.5

43

Carbondale

August 3-4

23

63.9-

Woodmen

August 26

3
2,989

Total

I

~

VI
\0

1/ Includes pigeons classified as subadults.
~/ All trapped at LaVeta in 1972, held in captivity over winter and released in New Mexico.

11 Includes 27 birds from aviary (24 adults, 3 immatures).

I

�-160-

Time of day was recorded for all pigeons captured in 1973. A summary of
these data is presented in Table 2. As in other years, more males than
females (64.8:35.2 percent) were trapped prior to 1000 hours MDT. Between
1000 and 1600 hours, more females than males were captured (68.0:32.0 percent)
while after 1600 hours, differences were not as great, as 59.0 percent of
the birds trapped were males, while 41.0 were females. Most adults were
trapped between 1000 and 1600 hours (48.9 percent of all captures), with
lesser percentages being trapped prior to 1000 hours (33.5 percent), and
after 1600 hours (17.7 percent). This probably explains the slight imbalance
in sex ratios of adult pigeons newly banded. Immatures were caught more
frequently before 1000 hours (53.8 percent of all immature captures) than
in any other time period. Reasons for this anomaly are not understood, but
are probably related to behavior.
During the 1973 trapping period, 1,017 previously banded birds were recaptured with all but 7 being initially banded in Colorado. Of the 7 "foreign"
bandings recaptured, 6 had been initially banded in New Mexico, while 1 had
been banded in Utah. Considering the 1,010 birds initially banded in Colorado, 24 were banded in 1969, 118 were banded in 1970, 237 were banded in
1971, 505 had been banded in 1972, while 126 were newly banded in 1973. In
all, 280 pigeons recaptured had moved at least one degree block from where
initially banded. Less than 100 had changed flock areas, a situation
previously reported (Braun 1972b). Thirty-nine birds banded in Colorado
were reported recaptured in New Mexico (35), and Utah (4). All but 3 had
been banded prior to 1973. It is probable that some of these pigeons were
migrating to or from Colorado when trapped, but it is also likely that some
flocks of pigeons occupy areas partly in two states. Some of these recaptures may represent birds that have changed flock areas. From examination
of the reported recaptures, it is obvious that pigeons living in the LaVetaFt. Garland area routinely move into New Mexico. This is also true for
some pigeons of the Monte Vista south flock and the Del Norte-MOnte Vista
north flock. The 4 recoveries reported in Utah (all at Monticello)
represent 4 different bariding locations in southern and southwestern Colorado. Of 39 birds that were recaptured out of Colorado for which sex was
ascertained, 21 were females, while 18 were males. This agrees with earlier
reports (Braun 1972b), that indicated no differences between sex classes in
those birds moving away from flock areas where originally banded.
Hunting Season
Band-tailed pigeon hunting, while no longer experimental, was closely
monitored in Colorado in 1973. Hunting was allowed from September 8 through
October 7, with a daily bag limit of 5 birds and a possession of 10. During
the season, all of the state west of Interstate Highway No. 25 was open to
hunting. As in previous years all hunters had to obtain a free permit
obtainable at 9 Division offices and the headquarters of the Monte Vista
National Wildlife Refuge. No limit was placed on total number of permits
to be issued, and 455 permits (an additional 7 permits may have been issued)
were obtained by hunters. This was a decrease of 107 (19.0 percent) from
1972.

�Table 2;

Time MDT

Time of day of band-tailed pigeon trapping, 1973.

Percen t 0 f All
Successful
Trap Attempts

Percent of All Adu1t1/
Captures
Males
Females

Percent of:l:./ Percent ofl/
All Females
All Males
Captured
Captured

Percent of
All Immatures
Captured

Percent ofl/
All Adults
Captured

0600-0959

26.6

64.8

35.2

45.4

22.6

53.8

33.5

1000-1559

45.5

32.0

68.0

32.7

63.6

28.5

48.9

1600-

27.9

59.0

41.0

21.8

13.9

17.8

17.7

1/ Including those birds classified as subadu1ts.

I

t-'
0\
t-'
I

~/ Adults and subadults only.

�-162-

Hunter Questionnaire Survey
The hunting season for bandtails in 1973 closed at sunset on October 7. On
October 8 and 9, letters containing a self-addressed, postage paid return
card were sent to all permittees. No questionnaires were returned for
lack of sufficient address. On November 6 a follow-up letter was sent to
the 109 permittees who had failed to respond to the initial letter. Results
of the questionnaire survey are presented in Table 3. Data for the 36 nonrespondents were calculated using mean values of the respondents to the
followup survey.

Table 3.

Harvest statistics, 1973 band-tailed pigeon season.-1/

Category

Response
First
Letter

Number of permittees
responding

346(76.0)

73(16.0)

419(92.1)

Number of permittees
hunting

173(50.0)

26(35.6)

199(47.5)

212(46.6)

Number of permittees
not hunting

173(50.0)

47(64.4)

220(52.5)

243(53.4 )

Number of successful
hunters

46(26.6)

9(34.6)

55(27.6)

60(28.3)

Number of hunter days

544

65

609

641

Days hunted per
hunter

3.1

2.5

3.1

3.0

Number of pigeons
bagged

273

36

309

329

Pigeons per successful hunter

5.9

4.0

5.6

5.5

Pigeons per hunter

1.6

1.4

1.6

1.6

Number of pigeons
crippled and lost

27

6

33

37

Pigeons crippled and
lost per hunter

.2

.2

.2

.2

Total harvest (bagged +
crippled and lost)
300

42

342

366

Percent crippling loss 9.0

14.3

9.6

10.1

Response
Second
Letter

1/ Values in parentheses are percentages.

.Sum
First and
Second Letters

Projected
For all (455)
Permittees

�-163-

Interest in pigeon hunting decreased in 1973 primarily because pigeons were
widely scattered due to excellent food conditions, and thus were not concentrated during the hunting season. Hunter success declined significantly
from 1972 (28.3 percent versus 41.6 percent) and total harvest fell from
822 in 1972 to 366 in 1973. As in previous years, a significant number of
people who obtained permits did not hunt (53.4 percent). These individuals
probably obtained a permit in order to hunt if they observed pigeons while
engaged in some other activity. Hunters did hunt longer in 1973 (3.0 versus
2.6 days) than in 1972 and number of pigeons bagged per successful hunter
did not materially change from 1972 to 1973 (5.9 to 5.5).
Percent crippling loss (total number of pigeons reported lost divided by
total harvest, i. e., bagged and crippled and lost) was 10.1 in 1973;
similar to the 11.3 percent reported in 1972. Reported crippling loss is
minimal and observations of actual crippling loss suggest that it may represent as much as 20 percent of the birds actually retrieved.
The questionnaire was devised in order to ascertain the proportion of the
harvest that occurred in given time periods. It was found that 45.6 percent
of the total harvest occurred on the opening weekend, 10.4 percent on the
second weekend, 11.1 percent occurred on the third weekend, while 6.7 percent
occurred on the last weekend of the season. Not surprisingly, only 21.2
percent of the total harvest occurred during the non-weekend portion of the
season (20 days). Harvest was fairly well spaced during the 1973 hunting
season, as only 65.1 percent of the total harvest occurred during the first
9 days. This suggests that either hunting was poor throughout the season
or that hunter interest remained somewhat constant as the season progressed.
Hunting pressure in 1973 was diffuse as 193 hunters reported hunting in
49 different locations. The Durango (12.5 percent) and Monte Vista (9.1
percent) areas continue to be preferred for hunting, and these 2 areas had
21.6 percent of all hunting trips. No other area had more than 5 percent of
the hunter activity. On a regional basis, 43.3 percent of the hunters
hunted in southwestern Colorado, 33.7 percent hunted in southeastern Colorado,
12.5 percent hunted in northeastern Colorado, and 10.6 percent hunted in
northwestern Colorado. These data are similar to that collected in all 3
of the previous seasons.
Most comments received from hunters concerning the hunting season referred
to their failure to locate large flocks of birds, and to the scattered distribution of those that they did see. Hunting was difficult.
Wing Survey
Packets containing 5 wing envelopes with instructions to place one day's
bag in each envelope were issued to most pigeon permit holders in 1973.
Envelopes containing 240 wings were received from 44 hunters. Additional
data were received concerning hours hunted (230.8), birds crippled and lost
(50), location of kill and banded birds harvested (43). Major harvest areas
were near Durango (39.1 percent of the harvest), LaVeta (21.7 percent),
and Cotopaxi (18.7 percent). All other areas contributed less than 6 percent
each of the total harvest. These data are somewhat different than the
distribution of hunters obtained from the questionnaire survey. Thus,

�-164-

Durango, with 12.5 percent of the hunters had 39 percent of the harvest,
while Cotopaxi and LaVeta each with less than 5 percent of the total hunters
each had about 20 percent of the total harvest. On a.regional basis, 53.5
percent of the harvest came from the southwestern part of the state, 41.8
percent came from the southeast, 4.3 percent was from the northwest, and only
0.4 percent occurred in the northeast. Crippling loss calculated from the
wing survey was 17.8, substantially higher than the 10.1 percent reported on
the questionnaire survey.
Forty-three of the 236 bandtails (excluding wings from 5 feral pigeons,
Columba livia) reported harvested on the wing survey were banded (18.2 percent).
This is the highest percentage recorded in any of the 4 years that hunting
has been allowed (6.3 percent in 1970, 9.3 percent in 1971, and 13.6 percent
in 1972). Percent of the birds harvested that were banded varied from 20.7
(N = 92) at Durango, 19.6 (N = 51) in the LaVeta area to 13.6 (N = 44) at
Cotopaxi. Bands were reported from 4 other locations but sample sizes of
wings received were too small (less than 10) for meaningful comparisions.
None of the banded birds were harvested within one mile of any trap site.
While it is not believed that 18 percent of the fall population was banded
percent banded birds in the population is undoubtedly greater than 10 percent.
Comparable data were available from both hunter questionnaire and wing surveys
concerning numbers of birds bagged and those crippled and lost by 38 individual hunters. A reporting difference of +5.8 percent (206 in wing survey
versus 218 in questionnaire survey) was found in number of birds killed, and
-46.3 percent (41 versus 22) in number of birds crippled and lost for those
hunters who sent in wings and also responded to the questionnaire. The
difference in birds reported killed could not be attributed to a few hunters
killing more pigeons than for which they had wing envelopes, since no
hunters reported killing more than 25 birds. It is interesting to note that
this group of hunters reported crippling only half as few (22 versus 41)
pigeons on the questionnaire as they reported on their wing envelopes.
All wings received were measured (carpal and all 10 primaries) and classified as immature, subadult and adult in order to derive age structure of
the harvest. Wings were received from 182 adults (77.1 percent), 4 subadults
(1.7 percent), and 50 immatures (21.2 percent). Percent of adults was
higher than in any year of the study (67.8, 68.0, 64.8 percent, respectively,
for 1970 through 1972), while percent immatures and subadults decreased
markedly in 1973. Age ratios in the 1973 harvest were different to that
expected based on trapping results and observations in August and early
September. From all appearances, 1973 was a good production year, but
obviously many immatures were not available to hunters. It is possible that
many immatures had migrated leaving only late nesting adults and their
progeny available to hunters. Upon examination of molt data for immatures,
the hypothesis that many early produced young had migrated is not supported.
Of the 50 immatures, only 12 (24.0 percent) had not molted primary 1, while
30 (60.0 percent) had molted primary 2 or beyond. All birds molting primary
2 or beyond were at least 59 days past hatching, and 29-34 days past fledging
(White 1973). Twenty of the 50 hunter harvested immatures (40 percent) had
hatched prior to July 1.

�-165-

Hunter Field Checks
During the 1973 season, emphasis was again placed on contacting pigeon
hunters in the field in order to examine harvested birds for crop gland
activity.
Crops of all adult pigeons checked were classified as being
active (crop gland with curds), stimualted (gland apparent, but no curds),
or inactive.
In addition, sex by gonadal inspection was obtained for as
many birds as possible.
In all, 122 pigeons were checked, with 27 being
immatures (22.1 percent), with the remainder (95) being classified as
adults.
Of the adults, 32.6 percent were classified as having active crop
glands, while 30.5 percent were classified as being stimualted.
Percent
of crops showing glandular activity (active + stimulated) was higher in 1973
than in 1972 (63.2 versus 50.9), and was the highest recorded in the 4 years
of the study. This supports field observations that indicated that nesting
and/or feeding of young continued late into August and September.
Data on sex was available for 88 adults.
Of this number, 61.4 percent were
males, and 38.6 percent were females.
Of the males, 63.0 percent exhibited
some glandular development, while 61.8 percent of the females showed development of the crop gland. These data are dissimilar to that collected in
previous years, which showed that more males than females were associated
with nesting at time of death.
When samples of adults examined by area are
compared, it is apparent that nesting was still occurring in most areas of
the state during the hunting season.
These data indicated that 61.5 percent
of the 52 adults examined at Durango were still associated with nest
activities, this percentage was 66.7 in the LaVeta area (N = 18), 85.7 at
South Fork (N = 7), and 60.0 (N = 15) at Cotopaxi.
With the exception of
Durango, samples were too small to draw meaningful conclusions concerning
differences between areas.
Mortality
Recoveries of 92 band-tailed pigeons initially trapped and banded in Colorado
have been reported to date in the 1973 recovery year.
Five of these were
initially banded in 1969, 9 were banded in 1970, 14 were banded in 1971, 45
were banded in 1972, while 19 were banded in 1973. All but 20 were shot
recoveries.
Of the 72 shot recoveries, 47 were recovered in Colorado, 17
in New Mexico, 4 in Arizona, 3 in Mexico, and 1 in California.
Hunting
mortality in New Mexico continues to be higher than initially expected (Braun
1972b).
Migration from Colorado by some pigeons appears to be underway by
September 1, as 7 banded birds were recovered in New Mexico during the week
of September 1 through 7. Only 1 of these 7 pigeons had been banded in 1973.
Of the 17 recoveries in New Mexico, most (16) were taken along a line southwest from LaVeta to Silver City.
Interestingly, 14 of the 17 recoveries were
banded in eastern slope areas of Colorado.
Of the 4 pigeons recovered in
Arizona, 3 were banded west of the Continental Divide, while the 1 California
recovery was banded east of the Continental Divide.
It would appear that
birds from easternslope areas migrate south along the Rocky Mountains in ColQrado, and then southwest through New Mexico.
Pigeons from western slope area~
move southwest through Colorado and then south along the eastern edge of
Arizona.
These patterns are similar to that reported for recoveries in 1972
(Braun 1973a).
During the 1973 hunting season, one pigeon banded in Ari~ona

�-166-

(1971) was recovered near Durango. This is the first shot recovery from
Arizona and only the second movement from Arizona to Colorado that has been
documented.
Breeding Phenology
Few data on breeding pheonology were available as this portion of the study
was completed in early 1973 (Gutierrez 1973). Data were available from examination of 41 birds killed in trapping operations (24), highway casualties
(11), poaching (2), presumed electrocution (2), and found crippled or dead
(2). Data from these pigeons indicated that nesting started in mid- to late
May and continued into September. Observational data indicated that a few
pigeons started arriving in Colorado by early April (April 3 at Evergreen),
with peak spring migration in mid- to late May, with some numbers of pigeons
remaining as late as October 18. Thus, observational data support those
obtained from gonadal and crop examinations. Observational data for previous
years of the study have been reported by Braun (1973b). During 1973, 1
active nest containing 1 egg was found by W. Stanley of Collbran on Baxter
Point on 22 August. This nest and egg were examined, described and photographed. It was unfortunately abandoned, possibly due to activities of the
observers.
Wing Measurement and Molt Data
Data concerning length of primaries, rectrices, total body length, carpal
length and bill length were obtained from most dead pigeons examined in
1973. Once adequate samples are available, these data will be analyzed to
examine differences between age and sex classes of pigeons. Prior to
dissection of collected birds, plumage characters of each adult specimen
were examined and recorded. Each bird was then classified as either male or
female. Upon dissection, gonads were examined and actual sex of each bird
was compared with the classification based on plumage appearance. Comparative data were available for 39 adults and subadults (18 males and 19
females), and 2 immatures. Only 1 bird was inaccurately sexed. This bird
appeared to be a male by plumage, but was actually a female. This percentage of error (2.6) is similar to that found in prior years.
Information concerning primary molt was available for 623 immatures (569
banded and released, 50 from the hunter wing survey and 4 from birds necropsied). Hatching dates were estimated using data from White (1973). Estimated haoching dates for all immatures for which data were available are presented in Table 4. As in earlier years, hatching was spread over a long
time period (April 12 to September 11). Two major peaks (mid-June and early
July) in hatching can be inferred from data in Table 4, but the interval is
too short to represent two nests by the same group of pigeons. It would
appear that the hatching data in Table 4 only represent the range of hatching
dates and give some idea when most pigeons are nesting. These data are
probably not representative of the actual nesting peaks in Colorado in 1973.
Of importance in Table 4 are the data that suggest a steady decrease in
numbers of pigeons hatching after mid-July. All observational and crop data
collected in August and September refute the hypothesis that most (over 99
percent) nesting terminated prior to September 1.

�-167-

Table 4.

Estimated hatching dates for wild band-tailed pigeons 1973.
Number
Hatching

Percent

Prior to April 11

0

0.0

April 12-21

2

April 22-May 1

Number
Hatching

Percent

July 1-10

147

23.6

0.3

July 11-20

28

4.5

2

0.3

July 21-30

15

2.4

May 2-11

4

0.6

July 31-August 9

8

1.3

May 12-2 i

17

2.7

August 10~19

6

1.0

May 22-31

33

5.3

August 20-29

4

0.6

June 1-10

91

14.6

August 30-September 8

2

0.3

June 11-20

179

28.7

September 8-17

3

0.5

June 21-30

82

13.2

Time Period

Time Period

Parasitic Load
Information on helminths was available for 36 pigeons necropsied. Five (all
adults) of the 36 birds examined had helminth infections. The infection rate
of 13.9 percent is similar to that found in 1969 (13.2 percent), higher than
that found in 1970 and 1972 (10.4 and 9.5, respectively), but lower than the
18.3 percent observed in 1971. Three of the 5 birds infected were females,
while 2 were males. Four of the pigeons had nematodes, and 2 had tapeworms,
(1 female had both). No adverse effects due to the observed parasites were
noted. All worms were preserved for later identification. Materials from
all 5 years of the study are presently being identified by Dr. O. W. Olsen,
Colorado State University.

LITERATURE CITED
Braun, C. E. 1970. Band-tailed pigeon investigations. Colo. Div. Game,
Fish and Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. p.
151-171.
1971. Band-tailed pigeon investigations. Colo. Div. Game, Fish
and Parks. Game Res. Rept., Fed. Aid Proj. W-88~R. October.
p. 209-236.

�-168-

Braun, C. E. 1972a. Band-tailed pigeon investigations. Colo. Div. Wildl.
Game Res. Rept., Fed. Aid Proj. W-88-R. October. p. 123-142.
1972b. Movements and hunting mortality of Colorado band-tailed
pigeons. Trans. No. Am. Wildl. and NaL Resour. Conf. 37:326-334 .
1973a. Band-tailed pigeon investigations. Colo. Div. Wildl. Game
Res. Rept., Fed. Aid Proj. W-88-R. October. In Press.
1973b. Distribution and habitats of band-tailed pigeons in Colorado.
Proc , Western Assoc. State Game and Fish Comms. 53:In Press.
I

Gutierrez, R. J. 1973. Band-tailed pigeon investigations:breeding and
nesting chronology studies. Colo. Div. Wildl. Game Res. Rept., Fed.
Aid Proj. W-88-R. October. p. 153-177.
White, J. A. 1973. Band-tailed pigeon investigations:plumage studies ..
Colo. Div. Wildl. Game Res. Rept., Fed. Aid Proj. W-88-R. October.
p. 125-152.

Prepared by

~jld~~~

~_~,rv~t?t~~
__ ~~~~.~~~~~~
Clait E. Braun
Wildlife Researcher

_

�October 1974

-169-

JOB l:'ROGi\.E~SLETORT

State of'
Project

CO1010\]) 0
------~~~-~--------------

work Plan

No.

Migratory

W-88-R-19

No.

5

Investigations

Period Covered:

September

1

-----of Snipe in Colorado
._---

-----.'------------

Job Title

Job No.

Bird Investigations

1, 1973 to March 31, 1974

Per sonne.l.: C. 'E. Braun, H. D. Funk, and B. R. Johnson, Colorado Division
Wildlife; and R. A. Ryder, Department of Fisheries and Wildlife Biology,
Colorado State University.

of

Investigations of connnon snipe (Capella gallinago de1icata) were initiated in
September 1973. Field work conducted thus far concerned selection of study
areas in the San Luis Valley, North Park, Yampa Valley, and the vicinity of
Fort Collins. Areas with apparently good snipe population densities during
September and October generally provided unfavorable habitats during spring
because of poor moisture conditions.
Selected study areas will be censused
during the breeding season through territorial responses of snipe elicited
by playbacks of recorded territorial calls.
In addition, each area will be
described using such parameters as vegetative dominance, height and density,
habitat modifications
such as lOOwing, burning, or grazing, water chemistry,
water depth and permanence.
Evaluations of habitat preferences of breeding
snipe will be made based on these data. Censuses on snipe wintering in the
Fort Co11irisarea from January through March indicated high (20 snipe flushed
per hour) and stable population densisites on preferred habitats.
Efforts
to document hunting pressure and estimate harvest and hunter success were
largely unsuccessful because of the dearth of respondents to Colorado small
game hunter questionnaires.

�-170RECOMMENDATIONS

1.

It is recommended that a description of the obj ect Lves of this study
by sent to all field personnel of the Colorado Division of Wildlife.
Help should be requested from field personnel in describing where snipe
occur in their districts, when they are present arid major habitat types
occupied.

, ",

�-171-

INVESTIGATIONS

OF SNIPE IN COLORADO

Bruce R. Johnson

Members of the genus Capella have an almost worldwide distribtuion, however,
the common snipe (Capella gallinago delicata) is the only snipe subspecies
indigenous to North America.
Although snipe are members of the family
Scolopacidae, they have few characteristics usually attributable to shorebirds.· As an example, snipe do not frequent shores of lakes and coasts.
Rather, they prefer bogs,low
moist meadows, or wet pasturelands.
They are
seldom seen in large concentrations as are other shorebirds and are primarily
crepuscular, remaining in cover during bright daylight.
Furthermore, snipe
are one of only two shorebirds hunted as a game bird, the other being the
woodcock (Philohela minor).
Prior to the 1940's, snipe hunting was quite popular as witnessed by Bent's
observation in 1927 that "probably more snipe have been killed by. sportsmen
than any other game bird."
However, with closure of the snipe season from
1941 through 1952, interest in snipe hunting greatly diminished.
After two
decades· of open seasons and an almost tripling
of U. S. hunters, interest
in snipe hunting is increasing.
Estimates of the annual snipe harvest have
increased twofold from 263,900 in 1965, to 524,600 in 1970 (Clark 1971).
In. the mid-1960's, interest in migratory webless game bird management increased,
and consequently, the Bureau of Sport Fisheries and Wildlife initiated an
accelerated research program designed to supplement research programs on game
species such as snipe. The results of such research should increase knowledge of webless game birds, thus allowing management to be refined and
intens ified.
P. S. OBJECTIVE
To study the distl;'ibution and status of common snipe in Colorado in order
that data might be made available to improve management of the species.
SEGMENT OBJECTIVES
1.

To identify

and delimit range of common snipe in Colorado.

2.

To document seasons of occurrence
major areas.

3.

To document hunting

4.

To compile data, analyze results,
findings in appropriate technical

pressure

and estimate

and estimate

relative numbers

harvest

prepare progress
journals.

and hunter

in 3

success.

reports and publish

In addition, evaluations will be made concerning: (1) habitat preferences of
common snipe breeding in various areas throughout Colorado, and (2) density
and stability of wintering snipe populations on selected study areas near
Fort Collins.

�-172-

REVIEW OF LITERATURE
Description
Blairet
al. (1957) give the following description of the connnon snipe:
"Crown sooty, with a median buff stripe; hind neck streaked black and buff;
back and tertials black barred with brown, with four long buff stripes;
wing coverts dark brown, spotted with whitish; tertials black, edged with
white and barred with brown; remiges dusky; edge of outer primary and tips
of primary coverts and secondaries white; upper rump black, narrowly barred
with white; lower rump and upper tail coverts rich buff, irregularly barred
with black; tail black, rufous terminally, where interrupted by a black bar;
lateral pair of rectrices black-and-white-barred;
next two pairs white
beyond subterminal black bar; superciliary buff; lores brown; chin buffy
white; throat and upper breast buff streaked or barred with brown; belly
white; sides black and white barred; crissum buff with dusky squamations;
axillars and wing lining barred black and white; iris brown; tip of bill
dusky; base of bill and feet greenish.
Wing 118-135 nnn, tail 50-63 nnn,
culmen 58-73 nnn, tarsus 28-33 nnn."
Origin
Tuck (1972) suggests that the shorebird ancestors from which the· genus
Cappella ev lved were a very primitive group.
It is quite likely that these
primitive shorebirds inhabited the shores of vast cretaceous salt basins as
many present-day snipe species posSess vestigial characteristics such as
nasal salt glands and rudimentary membranes between the toes •. However,
Capella gallinago does not have nasal salt glands or rudimentary membranes
which suggests it evolved as a species rather recently (Tuck 1972).
Unlike most shorebirds, there is no seasonal and little sexual dimorphism
in Capella, suggesting that this genus evolved in boggy areas associated
with forests or in regions of low cover.
In any case, taken as a group, the
present snipes are more forest birds than shorebirds (Tuck 1972).
Taxonomy
The extensive radiation and divergence of snipe and snipe-like birds has
perplexed those wishing to classify snipe into conventional taxa. Tuck
(1972) has separated the snipes into three descriptive groups; semi-snipes,
a seemingly primitive stock; aberrant snipes, which appear to have diverged
from the main stock; and true snipe, apparently of recent radiation.
Tuck (1972) considers the common snipe to be a true snipe which, according
to the American Ornithologists'
Union (1957), belongs to the order Charadriiformes and Family Scolopacidae.
The true snipes are included in one genus,
Capella, and divided into nine species which are remarkably alike.
Species
s~paration is ..based principally on the number of tail feathers each species
possesses (Tuck 1972).
Capella gallinago is further divided into eight subspecies whose members
presently indigenous to every continent except Australia and Antarctica
(Tuck 1972).
The common snipe, Capella gallinago delicata, is the only
member indigenous to North America.

are

�-173-

Dist ribut ion
Common snipe occur at one season or another, in nearly every part of North
America (Bent 1927). The American Ornithologists'
Union (1957) reports
the breeding range extends from northwestern Alaska, northern Yukon,
northern and eastern MacKenzie, northeastern Manitoba, northern Ontario, both
coasts of James Bay, northern Quebec, and central Labrador south to northern
and eastern California, central eastern Arizona, northern Colorado (casually
south to San Luis Lakes), western Nebraska, central Iowa, northeastern
Illinois, northern Indiana (formerly), southern Michigan,. northern Ohio,
northern West Virginia, northwestern Pennsylvania, northern New Jersey
(rarely), southern New York, Connecticut, Massachusetts, Maine, New Brunswick,
Nova Scotia and Newfoundland.
It has also been reported to breed in central
Mexico.
Snipe normally begin their fall migration from the northern portions of their
breeding range during early September and the bulk of the North American population winter in the rice fields, wet cattle pastures, and coastal marshes
of Louisiana (Tuck 1972). Whitehead (1965) noted that males arrive in Louisiana during October and leave in March, while females arrive in November and
leave in April.
The primary wintering range of common snipe extends from southern British
Columbia,western
Washington, Idaho, Oregon, northern Utah, northern Colorado, central Nebraska, Missouri, Illinois, Indiana, western Kentucky,
western and central Tennessee, northern Alabama, northern Georgia, South
Carolina, central North Carolina, and Virginia south to the West Indies,
Colombia, Venezuela, and southern Brazil.
It has been casually or irregularly reported to winter north to southwestern Alaska, Montana, Wyoming,
North Dakota, Minnesota, Wisconsin, southern Ontario, Ohio, West Virginia,
New York, Connecticut, Massachusetts, New Hampshire, Maine, and Nova Scotia.
Common snipe have also been recorded casually in Greenland, and less frequently in Hawaii, Bermuda, England, and Scotland (Stout 1967).
The distribution of common snipe in Colorado has not been well documented.
Sclater (1912) notes that snipe are perhaps most connnon during migration
when they can be found throughout the state.
They can also frequently be
found wintering near unfrozen springs, both on the plains and in the mountains
up to 9,500 feet. Bailey and Niedrach (1965) observed that snipe are sunnner
residents in small numbers from the plains to the Lowe r Canadian zone;
numerous during migration, and often seen wintering at low elevations.
There are no published estimates as to the number of snipe in Colorado
during a particular season.
Boeker (1954) censused the Yampa Valley in
northwestern Colorado during the spring and sunnner of 1953 and estimated
that a breeding population of 200 snipe utilized the Valley at that time.
Published reports of snipe wintering in Colorado date to, at least,. 1886 as
two common snipe were observed.near Silverton in midwinter at an elevation
.of 9,500 feet (Anon. 1886). The editor of this publication (Forest and
Stream) reported that snipe were not uncommon in the Rocky Mountain regions
in the coldest winter weather in the neighborhood of warm springs having
open water and soft mud.

�-174-

Aiken and Warren (1914), in cataloging the birds of El Paso County, Colorado, noted that along the banks of Fountain Creek, which rarely froze
over, there were many places where water seeping through sand came to the
surface and formed marshy areas.
These areas became winter feeding grounds
for normally one or two snipe. However, during severe cold, many smaller
areas froze forcing snipe to concentrate at places where a larger flow of
water kept the area open.
More recently, Bailey and Niedrach (1965) reported that, although many
sn Ipe leave Colorado by November, there is a scattered winter population
wherever marshes and borders of sloughs remain unfrozen.
Ecology
The common snipe is an early spring migrant, leaving its wintering grounds
in the southern United States, Central America, and northern South America
as early as February (Stout 1967). The principal migration period occurs
from March through mid-April, with males arriving on breeding grounds approximately ten days before females (Tuck 1972).
Shortly after arrival, males begin winnowing over relatively large areas.
Upon arrival of females, males reduce the size of their territories and
intensify their winnowing.
Territories are generally located in bogs,
marshes, peatlands, or other wetland types having low vegetation.
Territories become well defined after females begin incubation.
Theprimary
requisites for territories appear to be: (1) a suitable nest site, (2) a
suitable area for the males' station during incubation, normally in dense
cover, (3) a suitable retreat for the female, and (4) a feeding area (Tuck

1955).
Nests are usually built of fine grasses, but occaSionally of fine; dry
sedges (Tuck 1955) and are so cohesive that they may be lifted intact (Stout
1967). Average clutch size is four eggs, although three are not uncommon
(Tuck 1972). Eggs are ovate pyriform in shape, slightly glossy (Bent 1927),
dark ochre or light ochre in color and boldly spotted or blotched in dark
shades of brown, brownish black, or black (Tuck 1972).
Incubation usually requires from 18 to 20 days and is done only by the
female (Tuck 1972). Upon hatching, the brood is shared, although not necessarilyevenly,
between the male and female (Stout 1967). To date, there is
no evidence that snipe produce more than one brood annually.
Development of chicks is quite rapid, and within one week after hatching
chicks double their weight.
By three weeks, they are as heavy (90 grams) as
their moulting parents (Tuck 1972). The chicks' first flights are taken
between 15 and 18 days of age (Stout 1967). When the young are about six
weeks old, adults begin mOUlting and become solitary.
Immatures begin aggregating by mid-August, and may occur in flocks of one hundred or more. There
are indications that young migrate prior to adults (Tuck 1972).
Pre-migratory restlessness first appears in common snipe in mid-August (Tuck
1972), and it is likely that migration begins with the first early frosts
(Bent 1927). The main flights in southern Canada and the upper half of the

�-175-

conterminous United States occur from September to early October (Stout
1967).
During this time, the Colorado snipe population is probably at its
height (Bailey and Niedrach 1965).
Although most common snipe migrate to wintering grounds in southern United
States, Central America, and northern South America, some overwinter
regularly on wet cattle pastures of Vancouver Island, British Columbia, and
less frequently, on tule marshes, open creeks, and open springs of Oregon
and Washington (Tuck 1972).
Snipe are commonly found in the Humboldt region
of northwest California (White and Harris 1966).
However, they are reportedly rare in winter east of California, because of continental climatic
conditions (Tuck 1972).
In Colorado, Sc1ater (1912), and Bailey and Niedrach
(1965) report that common snipe frequently winter in areas where open springs
occur.
Early naturalists assumed that food habits of common snipe were similar to
those of woodcock with earthworms constituting their principle food (Bent
1927).
Later findings based on stomach analyses (Whitehead 1965, White 1963,
Van Owens 1967, Booth 1968) revealed that between 42.7 percent (Booth 1968)
and 63.1 percent (White 1963) of the stomach contents were plant material.
Investigators believe that a large proportion of this plant material is
accidentally ingested and the fact that the plant fibers remain relatively
unchanged by any digestive process supports this hypothesis (Tuck 1972).
Composition of animal and plant material ingested by common snipe varies
with season and location.
Insect food, particularly larvae, often comprised
more than 80 percent of the total animal material ingested.
However, this
may be less than 15 percent in salt marshes and rice fields (Tuck 1972).
Other foods include some crustaceans, earthworms, mollusks, and even an
occasional fish (Stout 1967).
Common snipe obtain food primarily by probing upper levels of soft organic
soils. The snipe's bill, which is one-fifth of the total body length, has
a highly developed sense of touch.
Furthermore, the distal portion of the
upper mandible is pliable and can be elevated upward (Tuck 1972).
This
action enables the snipe to grasp food items and swallow without removing
its bill from the. ground (Erickson 1953).
Snipe are also surface feeders
(Stout 1967), and have been reported during migration to run along stream
borders picking up food particles (Bent 1927).
Erickson (1941) examined 76 snipe and found 80 percent infected with parasites. More recent accounts of ectoparasites and endoparasites are reported
by Whitehead (1965), Schmidt (1962), Booth (1968), and Tuck (1972).
Census Techniques
With the exception of spring winnowing, snipe are normally secretive and
seldom seen unless flushed.
Consequently, censusing snipe populations is
difficult,..especially._dliring,
..late summer •.fall and winter.

�-176-

Snipe Winnowing

Counts

Perhaps the first technique employed on a large scale to census breeding
snipe was the "winnowing" count.
One adaptation of this method was employed
by Burleigh (1952). After determining a starting point, he established
successive stations at half-mile intervals.
The half-mile intervals were
decided upon after a ntimberof tests were made to determine how far away a
winnowing snipe could be distinctively heard.
Closer intervals would have
involved possible duplication, while longer distances would have unquestionably meant an incomplete count.
Solman (1953) concurred with the one-half
mile distance.
Burleigh also investigated
factors are:

factors necessary

for an accurate

count.

These

1. The actual hour at which each was made. Morning hours proved entirely
unsatisfactory and evenings appeared to be the best times for accurate
censusing. .Solman (1953) also stated that part of the post-sunset winnowing
period was likely to be the most useful as a popUlation index.
2. Weather conditions.
whereas adverse weather

Maximum counts were made on still, clear nights,
reduced the count materially.

3. Activities of the individual bird. Although male snipe remain in the
air for long intervals, they were noted from time to time dropping to the
ground where they would remain for an indefinite period.
Consequent1y,the
count was reduced.
Finding a bird in the air when a scheduled stop was made
was apparently a matter of arriving at an established station at an opportune
moment.
4. Breeding activities of individual pairs of snipe. A possible factor in
an accurate count is the possibility that males cease "winnowing" flights
when the young have hatched and demand the attention of both parents.
The winnowing count was further evaluated by Robbins (1953). He described
his censusing procedure as follows:
Each transect was covered by car, with
two minute stops at each half-mile interval.
The number of birds winnowing
and the number of winnows heard from each bird were recorded for each minute.
It also proved helpful to indicate the direction and approximate distance of
each winnowing bird.
In this way, any significant change in position during
the two-minute period could be detected and nearly all duplications with
birds at the preceding and following stop could be recorded as such. Temperature, wind, and sky conditions were recorded at the beginning and end of
each transect along with notations of significant weather changes during
the transect census.
Robbins noted that in Ontario and New Brunswick (460 N latitude) winnowing
was consistent enough to serve as a population index only during a half-hour
period from 75 minutes to 40 minutes before sunrise, and from 30 minutes to
60 minutes after· sunrise.

�-177-

Regarding weather conditions, Robbins considered wind velocity to be, by
far, the most important factor.
Winds of 5 mph or more invariably reduced
·winnowing activity and increases to 10mph generally stopped all winnowing.
Morning temperatures ranging from·300 to 500 F. produced near or below
average resu Lt s; and above 500 F. the counts were especially low. No correlation was found between evening temperatures and snipe winnowing activity.
Clear skies in the early morning .were associat.ed with average occurrences
of winno~ng
(within 20 percent), with a tendency to be above average
rather than below.
Clear skies in the evening were associated with much
more variable results (from 40 percent below average to 40 percent above
average).
On calm, overcast evenings and mornings, winnowing was very
variable (from 55 percent below to 45 percent above average).
Rain reduced
the occurrence of winnowing, whereas morning fog was associated with near
to above average results.
Tuck (1972) observed that although snipe winnow any time of day or night
during the breeding season and sporadically during migration, the maximum
winnowing period is during the twilight hours of evening and morning.
Winnowing reaches a peak soon after the females arrive and counts for the
following.IO days are quite reliable.
Timing of the peak period varies, and
in Newfoundland, is from May 5 to May 15. In more southern latitudes, the
peak period is considerably earlier •.
Tuck also found that winnowing may be suppressed by wind, rain,or
dense
fog. An.overcast day will precipitate evening winnowing by half an hour
or so, a bright; hot day will retard it by a similar length of time. Both
sexes winnow, although the female apparently does not winnow. for very long.
Females sometimes winnow only a half dozen times, but may remain in the air
for considerable periods diving, but not winnowing.
Tuck concluded that the post-sunset winnowing period was more reliable
than that of the pre-daWn period.
He also reported that all birds heard
winnowing at one time are not necessarily paired, and that in marginal
habitat, birds frequently winnow, but move elsewhere to breed.
However,
the above problems are taken into consideration, a comparative index of
snipe found in a relatively large area can be calculated each year.
Territorial

if

Response

A second method of censusing breeding populations is the territorial response
technique.
This technique
is relatively new and is considered by Tuck (1972)
to be the only precise method of censusing snipe. Although the details are
not available, Tuck describes the technique as an estimation of the breeding
population through territorial responses of snipe elicited by play-backs of
recorded territorial calls. ·These calls are described as "yakking" and are
heard most frequently during the twilight periods before winnowing begins.
These ground calls suddenly
reach a peak of urgency and cease abruptly. as
the snipe springs into the air. Both sexes "yak" and the techniques can be
effectively used the entire day during the early part of the breeding season,
but is more time consuming than other techniques.

�-178-

The above two techniques are restricted to censusing breeding populations.
The remaining techniques can, for the most part, be used to census either
a breeding or wintering population, but are better suited to the latter.
Snipe Abundance

Index

The snipe abundance index has been used extensively to estimate wintering
snipe populations in southern states.
The census is accomplished by driving
or walking a specified road or transect and recording the number of snipe
seen per hour. Robbins (1957) criticized this method because 30 to 40 percent
of the winter snipe counts cannot be used for year-to-year statistical comparison because they either are not taken in the same areas for two or more
successive years, or they are taken entirely by car or boat instead of on
foot and therefore cannot be used in statistical comparison which are based
on number of party-hours of foot coverage.
Random

Ground Transects

Robbins (1955) pointed out that one of the major fallacies in the snipe
abundance index is that the majority of areas covered include optimum
wintering habitat and there is insufficient information from large areas of
low abundance.
Observers asked to census a certain portion of the country
usually select an area where they can record the largest number of birds
with a minimum amount of effort.
If all areas were selected on a purely
random basis a large increase in personnel and expense would be required.
The practicality of random transects was tested in wintering areas where
the snipe popUlation was relatively widespread.
Fifty 5-minute intersections
of latitude and longitude in southern Louisiana were selected by a random
drawing.
Each location was then marked on a topographic map and a 20-mile
census route was laid out starting at the nearest road and running as
closely as possible in a northerly direction.
Robbins found that average number of snipe seen per hour on the random
ground transect was almost identical to the number determined by the snipe
abundance index. He concluded that with more intensive coverage it should
be possible to obtain rough estimates of the total population, although
important biases, such as all transects·parallel roads, may always be present.
More recently, Martin and Goudy (1965) investigating the feasibility of
randomized breeding survey routes found that population indices obtained
from counts on randomly selected routes were more representative of actual
breeding popUlation sizes than indices from non-random routes.
Air Transects
The use of air transects is another technique for estimating wintering snipe
popUlations.
Robbins (1955) described the following method for conducting
an aerial census of snipe. _ All flights were made at .an air speed of 85 mph
and a height of 70 to 120 feet. Transects were counted during all daylight
hours expect late afternoon and under a variety of temperatures, winds, and
sky conditions.

�-179-

Transects were run primarily through rice fields in Louisiana.
Nearly all
the birds flushed came from a strip 300 feet wide directly under the plane
and had to be flushed to be seen. Robbins listed the following conditions
to be avoided as they seriously reduced the number of snipe recorded by
observers:
(1) overcast skies; (2) winds in excess of 15 mph; (3) freezing
temperatures or frozen ground; and (4) late afternoon or pre-sunrise coverage. Robbins also notes that air transects in Louisiana should preferably
be made during January and the first week in February, as there is a decided
influx of other shorebirds into rice fields after this date.
Perhaps the chief advantage in aerial censusing is the ability to cover
large areas in relatively little time.
Smail (1968) comments that to census
on the ground areas normally covered in 90 minutes by air, would take about
4 days, even if access could be gained.
Strip Census
The strip census or King method is advantageous in open bogs or other areas
of low vegetation.
It is conducted by walking a predetermined route through
snipe habitat and counting the number of birds and the distances at which
they flushed.
The average flushing distance is determined and used to calculate the effective width of the census strip. The population is the number
of birds flushed divided by the area of the strip and multiplied by the total
ar~a (Tuck 1972).
AZ

p

2YX

where:

P
A
Z
Y
X

population
total area of study
number flushed
average flushing distance
length of strip

K. A. Arnold (personal communication 1973) found that on his study areas
in Texas the only reliable method of censusing wintering snipe was by
flushing the birds and observing where the flushed birds alighted.
Detonating

Devices

Tuck (1972) states that on fall migration and wintering grounds, delayed
detonators fired from a shotgun into a field of known size will cause snipe
to flush so they can be counted.
Annual Wing Survey
An annual wing survey, similar to the waterfowl wing survey conducted by the
U. S. Fish and Wildlife Service, would provide a ratio of adults to juveniles
and an index to annual production.
This method is not presently used and
is not possible until a system of contacting migratory game bird hunters is
devised (K. A. Arnold, personal communication).

�· -180-

Wetland

Classification

Methods for classifying wetlands are numerous and oftentimes dictated by
the intensity, scope, and resources of the study. Each method incorporates
characteristics common to other wetland classification sy~temsand
conversely, each method may contain unique characteristics or unique procedures
in summarizing characteristics into a concise classification scheme.
Sanderson and Bellrose (1969) define "wetlands" as lowlands covered with
shallow and sometimes temporary or intermittent water - the waterlogged or
shallow water areas known as wet meadows, marshes, sloughs, potholes, swamps,
bogs, and river-overflow lands.
Shallow lakes and ponds, often with emergent
vegetation, are included in the definition, but· open-water areas of streams,
reservoirs. deep lakes, and bays are not included, although such areas may
contain valuable wetland areas around their margins.
These open water areas,
however, have been included in several broad classification schemes •.
An early attempt to classify wetlands was devised by Bach in 1951 (Leitch
1966). Bach's classification divides prairie wetlands into descending
classes of permanency.
These classes areas
follows:
a.

Areas with permanent water

b.

Areas which normally

c.

Areas which are normally

d.

Areas having only transitory

hold water all year except in severe droughts
dry by July or August
water

Another system, developed by Martin et ale in 1953 is presently used by the
Bureau of Sport Fisheries and Wildlife.
This system categorizes wetlands
primarily upon evaluation of water depth and permanence, water alkalinity,
geographic location and dominant vegetation.
Wetlands were initially divided
into twenty categories ranging from seasonally flooded basins or flats to
mangrove swamps.
Five additional categories have been added to include
streams, rivers, and artificial wetlands (Fredrickson 1967).
The initial 20 wetland types were listed. under four regional categories:
Inland Fresh, Inland Saline, Coastal Fresh, and Coastal Saline.
The following
categories include wetland types and descriptions of water levels during
various times of the year. Martin .et a1.have also given further descriptive
information including location of each type in the United·State,s, prominent
vegetation, and each type's value to various wildlife species which have not
been included here:
Inland Fresh Areas
1. Seasonally Flooded Basins or Flats: Soil covered with water or waterlogged during variable seasonal periods; usually well drained during much
of the growing season.
Along river courses, flooding ordinarily occurs in
late fall, winter, or spring; in upland areas, basins or flats may be filled
with water during periods of heavy rain or melting snow.

�-181-

2. Fresh Meadows:
Soil without standing water, but water-logged
at least a few inches of its surface during the growing season.

within

3. Shallow Fresh Marshes:
Soil normally water-logged during the growing
season; often covered with as much as 6 inches of water.
4. Deep Fresh Marshes:
growing season.
5.

Open FreshWater:

Soil covered with ~ to 3 feet of water during the

Water of variable

depth.

6. Shrub Swamps:
Soil normally water-logged during the growing season;
often covered with as much as 6 inches of water.
7. Wooded Swamps:
Soil water-logged at least to within a few inches of
its surface during the growing season, often covered with as much as 1 foot
of water.
8. Bogs: Soil usually water-logged, generally blanketed
covering of mosses or other plant material.

with a spongy

Inland Saline Areas
9. Saline Flats:
Soil without standing water, but water-logged
at least a few inches of its surface during the growing season.
10. Saline Marshes:
Soil normally water-logged
often with as much as 3 feet of water.
11.

Open Saline Water:

Water of variable

to within

during the growing season,

depth.

Coastal Fresh Areas
12. Shallow Fresh Marshes:
Soil always water-logged during the growing
season, may be covered at high tide with as much as 6 inches of water.
13. Deep Fresh Marshes:
Soil covered at average
feet of water during the growing season.
14.

Open Fresh Water:

Water of variable

high tide with ~ to 3

depth.

Coastal Saline Areas
15. Salt Flats:
Soil almost always water-logged during the growing season,
sites varying from those submerged only by occasional wind tides to others
that are covered fairly regularly with a few inches of water at high tide.
16. Salt Meadows:
Soil always water-logged
rarely covered with tide water.

during the growing season,

17. Irregularly Flooded Salt Marshes:
Soil covered by wind tides at
irregular intervals during the growing season.

�-182-

18. Regularly Flooded Salt Marshes:
Soil covered at average high tide
with ~ foot or more of water during the growing season.
19.

Sounds and Bays:

Water of variable

depth.

20. Mangrove Swamps:
Soil covered at average high tide with ~ to 2 feet
of water during the year round growing season.
The U. S. Fish and Wildlife Service (1955) in an attempt to determine wetland
types found in Colorado, surveyed North Park, the San Luis Valley, the
Arkansas Valley, and the South Platte Basin. These 4 areas encompass 77
percent of the Colorado's wetlands.
Each wetland was classified using the
above classification.
The nine types of wetlands found in Colorado with their percent occurrence
as determined by the survey are listed below:
Description

Percentage

1

Seasonally

2

Fresh meadows: while none of this type
occurred on areas covered by the inventory,
it is found in other sections of the state.

3

Shallow fresh marshes

18

4

Deep fresh marshes

3

5

Open fresh water

14

6

Shrub swamps

6

9

Saline flats

14

10

Saline marshes

1

11

Open saline water

(+)

flooded basins or flats

44

In addition to the above types, lands regularly flooded in connection with
agricultural practices, such as pastures and haylands, were included
separately and deSignated as Type I-A. Hopper (1968) describes Type I-A
wetlands as pastures and hayland periodically flooded with irrigation water
during spring, summer, and early fall for producing livestock forage. Vegetation consists primarily of a mixture of grasses, sedges, and rushes.
Martin et al.'s system is advantageous for rapidly classifying large numbers
of wetlands, however, the categories are too broad for detailed habitat
investigations.
Quite often, the Martin et al. system serves as a basis upon
which additional measurements or estimates are added.
Such additions may
include soil and geological maps, land capability classifications, densities
and distributions of wild and domestic animals, water chemistry, and temperature and precipitation fluctuations (Mann 1955).

�-183-

Stewart and Kantrud (1971) developed a more dynamic and complex classification system that more precisely
reflected seasonal, regional, and local
variations.
They stated that the classification system of Martin et al.
(1953) was too general for detailed investigations of wetlands and has
often been misinterpreted by placing too much emphasis on water depth and
cover interspersion.
Furthermore, they argue that water depth of a given
wetland type varies considerably.
By itself, it is generally a poor indicator of prairie wetland types and should not be equated with water permanence.
The classification system devised by Stewart and Kantrud is suited to natural
ponds and lakes in the glaciated prairie region.
Briefly, the system
involves seven major classes of wetlands, each distinguished by the vegetational zone· occurring in the central or deeper part of the wetland and
occupying 5 percent or more of the total wetland area being classified.
Plant species characteristic of each classes were included in the system.
The classes are:
Class I

Ephemeral

ponds

Class II

Temporary

ponds

Class III

Seasonal ponds and lakes

Class IV

Semipermanent

Class V

Permanent

Class VI

Alkali ponds and lakes

Class VII

Fen (alkaline bog) .ponds

ponds and Lakes

ponds and lakes

In addition, distinCt subclasses may be recognized within several of these
major classes.
Stewart and Kantrud based these subclasses on differences
in species composition of plant connnunities within wet meadow, shallow marsh,
or deep marsh zones that· are correlated with variations in average salinity
or surface water.
These subclasses are designated as:
Subclass A

Fresh

Subclass B

Sl.Lght.Ly brackish

Subclass

Moderately

C

Subclass D

Brackish

Subclass

Subsaline

E

brackish

The classification system also considered four cover types. These cover
types represent differences in the spatial relationships of emergent cover to
open water or exposed bottom soil and ranged from closed stands of emergents
with less than 5 percent open water or bare soil. to 95 percent open water or
bare soil.

�-184-

To use this system in the field, each pond or lake was classified by designating the class, subclass (if differentiated), and the cover type, in
.
that order.
The system was found to be both expedient and realistic in
classifying the ponds and lakes in Worth Dakota.
Cowardin and Johnson (1973) found Stewart and Kantrud's system had disadvantages. in its restriction to the glaciated prairie and omission of many
wetland types important to their area (north-central Minnesota).
The
classification system devised by Cowardinand
Johnson used the stand as
the basic unit which they defined as an area of wetland vegetation with
sufficient uniformity and size so as to be recognizable both on the ground
and on aerial photographs.
During field surveys, each plant species within a stand was described in
terms of the percent of area coverage and placed into one of the following
.categories: 0 = absent; 1 = rare, a few scattered individuals; 2 = occasional, less than 1 percent; 3 = fairly common, 1-10 percent; 4 = common,
11-50 percent; 5= abundant, 51-100 percent. . Stands were assigned. to communities according to their physical and botanical characteristics.
Vegetative
species which were used as indicators of water permanence by Stewart and
.
Kantrud (1971) were .used similarly whenever possible.
Communities were .
divided into those haVing permanent water and those without permanent water.
The communities were further separated on the basis of the amount of emergent cover present.
This separation was similar to the cover types described by Stewart and Kantrud (1971). .On the basis of these collective
characteristics,
each wetland was classified as one of 21 community types.
The communities without permanent water included: (1) ephemeral; .(2) temporary; (3) seasonal-closed;
(4) seasonal-patchy;
(5) seasonal-open; (6) semipermanent-closed;
(7) semipermanent..,.patchy; (8) semipermanent-open;
(9) shrub
swamp; (10) hardwood swamp; (11) circumneutral bog-sedge bog; (12) circum.neut ra.L bog-shrub phase; (13) circumneutral bog-ericaceous phase; (14) acid
bog; and (15) softwood swamp. Communities with permanent water included:
(16) open water; (17) bulrush; (18) ph ragmf.t.ea
;" (19) wild rice; (20) emersed
aquatics, and (21) other emergents.
Wetland surveys conducted by the National Wildlife Federation (Morgan and
Rose 1967) included 7 categories upon which wetland evaluations were made.
These categories were as follows:

. . .~.

Category I was a soil texture classification containing 10 gradations
ranging from sand and gravel to heavy clay. Category II was a land utilization classification of 8 classes varying from excellent agriculture
soils to soils impossible to farm and des Lgnat.ed for wildlife and recreational
use. Category III consisted of7 classes of natural topography and drainage
ranging from undulating and poorly drained to level and well drained •
Category. IV consisted of 10 divisions of the annual average precipitation
from below 5 inches to 25 inches or more. Category V was an estimate of
evaporation as indicated by the mean July temperature.
This category is
divided into 10 classes varying from 63.50 F. or lower, to 72.60 F or greater.
Category VI was a measurement of the number of water areas per square mile
determined from aerial photograph interpretation.
The category consisted
of 10 classes ranging from 4 water areas or less, to 45 or more. Category
VII consisted of 5 wetland types described as follows:

�-185-

We tland Type I:

Shallow depressions, usually holding water in the
spring or after heavy rains.

.Wetland

Type II:

Shallow fresh marshes holding surface water during
spring and early summer.
Often dry by midsummer.

.Wetland

Type III:

Deep fresh marshes normally holding surface water yearround, except during extended periods of drought.
Usually have cattails or bulrushes around edge and
often over the entire marsh.

Wetland

Type IV:

Open fresh water of variable depth.
Often surrounded
by marsh vegetation.
Seldom go dry except during
periods of extreme drought.

Pothole

Community:

A group of water areas of various
mile diameter circle.

types within a 4

Areas were then divided into 6 categories varying from one to 20 points based
upon the wetland type or combination of wetland types located within a specified wetland area •.
These 7 categories represent a possible 100 points on rating system.
Categories I through III considered soils, land use, and topography for a maximum of 50 points.
Categories IV through VII considered climatic influences
and the numbers and types of water areas for the remaining 50 points.
Each
wetland area was then assigned a priority rating based on its total point
value. The priority system ranged from Priority I (80-100 points) to
unimportant (below 40 points).
In an attempt to categorize each wetland area concisely, a classification
system utilizing a.chain formula has been devised (U.S.D.I. ND). This chain
formula was devised to provide a mental image as well as factual data for a
wetland area.
In devising the chain formula, 6 basic factors were selected
consideration.
These included:
1.

Acreage

2.

Permanency (as determined
a1. "s 0 r Bach's)

3.

Vegetation

4.

Species of vegetation

5.

Occurrence

6.

Percent of open water

density

by a classification

(sparse,moderate,
(abbreviated

of vegetation

such as Martin et

or. dense)
if possible)

(border or throughout)

for initial

�-186-

An example of the chain formula could be written as 4-4-II-Typha-Bo-75.
This would describe a four acre deep fresh marsh (using Martin et al. 's
classification)
with a moderately dense cattail border and 75 percent open
Wa ter. Additional factors could be added to the formula corresponding to
the necessary parameters required by the investigation.
A classification
system not strictly limited to wetlands but which has
application to a variety of vegetative types throughout the world is a
physiognomic classification devised by Kuchler (1949). This classification
is necessarily broad since it is applicable to all types of vegetation in
all regions throughout the world.
In this classification, plants were
divided intowoody
and herbaceous plants.
Woody plants were subdivided
.into evergreen broadleaf, deciduous broadleaf, evergreen needleleaf,
deciduous needleleaf, and without leaves; herbaceous plants were subdivided
into graminoids, forbs, and lichens and mosses.
The vegetation was further subdivided according to plant height, density
and special features such as epiphytes, succulents, or aquatic vegetation.
Each subdivision was indicated by a letter, thus vegetation over an area
is described by a formula.
The system is advantageous in that it is a well known classification system
and can be readily applied to any location.
Furthermore, the system is
adaptable to a.ny scale of investigation and can be used to classify small
as well as large areas.
However, because of its broadness, the system may
require additional parameters to increase number of classifications
for
more intensive studies.
The use of soil types for classifying areas is an important parameter in
several classification systems and served as a basis for a rating system
devised by Allan et al. (1963).
In this study, eight habitat elements were
selected for rating via soil conditions.
These elements were (1) grain and
seed crops, (2) grasses and legumes,(3)
wild herbaceous upland plants,
(4) hardwood woodland plants, (5) coniferous woodland plants, (6) wetland
food and cover plants, (7) shallow water developments, and (8) excavated
ponds.
First, soil types were ra t ed in terms of suitability for these
habitat elements.
For example, the category wetland food and cover plants
which included annual and perennial wild herbaceous plants of moist to
wet sites, the rating criteria were as follows:
1.

Soil cqnditions suitable for the growth cif a wide variety of climatically adapted species particularly food producing plants.

2.

Soil conditions
.perennials.

3.

Soil conditions· that tend to produce dominant stands of a few vigorous
perennial species, generally of low food production value.

4.

Soil conditions under which wetland plants do not grow or are so sparse
as to be of no significance to wildlife.

suitable

for a wide variety

of species particularly

�-187-

Next, various combinations of habitat elements were considered for their
significance to and rating for general classes of wildlife.
These classes
were: (1) openland wildlife, (2) woodland wildlife, and (3) wetland wildlife. The ratings of the suitability of soils for these wildlife classes
were made on the basis of weighted values assigned to a selection of habitat
elements appropriate to a particular class of wildlife.
For example, grain
and seed crops were given a greater weight than hardwood woodland plants as
habitat elements for open1and wildlife.
Each soil type is then classified
as either well suited, suited, poorly suited, or unsuited as to these wildlife habitat elements and kinds of wildlife.
Other wetland inventories (Farmes 1956, Leitch 1966, Smith 1971) have incorporated various aspects of the above classifications systems and also
included other descriptive characteristics such as available nesting cover,
adjacent land. use, historical data, and water basin shape depending upon the
intensity of the. study and the wildlife species of interest.
Since vegetation often plays an important role in wetland classification,
factors influencing the vegetative composition of a wetland are also of
interest.
Each plant species has its own particular level of tolerance to
inundation.
Those with similar moisture requirements and limitations are
characteristically
found in locations with the same moisture regime. As a
result, pothole vegetation typically occurs in zones that can be correlated
with depth and duration of submergence.
This relationship has frequently
been used as a factor in classifying wetlands.
.
Millar (1969) discussed several of the important factors which affect distribution and density of plant species. Briefly, these factors are:
1.

Availability of Seed: The absence of plant species in certain areas may
be cause by absence of viable seed rather than to deficiencies in the
environment.

2.

Moisture Regime:
Each plant species will tolerate different depths and
durations of flooding, and these differences are reflected in the rather
definite zonation of wetland vegetation.

3.

Chemistry of the Basin Soil: This is examplified by the effect of
extreme salinity in reducing the complexity and luxuriance of marsh
vegetation.

4.

Climatic Factors:
Such factors include length of growing season and
severity of temperature extremes but little .is known about requirements
of individual species.

5.

Mode of Reproduction:
This is important in determining the survival
and proliferation of a species.
Perennials spreading by rhizones and
seeds may perpetuate themselves more easily than annuals or biennials.

6.

Inter-and
Intra-specific Competition:
This factor plays an important
role in limiting the density and distribution of species but as yet
knowledge of the factor is negligible.

�-188-

7.

Disturbance:
Disturbances such as cultivation, mowing, grazing, burning,
drainage, and fluctuation of water levels tend to alter the environment
of a basin in such a way as to adversely affect existing vegetation.

These classification systems are only a portion of many which have been
devised.
Each system has been described only briefly and for a complete
description of each, the reader is referred to the orignial reference.
METHODS

AND MATERIALS

Selection

of Study Areas

Initial investigations on Connnon snipe in Colorado began in September, with
assessment of possible study areas for use in evaluating the importance of
representative habitat types to breeding snipe. Major geographic areas in
Colorado in which sites have been investigated include the San Luis Val~ey,
North Park, Yampa Valley and neat Fort Collins.
Each potential study area was grossly examined and factors such as drainage,
water levels, vegetation type and densd ty, domestic animal grazing, and
ownership were noted.
Areas appearing favorable as snipe habitat were
censused by walking circumferences of ponds or banks of streams.
Numbers of
.snipe flushed, :if any, were recorded.
If no snipe were flushed, the location
of the area was recorded so that it could be recensused at a later date,
provided the area appeared to contain excellent habitat, or if the habitat
appeared to be suitable for snipe following grazing or flooding.
Since these .
censuses were made only to determine the potential of an area to serve as a
spring study area, most areas were censused only once and no attempt was made
to quantify the data respective of time or area.
Each area censusedwas
then
recorded on a map for future reference.
Selected areas will be censused
regularly to estimate breeding densities on each site. These and/or additional study areas will also be censused to estimate fall and winter
densities.
Censusing

Techniques

Spring censusing of all breeding snipe will be based on a technique desribed
by Tuck (1972) in which population densities are estimated by territorial
responses elicited by play back of recorded territorial calls. Mechanics of
this technique are not known in detail, however, a letter has been sent to
Dr. Tuck requesting the information needed.
Fall and winter censusing will
utilize either the strip census or a birds observed per hour index.
Hunting

Pressure

Hunting pressure and harvest was estimated through questionnaires mailed to
hunters Who responded on the 1972 Colorado small game questionnaire that
they had harvestad snipe. A copy of the snipe questionnaire and cover letter
are included in the Appendix.
Wing samples from each bird harvested by the
hunters were also requested.
Wings were examined to provide an estimate of
the juvenile:adult
ratio in the hunted population.

�-189-

Winter

Census

Investigations of common snipe wintering in the Fort Collins vicinity were
initiated in mid-December and continued through March.
Four areas having
open water and low vegetatiOn were censused to estimate numbers on each site.
Two contrasting study areas were selected for biweekly censusing to evaluate
density and stability of populations in each habitat.
Each census was conducted as follows:
The starting time was recorded and
the census route paralleled the stream varying in distance from 3 feet to 1.5
feet from the shoreline, depending on topography of the stream bank.
The
observer walked at a moderate pace periodically clapping his hands to flush
snipe. When a snipe was flushed, the time and the estimated distance from
the observer to the flushed· snipe were recorded.
The location in which the
flushed snipe alighted was noted to avoid duplicate counts. At the route's
end the time was again recorded.
Data collected during each census weTe summarized as total snipe flushed
during each census (Fig. 1) and snipe flushed per hour. When the length
of each route has been ascertained, .the data will be quantified into a
snipe flushed per kilometer index.
Description

of Winter Study Areas

The first study area, designated as Area 1, is a southeast flowing drainage
near the Fort Collins sewage plant. The area is characterized by a. narrow
(4 to 6 feet), shallow, flowing stream with steep to perpendicular banks .
from 4 to 8 feet in height.
Vegetation is coarse and partially grazed, few
exposed mud flats are present
in the stream channel.
The second site, Area
2 is a. portion of Spring Creek within the Fort Collins city limits which
begins at Taft Hill Road and ends immediately west of South College Avenue.
This route contains a variety of habitats, the majority of which are good :
to excellent for wintering snipe. Portions of the route are grazed and many
exposed sparsely vegetated mudflats
occur.
It is in these areas that snipe
are most commonly found.
Wetland

Classification

Methods to be used in this study for evaluating and classifying wetlands
have not been field tested.
From the wetland classification systems reviewed,
parameters were chosen for consideration evaluating habitat preferences of
snipe. These parameters are as follows: (1) acreage of study areas; (2)
County and location of study areas; (3) ownership of study areas; (4) elevation; (5) water permanence, based on Martin et al.,"s (1953) classification;
(6) water depth and fluctuations; (7) conductivity, alkalinity, and pH of
water; (8) water turbidity; (9) natural drainage of study areas; (10) stream
flow velocity and volume; (11) hazard of flooding; (12) stream bank or wetland shoreline topography and erosion; (13) percent open water and percent
emergent vegetation; (14) coverage and distribution of dominant species of
vegetation; (15) height of dominant vegetation; (16) phenology record of
dominant plants; (17) tree heights within study areas; (18) frequency or

�-190-

40

35

·Area 2

30

Cf)

0
Cf)

z 25

f.&gt;Q
U

r::t::
r:t:l
P;
q
;.:!

:I:: 20
Cf)

0

.-:l

~

r:t:l
P;
H

z

Cf)

15

'-

~

0
r::t::
r:t:l

~

0
z
10

5

Area 1

20

27

January

3

10

14

24

9

February

17

25

29

Harch
CENSUSING

Fig. 1.

3

DATES

Number of snipe flushed per census from January

through March,

1974.

�-191-

density of aquatic invertebrates; (19) soil types within study areas; (20)
classification of land capability; (21) available moisture within 12 inches
of soil surface; (22) mean annual precipitation;
(23) evaporation as estimated by mean monthly temperatures; (24) composition and juxtaposition of
habitat types; (25) domestic animals present; '(26) degree of freedom from
human disturbance; (27) habitat modifications such as mowing, burning,
grazing, cultivation, or pollution; and (28) photographs of ea.ch study site.
Each of these parameters is currently being evaluated in respect to importance of the parameter to breeding snipe and simplicity with which a
reliable estimation or measurement of the parameter can be made. Because
several widely distributed study areas will be used, intensive evaluation
of each area will not be possible.
RESULTS AND DISCUSSION
Selection

of Study Area

In September and October of 1973, ten potential study areas having sizable
snipe populations were located: four areas in the San Luis Valley, three
near Fort Collins, and three in North Park. Each area was classified into
one of the following categories: (1) flooded hayland; (2) wet grazed pastureland; (3) wet ung raz ed pastureland, and (4) grazed river oxbows.
Elevations
of these areas ranged from 5, 000 ft in the Fort Collins vicinity to 8,200 ft
,in North Park. The Yampa Valley was also investigated.
However, all areas
examined were devoid of standing water.
These poor moisture conditions
precluded selection of potential st udy areas.
Field investigations were resumed in mid-March in an effort to locate
additional study areas to serve in evaluating habitat preferences of breeding
snipe. Thus far, all areas examined were near Fort Collins.
Eleven
potential study sites were censused, however, no snipe were flushed on any
area.
Several of these areas appeared to contain excellent snipe habitat
and will be recensused in April.
During several censuses, a recording of a snipe territorial call was played,
but no snipe responded, perhaps because the areas may not have contained
snipe. Also, no snipe have been heard winnowing during the censuses, but
it should be noted that winnowing most often occurs during twilight periods
which did not coincide with the times of the censuses.
Hunter Questionnaires
Thirty-eight questionnaires were distributed in September to obtain an
approximation of hunter interest in snipe hunting, areas most frequently
hunted, days afield, hunter success, and number of snipe seen and bagged.
Results of this survey were disappointing as only 3 questionnaires were
returned by hunters.
Of these 3, 2 did not hunt snipe in 1973 and the
third individual hunted a total of 4 days and bagged an estimated 25 snipe.
Obviously, with a sample size of 3 no inferences can be made.

�-194-

Hopper, R. M. 1968. Wetlands of Colorado.
Parks, Tech. Publ. 22. 88 p.
Kuchler, A. W. 1949. A physiognomic
Assoc. Amer. Geogr. 39:201-210.
Leitch, W. G.
inventory.

Colo. Dept. of Game, Fish and

classification

of vegetation.

Anns.

1966. Historical and ecological factors in wetland
Trans. N. Am. Wildl. and Nat. Resour. Conf. 31:88-96.

Mann, G. E. 1955. Wetlands inventory of Wisconsin.
Servo Office of River Basin Studies.
63 p.

U. S. Fish and Wildl.

Martin, A. C., N. Hotchkiss, F. M. Uhler, and W. S. Bourn. 1953. Classification of wetlands of the United States. U. S. Fish and Wildl. Servo
.Spec. Sci. Rept. Wildl. No. 20. 14 p.
Martin, F. W., and M. H. Goudy. 1965. Feasibility of randomizing singingground survey routes in important woodcock breeding areas. p. 62. In
Ann. Prog. Rept , Fiscal Year 1965. U. S. Bureau Sport Fish. and Wildl.
Migr. Bird Pop. Sta., Laurel, Maryland.
Millar, J. B. 1969. Observations on the ecology of wetland vegetation.
Saskatoon wetlands seminar.
Can. Wildl. Servo Sere No. 6:49-56.
Morgan, H. R., and B. J. Rose. 1967. Wetland preservation rating system
for Manitoba and Saskatchewan.
National Wildl. Federation, Washington,
D.C.2Ip.
Robbins, C. S. 1953. Further investigations on winnowing methods of
measuring Wilson's snipe populations.
p.61-65.
In Investigations of
woodcock, snipe, and rails in 1953. U. S. Fish and Wildl. Servo Spec.
Sci. Rept. Wildl. No. 24. 68 p.
1955. Wilson snipe wintering ground studies, 1954-1955.
p. 47-54.
In Investigations of woodcock, snipe and rails in. 1955. U. S. Fish and
Wildl. Servo Spec. Sci. Rept. Wi1d1. No. 31. 54 p.
1957. Wilson snipe wintering ground studies, 1955-56.
In Investigations of woodcock, snipe and rails in 1956.U.
Wildl. Servo Spec. Sci. Rept. Wi1dl. No 34. 85 p.

p. 57-65.
S. Fish and

Sanderson, G. C., and F. C. Be1lrose.
1969. Wildlife habitat management
of wetlands.
Ill. Nat. Hist. Surv. No. R272. Urbana.
p. 154-204.
Schmidt, G. D. 1962. Endoparasites of Wilson's snipe Capella gal1inago
delicata Ord, in northeastern Colorado.
M. S. Thesis.
Colo. State
Univ., Ft. Collins.
65 p.
Sclater, W. L. 1912.
London.
576 p.

A history of the birds of Colorado.

Witherby and Co.,

�-195-

Smail, J. 1968. Aerial census of shorebirds, an evaluation.
Reyes Bird Observatory Newsletter No. 11:45.

Point

Smith, A. G. 1971. Ecological factors affecting waterfowl production in
the Alberta parklands.
U. S. Fish. and WildL
Serv. Resour • Pub!. 98.
49 p.
Solman, V.E.F.
1953. Wilson's snipe "winnowing" counts in eastern Canada,
1953. p. 57-59.
In Investigations of woodcock, snipe and rails in 1953.
U. S. Fish and Wildl. Servo Spec. Sci. Rept. Wildl. No. 24. 68 p.
Stewart, R. E., and H. A. Kantrud.
1971. Classification of natural ponds
and lakes in the glaciated prairie region.
U. S. Fish and Wildl. Servo
Resource Publ. 92. 57 p.
Stout, G. D. 1967.
York. 270 p.

The shorebirds

of North America.

Viking Press, New

Tuck, L. M. 1955. Some observations on nocturnal activities of Wilson's
snipe - Newfoundland (1953). p. 41-50. In Investigations of woodcock,
snipe and rails in 1954. U. S. Fish and Wildl. Servo Spec. Sci. Rept.
Wildl. No. 28. Washington, D. C. 62 p.
1972. The snipes: a study of the genus Capella.
Servo Monograph Series No.5, Ottawa.
429 p.
U. S. D. 1. [po Dat~
.
Servo 5 p. Mimeo.

Wetland classification.

Canadian Wildl.

U. S. Fish and Wildl.

U. S. Fish and Wildlife Service.
1955. Wetlands inventory: Colorado.
U. S. Fish and Wildl. Serv., Office of River Basin Studies. Albuquerque.
19 p. + 16 p. of Figs.
Van Owens, J. 1967. Food habits of the common snipe (Capella gallinago
delicata) in the pastures of south-central Louisiana.
M. S. Thesis.
Louisiana State Univ., Baton Rouge. 108 p.
White, M. 1963.
Occurrence, habitat, ecology, food habits and sex and age
studies of the Wilson's snipe in the Humboldt Bay Region.
M. S. Thesis.
Humboldt State College, Arcata.
90 p.
White, M., and S. W. Harris. 1966. Winter occurrence, food and habitat use
of snipe in northwest California.
J. Wildl. Mgmt. 30(1):23-24.
Whitehead, C., Jr. 1965. Foods and feeding habits of the common snipe
(Capella gallinago delicata) in Cameron parish, Louisiana, with ecological notes and a discussion of methods of sexing and aging. M. S. Thesis.
Louisiana State Univ., Baton Rouge. 200 p.

"l)

_(~

~t\.A"l.t:;,x)""".

Prepared by
.

\\

J

~)

c=t\\i\~\.:.£\c:V~

Bruce R. JohnsbnL\
Graduate Student

�-196. STATE OF COLORADO

APPENDIX A

.John A. Love, Governor
DEPAHTMENTOF NATURAL RESOURCES

DIVISION OF "VILDLIFE
Harry n. Woodward, Director
6060

Broadway ..

Deriver, Colorado 80216

• B25~1192

August 27, 1973

Dear Sir:
Because of your responses on the 1972 Colorado Division of Wildlife
Small
Game quest:ionnai.re~we
are asking for you!' assistance
in obtaining
information on the common snipe.
As you may know, . snipe maybe hunted f rom
September 1 through November 4 east of the Continental
Divide, and September 29 through October 12, 1973 'andNovember3,
1973 through January
20, 1974 'v'est of the Continental
Divide.
We would like you to read the
en cl.osed q~estiq!'-!!.aire, .keep t he q~.!~sticn~·-in,
and return
qu~stiuruiai.i.·i:: w.i.Lll your response:,;cu: the se ason+s end.

-~nd"

We have also enclosed a large envelope and ~~uid like to ask you to
return one .wing £romeach snipe you bag during the coming season.
Please
returntheeIivelop~
at the close of the season,
This will proVide us
with the' age of each bird you bagged.
.
Your cooperation will greatly aid usd.n understanding
this relatively
unknown game species
and provide a basis for its 'future. management.
Sincerely,

Howard D.'Funk
Section Chief
Small Game Research
CEB:al
Encl.

DEPARTMENT OF NATURAL RESOURCES,T. W. Ten Eyck, ExecutivoDirector.
Wiliiam VI/. Robinson, Vic.eChairman. Ford Strong, Secretary.
Harry Cum!"" Mnmber.

R. Wil/)nrs Cool. Member. Criarte-

WILDUFECOMMISSIOr,,;

Dp.anSuttle, Chdirman .

Dr. .J.K. Childre.:;,;,Mornbor « Wilton W. C09swell.Jr., Momber

A.GHhauer, Mnmbor. OrcSI Gorbaz, Member .LeRoy r~oh"on, Momhor

I'

�-:.197,..
APPENDiX B

SNIPE HUNT~R
REPORT

1.

Did you hunt snipe during the 1973 season?
Yes
No
If answer
was "No" you need not fill
out the remainder of the questionnaire.
(If
you hun'ted or did not hunt,please
return the questionnaire
in the small
stamped,. self addressed envelope).

2.

Countyor

are~ most frequently

hunted.......;..._.--:..

.......;..._-"--

3. Howmany days did you hunt snipe in the 1973 season?

4.

Howmany hunters

5.

What waS the approximate
snipe season ?
--

6.

How many srripe

7.

How often

were in your party

di.d ~

during

total

while

snipe hunting

number of snipe

bag during

the 1973 snipe

I

Often (10 or more days)

I

I

Frequently

NAME:

~LING

note

'--- __

ADDRESS:

'---

that

_

the 1973

season?

_
P1ease

(4 - 9 days)
(1

3 days)

Rare ly (only if accidentally
other game species)
Optional-Please

year?~

did you hunt only snipe?

I

Occasionally

this

yoti saw during

the 1973 snipe

season

-------------

flushed

while hunting

you are not ob.l Lgat ed to give your name and address.

~~-----,..-------~----.,.--

_

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                  <text>-1-

January
JOB PROGRESS

State

of

Work

REPORT

----=CO:::::.;:LO;:::.::RAD=.;:O
_
No.

W-4l-R-24

Bighorn

Sheep &amp; Mountain

Plan No.

1

Job NO.

1_6

Project

1975

Goat Inyestigations

~----------------

Job Title _E:.v.:...a:.l:.u.:..a.:..t.:..l.:..·
o:..:n~o:..:f=-t:..:h_e~N_u:....t_r_i_t_i_o_n
....
a....
l_R:....e....:q:....u_i
....
r..:.e_m_e
....
n....
t..:s--:.o
....
f--=-B..::i£g.;.:h.;;..o..::rn~;;:.Sh~e.;;.
_
Period

Covered:

Personnel:

June 1, 1973 - May 31, 19,/4

Gene G. Schoonveld

ABSTRACT
Samples of majoI' forage species used by bighorn sheep on a year-round basis
were chemically analyzed to determine nutrient content.
Forages analyzed
were Carex spp., Kobresia beliardi, Poa spp., Fescue spp., Agropyron spp.,
and Salix glauca collected on the Buffalo Peaks and Pikes Peak sheep ranges
and Carex spp., Fescue spp., Muhlenbergia
spp., Bouteloua gracilis, Artemisia
frigida and ~olodiscus dumosa on the Trickle Mountain sheep range.
Chemical
analyses were for amounts of soiuable and insoluable ash, crude protein and
relative percentages of plant cell walls and cell contents.
The cell wall
fraction was further analyzed for amounts of cellulose, hemicellulose,
and
lignin and the.ash fraction for amounts of calcium, phosphorus, sodium,
manganese, zinc, iron and copper.
Evaluation of the nutritional
quality of forages used by bighorn sheep on
the Buffalo Peaks, Pikes Peak and Trickle Mountain ranges was made by comparing nutrients available in forages to minimum nutrient requirements
that
have been established for domestic ruminants.

��-3-

EVALUATION OF THE
NUTRITIONAL REQUIREMENTS OF BIGHORN SHEEP
Gene G. Schoonveld

P. S. OBJECTIVE
To develop methods: (a) for the evaluation of nutritional requirements of
bighorn sheep under natural conditions, and (b) to measure the adequacy of
selected ranges to meet these requirements.

SEGMENT OBJECTIVES
1.

To estimate the nutritional
sheep.

level of forage species utilized by bighorn

2.

To evaluate if the nutritional level of selected bighorn sheep ranges
is adequate to meet nutrient requirements of bighorn sheep at all seasons
of the year.

METHODS AND MATERIALS

Nutritional Quality of Forage
Species Used by Bighorn Sheep
Methods used to determine forage species used by bighorn sheep and means of
collecting representative samples were previously described (Schoonveld 1973,
Keiss and Schoonveld 1974). During this segment, samples of major forage
species used by sheep were chemically analyzed to determine nutrient content
using procedures detailed by Keiss and Schoonveld (~cit).
Forages analyzed
were Carex spp., Kobresia. bellardi, Poa spp ,, Fescu~ spp ,, Agropyron spp ,,
and Salix glauca collected from the Buffalo Peaks ffildPikes Peak sheep ranges
and Carex spp., Fescue spp., Muhlenbergia spp., Bouteloua gracilis, Artemisia
frigida and Holodiscus dumosa from Trickle Mountain.
~hemical analyses were
for soluable and insoluable ash, crude protein, ether ext~act and relative
percentages of plant cell walls and cell contents. The cell wall fraction was
further analyzed for amounts of cellulose, hemicellulose, and lignin constituents and the ash fraction for amounts of calcium, phosphorus, sodium, manganese, zinc, iron and copper.
Relative proportions of plant species comprising sheep diets would not be
reliably estimated (Schoonveld 1973, Keiss and Schoonveld 1974) and actual
diets simulated, therefore only mean values of the nutrients contained in
major forage species used by sheep are reported. The actual nutrient intake
of sheep would most likely be greater than values reported due to selectivity
and ingestion of greater proportions of the more nutritious plant species.

�-4-

Evaluation of the nutritional quality of bighorn sheep ranges was made
by comparing nutrient levels available in forages used by sheep on the
Buffalo Peaks, Pikes Peak and Trickle Mountain sheep ranges. Where
possible, levels of nutrients contained in forages used by bighorn sheep
were compared to minimum requirements established for domestic ruminants.
Hopefully, such cqmparisons will offer clues as to possible nutrient
deficiencies.

RESULTS AND DISCUSSION

Forage Analysis
Crude Protein
The amount of crude protein contained in a forage, together with the relative percent of cell contents may be used as an arbitrary indicator to predict nutritive value (Van Soest and Moore 1965), higher amounts of protein
and cell contents indicative of better quality.
Protein availability (means) in representative forage samples used by bighorn sheep and collected from the three study areas is graphed in Figure 1.
Percentages of crude protein contained in all forage species were highest
in the leaf and seed stage, then declined with advancing maturity.
Protein
content and seasonal trends observed in this study were similar to values
reported by Johnston et al. (1968) in studying nutrient quality of alpine
plants.

Plant Cell Walls and Cell Contents
Plant cell wall constituents contain partially nutritive and non-nutritive
material.
Nutritive material includes cell wall carbohydrates, cellulose,
and hemicellulose which are partially digested by microbial firmentation
within the rumen. Non-nutritive material is composed of lignin and insoluable ash (mostly silica). These substances have no known nutritive
value to ruminants (Harris 1970).
Cell contents of plants include readily available nutritive matter (carbohydrates,proteins, lipids and soluable ash) that are digested by rumen microbes
or enzymes secreted in the digestive tract. Ruminant digestion trials have
shown cell contents of all forages are almost completely digestible (Van
Soest and Moore 1965). The relative amounts of cell walls and cell contents
contained in forages may therefore be used to predict nutritive value.
Relative proportions of cell walls and cell contents contained in forages
used by bighorn sheep are shown in Figure 2. Percentages of lignin, hemicellulose, cellulose and total ash contained in the cell wall fraction are
also shown.

�-5-

Soluable and Ihsoluable Ash
The insoluable ash fraction contains minerals that are bound in complex
compounds which cannot be extracted from the food and consequently have
no nutrient value. The soluable ash fraction contains those minerals
which can be extracted and are essential for proper nutrition.

Calcium and Phosphorus

(Ca,P)

Adequate amounts of calcium (Ca) and phosphorus (P) are especially important for ewes during pregnancy and lactation.
Both elements are required
for skeletal development and production of milk. Phosphorus is also vital
in vitamin and enzyme activity and therefore related to metabolism of almost
all other nutrients.
Levels of both Ca and P contained in forage species
used by bighorn sheep are shown in Figure 3.

Sodium (Na)
Sodium is essential in the osmotic regulation of body fluids and maintaining
the acid-base balance of the body. Some sodium required by an animal is
ingested in the form of sodium chloride (Na Cl), however much Na is supplied
in forages. Levels of Na found to be present in for'age.sused by sheep during
this study is shown in Figure 4.
Copper (Cu)
Copper is known to be essential in haemoglobin formation and production of
red blood corpuscles.
It is also a component of many enzyme systems and is
necessary for hair pigmentation (NRC 1968). Amounts of Cu contained in
samples of forage specieH utilized by bighorn sheep are graphed in Figure 5.
Zinc (Zn)
Several enzymatic reactions within the body require zinc. Diets of ruminants
normally furnish adequate amounts of zinc, however minimum requirements have
not been well established.
Amount of Zn available in forages used by bighorn
sheep are given in Figure 6.

Iron (Fe)
Iron is also essential in the formation of haemoglobin and is a component of
many enzymes. The bodies requirement for additional iron is normally low
except as a result of hemorrhage or pregnancy.
Normally iron is available
for recycling within the body upon destruction of haemoglobin (McDonald et
al. 1969). Amounts of iron contained in forages used by bighorn sheep are
given in Figure 7.

�-6-

Manganese

(Mn)

Manganese serves as an activator of enzyme reactions involving carbohydrates, protein and lipid metabolism.
Amounts of manganese determined
to be in forages used by bighorn sheep on the Buffalo Peaks, Pikes Peak
and Trickle Mountain ranges were 72-160 p.p.m., 94-330 p.p.m., and 81-205
p.p.m., respectively.

Nutritional Quality of Bighorn Sheep Ranges
Nutrient requirements of bighorn sheep would vary throughout the year
changing as the animal advances in age or as it proceeds through its
annual life cycle (growth period May-October; breeding November-December;
and gestation or lactation for ewes from December·-June). Minimum nutrient
requirements for bighorn sheep during these periods however, have not been
established.
It is necessary therefore to compare data on nutrients available in forages used by bighorn sheep on the three study areas to minimum
requirements established for domestic ruminants.
It io unlikely bighorn
sheep and domestic ruminants would have the same minimum requirements for
all nutrients, however such comparisons may indicate possible deficiencies
and levels of nutrients which require more in-depth investigation.

Crude Protein
Insufficient protein intake results in poor feed efficiency and reduced food
intake that eventually limits growth, muscle development and reproduction
(NRC 1968). Minimum yearly protein requirements for bighorn sheep are not
known. Hebert (pers. comm.) fed yearling bighorn ewes a 4 percent protein
ration during an extended period over winter and observed negligible body
weight changes. Based upon this work and known protein requirements of
domestic ruminants it appears protein availability in forages used by bighorn sheep on the three study areas would be sufficient in furnishing minimum
body requirements (Fig. 1). Percent protein contained in forages used by
bighorn sheep on all areas would also appear to be adequate for ewes during
gestation and lactation when compared to requirements of about 8 percent
established for domestic sheep (NRC 1968). Although data suggests Trickle
Mountain sheep receive less protein in their diets compared to either Pikes
Peak or Buffalo Peaks sheep, more definitive data need to be obtained relating to body condition indices before effects of less protein intake can
be assessed.

Plant Cell Wall Constituents

and Cell Contents

Amounts of cell wall constituents were higher in forages used by Trickle
Mountain sheep than were constituents in forage~ used by either the Buffalo
Peaks or Pikes Peak sheep (Fig. 2). These data indicate alpine forages used
by Buffalo Peake, and Pikes Peak sheep would be higher in nutritional quality
and more easily digestible than forages used by Trickle Mountain sheep,
however data needs to be analyzed for both statistical and biological significance. Biological significance of these results would need to be further

�-7-

evaluated by means of digestion trials preferably using sheep to determine amounts of cell wall constituents that contribute to the nutritional
level of the animal.

Calcium (Ca) and Phosphorus

(P)

Intake levels of 0.24-0.32 percent Ca and 0.16-0.20 percent P are considered minimum requirements for domestic ewes during gestation and lactation (NRC 1968). Defiency symptoms of inadequate Ca and P in an animal's
diet are slow growth, roegh appearance and listlessness.
A defiency of P
results in weak lambs due to inade.quate milk production by its dam and
failure of animals to fatten. Severe P deficiency is associated with increased parasitism and often diagnosed as starvation (Tillman 1961). The
Ca:P ratio must also be considered when determining adequacy of these two
elements in a diet. A high ratio may be as harmful as a defiency of either
element. The optimum Ca:P ratio is 1:1 or 2:1 for domestic livestock.
Forages used by bighorn sheep on the three study areas would supply ample
Ca on a year-round basis (Fig. 2) when compared to minimum requirements
established for domestic sheep. Amounts of P furnished in forages would
appear to be adequate only during summer for Pikes Peak and Buffalo Peaks
sheep and deficient during all other periods. Forages used by Trickle
Mountain sheep would appear to be P deficient on a year-round basis (Fig.
3). Calcium to phosphorus ratios in forages are also extremely wide for
all ranges which may further compound a P deficiency.
It is not known to what extent free ranging animals can select forages high
in P or to what degree they can accumulate P in storage depots for use when
a deficiency exists. Simesen (196l) states, "bone mineral is readily mobilized
to maintain the level of serum calcium, but less readily to maintain that of
P." Serum inorganic phosphorus levels,should therefore be established in bighorn sheep from the three study areas and correlated with P levels in forages
to better evaluate possible P deficiencies.

Sodium (Na)
Inadequate intake of Na by an animal reduces efficiency of food metabolism and
thereby slows growth. Generally, a diet containing 0.6 percent Na is considered adequate for domestic sheep, however minimum requirements have not
been established (NRC 1968). Forages used by bighorn sheep on the three study
areas would furnish less than 0.6 percent Na (Fig. 4), however use of natural
mineral licks by bighorn sheep and salt supplementation would increase Na intake to much higher levels.

Manganese

(Mn)

Minimum dietary requirements of Mn for domestic sheeF are unknown. In research studies with cows, Bentley and Phillips (195l) produced deficiency
symptoms with natural forages containing less th~~ 10 p.p.m. Mn. Generally,
dietary intake of 15-30 p.p.m. Hn would furnish sufficient amounts for most

�-8-

domestic ruminants (Tillman 1961). Forages used by bighorn sheep on the
Buffalo Peaks, Pikes Peak and Trickle Mountain ranges contained 72-100
p.p.m., 94-330 p.p.m., and 81-205 p.p.m., Mn respectively, and it is
therefore unlikely that sheep utilizing these ranges would suffer Mn
deficiency.

Copper (Cu)
Underwood (1966) pointed out that minimum Cu requirements for ruminants
cannot be clearly established due to various interfering factors. NRC
(1968) suggests at least 5 p.p.m., Cu is essential in diets of pregnant
and lactating domestic ewes, and nursing lambs. Continuous intake of Cu
in excess of bodily requirements may however be toxic. Susceptibility
of ruminants to Cu poisoning is partly due to excess Cu being stored in
the liver where it becomes a cumulative poison (Tillman 1961). Symptoms
of inadequate copper in the diet are generally seen in young animals;
deficiencies are evidenced by poor growth, bone disorders, scours, lack
of hair pigmentation, anaemia, gastro-intestinal disorders and lesions in
the brain stem and spinal cord (NRC 1968, Tillman 1961, Church et al. 1971).
Based upon requirements of domestic sheep, amounts of Cu available in forages
used by bighorn sheep and collected on the three study areas are sufficient
to meet bodily requirements (Fig. 5). No observable deficiency symptoms were
evidenced by bighorn sheep during the field collection phases of the study
or when lambs were collected for pathological investigations.

Zinc (Zn)
Itching, parakeratosis, retention of the placenta, loss of body wool and
development of thick, wrinkled, pink skin in domestic sheep are thought to
be caused by insufficient amounts of Zn in the diet (Church et al. 1971).
Minimum dietary requirements of Zn by domestic livestock however has not
been well defined due to many interfering factors such as the effort of Ca
or Zn requirements.
Generally, forages containing 30 p.p.m., are considered
adequate in supplying minimum dietary requirements of Zn for domestic
ruminants (Tillman 1961). Based on these requirements, data obtained from
analysis of forages used by sheep would suggest animals receive ample amounts
of Zn on a yearly basis (Fig. 6).

Iron (Fe)
With rare exceptions, Fe deficiency does not appear to be a problem in
ruminants consuming solid foods (Church et al. 1971). Til:man (1961)
suggests diets of grazing animals should contain 80-150 p.p.m., Fe to supply
bodily requirements.
Compared to these requirements, data obtained from
forage analysis suggests bighorn sheep on the three study areas would be
receiving insufficient Fe in theird~ets
(Fig. 7), especially sheep on the
Buffalo Peaks and Pikes Peak ranges. Again, use of natural mineral licks
by sheep could supply additional Fe in their diets.

�-9-

LITERATURE

CITED

Bentley, O. G., and P. H. Phillips.
1951. The effect of low-manganese
rations upon dairy cattle. J. Dairy Sci. 34:396-403.
Church, D. C., J. P. Fontenot, G. E. Smith, and A. T. Ralston. 1971.
Digestive physiology and nutrition of ruminants, Vol. 2 - nutrition.
D. C. Church, O.S.U. Book Stores, Corvallis, Ore. 80lpp.
Harris, L. E.
animals.

1970. Nutrition research techniques for domestic and wild
L. E. Harris, Logan, Utah.

Hebert, D. M. (personal comm.). Dept. of Zoology, Univ. of British
Columbia, Van., B.C. Canada.
Johnston, A., L. M. Bezeau, and S. Smoliak. 1968. Chemical composition
and in vitro digestibility of alpine tundra plants. J. Wildl.
Manage.
32(4):773-777.
Keiss, Robert E., and G. G. Schoonveld.
1974. Bighorn sheep and mountain
goat investigations - evaluation of the nutritional requirements of
bighorn sheep. Colo. Div. of Wildl., Game Res. Div., Fed. Aid Proj.
W-4l-R-23, Game Res. Rpt. (in press).
McDonald, P., R. A. Edwards and J. F. D. Greenhalgh.
tion. Oliver and Boyd. Edinburgh.
407pp.

1969.

Animal nutri-

National Research C&lt;:uncil. 1968. Nutrient requirements of domestic animals.
No.5.
Nutrient requirements of sheep. Nat. Acad. of Sei., Washington,
D.C. Pub. 1693.
Schoonveld, Gene G. 1973. Bighorn sheep and mountain goat investigations
evaluation of the nutritional requirements of bighorn sheep. Colo.
Div. of Wildl., Game Res. Div., Fed. Aid Proj. W-4l-R-22.
Game Res.
Rept., Jan. Part 2. pp. 75-87.

-

Simesen, M. G. 1961. Calcium, inorganic phosophorus, and magnesium metabolism
in health and disease. In Clinical Biochemistry of Domestic Animals.
Edit. C. E. Cornelius and J. J. Kaneko. Acad. Pr.ess, New York.
Tillman, A. D. 1961. Mineral nutrients for livestock.
Assoc. 138(11):600-602.

J. Amer. Vet. Med.

Underwood, E. J. 1966. Trace elements in human and animal nutrition.
Academic Press, New York, N.Y.
Van Soest, P. J., and L. A. Moore. 1965. New chemical methods for analysis
of forages for the purpose of predicting nutritive value. Proc. 9th
Inter. Grassl. Congr., Sao Paulo, Brazil. pp. 783-789.

~/~:~lU«&gt;;;/

Prepared by /~_
Gene G. Sc oonveld
Asst. Wildlife Researcher

�PROTEIN

14\
I

12

10

~/
W

u

6

a:

~

~O

________

----.:~--

I

C~

W

n. 6

C______

X

0

0-0

x-x
0'-0

JULY

SEPT

NOV

I

---0

0

~

X

a

4 .•

MAY

t-'

____

C

BUFFALO
PKS,
PIKES
PI&lt;.
TRICKLE
MTN.

"AN

MAR

Fig. 1. Protein availability (means) in major forage species used by bighorn sheep and collected from
the Buffalo Peaks, Pikes Peak and Trickle Mountain ranges May 1972-March 1973.

�~~.:!.hJ

..•k S r-i

SSSsss:s:J- C E LL.ULOSE

L

!-

H~MIC ElLY! ..OS \:!

WOW.\K&amp;.l"

L UUH ~

80
TN
70

BP

TM
TM

CD

..l

...I
C

•

ppTM

TM

TM

PP

60

tiP

pp

8P

PP
BpPP

BP
PP

50

..l
...I
..,

I

(,) 40

t-£
t-£
I

•••
z

.., 30+
o

CII:
III

A.

20
10
l.

t;! VI _J,k==-.

MAY

==

~\ID ~;I ="4=
JULY

Jl .~ ~1._._._
SEPT

:i

~~1._

NOV

.. .
JAN

MAR

Fig. 2. Percent cell walls (means) contained in major forage species used by bighorn sheep on the
Buffalo Peaks (BP), Pikes Peak (PP), and Trickle Mountain (TM) sheep ranges. Relative amounts of
lignin, hemicellulose, cellulose, and ash contained in the cell wall fraction are also shown.

�CALCIUM
0;8

I

0---

+

0.7

/

X/

0.6

0.5

e

min.

0.4
I

PHOSPHORUS
~
Z
UJ

o

I

N
I

-

0.18

&lt;,&lt;,

0:

~

I

t-'

0.14f~

0-0

au FFALO

X-x

PI KfS

t::t-oTRICKlE
0.1 0 ~

&lt;,

"~

0.06t

PKS.

PK.
MTN.

--------o~

~

o

X

tJ

-x:
min. P r e q,

e

X
O. I 6- O. 20 0/0

( NAS 1968)
MAY

JULY

SEPT

NOV

JAN

MAR

Fig. 3. Calcium and phosphorus content (means) in major forages used by bighorn sheep and collected
from the Buffalo Peaks, Pikes Peak and Trickle Mountain ranges, May 1972-March 1973.

�SODIUM
0.7
min. Na r e q, or 0.6%

0,6

(HAS 1968)

0.5

c

~o
I

•...
0.4

too"

z

t:J

w
u
c:

W
I

............................
------x
.:---------

........

w 0.3

Q"

0.2

0

BUFFALO
PKS.
PIKES
Pt&lt;.
0-0 TRIC Ir1..E MTN
0-0

x-x

Orl

MAY

JULY

SEPT

NOV

JAN

hi A~

Fig. 4. Sodium availability (means) in major forage species used by bighorn sheep and collected
from the Buffalo Peaks, Pikes Peak, and Trickle Mountain ranges, May 1972-March 1973.

�-14-

COPPER

tu)

25

O-OBUFFALO

PKS.

PIKES
Pt(.
a-a TRICKLE NTH.
~Jt

~115.~

~X~:.

~

0--

10

_

~

5

".-

0

MAY

a----------D---------~

J"LY

X.............

0-

~

~~o

·0

NOY

SEPT

JAN

MAR

Fi.g. 5. Amounts of copper (means) contained in major forage species
used by bighorn sheep and collected from the Buffalo Peaks, Pikes
Peak, and Trickle Mountain !anges, May 1972-March 1973.

ZI NC (Zn)

100

1

min Zn re q. ;;! 30 PP'I'l

80~

~:------x

60

-===-~7'

-~

40
0-0 BUFFALO

PKS.
PK.
c-/:l TRI CKLE MTN.

x-x PIKES

20

MAY

JULY

SEPT

NOY

JAN

MAR

Fig. 6. Amounts of zinc (means) contained in major forage species
used by bighorn sheep and collected from the Buffalo Peaks, Pikes
Peak, and Trickle Mountain ranges May 1972-March 1973.

�-15-

I RON

(Fe)

80

10

MAY

)f--)(

BUf'FAL'() PKS.
PIKES
PIC_

CI-Q

TRICKLE

0--0

'0
JULY

SEPT

NOV-

JAN

M TN.

MAR

Fig. 7. Amounts of iron (means) contained in major forage
species used by bighorn sheep and c31~eeted from-the Buffalo Peaks,
Pikes Peak, and Trickle Mountain ranges May 1972-March 1973.

��-17-

January 1975

JOB PROGRESS REPORT

State cf

.::CO:::..;:LO:;::,RAD:::.::::.O:::------

Project No.

W-41-R-24

Work Pl.an No.

1

Job TLtle

Bighorn Sheep &amp; Mountain Goat Investigations
Job No.

17

---------------------------

Ma
__n_i~p_u_l_a_t_i_o_n
__o_f
__v_e~g~e_t_a_t_i_o_n
__o_n
__B
__i~g_h_o_rn
S_h_e_e~p
__R_an~g~e_s

Period Covered:

_

June 1, 1973 to May 31, 1974

Personnel: George Bear, Julius Klein, Judson Brown, Dale Stahlecker, William
Adrian, llobert Keiss, Thomas Poj ar, .and Steve Steinert.

ABSTRACT

Study plots near Cathedral, Colorado were treated with varied applications
of nitrogen and phosphorous fertilizers, and 2,4-D herbicide in 1971-72.
Two areas were selected for study sites: (a) c~ alpine range, and (b) a
ponderosa pine-bunchgrass range. Nitrogen fertilizer was applied at 0, 30,
60, and 90 lbs. per acre; phosphorous fertilizer at 0 and 30 Ibs. per acre;
and 2,4-D herbicide at 0 and 2 Ibs. per acre. The effects of these treatment·s are being monitored.
There were few changes in plant composition que to the nitrogen and phosphorous
treatments. Chenopodium sp. increased on nitrogen plots. Herbicide treatments caused decreases in forbs and shrubs.
The alpine plots did not measurably respond to any of the fertilizer treatments.
On the lower study area herbage production generally increased with increased
nitrogen rates. Production also increased with the addition of the herbicide.
Plant species were collected and analyzed for protein, phosphorous, and calcium.
On the lower area protein levels were higher in plants collected from the
nitrogen-treated plots. These higher protein levels were maintained throughout
the winter. Plant phosphorous and calcium levels appeared to be unaffected by
the various treatments. The chemical content of the alpine plants were not
obviously influenced by the fertilizer and herbicide treatments.
Elk on the lower area exhibited a pronounced ~reference for the nitrogentreated plots. So few observations were made of deer and bighorn use of both
study areas that no inferences of treatment preferences could be drawn.

��-19-

MANIPULATION

OF VEGETATION

ON BIGHORN SHEEP RANGES

George D. Bear

P. S. OBJECTIVE
Improve the herbage yield, vegetative density and vegetative
selected bighorn sheep ra~ges in Colorado.

composition

on

SEGMENT OBJECTIVES
1.

Alter the quality and quantity of forage on specific study areas with
varied treatments of chemical fertilizers and herbicides.

2.

Determine the effect of each treatment on the composition,
and chemical content of plant species.

3.

Determine grazing preferences
to each treatment.

of wild bighorns

production,

on the area with respect

METHODS AND MATERIALS
The study areas are located about 35 miles south of Gunnison, Colorado.
This
bighorn herd winters in two areas: in a low (8,000 feet elevation) bunchgrass
and ponderosa pine type, and in a high (11,000-14,000 feet elevation) alpine
type. This is the third year of a five-year study, therefore, the procedures
outlined here are being used in a continuing study. The following work is
being conducted on each winter range.
1. Bighorn preference and response of the vegetation was measured on
plots treated with 2,4-D (herbicide), nitrogen, and phosphorus.
Two levels
of 2,4-D treatment (0 and 2 lbs./acre) 'were evaluated.
Four levels of
nitrogen treatment (0, 30, 60, and 90 lbs./acre); and two levels of phosphorous
treatment (0 and 30 lbs. /acre) were examined.
These we're evaluated in all
possible combinations, or a 2x4x2 factorial for a total of 16 treatments, as
outlined in the following table:
Treatment
Number
1
2
3
4
5
6
7

8
9
10
11
12
13
14
15
16

2,4-D
0
0
0
G

0
0
0
0
2
2

2
2
2
2
2
2

Treatment
Nitrogen
0
0
30
30
60
60
90
90
0
0
30
30
60
60
90
90

Phosphorus
0
30
0
30
0
30
0
30
0
30
0
30
0
30
0

30

�-20-

Treatment plots, measuring 50 feet by 100 feet, have been established.
Each plot corner was marked with a metal fence post. These markers were
numbered to assist in identifying plots when bighorns are grazing on them.
Each treatment was randomly r.eplicated 3 times on each study area. Each
replication consisted of the 16 treatment plots listed on the preceding
page. Plots in each replication were arranged in a rectangular area eight
plots long and two plots deep. Randomized selection of each plot treatment
was determined by using a table of random numbers.
2. Frequency sampling. Transects were established on each plot to
determine changes in vegetative frequency which may result from the various
treatments.
Frequency (p) is defined p=m/n, where m is the number of quadrats
containing a given species and n is the total number of quadrats observed in
each treatment.
Four transects were established in each plot. Transects
paralleled the base of the plot with approximately 10-foot spacing between
transects.
A total of 15 quadrats are located at 6-foot intervals along each
transect, or a total of 60 quadrats for each treatment plot. Preliminary
sampling work indicated a quadrat size measuring l6x!6 inches should be used
on the low winter range at Cebolla Creek; and a split-sampling scheme should
be used on the Alpine Area. Two quadrat sizes are used on the alpine: Lxl, inch
for measuring Trifolium nanum and Carex sp., and 8x8 inches for all other
species.
Frequency percentages were derived for all plant 3pecies encountered
on the quadrats.
To be considered as occurring on a given quadrat, the center
of an individual plant or one-half of its basal area must be inside the quadrat.
An "individual" for bunchgrasses was a clump; for single stem species, such as
Western wheatgrass, it was individual stems; for cactus it was a lobe; and for
shrubs and forbs it was the basal area of all stems. Obviously, these data
are acceptable for determining changes in frequencies for each individual
species only, and cannot be used to compare one species to another.
These
transects were read in June and July, when the vegetation was growing well,
but before the crowns became so large they interfered with observations.
Yield plots. Quanfity of forage produced on the treatment plots was
determined by using a Neal Electronics Herbage meter. The herbage meter
measures an area 24x12 inches x 18 inches, which is designated as yield plot.
The 60 plots established on each treatment for the frequency plots above were
also used for the yield plots. Obviously the 60 plots is considerably greater
than the recommended numbers given above; however, the vegetation will be
changing due to treatment and larger samples will be needed as variation increases.
At every twelfth meter read plot, all vegetation was clipped and weighed.
All green vegetation on these plots was clipped at the root crown and placed
in a labeled paper sack. These samples were oven dried and weighed to the
nearest gram. The weights were used to compute regression equations for
converting herbage meter readings into pounds of forage per acre.
Wire cages were placed on the study areas to monitor the effect of
grazing on the. area. Cages measuring 3 feet by 3 feet by 1 foot high were
placed on each area. Vegetation produced inside the cages (no grazing) was

�-21-

visually compared with vegetation produced on areas outside the cages
(grazed areas) to get a general idea concerning effects of grazing.
Photo points were established on each treatment. Photos were
taken at the end of the growing season to establish visual records of
changes occurring:in the vegetation due to various treatments.
Chemical analysis. Vegetation samples of the most common plant
species were collected from the treated areas. Samples were analyzed
for phosphorus, calcium, and protein content. This work was done at the
Fort Collins Research Center Laboratory.
Since the study area is a bighorn winter range, the vegetation was sampled at the completion of the
growing season, again in mid-winter, and in late winter.
3. Bighorn preference or selection of treatment plots was measured
by recording number of bighorns on each treatment plot at IS-minute
intervals during observation periods. Sex and age of the sheep, time of
day, general activity of the animals, weather conditions, and snow depth
(if present) were also recorded. Observations were made with a spotting
scope during daylight hours. Sampling periods extended throughout the
time the sheep occupied the winter range. Notations were also kept on
other animals (deer or elk) that use the study plots.
Records were kept on the snow depth on each treatment plot to relate
to bighorn preference for given plots. The fence post plot corner markers
provided permanent snow depth markers.

RESUL7S AND DISCUSSION
Fertilizer and herbid.de treatments were applied between the 1971 and 1972
growing seasons. Data presented here represent responses during two growing
seasons.

Monitoring Treated Plots

Frequency Sampling
Percent frequency of occurrence for plant species on the study plots are
presented in Tables 1 and 7.. Data from the three replications (180 sampling
plots) were combined to give a single percentage-figure for each plant
species within each level of treatment. This information is to be used to
measure changes that occur in vegetation frequency due to the various treatments.
To date, no measurable changes in plant composition are attributable to
nitrogen and phosphorous treatments. There was one excep td.on , Chenopodium
sp. on the Cebolla Creek area showed a marked increase on all nitrogentreated plots, with a greater increase on the higher treatments (90 lb. A.).
Also, Arabis crandelli increased on all plots, irrespective of treatment.

�-22-

As might be expected, the herbicide treatments caused some changes on
both areas. Artemisia·frigida and all shrubs on the Cebolla Creek area
decreased following the herbicide application.
The following species
showed significant decreases on the Alpine Area: Androsace ~eptentrionalis,
Artemisia scopulorum, Cerastium beeringianum, Erigerortpinnatisecta,
Erigeron simplex, .~eum turbinatum, Phlox caespitosa, Potent ilIa concinna,
Potentilla diversifolia, Silene acaulis, and Trifolium nanun.
In some
instances a few individual plants of the forbs persisted on herbicidetreated plots and were tallied during frequency sampling, but these
survivors were generally of poor vigor and contributed little to the overall production on the plot.

Yield Plots
Relationships of herbage meter readings to herbage weights on the clip-plots
are presented in Table 3. Many of the correlation coefficients are disappointingly low; however, when comparing the 1973 data with 1972 data it
would appear the poor correlations between meter readings and actual weights
is inherent in the plots themselves.
Regression lines were plotted for
each of the sixteen treatments.
There were sufficient differences between
these lines (apparently due to treatment) that it appeared these data
should not be combined.
Herbage production on control plots (treatment 0-0-0) has declined steadily
on both study areas during the past three summers (Table 4). Data from two
U. S. Weather Bureau Stations in the area also indicates a downward trend
in total precipitation (Table 5). Data in Table 5 were computed on the
basis of a period from September one year through August the next year,
which corresponds more closely to the vegetative growing season. Production
on treated plots showed a downward trend in 1971-1972, however, nearly all
plots increased in 1973. This increase. is apparently due to fertilizer and
herbicide treatments.
Due to differences in potential production on the various plots and annual
va~iations in production, figures in Table 4 were converted to a weighted
percentage (Table 6). The following formula was used to compute percentages:
b
a

x
c

Where: a = production
b

on control plot in 1971

production on control plot in
year under inspection

c = production
1971

on the treated plot in

x = estimate of production for the
treated plot in the year under
inspection
Then:

x - d
x

- y

Where: d

actual measurement of production
on the treated plot

y

percent variation from predicted
production due to treatment

�-23-

Herbage production on treated plots on Cebolla Creek showed a marked
increase from 1972 to 1973. In 1973 production responded to nitrogen
fertilizer and herbicide, generally increasing with increased nitrogen
rates; also production increased with the addition of the herbicide.
Plant production did not respond to phosphorous fertilizer treatments.
Plants on nitrogen treated plots started growth approximately 4 weeks
earlier in spring:1973 than plants on surrounding untreated areas; also
plants on the nitrogen plots were greener in color and more vigorous
looking. A more rigorous analysis of the data are needed to adequately
test treatment effects.
The Alpine Study Area does not appear to be undergoing any noticeable
changes in production readily related to treatment levels. Nearly all
treated plots showed an increase in production from 1972 to 1973, whereas, the control plots decreased in production.
Visually, the highnitrogen and phosphorus-treated plots were greener this year than in 1972.
Perhaps the Alpine Area will require a longer period of time than the
Cebolla Creek Study Area to respond to the treatments.

Chemical Analysis
Samples of the following plant species were collected from the Cebolla Creek
Study Area and analyzed for protein, phosphorous, and calcium: Artemisia
frigida, Muhlenbergia montana, Festuca arizonica, and Bouteloua gracilis
(Tables 7-10). Samples were collected at three times during the winter,
but only plants collected in the fall and mid-winter have been analyzed.
Protein levels were higher in plants collected f~om nitrogen-treated plots.
Plants collected from the 90-pound nitrogen treatments tended to be higher
in protein than those from other nitrogen treated plots. This basic pattern
persisted into mid-winter even as overall protein levels decreased.
The
1973 data indicated lower protein levels in these plants than in 1972; however, the plants from the control plots also e~~ibited lower protein levels.
Phosphorous and calcium levels in these plant species appeared to be unrelated to the various treatments.
Fall samples were reasonably close to
the 2:1 ratio (Ca:P) recommended as a minimum for domestic livestock; however,
winter samples had considerably wider ratios. The 1973 data indicated Ca:P
ratios nearer to a 1:1 ratio than the 1972 data, again the controls were
also lower.
The following plant species were collected from the Alpine Study Area for
chemical analysis: Kobresia bellardi, Poa sp., Trifolium nanum, Oreoxis
bakeri, and Geum turbinatum (Tables 11-15). The latter three species were
collected in fall samples only. Protein level~: in Poa and Kobresia showed
a minor increase on nitrogen-treated plots; but protein levels in forbs
were unaffected by nitrogen treatments.
However, these plants appeared to
be affected by phosphorous treatments, because calcium:phosphorous ratios
were lowered in all plant species. As on Cebolla Creek, the Ca:P ratio for
Poa and Kobresia during mid-winter were well outside that recommended for
livestock, which further points out the need for additional information on
the basic nutritional requirements for bighorn sheep.

�-24-

Grazing Preferences
Observations of animal preference during this segment was limited to elk
and deer. No bighorn sheep were observed on the study areas, but tracks
were observed.
Trend cotmts conducted last summer indicated this bighorn
herd decreased du~ing the previous year, and fewer sheep were seen in the
vicinity of the study plots this winter.
Only two rams were observed on
the Cebolla Creek Wildlife Area. Eighteen sheep were in close proximity
to the Alpine Study Area in January. Then, all but six dispersed to
surrotmding areas for the remainder of the winter.
Approximately 50 percent of the alpine range had snow accumulations of six inches or less throughout the winter.
This was largely due to high winds and light snowfalls.
Therefore, bighorns were able to remain dispersed and in small groups.
South-facing slopes on the lower Cebolla Creek Area remained free of deep
snow accumulations throughout the winter.
The only observed bighorn use on the Alpine Area was recorded on January 23.
There had been a new snowfall (2-3 inches) during the night. Fresh tracks
indicated a band of 18 sheep had moved across many of the plots on Replication 1 but the group had concentrated on the 90-30-0 plot, apparently feeding.
During the remainder of the winter sheep were observed near the area but
not actually on the plots. This may indicate the plots offered no pronounced
attraction to bighorns.
Elk on Cebolla Creek Area showed a definite preference for nitrogen-treated
plots. It was difficult to accurately record deer and elk usage on the plots
since they were most active during the nights. Most of the observations in
Table 16 were recorded during March when the elk started feeding later into
the daylight hours. The March data suggest elk preferred the 60 N plots,
but by this time the 90 N plots had already been heavily grazed.
Grazing behavior of elk on the plots was definitely related to treatment.
Elk would move along with their noses to the ground, taking an occasional
bite of forage tmtil they reached a plot treated with the higher (60-90 lbs.)
levels of nitrogen.
Then they would pause and begin grazing intensively.
They would graze to the edge of the plot then abruptly turn about so as to
continue grazing on the plot.
Visual inspection of the Cebolla Creek plots in April also indicated definite
preferences for nitrogen-treated plots. Untreated Arizona fescue plants
were 10-14 inches tall, whereas the fescue plants treated with nitrogen were
grazed down to 3-4 inch heights by spring. The boundary-line contrast was
very pronotmced.
Elk apparently are able to recognize and select nitrogen treated vegetation.
Similar preferential foraging by bighorn sheep and deer was not obvious.

Prepared by

~~c::P,~.

ceorg£.

Wildlife

Bear
Researcher

�Table 1.

Percent frequency of occurrence

Species

for plant species on the study plots treated with fertilizers

Treatment. Pounds Per Acre. Nitrogen - PhosEhorus - Herbicide
0-0-2 0-30-2 30-0-2
30-30-0 60-0-0 60-30-0 90-0-0 90-30-0

0-30-0

30-0-0

21
16

13
13
1

21
3

19
20

54
5
58

48
3
61

45
2
67

-

-

2

1

3
31

25

0-0-0

and herbicide,

Cebolla Creek, 1973.

30-30-2

60-0-2

60-30-2

90-0-2

90-30-2

G~
Agropyron smith!i
Bouteloua gracilis
Carex sp.
Danthonia Earryi
Festuca arizonica
Kocleria cristata
Muhlcnbcrgia montana
Muhlenhergia
richardsoni
Muhlcnbergia torreli
OryzoEsis hymenoides
Poa secunda
Sitanion hystrtx
Stipa ~
Unid. Grasses

-

-

-

1

-

-

-

-

1
2
42

-

-

-

-

-

8
6

3
13
9

16
8

12
15

51
7
53

72
2
75

46
0
43

49
3
58

52

--

-

-6-

1
29·

2
20

-

2
1
27

-

-

--

-

-

--

-

-

1
1
2
46

--

-

-

57

16
6
1
44
1
76

57
1
67

1

1

2

-

-

--

--

-

-

37

-

-

28

5
2

-

-

17

- . --

-

8
3

10
17
16

15
10

22
5

47

55

46

45

67

72

57

71

70

38
1
60

--

-

6
11
6

3
20
8

59

-

-

22
-

-

---

29

,

1

-

1
25

--

-

2
40

--

-

1

-

1

--

42

-

1

-

.1
1
50

--

\.II

I

Forbs
Androsace sp.
Arabis crandallii
Cas t i lle j a sp.
Chaenactis douglasii
ChenoEodium sp.
Erigeron sp.
Eriogonum sp.
Fragaria sp.
Geranium sp.
Heuchera sp.
Hymenoxys richardsoni
Lappu1a sp.

-----

I
N

-

7

-

1

-1

-

--

-

-

3
1

7
1

2

8
1

-

--

---

-

---

-

-

-

-

-

-

1

1
6
1

6

11
1

20
1

31
3

3

1
6
1

--

1
1
1

7

-2
3

1

-

1

-

1

-7

--

---

3

-

-

---

1

1

2

-

-

3

5

7

4

2

3

1

5

10

10

28
1
1

29

---

1

---

1

1

-----------------------

--

--2

�Table 1.

Percent frequency

Species
~

0-0-0

of occurrence

for plant species on the study plots treated with fertilizers

0-30-0

30-0-0

---

1

and herbicide,

Treatment. Pounds Per Acre. Nitrosen - PhosEhorus - Herbicide
90-0-0- 90-30-0
30-30-0 60-0-0 60-30-0
0-0-2 0-30-2 30-0-2

Cebolla Creek, 1973 (continued).

30-30-2

60-0-2

60-30-2

-

1

1

-

T

90-0-2

90-30-2

-

1

(continued)
Lepidium montanum
LeEtodactxlon Eungens
Mertensia lanceo1ata
Penstemon teucrioides
Potentilla concinna
Solidago decumbens
Senecio mutabi1is
Senecio sEartioides
Smelowskia calycina
Sphaeralcea coccinea
Va1eriana edulis Unid. Forb-s---·-

3
3

2

-

1

-T

-1
T

1

T

2
6
1

--

-

1

6
11

--

1

-

---

-

1

T

48

-

46

56

43

50

1

6

2

5

4

-

T

--

--

-

-

-

T

5
12
1

-

16

-3
-

1
1
1

2

-

-

-

-

1

-

-6
-

-T
-

-

1

-

-

T

1
1

1

1

-

---

---

31

33

-

43

1

-

3

3

T

60

48

40

1

1

2

-

-

-

---

1

1
T

-

--

'1-

1

-

2

40

32

32

1

-

25

1

-

2

1

1

--

T

T

~
Artemisia frigida
Artpmisin tridentata
Chr~sothamnus ~
Chrysothamnus
viscidiflorus
Echinocactus sp.
Jamesii americana
Rhus trilobata
Ribes .£!!.!!!!!

T

-1

-

T

--

-

----

T
4
2

1

-

T

31
T
5
T

-

4
T
1

-

---

T

--

-

--

-

-

-

-

T

---

-

-

T

T

-

-

I
N
C1\
I

�Table 2.

Percent frequency of occurrence for plant species on the study plots treated with fertilizers and herbicides,

Species

0-0-0

0-30-0

30-0-0

13

4
6
2

Treatm~~tl Pounds Per Acrel Nitrogen - PhosEhorus - Herbicide
0-0-2 0-30-2 30-0-2
30-30-0 60-0-0 60-30-0 90-0-0 90-30-0

Alpine, 1973.

30-30-2

60-0-2

60-30-2

90-0-2

6
4

T
5

28

9-30-2

--Grasses
1
Agropyron scribneri
17
~sp.
Deschampsia caespitos8
8
Festuca ovina
HelictotrICh'On
T
mortonianum
2
~
sEicatum
63
.f2! sp.
21
Tr1setum spicatum
51
~egi'l
bellardi

-

2
Achillea lanulosa
Androsace
seJ2tentrionalis
7
Arenar1a fendleri
Artemisia scoJ2ulorum 41
~
leJ2toseJ2a1a
Castilleja
occidentalis
Castilleja rhexifolia
Cerastium
17
beeringianum
Erigeron J2innatisecta 10
5
Erigeron simJ2lex
8
Eritrichium elongatum
Gentiana J2rostr~
21
Geum turbinatum
HaploEaEpus pygmaeus
2
Hj~enoxys grandif10ra
4
Hertensia ~

-

-

-2

-

-

-

14
3
22
1
T
51

12
27
3
37

-~

1
1

I

9

4

-

----------------------------

-

7
22

-

-.

-

8
35

-

T

2
2

2
15
4
4

5
12
2
5

10
11
2
7

22

24
4
1
4

25
T
3
2

-

-4
-

17
3
61
3
59

-

7

2

T

2

T
9
9

T
6
2

T
17
25

1

6
3

6
T
70
21
37

-

.,.

-

9
3
69
15
54

-

2

-T

4
3
62
11
4~

-

53
3
62

-

3
3
56
5
42

-

51
23
56

4
3
66
15
52

-

17
T
53
T
32

-

4

7
27

23

12
8
66
7
51

-3

77
T

13

6
4
66
7
47

17

2
2
18

16

10
7
'I
7

~5
7
67
13
56

18

~

8
6

12

-

-

3
20

-

-

8
5

2
12
3
23

2
1

1
7

6
36
T
14

4
7
T
9

7
9

-

T

-

-

14
23
1
2
2
24
2
2
6

16
15
5

-3
26
2
6

-

28
4
2

-

4
3

-

-

-

T

-

T

11

-

12
1
7
T

-3

-3
8

4

2

-

1
10

4

1

-

-

1
4
2
12

-

-

T

6
T

-----------------------------

-

-

-

17

23
8
15

.,68
21
53

4
62
27
51

T
76
16
56

2

2

--

-

6

-

'I
1
24
T

-

14
2
T

12

-

2
15
'I

-

-

--

T
T
18

T

25
1
11
3

2

7

-

-

7

6

t

T

5

1

8
'I
2
4

2

~

-2
-3
2

-

2
6

-

4
4

7

i
6

---

••
N
-...J

•

T
4

�Table 2.

Percent frequency of occurrence

Species

~

0-0-0

0-30-0

for plant species on the study plots treated with fertilizers and herbicides, Alpine, 1973 (continued).

30-0-0

Treatment I Pounds Per Acrel Nitrogen - PhosEhorus - Herbicide
30-30-0 60-0-0 60-30-0 90-0-0 90-30-0
0-0-2 0-30-2 30-0-2

30-30-2

60-0-2

60-30-2

90-0-2

90-30-2

47

50

62

61

83

64

-

T
24

26

-

31
T

17
T

23
10

11

14
13
16

--

11

7
3

30
1
3

-

-

2

T

-T

4

3

2

--

T

---

1
3

2

T

(continued)
Oreoxis bakeri
66
PedicularrsscoEulorum
Phacelia sericea
Phlox caespitosa
37
Podistera eastwoodae
Polemonium viscosum
2
P01ygonum bistortoides
4
Polygonum viviEarum
Potentilla concinna
24
Potentilla
diversifolia
T
Potenti11a
rubricaulls
11
Ranun cu Iua tnamoenus
3
~l_frllen cerna
Saxifraga chrysantha
Saxifraga flngcllarls
T
Saxlfraga rhomboidea
22
Sedum stenonetalum
SI'le'iie
acau Lfs
12
~skia
cnlycina
21
Thalictrum alpinum
30
Thlaspi alEestre
1
Trifolium dasyphy11um
3
Trifolium nanum
36
Trifolium Earryl
Unid. Forbs

-

-

-

-

-

Shrubs
~

niva11s

68

44

57

22
2

38

22

-

10

--

-

-3

57

-

25

-

14
9

14
T
39

-

1

-

25
-

1
7
T

10
3

3
T

3
1

6

30
2

-

28
T

-

28
16

-

T

1
22
T
T
22
24

--

T
29

-

-

T
13
2
11
21
11
T

-

24

30

-

-

-

-

-

-

-

31

-

8
24
11

-

32

--

59

61

57

-

21

-

35

38

-

-7
1
26

2
2

-

-

2
12
1
2
19
22

--

23

-

-

-

58

-

24
T

43

-

26

53

22

11

14

T

-

-

-

-

4

2
12

-

-

2
T

-

4
1
33

-

23
2
8
9
8
T

-

28

-

28

-

T

-

T
8

T
30
10
1

-

26
16

-

--

1
17
2
4

-

10

-

21

6
1
13

T
3
14

16

-

-

-

-

-

20

14

13
7
14

22

-

-

--

-

14
T
9
4
12
5
11

-

-

---

T

T

---

-1

-

2

--

21

3
T
16

21
T
6

T
12
2
11

15

29

14

14

17

11
1

3

19

2

T

-

-

1

T

I
N
00
I

�-29-

Table 3. Correlation coefficients (r) of herbage meter readings to oven-dry
weights of forage on meter-read plots.

Treatment
Nitrogen-Phosphorqs
Herbicide*

Cebolla Creek
1972
1973
r
r

AlEine
1972
r

1973
r

0-0-0

.70

.84

.87

.68

0-30-0

.78

.54

.83

.83

30-0-0

.70

.83

.80

.78

30-30-0

.57

.34

.67

.66

60-0-0

.87

.86

.75

.88

60-30-0

.68

.68

.83

.90

90-0-0

.86

.70

.88

.92

90-30-0

.81

.72

.53

.69

0-0-2

.74

.70

.56

•76

0-30-2

.57

.74

.85

.67

30-0-2

.81

.82

.75

.63

30-30-2

.85

.56

.55

.52

·60-0-2

.71

.65

.69

.56

60-30-2

.72

.75

.78

.68

90-0-2

.80

.67

.78

.79

90-30-2

.83

.79

.88

.82

* Treatment rates are in pounds per acre.

�-30-

Table 4. Herbage production (pounds per acre, dry weight) on the study
areas. Treatments were applied following the 1971 growing season and prior
to the 1972 growing season.

Treatment

.'1971

Al:eine
1972

1973

1971

Cebolla Creek
1972

1973

0-0-0

2158

1145

918

826

694

598*

0-30-0

2195

1663

1662

1455

627

872

30-0-0

848

1884

1270*

715

794

1743 **

30-30-0

2665

1234

1030

825

757

683

60-0-0

1444

1155

2293**

1241

827

1739*

60-30-0

1758

1238

1336

998

774

1200**

90-0-0

1667

1055

1251

998

801

1357*

90-30-0

1077

1456

773*

860

699

1577**

Mean

1726

1354

1316

990

734

1221

0-0-2

1934

871

1005*

998

577

1989**

0-30-2

1323

689

1234

1498

731

1538**

30-0-2

1247

837

1023

908

857

1657*

30-30-2

1051

841

845

1009

999

1148

60-0-2

1308

915

949

708

770

1639*

60-30-2

1178

1024

1134

917

908

1707*

90-0-2

1678

1023

999

1235

797

1194**

90-30-2

1562

1496

2312**

998

697

1905**

Mean

1410

962

1188

1034

792

1597

1973 production was significantly

** 90% level); paired - T test.

different from the 1972 production

(*95% level,

�-31-

Table 5. Climatological data reported by the U. S. Weather Bureau.
Average
temperatures and total annual precipitation was computed for a period from
September through August each year.

Year
Sampling Station

1969-70

1970-71

1971-72

1972-73

temperature

36.4

36.7

37.4

35.4

Total precipitation

13.8

11.6

8.9

12.3

37.6

34.9

10.2

12.6

Cochetopa Creek
Average

Lake City
Average Temperature

38.6

Total precipita.tion

20.4

14.9

�-32-

Table 6. Percent change in herbage production on the study plots
computed
from the base-year (1971) and the control-plots each year.
Treatment
N - P - H

Cebolla Creek
1972
1973

AlEine
1972
1973

0-30-0

.-49

-17

42

78

30-30-0

9

14

13

9

60-30-0

- 8

66

33

79

90-30-0

- 3

153

155

69

0-30-2

-42

42

- 2

120

30-30-2

18

57

51

89

60-30-2

18

159

64

126

90-30-2

-17

164

80

248

30-0-0

32

237

319

57

60-0-0

-21

94

51

273

90-0-0

- 4

88

19

77

0-0-2

-31

137

-15

24

30-0-2

12

152

26

93

60-0-2

29

220

32

71

90-0-2

-23

34

15

40

�-33-

Table 7. Chemical analysis of Festuca arizonica
Creek study area, 1973-7L;-.

Treatment
N - P - H

Protein

collected

c:;ollection Date and Anallsis
SeEtember
Ca:P
Phose Calcium Ratio
Protein

on the Cebolla

(Percent2
J'anuarl
Phose

Calcium

Ca:P
Ratio

0-0-0

5.9

.15

.19

1. 3:1

2.2

.05

.22

4.4:1

0-30-0

6.2

.19

.20

1.1:1

2.4

.05

.30

6.0:1

30-0--0

7.2

.17

.20

1.2:1

2.6

.04

.24

6.0:1

30-30-0

8.5

.18

.28

1. 6:1

3.0

.05

.28

5.6:1

60-0-0

10.4

.17

.20

1.2:1

3.5

.07

.20

2.9:1

60-30-0

9.4

.17

.25

1.5:1

3.9

.05

.34

6.8:1

90-0-0

8.7

.12

.18

1.5:1

4.3

.05

.35

7.0:1

90-30-0

10.8

.16

.26

1.6:1

6.2

.07

.25

3.6:1

0-0-2

6.4

.18

.24

1.3:1

1.8

.04

.19

4.8:1

0-30-2

7.4

.16

.21

1.3:1

1.8

.05

.18

3.6:1

30-0-2

8.0

.15

.30

2.0:1

2.2

.05

.29

5.8:1

30-30-2

8.6

.18

.30

1. 7:1

2.6

.06

.30

5.0:1

60-0~2

9.5

.17

.20

1.2 :1

3.2

.04

.34

8.5:1

60-30-2

8.4

.16

.25

1. 6:1

3.4

.05

.29

5.8:1

90-0-2

11.9

.18

.31

1. 7:1

4.8

.05

.31

6.2:1

90-30-2

12.3

.20

.31

1.6:1

4.4

.05

.29

5.8:1

�-34-

Table 8. Chemical analysis of Boute1oua
Creek study area, 1973-74.

gracilis collected on the Cebolla

Treatment
N - P - H

Protein

Collection Date and Ana1l:sis (Percent)
SeEtember
Januarl:
Ca:P
Phos. Calcium
Ratio
Protein Phos. Calcium

0-0-0

12.3

.18

.46

2.6:1

0-30-0

9.8

.23

.50

2.2:1

30-0-0

12.0

.13

.46

30-30-0

10.1

.18

60-0-0

11.4

.17

Ca:P
Ratio

3.5

.07

.41

5.9:1

3.5:1

5.4

.07

.40

5.7:1

.64

3.6:1

4.2

.06

.54

9.0:1

.54

3.2:1

5.0

.06

.35

5.8:1

4.2

.06

.58

9.7:1

60-30-0
90-0-0

12.3

.15

.58

3.9:1

7.1

.07

.41

5.9:1

90-30-0

14.9

.22

.50

2.3:1

5.8

.06

.45

7.5:1

0-0-2

9.2

.16

.58

3.6:1

3.9

.06

.40

6.7:1

0-30-2

9.3

.20

.57

2.9:1

3.8

.05

.42

8.4:1

30-0-2

12.7

.15

.64

4.3:1

5.1

.05

.45

9.0:1

30-30-2

11.5

.19

.46

2.4:1

5.1

.06

.45

7.5:1

60-0-2

15.0

.15

.64

4.3:1

6.2

.07

.41

5.9:1

60-30-2

14.7

.20

.60

3.0:1

5.4

.08

.47

5.9:1

90-0-2

14.1

.16

.55

3.4:1

6.7

.07

.51

7.3:1

90-30-2

14.0

.24

.51

2.1:1

6.4

.06

.42

7.0:1

�-35-

Table 9. Chemical analysis of Mublenbergia
Creek study area, 1973-74.
..

Treatment
N - P - H

Protein

montana

collected on the Cebolla

Collection Date and Anal~sis (Percent)
SeEtember
Janua!1
Ca:P
Phose Calcium
Ratio
Protein Phose Calcium

Ca:P
Ratio

0-0-0

4.2

.13

.37

2.8:1

2.1

.07

.31

4.4:1

0-30-0

6.8

.15

.36

2.4:1

2.3

.05

.31

6.2:1

30-0-0

8.6

.11

.31

2.8:1

3.3

.05

.29

5.8:1

30-30-0

8.2

.06

.34

5.7:1

2.5

.04

.35

8.8:1

60-0-0

9.9

.10

.35

3.5:1

3.8

.05

.28

5.6:1

60-30-0

8.9

.12

.33

2.8:1

4.8

.05

.36

7.2:1

90-0-0

9.6

.11

.31

2.8:1

5.2

.05

.38

7.6:1

90-30-0

10.6

.13

.40

3.1:1

4.5

.06

.29

4.8:1

0-0-2

7.9

.14

.29

2.1:1

3.0

.05

.30

6.0:1

0-30-2

7.3

.12

.40

3.3:1

2.7

.05

.31

6.2:1

30-0-2

9.9

.13

.33

2.5:1

2.7

.04

.34

8.5:1

30-30-2

9.0

.12

.31

2.6:1

2.4

.04

.31

7.8:1

60-0-2

10.0

.13

.46

3.5:1

3.3

.05

.40

8.0:1

60-30-2

8.4

.12

.30

2.5:1

4.2

.05

.35

7.0:1

90-0-2

10.0

.13

.43

3.3:1

6.4

.05

.42

8.4:1

90-30-2

10.9

.15

.43

2.9:1

4.2

.04

.42

10.5:1

�-36-

Table 10. Chemical analysis of Artemisia
Creek study area, 1973-74.

frigida collected on the Cebolla

Treatment
N - P - H

Protein

Collection Date and Ana1~sis (Percent)
SeEtember
Januar~
Ca:P
Phos. Calcium
Ratio
Protein Phos. Calcium

0-0-0

11. 0

.23

.54

2.3:1

6.4

.13

.58

4.5:1

0-30-0

9.8

.27

.54

2.0:1

5.8

.16

.64

4.0:1

30-0-0

11.8

.23

.64

2.8:1

7.8

.15

.55

3.7:1

30-30-0

13.6

.30

.55

1.8:1

6.8

.12

.73

12.0:1

60-0-0

14.2

.25

.58

2.3:1

8.0

.12

.54

4.5:1

60-30-0

11.8

.21

.53

2.5:1

7.8

.11

.74

6.7:1

90-0-0

15.8

.26

.45

1. 7:1

8.4

.10

.53

5.3:1

90-30-0

15.4

.30

.58

1. 9:1

9.2

.13

.68

5.2:1

Ca:P
Ratio

�-37-

Table 11. Chemical analysis of Kobresia be11ardi collected on the Alpine
study area, 1973-74.

Treatment
N - P - H

Protein

Collection Date and Analx:sis (Percent)
SeEtember
Januarx:
Ca:P
Pho s ,
Calcium
Ratio
Protein Phos. Calcium

0-0-0

14.1

.15

.53

3.5:1

5.3

.06

.65

10.8:1

0-30-0

14.4

.22

.53

2.4:1

5.3

.06

.75

12.5:1

30-0-0

15.3

.24

.58

2.4:1

6.7

.10

.70

7.0:1

30-30-0

14.2

.21

.54

2.6:1

7.4

.10

.71

7.1:1

60-0-0

14.6

.13

.53

4.1:1

9.3

.08

.71

8.9:1

60-30-0

17.2

.24

.46

1. 9:1

5.9

.07

.70

10.0:1

90-0-0

17.0

.28

.61

2.2:1

6.1

.06

.75

12.5:1

90-30-0

16.2

.28

.55

2.0:1

8.6

.10

.78

7.8:1

0-0-2

16.2

.16

.51

3.2:1

7.2

.06

.65

10.8:1

0-30-2

14.8

.23

.46

2.0:1

6.8

.06

.46

7.7:1

30-0-2

16.2

.20

.50

2.5:1

6.3

.08

.66

8.3:1

30-30-0

14.4

.24

.46

1. 9:1

5.7

.10

.63

6.3:1

60-0-2

15.8

.18

.50

2.8:1

6.4

.08

.66

8.3:1

60-30-2

12.6

.24

.46

1. 9:1

7.5

.06

."13

12.6:1

90-0-2

15.6

.19

.58

3.1:1

6.7

.05

.75

15.0:1

90-30-2

15.2

.25

.46

1. 8:1

8.2

.10

.65

6.5:1

Ca:P
Ratio

�-38-

Table 12. Chemical analysis of Poa sp. collected on the. Alpine Study
Area, 1973-74.
Collection Date and Ana1~sis ~Percent)
SeEtember
Januar~
Ca:P
Ratio
Phos. Calcium
Protein Phos. Calcium

Treatment
N - P - H

Protein

0-0-0

10.2

.15

.38

2.5:1

3.0

.03

.61

20.3:1

0-30-0

12.8

.26

.35

1. 3:1

4.4

.07

.41

5.9:1

30-0-0

10.6

.15

.30

2.0:1

4.5

.04

.47

11.8:1

30-30-0

11.6

.22

.28

1. 3:1

4.7

60-0-0

12.4

.15

.38

2.5:1

5.1

.03

.53

17.7:1

60-30-0

19.4

.29

.40

1.4:1

5.5

.06

.53

8.8:1

90-0-0

12.0

.20

.31

1.6:1

5.2

.03

.58

19.3:1

90-30-0

11.0

.25

.28

1.1:1

4.6

.04

.51

12.8:1

0-0-2

12.4

.15

.38

2.5:1

4.7

.03

.63

21.0:1

0-30-2

15.0

.25

.46

1.8:1

5.6

.05

.61

12.2:1

30-0-2

13.4

.14

.36

2.6:1

5.2

.05

.65

13.0:1

30-30-2

13.0

.23

.29

1. 3:1

5.0

.04

.58

14.5:1

60-0-2

13.4

.16

.38

2.4:1

6.3

.06

.58

9.7:1

60-30-2

17.4

.29

.38

1.3:1

6.6

.03

.53

17.7:1

90-0-2

17.6

.19

.34

1.8:1

5.8

.05

.55

11.0:1

90-30-2

14.4

.26

.26

1.0:1

Ca:P
Ratio

.46

�-39-

Table 13. Chemical analysis of Trifolium
Study Area, September, 1973.

nanum collected

on the Alpine

Protein

Ana1xsis ~Percent)
Phosphorus

Calcium

Ca:P
Ratio

0-0-0

20.0

.23

2.20

9.6:1

0-30-0

19.8

.28

2.32

8.3:1

30-0-0

19.2

.24

2.34

9.8:1

30-30-0

19.4

.25

2.20

8.8:1

60-0-0

18.6

.23

2.25

9.8:1

60-30-0

19.2

.30

2.30

7.7:1

90-0-0

18.8

.26

2.30

8.8:1

90-30-0

19.6

.27

2.35

8.7:1

Table 14. Chemical analysis of Ore ox is bakeri collected
Study Area, September, 1973.

on the Alpine

Treatment
N - P - H

Protein

Ana1xsis ~Percent~
Phosphorus

Calcium

Ca:p
Ratio

0-0-0

16.2

.24

1.05

4.4:1

0-30-0

15.8

.31

1.13

3.6:1

30-0-0

14.4

.25

1.05

4.2:1

30-30-0

15.0

.30

1.08

3.6:1

60-0-0

15.4

.26

1.38

5.3:1

60-30-0

15.4

.34

.90

2.6:1

90-0-0

16.0

.25

1.05

4.2:1

90-30-0

14.4

.27

1.00

3.7:1

Treatment
N -

H - H

�-40-

Table 15. Chemical analysis ofGeum
Study Area, September~ 1973.

ttirbirtatumcollected on the Alpine

Protein

Analxsis (Percent~
Phosphorus

Calcium

Ca:P
Ratio

0-0-0

13.2

.21

.80

3.8:1

0-30-0

12.8

.22

.90

4.1:1

30-0-0

11.0

.19

1.05

5.5:1

30-30-0

12.6

.23

1.05

4.6:1

60-0-0

11.6

.19

.88

4.6:1

60-30-0

17.0

.28

.83

3.0:1

90-0-0

13.4

.21

.99

4.7:1

90-30-0

12.6

.19

.88

4.6:1

Treatment
N - P - H

Table 16. Number of bighorn sheep, deer, and elk observed grazing on the
Cebolla Creek Study Area during the period January - May, 1974.

Treatment
N -

P - H

0-0-0

Ewes

Bighorn Sheep
Lambs
Rams

Total

Elk
No.

Deer
No.

7

1

0-30-0
30-0-0
30-30-0
60-0-0

13

60-30-0

6

90-0-0
90-30-0
0-0-2
0-30-2

1
2

30-0-2
30-30-2

5

60-0-2
60-30-2

17

90-0-2

3

90-30-2

3

�-41January
JO~ PROGRESS

Sta.te of
Project

REPORT

CO:::.=.:LO:::::.::I&lt;AD=:..::O~----

Job Title

Job No.

1

Trapping,

Covered:

P.ersonnel:

Bighorn Sheep &amp; Mountain

W-41-R-24

No.

Work Plan No.

Period

1975

Marking,

and Collecting

Goat Investigations

~2~1~

_

Bighorn Lambs

June 1, 1973 to April 30, 1974.

R. L. Schmidt, J. Rodgers,

J. Morris,

P. Neil, and T. Spraker.

ABSTRACT
Eight bighorn lambs were collected for necropsy and histopathological
studies
of lungworm infection.
Results of the necropsy ani histopathological
investigations are included under the Work Plan 1, Job CONTRACT Progress Report submitted by Colorado State University.

��-43-

TRAPPING,

MARKING, AND COLLECTING

BIGHORN SHEEP

Robert L. Schmidt

P. S. OBJECTIVE
To trap or collect bighorn sheep for adding to the captive flock at Little
Hills Experiment Station, for obtaining nasal swabs and blood samples, for
marking and treatment with therapeutic drugs, or (in the case of a dead
animal collection) for necropsy examination.
SEGMENT OBJECTIVES
1.

Collect bighorn lambs at intervals and locations prescribed
agreement.

2.

Trap an indefinite number (as many as can reasonably
handled) of bighorn sheep of all age classes.

by contractual

be caught and! or

METHODS AND MATERIALS
From June 1 to August 15, 1973, 7 bighorn sheep lambs were collected by
shooting from Pikes Peak ewes. Attempts were made to shoot all lambs in
the neck. to minimize damage to tissues and organs of interest.
Lambs were
collected only from ewes without collars so that data concerning parasitology
and pathology were not biased by drug treatments.
Trapping and marking procedures
Progress Report.

are described

in the Work Plan 1, Job 22

RESULTS AND DISCUSSION
From June 3 to August 14, 1973, 8 bighorn sheep lambs were necropsied from the
Pikes Peak sheep herd. Seven of these lambs were shot and 1 was found approximately 12 hours after it had died. Four of the lambs were males and 4 were
females. Three were collected in June, 4 in July, and 1 in August (Table 1).
All 8 lambs had numerous first stage lungworm larvae (Protostrongylus stilesi)
in their lungs upon examination.
A more detailed summary of lamb necropsy
results are presented in Work Plan 1, Job CONTRACT report.
Summaries of the trapping and tagging results are presented
Job 22 Progress Report.

Prepared by
Robert L. Schmidt
Sr. Conservation Aide

in Work Plan 1,

�Table 1. Dates and locales of collection, sex, ages, and weights of bighorn lambs, collected from Pikes
Peak, June-August 1973.
Lamb
Number

Collection
Date

Collection
Locale

Sex

Weight
(Lbs.)

Estimated
Age (Wks.)

3-1/2 - 4

18

73-BHL·ll

6-03-73

N. side Bison Reservoir

F

73-BHL-12

6-19-73

E. face W. Beaver Creek

M

4 -

5

23

73-BHL-13

6-28-73

E. side W. Beaver Creek

F

6 -

7

26

73-BHL-14

7-12-73

W. face W. Beaver Creek

F

8 -

9

32

73-BHL-15

7-24-73

Pikes Peak Toll Road below
gravel pits

M

8 - 10

34

"

I

73-BHL-16

7-26-73

Pikes Peak Toll Road below
Boulder Park

M

8 - 10

28

73-BHL-17*

7-26-73

E. side Pikes Peak Summit

M

8 - 10

21

73-BHL-19

8-14-73

Cog Rail near Pikes Peak
Swmnit

F

10 - 12

36

* Found dead.

I
.p.p-

�-45-

January 1975

JOB PROGRESS REPORT

State of

..::;CO;:..LO=RAOO=~----Bighorn Sheep
W-41-R-24

Project No.
Work Plan No.

1

Job No.

Job Title

Monitoring

Period Covered:

June 1, 1973 - May 31, 1974

Personnel:

&amp; Mountain Goat Investigations

22
----------------------------

Bighorn Sheep Populations

Robert L. Schmidt, Gene G. Schoonveld, Terry Spraker, and R. Bruce
Gill.

ABSTRACT
Bighorn sheep numbers in the Four Mile area of Pikes Peak. were estimated using
a simple Lincoln index. The Lincoln index estimated the population at 43 bighorn sheep (31 &lt;
60; p&gt;. 95). Ground counts indicated a population of
about 45 animals. Age and sex composition counts during the period Nov. 15,
1973 _ April 15, 1974, indicated 11.5 percent of the Pikes Peak population
were adult rams, 0.9 percent were yearling rams, 82.3 percent were adult ewes,
0.2 percent were yearling ewes and 5.1 percent were lambs.

N.s..

Ratios of lambs:100 adult ewes of bighorn sheep treated witlt chemotherapeutic
drugs were nearly double those of untreated sheep (49 lambs:lOO adult ewes vs·
25 lambs:lOO adult ewes).

�-48-

Lambs
Lambs were distinguished pr:lmarily on the basis of their diminutive size.
At approximately 6 months, lambs are about one-half the size of an adult
female and have the characteristic facial features of a young animal (Fig.
1).

Yearling

Females

Yearling females resemble adult females except they are not as large
(approximately 27 inches at the shoulder at 1 year) and they retain a juvenile
short face. They differ from yearling males in appearance in that yearling
females appear thin and sleek-looking with a long thin neck and slim legs.
Horns of yearling ewes are approximately 5-7 inches long and are thin and
sharp pointed (Fig. 2).

Yearling

Rams

This age class of bighorns is most easily misidentified as an adult ewe.
Yearling rams differ from adult ewes by having shorter noses shaped more
like that of a lamb. The nose appears triangular compared to the long nose
of an adult ewe. Horn bases are slightly wider and longer and horns are more
divergent than those of adult ewes. According to Geist (1971) yearling rams
differ from adult ewes behaviorially by using the ram urinating posture,
courting females, and lip curling ov~r female urine (Fig. 3).

Adult Ewes
Adult ewes measure approximately 34 inches at the shoulder. Horns are nearly
6-7 inches in length and appear smaller'and are less divergent than yearling
ram horns.
The nose appears long and slim. Adult ewes range in weight from
120-200 pounds. This is the most frequently observed class of mountain sheep
(Fig. 2).,

Adult Rams
Adult rams are noticeably larger th&amp;~ adult ewes or yearling rams. Horns
appear larger and more massive than any other class of sheep. Body conformation has a blocky appearance.
Horn configuration varies between 1/2 to full
curl or better in exceptional rams. Testes are obvious. The body color varies
with age, but generally adult rams are darker and have less white on the belly
and rear margins of the front legs than adult ewes (Fig. 3).
Since these sex and age criteria are entirely subjective, their reliability is
unknown in the absence of objective tests on populations of known sex and age
composition.
Smith (1954) questioned the reliability of distinguishing yearlings (particularly yearling rams) from adult ewes, especially during the
spring-summer months (March - August).
Because these classification criteria
are subjective the data herein sUDllitarized
may just as readily reflect observer
biases as real population composition.

�-49-

Lamb - 6 Weeks

r3JY

r31lI

Age in 8-16

6-6

years

(Geist 1971)

r3I
2,5

0r.egrling
(ffy.J
1,5

~

~ yearling

ts»)
1.5

Lamb.
0,5

(from Geist 1971)

Fig. 1. Comparisons of body sizes, horn and face characteristics
the various sex and age classes of bighorn sheep.

of

�-50-

Adult Ewe (Moser 1962)

3 Ewes and 3 Yearling Rams
(Geist 1971)

Fig. 2.
rams.

19 Mo. Yearling Ewe (Geist 1971)

Adult Ewe and 4 Young Rams
(Geist 1971)

Comparisons of adult ewes, yearling ewes, yearling rams and young

�-51-

Adult Ram (Geist 1971)

22 Mo. Yearling Ram (Geist 1971)

Ie Mo. Yearling
(Smith 1954)

Fig. 3.

Comparisons

of adult rams and yearling rams.

Ram

�-52-

In addition to potential observer bias, there is a sampling deficiency.
Data were gathered by traveling in a vehicle, on horseback, or afoot.
No serious attempt was made to sample different sex and age categories
by stratifying the sampling effort according to known or suspected
differences in distribution of the various sex and age classes.
Smith
(1954) reported mature rams were --- "consistently less 'observable' than
females and juveniles"---except
possibly during the fall rutting season
and early winter period when rams closely associated with groups of ewes
and young sheep.
Another potential bias exists in early to mid-summer (June and July) lamb:
adult ewe ratios.
During this period lambs are occasionally seen with a
single ewe while actual mothers of these lambs may be as much as 1 mile
distant from their lambs. This tendency for adult ewes to "baby-sit" lambs
of other adult ewes could result in unrealistically high or low ratios of
lambs:adult ewes depending upon the frequency with which one encountered
the ewes-only groups or the "baby-sitting" groups.
All of the aforementioned
potential biases should be carefully considered befora conducting bighorn
sheep classification surveys.
Trapping

~id Drug Treatment

Bighorn sheep were trapped on four areas on Pikes Peak: the Beaver site,
the Tunnel site, the Cog Rail site, and the 4 Mile site (Fig. 4). These
sheep were trapped with a modified version of the drop-net trap described
by Erickson (1970) us.lng apple pulp mash as a bait. Captured bighorns were
marked with colored neckbands lrith individually identifi.able symbols and
numbers (Table 1). Each bighorn captured was also treated with a single
injection of a candidate chemotherapeutic
drug for lungworm control.
Five
drugs were used (thiabendazole, dichlorvious, cambendazole, tramisol, and
diethylcarbamizine),
the color of the collar denoting the particular drug
tested (Table 1). Ewe:lamb ratios of, treated and untreated groups of sheep
were used to compare the effectiveness of drug treatments.
RESULTS AND DISCUSSION
Bi.ghorn Sheep Numbers
Estimating numbers of bighorn sheep populations is difficult under the most
favorable circumstances.
On Pikes Peak the availability and observability of
marked bighorn sheep provided an opportunity to experiment with capture-recapture techniques for estimating population numbers.
An example of the
potential of this approach was afforded from data on the Four Mile bighorn
sheep herd on Pikes Peak. This bighorn herd remained fairly discrete from
other Pikes Peak bi.ghorn herds during the 1973-74 winter.
Repeated counts
placed the number of sheep m this group at approximately 45 animals.
On
April 4, 1973, 15 bighorn sheep were trapped and fitted with individually
recognizable collars.
During the period December 12, 1973 to April 11, 1974,
113 observations of bighorn sheep were recorded, of which 39 were observations
of collared sheep.

�•• I
•

I

t'

",

I

••

I

I

•

I

••

I

••

·.W;"
•• ~ 1 ~ 11-(
.... 1_'
_ .~~~I
.. _ 1 I

0/

1

"

JJi

r

n ....

I

• • ••

••

I

!I

I

10 \

..

!'.'

/l

~:"'&lt;X&gt;3'

I

/,.&lt;j ffit;'
.

1 ·

~

;;/

, J!~
·1":""
1.. '

•
.,

/VA

---

,

I!~

I

)

, .,.-~~r.."/

~

I
V1
1,.0.)

I

-

••

•

Fig. 4. Bighorn sheep trapping areas, Pikes Peak 1973-74, (1) Beaver site, (2) Cog Rail, (3) Tunnel
site, and (4) 4-Hile.

�-54-

Table 1. Bighorn sheep trapped, marked and treated on Pikes Peak from
February - April 1974.

Date

Collar and
Number

Age

Location of
Capture

Treatment

Bighorn Rams Captured on Pikes Peak
2-5
2-15
2-21
2-21
2-21
3-20
4-4
4-9

Lamb
5 Yrs.*
1 Yr.*
6 Yrs.
6 Yrs.
1 Yr.
1 Yr.
5 Yr.

Eartag 084
White.
1
Eartag 090
White.
7
W!.lite. 8
Eartag 089
Pink BS
White.
1

Beaver
Tunnel
Tunnel
Tunnel
Tunnel
Cog Rail
Cog Rail
Cog Rail

Camb 115 cc only
Camb 270 cc only
Triben 60 cc only
Triben 120 cc only
Triben 120 cc only
Rec. Lost 090, New 089
Camb 70 cc
Rec. - Tram 10 cc

Bighorn Ewes Captured on Pikes Peak with Blue Collars
2-21
2-21
2-21
2-21
3-12
3-12
4-10
4-12
4-12
4-12

6 or 7*
6 or 7
5 or 6*
5 or 6***
6 or 7
6 or 7
6 or 7
5 or 6
6 or 7
6 or 7

Blue. 1
Blue. 2
Blue
3
Blue. 4
Blue.ll
Blue.12
Blue. 5
Blue. 6
BJ_uee 7
Blue. 8

Tunnel
Tunnel
Tunnel
Tunnel
Cog Rail
Cog Rail
4 Mile
4 Mile
4 Mile
4 Mile

Triben
Triben
Triben
Triben
Triben
Trioen
Triben
Triben
Triben
Triben

120 cc - Diethy 3. 75 mg
120 cc - Diethy 3. 75 mg
120 cc - Diethy 3.75 mg
120 cc - Diethy 3.75 mg
120 cc - Diethy 3. 75 mg
120 cc - Diethy 3. 75 mg
120 cc - Diethy 3. 75 mg
120 cc - Diethy 3. 75 mg
120 cc _ Diethy 3. 75 mg
120 cc _ Diethy 3. 75 mg

Bighorn Ewes Captured on Pikes Peak with Yellow Collars
2-5
2-5
2-15
2-15
2-15
2-15
3-1
3-1
3-1
3-20
3-20
3-20
3-20
3-20
3-20
3-25
3-25
3-25
3-25
3-25

2 Yrs.
6 Yrs.
6 or 7 Yrs.
6 or 7
6 or 7
6 or 7
6 or 7**
6 or 7
5 or 6*
5 or 6
6 or 7
6 or 7*
6 or 7
6 or 7
5 or 6
6 or 7
5 or 6
6 or 7
6 or 7
6 or 7

Yellow.
1
Yellow
2
Yellow
3
Yellow.
4
Yellow
5
Yellow.
6
Yellow.
7
Yellow.
8
Yellow.
9
YellowO 11
Ye 11owe 12
Yellow
13
Yellow
14
Yellow. 15
Yellow. 16
Yellowe17
Yellow. 18
Yellow. 19
Yellow. 20
Yellow. 21

Beaver
Beaver
Tunnel
Tunnel
Tunnel
Tunnel
Tunnel
Tunnel
Tunnel
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
4 Mile
4 Mile
4 Mile
4 Mile
4 Mile

Task
Task
Task
Task
Task
TaGk
Task
Task
Task
Task
Task
Task
Task
Task
Task
Task
Task

120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
120 mg
Task 120 mg
Task 120 mg
Task 120 mg

�-55-

Table 1. Bighorn sheep trapped, marked and treated on Pikes Peak from
February - April 1974 (continued).

Date

Collar and
Number

Age

Location of
Capture

Treatment

Bighorn Ewes Captured on Pikes Peak with Red or Orange Collars
2-5
2-15
2-15
3-15
3-15
3-25

Lamb
6 or 7 Yrs.
6 or 7*
5 or 6
6 or 7
4

4-4

8

4-9
4-9
4-9
4-9
4-10
4-10
4-10

6 or 7

5 or 6
6 or 7
6 or 7
5 or 6
6 or 7
5

Eartag (RE) 081
Orange. 4
Orange. 5
Orange. 2
Orange. 6
Blue 6 with
Yellow Patch****
Red 9 with
Yellow Patch****
Red. 1
Red. 6
Red. 3
Red. 5
Red. 7
Red. 8
Blue 1 with
Yellow Patch****

Beaver
Tunnel
Tunnel
Cog Rail
Cog Rail
4 Mile

Camb8J
cc - Diethy
Camb 270 cc - Diethy
Camb 270 cc - Diethy
Camb 270 cc - Diethy
Camb 270 cc - Diethy
Camb 270 cc - Diethy

Cog Rail

Camb 270 cc - Diethy 3.75 mg

Cog Rail
Cog Rail
Cog Rail
Cog Rail
4 Mile
4 Mile
4 Mile

Camb
Camb
Camb
Camb
Camb
Camb
Camb

270 cc - Diethy
270 cc - Diethy
270 cc - Diethy
270 cc - Diethy
270 cc - Diethy
270 cc - Diethy
270 cc - Diethy

3.75 mg
3.75 mg
3.75 mg
3.75 mg
3.75 mg
3.75 mg

3.75 mg
3.75 mg
3.75 mg
3.75 mg
3.75 mg
3.75 mg
3.75 mg

Bighorn Ewes Captured on Pikes Peak with White Collars
2-21
2-21
2-21
2-21
2-21
3-25

6 or 7*
5 or 6*
6 or 7*
6 or 7*
6 or 7*
6*

3-25

6*

4-4

7

4-12

5

White. 2
White. 3
White. l+
White. 5
White. 6
White 16 with
Yellow Patch****
White 17 with
Yellow Patch****
White lR8 with
Yellow Patch****
White 15 with
Yellow Patch

Tunnel
Tunnel
Tunnel
Tunnel
Tunnel
4 Mile

Tram
Tram
Tram
Tram
Tram
Tram

4 Mile

Tram 10 cc - Diethy 3.75 mg

Cog Rail

Tram 10 cc - Diethy 3.75 mg

4 Mile

Tram 10 cc .-Diethy 3.75 mg

10 cc - Diethy
10 cc - Diethy
10 cc - Diethy
10 cc - Diethy
10 cc - Diethy
10 cc - Diethy

3.75 mg
3.75 mg
3.75 mg
3.75 mg
3.75 mg
3.75 mg

Other Bighorn Sheep Captured on Pikes Peak (Ewes)
2-5
2-5
2-5
2-5
2-5

2-5
3-1
3-1

6 or
6 or
5 or
5 or
6 or
6 or
8

7 Yrs.
7
6
6
7
7

5 or 6

White/Red
Blue. 4

7

Beaver
Beaver
Beaver
Beaver
Beaver
Beaver
Tunnel
Tunnel

To Fort Collins
To Fort Co l.I ins
To Fort Call ins
To Fort Collins
Net Loss
Net Loss
Net Loss
Rec. - No Treatment

----------------------------_._-----------------------------------------------------

�-56-

Table 1. Bighorn sheep trapped, marked and treated on Pikes Peak from
February - April 1974 (continued).

Date

Collar and
Number

Age

Location of
Capture

Treatment

Other Bighorn Sheep Captured on Pikes Peak (Ewes) (Continued)
3-12
3-12
3-12
3-12
3-12
3-20
3-20
3-20
4-4
4-4
4-4
4-9
4-9
4-9
4-9
4-9
4-9
4-9
4-12

5 or 6
6 or 7
6 or 7
5 or 6
6 or 7
6 or 7
6 or 7
6 or 7
6 or 7
5 or 6
5 or 6
6 or 7
6 or 7
5 or 6
6 or 7
6 or 7
6 or 7
5 or 6
6*

4-12

6*

Blue.4
White.

6

3lue • 1
Yellow.
7

Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
Cog Rail
4 Mile

Yellow.13
Yellow. 9
Blue e· 3
Wllite e 4
White.5
White. 3
White
2
Yellow.
7
Orange. 5
Blue. 4
White 16 with
Yellow Patch****
White 17 with
4 Mile
Yellow Patch****

e

Rec. - Triben 120 cc
Rec. - Tram 10 cc
To Fort Collins
To Fort Collins
To Fort Collins
Rec. - Triben 70 cc
Rec. - No Treatment
Net Loss
Rec. - No Treatment
Rec. - No Treatment
Rec. - Triben 120 cc
Rec. - Tram 10 cc
Rec. - Tram 10 cc
Rec. - Tram 10 cc
Rec. - Tram 10 cc
Rec. - No Treatment
Rec. - Camb 270 cc
Rec. - Triben 70 cc
Rec. - No Treatment
Rec. - No Treatment

Other Bighorn Sheep Captured
45 bighorns were captured in South Dakota. 24 rams and 2 ewes were brought to
Fort Col11lls. 6 yearlings were sent to University of South Dakota. 13 sheep
were marked and released in South Dakota.
Other Bighorn Sheep Treated
Feb. 1, 1973
18 sheep were treated on Rampart Range.
Mar. 1, 1973
15 sheep were treated on Rampart Range.
Apr. 3, 1973
12 sheep were treated on Rampart Range.
All of the Rampart Range sheep were treated with 270 cc Camb in apple pulp.

*Recaptured ~
**

Recaptured

in 1974.
twice in 1974.

***

Recaptured three times in 1974.

****
Caught in 1973 and again in 1974.

Triben
Task
Camb
Tram
Die thy

Thiabendazole
Dichlorvious
Cambendazo1e
Tramiso1
Diethy1carbamzine

�-57-

A

A simple Lincoln index was constructed so that N =

A

nM where: N is an
x
estimate of the size of the bighorn sheep population; n is the total number
of bighorn sheep observations during the sample period; M is the total number
of collared sheep in the population; and x is the number of collared bighorn
sheep observations
"'"
. during the sample period (Overton 1971). In this particular case: N = 113 x 15 = 43 bighorn sheep in the Four Mile population.
39
Confidence limits of this estimate are defined by the furmula:

'"N

A

N
L' U

= nM (x+2) ±\J
x

A

x+l

A

N ,N
L U

Again in this particular case:

2

113 x 15 (4l)

±'.J 40

1,695 (.0353; .0186) or

392
P = .95 that 3l~ N~60.
This simple model serves only to illustrate the
potential utility of a capture-recapture index for estimating bighorn sheep
numbers. This potential should be explored further.

Age and Sex Composition
Based upon a sample of 516 observations of bighorn sheep during the period
November 15, 1973 to April 15, 1974 the age and sex composition of bighorn
sheep on Pikes Peak was estimated to be: 11.5 percent adult rams, 0.9 percent yearling rams, 82.3 percent adult ewes, 0.2 percent yearling ewes, and
5.1 percent lambs. The ratio of 1ambs:lOO adult ewes was only 6:100 and the
ratio of yearlings:lOO adult ewes was only 1:100. Even considering the
potential biases mentioned before, the lamb and yearling survival on Pikes
Peak is extremely low. With ratios of this magnitude severe population declines of bighorn sheep on Pikes Peak are inevitable.
The ratios of adult rams:lOO adult ewes declined on Pikes Peak from 82:100 in
1972-73 to 14:100 in 1973-74. Observations of collared rams indicated only
20 percent of those rams present in 1972-73 were accounted for in 1973-74.
It was hypothesized that most of this decline was attributable to migrations
of adult rams from the Pikes Peak area.

Trapping and Drug Treatment
During the period February 5 - April 12, 1974, 99 bighorn sheep were captured,
marked, treated with drugs, and released on Pikes Peak (Table 1). Eighteen
of these sheep were recaught from 1 to 3 times during this period and 8 sheep
were originally trapped in 1973 and recaught in 1974. In addition, 45 bighorns were captured in South Dakota, 26 of which were transported to Fort
Collins for experimental lungworm control research, 6 were shipped to the
University of South Dakota, and 13 were marked and released at the capture
site in South Dakota. Approximately 45 sheep were treated with cambendazole
in the Rampart Range by mixing cambendazole in apple pulp mash and offering
it ad libitum. Each sheep received an estimated 270 cc of the drug.

�-58-

The effects of drug treatment upon lamb survival of wild bighorn sheep
is still in the process of being analyzed. But classific.ation data from
the period November 19, 1973 to March 15, 1974 revealed a ratio of 45
lambs:lOO adult ewes (n = 32 observations) for collared ewes compared to
5 lambs:lOO adult ewes (n = 240 observations) for unco.lLarad ewes. This
could indicate b~tter lamb production and/or survival from treated vs. untreated ewes.

LITERATURE CITED
Bell, G. 1974. Population estimates from recapture studies in which no
recaptures have been made. Nature 248:616.
Darroch, J. M. 1959. The mUltiple recapture census. II. Estimation
where there is immigration or death. Biometrika 46:336-351.
Geist, V. 1971. Mountain sheep, a study in behavior and evolution.
The Univ. of Chicago Press. 383p.
Hansen, C. G. 1965. Grcwth and development
J. Wild1. Manage. 29(2): 387-391.

of desert bighorn sheep.

Jolly, G. M. 1963. Estimates of popUlation parameters from multiple
recapture data with both death and dilution - deterministic model.
Biometrika 50:113-128.
1965. Explicit estimates from capture-recapture data with both
death and immigration - stochastic model. B'Lome trr Lka 52 :225-247.
Lincoln, F. C. 1930.
banding returns.

Calculating waterfowl abundance on the basis of
U.S.D.A. Circular No. 118. 4p.

Manly, B.F. J. 1970. A simulation study of animal population estimation
using the capture-recapture method. J. Appl. Eco1. 7(1):13-19.
" and M. J. Parr. 1968. A new method of estimating popul.atLon size,
---survivorship,
and birth rate from capture-recaptur.e data. Trans.
Soc. Brit. Ent. 18:81-89.
Moser, C. A. 1962. The bighorn sheep of Colorado.
and Fish. Tech. Pub1. 10. 49p.

Colo. Dept. Game

Overton, W. S. 1971. Estimating the numbers of animals in wildlife populations. p , 403·-456. In Giles, R. H., Jr. (Ed.). Wildlife management
techniques.
The Wildlife Society, Washington, D.C. 633p.
Parr, M. J., T. J. Gaskell, and B. J. George. 1968. Capture-recapture
methods of estimating animal numbers. J. BioI. Educat. 2:95-117.
Smith, D. R. 1954.
and management.

Prepared by

The bighorn sheep in Idaho, its status, life history,
Idaho Dep. Fish and Game, Boise. l54p.

�-59-

January 1975

JOB PROGRESS REPORT

State 0 f

---.:C:;:O:,:LO::::..::;RAD:::::::.:::.O
_

Project No.

W-4l-R-24

Work Plan No.

1

Job Title

Bighorn Sheep &amp; Mountain Goat Investigations
Job Noo

C_ONT_RA_C_T

S_p_o_n_t._an
__e_o_u_s
__D_l_·s_e_a
__s_e_s
__o_f
__B_i_g_h_o_rn
S_h_e_e_p

_
_

Period Covered: July 1, 1973 - June 30, 1974
Personnel:

Colorado Division of Wildlife - Gene G. Schoonveld, Robert L.
Schmidt, and Robert E. Keiss.
Colorado State University - C. P. Hibler, T. R. Spraker, L. H.
Lauerman, R. E. Lange, C. J. Metzger, F. ~. Latson, L. R. Clum,
and J. G. Wegryzn.
ABSTRACT

The primary objectives of the research this past year were: (1) Treat sheep
with antihelminthics in an effort to find a suitable compound; (2) Obtain
more information on transplacental transmission, especially in'regards to
the strong possibility that lambs were tolerant to the larva with which
they were born; (3) Locate the "hot spots" for lungworm transmission on Pikes
Peak and describe these areas; (4) Obtain additional information on the cause
and nature of lamb mortality by postmortem examination of lambs; and (5)
Establish a captive herd of sheep for experimental infection and chemotherapy
studies.
Cambendazole, when used on ewes in early pregnancy seemed to have little or
no efficacy against the adult parasite,- but did depress fecal output of first
stage larvae for periods of about 6 weeks; moreover, lambs born of ewes treated
with this drug during early pregnancy died of a typical verminous pneumonia.
However, when ewes in late pregnancy were treated. the. lambs were born healthy
and survived the winter in good shape. Tramisol, the only other compound
evaluated. during 1972-73, did not show any efficacy (all drug evaluation
reports are one year after they have been evaluated),
Lamb collections in 1973 revealed that there is indeed an immune tolerance
to the larva with which they are born. The ewe pas-ses the infective larva
she has stored to her fetus in the third trimester of pregnancy,
These
larvae are stored in the liver until birth, at which time they go to the lungs
and begin to develop, Development from 3rd to 4th and finally to the 5th stage
(adult) requires about three weeks. There is no host Tesponse to these larvae
when they are developing, which means they are- tolerant and do not recognize
the larva as a foreign body; rather, they recognize it as "self", On the
third to fourth week of life, the worms are mature and start laying eggs.
At this time there is a very severe host response to the eggs. This means
the lamb is immunologic~lly competent and can recognize foreign protein, it

�-60-

simply does not recognize the worm as such. Once eggs are deposited however, this represents a new, definitely foreign, protein and it responds.
The type of response is a "delayed-type hypersensitivity" and is so severe
it almost overwhelms the lamb. By the time the eggs develop, and hatch
into larvae, the lamb is thoroughly sensitized to these large amounts of
antigen. As the Larvae pass up the bronchial t ree , many are accidentally
aspirated into the ventral lobes of the lungs. These larvae, in the
sensitized lambs, initiate a granulomatous response. This then creates
an ideal environment for Pasteurella and/or Mycoplasma to become entrenched
and kill the lamb. Thus, transplacental transmission of lungworm predisposed the lamb~ to a fatal pneumonia by other agents.
The "hot spots" for lungworm transmission on Pikes Peak are the winter bedding
grounds. Ewes spend all winter on these bed grounds, leaving for lambing
grounds in the spring.. From there they go to smmner range and do not return
to the winter bedding grounds until August. Lungworm larvae enter snails in
the spring and are ready to infect ewes when they return in August.
Most of the information regarding the postmortem examination of lambs is in
tabular form immediately following the Abstract (Table 1).
A captive herd of bighorn sheep obtained in South Dakota, and a herd of
mouflon/bighorn hybrids obtained from Pennsylvania are currently being infected with lungworm to do immunology and chemotherapy studies.

�Table 1.

Summary of lamb examinations

Sheep

1973.

Wt.

Date
Examined

Sex

Age

(II)

Pneumonia

Unknown
Isolates
Mycoplasma
Virus

Nasal

Pasteurella
Bronchi
Trachea

Lungs

Lungworm
(1st Stage
Larvae)

+

682.,331

+

-

-

1,286,900

+

+

+

+

5,774,710

+ (

+

-

-

3,947 &amp;
many 3rd &amp; 4th

-

+

-

-

507,020

-

-

-

-

-

259,490

-

73BHL-5

(H)

10-2-73

M

21 Wk.

46

Yes

-

+

+

+

73BHL-7

(H)

8-25-73

F

15 Wk.

40

Yes

+*

+

·73BHL-8 (H)

8-1-73

M

10 Wk.

18

Yes

+

73BHL-ll

6-5-73

F

3~-4
Wk.

18

No

+

-

73BHL-12

6-19-73

M

4-5 Wk.

23

No

+

73BHL-13

6-28-73

F

6-7 Wk.

26

No.

-

73BHL-14

7-12-73

F

8-9 Wk.

32

Yes

+

+

+

+

-

73BHL-15

7-24-73

M

8-10 Wk. 34

Yes

+

+

-

+

73BHL-16

7-26-73

M

8-10 Wk. 28

Yes

+

+

+

+

-

+

470,625
898,650

2,279,800

(t)

7-26-73

M

8-10 Wk. 21

Yes

+*

+

-

-

-

-

73BHL-18

(H)

8-10-73

M

6 Days

5

No

-

+

+

+

+

+

73BHL-19

8-14-73

F

10-12
Wk.

36

Yes

+

-

+

-

+

+

960,412

George

5-20-72

M

1 Yr.- 110
6 Mos.

Yes

-

+

+

+

+

+

Very few

H

arginini.

- Born in captivity

from ewes captured

on Pikes Peak.

t

- Found dead on Pikes Peak and was dead too long for thorough

postmortem

examination.

t-'
I

75,640

73BHL-17

* -~.

I

0\

M:my 3rd &amp; 4th

��-63-

SPONTANEOUS DISEASES OF BIGHORN SHEEP
C. P. Hibler
P. S. OBJECTIVE
To determine causes, nature and effects of spontaneous diseases in Colorado
bighorn sheep with particular reference to mortality in lambs, and study
various means of controlling these diseases.

SEGMENT OBJECTIVES
1.

Complete tissue examination of bighorn lambs col:ected during 1972.

2.

Collect a minimum of 6, and a maximum of 9 lambs from Pikes Peak earlier
in the summer for a detailed postmortem examination in order to examine
specific neo-natal pathological processes suspected to play an integral
part in the inception of chronic lamb pneumonias.

PROCEDURES
1.

Complete tissue examination of bighorn lambs collected in 1972.

2.

All lamb collections tae last two years have been in late summer and
early fall. Histopathology, virology, bacteriology and parasitology
studies conducted on these animals strongly suggest that earlier
collections would yield information which woule help delineate the
initiation and progression of the pathologic process resulting in
almost total loss of lamb crops. As in 1972, a field research team
will conduct the postmortem examination.
Blood samples will be collected
for clinical pathology studies, serum for bacteriological, virological,
clinical pathology, physiological baseline data, and trace mineral evaluation. Selected tissues will be taken for bacterial and viral isolation,
and trace mineral evaluation. A complete postmortem for histopathology
studies will be performed. The remainder of the carcass (liver, lungs,
muscles and lymph nodes) will be further examined for additional evidence
of parasitism by Protostrongylus.

The proposed collection schedule is:
Lamb
Lamb
Lamb
Lamb
Lamb
Lamb

111
1st week in June, a 1 or 2 day old lamb.
112
2nd week in June, a l-l!-:!
weoSk old lamb.
113
3rd week in June, a 2Yz-3 week old lamb.
114
1st week in July, a 4-5 week old lamb.
115
3rd week in July, a 6-8 week old lamb.
116 - 1st or 2nd week of August, an 8-10 week old lamb.

Additional lambs may not be necessary, but in the event that a significant
observation is made on lambs collected before mid-July,.an additional 1 to 3
lambs should be collected at the discretion of the rese~rch team.

�-64-

RESULTS
The results are given in case history form for each lamb collected and
examined. A tabular summary (Table 1) precedes the detailed results.
In addition, a more complete written summary follows each case history.
During the summer ·of 1973 seven bighorn lambs were collected by Robert
Schmidt, Colorado Division of Wildlife. These lambs were collected at
two week intervals starting the 1st week of June through the second week
in August. The only organ which had lesions was the respiratory system.
A detailed gross and histological examination was done on all lambs.
Also on July 26, 1973, a small ram lamb was found dead and subjected to
the same necropsy as a collected lamb. The results are presented separately for each sheep.

73BHL-ll

June 5, 1973

History: The age of this lamb was estimated to be about 3 weeks. It was
collected just above the cabins on the north side of Bison Reservoir. The
lamb was shot in the nasal turbinates and has aspirated blood into her lungs.
Postmortem:
PMI - 20 minutes
Sex - Female
Body Temp. at Death - 104.4 degrees F
Weight - 18 pounds
Body Condition:

The hair coat is smooth and has a dark tan color.

Cardiovascular System: The heart is normal in size, shape, color and texture.
Vessels are unremarkable.
Lymphohemopoietic System: The mediastinal lymph nodes are slightly enlarged,
especially on the posterior aspect of the chain; they are 1 cm in diameter
from the posterior aspect. The retropharyngeal lymph node is normal size
(~ cm diameter). The hilar lymph nodes are slightly enlarged. The prescapu Lar lymph nodes are both enlarged and slightly moist. Mandibular,
popliteal, and parieta lymph nodes are unremarkable.
Spleen is normal in
size, shape, color (6-8 cc x 3 cc x lcc). Thymus has no gross lesions, it
extends from the base of the heart to ~ way up the trachea. Bone marrow is
red and active.
Digestive System: The rumen is the size of the abomasum and filled with ingesta. Blood is present in the rumen because of swallowed blood. The abomasum
is filled with ingesta and milk curd and some hair. The remaining digestive
system is unremarkable.
Formed fecal pellets are in the colon. There are
six ~ to 1 mm white foci just under the capsule of the liver.
Respiratory System: The lungs do not collapse well, but this is due to the
blood aspirated into sma.lL bronchioles thus trapping air. The trachea and
bronchi are filled with a red tinged froth (blood with bubbles). There are
mUltiple bright red foci scattered diffusely over mid through the parenchyma

�-65-

of both lungs (aspirated blood). No gross evidence of 1ungworms are present except for 3-5 small dark spots located on the posterior dorsal aspect
of the diaphragmatic lobes.
Musculoskeletal

System:

Unremarkable.

Urogenital Syst.em;. Kidneys are normal in size, shape and color. A small
amount of fat is located just under the capsule. The remaining urogenital
system is unremarkable.
Endocrine System:
Nervous System:

Unremark.ab1e.
Eyes and brain are unremarkable.

Histological Findings of Importance:
1.

Lungs - There are multiple small cellular infiltrations in the
posterior dorsal aspect of both diaphragmatic lobes. These
cellular infiltrations are nodules composed of mononuclear
inflammatory cells and eosinophi1s; some areas are predominantly
mononuclear cells while in other areas eosi~ophi1s predominate.
There are several areas of thickened alveolar septa due to the
infiltration of the same inflammatory cellular components.
Parasites also occur in the same general areas as the inflammatory
nodules. These parasites are immature 4th and 5th stage
Protostrongylus sti1esi. There is only a very mild to no cellular
response to these parasites. The minimal response seen is due to
an infiltration of mvnonuc1ear inflammatory cells, lymphocytes,
and eosinophi1s.

2.

Liver - There are several areas of hepatic necrosis that has been
replaced by mononuclear cells, few lymphocytes, eosinophils and
fibroblasts. There are probably areas where the 3rd stage
Protostrongy1u~ sti1esi larvae had been migrating during fetal
life.

3.

Mediastinal Lymph Nodes - The mediastinal lymph nodes are hyperplastic and very active.

4.

ThymUS - There are neets of eosinophi1s in the medulla of the thymus
but the significance is unknown.

Summary: This 3-3~ week old lamb has developing L·-4 to L-5 Protostrongylus
sti1esi larvae in the dorsal posterior aspect of the diaphragmatic lobes.

73BHL-12

June 19, 1973

This lamb was seen with seven ewes (one was an orange tag #1 with one lamb)
and one other lamb. These lambs ran up and down on the rocks, played and
rested. No coughing was noticed. The age was estimated to be about 4-5
weeks. This lamb had a slightly darker tan hair coat than the other lamb,
and it was also a slightly rougher hair coat. The location of the collection
was on the east face of West Beaver Creek, just above the old cabins and
beaver dams in the Bou Lde r field. The lamb was shot at 1:45 PM.

�-66-

Postmortem:
PMI - I hr. 35 minutes
Sex - Male
Body Temp. at Death - 104.2 degrees F
Weight - 23 pounds,
Body Condition: The hair coat is slightly rough and darker than most lambs.
Very little fat is present in the subcutis.
Cardiovascular System: The heart is normal in size, shape, texture, and
color. There is a normal amount of fat in the coronary bond.
Lymphohemopoietic System: The spleen is normal in size, shape, color, and
consistency.
There are multiple red foci located along the periphery of the
organ (normal). Bone marrow normal. The thymus is normal in size and extends
from the base of the heart to about ~ way up the trachea. The mediastinal
lymph nodes are enlarged, wet, and have an exaggerated prominence of the
cortex (size - 7 cm x l~ ern x l~ cm). Mesenteric lymph nodes are normal in
size, shape and internal structure.
Digestive System: The rumen and abomasum are filled with ingesta. There is
a small amount of milk curd in the abomasum. Formed feces are present in the
colon. There is one small (~ to I mm) white focus located just under the
capsule of the liver.
Respiratory System: The nasal cavity is normal, the mucosa is unremarkable.
The trachea has no gross lesions, and its mucosa is moist and shiny. The
lungs only partially collapse on opening the thorax. There are many
irregular firm raised grey-white speckled nodules located on the posterior
dorsal aspect of both diaphragmatic lobes. These reactive nodules are
surrounded by a red zone (hyperemia). These are believed to be lungworm
nodules caused by Protostrongylus stilesi.
(See Figure 1). The bronchi
are clean and moist.
Musculoskeletal Sys tenu
The bones and joints are normal.
a sharp 'snap. Joints are clear. and normal.

Bones break with

Urogenital System: Both testicles are located in the scrotum. Both kidneys
have a small amount of subcapsular fat. The right kidney has two small (l~
cm diameter, other 3/4 cm diameter) cystic areas in the cortex.
Endocrine System:

Thyroid, adrenals, pituitary glands are unremarkable.

Nervous System: The brain and eyes are normal.
C-4 and C-5 cervical vertebra.
Histopathological
1.

The lamb had been shot in

Findings of Importance:

Lungs - The grey-white firm nodules are characterized by mature lungworms with production of numerous eggs. The majority of eggs have
not as yet matured to active 1st stage larvae. The cellular reaction
is mainly mononuclear inflammatory cells, composed of lymphocytes
(in nodules and in the septa), plasma cells, a few eosinophils and
macrophages.
There are also many hemorrhagic foci present through
all lobes of-both lungs.

�-67-

L u.f\:\ wOt"lY\
f\O~u. \~S

."

./

(
I
L e -+ -\- Lu.~~
Fig. 1. Right and left lungs showing lungworm nodules believed caused by
protostilesi.

�-68-

2.

Thymus - The thymus has many nests of eosinophils

in the medulla.

3.

Liver - There is one lymphocytic nodule present in the liver
parenchyma.
The capsule is thickened in one area.

4.

Mediastinal Lymph Nodes - The mediastinal
hyperplasti~ lymphocytes.

lymph nodes have

Summary: The lungworms are mature in this lamb and have begun to produce
eggs. The eggs must take 4 to 7 days to develop however, because no
larvae are present.

73BHL-13

June 28, 1973

History:
This lamb was seen with two ewes and a yearling on the east side
of West Beaver. The lamb ~as collected in the southeast facing rocks on
the east side of West Beaver. The lamb was estimated to be 6-7 weeks old
because she would kneel to nurse and the back of the lamb was l~ to 2~
inches above the ventral aspect of the belly of the ewe. The lamb was
collected about 5:45 PM.
Postmortem:
PMI - 1 hour 20 minutes
Sex - Female
Body Temp. - 104.4 degrees F
Weight - 26 pounds
Body Condition:
The hair coat is slightly roughened and has no sheen. There
is a very small amount of fat in the subcutis, especially around peripheral
lymph nodes.
Cardiovascular System: The heart is normal in size, shape, color and texture.
No excessive pericardial fluid is present in the pericardial sac. A normal
amount of fat is present in the coronary band around the heart.
Lymphohemopoietic System: The spleen is normal in size, shape, color and
texture. There are many small red foci located around the periphery of the
organ (normal). The mediastinal lymph nodes are enlarged. The cut surface
is moist, the cortex is very prominent and granular due to many hyperplastic
germinal centers (Le., the cut surface is "nobby" or roughened due to small
rounded nodules, size 6~ cm x l~ cm) , The prescapuJ.ar nodes are also enlarged
(4 cm x 1.5 em x 1 cm). The cortex is prominent and has a moist cut surface.
The mandibular, parotid, mesenteric, popliteal and prefemoral lymph nodes are
unremarkable.
Digestive System: The rumen and abomasum are filled with vegetation.
There
is only a trace of milk curd in the abomasum.
Formed fecal pellets are present in the colon. There are three small
to 1 mm) white foci under the
liver capsule. A small amount of fat is present in the mesenteries, and the
lacteals are filled with chJllle.

r~

�-69-

Respiratory Syste~: The nasal cavity and turbinates are normal in color
and have no exudate on their mucosal surface. The mucosa of the trachea
is also normal. Ihe lungs only partially collapse. White firm, raised,
irregular areas (lungworm nodules) are located on the posterior dorsal
edge of both diaphragmatic lobes (Figure 2). The firm areas bulge on
their cut surfaces. This cut surface is white and finely granular. No"
inflammatory ring occurs around these lungworm nodules compared to 73BHL12. The remaining lung parenchyma is sprinkled with tinged red to grey
foci (size 1 to 2 mm). The bronchioles and bronchi are cleml and only
slightly moist.
Musculoskeletal System: The bones and ribs have a normal snap. Bone
marrow is red and active. No lesions were seen in the musculature.
Urogenital System: A small amount of fat is present under the capsule
of both kidneys. No lesions were present in the kidneys.
Endocrine System: The thyroid, adrenals and pituitary glands appear to
be normal in size, shape, and color.
Nervous System:

The brain and eyes are unremarkable.

Histopathology:
1.

Lungs - The ventral aspects of the apical, cardiac and intermediate
lobes and the anterior ventral aspect of the diaphragmatic lobes
all contain small granulomas consisting of many epithelioid cells,
multinucleated giant, cells, eosinophils and a few nests of
neutrophils. First stage larvae can be found in the majority of
these granulomas. A few of the nodules have not yet developed into
granulomas and they are characterized by edema, eosinophils,
macrophages, few neutrophils,"hemorrhage, but no multinucleated
giant cells. Several vessels in these areas are cuffed with lymphocytes. The lobules grossly observed to be darker than normal
histologically are seen to be areas of pulmonary edema and hemorrhage
with a moderate number of alveolar macrophages and a few neutrophilso
The white firm lungworm nodules characteristically have many mature
Protostrongylus stilesi eggs and 1st stage larvae. Many of the
bronchioles leading away from these nodules are filled with mucous
and 1st stage larvae. The mucosa of these bronchioles is hyperplastic.
The lungworm nodules are characteristically areas of near complete
consolidation of lung parenchyma and consist of adult worms, eggs and
1st stage larvae. There is a severe cellular reaction towards these
parasites, composed of macrophages, lynlphocytes, eosinophils, multinucleated giant cells, and very few neutrophils. Many bronchioles
and blood vessels are cuffed with lymphocytes.

2.

Lymphohemopoietic System - The mediastinal lymph nodes have undergone
reticuloendothelial and lymphocytic hyperplasia. Many germinal centers
are present in the cortex and a moderate number of 1st stage larvae
can easily be found in the node and the afferent lymphatics. Also
several 1st stage larvae are present in the mesenteric lymph nodes.
Most of the eosinophils are located in the medulla of the thymus.

�-70-

(£fP

tJJ

....

e,

L

»:

"""

J"

./

&lt;;

'"

Fig. 2. Right ~ele~t ~ungs ~~i~g~irm,
nodules)

located on the posterior

raised, irregularareas (lungworm

dorsal edge of both diaphragmatic

lobes.

�-71-

Summary: The primary histopathologic findings are again in the lungs.
The parasites located in the posterior dorsal aspect of the diaphragmatic
lobes are now fully mature and producing tremendous numbers of eggs.
First stage larvae are matu~e and are leaving the lungworm nodules via
horizontal bronchi and are being aspirated into the ventral aspects of
the lungs. These pspirated 1st stage larvae are causing a granulomatous
response. The ventral aspects of the cardiac, apical, and diaphragmatic
lobes are diffusely altered by these multiple granulomas.

73BHL-14

July 12, 1973

History: This lamb was found on the west facing rock ~lope of West Beaver.
The lamb was with 5 ewes and 3 lambs. All lambs appeared to have a slightly
rough hair coat but no coughing was observed. The lamb was shot at 1:30 PM.
The animal had not been running prior to collection.
Postmortem:
PMI - 2 hours 15 minutes
Sex - Female
Body Temp. - 106 degrees F
Weight - 32 pounds
Body Condition: This lamb has a slightly rough hair coat. A little fat is
present in the subcutis especially around the peripheral lymph nodes.
Cardiovascular System: A small amount of fat is present in the coronary'band
of the heart. The heart is normal in size, shape, color and texture.
Lymphohemopoietic System: Both retropharyngeal lymph nodes are enlarged and
juicy (size 2 cm x l~ cm x 1 em). The right node is slightly more red in
color than the left. The pre scapular lymph nodes are both enlarged and wet
(size - ~ cm x 2 cm x l~ cm). Mediastinal lymph nodes are enlarged and wet.
The cortex bulges on the cut surface and has a slightly granular appearance
(9~ em x l~ cm x 1 cm). The spleen is normal in size, shape, co lor and consistency. There are mUltiple red foci located along the periphery of the
organ (normal). The bone marrow is red and very active, no fat is present
in the marrow.
Digestive System: The rumen and abomasum are filled with ingesta and very
little milk curd is present in the abomasum. A small amount of mesenteric
fat is present. No parasites are noticeable in the alimentary tract.
Respiratory System: There is a very small amount of white exudate in both
nasal turbinates. The trachea has no exudate and the tracheal mucosa is
unremarkable. The lungs did not collapse when the thorax was opened. There
were no adhesions present and no excessive thoracic fluid. The ventral
aspect of the cardiac and apical lobes and anterior ventral aspect of the
left diaphragmatic lobes did not collapse, were red, firm and covered with
a dull pleura. The cut surface was dark red and has a slightly moist
granular appearance but no exud~te could be expressed from small cut
bronchioles. The consolidated areas were sharply delineated from the normal;

�-72-

i.e., one lobule would be affected, while the adja~ent lobule would be
normal.
The posterior dorsal aspect of both diaphragmatic lobes were
white, firm, and bulged on cut surface. The cut surface had a dull
white coloration and a granular appearance (Figure 3).
Musculoskeletal

S~stem:

The muscles,

joints and bones were unremarkable.

Urogenital System: There was a small amount of fat just under the peritoneal covering of b~th kidneys. The kidneys were normal in size, shape,
color and consistency.
Endocrine System: The adrenals, thyroids, and pituite.ry glands were normal
in size, shape, color and texture.
Nervous System: The lamb was shot in the mid-cervical
The brain and eyes were unremarkable.
Important Histopathological
1.

area (C-2 to C-4).

Findings:

Respiratory System - The ventral red-brown consolidated area is
characterized by distended alveolar capillaries.
The alveoli are
filled with neutrophils, alveolar macrophagcs, fibrin and edema.
The alveolar epithelium is hyperplastic.
Many first stage larvae
occur in the depths of the consolidated area.
The reaction is centered around bronchioles, t.uus the spread of
the pneumonic process is via peribronchiola: and down the
bronchioles.
Many bronchioles are filled with cellular debris.
The lungworm nodules are again filled with many adult, males and
females, eggs and 1st ~tage larvae. The cellular response to the
parasites are mononuclear cells, alveolar epitbelial hyperplasia,
lymphocytes and lymph nodules, and'plasma cells. The mediastinal
and prescapular lymph nodes are hyperplastic.
The remaining
systems are normal.

Summary:

Same as 73BHL-13

73BHL-15

July 24, 1973

History:
This lamb was seen at 7:30 AM with 4 ewes and 2 yearlings.
They
were about 75 yards off the Pikes Peak toll road belDw the gravel pits eating
salt (salt block) on the old trap site. The lamb was estimated to be 8-10
weeks of age and he had a rough hair coat but was not observed coughing.
The lamb was shot at 7:45 AM. First shot hit the la~b's lower jaw. Second
shot hit in the base of the skull. Temperature at collection was 104.8 degrees

F.
Postmortem:
PMI - 40 minutes
Sex - Male
Body Temp. - 104.8 degrees F
Weight - 34 pounds

�-73-

R\~ k\-

L\'\'~~
f\

od~ \~.s

U-eY\+ro. \ eon So ltda fed
CLY'ea,..

Le t-+ Lu J
1'\

Fig. 3.

Right and left lung showing ventral consolidation.

�-74-

Body Condition:
The lamb has a dull, yellow-colored rough hair coat.
There is little to no fat in the subcutis. The entire carcass is pale.
About 400 cc of blood was removed from the lamb at the collection site.
Cardiovascular System: There is one circular area of pericardium that is
thickened (1 to l~ mm) and is loosely adhered to the medial aspect of the
right apical lobe~ A small amount of fat is present in the coronary band.
The heart is normal in size, shape, color and texture. No excessive fluid
is present in the pericardial sac.
Lymphohemopoietic System: The spleen is normal in size, shape, color and
consistency.
The lymph nodules of the spleen are not prominent.
Red foci
are located along the periphery of the organ. The prescapular lymph nodes
are bilobed, enlarged (l~ in. x 3/4 in.) and moist. There are several red
foci visible on the cut surface of these nodes. Also, the cortex of the
node is prominent.
The ~etropharyngeal lymph nodes are enlarged (2 cm x
l~ cm x 1 em) and moist. The cortex is also prominent. Mediastinal nodes
are enlarged (4 in. x ~ in. x 3/4 in.), and moist. The cortex is also
prominent.
The prefemoral nodes are normal (size 2~ cm x 1 cm). The bone
marrow is red and active; no fat is present. The thymus extends 4 inches
up from the thoracic inlet. Mesenteric nodes are normal.
Digestive System: The rumen is filled with ingesta. No milk curd is present in the abomasum. No fat is present in the mesenteries.
Formed feces
are present in the colon. The liver is normal in size, shape, consistency
and color. However, there is one white focus (1 mm diameter), just under
the capsule of the liver. The gall bladder is small and unremarkable.
Respiratory System: ThP-nasal mucosa is reddened; hovever , he has been shot
both in the mandible and at the base of the skull. There are several white
flecks of material present in the trachea and blood (due to the shot) is
present in the upper half of the trachea. The lungs do not collapse. The
entire anterior-ventral aspect of the right lung is covered with a dull
yellow, rough, friable material (fibrin). This entire area is adhered to the
costal pleura; however, these adhesions are not tight and can be easily
separated. When this fibrinous coat is peeled away the pleura has a dull
sheen. ,The lung parenchyma is light grey and has a nobby or granular appearance when cut. A small amount of a slightly mucoid white material can be
expressed from small cut bronchioles.
The light grey consolidated areas have
a very meaty texture. There is no excessive fluid in the pericardial sac.
There are three areas of lungworm involvement in the right diaphragmatic lobe
These nodules are white, meaty, and elevated on pleural and cut surface. Those
nodules are speckled grey and white on cut surface. The left lung is similar
in appearance except that there is no fibrin coating the pleura. There are
four areas of lungworm involvement on the left diaphragmatic lobe. These
nodules are similar to the ones just described.
Musculoskeletal System: The femur has a normal thickness to its cortex and
the bone appears to be of normal strength. The ribs have a normal snap.
The joints and muscles are normal.
Urogenital System: The ra is a very small amount of fat covering the capsule
of the kidney just under the peritoneum.
The kidneys are normal in size,
shape, color, and texture.

�-75-

Endocrine System: The thyroids,
size, shape, color and texture.

adrenals,

and pituitary

are normal in

Nervous System: The eyes are unremarkable.
The brain is reddish in color
due to hemorrhage.
The cerebellum and brain stem was destroyed by the shot.
Histopathology:
There is depletion of thymocytes in the thymus. There are many nests of
eosinophils in the medulla also. The respiratory system has the most
important lesions.
Grossly, the normal portion of the lung has histological
alterations characterized by a thickened alveolar septa due to infilitrators
of mononuclear cells, hyperplasia of alveolar epithelium, hyperemia and a
few neutrophils.
Thus the reason the lungs do not collapse.
The ventral
aspects or consolidated area is characterized by bronchioles filled with
cellular debris and neutrophils, and the mucosa is hyperplastic.
Nearly all
of the bronchioles are surrounded by mononuclear cells, lymphocytes and a
few plasma cells. These cells have also infiltrated into the lamina propria
of the bronchioles.
Blood vessels are cuffed with lymphocytes.
The alveoli
are filled with nests of neutrophils and mononuclear inflammatory cells. The
alveolar capillaries are congested.
The lungworm nodules are filled with
adult worms, eggs, and first stage larvae. The primary cellular reaction is
nodules of lymphocytes, mononuclear inflammatory cells, multinucleated giant
cells and a few plasma cells. There are several areas of emphysema and hyperplasia of the smooth muscle of the terminal bronchioles.
The mediastinal and
retropharyngeal lymph nodes have a lymphocytic hyperplasia.
Summary:
The main gross and histological lesions are restricted to the respiratory and lymphohemopoietic systems. The lung has lungworms (in the
dorsal posterior portion of both diaphragmatic lobes), and an interstitial
reaction in the normal colored non-collapsed areas. There is an exudative
pneumonia in the ventral aspect of the lungs.
The lamb, considering
a week.

73BHL-16

the extent of pneumonia,

would not have lived much over

July 26, 1973

History:
This lamb was seen at 6:00 AM above the Pikes Peak toll road just
below Boulder Park. He was with one other lamb and two ewes. The ewes were
about 1/3 shed of their ~rlnter coat. Both lambs were small and estimated to
be 8-10 weeks of age. Both lambs had a rough hair coat but no coughing was
observed.
The lamb was shot in the neck at 7:00 AM.
Postmortem:
PMI - 1 hour
Sex - Male
Body Temp. - 104.8 degrees F
Weight - 28 pounds
Body Condition:
The lamb has a dull yellow roughened,
fat is present in the subcutis.

brittle hair coat.

No

�-76-

Cardiovascular System: The heart is normal in size, shape, color and
texture. No excessive pericardial fluid is present. There is a small
amount of fat in the coronary band.
Lymphohemopoietic System: The spleen is normal in size, shape, color and
texture. The malpighian corpuscles are not prominent. The prescapular
lymph nodes are s~.ightly enlarged and wet (3 em x l~ em x l~ cm). The
retropharyngeal lymph nodes are enlarged, white colored and moist. The
mediastinal nodes are enlarged, moist and have a slight bulge on cut surface (size 0 5~ cm x l~ cm x l~ cm). The thymus is normal in size and
extends from the base of the heart to within two inches above the thoracic
inlet. The bone marrow is red and active. Mesenteric nodes are normal.
Digestive System: The rumen is filled with green grass. The abomasum is
filled with green grass and with a moderate amount of milk curd. No fat
is present in the mesenteries and formed fecal pellets pre in the colon and
rectum.
Respiratory System: The nasal mucosa is reddish-colored and white, mucoid
exudate is present on the turbinates. The trachea is slightly red in color,
but no exudate covers the mucosa. The lungs do not collapse when the thorax
is opened. The anterior 2/3 of the right apical and lower ~ of the cardiac
lobes are red, firm and covered with a shiny pleura. There are many small
grey clover-shaped foci r~to 2/3 mm) located in the consolidated area. A
white mucoid material can be expressed from small cut bronchioles. The cut
surface also has these small grey nodules, giving the cut surface a nobby
appearance. Many of the septa between the lobules in the consolidated part
have enlarged due to an accumulation of a clear yellow gelatinous material.
The altered portion of the lungs are sharply demarcated from the noncollapse pale pink lung parenchyma. Lungworm nodules are present in the
posterior dorsal aspect of both diaphragmatic lobes. These nodules are
white, firm and raised from the cut surface.
Musculoskeletal System: The muscles and bones are unremarkable. The cortex
of the femur is normal in thickness. The joints are clean and fat is present in the stifle joint.
Urogenital System: There is no fat around the kidneys. Both kidneys are
normal in size, shape, color and texture. Reproductive tract is unremarkable.
Endocrine System: The adrenals, thyroids, adrenal glands are normal in size,
shape, color and texture.

Histopathology: The primary changes in this lamb are identical to 73BHL-15
and will not, therefore, be repeated.

�-77-

73BHL-17

July 26, 1973

History: This lamb was found dead on 26 July 1973 under some rocks (in
the shade) at 1:00 PM. He must have died that night because few postmortem changes had occurred.
The reason the lamb was found was because
the ewe would not leave. Also this ewe was a white/red stripe #7,
(tramisol treated). The ewe was caught 14 May 1973 at the gravel pits.
The lamb was found northeast of the cog rail and just southwest of Bar
trail (near the summit) in the steep rocky benches.
Postmortem:
PMI - Estimated to be 14 to 18 hours
Sex - Male
Body Temp. - ?
Weight - 21 pounds
Body Condition:
This lamb has a rough, dull yellow hair coat. A few body
lice are present between the hind legs. The ears, anterior dorsal part of
the muzzle and hoofs have been chewed by rodents. The eyes are slightly
sunken and the animal is emaciated.
Cardiovascular System: The heart is normal in size, shape, color and texture. No fluid is present in the pericardial sac. There is a very small
amount of fat in the coronary band.
Lymphohemopoietic System: The spleen is slightly enlarged (11 cm x 8 xm x
3 cm) and the cut surface is mushy (due to postmorten autolysis).
The
retropharyngeal lymph nodes are only slightly enlarged (2 x l~ x 3/4 cm)
and their cut surface is dry. The prescapular nodes are slightly enlarged.
The prefemorals are unremarkable.
The mediastinal lymph nodes are enlarged
and wet (size 9~ em x l~ cm x 3/4 em). The thymus is extremely small. None
can be found beyond the thoracic inlet and only a small piece can be found
just anterior to the heart.
Digestive System: The rumen is half filled with dry ingesta. No fluid is
present.
The abomasum is empty, indicating he has not had any water for
the last day or two preceding death. Only few formed fecal pellets are present in the colon and rectum. No fat is present in the mesenteries.
The
liver has a distended gall bladder (indicates that the lamb has not eaten
for a day or two prior to death). The liver is otherwise normal. No white
foci, as with other lambs, were found.
Respiratory System: The nose and anterior dorsal portion of the muzzle has
been eaten by rodents. The nasal mucosa is red-colored and several white
flecks of exudate are present on the surface. A small amount of exudate
is covering the nasal turbinates and ethimoid bones. The trachea also has
several flecks of white exudate on its mucosa.
The mucosa has a slight
red coloration.
The lungs do not collapse when the thorax is opened. The
anterior and ventral aspect of the right apical, right and left cardiac and
diaphragmatic lobes are similar to other lambs. The consolidated areas are
reddish-brown, firm and the pleura covering it is shiny. The cut surface

�-78-

is moist and dark rel and has a knobby appearance.
Mucoid exudate is not
easily expressed from some cut bronchio1es.
Three small (1 to 3 cm) lungworm nodules are present on the posterior dorsal aspect of the right diaphragmatic lobe and two on the left. They are raised from the pleural surface, and
their cut surface is speckled grey-white.
These nodules are closer in appearance
to 73BHL-12 than to the white firm nodules of 73BHL-13, 14, 15, 16.
Musculoskeletal System: There is less muscle mass than in other lambs. Also,
the muscle appears slightly dry. The joints are clean, and the bone has a normal thickness to the cortex. The ribs have a slight bend than a snap.
Urogenital System: The kidneys have a normal color, texture, size and shape.
There is no fat on the kidneys.
About 40 cc of urine is present in the bladder.
Both testicles are normal.
Endocrine
Nervous

System:
System:

The thyroids,

adrenals, pituitary

glands are unremarkable.

Unremarkable.

Remarks:
This lamb died from the same disease (pneumonia) that is affecting all
lambs on the Peak. This lamb evidently did not eat one or two days prior to death
and died during the ~ight. The histopathology is similar to 73BHL-15.
73BHL-19

August 14,1973

History:
This lamb was seen with 6 ewes and one other lamb by the cog rail near
the summit of Pikes Peak. It did not look ill, but the other lamb had a dull
yellow, rough hair coat. The lamb was shot low in the neck and she ran over the
near Bar trail and had to be shot again. The lamb was shot at 2:00 PM.
Postmortem:
PMI - 1 hour 35 minutes
Sex - Female
Body Temp. - 104.8 degrees F.
Weight - 36 pounds
Body Condition:
This lamb has shed about half its lamb pelage showing a smooth,
slick ~oat about the chest and neck.
All systems grossly and histologically

are similar to 73BHL-16.

SEGMENT OBJECTIVE

#3

Continue visual surveillance of the Pikes Peak herd to further elucidate the
characteristics of the lamb mortality currently underway, and intiate a visual
surveillance program in the Saguache herd. These observations may help us relate
histological lesions to behavioral changes of lambs.
PROCEDURE
Continue a visual surveillance of the Pikes Peak herd, and initiate a visual
surveillance of the Saguache herd. Accurate documentation of the die-off currently
underway at Pikes Peak necEssitates a close surveillance of this herd to evaluate
all the factors involved.
The Saguache herd should likewise be visually surveyed
and contrasted with the Pikes Peak herd. In the ev~nt that any morbidity becomes
evident in the Saguache herd, or mortality of lambs occurs, a course of action
similar to the study on Pikes Peak should be initiated.

�-79RESULTS
A visual surveillance of the Saguache herd was not initiated because of time
and personnel limitations.
However, the Pikes Peak herd was closely monitored
by observations, trapping and tagging. Robert Schmidt, Division of Wildlife
was in charge of this operation and the results are incorporated in his report.
SEGMENT OBJECTIVE

#4

Continue pathology support for experimental projects in parasitology, virology, bacteriology, immunology, trace mineral evaluati0n and physiological
baseline studies.
PROCEDURE
Continue pathology support for experimental projects in parasitology, virology, bacteriology, immunology, trace mineral evaluation and physiological
baseline studies. Details are given under Procedure #2.
RESULTS
This is a routine objective incorporated in the postmortem procedure used to
determine the cause of illness and/or death and tt.e events leading to death.
The results are given under the various headings; i.e., virology, bacteriology,
parasitology and pathology.
SEGMENT OBJECTIVE

#5

Continue clinical pathology studies, especially on blood serum, urine, and
tissue fluids from experimental bighorn sheep, field collections of bighorn
lambs, and all other bighorn sheep dead or dying from any cause.
PROCEDURE
Continue clinical pathology studies especially on blood, serum, urine and
tissue fluids from experimental bighorn sheep, field collection of bighorn
lambs, and all other bighorn sheep dead or dying from any cause. Most of
this data will be determined by the research team in cullaboration with the
Clinical Pathology Laboratory at CSU.
RESULTS
The majority of the postmortem examinations
from Pikes Peak lambs and the ewes captured
data are given in summary form below.

conducted during 1972-73 were
in January-February, 1973. These

Acute Septicemic Pasteurellosis
Nine bighorn sheep were captured at Pikes Peak and placed in the fenced area
at the Division of Wildlife bird pens on Highway 25 in January and February
1973. These sheep consisted of 6 ewes, 2 yearling rams and one 3 year old
ram. An additional yearling ram from the Denver zoo was also penned here.
These sheep were doing quite well and had no problems until the night of 28

�-80-

October when a cold rain with lots of wind began. These weather conditions
continued through the next day (29 October).
The next morning, 30 October
Archie (Denver Sheep) and one ewe were found to be very ill and they died
within 30 minutes.
The next day two more sheep were found dead. Then one
sheep died each day until all had died except one ewe (Tag #5). These
sheep all died of ~he same disease - a typical necropsy report follows.
The hair coat is smooth and slick. There is an abundance
Body Condition:
of subcutis fat. The animal is in excellent condition.
Cardiovascular System: There is an excessive amount of pericardial fluid
with fibrin present in the fluid. There are massive hemorrhages on the
epicardium and endocardium.
The myocardium has a slightly mottled appearance.
Massive hemorrhages ar.e present on the parietal pleura of the ribs. An
abundance of fat is present in the coronary band.
Lymphohemopoietic
System: Most of the lymph nodes are enlarged, wet, and
red. A reddish fluid exudes from the cut surface of these nodes.
The
spleen is normal in size and shape. The bone marrow is white opaque (normal)
and filled with fat.
Digestive System: The liver is normal in size, shape, and color. Several
of the sheep have a tapeworm present in the bile ducts (Wyominia tetoni).
The digestive system has multiple hemorrhages on the serosal surface,
especially on the rumen. Formed feces are present in the colon and rectum.
Respiratory System: The nasal mucosa and trachea are highly inflammed with a
red froth covering the s~rface. The lungs do not collapse any when the thorax
is opened. A slight but excessive volume of a clear yellow fluid is present
in the thorax of several sheep. This fluid clotted on exposure to air
(indicates high fibrin content in fluid). The anterior ventral aspect of the
lungs are usually covered with a thin layer of yellow, friable material which
can be easily peeled away (fibrin). The lung under this fibrin is red and
firm, and has a fine granular appearance.
A few septa are widely distended
by a yellow gelatinous material.
A small amount of watery red fluid can be
expressed from bronchioles.
Lungs from some sheep are soft, but not collapse
(due to pulmonary congestion and edema). All sheep have mUltiple lungworm
nodules in the posterior dorsal aspect of the diaphragmatic lobes.
Protostrongylus rushi is not present in any of the sheep.
Musculoskeletal System:
The bones are normal.

There are a few hemorrhages

present in muscles.

Urogenital System:
There is a moderate

The kidneys are normal in size. shape, color and texture.
amount of fat around both kidneys.

Endocrine System:
due to congestion.

Adrenals

Nervous System:

are slightly

larger than normal and red in color

The brain is mildly congested.

Eyes are unremarkable.

Remarks:
Pure cultures of Pasteurella hemolytica were isolated from the spleen,
peripheral blood and lung tissue. Septicemia pasteurellosis is an acute,
highly contagious disease of bighorn sheep. The mortality is nearly 100 percent. The cold rains of 29 October must have triggered this disease in our

�-81-

captive sheep. A virus was isolated from one of the sheep t.hat died but
it has not been characterized to date; its role in Acute Septicemic
Pasteurellosis is unknown. This disease claimed 8 of the 9 sheep present.
We now have an autogenous vaccine, but do not know if the vaccine will
protect the animals as of yet.

Mouflon/BighornSheep
Twelve mouflon/bighorn cross ewes and 4 rams are located in the pens behind the Division of Wildlife building on Prospect Street. The adult
animals are doing fine. Lambing began during May and continued until midJune, 1974. The first four lambs that were born died just minutes after
birth or were stillborn.
Four lambs were taken from the ewes because two
ewes had mastitus and two refused to accept the lambs. These four lambs
are healthy to date. Four other lambs were left with their ewes. They
all died at 7 to 8 weeks of age from an interstitial pneumonia.
Culture
and histopathology has not been completed, so more detail on etiology is
not possible at this time. In summary, 12 lambs were born this year, 4
were stillborn or died within minutes of birth, 4 died at 7 to 8 weeks
of age due to an interstitial pneumonia, and 4 are being hand-raised and
are healthy to date. The 12 ewes and 4 rams are also in good shape now.
However, when the lambs were born the ewes were in poor body condition,
a result of inadequate nutrition (inadequate amount).

South Dakota Bighorn Sheep
Bighorn sheep were live trapped in Custer State Park, South Dakota during
January of 1974. Twenty-six of these sheep were loaded ~d transported to
Fort Collins. Two ewes died within 3 hours of capture, and one yearling
ram died shortly after arrvTal in Fort Collins. The remaining sheep were
released in the bird pens at the Research Station on Interstate 25. Following release, seven sheep ruptured their gastrocnemius muscles.
All of these
sheep were cast and only one (White/Red #12) is still alive and doing well;
the other six died (5 ewes and a large 4 year-old ram). One of the ewes that
died ha4 a Wyoming ear tag #D-1870, Wyoming Game and Fish Co~.ission.
The problem that caused the death of these 6 sheep appears to be a "capture
myopathy".
The cause and nature of "capture myopathy" is unknown, but it has
been seen in other situations (wildlife and zoo animals) allover
the world.
The remaining sheep include 13 ewes, 3 yearling ewes and one two-year old ram.
All are healthy to date. During May and June, a total of six lambs were born,
the first lamb was abandoned and died from starvation.
The second lamb died
during parturition from a dystocia caused by the umbilical cord wrapping around
the hocks with the lamb in a breech presentation; thus the lamb never breathed.
One lamb that had been refused by the ewe was hand-raised and was doing very
well but died at 9 weeks of age due to enteritis caused by Salmonella typhimurium
var. copenhagen.
Presently, there are three lambs, all doing well at the bird
pens.
The adult sheep have Meullerius capillaris, a lungworm that inhabits the same
area (posterior dorsal aspect of the diaphragmatic lobes) in the parenchyma
of the lungs as P. stilesi. This worm as far as we know does not cross the
placenta and does not predispose lambs to verminous pneumonia.
Thus, lambs
born of these ewes (and the ewes themselves) can be infected with P. stilesi

�-82-

for chemotherapy

studies.

This is currently underway.

SEGMENT OBJECTIVE

#6

Evaluate the role of transplacental transmission of Protostrongylus as
a predisposing factor in the etiology of "Spontaneous Lamb Mortality".

PROCEDURE
Evaluation of the role of transplacental transmission of Protostrongylus
as a predisposing factor in the etiology of "Spontaneous Lamb Mortality".
In late 1972, before the bighorn sheep ewes at Little Hills became pregnant,
two ewes were given a large dose of infective Protostrongylus larvae, the
purpose being to determine if they were capable of storing these larvae in
their tissues and transmitting them during late pregnancy to the foetus.
Two other ewes were given "trickle" doses of infective larvae from midpregnancy to parturition to determine if they passed the larvae directly
to the foetus rather than store them. Four sheep are an insignificant
number of animals for this purpose, but hopefully will serve the purpose
as a pilot experiment.
During the summer, fall and winter of 1973, the 4 tamest ewes at Little
Hills will again be used for this purpose. Two will be fed massive doses
of infective larvae before pregnancy, and one ewe and one ram kept as uninfected controls. Lambs born of these ewes will be closely monitored for
evidence of "Verminous Pneumonia" as a result of transplacental infection.
Since the factor predisposing lambs to pneumonia may be a combination of
transplacental transmission, and a later "triggering response" from infection
acquired after birth, an additional two lambs (possible from the captive
Pikes Peak animals) will be challenged .at the age of 3-4 weeks with a dose
of infective larvae. One will be given a small dose, the other a massive
dose. We will do this with two lambs this summer at Little Hills.

RESULTS
As noted in the 1973 annual report (Segment Objective #4) the data from
lambs transplacentally infected with lungworm in experiments conducted at
Little Hills was incomplete.
This experiment was completed in 1973 however,
and is incorporated as a portion of a M.S. thesis at C.S.U. submitted by
Robert E. Lange. The thesis is entitled "Epidemiology of Lungworms
(Protostrongylus stilesi and rushi) in Rocky Mountain Bighorn Sheep (Ovis
canadensis canadensis."
That particular experiment is given below.
Two ewes (Ewe I and Ewe II) were selected at Little Hills Game Experiment
Station near Meeker, Colorado, to test somatic storage of third-stage
protostrongylin larvae.
Lltermediate host snails were cultured and infected
in the laboratory at Colorado State University.
Larvae were permitted to
mature in the snails, Vallonia pulchella and Vallonia cyclophorella for
known periods of time. Infected snails were periodically transported to
Little Hills where they were fed to the test sheep. Infected snails were
placed in number one gelatin capsules and there were secreted in hollowedout alfalfa cubes. The alfalfa cubes were fed by hand to the test sheep

�-83-

in most cases. Administrations were made by Steve Steinert~ Research
Assistant at Little Hills, and by this author. Total dOBages to the two
ewes were completed before breeding season. Ewe I received 754 and Ewe
II 760 mature third-stage lungworm larvae. The test was devised so that
lambs of these two ewes, in order to pass first-stage larvae the following
summer, could only d0 so through the mechanism of somatic storage of administered third-stage larvae and prenatal infection of the fetus during pregnancy.
In the second portion of the study, two other ewes (Ewe III and Ewe IV)
were selected for infection after they were bred. These ewes were administered third-stage larvae between February and late April. Ewe III received
1,643 larvae and Ewe IV received 1,489. Each of the larvae matured in snails
for a specific length of time. Lambs from these two ewes were used to demonstrate direct transplacental transmission if somatic storage did not occur.
However, lambs from these two ewes were unable to rule out somatic storage
if they did pass first-stage larvae. Only lambs born to Ewes I and II could
do that. The ram in the herd at Little Hills Eerved as a control because
he never had a patent lungworm infection.
Plans to test somatic storage and transplacental transmission of third-stage
lungworm larvae were bolstered when all four test ewes bore lambs. Ewe I
born a ewe lamb between June 19 and 24. Ewe II had a ewe lamb between June
2 and 8. Ewe III had a ram lamb between May 20 and 31 and Ewe IV had a ewe
lamb between May 2C and 31.
Ewe II was the only ewe that passed obviously increased levels of first-stage
larvae as a result of feeding third-stage lungworm larvae. This ewe averaged
no more than six larvae per g of feces during 15 months before administration
of third-stage larvae began. A dramatic increase in first-stage larvae output followed administration of infective lungworm larvae. The increase in
larva output followed administration of third-stage larvae by 6-8 weeks.
Seven hundred sixty infective third-stage larvae were fed to this ewe. Ewe
II was less than two years old during the study and her lungworm load, as
measured by output of fir.st-stage larvae, was very low until treatments began.
Other ewes (I, II~ and IV), to which equal or larger numbers of third-stage
larvae were administered, failed to show readily detectable increases in
lungw~rm loads. These ewes were administered 760, 1,643, and 1,489 thirdstage larvae respectively.
Higher lungworm loads existed in these ewes than
in Ewe II prior to treatments.
Completion of the lungworm life cycle was
accomplieshed in Ewe II. It is uncertain that artificial increase in lungworm loads was accomplished in Ewes I, III and IV.
Transplacental transmission of lungworms was experimentally reproduced in
the study. Lambs were born to each of the ewes administered third-stage
larvae. Lambs were assigned the same experimental number as their dam.
Lambs 2, 3 and 4 began passing first-stage larvae 33-54 days after birth
(Table 2). Lambs 3 and 4 were from ewes who had received third-stage
larvae during gestation.
Both of these lambs passed larvae (Table 2).
Lamb 2, which was a test of somatic storage of third-stage lungworm larvae,
also passed first-stage larvae giving some indication that the mechanism
does occur. Lamb 1 had passed no larvae at writing.
The number of larvae passed by lambs 2, 3 and 4 were quite low considering
the numbers of infective third-stage larvae their dams received.

�-84-

As a result of unforeseen difficulties originating in other aspects of
this research, the plans to repeat the experiment at Little Hills was
not possible.
Therefore, these animals will be transferred to Fort Collins
facilities and the experiment conducted there.

Table 2. Passage of first-stage larvae in numbers per g of air-dried
feces by lambs at Little Hills Experiment Station.
Date

Lamb 1*

Lamb 2**

6-15-73
6-19
6-20
6-21
6-24

0

6-25
6-26
6-27
6-28

0
0
0
0

,Lamb 3t

0
0
0
0
0

0
0
0 &amp; 0
0

0

0
0
24

7-11
7-13
7-14
7-15
8-1

0
0

0

8-5

0

0

*Lamb 1 was born between June 19-24.
**Lamb 2 was born between June 2-8.
tLamb 3 was born between May 20-31.
tt

0
0

6

0

7-6
7-7
7-8

0
0

7-1
0

0

0

6-29
7-2
7-3
7-4
7-5

Lamb 4tt

Lamb 4 was born between May 20-31.

0

1
2
77

7
35
6
8
3 &amp; 9
2
1
0
16

�-85SEGMENT OBJECTIVE

117

Attempt to comvlete the biological cycle of Protostrongylus
domestic sheep or in bighorn-domestic crosses.

stilesi in

PROCEDURE
Attempt to complete the biological cycle of Protostrongylus using domestic
sheep (or half-breeds).
Considerable information can be obtained about
lungworm by using an abnormal host, providing infection can be established.
All previous attempts to infect domestic sheep have failed (but have been
successful in half-breeds); thus they are considered to be refractory to
infection.
However, infection with parasites has been accomplished in
animals normally refractory to infection by the administration of immunosuppressant compound~ such as cortisone or massive protein injection; moreover, drainage of the thoracic duct is also an effective means of establishing
otherwise refractory infections.
RESULTS
Between the time Objective 7 was written and before domestic sheep could be
bred to fulfill this objective, we fortunately obtained some mouflon/bighorn
hybrids from Pennsylvania.
Twelve of these animals were used for the experiment; four were infected and 8 were controls.
The experiment had a two-fold
purpose: (1) Establish infection in the animal, and (2) Attempt to induce
transplacental infection.
The 4 ewes infected with larvae of ~. stilesi (via snail-feeding) began to
pass first-stage larvae of f. stilesi on schedule (30 to 40 days postinfection).
Thus, the life-cycle of this parasite has been completed in
hybrid sheep. Attempts to induce transplacental infection of the lambs did
not appear to be successful with the initial infection, but we did not feel
that transplacental would occur until the second exposure (2nd year of
infection) • At present, two of the fufected ewes are being "trickle dosed"
with infected snails to induce transplacental transmission; two additional
(uninfected) ewes will be started shortly.
SEGMENT OBJECTIVE

#8

Continue studies on the epizootiology of Protostrongylus in bighorn sheep at
Little Hills Experiment Station, including transplacental transmission,
efficacy of chemotherapeutic agents and of intermediate host population fluctuation.
PROCEDURE
Continue studies on the epizootiology of Protostr.ongylus in bighorn sheep at
Little Hills. Lungworm larval output from these sheep has been monitored for
two years. Monitoring of the adults will continue on a bi-month1y basis.
The
date that lambs born of these ewes begin p~oducing larvae, and the level of
infection, will be monitored on a weekly basis. Lungworm larval output from
animals reaches a peak in the winter months, and decreases in the summer. This
may be a reflection of increased fecundity of lungworm in the winter, or a loss
of resistance by the adult as a result of stress. Winter is obviously a period
of considerable stress which may lower body resistance, thus transplacental
transmission may occur during this period simply because the adult is incapable

�-86of destroying the larvae she ingests. While information of thiS nature is
merely suggestive, all supporting observations will help to piece together
the puzzle. Moreover, these data are necessary support for the objectives
given in number 6 above.
RESULTS
This study is comPlete and is the subject of a Master's Thesis by Robert
Lange. The title is "Epidemiology of Lungworm (Protostrongylus stilesi
and rushi) in Rocky Mountain Bighorn Sheep (Oviscan:adensis canadensis)".
This thesis may be obtained from the library at Colorado State University.
It will not be incorporated in this report, resulting in unnecessary expense.
SEGMENT OBJECTIVE

#9

Evaluation of chemotherapeutic agents for treatment of bighorn sheep to kill
adult lungworms and the third-stage infective larvae stored in somatic tissues.
This work will be conducted at Little Hills and in Fort Collins through carefully selected field trials of chemotherapeutic agents.
PROCEDURE
Evaluation of chemotherapeutic agents to kill adult lungworm and the larval
stages stored in tissues is currently underway with the 12 sheep captured on
Pikes Peak. Four of these sheep were treated with Tramiso1 and four with
Cambendazole.
Lungworm larval output is being monitored, and the lambs born
of these ewes will be followed closely for evidence of transplacental infection
beginning 30-40 days after birth. Lambs from untreated ewes, and showing a
heavy infection, will be treated in late summer or early fall and monitored
for efficacy of the chemotherapy.
Three anthelmintics will be used: Tramisol,
Cambendazole and Hygromycin B.
In addition to evaluating the efficacy of the drug, a vehicle suitable for
administration will likewise be evaluated.
The vehicle showing the most promise at present is "Apple Mash". This coming year it will be the vehicle of
choice for all drug trials.
RESULTS
The chemotherapeutic trials have been undertaken by personnel in the University
and in Division of Wildlife; consequently, there may be some overlap between
this report and the ~ne submitted by Robert Schmidt.
Ewes were treated on Pikes Peak during the winter of 1972-73 with Cambendazole
and Tramisol.
Most of these animals were treated in early pregnancy.
As near
as can be determined, all lambs from these ewes died. One group of 7 ewes
were treated in late pregnancy with Cambendazole.
Five of these ewes are
known to have had lambs. All five survived the winter and the last time they
were observed (Spring, 1974) all were quite healthy.
This limited trial
strongly suggests that Cambendazole is effective in killing the third-stage
larvae of Protostrongylus stilesi. Apparently the drug does not effectively
kill the larave stored in the ewe; however, this drug is known to cross the
placenta.
The results suggest that the larva, once in the lamb, is susceptible
to treatment by this drug.

�-87-

Of the 12 ewes captured on Pikes Peak and retained in captivity, four were
treated with Tramisol, four with Cambendazole and four were untreated controls. All of the lambs born (alive and/or stillbirths) of the treated ewes
had heavy lungworm loads and all eventually died of a pneumonia identical
to the disease seen on Pikes Peak.
During the winter of 1973-74, the drugs evaluated were: Tramisol and Diethylcarbamazine, Thiabendazole and Diethylcarbamazine,
Cambendazole and Dichlorvos.
These trials are not yet complete and will be reported in 1975.
SEGMENT OBJECTIVE

#10

Surveillance of captive bighorn sheep ewes and lambs for evidence of the disease
responsible for the high rate of mortality on Pikes Peak.
PROCEDURE
Surveillance of captive bighorn sheep ewes and lambs for evidence of the disease
responsible for lamb mortality.
Initial plans for the lambs born of the ewes
captured on Pikes Peak w·as to infect them with various combinations of the three
agents considered to be responsible for the mortality.
However, when faced with
the fact that we were going to attempt infection with pathogens without first
doing a surveillance of a captive herd of ewes and lambs for a year to determine what they became infected with, and when, we realized th~t any attempt
at infection during the first year without this information could possibly
result in difficult interpretation of data. Therefore, in 1973, the surveillance approach will be utilized.
Lambs from these ewes will be given a complete
clinical examination every two weeks. Not only will this furnish sound baseline data for future studies, it may possibly furnish information on the development of the pneumonia responsible for lamb mortality.
These monitoring
plans could be changed if increased mortality results in our holding pens in
Fort Collins.
In such an event some or all of the ewes and lambs might be
transported to Little Hills.
RESULTS
The decision to pursue Objective #10 by the procedure given above was fortunate.
All of the lambs died of a typical "vermir..ouspneumonia" characteristic of the
die-off among the free-ranging wild sheep on Pikes Peak. The results from
these lambs were reported last year (1973 Annual Report) under Segment Objective
#8. At the first onset of cold weather (first winter storm) in November of
last year, all except one of the captive sheep from Pikes Peak died of
Pasteurellosis within a 5 day period. A summary of the postmortem results is
given in this report. Had the initial plans been attempted (to infect these
lambs with the various pathogenic agents) the results would have been impossible
to interpret.

SEGMENT OBJECTIVE

#11

Monitor two herds for evidence of any factors predisposing
a die-off.

the population

to

�-88-

PROCEDURE
Monitor two herds of bighorn sheep for evidence of the factors predisposing
the population to a die-off.
Trapping operations should be continued on
Pikes Peak and trapping should be initiated at Saguache.
The Pikes Peak
herd well represeqts a dying herd, while the population at Saguache is an
example of a thriving herd. A thorough job of monitoring necessitates that
clinical information be obtained fr0m two such herds. From 25-50 animals
in each herd should be trapped and examined: (1) Age, (2) Body condition,
(3) Nasal and rectal swabs for bacteria and virus, (4) Blood for complete
clinical pathology evaluation, (5) Serum for bacterial and viral titers,
arid physiological baseline data, and (6) Feces to dete~ine level of lungworm and gastrointestinal parasite burden.

RESULTS
Monitor of two herds was found to be impossible.
While trapping operations
were scheduled and conducted on Pikes Peak this last year - and information
was obtained - we were not notified of, nor invited to, the trapping operations underway at Saguache; therefore, we had no basis for comparing the
health and disease status of a healthy and dying herd. While we know that
pathogenic agents (and in what combinations) are found at Pikes Peak, we
do not know what is present at Saguache.
Thus, the efforts made on Pikes
Peak were essentially fruitless.

SEGMENT OBJECTIVE

#12

Determine the types of immune ~esponses that are stimulated in bighorn sheep
by the three pathogens (1) PI-3, (2) Pasteurella hemolytica, and (3) Protostrongylus stilesi.

PROCEDURE
The h4moral response will be detected by immunoelectrophoresis
using specific
soluble antigen from the pathogen.
Specific antibody reagents will be produced to detect the various immunoglobulin classes of bighorn sheep. These
class specific anti-immunoglobulin reagents will be used to verify the immunoelectrophoresis observations and aid in direct and indirect fluorescent antibody studies of diseased tissues for the pathogens.
The cell-mediated response will be detected by skin tests in vivo and macrophage migration inhibition tests in vitro. The cytophilic antibody response
will be detected by passive cutaneous anaphylaxis tests and by development
of a specific anti-IgE reagent.
The cytophilic antibody could also be detected indirectly by specific macrophage attachment of antibodies to Protostrongylus stilesi and its inactivation.

RESULTS
Segment Objective #12 has not been attempted; however, serum from all of the
sheep collected thus far has been stored for this purpose.

�-89-

SEGMENT OBJECTIVE

#13

Determine the bacterial agents involved in bighorn sheep disease by
continuation of serological survey and isolation attempts.

PROCEDURE
Serology will be performed by screening sera from bighorn sheep for precipitating, agglutinating or complement fixing antibodies to antigens from
the various potential pathogens isolated previously from bighorn sheep.
Attempts to isolate pathogens from nasal swabs and lesion material will be
continued using standard bacteriological isolation and identification
techniques.

RESULTS
These data are summarized below. A table, noting the location of Pasteurella
spp. in these lambs is included.
Note that as the summer progressed, and the
observed pneumonia became more severe, Pasteurella moved into the lung proper.

Bacteriological

Summary

73BHL-ll
Nasal:

Trachea:

Staphylococcus sp.
Alpha Streptococcus
Pasteurellamultocida

sp.

Escherichia coli
Corynebacterium sp.
Staphylococcus sp.
Pasteurella multocida

Nasal:

Pasteurella multocida
Streptococcus sp.---Mima sp.
Micrococcus sp.

Lungs:

Staphylococcus sn.- rt. cardiac
Bacillus sp. - rt. diaphragmatic
Staphylococcus sp. - rt. diaphragmatic
Corynebacterium - rt. diaphragmatic nodule
unknown (not Pasteurella) - rt. diagphragmatic

Spleen:

Bacillus sp.
Clostridium (suspect)

Retropharyngeal

lymph node:

Lactobacillus

sp.

nodule

�-90-

73BHL-13
Nasal:

Bacillus sp.

Spleen:

Staphylococcus

sp.

73BHL-14
Nasal:

Staphylococcus sp.
Pasteurellamultocida
Alk.aligertes sp ,
Corynebacterium sp.

Trachea:

Pasteurella multocida
Corynebacterium sp.
Mirna sp.
Herellea sp.

Liver:

Escherichia

coli

73BHL-15
Nasal:

Streptococcus
Bacillus sp.
Mirna suspect

Trachea:

sp.

Neisseria sp.
Bacillus suspect
Alpha Streptococcus sp.
Pasteurella multocida

Retropharyngeal

lymph node:

Main stem bronchus:

Escherichia

coli

Bacillus sp. (contaminant)

73BHL-16
Nasal:

Trachea:

Pasteurella hemolytica
Pasteurella multocida
Bacillus sp.
Staphylococcus sp.
Acromonas sp.
Staphylococcus sp.
Acromonas sp.
Pasteurella hemolytica suspect
Alpha Streptococcus sp.

Main stem bronchus:

Lungworm nodule:

Corynebacterium sp.
Alpha Streptococcus sp.

Pasteurella hemolytica

Lobe of lung (consolidated):

Pasteurella

multocida

�-91-

73BHL-17
Found dead on Pikes Peak.
Pasteurella not isolated.
Many opportunists in various

tissues

(probably ,spread after death).

73BHL-18
Nasal:

Trachea:
Lungs:

Staphylococcus sp.
Bacillus sp.
Corynebacterium p'yogenes
Pasteurella multocida
Pasteurella
Pasteurella
Pasteurella

~ultocida
sp. - rt. diaphragmatic
mu1::ocida - rt. ventral

Main stem bronchus:

Pasteurella

multocida

73BHL-19
Nasal:

Trachea:

Pneumococcus sp.
Moraxel1a sp. (suspect)
Pasteurella hemolytica
Corynebacterium
sp.
Alpha Streptococcus sp.
Bacillus sp.

Lobe of lung (consolidated):
Lungworm

nodule:

PasteureLla

hemo1ytica

Mima po1ymorpha

Main ~tem bronchus:

Pasteurella

hemolytica

SEGMENT OBJECTIVE
Determine the viral agents involved
of serological survey and isolation

#14

in bighorn
attempts.

sheep disease by continuation

PROCIDURE
Serological survey will be performed by screening sera from bighorn sheep for
virus neutralizing antibodies in tissue culture systems.
The virus to be used
in the survey are: (1) Para-inf1uenza-3;
(2) Bovine Virus Diarrhea; (3) Infectious Bovine Rhinotracheitis;
(4) Blue Tongue Virus; (5) Epizootic
liemorrhagic Disease of Deer; (6) Parvovirus; (7) Adenovirus and any other virus
that becomes available for study that is considered a potential pathogen.
Virus isolation attempts will be made in various tissue culture systems in
search of the above named viruses.

�-92-

RESULTS
As in previous years, a concentrated effort was made to recover all possible
infectious agents potentially involved as pathogens in lamb mortality.
In
the past, bighorn sheep have had titers to a number of different viral agents
(see previous reports) and so~e viral agents (PI-3) have been isolated.
Year
after year however, our results are inconsistent - we get different viral
agents - making the situation questionable regarding viral agents as being
detrimental to the host. Every year that we find a different virus, however,
the question arises once again. This last year an extremely pathogenic virus
was found in a considerable number of sheep from Pikes Peak (including George).
Unfortunately, we have not yet identified this virus. We know that it is not
PI-3, BVD (Bovine Viral Diarrhea) or IBR (Infectious Bovine Rhinotracheitis).
It is still in culture and we hope to get an answer this year. As with PI-3,
it may be insignificant.
From the inception of this project we felt that we must consider Mycoplasma,
because some species, e.g. M. arginini is a pathogen responsible for pneumonia.
The second year an unsuccessful attempt was made. The third year, however,
was successful.
As indicated on the table, a species was receovered from the
respiratory system in 9 of 13 sheep examined.
Two (7 and 17) have been identified as~. arginini, a known pulmonary pathogen.
Tnis role in lamb deaths (in
the wilds) is still unclear, but we know that bighorn/mouflon hybrids and pure
mouflons are extremely susceptible and that this sPecies will kill lambs if
they remain on the ewes (in captivity).

Parasitology
The parasitology efforts this past year have been confined to (1) Recovery of
lungworm larvae from transplacentally infected lambs dying spontaneously,
(2) Recovery of lungworm larvae from lambs collected at Pikes Peak, and (3)
Attempts to find the site of somatic storage of third-stage larvae in mature
ewes.
Lambs 7, 8 and 18 were born in captivity from Pikes Peak ewes at the Bird Farm.
Nume~ous eggs and first-stage larvae were collected from 7 and 8, while many
fourth stage were found in lamb 18, indicating in all three that heavy transplacental transmission had occurred.
Data from lambs collected at Pikes Peak once again reveals an extremely heavy
burden of lungworm (see below). Lamb 11 was quite young when collected and
thus has few lungworm eggs and larvae, but many third and fourth stages. After
that collection, the larval output per lamb increased to very high levels,
indicating heavy loads of adult parasites.
Lamb 15 is an exception, yet the
gross and histopathology picture indicates he was as heavily infected as the
other lambs; possibly a mistake was made in the count.

�-93-

Lamb Number
*73BHL- 7
*73BHL- 8
73BHL-ll
73BHL-12
73BHL-13
73BHL-14
73BHL-15
73BHL-16
73BHL-17
*73BHL-18
73BHL-19

Larvae per Lamb
1,286,900 first-stage
5,774,710 first-stage
Many 3rd, 4th and 3,947 first
507,020 first-stage
259,490 first-stag~
2,279,800 first-stage
75,640 first-stage
470,625 first-stage
898,650 first-stage
Many third and fourth stages
960,412 (partially digested)

*Born of captive bighorn sheep from Pikes Peak and all (except
18) died of verminous pneumonia in captivity.
Lamb 18 was not
claimed by the ewe and starved. These animals (7, 8, 18) were
not reported in 1972-73 report. For lambs in the series prior
to BHL-7, check last year's annual report; all of these lambs
were from Pikes Peak ewes and were either still-births or died
from other neonatal problems. All were heavily infected with
transplacentally transmitted lungworm.

SOMATIC STORAGE OF PROTOSTRGNGYLUS

IN BIGHORN SHEEP

INTRODUCTION
Lambs born on Pikes Peak are heaVily infected with lungworm as a result of
transplacental transmission during the 'last trimester of pregnancy.
The
only method by which such a phenomenon could occur is for the ewe to become
infected by eating snails containing third-stage larvae. She must then store
these larvae somewhere in he r body (somatic storage) until late pregnancy.
Snails are not available in the winter months; therefore, ewes must eat infectea snails in spring, sunnner and/or fall and store the larvae.
There are two reasons for seeking the site of somatlc storage in the ewe.
From an applied standpoint, antihelminthics can be more accurately evaluated
if the site is known. From a basic standpoint, the site of storage may offer
some clues regarding innnunology to the parasite.
METHODS AND MATERIALS
Various tissues from each ewe necropsied in late August, 1973, through March,
1974, were ground or minced and placed in 1 gallon jars containing water for
24 hours to allow any larvae present to exit. The ground tissue was removed
with a strainer and then washed several times to remove any larvae remaining
on tissue surfaces. This tissue was held for digestion.
The washings and
original soaking fluid were allowed to sit for 30 minutes to permit any larvae
to settle to the bottom. Excess fluid was then decanted.
Sediment and remaining fluid were examined for larvae. The tissues were then placed in 1

�-94-

gallon jars and Pepsin-HCI solution added to digest tissues, allowing any
larvae still trapped within to be released.
After 24 hours sufficient
formalin was added to create a 10 percent formalin solution and stop
digestion, preserving any larvae present.
Again, the decanting procedure
was followed.
Since the sediment remaining was very thick, anhydrous ether
was added to 20 percent total voltune, to float the di.gested organic matter,
leaving ltmgworm larvae in the aqueous layer below.
The organic layer was
removed by aspiration with a pipette and rubber bulb. The aqueous layer
was examined for presence of larvae.
Specific tissues examined for larvae
were as follows:
bone and bone marrow, muscle of the front and hind
quarters, facial muscle, cervical muscle, muscle of the back, lOins, limbs
and thoracic area, abdominal muscle diaphragm, soft and hard palates of
the mouth, tongue, gums, salivary glands, trachea, pharynx, nasal turbinates
and sinus scrapings, lungs, esophagus, rtunen, reticultun, omastun, ab omas tun,
small and large intestines, pancreas, liver, gall bladc'.er,spleen, perit oneum , omen tum , thy.:-oid,thymus, kidneys, ureters, urinary bladder, adr-ena.l s ,
ovaries, uterus, vaginal tissue, mammaries, body fat, beart, pericarditun,
major arteries and veins, lymph nodes, brain, spinal cord, eyes and integtunent.

RESULTS
Case #1:

In late August, 1973, a ewe trapped on Pikes Peak the previous
winter, and held in captivity at the Colorado Division of Wildlife Research Station at Fort Collins, died of acute Pasteurellosis.
One Protostrongylus sp. L3 was found in lung tissue and all other
tissues were negative.

Case #2:

In October 1973, a ewe trapped in Pikes Peak the previous winter,
and held in c3.ptivity at the Colorado Division of Wildlife Research
Station at Fort Collins, died of acutePasteule1losis.
One Protostrongylus sp. L3 was found in lung tissue and all other
tissues examined were negative.

Case #3:

In early November, 1973, a ewe trapped on Pike~ Peak the previous
winter and held in captivity at the Colorado Division of Wildlife
Research Station at Fort Collins, died of acute Pasteurellosis.
All tissues examined were negative

Case #4:

for Protostrongylus

sp. L •
3
In November, 1973, a ewe trapped on Pikes Peak the previous winter
and held in captivity at the Colorado Division of Wildlife Research
Station at Fort Collins died of acute Pasteurellosis.

Two Protostrongylus Bp. L3 were found in lung tissue and all other
tissues examined were negative.
Case #5:

In November, 1973, a ewe trapped on Pikes Peak the previous winter
and held in captivity at the Colorado Division of Wildlife Research
Station at Fort Collins, died of acute Pasteurellosis.
Three Protostrongylus sp. L3 were found in lung tissue and all
other tissues examined were negative.

�-95-

Case #6:

In early February, 1974, a ewe that was a trap mortality on
Pikes Peak was examined.
She was 2-4 weeks pregnant.
Her
lung tissue showed 124 Protostrongylus Spa L •
All
other
3
tissues including the fetus were negative.

Case #7:

In earlJ February, 1974, a ewe that was a trap mortality on
Pikes Peak was e~~ined
and showed 55 Protostrongylus Spa L3
in lung tissue. All other tissues examined were negative.
This ewe was not pregnan t ,

Case #8:

In early February, 1974, a ewe recently trapped on Pikes Peak
died of ventricular fibrillation while being unloaded at the
Wild Animal Disease Center Pens at Colorado State University
at Fort Collins.
She was 8-10 weeks pregnant.
Her lung tissue ,;howed 124 Protostrongylus Spa L3'
All
other tissues examined, including the fetus were negative.

Case #9:

In early March, 1974, a ewe that was a trap mortality on Pikes
Peak was examined.
She was 12-16 weeks pregn~nt and showed 34
Protostrongylu~ Spa L3 in lung tissue and three Protostrongylus
Spa L3 in the cotyledous.
All other tissues examined including
the fetus were negative.

Case #10: In mid-March, 197~, a ewe at the Wild Animal Disease Center Pens
at Colorado State University died as a result of injuries suffered
when previously trapped on Pikes Peak. She had been treated with
Tramisol at the time of her capture.
Her fetus was aborted before she died and was negative for Protostrongylus Spa L3•
He~ lung tissue showed 20 Protostrongylus Spa
L3 and all other tissues examined were negative.
Case #11: In mid-March, 1974, a ewe trapped on Pikes Peak aborted her fetus
some 3-5 days after being released into penn at the Wild Animal
Disease Center facility at Colorado State University.
The aborted
fetus was found the morning of the ewe's death.
This fetus was
about 12-16 weeks old and was negative for Protostrongylus Spa L3 .
The ewe's lung tissue showed 26 Protostrongylus Spa L3 and the
cotyledons showed 3 Protostrongylus Spa L3.
All other tissues
examined were negative.
Case #12: In late March, 1974, a ewe trapped on Pikes Peak and released into
pens at the Wild Animal Disease Center at Colorado State University
began to abort her fetus. Her condition deteriorated and she died.
She had been treated with Task at the time of her capture.
Her
lung tissue and cotyledons showed 20 and 4 Protostrongylus Spa L3 ,
respectively.
All other tissues examined

including

the fetus, were negative.

�-96-

Case #13: In late March, 1974, a ewe trapped on ?ikes Peak bloated and
died while still in the net. Her lung tissue contained 23
Protostrongylus sp. L3 and the fetal liver contained 6 Protostrongylus sp. L3 •
All other tissues examined were negative.

DISCUSSION

OF RESULTS

The data observed on the ewes which died in late 1973 of acute Pasteurellosis
at the Colorado Division of Wildlife's Research ftation at Fort Collins,
showed lung tissue to be a storage site of ProtostrongYlus sp. L3 •
Ewes examined through the middle of February, 1974, showed relatively high
numbers of Protostrongylus sp. L3 within their lung tissue at a time when
exposure to snails containing these larvae was greatly reduced due to winter
climatic conditions on Pikes Peak. Ewes examined after early March, 1974,
showed relatively low numbers of Protostrongylus sp. L3 in lung tissue but
L3 larvae began to appear in their reproductive tissues and eventually the
liver of the fetus.

CONCLUSIONS
These data to suggest that Protostrongylus sp. L3 migrate to the lungs of
ewes but do not develop to mature adults; rather they remain dormant until
some mechanism triggers them to move (by an unknown route) to transplacentally
infect the fetus.
Obviously, much more research is needed to ver:l.fythese phenomena and to show
the route that L3 take from storage si~e(s) to the fetus. Hopefully, this
may be accompli~hed in 1974-75.

�-97-

JOB PROGRESS REPORT

State of

Colorado

Project No.

W-124-R-2

Raptor Investigations

Work Plan No. __~l~
Job Title:

Job No. 1

Statewide Raptor Populations and Characteristics Studies

Period Covered:
Personnel:

_

February 1, 1973 through January 31, 1974

Gerald R. Craig

ABS~RACT
One hundred and thirty-nine golden eagle nests were surveyed .for activity
and productivity in 1973. An average of 1.13 young golden eagles per active
site were estimated to have fledged.

Productivity of four osprey nests was

reduced to two young fledged from one site.

Twenty-three peregrine sites were

surveyed and 11 pairs were located which r~presented a 50% decline in known
breeding pairs.
young.

Productivity was poor with the 11 sites producing only two

Data for eggshell thinning and pesticide content is given.

Sixty-one

prairie ~alcon sites succeeded in producing an estimated 94 fledglings (an
average of 2.84 young fledged per productive site).

Wintering bald and golden

eagle trend information was obtained through aerial cen~uses of northeastern
Colorado.

An experimental aerial eagle census of southeastern Colorado did

not produce sufficient results to justify adoption.

Statewide eagle concentra-

tion counts were made in February and March by field personnel.

The statewide

concentration counts also yielded information about habitat preferences and
activity of eagles.

Mortality factors were obtained from reports of injured

and dead raptors turned into the Division and its cooperators.
and one technical articles were written.

Two popular

��-99-

STATEWIDE RAPTOR POPUlATIONS AND CHARACTERISTICS

STUDIES

Gerald R. Craig

To investigate

status of current populations,

including distribution,

characteristics

migration, patterns and chronology

and trends

of Bald and Golden

Eagles, Osprey, Peregrine and Prairie Falcons and Burrowing Owls in or through
Colorado.
SEGMENT OBJECTIVES
1.

Develop and refine methods for determining breeding numbers of Bald and
Golden Eagles, Osprey, Peregrine and Prairie Falcons and Burrowing Owls
in Colorado.

2.

Develop and refine methods for determining winter trend information

for

Bald and Golden Eagles, Peregrine and Prairie Falcons and Burrowing Owls
in Colorado.
3.

Locate and categorize major and important habitats and associated Bald
and Golden Eagle, Osprey, Peregrine and Prairie Falcon and Burrowing Owl
populations

4.

in Colorado.

Establish and maintain annual or periodic inventories

of Bald and Golden

Eagles, Osprey, Peregrine and Prairie 'Falcons and Burrowing Owls in
Colorado from inventory methods selected from Project Statement Procedures
1 and 2.
5.

Monitor mortalities and inspect injured and dead raptors reported or
brought to the Division to determine mortality

factors of Bald and Golden

Eagles, Osprey, Peregrine and Prairie Falcons and Burrowing Owls and other
species that are obtained incidentally.
6.

Initiate and maintain a IO-year raptor banding program to determine migration, chronology, movement, life tables and mortalities

of Bald and Golden

Eagles, Osprey, Peregrine and Prairie Falcons and Burrowing Owls and other
species that are encountered accidentally.
7.

Prepare a series of comprehensive

information leaflets and recreational

guides at intervals when the information becomes available.

�-100-

METHODS AND MATERIALS
Breeding

Inventory

Bald Eagles
As confirmed
nesting

nesting bald eagle observations

investigations

and little actual
Golden

are nonexistent

have been limited to inquiries

of any nesting

attempts

field work.

Eagles

The nesting
gressed

survey of golden eagles commenced

through June 30.

Two temporary

in March,

employees

to time limitations

and existence

of extensive

drive,

trail bike or on foot.

and 15 to 60x variable

spotting

nest sites, the attached
ability

mapping

data sheets

of the observers.

investigators

active and inactive

populations

were made using binoculars

Upon discovery

of active or inactive

(Appendix A) were completed

While

and Rio Blanco Counties

early in the nesting

nature,

in northwestern

productivity,

Where

such observations

the nest, thus many nestlings

entering

dense nesting

sites which were determined

season in order to establish
feasible,

the

Colorado.

prey remains and band nestlings.
without

the

amount of their time to

to the particularly

An attempt was made to revisit more accessible
to be active

to the best

some time was spent throughout

of the effort was confined

in Moffat

investigations

Observations

to devote the greatest

sites.

Due

several hours either

As the survey was of an extensive

were encouraged

state, the majority

scopes.

the period.

nesting habitat,

limited to those sites that could be reached within

by 4-wheel

1973, and pro-

equipped with a 4-wheel

drive vehicle and trail bikes were in the field throughout

were

in Colorado

efforts were made to make
were not

banded.
Peregrine

Falcon

The peregrine
continued

through July 31, 1973.

cooperation
falcons.

falcon nesting

survey was initiated March
The primary

with James H. Enderson,

All known historic

surveyed

eyrie sites

investigator

1, 1973 and

(G. R. Craig),

in

the state for nesting peregrine

(a total of 23) were visited by the

investigators

and each site including

adjacent

The procedure

established

in determining

vantage points

overlooking

known or suspected

suitable habitat,

activity

was to position

was surveyed.
oneself at

eyrie sites and keep the area

�-101-

under observation with binoculars and spotting scopes for six to eight hours
minimum if falcons were not seen prior to the cutoff time.

The activity of

the falcons, extensive and often convoluted nature of the terrain, great distance of vantage points from nest sites (often in excess of ~ mile) and
observer fatigue made the presence of two observers almost mandatory.

Sub-

stanciation of presence of nesting falcons was often facilitated by inspecting
nesting areas for excrement on potential roost sites and noting activity of
other raptors and prey species.
Upon confirmation of presence of peregrines,
periodically

the site was revisited

to establish nesting activity, initiation and completion of egg laying

behavior of the pair.

Hatching time was estimated for those pairs apparently

incubating eggs and the nest site was climbed into to collect unhatched eggs,
shell fragments and prey remains.

Duration of visits was limited to less than

30 minutes at the nest cliff and only several minutes at the nest site.
were also visited that were occupied by unsuccessful

Sites

falcons to determine the

extent of each nesting attempt, collect infertile eggs and shell fragments, and
collect prey remains.
Sites occupied by successful pairs were again revisited at time of fledging
and the number of fledglings was determined from a distant vantage point.

Thus,

it was necessary to actually visit the nest site and disturb the falcons only
one time.
Prairie Falcons
Nesting investigations of prairie falcons were carried out concurrently
with studies of golden eagles and peregrines.

Intensive field work on prairie

falcons was associated with Work Plan I, Job No. 2 of W-l24-R-2 and is described
in the Job Progress Report for that study.
Historic prairie falcon eyr.ies along the Front Range of Colorado were
visited at time of fledging to determine productivity.

Nest cliffs were

examined with spotting scope and binoculars to locate active nest ledges and
determine presence of tLe adults.

Whether or not falcons were sighted, the

investigator climbed to the base of the cliff and then worked his way to the
cliff top.

Invariably, active sites would be disclosed when an adult would

fly by to protest the investigator's presence.

If time permitted, the inves-

tigator would then rappel into the eyrie to establish brood Size, band the
young and identify prey remains.

Nest site information, surrounding habitat

�-102-

and disturbance
Appendix A.

factors were recorded on forms similar to that described in

Osprey
Nesting investigations
through July 31, 1973.
productivity,

of osprey commenced early in June and continued

In mid-July, known nest sites were visited to ascertain

band nestlings, collect eggshell fragments and record data on the

nest sites, habitat and disturbance factors.

(Preliminary nest locations and

determination

of activity in North Park, Colorado was accomplished by Wildlife

Conservation

Officer Steven Porter).

Due to low nesting densities and the

nesting habitat utilized by Colorado osprey, nest searches designed specifically
for osprey are not feasible, the most effective method appears to be solicitation
of observation from field personnel who are likely to encounter the birds in
the course of their work.
Burrowing Owls
Burrowing owl nesting investigations were restricted to recording locations
of burrowing owl colonies reported by field personnel and the public.
WINTER INVENTORY
a)

Aerial Golden Eagle Census in NE Colorado
The same procedure established in 1972 was followed.

Briefly, an area of

3,000 square miles was des Lgna t ad in northeastern Colorado.

The area is bounded

on the west by U. S. 85 from Greeley to th~ Colorado-Wyoming

border; the north

boundary is the eastern boundary is Highway 71 south to Colorado 14, thence south
to Interstate 80-S; the southern boundary is Colorado 36 from I-80-S to Greeley.
Thirteen randomly selected south transects represent a total of 600 linear miles.
The tra~sects are flown with a Cessna 185 at an altitude of 100 to 200 feet with
speeds averaging

100 to 120 miles per hour, and all raptors within ~ mile of

either side of the aircraft are counted.
for the entire 3,000 square mile area.
b)

The results obtained are then extrapolated

Aerial Golden Eagle Census in SE Colorado
A 10,000 square mile area in southeastern was delineated for an aerial census

similar to that described for the northeastern Colorado cenSUG.

The area is

bounded on the west by Interstate 25 from the New Mexico border to Pueblo; the
northern boundary approximates

the Arkansas River from Pueble east to the Kansas

border; the eastern border is the Coljrado-Kansas
is the Colorado-Oklahoma

border; and the southern border

and Colorado-New Mexico borders.

Twenty randomly selected

�-103-

north-south transects represent a total of approximately

1,000 linear miles.

The

transects are flown in the same manner described for the northeastern aerial
census.

The actual coverage of 500 square miles (~ mile on both sides of the

aircraft for 1,000 miles) represents a 10% sample of the study area.
c)

Aerial Bald Eagle Census of South Platte River from Greeley to Fort Morgan
Upon completion of the northeastern Colorado aerial census, a flight was

made of the South Platte River from Fort Morgan to Greeley.
identical from one year to the next.

The procedure is

The aircraft is flown above the river at

altitudes of 150 to 300 feet and at speeds of 100 to 120 miles per hour.

All

eagles observed on either side of the aircraft are classified as to species and
age.
d)

Aerial Bald Eagle Census of Arkansas River from Pueblo to Colorado-Kansas
Border
Upon completion of the experimental Southeastern Colorado aerial flight,

the aircraft was flown along the course of the Arkansas. River and an attempt was
made to count eagles present along the river.

The same technique was used as

described in the South Platte bald eagle census.
e)

Statewide W.C.O. Eagle Concentration Counts
For the second year, all Wildlife Conservation Officers were requested to

record all eagles observed in their districts during the periods of February 3-9
and March 3-9.

The officers were instructed to survey those areas where greatest

concentrations of eagles were likely to occur.
LOCATION OF IMPORTANT HABITATS
Data as to land use, begitative type, disturbance factors were recorded on
nest report forms (Appendix A) which were completed for each raptor nest that
was investigated.

In addf.t Lor, , the major vegitation types in which raptors were

observed were recorded on migration and winter raptor population censuses.
MORTALITY
Reports of dead raptors were obtained from field personnel, where possible
they attempted to determine and report the cause of mort~lity.

When feasible,

carcasses were brought into the investigator for examination to ascertain cause
of mortality.
A variety of injured raptors were also brought into several veterinarians
authorized to treat and rehabilitate wildlife.

Species, sex, age, cause of injury

�-104-

location of discovery, party turning the raptor in and other pertinent data were
recorded by the veterinarians
poses of computation,

and turned into the Division of Wildlife.

such incapacitated

since they were incapacitated

For pur-

raptors were considered as mortalities,

sufficiently to permit their capture.
BANDING

Nestlings were banded when nest sites were visited to obtain productivity
and preY,data.

In regard to E:agles, sites were not visited for banding purposes

if the 'eaglets were within two weeks of fledging.
caus.tng them to fledge prematurely.
I

t Icn

This precaution was to avoid

Raptors were also trapped during fall migra-

at several localities where trapping stations were erected.

Further, inca-

pacitated raptors that had beE'.ll
rehabilitated were banded upon release to monitor
/the success of rehabilitation.
./

/

RESULTS AND DISCUSSION
Breeding Inventory
Bald Eagles
One probable bald eagle nest was located on the Colorado River west of
Rifle, Colorado.

An interview with the landowner revea Led that the site had

been occupied by a pair of bald eagles in 1968, 1969 and 1970.
been successful in rearing several young in that pe,iod.

The pair had

As the region is

frequented by wintering bald eagles, the stockman was acquainted with the
difference between bald eagles and golden eagles, thus the chances of minidentification are remote.

He has made special note that the pair had white heads

and tails indicative of adult plumage.

The site was vinited on several occasions

in the &amp;pring and summer of 1913, and while bald eagles were observed along
the river early in the spring, no eagles were observed to be attached to the
site.

The nest is locat~d in a large cottonwood tree on an island in the

center of the Colorado River.

It is quite large and bulky and unlike the

average golden eagle nest, appears to be construc.ted almost entirely of large
sticks which are uniform in diameter.
inspected by authorities

Photographs of the nest have been

experienced with bald eagle nests and most have observed

that it is typical of nests constructed by bald eagles.
Golden Eagles
A total of 139 golden eagle nest sites were surveyed for activity in 1973.
Fifty-two percent of the sites were located in northeastern Colorado

{Rio Blanco

�-105-

and Moffat Counties).

As many sites as possible WEre located early in the nesting

season and an effort was made to return to the sites prior to fledging to delineate
productivity.

Unfortunately,

this could not be accomplished

in all cases and

fledging success was estimated by visiting the sites after fledging and inspecting
for prey remains and ~ther signs of the former presence of young.
Seventy-nine percent of the sites were active (activity was defined as a
site with fresh decorations or an incubating adult),

A total of 86 young were

estimated to have been produced by 68 sites (54 young were observed in 36 nests),
which averages 1.26 young per site (an average of 1.50 young per site were
observed).

Brood sizes of 1 were estimated for those sites for which no early

productivity data was obtained.

Sixty-six young were estimated to have fledged

from 59 sites (59 young were observed to have fledged from 52 sites), which
averages 1.11 young fledged per site (an average of 1.13 young per site were
observed to have fledged).
Osprey
Little is known of osprey nesting activities in Colorado.

In the past,

only two sites were known, one on Grand Lake in Grand County and the other
on Electra Lake in LaPlata County.

Both sites have been active at least from

the mid-1960's and on several occasions, at least two yo~ng were fledged from
both sites.

The Electra Lake nest supposedly has fledged at least 2 young

each year.
While sightings of cspr ey are fairly common during migration, summer
sightings are few.

Jim Webb of the U. S. Forest Service, stationed at Steam-

boat Springs provided a number of such summer observations in Routt and Jackson
Counties.

John Ellenberger, Wildlife Conservation Officer in North Park

reported one osprey nest recently discovered by Forest Service personnel, as
well as a second site he had located.

Both sites were visited by a second

Wildlife Conservation Officer, Steve Porter, in the spring of 1973 and were
determined to be active as adults were present on the nests during incubation.
Later, when young should have hatched, the trees were climbed to determine
productivity.

Neither site produced young, nor could eggshcll fragments be

discovered.
Table 3 provides productivity

figures for the four known osprey nests.

A pair of adults were present early in incubation at the Grand Lake site,
but abandoned nesting activities.
cessful in producing young in 1973.

Thus, only the Electra Lake pair were suc-

�-106-

Peregrine Falcons
In cooperation with J. H. Enderson the history of occupancy of historic
peregrine

eyries was compiled which is summarized in Table 4 .

All historic peregrine
total of 23 sites surveyed,

falcon eyrie sites were visited in 1973 and a
11 were found to have pairs present with another

site occupied by a lone male.

This represents a 50% decline in number of

known breeding pairs in Colorado frcm 1964 to 1973.

In addition to investigating

known eyries sites, efforts were expended to survey suitable habitats for new
eyrie sites.

A total of 19 such sites were visited which contained ideal nest

cliffs and hunting areas, but for whi.ch no peregrine sightings had been reported.
One new pair was located with a possible second nesting pair (both falcons were
observed early in the nesting season, but later searches failed to reveal the
pair) and a possible adult at another.

The terrain did not permit confirmation

of the observation and the falcon was not sighted again.
Total productivity
site fledging 2 young.

for the 11 pairs of peregrines was meager with only one
By making visits to sites and observing pairs through

spotting scopes, it was determined that at least 6, and possibly 7, pairs were
incubating

eggs (eyries 1, 2, 5, 6, 7, 8 and possibly 9).

At least 2 pairs

(eyries 2 and 6) failed during incubation of the first clutch and renested on
another ledge.

Nesting faflure eyrie 1 may be attributed

to disturbance by

rock climbers, but failure of other sites. could not be attributed
disturbance.In

to human

fact, the single pair that succeeded in fledging young received

a great deal of indirect human harrassment

by rock climbers.

Two infertile and one E.bandoned egg were each collected from different
eyries (eyries 1, 2 and 8) a.nd eggshell fragments were found at another eyrie
(No.6).

The thickness index (described by Ratcliffe

with a micrometer,

1967), thickness measured

and eggshell weight were compared with similar data for

eggshells of the subspecies available in the literature

(see Table 6).

Egg-

shell thinning on the order of 20% of pre-1947 eggshells is evident which is
about that averaged by the eastern population of peregrines as it was declining
to extinction.
Contents of the three intact eggs were analyzed for chloronated hydrocarbon residues by the Denver Wildlife Research Center.

DDE residues accounted

for 90% of the pesticide residues and the levels were found to be similar to
those found in interior Alaskan and Canadian peregrine populations.

�-107-

Among prey taken by peregrines
(Aeronautes saxatalis),

at eyries were White-throated

swifts,

rock dove, (Columba livia), Brewer's blackbirds,

(Euphagus cyanocephalus),

Clark's nutcracker,

jays (Gymnorhinus cyanocephala),

(Nucitraga columbiana),

pinon

and mourning doves (Zenaidura macrura).

Prairie Falcons
A total of 61 prairie falcon nesting
in J.973. The majority

of the sites were located east of the Continental

The number of sites surveyed is probably
population,

but time limitations

known sites.

locations were surveyed for activity

less than one-sixth of the entire

did not permit detailed investigations

Primary purpose of the visits was to establish presence

and secondarily
Fifty-four

Divide.

of all
of falcons

to estimate productivity.
of the 61 sites were determined

to be active

(an active site was

defined as one having at least an adult male or female attached

to a nest cliff).

A total of 85 young were known to have been produced by 37 sites (average of
2.3 young per site).

However, the actual number of young produced by 14 of the

37 sites was not established,

thus actual production

of 23 sites is 3.7 young

per eyrie which is probably more indicative of productions

of tile population.

Known fledging success could be determined

from 22 sites '~ich fledged 79 young

(average of 3.69 young per site).

success of 11 additional

Fledging

not available due either to lack of observation
true of 6 sites), or disappearance
young but for which production

at time of fledging

information

is lacking (5 sites).

lings for each of the 11 sites, thus yielding an adjusted
The adjusted

(this was

of young from sites that obviously produced

fledging success may be adjusted by assuming average production

2.84.

sites was

Thus, known
of 3.69 nest-

fledging success of

fledging success of 2.84 per eyrie is 0.85 o~ a bird less

than the estimated fledging success of 3.69 for all 33 sites.
from 5 and possibly 6 sites between hatching and fledging.
persons may be a factor in at l~ast four and possibly

Broods disappeared

Removal of young by

six cases.

Production

of prairie falcons in 1973 appeared to be good with little nat-

ural mortality

occuring between hatching and fledging in eyries that were remote

from human disturbance.

�-108-

Burrowing

Owls

As is common of the species, burrowing
prairie dog (Cynomy~~)

colonies.

owls were commonly associated with

Burrowing owl populations

appear to be

stable with owls being abundant in some localities and nonexistent
despite apparently

ideal conditions.

in others

Sightings were made throughout

the

eastern plains region of the state and were generally associated with blacktailed prairie dog (Cynomys ludovic~)
8 to 10 miles north of Fort Collins
Broomfield
Animas

towns.

Concentrations

(Larimer Co.), east of Timnath

(Boulder Co.), and the Comanche National Grasslands

counties).

with white-tailed

(Larimer Co.),

(Otero and Las

Owls were also observed along the Colorado

Valley (Garfield Co.) in association
leucurus).

were reported

River near Grand

prairie dogs (Cynomys

WINTER INVENTORY
a)

Aerial Golden Eagle Census in NE Colorado
Due to difficulty

peak concentration

in scheduling an aircraft more than one time during the

months of December, January and February,

were reduced to one in mid-.January.
establish

As the primary purpose of the count is to

trend data from year to year to determine possible

the reduction

the number of flights

of flights to one should not alter the results.

long range fluctuations,
Table 10 summarizes

the results of the 1972-73 winter flights and the January 1974 flight.
b)

Aerial Golden Eagle Cens~s in SE Colorado
The flight was initiateo January 23, 1974, but due to the length of time

required' to complete the census, an additional
flight.

Unfortunately,

day was required to complete

pressing ~atters required delaying completion

one day until January 25, 1974.

The investigators

number of eagles observed despite ideal habitat.
an extrapolated

of the flight

at the small

Only 10 eagles were observed for

number of 100 eagles in the entire 10,000 square mile study area.

This was only one eagle per 100 square miles.
decided that this flight would discontinued
c)

were su~rised

the

Due to the poor results, it was

in the future.

Aerial Bald Eagle Census of South Platte River from Greeley to Fort Morgan
As the flight is integral with the northeastern

Colorado aerial census, the

number of flights were reduced from three of the previous winter to one only in
January.

This flight was made January 16, 1974, counting ccnditions were ideal

�-109-

and it was estimated

that the census of the riverbottom

of the total population.

The irmnature population

was accurate

increased

within 5%

slightly

from 42%

in 1973 to 48% in 1974 (see Table 11).
d)

Aerial

Bald Eagle, Census of Arkansas

River from Pueble to the Colorado-Kansas

Border
The count of the Arkansas
tion of the southeastern

River was made on January

Colorado

flight.

only three (2 adults and 1 immature)
eagle and one unknown
made to establish
e)

Statewide

The results were disappointing

bald eagle observations,

eagle sighting.

Consequently,

W.C.O. Eagle Concentration
50% of the Wildlife

Counts
Conservation

Officers

forms, the results of which are presented

(61 bald eagles and 157 golden eagles).

269 eagles were observed,
gathered

one adult golden

no future plans have been

responded

in Table 12,

ruary 3-9 count period, a total of 225 eagles were observed,
eagles

observed

by returning

In the Feb-

of which 27% were bald

In the March 3-9 count period,

of which 17% were bald eagles.

As compared

in 1972, Table 12, the total number of eagles observed

period decreased

with

trend data in this region.

Approximately
observation

25, 1974, upon comple-

with data

in the February

51% (466 in 1972 to 225 in 1973) and the total nUmber of eagles

in the March period declined

19% (333 in 1972 to 269 in 1973).

When

the average number of eagles sighted per observer are compared

for 1972 and 1973

(Table 12) the cause for the decline in total eagles obseyved

is apparently

to a decrease

in the wintering

The observation

bald eagle population.

forms also provided

for information

and habitats

over which they were Sighted.

feeding,upon

carrion.

Road-killed

eagles were observed

The activity

Trees were preferred

in trees), 870were perched

of eagles

rabbits were fed
In 34% of the

sharing kills with other eagles.
(87 were bald eagles and 337 were

28 were unidentified

to bald eagles, 34% were observed
were perched.

and unidentified

for 27% and elk 5%.

of 452 eagles were recorded

golden eagles, the remaining

as to activity

Table 14 details of eagles observed

jackrabbits

upon by 47% of the eagles, deer accounted
observations

due

as to species).

With regard

flying, 10% were feeding upon carrion,
perch sites of bald eagles

on ice and 6% were perched

and 55%

(79% were perched

on the ground.

Of golden

eagles, 37% were seen flying, 9% were feeding upon carrion and 52% were perched.
Thirty-five
ground,

percent

of the golden eagles were perched

on power poles, 26% on the

11% in trees, 10% were on fence posts and 8% were perched

croppings.

on rock out-

�-110-

LOCATION OF IMPORTANT HAB:::TATS
a)

Habitat Preferences

of Wintering

Data recorded by Wildlife
is summarized
accounted

Conservation

were observed in that vegetative
in riverbottom

that type of habitat.
grassland

Officers on eagle concentration

in regard to habitat preferences

for 37% of the sightings

were observed

Eagles

in Table 16.

counts

Pure sage stands

of golden eagles while Wo of the bald eagles

type.

Thirty-four

percent of the bald eagles

habitats while only 4% of the golden eagles were in

Fourteen percent of the golden eagles were located in

situations and p% of the bald eagles were in similar areas.
MORTALITY

Of 187 dead and injured raptors reported in Colorado
December

from January 1 through

31, 1973, 22% were caused by collision with objects, 20% were caused by

gunshot and 17% were caused by vehicle strikes

(Table 17).

Golden eagles accounted

for lWo, red-tailed hawks for 15% and great horned owls for 14% and American

kes-

trels for 12% of the dead and injured raptors.
Figure 1
Breakdown by Month of Dead or Injured Raptors Reported to the Division
Number
Raptors
30
28
26
24
22
20
18
16
14

12
10
8

6
4
2

Jan.

Feb.

Mar.

Apr.

May

Jun.

Jul.

Aug.

Sep.

Oct.

- 1973

Nov.

Dec.

�-111-

The graph depicts seasons when injured or dead raptors were reported.

The

graph follows a trend similar to 1972 in that there is a general decline in
mortalities

from January through May when the raptors have already undergone

great mortality
unavailable

of f~ll migration

in nesting

and winter.

Raptors,

season and are relatively

November

when migrating

trusting,

encounter humans.

and man.

adults, are

secure from mortality.

and injuries climb sharply as fled lings first encounter
and quickly succumb to the environment

especially

Death

the world on their own

The next peak occurs during

immature raptors from the north, which are relatively

BANDING
Table 19 lists the species and numbers of raptors banded in Colorado
the Division

of Wildlife

from January through December,

ARTICLES.

INFORMATION

the bi-monthly

publication

of the Division

Hawks and Falcons of Colorado"
the March-April
peregrines

1973 issue.

1973.

for Colorado Outdoors,

of Wildlife,

and "Colorado's

entitled

"The Eagles.

Raptor Prog'Iam", published

A technical article on the status of nesting

was co=aut ho re.I with Dr. James H. Enderson and published

October 1974 issue of Auk, Vol. 91, p. 727-736.
Prepared

by,

C. IZ.

e.a.:,

Gerald R. Craig

by

GUIDES AND LEAFLETS

Two popular articles about raptors were written

the

in the

in

�-112TAB1E 1

Activity

and Productivity

of Colorado Golden Ellg1es

--

Symbols:

A = active site; IA = inactive site;

Site
Number

Activity

CH-2

A

CH-3

A

FA-2

IA

FA-4

A

FA-5

A

FA-8

IA

EP-3

A

GA.-3

A

Adult incubating

GA.-8

A

Adult incubating

GR-1

A

GR-9

IA

HU-3

A

HU-8

A

1S-1

Number
of young

Number
Fledged

2

2

= no ohservations

Comments

Adult brooding

1

0

1 wk chick dead below nest

1+

1 chick 1 wk. of age

1+

Adult brooding

A

1+

4-5 wks. of age

1S-2

A

0

0

1S-5

A

1

1

ME-5

A

1

1

ME-6

IA

ME-7

A

ME-8

IA

MI-l

A

1+

MF-l

A

2

2

MF-2

A

1+

1

MF-5

A

0

0

MF-6

A

2

1

MF-8

A

2

1

MF-ll

A

0

0

MF-13

A

1+

1

MF-15

A

1+

1

MF-18

A

1+

1+

1 spoiled egg

Adult incubating

Abandoned

No eggs laid

�Activity
Table 1

and Productivity

-113of Colorado Golden Eagles

page 2

Site
Number
Number
_N~u=m~b~e~r
__ ~A~c~t~i~v~i~t~y
o~f~y~o~u_nag
__ ~F~l~ed~g~e~d~
__ ~C~omm==e~n~t~s~
MF-19

A

1

1

MF-2l

A

1

1

MF-22

A

1

1

MF-23

A

2

2

MF-24

A

2

2

MF-25

A

o

o

MF-26

A

1

1

MF-27

A

MF-28

A

1+

1

MF-29

A

2

1

MF-3l

A

1

1

MF-33

A

1+

1

MF-35

A

MF-36

IA

MF-37

A

1+

1

MF-38

A

o

o

MF-39

A

2

2

Adult incubating

5 wk. chick dead below nest

Adult incubating

2 spoiled eggs

MF-40
MF-4l

A

MF-42

IA

MF-43

A

1+

1+

MF-44

A

1+

o

MF-46

A

1+

1

MF-47

A

o

o

MF-48

IA

MF-49

A

MF-SO

A

l-1F-Sl
MF-S2

Adult incubating

MF-45

Nest filled wlsand
Pair shot in 1972

o

o

A

1+

1+

A

1+

1

MF-S4

A

1+

1

MF-SS

IA

MF-56

A

1+

1

MF-S3

Abandoned

- human disturbance

_

�-114Activity
Table 1

2',ndProducti vi ty of Colorado

Golden Eagles

page 3

Site
Number

ACtiTity

Number
of young

Number
Fledged

MF-57

A

0

0

MF-58

IA

pu-o

A

RI-l

A

0

0

1 spoiled egg

RI-3

A

2

1

Dead chick below nest

RI-6

A

1+

1

RI-7

A

1+

1+

RI-lO

A

RI-12

A

1

1

RI-13

A

1

1

1 spoiled egg

RI-14

A

0

0

2 spoiled eggs

RI-15

A

0

0

Abandoned

RI-16

IA

RI-17

A

RI-l8

IA

RI-19

A

RI-20

IA

RI-2l

IA

RI-22

A

RI-23

IA

RI-24

A

RI-25

incubating

1+

1

0

0

A

0

0

RI-28

A

2

RI-29

A

0

0

RI-30

A

1+

1

RI-3l

A

1+

1

RI-32

A

2

2

RI-33

A

2

2

RI-34

--

1+

1+

RI-26
RI-27

RI-35
RI-37
A

2 spoiled eggs

Adults

1+

RI-38

Comments

1 spof.Led

egg

�-115Activity
Table 1

and Productivity

of Colorado Golden Eagles

Site
Number

Activity

Number
of young

Number
Fledged

RI-39

A

1+

I

RI-40

IA

RI-41

A

1

SA-2

A

2

2

SA-3

A

2

2

SA-4

IA

WE-1

A

1

1

WE-2

A

0

0

WE-3

A

0

0

WE-4

A

0

0

WE-5

A

0

0

WE-6

IA

WE-7

IA

WE-8

A

2

2

WE-9

A

2

1

WE-lO

A

2

2

WE-ll

A

0

0

WE-12

IA

WE-13

IA

WE-14

IA

WE-15

A

1+

I

WE-16

A

1

1

WE-17

A

0

0

WE-18

A

2

2

WE-19

A

2

0

WE-20

A

1

1

WE-21

A

0

0

WE-22

IA

WE-23

A

1+

1

WE-24

IA

WE-25

IA

Comments

Fledgling dead below nest

Nest gone
Nest gone

WE-26

A

1+

WE-27

A

0

0

WE-28

A

1+

1

1

page 4

�-116Activity
Table 1

and Productivity

of Colorado Golden Eagles

Site
Number

Activity

Number
of young

Number
Fledged

WE-29

A

1+

1

WE-30

IA

WE-31

A

WE-32

IA

lVE-33

A

WE-34

IA

WE-35

IA

WE-36

A

LO-1

A

1 egg

10-2

A

2 eggs

LO-3

A

1 egg

LO-4

A

1

LO-5

A

0

0

LO-6

A

lA-I

A

1

1

Comments

Nest gone
1+

1

0

0

1

1

page 5

�-117-

TABLE 2
Summary of Actual

and Estimated Productivity

110 of 139 sites (79%) were determined

of Colorado

Golden Eagles

to be active.

80 (73%) of the active sites were known to have produced

eggs.

8 (7%) additional

eggs as was indicated

sites were suspected

by incubating

to have contained

adults.

68 (61%) of the active sites produced young.
59 (54%) of the active sites fledged young.
A total of 54 young were observed in 36 nests prior to fledging.
34 additional

young were assumed to have been produced by 33 other sites.

estimated productivity

of
Total

was 86 young produced by 68 sites.

A total of 59 young were observed
7 additional

A minimum

to have fledged from 52 sites with at least

young fledged from 7 other sites.

success was 66 young fledged from 59 sites.
Averages.
Average

observed brood size:

1.50 young/site

Average

estimated brood size:

1.26 young/site

Average

observed fledging Success:

1.13

Average

estimated

1.11

fledging success:

Total estimated

fledging

�-118-

TABLE

Activity

Symbols:

and Productivity

A = Active

3

of Colorado Osprey - 1973

site

Activity

No. of
Young

Number
Fledged

Comments

GR-2

A

0

0

Adults

JA-2

A

0

0

Adults incubating.

JA-3
LP-l 1.1

A

0

0

Adult incubating

A

2

2

Eyrie
Designation

1/ Reported

to investigators

incubating.

by reputable observer.

�TABLE 4
Table 4 - Occupancy
Site
1
2
3
4
5
6
7
8
9
*10
*11
12
13
14
15
16
17
18
19
20
21
22
23
*

of Colorado

Peregrine

Pre-1964

1964

1965

P

P

+
+
+

p

p

P

A
A
M

Falcon Eyries
1966

1967

1964-1974
1968

1969

1970

1971

1972

1973

P
P

P

P
P
P

M
P
P

P

P
P

P
P
P
A
P
P
P
P
P

P
P
P
P
P
P
P
P
P

P
P
A
P

p

P

p

+

V

p

P
A
F

+
+
+
+
+
+
+
+
+

V
V

+

V

P
M
P

p

V

A

V
V

V
V

V
V
A

1./

V
V
V
V
A

These are neighboring

V
V
V
V

sites and this could represent

V

V
A

a shift of one pair from site 10 to site 11

1/ An adult female w~s found dead in the vicinity of th~ eyrie.
+ = Known to have had pairs present prior to 1964.
Blank Spaces = No data available for that year.
P = Pair.
M = Male .
F = Female.
A = Lone adult·
V = Site was vacant.

P
V
A
V
V
V
V
V
V
V
V
V
V

I
•.....
•.....
\0
I

�-120TABLE 5

Results of Nesting Colorado

1/

Peregrines

in 1973

Visits

Eggs

Young

1

7

3

o

Adult present 4 March; copulation, incubation 7 May; 3 eggs (1 spoiled) 17 May;
pair inattentive 5 June (zock climbers in
eyrie).

2

lO

+2

o

Copulation 5 April; incubation 2 May;
clutch broken 7 May; shell fragment and
spoiled egg on new ledge 25 May.

3

4

?

o

Pair absent early in season; not incubating
in middle of incubation period; not present
in early summer, etc.

4

7

o

o

Pair seen once, lone birds on two visits;
no evidence of nesting.

5

7

+'!J

o

Copulation seen 1 March, 13 May; pair
visited ledges in mid-May; incubated in midJune in apparent renesting attempt; pair
absent 19 July.

6

13

3

2

Copulation 10 March and through early April;
laying 11 April; chose new ledge 16 April;
incubating 21 April; 3 eggs 19 May; two
2-wk. young 5 June; young near fledging in
late June; young flying early July.

7

4

+

o

Probable incubation 5 May, incubation
ledge empty, female passive 31 May.

8

2

1

o

Female carried food to ledge 25 May; one
spoiled egg only 31 May, female incubating.

9

2

?

?

Male on cliff 14 May, incubation possible;
pair flying and p~rching, young unlikely
9 June.

lO

2

?

o

Copulation and ledge scraping
14 May;
lone adult passive 18 July, no young.

11

3

?

o

Pair flying and vocal 12 April; present,
possibly pair 16 June; no young 17 June.

Eyrie

1/ eyrie designation

corresponds

~/ number of eggs undetermined.

Remarks

to designation

in Table

4.

26 May;

�-121-

TABLE

Eggshell

Condition

.

6

of Colorado

Peregrine

Eggs

*

Eyrie

Year

Thickness
Index

Thictyess
(mm)-

Shell Weight
(g)

1

1973

1.62

0.325

3.86

2

1973

1.52

0.305

3.66

8

1973

1.33

0.254

2.92

6

1971

0.284'1-'/
0.

294'l:./

+
Meaxr-sE

+
1.49-.08

Information
Albta.,

below

Saska.,

from other studies

for comparison

3/

3/
pre '47-

Mass.,

N. J ., 1947-502.1

Ungava,

1967, 1970f!:.!

+
1.87..:..0.03

+ •005
o •359..:..0

+
4.12~.09

+
1.99..:..0.01

0.375±o.005

+
4.47..:..0.04

+
1.54..:..0.09

+
0.310..:..0.025

+
3.35..:..0.20

0.291±o.013

l/

Shell plus shell membrane

~/

Shell membranes. absent; values are for shells plus the mean membrane
(O.06mm; range 0.05-0.07) for the other 5 shells.

1/

Anderson

and Hickey

~/

Berger

et al (1970).

*

Table

removed

+
3.4~.28

pre '47-

Montana
East U.S.,

is included

+
0.292-.012

(1972).

from data provided

in W~rk Plan I, Job 1 of W-124-R-2

thickness

�-122TABLE

Chlorinated

Hydrocarbon

7

Residues

in Colorado Peregrine

Eggs*

ug/g
DDE

PCB

DieldIio

Eyrie

Percent
Moisture

Percent
LiEid

1

81

5.2

28

538

0.1

6.4

123

2

82

4.2

16

381

0.1

3.6

86

8

76

5.0

40

571

ND

5.9

84

Mea~E+

+
79.7-1.87

+
4.S-:-.3

28:;'.9

496'±58.6

+
5.r.86

+
97.6-12.7

Information

below from other studies is included

we,t
Wt.

Lipid
Wt.

Alaska!/

Wet
Wt.

for comparison
407

2/
N.W. Can.- G r. B·r~t.-3/

+
17.8-10.9

+
0.s-:-Q.7

11. 9

0.6

Alaska~/
5/
UngavaAlaska.§.1

+
27.4-13.0

46g2:l94

12.7

253

Tundra

889

Taiga

673

Aleutian

.167

1.1

Calculation

~/

Enderson and Berger

1/

Ra t c Lf f f e

!!./

Lincer et al (1970), n=3.

from Cade et al (1968) , n=~.
(1968) , n=f,

(1965), n=13.

2.1 Berger et al (1970) , n=1O.
.§../Cade et al (197i), n=19,
14, 11 respectively.

*

Table removed from data provided in Work Plan I, Job 1 of W-124-R-2

Wet
Wt.

Lipid
Wt.

�-123TABLE 8

Activity

and Productivity

= Active

site; ~

Activity

Number of
eggs

of Colorado

Prairie Falcons

Symbols:
A
Site No.

= Inactive

site; -- = No observations.

Number of
young

Number
fledged

1+

1+

1 young observed

1+

o

Young hatched,

Comments

DO 1

A

DO 2

A

DO 3

A

1+

1+

1 young observed

DO 4

A

3

3

(2 females,

DO 5

A

4

4

(2 females, 2 males).

DO 6

A

1+

o

Young hatched,

DO 7

A

3

3

(1 female,

2 males).

DO 8

A

3

3

(2 females,

1 male)

DO 9

A

1+

o

Young hatched,

DO 10

A

1+

DO 11

IA

DO 12

A

DO 13

4

disappeared.
in vicinity.

1 male).

disappeared.

disappeared.

Young heard in eyrie.

1+

o

A

1

1

DO 14

A

1+

DO 15

A

1+

DO 16

IA

4

in vicinity.

Young hatched,

disappeared.

o

Young hatched,

disappeared.

o

Young hatched,

disappeared.

DO 17

EP 1

A

3

GR 1

A

1+

HU 1

IA

HU 2

IA

JE 1

A

LS 1

A

3

Female incubating.

1+

Young heard in eyrie.

LS 2
LS 3

A

MF 1

~

PA 1

IA

PA 2

A

1+

PA 3

A

1+

1+

Young

apparent ly fledged.

Adult carried prey to ledge.

�Activity
Table 8

and Productivity

Site No.

Activity

of Colorado

Number of
eggs

-124Prairie Falcons

Number of
young

Number
fledged

page 2

Comments

PA 4

A

1+

PA 5

A

1+

1+

Young ~pparently

PU 1

A

Eggs failed to hatch.

Young heard in eyrie.
fledged.

RI 1

11/

A

o

o

WE 2 1/

A

4

4

31/
WE 41/

IA
A

3

2

WE 5 1/

A

5

5

WE 6 1/

A

WE 7 1/

A

3

o
o

Eggs failed to hatch.

81/

A

4

o
o
o

o

Eggs failed to hatch.

WE 9 1/

A

5

5

WE 10 1/

A

5

o

o

WE 11 1/

A

5

3

3

WE 12 1/

A

4

4

WE 13 1/

A

o

o

WE 14 1/

A

5

5

WE 15 1/

A

1

1

WE 16 1/

A

4

4

WE 17 1/

A

o

o

WE 18 II

A

4

4

WE 19 1/

A

WE 20 1/

A

o
o

Male p rcaent ,

WE 21

A

o
o
o

o

Pair present.

WE 22 1/

A

4

3

1 young dead below eyrie.

WE 23 1/

A

o

o

Female present.

WE 24 1/

A

4

4

(2 females, 2 males)

WE 25 1/

A

o

o

Pair present.

WE 26 1/

A

4

4

(3 females, 1 male)

WE 27 1/

A

5

5

(1 female, 4 males)

WE 28 1/

A

4

4

(3 females,

WE 29 1/

A

4

4.

(1 female, 3 males)

WE 30 1/

A

4

4

WE

WE

WE

u

1/ Site data originally
(Raptor Populations

2

1 chick found dead below eyrie.

Eggs failed to hatch.

Eggs failed to hatch.

Female present.

1 male)

obtained from Work Plan 1, Job 2 of W-124-R-2,
and Productivity

is included for continuity

Studies in Northeastern

of statewide productivity

data.

Colorado)

�-125TABLE 9
Summary of Actual and Estimated Productivity of Colorado Prairie Falcons
61 of 68 sites (89%) were determined to be active.
44 (72%) of the active sites produced eggs.
37 (60%) of the active sites hatched young.
33 (54%) of the active sites fledged young.
Total number of young observed prior to fledging = 85
Average brood size of 23 sites with known production

= 3.69

Adjusted young production for all active sites hatching young (37 sites) = 137
(This figure was obtained by including average brood size of 3.69 for 14 sites.
which were known to have hatched young, but for which broods had not been counted).
Total number known fledged from 22 sites = 79 (average fledging Success of 3.69)
Adjusted fledging success of all productive sites (assuming an average brood size
of 3.69 young for 4 sites known to have hatched young, but for which the number
fledged was not known) = 94
94 = 2.84
Thus, the adjusted average fleding Success of all productive sites = 33

�-126-

TABLE 10
Results of Aerial Flights for Golden Eagles in NE Colorado
Date

1972-73 and 1974

Adults

Juveniles

Unknown

Total Area
Estimate·

Dec. 19, 1972

22

6

4

320

10 .6

Jan. 24, 1973 1/

16

8

0

240

8.0

Feb. 16, 1973

18

7

0

250

8.3

Jan. 16, 1974

19

3

0

220

7.3

Eagles per
100 s9. mi.

1.1 This flight will be the only one of the winter 1972-73
used for comparison
in the future.

TABLE 11
Results of Aerial Bald Eagle Census in Northeastern
Date

Adults

Juveniles

Dec. 19, 1972

23

15

Jan. 24, 1973

18

Feb. 26, 1973
Jan. 16, 1974

Unknown

Colorado

- 1972-1974

Total

% Juveniles

0

38

39

13

0

31

42

47

19

0

66

29

16

15

0

31

48

�TABLE 12
Summary of Statewide W.C.O. Eagle Concentration Counts, 1972-1973
Count Period

Number of
Respondents

Feb. 12-18, 72

57

150

17

64

231

153

39

Mar. 1-6, 1972

54

101

32

0

133

137

Feb. 3-9, 1973

51

46

6

9

61

Mar. 3-9, 1973

46

40

7

0

47

Bald Eagles,
Ad Imm Unk

Subtotal

Golden Eagles
Ad Imm Unk

Subtotal

Unknown

Total

14

206

29

466

26

21

184

15

333

117

27

14

157

6

225

144

41

22

207

15

269

I
•....
N

-..,J

I

TABLE 13
Average Number of Eagles Sighted per Observer (Eagle Concentration Counts
1972-1973
Number of
Respondents

Bald
Eagles

Golden
Eagles

Total

Feb. 12-18, 1972

57

4.05

3.61

8.17

Mar. 1-6, 1972

54

3.40

6.16

Feb. 3-9, 1973

51

2.46
1.19

3.07

4.41

Mar. 3-9, 1973

46

1.02

4.50

5.64

Count Period

�-128-

TABLE 14
Eagles Observed Feeding on Carrion February 3-9 and March 3-9, 1973
Species of Eagle
Golden
Bald
1

4

Type of
Carcass*

1

Cow

2

2

Elk

1

1

Elk

1

5

Deer

1

1

Deer

1

1

Deer

1

1

Deer

1

1

Deer

1

1

Deer

1

Deer

2

2

Jackrabbit

2

2

Jackrabbit

2

2

Jackrabbit

1

Jackrabbit

4

Rabbit

2

2

Rabbit (Unk. ssp.)

2

.2

Rabbit

(Unk ,

1

1

Rabbit

(Unk. s sp , )

1

1

Rabbit (Unk. ssp .)

1

1

Rabbit (Unk. ssp.)

1

1

Rabbit

1

1

Pheasant

2

2

Unknown

1

1

1
3

TOTALS

Total Number of
Eagles per Carcass

1

1

Unknown

1

1

Unknown

1

1

Unknown

40

26

31

*Note:

9

The majority of the rabbit carcasses were apparently

on the shoulders of roads.
kill.

(Unk. spp.)

(Unk ,

ssp.)

ssp. )

roadki11ed as they were

The cow, deer and elk were probably victims of winter-

�-129TABLE 15

Behavior of Eagles Observed February 3-9 and March 3-9, 1973

Activity

Bald

Golden

Unknown

Total

Flying

(flapping)

16

76

16

108

Soaring

(circling)

14

52

8

74

30

128

24

182

Feeding

9

31

0

40

Perched
Trees

38

21

1

60

Power Poles

0

63

0

63

Ground

3

47

2

52

Fence Post

0

18

1

19

Rock Outcropping

1

15

0

16

Ice

4

2

0

6

Haystack

0

2

0

2

Windmill

0

2

0

2

Pile of Wood

0

1

0

1

Bush

0

1

0

1

Snowbank

0

1

0

1

Unknown

2

5

0

7

Subtotal

48

178

4

230

87

337

28

452

Subtotal

Total

�-130TABLE 16
Habitat Preferences

of Eagles as Observed on Statewide W.C.O. Eagle Concentration
Counts (1973)

Habitat Type

Golden

Bald

Unknovm

Total

Agricultural
Pasture
Grassland
Grassland Pasture
Riverbottom
Riverbottom-Cottonwoods
Riverbottom-Coniferous Forest
Riverbottom-Meadow
Cottonwood-Willow-Meadow
Cottonwood-Lakeside
Cottonwood-Cedar
Coniferous Forest
Pine-Pinyon-Juniper
Pine-Aspen
Pine-Aspen-Sage
Pine-Oakbrush
Aspen-Sage
Pinyon-Juniper
Pinyon-Juniper-Cottonwood
Pinyon-Juniper-Sage
Oakbrush
Oakbrush-Sage
Sage
Sage-Cedar
Sage-Bitterbrush
Sage-Greasewood
Sage-Willow
Sage-Serviceberry
Sage-Meadow
Meadow
Meadow-Wi.1low

12
12
48
10
4
3
3
6
5
5
2
5
3
3
3
6
5
19
0
9
5
5
111
9
10
9
8
5
8
5
1

12
4
7
0
20
10
5
0
7
4
0
5
0
0
0
3
0
5
1
3
2
0
10
1
0
0
0
0
0
0
0

2
2
1
0
0
1
0
0
0
0
0
2
0
0
2
3
0
1
0
0
0
0
13
1
0
0
0
0
0
0
0

26
20
56
10
24
14
8
6
12
9
2
12
3
3
5
12
5
25
1
12
7
5
134

339

101

28

468

Total

11

10
9
8
5
8
5
1

�-131TABLE 17
of Dead and Injured RaEtors ReEorted

Summa~

-

to Division

01

~
o,.f

~

.&amp;..I

s::

00

$.I

0
o,.f
1Il.&amp;..l
.r-! 0
.-I 01
.-I •...,
0,.0

uO

.&amp;..I

0
..c
III

§
C!&gt;

01
.-I
0
o,.f

..c:

01
:::-

s::
0

.r-!
.&amp;..I

s::

0
o,.f
.-I III
C1I III

tlOO1

01 III
.-I III
.-I 0
1-1 Po.

::s

~

]

§

Golden Eagle

2

10

7

Red-tailed

Hawk

9

4

6

3

4

Great Horned Owl

11

2

5

2

4

Am. Kestrel

6

2

5

3

2

Swainson s Hawk

1

1

1

9

Rough-legged

3

7

I

Hawk

3

0
0
$.I
.&amp;..I

0
01
.-I

c:l.

.&amp;..I

C1I
$.I
E-t

C1I
U

"0
.-I
0

.&amp;..I

..c

I

tlO

01

C1I
U

~

.&amp;..I

s::
s::

01

E!

$.I
01

.-I
C1I

.&amp;..I

0

0
E-t

2

·37

1

2

29

1

1

27

4

24

1

13

1

12

tlO

~

7

4

::s

0
III
o,.f
0
Po.

01
.-1$.1
001

s::..c:

1-1.&amp;..1

2

1

1

1

Bald Eagle

1

Screech Owl

1

4
2

2

1973

I
C1I
01

&gt;.c

..c
...:I

1

during Jan.-Dec.

1

..c:

1

.&amp;..I

5
5

Prairie Falcon

1

1

Marsh Hawk

1

4

Cooper s Hawk

1

1

1

1

2

1

3

2

1

3

I

Sharp-shinned
Short-eared

Hawk
Owl

Goshawk
Burrowing Owl

1

5
5
2

4

1

3

1

Saw-whet Owl

1

Long-eared Owl

1

1

2

1

2
2

1

Barn Owl

2

Peregr:i,.ne
Falcon
Ferruginous

Hawk

Broad-winged

Hawk

F1ammulated

Owl

Total

1

1
1

1
1

1

1
42

2

39

32

18

18

10

-s

4

1
3

2

13

187

�-132TABLE 18
Seasonal Occurence

Cause of Mortality

of Mortality

and Injury to Raptors - 1973

Jan

Feb

Mar

Apr

May

1

2

1

3

2

Gunshot

6

1

4

4

1

Vehicle Strike

5

1

2

3

4

1

2

1
3

Collision

w/Object

Illegal Possession
Unknown

1

2

3

1

Electrocution

1

4

2

1

Fall from Nest
Leg-hold Trap

1
2

1

1

Ju1

Aug

Sep

Oct

Nov

Dec

Total

5

3

3

6

12

4

42

1

2

3

2

9

6

40

6

3

2

1

3

2

30

6

4

1

2

2

2

1
2

1

3

Jun

1

18
2

1

1

11

6

9

1

Caught by Cat

1

1

6
1

1

4

Poison
Inclement Weather

3
1

1

3 3/

-

4

29

16

187

""i:./

17

1/ Thorn in leg
2::/ Fowl pox
1/ Overgorged
!if Plant burr between legs

13

14

15

14

6

26

14

11

13

3
2

2/

11/

Other

Total

18

�-133TABLE 19

Raptors Banded by the Division
Species

in 1973

Number Banded

Prairie Falcon

51

Ferruginous

Hawk

26

Red-tailed

Hawk

14,

Kestrel

12

s Hawk

12

American
Swainson

I

Golden Eagle

11

Cooper s Hawk

4,

Marsh Hawk

2

Barn Owl

1

I

Broad-winged

Hawk

1

F1ammu1ated

Owl

1

Goshawk

1

Great-Horned

Owl

1

Harris Hawk

1/

1

Total

11 Harris hawk was held in zoo for several years,

138

then banded and released.

�-134APPENDIX

A

Date

NEST DATA

_

Time

_

Landowner
Species
Location:
County

I.

~~~----.----_ N or S; R._,

T

Supporting

_

W; Sec ,

_

Additional

_

_____

1/4

DetaUs

_

Structor Information

CLIFF OR OUTCROPPING
height

_

width

length,

_

Overhand: width

length,

_

Pothole:

width

Ledge:

depth.

_

ht to overhand.

_

Other:
Direction of Avis
Geology:
TREE
~:pecies:

_

height

No. of trees in vicinity
O~HER STRUCTURE

_

_

(describe)

_

Ht. above ground.

Ht. to top of structure

E:;{p&lt;.)sure
(direction)
Nest Materials
II.

rBH~

_

Size:

_

~

_

Habitat Data
Topography

_

Dominant Vegetation,

_

G~ologic Formations

_

Drainages/Impoundments:
Soil types

Direction~

_ Distance,

_
_

Land Use
Ill. Behavior
Adult s Acti vi ty:

Young Activity:

_

Ma1e.
Female.

_
_

�Apt;endix A - page 2

-135IV.

Prey Items
No. of Individuals

Species

Unidentified
V.

Age

------------------------_ Sexo

_
o

_

Nest Failure
No. of unhatched eggs

_

No. of destroyed eggs

No. of dead young~

_

Age

_

_

Cause of Destroyed eggs

_

Cause of Dead Young

_

Rerra Lns

VII.

item I.D. No.

Eggs/Young
No. Present
Band No.

VI.

Portion Remaining

and Shell 1.D. No.

_

uisturbance Factors
~oad/Highway

type__________

Degree Use

,Trail/Track
Habitation:

type.__________

Degree Use.
_ Dist.&amp; Direct.
Dist. &amp; Direct.

o

_ Dis t.&amp; Direct.

-----_
_

Other Human Activiti~s
VII.

Additional

_

Comments:

--_.-_. ---------_ .. - . - --------_. --

----''--------------------------

�Appendix

-136-

A - page 3

Date:
Property

_

Owner Data:

Name:

Phone:

_

Mailing
Address:

_

Location

of Residence:

_

Attitude

toward raptors and willingness

to protect raptors nesting on

his property:
Attitude

_

toward public accesS:

Description

----------------------.------------------

(sketch map) of land he holds in the vicinity

of the nest

site(s) •

List of rapt or nests on his property
Location

Species

Were the following

left with the landowner?

T;ivision Identification
Violation

Report Card

Card

Yes

,=,

No

P

Yes

'--:J

No

0

�-137-

JOB PROGRESS

REPORT

sea te of _--=C:;:o:..:l:,.:o:.,:r:.:a:.;d:.;o:.._
Project
Work

Raptor Investigations
No. ~W:...-..:::l:::..24:::..-.....:R::.--=2=--_

Job Title:

Raptor'Population
eastern

Period

Job No •.
--=2~

1

Plan No.

Covered:

Personnel:

Studies

in North-

ColoraJo

February

Gerald

and Productivity

_

1, 1973 through

January

1, 1974

R. Craig

ABSTRACT
Twenty-four

golden

eagles,

falcon

and 17 great horned

square

mile

37 ferruginous

owl nests

study .irea in Northeastern

and fledging

success

Prey items

for golden

cons were

enumerated.

were determ~ned
eagles,

were

hawks,

occupied

Colorado.
for the above

ferruginous

29 prairie

on a 2,000
Productivity
four species.

hawk and prairie

fal-

��-139RAPTOR POPULATION AND PRODUCTIVITY

STUDIES IN NORTH~STERN

COLORADO

Gerald R. Craig
To intensively

document nesting and wintering

ology, habitat requtrement s , productivity
the Pawnee National Grasslands

densitie~,

migration

chron-

and prey pref exenc es of raptors on

and adjacent prairie

lands.

SEGMENT OBJECTIVES
1.

Document

exact nesting densities and productivity

the Pawnee National

Grasslands

of raptors present

and adjacent prairie

2.

Document prey items at nest sites.

3.

While at nest sites, record disturbance

on

lands.

factors, land use practices,

habi-

tat type and nest site features.

METHODS AND MATERIALS
Figure

1 depicts the study area of approximately

and east of Fort Collins.
National

Grasslands

cropping

capped by volcanic

The area encompasses

which is predominantly

2,000 square miles north

the majority

of the Pawnee

shortgrass prairie.

Sandstone

out-

rock borders the area along the north and dips into

the center at about Grover.

The western half of the area (approximately

square miles) was intensively

surveyed for nesting rapto~s while the eastern

half of the area was not covered as extensively

due to limited manpower

1,000

availa-

bility.
The investigation

commenced

in April and proceeded

through mid-July.

This

did not permit the effective .survey of great horned owls nor that of Swainson' s
hawks which fledge young late in July.

However, all known nest sites that have

been occupied over the last three years were surveyed for activity.
four-wheel

Use of a

drive vehicle, a trail bike and a spotting scope were used to make

more effective use of limited time available.
Nest sites were observed from a distance during incubation
occupancy.

Several visits were made to the nests betweell hatching

to ascertain hatching
Fledging

to ascertain

success, monitor nestling mortalities

success was determined

and fledging

and prey items.

by visiting the nests after the young had fledged

and counting young in the vicinity.

�-140-

RESULTS AND DISCUSSION
Golden Eagles
Twenty-four

out of 36 sites were occupied.

A total of 20 young were pro-

duced and 18 young w~re fledged for 16 sites for an averaga brood size of 1.33
and fledging

success of 1.20 young per successful

site (Table 2).

density of 1.60 pairs per 100 square miles, production
fledging

of 1.60 young and

success of 1.40 young per 100 square miles were established

Young eagles were observed up to five weeks after f~edging.
were not very active, would not fly great distances although
of sustained
returned

A nesting

(Table 9).

In all they

they were capable

flight, and wer~ dependent upon adults for food.

Several young

to the nest for short periods after they had b~en out for several weeks.

One fledgling was seen to return to the nest after being out for five weeks.
Prairie Falcons
Ninety-seven
sites produced

% (29) of 30 sites were determined

67 young (average 3.94 young/site)

3.82 young/site).

It was calculated

to be active.

Seventeen

and fledged 65 young

(average

that there wer~ 1.70 nesting pairs per

100 square miles, 3.80 young were produced per 100 square miles and 3.70 young
were fledged per 100 square miles.
Post fledging behavior
nests.

of young prairie

falcons wc:s investigated

at three

Upon fledging at 29 to 33 days of age, the young falcons still have one

inch of feather growth to complete

on their tails and wings.

maries give the wings of fledglings

a rounded appearance.

were still in family groups 35, 39 and 40 after fledging.

The shorter pri-

The prairie

falcons

Young were still

begging food from adults at that period.
Ferruginous

Hawks

Thirty-seven
55 young
Tabulation

(80%) of 46 sites were occupied.

(1.4~ young/nest)
revealed

Twenty-two

nests produced

and 21 sites fledged 45 young (1.22 young/nest).

2.40 pairs per 100 square miles, 3.40 young produced per

100 square miles, and 2.70 young fledged per 100 square miles.
The following causes of nesting failures were noted:

four nests were blown

down, one nest abandoned

during incubation,

tion.

took young from one nest ~nd two young from a second

Coyotes probably

nest were discovered

and two nests failed due to preda-

in a golden eagle eyrie.

�-141-

Swainson's

Hawks

Data on this species is incomplete
to young production

of most sites.

due to termination

Table 7 sUmmarizes

of the study prior

those findings available

upon termination.
Great Horned Owls
Fifteen

(88%) of 17 sites were occupied by pairs.

duced 19 young and 8 sites fledged 16 young.

Ten of the sites pro-

Table 9 reveals 0.60 pairs of owls

per 100 square miles, 0.60 young per 100 square miles and 0.60 fledglings
100 square miles.
As fledging of many owl nests occurred before the investigation
taken, productivity

per

was under-

data is meager.
RECOMMENIlA.TIONS

1.

If present manpower

limitations

horned owls and Swainson's
investigation
2.

several species such as great

hawks should be deleted to permit more intensive

of golden eagles, prairie falcons and ferruginous

Use of aircraft

to determine productivity

survey remote portions

Prepared by:

are continued,

or activity

hawks.

of known sites and

of the study area for new nests might be more efficient.

e. 12. ~

(J ; ,

Gerald R. Craig

Date:

2,/~
"/7!C
~
I

�FIGURE
INTENSIVE

~_, o,r

~

Vi6r ,/)00
r~t ' 0

,;

"\

"

...

"

',,,-',

-r

./

ti: II.-~~OV15 ~ ,J
. ko Olk
.' \'0

,\

,

\

,I,

"

).

f'

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COLORADO

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00

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-

NORTHEASTERN

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STUDY AREAS

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AND EXTENSIVE

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�-143TABLE 1
Activity and Productivity
Symbols:
Site
Designation
WE-l
WE-2
WE-3
WE-4
WE-S
WE-6
WE-7
WE-8
WE-9
WE-lO
WE-II
WE-12
WE-13
WE-14
WE-IS
WE-16
WE-17
WE-18
WE-19
WE-20
WE-21
WE-22
WE-23
WE-24
WE-2S
WE-26
WE-27
WE-28
WE-29
WE-30
WE-31
WE-32
WE-33
WE-34
WE-3S
WE-36

A

Activity

active site; IA
Number of
Young

A
A
A
A

A

= inactive site; -- = no observations
.Numb.er .
Fledged

o
o
o
o

o
o
o
o

2
2
2

2
1

IA
IA

A
A

Comments

Spoiled egg

Fledgling dead below nest

2

A

o

o

IA
IA
IA
A

1+

1
I

A
A
A
A
A
A

1

o

o

2
2
1

2

1
I

o

o

1+

1

IA

A

Nest gone
Nest gone

IA

IA
A
A
A
A
IA
A
IA
A
IA
IA
A

Colorado

I

I

A

of Golden Eagles in Northeastern

1+

1

o

o

1+
1+

1

1+

I

o

o

:I.

1

1

Nest gone
Nest gone

�-144TABLE 2

Summary of Actual

and Estimated Productivity
Colorado

24 of 36 sites (67%) were determined

of Golden Eagles in Northeastern

to be active.

16 (67%) of the active sites were known to have produced
15 (62~of

eggs.

the active sites produced young.

15 (62%) of the active sites fledged young.
A total of 14 young were 'observed in 9 nests prior to fledging and a minimum
of 6 additional
minimum

young were known to have been produced by 6 other sites.

total productivity

was 20 young produced by 15 sites.

A total of 18 young were fledged from 15 sites.
Average

brood size:

Average

fledging

1.33

success:

1.20

Thus,

�-145TABLE 3
Activity and Productivity of Prairie Falcons in Northeastern
Colorado
Sy.mbols: A
Site
Designation
WE-I
WE-2
WE-3
WE-4
WE-5
WE-6
WE-7
WE-S
WE-9
WE-lO
WE-ll
WE-12
WE-13
WE-14
WE-IS
WE-l6
WE-17
WE-IS
WE-19
WE-20
WE-21
WE-22
WE-23
WE-24
WE-2S
WE-26
WE-27
WE-28
WE-29
WE-30

active site; IA = inactive site;

Activitl
A
A
IA
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A

Number
of Eggs

3
4
5
5

2

Number of
Young

Number
Fledged

= no observations
Comments

0
4

0
4

Eggs failed to hatch

3
5
0
0
0
5
0
3
4
0
5
1
4
0
4
0
0
0
4
0
4
0
4
S
4
4
4

2
5
0
0
0
5
0
3
4
0
S
1
4
0
4
0
0
0
3

I chick found dead below eyrie

0

4
0
4
S
4
4
4

Eggs failed to hatch
Eggs failed to hatch
Eggs failed to hatch
Eggs failed to hatch

Female present
Male present
Pair present
1 young dead below eyrie
Female present
(2 females, 2 males)
Pair present
(3 females, 1 male)
(1 female, 4 males)
(3 f~males, 1 male)
(1 female, 3 males)

�-146-

~BLE
Summary of Actual

4

and Estimated Productivity

29 of 30 sites (97%) were determined

of Colorado Prairie Falcons

to be active

25 (86%) of the active sites were known to have produced
nests

eggs while 4 additional

(14%) had pairs present and probably produced

17 (59%) of the active sites produced young.

The same number of sites succeeded

in fledging young.
Total number of young produced

= 67

Total number of young fledged

= 65

Average

clutch size known from 5 sites

Average

brood size

Average

fledging

= 3.80 eggs/site

= 3.94 young/site
success = 3.82 young/site

eggs.

�-147TABLE S
Activity and Productivity
Symbols:

Site
Designation
WE-I
WE-2
WE-3
WE-4
WE-S
WE-6
WE-7
WE-S
WE-9
WE-IO
WE-II
WE-12
WE-13
WE-l4
WE-IS
WE-16
WE-17
WE-IS
WE-19
WE-20
WE-21
WE-22
WE-23
WE-24
WE-2S
WE-26
WE-27
WE-28
WE-29
WE-30
WE-3I
WE-32
WE-33
WE-34
WE-3S
WE-36
WE-37
WE-3S
WE-39
WE-40
WE-4I
WE-42
WE-43
WE-44
WE-4S
WE-46

of Ferruginous Hawks in Northeastern

A = active site; ~

Activity
A
A
A
A
A
A
~
A
A
A
A
~
A
A
~
A
A
A
IA
A
~
A
A
A
A
A
A
A
A
A
A
A
A
A
A
~
~
A
A
A
A
A
A
~
~
A

~ inactive site; -- ~ no observations

Number of
Young

Number
Fledged

3

2
4

4
2

3

o

o

3

Colorado

2
1

o
o

Connnents
Nest fell, young unhurt
1 unpiped egg in nest
Adult incubating
Adult incubating

3

o

o

3
2

2
2

2
2

2
2

o

o

3

3

3

2

Pair present, nest gone
Nest fell down
2 young were melanistic
1 young was a runt

o

o

Adult incubating

o

o

2+
3
3
3
1
1+

2

Adults in vicinity
1 melanistic young

3
3

o

1
1

o

o

3

3

o
o
o

3

Adult incubating
1 spoiled egg in nest
Both young were melanistic

o

o

o

Nest down, chicks dead
1 egg spoiled
Nest tilted, eggs on ground
3 eggs
Adult incubating
Adult incubating
Failed during incubation

3

o

o

Adults present, failed

o

o

Building nest, failed

1+
3

1
1

1 sp~iled egg in nest

2

2

o

o

o
o

o

o

Adult incubating
Adult incubating

�-148-

TABLE 6
Summary of Productivity

of Ferruginous
Colorado

37 of 46 (80%) of the sites were active
22 (59%) of the active sites produced young
21 (57%) of the active sites fledged young

Minimum number of young produced
Total number of young fledged

=

55

= 45

Average brood size = 1.49 young/nest
Average

fledging success

= 1.22 young/nest

Hawks in Northeastern

�-149-

TABLE 7

Activity

of Swainson's

Symbols: A
Site
Designation
WE-I
WE-2
WE-3
WE-4
WE-5
WE-6
WE-7
WE-8
WE-9
WE-lO
WE-II
WE-12
WE-13
WE-14
WE-15
WE-16
WE-17
WE-18
WE-19
WE-20
WE-2l
WE-22
WE-23
WE-24
WE-25
WE-26
WE-27
WE-28
WE-29
WE-30
WE-3l
WE-32
WE-33

active site; IA

Activity
IA
IA
A
A
A
A
A
A
A
A
IA
IA
IA
IA
IA
IA
IA
IA
IA
IA
A
A
A
IA
IA
A
A
A
A
A
A
A
A

Hawks in Northeastern

Number
of Young

Colorado

= inactive site; -- = no observations
Number
Fledged

Comments

Constructing

nest

Adult incubating
Pair present
Pair present
Adult incubating
Adult incubating
2

Nest gone

Nest gone
Nest gone
Nest gone
New nest constructed
Pair present
Pair present
Nest gone
Nest gone
Adding to nest
Pair present
Adult incubating
Adult incubating
Adult incubating
Constructing nest
Adults incubating
Constructing nest

�-150TABLE 8

Activity

and Productivity

Symbols:

A

of Great-Horned

Owls in Northeastern

= active site; IA = inactive site;

Colorado

= no observations

Activity

Number of
Young

Number
Fledged

WE-l

A

1+

1+

WE-2

A

2

2

WE-3

A

2

2

WE-4

A

2

2

WE-5

A

Adult incubating

WE-6

A

Adult incubating

WE-7

IA

WE-8

IA

WE-9

A

2

2

WE-lO

A

2

2

WE-ll

A

Adults

WE-12

A

2 eggs

WE-13

A

WE-14

A

WE-15

A

1

WE-16

A

2

WE-17

A

2

Site
Designation

3

Comments

Adult incubating
in vicinity

3
Adult incubating

2

TABLE 9

Summary of Productivity

of Great Horned Owls in Northeastern

15 of 17 sites (88%) were active
10 (67%) of the active sites produced young
8 (53%) of the active sites fledged young
Total number of young produced =.19
Total number of young fledged

16

Average

brood size of 9 sites

2.0 young/site

Average

fledging success of 7 sites = 2.14 young/site

Colorado

�-151-

TABLE 10

Raptor Nesting Densities on the 1,000 Square Mile
Study Area

Species

Active Nests
per 100 mi2

Young ~er
100 mi

Fledglings per
100 mi2

Golden Eagle

1.60

1.60

1.40

Prairie Falcon

1.70

3.80

3.70

Ferruginous Hawk

2.40

3.40

2.70

Great Horned Owl

0.60

0.60

0.60

6.30

9.40

8.40

Total

�-152-

TABLE 11
Prey Taken by Golden Eagles, Ferruginous

Prey Species

Golden Eagles

Manunals
Cottontailed Rabbit
Jackrabbit
Northern Pocket Gopher
Long-tailed Weasel
Black-tailed Prairie Dog
Fox Squirrel
Muskrat
Thirteen-lined Ground Squirrel
Ord's Kangaroo Rat
Antelope (fawn)
Birds
Mallard
Pintail
Ferruginous Hawk
Mountain Plover
Horned Lark
Western Meadowlark
Lark Bunting
Reptiles
Gartersnake
Bullsnake

Total Prey Items

Hawks and Prairie Falcons in 1973
Ferruginous
Hawks

35
38

5

1

8

Prairie Falcons

1
2
2

1

1
2

9

13

1
2

2

1
2

1

1

1

6
6

3
5

31

27

1
2

94

�-153-

JOB PROGRESS REPORT

State of

~C~o~l~o~r~a~d~o~

Project No.

W-124-R-2

Work Plan No.

1

Job Title:

_
Raptor Investigations
Job No. _--=3

_

Raptor Population and Productivity Studies in Southeastern Colorado

Period Covered:
Personnel:

July 25, 1973 through January 31, 1974

Gerald R. Craig, William C. Andersen

ABSTRACT

Occupied nests of 8 golden eagle s, 36 ferruginous hawks, 8 redtailed hawks and 176 Swainson's hawks were located in an area of
4,421 square miles in Southeastern Colorado.
established for each of the above species.

Fledging success was
Decline in fledging success

of Swainson's hawks were related to strong wind storms of the hatching.
Weekly migration and winter trend counts of raptors were also undertaken.

��-155-

RAFTOR POPUlATION AND PRODUCTIVITY STUDIES IN SOUTHEASTERN COLORADO
William C. Andersen and Gerald R. Craig
To intensively document nesting and wintering densities, migration chronology, habitat requirements, productivity

and prey preferences

of raptors on the

Comanche National Grasslands and adjacent prairie lands.
SEGMENT OBJECTIVES
1.

To continue documentation

of nesting densities and productivity

of golden

eagles, prairie falcons, ferruginous hawks, Swainson's hawks, red-tailed
hawks, great horned owls, and burrowing owls on the Comanche National
Grasslands and adjacent prairie lands.
2.

To continue documentation

of the timing of breeding cycles of golden eagles,

prairie falcons, ferruginous hawks, Swainson's hawks, red-tailed hawks,
great horned owls and burrowing owls.
3.

To continue documentation

of prey items at nest sites.

4.

To investigate and categorize nest sites by such features as disturbance
factors, habitat type, human use, dominant vegetation, relief, etc.

5.

To continue documentation

of migration chronology.

6.

To continue to gather trend information on wintering raptor populations.

7.

To categorize important wintering habitats.
METHODS AND MA.TERIALS
Breeding Inventory
Nesting investigations commenced in March of 1973 when great horned owls

initiated nesting activities and continued through July when the young of late
nesting raptors, such as Swainson's hawks, fledged.
area is Southeastern Colorado.

Figure 1 depicts the study

The primary vegetative type in this region is

shortgrass prairie interspersed with cropland and deciduous riverbottom along
the Arkansas River.

All public roads were traveled and all structures capable

of supporting nests which were within sight from the roads were scrutinized
with binoculars and a spotting scope.

When nests were located, they were

�-156-

observed from a distance to determine presence of nesting raptors.

After com-

pletion of hatching, the sites were then visited to ascertain actual productivity.
The purpose of delaying actual visits to nests were delayed until after hatching
to avoid possible disturbance that might cause abandonment of nesting attempts
by incubating adults.

Near fledging time, those active sites which contained

young were revisited and young were banded and presence of prey items was recorded.
Migration and Winter Population Trends
Generally, the censuses commence early in October and continue on a weekly
basis through mid-March.

A roughly circular route of approximately

150 miles

(Fig. 2) were driven at speeds between 30 and 35 mph with two observers.
a raptor was sighted, the vehicle was stopped and identification

When

confirmed using

binoculars and a spotting scope.

Excursions were made off the road when identi-

fication could not be determined.

Species, age, activity and habitat were recorded

for each observation.
RESULTS AND DISCUSSION
Breeding Inventory
Due to reduced funding, the nesting investigations were limited in scope.
Approximately

26.7% (4,421 mi.) of the study area was intensively investigated

for presence of nesting raptors (see Table 1).
In all, 8 active golden eagle nests were located which averaged a fledging
success of .50 young.

Thirty-six ferruginous hawk nests were active and fledged

an average of 1.47 young per nesting attempt.

Eight red-tailed hawks nested and

fledged 1.88 young for each nesting attempt.

Only 62 young fledged from 176

active Swainson's hawk nests, which averaged .35 young fledged per nesting attempt.
Table 2 summarizes the results.

The extremely poor reproductive Success of

Swainson's hawks and ferruginous hawks was attributed to two storms in which wind
velocities exceeded 100 mph in the region and particularly Lincoln and Kiowa
Counties.

Severe winds destroyed 84% of the Swainson's hawk nests and 53% of

the ferruginous hawk nests.

Swainson's hawks were hardest hit because they

build relatively fragile nests near the tops of trees.

Ferruginous hawks con-

struct bulky nests which are situated near the tree bole.
Low producLivity

of golden eagles could not be adequately explained.

Prey

items were common in those nests that produced young and the young at each site
were in excellent condition.

Human interference was responsible for two nesting

failures, one caused by campers and another caused by construction while the pair

�-157were engaged in nest building activities.
Migration

and Winter Population Trends

Table 3 summarizes censuses begun October 7, 1972 which continued on a
weekly basis until March 10, 1973.
January 27.

Counts were not made November

25 and

Peaks in migrating raptors were observed in late October which

were due to increases in kestrels, ferruginous hawks and golden eagles.

In

early December, another peak was due to an influx in ferruginous hawks, bald
eagles and rough-legged hawks.

The peaks in late December and January were

caused by bald eagles and rough-legged hawks.

Raptor numbers gradually dropped

in late January, February and March.
Table 4 provides results of trend counts initiated October 5, 1973 and
completed March 2, 1974.

As in 1972-73,censuses

except November 24 and December 22, 1973.

were made on a weekly basis

The highest peak of 101 raptors occurred

December 8 and was due primarily to an increase in rough-legged
The peak in the fall migration occurred November
legged hawks and kestrels were most abundant.

Prepared by:

GRMO'"

12. ~al ,

Gerald R. Craig

hawks and kestrels.

10 when prairie falcons, rough-

�FIGURE 1

Colorado Counties
SEDGWICK
WELD
MOFFAT

LOGAN

LARIMER

PHilLIPS

ROUTT
MORGAN
YUMA
ADAMS

I WASHINGTON

ARAPAHOE

-

ELBERT
I

co
LI"I

.-!
I

I

fl.

I'll KIN

I _~f

I

ILINCOLN I KIT CARSON

IDOUGLASI

MESA

"

I

MONTROSE
~OURAY

DOLORES

-

MONTEZUMA

,.,

- ""

~

I

DFITA

""

It.LLt.KI
EL PASO

' ~

~

I
I

c:,u,"

I

•",-,y,v."

"

I

J

rr-

I

CHEYENNE

KIOWA

I

PUEBLO

y-

1-.

/

~

I

n

OTE~O

I

BENT

I PROWERS

.-.;r
LAS ArtlMAS

I

SACA

Study Area Boundary

�-159-

FIGURE 2

RESEARCH AREAS

:GRASSLAND

\
-

1
Area boundaries

-1

II

•
-

COLORADO

I

••••• Census route

•

Otero Junior College

�-160-

TABLE 1
Counties Covered by Breeding Inventory

(1973)

within Area

Mi2 of
County
within Area

% of
County
Studied

Mi2 of
County
Studied

2,565

100.0

2,565

38

952

2.24

Bent

1,517

100.0

1,517

33

501

2.28

Cheyenne

1,772

90.4

1,602

11

167

1.39

Crowley

803

86.0

691

44

311

3.92

Elbert

1,865

7.0

131

2

3

2.10

Kiowa

1,795

100.0

1,795

18

323

1.14

Las Animas

4,794

46.6

2,234

26

580

1.20

Lincoln

2,593

56.8

1,473

46

659

1.86

Otero

1,267

90.0

1,140

74

844

8.71

Prowers

1,626

100.0

1,626

5

81

3.51

Total

20,597

.1/ Weighted

average

% of County

County

Total Area
within
2
County(Mi )

Baca

16,546

4,421

Human Popu1ation
per Mi2

3.32 1/

�-161-

TABLE 2
Activity and Nesting Success, 1973

SEecies

Count~

Numb ar
Nest
Atte!!!Ets Fledged

Young/Nest
AttemEts

Active
Nest/IOO mi2

Yo~ne
Pro
u e~
/l00 mi

Golden Eagle
Baca
Bent
Las Animas
Otero

2
2
3
1
8

2
1
1
0
4

1.00
0.50
0.33
0.00
0.50

0.27

0.10
0.19
0.17
0.00
0.13

9
5
1
3
2
1
9
6

14
0
0
3
0
2
21
13

1.56
0.00
0.00
1.00
0.00
2.00
2.33
2.17

0.94
0.99
0.59
0.96
0.61
0.17
1.36
0.71

1.47
0.00
0.00
0.32
0.00
0.34
3.18
1.54

Jb

TI

1:47

0":1r1"

---r.rr

2
3
3
8

5
5
5
15

2.50
1.67
1.67

0.39
0.51
0.35
0.41

0.99
0.86
0.59

57
5
26
3
28
3
46
8
176

23
3
16
4
3
5
8
0
62

5.98
0.99
8.36
100 .00 1/
8.66
0.51
6.98
0.94
4.21

2.41
0.59
5.14
133.00 1/
0.92
0.86
1.21
0.00
1.48

0.21
0.39
0.51
0.11

Ferruginous Hawk
Baca
Bent
Cheyenne
Crowley
Kiowa
Las Animas
Lincoln
Otero

Red-tailed Hawk
Bent
Las Animas
Otero

l.8s

0.77

Swainson's Hawk
Baca
Bent
Crowley
Elbert
Kiowa
Las Animas
Lincoln
Otero

0.40
0.60
0.62
1.33
0.11
1.67
0.17
0.00
0.35

]) Figures are inaccurate due to extremely small sample size of 3 nests
located
2
on 3 mi

of Elbert County.

�TABLE 3
Migration and Winter Populations Trends, 1972-1973

7
Golden Eagle
(adult)
(imm.)
(unknown)
(total)

-

--

Bald Eagle
(adult)
(Lmm ,)
(total)
Ferruginous
Hawk

7

14Y21

1
-

3
3
1
7

-

1

1

1

18'1:./25

2

3
2

4

1
2
2
5

5

28

4

11

6
5

1
3
1
5

2
2

-

11

-

-

g'l:./16

23

30

6

5
3

7
2

2
2
1
5

7
1
8

3
1
3
7

15

8

-

1~/

20'l:..£.7

1

1

-

8

-

9

-1

-

--

1

3

8
1

7 3

1
1

3

1

2

-

- -2 3- -1
-

-

1

2

-

3

2

1

2

3

- - 1

4

- 4 2
4 19 10

7
2
9

21
4
25

-

2

1

10
3
13

-

2

2
1
3

13

17

6

9

1

13

13

5

8

2

3

7

4

2
1

-

4
1
5

18

1

10

8
1
9

-

-

6

3

-

-

1

10 17'1:./24'1:..1

-9

1
2
3

-

March

February

January

December

November

October

-

-

1

-

I
•.....
0N

Rough-legged
Hawk

-

-

1

9

4

5

11

12

8

14

10

9

14

2

6

Red-tailed
Hawk

5

6

6

7

4

4

-

6

3

3

-

7

18

2

Marsh Hawk

2

5

10

9

6

10

2

7

7

2

3

6

18

Prairie Falcon 2

3

2

3

1

-

1

4

5

Merlin

- 2 :2 2
- - - - -

1

-

Kestrel

26

5

12

22

13

23

2

Cooper I s Hawk

-

-

-

- -

2

Great Horned
Owl

2

-

1

Short-eared ~

-

-

-

-

Total
Raptor/mi

-

44 18 52 81 41
.29 .12 ,35 .54 .27

3

1

6

1

1

-

5

5

2

2

4

-

2

-

2

3

-

-

1

1

3

1

1

2

3

1

-

-

-

-

-

- -

-

-

-

3

-

2

-

2

-

2

-

- - - - 6 20
11
6
7
5
- - - - - - -

-

-

- -

-

-

-

-

2

3

3

2

2

2

-

2

-

- - - - -

2

-

-

- -

-

-

- -

61 30
.41 .20

1/ Heavy cloud cover and light mist.

-

1

3

2

2

-

73 51 73 41 57 121
9
,49 .34 .49 .27 .38 .81 .06
'1) Snow, wind,

25
,17

34 27 14
.23 .18.09

-

14
9 11
.09 .06 .07

I

�11\BLE 4

Migration

Golden Eagle
(adult)
(Lmm, )
(unknown)
(total)
Bald

12

19

26

3

10

- -

1

1

Eagle
(adult)
(Lmm, )
(Total)

3
1

7
1

4
1

Hawk

5

3

4

3

5

10

6

Rough-legged

Hawk

- -

-

1

6

12

7

2

6

7

8

7

8

Red-tailed
Marsh

Hawk

Hawk

Prairie

24

17

Ferruginous

Falcon

5
6

3

5

7

- 2 4
- - - 3

9
3

8
9

Trends,

.Janua ry

-

15

22

29

5

2

6
1

4
2

-

7
4

6
2

9 17

15

7

10

4

-

1

8

-2 7- -6 - 116
5 8
- 91
1 1
2
6 9
8
- 10
8 9
7
- 6

-

6

1

2

9

-

5

7

.5

6

1

- - -

2

-

----

--

-

------

-

-

-

34

16

30

30

47

79

56

63 101 92

Raptors/mi.

.23

.11

.20

.20

.31

.53

.37

.42

.67.61

-

---

- - -----

-

9

5
1

1

2

16

1

-

1

2

4

1

-

-

8

3

7

6

5

6

1.

2

-

1

1

-

-

3

5

5

7

3

9

17

6

3

7

6

9
3
2

2

2

9

4

-

3

2

6

-1

2

1

2

1
1

1

-

5

2

-

-

3

23

- - - -

3

6

-

2

2

11

3

26

4
1
5

3

-

19

March

February

- - - -8 4- 6- -1 -2 -1 -3
1

7
1
8

I

Total

12

8
2
10

- - - - 13
8 10
6
6 16 10
Kestrel
- 15 20 17 - 9
Cooper s Hawk
- - - - 1 - - - - - - - Great Horned Owl
- - 1 - - 1 2 - 2 2 4 - 1
Short
eared Owl
- - - - - - - - - - - - 1

Merlin

1973-74

December

- - - - - - - - 1
- 1 1 4 8 5 - - - - - - 2 - - - - - - - - - - - - - 2 1

Populations

November

October
5

and Winter

5

1

-

1

-

-

3

- - - - - - - - - - - - - - - - - 1

- -

~. -

-

-

-

3

2

5

9

2

4

1

3

2

1

2

2

1

75

72

64

50

46 18 23

20

15

16

.50

.48

.43

.33

.31.12.15

.13

.10

.11

I
f-I
0\

(,.oJ

I

�-164-

TABLE 5

Raptors Banded in Southeastern

Species

Colorado

Number Banded

Swainson's

Hawk

70

Ferruginous

Hawk

56

Great Horned Owl

29

Red-tailed

20

Hawk

Golden Eagle

4

American

3

Kestrel

Long-eared

Owl

Rough-legged
Barn Owl
Cooper's

Hawk

2
2

1
Hawk

TOTAL

1
188

�-165-

JOB PROGRESS REPORT
State of

~C~o~l~o~r~a~d~o~

_

Project No. ~W~-~1~2~4_-~R~-~2

_

Raptor Investigations

Work Plan No.

_

Job No. 4

Job Title:

Roadside Raptor Censuses

Period Covered:
Personnel:

~l~

October 11, 1973 through January 31, 1974

Gerald R. Craig

ABSTRACT
Contracts were developed, but due to inability to locate
qualified persons to undertake the censuses at such a late date,
no contracts were let on this project during this segment.

��-167-

JOB PROGRESS REPORT

State of

~C~o~l~o~r~a=d~o

_

Project No.

W-124-R-2

Raptor Investigations

Work Plan No.

2
------=-------------

Job No. 1

Job Title: Peregrine Falcon Propagation and Reintroduction Studies
Period Covered:
Personnel:

October 11, 1973 through January 31, 1974

Gerald R. Craig, James H. Enderson

ABSTRACT

Twenty-six eggs were obtained from two pairs of captive anatum
peregrine falcons.

Nine eggs were fertile and four young were hatched

of which three were successfully reared.

These were the first anatum

peregrine falcons ever produced in captivity.

Due to the low number

of breeding stock in captivity, all young were retained for further
propagation rat.her than being introduced into the wild.

Three pairs

of prairie falcons in the same project laid 22 eggs of which six
were fertile and two young were reared.

��-169-

PEREGRINE FALCON PROPAGATION AND REINTRODUCTION STUDIES
Gerald R. Craig

To captive1y produce eggs and/or young from peregrine falcons for reintroduction
into the nes~of

wild peregrine falcons that have low or no natural productivity.

SEGMENT OBJECTIVES
1.

To establish the cause for reproductive failure of wild peregrines.

2.

To produce young or eggs from peregrine falcons being held in captivity at
the facilities of Dr. James H. Enderson.

3.

To visit wild peregrine falcon eyries where adults are present and replace
spoiled eggs with captive1y produced eggs or young.

4.

To keep those eyries containing captively produced eggs or young under
observation to ascertain the success of the endeavor.

METHODS AND MATERIALS
Dr. James H. Enderson was awarded the contract to captively produce peregrine
falcons for reintroduction efforts.

Enderson's facility at Colorado Springs

consists of a building containing five lofts 21 feet in length, 6 feet in width
and 14 feet in heighth.
the facility.

A sixth room of slightly smaller dimensions completes

Nest ledges approximately 3 feet deep and six feet in width are

provided near the ceiling at the end of each 10ft.
with stream sand.

The nest ledges are covered

One-way glass at the rear of each nest ledge permits obser-

vation of the nest ledge and much of the loft.
locations in each loft.

Perches are placed at various

Two x eight foot openings, covered with heavy gauge wire,

provide light and ventilation to each loft.

Pairs of falcons in each 10ft are

fed five week old cockrels and coturnix quail.

Bathing water is continually

supplied to each room.
In 1973, two pairs and a single two-year-old male anatum peregrine (Falco peregrinus anatum), one pair of Peale's peregrines (Falco peregrinus pealii), one
immature female arctic peregrine (Falco peregrinus tundrus) and three pairs of
prairie falcons were maintained at the facility.

The prairie falcons were used

to test artificial insemination techniques as well as develop artificial incubation and brooding methodology.

�-170Several peregrines were artificially

inseminated. to apparent Success.

Three

days to a week after completion of clutch of eggs, the entire clutch was removed
and artificially

incubated.

Within a week to ten days after removal of the first

clutch, the falcon laid a second clutch. Thus "recycling" is an excellent method
of increasing productivfty of captive falcons.
RESULTS AND DISCUSSION
The prairie falcons laid a total of 22 eggs of which 6 were fertile.
were permitted to incubate the eggs naturally.
and two chicks did not survive to hatching.
reared and fledged.

The parents

Of the six fertile eggs, two froze

Two chicks were successfully hatched,

The anatum peregrines succeeded in laying 26 eggs, of which 9 were fertile.
eggs, except the last clutch were artificially
were successfully hatched, but died.

incubated.

All

Four young peregrines

The remaining three females were all reared

and are being held for further propagation.

These are the first anatum peregrines

ever produced through captive propagation.

Due to scarcity of captive anatum

peregrines for use in breeding programs (there are less than a dozen pairs in
captivity at this time) it was decided to hold those progeny produced for the
first few years to increase breeding stock.
Information gathered in the course of Work Plan 1, Job 1 (Statewide raptor population and characteristic
in wild peregrines.

studies) indicates that pesticide levels are critical

Tables 1 and 2 provide details of the analysis.

are also included in Job Progress Report for Work Plan I, Job 1.).

(These tables

�-171Decreased production relates directly to decreased eggshell thickness which is
about 20% less than pre-1948 samples.
reported for peregrines.

DDE

residues are as high as levels ever

The next step will be to attempt to trace down the

source of prey containing high pesticide residues and continue to monitor pesticide
levels to document possible changes.
RECOMMENDATIONS
1.

Continue collecting and analyzing infertile or addled peregrine eggs and
shell fragments to increase sample size of data base.

2.

Collect and examine representatives from the prey population in areas
hunted by nesting peregrines to determine pesticide levels.

3.

Maintain captively produced peregrine falcons in the breeding program
for at least another year to increase the presently meager captive
breeding stock.

4.

Investigate methods of increasing egg fertility through artificial
insemination.

Prepared

by,

a IZ ~
Gerald R.~

Date:

�-172TABLE 1

Eggshell Condition of Colorado Peregrine Eggs
Eyrie

Year

Thickness
Index

*

Shell Weight
(g)

1

1973

2

1973

8

1973

6

1.62
1.52
1.33

1971

0.325

3.86

0.305

3.66

0.254

2.92

0.284'1:/
0.294~/

+
Mean-SE

+
1.4~.08

+
0.292-.012

+
3.4S-:-.28

Information below from other studies is included for comparison
3/
Albta., Saska., pre '47Montana

+
1.87~.03

East U.S., pre '472/

+
1.9~.01

Mass., N. J. , 1947-501/

+
1.54~.09

+
0.35~.OO5

+
4.l2~.09

0.3752:0.005

+
4.47~.04

+
0.31O~.025

+
3.35~.20

Ungava, 1967, 1970!!.!
0.2912:0.013

.Y

Shell plus shell membrane

'!:..!

Shell membranes absent; values are for shells plus the mean membrane thickness
(0.06mm; range 0.05-0.07) for the other 5 shells.

3/

Anderson and Hickey (1972).
Berger et al (1970).

Table removed from data provided in Work Plan I, Job 1 of W-124-R-2

�-173TABLE

Chlorinated

2

Hydrocarbon Residues in Colorado Peregrine

Eggs*

uglg
DDE
We.t
Wt.

PCB

DiddtiD

Eyrie

Percent
Moisture

Percent
LiEid

1

81

5.2

28

538

0.1

6.4

123

2

82

4.2

16

381

0.1

3.6

86

8

76

5.0

40

571

ND

5.9

84

+
Mean-SE

+ 87
79.7-1.

+
4.S-:-.3

28~.9

496±58.6

Lipid
Wt.

Wet
Wt.

+

5.r.86

Information below from other studies is included for comparison
A 1aska-II

21
N.W. Can.-

407

-

G r. B'r1t.-31
A1aska-41
Ungava-51

+
17.8-10.9

+ 7
o. S-:-O.

11.9

0.6

+
27.4-13.0

469±194

12.7

253

Alaska§.I
Tundra

889

Taiga

673

Aleutian

167

}j

Calculation

11

Enderson and Berger

from Cade et a1 (1968), n=2.
(1968), n=5.

_~I Ratcliffe (1965), n=13.

1::.1

Lincer et a1 (1970), n=3.

'il

Berger et al (1970) , n=lO.

£1

Cade et al (1971), n=19, 14, 11 respectively.

Wet
Wt.

'[able removed from data provided in Work Plan I, Job 1 of W-124-R-2

Lipid
Wt.

+

97.6-12.7

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                  <text>April 1975

-1-

JOB FINAL REPORT

State of

COLORADO
----~~~~~----------

Project No.

Game Bird Survey

W_-_3_7_-_R_-_2_8 _
1

18

Work Plan No.

---------------------

Job Title

'Pheasant Roadside Cover Evaluation Study

Job No.

-------------------------------

------------------------------------~-----------------

Period Covered:
Personnel:

April 1, 1974 to March 31, 1975

Oliver B. Cope, Margaret Stumpf and Warren D. Snyder.

ABSTRACT
Information derived from this study was prepared and published during this
Segment as follows:
Snyder, W. D. 1974. Pheasant use of roadsides for nesting in northeast
Colorado. Colorado Division of Wildlife Special Report #36. 24 p.
Snyder, W. D. 1974. Seeding roadsides for pheasant nesting cover.
Division of Wildlife Game Information Leaflet #99. 3 p.
Snyder, W. D.

1974.

The beleaguered ringneck.

Colo

.do

Colorado Outdoors 23(6) :5-8.

��-3-

PHEASANT ROADSIDE COVER EVALUATION STUDY
Warren D. Snyder

P. S. OBJECTIVE
To compare pheasant production under natural roadside conditions with the
following cover types to be established along roadsides: (1) grass, and
(2) grass-legume mixtures.

RESULTS AND DISCUSSION
Information derived from this study was prepared and published during the
work segment as follows:
Snyder, W. D. 1974. Pheasant use of roadsides for nesting in northeast
Colorado. Colorado Division of Wildlife Special Report #36. 24 p.
This report covers all aspects of the study with recommendations
ment use.

for manage-

Snyder, W. D. 1974. Seeding roadsides for pheasant nesting cover.
Division of Wildlife Game Information Leaflet #99. 3 p.

Colorado

This publication summarizes benefits and recommended procedures for use by
landowners and others in seeding roadsides to provide pheasant nesting cover.
Snyder, W. D.

1974.

The beleaguered

ringneck.

Colorado OutdOors 23(6):5-8.

This article briefly summarizes the ~light of the pheasant and proposes use
of roadsides as a partial answer to the problem of diminishing habitat.
A presentation summarizing findings of this study was presented at the.18th
annual meeting of the Midwest Pheasant Council in North Platte, Nebraska,
Ap ril 9, 1975.

ACKNOWLEDGEMENTS
Oliver B. Cope, editor, and Margaret Stumpf, typist handled their respective
duties with prompt and thoro~gh efficiency.
Their combined efforts were
invaluable in completing the above listed publications within this work
segment.

Prepared by

~"'-7JyJ

Warren D. Snyder
Wildlife Researcher

��April 1975

-5-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

~W~-~3~7_-~R~-=2~8

Work Plan No.

_

-=l__~-----------

Game Bird Survey
Job No.

21
----------~------------------

Job Title

Pheasant Mortality Investigation

Period Covered:

April 1, 1974 to March 31, 1975.

Personnel:

Howard Funk, David Bowden, Warren Snyder, Lucien Brevard, Larry
Crooks, Stanley Haskell, Gary Rafert, Steven Steinert, Bruce
Trindle, Daniel Walsworth, and Donald Hoffman.

ABSTRACT

Pheasant populations within the study area and within ,the prime pheasant range
in northeastern Colorado reached the lowest level in over 20 years during
1974, based upon census indices and hunter success. Data gathered during this
study indicated at least a 45 percent reduction in the spring 1975 breeding
population, in comparison with the spring 1974 breeding population due to
heavy losses from an extremely severe blizzard on March 27, 1975. A total of
534 pheasants were captured and banded within the four township study area
using nightlighting techniques during the fall period of 1974. A reward band
system was instigated to correct for non return of bands by hunters. Males
constituted 44 percent of the total fall banded sample and females 56 percent.
Thirty-six bands, including 23 reward bands, were returned by successful
hunters. Upon correcting band recovery data for non-reporting of non-reward
bands, it was estimated that hunting mortality on immature males was approximately 19 percent, 15 percent for mature males and 18 percent for all males.
Crippling loss on males was estimated at 10 percent of the retrieved bag for
both age classes, based upon hunter questionnaires. Thus, total hunting season
mortality through harvest and crippling loss was estimated at about 20 percent
for all males combined. This information also suggested that immature males
were about 1.3 times more vulnerable to shooting than adult males. A total
of 173 pheasants were captured during the winter period of 1975, again using
nightlighting techniques. Of these, 149 were newly banded and 24 were recaptured from the fall, 1974 banded sample. Males constituted 19 percent of
the total winter banded sample and females 81 percent. A total of 167 incidences of pheasant mortality were recorded within the study area from fall,
1974 through early spring, 1975. Banded males shot off the study area are
included in this sample. Incidences of mortality from the banded segment included: hunting (36),winter killed (9), road killed (1), and probable hunting
crippling loss (1). Incidences of mortality from the unbanded segment included:
hunting (62), winter killed (37), trapping casualties (9), road killed (6),
and unknown (6).

(Abstract continued on following page)

�-6ABSTRACT

(Continued)

Immature males banded during fall, 1974 and harvested during the open
season, 1974 dispersed more widely (average movement of 5.1 miles) than
mature males (average movement of 2.6 miles).
Twenty-six percent of the
males harvested, where movements could be determined, moved entirely off
the study area. In contrast, the winter, 1975 recapture and early spring,
1975 recovery samples of both males and females showed little movement
from the sites where originally banded in the fall of 1974. Based upon
this movement data gathered, this population of pheasants was considered
open rather than closed.

RECOMMENDATIONS

An important requirement for future statistical analyses is that hunter
success within the study area be sampled randomly. Random sampling will
allow for determination of confidence limits for popUlation estimates.
Random sampling through a hunting permit system is not recommended since
undue attention would be drawn to the study area.
A portion of this study depends upon marking and recapturing random
samples of birds for several years to estimate popUlations through changeill-ratio methods.
Statistical analyses were limited with only one year's
data. Preliminary data, however, indicated that sample sizes required to
estimate populations even with 25 percent accuracy may be unrealistically
large for this low density pheasant population.
This population is, however, representative of the northeastern Tablelands, long considered
Colorado's best pheasant population.
It is recommended that statistical
analyses be continued following the second year of data collection to
determine the reliability of the data being collected.

�-7-

PHEASANT MORTALITY

INVESTIGATION

Donald M. Hoffman

P. S. OBJECTIVE
To investigate mortality rates of a population of ring-necked pheasants.

SEGMEh~ OBJECTIVES
1.

Review relevant literature concerning pheasants and studies of mortality and survival of birds.

2.

Select study area.

3.

Monitor population status on selected areas.

4.

Develop new and/or improve existing trapping techniques.

5.

Investigate mortality factors and estimate mortality rates of marked
individuals.

6.

Prepare annual progress report.

METHODS AND MATERIALS
A search of literature was made and brief abstracts of pertinent articles
were filed on 5 by 8 inch "Unisort" cards.
Field surveys of several areas within the prime pheasant range in the northeastern Tablelands were conducted along with preliminary population monitoring
using crow counts and roadside counts. A study area was selected during the
sun er of 1974 on the basis of representative pheasant populations, landowner
cooperation, and habitat types (extensive wheat stubb1efields) needed for
nightlighting-trapping operations.
Pheasant populations within the selected study area were monitored through
crow counts made along one trend route from April 25 to June 4, 1974, brood
counts along one trend route from August 7 to August 28, 1974, and sex-ratio
counts made in various sites within the study area from January 29 to March
20, E 75.

Pheasants were captured using nightlighting techniques developed in Illinois
(Labisky 1968). Two crews of two men each and two 4-wheel drive vehicles
were used for a portion of this work and one crew of two men with one 4-wheel
drive vehicle for the remainder. All pheasants were leg banded with aluminum
colored bands and approximately one half were also leg banded with gold colored
aluminum reward bands. An intensive effort was made in January, 1975 to
attract pheasants to whole shelled corn baitsites for winter period bait
trapping attempts with little success. Abundant waste grain in the large
wheat stubblefields combined with an open winter were considered to be major

�-8-

factors why the birds did not readily utilize the bait stations.
This
method was therefore discarded in favor of winter period nightlighting
trapping.
Banding efforts were concentrated from mid-August to mid-October, 1974
(pre-season) and from early February to late March, 1975 (post-season).
Brood counts indicated the average age of the young birds was approximately ten weeks in mid-August, 1974. Pheasants were aged by the length
and shaft diameter of the first primary according to Wishart (1969).
Hunting mortality data were collected through field contacts of hunters,
check station operations, and hunter questionnaires.
Hunters reporting
banded birds were surveyed to obtain banded-unbanded ratios of birds in
the bag, success and number days afield. They were also asked to supply
names and addresses of their hunting companions and these, too, were surveyed to obtain additional data. The reward given a hunter for returning
a gold colored reward band and the corresponding aluminum colored regular
band during 1974 was a 144 page book Cooking the Sportsmen's Harvest published by the South Dakota Department of Game, Fish, and Parks.
Non-hunting mortalities of both banded and unbanded birds were documented
by date, age, sex, and cause of mortality whenever possible.
Field searches
of wheat stubblefields, draws, windrows, old homestead sites, and along roadways were made by four men, using three bird dogs and two 4-wheel drive
vehicles following a severe blizzard in late March, 1975 to secure winter
kill mortality information.
Data were assembled and compiled.
Hunting mortality rates utilizing fall
banding data, including reward and non-reward band recoveries, were estimated.
RESULTS AND DISCUSSION
Selection

of Study Area

A study area of four townships in size and located in southwestern Phillips
County (Fig. 1) was selected following preliminary population monitoring,
landowner contacts, habitat inspection, and conferences with Wildlife Researcher
W. Snyder and Wildlife Conservation Officer T. Lines. The western border of this
study area is the Logan County line and the southern border is 'the Yuma County
line. Haxtun, Colorado, located two miles north of the northern boundary, was
used for a base of operations during 1974. Pheasant populations for 1974 proved to be highest in the two southern townships of the study area.
Crow Counts, 1974
A tally of the highest number of crows heard per two minute period during four
counts made along a 19.5 mile trend route (Haxtun-South) within the study area
showed a mean of 20.6 calls compared with a mean of 26.0 calls recorded by
Wildlife Conservation Officer T. Lines along the Wages-Clarkville-Haxtun
Management Trend Route (Table 1). Crow count indices show a continued decline
of pheasant numbers within the study area and also within the prime northeastern pheasant range of Colorado.
Snyder (1974b) stated the 1973-74
pheasant population level in northeastern Colorado was the lowest in over 20
years.

�1 7 N.

(9

o

R.t't 7 VI.

F&lt;.46W

11

YUMA

Fig. 1. Location of study area, consisting
County, is outlined in solid black line.

COUNTy

of four townships in southw stern Phillips

�-10-

Brood Counts,

1974

Table 2 summarizes brood counts completed within the study area for 1974.
An average of 1.86 birds per mile was tallied in four counts along the
Haxtun-South trend route compared with 1.89 birds per mile tallied by W.
Snyder in three counts along the nearby Wages-Clarkville-Haxtun
Management
Trend Route. Of 31 hens tallied along the Haxtun-South route, 20 (65%)
had broods and 11 (35%) apparently were unsuccessful.
An average of 5.75
young per brood was tallied along this route.

Pheasants

Banded on Study Area, Fall, 1974

Table 3 summarizes data from 534 pheasants captured and banded on the study
area using nightlighting techniques during the fall period of 1974. Males
constituted 44 percent of the fall sample and females 56 percent.
A young:
adult ratio of 1.80:1.00 was suggested from this trapping sample with an
indicated young:mature hen ratio of 2.40:1.00.
A male pheasant being
captured with a hand net using nightlighting equipment is shown in Figure 2.
A view of a wheat stubblefield intersected by a draw which was trapped during
both fall and winter night-trapping operations is shown in Figure 3.

Pheasant

Season, 1974

A regular open season was held extending from noon on November 16 through
November 28, 1974 (12.5 days) and an extended season from November 29
through December 15, 1974 (17.0 days) with a bag limit of three males and a
possession limit of six (Table 4). Numbers of hunters (Fig. 4) were tallied
periodically along roadsides (Table 5). These counts indicated that most
hunters hunted during the opening weekend only and hunting pressure-was
minimal during the remainder of the season. Success in the general area
was poor on opening day (0.25 birds per hunter at the Fleming Check Station),
apparently resulting in many hunters losing interest early and many not
returning.
A maximum average of two hunters per square mile in the study
area indicated only a fair amount of hunting pressure, even during the opening
day of the 1974 season. Hunter questionnaire data suggested that the average
time spent per hunter in the study area was 2.1 days. However, the sample of
hunters was biased toward the successful hunter and the time spent was also
probably biased on the high side.
Thirty-six total bands, of which 23 were reward bands, were returned or recovered from successful hunters (Table 6). This information indicates a
discrepancy in reward to non-reward band ratios according to method in which
the bands were obtained.
Of 13 bands secured from hunters through field
contacts or at the Fleming Check Station, nine were regular bands and four
were reward bands. Of 23 bands mailed or brought in voluntarily by hunters,
four were regular bands and 19 were reward bands. A significant difference
(10.63; P&lt;O.Ol) was determined, using a chi-square goodness of fit test,
comparing the number of reward to non-reward bands collected by field or
check station contacts with those voluntarily returned by hunters.
As shown
in Table 3, about half of the immature and 69 percent of the adult males were
reward-banded.
However, there were more immatures banded than adults. Thus,
the rate of encountering reward to non-reward bands would be expected to be
close to 50-50 either in the field or for those reported by hunters through
the mail.

�Table 1.

Summary of pheasant

Route

Haxtun-South

Wages-ClarkvilleHaxtun 1/

crow coun r s , 1974.

Period of
Counts

No. of
Counts

No. of
Stations

Tally of
Highest Count
Per Station

Mean

Cocks

4-25-74 6-4-74

4

10

206

20.6

11

4-23-74 6-5-74

3

10

260

26.0

Observed During Counts
Sex-Ratio
Hens
.48 : 1.00
(Males : Females)

23

Data Not Recorded

1/
- Counts were made by Wildlife

Conservation

Officer T. Lines along this established

Pheasant

Management

Route.
I

f-'
f-'

Table 2.

Route

Haxtun-South

Wages-Clarkvil1eHaxtun 1/

1/

I

Summary of pheasant brood counts, 1974.
Observed During Counts
Sex Ratio
UnYoung/
No.
Brood Adults (M : F)
Broods
class. Total

Period of
Counts

No. of
Counts

Total
Miles

Average
Birds/
Mile

Cocks

Hens

Young

8-7-74 8-28-74

4

98.3

1.86

10

31

115

27

183

20

5.75

.32 : 1.00

3

91.0

1.89

11

32

126

3

172

25

5.04

.34 : 1. 00

8-6-74 8-20-74

- Counts were made by Wildlife

Researcher

W. D. Snyder along this established

Pheasant

Management

Route.

�Table 3.

Pheasants banded on study area, August 21, 1974 through October 16, 1974.

Age

Sex

Number Banded

Percentage

Number Reward Banded

Percentage

Juvenile

Male

185

34.64

93

50.27

Male

48

8.99

33

68.75

Juvenile

Female

158

29.59

64

40.51

Mature

Female

143

26.78

66

46.15

534

100.00

256

Mature

Totals
Average

47.94
I
•...
N

Summary:

I

Young: 'Adult Ratio = 343 : 191 or 1.80 : 1.00
Young: Mature Hen Ratio = 343 : 143 or 2.40 : 1.00
Percent Males in Total
= 43.63
Percent Females in Total = 56.37

Table 4.

Summary of pheasant hunting season pertaining to study area, 1974.
Length
(Days)

Bag
Limit

Possession
Limit

11-16-74 - 11-28-74 (regular)

12.5

3 (males only)

6 (males only)

11-16-74 Only-Noon to Sunset
All Other Days - Sunrise to
Sunset

11-29-74 - 12-15-74 (extended)

17.0

3 (males only)

6 (males only)

Sunrise to Sunset

Dates

Hours

�-13-

Fig. 2. A total of 683 pheasants, including 534 in the fall and 149 in the winter,
were captured using night lighting techniques and banded with aluminum leg bands
during the first year of the study. Approximately one half were banded with reward
bands.

�I

I-'
+:I

Fig. 3. A wheat stubblefield intersected by a draw, located within the study area. This field was trapped
during both fall and winter nightlighting-trapping
operations.
The numerous draws provide drainage to the
land and cover and water for pheasants.

�I
•....•
I.n
I

Fig. 4.

Pheasant hunters searching a wheat stubblefield

during open season, November, 1974.

�Table 5. Pheasant hunter counts in study area, 1974.

Date

Time

No.
Miles

No. Sq.
Miles 1/

No.
Hunter
Vehicles

Ntnnber
Hunter
Parties

Ave. No.
Hunters/
Vehicle

No. Hunters
in
Sample

No. Hunters
in 144 Sq.Mi.
Study Area

Counted
By

11-16-74
(Saturday)

12:15 PM1:15 PM

31.0

31

26

20

2.43

63

293

D. Hoffman

11-17-74
(Sunday)

8:45 AM10:30 AM

31.0

31

6

6

3.00

18

84

D. Hoffman

11-21-74
(Thursday)

8:45 AM9:45 AM

31.0

31

0

0

--

0

0

B. Trind1e

11-23-74
(Saturday)

8:30 AM11: 30 AM

58.0

58

2

2

2.00

4

10

w. Snyder

I
•....

aI

11-23-74
(Saturday)

12:15 PM1:30 PM

31.0

31

0

0

--

0

0

D. Hoffman

11-24-74
(Sunday)

8:10 AM9:45 AM

31.0

31

0

0

--

0

0

D. Hoffman

1/
- Hunters and vehicles tallied one-half mile on each side of route.

�-17-

Table 6.

Band recovery information, 1975 season.

Recovered

Bands

Non-Reward
Bands

Field Checks and
Fleming Check Station

13

4

9

Reported by Hunters

23

19

4

Totals

36

23

13

Source

Even with the possibility of bias as brought out in Table 6, the band recovery
information has to be the best data to utilize in estimating percent of the
male population harvested.
The purpose of the reward band system is to obtain
a non-reporting rate for harvested, non-reward banded birds so that total
banded birds harvested can be estimated. This method includes the assumption
that all reward bands from harvested birds are reported by hunters. Therefore, non-reward bands were corrected for non-reporting on the basis of returned reward bands utilizing information on numbers banded with reward and
non-reward bands by immature and adult age class from Table 3.
The projections for correcting for non-reporting are illustrated in Table 7.
For immature males, 18 reward and 12 lion-reward bands were obtained from
hunters. Because immatures were banded at very close to a 50-50 ratio with
reward and non-reward bands, the reporting rate of non-reward bands was indicated to be only about 67 percent. Correcting for this, the tQtal number
of recovered bands for immatures was estimated at 36 rather than 30. Reward
bands were placed on 69 percent of the adults so the projection for non-reward
bands was estimated at 2 with a total estimated at 7 rather than 6. Thus,
the estimate of total banded birds harvested and retrieved by hunters was 43
rather than 36.

Table 7. Projection estimates for non-reporting
age class.

No.
Banded

Immature

Reported
Recovered
Reward Non-Reward

of banded males harvested by

Projected
Non-Reward
Bands

Total
Recoveries

Percent
Recovered

185

18

12

18

36

19.5

Adult

48

,?

1

2

7

14.6

Totals

233

23

13

20

43

Average

18.5

�-18-

Data in Table 7 suggest that 19.5 percent of the immature segment of the
male population was harvested and retrieved by hunters and 14.6 percent
of the adult male population.
This averages 18.5 percent for all males.
Comparison of the recovery rates by age also suggests the immatures were
1.34 times as vulnerable to hunting as adults (19.5 + 14.6).
Hunter questionnaire information suggested that crippling loss was about
10.3 percent of the retrieved bag. This would add 1.9 percent mortality
to the average computed for all males harvested (18.5 percent) making the
estimate of all mortality due to hunting total 20.4 percent. An apparent
hunting season crippling loss (male) is shown in Figure 5.
All successful hunters returning bands indicated they had killed only one
banded male each, even though some killed as many as six males during the
season. This was considered very unlikely, considering the number of banded
.males in the study area. It is also of interest that no bands from female
pheasants were returned by hunters even though 301 were banded and reward
bands were placed on 43 percent of these.
Hunters who were sent questionnaires following the close of the 1974 season,
were asked to comment on the season. Of 52 hunters who returned their
questionnaires, 26 (50%) made no comment.
Of the 26 hunters who made comments,
22 (85%) indicated the hunting was poor, and only 4 (15%) indicated the hunting
was good. Several hunters commented on the deterioration of habitat and low
populations of pheasants compared with previous years. Habitat deterioration
and declining pheasant populations are occurring throughout its entire range
(Snyder 1974a), so this trend is not unique to Colorado.

Pheasants

Banded on Study Area, Winter, 1975

Table 8 summarizes data from 173 pheasants captured on the study area during
the winter of 1975. Nightlighting-trapping
techniques were again used during
this period. Of these, 149 were newly banded birds, of which 75 were also
banded with reward bands, and 24 were recaptures from the fall, 1974 period.
Utilizing newly banded and recaptured birds, a sex ratio (males:females) of
0.23:1.00 was determined for the period and the young:adult ratio was determined to be 0.88:1.00.

Winter Sex-Ratio Counts, 1975
Winter period sex ratio counts completed in January through March, 1975 by
searching draws, windrows, old homesites, and recording birds along roadsides
are listed in Table 9. A sample of 262 pheasants (36 males, 179 females, and
47 unclassified) indicated a sex ratio (M:F) of 0.20:1.00 for the study area.
This compares with a sex ratio of 0.23:1.00 for the winter trapping sample
(Table 6). Thus, males which appeared to be more difficult to capture than
females when fully grown, were actually trapped at a slightly higher rate
than observed in the wild population.
Males were, however, somewhat easier
to observe than hens during the nightlighting-trapping
operations because of
their coloration.

�I

I-"

\0

I

Fig. 5. A male pheasant carcass found following the 1974 open season. Crippling loss of males was estimated
at ten percent of the retrieved bag from a questionnaire survey of hunters.

�Table 8.

Pheasants banded on study area, February 5, 1975 through March 20, 1975.

Age

Sex

Number Banded

Percentage

Number Reward Banded

Percentage

Juvenile

Male

18

10.40

10

55.56

Recaptures 1./

2

1.16

1

50.00

Sub-totals-

20

11.56

11

10

5.78

9

90.00

Recaptures 1./

2

1.16

0

0.00

Sub-totals

12

6.94

9

Mature

Male

I
N

Juvenile

Female

53

30.64

23

43.40

Recaptures 1./

8

4.62

3

37.50

Sub-totals

61

35.26

26

68

39.30

33

48.53

Recaptures 1./

12

6.94

5

41.67

Sub-totals

80

46.24

38

173

100.00

84

Mature

Female

Totals
Sunnnary:

Young:Adult Ratio = 71 : 78 or 0.91 : 1.00; W/Recaptures = 81 : 92 or 0.88 : 1.00
Young:Adult Female Ratio = 71 : 68 or 1.04 : 1.00; W/Recaptures = 81 : 80 or 1.01 : 1.00
Sex Ratio (Males: Females) = 28 : 121 or 0.23 : 1.00; W/Recaptures = 32 : 141 or 0.23 : 1.00
Percent Males in Total = 18.79; W/Recaptures = 18.50
1/

- Included only birds recaptured from Summer, 1974 banding sample.

o
I

�Table 9.

Pheasant

sex-ratio counts obtained during daytime by date and cover type, winter, 1975.

Cover Type

No.
Males

No.
Females

No.
Unc1ass.

Total

2

Draw Bottom

20

115

45

180

0

1

Roadside

10

24

0

34

2

5

8

Old Homesite
and/or Windrow

6

40

2

48

2

5

0

7

2-11-75

2

12

0

14

Totals

36

179

47

262

2-12-75

0

5

0

5

2-13-75

2

0

0

2

No.
Males

No.
Females

No.
Unc1ass.

Total

1-29-75

1

1

0

1-30-75

0

1

1-31-75

1

2-6-75

Date

I
N

~

Summary:
2-18-75

5

48

20

73

2-19-75

5

18

7

30

2-20-75

5

14

8

27

3-3-75

5

33

5

43

3-6-75

2

0

0

2

3-18-75

3

25

1

29

3-19-75

0

3

0

3

3-20-75

3

12

1

16

Totals

36

179

47

262

I

Male

-----.

: Female Ratio
.20 : 1.00

= 36 : 179 or

�-22-

Ratios: Banded toUribartded aridYotirigto Adtilt
Information gathered on banded to unbanded ratios is listed in Table 10.
Hunter band returns plus hunter questionnaires indicated a banded to unbanded ratio of 1.00:3.35 for males during the open season, 1974. This
estimate was, however, secured from a non-random sampling of hunters and
was not used in statistical analyses.
The banded to unbanded ratio for the entire winter trapping period was
1.00:6.21, with recaptures from the winter period excluded, and 1.00:2.29
for the final two weeks of winter trapping.
This last ratio included recaptures from both the fall, 1974 and recaptures during the final two weeks
from the winter, 1975 periods and indicated approximately 30 percent of the
wintering population was banded.
The banded to unbanded ratio from 46 carcasses found within the study area
following the March 27, 1975 blizzard was 1.00:4.11 and indicated approximately 20 percent of the wintering population was banded.
Age ratio information gathered during the first year of the study is listed
in Table 11. The fall, 1974 banding sample indicated a ratio (young:adult)
of 1.80:100 compared with an indicated ratio of 0.91:100 for the winter,
1975 banding sample.

Losses from an Early Spring, 1975 Blizzard
Freezing, driving rains with high winds, followed by drifting snow may result in high losses of pheasants through their freezing to death, according
to Edminster (1954). Such was the unusual weather combination which occurred
in northeastern Colorado in late March, 1975. A very severe blizzard occurred
on March 27, 1975, but this was preceded by driving rain during the night of
March 26, 1975. Counts of live birds made, in known concentration sites within
the study area before and after the storm indicated approximately one half
of the pheasants died. Forty-six carcasses, of which nine were banded, were
found on the study area following the storm (Fig. 6 and Tables 12 and 13).
Eleven additional pheasant carcasses (4 immature males, 3 immature females,
2 mature females, and 2 unclassified females) were found within one-half mile
of the north boundary, but outside the study area. Predators such as hawks,
owls, and coyotes were more efficient searchers than a crew of four men with
three bird dogs and two 4-wheel drive vehicles, so the extent of the losses
could not be measured on~y through numbers of carcasses found. Also, continuing inclement weather closed secondary roads and prevented field searches
for almost a week ,
Figures 7 through 9 are included to show effects of this storm on habitat,
cattle, and Hildlife.
Many individuals considered this storm to be the most
severe since the January, 1949 blizzard, which also caused heavy losses to
pheasant populations of northeastern Colorado.

�Table 10.

Banded to unbanded

rat '.os of pheasants

by period.

Banded/Unbanded
Ratio

Period

11-16-74 to 12-15-74
(Open Season, 1974)

26 : 87 or 1.00 : 3.35

2-5-75 to 3-20-75
(Entire Winter Trapping
Period)

24 : 149 or 1.00 : 6.21-

3-10-75 to 3-20-75
(Final Two Weeks of
Winter Trapping Period)

Estimate
Percentage

1/

2/
21 : 48 or 1.00 : 2.29 -

of
Banded

How Obtained

23.01

Hunter band returns plus
hunter questionnaires

13.87

Winter trapping

(night1ighting)

30.43

Winter

(nightlighting)

trapping

I

N

W
I

4-2-75 to 4-10-75
(Following the March 27,
1975 Blizzard)

9 : 37 or 1.00 : 4.11

19.57

Tally of winter killed pheasants
following March 27, 1975
blizzard (on study area).

1/
- Ratio excluded recaptures from winter period since these were tallied during final two weeks of trapping
only. Twenty four recaptures from fall, 1974 period used to determine ratio.

2/
- Ratio included

8 recaptures

from fall, 1974 period plus 13 recaptures

from winter trapping

period.

�Table 11.

Young to adult ratios of pheasants by period.
Number
Young

Number
Adults

Total

Percent
Adults

Fall Banded, 1974

343

191

534

35.77

1.80 : 1.00

Open Season, 1974
(Banded Sample, Cocks)

30

6

36

16.67

5.00 : 1.00

Winter Banded, 1975

71

78

149

52.35

0.91 : 1.00

Sample

Young : Adult Ratio

I
N

+:I

�I
N
\J1

I

Fig. 6. One of 46 pheasant carcasses found within the study area following a blizzard on March 27, 1975.
Approximately one half of the pheasant population of the northeastern Tablelands died during this storm.

�Table 12.

Incidences of pheasant mortality from band returns, fall, 1974 through early spring, 1975.

Date

Aluminum
Band No.

Gold
Band No.

Sex

Age

9-10-74

5089

0070

Male

Immature

Road Killed

5005
5017
5021
5030
5064
5066
5082
5083
5085
5116
5120
5134
5137
5160
5185
5191
5227
5266
5276
5289
5307
5332
5340
5344
5348
5358
5367
5390
5395

----

Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male

Immature
Immature
Immature
Mature
Immature
Immature
Immature
Immature
Immature
Immature
Immature
Mature
Immature
Immature
Mature
Immature
Immature
Immature
Immature
Immature
Immature
Immature
Mature
Immature
Immature
Immature
Immature
Mature
Immature

Hunting (Open Season-1974)

11-16-74 to
3-24-75

--

--

--

--

0067
0068
0033
0034
0040
0042

-0093
0096
0113
0158
0163

--

0179
0253
0190
0192
0194
0129

--

0145
0148

Cause of Mortality

"
"
"

"

"

"
"
"
"
"

"

"

"
"
"
"

"

"

"

"
"
"

"

"

"

"

"
"

"
"

"

"
"
"
"

"
"

"

"

"

"

"

"
"

"
"

"

"

"

"

"

"
"

"

"
"

---------------------------------------------------------_._--------------------------------------------------

I
N
C1\
I

�Table 12. Incidences of pheasant mortality from band returns, fall, 1974 through early spring, 1975
(continued).

Date

Aluminum
Band No.

Gold
Band No.

5400
5401
5412
5415
5497
5505
5520

0150
0256

11-16-74 to
3-24-75 (cant)

1-21-75

5004

0217
0274

0002

Sex

Age

Cause of Mortality

Male
Male
Male
Male
Male
Male
Male

Immature
Immature
Immature
Immature
Mature
Immature
Immature

Hunting (Open Season-1974)

Male

Mature

Crippling Loss (Open Season-1974)

"

"

"
"
"
"

"
"

"
"

"

"

I

4-2-75 to
4-10-75

5055
5154
5516
5531
5591
5602
5610
5625
5655

0053

Male
Female
Male
Male
Female
Male
Female
Male
Female

Immature
Immature
Immature
Immature
Mature
Immature
Immature
Mature
Immature

Winter Killed (3-27-75 Blizzard)

"

"
"

"
"
"

"

"

"

Number

Percent

Legal Hunting (Harvest 36 and Crippling Loss 1)
Winter Killed (March 27, 1975 Blizzard)
Road Killed (on Paved Highway)

37
9
1

78.72
19.15
2.13

Total

47

100.00

0230
0291
0295
0310
0304

"
"
"

"

"
"

"

Summary:
Cause of Mortality

_-_.--------------------------------------------

_ ..

..

'

N
.•....•

I

�-28-

Table 13. Incidences of pheasant mortality
through early spring, 1975.

Date (s)

Sex

Age

No.

8-21-74
8-21-74
8-28-74
9-10-74
9-10-74
9-12-74
9-17-74
9-18-74
9-19-74
9-19-74
10-3-74
11-16-74 to
12-15-74

Male
Unk.
Female
Male
Male
Male
Male
Female
Female
Male
Female

Immature
Immature
Immature
Immature
Immature
Immature
Immature
Immature
Unknown
Immature
Immature

1

Male

11-23-74
12-12-74
1-29-75
1-30-75
2:...12-75
2-12-75
2-13-75
2-20-75
3-11-75
3-18-75
3-20-75
4-2-75 to
4-10-75

from unbanded birds, fall, 1974

Cause of
Mortality

Remarks

1
1
1

Road Killed
Trapping Casualty
Road Killed
Trapping Casualty
Trapping Casualty
Road Killed
Trapping Casualty
Trapping Casualty
Trapping Casualty
Road Killed
Road Killed

Found on Paved
Stepped On
Found on Paved
Suffocated
Heart Attack
Found on Paved
Heart Attack
Suffocated
Hit with Net
Found on Paved
Found on Paved

Unknown

61

Legal Hunting

Male
Male
Unk.
Unk.
Female
Male
Male
Unk.
Female
Female
Female

Unknown
Immature
Unknown
Unknown
Unknown
Unknown
Unknown
Immature
Mature
Mature
Mature

1

Legal Hunting
Road Killed
Unknown
Unknown
Unknown
Unknown
Unknown
Unknown
Trapping Casualty
Trapping Casualty
Trapping Casualty

Hunter Contacts and/or
Questionnaires
Probable Crippling Loss
Found on Paved Highway
Feathers Only Found
Feathers Only Found
Feathers Only Found
Feathers Only Found
Feathers Only Found
Feathers Only Found
Hit With Net
Heart Attack
Heart Attack

Male
Male
Male
Female
Female
Female

Immature
Mature
Unc1ass.
Immature
Mature
Unc1ass.

Winter
Winter
Winter
Winter
Winter
Winter

March
March
March
March
March
March

1
1
1
1
1

1
1

1

1
1

1
1
1
1
1
1

1
11
7
1

5
4
9

Killed
Killed
Killed
Killed
Killed
Killed

27,
27,
27,
27,
27,
27,

1975
1975
1975
1975
1975
1975

Highway
Highway

Highway

Highway
Highway

Blizzard
Blizzard
Blizzard
Blizzard
Blizzard
Blizzard

Summary:
Cause of Mortality
Legal Hunting (Harvest 61 and Crippling Loss 1)
Winter Killed (March 27, 1975 Blizzard)
Trapping Casualties (Various Causes)
Road Killed (All on Paved Highways)
Unknown (Feathers Only Found)
Total

Number

Percent

62
37

6

51.67
30.83
7.50
5.00
5.00

120

100.00

9

6

�I
N
10

I

Fig. 7. Sites such as shelterbelt tree plantings, draws, and wheat stubblefields, which normally serve as
winter cover for pheasants, filled with snow drifts. The birds were then forced into a variety of exposed
situations, resulting in heavy losses.

�I
W

o
I

Fig. 8. Secondary roads were still in the process of being opened within the study area one week after the
blizzard had ended.

�I
W

•.....
I

Fig. 9. County Agents estimated 19,000 cattle (mostly calves and yearlings) were killed outright in northeastern Colorado by the severe blizzard. Pheasants, cottontails, jackrabbits, hawks (marsh, rough-legged,
and sparrow), crows, and meadowlarks also suffered losses from the storm.

�-32-

InstartcesofMotta1ity
Information on pheasant mortality obtained through bands
covered is listed in Table 12. Numbers tallied by cause
included: hunting during open season (36), winter killed
1975 blizzard (9), road killed (1), and probable hunting
loss (1).

returned or reof mortality
from March 27,
season crippling

Information on pheasant mortality obtained from unbanded birds is listed
in Table 13. Numbers tallied by cause of mortality included: hunting
during open season (61), winter killed from March 27, 1975 blizzard (37),
trapping casualties (9), road killed (6), unknown (6), and probable hunting
season crippling loss (1).

Movements

of Banded Pheasants

Thirty-three fall, 1974 banded males (both immature and mature) moved an
average of 4.7 miles (range&lt;l to 12 miles) from the site of banding prior
to the open season (Table 14). Twenty-six percent of the males (8 immature
and 1 mature) moved more than one mile off the study area. By age class,
27 immature males moved an average of 5.1 miles (range&lt;l to 12 miles) and
six mature males moved an average of 2.6 miles (range &lt;1 to 7.8 miles).
Thus, immature males were found to disperse more widely than mature males
from the various banding sites. Because of the movement noted, particularly
with immature males, this population of pheasants was considered as open in
statistical analyses.
In sharp contrast, the sample of both male and female birds (a) banded during
the fall, 1974 and recaptured during the following winter period or (b) banded
during either the fall, 1974 or winter, 1975 periods and recovered 4uring the
early spring period following the March 27, 1975 blizzard showed far less
movements (Table 15). Twenty-two of the 33 birds tallied (67%) were recaptured or recovered in the same section where banded and only two (one
immature female and one immature male) moved more than two miles from the
section where banded.
The reason for this, of course, has to be that winter
banding and the search for dead birds after the storm were accomplished only
on the study area and reflected the movements of only those which stayed on
the study area after fall banding.

Harvest and Population

Estimates

When this study was initiated, the plan was to trap and band pre-season and
post-season to allow, not only estimates of percentages of males taken by
the gun, but also to estimate survival from fall to winter through recapture
data. As per Table 3, 233 males were banded pre-season which was not sufficient
to give the total of 50 recoveries hoped for to allow more confidence in
estimates of percent of the male population taken by hunting.
However, good
experience was obtained as far as trapping methods and what to expect in the
way of hunting pressure and sample sizes needed.

�-33-

Table 14. Movements of 34 male pheasants, banded in the fall, 1974 and
harvested in open season, 1974.
Aluminum
Band No.
5005
5017
5021
5030
5064
5066
5082
5083
5085
5116
5120
5134
5137
5160
5185
5191
5227
5266
5276
5289
5307
5332
5340
5348
5358
5367
5390
5395
1400

Gold
Band No.

0067
0068
0033
0034
0040
0042
0093
0096
0113
0158
0163
0179
0253
0190
0194
0129
0145
0148
0150

.1.2
!~:~

5415
5497
5505
5520

0217
0274

Sex

Age

Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male
Male

Immature
Immature
Immature
Mature
Immature
Immature
Immature
Immature
Immature
Immature
Immature
Mature
Immature
Immature
Mature
Immature
Immature
Immature
Immature
Immature
Immature
Immature
Mature
Immature
Immature
Immature
Mature
Immature
Immature
Immature
Immature
Mature
Immature
Immature

Moved from Section Banded
Distance (mi.)
Direction
1.4
NE
1.4
SW
4.4
SSW
4.0
S
1.4
SE
4.0
SE
10.3
ESE
10.3
ESE
2.0
N
10.0
liE (off study area)
(Killed in Logan County)- NW (off study area)
2.0
N
6.2
ESE
10.0
SW (off study area)
7.8
WNW (off study area)
8.4
E
12.0
SSE (off study area)
(Killed in same section)
{Killed in same section}
1.0
N
8.3
NE
1.4
SE
1.0
W
8.2
WSW {off study area}
1.0
S
6.9
ESE
{Killed in same section}
10.6
SSE (off study area)
7.9
NE (off study area)
(Killed in same section)
1.8
SSE
1.0
NE
8~5
ENE (off study area)
1;4
NE

Summary:
33 males of both mature and immature ages moved an average of 4.7
miles (range &lt; 1 to 12.0 miles)
6 mature males moved an average of 2.6 miles (r'ange-C'l,
to 7.8 miles)
27 immature males moved an average of 5.1 miles (range &lt; 1 to 12.0 miles)
9 of 34 (26%) of the males (1 mature and 8 immature) moved more than
1 mile off the study area
Direction of movements was not consistent but the majority moved
generally southeastward from banding sites.
1/
- Location of this bird not listed other than killed in Logan County, south of
Fleming.

�-34-

Table 15. Movements of 33 banded pheasants, recaptured
1975 or recovered during the early spring, 1975.
Aluminum
Band No.

Gold
Band No.

Recaptured

During Winter Trapping:

5032
5036
5055
5088
5105
5131
5132
5155
5157
5318
5347
5349
5357
5417
5422
5432
5443
5445
5469
5485
5486
5488
5506
5523
Recovered
5055
5154
5516
5531
5591
5602
5610
5625
5655

Sex

during the winter,

Moved from Section Banded
Distance (mi.)
Direction

Age

!/

Female
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Male
Female
Female
Female
Female
Female
Male
Female
Female
Female

Mature
Immature
Immature
Mature
Innnature
Mature
Mature
Innnature
Innnature
Mature
Mature
Immature
Innnature
Immature
Mature
Mature
Innnature
Mature
Innnature
Mature
Mature
Mature
Mature
Mature

Male
Female
Male
Male
Female
Male
Female
Male
Female

Immature
Immature
Innnature
Innnature
Mature
Innnature
Immature
Mature
Immature

(Recaptured in same section)
1.0
S
0053
2.0
N
(Recaptured in same section)
0028
(Recaptured in same section)
(Recaptured in same section)
0039
(Recaptured in same section)
0078
(Recaptured in same section)
0079
(Recaptured in same section)
(Recaptured in same section)
1.0
S
(Recaptured in same section)
1.0
S
(Recaptured in same section)
(Recaptured in same section)
(Recaptured in same section)
2.5
NW
(Recaptured in same section)
(Recaptured in same section)
0211
(Recaptured in same section)
(Recaptured in same section)
0213
(Recaptured in same section)
0275
(Recaptured in same section)
0226
1.4
SE
(dead) Following March 27, 1975 Blizzard: 2:.../
0053

0230
0291
0295
0310
0304

2.0
1.0
(Recovered
7.5
2.0
2.0
(Recovered
(Recovered
(Recovered

S
S
in same section)
NNE
N
E
in same section)
in same section)
in same section)

Summary:
22 of the 33 birds (67%) were recaptured or recovered in same section
where banded.
9 of 11 birds which moved (82%), moved 2.0 miles or less from section
where banded.
1 innnature female moved 2.5 miles northwest and 1 immature male moved
7.5 miles north by northeast.
l/These

pheasants were banded during fall, 1974 period.

J:.../Aluminum band numbers 5055 through 5531 were banded during fall, 1974 and
aluminum band numbers 5591 through 5655 were banded during winter, 1975.

�-35-

The winter banding period was not as fruitful as the fall period in
several ways. Trappedmalestlumberedonly
32,. of which only 4 were retrapped from the fall period (Table 8). Thus, males were not captured
in the same ratio to females as in fall, possibly reflecting differential
movement by sex, and the sample sizes were considered too low to enable
trap-retrap estimates to estimate survival of males from fall to winter.
Since trapping was accOmplished only on the study area, there was no way
to estimate how many banded males had left the area and no' way to estimate
how many unbanded birds had come into the study area from fall to winter.
Therefore, attempts to estimate survival will have to wait until results
of the pre-season banding period in 1975 are obtained.
Also, with so
little data to work with, attempts to estimate the pre-season population
of males produced such variable results that they were abandoned at this
time.

LITERATURE

CITED

Edminster, F. C. 1954. American game birds of field and forest:their
habits, ecology and management.
Chas. Scribner's Sons, N. Y. pp. 1-57.
Labisky, R. F. 1968. Nightlighting: its use in capturing pheasants,
prairie chickens, bobwhites, and cottontails.
BioI. Notes No. 62.
Illinois Natural History Survey, Urbana.
l2p.
Snyder, W. D.

1974a.

The beleaguered

ringneck.

Colo. Outdoors

23(6):5-8.

1974b. Pheasant use of roadsides for nesting in northeast Colorado.
Spec. Report No. 36. Colo. Division of Wildlife, Denver.
24p.
Wishart, W. 1969. Age determination of pheasants by measurement
primaries.
J. Wildl. Manage. 33(3):714-717.

Prepared by

(~?!l~a.-u
Donald
• Hoff~
Wildlife Researcher

of proximal

��April 1975

JOB PROGRESS REPORT

State of

~C~OL~O=RAD~~O

Project No.

_

W~-~3~7~-~R~-~2~8~ _

Game Bird Survey

Work Plan No.
Job Title.

~~3----~~--~Job No.~------------~8~a-------------Effects of Sagebrush Control
_
~o~n~·~D~l~·s~t~r~i~b~u~t~i~o~n~an~d~~A~b~un~d~an~c~e~·~o~f~S~a~g~e~G~r~o~u~s~e~

Period Covered:

April 1, 1974 to March 31, 1975

Personnel: Fred Giese and Kenneth Russell, U.S.F.W.S.; Thomas Beck, Don Benson,
Clait Braun, Lonnie Brown, Thomas Campbell, Jack Corey, Heather Flanagan, Howard
Funk, Don Gore, John Hobbs, Jim Jackson, Bruce McCloskey, Mike McLain, Ken
Miller, Sig Palm, Steve Porter, Howard Spear, John Wagner, Colorado Division of
Wildlife.
ABSTRACT
In all, 172 sage grouse (Centrocercus urophasianus) were banded in North Park
during this segment, 158 on the Lake John study area strutting grounds and.14
on wintering areas. Eight active strutting grounds were located within or
adjacent to the study area, one of which was a new ground. Of the 12 active
strutting grounds on the study area in 1963, only 4 were still active, with
one having only two attending males. The peak male count was 283. Two strutting
grounds accounted for 50.0 percent (141) of the cocks observed. Total cock
counts varied 28 percent during the peak week of female attendance. Over 32
percent (154.7 mi2) of sagebrush lands in North Park have been sprayed or
disked since 1957. Seven hundred and thirty hunters checked 746 sage grouse
through check stations on the opening weekend of a three day season. Age
composition of 698 wings collected WqS 50 percent immatures, 20 percent
yearlings, and 30 percent adults. Females comprised 73 percent of the subadult
and adult birds harvested. Peak of hatch was the week of June 19-25. Permits
were required of all sage grouse hunters with questionnaires sent to all
permittees. Of 1,184 permittees, 1,061 (89.7%) responded to the questionnaire.
Estimated harvest was 1,174 birds, a rate of 1.2 grouse per active hunter.
Winter distribution was plotted on the basis of 70 flocks, amounting to 2,327
birds. Strong segregation by sexes was evident in sage grouse flocks. Of the
60 flocks classified, 60 percent (36) were composed of at least 90 percent
birds of like sex. Mean size of male flocks was 21 birds, that of female flocks
48 birds. Flock size was less than 50 birds in 81 percent of all observations.
No differences could be detected in vegetal characteristics of winter sites used
by male and female flocks. Sixty-nine percent (36) of the winter use sites
had slopes less than 5 percent while only 4 percent of the areas had slopes
greater than 15 percent. Forty-eight percent (N = 44) of the winter flock
locations were on south, southwest, or west facing slopes. Sex ratio of 1,984
birds classified during the winter of 1974-75 was 61 females:39 males.

��-39-

EFFECTS OF SAGEBRUSH CONTROL
ON DISTRIBUTION AND ABUNDANCE OF SAGE GROUSE
Thomas D. I. Beck

An investigation of sage grouse abundance and distribution in North Park has
been in effect since 1963. Various topics have been concentrated on through
the years. Major emphasis was placed on winter distribution during this
segment. Continued emphasis was also placed on spring strutting ground counts,
brood counts, and harvest statistics with major revisions in collection of
harvest data.
P. S. OBJECTIVE
To determine the effects of 2,4-D spraying of sagebrush on (1) sage grouse
abundance, (2) sage grouse distribution, and (3) vegetative composition and
density.
SEGMENT OBJECTIVES
1.

To investigate seasonal numbers of sage grouse present by age and sex
throughout the year.

2.

To investigate the seasonal distribution of sage grouse throughout the
year.

3.

To investigate the current correlation between numbers and distribution
of sage grouse in relation to vegetative and habitat composition, patterns
and characteristics and initiate comparisons of current results with those
obtained in previous segments .

.4.

To prepare the required annual job progress report.
METHODS AND MATERIALS

Data was collected on sage grouse distribution during spring, summer, and
winter periods with major emphasis on winter. Age and sex composition of the
sage grouse population were investigated during spring, late summer, and
winter periods. Weather data was obtained from U. S. Weather Bureau climatological stations at Walden and Spicer. Extent of vegetative disturbance
was invest igated through field work and search 0 f records of Bureau of Land
Management and Soil Conservation Service. Methodology was identical to that
reported in the 1974 annual report (Beck 1974), except for changes in harvest
statistics methods. In 1974, four check stations were operated on opening
weekend. These stations were located at the State Line, Gould, Muddy Pass and
Willow Creek Pass. Each station was operated from about 1000 to 1800 MDT.
Data obtained per party were: County of origin, number of hunters, hours

�-40-

hunted, birds observed, birds bagged, birds lost, number of banded birds,
location of where banded birds were taken, and area hunted. One wing was
obtained from each bird checked. Few birds (48) were completely wingless.
Free, unlimited permits were required of all North Park sage grouse hunters.
Questionnaires were sent to all permittees within one week of the season end.
Data collected from questionnaires were: number of hunters, area hunted,
number of days hunted, birds bagged, birds lost, number of banded birds shot,
harvest location of banded birds, and county of origin.
RESULTS AND DISCUSSION
Capturing and Banding
One hundred seventy-two sage grouse were banded in this segment, 158 on
strutting grounds and 14 on wintering areas. Data on sex, age and specific
locations of the spring caught birds are presented in Table 1. Twenty-four
birds were recaptured, of which 14 were banded in 1973. When trapping
personnel could see bands on a bird no effort was made to capture the bird,
thus the number of recaptured birds was appreciably lessened. Only one bird,
an adult male, was killed during capture operations. Two adult females TTere
dispatched due to injuries received prior to being captured. The peak week
for capturing was April 18-25, when 79 birds were banded. This coincides
with the peak of male attendance on the strutting grounds as determined from
observation. Of the 24 males recaptured on strutting grounds and the one
mortality, only two (one adult, one sub-adult at initial capture) were recaptured on strutting grounds different from the ground of initial capture. Both
birds were caught five airline miles from the initial capture site.
All males captured were aged by techniques described by June (1967). These
techniques are based on primary feather wear. Some researchers, primarily
behaviorists, have also aged males by. the relative length and shape of the
rectrices (Wiley 1973). This method was also used on all captured males for
comparison. Only two of the 184 males examined could not be accurately aged
by the rectrices. In both cases adult males, as indicated by primary feather
characteristics, had rectrices characteristic of sub-adult males.
Strutting Ground Counts
Known strutting grounds in the area were censused regularly from March 13
until most strutting activity had ceased in late May. Strutting activity
began approximately three weeks earlier in 1974 than in 1973, presumably due
to warmer temperatures and earlier melting of snow. Eight active grounds
were located in or adjacent to the original study area. One new ground was
located, bringing to four the number of new strutting grounds located in the
last two springs. Of the 12 active strutting grounds marked by Gill (1965)
only four were still active. The peak male count for the eight strutting
grounds was 283, which included counts made from March 26 to May 14 (Table 2).
Coun ts were made twice.on all grounds and on the largest five grounds three
times during the peak week of male attendance as determined from counts and
capture data, thus providing some data on the stability of numbers of males

�-41-

on the grounds during this week. The maximum count for the eight grounds
during this week was 267, whereas a minimum count of 193 was recorded. Thus,
daily variation during the peak week could result in a 28 percent reduction
in the count (267-193)
267
.

Table 1.

Age and sex classes of birds banded in North Park, spring 1974.

Area

Adult

Male
Sub-adult

Adult

3

3

Alkali Lake Strutting Ground

8

3

2

Boettcher Junction Strutting
Ground

11

8

10

Walden Strutting Ground

4

6

Coalmont Strutting Ground

16

3

Cowdrey #5 Strutting Ground

2

Riley Strutting Ground

2

8

4

4

Hound Strutting Ground

12

3

Fish Hatchery Strutting Ground

18

12

Jackson County 5

1

Ridge Road Strutting Ground

2

10
20

1

7

1

1
1

5

3

2

20

1

3

34
1

3

Walden Reservoir
46

6

1
1

1

1

1

2

20

16

165

Butte Strutting Ground

83

37

2

Roth Strutting Ground

-----.

13

1

Monahan Draw Strutting Ground

Total

6

Wattenberg Strutting Ground

Total

Female
Sub-adult

�-42Table 2.

Summary of strutting ground counts, northwest North Park 1974.

Strutting Ground

Alkali Lake
Boettcher Lake Junction
Hound
Walden
Wattenberg
Cowdrey 115
Riley
Monahan Draw

Total

During 4/18-4/25
Maximum Males
Minimum Males

Maximum Males

Date

72

5-14

69

48

46

4-11

44

42

69

4-18

69

57

37

4-18

37

29

22

3-26

18

14

19

4-18

19

15

15

5-12

8

5

3

4-18

3

3

267

193

283

Strutting ground counts were initiated as a population index for management
purposes. A variation of as much as 30 percent in the counts during the
supposedly most stable week of male attendance, however, causes one to question
the value of the strutting ground counts. Few agencies have the maripower to
count all strutting grounds in an area more than once during the peak week,
if that often. Analysis of past strutting. ground counts in the study area
shows only two years since 1959 when the percent increase or decrease in
counts between subsequent springs was greater than 30 percent (discounting 1972
when all grounds were not counted). Thus, most of the supposed population
fluctuations pOSSibly, though not probably, can be attributed to daily variation
in attendance and the chance of an observer being present on the peak day.
Other factors that tend to cast doubt on the value of strutting ground counts
as an indicator of population levels are multiple changes in personnel and
little or no effort to locate new grounds since 1965 until 1973. Had not the
four new grounds been located, counts would have been even lower in 1973 and
1974, 177 and 143, respectively.
The Monahan Draw strutting ground, which is in the large block spray area, had
only 3 cocks present this spring. This ground had maintained high numbers of
birds in the years since the spraying (64 in 1966, 74 in 1967,53 in 1968,
80 in 1969, 57 in 1970, 81 in 1971), and the sudden drop in 1972 (10), and
1973 (11), is at present unexplainable.
The new ground located this spring is
approximately 5.5 miles southeast of the Monahan Draw ground and had a peak
of 69 cocks. In addition, one of the new grounds located in 1973 (15 cocks in
1974) is only 0.5 mile east of the Monahan Draw ground. Counts for the southern

�-43-

portion of North Park have likewise been appreciably affected by the discovery
of a new ground in 1970. In summary, the value of strutting ground counts
needs to be seriously examined in light of the high degree of variation from
numerous sources before using these data for management purposes. Serious
consideration should be given to examining evidence for possible distributional shifts in the last decade.
A minimum of 32 birds banded in 1973 were observed on strutting grounds in
1974. Of these, 27 showed fidelity to the ground of capture, whereas five
cocks were observed on grounds different from the ones where banding took place,
movements ranging from 2-9 miles (5, 5, 2, 9, 2). Combining observation and
recapture data, a minimum of 27 males were loyal to the strutting ground where
initially captured, while at least 7 males changed strutting grounds.

Harvest Data
Results of the 1974 check station operations are presented in Table 3, along
with data from all years since the inititation of this study. The age ratio
from 698 sage grouse wings collected was 50 immatures:20 yearlings:30 adults.
Sex ratios were determined for each age class, with a ratio of 51 females:49
males for immatures, 65 females:35 males for yearlings, and 79 females:21
males for adults. Data from 350 wings from immature birds indicated the peak
week of hatch was June 19-25 (34%), while 68 percent of the young hatched
between 12 June and 2 July. Results of the 1974 hunter questionnaire are
presented in Table 4.
Winter Distribution
Seventy sage grouse flocks were sighted, amounting to 2,327 birds. Weather
conditions and calendar dates could not explain patterns of observatLon nor
distribution. Distribution of sage grouse 'flocks was limited to that portion
of the Park north of Owl Ridge and Peterson Ridge. All areas south of this
ric./'line had sparse available sagebrush.
App rc xtmat ef.y 483 mi2 of sagebrush lands are suitable summer habitat, of which
approxf.m s t eLy 50 percent is available during winter. Over 154 mi2 of sagebrush has been sprayed or plowed since 1957. Some of this total has undoubtedly
reverted to sagebrush stands suitable for sage grouse use.
Characterization of Winter Flocks
Sage grouse in North Park are gregarious, being found in flocks most of the
time. Only 7 single birds (3 males, 4 females) were observed in the winter of
1974-75 out of a total of 2,327 birds observed. Strong segregation by sexes is
evident, with 60 percent (36) of the 60 flocks classified being composed of
at least 90 percent birds of like sex. Male flocks were smaller than female
flocks, averaging 21 birds, compared to 48 females per flock. Flock size was
less than 50 birds in 81 percent of all observations. Sex ratio of 1,984 birds
cLassi fLed during the winter of 1974-75 was 61 females: 39 males.

�Table 3.

Year

1963
1964
1965.Y

1966
1967
1968
1969
1970
1971
1972
1973
1974

North Park sage grouse hunter checks, 1963-1974.-1/

Bag
Limit

Hunters
Checked

Hours
Hunted

Adult
Cocks

Adult
Hens

Percent
Adults

Juvenile
Cocks

Juvenile
Hens

Percent
Juvenile

2
2
2
2
2
2
2
2
2
2
2
2

592
217
150
306
300
546
662
564
357
452
366
730

2,460
624
626
1,227
1,177
2,604
2,936
2,617
1,802
2,377
1,599
5,912

62
25
27
31
50
80
79
58
28
50
24
94

150
81
30
116
127
135
180
89
67
69
54
254

42
59
49
56
67
42
70
55
51
67
52
50

113
28
26
45
42
156
37
53
54
40
29
171

181
45
26
71
46
141
74
66
39
18
42
179

58
41
51
44
33
58
30
45
49
33
48
50

Unclassified
(Dressed)
Birds

- 3/

- 3/
- 3/
- 3/
- 3/

-1/

199

76
76
44
48

Total
Birds

Birds
per
Hunter

Hours
Hunted
per Bird

506
179
116
263
267
512
569
266
264
253
193
746

1.03
0.82
0.77
0.86
0.89
0.94
0.86
0.47
0.74
0.56
0.53
1.02

4.86
3.49
5.40
4.67
4.41
5.09
5.16
9.84
6.83
9.40
8.28
7.92

1/ Data from 1963 through 1973 are based on Walden and Cowdrey check stations operated during opening weekends.

(These
stations check harvested birds primarily in the northwest part of North Park).
Data from 1974 are based on four check
stations (State Line, Gould, Muddy Pass and Willow Creek) operated during the opening weekend.
(These stations sampled
all of North Park).

1/ A one day season, hence only a one day check.
1/ Composition of kill projected from wing samples, hence no unclassified birds.

1-

t

�-45-

Table 4.

North Park sage grouse hunter questionnaire data, 1974.
Projected
For

886
No. in sample
Percent total of
74.9
permittees
749
No. hunters
84.5
Percent hunters
137
No. non-hunters
15.5
Percent non-hunters
No. successful
398
hunters
Percent hunters
53.1
successful
1,217
No. hunter days
1.62
Days/hunter
No. sage grouse
859
bagged
.97
Grouse/permittee
1.1
Grouse/hunter
Grouse/successful
2.2
hunter
48
No. grouse los t
.06
Crippling loss/hunter
907
Total kill
Percent crippling
5.3
loss
Percent success of
44.9
permittees
28
Banded birds
Percent non12
deliverable

1,184

123

1,061

175

Projected For
1184

14.8
124
70.9
51
29.1

89.7
873
82.3
188
17.7

10.3
87
70.9
36
29.1

100
960
81.1
224
18.9

70

468

49

517

56.5
199
1.60

53.6
1,416
1.62

56.5
139
1.60

53.9
1,555
1.62

151

1,010

106

1,116

.86
1.2

.95
1.2

.86
1.2

.94
1.2

2.2

2.2
54
.06
1,064

2.2

2.2
58
.06
1,174

6

.05
157

4
.05
110

3.8

5.1

3.8

4.9

40.0

40.0

3

44.1
31

43.7
33

o

12

2

1. 01% (12)

Characterization of Winter Use Sites
Vegetal characteristics for 60 flock locations were not significantly different
for male and female flocks. Measurement of vegetal and physical characteristics
of the environment for all flock locations in the winter of 1974-75 appear to be
significantly different than similar measurements in 1973-74, possibly a reflection of total snowfall. Statistical tests have not been finished on these data.
Slope appears to be a significant factor in selection of a flock use site.
Sixty-nine percent (36) of the winter use sites described had slopes less than
five percent, while only four percent of the areas had slopes greater than 15
percen t .

�-46-

Aspect also appears to be a significant factor on site selection.
Forty-eight
percent (N = 44) of winter flock locations were on west, southwest, or south
slopes. The prevailing wind is from the southwest.
Further analysis of data collected during this and previous segments will be
presented in two subsequent reports; a Master's Thesis by Thomas D. I. Beck,
and the final comprehensive report, also to be prepared by T.D.I. Beck.
Completion dates for the two reports are 31 May 1975 and 14 August 1975,
respectively.

LITERATURE CITED
Beck, T.D.I. 1974. Effects of sagebrush control on distribution and abundance
of sage grouse. Colorado Fed. Aid Proj. W-37-R-27, Work Plan 3, Job 8a.
42 pp.
Gill, R. B. 1965. Effects of sagebrush control on distribution and movements
of sage grouse. Colo. Game, Fish and Parks Dept., Game Res. Rept., Part
3. W-37-R-17.
185 p.
June, J. W. 1967. Sage, blue and ruffed grouse sexing and aging characteristics.
Wyoming Game and Fish Comm. 59 p.
Wiley, R. H. 1973. Territoriality and non-random mating in sage grouse,
Centrocercus urophasianus.
Animal Behaviour Monographs.
Vol. 6, Part2.
P. 87-169.

Prepared bY~\~.~~~_~
;~
__
'~_l_~~~~__._~~_-__~__~__~_;_-~~~~_~
__
(
Thomas D. I. Beck
Graduate Research Assistant

_

�-47-

State 0 f

C_O_L_O_RAD
__ O

_

Project No.

W_-_3_7_-_R_-_2_8

_

Work Plan No.

3

------------------

Job Title:

Job No.

8a

_

Effects of Sagebrush Control on Distribution and Abundance of Sage
Grouse: Attributes of a Wintering Population of Sage Grouse,
North Park, Colorado

Period Covered:
Personnel:

Game Bird Survey

April 1, 1973 to March 31, 1975

Thomas Beck, Don Benson, Clait Braun, Lonnie Brown, Thomas
Campbell III, Jack Corey, Courtney Crawford, John Ellenberger,
Heather Flanagan, Howard Funk, Fred Giese, Don Gore, Jack
Gustafson, John Hobbs, Richard Hoffman, Jim Jackson, Russell
Kozacek, Bruce McCloskey, Mike McLain, Ken Miller, Ron Oakleaf,
Sig Palm, Steve Porter, Kenneth Russell, Wayne Russell, Howard
Spear, Robert Streeter, Michael Szymczak, John Wagner, Richard
Wenger.

ABSTRACT
Wintering habits of sage grouse (Centrocercus urophasianus) were studied in
North Park, Colorado during 1973-74 and 1974-75. Distribution was plotted
from sightings of 199 flocks and 17 single birds, totaling 5,080 birds.
Distribution in winter was restricted to the portion of the park north of
Owl and Peterson Ridges due to lack of available sagebrush (Artemisia spp.)
south of the ridges; a result of greater snowfall in the south. Further
restrictions of range occurred in the north area due to snow cover, steepness
of a slope, and unknown factors. Thus, only 50 percent of the 1,251 sq. km.
of sagebrush-dominated lands in North Park sustained winter use by sage grouse.
Seven high-use areas comprising approximately 85 sq. km. were delineated.
These areas accounted for 74 and 37 percent of all flocks and 79 and 51 pe r-:
cent of all birds observed in the winters of 1973-74 and 1974-75, respectively.
Vegetal characteristics of the high use areas were not significantly different (P&gt;. 05) from characteristics of other areas of grouse use. Over 32
percent (401 sq. km.) of sagebrush lands in North Park have been disturbed by
2,4-D spraying, plowing and seeding, and burning since 1957, and only four
of 199 flocks were observed in areas that still showed evidence of alteration.
Flocks were the dominant social unit as only 17 single grouse were observed
in a total of 5,080. Strong segregation by sexes was evident. Of 176 flocks
classified as to sex, 70 percent (123) .were composed of at least 90 percent
birds of like sex, of which 81 percent (100) were unisexual. Segregation was
more pronounced in male than in female flocks. Flock size was 50 or less in
88 percent of all observations. Mean size of flocks in 1973-74 was 12.2 for
males and 29.2 for females, whereas mean flock sizes in 1974-75 were 21.2
for males and 48.1 for females. The increase the second winter was possibly
the result of excellent production in the summer of 1974. Females had a
stronger tendency to form large flocks (:&gt; 75) than did males. In 1973-74

��-49TABLE OF CONTENTS

Introduction .
Description of Study Area
Methods
Results
Delineation of Winter Distribution
Flock Composition and Size . . • •
Physical Characteristics of Winter Use Sites
Vegetal Characteristics of Winter Use Sites
Movements . .
Discussion
Literature Cited

LIST OF TABLES
Table

Page

1

Winter precipitation and temperature, North Park,
Colorado . • . • . • . • . . • . . .

2

Location and size of areas of high use by wintering sage
grouse, North Park, 1973-74 and 1974-75 ..•..

3

Number, frequency of encounter, and percentage of total
sage grouse observed as singles and flocks, North Park,
1973-74 and 1974-75 . . .
. . . . . . .

4

Sex composition of winter flocks of sage grouse,
North Park, 1973-74 and 1974-75 ..
. ....

5

Temporal trends in flock size 'of sage grouse in
North Park, 1974 . . . . . . . . . . . . • . .

6

Temporal trends in flock size of sage grouse in
North Park, 1975
. . • . .

7

Vegetal characteristics of winter use sites, 1974 and 1975

8

Vege te.lcharacteristics of high grouse use areas, North
Park, 1974 and 1975
.

9

Minimum distances traveled from spring to winter by male
and female sage grouse banded near Lake John North Park,
1973-74 and 1974-75 .........•.........

LIST OF FIGURES

1

North Park study area . . .

2

Cumulative precipitation at Walden, Colorado during
December, January and February 1973-74 and 1974-75 .....

�-50LIST OF FIGURES (Continued)
Figure
3

Location of sage grouse flocks during winters of
1973-74 and 1974-75, North Park, Colorado ••

4

Aspect and percent slope of areas utilized by flocks
of male sage grouse in winter • • • • • • "
• • • •

5

Aspect and percent slope of areas utilized by flocks
of female sage grouse in winter.
• •.•

6

Radiation indexes for various facets at Latitude 400 N
ACKNOWLEDGMENTS

I wish to thank Dr. Ronald A. Ryder, Professor, Department of Fishery and Wildlife Biology, Colorado State University, for serving on my committee, for providing advice when I most needed it, and for editing this thesis. Thanks are
expressed to Dr. Philip N. Lehner, Assistant Professor, Department of Zoology,
for serving on my committee and editing this thesis. Howard Funk, Project
Leader, Colorado Division of Wildlife, provided valuable administrative assistance, served on my committee, and edited this thesis.
I am grateful to Stephen Porter and John Wagner, wildlife conservation officers,
along with Fred Giese, U.S. Fish and Wildlife Service, for assistance in field
work and friendships that made the long North Park winters easier. Special
thanks go to their respective wives, Betsy, Kathy, and Marcia, for their patient
understanding of my taking their husbands out at all hours of the day and night
and for their warm hospitality.
I am grateful to Russ Kozacek, Dick Wenger, Heather Flanagan, and Tem Campbell
for their diligent assistance in field work. My appreciation is extended to
the many Division of Wildlife personnel who contributed to this project.
Finally, to Dr. Clait E. Braun, Wildlife Researcher, Colorado Division of
Wildlife, a very special appreciation for his many helpful suggestions, his
sincere interest in the project, and most importantly, his friendship. His
concern for me as a scientist and an individual will always be remembered and
appreciated. Though not on my committee, Dr. Braun was instrumental in development of the study, supervision, and editing of this thesis .

�-51-

ATTRIBUTES OF A WINTERING POPULATION
OF SAGE GROUSE, NORTH PARK, COLORAOO
Thomas D. 1. Beck

INTRODUcrION
Sage grouse, largest North American tetraonid, inhabit areas dominated by
sagebrush (Artemisia spp.) and subdominant grass types with interspersed
native and cultivated hay meadows. Dependence on sagebrush for food and cover
has been well documented in all seasons (Griner 1939, Patterson 1952, Dalke
et aL 1963, Gill 1965, Autenrieth 1969, Klebenow 1969, Eng and Schladweiler
1972). As a result of agricultural demands, livestock grazing, and encroachment of man-related developments, there has been a continued destruction of
sage grouse habitat. In the face of changing land use patterns, sage grouse
have continued to be highly specialized in their habitat requirements. They
have not adapted to utilize land not dominated by sagebrush. Climax vegetation types, such as sagebrush, are hazardous food sources due to their
slow recovery after natural or man-caused catastrophies (Daubenmire 1970).
Due to pressures to alter vegetative composition of sagebrush dominated rangelands, the Colorado Division of Game, Fish and Parks (now Colorado Division
of Wildlife), in cooperation with the Bureau of Land Management, initiated a
study in North Park, Colorado, to document the effects of 2,4-D spraying of
sagebrush on sage grouse abundance and distribution. At the time of the
study's initiation in 1963, 2,4-D spraying to reduce or eliminate sagebrush
was a popular tool of land management agencies.
Pre-treatment studies were accomplished in 1963-65 with study patterns for
spraying designated, and actual spraying accomplished in June 1965 (Gill 1965).
Two years of study on the immediate effects of spraying sagebrush were completed in 1967 (Carr 1967). During the summers of 1968 and 1969, studies
were conducted on distribution of females in relation to sprayed areas by use
of radio telemetry (Poley 1969, May 1970).
Even [hough changes in sage grouse abundance and distribution due to habitat
changes are likely to occur during a time of limited habitat, i.e., winter,
little scientific effort has been directed toward winter ecology of sage
grouse. Numerous references have referred to winter distribution of sage
grouse but few quantitative data have been presented. In areas of greatly
varying elevations, winter distribution and movements of sage grouse are
dependent upon snow depth, with birds moving to lower elevations as snow
covers the sagebrush on higher ranges (Griner 1939, Patterson 1952, Dalke et
al. 1963). Eng and Schladweiler (1972) studied five radio telemetered
females and found little movement during the winter. Thei~ central Montana
study area does not have elevational variation nor snow depths as great as
the areas studied by Griner, Patterson, and Dalke et al. The birds primarily
used areas of greater than 20 percent sagebrush canopy coverage. Eng and

�-52-

Schladweiler reported wintering sage grouse were "never" observed on steep
slopes of dense sagebrush, with all feeding and roosting areas on level or
gently sloping ground. The possibility of limited quantity of food in winter
as a population regulation mechanism for sage grouse has not been reported
in the published literature.
Specific studies on sage grouse have not been directed at winter flock composition. However, Patterson (1952) referred to partial segregation of sexes
in winter flocks in Wyoming".
. • flocks composed predominately of males or
females were commonly seen • • ." Quantitative data indicating the degree
of segregation were not presented. Patterson further stated, "there was a
pronounced tendency for adult birds to segregate themselves according to sex
on their winter grounds." Patterson observed that the prevalent social unit
was the flock and that flock size varied between 10-100 birds, with occasional
flocks of up to 200 birds. Autenrieth (1969) and Bailey (1925) also commented on segregation by sex in winter flocks in Idaho and Colorado, respectively,
but presented no data. Investigations on other grouse species indicate
strong segregation by sex in winter is associated with differential habitat
utilization by sexes (Koskimies 1957, Seiskari 1962, Weeden 1964, Braun and
Schmidt 1971).
In an effort to increase knowledge of sage grouse winter ecology, this study
was initiated in 1973. The following hypotheses were developed: (1) distribution of sage grouse in winter is directly related to available sagebrush,
(2) strong segregation by sex occurs, and (3) male and female sage grouse
utilize areas of differing vegetative and physical characteristics during
winter.
In addition, winter flock size and movements of banded birds were
investigated.

DESCRIPTION OF STUDY AREA
North Park (Jackson County) was chosen for study due to a plan proposed in
1962 by the Bureau of Land Management (BLM) that would result in over 1,192
sq. km. of BLM controlled sagebrush lands in North Park being sprayed. This
area supported the highest population of sage grouse in Colorado during the
early 1960's (Rogers 1964:47).
North Park is northernmost of Colorado's four large inter-montane parks, and
is located immediately west of the Front Range of the Rocky Mountains.
It
lies within townships 5 to 11 north, ranges 77 to 82 west, sixth principal
meridian. The park is completely enclosed by mountains. The west boundary
is formed by the Park Range which runs north-south parallel to the Medicine
Bow Range which forms the east boundary 80 km. distant. The south end of the
park is enclosed by the Rabbit Ears Range. Independence Mountain, approximately 90 km. to the north, forms the north boundary.
The topography of Nor-th Park is relatively flat in contrast to the surrounding
mountains.
In general, it consists of low, flat to undulating benches and
ridges separated by the main drainages. The main streams form a meandering
pattern on low alluvial flood plains up to one-half mile wide. Elevation of

�-53most of the sagebrush lands in North Park is between 2,400 and 2,585 m. above
sea level. The principal exception is Owl Ridge-Peterson Ridge, two en:-toend ridges running northwest from Owl Mountain on the east to Delaney Butte
on the west and rising 61.5-154 m. above the surrounding terrain (Fig. 1).
Sixty-four percent of the park lies north of this ridge line. The mountains
encircling the park rise steeply to elevations of 3,077 to 3,846 m. above sea
level. Geology of the park has been described in numerous publications (Beekly
1915, Finch 1957, Hail 1965). Major drainage of the park is to the north via
the North Platte River. Main tributaries to the North Platte include the
Roaring Fork, North Fork, Canadian, Michigan, and Illinois Rivers and Grizzly
Creek.
The park has over 1,870 sq. km. of sage grouse habitat as all areas not timbered
are considered potentially usable by sage grouse. Of this, about 1,252 sq.
km. are dominated by sagebrush, while about 620 sq. km. are irrigated meadows
of native sedges and grasses.
With the exception of hay meadows along watercourses the vegetation of North
Park is mainly composed of the shrub-bunchgrass type dominated by species of
sagebrush. Five species of sagebrush dominate, with Artemisia tridentata
vaseyana occupying approximately 90 percent of the sagebrush type in North
Park. Artemisia longiloba, ~. cana viscidula, ~. argilosa, and ~. ~
are
dominant on selected smaller areas with certain soil conditions favorable to
the species (Beetle 1960, Smith 1966). Other important shrubs are greasewood
(Sarcobatus vermiculatis), rabbitbrush (Chrysothamnus spp.), willow (Salix spp.),
and bitterbrush (Purshia tridentata). Herbaceous vegetation consists primarily
of low growing perennial forbs and perennial bunch grasses with few annual forbs.
Climatic conditions in North Park during winter (December-February) vary considerably north and south of Owl and Peterson Ridges. Winter precipitation in the
north is normally 45.2 percent of that in the south, based on 30-year averages
(U. S. Weather Bureau files, Walden) (Table 1). Precipitation in winter normally accounts for 15.3 and 22.7 percent of the yearly precipitation in the
north and south areas, respectively. Snowfall in the south usually is sufficient
to cover most of the sagebrush not on steep slopes. Precipitation in the north
(Walden) is relatively uniform throughout the winter, whereas precipitation in
the south (Spicer) is greatest in December and decreases steadily through February. Harch typically has frequent snowstorms followed by warm days. Much of
the ncrth area is usually snow free by March 7 except immediately following
snowstorms. Ha rch was not included in the analysis of winter moisture as precipitation during the month does not normally contribute to snow accumulation due
to rapid melting.
Winter precipitation in the north was 26.2 percent above the 30-year norm in
1973-74, while it was 25.9 percent below the norm in 1974-75. The major difference in precipitation patterns in the north between the two winters was during
late December and early January (Fig. 2). Winter precipitation in the south
was within 5 percent of the norm in both 1973-74 and 1974-75. Changes in
Hinter precipitation in the south between years did not affect the amount of
sagebrush available during the winter months as snowfall both years was sufficient to cover nearly all sagebrush on moderate slopes (10-30 percent).
Winter temperatures vary little between the north and south areas. Temperatures
during the two winters were similar but the winter of 1974-75 was slightly warrae r , having had four more days with maximum dail temperatures greater than -loCo

�-54-

Figure 1. North Park study area.
1:250,000.

U. S. Geological Survey map, scale

�Table

1.

Winter

precipitation

and temperature,

North Park,

Colorado.

--Station

Mean Daily Temperature

Precipitation

(cm)

December

January

February

Total

December

January

February

year

(0C)

Days above
_1°C

--

vValden,

1973-74

1. 75

2.51

0.46

4.72

- 7.2

-10. 0

-8. 9

40

,V aid en,

1974-75

0.36

1. 14

1. 27

2.77

-10.2

- 8.6

-8. 6

44

Walden, 30-year
average

1. 37

1. 30

1. 07

3.74

- 7.4

- 9.2

-7. 7

I

\J1
\J1

I

Spic er, 30-year
average

3.20

2.69

2.39

8.28

- 7.4

- 8. 6

-6.6

�-56-

1974-75
I--~---,
~

,- --~

___

___

,.-

,

,- -,
_.I

I

.J

.J"

12-112-1012-2012-30 1-9 1-19 1-29 2-8

2-18 2-28

DJU"E
Figure

2.

Curnu Ia t iv e precipitation

during December,
1974-75.

January

at Walden,

and February,

Colorado

1973 -74 and

�-57-

METHODS
Winter observations were conducted from 8 January to 8 March 1974, and 24
December to 1 March 1975. Distribution of wintering sage grouse in North Park
was investigated by searching all sagebrush lands and all probable use areas
were searched equally. An area was considered potentially usable if any sagebrush was visible above the snow. Limited searches were also made in areas
not considered usable. Areas inaccessible by four-wheel drive vehicle were
searched from snowmobile or snowshoes. Aerial search (with two observers)
along random transects (covering 10 percent of the total area in the park)
were conducted on 21-22 February and 18 March 1974. Data on grouse flocks
observed from the air were used in mapping distribution but were disregarded
when analyzing flock sizes because of inaccuracies involved in counting flying
grouse. Once grouse were located from the ground they were first observed
through 7 X 50 binoculars to obtain a total count and to determine sex composition. Attempts to observe colored leg bands were made through 15-60
variable power scopes. Written records were kept on both grouse observations
and grouse sign (tracks, droppings, roosts). Observations reported by reliable
observers were followed by intensive searches of reported locations within one
day of the initial sighting. All locations were plotted on U. S. Geological
Survey topographic maps (7.5 minute series).
A flock was considered to be any group (two or more) of birds in close proximity to each other. The decision as to flock integrity was made by the observer as flocks often ranged over distances from 10-200 m., thus inhibiting use
of standard distances between birds as criterion for flock integrity. Flocks
comprised of greater than 50 percent of one sex were referred to as female or
male flocks depending upon which sex predominated.
Sage grouse are sexually dimorphic, thus allowing sex of birds to be visually
determined up to 200 m. Male sage grouse in breeding plumage are characterized by contrasting color zones of black and white from the base of the lower
mandible to the abdomen. The head is dark, in appearance, with a mixture of
gray and black feathers. The black and white speckled area at the base of the
lower mandible is separated from the black throat by a narrow white collar.
This collar runs from the eye, under the head, and is visible from the front
and side. Lines of demarcation are distinct on the collar. The black
feathers of the throat are white tipped, but contrast sharply with the solid
white breast feathers. The feathers on the lower breast are white with black
tips. The black portion of these feathers become greater towards the abdomen
which appears solid black. Males have long (10-15 mm.) filoplumes extending
from the side 0 f the neck which are easily visible, especially on windy days.
Males are larger in size, averaging twice as large and heavy as females.
Adult sage grouse undergo a complete post-nuptial molt which is completed by
late September. Juvenile birds acquire their first adult or winter plumage
hy late October. Thus, all males were in breeding plumage during the study
period (Bent 1932, Patterson 1952).
Female sage grouse are smaller and more drab colored than males. The abdomen
is black, but not as solid as in males, being splotched with gray and white.
The breast is white, gray and tan, giving a splotched appearance \vith no
discrete color patterns visible. Females do not have the white neck collar

�-58-

nor filoplumes as do males. The basal area of the lower mandible is more
white in females than in males. The back and sides of females are lighter
in color, and are less black than males.
The following physical and vegetal characteristics of each site where birds
were observed were recorded: aspect, slope, snow depth and condition, sagebrush
density, crown breadth, and height of sagebrush above snow. Slope and aspect
were measured with an Abney level and compass, respectively. Snow condition
was rated as crusted, powder, or melting. Sagebrush density was measured as
number of plants per circular 0.004 ha. plot. Crown breadth was calculated
as the mean of 10 representative shrubs arbitrarily chosen by the observer
from within the plot. Height of sagebrush above snow was calculated as the
mean of three shrubs within the plot which the observer chose as being representative of shrub height within the utilized area.
The following weather conditions were recorded at each site: wind direction,
temperature, cloud cover, and precipitation.
Movement data were obtained from observations of birds banded with colored
bandettes coded to banding sites. The location of each sighting was plotted
on a 7.5 minute U.S.G.S. topographic map and the airline distance from banding
site measured; thus, all movements represent minimum distances traveled.
Directions of movements were determined from banding sites near Lake John to
wintering locations.
Capturing and banding were conducted during the spring seasons of 1974 and
1975 in conjunction with long term study procedures (Gill 1965). Birds were
located by spotlighting and were then netted as described by Gill (1965).
All captured birds were banded with aluminum, serially numbered bands and
colored plastic bandettes. Sex and age classification was based on plumage
characteristics described by June (1967). Age, sex, weight, dat~, and location
of capture were recorded for each bird. The maximum number of banded birds
present in the winter of 1973-74 was 268 (166 males, 102 females), the result
of banding in spring, 1973. The corresponding figure for the winter of 197475 was 289 (186 males, 103 females) from additional spring banding in 1974.

RESULTS
Delineation of Winter Distribution
During 8 January to 8 March 1974, 129 sage grouse flocks and 10 single birds
were sighted, amounting to 2,753 birds. Between 1 January and 1 March 1975,
70 flocks and 7 single birds were sighted, involving 2,327 birds. Plotting
of flock locations (Fig. 3) indicates three major areas where no flocks were
observed, seven areas of high use, and scattered observations within remaining areas of sagebrush.
The largest area of no observed use was the area south of Owl and Peterson
Ridges, which amounts to 36.0 percent (446 sq. km.) of the sagebrush lands in
North Park. Only one of 216 sage grouse sightings was south of this ridge
line. Very little sagebrush was available in this area due to extensive snow

�-59-

Figure 3. Location of sage grouse flocks during winters of 1973-74 and 1974-75,
North Park, Colorado. U. S. Geological Survey map, scale 1:250,000. Each
dot represents one flock. Numbered areas correspond to high use areas
described in Table 2.

�-60-

cover. The area between the Canadian River and Medicine Bow Range had ample
available sagebrush throughout winter but no observed grouse use. This area
accounts for 9.1 percent (114 sq. km.) of the sagebrush lands in North Park.
The third area of no observed grouse use was the broken ridges south of
Independence Mountain and north of Jackson County Road No.6, encompassing
5.0 percent (63 sq. km.) of the sagebrush lands. Extensive stands of tall,
dense sagebrush on relatively steep slopes were common in this area. Thus,
50.1 percent of the sagebrush dominated lands in North Park received no
observed use by sage grouse during winter months in both years of study.
Seven areas of high use were evident and comprised 6.8 percent (85 sq. km.)
of sagebrush lands in North Park. These seven areas were not searched more
frequently than other less productive areas, thus eliminating the bias of
implied heavy use due to high frequency of search time. Searches for sage
grouse were conducted during 60 days each winter. At least one or more of
the seven areas was searched on 30 days each year. Areas subsequently determined to be used lightly or not at all were searched on at least 20 days
during each season.
Nearly 75 percent (74.4) of all flocks and 79.2 percent of all grouse
observed in 1974 were within these seven areas, whereas only 37.1 percent of
all flocks and 51.2 percent of all grouse observed in 1975 were in these
areas (Table 2). Sage grouse flocks were mobile as birds could not always
be found in high use areas. Flock size varied considerably between days in a
given area. Since grouse were not individually marked it was not possible
to determine if a particular flock was habitually in an area. As further
evidence of mobility from these and other areas, on 14 occasions flocks, upon
flushing, flew from sight beyond ridges 13 or more kilometers distant.
Thirty-two percent (401 sq. km.) of the sagebrush lands in North Park have
been disturbed by 2,4-n spraying, plowing and seeding, and burning since 1957
(Beck, unpub. data). Only four flocks were, observed in areas that were still
noticeably altered. Three flocks were observed on 1 January 1975 in a crested
wheatgrass (Agropyron desertorum) field which was planted in 1962. Snow depth
was less than 2 em. and exposed young sagebrush plants were common. One flock
was observed in a crested wheat grass seeding on 9 January 1974. Snow depth
was 15 cm. and the closest available sagebrush was 40 m. distant.
Flock Composition and Size
Sage grouse in North Park were gregarious, the flock being the prevalent
social unit (Table 3). Over 85 percent (85.4) (2,350) of the 2,753 birds
observed in 1974 were classified as to sex. Males numbered 889 and females
1,461 for a pooled sex ratio of 62 females:38 males. Approximately 85 percent
(84.3) (1,984) of the 2,327 sage grouse observed in 1975 were classified as
to sex with males numbering 773 and females 1,211. The pooled sex ratio for
1975 was 61 females: 39 males.

�~-~~~.
~~~~~~~==~~~~~
T'abla 2.

Location and size of areas
1973 -74 and 1974-75.

of high use by wintering

sage grouse,

Approximate
Area

Location

NE+ S13,

Et S12,

RSOW, T10N; NEi S19, SlS,
TI0N

sw ; 817, NWi, si S7, R79W,

Park,

No. of Flocks
. Obs erved

Total Birds
Obs erved

1974

1975

1974

7. S

14

°

114

°

16.9

16

1

214

60

km.)

(Sq.

1.

North

2. SEi S 19, si S20, SWi S27, si,

NWi S2S, S29, E~
S32, S33, S34, R79W, TI0N; Nt S3, Nt S4, NEi

S5, R79W,
3.

T9N

SEisl,
Eisl2,
NEis13,
RSIW,
1
S7, NW4"S20, S21, RSOW, T9N

4. SEis7,

siss,
S20, S21, S2S,

5.

6.

. 7.

T9N;

wi,

1975

•....•

NEi

I

'

Ni, SEis17, WiSI6, SEis19 .•
NEi S29, Ni S33, RSOW, TI0N

Eis7, SWiss,
SWiS16, S17, Eisl8,
Eis19,
S20, S21, sw] S22, wt S27, S2S, S29,
S30,
NEi S33, R7SW, T9N; S 13, wi S24, Ei S23, R79W,
T9N

7.S

19

1

223

11

14.3

S

10

179

44S

26.6

19

8

S66

521

9. 1

13

6

371

152

2.6

7

o

213

o

85. 1

96

Z6

Z, 180

I, 192

Nt

si

S~ S31,
S32, R7SW, TSN;
N} S7, Nwi S8, R7SW, T7N

S2, R81W,
Total

TION

wi,

I
0\

NEi S5, S6,

�-62-

Table 3.

Number, frequency of encounter, and percentage of total sage grouse
observed as singles and flocks, North Park, 1973-74 and 1974-75.

Number of encounters
Frequency of encounter

a

Percent of all birds
observed
a

1

2-25

Size of Unit
26-50
51-100

17

153

21

16

9

7.9

70.8

9. 7

7.4

4.2

0.3

32.5

14.5

22.0

30.7

&gt;100

In percent of total.

Strong segregation by sexes was evident in sage grouse flocks. Of 176 flocks
classified in 1974 and 1975, 69.9 percent (123) were composed of at least 90
percent birds of like sex, of which 81.3 percent (100) were unisexual (Table
4). Segregation was more pronounced in male than in female flocks as 71.6
percent of male and 39.5 percent of female flocks were unisexual. Only one
flock (six birds) was equally composed of males and females.
Flock size was 50 or less in 88.4 percent of all observations (Table 3). Mean
size of flocks in 1974 was 12.2 for males and 29.2 for females, whereas, mean
sizes in 1975 were 21.2 for males and 48.1 for females. Differences between
male and female flock size during the two winters were significant (P &lt; 0.05).
Increase in size between years (1974 vs. 1975) was significant (P~ 0.05) for
male but not for female flocks. Median size for male flocks was 10 in 1974
and 15.5 in 1975, while corresponding values for female flocks were 15 and
20.5. No obvious patterns in flock size variation and frequency of sightings
were detected through the 60 days of observation each winter (Tables 5 and 6).
A large increase in total numbe r of female and female flocks observed occurred
during the last fortnight of each study period.

Physical Characteristics

of Winter Use Sites

Flocks of sage grouse were typically located on south, southwest, or west
faCing slopes of less than 5 percent (Figs. 4 and 5). The average percent
slope for areas used by female flocks was 4.9 (N = 74) and for male flocks was
6.0 (N = 82). Sixty-six percent of the areas had slopes less than 5 percent,
while only 12.8 percent had slopes greater than 10 percent. On areas with
slope greater than 5 percent, 66.7 percent of female and 93.1 percent of male
flocks were observed on west, southwest, south or southeast facing slopes.
Wind was present during 73.7 percent of 152 observations for which physical
data were collected. Wind direction was from the southwest 60.7 percent of
the time, west 17.0 percent, and south 12.5 percent. No data on wind velocity
were collected.

�-63-

Table

4.

Sex composition
of winter flocks
Park,
1973 -74 and 1974 - 75 a.

Siz e of Flock

51 -59

60-69

of sage grouse,

Percent
in Flock
70 -79
80-89
90-99

North

100

Total

4
2

36
24
5

42
34
11
1

2

2

6

Males
2 -9
10-19
20 -29
40-49
50-74
75 -99
Sub -total

I

2
1

2
1
2

2
3
2

I

1
3

1
4

5

7

I

I

8

68

95

16
5
1
1
3
2
1
2
15

II

1
32

24
23
8
5
5
4
3
9
81

176

Females
2 -9
10-19
20-29
30-39
40-49
50-74
75 -99
&gt;100
Sub -total

1
1

10

6

4
17

Total

4

14

11

24

23

100

Percent of
all flocks

2

8

6

14

13

57

aExcluding

5
1
2
1

1

2

I

5

1

3
1

I

2
2
1

one flock of 50:50 males

and females.

2
1
1

�-66-

N

s

• 2-9
0

10-19

0

20-29
2:30

A

Figure
5. Aspect and percent
slope of areas utilized
by
flocks of female sage grouse in winter.
Each concentric
circle
represents
5 percent
slope with the center
representing zero slope.
Symbols representing
different
flock sizes are given in the legend.

�-67-

Vegetal Characteristics of Winter Use Sites
Vegetal measurements were taken at 100 sites in 1974 and 51 sites in 1975.
Mean sagebrush density, mean height of sagebrush above snow, mean crown
breadth, and mean cover (ht. x density) were used in comparing sites utilized
by male and female flocks. Significant differences (P ~ 0.05) were documented
in sagebrush density and cover in 1974, when females used denser sagebrush
stands than males. Significant differences (P ~ 0.05) between years occurred
for sagebrush density on sites used by male flocks, height of sagebrush above
snow and cover for sites used by both male and female flocks (Table 7). Comparisons of vegetal characteristics for flocks of similar size but different
sex resulted in no differences other than those mentioned above. Vegetal
characteristics of each high-use area were compared to those of the combined
values for all locations not within a high use area. Only one high use area
had vegetative characters significantly different (P &lt; 0.05) from other high
use areas and the scattered use locations (Table 8). -This area was located
in the McCallum oil field where earlier researchers also observed wintering
flocks of sage grouse (Gill 1965, Carr 1967). Sagebrush in this area was
less dense than in other areas of flock sightings.
Twenty-six of the 199 flock locations were used as roosting sites, the remainder being feeding-loafing sites. Vegetative differences between the two
groupings were not significant (P&gt;0.05).
Roosting depressions in the snow
occurred at 13 of the 26 sites. Snow at 8 of the sites was crusted and at 5
sites was powder. These values were similar to the frequency of crusted and
powder conditions recorded at all sites (64.9 percent crusted, 29.8 percent
powder, 5.3 percent melting). Depressions to depths of 30 cm. were observed.

Movements
Seventy-one banded birds were observed in the winter of 1973-74, ·of which only
5 were females. Twenty-five banded birds were observed in the winter of 197475 and again, 5 were females. More emphasis was placed on obtaining observations of banded birds during 1973-74 than in 1974-75. Analysis of observations of banded birds indicated that sage grouse were moving freely throughout North Park. Thus, all sage grouse in the park should be considered as
one population rather than several localized, sedentary subpopulations.
Delineation of population boundaries was the primary objective of band sightings
and as the delineation was made after 1974 the time spent on obtaining band
observations was greatly reduced in 1975.
Average distance traveled from banding sites for males was 10.8 km. (N = 51)
in 1973-74 and 5.3 km. (N = 17) in 1974-75. Corresponding values for females
were 19.2 km. (N = 5) and 15.8 km. (N = 5). Not all banded birds were banded
with colored bandettes coded to capture locations, thus explaining the difference in banded birds observed and banded birds used for movement analysis
(66 vs. 51 and 20 vs. 17). Distribution of distances traveled is presented
in Table 9. Shorter distances traveled in 1974-75 probably resulted from the
milder winter. Direction of movement (N = 73) was northerly 17.8 percent of
the time, northeasterly 26.0 percent, easterly 19.2 percent, and southeasterly
21.9 percent. This corresponds strongly with the directions of the main
winter use areas from banding sites near Lake John. Maximum distance traveled
was 35.1 km. by an adult male in 1973-74.

�Table 7.

Year

Vegetal

Sample
Size
a

characteristics

of winter

Mean Sagebrush
Density
(Plants / O. 004 ha. )
M
F

use sites,

1974 and 1975.

Mean Height of
Sagebrush Above
Snow (cm)
M
F

Mean of Average
Crown Breadth
of Sagebrush
(cm)
M
F

Mean Cover
(Density x Height)
(cm-Plant/O.
004 ha , )
M
F

M

F

1974

58

42

46.0

6'8.2b

19. 9

23.9

37.6

43.9

991

I, 700b

1975

22

29

63.7c

77.5

29.6c

3S.6c

40.8

44.4

2, 048c

2, 906 c

aM denotes

flock of &gt;50 percent

males,

F denotes

bSignificant

difference

between

s~xes

during a year,

CSignificant

difference

between

years

within a sex,

flock of &gt;50 percent
P &lt; 0.05.
P~

0.05.

females.

I

0\

oe
I

�Table

8.

Area

Vegetal

characteristics

of high grouse

North Park,

Sagebrush
Density
(Plants /0.004 ha , )

No. of Birds

No. of Flocks

use areas,

1974

1975

1975

1974 and 1975.

Sagebrush
(em)

Ht.
1975

1974

Cover (Ht. x
Density) (cmPlant/O. 004 ha. )
1975
1974

1974

1975

1974

1

14

0

118

0

47.4

2

14

1

196

60

30.3a

62.0

17. 5

10.0

494a

620

3

17

1

207

11

72. 1

78.0

22.5

13.0

1, 675

1, 014

4

8

10

181

448

52. 0

63.5

26.3

35.7

1, 484

2, 328

5

18

8

857

521

54.9

87. 1

29.6

41.7

1, 846

3, 752

6

13

5

371

152

77.9

38.0

16. 2

21.3

1, 564

1, 165

7

7

0

213

0

66.5

38

45

610

1, 129

52.4

Remaining
locations

aSignificantly

different

from

other

high use areas

803

17.4

804

13.5

72.8

23.2

and/or

remaining

33.0

locations,

1, 309
P -&lt; 0.05.

Table 9. Minimum distances traveled from spring to winter by male and female
and female sage grouse banded near Lake John, North Park, 1973-74
and 1974-75.
: Distance Traveled (km.)
Year

0-5

5.1-10

10.1-15

15.1-20

20.1-30

&gt; 30

1973-74

13

10

20

9

4

1

1974-75

10

6

3

1

2

o

~

-------_._,-----------_.--.-----

__ .•

..

---

I

0\
\0

I

2,545

�-70-

DISCUSSION
Distribution of sage grouse in North Park during winter was primarily a
reflection of availability of sagebrush above snow, slope, and aspect. In
both years of study, snowfall was sufficient to cover nearly all sagebrush
on level and moderate slopes in the south portion of the park. Tall sagebrush
in swales and draws was covered with snow due to drifting. Although sage
grouse are capable of scratching into crusted snow to depths of 30 em., areas
without protruding sagebrush offered little cover and no indication that sagebrush was present beneath the snow. The south portion of the park provided
little, if any, habitat for sage grouse during winter due to snow accumulation.
This was true even in a mild winter (1974-75).
Distribution of sage grouse in the north area was also greatly affected by
snow accumulation. During the mild winter of 1974-75 less sagebrush was
completely covered. Consequently, a larger area was available for foraging.
More available sagebrush accounts for the reduction in use of the high use
areas delineated in 1973-74 and the increased difficulty in locating flocks.
As the winter of 1973-74 was more severe than normal, some high use areas
possibly were the last suitable habitat. Only 3 of the 7 1973-74 high use
areas sustained high use in 1974-75, thus discounting the possibility of
affinity for specific wintering grounds. Wintering areas in Idaho and Wyoming
were also determined by snow accumulation rather than affinity for specific
areas (Dalke et ale 1963, Patterson 1952). The high use area near California
Gulch sustained high use throughout the winter of 1974-75 but only during the
last 3 weeks of winter in 1973-74. During this 3-week period snow was melting
in the area and cover conditions were very similar to those found throughout
1974-75. The California Gulch area was the first part of North Park to become
snow free in both springs of study. Possibly the birds observed in both years
were associated with the 5 strutting arenas in the area. The other 6 high use
areas sustained heaviest utilization prior to any melting and likely were
important wintering areas. Thus, the hypothesis that distribution was directly
related to available sagebrush was accepted.
Although snow cover was the major determinant in the amount of available sagebrush, slope and aspect appeared to further restrict the amount of sagebrush
range suitable for wintering sage grouse. No data were available on relative
abundance of varying slopes although numerous steep slopes dominated by sagebrush exist. No observed use occurred on steep slopes (&gt; 30 percent) and
most observed use was on areas with less than 10 percent slope. A tendency
for utilization of southerly aspects was documented for sites with slope
greater than 5 percent. Southerly exposures receive much greater solar
irradiation (Fig. 6) which may affect the following site characteristics:
temperature, soil characteristics, moisture, and vegetation.
The prevailing southwest winds probably exert similar influences. A plausible
explanation for use of south aspects involves the direct effects of wind and
solar irradiation on snow depths and the indirect effects manifest through the
vegetation. Based on data presented, steepness of slope, modified by aspect,
eliminated many sagebrush areas from suitable habitat status. This was

�RADIATION INDEXES FOR VARIOUS FACETS AT
LATITUDE 400 N
60

x 50

IJJ

C

- 40
Z

2!

0

S
o SSE-SSW
o

~

~ 30

I

• SE-SW
Cl ESE-WSW

a::
...J
&lt;t

~ 20

z
&lt;t

10-/

0

-...JI
I-'

• E
• ENE-WNW
o NE-NW
0 NNE-NNW
• N

I
10

20

30

40

50

60

70

80

90

100

SLOPE INCLINATION, PERCENT
Figure 6. R~diation indexes for various facets at Latitude 40oN.
Radiation index i6 the ratio of the total annual potential insolation
to the maximum potential insolation at the site.
Adapted from
Frank and Lee 1966.

�-72-

probably the reason why the broken ridges south of Independence Mountain
received little use. Avoidance of steep slopes may be an adaptation against
avian predation as golden eagles (Aquila chrysaetus) were commonly seen
hunting along ridge crests, then plummeting down the slope in apparent
attempts to capture prey. Adaptations by sage grouse to avoid avian predation
during spring were discussed by Hartzler (1974).
The area of no observed use between the Canadian River and North Sand Dunes
was not an area of deep snow or steep slopes. Quantity of sagebrush above
snow was apparently equal to or greater than other areas where use occurred,
although no vegetal data was obtained from this area. Possibly quality of
sagebrush, primarily chemical composition, was a determining factor. An
interesting point is that nearly all soils within the sagebrush type of elevation below 2,743 m, are alkaline (pH&gt;7.0) except the sandy soils in this
area of no observed use where the soils were neutral (pH = 7.0) (Soil Conservation Service, unpublished soil type handbook for Jackson County). Possibly
the minerals responsible for alkalinity are needed by sage grouse in quantities
not available in the vegetation of this sandy soil area. Studies on red
grouse (Lagopus lagopus scoticus) indicate selection for plants high in
nitrogen and phosphorus (Moss 1972) while selection for calcium rich vegetation has been amply demonstrated for many species (Scott 1972).
No differences in vegetation and topography were detected between the 7 high
use areas except the one area where sagebrush cover was low. This area was
characterized by bare ridge tops with short, sparse sagebrush and swales
filled with snow. Sage grouse were observed feeding along the ridge tops but
seldom loafing or roosting. In winters of heavy snowfall with great drifting
such wind-blown areas may receive greater use as the ridges are quickly blown
bare following storms.
Several areas of occasional use had characteristics similar to the high use
areas. Whether these areas would receive high use in more severe winters is
purely speculative.
Reasons for selection of specific high use areas from
the available areas are unknown. Lack of observed use in disturbed areas
was likely a reflection of lack of sagebrush for food and cover. Utilization
of disturbed areas probably occurred until snow accumulation covered the
young sagebrush plants as evidenced by the early winter use of a crested
wheatgrass field. Dense stands of sagebrush were usually the target of vegetative control programs and possible much formerly used winter range is not
presently suitable due to slow regeneration following control measures. However, quantity of sagebrush in winter did not appear to be in limited supply.
With continued regeneration of disturbed areas, distribution of sage grouse
within North Park may change in the future, especially locations of high use.
However, areas in the south portion will continue to provide habitat only in
spring, summer, and fall. Vale (1974) summarized season use patterns of sage
grouse and observed that use of stands of less than 30 percent canopy coverage
during all periods except winter was extreme. It is possible that well
designed sagebrush control programs in the south portion of North Park may
actually benefit sage grouse by reducing plant density in such stands and thus
result in better nesting and brood rearing habitat. Maintenance of dense
stands for wintering birds is not needed in this area because snow covers nearly
all the sagebrush each winter.
Use of habitat by sage grouse has been based on observations within a given
area. Unfortunately, long term experimental manipulation of various sagebrushgrassland components has not been conducted to delineate necessary minimum

�-73-

seasonal requirements.
Such experimentation is dependent upon development of
accurate and sensitive population monitoring techniques.
The most striking feature of sage grouse flocks in winter was the strong
sexual segregation.
Segregation was as great as, or greater than that reported
for other grouse species (Koskimies 1957, Seiskari 1962, Weeden 1964, Braun
and Schmidt 1971) and the hypothesis that strong segregation occurred was
accepted. Segregation was greater in male than female flocks as evidenced by
the much higher percentage of unisexual flocks (71.6 vs. 39.5). Knowledge of
seasonal behavior of sage grouse provides a plausible explanation for this
behavior pattern. Males form flocks upon leaving leks and since most females
are involved in nesting activities at the time such flocks are characteristically unisexual. Hens unsuccessful at nesting form flocks of "bachelor"
hens or join with male flocks. Successful nesters remain with their broods
at least until early September, but little information has been published on
sage grouse life history during autumn. Much of the segregated flock formation
evidently occurs during autumn because flocks observed in late December are
strongly segregated by sex. Female flocks could not be fully comprised of
entire family units (broods) because juvenile sex ratios are nearly 50:50
(unpubl. check station data, files of Colorado Division of Wildlife).
It is
thus concluded that many immature males leave the brood structure and find
male flocks prior to winter. The fact that female flocks had one or more
males more often than male flocks had a female was probably due to some
immature males flocking with sibling groups and successful hens. The mechanism for segregation of immatures is presently unknown while the adult segregation likely occurs during mating and nesting activities.
The reason for such strong sex segregation in sage grouse winter flocks is a
question of great interest. Review of the literature indicates sex segregation by grouse is accompanied by differential habitat utilization (Koskimies
1957, Seiskari 1962, Weeden 1964, Braun and Schmidt 1971). Data frQm the
winter of 1973-74 indicate female flocks were utilizing denser stands of sagebrush than were males. Patterns of use were similar in both winters, although
differences in density and cover in 1974-75 'were not significant (P&gt; 0.10) . The
lack of statistical differences is possibly due to smaller sample size in 197475 (99 vs. 50). Although the difference in mean values for male and female
flocks was less in 1974-75, variability in data was also less. Density and
cover values were much higher in 1974-75, especially for male flocks. This is
probably a reflection of the low snowfall the second winter. Selection for
differing densities of sagebrush may also have been tempered by the mild
winter in 1974-75. Density and cover values did not increase as much for
female as for male flocks in 1974-75, suggesting that female flocks may be
utilizing sagebrush stands approaching the maximum density available in North
Park. An unfortunate weakness in the data was that measurements of sagebrush
stands were not taken at available areas of no observed use. Thus, although
female flocks were using denser stands than males, there are no data to indicate where these densities align on the density continuum of available sagebrush. Such information would be of value in evaluating the differences
observed between sexes and winters. It is inferred from the data that segregation occurs by habitat types as well as sex. Due to the variability and
short term nature of the data, judgment is reserved on the hypothesis regarding
different habitat types.

�-74-

Evaluation of mean and median flock sizes indicate females had a greater
tendency to form very large flocks (&gt;150) although most female flocks were
within the same size range as male flocks (~50).
Possibly the tendency
toward larger flock size shown by females was associated with the use of
denser sagebrush stands. Such an association
can be attributed to either
large groupings selecting dense sagebrush for cover and food or the use of
dense stands allowing the congregation of large numbers.
The large increase in flock size for both sexes during 1974-75 can possibly be
attributed to excellent production the preceding summer and/or the mild winter
of 1974-75 which resulted in more available sagebrush and thus, larger flocks.
White-tailed ptarmigan (Lagopus leucurus) apparently form more flocks rather
than increase flock size in winters following summers of excellent production
(Hoffman 1974). However, the increase in sage grouse flock size observed
between the winters of 1973-74 and 1974-75 was probably due to both excellent
production and mild winter which resulted in more extensive stands of available
sagebrush. Fewer flocks were found in 1974-75, following excellent production
in 1974, possibly the result of the more extensive winter range due to less
snowfall during 1974-75.
The large increase in observations of females during the last two weeks of
each winter was an enigma. Several possibilities exist, such as migration
of females out of North Park in late fall with subsequent return in late
winter, increased mobility of female flocks in late winter, thus increasing
change of observation, and congregation of many female flocks in staging areas
prior to the mating season. Migration is possible as movements of 80-160 km.
were documented during winter in Idaho and Wyoming (Dalke et al. 1963,
Patterson 1952). Several suitable winter ranges (Middle Park, Colorado;
Laramie and Saratoga, Wyoming) are within this distance of North Park. However, movements in Idaho and Wyoming were apparently a direct response to snow
accumulation gradually reducing the range and did not involve yearly. movements
to a specific area and a return. The winter of 1974-75 was relatively mild
and no obvious shortage of sagebrush was observed. However, quality of sagebrush may be important as well as quantity; The lack of affinity for wintering
areas within North Park tends to discount the possibility that birds within
the same population would have an affinity for an area 80-160 y~. distant.
Congregation of females at staging areas, though possible, probably was not
the cause for the observation pattern. Negating evidence was that locations
of females observed during the two-week period were different each year and
that females do not assemble in mass on strutting arenas until mid- to late
April, 6 weeks after the winter observations. Flocks of females did not remain
in these areas for 6 weeks.
A third possibility is the increased mobility of female flocks in late February. Increased mobility could be a response to environmental or internal
stimuli causing hens to seek locations of male activity. The lack of a corresponding increase in male observations suggests that males were relatively
sedentary during this period. In each of the two years, male activity on
strutting arenas commenced within two weeks of the cessation of winter studies.
Movements by females were generally greater than males during this study,
although sample sizes are not large. Unfortunately, lack of data precludes a
valid, definitive explanation of the large increase in observations of females
during late winter.

�-75Comparison of pooled winter sex ratios during each of the two winters with
harvest sex ratios was enlightening.
Sage grouse researchers have long noted
a disparity in harvest sex ratios favoring females and expressed great concern
over the assumed over harvest of females (Pyrah 1963). Such concern assumed
selectivity for smaller birds by hunters. The pooled harvest sex ratio for
North Park for the years 1955-74 was 64 females:36 males, a significant deviation «
0.05) from 1:1 and quite similar to the 62:38 and 61:39 ratios
observed-during the winters of 1973-74 and 1974-75, respectively. The sex
ratio of immature birds in the harvest in 1974 in North Park was not significantly different (P&gt; 0.10) from 1:1 and juvenile birds comprise approximately
50 percent of the population in fall. Thus, the disparity in adult sex ratios
may result from differential natural mortality rather than differences at
hatching or hunter selection. Apparently, differential natural mortality
favors females and hunter selectivity is less than formerly believed.
Two major points should be discussed in reference to management of sage grouse
in North Park. First, distribution of sage grouse is limited to the north
portion, and specifically to the northeast quarter of the park in severe winters.
The northeast area is underlain by extensive low sulfur coal deposits (Beekly
1915) and thus, may be subject to extensive vegetation disturbance should strip
mining operations develop. Because of the slow regeneration of disturbed
sites, caution should be taken to conserve suitable sagebrush in these areas.
If mining occurs, provisions to limit the acreage of total disturbance should
be initiated to preserve critical winter habitat. Human activity in this area
during winter should also be minimized.
Second, there was no evidence that the gross quantity of sagebrush available
for sage grouse was limiting. However, the quantity of sagebrush of sufficient
nutritional quality is not known. Such information, from a management viewpoint, may be important. If areas of sagebrush of insufficient nutrient quality
do exist, then procedures can possibly be implemented to improve the quality,
thus increasing the quantity of suitable habitat.
Reducing the extent of dense sagebrush stands in the south part of the park
could possibly provide more nesting habitat and summer range and result in
better utilization of public land. It is questionable that either improvement
of sagebrush rangelands by fertilization or reduction of large blocks of
dense sagebrush are feasible management procedures since hunting pressure is
not presently adequate to remove the annual surplus.

Prepared

�-76-

LITERA

Autenrieth,
Prog.

R. E.
Rep.

TURE

1969.
Sage grouse
Idaho Fish &amp; Game

Bailey,

A. M.
1925.
Segregation
Condor 27(4):172-173.

Beekly,

A. L.
Colorado.

1915.
U.S.

CITED

investigations.
Dept.
25 pp.

of the sexes

Third
Mimeo.

in the sage

Annu.

hen.

Geology and coal resources
of North
Geol. Surv.
Bull. 596.
121 pp.

Park,

Beetle,

A. A.
1960.
A study of sagebrush.
The section
Tridentatae
of Artemisia.
Wyo. Agr. Exp. Sta , Bull. 368.
83 pp.

Bent,

A. C.
birds.

Braun,

C. E. and R. K. Schmidt,
Jr.
1971.
Effects
of snow and wind
on wintering
populations
of white -tailed ptarmigan
in Colorado.
Pages 238 -250 in A. O. Haugen,
ed , Proc.
Snow and Ice Symp.,
Iowa State University,
Ames.

Carr,

H. D.
1967.
Effects
of sagebrush
spraying
on abundance,
distribution
and movements
of sage grous e. M. S. Thesis.
Colorado
StateUniv.,
Ft. Collins.
106pp.

Dalke,

P.

D.,

1932.
Life his tories
of North American
gallinaceous
Smithsonian
Inst.,
U.S. Nat. Mus. Bull. 162. 490 pp.

D.

B.

Pyrah,

E. F. Schlatterer.
ment of sage grouse
Daubenmire,
Agric.

D. C. Stanton,

J.

R.
1970.
Steppe vegetation
Exp. ses. Tech.
Bull. 62.

Crawford,

of Washington.
131 pp.

Eng,

R. L., and P. Schladweiler.
1972.
Sage
ments and habitat use in central
Montana.
36(1}:141-146.

Finch,

W. C., ed.
Middle Parks
Denver.

1957.
Basin,

E.

1963.
Ecology,
productivity
in Idaho.
J. Wildl. Manage.

Guidebook
Colorado.

and

and manage27(4):81! -841.
Washington

grouse
winter moveJ. Wildt. Manage.

to the geology of North and
Rocky Mtn. Assoc.
Geol.,

�-77-

Frank,

E. C., and R. Lee.
on slopes.
U.S. For.

1966. Potential solar beam irradiation
Servo Res. Paper RM-18.
116 pp,

Gill,

R. B. 1965. Distribution and abundance of a population of sage
grouse in North Park, Colorado.
M. S. Thesis.
Colorado State
Univ., Ft. Collins.
187 pp.

Griner,
L. A. 1939. A study of the sage grouse (Centrocercus
urophasianus),
with special reference to life history, habitat
requirements,
and numbers and distribution.
M.S. Thesis.
Utah State Univ., Logan. 83 pp,
Hail,

W. J. 1965. Geology of northwest
Geol. Surv. Bull. 1188. 133 pp.

Hartzler,
J. E. 1974. Predation
leks.
Auk 91(3}:532-536.

North Park,

Colorado.

U.S.

and the daily timing of sage grouse

Hoffman, R. W. 1974. Characteristics
and migration of wintering
populations of Colorado white -tailed ptarmigan.
M. S. Thes is.
Colorado State Univ., Ft. Collins.
58 pp.
June,

J. W. 1967. Sage, blue and ruffed grouse sexing and aging
characteris tics.
Wyoming Game &amp; Fis h Comma Cheyenne.
59 pp.

Klebenow, D. A. 1969. Sage grous e nes ting and brood habitat in
Idaho. J. Wildl. Manage. 33(3):649-661.
Koskimies,
J. 1957. Flocking behavior in capercaillie,
Tetrao_
urogallus (L.), and blackgame,
Lyrurus tetrix (L.).
Finnish
Papers on Game Res.
18. 32 pp.
May, T. A. 1970. Effects of sagebrush control on distribution and
abundance of sage grous e. Colo. Div. Game, Fish &amp; Parks,
Game Res. Rep., Fed. Aid Proj. W-37 -R. April.
pp. 115138.
Moss,

R. 1972. Food selection by red grouse (Lagopus lagopus
scoticus) in relation to chemical composition.
J. Anim. Ecol.
41(2}:411-428.

Patterson,
Inc.

R. L. 1952. The sage grouse
Denver.
341 pp ,

in Wyoming.

Sage Books.

�-78-

Poley,

B. E. 1969. Effects of sagebrush control on distribution and
abundance of sage grouse.
Colo. Div. Gam e, Fish &amp; Parks,
Garne Res. Rep., Fed. Aid Proj. W-37 -R. April.
pp. 61 -85.

Pyrah,

D. B. 1963. Sage grous e i nves tigations.
Idaho Fish and Game
Dept., Wildlife restoration Div. Job Cornpl , Rep , , Fed. Aid
Proj. W-125 -R. 71 pp.

Rogers, G. E. 1964. Sage grous e inves tigations in Colorado.
Colorado Garn e, Fish &amp; Parks Dept. Tech. Publ. 16. 132 pp.
Scott,· M. L. 1972. Nutrition in reproduction - direct effects and
predictive functions.
Pages 46-59 in D. S. F'ar-rner, ed.
Breeding biology of birds.
Natl. Ac ad , Sci. Washington, D. C.
Seiskari,
P. 1962. On the winter ecology of the capercaillie.
Tetrao
urogallus,
and the black grouse, Lyrurus tetrix, in Finland.
Finnish Papers on Garne Res. 22. 119 pp.
Srni.th, E. L. Jr.
1966. Soil-vegetation
relationships
of some
Arterrlisia types in North Park, Colorado.
Ph. D. Dissertation.
Colorado State Univ., Ft. Collins.
203 pp.
Vale,

T. R. 1974. Sagebrush conversion projects:
An el ern enj; of
c ont ernpor a r-y env i r onm enta l change in the western U. S.
Biological Cons ervation 6(4) :274 -284.

Weeden, R. B. 1964. Spatial separation of s exes in rock and willow
pta rrn iga n in winter.
Auk 81 (4}:534 -541.

�April 1975

-79-

JOB FINAL REPORT

State of

~C~OL~O~RAD~O=_

_

Project No.

~W_-~3~7_-~R~-=2~8

_

Work Plan No.

=l~O

_

Game Bird Survey
Job NO.

-=2

_

Job Title.__~E~x~p~e~r~i~m~e~n~t~al~~B~r~e~e~d~l~·n~g~o~f_H~un~g=a=r=i=a=n~P~a~r~t~r~i~d~g~e~
_
Period Covered:
Personnel:

April 1964 to March 1973

Many individuals have contributed to the completion of this study.
For a complete list see Hoffman, 1973a in Literature Cited of the
special report listed below.

ABSTRACT

The objectives of this study have been completed, and manuscript written,
and results were published in the Division's Special Report series as
follows:
Hoffman, D. M., C. L. Quarles, L. A. Webster, and L. R. Crooks. 1974.
Experimental propagation of gray partridges.
Special Report No. 35,
Colorado Division of Wildlife, Denver. 26 p.

/}

yjl

Cf70~

~/

Prepared by __~c:::\t"-=-ti~~..::z.:.:a:::-i.:::v,_±_t--..:./~
/7~
/-,v~' /-=-~.7/9~·'+-"'::;':".y"..:.· -=.:::z:..-,=.:r_·~=' =-.'__
Donal d M. Ho ffman 2~~
Wildlife Researcher

��April 1975

-81JOB PROGRESS REPORT

State

0

f

Project No.__
Wo rk Plan No .,

..::CO~L~O~RA=DO~

_

~W_-~3~7_-~R~-=2~8

_

--=1:-:.7

_

Game Bird Survey
Job No.

2
-----------=~----------------

Job Title~~C~o~n~t~i~n~u~~ed~I~n~v..::e~n~t~o~ry~_o~f~S~e~l..::e~c~t~e~d~P~t~a~rm~i~g~an~~P~o~p~u
_
Period Covered:
Personnel:

April 1, 1974 to October 20, 1974

W. John Arthur, Clait E. Br aun , Richard W. Hoffman, Tom Lytle and
Ron Oakleaf.

ABSTRACT

Investigations of white-tailed ptarmigan (Lagopus leucurus) populations in
Colorado initiated in 1965 were continued in 1974 with major emphasis in
ascertaining population levels and relationships of nesting success and production to climatic conditions. Breeding densities in 1974 were similar to
those recorded in 1973 on all areas. Thus the decline in breeding densities
first observed in 1970 halted. Reasons for attainment of stability are not
fully lll1derstoodbut were believed due to the hunting closure in 1973 at Mt.
Evans, low hlll1terpressure at Crown Point and Independence Pass and increased
survival of females in Rocky MOlll1tainNational Park. Nesting success was poor
(Independence Pass), to good (Crown Point and Rocky MOlll1tainNational Park)
in 1974. Failure of nests at Independence Pass was possibly related to age
of females while success at Crown Point and Rocky Molll1tainNational Park was
possibly related to weather conditions which were warmer throughout the
summer than normal, with below average moisture in May. Apparent hunting
pressure was low at Crown Point and Independence Pass. At Mt. Evans, the
season was opened for 3 days and hunters harvested an estimated 32 percent
of the fall population.

�-82RECOMMENDATIONS

1.

It is recommended that
for preparation
of the

2.

It is recommended that data from Work Plan 17, Jobs 1, 2, and 3 be
incorporated
into two major manuscripts.
Tentative
titles
of these
manuscripts
would be: "Population
Dynamics of Hunted and Non-hunted
Populations
of Uhite-tailed
Ptarmigan in Colorado",
and "Relationships
of {&lt;leather and Age of Females to Nesting Success of White-tailed
Ptarmigan" .

this project
final
report.

be extended

until

March 31, 1977

�-83-

CONTINUED INVENTORY OF SELECTED PTARMIGAN POPULATIONS
C1ait E. Braun

Knowledge concerning population levels of white-tailed ptarmigan and factors
influencing observed densities is essential for proper management.
This
project was undertaken to examine long term population trends and effects of
climatic conditions on white-tailed ptarmigan in Colorado. The initial 5
years of research on this grouse have been previously reported (Braun and
Rogers 1971). Progress reports on work since 1969 have been prepared (Braun
1971, 1972, 1973, 1974).
P. S. OBJECTI VES
To test the hypotheses that (1) populations of white-tailed ptarmigan in
Colorado are not cyclic, (2) production and fall population densities of
this species in Colorado are related to temperature and moisture conditions
during June and early July of the same year, and (3) harvest of over 50
percent of the fall population adversely affects breeding densities the
following spring.
SEGMENT OBJECTIVES
1.

To estimate breeding population levels of ptarmigan in 4 study areas (Ht.
Evans, Crown Point, Independence Pass, and Rocky Mountain National Park).

2.

To estimate nesting success and production in the above areas:

3.

To obtain weather data from a representative alpine area for use in
determining correlations between spring weather conditions and fall ptarmigan populations.

4.

Manipulate hunting seasons at Mt. Evans through closures, reduced length,
late timing, etc., in order to affect breeding densities.

5.

To compile data and prepare progress report.

METHODS AND MATERIALS
Techniques used were essentially those developed under Work Plan 17, Job 1
and reported in detail by Braun and Rogers (1971), and updated by Braun (1971).
In 1974 a combination of blue and white bandettes with black numerals were
used to mark all newly banded birds and those whose older bandettes had
become too worn for individual recognition on all areas except Crown Point.
At this area, only white bandettes were used. Weather data were obtained from
the Institute of Arctic and Alpine Research (INSTAAR), University of Colorado,
for reasons previously explained (Braun 1971). Hunting statistics were collected through return of bands from successful hunters (Independence Pass and
Crown Point) and a check station on the only access road at Mt. Evans.

�-84-

Description of Area
Areas intensively investigated have been described in detail by Braun and
Rogers (1971) and have been presented by Braun (1971).

RESULTS AND DISCUSSION
Breeding Densities
Censuses of breeding birds were initiated on 9 May at Mt. Evans, 10 May at
Independence Pass, 16 May in Rocky Mountain National Park and 17 May at Crown
Point. Pairing was still occurring as late as 17 May, with most pairs being
formed by 20 May. Timing of breeding events was similar to that of previous
years, but was somewhat earlier (about 5 days) than in 1973. Densities
observed are presented in Table 1.

Table 1.

White-tailed ptarmigan breeding denSities, all areas, 1974.

Study Area

No. of
Breeding
Pairs

Unmated
Males

Total
Breeding
PopUlation

Birds per
Square
Mile

6

5

17

13.6

.19

4

5

13

68.4

.70

6

2

14

20.0

2.14

16

12

44

20.6

1.93

2

3

7

3.6

1.54

11

3

25

16.2

1.12

5

8

18

16.1

S.
~ze-1/

Rocky Mountain National Park
Tombstone Ridge-Sundance
Mountain
Toll Memorial
Fall River Pass
Total - Rocky Mountain
National Park
Crown Point
Mt. Evans
Independence Pass

1.25

1/ In square miles.

Changes in breeding densities between 1973 and 1974 were not significant.
Densities at Mt. Evans decreased slightly, an unexpected event since the area
had been closed to hunting in the fall of 1973. However, nesting Success and
production in 1973 were poor on all study areas including the adjacent Guanel1a
Pass area CR. Hoffman, pers. comm.). Thus, few yearling birds were available

�-85-

for recruitment into the spring population. Densities at Crown Point and
Independence Pass were also slightly down, despite low hunting pressure and
poor hunter success in 1973. Only at Rocky Mountain National Park did
densities increase (only one bird increase in total numbers). Of significance,
however, was the stabilization of spring to spring mortality rates in Rocky
Mountain National Park. Breeding densities at all study areas from 1966
through 1974 are presented in Table 2.
Nesting Success and Production
In 1974, the month of May in alpine areas was dry but cool (Table 3). While
snow melt was retarded, preparation for nesting by females was advanced
because of early plant phenology and abundance of bare ground (due to low
overwinter snowfall). Some females started laying in late May and most started
incubating by 9 June when a major snowstorm occurred, depositing over one
foot of snow in alpine areas. Temperatures were warmer in June 1974 than in
June 1973; consequently, the snow rapidly melted and the effects of the 9 June
storm were not disasterous in terms of nesting success. Some hens apparently
renested following the storm, but most continued nesting during the period
the ground was snow covered. If nesting events had not been advanced in 1974
the 9 June storm would have occurred when most hens were laying and the effects
of the storm would have been pronounced. One nest located on 13 June (by T. A.
May) when incubation was well underway, hatched on 3 July. Estimated peak of
hatch in 1974 was between 5 and 10 July, almost two weeks earlier than in 1973.
Nesting success was good at Rocky Mountain National Park (16 of 26 hens observed were successful), and Mt. Evans (5 of 8 hens observed were successful).
Only 2 hens were observed at Independence Pass during the brood period and
one was successful. At Crown Point only one of the two hens known to have
been present was observed during the brood period and this hen was successful.
Overall nesting success for 37 hens observed during the brood period was 62.2
percent. This is a substantial increase from the estimated 25 to 30 percent
success documented in 1973.
Initial brood size in 1974 is not exactly known but was believed to be between
5 and 6 chicks. Survival of chicks to 1 September was excellent (Table 4) and
dispersal did not occur until about 20 September. Data from 1974 concerning
brood size to 1 September are similar to most years of the study. However,
average brood size and number of broods was greater in 1974 than in 1973.
Fall Densities
Estimates of densities of white-tailed ptarmigan on September 1, while useful
in illustrating population gain through production, are difficult to accurately derive due to a number of variables. Estimated fall densities for each
area studied are presented in Table 5. These densities were calculated
following the three basic assumptions discussed in detail by Braun and Rogers
(1971). Percent gain was more than adequate at all areas except Independence
Pass to maintain and increase populations the following spring provided overw.Ln ter mortality was not greater than in most years.

�Table 2.

White-tailed

ptarmigan

breeding

densities

1966-74.

1966

1967

1968

Birds Per Sguare Mile
1969
1970
1971

1972

1973

1974

29.0

25.2

29.4

30.8

24.8

23.4

22.4

20.1

20.6

14.5

21.2

18.1

14.5

10.4

5.7

1.0

4.7

3.6

7.8

7.1

7.1

5.8

5.2

11.0

19.5

16.2

16.2

18.7

18.7

18.7

17.9

10.7

21.4

18.7

17.0

16.1

------Study

Ar ea

Rocky Mountain

National

Park

Crown Point
Mt. Evans
Independence

Table 3.

Pass

Weather

data, Niwot Ridge, 12,300 ft, May-July,

1974.
I
00

Month

Maximum
0

Temperature
Mean Daill
Minimum
Maximum
Minimum

Number Days
Minimum
Temperature
Less Than 32° F

0'\

Precipitation/
Month
(Inches)

I

Number Days
Relative
Humidity
90% or Above

Wind Speed
Month
(mph)

May

54

F

8° F

40.8° F

25.2° F

26

1. 63

13

June 2:./

1/

690

F

12°F

51.0° F

35.0° F

5

3.92

15

1/

60° F

35° F

53.0° F

41.0° F

0

3.76

24

Y

July

1/
- Data not available.
2/
- Data are not available

for 17 June.

�-87-

Table 4.

Number of broods and average brood size, all areas, 1974.1/

Number of Broods
Observed

Average Number of
Chicks per Brood

July

4

4.2

August

18

3.8

September

5

4.2

Month

1/ Only distinct broods are included.
Hunting MOrtality
In 1974 all alpine areas of the state were open to ptarmigan hunting from
14 September through 8 October, except for the area lying within 1/2 mile of
the Mt. Evans highway. This area was open only from 17 through 19 September.
As in recent years, no ptarmigan hunting was allowed during any of the early
big game seasons. Bag and possession limits for ptarmigan were 3 and 6.
No bands were received from hunters in the Crown Point area in 1974. Thus,
hunting mortality at this site in 1974 appears to have been non-existent.
At least nine ptarmigan were harvested at Independence Pass in 1974 of which
3 were banded. This represents 14.3 percent of the 21 banded birds known to
be available to hunters at this site. This value is considered minimal as it
now appears that a substantial number of hunters at this site fail. to report
bands in the year the birds were shot. The best estimate of the fall hunting
mortality of this population in 1974 is from 30 to 40 percent.
At Mt. Evans a check station was operated from dawn to dusk all three days of
the season on the only access road into the area. All hunters going into the
area were recorded and accounted for each day. A check was made each evening
to insure that no hunters were still in the area after closure of the check
station. In all, 104 hunters were checked who hunted 289 hours, observed 53
ptarmigan, of which 22 were bagged. Eleven of the 22 ptarmigan bagged were
banded, thus an estimated 32 percent (11 of 34 banded birds known to be present)
of the fall population was harvested. This approximates the one-fourth to
one-third of the fall population loss to hunting that an individual population
of ptarmigan can sustain without detriment to the subsequent breeding population. Of the 22 ptarmigan harvested, 6 were young of the year, five were
adult males and 11 were subadult and adult females. This does not adequately
represent the population due to differential behavior patterns.

�Table 5.

Estimated

fall densities

of white-tailed

Estimated
Percent
Nesting
Success

Average
Brood Size
on
September 1

Rocky Mountain
National Park

60

Crown Point

Area

Mt. Evans
Independence

Pass

ptarmigan,

1974.

Total
Production

Total
Breeding
Population

Total
Population
on
September 1

4.0

40

44

80

37.4

50.0

50

4.0

4

7

10

5.2

40.0

60

4.0

28

25

50

32.5

56.0

40

2.5

5

18

21

18.7

23.8

Birds Per Percent
Square Mile Gain

I

co
co
I

�-89-

LITERATURE CITED

Braun, C. E. 1971. Continued inventory of selected ptarmigan populations.
Colo. Div. of Game, Fish and Parks, Game Res. Rept. Fed. Aid Proj.
W-37-R. April. p. 91-106.
. 1972. Continued inventory of selected ptarmigan populations. Colo.
Div. of Wildl. Game Res. Rept. Fed. Aid Proj. W-37-R. April. p. 91-98.
1973. Continued inventory of selected ptarmigan populations.
Colo.
Div. of Wildl. Game Res. ~pt. Fed. Aid Proj. W-37-R. April. p. 355361.
_____

1974. Continued inventory of selected ptarmigan populations.
Colo.
Div. of Wildl. Game Res. Rept. Fed. Aid Proj. W-37-R. April. p. 65-72.

_____ , and G. E. Rogers. 1971. The white-tailed ptarmigan in Colo.
Game, Fish and Parks Tech. Publ. No. 27. 80 p.

Prepared by

u_h_::J_/'_J_;-_... --1t.&amp;~/....:..--Z4tUU-,-,Clait E. Braun
Wildlife Researcher

_

Colo.

��April 1975
-91-

JOB FINAL REPORT

State of

COLORADO

Project No. ~W~-~3~7_-~R~-~2~8
Work Plan No. __~1~7
Job Title
Period Covered:
Personnel:

_

Game Bird Survey

_
3
Experimental Removal of a Breeding Population
of White-tailed Ptarmigan

April 1, 1971 to March 31, 1975

Clait E. Braun, Richard W. Hoffman and Terry A. May

ABSTRACT

All white-tailed ptarmigan (Lagopus leucurus) located were removed by shooting
from Crown Point, a 1.93 sq. mile isolated study area in northern Colorado,
in early June 1971. No replacement occurred in 1971 and only two males were
present in 1972. Numbers of birds present during the breeding season slowly
increased to removal levels in 1973 before decreasing in 1974. The first
successful nesting occurred in 1974; three years after all breeding birds
were removed. Repopulation of the area was slow due to the affinity of
yearling male ptarmigan to natal areas and the distance between the area
studied and surrounding occupied habitats. Presence of suitable males (adults)
was most critical in attracting females to the area. Male ptarmigan selected
areas for breeding which were at least 40 percent snowfree by mid May in
which bushes of Salix greater than 10 but not exceeding 26 inches in height
were a major component of the vegetation. Wind was the most important factor
affecting location of breeding territories.

�-92-

RECOMMENDATIONS

1.

It is recommended that data from Work Plan 17, Job 3 be incorporated
with that from Work Plan 17, Jobs 1 and 2 into a manuscript titled
"Population Dynamics of Hunted and Non-hunted Populations of White-tailed
Ptarmigan in Colorado". An oral presentation with abstract only to be
published discussing the results of this job would also be desirable.

2.

If white-tailed ptarmigan are to be released in isolated mountain ranges
where they do not occur, evaluation of the release site(s) should include
estimation of bare ground present by 15 May and actual measurement
of the height of available bushes of Salix. These criteria should then
be compared with results of this study to determine suitability. Areas
smaller than 3 sq. miles of suitable habitat will probably not support
huntable populations on a sustained basis. Releases should include
approximately 50 percent each adults and subadults of both males and
females, in both fall (August-September) and spring (May-June). Time of
initial release (either fall or spring) is not important for establishment.

�-93-

EXPERIMENTAL REMOVAL OF A BREEDING POPULATION
OF WHITE-TAILED PTARMIGAN
Clait E. Braun

Management of wild animals is dependent upon knowledge about population
levels, reproductive capacity and success, survival and mortality rates, and
habitat requirements. Without adequate habitat for any given species there
would be no need to investigate other aspects of the biology of a given
animal. In addition to knowledge about the particular habitat requirements of
a species, knowledge must also be available on the pioneering abilities of a
species and age of pioneers.
In recent years intensive studies of grouse (Family Tetraonidae) have focused
on the apparent non-breeding segment of the male population. Non-displaying
males have been documented for red grouse (Lagopus lagopus scoticus) (Watson
and Jenkins 1967), black grouse (Lyrurus tetrix) (Robel 1969), blue grouse
(Dendragapus obscurus) (Bendell et al. 1972; Zwickel 1972), sharptailed grouse
(Pedioecetes phasinellus) (Rippin and Boag 1974) and ruffed grouse (Bonasa
umbellus) (Dorney and Kabat 1960; Boag and Sumanik 1969; Rusch and Keith 1971;
Fischer and Keith 1974). Non-territorial males of white-tailed ptarmigan
were observed by Braun (1969) and Schmidt (1969) but whether or not these
males would become territorial upon removal of established males was not tested.
Habitats seasonally occupied by white-tailed ptarmigan in Colorado have been
described (Braun 1971b). Analyses of ptarmigan movements in relation to
vegetative communities suggest that ptarmigan seek and utilize certain areas
depending upon season and that the same patterns prevail throughQut the
alpine areas of Colorado (Braun 1969). Since number of suitable breeding
territories may control breeding densities, potential productivity, and fall
densities, an experiment was designed to investigate the exact requirements
for breeding territories. Crown Point-Crown Mountain was chosen as the site
for the experiment, as it is somewhat isolated from adjacent alpine areas,
thus reducing the possibility of ingress between the breeding period and the
early fall. This area had been studied from 1966-70 (Braun and Rogers 1971;
Braun 1971a), and locations of most previously occupied territories were known.
Additionally, hunters removed about 50 percent of the fall population in 1970
(Braun 1971a), and it was anticipated that the population would be low in
the spring of 1971. This would reduce the number of birds to be removed.

P. S. OBJECTIVE
To test the hypotheses that (1) ptarmigan select sites for breeding territories which contain readily available bushes of Salix spp. (&gt; 10 inches in
height) and snowfree locations, (2) immature birds initially pioneer new or
empty habitats, and (3) number of occupied territories is affected by age
class of male ptarmigan.

�-94-

METHODS AND MATERIALS
Ptarmigan were located during the breeding and brood seasons through use of
tape-recorded male challenge and chick distress calls as described by Braun
et al. (1973). At least two censuses were conducted each year during both the
breeding (May-June) and brood (August-early September) seasons. Locations
of all birds observed were plotted on standard 7.5 minute U. S. Geological
Survey topographic maps (scale 1:24000). In 1971, all birds located were
immediately shot with a .410 shotgun. After 1971, all birds observed were
pursued until caught or flushed out of view. Capture was accomplished with
22-foot telescoping fiberglass fishing poles (Zwickel and Bendell 1967).
Each bird was banded with a gold aluminum, serially numbered band and
individually numbered plastic bandettes. Colors used were pink in 1972, green
in 1973 and white in 1974. All vegetation mapping was accomplished in 1967-68
following proced~res outlined by Kuchler (1955) as presented by Braun (1969).

LOCATION OF STUDY AREA
The Crown Point-Crown Mountain area (hereafter referred to as Crown Point)
is on the Poudre District of the Roosevelt National Forest, approximately
six miles north of the north edge of Rocky Mountain National Park (T8N,
R74W, parts of Sections 27, 28, 33, 34 and 35, Larimer County). This area,
at the northeast end of the Mummy Range, lies at 105°37' W long. and 40°37' N
lat. The area studied, as planimetered from U. S. Geological Survey topographic
maps, scale 1:24000, was 1.93 square miles (Fig. 1).

RESULTS AND DISCUSSION
Population Characteristics
In 1971, breeding surveys at Crown Point were delayed in order to have assurance that territory formation and pairing was complete. Since Schmidt (1969)
reported that some territory establishement and pairing may occur as late as
25 May, delay was deemed essential if all birds were to be removed. On 2 June,
removal of all birds located (nine) was accomplished.
Subsequent surveys
during June, July and August did not result in the location of any ptarmigan
or recent sign (feathers or droppings). No ptarmigan were reported harvested
on this site in 1971.
The nine birds removed consisted of 3 pairs and 3 unmated males. Six of the
nine were banded. Ages of the males were: 6+ (1), 4+ (1), 4- (1), 3- (2)
and 2+ (1). Ages of the three females were 3- (1), and 1- (2). All of the
males were territorful as determined by response to tape-recorded calls and
all but one had been on the same breeding territory for at least two consecutive years.
In 1972, the study area was systematically surveyed 3 times during the breeding
season and twice during the late summer brood period. Only two males were
located of which one was captured and banded. This bird was a subabult (1-).

�-95-

CROWN

f

o
E+3

Fig. 1.

Crown Point Study Area.

POINT AREA

COLORADO

N

-

Study Area Boundary
SCALe: IN MILlS
I/I!

ES3

F*3

E*3

�-96-

The other male was repeatedly observed but could not be captured. This
individual exhibited the characteristics of an unmated, non-territorial
subadult. Male P3l was territorial by early June.
During 1973, the Crown Point area was systematically surveyed 3 times during
the breeding season and twice during the late summer brood period. A total
of 13 different birds were observed of which 8 were newly banded. Six males,
occupied territories in 1973 of which 1 was banded as a subadult in 1972,
1 was banded as a subadult in 1973, and 4 were unmarked. These four males
were extremely wild and were thought to be subadults based on behavior and
reaction to the tape-recorded male challenge call. It is possible that one
of these males was the unmarked bird observed in 1972. Three of these males
were unmated.
Seven females were banded on the study area, and 2 additional
unmarked hens were observed in early May. Only 3 hens became paired and
remained on the study area. All other females disappeared in late May and
were presumed to have moved west into the Mummy Range in search of suitable
males and/or territories. None of the three hens that became established was
successful in bringing off a brood. Observations indicated that most, if
not all birds had left the area by early August.
As in previous years, the study area in 1974 was systematically searched 3
times during the breeding season and twice during the late summer brood
period.
Eleven birds were identified of which four were young produced
in 1974. During the breeding season, the presence of only 7 birds (2 pairs
and 3 unmated males) could be documented. Four of these birds (3 males
and 1 female) were not previously marked and could not be captured. They
possibly were subadults although the males may have been those observed but
not marked in 1972 and/or 1973. No marked birds from either 1972 or 1973 were
observed in 1974. Of the three birds captured during the breeding season,
one was a subadult (male) while two were adults (one male and one female).
The adult male could have been one of thos~ that could not be captured in 1973
as it occupied a territory that had been defended in 1973 by an unmarked male.
It is reasonably certain that the adult hen marked in 1974 was not present
on the area in 1973. All of the birds except two adults (marked) and the
four chicks (marked) produced in 1974 had left the area by early September
1974. The nesting success documented in 1974 was the first at this site since
before the removal experiment began (1970).
Population statistics from the four years of study are given in Table 1.
Several problems arose during the study that were not initially anticipated.
These were the inability to capture all pioneering birds each year and the
inability of birds to become established. The limited data indicate that
males are most important in re-establishment of a population. Extensive
banding data (chicks) from all areas studied from 1966 through 1974 indicated
that males banded as chicks initially return to their natal areas in search of
territories while females move substantial distances (from 1 to over 15 miles)
before becoming established (Braun unpublished data). Once subadult males are
not successful in becoming established on areas close to where they were chicks,
some then disperse into adjacent habitats in search of vacant territories.
Those moving the farthest are the least dominant.

�-97-

Table 1.

Population statistics of white-tailed ptarmigan, Crown Point, 1971-74.

Year

Number of
Males
Females
1- 2+ UnK
1- 2+ Unk

1971

0

6

0

2

1

0

3

3

9

4.7

1972

1

0

1

0

0

0

0

2

2

1.0

1973

1

1

4

3

0

0

3

3

9

4.7

1974

1

1

3

0

1

1

2

3

7

3.6

No. of
Pairs

No. of
Unmated
Males

Total
Breeding
Population

Birds Per
Square Mile

Since reproduction did not occur at Crown Point from 1971 through 1973, there
were no subadult males produced on the area available to occupy vacant territories.
Thus all males pioneering to Crown Point had to move at least two miles from
other occupied habitats west of the study area. Further, males arrived
at Crown Point late (mid Mayor later) after most females had become established
on territories elsewhere. Data from other areas in Colorado and Schmidt (1969)
indicate that females do not become territorial until a suitable territory
occupied by a male is located. Since most males arrived at Crown Point
after the peak period of hen dispersal, few pairs were formed at this site.
It is possible that more pairs would have been formed if experienced (2+)
males had been present. It would appear then that distance from other occupied
habitats and age of males are of major importance in re-establishment of
breeding populations of white-tailed ptarmigan. Therefore the hypotheses
that immature birds initially pioneer new or empty habitats and number of
occupied territories is affected by age class of male ptarmigan are accepted.
Since production occurred at Crown Point in 1974 and was considered good to
excellent elsewhere in northern Colorado in 1974 (Work Plan 17, J 2), it
is expected that number of birds at this site will significantly increase
in the spring of 1975.

Description of Breeding Territories
Vegetative type mapping of the Crown Point Study Area was accomplished in
1967-68 under a grant from the National Science Foundation (Braun 1969).
These data are included for descriptive purposes in Figure 2 and Table 2.
Territories occupied during the 1966-1971 period are given in Figure 3.
Territories occupied during the 1972-74 period are shown in Figure 4. Every
territory was not occupied each year. From 1966 through 1971, 15 different

�-98-

territories were occupied for at least one breeding season. During the
1972-74 period, seven different territories were occupied during at least one
spring. Of the 7 territories on Crown Point itself during 1966-71, none
was reoccupied in 1972-74 while 7 of the 8 former territories on Crown Mountain
were occupied. None of the males during 1972-74 had prior experience in the
Crown Point Area and initially could not have "learned" from other males
about locations of territories. Thus it is significant that old territory
sites were selected on Crown MOuntain and no territories were established
on Crown Point. This strongly suggests that Crown Point is poorer habitat
for breeding birds than Crown Mountain, a hypothesis developed during 1966-71
that was not tested. Also pioneering males selected sites for territories
that had been previously utilized by other males. It is probable that
pioneering males were selecting territories by aspect and structure of the
vegetation.
Territories selected for and those used previously were principally located
along the upper edge of the Picea-Salix krummholz, Dryas-Picea-Salix rock
krummholz and Salix-Picea-Trifolium-Poa krummholz (Units C-3, C-8 and C-15)
Figure 2, Tabl~
While the vegetative composition of these units differ,
the structure of each is similar and Salix is a dominant. Of importance is
the observation that wind action along the upper fringe of the communities
prevents snow accumulation. Thus the edges of these communities are among
the first areas to become snowfree in late April and early May. Some of these
areas remain essentially snowfree throughout the year except following
major periods of snowfall.
Systematic observations of percent of ground covered by snow and height of
bushes of Salix were recorded for 10 territories used at least during two
consecutive years. Bare ground approximated at least 40 percent of ~ach
territory by 15 May during each year. Removal of snow cover was primarily
by wind action prior to 15 May and by melting after that date. Measurements
of 10 randomly selected bushes of Salix within each territory gave a range
of mean heights of Salix bushes per territory from 11.05 to 13.03 inches
(Table 3). Ranges in height of bushes within territories were from 4.0 to
26.0 inches. These data support the hypothesis that ptarmigan selected
sites that were partially snowfree by 15 May which were dominated by bushes of
Salix that were 11 to 12 inches in height. Typically these sites were near
the crest of ridges where some protection from the prevailing wind (westerly)
was available but where snow depths were not great enough to completely cover
all bushes of Salix. It would appear that wind direction indirectly controls
location of breeding territories of white-tailed ptarmigan through its
effects on plant distribution and height and distribution of snow.

�-99-

Fig. 2. Vegetation units, Crown Point.
C-l

Salix-~-~

Alpine Meadow

C-2

Poa-~

Alpine Meadow

C-3

Picea-~

Krummholz

C-4

~-Poa

C-5

Trifolium-~-~

C-6

Trifolium-~

C-7

Silene-Trifolium-Carex-PoaFellfield

C-8

Dryas-Picea-~

C-9

Dryas Turf

C-lO

~-~

C-ll

~

C-12

Dryas-Pinus Subalpine Meadow

C-13

Sibbaldia Snow Accumulation Area

C-14

~-~-Betula

C-15

Salix-Picea-Trifolium-PoaKrummholz

C-16

Artemisia-Trifolium Snow Accumulation Area

Alpine Meadow
Alpine Meadow
Alpine Meadow

Rock Krummholz

Krummholz
Subalpine Meadow

Alpine Meadow

�CROWN POINT

I
I-'

o
o
I

�-101-

fable 2.

Descriptio':',
of vegetation units, Crown Point.

UNIT
C-l

DESCRIPTION
Salix-Poa-Carex Alpine Meadow: Salix spp. and Poa spp.
each comprise 5-25 percent of the total coverag~with
Carex app. accounting for at least 20 percent of the
cover. Shrubs are less than 1 m in height, with
herbaceous vegetation being less than 1/2 m tall.
Plant density is primarily continuous.
Rocks cover at
least 5-25 percent of the area, with small areas approaching 75 percent rock cover. Rocks larger than 12
inches in diameter are dominant. Lichens are about
equally green and black and cover up to one-half the
surface of rocks. Percent slope is variable and averages about 10 percent.

tot;I

C-2

Poa-Carex Alpine Meadow: Between 25 and 50 perce~t of
the area is covered with Poa spp., with ~
spp.
covering 5-25 percent of the total area. The vegetation is short, less than 1/2 m in height, and density
is mostly continuous.
Rock's cover less than 25 percent
of the total area and are primarily in the 6- to 12inch diameter size class. Green and black lichens predominate, each covering 5-25 percent of thp expospd
rock surface. Slopes are gentle and average about 7
percent.

C-3

Picea-Salix Krummholz:
Picea engelmannii cove~s 25-50
percent of the area, while Sa"lix spp. covers 5-25 percent of the total area. Appearance of this unit is
ragged, as some shrubs attain a height of 4 m. Most of
the vegetation is less than 2 m tall, and the der.sity
is mostly continuous. The amount of rock is variable,
with small areas covering 75 percent of the total area.
Rocks larger than 12 inches are dominant.
Lichens are
primarily green and black, with each covericg up to 5
percent of the exposed,rock surface. Slopes are variable but average 16-20 percent.

C-4

Salix-~
Alpine Meadow: More than 75 percent of the
area is covered with ~
spp., with species of Poa
amounting to about 5 percent of the total coverage.
Plant density in this unit is continuous, and average
height of the vegetation is 2-3 m. Few rocks are
present and slope averages about 7 percent.

(-5

Trifolium-Poa-Salix Alpine Meadow: Each of the three
dominant genera comprises 5-25 percent of the coverage
of the area. Most plants in this unit are less than
1/2 m tall; although the shrubs exceed this height,

�-102-

Table

2. Con t ,

UNIT

DESCRIPTION
they do not attain 1 m. Density of the vegetation is
continuous.
Rocks cover less than 25 percent of the
area and are primarily larger than 6 inches in diameteL
Green and black lichens dominate, and each covers up to
25 percent of the exposed rock surface. Slope gradient
averages about 10 percent.

C-6

Trifolium-Poa Alpine Meadow: Trifolium spp. and ~
spp. comprise 5-25 percent each of the total coverage
of the area. The dominant clover is I. dasyphyllum in
contrast to
Earryi in C-5. Veg~tation in this unit
is less than 1/2 m in height, and the density is conti~uous. Rock~ cover 5-25 percent of the total area
and are primarily in the 6- to 12-inch diameter ciass.
Lichens are predominately g resn and black arid cover up
to 50 percent of the surface of exposed rocks. Average
perc~nt slope is about 15.

I.

C-7

c-S

C-9

Sile!!e-Irifolium-~_Poa
Fellfield:
Rocks dominate
this arEa and account for about 75 percent of the total
coverage.
Rocks are primarily larger than 6 inches in
diameter.
Four plant genera comprise less than 5 percent each of the covarage. Plants are short and occur
singly or in small pa tches. Green and black Li che ns
each covers up to 25 percent of the exposed rock surface. Slopes are smooth and average about 30 percent.
Dryas-~-~
Rock Krurnrnholz: Dryas octopetala
comprises 25-50 percent 01 subunit C-Sa, with Picea
engelmanrii and ~
spp. each covering up to 5 percent of the total area. Appearance of this u~it is
irregular as some Picea bushes attain a height of 4 m.
Density is variable, but is mostly continuous or in
small patches. Rocks cover at least 5-25 percent of
the area with no one size class dominating.
Liche~3
are few but are mostly green and black. Slope gradient varies up to 60 percent but averages about 20
percent. Subunit C-Sb differs from C-Sa in haVing
about equal amounts of the three dominant genera.
~
e~gelmannii dominates the overstory of the
Kru~~holz, whilt Dryas octopetala dominates the
interspersed windswept areas.
Dryas Turf: Mats of Dry~s octopetala cover 5-25 percent of the tritalarea, with rocks covering an equal
area. Vegetation of this unit is less than 1/2 m in
height, although scattered Salix bushes attain a
height of 1 m. De~~ity of plants is mostly continuous)

�-103-

Table
UNIT

2.

Cont.
DESCRIPTION
although many plants occur in small patches. Rock size
is variable, but most rocks are in the 0-6 and larger
than 12 inches diameter size classes. Green and black
lichens each covers 5-25 percent of the exposed rock
surface. Slope~ average 10-12 percent. This unit is
similar to R-l ~n RMNP.

G-lO

Pinus-Carex Krummholz: Pinus flexilis and Carex spp.
each covers 5-25 percent of the area, with rocks
amounting to 25-50 percent of the total coverage. Some
shrubs attain a height of 3-4 m, but much of the vegetation is less than 1/2 m tall. Plarrtgrowth is primarily interrupted, although small patches do occur.
One large bOulder outcrop is included, and rocks are
primarily larger than 12 inches in diameter. ~ichens
are primarily greer:.and black and cover up to 50 percent of the exposed rock sur~ace. Slopes are variab~e
and average i5-20 percent.

C-ll

Carex Subalpine Meadow: Different species of Carex
cover 5-25 percent of this area. Other subdominscts
are Poa spp. and Sibbaldia procClmbers, Plant density
is mostly continuous~ and the majority of the plants
present are less than 1/2 m tall. Scattered shrubs
attain a height of 3-4 m. Rocks comprise less than 25
percent of the entire cover~ with all diameter cla3ses
being about equal in cover. Lichens are u.C'common.
The slope gradient averages 17-18 percent.
.

C-12

Dryas-Pinus Subalpine-Meadow:
Dryas octopetala cover;;
25-50 percent of this area, with Pinus flexilis covering up to 5 percent. Appearance of this urrit is ragged)
with most plants being shorter than 1/2 m. Density of
the vegetation is pri~cipally continuous, but areas of
interrup ted growth are present. Tota 1 rock coverage is
5-25 percent. with all diameter classes being equally
represented. Coverage of exposed rock surfaces is Less
than 25 percent~ with green and black being the domirrant colors. Slopes average 12 percent.

C-13

Sibbaldia Snow Accumulation Area: This area is dominated by rocks which cover 50-75 percent of the entire
area. No one rock diameter class predominates.
Sibbaldia procumbe~s is the dominant plant present,
covering 5-25 percent of the area. Vegetation in this
unit is short, le~s than 1/4 m in height, and the der:.sity is interrupted as plants occur singly or in small
groups. Lichen" are uncommon on rocks, but a gray
lichen covers u~ to 5 percent of the exposed soil. The
average slope gradient is about 15 percent.

�-104-

Table

2.

Cont.

UNIT

DESCRIPTION

C-l4

Salix-Poa-Betula Alpine Meadow: This area is similar
to C-4 except Salix spp. covers 5-25 perce~t of the
area and Betula glandulosa occurs only in this unit.
Most shrubs in this unit are less than 1 m tall. but
scattered conifers extend to a height of 2 m. Density
of the vegetation is mostly continuous. Rocks larger
than 12 inches in diameter cover 5-25 percent of the
area. Black and green lichens are common, with each
covering up to 5 percent of the exposed surface of
rocks. Slopes are gentle and average 5-7 percent.

C-15

Salix-Picea-Trifolium-Poa Krummholz: Salix spp. covers
25-50 percent of the area, with the other three dominant genera each amounting to 5··25 percent of the total
cover. Vegetation of this unit is irregular as some
groups of conifers reach a height of 3 m, but much of
the vegetation is shorter than 2 m. Plant density is
primarily continuous. Rocks of all size classes are
common but make up less than 25 perce~t of the coverage. About 50 percent of the exposed surface of sto~es
is covered with green and black lichens. Slope gradient averages 12 percent.

C-l6

Artemisia-Trifolium Snow Accumulation Area: Species of
Artemisia (pri~cipally A. sconulorum and A. norvegica),
and Trifolium parryi each covers 5-25 perce r.t of t.he
area, with Artemisia occurring singly. while !. parryi
is grouped in small patches. Plants in this uru t do
not exceed 1/2 m in height, and the density is frequently interrupted. Stones of all size classes are
common and cover at least 25 percent of the area.
Lichens are uncommon, with green and black bei~g the
most prevalent colors. Slope gradient averages 10-12
percent.

�-105-

CROWN

COLORADO

N

o

t

-

E±3

Fig. 3.

POINT AREA

Study Area Boundary
ICALI I" MIL!S
1/2
E'"""3
E?3

'=='

Territory locations, Crown Point Area, 1966-71.

I

�-106-

CROWN
N

o

f

FS3

Fig. 4.

r

POINT AREA

COLORADO
---- Study Area Boundary
SCALI!

F+3

IN MILI!S
1/2

E'"±3

e+3

Territory locations, Crown Point Area, 1972-74.

F+3

�-107-

. h t1/ of Salix, white-tailed ptarmigan breeding territories,
Table 3. H elg
Crown Point Area.

Territory

X

S.D.

Sem

Range

Sample
Size

1

11.05

1.98

.63

8.0 - 15.0

10

2

11.88

2.92

.65

8.5 - 21.0

10

3

11.72

3.03

.55

7.0 - 18.5

10

4

12.31

3.87

.61

7.0 - 26.0

10

5

12.29

3.87

.54

5.0 - 17.5

10

6

12.16

3.80

.49

4.0 - 17.0

10

7

12.24

3.73

.45

7.0 - 20.0

10

8

12.56

4.19

.47

7.5 - 24.0

10

9

12.93

4.33

.46

10.0 - 23.0

10

10

13.03

4.27

.43

8.5 - 20.0

10

1./ All measurements are in inches.

�-108-

LITERATURE CITED

Bendell, J. F., D. G. King, and D. H. Mossop.
of bluegrouse in a declining population.

1972. Removal and repopulation
J. Wildl. Manage 36(4):1153-1165.

Boag, D. A., and K. M. Sumanik. 1969. Characteristics of drumming sites
selected by ruffed grouse in Alberta. J. Wildl. Manage. 33(3):621-628.
Braun, C. E. 1969. Population dynamics, habitat, and movements of white-tailed
ptarmigan in Colorado. Ph.D. Dissertation. Colorado State Univ., Ft.
Collins. 189 pp.
1971a. Continued inventory of selected ptarmigan populations. Colo.
Div. Game, Fish and Parks, Game Res. Rept., Fed. Aid Proj. W-37-R. April.
pp. 91-106.
1971b. Habitat requirements of Colorado white-tailed ptarmigan.
Western Assoc. State Game and Fish Comms. 51:284-292.

Proc.

_____ , and G. E. Rogers. 1971. The white-tailed ptarmigan in Colorado.
Colo. Div. Game, Fish and Parks Tech. Publ. No. 27. 80 pp.
_____ , R. K. Schmidt, Jr., and G. E. Rogers. 1973. Census of Colorado whitetailed ptarmigan with tape-recorded calls. J. Wildl. Manage. 37(1):90-93.
Dorney, R. S., and C. Kabat. 1960. Relation of weather, parasitic disease
and hunting to Wisconsin ruffed grouse populations. Wisc. Conserv. Dept.
Tech. Bull. 20. 64 pp.
Fischer, C. A., and L. B. Keith. 1974. Population responses of central Alberta
ruffed grouse to hunting. J. Wildl. Manage. 38(4):585-600 .

..

Kuchler, A. W. 1955. A comprehensive method of mapping vegetation.
Assoc. Amer. Geogr. 45~404-4l5.

Ann.

Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male
sharp-tailed grouse. J. Wildl. Manage. 38(4):616-621.
Robel, R. J. 1969. Movements and flock stratification within a population
of blackcocks in Scotland. J. Anim. Ecol. 38(5):755-763.
Rusch, D. H., and L. B. Keith. 1971. Seasonal and annual trends in
numbers of Alberta ruffed grouse. J. Wildl. Manage. 35(4):803-822.
Schmidt, R. K., Jr. 1969. Behavior of white-tailed ptarmigan in Colorado.
M.S. thesis. Colorado State Univ., Ft. Collins. 174 pp.
Watson, A., and D. Jenkins. 1967. Experiments in population control by
territorial behavior in red grouse. J. Anim. Ecol. 36(4):595-614.

�-109-

LITERATURE CITED (continued)

Zwicke1, F. C. 1972. Removal and repopulation of blue grouse in an increasing population.
J. Wi1d1. Manage. 36(4):1141-1152.
, and J. F. Bende11. 1967. A snare for capturing blue grouse.
--Wildl.
Manage. 31(1) :202-204.

Prepared by __

~.:......:...:::~=' _ •

....:::r~--I,.£/~~

C1ait E. Braun
Wildlife Researcher

_

J.

��-111April, 1975
JOB FINAL REPORT

State 0 f

--=-CO.=..L::.O.=..RAD==-O"--_
W-37-R-28

Project No.

Game Bird Survey

Work Plan No.

17
Job No.
4
Characteristics and Migration~-o~-a-------------------Job Title: Wintering Population of Colorado White-tailed Ptarmigan
Period Covered:
Personnel:

April 1, 1972 through March 30, 1975

C. Braun, H. Funk, R. Hoffman, R. Oakleaf, and C. Wagner, Division
of Wildlife; D. Hein, P. Lehner, S. Martin, and R. Ryder, Colorado
State University; T. May, University of Colorado.

ABSTRACT
Studies of the characteristics and migration of wintering populations of
white-tailed ptarmigan (Lagopus leucurus) were conducted in north-central
Colorado from April 1972 through April 1974 primarily at Guanella Pass. In
all, 293 km2 of alpine habitat were selected for studying movements from and
to the wintering area.
Guanella Pass was utilized principally by females (80 percent of 799 birds
identified) and primarily adults (65 percent). Numbers of subadults varied
with year depending upon production success. The 172 winter flo~ks observed
ranged from 2 to over 80 birds per flock, but most contained 2 to 25. Females
typically congregated in larger flocks (20-30 birds) than males « 15 birds);
probably the result of differences in Winter habitat preference as partial
segregation of sexes occurred. Apparently, winter flocks were not composed of
family units nor were the same group of individuals associated together throughout the winter. Sixty percent of the ptarmigan banded at Guanella Pass and
relocated on breeding or summering sites returned in subsequent winters. Adults
exhibited the greatest affinity for the wintering grounds with 73 percent returning, while only 51 percent of the subadults returned. Willow (Salix spp.)
was the most important vegetation found on wintering areas. Essentially all
sites on the winter range were utilized, but some preferences were noted.
}fujor factors affecting distribution were availability of willow and snow quality.
Eight chick movements were documented to wintering or breeding sites. Hunter
recoveries indicated chicks were relatively sedentary prior to brood dispersal.
Females banded as chicks traveled greater distances than males. Hales showed
high fidelity to natal areas. Ninety-nine and 45 movements were documented
from and to winter use sites, respectively.
Subadults were more. mobile than
adults, with subadult females traversing the greatest distances. Female grouse
were consistently more mobile than males. Average movements of males were
generally restricted to less than 2.5 km for adults and 4.0 km for subadults,
while movements of females frequently exceeded these distances. The longest
movement was 22.7 km for a subadult female. Seventy-four percent of the movements from Guanella Pass were in westerly directions with 28 percent to the
northwest.

�-112-

RECOMMENDATIONS
1.

Any proposed project with possible adverse effects on willow dominated
communities in riparian, subalpine or alpine areas should be carefully
investigated.

2.

Continued research should emphasize investigating dispersal of broods
and identifying factors responsible for major losses of young ptarmigan
prior to arrival on wintering areas.

3.

Movement studies at Guanella Pass should be discontinued with the
exception of collecting information on hunter recoveries of banded birds.

4.

Information concerning wintering populations of ptarmigan in Colorado
should be made available in form of a special repprt.

�-113CHARACTERISTICS AND MIGRATION OF A WINTERING
POPULATION OF COLORADO WHITE-TAILED PTARMIGAN
Richard W. Hoffman and Clait E. Braun

P. S. OBJECTIVE
To document movements of white-tailed ptarmigan by age and sex class to
and from a major wintering area in order to evaluate importance of such
areas and distances involved in annual migrations.

RESULTS AND DISCUSSION
The objectives of this study have been completed and the results have been
previously reported (Hoffman 1974). An article was submitted to and accepted
by the Journal of Wildlife Management; the title will be "Migration of a
Wintering Population of White-tailed Ptarmigan in Colorado".
A typed manuscript
of the final report was prepared in thesis form and was accepted in partial
fulfillment of the requirements for the degree of Master of Science. A second
paper entitled "Characteristics of Wintering Populations of Colorado Whitetailed Ptarmigan'! is in final preparation and will be submitted to Condor for
publication.

PUBLICATIONS

RESULTING FROM STUDY

Hoffman. R. W. 1974. Characteristics and migration of wintering populations
of Colorado white-tailed ptarmigan.
M.S. Thesis. Colorado State Univ.
58pp.
___

• and C. E. Braun. 1973. Migrat.ion of white-tailed ptarmigan to and
from a major wintering area. Trans. Central Mountains and Plains Section
of the Wildl. Soc. 18:16.
(Abstract).
1974. Composition.
white-tailed ptarmigan.

distribution. and size of wintering flocks of
J. Colo. Wyo. Acad. Sci. 7(5}:68.
(Abstract).

1975. Migration of white-tailed
Wyo. Acad. Sci. (In Press).

ptarmigan

in Colorado.

1975. Migration of a wintering population of white-tailed
in Colorado.
J. Wildl. Manage. 39(3}. (In Press).

LITERATURE

J. Colo.

ptarmigan

CITED

Hoffman. R. W. 1974. Characteristics and migration of wintering populations
of Colorado white-tailed ptarmigan.
Colo. Div. Wildl., Game Res. Rept.
Fed. Aid Proj. W-37-R. April. p. 79-116.

Wildlife Researcher

Candidate

��-115-

April 1975

JOB PROGRESS REPORT

Stat e 0 f

C;::O::;L=.:O:.:RAD=:.:O=-------

Project No.

W-37-R-28

Work Plan No.
Job Title __

1
Job No.
-----------------Investigation of Population
~S~t=a=t=u=s~o::;f~B::;o::;b::;w~h~i_t_~e~Q~u~a~i=l~i=n~E~a~s~t~e~r~n~C~o~l~o~r~a~d~o~
_

Period Covered:
Personnel:

Game Bird Survey

20

April 1, 1974 to March 31, 1975

Donald Hoffman, Bruce Tryndle, Jim Rogers, Mike Telck, Marvin
Gardner, Dan Walsworth, Steve Steinert, and Warren Snyder.

ABSTRACT
Approximately 100 to 120 bobwhite quail on the Tamarack Wildlife Area survived the fall and winter of 1973-74 from an October, 1973 population level
of about 722 quail. Harvest mortality of approximately 31 percent combined
with natural fall-winter mortality of about 52 percent to reduce the fall,
1973 population by about 83 percent. Additional mortality among survivors
was low over spring and summer 1974 so that the annual mortality rate
approximated 87 percent. Summer whisting counts in 1974 were indicative
of the low resident breeding population.
Extremely dry weather, with total
1974 rainfall approximating 7 inches, did not hamper reproduction.
Hatching
reached a rapid and dramatic peak in mid-June.
By October over 400 young
had been added to the population bringing the fall level to approximat"ely
499 quail. Bands were placed on 197, or 39 percent of these. Hunters and
dogs were handicapped by extremely dry fall weather so that hunting removed
only about 18 percent of the fall population.
Relatively mild weather persisted through the winter resulting in good survival up to late March.
Ninety quail were banded or retrapped in February and early March, 1975.
A late March snowstorm, which killed numerous livestock, waterfowl and
pheasants in northeastern Colorado, may have also dramatically reduced the
bobwhite population along the Tamarack and South Platte River. Some coveys
survived, but the percentage is unknown at this time. Winter snow stress
continues to appear as the major limiting factor to northeastern Colorado
bobwhite.

��-117-

INVESTIGATION OF POPULATION
STATUS OF BOBWHITE QUAIL IN EASTERN COLORADO
Warren D. Snyder

P. S. OBJECTIVE
To investigate (1) the distribution and relative density of bobwhite quail
in eastern Colorado, and (2) population structure and level, and rate of
harvest, and identify key factors that limit bobwhite on the Tamarack Management Area.
SEGMENT OBJECTIVES
1.

To investigate
Colorado.

the distribution

and density of bobwhite

in eastern

2.

To investigate population structure and level, and rate of harvest of
bobwhite quail on the Tamarack Management Area.

3.

To identify key factors limiting bobwhite
Area.

on the Tamarack Management

METHODS AND MATERIALS
Methods and materials used in this investigation were listed by Snyder
1973, 1974) in previous segment progress reports.

(1972,

RESULTS AND DISCUSSION

Bobwhite Distribution

and Density in Eastern Colorado

1974 Census
Four Wildlife Conservation Officers, Clayton Wetherill, Bill Olmstead, Larry
Budde and Brett Petersen, conducted bobwhite whistling counts on established
routes along the South Platte River in 1974. These counts were supplemented
by others on all transects along the river. Results, shown in Table 1, indicate the breeding population was approximately one-fourth below the previous
three year average, excluding Tamarack data. The Stateline-Ovid route showed
some increase whereas tallies on the proximal Tamarack study area were only
half as high as found the previous year. Counts were not conducted in the
Arkansas, Cimarron and Republican river drainages in 1974.

�-118-

Table 1. A four year comparison of bobwhite quail whistling counts completed along the South Platte River.

Route

Stateline-Ovid

East Tamarack

West Tamarack

Sterling-Iliff

Atwood-Merino

Brush-Ft. Morgan

Goodrich-Orchard

Percent
Change From
3 Year Average

Year

Number of
Counts

Number of
Stops

1971
1972
1973
1974

2
2
4
3

20
20
40
33

78
65
93
115

3.90
3.25
2.33
3.48

+ 18.0%

1971
1972
1973
1974

2
4
6
5

20
45
64
54

67
202
135
65

3.35
4.49
2.10
1.20

- 61.7%

1971
1972
1973
1974

6
7
5
5

75
79
57
64

219
312
109
53

2.92
3.95
1.91
0.83

- 72.6%

1972
1973
1974

2
2
4

19
20
41

63
73
133

3.33
3.65
3.24

-

1971
1972
1973
1974

2
2
3
2

20
20
30
28

108
182
246
77

5.40
9.16
8.20
2.75

- 64.1%

1971
1972
1973
1974

2
2
3
2

20
20
30
20

77
163
60
72

3.85
8.15
2.00
3.60

- 16.1%

1971
1972
1973
1974

2
2
1
1

20
23
11
12

23
32
7
19

1.15
1.39
0.64
1.58

+ 37.4%

572
1,019
723
534

3.27
4.51
2.87
2.12

- 40.0% 1/

Natural
Tally Mean

7.2%

-------------------------------------------------------------------------------S. Platte River X

1971
1972
1973
1974

16
21
24
22

175
226
252
252

1/
- The percent change skewed disproportionatelyby extra counts on the Tamarack.
Excluding the Tamarack, the 1974 index was 23.6 percent below the three year
average along the South Platte River.

�-119Relation of the Number of Males to Total Calls Heard Per Station
Ellis and Thomas, 1972, stated that the average number of calls per stop
was more accurate than the average estimated number of calling males per
stop as an index of fall population. They stated that the number of
whistling males per stop became difficult to use when over seven calling
males were within hearing distance. Both calling males and total calls
per three minute time interval per station were tallied by the author
during the 1974 census. When males and calls were plotted on a scatter
diagram, a direct positive relationship was evident. However, the tally
exceeded seven males per stop only twice out of 141 samples where one or
more quail were tallied.
Tallies of ten or more males per stop were common during previous years
when popUlations were higher. Therefore, recording of both total males
and total calls per three minute time interval should be continued and evaluated for future use.

Trend Comparisons of Census and Harvest in Northeast Colorado
The bulk of the bobwhite quail harvest in northeastern Colorado occurs along
the South Platte River. Whistling counts conducted there from 1971 through
1973 were compared to harvest projections taken from Small Game Hunter F~rvest
Surveys. Data illustrated in Table 2 show some evidence of comparable trends
between the call index and total harvest, but there is no evidence of a relationship between the call indices and birds per hunter. Hunter harvest survey data
for 1974 were not available for use at the time this report was written.

Table 2. A comparison of the bobwhite whistling call index and s~bsequent harvest data obtained in northeastern Colorado. 1/

Item

1971

Year
1972

1973

Call Index l/

3.53

4.81

3.15

Total Harvest

12,512

12,658

8,967

Season Bag/Hunter

5.27

4.48

4.13

1/
- Harvest data from Management Units #1 and #2 which include all quail range in
northeastern Colorado.

2/

- Trend data from along the South Platte River. Data from only the east part of
the Tamarack were included to more uniformly spread sampling.

Trend Comparisons on the Tamarack Management Area
Figure 1 shows a close approximation of fall population level to the preceding
summer call index on the Tamarack. Harvest data also conform to this trend.
These data indicate that the whistling call index is a fairly reliable method
for predicting fall population level and harvest potential.

�-120-

600

3,000

6

.....
500

2,500

....

5

&gt;&lt;
Ul

0

400

2,000

~

4

...:l
{/)

~
~
::t:
::.::
~

~

~

8

Ul
...:l

1,500

\
~

Harvest

..

~
{/)
H

3
Whistling
Call Index

~

§

8

~
Ii.
200

\ ..~
\ .
\

~

0

0.

~

,

H

H

~
~
0.

~

,., ..
,, \

Z
'"

Z
0

300

~

U

~
,
~

..

1,000

100

500

o

o

Population

Ul

~
::t:
~
~
m

2

Leve1~\
\
\

.

\.-........... e. .
,.,

01...---__ "'1971

.

.-..;:

1

1972

..

"'-

..&amp;...__

1973

1974

Fig. 1. The relationship between the bobwhite whistling index and subsequent
fall population and harvest levels on the Tamarack Wildlife Area.

�-121-

The Tamarack data combined more intensive census (Table 1) with accurate
population and harvest information.
Management census previously discussed was not as intensive on several routes and Hunter-Harvest Survey
data were highly questionable.
A minimum of three counts per route should
be obtained in the future on all management routes and census-harvest
evaluation should be continued.

Population Structure, Level and Rate
of Harvest on the Tamarack Wildlife Area

Past Segment Band Recoveries,

Mortality

Rates, and Population

Estimates

Previous Segment Band Recoveries--High fall-winter mortality levels during
the 1972-73 and 1973-74 intervals left few banded quail surviving by the
spring of 1974. Recoveries from preceding trapping periods are summarized
in Table 3. These few recoveries do not provide enough information to warrant
revisions of fall and winter population estimates previously reported (Snyder
1974).

Table 3. Number of quail band recoveries and retrap information
banding periods obtained during 1974 and the winter of 1975.

from previous

Period Obtained
Period
Banded

Late
Winter 1974

Fall 1974

Fall, 1971

No Bands Recovered

Winter, 1972

No Bands Recovered

Harvest 1974

Late
Winter 1975

Fall, 1972

2

1

1 (off area)

0

Winter, 1973

1

0

0

0

Fall, 1973

3

1

0

4

1

2

2

Winter, 1974

Fall, 1973 to Fall, 1974 Mortality and Population Level Estimates--Our best
estimate of the 1973 pre-season population level is 722 quail, based on changein-ratio data presented in Snyder, 1974. The pre-season population level one
year later approximated 500 birds of which 18.18 percent or 91 were adults.
This indicates an annual mortality rate of 87.4 percent which was slightly
below the 93.7 percent level the preceding year. However, it was still a high
mortality rate and, in spite of excellent reproduction in 1974, a declining
population resulted.

�-122-

Our failure to find or to catch many quail during late winter trapping
in 1974 provides evidence that most of this mortality of approximately
631 quail occurred prior to the end of the winter. Only eleven quail
were banded and six were retrapped from previous work segments. After
tenacious searches using dogs under partial to excellent snow cover,
the resident population of quail on the Tamarack was estimated at
approximately 100 birds. In contrast, proportional recovery ratios of
fall and winter marked birds, based on inadequate samples, placed the
late winter, 1974 population at only 41 birds. Only six band recoveries
from fall, 1973 and six from winter, 1974 were obtained, illustrating
the small, inadequate sample size and high probability of error. Therefore, this population projection will be ignored.
A closer approximation to the actual late winter, 1974 population level
was attained by comparing banded-unbanded ratios of adults in the subsequent fall and winter (1974-75) trapped and harvested samples. These
data show that 6 adult quail, banded during late winter, 1974 were retrapped or harvested among 42 adults taken during the subsequent fall and
winter.
Projection would place the late winter, 1974 population level
at about 119 birds using this method.
Factors including immigration of
unbanded birds, emigration of banded birds and trap bias toward capture
of a higher proportion of unbanded quail might yield a population overestimate.
However, since about 91 adult quail were present in fall, 1974
the above quoted figure seems feasible and will be used as our best estimate of the late winter, 1974 population level. If there were 119 quail
alive in late winter, then the natural mortality rate up to that time
approximated 52 percent and when added to 31 percent harvest mortality,
brought total fall-winter mortality to 83.5 percent (Table 4).
Survival of adults through the dry spring and summer of 1974 was excellent.
Data in Table 4 indicate approximately 3.9 percent of the original-fall
population were lost during that interval.
Bobwhite sex ratios did not
change significantly from the fall-winte~ (1973-74) interval through the
subsequent fall-winter interval (Table 11). This provides some evidence
of excellent over-summer survival, since summer mortality usually takes
a much higher proportion of the hens than of the cocks.
Fall Banding, 1974
Warm, dry weather, an apparent adequate food supply, older than average
juvenile quail at the time of banding, and a relatively low population
level combined to reduce fall trapping success in 1974. However, it was
estimated that about 39 percent of the population was trapped prior to
the hunting season. On the west section of the Tamarack, 91 quail were
banded, and 103 quail were banded on the east section. Only three previously
banded adults were retrapped bringing the fall total to 197 quail (Table 5).
The number of quail banded per hunting area is illustrated in Table 6. As
shown, many of the West Tamarack hunting areas were apparently void of quail,
whereas quail were trapped in nearly everyone of the east areas.

�-123-

Table 4. Projected mortality of bobwhite quail from October,
through October, 1974 on the Tamarack Wildlife Area.

1973

West
Tamarack

East
Tamarack

Total
Area

Total Fall, 1973 Population
Estimate

440

282

722

Hunting Season Removal

155

71

226

Remainder Subject to Natural
Mortality

285

211

396

Late Winter, 1974 Population
Estimate 1./

60

59

119

Natural Fall-Winter Mortality

225

152

377

Total Fall-Winter

380

223

603

Percent Harvest Mortality

35.26

25.24

31.30 'l:.1

Percent Natural Mortality

51.14

53.90

52.22

Percent Combined Fall-Winter
Mortality

86.40

79.14

83.52

Spring-Sunnner Mortality

14

14

28

Percent Spring Sunnner Mortality

3.18

4.96

3.88

Fall, 1974 Adult Population
Estimate

46

45

91

Annual Mortality Rate

89.54

84.04

87.40

Item

Mortality

11

- Samples obtained during the fall and winter (1974-75) indicate that
approximately equal numbers of adult quail were present on the two sections
of the Tamarack.
'l:./percentagesvary slightly based on the method that data from disproportionate
populations were combined for calculation.
Percentages from the east and west
sections, when combined, yield a slightly different average than data calculated
from the total area figures in the last column.

Fall Movement
There was no evidence that banded quail moved out of the study area after fall
trapping in 1974. Therefore, corrections for movement were not applied to
fall population estimates (Table 5). Low quail densities and a shorter time
lapse between trapping and hunting may have reduced movement.

�-124-

Table 5. A summary of banding, harvest and population projections
the Tamarack based on fall, 1974 and winter 1975 data.

from

West
Tamarack

East
Tamarack

Total
Area

Number of Quail Banded

91

103

194

Number of Retrapped Adults

2

1

3

Total Marked

93

104

197

Fall, 1974 Banded

18

8

26

Previously

2

o

2

20

8

28

Crippling Loss (Projected Total)

14

6

20

Known Harvest

45

28

73

Projected

Item

Fall, 1974 Banding Information

Harvest Band Recoveries

Banded Adults

Total Band Recoveries
Harvest Statistics

Total Harvest

59

34

93

Percent Crippling Loss

31.11

21.43

27.40

Projected

26

10

36

44.44

28.57

38.71

2}.96

9.62

18.27

16

22

38

Other Quail Banded or Retrapped

29

23

52

Total Quail Handled

45

45

90

Percent Fall Banded

35.56

48.89

42.22

Total Band Recoveries

Percent Banded

(Fall and Previous)

Percent Harvest Mortality
(Includes Cripples)
Winter, 1975 Banding
Retrapped

Population

from Fall, 1974

Projection

for Fall, 1974 Using Combined Harvest and Winter Data

Fall Banded Harvest and Winter
Recoveries

34

30

64

Total Quail Harvested
Trapped

90

73

163

253

499

±70.92

±97.74

Fall Population
Confidence

or Winter

Estimate

246

Limits

±66.6l

+

�-125Table 6. A summary by hunting area of fall banded bobwhite, hunter use,
total harvest and banded harvest recoveries on the Tamarack Wildlife Area
in 1974.
Hunter
No. Quail
No. of
Total Quail Banded Quail
Area
Banded
Harvest '];/ Harvest 11
Hunters 1}
Hours 1/
West Tamarack
1

2
3
4
5

1

6
21

4

2

1

14

34
59
72
72
77
47
36
26
32
33
43

95
115
99
96
54
93
157
113
198
162
141
231
264
278
313
189
166
137
149
165
178

91

973

3,589

10

7
8

55

9

4
34

10
11

12
13
14
15 4/
18 19
20
21
22
23
24
25
Subtotal

112

84

13

6

40
30
31
22
38
43
20
30
33
56

44

12
4

5

1
1
2

1
2

1

4
4

3
2

1

1
45

18

East Tamara.ck
Braddock
Pasture
1
2
3

2 Ii..!

1
1

9

14

10
11

6
2

46
38
28
15
46
38
32
25
39
46
77

Subtotal

103

430

1,432

28

8

Total

194

1,403

5,021

73

26

9

5
25

4
5
6
7

11

5
24

8

133
147
74
103
132
123
115
111
94
156
244

1/ Includes only known hunters ruld hours reported

3
1
6
1

4

1

1

3
4
1
5
3

1

through the check station.

21 Includes only known harvest. Some hunting probably went unrecorded.
3/ Approximates the location where quail were shot. Exact area is unknown.

il Braddock Pasture and areas 16 and 17 West are closed to public hunting.

�-126-

Fall Population

Estimate

The 1974 fall population estimate of 499 quail again declined from that
of the preceding year (Table 7). Change-in-ratio calculations used in
the derivation were based on a 39 percent marking rate and a 33 percent
recovery rate (Table 5). Both harvest and winter trapping were combined
for the latter. Marked and recapture samples were not sufficient to prevent errors greater than 10 percent with 95 percent accuracy as recommended
by Robson and Regier (1964). However they did approach this level of
accuracy.
Confidence limits are provided in Table 5.

Table 7. Bobwhite fall population and harvest statistics
1974 on the Tamarack Wildlife Area.
Item

Fall Population
Season Length

Estimate

for 1971 through

1971

1972

1973

1974

1,745

2,319

722

499

(days)

20.5

31.5

32.5

29.5 11

Known Harvest

411

548

202

73

Projected Total Harvest
Mortality 11

472

634

227

93

27.0

26.7

31.7

18.3

Percent Harvest Mortality

1/

11
- Quail and pheasant hunting terminated at 2:00 P.M. from November
December 15, 1974 on the Tamarack.
'

29 through

21
- Includes projected

crippling loss.

The 1974 Hunting Season
The 1974 hunting season started at noon, November 16, ran for 30 days, and
terminated December 15 in the northeast part of Colorado.
Pheasant season
coincided with quail season. Season splits during previous years of study
extended the quail season up to or into early January.
However, management
personnel adopted recommendations for an earlier season termination based on
study findings discussed in a subsequent section of this report.
A special regulation applying only to the Tamarack Wildlife Area was also
implemented in 1974. It closed the pheasant and quail season at 2:00 P.M.
each day starting November 29 and continuing to season termination.
The
basis for this regulation will be covered later in this report.

�-127-

Hunting Pressure and Hunting Success--Average daily hunter use of the
Tamarack Area in 1974 declined by 4.34 percent from the 1973 average and
13.8 percent from the combined average of the three previous years (Table
8). Bobwhite quail harvest declined by about 64 percent from the 1973
harvest and approximately 80 percent from the previous three year average
(Table 9).
Table 8. Hunter success ratios on bobwhite
of hunting seasons from 1971 through 1974.

Interval

Number of
Hunters

y

Average
Hunter Use

quail with the progression

No. of Quail
Harvested

Hunters per
Quail Harvested

1971 Season
First 8 Days

638

79.75

258

2.47

Second 8 Days

363

45.38

100

3.63

Last 5 Days

258

51.60

53

4.87

Season Average

59.95

3.06

1972 Season
First 8 Days

606

75.75

230

2.63

Second 8 Days

386

48.25

167

2.31

Third 8 Days

369

46.12

100

3.69

Fourth 8 Days

392

49.00

36

10.89

Season Average

3.25

54.78
1973 Season
102.87

142

5.80

339

42.38

36

9.42

Third 8 Days

161

.20.12

10

16.10

Fourth 9 Days

303

33.67

13

23.30

First 8 Days

823

Second 8 Days

Season Average

8.09

49.27
1974 Season

First 8 Days

686

86.75

43

15.95

Second 8 Days

414

51. 75

17

24.35

Third 8 Days

138

17.25

9

15.33

Last 6 Days

176

29.33

4

44.00

Season Average

47.13

19.37

1/
- Includes all hunters using the Tamarack.
During 1972 and 1973, most
hunters late in the season were after ducks.

�-128-

As illustrated in the last column of Table 8, the ratio of hunters
checking into the Tamarack per each quail taken increased sharply in
1974. This was primarily due to fewer quail present but weather variables must also be considered.
Pheasant harvest was nearly equal to that of the previous year (Table 9),
but was well below harvest levels in 1971 and 1972. Cottontail and waterfowl harvest also declined dramatically from harvest in previous years
(Table 9). Dry, relatively mild weather was the factor believed responsible
for major declines in harvest of these two species.
It apparently increased
fox squirrel activity and subsequent harvest as noted in the table.

Table 9. A comparison of small game harvest on
during the 1971 through 1974 quail and pheasant the Tamarack Wildlife Area
seasons.
Year

Season Interval

Quail

Pheasants

Ducks

Rabbits

20.5 days

411

87

196

30

9

First 20.5 days

480

130

555

126

4

31.5 days

548

143

704

156

4

First 20.5 days

184

53

283

40

3

1973

32.5 days

202

62

441

63

3

1974

First 20.5 days

64

49

69

19

11

29.5 days

73

59

141

23

15

1971
1972
1972
1973

1974

Squirrels

The extreme dry conditions made it difficult for dogs to locate pheasants
and quail. As a result, when compared to previous years, the percent harvest
declined in greater proportion on quail than hunting pressure declined.
Quail harvest, including crippling loss, approximated only 18 percent of the
fall population level in 1974 compared to a 31 percent harvest rate in 1973
(Table 7).
The reported 1974 crippling loss was much higher than that reported in previous years.
It approximated 27 percent of the reported take (Table 5).
Warm dry conditions probably were a factor in making .it difficult for dogs
to locate downed birds.

�-129-

Hunter Use and Success by Hunting Area--A summation of hunter numbers,
hunter hours, quail banded in fall, banded quail, and total quail harvest
by hunting area is illustrated in Table 6. It shows that most of the
harvest and band recoveries occurred in the same or proximal areas to
where the quail were banded earlier in the fall. Most of the coveys were
only lightly harvested, however, 7 of 13 quail originally banded on Area
4-West were harvested.
This is the only location where hunters removed
a fairly high proportion of the bobwhite present.
Information presented in Table 6 indicates that the west section of the
Tamarack received proportionately greater hunting pressure than the east
section. Each of the 23 west areas received approximately 156 hours of
hunter use compared to 130 hours per each east area. In contrast, 103
quail were banded on the 11 east areas, compared to only 90 on the 23 areas
to the west. This indicates that the section containing the 'highest quail
density received the least hunting pressure.
This variable probably accounted
for part of the low harvest rate.

Winter Banding, 1975
High river levels, ice jams, minor flooding and low population levels made
winter trapping difficult, however 90 quail were taken. Bands were placed
on 46, 38 were retraps from the previous fall, and the remaining 6 were retraps from other trapping periods (Table 5). Equal numbers of trapped birds
were handled on both the east and west sections of the Tamarack.
Trapping,
initiated in early February, was terminated in mid-March.
It revealed that
fall-winter survival to mid-March was proportionately much better than that
occurring during the two previous years.

Age Ratios - Fall to Winter, 1974-75
Table 10 summarizes age ratio information obtained during the work segment.
It illustrates that the relatively low breeding population was able to reproduce quite successfully.
Nine young for each pair of adults were estimated
to have been present in the fall population from the ratio of adults to young
in the harvest (1 Ad:4.5 Y). Only in the fall of 1972 was there a lower indicated percentage of adults in the fall population.
That year there were
16.3 percent adults compared to 18.18 percent adults in 1974. However the
low percentage of adults in 1972 followed a summer when 35 percent of the
adults were lost between late winter and fall. In 1974, adult mortality over
the same interval approximated 4 percent.
Therefore, reproduction success
per adult may have been higher in 1974 than in 1972 assuming most 1972 adult
mortality followed reproduction instead of preceding it. There were 19.2
percent adults in the 1971 fall population and in 1973 the fall population
contained approximately 20.3 percent adults. There is little evidence that
significant differentially higher mortality of young occurred during any of
the fall periods to affect age ratio percentages.

�-130-

Table 10. Bobwhite age ratio data obtained
during the fall and winter
of 1974-75 on the Tamarack Wildlife Area.
Number
Adults

Number
Young

Total

Percent
Adults

14

183

197

7.11

Banded

3

24

27

11.11

Unbanded

9

30

39

23.08

Total

12

54

66

Winter Banded

19

71

90

Sample

Fall Banded

Adult : Young
Ratio

1

13.07

18.18

1

4.50

21.11

1

3.74

Hunting Season

Sex Ratios
Harvest and late winter, 1973-74 samples showed roughly 54 percent of the
population to be males and 46 percent to be females (Table 11). By the
subsequent fall among surviving adults, about 57 percent were males and 43
percent were females.
Although fall trapped juveniles occasionally can be inaccurately sexed, most
of those obtained during the fall of 1974 were mature enough to en~ble
.
accurate sex determination.
A high proportion of males were found then and
again in late winter trapped samples (Table 11). The reason for this skewed
sex ratio is unknown. One fall banded covey was entirely composed of males.
This high proportion of males may increase whistling call indices disproportionately in relation to reproduction in 1975, since young approximate 80 percent of the 1975 breeding population, and about 57 percent of surviving adults
were males.

Table 11. Sex ratios of bobwhite quail from fall-winter,
winter, 1974-75 intervals.
Item

Male

1973-74 to fall-

Female

Total
214

Harvest and Late W'inter, 1973-74
Percent

116

98

54.21

45.79

Surviving Adults, Fall-Winter,
Percent

25

14

44

Fall, 1974 Banded Juveniles
Harvest, 1974 Juveniles
Winter, 1975 Juveniles

56.82
112
26
44

43.18
62
28
27

174
54
71

Combined Total of Juveniles
Percent of Total Juveniles

182
60.87

117
39.13

1974-75

299

�-131-

Factors Limiting Bobwhite on the Tamarack

Fall and Winter Mortality
Contrasts in fall-winter weather and quail survival among the years of
study are summarized in Table 12. Quail survival was excellent in 197172 during the first year of study which included an essentially snow-free
winter. No heavy snows (four inches or more) were recorded that fall and
winter.
Snows starting in November, accompanied by severe cold in December, were
evident during the fall and winter of 1972-73.
There were 76 days with
snow cover (Table 12) and 37 days with accumulations of four inches or more.
This severe weather coincided with hunting seasons to a large degree, creating
a multiplication of stress. As a consequence, natural plus hunter induced
mortality increased from about 16 percent in 1971-72 to approximately 60 percent in 1972-73.

Table 12. A comparison of fall-winter natural mortality,
and subsequent call index change from the previous year.

Item

1971-72

snow conditions,

Fall~WinterPeridd
1972-73

1973-74

11

52

50

Days With Snow Cover

14

76

54

Days With 4" + Snow Cover

0

37

30

16.3

60.0

52.2

+37.5

-51.3

-50.5

Total Snowfall

Fall-Winter

(Inches)

Mortality Rate 1/

Percent Change in Subsequent
Call Index From Previous Year

1/
- Does not include harvest mortality.

Similar high fall-winter mortality occurred in 1973-74. The major severe
weather stress period did not begin until December 18, after the major hunting
stress had occurred.
But snow depths to 10 inches or more persisted in
December, 1973 and January, 1974 for almost a month blocking off food supplies
during a period of severe cold. Approximately 119 quail remained on the
Tamarack in scattered small coveys in contrast to a late winter population of
nearly 1,000 two years earlier. Most survivors of the 1973-74 winter used
the tall Phragmites grass as survival headquarters in accompaniment with
pheasants, cottontails and deer.

�-132-

The fall-winter period of 1974-75 was essentially snow free up to late
March when a severe storm with cold rain, heavy snow, and high winds hit.
Winter covey searches and trapping, completed prior to the storm, revealed
survival of an estimated 200 to 250 quail. Post storm searches did not
reveal evidence of quail mortality but such mortality is difficult to detect. Numerous waterfowl, pheasants and small passerines were found dead
in the storm devastated area where large numbers of cattle also were killed.
Pheasants were not found dead in the riverbottom, but a high proportion
were lost in adjacent tablelands.
At least some of the quail coveys survived, and summer call indices will provide further evidence of the storm's
impact on the quail population.

Hunting As An Additive Stress--There is evidence that the relatively high
hunting pressure, to which Tamarack quail were subjected during periods of
snow stress, heightened mortality there. This is shown by comparison of
whistling call indices up and down river with indices obtained on the
Tamarack (Table 13). High mortality occurred on the Tamarack during fallwinter periods of 1972-73 and 1973-74 reSUlting in subsequent respective
call index declines of 51.3 and 50.5 percent from preceding years. Combined data from upstream and downstream census routes indicate call indices declined 34.4 percent after the first severe winter and 17.3 percent
after the second severe winter.
Quail hunting pressure on these locations
were light in comparison to that on the Tamarack.
The latter received
approximately 50 hunters per day during quail season.

Table 13. A comparison of bobwhite whistling counts on the Tamarack Wildlife Area with counts located up and down river.
Routes UE and Down River !!
No.
Count!
No.
Percent
Stops
Stop
Change
Stops

X

Year

1972

79

5.99

1973

120

3.93

1974

122

3.25

Routes on Tamarack

X Count!
Stop

Percent
Change

124

4.15

-34.4

121

2.02

-51.3

-17.3

118

1.00

-50.5

lIIncludes average of Brush-Ft. Morgan, Atwood-Merino,
Ovid-Stateline routes.

Sterling-Iliff

and

Review of U. S. Weather Bureau records indicates that if hunting seasons can
be held earlier in the fall there is less chance of overlaying hunting stress
with the impacts of snow and accompanying severe cold weather.
Data from
1955-56 through 1973-74 (19 years) showed that November averaged 4.3 days
with snow on the ground, December averaged 10.1 days, January averaged 13.2

�-133-

days and February averaged 8.8 days in the vicinity of the Tamarack.
Average minimum temperatures for November, December, January and February
were 24.8 degrees, 17.2 degrees, 13.8 degrees and 18.5 degrees, respectively, based on 22 years of data. These data illustrate that hunting
seasons should be started as early in November as possible and that they
probably should not be extended beyond mid-December to obtain maximum
hunting opportunity with minimum quail population depletion due to weather
related stress factors.
Current policy is for the quail and pheasant season to open on the third
Saturday in November so as to not conflict with harvest of farm crops.
Preferably, this opening should be moved forward. But if this opening
date is to continue, the quail season should run continuous to termination.
Division management personnel submitted recommendations for a continuous
season in 1974.
Division management personnel felt that special regulations were needed
to attempt to increase survival and retain a higher breeding population
on the Tamarack since the population there was so low. Rather than reduce season length, recommendations that called for an early (2:00 P.M.)
closure were implemented starting on November 29 and running through the
remainder of the season. This regulation applied to both pheasants and
quail. However, due to low populations, hunting pressure was low and
the regulation did not restrict much hunting opportunity.
This early
P.M. closure each day permitted hunted coveys to regroup and feed. Since
there are only 8-9 hours of daylight in December in contrast to 15 to 16
hours of darkness, adequate evening feeding opportunity and time to regroup are critical to survival. This is especially true during periods
of severe cold and snow.

Spring and Summer Mortality and Reproduction
Available information does not indicat~ that any significant adult mortality occurred during the spring and summer of 1974. Apparently a majority
of the quail were able to nest successfully and hatching success was good.
There were no severe thunderstorms, hail or flooding conditions during 1974.
The low adult population probably survived better than a higher popUlation
would have since the number surviving was far below carrying capacity and
was therefore subject to less environmental and competitive stress.

Projected Period and Peak of Hatch
A sample of 177 young bobwhite aged in early fall provided evidence that
nearly all young were hatched in June, 1974 (Fig. 2). As this figure
illustrates, hatching peaked three weeks earlier than in 1972 and over a
month earlier than in 1973 when floods delayed nesting.
It is probable
that the dry spring and summer weather (Table 14) prompted early reproduction
in 1974. In 1974, 3.43 young were alive in fall per adult alive prior to
the start of nesting season. This compared to 2.03 young per adult in 1973
and 1.96 young per adult in 1972. Apparently dry conditions which existed
during 1974 (Table 4) were not detrimental to reproduction or survival of
young to fall was better than in 1972.

�-134-

34
32
30
Sample Size
28
N •• 768 in 1972
26

N = 288 in 1973
N •• 177

24

in 1974

22
20
'0
GI

-6 18

~fO
:c 16

....

r-t

fO

~,. '.• '\91~••.
,',
0\,

+'
0 14

=: '. i,

••
IH
0

I

J&gt; ••••
_

.'

12

I

I

'. I,

I

+'
c

"

,"

'.

I

,

I'\ ,

'\

II

GI

u 10

I

I
I

~

"" 8

.
.. .

18

,,

,,

'

o "

,
-....
\ II

·t~.

..~

'•......•. .

,--- -~,
•
,,•

4

25

May

,

.'.

I

6

2

.

"

•

!

,•

,
,•

••.•.
",
•

/

... ,

"

I \

.~ "I

I~""

2

\

,

". \ ,

.. ,,

,

/

--

\

'.

.

\

/

"

_/
9

17
June

24

3

11

19
July

27

4

12

20 28
August

7 14
Sept.

Weekly Mid-points

Fig. 2. Period and peak of bobwhite quail hatch projected from primary
feather growth for 1972, 1973 and 1974 on a percent of total basis.

�-135Table 14. Precipitation
1971 through 1974. II

received in the vicinity of the Tamarack

from

Year
Month

1971

1972

1973

1974

January

0.46

0.25

0.68

0.25

February

1.20

0.09

0.12

0.67

March

1.98

0.08

3.09

0.91

April

2.71

1.95

2.50

0.45

May

3.71

2.20

3.05

0.70

June

2.54

3.13

2.26

2.05

July

3.40

2.88

2.54

1.15

August

1.28

3.57

0.23

0.40

September

2.10

0.64

4.75

0.15

October

1.42

0.64

1.10

Tr.

November

0.36

2.40

0.54

0.25

December

0.12

0.34

0.89

0.20

Total Annual

21.28

18.17

21.75

7.18

1/

- Information up to April, 1974 taken from U. S. Weather Bureau Station
located 5 miles south of Sedgwick, Colorado.

Winter Site Use Preferences
The values of tall Phragmites cover for winter survival during periods of
snow stress was previously mentioned.
However, during mild winters without
deep, prolonged snow cover, bobwhite survive well elsewhere along the Tamarack.
Figure 3 shows winter coveys located during the winters of 1971-72 through
1974-75. Although some locations, which contain both heavy cover and an
adjacent or intermingled food supply, were used consistently, other coveys
have been found throughout most of the riverbottom.
There were a few exceptions. The Braddock Pasture on the west end of the East Tamarack was grazed
lightly year-long and therefore lacked winter cover needed to sustain quail.
It provided an excellent late-summer and early fall feeding area, but quail
moved elsewhere to spend the winter. The far east portions of the Tamarack
(Fig. 3) on the north side of the river generally contained abundant cover.
But these locations lacked food supplies needed to hold quail through the
winter and repeated trapping efforts and searches with dogs failed to locate
quail there during the past four winters.

�TAMA RACK

MANAGEMENT

AREA

~~S1 AREAS

I
•....•
W
0\
I

tv

/
EAST

AREAS

o

SCALE
Yo

Mi.

LEGEND
1

HUNTING

~UNFARMED
~
AND

AREA

RIVER

RIVER BOTTOM
CHANNELS

aD = COVEY LOCATION AND NUMBER OF

~
ALFALFA
L........J

WINTERS
f.]GRASSLAND
c:=:JHAY

Fig. 3. Locations of bobwhite
from 1971-72 through 1974-75.

~
MEADOW

FOUND THERE

ROW CROPS
PLANTING

coveys found on the Tamarack

Wildlife

Area during winter

trapping

periods

�-137-

LITERATURE

CITED

Ellis, J. A., K. P. Thomas, and P,'Moore.
1972. Bobwhite whistling
activity and population density on two public hunting areas in
Illinois. National Bobwhite Quail Symposium.
1:282-289.
Robson, D. S., and H. A. Regier. 1964. Sample sizes in Peterson markrecapture experiments.
Trans. Amer. Fish Society. 93(3):215-226.
Snyder, W. D. 1972. Population status of bobwhite quail in eastern
Colorado.
Colo. Div. of Game, Fish and Parks. Game Res. Rept.
April. pp. 121-150.
1973. Population status of bobwhite quail in eastern Colorado.
Colo. Div. of Wild1., Game Res. Rept. April. pp. 253-296.
1974. Population status of bobwhite quail in eastern Colorado.
Colo. Div. of Wildl., Game Res. Rept. April. pp. 119-147.

Prepared by

2V~/2.

~

Warren D. Snyder
Wildlife Researcher

��April 1975

-139-

JOB PROGRESS REPORT

S tate of __

--=C:.::O.::L:.::O.::RAD=...::O
_

Project No.__ W_-_3_7_-_R__28
Work Plan No.

Game Bird Survey

2_1
~~~Job No.
l
Minimum Tillage Techniques for
Establishing Shrubs in Clump Plantings

Job Title
Period Covered:
Personnel:

_

_

April 1, 1974 through March 31, 1975

Donald M. Hoffman and Warren D. Snyder

ABSTRACT
Shrub plots were established on six sites in northeastern Colorado during the
spring of 1974. Two plots were respectively placed in each of three different
site situations represented by: (1) upland shortgrass vegetated loam soils,
(2) sandhill with sandy loam soil vegetated by mid- and short grasses, (3)
riverbottom containing clay loam soil, high water tables and high alkaline
condi tions. Each site contained seven plots to be treated by different mulch,
herbicide and cultivation methods for evaluation of weed competition, shrub
survival and growth. Hansen rose (Rosa sp.) and wild plum (Prunus americana)
were used in equal proportions in each of the forty-two plots. All chemical
pLo t s were treated with a combination of Eptam and Treflan during this first
year. In subsequent years additional chemical treatments will be 'used and
evaluated. Slightly over seven inches of precipitation were received during
1974, which is less than half the average annual amount. As a consequence,
shrub growth and survival was poor. Lowest shrub survival was observed in
the plots treated with organic mulch. The Eptam-Tref1an herbicide did not
reduce shrub survival. Mulch treatments were highly effective in control of
annual forbs. Noxious perennials present on the riverbottom sites were not
controlled effectively by any of the 1974 treatments. One additional site in
each of the three soil types will be established in 1975 and evaluations will
continue.

��-141-

MINIMUM TILLAGE TECHNIQUES FOR
ESTABLISHING SHRUBS IN CLUMP PLANTINGS
Warren D. Snyder

P. S. OBJECTIVE
To evaluate ground cover techniques, pre-emergent herbicide techniques,
combinations of the two, and cultivation techniques for establishing shrubs
in clump plantings in eastern Colorado.

SEGMENT OBJECTIVES
To prepare six planting sites and establish clump plantings of Hansen rose
and wild plum in three soil types. To apply pre-emergent herbicide, mulch,
mulch-herbicide, and cultivation treatments to clump plantings within the
test sites. To tally survival of shrubs within respective clumps, measure
shrub growth, and measure weed competition.

METHODS AND MATERIALS
Two sites within each of three soil types were selected on Division owned
Duck Creek and Tamarack Wildlife Areas in northeastern Logan County for initial
development. Three additional sites have been added for development in 1975.
The approximate locations of these is illustrated in Fig. 1.
Four of the six initial sites were rototilled with a tractor mounted rototiller
to a 4 to 6 inch depth in October 1973. All sites were worked again in March
1974 toobreak up sod and to prepare the sites for planting. Seven square plots
each 27 ft per side (729 square ft) were ali~led adjacent to each other in a
linear group within each site. Therefore, outside dimensions of each site
were 27 ft by 189 ft.
To facilitate application of chemicals to four plots during the first year of
study, these plots were placed in consecutive order by random selection at one
end of the plot series. Individual treatments to be applied in subsequent
years were selected by use of a random numbers table among the chemical group
and among the remaining three non-chemical plots. Placement sequences of
plots within sites, within soil types are illustrated in Table 1. The bottom
of this table also illustrates the plot sequences for sites to be planted in
1975.
A mixture of two pre+eme rgent; herbicides was applied to the four chemical plots
in late March 1974, or approximately one week before the shrubs were planted.
An application equivalent to two pounds of Eptam and one-half pound of Treflan
(active ingredients) per acre was applied in liquid form with a hand sprayer.
T.he sprayer contained three spray nozzles to permit rapid even application over
the entire area. Calibrations of water and coverage were made prior to treatment to obtain correct application and distribution rates. The chemicals were
rototilled into the top soil by shallow tillage immediately after application.

�-142-

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TION.

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~------~-------~------~------~------~\~=====db=
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Fig. 1. Location of shrub clump sites on Division of Wildlife properties
in northeast Logan County, Colorado.

�-143-

Table 1. Reference sequence of test shrub clump plots within sites for 1974
plantings and 1975 plantings.

Plot
Number

Sandhills
Red Lion
Sharptail

River Bottom
Tamarack
Smith Meadow

Tableland
Karg 111
Karg 112

1974
1

orgam.c-. 1/

Plastic

Control

Plastic

Ep - Tref

Sim - M

2

Plastic

Organic

Plastic

Organic

Simazine

Simazine

3

Control

Control

Organic

Control

Sim - M

Casoron

4

Ep - Tref

Simazine

Casoron

Casoron

Casoron

Ep - Tref

5

Simazine

Ep - Tref

Simazine

Ep - Tref

Organic

Plastic

6

Sim - M

Casoron

Sim - M

Sim - M

Plastic

Control

7

Casoron

Sim - M

Ep - Tref

Simazine

Control

Organic

Sequence
Direction E - W

NE - SW

NE - SW

E - W

N - S

NE - SW

Plot
Number

I - 76

Sonnenberg

Karg 113

1975
1

Organic

Sim - M

Plastic

2

Plastic

Ep - Tref

Organic

3

Control

Simazine

Control

4

Simazine

Casoron

Casoron

5

Sim - M

Control

Ep - Tref

6

Ep - Tref

Plastic

Simazine

7

Casoron

Organic

Sim - M

E - W

NE - SW

NE - SW

Sequence
Direction

1/ Organic and plastic are mulch treatments, Ep - Tref represents EptamTreflan herbicides and Sim - M represents Simazine overlaid with organic mulch.

�-144-

Fig. 2. Photo taken on Karg ill site while standing among wild plums
recently planted on the cultivated control. Hansen rose to the left
are less conspicuous.

Fig. 3. A thicket of wild plum. is one of the mos t highly utilized
covers that can be provided for pheasants, quail, rabbits and other
wildlife.

�-145-

A second application of granular Eptam 10-G was applied at the rate of one
pound per plot by broadcast and shallow tillage in late July, 1974. Only
two of the chemical plots within each of the six sites were treated and
tilled.
As illustrated in Table I, additional chemical treatments will be applied in
late winter or early spring approximately one year after the shrubs were
planted. One randomly selected plot within each site will receive a repeat
application of Eptam and Treflan at previously listed rates. One plot will
be treated with the pre-emergent herbicide, Casoron, applied at the rate of
approximately 150 pounds per acre. Two plots will receive treatments with
the chemical Simazine applied at the rate of 2 pounds (active ingredient)
per acre. One of these plots will be overlaid with a thin layer of organic
mulch after the chemical is applied. Chemical application rates were obtained
from the Colorado Weed Control Manual authored by E. Heikes and J. Fults and
updated each year.
Two shrub species, Hansen rose and wild plum were used in equal numbers on
the plots. Four rows of each species were spaced at three ft intervals
starting three ft from the plot boundaries (Fig. 2). A rope marked at three
ft intervals was placed acrost the site in linear manner to facilitate
planting one row in each plot before going to the next row. Bias among plots
due to variation in the quality of different seedling bundles was reduced by
this method. Bare rooted shrubs were obtained from the Colorado State Forest
Service Nursery located at Fort Collins.
Both shrub species were hand planted, however, two different planting techniques were used. Holes were dug using a narrow spade and post hole digger for
placement of the heavily rooted plums. The narrow spade was inserted into
the ground and worked fore and aft to open a narrow vee into which the poorly
root.edHansen rose were inserted. The ground was firmed around all seedlings.
No water was applied. Two men were able to plant 2,688 seedlings in less
than five days during the first week in April, 1974.
An overlay of wood chips or corn cobs were placed on the organic mulch plot
within each site in ~ate April or early May 1974. Each site received two
pickup loads which covered the plot to a depth of four to six inches.
Plots to be overlayed with black plastic (4 millimeter thickness) were ridged
before planting using a garden rototiller with furrower attachment and a hand
rake. Seedlings were placed in the furrows and 30 inch wide strips of plastic
were placed over the ridges to direct precipitation toward the shrub rows.
An overlay of wood chips or corn cobs was placed over the plot to hold the
plastic in place. Applications were completed by early May 1974.
Post-planting height measurements were obtained in May and repeat height
measurements were obtained in late fall 1974. A yardstick was used to obtain
the measurements which were subsequently averaged per plot and species.
The initial survival tally of shrubs was delayed to mid-July due to uncertain
survival status of many of the plants. A second tally of survival was
completed in mid-September 1974.

�-146-

Forms were prepared to record treatments, man-hours of effort and estimated
costs per treatment along with weed competition impact. All plots were
checked in mid-June, mid-July, mid-August and early September for weed
growth. Weeds were tallied and removed if present during the June, July
and September inspections on the loam and sandy loam sites. Perennial
weeds and grasses, not destroyed in preplanting tillage, were necessarily
removed more frequently in the two bottomland clay-loam sites.
An automatic recording rain and snow gauge was set up in mid-April 1974 at
the Tamarack headquarters, near centrally located among the sites. This
recorder was serviced monthly. Precipitation received prior to recorder
activation in 1974 was obtained from a U. S. Weather Bureau station located
five miles south of Sedgwick, Colorado.
All shurbs were treated with a rodent and deer repellent which was applied
in mid-fall. Application was made with a hand sprayer.
All shrubs in the loam and sandy loam sites were watered in mid-March 1975
because of the prolonged drought conditions that persisted to that time.
Each shrub received approximately two gallons of water. River bottom sites
were not watered because of high ground water levels at the locations.

Study Area
This evaluation study was established on Division of Wildlife properties
along the South Platte River in northeastern Logan County. It was designed
to compare treatments under three soil types. Fine sandy loams, which are
abundant in eastern Colorado, were represented by test sites located in the
sandhills along Interstate Highway 76 in the south part of the Tamarack
Wildlife Area (Fig. 1). Loam soils sub tended by clay or clay loam were
found on the Karg addition of the Duck Creek property in shortgrass sodded
pastureland approximately two miles north of the town of Crook. The third
series of sites lay along the valley of the South Platte (Fig. 1) and contain
clay loam soils with fairly high alkaline salts due to high water tables.
Inland saltgrass (Distichlis stricta) is the most ~ypical vegetation along
much of the valley area. Soil characteristics are described in Table 2.
Elevations in the study region approximate 3,700 to 3,800 ft. Precipitation
in the region averages about 17 inches, although considerable annual and
seasonal variation is common.

RESULTS AND DISCUSSION
Shrub Establishment and Survival
Drought Impact
Near record drought conditions, unforeseen during study implementation,
created a major stumbling block toward shrub establishment and study progress.
Average annual precipitation in the study region approximates 16 to 17 inches.

�-147However, only a little over 7 inches were received in the vicinity from
January 1 through December 31, 1974 (Table 3). Soil conditions were near
adequate at planting time, although pre-planting tillage dried the soil to
a degree. Regrettably, little precipitation was received in April and May
following planting when it was critically needed to moisten and firm the
soil around the shrub roots. As a result, many of the Hansen rose seedling
did not show evidence of life until after the June 8 rain which deposited
approximately two inches of precipitation. This was both the first and last
substantial precipitation received during the year. As shown in Table 4
about 93 percent of the plums and 80 percent of the roses were at least
partially alive in mid-summer. By mid-September survival had dropped to
about 82 to 87 percent for the plums, and 73 to 75 percent for the roses on
all plots. Due to lack of any significant moisture during the last half of
1974 and early 1975 over winter survival of many of the remaining shrubs
was uncertain at the end of the work segment. Many of the Hansen rose shrubs
in the sandy loam (Red Lion and Sharptail) sites were killed over winter by
burrowing rodents which consumed the roots of this species. Rodents only
occasionally ate the roots of the wild plum shrubs in these sites. The
result of these various mortality factors is relatively low survival during
the first year, especially on Hansen rose.

Table 2.

Soil analyses completed on the six shrub clump sites.l/

Analysis

Sandhi11s
Red Lion Sharp tail

Tableland
Karg 1
Karg 2

Riverbottom
Tamarack Smith Meadow

pH level

6.2

6.0

7.6

6.9

7.7

7.6

Soluble salts
(mmhos/ em)

1.7

1.6

1.1

1.5

3.4

3.3

Organic matter
(percent)

2.3

1.7

3.2

5.4

6.3

7.6

Percent sand

76

82

48

46

22

22

Percent silt

16

11

30

35

41

39

Percent clay

8

7

22

19

37

39

Sandy
loam

Sandy
loam

Loam

Loam

Clay
loam

Clay
loam

Texture

l/ Information from Soil Testing Laboratory, Colorado State University.

�-148-

Table 3.
1975. 1/

Precipitation

received near the study sites in 1974 and early

Month

Precipitation
1974

January

0.25

February

0.67

March

0.91

April

0.45

May

0.70

June

2.05

July

1.15

August

0.40

September

0.15

October

0.00

November

0.25

December

0.20

1974 Total

7.18
1975

January

0.05

February

0.15

March

1.25

1975 - 3 month total

Grand Total

1.45

8.63

Recorded

�Table 4.

Shrub survival on the shrub clump plots in mid-July

and late September 1974.

Cultivated
Control

Organic
Mulch

Plastic
Mulch

1

2

Chemical
3

4

x

Rose

78.6

72.4

77.1

85.9

90.1

79.2

80.7

80.6

Plum

95.3

88.0

91.7

95.8

93.7

95.3

95.8

93.7

73.4

57.8

72 .9

81.8

82.8

80.7

79.7

75.6

77.6

86.4

88.5

85.4

91. 7

85.9

86.9

Time

Mid-July

Late September1=!
Rose
Plum

l/ An additional
September.

92.7

1.8 percent of the roses and 4.5 percent of the plums were uncertain

as to survival

I
•....•

~

1.0
I

in

�-150-

As illustrated in Table 4, wild plums showed better survival than rose
shrubs during the 1974 growing season. This probably was due to better root
structure on the larger plum seedlings which subsequently were planted to a
greater depth and therfore had better access to available soil moisture.
Many of the rose seedlings did not have well developed root structure.
Treatment Impact on Survival
Hansen Rose - As illustrated in Table 4, Hansen rose survival was lower than
that of wild plum on the study plots. The four chemical plots, which all
received the same Eptam-Treflan treatment, showed similar survival rates
approximating SO percent. Cultivated control and plastic mulch survival
approximated 73 percent and organic mulch plot survival dropped to 57.S
percent for roses. This level was significantly lower than that of the
combined control and chemical plot survival average of 79.7 percent (Chi
square = 6.01, 1 d.f.).
Wild Plum - Organic mulch plots again showed the lowest average survival
among the four treatments, although the difference was not significant (Chi
square = 1.41, l.d.f.) between organic and combined control and chemical
plot comparisons.
Cultivated control survival of plum was the highest
recorded (Table 4).
Treatment Effects - It is probable that the organic mulch, which was applied
shortly after the plots were planted in April, withheld moisture vitally
needed by the shrubs during the subsequent dry periods of spring and summer.
Mulch applications probably should have been delayed until substantial
moisture had been received and the shrubs were rooted and established.
The plastic mulch, overlaid with organic mulch, did not show evidence of a
significant effect in reducing shrub survival. The plastic dire~ted at
least some of the moisture toward the shrubs since it was placed over ridges
between rows, however, at the same time the organic mulch overlay absorbed
and held some moisture. The organic overlay was considered essential to
reduce soil temperatures that would otherwise build under the black plastic
and it was also needed to hold the plastic in place.
Survival of both species in the organic and plastic plots within the Red Lion
sandy loam site was very low by the end of the year. Therefore, these two
plots will be completely replanted during the spring of 1975.
There was no evidence that applications of Eptam and Treflan had any significant impact on shrub survival (Table 4). Rose survival was higher in the
chemical plots than in the controls and plum survival averaged only slightly
lower in the chemical plots.
Shrub Growth
There was no appreciable shrub growth during the 1974 growing season for
either shrub species. Wild plum height averaged approximately 20.85 inches
in May and 21. 32 inches in September. Hansen rose height was about 20.5Sin.
in May and lS.50 inches among September survivors of the prolonged drought.

�-151Weed Competition
Noxious Weed Problems
Study sites were selected and cultivated during the fall and/or early spring
prior to planting in April 1974. Preferrably, the sites should have been
selected and summer fallowed during the growing season prior to planting.
This became evident when field bindweed (Convolvulus arvense) and white top
(Cardaria draba), both deep rooted noxious perennials showed up en masse in
the Tamarack Meadow test site. In addition, saltgrass (Distichlis stricta)
whorled milkweed (Asclepias subverticillata), Canada thistle (Cirsium arvense)
and sedge (Carex sp.) among others occupied the Smith meadow site on the Duck
Creek property. These perennials were not killed during pre-planting tillage
and subsequently persisted as weed problems on the bottom land clay loam
sites throughout most of the 1974 growing season. Repeated efforts to control
them using cultivation and 2-4-D Amine contact herbicide were only partially
successful. High ground water levels in the river valley provided moisture
prompting successive regrowths of these deep rooted perennials. As a consequence, these two sites have been excluded in treatment evaluations presented
in Table 5 and subsequent paragraphs of this section.
Weed Competition in Sandy Loam and Loam Sites
The sandy loam plots were vegetated with short and mid- grasses and sandsage
(Artemisia filifolia) pri.or to pre-plant cultivation. Loam soils on the
Karg sites were densely sodded with buffalo grass (Buchloe dactyloides) and
blue grama (Bouteloua gracilis). Most of these perennials were killed by
cultivation prior to shrub planting. Some of the deep rooted sandsage
attempted recovery and occasionally cactus plants (Opuntia sp.) remained,
although neither presented significant problems. In the sandy loams western
ragweeds (Ambrosia psilostachya) were most troublesome among the perennials.
In the loam shortgrass sites, skeleton weed (Lygodesmia juncea), and wild
alfalfa (Psoralea tenuiflora) were the deep-rooted perennials most commonly
present. Neither significantly competed'with the shrubs, but shallow tillage
and chemical pre-emergents were not effective controls.
Two factors suppressed weed growth on these upland sites. The dry weather
reduced weed germination throughout the growing season. In addition, annual
weed growth was reduced because the sites were newly broken out of, and
surrounded by grassland. Milk purslane (Euphorbia supina) was most abundant
among the annuals although the individual plants remained relatively small
and not highly competitive. Croton or doveweed (Croton texensis) and purslane
(Portulaca oleracea) were common in the sandy loam sites. Pigweeds (Amaranthus
spp.), Russian thistle (Salsoli kali) and lambsquarter (Chenopodium spp.) were
few in number but obtained large competitive status rapidly in the plots.
Effects of Treatments
As illustrated in Table 5 the mulch plots on the sandy loam and loam soil
types were essentially weed free throughout the 1974 growing season. Both
mulch treatments were highly effective in suppressing weed growth, but as
previously noted, the organic mulch withheld moisture reducing shrub survival
during this unusually dry year. It should be noted that the bindweed on the
Tamarack meadow site was not significantly retarded by mulch treatments.
Saltgrass on the Smith meadow was retarded but not totally suppressed by the
mulch overlays.

�-152-

Eptam-Treflan pre-emergent herbicides suppressed annual weed growth as is
shown in both the mid-June and mid-July figures in Table 5. The average
number of forbs were tallied before they were removed on each plot. The
average among the four plots and four sites was compared with the number
of forbs found on the four cultivated controls. Chi square values were
highly significant during both periods (Chi square = 654, 1 d.f. for June,
Chi square = 113, 1 d.f. for July). The chemicals significantly reduced
but did not totally eliminate forbs on any of the plots. Some species
such as milk purslane were apparently little affected by this chemical
combination.

Table 5. Weed competition on four shrub clump sites during the 1974 growing
season.

Shrub
Group

Cult.
Control

Organic
Mulch

Number of Weeds Removed
Plastic
Eptam-Treflan
Mulch
No. 1
No. 2

Mid-June Weed Competition
Red Lion
Sharptail
Karg #1
Karg #2
Sub-total

Treated Plots
No. 3
No. 4

Tally

955
27
149
256

o
o
o
o

o
o
o
o

132
44
158
104

237
10
168
86

200
30
68
64

225
72
65
76

1,387

o

o

438

501

362

438

175
360
100
24

318
384
40
46

479
373
34
48

659

788

934

85
78
18

100
113
15

o

o

Mid-July Weed Competition Tally
Red Lion
Sharptail
Karg #1
Karg #2
Sub-total

78
662
83
292

o

O.

6
10

10

106
416
77
58

1,115

16

10

657

o

o
o

Early September Weed Competition
Red Lion
Sharptail
Karg 111
Karg #2
Sub-total

Totals

o
o

Tally

o

o

o
o
o
o

o
o

0
2
0
0

13

10

o

3

2

181

228

2,515

26

10

1,098

1,162

1,331

1,600

13
0
0
0

10

3

�-153-

The impact of a late July application of Eptam in granular form to two of
the four plots in each site could not be evaluated.
Persisting hot dry
weather prevented weed germination on any of the plots in late summer.
Further tests and evaluations are needed under different weather conditions
prior to drawing conclusions on the use of mulch and chemical treatments.
Time did not permit careful site selection and summer fallow to avoid and/or
eradicate deep-rooted noxious perennial weeds prior to shrub planting.
However, neither mulches nor selective herbicides can be expected to control
bindweed and other noxious perennials.
Sites containing noxious perennials
should be avoided if use of these techniques is considered.
Impacts of
Simazine, Simazine-mulch and Casoron are yet to be measured.

Cost Analysis
Preliminary cost and man-hours figures have been worked up and placed on file
for all treatments.
These will be supplemented during successive years of
treatment for use in future evaluations.

LITERATURE

CITED

Heikes, E., and J. Fults. 1974 (updated annually).
Colorado Weed Control
Manual.
Colorado State University, Extension Service Publication.
Mimeo.

Prepared by·

.7d.rMW-:IL¥
Warren D. Snyder
Wildlife

Researcher

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                  <text>July, 1975

-1-

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~--------W-lOl-R-17

Project No.
Work Plan No.
Job Title

4

J~

Inventory of Range Manipulation

Period Covered:
Personnel:

Game Range Investigations
~.

la

-------------------------------

Projects in Colorado

April 1, 1974 through March 31, 1975

Roland C. Kufe1d and Regional Game Biologists.

ABSTRACT
An inventory was made of all range vegetation-modification
projects completed
during 1973 in the western half of Colorado, on lands administered by the
U. S. Forest Service and Bureau of Land Management.
Acreages treated were:
Forest Service - 2,410 and Bureau of Land Management - O.

��-3-

INVENTORY OF RANGE MANIPULATION

PROJECTS IN COLORADO

Roland C. Kufeld

P. S. OBJECTIVE
To collect information which describes proposed and completed range vegetation modification projects that are located in Colorado, and their effects
on the range, wildlife and livestock; and to provide desired IBM listings of
these data to cooperating agencies upon their request.

SEGMENT OBJECTIVES
1.

To collect information which describes proposed and completed range
vegetation modification projects that are located in Colorado, and
their effects on the range, wildlife and livestock.

2.

To compile, codify, process and analyze inventory data.

3.

To provide desired IBM listings of inventory data to cooperating
agencies upon their request.

METHODS AND MATERIALS
An inventory was made of all range vegetation modification projects completed
through 1972, in Colorado, west of Interstate Highway 25 on lands administered
by the U. S. Forest Service and Bureau of Land Management (Kufeld 1968, 1970,
1971, 1973 and 1974). Kufeld (1968 and 1970) also covered projects completed
on the Southern Ute and Ute Mountain Indian Reservations through 1969.
Indian Reservation projects were dropped from the inventory after 1969. This
report concerns vegetation modification projects completed on U. S. Forest
Service and U. S. Bureau of Land Management lands west of Interstate Highway
25 during 1973.
Data were collected using procedures outlined in Colorado Division of Wildlife Administrative Directive No. 204 entitled "Range Vegetation Modification
Projects", and by Kufeld (1968, 1970 and 1974). Inventory data were transferred from original data sheets to Mark Page Reader Forms, and then to IBM
cards.
Procedures outlined in Administrative Directive No. 204, call for evaluations
to be made on each vegetation modification project at the end of the 2nd, 5th,
and 10th years following treatment. During 1973,2 year post-treatment evaluations were made on projects completed in 1971, and 5 year post-treatment
evaluations were made on projects completed in 1968. Data were recorded on
a form described by Kufe1d (1971).

�-4-

RESULTS AND DISCUSSION
Acreages treated during 1973, on all lands administered by the U. S.
Forest Service west of Interstate Highway 25, are shown by vegetation
type and kind of treatment in Table 1. These acreages are shown by individual National Forests in Table 2. No vegetation modification projects
were completed on Bureau of Land Management lands in Colorado during 1973.

Table 1. Acreages of range land treated during 1973 in Colorado by the
U. S. Forest Service.
Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded As
Part of Treatment

Grass

Seed Only
Contour

2
1

225
600

225
600

3

825

825

6
1

965
600

965
0

7

1565

965

1

20

20

1

20

20

10

2410

1810

Browse }j

Conifer

Chain :?:../
Burn

Seed Only

All Vegetative Types
GRAND TOTAL

1/
- All vegetation modification projects conducted during 1973 in the "browse"
vegetation type were in the "oakbrush" subtype.

:?:../
Chaining includes chaining, cabling, railing and bulldozing.

�-5-

Table 2. Acreages of range land treated during 1973, within each National
Forest in Colorado.
National
Forest

Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded As
Part of Treatment

Rio Grange

Grass

Seed Only

1

25

25

1

25

25

1

600

600

1

600

600

1

200

200

1

200

200

3
1

665
600

665
0

4

1265

665

1

20

20

1

20

20

San Isabel

San Juan

Seed Only

Grass

Seed Only

Grass

Browse

];/

Conifer

White River

Total All
Forests

Chain Y
Burn

Seed Only

All Veg.
Types

All Treatments

6

1485

885

Browse ];/

Chain ];/

3

300

300

3

300

300

11

2410

1810

Total All
Veg. Types

Total All
Treatments

1/
All vegetation modification projects conducted during 1973 in the "browse"
vegetation type were in the "oakbrush" subtype.

~I
Chaining includes chaining, cabling, railing and bulldozing.

�-6-

A total of 2,410 acres were treated on National Forest lands in 1973.
To date (through Dec. 31, 1973) a total of 584,482 acres have been treated
in Colorado west of 1-25 on lands administered by the U. S. Forest Service
and Bureau of Land Management.
Of this 244,975 acres were on Forest Service
and 339,507 acres were on Bureau of Land Management lands.
Since 1973 was only the fourth year for 2 year post-treatment evaluations
of completed vegetation modification projects, and since no 5 year evaluations have yet been made, data from post-treatment evaluations of 1968,
1969, 1970 and 1971 projects will be kept on file and covered in a future
report when enough 2 and 5 year post-treatment evaluations have been made
to provide a basis for a meaningful data analysis.

LITERATURE CITED
Kufeld, Roland C. 1968. Inventory of range manipulation projects in
Colorado.
Colo. Div. Game, Fish and Parks. Game Res. Rept. P.R. Proj.
W-lOl-R-lO, WP 4, Jla, July, Part 1. p. 1-121.
1970. Inventory of range manipulation projects in Colorado.
Colo. Div. Game, Fish and Parks. Game Res. Rept. P. R. Proj. W-lOl-R12, WP 4, Jla, July, Part 1. p. 59-94.
1971. Inventory of range manipulation projects in Colorado.
Colo. Div. Game, Fish and Parks. Game Res. Rept. P.R. Proj. W-I0l-R-13,
WP 4, Jla, July, Part 1. p. 1-15.
1973. Inventory of range manipulation projects in Colorado.
Colo. Div. Wildl., Game Res. Rept. P.R. Proj. W-lOI-R-15, WP 4, Jla,
July, Part 1. p. 1-8.
1974. Inventory of range manipulation projects in Colorado.
Colo. Div. Wildl., Game Res. Rept. P. R. Proj. W-I0l-R-15, WP 4, Jla,
July, Part 1. p. 1-11.

Prepared by ~L-~~~~~~~~~~~~~~_
Roland C. Kufeld
Wildlife Researcher

�-7-

July 1975

JOB PROGRESS REPORT

State of

COLORADO
------~~~~~--------

Project No.

W_-_l_0_l-_R_-_l_7_

Game Range Investigations
2

Work Plan No.
Job Title

~4----~--~~=
Job No.
Experimental Improvement-~o~f~O~ak~b~r-u~s~h--------------------on Deer and Elk Winter Ranges-Beaver Creek

Period Covered:

April 1, 1974 through March 31, 1975

Personnel:

Roland C. Kufeld, John Klein, Judd Brown, and Larry Stewart.

ABSTRACT
Effects of spraying of Gambel oak vegetation on forage production and elk and
deer use were evaluated 2 and 5 years after treatment. The area~ located 28
miles southwest of Hayden, Colorado, was aerially sprayed with 2,4,5-TP in
1969.
With adjustments for normal vegetation production changes measured on a control
area, production of grasses increased 43 percent after 2 years, and shrubs and
forbs decreased 29 and 15 percent, respectively. After 5 years grass and
shrub production were 17 and 7 percent above pre-treatment levels and forbs
were 4 percent below. Total vegetation on the sprayed area de~reased 4 percent after 2 years, while a 5 percent increase was recorded after 5 years.
Data were grouped according to elk, deer and cattle forage. Definitions of
elk, deer and cattle forage are presented in the test. Spraying did not substantially increase elk, deer, or cattle forage. Increases in preferred
grasses after 2 years were offset by decreases in preferred shrubs and forbs.
After 5 years, the decline in preferred grasses was offset by increases in
preferred shrubs and forbs. Spraying did, however, change the forage composition, creating a variety of forage choices from that found in surrounding
unsprayed oakbrush types. After 2 years production of available, seasonal elk
forage ranged from 70 percent of pre-treatment levels during winter to 109
percent pre-treatment levels during fall. After 5 years elk forage production
in relation to pre-treatment levels ranged from 99 percent during fall to 108
percent during winter and summer.
Two years after treatment available deer forage had decreased for all seasons.
Seasonal deer forage production compared to pre-treatment levels ranged from
70 percent in spring to 90 percent during summer. After 5 years shrub regrowth elevated seasonal deer forage production to between 102 percent of pretreatment levels in spring and 107 percent during winter. Cattle forage
production after 2 and 5 years was 109 and 104 percent pre-treatment levels,
respectively. Comparison of control and spray areas indicate elk and deer use
between September 1 and mid-May increased 73 and 16 percent respectively, 2
years following spraying. Five year elk and deer use evaluations have not
been completed as of the date of this report. Cattle use was not measured
because the area was closed to grazing.

��-9-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH
ON DEER AND ELK WINTER RANGES-BEAVER CREEK
Roland C. Kufeld

p. S. OBJECTIVE
To determine if deer and elk forage production and game use can be increased
on overage Gambel oakbrush winter game ranges by spraying with 2,4,5-TP to
induce sprouting.

SEGMENT OBJECTIVE
To determine the extent of vegetation composition and production changes, and
the extent of elk and deer use changes which have resulted from spraying with
2,4,5-TP in 1969.

METHODS AND MATERIALS
Study area design, sampling scheme and procedures used in spraying on July 10,
1969 have been described by Kufeld (1970).

Vegetation Measurements

2 and 5 Years After Spraying

Two and 5 year post-treatment vegetation measurements were made on permanent
point-quadrat plots established in 1968. Procedures for measuring vegetation
were the same as those used in 1968, when vegetation was measured before treatment (Kufeld 1970). Two year post-tr~atment measurements began August 6, 1971
and were completed August 26. Measurements 5 years after treatment began
August 12, 1974 and were completed August 26.
Pre- and post-treatment vegetation production data were computerized and
subjected to covariance analysis by the Colorado State University Statistics
Department. Statistically significant changes in vegetation production was
tested at the .05, ..
10, .15, .20, and .25 confidence levels. Tests for
significance at lower levels were made because the study involved only one
spray and one control area. Inclusion of more replications would have
increased the study area size, workload and manpower requirements to the point
the study would not have been feasible.
Vegetation conditions in the spray and control areas were rephotographed
between August 30, and September 1, 1971; and August 21 through 23, 1974, at
67 permanent photo points established in 1968.
Elk and Deer Use Measurements 2 and 5 Years After Spraying
One hundred fifteen, .001 acre, circular pellet plots were cleared of elk and
deer pellet groups on September 1, 1971, and accumulated pellet groups were

�-10-

counted on May 8, 1972 to measure changes in deer and elk use 2 years after
spraying. These plots were recleared on August 25, 1974, and accumulated
pellets will be counted in May, 1975 to measure changes 5 years after treatment.

RESULTS AND DISCUSSION
Vegetation Changes 2 and 5 Years After Spraying
Vegetation production measurements discussed throughout this report refer to
vegetation below a level of 5~ feet. All vegetation change data have been
adjusted for normal year to year fluctuations in plant growth as measured on
the control area. This was done by computing the level of vegetation production one would expect to find on a treated area following treatment had treatment not occurred. This expected value, hereafter referred to as~, is
predicted from the regression of pre- to post-treatment vegetation data on
the control, coupled with pre-treatment data from the sprayed area. The dif"ference between the expected post-treatment production (E) on the sprayed
area and the actual production recorded there is presumed to be the change
resulting from spraying. Production as referred to in this report refers to
changes in frequency of occurrence of plants.
Total vegetation production changed very little following spraying. After 2
years total vegetation on the sprayed area decreased 3.94 percent, while an
increase of 5.34 percent was recorded after 5 years (Table 1).
Relatively large changes in production of forbs, grasses and shrubs occurred
after spraying. Grasses increased greatly, while shrub production decline d
markedly during the first summer after spraying. By the second summ~r when
the first post-treatment vegetation measurements were made grass production
had declined somewhat and shrub regrowth was underway (Fig. 1). Despite the
decline of grasses and regrowth of shrubs between the first and second years
after spraying, production of grasses had increased 43.92 percent and shrubs
had decresed 28.51 percent 2 years after spraying. By the end of 2 years,
forbs showed a 14.72 percent decrease due to spraying (Table 1).
Between 2 and 5 years after spraying grass production continued to decline
toward pre-treatment levels and shrub and forb production continued to rise
(Fig. 1, Table 1). Grass production after 5 years was still higher than
adjusted control area values, but only by 17.05 percent. This difference was
significant (P&lt;.05) but most of the difference was attributable to the large
increase in grass yields measured 2 years after treatment.
Shrub production after 5 years was 6.86 percent higher than before spraying,
and forbs were 3.93 percent below pre-treatment levels. The post-treatment
change in shrubs was significant at the a.05 level primarily due to the
large decrease in shrubs observed after 2 years. The forb decline was significant at the a.15 level, again due primarily to the decrease in forbs after
2 years.
The principal grass and grasslike plants on the study area are Bromus anomalus,
Elymu~El~~_,
Poa pratensis and Carex ~~~.
Poa sp. and Carex geyeri

�-11-

were lumped together in this study because it was often difficult to distinguish between them when making vegetation measurements.
Bromus anomalus
and Elymus glaucus accounted for most of the large increase in grass production after 2 years. These species increased 80.91 and 190.98 percent,
respectively. Production of Bromus anomalus continued to rise and after 5
years it had increased 195.62 percent. Five years after treatment production
of Elymus glaucus was only 28.33 percent above pre-treatment levels. Po aCarex geyeri declined by 1.02 percent after 2 years and 15.55 percent after
5 years (Table 1).
Production of primary shrubs declined substantially after 2 years. Amelanchier
alnifolia, Prunus virginiana, Quercus gambellii and Symphoricarpos Spa declined
67.15, 49.34, 12.68 and 18.64 percent, respectively.
The decreases in these
shrubs were probably even greater the first year after spraying than after the
second (Fig. 1). As a result of sprouting and regrowth, production of
Amelanchier alnifolia and Prunus virginiana after 5 years was only 12.34 and
20.52 percent below pre-spray levels, and Quercus gambellii and Symphoricarpos
Spa were 34.26 and 11.99 percent above production levels recorded before
spraying (Table 1).
Two of the primary forbs on the study area Agastache urticifolia and Mertensia
franciscana showed continued declines throughout the 5 year post-spraying
period. These were down 15.25 and 32.64 percent, respectively after 2 years
and down 31.30 and 46.84 percent after 5 years. Three others, Erigeron sp.,
Lathyrus leucanthus, and Lupinus argenteus showed decreases after 2 years
(-26.86, -32.16, and -41.18 percent, respectively) but by 5 years after
spraying Erigeron Spa and Lathyrus leucanthus were more abundant than before
(+11.74 and +29.19 percent, respectively), while Lupinus argenteus was still
below pre-treatment levels (-23.89), but still more abundant than it was after
2 years. Galium boreale and Thalictrum fendleri production increased 2 years
after treatment (+16.16 and +59.31 percent, respectively) and these continued
to become more abundant up to 5 years after spraying when their production was
+39.12 and +100.42 percent above pre-treatment levels, respectively. Achillea
lanulosa was 5.99 percent more abundant after 2 years, but 14.02 percent less
abundant after 5 years (Table 1).
Changes in Elk, Deer and Cattle
Forage 2 and 5 Years After Spraying
Definition of "Elk, Deer and Cattle Forage"
Classification of study area plants as elk or deer forage for pusposes of
determining production changes was based on the average, relative consumption
of those plant species by elk and deer in 48 elk and 99 mule deer food habits
studies reported in the literature (Kufeld 1973b; Kufeld, Wallmo and Feddema
1973). Plants considered cattle forage are those classified as "desirable"
and "intermediate" by the U. S. Forest Service in their Region 2 Range
Analysis Handbook for the foothill shrub, aspen weed, and mountain meadow
vegetation types.
Since animal preference for individual plant species varies from location to
location, the validity of applying "average" consumption data to this study

�-12-

area may be open to question. However, these averages do represent a great
deal of food habits research, and constitutes the best information available
to date on the foods these animals have been known to consume and the relative
amounts consumed.
For elk the following classifications are used: highly Valuable, valuable, and
least valuable (Kufeld, 1973b). For deer plants they are classified as heavily
eaten, moderately eaten, or lightly eaten (Kufeld, Wallmo and Feddema 1973).
Although the terminology used in the elk and deer references is different, the
basis for classifying plants according to relative consumption is exactly the
same. "Highly valuable" plants for elk, and "heavily eaten" plants for deer
by definition, generally comprised a major part of a food sample (usually at
least 20 percent) in food habits studies where reported. In a few cases,
plants which comprised less than a major portion of the food sample were
classified as "highly valuable" or "heavily eaten" if their reported contribution to the diet was far in excess of their reported vegetative composition.
"Valuable" plants for elk and "moderately eaten" plants for deer usually comprised between 5 and 20 percent of the food sample. "Least valuable" plants
for elk and "lightly eaten" plants for deer comprised less than 5 but more
than 1 pe rcen t.
Definition of "Available Forage"
Data and discussions relating to available forage are based on the assumption
that due to factors such as degree of plant maturity and snow cover, plant
availability to elk and deer on the study area will consist only of shrubs
during Win t er , grasses and shrubs during sp rtng and fall, and forbs, grasses
and shrubs during Summer. Discussions of cattle forage relate only to Summer
at which time all forage is available. These categorizations are somewhat
arbitrary and probably under-represent actual forage availability.
Effects of Spraying on Available Elk Forage.--Pre- to post-treatment ratios
using 1.00 as a basis to represent the level of elk forage production before
spraying with adjustment for normal year to year changes measured on control
areas are shown in Table 1. A single value for elk forage was derived by
weighting the number of hits on "highly valuable" forage by a factor of "3",
valuable by "2", and least valuable forage by "1". A total.weighted value
was then computed (Table 2).
Two years after treatment production of available elk forage was lower on the
sprayed area during winter and slightly higher during spring, summer and fall
than existed before spraying. The level of available forage compared to 1.00
after 2 years was 0.70 for winter, 1.07 for spring, 1.04 for summer and 1.09
for fall. This spray project was much less effective in stimulating elk
forage production after 2 years than was the spraying portion of another oak
control study conducted by Colorado Division of Wildlife near Hightower
Mountain, 20 miles east of Collbran, Colorado (Kufeld 1975). In that study
the following pre- to post-treatment ratios were observed 2 years after
spraying: winter, 1:1.01; spring, 1:2.09; summer, 1:1.72; and all, 1:1.94.
Grass production after 2 years on the Hightower Mountain study increased 295

�-13-

percent compared with the 44 percent increase observed on Beaver Creek.
During the first summer after spraying the kill on shrubs, particularly oakbrush, appeared about the same on both projects, however, after 2 years the
kill on the Hightower study area was obviously better than on Beaver Creek.
Both studies involved the use of 2,4,5-TP at a rate of 2 lbs/acre.
Herbicide
was mixed with 4 gallons of water and one-half gallon of diesel fuel on the
Hightower study while on Beaver Creek herbicide was mixed with 10 gallons of
water per acre. Both areas were sprayed between July 10 and 12, but in
different years. Equipment difficulties which resulted in completion of
spraying later in the day on Beaver Creek than on Hightower MOuntain may have
contributed to reduced effectiveness of the Beaver Creek treatment.
Except for winter forage very little change occurred in production of elk
forage on Beaver Creek between 2 and 5 years after spraying.
After 5 years
winter elk forage had increased to 1.08 times pre-treatment levels.
Pre- to
post-treatment ratios for spring, summer and fall were 1:0.99, 1:1.08, and
1:1.02, respectively (Table 2).
The 44 percent increase in grasses after 2 years did not substantially
increase the overall production of elk forage because this was offset by the
reduction of several shrubs preferred by elk. The reversal of this situation
occurred after 5 years when the increase in preferred shrubs was offset by
the decline in several preferred grass species.
Thus, during the 5 year posttreatment period spraying had little influence on the overall production of
elk forage. It did, however, change the forage composition, creating a greater
variety of elk forages compared to surrounding unsprayed oakbrush type areas.

Effects of Spraying on Available Deer Forage.--Pre-to
post-treatment ratios
for deer forage derived as described under the section on elk forage are
shown in Table 2. After 2 years, the sprayed area supported less available
deer forage during all seasons of the year than existed before spraying due
to the initial shrub kill. The level of available forage compared to 1.00 was:
winter 1:0.70, spring 1:0.83, summer 1:0.90, and fall 1:0.84. Available deer
forage increased as shrub resprouting and regrowth occurred between 2 and
5 years after spraying.
However, after 5 years available deer forage was
only slightly more abundant than before treatment.
Pre- to post-treatment
adjusted forage production ratios after 5 years were: winter 1:1.07, spring
1:1.02, summer 1:1.06, fall 1:1.05.

Effects of Spraying on Cattle Forage.--Spraying had little effect on production
of cattle forage. Pre- to post-treatment adjusted forage production ratios
computed after 2 and 5 years were 1:1.09 and 1:1.04. These compare to a ratio
of 1:2.17 found on the Hightower Mountain oak study area (Kufeld 1975) where
spraying was much more effective after 2 years in stimulating production of
cattle forage. Increased production of grasses on Beaver Creek, particularly
after 2 years, was offset by initial reductions in several shrubs and forbs
considered to be preferred cattle forage. Production of least desirable
plants decreased 15.31 percent after 2 years, but had increased 5.74 percent
after 5 years. The change in production of least desirables after 5 years
was significant at the a .05 level (Table 2).

�-14-

Elk and Deer Use Changes 2 and 5 Years After Spraying
Responses of elk and deer to the sprayed area 2 years after treatment were
described by Kufeld (1973a). Comparison of control and spray areas indicated
elk use from September 1 through mid-May increased 73 percent and deer use
during that period increased 16 percent following spraying.
Five year post-treatment elk and deer use data are not currently available as
accumulated pellets in plots cleared in August 1974 will not be counted until
May 1975.
Although vegetation data were categorized according to cattle forage preference it was not possible to measure the response of cattle to spraying in this
study as the area is closed to grazing.

LITERATURE

CITED

Kufeld, Roland C. 1970. Experimental improvement of oakbrush on deer and
elk winter ranges - Beaver Creek.
Colo. Div. Wildl., Game Res. Rept.,
Proj. W-lOl-R-12, WP 4, Job 2. July 1970. Pt. I. pp. 95-112.
1973a. Experimental improvement of oakbrush on deer and elk winter
ranges - Beaver Creek. Colo. Div. Wildl., Game Res. Rept., Proj. W-lOlR-15, WP 4, Job 2. July 1973. Pt. I. pp. 9-14.
1973b. Foods eaten by the Rocky Mountain
26 (2) :106-113.

elk.

J. Range Manage.

1975. Experimental improvement of oakbrush on deer, elk, and cattle
ranges - Hightower Mountain.
Colo. Div. Wildl., Game Res. Rept., Proj.
W-lOl-R-17.
WP 4, Job 3. July 1975.
(In Press).

--- , o. C. Wallmo, and Charles Feddema.

1973. Foods of the Rocky Mountain
mule deer. U.S.D.A. Forest Service, Rocky Mountain Forest and Range
Experiment Sta. Res. Paper. RM 111. 3lp.

Prepared by

-£ttl-~I c. %{del
Roland C. Kufeld
Wildlife Researcher

�Table 1. Production of summer vegetation on control and sprayed areas before and 2 and 5 years after spraying.

Species

r968
Percent
Hits compo

Control area
SQra~ area
1971
i974
i968
1971
Percent
Percent
Percent
Percent Expected
Hits camp. Hits camp. Hits camp. Hits compo 71 hitsl!Hits

1974
Percent Expected
camp. 74 hits

Percent change
Significance
due to
of change
sEra~ing 2/
After 2 After 5 over 5 year
peri od '}j
years
years

FOrbs
A cJUUea. tan.u1.0-60.
86
AgM.tache wr.:tJ..ci.6oUa
40
AgMeJ1M gtauca
0
AM:em.u.,'&lt;'a dltacwtcu1.l1.6
2
A:Ltem.u.,'&lt;'atudov.&lt;.ci.an.a
1
Mtelt .6p.
10
Chenopod.&lt;.u.m beJttan.d.&lt;.eJt.&lt;.
1
ClN., -tu.m can.es cen.6
26
Co.e.tom1a .e.J.neaJt1.6
9
C1lep.u., ltunci.l1a.ta
0
Vuph.&lt;.neu.m occi.den.ta.e.e
0
V e.6c U)la1n1a Jt.&lt;.c.hMManJ.i.
0
[JUgolOn .6p.
22
c.uogonu.m wnbe.e.ta.tu.m
2
F/tag ard.o: ovaU.6
0
GaUu.m bolteate
42
Gayophytu.m nutta.e..e.l.&lt;.
0
GeJta.n.1wn Jt.&lt;.ChMM 0 n.&lt;..&lt;.
2
Heten1wn hoope.6u
5
HeJtac.teu.m ta.n.a.tu.m
8
Hydlto phy.e.tu.m 6 endteJt.&lt;.
0
Lac..tuca pu1.c.he.e.ta.
6
Lathnus Leucanthu:
.32
Ugl1.6Uc.u.m
3
L1nM.&lt;.a. v u1.gcrM.6
16
Linum .te.v0.,u
1
Lup.&lt;.nl1.6Mgentel1.6
34

6.02
2.80
0.00
0.14
0.07
0.70
0.07
1.82
0.63
0.00
0.00
0.00
1.54
0.14
0.00
2.94
0.00
0.14
0.35
0.56
0.00
0.42
2.24
0.21
1.12
0.07
2.38

152
58
1
9
1
23
5
23
15
3
7
0
32
0
0
56
0
4
2
17
3
1
155

a
a

13
69

6.36
2.43
0.04
0.38
0.04
0.96
0.21
0.96
0.63
0.13
0.29
0.00
1.34
0.00
0.00
2.34
0.00
0.17
0.08
0.71
0.13
0.04
6.49
0.00

0.00
0.54
2.89

89
80
0
18
0
4
3
13
22
10
3

0
28
0
0
22
3
6
2
10
0
0
98

a

5
1

56

4.66
4.19
0.00
0.94
0.00
0.21
0.16
0.68
1.15
0.52
0.16
0.00
1.47
0.00
0.00
1.15
0.16
0.31
0.10
0.52
0.00
0.00
5.14
0.00

0.26
0.05
2.94

77
42
0
1
0
14
2
8
5
3
0
3
43
10
5
42
0
11
1
6
0
6
71
8
0
0
21

5.55
3.03
0.00
0.07
0.00
1.01
0.14
0.58
0.36
0.22
0.00
0.22
3.10
0.72
0.36
3.03
0.00
0.79
0.07
0.43
0.00
0.43
5.12
0.58

0.00
0.00
1.51

152
49
0
0
0
38
1
13
18
1
1
0
28
5
1
64
0
4
1
11
2

1
131
1
0

a

35

6.81
2.20
0.00
0.00
0.00
1.70
0.04
0.58
0.81
0.04
0.04
0.00
1.25
0.22
0.04
2.87
0.00
0.18
0.04
0.49
0.09
0.04
5.87
0.04
0.00
0.00
1.57

143
57

25

74
54
0
1
0
35
0

17
14

38
55

8

5
4
3
2
35
6
5
30
3
5
2

12

15
1
0

193

149

a

0
1

59

38

3.78
2.76
0.00
0.05
0.00
1.79
0.00
0.41
0.26
0.20
0.15
0.10
1.79
0.31
0.26
1.53
0.15
0.26
0.10
0.77
0.05
0.00
7.62
0.00
0.00
0.05
1.94

86
78

+

5.99 - 14.02
- 15.25 - 31.30
.05
.15

4
11
21

.15
.20
.20
.15

31

- 26.86 + 11.74

21

+ 16.16 + 39.12

.20
.15
7
115

- 32.16 + 29.19

49

- 41.18 - 23.89

.05

----------_._-------------------------------------------------------------------------------------------------------------------------

I
I-'
V&gt;
I

�Table 1.

Production of summer vegetation on control and sprayed areas before and 2 and 5 years after spraying (contd).

Species

1968
Percent
Hits
compo

Control area
Percent change
S[!ra~ area
1971
due to
1974
Signifi cance
1968
1971
1974
Percent
S[!ra~ingY
Percent
of change
Percent
Percent
Expected.
Percent Expected After 2 After 5 over
Hits
compo Hits compo Hits compo Hits compo
5 year
71 hitsl/ Hits compo 74 hits
years
years
period Y
Forbs (coritd2

MeJtte/'l.6.ut 6JLancUc.ana
04moJziUza oc.c1.den.taU..6
Ped.i.c.ui.aJU.6 i11taJj,t
Pvu.dvu.d.i.a giWr.dnvu.
Poiygonum dougla4'£'£
Potent.U.ea. pulc.heJr.JWna
Ptvu.d.i.um aquil.Utum
Rudbec.kla montana
Senec1.o 4 VtJta.
SoUdago c.anaden616
Ste.UalLi.a j amU,tana
SweJl.Ua. ltad.i.a.ta
Tevr.a.xJ.c.um4 p.
ThctUc..tJtum 6 endtvu.
T~agopogon P~en6-i4
UILUc.a gltac,Uen.ta
V,tgu.L~a muUi6.tolta
WyetlUa. alLi.zon-ic.a.

Unknown forb
lotal Forbs

49
3.43
10
0.70
0
0.00
0
0.00
0
0.00
0
0.00
8
0.56
0
0.00
1
0.07
1
0.07
0
0.00
0
0.00
7
0.49
1.47
21
0
0.00
3
0.21
1.26
18
0
0.00
0
0.00
--466 32.61

72
21
0
3
8
1
28
0
6
2
10
0
9
38
3
0
18
0
0
'"

3.01 63
0.88 18
0.00
2
0.13
1
0.33
2
0.04
0
1.17 45
0.00
8
0.25
2
0.08
6
0.42 11
0.00
3
0.38
0
1.59 26
0.13
1
0.00
5
0.75 42
0.00
1
0.00
0

3.30 49
0.94
5
0.10
2
0.05
0
0.10
0
0.00
0
2.36
0
.0.42
0
0.10
0
0.31
0
0.58
0
0.16
1
0.00
0
1.36 29
0.05
1
0.26
3
2.20 28
0.05
0
0.00
3

3.53 48
0.36 29
0.14
0
0.00
3
0.00
6
0.00
0
0.00 14
0.00
0
0.00
8
0.00
5
0.00
3
0.07
5
0.00
2
2.09 72
0.07
0
0.22
1
2.02
3
0.00
2
0.22
0

2.15
1.30
0.00
0.13
0.27
0.00
0.63
0.00
0.36
0.22
0.13
0.22
0.09
3.23
0.00
0.04
0.13
0.09
0.00

37.16 500

36.02 758

33.96

-

-------

868

36.33 709

,

71
16

33
12

1
0
4

45

0
21
7
19
2
20
4
0
59
0

21

2
25

9

a
--

---

888

694

1.69
0.61
0.05
0.00
0.20
0.00
1.07
0.36
0.97
0.10
1.02
0.20
0.00
3.02
0.00
0.10
1.28
0.46
0.00
35.48

62
16

- 32.64 - 46.84

.25

....
I

0\
I

.20
.15
29

+ 59.31 +100.42

43

722

.10
.25
.25
.05
.20

- 14.72

- 3.93

----------------------------------------------------------------------------------------------------------------------------------------------------------

-.15

�Table 1. Production of summer vegetation on control and sprayed areas before and 2 and 5 years after spraying (contd).
Percent change
Significance
due to
Control area
SEra~ area
of change
sEra~ing y
1974
1971
1968
1968
1971
1974
Percent Expected After 2 After 5 over 5 year
Percent Expected
Percent
Percent
Percent
Percent
period 3/
years
years
Hits compo Hits compo Hits compo Hits conp, Hits comp, 71 hits lJ Hits compo 74 hits

Species

Grasses
AgltopyJtOYl sp,
BltomUl&gt;anoma.lUl&gt;
CaJtex &lt;&gt;p.
ECymUl&gt; uYlVteUl&gt;

EI'_ymw) g.taUCUl&gt;
Ph-eeum pltctten&lt;&gt;e
Poa-CaJtex geyeM.
S~OYl
hy~~
S;(;{.pa.f.e;ttVtmanl..

Unknown grass

29
15
8
11
21
2
249
0
0
2

2.03
1.05
0.56
0.77
1.47
0.14
17.42
0.00
0.00
0.14

101
31
8
3
93
7
390
0
6
0

4.23 47
1.30 33
0.33
0
0.13
1
3.89 53
0.29 13
16.32 249
0.00
0
0.25 18
0.00
0

2.46 28
1.73 41
0.00 16
0.05
2
2.78 57
0.68
3
13.05 142
0.00
0
0.94 . 2
0.00
3

2.02 101 4.53
2.95 58 2.60
1.15 31 1.39
0.14 13 0.58
4.11 313 14.02
0.22
7 0.31
10.23 323 14.47
0.00
3 0.13
0.14 23 1.03
0.22
0 0.00

337

23.58

639

26.75 414

21.70 294

21.18 872 39.07

1.03
0.00
0.22
2.78
7.75
0.36

37
0

1.94
4.81
0.00
0.51
4.96
0.05
9.10
0.00
1.89
0.00

605

455

23.26

388

+ 43.92

+ 17.05

.05

70

46

2.35
0.00
0.10
3.02
11.81
0.15

52

- 67.15

- 12.34

.05
.20
.20

74
172

- 49.34 - 20.52
- 12.68 + 34.26

98
32
7

38
94

107

10
97
1
178

326

0

0

45
31

+

75

+190.98 + 28.33

2.10

- 1.02 - 15.55

2.77 - 17.03
+195.62

+ 80.91

.25
.15
.15
.05
.15

I

.25

---Total Grasses

Shrubs
Ame.taYlciUVt
a1.nl6olia
M-te.mu.,ia rudeYl-ta.ta.
POpU£Ul&gt; tJtemu.R.oidu
PItUl1Ul&gt; v.£ltg .£n.£ana
QuVtCUl&gt; gambeUU.
iUbu sp;

33
8
1

103
136
6

2.31
0.56
0.07
7.21
9.52
0.42

45
18
0
151
247
2

1.88 33
0.75 10
0.00
1
6.32 100
10.34 216
0.08
7

1.73
0.52
0.05
5.24
11.32
0.37

64
4.61 23
0.07
0
1
0.07
1
5
65 . 4.68 62
6.99 173
97
0.07
8
1

0

122
198

2
39
231

3

I
•....
-..)

---------------------------------------------------------------------------------------------------------------------------------------------

.25
.20

�Table 1.

Production of summer vegetation on control and sprayed areas before and 2 and 5 years after spraying (contd).

1965
Percent
Hits camp.

Species

Control area
S~ra~ area
,971
,97il·
,~~8
,9'1
1974
Percent
Percent
Percent
Percent Expected
Percent Expected
Hits camp. Hits camp. Hits compo Hits camp. 71 hits l! Hits camp. 74 hits

Percent change
due to
Significance
sEra~ing y
of change
After 2 After 5 over 5 year
years
years
period ~

Grasses· (COlitd)
ROM . .6p.
Sif/nphoJt.i.c.aJtpo.6

.6p.

Total Shrubs
TOTAL ALL PLANTS

8 0.56
328 22.95

19 0.80
394 16..49

623 43.60

876 36.67

1426 99.79

2383 99.75

15 0.79
402 21.07

34
324

2.45 14 0.63
23.34 315 14.11

35
387

442

0.97
22.60

16
394

- 18.64

+ 11.99

.20
.10

587

-- -42.29 600 26.88

839

802

41.00

750

- 28.51

+

.05

-2318

--

1907 99.95 1381

-- ---99.50 2230 99.91

--99.74 1856

784 41.09

19

1951

6.86

- 3.94 + 5.34

--

lIReflects the amount of vegetation one would expect to find on the sprayed area following spraying had spraying not occurred.
~Derived through comparison of sprayed and control areas with adjustments for normal year to year changes in forage production recorded on the control
area. Percent changes are only shown for the more abundant plants.
1Irndicates the highest confidence level at which changes due to spraying during the 5 year period following spraying are significant. Data in the
preceding column show whether most of the significant change occurred after 2 or 5 years. Only changes t,at are significant at the a .05, .10, .15,
.20 or .25 level are shown.
.

I
I-'

oo
I

�Table

2.

Two and 5 year

changes

recorded

T968-Season

11

Forage
value

Hits

Percent
camp.

in available,

seasonal

elk,

Control area
1971
1974
Percent
Percent
Hits
compo Hits
compo

deer

and cattle

1968
Percent
Hits
compo

forage.

1/, £I

Hits

Spray area
1971
Percent
Expected
compo 71 hits ~Hits

1974
Percent Expected
compo 74 hits

Percent change
due to
Significance
sprayi ng §J
of change
After 2 After 5 over 5 year
years
years
period
§j

263
329
0

11.78
14.74
0.00

17.28
23.57
0.00

- 34.06
- 24.88
0.00

Elk Forage
Winter

Highly valuable
Valuable
Least valuable
Pre-

Spring

Summer

327
598
139

to Post-Treatment

Hi9hly valuable
Valuable
Least valuable
Pre-

250
324
16

to Post-Treatment

Highly valuable
Valuable
Least valuable
Pre-

Fall

to Post-Treatment

Highly valuable
Val~able
Least valuable
Pre-

273
344
0

351
439
0

to post-Treatment

19.10
24.07
0.00
Ratio
17.49
22.67
1.12

443
431
0

18.54
18.04
0.00

350
427
0

18.34
22.38
0.00

227
359
0

16.35
25.86
0.00

398
437
0

338
46i
0

312
432
0

ZI
390
650
26

16.32
27.21
1.09

250
480
10

13.10
25.16
0.52

143
316
17

10.30
22.77
1.22

328
702
31

14.70
31.45
1.39

326
633
27

180
509
0

9.20
26.02
0.00

211
467
10

Ratio
22.88
41.85
9.73

571
978
270

23.90
40.94
11.30

376
820
180

19.71
42.98
9.43

303
658
113

21.83
47.41
8.14

712
853
217

31.90
38.22
9.72

548
1000
244

364
953
152

18.61
48.72
7.77

361
839
168

Ratio
24.56
30.72
0.00
Ratio

669
563
0

28.00
23.57
0.00

429
512
0

22.48
26.83
0.00

347 25.00
390. 28.10
0
0.00

840
377
0

37.63
16.89
0.00

654
524
0

418
501

c

21.37
25.61

0.00

419
475
0

+
+

8.20
6.66
0.00

1 : 0.70

1 : 1.08

+ 0.52
+ 10.85
+ 12.54

- 14.87
+ 8.82
-100.00

1 : 1.07

1 : 0.99

+ 29.88
- 14.75
- 11.26

+

0.63

-

9.72

+ 13.46

1 : 1.04

1 : 1.08

+ 28.43
- 28.07
0.00

+

1 : 1.09

-

0.40
5.29

0.00
1 : 1.02

-----------------------------------------------------------------------------------------------------------------------------------------------------------

.10
.05

~
.20

.05
.05

.05
.10

'f

�Table 2. Two and 5 year changes recorded in available, seasonal elk, deer and cattle forage (contd). l/,Y

Season Y

Forage
value

Percent change
Control area
S~ra~ area
due to
Significance
1968
1971
1974
1968
1971
1974
s~ra~in9 §j
of change
Percent
Percent
Percent
Percent
Percent
Expected
Hits camp. Hits comp. flits comp, Hits comp, Hits compo 71 hits l.IHits Percent Expected After 2 After 5 over 5 year
compo 74 hits
years
years
period §!
Deer Forage

\&gt;Jinter

Heavi ly eaten
t~od.eaten
Ughtly eaten

8
273
342

0.56
19.10
23.93

. 18
443
415

0.75
18.54
17.37

10 0.52
1
350 18.34 227
424 22.22 359

0.07
16.35
25.86

0 0.00
263 11.78
337 15.10

2
398
423

0 0.00
338 17.28
464 23.72

3
312
431

Pre- to Post-Treatment Ratio ZI
Spring

Heavily eaten
Mod. eaten
Lightly eaten

8
889
14

0.56
62.21
0.98

18
1354
10

Heavi ly eaten
Mod. eaten
Lightly eaten

388
696
217

27.15
48.01
15.19

0.75
56.68
0.42

10 0.52
1
1076 56.39· 758
7 0.37 17

0.07
0 0.00
54.61 1023 45.83
1.22
42 1.88

Heavily eaten
Mod. eaten
Lightly eaten

415
488
6

29.04
34.15
0.42

Pre- to Post-Treatment Ratio

I
N

0
I

2
1255
9

0 0.00
1016 51.94
3 0.15

3
991
7

-100.00 -100.00
- 18.53 + 2.47
+337.93 - 59.61

.20
.05
.10

- : 0.83 1 : 1.02
637
1047
441

26.66
43.83
18.46

466 24.42 247
934 48.95 732
295 15.46 260

17.80
52.74
18.73

499 22.36
824 36.92
660 29.57

537
1065
476

418 21.37
969 49.54
343 17.54

381
947
310

Pre- to Post-Treatment Ratio
Fall

.20
.10
.10

1 : 0.70 1 : 1.07

Pre- to Post-Treatment Ratio
Summer

-100.00 -100.00
- 34.06 + 8.20
- 20.49 + 7.56

- 7.23 + 9.52
- 22.65 + 2.27
+ 38.43 + 10.63
1 : 0.90

738
635
2

30.89
26.58
0.08

513 26.89 268 19.31
560 29.35 504 . 36.31
7 0.37
1
0.07

602 26.97
445 19.94
8 0.36

643
636
1

449 22.96
566 28.94
3 0.15

413
562
5

.05
.05

1 : 1.06

6.50 + 8.67
-- 30.14
+ 0.64
+304.82

- 43.12

1 : 0.84

1 : 1. 05

--------------~------------------------------------------------------------------------------------------------------------------------------------------

.05
.20

�Table 2. Two and 5 year changes recorded in available, seasonal elk, deer and cattle forage (contd). 11, gj

Season 11

Forage
value

Spray_ area
Control area
1974
T971
T9~W6-g
1971
1968
Percent Expectea
Percent
rxpecteal
Percent
Percent
Percent
P-ercent
compo 71 hits ~Hits comp. 74 hits
Hits compo Hits compo Hits compo Hits compo Hits

Percent change
due to 51 Significance
~ spraying :::.J
of change
After 2 After 5 over 5 year
years
years
period §!

Cattle Forage
Summer

Desirable
108
Intermediate
456
Least desirable 860

7.56 232
31.91 773
60.18 1378

9.71 174 9.12
32.36 503 26.36
57.68 1230 64.47

134 9.65
438 31.56
803 57.85

244 10.93
871 39.02
1115·49.96

247
749
1316

178 9.10
527 26.94
1246 63.70

179
487
1178

- 1.38
+ 16.24

- 15.31
1 : 1.09

- 1.04
8.21
5.74

+
+

.05

: 1.04

Pre- to Post-Treatment Ratio II

----------------------~~~~~~~
lIClassification of study area plants as elk and deer forage was based on the average, relative consumption of those plant species by elk and deer in 48
elk and 99 mule deer food habits studies reported in the literature (Kufe1d 1973b; Kufe1d, Wallmo and Feddema 1973). Plants considered cattle forage are
. those classified as "desirable" and "intermediate" by the U. S. Forest Service, Region 2, Range Analysis Handbook for the foothill shrub, aspen weed and
mountain meadow vegetation types.
£!Table is based on the assumption that due to such factors as degree of plant maturity and snow cover, plant availability to elk and deer will consist
of only shrubs during winter; grasses and shrubs during spring and fall, and forbs, grasses and shrubs during summer. Data are derived from measurements
made during summer.
1!seasons:

Winter = December, January, February; Spring = March, April, May; Summer = June, July, August; Fall = September, October, November.

9:JRef1ects the amount of vegetation one would expect to find on the sprayed area following spraying had spraying not occurred.

?I Derived through comparison of sprayed and control areas with adjustments for normal year to year changes in forage production recorded on the control
area.
(footnotes continued on following page)

I

�Table 2. Two and 5 year changes recorded in available, seasonal elk, deer and cattle forage (contd).
(footnotes contd)
6/

- Indicates the highest confidence level at which changes due to spraying during the 5 year period following spraying are significant. Data in the
preceding
show shown.
whether most of the significant change occurred after 2 or 5 years. Only changes that are significant at the a .05, .10, .15,
.20 or .25column
level are
7/

- The ratio·of elk, deer and cattle forage produced before and after treatment adjusted for normal year to year changes measured on the control areas.
A single value for elk and deer forage was derived by weighting hits on highly valuable elk forage, and heavily eaten deer forage by a factor of "3",
valuable and moderately eaten forage by "2", and least valuable and lightly eaten forage by "1". A.total weighted value was then computed. For
cattle desirables were weighted by "2" and intermediates by "1". Least desirables are not considered cattle forage.
I

N
N
I

�-23-

One year before

spraying

(August 1968)

One year after spraying

(August 1970)

Fig. 1. Vegetation conditions at a permanent photo point on the Beaver Creek
oak study area 1 year before and 1, 2, and 5 years after spraying (Fig. 1
continued on next page).

�-24-

Two years after spraying

(August 1971)

Five years

(August 1974)

after spraying

Fig. 1. Vegetation conditions at a permanent photo point on the Beaver
oak study area 1 year before and 1, 2, and 5 years after spraying.

Creek

�-25July, 1975
JOB PROGRESS REPORT

State of

COLORADO
--------~----~~-----------W-lOl-R-17

Project No.

Game Range Investigations

4

Job Title

Job No.
3
------~------------------Experimental Improvement of
Oakbrush on Deer, Elk and Cattle Ranges - Hightower Mountain

~==~~~~~------~----~~----~~~~~~-----

Period Covered:
Personnel:

April 1, 1974 through March 31, 1975

Roland C. Kufeld and Larry Stewart

ABSTRACT
Effects of burning, spraying and chaining of Gambel oak vegetation on forage
production and elk, deer and cattle use were evaluated 2 years after treatment.
With adjustments for normal vegetation production changes measured on control
units data indicate that summer vegetation increased 50 percent on sprayed
areas, 20 percent on chained areas, and decreased 17 percent on burned sites.
Most of the increase following spraying occurred among grasses which increased 295 percent, and produced 658 more pounds of air dry forage per acre.
Most of the increase on chainings and decrease on burns occurred among shrubs.
Data were grouped according to elk, deer and cattle forage. An explanation
of these terms is presented. Among the treatments tested spraying produced
the largest increases in available elk forage during spring, summer and fall
and particularly spring when production was 2.09 times pre-treatment levels.
Chaining produced the most elk forage during winter with 1.24 times as much
forage. Chaining also increased elk forage production during the other 3
seasons, but was second to spraying. Burned areas supported less available
elk forage during all seasons than existed before treatment. This ranged
from 64 percent of pre-treatment forage production levels during winter to
84 percent during spring.
Chaining resulted in the most available fall and winter deer forage with
1.26 and 1.23 times as much respectively as occurred before treatment.
Spraying yielded the most spring and summer deer forage with 1.40 and 1.13
times as much as prior to treatment. Burning decreased deer forage during
all seasons. On burned areas production ranged from 54 percent of pretreatment levels during winter to 84 percent during summer.
Sprayed plots produced 2.16 times as much summer cattle forage as before
compared with 1.24 times as much for chainings and only 95 percent of pretreatment levels for burns.

(Abstract continued on next page)

�-26-

Burned and chained areas when compared with control and spray areas showed
increased levels of total cell contents, soluble carbohydrate, copper and
zinc in most plant categories (forbs, grasses and shrubs).
Compared with controls elk density per square mile during fall, winter and
spring increased 45 percent on burns, decreased 48 percent on spray areas
and decreased 70 percent on chainings.
Deer density per square mile
during fall, winter and spring decreased 25 percent, 6 percent and 53 percent on burned, sprayed and chained plots respectively.
Relatively low
elk and deer use in the chainings may have been partly due to exceptionally
heavy hunting pressure in the study area vicinity during the fall of 1973,
when game use measurements were made.
Cattle showed a preference for burned areas. Cattle use on the two burns
increased 18.7 cow days per acre compared with a 3.3 cow~day increase on
sprayed areas. Cattle use in one chaining decreased 9.6 days per acre.
However, since cattle were excluded from the other chaining by high creek
flows during the summer of 1973, it is not valid to conclude that chainings
were less preferred than sprayed areas.

�-27-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH
ON DEER, ELK AND CATTLE RANGES - HIGHTOWER MOUNTAIN
Roland C. Kufeld

P. S. OBJECTIVE
To determine the extent to which deer, elk and cattle forage production
and game use can be increased and ~intained by chaining, spraying and
controlled burning on overage Gambel oak winter game ranges.

SEGMENT OBJECTIVE
To determine the extent of vegetation composition, production and nutrient
content changes, and the extent of deer and elk use changes which have resulted from burning, spraying and chaining.

METHODS AND MATERIALS
Study area design and sampling scheme have been described by Kufeld (1970
and 1971). Implementation of burning, spraying and chaining treatments on
the study area as well as procedures and species used in reseeding have been
discussed by Kufeld (1972).

Vegetation Measurements 2 Years
After Burning, Spraying and Chaining
Two year post-treatment vegetation measurements were made in 600 permanent
meter square plots on the Hightower Mountain oak study area during the summer
of 1973, using the same procedures employed during the 1970 pre-treatment
evaluations as described by Kufeld (1971). The 2 year post-treatment measurement period was July 9, 1973, through August 16, 1973. Prior to beginning
vegetation measurements observers were trained in use of the vegetation
measurement technique, and in identification of all plants on the study area.
Pre- and post-treatment vegetation production data were computerized and subjected to covariance analysis by the Colorado State University Statistics
Department.
Statistical significance of vegetation production changes was
tested at the .05, .10, .15, .20 and .25 confidence levels. It was necessary
to test for significance at lower levels because the study involved only 2
replications of each treatment. Inclusion of more replications would have
increased the study area size, workload and manpower requirements to the point
the study would not have been feasible.

Determination

of Nutrient Quality Changes

Samples of plant species designated by Kufeld (1971) as "moisture indicator
plants" and used to reflect changes in vegetation moisture content during

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measurement periods were collected in one unit representing each type of
treatment (Units 5, 6, 7 and 8) between July 24, and July 27, 1973 inclusive and on November 12, 1974. Composite samples of each species
weighing approximately 100 grams, green weight, were collected throughout
each of the 4 habitat improvement units. In treated units samples were
collected from those areas where it was certain burning, spraying and
chaining had been severe.
Samples were analyzed by the Division Research Laboratory to determine
changes in content of dry matter, protein, energy and trace mineral values
2 summers and 3 winters after treatments were applied.
Soil samples were also collected from each of the 4 areas from which plant
nutrient samples were obtained. These were analyzed by the Colorado State
University Soils Laboratory to determine any differences among areas in
soil properties.

Elk, Deer and Cattle Use Measurements
2 Years After Burning, Spraying and Chaining
Accumulated elk and deer pellet groups were removed from all pellet plots
on September 3, 1973. Groups deposited during the fall, winter and spring
were counted May 8, 1974. Data are compared with pre-treatment measurements
made during the same period from the Fall of 1970 to the Spring of 1971.
Two-year post-treatment cattle use changes were measured during the previous
project segment and results reported by Kufeld (1974). Since two year evaluation data on vegetation and elk and deer use were not available last year
the 2-year cattle use evaluation is rediscussed in this report.
When pellet plots in units 1, 5, 6, 7, 8 and a portion of 3 were cleared
September 3, 1973, a record was kept of the number of current year cow chips
in each plot in order to provide an index to cattle use. Cow chip counts in
units 2, 4 and the remainder of 3 were delayed until October 3, because these
units are in a different cattle allotment, and cattle were still grazing
until the end of September.

RESULTS AND DISCUSSION

Vegetation Changes 2 Years
After Burning, Spraying and Chaining
Vegetation production estimates discussed throughout this report refer to
vegetation below a level of five feet.
All production changes have been adjusted for normal year to year fluctuations
in plant growth as measured on control units. This was done by computing the
level of vegetation production one would expect to find on a treated area
following treatment h~d treatment not occurred. This expected value, hereafter referred to as E, is derived by applying the ratio of pre- to posttreatment forage production on the control areas to pre-treatment production

�-29-

on the treatxd areas. The difference between the expected post-treatment
production (E) on the treated areas and the actual production recorded
there is the change produced as a result of treatment.

Effects of Burning on Forbs, Grasses and Shrubs
Summer vegetation production decreased 17 percent 2 years after burning.
Most of the reduction occurred among shrubs which were down 38 percent,
a decrease of 305 lbs of air dry vegetation per acre. The shrub decline
was significant at the a .05 level (Table 1).
Vegetation production changes which occurred within individual plant species
under untreated conditions between the pre- and 2 year post-treatment period
are shown in Table 2. Comparable data for burned areas including levels of
production expected under untreated conditions (E) are shown in Table 3.
The shrub species mainly responsible for the overall decrease in shrubs
as well as in overall vegetation production were Artemisia tridentata
and S)~phoricarpos albus. These produced 153 and 182 fewer pounds of air
dry forage per acre respectively.
Artemisia tridentata did not resprout
after burning. Those plants exposed to fire were completely killed.
Symphoricarpos albus sprouted profusely, especially the first summer after
burning, however overall production was still significantly lower at the
a .10 level 2 years after treatment. Other shrubs among which much sprouting
and regrowth was noted after burning were Amelanchier alnifolia, Artemisia
cana, Prunus virginiana and Quercus gambellii. Like Symphoricarpos albus
most of the sprouting of these species occurred during the first summer
and slowed considerably the second summer. Forage production of several
had returned to or slightly exceeded pre-treatment levels by ~he end of
the second growing season (Table 3). Extensive resprouting of many shrub
species particularly during the first year can be seen in Figure 1, which
shows photos taken at a permanent photo point 1 year before and 1 and 2
years after burning.
Grasses on burned areas decreased 5 percent or 16 lbs/ac, and forbs increased
14 percent or 63 lbs/ac (air dry wt) (Table 1). Two year post-treatment
changes in production of individual grass and forb species on the burned
areas in terms of pounds per acre were relatively minor, even though some
showed fairly large percentage changes (Table 3).

Effects of Spraying on Forbs, Grasses and Shrubs
Spraying resulted in the largest total vegetation increase of the 3 treatments tested. Vegetation production increased 50 percent 2 years after
spraying. Most of the increase occurred among grasses which were up 295
percent and produced an additional 658 lbs/ac (air dry wt). This was significant at the a .10 level (Table 1). Stimulated grass production is evident
in Figure 2, which shows photos taken at a permanent photo point 1 year
before and 1 and 2 years after spraying.
Individual grasses and grasslike plants most responsible for the increase
were Poa sp. and Carex geyeri which together produced 212 percent or 489

�-30-

lbs/ac more air dry herbage (Table 4). Poa sp. and ~
geyeri were
combined in this study because it was often difficult to separate them
when clipping vegetation in meter square sampling plots. Other grasses
which showed proportionately large increases in air dry herbage after
spraying were Agropyron sp. (+42 lbs/ac), Bromus anomalus (+28 lbs/ac),
Dactylus glomerata (+8 lbs/ac), Elymus glaucus (+27 lbs/ac) and Stipa
sp. (+24 lbs/ac) (Table 4).
Forbs also increased after 2 years although not as dramatically as did
grasses. Forbs were up 15 percent or 79 lbs/ac, air dry wt (Table 1).
Those reflecting the largest increases were Achillea lanulosa (+49 lbs/ac),
Galium sp. (+39 lbs/ac), and Taraxacum officinale (+30lbs/ac).
Some
forbs which decreased by relatively large amounts were Lathyrus~
(-53 lbs/ac), and Lupinus argenteus (-37 lbs/ac).
The kill on Quercus gambellii was estimated at 80 to 90 percent during
the first summer after spraying. Extensive initial kills were also observed for Amelanchier alnifolia, Artemisia tridentata, Prunus virginiana
and Symphoricarpos albus. Two years after treatment total shrub production
on spray areas exceeded by 4 percent or 35 lbs/ac that which would have
been expected without spraying. This relatively minor change reflects a
situation where many shrubs, and particularly those just mentioned, have
sprouted back within 2 years to produce approximately as much herbage
as before treatment (Fig. 2).

Effects of Chaining on Forbs, Grasses and Shrubs
Production of vegetation on chained sites increased 20 percent 2 years
after chaining.
Shrubs which produced 19 percent or 167 more lbs/ac of
air dry vegetation accounted for most of the increase. The increase in
shrubs was significant at the a .15 level (Table 1).
Most of the higher shrub production was due to increases of 33 percent or
107 lbs/ac in Quercus gambellii, 65 percent or 90 lbs/ac in Prunus virginiana
and 33 percent or 35 lbs/ac in Amelanchier alnifolia. These 3 species sprouted
profusely after chaining (Fig. 3). Symphoricarpos albus produced 14 percent
or 31 lbs/ac less forage 2 years after chaining, and-resprouting was not as
noticeable on this species as it was in burned and sprayed areas (Table 5).
Forbs increased 25 percent or 108 lbs/ac on chainings (Table 1). Those
making up the bulk of the increase were Achillea lanulosa (+ 29 lbs/ac),
Aster-Erigeron (+16 lbs/ac), Galium (+25 lbs/ac) and Lathyrus-~
(+20 lbs/
ac) (Table 5).
Chainings also supported higher grass production after 2 years with an increase of 17 percent or 65 lbs/ac (Table 1). Post-treatment changes in production of individual grass species on the chainings in terms of pounds per
acre were relatively minor.

Contribution of Seeded Species to Vegetation Production in Burned, Sprayed
and Chained Areas
Plummer et al. (1970) were able to retard regrowth of Gambel oak thickets

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for 14 years by seeding heavily after treating by burning or mechanical
means. Therefore, all areas treated as part of the Hightower Mountain
oak study were seeded at a rate of 18.9 lbs per acre with the following
species! Melilotus officinalis, Medicago sativa (Ladak, Rambler and
Nomad strains), Agropyron cristatum, Agropyron intermedium,Bromus
inermis,
Dactylus glomerata, Festuca ovina, Cercocarpus montanus, and Purshia
tridentata.
Seeding rates for each species were described by Kufeld (1972).
Seeding success after 2 years was negligible, and part of the reason is
believed to be extremely dry weather conditions during the winter, spring
and summer of 1972, the first year after seeding in November, 1971.
U. S. Weather Bureau records from Collbran show precipitation during
January through August, 1972, was only 31 percent of normal and only 53
percent of normal at Glenwood Springs.
During the first summer new plants of seeded species could only be found
in the fire lanes constructed around the burns, and in small patches within
one burn. These consisted of only sparse stands of Agropyron intermedium
and Bromus inermis.
Precipitation improved somewhat the second year. Between January through
August, 1973, precipitation was 84 percent of normal in Collbran and III
percent of normal in Glenwood Springs. Seeded species began to appear the
second summer and Agropyron intermedium, and Bromus inermis began to form
fair stands in the fire lanes and part of one burn (Fig. 4). Agropyron
intermedium only produced 12 Ibs/ac of air dry forage in the burns including
fire lanes, and Bromus inermis produced 3 lbs/ac. The only other seeded
species which produced more than 1.00 lb/ac was Dactylus glomerata which produced 1.12 lbs/ac in the chainings and 8.09 lbs/ac in spray areas. Since
this grass was already present in one spray area it is felt that higher
production was due to existing plants being released by spraying and not
to the seeding effort (Tables 3, 4, 5).
Due to the poor seeding success resprouting of existing shrubs have gotten
a head start, and seeded species will probably not provide enough competition
to retard sprout growth even if the seeded species later become established.
In light of the observation that seeded species have only established themselves in part of one burn, in heavily disturbed spots such as fire lanes,
and parts of the chainings where bulldozer tracks had dug deeply it appears
that better seedbed preparation is needed than was afforded by the treatments used in this study. It may be more effective to burn, aerially seed
as soon as the ashes are cool, and pipe harrow the better burned areas.
This would plant seed as well as clear out burned snags.

Vegetation Responses After Burning Versus Spraying Versus Chaining
Statistical comparisons of vegetation production changes on burned, sprayed
and chained areas are presented in Table 1. Forbs increased from 14 to 25
percent after 2 years on all three treatments. There were no significant
differences among treatments in forb production.
Grass production 2 years after spraying was significantly higher than
burning (P &lt;.10) or chaining (P &lt; .15) . Air dry production of grasses on
the sprayed areas increased 295 percent or 658 lbs/ac compared with a 5

�-32-

percent or 16 lb/ac decrease on burns and 17 percent or 65 lb/ac increase
on chainings.
Shrub production in burns 2 years after treatment was significantly lower
(P &lt; .05) than in sprayed or chained areas. The difference in shrub production between sprayed and chained units was also significant, but only
at the a .20 level.
Changes in Elk, Deer and Cattle Forage 2 Years After Burning, Spraying
and Chaining

Definition of "Elk, Deer and Cattle Forage"--Classification of study area
plants as elk or deer forage for purposes of determining production changes
was based on the average, relative consumption of those plant species by elk
and deer in 48 elk and 99 mule deer food habits studies reported in the
literature (Kufeld 1973; Kufeld, Wallmo and Feddema 1973). Plants considered cattle forage are those classified as "desirable" and "intermediate"
by the U. S. Forest Service in their Region 2, Range Analysis Handbook for
the foothill shrub, aspen weed, and mountain meadow vegetation types.
Since animal preference for individual plant species varies from location
to location the validity of applying "average" consumption data to this
study area may be open to question. However, these averages do represent
a great deal of food habits research, and constitutes the best information
available to date on the foods these animals have been known to consume and
the relative amounts consumed.
For elk the following classifications are used: highly valuable, and least
valuable (Kufeld 1973). For deer plants are classified as he9vily eaten,
moderately eaten, or lightly eaten (Kufeld, Wallmo and Feddema 1973).
Although the terminology used in the elk and deer references is different
the basis for classifying plants according to relative consumption is exactly
the same. "Highly valuable" plants for elk and "heavily eaten" plants for
deer, by definition, generally comprised a major part of a food sample (usually
at least 20 percent) in food habits studies where reported. In a few cases,
plants which comprised less than a major portion of the food sample were
classified as "highly valuable" or "heavily eaten" if their reported contribution to the diet was far in excess of their reported vegetative composition.
"Valuable" plants for elk and "moderately eaten" plants for deer usually
comprised between 5 and 20 percent of the food sample. "Least valuable" plants
for elk and "lightly eaten" plants for deer comprised less than 5 but more than
1 percent.
Not all plants on the Hightower Mountain oak study area were classified
according to their relative preference as elk and deer forage as some had
never been reported in the literature as eaten by elk or deer during some
seasons or at all during the year. The absence of such plants from the
literature possibly suggests that they are of minor importance as elk or deer
forage. Therefore, for purposes of the Hightower Mountain oak study they will
not be considered elk or deer forage.

Definition of "Available Forage"--Data and discussions relating to available
forage are based on the assumption that due to factors such as degree of plant
maturity and snow cover, plant availability to elk and deer on the study area

�-33-

will consist of only shrubs during winter; grasses and shrubs during spring
and fall, and forbs, grasses and shrubs during sunnner. Discussions of
cattle forage relate only to summer at which time all forage is available.

Effects of Burning on Available Elk Forage--After 2 years burned areas
supported less available elk forage during all 4 seasons of the year than
existed before treatment. A pre- to post-treatment ratio using 1.00 as a
basis to represent the level of elk forage production before treatment with
adjustment for normal year to year changes measured on control areas is
shown in Table 6. A single value for elk forage was derived by weighting
Ibs/ac of highly valuable forage by a factor of "3", valuable by "2", and
least valuable forage by "1". A total weighted value was then computed.
Two years after burning the level of available forage production compared
to 1.00 was 0.64 for winter, 0.84 for spring, 0.80 for summer and 0.67 for
fall (Table 6). The largest Ib/ac decreases on burns during winter, sunnner
and fall occurred in the "valuable" category. The decrease in valuable fall
forage was significant at the a .10 level. During spring the largest decrease occurred in the least valuable category although this was not significant even at the a 2.5 level (Table 6).

Effects of Spraying on Available Elk Forage--Spraying produced the most
available spring, summer and fall elk forage of the 3 methods tested after
a period of 2 years. Pre- to 2 year post-treatment ratios of elk forage
production were 1:2.09 for spring, 1:1.72 for summer and 1:1.94 for fall
(Table 6). This was mainly due to the large increases in grasses which
occurred after spraying. Most of the increased spring, summer and fall
forage consisted of plants considered highly valuable elk forage. Covariance analysis indicated the increases in highly valuable .spring, summer
and fall forage were significant, but only at the a .20, .15 and .25 confidence levels respectively.
The increase in valuable fall forage was also
significant at the a .20 level.
Winter elk forage production in sprayed areas was almost unchanged 2 years
after treatment with a pre- to post-treatment ratio of 1:1.01 (Table 6).

Effects of Chaining on Available Elk Forage--Chained areas supported increased amounts of available winter, spring, summer and fall elk forage two
years after treatment. Respective pre- to post-treatment elk forage production ratios were 1:1.24, 1:1.21, 1:1.13 and 1:1.12. Chaining produced
the largest increases in available winter elk forage of the three methods
tested. Among forage categories the only increase due to chaining which
tested statistically significant was at the a .20 level in valuable spring
elk forage (Table 6).

Elk Forage Responses After Burning Versus Spraying Versus Chaining--The
greatest difference between treatments in production of available elk
forage was between burning and sprying, since production of elk forage
for all seasons of the year decreased in burns and increased in spray areas.
Production of available, air dry winter, spring, summer and fall elk forage
in sprayed areas exceeded that of burns by 334, 802, 921 and 991 lbs/ac

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respectively.
These production values represent the totals of highly
valuable, valuable and least valuable forage not weighted for relative
animal consumption.
Covariance analysis showed spray areas significantly higher than burns in the six following classes of forage. Some
of the confidence levels were rather low, however, despite large differences in terms of lbs/ac: highly valuable spring forage (P&lt;.15), least
valuable spring forage (p &lt; .25), highly valuable summer forage (P &lt; .25) ,
valuable surmner forage (P &lt; .15), highly valuable fall forage (P &lt; .20),
and valuable fall forage (P &lt;.05) (Table 6).
Available elk forage production for all seasons in terms of lbs/ac was
also much higher on chainings after 2 years than on burns. Production
of available, air dry winter, spring, sunnner and fall elk forage in
chainings exceeded that of burns by 515, 436, 444 and 451 lbs/ac
respectively.
Production on chainings was significantly higher at the
a .20 level in valuable sunnner forage, and at the a .05 level in valuable
fall forage (Table 6).
Spray areas were higher than chainings in available spring, summer and
fall elk forage by 260, 204 and 211 pounds of air dry forage per acre.
Available winter elk forage in chainings exceeded that of sprayed areas
by 152 lbs/ac (air dry wt) after 2 years. Sprayed areas were significantly higher at the a .20 level than chainings in production of highly
valuable spring and highly valuable surmner elk forage (Table 6).

Effects of Burning on Available Deer Forage--Two years after treatment
burned areas supported less available deer forage during all seasons of
the year than existed before treatment. Relative available deer forage
in burns after 2 years compared to a pre- treatment basis of 1:00 was
as follows: Winter, 1:0.54; spring, 1:0.59; sunnner, 1:0.84; £all, 1:0.66.
Decreases in moderately eaten winter and spring deer forage were significant at the a .15 level, and at the a .05 level in moderately eaten fall
forage (Table 7).
.

Effects of Spraying on Available Deer Forage--Spraying produced increased
amounts of available spring, summer and fall deer forage. Pre- to posttreatment forage production ratios were 1:1.40, 1:1.20 and 1:1.13
respectively.
Available winter deer forage was about unchanged 2 years
after spraying with a pre- to post-treatment ratio of 1:1.02. The increases in moderately eaten and lightly eaten spring deer forage were
significant at the a .15 and .10 levels respectively.
The increase in
lightly eaten sunnner deer forage was significant at the a .15 level (Table

7).
Effects of Chaining on Available Deer Forage--Chaining increased available
deer forage during all seasons after 2 years. Pre- to post-treatment
forage production ratios in chainings were: Winter, 1:1.23; spring, 1:1.17;
summer 1:1.18, and fall, 1:1.26. The only increase which was statistically
significant was at the a .10 level in moderately eaten spring forage (Table

7).

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Deer Forage Responses After Burning Versus Spraying Versus Chaining-Available deer forage 2 years a.fter treatment was much more abundant
in sprayed than burned areas during all seasons. Production of available, air dry deer forage in sprayed areas exceeded that of burns during
winter, spring, summer and fall by 392, 988, 894 and 422 Ibs/ac respectively.
The following differences in available forage production between burn and
spray areas were statistically significant:
Lightly eaten winter forage
(P &lt; •OS), heavily, moderately and lightly eaten spring forage (P &lt;.25,
P &lt; .05, and P &lt; .10 respectively), lightly eaten summer forage (p &lt; .10) ,
and moderately eaten fall forage (P &lt; .05) (Table 7).
Production of available, air dry forage in chainings was also higher than
burns during all seasons by the following amounts in lbs/ac: Winter, 546;
spring, 562; summer, 518; and fall, 532. Production of available moderately
eaten spring, summer and fall deer forage on chainings was significantly
higher than burns at the a .05, .20 and .05 levels respectively (Table 7).
Spraying produced 426 lbs/ac more available air dry forage than chaining
during spring, and 376 lbs/ac more during summer. Production on spray
areas was significantly higher than chainings in lightly eaten spring and
summer deer forage (P &lt; .10 and P &lt; .15). Chaining produced 153 lbs/ ac more
available air dry forage than spraying during winter, and 110 lbs/ac more
during fall. Production on chained areas was significantly higher (P&lt;.lO)
than spraying in lightly eaten winter forage (Table 7).

Effects of Burning on Cattle Forage--Slightly less summer cattle forage was
found in burned areas after 2 years. The pre- to 2 year post-treatment
forage production ratio for desirable and intermediate plants was 1:0.95.
Burning, however, resulted in a significant decrease (P &lt;.05) in abundance
of least desirable plants after 2 years (Table 8).

Effects of Spraying on Cattle Forage--Substantial increases in summer cattle
forage were observed on sprayed areas after 2 years. The pre- to 2 year posttreatment forage production ratio was 1:2.17. Among intermediate plant species
the increases were significant at the a .15 level (Table 8).

Effects of Chaining on Cattle Forage--A pre- to post-treatment forage production ratio of 1:1.24 for desirables and intermediates was recorded for
chained areas 2 years after treatment. Chaining, however, also brought
about an increase in least desirable plants that was significant at the a .15
level (Table 8).

Cattle Forage Responses After Burning Versus Spraying Versus Chaining--Sprayed
areas contained 653 more lbs/ac of air dry, summer cattle forage 2 years after
treatment than was found in burns. Most of the spray area increase occurred
among intermediate species. These were significantly more abundant at the
a .15 level in spray areas than in burns. Least desirable plants, however,
were also more abundant in spray areas by 375 lbs/ac. This was significant at
the a .05 level.

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Production of summer cattle forage was 193 lbs/ac higher in chainings
than burns, although this was not significant even at the a .25 level.
Least desirables were 405 lbs/ac more abundant in chainings than burns
and this difference was significant at the a .05 level.
Summer cattle forage production in sprayed areas exceeded that of chainings
by 461 lbs/ac. Among intermediate plants the increase was significant at
the a .15 level. Least desirables were 30 lbs/ac more abundant in spray
than chained units and this difference was significant at the a .15 level
(Table 8).

Nutrient and Mineral Content of Forage Collected on Control, Burned, Sprayed
and Chained Areas During the Second Summer and Third Winter After Treatment
Results of soil tests are presented in Table 9. These tests show no major
differences between areas in soil nutrients or other properties except for
nitrogen. Tests showed 24 ppm of nitrogen in the control compared with 2 ppm,
1 ppm and 4 ppm in the burned, sprayed and chained areas respectively.
Due
to the similarity of other soil property values among areas, and the fact
that this control area is located directly between the sprayed and burned
units the author doubts if the 24 ppm value in the control really represents
a true measurement of nitrogen in the control.
Summer plant collections for nutrient analysis (July 24 through 27, 1973)
consisted of the following species: Forbs - Achillea lanulosa, Agastache
urticifolia, Cirsium canescens, Galium boreale, Grindelia aphanactis,
Helenium hoopesii, Iris missouriensis, Lathyrus leucanthus, Ligusticum
porteri, Lupinusargenteus,
Rudbeckia montana, Swertia radiata, and Thalictrum
fendleri. Grasses - Agropyron smithii,Agropyron
trachycaulum, Bromus anomalus
and Poa pratensis. Shrubs - Amelanchier alnifolia, Artemisi~ cana,
Chrysathamnus viscidiflorus, Prunus virginiana, Quercus gambelIII: Rosa nutkana,
and Syrnphoricarpos albus.
Winter plant collections (November 12, 1974) consisted of the same shrub
species collected during summer and 2 grasses (or grasslikes) Carex geyeri
and Poa pratensis. Since fewer grasses including one of a different species
were collected during winter, summer and winter data for grasses are not
directly comparable. Forbs were not collected during winter as they were
not available.
Nutrient analyses for plants collected during summer are presented in Table
10, and in Ta~le 11 for plants collected during winter. All significant
differences shown in these tables and discussed in ensuing paragraphs on
nutrient and mineral analysis refer to the 5 percent confidence level.
Except for sodium in forbs, nutrient and mineral content of plants in the
spray area was not significantly different from plants in the control. Most
of the significant differences between treatments in plant nutrient and
mineral content occurred in forbs. Most of the significant differences in
nutrient and mineral content occurred between control and burn; control and
chain; burn and spray and chain and spray. Except for iron in shrubs during
summer and copper in grasses during winter there were no significant differences in plant nutrient and mineral content between the burned and chained
area. In general, burned and chained areas when compared with the control

�-37-

and spray areas showed increased levels of total cell contents, soluble
carbohydrate, zinc and copper in most plant categories {forbs, grasses
and shrubs (Tables 10 and 11).
Specific differences in plant nutrient and mineral content are discussed
in detail in the following paragraphs.

Comparison of Control and Burn Areas in Plant Nutrient and Mineral Content

Protein--Protein content of shrubs in the burn was significantly lower than
the control during summer and slightly lower during winter. During summer
forbs and grasses in the burn also had lower protein levels, but these were
not statistically significant.

Total Cell Content and Soluble Carbohydrate--During summer total cell content
and soluble carbohydrate levels in the burn were higher in grasses and significantly higher in forbs. These values were also higher in the two grasses
collected during winter. Among shrubs total cell content and soluble
carbohydrate levels in the burn were lower than controls in summer, but higher
in winter although neither difference was significant.

Copper and Zinc--During summer copper and zinc levels in forbs and grasses
were significantly higher in the burn than the control. During winter significantly higher copper levels were found in the two grasses collected in
November. Zinc levels in those grasses were also higher, but the increase
in zinc was not significant. Among shrubs on the burn copper and zinc was
higher in summer. Zinc was also higher in shrubs in winter, ~ut copper
dropped slightly below control area levels. The differences between burn
and control areas in copper and zinc levels in shrubs did not show statistical
significance.

Comparison of Control and Spray Areas in Plant Nutrient and Mineral Content
Sodium--During summer sodium levels among forbs in the spray area were significantly higher than on the control, and slightly higher in grasses. Sodium
levels in shrubs were slightly lower than in controls during summer, but
considerably higher in winter, although neither difference was statistically
significant.

Comparison of Control and Chained Areas in Plant Nutrient and Mineral Content

Total Cell Content--During summer total cell content of forbs on the chaining
was significantly higher than the control. Higher total cell content levels
were also found in shrubs and grasses during summer and shrubs during winter,
but these higher levels were not significant at the a .05 level.

Protein--Forb protein levels on the chaining during summer were significantly
higher than the control. Protein levels in grasses and shrubs during summer

�-38-

and shrubs during winter were slightly lower than the control.

Ether Extract--Forbs on the chaining had significantly higher ether extract
levels than the control during summer. Compared with the control ether
extract in shrubs was slightly higher in chainings during summer and winter,
but grasses during summer contained slightly less ether extract than on the
control.

Soluble Carbohydrate--Soluble carbohydrate levels in forbs and grasses were
'significantly h~gher than control area levels during summer. Shrubs also
contained more soluble carbohydrate than the control during summer and
winter, but the increased amount was not significant at the a .05 level.

Iron--Iron content of forbs and grasses on the chaining was significantly
below those of the control during summer and was the lowest of the four
types of treatments. Shrubs also contained less iron than the control or
other treatments but the lower iron levels for shrubs were not significant
at a .05.

Copper and Zinc--During summer copper and zinc levels in forbs and grasses
were significantly higher in the chaining than in the control. Higher
levels of copper and zinc were also found in shrubs on the chaining during
summer and winter, but the increases were not statistically significant.

Potassium--Less potassium was found in forbs, grasses and shrubs during
summer on the chaining as compared with the control. The decreased potassium
levels for forbs was significant.
During winter, potassium levels in shrubs
were slightly higher than in the control.

Comparison of Burned and Sprayed Area in Plant Nutrient and Mineral Content

Total Cell Contents and Soluble Carbohydrate--Forbs in the burn contained
significantly more total cell contents and soluble carbohydrate during summer
than those in the sprayed area. Total cell contents in grasses and shrubs
were also higher during summer in the burn than the sprayed area, but not
significantly higher. During winter total cell content and soluble carbohydrate in shrubs were slightly higher in the spray area.

Copper and Zinc--During summer copper and zinc levels in forbs, grasses and
shrubs were higher in the burn than the sprayed area. Levels which were
significantly higher in the burn during summer were copper in forbs and
grasses and zinc in forbs. During winter copper content in the two grass
species collected in November was significantly higher in the burn than the
spray, but winter copper as well as zinc content of shrubs in the burn was
slightly lower than in the spray.

�-39-

Manganese--During summer manganese content of forbs, grasses and shrubs
was higher in the sprayed area than the burn. This was significant for
forbs. During winter, however, manganese in shrubs was slightly lower
in the spray area than the burn.

Sodium--During summer sodium was significantly higher in forbs on the spray
area and slightly higher in grasses than on the burn. Among shrubs sodium
was slightly lower on the spray area during summer but higher than the burn
during winter. The sodium differences in shrubs, however, were not significant at a .05.

Potassium--Forbs, grasses and shrubs in the sprayed area contained more
potassium than in the burn during summer. Increased potassium in summer
was significant for forbs but not for shrubs and grasses. During winter
potassium in shrubs was slightly lower in the sprayed area.

Comparison of Burned and Chained Area in Plant Nutrient and Mineral Content

Iron--During summer iron content of forbs on the chaining was significantly
lower than on the burn. Iron was also lower in shrubs and grasses on the
chaining during summer and in shrubs during winter though these differences
were not significant at a .05.

Copper--Copper content of the two grasses collected in November was significantly higher on the burn than on the chaining.

Comparison of Sprayed and Chained Area in Plant Nutrient and Mineral Content

Total Cell Contents and Soluble Carbohydrate--Total cell contents and soluble
carbohydrate in forbs, grasses and shrubs were higher in the chaining than
the sprayed area during summer. The higher levels for forbs were significant,
but not for grasses and shrubs. During winter total cell contents and soluble
carbohydrate were slightly higher in the sprayed area.

Protein--Protein level in forbs on the chaining was significantly higher than
on the sprayed area during summer. During summer grass protein levels were
slightly higher on the sprayed area. Among shrubs protein was slightly
higher on the chaining during summer, but during winter it was higher on the
sprayed area.

Ether Extract--Ether extract content of forbs during summer was significantly
higher on the chaining than the sprayed area. It was slightly higher in
shrubs on the chaining during summer and winter. Among grasses collected
during summer ether extract was nearly the same level on both the sprayed
and chained areas.

�-40-

Iron--Iron levels in forbs, grasses and shrubs on the chaining were lower
than sprayed areas during summer, and iron content of shrubs was also
lower on the chaining in winter.
Iron was significantly lower in chained
area forbs.

Copper--The chaining showed significantly higher copper levels in forbs
and grasses during summer than did the spray area. Copper levels in
shrubs were also slightly higher in the chaining during both summer and
winter.

Zinc--Zinc levels in forbs on the chaining were significantly higher during
summer than on the sprayed area. More zinc was also found during summer
in grasses on the chaining and in chained area shrubs during both summer
and winter, but these levels were not significantly higher than the spray
area.

Elk Use Changes 2 Years After Burning, Spraying and Chaining
Despite declines in available, seasonal elk forage on burns 2 years after
treatment compared with sizable increases on sprayed and chained areas elk
exhibited a significantly greater (P&lt;.lO) preference for burned units.
Elk per square mile increased 45 percent on burns during the fall, winter
and spring period, and decreased 48 percent on sprayed areas and 70 percent
on chainings (Table 12). Compared with control areas, however, the changes
in elk use due to burning, spraying and chaining, though relatively large
percentagewise, were not statistically significant at the a .10 level.
Reasons for apparent preference of elk for burned areas over those sprayed
and chained cannot be pinpointed.
Two factors, however, which may have
influenced the attractiveness of burned areas to elk are changes in cover
and plant nutrient content.

Cover Changes
Burning did not denude the area but varied from almost total incineration
of all vegetation with just a few dead snags left standing (Fig. 5) to only
partial removal of the understory with little or no damage to the larger
oaks (Fig. 6). On much of the area most of the understory was removed with
partial destruction of the overstory (Fig. 7). The overall result of burning
was creation of openings interspersed with patches of standing oak vegetation
suitable for escape cover and shade (Fig. 8).
In sprayed areas most of the shrub vegetation was either totally or partially
killed, but the dead material remained standing leaving a thicket only
slightly more open and accessible to large animals than before treatment (Fig.
2) .

After 2 years chained areas were very open with little escap~ cover and no
trees large enough to provide shade (Fig. 3). Forest openings have been
found to receive increased use by elk and deer (Patton 1974; Pearson 1968;
Reynolds 1962a, 1962b, 1966a, 1966b; Wallmo 1969). However, during the
fall of 1973, when 2 year post-treatment game use measurements were being

�-41-

made this study area received unusually heavy hunting pressure particularly for deer as efforts were being made to reduce the deer herd.
Heavy hunting pressure coupled with the lack of cover in chainings may
have caused elk to avoid them during that period, and resulted in the
substantial drop in elk use recorded on chainings 2 years after treatment (Table 12).

Plant Nutrient Content Changes
Elk preference for burned areas may have been influenced to some extent
by increased levels of certain nutrients and minerals such as total cell
contents, soluble carbohydrate, copper and ZLqC. As described in detail
in the section on plant nutrients increased amounts of these were found
in some plant categories (forbs, grasses and shrubs) on the burned areas
as compared with control and spray areas. Since relatively little difference was observed between the burn and chaining in plant content of these
nutrients and minerals, however, any influence exerted by increased plant
nutrient and mineral content must have been coupled by other attracting
factors such as the previously mentioned cover changes.

Deer Use Changes 2 Years After Burning, Spraying and Chaining
Deer use on the study area during the 2 year post-treatment game use measurement period (Sept. 3, 1973 to May 8, 1974) was very light due to 2 factors.
Unusually heavy hunting pressure during the fall of 1973, moved deer onto
adjacent private lands, and record snowfall during the winter of 1973-74
forced deer to leave the area earlier and return later than normal. Because of low deer use it was felt necessary to make measurements another
year to determine if 1973-74 data provide a true picture of deer response
to the various treatments. Thus, pellet plots were cleared in September,
1974, and accumulated pellets will be counted in May, 1975.
Data on deer use 2 years after treatment are presented in Table 13 of this
report for purposes of maintaining a record in progress reports even though
the validity of these data is presently unknown. One facet of the data
which also occurred in elk use data is a large drop in deer use on chainings
after treatment. When tested statistically this drop was significant at the
a .05 level. As discussed in the section on elk use this is attributed, at
least in part, to unusually heavy hunting pressure in the study area vicinity
during the fall of 1973, coupled with the lack of escape cover in chained
areas.

Cattle Use Changes 2 Years After Burning, Spraying and Chaining

Control Areas
Data show cattle use increases for both control areas with an average increase of 4.1 cow days use per acre. Much of the increase in control unit
6 is believed due to movement of cattle around unit 7 (a burned area) and
between units 5 (a sprayed area) and 7 to which cattle were attracted.
Unit 6 lies directly between units 5 and 7. Thus, there was very little,
if any, actual change in cattle use on the control areas.

�-42-

Burned Areas
The most preferred treatment by cattle two years after treatment was
definitely burning, even though cattle forage production decreased on
burns compared to sizable increases on sprayed and chained areas. Both
burns showed the largest increases in cattle use after treatment, and
these were significant at the a .10 level (Table 14). Cattle use in
burned units increased 15.2 and 22.1 cow days use per acre respectively,
and averaged 18.7. Cattle preference for burned areas was also obvious
from field observations.
Preference for burns was approximately 6 times
greater than for the sprayed or chained areas.
The reasons for cattle preference for burns over sprayed and chained
areas have not been specifically identified. Possible reasons are creation of openings interspersed with patches of cover and shade in burned
areas, and increased amounts of certain nutrients and minerals in plants
found on burns. These changes may also have been responsible for increased
elk preference for burned areas over those sprayed and chained, and are
discussed in detail in the previous section on "elk use changes 2 years
after burning, spraying and chaining."

Sprayed Areas
One spray area (unit 5) showed a substantial increase in cattle use as a
result of spraying while the other (unit 2) showed a slight decrease (Table
14). According to the data, which show an average increase of 3.3 cow days
use per acre in sprayed areas, spraying would appear to be the second most
preferred treatment by cattle. Since the cattle use change on the combined
spray areas is not significant at the a .10 confidence level, and in light
of circumstances described in the following discussion on chaining, it is
not valid to conclude that cattle preferred the sprayed areas more than the
chained areas at the end of the second year following treatment.
Chained Areas
No cattle use was recorded in chained unit 3 in 1973 (Table 14) partly due
to the fact it is on the south side of Buzzard Creek, and high water during
the spring and summer of 1973 kept Buzzard allotment cattle on the north
side. This unit lies in two cattle allotments and is difficult for cattle
from both allotments to find. The almost impenetrable oak thicket in unit
3 was opened substantially by chaining, and it is felt cattle use would have
occurred in 1973, if cattle had been able to get to the area.
Cattle use in the other chaining (unit 8) dropped from 18.3 cow days use
per acre in 1970 to 8.7 in 1973. It appears that cattle were simply drawn
off unit 8 into the more preferred burned area of adjacent unit 7. This
occurred even though forage conditions in unit 8 were improved for cattle
by chaining. Because the change in cattle use on chainings is not significant at the a .10 level, and due to inability of cattle to find and get
into unit 3, it is not valid to conclude that chained areas were any less
preferred by cattle than sprayed areas. Had it not been for existence of
the highly preferred burned areas cattle may have exhibited a greater
preference for areas treated by spraying and chaining.

�-43-

LITERATURE CITED

Julander, Odell. 1955. Determining
J. Range Manage. 8(4):182.

grazing use by cow-chip counts.

Kufeld, Roland C. 1970. Experimental improvement of oakbrush on deer,
elk and cattle ranges - Hightower Mountain.
Colo. Div. Game, Fish
and Parks. Game Research Rept. Proj. W-10l-R-12, WP 4, J3. July,
1970, Part 1. pp. 113-126.
1971. Experimental improvement of oakbrush on deer, elk and
-----cattle ranges - Hightower Mountain.
Colo. Div. Game, Fish and
Parks. Game Research Rept. Proj. W-10l-R-13, WP 4, J3. July, 1971,
Part 1. pp. 23-86.
1972. Experimental improvement of oakbrush on deer, elk and
cattle ranges - Hightower Mountain.
Colo. Div. Wildlife.
Game
Research Rept. Proj. W-10l-R-14, WP 4, J3. July, 1972, Part 2.
pp. 81-97.
1973. Foods eaten by the Rocky Mountain elk.
26 (2)106-113.

J. Range Manage.

1974. Experimental improvement of oakbrush on deer, elk and
cattle ranges - Hightower Mountain.
Colo. Div. Wildlife.
Game
Research Rept. Proj. W-10l-R-16, vW4, J3. July, 1974, Part 1.
pp , 17-23.
Kufeld, Roland C., O. C. Wallmo and Charles Feddema. 1973. Foods of
the Rocky Mountain mule deer. U.S.D.A. Forest Service, Rocky Mtn.
For. and Range Exp. Sta. Res. Pap. RM-lil. 3lp.
Plummer, A. Perry, Donald R. Christensen, Richard Stevens and Kent R.
Jorgensen.
1970. Intermediate wheatgrass, smooth brome and other
herbs open and control Gambel oak thickets. In Highlights, results
and accomplishments of game range restoration studies. Utah Div.
of Fish and Game. Pub. No. 70-3. pp. 26-30.
Patton, David R. 1974. Patch cutting increases deer and elk use of a
pine forest in Arizona. J. For. 72(12):764-766.
Pearson, Henry A. 1968. Thinning, clearcutting and reseeding affect
deer and elk use of ponderosa pine forests in Arizona. U.S. For.
Ser., Rocky Mtn. For. and Range Exp. Sta. Res. Note RM-119, 4p.
Reynolds, Hudson G. 1962a. Use of natural openings in a ponderosa pine
forest in Arizona by deer, elk and cattle. u.S. For. Ser., Rocky
Mtn. For. and Range Exp. Sta. Res. Note No. 78, 4p.
1962b.

Effect of logging on understory vegetation and deer use

in a ponderosa pine forest in Arizona by deer; elk and cattle.

U.S. For. Ser., Rocky Mtn. For. and Range Exp. Sta.
80.
7p.

Res. Note No.

�-44-

Reynolds, Hudson G. 1966a. Slash cleanup in a ponderosa pine forest
affects use by deer and cattle. U.S. For. Ser., Rocky Mtn. For.
and Range Exp. Sta. Res. Note RM-64, 3p.
1966b. Use of openings in spruce-fir forests of Arizona by
elk, deer and cattle. U.S. For. Ser., Rocky Mtn. For. and Range
Exp. Sta. Res. Note RM-66, 4p.
Wal1mo, O. C. 1969. Response of deer to alternate strip c1earcutting
of lodgepole pine and spruce-fir timber in Colorado.
U.S. For.
Ser., Rocky Mtn. For. and Range Exp. Sta. Res. Note RM-141, 4p.

Prepared by

;e~-udC,.~
Roland C. Kufeld
Wildlife Researcher

�Table 1. Changes in air dry forage production recorded within forb, grass and shrub categories two years after burning, spraying and
chaining.

Plant
type

Change due
to burnt nq J.I
lbs/ac
percent

Change due
to s[:!ra~i
ng
1bs/ac
percent

Change due
to chaining
16s7ac
percent

Confidence level where treatment difference is significant ~
Control
Control
Control
Burn
Burn
Spray
&amp; Burn
&amp; Spray
&amp; Chain
&amp; Spray
&amp; Chain
&amp; Chain

Forbs
Grasses
Shrubs

+ 63.47
- 16.48
+305.36

+ 78.51
+658.05
+ 34.83

+ 14.96
+295.33
+ 4.28

+107.53
+ 64.60
+166.83

.05

Totals

-257.37

+780.48

+ 50.27

+340.08

+14.49
- 5.24
-38.18

--16.59

+25.34
+16.70
+18.54

.10
.15

.10
.05

.05

.15
.20

-+19.89
I

.pVI
I

lJDerived through comparison of treated and control areas with adjustments for normal year to year changes in forage production recorded on
the control areas.
~Indicates the highest confidence level at which treatment differences are Significant.
the a .05, .10, .15, .20 or .25 level are shown.

Only treatment differences that are significant at

�Table 2. Production
of air
spraying and chaining.

Plant

species

dry summer vegetation

Percent
1970

recorded

in the control

composition
1973

areas

before

and two years

following

Lbs of vegetation
produced.per
acre
1970 .
1973

1bslac

24.15
4.52
0.00
0.30
0.18
0.00
0.18
0.18
1.49
0.00
0.00
0.42
63.64
0.00
0.00
0.18
1.19
0.00
1.31
0.71
7.26
0.00
1.84
0.18
0.00
6.90
0.00
0.00

28.13
- 0.36
+ 0.59
+ 1.61
- 0.18
0.00
- 0.18
- 0.12
+ 0.71
0.00
0.00
- 0.06
-18.14
0.00
0.00
+ 0.06
+ 0.24
0.00
- 0.30
- 0.12
+ 2.32
+ 0.12
+ 0.95
+ 0.06
0.00
+ 3.75
+ 0.12
0.00

burning,

Change 1970 to 19.73 ]j
percent

~
AcJUUeo.
.tal1LLi'.oM.
Ag ct6.to.cJt e f.lIl.UcJ. 6 oUa
Ag 06 (21(M 9 .e.o.u.co.
AUlWII o.cwnbta..tLUn
Al'lC:VW.6ace .6 ep.tentJUa naLU
AQ[(.Ue.9&lt;'0. co eJtu..f.ea.
Alto.D-i..6 ciltwwnoneU
Alto.DM rUM u..to.
AJU~l'ta.Jt&lt;.o.
co I1ge6.ta.
AJt.te.Jnl..6.i.0.b.i.el1~

AJt.tcn0.,-Lo. ciJto.cuncu.£.u.6
AJt.te.Jn~.i.a fudo vo do.na.
A~;(:eJt - EJt.i.geno n
A6 .tl!.aga.e.u..6hay den.&lt;.o.nu..6
Ba..f..6wno Ith.i. ZOo .6 ag.&lt;..t:ta.:ta.
BeJtb~
Itepelu,
Ca.e.OcJlOJt.tu..6gu.n~onU
Co.p.6 eUo. buMo.-pct6.tOItM
Ca.6.t.i.l.e.eja .e..I.na.Jt&lt;.a.e60Uo.
CItcn a po eUwn b eJt.f.a.neUeJt.i.
C.Uv~.i.wn cen.to.VJ1.eae
Coilom&lt;.a linea.Jt.i..6
Como.l1ciJto.wnbe..Ua:to.
Cymop;(:VLU6 pUltpUlteu..6
Ve.e.pfz.i.n.i.wn I!Won.&lt;.
Vefph-i.nLum accJ.del1.;(:a.e.e
Ve6 cuJt£U.lUo. .6Oph-i.o.
V-i..6POltwn .tJtacJLYcCVtpwn

1.41
0.26
0.00
0.02
0.01
0.00
0.01
0.01
0.09
0.00
0.00
0.03
3.66
0.00
0.00
0.01
0.07
0.00
0.07
0.04
0.43
0.00
0.10
0.01
0.00
0.40
0.00
0.00

3.26
0.26
0.04
0.13
0.00
0.00
0.00
0.00
0.15
0.00
0.00
0.02
2.82
0.00
0.00
0.02
0.09
0.00
0.06
0.04
0.59
0.01
0.17
0.02
0.00
0.65
0.01
0.00

52.28
4.16
0.59
1.90
0.00
0.00
0.00
0.06
2.20
0.00
0.00
0.36
45.50
0.00
0.00
0.24
1.43
0.00
1.01
0.59
9.58
0.12
2.80
0.24
0.00
10.65
0.12
0.00

+116.50
7.89

-

+ 48.00

I

~

'"
I

- 28.50

+ 20.00
- 22.73
+ 31. 97
+ 51.61
+ 54.3',

-----------------------------------------------------------------------------------------------------------------------------------------------

�Table 2. Production of air dry summer vegetation recorded in the control areas before and two years following burning,
spraying and chaining (contd).

Plant species

Percent comQosition
1970
1973

Lbs of vegetation
~oduced
Qer acre
o
1973

Change 1970 to 1973 JJ
percent

, bs/ac

Forbs (contd)
~Logonum unbettatum
Ep.uob.[um paMQufa.tum
Fn4gak.[a amen1Qana
Ga.Uwn
Gentiana he;tvr.of.Jepa.la
Gvw.111um
Gilia
a.gg.l!.£gaia
GJt.[nde-Ua aphanaQ;Uf.,
HaQ/~e-Ua .f.ep:tophyUa.
He.f.en-&lt;-um hOOPMU
HydJtophy.e.tum Qap-i:ta.tum
HydJtophy.e.tl@ 6end.een1

Iw

m..Wf.JOuMeMM

Lac.tu.Qa .s Qak.[o.ta
Lappufa .l!.edowf.Jk1i
La.thy.l!.u.f.J
- V.[ua
Lep.[dium deM16.f.o.l!.um
Ugu.f.JtiQum
Unum .f.ew.u.,U
Lomatium .f.ep:toQanpum
Lu.p.[nu.f.JaJl.gen:teLV~
Madia g.f.omvr.a.ta
MediQago f.Jativa
Me.e-Uo.:tu.f.Jo6Mun~
MeA:tel1,6.[a 6JtanUf.JQana
MeA:teM.[a .tan c.eo.ea.:ta
Mo.edav.[Qa panv.[6.f.oJta

0.02
0.00
0.00
1.08
0.01
0.37
0.00
0.21
0.05
0.30
0.00
0.12
0.00
0.00
0.00
7.32
0.00
1.57
0.20
0.00
7.58
0.23
0.00
0.00
0.49
0.00
0.00

0.07
0.04
0.00
1.01
0.02
0.18
0.01
0.28
0.00
0.12
0.01
0.10
0.00
0.00
0.00
6.78
0.01
1.29
0.19
0.00
4.63
0.01
0.00
0.00
0.48
0.00
0.00

0.42
0.00
0.00
18.62
0.12
6.30
0.00
3.45
0.83
4.94
0.00
2.02
0.00
0.00
0.00
124.43
0.00
27.12
3.63
0.00
131.50
3.75
0.00
0.00
7.97
0.00
0.00

1.13
0.65
0.00
16.12
0.30
2.85
0.12
4.22
0.00
1.78
0.12
1.49
0.00
0.00
0.00
107.83
0.12
20.82
3.33
0.00
75.83
0.18
0.00
0.00
7.14
0.00
0.00

+ 0.71
+ 0.65

0.00
- 2.50
+ 0.18
- 3.45
+ 0.12
+ 0.77
- 0.83
- 3.15
+ 0.12
- 0.54
0.00
0.00
0.00
-16.59
- 0.12
- 6.30
- 0.30
0.00
-55.67
- 3.57
0.00
0.00
- 0.83
0.00
0.00

- 13.42
- 54.72
+ 22.41
I

- 63.86
- 26.47

- 13.34
- 23.25
- 8.20
- 42.33
- 95.24
- 10.45

-----------------------------------------------------.---------------------------------------------------------------------------_._---------

.p-

....•
I

�Table 2. Production of air dry summer vegetation recorded in the control areas before and two years following burning,
spraying and chaining (contd).

Plant species

Lbs of' vegetation
,roduced ~er acre

Percent com~osition
1970

1973

, 9 1i

1973

0.00
0.00
0.00
16.06
4.88
0.00
3.09
0.54
0.00
0.00
0.06
0.00
0.00
3.45
0.00
0.00
2.02
0.00
0.00
0.00
0.00
1.01
1.90
13.44
0.00
2.85
32.71

0.00
0.06
0.00
13.74
1.31
0.30
0.65
0.30
0.00
0.00
0.54
0.00
0.06
4.10
0.00
0.00
0.71
0.00
0.12
0.89
0.00
0.83
0.83
27.30
0.06
6.66
24.92

Change )970 to 197311
16s7ac
percent £(

Forbs {contd}
Oenotheha cae6pLto~a
Opwt-tLa
OMItOC.MpU6 .eu,tew.,
Osmonncza
Pac.rw,:t£ma mYJU&gt;-Ln-Ue6
PecU.c.u.e.cvw, gJtay-L
Pern&gt;-temon c.Jtan cia..e..eU
PeM.temon ~bUc..tU6
Phac.e.e-La he.teJtophyLea
Ph.eox .eong-L6oUa
Po.eemoYiA.um 6oUo.6-W.6-i.mum
Po.eygonum av-Lc.u.tMe
Po.eygonum dougla.6-i.-i.
Po.tentiUa
pu.tc.heJUWna
Ranunc.u.tw., -Lnamoenw.,
Rumex c.Jt-i..6pw.,
Rudbec.k-i.a montana
Senec.-i.o ambJtO.6-Lo-Ldu
Senec.-i.o -Ln.tegeJUWnU6
S enec.-i.o .6 eJtJta
SUene menUe6-i.-i.
SmUac.-i.na .6 .teUata.
SoUdago .6 paJU&gt;-L6.eoJta
S.teU.a.tl..La
SWeJL.tLa Jtadia.ta
TaJtaxac.um 06 6-Lc.-i.na.ee
ThaUc.:tJtum 6 end.evu.

0.00
0.00
0.00
0.95
0.28
0.00
0.19
0.03
0.00
0.00
0.00
0.00
0.00
0.21
0.00
0.00
0.12
0.00
0.00
0.00
0.00
0.06
0.11
0.80
0.00
0.17
1. 95

0.00
0.00
0.00
0.87
0.09
0.02
0.04
0.02
0.00
0;00
0.04
0.00
0.00
0.27
0.00
0.00
0.05
0.00
0.01
0.06
0.00
0.05
0.05
1.73
0.00
0.44
1.59

0.00
0.06
0.00
- 2.32
- 3.57
+ 0.30
- 2.44
- 0.24
0.00
0.00
0.48
0.00
0.06
0.65
0.00
0.00
- 1.31
0.00
+ 0.12
+ 0.89
0.00
- 0.18
- 1.07
+13.86
+ .06
+ 3.81
- 7.79

- 14.44
- 73.17
- 78.85
I

-I'-

00
I

+ 18.97
- 64.71

+103.10
+133.33
- 23.82

-----------------------------------------------------------------------------------------------------------------------------------------

�Table 2. Production of air dry summer vegetation recorded in the control areas before and two years following burning,
spraying and chaining (contd).
Lbs of vegetation
produced per acre

Percent composition
Plant species

1970

1970

1973

.

1973

1/
Change 1970 to 1973 lbs/ac
percent fI

0.30
0.00
1.25
2.44
0.06
0.00
3.39
2.02
5.65
2.14

+ 0.30
0.00
0.00
+ 0.12
+ 0.06
0.00
- 0.48
+ 1.01
+ 2.74
- 0.12

Forbs (contd)
T4agopogon dubium
TM.gopogon pJta.ten6M
UJr-uca dcocca
VaJ..eJtiana ocudentctU6
VeAonica ameJticana
Ve/tonica bilo ba
Vig u,,{.eJtamu1ti6R.oJta
V,tofu JtUg/LtOI.&gt;
a
V,to.e.aI.&gt;heUonU
Wlje..tJua amp.e.eucauU'.&gt;

Total Forbs

0.00
0.00
0.08
0.14
0.00
0.00
0.22
0.06
0.18
0.14

0.02
0.00
0.08
0.16
0.00
0.00
0.22
0.14
0.37
0.1'4

0.00
0.00
3.87
1.01
2.91
2.26

--

--

31.86

30.03

545.35

478.50

-66.85

- 12.26

84.52
0.00
0.00
36.58
0.18
0.59
0.00
0.00
16.18
0.00
0.06

35.27
0.00
·0.00
2u.40
3.81
0.00
0.00
0.00
4.10
0.00
0.36

-49.25
0.00
0.00
-16.18
+ 3.63
- 0.59
0.00
0.00
-12.07

- 58.27

0.00
0.00
1.25

2.32

+

5.13

- 12.31
+100.00
+ 93.88
5.26

-

Grasses
AgJt°pYJton
AgJtopYJton CJtiI.&gt;tatum
AgJtopyltOll intvunecUum
BJtomUl.&gt;anomaJ..UI.&gt;
BJtomUl.&gt;,tneJtmi!.&gt;
Bltomu-s te.ctOltum
CMex
Vacty.e.U6 g.e.omeJtata
Ef.ym UI.&gt;9 fuuCUl.&gt;
FeI.&gt;tuca oV,tna
FeI.&gt;tuca thUltbeJti

4.85
0.00
0.00
2.15
0.01
0.04
0.00
0.00
1.00
0.00
0.00

2; 17
0.00
0.00
1. 31

0.23
0.00
0.00
0.00
0.28
0.00
0.02

- 44.23

- 74.63

0.00
+ 0.30

-----------------------------------------------------------------------------------------------------------------------------------------,..

I

.".
\!)

I

�Table 2. Production of air dry summer vegetation recorded in the control areas before and two years following burning,
spraying and chaining (contd).

Plant species

Percent com~osition
1970
1973

Lbs of vegetation
~roduced ~er acre
1970
1973

Change 1970 to 1973 l!
16s/ac
percent ~

Grasses (contd)
Ho~deum b~aehyanth~um
JuneU6 ba.lUeU6
Koe..e.vu:.a eJUJ.,.taJ:.a.
O~ZUOP-6-i/.,hymeno..i.du
Phte.wn pJz.a.ten.oe
Poa-C~ex geyvu:.
SLtan..i.on hy-6bUx
Supa

0.25
0.01
0.01
0.00
0.00
12.74
0.00
0.97

1.00
0.00
0.05
0.00
0.00
13.52
0.00
0.91

4.10
0.12
0.24
0.00
0.00
221.61
0.00
16.06

14.87
0.00
0.89
0.00
0.00
218.64
0.00
14.45

+10.77
- 0.12
+ 0.65
0.00
0.00
2.97
0.00
- 1.61

-

+262.32

- 1.34
- 10.00

I
IJ1

0

Total Grasses

22.04

I

19.49

380.24

312.79

0.00
123.95
1.07
7.20
0.95
0.00
0.30
2.08
0.00
0.00
103.91
1.78

0.00
115.39
0.48
9.16
1.67
0.00
0.12
3.75
0.00
0.00
92.67
2.91

- 67.45

- 17.74

Shrubs
Ae~ g.e.ab~um
AmUa.nc.h..i.~ a.tn..i.6oUa
,."-t(:J~..i.a c.ana
Mte~..i.a bUden-ta.ta
C~eOc.~pU6 montanU6
CMY-6othamnU6 de~U-6U6
CMY-6othamnU6 naU6W6 U6
CMY-6othamnU6 v..i.-6c...i.d[6.e.O~U6
CJz.a.taegU6 ~ytMopoda
PoputU6 ~emuto..i.du
P~unU6 v..i.kg..i.n..i.ana
PulL6 h..i.a bUde.n:ta.ta

0.00
7.27
0.06
0.40
0.06
0.00
0.02
0.12
0.00
0.00
5.94
0.11

0.00
7.13
0.03
0.53
0.11
0.00
0.01
0.22
0.00
0.00
5.58
0.20

0.00

- 8.56
0.59

+ 1.96
+ 0.71
0.00
0.18
+ 1.67
0.00
0.00
- 11.24
+ 1.13

-

- 6.91
+ 27.27

+ 80.00
- 10.82
+ 63.33

----------------------------------------------------------------------------------------------------------------------------------------

�Table 2. Production of air dry surrmer vegetation recorded in the control areas before and two years folIowinq burning,
spraying and chaining (contd),

Plant species

Percent com~osition
1970
1973

Lbs of vegetation
produced per acre
,970
i973

l/
Change 1970 to 1973 iEs/ac
percent ~

Shrubs (contd)
QUVl.CU6 gambeUM.
RA:.bIU
R0.6a. Itutkana.
SymrhoJUcCUtpo.6
a1.bU6

Total Shrubs
TOTAL ALL PLANTS

17.92
0.43
1.35
12.44

18.75
0.40
2.01
15.51

- 6.13

308.57
6.96
23.43
213.35

302.44
6.13
32.00
248.62

- 0.83

+ 8.56
+ 35.27

- 1.99
- 11.97
+ 36.55
+ 16.53

--

--

46.10

--

50.48

793.55

815.32

+ 21.77

+

100;00

1719.14

1606.61

-112.53

- 6.55

--

100.00

1/

- Reflects normal year to year changes in forage production recorded on the control areas.
2/

- Percent changes are only shown for the more abundant plants.

2.74

I
Vl

•...
I

�Table 3. Changes in production of air dry summer vegetation recorded on the burned areas two years following burning.
Percent com~osition
Plant species

1970

1973

E" 73 11

Lbs of vegetation
Qroduced Qer acre
1970

1973

E 73

1bs/ac

111.82
0.30
0.95
8.80
0.00
0.00
0.00
0.00
4.28
0.00
0.18
3.81
105.33
0.00
0.48
0.00
1.07
0.06
1.67
8.92
8.15
0.24
0.30
2.02
0.06
0.00
0.36
0.00

92.84
0.05
21.23
18.27
0.00
0.00
0.03
0.10
4.40
0.00
1.78
2.96
77.18
0.00
0.24
0.00
1.28
0.59
4.32
0.20
22.84
0.12
1.80
1.19
0.01
0.00
1.42
0.00

+ 18.97
+ 0.24

Change due to burning fj
Confidence level
percent 1/ where significant if

Forbs
AcJUllea. fanu£o.o a.
Agcv~.ta.che {).)(;tiun0-Uo.
Ag 0.0vrM 9 fauca.
Allium a.cum-i.ncU:um
And!to.oa.ce .0 e.pte.rlbUo na,.U;.,
Aqu-i..te.g-i.a cOe.ku.tea.
Alta.bb., dJr.ummoI1fU
Alta.bb., hJ.M u..ta.
AltencvUa. conglUta.
MtemM-i.a. b-i.e.nnM
A'Lte.mM-i.a dJr.a.cunCu.tU6
Mte.mM-i.a fudov-i.uana.
Mte.k - EJUgVlon
M.tIta.ga..f.U6 ha.yden-LanU6
Ba.£6amoJrJUza -6agU.ta..ta.
Be.kbvrM Jz.epen.o
Ca..f.ocholL.tu6 gunnMonil
Cap.6eUa. bwv.,a-pa..otoJz.M
Ca..otiUeja unaJt.i.a.e60-Uo.
Chenopod,(.um bwand,(.e.Jz.,(.
C,(.Jz.;.,-i.um
centauJz.ea.e
Co.t.e.om-i.auneaJz.£.6
CommandJr.a.umbe.t.e.a.ta.
Cymopte.kU6 puJz.puJz.e.U6
Ve.f.phbuum nwon-i.
Ve.f.ph,Ln-i.um 0cc,(.den.ta..te
VIUc.uJz.a.-i.n-i.a.6 0plUa.
Vb.,poJz.um .tJz.achyca.Jz.pum

2.57
0.00
0.02
0.17
0.00
0.00
0.03
0.02
0.18
0.00
0.11
0.21
6.50
0.00
0.01
0.00
0.06
0.04
0.34
0.01
1.05
0.00
0.07
0.05
0.00
0.00
0.01
0.00

8.6~
0.02
0.07
0.68
0.00
0.00
0.00
0.00
0.33
0.00
0.01
0.30
8.14
0.00
0.04
0.00
0.08
0.00
0.13
0.69
0.63
0.02
0.02
0.16
0.00
0.00
0.03
0.00

5.96

5.00

42.88
0.06
0.36
2.85
0.00
0.00
0.48
0.30
2.97
0.00
1.78
3.45
107.95
0.00
0.24
0.00
1.07
0.59
5.59
0.24
17.31
0.00
1..19
0.89
0.00
0.00
0.12
0.00

+ 20.44

- 20.27

9.47
- 0.00

- 51.82

.10

0.00

- 0.03

- 0.10

0.12
- 0.00
- 1.61
+ 0.85
+ 28.15

0.00
0.24
0.00
0.21
- 0.54
- 2.65
+ 8.72
- 14.69
+ 0.12
- 1.51
+ 0.83
+ 0.05
0.00
1.06
- 0.00

- 2.70
- 90.00
+ 28.74
+ 36.48

+

-

- 16.67
- 61.45
- 64.33
- 83.51

----------------------------------------------------------------------------------------------~--------------------------------------------

I
Vl

'"
I

�Table 3.

Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 7311

1970

1973

'"

.

E 73

lbs/ac

16.47
11.67
0.12
9.89
0.00
0.70
0.00
10.05
0.00
0.37
0.00
0.09
18.85
0.83
0.00
31.65
0.00
1.51
8.19
0.00
77.58
0.00
0.01
0.00
0.01
0.00
0.01

- 14.09
9.24
0.06
+ 5.16
0.00
+ 1.08
0.00
7.31
0.00
0.19
0.00
0.03
- 11.42
0.83
0.00
+ 4.75
0.00
0.38
5.87
0.00
+ 0.93
0.00
+ 0.29
0.00
+ 0.59
0.00
+ 0.11

Change due to burning ~
Confidence level
percent Y where significant ~

Forbs (contd)
EM.og0 nwn wnbeil.a..twn
Epltob~wn pa~cutatum
FJtO.gcvUaameJUcana
Giliwn
Gel1..uana he;tvLO,~epa£a
GeJtwuwn
GULa aggltega:ta.
G~ndet£a aphana~
HacRet£a ieptophylla
Hue~wn hoopruU
HydltophyUwn capUa.twn
HydltophyUwn 6enct.eeJU
Iw mLMouJUeVl.6,u,
Lactuca .6CcvUoia
Lappuia ltedoLUQ
W
uuhipius - V~ua
LepMiwn deVl.6iMOJtwn
UgU6.ti..cwn
Unwn ie.w.i.6U
Loma.ti..wnleptocaltpwn
Lup~nU6 altgenteU6
Mad.i.a glomeJta.ta
Med.i.cago ~a.ti..va
~!Uilo;(:U6 o6Muna£.w
MeJtteVl.6~a olta.YLWcana
MeM:en6ia funceota:ta.
Moldavica paltvl6lolta

0.37
0.01
0.01
0.68
0.00
0.09
0.00
0.50
0.00
0.06
0.00
0.01
1. 12
0.05
0.00
2.20
0.00
0.12
0.54
0.00
8.13
0.00
0.00
0.00
0.00
0.00
0.00

0.18
0.19 .
0.00
1.17
0.00
0.14
0.00
0.21
0.00
0.01
0.00
0.01
0.58
0.00
0.00
2.81
0.00
0.09
0.18
0.00
6.04
0.00
0.02
0.00
0.05
0.00
0.01

2.04

4.98

6.07
0.18
0.12
11.42
0.00
1.55
0.00
8.21
0.00
1.01
0.00
0.12
18.85
0.83
0.00
36.52
0.00
1.96
8.92
0.00
134.54
0.00
0.00
0.00
0.00
0.00
0.00

2.38
2.44
0.06
15.05
0.00
1.78
0.00
2.74
0.00
0.18
0.00
0.12
7.43
0.00
0.00
36.40
0.00
1.13
2.32
0.00
78.51
0.00
0.30
0.00
0.59
0.00
0.12

-

-

-

--

- 85.55
+ 52.19
+154.81
- 72.77
I

'"

""I

- 60.57
+ 15.01
- 24.99
- 71. 68
+

.25

1.20

-_._-----------------------------------------------------------------------------------------------------------------------------"-----------------

�Table 3. Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
eroduced eer acre

Percent comeosition
Plant species

1970

1973

E 7311

1973

E 73

1bs/ac

0.00
2.14
0.00
0.18
0.00
0.00
2.80
1.31
0.00
4.22
0.06
0.06
0.00
5.95
0.00
0.48
1.49
0.30
1.19
0.18
0.00
0.06
1.43
28.01
0.00
23.02
2.02

0.00
33.61
0.30
1.53
0.00

0;00
- 31.47
0.30
- 1.35
- 0.00
0.00
+ 1.94
0.28
-+ 0.30
4.21
0.03
- 1.72
7.43
1.84
0.00
+ 0.36
+ 0.46
+ 0.29
+ 1.07
0.71
0.00
+ 0.05
- 0.39
+ 1.68
0.00
- 15.70
- 0.52

1970

Change due to burning Y .._____
Confidence level
percent '§j where significant if

Forbs (contd)
Oeno~heka cae6pito6a
Opu.n.tia
Olt.-thoC.MpU6 .tu:teU6
O~mottiuza
Pac.l~:t£ma myMinLte6
Pedic.ularJ~ gnay~
Pen6~emon ~nda.ttU
Pen6~emon ~:tJUdU6
Phac.e£ia het~ophy£la
Pheo x to ng.{·6o-f-&lt;.a
Potemol'l-&lt;'um6oUo~~-6~um
Potygonum av~c.ulafr.e
Potygonum dougta-6~
Po~e~
putc.h~a
Ranunc.utU6 ~namoenU6
Rumex. c.JW, pU6
Rudbec.kla mo~ana
Sene~o amb~o~~o~de6
Sene~o ~~eg~U6
Sene~o
-6~a
SUene menz~e6~
SmUaUna -6 ~e.Ua:ta
SoUdago -6PaM~MOM

s~eU~
SweM:.{a ~a~
TMaxac.um o6M~Y!a.ee
ThaU.cbtum 6end.e~

0.00
0.34
0.02
0.11
0.00
0.00
0.25
0.17
0.02
0.00
0.00
0.11
0.07
0.39
0.00
0.01
0.17
0.00
0.00
0.00
0.00
0.00
0.25
0.78
0.00
0.99
0.20

0.00
0.16
0.00
0.01
0.00
0.00
0.21
0.10
0.00
0.32
0.00
0.00
0.00
0.46
0.00
0.04
0.12
0.02
0.09
0.01
0.00
0.00
0.11
2.16
0.00
1.79
0.16

0.00
5.65
0.30
1.78
0.00
0.00
4.04
2.85
0.30
0.00
0.00
1.78
1.25
6.54
0.00
0.12
2.91
0.00
0.00
0.00
0.00
0.00
4.16
12.97
0.00
16.59
3.33

O,OJ

0.86
1.59
0.30
0.01
0.09
1.78
7.43
7.78
0.00
0.12
1.03
0.01
0.12
0.89
0.00
0.01
1.82
26.33
0.00
38.72
2.54

-

- 93.63

-

+226.74
- 17.50

--

.05

I
\Jl

~
I

- 23.58
+ 44.56

-

- 21.63
+ 6.38
- 40.55
- 20.30

----------------------------------------------------------~------------------------------------------------------------------------------------

�Table 3.

Changes in production

of air dry summer vegetation

recorded on the burned areas two years following burning (contd).
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

Tltagopogoll dub-i.um
TltagOpOgOH plULteno-W
Wz-Uea cU.o-i.ea
VatvUana. oeudelU:aLW
Vvwlu.ea amV'vteana
VvwvU.ea b..Le.oba
V-i.gu.i.eIta muLU6folta
V-i.o.ea Itug utO.6a
V-i.o.ea .6 he.Uo vU.-L
Wye.th-i.a a.mp.f.ex..i.eaut-i..6
Tota 1 Forbs

E 73 Jj

Change due to burni ng Y . _____
.Confidence level
percent l! where significant if

1970

1973

0.02
0.00
0.00
0.02
0.00
0.02
0.34
0.04
0.27
0.14

0.06
0.00
0.00
0.03
0.01
0.06
0.15
0.39
0.30
0.25

Forbs (contd)
0.30
0.06
0.00
0.36
0.00
0.30
5.59
0.71
4.46
2.32

--

--

--

--

--

30.09

38.76

499.31

501.57

438.10

+ 63.47

+ 14.49

30.27
0.06
12.07
5.35
2.91
0.83
0.00
0.30
1.43
0.00
0.00

29.21
0.01
0.01
5.57
0.21
0.07
1.61
0.01
0.00
0.00
49.96

+ 1.06
+ 0.05
+ 12.06
0.22
+ 2.70
+ 0.76
1.61
+ 0.29
+ 1.42
0.00
- 49.96

+

1970

1973

E73

lbs/ac

0.83
0.06
0.00
0.36
0.18
0.77
l. 96
5.12
3.87
:3.21

8.84
0.06
0.00
0.38
0.06
0.30
4.90
l.43
8.65
2.20

- 8.01

-

+

-

+

-

+

0.00
0.00
0.02
0.12
0.48
·2.94
3.69
4.78
1. 01

- 59.97
- 55.30
+ 46.15

Grasses
AgltopyltOYl
AgltopyltOYl ~tatum
AgltopyltOYl -i.ntvvnecU.um
BltOmM anomatM
B!WmM -i.Ylvvn-W
BItOmM .:tee-toltum
CMex

Vae.:ty.e.U.6g.i:.omelta..:ta.
UymM 9 .f.aUC.M
Fu.:tuc.a OV-Lrla.
Fu.:tuc.a .:thultbvU
------------------------------

4.22
0.00
0.00
0.60
0.00
0.26
0.10
0.00
0.00
0.00
0.50
•. -------------------------

2.34
0.00
0.93
0.41
0.22
0.06
0.00
0.02
0.11
0.00
0.00

l.89

70.01
0.00
0.00
9.99
0.00
4.34
1.61
0.00
0.00
0.00
8.33
_________________________

-

3.63

- 3.95

-

e. ___________________________________________________________

I

'"'
'"'
I

�Table 3. Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
produced per acre

Percent composition
Plant species

HOIJ.deum bJUJ..chyantheJWm
] unc.Ui&gt; ba..Ui.c.UI.&gt;
Koe.lvUa
vW,,-tCLta
OIJ.zyoy.u,,u, hymeYto..Lde6
Pheewn pIJ.CLteiU e
Poa-CMex
geyvU
SUaiUoil
hY-6vuX
Stipa

Total Grasses

1970

1973

E 7311

1973

E 73

0.18
0.11
0.28
0.00
0.02
15.65
0.19
0.89

00.03
0.13
0.09
0.00
0.03
17.72
0.03
0.89

Grasses (contd~
3.03
0.36
1.90
1.67
1.13
4.64
0.00
0.00
0.36
0.42
16.60
259.98 229.23
3.15
0.36
14.87
11.54

10.99
0.16
17.40
0.00
0.36
256.49
3.15
13.38

1970

23.01

23.03

382.20

-- --

1bs/ac

Change d ue to burnt
urnolng -2/
3/ Cot'lrfClence
1eve 1
percent - where significant iI

- 1O.63

- 96.75

- 16.27
0.00
+ 0.06
- 27.26
2.80
- 1.84

- 93.50

+ 1.51

-

.15

- 10.63
- 13.78
I

VI
0\

297.92

314.41

- 16.48

0.00
41.57
10.53
12.55
0.12
1.07
15.23
2.74
0.71
0.00
9.87
0.00

0.00
52.82
4.23
165.93
0.62
0.01
0.40
27.30
0.12
0.00
10.61
0.00

0.00
- 11.25
+ 6.30
-153.38
0.51
+ 1.06
+ 14.82
- 24.56
+ 0.59
0.00
- 0.74
0.00

- 5.24

Shrubs
AceIJ. g.f.abIJ.um
Ame.f.ancJUe/J.
a.f.iU6oUa.

A!i..tem,0.,.i.a carta
A!i..tem,0.,.i.a vudeYL.ta.ta.
CelJ.COCMPUl.&gt; moYttaYtU6
ChlLY-6othamYtU6 depILe6-6 U6
ChIJ.Y-6othamYtU6 !1aU6 1'.0-6UI.&gt;
ChIJ.Y-6othamYtUl.&gt; vcs c..i.d.i.6R..OIJ.U6
Cli.a.tae.gu6 e/J.y-thILopoda
PopuiU6 .tIJ.emuio..Lde6
PIJ.u.YtU6v.i.lJ.g.i.iUa.Yta
PUM h.i.a vude.YttCLta

0.00
3.45
0.58
7.90
0.02
0.00
0.06
0.92
0.01
0.00
0.71
0.00

0.00
3.19
0.81
0.96
0.01
0.08
1.17
0.21
0.06
0.00
0.77
0.00

3.39

0.00
56.74
9.52
130.37
0.36
0.00
1.01
.15.17
0.12
0.00
11.90
0.00

-

- 21.29
+148.91
- 92.44

- 89.98

- 6.93

-----------------------------------------------------------------------------------------------------------------------------------------

I

�Table 3. Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
Qroduced Qer acre

Percent comQosition

1973

g 73

lbs/ac

Shrubs {contd}
174.98 190.15
14.10
40.39
2.60
29.20
17.84
23.11
308.63 177.90

171 .51
35.55
39.88
359.65

+ 18.64

+ 10.87

- 21.46
- 22.03
-181. 75

- 60.35
- 55.25
- 50.54

Plant species

1970

1973

~ 73 Jj

gambeUU

10.56
2.41
1.74
18.54

14;70
1.10
1.38
13.78

11.06

--

--

46.90

38.21

QUeJtCU6

TUbe;.,

RO-6a ntd:hana
~!JmphoJUc.aJl.po-6 a£bU6

Total Shrubs

Change due to burning Y
Confidence level
percent '}f where significant iI

1970

.10

-778.38

494.38

799.73

-305.36

- 38.18

.05
I
Vl
-..J

TOTAL ALL PLANTS

100.00

100.00

1659.90

1293.87

1551.24

-257.37

- 16.59

lfReflects the amount of forage one would expect to find on a treated area following treatment had treatment not occurred.
YOerived through comparison of burned and control areas with adjustments for normal year to year changes in forage production recorded on
the control areas.
'}fPercent changes are only shown for the more abundant plants.
illndicates the highest confidence level at which changes due to burning are significant.
a .05, .10, .15, .20 or .25 level are shown.

Only changes that are significant at the

I

�Table 4. Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying.
Lbs of vegetation
2roduced Eer acre

Percent comEosition
Plant species

1970

1973

E 73 ]j

Change due to s2ra~in9 g(
Confidence level
percent 11
where significant ~

1970

1973

E 73

1bs/ac

25.69
0.18
0.06
0.48
2.20
0.00
0.06
0.06
0.24
'0.00
7.61
0.65
61.44
0.00
0.00
0.18
1.61
0.00
4.40
0.12
14.99
0.00.
0.00
0.00
0.00
17.66
0.18
0.18

104.86
0.48
3.15
1.01
0.00
0.95
0.00
0.00
3.21
0.00
0.18
1.90
47.34
0.65
1.13
0.06
1.19
0.00
0.83
0.00
29.86
9.99
1.78
0.24
0.06
20.58
2.26
0.12

55.63
0.16
3.54
3.05
0.12
0.01
0.00
0.02
0.35
0.00
7.61
0.56
43.93
0.01
0.01
0.24
1.93
O.CO
3.40
0.10
19.78
0.12
0.02
0.01
0.01
27.26
2.12
0.18

+ 49.23
+ 0.31

+ 88.50

- 2.03

- 10.89
- 66.80

~
Ac.hde.ea .tal1u.£.0-6
a !&lt;AgM.tacJte U/(M.uooua
Ago-6eJtM g.tauc.a
AWwn. ac.um.tHa..twn
AvtMO-6ac.e -6ep.tentJUoi1a..U-6
Aquileg-ia co C/Llilea
A'Ul.bi..-~ Mummo11cU.
AM.b~ hhz.6lU:a
A!z.el1cvz..{.a co I1g e6.ta
AI[,t(')n~-ia b.LemU-6
Al[,tcnl~-ia dttacuncu.£.U6
A.Ucnl~-ia ludov.tc..&lt;.avta
A-6.teA-EJUgvwvt
A.~t!z.aga£U6 haydel1.tMU6
BaLI, amoI!.fUz a -6 agi..-Ua..ta
BC/Lbew Jr.epe!1.-6
ca£OC.hol[,tU6 gUl1wovti..-[
Cap-6e.t.e.a bUMa-pa..6.toW
Ca..6.tL.t.e.eja UvtcvUae6o.e..&lt;.a
ChevtopocU.um beAlal1cU.vU
c.{Jr..~.twnc.elU:auJr.eae
CoUom.i.a UnecvU.6
Comandlta. umbC-Ua..ta.
CUmOp.tC/Llv" puJr.puJr.e.U6

Velplul1.tum vtWOM
Velph ..i.n-tW11oc.udenta£e
Vc.~C.c.Ula..{.l1.{a .~0ph.ta
V~pOJr.um w:.c.lU/C.CVlPW1

1.56
0.01
0.00
0.03
0.13
0.00
0.00
0.00
0.01
0.00
0.46
0,04
3.70
0.00
0.00
0.01
0.10
0.00
0.26
0.01
0.92
0.00
0.00
0.00
0.00
1.08
O.01
0.01

4.15
0.02
0.12
0.05
0.00
0.05
0.00
O~OO
0.14
0.00
0.01
0.07
2.00
0.02
0.06
0.00
0.05
0.00
0.04
0.00
1.10
0.40
0.08
0.01
0.00
0.76
0.10
0.01

3.61

2.85

- 0.39
- 0.12
+ 0.94
0.00

- 0.02

+

2.86
0.00
7.43
+ 1.34
+ 3.42
+ 0.64
+ 1.12
- 0.18
- 0.74
0;00
- 2.57
- 0.10
+ 10.08
+ 9.87
+ 1.77
+ 0.22
+ 0.05
- 6.68
+ 0.14
- 0.06

.20

00
I

-

+

7.78

-38.27

+ 50.95

- 24.50

----------------------------~-------------------------------------------------------------.--------------------------------------------------------..

I
V1

�Table 4.

Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying
Lbs of vegetation
Qroduced Qer acre

Percent com~osition
Plant species

1970

1973

E 73 ]j

1970

1973

E 73

0.19
0.00
0.00
59.12
0.30
3.87
0.00
1.84
0.59
0.00
1.84
3.33
0.00
0.06
1.90
34.68
0.77
8.74
1.25
0.00
67.39
0.00
0.00
0.06
4.76
0.95
0.00

0.81
0.00
0.18
20.55
0.59
2.80
3.54
1. 38
0.00
0.02
0.12
0.52
0.00
0.01
0.48
87.37
0.12
13.56
2.46
0.00
104.34
0.03
0.00
0.01
6.18
0.01
0.00

(contd).

y

1bs/ac

Change due to sQra~ing
.
Confidence level
percent '}j
where significant i!

Forbs (contd)
Wogonwn wnbde.a.twn
Ep~ob~wn pa~cutatum
FMga/ua amvUco.na
GaLLwn
'Ge.n.tiana he.:teJto.6
e.pa.ta
GeJta~wn
G~
aggl!.e.ga..ta.
Gkinde.t£a aphanactl6
Hacketia le.ptophytea
He..e.e.~wnhoopuil
Hljd/tophytewn cap~wn
Hyd/tophytewn 6e.ndlvU
Iw mM.60wUe.n.6M
Lac..tu.ca .6 ca/Uola
Lappula ke.dow.6kil
La.thYkU6- (!~Ua
Le.p~~wn de.n.6~6lokwn
UgU6Ucwn
u.nwn l~il
LomaUwn le.ptocakPwn
LUP~f1U6akge.nte.U6
Ma~ g.i'.omeJta..ta.
Me.~c.ago .6a.tiva
M~O.tU6 o6Mun~
MeJt.te.n.6~a6Mnw cana
MeJt.te.n.6ialance.ola..ta.
Moldav~ca pakvi6l0M

0.02
0.00
0.01
1.44
0.01
0.38
0.02
0.07
0.01
0.00
0.00
0.04
0.00
0.00
0.03
6.08
0.00
1.06
0.16
0.00
10.85
0.04
0.00
0.00
0.41
0.00
0.00

0:06
0.00
0.00
2.35
0.02
0.15
0.00
0.07
0.02
0.00
0.07
0.17
0.00
0.00
0.07
1.40
0.03
0.37
0.06
0.00
3.16
0.00
0.00
0.00
0.24
0.04
0.00

1.35

5.63
0.88
6.64

0.30
0.00
0.18
23.73
0.24
6.19
0.30
1.13
0.12
0.06
0.00
0.71
0.00
0.00
0.48
100.81
0.00
17.66
2.68
0.00
180.93
.0.59
0.00
0.00
6.90
0.00
0.00

+

0.38
0.00
0.18
+ 38.57
0.30
+ 1.06
3.54
+ 0.46
+ 0.59
0.02
+ 1.72
+ 2.81
0.00
+ 0.05
+ 1.43
- 52.69
+ 0.65
4.82
1.21
0.00
- 36.95
0.03
0.00
+ 0.05
1.42
0.94
0.00

-

-

+187.73

.15

+ 38.02
+ 33.28

I
V1
\0
I

-

-

-

- 60.31

.10

- 35.52
- 49.17

.15

- 35.41

- 22.99

-------------------------------------------------------------------------~-------------------------------------------------------------------

�Table 4. Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying (contd).
Lbs of vegetation
Eroduced Qer acre

Percent comEosition
Plant species

1970

1973

E 73 Jj

1973

E 73

1bs/ac

0.00
0.00
0.00

0.00
0.00
0.00

4.58
5.29
2.32
1.49
3.15

0.00
0.00
0.00

8.33
3.15

0.00
0.00
0.00

3.92
1.42
68.98

4.41
1.73
- 68.98
+ 0.16

1970

Change due to sQrating ~
onfidence level
percent ]I
where significant 11

Forbs (contd)
Oen.o.theJto cae.6 pU0.6 a
OPluttia.
O,'t.thocaJtpw, Me.w,
0-6moJrJuza
pachM.tUna myM.&lt;.nLte.6
Pe.cUc.U£M.&lt;.-6gJtay.&lt;.
Pe.YL6.te.mon c.Jtanda.te..U
Pe.I1.-6.te.mOH -6vuc.tw,
Phac.e..Ua he..tvwphyUa
Phiox iong'&lt;'6ot.&lt;.a.
Po.te.mon.&lt;.um 6o.e..&lt;.o-6.&lt;.-6I.&gt;.&lt;.mum
Poiygonum av.&lt;.cU£aJte.
Po.tygonwl1 dOug.tMU
Pote.n.t.&lt;.t.e.a pU£ch~a
RMWlC.U£W, .{.vlamoe.nw,
Rume.x eM..6 pw,
Rudbe.clUa montana
Se.ne.c.&lt;.o ambkO-6.&lt;.o.&lt;.de.6
Se.ne.c.&lt;.o .&lt;.n.te.gVl/UmU/.)
S en e.c.&lt;.o I.&gt; eJtJta
SUe.ne. me.nue.6U
Sm.&lt;..tac.&lt;.na-6.te.Ua..ta.
SoUdago -6paM'&lt;'6.f.olta
S.te.UM.&lt;.a
SweJtUa JtacUa..ta
Tal!.axacum o66.&lt;.c.&lt;.nale.
ThaUc..tkum· 6e.ndleJt.&lt;.

0.00
0.00
0.00

0.28
0.31
0.14
0.09
0.19
0.00
0.00
0.02
0.00
0.02
0.62
0.00
0.00

0.10
0.02
0.00
0.00
0.00

0.01
0.10
0.76
0.18

0.41
1.85

0.00
0.00
0.00

0.32
0.16
0.00
0.02
0.04
0.00
0.02
0.13
0.01
O. Ol
0.10
0.00
0.00

0.00
0.00
0.30
0.00
0.30
10.11
0.06
0.00

0.12
0.01
0.00
0.07
0.00
0.03

0.14
1.65
0.16
1.70
0.86

1.51

0.00
0.48

1.75

0.00
0.36

0.00
0.01

3.21

2.68

0.36
0.18

0.01

2.68

12.03

0.00
0.00

0.06

1.67
0.36

2.91
0.24

0.00
0.00
0.00

0.06

0,18
1.67
12.55
2.91
6.72
30.63

0.31

1.13

1.37

0.00
0.54

2.80
40.80

4.16
45.56
21.65

1.77
0.00
0.59
0.25

0.12
0.89

- 0.62

+
+
+

+112.51
+121.96
- 35.47

I

0.00
0.35
0.54
0.35

1.59
9.35
-- 0.06
- 0.00

+ 2.33
- 0.12
-+ 0.06
0.48

0.00

0.00

0.15
0.73
25.49
17.33
15.68
23.34

+ 0.39
+ 2.07
+ 15.31
- 13.17
+ 29.88
- 1.69

'"

0
I

- 77.75
+395.83

+283.67
+ 60.08
- 75.98

.15

+190.52

- 7.22

----------------------~-----------------------------------------------------------------------------------------------------------------------

�Table 4.

Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying
Lbs of vegetation
Qroduced Qer acre

Percent comQosition
Plant species

1970

1973

E 73 ]j

1970

1973

E 73

Ibs/ac

(contd).

Change due to sQra~ing fI
Confidence level
percent '}j
where significant ~

Forbs (contd)
Tkagopogon dubIum
Thagopogon pnateno~
Uwea
cUo.iea
Va.ivUana. oecJ.de~
Vruw Meet o.mvUeana.
VVWMea bUoba.
VJ.guJ.eha mu.Ui.6.toll.a
V.io.e.a.lLugu.toJ.&gt;a
V.io.e.a J.&gt;hUton,u
Wye.-tlua o.mp.e.exJ.ea~

Total Forbs

0.06
0.00
0.08
0.50
0.00
0.00
0.00
0.33
0.25
0.77

0.04
0.00
0.33
0.64
0.00
0.00
0.00
0.38
0.22
0.11

0.95
0.00
1.43
8.27
0.00
0.00
0.06
5.53
4.04
12.67

1.07
0.00
6.54
14.27
0.00
0.00
0.06
8.86
5.29
2.85

28.30
0.00
1.43
8.69
0.00
0.00
0.05
11.06
7.84
12.00

- 27.23
0.00
+ 5.12
+ 5.58
0.00
0.00
+ 0.01
- 2.20
2.55
9.15

-

-

- 19.89
- 32.49
- 76.21

36.07

24.82

598.22

603.40

524.89

+ 78.51

+ 14.96

47.52
0.00
0.48
32.53
0.00
5.71
0.00
9.69
28.97
0.00
0.00

5.98
0.00
0.01
4.31
0.00
0.00
0.00
1.61
1. 74
0.00
0.00

+ 41.54
0.00
+ 0.47
+ 28.22
0.00
+ 5.71
0.00
+ 8.09
+ 27.23
0.00
0.00

+694.45

+358 ..33
+ 64.24
.05
I

....
'"
I

.15

Grasses
Agll.OpY'wn
Agll.0pYll.on Cll..iJ.&gt;ta.tum
Agll.opYll.on .inteJr.mecUum
BlZ.omUJ.&gt;
anoma..tM
BlZ.omUJ.&gt;
.in~
BlZ.omUJ.&gt;
tectolZ.um
CMex
Vacty.e.UJ.&gt;g.e.om~
UymUJ.&gt;g.e.aueUJ.&gt;
Futuea
ov.ina
Futuca
thulZ.beJz.J.

0.86
0.00
0.00
0.47
0.00
0.00
0.00
0.10
0.41
0.00
0.00

2.06
0.00
0.02
1. 33
0.00
0.25
0.00
0.36
1.39
0.00
0.00

14.33
0.00
0.00
7.73
0.00
0.00
0.00
1.61
6.84
0.00
0.00

+654.44
.05
+503.70
+1569.36

------------------------------------------------------------------------------------------------------------------------------------------~---

--

..

�Table 4. Changes in production of air dry summer vegetation recorded on the sprayed areas two years fo11owing spraying (contd).
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 73 ]j

1973

E 73

1bs/ac

0.00
0.00
4.94
0.00
2.91
719.50
0.00
28.61

0.00
0.00
3.12
0.00
0.36
230.73
0.06
4.71

0.00
0.00
+ 1.81
0.00
+ 2.56
+488.77
0.06
+ 23.90

1970

Change due to s~ral:ing Y__~
Confidence level 4/
percent 11
where significant -

Grasses (contd)
Hokdeum bhachyanthekUm
J UYlCu,~ ba..Uicu,o

Koe£vUa c.JUJ.,:taXa
OJtzyop.6-u' hymevwJ.du
PiLe.eum pJta:ten~ e
Poa-CMex geyvU
S.{;taiU.on hy.6:tJUx
S:t.{pa

0.00
0.00
0.05
0.00
0.02
14.15
0.00
0.32

0.00
0.00
0.25
0.00
0.11
29.52
0.00
1.15

15.01

0.00
0.00
0.83
0.00
0.36
233.87
0.06
5.23

-

+211.84

.15

+507.32

I

CI\

'"
I

Tota 1 Grasses

16.37

36.43

270.86

880.86

222.81

+658.05

+295.33

.10

Shrubs
Acvr.. g.tabJtum
Ame.tanciuvr.. a.tiU.6oUa.
AJt:tem.u..{a eana
AJt:tem.u.J.a :tJUden:ta.:to.
Cvr..COCMpu,o montanu,o
Ch.Jt.Y.6o:thamnu,odepJtu.6U.6
CIvrY.6o:thamnu,o Mu,oeO.6U.6
ChILY.6o:thamnu,o v.i.6 cJ.dJ.6.t0ltU6
CJta:taegu,o vr..y:thILopoda
Popui.u,o :tJtemui.oJ.du
PILUl1u,o v.{Jtg.{n.{ana
PWl6h.{a :tJUd en:ta,ta

0.01
8.13
0.00
2.56
0.00
0.04
0.00
0.48
0.00
0.00
6.10
0.00

0.00
0.24
0.00
0.24
- 0.24
5.44
7.98
136.03 115.33 126.63
- 11.30
- 8.92
0.00
0.00
0.00
0.00
0.00
3.34
43.12
65.13
54.88
+ 10.25
+ 18.67
0.00
0.00
0.00
0.00
0.00
0.03
0.65
0.59
0.65
0.06
0.09
1.84
0.00
'0.00
+ 1.84
0.37
7.85
9.16
14.13
- 4.97
- 35.19
0.00
0.00
0.00
0.00
0.00
0.02
0.00
0.42
0.01
+ 0.41
3.96
5.73
100.34 100.87
89.48
+ 11.39
+ 12.73
0.00
0.00
0.00
0.00
0.00
-------------------------------------------------------------------------------------------------------------------------------~-----------

-

�Table 4. Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying (contd).
Lbs of vegetation
produced per acre

Percent composition
Plant species

1970

1973

E 73 1I

1970

1973

E 73

202.16
6.42
9.28
337.53

202.46
9.84
31.67
306.49

1bs/ac

Change due to spraying fI
Confidence level
percent ~
where significant 11

Shrubs {contd}
QueJtc.u,; gambe..UU
7U.bv.,
Ro.6 a. nu-tll.ana.
SymphoJUc.aJtpo.6

a..tbu,;

Total Shrubs

12.33
0.66
1.40
15.85

--

47.56

9.7l0.33
0.43
15.02

--

38.75

12.90
2.08
19.75

206.57
11.18
23.20
263.01

--

792.18

-- 0.30
3.42

-

- 22.40
+ 31.04

0.15
- 34.74
- 70.71
+ 10.13
+

--

848.74

-813.91

+ 34.83

2333.00

1552.52

+780.48

--

.10

4.28

-TOTAL ALL PLANTS

100.00

100.00

1661.27

+ 50.27

I

0\
...,
I

1/

- Reflects the amount of forage one would expect to find on a treated area after treatment had treatment not occurred.
flOerived through comparison of sprayed and control areas with adjustments for normal year to year changes in forage production recorded
on the control areas.
3/

- Percent changes are only shown for the more abundant plants.
~Indicates the highest confidence level at which changes due to spraying are significant.
a .05, .10, .15, .20 or .25 level are shown.
.

Only changes that are significant at the

�Table 5. Changes in production of air dry summer vegetation recorded on the chained areas two year3 following chaining.
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 73]J

A

1970

1973

E 73

1bs/ ac

26.82
2.50
0.18
0.65
0.30
0.00
0.36
0.24
1.13
0.00
0.00
2.14
85.94
0.00
0.00
0.00
0.48
0.00
3.87
0.18
3.75
0.00
2.62
0.00
0.00
1.07
0.00
0.00

86.78
5.77
0.06
3.45
0.00
0.00
0.00
0.06
5.06
0.00
0.18
2.91
77.86
0.00
0.00
0.06
0.71
0.18
0.65
0.54
2.26
0.12
1.07
0.00
0.00
0.12
0.59
0.00

58.08
2.30
.1O.61
4.19
0.02
0.00
0.02
0.08
1.67
0.00
0.01
1.84
61.45
0.00
0.00
0.01
0.57
0.01
2.99
0.15
4.94
0.,12
3.97
0.00
0.00
1.65
0.12
0.00

+ 28.70
+ 3.47
- 10.55
0.74
- 0.02
0.00
0.02
0.02
+ 3.38
0.00
+ 0'.17
+ 1.08
+ 16.41
0.00
0.00
+ 0.05
+ 0.14
+ -0.17
2.33
+ 0.39
2.68
0.00
2.90
0.00
0.00
1.53
+ 0.48
0.00

Change due to chaining £I
Confidence level
percent Y
where significant Sf

'Forbs
Ac}uUe.a tal1ulo.6a
AgM.tacJle. uM:.&lt;:.ci6o.eA.a
Ago.6vU.6 gtauc.a
Ae.Uwn ac.wn,[l1atwn
AI1M0.6 ac.e. .6e.pte.rWUOI1a.U6
Aqu.,{..te.g,[a c.oe.nule.a
Mab-&lt;..6 MWnmOl1cU
AlU1b-&lt;..6IWwu..ta
A/~e.flcvUa c.o;lg e..6ta
AIt.te.m,{.J.&gt;,[a
b,[e.nrU!.,
AIt.te.m,{.J.&gt;,[a
dJr.ac.u.nc.U£U.6
Alt.te.m,{.J.&gt;,[a
tudov,[c.,{.a/ta.
M teA - EIUg e.nOfl
M.tIU1g atU.6 hayde.MaYtU.6
Ba.t.6amoiliza .6ag,{..t.tata
Be.nb~
lte.pe.11.6
Cato c.hoJt.tu~ 9 UftrU!.,0 n,{.,{.
Cap.6e.Ua. buMa-pMtow
CM UUe ja. .eA.naft,[ae.6ow
Cite.l1opocUwn be.n.tancUru
C,{.M,[um c.e.n.taUlte.ae.
CoUom-ta. .eA.11e.aW
ComaHdJr.awnbe.Uata
Cymopte.nU.6 pUltpUlte.U.6
Vupluft,[wn
I1WOM
VUpluMwn
oc.cidentate.
V&lt;Y~CUll ct-(J,"ia.~ 0 ph.i..a
V.v., POltWl1 .tJtac.huc.Mpwn

1.50
0.14
0.01
0.04
0.02
0.00
0.02
0.01
0.06
0.00
0.00
0.12
4.78
0.00
0.00
0.00
0.03
0.00
0.21
0.01
0.20
0.00
0.15
0.00
0.00
0.06
0.00
0.00

4.38
0.30
0.00
0.16
0.00
0.00
0.00
0.00
0.25
0.00
0.01
0.15
3.93
0.00
0.00
0.00
0.03
0.01
0.03
0.03
0.11
0.01
0.05
0.00
0.00
0.01
0.03
0.00

3.47

3.68

+ 49.42
+150.75

.25

+202.28

.15

-

-

-

I

0\
.,..
I

+ 58.80
+ 26.71

.20
- 54.29
- 73.02

-

------------------------------------------------------------------------------------------------------------------------------------------------

�Table 5. Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining (contd).
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 73 !I

1973

E 73

1bs/ec

1.49
2.38
0.00
44.79
0.00
7.32
0.00
0.00
0.42
0.89
0.00
0.18
6.25
0.00
0.00
104.20
0.00
23.08
0.71
0.00
40.68
0.00
0.00
0.18
3.63
0.00
0.18

12.27
0.65
0.00
19.88
0.00
2.83
0.00
0.36
0.02
1. 05
0.00
0.74
7.26
0.06
0.00
83.86
0.00
17.35
0.87
0.00
54.09
0.00
0.00
0.01
0.53
0.00
O.01

- 10.78
+ 1.72
0.00
+ 24.91
0.00
+ 4.49
0.00
0.36
0.40
- 0.16
0.00
0.57
1.01
0.06
0.00
+ 20.34
0.00
+ 5.73
0.16
0.00
- 13.41
0.00
0.00
+ 0.17
+ 3.10
0.00
+ 0.17

1970

Change due to chaining Y ____
Confidence level
percent '}j
..where significant y

Forbs (contd}
EJUogonwn wnbe.UtLtwn
Ep-Leob-i.wlIparucu1a-twn
Fltagcvda amvu.cana
Ga.uwn
G~nUaY'.a heteJto~ epa..f.a
GVtaIUwn
Gilia ((ggltegata
GJt-i.nde£ia aphana~
Hack~
teptophylta
Hueluw)1 hoop~U
Hyd!tophyUwn capUatwn
Hyd!tophyUwn 6endlvu.
lw m-i.MowUel'L6M
Lac.:tuca &lt;I caJt-i.ota
Lapputa ltedo~V-6
W
LathyltU4 - V-i.c.-i.a
Lep-i.d-i.wndeM-i.6toltwn
Ug~ticwn
Unwn te.w.U,U
Lomatiwn teptocaltpwn
Lup-i.n~ Mg ente~
Mad-i.agtomeJtata
Med-i.cago &lt;lativa
"1e.Uto,t~ o66-i.c.-i.~
MeJt:teM-i.a 6Mnwcana
MeJt:teYL6-i.atanceola.tct
MoUav.ica paltv-i.61Ol!a

0.25
0.00
0.00
1.26
0.00
0.34
0.00
0.02
0.08
0.16
0.00
0.06
0.41
0.00
0.00
5.20
0.00
1.21
0.05
0.00
5.24
0.00
0.00
0.00
0.03
0.00
0.00

0.07
0.12
0.00
2.21
0.00
0.36
0.00
0.00
0.02
0.05
0.00
O.01
0.32
0.00
0.00
5.02
0.00
1.10
0.04
0.00
2.04
0.00
0.00
0.01
0.19
0.00
0.01

1.19

4.82
1.00
3.20

4.52
0.00
0.00
22.96
0.00
6.25
0.00
0.30
1. 37
2.91
0.00
1.01
7.26
0.06
0.00
96.77
0.00
22.60
0.95
0.00
93.80
(\00
0.00
0.00
0.59
0.00
0.00

- 87.88
+ 125.31
+158.69

.25

-

--

-

I
0\
V1
I

- 13.93
+ 24.25

+ 33.03

.15

- 24.79

--------~.------~----------------------------------------------------------------------------------------------------------------------------

�Table 5. Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining (contd).
Lbs of vegetation
produced per acre

Percent composition
Plant species

1970

1973

E 7311

E 73

1bs/ac

0.00
0.00
0.00
0.00
0.54
0.12
4.10
5.24
4.46
3.46
0.00
0.00
0.00
0.08
0.00
0.10
0.77
0.30
0.30
0.01
6.42 21.95
1.31
0.36
0.48
0.71
13.20
6.51
0.00
0.00
0.00
0.00
2.97
0.55
0.06
0.01
0.12
0.71
0.06
0.89
0.00
0.18
0.00
0.00
1.01
0.00
27.18 15.82
0.00
0.00
4.82
9.71
17.72 13.82

0;00
0.00
+ 0.42
- 1.14
+ 1.00
0.00
- 0.08
- 0.10
+ 0.48
+ 0.29
- 15.52
- 0.95
-+ 0.23
6.69
0.00
0.00
+ 2.43
+ 0.05
0.59
0.83
0.18
0.00
+ 1.01
+ 11.36
0.00
-+ 4.90
3.90

1970

1973

Change due to chaining £I
.3
Confidence level
percent -I
where significant i!

Forbs (contd}
Oeno.t!teJta. caes p.Uc~4
Opuntia
OlZ/thocMpuo .t'.u.:teuo
0'6moltluza.
Pa.ch-iAlWna mYM.£nJ;tu
Pedicuta!1M g!lay-i..
Pen6temo.11 c.~'Vlda..e..e.,u
Pen6temoH ,s.tuuuo
Phac.eUa heteJtophyUa.
Ph.t'.ox .t'.ong-i..6oUa
Po.t'.emoluwlI ~olio-6~-6hnum
Po.t'.ygonum av-i..cutMe
Po.t'.ygonum dOug.t'.MU
Poten..tLUa putclteJ!.!t.i.ma.
Ra.nw1cu.t'.uo -i..ltamoenus
Rumex ~puo
Rudbec~
montana
Senec,£o amblto-6-i..o-i..du
Senec,£o .£ntegeJ!.!t.i.muo
S enec,£o -6eJtJta
Sdene menz.£u«
SJJl-i...t'.ac,£no.
-6teUato.
Solidago -6PCI.M-i..o.t'.OItCl.
·SteUwUa
Swvvt.-io. ItCI.diato.
Ta/to.xa.cum o66-i..c,£no.te
ThaliCMum 6 end.t'.e«

0.00
0.00
0.01
0.33
0.68
0.00
0.02
0.01
0.02
0.00
0.14
0.07
0.01
0.31
0.00
0.00
0.09
0.00
0.00
0.00
0.01
0.00
0.00
0.42
0.00
0.23
0.98

.'

0.00
0.00
0.03
0.20
0.21
0.00
0.00
0.00
0.04
0.02
0.33
0.02
0.02
0.67
0.00
0.00
0.15
0.00
0.01
0.00
0.00
0.00
0.05
1.32
0.00
0.24
0.84

0.00
0.00
0.12
6.13
12.91
0.00
0.36
0.18
0.30
0.00
2.44
1.31
0.12
5.47
0.00
0.00
1.55
0.00
0.06
0.00
0.18
0:00
0.00
7.79
0.00
4.16
18.14

- 21.70
+ 28.82
I

0\
0\
I

- 70.73
+102.84

---

+ 71.77

.15

- 50.41
+ 28.25

-----------------------------------------------------------------------------------------------------------------------------------------------

�Table 5.

Changes in production of air dry summer vegetation

Percent
Plant species

1970

recorded on the chained areas two years following chaining
Lbs of vegetation
~roduced ~er acre

com~osition
1973

E 73 ]J

1970

1973

"
E 73

1bs/ac

2.44
0.00
3.03
0.00
0.00
0.00
1.49
0.59
1. 72
13.26

53.06
0.00
4.22
0.00
0.00
0.00
1.25
0.59
7.84
16.23

- 50.63
0.00
1. 19
0.00
0.00
0.00
+ 0.24
0.00
6.12
2.96

(contd).

Change due to chaining ~
Confidence level
percent ~
where significant

1/

Fo rbs (contd}
Tkagopogon dubium
Tkagopogon pkaten6~
UJL.ti.ca dioica
va£vUana
ocuden.ta1~
VVtontca
ClJllvUcana
Vekol1ica bUoba
V-i.gu.i.eM mu.f.tiMoka
Viofu kugu1.0.6a.
V-i.o.f.a .6he.UonU
Wyeth-ta amp.f.exica~

Tota 1 Forbs

0.10
0.00
0.24
0.00
0.00
0.00
0.08
0.02
0.22
0.91

1. 78
0.00
4.22
0.00
0.00
0.00
1.43
0.30
4.04
17.13

0.12
0.00
0.16
0.00
0.00
0.00
0.07
0.03
0.08
0.62

-- --

-- --

26.56

483.67

26.29

- 95.40

-

+ 18.79

-

- 78.00
- 18.27

531.91

424.38

+107.53

+ 25.34

21.00
0.18
0.00
2.74
3.63
0.36
0.00
1.13
0.12
0.00
12.85

9.13
0.01
0.00
2.22
7.61
0.01
0.00
0.01
0.54
0.00
0.06

+ 11.86
+ 0.17
0.00
+ 0.51
3.98
+ 0.35
0.00
+ 1.12
0.42
0.00
+ 12.79

+129.86

.10

Grasses
AgMPYkon
AgkOPYMIt ~.ta.tum
AgkoPYMn '&lt;'n.tekmedium
BkOmU.6anoma£U.6
BMmU.6 '&lt;'nekm~
BlLomU.6tectOkum

CMex
Va.cty.f.U.6 g.f.omekata.
UymU.6 g.f.aucU.6
Fe.6tuca ov.&lt;.na.
Fe.6tuca. tilUkbvU

1. 22
0.00
0.00
0.22
0.02
0.04
0.00
0.00
0.12
0.00
0.00

1.05
0.01
0.00
0.14
0.17
0.02
0.00
0.05
0.01
0.00
0.66

0.55

21.89
0.00
0.00
3.98
0.36
0'.65
0.00
0.00
2.14
0.00
0.00

-

+ 23.10

-

-----------------------------------------------------------------------------------------------------------------------------------------------

I

'"

"-I
I

�Table 5. Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining (contd).
Lbs of vegetation
~roduced ~er acre

.Percent com~os;tion
Plant species

1970

1973

E 73 Jj

1973

E 73

0.30
0.12
0.83
0.00
0.00
402.25
0.00
5.89

9.70
0.00
0.04
0.00
0.00
425.78
0.30
5.94

1970

1bs/ac

Change due to chaining ~
Confidence level
percent Y
where significant i!

Grasses (contd}
Ho~deum bkachyanth~

J WlcU,~ ba£:ti..cw..
KouvUa.
CJIM.ta:ta.
O~ZUOpf.,.u.,hymeno,[du
PiLteum pka.teY!f.J e
Poa-Cakex gey~
S.uc:n,[on hY.6bU.X
SUpa

0.15
0.00
0.00
0.00
0.00
23.77
0.02
0.37

0.02
0.01
0.04
0.00
0.00
19.74
0.00
0.30

25.92

22.21

25.22

2.68
0.00
0.00
0.00
0.00
431.57
0.30
6.60

- 9.40

+
+

0.12
0.80
0.00
0.00
- 23.53
- 0.30
- 0.05

- 5.53
- 0.90
I
0'\

Total Grasses

470.17

ce

451.37

386.77

0.00
0.00
1ll.58 138.52
7.49
2.62
34.91
35.33
0.00
0.00
1.07
0.00
0..00
0.00
0.95
0.00
5.29
5.17
0.00
0.00
154.88 228.57
0.00
0.00

0.00
103.87
3.33
44.44
0.00
0.01
0.00
1.71
5.29
0.00
138.12
0.00

+ 64.60

+ 16.70

Shrubs
Ac~ g.ea.b~um
Amuanch,[~
a1.n,Lno.Ua.
AlI-temiJ.,,La cana
MtemiJ.,,[a. bU.derz;ta..ta.
C~COCakPw.. morz;tanw..
C~f.,othamnw..
dep~uf.,w..
CMyf., othamnus naw.. eo.6w..
CMyf., othamnus
UCU6.eO~W..
Cnataegw.. ~y~opoda.
PopU£w.. ~emU£o'[du
P~Wlw.. v~g'[n,Lana.
PU/L.6h,[a ~en-ta..ta.

v.u.,

0.00
5.98
0.42
1.95
0.00
0.00
0.00
0.05
0.30
0.00
8.42
0.00

0.00
6.52
0.13
1.81
0.00
0.05
0.00
0.00
0.27
0.00
11.19
0.00

5.86

7.91

0.00
+ 34.65

- 0.71
- 9.11

0.00
1.06
0.00
1.71
- 0.12
0.00
+ 90.45
0.00

+ 33.36

- 21.43
- 20.49

+

-

- 2.25
+ 65.48

----------------------------------------------------------------------------------------------------------------------------------------------

I

�Table 5.

Chunges in production of air dry summer vegetation recorded on the chained areas two years following chaining (contd).
Lbs of vegetation
Qroduced Qer acre

Percent comQosition
Plant species

1970

1973

E 731/

Change du~ to chaining gj
Confidence 1eveb
percent Y
where signiflcan 4/

1973

E 73

lbs/ac

331.82
0.00
43.54
185.45

432.22
0.00
37.77
185.51

325.24
0.00
59.45
216.11

+106.99
0.00
- 21.68
- 30.60

- 36.47
- 14.16

875.93

1066.79

899.95

+166.83

+ 18.54

1970

Shrubs (contd)
QuVtCtM gambe.LtU
lUbv.,
ROMl i1utkw1a
SymplwIUCMP0,6
a1.btM

Total Shrubs

17.81
0.00
2.40
10.18

20.54
0.00
1.89
9.11

-- -47.51

51.50

18.64
3.57
12.68

+ 32.89
.20

.15

I

0-

'"

I

Total All Plants

100.00

100. 00

1829.77

2050.07

1709.99

+340.08

+ 19.89

1/

- Reflects the amount of forage one would expect to find on a treated urea following treatment had treatment not occurred.
gjoerived
through
comparison of chained and control areas with adjustments for normal year to year changes in forage production recorded
on the control
areas.
3/

- Percent changes are only shown for the more abundant plants •.
~Indicates

the highest confidence level at which changes due to chaining are significant.

ex .05, .10, .15, .20 or .25 level are shown.

Only changes that are significant at the

�Table 6.

J!

Season
Winter

recorded

Forage
value

to Post-Treatment

Spring

to Post-Treatment

Summer

Fall

to Post-Treatment

- 4.59
+ 24.02
- 4.97

+ 3.71
- 58.69
- 89.98

1 : 0.64
- 15.20
+ 0.90
- 83.14

1:

0.84

- 65.13
-111.07
- 63.98
Ratio

Highly val.
Valuable
Least val .

1:

- '17.34
- 20.92
- 23.06
0.80

- 30.88
-309.59
0.00
Ratio

1:

- 8.23
- 59.92
0.00
0.67

continued

on following

page)

dry elk forage.

%

- 1.08
+ 6.16
- 35.19

+488.81
+124.54
+ 13.18

+203.44
+ 30.58
+ 24.50

1 : 2.09
+484.52
+177.22
+ 19.46

+118.17
+ 32.68
+ 8.57

1 : 1.72
+588.73
+ 61.93
0.00

+161.61
+ 13.59
0.00.

1 : 1.94

as degree
and Fan;

]J ,_ 2/

Change due
to chaining
lbs/ac
+227.76
- 59.98
- 1.71

1 : 1.01

1/
,
- Table is based on the a'ssumptjon that due to such factors
only shrubs during Winter; grasses and shrubs during Spring
on measurements made duri ng Summer.
'
(footnotes

air

+ 8.64
-332.55
- 24.56

- 41.58
+ 2.68
-137.01

Highly val.
Valuable
Least val.

seasonal

Change due
to spraying
lbs/ac

Ratio

to Post-Treatment

in available,

Change due
to burning jJ
lbs/ac
%

Ratio ~

Highly val.
Valuable
Least val.

Pre-

Pre-

changes

Highly val.
Valuable
Least val.

Pre-

Pre-

Two year

%

Confidence level where treatment
difference
is significant
Control Control
Control
Burn
Burn
Spray
&amp; Burn &amp; Spray
&amp; Chain
&amp; Spray
&amp; Chain
&amp; Chain

5/ '

+ 39.85
- 18.67
-100.00

1 : 1.24
- 27.76
+295.87
- 7.92

- 6.46
+ 54.36
- 18.31

.20

.15

.20

.20
.25
I

?

1 : 1.21
+ 10.73
+156.90
+ 36.02

"

+ 1.83
+ 29.62
+ 25.42

.15

.25
.15

.20

.20

.20
.05

.05

1 : 1.13
+ 70.23
+ 40.52
0.00

+ 13.12
+ 9.80
0.00

.10

.25
.20

1 ~ 1.12

of plant maturity and snow cover, plant availability
to elk will
and forbs, grasses and shrubs during summer. Data in this table
.

,
consist of
are based

�Table 6. Two yea!' changes recorded in available, seasonal air dry elk forage (contd).
(footnotes contd)

y
Classification of study area plants as elk forage was based on the average, relative consumption of those plant species by elk in 48 elk
food habits studies reported in the literature (Kufeld 1973).
.
3/

- Seasons:

Winter = December, January, February;

Spring = March, April, May:

Summer = June, July, August;

Fall = September, October, November.

4/

- Derived through comparison of treated and control areas with adjustments for normal year to year changes in forage production recorded on the
control areas.
~/Indicates the highest confidence level at which treatment differences are significant.
a .05, .10, .15, .20 or .25 level are shown.

Only greatment differences that are significant at the

6/

- The ratio of elk forage produced before and after treatment adjusted for normal year to year changes measured on the control areas. A single
value for elk forage was derived by weighting pounds per acre of highly valuable forage produced by a factor of "3", valuable by "2", and least
valuable forage by "1". A total weighted value was then computed.

I
....•

....
I

�Table

7.

Season ]I

changes

recorded

Forage
value
Hea vi ly ea ten
Mod. eaten
Lightly eaten

Hinter

Pre-

Two year

to Post-Treatment

Spring

SUlTTIler

Heavi ly eaten
Mod. eaten
Lightly eaten

Pre- to Post-Treatment
Fall

Heavi ly eaten
Nod. eaten
Lightlyeaten

Pre- to Post-Treatment

Ratio

dry deer forage.

1/, Y

Change due
to chaining
lEis/ac

-157 ..99
+ 12.62
-235.11

+

9.11

+

3.38

- 16.84
+228.34
- 46.11

+ 87.20
+ 5.09
- 52.84

- 92.40
- 27.08
- 13.93

1 : 0.59

+ 19.35
-186.59
- 98.53

Ratio

air

Change due
to s~raxi n9
1bs/ac
%

-153.25
-179.64
- 41. 91

Ratio

seasonal

Change due
to burni n9 ~
lbs/ac
%

Ratio ]/1 : 0.54

Heavily eaten
Mod. eaten
Lightly eaten

Pre- to Post-Treatment

in available,

+ 10.81
- 25.63
- 18.72

1 : 0.84

+ 9.08
-327.18
- 21. 47

+ 4.30
- 53.05
- 60.36

1 : 0.66

- 0.57

+ 16.27
0.13
+ 0.97

-

1 : 1.02
+ 10.27
+116.17
+486.92

+ 18.72
+ 15.93
+202.17

1 : 1.40·

- 7.88

- 1. 94

+169.82

1 : 1.20
+ 59.54
+ 26.29
- 3.43

- 30.74
+ 39.92
- 17.12

.15

.05

.10

1 : 1.23

-

- 18.70
- 16.74
+663.78

%

Confidence level where treatment
difference
is significunt?J'
Contra 1 Control
Control
Burn
Burn
Spray
&amp; Burn &amp; Spray
&amp; Chain
&amp; Spray &amp; Chain
&amp; Chain

+ 31.24
+ 4.25
- 34.82·

1 : 1.13

-

8.90
+218.45
- 22.11

- 20.04
+ 26.32
5.21

-

.15

.15
.10

.10

.25
.05
.10

.05
.10
I

-.J

1 : 1. 17
+110.88
+103.02
+ 38.66

+ 30.31
+ 13.46
+ 6.82

N
I

.20

.15

.10

.05

.05

.15

1 : 1.18
+148.10
+ 44.45
0.00

+ 48.54
+ 7.60
0.00

.05

1 : 1.26

1/
- Table is based on the assumption that due to such factors
only shrubs during Winter; grasses and shrubs during spring
on measurements made duri ng Summer.
(footnotes

conti nued on fa l Iowi ng page)

as degree
and fall;

of plant maturity and snow cover, plant availability
to deer will consist
of
and forbs, grasses and shrubs during summer. Data in this table are based

�Table 7. Two year changes recorded in available, seasonal air dry deer forage (contd).
(footnotes contd)
2/

- Classification of study area plants as deer forage was based on the average, relative consumption of those plant species by deer in 99 deer
food habits studies reported in the literature (Kufeld, Wallmo and Feddema 1973).
3/

- Seasons:

viinter = December, January, February;

Spring = March, April, May; Summer = June, July, August;

Fall = September, October, November.

4/

- Derived through comparison of treated and control areas with adjustments for normal year to year changes in forage production recorded on the
control areas.
5/

- Indicates the highest confidence level at which treatment differences are significant.
0;
.05, .10, .15, .20 or .25 level are shown.

Only treatment differences that are significant at the

6/

- The ratio of deer forage produced before and after treatment adjusted for normal year to year changes measured on the control areas. A single
value for deer forage was derived by weighting pounds per acre of heavily eaten forage by a factor of "3", moderately eaten forage by "2", and
lightly eaten forage by "1". A total weighted value was then computed.

I
....•

...,
I

�Table 8. Two year changes recorded in air dry, summer cattle forage. l!

Season
Summer ~

Forage
value
Desirable
Intermediate
least desirable

Change due
to burni ng 2/
lbs/ac
%

l bs/ac

%

+ 3.66 + 6.21
- 34.56 - 7.55
-259.21 - 24.29

+ 83.98
+538.57
+115.48

+254.37
+100.52
+ 11.25

u

Pre~ to post-treatment ratio ~

1 : 0.95

Change due
to sprayi ng_

2.17

Change due
to chaining
lbs/ac
%
+ 41.36
+120.47
+145.58

+119.59
+ 15.60
+ 15.56

Confidence level where treatment difference is significant
Control Control
Control
Burn
Burn
Spray
&amp; Burn &amp; Spray &amp; Chain &amp; Spray &amp; Chain
&amp; Chain

.15
.05

.15

.15
.05

.05

11

.15
.10

1.24

1/

- Plants considered cattle forage are those classified as "desirable" and "intermediate" by the U. S. Forest Service, Region 2, Range Analysis
Handbook for the foothill shrub, aspen weed, and mountain meadow vegetation types.
gjDerived through comparison of treated and control areas with adjustments for normal year to year changes in forage production recorded on the
control areas.
1IIndicates the highest confidence level at which treatment differences are significant.
a .05, .10, .15, .20 or .25 level are shown.

Only treatment differences that are significant at the

4/

- Summer = June, July and August.
~The ratio of cattle forage produced before and after treatment adjusted for normal year to year changes measured on the control areas. A
single value for cattle forage was derived by weighting pounds per acre of desirable forage produced by a factor of "2", and intermediate
forage by "1". A total weighted value was then computed for desirables and intermediates. Production of least desirable was excluded from
the ratio since least desirables are not considered cattle forage.

I
....•
-I'-I

�Table 9. Comparison of soil properties on control, burned, sprayed and chained areas from which plants were collected for nutrient analysis.

Treatment

pH Conductivity l!

Percent
Organic Nitrorn
Matter
(ppm

Control
(Unit 6)

7.5

.7

7.8

24

27

780

4.10

4.30

0.94

4.00

3.69

1.23

0.86

Low

Silty clay
loam

Burn
(Unit 7)

8.0

.5

8.0

2

27

730

2.40 34.2

0.76

3.75

3.14

0.91

0.75

Low

Silty clay
loam

Spray
(Unit 5)

7.5

.3

6.8

1

13

418

2.70 41. 0

0.94

4.65

1.95

0.66

0.60

Low

Silty clay
loam

Phosphorous Potassium Zinc Iron Copper Manganese Calcium Magnesium Sodium
(ppm)
(ppm)
(ppm) (ppm) (ppm)
(ppm) (meg/L) Y (meg/L) (meg/L) Lime

Texture

I
....•
\J1

Chain
(Unit 8)

7.6

.4

lI1~i1imhos per centimeter.

YMilequavalent

per liter.

8.0

4

27

400

3.50 42.0

0.70

5.00

2.00

I

0.91

0.45

Low

Silty clay
loam

�Tab 1e 10. Nutrient and mineral content of summer collected plants 2 years following treatment (collected July 24-27. 1973).

Average cell content
Control Burn Spray Chain

Plant
type

Nutrient or
mi nera 1

Measurement

Dry or green
wt basis

Forb

Total cell contents
Protei n
Ether extract
Soluble cho.
Soluble ash

%
%
%
%
%

Green
Green
Green
Green
Green

14.19
3.45
1.95
6.22
2.57

16.28 13.87 16.25
3.63 3.30 3.91
2.37 1.96 2.52
7.78 6.18 7.35
2.50 2.44 2.46

Total cell walls
Insoluble ash
Lignin
Holocell ulose

%
%
%
%

Green
Green
Green
Green

11.49
.25
2.93
8.30

11.84 12.36 11.54
.14
.26
.13
2.69 2.88 2.68
9.02 9.27 8.71

~loisture

%

Green

74".32

71.87 73.01 72.21

Nitrogen
Calcium
Phosphorous
Cal/phos ratio
Iron
Zinc
Copper
Magnesium
Manganese
Sodium
Potassium

%
%
%

ppm
ppm
ppm
ppm
ppm
ppm
ppm

Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry

2.14
2.03 1.99 2.24
1.45
1.37 1.39 1.31
.16
.13
.14
.14
10.18
9.98 11.18 11.17
143.50 116.20 98.80 81.50
29.85 47.15 34.69 50.62
9.15 17.62 11.46 19.77
.26
.26
.24
.28
36.46 26.85 36.92 31.31
415.38 403.08 543.08 461.54
2.86
2.18 2.77 2.34

Total cell contents'
Protein
Ether extract
Soluble cho.
Soluble ash

%
%
%
%
%

Green
Green
Green
Green
Green

13.76
4.59
2.82
4.14
2.21

Significance at the a .05 confidence level 1.1
Contro 1 Contro 1 Contro 1 Burn
Spray
Burn
&amp; Burn &amp; Spray &amp; Chain &amp; Spray &amp; Chain &amp; Chain
L-H

L-H
L-H
L-H
L-H

L-H

H-L
H-L

L-H
L-H
L-H
L-H

...•
I

Grass

14.09 13.74 15.24
4.07
4.28 4.13
2.65 2.52 2.53
5.47 4.90 6.32
1.91 2.04 2.27

H-L

L-H
L-H

0\
I

-

H-L
L-H
L-H

H-L
H-L

L-H
l..-H

H-L

L-H
L-H
L-H

H-L

H-L
L-H
H-L

L-H

-----------------------------_._--------------------------------------------------------------------------------------------------_._-----------------

�Table 10.

Plant
type

Nutrient and mineral content of summer collected plants 2 years following treatment (collected July 24-27, 1973) (contd).

--

Nutrient or
mineral

Measurement

Dry or green
wt basis

Average cell content
Control Burn Spray Chain

Significance at the a .05 confidence level l!
Control Control Control
Burn
Burn
&amp; Burn &amp; Spray &amp; Chain &amp; Spray &amp; Chain &amp;Spray
Cha in

Grass (contd)

Shrub

Total cell walls
Insoluble ash
Lignin
Holocellulose

%
%
%
%

Green
Green
Green
Green

24.97
1.01
4.15
19.81

~loisture

%

Green

61.28

Nitrogen
Calcium
Phos phorous
Cal/phos ratio
Iron
Zinc
Copper
Magnesium
t1anganese
Sodium
Potassium

%
%
%

ppm
ppm
ppm
ppm
ppm
ppm
ppm

Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry

59.78 61.05 60.70
1.83
1.62 1.79 1.65
.50
.41
.47
.42
.14
.13
.12
.13
3.55
3.10 3.88 3.30
138.80 95.00 78.80 51.30
22.50 35.75 25.00 37.00
7.00 14.00 7.75 13.50
.12
.11
.12
.12
26.00 30.25 33.50 25.25
385.00 380.00 462.50 417.50
1.87
1.69 1.84 1.43

Total cell contents
Protein
Ether extract
Soluble cho.
Soluble ash

%
%
%
%
%

Green
Green
Green
Green
Green

23.88
5.68
2.03
13.82
2.34

22.81 22.48 23.96
4.99 5.20 5.47
1.97 1.97 2.22
13.58 13.06 14.07
2.20 2.26 2.06

Total cell walls
Insoluble ash
Ligni n
Ho 1ace 11ulose

%
%
%
%

Green
Green
Green
Green

16.28
.15
5.57
10.55

15.50 16.41 16.20
.15
.13
.11
5.02 5.30 5.41
10.33 10.97 10.69

Moisture

%

Green

59.84

61.69 61.11 59.84

25.21 25.21 24.98
1.08
.87
.73
4.44 4.30 4.34
19.70 20.06 19.91

-..JI
-..J
I

L-H
L-H

H-L
L-H
L-H

H-L

H-L

.,.

----------------------------------------------------------------------------------------------------------------------------------------------

L-H

�Table 10. Nutrient and mineral content of summer collected plants 2 years following treatment (collected July 24-27, 1973) (contd).
Plant
type

Nutrient or
minera1

Measurement

Significance at the a .05 confidence level l!
Dry or green
Average cell content
Confr-ol .Control Control -Burn
Burn
Spray
wt basis
Control Burn Spray Chain &amp; Burn &amp; Spray &amp; Chai.n &amp; Spray &amp; Chain &amp; Chain

Shrub (contd)
Nitrogen
Calcium
Phosphorous
Ca1/phos rati0
Iron
Zinc
Copper
r~agnesium
Manganese
Sodium
Potassium

%
%
%

ppm
ppm
ppm
ppm
ppm
ppm
ppm

Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry

2.26
2.16,\2.17
2.25
1.15
1.07 1.07 1.03
.16
.17
.15
.14
7.59
6.83 7.57 7.90
77 .10 83.60 88.60 61.40
33.57 39.14 36.43 37.71
9.43 11.14 10.00 10.29
.24
.26
.26
.24
61.29 41.57 56.00 46.00
455.71 487.14 447.14 468.57
. 1.36
1.42 1.45 1.27

H-L

lIWhere
differences
designated
by "H". significant at a .05 occur the treatment with the lower value is designated by "L" and the higher value treatment is

H-L

I
......
00
I

�Table 11.

Nutrient

and mineral

content

of winter

collected

plants

3 years

following

treatment

Plant
type

Nutrient or
minera 1

Measurement

Dry or green
wt basi s

Grass

Total cell contents
Protein
Ether extract
So1ub1echo.
Soluble ash

%
%
%
%
%

Dry
Dry
Dry
Dry
Dry

34.64
6.50
6.09
19.42
2.62

37.78
7.40
7.35
19.85
3.24

34.23
6.90
7.26
17 •45
2.62

33.98
7.80
6.55
16.50
3.14

Total cell walls
Insoluble ash
Li gnin
Holocellulose

%
%
%
%

Dry
Dry
Dry
Dry

65.37
3.42
9. 05
52.90

62.23
2.63
11 .75
47.85

65.78
4.21
7.57
54.00

66.02
2.80
10.50
52.73

Nitrogen
Ca1ci urn
Phosphorous
Cal/phos ratio
Iron
Zinc
Copper
Magnesium
Manganese
Sodium
Potassium

%
%
%

ppm
ppm
ppm
ppm
ppm
ppm
ppm

Dry·
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry

1. 04
1. 18
1.10
1. 25
.35.36.29.35
.16
.21
.18
.17
2.30
1.80
1.70
2.10
165.00 250.00 198.00 193.00
33.00
38.00 28.00 43.00
05.00
22.00 04.00 08.00
.115
.138.130.158
45.00
50.00 20.00 40.00
225.00 135.00 225.00 225.00
.63
.76
.60
.89

Total cell contents
Protein
Ether extract
Soluble cho.
Soluble ash

%
%
%
%

Dry
Dry
Dry
Dry
Dry

37.75'
6.20
6.72
21.39
3.44

(collected

November 12, 1974).

Significance
at the a .05 confidence level
Average cell content
Control
Control
Control
Burn
Burn
Control
Burn Spray Chai n &amp; Burn
&amp; Spray
&amp; Chain
&amp; Spray &amp; Chain

l!

Spray

&amp; Chai n

....•
'P
I

Shrub

%

40.38
5.80
7.22
23.97
3.36

40.88
6.10
7.25
24.83
2.66

L-H

H-L

H-L

40.43
5.90
7.52
23.92
3.13

-------'---------------------------------------------------------------------------~--------------------------------------------------------------

�Table 11. Nutrient and mineral content of winter collected plants 3 years following treatment (collected November 12, 1974) (contd).
Plant
type

Nutrient or
mineral

Measurement

Dry or green
wt basis

Significance at the a .05 confidence level 1/
Average cell content
~ontro1 .Contra·l Control -Burn-Burn
Spray
Control Burn Spray Chain &amp; Burn &amp; Spray &amp; Chain &amp; Spray &amp; Chain &amp; Chain

Shrub (contd)
Total cell walls
Insoluble ash
Lignin
Ho1oce 11u1ose
Nitrogen
Calcium
Phosphorous
Ca1/ phos rati0
Iron
Zinc
Copper
Magnesium
t~anganese
Sodi urn
Potassi um

%
%

%
%
%
%
%

ppm
ppm
ppm
ppm
ppm
ppm
ppm

Dry
Dry
Dry
Dry

62.25
.46
20.36
41:43

Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry
Dry

.99
.93
.98
.94
.56
.58
.53
.51
.14
.15
.13
.14
4.70
4.50 4.70 4.10
119.00 163.00 139.00 97.00
37.00 40.00 44.00 51.00
12.00 10.00 15.00 16.00
.142
.160 .159 .169
81.00 59.00 43.00 60.00
223.00 279.00 370.00 204.00
.49
.49
.41
.52

59.62 59.12 59.57
.47
.55
.37
17.40 17.19 18.80
41.74 41.37 40.42

1/Where
differences
designated
by "H". significant at.a .05 occur the treatment with the lower value is designated by "L" and the higher value treatment is

I

00
0
I

�Table 12. Elk use recorded on treated areas before and 2 years after burning, spraying and chaining.

Year )J

Control
Block 1 Block 2

Ave.

Block 1

Elk ~er sguare mile
Burn
S~ra:t
Block 2 Ave. Block 1 Block 2

Ave.

.B10ck

Chain
Block 2

Ave.

1971

8.4

10.0

9.2

11.8

18.6

15.2

11.8

13.6

12.9

21.8

18.6

20.2

1974

6.5

18.4

12.5

24.9

35.0

30.1

6.5

11.7

9.1

6.5

9.8

8.3

1971 to 1974 change

+3.3

+14.9

-3.8

-11.9

+36%

+98%

-29%

-59%

-

+9.4

-8.4

-19.1

Percent change 1971 to
1974

Change due to treatment
1971 to 1974 Y

I

Percent change due to
treatment 1971 to 1974

co
....

-

+45% 21

-48%

I

-70%

)JDuring the pre-treatment evaluation phase pellet plots were cleared Sept. 3, 1970, and accumulated pellets counted May 5, 1971.
During the post-treatment phase plots were cleared Sept. 3, 1973, and pellets counted May 8, 1974.
.
YDerived through comparison of treated and control areas with adjustments for normal year to year changes in elk use recorded
on the control areas.
2!Two years after treatment significantly more elk use (p&lt; .10~ was recorded on burns than sprayed or chained areas. However,
changes due to burning, spraying and chaining were not significant at the a .10 level when compared to control areas.

�Table 13.

Deer use t'ecorded on treated areas before and 2 years after burning, spraying and chaining. l!
Deer per_square mi 1 e

Year Y

Control
Block 1 Block 2

Ave.

Block 1

1971

13.6

25.2

19.4

30.2

1974

21. 5

16.6

19.1

15.0

SprayChai
Block 2 Ave.

Block 1

n
Block 2

16.8

15.2

16.0

25.2

6.5

15.9

13.5

16.6

15.0

4.9

9.8

7.4

Ave.

Block 1

15.2

22.7

18.4

16.7

Ave.

--

-0.3

-6.0

-1. 0

-8.5

0%

-26%

-6%

-53%

Change due to treatment
1971 to 1974 ]I

-5.6

-0.8

-8.3

Percent change due to
treatment 1971 to 1974

-25%

-6%

-53%

1971 to 1974 change

Percent change 1971 to
1974

1/

- Due to very low deer use on the area when 2 year post-treatment deer use measurements were made the validity of these data in
providing a true picture of deer response to the various treatments is unknown at this time. Additional data are currently being
collected for verification of data presented- in this table.
YDuring the pre~treatment evaluation phase pellet plots were cleared Sept. 3, 1970, and accumulated pellets counted May 5, 1971.
During the post-treatment phase plots were cleared Sept. 3, 197J, and pellets counted May 8, 1974.
]lDerived through comparison of treated and control areas with adjustments for normal year to year changes in deer use recorded
on the control areas.

I

00
N
I

�Table 14. Cattle use of control and treated areas before (1970) and 2 years after (1973) treatment.

Unit

Treatment

4
6

Control
Control

4 &amp;6

Control

1
7

Burn
Burn

&amp;7

Burn

2
5

Spray
Spray

2 &amp;5

Spray

3

8
3 &amp;8

Cow days use
per acre during
1970 season 11

Cow days use
per acre during
1973 season ?-J

Change in cow
days use per acre
from 1970 to 1973

0.0
1.3

3.0
6.5

+ 3.0
+ 5.2

--

0.7

-4.8

-+ 4.1

4.4
9.6

19.6
31.7

+15.2
+22.1

--

--

--

25.7

+18.7

7.0
8.7
3.0

7.4
10.9

Yes

- 1.3
+ 7.9

5.9

-9.2

+ 3.3

Chain
Chain

0.0
18.3

0.0
- 9.6

Chain

-- '#

0.0
8.7

--

Significance of 1970-1973
change from control
areas at a .10 level

No

-No

1Icow defecation rate is 11.5 chips per day as determined by Ju1ander (1955).
~Post-treatment data for all units have been adjusted for differential.cattle stocking rates which existed during 1970 and 1973.
Data have also been adjusted for differences in stocking rates between two cattle allotments which comprise the study area. Units
1, 5, 6, 7,8 and a portion of 3 are in Buzzard allotment, and 2, 4 and the remainder of 3 are in Porter allotment.
'#Data for chained areas w~re not averaged. High creek water and difficulty for cattle in findinqhUQit 3 caused the absence of
cattle in Unit 3 during 1973, even though the area had been substantially improved forcat~ e by C alning.

I

co
W
I

�-84-

One year before burning (July, 1970)

One year after burning (July, 1972)

Fig. 1. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1 and 2 years after burning
(Fig. continued on next page).

�-85-

Two years after burning (July, 1973)

Fig. 1. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1 and 2 years after burning.

�-86-

One year before spraying (July, 1970)

One year after spraying (July, 1972)
Fig. 2. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1 and 2 years after spraying
(Fig. continued on next page).

�-87-

Two years after spraying (July, 1973)

Fig. 2. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1 and 2 years after spraying.

�-88-

One year before chaining (July, 1970)

One year after chaining (July, 1972)

Fig. 3. Vegetation conditions at a permanent photo point on the Hightower
mountain oak study area 1 year before and 1 and 2 years after chaining
(Fig. continued on next page).

�-89-

Two years after chaining (July, 1973)

Fig. 3. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1 and 2 years after chaining.

�-90-

Fig. 4. Section of one burn where Agropyron intermedium and Bromus
inermis began to become established two years after burning (July, 1973).

Fig. 5. Most effective burn obtained in burned units. Vegetation
almost completely burned with just a few dead snags left standing.

was

�-91-

Fig. 6. Least effective burn obtained in burned units.
Only part of the
understory was removed with little or no damage to larger oaks.

Fig. 7. On much of the burned area most of the understory
with partial destruction of the overstory.

was removed

�-92-

Fig. 8. The overall result of burning was creation of openings interspersed
with patches of standing oak vegetation suitable for escape cover and shade.

�-93July 1975
JOB FINAL REPORT

State of
Project

C=..O::.;L=-O::.;RAD=:..:O=------

5

Work Plan No.
Job Title:
Period

Bighorn

Covered:

Personnel:

Game Range Investigations

W-lOl-R-17

No.

Job No.

1

Sheep Range Inventory

April 1, 1974 through March

31, 1975

Harold R. Shepherd

ABSTRACT
A final draft has been completed and is now in the editing stages by Dr. O. B.
Cope, and will be published in this segment as a Miscellaneous Report to Federal
Aid Project W-lOl-R-17.

��-95July, 1975
JOB PROGRESS REPORT

State of

~CO~LO~RAD~~O~

Projec.t No.

W-lOl-R-17

_
Game Range Investigations
Job No.

6

Work Plan No.
Job Title

Game Management Unit Inventory

Period Covered:

March 1, 1974 to September 30, 1974

Personnel:

l

_

William T. McKean and Paul H. Neil

ABSTRACT
Broad descriptive and wildlife species management information were compiled
for Wildlife Management Unit 21 (Douglas Creek). Subjects discussed included; Unit Description (Boundaries, Size, Physical Features and Climate);
Landownership Status; Land Use Status;"Human Populations (by County); Wildlife Species Checklists; Hunting Pressure, Harvests and Seasons; Narrative
and Map Descriptions of Game Species Distribution and Abundance; Introductions
of Game Species; Census Areas and Routes; Habitat Restoration Projects;
Management Problems Checklist; and Pertinent Research References by Game
Species and Author.

��-97-

GAME MANAGEMENT

UNIT INVENTORY

William T. McKean and Paul H. Neil

P. S. OBJECTIVE
To compile broad descriptive and wildlife species management
by wildlife management unit.

information

SEGMENT OBJECTIVES
1.

Describe Wildlife Management Units by name, number, boundaries,
physical features, and climate.

size,

2.

Inventory by unit the cultural and physical characteristics including
landownership status, land use status, and human population.

3.

Inventory by unit the.wildlife characteristics including species lists,
distribution and abundance, harvest data, introductions and/or past
species records, census areas and routes, and research and/or management studies bibliographies.

4.

Inventory habitat (range) characteristics of vegetation types, vegetation condition and trend, restoration projects, and transect records,
.all from existing sources,if available.
.

5.

Inventory present and future management

6.

Test application of any item of information compiled in steps 1-5
above in the computerized map project now in operation (Division Planning
Services activity).

problems.

METHODS AND MATERIALS
Five steps were necessary to produce the desired information about each
Wildlife Unit.
1.

The priorities of units were determined, based upon recommendations
field and administrative personnel.

of

2.

Working with sources of information in Fort Collins and Denver, as much
data as possible concerning each item were assembled and condensed.

3.

First-draft copies of descriptive material were submitted to appropriate
Northwest Regional personnel individually and to knowledgeable Denver
office persons for review and correction.

4.

After corrections were entered into the first draft, group meetings were
arranged with field personnel and others to make pencilled corrections on
the l/2"/mile game distribution maps. Further refinements to the narrative portions of the inventory were added.

�-98-

5.

Final revisions to maps and narratives were made and all information
gathere~ into file binder folders, except for the 1/2"/mile distribution maps which were reproduced by hand and bound in a separate
folder. Three copies of all this information were prepared and
personally delivered to: 1) the Northwest Regional office, 2) the
Denver office, 3) the Federal Aid section. Distribution maps at 1/4"/
mile scale were prepared on Forest Service base maps cut to 8"xlI " size.
Xeroxed copies of these were inserted into the file binder folders.
Addit10nal xeroxed copies of these maps and of the narrative were made.
These were delivered to the local W.C.O. 's, Bureau of Land Management,
and Forest Service personnel. Many additional copies have been
xeroxed for general distribution.

RESULTS AND DISCUSSION
Having completed write-ups for Wildlife Management Units 31 and 32 during
the 16th segment, there remained only Unit 21 (Douglas Creek) to be completed
as specified under the Job Description for Segment 17. This was completed
by September 1974. APPENDIX A is a copy of the narrative and distribution
maps as submitted. An extensive bibliography was included with all Division
copies, but is deleted from this progress report to save space and paper.
Again, no information was obtained concerning habitat (range) characteristics
of vegetation types (Objective 4). The "Land Use Status" tabulation is
essentially a listing by type. It lacks data on densities, species composition, condition and trend, etc. There were none available.
Data gathered under this project should be useful in the pilot computerized
map project now in operation by the Division Planning Section [Objective 6)
but this has not been realized as yet.
There have been numerous requests for copies of the six Wildlife Management
Unit Descriptions thus far prepared. As of this reporting time, requests
have been received from the following agencies or groups: Private consultants
and corporations - 40; Bureau of Land Management - 17; Soil Conservation
Service - 17; Colorado Division of Wildlife (other than Unit Area personnel) _
14; Fish and Wildlife Service - 8; c.s. U. Coop. Wildlife Unit - 6; Forest
Service - 4; C.S.U. library - 1; Others - 3; Total - 110.
\.

Prepared by

-,

~:t:21t~
William T. McKean
Wildlife Researcher

�-99-

APPENDIX

WILDLIFE MANAGEMENT

UNIT 21 (DOUGLAS)

Rio Blanco and Garfield

Information

Counties,

on Unit Description,

Human Population;

Wildlife

and Seasons; Narrative
and Abundance;
Management

A

Landownership,

Species Checklists;

and Map Descriptions

Land Use,
Harvests

of Distribution

Census Areas; Habitat Restoration

Problems Checklist;

Compiled by:

Colorado

References.

W. T. McKean and P. H. Neil

To:

Projects;
August 1974.

��-101-

Item
U;UT DESCjnpT[():~

(Boundaries,

Page

s Lz c , physical

features

and

1

climate)

LlJmOli')JI RSEI l' STATUS --- -.-----------------------------------_.___________

4

LAND USE STATUS -----------------------------------___________________

5

1m'IAN

POPUL\TION

GN·lE SPECIES

(Rio

Blanco

(Big game mammals,

OTHER l·fJ\H:·;ALL\I"~SPECIES
mamma l.s )

County,
small

(Furbearers,

1960-1970

and

game mammals
"varmint:"

projections)
and

game

mammals,

("Varmint"

nongame

Ai])

,!:&gt;irds,

nongame

birds,

11

raptores)
elk,

DIS'l'Klhul1uN
iL.'i.Ll
iiliUi~.0ANCE or. Sl-1..:\1..L GANE lHIZ1:;
necked
phe a sanr , band-tailed
pigeon,
mourning
HARVEST, SEASONS, MTD HlTh'TING PRESSURE

ARCHEWt

SEASONS, AND h1J't-tlING PRESSURE

BLACK BE.l\R lLWVEST AND SEASONS (1955-1973)

(1956-1973)

------------___

jj\Ti-:;ODUCTW:;S

OF G./\!'iE SPECIES

34

38
40

42
44

game rnan ag cmon r Un:i,!.
1+6
B1a.'1cO County,

------------------------------------___

i

-------

---------------------____

IL\.RVEST Al:D HlHITING PRESSURE (Rio
------------------------------------------------------_____

A.:;~) COllS;:

27

36

-------------------------__

LIO;~ HARVEST AND SEASO~~S (1965-1973)

195ft -1973)

24

-----------------------___

S:-:ALL GNfE 1l,\1zVEST pliDifCNTING
PRESSURE (Small
§_, 19 ()8-·.l9 j:5) ----------------------------------------------_________
Dl!CK

rabbit,

-----------___

ARCHERY ELK HJ\RVEST, SEASONS, Al'.TDHl}NTING PRESSURE (1956-1973)

Nom':TAf1':

23

(Grouse,
chuk ar , ringdove.
w-aterfowl)
-____

(1956-1973)

DEER HAi,\VEST AND SEASONS (1956-1973)

ELK Hl'JWEST,

18

ABUND•.
\NCE OF 1-111D HORSE ---------------------------___

DISTRIBUTION A::-';D ADUNDfu~CE OF Si:L\LL Gk'1E MAI1HALS (Cottontail
snowshoe
hare,
red (pine)
squirrc1)
----------------------------____

DZER

9

~_._

DISTRIBUTION AND ABUNDANCE OF BIG GAt'-'lEHAl&gt;["fALS (Black
bear,
rnoun t a in lion,
inu Le deer)
----------------------_____________________
DISTRIBUTION

7

birds)

---.-.----------------------------------------------

OTHER AVIAN SPECIES

6

--____

47
48

�-102-

ltUTI

- - - - - - - - - - - - - - - - - - - - - - 50
HAl&gt;ITAT

RESTOR\.TIO:~

:-lA:';L\GE;-IUT
BIG

PIWBLPiS

PROJECTS

-

PESEiuZCII

GA.,'-fE: B lRD

REFERENCES

HN2'!!\L

FURBE!\RER

-

RESEARCH

(;Eln::RAL AVIAS REFERENCES

REFERENCES

__

-

REFERENCES

RESEARCH REFER.ENCES

-

-

-

-

-

-

-

-

-

-

52

-

- - - - - - - - - - - - - - - - - - 54

HEFEREI'l'CES

S~'fALL Ci-]·rE WIl-L':AL

NmGP':·Ir:.:

-

CHECKLIST

GX-rt: l·U~{:-IAL RESEll-Rea

I;.r:SEARCH

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

57
-

_ _ _ _ _ _

_

70

- - - - - - - - - - - - -------

71

- - - - - - - - - - - --------

81

- - - - - - - - - - - - - - - - - - - 82

- - - - - - - - - - - - - - - - - - - - - - 85
- - - - - - - - - - - - - - - - - - - 86

ii

�-103-

•
DESCRIPTION

Boundaries

JJ.

- HILDLIFE H:INAGEHENT UNIT

21

(DOUGlAS)

1/

"That portion of Rio Blanco County south of the lfuite River

from the Colorado-Utah

State line east to Boise Creek, west of the divide
/.

between

the Yellow-Piceance

Creeks and the Douglas Creek drainages.from

Boise Creel: to the Garfield County line; that portion of Garfield
lying within the Evacuation

County

Creek and Douglas Creek drainages."
o

Size. -- 865.6 square miles.

Physi~.al Featm:es.--I.Jildlife Unit 21 corresponds
the Douglas Creek basin.

to what geologists

A major tributary of the \fuite River,

Creek drainage is underlain by rocks with wide differences
to erosion.

This has resulted in the more resistant

call

the Douglas

in resistance

rock layers of the

Green River Formation standing in relief while broad open valleys have
been cut in the softer rocks of the Wasatch l:lHlllation.

The Ncsa Verde Group of the Mesozoic occurs across broad areas of Douglas
Creek basin, consisting mainly of sandstone
and several coal beds.

w i.t h

much interbedded

These rocks are usually resistant

often form prominent mesas, ridges, and cliffs.

shale

to erosion and

The thickness

of Nesa

Verde ranges from 2,500 feet to 5,600 feet (U.S.D.A. 1966).

1/ Lega Lly t e ru.e d "Big Game }~anagement Unit (see footnote 2/ b e Low) but
'lcntatively called "l.Jildlifc~lan;)gementUnit" here to incl~de broader
aspects of animal life present.

?)

Colorado Div. Hildlife Laws and Regulations

Bi.g c.?:,,(~).

1

Hdbk., 1973 (p.S, Chap. 2 -

�-104-

Quoting.
out

U.S.D.A.

around

mainly

the mar g Lns of

of red,

These. rocks
valleys

1966 further,

purple,

are

the

Piceance

and gray

relatively

or badland

"11lC Hasatch

areas

shale

erodible

Formation,

which

also

Creek. basin

(Douglas

and clay

some sandstone

and this

be twe.en cliffs

with

crops

Creek),

fo rmat Lon generally

consists
layers.

forms broad

or

ridges

of the

Green River

valuable

oil

shales,

is

Formation

and the Hesa Verde Group."

The Green River
the

southern

Roan Plateau

Unit

its

found

only

around

This

caps

the

Creek

Divide

on the west.

a site

a few miles

below

Rangeley

on the

divide

periphery

of Douglas

on the

and the

Evacuation-Douglas

south

range
River

from 5,224

feet

upwar d to 9,090

West Fork of Douglas

Other

with

and western

elevations

on. the Ifnite

Formation,

Creek near

thr n '1'~~p D~~!!;las Creek,

at

feet

Douglas

Pass.

with

east~n0

its

drainages

i.n Wildlife

Unit

21 are:

Cathedral

Creek - all

cast

of main Douglas

draining

into

Creek arc

Horse

DraH; draining

Big Trujillo
west

are

vfuiskcy

Soils

Douglas

Shavetail

Park Canyon,

Fletcher

north

into

wes t branches

Gulch,
Creek;

Lower Horse

be twe en East

Spring
to

, the

Creek

the 'vest

Draw, Four Hile
are

Wash and Cottonwood

Creek;

draining

Cr~ck,

and

Draw and Big

River

Missouri

maIn

and also

the h'hite

Texas Creek,

Fork and

Johnson

Evacuation

Draw,
to

Creek,

the
and

Creek.

data

Sorv ice

Hash,

to the

Creek Basin.

for

Unit

Work Unit

21 can be obtained

p er's onne L located

b roo d soil

c Las s Lf i c a t i on s also

lkp:1rtr':ent

of Agriculture,

1966.

at

by consulting
Meeker.

can be found

Soil

Descriptions

in previously

\·.'ater and related

2

U.S.

land

Conservation
and a map of

cited:
resources,

U.S.
Whi.te

�-105River Basin in Colorado, Coop. Study Rep. of Co Lo ,, 'yater Conserve Board.
i!Ild

H.S.D.A. Econ. Res. Service. Forest Service, and Soil Conserve Service,

November.

Denver.

Climate.--The

92pp (Looseleaf).

climate of Unit 21 is definitely-semi-arid.

As in adjoining

areas, Unit 21 experiences wide extremes of precipitation

and temperature.

Hean annual precipitatio~ at Rangely, the only National Weather Station
in the unit, averaged 8.92 inches (1951-1972).
3.76 to 17.5 inches.

PreCipitation

ranged from

During the period 1930-1940 inclUSive, a mean annual

precipitation of 10.36 inches was r2corded.

Data are lacking for the forties.

Precipitation on Roan Plateau at the head of Douglas Creek is estimated at
25 inches annually.

Cloudburst storms of high intensity a~d short duration

occur over small areas and result from convective conditions.

Seasonal

distribution of precipitation is fairly uniform, higher elevations

receiving

a greater proportion in winter in the form of snow.

Temperature records are available for fifteen years of the twenty two-year
period 1951-1972.

The mean annual temperature for those years was 46.33

degrees, ranging from 44.0 to 49.8.

Frost free days recorded for the

period 1961-1972 averaged 120, ranging from 66 to 157.

Literature Cited
U.S. Department of Agriculture.

1966.

White River Basin in Colorado.

Water and· related land resources,

Coop Study by Colo. '-laterConserve

Board and H.S.D.A. Econ. Res. SerVice, Forest SerVice, and Soil Conserve
Service.

November.

Denver.

92 pp. (processed).

U.S. Department of Commerce. (Continuing).
HOllthly and Annual Summaries.

Climatological

data.

Colorado.

National Climatic Center, Fed. Bldg.

AsheVille, N.C. 28801

J

H. T. HcKeao
Hardl, 197/1

�-106-

LANDQl..7NERSIlTP
STATUS - HILDLIFE

HA.i.~GEHENT maT

21

Acres

.

"Bureau of Land HanagetJent

484,382

Private Lands

74,892

and Cou.nty Lands

13

Other Federal lands including National
Park Service, Forest Service. Indian
and Hi1itary Reservations and Bureau
of Sport Fisheries and Wildlife

o

Municipal

Co lorado Division

2,710

of 'Hildlife Lands

State L&amp;nd Board Administration

o

Lands

* Total

561,997
(878.12 sq. miles)

*Total Land acreage differs from teta1 in J1E:i-t pesc~~E-tl,~!::._
.~~ct~on (S5l

f,
026
or 856.6 square miles).
Bureau of Land Management lands and private lands
were derived from 1/2" = 1 mile B.L.M. planning unit base map (1965).

P. H. Neil
April 1974

I
"T'

�-107lAND USE AND VEGETATIVE
\\'JLDLIFE

COVER

HANAGENENT UNIT

Irrigated cropland
./

.

21

5,432 Acres

Craas Land with half-shrub mixtures

29,193 Acres

Sagehrush

73,994 Acres

BrUE;h (desert and mountain)

123,280 Acres

Woodland

257,181 Acres

(Pinon-juniper and oakbrush)

Commercial

timber to include aspen

River bottom vegetation

40,568 Acres
1,356 Acres

Urban

446 Acres

Miscellaneous

22,576 Acres

.

Total

554,026 AC1.eS *
(865.6 sq. miles)

*Cover type categories we re derived from 1/'2" = 1 mile S. c. S. Land Use
and Cover Type Haps, 1953, for Rio Blanco and Garfield counties. A
planimeter and grid were used to derive and compute acreages ..

1/ Urban - Those areas used for residential and municipal purposes as
illustrated on the 1/2" = 1 mile B.L.M. planning unit base map for the
town of Rongely.

2.:/

Hiscellaneous - Interpreted to consist mainly of bare ground with
sparse, scattered vegetation of various types.

P. H. Neil
April 1974

5

�-108Hl.NAN POPITLAT10N

-

I.JILDLIFE HANACnmNT UNIT 21

The rural popu1ation of Rio Blanco County has decreased from .62 to .51
persons per square mile b:::twcen1960 and 19711. This trend probably tipplies
to the Unit 7.1 porti.on of it, though temporary increases have been known to
occur because of 011, gas, and oil shale exploration. l/
Table 1.

Urban-rural

population,

He.eker
Rnnge1y
Rural
TOTAL

Rio Blanc~ County,

1960-70.

1960

1970

1,655
1,464
_2,031_
5,150

1,597
1,591
1,654
4,842

5,150

3,135

2,980
~943

1,690

1900
'Fig. 1.

1910

Population

One projection
1980 - 6,375,

1920

1930

1940

1950

1960

1970

trend of Rio Blanco County.

of human population

J:..I for Rio Blanco

County is: 1975 - 6,018;

Another projection of human population 11 for the White River Basin is: 1980 _
20,000; 2000 - 186,000; 2020 - 204,000.
A third projection !!..I for the oil shale area, based upon cumulative oil
production in barrels per day, is: 1974 - 6,631; 1980 - 46,738; 1985 - 100,000.

II
- U. S. Bureau of the Census and Colorado Yearbook,

1959-61.

2/
- Colorado Slate P1anning Office, advance population

projections,

]/ u. S· Burcau of the Census, Colorado Hater Conservation

Eoard

unpublished.
and

USDA Field

Party.

!!../u.

S. ])',&gt;01:. of Interior £1972)
Draft cnv Lr onrnent aI statement for the propo scd p ro t o t vne oil s h a.l e c:jsill;~ pro/.:rilf;l.
·Vol. To hTasldngton,
D.C, '+65 p ,

6

�-109GAJ'lE SPECIES - HILDLIFE

Big game ma~~als

~L\NAGEHENT UNIT

21

II

Black bear (QE.~t1S_ ~~~ricanus) Uncommon ,
Elk (CCT'TJS can adcnsLs) Unc or.unon ,
Hount:~'1-ill
J_ion(l~hs
--Zoncolor) Uncommon.
Mule deer (Odor:ilie::;-s
hC:!li~nus)Commo~.
Small game mammal.s II
Cottontail rabbit (SylvHag ,:!s.§.ud1.lbonii_;
~. n1.lttal~i.i)
Common.
Pine (nod) squirrel (TamiascJ.~!.:.~'s
hudsonicus) Common.
SUOv,1S11Oe hare
(Lepus ame ricanu s) Common •
4

.3mall game birds ]j
Migratory vat e rf'owl, and shorebirds
Great Basin Canad::igoose (Brant a canadensis moffitti) Uncommon.
Black brant (J!ran_~~T1.igrican-;)-ilPossibl-;-rare mig~~ant.
White-fronted goose (Rranta albifrons frontalis) 31 Possible rare migrant.
Snow goose (Q1en E~2ruJ.escC;n-s~
caerulescc~s) )j, !!l. Possible rare migrant.
Whistlinz S\o1an(Olor coluf.lbinnl..'.s)
Rare migrant.
Mallard (Arias.£.~&lt;1tYEEynchosPl?.!::'rhvnchos)Common resident.
Cadwa l.L (.:6:::i~_::: E~;-ep_~~) Common spring_ and fall migrant.

1/.

- Nomenclature according to Lechleitner~ R. R. 1969.
Colorado. Pruett Publishing Co., Bould~r.
254 pp.

Wild mammals

of

]j
Nomenclature from Bailey, A. M., and R. J. Niedrach.
1967. Pictorial
checklist of Colorado birds. Denver Hus. Nat. Hist. 168 pp. Information on
occurrence and status adapted from the above reference and Cringan, A. T.,
and L. C'etrlson.1973.
Wildlife in the Piceance Creek Basin, In: An
environmental reconnaissance oi the Piceance Basin, Rio Blanco and Garfield
counties, Colorado. A report on the completion of Part 1, Phase One of the
environmental inventory, analysis and impact study portion of the Regional
Oil Shale Study being done for the State of Colorado by the Thorne Ecological Instulite, Boulder, Colorado, 144 pp. Additional information on
occurrence, in employing the term "possible", is adapt:ed from the foregoing
refercnc es and Davis, loJ. A. 1969. Birds in we s t ern Colorado.
Colo. Field
O'rn Ltho l.ogist s , 61 pp . "There adjective "possible" is absent, actual
sightings have heen reported verbally by anyone or more Division personnel
Glenn E. Rogers, CLaude E. Hhite, George E. Steele. Louis D. Vidakovich, or
qualified by additional footnotes that follo\o1.

,..,
( -

�-110G&lt;:meblrds

(continued)

Pintail
(An~~~ ·:l~..t.::~,:3) Common spring
and fall
migrant.
American green-Hinged
teal
(/~1C1~ crecca
c:&lt;::rolinensis)
Common migrant
and un common year Lon~ resident
4/
Blue-Hinged
teal
(An a s - d Ls cors di.~ors)
Common migrant.
Cinnamon teal
(An~'-~:;--~'1D(lDtcr&lt;;sentcntr:i.onalium)
Common migrant •
.American w I geon -U~~-~-;-;;~~~:lo.)4/
Common migrant
and rare winter
resident.
No r t he rn shoveler
(J-.no-;' cl~jX~~~a)~4j
Common migrant
and uncommon summer
r es Ldent .
Redhend (;\ythya a:nericana)
Uncommon migrant.
Ring-neck_Q'(r--cf~ck(:\vti~~~
£olb!:..is)
Uncommon migrant.
Canvasback
(Avt hva vali.sinoyjti)
Unc ommon to rare migrant.
Greater
scau-p~;)ya
m;rila
n-carctica)
3/ Rare migrant.
Le s s e r scaup
(~'thya
affini~)
Un c orrrmon migrant.
Common golden-eye
(Buceohala
clangula
americana)
Common migrant
winter
resident.
Barro"r's
golden-eye
(Bucc_,:?hal&lt;l:.islandica)
)../ Rare Hinter
visitor.
Bufflehead
(~uceohala
a1beola)
Uncommon spring
and fall
migrant
rare winter
resident.
Ruddy duck (~):-:vur..!:!:.
j amaic(~nsis
summer resident.

rubida)

Common migrant

Hooded merganser
(~~ophodytes., cucullatus)
Rare
Common merganser
(;~.2~ mergcl!1ser americanus)
CO~~on SWlliuer resident.

and

and

and occasional

winter
visitor
Common ''linter

on river.
resident,

un-

Red-breasted
merg,1User (Men-;us serra tor serrator)
Uncommon winter
resident.
American coot (Fuljca
'americana
americana)
Common migrant
and summer
resident.
Common \';i1son'

s snipe

(Capella

.8allinago

de1icata)

Common migrant

and rare

Sandhill
crane
(Grus canadensis
canadensis)
Regular
migrant.
Vh'ginia
rail
(Ralus limico1a
limicola)
Possible
uncommon summer resident.
Sora (Po rz ana c:;U:-OTina) Possible
uncommon summer resident.

Upland

game birds

Blue grouse
(Dendri-lr;apus obscurus
obscurus)
Common.
Sage grouse
(~entrocercus
uroohasianus
urophasianus)
Uncommon to
Ring-necked
pheasant
(Phasianus
colchicus)
Uncommon.
Chukar (AlectoY.'is chukar)
i/ Uncommon.
Band-tailed
and fall
Mourning

)'/Unverified

pigeon
migrant.
dove

(Columba

(Zenaida

in hunters'

fasciata

macroura

bag checks·

fasciata)

legal

uncommon summer

if Common summer resident.

margine1la)

but

Possible

common.

game

1972-73.

ljf

- Ckmges in nomenclature
f oLl.ow the thirty-second
AIDerLean Ornithologists
Union check-lis
t of North
in Auk 90:411-419, April,
1973.

supplement
to the
American birds
published

W. T. HcKc&gt;an
1974

• June

8

�-111OTHER tfN-["1i\I,IAN SPECIES
Furbec re rs

II - w'ILDLIFE HANAG8-fENT

UNIT 21

2-../

Short-haired
Beaver (C:l~~.or ~&lt;!E.&lt;J.d.?!~sis)Common.
Mink (Muslcla vison)
Uncertain.
Muskrat"- (.0~G~6a Z:LbCt:.licllS) Unc ommon,
Ringtail
(Bassariscus
25tutllS)
Rare.
'l-lcasels O·;~~tcl~·
el':;i~e'~- 2i,.-frenata)
M. enninea
Uncommon.
Black-footed

ferret

(H&lt;l~tela

nigri~)

Possible

Uncertain;
rare

resident.

M.

frenata
Endangered.

Loug-ha Lred
Kit fox (.Y2-:!.:..~ ve102:) Uncommon.
Gray fox (.ll_~!?cyo~ci~~~~rgen_tetl~)
Uncommon.
American badger
(Taxice.a t3XU8) Corr.mon to uncommon.
Spotted
skunk (~~31e
~_oriu~)
Uncommon.
Striped
skunk Cl'lephitis mephitis)
Common.

"Vannint"

mammals

Coyote (Canis latrans)
Common.
Red fox (Vu Lpe s ful V2) Unc ormnon,
R2'~C)C"i1 (:rT·:~'(:-~~;~~"1
:Gtor) Unc ommon .
Porcupine
([r~izon
dorsatum)
Common.
Bobcat (\vildcat)
(Lynx rufus)
CODInOn.
W'nite-tailed
jack ~it
(Lenus tov.'l1sendii)
Common.
Ye Ll.ow-be Ll.Led mannot (Narmo t a flaviventris)
Common.
White-tailed
prairie
dog (Cvno;:IV8 Le uc uru s ) Uncommon.
Richardson's
ground squirrel
(Spe~onhilus
richardsonii)
Common.
Thirteen-lined
ground squirrel
(Snermonnilus
tridc.cemlineatus)
Common.
Rock squirrel
(Sper:::nopililus varie~atus)
Cornmon ,
Horthern. poc:ket gopher (Thomomvs talpoides)
Common.

!lThese
species,
grouped separately
as "Furbearers",
"Varmints"
and "Nongame
mammals" and out s i de of "game categories,
follow Chapter
62, Colo. Rev. Statutes
1963 As Amended, in Colo. Game, Fish and Parks Div. Laws and Regulations
Hdbk.,
1973 (Art. 1, Items 13, 17, and 18, Defini:::ions,
p , 3).

'!:../
Nomenclature
from Lechleitner,
R. R. 1969.
Wild mamma Ls of Colorado.
Pruett
Publishing
Co , , Boulder.
254 pp.
Information
on occurrence
and status
from
the abo ve r c f e r e n ce 3:1d: Cr Lngan , A. T., and L. Carlson.
1973.
Wildlife
in the
Pf.ccanc c Cr co k Basin,
In_: An cnv fr cnrnen t aj, reconnaissance
of the Piceance
Basin,
Rio Inaneo i!!lll Garfield
counties,
Colorado.
A report
on the completion
of Part
1, Ph a s e One of the env Lr onrnanr n I inventory,
analysis
and impact study portion
of the, ne~~i0;;.11OLl Sh31e Study being d ccnc for the State
of Co Lo'rad o by t he
Tho r nc Ecolo;~ic.1J. Institute,
Boulder,
Colorado,
144 pp.
Al so , Armstrong,
D. H.
1972, Dif:ldl:uliG'l
of mamrnaLs in Colorado.
:'!onograph of the ~luseum of Natural
History,
the Univ. of Kans a s , Numbcr 3, 1972.
1,15 pp.

9.

�-112-

Nongame mammals
Co Lderr-rnan t Lcd ground squ:Lrrcl
(Sr.,err.~QPMlus latcralis)
COnll'TIOn.
l-;rhite-tailed
antelope
squirrel
U-Tf:1;w..:.~cnnophilusleucurus)
Uncommon.
Leas t ch i praunk (}':ut:0:nias m:inimus) Corrrnon,
CoLc rado ch i pn.un k (~1l:ami;'2 oU2drivj~_~atus)
Common.
Uinl:a ch Lpraunk
(Eut arm.as u.cb r inus
) Un common
to uncertain.
Vagrant sh r ew (S;-re;.:--~;;;-r;;s)
Uncommon:
Herriams shrew (so~-;x merriani)
1/ Uncertain.

Townsend! s b i.g+ea rcd bat (Plecotus
t ownscnd Lf ) Common.
Silver-hairerl·
bat (Lasionv~~is
noct'~'lg,ms)
Common not abundant.
Hoary bat (Lao i u ru s cinereus)
Uncommon - common.
Big b rown h-at (i::;~i~stcus
fu~S~,§,) Cornmon,
Hestcrn
p Lpi s t r e L'le (~..P2:strcl1us
h~sperus)
Common.
Long-legged
mvo t i.s C'fyotis voLJ.~:~) Uncommon.
Ca Ldfo rn La myotis
Q'hrC2..!is~alifornicus)
Common - not abundant.
Small--footed
myotis
(Hyotis Lc LbLf.) C01JIllOn.
Long+e ar cd myotis
(Hyo t i.s evotj~)
Uncertain.
Little
bro •..
'11 myotis
(Hyotis lucifugus)
Uncertain.
Ord's

kangaroo

rat

(Dipodomys

ordii)

Unconnnon - Uncertain.

\\Testern harvest
mouse (Reithrodontom~
magalolis)
Uncertain.
Canyon mouse (Pe r omvs cus crinitus)
Common - unc ommon •
Deer mouse (Pero~~\:s~CllSmaniculatus)
Common.
Pinon mouse (Pe·.Lo;ys~-;:;:-struei)
Comrnon ,
Bushy-t a Ll.ed c'!ood rat(Neoto.~~ine~f;:.a)
Common.
Gapper's
red-backed
vole (Clethrionomys
gapperi)
HeadovJ vole (Hi c:rotus pennsy]vanicus)
Uncertain.
Mont:;;.ne vole G:!)crotu2 montanus)
Uncertain.
Long-tailed
vole (Hicrotus
lonr~icaudus)
Common.
Sagebrush vole (~curtatus)
Uncertain.
House mouse (~~u~~usculus)
Uncommon.
Western jumping mouse (Zapus princeps)

Uncommon.

Uncommon.

3/
- Occurrence
listed
by the Colo. Div. of Wildlife
as extremely
unusual
_
very f ew xlo cumen t e d records
wi.th Ln the past decade.
(1972 Status
Evaluation
for Selected
Colorado Species)
appended to: 1973 Wildlife
Operations
Work
Plan, Field Order No.4
- 1973).

W. T. McKean
June

10

1974

�-113OTHERAiJIAN SPECIES 1./
"Varmint"

- HILDLIFE Hl\.1'jAGE11Ei\T UNIT

21

birds

lHack-·billed
I,;:lgpie (Pic;:! l~ica hudsonii0
Common resident.
Starling
(StU1:-..Q~1'; .vulgar:i.s)
Common resident.
Nongame birds

2:../

COD"UllOn
loon (I.r~v:i::tJ8i'1e]J Possible
rare migrant.
Horned grebe
(~&gt;()dic:.~_~ ~_~.tus
cornutus)
Possible
rare migrant.
Eared g r cbe (}~C251}c~~ .r:_:ij:2J~~ollis ~a~ifon!icus)
Possible
common migrant
1/.
Western grebe (£,:.~:cb'!~::"C2.~)ho"£~5
.occidentaJ.is)
Possible
rare migrant.
Pied-billed
gre:)(~ (i:\?-,"hh~~?\lspodiceJ2£ ~diceps)
Possible
uncommon migrant
and rare
SFc~~:fler
resident:.
Double-crested
cormorant
(!'halacrocorax
auritus
auritus)
Possible
rare
migrant.
Great blue heron (Ardea herodias
treganzai)
Common summer resident.
Snowy egret
(Lgrett~'9. thu10~teri)
Uncommon surruner r es Ldcn t ;}j.
B'l ack+c rown e d nLgh t; heron
(Nvc t Lcor ax nycticorax
hoactli)
Possible
common
summe r resident.
Least bittern
(Ixobrvchus
e:-:i1is exilis)
Possible
rare summer migrant.
American bi tter';'- (.iis.~ur~j_~-le_ntiginosus)
Possible
rare summer migrant.
Hhite-faced
ibis
(PJ.egadi2 c!!:.~bi) Possible
rare migrant.
i-lhistling
swan (Olor co Lumbi.anus ) Uncommon migrant.
Semipalmated
plover
(Cha~lclrius
semi.pal.mat u s ) Possible
rare migrant.
Ki11deer
(Ch a r ad iu s vocifcrus
vociferus)
Corrunonsummer resident
and rare
wiater
resident.
Mountain plover
(~h,~xa4rj \1..~montanus)
Possible
rare migrant
;}./.
Elu~k··:':2:1icJ p l ove r (Plu'li::..lis
s::n:at2i:o:'.:::) Possible
uncommon mi[;-;:'2::t '&gt;./
Long-billed
curlew
(Nurneni.us americanus
amer Lcanus ) Possible
rare mf gran t ,
Spotted
s andp i pe r (~~~m~';:-L:~)
Possible
common summer- resident.
Solitary
sandpipe.r
(Tring.Q. solitClria
cinnamomea) Possible
common ruigrant
and occasional
summer resident.

1/
- The.se species,
grouped separntely
as "Varmint" birds,
"Nongame birds"
and
"Rap t or cs " and outside
of "g ame" categories,
fol1m .• Chapter
62, Colo. Rev.
Statutes
1962 As Amended, in Colo. Game, Fish and Parks Div. Laws and Regulations Hdbk., 1973.
(Art. 1, Items 13, 18, ~nd 15, Definitions,
p. 327).

2/
- Nomenclature
from Bailey,
A. M., and R. J. Niedrach.
1967.
Pictorial
checklist
of Colorado birds.
Denver !'lus. Nat. Hist.
168 pp ,
Information
on occurrence arid status
adrip t e d from the above reference
and Cringan,
A. T., and L.
Carlson.
1973.
Vi.ld1::fe in the Piceance
Creek Basin,
In: An environmental
reconnaissance
of the Piceance
Basin, Rio Blanco and Garfield
counties,
Colorado.
A report
on the c or.ipl e t Lo n of Part 1, Phase one of the environmental
inventory,
analysis
and impact study portion
of the Regional Oil Shale Study being done for
the Stat ..2 of CoLor ado by the Thorne ExoJ.ogical
Institute,
Boulder,
CoLor ad o , 144
pp.
Add i ti.UI1:J.l Luf o rma t i on on occurrence,
in employing
the term "possible",
is
adopted f ro.n the foregoing
references
and Davis, H.. A.
1969.
Birds in wes t er n
Colo r ado . Colo. F:i.vlJ Ornithologists.
(,i pp,
Hhere adjective
"possible"
is
ab scn t , .ictun l ~;:if',htings
have be en reported
verbally
by anyone
or more Division
pc r s onn cI Clenn I:. ;~Ci~Cr!j, Claude E. \{id t c , Ceo r ge E. Steele,
Louis D.
Vid";,(lv.ic:h
or qu.i lLfio d Ly add it Lonn L [001:1:01:es that f ol l.ow, Notations
concc r r.i n-. ,;(:;l';l1Jl."il ;·bul\(.i&lt;1nc(&gt;
of r ap t o r e s a r c by Gerald R. Craig,
Hi1dlife
llioJ,);:,j~;t .

11

�-114Nongame b Lrd s (oont Ir.ucd)

,,~:alet

Lno rno t us ) Possible
rare mi.grant .
common migrant 1/·
Lesser ycl1m.;}.('./,;s
(T1.:".:~'22 fJ a-'~:~22)Possible
uncommon migrant ]j.
Red knot (Ca1i0r1.:&gt; c anur us r u ra ) Possible
rare migrant
3/.
Pe c t or a I. s-;·ndpiT~r.:-;: Tc:~iidris-;;::];motos)
Possible
rare mIgrant
3/.
Baird "s sandpiper
u.:8iT;-GTs·b~:-;-i-;ZCiiyrossible
common migrant
37.
Least ~:andpipc::r (~,-,LI0;=-j-; i~i~l~~~l:TlJQ.)
Possible
common migrant
}/.
Long-bj 11e d dowi t: cl.c r (I~jYmojEoml!s _sco 10D&lt;lCells) Pos s ib Le uncommon nu grarrt ,
Stilt
sandpiper
C:~:~S:£()Jl&lt;d~:l:1l~~Ii,,'ntorlUs) Possible
rare migrant.
Seminalmatcd
sandoiucr
(Calidris
pus ilIa)
Possible
rare migrant
3/.
Hest~rn sandpipe,.'· (Cc.l.:i dris-;;,wri)Possible
uncommon migrant
3/.-NarbJed godwit (L~1;·Q.S;fedo2.Y-Possible
rare spring migrant.
Sanderling
(Cali'::~;:i-~--:~Jb,3.)Possible
rare migrant
3/.
American ;rvocet --(:·Gc;u;.:;:-,-j~Y-ostTa
arne r i.cana ) Rare mi-grant.
Black-necked
stil-t
(H:;~::;:nto';\~~mex i canus ) Possible
rare migrant.
viilson' s pLalar0~;e (~_~cr:ano;)t~;;-rr:i-;;To-;;Y' Possible
C0UU11on
migrant
and
uncommon summer resident.
Northern phalarope
(Lo~i1?e:=; lobatus)
Possible
uncommon migrant.
Pomar i.ne j aeger U;tc:-.£~orar~~.2_POtlar:0,lUs) Possible
rare mi gr ant .
Herring gull (L2EuS ar"Jmta~'_~~ .:?Tlithsonir-il1us) Possible
uncommon migrant.
Ca l.Lf ornd a gull
(;'a;.u~ caHfoc:nicu~)
Possible
rare migrant.
Ring...,billec1 gull
(L2121S del~'Jar:2nsis)
Possible
uncommon migrant.
F'r ank.Li.n' s gull (L,~nls .p2:..J2i:5~2n) Possible
uncommon migrant.
Bonaparte's
gull
(La.!u,2 .£.l1i-G~~~c1nhia)Possible
rare migrant.
Sabine's
gull (X:::'1a sab i.nL s ab i.n i.) Possible
rare nigrant.
Forste.:r's
tern -rS'tcrna rorsteri)
Possible
rare migrant.
Common tern (?te1=-:.12:·hlr~Ddo h:i"~undo) Possible
rare migrant.
Least tere (Ster~3 a:b2frons 2thn13ssos)
Possible
rare migrant.
Black tern (Chi~"-l~~~i-i~tc;er
s ur i.narnens i.s ) Pos s LbLe rare migrant.
Rock dove (COl,U1j.&gt;.-:-C:.l:ivj
..£!-) Pc s s i.bLe common resident.
Unprotected.
Ye.Ll.ow-b i.Ll.ed cuckoo
(Coccvzus americanus
amer i canus ) Possible
uncommon
summer rcsidc~:t.
Poor-will
(P}lalae:1oDtilus
nuttallii
nuttallii)
Commonsummer resident.
Commonn i.ght hawk (C;~dciles
minor he spe r i.s ; ~. ~. howelli)
Common
SUiJI1Iler
r os i.de a t ,
White-throated
s~~£t (Aeronautes
saxatalis
sclateri)
Common summer resident.
Black-chinned
hurzrai ngb Lr d (Ar ch i Lochus alexandri)
Commonsummer resident.
Broad-tailed
hurru i.ngb Lr d (S~].&lt;:!sj)horus p:GtY~~s
platvcercus)
Common
summer r e s i dcn t ,
Rufous hummi.ngb i.r.d (Se Las pho rus rufus)
Possible
common late
summer migrant.
Calliope; hUr:1miTI[;oird-(Stellula
calliOpe)
Possible
rare migrant
and
summer resident.
Rivoli's
humminzbird
(~ugene.:? fulgcI1s aureo'liridis)
Possible
rare summer
visitor.
Belted kingfisher
(l-b~acer\'lc
a l.cvon alcvon) Possible
common resident.
Common fli~ker
(COLlDt(';-;t~;-i~[~s~l~s-)ColrJnon
resident
3/
(C. a. luteus)-});-;~-j~bie-r~~;e
migrant
s]:
Lew.i s t \,roodpccker U.svnJesT'lus le\"is)
Possible
common summer resident.
(C;ttOTltr0·.·,1~()nrs

[;&lt;.:mil);}lmalm-:

Grcnt cr )'(.'J.l;;i·~~;;--(}~~~~}iS~
:i~~~~;!~~~ie~ca)
-f-~~-i-ble

1/Changc s in ncrncn cl.c t.u r c follow
O'rn i t.ho Lo.t ist s Un Lou Checklist
411-419.
April,
1973-

the Thirty-second
of North American

12

Supp.l.crnen t to the American
HirJs published
in Auk 90(2):

�-115Non~;'J'lC.birds

(continued)

Ye l.Low+bc
l L'l c d sapauckc
r (~.E.:'::Yr;:\lYi.c:lIs
varills
nucha
lLae ) Common summer
and possible
occasional
winter
resident.
WillL~s.son' s s ap s u c ke r (Sj)hvriln_~~
£!::.Y~i.~~us nataliae)
Possible
c ommon
summer resident.
Hairy
woodpecker
(9_c:!.l_~,X()C~ vi] ~osus mon t Lco La ) Possible
common resident.
Downy wo odpcck e r (D0.n~_;;,oc~
pllbcscC!ns leucur_lls)
Possible
common resident.
Northern
t hr cc+t
ocd wo odpe
c ke
r
(P icoid&lt;:,s tridactvlus
dorsalis)
Possible
rare m:i,gra!:lt.
Eastern
kinghinl
(TY_:;-..0.:~'u~
~nus)
Possible
uncommon summer resident.
~·jestern kingbird
(1:"'0:i::.'IIl1~':. verticalis)
Common summer resident.
Cassin f s kin~bird
(~::mnus
::'2£iferans
vociferans)
Possible
uncommon
suume r resident.
Ash-throated
flycatcher
(Hviarchus
cinerascens
cinerascens)
Common summer
resident.
Say's phoebe (~~i2
~;aya _saya) Common summer and possible
rare winter
resident.
.
Hlllovl flycatcher
(Emp i.d on ax traillii)
Possible
uncommon summer resident.
Harnrnon d ' s flyC'.at:cher(L~nax'
hamrnondii) Possible
migrant
and uncommon
SUmrH'."f.'
resident.
.
Dusky flycatcher
(Empidonax oberholseri)
Possible
summer resident.
Gray f Ly ca t che r (E~-;-id~nax wr i.ch t Lf.) Possible
summer resident.
Weste.rn flycatcher
(~iclonilx
difficilis
hellmayri)
Cornman summer resident.
Hestern
Hood peewee (~gllt_t?J2.ussO,rclidulus veliei)
Common summer resident.
Oljve-sided
flycatcher
(Nut t a Ll.o rn i.s borealis)
Possible
COI!UT10n
summer resident.
Horned La rk (Ere!1'1:S2.E..i:~ila
l:l_~trjs
Lcuco Laerna ) Common resident.
Violet-green
sva Ll.ow
(Trachvcineta
thalassina
Leo i.d a ) Common summer resident.
Tree swa LLow
(l~_~~~~('ne
bicolor)
Common migr;;t
and summe
r resident.
Bank. 8\-13110\07 (Rina'r i.a E~'.!"L~ r~E2ri~)
Possible
uncormnon migrant
and
summei

1'&lt;=::; Ld en c ,

Rough-s ...•
in8ed SHallow (Stelgidopteryx
ruficollis
serripennis)
Uncommon
g
and sumner
i.den t ,
Barn swa L Lov (Hi.rundo Eustica
e rvt h rorras t e r ) Common summer resident.
Cliff
swallow (Petrochelidon
pvrrhonota
flvrrhonota)
Common summer resident.
Purple martin
(Prognc subis .?_ubi:~) Poss-ible
rare s~rmner migrant.
Gray Jay (.?_ed,sor~Jl~ c,madensis
,s:apitalis)
Possible
uncommon resident.
Steller's
jay (~vanClcitt"!: stellcri
TTl3crolopha) Common resident.
Scrub jay (Aphclocoma ~o,=--~lescens ,...•
oodho us e i I ) Common resident.
Common raven (Corvus c or ax s i.nua t us ) Cornmon . resident.
Pinon jay (Gyr:l;lOr;li;-tl-;'-;;'~nOCeDhalus)
COr.JIIlotJ.
SUTIUller3/ and winter
resident.
Clark's
nutcrDc kiOr C&lt;ucifra2.3. co l.uub i.ana ) Common resident.
Black-cDpped
chickadee
(Par~s arti.caDillus
garrinus)
Common resident.
Mountain chickadee
(Pa r us ;:a.l:'!beli z arab e Lf.) -Cor:-unon
resident.
PTa In titmouse
(Parl!; inorn:1tus
r-jd~";'3vi) Common resident.
Bush t I t (Ps&lt;11tri~;irus
rr;ini:;~':~'spl~~::1[~-2us)Possible
co:r:mon resident
3/.
\·Ildte-brc';-,st-ed nuthatcTl(Sitt.:l-c;roTii1ensis
nelsoni)
Possible
unc-;;-mmon
r e s f.dcn t ,
--Red-breasted
nuthatch
(Sitta
~anadensi.s)
Possible
rare resident.
Pygmy nu t ha t ch (S'i.t t a fly!~r:!Cle.il mclanot:Ls) Possible
unc ornmon resident.
Brown creeper
(~crtt~fi'-filmi]
iaris raon t an a ) Possible
uncommon resident
and
common migrant.
Dipper (~:-i~~:.:1_tJs
!T,ex:Lcal~ un !color)
Common resident.
House wr en (Ir()(,lo(~;ly.5~ &lt;1C~::il1 p a r kmau i i ) Possible
common summer resident.
mf

raut

r-e

s

13

�·

~ongame birds

-116-

(continued)

Bewick r S wren (!!L~:Z(J_~.an.£.:2.be~':lckii
resident
and rare winter
resident.
Long-billed
marsh wren
winter
resident.

(Telrnatodvtes

Canyon vr en (Scllvinctes
resJoE'nt.

mexicanus

Rock '·,TF:n (SalDil,ctcs
rare ~inter
resident.

obsoletus

eremophilus)

Possible

palustris

pleusius)

consperus)

obsoletus)

Possible

common summer
Possi~le

rare

uncommon su:nmer

Common summer and possible

Hockingbird
(~!i.21:~.'pol vglottos
leucopterus)
Uncommon summer resident.
Gray catbird
(Dc:::--:&gt;tzLlacarolinensis)
Possible
rare summer resident
3/.
Sage thrasher
-(O;Z~;~-;:;-tes montanus) Possible
common summer resident-:
American robin Ti~-;Jus migratorius
propinouus)
Common summer and winter
r e s Lde n t 3/.
----Hermit thruSh (Catharus
resident
3/.

,8uttatus

audubonii)

SwaLns on I s "t-hrush (Catharus
ustulatus
Veery
(Catharus
fucesce~alicjcola)
resid-;"~It~
-------_.
------WestcJ~ bluehird
(Sialia
~~
unco,m;'on SUr.u1lsrr'e s Lden t ,
Mountain bluebird
(Sialia
and possible
occ::sional

Possible

Cornmonsunnner

nlmae) Possible
uncommon migrant.
Possible
common migrant
and summer

baird i)

Possible

comoon migrant

currucoides)
Connnonmigrant
winter
resident.

Townsend T s solitaire
resident.

Ofyadestes

tmmsendi

tm.msendi)

Blue-zroy
gnat catcher
summer resident.

(Polioptila

caerulea

amoenus)

RUDy-crmmed klnglet
migrant.

cineraceus)

(Regulus

Bohemian vlaY.1ving(BombvcHla
winter migrant
3/.

calendula
gar~

and summer resident
Possible

amoenissima)

Golden-crm,rneci kinglet
(!:.-egul:~ satrapa
m:ig:C3rtt an d ran= SUITll!1er
resident.

pallidiceps)

and

uncommon

Possible

Possible

uncommon

Possible
Possible

common

common
irregular

Cedar ,,,aX-wing (Bo;-!~cilla
cedrorum) Uncommon resident.
Northern shrike
(Lanius excubitor
invictus)
Common w i.n t e r resident.
Loggerhead shrike'-{L-;r!iu~~;i~mubitorides)
Possible
uncommon
summer and common 'iVinter resident.
Gray vireo
(Vireo vicinior)
Possible
uncommon summer resident.
Solitary
vireo
(Vi~eo solitarius
plumbeus) Possible
Cow~on summer
resident.
Red-eyed vireo
(V'ir-oo olivaceus)
Possible
rare summer resident.
Warbling vireo
(Vireo gilvus
sl-lainsonii)
Common summer resident.
Tennessee ,"arbler
(Vermivor~ peregrina)
Possible
rare but regular
migrant.
Orange-crowned
war bl.e r (Ver.::i_vora ceL.lta orestera)
Possible
uncommon
migrant and surmner resident.
Nashville
warhler
(Vermivora ruficanilla
ridgHayi)
Possible
rare migrant.
Virginia I s war bLer (Vermiv01:-a virg-i.niae)
qossible
common summer resident.
Yellow warbler
(Dcndroica
oct~chia
~cstiva)
Common summer resident.
Yellow-rurnped wa-;:-h-~-:-~--('Dend;;i-(:~-ZorO!lClt~--l'1emorabi1
is) Common summer
resident
J!; (D. c. C~lta)
Possible
coaT.on migrant
3/.
Black--tlnoatcd
g~aY-".'arbler
(])-cndroica nigrescens)
Possible
common summer
resident.
To'-.'11send's warb l.or (])(~ndrojca tmmsendi)
Possible
uncommon fall
mi.grant.
r-klcGi11ivray's
\·;;:n-blcr (Opo;~rnis -tol~ic~ r.-:onticola)
Possible
Common
&lt;&gt;umnwrresilient.

�-117C0I1'JnonycLl oc t h ro a t (G(?~_U0-5..rJ.-s ~ri(Jl&lt;1s ?cd(!~ntalis;
G. t. campLcol.a)
Possible
uncommon sur.cnc r r cs i.dcn t 3/.
Ye Ll,o•.
!-br(:~Jstc;d chat (Ictc!"ia
v i r en s ?uricolJ.is)
Possible
common summer
resident.
Hilson's
T.Je, r bLcr (I-iilsoni a E::.:silla .Eileolata)
Possible
common migrant
and
American r eds ta r t; (Set~E ..!t0.S.~ !uticilla
t r Lcol cr a) Possible
rare migrant.
House spar r o-» (P&lt;1:05er (-i();":(~3ci cus dcrnes t i.cus )
Common resident.
Boholink (D0Lic1~-~-;-01:-::,;~Tvort1s) Po'sslble rare summer migrant.
H,~stern !:1e·~~-c-:-c-;~.'l":;'''C(''Stu~I!..~ll-;
ncO'.Jecta rieg Lcc t a) Common summer and
possible
uncommon wi nt e r r e s i.den t .
YclJ.OI:-head(;d blackbird
(Xanthoceyhalus
xant ho cephaLus ) common summer
resident.
Red-winged b.lackb i.r d (~gcla!~~ phoen:i ceus fortis)
Common resident.
Nor the rn oriole
(Icterus.
Balbul3. bu.l Lock i L) Possible
common summer
resident
1/.
Rusty b l.ackb f.r d (Eu.Eh~~1..s..
carolinus
carolinus)
Possible
rare winter
migrant.
Brewer' s b Lackb Lr d (~lJa~
cyanoc eph aLus ) Common resident.
B'rcwn=headed coxrb Lr d (Nolothrus
ater a r t emi.s Lae) Common summer
resident.
I'lestern tanager
(Pira!lga ludoviciana)
Possible
common migrant
and summer
resident.
Scarlet
tanager
(P'i.r ang a olivacea)
Possible
rare migrant.
Black-headed' grosb2alZ(-Pheucticus
melanocephalus
melanocephalus)
Possible
common surnmer resident.
Blue gro sb e alc (Guiraca cacrulea
interfusa)
Possible
uncommon sunnner
resident.
Lazu.l L. bunr Lng (Pa s s e r I.na amoen a) Commonsummer resident.
Evening grosbeak
(Hesperiphona
vesperr:inabrooksi)
Possible
Lr r egu l ar
,.;inter resident.
Cassin's
finch (Carpodaeus
cassinii)
Possible
common resident.
House finch (0~rpodacu~ i-:C-xicanus frontal::'s)
Common summer and possible
uncommon ,-linter resident.
Pine grosbeak
(Pinicola
enucleator
Plontana)
Possible
unconnnon resident.
Gray-crowned rosy finch ~Leucosticte
tenhrocotis
tephrocotis;
L. t.
littoral:is)
Possible
COITJllOn
winter
resident-.Black-rosy
finch (Leucosticte
atrata)
Possible
COTI1Il!on
winter
migrant.
Br own+cappe d rosy ~finch-~o;-~
australis)
Possible
common winter
migrant.
Commonredpoll
(Acanthis
flammea flammea) Possible
rare winter m.i.gr.mt ,
l'jne siskin
(.§.I?
..::i:.nus.E,inus pinus)
Commonresident.
Americ&amp;n goldfinch
(Spinus tristus
tristus;
~ . .!.. pallidus)
Possible
COIl'J'1l0n
s ummer and unccramorr w.inter res iden t .
Lesser i,;oJ.dfinch (~..E.~~':&gt;_
psaltria
psaltria)
Possible
uncommon summer and
rare winter
resident.
Red crossbill
(Loxia curvirostra)
Possible
rare winter
resident.
h'hite·-\v:i.nged cr'ossbill-(LClxi~
Lei.co o t e r a leucoptera)
Possible
rare
w i.n t er migrant.
Grccn=LafLe d t owhee (Chl.orur a ell] or ur a) Possible
common summer resident.
Rufous-sided
t owhce (P~ilo
ervti';-;:m~halmus
morit anus ) Cammon summer
and rare winter resident.
La rk bunting
(C:ll.J!:~c!'·?l'i&lt;:..~~
!::cL:Jl1o~o_r~) Possib Le uncommon summer resident.
Savannah sp ar rcw (J_~..~~-,-~~~~~~~Q..ll~
.,:..'.,.:~;\;j.(Jlel:sis ncva~~nsis;
P. E... an thLnus )
Pos s i.bI e COli1Ji1Ol, mi~'.raut a!Hj SU!jL1Cr resident.
G[,1~sllC!pl'er Sp.:llTC'.,1 ({\r.'J:~o.'lE~
s;!'yal1l~l~~ Eerpallidus)
Possible
uncommon
sumner' r es i den t .

1.5.

�-118Nongame birds

tiUiJl.Iller

(continued)

re s Ld cn t .

Lar k s pa r row (.f;lCrlde3tc:..~.. t:!arr:-;:-:..'.l_c~~strig&lt;1tu~)
Possible
common mig r ao t
and s\!r":~lc!rr c s i den t ,
.
Black-throio'led
spar row (Amnh f.soLz a bilin2ata
desertico1.~)
Possible
common
sumne r res iden t .
Sage SPdlTOH (AI,:.!:.hJ.:?pizD. belli
nevadensis)
Possible
common summer resident.
Dark-eyed junco (~!E~
.0yemal:is a Lkcn L) possible
rare wf.nt e r resident
]./;
(J. h. hverealis;
J. h. cisrr:on~anus) Rare winter
resident
3/;
(I. h. '';;-·-:;':u:t;:,~-) C-OJ[,;on'wint;r-';;e;;-ident 3/; (J. h. o , var-:mearnsi)
C'rnillr;~l:'lintr;r:-;eside:nt
3/.
- - Gray-headed junco (~nco ~~niceos_ canic~s)
Commonsummer and winter
resident.
Tree spax~ow (Soizella
arborea
ochracea)
Possible
unco~non winter migrant.
Cby-colored
sp;rr;;--("s);iwJ:I; pa Ll.Ld a ) Possible common migrant.
Possible
common summer
Chipping sp ar row (§22.ze!.la pass7;rin:-boreophila)
resident.
Brewer t S si)arrow
resi0ent.

(Spizella

br ewer I prev.Teri)

Possible

.connnon summer

Harris'
spar row (Zonotr_ichia
Querula) Possible
uncommon mdgr an t and rare
winter resident.
l-,rhite-cro!med spar row (Zonotrichia
leucroohrys)
Commonresident.
White-throated
s pa r row (~onotrichia
albicollis)
Possible.
rare migrant.
Fox spar rev (Pns s e r e Lla iliaca
schistacea)
Possible
rare summer resident.
Lincoln I~; spai"roH C-Tc~lospizalincolnii
alticola)
Possible
common migrant
and su.amer resident.
Song sp ar row (Helospi~
melodia)
Corrnnonsummer and uncorrnnon winter
resident.
Lapland longspur
(Calcarius
lapponicus
alascensis)
Possible
rare winter
migrarl.l: •

Raptores

2:../

Turkey vulture
(Cathartes
winter resident.
Coshawk (ACCipiter
resident.

~

"?,entilis

Sharp-shinned
haHk (Accipiter
w:int e r resident.

meridionalis)
atricapillus)
striatus

CommonSummer and rare
Uncommon summer and common ~dnter

velox)

Uncommon summer and connnon

'Cooper's
hawk (Accipi ter coope r Lj ) Commonsummer and winter
Red-tailed
hawk (13~l-teoi amai c ens t s calurus)
Commonresident.
SwaLnson I s hawk (Buteo ~Hainsoni)
Uncommonsummer resident.
Rough-legged hawk (Bute-;;--lagopus ~. j ohannis)
CommonHinter
migrant.

resident.

resident

and

Ferruginous
hawk (But~~ regalis)
Uncommonsummer and common winter
resident.
Golden e ag Le (.:2.quila. _ci1rys:.letos canadensi~)
Commonresident
Balel e ag Le (Ibliaeetus_
.!-E-ucocephalus a l.a s can us ) Commonwinter
resident

i/.

i/.

!!..I
Goldell and bald eagle specifically
as ciled
in footnote
l/ but herein

excluded from statutes
listed
to avoid omission.

16

defining

"Raptore"

�-119Raptores

]j

Harsh

(continued)
hawk (Circ_~_ li)'aneus

hudsonius)

Common sununer and uncommon winter

r c s Lden t.

Osprey

(Pand ion .h"li:]ctus

carolinensis)

Uncommon migrant,

rare

summer

rcs idcn t .
Pr a Lr Lc falcon
(Fa Leo !!1C:.:iC3D.US)
Uncommon resident.
Peregrine
falcon--(FCl1c-;p-;;-~nus
an a t urn) Rare migrant,
endangered.
Herlin
(Fa Leo col\}~'ibarius)
Uncommon wintc"r migrant.
Arner Lcan kestr~T."·(Faico
;-na:-verius
sparverius)
Common summer and uncommon
\·,i.nter resident.
Screech owl (Or us as io ) Common resident.
Flanu:nulated o\ll-"(5:~tus .flar::r.me_~0~ f Lammeo Lus ) Rare summer resident.
Great horned owl, (Eub o v Lr c Lnianus ) Common resident.
Pygmy owl, (G1:..~1~"L~~~: gno:na. _&lt;:"01~SomicUin) Rare resident.
Bu r r ovr.in g owl. (~Eeot:..~~
cunicu1ad
a hypug ae a ) Conunon summer and possible
rare winter
resident.
Long-eared
0\011
Usio otus w i.Ls on.Lanus ) Unconunon resident.
Short-eared
oHI -(ji.sio..- _fla~
f Lainmeus) Conunon summer resident
and
uncommon w int.c r migrant.
Saw-whet; oHl (Aegolius_ acadicus
acadicus)
Common resident.

W. T. McKean and P. H. Neil
June

,,.,
_L:

1974

��-121DISTRIBUTION

AND AHmmANCE

WILDLIfE

OF HIe

GA.11E HAHtlALS

Nl\NAGENENT UNIT 21

BLACK

BFAR

Black bears generally occupy the areas between 6,000
tion in Unit 21, although on occasion,
entire unit.

and 9,000

feet eleva-

they may wander over almost the

Their yearlong habitat is largely within the timbered and

brush covered lands, whe re fair combinat ions of food ar a available.

such

as fruit be ar Lng shrubs, small mamma l.s, and carrion from livestock

and big

game animals.

At least three-fourths

of the unit is considered

bear habitat,

but human activity has restricted bears to the higher, rougher,

and most

isolated s egmerrt s (see accompanying maps).

trails and

jeep-type

Minnich

There are numerous

roads within, or upon almost all major drainages

(1973)

calculated a bears-per-square

and ridges.

mile figure for a large area

extending from Rifle to the Utah line, north of the Colorado River, and ineluding all higher areas of the Roan Plateau.

Applying

available bear habitat in Unit 21 as.mapped, produces

his ratio to the

an estimated

13 bears

w-ith a density of 0.05 bears per square mile.

Estimated annual bl~ck bear harvest for the 19 year period 1955-1973
'vas 4.4 bears, based upon hunter report card surveys.

inclusive

Annual harvest ranged

from 0-16; there were six years out of nineteen when no bears were reported
taken.

Bears have occC).sicnallybeen removed because of damage complaints.

w.e.c.

Vidakovich

reported none removed during 1972

by state or federal

trappers and only three estimated removed by landowners;
killed by ranchers and unreported.

13

each year bears are

The same report indicated a popUlation

�-124MULE DEER

Almost every square mile of Unit 21 is potential mule deer summer range,
but greater numbers generally occur in areas above 7,000 feet elevation
where they can avoid insects, high temperatures,
obtain better forage.

(1974 pers. corum.) delineated deer

Vidakovich

summer range as occurring

lack of water, and can

approximately

across the southern one-third

the unit and along a narrow band below the Cathedral
side of the unit.

Bluffs on the east

lIe also reported that some resident deer occur along a

n arrow zone adjacent and parallel to the White River
maps).

of

"Overall Distribution"

(see accompanying

maps provided by Minnich

(1973 and 1974)

show the entire ~~it as summer range.

"loC.O. Vidakovich

(1974 pers. comm.) indicated deer winter range as being

the central wes t ern and northern portions of the unit, including
]~iv2r bO~':cD!~lnr1.3.

He. end Minnich

(1973. 197/;),

the White

concur on this dcsc.r Ip t r.on

(see accompanying maps).

Vidakovich

(1974 pers. co~~.) estimated a summer (post-fa1vuing) population

of 6,000-7,000 deer.

Data provided by Minnich

(1973, 1974) indicated an

"Overall Populationlt

(post-fawning) of 13,112 with a density of 15.1 per

square mile.

Winter population estimated by Vidakovich
animals.

By comparison, Hinnich

(1974 pers. carom.) were 3,000-4,000

(1973, 1974) indicated 9,652 animals with a

density Qf 12,7 deer per square mile.

Heavy w1nt~r lQsse , !loth 1n lnZ-7'J

and in 1973-74, possiLly account for these differences

in estimates.

MeaD annual deer harvest was 2,080 for years 1963-1973, including archery.
Annual harve s t ranged from 1,282 to 3,439 (see Jbrvest Data) present in this
report.

2.1

�-125Literature

Armstrong, D. H.

1.972. Distribution

Kansas Printing Service, Lawrence.

Cited

of mammals in Colorado.

Univ. of

.

4l5pp.
/.

Dixon, K. R.

1967.

141.-16/..

Evaluation of effects of mountain lion predation,

In G3mC! Res. Rep ;, July - Part II.

and Parks.

pp.

Colo. Div. Game, Fish

pp.73-310.

Le chIeLtner , R. K.

1969.

iVild mammals in Colorado.

Pruett Pub. Co.

23Lipp.

Boulder.

Hinnich, D. H.

1973 and 1974.

Statewi de mammal and bird distribution

maps

and data compiled from H.C.O., Area Supervisor, and Regional Director
input.

Located at Denver Office, Colo. Div. of Wildlife.

W. "T. McKean

May 1974

22·

Unpublished.

and P. H.

Neil

��-127DISTRIRUTTO~;\:m A13mIDANCE
OF "WILD HORSE"
WILDLIFE

(1974 pe r s . comm.) reported

t-l.C.O. Vidakovich
horses

scattered

TeX3.S

Hountain

on East
lower

Hithin
area,

Douglas

Spring

Mr. Bill
counts

Creek

Lawhorn,

Little

Fourmile

observed

Bridge

the

"wild

map).

move to and from the

Piceance

firm

this

present.

Feral

or "oLLd horses"

laws,

but

In light
\~est,

movement at

decisions

should

Gulch

herd

the

of controversies

Creek,

In addition
in the

West and

and be twe en upper

main Doug Las Creek

be noted

and Spring

of

February

flight

and 55 for

the

using

East

but

lacks

the

Douglas

sufficient

"wild

August

Creek may

data

to

con-

do not

have legal

status

under

Colorado

Federal

Statutes

by the

Bureau

of Land

of horses

concern

about

the

possible

effects

might

have on Unit

21 range

resources.
in the

t ha t

"wild

horses"

have caused

elsewhere

BLH and Division

very

probably

are

future

horse"

Gillam

counts

horses

that

to

Total

has expressed

numbers

1974.

aerial

on both -flights.

drainage,

presently

conducted

were

The Division

in "wf.Ld

in

horses

under

the

the

the

the

that

(Meeker)

were observed

between

administered

increasing

both

It

21 we r e 57 for

Hr , Lavho rn believes

the

in

and a small

and August,

horses"

areas

flight.

that

B.L.H.

ranging

Draw,

for

}fanagement.

Bluffs,

on main Douglas

horses"

are

Cathedral

Biologist,

general

S2IllE:

Unit

of wild

be tween TommyI s Draw and Rocky Point
the

areas,

accompanying

LT)

herds

area.

Draw area

Dz'aw and State

several

One group- has been observed

21 and 22 in February

above mentioned

(see

21.

another

hTildlife

the

Creek

Unit

Creek under

of Units

I'lA:--lAGE~!ENTUNIT 21

management here

and at

in for
other

Colorado.

23.

P. H. Neil
August 1974

locations

difficult
in western

�\

�-129-

I'I~}:TJ:_:nl1TIS~~\iJ?.JJm:lTlA~E_.9~_S~1ALL
WILDI.lTE ).j;\.NAGE.~·:E:;rUNIT

GM1E }W'2!ALS
21

RABBIT,.

COTTONTAIL

Nearly every squar e mile of Unit 21 is considered capable of auppo r t f.ng
a.tleast some cottontail rabbits yearlong, however, it is believed
they are more abuodan

t

at the Lowe r elevations.

that

Both the desert cotton-

tail and the Nuttall's cottontail should occur; the desert cottontail being
common at the Imler elevations and the Nuttall's cottontail being less
abundant on the plateau areas (Lechleitner 1969, Armstrong

1972).

See

accompanying maps.

No specific studies have been conducted regarding cottontail rabbit density
anywhcr

e

in we s t e rn

Colorado,

howeve

r , Shepherd

(1965a)

concluded

~ after

a

}Lt.era t.ure survey, that .E..ossible
high-cycle densities of 150-200 cottontails
per square mile and low cycle densities of 15-20 cottontails per square mile
were reasonable for much of Colorado.

These estimates may be' applicable

the Lowe r elevations of Unit 21 in par t Lcul.ar. According

to tv.C.O.

to

Vidakovich

(1974 pcrs. comm.) cottontail densities were extremely low in 1973-74.

For abundance, as reflected in harvest data, see tables on Hunter Harvest
for Small Game Hanagement Unit 8, following.

SNOWSHOE
Snowshoe

hares are

known

to occupy

.

HARE

the Douglas-fir, Douglas-fir-aspen

mix

areas on the Roan Plate&lt;lu ~long the southeastern and southern border of Unit
21

(~;ce accorapany i.ug map s).

snowshoe

hare range

occur

She phe rd (1965b) s howe d occupied

and potential

rJng at the above mentioned a rcas , but also included

some ca st ern p or t Lons of the un:Lt.

�-130Minnich

(1973) indicated

mile of occupied
reported

an est~nated

range in Unit 21.

period 1968-1972,

of abundance,'shows

vest of 493. snowshoe s by 109 hunters

and 32 across

that for the flve year

Unit 8, an average

uccurred.

annual har-

This would probably

re-

Units 21, 22, 30, 31,

in i-lild1ifeHanagement

the Roan Plateau

(1974 pers. comm ,)

along the Roan Plateau.

in Small Game Nanagement

the total harvest

of 2.7 hares per square

H.C.O. Vidakovich

they are f a Lr Ly abundant

that

Harvest data, as an indicator

present

density

from the Utah line to Highway

13 north of

Rifle.

.RED (PINE) SQUIRREL
Armstrong

(1972), Lcchleitner

squirrel

should be fairly common

Douglas-fir,

stable populations

D. H.

1911.

Cited

Distribution

of mammals

Service, Lawrence.

415pp.

1972.

A biological

U. S. Dept. Agr.,

Office.

256pp.

Boulder.

prinCipally

R. R.

1969.

Hild

mammals

254pp.

25

See

PresUmAbly

in Colorado.

survey of Colorado.
Bur , of BioI.

that this

small

types.

Literature

No. 33,

L~ch1eitncr,

forests,

or hunter harvest are '1vailable.

exist in the Doug~as-fir

Kansas Printing

Cary, H.

in the coniferous

on the higher mesas and plateau areas in Unit 21.

No data on abundance

Armstrong,

(1969), and Cary (1911) suggested

Univ. of

In North American

Fauna

Sur •• Hash., D.C., Govt. Printing

of Colorado.

Pruett Pub. Co.,

�-131-

Hinnich, D. H.
data

1973.

Statewide mammal and bird distribution

comp Ll.ed f r ora H.C.O.,

Area Supervisor,

and Regional Director

input, located at Denver office, Colo. Iliv. ofoHildlife.

Shepherd,

H.

R.

1965a.

plans, 1965-1975,
Parks

maps and

Unpublished.

Colorado long range game speci.es management
for cottontail

rabbit.

Colo. Game, Fish and

Dept.

rep.

1965b.

Colorado long range game species management

Ll.pp ,

1975, for snowshoe hare.

(Himeo).

plans, 1965-

Colo. Game, Fish and Parks Dept. rep. llpp.

CHime 0 ).

P. H. Neil

May 1974

2}.&gt;

��-133-

DISTRUUTHn
f,~m ABmmAl';CE
--------_
_.- _----...

...

OF- SHALL CANE BIRDS
-

BLUE GROUSE
The distribution maps provi.ded by Rogers

(1965) and Minnich

(1973) in-

dicated that the forested and brush covered portions of the Roan Plateau
in the southern .and eastern part of Unit 21 are occupied blue grouse
habitat.

"1.C.O.

Vidakovich

(1974 pers. comm.) reported that the head-

vat er-s of Spring Creek appear to be the northern

boundary

of their

in Unit 21.

distribution

Data provided by Minnich

(1973) estimated blue grouse density at 4.9 birds

per square mile of blue grouse habitat in the Unit which results in an
estL~ated population of 1,255 birds for Unit 21.
pe rs . counn , )

[c.:lti:.hat,
01.::he

average,

Lou Vidakovich

t he re we re

(1974

f ev er blue grou.:::e
than

this, but their numbers fluctuate greatly from year to year.

Ha1~est of blue grouse Ln Unit 2lc~

be estimated

from the Small Game

Harvest and Hunting Pressure Tables for Unit 8 included in this report.

SAGE GROUSE
Rogers (1964) indicated several areas within Unit 21 where small populations
of sage grouse are present.
headHaters

of

The top of the Book Cliff Plateau near the

Douglas Creek and the Cathedral

Yellow Cr eek s are

Bluffs between Douglas

two specific areas whe re sage grouse have been observed.

He also reported that a small population

is present along both sides of

the ~lite River about 15 miles east of Rangely.
by l-ij_fiuich
\;,(:.0"

(19";3)

Vld:Jbwich

s rar t.c red 0;)

and

concur

\6(11

those of Rogers

Distribution

maps provided

(see accompanying

maps).

(1974 p ers , comm.) reported that sage grouse are well

t h« 1:0&lt;111 Plateau.

27

�-134Rogers

(1964) suggested

that fall densities

of sage grol1se range from

1-10 birds per square mile in occupied range in Unit 21.
provided

by Ninnich

birds per square
estim~tcd

(1973) suggested an approximate

mile of occupied

population

r ang e within

..of data

Analysis

of 6.4

mean density

the unit, resulting

of 400-500 sage grouse in Unit 21.

in an

Vidakovich

(pers.

corum.) est:i.r.lwted
200-300 birds as an average.

Harvest of s~~e grouse in Unit 21 can be estimated
vest T~lblE!for Un i.t; 8 included

from the Small Game Har-

in this report.

CHUKAR
During the period 1960-1965, a total of 491 chukars were released
Unit 21 (see Game Species
co,um.) reported

Introductions).

W.C.O. Vidakovich

that he has not observed

any chukars within

last three years and believes
or perisheci.
stantially

within

(1974 pers.
the unit in the

that they have either moved out of the area

He added that the cold, damp spr ing of 1966 may have sub-

reduced their numbers.

The accompanying

distribution

map and

1/2" = 1 mile mylar overlay mainly depict where chukars we re known to occur
:in the past.

Chukar censuses are no longer taken in Unit 21 due to the bird's

RING-NECKED

10\-1

population.

PHEASANT

(1973), ring-necked

According

to Hinnich

bordering

the I·lhiteRiver from Boise Creek on the east to a point approximately

four miles west of Rangely,
agricultuT"a13reas

(see accompanying

di.stribution, but indicated
t e nnf nat cd

bein~ mainly

pheasants

confined

maps).

SHope

occupy available

to the river bottom and

(1965) showed a similar

that the. western boundary

of occupied

a few miles west of R..lngely,near Big Trujillo

28

habitat

range

'\']ash.He classified

�-135this range

as poor quality habitat ~dth a spring breeding index of 0-15,

based on spr l.ngcrow counts and winter sex-ratio counts
veys).

t-l.C.O.

Vidakovich

(from 1957 sur-

(1974 pers , comm.) concurred with Minnich's

distribution maps, adding that there have been recent transplants

by local

4-U Clubs along the ~·fuiteRiver.

Analysis of data provided by Hb1.nich (1973) suggested an estimated mean
density of 3.!f birds per square mile of occupied range along the river
vlithin the Unit boundaries, wh Lch yielded an estimated population

of 200+

ring-necked pheasants for Unit 21.

Harvest of ring-necked pheasants can be estimated from the Small Game
Harvest Table for Unit 8 included in this report.

BAND-TAILED PIGEON
Br~,t.'n (1973) ':LndiC''1te.d
that band-tailed pigec'm distribution
mainly confined to the eastern portion of the unit.

should be

He also suggested

that generally, they are extremely uncommon in the sagebrush; pinon-juniper
areas of nor t hve s t ern Colorado.

Be Ing a migratory

species, they arrive in

Colorado in mid to late April and according to Braun (1973) the timing of
fall migration is apparently related to the timing of cold fronts in October.
The availability and crop success of Gambel's oak and various ripening berries
within the unit influence the distribution

of these birds; thus, because of

their migratory nature and the distribution

of available foods, an occasional

observation of band-tailed p i geons anywhe re in the unit could be expected.
The cultivated croplands and livestock feeding areas along the Douglas Creek
drainage and the 1-.'hite
River provide a source of food for band-tailed
especially during the breeding season (Braun 1973).

29

pigeons,

�-136Lf t t Le data are available on the abundance of band-tailed
Uait21,

pigeons Ln

hm,ever, Braun (1974 pers. comm.) suggested that they are un-

common in this area, largely because of the lack of ponderosa
association w LthGambel 's oak.

It is reasonabLe to expect that their

densities fluctuate in proportion "lith availability
(berries,

aCOTI1S,

pine in

of food resources

new conifer buds, and insect larvae), and during

migration periods.

Lou Vidakovich reported he has not seen any bandtails

since 1963.

MOURNING DOVE
Grieb (1965) indicated that the breeding range of mourning
throughout Unit 21.

Migratory

doves extends

in nature, mourning doves are common during

spring, summer, and fall in Colorado.

According to Bailey and Niedrach

(1965) a few may remain over the Hinter months in sheltered areas at lower
elevations

III

Cc Lcr ado

v!hieh, during mild v:Lnters, may apply to the J Olver

elevations of Unit 21.

Mourning doves have been observed throughout

tmit but are more abundant at the lower elevations.

See accompanying

Actual abundance of mourning doves in Unit 21 is difficult

the
maps.

to determine,

however, according to Davis (1969), Braun (1974 pers. comm.) and W.C.O.
Vidakovich

(1974 pers. comm.) they are very common during the spring and

summer months.

i1-f.nnich

.9 doves per square mile

(1973) estimated popu Lat Lon densities ranging from
at the higher elevations in the southern three-

fourths of the unit, to 15.4 doves per square mile along the Lowe r elevations
of Douglas Creck, and upwards to 42.3 doves per square mile in areas along
the \'mite River on the northern boundary of the.unit.

Resulting population

estimates ranged from 500 to 2,000 and upwa rds to 5',000 birds respectively
for the areas mentioned above . Due to their migratory nature, population
densities

~Jj

t.h
i.nthe unit var y cons] derab ly scnsona l l.y and from year to year.

30

�-137Harvest of doves for 2Jinll Came Hanagerneut
21.

Unit 8, which includes Unit

is Ll Iuc t r at e d in the Small Came Harvest Table in another

section of

this report.

WATERFOHL
Ereec1i~.cI

sum:;~erpopu La t Lon s ;

wat erf owl, brccd Lng habitat.

Generally,

Available

Unit 21 offers little good

range is mainly confined

to isolated

river bo t tom and irrj gated meadow areas along the ~fuite River and Douglas
Cree.k and some of their tributaries.

W.C.O. Vidakovich

(1974 pers. corum.)

reported t.hat; various species of wat er f owl, may be observed
during migration

and breeding periods to include such species as mallard,

g reen-w ing ed teal, cinnamon

and American coots.

teal, pintail, gadwal.L,

golden-eye,

He added that sandhill cranes, whistling

snowy egrets are also common migrants

in this area.

that the b rced irig range of the grcen-wtnged

Grieb

Szymczak

swan s , and

(1965) indicated

(1974 pers. comm.) suggested

species

down from the Yampa Rfvar area arid nest near sheltered

tv. C.O. Vidakovich

Jobman

teal to drift

stock ponds and beaver

(1974 pers. coram,) reported observing

33 goslings in 1974 in Unit 21 which indicates
pairs of geese is increasing

in the

that it would not be'unreason-

able to expect these two species and others like the cinnamon

ponds in Unit 21.

mergansers

teal extends over all of Unit 21

and stated that the mallard remains the most common nesting
state.

in these areas

that the number of breeding

in this area.

(1967). in a duck production study on irrigated meadows of Picea~ce

and Dry Fork Creeks, estimated a peak summer population
8 mile segment.

Assuming

that Douglas Creek-White

what comparable, a projected population
approxim2tcly

of 138 ducks on an

River wetlands

are some-

estimate for this area would be

500-1,000 ducks, depending mainly on the amount of water that

31

�-138is ava Ll.ab Le ,

Wint:.crir:gpopu lat Lons :
within

(197!!pers. comm ,) estimated

Vidakovich

The only available

the unit are the open

open stock or beaver

the mallard

but a fel-lother species

months.

Grand .Junct Lon OJ:"gel further

south.

No aerial

Bailey,

Iiist.

2 Vol.

Brailll,C. E.

1973.

Distribution

Colorado.

Davis, W. A.

wintering

and green-winged

teal

The geese from this area winter

near

of the ~Thite River.

Cited

A. H., and R. J. NiedJ:"ach. 1965.

of Nat.

some

to H.C.O.

is the principal

trend counts are condt::cted on this segment

Literature

for waterfowl

and perhaps

According

such as golden-eye

also remain over the winter

areas

of the t·lhiteRiver

ponds on its tributaries.

(1974 pers. comm.),

Vidakovich
specIes

wa ters

wintering

300-500 birds.

Birds of Colorado.

Denver

Mus.

895 pp.

Proc. Western

and habits

Assoc.

1969.

Birds of Hestern

1965.

Colorado

of band-tailed

Game and Fish Coroms.

Colorado.

pigeo~s

in

53:336-344.

Colo. Field Ornithologists.

61 pp.

Grieb,

J. R.

1965-1975, for migratory

long-range
birds.

game species

management

plans,

Colo. Dept. Game, Fish and Parks.

36 pp.

.Jobman , H.

C.

1967.

Waterfm"l

for state-m-illed lands
Color~do.

n ea r

production
Little

and management

Hills Experiment

CSU ten: paper for FW-495bx-Spcc.

38 pp. ('£YPcl-lrittcn).

32

Studies

recommendations
Station,

Meeker,

in Wildlife.

�-139111nnich, D. W.

1973.

da t a comp Lle d

Statewide mamma I and bird distribution

from HCO,

Area Supervisor,

and Regional Director input.

Located at Denver Of f Lce , Calc. Div. of Hildlife.

(Unpublished).

Rogers, G. E.

1964. Sage grouse investigations

in Colorado.

No. 16.

Colo. Dept. Game, Fish and Parks.

132 pp.

1965.

Dept. Game, Fish and Parks.

1965.

grouse, ptarmigan.

Colo.

16 pp ,

Colorado long-range game species management

1965-1975, for ring-necked pheasant.
Parks.

Tech. Pub.

Colorado long-range game species management plans, 1965-1975,

for blue grouse, sage grouse, sharp-tailed

Swope, H. M.

maps and

plans,

Colo. Dept. Game, Fish and

54 pp.

P. H. Neil
Ma~_ 1974

33

and W. T. McKean

��-141-

(From Colorado

Big

GaEl8

Harvest

HARVEST
----DOES
FAHNS
BUC~S

1956-1973)

PUNT

TOTAL

AND SE.:\SO:J

1956

2763

2223

1316

256

'1795

2ES

10/15-10/31

1957

7.951

2060

1333

396

3839

2DX
2DH

10/1-10/14
10/15-11/17

1958

5134

22.31

549

4119

1DM

10/15-11/2

1959

5675

2713

1713

594

5020

1D~·110/17-11/3
IDH 11/4-12/2, In part

1960

2858

2211.

1401

371

3986

IDH

10/17-11/6

1961

3506

2644

1657

389

4690

IDM

10/21-11/8

1%2

3801

3237

2293

380

5910

2ES

10/20-11/4

1963

3924

2336

2252

560

5148

1DM

10/19-11/7

2279

842

500

90

1432

ES

10/17-11/5

1965

3330

1196

1876

3~9

3401

1DMA 10/16-11/5

1966

2261

645

569

95

1309

ES

10/15-10/31

1967

2084

775

494

'115

1384

ES

10/21-11/9

1968

3343

2084

1081

274

3439

ES
2ES

10/19-10/27
10/28-11/7

1969

2978

1119

686

155

1960

ES

.L0/18-11/6

1970

3217

1481

551

38

2070

ES

10/17-1.1/6

1971

2586

1528

1528

IDB

10/30-:1/11

1972

3081

1589

692

22

2303

ES

10/14-10/23

1973

2439

1025

194

22

1241

IDB
ES

10/13-10/1B
10/19-10/23

Jj !;ased upon combLned t o t a.L resident
in nUffiber of licenses,

(Foo t not.cs continued

not

on

including

and non-resident
archery hunters.

following page).

34

license

sales

and expressed

�-142-

DE!-:RJ~.gl{£.S-r:L~r:/:~.O:rS
_.:''''-''ro J.!..l~~:!.TJNr
__
1'REs~::!;m::
t-lILDLIFE

l-!:\:;/,C;;:;;:;;T

U;;IT

21 Cuntinued

Note:

Hunt syr:;bolsexplained as follows:

ES

Either sex, one deer (one license).
Either sex, two deer (one license).
One deer, buck only (one license).
One deer multiple, eith8r sex (2 licenses and 2 deer per individual).
One deer multiple, Olle must be antlerless (2 Licenses and 2 deer per
individual).

2ES
lDE
11.11-1

IDHA
2DH

Hote:

Two deer multiple,

either sex (2 licenses and 4 deer per individual).

The term IDH evolved into 2n, He or two deer, hunters
and 2 deer per individual) and is synonymous.

P. H. Neil

Harch 1974

35

choice (2 licenses

�-143Ai~CHER·[ D!':ETt H/.r~VEST l\~,nJ SEJ\SO;,7S
---'--"'- ---.-----.~-_ .. --.------"-"._--_.--,- .._-- - ".'---

lHLUl.IFE ;·t\:&lt;AGE;'lE~\:T V';IT 21
------_._---------.

(From
YF'ill
~
.'

\.

1:../

Hl'CKS

CoJ.oY.:1do Big

DOES

1956-1973)

Game Harvest

FAUNS

TOTAt 2J
---.--

HUNT

lum SEASON

1956 1/

ES
2ES

10/1-10/14
10/15-10/31

1957

2DH

.

10/1-11/17

ES
IDH

9/1-9/30
10/15-11/2

ES
Lj}1
IDH

9/15-9/30
10/17-11/3
11/4-12/2,

1958

8

2

0

10

1959

4

0

0

4

•.

1960

2

6

0

8

1DN
IDN

9/10-9/30
10/17-11/6

1961

22

5

0

27

IDH
lDM
IDH

8/26-9/10
10/21-11/8
11/11-11/26

1962

30

7

0

37

ES
?ES

8/25-9/23
10/20-11/4

1963

26

10

4

40

ES
IDH

8/li-9/8
10/17-11/1

1%4

"&gt;'7

12

10

59

ES
ES

8/15-9/13
10/17-11/5

1965

13

10

6

29

ES
1DNA

8/2J-9/12
10/16-11/5

1966

54

36

11

101

ES
ES

8/20-9/18
10/15-10/31

1967

52

28

11

91,

ES
ES

8/19/9/17
10/15-10/31

1968 !!.../

27

17

7

51

ES
ES
2ES

8/17-9/15
10/19-10/27
10/28-11/7

1969 -~/

42

0

0

42

1DB
ES

8/16-9/14
10/18-11/6

1970 fll

20

0

0

20

IDB
IDB
ES

8/15-8/31
9/1-9/20
10/17-11/6

36

In part '"

�-144DEER HARVEST --_.-_._--AIm SEASONS
..ARCi:FTi.Y
-.-~.-. ----_._-_._._-----------_.HILDL iT }J!,?;/J;r:';UiT UiUT 21 cou t Lnucd

(From Colorado Big Game Harv£:st 1956-1973)

YEAR 1./

BUCKS
---

DOES

FA~·ms
--

TOTl\L

J971 Jj

18

0

0

...
18

!if

41

·15

0

56

1972

JJ

HUNT AND SEASON

IDB
IDB

8/21-9/6
10/30-11/11

lDB
ES

8/19-8/31
9/1-9/20
10/14-10/23

lDB
ES
!DB
ES

8/18-8/31
9/1-9/23
10/13-10/18
10/19-10/23

E$

1973 2)

19

16

6

41
•

--_._---

1/ Free permits issued to holders of regular big game hunting

licenses by
applico.tion only. Years 1957-1960, no issue of special permits; archery
hunting allowed to holders of valid deer licenses. Years 1961-1972, separate
archery license regulations in effect.

I/

Inconclusive harvest data omitted for years 1956 and 1957.

1/ For years 1956 and 1957 no hunter pressure data available.
!!.,/ 166 total resident and non-resident archery licenses issued for Unit.

2/ 847 total hunts for 1969 archery season in Unit 21, total does not reflect
number of hunters.

~/ 106 resident and non-resident archerY hunters.
7/ 124 hunters based on number of hunts; 91 total resident and non-res i den t
archery licenses issued for Unit 21.
8/ 145 resident and non-resident licenses issued for Unit.

2/ 225 resident and non-resident
Note:
ES

2ES
lDB
1DH
1DNA
2DH

Note:

archery hunters.'

Hunt s)~bols are explained as follows:
:::

:::

Either sex, one deer (one license).
Either sex, two deer (one license).
One deer, buck only (rne license).
One deer multiple, either sex (2 licenses and 2 deer per individual).
One deer m~ltiple, one must be antlerless (2 licenses and 2 deer per
individual).
Two deer mu l.tLpLe , either sex (2 licenses and 4 deer per indivtdual).
The term 1m! evolved into 2D, He or tHO deer, hunters choice (2 Lf.ce ns r-«
and 2 deer per individual) and is synonymous.

37

P. H. Neil
Ha r ch 197/.

�-145iIU&gt;iTING T'Iu:Ssurm
!I:l~_c:·\Z0::~s2:.,__~~2~~~
q~:£~~:'~'l'}_
.---.-----.-~------

__

HlLrL;
ri: ~~\;·:/\.C L=&lt;L~:'1'
.•.._-----------_.

_.

(Trela

mIYTI!'~G
YEAR

}';:~LSSU1\E

J/

Co Lo rr.do

Big

El.i1.LS

CCy":

Grtme

~::rr 21

Harvest

1956-1973)

HAl\VI~ST

CALVES
-----

/

TOTAL

19S6

Closed

j957

Closed

1958

Closed

1959
1960

HUNT AND SEASON

"---'-

1

1

0

0

1

AD

10/17-11/3

60

3

0

0

3

AD

10/17-11/6

9

6

0

0

6

AD

10/21-11/8

1961
1962

Closed

1963

Closed

1964

Closed

]:./

1965

4

4

0

0

4

25B + 25C

1966

0

0

0

0

0

25B + 25e

0

0

0

0

0

30B + soc 1/

10/21-11/9

0

6

0

6

30B + 30e 1/

10/19-11/7

0

0

0

0

30B + 30e 1/

10/18-11/6

5

2

0

7

AD i/

10/17-11/6

In part

11

1

0

12

AD i/

10/16-10/25

In part

0

0

0

0

AD !i/

10/28-11/12

In part

0

3

0

3

AD + 10C 4/

1967

1968

50

196)

0

1970

2/;

1971
1972
1973

70
9
38
"

2./ Based upon

10/16-11/5

1./

lO/i.3-H/3

10/27-11/11
In part

cODbjned r e :denl
and non-resident
license
sales
and expressed
i.n
number of licenses;
not including
archery hunters.
During the period
1956-1970,
~11cn elk season coinciducl with d0cr sea~on,
the hunting
pressure
statistics
were
very p rob ab l.y con s c rvn rLvo . Despi.te difficultips
verifying
that assumption,
un known [';,!bstal1tial
l:t:mbcrs of deer hunters
may also have carried
elk licenses
hoping for a bull (:.L;~ under antlered-only
regulations.
This latter
happens rC[;i.lrdLes s of 10'., elk ?0puOia(·iol1s.
Thus, because
of low succcs s an d tendency
(above
rlOn-rc;"orling
statd·:i.ci(:
fi.::;t,;res used Ln l1l'ojcctj.on~~) for non+s uc ce s s f uj, elk
license
holdcr~, not ro r e nor t; by card, e Ll; present
in Unit 21 probably
received
c&gt;:tra pr cs sur o over t.h a t Lndi c a t od by ca r d proj(~ction
y e a r s of antlered-only
(Foo r no t cs co n t Lnucd

on follmdng

P'!f,c).

33

�-146-

1/ Units
1/

21,

Units

22,

30,

31,

21,22,31,32

!if Units

22,

31,

32,

Note::

Hunt symbols

AD

=

32 combined.
./

combined.
and part

are

of Unit

explained

21 (east

.

of State

Highway 139).

permits

only

as follows:

Ant Ler ed only.

B + C

Bull

AO + C

An~lered

(antlered)
only

and cow (antlerless)
plus

specified

number of cow permits.

P. H. Neil
April 1974

(specified).

�-147J&gt;Y.CHERY ELK JL\\:V~_:'_~~~ __ ~~:~0qJ;~
__J\~.0) llTJNTI;~G
---···----···---\7J-:r
..
J
:j·l.:-L]"'
.:
!·~/\;:~:\Gl.&gt;i
;:,;;T lJ;'~IT? 1
-_._ _-------------(From

YEt,R

HUl:T11iG
]/
pT'
r_, SC'V;)E.
.:
..•• ..•
-...J

---J.\.

~

Col.o r adc

Big

B1JLLS
--

CO'NS

Came Harvest

PRESSURE

1956-1973)

HUNT .AND

CALVES
TOTAL
----.-

1956

Closed

1957

Closed

1958

Closed

1959

----

No Data

196(1

----

No Data

---- --- - -

No D",ta

1961

AD

10/l7-ll/3

AO

10/17-11/6

AD

10/21-11/8

1962

Closed

1963

Closed

1964

Closed

1965

- - -- -

1966
1967

1968

---3

25B + 25C Jj

No Date:.

----

25B

No Data

----

ES
30B + 30C :'1,/

0

0

0

----

No Data

1969

+ 25C 1/

No Data

0

SEi\SOl~

10/16-11/5
10/ lS-:L:l/ 3

. 9/2 -9/17
10/21-11/9

30B + 30e 1/

10/19-11/7

30B + 30C 1/

10/18-11/6

1970

----

No Data

AD !±I

10/17-11/6

In part

1971

----

No Data

AD !!J

10/16-12/25

In part

1972

0

0

0

0

0

AD
AD i/

8/19- 8/31
10/28-11/12 In part

1973

2

0

0

0

0

AD

8/18- 8/31
9/1 - 9/23
AD + laC ~/ 10/27-11/11
ES

-1/ Base d on combi ne d. tota 1 r cs Lcdent
in numhcr of licenses.

an d non-resident

(Footnotes continued on folloHing p.:1ge).

license

sales

and expressed

In

p.}';

�-148-

:.Y
}j

Units

21, 22, 30, ai , 32 combined.

Units

21, 22, 31, 32

comb f.ned ,

./

!.!..l Units 22, 31, 32, and part of Unit 21 (east of State Highway
Note:

Hunt symbols

139).

f'x.plainedas [ollO'l"s:

AO

Antlered

only.

B + C

Bull (antlered) and cow (antlerless) permits only (specified).

AO + C

Antlered only plus specified number of cow permits.

ES

One elk, either sex.

P. H. Neil

April 1971f.

�-149-

J}J,.A'?".:....1~J:(~'R:._!..1_t0~Yi:~~:~!:~_:.I~
__0L.2.:~0 ~,:s
\:lLDLJ I'J: }~\:;N;E:i:·,:;T

r.nr 21

(From C0lorad0 Bi~ Game Harvest 1955-1973)

.Y

HUNT !\J.~D SEASON

]j, _'l/

YE.A.R

IlARVIST

1955

6R

8/15--10/1 and the regular deer season.

1956

5R

5/15-8/15 and th~ regular deer season.

1957

o

1958

98 + 3R

1959

2R

No da t a on special season; any regular or post
deer and/or elk season.

1960

3R

4/1-9/15 and the regular deer and/or elk seasons.

1961

3R

Idl-9/15

1962

3R

4/1-9/15 and the regular deer season.

+ 3R

4/1-9/15 and the regular dec~ season ending 11/7/63.

o

4/1-9/15 and the regular deef."season ending 11/')/64.

+ 2R

4/1-9/30 and (he regu 1ell. deeL'and/or
ending ll/5/65.

1966

10S

4/1-9/30 and the regular deer and/or elk seasons
ending 11/3/66.

1967

16

Special season probably 4/1-9/30 and the regular
deer and/or elk seasons ending 11/9/67.

1968

o

4/1~9/30 and the regular deer and/or elk seasons
ending 11/7/68.

1969

o

4/1-6/30 and th~ regular deer and/or elk seasons

1963

68

19M
190j

13S

4/1-11/15 and the pre and regular deer seasons.
4/1-11/15 and the regular deer season.

and the regular deer and/or elk seasons.

eLk sedSOilS

ending 11/6/69.
1970

o

4/1-6/30 and the regular deer and/or elk seasons
ending 11/6/70.

1911

o

4/1-6/30 and the regular deer and/or elk seasons
ending 11/11/71.

+ 2R

4/1-6/30 and the regular deer and/or elk seasons

1972

2S

ending 11/12/72.

lR

1973

4/1-6/30 and the regular deer and/or elk seasons
ending 11/11/73.

-----------------------------------------------(Fco t no t eu

0:-[

f o l Low i ng page).

�-150Bj"ACj~ I.',U,R
1I/1.1'VEST 11:1)) SEASm;S
._._-_.
__ ._--_._--._--------

li-EJ2LIFE :;0.;:t~;I~~~rE:;T
m:l'l' 21 (cl)ntin~~~iL
•

1/

- R = Regular big same (Deer and/or Elk) licenses; S = Bear license for
Special SpriEg, SUDDer, or Sp r i.ng-Bunune r Seasons; Also see annual regulations
for archery hunting.

.

2/ From 1955-1959,

1 bear per special bear license and/or 1 bear per hear
coupon on either or both deer 3nd elk licenses; from 1960-1965, same as
1955-1959, except special bear license variously invalid after September 15
or 30, when bear coupon on either or both deer and elk license covered bag
and possession limits; 1966, s arne as 1960-1965, excep t one bear, hunter's
choice, per person per calendar year; from 1967 to present, bear coupon
removed from regular deer and elk licenses; thus 1 bear, hunter's choice,
per SPecial be2r or sportsman's license per person per calendar year on1z
during special bea~ and regular deer and/or elk seasons.

3/

.

- Dogs perml.tted except when bear seasons concurrent with deer and/or elk
seasons.

P. H. Neil
April 1974

43

�-151MOUNTAIN LION HARVEST AND SEASONS
WILDLIFE MANAGEHENT UNIT 21

YEARY

HARVEST

1965

2F 1/
IF 1/

1966

3M; 3F 1//
5M; 2F 1

HUNT AND SEASON

Rio Blanco Co.
Garfield Co.

10/16/65-3/31/66 West Slope; 10/23/653/31166 East Slope, 1 either sex.

Rto Blanco Co.
Garfield Co.

-

1/1/66-11/31/66 in all counties west of
the Continental Divide and in Jackson,
Conejos, Alamosa, Mineral, Saguache,
Rio Grande, Costilla, Archuleta, Hinsdale,
and San Juan counties, 3 either sex.
10/22/66-2/28/67 in all counties east
of the Continental Divide except those
specifically listed above; 3 either sex.

1967

No Data

9/1/67-3/31/68 statewide; 9/1/67-5/31/68
west of State Hwy. #13 and north of
U.S. Hwy. #6; 1 either sex.

1968

3M + 3F

9/1/68-3/31/69

statewide;

1 either sex.

1969

1M + 3F

9/1/69-3/31/70

statewide;

1 either sex.

1970

1M

1971

o

3/1/71-4/30/71, 1 male only, designated
areps including Unit 21; 9/1/71-10/12/71·
and 11/12/71-12/31/71,1
either sex,
designated areas including Unit 21.

1972

5M

2/1/72-4/30/72, 1 male only designated
areas including Unit 21; regular deer
and elk seasons statewide, 1 either sex;
11/13/72-12/31/72,
1 male only,
designated areas including Unit 21.

1973

4M

10/13/73-10/23/73
and 10/27/73-11/11/73,
statewide, 1 either sex; 11/12/73-12/31/73,
six male only permits issued for Unit 21.

9/1/70-12/31/70 designated areas, and'
regular deer and elk seasons statewide;
1 either sex.

1/
- Bounty of $50 per lion paid within period May 7, 1929-March 12, 1965;
protected by statute as of July 1, 1965, when mountain lion license was ..
established.

(Footnotes continued

on following page).

44

�-152MOUNTAIN LION HARVEST AND SEASONS
WILDLIFE MANAGEMENT UNIT 21 (continued)

~/
Credited to "1965-1966" and "1966-1967" for Rio Blanco and Garfield Counties
as reported by Dixon, K. R. 1967. Evaluation of effects of mountain lion
predation.
p. 147. In Game Res. Rep., July-- Part II. Colo. Div. Game, Fish
and Parks.
pp. 73-31~
No data available in reports from Section of Game
Management for 1965-1967. Dixon's information is probably conservative in
that the damage law allows for lax reporting of kills of marauders by stockmen.
For years 1968-1972 data are from Colorado Big Game Harvest Surveys.

P. H. Neil
April 1974

45

�SMALL GAME HARVEST AND HUNTING PRESSURE
SMALL GAME MANAGEMENT UNIT 8, 1968-197311
1968

1969

NUMBER OF HUNTERS
1970
1971
1972

1973

Average-

1968

1969

1970

HARVEST
1971

1972

1973

Average

1984

1328

1319

1411

1502

1070

1435

24813

11606

9276

14148

13511

7139

13415

Snowshoe hare

101

30

73

95

244

253

133

134

78

140:0.1 838

1575

678

Blue grouse

298

154

213

124

442

140

229

1172

73

368

. 78

815

144

442

Sage grouse

190

301

258

267

537

183

289

603

767

724

597

1134

584

735

Chukar

122

453

151

223

NOS 1/ NOS

238

76

606

247

358

NOS

NOS

322

27

NOS

61

NOS

NOS

NOS

44

31

NOS

153

NOS

NOS

NOS

92.

313

621

129

250

256

71

274

586

1040

127

246

446

212

443 If

NOS

NOS

NOS

NOS

NOS

NOS

--

NOS

NOS

NOS

NOS

NOS

NOS

--

276

290

242

447

107

272

--

5828

2412

2091.' 4373

920

SPECIES
Cottontail

Gambe1's

rabbit

quail

b:-Pheasant
Band-tailed
Mourning

pigeon

dove

--21

oil

3125

1/

- See Chap. 3 - Colo. Game, Fish and Parks Division laws and Regulations Handbook; Wildlife Management Unit 21 is wholly
contained within Small Game Management Unit 8 and approximates 17.4 percent (17.4%) of Unit 8's total area (300 sq. mi.
+ 4,970 sq. mi.) as planimetered .from Colorado topographic maps 1:500,000 scale.
Data from Colorado Small Game Harvest
Surveys 1968-1973.

21

- ~!ean for years data is shown.
3/
- No data or No open seasons
4/
- Possible

(NOS).

sample size error.

W. T. McKean
June 1974

I-"
U'1

�-154DUCK AND GOOSE HARVEST AND HUNTING PRESSURE
RIO BLANCO COUNTY. 1954-1973 1/
(From Colorado Waterfowl

YEAR
1954

DUCKS
EST. NO. HUNTERS

Kill Surveys)

EST. HARVEST

No Data:

GEESE Y
EST. NO. HUNTERS

EST. HARVEST

162

No Data

No Data

1955

14

1,976

"

"

1956

No Data

854

"

"

1957

.No Data

1,757

"

"

1958

135

824

"

"

1959

81

627

"

"

1960

38

516

"

"

1961

64

432

"

"

1962

98

252

"

"

III

531

'!

"

1964

170

645

"

"

1965

73

226

"

"

16

587

31

"

105

267

25

"

106

1,180

13

19

571

No Data

No Data

1,494

25

19

724

16

17

1.430

33

63

901

None

None

1963

1966
1967
1968
1969
1970
1971
1972
1973

1/
- Waterfowl

64
161
135
99
71

kill

Jilla are available only by county.
2/
- Small populatiol\!1 of geese and hunters alike create sampling difficulties.

47

W. T. McKean
May 1974

�G&amp;~
SPECIES

DATE

MALES

Chukar

3/30/60

45

3/24/61

3/19/62
~

SPECIES INTRODUCTIONS - WILDLIFE MANAGEMENT UNIT 21

FEMALES:

TOTAL

35

80

58

.50

75

75

•

RELEASE AREA

•

SOURCE OF STOCK

REFERENCES

Wardell's Ranch, 13.3 miles
west of jet. of Colo. 64 and
Rio Blanco 2, approx. 1.5
miles from Utah line.
(TlN Rl04W S25)

Little Hills

Sandfort's Files
(Ft. Collins Research Center)

i08

Wardell's Ranch, near small
stockyard about 0.3 mile
above Wardell's buildings
at foot of the north facing
hillside (TIN RI04W S25)

Little Hills

Sandfort's Files
(Ft. Collins Research Center)

150

13.3 miles west of the main
road jct. at west end of
Rangely (TIN RI04W S27)

Little Hills

t-4
Sandfort's Files
(Ft~ Collins Research Center) Y'

Sandfort's File
(Ft. Cpllins,Research Center)

3/15/65

27

26

53

West side-of Colo. Hwy. 139.
5 miles south of Rangely
(TIN RlOIW S29-30)

Little Hills

• 3/16/65

50

50

100

One-half mile from jct. of
Texas and Missouri Creeks
near Allen Gentry cow camp.
(T3S RlO4W_S12)

Lit tle Hills

&amp;

.

.

I

\J\

. Sandfort's Files
(Ft. Collins Research Center)

Literatur~ Cited
Ev~s,

R., K. Wagner, and B. Rowe. 1961. Chukar planting ~ecord.
files at Fort Collins Research Center.

March.

Rogers, C. E., K. Wagner, and T. Wardell. 1960. Chukar planting record.
'
Sandfort's files at Fort Collins Research Center.
(Citations ('ontinued on fo11c'\o11ng
page.)

Original rel~ase records located in Sandfort's
Mar,ch. Original release records located in

�GAME SPECIES INTRODUCTIONS - WILDLIFE MANAGEMENT UNIT 21 (continued)
~ature
. , and B. Rogers. 1962. Chukar planting record. March.
-----Fort Collins Research Center.

Cited
Original release records located in Sandfort's files at

Vidakovich, L. D., and H. Eddy. 1965. Chukar planting rec0rd. March.
files at Fort Collins Research Center.
-----, and G. Peters. 1965. Chukar planting record. March.
Fort Collins Research Center.

~

Original release records located in Sandfort's

Original release records located in Sandfort's fil~s at

P. H. Neil
April 1974

I

t-'

IJl

0'1
I

�-157CENSUS AREAS AND ROUTES - WILDLIFE MANAGEHENT UNIT 21

•
•

MULE DEER

Annual spring meadow counts are conducted on East and Main Douglas
'.

Creeks, Evacuation Creek in the western portion of the unit, and along
the White River.

An annual winter trend count is conducted along the

White River only.

These counts are made from a ground vehicle and the

results can be obtained from W.C.O. Lou Vidakovich in Rangely •

•
Aerial pre and post season sex-ratio counts have not been es t abLd.shed in
Unit 21, nor have aerial trend counts.

ELK
Because of the relative low density of elk in Unit 21, specific winter
aerial trend counts have not been established for the unit.

Ground

obnerlations of elk and their locat.ions have been recorded on an annual
.basis and available information regarding the results of these observations
are kept on file with W.C.O. Vidakovich in Rangely.

BLACK BEAR
Incidental observations and reported observations are recorded and kept on
file with W.C.O. Vidakovich in Rangely.

In addition, an annual estinate,

by W.C.O. district, of bear populations was begun in 1972.

This estimate

also included estimated harvest by trappers and others, and management
objectives for the species.

CHUKAR
An annual production count was established on the Wa!de11 Ranch, up Cottonwood Creek, beginning in 1961, but because of a substantial decline in

50

�-158-

population, was discontinued in 1968 or 1969 (Vidakovich 1974 pers. '.
comm.).

Another annual census route was established

Staley Mine area and was also discontinued

in 1960 in the

in 1968 or 1969.

Only a

'.

portion of this route, along Highway 64, was within the boundaries

of

Unit 21.

Details of technique, maps of the areas counted, and the results may be
found in the Northwest Regional Office, in ~he Denver Office, and in
Sandfort's files on chukars in the Fort Collins Research Center.

WATERFOWL
Aerial counts are not conducted in this wildlife management

P. H. Neil
t1ay 1974 .

51

unit.

�HABITAT RESTORATION
DATE
11/1941

LOCATION

PROJECTS - ~rrLDLIFE MANAGEHENT

VEGETATIVE
TYPE

TREATMENT

NO. ACRES
TREATED

PURPOSE OF
TREATHENT

AGENCY

Douglas Cr.-West, Black
Sulphur Cr.-East. Scattered
in TIN R98W, TIS R97,98W,
T2S R97,98,lOOW, T3S R99W,
T4S R94,95,99W.

Sagebrush

Seeded ~.;rith Agropyron
cristatum, Agropyron smithi,
Br\~
inennis, Helilotus
offi~ina1is, Melilotus alba.

298 total
(Scattered)

Increase
forage

BLM

Texas Cr. and Missouri Cr.west fork. Scattered in T2S
RI03W; T3S R9~,100W; T4S
R101,102,103W; T5S RI02,

Sagebrush

Se2ded with Agropyron
cris~,
Agropyron smithi,
Bro~
inermis, lfelilotus
officinalis, Melilotus alba.

1387 total
(Sea ttered)

Increase
forage

BLM

11/10-'20/ Calami ty Ridge
1942
TIN R99W 57,8,9,16,17,18;
TIN R100W Sl,12,13,24; T2N
R99W S6,7,18,19; TIS RI00W
S10,11,14,22,23,24.

Sagebrush

Seeded with Agropyron
cristatum, Agropyron smithi,
Melilotus officinalis.

500

Improve
forage

BLM

9/1943,
6/1944

Texas Cr. &amp; Missouri Cr.west fork. T3S Rl01W S18;
T3S RI02H S33; T4SRl02W
S18; T5S R102W S14.

Sagebrush

Seeded with Agropyron
~~thi,
Bromus inennis,
MeIi10tus officina1is.

8/20 &amp;
28/1959

Fletcher and Hammond Draws.
T2N RI00w SI5,16,17,18,19,
20,21,22.

Pinonjuniper

Chain

11-12/
1962

Gillam Draw
TIN RIOHl SI,2,11,12.

Pinonjuniper

Chain and seed (aerial
broadcast) with Agropyron
cristatum.

11-12/
1941

lOW.

~

UNIT 21 1/

I
to-'
\J1

\0
I

.

(I-way)

.•

100

__Improve
, forage

632

' Pinon
juniper
control

BLM

900

Improve
grasses

BLM

------------------------------- .._----------------------------------------------------------------

BLM

�HABITAT RESTO&amp;\TION PROJECTS - WILDLIFE MANAGEMENT UNIT 21 1/ (Continued).

DATE

V\
I.&gt;J

LOCATION

VEGETATIVE
TYPE

TREATMENT

NO. ACRES
TREATED

PURPOSE OF
TREATMENT

Chain

306

Increase
grasses

BLM

AGENCY

11/1962,
4/1963

Gillam Draw
TlN R100H S18,19; TIN
R101W S13,24.

Pinonjuniper

10/1711/13/
1964

T2N R100W Sl,2,11,12,13
14,15,23,24.

Pinonjuniper

Chain and seed (aerial
broadcast) with Agropyron
cristatum.

1512

Pinonjuniper
eradication

BLM

11/20/65

Douglas Creek
T3S R100W Sl,7,18,19,30;
T3S R101W S10,11,12,13,
14,15,16,21,22,23,24,25,
26,27,28,35,36.

Pinon- ••
juniper 1

Chain and seed (aerial
broadcast) with Agropyron
cristatum.

2100

Pinonjuniper
control

BLM

Douglas Creek
T3S R101W S8,9,16,17,18,19,
20,21; T3S R102W S10,12,13,
14,15,23,24,25,26.

Pinonjuniper

1889

Pinonjuniper
control

BLM

12/4/65

I

Chain

1/
- If details are desired on any of these projects, make requests directly to Mr. Ron Kufe1d, Colorado Division
of Wildlife, Research Center, Fort Collins, Colorado 80521.

P. E. Neil
May 1974

I
•....
0\

0
I

�-161-

MANAGEMENT PROBLEMS CHECKLIST

A.

- lVILDLIFE

MANAGEMENI' UNIT

•

Present
1.

21

Road development - as related to oil and gas exploitation.
The East Douglas and Main Douglas Creek areas in particular,are
heavily laden with numerous secondary roads leading to oil and
gas drilling sites, wells, and pumping stations.

The majority

of these roads intercept mule deer winter range and are left
open to travel even though the well or drill site has been abandoned.

w.e.o. Vidakovich

(1974 pers. comm.) believes that pre-

sently there are too many roads in these areas which in turn
provide excessive access to the area during hunting seasons.
2.

Deer-auto collisions along Highway 64.
At present, deer-auto collisions along Highway 64 are a minor but
continuing problem.

Data are lacking regarding highway mortality

in this area, however,

w.e.o. Vidakovich (1974 pers. comrn ,-) re-

ported that the majority of the d~er-auto collisions occur when
deer are on the winter range.

3.

Game damage to farm crops.
In general, the problem of deer and elk damage to farm crops in
Unit 21 is relatively small in comparison to some other units.

w.e.o. Vidakovich

(1974 pers. corom.) reported that he has not had

very many damage complaints in recent years because most problem
areas have been re@edied with fencing.
B.

Fu t.u. e

1.

Oil shale development.
Future oil shale development in the PIceance Basin will undoubtedly

54

�-162-

affect the Rangely area and Unit 21, creating a multitude of
management problems, not only for all species of wildlife but
all components of the environment as well.

For example, a pro-

posed oil pipeline which may exten~.along

the Roan Plateau from

Dere Cabin Ridge to Utah will certainly have adverse effects on
the wide diversity of wildlife now utilizing the plateau areas.

A proposal to dispose of spent oil shale from Piceance tract
C-a into tributaries of Do'uglas Creek by means of a conveyor
system will, if implemented, have far-reaching effects upon
wildlife in Unit 21, particularly mule deer.

Large acreages

of canyon land in the following drainages would be affected:
Left Fork Fourmile Draw, Right Fork East Fourmile Draw, State
Bridge Draw, Vandamore Draw, Philadelphia Creek, and Cow Canyon.
Obviously range carrying capacity wou.ld be reduced for mule
deer (both summer and winter range), elk (summer), and perhaps
wild horses.

Other game species wh Lch would be adversely

affected include chukar, sage grouse, cottontails, and mountain
lion.

Industrialization

of central and northern Piceance Creek areas

may force more game animals, particularly mule deer, to winter
in eastern areas of Douglas Creek, increaSing still further the
pressure on winter range there.

The increased human and vehicular activity throughout Unit 21,
to include residential and commercial developments, associated
with oil shale development, will playa

major role in reducing

the carrying capacity of the existing elk and mule deer winter

55

�-163range and result in loss of habitat for many small game and
non-game species of mammals, birds, reptiles, and amphibians.

A more detailed analysis of proposed oil shale development
,

and possible effects on the environment can be found by consuIting Thorne Ec.ological Institute (1973), Tweedy (1971)
and U. S. Dept. of Interior (1973), which are cited in greater
detail in the Reference Section at the end of this discussion.

2. Deer-auto collisons - as related to oil shale development.
It is inevitable that the increased human and vehicular activity
associated with oil shale development will increase deer highway mortality through deer-auto collisions in this area.
tective measures such as fencing and under/overpasses

Pro-

may have

to be developed similar to those used in the Vail area.

P. H. Neil and W. T. McKean
M?y 1974

��-165-

BIG GAME DISTRIBUTION

MAPS

WILDLIFE MANAGEMENTUNIT 21 (DOUGLAS)

Black bear year-round distribution ----------------___________________
Elk summer distribution -----------------------

1

~_____ 2

Elk winter distribution ----------------------------__________________

J

Mountain lion year-round distribution --------------------____________

4

Mule deer summer distribution -------------------------

~_________ 5

Hula deer winter distribution and comparative abundance

6

Wild Horse year-round distribution -------------~--------------------_

7

��1: 2 N.

tiN.·

1: I. s.]

t 2 S.

T :3 S.

t 4 S.·

1 ::,
S

Page 1

�J ./'._

,

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.,t

.,"

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,...• : ...•
-

I:
- 1 It

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�I _-'._

•••

R.I04 W. -f"-~--'.:..c..,.;r-:r-r-=---'---'rtY

r

o
•

o.

...

I

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,-..

L'

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I

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t 55.

Page 3

��;r

I

?'.:C)4·W··

~

J

-.-

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•i":... --::' _.•.~•. ·.J..-----.:...
..

Ai-;··!.~~

•

l. HILDLIFE I1ANA;~~'
2 N. .~"

MULE DEER

k-

5 S.

Page

5

��t 2

t I. s.]

t 2 S.

T 3 S.:

T 4 S.·

1: ~ S

Page 7

�•

�-175-

SMALL GAME MAMMALAND GAME BIRD DISTRIBUTION

MAPS

WILDLIFE MANAGEMENTUNIT 21 (DOUGLAS)

MaR
Small Game Mammals
Cottontail rabbit year-round distribution; comparative abundance _ 1
Red (Pine) Squirrel year-round distribution --------

2

Snowshoe Hare year-round distribution ------------~------------___

3

Game Birds
Blue grouse year-round distribution -------~-~--------------------

4

Sage grouse year-round distributi~

-------------------------_____

5

Chukar year-round distribution --------------------_______________

6

Ring-necked pheasant year-round distribution -----------__________ 7
Band-tailed pigeon summer distribution (variable) -

8

Mourning Dove summer distribution and comparative ab1.Uldance------~9
Water£owl year-round distribution ---------------________________

10

�·/.

-0.

R.I04 If( :.....
-f'-

Z----!..:..::...;..r:;...:;:..!:=:::....:&lt;--.:!.,-l!-(.

r
~ }IILDLIFE

t:
T. 2 N.' (.

l-~.. !'

•'

J,t

t~

t r x '

t 1.. s.]

t 2 S.

T ~ S.'

T.4 S.

! ~ S.

MANAGUvlENT UNIT: 21

�i

.r: .v: \Iff"

R.I04 W ~
. .f'.&lt;--?~---...::.::...r:;:;...:r=:""""-\-h

r.·

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~. WILDLIFE NilNAGEHENTUNIT 23. (DOUGLAS)

l;
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�.#/

I

.-,;-;~~~~-.
~ \~ILDLlFE MJ..NAG£1t1Et'IT UNIT 21
1.2 N. .

l:
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RING-NECKED

!::

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R.I04

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. HILDLIFE

l1ANAGEMENTUNIT 21
110URNING DOVE

l:'
1-:

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Su}WER DISTRIBUTION

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t

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Abundant ~

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Page 10

��-187July, 1975
JOB PROGRESS

State of
Project

REPORT

C:::;;O:.,:LO::::,:RAD=:.;:O
_
W-lOl-R-17

No.

Work Plan No.

Game Range Investigations
Job No.

7

Job Title
Period

l

Reprint Book: Winter Guide to Native Shrubs of the Central
Mountains
Covered:
July 1, 1974 to March 31, 1975

Personnel:

William

T. McKean

_
Rocky

and Paul H. Neil.

ABSTRACT
Work on refurbishing or replacing all illustrations
(107) was completed
during the segment and work on the narrative portions was approximately
one-half done. Details of printing and distribution remain to be done.

��-189-

REPRINT BOOK: WINTER GUIDE TO
NATIVE SHRUBS OF THE CENTRAL ROCKY MOUNTAINS
William T. McKean and Paul H. Neil

P. S. OBJECTIVE
To revise and reprint the book: Winter Guide to Native Shrubs of the
Central Rocky Mountains.

SEGMENT OBJECTIVE
To continue art work and make corrections

to text of book.

METHODS AND MATERIALS
A second conference with Division Publications Chief, Charles Hjelte, in
July 1974 resulted in a major procedural change: namely, to return to the
original plan of remounting photos and line drawings on new illustration
board rather than trying to re-photograph directly the pages in the first
edition. This procedure will ensure higher quality illustrations.
Many fresh specimens were collected in Colorado.
Herbarium specimens
were obtained from Colorado State University, and especially, from the
National Herbarium of the United States Forest Service, now located in
Fort Collins.
Changes in descriptions of shrub species are being entered directly upon
pages of the first-edition book as copy. However, this procedure may not
be acceptable to the printing contractor, in which case additional time
will be required to re-copy these descriptions of 107 shrub species.
Knowledgeable persons throughout the region were solicited for corrections
and changes in the book. Response was good, particularly from Forest
Service personnel.

RESULTS AND DISCUSSION
As of this segment ending, new or refurbished photos or line drawings of
all shrub species (107) have been mounted on illustration board and labels
affixed. Paste-ups for each species are of 6"x9" dimension.
Two diagrammatic
figures concerning leaf and twig characteristics have also been prepared.
Descriptive detail concerning the species have been checked for about half
the plants.
Special effort was made to up-date paragraphs on distribution
and palatability by means of literature review. Considerable work remains
to be done on these items.

�-190-

Work also remains, during the next segment, on: (1) introductory portions
of the book, (2) revision of the keys where possible, (3) revision of the
glossary, index, and a "Useful References" page.
A decision to finance printing costs with Division funds only has been
made by the Director.
Duties related to proof-reading, printing, and
distribution also remain during the next segment.

-;

Prepared by

?;;:.dYcCzne 7.'P/¢~
William T. McKean
Wildlife Researcher

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                  <text>July, 1975

-191-

JOB FINAL REPORT

State of

COLORADO
--------~~~~.~----------

Project No.

W-38-R-29

Work Plan No.

llC

, Job Title:

Evaluation

Period Covered:
Personnel:

Deer-Elk

Investigations

Job No.
of Experimental

2

Elk Harvest Regulations

April 1, 1974 through March 31, 1975

Raymond J. Boyd

ABSTRACT
A rough draft of the manuscript entitled "A Harvest Formula for the White
River Elk Herd" has been submitted as a final report for this job. When
the co-author finishes his rewrite the article will be submitted to the
Journal of Wildlife Management for pUblication.
An article entitled "An
Evaluation of Yearling Bull Elk Hunting Restrictions in Colorado" has been
sent to the Wildlife Society Bulletin for. publication. The article has been
accepted for publication and editing according to referee comments are
currently underway.

��-193-

EVALUATION OF EXPERIMENTAL

ELK HARVEST REGULATIONS

Raymond J. Boyd

P. S. OBJECTIVE
To determine if modified specified permits for elk will enable game managers
to change the bull/age structure of the herd.

METHODS AND MATERIALS
1.

The co-author has the final report which is in the form of a major
article for the Journal of Wildlife Management.
When he finishes his
rewrite, it will be retyped and submitted.

2.

An article on the changes in the bull/age structure of the White River
elk herd under three types of bull hunt regulations will be submitted
to the Wildlife Society Bulletin.

RESULTS AND DISCUSSION
The manuscript entitled "A Harvest Formula for the White River Elk Herd" is
in the hands of the co-author for his rewrite.
His rewrite has been delayed
due to his reassignment to the Planning Section of the Colorado Division of
Wildlife, but should be completed by June 1, 1975.
An article entitled "An Evaluation of Yearling Bull Elk Hunting Restrictions
in Colorado" was submitted to the Wildlife Society Bulletin.
The article was
accepted for publication and is being reworked according to the comments sent
to us by the various referees.

Prepared by

tZV7j:t!~
Raymond J.. oyd I
Wildlife Researcher

.

��-195July, 1975
JOB FINAL REPORT

State of

COLORADO
----~~~~~----------

Project No.

W-38-R~29

Deer-Elk

Investigations

Work Plan No.

14

Job No.

1

Job Title

Middle Park Deer Study - Population Distribution

----------------------~--------------------------

Period Covered:
Personnel:

-------------------------

April 1, 1974 through March 31, 1975.

W. M. Brandes, G. L. Brown, L. H. Carpenter, R. D. Clippinger,
M. C. Coghill, J. Cooney, J. J. Dorrance, P. F. Gilbert, R. B.
Gill, D. Hoart, S. Horn, J. J. Klein, Jr., D. Luce, P. K. Mason,
D. Merriam, K. Moser, L. A. Roper, R. L. Schmidt, B. C. Sigler,
W. Travnicek, M. Ward, J. B. Weir, J. L. Wolfe, and W. B. Woodward.
ABSTRACT

A final draft of a manuscript entitled "Mule Deer Movements and Distribution
in Middle Park Colorado" has been completed and submitted for publication.

��-197July, 1975
JOB FINAL REPORT

State of

COLORADO
--------~~~~----------W-38-R-29

Proj ect No.
h'ork Plan No.
Job Title

14

Job No.

4

---------------------------

Middle Park Deer Study - Physical Characteristics

Period Covered:
Personnel:

Deer-Elk Investigations

and Food Habits

April 1, 1974 through March 31, 1975

L. A. Roper

ABSTRACT

A final draft of the work on the Physical Characteristics portion of this
job has been completed entitled "Kidney Fat and Bone Marrow Fat Values for
Hule Deer in Middle Park, Colorado" and submitted for publication as a
Special Report.
The food habits portion of Job 4 were presented
Segment (W-38-R-27, WP 14, Job 4).

in full in the 1972-73

��-199July, 1975
JOB PROGRESS REPORT

State. of

COLORADO

P:;:-oject
No.

W-38-R-29

Hark Pla!l No.

14

Job Title

Job No.

Middle Park Deer Study - Experimental

Period Covered:
Personnel:

Deer-Elk Investigations
5a
Range Fertilization

April 1, 1974 to March 31, 1975

L. H. Carpenter and o. C. Wallmo

ABSTPJ.•..
CT
Cover measurements were made on the second-year response of native sagebrush
vegetation to combined fertilizer-herbicide treatments applied at six dates
during Spring of 1973. Vegetation responses were similar to those measured
during the first growing season after treatment.
Forb cover was greatest
on plots treated earliest (April 4) and least on plots treated May 31.
Shrub cover was greatest on plots treated April 4 but least on plots treated
June 13. Grass cover was least on plots treated April 4 and greatest on
plots treated May 31. The April 17 treatment appeared to be most promising
for producing an admixture of vegetation species composition beneficial to
mule deer.

��-201-

MIDDLE PARK DEER STUDY EXPERIMENTAL RANGE FERTILIZATION
Len H. Carpenter

P. S. OBJECTIVE
To test the effects of fertilizer on deer forage and deer feeding responses
to fertilization of critical winter range areas.

SEGMENT OBJECTIVE
Investigate the response of deer forage species to combined fertilizer
herbicide treatments applied at different periods during the growing season.

METHODS AND HATERIALS
Methods and materials have been presented
Exceptions are:

previously

by Carpenter

(1974).

1. During the 1974-75 segment it was determined that the control
area for the May 16 treatment in replicate 3 had received nitrogen fertilizer during initial applications.
As a result, it was not a valid control
area so measurements for the five other control areas in replicate 3 were
averaged and the resulting value used for the May 16 control.
2. The 1974-75 data analysis indicated that covariance analyses used
for 1973-74 data were not reducing error terms below those of an analysis
of variance.
Consequently, the covariance analysis was not used and all
data were analyzed by an analysis of variance.

RESULTS AND DISCUSSION
Aerial cover of forbs, measured in August of the second growing season after
treatment. was greatest on plots treated April 4 and least on plots treated
May 31 (Fig. 1). This was very similar to the response of forbs measured in
1973 (Carpenter 1974). Plots treated on June 13 had more forb cover than
those treated on May 31 (Fig. 1). As reported by Carpenter (1974) most of
this increase was attributable to mat penstemon (Penstemoncaespitosus)
(Fig. 3). Sulphur flower (Eriogonum umbellatum) another important forb on
the study area did not show this increase (Fig. 3). Percent change in forb
cover from the mean of the three control plots for each treatment is graphed
in Figure 2. Only the April 4 treatment had more aerial cover of forbs than
did the mean of the controls. The May 31 treatments decreased 94.6 percent
in aerial cover as compared to the mean of the three controls for those
treatments.
Table 1 presents the percent change from mean of controls for
shrubs, forbs, and grasses at all treatment dates for both 1973 and 1974
measurements.

�300

q
250

\
\

\
\
\

\
a:
w

200
GRASSES

&gt;

V

\

u

•...
o
-'
CL

\
\

150

.•.•.....•

~

\\

.........
,0- \

o

.

a:

.--.--.. .....•..

",
'.'.
,
""
" '0...
'~ "

U

~

"

\

100

I

N

o

N
I

\

.........

..•

...........
o,

50

"

" &lt;,
' ,,

FORBS

"0- . -. _.:;-. ~SSHRUBS

,

',0""

APRIL
4

APRIL

MAY

17

2

DATE

OF

MAY

MAY

16

31

'"

... ~ ~

JUNE
13

TREATME NT

Fig ••1'. Aerial cover mea:sured in AugWJto 1974 of gr/llSRS, forb,
and shrubs Ort
plots treated mth 2,4-D herbicide at si~ dates in Spring of 1197J. EaJch trea.tment
value i~ a mean of thre~ replications.

�-203-

Table 1. Percent change from mean of controls for shrubs, forbs, and
grasses for all treatment dates for years 1973 and 1974. Each value
represents a mean of three replications.

Treatment

1973

Shrubs
1974

1973

1974

Grasses
1973
1974

Forbs

April 4

-25.1

-11.5

+18.1

+26.6

-12.8

+ 1.3

April 17

-39.9

+20.2

-27.8

-32.7

+37.1

+90.2

May 2

-47.7

+12.3

-74.2

-73.1

+88.3

+118.6

May 16

-84.3

-46.9

-75.7

-74.5

-13.8

+89.0

May 31

-82.3

-62.6

-87.2

-94.6

-48.4

+214.7

June 13

-66.4

-67.4

-52.6

-78.3

- 4.6

+81.8

Aerial cover of shrubs showed a similar response to treatment dates (Fig. 1).
Considering only treated plots (excluding comparisons with controls) aerial
cover was greatest on plots treated April 4 and least on plots treated June
13. But comparing treatments to controls there was a decrease in shrub
cover from the mean of the control plots for treatment dates April 4. May
16, May 31, and June 13 (Fig. 2, Table 1) and there was an increase in shrub
cover on April 17 and May 2. The April 17 and May 2 increases were partially due to an increase of rabbitbrush (Chrysothamnus vaseyi) (3.8% April
17; 178.9% May 2) (Fig. 3). Big sagebrush (Artemisia tridentata tridentata),
the dominant shrub on the area, also increased 9.7 percent over the mean of
controls on April 17 treatments but decreased 51.1 percent on May 2 treatments. Apparently, the positive response of both rabbitbrush and big sagebrush (relative to controls) at the April 17 treatments and the large increase of rabbitbrush at the May 2 treatments resulted in total aerial cover
of shrubs exceeding the mean of the controls for these two dates.
Aerial cover of rabbitbrush decreased rapidly after the May 2 treatment (Fig.
3). It would appear that rabbitbrush was not harmed by herbicide prior to
May 2 but soon thereafter the effect became quite negative.
The positive
response of big sagebrush at the April 17 date as compared to the controls
is suspected to result from a large growth response of remaining sagebrush
plants and live crown portions of partially killed plants to the soil nitrogen that was added at the beginning of the experiment.
The negative effects
of herbicide at this date may not have been sufficient to overcome the positive
effects of reduced competition (fewer competing plants) and the added nitrogen.
At the next treatment date (May 2) the herbicide did suffiCiently reduce sagebrush cover to a point where the above mentioned positive factors could not
compensate for the decrease.
The relationships between aerial cover of big
sagebrush and total shrubs for the years 1973 and 1974 are graphed in Figure 4.
The 1974 analysis showed that date of herbicide application had a significant
(P&lt;.05) negative linear effect on aerial cover of big sagebrush, shrubs,
sulphur flower, and mat penstemon.

�\

')

+3 0

+200

w

o

+ 100
GRASSES

Z
~

:I:

...-'..A-._._. A... .

U

0

•...

t."'-

' .&lt;,

..............•
0...

Z

w
U
DI:
w
CL

'~'-

. -'--b,-. -' -.~-0SHRUBS
RBS

...........
00- - - - ---- _

-100

--C&gt;- -

__ - -

F0

-200

APRIL

APRIL

"

17

OATE

MAY
2

OF

MAY

MAY

16

31

JONE
,3

TREATMENT

Fig. 2. Percent change from meAn of controls for gr~$, shrub, and forb cover for
all treat~~nts. Measurement&amp; mads in August. 1974 and rQpresant means of three ~eplicationso

I

N

~
I

�200

PECA = Pens.temon caes-pitosus

ARTR = Artellisia tridenta ta
160

CHVA

= Chrysothamnu~ vasey1

EROM = Eriogonum umbellatum

IX
W

&gt;

0

U

120

~

·0
...•
e,

o!'RTR

0

••...

IX

-

U

I
N

" ••... ....

80

o
V1

~

I

/~
./'

\
40

\

CHVA
._./
.-.-.0"

ERUM

v-, 0-- ~--o..._
\

0

.

...0 .......•..

_

--

' ,.0 ... . :--'

......

··0

APRIL

APRIL

MAY

MAY

MAY

JUNE

4

17

2

16

31

13

DATE

OF

TREATMENT

Fig. J. Aerial cover measured in August, 1974 of four plant species on plots
treated with 2,4-0 herbicide at six d~t6$ in spring of l~J.
Each treatment
v~ue is a mean of three rep1ic~tions.

�200

160
at:
W

, ...••...••'"

&gt;
o

v--

u
••••

o
...J
e,
o
at:

-

U
~

q

120

,,
\

\
c-,\

\

\

\

'.

\

\'"

80

,

\

,

I

N

o
0'\
I

~

o,

1973
TOTAL SHRUBS

\
\

40

•.....•

.

"-.

c··

c..~

.'
- .... ~

1974
'0-.7- --0 TOTAL
SHRUBS

.. -

"-.'w-' _ -' ---. -... ".'
....•..

-'---

c 1974 BIG SAGEBRUSH

.......
--.-.

1973 BIG SAGEBRUSH

APRIL

APRIL.

MAY

MAY

MAY

J,LJNE

"

17

2

16

31

13

DATE

OF

T ~EATMENT

Fig. 4. Aerial cover of big smgebrush and total shrubs for years 1973 and 1974
on plots treated with 2.4-D herbicide at six dates in spring of 19730 Each
treatment value 15 a mean of three replications.

�-207-

Aerial cover of grasses measured in the second growing season after treatment was greatest on plots treated May 31 and least on plots treated April
4 (Fig. 1). Plots treated April 17, May 2, and May 16, all had similar
aerial cover measurements for grasses.
There was a decrease in grass
cover on June 13 plots as compared to May 31 plots. This decrease corresponded to the diminished effect the herbicide had on forbs at this date
(Fig. 1). Increases in aerial cover of grasses for all treatment dates
(except April 4) as compared to the mean of the control plots are presented
in Figure 2 and Table 1. Western wheatgrass (Agropyron smithii) and muttongrass (fQa fendleriana) were the species responding most to the treatments.
It would appear from Figure 1 that the April 17 herbicide treatments most
closely accomplished the objective of reducing sagebrush cover and minimizing
negative effects on forbs (Carpenter 1974). After this date aerial cover
of forbs decreased rapidly.
The increase in grass cover on plots treated
April 17 would also be considered beneficial to deer based on previous
determinations of mule deer food habits made on winter ranges in Middle
Park (Wallmo and Gill 1973).
Since this is an interim report, ultimate interpretations and statistical
comparisons of treatment dates relative to mule deer winter ranges will be
made in the final report submitted next segment.

,
LITERATURE

CITED

Carpenter, L. H. 1974. Middle Park deer study - range fertilization.
Game
Res. Div., Fed. Aid Proj. W-38-R-28.
Game Res. Rep., July, Part 2.
pp. 183-195.
Wallmo, o. C., and R. B. Gill. 1973. Middle Park deer study - Physical
characteristics and food habits.
Game Res. Div., Fed. Aid Proj. W-38R-27. Game Res. Rep., July, Part 2. pp. 83-103.

Prepared

by

h0-A/ If ~tk,
Len H. Carpente
Asst. Wildlife Researcher

��-209-

July, 1975

JOB PROGRESS REPORT

State of

COLORADO

Proj ect No.

W-38-R-29

Work Plan No.

14

Deer-Elk Investigations
Job No.

8

------------------------------Job Title
Middle Park Deer Study - Experimental Harvest Regulations
------------------~~--~----------------~------------Period Covered: April 1, 1974 through March 31, 1975
Personnel:

D. Freddy, L. Carpenter, L. Roper, R. Hoffman, W. Adrian, H. Riffel,
D. Mueller, J. Tulloch, D. Bowden, J. Neisius, F. DeiTIes, J. Brown,
D. Roberts, J. Klein, D. Clippinger, D. Luce, J. Wolfe, B. Gill,
D. Bartmann, and D. Walsworth.

ABSTRACT
Estimated mean deer density in Middle Park in January, 1975 was 10.217 ± 1.925
(P &lt;.10) deer per square mile of winter range yielding a projected population
estimate of 6,005 ± 1,881 deer (P ~.10). Net productivity based on pre-hunting
sex and age counts was 120 fawns:lOO does. Pre-natal productivity was estimated to be 157 fawns:lOO does. Winter mortality estimate was 420 deer or 4.3
percent of the January, 1974 population. Total estimated harvest during 1974,
including a 20 percent wounding loss, was 2,320 deer, 57 percent below the harvest objective of 5,390 deer. Of those persons hunting during the regular
deer season, 17.4 percent;were successful, while 35.5 percent- were successful
during the late-season. An estimated 87.9 percent of the permitted hunters
participated in the regular season, ,while 73.7 percent participated in the
late-season. Weather, deer distribution, hunter participation, and hunter
success appear too variable to achieve predetermined harvest objectives within
desired limits.

��-211-

MIDDLE PARK DEER STUDY EXPERIMENTAL HARVEST REGULATIONS
Dave Freddy

P. S. OBJECTIVE
To adjust a mule deer population

to a specified

density on a winter

range.

SEGMENT OBJECTIVES
1.

Estimate deer density,
in Middle Park.

structure,

productivity,

and winter mortality

2.

Recommend season regulations and publicize procedures for hunters
obtain permits and participate in the experimental hunt.

3.

Measure and estimate
of hunters.

4.

Determine if the hunting public will participate
to achieve adequate harvests utilizing specified
hunts.

5.

Determine if specified
harvest goals.

to

the harvest by sex and age class and the success

on a continuing basis
permits in post-season

permits are capable of achieving

predetermined

METHODS AND MATERIALS
Methods for estimating deer density, structure, productivity and winter
mortality have been previously reported by Gill (1969a, b, 1971). One change
was the use of a Hewes 500C turbine-powered helicopter to fly census quadrats
in the Troublesome sub-unit and on the second flight of the Muddy Creek subunit because maintenance schedules prevented using the standard Bell B-1
helicopter which was used to fly the Blue River, Williams Fork, Granby, and
Muddy Creek sub-units (first flight).
.
Methods for scheduling and publicizing season recommendations
viously reported by Roper (1973, 1974).

have been pre-

Methods for estimating harvest, success of hunters, and hunter participation
have been previously reported by Roper (1973, 1974). Exceptions are:
permit
harvest.

l.

A mail questionnaire was sent to all regular season either-sex
holders to obtain an estimate of the regular season either-sex

2.

A mail questionnaire was sent to 29 percent of regular season antleredonly permit hunters to estimate the regular season antlered-only harvest.

3.

A mail questionnaire was sent to 50 percent of the late-season
sex permit holders to estimate the late-season harvest.

either-

�-212-

The following equations have been used to estimate the number of deer
harvested and the number of hunters participating in a hunt based on returned survey questionnaires.
1.

Estimating

A.

the harvest:

For sampling from a finite population, that is, the population
Middle Park either-sex deer permit holders;

of

h = number of surveys returned
x
number of deer killed as reported on surveys
N
total number of permits issued
p = x/h = success ratio
reciprocal = (l-p)
x mean harvest = (p)(N)

+ 1

standard error of success ratio

2h
95 percent confidence limits for number of deer harvested;
Ll and L2
B.

= (p ± 1.96 (sp») N

For sampling from an infinite population,
number of permits;

that is, an unlimited

h
number of surveys returned
x = number of deer killed as reported on surveys
N = total number of permits issued
p
x/h = success ratio
reciprocal = (l-p)
x

mean harvest = (p)(N)

sp =~P(lh-

p)

= standard error success ratio

95 percent confidence
Ll and L2 = ~
2.

limits for number of deer harvested;

± 1.96(SP~

N

Estimating hunter participation:
Use the same formulas as above only (x) becomes the number of hunters
(participating) out of (h) (hunters returning surveys) and x/h becomes
a participation ratio.

Methods for determining if specified permits are capable of achieving predetermined harvest goals have been previously reported by Roper (1973, 1974).
Tables found to contain inaccurate data in previous reports have been corrected
and updated in this report.

�-213-

RESULTS AND DISCUSSION

Population

Density and Size

The Middle Park deer population was estimated from helicopter counts of
deer on square-mile sample quadrats from January 23-30, 1975 (Table 1).
Estimated mean deer density for 1975 was 10.217 ± 1.925 (P&lt;.lO) deer per
square mile of winter range. Projecting this density estimate
over 587.7
square miles of winter range yielded a mean population estimate of 6,005 ±
1,881 deer (P~.10), an apparent decline of nearly 38 percent compared to
the 1974 estimate of 9,670 deer (Table 3; Fig. 1).
All five sub-unit sample areas exhibited population decreases compared to
1974 estimates (Table 2). The population estimate indicated declines of
43, 49, 19, 8, and 45 percent in the Muddy, Blue, Williams Fork, Troublesome, and Granby sub-units, respectively.
However, empirical evidence indicated an increasing or at least stable
population level compared to 1974. The expected post-hunt population level,
estimated from 1974 winter and harvest mortality and net productivity implied
an increase in population size (Table 3). Utilizing a conservative net productivity estimate of 70 fawns:100 does, instead of the observed 120 fawns:
100 does (Tables, 4, 6), resulted in an expected 1974 post-hunt population
of 10,548 deer, not indicative of a significant population change. Although
one cannot preclude a population decline, the decline as indicated by the
census appears too large.
One possible explanation for the conservative census estimate
could be
attributable to deer distribution.
During the census, deer appeared to be
aggregated in distribution, probably due to deep or crusting snow conditions.
One basic assumption of the census sampling design is that deer are randomly
distributed throughout the winter range at the time of the census.
The
aggregated distribution negated the validity of this assumption and could
have accounted for the indicated population decline because the probability
of encountering quadrats with no deer would be increased and if deer aggregations were not randomly distributed then the probability of encountering
sections with large numbers of deer could have declined.
The Muddy sub-unit was flown twice during the census. The first flight on
January 23 was considered unacceptable due to poor weather conditions and
resulting poor visibility which hindered detection of deer and quadrat corner
markers.
Good visibility prevailed during the second flight on January 30.
Only high density quadrats were flown during the second flight because the
first flight revealed no deer activity on low density quadrats (Table 1).
The replicate flight showed that general deer distribution in the Muddy subunit had not changed appreciably during the one week interval between flights.
Only quadrats 1M and IBM varied between flights in terms of the absence or
presence of deer (Table 5). However, the distribution of deer in relation to
being on or off a specific quadrat varied considerably.
On the first flight
approximately 65 deer were seen just off quadrat 9M, whereas these deer were
apparently on the quadrat during the second flight.
Similar shifts in deer
distribution apparently occurred on quadrats 8M and 10M (Table 5).

�-214-

The discrepancy between sub-unit popUlation estimates between flights
(Table 5) revealed that a popUlation estimate can be inflated or deflated
by a slight shift in location of deer at the time of the census. Extreme
care must be taken when counting individual quadrats to include only deer
that are actually within quadrat boundaries.
Also, more information is
needed about deer distribution in relation to sampling design and the time
of day that sampling occurs.

Sex and Age Composition

Pre-Season Counts--The 1974 pre-hunting season deer sex and age classification counts were conducted from October 7 through 9 by two observers in a
helicopter.
A total of 251 deer were observed of which 225 were classified.
The ratio of 76 bucks:lOO does was considerably higher than ratios in all
previous years except for 1972 (Table 6). The ratio of 120 fawns:lOO does
was 27 percent higher than any previous pre-season fawn:doe ratio (Table 6).
Observed fawn:doe ratios and buck:doe ratios appear to have increased since
1971 (Fig. 2).

Post-Season Counts--Deer sex and age classifications prior to the 1974 latehunting season were obtained from December 2 through 12 via helicopter and
ground surveys. During ground surveys, four observers classified 793 deer
resulting in ratios of 53 bucks:lOO does and 65 fawns:lOO does (Table 7).
During helicopter surveys, one observer classified 1,417 deer resulting in
ratios of 49 bucks:100 does and 75 fawns:lOO does (Table 7). Post-season
buck:doe ratios apparently declined from 1972 through 1974 (Fig. 3). Fawn:
doe ratios apparently increased from 1971 through 1973 but dec1ine~ in 1974
(Fig. 3). Since 1968, the trend in post-season fawn:doe ratios has tended
to mimic the trend of the census population estimate (Figs. 1, 3). None of
the classification trends have been tested for statistical significance, so
inferences about apparent increases and decreases are only tentative.

Productivity
The pre-natal productivity estimate for 1974, based on 11 fetuses present
in 7 mature road-killed does collected from January through May, 1974, was
157 fawns:lOO does (Table 8). Pre-natal productivity appears to have declined
since 1972 (Table 8) but sample sizes have decreased also and hence the
apparent declines may not be significant.
The net-productivity estimate for 1974 based on the pre-hunting season fawn:
doe ratio was 120 fawns:100 does (Table 6). This was the highest pre-season
fawn:doe ratio observed since 1969. The high ratio was probably due to an
inadequate sample rather than a large increase in productivity.

Winter Mortality
Estimated over-winter mortality for 1974, projected from 15 dead deer found
on 93 randomly located mortality transects, was 420 deer or 4.3 percent of
the January, 1974 popUlation of 9,670 deer (Table 3). This estimated overwinter mortality was slightly lower than the estimate in 1973 (Fig. 4).

�-215-

During 1974, there were records of 12 deer dying from deer-auto collisions,
17 from deer-train collisions, and 1 that was inextricably entangled in a
fence. All of these figures were of deer that were actually found and, as
such, are minimum estimates of actual numbers of deer which died from these
various causes.

Harvest Mortality
Harvest Objective and Permit Numbers--The harvest objective for 1974 in
Middle Park was 5,390 deer including an arbitrary 20 percent inflation-factor
for wounding mortality.
This objective was based on an estimated pre-season
population of 15,390 deer (Table 3) and a desired winter population level
of 10,000 deer. Archery, high-country buck, regular deer season, and late
deer season hunters were all required to obtain a permit prior to hunting in
Middle Park. Regular and late-season deer hunters were restricted to hunting
only in Middle Park.
Of the permits issued, 533 were to archery hunters, 70 were to high-country
buck hunters, 3,543 were to regular season deer hunters, and 5,199 were to
late-season deer hunters.
Of the 3,543 regular deer season permits, 2,861
were antlered-only permits and 682 were either-sex permits (Table 9).
Originally, 2,000 regular season either-sex permits were available by drawing,
but applications were received for only 682 and issued. Antlered-only permits were unlimited in number. All late-season permits were for deer of
either-sex (Table 9).

Harvest Size and Composition--Harvest
size and composition were estimated
from questionnaires returned by hunters.
Of the 682 regular season eithersex permit questionnaires sent, 537 or 78.7 percent were returned.· In the
regular season, 831 antlered-only permit questionnaires were sent, and 537
(65 percent) were returned. Of the 2,600- late-season permit questionnaires
sent, 2,085 (80.2 percent) were returned.
Archery and high-country buck
hunters were surveyed by a state-wide hunter questionnaire.
Only regular
and late-season surveys were conducted independently of the state~ide
survey.
Estimated harvests from these questionnaires indicated archery hunters harvested 28 deer and high-country buck hunters harvested 2 deer in Middle Park
representing 1.4 and 0.1 percent, respectively of the 1974 Middle Park harvest
(Table 10).
The projected 1974 regular season harvest was 541 deer (Table 11) representing
28.0 percent of the total 1974 Middle Park harvest (Table 10). Either-sex
permit holders harvested 184 ± 14 deer (P~.05) while antlered-only permit
holders harvested 357 ± 79 deer (P~.05) (Tables 12, 13). The projected
composition of the regular season harvest was 76.3 percent bucks, 18.9 percent does, 2.9 percent buck fawns, 1.7 percent doe fawns, and 0.2 percent
unknown (Table 11).
During the regular season, 273 deer harvested in Middle Park were checked at
the Idaho Springs check station west of Denver.
The composition of these
deer was 75.1 percent bucks, 17.9 percent does, 5.9 percent buck fawns, and
1.1 percent doe fawns (Table 14). Ninety-six deer were checked within Middle

�-216-

Park and were 68.8 percent bucks, 25.0 percent does, 6.2 percent buck fawns,
and 0.0 percent doe fawns (Table 15). A composite summary of all deer
checked is given in Table 16.
The composition of deer checked at Idaho Springs compared favorably with
the composition of the harvest as reported on returned questionnaires.
These data suggest the proportion of the fawns harvested may be underestimated by the questionnaire, but the differences probably are not significant
(Tables II, 14).
A regression estimate of the regular Middle Park deer season harvest was computed from 203 deer checked at Idaho Springs in Middle Park Game Management
Units 18, 28 and 37 (Gill 1970). This estimate was 826 deer taken during the
regular season
(Gill 1970). The regression estimate has usually estimated
a larger harvest than hunter questionnaires (Gill 1970, Roper 1973).
Mail-in tooth envelopes were issued with each regular season either-sex permit. Only 41 or 22.3 percent of 184 potential tooth envelopes were returned.
No fawns were reported in the tooth envelope sample (Table 19).
The projected 1974 late-season harvest was 1,362 ± 80 deer (P&lt;.05) representing 70.5 percent of the total 1974 Middle Park deer harvest (Tables 10,
17). Composition of the late-season harvest projected from questionnaire
responses was 49.4 percent bucks, 42.7 percent does, 5.1 percent buck fawns,
2.6 percent doe fawns, and 0.2 percent unknown (Table 17). There were no
check stations established for the late-season, so comparisons of harvest
composition between questionnaire responses and check station samples were
not available.
Mail-in tooth envelopes were also issued with each late-season permit. Of
1,362 potential tooth envelopes, 481 or 35.3 percent were returned:
Composition of the harvest based on returned tooth envelopes was 49.5 percent bucks,
46.6 percent does, 2.5 percent buck fawns~ and 1.4 percent doe fawns. Compared to questionnaire results, tooth envelopes appeared to underestimate
the harvest of fawns (Tables 17, 21).
Age pyramids derived from mail-in tooth envelopes and check stations indicated
a young, potentially expanding popUlation, assuming the harvest accurately
reflects the age distribution of the live deer population (Tables 18, 19, 20,
21, 22; Figs. 5, 6, 7). Yearlings comprised 36.3 percent, 2 year-olds 29.8
percent, and fawns 5.1 percent of the combined 1974 Middle Park regular and
late-season harvest (Table 22). The low percentage of fawns in the harvest
is assumed to reflect hunter preference for adult animals and therefore may
not be a true reflection of the number of fawns in the population.
It may
also be that hunters are reluctant to report killing fawns on a mail-in
questionnaire, inflating the percentages of older deer in the reported harvest.
Does were the oldest animals harvested (Fig. 7). Deer older than 10 years of
age were apparently more vulnerable during the late-season than during the
regular season (Figs. 5, 6).
The projected composition of the regular and late-season harvests combined
based on returned hunter questionnaires, was 57.1 percent bucks, 35.9 percent does, 4.5 percent buck fawns, 2.3 percent doe fawns, and 0.2 percent

�-217-

unknown (derived from Tables 11, 17). This compares favorably with the
composition of the harvest based on check station information and returned
tooth envelopes (Table 22).

Hunter Participation and Success--Hunter participation and success were
estimated from returned hunter survey questionnaires.
Of 682 potential
regular season either-sex permit holders, 588 ± 10 (P~.05) or 86.2 percent participated in the regular deer season. The success of all eithersex permit holders was 27 percent while the success of only those who
hunted was 31.3 percent (Table 9).
Of the 2,861 regular season antlered-only permit hunters, 2,525 ± 79
(P~.05) (88.3 percent) participated in the regular deer season. Antleredonly permit holders had a 12.5 percent success rate while the success of
only those who actually hunted was 14.1 percent (Table 9).
Of 5,199 potential late-season hunters, 3,830 ± 81 (P~.05) or 73.7 percent participated in the late-season.
Approximately 26 percent of the
hunters who were issued a late season deer permit were successful, while
the success of only those who hunted was 35.5 percent (Table 9). It is
interesting that the success of regular season either-sex permit holders
differed only slightly from the success of late-season either-sex permit
holders (Table 9). Deer were more concentrated during the late-season,
but not necessarily more vulnerable.
The comparable success rates may
indicate that hunters are not capable of achieving high success ratios
in Middle Park unless deer are both concentrated and vulnerable.
Also,
hunters may expect to see deer concentrated along roads during a lateseason, and when this situation does not exist, are less inclined to walk
in order to harvest a deer.

Wounding Loss--On survey questionnaires, hunters were asked to specify
the number of deer they wounded.
Based on hunter responses, wounding loss
during the 1974 regular Middle Park season was 7.5 percent of the reported
harvest of 212 deer (Table 23). If these reports are accurate incorporating
a 20 percent wounding factor into desired harvest levels appears sufficient
for the regular season. Wounding loss during the 1974 late Middle Park
season was 17.2 percent of the reported harvest of 546 deer (Table 23).
Assuming some people did not report wounding a deer or unknowingly wounded
a deer, a 20 percent wounding factor may be insufficient for computing total
harvest mortality during late season hunts.

Success of Controlled Harvest
The 1974 harvest objective of 5,390 deer was not achieved.
The harvest of
2,320 deer (including a 20 percent wounding loss), was 57 percent below the
stated objective.
In 1973, the harvest was 41 percent below the objective
of 3,000 deer (Roper 1974). Based on the results of these two years, it is
questionable whether a controlled permit-type hunt can consistently achieve
a harvest of several thousand deer in Middle Park within acceptable ranges
of precision.
Weather, deer distribution, hunter participation, and hunter
success may all be too variable to allow predetermined harvest objectives to
be achieved within desired limits.

�-218-

LITERATURE CITED
Gill, R. B. 1969a. Middle Park Deer Study - population density and
structure.
Colo. Game, Fish and Parks, Game Res. Sec., Fed. Aid
Proj. W-38-R.
Game Res. Rep., July, Part 1, pp. 105-122.
_____
1969b. Middle Park Deer Study - productivity and mortality.
Colo. Game, Fish and Parks, Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rep., July, Part 1, pp. 123-140.
1970. Middle Park Deer Study - productivity and mortality.
Colo. Game, Fish and Parks, Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rep., July, Part 3, pp. 337-354.
1971. Middle Park Deer Study - population density and structure.
-----Colo. Game, Fish and Parks, Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rep., July, Part 2, pp. 170-188.
Roper, L. A. 1973. Middle Park Deer Study - experimental harvest regulations. Colo. Div. Wildlife, Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rep., July, Part 2, pp. 147-162.
1974. Middle Park Deer Study - experimental harvest regulations.
Colo. Div. Wildlife, Game Res. Sec., Fed. Aid Proj. W-38-R.
Game
Res. Rep., July, Part 2, pp. 197-210.

�Table 1. Numbers of deer counted per quadrat w i t hf.n eight sampling strata in four deer population sub-units, Hiddle Park, Colorado,
1968-1975.
Quadrat
No.

1968

Huddy Creek High Density Strata
1969
1970
1971
1972
1973

1974

_
1975

Quadrat
No.

1968

Huddy Creek Low Density Strata
1969
1970
1971
1972
1973

o

11M

56

6

o

o
o
o
o
o

14M
20M

0
0

1

23M

Ncli
o
o

0

30

24M

NC

0

25M

o

0

o
o

NC

0

o

o
o

9M

124

99

20

28

24

10M

58

39

6

68

39

97

52

77

29M

o

0

o

12M

30

66

2

2

o

52

54

5

30M

NC

15

11

13M

32

46

2

o

5

46

66

o

31M

NC

0

o

o
o
o
o
o
o
o
o
o
o
o
o

15M

NC

87

42

22

35

42

32

4

16M

44

31

28

6

o

o

o

1m

NC

6

5

1

20

12

12

o
o

o

IBM

o
o
o
o

o

56

21

42

0

IH

2

2H

o
o
o
o

3M
4M
5.M

o
o
o
o
o

o
o
o
o
o

6M

8

9

9

7M

64

6

34

8M

32

18

9

2

o
o

o
o

1

o

8

40

13

1

26M

11

60

25

47

27M

o

0

132

125

86

28M

NC

0

3

O·

3

3

o

1

o
o

27

3

o

o

175

136

158

480

370

220

o

o
o

o
o

22H

1

22

5

32N

NC

37

442

478
23.9

o

o
o
o
o
o

o
o
o
o
o
o

o

27

26.0

o

o
o
o

19M

y

o

o
o
o

21M

i

o
o

o
12
4

8.8

o

6.8

1

7.9

24.0

18.5

o

1974

1975

o

o

7

o

o

o
o
o
o
o
o
o
o
o
o
o
o

8

6

0.7

0.5

12
1.0

o
o

o
2

o
o
o
o
o
o

o
o
o
6

o
o
o
6

o
o
o
o
o
o

4

o
o
o
o
o
o
o
1

o
o

11.0

9.3

1.8

3.5

0.0

-----------------------------------------------------------------------------------------------------------------------------------------

I
N
•....
\D
I

�Table 1- Numbers of deer counted per quadrat within eight sampling strata in four deer population sub-units, Middle Park, Colorado,
1968-1975 (continued).
Quadrat
No.

1968

Blue River High Density Strata
1973
1972
1971
1970
1969

1974

1975

Quadrat
No.

1968

1969

Blue River Low Density Strata
1972
1973
1971
1970

1974

1975

1B

14

19

14

47

9

25

14

5

2B

4

8

0

0

0

0

0

0

4B

25

0

0

0

0

0

0

0

3B

10

10

17

13

0

4

4

4

7B

0

1

0

0

6

63

36

0

5B

0

0

0

0

0

0

3

6

8B

76

36

24

145

19

50

65

48

6B

30

0

37

0

0

0

90

78

9B

31

88

77

54

82

156

128

52

12B

32

35

34

15

30

19

87

24

lOB

4

20

18

7

41

28

0

0

16B

0

0

0

0

0

0

0

0

llB

9

4

3

0

29

9

12

6

17B

0

0

0

0

0

0

0

0

13B

134

102

20

34

15

78

48

17

19B

31

36

26

20

2

24

91

27

14B

49

15

83

43

22

74

50

48

21B

0

29

0

0

0

3

0

0

15B

71

105

136

67

64

50

79

31

18B

124

58

49

104

52

102

65

38

20B

28

17

6

0

0

2

2

11

565

465

430

501

339

637

499

256

107

118

114

48

32

50

275

139

= 47.1

38.8

35.8

41.8

28.3

53.2

41.6 21.3

11.9

13.1

12.7

5.3

3.6

5.6

30.6

15.4

j.
Y

=

----------------------------------------------------------------------------------------------------------------------------------------

I
N
N

0
I

��Table 1.
Numbers of deer counted per quadrat within eight sampling strata in four deer population sub-units, Middle Park, Colorado,
1968-1975 (continued).
Quadrat
No.

1968

Troublesome Cr. High Densit1 Strata
1969
1970
1971
1972
1973

1974

1975

Quadrat
No.

1T

43

5

11

0

3

21

15

21

1G

2T

25

41

60

8

24

40

41

46

2G

0

0

0

2

2

0

0

3G

70

54

63

4G

0
0
rt
(\)

3T

0

4T

35

39

51

0

70

51"

50

76

42

43

20

55

30

85

5G

65

44

90

36

40

61

84

17

6G

0

15

7

14

27

31

14

0

7G

21

6T
7T
8T

9

36

8

21

21

31

41

9T

0

0

0

10

0

21

1

0

lOT

26

6

29

7

12

33

16

17

Y

1968

z
0

1969

Granb~ Low Density Strata
1970
1971
1972
1973

1975
0

7

6

0

1

3

0

2

23

0

0

0

0

0

2

0

13

1

11

20

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

11

NC

0

0

0

0

0

0

29

13

2

14

20

11

4.1

1.9

0.3

2.0

2.9

1.6

rt

§
""

Z

=

253

262

298

139

219

365

296

270

11

y

= 25.3

26.2

29.8

13.9

21.9

36.5

29.6

27.0

1.8

1968

1969

1970

1971

1972

1973

1974

1975

yst

=

20.75

15.52

12.29

9.75

9.24

17.14

16.455

10.217

2 S (yst )

=

11.35

4.25

2.55

3.29

1.96

5.24

6.93

3.71

sy

=

3.37

2.06

1.60

1.81

1.40

2.29

2.632

1.925

NC = No count.

1974

I
N
N
N
I

�Table 2.
1975.

Deer population

estimates by population

sub-unit in Middle Park, Colorado, January 1968-

POEulation Sub-unit
Williams Fork
Troublesome
River
Creek

Year

Muddy
Creek

Blue
River

1968

4,101

4,074

843

1969

2,994

3,531

1970

1,429

1971
1972

Granby

Total
Population

1,622

No Count

10,640

774

1,679

134

9,112

3,290

271

1,910

306

7,206

794

3,335

571

891

139

5,730

1,002

2.,259

742

1,404

22

5,429

1973

2,856

4,186

556

2,340

149

10,087

1974

2,272

4,663

622

1,897

216

9,670

1975

1,281

2,370

506

1,729

119

6,005

I
N
N

w
I

�Table 3.

Life equation

for deer in Middle Park, Colorado, winter 1967-68 through 1974-75.

Year

January
Census
Pop. Est.

Winter Loss
Percent
No. Jan. Est.

1967-68

10,640

1,706

1968-69

9,112

1969-70

Pre-Fawn
Pop.

Fawn Reproduction
Percent Increase
No.
of Spring Pop.

Pre-Hunt
Pop.

Harvest
Percent PreNo.
Hunt Pop.

16.0

8,934

3,810

42.65

12,744

2,909

23

582

9,253

9,112

+ 1.5

639

7.0

8,473

4,531

53.48

13,004

4,503

51

901

7,600

7,206

+ 5.2

7,206

1,135

15.8

6,071

2,753

45.35

8,823

2,436

28

487

5,900

5,730

+ 2.9

1970-11

5,730

489

8.5

5,241

2,482

47.36

7,723

516

7

103

7,104

5,429

+23.6

1971-72

5,429

50

0.9

5,379

2,159

40.13

7,538

356

5

71

7,111

10,087

-41.9

1972-73

10,087

647

6.4

9,440

4,715

49.95

14,155

1,771

13

354

12,030

9,670

+19.6

9,250

6,140

1973-74

9,670

1974-75

6,005

420

4.3

Wounding Loss Post-Hunt POE'
Census
(20% of
a
harvest)
Expected
Est.

Percent
b
Difference

66.37

15,390

1,933

13

387

13,070

a

6,005

+54.1

Formula used to calculate expected population level was: P = PI - Mw + b - ~, where P = expected population level; PI = census population
2
n
2
estimate from previous year; Mw = winter mortality; bn = net productivity derived from October pre-hunting season doe:fawn ratios; ~ = harvest
mortality.
b

Percent

difference

indicates

I
N
N

-I'-

the amount that the expected

population

estimate

differs from the quadrat

census estimate.

I

�Table 4.

Life equation for deer in Middle Park, Colorado, January, 1974 through October, 1974.

Population
Data
1974 Winter Population

Winter Mortality
(4.3%)

Spring Population

Pre-Fawning Population

Bucks

Does

Buck
Fawns

Doe
Fawns

Total

24.9%
2,408

39.7%
3,839

20.5%
1,985

14.9%
1,438

100.0%
9,670

40.0%
168

20.0%
84

22.0% *
92

18.0% *
76

100.0%
420

24.2%
2,240

40.6%
3,755

20.5%
1,893

14.7%
1,362

100.0%
9,250

44.7%
4,133

Fawn Crop
(1.2 Pawns /Doe)

Pre-Hunt Population

Note:

26.9%
4,133

100.0%
9,250

55.3%
5,117

33.2%
5,117

55.0%
3,377

45.0%
2,763

100.0%
6,140

21.9%
3,377

18.0%
2,763

100.0%
15,390

Percent Yearlings
Bucks-Does

I
N
N
VI
I

45.8%

26.6%

All unclassified dead deer in winte~ mortality data were excluded.

*Assumption which had to be made:

Because the summer crew did not classify current dead fawns to sex,
the 4 dead fawns found on transects contributed to 40 percent of winter mortality of which 55 percent
(or 22.0%) was assumed to be buck fawns and 45 percent (18.0%) was assumed to"be doe"fawns.

�-226-

Table 5. Comparison of two census flights in the Muddy Creek deer population sub-unit high deer density sampling strata flown on January 23 and
January 30, 1975, Middle Park, Colorado.
Muddy Creek High Density
Quadrat Number

Deer Counted
January 23

January 30

1M

0

0

2M

0

0

3M

0

0

4M

0

0

5M

0

0

6M

0

0

7M

0

1

8M

28

47

9M

10

86

10M

17

77

12M

2

5

13M

0

0

15M

20

4

16M

0

0

17M

0

0

IBM

6

0

19M

0

0

21M

0

0

22M

0

0

32M

0

0

490

1,281

Sub-unit Population

Estimate

�-227-

Table 6. Pre-hunting season deer sex and age classification counts obtained
from helicopter surveys during October 1969-1974, Middle Park, Colorado.

Year

Bucks

Does

Fawns

Total
Classified

1969

195

451

419

1,065

43

93

1970

152

361

258

771

42

72

1971

20

41

30

91

49

73

1972

73

101

91

265

72

90

1973

174

314

273

761

55

87

1974

58

76

91

225

76

120

Ratios Eer 100 Does
Bucks
Fawns

Table 7. Post-hunting season deer sex and age classification counts obtained
from helicopter and ground surveys during November and/or December, 1968-1974,
Middle Park, Colorado.
Type of
Count

Bucks

Does

Fawns

Total
Classified

1968

Aerial
Ground

316
147

558
396

501
~63

1,375
906

57
37

90
92

1969

Aerial
Ground

261
319

490
582

418
505

1,169
1,406

53
55

85
87

1970

Aerial
Ground

211
104

471
231

363
178

1,045
513

45
45

77
77

1971

Aerial
Grotmd

356

783

321

No Counts
1,460

45

41

1972

Aerial
Ground

362
818

693
1,281

540
1,047

1,595
3,146

52
64

78
82

1973

Aerial
Ground

396

674

594

1,664
No Counts

59

Aerial
Grotmd

309
194

1,417
793

49
53

Year

1974

632
363

476
236

Ratios Eer 100 Does
Bucks
Fawns

----

----

88

75
65

�-228-

Table 8. Fetuses per 100 mature female deer in Middle Park, Colorado,
January-May, 1969-1974.
Based on deer collected by selective shooting,
from highway accidents, and from trapping accidents.

Year

Number
Collected
Shooting

Number
Collected
Highway

1969

33

8

0

41

193

1970

33

9

0

42

164

1971

33

8

10

51

161

1972

32

7

0

39

187

1973

0

26

0

26

162

1974

0

7

0

7

157

Number
Collected
Trapping

Total
Deer

Fetuses:lOO

Females

Table 9. Hunter participation and success for the 1974 Middle Park October 26November 3, and December 14-22 deer seasons based on returns from hunter survey
questionnaires.
.Regular .Season ~.
(October 26-Nov. 3)

Late Season
(Dec. 14-22)

Total Number of Permits Issued
Ant1ered-Qnly Permits
Either-Sex Permits

3,543
2,861
682

5,199
0
5,199

Total Projected Hunters
Antlered-Qn1y Permits
Either-Sex Permits

3,113
2,525 ± 79*
588 ± 10*

3,830 ± 81*

Percent Hunters Participating
Antlered-Qn1y Permits
Either-Sex Permits

87.9%
88.3%
86.2%

73.7%

Success of All Permittees
Antlered-Only Permits
Either-Sex Permits

15.3%
12.5%
27.0%

26.2%

Success of Those Hunting
Antlered-Only Permits
Either-Sex Permits

17.4%
14.1%
31.3%

*P ~.05.

3,830 ± 81*

73.7%

26.2%
35.5%
35.5%

�-229-

Table 10.

Summary of deer harvested in Middle Park, Colorado during 1974.

Type of
Season

Bucks

Does

Buck
Fawns

Doe
Fawns

Unknown

Total

Percent Total
Harvest

Archery

19

6

3

0

0

28

1.4%

High-country Buck

2

0

0

0

0

2

0.1%

Regular Deer

413

102

16

9

1

541

28.0%

Late Deer

673

582

70

35

2

1,362

70.5%

1,107
(57.3%)

690
(35.7%)

89
(4.6%)

44
(2.3%)

Totals

1,933
(0.1%) (100%)
3

100%

Table 11. 1974 projected regular season harvest by Game Management Unit based
on reported harvest from returned regular season either-sex and antlered-only
hunter survey questionnaires, Middle Park, Colorado.

18

Management Unit
28
27

37

Totals

Bucks

139 (72.8%)

77 (78.6%)

116 (84.1%)

81 (71.1%) 413 (76.3%)

Does

42 (22.0%)

13 (13.3%)

18 (13.0%)

29 (25.4%) 102 (18.9%)

Buck Fawns

5 ( 2.6%)

6 ( 6.1%)

1 ( 0.7%)

4 ( 3.5%)

16 ( 2.9%)

Doe Fawns

4 ( 2.1%)

2 ( 2.0%)

3 ( 2.2%)

0 ( 0.0%)

9 ( 1.7%)

Unknown

1 ( 0.5%)

0 ( 0.0%)

0 ( 0.0%)

0 ( 0.0%)

1 ( 0.2%)

Totals

191 (100%)

98 (100%)

138 (100%)

114 (100%)

541 (100%)

(35.3%)

(18.1%)

(25.5%)

(21.1%)

(100%)

�.-230-

Table 12.
1974 projected regular season either-sex harvest by Game Management
Unit based on reported harvest from returned regular season either-sex hunter
survey questionnaires, Middle Park, Colorado.

Management
18

27

Bucks

27 (33.9%)

13 (43.5%)

Does

42 (53.2%)

BUCK Fawns

Unit
28

37

Totals

9 (29.2%)

17 (38.9%)

66 (35.9%)

13 (43.5%)

18 (58.3%)

24 (52.8%)

96 (52.4%)

5 ( 6.5%)

1 ( 4.3%)

1 ( 4.2%)

4 ( 8.3%)

12 ( 6.2%)

Doe Fawns

4 ( 4.8%)

2 ( 8.7%)

3 ( 8.3%)

0 ( 0.0%)

9 ( 4.8%)

Unknown

1 ( 1. 6%)

0 ( 0.0%)

0 ( 0.0%)

0 ( 0.0%)

1 ( 0.7%)

Totals

79 (100%)

29 (100%)

31 (100%)

45 (100%)

184 (100%)

(42.8%)

(15.8%)

(16.6%)

(24.8%)

(100%)

Table 13. 1974 projected regular season antlered-only harvest by Game Management
Unit based on reported harvest from returned regular season antlered-only hunter
survey questionnaires, Middle Park, Colorado.
Managemen t Unit
18

27

28

37

Totals

Bucks

112 (100%)

64 (92.3%)

107 (100%)

64 (92.3%)

347 (97.2%)

Does

0 ( 0.0%)

0 ( 0.0%)

0 ( 0.0%)

5 ( 7.7%)

5 ( 1.4%)

Buck Fawns

0 ( 0.0%)

5 ( 7.7%)

0 ( 0.0%)

0 ( 0.0%)

5 ( 1.4%)

Doe Fawns

0 ( 0.0%)

0 ( 0.0%)

0 ( 0.0%)

0 ( 0.0%)

0 ( 0.0%)

112 (100%)

69 (100%)

107 (100%)

69 (100%)

357 (100%)

(31. 3%)

(19.4%)

(29.9%)

(19.4%)

(100%)

Totals

�-231Table 14. Deer harvested in Middle Park, Colorado and checked at Idaho
Springs check station, 1974.

18

Management
27

Unit
28

37

Totals

Bucks

60

63

26

56

205 (75.1%)

Does

19

4

9

17

49 (17.9%)

Fawns

10

3

2

1

16 ( 5.9%)

Unknown

3

0

0

0

3 ( 1.1%)

Totals

92

70
(25.6%)

37
(13.6%)

(33.7%)

Table 15.
1974.a

Deer harvested

273 (100%)
74
(27.1%)

in Middle Park, Colorado and checked in Middle Park,

18

Management
27

Unit
28

37

Totals

Bucks

10

8

0

48

66 (68.8%)

Does

10

4

0

10

24 (25.0%)

Fawns

6

o.

0

0

6 ( 6.2%)

Unknown

0

0

0

0

0 ( 0.0%)

Totals

26
(27.1%)

12
(12.5%)

0
(0.0%)

"96 (100%)
58
(60.4%)

a

Hunters checked in Middle Park were given a slip of paper indicating they had
already been checked, therefore minimizing the possibility of deer being double
checked at one of the main check stations.

Table 16. Deer harvested in Middle Park, Colorado and checked at either Idaho
Springs or Middle Park check stations, 1974.

18

Management
27

Unit
28

37

Totals

Bucks

70

71

26

104

271 (73.4%)

Does

29

8

9

27

73 (19.8%)

Fawns

16

3

2

1

22 ( 6.0%)

Unknown

3

0

0

0

3 ( 0.8%)

Totals

118
(32.0%)

82
(22.2%)

37
(10.0%)

132
(35.8%)

369 (100%)

�Table 17. 1974 projected late-season either-sex harvest by Game Management Unit based on reported
harvest from returned late-season hunter survey questionnaires, Middle Park, Colorado.

18

27

Management Unit
28

37

Unknown

Totals

Bucks

357 (47.8%)

72 (46.0%)

92 (52.9%)

149 (54.1%)

3 (37.5%)

673 (49.4%)

Does

330 (44.2%)

72 (46.0%)

65 (37.1%)

112 (40.5%)

3 (37.5%)

582 (42.7%)

Buck Fawns

37 ( 5.0%)

8 ( 4.8%)

15 ( 8.6%)

8 ( 2.7%)

2 (25.0%)

70 ( 5.1%)

Doe Fawns

20 ( 2.7%)

5 ( 3.2%)

2 ( 1.4%)

8 ( 2.7%)

o ( 0.0%)

35 ( 2.6%)

Unknown

2 ( 0.3%)

o ( 0.0%)

o ( 0.0%)

o ( 0.0%)

o ( 0.0%)

2 ( 0.2%)
I

N
W
N

Totals

746 (100%)

157 (100%)

174 (100%)

277 (100%)

8 (100%)

1,362 (100%)

(54.8%)

(11.5%)

(12.8%)

(20.3%)

(0.6%)

(100%)

I

�Table 18. Age and composition of deer harvested in Middle Park, Colorado and checked at Idaho Springs
or Middle Park check stations during the regular deer season, October 26-November 3, 1974. Age determined by tooth replacement and wear and dental cementum.
Age (Years)
Bucks
Percent

F

1

2

3

4

5

6

7

8

9

10+

Totals

10*
5.3

82*
43.6

43
22.9

27
14.4

16
8.5

6
3.2

2
1.1

o

1

1

o

0.5

0.5

188
100%

9*

16*

3

3

0

2

0

o

o

o

o

33

27.3

48.5

9.1

9.1

-

6.0

19

98

46

30

16

8

8.6

44.3
20.8
(Bucks - 80.5%)

Does
Percent
Totals
Percent

13.6
7.2
3.6
(Does -10.9%)

100%
2

o

1

0.9
0.5
(Buck Fawns - 4.5%)

1

o

221
100%

0.5
(Doe Fawns - 4.1%)

*Age based on tooth replacement and wear.

I

N
W
W
I

Table 19. Age and composition of deer harvested in Middle Park, Colorado during the regular deer season,
October 26-November 3, 1974 based on returned either-sex permit mail-in tooth envelopes. Age determined by
dental cementum.
1

F

Bucks

o

Percent
Does

o

Percent
Totals
Percent

o

3

2

Age (Years)
456

7

8

9

10+

Totals

o

0

1

o

16

5

6

3

0

0

1

31.3

37.5

18.8

-

-

6.2

5

10

3

1

2

1

3

20.0

40.0

12.0

4.0

8.0

4.0

12.0

10-

16

6

1

2

2

3

24.5

39.0

14.6

2.4

4.9

4.9

7.3

(Bucks - 39.0%)

(Does - 61.0%)

100%

6.2
0

o

o

25
100%

0

(Buck Fat~s - 0.0%)

1

o

2.4
(Doe Fawns - 0.0%)

41
100%

�Table 20. Age and composition of deer harvested in Middle Park, Colorado during the regular season, October
26-November 3, 1974. Age determined by dental cementum or tooth replacement and wear. Teeth obtained at
check stations or from mail-in tooth envelopes.
Age (Years)
6

7

6

3

o

2.•9

1. 5

F

1

2

3

4

5

10
4.9

87
42.7

49
24.0

30
14.7

16
7.8

9

21

13

6

1

4

1

3

Percent

15.6

36.2

22.4

10.3

1.7

6.9 1.7

5.2

Totals
Percent

19*

108*

62

36

17

10

4

3

7.3

41.2

23.7

13.7

6.5

3.8 1.5

Bucks
Percent
Does

(Bucks - 74.1%)

(Does - 18.7%)

8

9

10+

Totals

o

204
100%

o

58

1

2

0.5

1.0

0

o

100%
1

2

o

262

1.1 0.4

0.8

-

100%

(Buck Fawns - 3.8%)

(Doe Fawns - 3.4%)
I

*A11 fawns and 91 percent of the yearlings were aged by tooth replacement and wear.

N
W
.pI

Table 21. Age and composition of deer harvested in Middle Park, Colorado during the late-season, December 1422, 1974. Age determined by dental cementum•. Teeth.obtained from mail-in tooth envelopes.
Age (Yeat:~
7

8

9

10

11

12

13

14

15

Totals

5

2

2

o

1

o

o

o

o

250

F

1

2

3

4

5

Bucks

12

86

83

21

14

15

Percent

4.8

34.4

33.2 8.4

5.6 6.0

3.6 2.0 0.8 0.8

7

76

76

10

6

Percent

3.0

32.9

32.9 9.5

Totals

19

162

159

Percent

4.0

33.8

33.1 8.9

Does

22

43

13

6

9

4

4

2

100%

0.4
3

2

2

2

1

1

231

4.3 5.6

,2.7 1.7

1.7 0.9 1.3

0.9

0.9

0.9

0.4

0.4

100%

24

15

6

3

3

2

2

1

1

481

3.1 1.9 1.2 0.8 0.6

0.6

0.4

0.4

0.2

0.2 100%

28

5.0 5.8

(Bucks - 49.5%)

9

(Does - 46.6%)

4

(Buck Fawns - 2.5%)

(Doe Fawns - 1.4%)

�Table 22. Age and composition of deer harvested in Middle Park, Colorado in 1974. Age determined by tooth replacement and wear and dental cementum. Teeth obtained at check stations and from mail-in tooth envelopes.

F

1

2

3

4

5

6

Age (Yeat:"~}
789

10

11

12

13

14

15

Totals

Bucks

22

173

132

51

30

21

12

5

3

4

o

1

o

o

o

o

454

Percent

4.9

38.1 29.1

11.2

6.6

4.6

2.6

1.1

0.7

0.9

Does

16

97

89

28

11

17

7

7

4

2

3

2

2

2

1

1

289

Percent

5.5

33.6

30.8

9.7

3.8 5.9

2.4

2.4

1.4

0.7

1.1

0.7

0.7

0.7

0.3

0.3

100%

Totals

38*
5.1

270*

221

79

41

38

19

7

6

3

3

2

2

1

1

36.3

29.8 10.6

5.5

5.1

2.7

12
1.6

0.9

0.8

0.4

0.4

0.3

0.3

0.1

0.1

743
100%

Percent

(Bucks - 58.1%)

(Buck Fawns - 3.0%)

(Does - 36.7%)

100%

0.2

(Doe Fawns - 2.2%)

I

N
W
\J1
I

*50 percent of the fawns and 36.3 percent of the yearlings were aged by tooth replacement and wear.
Table 23. Wounding loss incurred during the 1974 hunting seasons in Middle Park, Colorado based on returned hunter
survey questionnaires.
Deer Reported
Wounded perRe~onse
Hunter Responses

o

Late Season
Dec. 14-22
Either-Sex Permits

Percent
Total Deer
Hunters
Reported
Responding Wounded

Total Deer
Reported
Harvested

Percent
Wounded

1

2

3

4

5

12

o

o

4

o
o

443
441

95.7
93.0

4

145
67

8.3
6.0

868

16

o

o

o

o
o
o

12

o

884

94.3

16

212

7.5

1,335

77

3

2

o

1

1,418

92.3

94

546

17.2

Regular Season
Oct. 26-Nov. 3
Either-Sex Permits
431
Antlered-Only Permits 437
Total Regular Season

Total
Responses

o

�-236-

14

13

12

11

10

-

9

V)

Q

Z

8

-c

V)

:J
7

0
X

t-

6

C¥

w
w

5

Q

4

°\1
1\1
1\] 1
11 1
•

3

68

69

70

71

72

73

74

75

YEAR

Fig.

1.

Meanestimates

of total. Kiddle Park deer population bracketed

by 90 p~roent confidenoe 11m::1.
ts, JanlUll"Y 196&amp;-19'15.

�CIJ
Q)

-237130

0

0

120

0
0

110

CIJ

100

s:

0

90

m

80

::J

CIJ

c

70

~
ro

60

u.

/e
/

,/

Fawns __

V

50
69

71

70

72

73

Bucks--74

Year
Fig. 2. Pre-hunting season deer sex and age ratios obtained from helicopter
surveys during October 1969-1974, Middle Park, Colorado.
(/)

~100
0

o90
~ 80
(/)
70
~
~ 60

-

/

al

V

. 50

~.--...-

- •••

(/)

~ 40

Fawns--

ro 30
u.
.

Buc~s--6'8

7'1

Year

7'2

Fig. 3. post-hunting season deer sex and age ratios obtained during
Novemberand/or December, 1968-1974, Middle Park, Colorado. Ratios based on
ground surveys except in 1973 when only aerial surveys were conducted.

&gt;-

.•...

ro
•...
L.

0

~
c:
Q)

0
L.

Q)

0..

12

10
8
6
4
2
70

7 1

Year
F.ig. 4. Estimated over-winter deer mortality expressed as a percentage of
January population estimates, 1968-1974, Middle Park, Colorado. Based on
dead deer found on 93 randomly located mortality transects.

�-238-

9 VAS
BUCKS

50

40

n= 204

30

20

DOES

10

0

10

20

30

n=58

40

50

PERCENT

Fig. 5.. Age distribution
of deer harvested in Middle Park, Colorado during
the regular season, October 26-November), 1974. year classes 2 through 9
were aged by dental cementum, while 100 percent of the fawns and 91 percent
of the yearlings were aged b1' tooth replacement and wear. Teeth obtained froll
hunter mail-in tooth envelopes and check stations.

�-239-

15 YRS

BUCKS

n

= 250

DOES

n:231

I
50

30

20

10

0
10
PERCENT

20

30

40

50

~g •. 9. Age:distribution of deer h~sted.in
Middle Park; .Co19p~·dur:i:ng .the lateseason, December lq...22, 1974. Age' determ1liecf b;f dental cementum. Teeth
obtained from hunter mail-in tooth envelopes.

�-240-

15 VRS
BUCKS

50

40

n = 454

30

20

DOES

10

0

10

20

n e 289

30

40

50

PERCENT

Fig. 7•• Age distribution
of deer harvested in Middle park, Colorado, 1974.
Age determined by dental cementumexcept for 50 percent of the fawns and )6. J
percent of the 1 year age-class which were aged by tooth replacement and wear.
Teeth obtained at check stations and from hunter mail-in tooth envelopes.

�-241-

July, 1975

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-29

Work Plan No.
Job Title

14

Job No.

9

-------------------------------

Middle Park Cooperative Deer Study - Deer Habitat Evaluation

Period Covered:
Personnel:

Deer-Elk Investigations

April 1, 1974 - March 31, 1975

L. H~ Carpenter, D. L. Baker, O. C. Wa11mo, D. W. Reichert, D. J.
Freddy, D. Walsworth, D. Roberts,J. Brown, W. Rege1in, and R. B.
Gill.

ABSTRACT
During the 1974-75 segment, work on the physical plant at the Junction Butte
Research Center was essentially completed. Thirteen mule deer fawns were
successfully reared, trained, and added to the tame deer herd. The second
year of study on estimating energy requirements of mule deer fawns in winter
was completed. Digestible energy of the pe11eted ration fed the first year
of study (1974) was estimated at 75.01 percent (se ± 1.03). Approximately
a one-to-one ratio between apparent digestible dry matter and apparent digestible
energy of the pe1leted ration was found. An initial attempt was made to estimate forage intake and energy expenditure of free ranging deer during the winter
period. Four, tame mule deer were observed from an observation tower for seven
24-hour periods. Data were recorded on activities and distances traveled.
Diets, bite sizes, and number of bites per second of grazing were estimated
by accompanying the four deer as they grazed. Daily dry matter intake for these
4 deer averaged 765 g. Analyses of horizontal and vertical components of deer
movements are not yet completed, consequently estimates of energy expenditures
are not included in this report.

��-243-

MIDDLE PARK COOPERATIVE
DEER STUDY - DEER HABITAT EVALUATION
L. H. Carpenter

and D. L. Baker

P. S. OBJECTIVE
Develop procedures
to support deer.

for quantifying

the capacity of Middle Park winter ranges

SEGMENT OBJECTIVES
1.

Develop and test a method for estimating
ranging mule deer.

energy expenditures

of free-

2.

Estimate forage intake and energy expenditures
winter months (January-March).

3.

Estimate nutritional quality of ingested forage in terms of in vivo
digestible and metabolizable energy.

of deer during "critical"

METHODS AND MATERIALS

Physical Plant
A complete description of the research facility used in these studies has been
previously described by Gill and Baker (1974).

Development

Acquisition

of a Tame Deer Herd - 1974

of Fawns

Thirteen mule deer fawns were successfully hand-reared during the period June
I-September 1, 1974 at the Foothills Deer Pens, Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, Colorado.
The fawns
were reared for the purposes of (1) investigating the nutritional requirements
and energy budgets of mule deer in winter, and (2) supplementing a tame deer
herd for future nutritional research programs.
Mule deer fawns were obtained from two sources.
Twenty-three were taken from
semi-tame, captive does maintained at the Foothills Deer Pens, Fort Collins,
Colorado.
Twelve were obtained as orphans and collected from various areas in
Colorado by Division of Wildlife Conservation Officers.

�-244-

Fawn Rearing and Mortality
Approximately 3 hours after birth, each newborn fawn was weighed (nearest
gm), vaccinated with 3 cc ofClosttidiumpetfringens
antitoxin, types B,
C, D, and the navel swabbed with tincture of iodine. The fawn was then
returned to the dam for a period of 12-48 hours. At the end of this period
the fawn was taken from the dam and bottle-reared.
Birth weight, date of
birth, sex and weekly growth rates were recorded for each fawn.
Fawn feeding schedules and amounts fed followed procedures outlined by
Reichert (1972). Milk composition fed to fawns in 1974 is shown in Table
1 (Robbins 1973). At 10 days of age, fawns were given free access to a
commercially pelleted concentrate (Table 2) and third-cutting alfalfa hay.

Table 1.

Composition

of milk formula fed to fawns in 1974 (Robbins 1974).

Nutrient

Content Percent*

Water

75.65

Protein

10.14

Fat

5.83

Lactose

7.14

Ash

1.24

*Formulated by adding to each can of evaporated milk 29.5 g. casein, 121.6 g.
tap water.
In addition, 1 m1 of vitamin-mineral
each day until 60 days of age.

mixture was given to each fawn

Mean birth weight for the 23 fawns born at the Foothills Deer Pens in 1974 was
3.33 kg (se ± 0.11). Mean birth weight for nine males was 3.50 kg (se ± 0.29)
and 3.22 kg (se ± 0.32) for 14 females. Mean birth weights did not differ
significantly (P&lt;0.05) between sexes. Birth weights were not available for
the 12 orphan fawns. At the end of 6 months only 13 of the 35 fawns still
survived (38 percent).
The sex of surviving fawns at 6 months was 7 males and
6 females.
Survival for male fawns was 77 percent while survival for female
fawns was 23 percent.
Postmortem examinations were made on 13 of 22 fawns which died during the 1974
fawn rearing period.
Seven fawns died from chronic viral diarrhea and associated
hypoglycemia, two fawns succumbed to pneumonia; one fawn died as a result of
necro-bacillosis
(Fusibacterium mecrophotum) and three of unknown causes.
Gastrointestinal upsets and diarrhea were most evident during the first 2 weeks
of bottle feeding. Neomycin sulfate and Kaopectate proved successful in light
to moderate cases of diarrhea.
Severe cases usually resulted in death. Limited
success was obtained by changing the diarrhetic animal's diet from milk formula
to a fluid electrolyte solution and oral antibiotic until feces were again
normal.

�-245-

Table 2.

Composition analysis of pelleted basal ration.

Ingredient

Percent

Barley, pulverized

10.0

Corn, pulverized

30.0

Milo, pulverized

5.0

Oats, pulverized

7.5

Wheat middlings

6.5

Beat pulp, shredded

2.5

Brewers grain

35.0

Dicalcium phosphate

1.0

Molasses, cane

2.5

Vitamin A, D, E premix

Tr (0.002)

Trace mineral package
(Mg., Zn, I, Fe, Co, Cu, Ca)

Tr (0.005)

(wt •)

------------------------------------------------------------------------------Analysis

(Dry Matter Basis)

Percent

Dry matter

90.59

Crude protein

19.34

Ether extract

2.8.3

Ash

7.86

Cell wall constituents

52.81

Acid-detergent

13.58

fiber

Lignin

3.58

Apparent Dry Matter Digestibility

70.67

Gross energy

kcal/g

4.60

Apparent digestible energy kcal/g

3.45

Apparent metabolizable

3.05

energy kcal/g

100.0%

�-246-

•

Growth Rates and Training
Mean daily growth rates were calculated for 10 fawns (five male; five females) during a 3 month pre-weaning and 2 month post-weaning period. Up to
2 weeks of age all fawns were weighed by placing them in a wooden box on a
balance beam scale and recording weight to the nearest gram. At 2 weeks of
age, fawns were gradually trained to accept weighing while standing on a platform scale located beneath the floor of a wooden handling chute. The fawns
quickly accepted this procedure and would walk into the weigh chute with little
or no forcing from the handler.
Mean daily gain during the first 5 months was 0.19 kg (se ± 0.01) for females
and 0.20 kg (se ± 0.01) for males. Mean daily gain did not differ significantly (P &lt; O.05) between sexes. Daily gain for both sexes was 0.19 kg (se ±
0.01).
Similar growth rates were recorded for captive dam-reared Columbian
black-tailed fawns (Odocoileus hemionus columbianus) over a similar period
(Cowans and Wood 1955). The fawn growth rates for 1974 were greater than the
0.17 kg for males and 0.14 kg for females as reported by Gill and Baker (1974)
for bottle-reared mule deer fawns but less than the growth rates of 0.22 kg
for males and 0.21 kg for females reported for dam-reared mule deer fawns
(Robinette et al. 1973).
For the purpose of this experiment and future projects it was necessary to
train fawns from birth to accept intensive handling.
In order to accurately
measure weekly weight changes, "normal" blood metabolites, and digestion parameters it was essential that all experimental animals accept varying degrees
of confinement with a minimum of excitability.
To accomplish this, each individual fawn was bottle-fed at least once a day
inside both a digestion cage and handling chute. After weaning, each fawn
was placed in a digestion cage each day for gradually extended periods beginning with a minimum of 15 minutes to a maximum of 7 days. Acceptance of
the weighing and blood sampling chute was accomplished by weighing of all
fawns weekly in the weighing facility.
This training schedule began at 2
weeks of age and terminated at 6 months of age.

Energy Requirements

Objectives

and Experimental

of Fawns in Winter

Design

The primary objective of this study was to estimate metabolizable energy requirements of captive mule deer fawns for winter maintenance.
Subsidiary
objectives were: (1) determine minimum dietary energy requirements for survival
of mule deer fawns, (2) determine length of survival (days) on a given level
of energy intake; (3) relate dietary deficiencies to changes in blood metabolites,
and (4) measure and relate weather parameters (ambient temperature, relative
humidity and wind speed) to energy balance.
Methodology followed during the winter of 1974 is presented by Gill and Baker
(1974). During the winter of 1973-74, data relevant to the overall objective
as well as subsidiary objectives (3) and (4) were collected and are being
analyzed.
Objectives (1) and (2) were only partially answered during the winter

�-247-

of 1973-74 because assigned treatment levels did not sufficiently restrict
energy intake to ascertain minimum energy consumption for survival for a
given period of time during winter.
In order to adequately satisfy these
objectives the entire experiment was repeated during the winter months
(January-March) of 1975. Methods and materials for the 1975 winter trial
were followed as described by Gill and Baker (1974), with the exception
that only two treatments (ad libitum and 25 percent of ad libitum) with 6
deer in each treatment were used during the 1975 experiment.
Fawns were
paired primarily on the basis of similar body weights.
Four sets of twins
were used in the experiment and when weights of twins were similar, one fawn
from each set was assigned to a different treatment.
Fawns were randomly
assigned to one of eighteen isolation pens. Differences in treatment means
for blood metabolites, weight changes, and survival are being analyzed using
a paired t-test (Dixon and Massey 1969). Regression analyses (Draper and
Smith 1966) will be used in an attempt to correlate body weight changes,
blood metabolites, intake, maintenance requirements, and survival to changing
environmental conditions throughout the study period.
Although the trials for estimating maintenance energy requirements for mule
deer fawns were conducted in Middle Park in winter, digestion coefficients
and digestible energy (DE) and metabolizable energy (ME) values of the test
ration were not estimated until late spring or early summer. These values
were estimated from in vivo digestion trials conducted in Fort Collins because facilities were-available there. The validity of comparing results of
digestion trials conducted in warm temperatures with performance values from
winter feeding studies is questionable.
Young and Christopherson (1974) reported apparent dry matter digestibility of sheep and cattle diets decreased
as ambient temperatures declined.
Therefore, if deer digestion coefficients
responded to similar changes in ambient temperature, then maintenance requirements obtained for mule deer fawns during the winter, but baseciupon
DE and ME coefficients obtained in summer, would be in error. The apparent
maintenance requirement would be lower than the true maintenance requirement
because erroneously high digestion and metabolizable energy coefficients were
used.
To avoid this source of error, two digestion cages were constructed in 1975
similar in size to those described by Cowans et al. (1969) (Fig. 1). These
cages were transported to the Junction Butte Deer Research Center, Kremmling,
Colorado, to: (1) calculate digestion coefficients of mule deer fawns fed a
pe11eted concentrate ration during winter, (2) measure the effect of level
of intake on apparent dry matter digestibility (ADMD), (3) compare changes in
ADMD to changes in physiological condition of fawns in winter, and (4) test
the hypothesis that for deer, digestion coefficients decrease relative to a
decrease in average ambient temperature by comparing digestion coefficients
calculated in Fort Collins in spring to those calculated in Middle Park during
winter.
Fawns of similar age, weight and training as those used in the digestion trials
of 1974 were used in the 1975 experiment.
A pelleted ration similar to that
fed in 1974 was fed in the 1975 winter trial. Two fawns, one from each treatment, were placed simultaneously each into one of two digestion cages. A 5 day
fecal collection was conducted on each of 12 fawns from January 21 to March 4,
1975.

�-248-

Fig. 1.
Research

Digestion
Center.

cages used during

1975 winter

at Junction

Butte Deer

�-249Estimate of Forage Intake and Energy Expenditure

of Deer

Estimates of forage intake and energy expenditures of free~ranging mule
deer have not received much attention by wildlife researchers.
This portion of the carrying capacity study was initiated in the 1974-75 segment.
As a result, the greatest emphasis of the 1974-75 winter work was directed
toward developing methods and sampling procedures for describing activity
patterns of mule deer through 24 periods in winter.
Knowledge of activity
patterns will permit calculations of total energy expenditures over time
once energy costs have been established for various activities.
Energy
intake will be calculated from total forage intake by adjusting for digestibility and metabolizability of the energy fractions of these diets. Combinations of energy expenditure and energy intake values will result in energy
budgets which can be used to interpret the adequacy of winter forage toward
the maintenance of mule deer.
Four mule deer, all one and one-half years old, were used in this study.
The deer were placed in a 2.4 hectare conditioning pasture on January 10,
1975 and left until February 3 when they were removed to a 4 hectare energy
budget pasture.
After being placed in the energy budget pasture deer were
allowed to adjust for one week. Measurements were begun on February 11.
Deer received no artificial supplement while in either pasture and were forced
to subsist on native forage.
Two types of measurements were made during the study and these were alternated
from week to week. The first type of measurement involved recording observations of deer activity and travel from an observation tower over 24 hour
time periods.
These observations were conducted February 11-14.
Estimates of forage intake constituted
in this study.

the second type of measurements

taken

On the week of February 18-20 deer were accompanied by observers as they grazed
and bite sizes, total number of bites eqten per second, and total number of
bites of each plant species were recorded.
Then 24-hour activity and travel
observations from the tower were repeated the week of February 24-28.
To aid locating deer in the energy budget pasture, arcs and radii were marked
with respect to the tower location (Fig. 2). Four equally spaced arcs were bisected by 11 radii, each 15 degrees apart. Intersections of arcs and radii were
marked with posts and signs identifying arcs 1, 2, 3, or 4. This system permitted reasonably accurate determinations of deer locations over a 24-hour period.
An observation tower was positioned at the intersection of all radii (Fig. 2).
The tower was a portable fire observation tower on loan from the United States
Forest Service. This tower consisted of a glass-enclosed, 8 foot square tower
extending from the roof of a trailer house.
A night viewing device (Model 221) developed by Javelin Division, Appolo Lasers
in Los Angeles, California* was used to aid night observation.
This device

*The identification

of manufacturing companies or brand names does not necessarily
constitute an endorsement of such firms or products by the Colorado Division of
Wildlife.
They are identified here only for the reader's information and
convenience.

�-250-

- 4SM
-SCALE

Fig. 2. Arc and radii pattern used to locate deer trom tbe observation
tower in energy budget pasture.

�-251-

magnified available light 38,000 times and was equipped with a 300 mm
telephoto lens. The four deer were fitted with colored neck-collars equipped with battery-powered lights arranged in varying patterns to aid in
locating and distinguishing individual deer during both day and night
observations.
Binoculars and a variable power spotting scope were used in
daylight observations.
Snow depths were measured at the beginning and end of each week. Measurements were made one-meter south of each of the permanently marked radius
and arc intersections.
Pasture snow crust and hardness were also described
concurrently with depth measurements.
Continuous temperature and relative
humidity measurements were recorded on a hygrothermograph located 50 meters
south of the energy budget pasture. General weather conditions were described
throughout each 24-hour deer observation period.

Forage Intake
Estimates of forage intake were derived from combinations of both ground and
tower observations.
Tower observations provided estimates of total grazing
time in each 24 hour period whereas grour..
d observations provided quantification
of amounts (grams) of food ingested during periods of grazing. A stopwatch
was started as the deer began to graze and not stopped until the deer lifted
its head to end a grazing sequence. Number of bites taken and the seconds
elapsed were then recorded on tape. Number of bites of each plant species
eaten by deer was also recorded. Percentage of total bites comprised by each
plant species was also determined.
Thirteen separate trials averaging 63 minutes each were conducted during the
3 day period. Trials were conducted throughout daylight hours. Observers
would approach deer in the pasture and if they were grazing the trial would
begin immediately.
Observations continued as long as deer grazed.
Size (weLgh t ) of an average bite was determined by two observers each "plucking"
what they considered to be an average bite grazed by deer. Each observer
plucked 20 bites of various plant species each deer consumed and placed them in
a paper bag. Forty-two of these samples were collected. These 42 samples were
taken to the lab and weighed to determine the average weight per bite (wet
weight). The samples were then oven-dried and reweighed to ascertain percent
moisture and dry weight averages per bite.
Average daily intake per deer was calculated by multiplying the total time
spent grazing (seconds) in a 24-hour day by the bites per second to obtain
total number of bites taken per day. This value was then multiplied by the
weight of average bites to obtain average daily intake. Total daily food intake will be converted to total gross energy intake from determinations of
gross energy values from deer winter forage samples collected in 1973. Gross
energy will be converted to digestible energy from existing published information (Robbins 1973) and from trials currently underway in Middle Park relating
digestible dry matter of selected forage species to in vivo digestible energy
(Milchunas, D. G. Master of Science Program, Colo. State Univ.).

�-252-

Energy Expenditure
Energy expenditure estimates will be determined indirectly from measurements of travel distances and activity patterns of the pasture-enclosed
deer. Travel and activity were recorded from the tower during 24-hour
observation periods.
Four observers, each worked a six hour shift, every
day for one week. The four shifts were from 0 to 0600, 0600 to 1200, 1200
to 1800, and 1800 to 2400 hours. Five minute observations were made 4 times
each hour. These five minute periods were fixed within each hour and were
from 5 to 10, 20 to 25, 35 to 40, and 50 to 55 minutes past the hour. This
resulted in actual observation time of 20 minutes each hour or 480 minutes
(8 hours) each 24-hour period. Each hour one deer would be observed for
the four, five minute periods.
The following hour another deer would be
observed, until all four deer had been observed, then the rotation of deer
started again. During each observation period activities were recorded as:
1) bedding, 2) walking or running, 3) standing, and 4) grazing.
Durations
of each activity class were timed with a stopwatch.
Incidental information
on the three other deer, such as time bedded, time up, or any unusual activity
was recorded whenever it occurred.
This was done to provide additional information on activities over the total 24 hour period to compare with the
estimates of these times generated from the sampling procedure.
Detailed maps were prepared of routes traveled by deer during each observation period. These maps contained information from which average distances
traveled and the elevational changes that occurred during a 24-hour day could
be extrapolated.
A detailed contour map of the pasture will be prepared so
that vertical components of travel can be measured.
Travel maps will also be
used to correlate travel by deer to snow depths within the pasture.
Initially,
values for energy expenditure for the various activities will be approximated
from values reported in the literature (Moen 1973; Clapperton 1961; Graham
1964; Hammel 1962; Stevens 1972; Mattfeld 1973). These values will .be summed
and compared to the energy intake values 'to estimate net energy balance of
deer under conditions measured.

RESULTS AND DISCUSSION

Metabolizable Energy Requirements for
Maintenance of Mule Deer Fawns, Winter 1974
To quantitatively assess the maintenance energy requirements of fawns in winter,
it was necessary to determine energy balance by measuring both body weight
changes (kg) and daily intake (gm). In order to express intake in terms of
digestible energy (DE) or metabolizable energy OME) intake it was essential
to estimate the DE and ME values from digestion trials. Apparent dry matter
digestibility of the experimental ration in 1974 was 72.07 percent (se ± 1.24)
while the digestible energy and metabolizable energy value was 75.01 percent
(se ± 1.03) and 66.25 percent (se ± 1.22) respectively.
The percent of digestible energy available for metabolism was 88.25 percent (se ± se). The digestion
coefficients, DE and ME values, did not differ significantly (P&gt;O.05) between
levels of intake. Correlation (r = .94) of apparent dry matter digestibility
and apparent digestible energy was significant (p&lt;0.05) and positive (Fig. 3).

�-253-

80

•

oI!-

&gt;- 75

o
0::::
w

Z

w

W
...J

m 70

Y= 18.86 + .7791 X
r= 0.94

~
CJ)

w

.t:)

n z; 12

0
~

z

65

w

0::::

ct

a.
a.
ct

65
APPARENT

70
DRY

MATTER

75
DIGESTIBILITY

80
(

%)

Fig. J. Relationship between apparent dr,y matter digestibilit7
and a-pparent digestible energy for. 1974 data.

�-254-

Metabolizable energy requirements for winter maintenance were calculated
by combining the measured urinary energy loss and the estimated energy
loss in methane production (Thompson et al. 1973) and subtracting the total
from apparent digestible energy requirements.
The measured urinary energy
loss was 3.78 percent (se ± 1.67) of the gross energy consumed, while 5.49
percent (se ± .19) was lost to methane production.
Regression equations were used to estimate feed requirements for maintenance
by solving for X (average daily feed) when Y (average daily body weight gain
or loss) equals zero. Mean daily feed consumption for the 10 week period
and corresponding body weight (BWO·75kg) changes are presented in Table 3,
along with regression equations relating feed intake to maintenance requirements. These regressions are based on 11 points representing the average
daily intake and average daily body weight change of 11 mule deer fawns for
the entire 10 week period. Expressed in terms of metabolic size the metabolizable energy requirement for maintenance is 135.90 kcal per kg BwD·75 per day.
The regression of average daily bod, weight gain or loss on average daily
metabolizable energy per kg of BwO· 5 for 10 weeks is shown in Figure 4. The
correlation between these two measurements is 0.94 and is highly significant.
In order to estimate maintenance requirements relative to changing environmental conditions during winter, an attempt will be made to establish a
multiple regression model that will relate weekly maintenance requirements
to a weekly weather severity index. A weather severity index has not been
formulated at this time, but will be during the coming segment.

Blood Metabolites
Analysis of blood metabolites relevant to estimating the energy balance and
nutritional status of mule deer fawns in the 1974 and 1975 winter t~ials will
be completed during the coming segment.

ME Reguirements

for Maintenance

- Winter 1975

Data relevant to the objectives stated for the winter of 1975 have been
collected and are being analyzed.

Estimate of Forage Intake and Energy Expenditures

Forage Intake
A total of 816 minutes (13.5 hours) was spent tallying bite counts of forage
ingested by deer during 13 grazing trials. During this period 8,992 bites were
recorded. These bites were consumed during 25,745 seconds of deer grazing
activity. This resulted in a value of 0.35 bites/second or 1 bite every 2.9
seconds. The diet of the four deer for the period February 18-20, 1975 is presented in Table 4. Four species made up 90 percent of the total diet with vetch
(Astragalus convallarius) making up over 50 percent of the diet. Bite sizes
taken by the deer averaged 0.099 g (se ± 0.005) wet weight and 0.090 g (se ±
0.004) dry weight. Average noisture content of the 42 samples was 9.4 percent.
The four deer lost weight steadily during the 6 week period while they were
dependent on native forage. Near the end of the measurement period one deer
became too weak to continue and subsequently died after being removed from the
study pasture.

�-255-

Table 3. Mean daily energy consumption, body weight gain or loss, and
estimated maintenance requirements for 11 fawns for the 10 week winter trial
of 1974.

Deer
No.

DE Intake
kca1/day

DE Intake
kcal/BwO·75

ME Intake
kcal/day

ME Intake
kcal/BWO.75

Body Weight
Gain of Loss
BwO·75

1

3329.03

185.53

2940.25

167.79

+ .015

41

3219.58

190.90

2769.29

164.32

+ .014

24

3326.36

220.44

2934.49

199.43

+ .017

8

2863.77

172.08

2529.33

151.99

+ .007

13

2863.77

186.68

2529.33

164.87

+ .015

55

2863.77

169.10

2529.33

149.37

- .001

52

2863.77

208.98

2529.33

181.92

+ .016

7

1906.16

125.45

1686.20

110.87

- .010

20

1906.16

133.31

1686.20

117.74

- .003

43

1906.16

115.88

1686.20

102.35

- .014

44

1906.16

162.60

1686.20

143.61

+ .001

Coefficient of
Determination
r2

Maintenance
Requirement

Intake

Regression

Equation

ME kcal/BWO.75/day

Y =

•00035 (X) - .04746

.89

135.90

ME/kcal/day

Y =

.00002(X) - .03732

.73

1866.00

DE kca1/BWO. 75/day

Y =

.000315(X) - .048416

.89

153.70

DE/kcal /day

Y =

.000016 (X) - .03675

.74

2296.87

�-256-

Table 4. Diet of four tame mule deer in 4 ha energy budget pasture for period
February 18-20, 1975.

Total Bites

Percent
Total Bites

Vetch (Astragalus convallarius)

4,699

52.3

Big sagebrush

1,383

15.4

1,252

13.9

Food Item

(Artemisia tridentata tridentata)

Bluebunch wheat grass (Agropyron spicatum)
Serviceberry

(Amelanchier alnifolia)

742

8.3

Muttongrass

(Poa fendleriana)

354

3.9

205

2.3

54

0.6

Snowberry

(Symphoricarpos

oreophilus)

Unknown grass
Green rabbitbrush

(Chrysothamnus viscidiflorus)

47

0.5

Needle-and-thread

grass (Stipa comata)

39

0.5

Fringed sage (Artemisia frigida)

38

0.4

Deer fecal pellets

34

0.4

Rocky Mountain juniper (Juniperus scopulorum)

32

0.4

Phlox (Phlox bryoides)

31

0.3

Western wheatgrass

29

0.3

(Agropyron smithii)

Snakeweed

(Gutierrezia sarothrae)

17

0.2

Penstemon

(Penstemon spp.)

13

0.1

10

0.1

Blue grama grass (Bouteloua gracilis)

9

0.1

June grass (Koeleria cristata)

4

T

8,992

100.0

Bottlebrush

Totals

squirrel tail grass (Sitanion hystrix)

�-257-

.03

.02

.01

a::

MAINTENANC

o
z
c
e

X: 135.90

E
O 75
KC.I/KgBW .

.OO;-

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~

to-

.,~

%

e

-.01

~

I 00

1&amp;1

~

Il"\.

-4··11- "'•. ••

&gt;
Q

o
m

~
,

1
1
1
1

-.02

I

-.03

'I

1
1
I
1

-.04

1

I
o

50

100

METABOLIZABLE

150
ENERGY

200

250

INTAKE

Fig. 4. Regression of average daily body weight ~ain or loss on average
daily metabolizable energy intake in kcal/kg BwO· 5. Estimate of maintenance
requirement is 135.90 kcal/kg BwO·75/day.

�-258-

Grazing tower observations indicated deer spent 28.1 percent of their
total time grazing. On a 24-hour basis this would total 24,278 seconds
grazing. This total Eultiplied by the 0.35 bite/second value resulted
in an estimated total of 8,497 bites ingested by deer each day. Average
daily forage intake was 841 g wet weight and 765 g dry weight per day.
These estimates of intake are similar to those Eade by Alldredge et al.
(1974) using Cesium-137 concentrations in tissues and forages of deer
from the Cache la Poudre Drainage, Colorado. Alldredge et al. (1974)
determined a mean intake ratio of 17.3 g (dry weight) per kilogram body
weight/day for adult deer in the winter period. At the end of our
study, average body weight of deer was 49 kg. Multiplying 17.3 g/kg/day
by 49 kg, the estimated average daily intake would be 848 g (dry weight),
only 10 percent greater than the value we obtained from direct forage
observations.

Energy Expenditure
Seven 24-hour days were spent in the observation tower during two separate
weeks of observations.
Using a sampling strategy of 4 five minute observation periods each hour, 3360 minutes (56 hours) of observations were possible.
One five minute observation period was unavoidably omitted, so actual observation time was 3355 minutes.
Summaries of the 5 minute samples indicated deer spent 41 percent of their
time grazing, standing, or moving, and 59 percent of their time bedded. A
summary of total activity observations for the entire 24 hour period (no
sampling involved) indicated that the deer spent nearly 34 percent of their
total time up and 66 percent bedded. This suggested the 5 minute sampling
scheme underestimated bedding time and overestimated "up activities".
These
discrepancies could have been attributable to sampling error or to artifacts
of the sampling procedure.
In some instances the deer being observed in the
5-minute sample period was the only one of the four either active or bedded.
Then when observations were shifted to another deer at the end of the hour,
the new deer was the only deer active or bedded. This anomaly may have partially accounted for the discrepancies between sampled activity observations and
continuous activity observations.
Rotating observations among the four deer
every 5 minute observation period might reduce the magnitude of this difference.
Twenty-four hour activity and grazing patterns of the deer in the pasture are
graphed in Figure 5. Maximum up (not bedded) activity occurred during hour
10 (10:00 a.m.) and hour 17 (5:00 p.m.). Maximum bedding activity occurred
at hour 23 (11:00 p.m.) and throughout the early morning period (1:00-7:00
a.m.). The average length of time spent up by the four deer during both night
(1800-0600 hours) and day (0600-1800 hours) periods was calculated from the
continuous activity records. The average length of time spent up at night was
49.4 minutes (se ± 6.3 minutes) and 129.4 minutes (se ± 12.1 minutes) for the
daytime periods.
Grazing activity closely parallels up activity.
In Figure 5 the difference
between up activity and grazing activity represents time spent standing or
traveling but not foraging. The IDaximum period for these activities as shown
by the graph would be hour 12 (noon) and at hour 17. It is suspected that

�-259-

observer activities during hour 17 contributed to the low grazing total.
Several times during the trials adjustments of neck collars and lights on
the collars was necessitated prior to night observations and as a consequence hour 17 activities were influenced by people in the pasture. The
percentage of time spent each hour in various activities is summarized in
Table 5. Data on horizontal and vertical components of travel during each
observation
have not been summarized. As a result no computations of
energy expenditure will be ~de in this report. These values will be reported in the following segment.

Physical Plant Improvements
Construction of the facilities at the Junction Butte Research Center as
initially designed was essentially completed in the 1974-75 segment. Several
gates were constructed and erected. The entire intensive study area facility
was dog and coyote proofed by attaching poultry mesh aprons to the bottoms of
pasture fences. The pasture complex was checked for small gaps or openings
at the bottom of fences and sealed to prevent predator entry or deer escape.
It was determined that the portable fire observation tower would be superior
to, and less expensive than, construction of a permanent tower. Therefore,
plans for constructing an observation tower were abandoned. A proposed lab
and storage building was not completed but bids have been let and construction
should be completed by next segment. Remaining fencing and gates adjacent to
the storage building will be completed as soon as the building is constructed.
Needs for certain additional facilities became apparent during the past winter.
To facilitate movement of deer between pastures and the pastures and the
isolation pen complex building a narrow alley is planned from the isolation
pens to the gate into the energy budget pasture. This alley will be constructed of steel posts and woven wire and will be located along the west
fence of the behavior-cover pasture. A small loading ramp is planned from
the main alley of the isolation pen complex to each digestion cage. This ramp
will facilitate the entry and exit of deer to and from the cages. A permanent
roof to protect the digestion cages will also be built. Most of the materials
needed for these projects are available. Only manpower and time are necessary
to complete the construction of the facilities. No progress has been made on
installation of an electric power or water system.

�BEDDED
w
~

•-...

,~
,

....•

,,

~

•...

,
I

0
•...

\

"

\

""":
v \

I

\

I

", _'\

\

z

\
\

w

U

\

I

•...
~
w
0..

I

UP

\

.'\

,,

\- __1

UP

\

,

o

\
\

r GRAZI NG

\

I

\

I

\
I

20
10J

I
1/

.,~.'

GRAZING

....

2

3

4

5

6

7

8

9

,
N
0\

10 11 12 13 14 15 16 17 18 19 20 21 22 2324

HOUR
Fig. 5.. Hourly (Mountain Standard Tiae) activity and gradong patterns
tame mule deer in 4 M. energy budgeti pasture during February, 1975.

of four

�-261-

Table 5. Percentage of time spent each hour in various activities by four
tame deer in 4 ha energy budget pasture for periods February 11-14, and
February 24-28, 1975 combined. All times are Mountain Standard Time.
Percent Total
Time Up

Percent Total
Time Bedded

Percent Total
Time Grazing

Percent Up
Time Grazing

1 (1 AM)

15.0

85.0

13.5

83.5

2

15.0

85.0

10.4

64.3

3

31. 7

68.3

23.2

68.0

4

17.9

82.1

14.0

72.9

5

21.4

78.6

18.6

80.6

6

18.6

81.4

13.0

65.1

7

14.3

85.7

13.1

91. 7

8

36.6

63.4

30.2

82.5

9

35.7

64.3

28.7

80.3

10

85.0

15.0

66.6

78.3

11

56.4

43.6

42.0

74.4

12

64.4

35.6

33.3

51. 7

13

60.7

39.3

37.1

61.1

14

39.9

60.1

21.6

54.2

15

32.1

67.9

21.4

66.7

16

73.4

26.6

52.7

71.9

17

78.6

21.4

47.3

60.2

18

73.5

26.5

52.8

71.9

19

57.1

42.9

34.3

60.1

20

42.9

57.1

15.3

35.8

21

32.1

67.9

19.7

61.1

22

40.0

60.0

24.7

61.8

23

7.1

92.9

6.4

89.2

24

35.7

64.3

28.1

78.7

Hour

-

�-262-

LITERATURE

CITED

Alldredge, A. W., J. F. Lipscomb, and F. W. Whicker.
1974. Forage intake
rates of mule deer estimated with fallout Cesium-l 37. J. Wildl.
Manage. 38(3):508-515.
Clapper ton , J. L. 1961. The energy expenditure of sheep in walking on
the level and on gradients.
Nutr. Soc. Proc. 20:31-32.
Cowan, R. L., E. W. Hartsook, J. B. Whelan, T. A. Long, and R. S. Wetzel.
1969. A cage for metabolism and radioisotope studies with deer. J.
Wildl. Manage. 33(1):204-208.
Cowan, I. McT., and A. J. Wood. 1955. The growth rate of the black-tailed
deer (Odocoileus hemionus columbianus).
J. Wildl. Manage. 19(3):331336.
Dixon, W. J., and F. J. Massey, Jr.
analysis.
3rd ed. McGraw-Hill,

1969. Introduction to statistical
New York, N. Y. 638pp.

Draper, N. R., and H. Smith. 1966. Applied regression
Wiley &amp; Sons, Inc., New York, N. Y. 407pp.

analysis.

John

Gill, R. B., and D. L. Baker. 1974. Middle Park deer study - deer habitat
evaluation.
Colo. Div. Wildlife.
Game Res. Div., Fed. Aid Proj. W-38R-28. Game Res. Rep., July, Part 2. pp.2ll-225.
Graham, N. M. 1964. Energy costs of feeding activities and energy expenditure of grazing sheep. Aust. J. Agr. Res. 15(6):969-973.
Hammel, H. T. 1962. Thermal and metabolic measurements of a reindeer at
rest and in exercise.
Arctic Aerodynamic Lab. Ft. Wainright Alaska.
Tech. Rep.
Mattfeld, G. F. 1973. The effect of snow on the energy expenditure of
walking white-tailed deer. N.E. Sec., The Wildl. Soc., Trans. 30:327343.
Moen, A. N. 1973. Wildlife
and Co., San Francisco.

ecology, an analytical
XVIII + 458 p.

approach.

W. H. Freeman

Reichert, D. W. 1972. Rearing and training deer for food habits studies.
USDA Forest Service. Rocky Mountain For. and Range Exp. Sta. RM Note
208. 7p.
Robbins, C. T.
capacity.

1973. The biological basis for the determination
PhD. Thesis, Cornell University.
239pp.

of carrying

Robinette, W. L., C. H. Baer, R. E. Pil1more, and C. E. Knittle.
1973.
Effects of nutritional change on captive mule deer. J. Wildl. Manage.
37(3):312-326.
Stevens, D. S. 1972. Thermal energy exchange and the maintenance of hemeothermy in white-tailed deer. Ph.D. Thesis, Cornell Univ., Ithaca, N. Y.
23lp.

�-263-

Thompson, C. B., J. B. Holter, H. H. Hayes, H. Silver, and W. E. Urban,
Jr. 1973. Nutrition of white-tailed deer. 1. Energy requirements
of fawns. J. Wildl. Manage.
37(3):301-311.
Young, B. A., and R. J. Christopherson.
1974. Some effects of winter on
cattle. Univ. of Alberta Agri. Bull. Summer, p.3-5.

Prepared by --.!.l~~:::..!...:~'
=,f.lw-~_~C=..;a:..:.....-1;..;~~ ~_' _' __
Len H. Carpenter
Asst. Wildlife Researcher

��July, 1975

-265JOB PROGRESS REPORT

State

0

f

-=-CO.=..L=.O:....:RAD=:.:....:O'-------

Work Plan No.
Job Title

Investigations

14
Job No._-=__~-1_0------------Middle Park Cooperative Deer Study Junction Butte Wildlife Habitat Improvement Project

Period Covered:
Personnel:

Deer-Elk

W-38-R-29

Project No.

April 1, 1974 to March 31, 1975

W. L. Rege1in, 0. C. Wa1lmo, and D. R. Reichert

ABSTRACT
Snowdrift accumulation behind the snowfences on Junction Butte reached a
peak average depth of 30.6 inches on February 16, 1975. Snow depth was
reduced by approximately 12 inches in the shrub stands protected by the
snowfences compared to control areas. The survival of Purshia tridentata
seedlings 2 years after planting was only 15 percent.
The best survival
occurred in seedbeds where all competing vegetation had been removed in
1971. The stands of Agropyron desertorum are growing vigorously and firmly
established.
The Meli10tus officina1is stands were nearly nonexistent in
1974 after good production in 1973.

��-267-

JUNCTION

MIDDLE PARK COOPERATIVE DEER STUDY BUTTE WILDLIFE HABITAT IMPROVEMENT PROJECT
Wayne L. Rege1in

P. S. OBJECTIVE
To increase availability of forage on critical winter ranges by relocating snowdrifts, and determine if changes in vegetative composition
occur due to altering snow depth.

SEGMENT OBJECTIVES
1.

Measure snow depth and density and configuration of snowdrifts in
relation to presence or absence of snowfences and in relation to shrub
stands protected and unprotected from the accumulation of snowdrifts.

2.

Measure occupation by deer of shrub stands protected
from the accumulation of snowdrifts.

3.

Measure the relative use of browse by deer in shrub stands protected
and unprotected from snowdrifts.

4.

Measure the response of native vegetation (composition
in relation to increased or decreased snow depth.

5.

Evaluate the success of forage species planted on sites of created
snowdrifts and on control areas not covered with drifts.

6.

Measure soil moisture with relation to increased or decreased
accumulation.

7.

Extend the p1antings of the most successful forage species on sites
where they were most productive in preliminary trials.

and unprotected

and density)

snow

METHODS AND MATERIALS
The methods and materials used to meet all segment objectives, except
number 7, are reported by Rege1in (1974). The study plan for segment
number 7 was written during the current segment and is included in Appendix

A.
RESULTS AND DISCUSSION
Weather and Snow Measurements
Snowfall on Junction Butte during the early winter of 1974-75 was extremely
slight. No significant amounts of snowfall occurred during November,

�-268-

December or early January. On January 4, no drifts were present behind
any of the snowfences.
Several small snowfalls occurred between January
14 and 20, accompanied by strong winds. By January 26 the snowfences
had accumulated drifts averaging 16 inches in depth, but this was only
about half of the drift depth recorded on this date during the past 3
years. During February the incidence of snowfall increased and the
drifts behind all snowfences reached near normal depths. The maximum
snow accumulation was recorded on February 16, 1975. At this time, the
average depth of created drifts was 30.6 inches. The largest drift was
51 inches deep. The average peak snow depth in the shrub stands protected
by the snowfences reached 16 inches. Relative to control areas, the snowfences reduced snow depth an average of 12.5 inches in the protected shrub
stands.
The density of the snow in the created drifts, protected shrub stands and
control areas was measured on March 2, 1975, using a Mount Rose Snow Sampler.
Mean snow density in the protected shrub stands was 26.3 percent compared
to 37.1 percent in the unprotected control shrub stands. The created drifts
had the highest snow density, 39.4 percent.

Deer Occupancy of Snowfence Sites
Due to the slight snowfall during the winter the south facing slopes remained free of snow throughout the winter.
The deer stayed on these slopes
the entire winter and no tracks were observed on any of the snowfence areas.

Soil Moisture
Measurements of soil moisture were made at monthly intervals during May,
June, July, and August on sites 2, 3 and 4. Bouyocous soil moisture blocks,
buried at depths of 16, 22, 34 and 46 inches, were used to make these measurements.
Calibration of these units has not been completed so the data will
not be included in this report.

Growth of Native Plants
Current annual growth (CAG) on the tagged branches of five shrub species
was measured in September 1974 and is summarized in Table 1. The amount of
CAG was not significantly different (P~.05) between quadrants for any
species. Data on species composition and density of native plants in each
quadrant were collected for the fourth year during July 1974. Analysis of
these data will be done after the final year of data collection in July
1975.

�-269-

Table 1. Average length of CAG per shoot in cm of each species by
quadrant during the 1974 growing season.

Amal

Syor

SEecies
Rice

A (created drift)

1.4

1.5

1.1

C (control)

2.0

1.7

1.1

!/
Prvi

Putr

Quadrant
3.0
4.2

B

(reduced drift)

1.4

1.9

1.2

2.9

D

(normal drift)

1.4

1.9

1.6

1.8

!/ Amal - Amelanchior

Syor - SymEhoricaEEos
alnifolia;
virginianus ; Putr -Purshia
Ribes cereum; Prvi - Pi'linus·

Establishment

oreoEhilus;
tridentata.

Rice -

of Planted SEecies

The survival of bitterbrush (Purshia tridentata) seedlings planted on May
1, 1973 was recorded on July 30, 1974 and the results are summarized in
Table 2. The average survival rate in all seedbeds was 15 percent.
Equal
survival of plants was found in seedbeds covered by the created snowdrifts
and in control seedbeds.
The seedbeds which were cleared of all vegetation
in 1971 had a significantly (P&lt;.05) higher survival rate in 1974 (25%
versus 5%) than the furrowed seedbeds where competing vegetation was not
removed. Application of nitrogen fertilizer at the rate of 120 pounds of
actual N per acre had a negative effect on seedling survival, 21.9 percent
survival in unfertilized seedbeds compared to only 8.1 percent survival in
the fertilized seedbeds. This difference is statistically significant at
the 90 percent confidence level. None of the seedlings exhibited good
vigor and very little growth was noted on any surviving seedling.
The stands of crested wheatgrass (AgroEyron desertorum) established in 1971
continued to exhibit good growth and persistence, but only slight spread of
these plants was noted during the 1974 growing season. Table 3 summarizes
the linear ground cover per 30 foot row for the 3 years in which measurements
have been made. In 1974, the linear ground cover per row in the cleared
seedbeds was significantly higher (P~.05) than in the furrowed seedbeds.
The fertilized seedbeds had more linear ground cover per row than the unfertilized seedbeds and the control seedbeds had more cover than the snowdrift covered seedbeds, but these differences were not statistically significant at the 95 percent confidence level.

�-270-

in various types of seed-

Table 2. Survival of bitterbrush seedlings
beds 2 and 14 months after planting.

Percent of Seedlings Surviving
Ave. 2 Sites
Site 5
Site 1
1974
1974
1973
1973
1974
1973

Seedbed Type

Seedbeds covered by drift

37.5

18.8

20.0

11.2

28.8

15.0

Seedbeds in control

23.7

8.8

39.0

21.2

31.4

15.0

All fertilized

28.8

10.0

16.2

6.2

22.5

8.1

32.5

17.5

42.5

26.2

37.5

21.9

All cleared seedbeds

51.0

26.2

44.0

23.8

47.5

25.0

All furrowed seedbeds

10.0

1.25

15.0

8.8

12.5

5.0

seedbeds

All unfertilized

seedbeds

Table 3. Establishment of Agropyron
and 3 years after planting.

desertorum

in seedbed types 1, 2,

Total Feet of Linear Ground
Cover Per 30-Foot Row
·1974
1973
1972

Seedbed Type

Seedbeds covered by snowdrift

9.91

12.92

12.96

Seedbeds in control

9.70

15.75

16.84

All fertilized

10.66

16.67

17 .27

8.95

12.00

12.54

All cleared seedbeds

14.48

18.36

19.28

All furrowed seedbeds

5.12

10.32

10.54

All unfertilized

seedbeds
seedbeds

The yellow sweetc10ver (Me1i1otus officinalis) planted in the fall of 1971
produced excellent stands in 1973 and most plants produced large quantities
of seed. Since yellow sweetclover is a biennial, natural reseeding must
occur to perpetuate the stands. Measurements in 1974 indicated that few of
the seeds produced seedlings and the stand is in danger of extinction.

�-271-

Measurements
germinate.

will be made again in 1975 to determine

if more seeds

Native revegetation of the seedbeds which were cleared of all plants
in 1971 is continuing to occur, but not at the rapid rate exhibited
during the second growing season after clearing.
Table 4 summarizes
the reinvasion by native species in the different type of seedbeds.
The seedbeds covered by the created drifts continued to have the greatest number of native plants per unit area.

Table 4. Average number of native plants per square foot plot on
cleared seedbeds 1, 2, and 3 years after clearing.
The average is
obtained from 90 lxl foot plots per seedbed.
Average number of plants
Eer sguare foot

1972

1973

1974

All drift covered seedbeds

3.42

6.95

7.10

All control seedbeds

1.58

3.30

4.42

2.84

6.57

6.32

2.17

4.53

5.56

4.00

7.33

7.87

1.·00

2.07

3.28

Drift covered seedbeds,
Control seedbeds,

site 1

site 1

Drift covered seedbeds,

site 5

Control seedbeds, site 5

Experimental

Snowmelt

A manuscript entitled "The use of carbon black to produce mid-winter snowmelt on deer winter ranges" by W. L. Regelin and o. C. Wallmo has been
prepared and will soon be published as a U. S. Forest Service Research Note.

Prepared

by

Iclyt"'Wa

of.

~iL

e L. Regelin

��-273Study No. FS-R~1-l251.21
Amendment No. 2
APPENDIX

A

STUDY PLAN

SEEDING DEER HINTER RAl~GES
IN HlDDLE PARK, COLORADO

by
Wayne L. Regelin

Suhmitted:

~~ /1-&lt;.,',"?,-"
(/
j~{
~I/,£j Date /;2 _;.&gt;&lt;'7 '/'Biologice/ Technician
;/
-

r·,

',,'\

~'

-.

-,

7/

Da te,_~_--,,--......;I
&gt;

) "

Revie\ved:'
.':,. ..~'.
Chief, Big Game Research
Colo. Division of Wildlife

Date~W
Date /
Principal Hildlife Biologist

Approved :.
__ ...:::.('.....;?::.....,_((_(_f_{.....;L_(~_,
Project Leader

J_~(_i_l._,~_._i

_

/

I

;'
i"?
.:

)

��-275-

THE PROBLEH
Most critical deer winter ranges in western Colorado occur in
windswept,

south~facing slopes where vegetative productivity

low due to shaLl.ow soils and limited soil moisture.

is often

Efforts to

improve deer winter ranges have relied heavily on planting of shrub
species, primarily bitterbrush

(Purshia tridentata).

Successful

establishment of shrubs on such ranges is difficult because the
environmental

conditions found on steep, south-facing

not conducive to shrub growth.

slopes are

If shrubs are successfully

estab-

lished, they need to be protected from grazing for several years,
a task which is often impossible or very expensive.

Compared to

grasses and forbs, shrubs produce only small amounts of available
forage

for SeVeral

critical deer winter ranges with grasses and forbs has received
little attention, even though they are extensively utilized by
deer during ~vinter (Kufeld et a1. 1973).

The purpose of this

study is to determine the feasibility' of improving critical deer
winter ranges by planting a mixture of grass, forbs, and shrub
species.
}liddle Park is a valley in central Colorado dominated by big
sagebrush (Artemisia tridentata) and enclosed by high mountains
which trap all deer within the Park during the winter.

The study

area is on Junction Butte, a small isolated mountain in the lowest
winter range in Middle Park.

The southerly slopes on Junction

�-276-

Butte are heavily occupied by deer during winter because of minimal
snow accumulation.

Bluebunch wheatgrass

(Agropyron spicatum) is

the most abundant species on such slopes.
principally

green rabbitbrush

horsebrush

(Tetradymia canescens).

Shrubs occur only sparsely,

(Chrysothamnus viscidiflorus)

and

These southerly slopes offer

areas where planting of forage species to increase forage supply
can be evaluated.

LITERATURE REVIEW
Several range seeding guides (Plummer et al. 1968, Hull and
Johnson 1955, Holmgren and Basile 1959) suggest that range seeding
be restricted to sites with level terrain and deep fertile soil.
Such a

Lt:!l:UUllll8w.laLiull

cannot be adhered to when attempting

improve critical deer winter ranges by seeding.

to

Sites accessible

to deer during winter, most often south-facing slopes free of deep
snow accumulation,

are generally moderate to steep slopes and

have gravelly or rocky soil of variable fertility.

However, there

is some experience with seeding less productive sites which suggest
that it can be feasible.
Bitterbrush

(Purshia tridentata) has received more attention

than any other shrub species in big gcme revegetation

studies.

It

is reviewed in detail primarily to explain its limitations, which
also apply to other popular browse plants such as Cercocarpus montanus

�-277-

and Anlelanchier alnifolia.

Bitterbrush occurs naturally on many

acres of western range land and has long been considered a very
desirable shrub on big game winter ranges because of its palatability
and high nutrient content (Hubbard 1962).
Spring versus fall planting of bitterbrush has been compared
by Hubbard and Sanderson (1961) in California.

They concluded

there is no one best time to plant and that several factors need
to be considered when choosing the planting date for any site.
Adequate soil moisture during germination and emergence is critical
for successful seedling establishment.

If soil moisture is rapidly

depleted on a site in the spring, fall planting is best because
the seeds are in the ground ready to germinate as soon as the
soil warms up.

Spring planting requires longer lasting spring

soil moisture because the soil must be allowed to dry to a certain
extent before planting.

Spring planted seeds must have their

dormancy broken artificially before planting.

Artificial

strati-

fication can successfully break the seed dormancy, but then the
seeds are too soft to be planted by mechanical means (Hubbard 1958).
Soaking seeds for a short time in thiourea can also break seed
dormancy, but without softening the seeds (Pearson 1957).

Planting

in the fall allows the seed dormancy to be broken naturally by
changes in ground temperature.

However, seeds planted in the fall

are often subjected to severe rodent depredation.

Casebeer

(1954),

�-278-

working in Idaho, found that 98 percent of the hills in a spot-type
fall planting had been dug up by rodents within six days after
planting.

Treatment of seeds with endrin, an insecticide which also

acts as a rodenticide, has proven effective in controlling rodent
depredation on bitterbrush seeds (Brown and Martinsen 1959); but
such treatment is expensive.
Planting depth of seeds affects seedling emergence.

A planting

depth of one-half to two inches is usually recommended, depending
upon soil moisture conditions

(Plummer 1968, Holmgren and Basile 1959).

Hubbard (1956) compared seedling emergence from nine planting depths
in California.

He found little difference in emergence of seeds

planted from one-half to two inches deep.

Seeds planted over t\VO

inches deep did not produce satisfactory results and broadcasting
seeds on the soil surface produced no seedlings.
Initial establishment of bitterbrush stands depends to so:ne
extent upon the amount of plant competition in the seeding sites.
Holmgren

(1954) compared bitterbrush establishment under three levels

of competition.

Interseeding into a cheatgrass dominated range

produced no seedlings.

The second treatment consisted of discing

and later hoeing around each seedling to remove all competition.
this weed-free treatment 62 percent of the seedlings which emerged
had survived for two years and averaged seven inches in height.
Data were not collected after the second year.

On

�-279-

Hubbard

(1964) estimated

that 200 to 600 mature bitterbrush

plants in northern California

are required to feed one laO-pound

deer for one month if it eats only bitterbrush.
of mature bitterbrush
California

The average height

plants ranges from two to six feet in

(Hormay 1943).

Hubbard

(1964) considers

initial estab-

lishment of 500 seedlings per acre as good success and a worthwhile
investment

to improve deer winter range.

Very seldom has such plant

density been achieved even when great care has been taken to remove
competing vegetation.
from bitterbrush

Brown and Martinsen

(1959) report results

seeding on 2,000 acres in north-central

They consider 25 percent of these p1antings

as total failure.

remaining acres support 10 to 60 plants per acre.
one 40-acre plot as having highly successful

Washington.
The

They classified

seedling establishment

three years after planting, but it supported only 270 plants per acre.
Brown and ~~rtinsen

(1959) concede that forage production

of these

plants is not great, but they do have value as a seed source.
However, Hormay

(1943) and Nord (1965) claim plants seldom produce

seeds for 10 years after establishment.
Several years are required before bitterbrush
substantial

forage yields.

protection

from grazing.

Growth rate is highly dependent

upon

Holmgren and Basile (1959) measured

three and five years after planting
Three years after planting

plants contribute

plants

in both fenced and unfenced plots.

the average height of protected

plants was

�-280-

eight inches compared to two inches for grazed plaGts.

Five years

after planting, protected plants averaged 20.75 inches in height
and unprotected plants only 8.25 inches tall.
Hubbard (1964) summarizes the results of bitterbrush

seeding at

19 locations in California.

He rated the survival on each site in 1962

as poor, fair or excellent.

The number of surviving plants per acre

was not reported.

Initial establishment

(first year after planting)

was reported in number of seedlings per acre.

Seven of these sites

were planting in 1954 or 1955; survival was rated as poor on all of
these sites even though initial seedling establishment
212 to 1025 seedlings per acre.

ranged from

Of the 19 sites evaluated only one

was rated as excellent and it had been planted the previous fall.
These data indicate that initial establishment

of large numbers ot

seedlings does not guarantee long-term establishment.

Few studies

have observed seedling survival for more than two or three yea~s.
After reviewing the extensive literature on bitterbrush seeding
experiments

I have concluded that bitterbrush

is not a suitable

species to plant when attempting to improve big game winter ranges.
Rubber rabbitbrush

(Chrysothamnus nauseosous)is

usually considered

an undesirable species and much effort has been expended trying to
eliminate it from western range lands.

However, it has frequently

been identified as a preferred deer food (Kufeld et al. 1973).
PluIDlller
et al. (1968) describe rubber rabbitbrush which grows in
a great variety of soils and sites as a highly nutritious plant
throughout the year.

They state that initial establishment

is good

�-281-

to fair, but growth is rapid and natural spread is abundant after
establishment.
planting.

Plants often reach full growth four years after

Only two attempts to plant rabbitbrush

range have been reported

(Plummer et ale 1970 and Holmgren

In Utah Plummer et al. (1970) transplanted
on three dry land sites.
3,500 plants per acre.

on deer winter

one-year-old

1954).

seedlings

Initial stocking rate was approximately
Seven years after planting,

survival on the

three sites averaged 48 percent and mean height of plants was 39
inches.

The average yield per plant on the three sites was 574, 700,

and 2,350 grams.

Initial establishment

of these plants was aided by

scalping 24 inch wide bands along each row, but competing vegetation
was not disturbed

thereafter.

The plants were not protected

grazing; in fact, the authors mention that rabbitbrush

from

yield was

reduced due to heavy grazi.ng by deer.
Holmgren

(1954) planted rubber rabbitbrush

seeds in Idaho

and observed seedling survival and growth under three levels of
plant competition.

Best seedling survival occurred in weed-free

treat-

ments, but some seedlings survived for two years under heavy competition
with cheatgrass

(Bromus tectorum).

After one growing season, the

seedling height varied from 2.5 to 6.5 inches.
Yellow sweet clover (Melilotus officinalis)
biennial

that grows well on disturbed

moisture

requirement

is a rapid-growing

sites without a high soil

(Plummer et ale 1968).

The second growing

�-282-

season

sweetclover

produces

The major disadvantage

large amounts of nutritous

of this species is that natural reseeding

must occur to perpetuate

the stand.

conditions

the seeds may lie dormant

in the spring

for several years until conditions
exist.

forage.

Yellow sweetclover

Without favorable moisture

conducive

to germination

do

is often planted in combination

with

other grasses or legumes.
Four new strains of alfalfa

(Nedicago sativa) have recently

been developed which are well suited for planting on western
range lands.
Teton--have

These four strains--Rambler,
a rhizomatous

from crown sprouts.

Travoes, Nomad and

root system which allows plants to spread

Rambler appears best suited for planting on

deer winter ranges in Colorado because it is drought resistant,
very winter hardy and persists
with other species

for long periods in association

(Heinrichs and Bolton 1958).

Standard crested wheatgrass
used extensively

to rehabilitate

(Agropyron desertorum)

western range lands because

it is easy to establish and provides nutritious
1968).

has been

forage

(Plummer et al.

It grows well on hot, dry sites where other grasses often fail

to survive.

A very desirable

characteristic

of crested wheatgrass

its ability to begin growth early in the spring.

is

On

winter ranges it can provide deer with succulent green forage before
other plants have begun growth.

�-283-

Gorum (1964) planted yellow sweetclover,
wheatgrass

alone and in mixtures

Lodak alfalfa and crested

in Hontana on abandoned

cropland.

He found that mixtures of either clover or alfalfa with crested
wheatgrass

produced more forage than either

legume or wheatgrass.

Yields two years after planting were over 4,000 pounds per acre.
Best establishment
one inch depth.

and yield occurred when seeds were planted at
Broadcasting

seeds on the soil surface produced

only small amounts of forage.

Seeds planted deeper than two inches

failed to emerge in most instances.

Gomm (1964) also discovered

that crude protein content of grass gro,~ in mixtures with legumes
was higher than when grown in pure stands.
Nichols and Johnson
sweetclover

(1969) inter seeded a mixture of yellow

and crested wheatgrass

Combined grass-sweetclover

on rangelands

production

in South Dakota.

averaged 1$04 pounds per

acre over a five-year period compared to only 750 pounds per acre
of herbage on control areas.

Again the crude protein content of

grass was higher when grown in association
pure stands.

with sweetclover

than in

The crude protein content of yellow sweetclover

remained high throughout

the year.

The lowest value was measured

in February when the crude protein content was 8.5 percent.
Kilcher and Heinrichs

(1966) compared yields of six grass-alfalfa

mixtures planted in southwestern
or intermediate

wheatgrass

Saskatchewan.

Crested wheatgrass

(Agropyron intermedium)

. .-

with alfalfa gave

~.-.... ,--.-.", .~.... .. --,-.-~--.-.-.-- .---".-.~..
,

~.'~-'"

."

.

�-284-

the highest yields.

A mixture of 5.6 kg of crested wheatgrass seed

and 1.1 kg of Lodak alfalfa seed per hectare yielded an average of
853 kg/ha of forage over a seven-year period during extreme drought.
Crested wheatgrass alone produced an average of 717 kg/ha over the
same period.

The alfalfa comprised 12 to 25 percent of the stand.

A seeding rate of one pound of alfalfa in combination with six
pounds of grass seed per acre produced the highest herbage yield in
Saskatchewan

~i£her

and Heinrichs 1968).

They tested five rates of

seeding alfalfa, 0.25, 0.5, 1.0, 1.5 and 2.0 pounds per acre, all
mixed with three pounds of crested wheatgrass and three pounds of
Russian wildrye seeds (Elymus ;unceus).

The mixture containing one

pound of alfalfa per acre out yielded those seeded at 0.25 and 0.5
lb/acre; but there was no difference in yield between the three
highest rates of seeding treatment.

OBJECTIVES
The specific objective of this study is to compare the establishment, spread and herbage yield of planted species using three methods
of planting seeds (drilling, broadcasting and harrowing, and
broadcasting only).

These treatments will be applied to different

plots for 3 consecutive years to determine if treatment response is
influenced by yearly variations in weather conditions.

�-285-

METHODS
Twelve blocks of land, each l5x90 meters, will be located on
Junction Butte in May, 1975.

These blocks of land will have

different slopes and aspects, but similar vegetation composition
and density.

Each block will be randomly assigned one of three

planting dates, September 1975, 1976 or 1977.
be divided into four plots, each 15 m2

Each block will

One of four treatments

will be randonly assigned to each plot within each block.
treatments will consist of:

The

(1) drill rows, (2) broadcast

seeded and harrowed, (3) broadcast only and (4) control plots.
A mixture of three species will be planted in the nine treatment
plot~; control plots will not be disturbed.
to be planted are yellow sweetclover

The three species

(Melilotus officinalis),

Rambler alfalfa (Medicago sativa) and standard crested wheatgrass
(Agropyron desertorum).

Seeding rate in the drill row treatm~nt will

be 4.5, 1.7 and 1.7 kg/ha of grass, clover and alfalfa seeds,
respectively.

Seeding rates in both broadcast treatments will be

three times the drill row rate or 13.5, 5 and 5 kg/ha of grass,
clover and alfalfa.
In both broadcast treatments the seed will be applied using
a small, handheld, whirlybird fertilizer spreader.

Each species

of seed will be applied separately to insure even distribution of

�-286-

seed.

On the broadcast

harrowed

immediately

and harrow treatment the plot will be

after the seed has been broadcast.

The

harrowing will break up the soil surface crust to provide a
better seedbed and cover the seeds with soil.
The drill row treatment will not be planted with a drill due
to the small size of the treatment plots.

To simulate planting

with a rangeland drill the plot will be furrowed using hand tools.
The furrows will be approximately

three inches wide, two inches

deep and spaced 30.5 cm (12 inches) apart.
dispersed

The seeds will be

in the furrow using an agri-planter.

seeds will be planted separately.

Grass and legume

The seeds will be covered by

raking one to two cm of soil over the furrow.

Species Establishment
To determine

the establishment

of each planted species and

the response of the native vegetation

to the disturbance

caused

by planting, aerial cover of all species will be estimated using
the rated microplot

technique

(Morris 1973).

In each treatment

and control plot, the aerial cover of each plant species will be
rated to the nearest one-tenth of the area occupied
inch microplot.
20 microplots)

Two hundred microplots

(10 transects each containing

will be located in a systematic

cover each treatment in a representative

in the lx2

fashion so they

manner.

In the drill row

�-287-

treatments, the transects will run perpendicular

to the drill rows.

Pretreatment measurements will be made prior to planting each series of
blocks.

Post-treatment measurements will be made on each series of

blocks the first and second year after planting.
will be made in late July.

All Cover estimates

The cover ratings for each species will be

addec together for the 200 microplots in each plot.

Cover ratings will

also be compiled for all native species combined and for all planted
species combined.
Analysis of Cover Data
The differences in aerial cover ratings between treatments within
each year will be compared using a randomized block analysis of variance.
The components of variation and degrees of freedom are shown below:
SourcpTreatments (T)
Replicates (R)
TxR

d.f.
3

3
9

This analysis will provide a method of comparing treatments before
conclusion of the study.

However, aft~r aerial cover ratings have been

collected from all three year replications, the three independent yearly
experiments will be pooled to provide a stronger test of treatments and
determine if a treatment by year interaction occurred.

The components of

variation and degrees of freedom for the pooled analysis are shown below:
Source
Treatment (T)
Year (Y)
TxY
Replicates (R)
YxR
TxR
TxYxR

d.f.
3

2
6
3
6
9
18

27 pooled d.f. in error term.

�-288-

Herbage Yield
Standing crop will be measured on all treatment and control
plots the July prior to planting (pretreatment) and the July two growing
seasons after planting.

A Neal Electronics herbage meter will be

used to estimate herbage yield.

This instrument measures capacitance

of all vegetation within the meter's probes which is closely correlated
to weight of that vegetation (Neal and Neal 1973).

A double sampling

system will be used in which a fraction of the meter-read plots will
be clipped to provide an estimate of the relationship between meter
readings and dry herbage weight.
ratio will be 4:1.

The meter-read to clipped plot

Within each treatment plot, 100 plots will be

read with the meter and 20 of these clipped.

All plots will be

located

eaL:h treatment

in

.:!

sYGtcmatic

representative manner.
treatment plot.

fashion

so they cover

in a

Ten transects will be established in each

The first transect will be 0.75 meters from the plot

edge and the next nine transects all 1.5 meters apart.

Ten meter-read

plots will be located along each transect by pacing approximately 1.5
meters and placing the herbage meter next to the right foot of
the meter reader.

On seeded areas placement of the meter is important

in relation to drill rows.

To obtain an unbiased estimate of yield,

the meter will be systematically placed in each of 12 positions
corresponding to the 12 numerals on a clock face.

This procedure

insures that the meter is not always placed between or directly over
dri.ll rows.

�-289-

Two plots in each of the 10 transects will be clipped to obtain
regression estimates of yield.

The plots to be clipped in each

.transect will be determined randomly.
within an area l2x24x18 inches.

The meter senses all vegetation

All vegetation within this three-

dimensional hexaedron will be clipped according to the three-dimensional
clip method described by Currie (1973).

The clipped vegetation will

be divided into two groups, planted species in one sack and all other
species in another sack.

This vegetation will be dried at 1050C

for 24 hours before weighing to obtain dry weight measure of herbage.
For the regression analysis, only the total dry weight will be
used for the Y value.

Meter reading will be the X value.

No

regression will be done on separate sack weights, but the percentage
of total herbage weight comprised of planted species will be
calculated.
Herbage Yield Data Analysis
The herbage yield in each treatment and control plot wilY be
estimated by regression analysis using the Dub Sa~1

computer program.

Past experience indicates that the sample size of 20 meter-read and
clipped plots and 80 meter-read only plots should estimate the
herbage yield within 10 percent of the mean at the 95 percent
confidence level.

II

Analysis of these data will be handled similar to

This program, developed by Dr. J. L. Kovner, Principal

Biometrician, is on file at the Rocky Mountain Forest and Range
Experiment Station, Ft. Collins, Colorado.

_______

•__ '00_- 0"0" __

0'_0

••0

�-290-

the cover data analysis, the same AOV table applies.

However,

covariance analysis will be done on the mean yield in each
treatment plot (yields will be adjusted for pretreatment values).
This procedure will remove one degree of freedom from the error
term.

Tukey's multiple range test will be used to make comparisons

between treatments.

The comparison will

be treatments--l vs. 2,

1 v&amp; 3, 1 vs. 4, 2 vs. 3, 2 vs. 4, and 3 vs. 4.

Scheduling
Layout of blocks

May 1975

Pretreatment vegetation measurements

July 1975, 1976, 1977

Planting treatment plots

September 1975, 1976, 1977

Post-treatment vegetation measurements:
Cover estimates

July 1976, 1977, 1978, 1979

Herbage yield

July 1977, 1978, 1979

Analysis of data and writing manuscripts - Sept.-Dec. 1979
Completion date

January 1980

Responsibility
W. L. Regelin

Investigation leader

D. W. Reichert - Field assistance
O. C. Wallmo

- Consultation

�-291-

Estimated Cost
Man weeks
each year
Field work

12

Compiling and analyzing data

10

Writing reports

6

Equipment and supplies
1,000

Seeds
Mileage and per diem
2/

1,500

Costs are covered by Cooperative Agreement No. 16-216 with

Colorado Division of Wildlife.

�-29218

LITERATURE CITED
Brown, E. R. and C. F. Martinsen.
1959. Browse planting for big game in the State of Washington.
Wash. State Game Dep. BioI. Bull. 12, 63 p.
Casebeer, R. L.
1954. The use of tetramine in bitterbrush
J. For. 52: 829-830.

revegetation.

Currie, P. 0., M. J. Morris, and D. L. Neal.
1973. Uses and capabilities of electronic capacitance
instruments for estimating standing herbage. Part 2.
Sown ranges. J. Br. Grassl. Soc. 28(3): 155-160.
Daubenmire, R. F.
1959. A canopy coverage method of vegetational analysis.
Northwest. Sci. 33: 43-64.
Gomm, F. B.
1964. A comparison of two sweetclover strains and Lodak
alfalfa alone and in mixtures with crested wheatgrR~s
for range and dry land seeding. J. Range Manage. 17:
19-23.
Heinrichs, D. H. and J. L. Bolton.
1958. Rambler alfalfa. Can. Dep. Agric. Publ. 1030, 15 p.
Holmgren, R. C.
1954.

A comparison of browse species for revegetation of biggame winter ranges in southwestern Idaho. U. S. Dep.
Agric. For. Serv., Intermt. For. and Range Exp. Stn.
Res. Pap. 33, 12 p. Ogden, Utah.

Holmgren, R. C. and J. V. Basile.
1959. Improving southern Idaho deer winter ranges by
artificial revegetation.
Idaho Dep. Fish and Game
Wildl. Bull. 3, 61 p.
Hormay, A. L.
1943.

Bitterbrush in California. U.S. For. Serv., Calif.
For. and Range Exp. Stn. Res. Note 34, 13 p. Berkeley,
Calif.

�-293-

Hubbard, R. L.
1956. Effects of depth of planting on the emergence of bitterbrush seedlings.
U.S. For. Serv., Calif. For. and Range
Exp. Stn. Res. Note 113, 6 p.
Berkeley, Calif.
Hubbard, R. L.
1956. Effects of plant competition upon the growth and survival
of bitterbrush seedlings.
U.s. For. Serv., Calif. For. and
Range Exp. Stn. Res. Note 109, 9 p. Berkeley, Calif.
Hubbard, R. L.
1962. The place of browse seeding in game range management.
North Am. Wildl.and Nat. Resour. Conf. Trans. 27: 394-401.
Hubbard, R. L.
1964. A guide to bitterbrush seeding in California.
U.s. For.
Serv., Pac. Southwest For. and Range Exp. Stn., and
Calif. Dep. Fish and Game Resour. Agency, 30 p.
Hubbard, R. L. and B. O. Pearson.
1958. Germination of thiourea-treated bitterbrush seeds in the
field. U.s. For. Serv., Calif. For. and Range Exp. Stn.
Res. Note 138, 6 p. Berkeley, Calif.
Hubbard, R. L. and H. R. Sanderson.
1961. When to plant bitterbrush--spring or fall? U.S. For. Servo
Pac. Southwest For. and Range Exp. Stn. Tech. Pap. No. 64,
21 p. Berkeley, Calif.
Hull~ A. C. and W. M. Johnson.
1955. Range seeding in the ponderosa pine zone in Colorado ..
U.S. Dep. Agric. Circ. 953, 40 p.
Kilcher, M. R. and D. H. Heinrichs.
1966. Performance of some grass-alfalfa mixtures in southwestern
Saskatchewan during drought years. Can. J. Plant Sci.
46: 177-184.
Kilcher, M. R. and D. H. Heinrichs.
1968. Rates of seeding Rambler alfalfa with dryland pasture
grasses. J. Range Manage. 21: 248-249.
Kufeld, R. C., O. C. Wallmo, and C. Feddema.
1973. Foods of the Rocky Mountain mule deer. USDA For.
Servo Res. Pap. fu~-lll, 31 p. Rocky Mt. For.
and Range Exp. Stn., Fort Collins, Colo.

�-294-

Morris, M. J.
1973. Estimating understory plant cover with rated microplots.
USDA For. Serv. Res. Pap. 1C'1-l04,12 p , Rocky Nt. For.
and Range Exp. Stn., Fort Collins, Colo.
Neal, D. L. and J. L. Neal.
1973. Uses and capabilities of electronic capacitance instruments
for estimating standing herbage.
Part 1. History and
Development.
J. Br. Grassl. Soc. 28(1): 81-90.
Nickols, J. T. and J. R. Johnson.
1969. Range productivity as influenced by biennial sweetclover in western South Dakota. J. Range Manage. 22:
342-347.
Nord, E. C.
1965. Autecology of bitterbrush
35: 307-334.

in California.

Ecol. Monogr.

Pearson, B. o.
1957. Bitterbrush seed dormancy broken with thiourea.
J. Range Manage. 10: 41-42.
Plummer, P. A., D. R. Christensen, and S. B. Monsen.
1968. Restoring big game range in Utah. Utah Div. Fish and
Game Publ. No. 68-3, 183 p.
Plummer, P. A., D. R. Christensen, R. Stevens, and K. R. Jorgensen.
1970. Highlights, results and accomplishments of game range
restoration studies. Utah Div. Fish and Game Publ.
No. 70-3. 94 p.

�July, 1975

-295JOB PROGRESS REPORT

State of

COLOPJillO
--------~~~~-----------

Project No.

W-38-R-29

Work Plan No.
Job Title

Personnel:

15

Monitor Potentially

Period Covered:

Deer-Elk

and 30

Job No.

Investigations
la

-------------------------------

Critical Deer-Vehicle

Accident Areas Statewide

April 1, 1974 through March 31, 1975.

James R. Adams, Albert E. Byrne, James D. ~leming, Larry L.
Green, William R. Heicher, David C. Hoart, Richard ~. McDonald
(and all Wildlife Conservation Officers in his area), Sharon L.
McDonnell, J. Kris Moser, John W. Seidel, Susan A. Smith, Dale F.
Reed, and Thomas N. Woodard
ABSTRACT

The number and location of deer-vehicle accidents were monitored in six
general areas of Colorado. A total of 542 deer and 14 elk were killed by
vehicles in these areas. Recommendations for installation of devices to
reduce deer-vehicle accidents were prepared.

��-297-

MONITOR POTENTIALLY CRITICAL
DEER-VEHICLE ACCIDENT AREAS STATEWIDE

Thomas N. Woodard

P. S. OBJECTIVE
To establish statewide, the location and number of deer-vehicle
in areas appropriate to the evaluation of devices.

accidents

SEGMENT OBJECTIVE
1.

Examine potentially critical deer-vehicle accident areas reported
by management and as determined by Job 1.

2.

As appropriate,

record all deer found killed in selected highway

areas.

3.

As appropriate,

estimate deer densities

area.

4.

Measure deer activity along the roadside or in the median of each
selected highway area when feasible.

5.

Summarize the Highway Department's
selected highway area as needed.

in each selected highway

traffic volume data for each

METHODS AND MATERIALS
Methods and materials

have been described by Pojar

(1972).

The locations and numbers of deer-vehicle accidents during 1973 and 1974
on U. S. Highway 6-24 from Eagle to Glenwood Canyon were plotted on general
highway maps for Eagle and Garfield Counties.
Results of spotlight counts
conducted during winter and early spring months of 1974 and 1975 on
Highway 6-24 from Gypsum west 15.6 km to the Eagle-Garfield County line
were graphed. Clover traps (Clover 1956) were used to capture deer
during winter months in 1973-75 from Eagle to the west approximately
8.8 km along the future alignment of Interstate Highway 70 and in 1974-75
from Gypsum to Dotsero adjacent to and north of Highway 6-24. Captured
deer were neckbanded and/or eartagged.
Neckbands placed on deer west of
Eagle were white and west of Gypsum were green. Black numbers and/or
letters were sewed on in three places.
Recoveries and reliable sightings
were documented.
~eadow counts were made in the springs of 1972-74
immediately west of Eagle to document neckband sightings and number of
deer present.

�-298-

DESCRIPTION

O~ AREAS

Highway 6-24 (1-70) Rifle West
See Pojar (1972) and Woodard (1973). Preliminarv planning and design of
the interstate highway which will replace Highwav 6-24 is completed and
construction is in progress from Rifle to the wes t 4.8 km,

Highway 1-70 Eagle East
Pojar (1972) and Woodard

(1973,1974) described

this area.

Highway 1-70 Avon-Wolcott
Pojar (1972) and Woodard

(1973) described

this area.

Durango Area
Pojar (1972) described

this area.

Highway 82 - Glenwood-Basalt
Meyers (1969) described this area. Two 2.44-m fences, 0.40 and 1.77 km
long, were installed adjacent to the h Lghwav in this area in lQ74.

Highway 6-24 Eagle-Glenwood

Canyon

This area extends from Eagle to the ~est approximately 26.6 km to the
Eagle-Garfield County line. The valley floor is irrigated hayland with
some sagebrush (Artemisia tridentata) flats. Adjacent south facing slopes
to the north typically have sagebrush and pinyon-juniper (Pinus spp. _
Juniperus spp.) communities.
Deer winter on the south facing slopes and
utilize the irrigated hay land in spring and winter unless precluded by
snow depth and condition.
Highway 6-24 from Eagle to the west 9.9 km to Gypsum is south of the
Eagle River. Preliminary planning and design of Interstate Highway 70
which will replace Highway 6-24 is completed and the alignment will be
north of the river.
Highway 6-24 from Gypsum to the west to the Eagle-Garfield County line is
north of the Eagle River but the interstate highway alignment is unknown
at the present time.

�-299-

RESULTS AND DISCUSSION
Highway 6-24 (1-70) Rifle West
A total of 31 vehicle-killed deer were documented on 29 km of highway
in the Rifle West study area; an increase of 10.7 percent over the
preceding segment.
Seventy-one percent of the deer were killed from
October 1, 1974 to February 28, 1975.

Highway

1-70 Eagle East

A total of 23 vehicle-killed deer were documented On 13.7 km of highway
in the Eagle East study area. Deer kill was reduced on the portions of
highway opposite the 7.7 km barrier fence (Reed 1975).

Highway

1-70 Avon-Wolcott

A total of 60 vehicle-killed deer were documented on 22.5 km of highway
in the Avon-Wolcott study area; a reduction of 20.0 percent from the
preceding segment.
Deer kill continued to be reduced on the portions of
highway opposite two sections of 2.44-m fence (Reed 1975).

Durango Area
A total of 228 deer and 14 elk were killed by vehicles on the 190 km of
highway on the Durango study area during this segment.
The kill was 114
deer and three elk, 58 deer and three elk, 35 deer and eight ~lk, and 21
deer, on Highways 160 east, 160 west, 550 north, and 550 south of Durango,
resoectively.
Evaluation of the deer underpass under Highway 160 with
associated 2.44-m fenCing west of Durango began during this segment and
was reported by Reed (1975).

Highway

82 - Glenwood-Basalt

Ninety-seven vehicle-killed deer were documented on 29 km of highway in
the Glenwood-Basalt study area; an increase of 40.6 percent from the
preceding segment.
Sixty-seven percent were killed during ~ebruary and
March.
Deer-vehicle accidents were reduced on the portions
two 1.77 km lengths of 2.44-m fencing (Reed 1975).

of highway

opposite

�-300-

Highway 6-24 Eagle-Glenwood

Canyon

Fifteen deer in 1973 and five deer in 1974 were killed by vehicles on
Highway 6-24 from Eagle to the west 9.9 km to Gypsum.
Fifty-eight
deer in 1973 and 25 deer in 1974 were killed by vehicles on Highway 6-24
from Gypsum to the west 15.0 km to the beginning of Glenwood Canvon.
February in 1974 and March in 1975 had the greatest mean number of deer
observations counted by spotlight adjacent to Highway 6-24 from the
Bureau of Land Management Gypsum Recreation Area to the west 6.4 km
(Fig. 1). February in 1974 and 1975 had the greatest combined number of
deer and elk observations counted bv spotlight adjacent to Highway 6-24
from the Colorado River Bridge at Dotsero to the west 5.6 km to the
Eagle-Garfield County line (Fig. 2).
One-hundred-seven
deer of both sexes and various age cLasses were
neckbanded and/or eartagged during winters of 1973-75 from Eagle to
the west.
Seventeen recoveries or sightings have been made to date on
summer or transitional range. One deer moved to the northwest approximately
11 km. Five deer moved north approximately 11 km to Castle Peak. Eleven
deer moved to the east 24 to 37 km. No confirmed recoveries or sightings
of these neckbanded deer have been made south of Highway 6-24 or Interstate
70.
Fifteen deer were neckbanded during winters of 1974-75 from Gvpsum to
Dotsero adjacent to and north of Highway 6-24. One recovery made from
this banding program was south of the highway and east approximately
11 km,
The greatest number of deer observed during meadow counts west of Eagle
was on 22 March 1972 when 124 deer were counted.
These deer w~re on
pastureland which will be bisected by Interstate Highway 70. Recommendations for mitigating structures for this area will be made to the Colorado
Division of Hf.ghways during the next segment.

LITERATURE
Clover, M.R. 1956. Single-gate

CITED

trap. Calif. Fish and Game. 42:199-201.

Myers, GaryT.
1969. An investigation of deer-auto accidents. Colo.
Div. Game, Fish and Parks. Game Res. Div. Fed. Aid Proj. \.J-38-R-23.
Game Res. Rep., July, Part 2. pp. 147-178.
Pojar, Thomas. M. 1972. Monitor potentially critical deer-vehicle accident
areas statewide. Colo. Div. Wildl. Game Res. Div. "Fed. Aid P'ro],
W-38-R-26. Game Res. Rep., July, Part 3. Pp. 305-310.

�-301-

Reed, Dale F. 1975. An evaluation of deer-proof fence length required
to prevent deer movements on or across high speed highways. Colo.
Div. Wildl. Game Res. Div. "Fed. Aid Proj. W-38-R-29 and 30. Game
Res. Rep. In Progress.
Woodard, Thomas N. 1973. An evaluation of deer-proof fence length
required to prevent deer movements on or across high speed highwavs.
Colo. Div. Wildl. Game Res. Div. Fed. Aid Proj. 1..J-38-R-27.Game
Res. Rep., July, Part 3. pp. 197-202.
Woodard, Thomas N. 1974. Monitor potentially critical deer-vehicle
accident areas statewide. Colo. Div. Wi1d1. Game Res. Div. Fed.
Aid Proj. W-38-R-28. Game Res. Rep., July Part 2 pp. 293-298.

1

.__ ..---1Prepared

by

/

W~

.- / I
AJ

,)

/~;

'J.

.

'/

i- (I (y-~-cJ(~

YThomas N. Woodard
Assistant Wildlife

Researcher

�-302-

50
46
44

r-

..-

41
40

-

30

-

-

44

..-

39
r-

24
20
20

-

r-

10 -

.

.
JAN

FEB
1974

MAR

APR

JAN

FEB

MAR

1975

Fig. 1. The mean number of deer counted per month from the Bureau of Land
Management Gypsum Recreation Area to the west 6.4 km.

�-303-

120

0
100

Deer
102
r

U

90

Elk

85
r

80

-

E-&lt;

S
c

u

63
60

-

40

-

~
~

37

37

37

r-

,

23
20

-

,

11

I
I
I

JAN

18

I
I

12

I

1

I

2

I

I,

1&lt;'EB MAR
1974

I

o

0

APR

DEC

o
.TAN

I

()

~
1&lt;'F.B MAR
1975

1&lt;'ig.2. The mean number of deer and elk counted oer month from the
Colorado River Bridge at Dotsero to the west 5.6 km to the Eagle-Garfield
County line.

��-305-

July, 1975

JOB PROGRESS REPORT

State of

COLO~~O
--------~~~~----------W-38-R-29

Project No.
~rknan

~.

and 30

15

Deer-Elk Investigations
Job No.

6a

--------------------------------

Job Title

Evaluation

Period Covered:

April 1, 1974 through March 31, 1975

Personnel:

of Deer Underpasses

Thomas N. Woodard, James D. Fleming, Arthur J. Gresh, and Dale
F. Reed

ABSTRACT
Six underpasses were monitored for deer use and supplemental early morning
observations were made at the Vail deer underpass.
Eight deer passages
occurred through four Interstate 79 concrete box underpasses.
Passages to
the south were considered disadvantageous since the 2.44-m fence was installed only on one side of the highway.
Forty-six passages occurred through
the arch deer underpass located west of Durango.
No flight reaction was
observed during 97 percent of westbound traffic when deer were present at the
Vail deer underpass.
The behavior modes of deer exits from the underpass were
71 (n=25), 23 (n=8), and 6 (n=2) percent, trotting, walking, aud bounding,
respectively.

��-307-

EVALUATION

OF DEER UNDERPASSES

Dale F. Reed

Mule deer response to an underpass 3.05 m (10 feet) by 3.05 m and 30.48 m
(100 feet) long under Interstate 70 has been reported (Reed et al. 1975).
This study purports to evaluate underpasses of different sizes and design
and to further evaluate deer behavioral responses when feasible.

P. S. OBJECTIVE
Determine the use of underpasses by deer in critical highway
areas and improve the effectiveness of these structures.

deer kill

SEGMENT OBJECTIVE
Measure deer use of underpasses
associated with their use.

and the extent of behavioral

reluctance

METHODS AND MATERIALS
Six underpasses which were potentially used by deer were monitored for use
by track-count methods previously described (Reed 1971). Four concrete
box underpasses under 1-70 east of Eagle, an open 3-span structure at
Chaffee Gulch north of Ridgway, and the arch (structural plate pipe arch)
deer underpass west of Durango were evaluated.
In addition, ~upplemental
observations were made in the early morning hours at the Vail deer underpass. A 20X spotting scope was use4 to observe deer at the underpass from
about 0.5 km away. Vegetational cover and camouflaging were utilized.
The
instances of vehicle caused flight reaction at the underpass entrance and
the mode of movement at the exit (walk, trot, or bound) were tallied. Use
of and behavioral response to this underpass during a previous 4-year study
has been reported (Reed et al. 1975).

RESULTS AND DISCUSSION
The four underpasses east of Eagle received few to no passages (Table 1).
No deer tracks were detected in or at the ends of underpass No.1.
Six
passages occurred through underpass No.2, three to the south and three to
the north, when the 2.44-m (8 foot) vehicle gate was left open by the landowner. Of special interest are the passages that occurred to the north since
they took place without the effect of a 2.44-m (8 foot) fence along the south
side of the interstate.
Only the north side of the interstate is fenced with
2.44-m fencing in the Eagle east area. The north end of underpass No.3
(Fig.l)

�-308-

received deer pressure (188 deer tracks) for over a 1 1/2-month (June 14Aug 9) period.
During this relatively dry period two passages occurred
through the underpass to the south. Underpass No. 4 received no passages.
Most of the activity at this location was deer movements north of and
parallel to the 2.44-m fence whether or not the 2.44-m vehicle gate was
open or closed.
The Chaffee Gulch underpass was checked on1v three times. These checks
yielded 14 passages, seven east and seven west, and were considered
sufficient indication of deer movement by the Division of Highwavs to
j us t Lfy replacing the sub-standard bridge with an equally open+snan bridge.
No 2.44-m fencing was associated with this structure.
The arch deer underpass located west of Durango and the Cherrv Creek campground, received 46 passages.
The deer activity adjacent the underpass
has been light. Considerable human and cattle use has possib1v affected
deer movements in this area.
Daily observations (n~22) were made at the Vail deer underpass from June
3 to June 25. Of 198 instances of vehicles traveling west when deer were
near the underpass entrance and the west bound lanes, 193 did not elicit
flight reaction (Hediger 1950) from 35 different individuals or groups
(range 1 to 11 deer). Four vehicles elicited flight reaction from about
50 percent of the individuals from four different groups ranging in size
from two to eight deer. One vehicle elicited flight reaction from 100
percent of a group of two individuals.
Previous to these observations it
was hypothesized that deer at proximal distances of 10 to 20 m from moving
traffic were frequently caused to leave the area. Other observations
indicated that when vehicles stopped at or when bicyclists traveled by the
underpass, most deer elicited immediate and intense flight reaction.
The
behavior modes of deer exits (leaving the underpass to the south) were 71
(n~25), 23 (n~8), and 6 (n~2) percent, trotting, walking, and bounding,
respectively.
The predominant exit behavior of trotting sU'Dports the
reported reluctance (Reed et a1. 1975) of deer to use a structure of this
size and character.

LITERATURE
Hediger, H.

1950.

CITED

Wild animals in captivity.

Butterworth,

London. p. 19.

Reed, Dale F. 1971. Deer underpass evaluation.
'D. 341-351.
In Game
Research Report, July Part 3. Colo. Game, Fish and Parks Div. Fede raI
Aid.
Reed, Dale F., T.N. Woodard, and T.M. Pojar.
1975. Behavioral response
of mule deer to a highway underpass.
J. l-1ildl.Manage. 39 (2) :

Wildlife Researcher

......

�-309-

Table 1. Highway underpasses, height and width dimensions, number of deer
passages, tunnel effect, and the deer activity adjacent to the underpasses.
Only the last two structures were intended for regular deer use.

No. deer
passages

Tunnel
effect.!.!

Adjacent
activity

0

0.3

None

l4X14

6

1.4

Light

Eagle 1-7n No. 3

8X8

2

0.3

T-1oderate

Eagle 1-70 No. 4

l4X14

0

0.9

Light-moderate
movements parallel
to 2.44-m fence

Chaffee Gulch
(N. Ridgway)

12XlO+2

6.9

Light, no fencing

Arch deer
(w. Durango)

10X20

46

2.1

Light

Vail deer (1-70)

10XlO

345.1 ~I

1.0

Concentrated, highly
motivated migration

Underpass

height X width

Eagle 1-70 No. 1

8X8

Eagle 1-70 No. 2

]j

e oxl 0) '!:..I 14 ]/
2

An expression

of openness

height X width

calculated

deer

as follows:

(or open end surface area)
length

'!:..I
Three-span

bridge with fill-slop under spans 1 and 3.

11 Deer passages

were monitored from 8 Oct 73 to 6 Apr 74. nocumented deer use
during this period was sufficient data for the Division of Hi~hways to justifv
replacing the sub-standard bridge with an equallv open span-bridge.

41

-

Seasonal mean resulting

from a 4-year study.

�I

o
......•
1:""1
I

Fig. 1. The north end of Eagle 1-70 underpass No. 3 received deer pressure for over a 1 1/2-month
period. Two passages occurred through the structure to the south during this period (Photo by
D.F. Reed).

�-311July, 1975
JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No. W-38-R-29 and 30
Work Plan No.
Job Title

15

Job No.

7

---------------------------

Effects of Highway Lighting on Number of Deer Killed by Vehicles

Period Covered:
Personnel:

Deer-Elk Investigations

April 1, 1974 through March 31, 1975.

Claudia A. Doose, James D. Fleming, Larry L. Green, Sharon L.
McDonnell, Allen F. Whittaker, John W. Seidel, Dale F. Reed,
and Thomas N. Woodard

ABSTRACT
The effect of highway lighting on the rate of deer-vehicle accidents was
evaluated. Estimated deer crossings per kill decreased by 2.0 percent
with the lights on compared to lights off. There was no significant
difference in the crossings per kill ratios (X2 = 0.003, P&gt;0.90).
The
amount of light available at 12 known kill locations during lights on
nights ranged from less than 0.01 to 1.41 foot candles. There was a
highly significant reduction in mean vehicle speeds (P&lt;O.OOl) when a deer
simulation was placed on the shoulder of the highway with lights on.

��-313-

EF~ECTS O~ HIGHWAY LIGHTING
ON NUMBER O~ DEER KILLED BY VEHICLES
Thomas N. \-Toodard

p.

S. OBJECTIVE

Determine if highway lighting affects the rate of deer-vehicle
on a portion of Highway 82.

accidents

SEGMENT OBJECTIVES
1.

Record the number of deer killed by vehicles at each end of, and
within a 1.1 km lighted portion of Colorado 82.

2.

Estimate deer densities
by the lighting.

3.

Compare pre-treatment
and count data.

4.

Gather additional information which may be needed to understand why
deer-vehicle accident rates are or are not affected by lighting.

adjacent to the portion of highway

affected

kill and count data to post-treatment

kill

METHODS &amp;~D MATERIALS
Methods and materials

have been described by Woodard

(1974).

The effect of lighting was evaluated from January 2, 1975 to March 31,
1975, during this segment.
Data collected during segment 28 and from April
1, 1975 to April 9, 1975, are also included in this report.
A deer simulation (transverse section of full taxidermy mount) was placed
in the emergency lane of the lighted portion of the highway with the lights
on between 1900 and 2212 DST on March 17, 1975. The simulation was located
24.1 m beyond the first and 20.1 m before the second of two magnetic loops
which activate and deactivate a vehicle speed recorder (Newmeyer Model NH-l).
Motorist response to the simulation was measured by comnaring the vehicle
speeds collected on March 17, 1975, (treatment) to vehicle speeds collected
during the lights on night of March 3, 1975, (control) and by making visual
observations.
Only speeds recorded from 2018 to 2042 and 2118 to 2149 were
utilized in tabulations because of probable bias from motorists interference
during the other time periods.
The amount of light (foot-candles) present at each kill location during lights
on nights was measured with a light meter (General Electric Street Lamp Meter
SL480A).

�-314-

RESULTS AND DISCUSSION
Estimated crossings per kill was only 2.0 percent lower with the lights
on compared to lights off (Table 1). There was no significant difference
in the ratios (x2 = 0.003, P&gt;O.90).
The amount of light available at 12 known kill locations during lights
on nights ranged from less than 0.01 to 1.41 foot-candles.
Available
light at seven of these kill locations was less than 0.01 foot-candles.
Mean vehicle speeds decreased by 8.61 mph when the deer simulation was
present.
The difference was highly significant (P&lt;O.OOl).
Brake lights
were observed on 50.6 percent of the vehicles approaching the simulation
(n=85) from 2118 to 2149 DST.

Table 1. Estimated deer crossings and total kill in the Highwav
lignting study area during 1974 and 1975.
Lights o~~
Lights ON
Estimated

Crossings

82

1378

Total Kill

17

16

Crossings/Kill

87.9

86.1

LITERATURE

CITED

Woodard, Thomas N. 1974. Effects of highway lighting on number of deer
killed by vehicles. Colo. Div. l.Ji1dl.Game Res. Div. ~ed. Aid Proj.
W-38-R-28. Game Res. Rep., July, Part 2. Pp. 307-312.

Prepared by

~r?~.".

l/ ~.

/ /

j,~

(.~;i..~·l/

Thomas N. Woodard
.
Assistant ~1ildlife Researcher

�-315July, 1975
JOB PROGRESS

State of

REPORT

COLORADO
--------~~~~----------W-38-R-29

Project No.

Job Title

Deer-Elk

Investigations

15
10
Job No.
An Evaluation of 8-Foot (2.4'4---m')~F~e-n-c-e~L-e-n-g~tTh--------------Required to Prevent Movements on or Across High Speed Highways

Period Covered:
Personnel:

and 30

April 1, 1974 through March 31, 1975.

Thomas N. Woodard, Sharon L. McDonnell, William R. Heicher,
David C. Hoart, Larry L. Green, Claudia A. Doose, James D.
Fleming, and Dale F. Reed.

Annual or seasonal deer kill caused by vehicles was reduced by 79, 77, 84,
59, and 76 percent along 8-foot (2.44-m) fence lengths of 1.8, 1.8, 3.6,
3.6, and 7.7 kilometers, respectively.
Ninety-five, 17, and 34 deer escaped
the highway rights-of-way adjacent to the two 2.25 and the 4.8 mile long
8-foot fences through 17 one-way deer gates, respectively.
Does (N=13) and
bucks (N=5) moved laterally or parallel to the 8-foot fence for mean distances
of 0.30 and 0.48 miles, respectively.

��-317-

AN EVALUATION OF 8-FOOT (2.44-M)
FENCE LENGTH REQUIRED TO PREVENT DEER
MOVEMENTS ON OR ACROSS HIGH SPEED HIGHWAYS
Dale F. Reed

P. S. OBJECTIVE
To evaluate length of 8-foot (2.44-m) fence in relation
on or across high speed highways.

to deer movement

SEGMENT OBJECTIVE
1.

Install 8-foot fence along appropriate
mined by Job 1a.

2.

Evaluate the effectiveness
affecting deer movements.

sections of highway as deter-

of various lengths of 8-foot fence in

METHODS AND MATERIALS
Methods and materials
(1974a).

have been described

DESCRIPTION
Highway

by Pojar

OF AREA

82 - Diamond

The Highway 82 study area has been described

Interstate

(1972) and Reed et al.

S
by Pojar

(1972).

70 - Avon. Edwards. Vail and Eagle

The Interstate 70 - Avon and Edwards study areas have been described by
Woodard (1973). the Vail study area by Reed (1971) and Reed et a1. (1974a).
and the Eagle study area by Reed (1974).

Highway

82 - Carbondale

The Carb-82 study area consists of a segment of Highway 82 from about
1.3 kilometers southeast of the junction with Highway 133 to about 0.2
kilometers up Crystal Springs road. An 8-foot (2.44-m) fence was completed
along the north side of the highway rights-of~ay
October 17, 1974. Alfalfa
fields are prevalent north of the highway with big sagebrush occurring on an
abbreviated south facing slope behind the fence.

�-318-

RESULTS AND DISCUSSION
Highway 82 - Diamond S
Deer concentrate in the crested wheatgrass fields northeast of the
highway in late winter and early spring (Table 1). The mean number of
deer crossings during March - May periods in 1971 before the 8-foot
fence was installed was 11.7 (N=32, where N=number of 24 hour periods)
per day. After the fence was installed the means were 2.1 (N=38) in
1972, 5.5 (N=34) in 1973, and 10.3 (N=38) in 1974. This represents
an 82, 53, and 12 percent reduction in crossings in 1972, 1973, and
1974, respectively.
The mean deer kill per year before the 8-foot fence was installed
(October, 1968 - September, 1971) was 10.0. The mean deer kill per
year after fence installation (October, 1971 - September, 1974) was
2.33; a mean reduction of 77 percent.

Table 1. The March-May mean number of deer observed on spotlight
counts between quarter-mile section markers 25 to 30 for 1968 through
1973 on Highway 82 (n=number of counts).

Month

1968

1969

Mean Count
1970
1971

1972

1973

1974

March

134.8
(n=4)

151.2
(n=4)

104.5
(n=4)=

66.8
(n=4)

102.2
(n=4)

137.4
(n=S)

143.5
(n=4)

73.0
(n=4)

34.0
(n::05)

56.0
(n=5)

51.4
(n=5)

4.5
(n=4)

47.0
(n=4)

52.3
(n=3)

6.0

(n=5)

0.0
(n=L)

1.5
(n=2)

0.0
(n=2)

66.2
(n=13)

77.5
(n=10)

63.7

47.6

53.4

(n=Ll.)

(n=L'l )

(n=B)

97.2
(n=9)

104.4
(n=7)

April

May

MEAN
TOTAL

Interstate

70 - Avon

The mean number of deer observed on spotlight counts during the August December periods was 15.5 (n=17) and 13.4 (n=21) in 1971 and 1072,
respectively, and 10.8 (n=ll) in lq73 for October - December and 7.3 (n=7)
in 1974 for November - December. Ninety-five deer made passages to the
north through six one-way gates (Reed et a1. 1974b) located in the 8-foot fence
during this segment.
Twenty-eight deer were killed from October 5, 1971
to October 5, 1972, before the fence was installed.
Nine were killed from
October 5, 1972 to October 5, 1974, after the fence was installed; a
reduction of 84 percent.
Two deer crossed the deer guard in the Avon 8-foot fence. Deer use and
response to other prototype test guards will be analyzed and reported in
future segments.

�-319-

Interstate

70 - Edwards

Twenty-seven deer were killed on this segment of highway before the
fence was installed from October 5, 1971 to October 5,1972.
Twentytwo (11 per year) were killed after fence installation from October 5,
1972 to October 5, 1974; a reduction of 59 percent.
Seventeen passages
occurred through four of seven one-way deer gates located in the 8-foot
fence.
Interstate
No additional
length.

data were collected

70 - Vail
in this study area in regard to fence

Interstate

70 - Eagle

The mean number of deer observed on spotlight counts during OctoberMarch 1972-73 was 80.5 (n=25). One hundred sixty-seven deer were
killed from October 5, 1972 to October 5, 1973, before the fence was
installed.
Forty were killed from October 5, 1973 to October 5, 1974,
after the fence was installed; a reduction of 76 percent.
An estimated
90-100 deer, including several collared animals, came across the highway from the south and either 1) passed through the one-way deer gates
or &amp;1 opening in the fence, 2) went around the ends of the fence, 3)
went back to the south to or across the Eagle River, or 4) were killed
on the highway.
Thirty-four deer made passages to the north through
seven of ten one-way gates located in the 8-foot fence.
According to 69 observations of collared deer north of the 8-foot fence,
does (N=13) and bucks (N=5) have moved laterally or parallel to the fence
for mean distances of 0.30 and 0.48 miles, respectively.

Highway 82 - Carbondale
The mean number of deer observed on spotlight counts during January April periods of 1968 through 1973 was 15.7 (n=99). Seventy deer were
killed from January, 1968 to April, 1973 (14 deer per year), before the
fence was installed.
Three were killed from October 17, 1974 to March
31, 1975, after the 8-foot fence was installed; a reduction of 79 percent.

LITERATURE

CITED

Pojar, Thomas M. 1972. An evaluation of deer-proof fence length required
to prevent deer movements on or across high speed highways.
Colo.
Div. Wildl., Game Res. Div. Fed. Aid Proj. W-38-R-26, Game Res. Rep.,
July, Part 3. pp. 305-310.

�-320-

Reed, Dale '1&lt;'.
1971. Deer tmderpass evaluation. Colo. Div. 'tU1d1. Game
Res. Div. ~ed. Aid Proj. W-38-R-25. Game Res. Rep. July, Part 3.
pp , 341-351.
Reed, D.~., T.M. Pojar, and T.N. Woodard. 1974a. Mule deer responses
to deer guards. J. Range Manage. 27(2):111-113.
Reed, D.F., T.M. Pojar, and T.N. Woodard. 1974b. Use of one-way
by mule deer. J. Wildl. Manage. 38(1):9-15.

gates

Reed, D.'I&lt;'.,1974. An evaluation of 8-foot fence length required to
prevent deer movements on or across high speed highwavs. Colo.
Div. Wildl. Game Res. Div. Fed. Aid Proj. W-38-R-28. Game Res.
Rep., July, pp. 313-320.
Woodard, Thomas N. 1973. An evaluation of deer-proof fence length
required to prevent deer movements on or across high speed
highways. Colo. Div. Wildl. Game Res. Div. Fed. Aid Proj.
W-38-R-27. Game Res. Rep., July, Part 3. pp.

Prepared

~/.~
by . /f~'-';1
I
Dale F. Ree
Wildlife Researcher

JZ(L~ .

.J

)
(

/

�July 1975

-321-

JOB FINAL REPORT

State of

C_o~L~O~RAD~O
W-38-R-29

Project No.

Deer-Elk Investigations

11
Job No.
Effects of a Simulated 8-Foot Fence Angle
in Diverting Deer from Their Established Direction of Movement

Period Covered:
Personnel:

and 30

15

Work Plan No.
Job Title:

_

April 1, 1972 through March 31, 1975

Thomas N: Woodard, Thomas M. Pojar, and Dale F. Reed

ABSTRACT

Observations were made on time and nature of mule deer (Odocoileus hemionus)
responses to five simulated fence angles in 1972 and three simulated fence
angles in 1973. Mean time needed for deer to successfully travel the 27.42-m
length of 900 fence angle was significantly greater (P &lt;0. 001) than the
mean control time (00 angle). The number of deer that failed to move through
the zone during the 900 fence angle measurements was not significantly
greater (X2 = 0.202, P&gt;0.50)
than the number during control. No additional
data were collected during the last segment.

��-323-

EFFECTS OF A SIMULATED 8-FOOT FENCE ANGLE
IN DIVERTING DEER FROM THEIR ESTABLISHED DIRECTION OF MOVEMENT
Thomas N. Woodard

P. S. OBJECTIVE
Evaluate the effectiveness of three angles of a simulated 8-foot fence to
divert deer from their established direction of movement.

SEGMENT OBJECTIVES
1.

Install a movable 8-foot section of 4-inch square netting adjacent to
and across heavily traveled deer trails and concentration points.

2.

Test the effectiveness

of three angles under field conditions.

METHODS AND MATERIALS
An 8-foot (2.44 m) high simulated fence made of 4-inch netting (350 lb test)
was easily installed for a length of 90 feet (27.42 m) at one of three
angles (450, 600, and 900; plus converse angles of 1200 and 1350) in relation
to a permanent 8-foot wire deer fence (Fig. 1). The permanent wire fence had
been placed across a deer migration route to divert migrating deer. There
was a heavily traveled deer trail along this fence.
The time for a deer to travel the 90-foot length of permanent 8-foot deer
fence (hereafter referred to as the "control"), and the same length of
simulated fence at each angle (variables 450,600, and 900, plus converse
angles of 1200 and 1350) was measured by making early morning observations
and recording with a stop watch the time each deer spent between two clearly
marked points. The amount of time was recorded to the nearest second for
each animal. If a group approached, the first animal moving into the marked
area (time zone) was chosen for timing. When two or more deer moved into
the zone simultaneously either the "lead" animal or a tagged animal was used.
Observations were made during early morning hours for 33 days from June 6 to
July 19, 1972. The sequence of the simulated fence angles tested was: 00
(control), 450, 600, and 900• This sequence was repeated throughout the
above period. The simulated fence was installed at the angle specified by
the above sequence during the evening before the day's observation.
Factors such as the wind, light, natural cover, and camouflage clothing
However, if the observer
were utilized to advantage during the observations.
was detected by the animals, the day's observations were terminated.

�-324-

Fig. 1. A permanent 8-foot wire deer fence is shown in the foreground.
The
simulated fence (netting) is shown installed at the 600 angle (right center)
from the permanent fence. The wires to the left and the far right are the
900 and 450 angles, respectively.
(Photo by D. F. Reed)

�-325-

Observations were made during 21 days in 1973 from June 12 to July 9. The
600 angle was omitted in 1973. No observations were made during 1974.
DESCRIPTION OF AREA
The observations on deer response to the 8-foot fence angle simulation
were made near Vail, Colorado, adjacent to the north 8-foot fence of
the Vail underpass - fencing complex (Reed 1971). The observation site
was located on the edge of a conifer community with interspersed aspen
(Populus tremuloides) and big sagebrush (Artemisia tridentata). Sunrise occurred behind the observer's location and normal westerlv winds
or breezes placed the location On the lee side of most deer during the
observations.
A dirt road paralleled the 8-foot fence, and curved awav from it near
the point where the simulated fence was attached to the wire fence.
There was a gradual slope located adjacent to the road opposite the
observation point. The simulated fence angle of 900 was installed across
the road and up this slope. The fence angle of 600 was tangent to the
outer shoulder of the bend in the road. The fence angle of 450 was installed down an incline to the east and 20 to 30 feet from the road and
slope at the closest point.

RESULTS AND DISCUSSION
Time was measured on deer in the simulated fence angle area on 55 occasions
(Table 1). Twenty-nine
times were recorded during control measurements. 0
.
Mean time needed for deer to successfullv travel the 27.42-m length of 90
fence angle was significantlv greater (P&lt;O.OOl) than mean control time.
The number of deer that failed to move through the zone during the 900 fence
angle measurements was not significantlv greater (X2 = 0.202, P&gt;O.SO) than
the number during control; thus the simulated fence angle did not preclude
deer from their apparent direction of movement.
Statistical tests were not applied to the other angle variables because of
insufficient sample size.
LITERATURE CITED
Reed, Dale F. 1971. Deer underpass evaluation. Colo. niv. r.ame, Fish and
Parks. Game Res. Div. Fed. Aid P'roj , l-l-38-R-2S.r.ame Res. Ren ;, Julv,
Part 3. p . 344.

---I

Prepared by

- /1"." "\ I,,"
-I;.

I.

,(

•..

Thomas N. Woodard
Assistant Wildlife Researcher

�-326-

Table 1. The mean times (seconds) of successes or failures of deer moving
through the zone with either the simulated fence angle removed (control)
or installed (variables: 450,600, or 90°). Successful animals moved along
the permanent 8-foot (2.44-m) fence or along the simulated fence. ~ailure
animals moved into the zone, turned around, and left in the direction that
they had come.

Angles

Successful
75.8 (n=19) }j

85.6 (n=10)

66

450

168.2 (n=3)

52.0 (n=Z)

60

0

900
0

120

0

173.6 (n=S)

]j

1350 J:../

2/

Percent Successful

00

600

1./

~ailure

78.3 (n=6)

0

121.3 (n=4)

73.0 (n=L)

117.0 (n=L)

Number of observations

60

50

(n ) •

°

When deer approached the simulated fence angles of 60 o and 45 from the
opposite direction (east), they encountered converse angles of 120° and 135°.

�-327JOB PROGRESS

July, 1975
REPORT

State of

COLORADO

Project No.

W-38-R-29

Deer-Elk

Investigations

Work Plan No.

16

Job No.

1

Job Title

Piceance Deer Study - Population

Period Covered:

April 1, 1974 through March 31, 1975

Personnel:

Distribution

R. M. Bartmann, J. J. Klein, Jr., M. Alan, D. Smeltzer,
A. C. Koonce.

and

ABSTRACT
Deer trapping and banding work was accomplished from December 16, 1974 through
February 28, 1975. Three hundred one deer were both eartagged and neckbanded
and another eight eartagged only. Banding was completed in Area 11. This
leaves about 250 deer to be banded in three areas next winter.
The composition
of the total catch was 76 male fawns, 61 female fawns, 15 adult males, and 157
adult females.
Sightings of 153 banded deer were reported along with 51 recoveries.

��-329-

PICEANCE DEER STUDY - POPULATION

DISTRIBUTION

Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.

SEGMENT OBJECTIVES
1.

To delineate deer sub-population
the Piceance winter range.
a.

Define deer sub-population

boundaries

and concentration

areas on

boundaries.

METHODS AND MATERIALS
See Bartmann

(1972).

RESULTS AND DISCUSSION
Deer Trapping
Deer trapping and banding work was started December 16, 1974 and continued
through February 28, 1975. Trapping was initiated on Oak Ridge (Area 10)
east of Meeker, but deep snow forced elk into the trapping areas and operations were moved, after marking 15 animals, to Strawberry Creek (Area 11).
Snowmobiles were required to set and check traps into early February, after
which pickups could be used. The winter was considerably milder than the
previous two, particularly in terms of snow cover, and survival of banded
animals should be good.
Only 301 deer were both ear tagged and neckbanded and another 8 eartagged
only, which is 69 deer fewer than the previous low of 378 deer marked in
1972 (Table 1). This leaves about 250 deer to be neckbanded in three areas
next winter.
One hundred fifty-three sightings of banded deer were recorded during Segment 29 and 51 deer were reported killed or found dead (Tables 2 and 3).
The number of band recoveries was reduced by the antlered-only season, and
deer movement information from both sightings and recoveries limited by
the late season (October 26 - November 5) as many deer were already back
on the winter range.

�-330-

LITEFATURE

CITED

Bartmann, R. M. 1972. Piceance deer study - population distribution.
p. 315-337.
In Game Research Report. Colo. Div. of Wildlife.
Denver.
3(Part 3):253-377.

Prepared

bY~?g;2~
Richard M. Bartman
Wildlife Researcher

..&lt;...

�Table 1.
1974-75.
Date
(1974)

Record of deer trapped and marked on the winter range in Game Management Units 11 and 23, winter

Age

8ex

Eartag
No.

Neckband
No.

Twp.

Location
Range

Remarks

Sec.

Area 10 - Yellow Neckband
12-19
12-19
12-19
12-19
12-21
12-21
12-21
12-21·
12-22
12-22
12-22
12-22
12-22
12-23
12-23

Male
Male
Female
Male
Female
Female
Female
Female
Male
Female
Female
Male
Female
Female
Female

Summary:

Fawn
Fawn
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Fawn

5 Male Fawns;

L-2780
L-2781
L-2783
L-2785
L-2786
L-2784
L-2790
L-279l
L-2792
L-2793
L-2794
L-2795
L-2796
L-2797
L-2798
2 Female Fawns;

32
33
1
34
2
None
3
4
35
7
5
39
8
6
13

IS
1S
18
1S
18
18
18
18
18
1S
18
18
18
1S
18

o Mature Males;

93W
93W
93W
93W
93W
93W
92W
93W
92W
92W
92W
92W
93W
92W
92W

11
11
2
2
6
11
14
24
15
15
19
19
24
19
19

Found dead 3-75, possible road-kill.

Lower part of left hind leg missing.

I

w
w
•...
I

8 Mature Females

Area 11 - Red and Blue Block Neckband
12-30
12-31
1-1-75
1-1
1-1
1-1
1-2
1-2
1-2
1-2
1-2

Female
Female
Male
Female
Male
Female
Male
Male
Female
Female
Female

Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Mature
Mature
Fawn

L-2799
L-2800
L-2801
L-2802
L-2803
L-2804
L-2805
L-2806
L-2807
L-2808
L-2809

1
14
90
11,
94
None
110
102
12
13
15

2N
2N
2N
2N
2N
2N
1N
1N
2N
2N
2N

94W
95W
95W
95W
94W
94W
95W
95W
94W
95W
95W

17
1
25
25
17
17
13
13
17
1
1

Possible broken neck.

----------------------------------------------------------------------------------------------------------------

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 11 and 23, winter
1974-75 (continued).
Date
(1975)

Sex

Age

Eartag
No.

Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Male
Female
Female
Male
Male
Female
Female
Female
Male
Female
Male
Female
Male
Male
Male
Female
Female
Female
Female
Female
Female
Male

Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Fawn
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn

L-2810
L-2811
L-2812
L-2813
L-2814
L-2815
L-2816
L-2817
L-2818
L-2819
L-2820
L-2821
L-2822
L-2823
L-2824
L-2825
L-2826
L-2827
L-2828
L-2829
L-2830
L-2831
L-2832
L-2833
L-2834
L-2835
L-2836
L-2837
L-2838
L-2839
L-2840
L-2841
L-2842

Neckband
No.

Location
Twp.
Range
Sec.

2
5
3
8
104
7
6
10
17
18
19
116
4
20
95
97
None
None
22
88
23
108
24
109
106
111
30
25
26
27
28
29
112

IN
2N
2N
3N
3N
IN
IN
IN
2N
2N
2N
2N
2N
2N
2N
2N
2N
IN
IN
IN
IN
IN
2N
2N
2N
2N
2N
2N
3N
3N
3N
IN
IN

Remarks

-1-3
1-3
1-3
1-3
1-3
1-4
1-4
1-4
1-4
1-4
1-4
1-4
1-4
1-4
1-4
1-4
1-4
1-5
1-5
1-5
1-5
1-5
1-5
1-5
1-5
1-5
1-5
1-5
1-5
1-5
1-5
1-6
1-6

.

95W
94W
94w
95W
95W
95W
95W
95W
95W
95W
95W
94W
94w
95W
95W
95W
94w
94W
94W
95W
95W
95W
95W
95W
95W
95W
95W
94W
95W
95W
95W
94W
94W

12
6
6
26
26
13
12
12
25
25
26
17
17
1
1
1
6
17
17
13
13
12
25
25
25
1
1
6
26
26
26
8
8

I

\.oJ
\.oJ

N
I

Poor shape.

-----------------------------------------------------------------------------------------------------------------

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 11 and 23, winter
1974-75 (continued).

Sex

Age

Eartag
No.

Female
Female
Male
Female
Male
Female
Female
Male
Female
Female
Male
Male
Male
Male
Female
Female
Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Male

Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Fawn
Fawn

L-2843
L-2844
L-2845
L-2846
L-2847
L-2848
L-2849
L-2850
L-2851
L-2852
L-2853
L-2854
L-2855
L-2857
L-2856
L-2858
L-2859
L-2860
L-2861
L-2862
L-2863
L-2864
L-2865
L-2866
L-2867
L-2875
L-2869
L-2868
L-2870
L-2871
L-2872
L-2873
L-2874

Date
(1975)
1-6
1-6
1-7
1-7
1-7
1-7
1-8
1-8
1-8
1-8
1-9
1-9
1-9
1-9
1-10
1-10
1-10
1-10
1-10
1-11
1-11
1-11
1-12
1-12
1-13
1-13
1-13
1-13
1-13
1-13
1-14
1-14
1-14

Neckband
No.

Twp.

39
34
113
35
114
40
31
85
32
33
None
103
100
101
36
37
87
38
92
46
47
48
42
41
43
21
44 .
117
45
53
54
55
96

2N
2N
1N
1N
3N
3N
1N
3N
3N
3N
1N
1N
2N
2N
IN
IN
IN
IN
2N
IN
2N
2N
2N
2N
IN
2N
IN
3N
IN
IN
2N
2N
2N

Location
Sec.
Range
95W
95W
94W
95W
95W
95W
94W
95W
95W
95W
94W
95W
95W
95W
94W
95W
94W
95W
94W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W
95W

Remarks

26
1
8
12
26
26
17
26
26
26
8
12
25
25
17
12
17
12
17
13
25
26
26
26
12
25
16
26
29
29
26
26
26

----------------------------------------------------------------------------------------------------------------

I

w
w
w
I

�Table 1.

Record of deer trapped and marked on the winter range in Game Management Units 11 and 23, winter
1974-75 (continued).

Date
(1975)

Sex

Age

Eartag
No.

Neckband
No.

Twp.

1-14
1-14
1-14
1-15
1-15
1-15
1-15
1-15
1-15
1-15
1-15
1-16
1-16
1-16
1-16
1-16
1-16
1-16
1-17
1-17
1-17
1-17
1-17
1-17
1-17
1-18
1-18
1-18
1-18
1-18
1-18
1-18
1-19
1-19

Male
Female
Female
Female
Female
Female
Male
Female
Male
Female
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Male
Male
Female
Male
Female
Male
Female
Female
Male

Fawn
Mature
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn

L-2876
L-2877
L-2878
L-2879
L-2880
L-2881
L-2883
L-2884
L-2885
L-2886
L-2887
L-2888
L-2889
L-2890
L-2901
L-2891
L-2892
L-2893
L-2894
L-2895
L-2896
L-2897
L-2898
L-2899
L-2900
L-2902
L-2903
L-2904
L-2905
L-2906
L-2907
L-2908
L-2909
L-2910

93
56
57
49
50
58
122
52
123
51
59
105
60
62
71
16
63
64
66
65
67
68
69
70
107
115
86 .
79
120
72
121
73
74
118

2N
3N
IN
IN
2N
2N
IN
IN
IN
IN
1N
1N
IN
2N
1N
IN
1N
1N
1N
1N
1N
IN
1N
1N
1N
IN
1N
1N
1N
IN
1N
1N
2N
2N

Location
Range
Sec.
94w
95W
95W
94W
95W
94W
95W
95W
95W
95W
95W
95W
95W
94W
95W
95W
96W
96W
95W
95W
95W
95W
95W
95W
95W
96W
95W
95W
95W
95W
95W
95W
96W
96W

17
26
29
17
25
17
16
17
29
29
29
12
12
17
29
16
12
12
17
17
16
16
29
29
29
12
17
17
17
16
16
29
27
23

Remarks

I

w
w
.po.
I

May be neckband #76.

-----------------------------------------------------------------------------------------------------------------

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 11 and 23, winter
1974-75 (continued).
Date
(1975)

Sex

Age

Eartag
No.

Neckband
No.

Twp.

1-19
1-19
1-19
1-24
1-24
1-24
1-24
1-24
1-24
1-25
1-25
1-25
1-26
1-27
1-30
1-30
1-31
2-1
2-1
2-4
2-5
2-5
2-6
2-7
2-7
2-7
2-7
2-8
2-9
2-9
2-10

Male
Female
Male
Female
Female
Female
Female
Female
Male
Female
Female
Female
Male
Female
Male
Female
Female
Male
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Male
Male
Female

Fawn
Fawn
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature

L-2911
L-2912
L-2913
L-2915
L-2916
L-2917
L-2918
L-2919
L-2921
L-2922
L-2923
L-2924
L-2925
L-2926
L-2927
L-2928
L-2929
L-2930
L-2931
L-2938
L-2946
L-2947
L-2957
L-2962
L-2963
L-2964
L-2965
L-2973
L-2985
L-2986
L-2987

119
75
124
82
77
83
76
78
126
80
84
151
125
152
133
161
168
127
159
153
155
167
158
134
157
160 '
169
154
128
132
165

1N
IN
IN
2N
2N
2N
2N
IN
IN
2N
2N
2N
IN
IN
IN
IN
IN
IN
1N
IN
1N
1N
2N
2N
2N
2N
2N
2N
2N
1N
IN

Location
Range
Sec.

95W
95W
95W
96W
95W
95W
95W
96W
96W
96W
95W
95W
96W
95W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

17
16
16
21
30
19
19
4
1
27
30
19
1
6
14
12
4
4
4
9
4
4
32
32
32
32
32
32
32
9
9

Remarks

May be neckband #79.

I

w
w
VI
I

May be neckband #190.

---------------------------------------------------------------------------------------------------------------------

�Table l. Record of deer trapped and marked on the winter range in Game Management Units 11 and 23, winter
1974-75 (continued).
Date
(1975)

Sex

Age

Eartag
No.

Neckband
No.

Twp.

Location
Range

Sec.

Female
Male
Male
Female
Male
Female
Female
Female
Female
Female
Female
Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Male
Female
Male
Female
Female
Female
Female
Male

Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Mature
Fawn
Fawn
Fawn

L-2989
L-2997
L-2998
L-3005
L-3006
L-3007
L-3009
L-3012
L-3013
L-3015
L-3018
L-3019
L-3020
L-3026
L-3027
L-3028
L-3029
L-3032
L-3033
L-3034
L-3035
L-3036
L-3038
L-3037
L-3043
L-3044
L-3045
L-3046
L-3047
L-3048
L-3051
L-3052
L-3053

156
131
129
162
142
182
163
164
166
170
171
140
172
"139
175
176
177
178
179
173
174
130
180
181
183
141
184
145
185
186
187
188
136

2N
2N
2N
2N
IN
IN
IN
2N
2N
IN
IN
IN
2N
IN
IN
IN
IN
IN
IN
IN
2N
2N
IN
IN
IN
IN
IN
IN
2N
IN
IN
IN
IN

96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

32
32
32
33
4
4
4
21
21
4
15
15
21
14
4
4
4
23
23
14
32
28
4
4
23
14
14
14
28
4
23
14
12

Remarks

-2-11
2-12
2-12
2-13
2-13
2-13
2-14
2-15
2-15
2-15
2-16
2-16
2-16
2-18
2-18
2-18
2-18
2-19
2-19
2-19
2-19
2-19
2-19
2-19
2-20
2-20
2-20
2-20
2-20
2-20
2-21
2-21
2-21

-------------------------------------------------------------------------------------------------------------------

I
Vol
Vol

0\
I

�Table

J.

1974-75

Record

of deer

trapped and marked on the winter range in Game Management

UnI ts 11 and 2:3, w Ln t c r

(continued) .

Date
(1975)

Sex

Age

2-21
2-21
2-21
2-21
2-21
2-22
2-22
2-22
2-22
2-22
2-22
2-22
2-22
2-22
2-23
2-23
2-23
2-24
2-24
2-24
2-24
2-25
2-25
2-25

Hale
Female
Female
Male
Female
Female
Male
Hale
Female
Female
Female
Female
Hale
Female
Female
Female
Hale
Female
Female
Female
Male
Female
Female
Hale

Fawn
Hature
Hature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Hature
Mature
Hature
Mature
Hature
Mature
Fawn
Mature
Fawn
Hature
Fawn
Mature
Mature
Fawn

Summary:

Eartag
No.

Neckband
No.

Twp.

Location
Range

Sec.

L-3054
L-3055
L-3056
L-3057
L-3058
L-3060
L-306l
L-3062
L-3063
L-3064
L-3065
L-3066
L-3067
L-3068
L-3070
L-3071
L-3073
L-3075
L-3076

137
189
138
144
135
143
198
199
146
147
148
149
195
150
192
193
197
196
200
194
201
202
None
None

IN
IN
2N
2N
IN
IN
IN
IN
IN
IN
2N
2N
2N
IN
IN
2N
2N
IN
IN
IN
IN
IN
2N
IN

96W
96W
96W
96W
96w
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

15
15
28
28
9
23
14
14
12
12
32
27
28
9
23
28
27
12
12
4
4
23
27
4

L-30n

L-3078
L-3085
L-3087
L-3088

50 Hale Fawns;

44 Female Fawns;

11 Hature Males;

Remarks

94 Mature Females

Area 12 - White with Red Stripe Neckband
2-2
2-2
2-3
2-3

Female
Hale
Female
Female

Mature
Hature
Mature
Hature

L-2932
L-2933
L-2934
L-2935

10
1
11
12

2N
2N
2N
2N

96W
96W
96W
97W

30
19
19
4

---------------------------------------------------------------------------------------------------------------------

I
\.oJ
\.oJ

-.J
I

�Table 1. Record of deer trapped and marked on the winter range in Game Management Units 11 and 23, winter
1974-75 (continued).
Date
(1975)

Sex

Age

Eartag
No.

Neckband
No.

2-3
2-3
2-4
2-4
2-4
2-4
2-4
2-4
2-4
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-5
2-6
2-6
2-6
2-7
2-7
2-7
2-7
2-7
2-7
2-7
2-8
2-8
2-8
2-8
2-8

Female
Female
Female
Female
Male
Female
Male
Female
Male
Male
Male
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Male
Female
Female
Female
Male
Female
Male
Female
Male
Female

Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature

L-2936
L-2937
L-2939
L-2940
L-2941
L-2942
L-2943
L-2944
L-2945
L-2948
L-2949
L-2950
L-2951
L-2952
L-2953
L-2954
L-2955
L-2956
L-2958
L-2959
L-296l
L-2966
L-2967
L-2968
L-2969
L-2970
L-2971
L-2972
L-2974
L-2975
L-2977
L-2978
L-2979

17
18
13
14
2
15
4
19
3
8
7
47
16
105
'106
20
21
107
108
109
22
5

111
51
119
27
118
48
116
50
36
52
112

Twp.

Location
Range

Sec.

2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N

98W
98W
96W
96W
96W
96W
97W
97W
97W
96W
96W
96W
96W
97W
98W
98W
97W
97W
96W
97W
97W
97W
97W
97W
97W
98W
98W
97W
97W
98W
97W
97W
97W

1
1
30
30
19
19
4
6
6
30
30
30
30
6
1
1
17
17
19
36
6
36
36
6
6
1
1
17
4
1
3
3
36

Remarks

--------------------------------------------------------------------------------------------------------------------

•

I

w
w

00
I

�Table l.
197/;-75

Record of deer trapped and marked
(continued) .

Date

on the winter range in Game Management

(1975)

Sex

Age

Eartag
No.

Neckband
No.

Twp.

2-8
2-8
2-9
2-9
2-9
2-9
2-9
2-10
2-11
2-11
2-11
2-11
2-11
2-11
2-11
2-12
2-12
2-12
2-12
2-13
2-14
2-14
2-15
2-15
2-16
2-16
2-16
2-17
2-17
2-18
2-18
2-19
2-19

Female
Female
Female
Female
Female
Male
Female
Female
Female
Male
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Male
Female
Female
Male
Female
Male
Female
Female
Female
Male
Male

Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn

L-3001
L-3002
L-2980
L-2981
L-2982
L-2983
L-2984
L-2988
L-2990
L-2991
L-2992
L-2993
L-2994
L-2995
L-2996
L-2999
L-3000
L-3003
L-3004
L-3008
L-3010
L-3011
L-30l6
L-3017
L-3021
L-3022
L-3023
L-3024
L-3025
L-3030
L-3031
L-3039
L-3040

113
37
114
115
117
49
124
127
126
53
54
120
23
121
123
133
134
135
136
122
6
24
55
25
26
56
125
57
128
129
28
58
59

2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
IN
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N

Location
Range

Sec.

96W
96W
96W
96W
97W
98W
97W
97W
96W
96W
96W
96W
98W
97W
97W
96W
97W
97W
98W
97W
97W
96W
96W
97W
97W
96W
97W
97W
97W
98W
98W
97W
97W

30
19
30
19
36
1
17
36
30
30
30
19
1
17
17
19
36
36
1
8
36
30
19
4
36
30
17
36
17
1
1
36
36

Units 11 and 23, \-lintcr

Remarks

--

--------------------------------------------------------------------------------------------------------------------

I

w
w

\0
I

�Table 1. Record of deer trapped and marked
1974-75 (continued) .

on the winter range in Game Management

Age

Eartag
No.

Neckband
No.

Location
Twp.
Range

Sec.

Female
Male
Female
Female
Male
Female

Mature
Fawn
Mature
Mature
Fawn
Mature

L-3042
L-3041
L-3049
L-3050
L-3059
L-3069

l30
None
29
131
60
190

2N
2N
2N
2N
2N
2N

97W
97W
97W
97W
98W
97W

6
6
7
17
1
17

Female
Female
Female
Female
Female
Male
Female
Female
Female
Female
Male
Female
Female
Female
Female
Female
Female
Female
Female

Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature

L-3074
L-3079
1-3080
L-3081
L-3082
L-3083
L-3084
L-3089
L-3090
L-3091
L-3092
L-3093
L-3094
L-3095
L-3096
L-3097
L-3098
L-3099
L-3100

132
30
137
138
31
62
l39
140
141
142
63
32
33
34
35
143
144
38
145

2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N

97W
96W
96W
98W
97W
97W
97W
98W
98W
97W
98W
97W
97W
97W
97W
97W
97W
97W
97W

36
30
30
1
17
7
7
1
1
17
1
18
17
10
2
2
17
18
18

Date
(1975)

Sex

2-19
2-19
2-20
2-20
2-21
2-22
2-23
2-24
2-24
2-24
2-24
2-24
2-24
2-25
2-25
2-25
2-27
2-27
2-27
2-28
2-28
2-28
2-28
2-28
2-28

Units 11 and 23, \vinter

Remarks

Possible

broken neck.

Has a red &amp; blue neckband
be 160.

Summary:

21 Male Fawns;

15 Female Fawns; .4 Mature Males;

Grand
Summary:

76 Male Fawns;

61 Female Fawns; 15 Mature Males; 157 Mature Females

55 Mature Females

--------------------------------------------------------------------------------------------------------------------

and # may

I

w
.j:0
I

�Table 1.
197LI-75

Date

Record of deer trapped and marked on the winter range in Game Management Units 11 and 23, winter
(continued).

Sex

Age

Eartag
No.

Neckband
No.

Location
Twp.
Range

Sec.

Remarks

Corrections to Previous Years' TraEEing Records
(Corrections are Underlined)
12-13-72
12-16-72
12-16-72
12-16-72
12-16-72
3-5-73
2-5-74
2-12-74
2-13-74
2-14-74
2-14-74
2-14-74
2-15-74
2-16-74
2-17-74
2-17-74
2-18-74

Female
Female
Female
Female
Female
Female
Male
Female
Female
Female
Male
Male
Male
Female
Female
Male
Female

Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Fawn

L-1697
L-1744
L-1745
L-1746
L-1747
L-2316
L-2580
L-2654
L-2655
L-2657
L-2658
L-2659
L-2662
L-2665
L-2666
L-2667
L-2668

96
125
126
None
127
17
88
135
136
127
126
118
116
133
142
None
146

2N
2N
2N
2N
2N
18
18
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN

99W
99W
99W
99W
99W
96W
93W
94W
94w
94W
94W
94W
94W
94W
94W
94W
94W

36
14
14
14
14
8
5
36
36
36
36
36
36
36
36
36
36

I
Vol

.po.
t--'
I

�-342-

Table 2. Sightings of deer marked on the winter range in Game Management
Units 22 and 23, March 1974 through February 1975.
Neckband

Date
(1974)

3-3
3-6
3-16
3-19
3-19
3-19
3-19
3-19
3-19
3-25
3-25
3-25
3-25
3-25
3-25
3-25
3-25
3-25
3-25
3-27
3-27
3-27
3-27
4-1
4-1
4-15
4-15
4-15
4-15
4-15
4-16
4-16
4-16
4-18
4-18
4-18
4-18
4-18
4-18
4-18
4-18
4-18
4-18
4-18
4-18
4-18
4-18
4-18

Color

Yellow wIred stripe
Yellow wIred stripe
White
Blue and white
Blue and white
Blue and white
Blue and white
Yellow wired stripe
Yellow wired stripe
Blue
White
Blue
Blue
Blue
Blue
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
White
Orange &amp; white blocks
Blue &amp; yellow blocks
Blue &amp; yellow blocks
Blue &amp; yellow
Blue and white
Blue and white
Blue and white
Blue and white
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
White
White
White
Green wlwhite stripe
Green wlwhite stripe
Green wlwhite stripe
Green wlwhite stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wired stripe
Blue &amp; yellow blocks
Orange &amp; white blocks
Orange &amp; white blocks
Orange &amp; white blocks
Red wlwhite stripe

Location
Number

Twp.

Range

Sec.

5
60
111
94
44
102
36
104
31
120
14
103
40
82
69
110
118
71
60
100
84
11
8
15
30
16
55
110
118
71

IN
IS
IS
IS
IS
IS
IS
IS
IS
2N
2N
2N
2N
3N
3N
IN
IN
IN
2S
IS
IN
IN
IN
2S
2S
IS
IS
IN
IN
IN
IS
2S
2S
IS
IN
IN
IN
IS
IS
IS
IS
IN
IN
IS
2S
IN
IN
2S

96W
96W
98W
97W
97W
97W
97W
97W
97W
98W
98W
98W
98W
99W
99W
96W
96W
96W
97W
93W
94W
94w
94w
97W
97W
97W
97W
97W
97W
97W
97W
98W
98W
96W
97W
97W
97W
96W
96W
96W
96W
96W
97W
94W
93W
93W
93W
97W

11
13
21
22
22
22
22
22
22
3
3
5
5
36
36
31
31
31
5
10
28
28
28
10
10
11
28
36
36
36
32
12
9
11
36
35
35
10
10
10
10
32
35
2
6
32
32
6

?
?
?

36
63
250
254
32
36
82
20
100
74
21
68
6
124
107

----------------------------------------------------------------------------------

�-343-

Table 2. Sightings of deeI' marked on the winter range in Game Hanagement
Units 22 and 23, Harch 1974 through February 1975 (continued).
Neckband

Date
(1974)

Color

4-20
4-20
4-23
4-23
4-23
4-23
4-23
4-23
4-23
4-23
4-24
4-24
4-24
5-19
6-5
6-6
6-8
6-8
6-14
7-7
8-20
8-20
8-20
8-24
8-26
8-7
8-7
8-7
9-4
9-5
10-3
10-5
10-5
10-5
10-l3
10-19
10-19
10-19
10-27
10-7
10-7
10-7
10-?
10-7
10-7
10-7
10-?

Orange &amp; white blocks
Blue &amp; yellow blocks
Blue and white
Blue and white
Blue and white
Yellow wIred stripe
Yellow wIred stripe
Pink
Pink
White
Green w/white stripe
Green w/white stripe
Green w/white stripe
Orange &amp; white blocks
Red w/white stripe
Red w/white stripe
Blue and white
Blue &amp; yellow blocks
Green w/white stripe
Green w/white stripe
Blue
Red w/white stripe
Yellow &amp; blue blocks
Red w/white stripe
Green w/white stripe
Orange &amp; white blocks
Blue &amp; yellow blocks
Red w/white stripe
Pink
Red w/white stripe
Red w/white stripe
Orange &amp; white blocks
Blue and white
Red w/white stripe
Yellow wIred stripe
Red w/white stripe
Blue and white
Blue
Blue &amp; yellow blocks
Blue &amp; yellow blocks
Pink
White
Orange &amp; white blocks
Green w/white stripe
Yellow wIred stripe
Red w/white stripe
Red w/white stripe

Location
Number

Twp.

Range

Sec.

19
8
101
43
5
92
59
30
52
7
68
25
14
7
57
7
7
7
7
113
85
7
7
7

18
IN
IS
IS
IS
IS
IS
2S
2S
2S
IN
IN
IN
IS
IN
IN
2N
IN
IS
4S
5S
5S
5S
58
5S
28
28
IN
5S
5S
2S
2S
2S
2S
28
28
2S
48
IN
2N
4S
4S
38
2N
28
2S
4S

94W
94W
97W
97W
97W
97W
97W
97W
98W
97W
96W
96W
96W
9lW
89W
90W
96W
93W
92W
94W
94W
94W
94w
94W
94W
94W
94W
93W
94W
94W
94w
94W
94W
94W
91W
96W
96W
96W
92W
90W
98W
98W
91W
89W
95W
95W
95W

6
29
2
11
11
21
21
20
24
30
22
22
22
24
9
35
8
30
22
5
34
34
34
20
34
11
23
1
26
26
14
14
14
14
10
31
31
36
24
19
20
20
26
22
24
24

?

139
7
100
4
7
7
21
7
7
28
7
7
96
6
16
?

7
7
7
7
7
7

1

------------------------------------------------------------------------------------

�-344-

Table 2. Sightings of deer marked on the winter range in Game Management
Units 22 and 23, March 1974 through February 1975 (continued).
Neckband

Date
(1974)

10-?
10-?
10-?
10-?
10-?
10-?
10-?
11-12
11-?
11-?
11-?
11-?
12-1
12-11
12-12
12-17
12-17
12-17
12-17
12-17
12-17
12-17
12-17
12-17
12-18
12-18
12-18
12-18
12-18
12-18
12-18
12-18
12-19
12-19
12-19
12-19
12-19
12-19
12-19
12-19
12-24
1-30-75
1-31
1-31
1-31
1-31
1-31
1-31

Color

Red w/white stripe
Orange &amp; white blocks
Red w/white stripe
Orange &amp; white blocks
Orange &amp; white blocks
Green w/white stripe
Yellow wIred stripe
Pink
White
White
White
White
Yellow wIred stripe
Green w/white stripe
Pink
Green w/white stripe
Green w/white stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Red w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
White
Green w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Pink
White
Red w/white stripe
Green w/white stripe
Blue &amp; yellow blocks
Green w/white stripe
Yellow wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Blue and white
Blue and white
Blue and white

Location
Number

Twp.

Range

Sec.

?
104
?
67
?
?
?
15
?
?
?
?
10
160
55
?
?
?
?
?

4S
IS
5S
IS
IS
IS
IS
2S
IS
IS
IS
IS
IN
IN
3S
IS
18
2S
2S
28
28
18
18
18
28
28
28
2S
28
28
28
18
IN
IN
IN
IN
48
48
38
2N
18
18
IS
IS
18
2S
2S
2S

95W
93W
96W
93W
93W
93W
96w
97W
100W
100W
100W
100W
92W
95W
97W
95W
95W
94W
94W
94W
94W
95W
.95W
95W
96W
96W
96W
96w
96W
96W
96W
100W
96W
96W
96W
96W
97W
97W
96W
94W
93W
94W
96W
96W
96W
97W
97W
97W

21
7
7
19
19
19
22
35
26
26
26
26
23
36
11
6
6
17
17
17
17
1
1
1
26
4
4
2
11
11
11
34
17
17
17
17
23
23
24
5
3
17
5
5

t
?
?
?
?

?
?
?
?

?
?
?
?
?

?
?

?
?
?
?

16
?
?
?
?
?
?
?

---------------------------------------------------------------------

5

3
3
3

--------------

,

�-345-

Table 2. Sightings of deer marked on the winter range in Game Management
Units 22 and 23, March 1974 through February 1975 (continued) .
Date
(1975)

1-31
1-31
1-31
2-1
2-1
2-19
2-19
2-19
2-18
2-18

Neckband
Color

Number

Yellow wIred stripe
Pink
Pink
White
White
White
White
White
(Unit 11)
Blue &amp; red blocks
B1ue &amp; red blocks (Unit 11)

Corrections
4-27-73
4-27-73

Location

Blue and white
Blue and white

?
?
?
?
?

229
236
239
42
18

Twp.

Range

Sec.

2S
3S
3S
2S
2S
2S
2S
2S
IN
IN

97W
96W
97W
98W
98W
99W
99W
99W
96W
96W

24
6
2
3
3
4
4
4
14
14

97W
97W

11
11

to Previous Sighting Records
132
131

IS
IS

Both of the above sightings should be omitted as there are no such numbers.

�-346-

Table 3. Recoveries of deer marked on the winter range in Game Management
Units 22 and 23, March 1974 through February 1975.
Date of
Recovery
(1974)

10-17-73
4-23-74
9-?
10-14
10-26
10-26
10-26
10-26
10-26
10-27
10-27
10-27
10-27
10-28
10-28
10-28
10-29
10-30
10-31
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?

Neckband
Color

Number

Blue
91
White
118
White
226
Green w/white stripe
200
Blue
117
Green w/white stripe
14
Red w/white stripe
39
Yellow wIred stripe
87
Blue &amp; yellow blocks
30
Blue
69
Red w/white stripe
22
Blue &amp; yellow blocks
31
Blue and white
36
Red w/white stripe
127
No neckband - eartag L-2065
Orange &amp; white blocks
108
Yellow wIred stripe
104
Red w/white stripe
41
White
24
Yellow wIred stripe
8
Orange &amp; white blocks
63
White
191
Red w/white stripe
11
Red w/white stripe
28
Pink
21
Green w/white stripe
31
Red w/white stripe
113
Green w/white stripe
149
Red w/white stripe
44
Orange &amp; white blocks
118
No neckband - eartag L-2137
Orange &amp; white blocks
71
Yellow wIred stripe
37
Yellow wIred stripe
60
White
6
Blue
118
No neckband - eartag L-2080
Red w/white stripe
1
White
20
Orange &amp; white blocks
65
Blue &amp; yellow blocks
63
Red w/white stripe
18
Yellow wIred stripe
94
Yellow wIred stripe
96
Green w/white stripe
133
Orange &amp; white blocks
112
Blue and white
37

Location
Twp. Range Sec.

5N

IS
IS
2S

IN
2N
2S

IS
2N
3N
2S

IN
2S
2S
2S
3N

95W
97W
99W
93W
99W
94w
95W
96W
93W
97W
98W
92W
94W
95W
97W
90W

27
32
3
20
19
9

?

?

?

95W
98W
89W
94W
loOW
94W
94W

31
35
14

3S
2S
2S

4

19
.
11 Found dead.
11 Found dead in fence.

?

?

?

IN

IN
IN

96W
96W
94W
96W
93W
95W
93W
96W
96W
97W
97W
97W
96W
98W
94w
92W
97W
96W
96W

33
11
17
11
7
29
19
22
22
24
1
12
17
14
25
17
24
34
34

?
?

?
?

2S

94W

?
?
5

3S

IS
4S

IS
2S

IS
IS
IS
2N

IS
2S
2S
2N
2S

IN
2S

Found dead.
Died soon after marking.
Found dead.

31
31
32
28
11
24
18
24
23 Died soon after marking.
27

2S
2N
2N

IS

Remarks

Road-kill.

Died soon after marking.
Found dead.
Found dead.

Found dead.
Found dead.

------------------------------------------------------------------------------------

�-347-

Table 3. Recoveries of deer marked on the winter range in Game Management
Units 22 and 23, March 1974 through February 1975 (continued).
Date of
Recovery
(1974)

Color

Number

11-17
11-?
11-?
2-28-75

Pink
White
Red w/white stripe
Blue &amp; yellow blocks

6
138
70
1

Neckband

Location
Twp. Range Sec.

2S

IS
3S

IN

97W
100W
97W
93W

20
26
12
20

Remarks

Found dead.
Found dead.
Road-kill.

Corrections to Previous Recovery Records
10-12-73

Blue

114

t

t

?

Omit - neckband no. is
wrong.

10-?-72

Green w/white stripe

76

2N

94W

20

Omit - duplicated in
same report.

��-349-

July, 1975

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-29

Deer-Elk Investigations

16

Job No.

2

~rkPlm

~.

Job Title

Piceance Deer, Study - Population Density and Structure

Period Covered:
Personnel:

April 1, 1974 through March 31, 1975

R. M. Bartmann and J. J. Klein, Jr.

ABSTRACT
The 1974-75 winter deer count yielded an average 7.1 deer per quadrat for
a projected population of 18,886 ± 3,458 at the 90 percent confidence
level. This is about 6,500 deer fewer than the approximately 25,500 predicted. Several possible causes are postulated for this difference.
It
may relate to deer distribution, sampling errors, and/or the use of a
different type helicopter.
Post-season classifications in mid-December
resulted in a buck:doe:fawn ratio of 18:100:85 on the basis of 2,817 deer
classified.

_

��-351-

PICEANCE DEER STUDY - POPULATION DENSITY AND STRUCTURE
Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.

SEGMENT OBJECTIVES
1.

To establish mule deer population density and sex and age structure
estimate techniques for the Piceance winter range.
a.

Estimate the size of the total wintering deer population in Game
Management Unit 22.

b.

Estimate the sex and age structure of the wintering deer population
in Game Management Unit 22.

METHODS AND MATERIALS
Deer Density
See Bartmann (1974).

Population Structure
See Bartmann (1974).

RESULTS AND DISCUSSION
Deer Density
The 1974-75 winter count, made January 30 - February 2, 1975, yielded 848
deer for a mean 7.1 deer per quadrat (Table 1). This was 1.1 fewer deer
per plot than in 1973-74. The population estimate for the Piceance Basin
was 27.4 deer per square mile or 18,886 ± 3,458 deer at the 90 percent
confidence level.
Timing of the count was about two weeks late due to problems of scheduling
a helicopter and, as in 1974, many of the plots located near the upper winter
range limit contained no deer. Fifty-two plots were devoid of deer this year
compared to 42 last year. The difference, however, was not with higher elevation quadrats, but with about 12 plots in the northwest part of the study
area that averaged 8.8 deer in 1974 and contained no deer this winter.

�-352-

Table 1. Number of deer counted on 120 1/4-square-mi1e quadrats on the
Piceance winter range in Game Management Unit 22, January 30 - February
2, 1975.
Quadrat

Deer

Quadrat

Deer

Quadrat

Deer

Quadrat

Deer

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30

29
21
4
11
17
4
14
0
0
0
0
0
0
11
11
16
12
10
7
15
9
6
3
0
11

31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60

12
0
0
19
16
8
0
7
0
0
0
0
35
9
24
8
37
5
0
6
14
10
17
0
3
1·
0
0
0
7

61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90

0
6
21
19
3
32
21
13
29
9
0
7
6
3
0
16
0
0
0
0
0
0
0
0
0
5
0
7
24
0

91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120

8
26
0
0
11
0
0
0
0
0
0
0
11
0
0
0
2
0
14
0
14
0
7
2
0
19
19
0
17
1

0

17
0
0
10

=

n
~X

-x
~ X

120

s

848
7.1
2

CZ X)2

15,312
719,104

2

78.31

90% Confidence Interval

s

8.85

7.1 ± (1.658)(.81) = 7.1 ± 1.3

s-

0.81

Total Population Estimate

C. v.

125%

x

18,886 ± 3,458

�-353-

The projected population is roughly 6,000 deer fewer than the predicted
level of approximately 24,700 deer (Table 2).

Table 2. Calculations
for winter 1974-75.

of projected population level of Piceance deer

Item

Bucks

Does

Fawns

Total

(22 bucks:lOO does:63 fawns)
1973-74 winter population
estimate
1974 winter mortality
estimate

2,596

11,778

7,438

21,812

159

745

4 416

5 320

1974 summer population

2,437 ~3,022

Fawns apportioned 50-50

1,511

1,511~

Adjusted 1974 summer
population

3,948

12,544

16,492

1974 estimated fawn
production (85 fawns:
100 does)

---

16,492

10 662

1974 pre-season population

3,948

1974 harvest estimate
(archery and rifle plus
25% wounding loss)

2 401 }:./

Calculated 1974-75 winter
population

1,547

12,544

10,662

27,154

12,544

10,662

24,713

1/
Harvest estimate is based on random questionnaire survey estimates
by an arbitrary 20 percent to account for wounding loss.

inflated

Several possible causes for the large difference between the predicted and
estimated 1974-75 winter population figures are postulated.
Among these
are the unknown effects of different deer distributions, the one in ten
chance for sampling error, the chance that the 1973-74 winter population
estimate was in error, that one or more of the estimates plugged into the
population formula were .In error, the use of a different type helicopter,
or a combination of one or more of all the above.

�-354-

In regard to the use of a different type helicopter, a Hughes 500C jetpowered helicopter was provided in place of the usual Bell. Deer were
noticeably less disturbed by the Hughes helicopter and, on several
occasions, small groups of deer stood under trees and were quite reluctant
to leave even when hovered over. An observer in the rear seat on the last
day noted several occasions when deer would run out from under trees after
they were behind the view of observers in the front. Visibility was poor,
especially for the middle man, and the tight seating allowed little movement which contributed to early observer fatigue. Also, the extra power
of the Hughes enabled counting during wind conditions when animals might
be more reluctant to move.

Population Structure
Post-season sex and age classifications were made December 15-19, 1974.
The Northwest Region biologist did not make post-season classifications in
the southern part of Unit 22 this year and so the entire area was covered
by research personnel. Deer were widely scattered and many were found
between 7,500 and 8,500 feet elevation.
The 2,718 total deer classified consisted of 238 bucks, 1,343 does, and
1,137 fawns for a buck:doe:fawn ratio of 18:100:85. The deer hunting
season was limited to antlered-only, so no information was obtained from
which to project fawn sex ratios or the yearling doe component.
Application of the post-season sex and age ratios to the 1974-75 winter
population estimate, while assumed in gross error, indicated 1,700 bucks,
9,254 does, and 7,932 fawns. A 50-50 sex ratio is assumed in the fawn
component. Application of the post-season data to the calculated 1974
post-hunting season population derived from the population formula will
not be done until the 1974 harvest data are available.

LITERATURE CITED
Bartmann, R. M. 1974. Piceance deer study - population density and
structure. P. 363-370. In Game Research Report. Colo. Div. Wildl.,
Denver, July {Part 2):185-398.

Prepared by

~~??~~~~
Richard M. Bart nn
Wildlife Researcher

�-355-

July, 1975

JOB PROGRESS REPORT

State of

COLORADO

--------~~~~----------,.
W-38-R-29

Project No.
Work Plan No.
Job Title

Deer-Elk Investigations

16

Job No.

3

---------------------------

Piceance Deer Study - Productivity and Mortality

---------------------------------------------------

Period Covered:

April 1, 1974 through March 31, 1975

Personnel: R. M. Bartmann, S. F. Steinert, J. J. Klein, Jr., D. Walsworth,
D. Roberts, J. Brown, F. Dieness, and L. A.Roper.

ABSTRACT
A fetal:doe ratio of 160:100 was calculated from a sample of five does
during spring 1974. Post-natal productivity, estimated from post-season
classifications of 2,480 fawns and does in Unit 22, yielded 85 fawns:lOO
does. The fawn:mother ratio was 100:100. The 1974 regular season harvest
estimated for Unit 22 is not yet available and only 19 deer were estimated
killed during the archery season. Three Division check stations recorded
459 antlered deer from Unit 22 during the first nine days of the ll-day
season. Yearling bucks comprised 30 percent of the check. The 1973-74
winter mortality estimate was 5,320 deer. A possible eight percent underestimate of the 1972-73 winter mortality was suggested by the finding of
seven carcasses missed during the 1973 survey.

��-357-

PICEANCE DEER STuDY - PRODUCTIVITY

AND MORTALITY

Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.

SEGMENT OBJECTIVES
1.

To estimate increments and losses to the Piceance deer popUlation
allow evaluation of density estimate techniques.
a.
b.
c.

to

Estimate productivity rates.
Estimate hunter harvest rates.
Estimate winter mortality rates.

METHODS AND MATERIALS
Pre-Natal Productivity
See Bartmann

(1972).

Post-Natal

Productivity

Post-natal productivity in the form of fawn:doe ratios are estimated from
helicopter classifications during mid-December in Game Management Unit 22.

Hunter Harvest
See Bartmann

(1974).

Winter Mortality
Procedures for estimating deer winter mortality in Game Management Unit 22
can be found in Bartmann (1974). The only change in procedures is the
addition of 16 quadrats bringing the total to 56.

RESULTS AND DISCUSSION
Pre-Natal Productivity
Only six adult does were examined for pregnancy during late winter and
spring 1974. Four were road-kills and two were trap mortalities.
Regular
checks for road-killed deer could not be made and fewer road-killed deer
were secured from management personnel for various reasons.

�-358-

All six does were pregnant (Table 1). Fetuses were too small to determine
sex in three does and the uterus of another doe was damaged so that the
number of fetuses could not be reliably determined. A fetal:doe ratio
based on five does was 160:100. Collection of pre-natal productivity data
will not be continued because of the undependable supply of animals.

Table 1. Pre-natal productivity data for mule deer does dying from various
causes in and around Game Management Unit 22, 1974.
Collection
Number

Date

Cause

Number of Fetuses
Male
Female

P-154

1-09-74

Trap !/

1 (sex undetermined)

P-155

1-17-74

Trap

2 (sex undetermined)

P-156

2-04-74

Vehicle

2 (sex undetermined)

P-157

2-14-74

Vehicle

P-158

3- ?-74

Vehicle

P-159

4- ?-74

Vehicle

Died

'1:./

1

1
1

160 fetuses:lOO does
1/
- Killed in trap by dogs.

2/

- Uterus badly damaged - only one male fetus found but more could have been
present.

Post-Natal Productivity
Previously, post-natal productivity data in the form of fawn:doe ratios were
obtained from pre-season classifications made from a helicopter during
September and October. In 1974, ratios were estimated from post-season
classifications in December. Budget limitations caused management personnel
to eliminate both pre- and post-season classifications on the Roan Plateau
and the southern part of Unit 22, respectively.
This would have required
additional research time and money to cover these areas. We decided it
would be better for us to expend additional effort on the post-season classification which is more crucial to the Piceance Deer Study.
The 1974 post-season classifications made December 15-19, yielded 1,343 does
and 1,137 fawns for a fawn:doe ratio of 85:100. The assumed reduction in the
proportion of two-year old does that normally produce one fawn may tend to

�-359-

inflate the fawn:doe ratio and preclude valid comparisons with earlier
year's estimates. Calculation of the fawn:mother ratio (the non-productive
yearling component removed from the total doe population, Anderson 1965)
was based on the estimated number of adult does carried into the summer
of 1974 as the antlered-only hunting season precluded information on the
female segment. The resulting ratio was 100 fawns:lOO mothers.

Hunter Harvest
The 1974 regular deer season in the White River area and nearly all adjacent
Game Management Units was limited to antlered-only from October 26 through
November 5. The archery season, also limited to antlered-only, occurred
from August 17 through September 22. The total harvest estimate from hunter
questionnaire surveys for Unit 22 in 1974 was 2,001 deer. The archery harvest estimate in Unit 22 is only 19 animals (Table 2).

Table 2. Estimates of the 1974 archery deer harvest and regular deer harvest
in four upper White River Game Management Units.

Unit

Archery Season

Number of Deer Harvested
(Antlered-onl~)
Regular Season

11

8

1,556

1,564

22

19

1,982

2,001

23

16

672

688

24

3

47

50

Totals

Four hundred fifty-nine antlered deer from Unit 22 were recorded at three
permanent Division check stations during the first nine days of the II-day
season (Table 3). The unknown effect of eliminating the Rifle station on
the total deer check at Idaho Springs prevents projection of total kill from
check station data. The percent yearling bucks in the check station tally
(30%) appears lower than the average 48 percent for the four years immediately
preceding the severe winters of 1972-73 and 1973-74, again possibly reflecting
the high fawn loss last winter.

�-360-

Table 3. Estimated age composition of the 1974 regular deer harvest in
four upper White River Game Management Units based on check station data.

Unit

Yearlings

Bucks
Mature

Unknown

11

33

121

81

235

22

117

266

76

459

23

23

47

36

106

24

6

7

6

19

Totals

179

441

199

819

Total !/

!/
Five illegal does and fawns from Unit 22 and three from Unit 11 were also
checked.

Winter Mortality
Dead deer searches were made from April 22 through May 23, 1974 on 56 1/8square-mile quadrats. This was an increase of 16 quadrats over the 1973
total. Fifty-eight dead deer were judged as losses of the previous winter
for a mean of 1.0 deer per plot, or 5,320 total dead deer on the winter
range (Table 4).
The sex and age composition of the estimated loss with unknowns apportioned
among the various classes was 45 percent male fawns, 38 percent female fawns,
3 percent adult males, and 14 percent adult females. The fawn proportion
increased from the 72 percent found in 1973. As before, the estimated winter
loss includes an unknown proportion of wounding loss and illegal kill from
the previous hunting season.
An additional seven carcasses were found on the original 40 quadrats that
may have died during the 1972-73 winter and were missed on the 1973 mortality
survey. This amounts to a possible eight percent error in finding dead deer
that first year and would have increased the 1972-73 winter mortality estimate by 1,064 animals to 12,768. In the future, the crew size is being increased to a minimum of seven people for more thorough plot coverage.

�Table 4. Results of dead deer searches on 56 1/8-mi1e2 quadrats on the Piceance winter range during
spring, 1974.
Quadrat
No. }j

2
7

9
10
11
12
13
16
17
21
23
24
25
26
27
32
33
34
35
36
37
40
42
50
52
53
59
62

Hale

-

-

-

-

-

-

Adults
Female

1

1
-

-

-

1

-

-

Unk.

Hale

Fawns
Female

Unk.

-

1

1

1

-

-

-

-

-

-

1

-

1

-

-

-

2
-

-

-

-

-

-

1

-

-

1

1

-

1

-

1
-

-

1

-

-

-

1

4

-

-

2
2
2
1
1
1

-

-

-

-

-

1
2
1

-

-

1

2

Quadrat
No.

65
68
70
73
74
75
76
78
81
84
86
87
91
93
95
96
99
101
107
109
110
111
112
114
115
116
118
119
Totals

Hale

Adults
Female

Unk.

Hale

Fawns
Female

-

-

-

3

1

-

-

1

-

-

-

-

-

-

-

1

-

1

-

-

-

-

1

-

-

1

-

-

-

-

-

-

-

1
2
1

1
1

-

1

--

-

-

Unk.

-

2

1

-

-

1

-

-

1

1

3

6

13

11

24

};./

Quadrats were renumbered in 1974 after the addition of 16 plots.
each mortality plot is located within.

Numbers correspond

to the census quadrat

I
Vol

'"

I-'
I

�-362-

LITERATURE CITED
Anderson, A. E. 1965. Population density and structure. P. 47-74. In
Game Research Report. Colo. Game, Fish and Parks Dept., Denver.
January {Part 1):1-246.
Bartmann, R. M. 1972. Piceance deer study - productivity and mortality.
P. 345-350. In Game Research Report. Colo. Div. Wildlife, Denver.
July (Part 3):253-377.
1974. Piceance deer study - productivity and mortality.
380. In Game Research Report. Colo. Div. Wildlife, Denver.
(Part 2):185-398 •

Prepared by

~~----

...
I

R. M. Bartm~~
Wildlife Researcher

""--

P. 371July

�July, 1975

-363-

JOB FINAL REPORT

State of

COLORADO
--------~~~~-----------

Project No.

W-38-R-29

Deer-Elk

Investigations

Work Plan No.

17

Job No.

l

Job Title

S~y_s_t_e_m_s
__M_o_d_e_l_lll_·~g~B_~_·g~G
__am
__e__P_o~p_u_l_a_t_i_o_n_s

Period Covered:
Personnel:

~

April 1, 1974 through March 31, 1975

James F. Lipscomb

ABSTRACT

Due to a change in work assignments from the Wildlife Research Center to
the Division's Planning Office in Denver, work was delayed on the final
report of this job.
A detailed description of the analysis method will be presented in a manuscript, which is in preparation, to be submitted to an appropriate journal.

_
_

��-365-

July, 1975

JOB PROG£'U::SS
REPORT

COLORADO

State of

W-38-R-29

Proj ect 1'10.
Hork Plan No.

Deer-Elk Investigations
Job No.

18

Job Title

Deer and Elk Management Study

Period Covered:

April 1, 1974 - March 31, 1975

Personnel:

l

_

Raymond J. Boyd, Thomas M. Pojar, and Bertram D. Baker

ABSTRACT
Slight adjustments were made in Data Analysis Unit (DAU) boundaries after
new data were obtained from the four Regional Biologists. Initial population models were completed on all deer and elk DAU's and a list of basic
dat3 needed to initiate the model is included. Special population models
were run on three separate occasions for the Wildlife Management section
in Denver to assist them in solving problems arising from field recommendations.
PART II. Browse utilization-composition and pellet group count data were
compiled from records of 557 transects for Uncompahgre deer DAU D19 for the
period of 1967-1974 and 631 transects for White River elk DAU E6 for 19631973. Linear correlation regression and t tests were done on deer and elk
population parameters of mean animal days per acre use, aerial trend counts,
and antlered harvests in exploratory efforts to evaluate them as population
indicators. Relationships of key browse utilization and animal days use per
acre were also tested to determine correlation coefficients and feasibility
of use for prediction equations. In general, lack of both statistical significance and performance credibility provided justification for the recommended
conditional discontinuance of the utilization portion of present interagency
big game range surveys. Other recommendations are presented concerning the
future of the interagency cooperative range program.

��-367-

DEER AND ELK ~ANAGEMENT STUDY
Raymond J. Boyd
Thomas M. Poja~
Bertram D. Baker

There has always been a need to quantify management objectives for wild
populations. The difficulty of obtaining reliable information upon which
to base these objectives has long been realized by wildlife managers.
Methods for estimating various population parameters are available using
the computer population simulation approach.

P. S. OBJECTIVE
Devise and test a statewide deer and elk management system.

SEGMENT OBJECTIVES
1.

Divide the state into a system of data analysis units (DAU's) that are
deemed practical for gathering and analyzing population data.

2.

Based upon currently available information, estimate population parameters.

3.

Identify data that are most useful for improving estimates of population
parameters and recommend procedures for collecting such data by DAU.

4.

Project average population parameters on a long-term basis. for those
alternative management plans that are feasible for each DAU.

PART I

MODELING DEER AND ELK POPULATIONS

Initiation of Modeling Procedure
Several modeling workshops, supported by the Division of Federal Aid, U. S.
Fish and Wildlife Service, were attended by the regional biologists and the
project investigators. These workshops stressed the potential uses in game
management of the modeling approach. Several big game populations were
modeled during the workshops which demonstrated the type of information
necessary to initiate a model and to familiarize those in attendance with
the format of the output. In addition, the investigators assigned to this
project underwent a period of intensive training, to learn how to initiate
and manipulate the model data card deck.
To construct a data deck the basic information listed in Table 1 is needed.
Most of the items simply involve setting up the calendar of biological events
of the species being modeled. In general, this type of information is available for the State of Colorado and is not expected to vary a great deal from
one area of the state to another.

�-368-

Table 1. Checklist of big game population data used to initially set up
the model.

1.

Initial year the simulation is to start ·and how long the simulation is
to run (in our case, we start in 1963 and run for 16 years).

2.

Determine length of mortality periods (in our case we set three mortality periods; June 1 to the day before hunting season; the hunting
season period; and the day after hunting season through May 31). These
three periods are broken down into the percentage of the year they each
make up.

3.

Choose the minimum age class to receive hunting mortality

(in our case,

calf or fawn).
4.

Choose the minimum age class to receive sex-selective hunting mortality
(in our case, yearlings).

5.

Choose the minimum age class of males classed as trophies.

6.

Estimate the time of breeding.

7.

Estimate male·and female wounding loss.

8.

Determine time of year when pre-and

post-hunt mature/young

ratios are

counted.
9.

Estimate total numbers of males and females in the initial year and set
age structure for 10 age classes based upon survival rates from check
station aging data.

10.

Enter total antlered and antlerless kill for each year in the simulation.

11.

Enter average weight of males and females, by age class.

12.

Estimate the maximum percent of adult and yearling females that become
pregnant.

13.

Sex ratio of the young at birth.

�-369-

For example litter conception dates of 50 mule deer collected (1960-64)
from the Cache la Poudre drainage of north-central Colorado were estimated (Anderson 1972). Ninety-six percent of the fawns in these litters
were conceived between November 17 and December 22. Approximately 93
percent of 173 litters examined during 1969-72 in Middle Park were conceived from November 16 to December 15 (Gill 1972). Similar results were
reported from west-central Colorado during the period 1949-1955 (Anderson
and Medin 1966). Therefore, conception dates and consequently fawning
dates would seem to be fairly consistent at least in the northern half of
Colorado.
Other information, such as wounding loss, is known to be a necessary part
of the model, but even a crude estimate of the magnitude of wounding loss
in Colorado is lacking. Consequently, the number used for wounding loss
in the model is the result of opinion tempered with some information reported from studies in Utah (Costley 1948 and Stapley 1971).
The two above samples show the extremes in availability
of the information used as input data for the model.

and consistency

Data Analysis Unit Boundary Changes

Elk
After tagged elk kill returns were analyzed from the 1974 elk hunt, it was
obvious that DAU boundaries between E-3 (North Park) and E-8 (Troublesome)
would have to be modified.
Tag returns indicated that there was considerable
elk movement back and forth between units 17 and 18 that was not taken into
consideration when the original boundaries were drawn up. Therefore, unit
17 was taken out of the North Park DAU and included within the Troublesome
DAV. The North Park DAU now consists of units 6, 7, 16 and 161 while the
Troublesome DLV includes units 17 and 18 (Fig. 1). No other changes were
made in elk DAU boundaries during the past segment.

Deer
Three deer DAU boundary changes were made. All of unit 11 was included in
D-7 rather than splitting it and putting part in D-6 and part in D-7.
Approximately the west one-half of unit 65 was separated from D-25 and
called a separate DAU. The northern tip of unit 59 was placed in D-17
rather than D-16. These boundary changes are represented on the DAU map,
Figure 2.

Initial Population Models
Initial population models based on available information since 1963 were
run on all deer and elk Data Analysis Units (DAU's) in the state. The 1974
pre-hunt population sizes for all deer and elk DAU's are shown in Figures 3
and 4. All of the models were reviewed at least once with appropriate
Regional Game Biologists, with the exception of Northeast Region deer DAU's
which have not yet been reviewed. When necessary the models were revised
to conform more closely with the characteristics deemed to be most correct

�-370-

by the biologists.
The top ten DAU's (rank based on 10-year average
harvest) for both species were reviewed twice with Regional Biologists.
These DAU's account for 75.9 and 62.5 percent of the average ten-year
harvest of elk and deer, respectively.
When reviewing the DAU population simulation with the biologists, each
simulated population parameter was carefully scrutinized.
In most cases
there were no observed data available to compare with the simulated data.
To judge how realistic the simulated data were, we relied heavily on
comparisons between simulated values and comparable data reported in the
literature and on data from other parts of the state. This approach resulted in what was considered the best possible characterization of the
population inhabiting each DAU. Examples of simulated population data
are presented in the Appendix.
Near the end of Segment 29, a more versatile version of the model was made
available to the project. This model has the ability to plot the tabled
output (Fig. 5) and to plot simultaneously the simulated output and any
observed data that is available.
It also calculates the R2 value from
which the correlation coefficient for the relationship between observed
and simulated data points can easily be obtained (Fig. 6). These features
of the model enhance the simulation alignment process.

Response to Management Problems
If the model has been aligned to historically mimic the real population,
it can be used to estimate how the population would respond to candidate
management strategies. On three occasions during the last segment the
modeling approach was used to furnish Game Management with probable consequences of management strategies under consideration.
The first case involved a decision of whether or not a post-season in two
of the deer DAU's was merited. The second case concerned a management
decision on whether or not to close the hunting season in three deer DAU's
that had experienced high over-winter mortality for two consecutive years.
Several options for attaining desired population goals were generated and
provided to Game Management for each of the above situations.
In the third case, simulated elk population data at the close of the 1974
elk hunt were used in order to judge whether proposals for post-season
hunts in seven elk Game Management Units were justified.
Check station
data on numbers and sex of all elk taken from these areas were projected
by the Wildlife Management Office in Denver to furnish minimum, maximum
and mean harvest estimates for each of the seven areas. These harvest data
were programmed into existing models of each population, and simulations
were run with each harvest figure to estimate popUlation parameters under
each situation. The three runs for each area were sent to the Denver Office
to assist management personnel in their assessment of potential post-season
hunts. These data were made available to Denver one day after receipt of
the kill estimates, which is a marked improvement in rate of data processing.

�-371-

LITERATURE
Anderson, A. E. 1972.
Xerox. 23p.

CITED

Summary of results .of the Poudre Deer Study.

Anderson, A. E., and D. E. Medin. 1966. Colorado Game, Fish and
Parks Dep. Game Information Leaflet No. 60. 4p.
Costley, R. J. 1948. Crippling losses among mule deer in Utah.
Amer. Wildl. Conf., Trans. 13:451-458.

N.

Gill, R. B. 1972. Middle Park Deer Study - Productivity and Mortality
p. 179-198.
In Game Research Report, Part II. Colorado Division
of Wildlife. :federal Aid Proj. W-38-R-26.
Stapley, H. D. 1971. Deer illegal kill and wounding loss.
Job Final Report. W-65-R-D, Job A-8, Utah. 6p.

Federal Aid

~.~~~

Wildlife Researcher

Thomas M. Pojar
Wildlife-Researcher

�I

W

"
N
B

Fig.

1.

Elk Data Analysis

Units.

�I

W

"
W
I

Fig. 2.

Deer Data Analysis Units.

�140

n

130

en
o

z

40

.-...-

-

.--

ct

en
:::&gt;
o
:J:

~

a:

I

.--

30 '-

w
w
o
u..

W

.....•

~

.-- .-- .--

o
a:

w

20

I

.--

-

r--

CD

r-- r--

JE

:::&gt;

r--

-

z

10

r--

.--

r-r--

-

-

r--

.-- .--

r--

- .-- r--.--

.--

r-- r--

-

r--

o

I'-

Fig. 3.

~
(\J

State-wide

(\J

to

1974 pre-hunt

0

0

If)

(\J

to

(1)
(\J

deer population

v

(\J

v

I'(\J

(\J
(\J

(1)

DEER
DAU
NUMBER
estimate - by simulation.

I'-

o

If)

658,782

If)
If)

If)

(\J
If)

CD
(\J

total animals.

(\J

If)

It)

to

C\I

It)

If)
(\J

�25

u&gt;20
0
Z
&lt;t

, (f)

::&gt;
0

::I:

1 II
-'

I

15

w
-...J
Vl

:

I

W

LL

0

ex: 10
W

!D

~

::&gt;

z

5

OB

I II

II

U)
rO

II
V
-

II
10
N

II

II
N

N

II!I
V
N

N
rO

II
rO
rO

II
U)
N

II
10
-

II
U)
-

II
0

en

rO

N

II

II
I'-

ELK

Fig. 4.

II

II
~

0 AU

~

II

II

II

~

II

II

II
.-

II

II

II

II

-.-

NUMBER

State-wide 1974 pre-hunt elk population estimate - by simulation.

134,417 total animals.

IDODCJCJ

�'.'800.

~~=~C~S&amp;~C~&amp;.&amp;~S&amp;C.CCC=&amp;&amp;CC&amp;.&amp;C~==C&amp;C=~&amp;c.&amp;.&amp;a&amp;cc.:c.c.C&amp;C&amp;&amp;&amp;&amp;==~C

1

!

1

•• CUC~.C&amp;

••• &amp;~~~~.~~'.7C~~'"

_ •••

_.

1

!

'2~ao.

1----

•

!
!

,

"1:----

, '200.

"

"
"
'~40().

I!----

"
"
%00.0

~
~

•

t!----

~

"

. --

B

~

~

~

B

B

,

B

B

"

-··_··a

•
"

•

B

--

r.

4

(
(
(

~
o

"

"
"

(

(

"

(

"

0

(

B•

a----

-----.

0
0

B
'200.

- ...

B

B

1:l~OO,()

"

B

!!----

~
88()()./)

~

0
[

D

---~~

D
0

f,4()/),()

(

B

t!".

- -

-

(

----a "

0
(

r.6()(). ()

•
•
•
•

0

0

----.

,

o

1

48()0.(I
C~~~&amp;&amp;~~t~c~~C&amp;C&amp;c~&amp;.&amp;a.&amp;.cccc.c

'960.(1

1

1

,

,

,

1

,

1968.0

1966.d

'972.0
1970.0

1916.0

'914.0

•

198~.0
1978.0

T 11&gt;£

Fig. 5.

.•

••• cc •• &amp;&amp; •••••••••• &amp; •• &amp;.c=c.&amp;.cccc=c~~=~C&amp;C~===~&amp;c==cuc====c==~c~=c~&amp;

'96'.'0

'962.0

"
"

0

~

"
.----

"

Example of a computer plot of big game population

data taken from Table 2 in Appendix.

I

W

~
~
I

�======~~====~=~===~:===========~===================================c=====================~=======~==

2?7SP.

~

ill
1
1

:=.

s

1

1
1
1

1
1

1

1
1
1

1
1
1

1
1
,

=

1
1
1

=
=
==

1'105::&gt;.

=

=

1Cl341,.

==

=-.:--~

1503".

=

.

_ .._

17F,4L•...
__ ..

=
==

s,

,
::,
~
;:

=

,

....

;----,

}422Cl.

/\
./-

=

=

\

./

'9'

=
=

\
-\ ..

\
12523 •.....

\
\

\.

10817.

==

,,

=
=

=

=

9111.6

==
=
=

HOS.@.

'-

==
&lt;;700.0

:==================:==========================~
1 &lt;H, &lt;. r

lq",,,,.n

11=
11=
11;

~================================:===================
lQ7'.O

196°.(1
19,,7,&lt;;

1'-11'4.5

1971\."

107".0
lq7~.5

lQ78.0
1976.0;

TI~F
POST-Y4RV~ST

Table 6.

-...J
-...J
I

\

'._

I

W

(C~TTW"1

Example of a computer

PODUlATln~

StZF

plot of observed

rS=sIMuLATEO.o:DEpnoTFDI

and simulated

population

D~QR=

.015

sizes with the R2 value also computed.

��-3'19-

APPENDIX

��-381-

============================================================
============================================:===========:===

=

=s

=

RESULTS 01="
I MUJ-A1J ('IN TP I AL ~-!IJ~"RE~ __ I. ~

==============================
=========================
.

=

.._

.. __

..

E::UC

SPF:CIES

.

-.0_"

=

=========================
================:===========================================

============================================================
1
=============
AGE CLASS

STZES

AT START

OF EACH

VEAR

================:===========================~===============
============================================================

==== ======
1963 2000.0
1964 2048.~
-1965 _2285: 81966 2391.7
1967 2411.9
--1968 2409-~4
1969 2~61.3
1970 2276.?
-.-1971 24 37 ~5 -)972 2557.5
1973 2206.6
1974 2164.7
1975 2697.?
1976 22C)4.3
1977 2064.4
1978 2045.6

AGE CLASS
YEAR

==== ======
1963
1964
1965

291.0
313.4
4n'i.t;

19M)

487.1',

1967
1968
1969
1970

59".,
567.3
573.4
6;&gt;1',.0

1971

63').1',

1912
}973

f-Jt;f).r)

1q74

1975
lCl76

19-,7
}q78

_1\15.

________ ~14

YEAR

======

======

(H8.0

171}f1.0
16')1.1

1312.'i
1235.2
1218.9
1369.3 .
14qq.3
1495.,
1::l89.1
1257.9
1531.8
1679.1
llQO.9
1603.1
If,14~q
2012.1
1681.7

16-28.-9--

1801',.5
1889.4
,9f9~r
1899.?
1 78~. 1
-180i;;-~~
1941.1
?,023.4

1740-. (,
}753.4
2184.7
182f,.O
167?2
~IZES

AT

sTAIn

OF

EACH

._

-----------

-----------

614.0

412.0

7

C; '_1.?

lO::l~.c;

647.4
A05.6

_____
J9?

94?1
~ ~ n •~

.?~_~
__
.J

110".5
766.1
117A.8
A74.~
II 09 •f,
A8q •c:;
1016.0
~?6.7
IOC;1.R
859.9
12? 1 •A______ "56.4
1?96.3
955.6
IlnJ.CJ
1195.1
l476.6
__)1\ 19_.6
14R7.t;
1160.3
)~C:;~.4
1~70.3
YEAR (CONTTNlIED)

t-.17

NR

t-IO

======

======

======
, J4.0

======
At;.o

'?c;.o
13~.?

ql.R

1

H\C:;.~
111.0

21 1 • n
239.,
2AI;.C:;
311.,
37".9
467.1
42q.4
45C:;.1
504.1

154.0
175.4
1~3.S
20Q.4
?44.'
2~7.'?
357.7
320. :
31?7
36'"'.7

66".1\
619.5
i:l19.7
l022.A
879.1

5r.f,.-:)

39?~

500.4

387.;t
432.7

1162.(1

7SA.3

41q.A

551.,
7(n.c;
87,.4

475.7

6(l4.!1
749.7

'~A.7

,lC;9.9
q, .?
??4.n
?f"~.1
'?}C:;.C:;

?4':'."
'.l7A.4
?Q?4
"'l:::»4.7

'16:::».6
1Q7.7
1:;(\".5

.

97.~

1?4.7
143.q
16".9
195. ,
lR4."
1 R6.1

"'1'16.9
?43.:I
?f,9.9

1'11.'
12Q.R

.__ .

_

_

_

�-382-

======================================================:=====
============================================================
T A
-

8 L E

?

=============
- .. -.... ------

--------..

.

-_ _---..

.-

.--

POPULATION NUM~EP AT BEGINNrN~.
PRFHARVE~T.
°O~T~APVEST
AND END OF YFAR

= = = = = =====~==-~:= ==: ~::==:=::= == =-====::::=:==:: == ===-== =:: == = = ==:::===== ====- -------------------------=====================================~======================
C

===

(

_ ~JZf
AT

(

I

t
~

C

•

YEAR

STAPT

====
--1963
1964
1965

6559.0
123R.6
1932.3

-----

-rqtj6-

- 84jr~-7

1967
1968
---i969
1970
1971
--j"qj2
1973
1974
1975
)976
]977

8817.4
9342.1
9437.2
924".5
___9L?i? 7 _
.99115.6
_10010.9
9374.2
10722.5
11384.5
11807.7
1~12t3.4

]q18

===

SIZE

~ __

PREHUNT

---------

6325.7
6990.7
765~_.;l
8136.':'
8532.9

______
,!O~?_~_.3
9126.7
8937.1
_ ~~_55.5
9t;67.~
9f.9?5

9065.6

--}C14~. 6
1]027.9
11447.6
-1 "75-1 •..,

D

---

---

SIZE
AT
POSTHUNT

=:=

PERCENT
___~_l7E _ .
OT
END
.~.::==.==

=====

C H "N G E
(BAC:;EO
ON STAPT)
....========::;

..

5620.3
519~.4
/).0
l'112.8
564".~
10.4
6.&lt;)J ~ • n
Q54:~.6 _____
9. F,
._.
_
F-971 .5
64ct;.C;
F,.3
7521.A
69'3:;1.1=1
4.q
7?_~6.6
_.lJ]C:;.9
5. 1
_
-1573.5
697n.,
l.t}
7332.2
67315.3
-c.O
80_()5.0 __7_:34A.1 - --------------~- -~----------8490.6
7804.4
8.0
7849.4
721)9.~
1.1
~749.3
_.___
~(l?5. ,
______
.- 6.4
_
9915.4
9131'1.~
14.4
10634.Q
9741.1
6.~
11026.0
.
3.1.
_J OO}9.~ ___
1l3{\2.~
1(l3tl~.f
2.7

_

..

(
_

t:

_ E

••••••••

__

._

0·0

~

.__

�-383-

==========================================:~==========:=====
==========================================~=================
T

.A

'3 l E

3

- =============

...

OF A I\JTLERlE 5S AN IMAL &lt;; IN EACH AGe'
CONTAINED
IN HARV~5T

r-!11~8EQ

CLASS

=========================================~~========~========
==============================~==========~=~==========:=====
YEAQ

N1

1'1i?

~J3

====

======

=======
27.f-l
.___~c; .3

======

1963
1964
196~
1966
1967
1968
1969
1970
1971
1972
1973
1914
1975
1976
1977
1978

17.7

31.6
47.2
51.8
36.~
C;4.9
47 .•
7
39.7
28.6
50.4
4q.f)

.0

... - ._ ..

...

71 .7
,?:;&gt;.O
-

80.::?
73.4
__?7.
3A.R
70.4

c

__ !i?_~3._
.0
.0

.1}

.n

20.7
43.~
~3.1
55.1
42.1
69.1
62.9
51. q
33.3
58.0
68.0
.0
.1

61 .6

.0

.n

._-

.

.---~I)

.!l
'.0
.... _"-_ ... _-_.-

NIJ~BEP
CLASS

"J4

-----------

OF FEMALES
cnNTAINE~

-----------

lS.r:;
3? .~

11. S
24 •!L_. _

S".C;

17.5

47.6
:3? •6

44.9

.0

2 7. B __ - -42.9
43.7
37 .cL
2S.F-J
44.7
.___ 47.5
.0
.0

()

.• I) .

._._
..•0.

• I}

.0

.n

.0

.0

•• 1'1

.

c;~.3

_

1:)4.?
4 4 •4 __
3n.?
49.7
~_5~. 0 ..

.n

I"l EACH At';F:
IN HARVEST(~0NTINUED)

N6

N7

N8

----

======

======

------

lq63
1964
1965
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
916
1977
1978

~.~

16.q
26.4

31.~
24.A
"34.6

::n .n
~8.9
20.7
'3t;.8

.

---_ 6.5
..

12.7
P~.5
2~.2
17.5
30.g
25.')
20.q
11:).8
2q.1)
3?4

.-

4.7
9.~
13.t5

.__ .. -. 15.1

11.q
21.1
22.!J

16.1

.0

.0

.n

.0
.0

.0
.0

.n

____
2.6 "-----.----_
4.9
7.0

...- •..... - .
'

9- .7 ••

..

-_

q.s

7.4
11.6 ---------- .. ...- --- .
11.n
-

./')

.0

.n
.n

.t)

.0

. -.-...

•....._-_.

.0
.0
.0_

_.--_ ...--

...•

.---_._._-- -_.- ---_ .. , ..

9.7
10.1 ----_.'- ...

l~.A

,.

....

S.A

FI.6

.0

.1)

.

A.l

It;.O

.1)

....
__
._-----.

14.4
It;.t'J
14. 1

------_._-

-----------

1,.7
o.B
11. n

_

~!10

3.5

11 •1

21.4
25.4
.0
.0
.0
.0

40.4
.n

======

.._..

__ _-----_._---_._._--_

--_ .•

P&lt;.jq

YEAR

.__
~.

----- -

--".

..

-- - --.

-

�-384-

==================================:=========================

==~===================================================
======
TAg
L E
4
=~===========
NUM8EO OF ANTL~PED ANI~AlS
CLASS CONTAINED IN H~pVFqT

IN EAC~

AGE

===~======================:====~==================~:========

====================::=================~===========:===:=====
YEAR

1

====
1963

====~=

•n

N::&gt;.
:=====
231.4

1964
1965
1966
1967
19n8
1969
1q70
1971
1972
1973
1974
1975
lq76
1977
1q78

.n

_20?_.JL_

.1)

.n

227.3
210.6
__1q5 •.
_5..._
lqR.q
296.7
341 __
4- ____
95.8
4S.5

.n.

_f&gt;lQ .7 __
.0

'\!

.0
• Q_---

.n
.0

•.n
.t)

.,:, ._--.

.0
.IL

~n
-----93.4
138.q
149.::,
133.4
124.2
139.3
213.1
221.2
78.q
136.4
188.5 _
.0

.0
.0___
.0
.0

.0

.n
"'

.--"----

NU~RER

r-Jf,

---- ======
1963
1964
19115
1966
1967
1968
1969
1970
1971
1972
1973
Iq74
1975
1976
1977
1978

t;.q
7.9
14.7
:?l .c;
2q.o
VI.l
4q.2
~1.5
?1. 1

?~.c;
34.n

1B.3

.n.
.n

2?8
.0
.0
.0
.0

•n

.0

.n
• f)

_2l.8

.n
- -----.

-----. (!

.0

.~

.4
.q
..
__
1.1
1 .4
~.4

'=.5

co.o

l?l

7.1
10.5
14.5

4.1
5.4

._._---.

....

._-

-n 0

======
•

0 ___

"_._--

-.0 ----------- -.--- ---

.1
._ .....

.- .-

..

~.,

3.1
4.?
.0

.0

./}

.1"

.0

.n

---- '-----_ •..

.1

-------.---..
-______
•4_--- -------._--- ..
.5
.9
....... -._---2.3
.. --_._-_ ----.4.3

.n

.n

....••..

__ ----------._.- -_._-- --._---- ------_._----- "---"'-

.0
.0
.0

-. ---- .------_ .

----- .-.. "---- --- _------_.-

.'.-._.'-"-

R.l
.0

.r}

.

.n

======
-

.•.

n
.n

.n

~.•-.. --- --_.

..

.0
.0

NQ

]8.4-

....

__

.n

.

1 .1
2.?
2.8
3.Q
6.9

_._---------

.

.

IN EACH AGE
IN HARVF5T(C~NTINU~O)

======
____
.8

..

-

.0

======

.

n;:&gt;.?

-_ ..

N8

----- --- -.-.-

___
_._------- --------_.- .40.0
P,7.,
5R.7
71.8
_46.2
-- -.. _----- _._-p,R.S
50.1
14Q.3
q?.Q
-158.9 ---- -- ---1- 09. n -------------_ _- -------t;1 .1.
3n.f)
6R .•l
43.::&gt;
.---1-19.3
-------.--58.:1 ....• ._-----.- ---- ._--------

.0

N7
2./0. ___
3.2
5.3
7.9
10.6
lR.q
2q.~
33.6
13.4

,

37.::1
SI5.6

.0

OF MALF~
CL,sS CONTAINED
YEAR

======

__0 ___

.

',15
=::====
14.'5

"14

.0
.0

..

-----_._----_ ....

.

_.-.

. ---_ ..

-

-.--

�=====================

=============================
=============================

-385-

=====================

5

T A 8 L E

=============.
·TOTALS A~O PERCENTAGES
ANn FE~AlE~
HAPVESTEO

OF MAL~~
AND IN HaRVE~T

===================================================~========
============================================================

===

===

NU"1gFP
OF

YEAR

I(

(:'

I'

f\1l.JM~EP
FEM.ALES

HI
HAPVF~T

TN
HAP\jtS·T

HApVEST

=======

()F

TN

=======
11 o ,
_ .. -

3q7.
448.
S660

17.4
_ _..- _
17.3
19.3

S7~.

1 ~ .!.g. __ .

501.
563.

14.9

8A~.!,.

984.
326.

...

..• ...

,·5.6
?~_~..EL
?A.l

===

PFq("Ei'JT
OF .

PEPCF:~!T

MALES

OF
TOTilL

POPIJLATION

------------5n7•

==========

=======
2~A

?1l.

4.q

101.
"33:1.
?41.
.387.
362 •

~.S

"301.

TnrAL

TOUL

FEMALE&lt;=; NUMRF:°
IN
IN
HARVEST
HAOVFC;T

6.8
4.7
7.2
...6.•q
5.6

3.7

6O:;Q.
867.
91l~.

74? •.
950.
.J 244 e .; _
l?A;.
532.
.745.
1494e
n.

9.9
'0.2
?7.4

:"06.
3f&gt;1.
409.

.0
..• 0
.0

().
O.

o,

.0

fl.
0•

n.

• (;

0 ••

.0

n.

o.

_.•0

O.

.0

O.

38~.

1085.

19i'6

1977
1978

PEQCENT
nF

T"TAL
MflLES

=======
1963
19(A
1965
1966
i967
1968
:969
1970
1971
1972
1973
1974
1915

E

D

A

6.3
7.1!
.0
.0

H'
HflPVEST

B.l
9.5
]1.4
Ll.2
8.8
10.6
._13.•7
14.5
6.1

0_·'·

_

•••

•

•

_

7.8
15.5
.0
...• 0

.0
.0
.0

......

0_--·_·_--_··- ....-----

..-.-

..-.

�================================,

================================~
T A 8 L E
=============
TOTALS
ANn

- -AND

PEPCEf\;T

-386£,

AG~S

nF

A", T L E P L E 5 S H A R V E c:; T E ('l

::===================

~====================
ANT'-~RFD
11N f)

pJ

HAP V ~ S T

======================================================:::===:
=

�-387-

============================================================
====~=======================================================
T

A

8

I

E

=============

7

pEPCENT~GE
OF MAL~S TN ~G~
CLASSFS AT START nF HUNTrN~

==================================:==:======================
============================================~===============
YEAR

N1

I\I?

1\;1

==:=
1963
1964

======
C:;0.0

======

======
3.•.•
5

------

19h5
19~6

S11. n

41.6
41.0

27.5

1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978

c:;o.n

49.7
49.7
49.7
49.7
4°.7
49.7
49.7
49.7
49.7
49.7
4q.7
49.7
49.7
~9. 7_..
49.7
49.7

SO.O
SI).n
511 •.0

50.a

SO.n.

So.n
SO.n
50.n
51).0

50.0
5('.0

So.n
So.n
_.

4-?1'.
43.1S
44.f1

45.4
41.4
3.3.?
47.4
50.0
27.6
49.2
49.2
49.?
49.?

~.!'"

;&gt;?('I

~IS

-----------

l/'\.A

1?9
16.7
2?'
?A.7
30.4
31.R
14.;&gt;
32.3
26.'1

It...4

;n.1

43.4
4C:;.n
;&gt;7.;;&gt;

30.?
111.":1
44.2
26.6
47.9
-47.9

"=IC:;.?
~.;.?

~';.7
~A.tS
41~. ,

:F.?

4g.7
4A.7
4R.7

-

pERCENTdGt:
0F fl.1ALESIN AGE rLllSSEC:;
OF HUNT rNG (CONTI r.JIII='O)
AT START
YEAR

\if,

------- ==::===
1963

7.4

1964
19,,5
}'&lt;66
1967
lC168

1969
1970
lq71
1972
1973

1974
1975
1976
1977
]978

9.4
13 ••n
17.3

;:&gt;3.",
:?C:;.~
;&gt;7.6
?f).7

22.c;
2'3.3
23.M
?"&gt;.1

:n.l
4 j •1

?S.~
(, f., • ,Co.

"'7

N8

NO

"'to

-======

--~--------

-----------

::.::=====

4.2

1.9

3."

~9
1 .4

.t:..

4.1

?3

1 •()

5.S

-.('1

1.7

7.Cl
11. S
17."

4.?

e,.,

?4
':; "-

21 • t)
1'1.;;&gt;

] f:. • 9

1? .•5

7.f)

]4. ,

R. 1

19.c;
20.2

i 5 .,~

lC'l.A
cS.l

17 •,
19.4

, 1 • /'\
1 ? •1
11.S
14. 1

3A.)
4;:&gt;.1
24.9

24.9
3S.~
41 •&lt;;

5.,
7.2
9.9
11.A
19.1)

2?.7'

17. I

9.e,

1':'.5
1".7

.0

.

5.,
9.4
1('1.4

11. ?
J3.7

If,.r.

?4.f'l

lB.?

,':".7

;;&gt; ••

4

�-388-

f

==============================~=======================
:=====
=====================================================:======

'-

T

8 L E

A

===========::=
OF

YOUNG

PPOOIJrED

INDIVIDUAL

AND

CO~8IN~D

NuMqF:P

(

c

_________ 8_

--____YDUNG

-

pFP

ADIJLT

(

YEAR

(

-------

(

(

(

(

(

(

(

(

e
.f.

"

I

*
(

(
(

(
_:"".

""--

YOIINr,

Y('lUNG

YOUNG

PER

PER
YEAR.

PER

AV!="PAGE

FA\.jN

I='F:~ALE

~F-~ALE

AT
BIPTH

AT
RlPTH
-----------

1963
.95
1964
•9,
1965
.9S
-1966
.93
1967
.88
1C) &amp;(~
.82
19b9
.fil
197()
_____________
.83
!971
.R4
1q72
.7(-,
1973
.•
74
1974
.82
1975
.6~
1976
.5~
FH7
.C,3
1978

(

---

Y_OUI\lt;

__
fF-~ALE

-----n.oo

E

D

(

,t

~LASSEc

== = = = = = = == == = = = = == =~:; = == == = = = =.:; ~.=== = = = = = =:: = = = = = = = = == = == = = = = = = _...__._===========~================================================

(

•

pc-p

AG~

0.00

.~()
•~0

- -

PER

AvERAGE
FFMALE

Ftf-1t1Lf __
AT

aT

6. T

pD!="_

DOST-

BIRTH

HUNT

======

.======~.

n.on
o.on

_(l •0 0______

.c:;q

.513

.58

.~fl

0.(\0

.57

.?,A

0.00

o .0(, __

.SS

.?O

.10

0.00

.52
.48

.nF.

0.00

.48

•l?

{I. 0 Q

.14

0.00
0.00

.51
.51
.43
.43

o • {HI
(l.on

o.on

1).00

.t;l
.40

G. 0 (\

(l.on

.35

(,.(1(\

o.on

.34

_

HUNT

--------- ---- -

-------

.61

.57
.61
.61
.137
.55

.51
.130
.53
.54
.46

.44
.52
.•
40

.36 __
.35

_

_

�-------======================
(

--

~=======================

-389-

T A 8 L E

9

=============
(

============================================================
============================================================
A

(

(

(

(

(

(

(

(

(

(

(

(

(

(

(

-.---.. _.--_.:: =

---

B

---

c

---

t~ALES
YOUNG
TOThL
(u(1t.
(AGE
YEt,p.
CLASS 2+) CU\2.~ 1)
ANI)
PER lon
PEo 1co
AOlH_T
~EMALEc;
~~AL F.s
F E ~~A Lf C;
(AGF::
(AGE
(AGF
YR CLASS
2+) CLASS
2+) CLASe; ~+)

=========
-- --------44.8

63
64
65
66
67
68
69
-70
71
72
73

74
75
76
77
78

49-:0
52.4
54.f,
57 ~l
59.6
62 ••
2
56 .•
Q
49.6
56~fl

63~5
54.7
63.5
71~7
77.3
81.'5

5Q.2
54-:-8
57.5
57.2

=========

1390.4
IAAO.2
1910.P
'2')97.4
55~-(f-· 2297.~
2'545.7
51.8
47.7
2703.A
4-]-:-8- 24A2.9
50.5
2193.6
50.9
2('10.6
2Q7q.4
43.2
2'3(\4.4
43.4
51.4
30S7.?
3739.9
39.6
3,;;;.2 41f,5.4
34.5
4433.3

D

F

---

---

TOTAL

T(lTAL
ADULT
FEMALES

YEAPLING
F Hill l. F. C;

F

---

- . _. -'----0-.-_- __ ·_,___ ._

-·--0-

._P.

•

.0

___

TnTAL
Y()lJ!lIG

(AGr

(AGE

(lIGF.

CLASe:; ~)

i.LAS~ 3 +)

CLIle:;S 1 )

=======:=

========

??t;7.0
?A16.n
?A::I9.2
~Q46.A
~()1=\4.8
-=!::-19.(\
~4(16.0
'3479.7
3';;;~9.7
":lA44.?
::l6P4.9
":1711.7
3946.9
4136.0
44~?3
4611.9

18::1C::;.2
__ _-1879.5
2097.5
'2196.L_. ---. ------.- - ..----. -22,1.2
'22}O.9
207_SO!J!___
....__________
._. ___
20BA.7
22~6.7
?34A _.8
..---~, _- _. --".,_.,._ .. 20~4.8
19p('.4
2475.0 _____
~OAA.6
1894.3
1877 ••
1
._.--_._., -- -.
--

========

843. )
795.0
8(17.8
R95.9 _
937.0
Qt:;1.7
941.9
~85.8
895.5
965.A

11)03.4

863.2
A6q.6
1()8~.4
905.5
829.3

.. _ ..

.. . ---p ..,

.

...

..

"

__

.

..

..

----.

�================================
============================~===

-390io

TABLE

:======:=======:=====
~=====~=====:=========

=============
. P('lS THlJ~ITI

Nr,-SE

A SON

MANAGr:/~r:tlJT

ST ~ T T S T r CS

================~===========================================
= = = = = = = =-~=::-~.= =- = = = == == = =~ = = = = = == = = = = = = = = ~ = = :: = = = = = = = = == ~
=::

A

B

C

===

::==

M~L~S

Y0UNG

===

TOTAL

(AGE

(~GE

YEAR.

CLASS

2+ 1

PERfo-n

FEMALES

CL~&lt;;S

II

-r6-n-----

PER
FEMALES

CAGE
Ct.0.S __ .
YR CLASS -i+-,- CLASS
2+l

AND
Af)UL T
~ALES

. (AGF
CLASS

2+)

D

r

F

TOTAL_

TOTAL
tlDtlL T
FE"MALt:S

T0TAL
Y(')llr-.IG

YFAPL H!G
Fr=:~AtlL ES

(AGF
( ~GE __
Cl.~&lt;;S 2) CLt.SS 3+l

_-- ::::-=::====== =======::.;= ========
---------- -----_
..

63
---64

65

,(

66
67
68
69
70
11

I(

73
'74

i(

72
75"

76
\ {

77
78

. (

33.5
38 •.
7
40.8
44.2
48 ••
1
Sl ••
8
4~_;-_Q
____
3~~9
45.1

53.5 __ .
44.4
55.7
64.6
72'.9
78.7
82.7

.....

6;' 5_---- 970.]
57.1
1;;:»09.A
60.9
1~22.6
6 .T ___ J ~!1? • :=;
57.3
17A}.2
5~.2
1949.2
5 0 .~. ____ t772. L
50.5
1444.7
5~.6
1.'126.5
54.1
21R7.Q
4A.2
lR~8.1
43.7
2443.'5
5;.8
2977.}
40~O------364-2.
Q
3r:;.5
41l4Q.?
34.9_:.
__ 4294.f-

n

=:: ':

==

( fI Gt:

CLASS

.

----------------- ==:=====

Q04.7 __ 20Q2.3
7J~.~
;:&gt;3~3.3
734.1
?S11.2

17,3.2
17ql.4
1975.6

H}n.A

2066 •.3 __
.__
2098 •.9

g7?O

R57."
gS4.1

_

II

__
._ ?5Q2.0

?7R9.9
~9"3.4
_ 2993.8
'~06.9

_

?077.0
.__19C;3•.3 ..__._.__.. _. .
= r s ,»
1974.R
844.8
1204.,
21?9.4
MA".9
32118.9
221 ? B _.____ ___.
~05.j:J 3211.7
19n3.6
~5"3.';
~S~4.7
1917.6
'159.9__
. 1749 •2_ _ 2389 •2.
11)71.4
39?3.7
1996.9
RQS.C;
42r:;2.6
18?A.7
'12(1.0_
437C5.C;
lAl2 •.1 _

_

_

__ ._

�==============================
(

.:============;========
T A 8 '-

============::
SU •..•
MARy
(

- -

PRII\JTOUT OF TRIAL

============================================================
============================================================
C

0-

===

=:-=

E
.---:----

TOT ~I_

(

TOT !I_
--Af\jII'-11.L
(

·"7--

TRIAL
---------

KILLEn
-------------

TOTAl_
VALUE

TOTAL
&gt;.1ALE.S
'&lt;ILLEO

==::==

======

-TROPHY

1

TRooHV

TOTAL.

~tlL~S

FFMflLFS

KILI_F'n

KILLEn
=====:=
~221.6

=====-=

(

(

TRIAL

=====

1

c

,
,

TOTllL
POUNns
ME:ilT
PROOUCED
---------------4450158.9

USr:D

U~En

o.

======

---------

--------

=======

-:16514.1

3n764.6

K

L

===
TOTAL
REFUGr::
BENf~TT

UNTTe;
------------.'107F+OR

AtJMC;

6750113.S

TOTAL
ANIMALS
DYING

PROf)UCEO

1

TOTI\L

TOTI\L

---------

==::==

TOTAL

---------ENI:PGY

J

TRIAL

TRIAl.

H

t

YOI'''II;

1

G

REFUGE
RENEFIT
UNTTS
(FRO"'1
SALfS)

M

===
REF '-'t:; f.
Bt:NEF IT

UNTTS
(F ROI&gt;l
IJC;F:)

-------

--------------

.9G1F+OA

.491F+(l8

N

--REFUGf

RE'JE~rT
UNITS
(FPOM

STOrK)

=======

.993F+(I4

------- .

. -_.-

-

.

��-393-

PART II

EVALUATION OF INTERAGENCY BROWSE
UTILIZATION AND PELLET GROUP COUNT METHODOLOGY

INTRODUCTION

For several years the Colorado Division of Wildlife has surveyed deer
and elk ranges in Colorado cooperatively with the federal land management
agencies, primarily the U. S. Forest Service and the Bureau of Land Management. The methodology of these surveys has been detailed previously (Denney
1962). The stated intent of the system is to measure changes in carrying
capacity of big game winter ranges over time. Generally, the methods employed
consist of measuring or ocularly estimating utilization (expressed as percentages of current annual growth) of selected "key" shrub species on selected "key" winter range sites. The term "key" implies that both selected
shrub species and selected winter range areas are of primary importance in
maintaining the particular big game populations utilizing them.
Several deficiencies in this approach to estimation of carrying capacity
seem obvious. First the selection of key areas for sampling predestines
biased information.
The patterns of animal distribution throughout large
expanses of so-called winter range are complex and dynamic (Gilbert et al.
1972; Lynch 1974; Bergerud 1974) making the term "key area" ambiguous if not
meaningless.
For instance in Middle Park, Colorado snow depths appeared to
regulate duration, intensity, and patterns of winter range use by mule deer.
Some areas which were used heavily and for prolonged periods during deep snow
winters were practically unused in the winters of shallow snow accumulation
(Gilbert et al. 1972). Those areas which were subjected to heavy, prolonged
use in deep snow winters would have been classed as "key areas" according to
the Interagenc.y Range Evaluation system and, thus, would have been selected
for sampling while those areas that were most intensively used in light snow
winters would not have been sampled at all.
Conversely, in heavy snow winters deer would concentrate on key areas with
resultant evidence of overutilization suggesting ranges were overstocked.
In both situations the conclusions would likely be erroneous for several reasons. First, little is known of the contributions of various zones of the
winter range in maintaining deer in winter.
It is far from certain that "key
areas" are any more important to the winter survival of deer t han areas used
earlier in the winter. Consequently, sampling procedures which measure
selected attributes of key areas only do not reflect the contributions of the
total winter range complex. Secondly, the interagency method probably does
not even reflect the contributions of the key areas. This is because both
sampling methodology and intensity are deficient.
Also since only selected
species of browse are measured, the method ignores the contributions of unmeasured browse species and herbaceous forbs and grasses.

�-394-

The previous discussion suggests that interactions between big game
animals and their winter ranges are complex and poorly understood.
To
assume that a simple, easy, quick system can be developed to predict
directions and magnitude of these interactions is unwise. Any scheme for
estimating carrying capacity of big game on winter ranges must be based
upon: a thorough knowledge of the vegetation species that contribute to
the total diet of big game animals, an understanding of the nature of their
individual and collective contributions, insights into factors that effect
changes in the relative contributions of each forage species (snow, temporal
changes in palatability and digestibility, big game range use patterns, etc.),
and nutrient requirements of each big game species.
It is not surprising that tests of the efficacy of the interagency system of
big game range evaluation or similar systems have been negative (Bartmann
and McKean 1969; Mackie 1975). A discussion of an empirical evaluation of
the system as used on the Uncompahgre Plateau and the Upper White River
drainage follows, but again the results are not encouraging.

METHODS
Procedures for obtaining browse utilization and pellet group count information evaluated in this report are described elsewhere (USDA Forest Service
1968). Despite official acceptance and application of the techniques, individuals and factions of all three agencies have intermittently been critical
of the program.
Some minor changes have been effected sporadically, but
methodology has remained essentially the same since about 1960.
As recent as 1968 (Bartmann, Kufeld), formal proposals for research and
development of range methodology have been made. Those proposals never
materialized, probably because estimated time needs ranged from 4 to 10
years. Thus, notwithstanding the obviously deficient time available for more
thorough study here, task force and supervisory staff felt that an attempt
should be made now to reawaken interest in the problem. Within the short
1-1/2-year span remaining for operation, it was deemed infeasible to accomplish
new field work. So, the next most logical alternative was to evaluate existing
data.
First in order was determination of how much, where, and from whom information
would be solicited.
Previously mentioned time limitations pegged data needs
small, which also meant that, at most, only a few areas of the state could be
represented.
Coincidental with quantity aspects of data selection was also who might have
the best quality, especially over time. Quality through time was critical
because of needs for determining existence of valid trends in forage utilization and deer and elk stocking indexes from pellet group counts. Picking
satisfactory data sources very obviously depended upon subjective judgment,
undesirable but necessary for doing this type of study.
In addition to finding usable browse transect data, perhaps next most important was the need for areas also having deer and elk population data. That
meant using information based primarily on aerial counts for trend or sex
and age tallies fundamental to change-in-ratio total estimations.

�-395-

Excepting the Middle Park Deer Management Study Area, which is under
separate intensive populations investigations, foregoing population
information needs immediately reduced the choice to two usable areas.
Of the two, one area is the upper White River Basin. The other area is
the Uncompahgre Plateau.

White River Area
The upper White River country in the Division Northwest Region is a locale
of much past and continuing study and management effort regarding elk populations and habitat. Complicating evaluation of animal and forage use relations here, however, is the presence of substantial deer numbers in joint
tenancy with elk. This dual range OCcupancy is yet more the rule than the
exception in Colorado. Thus, examination of White River area data was felt
to be advantageously representative of statewide conditions. And, wherever
analyses demanded, compensating allowances were possible to accommodate for
the dual range use happening.
I began compiling range data in March 1974 by exam1n1ng records filed at
the White River National Forest Supervisor's office in Glenwood Springs.
Information was available there for territory corresponding with Big Game
Management Units 25, 26, 33, and 34.
The White River National Forest Blanco Ranger District office in Meeker
was next visited for Units 23 and 24 data. Routt National Forest Bears
Ears and Yampa Districts' offices, respectively, in Craig and Yampa, yielded
data for Units 12, 13, and 26. Except for supplementary 1974 data for Units
23 and 24 obtained in November 1974 from Blanco District Ranger Dale Harthan
and Range Conservationist Dan Sommers, the Craig and Yampa offices data
compilation completed transfer of needed big game range records. The
eight units mentioned in these paragraphs comprise elk Data Analysis Unit
(DAU) E6.
Time span of DAU E6 records study was arbitrarily chosen to be 10 years,
extending from 1964 through 1973 (excepting 1974 Units' 23 and 24 data
previously mentioned). As will be seen later, there was little consistency
in transect sample sizes, particularly before 1969.
Uncompahgre Area
The Uncompahgre Plateau in the Division Southwest Region has long been a
very important deer area. In spite of recent noticeable gains in elk population numbers, dual winter range use by deer and elk apparently is still
comparatively limited.
Uncompahgre Big Game Management Units 61 and 62 range records were essentially
complete in Regional Biologist Hal Burdick's files, with data gaps easily
filled by goi11g nearby to the Montrose BLM District office. Units 61 and 62
comprise deer Data Analysis Unit (DAU) D19 (or elk DAU E20).

�-396-

RESULTS AND DISCUSSION
Numbers of transects read by big game unit by year are presented in Tables
1 and 2. The tables also combine unit data into data analysis unit totals,
respectively, for DAU E6 (White River elk) and DAU D19 (Uncompahgre deer).
The purpose of presenting sample size information is that information was
pertinent to initial discussions of approaches to data analyses with
Statistician Dave Bowden and task force coworkers Ray Boyd and Tom Pojar.
In addition to lacking confidence because experimental design was not evident in the big game range methodology, Dr. Bowden was critical of the
extreme variability in sample numbers from year to year. Data Analysis
Unit E6 samples were particularly poor in that respect. Excessive sample
size variation, added to lack of experimental design, inadequately trained
and supervised personnel, and inherent transecting weaknesses were very
discouraging factors from the outset. Besides, I had participated in annual
reading of transects with Southeast and Southwest Region field personnel
(Boeker and Baker 1960; Denney 1961, 1962) for several seasons and observed
methodology diversions that hindered making and applying valid management
conclusions for which I understood the techniques were intended.
Strong initial discouragement notwithstanding, data were subjected to
statistical tests by assuming, though questionable, ordinary requirements
of acceptable scientific inquiry had been met. Consequently then, specific
findings from testing for the two areas follow, with White River data being
treated first.

White River Area
A regression of White River elk DAU E6 aerial elk trend (Y) versus mean elk
days per acre (X) for the ten years of 1964-1973 (Tables 3,4) was an attempt
to evaluate these estimators of true elk population trend. There was no
significant relationship (P&gt;.20) between the two estimators (r = .381, 8
df). So, one of the estimators could be good while the other is poor, or
neither estimator is good (Pojar pers. corom.).
In a second approach, antlered elk harvest (excluding Units' 23 and 24 data)
(Y) versus mean elk days per acre (X) were tested. Resulting was r = .636,
which is not significant (P &gt;.05, 7 df).
Although not based upon nor involving range transect data, another regression
was computed (Pojar pers. corom.) from information shown in Table 3. That
test compared DAU E6 aerial elk trend (Y) versus antlered elk harvest (X)
for years 1964 through 1973 (also excluding Units' 23 and 24 antlered harvest
data). A resulting r = .595 was not significant (P&gt; .05, 7 df). Units' 23
and 24 data were excluded from this and previously mentioned tests because
bull licenses were issued in restricted numbers through "specified elk season"
procedures. Unrestricted hunter selectivity on male elk thus was not effected,
necessitating use of data only from the six adjoining management units where
free-choice antlered elk hunting was allowed.
Estimators of DAU E6 true elk population trend that were tested and had results
presented in preceding paragraphs are of questionable value. Techniques

�-397-

inadequacies already discussed, plus lack of continuity of personnel
and virtual nonexistence of quality control across too great an area and
over several agency administrative subunits were probable factors acting
to prevent any success in trying to find reliable indexes to elk population numbe rs .
Not yet willing to concede to desperation from results given in previous
paragraphs, I decided to attempt analysis of some selected data from a
subunit of DAU E6. Aim of that effort was to determine if a smaller management entity would improve statistical reliability, holding to the 1968-74
time span. Therefore, by subjective selection involving prime factors of
demonstrated interest over time by the same personnel and aerial trends
applicable to a single unit, I went to Management Unit 23 for additional
data.
In keeping with attempts to relate elk days per acre indexes with aerial
trend data (Table 3), a regression was computed for aerial 'trend (Y) versus
elk days per acre (X) for period 1968-74. The correlation coefficient was
not significant (P&gt; .05; r = .578, 5 df).
To this point, cursory examination of browse utilization portions of transect records indicated so much variability in species composition and
utilization that statistical computations seemed inappropriate.
For example,
relative to browse composition variation, 602 transects read from 1965 to
1973 showed species occurrence on transects as follows: oakbrush (Quercus
gambelii) 72%; serviceberry (Amelanchier spp.) 69%; big sagebrush (Artemisia
tridentata) 67%; true mountain mahogany (Cercocarpus montanus) 47%; bitterbrush (Purshia tridentata) 37%; and rabbitbrushes (Chrysothamnus spp.) 19%.
At least 15 other species, ranging from 0.2 to 9.1 percent frequency of
occurrence, were also recorded on those DAD E6 transects. Since, from the
foregoing analysis, oakbrush and serviceberry are the most abundant and also
happen to be high value forage species for elk (Kufeld 1973), I arbitrarily
chose them for statistical testing.
So, a regression analyses was done on data hopefully selected to be most
accurate, specifically 25 transects read in Unit 23 in 1971 (Table 5). A
correlation coefficient for mean oakbrush utilization (Y) versus mean elk
days per acre (X) was found to equal .624 which is significant (P&lt;.Ol, 23
df). This "r" value is, however, too weak for predictive purposes (Y = 3.73
+ .42 X; SE = 7.31, n = 25). A similar result came from comparison of mean
serviceberry utilization (Y) versus mean elk days per acre (X). Irrespective
of this species occurring on fewerA..transects (22 of 25) in 1971, significance
at .05 resulted (r = .497, 20 df; Y = 5.98 + .47 X; SE = 11.65, n = 22) but
not .01.
Elsewhere in Colorado, Anderson et a1. (1972b) found correlation coefficients
not significant (P&gt; .05) for mean utilization of true mountain mahogany and
bitterbrush (Y) when compared with mean deer nwnbers (X) based upon pellet
group counts. That investigation had stringent experimental control, something
very lacking in field work basic to this study.
It should be noted that in statistical comparisons done so far involving elk
days per acre use indexes, concurrent deer stocking was not accounted for nor
was it necessary. However, in comparing utilization with elk days per acre

�-398-

indexes in the preceding paragraph, it was pertinent that total browsing
pressure be included. Consequently, on 10 of 25 transects where deer
pellet groups were present, values for mean elk days per acre were adjusted
upwards by a factor of 0.286. The latter was computed from American Society
of Range Management (1964) animal unit conversion factors of 0.7 for elk
and 0.2 for deer (0.2 ~ 0.7 = 0.286).
Additional comparisons were made in efforts to learn if mean elk days per
acre values would be usable evaluators of elk trend between years. Results
of those analyses are presented in Table 6 and discussed as follows:
1. In two out of three sets of year comparisons (1971 and 1972 vs.
1973), upward trends of mean elk days per acre agreed fairly closely with
computer simulation and aerial trends for E6.
2. Comparisons of elk days per acre means for 1970 versus 1973 for E6
and Unit 23 yielded inconclusively contradictory results. Empirically, population simulation and aerial count changes would seem to be more reliable
indicators of trend in this instance.
3. Unit 23 mean elk days per acre data comparison for 1972 versus 1973
show significant change which is not what is indicated by the corresponding
aerial count. Explanation for this discrepancy could be that elk population
gains for E6 occurred on portions exclusive of Unit 23, and/or Unit 23 pellet
group data are out-of-line through sample error or other fault.
4. Other conclusions seem possible from Table 6 data; but, without a
highly reliable population yardstick upon which to depend, and none is yet
available, about anything deducted from present populations technology must
be considered as pure speculation.

Uncompahgre Area
In 1967, Manager C. E. Till and Game Biologist Errol Ryland of the Division
Southwest Region started a program of increased browse utilization and associated
pellet group count transecting. District Wildlife Conservation Officers were
made responsible for reading 30 transects each in their districts, the number
having arbitrarily been determined to fit average needs for all Southwest Region
district situations. Prior to 1967, available data were so meager and records
so scattered that I began data compilation with the 1967 upsurge in activity
and continued on through with 1974 spring field season information. Results
presented in following paragraphs therefore pertain to the eight-year span
from 1967 to 1974.
Preliminary to exploring Uncompahgre data with statistical computations, I
calculated and recorded (Table 7) deer pellet group statistics for Units 61
and 62 (DAU D19). Ignored were elk pellet group count statistics. As mentioned
previously, Uncompahgre Plateau elk numbers were generally so few within the
period of 1967-1974 that only sporadic and sparse pellet groups occurred.
For example, five of 80 transects (6%) in 1967 had elk groups on them. Percent frequency of occurrence of elk pellet groups for the other seven years
varied from 4% (1968) to 22% (1974), showing an overall frequency of about 12%
(64 of 553 transects). Perhaps as important as frequency of occurrence was

�-399-

intensity of occurrence, which is exemplified by the average low stocking
index of 3.5 elk days per acre (range : 1-15 ED/A) for those 64 transects
having elk pellet groups present.
Following the order of statistical comparison testing used previously for
White River data, a regression of mean deer days per acre (Y) versus aerial
trend (X) for 1967-1974 was computed for DAU D19 (data from Table 8). Again,
purpose of this mathematics was to help evaluate the population numbers
indexes as estimators of true deer population trend. Correlation was not
significant (P &gt;.05) of the two estimators (r = -0.206).
Much the same result was found when a Unit 62 mean deer days per acre (Y)
versus aerial trend (X) regression showed no significance (P&gt;.05; r = -0.377).
Perhaps of greatest interest in both of these results is not as much the weak
relationship but the negative correlation of the two parameters.
Inherent
deficiencies in methodology could account for the weakness in cor~elation,
but winter conditions causing deer to concentrate and help provide for high
aerial trend values should also help yield higher mean pellet group counts.
It is not easy to understand how that situation could be otherwise.
Next, antlered deer harvest (Y) versus mean deer days per acre (X) regressions
were done for DAU D19 and Unit 62 for the 7-year span of 1967-1973 (data from
Table 8). Both resulting correlation coefficients were not significant
(P&gt;.05, 5 df), with r = .744 and r = .715, respectively, for DAU D19 and Unit
62. The "r" values show strong positive relationship, however, with harvest
logically dependent upon preceding spring populations as indicated by pellet
group counts. These interactions could become more important and providers of
usable indexes to population yields with additional years of foundation data.
Information given i~ Table 8 regarding antlered deer harvests and aerial
trends, 1967-73, for DAU D19 and Unit 62 were tested to determine correlation.
First, for DAU D19, antlered harvest (Y) versus aerial trend (X) was significant (P&lt; .01, 5 df) with r
-.883 and Y = 7,041.10 - .67 X; SE = 380.50,
n = 7. With Unit 62 furnishing nearly two-thirds of the total DAU harvested
animals, it was not too surprising that results of a regression similar to the
immediately preceding one would yield close to the same results. Thus, for
Unit 62, antlered harvest (Y) versus aerial trend (X) was also significant
(P&lt;.Ol, 5 df), r = -.810 and'?'= 3,531.40 -.34X; SE = 250.80, n = 7.
The strongly negative correlation of antlered harvests and aerial trends presented in the previous paragraph are paradoxical inasmuch as one might expect
greater harvests following higher post-hunt aerial counts of the previous
winter. Tom Pojar (pers. comm.) suggested a plausible explanation by reasoning
that good aerial counting conditions (and thus, high counts) happen generally
in rough winters. The latter also cause greater animal losses through starvation and lowered subsequent production via stress on does. The higher winter
mortality and lower production would be reflected immediately by lower harvests
the following fall, everything else being equal. Again, despite high inverse
correlations, "r" values are not high enough to allow data use in prediction
equations.
Browse utilization comparison compuations with pellet group count indexes
involved only big sagebrush. This species overwhelmingly dominated as the one
key species, occurring on 94% (523) of a total of 557 transects. Other shrubs

�-400-

such as black sagebrush (~. nova), serviceberry, true mountain mahogany,
and oakbrush variously occurred on between IS to 20 percent of the transects which would largely furnish inadequate sample sizes for analyses.
Percent frequency of several other shrubs that equalled &lt; 1% to about 10%
included rabbitbrushes and fourwing saltbush (Atriplex canescens), both
about 8%.
Despite a fairly high correlation coefficient (r = .617) between percent
big sagebrush utilization (Y) and mean deer days per acre (X) (Table 9),
it was not significant (P&gt;.OS, 6 df). Considering difficulties involving
accurate estimations of utilization on this evergreen, early starting
shrub, apparent shortcut sampling procedures applied in making estimations,
and overall light mean utilization values obtained (all &lt; 27%),' this utilization phase seems to also yield irrelevant management information.
DAU D19 deer populations for years 1973 and 1974 were compared by three
different estimators (Table 10). The mean deer days per acre index showed
a significant (P&lt;.Ol) drop in deer stocking from 1973 to 1974, whereas
the computer model and aerial trends showed nearly static conditions (Unit
61 excepted). In addition, when seven years (1966-73) of data are pooled
for comparing with 1974 information (Table 11), mean deer days per acre was
significantly different (P &lt;.OS, but no t K .01; tSS2 df = 1.989&gt;1.96
.OS).
tOJ ,
Since, like the White River area elk population, there is no way of knowing
what actually is happening as Uncompahgre deer populations change seasonally,
one estimator technique appears now as good or bad as another.

CONCLUSIONS
1. Sampling and technique deficiencies in the interagency methodology,
as now performed, appear to preclude use of pellet group counts and browse
utilization estimates as valid indicators of population or utilization trends,
respectively. Mean utilization of primary (key) browse for both areas was
too light to cause general concern for their continued existence and well
being.
2. Lack of replication seriously detracts from aerial trend counting.
This indicator was not a principal target of the present study but demanded
attention because it has been and is still given much credibility in local
management considerations.
3. Antlered harvest of elk for the White River correlated positively
with pellet group count stocking indexes, although not significantly.
Uncompahgre antlered deer harvests also showed strong, though not significant,
positive relationship with mean deer pellet group counts. As these relationships now stand, however, much greater preclsl0n is needed before mean pellet
group indexes might be used to predict harvest.
4. Computer simulations appear to yield population data that, though
not easily refutable, nevertheless lack statistical sideboards and thus are
just as suspect as other currently applied indicators.

�-401-

RECOMMENDATIONS
1. Discontinue browse utilization estimations except where utilization on key shrub species occurs at intensities thought detrimental to
continued welfare of the forage species. Those occasions when excessive
utilization is believed to be evident must be verified through interagency
concurrence on the ground.
2. Pending possible further review of the procedures, accept other
inadequacies as they now exist and continue the pellet group count part
of instructions in at least present minimum intensity or solicit advice
for increased sampling from qualified personnel of anyone of the three
principal cooperating agencies.
3. Reinstate a Division-BLM-U. S. Forest Service panel to implement
a joint, three-agency, all-out research effort for review of big game range
analysis techniques in their entirety. Further, it is strongly recommended
that panel members, as well as researchers who are assigned to possible
subsequent investigations projects, never have been associated with the
fabrication of original methodology and/or subsequent revisions.
4. The widely accepted and applied pellet group count methodology
(Hart 1958, Smith et al. 1969, Anderson et al. 1972a,b) warrants further
attempts at large-scale application.
That application should concern at
least one Colorado big game management unit, or preferably more, either
singly, or in combination equaling a data analysis unit. Anderson (pers.
comm.) made similar recommendation in the past. This study proposal
would logically be integrated with any research program resulting from
action on preceding. item 3.

LITERATURE CITED
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1972a. Mule deer fecal
group counts related to site factors on winter range. J. Range Manage.
25:66-68.
1972b. Mule deer numbers and shrub yield-utilization
range. J. Wildl. Manage. 36:571-578.

on winter

Bartmann, R. M. 1968. Proposed research study: evaluation of big game range
analysis techniques. Colo. Dept. Game, Fish and Parks. April 4. lp.
(resubmitted in 1969).
Bartmann, R. M., and W. T. McKean. 1969. Paddock studies on effects of varying
intensities of deer use. Colo. Game, Fish and Parks Div. Game Res. Rep.,
Fed. Aid Proj. W-lOl-R, Job 4. July, Part 3. pp. 305-393.
Bergerud, A. T. 1974. The role of the environment in the aggregation movement, and disturbance behavior of caribou. pp. 552-584. In V. Geist,
and F. Walther (editors). The behaviour of ungulates and itS relation
to management.
Vol. 2. International Union for Conserv. of Nature and
Nat. Res. ICUN Publ. 24. Morges, Switzerland.
512-940 p.

�-402-

Boeker, H. M., and B. D. Baker. 1960. Browse transect analysis and application. Colo. Dept. Game and Fish. Fed. Aid Quart. Rep. (October),
Proj. W-lOl-R-2, WP-3, J-3, p. 35-36.
Denney, R. N. 1961. Transect analysis and application.
Colo. Dept. Game
and Fish, Fed. Aid Quart. Rep. (April), Proj. W-lOl-R-3, WP-3, J-3,
p. 37-47.
1962.
and Fish.
p. 51-96.

Browse transect analysis and application.
Colo. Dept. Game
Fed. Aid Quart. Rep. (April), Proj. W-lOl-R-4, WP-3, J-3,

Gilbert, P. F., O. C. Wallmo, and R. B. Gill. 1970. Effect of snow depth
on mule deer in Middle Park, Colorado. J. Wildl. Manage. 34(1):15-23.
Hart, R. D. 1958.
South Dakota.

Evaluation of deer pellet group census in the Black Hills,
M.S. Thesis, Colo. State Univ. 66p.

Kufeld, R. C. 1968. Proposed research study: development of improved deer
and elk census and range survey techniques. Colo. Dept. Game, Fish and
Parks. April 1. lp. (resubmitted in 1969).
1973.

Foods eaten by Rocky Mountain elk.

J. Range Manage.

26:106-

113.
Lynch, J. J. 1974. Merino sheep: some factors affecting their distribution
in very large pastures.
697-707. In V. Geist, and F. Walther (editors).
The behaviour of ungulates and its relation to management. Vol. 2.
International Union for Conserv. of Nature and Nat. Res. ICUN Publ.
24. Morges, Switzerland.
512-940 p.
Mackie, R. J. 1975. Evaluation of the key browse survey method.
Workshop, Proc. 5:28-33.

Mule Deer

Range Term Glossary Committee.
1964. A glossary of terms used in range
management.
D. L. Huss, Comm. Chairman. Amer. Soc. of Range Manage.
32p.
Smith, R. H., D. J. Neff, and C. Y. McCulloch.
1969. A model for the installation and use of a deer pellet group survey. Spec. Rep. No.1,
Ariz. Game and Fish Dept. 30p.
USDA Forest Service. 1968. Chapter 80 - Big game range analysis.
In
Range Environmental Analysis Handbook, Forest Service Handbook 2209.21,
Rocky Mtn. Region (Region 2), November. pp. 00-88.12.

Prepared by

---r--

!3e/lcdva'in!.

jJ

-'?,
I

Q1j

f

j)¥ 1&lt;'

Bertram D. Baker
Asst. Wildlife Researcher

�Table 1.

Annual big game range transecting

* - Data Analysis Unit E6 (White River) - 1963-1973.
Number of Transects

Big Game
Mgmt.
Unit

1963-64

1964-65

1965-66

1966-67

1967-68

1968-69

1969-70

1970-71

1971-72

1972-73

Totals
1963-1973

12

0

7

5

9

13

23

0

10

12

4

83

13

0

3

1

0

5

13

1

3

3

0

29

23

0

3

2

5

5

22

20

27

14

18

116

24

3

9

8

14

17

9

13

9

11

6

99

25

19

10

0

0

0

30

6

36

59

24

184

26

1

2

2

0

0

1

0

8

5

3

22
I

.t'

33

4

6

5

6

3

8

2

8

7

58

9

0

w
I

34

2

3

0

0

0

6

0

7

11

11

40

DAU
Total

29

43

23

34

43

112

42

108

122

75

631

*Includes Production-Utilization

(P-U) transects where current annual growth is measured pre- and post-use on
shrubs and extensive transects that give shrub species utilization estimates post-use in spring. Pellet group
counts were also done in conjunction with both types of spring shrub-utilization transecting.

�Table 2. Big game annual range transecting
1966-1974.

- Data Analysis Unit D19 (Uncompahgre)

or E20 (Uncompahgre)

_

Big Game
Mgmt.
Unit

1966-67

1967-68

1968-69

1969-70

1970-71

1971-72

1972-73

1973-74

Totals
1967-1974

61

24

26

33

20

27

20

20

20

190

62

56

58

55

42

39

48

37

32

367

DAU
Total

80

84

88

62

66

68

57

52

557

Number of Transects

I

.j:'-

g
I

�Table 3.

Elk Population

indexes - White River elk DAU E6 (1964-73) and Big Game Management

Elk DAU E6

Unit 23 (1968-74).

Big Game Unit 23
Mean
Aerial
Elk Days
Trend
Per Acre

Year

Mean
Elk Days
Per Acre

Aerial
Trend

Antlered
Harvest

Antlered
Harvest
Exc1. 23 &amp; 24

1964

8.38

3863

2501

1318

1965

16.77

5253

1993

1136

1966

13.78

6297

1873

1097

1967

19.94

6055

1737

849

1968

21. 21

5700

2184

1257

21145

29.20

1156

1969

20. 70

6371

2262

1424

20722

42.23

2037

Model
Pre-Calving
Total

I

~

0

20733

45.70

1996

21113

31. 70

1859

1203

23634

32.71

3168

3075

2318

25261

54.72

3156

-

-

24945

48.00

3270

1970

37.69

5560

2517

1971

19.52

7430

1039

1972

16.00

8339

1770

1973

31.00

10149

1974

-

-

*Figures not used, - spike elk protected.

1651

*

686

*

VI
I

�-406-

Table 4.

White River elk DAU E6 pellet group transecting
Elk Da~s Per Acre

Year

n

X

-6

- 1964-1973.

Deer Da~s Per Acre

-6-

X

X

-6

-6_

X

1964

29

8.4

9.4

1. 75

36.2

30.0

5.76

1965

43

16.8

19.0

2.90

23.5

25.8

3.90

1966

23

13.8

20.9

4.36

14.5

21.0

4.37

1967

34

19.9

19.9

3.41

13.9

21.8

3.74

1968

43

21.2

18.7

2.85

8.9

14.7

2.24

1969

112

20.7

22.0

2.07

13.9

19.3

1.82

1970

42

37.7

21. 9

3.38

5.6

11.2

1. 74

1971

108

19.5

15.5

1.49

11.4

15.6

1.50

1972

122

16.0

15.1

1.37

13.9

16.0

1.45

1973

75

30.9

26.6

3.07

21.6

30.6

3.53

1974*
(Unit 23)

21

48.0

20.6

4.50

3.24

5.3

1.15

1974*
(Unit 24)

13

58.9

20.3

5.62

1.62

2.9

0.60

*

Data compiled in 1974 only for Big Game Units 23 and 24, - included here
for convenience.

�-407Table 5. Elk days per acre stocking and oakbrush utilization - Spring
1971 - from 25 extensive transects in Big Game Management Unit No. 23
(Miller Creek).
Deer Days
Per Acre

Elk Days
Per Acre

Adjusted
Elk Days Per Acre *

Ave. % Utiliza.
Oakbrush

2.00

42.00

42.57

32.00

5.00

57.00

58.43

28.00

10.00

54.00

56.86

36.00

0.00

28.00

28.00

14.00

0.00

18.00

18.00

13.00

0.00

38.00

38.00

14.00

1.00

19.00

19.28

6.00

0.00

30.00

30.00

18.00

0.00

28.00

28.00

9.00

0.00

18.00

18.00

13.00

0.00

38.00

38.00

28.00

0.00

28.00

28.00

9.00

0.00

42.00

42.00

19".00

0.00

44.00

44.00

11.00

2.00

31.00

31.57

30.00

10.00

62.00

64.86

30.00

1.00

40.00

40.28

11.00

15.00

l3.00

17.28

30.00

6.00

33.00

34.71

20.00

0.00

18.00

18.00

15.00

2.00

26.00

26.57

12.00

0.00

31.00

31.00

12.00

0.00

15.00

15.00

3.00

0.00

11.00

11.00

7.00

0.00

28.00

28.00

11.00

X = 32.296

X = 17.240

*Obtained by mUltiplying deer days per acre by 0.286 (elk equivalent) and
adding result to value for elk days per acre.
,

�Table 6.
Comparison of three methods of estimating elk population trends - DAU E6 (White River) and Big Game
Management Unit 23 (Miller Creek) - years 1970, 1971, and 1972 versus 1973.

Mean Elk Dazs Per Acre
1970
1973
Percent Change

Est. Total Elk Population
ComE' Model at Preca1ving
Percent
1970
1973
Change

1970

1973

Percent
Change

DAU E6

37.7

30.9

- 18

20733

25261

+ 18

5560

10149

+ 45

BGMU 23

45.7

54.7

+ 16

-

-

-

1996

3156

+ 37

1971

1973

1971

1973

Percent
Change

1971

1973

Percent
Change

Management
Entity

N.S. (P&gt; 0.10;
t
= 1. 36)
115
N.S. (P&gt; 0.20;
t
= 1. 29)
36

Percent

Change

Elk Aerial Trend

I
.p-

o

(X)

DAU E6

BGMU 23

DAU E6

BGMU 23

30.9

+ 37

Sig. (P&lt; 0.001;
t181 = 3.87)

21113

25261

+ 16

7430

10149

+ 28

31. 7

54.7

+ 42

Sig. (P&lt; 0.001;
t
= 4.51)
43

-

-

-

1859

3156

+ 41

Change

1972

1973

Percent
Change

1972

1973

Percent
Change

N.S.* (p&gt; 0.05;
t
195 = 1. 82)

23634

25261

+ 6

8339

10149

+ 18

Sig. (P&lt;0.05;
t
= 3.21)
30

-

-

-

3168

3156

-0.4

1972

1973

16.0

30.9

+ 48

32.7

54.7

+ 40

*
Significant

I

19.5

(P &lt; 0.10).

Percent

�-409-

Table 7.
1974.

Uncompahgre deer DAU D19 pellet group transect data summary, 1967-

Deer Pellet Groups/Acre
s
X

Management
Entity

n

1967

Unit 61
Unit 62
DAU Dl9

24
56
80

13.71
16.41
15.60

9.32
14.09
12.85

1.90
1.88
1.44

1968

Unit 61
Unit 62
DAU Dl9

26
56
82

13.62
25.32
21.61

12.10
22.14
20.20

2.37
2.96
2.23

1969

Unit 61
Unit 62
DAU Dl9

33
54
87

16.48
26.17
22.49

8.80
22.27
18.89

1.53
3.03
2.03

1970

Unit 61
Unit 62
DAU Dl9

20
42
62

9.95
18.95
16.05

6.89
11.59
11.09

1.54
1. 79
1.41

1971

Unit 61
Unit 62
DAU D19

27
39
66

12.81
22.28
18.41

7.16
15.37
13.44

1.38
2.46
1.65

1972

Unit 61
Unit 62
DAU Dl9

20
48
68

10.55
31.44
25.29

7.40
17.36
17.86

1.66
2.51
2.17

1973

Unit 61
Unit 62
DAU Dl9

20
37
57

10.90
47.08
34.39

10.40
32.43
31.88

2.33
5.33
4.22

1974

Unit 61
Unit 62
DAU Dl9

20
31
51

9.60
20.23
16.06

10.25
11.95
12.37

2.29
2.15
1. 73

Year

sX

�Table 8. Deer population indexes - Uncompahgre
and Unit 62 (East Side Uncompahgre).

Deer DAU D19 - Big Game Management

Unit 61 (West Side Uncompahgre)

Ae rial Trend
Year

Mean Deer Days/Acre
Unit 62
DAU Dl9

1967

16.41

1968

Antlered Harvest
Unit 62
DAU Dl9

Total
Harvest
DAU Dl9

DAU Model
Pre-fawning
Total

Unit 61*

Unit 62

DAU Dl9
(Total)

15.60

932

5,391

6,323

1,553

2,072

3,562

19,158

25.32

21.61

1,250

6,960

8,210

1,060

1,562

2,870

19,770

1969

26.17

22.49

749

6,120

6,869

1,986

2,955

5,395

21,285

1970

18.95

16.05

1,058

5,280

6,338

1,603

2,697

3,097

19,897

1971

22.28

18.41

1,411

5,168

6,579

1,615

2,661

2,661**

21,247

1972

31. 44

25.29

939

5,474

1,698

2,892

3,202

23,561

2,456

4,165

5,824

25,611

47.08

34.39

1,006

3,560

4,566

1974

20.23

16.06

447

3,833

4,280

583

4,050

4,633

1975

*

For Uravan portion.

** Statewide

antlered only season.

t-'

6,413

1973

I

+:-

26,799

0
I

�-411-

Table 9. Deer days per acre stocking and big sagebrush utilization,
Deer DAU D19 (Uncompahgre) - 1967-74.
1967

1968

1969

1970

1971

1972

1973

1974

Mean Deer Days/A.

15.60

21.61

22.49

16.05

18.41

25.29

34.39

16.06

Mean Percent
Utilization

25.65

21.00

19.82

14.10

17.65

26.11

26.96

19.25

�Table 10. Comparison of three methods of estimating deer population trends - DAU D19 (Uncompahgre)
Big Game Management Units 61 and 62 (West and East Sides Uncompahgre) 1973 versus 1974.

Management
Entity

Mean Deer Dals Per Acre
1973
1974
Percent Change

DAU D19

34.39

BGMU 62

16.06

47.08

20.23

10.90

9.60

-53

Sig. (P&lt;O.01;
t106 = 3.71)

-57

t

Sig. (p&lt; 0.001;
= 4.37)
66
N.S.

BGMU 61

-12

t
38

Est. Total Deer Population
ComE. Model at Pre-Fawnins
1973
1974
% Change

and

Deer Aerial Trend
1973
1974
% Change

25611

26799

+ 4

4566

4280

-

6

-

-

-

3560

3833

+

7

-

-

-

1006

447

- 56

(p&gt; 0.01;

= 0.40)

I

+='
I-'
tv
I

Table 11. Deer pellet group count index statistics
(Uncompahgre).
Year (Period)
1966-73
1974

for comparing

7-year span of 1966-73 versus 1974 _ DAU 19

n

Mean Deer Days/Acre

S. D.

502

21.65

19.45

51

16.06

12.37

�-4l3-

July 1975
JOB PROGRESS REPORT

State 0 f __

--=CO=L:.::O=:RADO=-=-_

Project No.__
Work Plan No.

~W~-~3~8~-~R~-~2~9 _
~1~9

_

Deer-Elk Investigations
Job No.

1
-------=--------------

Job Title __ ~E~v~al~u~a~t=i~o~n~o~f~R~a~d~l~·o~=T~e=l~e=m~e~t~r~yL_
Period Covered:
Personnel:

_

April 1, 1974 - March 31, 1975

Allen E. Anderson, R. Bruce Gill, Charles A. DeYoung, Howard L.
Geduldig, Steven Steinert, George E. Stewart, Gail Vanderpool,
Charles Wallmo, Joe B. Wallmo, Colorado Division of Wildlife;
Louis R. Burke, Fred P. Dasch, G. L. Terrell, Colorado Department
of Agriculture.
ABSTRACT

Twenty-three newborn mule deer fawns and 14 coyotes were captured, aged,
weighed, measured, radio-collared, ear-tagged, and released. They were radiotracked with ground and air, low frequency (46.5 MHz) telemetry equipment
within a 12,045 acre study area about 30 miles northwest of Fort Collins. Ten
fawns died three to 127 days from their capture dates, four fawns lost their
collars, and two fawns were never contacted after release. In only three fawns
was the probable cause of death (starvation) established.
Fawns were relocated
165 times, and 76 relocations were achieved without telemetry. The approximate mean (± SD) df.stance from capture sites to relo.cation sites -was 2,288
1,506 ft with a range of 0 ft to 7,100 ft. One coyote was caught in a steel
trap and killed by a commercial trapper, one coyote lost a non-functional
collar, and three coyotes were never contacted. Coyotes were relocated 39
times; three of these without telemetry and the approximate mean (± SD) distance from capture sites to relocation sites was 8,223 ± 7,034 ft with a
range of 1,600 ft to 39,600 ft. Four mule deer-coyote interactions are
described in which the female adult deer appeared to be the aggressor. Our
results with telemetry suggest the need for additional manpower and use of
both low (46.5 MHz) and high (150 MHz) equipment. From June 1974 through
March 1975 the percentages of radio-collared fawns to all unmarked fawns
recorded ranged from 40 during June to 2 during November based on 2,023
sightings of fawns. Fawns seen per man hr increased each consecutive month;
June (.162 per man hr) through November (2.175 per man hr), even though the
actual number of fawns alive must have been decreasing.

±

��-415RECOMMENDATIONS

(Continued)

In addition, the methodology associated with estimating the age of fawns needs
investigation and improvement. Appropriate materials should be acquired for
disease diagnosis of each fawn at the time of capture. A more durable plastic
and numbers rather than geometric symbols should be incorporated into the
design of future expandable radio collars for fawns. Ear tags for fawns
should have larger numbers on both sides.

��-417-

EVALUATION OF RADIO TELEMETRY
Allen E. Anderson

Radiotelemetry has been used successfully in studies of neonatal mortality
in dense white-tailed deer (Odocoileus virginianus) populations occupying
Texas Gulf Coast habitats (Cook et al. 1967), and to estimate coyote (Canis
latrans) movements in the Intermountain West (Hibler and Knowlton 1975:15-16),
Minnesota (Chesness and Bremicker 1975:3), and Arkansas (Gipson 1975:4).
Knowlton (1968) and his colleagues studied white-tailed deer-coyote interactions within Texas Gulf Coast habitats by radio-telemetry of 120 fawns over
a three-year period and concluded (P. 66) that over 90 percent of the fawn
mortality occurred during the first three weeks of life and that: "coyotes
apparently were involved in 75-80 percent of the fawn mortality".
Except for
the ongoing and unpublished studies of the Steen's mountain, Oregon population
(Trainer and Torland 1974), however, rates of fawn mortality in free-ranging
Rocky Mountain mule deer (Odocoileus hemionus hemionus) have not been estimated directly with or without radiotelemetry.
This is probably due to the
practical difficulties of finding living or dead neonates by either visual
ground search or telemetry in the mountainous terrain typical of most mule
deer habitats.

P. S. OBJECTIVE
To test and improv~ the field and statistical methodology used to estimate
selected population parameters of mule deer and coyotes.

SEGMENT OBJECTIVES
1.

Test the feasibility of using and perhaps improve radio-telemtry to estimate fawn mortality rates and coyote activity in mountainous habitats.

2.

Analyze data and submit reports for publication.

METHODS AND MATERIALS
Mule Deer Fawns
Capture Techniques
Very young fawns were usually captured by two-man teams maintaining a dawn-todusk surveillance of apparent high density deer areas from elevated sites.
Three or occasionally, four persons used 7 x 35, 8 x 40, and 9 x 35 binoculars
and 20X or l5X-60X variable-power spotting scopes to locate and observe fawns
at distances of about one-quarter to one-half mile. Fawns were observed until

�-418-

they had bedded and the doe had left the immediate vicinity. We memorized
the location of the fawn(s) in relation to prominent and nearby landscape
features such as gulleys, logs, boulders, bushes or trees; then walked rapidly
to the proximate vicinity of their bedding site(s). Because fawns would sometimes move in the interim, we verified their 'exact postion then generally
positioned ourselves so that we were visible to the fawn while walking toward
it. All successful captures were made by hand. Two attempts with a longhandled fish net were unsuccessful.
Fawns were sexed and immediately placed
in laundry bags equipped with a drawstring.
Body Weights and Measurements
Body Weight.--Fawns were weighed in the laundry bag to the nearest 1/4-lb with
a spring scale graduated in 1/4-lb increments. The weight of the laundry bag
was subtracted from the scale reading. The accuracy and precision of the
scale had been checked with known weights.
Length of Hind Foot.--From the calcaneum (heelbone) to the tip of the outside
claw with the hoof and metatarsal aligned on the same plane (modified from
Anderson 1948:66) with a fexible steel tape to the nearest 0.1 inch.
Hoof Growth.--From the "hairline to the growth ring on the front edge of a
front hoof" of (Robinette et al. 1973:321) who cites Haugen and Speake (1958).
The above instructions of the first author's and failure to check the second
author's technique resulted in a set of invalid measurements.
These data,
taken to the nearest mm with a steel tape, are not presented.
Neck Circumference.--By inspection and measurement, the minimum and maximum
circumferences were taken with a flexible steel tape to the nearest 1/8-inch.
Estimation of Age
The body weight and the length of hind foot of each fawn were entered in the
prediction equations of Robinette et al. (1973:320) to estimate its age to
the nearest day. The condition of the umbilical cord was also recorded.
Ear Tagging
Each fawn was tagged in both ears with identically-numbered metal tags using
the procedures of tag placement described by Downing and McGinness (1969).
Each set consisted of one red and one yellow tag with one or two digit numbers
engraved in black, one-inch in height, and on one side of the tag. The
mailing address of the Research Center was stamped on the staple portion of
each tag.

Characteristics of Radio Collars
Each fawn was fitted with a 46.5 MHz transmitter powered by mercury batteries.
As supplied by Albertronics, Inc., College Station, Texas, the transmitter
and batteries were embedded as one Unit in acrylic. The unit was bolted with
a nylon clamp to a self-expanding and contracting ABS white plastic collar
0.0625 inches thick and l~ inches wide. Geduldig (1975:9) modified the basic,

�-419-

expandable collar design from that of Fashingbauer (1962). The approximate
designed range of collar expansion and contraction was 25 em (9.8 inches) to
75 cm (29. 7 inches) based on an extrapolation of the growth curves for the
neck circumferences of male and female mule deer one month of age and older
(Anderson et al. 1974:29, 30). Because of the expandable feature of the
collar, the light wire. antenna could not be protected by securing its entire
length to the collar. Therefore, about 4.5 inches of its length was
embedded in caulking compound and secured to the collar with vinyl electrical tape. About 7.5 inches of the antenna arced unattached over the dorsal
portion of the collar. To identify each fawn, one or two geometric symbols
were painted on each side of the collar with methyl ethyl ketone, a solvent
of ABS plastic, resulting in black symbols on a white background.
The
weight of six, complete transmitter-collar units averaged 151 g and ranged
from 137 g to 158 g.
Collaring and Release of Fawns
Just prior to placing the collar on the fawn, the transmitter was activated by
soldering the lead wires with portable-battery or truck-alternator-powered
soldering irons and dOUble-wrapping the exposed portion of the soldered wire
and the entire transmitter-battery unit with vinyl electric tape.
After collaring, the prepared form (Appendix 1) was checked for completeness
and the fawn returned to its bedding site. The mean, +SD, and range of time
(minutes) elapsed between capture and release of 23 f~s
were 26.9 + 42.5
and 7 to 215, respectively. An attempt was made to leave the vicinity of the
capture site without disturbing the fawn. The capture site was marked with
red surveyor's tape.
Assessing Causes of Fawn Mortality
Each dead fawn, collared or unco Llared, was examined for the probable cause
of death. Particular attention was given to possible evidence of coyote
predation according to the description of White (1973). Carcasses in good
condition were submitted to the Diagnostic Laboratory at Colorado State
University, Fort Collins for clinical examination.
Coyotes
Capture Technique
We were not permitted to trap coyotes within the Maxwell Ranch property so
permission was received to trap on five privately owned ranches south of,
and adjacent to the Maxwell Ranch. All coyotes were trapped by George E.
Stewart on a contract basis. Most traps were loaned by the Colorado Department of Agriculture.
Coyotes were captured primarily with Montgomery No. 4 steel traps with offset
jaws using drag hooks attached to eight foot lengths of chain as anchors. Trap
jaws were not padded from 3 July to 20 September 1974, but were padded during
November-December 1974. Both trap jaws were padded by several wraps of oneinch wide strips of chenille cloth on each jaw and securing these to the jaws

�-420-

with four ties of light nylon cord. The entire trap was dipped in hot
paraffin to eliminate odor. A variety of attractants were used including
decomposed egg, decomposed beaver, fish oil, Nelson's Lobo, Grawe's No.1
and No.2, Mast's No.2 and No.6, tonquin, and skunk urine. Coyote urine
was used both alone and in combination with mOst of the other listed attractants. Traps were checked, and if necessary, reset, adjusted, removed, or
moved every 24 to 48 hours. While tranquilizer tabs (Balser 1965) employing
600 mg of powdered propopromazine were used initially to reduce trap injury
and to aid in handling the coyote, their use was abandoned because the high
temperatures, general lack of cover for shade, and possible trampling by cattle
(Hawthorn 1970) seemed likely sources of mortality for drugged and trapped
coyotes. No coyotes were caught while tabs were attached to the traps.
Handling the Coyote
Coyotes were subdued by the use of a commercially made choker device. While
one person held the coyote with this device, another slipped a loop of light
nylon cord over its muzzle, thus securing the jaws. This cord was fastened
to a broomstick, which, when pushed into the soil and rotated slowly, effectively immobilized the head and jaws. The trap was not released until all
data had been obtained and the coyote ear-tagged and collared.
Body Weights and Measurements
Body Weight
Coyotes were weighed to the nearest l/4-lb with the same spring scale used
for fawns. A light cord looped to the hind foot was used to suspend the
coyote from the scale.
Body Measurements,

General

The measurements listed below were taken with flexible steel tapes to the
nearest l/8-inch or 0.1 em and are modified from Anderson (1948:65).
Length of Hind Foot.--From the end of the heel bone (calcaneum) to the tip of
the longest claw.
Total Length. --The coyote was laid on its back on a flat surface and is the
straight line distance from the tip of the nose to the end of the last tail
vertebra, exclusive of hairs.
Tail Length.--Base of the tail to tip of the last vertebra, exlusive of
hairs.
Length of Ear.--From the notch at the lower opening of the ear conch to
the tip of the ear, excluding hairs.
Girth.--Chest circumference immediately posterior to the foreleg with
tape held snugly but without excessive tension.
Circumference of Neck.--By inspection and measurement the maximum and
minimum values were measured with the steel tape held snugly without excessive
tension.

�-421-

Estimation of Age
Coyotes were segregated into two age classes, young born during 1974, and
those more than one year of age by reading directly from the growth curves
for body weight, total length and length of hind foot and the dental criteria
of coyotes (Gier 1957:50-51, 55). In Colorado, most coyotes are born during
April and May (Hamlett 1938:9).
Ear Tagging
Each coyote was tagged in both ears with identically numbered, red and yellow
metal tags with black engraved numbers one-inch in height. These were similar
to the ear tags used for fawns. Based on our earlier experience with fawns,
some coyotes were tagged with the numbers facing forward or back as an aid
to their identification. Unfortunately, this information was not recorded
for all coyotes.
Radio-Collars
Coyotes were fitted with 46.5MHz transmitter powered by mercury or lithium
batteries and embedded as one unit in acrylic. Two types of collar and
antennae were employed with this unit. In the first type, powered by either
mercury or lithium batteries, the transmitter-battery package and its light
wire antenna were taped to a heavy leather collar l~ inches to 2 inches in
width. The antenna followed the complete circumference of the collar , and
like the transmitter-battery unit was completely covered with two wrappings
of vinyl electrical tape. In the second type, the transmitter-battery
package was bonded with acrylic directly to the collar to form an integral
trasmitter-battery-collar unit. The collar material was food conveyor
belting 2~ inches wide and 3/8 inch thick. The stainless steel'antenna was
O.lOl-inch diameter and about 11 inches long and curved, so as to be about
2~ inches above the mid-dorsal portion of the collar at its midpoint. One
complete collar-transmitter-battery unit of the first type weighed 296 g and
the mean and range of weights for five complete units of the second type were
465 g and 424 to 513 g, respectively.
Collaring and Release of Coyotes
The coyote transmitters were activated by soldering and taping the lead wires.
For the transmitter-battery-collarunit of the first type, the process was a
duplicate of that for fawns. For the second type the process was the same
for the lead wires; but, in addition, they were sealed with a fast-drying,
semi-flexible type of epoxy to form an extension of the acrylic transmitterbattery package. Both types of collars were fitted snugly to the coyotes neck
and fastened with two pop-rivets through pre-punched holes. For two collars,
brass, screw-on type rivets were used through pre-punched holes. Once collared,
the coyote was released from the trap and notes were taken of trap-incurred
injuries to the foot. The record form (Appendix 2) was checked for completeness and the coyote released.

�-422-

Radio-Tracking

Fawns and Coyotes

Two Johnson receivers, 8-channel, 46.5 MHz&gt; models 350 DF and Messenger III
were used in conjunction with earphones and a truck-mounted, Hy-Gain, Model
64B, four element, six meter, beam yagi antenna, two truck-mounted whip
antenna, two hand-held whip and two hand-held, directional, loop antennae to
track radio-transmitter collared fawns and coyotes. In most ground work, a
signal from a transmitter was first picked up with the truck mounted nondirectional whip antennae which indicated by the pulse rate per 15 seconds
heard on a particular channel that the particular fawn or coyote was in the area.
The hand-held non-directional whip antenna was carried about until a stronger
signal was received. When the apparent maximum strength signal was received
the directional loop antenna was used to locate the radio-collared fawn or
coyote. These procedures involved considerable walking, some over steep
terrain, and we carried the receivers in chest packs designed and fabricated
by Joe B. Wallmo.
In addition to ground telemetry, aerial telemetry was sporadically utilized
in attempts to locate radio-collared animals that were difficult to find on
the ground. The aerial antenna was a light wire 61 inches in length and
fastened to the wing of the Cessna 180. Generally, aerial telemetry was
performed at speeds of about 170 mph about 1,000 ft above ground surface.
Once the observer obtained a strong signal he handed the earphones to the
pilot who was usually able to place the animals within a given mile2 or
perhaps a 1/4-mile2 on the basis of increaSing, decreasing, and static signal
strength over a specific ground area. Collared coyotes were never, and
collared fawns only rarely seen during aerial telemetry.
Each telemetric or visual observation of fawns or coyotes was plotted on U.S.
Geological Survey Quadrangles, scale 1:24000, 40 ft contour interval. A
coded grid system, about 1,300 ft per side, was inked on a transparent, plastic
overlay to provide a more precise location of each radio-collared animal.
Field Observations
Each person on the project recorded the time to the nearest minute on entering
and leaving the study area. Within the study area, all living and dead deer
and other vertebrates seen were recorded and identified by time, sex, and age
class, habitat, and legal description to the 1/4-mile2 section on prepared
forms (Appendix 3). As a possibly important food item of coyotes, all cottontail rabbits (Sylvilagus sp.) seen were recorded. Plant phenology and any
phenomena conceivably related to fawn mortality rates were described in detail
on this same form. Particular attention was given to the behavioral interactions between deer and between deer and coyotes.
During the limited (60 permits, hunter's choice) deer season held on the
Maxwell ranch property, 26 October 1974 to 5 November 1974, I interviewed
hunters relative to any observations of radio-collared or ear-tagged fawns
and coyotes. The approximate positions of each observation were plotted on
topographic maps.

�-423-

Collection of Coyote Scats
During April, May, June and July 1974 all coyote scats noted in the field were
collected, placed in plastic bags and labeled with the date, location and
habitat. In addition, six marked walking routes totaling about 20 miles were
established during May 1974, to systematically collect coyote scats once each
month. Our objective was to relate, through generic identification of food
items in the scats, the relativ~ qualitative importance of deer in the inferred
diet of coyotes prior to, during, and after the period of fawn birth. Analyses
of this material has not yet been attempted.
DESCRIPTION OF AREA
Several areas were examined with the following criteria in mind: (1) relatively
high densities of mule deer during the summer and of coyotes yearlong, (2)
moderately rugged, mountainous terrain with a heterogenous, fairly open vegetative cover, (3) elevated vantage points from which large areas of fairly
sparse vegetative and topographic cover could be scanned, (4) exploited mule
deer and coyote populations, and (5) cattle range not adjacent to domestic
sheep range. The area selected for study was a 8,205 acre portion of the
Maxwell ranch property of the Colorado State University Research Foundation,
plus about 3,840 acres of private lands comprising portions of five ranches
adjacent to the northern, southern and eastern boundaries of the Maxwell ranch
property. The total of about 12,045 acres ranges in elevation from about 6,480
ft on the south boundary to 7,400 ft on the north boundary, TllN, T12N, R70W.
The study area is about 30 miles northwest of Fort Collins in Larimer County,
Colorado and just east of U. S. Highway 287. Its northern boundary is about
2.5 miles south of the Colorado-Wyoming state line.
The landscape of the study area is characterized by high, well-dissected,
shrub covered buttes on the south, shrub and pine covered hogback formations
on the east, high, pine covered hills with grassy meadows on the north, and
rolling prairie interspersed with isolated, high shrub covered hills and rock
outcrops on the central and eastern portions. Stonewall, Tenmile and Lonetree
creeks are intermittent streams and they and their tributaries trend south on
the southern portion, while the South and North Forks of Boxelder are permanent
streams and trend southeast in the northern portion of the study area. Small
portions of most major drainages are narrow, steep-sided canyons, up to about
70 ft in depth, and often supporting a diverse and almost impenetrable vegetation. The shallow loam and loamy foothill soils are both granitic and sedimentary in origin and highly erodible.
Subjectively, the climate is characterized by hot summers, high wind velocities,
and high intensity thunderstorms. Complete, prolonged snow cover is rare.
According to O. Boyd, Maxwell ranch foreman, total annual precipitation has
averaged about 13 inches over the past several years. There are no adequate
records of yearlong ambient temperatures on or near the study area.
Some conspicuous plants on the study area are ponderosa pine (Pinus ponderosa),
Rocky ~fuuntain juniper (Juniperus scopulorum), aspen (Populus tremuloides~,
narrowleaf cottonwood (Populus angustifolia), plains cottonwood (Populus
sargentii), willow (Salix spp.), snowberry (Symphoricarpos sp.), chokecherry

�-424-

(Prunus virginiana), alder (Alnus tenuifolia), Rocky Mountain maple (Acer
glabrum), true mountain mahogany (Cercocarpus montanus), raspberry (Rubus
deliciosus), currant (Ribes spp.), rose (Rosa spp.), skunkbush (Rhus trilobata),
boxelder (Acer negundo~tbush
(Atripl;XCanescens),
rabbitbrush
(Chrysothamnus sp.), sunflower (Helianthus sp.); cinquefoil (Potentilla spp.),
pricklepoppy (Argemone spp.), clover (Trifolium sp.), beardtongue (Penstemon sp.),
milkvetch (Astragalus spp.), lupine (Lupinus sp.), miner's candle (Cryptantha
sp.), Russian thistle (Salsola kali), larkspur (Delphinium spp.), sweetclover
(Melilotus spp.), aster (Aster spp.), fleabane (Erigeron spp.), mountain
dandelion (Agoseris sp.), yarrow (Achillea lanulosa), thistle (Cirsium spp.),
phlox (Phlox longifolia), cowparsnip (Heracleum lanatum), fringed sage
(Artemisia frigida), locoweed (Oxytropis spp.), bistort (Polygonum spp.),
eriogonum (Eriogonum spp.), blue grama (Bouteloua gracilis), wheatgrasses
(Agropyron spp.), needle and thread (Stipa comata) , needlegrass (Stipa spp.),
bluegrass (Poa spp.), sedge (Carex spp.), mountain muhly (Muhlenbergia montana),
Indian rice grass (Oryzopsis hymenoides), fescue (Festuca spp.), cheatgrass
(Bromus tectorum), junegrass (Koeleria cristata), dropseeds (Sporobolus spp.),
oatgrass (Danthonia spp.), barleys (Hordeum spp.), rushes (Juncus spp.),
horsetails (Equisetum spp.), and moss (Selaginella spp.).
Eight habitats were recognized on the study area: (1) prairie, (2) open ponderosa
pine-true mountain mahogany, (3) dense ponderosa pine, (4) true mountain
mahogany-currant, (5) true mountain mahogany, (6) dry wash, (7) riparian, and
(8) irrigated pasture and hayland. The latter habitat comprises about 194 acres
of the Maxwell ranch. Within the study area of about 12,045 acres, the most
important plants on a ground cover basis are probably blue grama, wheatgrasses,
needlegrasses, true mountain mahogany, ponderosa pine, currants and fringed
sage. The latter species is found within all natural habitats and is especially
abundant on many sites within the prairie habitat.
The riparian and irrigated pasture and hayland habitats, while relatively
small in area, seem to be an important influence on the temporal and spatial
distribution of some groups of deer during the late spring and autumn. At
these times, deer were often observed to bed and feed in adjacent ponderosa
pine-true mountain mahogany and prairie habitats during the day, then move
to the more mesic sites during early darkness, apparently remaining during the
night and returning at, or shortly after, dawn. Year-long, however, deer were
occasionally observed to follow a similar movement to the prairie habitat
from adjacent habitats.

RESULTS AND DISCUSSION
Capture of Fawns and Coyotes
Twenty-three mule deer fawns and 14 coyotes were captured, ear-tagged, weighed,
measured, examined and released within or adjacent to the study area. The
capture sites of fawns and coyotes are described in Tables 1 and 2, respectively.
The effort expended to locate, capture, and radio-track 23 fawns during June
1974 approximated 593 man-hours. The trapping effort required to capture the
14 coyotes is documented by month in Table 3, and trap injury to coyotes in
Table 4. Other vertebrates known to have been killed, released, or escaped
from steel traps are listed by month in Table 5.

�Table 1.
Description of capture site of 12 male and 11 female mule deer fawns, ear-tagged and fitted with radio-collars
during 1974.

Ear
Tag
. No.

Sex

Capture
Date

Legal Description
1/4S
S
T
R

1

M

6-11

SE

17

llN

70W

6920

Dry wash

Rhus tri10bat a

2

M

6-11

SE

17

llN

70W

6920

Dry wash

Rhus trilobata

3

F

6-13

SW

16

llN

70W

7020

Shrub-prairie
ecotone

Boute1oua gracilis, Opuntia sp , , Cercocarpus
montanus

4

M

6-13

SE

16

llN

70W

7020

Shrub-prairie
ecotone

Cercocarpus montanus

5

F

6-13

NW

2B

11N

70W

6640

Shrub

Cercocarpus montanus

6

F

6-13

NW

10

llN

70W

7120

Open pineprairie ecotone

Pinus ponderosa

Elevation
(f t )

Habitat

Dominant Vegetation at Bedding Site

v •

I

~

N

V1
I

7

M

6-16

NE

4

llN

70W

7120

Dry gu11ey

Pinus ponderosa, Stipa sp., Agropyron s pp .

B

M

6-16

NE

4

11N

70W

7100

Open pine

Pinus ponderosa, Boute1oua gracilis, Bromus
tecto rum

9

M

6-16

NE

4

11N

70W

7100

Open pine

Pinus ponderosa, Bouteloua gracilis, Bromus
tectorum

10

F

6-17

NW

4

11N

70W

7320

Open pine

Cercocarpus montanus, Pinus ponderosa

11

F

6-1B

SE

4

11N

70W

7160

Dry wash

Bouteloua gracilis, Unident. grass &lt;20 em ht ,

12

H

6-1B

NE

4

11N

70W

70BO

Open pine

Cercocarpus montanus, Pinus ponderosa

13

F

6-1B

NE

4

llN

70W

70BO

-Op en pine

Agropyron spicatum, .Symphoricarpus s p ,

14

F

6-19

NE

4

11N

70W

7240

Open pine

Pinus ponderosa,Cercocarpus

15

H

6-19

SW

33

12N

70W

7120

Dry wash-open pine

Pinus ponderosa, Cercocarpus montanus
Stipa comata

16

F

6-20

SW

21

11N

70W

6B40

Shrub

Cercocarpus montanus, Rhus trilobata

montanus

----------._----------------------------------------------------------------------------------------------------------------

�Table
1 . Description of capture site of 12 male and 11 female mule deer fawns, ear-tagged and fitted with radiocollars during 1974 (continued).

Ear
Tag
No.

Sex

Capt ure
Date

Legal DescriEtion
1/4S
S
T

R

Elevation
(ft)

Habitat

Dominant Vegetation at Bedding Site

17

F

6-21

NW

28

llN

70W

6680

Shrub

Cercoc~rpus montanus

18

F

6-21

NE

19

llN

70W

6760

Shrub

Rhus trilobata, CercocarEus montanus

19

M

6-23

NW

3

llN

70W

7060

Dense pine

Pinus ponderosa, Cercocarpus montanus,
AgroEyron sp.

20

M

6-24

SE

17

llN

70W

6960

Prairie

Bouteloua gracilis, CercocarEus
montanus

21

F

6-25

SW

22

llN

70W

6960

Shrub

Pinus Eonderosa, Boute1oua gracilis

22

M

6-25

SE

21

llN

70W

6920

Shrub

CercocarEus montanus, Bromus tectorum

23

M

6-26

NE

28

llN

70W

6840

Shrub

Rhus trilobata, Cercocarpus montanus

I
.j:N

0\
I

�Table 2.
Description of capture site of six male and eight female coyotes ear-tagged and fitted with radio-collars
during 1974.

Ear
Tag
No.

Sex

Capture
Date

1/4S

24

M

7-31

SW

29

llN

70W

6520

Riparian

25

F

8-17

SW

27

llN

70W

6940

Dry wash

Rhus trilobata, C~rcocarpus montanus,
Chrysothamnus sp.
JuniEerus scoEulorum, CercocarEus montanus

26

M

8-24

SW

29

llN

70W

6520

Riparian

Salix sp , , Populos spp., Chrysothamnus

27

M

8-30

NW

28

llN

70W

6700

Dry washshrub ecotone

JuniEerus scopulorum,
Salix sp ,

CercocarEus montan~s,

28

F

9-2

NE

27

llN

70W

6840

Shrub

CercocarEus montanus,

JuniEerus scoEulorum

29

F

9-4

NW

27

llN

70W

6960

Prairie-shrub
ecotone

Bouteloua

LeSal DescriEtion
S
T
R

Elevation
(ft)

Habitat

Dominant Vegetation at Trap Site

sp .

gracilis, CercocarEus montanus

I

.j:N
-....J
I

30

F

9-4

SW

27

llN

70W

6940

Shrub

Juniperus scopulorum,

31

F

9-6

NE

29

llN

70W

6580

Shrub-pr airie
ecotone

CercocarEus montanus, Bouteloua ~racilis

32

M

9-15

NW

27

llN

70W

6920

Prairie

Bouteloua

gracilis

33

F

9-19

NE

32

UN

70W

6480

Prai rie-sh rub
ecotone

Bouteloua

gracilis, CercocarEus montanus

34

M

ll-ll

SW

29

llN

70W

6420

Prairie

Bouteloua

gracilis

35

M

12-7

SW

29

llN

70W

6720

Riparianprairie ecotone

Bouteloua gracilis, Salix sp.

36

F

12-13

NW

10

UN

70W

6300

Dry washshrub ecotone

Bouteloua

37

F

12-15

SW

26

llN

70W

6720

Dry washshrub ecotone

Chrysothamnus sp., Cercocarpus montanus,
Bouteloua gracilis

Cercocarpus montanus

gracilis, Chrysothamnus

sp.

�Table
3. The 1974 coyote trapping effort by month and individual coyote within Sections 10, 21, 26, 27, 28,
29, 32, T11N, R70W.

Cumulative Trap Days
Per CaEture
Coyote No.
No. Days

Month

No. Days Traps
Were Set

No. Traps Set
Per Day*

Total Trap
Days

Total Coyo tes
Captured

July

29

6-19

310

1

24

291

August

31

19-25

611

3

25

606

September

20

10-25

465

6

26

733

November

19

11-16

234

1

27

866

December

18
117

9-18

263

3

28

915

6-25

1833

14

29

940

30

990

31

1,040

32

1,266

33

1,370

34

1,405

35

1,639

36

1,729

37

1,765**

I

.j:'N

00

117

. 1,833

14

*
Trap numbers we re changed according to relative numbers of trap visitations by coyote, disturbance by cattle,
porcupine, rabbits; discovery of additional coyote "nms", or apparent trap malfunction.
**

An additional 118 trap days were spent attempting to capture one additional coyote.

I

�Table 4.

Description

of steel trap injuries

Coyote
No.

Trap Jawsb
Padded
Not Padded

to 14 coyotes during 1974.

Foot Caught and
Location of Trap Jaws

Description

of Damage

Trap Type and
Sizec

24a

x

Left front, just above
paw

Skin cut to bone on dorsal surface
but no cuts on ventral surface.
One or more tendons may have been
severed.
Severe swelling of eptire
foot, moderate bleeding.

Montgomery 114,
Offset jaws

25

x

Left rear

Coyote chewed off three pads and
three toes. Severe circumferential
tissue damage where jaws held tarsal,
slight bleeding.

Montgomery 114,
Offset jaws

26

x

Left front, one inch
above paw

Skin cut to ligaments on left dorsal
surface but no cuts on ventral surface, severe swelling of paw but no
bleeding.

Kangaroo,
Offset jaws

Montgomery

I
.p-

N
\0
I

27

X

Left front, just above
paw

Skin not broken, moderate

28

X

Left front, just above
paw

Skin cut only on dorsal and left
side, no swelling.
Bled profusely
when jaws released.

Montgomery 114,
Offset jaws

29

X

Left front, below
joint

Skin cut deeper on dorsal inside,
moderate swelling.
Bled profusely
when jaws released.

Blake and Lamb
113, with teeth

swelling.

-------------------------------------------------------------------------------------------------------------------

113

�Table 4.

Description

of steel trap injuries

Coyote
No.

Trap Jawsb
Padded
Not Padded

Foot Caught and
Location of Trap Jaws

Description

of Damage

Trap Type and
Sizec

30

x

Right front, two inches
below joint

Skin cut dorsally but not ventrally.

Montgomery

31

x

Right front, diagonally
across phalanges

One phalange broken, others exposed.
Dorsal and ventral cuts similar in
depth.

Montgomery 114,
Offset jaws

32

x

Not recorded

Lateral cut through tendons and exposed bones.

Montgomery 114,
Offset jaws

33

x

Not recorded

Two center phalanges broken and
lateral laceration above middle
toes, possible torn ligaments.

Montgomery 114,
Offset jaws

113

X

Right front

Lateral cut across entire paw but
no bones broken.

Montgomery 114,
Offset jaws

35

X

Left front, by two
inside toes

One-half inch cut in skin of one
toe and phalanges broken.

Montgomery 114,
Offset jaws

36

X

Right fron t paw on
upper portion

One-half inch cut, possibly one or
two phalanges broken. No bleeding.

Montgomery 114,
Offset jaws

37

X

Right front paw on
lower portion

Skin cut, one inch long, and one
bone in foot broken.

Mont gome ry 114,
Offset jaws

bSee text for description

trapper on January 21, 1975, the entire foot was gone below the dew claw, and the

of jaw padding.

In all cases, a drag hook attached

to an 8 ft chain served as the trap anchor.

I

-!:'-

w
o
I

34

aWhen caught by a commercial
woun d well healed.

c

to 14 coyotes during 1974 (continued).

�-431-

Table 5. Vertebrates other than coyotes killed, released, or escaped from
the steel traps set for coyotes, 1974.

Species

July

August

September

November

December

Total

Cottontail
(Sllvilagus sp. )

1

8

6a

4

3

22

Porcupine
(Erethizon dorsatum)

1

2

lb

0

0

4

Bobcat
(~
rufus)

1

0

lC

0

ld

3

Magpie
(Pica pica hudsonia)

0

0

0

0

1

1

Total dead

3

10

8

4

5

30

a One rabbit released alive.
b Porcupine ref.easud alive.
c Bobcat released alive.
d Bobcat

escaped from trap.

Physical Characteristics of Mule Deer Fawns and Coyotes
Some physical characteristics and relevant temporal data are listed for 23
"newborn" fawns and 14 coyotes in Tables 6 and 7, respectively.
Some of
these data provide criteria for estimating their ages.
Estimating Age of Mule Deer Fawns
We attempted to assign age in days from birth to each captured fawn using the
growth curve equations of male and female body weight and length of hind foot
in Robinette (et al. 1973). Their equation for the length of hind foot for
each of 12 male fawns yielded nonsensical values. Their equation for the
length of the hind foot for each of 11 female fawns yielded plausible values
for some individuals but not for others. For example, the umbilical cord
of fawn number 3 was not dried on capture and the calculated age was one day.
On the other hand, the age of fawn 16 was calculated as 904 days, clearly an
impossible value. The body weight equations for each sex yielded this same
mix of plausible and impossible values. Therefore, the estimated ages thus
derived are omitted.

�Table
6. Some physical characteristics of 12 male and 11 female newborn mule deer fawns captured within or
adjacent to the Colorado State University Research Foundation Maxwell Ranch property ear-tagged and fitted
with radio-transmitter collars during 1974.

Ear
Tag
No.

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23

Sex

CaEtured
Date
Time

BodZ Weight
kg
1b

M
M
F
M
F
F
M
M
M
F
F
M
F
F
M
F
F
F
M
M
F
M
M

6-11
6-11
6-l3
6-l3
6-l3
6-13
6-16
6-16
6-16
6-17
6-18
6-18
6-18
6-19
6-19
6-20
6-21
6.21
6-23
6-24
6-25
6-25
6-26

4.0
3.6
2.9
3.3
3.5
3.9
4.5
3.4
3.2
4.3
4.1
4.0
3.9
3.4
4.4
6.7
4.9
4.0
4.8
3.7
3.5
3.5
4.9

1245
1248
0800
0800
0940
1305
1130
2052
2052
1250
0907
1848
2015
0921
1042
0915
0745
2008
1855
0801
2034
2051
0913

8. 75
8.00
6.50
7.25
7.75
8.50
10.00
7.50
7.00
9.50
9.00
8.75
8.50
7.50
9. 75
14.75
10.75
8. 75 '
10.50
8.25
7.75
7.75
10. 75

Length
Hind Foot
(em)

27.3
25.1
24.5
25.4
25.6
26.0
28.0
25.0
24.8
26.8
26.2
26.5
27.5
25.5
27.0
28.5
26.5
27.0
27.3
26.8
26.0
25.5
28.5

Neck
Circumference (cm)
Min
Max

16.5
17.8
17.3
15.2
16.5
15.5
16.8
15.2
15.0
16.8
15.7
16.5
15.2
14.0
17.8
19.0
16.5
14.7
19.0
17.3
15.2
15.0
18.3

20.3
17.8
17.5
18.8
17.8
19.0
17.8
17.3
16.5
17.8
16.5
17.8
16.5
16.5
19.0
19.8
17.8
16.5
19.8
18.8
16.5
16.8
18.3

Remarks

No. 1 and 2 were twins

No. 3 and 4 were twins
I

~

w

N

I
No.8 and 9 were twins
Birth witnessed, 1940-2020

�7 . Some physical characteristics of 6 male and 8 female coyotes captured with steel traps immediately south
Table
of the Colorado State University Research Foundation Maxwell Ranch property, ear-tagged and fitted with radiotransmitter collars during 1974.

EarTag
No.

Sex

Date
Captured

24
25
26
27
28
29
30
31
32
33
34
35
36
37

M*
F
M
M
F*
F*
F*
F*
M
F
M
M*
F
F

7-31
8-17
8-24
8-30
9-2
9-4
9-4
9-6
9-15
9-19
11-11
12-7
12-13
12-15

Body Weight
kg
1b

10.9
5.9
7.0
8.2
10.9
7.5
8.6
9.8

24.0
13.0
15.5
18.0
24.0
16.5
18.5
21.5

-

-

6.4
8.4
12.5
10.0
10.0

14.0
18.5
27.5
22.0
22.0

Hind Foot

19.0

17.8
19.0
17.0
18.5
19.5
19.5
19.0
17.0

-

19.1
16.5
16.5

Lengths (em)
TaU
Ear

30.5
27.9
28.0
33.5
35.0
32.5
34.5
34.0
36.0
33.0
30.5
34.3
33.0
33.0

11.0
10.2
9.0
11.5
10.5
12.0
10.5
12.0
12.0
11.0
10.8
10.8
10.2
10.8

Total Body

Chest

-

47.0
38.1
38.1
37.5
44.4
43.2
45.7
41.9
48.3
34.3
47.0
55.9
50.8
53.3

91.4

-

-

108.0
111.8
106.7
106.7

Circumference
Min Neck

~em2
Max Neck

26.7
17.8
20.3

26.7
22.9
21.6
21.6
25.4
27.9
25.4
22.9
27.9
20.3
27.9
38.1
33.0
29.2

-

23.5
22.9
24.1
21.6
24.1
19.1
24.1
34.3
22.9
22.9

* Coyotes more than one year of age. All others were estimated to have been born during 1974 on the basis of the
growth curves and dentition criteria of Gier (1957:50, 51, 55).

I

.j:'-

w
w
I

�-434-

Estimating Age of Coyotes
Based on the very approximate criteria used, two males and four females
exceeded one year of age. Four males and four females were born during 1974.
The body weights of the 14 coyotes resemble those of similar approximate age
and season of collection from Kansas (Gier 1957:52) and California (Hawthorne
1970: 71) ,
Fate of Radio-Collared Fawns
Of the 23 fawns captured, 10 were found dead as of 31 March 1975; 3, 3, 3, 7,
7, 16, 26, 42, 86, and 127 days following capture and release. Two fawns
were never contacted following release. Four fawns were found very soon after
death; two visually and two using telemetry. Three carcasses were suitable
for certain clinical studies and starvation was stated to be the probable
cause of death (Table 8). Because fawns 8 and 9 were captured less than an
hour after birth, our disturbance may have been a factor in their deaths.
Four fawns lost their expandable collars. The cause(s) of mortality could
not be determined for seven fawns. Typically, these animals had been scavenged
or deteriorated to such a degree that positive identification of the causal
factor(s) of death was impossible (Fig. 1). This was also true of nine
uncollared fawn carcasses born during 1974 and found from 8 July 1974 to 14
March 1975. In Texas white-tailed deer, White et al. (1972:897), reported that:
"capture, and particularly the marking, of young fawns increased mortality."
Fate of Radio-Collared

Coyotes

Only one radio-collared coyote (No. 24) was known to have died as of 31 March
1975. This animal was trapped and killed for profit on 21 January 1975 about
l~ miles from its capture site. A minimum of 20 additional coyotes are believed
to have been shot during the summer, autumn, and early winter of 1975 within
or adjacent to the study area. None of these animals were reported as being
ear-tagged or radio-collared.
Radio-Tracking

Fawns and Coyotes

Some indices of signal distance and strength obtained with our telemetry equipment under ideal, line-of-sight conditions are given in Table 9. The time in
minutes expended obtaining signals with various combinations of antennae are
described statistically in Table 10. Relevant ear-tag, mortality, and radiocollar data for 23 fawns and 14 coyotes are given in Table 11. Capture sites,
death sites, and telemetric and visual locations of 23 fawns are plotted on
portions of U.S.G.S. Topographic Quadrangles,Figs. 2-18. Approximate distances of radio-collared fawns and coyotes between their capture sites and
subsequent telemetric and visual locations are described statistically in
Tables 12 and 13, respectively.
Observations of Deer and Coyotes
The total number and numbers of mule deer recorded per man hour within the
study area for each of 11 months are listed by sex and age class in Table 14.

�Table
8. Summary of clinical examinations of three male radio-collared mule deer fawns by the Diagnostic Laboratory,
College of Veterinary Medicine and Biomedical Sciences, Colorado State University, Fort Collins. *

Ear
Tag
No.

8

Date
Fo und

Sex

Date
Tagged

Dead

Bacteriology

Virology**

M

6-16

6-19

Intestine - Bacillus
and E. coli
Liver - negative
Lung - negative

FA for BVD - negative
FA-for PI3 - nega~ive
FA for IBR - negative

Pathology

No milk in stomach and the meconium was still
in the colon and rectum. No food particles
in intestine.
Histology - autolytic changes in intestine.
Congestion of centri-lobular areas and
vacuolation of hepatocyte cytoplasm of
liver. No visible lesions of lung, kidney,
thymus, thyroid. Summary: No signifiI
cant lesions were recognized histologi+:w
cally.
VI
Diagnosis: Starvation as the probable
cause of death.

9

M

6-16

6-19

Intestine, liver,
lung -~. coli
Ltmg - Bacillus subtilis

No tests

No milk in stomach or intesttne. Some
fecal material had grass and possibly
some milk residue. No other lesions
noted.
Histology - negative
Diagnosis: Starvation
cause of death.

as the probable

--------------------------------------------------------------------------------------------------------------------------

I

�Table 8.
Summary of clinical examinations of three male radio-collared mule deer fawns by the Diagnostic Laboratory,
College of Veterinary Medicine and Biomedical Sciences, Colorado State University, Fort Collins (continued).*

Ear
Tag
No.

4

Sex

Date
Tagged

Date
Found
Dead

Bacteriology

Virology**

M

6-13

6-20

No tests

No tests

Pathology

No milk in stomach or intestine, but a
2 em diameter ball of dry grass was
present. Most of the meconium was gone
and the remainder was soft. No
evidence of diarrhea.
Diagnosis: Starvation as the probable
cause of death.
I
.po.

*

Abstracted or paraphrased

** FA

=

from laboratory reports prepared by Dr. A. E. McChesney and Dr. W. W. Brown.

fluorescent antibody.

BVD = bovine viral diarrhea.
PI3 = type of influenza virus.
IBR

= infectious

bovine rhinotracheitis.

Vol
0\
I

�-437-

Fig. 1. Fawn number 13, captured and released 18 June 1974; located with
aerial telemetry 1, 2, 3, 4 July 1974, but never seen alive. Found dead
3:03 P.M., 14 July 1974, with ground telemetry about 1800 ft from its capture
site.

�Table 9.
Signal
with line-of-sight

distance*
and strength**
achieved by lithium
and mercury b a t t.e ry+powere d 46.5
orientation
over slightly
rolling
prairie
using three types of antennae.

Johnson
Receiver

Date

350 df
Channel

Type of
Battery

Signal Distance
Truck-Mounted Whip
1/2
1
1 1/4
1 1/2
1/4

MHz transmitters

(Miles)*
Hand-Held Whip
1/2
5/8
3/4

1

7-30-74

L

Lithium

E

G

W

0

E

E

G

0

0

1-21-75***

L

Lithium

0

0

0

0

0

0

0

0

0

E

G

0

0

,./

,

7-30-74

Mercury

E

G

E

Signal

W

0

Distance_Cft)

*

/
/

Directional

Hand-Loop,

,.,,/

E

.

/

I

+0W
OJ

"

I

I

/
/

/

1050

1320

/~

G

0

792

1

"

7-30-74

L

Lithium

E

1-21-75***

L

Lithium

G

W

W

7-30-74

E

Mercury

E

G

0

*

On the

ground within

**

E = excellent

signal;

the

study

area.

G = good signal;

W = weak s Lgnlal , 0 = no signal.

*** Same transmitter
used on coyote 1124;' signal
on l-21-,!'75 audible
at 2100 ft
1500 ft with the hand-held whiP. and 1650 ft with the directional
hand loop.
expiration
for this transmitter
was 1-23-75.

,

'I,
I

with the truck-mounted
whip, at
The theoretical
date of battery

�-439-

Table 10. The various combinations of four types of antennae uS~d and a
statistical description of the time (minutes) between receivin~ a signal on
the first antenna and a signal on the second antenna or visual or telemetric
identification of a particular mule deer fawn.

First

Antennae
Second or Identified Fawn

N

x

f3n

Min

Max

60

60

26.;0

10

42

53

79

Yagi to truck whip

1

Yagi to loop

2

Yagi to hand whip

0

Truck whip to yagi

2

66.0

Truck whip to visual

5

80.2

66.9

25

180

4

42.2

50.4

1

110

Truck whip to hend whip

2

4.0

4

4

Hand whip to loop

16

18.5

18.9

3

75

Hand whip to visual

10

64.5

68.3

9

180

Hand whip to telemetric only

2

184.5

120

249

Loop to visual

5

25.8

27.1

6

73

28

54.8

55.4

2

180

14

61.4

64.6

6

249

Truck whip to telemetric
identification

-'

First telemetric signal to
visual identification
First telemetric signal to
telemetric identification

�Table

11.

Description

Ear
Tag
No.

Species
and
Sex

of identically-numbered

Capture
Date

Date
Found
Dead

Date
Collar
Found

metal ear tags and radio-collars

Color of Ear Tags
L
R

fitted to 23 mule deer fawns and 14 coyotes during 1974-75.

Theoretical
No. of 46.5 MHz
Date of Battery
TransmitterExpiration
Battery Unit

Pu.l se

Rate per
15 Sec.

Battery
Type

Color of
Collar

Collar
Symbo I,
or 'l~o.

v

Remarks

FAWNS
1

M

6-11

2

M

6-11

3

F

6-13

6-29-74

4

M

6-13

6-20-74

5

F

6-13

6

F

6-13

7

M

8

9-6-74

Yellow

Red

8-28-75

lAl5

13

Hg

'White

Yellow

Red

9-4-75

2B16

13

}1g

White

Yellow

Red

9-2-75

4DlO

21

Hg

White
White

twins

White

s:

Collar broken, 4 pc.

20

Hg

White

o

5E19

14

Hg

White

L

No.8

Hg

White

U

twins

Red

7-2-75

6F12

2~

Hg

Red

9-28-75

4Dl8

13

Hg

White

Yellow

Red

7-30-75

5Ell

21

Hg

6-16

Yellow

Red

8-2-75

7G13

M

6-16

6-19-74

Red

Yellow

9-26-75

6-19-74

4

9

M

6-16

Red

Yellow

9-29-75

6 FZ:O

13

10

F

6-17

Red

Yellow

10-7-75

:,C17

13

Hg

White

X

11

F

6-18

Red

Yellow

9-1-75

2B8

21

Hg

White

X.

12

M

6-18

13

F

6-18

14

F

6-19

15

M

6-19

16

F

6-20

17

F

6-21

18

F

6-21

19

M

6-23

20

M

I

&amp; 9 were

Red

Yellow

11-10-75

7G21

15

Hg

White·

Red

Yellow

8-4-75

5E5

15

Hg

White

Red

Yellow

9-29-75

17

Hg

White

10-24-74

Red

Yellow

8-.5-75

8L22
3C9 .

'1-4
y"V
~D

22

Hg

White

XU

Yellow

Red

11-2-75

6F20

13

Hg

White

U

Collar from fawn 9 \

Yellow

Red

10-22-75

5E19

14

Hg

White

L

Collar from fawn 8 "

Yellow

Red

9-15-75

8L14

13

Hg

White

XL

Red

Yellow

7-12-75

6F12

20

Hg

White

7-3-74
8-1-74
2-3-75

Yellow
~

Red

9-11-75

15

3C3
~

L

Hg

White

•

t

o

7-14-74

6-24

_____________________________

collar in tact
No. 3 &amp; 4 were

•

Yellow
Yellow
2-11-75

•
•

No. 1 &amp; 2 were twins,

Co Ll.ar-broken

(from,

fawn No.4)

xy
_

�Description
Table

11.

Ear
Tag
No.

Species
and
Sex

~

(continued)

Capture
Date

of identically-numbered

Date
Found
Dead

Date
Collar
Found

metal ear tags and radio-collars

Color of Ear Tags
R
L

Red

fitted to 23 mule deer fawns and 14 coyotes

Theoretical
Date of Battery
Expiration

No. of 46.5 MHz
TransmitterBattery Unit

9-8-75

6F6

21

F

6-25

9-19-74

Yellow

22

M

6-25

6-28-74

Yellow

6-26

7-3-74

Yellow

Red

4D4

M

8-23-75

23
COYOTES

M

7-31

1-21-75

Yellow

Red

1-23-75

8L36

24
25
26
27

F
M
M

8-17

28

F

8-24
8-30
9-2

29

F

9-4

.

Hg

1.1.

17

Li

18

Li
Li

Yellow

Red
Yellow
Red

3-1-75

lA33

Yellow

Red

Yellow
Red

9-6
9-15

33
34

M

35

M

11-11
12-7

Yellow
Red
Yellow
Red

* Erratic pulse rate.

16

26

F
M
F

12-15

X.

White

2B38
lA37

31
32

F

*

4D40

Red

12-13

XI

White

5-13-75
4-9-75

9-4

F

White

Hg

2-3-75

F

36
37

lA7

Red
Yellow
Red

Yellow
Red
Yellow
Red

-

3C39

19
13
15

Hg

Remarks

None
Black
None
Red &amp; black
Black &amp; white None
None
Red &amp; orange
Red

I

•...

""
""

None

Li

Orange &amp; who

None
None
None
None

I

Trans.non-funct.
when found

2-}-75

7G37

15

Hg

Blue &amp; red

3-5-75
2-22-75

5E35
6F36

4-3-75

2B34

15
14
13

Hg
Hg
Hg

Blue &amp;'white
Black &amp; blue
White

Li
Li

White

Yellow
Yellow

7-23-75
8-26-75

8L45
G44

Red

9-1·2-75

6F43

20
20
21

7-21-75

3E42

20

Yellow

Collar
Symbol
or No.

Hg

Red

30

9-19

8-11-75

Color of
Collar

15

Li

Yellow

12-10-74

Red

Pulse
Rate per Battery
15 Sec. Type

during 1974-75. (cont.).

None

Li

White
White

1
2
None

Li

White

None

�-442-

.

I

.

(

.

",1.

.';j':'

~.
\I

II
_ t\7110

,; -;-.;;.

-

\\..

\'

-'

"

"

-,,-,---7//6

'j, ,~-

Fig. 2. Fawn No. 1 captured 11 June 1974 (solid circle) with telemetric
and visual observations by month. Last observation 15 September 1974,
TUN,

R70W.

�-443-

Fig. 3. Fawn No.2 captured 11 June 1974 (solid circle) with telemetric
and visual observations by month. Intact radio-collar found (half solid
circle), 6 September 1974, TllN, R70W.

�-444-

7168

17'~/,

/

"

.

-:

"

,/
II

~&lt;;

-"

',I

()

"

.

il

.lo

_JVItO

C'6

e : ."

'----

.;-7
-,~.:.

Ji.
~~ .•.
q.;.-~.--'--'---.-_ ~--

\

\\
,I

..../'

.p~~=f--'-,,,*o..

/-

Fig. 4.
Fawn No.3 captured 13 June 1974 (solid circle)
with telemetric
and visual observations
by month.
Found dead ("X" circle),
29 June 1974,
TUN,
R70W.

�-475July 1975

JOB PROGRESS

State

0f

Project

__

-=C.:::O=L.:::O::.:RA:.:DO=-=_

No. __ W~-~3~8-_R~-_2_9~

Work Plan No.
Job Title
Period

REPORT

~1~9~

_
_

Deer-Elk
Job No.

Investigations

2
-------~~------------------

Evaluation of Herd Structure Methodology
---~~~~~~--~~~~~~--~~~~~---------------------

Covered:

Personnel:

April 1, 1974 - March 31, 1975

Allen E. Anderson, David C. Bowden, Eugene
Medin, Elmer Remmenga, Charles Wallmo.

G. Johnson,

Dean E.

ABSTRACT
The validity of the assumptions or models for sample observations from each
of two sampling plans which permit the development of measures of reliability
for estimates of fawn to doe ratios of Rocky Mountain mule deer (Odocoileus
hemionus hemionus) sampled after the autumn hunting season are examined and
the results of data analysis under an appropriate model are presented.
The
first plan (route counts) consisted of ten, dawn-to-dusk, circular routes
designed to yield the maximum number of classified deer on representative portions of the Cache la Poudre winter range. The routes were walked by two
observers twice each breeding season; (1) during the estimated peak, and (2)
during the late portion, 1962-65.
The second plan (quadrat counts) consisted
of classifying deer within randomly-selected,
l/4-mile2 quadrats, stratified
by subjective estimates of low and high deer densities and proportionally
allocated within each of eight topographic quadrangles on Cache la Poudre
winter range and two quadrangles within, and adjacent to the Maxwell Ranch,
18 November 1974 - 9 January 1975.
Because of the statistical models examined, fawn to doe ratios are stated in
terms of PF = F/(F + D) where PF = the proportion of fawns, F = the number of
fawns, and D = the number of does. Tests indicated significant (P &lt;0.01)
departure of the observations from the binomial distribution and simple linear
regression through the origin of fawns per group on does and fawns yielded
variances about one-half of those derived from binomial models.
RegreSSion
analysis allowed valid comparisons of routes within years and between the peak
and late breeding periods on the same route.
Among pooled route counts, the
approximate peak period of breeding exhibited significantly
(P &lt; 0.05) larger
proportions of fawns (PF) than the late period of breeding.
Among the quadrat
coun ts, the proportion of fawns was significantly (P &lt; O. 01) higher .within the
low deer density quadrats on Cache la Poudre winter range. The relative merits
of the route and quadrat count sampling plans require testing with a random
selection of routes.
However, one important feature of the quadrat count
sampling plan is that both minimum deer densities and fawn to doe ratios may
be inferred from the same sample.
The minimum estimates of the mean (+SE)
numbers of mule deer per mi1e2 during November-December
1974 were 25.5-+ 2.4
(Cache la Poudre), and 26.8 + 12.2 (Maxwell).
Deer on an estimated lll-quadrats would require classific~tion for the proportion of fawns (PF) to within
5 percent of the true value at the 90 percent level of confidence.

�-476-

RECOMMENDATIONS
Because many of the field and statistical problems
be fully anticipated the following recommendations

involved cannot now
are tentative:

1.

The relative merits of the two sampling plans examined herein, route
counts and quadrat counts; require further work involving random
selection of routes.

2.

One assumption of the regression model presented; that groups of deer
are encountered at random should be tested with known numbers of deer
within large enclosures of natural range.

�-477-

EVALUATION OF HERD STRUCTURE METHODOLOGY
David C. Bowden and Allen E. Andersona

Sample ratios of sex and age of Rocky Mountain mule deer are routinely
obtained by many agencies concerned with its management.
In some cases,
the ratios alone are believed important in evaluating the effects of
hunting or observed environmental stress. In other cases they form the
basis for calculating population characteristics difficult to measure
directly. Hanson (1963) reviewed the rationale and methods of calculating
the equations presented by Rasmussen and Doman (1943), Kelker (1944),
Robinette (1949), and Lauckhart (1950); which use sample ratios of sex and
age to calculate one or more of the following population characteristics
of mule deer (Odocileus hemionus); (1) productivity (p. 21), (2) survival
rates (p. 30), and (3) absolute abundance (p. 45). This author, presumably
referring primarily to sex and age ratios of vertebrates generally, states
(p. 6): "Methods for obtaining valid data are not covered here because they
have been treated in many other publications, ---", and that (p. 7) "Procedures for calculating confidence limits of estimates are not considered
in detail
"We
are not aware of any published statistically adequate
procedure for collecting "valid data" on sex and age ratios of mule deer.
Recently, however, Paulik and Robson (1969) and Seber (1973:353) have presented the assumptions, models and methods of computing confidence intervals
and sample size requirements for the change-in-ratio methods of estimating
absolute abundance of many vertebrates including deer. Actually, much of
the literature reporting sex and age ratios of deer has been unpublished
with little or no discussion of sampling methodology or statistical attributes. Some exceptions were: Leopold et ale (1951:118) who attempted to
estimate sample size requirements, Riney (1956) who presented a formula
for calculating approximate confidence limits about sample ratios, and
Wallmo (1964) who discussed probable sources of bias and sampling errors.
Overton (1971:455) and Eberhardt (1971:494) mention sex and age ratios of
deer briefly but do not discuss sampling methodology or statistical attributes. Caughley (1974) examined by simulation the assumption that age
ratios reflect the dynamics of a population and concluded (p. 562) that;
"age ratios cannot be interpreted without a knowledge of the rate of increase,
and if we have an estimate of this rate we do not need age ratios." However,
this author also states (p. 562) that a "sudden change in a ratio --- indicates
that something has happened," and there is some evidence that fawn to doe
ratios may reflect the dynamics of some deer populations.
For example, Smith
et al. (1969) documented a marked decline in a deer population subjected to
controlled, any-deer hunting regulations and believed that concurrent and
inexplicably poor fawn to doe ratios (11:100-49:100) partially responsible.
Swank (1958:78-79) associated summer range drought as one important factor in

a
Dr. Bowden, Dept. Statistics, Colorado State University, wrote the METHODS
AND MATERIALS and nearly all the RESULTS AND DISCUSSIONS sections and with
the assistance of Graduate Student Gene Johnson developed the statistical
rationale described herein. A. E. Anderson wrote the ABSTRACT, INTRODUCTION,
and RECOMMENDATION sections. The field work was performed by A. E. Anderson
and D. E. Medin during 1962-65 and by A. E. Anderson and Charles ~-lallmoduring
1974-75.

�-478-

the decrease from 90:100 to 50:100 in winter fawn:doe ratios and a concurrent,
calculated loss of 18,000 deer on the Kaibab North, 1953-1954. Taber and
Dasman (1958:129) found large differences in fawn :doe ratios (71:100, 115:
100, 145:100) among three adjacent range types. We hypothesize that estimated
fawn to doe ratios may be useful in assessing rates of fawn mortality if the
data can be obtained under conditions which permit the valid estimations of
the standard errors of the estimated fawn to doe ratio and the sampling
intensity for small coefficients of variation of the fawn to doe ratio can be
practically achieved.

P. S. OBJECTIVE
To test and improve the field and statistical methodology used to estimate
selected population parameters of mule deer and coyotes.

SEGMENT OBJECTIVES
1.

Study existing herd structure data to provide an improved statistical
rationale for analyses of fawn:doe and buck:doe ratios.

2.

Analyze data and submit reports for publication.

METHODS AND MATERIALS
An ecological study of the Rocky Mountain mule deer population w~thin a
1376.5 km2 portion of the Cache la Poudre River drainage in Roosevelt National
Forest in Larimer County, northcentral Colorado was conducted from 1961 to
1965. In one facet of the study it was of interest to examine the fawn, doe,
buck components of population at specified time periods (Anderson and Medin
1965). Since it was not feasible to estimate the actual number of deer in
each category, the fawn to doe ratio of population sizes and the buck to doe
ratio of population sizes were studied.

Deer were classified into fawn, doe, buck categories in late November through
December and again in January through early February for 3 winter seasons
beginning in 1962: 11-29-62 to 12-18-62, 1-3-63 to 1-27-63, 11-21-63 to 12-14-63,
1-9-64 to 1-31-64, 12-3-64 to 12-17-64 and 1-13-65 to 1-27-65. The winter
range of the deer population in the study area was sampled by selecting 10
representative sections of the winter range (Fig. 1). A route to be traversed
by walking was outlined on aerial photographs of each representative section.
The routes were located to classify the maximum number of deer. The length
of the route was determined by the distance which could be covered in the
given terrain by walking from dawn to dusk. The routes were covered by two
observers working together, generally with one observer maintaining a vantage
point while the other observer attempted to bring hidden deer into view.
Classifications of deer were made by using 8X40 or 7X35 binoculars and 20X
spotting scopes. Deer were classified as fawn, doe, buck or unidentified by
groups on the basis of relative body size, presence or absence of antlers

�so:

!.o'

105' 30'

w'

::0'

-l
-j-+,=*-~,,,,,,,,~jl-"~t~~~
T10N

~gl

\f~Ti..Ff~t~~1
T 9 N

••'d"\ R

\\I!)u:- ••
_..JI

l~r

••
~-""'~

I

I

I

I
.p..

-...J

1.0

~40

I

T8Ni
,

7N

GAME

MANAGEMENT
DEPT. OF

SIXTH

PRINCIPAL

UNIT

19

GAME AND FISH

MERIDIAN

Ie:.;,,-t.-=_';;.=:=,~~""=,,_~~=-J~_~

Mill!s

40'
30'

LEGEND

~
~"'~..:i7~.-

'.
"!~

..•;-;

'
--~'---

"

_

_I

-'-r3-:rIct-,,:-a"hn:;:;r--

' -

--'-e~.t-"

"" - ,

a~r---:=::"~-:'-~

Management

Unit

AppralllTlale

Tronsihonot

~!~~

TeN

Boundary

~_~_~.8~_~d_.~m~~

Zone
__Rono~s

_

p'.:IJt'.t'('f,
,..,rll. A",,,r'&lt;'8r dOl,.""
;"&gt;,,,'y,.,,,""(

Fig. 1.
The location
of 10, all-day walking routes designed to classify
during the approximate
(1) peak, and (2) late portion of their breeding

the maximum number of mule deer
season,
1962-65.

�-480-

and the applicable criteria listed by Dasmann and Taber (1956:80). A
group was identified by observing the joint behavior and spatial distribution of the deer. Only those groups in which all deer were classified
were used in this analysis. We refer to this sampling plan as the route counts.
In 1974 a new sampling plan was applied to the study area. The deer winter
range was divided on 1:24000 scale U. S. Geological Survey Topographic
Quadranges into square sampling units (1/4 mile2 or one-half mile per side)
and the sampling units placed into strata. Each sampling unit was classified
subjectively into high and low deer density strata by eight Topographic Quadrangles (16 strata). Counts started 22 November and were concluded 31
December except for two sampling units covered 9 January, 1975. Sampling
effort was proportionally allocated according to the total number o£ sampling
units per stratum (Topographic Quadrangle) except that at least two sampling
units were taken from every stratum. Two walking observers classified deer
on each selected sampling unit. The sampling unit was approached and traversed to classify the maximum number of deer. Four sampling units were
usually counted beginning at dawn and finishing at dusk each day. The
counting sequence of the sampling units was arranged so that adjacent or
nearby sampling units were counted on the same day to minimize duplicate
classification of the same deer and to allow for efficient use of available
time. Deer were recorded by groups as before. We refer to this sampling plan
as the quadrat counts.
In this report we present these data and examine the validity of assumptions
or models for the sample observations from each sampling plan which permit
the development of measures of reliability of estimates of ratios of the deer
population size components.
In addition, the results of the data analysis
under a model not contradictive to the data will be presented. All analyses
are limited to the fawn to doe ratio. The buck to doe ratio will be examined
in subsequent reports.

DESCRIPTION OF AREA
That portion of the Cache la Poudre winter range sample has been described
by Anderson et ale (1970:390). The Maxwell ranch area is described in the
Work Plan 19, Job 1 Progress Report accompanying this report.

RESULTS AND DISCUSSION

Statistical Models for the Observations

- Route Counts

Although the stated parameter of interest is the fawn to doe ratio, there are
equivalent parameters which are preferable under some statistical models.
The following notation permits an easy definition of the equivalent parameters.
Consider the population of deer at a specified point in time and define F, D
as the number of fawns, does, respectively, in the population at this time.
The stated ratio of interest is RF = F/D.
Consider instead PF = F/(F + D).

�-481-

Then clearly, ~ = PF/(l - PF). The statistical models given here will be
stated in terms of PF so that the estimation problem will be to determine
the proportion of fawns (relative to does and fawns). Estimates of PF can
be converted into estimates of RF.
Much of the difficulty in objectively sampling natural wildlife populations
arises in determining a sampling frame (a list of sampling units) which can
be sampled by using statistical sampling plans while at the same time providing sampling units which when measured provide data on the target population of interest. A statistical sampling plan is a scheme of sampling unit
selection such that the probability of obtaining each individual sampling
unit in a sample of a given size is known or can be calculated. Conceptually,
the use of the animals themselves as a sampling unit is an appealing sampling
frame. In its simplest form each animal is a sampling unit. A desirable
property of having the animals as sampling units is that conceptually the
value of PF could be determined without error if all the sampling units were
measured. The practical problem with such sampling frames is that once a
specified sample of animals was determined, the animals have to be located
and classified. Such a sampling frame also implies that the population size
is known and that the animals or groups of animals (sampling units) can be
listed.
As an alternative to forming sampling units from or of the animals themselves,
sampling frames are constructed by considering a given area or region, perhaps volume, that the population is associated with, restricted to, or contained in. As in the 1974 sampling of the Poudre drainage, the region is
subdivided into parts which are considered as the sampling units. The construction of such a sampling frame and the actually selection of the sampling
units to be measured is not difficult. However because of the movement of the
animals among the sampling units and because the animals behavior limits their
detection by human observers, measurement errors on such sampling units can be
quite important. The explicit or implicit use of these two types of sampling
frames will now be examined relative to the data of the present study.
First, statistical models or sets of assumptions which give rise to the use of
a sampling frame dependent on the animals themselves will be examined. Thus,
let n represent the number of animals identified as either does or fawns while
d and f will be the numbers of does and fawns in the sample of n.
Consider the following assumptions:
1. The classification category for each animal is independent of the
other animals in the sample. Alternatively the probability that an individual
animal is a fawn does not depend on the other animals in the sample.
2.

The probability that an individual animal is a fawn is PF.

It is well known that these assumptions imply that the probability distribution
of f is given by the binomial distribution where n is the number of trials,
PF is the probability of success on an individual trial and f is the number of
successes out of n. In the sampling frame terminology this same binomial distribution is arrived at by assuming that a simple random sample without replacement
is taken from a sampling frame which lists the individual animals. The sampling

�-482-

procedure then implies the distribution of f is given by the hypergeometric
distribution.
But in most situations this hypergeometric distribution can
be adequately approximated by the binomial distribution given above.
In general, the practicality of actually constructing and using such a sampling
frame is nil, but is is common practice to assume the collected data, however gathered, satisfy assumptions 1. and 2. and in essence claim that the
collection procedure operates as if a simple random sample of the individual deer was taken. It remains to establish that the data are not in
strong disagreement with the implications of these assumptions.
Since the
alternatives to accepting assumptions 1. and 2. are complicated, the disagreement of the actual data with assumptions 1. and 2. will be explored
next.
The assumptions will be examined for the data collected on the route counts
from 1962 to 1965. On each route the deer were observed in natural occurring
groups, that is, classified into groups by their behavior and the proximity
of individuals to each other. A group in the present case would be one or
more does or fawns. Now if assumptions 1. and 2. are true, then the distribution of the number of fawns within groups of fixed size should be binomial.
If adequate numbers of groups were observed at each given group size, then
a chi square goodness of fit test could be applied at each group size and
pooled. In the actual data, the limited frequency of groups of a given size
prohibited the use of this'test. However, Cochran (1936) presented the index of
dispersion test for such cases of small expected frequencies. The index of
dispersion test can be written as:
m

2

L

X (m-l)
i

(fi - ni p)2

= 1

ni P (1 - p)

where m is the number of groups observed, fi is the number of fawns in the
ith group, ni is the number of fawns and does in the ith group and p = L fil
L ni. The distribution of the index of dispersion is approximated by the
chi square distribution with m-l degrees of freedom. Cochran (1954) suggests
when m &gt; 15
that the mean and variance of the index of dispersion test
should be used to form a standard normal test statistic (~test).
Rejection
of a common binomial distribution for groups of all sizes can occur in the
use of the statistic even if the distribution of the fawns at a given group
size is binomial if the proportion of fawns change over or among the group
sizes.
The index of dispersion can be written in two steps to examine the alternative
just raised. First calculate:

X2

(m-K)

-

k

K

mk

l:

l:
1

i

= 1

(fki - nk Pk)
nk Pk (l-Pk)

K
L:

k

1

(mk - 1)sk2
nk Pk (l-Pk)

Where K is the number of different group sizes, nk is the size of the groups
indexed by k, mk is the number of groups of size nk, fki is the number of

�Table 1. Values for the index of dispersion
on Cache la Poudre winter range, 1962-65.

(X2m_l) test for does and fawns recorded on 10, all-day walking

routes

Seaman
Reservoir

Washout
Gulch

Sevenmile
Creek

Bennett
Creek

Kelly
Flats

E. Elkhorn
Creek

E. Hewlett
Gulch

W. Hewlett
Gulch

Youngs
Gulch

Livermore
Mountain

Index

8.36

2.22

1. 09

8.14

18.92

11.90

8.13

6.79

11. 99

12.81

df

13

2

3

26

43

27

19

14

25

25

1962 Novembe r

I

+:0)

w

1963 November

I

.44

Index
df

1

0
1

3.94

4.56

11.21

3.13

5.06

4.00

7.53

4.23

3

16

33

11

7

1

19

13

�-484-

Z

fawns in the ith group of size nk, Pk is given as Pk = i fki/mknk and sk2 is
the sample variance of the number of fawns in the sample of mk groups of
size nk. This criterion tests whether the distribution of the number of
fawns at a fixed group size is binomial. The significance levels of the
criterion can be approximated by using a chi square distribution with m-K
degrees of freedom. Since the K components of this test are independent
the lack of fit of the binomial distribution can be made at each group size
where each individual component has a chi square distribution (approximately)
with mk-1 degrees of freedom.
Secondly calculate:
K
2

X (K-1)

Z nkmk (Pk-p)2
p

(l-p)

If one does not reject the binomial distribution at each groups size, then
this criterion tests the equality of the proportion of fawns among groups
of different size. If at each group size the number of fawns is binomially
distributed, then the criterion above is approximately distributed as a chi
square distribution with K-1 degrees of freedom. If in the first step the
denominator of X2m_k is changed from nkPk (l-Pk) to nkP (l-p), then

The values of the index of dispersion test for the does and fawns recorded
on individual routes are given in Table 1. The calculations were performed
only for the 1962 and 1963 counts, since the uniformity of the rejections of
the binomial distribution clearly indicated an alternative model or set of
assumptions was needed. The component of the index of dispersion X2m-K, was
calculated for three route counts, which were selected only because they had
a fair range in group sizes to check on the possibility of changing proportions of fawns with groups size (Table 2). Again the binomial distribution
is clearly in disagreement with the data.
The next set of assumptions will serve as a basis for the analysis of the
present data set. Therefore, some practical aspects of route sampling will
be examined here, which were not mentioned when the binomial distribution
was discussed. These aspects would also be important if one had accepted the
binomial distribution for a description of the number of fawns given the
group sizes. The area of winter range covered by walking along a route for
one day is obviously limited. Hence, the area sampled has various relationships to the home range of the deer being classified. A few animals may
have their home range entirely within the area sampled while some animals
may have most of their home range within the sampled area and some may have
only a limited part of their home range inside the sampled area. The
probability of classifying a particular animal may depend on the proportion
of his home range within the area sampled and the proportion of time the
animal occupies this (included) part of the home range.

�-485-

Table 2. The component of the index of dispersion, X2 m-K, for three routes
selected because of their wide range of group sizes of does ~d fawns.

Sampling
Period

Route

2
X m-K

E(mk-l) Sk2/nkPk (l-Pk)

d.f.

Sig.
Level

1962 November

E. Elkhorn Creek

18.06

~

P - .01

1964 December

E. Elkhorn Creek

18.74

36

P - .01

1963 January

Livermore Mountain

10.62

24

P - .01

Now the primary interest in this study is with PF. Define F(t) and D(t) to
be the number of fawns and does, respectively, present in the sampled area
at time (t). Also, define PF(t) = F(t)/(F(t) + D(t».
It will be assumed
that although individual animals move into and out of the sampled area, PF(t)
will remain const~t at the value PF. In addition, given that ~ animal is
present in the area when the classification is performed, its behavior may
be such that the chance of classifying the ~imal is greater or less than the
mean probability of classifying an animal given it is present. Actual knowledge of the individual probabilities of classification are not necessary for
unbiased estimation of PF, but knowledge of the difference in the mean probability of classifying a doe versus a fawn would be necessary.
The present data does not allow estimation of the difference in the mean classification probabilities. Hence, it will be tacitly assumed that the me~ probability of classifying a fawn is equal to "the mean probability of classifying a doe.
It will be again assumed that during the count day m groups of deer are
completely classified. It will be further assumed that
fi = Sni + €i,
i

1, 2, ...

, m

where fi is the number of fawns in the ith group with ni total does ~d fawns,
S is equal to PF and €i is a random residual component. It is assumed that
the expected value of €i is zero with the variance of €i = 02 and the covariance between €i and €i~, i+i~ is zero. With the ni considered as fixed, the
model is that of a linear regression through the origin. If the distribution
of fi given ni was binomial, then the variance of €i would be ni PF (l-PF) or
proportional to ni. Examination of plots of fi versus ni, equivalently Table
3, and other calculations indicate that the variance of €i is const~t, say
02 thus providing further evidence against the binomial distribution of fi
given ni. The values of S = E fini/Eni2, &amp;2 = (Efi2 - SEfini)/(~l),
and the
st~dard error of S = v!&amp;2/Eni2 were calculated for each route at each of the
six sampling times.

I

I

A test of equality of S values among routes was performed

�Table
3 . Frequency of total groups of doe and fawn mule deer recorded on 10, all-day, walking routes on Cache 1a
Poudre winter range during each of six sampling periods, 1962-65.
Sample Dates:

Nov. 29, 1962 to Dec. 18, 1962

Sample Dates:

Jan. 3, 1963 to Jan. 27, 1963

No. Fawns
0

1

1
2
3
4
5
~ 6
I'L&lt;
7
+ 8
&lt;J)
9
8 10
• 11
.g 12
13
14
15

44
4
2

10
47
15
3

E

50

(/)

1

(/)

76

2

5
20
19
7
1
1
1

54

3

1
2
3
4
2

12

4

No. Fawns
5

6

7

E

1
1

54
56
37
23
9
5
7
4
1
2
1
5
2
0
1
207

2
1
1
2
1

1
3

7

4

1
2

3

0

1

1
2
3
4
5
6
7
8
~
I'L&lt;
9
+ 10
gj 11
o 12
~ 13
o 14
z 15
16
17
18
19
20

21
9
8
1

2
28
33
19
6
1

E

40

1

(/)

1

2

14
12
22
6
4
1
1
1

3

2
7
6
6
3

4

6

7

8

E

1
1

23
37
55
32
30
16
11
10
5
2
6
4
0
0
0
0
0
1
0
1
233

1
1
2
1
2
2

2

5

1
4
2

1

90

63

24

9

5

1

0

I

.j:'00
0I

�Table
Poudre

3,

Frequency of total groups of doe and fawn mule deer recorded on 10, all-day, walking routes on Cache 1a
range during each of six sampling periods, 1962-65 (continued).

wi.n t er

Sample Date~:

SamE1e Dates:

Nov. 21, 1963 to Dec.
14, 1963

Jan. 9, 1964 to Jan. 31, 1964

No. Fawns

No. Fawns

0

1

1
2
3
CIl
4
~
~ 5
+ 6
7
CIl
8
8 9
10
z11
12

18
2

8
27
16
3
1
1

L

21

1

Q)

2

10
16
2
1
1

3

30

5

1
2
1
3

o

56

4

7

1
1
1
1
3

1

L

26
29
26
21
5
2
1
5
0
1
1
1
118

1
2
3
4
5
6
7
8
9
CIl
10
~
~ 11
12
+ 13
CIl
~ 14
0 15

0

1

24
7
3

4
28
22
8

1

2

19
17
14
5
6

3

3
3
5
7
6
2
1

1
9
4
2
2
1

5

6

62

61

27

21

8

9

L

1
1

28
35
44
28
17
11
14
15
7
4
3
3
1
0
2
0
2
0
0
0
0
0
0
1
215

1
2
1
1
1

1

35

7

1
1

1

" 16
z 17
18
19
20
21
22
23
24
L

4

2

4

2

0

I

.j::-

ex&gt;
....•
I

�3. Frequency of total groups of doe and fawn mule deer recorded on 10, all-day, walking routes on Cache 1a
Table
Poudre winter range during each of six sampling periods, 1962-65 (continued).

~ ample Dates:

Sample Dates:

Dec. 3, 1964 to Dec. 17, 1964

Jan. 13, 1965 to Jan. 27, 1965

No. Fawns

No. Fawns

1
2
3
4
01
5
1
~ 6
&gt;x..
7
+ 8
01
9
~ 10
°ll
12
:z 13
14
15
16
L:

0

1

43
12
6
2

5
47
29
II

3
2
1

2

19
23
13
6
2
1

3

4

5

6

7

8

L:

1
1

48
59
54
36
20
13
5
8
4
4
0
0
0
0
0
1
252

4
4

3
5
1

1
1
1
3

2
1

01

63

98

64

17

6

3

0

0

1
2
3
4
5
6
01
7
8
~
~ 9
+ 10
~ll
o 12
o 13
o 14
:z 15
16
17
18
19
20
L:

0

1

18
6
3
1

3
27
26
7
5
1
1

2

12
14
4
4
2
5
1
1

3

1
4
5
3
1
2
1

4

6

7

2
1
1
2

1

5

1
1
1
1

1
1

28

71

43

17

7

1
3

1

2

L:
21
33
41
22
10
9
8
10
3
3
3
4
0
1
1
1
0
1
0
1
172

I

-!:'
00
00
I

�-489-

at each sampling time. Significant differences (P&lt;O.Ol) were noted for the
1962 fall and 1963 winter counts, while for the other four times no significant differences (P&gt;0.25) were found. Field experience indicates that the
probabilities of classifying deer appear to depend largely on environmental
conditions so that the number of groups which would be classified from day to
day may not be in a constant proportion to the number of groups present.
Although this is not a problem for sampling within a route, it is important
when route information is combined. Since the routes were not randomly
selected from the entire winter range but picked as "representative" routes
and since for two count times it is indicated that the values of PF vary
within the winter range, there is also the problem of what a combined sample
represents. However, the data will be examined as if the pooled counts of
all the groups is a random sample of all the deer in the area sampled by the
10 routes, that is, one may write
fi = Bni + E:i
for all classified groups combined with identical properties as before. The
~ ~2
~
values of B,a and the S E (B) are given in Table 4. Test of equality of PF
values may then be made among count periods. There is a significant difference
(P&lt;O.Ol) among early counts for each winter and among late counts for each
winter. Also within each winter the PF value for the early count is significantly (P&lt;0.05) larger than the PF value for the later count.
Since routes were not randomly selected, no attempt will be made to determine
adequacy of 10 routes as giving adequate precision for a PF estimate of the
entire winter range. For the pooled route data at each count time, the number
of randomly selected groups, from the composite area of the 10 routes needed
to estimate B within 5, 10, and 20 percent at the 95, 90, and 80 percent
confidence level were calculated. Since there was little variation among
count time sample sizes the mean sample sizes for the six count times are
listed below as the number of groups required to be within X percent of
PF at the (l-a) confidence level.
Confidence Level (I-a)
X

80

90

95

20
10
5

5
18
69

8
29
114

11
41
162

Statistical Model for the Observations - Quadrat Counts
The model for the observations obtained in 1974 on the 1/4-mile2 sampling units
(Table 5) is given as
i=l,

...

,m

where m is the number sampling units observed, fi is the total fawns out of
ni total does and fawns observed on the ith sampling unit. Again, the mean
and variance of f:iare assumed to be 0 and a2, respectively, and Ei &amp; Ei i+i ~

�-490-

S,

Table 4. Values of
&amp;2, SE (S), differences between early and late S values, and
tests of the equality of early and late PF's(F/(F + D»of the pooled counts of all
groups of does and fawns classified on 10, all-day walking routes on the Cache 1a
Poudre winter range, 1962-65.

Nov. -Dec.

6
n
l:X
l:X2

l:Y
l:y2

l:XY

&amp;2

1962-63
January

.4413
207
666
3,668
293
769
1,619
.2641

.4079
215
984
7,014
406
1,258
2,861
.4252

1963-64
Nov. +De c,
January

December

.4114
118
363
1,643
154
312
676
.2894

.4046
252
841
4,051
325
747
1,639
.3342

.3446
171
757
5,359
271
717
1,847
.4731

S Difference ± 95% C. L.
Year
1962-63

Nov.-Dec.

January

.44l3

.4079

+ 1.96

.0334 + .0226
1963-64

.4114

1964-65

.4046

.3742

.0304 -+ .0233

.2641 + .4252
3668
7014

.2894 + .4731
--1643
5359

.0668 -+ .0319

.3742
233
1,008
6,302
369
969
2,358
.3738

Test of Egua1ity of Pf 's, 1962-65

+ 1.96

.3446

1964-65
January

+ 1.96

j.3342 + .3738
4051

6302

Nov. -Dec.

January

577
1,870
l:X2
9,362
l:Y
772
l:y2
1,828
l:XY
3,934
ss (within)l72 .1486
"'2
Pooled a .2999
Combined
Data ss
174.8964
4.58
F2,574

619
2,749
18,675
1,046
2,944
7,066
258.1414
.4191

n
l:X

270.4602
F2 ,616 = 14.70

�Table
5. Frequency
q u ad r at.s subjectively
22, J.974 to January

of

total
groups of
assigned
to high
9, 1975.

doe and fawn mule deer counted on 106, 1/4-mi1e2
randomly-selected
and low deer density
strata,
Cache 1a Poudre winter
range,
November

High Deer_ Densi tl': guadrats

0

(fJ

§
C1:I

~
+
~
0

Q

0

:z;

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19

19
16

"

40

w

3
1

1

18
14
3
3
3
1

1

2

9
11
3
1
2
1

Low D~e~ pensi~y

N = 50

N = 56

No. Fawns

No. Fawns

3

4

3
1
1
2

1

1

1

5

6

2:

0

19
34
23
17
6
9
4
3
3
1
2
1
0
2
0
0
0
0
1
125

1

1
1
1

43

28

Quadrats

9

1

3

1
2

0
1
2
3
~
~ 4
+ 5
6
C/)
Q)
7
0
Q
8
9
0
z 10
11

11
4

2:

15

C/)

1

2

7
8
4
3

1
3
2
1
1

.

22

8

3

4

5

2:

2
1
3

11
12
11
6
4
3
2
1
.0
2
1
53

1
2
1
1

5

0

I
.p.
\0

t-'
I

�-492-

S,

are uncorrelated.
52 and the standard error of 8 were calculated for each
strata. Since proportional allocation was used (Fig. 2), the low density
strata data (Table 6, Fig. 3) was pooled and similarily for the high density
strata (Table 7, Fig. 4). Again S, 52 and the standard error of 8 was calculated for each group. The proportion of fawns was significantly higher
(P&lt;O.Ol) in the low density quadrats than in the high density quadrats. The
entire pooled sample gives 8 = .3132, 52 = 1.087 and S.E. (8) = 0.00973. In
contrast to the route samples,
is an estimate of PF for the entire winter
range. The sample size or number of quadrats required to be within X percent
of PF at (I-a) confidence levels is as follows:

6

Confidence Level (I-a)
X

80

20
10
05

90

95

5

7

17
68

28
III

10
40
158

One of the advantages of the quadrat-sample design is that IDln~mum bound
estimates of the popUlation may be made from the total deer counts on each
sampling unit using the stratified sampling estimate of the population total.
The estimates along with their standard errors for the Poudre and the Colorado State University Research Foundation Maxwell property and vicinity are
all given in Table 8.

LITERATURE CITED

Anderson, A. E., and D. E. Medin. 1965. Population density and structure.
Work Plan 4, Job 1, Job Completion Report. Pp. 405-427. In Colorado
Game, Fish a~d Parks Dept. Game Research Report, January, Part 3.
327-501 p. (Processed).
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1970. Erythrocytes and
leukocytes in a Colorado mule deer population. J. Wildl. Manage. 34(2):
389-406.
Caughley, G. 1974.
557-562.

Interpretation of age ratios.

J. Wildl. Manage. 38(3):

Cochran, W. G. 1936. The chi square distribution for binomial and Poisson
series with small expectations. Annals Eugenics 7:207-217.
1954. Some methods for strengthening the common chi square tests.
Biometrics 10:417.
Dasmann, R. F., and R. D. Taber. 1956. Determining structure in Columbian
black-tailed deer popUlations. J. Wild1. Manage. 29(1) :78-80.

�-493-

Eberhardt, L. L. 1971. Population analysis. Pp. 457-495.
In R. H. Giles,
Jr. (Ed.). Wildlife management techniques. The Wildl. Soc., Washington,
D. C. 633 p.
Hanson, W. R. 1963. Calculation of productiVity, survival and abundance of
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Kelker, G. W. 1944. Sex-ratio equations and formulas for determining wildlife
populations.
Proc. Utah Acad. Sci., Arts, and Letters 19-20:189-198.
Lauckhart, J. B. 1950. Determining the big game population from the kill.
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Leopold, A. S., T. Riney, R. McCain, and L. Tevis, Jr. 1951. The Jawbone
deer herd. California Div. Fish and Game, Game Bull. No. 4:1-139 p.
Overton, W. S. 1971. Estimating the numbers of animals in wildlife populations. Pp. 403-455.
In R. H. Giles, Jr. (Ed.). Wildlife management
techniques. The Wildlife Soc., Washington, D. C. 633 p.
Paulik, C. J., and D. S. Robson. 1969. Statistical calculations for changein-ratio estimators of population parameters.
J. Wildl. Manage. 33(1):
1-27.
Rasmussen, D. I., and E. R. Doman. 1943. Census methods and their application
in the management of mule deer. Trans. N. Amer. Wildl. Conf. 8:369-379.
Riney, T. 1956. Differences in proportions of fawns to hinds in red deer
(Cervus elaphus) from several New Zealand environments.
Nature 177:
488-489.
Robinette, W. L. 1949. Winter mortality among mule deer in the Fishlake
National Forest, Utah. U. S. Fish and Wildl. Serv., Spec. Sci. Report65. 15 p ,
Seber, G.A.F. 1973. The estimation of animal abundance and related parameters. Hafner Press, New York. 506 p.
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deer population. Arizona Game and Fish Dept. Wildl. Digest Abstr. 3:1-8.
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other important deer herds. Arizona Game and Fish Department Wildl.
Bull. No. 3:1-109.
Taber, R. D., and R. F. Dasmann. 1958. The black-tailed deer of the chaparral:
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New Mexico-Arizona Section, The Wildl. Soc., Safford, Ariz. 10 p.
Prepared by

~

£. ~
Allen E. Anderson
Wildlife Researcher

�REDFEATHER

R-2 Forest Service.

DISTRICT

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Not'l

Forest
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I

I
1

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,

Fig. 2. The location of 121 stratified (solid black-high deer density, half black-low deer density),
proportionally allocated, and randomly-selected 1/4-mi1e2 quadrats used to sample herd structure and minimum
population densities of mule deer on Cache 1a Poudre winter range and the Maxwell ranch area (upper right)
by two walking observers, 18 November 1974 to 9 January 1975.

40

I

�I'J
12j
III

B

=

0.3607

(J2

=

0.4718

SE(B)

=

0.1536

n

=

56

I

s+

G
G

~61
a:

La...

I

. 'it

VI
I

G

-

21 QQ-X
III

NO.

/

GG

0

z

I
.j:'\0

DOES

G

CSl

AND

lit

CSl
N

III

N

CSl
M

III

M

CSl

:r

FAWNS

Fig. 3. Simple linear regression of fawns pe~ group on does and fawns recorded on each of 56, randomly
selected, low deer density, 1/4-mi1e2 quadrats on Cache 1a Poudre winter range, 22 November 1974 to 9
January 1975. No deer were classified on 30 quadrats.

�A

= 0.30316
2
= 1.6879
SE(S) = 0.0133
S

a
12+

=

n

+,/+

50

+2
18

I

B

+
+

.&gt;".

U1

Z

3&amp;

a:

L..

+

2

.it
Cl
Z

+

+7

2

-

I
.po

\0
0\
I

+

+

+ -:

2

+

L. L.

+

+
lit

NO.

DOES

IS

AND

FAWNS

III

IS

'"

III

'"

IS
J'1

III

M

IS
:r

Fig. 4. Simple linear regression of fawns per group on does and fawns recorded on each of 50, randomly
selected, high deer density, 1/4-rni1e2 quadrats on Cache 1a Poudre winter range, 22 November 1974 to
9 January 1975. No deer were classified on 7 quadrats.

�-497-

Table 6. Raw data and regression computations of mule doe and fawn groups
counted on 56, randomly-selected, 1/4-mi1e2 (one-half mile per side) low
deer density quadrats on Cache 1a Poudre winter range, November 22, 1974 to
January 9, 1975.
No. Does + Fawns (X)

No. Fawns (Y)

5
10
0
0
0
0
11
12
0
3
0
0
0
22
2
0
0
8
17
3
7
5
0
0
0
0
10
3
0
0
0
0
8
0
2
15
6
3
1
0
0
0
0
0
0
0
0
11

1
4
0
0
0
0
5
3
0
1
0
0
0
8
1
0
0
3
7
1
3
3
0
0
0
0
5
1
0
0
0
0
2
0
0
2
3
2
0
0
0
0
0
0
0
0
0
4

No. Does + Fawns (X)

No. Fawns (Y)

5

3
2

4

o
o

1
1
8

4

o
o

o

o

o
o

183

68

Computations:
N

56
183
1,999
68
286
721

LX
LX2
LY
Ly2
LXY
~

f3 =

LXY

W = 0.3607

02

;=

0.4718

With Zero Deer Quadrats:
r
b
a

=

0.9343
0.3560
0.0508

Without Zero Deer Quadrats:
r
b
a

=

0.8741
0.3409
0.2158

�-498-

Table 7. Raw data and regression computations of mule deer doe and fawn
groups counted on 50, randcmly-se1ected, 1/4-mi1e2 (one-half mile per side)
high deer density quadrats on Cache 1a Poudre winter range, November 22,
1974 to December 31, 1975.
No. Does + Fawns (X)

28
37
5
11
2
8
6
5
8
0
6
2
4
19
0
3
2
0
1
8
28
5
0
8
9
1
21
21
8
11
41
19
12
9
0
2
2
6
4
5
12
32
17
5
11
2
16

No. Fawns (Y)

11
12
2
2
0
2
3
2
4
0
2
0
1
8
0
2
1
0
0
1
11
2
0
4
2
0
5
4
2
3
12
6
4
2
0
0
0
3
2
2
4
5
6
1
5
1
7

No. Does + Fawns (X)

16

No. Fawns (Y)

2

o
o

o

o

478

148

Computations:

50
478
9,480
148
954
2,874

0.30316

r
b
a

0.9167
0.2972
0.1192

1.6879

�-499-

Table 8. Estimated man1mum densities of mule deer on the Cache la Poudre
winter range and the C.S.U.R.F. Maxwell Ranch property and vicinity based
on ground counts of deer within 121, randomly-selected, proportionally
allocated, and subjectively stratified, 1/4-mile2 quadrats during the winter,
1974-75.

Strata

Total No.
Units
(1/4 mi2)

Population
Estimate

SE
(Pop. Est.)

Density/
Sq Mile

SE
Density

Cache la Poudre Winter Range
Nov. 22, 1974 to Jan. 9, 1975
Low deer density

230

946.4

162.0

16.46

2.82

High deer density

259

2168.7

244.6

33.49

3.78

Total

489

3115.1

293.4

25.48

2.40

Maxwell Ranch and Vicini t~
Nov. 18, 1974 to Nov. 21, 1974
Low deer density

30

171.9

94.8

22.92

12.64

High deer density

31

237.5

160.6

30.65

20.73

Total

61

409.4

186.5

26.85

12.23

��-501-

July, 1975

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No.

W-38-R-29

Work Plan No.
Job Title

20

Job No.

Brucellosis-Leptospirosis

Period Covered:
Personnel:

Deer-Elk Investigations
1

--------------------------------

Survey of Big Game Animals in Colorado

April 1, 1974 through March 31, 1975

William J. Adrian, Eileen Vostinak, Richard Ford, Gene Grenerd.

ABSTRACT
During the 1974 season we checked antelope for brucellosis and five species
of leptospirosis (leptospira canicola, grippotyphosa, hardjo, icterohemorrhagiae,
and pomona). We also attempted a check for bluetongue and anaplasmosis, however, the antigen was not compatible with antelope serum, and we have no results for these two diseases.

��-503-

BRUCELLOSIS-LEPTOSPIROSIS SURVEY
OF BIG GAME ANIMALS IN COLORADO
William J. Adrian

P. S. OBJECTIVE
To survey big game populations within the State of Colorado for the presence or absence of brucellosis and leptospirosis.

SEGMENT OBJECTIVE
1.

Survey Colorado's antelope populations
during the annual hunting season.

for brucellosis and leptospirosis

METHODS AND MATERIALS
Blood sample kits were mailed to antelope hunters with the request they draw
a blood sample from their kill and return the kit to the Colorado Cooperative
Brucellosis Laboratory in Denver, Colorado for processing.
A maximum of 300
sampling kits were sent to hunters in each antelope unit. If more than 300
permits were issued for one unit the names were randomly selected.
The kit consists of a cardboard mailer, blood tube and stopper, plastic bag,
rubber band, paper towel and instructions for obtaining the sample and mailing
the kits.

RESULTS AND DISCUSSION
The Wildlife Research Laboratory in cooperation with the Cooperative Brucellosis
Laboratory in Denver has, in 1974, completed its sixth annual survey for
brucellosis and leptospirosis in Colorado's big game populations.
Brucellosis is a specific contagious disease which primarily affects cattle, swine,
and goats. It can, however, affect many other animals including elk, deer,
antelope and man (undulant fever, Malta fever and Bang's Disease).
It is
caused by bacteria of the Brucella group and is characterized by abortion in the
female, and to a lesser extent, inflammation of the testes and infection of the
accessory sex glands in the male and infertility in both sexes.
Leptospirosis is a febrile (feverish) disease caused by certain species of
leptospirae.
The disease occurs with various manifestations in dogs, cattle,
pigs, sheep, goats, horses, and wildlife and is transmissable to man.
During the 1974 season we checked antelope for brucellosis and five species of
leptospirosis (leptospira canicola, grippotyphosa, hardjo, icterohemorrhagiae,
and pomona). We also attempted a check for bluetongue and anaplasmosis, however, the antigen was not compatible with antelope serum, and we have no results

�-504-

for these two diseases.
A total of 3,995 kits were sent out and we received 2,044 (51%) blood
samples, of which 1,383 were unfortunately badly hemolyzed and were
not usable. We also received 214 (5%) empty kits. Again in 1974, all
samples tested were negative, with one sample having a marginal 1-50
titer for brucellosis.

~

(

Prepared by'_-~~~~~~~~~

__~ __~~~~

Sr. Research Assistant

j

,

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                  <text>October 1975

-1-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W_-_8_8_-_R__20

_

Work Plan No.

-=l

_

Migratory Bird Investigations

1
---------=----------------------Waterfowl Production Surveys
Job Title
--------------------------------------------------------------------Period Covered:

J~

No.

May 13, 1974 to June 30, 1974

Personnel: C. Bryant and staff, Monte Vista National Wildlife Refuge; F. N.
Folks, Utah State Division of Wildlife Resources; R. Blumberg, R. Clark, R.
Desilet, W. Dolezal, H. Funk, R. Hopper, J. Lorentzson, R. Lowry, W. Russell,
G. Saville, R. Velarde, K. Wagner, R. Weldon, M. Zgainer, and M. Szymczak.

ABSTRACT
Water conditions for waterfowl production are considered good in North Park,
Brown's Park and the Cache la Poudre and Yampa River Valleys; below average
in the South Platte Valley and very poor in the San Luis Valley. The
number of estimated duck breeding pairs reached 64,484 in 1974; about 15
percent above the 1973 level and the long term average. Increases in the
number of breeding pairs were recorded in all areas except the Cache la
Poudre Valley. The mallard, gadwall, lesser scaup and blue-winged and
cinnamon teal showed significant numerical increases in comparison to 1973
levels. The post-nesting season population of Canada geese in northwest
Colorado is estimated to be 22 percent below the 1973 level, but 23 percent
above the long term average. Canada goose production in north central
Colorado is down nearly 40 percent from 1973 and long term levels, with
total numbers of geese observed down about 32 percent.

��-3-

WATERFOWL PRODUCTION SURVEYS
Mi chael R. Szymczak

P. S. OBJECTIVE

To formulate waterfowl harvest regulations for Colorado.

SEGMENT OBJECTIVES

To estimate the number of duck and goose breeding pairs, by species, in each
of Colorado's major waterfowl breeding areas.

METHODS AND MATERIALS

Present duck breeding pair and production surveys consist of a breeding pair
inventory of only major production areas.
The 1974 duck breeding pair surveys were conducted during the period of May
13 to June 30. Surveys in North Park and the Cache la Poudre and South Platte
Valleys were conducted exclusively from the air. Ground counts were made in
the Yampa Valley and Brown's Park. In the San Luis Valley intensive aerial
counts were adjusted for visibility by air-ground comparison studies. Pair
estimates for the Monte Vista National Wildlife Refuge in the San Luis Valley
were obtained from nesting transects. All survey methods and sample areas
for ducks remained the same as in previous years.
Surveys of Canada goose production were conducted within the period of May 20
to June 14. Because of mechanical difficulty goose surveys in Moffat County,
Colorado were conducted only in the Lily Park section of the Yampa River and
in the Brown's Park section of the Green River. Population estimates for
other portions of the Yampa and the Little Snake River are based on the percent
of birds in the entire Yampa-Little Snake River Complex that were located in
the Lily Park portion during the 1969 through 1973 period. The calculated
percentage is then applied to the 1974 Lily Park data to obtain an estimate
for the entire complex.
All flying was accomplished with a Cessna 185 aircraft. Areas sampled by
section or block were flown with one observer, while two observers were used
in sampling by transect.
On the basis of these studies, a report is submitted to the Bureau of Sport
Fisheries and Wildlife, which constitutes Colorado's part in the annual
continental cooperative breeding ground survey.

�-4-

RESULTS AND DISCUSSION

Water conditions for duck nesting and production are considered good in
North Park, Brown's Park and the Cache la Poudre and Yampa River Valleys.
In the South Platte Valley many small marshes and most drainage basins are
dry. Water conditions are extremely poor in the entire San Luis Valley, with
the exception of the river bottoms and adjacent sloughs. High water on
rivers in northwestern Colorado resulted in a considerable amount of flooding
of goose nes ts.
The number of estimated duck breeding pairs reached 64,484 in 1974, about 15
percent above the 1973 level and the long term average (Table 1). In comparison with 1973 totals, increases were recorded in all areas except the Cache
la Poudre Valley.
The mallard, gadwall, lesser scaup and blue-winged and cinnamon teal showed
significant numerical increases in comparison to 1973 levels. The redhead,
which had reached a record 11 percent of the breeding population in 1973,
declined nearly 6,000 pairs in 1974 (Table 2). The mallard continues to be
the major breeding species in the State, making up about 44 percent of the
population.
The post-nesting season population of Canada geese on the Yampa and Little
Snake Rivers totaled an estimated 689 birds (Table 3). On the Green River
in Brown's Park there are an estimated 326 birds (Table 3). The total
estimated gosling production in Moffat County is down about 26 percent below
the 1973 level (Table 4). The total estimated goose population is down about
22 percent below 1973, but approximately 23 percent above the long term
average (Table 5).
The results of the 1974 Canada goose production census in north central Colorado
are presented by individual area in Table 6. Gosling production on north
central Colorado trend areas is down nearly 40 percent from the 1973 and long
term levels (Table 7). All five trend areas had decreases in production and
in the total number of birds observed (Tables 7 and 8).
Fall Flight Predictions
Although the duck breeding population increased over the 1973 level, water
conditions indicate a definite decline in the fall flight. SpeCifically,
North Park, Brown's Park and the Yampa Valley should experience good duck
production and brood survival. Extensive irrigation in the Cache la Poudre
and South Platte Valleys should insure good brood survival in those areas.
But prospects in the San Luis Valley are not good, because of very poor water
condi tions.
The Canada goose flight from both northwest
be below the 1973 level.

and north central Colorado will

�-5-

Table 1. Summary of Colorado's
selected areas, 1974.

duck breeding

ground population

Total Estimated

estimates

in

1974

Breeding Pairs
Long Term
1973
Average 1:./

Percent Change
From
From Long
1973
Term Ave.

San Luis Valley

29,695

24,942

27,096

+19.1

+ 9.6

North Park 1/

16,657

14,255

15,856

+16.9

+ 5.1

South Platte Valley

8,457

7,917

6,155

+ 6.8

+37.4

Cache 1a Poudre Valley

5,713

5,852

3,244

- 2.4

+76.1

Yampa Valley

2,834

2,571

2,802

+10.2

+ 1.1

Brown's Park

1,128

632

1,024

+78.5

+10.2

Total

64,484

56,169

56,177

+14.8

+14.8

Area

1:./
San Luis Valley and North Park averages are based on results of 1964 through
1973 and 1968 through 1973 surveys respectively, because of changes in survey
methods utilized prior to those dates. Figures for other areas are 20 year
averages.

Prepared by

·_·~~1.A~·~~~.j~~~S~~~·~_!=--~~7r_7~.~~~~~~~~
_
Michael R. Szymcza_
Assistant Wildlife Researcher

�-6-

Table 2.

Species composition of Colorado's 1974 duck breeding population.

Number of Breeding Pairs
1954-731/
1974
1973
Ave rage-

Species

Percent SEecies ComEosition
1954-73
1974
1973
Average

Mallard

28,427

22,608

27,654

44.1

40.2

55.3

Blue-winged and
Cinnamon teal

11,441

6,374

4,928

17.7

11.3

9.9

Gadwall

8,465

7,400

5,243

13.1

13.2

10.5

Pintail

4,344

5,794

3,521

6.7

10.3

7.1

565

1,161

2,227

.9

2.1

4.5

Shoveler

3,918

4,556

2,302

6.1

8.1

4.6

Redhead

716

6,416

2,042

1.1

11.4

4.1

American wigeon

1,087

789

874

1.7

1.4

1.8

Other divers

5,521

1,071

1,138

8.6

1.9

2.3

Total

64,484

56,169

49,839

Green-winged

teal

1/ Species composition

computed from data from all areas for the 20 year period
regardless of changes in survey methods.

�-7Table 3.

Total estimated Canada geese, Mo ffat County, Colorado 1974.

Area

Nesting
Pairs

Non-nesting
Birds

8

99

Total
Adults

Estimatedl/
No. Goslings-

Total
Birds

115

27

142

5751/

1141/

68~/

207

119

326

Yampa River
Craig to Juniper
Springs ])
Juniper to Cross
Mountain '!:.../
Lily Park
Little Snake River-2/
Subtotal Y ampa-Li ttle
Snake
Green River
Brown's Park

30

147

Dinosaur National
Monument

l/ Calculated using average brood sizes and number of successful nests.
'!:.../

Not surveyed due

to mechanical difficulty.

1/ Figures for all areas other than Lily Park are based on the percent of
birds on the Yampa and Little Snake Rivers located in Lily Park during the
1969 through 1973 periods; that percent is then applied to the 1974 Lily Park
data to obtain an estimate for the entire complex.

�-8-

Table 4.
1974.

Estimated number of Canada goose goslings, Moffat County, Colorado,

Number of Goslings
1956-73
Average
1973
1974

Area

Yampa and Little
Snake Rivers

Percent Change
From 1956From 1973
1973 Average

114

166

217

- 31.3

- 47.5

119

151

59

- 21.2

+101. 7

105

128

317

276

- 26.5

- 15.6

Green River
Brown's Park
Dinosaur National
MonumeI'lt
Total 1:./

233

1/ Includes all areas except Dinosaur National Monument.
Table 5.

Estimated total Canada geese, Moffat County, Colorado, 1974.

Area

Yampa and Little
Snake Rivers

Estimated No. Geese
1956-73
Average
1974
1973

Percent Change
From 1956From 1973
1973 Average

689

857

672

- 19.6

+

2.5

326

437

156

- 25.4

+109.0

246

334

1,294

828

- 21.6

+ 18.4

Green River
Brown's Park
Dinosaur National
Monument
Total 1.1

1,015

Jj Includes all areas except Dinosaur National Monument.

�Table 6.

Production
Area

Wellington

Results of the northcentral

Colorado goose census June 1974.

Water Area

Terry Lake
Water Supply and Storage
No. 4
Deines Reservoir
Launer Pond
Douglas Lake
North Poudre No. 3
North Poudre No.5
Divide No. 8
Elder Reservoir
Annex No. 8
Van Sant Pond
Cobb Lake
Dale Pond
Watson Lake
Curtis Lake
Begtho1 Lake

No.
Broods

Peterson Ponds
Herring Lake
College Lake
Dean Acres
Claymore Lake
Sterling Gravel Pits
Lindenmeier Lake
Grey Lakes
Winnick Ponds
Flatiron Gravel Pits
Anderson's Pond
Timnath Reservoir
Parkwood Lake
Kitchel Lake

Total No.
Adults &amp; Yrlgs.

Total No.
of Birds

10
2

43
6

87
10

130
16

3
1
3
0
3
0
0

12
2
16
0
11
0
0
23
4
23
9
38
0
0

11
2
23
6
8
33
79
61
14
115
5
104
75
2

23
4
39
6
22
33
79
84
18
138
14
142
75
2

187

635

825

6
0
3
39
18
40
18
40
7
0
29
4
0
16

8
13
109
29
34
53
49
26
8
5
29
101
2
7

14
13
112
68
52
93
67
66
15
5
58
105
2
23

2
6
2

0
0

Total
Fort Collins

Total
No. Goslings

1
0
1
7
4
9
4

2
0
4
1
0
3

---------------------------------------------------------------------------------------------------~---------

I

\0

I

�Table 6.

Results of the northcentral

Colorado goose census, June 1974 (continued).

Production
Area

Water Area

No.
Broods

Fort Collins
(continued)

Schuelke Reservoir
\.JolaverPonds

1
1

Total
Loveland

Boedeker Reservoir
McNeil Reservoir
Flatiron Gravel Pits
Welch Reservoir

3
10
3
5

Total

Total
No. Goslings

5

Total No.
Adults &amp; Yrlgs.

Total No.
of Birds

7

4

2
2

6

229

477

706

15
43

77

92

134

177

12
20

7

21

19
41

90

239

329

144

350
101
21
25

10
4

206
80
21
13
29
13
3

206

365

571

9
2

45
6

54

I

~

o
I

Boulder

Valmont Reservoir
Terry Lake
Crystal Lake
Ish Lake
Faivre Ponds
Boulder Valley Farm Pond
Eddy Pond

21
3

15
3

Total
Denver

South Colorado Blvd. &amp; Quincy
Bl ackmer Lake
Reservoir No. 3
Denver City Park
Sloans Lake
Federal Center Pond
Claifield Reservoir
Gallup Reservoir
Columbine Country Club
Bowles Lake

12

3
2
0
0
2
11
2
7
2

o
o

44
23
7

8
21
100
202
27

53
4

21
100
149
23

44

43

87

13
23
8

76
12
166

89
35
174

-------------------------------------------------------------------------------------------------------------

�Table

6.

Resul ts of

the northcen

Production
Area

Denver
(continued)

Water

Colorado

goose

census,

No.
Broods

Area

Kings Pond
Marston Reservoir
Grant B Reservoir
Pinehurst
Country
Kendrick Lake
Mohn Estate
Standley
Lake

Total

Grand Total

tral

Club

0
0
3
4
0
2
0

June

1974

Total
No. Goslings

(continued).

Total No.
Adults &amp; Yrlgs.

Total No.
of Birds

0
0
10
13
0
6
0

90
110
88
41
11
19
4

90
110
98
54
11
25
4

185

1,004

1,189

897

2,720

3,620

I
•....
•....
I

�-12-

Table 7. Number of Canada goose goslings produced in north central Colorado
production trend areas, 1974.

Number of Goslings
1969-73
1974
1973
Average

Percent Change
From 1973
From 1969-73

Wellington

187

347

285

-46.1

-34.4

Fort Collins

229

347

283

-34.0

-19.0

Loveland

90

106

85

-15.1

+ 5.8

Boulder

206

270

244

-23.7

-15.5

Denver

185

380

300

-51.3

-38.3

Total

897

1,450

1,476

-38.1

-39.2

Area

Table 8. Number of Canada geese observed in north central Colorado production
trend areas, 1974.

Area

Number of Geese
1969-73
1974
1973
Average

Percent Change
From 1973
From 1969-73

Wellington

635

828

848

-23.3

-42.6

Fort Collins

477

843

766

-43.4

-60.7

Loveland

239

283

198

-15.5

+20. 7

Boulder

365

827

739

-55.9

-50.6

Denver

1,004

1,248

1,359

-19.6

-26.1

Total

2,720

4,029

3,9l0

-32.5

-30.4

�-13-

October 1975

JOB PROGRESS REPORT

S tate 0 f __

--=CO=L.::.O:.:RADO:..=.=-_

Project No.__

W~--.::.88~-~R~-~2~0=__
_

Work Plan No.

-------------

Job Title-

1

Migratory
Job No.

Bird Investigations
~2=__

~T~r~a~p~p~l~·n~g~a~n~d_B==an~d=i:.:n~g~D~u~c~k~s~an==d~G~e~e=s~e~

Period Covered:

_
_

April 1, 1974 to March 31, 1975

Personnel:

C. Bryant and Staff of Alamosa and Monte Vista National Wildlife
Refuges, S. Bissell, R. Blumberg, J. Carsella, G. Claassen, J.
Corey, D. Coven, C. Crawford, M. DePra, R. Desilet, K. Dillinger, W. Dolezal,
G. East, G. Eyre, H. Funk, J. Gray, A. Hemmert, J. Hicks, R. Holder, J.
Lorentzson, R. Mason, J. Miller, K. Miller, W. Mink, R. Oakleaf, J. Olterman,
D. Owens, B. Peterson, S. Porter, F. Rinella, Jr., R. Rouse, H. Spear, S.
Steinert, M. Stone, M. Szymczak, J. Wagner, K. Wagner, C. Wetherill, and
R. Hopper.
ABSTRACT
Trapping efforts in Segment 20 resulted in the banding of 17,806 ducks of
nine species at eight locations in Colorado. The major species in the banded
sample was the mallard, which contributed 11,701, or about 66 percent of the
total. Pre-season banding in the high country yielded 11,185 ducks, including
3,590 in North Park, 2,666 in South Park, and 4,924 in the San Luis Valley.
Mallards and pintails were the dominant species in the samples in these three
areas. Post-season banding produced a banded sample of 8,416 mallards,
including 6,558 in eastern Colorado (Central Flyway) and 1,858 in west central
Colorado (Pacific Flyway). Goose trapping- activities resulted in a banded
sample of 1,946 Canada geese in Segment 20. Included in this total were
200 goslings transplanted to new production sites in South Park and the South
Platte Valley. Summer banding on established production areas in the San Luis
Valley and in north central Colorado accounted for 1,077. The remainder were
banded post-season in the Cache la Poudre and South Platte Valleys (241) and
the Arkansas Valley (428).

��-15-

TRAPPING AND BANDING DUCKS AND GEESE
Richard M. Hopper

This report summarizes all waterfowl banding activities conducted under
Federal Aid Project W-88-R-20 for the segment year April 1, 1974 to March
31, 1975. Band recovery data are analyzed as part of other jobs (Work Plan
1, Job 3; Work Plan 2, Job 6; and Work Plan 3, Job 6) and thus, will not be
included in this report. This report merely presents a listing of numbers
of ducks and geese banded by species and location during the segment year.
P. S. OBJECTIVE
To formulate waterfowl harves t regulations for Colorado.

SEGMENT OBJECTIVES
1.

To provide adequate samples of banded ducks and geese in Colorado's major
breeding and harvest areas for the purpose of accumulating migration and
annual mortality data.

2.

To report species and numbers of mallards winter banded in the South
Platte Valley, Arkansas Valley, and Bonny Reservoir as part of Work Plan
3, Job 6.

3.

To report transplant geese banded under Work Plan 2, Job 2.

4.

Prepare progress report.
METHODS AND MATERIALS

Three major phases of trapping and banding operations were continued in
Segment 20 as in recent years. These included: (1) mf.d+aummer banding of
Canada geese, (2) late sUlIUD.er
(pre-season) duck banding, and (3) winter (postseason) duck and goose banding. Goslings for transplanting were hand-reared
and also obtained from the wild by drive-trapping prior to the time the birds
were capable of flight. Molting adults and their flightless goslings were
captured and released on north central Colorado production areas by drivetrapping. Pre-season banding of ducks was conducted in August and September
using baited Salt Plains traps (cage type) to capture birds in North Park,
South Park, and the San Luis Valley. Night~lighting was again employed
experimentally to capture ducks in North Park during the pre-season period.
Post-season banding (January-February) utilized baited Salt Plains traps
and cannon-nets for duck trapping, and baited cannon-nets for goose trapping.

�-16-

RESULTS AND DISCUSSION

Ducks

Nearly 18,000 ducks of nine species were banded in Segment 20 (Table 1).
The South Platte Valley produced the largest sample (3,750) of the eight
banding locations in Colorado, followed by North Park (3,590), the San Luis
Valley (3,134), South Park (2,666), and the Grand Junction-Delta area of the
western slope (1,858).
The mallard continued to dominate the banded sample, compr1s1ng 11,701, or
about 66 percent of the total. Pintails and green-winged teals were the
next most important contributors to the banded sample with 3,736 and 1,615,
respectively.

High Country Study

Objectives of the high country study of duck populations are presented by
Hopper (1972). Species composition by age and sex for 11,185 ducks banded
in the three high country areas during 1974 is shown in Tables 2-5. San
Luis Valley bandings were accomplished by both State and Federal personnel
(Table 4). The U. S. Fish and Wildlife Service continued to cooperate in
this study by banding a portion of the Valley quota on the Monte Vista and
Alamosa National Wildlife Refuges.
Mallards and pintails were again the main species in the banded samples in
all three high country areas (Tables 2-4). They contributed 3,986 and 4,084
birds, respectively, for all three areas combined (Table 5). The only other
species for which significant banded samples were obtained included greenwinged teals (1,928) and blue-winged and cinnamon teals (1,010).

Winter Banding

Post-season banding of terminal wintering populations of mallards was continued in eastern Colorado for the twelfth consecutive year under Work Plan 3,
Job 6. The total banded sample amounted to 6,558 mallards for this period
in Segment 20 (Table 1). A detailed breakdown of this sample by age, sex
and location, as well as a discussion of other aspects of the study, is
presented in the Job Progress Report for Work Plan 3, Job 6 (Investigation
of Mallard Management Units of Eastern Colorado).
An additional 1,858 mallards were banded on the Western Slope of Colorado
during Segment 20 (Table 1). This was the second consecutive year of banding
in this area as part of a study described by Hopper (1974) in the previous
Job Progress Report for Work Plan 1, Job 2. Age and sex ratios obtained in
this year's banded sample are shown in Table 6 by location.

�Table 1.

Number of ducks banded by species,

location and period of year 1974-75.

Number of Ducks Banded
Pos t-season 27
Cache la
So. Platte
Bonny
Valley
Poudre Valley
Reservoir

North
Park

Pre-season I7
South
San Luis
Park
Valley

Hallard

1,304

992

989

825

3,750

993

Gadwall

15

1

8

-

-

American Wigeon

23

-

-

-

Green-winged

367

753

495

-

Species

Teal

Blue-winged and
Cinnamon Teal

West 3/
Slope-

Total

990

1,858

11,701

-

-

-

24

-

-

-

-

23

-

-

-

-

1,615
I
•....•

-

-

597

-

3,736

-

-

109

-

-

-

1

993

990

1,858

17,806

81

334

182

-

-

-

Pintail

1,779

580

1,377

-

-

Redhead

21

5

83

-

-

-

Shoveler

-

1

-

-

-

3,590

2,666

3,134

825

3,750

Total

Arkansas
Valley

1/ August-September.
'1:/ January-February.
1/ Colorado-Gunnison-Uncompahgre

River Complex.

"
I

�-18Table

2.

Sex and age composition

of ducks banded

pre-season

in North

Park,

1974.

Species

AM

Number Banded
Age and Sex
IM
AF
IF

Mallard

317

362

227

398

1,304

Gadwall

2

4

6

3

15

0

13

1

9

23

119

118

50

80

367

3

39

1

38

81

Pintail

441

482

389

467

1,779

Redhead

13

1

4

3

21

Total

895

1,019

678

998

3,590

pre-season

in South

American

Wigeon

Green-winged

Teal

Blue-winged

and Cinnamon Teal

Table

3.

Sex and age composition

of ducks banded

Total

Park,

Species

AM

Number Banded
and
Sex
Ase
,1M
AF
IF

Mallard

325

260

257

150

992

Gadwall

1

0

0

0

1

Total

Green-winged

Teal

339

226

55

133

753

Blue-winged

and Cinnamon Teal

47

162

14

111

334

0

0

0

1

1

Pintail

156

188

87

149

580

Redhead

1

0

4

0

5

869

836

417

544

2,666

Shoveler

Total

1974.

�-19-

Table 4. Age and sex composition of ducks banded pre-season in the San
Luis Valley, 1974. !I

AM

Number of Ducks Banded
Age and Sex
1M
AF
IF

Total

Mallard

324

484

404

478

1,690

Gadwall

0

2

4

10

16

Green-winged Teal

360

223

69

155

807

Blue-winged and Cinnamon Teal

57

291

24

223

595

Pintail

390

597

248

490

1,725

Redhead

36

5

42

8

91

1,167

1,602

791

1,364

4,924

Species

Total

!/ Includes ducks banded by Federal personnel on Alamosa
National Wildlife Refuges as part of high-country

and Monte Vista
duck study.

Table 5. Sex and age composition of ducks banded pre-season in the three
high country areas combined, 1974. 1/

Number Banded
Age and Sex
1M
IF
AF

Species

AM

Mallard

966

1,106

888

Gadwall

3

6

American Wigeon

0

Green-winged Teal
Blue-winged and Cinnamon Teal

Unk.

Total

1,026

0

3,986

10

13

0

32

14

1

12

0

27

818

567

174

368

1

1,928

107

492

39

372

0

1,010

Shoveler

0

0

0

1

0

1

Pintail

987

1,267

724

1,106

0

4,084

Redhead

50

6

50

11

0

117

2,931

3,458

1,886

2,909

1

11 ,185

Total

1/ Includes 1,795 ducks banded by Federal personnel on Alamosa and Monte Vista
Wildlife Refuges in the San Luis Valley as part of the high country duck study.

�-20-

Table 6. Age and sex composition of mallards banded post-season in west
central Colorado, 1974-75.

AM

Number Banded
Age and Sex
IF
1M
AF

Total

Grand Junction-Mack

223

265

131

447

1,066

Delta

167

276

67

2~

792

Total

390

541

198

729

1,858

Location

Geese
Nearly 1,950 Canada geese were banded during Segment 20 (Table 7). The Cache
1a Poudre and South Platte Valleys accounted for the largest number with
1,408, while the Arkansas Valley and South Park followed with 428 and 100,
respectively.

Summer Transplants
Two hundred Canada goose goslings were banded and transported from Front
Range production areas and released on two transplant sites having promise
as future production areas (Table 8). These two sites, Antero Reservoir in
South Park and Prewitt Reservoir in the South Platte Valley each received
100 goslings. The number of males and females in each group of transplants
was nearly equal.
The 1974 releases marked the end of the Canada goose transplant program as
part of Federal Aid Project W-88-R. This program, involving annual releases
of goslings since 1957, resulted in the successful establishment of breeding
flocks of Canada geese in many areas of Colorado. The transplant program
is discussed in detail under Work Plan 2, Job 2.
Pre-season Banding on Colorado Production Areas
A study was initiated in 1974 to investigate the status of breeding populations
of Canada geese in the San Luis Valley and the Fort Co11ins-Bou1der-Denver
area of north central Colorado. Pre-season banding is a major component of
this study. Details of the overall investigation are discussed under Work
Plan 2, Job 6.

�-21-

Table 7.
1974-75.

Number of Canada geese banded by location and period of year,

Number Banded
Location

Sunnner
Transplants.V

Arkansas Valley

Pre-se asony

Pos t-se ason]:./ Total

0

0

428

428

Cache la Poudre and South
Platte Valleys

100

1,067

241

1,408

South Park

100

0

0

100

0

10

0

10

200

1,077

669

1,946

San Luis Valley

Total

Y June-July.
~/ January-February.
Trapping efforts proved unsuccessful in the San Luis Valley for the first
year's attempt, with only 10 geese being banded. The banding crew was late
in reaching the Valley and most geese were already flying by the time trapping
efforts were initiated. More time will be allocated for this endeavor next
segment.
Pre-season trapping operations were highly successful in north central Colorado and resulted in the banding of 1,067 adult and young geese (Table 9).
Four hundred of these birds were also color-marked with black on orange neck
collars for studies of goose distribution as part of the overall investigation.
Pre-season Banding on Alberta Production Areas
The Colorado Division of Wildlife assisted in a Central Flyway and Pacific
Flyway cooperative banding study of Canada geese on Canadian production areas
in southern Alberta. The major purpose of this study is to define the
breeding areas of the Great Basin and Hi-line Canada Goose Populations in
Alberta. This cooperative venture, irtvolving personnel from Colorado, Idaho,
Utah, Wyoming, Alberta, the U. S. Fish and Wildlife Service, and the Canadian
Wildlife Service, resulted in the banding of 1,346 Canada geese in a 10 day
period in June and July.

�-22-

Table 8.

Sex composition of goslings banded for summer transplants, 1974.

Release Site

South Park (Antero

Male

Reservoir)

Number Banded
Sex
Total
Female

48

52

100

South Platte Valley (Prewitt Reservoir)

55

45

100

Total

103

97

200

Table 9. Age and sex composition of Canada geese banded pre-season in north
central Colorado, 1974.

Age and Sex

Number Banded

Adult Male

393

Local Male

158

Adult Female

358

Local Female

157

Unknown

Total

1

1,067

Winter Banding
Post-season activities produced 669 banded Canada geese in the Cache la
Poudre and South Platte Valleys (241) and Arkansas Valley (428) (Table 10).
Overall sample sizes and age and sex ratios in the banded sample were not
satisfactory for either of the two banding locations.

�-23-

Table 10. Age and sex composition of Canada geese banded post-season in
eastern Colorado, 1974-75.

Number Banded
Age and Sex
AF
IF
UF

Location

AM

1M

Cache la Poudre and South
Platte Valleys

97

24

28

3

89

241

Arkans as Vall ey

193

31

164

40

0

428

Total

290

55

192

43

89

669

Total

LITERATURE CITED

Hopper, R. M. 1972. Trapping and banding ducks and geese.
Fed. Aid Game Res. Rept., Oct. pp. 13-21.

Colo. Div. Wildl.,

Hopper, R. M. 1974. Trapping and banding ducks and geese.
Wildl., Fed. Aid Game Res. Rept., Oct. pp.13-22.

Colo. Div.

Prepared by

.~da~

.chard M. Hopper
Wildlife Researcher

~'&lt;""L
0&gt;'

��-25JOB PROGRESS

October
REPORT

State of

C.::;,O::.:L:::::O::.:RADO=:::::..
_

Project No.__

~W_-~8~8_-~R~-=2~0 _

W~rk Plan No.
Job Title

Analysis

Period Covered:
Personnel:

=l

_
of Waterfowl

1975

Migratory
Job No.

Bird Investigations
~3~

_

Banding Data

March 1, 1975 to March 31, 1975

J. Gustafson

and M. Szymczak.

ABSTRACT

First year recovery rates for mallards (Arras platyrhynchos) and pintails (Arras
acuta) banded during August and September in the San Luis Valley and South
Park from 1968 through 1973 and in North Park from 1971 through 1973 are
presented.
Rates are categorized by age and sex for each species.
During
the 1975 banding season, 300 mallards of each age and sex grouping, excluding
locals, will be banded in the three areas. All pintai1s captured during the
1975 banding period will be banded.

�-26RECOMMENDATIONS

1.

Band 300 mallards in each of the following categories in each of the
three high mountain areas during the pre-season banding period:
immature males, adult males, immature females, adult females. Also
band all local mallards that are captured.

2.

Band every pintail, green-winged teal (Anas carolinensis), blue winged
teal (Anas discors), cinnamon teal (Ana~anoPtera),
gadwall (Anas
strepera), widgeon (Anas americana), redhead (Aythya americana), and
canvas back (Aythya valisineria) that are captured during the pre-season
banding period.

�-27-

ANALYSIS OF WATERFOWL BANDING DATA
Michael R. Szymczak

P. S. OBJEcrIVE
To formulate waterfowl harvest regulations for Colorado.

SEGMENT OBJECTIVE
To investigate migration, mortality, recovery distribution and relationships
among populations of ducks banded in Colorado's high country, specifically
North Park, South Park, and the San Luis Valley.

METHODS AND MATERIALS
All recoveries of mallards and pintails banded in South Park and the San Luis
Valley since 1968 and in North Park since 1971 and reported recovered through
the 1973-74 hunting season were sorted by age and sex. First year recovery
rates were calculated for all groupings.

RESULTS AND DISCUSSION
Because of the small number of ducks other than mallards or pintails which
have been banded in the high mountai~area,
first year recoveries for only
those two species were calculated. As many individuals as possible of the
other target species should be banded in all three banding areas.
The first year recovery rates for mallards for 1968 through 1973 in South
Park and the San Luis Valley and for 1971 through 1973 for North Park are
presented by age and sex in Tables 1 through 4. Because of small banded
sample size, first year recoveries of locals were not tabulated. First year
recovery rates in the three areas are generally quite variable from year to
year in any age or sex class.
When the original high mountain banding program was designed, the objective
was to band in the three areas for five consecutive years. Original quotas
for mallards were at least 250 of each age and sex grouping.
Quotas were
generally met in both North Park and the San Luis Valley. After reviewing the
number of first year recoveries from 1971 bandings, the quotas were increased
to 300 mallards of each age and sex grouping. The intent was to strive to
obtain 50 first year recoveries for each banding cohort. After reviewing the
results of three years of mallard banding in the three areas, we find that
acquiring 50 first year recoveries would require an increased effort in all

�-28-

three areas and may be logistically impossible, particularly for females.
Therefore, during the fifth year of banding, summer of 1975, the banding
quotas for each age and sex grouping of mallards will remain at 300 in
order to maintain a numerically consistent series of bandings.
Complete
analysis of these bandings and resulting recoveries after the 1975-76
hunting season using methods which will provide confidence intervals for
calculated mortality rates will give us a better indication of the appropriate
sample sizes for bandings in these areas.
The first year recovery rates for pintail banding in the three high mountain
park areas are presented by age and sex in Tables 5 through 8. With few
exceptions, the recovery rates of pintails of any age or sex banded in any
area are extremely low. In 1971, a general quota of 200 to 250 bandings
were established for each age and sex grouping. In 1972 the quota was
increased to 250. In 1973 the decision was made to eliminate quotas and band
every pintail captured during the entire banding period. That policy will
be continued through the 1975 banding season.

Prepared by

/ll~K~
Michael
SzymnG
R.

Assistant Wildlife Researcher

�Table 1. First year recovery rates for adult male mallards banded in North Park, South Park and the San Luis
Valley, 1968-73.

Recovery
Rate

Number
Banded

South Park
No. First Year
Recoveries

Recovery
Rate

Number
Banded

San Luis Valley
Recovery
No. First Year
Rate
Recoveries

Year
Banded

Number
Banded

North Park
No. First Year
Recoveries

1968

-

-

-

74

1

0.014

1,077

58

0.054

1969

-

-

-

339

13

0.038

1,250

102

0.082

1970

-

-

-

210

15

0.071

939

98

0.104

411

18

0.044

312

21

0.067

1971

245

9

0.037

1972

262

17

0.065

250

8

0.032

262

23

0.088

1973

315

16

0.051

250

15

0.060

444

19

0.043

Total

822

42

0.051

1,534

70

0.046

4,284

321

0.075

I
N
\0
I

�Table 2. First year recovery
Valley, 1968-73.

rates for immature male mallards banded

in North Park, South Park, and the San Luis

Number
Banded

South Park
No. First Year
Recoveries

Re&lt;":overy
Rate

Number
Banded

San Luis Va11eI
Recovery
No. First Year
Rate
Recoveries

Year
Banded

Number
Banded

North Park
No. First Year
Recoveries

-

-

-

111

3

0.027

1,087

85

0.078

1968

-

-

-

235

15

0.064

1,090

116

0.106

1969

-

-

-

235

17

0.072

877

93

0.106

1970

203

5

0.025

352

39

0.111

Recovery
Rate

I

1971

292

12

0.041

1972

285

12

0.042

209

12

0.057

229

19

0.083

1973

313

16

0.051

190

13

0.068

370

34

0.092

Total

890

40

0.045

1,183

65

0.055

4,005

386

0.096

w
0
I

�Table 3. First year recovery rates for adult female mallards banded in North Park, South Park, and the San Luis
Valley, 1968-73.
North Park
No. First Year
Recoveries

Number
Banded

South Park
No. First Year
Recoveries

Recovery
Rate

Number
Banded

42

0

0.000

1,281

48

0.037

151

2

0.013

1,238

54

0.044

171

5

0.029

1,148

39

0.034

373

8

0.021

305

9

0.030

0.032

285

4

0.014

Year
Banded

Number
Banded

1968

-

-

-

237

10

0.042

1969
1970
1971

-

San Luis Valle:i
Recovery
No. First Year
Rate
Recoveries

Recovery
Rate

.I

w
I

1972

285

5

0.018

250

8

1973

254

9

0.035

192

4

0.021

466

8

0.017

Total

776

24

0.031

1,179

27

0.023

4,723

162

0.034

�Table 4. First year recovery rates for immature female mallards banded in North Park, South Park, and the San
Luis Valley, 1968-73.

Year
Banded

Number
Banded

1968

-

1969

North Park
No. First Year
Recoveries

Recovery
Rate

South Park
NtUllber No. First Year
Recoveries
Banded

Recovery
Rate

San Luis Va11el
Recovery
NtUllber No. First Year
Rate
Recoveries
Banded

-

72

4

0.056

896

34

0.038

-

-

-

100

0

0.000

970

36

0.037

1970

-

-

-

151

5

0.033

881

67

0.076

1971

291

10

0.034

92

4

0.043

252

10

0.040

1972

253

16

0.063

159

7

0.044

196

18

0.092

1973

256

9

0.035

113

3

0.027

381

24

0.063

Total

800

35

0.044

687

23

0.033

3,576

189

0.053

I

w

N
I

�Table 5. First year recovery rates for adult male pintai1s banded in North Park, South Park, and the San Luis
Valley, 1968-73.

Number
Banded

South Park
No. First Year
Recoveries

Recovery
Rate

Number
Banded

San Luis Vallez:
Recovery
No. First Year
Rate
Recoveries

Year
Banded

Number
Banded

North Park
No. First Year
Recoveries

1968

-

-

-

155

3

0.019

207

3

0.014

1969

-

-

-

82 •

1

0.012

57

0

0.000

1970

-

-

-

97

2

0.021

35

5

0.143

70

0

0.000

359

6

0.017

Recovery
Rate

I

1971

205

2

0.010

1972

280

4

0.014

150

1

0.007

280

1

0.004

1973

791

8

0.010

186

4

0.022

375

3

0.008

Total

1,276

14

0.011

740

11

0.015

1,313

18

0.014

w
w
I

�Table 6. First year recovery rates for immature male pintai1s banded in North Park, South Park, and the San
Luis Valley, 1968-73.

Year
Banded

Number
Banded

North Park
No. First Year
Recoveries

1968

-

-

1969

-

1970

-

South Park
No. First Year
Recoveries

San Luis Valley
Recovery
No. First Year
Rate
Recoveries

Recovery
Rate

Number
Banded

1

0.015

224

11

0.049

80

0

0.000

77

2

0.026

76

2

0.026

73

7

0.096

286

7

0.024

Recovery
Rate

Number
Banded
67

-

-

-

-

I

1971

230

1

0.004

72

1

0.014

1972

270

7

0.026

125

1

0.008

264

11

0.042

1973

201

8

0.040

123

0

0.000

350

6

0.017

Total

701

16

0.023

543

5

0.009

1,274

44

0.035

w

~
I

�Table 7. First year recovery rat~s for adult female pintails banded in North Park, South Park, and the San
Luis Valley, 1968-73.
______~N~o~r~t~h
Park
Year Number No. First Year
Banded Banded
Recoveries

Recovery
Rate

Number
Banded

_
South Park
No. First Year
Recoveries

Recovery
Rate

Number
Banded

San Luis Valley
No. First Year
Recovery
Recoveries
Rate

1968

97

1

0.010

204

2

0.010

1969

76

1

0.013

120

1

0.008

1970

25

2

0.080

106

9

0.085
I

1971

223

1

0.004

28

0

0.000

346

2

0.006

1972

273

2

0.007

125

1

0.008

273

6

0.022

1973

462

7

0.015

147

2

0.014

429

5

0.012

498

7

0.014

1,478

25

0.017

----."

Total

---" -"--~--~--""-~-~~~-~~~~--"

958

10

0.010

~--~--~"~-,~~~~"

,~."&lt;--~~--~~-.--~~-"~--~"--~"'"~-~---~.~
-~.--~-~--~~--.--,---

w

\JI

I

�Table 8. First year recovery rates for immature female pintai1s banded in North Park, South Park, and the San
Luis Valley, 1968-73.

Year
Banded

Number
Banded

North Park
No. First Year
Recoveries

Recovery
Rate

Number
Banded

South Park
No. First Year
Recoveries

Recovery
Rate

Number
Banded

San Luis Va11eI
Recovery
No. First Year
Rate
Recoveries

1968

-

-

-

52

0

0.000

230

4

0.017

1969

-

-

-

75

1

0.013

133

5

0.038

1970

-

-

-

33

1

0.030

111

5

0.045

1971

201

2

0.010

74

0

0.000

258

5

0.019

1972

263

4

0.015

121

1

0.008

266

14

0.053

1973

202

8

0.040

109

0

0.000

329

5

0.015

Total

666

14

0.021

464

3

0.006

1,327

38

0.029

I

w
0'
I

�October 1975

-37JOB FINAL REPORT

State of
Project

C..:..O..:..I:.::.O..:..RA=D:.,..O:...._
W-88-R-20

No.

Migratory

1

Work Plan No.

Job No.

Job Title: San Luis Valley Cooperative
Period Covered:
Personnel:

Bird Investigations
12

Mallard Investigation

April 1, 1964 to March 31, 1975

Richard M. Hopper

ABSTRACT
The final report covering all segment objectives of this job was submitted
to Wildlife Monographs in November 1974. It was reviewed by an editorial
staff and returned to the authors in March 1975. At this time it was
considered to be acceptable for publication as a Wildlife Monograph, following
some r eva.sa.ons , The title of this paper is "Exper·imental Duck Hunting
Seasons -- San Luis Valley, Colorado, 1963-1970", with Richard M. Hopper,
Aelred D. Geis, Jack R. Grieb, and Lewis Nelson, Jr. as authors.
Additional time needed for revisions, editing, and proofing, as required
of the authors by Wildlife Monographs, will be accomplished in the next year
under Work Plan 6, Job 1 (W-88-R).

Prepared by

·Zwl~/~%
~'" " "
Richard M. Hopper'VGff
t.

o

Wildlife

Researcher

��October 1975

-39-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W~-_8~8-~R~-~2~0~

Work Plan No.

_

Migratory Bird Investigations

Job Title

-=-=2_____________
Job No.
~2~
Experimental Studies on Improving
the Status of Canada Goose Populations

Period Covered:

April 2, 1974 to February 17, 1975

Personnel:

_

S. Bissell, M. DePra, W. Dolezal, J. Hicks, R. Holder, T. Lines,
J. Lorentzson, W. Olmstead, S. Porter, S. Steinert, K. Wagner,
J. Wagner, L. Webster, and M. Szymczak.

ABSTRACT
Canada geese (Branta canadensis) were reported to be nesting in the Jumbo
Reservoir area in 1974. Reported hunting mortality of Canada geese released
at Jumbo Reservoir Annex has been comparatively light through the 1973-74
hunting season, with only 16 of 303 birds recovered. Band recoveries indicate that a portion of the birds released at Jumbo in 1972 have drifted east
of Colorado and are following a north-south migration pattern through the
central portion of the eastern states of the Central Flyway. About 10
percent of the 166 birds released at Prewitt Reservoir through the summer of
1973 were reported recovered by hunters through the 1973-74 hunting season.
An additional 100 goslings were released at Prewitt Reservoir on July 5,
1974. Canada geese began nesting at Lake John Annex in North Park as early
as March 9, 1974. A total of nine known nests were established at three
locations in North Park in 1974, however, only two of the nests hatched.
The six nests established at Lake John Annex were all destroyed by predators.
According to band recoveries, a portion of geese released on Walden Reservoir
in North Park in 1973 migrated into the Texas Panhandle in the fall of 1973.
Over 45 percent of the direct recoveries of all birds released in North Park
were reported taken in Arizona, while extreme southern California accounted
for 19 percent of the direct recoveries. Excluding Mexico, 78.7 percent of
the direct recoveries of birds released in North Park came from Pacific
Flyway areas. Colorado accounted for 29.4 percent of the indirect recoveries
.of geese released in North Park, while 26.5 and 17.6 percent of the indirect
recoveries came from California and Arizona, respectively.
Excluding Mexico,
63.6 percent of the indirect recoveries of North Park geese were taken in
the Pacific Flyway. Through the 1973-74 hunting season, six of the 156
Canada goose goslings released at Antero Reservoir had been reported
recovered. The largest number of Canada geese observed at Antero Reservoir
in the spring of 1974 was 120 birds sighted in mid-April. At least 40
goslings were produced at Antero Reservoir in 1974.

��-41-

EXPERIMENTAL STUDIES ON IMPROVING
THE STATUS OF CANADA GOOSE POPULATIONS
Michael R. Szymczak

P. S. OBJECTIVE
To investigate the success of attempts to establish breeding populations of
Canada. geese in suitable habitat where they do not presently exist in
Colorado.

SEGMENT OBJECTIVES
1.

Edit and publish a technical bulletin on the technique of establishing
a breeding population of Canada geese.

2.

To expand the breeding flocks of Canada geese in the South Platte Valley
and South Park.

3.

Prepare progress report.

METHODS AND MATERIALS
Canada goose goslings for transplant purposes were obtained from two different
sources: (1) goslings from the metropolitan Denver area, and (2) goslings and
some eggs from the Fort Collins area. All eggs were transported to the Fort
Collins Wildlife Research Station for incubation and subsequent raising.
All wild trapped goslings were held for variable periods of time before
release on the restoration areas. A mixture of hand reared goslings and wild
trapped goslings were released on each area.
An attempt was made to release an equal number of birds of each sex at each
location. Each bird was marked with a U. S. Fish and Wildlife Service leg
band.
Migration and mortality information was obtained from computer printouts
received from the Migratory Bird Population Station which present a list
of reported encounters with banded birds.

RESULTS AND DISCUSSION
Preparation of Technical Bulletin
Preparation of the bulletin is completed and is now in the final stages of
review.

�-42-

South Platte Valley-Jumbo Reservoir
By July 1973, a total of 303 Canada goose goslings had been released at
Jumbo Reservoir Annex located near Sedgwick along the South Platte River in
extreme northeast Colorado. In the spring of 1973 at least 12 nests were
established in the Jumbo Reservoir area (Szymczak 1974). No counts were
obtained of the number of nests established in the area in the spring of
1974, but sporadic reports indicated that birds were actively nesting.
Surveys of nesting birds in the Jumbo area are scheduled for spring 1975.
Reported recoveries of birds released at Jumbo Reservoir Annex indicate they
are being subjected to comparatively light hunting mortality, at least
through the 1973-74 hunting season (Table 1). The group released in 1972
has undergone the heaviest mortality with nine percent of the birds being
recovered during a two year period. Only six percent of the birds released
in 1971 had been recovered through three hunting seasons, while only a
single recovery was reported during the 1973-74 season of the 101 goslings
released in 1973. An area of about 250 square miles remains closed around
the release site.
The distribution of recoveries of birds released at Jumbo Reservoir Annex
does not indicate any specific migration patterns (Table 1). Indirect
recoveries of birds released in 1971 and 1972 indicate migration of at
least a portion of the birds away from the release site. Apparently, at
least a portion of the birds released in 1972 have drifted east of Colorado
and are following a north-south migration pattern through the central portion
of the eastern states of the Central Flyway. In addition to the indirect
recoveries of 1972 released birds, another bird was recaptured in June,
1973 in Kidder County, central North Dakota.

South Platte Valley-Prewitt

Reservoir

By late summer 1973, 166 Canada goose goslings had been released on small
ponds directly below the Prewitt Reservoir dam. One nest had been established
in the release area in the spring of 1973. Extensive surveys for locating
nesting geese were not conducted in the release area in 1974. Surveys of
nesting birds in the Prewitt Reservoir area are scheduled for spring 1975.
Through the 1973-74 hunting season about 10 percent of the birds released
at Prewitt have been reported recovered by hunters (Table 2). All but one
of the ed ght total recoveries have reportedly occurred in Colorado (Table 2).
Five of the birds have been recovered near the release site in Washington
and Morgan counties. An area of approximately 100 square miles around the
release site remains closed to goose hunting.
On July 5, 1974, 55 male and 45 female Canada goose goslings were released
on the ponds below the Prewitt Reservoir. This release brings the total
number of birds released at that location in three years to 266.

�-43-

Table l. Hunting season recovery locations of Canada geese released at
Jumbo Reservoir Annex, near Sedgwick in extreme northeastern Colorado.

Year Released
Number Released

1971
102

1972
100

1973
101

Area Recovered

No. Recovered
Direct
Indirect

No. Recovered
Direct
Indirect

No. Recovered
Indirect
Direct

Colorado
Logan County
Ot ero Coun ty
Baca County

1
0
0

0
0
0

0
0
1

0
0
1

0
1
0

Nebraska
Chase County
Keith County

0
0

1
1

0
0

0
0

0
0

W;yoming
Goshen County

0

1

0

0

0

Kansas
Pawnee County

0

0

0

1

0

Texas
Hockley County
Uvalde County
Zavala County

0
0
0

0
0
0

1
1
2

0
0
0

0
0
0

South Dakota
Buffalo County

0

0

0

1

0

North Dakota
McHen ry Coun ty

0

0

0

1

0

Minnesota
Chippewa County

0

1

0

0

0

Albertaz Canada
Beaverhill Lake
Area

0

1

0

0

0

Total

1

5

5

4

1

�-44-

Table 2. Hunting season recovery locations of Canada geese released at
Prewitt Reservoir in Washington County, northeastern Colorado.

Year Released
Number Released

1972
70

1973
96

Area Recovered

No. Recovered
Direct
Indirect

No. Recovered
Indirect
Direct

Colorado
Washington County
Mor gan Coun ty
o tero Coun ty
Bent County

o
o

1

o

Texas
--Randall County

1

o

o

Total

5

2

1

3

o

1

1

o
o
1

North Park
Spring Returns
A total of 931 Canada goose goslings were released in North Park during the
five year period, 1969 through 1973. In the spring of 1974, in response to
early open water and no residual snow cover, geese had begun nesting at Lake
John Annex as early as March 9. A complete clutch of five eggs was observed
at the Annex on March 16, 1974. At least eight birds were observed on that
same date on Pole Mountain Reservoir. By April 3, 1974, 30 geese were present
at Lake John Annex and 14 at Walden Reservoir.
Summer Residents
An extensive survey of summer residents was not conducted in North Park in
1974. However, periodic observations during August indicated that more geese
were present in North Park during at least most of the summer period than in
previous years. Geese were observed on Lake John Annex, Walden Reservoir,
MacFarlane Reservoir, Hebron Ponds, Pole Mountain Reservoir, and the Case
Flats Ponds throughout August and early September. The largest group of
birds, 127, was sighted on Hebron Ponds on August 9,1974.

�-45-

Nesting and Production
A total of nine known nests were produced at three locations in North Park
in the spring of 1974 (Table 3). Unfortunately, only two of the nine were
successful. All six nests at Lake John Annex were destroyed by predators
and one abandoned nest was found on Walden·Reservoir.

Band Recoveries
Fourteen of the 100 goslings released on Walden Reservoir in 1973 were
reported taken by hunters during the 1973-74 hunting season (Table 4). Ten
of the 14 birds were reported taken in counties in the Panhandle of Texas,
indicating that a portion of the 1973 released birds did not follow the
normal Pacific Flyway migration route apparently used by most of the birds
released in North Park. It appears the birds taken in Texas were not
accompanied by birds released in North Park in previous years, as no indirect
recoveries for North Park birds were recorded in the Panhandle during the
1973-74 season. A portion of the 1973 releases did follow the normal
migration route as indicated by the California and Mexico recoveries presented in Table 4.
A summary of the distribution of recoveries through the 1973-74 hunting
season for North Park released birds are presented in Figs. 1 and 2. Direct
recoveries in Fig. 1 depict the Pacific Flyway migration pattern mentioned
above. Over 45 percent of the direct recoveries were reported taken in
Arizona with the large majority coming from the extreme southwest corner of
the state (Table 5, Fig. 1). Extreme southern California accounted for an
additional 19 percent of the direct recoveries (Table 5, Fig. 1). Excluding
Mexico, 78.7 percent of the reported direct recoveries came from Pacific
Flyway lcoations. The only major recovery areas for birds recovered directly
outside the Pacific Flyway were the degree block in Wyoming just north of
the release site, generally thought to be the route used by geese leaving
North Park, and the Texas Panhandle, an apparent first year wintering area
for a portion of the birds released in 1973 (Fig. 1).
In plotting indirect recoveries, Colorado becomes the major recovery area
with 29.4 percent of the recoveries (Table 5). Birds taken in California
amount to 26.5 percent of the indirect total while the number of recoveries
in Arizona decrease compared to the direct rate to only 17.6 percent of the
total (Table 5). Excluding Mexico, 63.6 percent of the indirect recoveries
were reported taken in Pacific Flyway locations. Nearly 21 percent of the
indirect recoveries were reported taken in areas north of Colorado, indicating some birds are by passing the release site on spring migrations (Fig. 2).
Recoveries along the foothills of eastern Colorado indicate that some of the
released birds have returned to their area of origin. Three indirect
hunting season recoveries have been reported taken along the foothills. In
addition, in the summer of 1974, four birds released in North Park were
recapt~red during extensive banding operations in the Fort Collins area,
three at Watson Lake and one at Terry Lake. All four recoveries were
indirect, with two birds each being released in 1971 and 1972.

�-46-

Table 3.

Known nesting data for Canada geese in North Park 1971-1974.
Number Nests Established
1971
1972
1973
1974

Area

Lake John Annex

1

3

6

Boettcher Lake

0

2

0

Arapahoe National
Wildlife Refuge

0

1

1

Walden Reservoir

0

0

0

6

Number Nests Hatched
1971
1972
1973
1974

1

2

4

0

2

0

1

0

1

1

1

2

0

0

0

1

0

Table 4. Hunting season recovery locations for Canada geese released on
Walden Reservoir, North Park, Jackson County Colorado, 1973.

Area Recovered

Number Recovered 1973-74

Colorado
Gunnison County

1

California
Imperial

County

2

Texas
Randall County

4

Armstrong

County

3

Deaf Smith County

3

Mexico
Baja California

Total

1

14

�-47-

Table 5. The distribution by area of direct and indirect recoveries through
the 1973-74 hunting season of Canada geese released in North Park.

Area

Direct
Number
Recoveries Percent

Indirect
Number
Recoveries Percent

Total
Number
Recoveries
Percent

Arizona

54

45.4

6

17.6

60

39.2

California

23

19.3

9

26.5

32

20.9

Colorado

9

7.6

10

29.4

19

12.4

Texas

11

9.2

0

0.0

11

7.2

Wyoming

8

6.7

2

5.9

10

6.5

Chihuahua (Mexico)

5

4.2

1

2.9

6

3.9

Sinaloa (Mexico)

4

3.4

0

0.0

4

2.6

Utah

2

1.7

1

2.9

3

2.0

Baja California
(Mexico)

2

1.7

0

0.0

2

1.3

Montana

0

0.0

2

5.9

2

1.3

North Dakota

0

0.0

1

2.9

1

0.7

South Dakota

0

0.0

1

2.9

1

0.7

New Mexico

1

0.8

0

0.0

1

0.7

Saskatchewan
(Canada)

0

0.0

1

2.9

1

0.7

Total

119

34

153

Spring Returns
On February 17, 1975 a pair of Canada geese were observed on the Fuqua Ranch
along the Illinois River near Rand, Colorado. These birds were reportedly
feeding on hay being fed to cattle. By March 21, 1975, larger groups of
birds began to arrive at Lake John Annex.

�,

,

'1"

~t-,_

-'-I...

I
.j::o.
00
I

.~"

-®- - ~

------

-----®~-" •.

\

"\

Circled numbers are
recoveries which
occur somewhere
within the Mexican State
indicated.

/@"

.:
I

~.

-.
Fig. 1.

Distribution

of direct recoveries

of Canada geese released in North Park.

�/.

./
I

I

+:\0

I

'@.

,_
.'7-' ,
~®j .'

1",-,

~

,-.

~

\

Circled numbers are
recoveries which occur
somewhere within the
Mexican state indicated.

/®

i

®

!

0'(;
Fig. 2. Distribution
in North Park.

of indirect

recoveries

through the 1973-74 hunting season of Canada geese released

�-50-

South Park Antero Reservoir

By mid-summer 1973, a total of 156 Canada goose goslings had been released
on Antero Reservoir in Park County, Colorado (Table 6). Seventy-six birds
were released in July 1972, and 80 in July 1973.

Table 6. Hunting season recovery locations of Canada geese released at
Antero Reservoir in Park County, Colorado.

Year Released
Ntnnber Released

Area Recovered

1972
76

1973
80

No. Recovered
Indirect
Direct

No. Recovered
Indirect
Direct

Colorado
Fremont County
Alamosa County
Pueblo County

1
1
1

0
0
0

0
1
0

2

0

0

5

0

1

Texas
Reeves County

Total

Only one bird released at Antero was reported recovered during the 1973-74
hunting season. That bird was taken in Alamosa County in the San Luis Valley.
Previous hunting season recoveries are presented in Table 6. Two birds
released in 1972 were recaptured during the summer of 1973, one in the State
of Chihuahua, Mexico and one at Wheatland Reservoir, Wyoming.
The largest ntnnber of Canada geese observed at Antero· Reservoir in the spring
of 1974 was 120 birds sighted in mid-April. The following nesting season at
Antero resulted in at least 40 goslings being produced.
Reports indicate geese were sighted below the dam at Antero Reservoir on
March 21, 1975. A snowstorm the following day apparently resulted in the
birds leaving the area.

�-51-

LITERATURE

CITED

Szymczak, Michael R. 1974. Experimental
studies on improving status of
Canada goose populations.
Colo. Div. of Wildl., Game Res. Rep., Fed.
Aid Project W-88-R.
October.
p. 47-53.

Prepared

~

0

~

bY~In1M~:ld·4.~~oJ!:i:...k:~1;'-1..·':""
.::I:.:oCW'l~~""'4,J1a6C1:""'-'_
Michael R. Szym~lk1i'?)
Assistant Wildlife Researcher

��October 1975

-53-

JOB PROGRESS

REPORT

State of

.:::C.:::.O,::LO.:::.RAD=::.:O::...------

Project No.

W-88-R-20

Migratory

Work Plan No.

2

Job No.

Job Title

Studies of Canada Goose Populations

Period Covered:
Personnel:

Bird Investigations
6

in Colorado

_
Transplant

Areas

April 1, 1974 to March 31, 1974

C. Bryant and Staff, Monte Vista National Wildlife Refuge; R.
Staffon, Colorado State University; M. Babler, D. Benson, S. Bissell,
S. Brock, G. Brown, G. Claassen, R. Clark, C. Crawford, G. Crawford,
M. DePra, R. Desi1et, J. Frothingham, B. Goetze, V. Graham, J. Gumber,
T. Henry, J. Hicks, J. Hobbs, R. Hopper, J. Jackson, C. Leonard, M.
Loftsgard, J. Lorentzson, D. Lowry, T. Lynch, C. Manghan, K. Miller,
R. Oakleaf, D. Owens, S. Palm, B. Peterson, F. Rinella, C. Roberts,
L. Rottman, G. Saville, H. Spear, S. Steinert, E. Wagner, and M.
Szymczak, Colorado Division of Wildlife.
ABSTRACT

Analysis of band recoveries of 130 Canada geese (Brant a canadensis) banded
outside Colorado and recovered in northcentral Colorado during the 1973-74
season did not reveal any new Hi-Line population production areas. Banded
geese classified as experimental transplants that had been released in the
Manito Lake area east of North Batt1eford, Saskatchewan; in the Pelican Lake
area east of Moose Jaw, Saskatchewan; in Crook County in northeast Wy~ming;
and Harding County in northwest South Dakota were represented in the northcentral Colorado harvest.
Analysis of tail fans indicate that the percent
of harvest composed of "small" Canada geese in northcentral Colorado has
stabilized at about 15 percent.
Southwestern Saskatchewan continues as a
prominent northern recovery area of "large" birds banded post-season in
northcentral Colorado with size 8 bands while Alberta continues as the major
northern recovery area of small birds wearing size 7B bands.
Colorado is
the prime southern recovery area for post-season banded geese with 43 percent of the large birds taken in the northcentral part of the state and 19
percent of the small birds being taken in the southeastern portion of the
state. The December 1974 coordinated inventory in the Hi-Line wintering
area resulted in a total of 62,069 while the January count was 41,986.
About 77 percent of the population during both counts was found in Colorado.
Mortality rates based on recoveries of birds banded post-season in northcentral Colorado, calculated by the composite dynamic method resulted in a
year rate of .279 and .326 rate for all years. Observations during the fall
and winter of resident geese, neck-collared in June and July 1974 indicated
that: (1) geese reared on ponds inside the closed goose hunting area seldom
leave that area; (2) hunting mortality for geese reared inside the closed
area is lower than for geese reared outside the closed area; (3) resident
birds seldom flew further than 3 km from resting areas before landing to
feed; (4) feeding flocks of resident birds formed a very close-knit association

(Abstract continued

on next page)

�-54(ABSTRACT, Continued)
which was maintained from early fall until the end of winter; (5) although
feeding in the same fields, very seldom did resident feeding flocks intermingle with migrant flocks. Movement data for marked birds from ten
banding areas are summarized.
During the pre-hunting season period wintering
geese showed a preference to feed in chopped corn fields; selected barley
fields for feeding at the same rate in which they occurred; and avoided
picked-corn fields. During the hunting season when most geese fed within
the closed area, geese indicated a preference for picked corn, oats, barley,
lawn and selected chopped corn at essentially the same rate at which it
occurred.
The bulk of the feeding activity during the hunting season
occurred in well grazed pastures inside the closed hunting area, yet geese
selected pastures at a lesser rate than they occurred. After the hunting
season picked-corn fields, most of which had been heavily grazed, were
highly preferred while chopped corn, oat, and wheat fields were used at the
same rate as they occurred.
Throughout the fall and winter period alfalfa,
bean, prairie grass, and sugar beet fields were essentially avoided completely.
During wet, muddy periods geese avoided corn fields and shifted
to barley fields or pastures.
Mean feeding flock size was 119 during the
pre-hunting season period, 211 during the season and 193 after the season.
Analyses of fecal droppings collected at College Lake indicated diets of
geese roosting at College Lake during the respective periods were as follows:
pre-season - 50 percent corn, 30 percent wheat, 5 percent smooth brome, 4
percent sedge, 4 percent barley, 2 percent green bristlegrass, 1.5 percent
cattail and 1.5 percent oats, with eight other plants present in trace
amounts; in-season - 32 percent corn, 10 percent wheat, 36 percent smooth
brome, 4 percent alfalfa, 3 percent bluegrass, with seven other plants in
trace amounts; post-season - 55 percent corn, 11 percent smooth brome, 11
percent wheat, 3 percent cattail, 18 percent decomposed matter, with seven
plants in trace amounts. At Elder Reservoir fecal droppings indicated the
diets of geese resting there during the respective periods were as follows:
pre-season - 98 percent corn, with seven species in trace amounts; in-season no estimates; post-season - 61 percent co~, 9 percent wheat, 6 percent smooth
brome, 5 percent tansy mustard, 2 percent alfalfa, 14 percent decomposed matter
with eight species in trace amounts.
Seven of the nine geese banded outside the San Luis Valley that were reported
taken in the San Luis Valley during the 1973 hunting season had been banded
at Wheatland Reservoir in Albany County, Wyoming.
Goose counts in the San
Luis Valley during the winter of 1974-75 totaled 1,013 on November 22, 934
on December 19 and 709 on January 13. A record low 69 geese were estimated
harvested in the San Luis Valley during the 1974 season by 249 hunters for
a seasonal bag of 0.28 birds/hunter.
An estimated total of 85 geese were harvested in the westcentra1 Colorado
permit area, 66 in Mesa County and 19 in Garfield County, by 370 active
hunters for a seasonal bag of 0.28 birds/hunter.
A record high total of
1,515 geese were counted during the January Inventory conducted on January
10 and 11, 1975.

�-55-

STUDIES OF CANADA GOOSE
POPULATIONS IN COLORADO TRANSPLANT

AREAS

Michael R. Szymczak

P. S. OBJECTIVE
To investigate the status of resident and migrant Canada goose flocks
and their interrelationships in areas in which populations have been
established through transplant programs in Colorado.

SEGMENT OBJECTIVES
la.

To determine
Colorado.

breeding

areas of Canada geese wintering

in northcentral

lb.

To examine migration routes and/or harvest patterns and distribution
of Canada geese wintering in northcentral Colorado and the San Luis
Valley.

2.

To estimate hunting pressure on, and hunter harvest of, Canada geese
in northcentral Colorado, the San Luis Valley and westcentral Colorado.

3.

To estimate annual mortality
Colorado.

4.

To make recommendations for continuing Canada goose hunting seasons in
westcentral Colorado, northcentral, Colorado and the San Luis Valley.

5.

To document fall and winter movements and distribution
population of Canada geese in the Fort Collins area.

of the resident

6.

To document the content of Canada goose fecal droppings
Collins area during the fall and winter period.

in the Fort

of Canada geese wintering

in northcentral

METHODS AND MATERIALS
1

Band recoveries listed on computer printouts provided periodically by the
Migratory Bird Populations Station were sorted and plotted as to banding and
recovery location, sex, and age at time of banding, and estimated breeding
area for birds associated with the San Luis Valley and northcentral Colorado.
Tail fans reported from Canada geese harvested in northcentral Colorado and
collected through the U. S. Fish and Wildlife Service's Waterfowl Parts
Collection Survey were examined and classified as to whether the fans
originated from large or small Canada geese.
Canada geese in northcentral Colorado were counted on December 11, 1974 and
January 8, 1975. In the San Luis Valley geese were counted on November 22,

�-56-

1974, December 19, 1974, and January 13, 1975.
Permits to hunt geese in northcentral Colorado were not required during
the 1974-75 hunting season, therefore no special survey concerning goose
hunting activity in northcentral Colorado was conducted. However, information will be forthcoming from the regular small game survey of waterfowl
hunters. All hunters obtaining special permits to hunt geese in the San
Luis Valley and westcentral Colorado were mailed questionnaires inquiring
about their hunting activity and success.
Recoveries of Canada geese banded post-season in northcentral Colorado were
sorted by age and recovery year and mortality rates calculated by the composite dynamic and relative recovery rate method.
All pertinent data collected concerning the status of the northcentral,
west central and San Luis Valley Canada goose populations were presented to
management personnel in Colorado for use in establishing hunting seasons.
In order to document the fall and winter movements a total of 400 resident
Canada geese of both sexes and various age classes were trapped and marked
during the 1974 summer molt period in the Fort Collins area. On each goose
was placed a U. S. Fish and Wildlife Service leg band, and an individually
numbered, orange plastic neck collar. Geese were marked on five reservoirs
within and five reservoirs adjacent to the study area. The' locations of
the banding sites and the collar numbers applied at each site are listed in
Table 1.

Table 1.
site.

Location of banding sites and list of collar numbers used at each

Banding Site
Location

Series of Collar
Numbers Used

Total Number of
Geese Banded

College Lake
Dean Acres -1/

OlA - 50A

50

5lA - 85A

35

Dixon Reservoir

86A - OOA )
OlE - l3E )

28

Sterling Gravel Pits 1/

14E - 53E

40

Watson Lake

54E - OOE

Terry Lake -1/

47

OlF - 50F

Number 8 Annex -1/

50

51F - 75F

25

Cobb Lake
Elder Reservoir
Grey Lakes

76F - OOF
76C - OOC

)
)

50

1/
OlC - 50C

50

51C - 75C

25
Total

1/

Indicates banding sites located within the study area.

400

�-57-

Due to the large area involved no attempt was made to follow all marked
geese at each banding site. Instead, a study area of limited size was
selected in which observational effort could be concentrated.
The study
area was located just north of Fort Collins (Fig. 1). This area was
selected because its juxtaposition with the ten banding sites should result in a large number of marked birds feeding in the study area. The
study area was selected so that the lower third was located in the closed
hunting area (Fig. 1). This increased the likelihood that marked geese
would be present in the study area throughout the observation period including the hunting season. The study area contains a representative
array of field crops and reservoirs normally used by the wintering goose
population.
To systematically observe the feeding flocks of geese within the study
area, a designated transect of roads was driven three times per week on
randomly selected days. One weekend day was always included in each week
to observe the effects of increased human activity associated with weekends. The majority of field observations were conducted after the geese
left the reservoirs in the morning.
Thus, observations occurred when the
greatest percentage of the goose population was simultaneously in the
fields feeding.
Evening flights tended to be irregularly scheduled, and
frequently did not start until late in the afternoon or evening, thus
darkness would come before the transect could be completed.
For each flock of geese observed feeding along the transect route, the
following data were tabulated on field sheets and later coded on Unisort
cards for analysis:
(1)
(2)
(3)
(4)
(5)
(6)

Total number of geese in flock.
Collar numbers of marked geese present.
Crop type (corn, barley, wheat, etc.).
Field type (standing, stubble, plowed, etc.).
Field moisture condition (dry,"wet, frost, snow).
Weather conditions.

To locate roost areas of marked geese periodic early morning counts were made
before the feeding flights on all ten banding sites, and at any other reservoir within the study area that was being used as a roosting place.
The botanical composition of the diet of the wintering goose population was
estimated from fecal pellets gathered at College Lake and Elder Reservoir.
These two reservoirs were selected because the first lies within the closed
area, and the latter in the open hunting zone. Collecting pellets from these
two areas allows a comparison to be made between food habits of geese roosting
inside and outside of the closed area.
Feces were gathered at weekly intervals from circular plots located along
the shoreline in areas traditionally used as resting and loafing sites by
the geese. Originally, nine plots were set at College Lake and six at Elder
Reservoir.
The number of plots established depended on the number of likely
loafing sites at each reservoir.
Unfortunately, the location, size, and
total number of plots had to be continuously changed in order to collect
adequate quantities of fecal matter, because the geese constantly redistributed

�-58-

their loafing site locations in response to changing water levels, population size, and ice conditions.
After collection, each weekly fecal
sample was oven dried at 700 C, ground in a lmm screen Wiley mill, and
stored in paper bags.
The plant species in the feces were identified and quantified by Colorado
State University's Composition Analysis Laboratory using a microhistological
technique.
Individual plant species can be identified using a 100X microscope by the unique characteristics of cutinized epidermal fragments and
lignified cell walls that remain after digestion (Storr 1961, Williams 1969,
Ward 1970, Free et al. 1970, Stewart and Stewart 1970).
Before wild fecal samples could be analyzed a collection of reference plants
was gathered from all field types normally used by the feeding geese. Both
the dominant crop type and its associated weeds were collected.
The reference material was ground in a Wiley mill having a screen with lmm openings.
Microscope slides were made from the various plant parts (i.e. stems, leaves,
seeds) and these were studied by laboratory technicians to learn each plant's
characteristics.
Since it was too costly to analyze each weekly fecal collection individually
the samples were composited into three periods; a pre-hunting season period
(25 September, 1974 to 12 November, 1974), a hunting season period (13
November, 1974 to 31 December, 1974), and a post-hunting season period
(1 January, 1975 to 18 February, 1975). Each period is 49 days which is the
length of Colorado's northcentral area goose hunting season for 1974. The
hunting season was chosen as the base period for comparison because hunting
pressure greatly influences goose movements and field availability.
The
feeding habits of the wintering geese are suspected to be quite uniform
within each period, but to change considerably between periods.
For this
reason no feces were collected for one week after each period while the
geese were "adjusting" their feeding habits.
Each period sample was
composited by extracting a 20 percent drY'weight subsample from all weekly
collections made during that period.
These sub samples were homogenized in
a blender to randomly mix the plant fragments.
Microscope slides were
prepared from the fecal samples as described by Sparks and Malechek (1968),
Ward (1970), and Flinders and Hansen (1972).
The percent frequency of recognized fragments in each period.'s fecal s amp Le
was estimated by counting the number of identifiable fragments in 20 microscope fields on each slide. Twenty different slides were analyzed yielding
a total of 400 microscope fields examined per period.
Next, the mean density (D) of discerned particles
field is determined by the formula:
F = 100 (1 - e -D)
where (F) represents
density of particles
by:

of a species per microscope

the percent frequency and e = 2.71828.
Finally the
per field is converted to relative percent density (RD)
RD = (X + Y) x 100

where X is the density of discerned fragments of a species and Y is the sum
of the densities of discerned fragments for all species (Hansen 1974).

�lO

28

29

ilH

REED'3'

li»

po:·m0

"'AWELLlHGTON
0'MGMT.
AREA

HAGEN
POND

@

r'

-H---+----I~i

(legend
. ::::::::::JCilyor Town
-[""""JHighway or Co"nty Rood
I~i Lake, Pond or .Reservoir·

2

l

~Riv~r

_f=l Section line

Scale

1:208,644

I

II

!)E~::£;;t::·:'t;;;;:1=:=::::::3~Wles
TAKES@
, POND 0.

12

I:&gt;

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t

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a ~~-- ..
,I "
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~,",?,~~_;om~i!IlBIiW'

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~
••••••.
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Ij

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FI~

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__

""·""",,IUIIIo""JUI

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~ •.••••..
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Fig. 1. Location of the intensive study area and boundary
closed hunting area.

line of the

'~~L""A-K-E~"'"

�-60-

A problem encountered with fecal analysis is the likelihood of differential
digestibility of various foods in the diet. A food which is more easily
digested than others would be underestimated by the fecal analysis technique.
Correction factors for differential digestibility were estimated by feeding
a hand-compounded diet of known dry weight composition to 5 captive geese.
Fecal samples were collected from the captive geese, and analyzed.
The
estimated dry weight composition (RD) was then compared to the known
composition and the necessary correction factors determined.
Three different mixtures of hand-compounded diets were fed to the geese.
The important goose forages as identified by the analyses of the fecal
samples were each represented at a high, medium, and low level (with respect
to its average relative percent density) in one of the three hand-compounded
mixtures.
This was done because the digestibility of a food may differ
when that food makes up a larger percentage of the diet as compared to a
smaller percentage.
Each hand-compounded mixture was fed to the captive
geese for three days. No feces were collected during the first day.
Gooche's Best poultry food was used as a maintenance diet. This material
was completely voided from the digestive tract after the one day pre-trial
period since the through-put-time for geese is only about 2 hours (Mattocks
1970). During the second and third days, six fecal samples were collected.
After two days on the maintenance diet the procedure was repeated using the
second and third diet mixtures.
The relationship between the estimated percentage dry weight (X) and the
actual percentage dry weight (Y) was compared for the digested hand-compounded
mixtures using regression equations which fit a straight line through the
origin.
These equations determine the degree of under or overestimation
due to'digestibi1ity.
The relationship between the dry weight (Y) and the
estimated (X) percentages is expressed as:
Y

= BX + E

where, the residual E follows N (O,&lt;r 2). 'The least squares estimate of B is
b = ~ XY / 2- X2 and "b" (the slope of the line) is used as the correction
factor for differential digestion.
A similar procedure was used to determine over or underestimation due to
problems in distinguishing differences by the technicians.
The same handcompounded diets of known composition were given to the technicians to analyze
before digestion by the geese. Regression equations were again used to
determine correction factors.

RESULTS AND DISCUSSION

Northcentral

Colorado-The

Hi-Line Population

Breeding Range

Foreign Recoveries
During the 1973-74 hunting season 130 Canada geese that were banded outside
of Colorado were reported recovered in northcentral Colorado (Table 2).

�-61-

An additional 29 foreign banded birds were recaptured during 1973-74 postseason banding operations (Table 3). The general banding locations of
birds harvested or recaptured during the fall and winter of 1973-74 that
were classified as wild-trapped locals when banded are presented in Table

4.
Analysis of banding areas presented in Tables 2, 3, and 4 did not reveal
any new production areas. However, small numbers of birds that were banded
and released as transplants, apparently in restoration efforts, were reported
harvested or recaptured directly in northcentral Colorado.
Three of these
transplant birds were released in the Manito Lake area, located directly
east of North Battleford, Saskatchewan near the Alberta-Saskatchewan
border
while five of the geese were released in the Pelican Lake area, just east
of Moose Jaw, Saskatchewan.
Another group of birds released in restoration
efforts in Crook County in extreme northeast Wyoming were represented by
five direct and one indirect recoveries in the northcentral Colorado harvest.
The birds released in Crook County were originally trapped in goose production
areas in northcentral Colorado. Four of the five direct recoveries were released as goslings while the other two recoveries were released as adults.
In addition, four birds banded in April 1973, in Harding County in northwest
South Dakota and classified as experimental immatures were also taken in
northcentral Colorado.
These South Dakota birds are also believed to be
geese released in restoration efforts.
Canada goose nesting areas at Dowling Lake in Alberta, the Cypress Hills
area of Saskatchewan, and Phillips County and Bighorn County. Montana, continue to contribute birds to the northcentral Colorado wintering population.

Tail Fans
A total of 85 of the 95 tail fans collected through the "Waterfowl Parts
Collection Survey" from geese bagged in'northcentral Colorado were used to
establish the comparative rate of harvest of "small" Canada geese in northcentral Colorado.
The percentage of small geese in the harvest sample has
been generally consistent over the 1970 through 1974 hunting period when
comparing within counties by year when seemingly adequate sample sizes are
available (Table 5). Of the three major harvest counties, Larimer, Boulder
and Weld. only Weld seems to fluctuate dramatically between years in the
percent of "small" birds in the harvest.
The general trend over the five
year period tends to suggest that the percent of harvest of small birds in
northcentral Colorado has stabilized indicating that the percent of "small"
birds in the population is remaining the same from year to year.

Winter Banding - Northcentral

Colorado

In January 1973, 178 Canada geese were banded post-season in northcentral
Colorado.
Size 8 bands were placed on 170 birds that were subjectively considered large, while eight small birds were banded with size 7-B bands.
These bandings brought the three year total of birds banded post-season since
1971 in northcentral Colorado to 2,205; 1,490 with size 8 bands and 715 with
size 7-B bands.

�-62-

Table 2. Banding areas outside Colorado of Canada geese recovered
northcentral Colorado during the 1973-74 hunting season.
Area

Alberta
Edmonton - Camrose Area
Dowling Lake Area
Brooks Area
Saskatchewan
Manito Lake Area
Pelican Lake Area
Cypress Hills Area
Eyebrow Lake

in

Direct

Indirect

1
17
1

1

1
5
8

3
4

10
1

CENTRAL FLYWAY
Montana
Phillips County
Bighorn County
Garfield County
McCone County
Treasure County
Wyoming
Crook County
Albany County
Heridan County

9

4

30
1
3
1

1

5

1

New Mexico
Socorro County
San Miguel County
Chaves County

1
1
1

4
1

2

South Dakota
Harding County

4

Kansas
Rook County

1

1

PACIFIC FLYWAY
Washington
Grant County
Walla Walla County

1
1

Idaho
---OWYhee

1

County

Montana
Beaverhead

County

1

----------------------------------------------------------------------------

�-63Table 2. Banding areas outside Colorado of Canada geese recovered in
northcentral Colorado during the 1973-74 hunting season (continued).
Direct

Area
PACIFIC FLYWAY

Indirect

(Continued)

Wyoming
Lincoln County

1

Utah
Emery County

1

MISSISSIPPI

FLYWAY

Missouri
Chariton

1

County

59

Totals

71

Table 3. Banding areas outside Colorado of Canada geese recaptured,
season, in northcentral Colorado, January 1974.
Area

Direct

Alberta
Brooks Area
Saskatchewan
Cypress Hills Area
Manito Lake Area
Pelican Lake Area

Indirect

1

8
2
1

CENTRAL FLYWAY
Montana
--Phillips County
Bighorn County

13

1

Nebraska
Garden County

1

New Mexico
Mora County

1

PACIFIC FLYWAY
Oregon
Harney County
Totals

1
4

25

post-

�-64-

Table 4. Banding locations of Canada geese classified as wild-trapped
locals which were recovered or recaptured in northcentral Colorado
during the winter of 1973-74.

Year of Banding
Before
1967 1967

Location

Alberta
Edmonton-Camrose Area
Beaverhill Lake Area
Dowling Lake Area
Brooks Area
S. of Brooks
E. of Brooks
Saskatchewan
Cypress Hills Area
Maple Creek Area
Cypress Hills Park
N. of Eastend
Cypress Lake Area
Vidora Area
Masefield Area

1968

1969

1970

1

2

1971

1972

1973

1
16
3

1

1

4
1

1
1
1

3
10

2

CENTRAL FLYWAY
Montana
Phillips County
Bighorn County
McCone County

3

5

1

3

17

8

1

5

1

PACIFIC FLYWAY
Washington
Grant County

1

Oregon
Harney County
Wyoming
Lincoln County
Utah
--Emery

County

1

1

1

�Table 5. Classification of Canada goose tail fans collected through the Bureau of Sport Fisheries and Wildlife's
Parts Collection Survey, from birds harvested in north-central Colorado, 1970-1974 hunting seasons.
Year
1970

1971
Percent
Percent
Small
Large

1972
Percent
Percent
Small
Large

1973
Percent
Percent
Small
Large

1974
Percent
Percent
Small
Large

County of
Harvest

Percent
Large

Percent
Small

Larimer

92(36) 1/

8(3)

91(41)

9(4)

92 (22)

8 (2)

93(28)

7(2)

86 (31)

14(5)

Boulder

100(1)

0(0)

86(18)

14 (3)

100(10)

0(0)

84(42)

16 (8)

88(21)

12 (3)

Arapahoe

-

-

100(2)

Adams

100(2)

0(0)

100(4)

0(0)

100(1)

0(0)

-

-

100(1)

0(0)

Weld

82(18)

18(4)

59(13)

41(9)

67(2)

33(1)

79(11)

21(3)

67(16)

33(8)

Morgan

-

-

-

-

-

0(0)

100(4)

Totals

89 (57)

17(16)

92(35)

8 (3)

83(81)

17 (17)

81(69)

19(16)

1/

11 (7)

Sample size in parentheses.

83(78)

I

0'\

V1
I

�-66-

In Table 6, the three year scheme of the distribution of recoveries of
birds banded in January, 1971 in northcentral Colorado is presented.
Recoveries during the 1973-74 hunting season show Saskatchewan continuing
as a prominent recovery area for "large" geese wearing size 8 bands. A
few more birds, wearing size 8 bands, percentage-wise were recovered in
Alberta in 1973-74 than in 1972-73.
However, the rate of recovery of birds
wearing size 8 bands in Alberta in 1973 was still lower than the rate that
occurred in 1971-72.
Recoveries during the 1973-74 hunting season of birds banded with size 8
bands in January 1972 reflected nearly the same distribution as recoveries
of birds banded in January 1971 (Tables 6 and 7). For both banding years
slightly over 18.5 percent of the recoveries were reported taken'in
Saskatchewan.
Alberta recoveries during the 1973-74 season comprised 9.3
percent and 12.5 percent of the recoveries and 1971 and 1972 banding,
respectively.
Only six of the 178 birds banded in January 1973 were reported recovered
during the 1973-74 hunting season. All six birds had been banded with size
8 bands.
Five of the birds were taken in northcentral Colorado and one bird
was taken in Saskatchewan.
Recoveries in Saskatchewan continue to come from the southwest part of the
Province; south of the South Saskatchewan River and west of Swift Current.
Most Alberta recoveries reportedly came from the Edmonton area. The lack
of recoveries of northcentral Colorado banded birds in southern Alberta
along with the general absence of recoveries in northcentral Colorado of
birds banded in the Brooks-Lethbridge,
- Medicine Hat area of southern
Alberta indicate that geese nesting in southeast Alberta may not be members
of the Hi-Line population as are the geese nesting in adjacent southwest
Saskatchewan.

Migration

Routes and Harvest Patterns

During the 1973-74 season, northcentral Colorado accounted for nearly 52
percent of the recoveries of geese wearing size 8 bands that were banded
post-season in northcentral Colorado since January, 1971 (Table 8).
Saskatchewan was the second major harvest area accounting for 18.5 percent
of the birds wearing size 8 bands.
Alberta continues to be the major recovery area for small birds wearing size
7 bands. During the three year hunting period 1971 through 1973, about 37
percent of the recoveries of small birds have reportedly come from Alberta
(Table 8). During that same period Saskatchewan accounted for 17 percent of
the small bird recoveries.
The small birds are thought to be members of the
shortgrass prairie population.
However, during that same 1971 through 1973
period, while Alberta was accounting for more than 68 percent of the "shortgrass" birds banded in northcentral Colorado, and taken in Alberta and
Saskatchewan, only about 55 percent of the shortgrass birds banded in southeast
Colorado during a comparable banding period were being taken in Alberta
(Szymczak, unpublished).
These differences in percentage distribution may be
real, indicating the northcentral Colorado shortgrass birds are oriented more
to Alberta than the southeast Colorado shortgrass birds, or they may simply
be a function of a somewhat small sample size for northcentral short grass
geese.

�Table 6. Percentage distribution of recoveries from 745 large (8 band) and 148 small (7B band) of Canada
geese banded post-season in northcentral Colorado, January 1971.

Location

1971-72 (First Year~
Size 7
Size 8

Hunting Season
1972-73 (Second Year)
Size 7
Size 8

1973-74 ~Third Year)
Size 7
Size 8
9.3
18.6

(4)

50.0 (1)

(8)

50.0 (1)

2.3

(1)

(0)

(0)

12.5 (1)
(0)

(0)

(0)

2.1

(1)

(0)

11.6

(5)

(0)

2.1

(1)

(0)

2.3

(1)

(0)

Alberta
Saskatchewan

13.8 (11)

40.0 (6)

18.8 (15)

6.7 (1)

2.1 (1)
31.3 (15)

Montana
South Dakota

7.5
2.5

(6)

13.3 (2)

4.2

(2)

6.7 (1)

Wyoming

2.5

(2)

(0)

Nebraska

2.5

(2)

(0)

(2)

12.5 (1)
12.5 (1)

I

Colorado

0'1

Northcentral
Southeast
New Mexico

46.3 (37)
(0)
(0)

Arizona

1.3

2'0.0 (3)
13.3 (2)
(0)

(0) ~I

39.6 (19)
8.3 (4)
4.2 (2)

25.0 (2)
37.5 (3)
(0)

51.2 (22)
2.3 (1)
2.3 (1)

(0)
(0)

(1)

(0)

2.1

(1)

(0)

(0)

(0)

(0)

(0)

2.1

(1)

(0)

(0)

(0)

(2)

(0)

2.1

(1)

(0)

(0)

(0)

Pennsylvania

2.5
1.3

(1)

(0)

(0)

(0)

(0)

(0)

Unknown

1.3

(1)

(0)

(0)

(0)

(0)

(0)

80

15

48

8

43

2

Nevada
California

Total Recoveries

�Table 7. Percentage distribution of recoveries from 579 large (8 band)
and 463 small (7B band) of Canada geese banded post-season in northcentral
Colorado, January 1972.

Location

Htmting Season
1972-73 ~First Year~
1973-74 (Second Year~
Size 8
Size 7
Size 8
Size 7

Alberta

21.8 (12)

42.9 (9)

12.5

(4)

36.4 (8)

Saskatchewan

21.8 (12)

33.3 (7)

18.8

(6)

9.1 (2)

Manitoba

1.8

(1)

(0)

(0)

(0)

Montana

5.5

(3)

(0)

3.1

(1)

4.5 (1)

Wyoming

1.8

(1)

(0)

3.1

(1)

4.5 (1)

Idaho

1.8

(1)

(0)

(0)

(0)

South Dakota

1.8

(1)

(0)

(0)

(0)

Nebraska

1.8

(1)

(0)

3.1

(1)

(0)

(0)

(0)

3.1

(1)

(0)

Northcentral

27.3 (15)

4.8 (1)

46.9 (15)

18.2 (4)

Southeast

10.9

(6)

14.3 (3)

3.1

(1)

22.7 (5)

Other

1.8

(1)

(0)

(0)

(0)

(0)

(0)

(1)

(0)

(1)

(0)

(0)

(0)

Arizona

(0)

4.8 (1)

(0)

(0)

New Mexico

(0)

(0)

(1)

(0)

Texas

(0)

(0)

(0)

4.5 (1)

Total Recoveries

55

21

32

Kansas
Colorado

Nevada
California

1.8

3.1

3.1

22

�Table 8.

Percentage distribution

of recoveries of Canada geese banded post-season

in northcentral

Colorado, 1970-71, 1971-72 and 1972-73.

Hunting Season
Size 8

Size 7

Size 8

Size 7

Size 8

Size 7

All Years
Size 7
Size 8

Alberta
Saskatchewan

13.8 (11)
18.8 (15)

40.0 (6)
6.7 (1)

12.6 (13)
26.2 (27)

34.S (10)
27.6 (8)

9.9 (8)
18. S (IS)

37.S (9)
12. S (3)

12.1 (32)
21. 6 (57)

Manitoba
Montana

7.5

(6)
(0)

13.3 (2)
(0)

1.0

2.5

(0)
(2)

(0)
4.2 (1)

2.7
2.7

(2)

(0)

4.9
1.9

(0)
(1)

(2)

(0)

7.4

(6)

4.2 (1)

3.8 (10)

(0)

(0)

1.0

(1)

(0)

(0)

(0)

Area

Wyoming

2.5

Idaho

1973-74

1972-73

1971-72

(1)
(5)

3.4

(0)

(0)

0.4
1.1

2.5

(2)

(0)

2.3

(0)

1.2

(1)

(0)

(3)

(0)
(1)
(2)
(0)
(0)

(6)
(0)
(0)
(1)
(0)
(0)
(0)
(0)
(0)

51.9 (42)
2.5 (2)
(0)
2.5 (2)
(0)
(0)
1.2 (1)
(0)
(0)
(0)

103

29

81

South Dakota

2.5

(2)

6.7 (1)

1.0

(1)

(0)

Nebraska

2.S

(2)

(0)

1.9

(2)

(0)

(0)

(0)

(0)

46.3 (37)
(0)
(0)
(0)
1.3 (1)
(0)
(0)
2.5 (2)
1.3 (1)
1.3 (1)

20.0 (3)
13.3 (2)
(0)
(0)
(0)
(0)
(0)
(0)
(0)
(0)

33.0 (34)
9.7 (10)
1.0 (1)
1.9 (2)
1.0 (1)

Kansas
Colorado
Northcentral
Southeast
Other
New Mexico
Arizona
Texas
Nevada
California
Pennsylvania
Unknown

Total Recoveries

80

15

1.0
1.9

10.3
20.7

3.4

(7)

(7)

36.8 (2S)
17.6 (12)
2.9
2.9
1.S

(6)

(1)
(0)

(1)
(3)

(2)
(2)

1.5

(1)
(0)

(1)

(0)

16.7 (4)
20.8 (S)
(0)
(0)
(0)
4.2 (1)
(0)
(0)
(0)
(0)

42.8(113)
4.S (12)
0.4 (1)
1.S (4)
0.8 (2)
(0)
0.8 (2)
1.5 (4)
0.4 (1)
0.4 (1)

14.7 (10)
19.1 (13)
(0)
(0)
1.S (1)
l.S (1)

24

264

68

0.4

(0)
(0)
(0)
(0)

I

'"

\0
I

�-70-

Over the three year recovery period, Colorado has accounted for nearly
34 percent of the recoveries of small birds banded in northcentral Colorado
(Table 8). The distribution of these Colorado recoveries indicate that
small birds banded post-season in northcentral Colorado have no special
affinity for northcentral Colorado.
In fact the majority of the Colorado
recoveries have come from the southeast portion of the state. In turn
about four percent of the birds banded post-season in southeast Colorado
have been recovered in northcentral Colorado during the 1971 through 1973
period.
There seems to be a definite interchange by short grass prairie
geese from one wintering area to another on an annual basis.
Plotting the distribution of recoveries of northcentral Colorado banded
birds taken during the 1973-74 hunting season gave no indication of additional potential Hi-Line production areas.

Population

Distribution

The December 1974 coordinated inventory in the Hi-Line wintering area resulted in a total of 62,069 birds being recorded (Table 9). The total was
approximately 16 percent below the December 1973 record high count and
very similar to the 1970 and 1971 totals (J. Sands, unpublished data). However, the count in Wyoming in December 1974 was considered incomplete.
The
January 1975 count of 41,986 (Table 9) was 47 percent above the January 1975
level but very close to previous January count totals for the 1970 through
1973 period.

Table 9.

Results of Hi-Line Canada goose population

Location

December

11

inventories,

1974-75.

January 8

'l:./

Montana

7,231

4,146

Wyoming

5 ,800 1./

4,100 1/

Colorado

47,671

32,611

New Mexico

1,367

1,129

Totals

62,069

41,986

Incomplete

count.

Yellowstone

River, Bighorn River, Ft. Peck Dam.

'l:./

3/
- Ground count of two reservoirs,

Goshen County.

�-71-

During both December and January counts, about 77 percent of the population was in Colorado.
Normally the percent of the popUlation in Colorado
increases between the December and January inventories.
However, both
Wyoming and Montana had record high populations in their states during
the January count.
During the December inventory most birds in the Fort Collins-Loveland area
were located on reservoirs within the hunting closure (Table 10). The
January count, taken after the hunting season, indicated a wider distribution of geese in the Fort Collins-Loveland area.
Between the December and January counts, the population in Colorado declined
by 15,000 geese. The majority of the decline in the number of birds occurred
in the Fort Collins-Loveland area (Table 11).

Mortality

Estimates

Documenting estimates of mortality for the Hi-Line Canada goose population
was not an original objective of the post-season banding operation in northcentral Colorado.
However, a series of bandings and resulting recoveries
have accumulated which lend themselves to some preliminary mortality analyses.
Post-season banding operations have been conducted in northcentral Colorado
in six out of the eight years since the winter of 1965-66. No banding was
accomplished after the 1966-67 or 1968-69 seasons.
Beginning in January
1970, the birds banded were subjectively classified as "small" or "large"
Canada geese. Size 7 bands were placed on "small" geese while "large" geese
were given size 8 ba-:J.ds.Pooling recoveries of both "large" and "small"
birds and subjecting them to a composite dynamic mortality analysis resulted
in essentially the same mortality estimates as was calculated for recoveries
of large birds only. However, considering the differences in the geographic
distribution of recoveries of birds with size 7 and size 8 bands, particularly
on terminal wintering areas in Colorado, the two groups should definitely not
be pooled for mortality analysis.
A total of 2,322 Canada geese were banded during an eight year period beginning in 1966. Unfortunately the banding effort and success was extremely
erratic during that period (Table 12). First year recovery rates varied from
.041 to .115 and averaged .084. Mortality rates calculated by the composite
dynamic method for first year and all years were .279 and .326, respectively.

Northcentral

Colorado - Local Movements

of Resident Geese

To document the local movements and distribution of the resident flock I made
weekly visits to each reservoir on which geese had been banded during the
summer. Occasional observations were also made at other reservoirs within
and adjacent to the study area when they were used as roosting sites. Most
observations were made at dawn before the morning feeding flight. During
each observation the collar numbers that were visible were read and an estimation was made of the total number of geese on the reservoir.
Figures 2-14
summarize these reservoir observations.
On each map is plotted all the
locations recorded for all geese banded at one reservoir.
In the following discussion the general pattern of movements from each of the ten banding sites will
be described.
I will first discuss the six banding sites within the closed area,
then the four in the open area.

�-72-

Table 10. Results of northcentral Colorado goose surveys, 1974-75.
Area

December 11

January 8

Fort Collins - Loveland
2,848
Lindenmeier Lake
Longs Pond
702
178
Reservoir No. 6
1,897
Terry Lake
Watson Lake
1,667
Claymore Lake
1,950
1,084
Sterling Ponds
Ft. Collins City Park
340
College Lake
5,400
Fossil Creek Reservoir
7,480
820
Boyd Lake
Hollister Lake (Old Windsor Reservoir)
50
New Windsor Reservoir
3,650
Timnath Reservoir
76
Woods Lake
150
Black Hollow Reservoir
7
192
Cobb Lake
Elder Reservoir
0
Reservoir No. 5
0
0
Douglas Reservoir
Horseshoe Lake
0
Horsetooth Reservoir
0
Sub-total

2,509
0
0
543
1,572
0
194
110
3,375
1,256
4,200
0
1,800
425
250
31
8
317
650
500
19
227

28,491

17,986

1,860
780
110
14
1
30
7,747
0
0

745
370
415
0
67
0
6,893
385
2

10,542

8,877

Boulder-Longmont Area
Terry Lake
McCall Reservoir (Stamp)
Baller Reservoir
Union Reservoir
Boulder Reservoir
Baseline Reservoir
Valmont Reservoir
Faivre Ponds
Dodd Reservoir

Sub-total

�-73-

Table 10. Results of northcentral

Colorado

goose surveys, 1974-75

December

Area

11

(continued).

January 8

Denver Area
Standley Lake
Sloans Lake
Rocky Mountain Arsenal
Thornton Water Supply
Great Western Reservoir
Lowry Air Base
Cherry Creek Reservoir
Denver City Park
Public Service
Cooley Sand and Gravel
Bowles Lake Area
Green Gables Country Club
Kettring Lake
Norgren Reservoir
Youngs field Lake

180
197
10
73
24
255
110
320

235
105

o
42

o
o
280
350

5

o

35
2,900
425

42
2,750

o
o

o

87
25
197

4,534

4,103

Barr Lake
Horsecreek Reservoir
Milton Reservoir
Latham Reservoir
Riverside Reservoir
Empire Reservoir
Jackson Reservoir
Prospect Reservoir
Orchard Area-South Platte River

700
1,500
150
412
300
42
1,000
0
0

75
0
0
770
0
110
380
200
110

Sub-total

4,104

1,645

47,671

32,611

Sub-total

Brighton-Gree1ey-Ft.

GRAND TOTAL

o

Morgan Area

�-74-

Table 11.

Winter

inventories

of the Colorado Hi-Line Canada goose population.

Count Date

Fort Co11insLoveland

Longmont-Boulder
Denver

Brighton-Gree1eyFort Morgan

1967
January 9

9,739

2,883

991

13,613

1968
January 10

12,217

4,029

678

16,924

1968-69
November 20
December 19
January 2 &amp; 13

15,848
20,905
19,693

3,461
4,236
4,874

2,667
1,170
775

21,976
26,311
25,342

1969-70
November 5
November 28
December 23
January 6

8,737
31,350
18,522
30,650

2,255
3,782
5,668
5,060

390
1,374
1,259
1,914

11,382
36,506
25,499
37,624

1970-71
November 4
November '23
December 22
January 6

12,612
29,970
36,034
19,879

1,690
16,710
12,664
15,566

348
1,370
3,055
2,425

14,650
48,050
51,753
37,870

1971-72
November 4
November 23
December 21
January 10

25,699
31,072
31,516
19,117

1,815 !/
9,181'
19,525
10,742

6,905
13,525
4,476
3,661

34,419
53,788
55,517
33,520

1972-73
November 6
November 22
December 20
January 10

15,230
21,118
13,841
13,595

1 ,240 !/
5,764
19,290
18,709

1,050
4,981
1,225
1,405

17,520
31,863
34,356
33,709

1973-74
December 13
January 9

31,558
14,854

10,336
6,802

10,380
1,809

52,274
23,465

1974-75
December 11
January 8

28,491
17,986

15,076
12,980

4,104
1,645

47,671
32,611

Denver area not included.

Total

�-75-

Table 12. First year recovery rates for large Canada geese banded postseason in northcentral Colorado.

First Year Recovery

Year Banded

Number Banded

1966

163

1967

0

1968

480

1969

0

1970

185

.043

1971

745

.115

1972

579

.104

1973

170

.041

Rate

.041

.054

College Lake
Most geese banded at College Lake remained strongly attached to that reservoir
throughout the study (Fig. 2). They seldom, and in some cases, were never
observed to roost at another reservoir.
Only Dean Acres Pond and Sheldon Lake
in the City Park of Fort Collins were used to any extent by College Lake geese.
Furthermore, many of the observations at these two sites were made in mid-day;
suggesting that they are used for daytime resting areas more often than as a
night-time roost.
Two adult geese banded at College Lake did wander considerably.
The birds
wearing collar numbers 06A and 39A were located together at Parkwood, Elder,
and Divide #8 Reservoirs, and 39A also used Watson Lake. In fact, these two
birds accounted for most locations of College Lake bands outside of the three
roost normally used by College Lake geese.

Dean Acres
The birds banded at Dean Acres remained relatively close to their banding site
until cold weather froze over their small 2 ha pond (Fig. 3). Moving
principally as a group they alternately roosted at College and Sheldon Lakes
during the winter, staying at each site for a week or two before switching
back again. They returned to Dean Acres Pond in late winter as soon as the
ice began melting.
Dean Acres geese were never located on a reservoir outside of the closed area.

�DRY CREEK
~lS.
HORT
20)

OUDRE
NO. 12

OSIPES
POND

POUORE
~HO.l

II

,.

2)

HUTCHINSON
&lt;@~OND

JJ
,)0

2Z

NORTH I
POUDRE
NO ••

O-

HAGEN
POND

legend
~

City or Town

[=-1

Highway or County Road

©I Lake, Pond or .Reservoir"

"

~RiYc,

I

ISection line

Scale
!)

1:208,644
1

~

!OJ

"

2 Miles
I

H

II

COBB LAK E Me" 1
@AREA

TAK ES@
PONDO
II

--..
FISHER

~m.'~~
,

RE!..

~
"u
KITCHEL

RES.

.
IJ

DEMEREi'
~NDI

••

I
Fig. 2. Reservoir utilization by geese banded at College Lake.
Pre-Season observation; J = In-Season; S = Post-Season.

P

�DRY CREEK'
~I;S.

OUDRE
NO. Il

OSIPES
POND

'I

NORTH POUORE
2~
~NO.l

U

H

,.

HUTCHINSON
@J~OND

~~~g@

~
.JO

1·AWELLINGTON
-.,.WMGMT.
~
AREA

HAGEN
POND

Legend
~
City or To"'"
[e::z::.rIHighway or County Rood

[~i

Lake,

s

Pond or .Reservoir

z

~Riyer

I

Scale

ISection

Line

1:208,644

II

()E~S;;:;;:J.='~j;;lC==~::i~
Miles
N

TAKES@
POND 0.

I'

COSS LAKE MC,MT.

@ AR-fA

16

~~~tR~'~

i

~~"-+""""~~~"~"""~~"""~I~"----!-F-I

t.

Fig. 3. Reservoir utilization by geese banded at Dean Acres Pond.
P = Pre-Season observatibns; I = In-Season; S
Post-Season.

f'

�-78-

Watson Lake
The Watson Lake flock was extremely cohesive and their EOvements more
restricted than at any other roosting site (Fig. 4). The majority of
these geese roosted only at Watson Lake. A small portion of the flock
was seen at College and Sheldon lakes, and at Dean Acres Pond during preseason. This apparently was a very transitory movement since these geese
were seen only once on these lakes and were all later found back at Watson
Lake. During and after the hunting season Watson Lake birds confined themselves almost entirely to roosting at Watson Lake. Many times they all
fed together in the same field and were never observed feeding more than 2
km from the lake. Watson Lake geese were never located on a reservoir in
the open hunting area.

Sterling Gravel Ponds
The geese banded at Sterling Gravel Ponds typically stayed at that site
(Fig. 5). Many times most of them could be found feeding together in the
same field only a short distance away. Movements to other reservoirs inside
the closed area were uncommon, yet most of these birds were occasionally
seen at from one to three other roosting sites. These occasional wanderings
seemed to occur throughout the study period.
This group of birds was affected
very little by disturbance from heavy traffic, gravel pit workers and human
activity in general. Many times they fed within several meters of a road
busy with trucks and automobiles.
They were never observed on a reservoir
outside of the closed area.

Terry Lake
The geese banded at Terry Lake dispersed widely (Fig. 6). They did not remain closely attached to their banding site as did most other geese banded
inside the closed area. Two factors may account for this apparent discrepancy.
The size and shape of Terry Lake hindered reading of collar numbers so that the
total number of successful sightings made on the reservoir itself were limited.
But most important I feel is the fact that two sides of Terry Lake are close
to the border of the open hunting area. Continued hunter disturbance as Terry
Lake geese flew out to feed in these directions may have caused them to move
to roosting areas deeper within the closed area. Terry Lake geese frequented
the Sterling Gravel Ponds the most, but also made considerable use of College
Lake and were found in lesser numbers throughout the closed area. The postseason distribution is especially interesting in that the Terry Lake geese
were so widely dispersed, and also, because they were the only birds banded
within the closed area that moved north to roost sites in the open hunting
area. The reason for such movement is not clear. Possibly, Terry Lake
serves as a brood rearing area for some of the geese hatched on reservoirs
to the north.

�NORTH POUDRE
~NO.l

DRY CREEK'
~IOS.

OUORE
1'10.12

051f'E5
POND
21

u

20)

W

~~~g@

~
ZI

lO

,.

2)

HUTCHINSON
&lt;@~OND

HAGEN
POND

'~WELLtNGTOt&lt;.

@&gt;

~~~l'

legend
~
City or T o"'n
1c:x:wIHig!tway or Ca!.lnty Road

i('§;$j Lake, Pond or .Reservoir·

2

~Riyt:r

I

ISection

Scale

line

1:208,644
II

!)E~3'2=·~~lE:
==~1::'":::l7Miles
N

TAKE5@
POND 0

COBB

"
LAKE

@ AR-EA

MGMl

'5

10

29

"

FISHER
RES.

~

DEMERE' .
~NDI

It

,.
!,.
r ,

Timnath

••

I
!

u
Fig. 4. Reservoir utilization by geese banded at Watson Lake.
Pre-Season observations;.I = In-Season; S = Post-Season.

P

�DRY CREEK

~!;~.

OUORE
NO. Il

OSIf'ES
POND

NORTH POUORE
20
~NO.I

21

2)

I.

HUTCHINSOtl
&lt;@~OND

A

HAGEN
POtlO

30

Z2

NORTH
ApOUORE
ytlO.4

I

~g~g@
&gt;~:~~\~NC;TON
@)
AREA

legend

E::a City or Town
I==-I Highway or CO:Jnty Roc:id
l~iLo!ce,

Pond or.Reservoi;

2

[;.gJ River
(

I Section

Line

Scale 1:208,6~4
!)e~::£2:.::':'i~lE=~=::i~
Miles

"

It

N

_~

.~~~'f,~
"'HINK L [Y
NOVAK

RES,

RES.\

FISHER
RES,

a

Fig. S. Reservoir utilization by geese banded at Sterling Gravel
P = Pre-Season observations; I = In-Season; S = Post-Season.
Ponds.

�HORTH POUDRE
20

POUORE _
NO.1l

DAY CREEK 051"[5
~~S,
POND
21

~'Q'" NO. I
"

U

Z)

HUTCHINSON
OHD

@Ji

,: ,.

,

~
HAGEN
POND

Legend
E::::! City or T O"'n
I-==-I Hig!'woy or County Rood

I~i

2

Lalce, Pond or ,Reservoi;

(;;:J Rivet
I

I Section

Scale

Line

1:208,644

'II

~E~~=':'i=~1==~=3r
Miles

"

N

TAKES@
POND 0

COBS LAKE M(,MT

@ AR'EA

16

~~

-:I-

~'NORTH_+-_..m'GREY~
RES. ~

sg~~~'i~
RES.

DEMEREI

~NDI

)4

i

Fig. 6. Reservoir utilization by geese banded at Terry Lake.
Pre-Season observations; , I = In-Season; S = Post-Season.
,

P

.~~~~~
__

NOVAK
RES.\,

": ,

HINK L E1
RE~

�-82Dixon or Herring

Reservoir

Like Terry Lake birds, the geese banded at Dixon Reservoir were also
weakly attached to their banding site (Fig. 7). However, this is where
the similarity ends.
Instead of dispersing widely, Dixon geese were all
closely associated with College Lake.
All the geese I observed from
Dixon Res ervoi r were seen at leas t once on College Lake. Half of them
were never seen anywhere except at College Lake. Most Dixon geese had
moved to College Lake by early September and remained close to there
throughout the winter.
Some remained to nest on College Lake.
Seven
Dixon geese were only rarely found at College Lake, and seen once at
Horsetooth Reservoir.
This suggests that Horsetooth may also be a
wintering site for a few Dixon geese.
The very close association of
Dixon geese with College Lake suggests that Dixon Reservoir may serve as
a brood rearing area for some of the birds hatched at College Lake.

118 Complex
I will now describe the movements for those geese banded within the open
area. The 118 Complex involves three different reservoirs; Annex 118, Divide
118, and Elder. We banded birds at both Annex 118 and Elder Reservoir.
These three reservoirs are situated very close together.
It soon became
clear that geese using one of these reservoirs would also frequent the other
two. A daily interchange of geese between the three reservoirs normally
occurred.
Sometimes all the geese would concentrate on one or two of the
lakes, and other times they would be spread out on all three.
Because
this intermixing occurred, and since movements from these two lakes was so
similar I will consider them tpgether.
Most geese banded on the 118 Complex remained in that vicinity during the preseason until duck hunting season opened on 5 October (Figs. 8, 9). This was
not always the case, however, as several
118 Complex geese did move south
into the closed area in early and mid-September.
The reason for this early
movement is unknown.
Movement south accelerated after the start of duck
season, probably because of the disturbance factor.
Some of the geese did
not leave the area until goose hunting opened on 13 November.
During the hunting season College Lake was by far the most frequented roost
site for 118 Complex geese, but they did not restrict themselves entirely to
that site and most of the birds were occasionally seen at other reservoirs
throughout the closed area.
As soon as open water was available after the hunting season most 118 Complex
geese returned to their banding site.
A few did show up at Grey Lakes about
5 km southeast of the 118 Complex.
Grey Lakes
Grey Lakes geese behaved similarly to 118 Complex birds except that they
abandoned their reservoir quite early in the fall (Fig. 10). Most of them
had moved into the closed area by mid- to late-September.
They did not disperse widely throughout the closed area. Most of the birds utilized College
Lake and Dean Acres Pond before and during the hunting season.
They completely
deserted the closed area after the hunting season and were seen during the
post-season only at Grey Lakes.

�DRY CREEK

~I;S.

OUDRE 1010.12

OS'I'ES

•

2'

~
10

II

II

HAGEN
POND

,.

POND

NORTH POUDRE
~HO.l

2.,

ZZ

2S

HUTCH1NSOt-l
&lt;@~DND

I .rr--~--~--~-"~~~g@

~~~~~RE
yNO.4

&gt;~WELLINGTON

@)

'!~~l'

Legend
E:2 City or T O"'n
I=-I

Hig!.way or CO:lnty Rood

l~iLake,

z

Pond or.Reservoi;

t.;.;d River
I

ISection

Line

Scale 1:208,644
()f~5~::'~b;lE===::::lr
Miles
N

.,

"
TAKES@

POND 0

COBB LAKE

@ AR-EA

MC,MT.

III

~

,
i'

l

i
"
FISHER
RES.

~

~
I'

~
-,

i

,'.\

Fig. 7. Reservoir utilization by geese banded at Dixon Reservoir.
P
Pre-Season observations; I = In-Season; S ~ Post-Season.

=

�DRY

HORTH POUDRE
~NO.l

CREEK
~I;S.

OUDRE
NO. 12

OS.PES
POND

II

20)

U

H

I.

HUTCHINSON

&lt;@fO
~g~g@
ND

30

it

il

A

HAGEN
PONO

'~WELLINGTON

@J

'1~~~.

legelld
~City
or To"'n
[=IHig!,way or Counly Rood

!©iLake, Pond or .Reservoir"

z

r;_~ RiYer
(

ISection Line

Scale

1:208,644

••

()E~5-::2="~c;:,~lE=::::=:ir
Miles
N
TAKES@
PON 0

COBB LAKE
@AR-EA

MC,MT.

'11

SHELDON LAKE"
-:O~'1I

•.~

•••
~

,;,.Q [Fort Collins:

I

\BJ

r

II

r

'~-~-f

1
Fig. 8. Reservoir utilization by geese banded at Annex #8 Reservoir.
P = Pre-Season observation~;, I = In-Season; S = Post-Season.

�OUDRE
NO. 12

DRY CREEK
MORT POUDRE
20
~HO.'

~"~'I---~

21

~
HAGEN
POND

I.

@JiO
,

ZI

n
HUTCHINSON
ND

!W
)0

n

I

NORTH
POUDRE
yNO.4

rr\

~~~g@

I

&gt;~WELLINGTO"
@&gt; .

'1~~~.

legend
~
City or To"'"
I==-I Highway or CO,,"ty Rood

ii@jLake, Pond or ,Reservoi;

z

~Riyer

I

ISection Line

Scale

1:208,644

!)E~::£~::.~~lE=~=~1
Miles

"

II

N

i'

~

Fig. 9. Reservoir utilization by geese banded at Elder Reservoir.
P = Pre-Season observations; I = In-Season; S = Post-Season.

�"~:;~~E

DRY CREEK

12

NO.

NORTH POUDRE
20
~NO.l

~ES'J-

__

'"

21

Zl

,.

25
HUTCHINSON
ND

&lt;@iO

~~~g/Q)

~

se

"

"

HAGEN
POND

&gt;~W
@&gt;

LLINGTON
~~~~.

legend
~Cily
or Town
[==-IHig!,way or County Road
®jLake,

Pond or.Reservoir·

~River

[

J Section Line

Scale

1:208,644

!)E~~~::.:'tbd~l==~~?
Miles

II

I'

N
TpAOK'~DS@
n

COBS LAKE MGMT.
@AR-EA

15

WINICK

~
@'PONDS

FISHER
RE~

~

Fig. 10. Reservoir .utilization by geese banded at Grey Lakes.
Pre-Season observations; I = In-Season; S = Post-Season.

P

�-87-

Cobb Lake
Cobb Lake is unique among the reservoirs in the open area in that it
sometimes holds geese during most of the hunting season. Unfortunately,
it was nearly impossible to approach the geese on Cobb Lake. These birds
were extremely wary and nervous. Before I could ever get close enough to
observe collar numbers with a spotting scope, the geese would fly out into
the middle of this large lake.
The resident geese on Cobb Lake that could be identified followed a similar
movement pattern as the birds at #8 Complex (Fig. 11). Most Cobb Lake birds
remained there until the start of duck season. Then they gradually moved
into the closed area, with nearly all of them inside the closed area after
the start of goose hunting. Apparently those birds that remain on Cobb
Lake are mostly migrants. Once inside the closed area College Lake was the
preferred roosting site. They did occasionally use several other reservoirs,
especially Sheldon Lake. During the post-season the Cobb Lake birds dispersed widely. Many of them eventually congregated at the #8 Complex and
Grey Lakes. Possibly they use these sites as staging areas while waiting
for the ice to break on Cobb Lake.

Summary
In the preceding section I have described the general pattern of local movements by resident geese from their molting and rearing sites on which they
were banded. I feel that the typical pattern of local movements has been
described, but it should be remembered that many exceptions to the general
rule are certain to exist, especially with an adaptive, flexible species
like the Canada goose.
Several important factors have emerged. The evidence suggests that geese
hatched and raised inside the closed area may seldom or never leave that
area. Most remain closely associated with their rearing site, and usually
feed nearby. Although band returns are at this time incomplete, what I have
received further emphasizes this point. No collars have been returned by
hunters from geese that'were banded on Dixon Reservoir, College Lake, Dean
Acres Pond, or the Sterling Gravel Ponds. All these sites are located well
within the closed area boundary. In contrast, 16 collars were returned from
the other banding site which were either on the border of or just inside the
open hunting area.
Another interesting difference also exists. Figure 12 shows a map of distant
sightings. These are geese which moved long distances (at least 24 km (15 mi»)
from their banding sites. No distant sightings were made of geese from the
four banding sites located well inside the closed area boundary, but 11 geese
banded on the border of or outside of the closed area did make these long
distance movements.
Hunting mortality for geese reared inside the closed area is probably lower
than for outside of or on the edge of the closed area. Seventy-three percent of the geese banded on the four interior reservoirs of the closed area
and known to exist before or during the hunting season were still seen in
the post-season. Only 51 percent of the geese banded at the edge of the
closed area or in the open area could be found during the post-season.

�HORTH POUORE
20
~NO.l

DRY CREEK
~I;S'

OSIF'ES
POND

,..
~g~g@

~
lO

HAGEN
POND

II

'~WELLINGTON

@)

~~~~.

l&lt;Jl

Legend
~City
or To"'"
[c=olHighwoy or Counfy Rood

I~i
[;:-~

lak e, Pond or ,Reservoir"
River

I

ISection line

Scale

o

~

1:208,644
1

*1'1

2

2 Miles

12

"

N
TAKES@
POND 0

COBB LAKE MC".,i

@ "R'EA

16

NOVAK
RES.,

",

FISHER
RES.

~

DEMEREI
~NDI

It

34

Fig. 11. Reservoir utilization by geese banded at Cobb Lake.
Pre-Season observations; I
In-Season; S = Post-Season.

P

�Fig. 12, M~p
boundary.
,~

~--.

collar sJghtings

of

.---

I,r.l

loode

at dlstrutces greater than 24 km (IS mil from the Study Area

0
0:=7-:0··2:\0=;=:~I:=~,=x=,=(:;l:;~:F1'
·C~T
w,-----I·;i

1"r='-i-;"-O'=;C=ooo'-Co=o~:,

_l:~.

HIt.

_

ltIJ

.

_

. I"~

Observation Site

Key:

(1) Total Collars Seen At Site
4SF Individua1,Co11ar

.

5 ~.

-

Numbers

·I

i

• -

I

·

I
I ---""

Observed

I

1. Study Area

,·

2. Arapaho NlVR (2)

·

3. Terry Lake (1) 18C
4. McCnl1 Lake (12) 7SF

I

S. Platte River (1)

I

I---

I
00
10
I

(1)

6. Lower Latham Res.
7. McIntosh

I

·I

I

(1)

8. Sanborn Res. (3) 92F, 50C

,,

9. Faivre Ponds (1)
10. Gapter Lake (8) 87F, 88F
11. Valmont Res. (6) (15F, 35F in field)
12. Teller Lake (1)

I

.

I

,

.

14. Sloan Lake (12) l3C, 35C
15. Denver Federal Center (10)

I.

13C, 35C, 9IF
16. Green Gables Country Club (1)

\..._.--r.Iii1'-'-' L r-'-'-'-'....,.._ ..-

._.-

17. San Luis Valley (1)

I

l .t .... .-.---),---.-----------.--. ,'.;-------------..-.

'

I

13. Lee Lake (1) 9IF

i

~,

r---

I

I

I

..

I

"1

l·

I

I

j

I

I

-.------_1.,;

\

I

•• r-----:'"

,

I

, I

Outline ,\,,,P
"f

r.".,."

_

AM!.~~~~~.~~~~,t:?~~~~~:.~,...COLORAl)O

~'..:,:~:..;;:~) ,.".. " .: '~:

1

'H~...,

M'r No.IOOS

.-

I

SCHOOL 8Flm;S

,

I,

I

_____ -_
--

I --'

---

.

-

:!:I------'-. ---"-"------------r,~

�-90-

Finally, the heavy in-season usage of College Lake by all resident geese
except those from Watson Lake should be considered. The importance of
protecting this reservoir from disturbance is high for the resident
goose population. Although a great deal of intermixing of this resident
flock occurs during the hunting season most geese eventually return to
their home reservoirs. This was found to even hold true for those geese
that made long-distance movements.

Northcentral Colorado-Field
Feeding Behavior of Resident Geese
For each collared goose a map was prepared on which all field feeding
locations made of that bird were plotted. An example of one of these maps
is shown in Figure 13. The direction and distance of resident feeding
flights made from reservoir roost sites was determined by comparing the
field locations of the geese with the location of the roost sites they were
using at the same time. This information has been summarized in Figure 14
which shows all observations of marked feeding geese made during the study.
On the figure the broken lines circumscribe the typical maximum extent of
resident feeding flights originating from the major reservoir roost sites
encompassed by the line.
The feeding sites of the resident geese were normally located a short
distance from their roost. The resident birds seldom flew further than 3
km before landing to feed. The majority of flights were much shorter than
this with the geese usually selecting one of the nearest available fields
from their roost site.
The resident birds would use a certain group of fields repeatedly, often
completely ignoring similar fields nearby. Usually, the majority of resident
geese on a reservoir all used the same field on a given day, though seldom
did they all arrive at the site simultaneously.
Traditional use of fields by resident geese in the open hunting area ceased
when hunting season commenced and the birds moved into the closed area.
Yet, after the hunting season the resident geese frequently returned to the
same fields they had used earlier, especially if the field had not been
plowed in the meantime.
When migrant geese arrived in large numbers, the integrity of most resident
feeding flights was not destroyed. The resident birds usually did not disperse throughout the many flocks of migrants. Though migrants and residents
often fed together in the same fields they would not intermingle. The
resident birds would remain together in a loose group in the midst of the
migrant flock. On several occasions a car passing a field of feeding geese
would cause the migrant portion of the flock to flush and flyaway while the
resident birds, who were accustomed to traffic, would remain on the field.
When the migrants first arrived they appeared to "follow" the resident geese
and feed in the same fields with them. As time progressed the migrants
would pioneer into new feeding areas, but most of the residents would not
move with the migrants, apparently preferring to stay in their traditional
fields.

�-~--1·--1

DRY CREEK
~ES.
21

legend
~City

or Town

!=-IHighway

I~i

Lake,

~

or County Road

28

Pond or Reservoir

HAGEN
POND

~River

I

ISection

line

Scale

1:208,644

~::::;:;

1

o

!~7:.~Nas:::igz2I Miles

33

r- ------

,.
Z9

CLAYM~LAKE

ANO:IJESKI
MARSH

@

BAK

RE

.1

Fig. 13. Reservoir and field feeding locations made on goose with collar
number 42C (banded at Elder Reservoir).
Reservoir observations are
circled and field feeding locations are underlined.
P = Pre-Season; I =
In-Season: S = Post-Season.

�~----~----~--~~~~~-+----~
I
)RY CREEK'O,IPES
ES.
POND

«:

NORTH POUDRE
lJ
WNO.1

...•

~ .&gt;

:I
I

1

\

~

;
19

)0

·1
.1

_---

1

l8

ft.,__

HAGEN
POND

~~ . ...---~

I
i

i

I

I

___ L_~_
I

I

II

I
CURTIS LAKE

" \fJJ

16

i
I

5(2)

!

21

16

tlORTH

-.

~~P@I

~

,

.
125
51

,P(3) \
I I •

f

L

I L __

'-:-1
I

~-2~9

I

' -- 1'---4-

__

"5 ~
~~~-----o.l

-D\r--+-----r-" ••.
\-...iiiiii.i:~-_~:----'_~~-_(ja-~t-:~.

~"~'"

Fig. 14. Distribution of collar marked Canada geese observed feeding
in the fields. P = Pre-Season; I = In-Season; S = Post-Season. Number
equals the total collard read in that Section. Parentheses indicates
collars seen but not read. The broken line describes the typical
feeding ranges from the band~g sites and reservoirs within its boundary.

&lt;""""",",0

\~.
l: r-'-) .

�-93-

Many times resident birds would come from two or three different roost
sites to feed together in one field. This situation may have caused
much of the frequent switching of roost site locations which occurred.
When the time came to return to their roost after feeding there was a
tendency for the resident geese to follow those who left first. Sometimes they would follow geese who had come from a different reservoir
than they had. This seemed to be most prevalent when the geese were
flushed from their fields by a disturbance.
It seemed that the resident
geese were more closely associated with certain feeding sites than they
were with a particular roosting reservoir.
When resident birds which had been banded in the open area were forced
into the closed area by hunting pressure, they often joined the feeding
flights which had already been established by resident birds inside the
closed area. This relationship would last until after the hunting season
when most resident geese returned to the vicinity of their banding sites.
In summary, resident feeding flocks develop a very close-knit association
which is maintained from early fall until the end of the winter.

Northcentral Colorado-Observations of
Feeding Activities of the Wintering Goose Population

Field Feeding
The geese usually fed in the fields for no longer than three hours before
returning to their roosting sites. Unfortunately, the entire road transect
could not be traversed in this amount of time, so it was necessary to divide
it into two routes; one in the north half of the study area and the other
in the south half. One route was driven on alternate observation days.
The intention of the field feeding observations was to obtain a population
estimate of field utilization by feeding geese. It is important to remember
that each sub-flock within the population has a somewhat different pattern
of field utilization when compared to other sub-flocks. Most sub-flocks
associated with a particular reservoir normally flew less than 3 km from
that site to feed.
A sum of all the field feeding observations along the study transect includes several different sub-flocks. Thus, results reflect the "average"
of field utilization by several sub-flocks. This "average" hopefully represents field utilization at the popUlation level.
During the pre-hunting season period (pre-season: 25 Sept. to 12 Nov.)
18,314 geese (154 separate flocks) were observed in the fields feeding
(Table 13). Only four different field types were utilized to any degree.
Over 78 percent of the geese were in chopped corn stubble; 17 percent in
barley stubble; 4 percent in grass pasture; and 1 percent in picked corn.
One flock of 40 birds fed in a harvested sugar beet field.
A major shift in field utilization resulted with the start of hunting season
on 13 November 1974. In just one or two days the vast majority of geese
had been forced into the closed area. Since almost no geese remained along

�-94-

the north transect route this route was traveled only once per week and
the south route twice per week. During this in-season period (13 Nov.
to 31 December) 38,781 geese (184 flocks) were observed feeding (Table
13). Utilization of field types was much more diverse with 41 percent
of the geese in grass pasture; 18 percent in picked corn; 17 percent
in barley stubble; 7 percent in both chopped corn and lawn; 3 percent
in both alfalfa and wheat stubble and about 1.5 percent in oats.
After the hunting season the geese gradually dispersed from the closed
area to any reservoirs with open water. During this post-season period
24,102 geese (125 flocks) were observed (Table 13). Picked corn now
became the most important field type with 32.5 percent of the geese
feeding in it. Grass pasture had 20 percent; barley 14 percent; and
chopped corn 11 percent. Oats had 12 percent and wheat 5 percent, however, both these estimates were inflated by the presence of one very
large flock and a few small ones. Alfalfa was 2 percent and sugar beets
was 1 percent.
Interestingly, a different field type was used predominantly during each
period; chopped corn stubble (pre-season), pasture (in-season), and picked
corn (post-season).
Barley stubble was used consistently throughout the
study, but at a moderate level. Lawn (primarily golf courses) was important only when the geese were crowded in the closed area. Alfalfa, oats,
and wheat were avoided before the hunting season and thereafter only
lightly used. Sugar beets and field beans were rarely used.
Before discussing preference, an understanding is needed concerning the
availability of field types to the geese. The acreages of croplands within
the study area and within the Fort Collins closed goose hunting area were
measured by drawing all fields onto a U.S.G.S. Quadrangle map and then
using a dot grid. The results are shown in Table 14. However, total
acreage planted is not a~ accurate indicator of food availability to the
geese. Two other factors must be considered; crop harvest chronology and
plowing.

Harvest Chronology
Food availability is closely related to harvest because geese in this area
rarely utilize standing crops. Harvest chronology was estimated at monthly
intervals by counting the number of miles of each harvested crop along both
sides of the road transect.
The results are presented in Table 15.
When the study began on 25 September 1974, all fields of barley, wheat and
beans had already been harvested.
Seventy-five percent of the corn had been
chopped for ensilage and over 60 percent of the hay fields were cut. No
sugar beets, oats, or corn fields to be picked had been harvested.
The sugar beet harvest began the second week of October and was complete by
mid-November.
Corn picking started about the same time, but the harvest was
not complete until early December.
About 20 percent of the hay fields and
nearly all oat fields wer~ left standing to be grazed by livestock in winter.
Of the total acreage of corn harvested, slightly less than 20 percent was
picked and the rest chopped.

�Table 13.
routes.

Field utilization

by geese during each period of the study as observed

along the transect

Period
Pre-Season
Total
Relative
Flocks
Percent

Total
Geese

In-Season
Total
Relative
Flocks
Percent

Total
Geese

Post-Season
Relative
Total
Percent
Flocks

Field Type

Total
Geese

Alfalfa

None

--

--

1328

11

3.4

452

6

1.9

Wheat

None

--

--

1250

2

3.2

1115

2

4.6

Barley

3099

22

16.9

6517

22

16.8

3379

24

14.0

Oats

None

--

--

549

2

1.4

2912

8

12.1

Chopped Corn

14310

123

78.1

2799

5

7.2

2693

21

11.2

Picked Corn

175

1

1.0

6992

55

18.0

7831

33

32.5

Sugar Beets

40

1

0.2

None

--

--

304

5

1.3

Pasture

690

7

3.8

15817

78

40.8

4742

23

19.7

Lawn

None

--

--

2795

5

7.2

None

Bare Soil

None

--

--

734

4

1.9

674

3

2.8

18314

154

100.0

38781

184

99.9

24102

125

100.1

Total

I
\D

V1
I

�-96-

Table 14. Acreage of field types within the total study area and
within the closed portion of the study area.

Entire Study
Area

Closed Portion of
Study Area

(acres)

(acres)

Corn

5833.8

704.0

Pasture

4729.2

2234.7

Alfalfa

3898.4

855.0

Barley

3729.9

350.8

Sugar Beets

2314.2

125.4

Prairie

1940.5

0.0

Beans

373.8

0.0

Oats

245.8

35.9

Wheat

157.7

139.7

Lawn

151.0

151.0

Sorghum

37.9

0.0

Totals

23412.2

4596.5

Crop
Type

�-97-

Table 15. Harvest chronology of agricultural crops in the study area
presented as percent of annual acreage harvested by date.
Field
T~e

25 Sept. 1974

Corn

75

96

100

Alfalfa

62

72 !I

100 !I

Barley

100

100

100

a

100

100

Beans

100

100

100

Oats

a

6

85

Wheat

100

100

100

Sugar Beets

Percent of Harvest
15 Nov. 1974

1 Jan. 1975

11

- Harvest includes grazing by livestock.

Plowing
As harvesting of standing crops made more food available, plowing of harvested fields continued to reduce food supply. Plowing was monitored in
the &amp;ame way as harvest (Table 16). The fall of 1974 was quite dry and
mild. The field conditions remained excellent for plowing from September
until the ground froze in December. The majority of plowing occurred in
corn fields. The percentage of all croplands that had been plowed rose
from 2.7 to 23.0 percent from mid-September until mid-December.

Preference
The actual availability of a field type for goose feeding is the acreage
of that field type harvested minus the acreage that has been plowed.
Preference assumes that geese select their feeding places rather than making
a random sample of what is available. An index may be used to test this
by comparing (a) the percentage composition of the field 't~e available with
(b) the percentage composition of field utilization by the geese (Newton and
Campbell 1970). The index (b)/(a)
indicates the preference shown for different field t~es: an index of 1 means that the geese are selecting for that
field t~e in relation to its availability; more than 1 that they prefer
that field t~e over others; less than 1 that they choose that field type
less than would be expected in relation to its availability; and 0 that they
avoid it altogether.

�-98-

Table 16. Percentage of plowing complete
(plowing began in late September).
Harvested
Field
Type

in each harvested

field type

Datie '

15 Nov. 1974
(percent)

1 Jan. 1975
(Percent)

Picked Corn

2.3

1.5

Chopped Corn

36.0

53.0

Cut Alfalfa

1.7

3.3

Barley Stubble

9.1

24.8

Pulled Sugar Beets

5.1

25.4

Picked Beans

42.6

55.3

Oat Stubble

0.0

0.0

Wheat Stubble

0.0

0.0

The preference index values for each period of the study are shown in Table
17. The pre-season period should give the best indication of preference for
field types since this is when the geese are least restricted in their movements and when the widest variety and greatest quantity of foods are available. Under these conditions only two field types were selected at rates
equal to or greater than would be expected in relation to their relative
availability.
Chopped corn stubble was highly preferred with an index
value of 4.9 and barley was taken in equal respect with its availability
with an index of 1. Somewhat surprisingly, picked corn had a very low
index of 0.3. This was confirmed by several field observations of goose
flocks which fed entirely on chopped corn even though the fields they were
using contained strips of picked corn nearby.
With the start of hunting season the relative availability of corn and barley
fields was much restricted since the geese remained for the most part within
the boundaries of the closed area (Table 14). The major differences of
field composition between the open area and the closed area can be seen by
comparing the "Pre-Season (a)" colunm with the "In-Season (a)" column in
Table 17. Under the restrictive conditions within the closed area during
season the most highly preferred foods were picked corn (5.3), oats (2.8),
barley (2.5), lawn (1.9), and chopped corn (1.1). The shift from chopped
corn in the pre-season to picked corn during the in-season may have been
caused by the heavy pre-season goose grazing which simply removed most of
the available food in the chopped corn fields. It should be remembered that
the above fields, though showing high preference values, were in relatively
short supply inside the closed area. The bulk of the feeding activity took
place on the well grazed pastures which comprised more than 50 percent of
the cropland within the closed area. Pasture had a preference index of 0.7.

�Table 17.

Comparison of percent availability of croplands with percent utilization for feeding by geese.

Field Type

Pre-Season
(a)
(b)
Percent
Percent
Utili.
Avail.

Period
In-Season
(bl a)
(a)
(b)
Preference Percent Percent
Utili.
Index
Avail.

Post-Season
(bl a)
(a)
(b)
Preference Percent Percent Preference
Avail.
Index
Utili.
Index
(b/a)

Com

(Chopped)

15.9

78.1

4.9

6.6

7.2

1.1

11.6

11.2

1.0

Com

(Picked)

3.9

1.0

0.3

3.4

18.0

5.3

5.9

32.5

5.5

Pasture

24.3

3.8

0.2

54.9

40.8

0.7

24.5

0.8

Alfalfa

14.2

-

0.0

18.4

3.4

0.2

19.9

19.7
1.9

0.1

Barley

17.4

16.9

1.0

6.6

16.8

2.5

14.8

14.0

0.9

Sugar Beets

11.3

0.2

0.0

2.3

-

0.0

9.1

1.3

0.1

Prairie

10.0

-

-

0.0

10.2

Beans

1.1

-

0.0
0.0

-

-

0.0

Oats

0.1

-

0.0

0.5

1.4

Wheat

0.8

-

0.0

3.5

Lawn

0.8

-

0.0

SorghtDIl

0.2

-

0.0

0.9

-

0.0

2.8

1.3

12.1

9.31/

3.2

0.9

0.8

4.6

5.8 1./

3.8

7.2

1.9

0.8

0.0

-

-

-

0.0

0.2

-

0.0

Totals

II

100.0

100.0

100.0

- Index is highly inflated due to the occurrence of one abnormally large flock in this field type.

0.0

,
,

\0
\0

�-100-

By the time the hunting season was over and the geese were free to feed
without restriction, several changes had taken place to alter food availability on the fields in the open area. Many fields had been plowed and
had virtually no value as feeding sites. Even more fields had been disced
which covers a good measure of the spilled grain left after harvest. Most
harvested fields which are not plowed are grazed by herds of cattle and
some very large flocks of sheep. This is especially true in the picked
corn fields. We examined a few fields after they had been heavily grazed
by sheep and found almost no vegetation left on which a goose could feed.
Finally, the geese are returning to the same fields on which they have
already spent two months feeding before the hunting season and thus much
of the food supply has already been removed from the fields even if they
had not been disced or grazed. Most of the above described activities
occurred in the corn fields.
Thus, it seems somewhat surprising that picked corn with an index of 5.5
was the most highly preferred field type during the post-season.
The
reason for this is not clear. It is possible that the heavy grazing
pressure by sheep and cattle which knocked down and removed many picked
corn stalks had the effect of opening up the stand. This may have improved
visability enough so that the geese felt secure when feeding in these same
fields they had avoided before the hunting season. Chopped corn had an
index value of 1.0, barley 0.9, and pasture 0.8. The only other field
types with index values above 1.0 were oats and wheat.
In summary, when the geese are free to select feeding sites without hunting
pressure only chopped corn, picked corn, oats, and wheat are selected for at
a rate greater than their availability would suggest. When hunting pressure
restricts movements barley and lawn are also preferred.
Pasture seems to
become more preferred as the winter progresses.
As other food supplies
become depleted the young green shoots which develop in pastures during
warm spells may increase in importance to the geese. Alfalfa, beans,
prairie, and sugar beets have very low preference, and for the most part
are avoided completely.

Flock Size
The average size of field feeding flocks varied between periods. Mean
flock size in the pre-season was 119; in-season 211, and post-season 193.
Mean flock size was lowest during the pre-season when the geese were widely
dispersed over the study area and before large numbers of migrants had
arrived and big roosting concentrations were developed.
Highest mean size
was during the in-season when the birds gathered in mass inside the closed
area. Post-season flock size remained high. This was because cold weather
had frozen over most roost sites in the open area. Only a few reservoirs
remained that had enough open water for roosting.
This fact kept the geese
concentrated even though hunting season was over.
Throughout the study most flocks ranged in size from 25 to 350 individuals.
Only eight flocks larger than 1,000 birds were seen in one field during the
study. The largest flock was in an oat field and contained 2,300 birds.

�-101-

Weather Effects
Changes in temperature, wind speed, or cloud cover d~d not influence which
fields the geese chose to feed in. Howeve r , moisture conditions of the
surface soil d~d influence field selection.
This is shown in Figure 15.
When surface soil was dry or frozen the geese fed primarily in corn fields.
But, when surface soil was wet and muddy from rains or melting snows many
feeding flocks would shift to barley fields or pastures.
Two factors may
account for this. First, and most importantly I feel is the problem of
"mudballing".
The soil in this area has a high silicon clay content.
When it gets wet it forms a sticky gumbo that clings to everything.
When
feeding, as the geese walk along and probe the soil with their bills large
clods of mud develop on their feet and bill tips. A rather small amount
of rainfall is all that is needed to create conditions in which the geese
can be seen plodding along, barely able to navigate through the corn fields.
Apparently this creates enough inconvenience that the geese will leave the
corn and move to barley stubble or pastures which have thick ground litter
where "mudballing" is not a problem.
The second possible factor relates to palatability.
The coarse, rough
barley seeds may become easier to eat when softened by wetness.
It may
also be easier to pullout sprouted barley seeds and young grass shoots
when the soil is soggy and friable.

Northcentral Colorado-Botanical Composition
of the Diet as Estimated by Fecal Analyses
Twenty kinds of plants were identified in the five composite period samples
of goose feces (Table 18). Trace amounts of some unknown forbs and grasses
were also found. Plants showing less than 1 percent relative density (RD)
in the fecal samples were considered as trace amounts. Plants that equaled
or exceeded a 1 percent RD were considered to comprise a significant portion
of the diet.
Before discussing the importance of different plant species in the diet,
correction factors must be used. The results of the feeding and technician
trials indicated that correction factors need to be applied to the relative
densities given in Table 18. The analyses of the three hand-compounded diet
mixtures before and after digestion are shown in Tables 19-21. Differences in
discernible characteristics of the plants caused most of the error by the
technician rather than differential digestibility.
This is evident by noting
the small differences between the before and after digestion RD estimates.
Much greater differences exist between the actual dry weight percentages and
the estimated RD percentages before digestion.
This technician error results because certain plant species have outstandingly unique cell wall
characteristics which makes them very easy to identify in a microscope field.
Thus, a tendency exists to overestimate these readily identifiable species
and to underestimate those plants having less distinguishable cell wall
characters.

�100
90

n

80
70

60+
Vl
"'0
r-

Q}

.,.....

T I I

I I

II I I

I I

II

o

Corn Fiel ds

101 I I

~n

~

I I

~

Al fal fa and
Barley Fields

50

LL..
C
.,....
Q}
Vl
Q}
Q}
C,!)

4-

40
30

I I I I I I I ~I~

~ II

I
""'"
0
N
I

0
oj-)

c
u
s,
Q}

20

Q}
0-

10

Dry

Wet

Dry

Wet

Dry

Wet

Dry

-Soil Moisture Condition-

Sep. 25Oct. 11

Oct.
12-15

Oct.
16-23

Oct.
24-25

Oct.
26-29

Oct. 30-

Nov.

Nov. 8

9-12

-Time Span-

Fig. 15. Relationship
pre-season period.

of soil moisture

condition with field selective by feeding geese during the

�Table 18. Percent relative densities of discerned fragments
of goose feces. Based on 400 microscope fields per sample.

Food Type

Pre-season
College Lake
Elder Res.

Alfalfa
Barley
Blue Grama
Bluegrass
Cattail
Composite
Corn
Green Bristlegrass
Indian Ricegrass
Loco Weed
Moss
Muhly Grass
Oat
Rush
Sedge
Smooth Brome
Sugar Beets
Tansy Mustard
Wheat
Wheatgrass
Unknown Forb
Unknown Grass
Decomposed Matter

Period and Location
In-season 1/
College Lake

period samples

Post-season
College Lake
Elder Res.

10.2
0.5

2.4
1.5

2.3
0.9

7.7
3.6

1.4
7.6

5.0
1.2
0.1
1.6
2.5

91.6

25.7

45.7

46.5

4.5
0.4
3.6
0.4
31. 3
3.3
0.1

found in the five composite

-

0.1
17.9
1.4
2.1
15.6
2.9
0.1
0.3
15.9
0.3

,..aI

3.3
0.3
0.4
0.3
0.2

0.4
0.1
0.4

0
\..oJ

0.2

0.4

0.2
24.5

0.1
8.0

0.1
6.5
0.1
0.1
20.7

-

7.9
0.1
0.3
0.4
26.4

0.1
0.3
0.1
3.9

-

13.1
5.8
0.1
0.2
19.6

1/
- Elder Reservoir

was located

in the open hunting

area and was not used by geese during the in-season

period.

I

�-104-

The minor effect of digestion is not surpr1s1ng even though plants in
the hand-compounded diets were ground to a 2mm particle size. The
through-put-time of geese is so fast that little digestion of cell walls
could occur. Most nourishment must come from the contents of ruptured
cells, and from the endosperm of seeds crushed in the gizzard. Apparently,
the majority of cell wall material for most plant species passes through
the goose's digestive system nearly intact; or if some cell wall digestion
does occur its extent is about equal for different plant species.
To correct the estimated densities in Table 18 regression equations were
calculated for each plant species used in the hand-compounded mixtures.
The after-digestion RD estimate was the (X) value, and the actual dry
weight percentage was the (Y) value. The slope (b) of the regression
equation is the correction factor for the combined effect of technician
error and differential digestibility (Table 22). A perfect correlation
between true dry weight percentage and estimated percentage exists when
the sloye equals one. Slopes less than one indicate overestimation;
greater than one underestimation.
The magnitude of the difference of the
slopes value from one is directly related to the magnitude of the error of
estimation.
Barley, green bristlegrass, oats, and rush were closely estimated to their
actual dry weights. Alfalfa, bluegrass, cattail, sedge, and tansy mustard
were moderately overestimated while moss was greatly overestimated.
Corn
was underestimated because its grain has a high endosperm to seed-coat
ratio compared to other grass seeds. Smooth brome and wheat were also
underestimated.
These two plants like most other plant species were often
eaten by the geese as young shoots. At this early stage of development the
cell wall characteristics of these two plants were poorly defined compared
to other species, and could not be identified in the microscope fields.
They are represented by the large "unknown grass" category in Tables 19-2l.
This resulted in a very large underestimation.
Corrected Diet Composition
The adjusted RO percentage values for botanical composition were calculated
for each plant species by:
Di

bi X RDi
n

Z. (bi X ROi)
i

= 1

where Di is the corrected density of the ith species divided by the sum of
corrected densities for all plant species (n) in the sample (Table 23).
Pre-season
Corn and wheat dominated the pre-season diet of College Lake geese. About
50 percent was corn, 30 percent wheat, 5 percent smooth brome, 4 percent sedge,
4 percent barley, 2 percent green bristlegrass, 1.5 percent cattail, and 1.5
percent oats. Eight other plants were present in trace amounts.
At Elder Reservoir the pre-season diet was almost exclusively corn.
98 percent with seven other species in trace amounts.

Corn was

�-105-

Table 19. Estimated percent relative density (RD) of discerned
in Diet #1 before and after digestion.

Food Type
Alfalfa
Barley
Bluegrass
Corn
Green Bristlegrass
Moss
Oat
Rush
Sedge
Smooth Brome
Sugar Beet
Tansy mustard
Wheat
Unknown Grass
1/Based

Actual
Percent in
Diet 111

fragments

RD Before
Digestion 1/

RD After
Digestion

10.1
2.0
1.6
21.6
0.4
26.6
1.2
4.S
6.2
0.4
0.0
9.6
0.0
16.0

6.1
1.7
3.6
4S.4
1.3
18.0
1.0
7.0
6.9
0.4
0.1
4.S
0.4
3.7

2.0
1.S
1.S
80.1
0.7
O.S
0.7
S.O
2.0
2.0
0.7
1.S
2.0

on 100 microscope

fields per sample.

1.IBased on 400 microscope
six samples.

fields per sample.

1.1

Given value is an average of

Table 20. Estimated percent relative density (RD) of discerned fragments
in Diet #2 before and after digestion.

Food Type
Alfalfa
Barley
Bluegrass
Cattail
Corn
Oat
Rush
Sedge
Smooth Brome
Sugar Beet
Tansy mustard
Wheat
Unknown Grass

Actual
Percent in
Diet #2

RD After
Digestion

18.1
3.3
9.0
9.0
17.6
3.7
11.4
0.0
0.0
0.3
12.7
1.S
13.4

lS.S
3.7
12.0
lS.8
lS.9
1.3
10.3
0.1
0.1
0.1
16.0
1.9
7.4

6.0
4.0
S.O
6.0
40.0
1.S
9.S
0.0
10.0
0.0
10.0
8.0

11

- Based on 100 microscope

21

RD Before
Digestion 11

- Based on 400 microscope
six samples.

,..

1.1

fields per sample.
fields per sample.

Given value is an aver age of

�-106:'"

Table 21. Estimated percent relative density
in Diet #3 before and after digestion.

Food Type

Alfalfa
Barley
Bluegrass
Cattail
Corn
Green Brist1egrass
Oat
Rush
Sedge
Smooth Brome
Sugar Beet
Tansy mustard
Wheat
Wheatgrass
Unknown Grass

Actual
Percent in
Diet #3

1/

fragments

RD Before
Digestion Y

RD After
Digestion

27.3
7.2
7.2
15.5
5.0
0.0
5.4
1.5
0.4
0.0
1.9
11.4
0.7
0.0
16.7

20.9
6.3
15.2
20.9
2.5
1.7
3.1
1.1
1.1
0.0
0.0
12.0
1.4
0.0
14.0

12.0
8.0
10.0
12.0
10.0
1.5
2.5
1.5
0.5
20.0
0.0
5.0
16.0
1.0

- Based on 100 microscope

(RO) of discerned

2:/

fields per sample.

2/
- Based on 400 microscope
six samples.

fields per sample.

Given value is an average of

Table 22. Correction factors for differences in the discernible characteristics and digestibility for each plant species used in the hand-compounded
diets. The correction factor equals the slope (b) of the after-digestion
regression equations.
Plant Type

Alfalfa
Barley
Bluegrass
Cattail
Corn
Green Brist1egrass
Moss
Oat
Rush
Sedge
Smooth Brome
Tansy mustard
Wheat

Correction

0.5
1.0
0.5
0.5
1.8
0.7
0.03
0.5
0.8
0.3
2.1
0.5
2.1

Factor

�Table 23. The adjusted relative
samples of goose feces. II

Food Type
Alfalfa
Barley
Blue Grama
Bluegrass
Cattail
Composite
Corn
Green Bristlegrass
Indian Ricegrass
Loco Weed
Moss
Muhly Grass
Oat
Rush
Sedge
Smooth Brome
Sugar Beet
Tansy Mustard
Wheat
Wheatgrass
Unknown Forb
Unknown Grass
Decomposed Matter

density percentages

Pre-season
College Lake
Elder Res.

-

for botanical

composition

Period and Location
In=aeason
College Lake

of the five composite

Post-season
Elder Res.
College Lake

-

3.5
0.3

0.8
1.0

-

-

0.7
0.3

2.7
1.3

0.5
2.5

1.8
0.9
0.1
0.6
1.0

97.9

32.1

54.7

61.1

-,

-

-

0.1

0.2

0.1

4.1
5.4
0.1
0.2
29.5

0.1
0.2

0.1
35.7

0.1
11.2

0.1
0.1
6.0

-

0.3

0.1
0.2

0.1
9.5
0.1
0.1

4.0

0.2
1.6
0.2
49.8
2.0
0.1

-

0.4
0.6
1.5

-

-

-

-

-

0.5

-

-

0.1

-

14.4

-

11.0
0.1
0.2
0.3
17.5

4~8
8.9
0.1
0.1
14.3

II
- Correction factors for plant species not tested in the feeding trials were estimated from the most closely
related species that was tested. All plant species with an adjusted percent relative density of less than
0.1 were given a value of 0.1 in the table.

I

I-'
0
'-I

I

�-108-

In-season
During the in-season period at College Lake corn dropped to 32 percent of
the diet and wheat to 10 percent, while smooth brome rose to 36 percent.
Alfalfa was 4 percent, bluegrass 3 percent and seven other plants occurred
in trace amounts.
Decomposed plant material comprised 14 percent of the
diet. Apparently decomposed matter was inadvertently consumed by feeding
geese when hunting pressure forced them to use low quality sites, and
freezing weather killed existing green vegetation.
Elder Reservoir was not used by geese during the hunting season so no
fecal collections could be made there during the in-season period.
Post-season
During the post-season period College Lake geese increased their use of
corn to 55 percent, while smooth brome dropped to 11 percent of the diet.
Wheat was 11 percent, cattail 3 percent and decomposed matter about 18 percent. Seven plants were present in trace amounts.
At Elder Reservoir the post-season diet was about 61 percent corn, 9 percent
wheat, 6 percent smooth brome, 5 percent tansy mustard, and 2 percent alfalfa.
Eight plants occurred in trace amounts and 14 percent was decomposed matter.
Summary
Fecal analyses provided more information on food habits than did field feeding
observations.
Even though considerable technician error occurred, the fecal
analysis was still very sensitive.
The feeding trials demonstrated that
fecal analysis could detect plants in the ~iet that comprised as little as
0.5 percent of the food intake. The fecal analysis revealed that many plants
besides crops were consumed.
Marsh plants such as cattail, moss, rush, and
sedge were found in the fecal samples.
Field weeds like tansy mustard, loco
weed, and composites were also used. Many uncultivated grasses were eaten
including blue grama, green bristlegrass, Indian ricegrass, muhly grass, and
wheatgrass.
The results of the fecal analyses suggest that non-cultivated
plants do sometimes contribute significantly to the diet, but seldom or never
make up a major portion of the diet. Sedge and green bristlegrass may be
important during the pre-season at College Lake, and tansy mustard during
the post-season at Elder Reservoir.
Cattail comprised more than one percent
of all the diets except during the pre-season at Elder Reservoir.
Even though the fecal analysis provided a more complete and specific picture
of the fall and winter diet, the major categories of foods used remained
similar to what was indicated by the field feeding observations.
The field
feeding observations did miss the importance of wheat to the geese when they
are feeding inside the closed area. This was primarily because most of the
wheat fields used by the geese were located south of the study area. Only
a few wheat fields were located inside the portion of the study area closed
to hunting.

�-109-

Both field observations and fecal analysis techniques have shown the
diet of the wintering geese to be quite diverse, but only a few plants
are used extensively.
Corn, smooth brome, and wheat seem to comprise
the bulk of the diet of the wintering goose population.

San Luis Valley

Foreign Band Recoveries
Nine birds banded outside the San Luis Valley were reported recovered in
the Valley during the 1973 hunting season.
Seven of the nine birds had
been banded on the Wheatland Reservoir molting area in Albany County,
Wyoming (Table 24). The other two birds were banded in northcentral
Colorado in 1966. Both birds were experimental with one being neck-collared
and the other transported and released.
For the second consecutive year
no birds banded in Canada goose production areas north of Colorado were recovered in the San Luis Valley.

Population

Distribution

Inventories of Canada geese in the San Luis Valley during the winter of
1974-75 indicated the typical scattered distribution during counts in
November and December with a steady accumulation of birds on the Monte
Vista National Wildlife Refuge (Table 25). Comparing the periodic counts
with the previous years, totals indicated an increase in the number of birds
in the San Luis Valley in 1974-75 compared to 1973-74 totals (Table 26).
Harvest
For the second consecutive year a total of 300 permits were issued in the
San Luis Valley authorizing the taking of one goose. An estimated 249
active hunters harvested 69 geese for a record low success of 0.28 birds/
hunter; the lowest success ratio since permits were first issued in 1970
(Tables 27 and 28). Harvest totals in 1974 on a numerical basis were essentially the same as in 1973 in all counties except Rio Grande where harvest
declined 57 percent.
The total harvest declined approximately 38 percent
in 1974 compared to 1973 (Table 29).
In 1971, 1972, and 1973, 60 percent, 47 percent and 44 percent, respectively,
of the harvest was reported taken on or near the Monte Vista Refuge.
During
the 1974 season, only 16 percent of the harvest was reported taken in that
same area.

Westcentral
Hunter Activity

Colorado

and Harvest

Permits authorizing the taking of one goose in designated counties in westcentral Colorado were again issued in 1974. In Mesa County 300 permits were

�-110-

issued while 100 were issued in Garfield

County

(Table 30).

Hunting success in the entire permit area in 1974 was essentially the
same as in 1973, however, the distribution of harvest by county was somewhat different (Tables 30 and 31). An estimated total of 19 of the 85
geese harvested were taken in Garfield County with 87 percent of the birds
reported taken along the Colorado River between Rifle and Grand Valley.
The number of hunter trips in westcentral Colorado in 1974 increased about
eight percent above the 1973 level (Table 32).
The season in westcentral Colorado in 1974 opened on November 2, approximately two weeks earlier than in previous years, and closed at about the
same time as in previous years, December 15. The earlier opening date
apparently had very little effect on the harvest as less than 10 percent
of the birds in Mesa County and none of the birds in Garfield County were
reported taken during the first two weeks of the season.
The rr~jority of the Canada geese wintering in westcentral Colorado were
located along the Colorado River from Silt to the Utah State Line during
the January inventory (Table 33). A record total of 1,515 geese were
counted during the January inventory (Table 34).

LITERATURE

CITED

Flinders, J. T., and R. M. Hansen.
1972. Diets and habitats of jackrabbits in northeastern Colorado.
Range Sci. Dept. Sci. Ser. No.
12. Colorado State Univ., Fort Collins.
29pp.
Free, J. C., R. M. Hansen, and P. L. Sims. 1970. Estimating the dryweights of food plants in feces of herbivores.
J. Range Manage.
23:300-302.
Hansen, R. M. 1974. Dietary of the chuckwalla,
determined by dung analysis.
Herpetologica
Mattocks, J. G.
22:107-113.

1970.

Sauromalus obesus,
30(2):120-123.

Goose feeding and cellulose

Newton, I., and C. R. G. Campbell.
1970.
1967/68.
Scottish Birds 6(1):5-18.

digestion.

Wildfowl

Goose studies at Loch Levan in

Sparks, D. R., and J. C. Melechek.
1968. Estimating
in diets. J. Range Manage. 21:203-208.

percentage

dryweight

Stewart, D. R. M., and J. Stewart.
1970. Food preference data by fecal
analysis for African plains ungulates.
Zool. Afr. 5:115-129.
Storr, G. M. 1961. Microscope analysis of faeces, a technique for
ascertaining the diet of herbivorous mammals.
Aust. J. BioI. Sci.
14:157-164.

�-111-

Ward, A. L. 1970. Stomach content and fecal analysis: Methods of
forage identification.
U. S. Dept. Agr. Misc. Pub1. (1147:146158.
Williams, O. B. 1969. An improved technique for identification
fragments in herbivore feces. J. Range Mange. 22:51-52.

Prepared by

~~~l1[~~d4L
Asst. Wildlife

Researcher

of plant

~t:~.AJ..
R1Crd
Staffon
Graduate

~
::::::

~~

�-112-

Table 24. Banding areas outside the San Luis Valley of Ganada geese
recovered within the San Luis Valley during the 1970-71, 1971-72, 1972-73,
and 1973 hunting seasons.

1970-71

Area

Number of Recoveries
1971-72
1972-73

1973

Alberta
2

0

0

0

Phillips County

1

0

0

0

Garfield County

0

2

0

0

1

5

6

7

0

1

0

0

2

0

0

2

1

0

0

0

Soccoro County

0

1

2

0

Mora County

0

0

1

0

0

1

0

0

Dowling Lake

Montana

Wyoming
Albany County
Wheatland

Reservoir

Fremont County
Ocean Lake

Colorado
Northcentral

Texas
Panhandle

New Mexico

Wisconsin
Columbia County

�-113-

Table 25. Results of Canada goose surveys during the 1974-75 hunting season
in the San Luis Valley.

November 22

Area

December

19

1/
January 13-

130

377

623

468

113

0

Del Norte to Monte Vista

0

70

8

Monte Vista to Alamosa

0

340

0

Rio Grande Mgmt. Area

0

0

10

Other Areas

0

0

16

Conejos River

5

22

22

La Sauses Area

20

0

0

San Luis Lake

107

0

0

Smith Reservoir

220

10

0

Sanchez Reservoir

63

0

0

McIntire

0

2

26

0

0

4

1,013

934

709

Monte Vista National Wildlife Refuge

Rio Grande River
Alamosa to State Line

Springs

Mishak Lake

Total

})
Poor count.

Birds were not on water areas during the count.

�-114-

Table 26.
Colorado.

Year

Winter inventories

of Canada geese in the San Luis Valley of

Number of Geese !/
November
December

October

1970-71

January

1,490

1,050

1,261

1971-72

1,570

913

1,255

1,286

1972-73

1,006

1,196

1,079

1,621

1973-74

289

736

774

905

1,013

934

709 '1:./

1974-75

1/
- Compared monthly inventories
five days of each other.

for the five years generally

2/
- Geese not concentrated.

Poor count.

taken within

�-1l5-

Table 27.

Year

1970-71
1971-72
1972-73
1973-74
1974-75

Hunter activity and success in the San Luis Valley permit area.

Number
Permits
Issued

Est. Number
Active
Hunters

Average
Days Hunted

Average
Seasonal
Bag/Hunter

200
350
400
300
300

164
296
311
233
249

3.4
4.2
4.6
4.2
4.3

0.65
0.60
0.82
0.48
0.28

Table 28. Distribution of harvest, by county, in the San Luis Valley
permit area.

Year

Rio Grande

Alamosa

Saguache

Conejos

Costilla

Total

1970-71
1971-72
1972-73
1973
1974

74
121
195
62
27

31
50
51
40
37

2
1
2
3
2

0
5
1
1
3

0
0
6
6
0

107
177
255
112
69

Table 29. Hunting pressure and harvest in the San Luis Valley goose
permit area 1973 and 1974.

Estimated Number
1973
1974

(± 5)

Total Individual Hunters

233

Total Hunter Trips

974 (±65)

Total Geese Bagged

112

(± 6)

Percent change
from 1973

(± 5)

+ 6.9

1,068 (±40)

+ 9.7

(± 5)

-38.4

249

69

Average Hunter Trips/Hunter

4.2

4.3

Average Bag/Hunter

0.48

0.28

Average Bag/Hunter Trip

0.11

0.07

�-116-

Table 30. Hunter activity and success in the westcentral Colorado
permit area.

Year

1971-72
1972-73
1973
Mesa Co.
Garfield Co.
Entire Area
1974
Mesa Co.
Garfield Co
Entire Area

Number
Permits
Issued

Est. Number
Active Hunters

Average
Days Hunted

Average
Seasonal Bag/
Hunter

150
250

112
182

2.9
3.7

0.29
0.32

300
66
366

254
46
300

3.9
3.7
3.8

0.30
0.20
0.28

300
100
400

235
67
303

4.3
3.6
4.1

0.28
0.28
0.28

Table 3L Distribution
permit area.

Year

1971
1972
1973
1974

of harvest by year in the west central Colorado

Mesa County

Garfield County

75
66

33
58
84
85

9

19

Table 32. Hunting pressure and harvest in the westcentral
goose permit area, 1973 and 1974.

Colorado

Estimated Number
1974
Percent Change

1973
Total Individual Hunters
Total Hunter Trips
Total Geese Bagged
Average Hunter Trips/Hunter
Average Bag/Hunter
Average Bag/Hunter Trip

Total

300 (+ 6)
1,152 (+51)
84
7)
3.8
0.28
0.073

(±

303 (:t7)
1,242 (+51)
85
6)
4.1
0.28
0.068

(±

+ 1.0
+ 7.8
+ 1.2

�-117-

Table 33.
1975.

Canada goose inventory,

westcentral

Colorado, January 10, 11,

Area

Number of Geese

Uncompahgre River
Delta-Montrose
Adjacent Warm Springs

21

o

Gunnison River
Black Canyon to North Fork
North Fork to Hotchkiss
Confluence North Fork to Gunnison River to Delta
Delta to Mesa County Line
Mesa County Line to Grand Junction

27

o
117
32
12

Colorado River
Grand Junction-Horsethief Canyon
Horsethief Canyon-State Line
Silt to DeBeque
DeBeque to Grand Junction

48
784
409

o
65

Mack Area
Highline Lake

o

Total

1,515

Table 34. Winter inventories of Canada geese in westcentral

Colorado.

Number of Canada Geese

Date
1967

January 17

71

1968

January 16

92

1969

January 2

307

1970

January 19

339

1971

January 4

879

1971-72

November 24
January 5

485
989

1972-73

November 21
January 11

329
1,183

1974

January 11

965

1975

January 10, 11

1,515

��-119...:

October 1975

JOB PROGRESS REPORT

State of __

--==C:.::o~L:.::O:.:.RAD=:.::O:...._
_
Migratory Bird Investigations
~W~-:.::8:.::8~-:.::R~-~2:.::0--_------

Project No.
Work Plan No.
Job Title

Period Covered:
Personnel:

2

Job No.~-------~----------------7
Non-hunting Mortality
Investigations of Canada Geese in Southeastern Colorado
October 22, 1974 to March 31, 1975

D. Bowden, Colorado State University; Marilyn Stevens, W. Adrian,
J. Carsella, J. Dennis, G. Eyre, J. Gustafson, G. Nugent, R.
Oakleaf, S. Steinert, J. Stephenson, M. Szymczak, Colorado
Division of Wildlife.

ABSTRACT

The amount of surface lead present prior to the 1974-75 hunting season in the
field adjacent to the firing line east and southeast of Turk's Pond was only
14.7 percent and 31.6 percent, respectively of what was present after the
1973-74 hunting season. During the 1974-75 season, additional accumulation
of pellets increased the amount of available surface lead to 45.1 percent of
1973-74 levels in the east field and 80.4 percent in the southeast field.
Comparisons indicated the extreme eastern portion of the southeast field, an
area subject to very heavy shooting pressure, contained most of the additional
surface lead. A total of 129 of 337 Canada goose (Branta canadensis) carcasses removed from Turk's Pond were selected for examination.
Only 44 of
the carcasses were completely intact and of those, 26 showed visible signs of
severe wounding.
Lead pellets were found in four of the wounded birds and
steel pellets in three birds. Four of 48 birds having complete gastro
intestinal tracts had impacted proventriculi.
Of the 53 birds having gizzards,
nine contained lead pellet fragments. Liver tissue was collected from 59 of
the 129 birds sampled while the ulna was collected from the other 70 birds.
Quantities of lead in the livers of 16 of 59 geese were considered sufficient to classify the birds as lead poisoning victims. Lead levels in the livers
of poisoning victims ranged from 25 ppm to 220 ppm with a mean of 86 ppm.
Bone tissue (Ulna) was found to be not a definitive indicator of lead poisoning. However, it was estimated that from 10 to 26 of the 70 birds represented
in the sample by bone tissue succumbed to lead poisoning.
An estimated 67 to
110 Canada geese died of lead poisoning at Turk's Pond during the fall and
w Ln ter of 1974-75, only 7 to 12 percent of the poisoning that occurred in
1973-74.

�-120-

RECOMMENDATIONS

1.

Continue the ban on the use of lead shot at Turk's Pond.

2.

Concentrate law enforcement efforts in areas of heavy hunting pressure
in which the firing line is adjacent to agricultural fields.

3.

Encourage pre or during hunting season plowing of harvested fields as
eas1y as possible in order to reduce the attractiveness of the areas
to geese to reduce the amount of surface lead. Concentrate on fields
in areas in which heavy hunting pressure occurred or is occurring.

4.

Discourage the establishment of perennial crops in fields adjacent to
the firing line.

5.

If additional mortality of Canada geese occurs and lead poisoning is
suspected as the causative factor and confirmation is desired, a forty
percent random sample from the available carcasses should be obtained.
A portion of the liver from each bird sampled should be removed, frozen,
and sent to the Laboratory director at the Research Center in Fort
Collins.

�-121-

INVESTIGATI0NS

NON-HUNTING MORTALITY
OF CANADA GEESE IN SOUTHEASTERN

COLORADO

Michael R. Szymczak

Lead poisoning was documented as the major cause of Canada goose nonhunting mortality at Turk's Pond during and after the 1973-74 hunting season (Szymczak 1974). Examination of agricultural fields adjacent to the
firing line at Turk's indicated these fields contained a definite, available source of pellets on the surface of the ground. In an effort to reduce further lead poisoning losses, a ban was placed on the use of lead
pellets for the 1974-75 hunting season in a four square mile area surrounding
and including Turk's Pond. This report deals with the results of the lead
shot ban in regard to lead poisoning losses at Turk's.

P. S. OBJECTIVE
To identify non-hunting
in southeast Colorado.

mortality

factors of Canada geese at Turk's Pond

SEGMENT OBJECTIVES
1.

Determine the incidence of lead shot in the environment
and the local vicinity.

2.

Determine
Pond.

3.

Determine if other causative agent of disease or poisoning
for mortality of Canada geese at Turk's.

4.

Determine

5.

Analyze data and formulate recommendations for preventative measures designed to lessen non-hunting mortality of Canada geese at Turk's Pond.

if lead poisoning occurs in the waterfowl

the magnitude

of non-hunting

mortality

at Turk's Pond

population

at Turk's

are responsible

in the Turk's Pond area.

METHODS AND MATERIALS
On October 17, 1974, prior to the Canada goose hunting season, agricultural
fields to the east and southeast of Turk's Pond were sampled to determine the
incidence of available lead shot by the transect-plot method described by
Szymczak (1974). According to measurements after the 1973-74 season, the
selected fields contained the greatest density of lead pellets in areas
supporting annual crops. Thus, it seemed logical to select these fields in
order to measure the amount of residual lead remaining after the fields had
been subjected to an annual sequence of normal agricultural activities.
Fields were to be sampled for 500 feet in front and 300 feet in back of the
firing line; again the area of heaviest concentration of lead. However, by

�-122-

mid-October only that portion of the corn planted in the fields to the
southeast of the pond had been harvested.
Therefore measurements of
lead were taken only in a portion of the area scheduled for sampling.
The field to the east containing winter wheat and non-irrigated milo was
sampled as scheduled.
All pellets found were classified as to location
and shot size by plot.
After the 1974-75 season the essentially same portions of the same fields
were again sampled in order to document the magnitude of accumulations of
lead pellets during the 1974-75 season.
In addition, a field directly
south of the pond was also sampled. All steel and lead pellets found were
classified to size and location.
Data sets composed of pellets per plot at certain distances by transect
were compared statistically by field in order to document significant changes
in densities between the various sampling periods.
All Canada goose carcasses were removed from around Turk's Pond periodically
from fall through early spring. A total of 15 pick-ups were made with the
first one occurring on November 18, 1974 and the final on April 8, 1975.
A 40 percent random sample of the carcasses collected during each sampling
period was selected for necropsy and organ collection.
Each selected bird
was to be examined for evidence of gunshot wounds and/or visible signs of
lead poisoning.
In addition the contents of the gizzard was to be examined
for lead pellet fragments and a portion of the liver collected to document
lead concentrations.
However, many of the carcasses were composed only of
the skeletal portion, in which case the ulna was collected to document lead
levels. Tissues chemically analyzed for lead content were treated in the
same manner as those collected during the 1973-74 period (Szymczak 1974).
An accumulative total of the number of carcasses removed from Turk's Pond
was maintained throughout the collection period.
According to 1973-74 findings, lead poisoning was the major cause of nonhunting mortality of Canada geese at Turk's Pond. Therefore no plans were
formulated to collect tissues for analysis to document other possible causes
of mortality.

RESULTS AND DISCUSSION
Although the raw data on lead pellet distribution collected post-season 197374 indicated slight changes in density over distance as measurements progressed
away from the firing line, these changes were not significant in the area from
700 feet in front to 500 in back of the firing line. The same held true for
all fields sampled during the 1974-75 period.
The number of lead pellets found in the east field pre-season 1974-75, after
an annual period of normal maintenance, was only 14.7 percent of what was present after the 1973-74 season at comparable distances (Table 1). In the field
southeast of the pond, pre-season 1974-75 measurements amounted to 31.6 percent of what was present after the 1973-74 season (Table 2). The differences
between the mean number of pellets for the two measurements for each field
were highly significant (t
= 8.38, P&lt;.OOI for east field; t82 = 5.20,
P&lt;.OOl,
southeast field). 88

�-123-

Table 1. Comparative distribution of lead pellets in the field east of
Turk's Pond post-season 1973-74, pre-season 1974-75 and post-season
1974-75, for up to 700 feet in front and 500 feet in back of the firing
line, weighted for sample size.
Distance from
Firing Line
in Feet

Post-Season
1973-74

Number of Lead Pellets Found
Pre-Season
1974-75

Post-Season
1974-75

700

7

11

3

600

10

500

14

2

4

400

14

3

5

300

16

1

4

200

11

1

6

100

14

3

2

11

0

3

100

12

1

7

200

7

2

9

300

10

3

8

400

8

11

3

500

8

- 1/

6

0 (Firing Line)

11

4

1/

Not sampled.

During the 1974-75 season, additional accumulation of pellets increased the
amount of available surface lead in the east field to 45.1 percent of what
was available post-season 1973-74; an increase of some 200 percent from preseason levels (Table 1). Numerical differences in surface lead as measured
post-season 1974-75 in the east field were significantly greater (P&lt;.OOl)
than what was present pre-season 1974-75 (t143 = 4.48, P &lt;.001), yet still
lower than 1973-74 levels (t
= 6.09, P&lt; .001).
279

�-124-

Table 2. Comparative distribution of lead pellets in the field southeast of Turk's Pond, post-season 1973-74, pre-season 1974-75 and postseason 1974 for 500 feet in front of the hunting pits, weighted for sample
size.
Distance In Front
Firing Line
in Feet

Post-Season
1973-74

Number of Pellets Found
Pre-Season
1974-75

Post-Season
1974-75

500

8

3

9

400

14

2

14

300

8

2

4

200

10

2

5

100

6

5

5

11

4

0 (Firing Line)

1:./

1/
- Not sampled.

In the area of the southeast field which was measured in all three periods,
the amount of surface lead accumulation during the 1974-75 season brought the
numerical pellet total to 80.4 percent of what was present post-season 1973-74
(Table 2). The differences in the mean number of pellets found post-season
1974-75 were not significant when compared with post-season 1973-74 data but
were significant (t68 = 3.34, P &lt; .01) when compared with pre-season 1974-75
data. However, in comparing a larger area in the same southeast field which
was measured post-season only in both years, we found that available surface
lead post-season 1974-75 was only 53.2 percent of what was present post-season
1973-74 with the means being significantly different (Table 3) (t
= 2.54,
222 only a
P&lt;.Ol).
These differences by location within an area indicate not
variation in hunting pressure, but also pinpoint possible areas in which
lead accumulation may be reaching the critical level.
Mortality
Necropsy

- Gross Observations

A total of 337 Canada goose carcasses were removed from Turk's Pond from November 18, 1974 through April 8, 1975. A sample of 129 carcasses was selected
and examined in order to document the cause of death. Only 44 of the selected
carcasses were completely intact permitting examination of most anatomical
features.

�-125-

Twenty-six or 59 percent of the whole body carcasses examined showed some
visible signs of suffering from serious wounds. Only four of the 26 carcasses
were in poor body condition. Pellets were found in seven of the 26 birds,
of which four had been wounded with lead shot and three with steel shot.
The gastro intestinal tract was intact in 48 of the 129 carcasses, while
gizzards only were present in an additional five birds. Four of 48, or about
eight percent of the carcasses with complete GI tracts had impacted proventriculi. Nine of Lhe 53 or about 17 percent of the gizzards examined contailed lead pellets. One gizzard, from a bird which had definitely been
wounded, contained five steel shot pellets. Three geese had both an impacted
proventriculus and lead pellet fragments in the gizzard.
Lead Content of Tissues
Liver tissue was collected from 59 of 129 sample birds while the ulna was
collected from the other 70 carcasses. Lead levels in the sampled tissues are
presented in Table 4. Quantities of lead in 16 of 59 or 27.1 percent of the
livers were considered sufficient to classify the birds as lead poisoning
victims. Lead levels ranged from 25 ppm to 220 ppm with a mean of 86 ppm
(Table 5). All other livers contained less than five ppm. Seventy-one geese
diagnosed as lead poisoning victims at Turk's Pond during 1973-74 had livers
containing from 33 to 283 ppm with a mean of 96 ppm. Livers of 10 hand reared
birds used as controls contained from two to 20 ppm, with a mean of 5 ppm
(Szymczak 1974).
Unfortunately, bone is not a definitive indication of lead poisoning as is
liver. In addition, during 1974-75 supporting data such as impaction of the
proventriculus or lead fragments in the gizzard were not available for birds
in the sample represented by bone tissue. During 1973-74 bone tissue (ulna)
from 64 lead poisoned birds at Turk's contained lead concentrations which
varied from 7 to 389 ppm with a mean of 30 ppm. Concentrations in bones from
five hand-reared birds ranged from 6 to 10 ppm with a mean of 8 ppm.
Theoretically,
the rate of lead poisoning as indicated by bone tissue should
be the same as indicated by liver tissue throughout the entire collection
period. However, during each succeeding pick up, fewer complete carcasses
were available. Therefore, as the collection period progressed and the probability of lead ingestion increased, the chance of a sampled bird possessing
liver tissue generally declined (Table 6).
ASSuming that scavengers select both lead poisoned or non-lead poisoned birds
for consumption at an equal rate, birds represented in the sample by bone
tissue should have the same rate of poisoning as those represented by liver
tissue within each collection period. The collection period ending on 1-20-75
contained 63 birds represented by 31 in the sample which were analyzed.
Sixteen of the birds were represented by liver tissue and 15 by bone tissue
(Table 6). Five, or 31.3 percent of the birds represented by liver tissue
were definite lead poisoning victims (Table 7). But only one of the 15 birds
represented by bone tissue con~ained, at 30 ppm, a lead level indicative of
lead poisoning. To obtain an additional four birds, lead levels as low as 8
ppm must be selected. If levels above 10 ppm, the maximum level for hand
reared controls were selected as lead victims, three of 15 or 20.0 percent

�-126-

of the birds represented by bone in the sample would be classified as lead
poisoning victims. If the same lead concentration criteria were applied to
the sample selected for the period ending 1-31-75, 42.9 percent of the birds
represented by bone would be considered lead victims, compared to 36.4 percent
of the birds represented by liver.

Table 3. Comparative distribution of lead pellets in fields south and southeast of Turk's Pond, post-season 1973-74 and post-season 1974-75, for from
100 to 700 feet in front of the hunting pits, weighted for sample size.

Distance from
Firing Line in Feet

Number of Pellets Found
Post-season 1973-74
Post-season 1974-75

700

10

11

600

13

8

500

21

6

400

21

12

300

18

8

200

16

7

100

12

7

In 1973-74 only four of 65, or 6.2 percent of the victims of lead poisoning
at Turk's contained bone lead levels of &lt; 10 ppm. Only 3 of 65, or 4.6 percent had lead levels &gt;10 &lt;15 ppm, while 12 of 65 had levels &gt;15 &lt;20 ppm. Of
the non-lead poisoning vi~tims, including the controls, 11 of l2-birds had
bone concentrations 2 10 ppm. The twelfth bird showed 11 ppm.
Considering all the above information concerning lead levels in bones, the
1974-75 data were classified according to what is termed a minimum, medium
and maximum poisoning rate (Table 7). The minimum rate includes all birds
represented by bone tissue which contains &gt; 20 ppm. The medium rate includes
those birds whose bones register lead &gt; 15 ppm, while the maximum rate
includes all birds with lead levels&gt; 10 ppm (Table 7).

Magnitude of Non-Hunting Mortality
The magnitude of the lead poisoning losses is calculated based on the mlnlmum,
medium and maximum rates and presented by time period in Table 8. All three
rates include all victims of lead poisoning as diagnosed by liver tissue in
addition to victims diagnosed by bone tissue at the respective levels.

�-127-

Table 4. Lead levels in tissues from Canada geese sampled from birds collected
at Turks Pond, southeast Colorado, 1974-75.

Goose
Sample No.

A

B

C

D

E

1
2
3
1
2
3
4
5
6
1
2
3
4
1
2
3
4
1
2
3
4
5
6
7
8
9

10
11
12
13
F 1
2
3
4
5
6
8
9
10
G 1
2
3
4
5

Tissue

Lead (ppm)

Liver
Liver
Liver
Liver
Liver
Liver
Liver
Bone
Liver
Liver
Liver
Liver
Liver
Liver
Liver
Liver
Liver
Bone
Liver
Bone
Bone
Bone
Bone
Bone
Bone
Bone
Bone
Liver
Liver
Liver
Liver
Bone
Bone
Liver
Liver
Bone
Liver
Bone
Bone
Liver
Bone
Bone
Bone
Liver

&lt;
&lt;

1.2
1.4
2.5
1.4
&lt;
2.1
1.8
&lt;
1.4
20
3.4
&lt;
7.1
&lt;
32
63
1.9
&lt;
3.0
52
2.5
&lt;
3.7
&lt;
37
&lt; 12.6
6
4
8
9
5
5
11
6
4
65
1.6
&lt;
4
6
12
2.8
&lt;
3.1
&lt;
6
26.6
11
4
6.4
&lt;
9
5
8
1.4
&lt;

Goose
Sample No.

G

6
7
8
9
10
11
H 1
2
3
4
5
6
7
I
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31

Tissue

Lead (ppm)

Liver
Bone
Bone
Bone
Liver
Liver
Liver
Liver
Liver
Bone
Bone
Bone
Bone
Bone
Liver
Liver
Liver
Liver
Bone
Liver
Bone
Bone
Liver
Bone
Bone
Bone
Bone
Bone
Liver
Bone
Liver
Bone
Liver
Liver
Liver
Liver
Liver
Bone
Bone
Bone
Liver
Liver
Bone
Liver

&lt;

4.3
13
7
7
3.6
&lt;
171
2.5
&lt;
33
7.7
&lt;
11
4
14
8
5
1.0
&lt;
2.4
&lt;
1.8
&lt;
70
7
3.5
&lt;
8
30
4.5
&lt;
5
5
8
8
11
38
6
151
11
56
1.7
&lt;
3.6
&lt;
1.4
&lt;
2.3
&lt;
7
8
7
1.9
&lt;
113
4
2.6
&lt;

-----------------------------------------------------------------------------

�-128-

Table 4. Lead levels in tissues from Canada geese sampled from birds
collected at Turk's Pond, southeast Colorado, 1974-75 (continued) •
Goose
Sample No.

J

K

L

M

N

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
17
18
19
1
2
1
2
3
4
5
1
2
3
1
2
3
4
5
6
7
8
9

10
11
12

Tissue

Lead (ppm)

Bone
Liver
Liver
Liver
Liver
Bone
Bone
Liver
Liver
Liver
Liver
Liver
Liver
Liver
Bone
Bone
Bone
Bone
Bone
Bone
Bone
Liver
Bone
Bone
Liver
Bone
Bone
Liver
Bone
Bone
Bone
Bone
Bone
Bone
Bone
Bone
Bone
Bone
Bone
Bone

7
29
&lt; 3.0
&lt; 2.8
25
8
14
&lt; 3.4
220
&lt; 4.2
112
3
&lt; 3.3
&lt; 3.7
27
9
7.3
17
12
30
7
&lt; 3.9
47
8
96
16
45
81
6
5
8
13
33
15
9
10
70
4
13
30

Goose
Sample No.

P

1
2
3
4

Tissue

Lead (ppm)

Bone
Bone
Bone
Bone

7
40
7.3
9

�-129-

Table 5. Lead concentrations in the liver and the occurrence of other physical
indicators in Canada geese considered victims of lead poisoning that were
randomly selected f~om carcasses collected at Turks Pond from 11/18/74 to 4/8/75.
Lead Fragments
in Gizzard

Body
Condition

Proventriculus

63

poor

not impacted

yes

D2

52

poor

impacted

yes

12/17/74

E12

65

poor

impacted

yes

1/6/75

G11

171

poor

impacted

yes

1/20/75

15

70

poor

impacted

yes

116

38

118

151

poor

not impacted

no

120

56

129

113

J2

29

J5

25

J9

220

no

J11

112

yes

2/24/75

L5

96

3/10/75

M3

81

Bird No.

Lead Conc.
(ppm)

C2

32

C3
12/9/74

Last Day of
Sample Period

12/3/74

1/31/75

no
poor

not impacted

yes

good, had
also been
wounded

not impacted

no

poor

not impacted

yes

not impacted

yes

�-130-

Table 6. Representation of liver and bone tissue by collection in interval
samples selected at random for analysis of lead concentration.

Last Day of
Sample Period

Number
Collected

Sample Size

11/18/74

7

3

3

0

100

11/26/74

7

3

2

1

67

12/3/74

15

4

4

0

100

12/9/74

16

4

4

0

100

12/17/74

34

13

4

9

31

12/30/74

31

9

4

5

44

1/6/75

28

11

5

6

45

1/14/75

27

7

3

4

43

1/20/7 5

63

31

16

15

52

1/31/75

44

18

11

7

61

2/9/75

5

2

0

2

00

2/24/75

15

5

2

3

40

3/10/75

6

3

1

2

33

3/24/75

31

12

0

12

0

4/8/75

8

4

0

4

0

Total

337

129

59

70

46

Organ SamE1ed
Liver
Bone

Percent
Liver

�Table 7. Rates of lead poisoning, by sample period, as diagnosed for birds represented by liver tissue, compared to three arbitrary lead
concentration levels for birds represented by bone tissue from carcasses randomly selected from Turks Pond, 11/18/74 to 4/8/75.

Liver Tissue

Bone Tissue
Lead Victims
Med. (&gt;15ppm) Max. (&gt;lOppm)

Sample Size

Lead Victims

Rate oj'
Poisoning

11/18/74

3

0

.000

0

0

0

0

11/26/74

2

0

.000

1

0

1

1

12/3/74

4

2

.500

0

0

0

0

12/9/74

4

1

.250

0

0

0

0

Sample Period

Sample Size

Min. (&gt;20ppm)

Rate of PoisoninB
Max.
Med.
Min.

---------.000

1.000

2

.111

.111

I
•....
.222 w
•....

1.000

4

1

.250

9

1

1

12/30/74

4

0

.000

5

0

0

2

.000

.000

.400

1/6/75

5

1

.200

6

0

0

1

.000

.000

.167

1/14/75

3

0

.000

4

0

0

2

.000

.000

.500

1/20/75

16

5

.313

15

1

1

3

.067

.067

.200

1/31/75

11

4

.364

7

1

2

3

.143

.286

.429

2/9/75

0

0

-

2

1

1

2

.500

.500

1.000

2/24/75

2

1

.500

3

1

1

1

.333

.333

.333

3/10/75

1

1

1.000

2

1

2

2

.500

1.000

1.000

3/24/75

0

0

-

12

3

3

6

.250

.250

.500

4/8/75

0

0

-

4

1

1

1

.250

.250

.250

Total

59

16

.271

70

10

13

26

.143

.186

.371

12/17/74

I

�-132-

Depending upon the rate of lead poisoning which applies to the number of
victims of lead poisoning at Turk's Pond in 1974-75 ranged from an estimated
67 to 110. During the same period in 1973-74 using a much less sophisticated
method of sampling, an estimated 908 birds died of lead poisoning (Szymczak
1974). Therefore lead poisoning losses at Turk's in 1974-75 were only in the
range of 7 to 12 percent of what occurred in 1973-74.
It was estimated in 1973-74 that 62 percent of the birds succumbing to lead
poisoning died during or shortly after the hunting season. The percentage
was similar in 1974-75 with, depending on the rate of lead victims utilized,
from 66 to 68 percent dying prior to the first of February. In 1974 the season
ended on January 20, while it ended on January 19 in 1975.
On a numerical
basis in 1974-75, lead poisoning deaths reached a peak in
late January, shortly after the hunting season (Table 8). Throughout the
remainder of the winter the number of lead poisoning victims did not change
substantially between pick up periods, indicating a fairly constant exposure
of birds to lead pellets throughout the fall and winter period. Collections
on 3-24-75 included a number of birds that had apparently been missed on other
collections, and therefore the rate of poisoning may reflect the entire fall
and winter period rather than the individual collection period.

CONCLUSION
The loss of Canada geese to lead pOisoning was substantially reduced at Turk's
Pond during the winter of 1974-75, compared with the winter of 1973-74.
ObViously, the ban on the use of lead shot at Turk's had a major effect. In
addition, weather conditions in late winter enabled plowing to begin in
portions 0 f some picked cornfields directly after the conclusion of the
hunting season.
Hunters disregarding the ban on lead shot did result in accumulation of lead
pellets in portions of the fields southeast of .the reservoir which approached
the 1974-75 level. However, it seems probable that plowing directly after
the season eliminated most of the exposed lead in what may have been a critical area.
The use of steel shot at Turk's Pond apparently also had a favorable effect
on reducing the total number of birds mortally crippled. According to 197475 season estimates, between 227 and 270 geese were lost to crippling,
compared to an estimated 982 during the 1973-74 season.

LITERATURE CITED
Szymczak, M. R. 1974. Non-hunting mortality investigations of Canada geese
in southeastern Colorado. Colo. Div. of Wildl., Game Res. Rept., Fed.
Aid Project W-88-R. October. p. 107-123.

�-133-

Table 8. The number of lead poisoning victims found at Turk's Pond by sample
period based on three specific rates of poisoning.

Last Date of
Sample Period

Rate of
Lead Poisonin8
Min. Med. Max.

Total Lead
Poisonin8 Victims
Max.
Min. Med.

Col1ected

Sample Size

11/18/74

7

3

.000

.000

.000

0

0

0

11/26/74

7

3

.000

.333

.333

0

2

2

12/3/74

15

4

.500

.500

.500

8

8

8

12/9/74

16

4

.250

.250

.250

4

4

4

12/17/74

34

13

.154

.154

.231

5

5

8

12/30/74

31

9

.000

.000

.222

0

0

7

1/6/75

28

11

.091

.091

.182

3

3

5

1/14/75

27

7

.000

.000

.286

0

0

8

1/20/75

63

31

.194

.194

.258

12

12

16

1/31/75

44

18

.278

.333

.389

12

15

17

2/9/75

5

2

.500

.500 1.000

3

3

5

2/24/75

15

5

.400

.400

.400

6

6

6

3/10/75

6

3

.667 1.000 1.000

4

6

6

3/24/75

31

12

.250

.250

.500

8

8

16

4/8/75

8

4

.250

.250

.250

2

2

2

Total

337

129

67

74

110

-1_

Prepared by _'-,-"n-,-n;JL)~" .;..&gt;::;;" OC::=,-_fl..L...:.....:... ";::;;.;;,jJ~~~
Michael R. Szym~
Assistant Wildlife Researcher

��October 1975

-135-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-88-R-20

Work Plan No.
Job Title:

2

Job No.

8

Investigation of Canada Geese in the San Luis Valley

Period Covered:
Personnel:

Migratory Bird Investigations

April 1 to August 31, 1974

H. Burdick, B. Clark, R. Desilet, N. Forbes, J.Frothingham,
H. Funk, G. Hinshaw, D. Lowry, J. Rauch, G. Saville, W. Schuett,
M. Szymczak, R. Velarde, R. Weldon, B. Widhalm, M. Zgainer of
Colorado Division of Wildlife; C. Bryant, J. Sarvis, S.
Cornelius of Monte Vista National Wildlife Refuge; B. Darnell,
D. Ziegler of Alamosa National Wildlife Refuge; M. Strong,
R. Tragstaad of Bureau of Land Management.

ABSTRACT

Canada geese (Branta canadensis) resident in the San Luis Valley of Colorado
were studied in the spring and summer of 1973 and 1974. Main objectives
were to develop methods for estimating size, distribution and productivity
of the breeding population from aerial surveys. Survey data collected using
fixed-wing and helicopter aircraft were compared with results of ground
surveys. Preferred nesting habitats included river bottoms with adjacent
ponds or sloughs, and lakes and ponds with islands or emergent vegetation.
Reliable estimates of numbers of breeding pairs were obtained from surveys
using fixed-wing and helicopter aircraft; estimates from helicopter surveys
were most accurate. Most aerial counts of goslings and adults during the
brooding seasons were unreliable. A count on river bottoms in late summer
from a helicopter was higher than a count from a fixed-wing aircraft on the
same transects. In 1973 and 1974, nesting began in late- and mid-March and
lasted approximately 89 and 82 days, respectively. Of the 160 nests studied,
44 percent were successful. Success of 108 nests on natural sites was 56
percent, and of 52 nests on artificial structures was 21 percent. Hardstem
bulrush (Scirpus acutus) was the most frequent plant near natural sites. Mean
clutch size was 4.9 eggs in 85 completed nests found. Estimated success of
egg hatching was 87 percent with a mean clutch of 4.3 eggs in 84 successful
nests, and 46 percent with a mean hatch of 2.3 eggs in 175 successful and
unsuccessful nests. Estimated mortality of goslings was 31 percent between
hatching and flight stage. Results were discussed, and recommendations for
future management and research were presented.

�-136RECOMMENDATIONS
1.

To estimate the breeding and non-breeding goose population, conduct
annual aerial surveys of water areas known to have geese during past
and present nesting seasons. Divide areas into transects similar to
those in Figure 2. Count every other transect containing river bottoms,
and all transects containing lakes and ponds which are isolated from
river bottoms. Preferably, use a helicopter, count~mmediately
prior
to the expected peak hatching period, and follow aerial survey procedures
as described in METHODS AND MATERIALS,under Aerial and Ground Survey
Techniques, Breeding Pair Counts.

2.

To predict production, approximate (1) the number of successful nests
from annual breeding pair counts (68 percent success in wooded river
bottoms, 78 percent success in open lake and pond areas and 93 percent
success in open river areas--applicable to active nests immediately
prior to peak hatching period), (2) the average number of goslings to
be produced in successful nests (4.3 per nest), and (3) the number of
goslings to survive to flight stage (69 percent of those produced).

3.

To detect possible expansion of nesting range, conduct an annual
aerial search of water areas that have not held nesting geese.
Preferably, carry out search during incubation period (April) and
prior to counts of breeding pairs.

4.

To promote increased production, develop areas similar to those
used for nesting in the past (see RESULTS AND DISCUSSION concerning
Delineation of Nesting Habitat). Preferably, develop areas in open
habitat which provide natural nesting sites such as islands, and
emergent vegetation with muskrat houses.

5.

To learn whether avian or ground predators are primarily responsible
for destroying nests in artificial structures and if certain structures
can have higher nest success than others, construct and compare structures with guards defending against ground predators with structures
presently standing.

�-137INVESTIGATION OF CANADA GEESE IN THE SAN LUIS VALLEY
Dennis G. Lowry

This second report for Job 8 is a summary of results from field studies
conducted during the spring and summer seasons of 1973 and 1974. The previous
report summarized findings of the first year's field study (Lowry 1974).

P. S. OBJECTIVE
To investigate the size, distribution and productivity of the Canada goose
population in the San Luis Valley during the spring and summer period and
develop a management plan.

SEGMENT OBJECTIVES
1.

To develop a technique for estimating the annual size and distribution
of the breeding population of Canada geese in the San Luis Valley.

2.

To develop a technique for estimating the annual productivity of Canada
geese in the San Luis Valley.

3.

To compile data, analyze results and prepare a progress report.
METHODS AND MATERIALS
Aerial and Ground Survey Techniques

Aerial surveys were conducted from a Cessna 185 fixed-wing aircraft at
elevations from 30 to 45 m (lOOto 150 ft) above the ground and at speeds
from 120 to 160 km/hr (75 to 100 mi/hr), and from a Bell 47-G3Bl helicopter
at elevations from 6 to 45 m (20 to 150 ft) above the ground and at speeds
from 25 to 80 km/hr (15 to 50 mi/hr). Ground surveys were made primarily on
foot and occasionally from a four-wheel drive pickup. Veeder handcounters
aided in counting geese and U. S. Geological Survey 7.5 minute topographic
maps were used to locate and plot nesting and brood-rearing data. Other
ground survey aids included 7 x 35 binoculars, a 20-power spotting scope,
hip waders, chest waders, two- to four-man boats, a 30.5 m (100 ft) and a
3.7 m (12 ft) measuring tape. Weather information was obtained from U. S.
weather stations at Alamosa, Center, Monte Vista, Del Norte and Saguache
in the San Luis Valley.
Habitat occupied by nesting geese was identified by aerial and ground
inspection of all water areas in the Valley below 2,470 m (8,100 ft), except
in the case of the Rio Grande River which was examined below 2,500 m (8,200 ft)
(Figure 1). Flights in fixed-wing aircraft on 20 and 24 April 1973 provided
fundamental information on the distribution of occupied goose nesting habitat.
Ground reconnaissance throughout the field study supplemented aerial data.
Aerial counts of geese were made by systematically counting within transects,
or by locating geese and then counting without a predetermined flight pattern.

�-138-

N

,,'
COLO:&gt;ALJO
SAN

LUIS

VALLEY

SCALE

IN

o

KILOMETERS
8

16

HHHA!
SCALE

o

IN

I H

38°
00'

MILES

5

10

H ES'""'=3

RIO

GRANDE

LEGEND

o

Town

9 us
-0

37°
00'

Highway

State

Highway

Appr ox i rn cte

2469

m Contaur

(BIOOfl)

[JJ

Refuge,

Management

Area

37°
00'

COLORADO

ih"WM'E'Xico- ---IS'

Figure 1.

San Luis Valley,

Colorado.

�-139-

The latter method was tried only for late summer brood counts when the geese
were in large groups. When possible, in all counts, the pilots aided the
observer(s) in spotting or counting geese. Techniques used in the aerial
surveys were designed to maximize counts of geese in each transect or study
area. Goose counts and nest surveys from the ground were conducted by
traversing wetland transects. Manner of approach varied according to different
terrains, water depths, vegetative conditions and goose habits.
River bottom goose habitat was divided into transects 1.6 by 4.8 km (1 by 3
mi) whose lengths were roughly perpendicular to the mainflow of water
(Figure 2). These transects included most river bottom ponds and sloughs
that geese preferred for nesting and brood-rearing.
Boundaries of the transects were section lines, usually marked by roads and fence lines visible
from the air. During goose counts, certain transects were flown lengthwise
with both fixed-wing and helicopter aircraft in a zigzagging pattern (Figure 3).
The number of passes needed to scan surface parts of the transects varied
and depended on vegetative conditions and goose habits.
Lake and pond areas isolated from river bottoms which contained geese were
also divided into transects. These transects were 1.6 km (1 mi) wide and of
various lengths depending on the size of the wetlands (Figure 2). Counts
from fixed-wing aircraft were conducted in lake and pond areas using the same
pattern as in river areas (Figure 3). Counts from a helicopter were conducted
over one water body at a time, but still using a zigzag pattern.
During counts from fixed-wing aircraft, two observers scanned the area below
both sides of the aircraft. During counts from a helicopter, a single observer
scanned the area below the front and both sides of the aircraft. Additional
passes were made or areas were circled where sightings of geese or counts
of geese were uncertain. All aerial counts were made under clear to partlycloudy skies.
Breeding Pair Counts
In 1973, aerial counts of breeding pairs were not attempted due to the
investigator's unfamiliarity with the study area during the nesting season.
In 1974, breeding pair counts were made from both helicopter and fixed-wing
aircraft.
During fixed-wing counts, four passes per transect were made, requiring the
observers to scan out to 200 m (220 yds) from the aircraft to see all surface
area. On helicopter counts 8 to 12 passes per transect were made, requiring
the observer to scan out to 100 m (110 yds) and 67 m (73 yds), respectively,
from the aircraft to see all the surface area. These aerial counts were made
between 0730 and 1430 daylight savings time.
During aerial and ground counts, geese observed were recorded as (1) geese
on nests, (2) singles not on nests (usually mates of nesting geese), or
(3) paired and (4) grouped geese (apparently immature, nonbreeding birds).
During aerial counts, the numbers of geese on nests or of singles not on
nests, whichever number was greater on each transect, were used as indicators
of breeding pairs. After flushing a nesting goose during helicopter counts,
eggs were occasionally counted and recorded. During ground counts, only
the numbers of geese on nests were used as indicators of breeding pairs.

�-140-

Brood Counts
In 1973, aerial counts of broods were conducted using only fixed-wing aircraft.
In 1974, aerial counts were made from both fixed-wing and helicopter aircraft.
During fixed-wing counts of flightless geese, 2 passes per transect were made
during 1973 and 4 passes per transect were made during 1974, requiring observers
to scan out to 400 m (440 yds) and 200 m (220 yds), respectively, from the
aircraft to see all the surface area. During helicopter counts of flightless
geese, 7 to 22 passes per transect were made, requiring the observer to scan
out to 115 m (126 yds) and 37 m (40 yds) , respectively, from the aircraft to
see all surface area. These aerial counts were made between 0800 and 1630
daylight savings time.
During aerial and ground counts, flightless geese were recorded as adults
(fully-feathered geese) or goslings (downy to partially-feathered geese).
Post Brood-rearing Counts
On fixed-wing counts of geese capable of flight, 2 passes per transect were
made during 1973 and 4 passes per transect were made during 1974, requiring
observers to scan out to 400 m (440 yds) and 200 m (220 yds), respectively,
from the aircraft to see all the surface area. On helicopter counts, 4 to 6
passes per transect were made, requiring the observer to scan out to 200 m
(220 yds) and 134 m (147 yds), respectively, from the aircraft to see all
the surface areas. These aerial counts were made between 1000 and 1330 daylight savings time.
During aerial and ground counts of geese capable of flight, geese were recorded
as a combined total of adults and goslings. since the difference between the
two could not be recognized.

Nesting and Gosling Mortality Studies
With few exceptions, studies were carried out as described by Lowry, 1974. To
supplement descriptions of nesting sites in 1974, live plants found within 1
m (3.28 ft) radius of, or overhanging natural nests were collected and identified to genus, and to species when possible.
Gosling mortality between hatching and flight stage was estimated by two
methods. One method involved comparing periodic ground counts of goslings
on different isolated areas (Figure 4). The other method involved comparing
estimated numbers of eggs hatched and latest ground counts of goslings on
the same areas. Unhatched eggs were opened, contents examined, and eggs
classified as to fertile, (Cooper and Batt, 1972) or infertile or whether
the embryo had died at an early development stage. Nests were located and
examined by the investigator in goose nesting habitats excluding national
wildlife refuges (Figure 1). Refuge personnel collected nesting data on the
Alamosa and Monte Vista National Wildlife Refuges, and I labeled and used
these data when they could be compared or included with my data.

�-141-

R 5 E
N

o r;-)5M\iShoA

t
I

'"

q
I-

IN

KILOMETERS

E3

E'3

8

SCALE

o

z

SCALE

o
i

E3

~OASS

A~

16

IN

E3

F'3
Z

MILES

e

10
I

5

H

I

I-

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45'

Dol/or

o
Head

'if
E

-

z

- r ~
G I-

t.as»

H

Son ca/s

t.os»

•....•

z

I-

K

Dry
Loses

...•
I-

L

37·
30'

@z
•..
MONTE

...•

VISTA

I-

NATIONAL
WILDLIFE

•

Transects

surveyed

total

breeding

April

1974

Transects

surveyed

geese

on fiver

wing

and

August

pairs

for estimate
from

for

systems

hell-copter

of

helicopter,

estimate
from

REFUGE

R

[J-

of

fixed-

aircraft,

1974

..

z

...•
Manassa
R5E

R4E

R 9E.

10'5° 30

Figure

2.

Transects
used in aerial and ground surveys
of Canada
geese in the San Luis Valley,
1973 and 1974.

�-142-

1.6

km

( I mile)

(

\

i

I

I
Transect

I
!

~

Flight

Pattern

I

I

I

I

I

I

-~i
I

I

I

Figure 3.

I~

I

I
I

I

I

I
I

I
I

I
t
I

I
I
I

t
I

I

I

I

I

I

I

I

I

I

I

I
I

I

I

I

I

\

I

I
I

~

I
I

~

I

Scanned

I

I
I
I

Area

I
~

,f.

In One Pass

I
I

I

I

.

I

(

I
I

I

I

I
I

I

I
I

I
~

I
~

I

I

I

I

I

I

I

I

I

I
I

I

I

t

••....
----f

Flight pattern of lengthwise passes in a transect for aerial
surveys of Canada geese in the San Luis Valley, 1973 and
1974.

�-143-

45'

R 5 E
N

f
SCALE

IN

o

o Misho~
Lases

o

Wsismon
LoAo

KILOMETERS
9

16

1E3E3E3E3
SCALE

COo

~
~
z

IN MILES

o

5

e
&gt;-

10

tl:Et=1=r~E3~:jI~========31
o Center
Dol/or

o
Heod

O'~LOAO
Z

LoA

~

&gt;--

L
Son Luis

t.os»

Z

RIO GRANDE
WILDLIFE

MONTE

VISTA

NATIONAL
WILDLIFE
~

Areas

Studied

in 1973

~

A reas

Studied

in 1974

~

Areas

Studied

in Bo tb Years

REFUGE
NATIONAL
WI LOLIFE
REFUGE

.•,.,..
z

z
on
15'

z

"•..

R ~E.

R4£
106° 30'

Figure

4.

Areas where Canada goose goslings were studied for
estimates
of mortality
in the San Luis Valley,
1973
and 1974.

�-144-

Aerial and Ground Survey Comparisons
Aerial counts were compared with ground counts on the same selected areas
in order to evaluate the aerial counts. Ground counts on the same areas
counted from the air were made by the aerial observer(s) or by a team of
ground observers before or after the aerial count, or by teams of ground
observers during the aerial count. Ratios of air-ground counts were tested
using Chi-square tests with Yates correction factor for small samples or
lack of continuity.

Breeding Pair Counts
Aerial counts were compared with ground counts taken within one week of the
flights. The estimated number of breeding pairs from aerial observations on
each transect were compared with the actual number of nesting geese found
during the ground counts.

Brood Counts
Corresponding aerial and ground counts of flightless geese were conducted
simultaneously or within four hours on river bottom transects and within one
day on isolated lake and pond areas. Numbers of goslings and adults recorded
in both aerial and ground counts were compared.

Post Brood-rearing Counts
Corresponding aerial and ground counts of geese capable of flight were
simultaneous or within four hours. Numbers of total geese, combined adults
and goslings, recorded in both aerial and ground counts were compared.

Aerial Population Surveys
Breeding Pair Counts
The number of breeding pairs and the total goose population of the Valley,
excluding the Monte Vista National Wildlife Refuge, were estimated from
helicopter counts on 22 and 23 April 1974. Open lake and pond areas known
to have nest~ng geese were counted. River areas were sampled by counting within
all transect'; of sections believed to hold the majority of nesting geese
(transects 2i through 34, 42 through 45, 50 through 53, and 63), and by counting
within randomly drawn transects of remaining sections believed to hold few if
any nesting geese (Figure 2).
Helicopter-ground comparison results for all different areas (Tables 1, 2 and
3) were used in the estimation, including results in open river areas where air/
ground ratios between observations were assumed to be consistent. First,
breeding pairs were estimated on each transect sampled, then estimated within
the different river sections or lake and pond areas. Then, each section's
or area's total estimate was adjusted according to the habitat involved
(multiplied by the appropriate ground/air ratio: 1.00 in wooded areas, 1.16
in open lake and pond areas, and 1.75 in open river areas). Finally, estimates
for all sections and areas were totaled.

�-145-

Table 1. Aerial estimates of breeding pairs of Canada geese in wooded river
areas compared with ground counts on the same areas in the San Luis Valley,
1974.

Area

1

Active Breeding Pairs (No. of Active Nests)
Ground Count (Within
Estimate from Helicopter
One Week of Flight)
Count (23 April 1974)

3
6
13
22(S)
25(S)
26
27
29(N)
34(N)

1
1
1
1
6
4
2
2
2

Totals
Air/Ground Ratio

20

Area

1

1
2
1
2
7
3
2
1
1

Air/ground ratios
were consistent
for repeated counts
1.49
X2
8
d. L
0.
0.05

20
100.0%

Active Breeding Pairs (No. of Active Nests)
Ground Count (Within
Estimate from Fixed-Wing
One Week of Flight)
Count ( 1 May 1974)

3
6
13
25(S)
26
27

0
2
1
1
1
3

Totals
Air/Ground Ratio

8

0
2
1
6
1
2

Air/ground ratios
were consistent
for repeated counts
X2
4.13
a.r ,
4
0.
0.05

12
66.7%

lArea numbers refer to numbered transects in Figure 2, and (N) or (S) indicates
the north or south side of the river where aerial-ground comparisons were
made which did not include the entire transect.

1

�-146-

Table 2. Aerial estimates of breeding pairs of Canada geese in open lake and
pond areas compared with ground counts on the same areas in the San Luis
Valley, 1974.

Area

1

Active Breeding Pairs (No. of Active Nests)
Estimate from Helicopter
Ground Count (Within
Count (22 April 1974)
One Week of Flight)

A
B
C

1
5
9
0
0
1
2
1
1
0
2
2
1

D

E

F
G,H
I

L
0
P
30(N)
39(S)
Totals
Air/Ground Ratio

Area

1

25

Air/ground ratios
were consistent
for repeated counts
2.16
X2
d. f.
11
a
0.05

29
86.2%

Act tve Breeding Pairs (No. of Active Nests)
Estimate from Fixed-Wing
Ground Count (Within
Count ( 1 May 1974)
One Week of Flight)

B,C
D

P
30(N)
Totals
Air/Ground Ratio
1

1
5
10
1
0
1
2
1
1
1
2
3
1

9
1
1
2

17
1
2
1

12

21

Air/ground ratios
were consistent
for repeated counts
2
3.93
X
d. f.
3
a
0.05

61.9%

Area letters or numbers refer to lettered or numbered transects in Figure 2,
and (N) or (S) indicates the north or south extreme of a river transect where
an open lake or pond area was found.

�-147-

Table 3. Aerial estimates of breeding pairs of Canada geese in open river
areas compared with ground counts on the same areas in the San Luis Valley,
1974.

Area

1

Active Breeding Pairs (No. of Active Nests)
Estimate from Helicopter
Ground Count (Within2
Count (23 April 1974)
One Week of Flight)

42
43
44
45

Totals
Air/Ground Ratio

1
1

4
2
8

14
57.1%

lArea numbers refer to numbered transects in Figure 2.
2Data collected by cooperating observers was not comparable within each
separate transect.

�-148-

Post Brood-rearing Counts
The river population of adults and goslings capable of flight was sampled
from a fixed-wing count on 22 August 1974 and from helicopter counts on 27
and 28 August 1974. River areas were sampled in two ways. Aerial counts of
geese were made within every other transect of sections believed to hold
geese (transects 20 through 36, 42 through 45, and 50 through 53), based on
recent ground observations. Also, river bottom sections adjacent to the
sampled sections were scanned supetficially.
If geese were observed on
those sections they also were flown and counts included in the survey.
Then fixed-wing and helicopter counts of geese on the same river transects
and areas (Figure 2) were compared.
During 27 and 28 August 1974, helicopter counts of geese were also conducted
on all lake and pond transects excluding the Monte Vista National Wildlife
Refuge (Figure 2). The total goose population of the Valley was then estimated from the helicopter counts and reported numbers of geese on the refuge.
For helicopter estimates, it was assumed that all geese were seen and accurately
counted in all transects and river bottom sections which were sampled.

RESULTS AND DISCUSSION
Delineation of Nesting Habitat

Habitats used for nesting included river, lake and pond areas (Figure 5).
River bottoms With adjacent ponds or sloughs were preferred along river
courses. Lakes and ponds with islands or emergent vegetation were favored
open water areas.
Although nesting geese were found in many different areas of the Valley,
some nesting probably took place in additional areas and was not detected.
Possible nesting areas were those where geese were observed b~t nesting
was not confirmed during either or both nesting seasons (compare Figures 5
and 6). Areas where many geese were observed, along the Rio Grande River
near La Sauses and between Monte Vista and Alamosa, were considered likely
nesting areas. Examination from the ground of all possible nesting sites
during both nesting seasons was not possible because of the vastness of the
study area, and denial of access to certain lands.

Potential Nesting Habitat
Potential goose nesting habitat was abundant, but some areas are more
likely to be used than others. It seems likely that geese would prefer
potential areas similar to those in which they actually nested. Therefore,
river areas with adjacent sloughs and ponds, and lake and pond areas with
emergent vegetation or islands are more attractive than wetlands without those
features.

�-149-

,~.
R 6 E

R 5 E

R 4E

4~'
R7E
25 {o9i

II

z

LN

o

'"•...

"

(II

LokOS

~MiS/JO"
LO"8S

i
SCALE

Lose

KILOMETERS

a
SCALE

IN

a

.....

I E3 F3

.....
Z

IV/lisman

16

iE3E3E3E'3

z

R.ItE.

RIOE

Russoll

~Q.

~

N

R9E

R8E

N

COo

~
~

z

..

MILES

...

00

5

I

I

o Center

4~'

45'

Dol/or
0 Lase

o

..

z

Head

0

(o)

z
~

~

Lake ~

2(2)

~(3J~

Son Luis

c as«

•.'"
z

RIO

GRANDE

•...

WILDLIFE
MANAGEMENT

AREA

o

Monte

·"t

Dry
Lakes
I (3)

.•...'"

z

~

()

&amp;

.•'"
z

1(1)

II

]0'

z
•..

z
•...
...'"

•...
'"

•

Approximate

nestrnq

found

on 1973

of nests found

in 1974

Number of nests

z

area
z

'"'",.

(3) Number

••
'"•...
z

..,

z

'"•...

''""

I~'
I~'

..'"

..'"

Z

z

Monosso 0

"

R5E

R4E

R6E

R7E.

RaE

RilE

R 9 E.

106°30'

Figure

5. Areas where nesting Canada geese
San Luis Valley.
1973 and 1974.

were

found in the

�-150-

N
15'
COLORADO

SAN LUIS
vALLEY

SCALE

IN

KILOMETERS

o

8

H

H

SCALE

IN

o
I

16

H

HI

t=1

H

38°
00'

MILES

~

10

F""""SS"'3

Center
45'

RIO

d)pry

LEGEND

o

-9
-0

GRANDE

~La"{/s
Town
US

37°
00'

Highway

State

Highway

App r ox imote

CJ

Refuge,

~

Areas

2469 m Contour
(8100ft)

Management
Where

Geese

Area
Were Observed

15'

37°
COLORADO

NEWMEx'iCO--- -15'

Figure

6.

on'

---.5'

Areas where Canada geese were observed during nesting
seasons in the San Luis Valley, 1973 and 1974.

�-151-

Williams and Nelson (1943) described major requirements of good Canada
goose-breeding areas as:
••
.(1) a browsing area available to nesting birds
and to paired adults prior to the season; (2) an aquatic feeding
area during the breeding period; (3) a brooding environment
of open water and resting banks; (4) molting cover (emergents);
(5) a browsing area for broods after they are on the wing
(this may be the same as 1); (6) nesting sites isolated from
interference; (7) nesting sites providing firm foundations;
(8) nesting sites with good to excellent visibility. All these
items seem to be essential, or at least very important, in determining breeding populations. Nesting sites are also selected for
their proximity to channels and to open ponds that provide
avenues to the breeding areas."

San Luis Lake lacked adequate nesting sites and most nesting attempts
were unsuccessful.
Properly maintained, wind-firm structures or man-made,
wave-firm islands may increase gosling production there substantially.
~hak Lakes lacked water during the brooding period, and emergents for
molting cover were lacking due to non-permanent water. Water wells with
adequate flow and properly placed dikes would increase the potential for goose
nesting. Dry Lakes was similar to Misrnk Lakes but also lacked nesting sites.
There, the Bureau of Land Management, has been developing waterfowl production
areas in recent years through manipulation of water, and construction of
nesting islands and properly maintained nesting structures. As a result,
geese first nested there in 1973 and produced two goslings to flight stage
(Tragstaad, unpublished data), and gosling production increased to 14
in 1974 (Strong, unpublished data).
The Rio Grande River Canyon from La Sauses to the Colorado-New Mexico state
line may, in places, be suitable for geese willing to accept rocky ledges or
cliffs as nesting sites. However, use of these sites is not common and the
potential for high production within this area is uncertain.

Aerial Survey Techniques
During aerial surveys of geese, aircraft were flown at various speeds and
heights above the ground. The lower heights and slower speeds were favored
for observing and counting geese. Others have expressed the need to fly
low and slow for efficient aerial surveys (Ballou 1956, Benson 1962).
Transects and flight patterns in the Valley were different from those of
more extensive waterfowl breeding pair surveys in the prairie states and
provinces of the United States and Canada (Crissey 1957, Kaczynski and
Chamberlain 1968). The more extensive aerial surveys were conducted along
0.4 km (0.25 mi) wide transects over which just one pass was made with
aircraft, requiring the observers to scan out to 200 m (220 yds) from the
aircraft. However, obtaining an index to the size of the fall population
was the major objective of each of the studies.

�-152-

In my study, goose habits and wind conditions were primary factors in deciding
the time of day that survey flights were made. During counts of breeding
pairs (0730 to 1430 DST) and flightless broods (0800 to 1630 DST), the main
concern was to avoid rough air conditions which occurred mostly during
afternoons. Throughout the day, both breeding pairs and flightless broods
were essentially confined to the wetland areas; and therefore, their movements
did not influence time of count. However, post brood-rearing counts were
conducted only during midday (1000 to 1330 DST). Geese flew less and spent
more time on the water areas during midday than in early mornings or late
afternoons. The importance of time-of-day for aerial surveys of waterfowl
was also pointed out by others (Ballou 1956, Diem and Lu 1960).

Aerial and Ground Survey Comparisons
Diem and Lu (1960) pointed out that countability of waterfowl from aerial
surveys was influenced by many interacting factors. However, comparisons
of counts from air and ground make it possible to correct these influencing
factors. Aerial and ground counts, on the same area have been used since
1959 in Canada and the northern U. S. to calculate the proportion of waterfowl
observed by aerial crews (Martinson and Kaczynski 1967). The basic assumptions
were that the same waterfowl were counted from the air and the ground, and
that the ground counts were complete censuses. Results of aerial counts of
Canada geese in my study were also evaluated according to comparisons
with ground count.

Breeding Pair Counts
Estimates of breeding pairs from aerial counts best agreed with ground counts
on the same areas, within each different habitat using a helicopter. However,
estimates in open lake or pond areas and in wooded river areas from both
fixed-wing and helicopter aircraft produced air/ground ratios that were consistent (Cl. = 0.05) for repeated counts (Tables 1 and 2).
The estimate of breeding pairs from the helicopter on open river areas
agreed less with the ground count on the same areas than other aerial
estimates on different habitats (compare Tables 1 and 2 with 3). Chi-square
tests were not possible for aerial estimates in open river areas due to
lack of specific ground count data in each transect of the helicopter count
(Table 3).
Results of aerial breeding pair estimates and ground comparison counts (Tables
1 and 2) indicated that aerial estimates without correction from helicopter
counts in wooded area, and fixed-wing or helicopter estimates with correction
in different habitats were similar to ground counts. Accordingly, these
data could be used reliably for annual estimates of breeding pairs.
My results of the helicopter estimate and ground count on open river transects
indicated that the aerial count in that type of habitat was the least accurate

�-153-

of all aerial estimates of breeding pairs on different habitats (Tables 1,
2 and 3). The open river transects which were counted from air and ground
held vast areas of preferred nesting habitat as opposed to more concentrated
areas of preferred nesting habitat in other types of transects. Also,
these transects occurred on the Alamosa National Wildlife Refuge and I was
unfamiliar with the area having personally collected no ground data.
Therefore, it was possible that by not knowing where to concentrate observations in these transects, that the air/ground ratio of breeding pairs was
not similar to air/ground ratios in other habitat types, when counting from
a helicopter.
I was confident that the ground counts of breeding pairs were accurate and
complete within each area of comparison with aerial counts. This was because
nesting geese were confined to a specific site until termination of nesting,
and the status of most active nests did not change between aerial and ground
coverages. This allowed time for thorough ground searches in several different
areas.

Brood Counts
Aerial counts of flightless goslings and adults varied greatly from ground
counts. Air/ground ratios were inconsistent (a = 0.05) for repeated counts
with both aircraft types in both open and wooded areas during both years
(Tables 4, 5, 6 and 7) and indicated that aerial counts could not be
corrected to compare with ground counts and that these data could not be
used reliably for annual productivity estimates.

Post Brood-rearing Counts
Counts from fixed-wing aircraft of geese capable of flight varied from
ground counts (Tables 8 and 9). Air/ground ratios were consistent
(a = 0.05) for repeated counts only in the 1974 counts in open areas
(Table 9). No data comparing counts from helicopter with ground counts
were collected since many geese counted from the air flew out of the counting
area before ground counts could be made. Thus, only counts from fixed-wing
airsraft in open areas could be corrected to compare with ground counts
and that only these data could be used reliably for annual estimates of
goslings and adults combined.
The ground comparison counts of geese during the brood-rearing period and
post brood-rearing period were not reliable. The flightless geese were
restricted in movement but were secretive and many times not heard or seen
in the dense vegetation of brood-rearing areas. This was especially true
for ground observers who were unfamiliar with the study areas. Many times,
if I had not known from prior observations that flightless goslings and
adults were probably in a certain area, more geese would have gone unnoticed.
Geese capable of flight were easier to see and count, but their mobility
made comparisons of aerial-ground counts unreliable. The ground observer(s)
could never be certain that individuals had not left or joined a group of
geese unless ground and aerial counts were simultaneous. Unfortunately, only
three comparison counts of geese capable of flight were simultaneous
(Tables 8 and 9).

�Table 4. Fixed-wing counts of flightless Canada geese in wooded areas compared with ground counts
on the same areas in the San Luis Valley, 1973 and 1974.

Area

1

22(S)
23(S)
29(S)
30(S)
32 (S)
33(S)
34(N)

Fixed-wing Count
Adults Goslings

Date
18 June
18 June
18 June
18 June
18 June
18 June
18 June

73
73
73
73
73
73
73

Sub-totals

2
Ground Count
Adults
Goslin~s

14
14
22
25
31
2
6

0
0
0
0
0
0
0

113
03
03
03
03
03
2

753
03
03
03
03
03
3

114

0

l3

78

Air/Ground
Adults

Percentage
Goslings

Air/ground ratios were not
consistent for repeated counts
2
X
= 6.69
X2
= 76.09
d.E. = 1
d.E. = 1
CI.
CI.
= 0.01
= 0.005

876.9

0.0
I
•...•

.,..

'-J1

30(S)
25(S)
27(S)
22(S)
23 (S)

18 June
18 June
18 June
18 June
18 June

Sub-totals

74
74
74
74
74

0
0
0
0
5

0
0
0
0
12

10
6
0
14
10

10
26
0
10
15

Air/ground ratios were not
consistent for repeated counts
X2
= 29.11
X2
= 43.48
d.f. = 3
d.f. = 3
CI.
= 0.005
CI.
= 0.005

5

12

40

61

12.5

19.8

lArea numbers refer to numbered transects in Figure 2, and (N) or (S) indicates the north or south
side of the river where aerial-ground comparisons were made which did not include the entire transect.
2Ground counts were simultaneous

with aerial counts or within 4 hours.

3Ground counts were not made by observers who made respective

aerial counts.

I

�Table 5. Fixed-wing counts of flightless Canada geese in open areas compared with ground counts on
the same areas in the San Luis Valley, 1973 and 1974.

Area

1

B,C
E,F
G,H
I

M

Fixed-wins Count
Adults Goslinss

Date
30 May 73
30 May 73
30 May 73
30 May 73
18 June 73

Sub-totals

B,C
E,F
G,H
30(N)

18 June
18 June
18 June
18 June

Sub-totals

74
74
74
74

2
Ground Count
Goslings
Adults

Air/Ground
Adults

Percentase
Goslings

56
0
27
3
36

20
0
0
0
0

70
0
43
5
3
67

53
0
0
0
3
35

Air/ground ratios were not
consistent for repeated counts
= 53.94
X2
2
= 22.52
X
d.L = 1
d.L = 3
a.
= 0.005
a.
= 0.005

122

20

185

88

65.9

Air/ground ratios were not
consistent for repeated counts
= 12.22
X2
2
= 79.70
X
d.L
= 3
d.L = 3
a.
a.
= 0.01
= 0.005

19
34
10
0

28
2
0
0

46
35
81
4

34
1

63

30

166

48

5

8

37.9

22.7

62.5

1Area numbers or letters refer to lettered or numbered transects in Figure 2, and (N) indicates the
north side of the river where aerial-ground comparisons were made which did not include the entire
transect.
2Ground counts were simultaneous with aerial counts, or within 4 hours on river bottom transects,
or within 1 day on isolated lake and pond areas.
3Ground counts were not made by observers who made respective

aerial counts.

I
•....
VI
VI

I

�Table 6. Helicopter counts of flightless Canada geese in wooded areas compared with ground counts
on the same areas in the San Luis Valley, 1974.

1
Area

Date

6(N)
6(N)
21(S)
22(S)
22(S)
23(S)
25(S)
25(S)
30(S)

11 June 74
12 June 74
11 June 74
11 June 74
12 June 74
11 June 74
11 June 74
12 June 74
11 June 74

Sub-totals

Helicopter Count
Adults Goslings

2
Ground Count
Adults
Goslings

Air/Ground
Adults

Percentage
Goslings

03
53
2
113
103
83
03
0
6

03
133
0
103
93
113
03
0
5

5
5
2
11
11
84
6
6
12

14
14
4
10
10
134
26
26
18

Air/ground ratios were not
consistent for repeated counts
X2
= 16.90
X2
= 75.02
d.f. = 8
d.f. = 8
a
= 0.05
a
= 0.005

42

48

66

135

63.6

35.6

lArea numbers refer to numbered transects in Figure 2, and (N) or (S) indicates the north or south
side of the river where aerial-ground comparisons were made which did not include the entire transect.
2Ground counts were simultaneous with aerial counts, or within 4 hours on river bottom transects, or
within 1 day on isolated lake and pond areas.
3Aerial counts on same areas on different dates were by different observers.

4

Ground count shown was by observer who was familiar with the habitat, and the ground count of same
area by observer unfamiliar with the habitat was 0 adults and 0 goslings.

I
•....
Ln
0\.
I

�Table 7. Helicopter counts of flightless Canada geese in open areas compared with ground counts on
the same areas in the San Luis Valley, 1974.

Area

1

B

C
E
L
M
0

30(N)

HelicoEter Count
Adults Goslings

Date
10 June
10 June
10 June
10 June
10 June
10 June
10 June

Sub-totals

74
74
74
74
74
74
74

2
Ground Count
Goslings
Adults

Air/Ground
Adults

Percentage
Goslings

5
18
12
0
15
2
0

12
21
4
0
0
3
0

10
26
8
0
27
2
3

14
44
4
0
13
3
6

Air/ground ratios were not
consistent for repeated counts
X2
= 28.87
= 12.83
X2
d.L = 5
d.L = 5
a
a
= 0.005
= 0.05

52

40

76

84

68.4

47.6

.....I
lArea numbers or letters refer to lettered or numbered transects in Figure 2, and (N) indicates the
north side of the river where aerial-ground comparisons were made which did not include the entire
transect.
2Ground counts were simultaneous with aerial counts, or within 4 hours on river bottom transects,
or within 1 day on isolated lake and pond areas.

\.II

-...J
I

�-158-

Table 8. Fixed-wing counts of Canada geese capable of flight in wooded
areas compared with ground counts on the same areas in the San Luis
Valley, 1973 and 1974.

Area

1

22(S)
22(S)
22(S)
31(N)
32(S)
37(N)

Flight
Date
9 Jul
13 Aug
22 Aug
15 Aug
15 Aug
22 Aug

73
73
73
73
73
73

Sub-totals

30(S)
31(N)
34(N)
M

23 Aug
23 Aug
23 Aug
23 Aug

Sub-totals

74
74
74
74

Counts
Fixed-wing

Ground

2

Air/Ground

Percentage

39
19
10
26
47
118

119
120
16
66
93
151

Air/ground ratios were
not consistent for
repeated counts
2
192.06
X
d. f.
5
a.
0.005

259

565

45.8

273

353

6

o

20
17

20
105

Air/ground ratios were
not consistent for
repeated counts
X2
74.54
d.f.
2
a.
0.005

70

160

43.8

lArea numbers or letters refer to numbered or lettered transects in Figure 2,
and (N) or (S) indicates the north or south side of the river where aerialground comparisons were made which did not include the entire transects.
2

Ground counts were simultaneous with aerial counts, or within 4 hours.

3Aerial and ground counts were simultaneous.

�-159-

Table 9. Fixed-wing counts of Canada geese capable of flight in open areas
compared with ground counts on the same areas in the San Luis Valley, 1973
and 1974.

Area

1

M
M

Q
30(N)
30(N)

Flight
Date
13 Aug
15 Aug
9 Ju1
13 Aug
15 Aug

73
73
73
73
73

Sub-totals

M

N
N
31(S)
34(N)
35(N)

23 Aug 74
23 Aug 74
23 Aug 74
23 Aug 74
23 Aug 74
23 Aug 74

Sub-totals

Counts
Fixed-win~

Ground

49
86
25
22
25

75
75
28
20
7

Air/ground ratios were
not consistent for
repeated counts
54.23
X2
d .f.
4
a.
0.005

207

205

101.0

183
923
10
30
37
28

20
1103
123
31
34
42

Air/ground ratios were
consistent for
repeated counts
X2
7.61
d.f.
5
a.
0.05

215

249

86.3

2

Air/Ground Percenta~e

1Area numbers or letters refer to numbered or lettered transects in Figure 2,
and (N) or (S) indicates the north or south side of the river where aeria1ground comparisons were made which did not include the entire transect.
2Ground counts were simultaneous with aerial counts, or within 4 hours.
3Aeria1 and ground counts were simultaneous.

�-160-

Aerial Population Surveys

Breeding Pair Counts
From helicopter counts on 22 and 23 April 1974 (Figure 2), 115 breeding
pairs with active nests were estimated to be in the Valley, excluding
the Monte Vista National Wildlife Refuge; 76 breeding pairs were estimated
on the river systems and 39 on the lake and pond areas. This count included
estimates on open river transects where reliability of aerial counts had not
been thoroughly evaluated and, therefore, left in doubt. However, geese
were only found in such transects within the Alamosa National Wildlife Refuge,
and the estimate from aerial counts was corrected through ground counts by
refuge personnel. Therefore, the breeding pair estimate from helicopter
counts was consider~d reliable, though valid statistical confidence intervals
were not calculated due to sampling procedure.
Excluding the Monte Vista National Wildlife Refuge, 730 breeding and nonbreeding geese were estimated to be in the Valley from the same helicopter
counts, with 170 geese on lake and pond areas and 560 on river areas. Including
refuge estimates (Bryant, unpublished data), total geese numbered approximately 1,030 in the entire San Luis Valley.
Estimates of geese other than
breeding pairs were not compared with ground counts and not evaluated as to
reliability. It seemed that during the aerial count, most geese which
were present on each sa~pled transect were seen and counted. The dormant
and matted vegetation during this season allowed most of the surface area
of the transects to be scanned. Also, geese seen and counted were not
recounted in other transects because geese that flushed would circle and
soon return to the same general area. Therefore, I believed that the
estimate of total geese, from helicopter counts on 22 and 23 April 1974,
was valid.
Counting within every transect of certain sections of river bottom from the
helicopter was tedious for both the observer and the pilot. The continual
zigzagging pattern of flight and scanning for geese during long periods also
hindered the observer's judgement of where different transect boundaries laid.
I believe that counting within every other transect would have been easier
and less confusing.

Post Brood-rearing Counts
Counts from fixed-wing and helicopter aircraft, on the same river transects
(Figure 2), of adults and goslings capable of flight were different. Geese
seen during the fixed-wing count of 22 August 1974 totaled 271. Geese seen
during the helicopter counts of 27 and 28 August 1974 totaled 368. The
operation of the helicopter usually caused geese to fly from its path,
and geese were, therefore, seen and counted in flight. The fixed-wing
aircraft did not usually flush the geese, and many were, therefore, seen and
counted on the ground. Possibly this explains why more geese were seen
using a helicopter.
Counts from the helicopter of the same areas of river and additional lake
and pond transects totaled 438, resulting in an estimate of 625 geese on
all areas excluding the Monte Vista National Wildlife Refuge. Including

�-161-

refuge counts at this time (Cornelius, unpublished data), total geese
numbered approximately 875 in the entire San Luis Valley. These estimates
were based on the assumption that all geese within sample transects were seen
and accurately counted. However, reliability of this type of survey was not
evaluated, and the reliability of the estimates was, therefore, uncertain.
Counts within every other transect of certain sections of river bottom
from the helicopter were conducted. This method was less fatiguing for
observer and pilot and seemingly more practical than the method used in the
earlier breeding pair population survey on river bottoms.

Nesting and Gosling Mortality

Studies

Nesting Chronology
In 1973 and 1974, the earliest possible initiations of egg-laying were
24 and 18 March, the latest were 15 and 2 May, and the two-week peaks of
initiation of egg-laying were 26 March through 8 April and 23 March
through 5 April, respectively.
Average temperatures during March and
0
April were approximately 10 C (340 F) in 1973 and 40 C (39 F) in 1974.
In 1973 and 1974, the earliest possible hatching dates were 29 and 23 April,
the latest were 20 and 7 June, and the two-week peaks of hatching were
1 May through 14 May and 27 April through 10 May, respectively.
Brakhage (1965) defined the interval between the laying of the first egg
and the hatching of the last successful nest as the nesting period. The
nesting periods for 1973 and 1974 were, therefore, approximately 89 and 82
days, respectively.
By comparing initiation dates and lengths of nesting periods of Canada geese
in different areas of North America (Table 10), certain trends can be
detected. Generally, nesting starts earlier and lasts longer in southern
latitudes than in northern latitudes. However, there are exceptions to both
trends, such as the results of my two-year study (Table 10).

Nesting Sites
During both years of field study, 160 nests were found, 108 on natural sites
and 52 on artificial structures. Of the 108 nests on natural sites, 51 nests
(47.2 percent) were located in emergent vegetation of bulrush or cattails,
38 nests (35.2 percent) were on islands of lakes, ponds or rivers, 18 nests
(16.7 percent) were on mainland near water and one nest (0.9 percent) was in
a duck blind built over water. Nest bases consisted of bare ground for 53
nests (49.1 percent), muskrat houses for 29 nests (26.9 percent), matted
vegetation for 22 nests (20.4 percent), haystacks for two nests (1.9 percent),
an abandoned magpie nest for one nest (0.9 percent), and a wooden platform
for one nest (0.9 percent) (Table 11).

�Table 10. Initiation
North America.

dates and lengths of nesting periods for Canada geese at different

areas in

-Location
California
Colorado
Colorado
California
Montana
Colorado
Colorado
Illinois
Illinois
Colorado
Idaho
Wyoming
Utah
Idaho
Manitoba

Area DescriEtion
AEEroximate
Altitude
N. Latitude
1212 m
(4,OCO f t )
1545 m
(5,100 ft)
1545 m
(5,100 ft)
1212 m
(4,000 ft)
758 m
(2,500 ft)
2303 m
(7,600 ft)
1606 m
(5,300 ft)
242 m
(
800 ft)
242 m
( 800 ft)
2303 m
(7,600 ft)
1970 m
(6,500 ft)
1970 m
(6,500 ft)
1273 m
(4,200 ft)
1970 m
(6,500 ft)
242 m
( 800 ft)

Nesting Period
Initiation
Length
(Earliest)
(Da:t:s) Year (s)

Reference

40°

1 March

71

1939-40

Dow (1943)

41°

7 March

100

1968

Will (1969)

41°

12 March

88

1967

Will (1969)

42°

15 March

83

1952

Miller and Collins

48°

15 March

72

1953-54

Geis (1956)

(1953)
I
•....
0\
N
I

37°

18 March

82

1974

This study

40°

21 March

91

1966

Szymczak

(1967)

42°

23 March

68

1946

Kossack

(1950)

42°

24 March

61

1945

Kossack

(1950)

37°

24 March

89

1973

This study

43°

1 April

70

1951

Steel, Dalke and Bizeau

(1957)

43°

1 April

61

1947

Craighead

and Craighead

(1949)

41°

3 April

62

1937

Williams

and Marshall

43°

6 April

79

1950

Steel, Dalke and Bizeau

51°

9 April

61

1955

Klopman

(1958)

(1937)
(1957)

�-163-

Table 11. Natural sites and bases of Canada goose nests on San Luis Valley
study areas, 1973 and 1974.

1974
% of
No. of
Nests
Nests

1973
% of
No. of
Nests
Nests

1973 and 1974
% of
No. of
Nests
Nests

Nest Site
Emergent vegetation

25

(56.8)

26

(40.6)

51

(47.2)

Islands

14

(31.8)

24

(37.5)

38

(35.2)

Mainland

4

( 9.1)

14

(21.9)

18

(16.7)

Duck blind

1

( 2.3)

0

( 0.0)

1

( 0.9)

Totals

44

(100.0)

64

(100.0)

108

(100.0)

Nest Bases
Bare ground

16

(36.4)

37

(57.8)

53

(49.1)

Muskrat houses

10

(22.7)

19

(29.7)

29

(26.9)

Matted vegetation

15

(34.1)

7

(10.9)

22

(20.4)

Hay stacks

2

( 4.5)

0

( 0.0)

2

( 1.9)

Magpie nests

0

( 0.0)

1

( 1.6)

1

( 0.9)

Wooden platform

1

( 2.3)

0

( 0.0)

1

( 0.9)

Totals

44

(100.0)

64

(100.0)

108

(100.1)

�Table 13. Nesting success of Canada geese in different habitats
areas excluding national wildlife refuges, 1973 and 1974.

I

1973
Habitat
·2
Wooded

Both

and sites of the San Luis Valley study

Open

1974
Habitat
Wooded

Both

1973 and 1974
Habitat
Both
Open
Wooded

Nest Site

Open

Artificial Structures
No. of nests
No. successful
% successful

7
3
42.9%

22
0
0.0%

29
3
10.3%

5
4
80.0%

18
4
22.2%

23
8
34.8%

12
7
58.3%

40
4
10.0%

52
11
21.2%

Natural Sites
No. of nests
No. successful
% successful

35
19
54.3%

9
5
55.5%

44
24
54.5%

42
27
64.3%

22
9
40.9%

64
36
56.3%

77
46
59.7%

31
14
45.2%

108
60
55.6%

•

aa-

•

Total .(Both Sites)
No. of nests
No. successful
% successful
1Open habitat

42
22
52.4%

31
5
16.1%

73
27
40.0%

47
31
66.0%

40
13
32.5%

87
44
50.6%

had 10 or fewer large trees within 100 m (328 ft) radius of the nest.

2Wooded habitat

had 10 or more large trees within 100 m (328 ft) radius of the nest.

89
53
59.6%

71
18
25.4%

160
71
44.4%

�-167-

Nesting success was evaluated on the Rio Grande Wildlife Management Area
during 1973 and 1974. I found 23 nests each year, all of which were in wooded
river bottom habitat. Thirty-six of the 46 nests were on artificial standing
structures over water or land and 29 (80.6 percent) nests on structures
were destroyed. All structures were the type with four steel posts holding
three and one-half bales of hay or straw (Grieb and Crawford 1967). I
suspected the cause of destruction to be probably raccoon damage or possibly
magpie damage. In one case a goose was found dead beneath a destroyed
nest on a structure. Raccoons were probably capable of grasping the steel
posts of the structures and climbing to the nests. Magpie destruction was
possible if the nesting geese had left their eggs unattended for a short while.
Personnel of the Monte Vista and Alamosa National Wildlife Refuges contributed
data for 160 additional nests in 1973 and 1974, 93 on natural sites and 67 on
artificial structures (Bryant and Ziegler, unpublished data). Ninety-five
nests (59.4 percent) of the 160 nests were successful. Forty-six (49.5
percent) of the nests on natural sites were successful while 49 (73.1 percent)
of the nests on artificial structures were successful (Table 14). Nests
were located predominately in open habitat.
On the Alamosa National Wildlife Refuge in 1973 and 1974, 13 nests were
successful, 8 nests were destroyed and 3 nests were deserted (Ziegler, unpublished data). All nests were found on natural sites in 1973, and all nests
were found on elevated, artificial structtures which were designed to defend
nests from ground predators in 1974. The destroyed nests were on natural
sites and destruction was attributed to raccoons (Ziegler, personal communications).
On the Monte Vista National Wildlife Refuge in 1973, 52 nests were successful,
17 nests were destroyed and 10 nests were deserted (Sarvis, unpublished data).
Nine nests were destroyed by mammals and 8 nests by birds. Magpies were
believed to be the avian predators (Bryant, personal communications).
Success was calculated for nests active during the aerial-ground
comparison counts of breeding pairs. Of nests active on 22 and 23 April
1974, 60.0 percent in wooded river areas, 75.9 percent in open lake and pond
areas and 92.9 percent in open river areas were successful. Of nests active
on 1 May 1974, 75.0 percent in wooded river areas and 80.9 percent in open lake
and pond areas were successful. Peak of hatching was estimated to begin
on 27 April 1974, an intermediate date between those expressed above. Therefore,
I assumed the success of active nests at the beginning of peak hatch in 1974
to be approximately midway between those percentages calculated for the above
different dates (68 percent in wooded river areas, 78 percent in open
lake and pond areas, and at least 93 percent in open river areas).
Calculations of success for nests which were active on certain dates may be
useful for estimating total successful nests from aerial breeding pair counts.
It is important to note that in respective areas, the relative success of
nests active at the later date was higher than nests which were active earlier.
Nests nearing hatching dates had decreasing chances of disaster.

�Table 14. Nesting success of Canada geese in different sites of predominately open habitats 1 of the
Monte Vista and Alamosa National Wildlife Refuges in the San Luis Valley, 1973 and 1974.

Nest Site

MVNWR2

1973
Refuge
ANWR3

Artificial Structures
No. of nests
No. successful
% successful

28
21
75.0%

0
0
0.0%

28
21
75.0%

24
15
62.5%

15
13
86.7%

39
28
71.8%

52
36
69.2%

15
13
86.7%

67
49
73.1%

Natural Structures
No. of nests
No. successful
% successful

51
31
60.8%

9
0
0.0%

60
31
51. 7%

33
15
45.5%

0
0
0.0%

33
15
45.5%

84
46
54.8%

9
0
0.0%

93
46
49.5%

15
13
86.7%

72
43
59.7%

136
82
60.3%

24
13
54.2%

160
95
59.4%

Total (Both Sites)
No. of nests
No. successful
% successful

79
52
65.8%

9
0
0.0%

Both

88
52
59.1%

MVNWR

1974
Refuge
AmJR

Both

1973 and 1974
Refuge
MVNWR
ANWR
Both

57
30
52.6%

lOpen habitat had 10 or less large trees within 100 m (328 ft) radius of the nest.
2MVNWR stands for Monte Vista National Wildlife Refuge.
3ANWR stands for Alamosa National Wildlife Refuge.

I
I-'

~

00
I

�-169-

Clutch Sizes
Clutch sizes in all 85 completed nests found ranged from 1 to 8 eggs with
a mean of 4.9. Mean clutch size in 52 successful nests was 5.0 and in 33
unsuccessful nests was 4.9 (Table 15). Data from the Alamosa National Wildlife
Refuge in 1974 were included in these results (Ziegler, unpublished data).
Canada geese lay an average of five eggs per clutch, but sometimes lay as
few as one egg or as many as ten (Williams 1967).

Hatching Success
Hatching success in both years of the study was estimated to be 87 percent
with a mean hatch of 4.3 eggs in 84 successful nests. Hatching success
was estimated to be 46 percent with a mean hatch of 2.3 eggs in 175 successful and unsuccessful nests. Data from the Alamosa National Wildlife Refuge
in 1974 were included in these results (Ziegler, unpublished data). Estimated hatching success on the Monte Vista National Wildlife Refuge in 1973
was 52 percent with a mean hatch of 2.3 eggs in 79 successful and unsuccessful
nests (Sarvis, unpublished data).
Of 52 abandoned, unhatched eggs in otherwise successful nests, 26 (50.0
percent) were infertile or with embryos which died within the first nine
days of development and 26 (50.0 percent) were fertile. Assuming only 26
of the abandoned eggs were actually fertile, infertility and embryonic death
of eggs were each approximately 6 percent in successful nests and 3 percent
in all nests.
Of 88 abandoned, unopened eggs in deserted nests, 38 (43.2 percent) were infertile or with embryos which had died within the first nine days of development and 50 (56.8 percent) were fertile. Eggs in these deserted nests
accounted for approximately 10 percent of the eggs laid in all nests.
Of 8 abandoned, unopened eggs in otherwise destroyed nests, 6 (75.0 percent)
were infertile or with embryos within the first nine days of development and
2 (25.0 percent) were fertile. Unbroken eggs in these nests accounted for
approximately 1 percent of the eggs laid in all nests.
The above estimates of abandoned eggs totaled 17 percent of all eggs in all
nests. Failure of remaining eggs which did not hatch (37 percent of all eggs)
was attributed to egg destruction. However, some of the failure apparently
due to destruction may have resulted because eggs were first deserted and
then destroyed.

Gosling Mortality
Two methods of estimating mortalities of goslings were compared using results
(Table 16) of studies on different, isolated areas (Figure 4). Overall
gosling mortality between hatching and flight stage was estimated to be 16
percent by comparing periodic ground counts of goslings, and 31 percent by
comparing estimated numbers of eggs hatched and latest ground counts of goslings.

�-170-

Table 15. Clutch sizes of Canada goose nests in San Luis Valley study areas,
1973 and 1974.1

Year

Successful
Nests

Unsuccessful
Nests

All
Nests

22
4.8
5
2-7

18
5.3
6
3-8

40
5.1
5
2-8

30(18)2
5.1

15
4.3

45(33)2
4.8

1973
n

mean
mode
range

1974
n

mean
mode
range

6

4

6

1-7

1-7

1-7

52(40)2
5.0
5 s 6
1-7

33
4.9

85 (77) 2
4.9
6
1-8

Both Years
n

mean
mode
range

6

1-8

1

Data from the Alamosa National Wildlife Refuge in 1974 were included.

2

Mean was calculated from larger number of nests; other descriptive st~tistics
were based on smaller number of nests ,.in parenthesis, because data reported
by cooperating observers were not usable for all calculations.

�-171-

Table 16. Observed and estimated gosling mortality of Canada geese between
hatching and flight stage in San Luis Valley study areas, 1973 and 1974.

Year

No. of
Eggs 2
Hatched

Highest
Count of
Goslings

Lowest
Count of
Goslings

Estimated
3
Mortalit~

Observed
4
Mortalit~

A

1973
1974

61
99

58
72

51
44

16%
56%

12%
39%

B

1973
1974

8
13

7
9

4
9

50%
31%

43%
0%

C

1973
1974

4
13

3
14

2
14

50%
- 7%

33%
0%

D

1973
1974

11

4

4

10
4

0%

9%
0%

1973
1974

4

4
3

4
3

25%

0%
0%

1973
1974

4

5
4

5
4

0%

0%
0%

Area

E

F

1

G

1973

24

22

8%

H

1973

2

2

0%

I

1973

15

15

0%

J

1974

31

28

26

16%

7%

K

1974

13

13

13

0%

0%

L

1974

9

9

8

11%

11%

285

240

Comparable Observed Mortality
Comparable Esti- 263
mated Mortality

182

16%
31%

1
Letters refer to areas in Figure 4.
2
Number of eggs hatched was estimated by multiplying number of successful
nests
in the area times mean number of eggs hatched in
all successful nests in that
year.
3Estimated mortality based on estimated number of
eggs hatched and latest count
of goslings.
40bserved mortality based on earliest and latest counts of goslings.

�-172-

Most gosling mortality is believed to take place early in life (Williams
1967). In counting goslings early in the season for the best estimate of total
goslings hatched, some mortality had probably already occurred. Therefore, I
believed that my mortality figures which included only periodic counts of
goslings were less reliable than the mortality figures which included estimates
of numbers of goslings that had hatched in certain areas (Table 15).

LITERATURE CITED

Ballou, R. M. 1956. Nesting, distribution and mortality studies of Canada
geese. Wyoming Fish and Game Comm. P-R Proj. Completion Rep., W-50-R-5.
pp. 47-77.
Benson, D. A. 1962. Use of aerial surveys by the Canadian Wildlife Service.
Canadian Wildl. Servo Occas. Pap. 3. 35 pp.
Brakhage, G. K. 1965. Biology and behavior of tub-nesting Canada geese.
J. Wildl. Manage. 29(4):751-771.
Cooper, J. A., and B. D. Batt. 1972. Criteria for aging giant Canada goose
embryos. J. Wildl. Manage. 36(4):1267-1270.
Craighead, F. C., Jr., and J. J. Craighead. 1949. Nesting Canada geese
on the upper Snake River. J. Wildl. Manage. 13(1):51-64.
Crissey, W. F. 1957. Forecasting waterfowl harvest by flyways.
Am. Wildl. Conf. 22:256-268.

Trans. N.

Diem, K. L., and K. H. Lu. 1960. Factors influencing waterfowl censuses
in the parklands, Alberta, Canada. J. Wildl. Manage. 24(2):113-133.
Dow, J. S. 1943. A study of nesting Canada geese in Honey Lake Valley,
California. California Fish and Game 29(1):3-18.
Geis, M. B. 1956. Productivity of Canada geese in the Flathead Valley,
Montana. J. Wildl. Manage. 20(4):409-419.
Grieb, J. R., and G. I. Crawford. 1967. Nesting structures for Canada geese.
Colorado Game, Fish and Parks Dept. Game Information Leafl. 48. 4 pp.
Kaczynski, C. F., and E. B. Chamberlain.
1968. Aerial surveys of Canada
geese and black ducks in eastern Canada. U. S. Fish and Wildl. Servo
Spec. Sei. Rep. Wi1dl. 118. 29 pp.
Kerbes, R. J., M. A. Ogilvie, and H. Boyd. 1971. Pink-footed geese of
Iceland and Greenland; a population review based on an aerial survey
of Thorsarver in June, 1970. Wildfowl 22:5-17.
Klopman, R. B.
Manitoba.

1958. The nesting of the Canada goose at Dog Lake,
Wilson Bull. 70(2):168-183.

�-173-

LITERATURE CITED (continued)

Kossack, C. W. 1950. Breeding habits of the Canada goose under refuge
conditions. Am. MidI. Nat. 43(3):627-649.
Lowry, D. G. 1973. Investigation of Canada geese in the San Luis Valley.
Colo. Div. Wildl., Game Res. Rept., Fed. Aid Proj. W-88-R. Oct.
pp. 125-136.
Martinson, R. K., and C. F. Kaczynski.
1967. Factors influencing waterfowl
counts on aerial surveys, 1961-1966. U. S. Fish and Wildl. Servo
Spec. Sci. Rep. Wildl. 105. 78 pp.
Miller, A. W., and B. D. Collins. 1953. A nesting study of Canada geese on
Tule Lake and Lower Klamath National Wildlife Refuges, Siskiyou
County, California. California Fish and Game 39(3):385-396.
Steel, P. E., P. D. Dalke, and E. G. Bizeau. 1957. Canada goose production
at Gray's Lake, Idaho, 1949-1951. J. Wildl. Manage. 21(1):38-41.
Szymczak, M. R. 1967. Breeding biology of Canada geese of the Metro-Denver
area. M. S. Thesis. Colorado State Univ. 83 pp.
Will, G. C. 1969. Productivity of Canada geese in Larimer County,
Colorado. M. S. Thesis. Colorado State Univ. 144 pp.
Williams, C. S. 1967.
N.J. 179 pp ,

Honker.

D. Van Nostrand Co., Inc., Princeton,

, and W. H. Marshall.
1937. Goose nesting studies on Bear River
-----Migratory Waterfowl Refuge. J. Wildl. Manage. 1(3-4):77-86.
, and M. C. Nelson.

Prepared by

1943.

Canada goose nests and eggs.

~~.~Dennis G. Lowry
Graduate Student

Auk 60(3):341-345.

��-175October, 1975
JOB PROGRESS REPORT

State of

COLORADO
-----------------------------W-88-R-ZO

Project No.
Work Plan No.
Job Title

Bird Investigations

3

6

Investigation of Mallard Management Units of Eastern Colorado

Period Covered:
Personnel:

Migratory

April 1, 1974 through March 31, 1975

S. Bissell, J. Carsella, G. Claassen, R. Clark, J. Corey, D. Coven,
C. Crawford, M. DePra, R. Desilet, G. East, M. Elkins, G. Eyre, R.
Forbes, H. Funk, A. Hemmert, J. Hicks, G. Lorentzson, R. Mason, K.
Miller, R. Oakleaf, B. Peterson, F. Rinella, Jr., R. Rouse, W. Russell,
G. Saville, H. Spear, S. Steinert, M. Szymczak, K. Wagner, C.
Wetherill, and R. Hopper.
ABSTRACT

Post-season mallard trapping efforts resulted in the banding of 6,558 birds
in nine management units of eastern Colorado during the winter of 1974-75.
Banding quotas of 800-1,000 mallards were reached in all units except for
those in the Arkansas Valley.
The January inventory of mallards in eastern Colorado (28l,972) was well
above that of the previous year. November and December counts were incomplete.
Banding and recovery tapes were obtained from the Bird Banding Laboratory for
mallards banded post-season from the winter of 1963-64 through the winter of
1971-72, and for those recovered through the 1972-73 hunting season.· Over
51,000 mallards were banded during this period, resulting in nearly 4,400
recoveries.
Some major conclusions regarding the geographic distribution of mallard band
recoveries include: (1) eastern Colorado was the main recovery location for
all four age and sex classes, (2) males were less likely to be recovered in
Canada than females, and (3) adult males were more likely to be recovered in
eastern Colorado and the High Plains Region as a whole than any other age and
sex class.
Mean survival rates were high for both sexes, being about 75 percent for males
and 66 percent for females. There appeared to be no difference in recovery
or survival rates between immature and adult birds. The mean life span for
males was estimated at 3.43 years and at 2.28 years for females.

��-177-

INVESTIGATION OF MALLARD ~~GEMENT
UNITS OF EASTERN COLORADO
Richard M. Hopper

This report summarizes aerial survey and post-season banding activities
during Segment 20 as part of the study of terminal wintering populations
of mallards in eastern Colorado.
Also included is progress made on the
analysis of band recovery data. Past contributions of this study as part
of a cooperative endeavor with other Central Flyway States, were reviewed
in a previous report (Hopper 1973).

P. S. OBJECTIVE
To develop a harvest formula for eastern Colorado management

units.

SEGMENT OBJECTIVES
1.

To determine mortality rates for wintering mallards in each of nine
eastern Colorado study areas from banding data collected during the
1963-64 through 1973-74 hunting seasons.

2.

To determine the feasibility of requesting special hunting regulations
for individual mallard management units within Colorado.

3.

To trap, band, age, and sex samples of mallards during winter in each
management unit of study, based on results of final analysis, of
sufficient size to allow adequate monitoring of each population.

METHODS AND MATERIALS
Banding and recovery tapes were requested from the Bird Banding Laboratory
in April 1974. Specifications for these tapes were:
Banding tape - only normal, wild-caught mallards banded under
permit no. 6529 in Colorado east of longitude 1050 30' during
December, January and February of 1963-64 through 1971-72.
Recovery tape - only recoveries from mallard bandings specified
for above banding tape, recovered anywhere, and shot and/or found
dead during September 1 through January 31 of the 1964-65 through
1972-73 hunting seasons.
These tapes were received in May 1974 and existing computer programs were
employed under contract with Colorado State University to produce desired
printouts of numbers banded, recovery tables, and recovery distributions.
Computer programs designed to make survival rate estimates of bird populations using methods developed by Brownie and Robson (1974), Seber (1970)

�-178-

and Robson and Youngs (1971) are not yet available for general use.
However, we were fortunate to have Dr. David Anderson, Office of Migratory Bird Management, U. S. Fish and Wildlife Service, run our recovery
data through these programs.
Thus, we now have survival rate estimates
by sex and management unit for mallards banded post-season in eastern
Colorado.
Trapping and banding methods and banding locations remained the same as
in last segment (Hopper 1974). Periodic aerial censuses were conducted
using a Cessna 185 aircraft and 1-2 observers.

RESULTS AND DISCUSSION

Trapping

and Banding

Banding quotas of 800-1,000 mallards were set for each waterfowl management
unit of northeastern Colorado (1, 2, 3, 4, 6 and 9) and for all units in
the Arkansas Valley combined (10-13)(Fig. 1). These quotas were met in all
cases, however, as in most years, age and sex ratios in the banded samples
were not always desirable (Table 1).
Trapping efforts resulted in the banding of 6,558 mallards, or about 1,000
more, than last year (Table 1). Most samples were easily obtained, despite
generally mild weather conditions.

Winter Aerial Surveys
Dates and results of periodic aerial counts are shown by management unit
in Table 2. The November and Decembe~ surveys were incomplete because of
the absence of data in some units. Thus, totals for all units combined are
rather meaningless except for the January inventory.
The January count of
1975 (281,972) was considerably above those of the past two years, but interpretation of results between years is difficult at best.

Analysis

of Band Recovery Data

This preliminary analysis of band recovery data for eastern Colorado wintering
mallards was based on bandings from the winter of 1963-64 through the winter
of 1971-72 and on recoveries from the 1964-65 through 1972-73 hunting seasons.
Table 3 presents a tabulation of numbers of birds banded during this period
by age, sex and management unit. Samples were quite small for units in the
Arkansas Valley (10-13). As a result, this analysis combined these units
and treated the Arkansas Valley as one unit in most cases.
Samples of
females were nearly always smaller than those for males, largely because:
(1) trapping techniques were mostly selective toward males, and (2) females
represented a smaller segment than males in the overall population of mallards
wintering in eastern Colorado.

�"

«
0:

z

�Table 1.
1974-75.

Numbers and percentages of mallards in the banded sample by location, age and sex, eastern Colorado,

Management Unit

Number
Banded

(1) Sterling-Julesburg

Adults
Males
Females
Number
Percent
Number
Percent

Immatures
Males
Females
Number
Percent
Number
Percent

986

292

29.6

193

19.6

286

29.0

215

21.8

(2) Ft. Morgan-Sterling

894

248

27.8

135

15.1

289

32.3

222

24.8

(3) Greeley-Ft. Morgan

848

249

29.4

155

18.2

288

34.0

156

18.4

825

245

29.7

254

30.8

178

21.6

148

17.9

1,022

251

24.6

247

24.2

266

26.0

258

25.2

(4) Ft. Collins
(6) Denver-Greeley
(9) Bonny Reservoir
(Ll) Two Buttes
Reservoir
(12) Rocky Ford-Lamar
(13) Pueblo-Rocky Ford
Totals

993

259

26.1

216

21.7

250

25.2

268

27.0

600

120

20.0'

80

13.3

172

28.7

228

38.0

90

50

55.6

12

13.3

17

18.9

11

12.2

300

141

47.0

57

19.0

50

16.7

52

17.3

6,558

1,855

28.3

1,349

20.6

1,796

27.4

1,558

23.7

I
•....•
00
0
I

�-181-

Table 2. Aerial duck counts by date interval and study unit, eastern
Colorado, 1974-75.

Management

November
4

Unit

Number of Ducks Counted
December
December
January
11
23
7-8

(1) Sterling-Julesburg

17,000

54,013

44,656

(2) Ft. Morgan-Sterling

7,800

37,665

13,823

(3) Greeley-Ft.

16,800

4,104

20,485

54,880

(4) Ft. Collins

29,425

28,491

46,965

69,208

(6) Denver-Greeley

6,800

15,076

14,745

39,545

Morgan

(9) Bonny Reservoir

35,000

(10-13) Arkansas Valley

9,950

(15) San Luis Valley

14,910

Totals

77,825

47,671

173,873

281,972

�-182-

Table 3. Winter bandings of eastern
Colorado mallards,
un L t , 1963-64 through 1971-72.
Number of Mallards

by age, sex and

Adult

Banded
Females
Immature

Total

Grand
Total

4,066

970

1,542

2,512

6,578

1,881

4,502

1,367

1,300

2,667

7,169

2,274

1,857

4,131

1,364

1,733

3,097

7,228

7,024

5,675

12,699

3,701

4,575

8,276

20,975

2,012

2,000

4,012

1,236

1,778

3,014

7,026

2,449

2,093

4,542

1,457

1,744

3,201

7,743

2,714

1,898

4,612

1,176

1,545

2,721

7,333

7,175

5,991

13,166

3,869

5,067

8,936

22,102

398

444

842

192

379

571

1,413

188

151

339

375

330

705

1,044

1,441

905

2,346

756

699

1,455

3,801

782

399

1,181

380

347

727

1,908

Totals
(10-13)

2,809

1,899

4,708

1,685

1,755

3,458

8,166

Grand Total

17,008

13,565

30,573

9,255

11 ,397

20,670

51,243

Unit of
Banding

1
2
9
Totals
(1-2-9)

3
4
6
Totals
(3-4-6)

10
11
12
13

Adult

Males
Immature

Total

2,129

1,937

2,621

Over 51,000 mallards were banded during the above 9-year period. Males accotnlted
for 30,573 of this total. including 17,008 adults and 13,565 immatures.
The
remaining 20,670 birds were composed of 9,255 adult females and 11,397 immature
females.
The number of usable recoveries that resulted from these bandings are listed in
Tables 4 (first-year) and 5 (all years) for hunting seasons 1964-65 through 197273. First-year recoveries totaled 1,461 for the four age and sex classes combined, while nearly 4,400 recoveries were obtained for all years.

�-183-

Table 4. Number of recoveries resulting from eastern Colorado winter
mallard bandings by age, sex, and unit, 1964-65 through 1972-73 hunting
seasons (first year recoveries).
Number of Band Recoveries
Females
Total
Adult
Immature

Unit of
Banding

Adult

Males
Immature

1

74

85

159

19

2

84

68

152

9

42

60

Totals
(1-2-9)

200

3

Total

Grand
Total

34

53

212

26

23

49

201

102

23

33

56

158

213

413

68

90

158

571

85

68

153

26

41

67

220

4

76

78

154

38

45

83

237

6

105

91

196

17

37

54

250

Totals
(3-4-6)

266

237

503

81

123

204

707

10

7

7

14

1

2

3

17

11

3

4

7

4

9

13

20

12

38

23

61

9

17

26

87

13

29

17

46

5

8

13

59

Totals
(10-13)

77

51

128

19

36

55

183

Grand Total

543

501

1,044

168

249

417

1,461

�-184-

Table 5. Number of recoveries resulting from eastern Colorado winter
mallard bandings by age, sex and unit, 1964-65 through 1972-73 hunting
seasons (all years).
Number of Band Recoveries
Females
Total
Adult
Immature

Unit of
Banding

Adult

Males
Immature

1

263

233

496

31

316

226

542

174

167

753

Total

Grand
Total

86

117

613

59

61

120

662

341

60

77

137

478

626

1,379

150

224

374

1,753

248

233

481

65

98

163

644

281

256

537

80

86

166

739

374

251

625

46

80

126

751

Totals
(3-4-6)

903

740

1,643

191

264

455

2,098

10

31

24

55

5

6

11

66

11

4

5

9

5

13

18

27

133

87

220

30

35

65

285

13

77

54

131

11

16

27

158

Totals
(10-13)

245

170

415

51

70

121

536

1,901

1,536

3,437

392

558

950

4,387

2
9
Totals
(1-2-9)

3
4
6

12

Grand
Total

�-185-

A complete analysis of banding data resulting from the above bandings
and recoveries was not finished during Segment 20. Thus, this report
presents only some general data and does not consider the detailed information by management unit that will be a part of the final analysis.

Geographic

Distribution

of Recoveries

Tables 6 and 7 show the distribution of band recoveries from mallards
banded post-season in eastern Colorado for the first year after banding
and for all years, respectively.
The distribution of first-year recoveries
did not differ greatly from that of all years for each age and sex class.
One possible exception may be that about 43 percent of the first-year
recoveries of immature males occurred in the High Plains portion of Colorado,
whereas 52 percent of the all-year recoveries occurred here. The following
other conclusions seem 3pparent upon reviewing Tables 6-7.
1.

Males were less likely to be recovered

in Canada than females.

2.

Males were more likely to be recovered in eastern Colorado and
the High Plains Region as a whole than the corresponding age
class of females.

3.

Adult males were more likely to be recovered in eastern Colorado
and the High Plains Region as a whole than any of the other age
and sex classes.

4.

Immatures of both sexes were more likely than adults to be recovered in areas outside the Central Flyway, at least for firstyear recoveries.
Recoveries from all years showed the same
tendency, but perhaps mainly because of the influence of firstyear recoveries on the total.,

5.

Eastern Colorado was the major recovery
age and sex classes.

6.

Alberta was the most important province of recovery
for all four age and sex classes.

location for all four

in Canada

Recovery and Survival Rate Estimates
Recent developments in methods for estimating recovery and survival rates
of animal populations provide for the calculation of confidence limits to
apply to these rates. This is a definite advantage in estimating population
parameters, since for the first time we are able to see the accuracy of our
estimates.
Two such methods, one described by Seber (1970) and Robson and
Youngs (1971) and the other by Brownie and Robson (1974), are now commonly
in use. The former method requires that recovery and survival rates to be
independent of age, whereas the latter assumes age-dependent recovery and
survival rates.

�-186-

Table 6. Distribution of band recoveries from mallards banded postseason in eastern Co.l.o rado, 1963-64 through 1970-71 (first year recoveries).

AM

Percent of Total Recoveries
Age at Banding
1M
AF

IF

Alberta

9.4

12.6

15.4

16.5

Saskatchewan

5.5

4.8

6.0

0.8

Manitoba

0.7

0.6

0.0

6.8

Other Canada

0.2

0.2

0.0

1.2

15.8

18.2

21.4

25.3

Washington

0.2

1.4

0.0

1.2

Oregon

0.0

0.8

0.0

0.8

California

0.2

0.4

0.0

0.4

Idaho

0.6

1.2

1.8

2.8

Nevada

0.0

0.0

0.0

0.0

Utah

0.2

0.8

0.0

1.2

Arizona

0.2

0.4

0.0

0.0

W. Montana

0.7

2.0

0.6

2.8

W. Wyoming

0.2

0.2

0.0

0.8

W. Colorado

0.0

0.0

1.2

0.8

W. New Mexico

0.0

.0.0

0.0

0.0

2.2

7.2

3.6

10.8

E. Montana

1.3

0.4

1.2

1.2

E. Wyoming

1.8

3.2

0.6

1.6

E. Colorado

65.7

42.9

50.0

31.3

E. New Mexico

0.2

0.2

0.6

0.0

W. North Dakota

0.4

1.2

1.8

2.4

W. South Dakota

0.6

3.2

0.0

1.2

W. Nebraska

4.8

9.8

6.5

7.6

W. Kansas

0.4

1.8

1.2

1.6

W. Oklahoma

0.0

0.0

0.6

0.0

W. Texas

O.7

0.4

0.6

0.8

75.9

63.1

63.1

47.8

Recovery
Location

Total Canada

Total Pacific Flyway

Tota1 High Plains

�-187-

Table 6. Distribution of band recoveries from mallards banded postseason in eastern Colorado. 1963-64 through 1970-71 (first year recoveries)
(continued).
Percent of Total Recoveries
Age at Banding
1M
AF

Recovery
Location

AM

E. North Dakota

1.8

1.4

0.6

0.8

E. South Dakota

0.6

2.0

2.4

2.4

E. Nebraska

1.1

2.6

1.8

2.4

E. Kansas

0.9

2.2

1.2

3.6

E. Oklahoma

0.4

0.0

1.2

0.8

E. Texas
Total E. Central
Flyway

0.0

1.2

2.4

2.0

4.8

9.4

9.5

12.0

Minnesota

0.2

0.2

0.0

0.0

Wisconsin

0.0

0.0

0.0

0.0

Michigan

0.0

0.0

0.0

0.0
0.0

IF

Iowa

0.2

0.6

1.2

Illinois

0.2

0.2

0.0

0.4

Indiana

0.0

0.0

0.0

0.0

Ohio

0.0

0.0

0.0

0.0

Missouri

0.2

0.0

0.0

0.8

Kentucky

0.0

0.0

0.0

0.0

Tennessee

0.0

0.0

0.0

0.0

Arkansas

0.4

1.0

0.6

0.4

Louisiana

0.2

0.4

0.6

2.0

Mississippi

0.0

0.2

0.0

0.4

Alabama

0.0

0.0

0.0

0.0

1.3

2.6

2.4

4.0

Atlantic Flyway

0.0

0.0

0.0

0.0

Alaska

0.0

0.0

0.0

0.0

Mexico

0.0

0.0

0.0

0.0

Total Mississippi
Flyway

�-188-

Table 7. Distribution of band recoveries from mallards banded post-season
in eastern Colorado, 1963-64 through 1970-71 (recoveries from all years).
Percent of Total Recoveries
Age at Banding

Recovery
Location

AM

1M

AF

IF

Alberta

7.7

8.8

15.3

16.8

Saskatchewan

5.7

6.1

6.4

7.5

Manitoba

0.4

0.45

0.2

0.5

Other Canada

0.1

0.1

0.0

0.5

13.9

15.6

21.9

25.4

Washington

0.2

1.0

0.2

1.1

Oregon

0.05

0.5

0.0

0.7

California

0.15

0.3

0.0

0.5

Idaho

0.7

1.7

1.0

2.3

Nevada

0.05

0.0

0.2

0.0

Utah

0.2

0.5

0.2

0.7

Arizona

0.1

0.2

0.0

0.0

W. Montana

0.7

1.1

0.5

1.6

W. Wyoming

0.15

0.1

0.0

0.4

W. Colorado

0.2

0.2

0.8

0.5

W. New Mexico

0.0

0.0

0.0

0.0

2.5

5.5

3.1

7.9

E. Montana

0.8

0.7

1.0

1.1

E. Wyoming

1.5

2.0

0.8

2.0

E. Colorado

64.6

52.1

50.8

35.1

E. New Mexico

0.2

0.2

0.5

0.0

W. North Dakota

0.6

0.45

1.3

2.0

W. South Dakota

0.5

1.6

0.8

1.4

W. Nebraska

8.4

10.5

8.4

8.1

W. Kansas

0.8

0.9

1.0

1.1

W. Oklahoma

0.0

0.1

0.2

0.0

W. Texas

0.7

0.6

1.0

0.9

78.2

69.1

66.1

51.6

Total Canada

Total Pacific Flyway

Total High Plains

�-189-

Table 7. Distribution of band recoveries from mallards banded post-season
in eastern Colorado, 1963-64 through 1970-71 (recoveries from all years)
(continued) •
Percent of Total Recoveries
Age at Banding
1M
AF

Recovery
Location

AM

E. North Dakota

1.5

1.2

1.0

2.0

E. South Dakota

0.5

1.4

1.0

1.4

E. Nebraska

1.2

2.0

1.8

1.8

E. Kansas

0.6

1.4

0.8

2.2

E. Oklahoma

0.15

0.3

1.3

0.9

E. Texas

0.3

1.0

1.8

1.6

4.3

7.2

7.6

9.8

Minnesota

0.1

0.3

0.5

0.4

Wisconsin

0.0

0.0

0.0

0.0

Michigan

0.0

0.0

0.0

0.0

Iowa

0.05

0.5

0.5

0.2

Illinois

0.15

0.2

0.0

0.4

Indiana

0.0

0.0

0.0

0.0

Ohio

0.0

0.0

0.0

0.0

Missouri

0.1

0.2

0.0

0.7

Kentucky

0.0

0.0

0.0

0.0

Tennessee

0.0

0.1

0.0

Arkansas

0.4

0.6

0.2

0.0
1.8

Louisiana

0.3

0.6

0.2

1.2

Mississippi

0.05

0.1

0.0

0.2

Alabama

0.0

0.1

0.0

0.0

1.1

2.7

1.5

4.8

Atlantic Flyway

0.0

0.0

0.0

1.2

Alaska

0.0

0.0

0.0

1.2

Mexico

0.0

0.0

0.0

0.0

Total E. Central
Flyway

Total Mississippi
Flyway

IF

�-190-

The first step in the analyses of post-season banding of eastern Colorado
mallards was to select the proper analysis method of the two noted previously. This allowed the determination as to whether adult and immature
birds should be pooled or considered separately in the analysis. The two
age groups were treated separately by sex under the method outlined by
Brownie and Robson (1974), and a contingency chi-square test was applied
to the resulting data to test the null hypothesis that no difference
occurs in recovery and survival rates between adults and immatures (Table
8). The null hypothesis was accepted for both sexes, with chi-square
values of 15.26 (P&gt;0.50) and 19.68 (P&gt;0.25), respectively.
Thus, there
was no evidence that adult and immature mallards banded post-season in
eastern Colorado were different in regard to recovery and survival rates.
This conclusion prompte~ the decision to combine the two age classes and
use a method that assumes recovery and survival rates are not age-dependent,
i.e. that of, Seber (1970) and Robson and Youngs (1971).

Table 8. Comparison of adult and immature recovery and survival rate
estimates of mallards banded post-season in eastern Colorado, 1963-64
through 1970-71.
Sex
and
Age

Mean
Recovery Rate !/
(%)

Mean
Survival Rate !/
(%)
d.f.

3.21 ± 0.1

75.41 ± 1.70

2

X

P

17

15.26

&gt; 0.50

17

19.68

&gt; 0.25

Males
Adult

Immature

3.24 ± 0.2

74.05 ± 3.54

1.65 ± 0.1

69.37 ± 3.29

Females
Adult

Irrnnature

1.81 ± 0.1

68.88 ± 6.30

!/
± t.05 standard error.

The next procedure was to test for differences due to sex for both ages combined. This involved the analysis of the null hypothesis that recovery and
survival rates of males equals recovery and survival rates of females. The
resulting contingency chi-square test produced a highly significant chi-square

�-191value of 561.6 with 17 degrees of freedom. Thus, it was very clear that
males and females have different population parameters and that further
analysis should be done separately by sex.
Table 9 presents information on estimates of population parameters by sex
(ages combined) for mallards banded post-season in eastern Colorado, 196364 through 1971-72, according to two models under the method of banding
analysis by Seber (1970) and Robson and Youngs (1971). Modell assumes
time-specific recovery and survival rates, while Model 2 assumes recovery
and survival rates are constant over time. Both models seem to fit the
data in question, with each having certain advantages. Modell provides
an estimate of mean life span in addition to mean recovery and survival
rate estimates. Model 2 lacks the mean life span estimate, but estimates
of mean recovery and survival rates are often better than under Model I,
in that standard deviations are usually smaller and confidence intervals
narrower.
The mean recovery rates for females for Models 1 and 2, respectively,
(1.71 and 1.75%) were considerably less than those for males (3.06 and
3.23%) (Table 9). Survival was quite high for both males and females
during the 1964-72 period, with mean rates of about 75 and 66 percent,
respectively.
Their mean life spans as adults were estimated at 3.43 years
for males and 2.28 years for females. Note the relatively large difference
in mean life span between the sexes as compared to the difference between
survival rates. This exemplifies the tendency for a relatively small change
in survival rate to produce a sizable change in mean life span.
A more detailed analysis of these data will be conducted next segment, including comparisons of management units. Geographic distributions of recoveries by 10-day intervals of the hunting season will also be prepared
and compared by age, sex and management unit.

LITERATURE CITED

Brownie, C., and D. S. Robson. 1974. Models allOWing for age-dependent
survival rates for band-return data. Cornell Univ. Biometrics Unit
Mimeograph Series, Paper No. BU-5l4-M, 34pp.
Hopper, R. M. 1973. Investigation of mallard management units of eastern
Colorado. Colo. Div. of Wildl., Game Res. Rept. Oct. p. 83-98.
1974. Investigation of mallard management units of eastern Colorado.
Colo. Div. of Wildl., Game Res. Rept. Oct. p. 137-144.
Robson, D. S., and W. D. Youngs. 1971. Statistical analysis of reported
tag-recapture in the harvest from an exploited population.
Cornell
Univ. Biometrics Unit Mimeograph Series, Paper No. BU-369-M.

�-192-

Seber, G. A. F. 1970. Estimating time-specific survival and reporting
rates for adult birds from band returns. Biometrika 57(2):313-318.

",..,..:~~/10/~,-'

Prepared by --.Jqflh-· '.t.~t:;;4i~::M::'B.'c
~chard
M. Hopper~
Wildlife Researcher

.£..~~~

-~4-

�-193Table 9. Population parameters 1/ of male and female mallards banded postseason in eastern Colorado, 1963-64 through 1971-72.

Estimate

Standard
Deviation

90 Percent
Confidence
Interval

Mean recovery rate (%)

3.06

0.08

2.92-3.20

Mean survival rate (%)

75.43

1.23

73.41-77.45

Mean life span (years)

3.43

0.19

3.11-3.75

Mean recovery rate (%)

1.71

0.09

1.57-1.86

Mean survival rate (%)

65.58

2.27

61. 86-69.30

Mean life span (years)

2.28

0.18

1. 98-2.58

Mean recovery rate (%)

3.23

0.09

3.09-3.37

Mean survival rate (%)

74.91

0.98

73.30-76.52

Mean recovery rate (%)

1. 75.

0.09

1.61-1.89

Mean survival rate (%)

65.57

1. 78

62.66-68.49

Population
Parameter
2/
Model 1Males

Females

Model 2 ]./
Males

Females

1/ Estimates

after analysis method of Seber (1970) and Robson and Youngs

l/ Assume time-specific

recovery and survival rates.

1/ Assumes constant recovery and survival rates over time.

(1971).

��-195-

October 1975

JOB PROGRESS REPORT

State

....:C:..:O:.:L:.:O:.:Ri:.::,:.::ill=-O=-------

0f

Proj ect No.

W-88-R-20

Work Plan No.

4

Migratory

Bird Investigations

Job No.

Job Title

Trapping and Banding Doves

Period Covered:

April 1, 1974 through March 31, 1975

Personnel:

3

_

W. John Arthur, Clait E. Braun, Dwayne Finch, Howard D. Funk,
Richard W. Hoffman, J. Edward Kautz, Charles W. Loeffler, Ronald
Oakleaf, Willie Travnicek, and Kenneth C. Wagner, Colorado Division
of Wildlife.

ABSTRl\CT
Efforts initiated in 1964 to trap and band samples of mourning doves (Zenaida
macroura) in three geographic areas of Colorado were continued and terminated
in 1974. Efforts of State personnel resulted in 4,499 birds being newly
banded. Of this total, 2,021 were immatures, 1,415 were adult males, 1,057
were adult females, while age and/or sex were not determined for 6 birds.
Assigned quotas were accomplished for all areas except extreme eastern Colorado.
Wing molt data from trapped and harvested immature doves indicated that nesting
started earlier in 1974 than in 1973 and continued later. Hatching peaks
occurred in mid May, mid June, mid July 'and mid August. A sample of 1,165
hunter harvested birds, all from east of the Continental Divide, gave a young-toold ratio of 2.2:1. Wing molt data from harvested immatures in this sample indicated that few (less than 12 percent of those shot hatched prior to 1 July)
early hatched young were available to hunters in Colorado. Thirty-one band
recoveries from doves banued by Division personnel were reported to date in the
1974 recovery year. Fourteen (45.2 percent) had been banded in 1974. Of the
recoveries, 18 (58.1 percent) were recovered in Colorado, while 7 (22.6 percent)
were from Texas, with the remainder being from New Mexico (3), Arizona (2) and
Mexico (1). Twenty-seven (87.1 percent) of the recoveries were by hunters.
Apparent hunting mortality for doves banded in Colorado continues to be less
than 3'percent.

�-196-

RECOMMENDATIONS
1.

It is recommended that a final report exclusive of detailed recovery
and mortality analyses be prepared in 1975-76.

2.

It is recommended that this project be held inactive until 1977 or 1978
when final analyses of all bandings and recoveries will be conducted in
conjunction with the analyses of all Central Management Unit bandings
and recoveries.

�-197-

TRAPPING AND BANDING DOVES
C1ait E. Braun

Intensive efforts to trap and band mourning doves in Colorado as part of
a Central Management Unit banding program were continued in 1974. Data
presented in this report are those collected in 1974, the eleventh and final
banding year for this investigation.

P. S. OBJECTIVE
To investigate migration patterns and mortality
banded in Colorado by age, sex, and area.

rates of mourning

doves

SEGMENT OBJECTIVES
1.

To trap and band mourning doves in three selected areas of Colorado for
the purpose of obtaining migration, life history, and annual mortality
data.

2.

To estimate harvest size and hunter success.

METHODS AND MATERIALS
Methods and materials used in 1974 were similar to those described
reports and summarized by Braun (1970) .•

Description

of Trapping

in earlier

Sites

Trapping sites throughout the State were essentially
vious years, described by Braun (1970, 1974).

those utilized

in pre-

RESULTS AND DISCUSSION

Trapping

and Banding

Trapping and banding efforts in 1974 resulted in 4,499 doves being newly
banded. Doves were banded at five sites on the eastern slope, two sites on
the eastern plains and at two sites west of the Continental Divide.
The
banding goal of 4,000 birds was essentially accomplished as 1,200 were banded
in western Colorado (quota = 1,000), 900 were banded in extreme eastern
Colorado (quota = 1,000), and 2,399 were banded on the east slope of the
Continental Divide (quota = 2,000). Sex and age distribution of the banded
sample by area are presented in Table 1.

�-198-

Table 1. Sex and age distribution
Colorado, 1974.

Area

Adult Male
No.
Banded Percent

of mourning

doves banded by area in

Adult Female
No.
Banded
Percent

Immature
No.
Banded
Percent

Total

148

24.7

316

52.6

600

67

29.7

44

7.3

300

215

23.9

360

40.0

900

162

27.0

263

43.8

600

35

5.8

466

77.7

600

35

6.5

442

81.8

540

36.6

130

26.0

187

37.4

500

Eastern Colorado
Ft. Morgan
Vineland
Subtotal

136
189
325

22.7
63.0
36.1

Eastern Slope
Monte Vista
Ft. Garland
Denver

175

29.2

99

16.5

63

11. 7

Ft. Collins Area

183

Salida

73

47.7

54

35.3

26

17.0

159 ~/

593

24.8

416

17.4

1384

57.8

2399

43.8

247

41.2

90

15.0

600

39.0

179

29.8

187

Subtotal

---

Western Colorado
Durango
Montrose
Subtotal

TOTAL ALL AREAS

263
234
497

1415

41.4

31.5

426

35.5

-277

1057

23.5

2021

31.2

600

23.1

1200

45.0

4499

1/
Including 1 unknown age and sex and 5 unknown sex adults.

Numbers and percentages of adult males and females, and immatures banded in 1974
were similar to those banded in 1973 and in all years since 1969 when sample sizes
were enlarged through intensified trapping efforts.
Percentages of immatures in
trap samples were highest at Denver and Ft. Garland, sites that were trapped in
late July and August.
Sites trapped in early June (Vineland, Durango and Salida)

�-199-

had the lowest percentages of immatures in the banded sample. Thus, percent immatures banded was a function of time of banding. As in other years,
more adult males than adult females were banded (31.5 to 23.5 percent).
This disparity is probably related to timing of incubation and brood sharing,
as females would normally be exposed to only one trapping period per day.
During the 1974 trapping period, 58 mourning doves were recaptured that had
been banded in previous years. Of this sample, 55 returned to the area where
they were initially banded. One bird banded at Denver was recaptured at
Vineland while two doves banded outside of Colorado were recaptured.
One of
these recaptures (Montrose) had been initially banded at Eufaula, Alabama
in 1972 while the other (Salida) was originally banded in 1973 near Roswell,
New Mexico. Forty-three (74.l percent) of the 58 recaptures had been banded
in 1973, while 10 (17.2 percent) were banded in 1972, 4 in 1971 and 1 in 1969.

Hatching Data
Wing molt data were available for 2,012 of the 2,021 immature doves trapped
and banded in 1974 and from 797 hunter harvested immatures. Estimated hatching
dates (Allen 1963) are presented in Tables 2 and 3. Examination of data in
Table 2 indicates that few mourning doves trapped in Colorado hatched prior
to 1 May, with most trapped immatures hatching from mid May to late June.
While definite hatching peaks are difficult to ascertain from data in Table 2,
peaks are suggested in mid to late May and in mid June. Since trapping
ceases in early August, progeny of late nests are not represented in Table 2.
This can clearly be seen upon examining data in Table 3 as two peaks can be
observed (early July and early August). Data in Table 3 suggest that early
hatched doves have migrated from Colorado prior to 1 September.
It is
apparent that neither trapping or hunting data alone accurately represent
actual hatching dates as one method (trapping) fails to show late nesting,
while the other (hunter harvest in Colorado) misrepresents production early
in the nesting period. It should be remembered that neither trap or hunter
samples may actually represent doves produced in Colorado.

Harvest
Efforts were again made to collect wings from hunter harvested doves throughout Colorado during the 1974 season. Through this effort, 1,165 were classified as to age (adult or immature) and primary molt. All wings were from
east slope areas with 7 from North Park, 52 from near Vilas, and 1,106 from
Ft. Carson. All wings were from September with over one-third (38.5 percent)
being from after mid September. The overall immature to adult ratio was
2.2:1 but varied greatly by area and time. At Ft. Carson, where adequate
samples were obtained, immature to adult ratios were lowest early in September
(1.4:1 during 1 to 5 September and 2.3:1 during 6 to 12 September) and highest
later in the month (4.7:1 during 13 to 20 September and 3.2:1 during 21 to
30 September).
Sample sizes, while uneven (322, 109, 232 and 217 respectively)
were adequate in each time period.
Harvest data obtained through use of a mail survey of hunters after the 1974
hunting season are not presently available.
Final results of this survey are
normally not available for distribution until 30 June of the year following
the season (H. Riffel, personal communication).
Results of the 1973 survey
of three percent of the small game hunter license holders {about 80 percent

�Table 2.

Estimated

Table 3.

doves, 1974.

Prior
to
May 1

May
1-7

May
8-14

May
15-21

May
22-28

May 29June 4

June
5-11

June
12-18

June
19-25

June 26July 2

July
3-9

July
10-16

July
17-23

12

31

101

281

344

272

229

236

205

144

139

18

0

1.5

5.0

14.0

17.1

13.5

11.4

11. 7

10.2

7.2

6.9

0.9

0.0

Number
Hatching
Percent
of Total

hatching dates for wild trapped immature mourning

0.6

Estimated hatching

dates for hunter harvested

immature mourning

doves, 1974.
I

tv

Number
Hatching

0
0
I

Prior
to
June 12

June
12-18

June
19-25

June 26July 2

July
3-9

July
10-16

July
17-23

July
24-30

July 31August 6

4

28

41

23

89

70

75

95

131

71

3.5

5.2

3.0

11.2

8.9

9.4

11.9

16.5

8.9

Percent of
Total
0.1

August· August
7-13
14-20

August
21-27

August 28Sept. 3

84

42

44

10.6

5.3

5.5

�-201-

response) indicate that an estimated
484,660 mourning doves.

31,688 hunters harvested

an estimated

To date (30 April), recoveries of 31 mourning doves banded by Division
personnel have been reported in the 1974 recovery year (1 June 1974 to 31
May 1975). Of the 31 recoveries, one each was banded in 1969 and 1970,
five in 1972, 10 in 1973 while 14 were first year recoveries (banded in
1974). All but 4 were shot recoveries.
Three of the 31 recoveries were
banded west of the Continental Divide, with two being recovered in Arizona
and one in west Texas. While data are limited, recoveries of west slope
banded doves are uncommon east of the Continental Divide.
Considering all
recoveries, 18 (58.1 percent) were from Colorado, 7 (22.6 percent) were
from Texas, with the remainder being from New Mexico (3), Arizona (2), and
Mexico (1). Examining first year recoveries only, 7 of 14 were banded as
immatures (50.0 percent).
This is slightly higher than the percent immatures
in the 1974 trap sample (45.0) suggesting that vulnerability differences
between adults and immatures are not great.
Ten doves banded in other states were reported recovered in Colorado
Five were banded in Montana, three in New Mexico and two in Wyoming.
of the recoveries (all banded as immatures) were from 1974 bandings.

LITERATURE

in 1974.
Six

CITED

Allen, J. M. 1963. Primary feather molt rate of wild immature doves in
Indiana.
Ind. Dept. Conserv., Game Res. Sect. Circ. No.4.
Indianapolis.
4p.
Braun, C. E. 1970. Mourning dove trapping and banding.
Colo. Div. Game,
Fish and Parks, Game Res. Rept., Fed. Aid Proj. W-88-R.
Oct. p. 143149.
1974. Mourning dove trapping and banding.
Colo. Div. Wildl., Game
Res. Rept., Fed. Aid Proj. W-88-R. Oct. p. 145-152.

Prepared by

_~tfa.:::.::..:id~_·.=.T.;...;''-lL'~~~
Clait E. Braun
Wildlife Researcher

_

��October 1975
-203-

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No.

W-88-R-20

Work Plan No.

4

Migratory Bird Investigations
Job No.

Job Title

Band-tailed Pigeon Investigations

Period Covered:

April 1, 1974 through March 31, 1975

4

_

Personnel: Rick Adams, W. John Arthur, Clait E. Braun, Ron Desilet, Richard
Fentzlaff, Roward Funk, Bill Reicher, Tom Henry, J. Edward Kautz, Harold
Lanning, Ann Leckler, Don Masden, Sam Matthews, Ron Oakleaf, Charles
Reichert, Robert Rouse, Warren Rupke, Glen Smith, Ron Velarde, Charles
Wagner, Ken Wagner, Pat Waters, Al Whitaker, Mike Zgainer; Wildlife Conservation
Officers and other personnel of the Colorado Division of Wildlife.
ABSTRACT

Investigations of band-tailed pigeons (Columba fasciata) initiated in Colorado
in conjunction with the Four Corner States Cooperative Study were continued
in 1974. Major emphasis was placed on banding and continuation of analyses
of hunting statistics. Pigeons were trapped at 23 different locations in 1974
with 3,960 birds (2,468 adults, 580 subadults, and 912 immatures) being newly
banded. Over one thousand (1,229) pigeons were retrapped, including 13 banded
outside of Colorado (12 from New Mexico and 1 from Utah). During the hunting
season, 518 permits were issued, with 288 individuals actually hunting. Only
132 hunters were successful, each harvesting an average of 5.6 pigeons. Total
estimated harvest (including crippling loss) projected from the questionnaire
survey (89.8 percent response) was 792 birds, more than twice the number
harvested in 1973. Crippling loss approximated 8 percent of the birds shot and
retrieved. Wings were received from 33 successful hunters. Immature and subadult pigeons comprised 42.1 percent of the 171 bandtail wings received.
Field checks of 153 (101 adults, 52 immatures) hunter-killed birds indicated
that 52.5 percent of the adults were still involved with nest activities and/or
feeding of young. Recoveries were received from 137 pigeons banded in Colorado.
Most (94) were recovered in Colorado, with 39 being recovered in New Mexico,
2 in Mexico and 1 each in California and Utah. At least 11.2 percent of the
birds harvested during the hunting season in Colorado were banded. Hatching
dates were calculated for 998 pigeons with most hatching between 12 May and 10
July. Helminth infections continue to be low (15.8 percent).
The final report of the Four Corner States Cooperative Band-tailed Pigeon
Investigation was compiled, reviewed and submitted for editing and publication.
Two papers on parasites (one accepted to date) and one paper on reproductive
biology (accepted) were submitted ~to technical journals.

�-204-

RECOMMENDATIONS

1.

This project should be extended through 31 March 1976 for writing of
technical articles.

2.

Technical papers should be prepared and submitted on the following subjects
concerning band-tailed pigeons: mercury residues, parasites (2), weights,
molt (2), methods of trapping and location, time of day of capture, aviary
nesting, estimation of population size, mortality and survival, and general
ecology.

�-205-

BAND-TAILED PIGEON INVESTIGATIONS
Clait E. Braun

Intensive ecological studies of wild band-tailed pigeons which nest and reside
in Colorado from late April to early November, initiated in 1969, continued in
1974. Investigations in Colorado represent a portion of the regionwide Four
Corner Cooperative Band-tailed Pigeon Investigation which was initiated in
Arizona in 1967. Data presented in this report are those collected in 1974,
the last year of field work for this investigation.

P. S. OBJECTIVE
To investigate life history and ecological facts pertinent to band-tailed
pigeons in Colorado as follows: range; relative numbers; relation between
numbers of birds present by area and such physical factors as weather, food,
vegetative cover, elevation, and salt; breeding and nesting chronology;
migration patterns and chronology; harvest patterns; and mortality rates.

SEGMENT OBJECTIVES
1.

To trap, band, age, and sex as many birds as possible in selected areas
of Colorado to investigate migration patterns and chronology of Colorado
bandtails throughout their range.

2.

To conduct a hunting season, gather-harvest

3.

Compile and initiate analyses of all data collected in Colorado.
ate in preparing fin~l report for Four Corner States.

data, and evaluate results.
Cooper-

METHODS AND MATERIALS
Methods and materials used in 1974 were identical to those in previous years
of the study (Braun 1970, 1971, 1972, 1973, 1974). Written reports of pigeons
were not solicited from field personnel although communications were encouraged for ease in locating groups of pigeons in trappab1e locations. Trapping
methodology in 1974 was that used in previous years, with most birds being
captured through use of cannon nets. Evaluation of the hunting season and
disposition of collected birds were the same as in previous years.

Description

of Trapping Areas

In 1974, pigeons were trapped at 23 different locations of which only three
(Trinidad, Collbran and Sedalia) had not been trapped in previous years. All

�-206-

other sites (Table 1) have been previously described (Braun 1970, 1971, 1972,
1973, 1974). The three sites newly trapped in 1974 are similar to other sites
trapped in most years as one was a feedlot (Trinidad), one was a grain field
(Collbran) and one was in a. residential area (Sedalia). As in previous years,
all trap sites were immediately adjacent to readily accessible trees or power
lines used for perches.
Trees used were primarily ponderosa pine (Pinus
ponderosa), Gambel's oak (Quercus gambelii), pinon pine {Pinus edu~
cottonwoods (Populus spp.), and Engelmann spruce (Picea engelmannii).
It was difficult to ascertain feeding preferences, as about equal numbers of pigeons were
trapped in fields where they were feeding on waste barley (Hordeum vulgare),
and wheat (Triticum aestivum).
Whole or cracked corn (Zea mays) was frequently
used as an attractant and bandtails clearly indicated a preference for this
grain.

RESULTS AND DISCUSSION

Banding
Trapping and banding activities remained at the same level in 1974 as in 1973
although more pigeons were banded (3,960 versus 3,559). As in 1973, emphasis
was placed on banding samples of pigeons (usually less than 200 birds per site)
in as many locations as feasible. As a result, more than 200 birds were banded
at only 8 of the 23 trapping locations.
Reasons for more than 200 new bandings
per site were related to intensive studies of color marked pigeons (two sites),
efforts to mark large numbers of immatures (five sites), and the initial trapping
of pigeons in a new flock area (one site). Efforts were continued to maintain
banding at as many of the sites used in previous years as feasible.
Thus,
trapping was successful at 3 of the 5 sites used in 1969, 9 of the 13 sites
used in 1970, 13 of the 15 sites trapped in 1971, 14 of the 23 locations trapped
in 1972, and 18 of the 25 sites trapped in 1973. MOre importantly, only two
(Minturn, Rye-Beulah-Wetmore)
of the major flock areas in Colorado were not
trapped in 1974. In all, 3,960 pigeons were newly banded (Table 1). Of this
number, 912 (23.0 percent) were immatures, 580 (14.6 percent) were subadults
and 2,468 were adults. Percent immatures in trap samples in 1974 was the highest
since 1969 when immatures comprised 23.6 percent of 1,600 pigeons banded. Upon
examination of data in Table 1 it is apparent that percent immatures in trap
samples increased throughout the summer. The increase in immatures can clearly
be observed in data at sites where pigeons were periodically trapped during the
summer (Woodland Park, Chipeta Park, Del Norte, etc.). The initial immature
trapped was on 25 June (Del Norte) about 10 days earlier than in 1973. This
suggests that nesting was earlier in 1974 than in 1973, a hypothesis supported
by field observational data of breeding activities.
Sex of adult, subadult and some immature pigeons was determined through examination of external plumage characters.
Of the 2,468 adults, 1,417 were males
(57.4 percent) and 1,051 were females (42.6 percent), while 232 of the 580
subadults were males (40.0 percent), with the remainder (348) classified as
females (60.0 percent).
Of the 912 immatures banded, sex by plumage examination was ascertained for 460. Of this sample, 239 (52.0 percent) were males while
221 (48.0 percent) were females. The disparity in sex ratios of adults is
probably related to time of day trapping was conducted.
The variance in percent
of subadult males and females trapped compared to percent adult males and females

�Table 1. Number of band-tailed

Area

Date(s)

pigeons banded by area in Colorado,

AdultJ.1
Percent
No. Banded

Immatures
Percent
No. Banded

Totals

Estimated No. of
Birds Using
Trapping Sites

11

100.0
90.0
57.1
42.3

0
2
6
15

0.0
10.0
42.9
57.7

48
20
14
26

200+
50
30
70

May 23
May 30

75
14

100.0
100.0

o
o

0.0
0.0

75
14

200+
25+

May 24
July 11
August 9

60
64
41

100.0
64.6
41.8

o
35
57

0.0
35.4
58.2

60
99
98

150+
175+
200+

May 13
June 29
July 30
August 15

48
18
8

Gypsum

Chipita Park

Woodland Park

1974.

-------------------------~----------------------------------------------------------------------1

Del Norte

May 25
May 28
May 29
June 25
July 30
August 23

107
54
31
78
120
21

100.0
100.0
100.0
98.7
78.9
23.6

0
0
0
1
32
68

Uniweep

May 28 &amp; 29
June 17
July 22

15
15

100.0
78.9

4

Basalt

May 29 &amp; 30

124

100.0

Longmont

June 4
July 29

101
16

Salida

June 4
July 13

73
81

0.0
0.0
0.0
1.3
21.1
76.4

107
54
31
79
152
89,

350+
150+
65+
200+
350+
200+

0.0
21.1

15
19

100+
25+

o

0.0

124

200+

100.0
100.0

o
o

0.0
0.0

101
16

300+
25+

100.0
79.4

o
21

0.0
20.6

73
102

150+
150+

------------------------------------------------------------------------~----------------------150+
68
100.0
68
0.0
o
o

1
N

o
.....•

�Table 1.

Number of band-tailed

Area

Date(s)

Stoner

June 15 &amp; 16

Collbran

June 18

Aspen

Evergreen

Meeker-Buford

Montrose

Trinidad
Arboles

Adu1tJ/
No. Banded
Percent

88

100.0

Immatures
No. Banded
Percent

o

0.0

o

0.0

Totals

Estimated No. of
Birds Using
Trapping Sites

88

100+

----------------------------------------------------------------------------------~-----------69
100.0
69

100+

-----------------------------------------------------------------------------------------------June 20
July 24 &amp; 25

62
51

100.0
68.0

o

24

0.0
32.0

62
75

100+
100+

------------------------------------------------------------------------------------------------

June 21
July 2 &amp; 3
July 25 &amp; 26
August 29
September 5

32
90
62
9

27

100.0
84.1
62.6
40.9
50.9

o

17
37
13
26

0.0
15.9
37.4
59.1
49.1

32
107
99
22
53

100+
200+
150+
40+
75+

-----------------------------------------------------------------------------------------------June 25
July 20 &amp; 21

57
21

100.0
95.5

o

1

0.0
4.5

57
22

80+
40+

-----------------------------------------------------------------------------------------------June 26 to July 5
July 23
August 13

171
98
24

98.8
86.7
16.7

2

15
120

1.2
13.3
83.3

173
113
144

700+
500+
400+

------------------------------------------------------------------------------------------------

July 1 &amp; 2

207

89.6

24

10.4

231

400+

-----------------------------------------------------------------------------------------------July 10

41

89.1

5

10.9

46

60+

------------------------------------------------------------------------------------------------

\.Joodmen

July 10
August 7 &amp; 8

Ft. Garland

July 16-24

Monte Vista

pigeons banded by area in Colorado, 1974 (continued).

69
82

67.0
58.6

34
58

33.0
41.4

103
140

150+
200+

-----------------------------------------------------------------------------------------------158

66.1

81

33.9

239

500+

-----------------------------------------------------------------------------------------------July 29
August 12

68
29

53.1
37.7

60
48

46.9
62.3

128
77

600+
500+

-------------------------------------------------------------------------------------------------

I
N

o
co
I

�Table 1.

Number of band-tailed pigeons banded by area in Colorado, 1974 (continued).

AdultJ./
Percent
No. Banded

Immatures
No. Banded
Percent

Area

Date(s)

LaVeta

July 31

52

94.5

3

September 11

49

44.1

62

55.9

3 ,04~/

77.0

912

23.0

Sedalia
Bayfield
Estes Park
Total

5.5

Totals

Estimated No. of
Birds Using
Trapping Sites

55

75+

-----------------------------------------------------------------------------------------------35+
24
12.5
3
87.5
21
August 1
-----------------------------------------------------------------------------------------------300+
190
29.5
56
70.5
134
August 15 &amp; 16
-----------------------------------------------------------------------------------------------150+
111
3,960

~1~/--------------------------------------------------------------------------------------------------------'~
- Includes pigeons classified as subadu1ts.
_2/ Includes 17 adults released from the aviary at Evergreen (14) and Longmont (3).

I
N

o

�-210-

captured (difference of over 17 percent) is not Qqderstood especially when
percent juvenile males and females in trap samples was not significantly
different from 1:1. It is possible that differences exist in molting rates
of immature males and females and in survival to the subadult and adult age
classes.
Time of day was recorded for most pigeons (5,075) captured (newly banded
and recaptures) in 1974. A summary of these data is presented in Table 2.
As in other years, more males than females (76.4:23.6 percent) were
trapped prior to 1000 hours MDT. Between 1000 and 1600 hours, more females
than males were captured (70.3:29.7 percent) while after 1600 hours,
differences were not as great, as 58.7 percent of the birds trapped were
females while 41.3 percent were males. These data are similar to that from
previous years. Most adults were trapped between 0600 and 0959 hours
(50.3 percent of all captures), with lesser percentages being trapped from
1000 to 1559 hours (42.9 percent), and after 1600 hours (6.8 percent).
These differences probably explain the imbalance in sex ratios of adult
pigeons newly banded. Immatures were caught more frequently before 1000
hours (67.3 percent of all immature captures) than in any other time period.
Reasons for this anomaly are not und'ezs t ood but this pattern has remained
constant during all years for which data are available.
During the 1974 trapping period, 1,229 previously banded birds were recaptured
with all but 14 being initially banded by Division personnel. Of the 14
"foreign" bandings recaptured, 12 had been initially banded in New Mexico,
1 in Utah, while 1 had been banded by a private bander in Colorado.
Considering the 1,215 birds initially banded by Division personnel, 23 were
banded in 1969, 108 were banded in 1970, 192 were banded in 1971, 338 had been
banded in 1972, 340 had been banded in 1973, while 214 were newly banded in 1974.
In all, 230 pigeons recaptured had moved at least one degree block from where
'
initially banded. Of this number, 128 changed flock areas. Thirty-three
birds banded in Colorado were reported recaptured in New Mexico (31), and
Utah (2). All of these pigeons had been banded prior to 1974. While it
is probable that some of these pigeons were migrating to or from Colorado
when trapped, it is also likely that some flocks of pigeons occupy areas
partially in two states. This is especially true for flocks of pigeons near
Monte Vista-Del Norte, Durango-Arboles, and LaVeta-Ft. Garland as 25 of the
31 recaptures in New Mexico originated from bandings of these three flocks.
Of the 29 birds that were recaptured out of Colorado for which sex was ascertained at time of banding, 20 were females, while 9 were males. Only 3 of the
33 recaptures out of Colorado were banded as immatures while 1 was banded as
a subadult.

Hunting Season
Band-tailed pigeon hunting, while no longer experimental, was again closely
monitored in Colorado in 1974. Hunting was allowed from September 7 through
October 6, with a daily bag limit of 5 birds and a possession of 10. During
the season, all of the state west of Interstate Highway No. 25 was open to

�Table 2.

Time HDT

Time of day of band-tailed

Percent of All
Successful
Trap Attempts

pigeon trapping, 1974.

1/
Percent of all AdultCaEtures
Males
Females

1/
Percent ofAll Adult Males
Captured

1/
Percent ofAll Adult Females
Captured

Percent of
All Immatures
Captured

1/
Percent ofAll Adults
Captured

0600-0959

41.5

76.4

23.6

71.2

25.9

67.3

50.3

1000-1559

50.4

29.7

70.3

23.6

65.5

26.5

42.9

1600-

8.1

41. 3

58.7

5.2

8.6

6.2

6.8

1/ Including those birds classified as subadults.
I
N

~
~
I

�-212-

hunting. As in previous years all hunters had to obtain a free permit
obtainable at 9 Division offices and the headquarters of the Monte Vista
National Wildlife Refuge. No limit was placed on total number of permits
to be issued, and 518 permits (an additional 4 permits may have been issued)
were obtained by hunters. This was an increase of 63 (13.8 percent) from
1973.

Hunter Questionnaire

Survey

The hunting season for bandtails in 1974 closed at sunset on October 6. On
October 7 and 8, letters containing a self-addressed, postage paid return
card were sent to all permittees. Four questionnaires (0.8 percent) were
returned for lack of sufficient address. On October 28 a followup letter
was sent to the 167 permittees who had failed to respond to the initial
letter. Results of the questionnaire survey are presented in Table 3. Data
for the 53 non-respondents were calculated using mean values of the respondents
to the followup survey.
Interest in pigeon hunting increased slightly in 1974 over 1973 but did not
return to the previous high level reached in 1972. It would appear that the
novelty of having a pigeon hunting permit has dissipated and that only hunters
who believe they may see pigeons are now obtaining permits. For a large number
of hunters (44.4 percent), pigeon hunting remains an opportunistic event.
Less than 300 permittees can be considered serious pigeon hunters. The
number of hunters in this group has remained fairly constant since hunting
was initiated in 1970.
Hunter success increased in 1974 from 1973 levels (45.8 versus 28.3 percent).
Total harvest in 1974 increased significantly from the 366 birds reported
bagged in 1973. Number of days per hun~er remained at about the
same level observed in previous years. It would appear that regardless of
pigeon availability, most hunters will not hunt more than 2 or 3 days per
season. Number of pigeons bagged per successful hunter has also changed very
little from 1970-74 regardless of pigeon availability.
Percent crippling
loss reported in 1974 wa3 lower than in previous years and does not adequately
represent actual conditions. Observations of hunters and pigeons during 1974
and previous years suggest that actual crippling loss may be as high as 20
percent of the birds retrieved.
As in other years, hunter activity and harvest rapidly decreased after the
initial weekend. In 1974, 44.0 percent of the harvest occurred on the opening
weekend, 11.7 percent on the second weekend, 5.7 percent occurred on the third
weekend, while 4.8 percent occurred on the last weekend of the season. Not
surprisingly, only 29.7 percent of the total harvest occurred during the nonweekend portion of the season (20 days). About 68 percent (68.3) of the total
harvest occurred during the first 9 days of the season.
Hunting pressure in 1974 was diffuse as 251 hunters reported hunting in 43
different locations. The Monte Vista (15.1 percent) and Durango (9.2 percent)

�-213-

Table 3.

1/

Harvest statistics, 1974 band-tailed pigeon season.-

Category

Response
First
Letter

Response
Second
Letter

Sum
First and
Second Letters

Number of permittees
responding

351(67.8)

114(22.0)

465(89.8)

Number of permittees
hunting

203(57.8)

58(50.9)

261(56.1)

288(55.6)

Number of permittees
not hunting

148 (42.2)

56(49.1)

204(43.9)

230(44.4)

Number of successful
hunters

98(48.3)

23 (39.7)

121(46.4)

132(45.8)

Number of hunter days

564

145

709

776

Days hunted per
hunter
Number of pigeons
bagged

2.8

594

2.5

2.7

689

95

Projected
For all (518)
Permittees

2.7

734

Pigeons per successful hunter

6.1

4.1

5.7

5.6

Pigeons per hunter

2.9

1.7

2.7

2.5

Number of pigeons
crippled and lost

40

13

53

58

Pigeons crippled and
lost per hunter

.2

.2

Total harvest (bagged +
crippled and lost)
634

105

Percent crippling loss

12.4

6.3

.2

.2

739
7.2

792
7.6

1/ Values in parentheses are percentages.
areas continue to be preferred for hunting, and these 2 areas had 24.3 percent
of all hunting trips. Areas west of Pueblo and near La Veta each received
6.4 percent of the hunter trips while Bear Creek near Salida received 5.2
percent of the reported pressure. On a regional basis, 48.6 percent of the
hunters hunted in southwestern Colorado, 30.7 percent hunted in southeastern

�-214-

Colorado, 11.6 percent hunted in northeastern Colorado, and 9.2 percent
hunted in northwestern Colorado. Reported harvest paralleled hunter pressure
as 49.4 percent of the harvest occurred in the southwest, 29.8 percent in the
southeast, 10.6 percent in the northeast and 10.2 percent in the northwest.
These data are similar to that collected in all years since the hunting
season reopened in 1970.

Wing Survey
Packets containing 5 wing envelopes with instructions to place one day's
bag in each envelope were issued to a sample of hunters, mostly in the
Monte Vista and Durango areas. In all, packets were issued to 155 permittees.
Envelopes containing 138 wings were received from 33 hunters. Additional
data were received concerning hours hunted (148), birds crippled and lost
(15), location of kill and banded birds harvested (21). Wings from an additional 33 birds, including two that were banded, were received via non-wing
survey mail. Of the 171 wings received, 23 (13.4 percent) were aecompanied
by bands. Percent bands received varied from 19.4 in the San Juan Basin,
16.9 in the San Luis Valley to 11.8 near Salida.
Of the 171 classified as to age, 65 (38.0 percent) were from immatures, 7
(4.1 percent) were from subadults while 99 (57.9 percent) were from adults.
The 38.0 percent immatures was the highest recorded in the five years for
which data are available. While juveniles may be more vulnerable to hunting,
data from both trapping and field observations indicated that production in 1974
was above average. Molt data were obtained for the 65 immature wings processed.
Of this sample, 22 had not molted primary 1, 13 were molting primary 1, 15
were molting primary 2, 9 were molting primary 3, 4 were molting primary 4,
while 2 were molting primary 5. Thus 30 of the immatures had hatched prior to
1 July. These data do not support the hypothesis of early migration of immatures.

Hunter Field Checks
During the 1974 season, emphasis was again placed on contacting pigeon hunters
in the field in order to examine harvested birds for crop gland activity.
Crops of all adult pigeons checked were classified as being active (crop
gland with curds), stimulated (gland apparent, but no curds), or inactive.
In addition, sex by gonadal inspection was obtained for as many birds as
possible. In all, 153 pigeons were checked, with 52 being immatures (34.0
percent), with the remainder (101) being classified as adults. Of the adults,
19.8 percent were classified as having active crop glands, while 32.7 percent
were classified as being stimulated. Percent of crops showing glandular activity
(active + stimulated) was lower in 1974 than in 1973 (52.5 versus 63.2) and
was similar to that observed in 1970 and 1972. Data from all years except 1971
indicate that pigeons continue nesting in Colorado into September. This
is apparently a normal occurrence.
Data on sex of harvested pigeons were available for 91 adults. Of this number
54.9 percent were males and 45.1 percent were females. Of the males, 50.0

�-215-

percent exhibited some glandular development, while 53.7 percent of the females
showed development of the crop gland. These data are similar to that collected
in 1973 but dissimilar to that collected in 1970-71-72 which showed that more
males than females were associated with nesting at time of death. When samples
of adults examined by area are compared, it is apparent that nesting activity
during the hunting season varied by area. Thus only 20 percent of the 10
adults examined near Bayfield were still associated with nest activities.
This percentage ~as 75.0 in the Salida area (N = 32) but only 44.4 near
Monte Vista-Del Norte (N = 54). Realistically, samples were too small to
draw meaningful conclusions concerning differences between areas.

Mortality
Recoveries of 137 band-tailed pigeons initially trapped and banded in Colorado
have been reported to date in the 1974 recovery year (1 May 1974 to 30 April
1975). Six of these were initially banded in 1969, 9 were banded in 1970,
15 were banded in 1971, 38 were banded in 1972, 27 were banded in 1973 while
42 were banded in 1974. All but 12 were shot recoveries.
Of the 125 shot
recoveries, 82 were recovered in Colorado, 39 in New Mexico, 2 in Mexico
(Chihuahua), and 1 each in California and Utah. Hunting mortality in New
Mexico continues to be higher than initially expected as 31.2 percent of the
mortality due to hunting in 1974 was in that State. As expected, hunting
mortality was highest in Colorado (65.6 percent of all shot recoveries).
In 1974, New Mexico split their pigeon hunting season north and south of U.S.
Highway 60. The season in northern New Mexico started 1 September while it
started on 12 October in southern New Mexico. Of the 39 reported recoveries
in New Mexico 23 were from the first season while 16 were from southern New
Mexico. Seventeen of the recoveries were from September 1 through 6, before
the season had opened in Colorado. Thus, migration in 1974 had begun prior
to the hunting season in Colorado. Distribution of recoveries in New Mexico
indicates that the fall migration is characterized by slow movement in a
southwest direction from Colorado.
Of the 137 recoveries, 37 (27.0 percent) had been banded near Monte VistaDel Norte, 21 (15.3 percent) in the Pagosa Springs-Arboles-Bayfield-Durango
area, 17 (12.4 percent) near Salida, 11 (8.0 percent) near Montrose-Sanborn
Park, 10 (7.3 percent) near LaVeta-Ft. Garland, 9 (6.6 percent) near Evergreen
and 8 (5.8 percent) at Woodland Park. No other area provided more than 5 band
recoveries.
In all, 17 different flock areas provided recoveries in 1974 with
five flock areas providing 70 percent of the recoveries.
Of interest was the
contribution of Salida and Montrose-Sanborn Park in 1974 as these two areas
normally rank below LaVeta-Ft. Garland.
Slight differences were noted in vulnerability of adult males and females as
males provided 57 (55.3 percent) of 103 recoveries for which sex was ascertained
at time of banding. First year recoveries of birds banded as immatures totaled
16. This represented 38.1 percent of the 42 total first year recoveries of
banded adults and immatures. Thus immatures are indicated to be more susceptible
to hunting than adults as this age class represented only 23.0 percent of the
total bandings in 1974.

�-216-

Breeding Phenology
Few data on breeding phenology were available as this portion of the
study was completed in early 1973 (Gutierrez 1973). Data were available from examination of 53 birds killed in trapping operations (36),
trichomoniasis tests (13), poaching (2) and electrocution (2). Thirtythree of these birds were subadult and older. Data from these birds
indicated that nesting had started by 13 May and continued throughout
the summer into October. Observation data indicated that a few pigeons
started arriving in Colorado in early April with peak spring migration
in mid to late May. Some pigeons remained as late as early November
although most had left by 10 October. Thus, observational data support
those obtained from gonadal and crop examinations.

Wing Measurement

and Molt Data

Data concerning length of primaries, rectrices, total body length, carpal length and bill length were obtained from most dead pigeons examined
in 1974. These data will be compiled and analyzed with those from 196973 to examine differences between age and sex classes of pigeons. Prior
to dissection of collected birds, plumage characters of each adult specimen
and some juveniles were examined and recorded. Each bird was then classified as either male or female. Upon dissection, gonads were examined and
actual sex of each bird was compared with the classification based on
plumage appearance.
Comparative data were available for 32 adults and subadults (13 males and 19 females), and 4 immatures. Only 1 bird was inaccurately sexed. This bird appeared to be an adult male by plumage, but
was actually a female. This percentage of error (2.8) is similar to that
found in prior years.
Information concerning primary molt was available for 998 immatures (911
banded and released, 65 from the hunter wing survey and 22 from birds
necropsied).
Hatching dates were estimated using data from White (1973).
Estimated hatching dates for all immatures for which data were available
are presented in Table 4.
As in earlier years, hatching was spread over
a long time period (12 April to 2 October). Only one major peak can be
ascertained from data in Table 4 and the data best fit a bell shaped curve.
It would appear then that calculated hatching dates only represent the range
of nesting in Colorado. Of importance, data in Table 4 indicate that a sharp
decrease occurred in hatching after 10 July as only 10.5 percent of those
examined hatched after that date. All observational and crop data collected
in August and September refute the hypothesis that most nesting ceased in
early July. These differences relate to trapping effort as most trapping
terminated in mid August. Consequently progeny of nests after mid July
are not represented in the trap sample as it takes 25-30 days from hatching
to fledging. Examinations of hatching dates for the 65 immatures collected
in the wing survey (7 September and later) reveals that 35 (53.8 percent)
hatched after 10 July. However, only 24 (36.9 percent) of this sample
hatched after 31 July.

�-217-

Table 4.

Estimated hatching dates for wild band-tailed pigeons 1974.

Number
Hatching

Percent

July 1-10

118

11.8

0.3

July 11-20

34

3.4

11

1.1

July 21-30

24

2.4

May 2-11

26

2.6

July 3l-August 9

38

3.8

May 12-21

99

9.9

August 10-l9

5

0.5

May 22-31

120

12.0

August 20-29

3

0.3

June 1-10

234

23.5

August 30-September 8

0

0.0

June 11-20

150

15.0

September 8-17

0

0.0

June 21-30

132

13.3

After September 17

1

0.1

Number
Hatching

Percent

Prior to April 11

0

0.0

April 12-21

3

April 22-May 1

Time Period

Time Period

Parasitic Load

Information on helminths was available for 38 pigeons necropsied.
Six (all
adult or subadult) had infections of helminths. The infection rate of 15.8
percent is similar to that found in collections from 1969-73 (range 9.5 to
18.3 percent). One paper was prepared and submitted for publication on new
species of nematodes from band-tailed pigeons. Another paper on trichomoniasis
in band-tailed pigeons has been submitted and accepted for publication.

LITERATURE CITED

Colo. Div. Game, Fisb
Braun, C. E. 1970. Band-tailed pigeon investigations.
and Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. p. 151-171.
1971. Band-tailed pigeon investigations.
Colo. Div. Game, Fish and
Parks. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. p. 209-236.
1972. Band-tailed pigeon investigations.
Colo. Div. Wild1.
Res. Rept., Fed. Aid Proj. W-88-R. Oct. p. 123-142.

Game

�-218-

Braun, C. E. 1973. Band-tailed pigeon investigations.
Colo. Div. Wildl.
Game Res. Rept., Fed. Aid Proj. W-88-R. Oct. p. 107-123.
1974. Band-tailed pigeon investigations.
Game Res. Rept., Fed. Aid Proj. W-88-R. Oct.

Colo. Div. Wildl.,
p. 153-168.

Gutierfez, R. J. 1973. Band-tailed pigeon investigations:breeding
and
nesting chronology studies. Colo. Div. Wildl. Game Res. Rept.,
Fed. Aid Proj. w-88-R. Oct. p. 153-177.
White, J. A. 1973. Band-tailed pigeon investigations:plumage
studies.
Colo. Div. Wildl. Game Res. Rept., Fed. Aid Proj. W-88-R. Oct.
p. 125-152.

Prepared by __

"",,~~/_~_.
--:.f~~~l,,",!~~=~
Cla.it E. Braun
Wildlife Researcher

_

�October 1975

-219-

JOB PROGRESS REPORT

State of

C=-O::..;L::..:O::.::RAD=::.::O~-----

Project No.

W-88-R-20

Work Plan No.

;5

Migratory Bird Investigations
_

Job No.

1
----------~------------------

Job Title.__~I~n~v~e~s~t~i~g~a~t::.::~~·o~n~s~o~f~C=_omm~=_o~n~S~n;iLP~e~i~n~C~o~l~o~r~a~d~o~
_
Period Covered:

April 1, 1974 to March 31, 1975

Personnel:
C. E. Braun, H. D. Funk, and B. R. Johnson, Colorado Division
of Wildlife; and R. A. Ryder, Department of Fishery and Wildlife Biology,
Colorado State University.
ABSTRACT

Common snipe (Capella gallinago delicata) utilized open water wintering habitat
in the Fort Collins area in relatively high numbers (8-12 snipe per km) from
November through April. Censuses in 1974 on 12 study areas (total 244.3 ha)
in the San Luis Valley, North Park, Fort Collins area, and Yampa Valley
indicated overall snipe densities of 0.63 and 0.47 snipe flushed per ha in
May and June, respectively.
The peak fall migration period for 1974, based
on censuses in September and October, apparently occurred during the third
week in September.
Results of the 1972 and 1973 small game questionnaires
(N = 22) show a mean harvest of 3.7 snipe per hunter and 4.0 days spent
afield. Results from a 1973 and 1974 special snipe survey (N
19) indicated
eight had hunted snipe during this period and had a mean harvest of 10.8
snipe per hunter and spent 4.9 days afield.

=

��-221-

INVESTIGATIONS OF COMMON SNIPE IN COLORADO
Bruce R. Johnson

This second annual progress report on investigations of snipe in Colorado
concerns field activities such as selection of study areas, monitoring
numbers of snipe and evaluating important habitat characteristics on each
area. Data have been compiled on densities of snipe occurring on these
areas during spring and fall migrations and the breeding seasons. Additional
data concerning numbers of snipe wintering in the Fort Collins area are also
included.
P. S. OBJECTIVE
To study the distribution and status of common snipe in Colorado in order that
data might be made available to improve management of the species.
SEGMENT OBJECTIVES
1.

To identify and delimit range of common snipe in Colorado.

2.

To document seasons of occurrence and estimate relative numbers in four
major areas.

3.

To document hunting pressure and estimate harvest and hunter success.

4.

To compile data, analyze results, prepare progress reports and publish
findings in appropriate technical journals.
METHODS
Selection of Study Areas

Twelve study areas were selected in the San Luis Valley, Fort Collins area,
North Park, and the Yampa Valley. Selection of each area was based on (1)
presence of snipe performing mating and territorial displays, (2) permanence
of water, (3) similarity to general habitat types in each region, (4) contradistinction from other selected study areas, and (5) sufficient area to
provide adequate numbers of snipe.
Census Techniques
A systematic strip census was employed to ascertain numbers of snipe on
breeding ground study areas. In this method, census routes consist of
straight line transects at 20 to 25 meter intervals throughout the area.

�-222-

Field maps of all areas were used to maintain a consistent route and record
all snipe flushed and alighting, thus avoiding duplicate counts. A similar
method was used on the winter area, however, the census route adjoined the
stream channel. These above methods as well as those used in evaluating
habitat characteristics have been described in previous reports.
RESULTS AND DISCUSSION
Wintering Area Censuses
Censuses on a snipe wintering area near Fort Collins have been conducted from
January 27, 1974 to the present. Snipe utilize this open water area in
relatively large numbers (approximately 8 to 12 snipe per km) from November
through April. In 1974 snipe had essentially departed the area by May and
returned in small numbers in October. Censuses will continue on this wintering
area until spring departure of snipe occurs.
Spring Migration Period
Because of difficulties in selecting breeding ground study areas, censuses
were delayed until early May. Consequently, no data were compiled concerning
the 1974 spring migration. Currently, such data are being collected and will
be discussed in the final report.
Breeding Ground Censuses
Censuses were conducted from early May through early August to estimate numbers
of snipe breeding on the 12 study areas. Data from May and June censuses by
region have been summarized in Table 1.

Table 1.

Mean numbers of snipe flushed per hectare, Colorado 1974.

Major Region

Hectares

SniEe Flushed Eer Hectare
May
June

Fort Collins

25.1

1. 35

0.83

North Park

92.0

0.56

0.46

Yampa Valley

63.6

0.60

0.48

San Luis Valley

63.6

0.47

0.33

Total

244.3
0.63

0.47

Mean

�-223-

The progressive decline from May to June is probably a result of increased
nesting and rearing activities by snipe, consequently, snipe were more
difficult to flush. Data from July and August censuses were less reliable
because birds of the year were able to fly at this time. Eighteen snipe
nests were located during censuses with estimated onsets of incubation
ranging from May 1 to July 4, the majority of which occurred from mid- to
late May. Censuses in 1975 were initiated in late March and will continue
through mid-June to supplement 1974 data.
Fall Migration Period.--Data collected on 8 of the 12 breeding ground study
areas indicated maximum numbers of snipe were observed in the San Luis Valley
on September 15-16, in Fort Collins on September 17, and in North Park on
September 21. From this, the peak fall migration period for 1974 apparently
occurred during the third week in September.
Snipe Hunter Survey.--Hunters indicating they hunted snipe through the
regular annual small game harvest survey in 1972 and 1973 (N = 22) had a mean
harvest of 3.7 snipe per hunter and spent 4.0 days afield. Special snipe
hunting questionnaires were sent to specific hunters in 1973 and 1974. Of
those who responded to this special survey (N = 19) eight indicated that they
had hunted snipe. These respondents had a mean harvest of 10.8 snipe per
hunter and spent 4.9 days afield. Although sample sizes in both surveys
were small, the harvest of snipe by Colorado hunters appears small and hunting
is an insignificant mortality factor.

Habitat Characteristics
Data on habitat characteristics such as vegetation composition, soil types
and compaction, and water characteristics have been compiled. A summary of
these data will be reported in the final report.
Final Report Plans
The final report of this project in form of a M. S. Thesis will be prepared
after termination of field work in late June. This report will be available
by September 1, 1975.

Prepared by
Bruce R. Johnson
Graduate Research Assistant

��October 1975

-225JOB PROGRESS REPORT

State of

~C~O~L~O~ru~\D~O

Proj ect No.

W-88-R-20

\.;'ork
Plan No.

6

_
Migratory

Bird Investigations

Job NO.

Job Title

Migratory Bird Publications

Period Covered:

April 1, 1974 through March 31, 1975

Personnel:

~l~

_

Howard D. Funk

ABSTRACT
The Colorado Division of Wildlife cooperated with the remainder of the
Central Flyway States in the preparation of a waterfowl identification
guide to be distributed free of charge to hunters. Printing was completed during the segment with the South Dakota Department of Game, Fish
and Parks contracting with the printer and Colorado &amp;,d the other states
reimbursing South Dakota for their individual costs. The booklet is
titled "Waterfowl Identification in the Central Flyway".
It is 51 pages
in length and printed on plastic paper. Color plates throughout the
booklet include males in full winter and eclipse plumage. females in
winter plumage, and wings of each sex for each species. Text and plates
of birds in flight are also included to aid the hunter in identification.

Prepared by
Howard D. Funk
Small Game Section Chief

.:

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                  <text>January 1976

-1-

JOB PROGRESS

State of

COLORADO

Project No.

W-4l-R-2S

Work Plan No.

1

Job Title Evaluation

of the Nutritional

Period Covered:
Personnel:

REPORT

Bighorn·Shee~

&amp; Mountain Goat Investigations

Job No.

~l~

Requirements

of Bighorn Sbeep

June 1, 1974 - May 31, 1975

Robert E. Keiss

ABSTRACT
A study was designed and implemented to test for differences in major
nutritive components of forages from three bighorn sheep study areas.
The study was designed so that statistical comparisons could be made
between the three areas (Pikes Peak, Buffalo Peaks, and Trickle Mountain)
at three seasons of the year (Summer, Fall, Winter).
Chemical analysis
for percent composition of protein, soluble ash, etLer extract, soluble
carbohydrate, lignin, acid insoluble ash and holocel1ulose were made on
each sample. Mineral levels including calcium, phosphorus, sodium,
magnesium, iron, zinc, copper and selenium were determined in each sample.
Bighorn sheep lambs collected during 1972, 1973 and 1974 were subjected
to mineral level analysis on selected tissues. The mineral levels of
calcium, phosphorus, sodium, potassium, zinc, iron, and copper were determined in hair, liver, spleen, kidney, skeletal muscle and heart muscle
of each lamb.

_

��-3-

EVALUATION

OF THE NUTRITIONAL

REQUIREMENTS

OF BIGHORN SHEEP

Robert E. Keiss

P. S. OBJECTIVE
To develop methods: (a) for the evaluation of nutritional requirements of
bighorn sheep under natural conditions, and (b) to measure the adequacy of
selected ranges to meet these requirements.

SEGMENT OBJECTIVES
1.

To establish
horn sheep.

the nutritional

level of forage species utilized

by big-

2.

To evaluate if the nutritional level of selected bighorn sheep ranges
is adequate to meet nutritional requirements of bighorn sheep at all
seasons of the year.

METHODS AND MATERIALS
Methods used to estimate forage species utilized by bighorn sheep were
previously described (Schoonveld 1973; Keiss and Schoonveld 1974). During
this segment forage samples were collected from three study areas (Pikes
Peak, Buffalo Peaks, Trickle Mountain) following a sampling scheme which
would allow the analytical chemical data to be subjected to statistical
comparisons.
Collections were made at three seasons of the year (Summer - July; Fall September; Winter - November-December)
in order to evaluate the changes in
nutritional quality of bighorn sheep forages as the growing season progresses
from summer into fall and winter and plant species grow, mature and go into
dormancy.
Because of the complexity of the composition of different plant species on
the three study areas this phase of the study limited the nutritional comparisons to general plant types (grass and browse) rather than attempt the
complex comparisons between individual plant species. The main purpose was
to compare the gross nutritive quality of bighorn sheep forage on the three
study areas at three seasons of the year. For this reason samples were
designated as "grass" and "browse" and no further attempt was made to differentiate plant species composition.
Five transects were selected on each of the three study areas. These transects were situated to cover known winter range areas used by bighorn sheep.
A wooden frame measuring one meter on each side was used to aid in plant
collections.
The frame was placed on the ground at one end of the transect
and all grass within the square meter was clipped and placed in a paper sack.
After the grass within the frame was collected we moved one hundred paces
along the transect and at this point collected another one square meter grass

�-4sample. A total of ten samples, over a one thousand pace transect, were
obtained and returned to the laboratory for analysis.
Browse samples were collected from selected browse plants along "modified"
transects.
Because of the sporadic distribution of browse plants on the
bighorn sheep winter rauges, it was necessary to select groups of plants
to sample, and a rigid sampling scheme similar to that described for grass
was not possible.
For the most part browse was collected from plants
closest to points where grass samples were collected.
Current annual growth
was clipped from plants for chemical analysis.
All forage samples were returned to the laboratory and allowed to air dry.
They were then ground in a Wiley Mill (equipped with stainless steel screens)
and placed in glass jars, oven dried at 600c for 24 hours and stored for
future analysis.
Because of the large ·.numberof individual samples it was not practical to
do a complete chemical analysis on each one, so each transect was considered
as a sample unit for comparison purposes.
All samples collected along a
single transect were composited into a single sample. This was accomplished
by thoroughly mixing each individual forage sample and removing a ten gram
subsample.
These subsamples were then composited and the composite sample
was thoroughly mixed and submitted for chemical analysis.
The above described sampling scheme resulted in five grass samples and five
browse samples representing five transects from each of the three study areas
at three seasons of the year so that the study design resulted in a total of
ninety samples for complete laboratory analysis.
All forage samples were analyzed for their proximate constituents by procedures described previously (Keiss and Schoonveld 1974). These analytical
procedures partition forages into the following constituents:
Protein, Ether
Extract, Soluble Ash and Soluble Carbohydrates, Lignin, Acid Insoluble Ash
and Holocellulose which is composed of Cellulose and Hemicellulose.
All forage samples were analyzed for their trace mineral content by methods
previously described (Keiss and Schoonveld 1974). These minerals include
Calcium, Phosphorus, Magnesium, Sodium, Potassium, Zinc, Iron, Copper and
Selenium.
Tissue samples from bighorn sheep lambs collected during previous segments
of this study were analyzed for trace mineral levels. At the time these
lambs were necropsied tissues were collected and frozen for this purpose.
Seven lambs were collected during the summer of 1972, nine lambs collected
during the summer of 1973 and seven lambs collected during the summer of
1974. The following tissues were analyzed from each lamb: Hair, Liver,
Spleen, Kidney, Skeletal Muscle, and Heart Muscle.
The tissues were removed from the freezer, allowed to thaw and placed in a
drying oven at 600C until dry. Low drying temperatures were used because
of the reported volatility of selenium from tissues when dried at high
temperatures.
A one gram sample of each tissue was weighed into a dtgestion vessel and
digested by the method of Adrian 1973. This wet ashing was accomplished
by placing one gram sample in a 4 oz Nalge bottle.
Two milliliters of

�-5perchloric acid-sulfuric acid (7+l) plus four milliliters of nitric
acid was added to each bottle and allowed to pre-digest for twenty four
hours at room temperature.
The digestion bottles were stoppered, placed
in a water bath at 700C for 3 hours. It is important that the bottles
are tightly stoppered as the pressure within the bottle is essential to
the procedure.
After the digestion was completed each sample was diluted
with distilled water until the total volume equaled 100 mI.
All mineral determinations
following methods:

were made from the digested

sample by the

Phosphorus - vanadate-molybdate procedure.
Selenium
- fluorometrically using 3,3' Diaminobenzidine
Calciumm Magnesium, Potassium, Iron, Copper, Sodium and Zinc atomic absorption spectrophotometer
(Perkin-Elmer 303)

RESULTS AND DISCUSSION
The results of the analysis of grass and browse samples collected as a part
of this study are presented in Tables 1 through 12.
The statistical analyses of these data are still in progress and the results
of this work and its significance will be discussed in the job completion
report which will be prepared during the next segment of this study.
The bighorn sheep lamb tissue analyses for trace mineral levels have been
completed and the analysis of the data is in progress.
Table 13 lists the
animals which were examined as a part of this study and identifies them as
to their origin and date of collection.
Tables 14 through 20 give the data
on mineral levels of selected tissues from these bighorn sheep lambs as determined by analytical procedures.

BIBLIOGRAPHY
Adrian, W. J. 1973. A comparison of a wet pressure digestion method with
other commonly used wet and dry-ashing methods.
Analyst 98:213.
Keiss, R. E., and G. G. Schoonveld.
requirements of bighorn sheep.
W-4l-R-23.
January.
pp.3-78.

1974. Evaluation of the nutritional
Colo. Div. Wildl. Fed. Aid Proj.

Schoonveld, G. G. 1973. Evaluation
horn sheep. Colo. Div. Wildl.
2. pp. 75-88.

of the nutritional requirements of bigFed. Aid Proj. W-4l-R-22.
January, Part

Prepared by

~~~~~~ ~~~~_:&gt;~~~==~.~
_
-'fob!rt1.
KYs7"
z&lt;

Wildlife Researcher

�Table 1.
Pikes Peak grass - chemical partitioning of the nutritive constituents of grass samples from five
transects on Pikes Peak Study Area.

Season
of
Collection

Transect
No.

Percent
of
Total

I

Cell Contents
Ether
Soluble
Protein
Extract
Ash

Soluble
Carbo

Cell l.,Talls
Acid InHolosoluble
cellulose
Ash %
%

%

%

%

%

Percent
of
Total

34.72

8.80

6.73

7.70

11.49

65.28

6.53

5.85

52.90

II

35.16

8.80

8.37

6.72

11.27

64.84

6.48

5.94

52.42

III

34.12

8.50

9.62

6.91

9.09

65.88

6.92

5.63

53.33

IV

34.17

8.40

6.91

7.08

11. 78

65.83

6.38

5.73

53.72

V

35.63

8.30

7.37

7.21

12.75

64.37

6.84

4.62

52.91

Summer

Lignin
%

I

I

Fall

Winter

31.62

7.80

.9.94

4.56

9.32

68.38

8.37

5.87

54.14

I

II

30.63

7.70

9.27

4.63

9.03

69.37

8.58

5.78

55.01

III

31.37

7.10

8.63

5.92

9.72

68.63

8.43

5.73

54.47

IV

32.51

7.70

9.81

4.81

10.19

67.49

8.62

6.59

52.28

V

33.27

7.90

9.37

5.09

10.91

66.73

8.79

6.30

51.64

I

24.62

5.00

4.37

7.37

7.88

75.38

9.48

5.78

60.12

II

24.72

5.10

4.47

6.37

8.78

75.28

10.73

6.37

58.18

23.23

5.00

5.17

7.94

5.12

76.77

8.77

6.69

61.31

IV

23.76

5.70

6.21

6.38

5.47

76.24

8.92

6.30

61.02

V

24.89

5.10

4.21

5.91

9.67

75.11

10.81

5.31

58.99

III

'"

�Table 2. Pikes Peak grass - mineral analysis of grass samples from five transects on Pikes Peak Study
Area.

-Season
of
Collection

Sunnner

Fall

Winter

Potassium

Magnesium

%

%

Iron
ppm

Copper
ppm

Zinc
ppm

Selenium

%

Calcium :
Phosphorus
Ratio

.46

.17

2.7 : 1

.70

.18

25

3.7

50

.07

II

.46

.16

2.9 : 1

.71

.17

26

3.2

56

.07

III

.44

.17

2.6 : 1

.72

.18

28

3.3

53

.08

IV

.47

.17

2.8 : 1

.75

.18

27

3.7

52

.08

V

.49

.15

3.3 : 1

.77

.17

22

3.5

51

.06

I

.51

.12

4.3 : 1

.63

.14

22

3.2

34

.05

Transect
No.

Calcium
%

I

Phosphorus

%

II

.51

.11

4.6 : 1

.63

.15

23

3.2

36

.06

III

.53

.10

5.3 : 1

.62

.16

24

3.3

38

.07

IV

.51

.10

5.1 : 1

.66

.17

26

3.5

40

.05

V

.52

.11

4.7 : 1

.63

.16

22

3.6

38

.05

I

.62

.05

12.4 : 1

.45

.11

16

2.2

30

.03

II

.63

.04

12.6 : 1

.48

.11

17

2.7

38

.02

III

.62

.05

12.4 : 1

.45

.12

16

2.1

31

.01

IV

.64

.06

10.7 : 1

.49

.11

16

2.2

33

.05

X

.62

.05

12.4 : 1

.44

.10

1"

2.5

34

.03

I

"'-I

I

�Table 3. Pikes Peak browse - chemical partitioning of the nutritive constituents of the browse samples from
five transects on Pikes Peak Study Area.

Season
of
Collection

Summer

Fall

Winter

Cell Contents
Ether
Soluble
Protein
Extract
Ash

Transect
No.

Percent
of
Total

I

34.62

8.80

II

34.71

III

Soluble
Carbo

Cell Walls
Acid InHolosoluble
cellulose
Ash %
%

%

%

Percent
of
Total

12.17

4.32

9.33

65.38

8.73

5.31

51.34

8.80

12.72

5.21

7.98

65.21

8.37

3.31

53.53

35.12

8.80

13.36

4.83

8.13

64.88

8.38

5.20

51.30

IV

34.31

8.10

12.79

4.92

8.50

65.69

8.92

4.11

52.66

V

33.92

8.70

13.27

5.31

6.64

66.08

8.12

4.72

53.24

I

33.19

8.00

-13.27

5.96

5.96

66.81

8.25

5.29

52.27

II

32.31

8.10

11.73

6.73

5.75

67.69

8.93

4.37

54.39

III

34.17

8.00

12.17

5.81

8.19

65.83

8.79

5.18

51.86

IV

33.19

7.80

13.29

6.33

5.77

66.81

8.27

4.47

54.07

V

32.72

7.90

11.69

5.18

7.95

67.28

8.81

5.66

52.81

I

26.31

7.60

8.97

4.93

4.81

73.69

8.39

5.78

59.52

II

25.47

7.60

8.99

5.73

3.15

74.53

8.37

5.72

60.44

III

26.37

7.50

7.63

4.99

6.25

73.63

8.63

6.23

58.77

IV

24.72

7.40

7.83

5.07

4.42

78.28

9.08

6.44

62.76

V

24.37

7.10

7.17

6.31

3.79

75.63

8.81

5.66

61.16

%

%

Lignin
%

I
00
I

�Table 4. Pikes Peak browse - mineral analysis of browse samples from five transects on Pikes Peak Study
Area.

Season
of
Collection

SUIIDl1er

Fall

Winter

Potassium

Magnesium

%

%

Iron
ppm

Copper
ppm

Zinc
ppm

Selenium

%

Calcium :
Phosphorus
Ratio

.52

.19

2.7 : 1

.76

.19

28

3.5

49

.10

II

.51

.20

2.6 : 1

.78

.19

29

3.7

48

.11

III

.53

.19

2.8 : 1

.75

.20

31

3.6

47

.10

IV

.55

.21

2.6 : 1

.77

.19 .

31

3.9

49

.12

V

.56

.18

3.1 : 1

.76

.18

29

4.0

57

.10

I

.50

.15

3.3 : 1

.68

.18

27

4.1

53

.09

II

.51

.14

3.6 : 1

.69

.19

28

4.1

61

.08

III

.50

.15

3.1 : 1

.70

.19

26

4.3

58

.10

IV

.55

.14

3.9 : 1

.72

.20·

24

3.9

60

.10

V

.51

.13

3.9 : 1

.74

.18

25

4.4

53

.09

I

.51

.12

4.3 : 1

.69

.15

21

3.0

56

.07

II

.52

.11

4.7 : 1

.68

.14

22

3.0

50

.08

III

.55

.10

5.5 : 1

.69

.15

21

3.1

55

.07

IV

.51

.12

4.3 : 1

.64

.16

19

3.3

51

.08

V

.52

.12

4.3 : 1

.65

.14

20

3.5

54

.09

Transect
No.

Calcium
%

I

Phosphorus

%

I
\0
I

�Table 5.
Buffalo Peaks grass - chemical partitioning of the nutritive constitutents of grass samples from
five transects on Buffalo Peaks Study Area.

Season
of
Collection

No.

I

39.15

10.40

39.16

10.40

Transect

II
Summer

III
IV
V

III
IV
V
I
II

~"'""=-'-"-"='~

....•.
-=-.~.

41.27

31.63

II
Fall

40.37

39.31

I

Winter

Cell Contents
Ether
Soluble
Protein
Extract
Ash

Percent
of
Total

31.72
32.36

%

10.10
10.50
8.20

%

%

7.93

9.37

60.85

7.08

5.60

48.17

9.73

7.64

11.39

60.84

7.76

4.39

48.69

10.91

7.37

11.39

59.63

7.44

5.31

46.88

8.62

7.73

14.82

58.73

7.59

4.72

46.42

9.21

6.94

12.66

60.69

7.68

5.23

47.78

10.35

6.63

6.45

68.37

7.93

5.72

54.72

11.45

%

8.36

6.72

8.44

68.28

7.82

5.33

55.13

8.30

9.37

7.37

7.32

67.64

7.92

5.73

53.99

10.47

6.54

4.80

69.69

8.18

6.54

54.97

9.36

7.38

6.81

68.35

8.25

6.85

53.25

4.63

7.44

4.40

77.53

9.27

6.34

61.92

4.73

6.37

5.17

77.63

9.16

8.39

60.08

6.30

5.91

6.92

2.16

78.26

10.37

7.85

60.04

76.69
78.28

10.28
10.35

7.76
8.27

58.65
59.66

30.31

8.50

31.65

8.10

22.47

6.00

22.37

6.10

21. 74

IV

23.31

5.80

4.97

7.17

5.37

V

21.72

5.90

3.21

6.38

6.23

"--

Lignin

8.20

III

--

Cell Walls
Acid InHolosoluble
cellulose
Ash %
%

Percent
of
Total

%

10.70

Soluble
Carbo

.-~~ ..
-~.~.
--~=--'

I
I-'
0
I

�Table 6.
Area.

Season
of
Collection

Summer

Fall

Winter

Buffalo Peaks grass - mineral

analysis of grass samples from five transects

on Buffalo Peaks Study

Potassium

Magnesium

%

Calcium :
Phosphorus
Ratio

%

%

Iron
ppm

Copper
ppm

Zinc
ppm

Selenium
%

.61

.18

3.4 : 1

.55

.17

25

3.2

30

.11

II

.67

.19

3.5 : 1

.54

.18

26

3.3

31

.11

III

.63

.18

3.5 : 1

.57

.17

27

3.2

28

.13

IV

.66

.19

3.5 : 1

.56

.18

23

3.5

28

.14

V

.62

.18

3.4 : 1

.54

.17

22

2.9

27

.15

I

.63

.10

3.6 : 1

.48

.15

21

3.0

21

.10

II

.64

.11

5.8 : 1

.49

.14

21

3.0

21

.09

III

.60

.10

6.0 : 1

.47

.14

22

3.0

25

.08

IV

.65

.09

7.2 : 1

.44

.13

25

3.9

24

.10

V

.63

.10

6.5 : 1

.46

.14

22

2.8

30

.10

I

.65

.04

16.3 : 1

.36

.12

15

2.1

16

.09

II

.64

.03

21. 3 : 1

.38

.11

16

2.5

18

.07

III

.63

.04

15.8 : 1

.35

.12

15

2.2

21

.08

IV

.66

.05

13.2 : 1

.35

.11

17

2.1

20

.07

V

.69

.06

11.5 : 1

.35

.09

19

2.3

17

.06

Transect
No.

Calcium

Phosphorus

%

I

I
f-I
f-I
I

�Table 7.
Buffalo Peaks browse - chemical partitioning of the nutritive constituents of browse samples from
five transects on Buffalo Peaks Study Area.

Season
of
Collection

Tran3ect
No.

Percent
of
Total

I

35.94
35.63

II
Summer

Cell Walls
Acid InHo1osoluble
cellulose
Ash %
%

%

%

%

Percent
of
Total

12.20

13.36

5.17

5.21

64.06

8.76

4.54

50.76

12.00

12.37

6.73

4.26

64.37

8.29

4.63

51.45

12.39

5.92

4.96

64.63

8.73

4.95

50.95

%

Lignin
%

35.37

IV

36.37

12.30

13.63

4.86

5.58

63.63

7.98

4.36

51.29

35.17

12.30

12.91

6.21

3.75

64.83

7.87

5.87

51.09

I

33.27

12.10

I
•....

7.60

'13.83

5.07

6.77

66.73

6.38

5.19

55.16

7.60

12.37

5.09

8.66

66.28

6.37

5.36

54.55

II

33.72

III

34.31

7.80

13.17

6.21

7.13

65.69

7.21

6.77

51.71

32.36

7.70

12.39

5.79

6.48

67.64

6.84

5.28

55.52

33.37

7.10

11.62

6.17

8.48

66.63

6.39

5.89

54.35

27.62

8.20

7.95

5.82

5.65

72.38

7.63

5.72

59.03

II

26.51

8.10

8.93

6.09

3.39

73.49

7.78

6.78

58.93

III

25.37

8.20

7.64

5.72

3.82

74.63

7.29

6.39

60.95

IV

26.37

8.70

7.37

8.21

2.09

73.63

7.84

5.80

59.99

27.36

8.10

8.77

7.35

3.14

72.64

8.05

6.61

57.98

IV
V
I

Winter

Soluble
Carbo

III

V

Fall

Cell Contents
Ether
Soluble
Protein
Extract
Ash

V

------.........
..........--~
..,..,...."....---.-..-.. -. -~~

..-.--~~--=--~"""~-~-~--,_~'"--......--..-,
..."-'"~""'.....
..=z.~.,...._____ -'-.

N
I

�Table 8. Buffalo Peaks browse - mineral analysis of browse samples from five transects
Study Area.

Season
of
Collection

Summer

Transect
No.

Calcium

Phosphorus

%

%

Calcium :
Phosphorus
Ratio

I

.48

.21

2.3 : 1

II

.49

.20

III

.51

IV

on Buffalo Peaks

Copper
ppm

Zinc
ppm

Selenium

%

Iron
ppm

.65

.19

29

3.8

36

.12

2.5 : 1

.62

.19

29

3.8

37

.13

.22

2.3 : 1

.63

.19

30

3.7

38

.14

.52

.21

2.5 : 1

.66

.18

28

3.8

40

.15

V

.55

.22

2.5 : 1

.67

.18

31

3.6

33

.16

I

.49

.14

3.5 : 1

.57

.18

28

3.6

30

.14

II

.48

.15

3.2 : 1

.55

.18

27

3.7

33

.14

III

.50

.13

3.8 : 1

.57

.18

24

3.8

38

.13

IV

.44

.13

3.4 : 1

.54

.19

25

3.7

37

.12

V

.46

.14

3.3 : 1

.56

.20

22

3.8

36

.l3

I

.59

.10

5.9 : 1

.46

.16

23

3.5

35

.11

II

.58

.10

5.8 : 1

.47

.15

22

3.5

33

.10

III

.55

.11

5.0 : 1

.46

.14

23

3.6

40

.09

IV

.55

.12

4.6 : 1

.48

.16

20

3.6

38

.10

V

.54

.10

5.4 : 1

.44

.15

24

3.7

36

.08

Potassium

Magnesium

%

%

,

~

,

w

Fall

Winter

�Table 9.
Trickle Mountain grass - chemical partitioning of the nutritive constituents of grass samples from
five transects on Trickle Mountain Study Area.

Season
of
Collection

Transect
No.
I
II

Summer

III
IV
V

Percent
of
Total
28.37
27.72
26.31
28.17
27.21

Cell Contents
Ether
Soluble
Protein
Extract
Ash

Soluble
Carbo

Cell t.Jalls
Acid InHolosoluble
cellulose
Ash %
%

%

%

%

%

Percent
of
Total

6.20

8.72

3.42

10.03

71.63

5.27

6.38

59.98

6.20

8.37

4.72

8.43

72.28

5.61

6.79

59.88

9.59

5.63

4.99

73.69

5.93

7.92

59.84

6.20

7.37

4.37

10.23

71.83

6.27

8.27

57.29

5.90

8.77

6.91

5.63

72.79

5.33

8.63

58.83

6.10

Lignin
%

I
•.....

.s&gt;-

I
II
Fall

Winter

III

30.33

8.20

.8.81

I

4.21

9.11

69.67

7.45

8.27

53.95

7.63

5.37

11.01

67.69

7.62

9.98

50.09

32.31

8.30

28.27

8.10

7.93

4.24

8.00

71.73

7.38

7.73

56.62

8.40

8.27

5.63

5.20

72.49

7.91

9.21

55.37

8.59

6.27

5.25

71.79

7.53

8.16

56.10

3.11

8.72

5.14

79.63

9.39

9.37

60.87

4.73

6.37

7.12

78.27

8.92

8.93

60.42

IV

27.51

V

28.21

8.10

I

20.37

II

21.72

3.50

III

23.71

4.10

5.17

8.81

5.63

76.29

9.37

10.21

56.71

IV

21.21

4.20

3.37

5.27

8.37

78.79

9.01

8.62

61.16

V

19.63

4.40

3.38

6.37

5.48

80.37

8.35

9.93

62.09

3.40

�Table 10. Trickle Mountain grass - mineral analysis of grass samples from five transects on Trickle Mountain
Study Area.

Season
of
Collection

Sunnner

Fall

Winter

Potassium

Magnesium

%

%

Iron
ppm

Copper
ppm

Zinc
ppm

Selenium

%

Calcium :
Phosphorus
Ratio

.40

.14

2.9 : 1

.51

.16

26

3.8

28

.16

II

.41

.15

2.7 : 1

.55

.16

27

3.8

29

.17

III

.42

.16

2.6 : 1

.54

.14

26

3.7

30

.15

IV

.46

.15

3.1 : 1

.51

.15

27

3.8

25

.16

V

.48

.17

2.8 : 1

.53

.17

29

3.6

27

.15

I

.49

.12

4.1 : 1

.48

.14

21

3.7

21

.11

II

.51

.13

3.9 : 1

.49

.14

22

3.7

21

.11

III

.48

.14

3.4 : 1

.51

.15

27

3.6

22

.12

IV

.50

.11

4.5 : 1

.52

.16

21

3.7

18

.14

V

.52

.12

4.3 : 1

.48

.14

22

3.5

26

.15

I

.51

.08

6.4 : 1

.42

.10

18

2.5

16

.11

II

.55

.10

5.5 : 1

.45

.11

17

2.6

17

.12

III

.58

.11

5.3 : 1

.47

~10

16

2.7

19

.10

IV

.60

.10

6.0 : 1

.46

.09

19

2.B

16

.11

V

.59

.12

4.9 : 1

.45

.11

20

2.2

22

.13

Transect
No.

Calcium
%

I

Phosphorus

%

I
I-'

VI
I

�Table 11.
Trickle Mountain browse - chemical partitioning of the nutritive constituents of browse samples
from five transects on Buffalo Peaks Study Area.

Season
of
Collection

-

Transect
No.

I

40.83

II

39.37

9.70

41.73

Summer

III
IV
V
I
II

Fall

III
IV
V

Winter

Cell Contents
Ether
Soluble
Protein
Extract
Ash

Percent
of
Total

44.27
41.36
35.37

Cell Walls
Acid InHolosoluble
cellulose
Ash %
%

%

%

%

Percent
of
Total

14.01

4.79

12.23

59.17

7.37

7.47

44.33

13.03

5.37

11.27

60.63

6.91

7.30

46.42

9.40

12.27

6.39

13.67

58.27

7.28

8.71

42.28

9.60

12.91

5.41

16.35

55.73

7.59

8.62

39.52

12.81

5.45

14.00

58.64

7.63

5.53

45.48

%

9.80

9.10

Lignin
%

8.60

12.73

5.18

8.86

64.63

7.92

8.32

48.39

8.60

11.63

5.19

9.30

65.28

8.63

8.27

48.38

35.31

8.70

12.17

6.37

8.07

64.69

8.27

8.72

47.70

34.37

8.40

12.39

4.73

8.85

65.63

8.17

7.35

50.11

34.21

8.80

11.62

5.72

8.07

65.79

8.43

8.29

49.07

6.20

9.13

5.43

2.96

76.28

7.62

5.37

63.29

9.73

5.31

3.39

75.27

7.39

6.29

61.59

10.62

6.21

4.69

72.38

7.49

8.11

56.78

6.70

8.17

5.72

5.72

73.69

8.05

7.38

58.26

6.10

8.37

6.73

4.01

74.79

7.27

6.23

61.29

34.72

I

23.72

II

24.73

6.30

27.62

6.10

III

Soluble
Carbo

IV

26.31

V

25.21

I
I-'
0\
I

�Table 12. Trickle Mountain browse - mineral analysis of browse samples from five transec:tson Trickle
Mountain Study Area.

Season
of
Collection

Sununer

Fall

Winter

Potassium

Magnesium

%

Calcium: _
Phosphorus
Ratio

%

%

.47

.19

2.5 : 1

.63

.17

II

.49

.22

2.2 : 1

.64

III

.48

.23

2.1 : 1

IV

.44

.24

V

.48

I

Iron Copper
ppm
ppm

Zinc
ppm

Selenium

31

4.7

36

.15

.18

33

4.3

35

.14

.62

.17

31

4.5

33

.15

1.8 : 1

.62

.15

28

4.7

38

.14

.22

2.2 : 1

.63

.16

27

4.8

37

.15

.55

.15

3.3 : 1

.60

.16

23

4.2

35

.14

II

.57

.16

3.6 : 1

.62

.14

23

4.2

35

.13

III

.56

.16

3.5 : 1

.59

.15

24

4.1

36

.14

IV

.55

.15

3.7 : 1

.58

.16

26

4.2

39

.12

V

.57

.17

3.4 : 1

.62

.15

25

4.4

41

.16

I

.58

.11

5.3 : 1

.51

.15

21

3.9

29

.13

II.

.59

.12

4.9 : 1

.49

.14

22

3.8

28

.13

III

.61

.11

5.5 : 1

.52

.13

27

3.5

27

.12

IV

.65

.10

6.5 : 1

.54

.14

24

3.4

29

.13

V

.67

.11

6.1 : 1

.55

.15

25

3.3

31

.12

Transect
No.

Calcium
%

I

Phosphorus

,-

%

I
I-'

-..J
I

�-18Table 13. Bighorn sheep lambs necropsied during the course of the bighorn
sheep study and used for mineral level analysis in selected tissues.
Animal
No.

Study
Area

72BHL-2

Pikes Peak

8-15-72

72BHL-3

Pikes Peak

8-17-72

72BHL-4

Pikes Peak

8-18-72

72BHL-5

Pikes Peak

8-22-72

72BHL-6

Trickle Mountain

8-29-72

72BHL-7

Trickle Mountain

8-30-72

72BHL-8

Buffalo Peaks

9-13-72

Date
Collected

73BHL-7

Remarks

Born in captivity

73BHL-11

Pikes Peak

6-05-73

73BHL-12

Pikes Peak

6-19-73

73BHL-13

Pikes Peak

6-28-73

73BHL-14

Pikes Peak

7-12-73

73BHL-15

Pikes Peak

7-24-73

73BHL-16

Pikes Peak

7-26-74

73BHL-17

Pikes Peak

8-14-74

73BHL-19

Pikes Peak

8-14-74

74BHL-10

Pikes Peak

6-25-74

74BHL-5

Pikes Peak

6-12-74

74BHL-13

Pikes Peak

7-10-74

74BHL-14

Pikes Peak

7-17-74

74BHL-4

Pikes Peak

6-11-74

74BHL-12

Pikes Peak

7-09-74

74BHL-9

Pikes Peak

6-19-74

Found dead - partially
decomposed

Found dead

�-19-

Table 14. Phosphorus content of selected tissues from bighorn sheep lambs
(% dry weight).
Lamb
No.

Hair

Liver

Spleen

Kidney

Skeletal
Muscle

Heart
Muscle

72-2

.11

.12

.13

.42

.19

.22

72-3

.10

.14

.16

.30

.17

.29

72-4

.14

.18

.20

.18

.16

.21

72-5

.08

.24

.18

.11

.19

.24

72-6

.09

.11

.27

.29

.26

.26

72-7

.18

.12

.16

.14

.10

.15

72-8

.12

.11

.36

.22

.12

.17

73-7

.08

.35

.37

.38

.35

.28

73-11

.08

.19

.41

.33

.18

.25

73-12

.08

.12

.39

.35

.19

.32

73-13

.08

.19

.24

.30

.19

.25

73-14

.08

.15

.28

.37

.20

.26

73-15

.12

.10

.29

.31

.24

.31

73-16

.13

.25

.30

.24

.20

.36

73-17

.11

.29

.40

.19

.26

.28

73-19

.11

.14

.30

.40

.28

.29

74-10

.10

.15

.20

.41

.29

.32

74-5

.09

.10

.28

.30

.21

.23

74-13

.11

.16

.21

.35

.28

.30

74-14

.08

.24

.34

.30

.25

.24

74-4

.15

.11

.32

.42

.30

.28

74-12

.15

.30

.26

.26

.30

.26

74-9

.10

.24

.35

.22

.25

.30

�-20Table 15. Calcium content of selected tissues
from bighorn shee1 lambs
(% dry weight).
Lamb
No.

Hair

Liver

Spleen

Kidney

Skeletal
Muscle

Heart
Muscle

72-2

.15

.08

.09

.11

.08

.10

72-3

.17

.08

.09

.11

.08

.08

72-4

.17

.07

.09

.09

.06

.09

72-5

.15

.09

.09

.08

.07

.09

72-6

.16

.08

.17

.10

.09

.08

72-7

.16

.07

.10

.08

.06

.08

72-8

.22

.08

.11

.10

.07

.09

73-7

.18

.12

.10

.12

.14

.11

73-11

.16

.10

.10

.10

.07

.11

73-12

.17

.10

.19

.11

.08

.11

73-13

.18

.08

.09

.10

.09

.11

73-14

.26

.08

.09

.10

.09

.12

73-15

.17

.08

.07

.10

.10

.09

73-16

.17

.09

.10

.12

.10

.10

73-17

.18

.09

.10

.13

.09

.08

73-19

.29

.09

.08

.10

.12

.09

74-10

.17

.09

.08

.09

.09

.12

74-5

.16

.08

.13

.09

.09

.09

74-13

.17

.07

.09

.10

.13

.09

74-14

.39

.10

.15

.15

.17

.11

74-4

.22

.09

.14

.11

.11

.09

74-12

.16

.10

.14

.11

.09

.10

74-9

.21

.10

.09

.09

.09

.09

�-21-

Table 16. Sodium content of selected tissue s from bighorn sheep lambs
(% dry weight).
Lamb
No.

Hair

Liver

Spleen

Kidney

Skeletal
Muscle

Heart
Muscle

72-2

.03

.28

.32

.78

.17

.36

72-3

.03

.19

.28

.64

.18

.44

72-4

.05

.19

.29

.56

.19

.30

72-5

.05

.29

.31

.30

.22

.38

72-6

.04

.19

.36

.56

.21

.28

72-7

.07

.20

.29

.59

.19

.37

72-8

.15

.38

.69

.50

.20

.31

73-7

.05

.42

.33

.80

.19

.31

73-11

.05

.32

.24

.65

.22

.36

73-12

.03

.35

.35

.59

.19

.36

73-13

.03

.25

.33

.63

.20

.39

73-14

.06

.28

.37

.73

.23

.39

73-15

.05

.28

.37

.84

.18

.44

73-16

.05

.36

.37

.99

.46

.43

73-17

.05

.47

.38

.60

.56

.37

73-19

.10

.20

.26

.60

.35

.34

74-10

.02

.28

.32

.70

.19

.42

74-5

.03

.28

.32

.61

.29

.32

74-13

.04

.39

.35

.67

.21

.44

74-14

.04

.25

.30

.66

.19

.30

74-4

.04

.20

.23

.41

.17

.26

74-12

.02

.24

.35

.49

.16

.31

74-9

.03

.22

.19

.59

.24

.33

�-22Table 17. Magnesium content of selected tissues from bighorn sheep lambs
(mg/Kg dry weight ppm).
Lamb
No.

Hair

Liver

Spleen

Kidney

Skeletal
Muscle

Heart
Muscle

72-2

470

458

504

633

716

717

72-3

380

490

591

511

775

860

72-4

373

461

592

534

760

692

72-5

299

488

495

583

771

717

72-6

332

422

593

498

786

761

72-7

284

475

510

510

788

667

72-8

342

455

577

613

770

657

73-7

327

501

562

520

1027

674

73-11

338

551

649

591

·655

635

73-12

254

. 337

634

589

632

675

73-13

325

376

624

571

794

958

73-14

356

356

683

581

820

955

73-15

299

525

702

643

844

758

73-16

299

600

675

673

689

720

73-17

307

554

670

725

643

643

73-19

390

421

578

596

638

706

74-10

365

561

549

519

883

1023

74-5

322

529

946

581

807

838

74-13

291

531

629

533

792

684

74-14

1600

511

1127

866

1419

965

74-4

348

590

784

629

901

653

74-12

383

445

786

676

721

604

74-9

436

482

606

562

704

685

�-23Table 18. Iron content of selected tissues from bighorn sheep lambs
(mg/Kg dry tissue ppm).
Lamb
No.

Hair

Liver

Spleen

Kidney

Skeletal
Muscle

Heart
Muscle

72-2

174.7

312.4

638.4

372.5

240.9

385.3

72-3

186.3

302.8

336.5

396.1

241.9

256.6

72-4

615.4

339.6

366.2

302.4

222.6

331.5

72-5

172.4

378.4

332.5

354.9

224.9

321.0

72-6

259.0

268.9

577 .2

498.0

261.9

305.6

72-7

171.6

261.6

381.2

480.4

208.2

312.7

72-8

230.6

276.5

346.1

383.2

281.3

313.9

73-7

180.0

423.8

496.6

449.6

289.9

418.6

73-11

190.5

426.3

463.2

399.8

222.7

321.3

73-12

253.7

381.6

689.4

344.7

221.3

355.2

73-13

201.2

425.5

383.7

402.7

234.0

299.4

73-14

322.5

362.3

336.2

403.2

236.3

281.6

73-15

238.8

249.5

275.0

297.6

248.6

325.9

73-16

208.1

324.1

401.5

345.1

264.3

360.2

73-17

312.3

636.7

427.3

379.2

299.1

448.3

73-19

470.9

307.5

426.6

322.2

288.6

323.8

74-10

207.0

548.6

548.6

399.6

242.1

379.4

74-5

195.5

455.7

362.8

283.7

246.6

325.8

74-13

247.3

343.3

378.7

383.9

311.2

319.0

74-14

453.6

426.0

476.9

332.6

283.8

345.7

74-4

307.3

721. 7

392.2

449.4

270.2

356.9

74-12

173.4

899.3

655.2

655.2

301.8

402.4

74-9

224.6

332.2

317.5

287.4

273.9

327.1

�-24Table 19. Copper content of selected tissues from bighorn sheep lambs
(mg/Kg dry tissue ppm).
Lamb
No.

Hair

Liver

Spleen

Kidney

Skeletal
Muscle

Heart
Muscle

23.6

5.5

13.9

9.5

17.9

72-3

85.6
81.2

9.4

5.5

8.8

9.2

20.8

72-4

92.6

13.9

5.5

14.1

6.5

19.9

72-5

63.2

16.8

4.9

14.6

6.4

19.4

72-6

82.9

69.1

10.9

13.3

7.9

20.4

72-7

76.9

42.6

6.7

15.4

7.3

20.8

72-8

75.6

48.3

8.2

15.3

6.3

21.9

73-7

60.9

9.6

6.6

15.9

5.8

14.6

73-11

69.3

66.3

7.4

13.9

10.5

15.3

73-12

82.1

62.5

13.0

13.5

8.2

23.1

73-13

69.2

63.6

5.6

16.8

9.2

22.6

73-14

122.3

13.3

6.3

16.5

8.2

21.2

73-15

90.9

17.1

5.9

16.1

7.5

22.7

73-16

84.5

16.2

6.4

13.8

10.4

20.2

73-17

89.3

36.9

6.5

21.9

9.7

22.4

73-19

105.6

8.1

4.8

12.9

12.2

19.1

74-10

79.2

31.3

4.9

10.7

9.3

21.5

74-5

74.7

64.7

5.5

13.5

7.3

23.3

74-13

80.0,

6.5

5.6

12.5

6.6

19.8

74-14

106.7

24.6

8.5

15.5

8.4

20.9

74-4

148.5

95.1

6.1

14.4

15.0

19.7

74-12

66.2

6.5

6.2

17.6

8.4

18.3

74-9

93.1

82.6

7.4

15.0

10.2

22.8

72-2

�-25-

Table 20. Zinc content of selected
(mg/Kg dry weight ppm).

tissues from bighorn

sheep lambs

Lamb
No.

Hair

Liver

Spleen

Kidney

Skeletal
Muscle

Heart
Muscle

72-2

98.1

190.9

74.9

110.0

105.9

71. 7

72-3

91. 7

156.8

88.0

88.0

107.3

79.8

72-4

102.2

166.8

113.0

112.5

128.2

71.8

72-5

158.0

134.5

120.3

112.5

131.8

72.8

72-6

108.8

92.7

132.6

80.9

121.0

81.5

72-7

105.0

103.4

107.3

80.1

118.2

67.7

72-8

107.4

102.5

204.5

88.1

119.5

78.5

73-7

111.8

303.4

112.4

100.6

169.8

72.8

73-11

121.3

203.7

89.5

76.1

91. 7

77 .6

73-12

113.8

281.3

102.5

82.0

137.5

77.7

73-13

103.8

119.3

107.9

107.0

117.0

76.5

73-14

97.3

136.2

110.2

116.2

181.2

79.3

73-15

108.2

338.0

115.6

168.9

186.4

90.9

73-16

94.3

376.8

91.2

116.5

136.2

76.9

73-17

105.6

475.2

176.2

193.8

123.4

80.3

73-19

97.4

153.8

110.1

149.0

193.7

84.6

74-10

103.5

198.5

98.5

89.9

99.7

72.2

74-5

120.7

99.2

106.5

98.0

167.2

75.6

74-13

120.0

252.6

102.7

105.8

193.3

129.1

74-14

93.4

234.3

115.3

131.5

112.7

80.4

74-4

94.8

111.5

100.2

222.5

117.3

73.2

74-12

101.5

81.1

108.3

98.0

106.9

91.5

74-9

97.9

81.0

107.1

91.4

58.7

81.8

��January 1976

-27JOB PROGRESS

State

0f

REPORT

C.::.O:::.;L::;O:..:RAD=:...:O~
_

Project No.

w-41-R-25

Work Plan No.

1

Bighorn Sheep

&amp; Mountain Goat Investigations

Job No.

17

-------------------------

Job Title __~Man~=i~p~u=l=a=t~i=o~n~o~f~V~e~g~e~t~a~t~i~on--~on=-B~ig~h~o~rn~_S~h~e~e~p~Ran==~g~e~s~
~
_
Period Covered:
Personnel:

June 1, 1974 to May 31, 1975

George D. Bear, Dave Roberts,
Lynn Stevens

Thomas McClure,

William Adrian,

and

ABSTRACT
Study plots near Cathedral, Colorado were treated with varied applications of
nitrogen and phosphorus fertilizers, and 2,4-D herbicide in 1971-72.
Two
areas were selected for study sites:
(a) an alpine range, and (b) a ponderosa
pine-bunchgrass range. Nitrogen fertilizer was applied at 0, 30, 60, and 90
lbs per acre; phosphorus fertilizer at 0 and 30 1bs per acre; and 2,4-D herbicide at 0 and 2 lbs per acre. The effects of these treatments ·are being
monitored.
There were no major changes in the plant composition from the previous year.
There was a significant increase in forage production on the ponderosa pinebunchgrass area due to the nitrogen-phosphorus-herbicide
treatment.
Alpine
plots treated with phosphorus and nitrogen-herbicide
showed significant changes
in vegetative production.
Soil samples collected last summer indicated phosphorus and nitrogen fertilizers were still prevalent on the ponderosa pinebunchgrass study area, while only minimal amounts of fertilizer still persist
in the alpine area. Protein content of the plants collected from the ponderosa
pine~bunchgrass winter range increased with increased nitrogen application
rates. The increased protein levels persisted into the mid-winter sampling
period. Vegetation calcium:phosphorus
ratios were lowered by the phosphorus
fertilizer treatments.
Differences in the protein content and ca1cium:phosphorus
ratios of alpine vegetation due to the various treatments were erratic and not
as pronounced as in vegetation on the ponderosa pine-bunchgrass area. Deer and
elk were very selective for the nitrogen treated plots.

��-29-

MANIPULATION

OF VEGETATION

ON BIGHORN SHEEP RANGES

George D. Bear

P. S. OBJECTIVE
Improve the herbage yield, vegetative density and vegetative
on selected bighorn sheep ranges in Colorado.

composition

SEGMENT OBJECTIVES
1.

Determine the effect of each treatment on the composition,
and chemical content of plant species.

2.

Determine grazing preferences
to each treatment.

of wild bighorns

production,

on the area with respect

METHODS· AND MATERIALS
The study areas are located about 35 miles south of Gunnison, Colorado.
This
bighorn herd winters in two areas: in a low (8,000 feet elevation) bunchgrass
and ponderosa pine type, and in a high (11,000-14,000 feet elevation) alpine
type. This is the fourth year of a five-year study, therefore, the procedures
outlined here are being used in a continuing study. The following work is
being conducted on each winter range.
Bighorn preference and response of the vegetation was measured on plots
treated with 2,4-D (herbicide), nitrogen, and phosphorus.
Treatments were
made in 1971 and 1972., Two levels of 2,4-D treatment ( 0 and 2 1bs/acre) were
evaluated.
Four levels of nitrogen treatment (0, 30, 60, and 90 1bs/acre),
and two levels of phosphorous treatment (0 and 30 1bs/acre) were examined.
These were evaluated in all possible combinations, or a 2x4x2 factorial for a
total of 16 treatments, as outlined in the following table:
Treatment
Ntunber

2,4-D

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16

0
0
0
0
0
0
0
0
2
2
2
2
2
2
2
2

Treatment
Nitrogen
0
0
30
30
60
60
90
90
0
0
30
30
60
60
90
90

Phosphorus
0
30
0
30
0
30
0
30
0
30
0
30
0
30
0
30

�-30-

Treatment plots, measuring 50 feet by 100 feet, have been established.
Each plot corner was marked with a metal fence post. These markers were
numbered to assist in identifying plots when bighorns are grazing on them.
Each treatment was randomly replicated 3 times on each study area. Each
replication consisted of the 16 treatment plots listed on the preceding
page. Plots in each replication were arranged in a rectangular area eight
plots long and two plots deep. Randomized selection of each plot treatment
was determined by using a table of random numbers.
1. Frequency sampling.
Transects were established on each plot to
determine changes in vegetative frequency which may result from the various
treatments.
Frequency (p) is defined p=m/n, where m is the number of quadrats
containing a given species and n is the total number of quadrats observed in
each treatment.
Four transects were established in each plot. Transects
paralleled the base of the plot with approximately lO-foot spacing between
transects.
A total of 15 quadrats are located at 6-foot intervals along each
transect, or a total of 60 quadrats for each treatment plot. Preliminary
sampling work indicated a quadrat size measuring l6x16 inches sh~uld be used
on the lower winter range at Cebolla Creek; and a split-sampling scheme should
be used on the Alpine Area. Two quadrat sizes are used on the alpine: lx1
inch for measuring Trifolium ~
and Carex sp., and 8x8 inches for all other
species.
Frequency percentages were derived for all plant species encountered on
the quadrats.
To be considered as occurring on a given quadrat, the center
of an individual plant or one-half of its basal area must be inside the quadrat.
An "individual" for bunchgrasses was a clump; for a single stem species, such
as Western wheatgrass, it was individual stems; for cactus it was a lobe; and
for shrubs and forbs it was the basal area of all stems. Obviously, these
data are acceptable for determining changes in frequencies for each individual
species only, and cannot be used to compare one species to another.
These
transects were read in June and July, when the vegetation was growing well,
but before the crowns became so large they interfered with observations.
Yield plots. Quantity of forage produced on the treatment plots was
determined by using a Neal Electronics Herbage meter.
The herbage meter
measures an area 24x12 inches x 18 inches, which is designated as yield plot.
The 60 plots established on each treatment for the frequency plots above were
also used for the yield plots.
At every twelfth meter read plot, all vegetation was clipped and weighed.
All green vegetation on these plots was clipped at the root crown and placed
in a labeled paper sack. These samples were oven dried and weighed to the
nearest gram. The weights were used to compute regression equations for
converting herbage meter readings into pounds of forage per acre.
Wire cages were placed on the study areas to monitor the effect of
grazing on the area. Cages measuring 3 feet by 3 feet by 1 foot high were
placed on each area. Vegetation produced inside the cages (no grazing) was
visually compared with vegetation produced on areas outside the cages (grazed)
areas) to get a general idea concerning effects of grazing.
Photo points were established on each treatment.
Photos were taken
at the end of the growing season to establish visual records of changes
occurring in the vegetation due to various treatments.

�-31-

Chemical analysis. Vegetation samples of the most common plant
species were collected from the treated areas. Samples were analyzed
for phosphorus, calcium, and protein content. This work was done at the
Fort Collins Research Center Laboratory.
Since the study area is a bighorn winter range, the vegetation was sampled at the completion of the
growing season, agaiu in mid-winter, and in late winter.
2. Bighorn preference or selection of treatment plots was measured
by recording number of bighorns on each treatment plot at l5-minute intervals during observation periods.
Sex and age of the sheep, time of day,
general activity of the animals, weather conditions, and snow depth (if
present) were also recorded. Observations were made with a spotting scope
during daylight hours. Sampling periods extended throughout the time the
sheep occupied the winter range. Notations were also kept on other animals
(deer or elk) that use the study plots.

RESULTS AND DISCUSSION

Monitoring

Treated Plots

Frequency Sampling
Percent frequency of occurrence for plant species on the study plots are
presented in Tables 1 and 2. Data from the three replications (180 sampling
plots) were combined to give a single percentage figure for each plant
species within each level of treatment; however, in the final analysis data
from each replication will be regarded as an individual sampling unit.
Visual inspection of the mean percentages did Ilot reveal any gross changes
in plant frequency from the previous year. Data ~~ll be analyzed (analysis
of variance) in greater detail at the completion of this study to determine
the effects of treatment over a period of several years.

Yield Plots
Correlation coefficients (r) and regression equations were computed for
determining the relationship of the herbage meter readings to herbage weights
on the clip-plots and converting meter readings to herbage production (Table 3).
Last summer was very dry as compared to the previous summer and herbage production on the control plots (0-0-0) showed a marked decrease.
Also, there
were only minor differences in herbage production on the Cebolla Creek Area
due to the various treatments; however, statistical tests indicated a significant (P&lt;.lO) increase due to the nitrogen-phosphorus-herbicide
treatment.
It was interesting to note the apparent relationship of herbage production on
the plots treated with 60 lbs and 90 lbs of nitrogen to general precipitation
patterns. Precipitation in summers 1974 and 1972 was low, and herbage production on the plots treated with 60 lbs of nitrogen was greater than production
on plots treated with 90 lbs of nitrogen.
However, in 1973, a year with greater
precipitation, production was greater on the Cebolla Creek plots treated with
90 lbs of nitrogen.
Apparently the higher level of nitrogen has a detrimental
effect on herbage production in the drier years.

�Table l. Percent frequency of occurrence for plant species on the study plots treated with nitrogen fertilizer, phosphorus fertilizer, and herbicide,
Cebolla Creek Area, 1974.

Plant Species

0-0-0

Treatment ~Nitrosen - PhosEhorus - Herbicide)
60-0-0 60-30-0 90-0-0 90-30-0 0-0-2 0-30-2 30-0-2

0-30-0

30-0-0

30-30-0

-

-

20

1
14

-

21

-

-

3
1
48
3
63

1

7

39
1
64

14
2
54
6
52

-4

64
4
72

53
1
47

-

2

2
1

-

-

-

1

3

2

2

1
1
15
1

7
1
13

3
36

24

-

1
1

-

1

30-30-2

60-0-2

60-30-2

90-0-2

90-30-2

2
15

-

8
17

17

22

Grasses
1
AsroEyron smithii
23
Boute1oua sraci1is
Bromus sp.
13
Carex sp.
Danthonia Earryi
Festuca arizonica
50
11
Koe1eria cristata
Muh1enbergia montana 65
Muh1enbersia
richardsoni
Muh1enbersia torr~~i
OryzoEsis hymenoides
2
Poa secunda
24
Sitanion hystrix
Stipa~
Unid. Grasses

-

-

26
-

-

1
32

-

-

22
-

-

9

-

-

3
13

-

-

19

-

16

-

15
2
4

1

1

1

10

17
1

3

1

5
1

41
1
53

-2

1

2

1

9

-

41
1

2

1

5

7

6

16

12

1

-

-

1
1

-

1
1

-

1

2

-

1

1

1

1

3

-

1

-

-

--

-

2

-

1

1

1

1

2

44
12
66

-1

2

1
2

50
3
73

-1
13
-

42

-

13

21

1
58
3
74

49
5
53

-

9

1
65
1
66

1
49
10
60

-

5
11

10

16

-2

7

5

-6

-

-

-

83

1
30
-

-3

-

58
1
68

3
1

-

1
15

20

-

-3

54

-

56
2
2

21
-

-

27

-

18

4

-

Forbs
Achillea 1anu1osa
Androsace sp.
Arabis cranda11ii
2
Aster rubrotinctus
Castilleja sp.
Chaenactis cious1asii
ChenoEodium sp.
ChrysoEsid vi110sa
Draba sp.
Erigeron sp,
1
1
Eriosonum sp.
Frasaria sp.
Geranium sp.
1
Gilia
Heuchera sp.
-

-2

-

-

-

-

-

-

1

1

-

-

3

1

1

--

1
1

-

-

2

3

-

-

-

-

-

2

-

4

-

-

-

-

-

-

-

1
1

-

-

2
1

-

1

---------------------------------------------------------------------------------------------------------------------------------------------------------

I

w

N
I

�Table l. Percent frequency of occurrence for plant species on the study plots treated with nitrogen fertilizer, phosphorus fertilizer, and herbicide,
Cebolla Creek Area, 1975 (continued)•

Plant Species

0-0-0

0-30-0

30-0-0

30-30-0

1
1

1

-

Treatment (Nitrogen - PhosEhorus - Herbicide~
60-0-0 60-30-0 90-0-0 90-30-0 0-0-2 0-30-2

30-0-2

30-30-2

60-0-2

60-30-2

90-0-2

90-30-2

-

-

1

1

-

1

Forbs (continued)
Hymenoxys richardsoni LaPEula sp.
LeEidium montanum
LeEtodactylon Euugens 2
Hertensia 1anceo1ata 4
Penstemon teucrioides Potentil1a concinnea 2
Solidago decumbens
3
1
Senecio mutabi1is
1
Senecio sEartioides
Sme10wskia ca1ycina
SEhaeralcea coccinea
Valeriana edulis
Unid. Forb-s-----

-

-

-

4

-3
1
7

-

-

1
2

-

z

-I
1
3

-

1
1

-7

-

1
3

1
1
1

5
-

-

1

1
-

43
1
7

40

18

40

34

3

11

3

5

1
1
1

1
-

1

-2

1

-1

18

-

-

13
1

15

-4

2

1
1

1

-

-

-

-

-

-

39

45

5

2

-

--1

-

2

-1

-

-

-

1

1
1

-

-1

-

1

-

1

1
-

-

1
3

I

1
-

1
1

1
-

-

-

1

-

-2

2

-

1

1
1

-

-

1
1

-

-

-

-

-

31

20

22

24

-

1

-

Shrv\"os
Artemisia frigida
42
Artemisia tridentata Chrysothamnus parrvi 2
ChtJ::sothamnus
viscidif10rus
1
Echinocactus sp.
Jamesii americana
1
1
Rhus trilobata
1
Ribes ~

1
-

-

1

-3

-

.-

-5

-

2

-

-

-

35

-2
1
1

-

33

1

-

26

-7
--

32

-3

1

-

-

-l
-

-

-

4

-1
1

-6

-3

I

w
W
I

�Table 2. Percent frequency of occurrence for plant species on the study plots treated with nitrogen fertilizer, phosphorus fertilizer, and herbicide,
Alpine Area, 1974.

Plant Species

0-0-0

0-30-0

30-0-0

30-30-0

4
17
2
12

4
8
4
4

4
13

8
2
44
3
54

18
1
35
2
66

Treatment {Nitrogen - PhosEhorus - Herbicide~
60-0-0 60-30-0 90-0-0 90-30-0 0-0-2 0-30-2

30-0-2

30-30-2

60-0-2

60-30-2

90-0-2

90-30-2

8
12
2
21

7
7

1
9
1
26

12
13
38

5
3
62
30
39

2
3
71
28
38

~
4
AgroEyron scribneri
8
Carex sp.
De~chamEsia caesEitosa
6
Festuca ovina
He1ictotrichon
4
mortonianum
1
Luzu1a sEicatum
44
Poa sp.
6
Trisetum sEicatum
46
Kobresia be11ardi

-

-3

3
12

-

-

12

7

-8

11
2
49
5
43

2
6
54
4
47

16
4
48
15
40

1
3

10
21

8
11

10
6

9

2
12
6
8

36

29

15

26

24
6
41
7
51

20
1
42
2
49

16
7
57
17
48

9
3
58
19
48

25
1
67
15
37

8

3
7
1

-

-

-

-

76
21
32

29
3
61
4
44

-

39
13

58
28
42

I

Forbs
Achillea 1anu1osa
Androsace
seEtentriona1is
Arenaria fend1eri
Artemisia scoEulorum
Caltha 1eEtoseEa1a
Castilleja
occidenta1is
Castilleja rhexifo1ia
Cerastium
beeringianum
Erigeron Einnatisecta
Erigeron simE1ex
Eritrichium e10ngatum
Gentiana Erostrata
Geum turbinatum
HaE10EaEEus Elgmaeus
H~eno~s
grandif10ra
Mertensia bakeri

I

w
....

2

2

-

-

6
35

-

1
9
5
6

-

18
2
2
4

9
15

1
1
3
16

1
4

-

28

-

1
12

-

1

1

-

-

-

2
3
14

1
1
66

1
6
33
1

1
16
22

2
8

-

-

1
2

12
7
26
3
1
39
1

10
16

-

-

6
25
1
5

1
13
1
1

34
1
1

17
1
1

-

-

-

7

-

7

2

-

2

-

3

1

1

-

4
1
12

2
2
6

1
11
11

-7
-

-

1
1
11

-

20
1

1
15

2
1
28

1
1

3
1
6

1
1
7

6
1
3
3

11

-4
-

22
2
8
4

4

7

17

1
3

-8
-3

5
1

-

2

8

1
3
6

1
8

1
3

-

14
12
5

4
9
3
2

3
22
1
3

1
25
1
3

19
2
6

23
1
1
2

25
3
1

26
3

7

1

9
1
2
11

-

-

-

-

-

-

7

-5
1
2
1

1
2
10

-3

-

-

--------------------------------------------------------------------------------------------------------------------------------------------------------

�Table 2. Percent frequency of occurrence
Alpine Area, 1974 (continued).

Plant Species

Forbs

for plant species

on the study plots

treated

Treatment ~Nitrogen - PhosEhorus
90-0-0
60-0-0
60-30-0
90-30-0

0-0-0

o 30-0

30-0-0

30-30-0

47

15

40

51

54

52

45

18

29
-

-

-

-

-

25
1

19

-

-

24
21

2
3
2
18

13
17
9

14
9
27

3
4
22

38
1
5
10

18
-

-

20
13

22
-

3
28

25
15

with nitrogen

- Herbicide)
0-0-2
0-30-2

fertilizer,

phosphorus

fertilizer,

and herbicide,

30-0-2

30-30-2

60-0-2

60-30-2

90 ...
0-2

90-30-2

49

42

61

74

49

-

-

14
6

17
20

5
11
2

21
4

(continued)

Oreoxis bakeri
Pedicu1a~
scoEu10rum
Phace1ia sericea
Phlox caespitosa
Podistera eastwoodae
Po1emonium viscosum
Po1~gonum bistortoides
Po1ygonum viviEarum
Potent ilIa concinna
Potent LLl.a
diversifo1ia
Potenti11a
rubricau1is
Ranunculus inamoenus
Saxifraga ~
Saxifruga chrysantha
Saxifraga flage11aris
Saxifraga rhomboidea
Sedum stenoEetalum
Silene acaulis
Sme10wskia calycina
Thalictrum alEinum
Thlaspi alEestre
Trifolium das~ph~11um
Trifolium nanum
Trifolium Earryi
Unid. Forbs

-

-

11

1
11
14

-

-

-

-

1

-

-

1

5
1

2
4

7
2

5

4

4
1

2

3
10
1

-

1
3

-

11
17
18

5
1
2
14
15

-

-

-

-

1
16
21

-

25
12

30
-

-

-

-

3
25

-

4
1
8
18
11

-

-

2
14

-

12
13
9

-

24
-

14

-

29

-

1
18
1
1
13
27

22
-

Shrubs
Salix niva1is

-

2

14
1
8
8
15

25
-

-

-

14
1
1
16
14

-

26
17

54

42

53

-

-

2
-

27
3
-

10

10

21

44
-

1
22

-

4

-1

26

14

-

-

-

-

-

-

5
1

5

3

1

-

1
-

6
3
15

-

-

3
17

2
22

9
2
3
1
19

-

-

27

-

22
21
1

-

1
26
2
1

-

12

-

-

1

-

-

2

-

-

-

-

12
10
8

3
1
9
3
5

4
1
2

11
1
3

9
1
7

1
12
6
9

22

11

11

28

1
34

6
15

21
1

17

17

13
1

-

-

-

-

I

w

V&gt;
I

�Table 3. Mean herbage production (pounds per acre) and relationship (correlation coefficient) of meter
readings to oven-dry weights of forage on the treated plots.
Cebolla Creek
Herbage
Production
(lb/A)

A1Eine

Correlation
Coefficient

Herbage
Production
(lb/A)

Correlation
Coefficient

459

.71

1733

.62

30-0-0

456

.66

1594

.70

60-0-0

581

.55

1565

.74

90-0-0

518

.75

1577

.84

0-30-0

475

.80

2489

.42

30-30-0

576

.83

2065

.50

60-30-0

547

.67

1683

.75

90-30-0

499

.89

1767

.78

0-0-2

485

.76

1112

.90

532

.83

1303

.63

60-0-2

504

.85

1784

.73

90-0-2

494

.81

1896

.68

0-30-2

480

.82

1340

.65

30-30-2

547

.84

1173

.90

60-30-2

667

.77

1620

.71

90-30-2

475

.83

1950

.75

Treatment
(N-P-H)
0-0-0

30-0-2

I

w
0\
I

�-37There were significant differences (P&lt;.lO) in vegetative production on
alpine plots treated with phosphorus and nitrogen-herbicide.
It appeared
that increase due to phosphorous treatment was less pronounced on the plots
when the herbicide was introduced to the treatment than when phosphorus
was used as a single agent or phosphorus-nitrogen was applied.
Phosphorous
fertilizer appeared to have potential for increasing vegetative production
on alpine ranges and should be investigated in greater detail.
The herbicide treatments influenced vegetative production on the alpine
plots in two ways; first they reduced forb cover and secondly potentiated
the effects of nitrogen treatments.
Nitrogen applications alone had no
significant effect on production, however, when combined with herbicide,
production generally increased with increased nitrogen levels. Apparently
the competitive effect of the forbs or presence of plants with nitrogenfixing characteristics (legumes) minimize the effects of nitrogen fertilizer
when applied at the rates tested here. Use of herbicide in the alpine as
a management tool is somewhat questionable, since it reduces ground cover
and removes plants that may be important to wildlife.
Soil samples were collected last summer and sent to the Colorado State
University Soil Testing Laboratory for analysis.
Results of these analysis
indicated phosphorous and nitrogen fertilizers were still prevelant on the
Cebolla Creek plots, while only minimal amounts of fertilizer still persist
in the Alpine area (Table 4).

Chemical Analysis
Chemical analysis (protein, phosphorus, and calcium) for the plant species
collected are presented in Tables 5-14. Data for the fall and mid-winter
sampling periods are presented here. However, chemical analysis of the
late-winter samples (April, 1975) have not been completed and detailed
statistical analysis will be undertakeri when data collection is completed.
Protein content of the plants collected from the Cebolla Creek study plots
increased with increased rates of nitrogen.
However, in many cases there
were only minor differences in protein content between plants collected from
plots treated with 60 lbs/acre of nitrogen and plots treated with 90 lbs/acre
of nitrogen.
The increased protein levels persisted into the mid-winter
sampling period.
Vegetation-calcium:phosphorus
ratios were lowered by the phosphorus treatment
on the Cebolla Creek area. This was very evident in the mid-winter samples
for each plant species. Ratios appeared to be affected to a lesser degree in
the fall-samples when the ratios were low anyway.
Differences in the protein content and calcium:phosphorus ratios of alpine
vegetation due to the various treatments were erratic and not as pronounced
as on Cebolla Creek. This is likely due to the minimal amounts of fertilizers persisting in soils on alpine plots, as indicated by the soil samples.
Each of the species collected on the alpine area in fall showed a general increase in protein with the addition of nitrogen onto th.e plots; but this trend
was not duplicated in the mid-winter collection of Kobresia bellardi.
Again,
these data will be analyzed (analysis of variance) in greater detail for the
final report while comparing data from several years.

�Table 4.

Treatment
(N-P-H)
Cebolla
0-0-0

Analysis of soil samples collected in July, 1974.

NH4-AC

N03-N
ppm

NaHOO3
P
ppm

ppm

Zn
ppm

Fe
ppm

CU

Mn

2.5

1

9

295

0.58

10.4

0.7
1.0

16.0
15.0

0.8

18.0

O.M.

pH

Condo
mn hos/cm

%

6.5

0.5

K

0-30-0
30-0-0

6.7

0.6

2.3

1

20

430

0.45

6.4

0.5

2.4

2

9

278

0.63

9.7
11.1

30-30-0

6.4

0.4

2.3

3

25

308

0.84

60-0-0

13.2

6.6

0.8

0.5

16.0

2.7

8

9

225

60-30-0

10.7

6.6

0.8

0.6

22.0

2.6

4

18

350

0.60
1.35

90-0-0

10.6

6.6

0.9

0.6

17.0

3.0

7

10

380

0.63

90-30-0

13.6

6.6

0.7

0.5

2.6

8

18

380

0.61

15.8

0.8

22.0
21.0

0-0-0

5.9

0.4

5.7

1

11

188

1.01

0-30-0

76.5

4.9

0.7

0.8

6.5

1

14

120

140.0

30-0-0

6.0

0.5

6.1

1

13

198

30-30-0

65.0

5.7

0.3
1.4

0.85
1.29

0.6

6.5

1

27

135

0.53

60-0-0

70.0

5.4

0.7

6.5
1.6

0.3

5.7

&lt;1

10

120

0.70

60-30-0

73.5

5.8

0.8

0.4

9.8

13.0

1

11

195

0.91

90-0-0

56.0

5.7

0.6

0.3

8.6

9.9

8

8

125

0.32

90-30-0

5.4

3.5

7.0

5

8

133

0.78

37.0
41.0

0.4

0.3

0.5

5.2

AlEine
8.6
11.0

I

Vol

00
I

�Table 5.

Chemical

analysis

of Festuca

arizonica

collected

on the Cebolla
Collection

Treatment
(N-P-H)

0-0-0
0-30-0

Protein
2.6
2.6

30-0-0

2.9

30-30-0

2.6

60-0-0

3.6

April 1974
Phosphorus
Calcium
.03

.17

Ca:P Ratio

Protein

5.7 : 1

9.7

Creek Study Area.

Date and Analysis (%)
September 1974
Phosphorus
Calcium
Ca:P Ratio

.18

.38

Protein

January 1975
Phosphorus
Calcium

2.1 : 1

2.7

.03

.27

9.0

.27

1. 8 : 1

.30

5.0

.05

.16

3.2 : 1

9.1

.20

.22

1.1 : 1

3.1

.15

.03

.20

6.7 : 1

13.2

.18

.22

1.2

5.3

.06

.06
.02

.22
.18

Ca:P Ratio

: 1

: 1

: 1

3.7 : 1

14.1

.20

.28

1.4 : 1

4.7

.12

.22

1.8 : 1

9.0 : 1

12.2

.16

.25

1.6

1

4.9

.06

.27

4.5

1.6

: 1

2.7

.18

.20

1.1 : 1

: 1

60-30-0

3.4

.03

.19

6.3 : 1

14.3

.18

.28

3.4

.02

.12

6.0 : 1

14.5

.16

.25

1.6 : 1

3.7

.03

.18

6.0 : 1

90-0-0

1.4 : 1

5.9

.05

.27

5.4 : 1

90-30-0

3.2

.04

.16

4.0 : 1

14.1

.18

.25

0-0-2

2.7

.05

.19

3.8 : 1

10.1

.22

.28

1.3 : 1

3.3

.07

.25

3.6 : 1

2.7

.04

.18

4.5

1

8. 7

.20

.25

1. 3 : 1

3.3

.06

.27

4.5

1

0-30-2

.20

2.2

1

I
W

30-0-2

2.7

.04

.18

4.5

1

13.2

.18

.22

1.2

: 1

2.5

.09

3.1

.04

.18

4.5

1

13.6

.18

.25

1.4 : 1

4.9

.07

.27

3.9

1

30-30-2

16.1

.18

.28

1.6 : 1

5.1

.07

.20

2.9

1

.28

1.8 : 1

4.1

.05

.23

4.6

1

.28

1.6 : 1

6.9

.07

.27

3.9

1

8.8

.08

.32

4.0

1

60-0-2
60-30-2
90-0-2
90-30-2

3.2
4.6
3~6
4.6

.03
.06

.04
.06

.12
.17
.17

.18

4.0
2.8

4.3
3.0

1
1
1

,
.L

14.1
14.3
15.5

.16

.18
.18

.28

1.6 : 1

'"

I

�Table 6.

Chemical analysis of Muhlenbergia montana collected on the Cebolla Creek Study Area.

Collection Date and Ana~i~(%)
Treatment
(N-P-H)

April 1974
Protein

September 1974

Phosphorus Calcium Ca:P Ratio

Protein

Phosphorus

January 1975

Calcium Ca:P Ratio

Protein

Phosphorus Calcium Ca:P Ratio

0-0-0

2.7

.02

.30

15.0 : 1

8.5

.16

.25

1.6 : 1

3.5

.07

0-30-0

.32

2.6

4.6 : 1

.03

.24

8.0 : 1

7.4

.16

.34

3.5

.10

.35

3.6

.03

3.5 : 1

.30

10.0 : 1

9.3

.16

.22

2.1 : 1
1.4 : 1

4.9

.09

.30

2.9

3.3 : 1

.03

.30

10.0 : 1

11.4

.16

.22

1.4 : 1

5.1

60-0-0

.18

.27

1. 5 : 1

4.3

.03

.25

8.3 : 1

9.9

.16

.22

1.4 : 1

4.9

60-30-0

.16

.27

3.9

1. 7 : 1

.03

.26

8.7 : 1

9.9

.18

.25

1.4 : 1

4.9

.10

4.3

.27

.03

2.7 : 1

.37

12.3 : 1

10.1

.14

.25

1.8 : 1

5.3

.06

.35

5.8 : 1

.25

8.3 : 1

9.5

.14

.22

1.6 : 1

6.2

.07

.39

5.6 : 1
3.2 : 1

30-0-0
30-30-0

90-0-0
90-30-0

3.6

.03

0-0-2

2.7

.05

.31

6.2 : 1

8.3

.16

.25

1.6 : 1

0-30-2

3.7

.10

2.7

.32

.04

.33

8.3 : 1

7.9

.16

.25

1.6 : 1

3.3

.06

.30

.04

.25

5.0 : 1

6.3 : 1

9.3

.16

.28

1.8 : 1

4.3

.08

.39

4.9 : 1

30-0-2

3.4

30-30-2

3.6

.03

.24

8.0 : 1

10.5

.18

.20

1.1 : 1

60-0-2

5.7

.09

3.9

.04

.35

.25

3.9 : 1

6.3 : 1

8.1

.16

.28

1.8 : 1

5.5

.15

.39

60-30-2

3.8

.05

.24

4.8

: 1

9.5

.16

.22

1.4 : 1

90-0-2

6.4

.14

5.0

.05

.27

2.6 : 1
1.9 : 1

.30

6.0 : 1

12.2

.16

.31

1.9 : 1

5.5

.05

.32

.26

6.4 : 1

4.3 : 1

10.8

.16

.28

1.8 : 1

6.8

.07

.35

5.0 : 1

90-30-2

5.3

.06

I

.I&gt;-

o
I

�Table 7.

Treatment
(N-P-H)
0-0-0
0-30-0

Chemical analysis of Boute10ua gracilis collected on the Cebolla Creek Study Area.

Protein

A2ril 1974
Phosphorus Calcium Ca:P Ratio

Collection Date and Ana1~sis (%~
Se2tember 1974
Protein Phosphoru3 Calcium Ca:P Ratio

Protein

Januar~ 1975
Phosphorus Calcium

Ca:P Ratio
6.3 : 1

6.8

.08

.43

5.4 : 1

10.5

.18

.53

2.9 : 1

5.9

.07

.44

3.9

.09

.41

4.6 : 1

8.3

.18

.47

2.1 : 1

6.9

.17

.37

2.2 : 1

.53

2.9 : 1

7.4

.08

.39

4.9 : 1

.63

2.9 : 1

9.3

.18

.48

2.7 : 1
2.4 : 1

30-0-0

4.0

.07

.48

6.9 : 1

14.7

.18

30-30-0

7.7

.08

.53

6.6 : 1

15.3

.22

60-0-0

6.3

.06

.46

7.7 : 1

13.4

.18

.50

2.8 : 1

10.1

.18

.44

60-30-0

6.0

.07

.44

6.3 : 1

14.9

.20

.50

2.5 : 1

10.1

.13

.60

4.6 : 1

90-0-0

7.3

.09

.50

5.6 : 1

13.9

.18

.53

2.9 : 1

11.5

.09

.48

5.3 : 1

90-30-0

7.2

.08

.51

6.4 : 1

14.9

.20

.63

3.2 : 1

14.7

.11

.46

4.2 : 1

6.6

.09

.37

4.1 : 1

4.3

.08

.39

4.9 : 1
4.0 : 1

.•..
•...
I
I

0-0-2

3.9

.09

.43

4.8 : 1

10.3

.20

.47

2.4 : 1

0-30-2
30-0-2

3.6

.07

.47

6.7 : 1

9.9

.18

.56

3.1 : 1

4.6

30-30-2
60-0-2

5.6
6.8

60-30-2
90-0-2
90-30-2

15.1

.20

.53

2.7 : 1

0.6

.08

.32

6.7 : 1

13.2

.18

.50

.15

.44

2.9 : 1

13.4

.18

.53

?8 : 1
2.9 : 1

8.6

7.1 : 1

7.8

.10

.39

.46

5.1 : 1

15.5

.18

.63

3.5 : 1

9.7

.35

.46

3.9 : 1
1.3 : 1

.47

5.9 : 1

14.1

.18

.47

2.6 : 1

.41

1.2 : 1

.59

10.3
11.7

.33

3.0 : 1

.19

.46

2.4 : 1

.46

5.1 : 1

.07

.47

.07

.50

3.9

.09

7.3

.08

8.5

.09

.09

.51

5.7 : 1

16.1

.20

�Table 8.

Treatment
(N-P-H)

Chemical analysis of Artemisia frigida collected on the Cebolla Creek study area.
Collection Date and AnalYSis (%)
April 1974
Protein

Phosphorus

Calcium

September 1974
Ca:P Ratio

Protein

Phosphorus Calcium

January 1975
Ca:P Ratio

Protein

Phosphorus

Calcium

Ca:P Ratio

0-0-0

9.6

.14

.67

4.8 : 1

11.0

.20

.75

3.8 : 1

0-30-0

11.4

8.2

.21

.13

.66

.81

3.1 : 1

6.2 : 1

12.8

.26

.72

2.8 : 1

7.8

.31

.62

.81

3.6 : 1

2.6 : 1

14.3

.24

.69

2.9 : 1

7.8

.13

.61

.93

4.1 : 1

7.2 : 1

14.9

.26

.78

3.0 : 1

8.6

.23

1.00

4.3 : 1

.22

.75

3.4 : 1

11.0

.22

.79

3.6 : 1

30-0-0
30-30-0
60-0-0

12.3
8.7
10.8

.17
.15
.14

.62

4.4 : 1

14.5

N
I

60-30-0

12.0

.18

.63

3.5 : 1

15.3

.24

.75

3.1 : 1

90-0-0

11.3

10.1

.14

.13

.61

.79

4.7 : 1

5.6 : 1

15.5

.20

.59

3.0 : 1

9.5

.11

.81

7.4 : 1

16.3

.22

.72

3.3 : 1

9.9

.18

.87

4.8 : 1

90-30-0

14.2

.18

.61

3.4 : 1

..,..I

�-43-

collected on the Cebolla
Table 9. Chemical analysis of Chen01!2diumsp.
Creek Study Area in September, 1974.
Ana1Isis (Percent)
Calcium
Phosphorus

Treatment
(N-P-H)

Protein

0-0-0

21.1

.48

1.00

2.1

1

0-30-0

18.8

.26

1.24

4.8

1

30-0-0

21.5

.22

.91

4.1

1

30-30-0

22.1

.42

.97

2.3

1

60-0-0

19.2

.20

1.27

6.4

1

60-30-0

21.1

.26

1.13

4.3

1

90-0-0

21.1

.20

1.27

6.4

1

90-30-0

24.6

.26

1.03

4.0

1

Ca:P Ratio

�Table 10.

Treatment
(N-P-H)

Chemical analysis of Kobresia be11ardi collected on the Alpine Study Area.

Protein

~ril 1974
Phosphorus
Calcium

Collection Date and Anal~sis ~%~
SeEtember 1974
Ca:P Ratio Protein Phosphorus Calcium Ca:P Ratio

Januan: 1975
Protein Phosphorus Calcium

Ca:P Ratio

0-0-0

5.6

.03

.67

22.3 : 1

13.4

0-30-0

.16

.53

6.0

3.3 : 1

.07

7.2

.74

.07

10.6 : 1

.81

13.6

.24

11.6 : 1

.53

2.2 : 1

9.2

.06

.93

15.5 : 1
14.0 : 1

30-0-0

7.3

.06

.64

10.7 : 1

30-30-0

13.4

.20

.59

7.7

3.0 : 1

.07

7.4

.68

.05

9.7 : 1
21. 7 : 1

.84

11.4

.16

.53

3.3 : 1

8.0

.08

.60

14.3

.18

7.5 : 1

.41

2.3 : 1

6.2

.08

.72

9.0 : 1

60-0-0

6.0

.03

.65

60-30-0

7.0

.07

.68

9.7 : 1

15.3

90-0-0

.18

.47

7.0

2.6 : 1

.09

.72

7.2

8.0 : 1

.15

.77

15.3

.16

5.1 : 1

.53

3.3 : 1

5.7

.06

.77

12.8 : 1

.47

2.4 : 1

7.6

.18

.72

4.0 : 1

90-30-0

7.2

0-0-2

6.8

0-30-2

6.8

.09

.68

7.6 : 1

.08

.71

8.9 : 1

14.9

.20

.41

2.1 : 1

.06

7.4

.70

.08

11.7 : 1

.79

16.5

.24

9.9 : 1

.47

2.0 : 1

6.6

.10

.84

8.4 : 1

16.3

.20

..,.
..,.
I

I

30-0-2

7.7

.06

.66

11.0 : 1

14.5

30-30-2

.18

.53

6.5

2.9 : 1

.05

.70

7.2

.07

14.0 : 1

.68

12.6

.18

9.7 : 1

.50

2.8 : 1

5.3

.06

.65

10.8 : 1

60-0-2

7.7

.09

.72

8.0 : 1

14.5

60-30-2

.16

.5G

7.9

3.1 : 1

.06

.70

9.5

11.7 : 1

.12

12.6

.79

.18

6.6 : 1

.50

2.8 : 1

6.9

.07

.75

10.7 : 1

90-0-2

7.7

.07

.73

10.4 : 1

14.7

90-30-2

.20

6.2

.41

2.1 : 1

.09

.71

6.8

7.9 : 1

.05

14.9

.89

.20

17.8 : 1

.41

2.1 : 1

5.5

.13

.79

6.1 : 1

�Table 11.

Treatment
(N-P-H)

Chemical analysis of Poa sp , collected on the Alpine Study Area.

Protein

Collection Date and Anallsis ~%~
Se2tember 1974
AEril 1974
Calcium
Phosphorus
Protein
Ca:P
Ratio
Phosphorus Calcium

Ca:P Ratio

10.8
11.2

.20

.22

1.1 : 1

.24

.25

1.0 : 1

13.4

.16

.31

1.9 : 1

8.5 : 1

13.6

.18

.25

1.4 : 1

.39

13.0 : 1

12.4

.20

.20

1.0 : 1

.06

.57

9.5 : 1

16.3

.24

.28

1.2 : 1

5.5

.04

.43

10.8 : 1

19.4

.26

.38

1.5 : 1

5.8

.06

.45

7.5 : 1

15.5

.18

.28

1.6 : 1

0-0-0

4.6

.04

.50

12.5 : 1

0-30-0

4.7

.07

.52

30-0-0

5.8

.04

.45

7.4 : 1
11.3 : 1

30-30-0

6.1

.06

.51

60-0-0

6.4

.03

60-30-0

7.2

90-0-0
90-30-0

I

01:"V1

I

.18

.20

1.1 : 1

7.0 : 1

10.3
11.2

.18

.22

1.2 : 1

.45

9.0 : 1

14.1

.14

.25

1.8 : 1

.07

.34

12.6

.20

.22

1.1 : 1

5.1

.05

.55

4.9 : 1
11.0 : 1

17.2

.18

.34

1.9 : 1

60-30-2

8.5

.07

.45

6.4 : 1

10.3

.22

.18

90-0-2

7.2

.06

.42

7.0 : 1

14.5

.24

.25

0.8 : 1
1.0 : 1

90-30-2

5.6

.07

.43

6.1 : 1

11.0

.24

.22

0.9 : 1

0-0-2

4.8

.03

.50

16.7 : 1

0-30-2

7.5

.07

.49

30-0-2
30-30-2

8.4

.05

5.8

60-0-2

�-46Table 12. Chemical analysis of Oreoxis ~
September, 1974.
Treatment
(N-P-H)

collected on Alpine Study Area,

Analysis (Percent)
Protein

Phosphorus

Calcium

Ca:P Ratio

0-0-0

11.4

.22

1.13

5.1

1

0-30-0

13.4

.32

1.40

4.4

1
1

30-0-0

15.1

.22

1.34

6.1

30-30-0

17.0

.28

1.37

4.9

1

60-0-0

12.0

.22

1.27

5.8

1

60-30-0

14.5

.24

1.37

5.7

1

15.7

.24

1.34

5.6 : 1

90-0-0
90-30-0

Table 13. Chemical analysis of Trifolium ~
Area, September, 1974.

collected on the Alpine Study

Analysis (Percent)

Treatment
(N-P-H)

Protein

Phosphorus

0-0-0

20.3

.24

2.80

11.7

1

0-30-0

19.2

.22

3.40

15.5

1
1

Calcium

Ca:P RatIo

30-0-0

22.1

.24

2.20

9.2

30-30-0

21.5

.26

1.90

7.3

1

60-0-0

20.5

.24

10.4

1

60-30-0

24.0

.26

2.50
1.70

6.5

1

90-0-0

20.3

.24

11.7

1

90-30-0

23.8

.26

2.80
1.90

7.3

1

Table 14. Chemical analysis of Geum turbinatum collected on the Alpine Study
Area, September, 1974.
Analysis (Percent)

Treatment
(N-P-H)

Protein

Phosphorus

Calcium

0-0-0

10.8

.26

.87

3.3

1

0-30-0

.20

1

.20

.87
1.20

4.4

30-0-0

10.5
11.4

6.0

1

30-30-0

11.8

.26

.97

3.7

1

.97
1.24

3.7

1

6.9

1

Ca:P Ratio

60-0-0

10.3

.18

60-30-0

10.1

.18

90-0-0

12.4

.18

1.67

9.3

1

.20

1.37

6.9

1

90-30-0

14.5

�-47AnimalPreferertce
Few observations were made of animals grazing on the plots (Table 15).
Bighorn use was limited to three treatments in the alpine; however, this
was likely influenced by the location of the plots rather than sheep
selectivity.
These plots were most closely located to a preferred bedding
ground.
Sheep were frequently observed grazing in the area adjacent to the
plots and exhibited no pronounced effort to move onto the plots.
Visual
reconnaissance did indicate the sheep had grazed on the nitrogen treated
plots and apparently were seeking out the more vigorous stands of Po a sp.

Table 15. Number of bighorn sheep, deer, and elk observed
study plots during the period January - May, 1975.
Treatment
(N-P-H)

Alpine
Bighorns

grazing on the

Cebolla Creek
Bighorns
Deer

Elk

2

0-0-0

3

0-30-0
30-0-0

4

30-30-0
60-0-0

1

9

11

60-30-0
1

90-0-0
90-30-0

4

0-0-2
2

6

9

1

30-30-2

4

3

60-0-2

6

0-30-2
30-0-2

2

60-30-2
90-0-2
90-30-2

10

4

2
1

�-48-

Observations on the Cebolla Creek study area were limited to deer and
elk. Five rams wintered on the area~ but were not observed on the plots.
They were seen near the plots and tracks indicated they had frequented
the plots, however, a definite pattern of use could not be detected.
As in other years~ deer and elk were very selective for the nitrogen
treated plots. This was determined by the degree of grazing or utilization of Arizona fescue~ fringed sage~ and goosefoot~ rather than directly
observing animals on the plots. General inspection of the plots during
January and February found the plants on the plots treated with 90 and
60 lbs/acre of nitrogen were receiving heavy use. The animals were still
feeding primarily at night and seeking shelter in the timbered areas soon
after daylight.
By March and April the vegetation on the plots treated
with 90 and 60 lbs/acre of nitrogen had been reduced to a stubble and use
was increasing on the plots treated with 30 Ibs/acre of nitrogen.
At this
time the animals were becoming more active during daylight hours~ increasing
the number of observations on the plots. Therefore~ the data in Table 15
does not reflect the degree of use occurring on these plots. During the
observation period from January through April the plots without nitrogen
received very light use.

Prepared by

~6.~~
George D. Bear
Wildlife Researcher

�-49JOB PROGRESS

State of

January 1976
REPORT

COLORADO
--------~~~~-----------

Project No.

W-4l-R-25

Work Plan No.

1

Job Title

Trapping, Marking,

Period Covered:
Personnel:

Bighorn Sheep &amp; Mountain
Job No.
and Collecting

Goat Investigations

21

-----------------------

Bighorn Sheep

June 1, 1974 - April 15, 1975

G. D. Bear, R. Davies, J. Ellenberger, M. P. Elkins, J. Goodyear,
C. P. Hibler, R. Imler, D. Imler, E. Latson, J. Morris, P. Neil,
W. J. Olmstead, R. L. Schmidt, J. Wegryzn.

ABSTRACT
Six lambs were collected during the period covered by this report and were
examined for lungworm infections.
Fifty-four bighorn sheep were captured, marked, and released.
Thirty-eight
were captured, marked, and released in the Cache la Poudre Canyon and 16
were captured, marked and released at the Rampart Range Gravel Pit.

��-51-

TRAPPING, MARKING,

AND COLLECTING

BIGHORN SHEEP

Robert L. Schmidt

P.S. OBJECTIVE
To trap or collect bighorn sheep for adding to the captive flock at Little
Hills Experiment Station, for obtaining nasal swabs and blood samples, for
marking and treatment with therapeutic drugs, or (in the case of dead
animal collection) for necropsy examination.

SEGMENT OBJECTIVES
1.

Collect bighorn lambs at intervals and locations prescribed
tractual agreement.

2.

Trap an indefinite number (as many as can reasonably
handled) of bighorn sheep of all age classes.

by con-

be caught and/or

METHODS AND MATERIALS
The 1974 bighorn lamb collections were made in the same manner as reported
previously by Rutherford (1974). Briefly, this involved locating a group
of ewes and lambs in an area from which collected specimens were desired.
Then a lamb was selected for collection by observing them unnoticed from a
distance.
Lambs were selected for collection on the basis of outward
physical appearance.
Attempts were made to select lambs of contrasting
physical condition, ranging from healthy, robust appearing lambs to "scruffy"
looking, smaller lambs in apparent poor condition.
Next the shooter
approached the ewe lamb group on foot taking advantage of terrain to avoid
detection.
Lambs were shot with a 22 magnum caliber rifle usually from
distances of 100 yards or less. In all cases lambs were shot in the neck
to minimize bullet damage to tissues of interest.
Trapping techniques have been reported previously (Erickson 1970, Schmidt
1975) and involved baiting a trap site with apple mash, then capturing bighorn sheep with a remotely triggered drop net.

RESULTS AND DISCUSSION

Lamb Collections
Bighorn lambs were collected from the Pikes Peak area from June 12 to July
17, 1974. Collection dates for the six collected lambs were: 6-11-74 ram
lamb, 6-12-74 ram lamb, 6-19-74 ewe lamb, 6-25-74 ewe lamb, 7-10-74 ewe
lamb, and 7-17-74 ram lamb. In addition a ram lamb was found dead on 7-9-74
and was estimated to have died approximately 3 to 7 hours before discovery.
Detailed necropsy reports of post-mortem examinations of these lambs are
reported by Hibler (1976).

�-52-

Trapping and Marking
Bighorn sheep were trapped at two locations during the period covered
by this report. The first capture occurred in the Cache la Poudre River
drainage.
Twenty-one bighorn sheep were captured, and marked at a bait
site near the Poudre Ponds Fish Hatchery and 17 were captured at a bait
site in the Big Meadows area near Kinikinik (Table 1). Twenty-five of
these bighorns (15 from Poudre Ponds site and 10 from Big Meadows site)
were transplanted to a release-site near Big Narrows (Bear and Schoonveld
1976). The remaining sheep were released at their respective capture
sites.
The second capture occurred at the Rampart Range Gravel Pit near Colorado
Springs on April 4, 1975. Sixteen sheep ~ere captured at this location
and were collared and released (Table 2). These sheep previously had been
treated with chemotherapeutic drugs delivered in apple mash bait. They were
collared to assess the effects of drug treatment.

LITERATURE

CITED

Bear, G. D., and G. G. Schoonveld.
1976. Extending bighorn sheep ranges.
Colo. Div.Wildl.
Game Res. Rep. January.
(In press).
Erickson, J. A. 1970. Use of drop net and collars in a study of Dall
sheep. North Wild Sheep Council, Trans. pp. 20-21.
Hibler, C. P. 1976. Investigation of spontaneous diseases of bighorn
sheep. Colo. Div. Wildl. Game Res. Rep. January.
(In press).
Rutherford, W. H. 1974. Collection of bighorn lambs.
Game Res. Rept. January.
p. 108-110.
Schmidt, R. L. (Ms. in prep.).
in Colorado.

Sr. Conservation

Procedures

Aide

Colo. Div. Wildl.

for trapping bighorn sheep

�-53Table l. Sex, age and tagging information of bighorn sheep captured
Cache la Poudre River drainage, January 21, 1975.

Estimated
Age
Sex ~/

Ewe

4-1/2 yrs.

Ewe
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe
Ram
Ewe
Ewe
Ram
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe

5
5
6
5
3
6
2
6
4
2-1/2
1-1/2
Lamb
Lamb
Lamb
4
3
2-1/2
8
4
5
4

Ewe
Ewe
Ewe
Ewe
Ewe
Ram
Ram
Ram
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe
Ewe

5
2-1/2
3
2
3
7
1
1-1/2
Lamb
Lamb
Lamb
6
3-4
3
3
3

~/

in the

-Capture
Location

Neck Collar
Color

Neck
Collar No.

Ear
Tags

Remarks

Poudre Ponds

Blue

1

None

Transplanted
Big Narrows

"
"

2
4
5
6
7
9
10
11
12
13
18

"
"
"
"

"
"

"
"

II

"

"
"
"
"

"
"
"
"

"
"

"
"
"
"

"
"

"

"
"

"

"
"
"
"
"
"

"
"
"

"

Rig Meadows

"
"
"
"
"
"

"

"

"

"
"
"
"
"

"
"
"
"
"

"
"
"

"

"

II

"

"

"

"

"
"
"
"
"

"
"

"
"

No collar
No collar
No collar
Red

"

"
"
"
"
Blue

,,"
"
"
"

"

"
"

No collar
No collar
No collar
Yellow

1
3
4
5
7
8
14
15
16
17
19
20
No /I
No /I
No /I

i

"

2

"

II

"

"

3
4
5

"

"

"
"

"

to

"
"
"
"
"
"
"
"

"
"

"

Blue 111
Blue 112
Blue 113
None

"

"
"

"

"
"

"

"
"

"
"

"

"
"
"

"
"
"
"
"

Orange 113
None

"

Blue /14
Orange III
Orange 112
None

Transplanted
Big Narrows

"

"
"
"
"
"

"
"
"

"
"
"

"

"

"

"
"
"

"
"

"
"

Three previously collared sheep were recaptured: 1 - red/black stripe and 1
red/white 112 at Poudre Ponds and 1 white/red 112 at Big Meadows.

to

�-54-

Table 2. Sex, age, and tagging information
Rampart Range Gravel Pit, April 4, 1975.

Sex

Estimated
Age

Ram

1-1/2

White/green

Ewe

4-1/2

Ewe

Neck Collar
Color

of bighorn sheep captured at

Neck Collar
No.

Weight

Body
Temperature

4

118 lbs.

102.0

Yellow

6

149

102.2

6-1/2

Yellow

7

149

103.6

Ewe

6-1/2

Yellow

8

136

103.2

Ewe

4-1/2

Yellow

9

Not taken

Not taken

Ewe

4-1/2

Yellow

11

127

101.8

Ewe

4-1/2

Yellow

13

110

104.6

Ewe

3-1/2

Yellow

14

147

103.0

Ewe

7-1/2

Red

1

127

103.0

Ewe

1-1/2

Red

2

110

104.4

Ram

Lamb

No collar

Orange 114

70

104.0

Ram

Lamb

No collar

Orange 115

68

105.6

Ewe

Lamb

No collar

Orange #6

62

104.4

Ram

Lamb

No collar

Orange 117

48

104.8

Ram

Lamb

No collar

Orange 118

64

104.0

Ewe

Lamb

No collar

Orange #9

69

105.5

stripe

Eartags

�January 1976

-55JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No.

W-4l-R-25

Work Plan No.

1

Bighorn Sheep
Job No.

Job Title

Monitoring

Period Covered:

June 1, 1974 - May 31, 1975

Personnel:

&amp; Mountain Goat Investigations
22

---------------------------

Bighorn Sheep Populations

M. P. Elkins, C. P. Hibler, J. Morris, P. Neil, R. L. Schmidt,
J. Teeter.

ABSTRACT
Bighorn ewes on Pikes Peak which were treated with chemotherapeutic drugs
had nearly 14 times greater lamb survival than ewes which were not treated.
Treatment with cambendazole in combination with diethylcarbamazine apparently
was most beneficial for lamb survival of the drugs tested. Tramisol with
diethylcarbamazine was least beneficial.
The Pikes Peak bighorn sheep population was estimated at 163 animals of
which 27 were adult rams, 83 were adult ewes, 10 were yearlings and 43
were lambs, as of April 15, 1975.

��-57-

MONITORING

BIGHORN SHEEP POPULATIONS
Robert L. Schmidt

P. S. OBJECTIVE
To monitor bighorn sheep herds by visual surveillance
the characteristics of lamb mortality.

to further elucidate

SEGMENT OBJECTIVES
1.

Monitor bighorn sheep numbers, age and sex composition,
appearance, behavior and activity.

general body

2.

Monitor numbers, distribution, association with other sheep, and lamb
births and survival, of bighorn sheep which have been trapped, treated
with therapeutic drugs, neck collared, and released at the trap site.

METHODS AND MATERIALS
All of the information collected concerning numbers, age and sex composition,
and lamb survival of bighorn sheep populations in general was gathered on
populations which were being monitored for evaluation of drug treatment
effects. These results will be presented along with results of drug treatment evaluations.
In addition a simple Lincoln Index was used to estimate
size of the current Pikes Peak bighorn sheep population.

Drug Treatment Evaluation

Pikes Peak, 1975 Treatments
The objective of drug treatments during the 1974-75 segment was to treat
Pikes Peak bighorns with drugs administered in apple mash bait without
capturing the animals. The goal was to deliver 8.1 cc of cambendazole and
3200 mg of diethylcarbamazine to each adult utilizing bait stations. Drugs
were mixed with apple mash bait on five dates, March 14, April 10, April 12,
April 16, and April 17. Seven drug delivery stations were selected within
winter habitats frequented by various segments of the Pikes Peak bighorn
population (Fig. 1). Baiting with apple mash without drugs was begun the
first week in January. Bait was put out twice daily and observations were
kept on the amount of bait put out, the number of sheep using each bait site
after each bait placement, and the amount of bait consumed. These data were
used to compute the average daily consumption per individual bighorn per day.
From these data it was estimated that each adult sheep consumed about 5
pounds of apple mash per day for those days they visited the bait station.
Consequently, on days when drugs were delivered to sheep in the apple mash
sufficient cambendazole and diethylcarbamazine were added so that each 5
pounds of apple mash contained 8.1 cc of cambendazole and 3200 mg of
diethylcarbamazine.

�Manitou Springs
&amp;

Dome Rock

.0.

Sentinel
'0 Point

I
VI
00
I

o
• = Bait Site

Fig. 1.

Location of drug trea~ment bait stations. ,1975

�-59-

Evaluation of the effects of drug treatment will be made in two ways.
First, ratios of lambs per 100 ewes will be calculated from population
classification data gathered from June, 1975 to mid-February, 1976.
These ratios will be compared to ratios of lambs per 100 ewes in years
before drug treatment was begun. Secondly, 71 percent of the known
collared ewes were treated and 29 percent were not treated so r&amp;tios of
lambs per 100 ewes will also be compared this way. Fecal pellets will
be collected from collared sheep which were known to have ingested drug
treated apple mash. These will be analyzed for numbers of lungworm
larvae per gram of fecal material and compared to similar samples from
collared bighorn sheep which did not ingest drug treated apple mash bait.

Pikes Peak, 1974 Treatments
In 1974 during the period February-April, 52 bighorn ewes were treated
with various drug combinations (Gill 1975). These drug treatments were
evaluated during the 1974-75 segment by observing and recording lamb
natality and survival of collared treated ewes vs lamb natality and
survival of collared and uncollared untreated ewes. Natality and survival
of lambs was estimated by pairing each lamb with its dam every time population
composition classifications were conducted.
These pairings were done by
observing from a distance groups of bighorn sheep containing ewes and lambs
until every lamb in the group had nursed a ewe. It was assumed that each
ewe produced only one lamb and that a ewe would nurse only her own lamb.
This pairing process worked well through December. After that time it was
necessary at times to pair on the basis of maternal-filial behavioral
characteristics such as clos~ association during flight, periods of alarm,
or periods of foraging.

Rampart Range, 1975 Treatments
One bait station was established at the Rampart Range Gravel Pit during
the first week in January, 1975 (Fig. 1). Untreated ba.it was placed at the
bait site 5 days each week up to one week prior to drug delivery. At this
time bait was delivered for all seven days prior to treatment.
On February
11 the drug dichlorvos was placed in the bait according to procedures outlined for the 1975 Pikes Peak treatments. The Rampart treatment was designed to deliver 25 mg/lb. of body wt. to each sheep using the bait site
on drug delivery day. Untreated bait was then placed at the site 5 days a
week until February 24 and then daily up to March 5 when the bait was again
treated with dichlorvos. This process was repeated up to April 3 when
dichlorvos was again added to the bait. The intent was to treat all Rampart
bighorn sheep on all 3 drug treatment periods. On April 4 all sheep visiting
the bait station were trapped and all adults were marked with numbered colored
neck collars and lambs with numbered colored ear tags (Schmidt 1976). Evaluation of drug treatment efficacy was to be based upon lamb natality and survival and fecal larval loads.

Rampart Range, 1974 Treatment~
The 1974 Rampart Range drug treatment experiments were the first attempts
at delivering drugs to unrestrained bighorn sheep via apple mash bait.

�-60-

Procedures were the same as those employed in the Pikes Peak, 1975 and
Rampart Range, 1975 treatments except that 8.1 cc of cambendazole was the
intended dosage for each ~dult sheep (lamb dosages were two-third adult
rates). Treatment dates were January 31, February 28, and April 1, 1974.
None of the sheep receiving drugs were marked so the evaluation of drug
treatment was based upon natality and survival of young. In all of the
drug treatment trials where drugs were delivered in apple mash bait, the
intent was to treat ewes in early-, mid-, and late·-gestation.

Pikes Peak Bighorn Sheep Population

Estimates,

1974-75

Composited observational data on the proportion of collared vs uncollared
bighorn sheep observed over the period May 20, 1974 to March 6, 1975 were
used to estimate the size of the Pikes Peak bighorn sheep population.
A
simple Lincoln Index was used to project these data to population estimates.
Since all sheep which were collared were adults population projections were
pertinent only to adults. Proportions of yearlings and lambs in classification data were used to project to final estimates of total populations.

RESULTS AND DISCUSSION

Drug Treatment

Evaluation

Pikes Peak, 1975 Treatments
Data to evaluate the 1975 drug treatments will be gathered during the 197576 segment. Forty-five sheep were treated at all 7 bait stations during the
March 14 treatment date, 25 on April 10, 2 on April 12, 23 on April 16, and
23 on April 17.
.

Pikes Peak, 1974 Treatments
When lamb natality and survival data of all treated ewes are combined, regardless of drug type, and compared to natality and survival of lambs from
untreated ewes it appears that both natality and survival of lambs from
treated ewes is superior (Table I). Though it is not obvious from the
summaries in Table 1, the major period of lamb mortality occurred sometime
between July and August, 1974, and probably was well underway by the first
week in July. Ratios of lambs:lOO ewes remained high (Ca 70:l00) throughout
the September, 1974 - April, 1975 period for those ewes receiving drug
treatments.
Ewes not receiving drugs experienced extremely low lamb survival for the same period (5 lambs:lOO ewes).
It has been suggested that the high survival rates experienced by lambs from
drug treatment could have resulted from the high quality nutrient supplement
provided by the apple mash bait and not necessarily from drug treatment.
This is not likely since all ewes had access to bait sites prior to treatment.
Drugs were delivered either by injection or were force-fed orally to sheep
treated in 1974. So differences are likely due to the effects of drug treatment.

�-61Table 1. Comparisons of 1974-75 Pikes Peak bighorn sheep lamb survival
from ewes treated with drugs in 1974 and ewes not treated (treated ewes
include all drug treatments).
No.
No.
Untreated
Ewes 1:../ Lambs

Period

Lambs:
100
Ewes

No.
Treated
Ewes

No.
Lambs

Lambs:
100
Ewes

June-Aug. 1974

53

38

72 :100

41

40

98:100

Sept.-Dec. 1974

80

4

5:100

67

46

69:100

Jan.-April 15, 1975

122

6

5:100

93

62

67:100

1:../
Some of the ewes listed as untreated were treated in 1973 but did not receive treatment in 1974.

Treatment with cambendazole and diethylcarbamazine appeared to result in the
highest lamb survival rates though sample sizes were small (Table 2).
Thiabendazole and diethylcarbamazine and dichlorvos treatments appeared to be
nearly as beneficial.
Treatment with tramisol + diethylcarbamazine was least
efficacious.
Since diethylcarbamazine was not administered singly it is
possible that much of the beneficial results of the tramisol, cambendazole
and thiabendazole treatments could have been potentiated by the diethylcarbamazine.

Table 2. Summary of 1974-75 Pikes Peak bighorn sheep lamb survival comparing
various drug treatments.

No.
Ewes Treated

Drug
Tramisol 1:../

No. of Ewes
Individually Identified
At Least Once
After Sept. 1974

Lambs Positively
Identified With a
Treated Ewe From
Oct.1974-Mar.1975

Percent
Lamb
Survival

9

8

4

50

Cambendazole

1:../

13

7

7

100

Thiabendazole

11

10

6

5

83

20

14

12

86

Dichlorvos

Treatment contained diethylcarbamazine.

�-62-

Rampart Range, 1975 Treatments
These treatments will be evaluated in the 1975-76 segment.

Rampart Range, 1974 Treatments
Since none of the Rampart Range sheep were marked prior to or after
the 1974 drug treatment, there was some question whether subsequent
pairings of lambs and ewes were made on treated sheep. Because of
this, no good evaluation can be made of the results of the 1974 drug
treatments.

Pikes Peak Bighorn Sheep Population

Estimates,

1974-75

During the period of May 1974 to April 1975, 332 observations of collared
bighorn ewes and 10 observations of collared bighorn rams were recorded.
During the same period 282 observations of uncollared ewes and 35 observations of uncollared rams were recorded. Forty-five collared ewes
and 6 collared rams were known to be alive during the same period. Using
a simple Lincoln Index, the adult ewe population was estimated by the
formula:
45
X = 83 adult ewes.
332
332 + 282
The formula for estimating the adult ram population was:
6
X
X = 27 adult rams.
10 10 + 35
Thus the adult bigho.rn sheep population was estimated at 100 animals.
By using population classification data projections can be made for total
numbers of lambs and yearlings in the population as ·well. The overall lamb:
ewe ratio for the period Sept. 15, 1974 to April 15, 1975 was 33 lambs per
ewe. MUltiplying this figure by 124 ewes, as that was the 1974 ewe population,
the estimate of 41 lambs was derived. The yearling population estimate was
based on the April 15, 1974 population of 11 lambs assuming that mortality
in this age class was small from April 15 on, the yearling population estimate was 10 animals. Therefore, the Pikes Peak bighorn sheep popUlation
was estimated to contain 27 adult rams, 83 adult ewes, 10 yearlings and 41
lambs for a total of 163 bighorn sheep.

LITERATURE

CITED

Gill, R. B. 1975. Monitoring bighorn sheep popul.at Lons ,
Wildl., Game Res. Rep. January.
(In Press).

Prepared by

Colo. Div.

�-63January,

1976

JOB PROGRESS REPORT

State of

COLORADO
------~~~~---------

Project

No.

Bighorn Sheep

W-4l-R-25
1

Work Plan No.

&amp; Mountain Goat Investigations

23
------~--~----------Job Title Control of Snails and/or Lungworm Larvae on Bighorn Sheep Ranges
Period Covered:

Personnel:

Job No.

June 1, 1974 through May 31,- 1975

Gene G. Schoonveld

ABSTRACT
No work was accomplished in this segment due to a re-assignment of the
Principal Investigator Gene Schoonveld from the Research Section to
Management Section.
Study has now been assigned to Harold Shepherd and a detailed plan is
being developed to initiate the study in the present segment, and will
be reported on, at the end of the 1975-76 segment.

��.-65-

January 1976

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~----------Bighorn Sheep &amp; Mountain Goat Investigations

W-41-R-25

Project No.
Work Plan No.

1

Job No.

Job Title

Extending Bighorn Sheep Ranges

Period Covered:

June 1, 1974 to May 31, 1975

Personnel:

24

---------------------------

Gene Schoonveld, Paul Neil, Robert Keiss, Robert Schmidt, R. Bruce
Gill, and George D. Bear

ABSTRACT
Twenty-five bighorn sheep (17 ewes, 4 rams, and 4 lambs) were trapped in
the upper portion of Cache la Poudre Canyon in northcentral Colorado and
transplanted to a presumably suitable area lower in the canyon which was
void of sheep. This work is being done to evaluate the feasibility of
using such transplants to reduce concentrations of bighorns and extend
their distribution.
All sheep were marked with numbered collars and
eartags to assist in identifying individuals.
The reactions of displaced sheep were extremely variable.
Sixteen were
remaining at the release-site.
Four of them had traveled back up the
canyon, mingled with other sheep, then later returned to the release-site.
Others immediately returned to areas where they had been trapped.

��-67-

EXTENDING BIGHORN SHEEP RANGES
George D. Bear
Gene G. Schoonveld

P. S. OBJECTIVE
Determine if the distribution
be increased by transplantulg
present concentrations.

and population of bighorn sheep herds can
sheep into suitable areas adjacent to the

SEGMENT OBJECTIVES
1.

Trap and transplant

sheep in Cache la Poudre Canyon.

2.

Monitor survival, movements,
planted sheep.

distribution,

and reproduction

of trans-

METHODS· AND MATERIALS
In December, 1946, 16 bighorn sheep were removed from the Tarryall Mountains
and introduced into Cache la Poudre Canyon in northcentral Colorado.
This
herd has increased and now exceeds 100 animals. However, this herd occupies
only a limited portion of the suitable bighorn range in the canyon. This
study is exploring the feasibility of extending sheep distribution by transplanting animals into the lower portion of the canyon. The transplant-site
was located in Big Narrows, which is approximately seven airline miles below the normal range of the Poudre herd.
Bighorn sheep were trapped on bait-stations established near Poudre Ponds
Fish Hatchery and Kinikinik.
The bait-stations were established approximately
six weeks prior to trapping and regularly supplied with apple-mash, alfalfa,
and salt blocks. By mid January bighorn sheep were making daily visitations
to these stations; then drop-net traps were setup.
Forty-one bighorns were trapped on January 21; twenty-five (17 ewes, 4 rams,
and 4 lambs) were marked with collars or eartags and transported to the Big
Narrows area. Adult and yearling animals were fitted with collars made
from 4-inch wide canvas webbing with an outer layer of blue vinyl sewn on
the webbing.
A black numeral was painted on each collar so individual animals
could be identified (Table 1). Lambs were marked with blue 2-inch Richie eartags with black numerals painted on them.
The bighorns were transported from trap-site to release-site in a 2~-ton truck
with a box enclosed to accommodate the entire group of sheep. At the releasesite the truck was backed into the hillside and the tailgate opened, permitting
the sheep to run out; , They left the truck in two major groups with several
stragglers.
The single animals started rejoining larger groups within a few
hours after release.

�Table 1. Sex and age of bighorns marked with blue collars or eartags, then transplanted to the lower
Cache la Poudre River area.
Collar
No.

Sex

Age
(Years)

Collar
No.

Sex

Age
(Years)

Eartag
No.

Sex

Age

1

Ewe

4

14

Ewe

4

1

Ewe

Lamb

2

Ewe

5

15

Ewe

5

2

Ewe

Lamb

4

Ewe

5

16

Ewe

2

3

Ram

Lamb

5

Ewe

6

17

Ewe

3

4

Ewe

Lamb

6

Ewe

5

18

Ram

1

7

Ewe

3

19

Ewe

2

9

Ewe

6

20

Ewe

3

10

Ewe

2

None

Ram

7

11

Ewe

6

None

Ram

1

12

Ewe

4

None

Ram

1

13

Ewe

2

I
0\

ex&gt;
I

�-69-

The sheep were allowed a one week adjustment period before attempts were
made to relocate them. Observers traveled the main road in Cache la
Poudre Canyon and hiked up side canyons looking for collared sheep.
Whenever an individual was seen tne following information was recorded:
date, location, and collar number; the location was a!so plotted on a
map.
To simplify data analysis, the location of the sheep was expressed in
"miles from the release-site"; section lines on the topography maps were
used for this determination.
Since all observations were made within 1/2
mile of the Cache la Poudre River north-south variations were disregarded,
thus data were recorded only as miles east or west of the release-site.
The release-site was designated as 0 miles, while mil~s west (upstream)
were positive numbers and miles east (downstream) were negative numbers.

RESULTS AND DISCUSSION
The sheep moved around a great deal during the first thirty days after
their release.
Some individuals remained together, while others moved
between groups. Eighty days after the transplant it appeared that sixteen
sheep had settled in the vicinity of the Big Narrows release-site (Table 2).
Two of the transplanted ewes had traveled up the canyon, mingled with other
bighorns; then after 20 days (following release) started back down the canyon
and joined with the Big Narrows group. Also, the two yearling rams followed
a similar movement pattern in a 10-15 day period. Two other ewes remained
at the release-site for more than 60 days then traveled upstream 13 miles
and joined with another band of sheep.
Three ewes moved upstream 22 miles within 37 days after the release.
This
put them above the upper trap-site, where two of these ewes had been captured.
The urge to migrate back upstream must have been quite prominent, since they
had to pass by many other bands of sheep during the migration and did not
stay with any of these groups. There were no lambs transplanted with the
group taken from the up~er trap-site, thus, we speculate the ewes were trying
to return to their lambs. However, there were four other adult ewes transplanted from the same trap-site and they remained at Big Narrows.
Two of
these traveled half-way back up the canyon, then returned to the Big Narrows
release-site.
Twenty-three of the twenty-five bighorns have been reobserved since their release. A lamb and large ram have not been seen since the transplant.
The reactions of these displaced bighorns were extremely variable.
Observations will be continued during the next work segment to further evaluate the
effects of this transplant.

Prepared by __

~__ "~~_-_b_'
_'~_~
__ ' --=:=c.-_
GeorgEP'D.'""
Bear
Wildlife Researcher

�Numbers in
Table 2. Location of transplanted bighorns on days following release at Sheep Mountain.
table represent miles from release site; positive numbers being miles upstream and negative numbers
being miles downstream.
Day of
Observation
9
10

1

2

4

1

-

-

-

-

5

6

Collar Number
7

9

10

11

12

13

1
1

-

-

-

1

-

1

1

-

2

-

2

2

2

2

1

1

-

1

1

1

1

1

1

-

-

1

1

1

1

1

2

-

-

2

2

1

2

1

1

1

-

1

1

1

1

1

1

1

-

1

1

1

1

1

1

1

1

1

-

-

1

-

-

-1

-1

-1

-

-1

-1

-1

-1

-1

-1

-1

-

-1

-1

-1

-1

-1

-1

-1

-

-1

-1

-1

11
11

12

-

1

1

12

13

-

-

-

-

13

15

-

2

17

1

1

20

1

1

21

1

2

-

23

-

-

3

30

1

1

31

1

1

-

37

-

-

22

43

-

1

-

44

-

22

50

-

53

-

-

56

-1

-1

-

57

-1

60

-1

-1
-1

78

-

-

-

22

-

-

79

1

1

22

1

80

-

-

-

-

-

24

42

51

22
22
22

----------------------------------------------------------------------------------------------------------

I
....•
0
I

�Table 2. Location of transplanted bighorns on days following release at Sheep Mountain. Numbers in
table represent miles from release site; positive numbers being miles upstream and negative numbers
being miles downs t ream (continued).
Collar Number
18

19

20

Rams

Lambs**

-

-

7
7

1

-

7
7

-

-

-

9
7

1
1
1*

2
1

-

5

-

-

Day of
Observation

14

15

16

17

9

6

6

10

-

-

6

8
6

9
5

-

-

-

-

-

7
7

15
17
20
21
23
24
30
31
37
42
43
44
50
51
53
56
57
60
78
79
80

--

-

5

-

-

-

1
1

-

-

-

-

1

-

-

-

-

-

-

9

-

-

-

5

-

1

-

0
1
0

..

-

-

-

1

22

22

-

-

-

11

-

-

-

-

12

21

-

13

-

-

22
22

1

-

-

11

-

-

*Both unnumbered yearling rams were observed at the same location.
**
Maximum number of lambs seen at the same time was three.

1
1

-

-

22

-

-

-

1

1

-

0*
0*
0*

-1
-1
-1

22
22

1*

2

-

-

-

-

1
I

"
•.....
I

22
22
22

-

1

��January, 1976

-73JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Proj ect No.

w-4l-R-25

Work Plan No.

1

Job Title

Controlled

Job No.

&amp; Mountain Goat Investigations
25

---------------------------

Burning of Bighorn Sheep Ranges

--------------------------------------------------

Period Covered:
Personnel:

Bighorn Sheep

June 1, 1974 through May 31, 1975

Gene G. Schoonveld

ABSTRACT
No work was accomplished in this segment due to a re-assignment of the
Principal Investigator Gene Schoonveld from the Research Section to
Management Section.
Study has now been assigned to Harold Shepherd and a detailed plan is
being developed to initiate the study in the present segment, and will
be reported on at the end of the 1975-76 segment.

��January,

-75JOB PROGRESS

State of

1976

REPORT

COLORADO
------~~~~---------

Project No.

W-4l-R-25

Work Plan No.

1

Job Title

Investigation

Period Covered:
Personnel:

Bighorn Sheep &amp; Mountain
Job No.
of Spontaneous

CONTRACT

Goat Investigations
(26)

------------~~-------

Diseases of Bighorn Sheep

June I, 1974 - May 31, 1975

Robert Schmidt and Paul Neil, Colorado Division. of Wildlife;
C. P. Hibler, T. Spraker, R. Davies, J. DeMartini, Ed Latson,
J. Ellenberger, J. Wegryzn, L. Clum and J. Hurlock, Colorado
State University.

ABSTRACT
Seven bighorn sheep lambs were collected on Pikes Peak in June and early
July, 1974 to examine for early evidence of predisposition to verminous
pneumonia by·Protostrongylus.
Details are given in the text. Eighteen
bighorn sheep trapped in South Dakota durtllg January 1974 died during a
lO-day period in August as a result of stress induced Muellerius - Pasteurella
pneumonia.
Transplacental transmission was experimentally reproduced using
mouflon/bighorn hybrid sheep. The site of somatic storage of third-stage
Protostrongylus was determined to be in the lungs. Molluscacides and
larvicides experimentally evaluated failed to show any efficacy against
snails and/or larvae of Protostrongylus.
Bacterial isolations from various
sheep, including those trapped in Poudre Canyon and on Rampart Range are
detailed in the text.

��-77SPONTANEOUS DISEASES OF BIGHORN SHEEP
C. P. Hibler

P. S. OBJECTIVE
To determine causes, nature and effects of spontaneous diseases in
Colorado bighorn sheep with particular reference to mortality in lambs,
and to evaluate various means of promoting lamb crop survival (controlling
these diseases).

SEGMENT OBJECTIVE
1.

Obtain 4 to 6 lambs from the Pikes Peak herd during June for a detailed post-mortem examination to evaluate specific neo-natal pathologic processes suspected to play an integral part in the inception
of chronic lamb pneumonia (lambs will be collected under Work Plan
1, Job 21).

METHODS AND MATERIALS
Lamb collections the last three years indicate the major pathologic processes leading to pneumonia are initiated shortly after birth of the lamb.
Histopathology, virology, bacteriology and parasitology studies conducted
on these animals strongly suggest that earlier collections would yield
information which would help delineate the initiation and progression of
the pathologic process resulting in almost total loss of lamb crops.
Therefore, all collections are 3chedu~ed for the month of June. As in
1973, a field research team will conduct the post~ortem examinations.
Blood samples will be collected for clinical pathology studies, serum for
bacteriological, virological, clinical pathology, physiological baseline
data, and trace mineral evaluation. Selected tissues will be taken for
bacterial and viral isolation, and trace mineral evaluation. A complete
post-mortem for histopathology studies will be performed. The remainder
of the carcass (liver, lungs, muscles and lymph nodes) will be further
examined for additional evidence of parasitism by Protostrongylus.

RESULTS AND DISCUSSION
Results are presented in case·history form for each lamb collected and
examined. A tabular summary (Table 1) precedes the detailed results.

�Table 1.

Summary of lamb examinations,

1974.

---_.

-

-

-

---

First Stage
Lungworm Larvae
in Feces

Pasteurella
Nasal
Trachea
Lung

Sheep

Date Examined

Sex

Age

Wt II

Pneumonia

74BHL-4*

6-11-74

Male

3 weeks

20

None

-

-

-

Very few

74BHL-5*

6-12-74

Hale

4 weeks

2.8

Yes

-

-

-

Very few

74BHL-9",

6-19-74

Female

5 wee ks

21

Yes

-

+

+

Very few

74BHL-10*

6-25-74

Female

5 weeks

28

Yes

+

+

+

Very few

74BHL-12&gt;'&lt;*

7-9-74

Male

6 weeks

23

Yes

+

+

+

74BHL-13**

7-10-74

Female

6 weeks

36

Yes

+

+

-

74BHL-14

7-17-74

}!a1e

7 weeks'

34

Yes

-

+

+

,,&lt;

**

Too young for patent infections.
Pneumonia primarily bronchopneumonia.

,

Not examined
?
&gt;

1,000/g

I
--..J
00
I

�-79-

74BHL-4

6-11-74

History: This male lamb was found 6-11-74 about 7:30 AM west of
Bottomless Pits, in the lambing ground east of the toll road. The
lamb appeared healthy and was in a group with 6 other lambs that were
being "baby-sit" by two ewes. There were nine additional ewes nearby.
The lamb was killed at 8:15 AM.
Postmortem:

'

Began postmortem at 9:30 AM.
Body temperature - 102.8oF.
Body weight - 20 pounds.
Age estimated to be 3-4 weeks.
Integument and Subcutis: The hair coat is smooth. There is a small amount
of fat in the subcutis and around the peripheral lymph nodes.
Cardiovascular System: A small amount of fat around the coronary bond.
The heart is normal in size, shape and color.
Lymphohemopoietic

System:

Spleen--Normal in size, shape, color and consistency. Lymph nodules
are not enlarged. The typical red foci located around the peripheral of
the spleen are present. The parenchyma contracts from the capsule when
the organ is cut.
Bone marrow--Red, dull and granular, very active.
Retropharyngeal lymph nodes--Small,
architecture on cut surface.

1.5 x 1 cm, moist with normal

Prescapular lymph nodes--Non-reactive, bilobed, left 3.5 x 1.5 cm,
right, 2 x 1 cm; moderate fat around their cut surfaces; moist with
normal architecture.
Mediastinal lymph nodes--Small,
normal architecture.

6 x 2.5 cm; cut surface, moist with

Thymus--Normal in size and measured 15 x 2 cm.
third up the neck to the base of the heart.

It extended from one-

Digestive System: Rumen filled with grasses; small amount of milk curd in
abomasum; formed feces; liver normal - there are several white foci on and
through the liver, about 10 to 15 in number.
Respiratory System: Both lungs collapse but not completely; thus they may
contain slightly more fluid than a normal lung. Many bright red to dark
red foci (0.5 to 2 ~) are diffusely scattered under the pleura of both
lobes; some are deep in the lung parenchyma. There are also several small
pale foci that have a pale, white caseated core surrounded by a clear halo.
The bronchi, bronchioles and nasal turbinates are unremarkable.

�-80-

Musculoskeletal

System:

Bones--Thick,

Unremarkable.

difficult

to break; snap when broken .•

Urogenital System: Kidneys normal, small amount of fat over the capsule
of the kidney. Both testes have descended.
Endocrine
CNS:

System:

Thyroids - unremarkable;

adrenals

- normal, 1 x 0.75 cm.

Unremarkable.

Remarks:
Hemorrhage

74BHL-5

in the lungs probably due to migrating

lungworms.

6-12-74

History:
This male lamb was killed at 12:45 AM on 6-12-74 just above the
cabin on the beaver ponds in East Beaver Creek. This lamb was in with
a herd of 12 ewes, 8 lambs, 1 yearling ram and 1 yearling ewe. The lamb
was estimated to be 3.5-4.5 weeks of age. The body temperature at death
was 103.20F.
Postmortem:
PMI - 1:15 hours.
Weight - 28 lbs.
Integument and Subcutis:
The hair coat is smooth.
amount of fat in the subcutis.

There is a moderate

Cardiovascular System: The heart is normal in size, shape and color.
small amount of fat is present in the coronary groove.
Lymphohemopoietic

A

Syst em,

Spleen--Normal in size, shape, color, texture - the parenchyma contracts
(on cut surface) from the capsule. Lymph nodules are not prominent.
Lymph Nodes:
Retropharyngeal

lymph nodes--Destroyed

Pre scapular lymph nodes--Bilobed;
architecture; 3 x I em in size.

normal in size, shape, texture and

Mediastinal lymph nodes--Unremarkable;
normal architecture.
Mesenteric

by shot (but small) 1 x .75 mm.

small; cut surface moist with

lymph nodes--Normal.

Thymus--Normal;

extended from base of heart to one-third up trachea.

�-81-

Respiratory System: The lungs do not collapse well, especially the
diaphragmatic lobes. The lung is "speckled" with red to brown foci
0.5 to 1 mm in diameter. These focal hemorrhages are more numerous
in the posterior dorsal aspect of the lungs. They resemble very early
lungworm nodules. The bronchi are clean and the trachea is normal.
Digestive System: There is a small amount of curd in the abomasum;
abomasum about half the size of the rumen; rumen is filled with ingestion; formed feces in the colon.
Liver--Normal in size, shape, color and texture. There are 2
white foci (1 mm in size) just under the liver capsule.
Musculoskeletal

System:

Normal.

Urogenital System: Unremarkable;
Both testes descended.
Endocrine System:

small amount of fat around both kidneys.

Adrenals normal: 1 x 0.75 em.

Thyroid - unremarkable.

eNS: Shot in the brain; eyes unremarkable and the spinal cord is unremarkable.
Summary:
Numerous brown and red foci in lungs that appear to be early lungworm nodules.

�-82-

/~
Hemorrhage

(red-brown foci)

q

Area of atelectesis

J~ e..~ ----

74EHL-5
Areas from which tissue samples were collected.

~

�-8374BHL-9

6-19-]4,

History: Thi~ ew~ lamb was found east of the Bottomless Pit and east
of the toll ruad, The lamb was with 7 ewes. The lamb was shot but was
not found for 30 minutes. Body temperature was 102.4°F and weight was
21 pounds; the lamb was estimated to be 5 to 6 weeks old.
Postmortem:
Killed about 9;00 AM.
about 2:00 PM.

Postmortem started about 11:30 AM and finished

Integument and Subcutis: Hair coat normal, but not slick.
amount of fat was present in the subcutis.

A moderate

Cardiovascular System: Heart normal in size, shape and color; blood
vessels - unremarkable.
Lymphohemopoietic System: Spleen normal in size, shape, and color and
texture; the pulp contracts from the capsule when cut.
Retropharyngeal lymph nodes--Small
and size; moist surface.

(1.75 x I em); normal structure

Prescapular lymph nodes--Small

(unremarkable); 2 x 1 cm in size, bilobed.

Mediastinal

(unremarkable); 3.5 x 1 em in size.

lyu,ph nodes--Small

Mesenteric lymph nodes--Small

(unremarkable).

Prefemoral and Popliteal lymph nodes--Normal
Thymus--Extended

(unremarkable).

from heart base to one-thi.rd up the trachea.

Bone marrow--Red and active.
Digestive System: Rumen twice the size of the abomasum; only several small
flecks of milk curd present; formed feces in colon.
Liver--Normal in size, shape, color and texture; there are 3 small (lmm)
white foci just under the capsule of the liver.
Respiratory System: Nasal tract normal; trachea clear, normal. The lungs
do not collapse. Red to brown foci are scattered diffusely over the lungs.
Lungworm nodules are present in the posterior aspect of the diaphragmatic
lobe. The area is soft, and white-grey speckled; the cut surface bulges
and is encircled with reddened peripheral area. Bronchi are unremarkable.
Musculoskeletal System: Massive hemorrhage in muscles due to falling and
rolling about 100 yards from where it was shot. Femur 16 cm long, normal.
Urogenital System:
tract normal.

Small amount of fat around both kidneys, reproduction

�-84Endocrine
eNS:

System:

Adrenal glands normal, thyroi.d gland normal.

Eyes normal; cerebellum

with massive hemorrhage

from gunshot.

Summary:
Red-~rown foci in the lungs and lungworm nodules in the dorsal
diaphragmatic lobes.

�-85-

~~----------------~~~~
c~--------~

74BHL-9
Areas from which tissue samples

were collected.

�-86-

74BHL-IO

6-25-74

History:
This ewe lamb was killed at 7:30 PM 6-25-74 in the Bottomless
Pit. The lamb was with about 20 lambs and 20 ewes. All lambs appeared
to be in good condition and were large and growing well. The lamb was
shot in the base of the brain and fell 50 to 60 feet and rolled 100 yards.
Postmortem:
The lamb weighed 28 pounds and body temperature was 102.4°F (the body laid
in a snow bank about 30 minutes before temperature was taken). Lamb was
shot at 7:30 PM; necropsy started at 10:45 PM and ended at 12:30 AM.
Integument and Subcutis:
The hair coat is normal and smooth. A moderate
amount of fat is present around the prescapular and prefemoral lymph nodes.
Cardiovascular Systen: The heart is normal in size, shape and color. A
moderate amount of hemorrhage is present on the epicardium (due to fall).
Blood vessels unremarkable.
Lymphohemopoietic

System:

Retropharyngeal lymph nodes--Small,
x 1 cm in size.

moist and normal in structure,

1.75

Right prescapular

lymph node--Normal,

small, moist; size 2.5 x 1.25 cm.

Left prescapular

lymph nJde--Normal,

small, moist; ~ize 3.5 x 1.5 cm.

Thymus--Normal,
heart.

extended about 1/2 up the trachea and to the base of the

Mediastinal lymph node--Small, normal size, and structure with slightly
moist cut surface. Size - 6.5 x 1.5 cm.
Mesenteric

lymph nodes--Normal

size and texture.

Digestive System: The liver is ruptured and totally macerated from the fall;
part (25%) of the liver is located in the thoracic cavity because the
diaphragm ruptured.
Stomach and Intestines: Milk curd (small amount) is present in the abomasum;
formed feces in the colon.
Respiratory System:
The lungs partially collapse. They are filled with
blood, probably due to the fall. Lungworm nodules are present in the
posterior dorsal aspect of both diaphragmatic lobes. They are encircled
with a zone of colored tissue and the centers of the nodules are grey-white
speckled, elevated on the pleural surface, and bulge on cut surface. The
bronchi are clean and have no mucus. A slight amount of white froth is present in the nasal cavity and trachea, but not in the bronchi.

�-87-

Musculoskeletal System:
There are massive hemorrhages in the muscle
mass due to the fall. Ribs on this lamb seemed to bend and not snap
as well as the other lambs; joints and tendons are normal.
Urogenital System: Both kidneys are normal in size, shape, color and
texture.
A moderate amount of fat occurs around both kidneys.
Endocrine System: Thyroid g1and normal
Adrenal glands small, normal.

(mostly destroyed

CNS: Eyes - badly bruised;
completely destroyed.

brain - hemorrhage

Summary:
lobes.

are prominent

Lungworm

nodules

by the shot).

under dura; cerebellum

in both dorsal diaphragmatic

�--

-88-

74BHL-IO
Areas from which tissue samples were collected.

�-89-

74BHL-12

7-9-74

History:
This male lamb was found dead at 8:00 AM on 7-9-74 north of
the gravel pit in Bottomless Pit on Pikes Peak. The lamb was lying
on his left side and his mother was still with him.
Postmortem:
PMI - 8-10 hours, lamb died early that morning.
started at 11:45 AM; sex - male; weight - 23 pounds.

Necropsy

Body Condition:
The lamb has a slightly roughened, light-colored hair
coat. No fat is present in the subcutis except around the peripheral
lymph nodes.
Cardiovascular System: The heart is normal in size, shape, color and
texture. There is a slightly excessive volume of pericardial fluid, and
several very small, white flecks (fibrin) floating in the fluid. There
is a normal amount of fat in the coronary band.
Lymphohemopoietic System: The spleen is normal in size, shape, color
and texture. The retropharyngeal lymph nodes are moderately enlarged
and moist (2 x 1 cm).
Prescapular

lymph nodes--About

Mediastinal

lymphnodes--Enlarged

normal size and moist

(size: 3 x 1 cm).

and wet (size: 0.5 x 1.5 cm).

Remaining nodes are unremarkable.
The bone marrow is red and active.
The thymus is normal in size, shape and color, it extends 8 cm up from
the thoracic inlet.
Digestive System: The rumen is filled with ingesta and a moderate amount
of milk curd is present in the abomasum.
Formed feces are present in the
colon. The liver is normal in color, size, shape and texture. There are
several (.25 to .5 mm) white foci just under the capsule.
There are
several flecks of yellowish friable material in the peritoneal cavity;
they look like small flecks of fibrin.
Respiratory System: The lungs do not collapse when the thorax is opened.
About 33 percent of the lungs (involving all lobes) are slightly depressed,
red and firm. There is only a very slight excess of serosanguinous fluid
in the thoracic cavity. No fibrin is present on the pleura and no adhesions
are present. The cut surface of the red firm areas are moist and have a
fine nodular appearance (thought to be distended lobules).
A watery, white
mucoid material can be expressed from cut bronchioles.
There is no noticeable
lungworm nodules in the posterior dorsal aspect of the diaphragmatic lobes.
A white exudate is present in the nasal turbinates, trachea and etymoid
turbinates.
The mucosa in the nasal turbinates, trachea, and bronchi and
bronchioles is reddish colored.
Musculoskeletal System: The muscles and bones are unremarkable.
The bones
have their normal thickness; the ribs may be somewhat soft, but have a slight
snap when broken. Joints have several very small flecks of fibrin floating
in the synovial fluid.

�-90-

Urogenital System: There is a very small amount of fat between the
peritoneum and the capsule of both kidneys. Both testicles have
descended.
Endocrine System:
Nervous System:
also normal.

The thyroid, adrenals and pituitary are unremarkable.
The eyes are slightly dried but normal.

The brain is

Diagnosis:
1) Suppurative bronchopneumonia, 2) mild acute fibrinous
peritonitis, 3) mild acute fibrinous arthritis.

�-91-

Black (medial side)

s
Medial

Red,
firm,
Underside, no lesions
but emphysematous
Septa of the cardiac lobes are red and slightly distended.

---------...:~
No indication of lungworms

74BHL-12
Areas from which tissue samples were collected.

�-92-

74BHL-13

7-10-74

History: This lamb was found at 7:30 AM, July 10, 1974, on Bar Trail.
She was with 14 other lambs and about 22 ewes. Two lambs were noted
to experience severe coughing, had dull hair coats and lagged slightly
behind the band. This lamb was thought to be a large healthy lamb prior
to collection. The lamb was shot while lying down so no running or
excitement was incurred before death.
Postmortem: Killed 9:45, necropsy started 11:20.
temperature was 1030F and weight was 36 pounds.

Sex - female, body

Body Condition: The lamb has a dull hair coat but it is not roughened.
The lamb is large, especially for a female. There is a small amount of
fat in the subcutis around the peripheral lymph ~odes.
Cardiovascular System: The heart is normal in size, shape, color and
texture. No excessive fluid is present in the pericardial sac. There is
a small amount of fat in the coronary band around the heart. Vessels are
unremarkable.
Lymphohemopoietic System: The spleen is slightly larger than most spleens.
There appears to be a prominence of the germinal centers of the spleen,
causing a slightly "cobble-stoned" appearance of the cut surface. The red
foci, which are normally present along the periphery of the organ are
present. The retropharyngeal lymph nodes are slightly enlarged and wet,
(size: right, 1.75 x 1.5 cm; left, 2 x 1.75 cm). The prescapular lymph
nodes also are slightly enlarged and wet (size: right, 3 x 1.75 cm; left,
3.25 xl.75 cm). Both are bilobed. The mediastinal lymph nodes are enlarged and wet. They have an exaggerated appearance to the cortex; size
10 x 2 cm. The mesenteric lymph nodes are moist but have normal structure
and size. Prefemoral, popliteal and ileal lymph nodes are normal. The
thymus is large in this lamb, extending from the base of the heart of the
posterior aspect of the thyroid glands. Diameter is about 2 cm at the
thoracic inlet. Bone marrow is red and active.
Digestive System: The rumen is filled with green grass and the abomasum
also is filled with grass. A moderate amount of milk curd is also present.
The mesenterics have no fat. Formed feces are present in the colon. The
entire digestive tract is normal.
Respiratory System: The nasal mucosa is reddened and there is a small
amount of white tenacious exudate covering the surface. The ethmoid turbinates are filled with a white mucoid exudate. Frontal sinuses are normal.
The trachea is reddened and has a small amount of white mucoid exudate on
the surface. The lungs do not collapse when the thorax is opened (even after
20 minutes). The lower 75 percent of the right apical lobe, the entire
cardiac lobe, and lower 50 percent of the right diaphragmatic lobe have
contracted from the non-collapsed, dull pink lung. These areas are red
(dark reddish-brown) and firm. The pleura covering this area is not covered
with fibrin but is moist and shiny. The cut surface has a "knobby" or
"cobble-stone" appearance due to many small nodules (size: .5 to .75 mm in
diameter). There is a small amount of pearly white, thick mucoid exudate that

�-93-

can be expressed from small cut bronchioles.
The mucosa of the bronchi
is reddened and also has a moderate amount of this white, mucoid exudate.
Musculoskeletal System: The bones are normal in size and shape. There
is a normal thickness of cortex to the femur. The femus is 16-1/2 em in
length. The ribs bend, then have a sharp snap when they break. The
joints are clean and contain no fibrin. The muscles are unremarkable.
The lamb was shot just below the right ear.
Urogenital System: Both kidneys are normal in size, shape, color and
texture. There is a very small amount of fat under the peritoneal covering
of the kidneys. The ovaries are large on this lamb (1 em in diameter) and
have many small (1-2 mm diameter) cysts through both ovaries.
Endocrine System: The thyroid, adrenal and pituitary are normal in size,
shape, color and texture.
Nervous System: The brain and eyes are normal. There is considerable
blood just under the leptomeninges of the cord and cerebellum.
Diagnosis:

Suppurative acute pneumonia

(broncho pneumonia).

�· .

8

1
inside

\

L90l_..---J

Several
grey foci with white
center and translucent
ring.
~reamy

02~ ~~ --

_

Medial
consolidated

74BHL-13
Areas from which tissue samples were collected.

�-95-

74BHL-14

7-17-74

History: This lamb was found in the Bottomless Pits, just below the
stone wall. He was with several ewes and lambs. The lamb had not run,
nor was he excited in any way before death. The lamb was killed at
9:45 AM and shot in the cerebellum and C-l. Body temperature was 104.4°F
and weight was 34 pounds.
Postmortem:

PMI was 1 hour.

Integument and Subcutis: The lamb has a slightly dull and rough hair
coat. There is a small amount of fat in the subcutis around the peripheral lymph nodes, and eyes and sternum.
Cardiovascular System: The heart is normal in size, shape, color and
texture. No excessive amount of pericardial fluid is present in the
pericardial sac. A small amount of fat is present around the coronary
band.
Lymphohemopoietic System: The retropharyngeal lymph nodes are moist and
slightly enlarged (2 x 1.5 cm). The prescapular nodes are similar in
size. The right is 3.5 x 1.5 cm and is bilobed; the left is 3.25 x 1.5 cm
and is monolobed. The mesenteric lymph nodes are slightly enlarged and
moist, and the cortex bulges on cut surface (size: 10.5 x 1.75 x 1.5 cm).
The spleen is normal in size, shape, color and texture. The thymus is of
acceptable size, shape, color and texture. The thymus extends from the
base of the heart 7.5 cm beyond the thoracic inlet.
Digestive System: The rumen is filled with ingesta. The abomasum is also
filled with ingesta and milk curd. A large volume of milk curd is present
in the abomasum - about 100 g. The alimentary tract has no gross lesions;
no excessive fluid or fibrin is present in the peritoneal cavity. Formed
feces are present in the colon and rectum. No lesions were present on the
liver.
Respiratory System: The nasal mucosa is reddened and has a small amount of
white exudate covering the mucosa. The trachea is normal in coloration but
has a small amount of blood (due to shot) and white exudate covering the
ventral aspect of the trachea. The lungs do not collapse when the thorax
is opened. There is no excess fluid in the thoracic cavity. The lower 80
percent of the right apical lobe, the entire cardiac lobe, the lower 20 percent of the right diaphragmatic lobe and 50 percent of the intermediate lobes
of the lung are dull grey, firm and have a knobby cut surface. A small
amount of pearly white, mucoid exudate can be expressed from the cut
bronchioles. The bronchi and main stem bronchi have a white mucosa and a
small amount of white mucoid material in the lumen. The left lung is
similarly involved. The right diaphragmatic lobe has a white, firm triangular area located in the distal dorsal posterior aspect. This area is
white, firm, bulges 011 cut surface and a thick whitish mucoid exudate can
be expressed from bronchioles in this nodule. The left side is similar except
it has two of those nodules, on the medial side of the left cardiac lobe.
A 4 mm wh Lt.e , firm nodule (abscess) is present. Also, the most anterior
aspect of the right apical lobe has a small, early adhesion to the costal
pleura. The most remarkable total lung changes are: 1) ventral consolidation,

�-962) non-collapse

of all normal-colored

lung, 3) lungworm nodules.

Musculoskeletal System: The muscles and joints are unremarkable.
ribs have a slight bend, then a normal snap.

The

Urogenital System: Both kidneys have a small amount of fat under the
peritoneal covering. The size, shape, color and texture of both kidneys
are normal. The bladder is empty and its mucosa is unremarkable.
Both
testicles have descended and are normal.
Endocrine
eNS:

System:

Thyroids and adrenals are unremarkable.

Eyes and brain are unremarkable.

Summary:

Severe verminous pneumonia.

�-97-

:-

.;2L/.

c...nA-

----------_~

Many fresh
hemorrhages
are scattered
throughout the
collapsed pale
parenchyma

There are many small foci
that appear to the alveoli filled with
air-scattered diffusely in the
consolidated areas, of both lungs~

02..\ ~

~

G
A - white circular
4 mm nodule (abcess)

74BHL-14
Areas from which tissue samples were collected.

�-98-

In addition to the lamb collections on Pikes Peak in June, 1974, an
epizootic among the South Dakota captive bighorn sheep herd occurred
during August, 1974. The circumstances leading to mortality, and a
brief resume of the cause is given.

History and Clinical

Signs

Utilizing a drop net, personnel from the Colorado Division of Wildlife,
in cooperation with the South Dakota Game and Fish Department, captured
24 bighorn sheep (Ovis canadensis) from Custer State Park, South Dakota,
January, 1974. Th~ animals were transported to Fort Collins, Colorado
via an enclosed flat bed truck and released into three pens surrounded by
a 2.5 m (8 foot) chain link fence.
Six of the animals died shortly after arrival in Fort Collins due to a
degenerative muscle syndrome (capture myopathy).
Most of these animals
suffered ruptured gastrocnemious muscles and died in spite of treatment.
The remaining sheep appeared healthy and were fed free choice alfalfa hay
and dairy ration in addition to the grasses and forbs growing in the pens.
Salt and mineral blocks were also provided and the animals were observed
at least once a day. This herd was to provide lambs to become the nucleus
of a "tame" herd of bighorn sheep which would be utilized for lungworm
(Protostrongylus spp.) experiments, bacterial and viral diseases and
nutritional studies.
Several lambs were born during the spring of 1974,
of which three survived.
One ewe was found dead on 26 August 1974, and, at that time others were
noticed sick, with signs of dyspnea and open-mouth breathing, rapid,
shallow respiration and weakness.
Two more ewes died the next day. On
the following day, the remaining sheep were trapped under a drop net,
clinically examined, and injected with -5 ml of Oxytetracycline and 2 m1
of polyvalent past~urella bacterin.
Body temperatures were recorded
and blood samples were collected at that time. One lamb died the following
day and only one ewe died during the following week. The remaining animals,
including a ram, 2 lambs, and 12 ewes died during the next 10 days.

Summary
An epizootic of Muellerius capillaris-associated
bacterial pneumonia in 20
captive Rocky Mountain bighorn sheep (Ovis canadensis canadensis) was the
cause of the mortality.
In all adult sheep there were heavy infections with
~. capillaris.
First stage larvae were widely disseminated throughout the
lungs, including a granulomatous pneumonia.
Bacteria of several species
(Tables 2, 3, 3a, 3b) usually localized in bronchioles in area of larval
infection and were considered secondary invaders, although this important
role in death of the animals cannot be disputed.
Relative neutrophlia lymphopenia, and elevation of serum fibrinogen levels
were found in blood samples taken from several sheep 7-14 days prior to death.

�Table 2.

Bacteriological

summary of Pikes Peak bighorn lamb collections,

1974.

§
'M
H
0)

~

U
Cd

.c
0)

~

H
0

'Cl

~

~I~

...-I 0
04-1
U'M
...-I
• 0

u ~

~IU

1.

Nasal

1,2,4
5,6,7

1,2,3
4,5,6
7

2.

Trachea

2

3.

Tracheal
bifurcation

4.

Lung parenchyma
(normal)

5.

Lung parenchyma
(pneumonic)

3

PfJl

Cd fJl

I'~I

Cd
...-I
...-I
0)
H
0)

::r=

Cd
...-I
...-I
0)
'M
fJl

Cd
'M
H
0)
fJl
fJl

.c

0)
...-I
~

P-

~

.

PfJl

.

...-I U
0) 'M
H ~
::l
0)...-1
~ 0

~~

~I ~ m

~Or-!

p..,,.c::

Cd ::l
p..,

H :I
p.., &gt;

fJl

2,5,6

1,2,3

1,3,4

3,4,5

4,5,6

-

.~
::l H
0) Cd

~~
fJl...-l

O) PZ
fJl

3

1,4,5
6

3,5,6

2

3,4,5

4,6,7

1,2,4
5

4,5,7

5

5

3,4,5

4,5,6

5

4,5

5

Cd
...-I
...-I Cd
0)'Cl
H'M
::l U
0) 0

a

~

o

fJl
0

fJl
::l
U
U
0
U
0

::l
H
O)!
ctI

~
~Cd
~
tf.)

a 'M
~
.g~~
::l
0)
fJl
p..,

Peripheral

7.

Spleen

blood

u
0

~0)
H
~
tf.)

1,2,3
4,5,6
7

1,2,3
4,5,7

4,5,6

1,2,3
4,5,6
7

1,2,3
4,5,7

4,5,6

1,2,3 1,2,3
4,5,6-, 4,5,6
7
7

1,6

4,5

4,5

4,5,6

3,4,5
6

4,5

4,5,7

4,5,6

1,2,3
4,5,6
7

1,2,3
4,5,6
7

1,5

1,3,5

3,4,5
6

3,4,5
7

2,4,5
7

2,4,5
6,7

1,2,3
4,5,6
7

1,2,3
~,5,6

~,5,7

2,3,4
5,6,7

5,7

~,5

2,3,6

5,7

-

6.

fJl
::l
U
U
0

I

\0
\0

I

i
I

�Table 2.

Bacteriological

summary of Pikes Peak bighorn

lamb collections,

1974 (continued).

S

::I

'r-l

~
&lt;1.1

+J
()

t\l (/)
.0
&lt;1.1
&lt;1.1 C
C QJ

»00
~ 0
o »
Uo..
8.

Mediastinal
lymph node

9.

Retropharyngeal
lymph node

10.

Main stem
bronchus

.

'"c

0..

t\l

(/)

e

'r-ll 0
..-!
04-&lt;
()

'r-l

..-!
• 0

~Iu

(/)

I'~I

t\l
..-!
..-!
QJ

~
QJ
::c:

::I

t\l
..-!
..-!
QJ

'r-l

(/)

..0
QJ
..-! 0..
~
Ul

~
&lt;1.1
(/)
(/)

'r-l

QJ

0..
Ul

.

0..

Z (/)

4

4,6

t\l
..-!
..-!
&lt;1.1

!I ,
.

t\l
'r'i

~
::I
QJ

+J

Ul
t\l
Po&lt;

5

3,4,6

~
..c:

t\l
..-!
..-! t\l
&lt;1.1",
~

'r-l

::I ()
QJ 0

+J+J

(/)..-!
t\l ::I
Po&lt; S

o
()

(/)

t\l

t\l

I:l (/)

o

0

::I ~
QJ t\l

s

O..-!
~ ::I
Po&lt;

::I
QJ ~
Ul QJ
Po&lt; t\l

.~'"

.j..J

&gt;

C

'r-l

~

Ul

::I

o

0

()

()

0

0

()

0

~
~t\l

?

+J

.j..J

UJ

UJ

QJ

~

5,6

5,6

3,4,5
6,7

2,4,5
6,7

3,6
7

3

4,5,6
7

2,3,4
5,6,7

3,4,5
6,7

2,3,4
5,6,7

I
I-'

o
o

Key:
1.
2.
3.
4.
5.
6.
7.

(/)

I

74BHL-4
74BHL-5
74BHL-9
74BHL-lO
74BHL-l2
75BHL-13
75BB.L-l4

Bacillus sp. was found in nasal swabs of all lambs.

�-101Table 3. Bacteriological summary of cultures from captive South
Dakota bighorn epizootic.

I jI
!II

'"CI

..-l

p.
(IJ

0

~
..-l

,..
Q)

+J

o

!II

,.a

'8

.

(IJ.

'~l~
!II

u ~

r-I 0
O~
tJ .~
r-I
• 0
[%:II c.J

2,4

2,6

Q)

F.

,..
0

Iii

p.
.rJl
!II

r-I
r-I

,..
Q)

Q)

::I::

.

P.

.

P.

(IJ

(IJ
!II

r-I
r-I
Q)

..-l
(IJ

,.a
Q)

r-I

:&gt;&lt;:

.~
,..
!II

Q)

rJl

rJl
.~
z
Q)

I!I
~

~

r-I

.t::

~

!II

!II

r-I
r-I

Q)

,..

rJl

;j
Q)

;j
Q)

;j
Q)

+J

+J

+J

(IJ
!II

rJl
III

2,3
7

2,7

3

2

2

7.

Spleen

2

2,7

8.

Med iastina1
lymph node

2

9.

Peritoneum

10.

Pericardium

11.

Mammary
tissue

Code:

1.
2.
3.

74BHS-28
74BHS-29
74BHS-30

Po&lt;

Po&lt;

3

2

2

4.
5.
6.

74BHS-31
74BHS-32
74BHS-34

2?

7.

Q)

7

Peripheral
blood

3

,..

2,3, 3,7
7

6.

2

3

+J

1

Left lung
parenchyma

2

~

+J

1,6

5.

2

;j
Q)

(IJ

U)

U)

1,6

1,6

3

o

0

~
~III

0
'"CI

1,6

Left lung
bronchus

1,6

0

5,6

4.

6

;j

o

2,4,
5,6,
7

Right lung
parenchyma

1,6

(IJ

tJ

5,7 2,5,
6

3.

1

o

2,4

Right lung
bronchus

6

;j

tJ
0
tJ
0

rJl

Po&lt;

2.

1

,..0

0

S

(IJ

.

c,

Po&lt;

Upper resp.
tract

5

s::

Q)

l.

5

(IJ
!II

r-I
r-I

,..

.

o

+J

2

2

2?

2

74BHS-40

3,7

�-102Table 3a. Bacteriological
Dakota bighorn epizootic.

summary of cultures from captive South

I
I'd
"0
o,.,j

§

o,.,j

.

J.I
"0

Q)

.j..I

fii CIl

J.I 0
0
U

.-l 0
O~
Uo,.,j
.-l
• 0
r&gt;::l U

2

Iii

1,11

5

,0

Q)

~~~
~

1-

74BHS-28

3

2.

74BHS-29

5,1

3.

74BHS-30

4.
5.
6.
7.

74BHS-3l

e

P-

o,.,j

I'd
.-l
.-l
Q)
,...
Q)

::t:

PCIl

I'd
.-l
.-l

I'd

o,.,j

J.I
CIl
CIl

Q)

Q)

o,.,j

.-l

Q)

~

Z

3
11

1

I'd
.-l
.-l

Q)

Q)

J.I

J.I

CIl

='

.j..I

Q)

='

Q)

.j..I

.j..I

CIl
I'd

CIl
I'd

0
J.I

Q)

p..

CIl
I'd

r::

='

p..

1

4

1

3

2,3

Nasal cavity/upper respiratory
Right lung bronchus
Right lung parenchyma
4. Left lung bronchus
5. Left lung parenchyma
6. Peripheral blood
7. Spleen
8. Mediastinal lymph node
9. Peritoneum
10. Pericardium
11. Mannnary tissue

EI
"0

0

='
CIl
Q)

p..

1

4,5

.

PCIl
CIl

='

U
U
0
U
0

~

'¥
I'd

.

PCIl
CIl

='

U
U
0
U
0

e
Q)

J.I

.j..I

.j..I

tI)

tI)

3

3

1,6,
7

9

5

5

1
1

2,3
2,3, 2,3,
1
1,2

tract

0

3,2
5,4, 5,6,
1,6, 11
7,8,

5

ji

I'd
.-l
.-l

p..

2

~

1
1

3

I~l

~

5

4

--.--1.
2.
3.

CIl

CIl
,0

74BHS-40

Code:

.

P-

Q)
o,.,j

5

74BHS-32
74BHS-34

CIl

.

U
0

.j..I

.-l

1

1

1

1

1,3

1

2,1

3

�-103-

Table 3b. Bacterial isolates from the langs of bighorn sheep
with pneumonia.

r'requency

Organis~
Pasteurella

hemolytica

5/7

(71%)

Pasteurella

multocida

5/7

(71%)

3/7

(43%)

Corynebacterium

pyogenes

Neisseria

sp.

4/7

(57%)

Hemophilus

ovis

2/7

(29%)

Staphyloccus

sp.

4/7

(57%)

Streptococcus

sp.

3/7

(43%)

Mima polymorpha

2/7

(29%)

E. coli

1/7

(14%)

Herella sp.

1/7

(14%)

�-104SEGMENT OBJECTIVE

#2

Determine the bacterial, mycoplasma and viral agents involved in the
pneumonia responsible for excessive lamb mortality in the Pikes Peak herd.

PROCEDURES
Serology will be performed by screening sera froD bighorn sheep for precipitating, agglutinating or complement fixing antibodies to antigens from
the various potential pathogens isolated previously from bighorn sheep.
Attempts to isolate pathogens from nasal swabs and lesion material will be
continued using standard bacteriological isolation and identification
techniques.
Seriological survey for viruses will be performed by screening sera from
bighorn sheep for virus neutralizing antibodies in tissue culture systems.
The virus to be used in the survey are (1) Para-influenza-3,
(2) Bovine
Virus Diarrhea, (3) Infectious Bovine Rhinotracheitis, (4) Blue Tongue
Virus, (5) Epizootic Hemorrhagic Disease of Deer, (6) Parvo-virus, (7)
Adenovirus and any other virus that becomes av,~ilable for study that is
considered a potential pathogen.
Virus isolation attempts will be made
in various tissue culture systems in search of the above named viruses.

RESULTS
In addition to the examination of lambs collected on Pikes Peak during
June, 1974, a comple~e examination was conducted on all other sheep
handled during the 1974-75 year. The South Dakota epizootic (captive
sheep) results are given in Table 3. The trapping-banding-transplanting
in the Poudre River herd results are given in Table 4. The trappingbanding results for the Rampart Herd are given in Table 5.

SEGMF~

OBJECTIVE #3

Initiate an epizootiological study of bedding and lambing grounds to
characterize the foci of infection with Protostrongylus.

PROCEDURES
Preliminary studies conducted at Meeker, Colorado, and on Pikes Peak indicate that sheep are becoming infected with~. stilesi on certain preferred
bedding areas. In view of the strong possibility that control of spontaneous
lamb mortality can be achieved by eliminating the source of environmental
infection, these bedding areas found to be foci of infection must be
characterized.
This includes location, vegetation, size, usage, snails present and percentage infected and lungworm larvae present.

�-105Table 4.
results.

Poudre trapping operation - January 21, 1975, nasal bacteriological

-

.

to

~
~

to
I'd

~

..-I

E

..-l

I'd

0
J.I

Number

to

::I
..-I
..-I

-e+

0

~
..-I

(I)

0
I'd

~

&lt;

&lt;

75PR-5

+
+
+
+

75PY-l
75PY-2
75PY-4
75PY-5

+
+
+
+

75PR-l
75PR-3
75PR-4

+

----

7SPB-14
75PB-lS
7SPB-l7
75PB-l9
75PB-Ram

B

Y
R

Blue collar
Yellow collar
Red collar

to
I'd

Q)

tl
C'il
,0

to

(I)

Q

I'd

..-I

J.I

-

(I)

J.I

(I)

:x::

F. ~

(I)

to
to
..-l

~
..-l

J.I
0

a ~

(I)

z

~

~

to
to..-l
::I J.I

0

a0

"0

~
0..-1~

::I
to
p.,
(I)

~ ~

I~
I'd

..-I
..-I

0
0
0
0
0

Q

0

:x::

a
..-I

a

I'd

tI)

to
::I

"0

0
0
0
0

\1-1

Q)
..-I
..-I

~

-e

g

I'd

J.I

+J

+J

tI)

tI)

+

+

+
+

+
+

+
+
+
+
+
+

+

+

+

+
+

+

+

+
+

++

+

+

+

+

+
+
+
+
+
+
+
+
+
+
+
+

+

+

+

+
+

+
+

+

+
+

I
+
+
+

+

+

+

+

+
+
+

+
+

+
+

+

+
+

+
+

+
+

+

e

+J
Q 0
Q)\I-I
"0..-1
..-1..-1
Q 0
PU

0
+J

(I)

+

+

75PB-l
75PB-2
75PB-3
75PB-l,
75PB-5
75:?B-7
75PB-·8
75FB-9
75PB-I0
75PB-ll
75PB-12
75PB-13

..-I
..-I

+J

(I)

to
::I

Q)

�-106-

Discussion:
Bacteriological Results -- nasal swabs of sheep trapped
in Poudre Canyon, Jan. 21, 1975.

Most all bacteria isolated were normal flora for the nasal cavity and
upper respiratory tract. Though opportunists exists, there is no
evidence of primary etiological significance which can be attached to
any of these isolated organisms. Though no Pasteurella species were
confirmed the existence of Mima/Herella species points out the possibility of a morphologically similar organism's (such as Pasteurella sp.)
ability to step in and create secondary opportunistic bacterial complications. To this time I have not cultured any organism in diseased
sheep (particularly associated with lungworm-pneumonia complex) as often
as Pasteurella species even though this genus was not a common isolate
from routine nasal swabs.
I am at a loss at this time to explain satisfactorily the isolation of
Salmonella species from the nasal cavities of these sheep.

�-107Table 5. Rampa!'t Range trapping operation.
April 4, 1975.

Nasal bacteriological

results,

.

:;

W/GS=White with green stripe collar
Y=Yellow collar
R=Red collar
OT=Orange tag

.

•

0.
(/)
(/)

~

'"

Ir.l

C
0

6

~

0
H

Z

Q)

&lt;

(/)
Q)

0.

Q)

(/)

c

!l
.-4'" 0.

~

()

&lt;

::s
~

.

i-4

(/)

I:Q

75RW/GS-4
75RY-6

.

(/)

+

o.-l

6

H
0
4-&lt;
o.-l

()

.-4

'"

0
U

+

+

+

+

75RY-7

++

75RY-8

+

+

++

+

75RY-9

No

75RY-ll

+

~~

Q)

Q)

Q)
(/)

'"
C
H •.-l

•.-l

+J

o ()

::E: &lt;

.~

::s

H

Q)

Ul

+J

'"

Q)

H
p.,

::s
:&gt;

Z

o~

'"

C
0

6

0
-0

::s

Q)
(/)

p.,

()

i~,

()
()

0

~'"
~

+J

0

H

Q)

+J

0
0
0

~
~

Q»

::t: 0

'"

()

0.

H

+J

+J

CI)

CI)

+

+

Q)

::s
Q)
(/)

c,
'" ~

+

+

+

+

+

+

++

+

+

+

++

++
+

+

75RR-2

+

+

75ROT-4

+

75ROT-5

+

75ROT-6

++

+

75ROT-7

+

+-

+

+

+

+

+

75ROT-9

a!
•.-l.
H

(/)

::s

0

(/)

+

+

+

~'" .D'"
~ 0
&gt;&lt;

75RR-l

75ROT-8

..•..... ()

+

+
+

0.
(/)

+

++

+

+

+

+

+

+

++

+

Sample~

75RY-13
75RY-14

H
Q)

+J

+

+
+

+

+

+

+

+
+

+

+

+

+
+

++

++
+

+

+

+

+

+

++

++

+

+

�-108-

Table Sa.
4, 1975.

Rampart Range trapping operation.

Animal Number

Dry Wt (g.)
Feces Ana1ysed

Lungworm

fecal results, April

Total L1

L1/g. Dry Feces

75RW/GS-4

3.0

2

0.7

75RY-7

3.0

3

1.0

75RY-8

3.7

38

10.3

75RY-ll

4.0

7

1.7

75RY-13

4.0

1

0.3

75RY-14

2.7

0

0

75ROT-4

4.0

49

12.5

75ROT-6

2.2

205

93.2

75ROT-8

2.7

86

31.7

75ROT-9

3.0

52

17.3

�-109-

RESULTS
During July, August and September, 5 trips were made to Pikes Peak to
collect snails. Snails were found in 14 locations. Many areas were
examined where snails were not found. A total of 885 shells were
found, 361 snails were alive. Snails of the genus Pupilla were the most
numerous.
Twenty-nine snails were infected, including 24Ptipilla, 3
Vertigo and 2 Euconulus.
These contained 62 infective larvae and 18
immature larvae.
Observations indicated that Pupilla reproduces during wet periods of the
summer and that these moist times also allowed infection of the snails.
Several genera of snails were experimentally infected in the laboratory
including many Vallonia, several Pupilla and Vertigo, a single Gastrocopta,
and 3 Discus. The infection of Q. cronkhitei had not been previously reported. Several Vallonia were infected with 40 larvae.

SEGMENT OBJECTIVE 1t4
Evaluate chemotherapeutic compounds for efficacy against adult Protostrongylus
and third stage larvae in adult sheep and third stage larvae in the fetus.

PROCEDURES
Evaluation of chemotherapeutic agents in 1972 and 1973 for efficacy against
adult lungworm 'proved fruitless; however, preliminary results with cambendazole
indicate that when this compound is administered in the third trimester of
pregnancy the active ingredient crosses the placenta and kills third-stage
larvae stored in the fetal liver. Therefore, cambendazole will be further
evaluated, against captive sheep as well as against remnant herds, for
efficacy against this larval stage.

RESULTS
The bulk of these results have been included in the report by Schmidt

SEGMENT OBJECTIVE

(1976).

ItS

Experimentally infect captive bighorns and b~ghorn/mouflon hybrids with P.
stilesi to: (1) furnish experimental proof of transplacental transmissi~n,
(2) induce pneumonia, and (3) provide infected sheep for chemotherapeutic
studies.

PROCEDURES
A herd of bighorn/mouflon hybrids was obtained late in 1973 and a herd of
bighorn sheep was obtained from South Dakota in early 1974. Some of these

�-110-

sheep are currently being experimentally infected with Protostrongylus
stilesi, the remainder will serve as uninfected controls.
Once infected,
some of the infected sheep will continue to receive infective larvae in
an effort to induce somatic storage followed by transplacental transmission.
Once transplacental transmission has been accomplished, these sheep will
then be used in chemotherapy experiments.

RESULTS
The bighorn sheep brought in from South Dakota died. unfortunately,
any significant results could be obtained.

before

Severe mortality among captive snails hampered experimental infection work.
Mites, poor moisture control, nematodes and spoiled food were all considered
as possible causes of the mortality.
Severa.l insecticide preparations were
tested to find one which would kill mites without killing snails. None were
satisfactory.
Collection of wild snails was required to meet our needs.
Several trips were made to Glen Eyrie, northwest of Colorado Springs, to
collect snails. A closer site near Lyons was found in the spring of 1975.
Several thousand snails were collected at this location.
The difficulties in producing infected snails hampered further work on
transplacental transmission and infection of sheep, but a concerted effort
by many people allowed us to produce several thousand larvae. These snails
were used to infect sheep.
Date
Dec.
Jan.
Apr.
Apr.
Apr.
May.

1974
19-1975
8-1975
13-1975
17-1975
1975

/I Snails

Infective Larvae

80

401
300
1559
1095
279
60

150
248
95

Innnature Larvae

Sheep II

lamb
Given to Dr. Lauerman
35
R-76
82
1987
24
1988
Used in antigen prep.

Three lambs born in the spring of 1974 to bighorn-mouflon hybrid sheep were
found to have been transplacentally infected. Data on these lambs is:
Experimental
Ewe II

Date

F-11

Infection

II of Larvae

Date Lamb Born

2-22-74
3-1-74
4-3-74

513
748
569

5-10
74MX-lO
(Rebecca)

F-12

5-3-74

574

5-27
74MX-6

1987

4-30-74

729

6-7
Sugar

Results
On 7-29-74 she was
producing 210 larvae
per gram of feces;
died 7-29.
Died on 7-11. Found
to be infected by
histologic examination
of lungs.
On 8-10 she was producing 300 Ll/gram.

�-111Larvae from Sugar's feces were collected and used in experimental infections of snails. Gretchen, the other surviving lamb, was infected
on 12-3-74 by the administration of 401 infective larvae. Larvae from
her feces were also used in experimental infections. These larvae served
as our main source of larvae for infections performed in the spring of
1975.
Both of these lambs suffered the after effects of hypervitaminosis D which
led to the death of several other lambs. Their condition progressively
became worse and they were euthanized. Significant observations were made
on the difference in the infection in a pre-natally and post-natally infected animal.
Further information was gained in the 1975 lambs, this is summarized below:
Ewe

II of Larvae

Lamb Born

R-76

4-8-75
1559L

5-11

Difficult birth resulted in
death of lamb. Digestion
of tissues yielded 45L3 in
the liver, 1 in diaphragmatic
lobe of right lung and 4 from
placental tissues.

5-1-75
Xenobia

Patent at 45 days 150 larvae/g
on 6-24. By late August her
fecal output dropped to much
lower levels.

5-8-75

Not patent.

1

1988

60L3--9-19-74
279 L --4-17-75
3

1987

729 L --4-30-74
3

Results

1095 L3--4-13-75
F-ll

See previous table

Not patent.

F-12

See previous table

Not patent.

Red

60 L3--9-19-74

No lamb

At about 2 weeks of age Xenobia became anorectic. She became emaciated and
had subnormal temperatures. After a week she was given baby vitamins and
later that day she began to increase her intake of milk. None of the other
lambs showed any similar signs.

Raising of Lambs
All the lambs from the mouflon-bighorn hybrid and the 4 captive bighorn ewes
were hand raised. The lambs were left with the ewe for about 24 hours and
then captured and taken to a backyard. They were fed whole cow's milk with
1 teaspoon corn syrup, 1 tablespoon codliver oil, and a beaten egg added to
each 24 oz. of milk. Early in life the lambs were fed every 3 hours. Lambs
were weaned at about 2 months of age. All lambs born alive are still alive
and appear very healthy.
Nearly 50 percent of the l~mbs were stillborn. Three mouflon/bighorn cross
ewes and one bighorn ewe did not produce lambs. Eight live lambs were obtained.

�-112-

SEGMENT OBJECTIVE

#6

Evaluate molluscacides and larvicides for evidence of efficacy against
snails and lungworm larvae (will be closely coordinated with Work Plan
1, Job 23).

PROCEDURES
Molluscacides cmd larvicides are currently being evaluated in the laboratory. Known numbers of snails are exposed to small amounts of a molluscacide
by spraying them in their natural environment.
A constant amount of spray,
using varying dilutions of active ingredient is used. Larvicides are
applied in the same manner, but the number of la~~ae/gram feces is first
calculated.

RESULTS
The following molluscacides and larvicides were evaluated for their efficacy
against snails and for first-stage larvae of Protostrongylus:
1) Formalin
at 0.25 to 2% solution; 2) Sodium Bonate; 3) Zectran 2E; 4) Dursban; 5)
Balcite (5% para formaldehyde); 6) Vapotox (Parathion); and 7) Mesurol
(Methyl-dimethyl carbanate).
Formalin at all of the dilutions given above was effective in killing snails
and larvae; however, the solution also effectively killed herbage.
Sodium
Bonate was not effective against snails and/or larvae, but effectively killed
herbage even at weak solutions.
Zectran 2E and Dursban are Dow Chemical
products and both are insecticides.
Neither was mixable with water (although
advertised to be). While both are insecticides, both (supposedly) have
molluscacide properties.
Neither comp~und killed snails and/or larvae at
the recommended application.
Balcite and Vapotoxare
both designed to be
applied as a dry-mix and to be effective necessitates snail and/or larval
contact.
Both, when applied at the recommended level, were ineffective;
primarily because the organisms had to "crawl-over" this material.
At this
level the chances of a snail or larva coming into contact with the material
was negligible.
The latter compound (Vapotox) is a power insecticide
(parathion) that is {environmentally) too dangerous to be applied wholesale
at a higher level - if, indeed, at all.

SE&amp;"MENT OBJECTIVE

#7

Attempt to locate the site of somatic storage for third stage
adult sheep dying from any cause.

K. stilesi in

PROCEDURES
Large amounts of tissue from adult female bighorn sheep and fetal tissues are
macerated and soaked in physiological saline for 24 hours; following this
procedure, the tissues are digested with pepsin-hydrochloric acid. Using
these two procedures, eventually the site of somatic storage for third stage
larvae will be discovered; moreover, the number of larvae infecting the fetus

�-113can be determined.

RESULTS
Since July 1, 1974, 18 sheep have been examined for Protostrongylus sp.
third stage larvae (L3) somatically stored in body tissues. Of these
animals three sheep yielded L3 in tissue digestion fluids.
Case #1: In late January, 1975 a yearling ewe suffered a broken
femur in a trapping operation in Poudre Canyon. She died a week later
of pneumonia.
Four Protostrongylus sp. L3 were found in lung digestion
fluids.
Case #2: In late January, 1975 a ewe was brought in by Russ Mason,
WCO at Grant, Colorado, from the Indian Creek area near Mt. Evans. She
had been dead for several hours of a bronchopneumonia.
Lung digestion
fluids yielded 19 Protostrongylus sp. L •
3
Case #3: In mid May a lamb was born dead to an experimental bighorn/
mouflon ewe. Digestion fluids of fetal liver and lung yielded 45 and one
Protostrongylus sp. L3, respectively.
Digestion of the ewe's passed
placental membranes yielded 4 Protostrongylus sp. L3.

SEGMENT OBJECTIVE

#8

Study the epiz(lotiology of verminous pneumonia
herd.

in the Poudre bighorn sheep

PROCEDURES
Analysis of the epizootiology of Protostrongylus stilesi in the Poudre
River bighorn sheep herd is being measured by three parameters: analysis
of lungworm in fecal pellets; snail density and degree of infection of
snails by lungworm larvae; and lamb production and survival.

RESULTS
Larval output by the bighorn sheep herd has fluctuated throughout the
year as reported by other investigators.
Mean larval output for the time
period January 1, to March 31, 1975 fluctuated from a low of 1.2 larvae/gm
to a high of 1418.4 larvae/gm for individuals, with a mean of 208.8 larvae/
gm for all samples collected in that time period.
Snail densities and infections computed for 2 of 6 areas samples is
currently standing at 2.4 percent of the live snails collected being infected, and a density of 91 live snails per square meter in the areas
sampled.

�-114Observed 13mb per 100 ewes for the summer has averaged 26.9 lambs per
100 ewes, the period including June, July, and August.
June lamb/ewe
ratio - 23.2 lambs/IOO ewes. July lamb/ewe ratio - 47.2 lambs/IOO ewes.
August lamb/ewe ratio - 15.0 lambs/IOO ewes. These figures represent
incomplete work conducted on this portion of the study, and recent
observations have shown that lamb/ewe ratios may be considerably higher
than the three months of data indicates.

LITERATURE

CITED

Schmidt, R. L. 1976. Monitoring Bighorn Sheep Populations.
Div. Wildl., Game Res. Rept., January.
(In press).

Prepared by

Colo.

~~==~~,.~~~~~~.~~~~-__
C. P. Hibler

�-115-

JOB PROGRESS REPORT

State of

Colorado
--~--~~~~---------------------

Project No.

W-124-R-3

Work Plan No.

Raptor Investigations

1

Job No. 1

Job Title Statewide Raptor Populations and
Period Covered:
Personnel:

Characteristics

Studies

March 1, 1974 through February 28, 1975

Gerald R. Craig

ABSTRACT
Two hundred and forty three golden eagle ~ests were investigated,
of which 65

were occupied and 21 produced young with an average

brood size of 1.25.

The number of active peregrine falcon eyries

declined from 11 to nine, but production increased from 2 young
in 1973 to 12 young in 1974 (average brood size of 2.40).

Prairie

falcon productivity remained at 3.70 young/site for 34 of 60 active
sites.

A density of 9 eagles/IOO square miles was determined for

the trend area in Northeastern Colorado.

Wintering bald eagles

increased slightly along the trend area of 'the South Platte River.
Mortality figures are also provided.

��-117STATEWIDE RAPTOR POPULATIONS AND CHARACTERISTICS
Gerald

STUDIES

R. Craig

To investigate status of current populations,

characteristics

and trends

including distribution, migration, patterns and chronology of Bald and Golden
Eagles, Osprey, Peregrine and Prairie Falcons and Burrowing Owls in or through
Colorado.
SEGMENT OBJECTIVES
1.

Develop and refine methods for determing breeding numbers of Bald and
Golden Eagles,

Osprey, Peregrine and Prairie Falcons and Burrowing Owls

in Colorado.
2.

Develop and refine methods for determining winter trend information for
Bald and Golden Eagles, Peregrine and Prairie Falcons and

Burrowing

Owls

in Colorado.
3.

Locate and categorize major and important habitats and associated Bald
and Golden Eagle, Osprey, Peregrine and Prairie Falcon and Burrowing Owl
populations in Colorado.

4.

Establish and maintain annual or periodic inventories of Bald and Golden
Eagles, Osprey, Peregrine and Prairie Falcons and Burrowing Owls in
Colorado from inventory methods

selected from Project Statement Procedures

1 and 2
5.

Monitor mortalities and inspect injured and dead raptors reported or
brought to the Division to determine mortality

factors of Bald and Golden

Eagles, Osprey, Peregrine and Prairie Falcons and Burrowing Owls and other
species that are obtained incidentally.
6.

Initiate and maintain a lO-year raptor banding program to determine migration, chronology, movement, life tables and mortali~ies

of Bald and Golden

Eagles, Osprey, Peregrine and Prairie Falcons and Burrowing Owls and other
species that are encountered accidentally.
7.

Prepare a series of comprehensive information leaflets and recreational
guides at intervals when the information becomes available.

�-118-

RESULTS
Only 65 of 243 golden eagle nests were determined to be occupied in 1974.
Only 32% (21) of the eyries produced young.
site with a total of 25 young produced.

Average brood size was 1.25 young/

Fledging success was 1.21 young/site.

The steep decline in active sites (79% were active in 1973) and subsequent
meager production may be blamed upon a steep decline in jackrabbits, which are
the predominant prey of golden eagles.
An active bald eagle nest was located at Electra Lake, in La Plata County.
Two adult bald eagles were observed occupying an old osprey nest at the lake's
edge.

The female was incubating 1 egg which subsequently failed to hatch.

Cause for desertion of the nest was not determined.
The decline in nesting peregrines continued with a reduction from 11
active sites in 1973 to 9 in 1974.

Production, however, was significantly

improved with a total of 12 young having been hatched and fledged from 5 sites
(2.40 young/site).
Prairie falcon production continued at the same level as 1973 with 60
(83%) of 72 sites being active.

The average brood size was 3.70 young/site

with an average fledging success of 3.00 young/site.
Despite the decline in nesting success, the wintering golden eagle population in northeastern Colorado maintained levels similar to 1973, and 1974.

In

fact the density was slightly greater (9.0 eagles/lOO; sq. miles) in 1975.

The

number of wintering bald eagles along the South Platte River trend count also
increased slightly in 1975.
Data about mortality factors are presented in Tables

10 and 11.

�-119-

TABLE 1
ACTIVITY AND PRODUCTIVITY

Site No.

Activity

CH 1

IA

CH 2

IA

OF COLORADO GOLDEN

No. of Young

EAGLES

No. Fledged

CH 3

Comments

Active in 1973

CN 1

A

CS 1

A

CU 1

IA

CU 2

A

DO 1

IA

DO 2

A

EA 1

A

Adult incubating

EA 2

A

Adult incubating

EA 3

IA

EA 4

A

EA S

IA

EA 6

A

EA 7

IA

Adult incubating
1

1

Adult incubating

Adult incubating

EA 8
EA 9

IA

EA 10

IA

EA 11

A

EA 12

IA

EA 13

IA

EA 14

Young 7-8 weeks of age

1+

1+

A

1

0

EB 1

A

1

0

EP 1

IA

EP 2

IA

EP 3

IA

FR 1

IA

GA 1

IA

GA 2

IA

GA 3

IA

�-120Table 1 (Continued) Activity

Site No.

Activity

GA 4

IA

GA 5

IA

GA 6

A

GA 7

IA

and Productivity

of Colorado Golden Eagles

No. of Young

No. Fledged

1

1

GA 8

COIllII1ents

Active in 1973

GR 1

IA

GR 2

A

GR 3

IA

GR 4

IA

GR 5

IA

GR 6

IA

GR 7

A

GR 8

A

GR 9

IA

GR 10

IA

GR 11

IA

GR 12

IA

HI 1

IA

HI 2

IA

HU 1

A

HU 2

IA

HU 3

A

HU 4

IA

HU 5

A

HU 6

A

HU 7

IA

2

2

Adult incubating

Adult incubating

Adult incubating

Adult incubating

HU 8
JE 1

IA

JE 2

IA

JE 3

A

JE 4

A

JE 5

IA

La 1

A

Adult incubating
2

2

0

0

�-121-

Table 1 (Continued) Activity and Productivity

Site No.

of Colorado Golden Eagles

Activity

No. of Young

No. Fledged

LO 2

A

0

0

LO 3

A

2

2

LP 1

IA

LP 2

IA

LP 3

IA

LP 4

IA

LP 5

IA

Connnents

LP 6
LS 1

IA

LS 2

IA

LS 3

IA

LS 4

IA

LS 5
ME 1

IA

ME 2

IA

ME 3

A

ME4

IA

ME 5

Active in 1973

ME 6
ME 7

Active in 1973

ME 8
MI 1

A

MI 2

IA

MI 3

IA

MI 4

IA

MF 1

IA

Active in 1973

MF 2

IA

Active in 1973

MF 3

IA

MF4

IA

MF 5

IA

Active in 1973

MF 6

IA

Active in 1973

MF 7

IA

MF 8

IA

MF 9

IA

MF 10

A

1

1

Active in 1973

1

1

�-122and Productivity

Table 1 (Continued) Activity

Site No.

of Colorado Golden Eagles

Activity

No. of Young

No. Fledged

MFU

A

0

0

MF 12

IA

MF 13

IA

MF 14

A

MF 15

IA

MF 16

A

1

1

MF 17

A

0

0

MF 18

IA

MF 19

A

MF 20

IA

MF 21

IA

Active in 1973

MF 22

IA

Active in 1973

MF 23

IA

Active in 1973

MF 24

IA

Active in 1973

MF 25

IA

Active in 1973

MF 26

IA

Active in 1973

MF 27

IA

Active in 1973

MF 28

IA

Active in 1973

MF 29

IA

Active in 1973

MF 30

IA

Comments
2 spoiled eggs

Active in 1973
1

1
Active in 1973

Active in 1973
2 eggs, Success unknown

MF 31

Nest not located in 1974

MF 32

Nest not located in 1974

MF 33

Nest not located in 1974

MF 34

IA

MF 35

IA

MN 1

A

MN 2
MN 3

IA

MN 4

A

MN 5
MN 6

IA

MN 7

A

11N 8

A

PA 1

IA

PI 1

IA

PI 2

A

PU 1

IA

PU 2

A

PU 3

IA

Active in 1973
1

1

�-123Table 1 (Continued) Activity

Site No.

Activity

PU 4

IA

PU 5

A

PU 6

IA

PU 7

A

and Productivity

of Colorado Golden Eagles

No. of Young

No. Fledged

1

1

Connnents

RI 1
RI 2

IA

RI 3

IA

RI 4

IA

RI 5

A

RI 6

IA

Active in 1973

RI 7

IA

Active in 1973

RI 8

IA

RI 9

IA

RI 10

IA

RI 11

IA

RI 12

IA

Active in 1973

RI 13

IA

Active in 1973

RI 14

IA

Active in 1973

RI 15

IA

Active in 1973

RI 16

IA

RI 17

IA

RI 18

IA

RI 19

IA

RI 20

IA

RI 21

IA

RI 22

IA

RI 23

IA

RI 24

IA

Active in 1973

RI 25

IA

Active in 1973

RI 26

IA

RI 27

IA

RI 28

IA

Active in 1973

RI 29

IA

Active in 1973

RI 30

IA

Active in 1973

Active in 1973
1

1

Active in 1973

Active in 1973

Active in 1973

Active

in 1973

�-124Table 1 (Continued) Activity and Productivity
Site No.

Activity

No. of Young

of Colorado Golden Eagles

No. Fledged

Connnents

RI 31

IA

RI 32

IA

RI 33

IA

RI 34

IA

RI 35

IA

RI 36

IA

RI 37

IA

RI 38

IA

Active in 1973

RI 39

IA

Active in 1973

RI 40

IA

RI 41

IA

RO 1

IA

RO

2

IA

RO 3

IA

SA 1

A

SA 2

IA

Active in 1973
Active in 1973
Active in 1973

Active in 1973

1

1
Active in 1973

SA 3
SA 4

Active in 1973
IA

SN 1
SU 1

IA

SU 2

IA

SU 3

IA

WE 1

IA

WE 2

IA

WE 3

IA

WE 4

A

WE 5

IA

WE 6

IA

WE 7

IA

WE 8

A

WE 9

A

WE 10

A

WE 11

A

0

0

WE 12

A

1

1

WE 13

A

0

0

0

Nest blown down

0
2

2
0

�-125Table 1 (Continued) Activity and Productivity of Colorado Golden Eagles
Site No.

Activity

No. of Young

No. Fledfed

WE 14

IA

WE 15

IA

WE 16

A

0

0

WE 17

A

0

0

WE 18

A

1

1

WE 19

A

0

0

WE 20

A

0

0

WE 21

A

0

0

WE 22

IA

WE 23

IA

WE 24

IA

WE 25

A

1

1

WE 26

A

0

0

WE 27

IA

WE 28

A

0

0

WE 29

A

0

0

WE 30

A

0

0

WE 31

IA

WE 32

A

0

0

WE 33

A

0

0

WE 34

IA

WE 35

A

0

0

WE 36

IA

WE 37

IA

WE 38

IA

WE 39

IA

WE 40

A

0

0

WE 41

A

0

0

Connnents

�-126-

TABLE 2

SUMMARY OF PRODUCTIVITY OF COLORADO GOLDEN EAGLES
65 of 243 sites (26%) were determined to be active.
21 (32%) of the active sites were known to have
produced young.
19 (29%) of the active sites were known to have
fledged young.
A

total of 25 young were observed in 20 nests ;Jrior
to fledging
(brood size of 1.25 per productive site)

A total of 23 young were observed fledged froD 19 sites (fledging
Success of 1.21 young per productive site).

�TABLE 3
OCCUPANCY

Site
1
2
3
4
5
6
7
8
9
*10
*11
12
13
14
15
16
17
18
19
20
21
22
23
24

*

OF COLORADO PEREGRINE FALCON EYRIES

Pre-1964

1964

1965

P
P

P
P
A
A
M

+
+
+

P

1967

1968

1969

1970

1971

1972

1973

1974

P
P

P

P
P
P
P
P

M
P
P

P
P

P
P

P
P
P
A
P
P
P
P
P

P
P
P
P
P
P
P
P
P
P

V
V
V
V

V

M
P
M
V
P
P
P
V
P
V
P
V
V
V
P
V
V
V
V
V
V
V
V
V

P
P
A
P

V

+

P

+
+
+
+
+
+
+
+

1966

1964-1974

A
F
P
V

P
M
P

A

1/

V
V
V
V

+

V
V

+

V

A

These are neighboring

V

V
A

sites and this could represent a shift of one pair from site 10 to site 11.

1/ An adult female was found dead in the vicinity of the eyrie.
+ = Known to have had pairs present prior to 1964.
Blank Spaces = No data available for that year.
P = Pair.
M = Male.
F = Female.
A = Lone adult.
V = Site was vacant.

V
V
V

V
V
A

P
V
A
V
V
V
V
V
V
V
V
V
V

I

t--'
N

.....•
I

�-128TABLE 4
RESULTS OF NESTING ATTEMPTS OF COLORADO PEREGRINES IN 1974

Eyrie

Visits

Eggs

Young

1

4

o

o

Lone male observed in March

2

3

3

o

3 eggs, two broken, other infertile May 5.

5

3

3

2

Pair incubating AprilS; 2 one week old
young June 3; 2 large young June 26.

6

5

3

3

Pair incubating April 12; 3 eggs May 2;
3 large downy young May 29; 3 flying
young June 22.

7

5

2

9

2

t

o

Adults still courting in early June.

11

4

4

3

Pair copulation April 7; 4 eggs May 23;
3 young June 4; 3 young fledged June 27.

15

3

3

2

Pair copulating; possibly laying April 29;
2 small young and viable egg June 4;
2 large young June 26; young fledged and
observed in July.

Remarks

Pair copulating AprilS; 1 small young
and 1 spoiled egg June 3; 1 young fledged
June 25.

�-129TABLE 5
PRODUCTIVITY OF COLORADO PRAIRIE FALCONS

Site No.

Activity

AR 1

A

AR 2

A

CU 1

A

CU 3

A

CU 5

No. of Eggs

No. of Young

No. Fledged

Comments

5

5

5

A

4

2

2

DO 2

A

3+

3+

3+

3 yng. observed after fledging

00 3

A

3+

3+

3+

3 yng. obs. after fledging

004

A

2+

2+

2+

2 yng. ob s , after fledging

005

A

D06

A

DO 7

A

1+

1+

1+

1 yng. obs. after fledging

DO 8

lA

1+

1+

0

DO 9

A

00 10

A

00 11

A

00 12

A

EA 1

A

EA 2

A

EA 3

A

EB 1

A

EB 2

lA

EP 1

A

FR 1

A

GR 1

A

HU 3

A

HU 5

A

JE 1

A

LR 1

A

LR 2
LR 3

CU 6
DO 1

Apparently had produced young
since adults were defensive
3+

3

3

1+

1+

1+

1 young obs. after fledging

3

2

2

(1 male, 1 female)

A

4

4

4

(1 males, 2 females)

A

5

5

4

1 yng. dead (3 males, 1 female)

4

2+

�-130Table 5 (Continued) Productivity

Site No.

of Colorado Prairie Falcons

Activity

No. of Eggs

No. of Young

No. Fledged

LR 4

A

2+

2

2

Adult female with jesses

LS 2

A

LS 3

A

MF 1

A

PA 1

A

8

A

2+

2+

2+

At least 2 yng. heard
in nest

PU 9

A

2+

2+

2+

At least 2 yng. heard
in nest

WE 1

A

5

5

5

WE 2

A

3+

3

3

WE 3

IA

WE 4

A

3+

3

2

WE 5

A

4+

4+

4

WE 6

IA

WE 7

A

5

5

3

WE 8

A

5

5

4

WE 9

A

4+

4

1

WE 10

A

1+

1+

1

WE 11

A

WE 12

IA

WE 13

A

5

5

4

WE 14

IA

WE 15

A

WE 16

IA

WE 17

A

4+

4

3

WE 18

A

4+

4+

4

WE 19

A

4+

4

2

WE 20

A

1+

1+

0

WE 21

A

0

0

WE 22

A

4+

4

1

WE 23

IA

WE 24

IA

WE 25

IA

WE 26

IA

4+

4

4

WE 27

A

2+

2

2

PU

Connnents

Young taken by falconers

�-131Table 5 (Continued) Productivity

Site No.

Activity

of Colorado Prairie Falc.ons

No. of Eggs

No. of Young

No. Fledged

WE 28

A

0

0

WE 29

A

0

0

WE 30

A

3+

3

WE 31

A

WE 32

IA

WE 33

A

4+

4

4

WE 34

A

3+

3

3

3+

0

Connnents

�-l32-

TABLE 6

SUMMARY OF ACTUAL AND ESTIMATED PRODUCTIVITY OF COLORADO PRAIRIE FALCONS
60 of 72 (83%) sites were determined to be active.
36 (60%) of the active sites produced eggs.
34 (56%) of the a.ctive sites hatched young.
32 (53%) of the active sites fledged young.

= 106
Average brood size of 20 sites with known production = 3.70
Total number of young observed prior to fledging

Adjusted young production for all active sites hatching young (34 sites) = 125
Total number known to have fledged from 25 sites = 75 (average fledging success
of 3.00)
Adjusted fledging success of all productive sites = 96

�-1331974
OSPREY ACTIVITY AND PRODUCTIVITY

Site No.

Activity

No. of Eggs

No. of Young

No. Fledged

Comments

GR 1

A

GR

2

A

2

0

0

1 rotten egg in nest
1 broken egg on ground

GR 3

A

2

0

0

2 rotten

GR 4

A

LA 1

A

LP 1

lA

JA 1

A

JA

A

2

JA 3

IA

eggs in nest

Adult incubating
3

3

3

3

3

3

2

2

Adults present early
in season

�-134-

SUMMARY OF ACTUAL AND ESTIMATED PRODUCTIVITY OF COLORADO

6 of 9 (67%) sites were determined to

be act Lva;

4 (67%) of the active sites produced eggs.
3 (50%) of the acti've sites hatched young.
3 (50%) of the active sites fledged young.
Total number of young observed prior to fledging
Average brood size = 2.67 young/site
Average fledging success

= 2.67 young/site

8

OSPREYS

�-135-

TABLE 7

NORTHEASTERN

DATE:

January 22, 1975

FLIGHT TIME:
CREW:

COLORADO AERIAL EAGLE CENSUS

5 hours and 9 minutes

(8:14-11:30 and 1:22-3:15)

W. Russell, E. Boeker and G. Craig

Weather Conditions
The temperature rose gradually from 300F in the morning, 36°F at midday
and 400F at the end of the flight. Wind was light and variable. Visibility
was excellent with cloud cover increasing from 0% in the morning to 5% at
noon to 50% in the afternoon.
Counting Conditions
Counting conditions were excellent with bare ground over all but the northwestern portion of the census area.
Results
For details, see attached data sheet:
Golden Eagle

Bald Eag1e*

Adult
Immature
Unknown

22
5
0

Adult
Immature
Unknown

3
1
0

Total

27

Total

4

* All bald eagle sightings were along riparian habitat of the South Platte
River.
Extrapo1 ation
This 10% sample of a 3,000 square mile study area yields an estimated 270
golden eagles for the area.

�-136TABLE 8

BALD EAGLE CENSUS OF SOUTH PLATTE RIVER FROM FORT MORGAN TO GREELEY

DATE:

January 22, 1975

FLIGHT TIME:
CREW:

35 min. (3:20 - 3:55)

W. Russell, E. Boeker &amp; G. Craig

Weather Conditions
The temperature was approximately 40° F; wind was calm; visability was
fair to good since the inversion along the river was not as great and the
sun was sufficiently above the horizon so as not to blind the observer.

Counting Conditions
Good to fair with little backlighting.
Results
Golden

Eagle

Bald Eagle

Adult
Immature
Unknown

0
0
0

Adult
Immature
Unknown

28
14

Total

0

Total

42

0

Since this is an exact count, no extrapolation is necessary. This year, there
was little ice on the river to concentrate waterfowl and thus the eagles were
scattered along the length of the river. Further, fewer waterfowl were
present along the river this year because there was no late waterfowl season
to concentrate the waterfowl on more remote stretches of river.

�-137-

TABLE 9
RESULTS OF AERIAL FLIGHTS FOR GOLDEN EAGLES IN NE COLORADO 1972 - 1975

Total Area
.Estimate

Eagles per
100 s9. mi.

Adults

Juveniles

Unknown

Jan. 24, 1973

16

8

0

240

8.0

Jan. 16, 1974

19

3

0

220

7.3

Jan. 22, 1975

22

5

0

270

9.0

Date

RESULTS OF AERIAL FLIGHTS BALD EAGLE CENSUS IN NE COLORADO 1972 - 1975

Date

% Juveniles

Adults

Juveniles

Unknown

Total

Jan. 24, 1973

18

13

0

31

42

Jan. 16, 1974

16

15

0

31

48

Jan. 22, 1975

28

14

0

42

33

�-l38TABLE 10

SUMMARY OF DEAD AND INJURED RAPTORS REPORTED TO THE DIVISION JAN. - DEC. 1974

Q

&lt;,

~

Q

Q

0

.~

rn -IJ

.~ u

........,
••..• &lt;ll
0..0

Great Horned Owl
Golden Eagle
Red-tailed Hawk
American Kestrel
Rough-legged Hawk
Long-eared Owl
Barn Owl
Marsh Hawk
Prairie Falcon
Swainson's Hawk
Goshawk
Short-eared Owl
Bald Eagle
Burrowing Owl
Cooper's Hawk
Ferruginous Hawk
Flammu1ated Owl
Peregrine Falcon
Screech Owl
Turkey Vulture
TOTAL

-IJ
0

.c

rn
Q

;:l

UO

c.!&gt;

8
5
5
8
9
4
3
5

7
4
5

6
6

0
.~
-IJ

0
.~
•••.•rn
C1l rn
bO&lt;ll
&lt;ll rn

•••.•rn
•...•
0

::l

S

0
,...

0
,...

&lt;ll
••..• &lt;ll

0

-IJ

u~
.~
.~
.c,...

~

4-l

••..• -IJ

•••.•rn
C1l &lt;ll

Q

-E

Hp..

r.:.z

2-

1

3

4
1
3
2

1
(,

3
1

o

Q

u

&lt;ll

....•
~
2
8

~
....•
0
.c

Q

rn

0

,...
&lt;ll
.c

~
E-&lt;

:&gt;00

..d....t!..

.~
0

--E:!...-

-IJ

~

0

3
1
1

1
1
1

1
1

&lt;ll-IJ

, o,

bOC1l
&lt;ll ,...

E-&lt;

2
1

3

1

13

6

4

1

2
2
1

1

1
1

1
1

1
1

1
1
1
1
1
1
1
1
66

25

14

11

12

10

10

29
23
20
18

4

2

4

10
8
8
8
7
3
3
1
1
1
1
1
1
1
1
158

�-139TABLE 11

SEASONAL OCCURENCE OF MORTALITY AND INJURY TO RAPTORS - 1974

Cause of Mortality

Jan

Feb

Mar

Apr

May

Jun

JuJ. Aug

Sep

Oct

Nov

Dec

Total

Collision w/Object

9

7

6

1

3

7

5

11

11

2

2

3

67

Gunshot

4

3

5

1

3

3

3

3

1

26

3

10

1

Illegal Possession

3

Unknown

1

Electrocution

3

1

Vehicle Strike

3

1

11

3

1

1

2

1

2

1

1

2

1

1

1

11

1

10
1

1

9
1

11/

Other

20

1/ Tumor in throat
Starving

1/ Bumb1efoot infection

14

4
2

2

Poison

]J

3

1

Leg-hold Trap

TOTAL

5

3

Fall from Nest

14

16

9

8

I])

l'~/

27

9

I])

20

18

4

8

4

5

158

�-140-

TABLE 12

RAPTORS BANDED BY THE DIVISION IN 1974

No. Banded

Species
Swainson's Hawk

34

Ferruginous

22

Hawk

Prairie Falcon

14

Golden Eagle

6

Peregrine Falcon

6

Cooper's Hawk

5

American Kestrel

2

Great Horned Owl

2

Red-tailed

1

Hawk
TOTAL

99

�-141-

JOB PROGRESS REPORT

State of __~C~o~l~o~r~a~d~o~

_

Project No. ~W~-~1~2~4_-~R_-~3

_

Raptor Investigations

Work Plan No.

_

Job No. 2

Job Title:

~l

Raptor Population and Productivity

Studies in Northeastern

Colorado
Period Covered:
Personnel:

March 1, 1974 through February 28, 1975

Gerald R. Craig

ABSTRACT
Twenty-two golden eagle, 24 prairie falcon and six ferruginous
hawk nests were occupied on the 2,000 square mile study area in
Northeastern

Colorado.

Productivity of golden eagles and ferruginous

hawks greatly declined from 1973 and there was a minor decrease in
productive prairie falcon eyries.

��RAPTOR POPULATION AND PRODUCTIVITY STUDIES IN NORTHEASTERN

COLORADO

Gerald R. Craig
To intensively document nesting and wintering

densities, migration

chron-

ology, habitat requirements, productivity and prey preferences of raptors on
the Pawnee National Grasslands and adjacent prairie lands.
SEGMENT OBJECTIVES
1.

Document exact nesting densities and productivity of raptors present on
the Pawnee National Grasslands and adjacent prairie lands.

2.

Document prey items at nest sites.

3.

While at nest sites, record disturbance factors,

land use practices, habi-

tat type and nest site features.
RESULTS
While approximately

one half of the golden eagle eyries (22 of 41 sites)

were occupied by adults, only four sites succeeded in producing and fledging
5 young.

This rather drastic decline may be attributed to a crash in the

jackrabbit population which accounts for the major prey of golden eagles in
good years.

Little data was gathered on prey at eagle nests because so few of

the nests contained young.
Ferruginous hawk nesting numbers declined steeply from 37 occupied sites
in 1973 to six sites in 1974.

Not all sites were visited in 1974 (only 18 were

checked in 1974 when 47 were investigated in 1973) ~rhich biases the results.
However, the extremely poor productivity of the six active sites may be cause
for concern.
Activity of nesting prairie falcons declined slightly (29 sites active in
1973 and 24 active in 1974), but the average brood size of 3.92 young/site compares favorably with 3.94 young/Site of 1973.

Fledging success declined from

3.82 young/Site in 1973 to 2.94 young/site in 1974.

Slightly more pressure may

have been placed on the productivity by falconer&amp; in 1974.

�-144TABLE 1
ACTIVITY

AND PRODUCTIVITY

Site No.

Activity

OF GOLDEN EAGLE IN NORTHEASTERN

No. of Young

No. Fledged

WE 1

IA

WE 2

IA

WE 3

IA

WE 4

A

WE 5

IA

WE 6

IA

WE 7

IA

WE 8

A

WE 9

A

WE 10

A

WE 11

A

0

0

WE 12

A

1

1

WE 13

A

0

0

WE 14

IA

WE 15

IA

WE 16

A

0

0

WE 17

A

0

0

WE 18

A

1

1

WE 19

A

I)

0

WE 20

A

0

0

WE 21

A

0

0

WE 22

IA

WE 23

IA

WE 24

IA

HE 25

A

1

1

WE 26

A

0

0

WE 27

IA

WE 28

A

0

0

WE 29

A

0

0

WE 30

A

0

0

WE 31

IA

COLORADO

Comments

0

Nest blown down

0
2

2
0

�-145-

Table 1 (Continued) - Activity and Productivity

of Golden Eagles in NE Colorado

Activity

No. of Young

No. Fledged

WE 32

A

0

0

WE 33

A

0

0

WE 34

IA

WE 35

A

C

0

WE 36

IA

WE 37

IA

WE 38

IA

WE 39

IA

WE 40

A

0

0

WE 41

A

0

0

Site No.

Comments

�-146-

TABLE 2
SUMMARY OF ACTUAL PRODUCTIVITY OF GOLDEN EAGLES IN NORTHEASTERN COLORADO

22 of 41 sites (53%) were determined to be active.
4 (18%) of the active sites produced and fledged 5 young (1.25 brood
size and fledging Success per site).

�-147TABLE 3
ACTIVITY AND PRODUCTIVITY OF PRAIRIE FALCONS IN NORTHEASTERN COLORADO

Site No.

Activity

No. of Eggs

No. of young

No. Fledged

WE 1

A

5

5

5

WE 2

A

3+

3

3

WE 3

IA

WE 4

A

3+

3

2

WE 5

A

4+

4+

4

WE 6

IA

WE 7

A

5

5

3

WE 8

A

5

5

4

WE 9

A

4+

4

1

WE 10

A

1+

1+

1

WE 11

A

WE 12

IA

WE 13

A

5

5

4

WE 14

IA

WE 15

A

WE 16

IA

WE 17

A

4+

4

3

WE 18

A

4+

4+

4

WE 19

A

4+

4

2

WE 20

A

1+

1+

0

WE 21

A

0

0

WE 22

A

4+

4

1

WE 23

IA

WE 24

IA

WE 25

IA

WE 26

IA

4+

4

4

WE 27

A

2+

2

2

WE 28

A

0

0

WE 29

A

0

0

WE 30

A

3+

3

WE 31

A

WE 32

IA

WE 33

A

4+

4

4

WE 34

A

3+

3

3

3+

0

Connnents

Young taken by
falconers

�-148TABLE 4
SUMMARY OF ACTUAL AND ESTIMATED PRODUCTIVITY PRAIRIE FALCONS IN NORTHEASTERN COLORADO

24 of 34 sites (70%) were determined to be active.
19 (79%) of the active sites were known to have produced young.
Total number of young produced by
Total adjusted productivity

14 sites = 55 (3.92 young/site)

of 19 sites = 74

Total number of young fledged from 18 sites = 53 (2.94 per site).

�-149-

TABLE 5

ACTIVITY

Site No.

AND PRODUCTIVITY

OF FERRUGINOUS

Activity

HAWKS IN NORTHEASTERN

No. of Young

No. Fledged

0

0

3

3

LO 1
WE 1

IA

WE 2

IA

WE 3

A

WE 4

IA

WE 5

IA

WE 6

IA

WE 7

A

WE 8

IA

WE 9

IA

WE 10

IA

WE 11

A

0

0

WE 12

A

0

0

WE 13

A

0

0

WE 14

IA

WE 15

IA

WE 16

IA

WE 17

A

0

0

COLORADO

Connnents

�-150-

TABLE 6
SUMMARY OF PRODUCTIVITY OF FERRUGINOUS HAWKS IN NORTHEASTERN COLORADO
6 of 18 (33%) of the sites were active.
1 (16%) of the active sites produced young (3.0 young/site).
1 (16%) of the active sites fledged young (3.0 young/site).

�-151-

JOB PROGRESS REPORT

State of __~C~o~l~o~r~a~d~o~
Project No.

W-124-R-3

Work Plan No.
Job Title:

_
Raptor Investigations

1

Job No.

3

Raptor Population and Productivity Studies in Southeastern Colorado

Period Covered:
Personnel:

March 1, 1974 through February 28, 1975

Gerald R. Craig, William C. Andersen

ABSTRACT
Occupied nests of nine golden eagles, 38 ferruginous hawks,
nine red-tailed hawks and 176 Swainson's hawks were located
in a 4,421 square mile area in Southeas t ern Colorado.

Data

are provided about fledging success and nest density for each
of the four raptor species.
trend counts were continued.

Weekly migration and winter

��-153RAPT OR POPULATION AND PRODUCTIVITY STUDIES IN SOU7HEASTERN COLORADO
William C. Andersen and Gerald R. Craig
To intensively document nesting and wintering denSities, migration chronology, habitat requirements, productivity and prey preferences of raptors on the
Comanche National Grasslands and adjacent prairie landH.
SEGMENT OBJECTIVES
1.

To continue documentation of nesting densities and productivity of golden
eagles, prairie falcons, ferruginous hawks, Swainson's hawks, red-tailed
hawks, great horned owls, and burrowing owls on the Comanche National
Grasslands and adjacent prairie lands.

2.

To continue documentation of the timing of breeding cycles of golden eagles,
prairie falcons, ferruginous hawks, Swainson's hawks, red-tailed hawks,
great horned owls and burrowing owls.

3.

To continue documentation of prey items at nest sites.

4.

To investigate and categorize nest sites by such features as disturbance
factors, habitat type, human use, dominant vegetation, relief, etc.

5.

To continue documentation of migration chronology.

6.

To continue to gather trend information on wintering raptor populations.

7.

To categorize important wintering habitats.
RESULTS
Nine golden eagle nests were located of which two succeeded in fledging

3 young which provided an average of .33 young per nesting attempt.

Thirty-

eight nesting attempts by ferruginous hawks produced 72 fledglings for an
average of 1.89 young per nesting attempt.

This was a slight increase in pro-

ductivity from 1973 (1.47 young per nesting attempt).

Nine red-tailed hawk

nests succeeded in fledging 18 young (2.00 young/nesting attempt).

Swainson

hawk productivity increased dramatically with 176 nests producing 191 young

�-154(1.08 young/nesting

attempt).

This was due to the lack of severe wind and

thundersqualls which were prevalent in 1973.

In all, 328 raptors (including

191 Swainson's hawks, 72 ferruginous hawks, 22 red-tailed hawks, 19 great
horned owls and 13 kestrels) were banded.
The migration and winter trend counts were again run in 1974-75, although
vehicle difficulties and weather conditions did not permit the trends to be
run in February and March.

�-155TABLE 1
ACTIVITY

MlD

NESTING SUCCESS

- 1974

Nest
Number
Attem:Qts Fledged

S:Qecies

County

Golden Eagle

Baca
Bent
Las Animas
Otero
Lincoln

2
2
3
1
1
9

Ferruginous Hawk

Baca
Bent
Cheyenne
Crowley
Kiowa
Las Animas
Lincoln
Otero

9
5
1
3
2
3
9
6
38

Yng/Nest Active
NestL10Omi2
AttemEt

Yng/pr~duc.
/lOOmi

0
2
1
0
0
3

0.00
1.00
0.33
0.00
0.00
0.33

0.21
0.39
0.51
0.12
0.15
0.25

0.00
0.39
0.17
0.00
0.00
0.08

13
1
2
2
5
4
26
72

1.44
0.20
2.00
0.66
2.50
1.33
2.88
2.17
1.89

0.94
0.99
0.59
0.96
0.61
0.51
1.37
0.71
0.78

1.36
0.19
1.19
0.64
1.55
0.69
3.94
1.54
1.66

-1L

Red-tailed Hawk

Bent
Las Animas
Otero

2
4
_3_
9

3
9
6
18

1.50
2.25
2.00
2.00

0.21
0.69
0.36
0.45

0.59
1.55
0.71
0.93

Swainson's Hawk

Baca
Bent
Crowley
Elbert
Kiowa
Las Animas
Lincoln
Otero

57
5
26
3
28
3
46
8
176

90
3
24
4
31
1
28
10
191

1.59
0.60
0.92
1.33
1.11
0.33
0.61
1.25
1.08

5.98
0.99
8.36

9.45
0.59
7.72

8.66
0.51
6.98
0.94
4.21

9.59
0.17
4.25
1.18
4.58

�-156-

TABLE 2
RAPTORS BANDED IN SOUTHEASTERN COLORADO

Species

Number Banded

Swainson's Hawk

191

Ferruginous Hawk

72

Red-tailed Hawk

22

Great Horned Owl

19

American Kestrel

13

Barn Owl

4

Golden Eagle

3

Cooper's Hawk

3

Mississippi

Kite

1

TOTAL

328

�TABLE 3

MIGRATION AND WINTER POPULATION

TRENDS, 1974-1975

9

November
16

30

7

December
14

23

January
4

(Adult)

(-)

( 4)

( 7)

( 5)

( E)

( 7)

( 4)

(Immature)

(-)

( 1)

( 1)

( -)

( 1)

( 1)

( 4)

(Unknown)

(1)

( -)

( -)

(-)

( -)

(-)

( -)

Total

1

5

8

5

7

8

8

(Adu1 t)

(2)

( 4)

( 6)

(11)

(12)

(28)

( -)

(Immature)

(-)

( -)

( 1)

( 2)

( 2)

( 3)

(12)

Species
Golden Eagles

Bald Eagle
I

I-'

VI

2

Total

4

7

14

13

31

12

Ferruginous

Hawk

13

10

6

10

12

5

12

Rough-legged

Hawk

9

7

8

13

14

8

8

8

7

7

6

11

2

6

Marsh Hawk

4

5

4

2

7

5

Prairie Falcon

2

2

4

9

7

2

5

Merlin

-

-

-

-

-

-

2

Kestrel

12

9

12

16

16

8

13

Great Horned Owl

-

1

1

2

1

-

1

TOTAL

51

50

57

76

89

69

67

RAPTOR/MI.

.34

.33

.38

.51

.59

.46

Red-tailed

Hawk

-

I

.45

"
I

��-159-

JOB PROGRESS

Sta te of __

REPORT

-...;C:.,:o;,,:l:..:o:...,:ra:.=..:d::.:o=---_

Pro j e ct No. _W.:.:-=.1
=.24.:...-...,:R:.:,.-...,:3::_
Raptor
Work

Plan No.

Job Title
Period

Roadside

Covered:

Personnel:

~l

_

Raptor

March

Gerald

Investigations

Job No, 4

Censuses

1, 1974 through

February

28, 1975

R. Craig

ABSTRACT

Two migration
Weld

County

concerning

and winter

and the other
densities,

censuses

werE

in El Paso County.

behavior,

and habitat

conducted;

one in

Data are presented
preferences

of raptors.

��-161ROADSIDE RAPTOR CENSUSES
Gerald R. Craig, Babette Cranson and Patricia Kennedy
To continue roadside censuses currently in existence in
Northeastern

Colorado to obtain information on population fluctuation

and chronology of raptor movements.
SEGMENT OBJECTIVES
1.

Each route is established along lightly traveled dirt
roads.

2.

Each route is driven in a vehicle traveling between
20 and 35 m.p.h.

3.

All raptors observed with 1/4 mile cf either side of the
route are counted. This will provide an area as well
as a linear census.

4.

All pertinent information, such as sex, age and behavior
of the raptors, and the habitat over which it was observed
is recorded.

5.

Each route is traveled on a bimonthly basis throughout
the year.

6.

One census route runs in a northeasterly direction from
Riverside Reservoir to Sterling, Colorado, and the other
route travels north from just east of Greeley to Herford,
Colorado.

7.

Prepare monthly summary of results and furnish copies of the rough
data gathered.
EXPLANATION

In the winter of 1974-75, two census routes were established, one
in Weld County and the other in El Paso County.

Due to difficulty in

locating a person capable of running the second census in northeastern
Colorado, it was necessary to take over an old route southeast of Colorado
Springs in east-central Colorado.

The results for each of the censuses

are presented separately as follows:

�RAPTOR MIGRATION AND WINTER POPULATION TRENDS IN NORTHEASTERN

COLORADO 1974-75

by Babette Cranson
METHODS AND MATERIALS
Ten censuses were made on a route of approximately
County.

135 miles located in Weld

This route was established and censused in the winter of 1969-70 by

G. R. Craig.

The route was traveled at speeds of 20 to 35 miles per hour.

Binoculars and a spotting scope were used to identify those raptors that were
sighted from the vehicle.

Species, age, sex, activity, location and habitat were

recorded for each observation.
RESULTS AND DISCUSSION
A total of 273 raptors were observed on the censuses which began November 30 and ended March 7.

The counts were started on a bi-monthly basis with

the December 29th count missed.
weekly basis.

Beginning January 10, counts were made on a

Table 1 summarizes these censuses.

Fifty-two percent of the 273

raptors sighted were rough-legged hawks, 14% were golden eagles, 10% were marsh
hawks, 7% were American kestrels, 6% were prairie falcons, 4% were ferruginous
hawks, 2% were bald eagles and 1% were merlins.

Peaks in migrating raptors are

hard to assess from the data since low counts often result from high winds.

How-

ever a slight increase of raptors can be noted on the January 31 count.
Table 2 summarizes the behavior of raptors observed on the census routes.
Of the raptors Sighted, 67% were perched and 61% of those perched were on power
poles.

Trees were the next most preferred perch site with 20% of the raptors

perched there.

Seven percent of the rap tors were perched on fence posts, 4% were

perched on phone wires, 4% were perched on the ground, 3% were perched on windmills and 1% were perched on low vegetation.

Power poles and trees had the most

varieties of raptors perche~ on them with 9 and 7 of the 10 species perched on
each respectively.

Only four of the 10 species were perched on fence posts, 3 of

the 10 species were pe:rched on the ground and on windmills, 2 of the 10 species

�-163-

were perched on phone wires and only 1 of the 10 species was perched on low
vegetation.
Habitat preferences are summarized in Table 3.

Of the approximately

230

square miles covered by the censuses, approximately 47% of the habitat was
agricultural, 46% was grassland and 5% was cottonwood-riverbottom.
habitat areas accounted for the remaining 7%.

The other

Sixty percent of the raptors were

observed in the agricultural areas while 30% were observed in the grassland
habitat and only 7% were observed in the riverbottom-cottonwood

habitat.

The

agricultural and grassland areas supported the most species with 9 and 7 of the
10 species recorded for each respectively.

�-164TABLE 1

MIGRATION AND WINTER POPULATION TRENDS 1974-75

Golden Eagle
(Adu1 t)

Nov.

December

30

8

1511 22

6

(Innnature)

1

-

(Unknown)

1

(Total)
Bald Eagle
(Adult)

January
29

3

lOll 25

31

7

-

1

2

4

-

-

-

1

-

2

+

-

-

-

-

-

-

8

-

2

+

-

-

-

-

-

-

-

-

-

-

(Total)

-

-

-

(Innnature)

-

-

-

-

-

-

-

Ferruginous

Hawk

2

-

1

Rough-legged

Hawk

18

5

Red-tailed Hawk

1

Marsh Hawk

6

Prairie Falcon

4

Merlin

1

Kestrel

5

-

Great Horned Owl

-

-

1

-

1

-

-

-

February

-

-

-

-

-

14

7

2

5

-

4

-

3

-

2

1

2

2

-

1

1

-

-

2

8

2

8

2

4

2

1

2

-

1

-

-

1

-

3

-

-

-

-

-

3

-

1

1

9

19

22

21

-

-

-

1

1

-

5

-

3

-

2

19

2

11

17

-

-

3

2

-

1

4

8

-

1

4

-

2

2

-

1

-

-

-

-

1

1

4

2

3

2

-

-

3

1

-

2

-

1

-

-

-

2
45

Raptors/mi.

.33

II High Winds

-

10
.07

-

-

-

2

-

Unknown
TOTAL

March

2411 28

2

1

13

32

41

35

36

5

21

40

.10

.24

.30

.26

.27

.04

.16

.30

�-165TABLE 2

RAPTOR BEHAVIOR, 1974-1975
Activity

BE

GE

FH

RT

RL

MH

PF

AK

M

UNK

TOTAL

Flying (flapping)

1

4

2

0

27

24

2

4

0

0

64

Soaring (circling)

1

8

0

0

10

3

0

1

0

2

25

Hovering

0

0

0

0

1

0

0

0

0

0

1

2

12

2

0

38

27

2

5

0

2

90

Trees

4

2

2

5

19

0

1

2

1

0

36

Power Poles

0

10

4

0

78

0

12

6

1

1

112

Ground

0

5

0

0

1

1

0

0

0

0

7

Fence Post

0

6

1

0

4

0

0

0

1

0

12

Windmill

0

4

1

0

1

0

0

0

0

0

6

Bush

0

0

0

0

2

0

0

0

0

0

2

Phone Wire

0

0

0

0

0

0

1

7

0

0

8

4

27

8

5

105

1

14

15

3

1

183

6

39

10

5

143

28

16

20

3

3

273

Subtotal
Perched

Subtotal
TOTAL

�-166-

TABLE 3
HABITAT PREFERENCES

Habitat type

BE

GE

FH

RT

RL

MH

PF

AK

M

UNK

TOTAL

Agriculture

2

19

2

0

82

25

12

16

2

3

163

Elm - Agricul ture

0

0

0

0

1

0

0

0

1

0

2

Elm - Meadow

0

0

0

0

0

0

0

1

0

0

1

2

0

0

74

Grassland - Pasture

4

16

4

0

42

3

::;

Grassland-sage-pasture

0

3

2

0

2

0

0

0

0

0

7

Grassland-tall

grass

0

0

0

0

1

0

0

0

0

0

1

Grassland-deciduousconiferous

0

0

0

5

2

0

0

0

0

0

2

Riverbottom-cottonwood

0

1

2

0

9

0

1

0

0

0

18

Cottonwood-lakeside

0

0

0

0

2

0

0

0

0

0

2

Cottonwood-agriculture

0

0

0

0

0

0

0

1

0

0

1

Sage-willow

0

0

0

0

2

0

0

0

0

0

2

6

39

10

5

143

28

16

20

3

3

273

TOTAL

�-167RAPTOR MIGRATION AND WINTER POPULATION TRENDS IN EAST-CENTRAL

COLORADO

1974-1975

by Patricia Kennedy

METHODS AND MATERIALS
Areas southeast of Colorado Springs were invem':oried during the fall of 1974
and an 85 mile route that mainly parallels telephone transmission lines, was
established

(Figure 1).

Eight censuses were conducted along this route between

late September and mid-February.

The route

was censused by car in the mornings

and, with the exception of one census (January 9), censuses were conducted only
when the winds did not exceed 15 mph.

The route was driven at speeds between

30 and 40 mph, and all raptors within a quarter mile of the road were identified,
aided by binoculars and a spotting scope.
each observation.

Age

Species and habitat were recorded for

class and sex were recorded when feasible.

was reported for the last 5 censuses.

Activity

A horned lark count was also made along

a ten mile section of the route (Figure 1) and all larks observed were recorded
on a hand tally.
RESULTS AND DISCUSSION
A total of 181 raptors w~re observed, and of these 17% were rough-legged
hawks, 16% were American kestrels, 12% were prairie falcons, 12% were golden eagles,
11% were ferruginous hawks, 10% were marsh hawks, 8% were red-tailed hawks, 5% were
unidentified

raptors, 2% were me:r1ins and 1% were Swainson's hawks (Table 1).

The

highest raptor densities occurred in late October through early November indicating
a possible peak period in the fall migration.
in the area after October 12.

Swainson's hawks were not observed

Red-tailed hawks had left the area after November 1.

This indicates that Swainson's and red-tailed hawks are not found in the eastern
Colorado short-grass prairie during the winter months.

However, if the census had

traversed a riparean habitat, red-tailed hawks probably would have been observed.
Rough-legged hawks were not observed until November 1 and were present throughout
the winter in large numbers.

Merlins were not observed until January 9, but due

to the small number observed and lack of a December census, a migration trend for
for this species is difficult to assess.
year-round residents.

All other raptor species appear to be

�TABLE 2
RAPTOR BEHAVIOR

S ecies

Activit

Marsh
Hawk

G.Horned
Owl

Merlin

9

0

0

0

0

57

0

0

0

0

0

0

2

0

0

0

0

10

0

0

13

0

0

0

0

0

0

0

0

1

0

0

1

0

0

0

2

0

4

1

3

0

0

2

0

0

0

6

1

0

0

0

0

0

5

1

1

13

3

1

101

Ro.Legged
Hawk

Ferrug.
Hawk

10

21

10

2

0

0

Tree

3

0

Windmill

1

Fence Post

1

American
Kestrel

Golden
~le

Tel. Pole

7

Phone Wire

Prairie
Falcon

Unknown

Total

Perched

Ground

3

Soaring
Flying (flapping)
TOTAL

:1.4

1

0

2

0

4

5

0

14

27

12

13

7

10

I
I-"

.....•
0
I

�-171-

TABLE 3
PREY SPECIES TREND

Larks/mile

No. Mer1ins &amp; Prairie
Falcons Observed

Date

No. Horned Larks
Observed

9/26

145

14.5

1

10/12

202

20.2

2

11/1

271

27.1

5

11/29

97

9.7

5

1/9

60

6.0

1

1/11

84

8.4

3

2/2

77

7.7

5

2/16

348

34.8

3

�-172FIGURE 1

Larks/mile
= - No. of Falcons =
Numbers
Observed

35~-----------------------------------------------,,•
301~--------------------------------------------~---,,,
25------------~--~,------------~--------------~,~---~~
,
/
,,
20~-----,~~--------~,----------------------------~,------.:
I
,
,
l5,~~----------------~----------------------~------,
,
10~----------------------~\~--------------------~-------\

"

\

I

\

I

\

••••••
...,

...•.•

1'''

..•.•.... "

5

Dates

"'•••.-..-. •••• I

••

�-173JOB PROGRESS

REPORT

State of__ .--:C::.:o::.::l:..;o::.:r:..;a::.;d::.;o~
~
_
Raptor Investigations
Pro j ect No. _W~-.:.1=-24~-~R::..--,,3,-- _
Job No. 1
Work Plan No.
~2~
_
Job Title:

Peregrine

Period Covered:
Personnel:

Falcon Propagation

March

and Reintroduction

1, 1974 through February

Studies

28, 1975

Gerald R. Craig, James H. Enderson

ABSTRACT
Thirty-eight

eggs were laid by four female anatum peregrines

of which

14 eggs were fertile,

five hatched

and five young were reared.

Improper

incubator

and excessive

humidity

temperature

the cause for poor hatching
potential

for increasing

needs improvement.
augment

success.

Artificial

insemination

shows

egg fertility

although

the technique

still

All five young produced

captive breeding

may have been

stock.

in 1974 were retained

to

��-175PEREGRINE FALCON PROPAGATION AND REINTRODUCTION
James H. Enderson and Gerald R.

Craig

To captively produce eggs and/or young from peregrine
into

STUDIES

falcons for reintroduction

the nests of wild peregrine falcons that have low or no natural productivity.
SEGMENT OBJECTIVES

1.

To establish the cause for reproductive

failure of wild peregrines.

2.

To produce young or eggs from peregrine falcons being held in captivity at
the facilities of Dr. James H. Enderson.

3.

To visit wild peregrine falcon eyries where adults are present and replace
spoiled eggs with captively produced eggs or young.

4.

To keep those eyries containing captively produced eggs or young under
observation to ascertain the success of the endeavor.
SUMMARY OF RESULTS

Falcons in the project in 1974 included the following,
Peregrines:
3 pairs adult anatum peregrines
1 adult pair, anatum female and tundrius male peregrines
1 pair adult peale~ peregrines
1 adult anatum male
1 adult tundrius female
3 yearling Fl anatum females
1 adult anatum male (on loan to R.

Berry)

Prairie Falcons:
1 pair adults (inclJding an Fl male)
1 pair Fl adults (on loan to E. Freienmuth )
1 adult female (on loan to E. Freienmuth)
1 pair Fl yearlings

(on loan to U. S. Air Force Academy)

The four anatum females laid a total of 38
eggs and three which were broken by the birds.

eggs, including two malformed
Of the 38 eggs, 14 were known

�-176-

fertile, and five hatched and developed normally.
The Peale's falcon pair laid seven eggs in two clutches, two of which were
fertile by artificial insemination and a third probably so.

None of these

eggs

hatched although one developed to the point of hatching.
The technique of placing captively produced young in nests of wild falcons
was experimentally

tested with prairie falcons.

Twenty-one captively produced

prairie falcons, ranging in age from ten days to three weeks of age were flown
to Colorado from propagating

facilities at Cornell University.

Unfortunately

four of the young succumbed to overheating in the baggage compartment of a commercial airliner.

One of the young falcons was held for release at a later date,

and fifteen were placed in eyries of wild prairie

falcons.

One of the chicks

at a particular site died during the night when it fell from the nest ledge.
that site, the wild adult pair of falcons had

At

experienced nesting failure during

incubation when all their eggs were apparently removed by unknown persons.

The

adults still occupied the nest cliff and frequently perched on the ledge of the
eyrie.

The thought was to place the young prairie falcon chick on the nest ledge

in hopes that the adults would care for it.

The single chick apparently became

lonesome and wandered about the ledge, then fell before the adults were aware of
its presence.

This technique should be tested again since it may be useful if

wild peregrines are unsuccessful

in rearing young but are still occupying the site.

Captively reared young prairie falcons were placed in wild prairie falcon
eyries containing young.

In one case, the natural brood size of four young was

increased to seven and all fledged successfully.

At another site, an age disparity

of two weeks existed between the young, but the adults continued to rear the
younger chicks after the older young had fledged.

It was concluded that

�-177-

wild prairie falcons appear to be quite adaptable to manipulation

of brood sizes

and will accept captively produced young with little hesitation.
Of the total of 16 peregrine eggs known

fertile, only five hatched.

Of

the remainder, three were incubated by the Peales falcons and apparently received
inadequate incubation, five died at hatching after artificial incubation at
excessive humidity or temperature, or both.

The remaining three failing to

hatch died in the incubator midway in development.
The five hatched young were reared artificially by means of a hand-puppet
simulating an adult peregrine.
sight of people.

All

The puppet was used to feed the young out of

five young were fledged together in a loft and remain

there.
DISCUSSION
The disappointing results of the 1974 season are related to two unresolved
difficulties.

Only one pair of the anatum falcons copulate; nine fertile eggs,

in three clutches, resulted which also included three broken eggs.
fertile eggs were by artificial insemination.

The remaining

The success ratio of that tech-

nique was poor, only seven, probably eight out of a possible 21.

Discouraged

by this poor success, and similar difficulties in his project,

R. Fyfe dissected

a pair of breeding prairie falcons and discovered that we had been placing the
semen in the cloacal bursa of the female, not in the oviduct.

More dissections

will be done before the 1975 season to correct the imsemination procedure.
Problems were encountered with artificial incubation when eggs were run at
99.50 F. and 50-55% relative humidity.

Four chicks dying

at pipping of the

shell had large unabsorbed yolk sacs, indicative of excessive temperature or
humidity, or both.

Both R. Fyfe, and T.

Cade at Cornell, hatched peregrines

readily when conditions were about 98.50 F. and 45% relative humidity, values
which are planned for use in this project in 1975.
In summary, results i.n1974 were only fair, but were an improvement over
1973.

Four pairs of anatum peregrines have been produced

thus far.

I intend

�~178that two Fl pairs will remain here, and that two pairs will be lent to other
projects.

Assuming that this year's problems can

be resolved, a substantial

improvement in production is expected in 1975, and the intent is to place some
eggs or young in the eyries of wild peregrines.
Soon after the work with prairie falcons was completed, there was an opportunity to test the technique on wild peregrines.

One wild peregrine eyrie was

surveyed and found to contain two infertile eggs and the remains of a third egg.
The infertile peregrine eggs were replaced with two fertile eggs which had been
removed from a prairie falcon nest.
one of the prairie falcon eggs.
of age, it was removed

The wild peregrines succeeded in hatching

When the prairie falcon chick

was three weeks

and replaced with two captively produced peregrines which

were ten days of age and had been flown to Colorado from Cornell University.
The wild peregrines proceeded to rear and fledge both peregrine chicks without
complication.
RECOMMENDATIONS
Due to difficulty encountered in obtaining and hatching fertile peregrine
eggs in captivity, several more years will be required before sufficient numbers
of young peregrines are produced to permit an extensive reintroduction program.

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                  <text>-1-

April

1976

JOB PROGRESSREPORT

State

of

Project

COLORADO
----------~~~~-----------

Work Plan
Job

Pheasant

Covered:

Personnel:

1

No.

Title

Period

Game Bird

W-37-R-29

No.

April

Job No.
Hortality

1,

Survey

21

--------------------~--

Investigation

1975 to Harch

31,

1976

Donald 11. Hoffman, Jack Corey, Lucien
Daniel Walsworth and Warren Snyder.

Brevard,

Larry

Crooks,

ABSTRACT
Phe as ant; populations
dropped to reco r d Lows in the study area and surrounding
:Legion in 1975.
Crowing indices
were 68 percent
Lower than those obtained
in
1974 and 84 percent
below 1963-68 levels.
Triangulation
methods indicated
most of the study contained
less
than a roos ter per square mile, whereas
the southeast
corner contained
up to four roosters
per square mile.
Limited
sex ratio
data obtained
in April and again in fall
showed approximately
3.4
hens per adult
rooster.
The birds-per-mile
index of 0.38 derived
from August
production
counts was only about a fifth
of the level obtained
in the preceding
year.
Trapping efforts,
which extended
from mid-August to late October
reflected
the low population
density.
Only 109 roosters
were banded along
with four retraps.
In addition,
150 hens were banded and 7 retraps
were
taken.
Young per adult ratios
were higher
in both production
counts and fall
trapping
indicated
some improvenent
in reproductive
success
over the preceding
year.
The·1975 pheasant
season opened on Wednesday November 19 in conjunction
with a heavy snow wh.ich closed most roads reduced hunter effort
and bunched
pheasants
so that they were highly vulnerable
to road hunting.
Heavy snow
cover persisted
throughout
the short twelve day season.
Thirteen
reward bands
wer e obtained
from fall banded birds,
projecting
an approximate
23 percent
harvest.
Four regular
bands were obtained
by field
contact
of hunters
and
one hunt.e r returned
a regular
band by Hail several
months after
season closure,
bringing
the number of 1975 fall
trapped
regular
band recoveries
to five.
High movement of birds
from banding to harvest
was evident
in repetition
of
1974 results.
Some natural
mortality
was reported
as a result
of the November
snows t orm, but its
impact could not be accurately
assessed.
Late winter
sex
ratio
counts showed an excess of three hens per cock surviving
the winter,
w i th very low over-win tering
populations
present.

��-3-

PHEASANT MORTALITY

INVESTIGATION

Warren D. Snyder

P. S. OBJECTIVE
To investigate

mortality

rates of a population

of ring-necked

pheasants.

SEGMENT OBJECTIVES
1.

To review relevant literature
ity and survival of birds.

concerning

2.

Monitor

population

3.

Develop

new and improve on existing

4.

Investigate mortality
individuals.

5.

Prepare

pheasants

and studies of mortal-

status on study area.
trapping

techniques.

factors and estimate mortality

annual progress

rates of marked

report.

METHODS

AND MATERIALS

Reference is made to Hoffman (197.5) concerning methods used in this study.
Deviations or additions to these are summarized as follows.
Spring sex ratio counts were attempted, but study area pheasant populations
were too sparse to permit attaining an adequate sample so efforts were discontinued.
Limited data were obtained in early April under snow cover conditions.
Two 10-station crowing routes were established to distribute sampling throughout the four township study area (Fig. 1). The starting and ending locations
on these routes were varied to partially correct for call variance relative
to time within counts.
Efforts were made to tally total calling males on forty square miles within
the study area. Four square miles surrounding crowing stations numbers 1, 3,
5, 7, and 9 on each of the two routes were checked using triangulation
methods (Fig. 2). Two men in separate vehicles with radio communication pinpointed each male during early A.M. calling periods.
Repeat counts were made
on different mornings to increase accuracy.
Four production routes were censused with two rep1i cates on each being comp1eted during August 1975 (Fig. 3). Pheasant sightings were segmented at
2 mile intervals along the routes to pinpoint potential trapping areas.
Because of low populations, pheasant trapping was extended to locations outside the study area during the fall. Winter trapping was not attempted
because of lack of birds.
Late winter sex ratio counts were also extended
to locations south and east of the study area to increase sample size.

�-'t-

t 7 N.

(.9

o
.J

f-\.46\V~YUMA

Fig. 1. Pheasant crowing count routes with listening
the pheasant mortality study area in 1975.

stations

COUNTY
established

in

�-5-

-r---'.
.

6

&gt;IZ

-

::&gt;
'7

0
U
:=:~-

.

to'

.

.
":J
.
.
.
.

5

~
;::

•

.

9

:

I~~-/-Y;Lr&lt;
L--.~
T. 7 N
19

20

Z

&lt;X:
C)

o
,

-1

·T. 6 N

R.47VJ.

HA6 W

II

YUMA

COUNTY

blocks censused within the study are·a- during---th-e-s-p-Hn--g-------·
Fig. 2. Triangulation
Numbers
represent
the total roosters recorded per four square mile
of 1975.
1_ 1 __

,_

�-6-

RESULTS AND DISCUSSION
Pheasant
Pheasant

.10

Crowing

Population

Monitoring

Counts

The impact of the snow storm of March 27, 1975 whichkilled many of the
pheasants, was verified by crowing counts in late April and May.
Results
from crowing count routes (Fig. 1), illustrated in Table 1, show an overall
station 'average of 6.12 calling males per two minutes for the east route,
and 2.04 calling males per station on the west route. High count per station
averages were 10.8 on the east route, and 3.8 on the west route.
These are
somewhat comparable to high count per station indices obtained in previous
years (Table 2), but are biased upward for two reasons.
First, starting
points per route were rotated in 1975 so the peak calling rate was potentially
distributed to all stations not just the middle stations as happens when each
replicate is started at station number one.
Second, conducting more replicates, as made in 1975, tends to increase the high count per station mean.
Even with these biases the 1975 spring crowing count was 68.3 percent below
1974 data and 83.9 percent below 1963-68 data in the study area (Table 2).
This decline of 68 percent from 1974 to 1975 for the most part is attributable
to decimation of the population by the March blizzard, but poor reproduction
due to drought conditions in 1974 also may be a contributing factor.
Triangulation

Inventory

Crowing counts yield an index of rooster density, and when combined with sex
ratio data give a clue to relative breeding hen density among years or areas.
Triangulation methods were employed in 1975 to relate crowing indices to
actual pheasant abundance (Fig. 2). Results were believed to be closely
proximal to the actual population since rooster densities were very low.
Pheasant roosters do not crow consistently, so there is always the chance of
misSing a bird.
Counts were repeated on subsequent mornings to correct for
this.
In a few instances this may have resulted in a higher than actual
tally due to a shift in territory by a rooster so that he was tallied more
than once.
The triangulation methods indicated that roosters can be heard for over a
mile in distance under certain conditions, but may not be detected at even a
half mile in other circumstances.
The primary variables are wind and the
direction the birds are facing.
Data illustrated in Fig. 2 show there were almost no roosters present in the
northwest part of the study area, with two sample blocks (8 square miles)
containing no roosters.
Blocks in the southeast township were much higher.
This differentialin
density was primarily due to the greater severity of the
March blizzard in the northern part of the study area.
Projections based on data in Fig. 2 indicate that if 63 pheasant roosters were
present in 40 square miles (27.8 percent of the total area), then approximately
227 pheasant roosters were present in the total 144 square miles or there were
1.58 roosters per square mile.
In general, the northwest half of the study
area averaged less than a rooster per square mile.
The eight square miles
sampled in the southeast township contained 34 roosters in comparison to
only 29 roosters in the remaining 32 square miles sampled.

�-7-

&gt;-

'.

I-

z -:J

o

10

9

(J

:

T. 7 N,
19:

~l/~v~

&lt;,
20

,,:.

...•..

&lt;,... 21 ;,-

.
~

:"

For,', _

F(ett~~.
~-"23.

v--,)' :
24

•••
:
•

YUMA

Fig. 3.

a. \.

•
20

21

'\
II

2'

II
•

II

c OU I'J T y

Pheasant production routes established and censused during August 1975.

�-8-

Table 1.
routes.

1975 pheasant

Date

1

2

crowing

indices

3

Station
5

4

obtained

Number
6
7

on the east and west study

8

9

10

Mean

24

4

11

8

6.7

14

27

6

10

9

8.3

4

8

11

4.2

Y 4

4

7

2.5

East Stud;::Route
4-22

1

3

1

1

:J-/ 11

4-22

5

4

1

3

41./

5-1

11:./

1

3

0

0

5

9

5-8

1

0

1

3

1

1

3

5-18

31/ 4

4

2

3

11

14

8

15

12

7.6

6-1

1

0

5

8

6

17

13

9

11

41/

7.4

High/stat.

5

4

5

8

6

17

27

9

15

12

Overall

Average

10.8
6.12

West Stud;::Route
4-24

0

4-30

01:./

5-1

oy

01-/

1.5

0

0

1.5

2

4

1

2.3

1

2

4

1

2.2

41/ 0

2

5

2

0

2.6

0

4

0

2

2l/

1

1

2.2

4

2

2

1

4

4

0

1

2.0

6

3

5

2

4

5

4

1

3.8

1

6

0

3

1

1

3

0

1

2

1

5

2

1

3

8

3

2

1

1

1

5-1

01/ 5

4

3

1

1

5-8

0

7

6

0

5-14

0

7

5

5-30

al/

2

High/stat.

0

8

Overall

Average

1/ Location

on route where

2.04

count was started.

�-9Table 2. Pheasant
pheasant mortality

crowing counts as indices of population
study region.

Interval

Route Name

1963-68

Cent ral con trol

1974

Wages-Haxtun

and study route

change in the

High Count/Station

45.3

}

Percent

Change

-49.2%
-83.9%
-68.3%

1975

East and West study routes

1/ Starting

points were rotated around the routes in 1975 so high count per
station data are skewed slightly upward in relation to counts made in previous
years.
The mean in 1975 was 6.12 on the east route and 2.04 on the west
route.

Pheasant

Production

Counts

A combined total of 15 replicates were completed on four census routes (Fig.
3) in August, 1975. Only 0.38 pheasants per mile were observed as shown in
Table 3. Comparisons with census in previous years indicates a very low
pheasant population present in 1975 (Table 4). Results were only a fifth of
the index level obtained in 1974 and only a seventh of that obtained in the
late sixties.
In general, results showed the 1975 hatch was later than
average with a fairly high proportion of unsuccessful hens.
Pheasant

Sex Ratios

Late Winter to Summer Breeding Population in 1975--Pheasant sex ratio observations, reported by Hoffman (1975), indicated approximately 4.44 to 4.97
hens per cock were present in late winter 1975. This proportion of hens was
greater than that found during subsequent samples in spring and late summer
(Table 5). Although all samples were small, post-blizzard ratios obtained
in April and during fall trapping were similar, indicating between 3 and 3.5
hens per cock were present during the breeding season.
Using 3.44 hens per
rooster as the spring sex ratio, triangulation data project 5.4 hens per
square mile or 781 hens present in the 144 square mile study area.
Fall 1975 to Late Winter 1976--Data presented in Table 5 show there were
approximately 1.38 to 1.46 hens per cock present in the prehunting season
population.
The mid-November sample was obtained with heavy snow conditions
present prior to onset of significant harvest removal.
Late winter sex
ratios were primarily obtained after a 3-4 inch snow in late February.
Efforts to observe birds under bare ground conditions were futile due to
extremely low densities.
A large portion of the late winter sample was
obtained up to ten miles or more south of the study area in Yuma County and
in other areas to the east and southeast to increase sample size.

�-10Table 3. A summation of pheasant production data on the four census
routes
within the mortality study area.
Route

Replicate

Miles

Cocks

Hens

Young

Unclass.

Total

Broods

East open

3

78

0

6

30

3

39

6

East closed

4

104

1

8

34

0

43

6

West open

4

104

3

6

19

0

28

3

West closed

4

104

0

6

33

0

39

5

Total

15

390

4

26

116

3

149

20

Young/mile

0.30

Young/hen

Birds/mile

0.38

Broods/mile

=

0.051

= 4.46

Table 4. A comparison of pheasants observed per mile during production counts
in the mortality study area.
Interval

Miles

Birds Observed

Birds per Mile

1966-68

515

1,383

2.69

1974

189

355

1. 88

1975

390

149

0.38

Within the study area, 45 cocks and 153 hens were tallied, projecting a ratio
of 3.40 hens per rooster. There were 74 roosters and 220 hens recorded outside the area yielding a ratio of 2.97 hens per rooster. Combination of the
two samples projects a ratio of 3.13 hens per rooster in a total sample of
492 pheasants (Table 5).
Assuming 1.38 hens per cock as representative of the fall 1975 sex ratio, and
3.13 hens per cocks as the late winter 1976 sex ratio level then the number of
cocks per 100 hens would change from 72.4 in fall to 31.9 in late winter.
This would mean that 40.5 roosters, 55.9 percent, were lost from fall trapping

�-11-

to late winter in excess of the hen loss for the same interval.
Band return
information indicates that at least 24.7 percent of the original population was
removed by hunters, assuming that all reward bands were reported.
A greater
proportion, 31.2 percent, of the differential rooster loss was attributed to
other causes. No consideration is given in the above projections to hen loss
due to hunting.
While some is known to occur, the percentage of the total
population is believed small, based on the low percentage rooster kill.

Table 5. Pheasant sex ratios obtained from late winter,
winter, 1976 on, and proximal to, the pheasant mortality

Sample

.-

Roosters

1975 through
study area.

late

Hens

Total

Percent
Males

Percent
Females

Ratio

Late winter 1975,
observed 1/

36

179

215

16.7

83.3

1:4.97

Late winter
trapped

32

141

173

18.5

81.5

1:4.44

April 1975, observed

34

117

151

22.5

77 .5

1:3.44

Fall 1975, trapped,
adults

19

63

82

23.2

76.5

1:3.32

Fall 1975, trapped,
total

114

157

271

42.1

57.9

1: 1. 38

November

1975 observed

71

104

175

40.6

59.4

1 :1.46

Combined

fall total

185

261

446

41.5

58.5

1:1.41

Late winter, 1976,
observed '1:.../

119

373

492

24.2

75.8

1:3.13

1975,

!/ Late winter

1975 observed

2:./ Late winter

1976 data from both within

and trapped ratios taken from Hoffman
and proximal

(1975).

to the study area.

Hoffman (1975) observed 1.28 hens per cock in the fall 1974 banded sample, in
contrast to 1.38 in fall 1975. His late winter trapping and sex ratio counts
placed the number of hens per cock in the heighborhood of 4.44 to 4.97 (Table
5). Compared to 1975 results he found slightly more roosters present in fall,
but a much greater deficiency of roosters in late winter.
Yet total hunting
mortality of roosters,including
crippling loss, in 1974 was projected at 20.4
percent, or slightly below the 24.7 percent level recorded a year later.
This
fall to winter differential loss of roosters approximated 70 percent in 1974-75
compared to a 55.9 percent loss over the same interval in 1975-76.

�-12-

Explanation of the reasons for the differential fall to winter loss of
roosters is beyond the scope of this study.
The accuracy of the samples,
upon which previous data are based, is not beyond question.
In addition,
losses of roosters may have resulted from fall movement of roosters out of
the study area without replacement, to increased mortality, or to a combination of movement and mortality factors.
As discussed in a following section
of this report, fall 1974, and fall 1975 movements of roosters averaged
several miles generally in a southeast direction.
Since hens were not
hunted, it is not known whether they also moved in what might be called a
fall shuffle.
Without this information we can only speculate.
The gaudy male pheasant may also be more vulnerable to predation, especially
that by avian predators than his female counterpart.
Numerous rough-legged
hawks, prairie falcons and some golden eagles migrate and reside in the study
region through the fall and winter.
Their harrassment and taking of pheasants
has been observed on occasion.
Pheasant
Fall 1975 Efforts

Trapping

and Banding

and Results

Pheasant trapping efforts commenced August 18 and terminated October 23,
1975. Results of the eight weeks of effort by two 2-man crews were poor,
with only 109 males and 150 females banded.
Four roosters, banded during the
preceding fall and winter, were retrapped, as were seven hens, bringing the
totals to 113 roosters and 157 hens handled.
Six birds were lost to injuries
inflicted during netting and holding operations.
Results of the fall trapping effort are summarized in Table 6. Since reward
bands from hens banded in 1974 were not obtained except when retrapped by
Division personnel, reward bands were not placed on hens in fall 1975.
Table
6 shows that 10 old roosters were reward banded of the 15 banded.
One-half
of the 94 young roosters were reward bmlded, yielding a 52.3 overall percentage with reward bands in the fall 1975 sample.
Roosters comprised 41.9
percent of the fall banded sample and hens totaled 58.1 percent.
Table 7 summarizes retrap information.
Six of the eleven retraps were initially banded as adults.
Only one banded juvenile male was recaptured.
Only
one of the eleven moved a considerable distance between retrap and recovery.
Since trapping was largely confined to the study area, those birds moving
considerable distances elsewhere were less vulnerable to retrap.
Trapping

Success per Unit Area

Trapping crews searched approximately 10,340 acres, including numerous second
searches of fields where higher numbers of birds were initially found.
Approximately 8,910 acres of wheat stubble were searched within the study area with
most efforts concentrated in the southeast township.
An additional 1,430 acres
outside the study area, primarily to the east, were searched.
The total acreage
was equivalent to about 123 stubble fields.
Based on 35 percent stubble per
section, search efforts covered the equivalent of 46 square miles with 2.46
roosters and 3.41 hens trapped per section.
Trapping success outside the study
areas was little better than that obtained in the area. An average of less
than one rooster was obtained per field searched.

�-13Table 6. Pheasants banded and retrapped in and adjacent
study area, August 18 to October 23, 1975. 1/

Total

Percent
Total
by Sex

Number
Reward
Banded

Percent
Reward
Banded

19

16.8

10

66.7

94

83.2

47

50.0

57

52.3

Age

Sex

Ntnnber
Banded

Adult

Male

15

Juvenile Male

94

Subtotal

109

4

113

100.0

7

63

40.1

94

59.9

157

100.0

Adult

Female

56

Juvenile

Female

94

Retrapped

4

Subtotal

150

7

Ratio:

Adult Males:Juvenile

Males

Adult Females:Juvenile
Adult Females:Total
Males: Females

to the mortality

Females

Young

Hens not reward banded

1.00:4.95
1.00 :1.49
1.00:2.98

1.00: 1.39

1/ Retraps

from both fall 1974 and winter 1975 as stnnmarized in Table 7.

Comparison

of Fall 1974 and Fall 1975 Age Data

Young to adult ratios obtained during production counts and fall trapping
show a higher young-per-adult
ratio in 1975 than in the preceding year (Table
8). The extremely poor production in 1974 was believed primarily a result of
severe drought conditions persisting through most of the spring and summer of
that year.
More young per adult were tallied during production counts than during subsequent trapping operations as shown in Table 8 for both years.
This was to
be expected, as broodless hens and moulting roosters were difficult to locate
and observe in comparison to young broods.
Recovery of Banding
The 1975 Hunting

Information

Season

Hunting Conditions--Efforts were made by the Division to divert hunting pressure
away from the storm decimated northeast corner of the state. Whereas, the
season opened at noon Saturday November 15 throughout the rest of the state, a
delayed opening to sunrise Wednesday November 19 was implemented in the study
region.
The season was terminated at the end of twelve days on November 30.
A bag limit of three roosters, with six in possession duplicated that of
previous years.

�-14Table 7. A sunnnary of Fall 1975 retrap information
Fall 1974 and Winter 1975.

Band
Number

Adult
Male

5003

x

Adult
Female

Banded As
Juvenile
Male

Juvenile
Female

from pheasants

Period Banded
Fall
Winter
1974
1975

banded

Distance
Moved
to Retrap

x

0

5051

x

x

0

5219

x

x

7+

5346

x

x

1

5347

x

x

1

x

1

5488

x

5543
5554

x
x

5555

x

5611

x

5650

x

Total

3

3

1

4

6

x

1

x

2

x

0

x

2

x

2

5

in

X = 1. 54 miles

A major snow storm hit the study region the night before the season was to
open and persisted throughout the opening day. Approximately ten inches of
snow were deposited, filling all wheat stubble fields and other cover, and
closing access to most of the area.
County roads were opened slowly and with
difficulty, so that no significant hunting occurred until the first Saturday,
November 22 (Table 9). By the first weekend most roads were open so that
considerable road hunting occurred.
Pushed out of their normal wheat stubble habitat by the deep snow, the pheasants
concentrated along roads, old building sites and other meager cover. As a
result, they were highly vulnerable to hunting, and when pushed out of available cover, most had no other place to go. One party of three hunters was
checked the second day of season with twelve birds.
All had been shot along
the highway south of Haxtun or adj acent access roads.
A majority of those hunters afield on Saturday the 22 had good to excellent
success.
Many simply drove the roads and removed exposed roosters as they
found them.
A few attempted walking, but the deep snow made hunting afoot
difficult.
It is estimated that at least 90 percent of the harvest was taken
adjacent to roadways.
Although hunting success was good among the hunters
present, the quality of the sport would have to be rated poor.

�-15Table 8. Comparisons of age ratios obtained during production
fall trapping in 1974 and 1975 on the mortality study area.

Year

Total Adults

Adult Hens

Production

Total Young

counts and

Ratio

Data

1974

84

241

1.00:2.87

1975

30

116

1.00 :3.87

1974

63

241

1.00 :3.83

1975

26

116

1.00:4.46

Banding

Data

1974

19l

343

1.00:1. 80

1975

82

188

1.00 :2.29

1974

143

343

1.00:2.40

1975

63

188

1. 00:2.98

Hunting pressure dropped off markedly by Sunday November 23, the 5th day of
season (Table 9). No significant hunting occurred thereafter.
Efforts to
contact hunters during the second weekend were stymied by cold winds and
light snow which once more closed many of the county roads.
Over half the
band return kills were taken on Saturday November 22, and only one banded
bird was reported taken after the first weekend of season.
Hoffman (1975) reported a maximum average of two hunters per square mile
during the opening Saturday of pheasant season in 1974 within the study area.
Information presented in Table 9 would indicate there were less than a tenth
as many hunters present during the first Saturday in 1975.
Road closures and
other impact of the 1975 snowstorm make it difficult to compare 1974 and 1975
hunting pressure data. In addition, access difficulties, low hunting pressure,
road hunting characteristics of the majority of the hunters, and their movements in and out of the study area made it unfeasible to attempt to measure
man-days of hunting within the study area.
Hunter Success--Field
contacted hunters, numbering 57 reported hunting roughly
201 hours and bagging 47 roosters in and near the study area. This approximated 0.82 pheasants per hunter.
Hunter questionnaire data showed that 34
hunters in and near the study area made 51 hunting trips totaling 65 days and
335 hours to obtain 85 roosters.
This equals 2.50 roosters per hunter for
the season, and approximately one rooster bagged per four hours of hunter
effort.
Seven roosters were reported crippled and not retrieved approximating
eight percent of the reported harvest.

�-16-

Table 9. Hunting pressure surveys during the 1975 pheasant season in the
mortality study area. !/
Date

Miles

Hours

Parties

Hunters

11-20

55

3

2

5

11-21

76

6.5

1

2

11-22

127

7.5

7

22

11-23

131

6

3

8

11-24

30

2

o

o

11-26

56

2

o

o

11-27

59

3

1

1

11-29

90

4

2

6

11-30

45

3

o

o

Total

669

37

16

44

Miles per Hunter

1st Saturday

5.8

1st Weekend

8.6

Remainder of
Season

29.4

15.2

1/ Most of the sample was obtained in the east and southeast parts of the
study area and in bordering regions to the east and south.
was lighter to the west and north.

Hunting pressure

Hunter success was better in 1975 than during the 1974 season based on
Hoffman1s questionnaire summary. He found that 52 hunters killed 87 pheasants,
or 1.67 roosters per hunter during the 1974 season. Approximately 4.68 hours
per hunter were expended per bird bagged in 1974.
Percent Banded in Harvest--Several factors make it impossible to obtain a
reliable banded-unbanded ratio from which to project fall population estimates.
These include: (1) high movement of birds between banding and hunting season;
(2) movements of hunters into and out of the study area while road hunting;
and (3) fall banding in locations outside the study area in 1975 in efforts
to increase sample size.
Hunter Band Return and Harvest Projection
Proportional Band Recovery--Contacts of hunters in the study area and in
proximal locations to the east and south were conducted during the 1975 season.
These contacts yielded four regular band and four reward band recoveries from

�-17fall 1975 banded roosters.
This ratio of reward to regular recoveries was
expected since 57 of 109 (52.3 percent) of the males contained reward bands
(Table 6). In addition, one reward band from fall 1974 was collected during
field contacts.
Among bands subsequently mailed in by hunters from the fall 1975 trapping
period, nine were reward bands and only one was a regular band.
The latter
was not received until several months after termination of the hunting season.
One regular band from fall 1974 was received by mail in addition to the above.
Twenty-eight roosters were banded in winter 1975, but no bands, reward or
regular, were recovered from hunting.
The above information clearly shows reporting rate bias between reward bands
and regular bands mailed in by hunters.
Originally, 57 reward bands were
placed on roosters in fall -1975 , of which 13 (22.8 percent) were recovered.
Fifty-two regular bands were placed on roosters in fall, of which only 5
(9.62 percent) were recovered.
By direct proportional probability we would
have expected to receive 12 regular bands rather than the 5 obtained.
This
represents a 41.67 percent reporting rate on regular bands.
Hoffman (1975)
showed a first year bias wherein only 66. 7 percent of the expected number of
regular bands were obtained from hunters.
The voluntary hunter reporting rate of regular bands was actually much lower
than the above figures indicate for both 1974 and 1975.
Many of the bands
were obtained during field contacts and in 1974 some were obtained at the
Fleming Check Station.
Looking only at mail in results for the 1975 fall
trapped birds, nine reward bands were recovered, compared to only one regular
band.
By direct proportional projection, eight regular bands should have
been received by mail, or the mail in reporting rate was about 12.5 percent
of the expected on regular bands.
Hoffman (1975) reported that 19 reward
bands were received by mail in 1974 in comparison to only 4 regular bands.
Direct proportional projection would place the expected mail in recovery at
16 bands.
In other words, the mail in recovery of regular bands in 1974 was
only 25 percent of the expected.
Sample sizes in both 1974 and 1975 were rather small, but they clearly point
out the failure of hunters to return bands to the Division by mail unless a
reward is offered.
There is no reliable way of knowing whether 100 percent
of the reward bands were returned, but the preceding and following projections assume that near 100 percent were recovered.
If reward bands had not
been placed on a portion of the roosters and field contacts had not been made,
then very little information would have been recovered and there would have
been no basis for projecting percent harvest or annual mortality rates.
It
seems certain that use of reward bands must be continued in subsequent years
of study.
Potentially, all roosters could receive a reward band, and if a
100 percent recovery rate is to be sustained over several years, then a
monetary value of five or ten dollars should be stamped on the reward band.
Continued issuance of the present reward, a wild game cookbook, or similar
reward over a period of years potentially would reduce hunter incentive to
submit reward bands.

�-18Differential Harvest by Age--Hoffman (1975) observed a difference in harvest
mortality in 1974 between adult and juvenile roosters.
Juvenile males sustained a 19 percent harvest mortality, whereas adults sustained about 15
percent harvest.
In 1975, harvest mortality of adults was 20.0 percent, and
that of juvenile males was 23.4 percent.
These 1975 data are based on
reward band return sample sizes of only 2 for adults and 11 for juveniles.
Inadequate sample size prevents drawing the definite conclusion that juvenile males were more vulnerable to hunting than adults, but similar trend
in recovery rates by age suggest this is the case.
1975 Harvest Estimate--Thirteen
reward bands were returned from the
original 57 placed on roosters during the fall of 1975. Assuming all recovered reward bands were reported, this represents an estimate of 22.8 percent
harvest of the original fall population.
Hunters reported an 8.2 percent
crippling loss in addition to the retrieved harvest.
Combined crippling loss
and retrieved harvest would bring the total hunting season mortality to 24.7
percent among roosters.
Hoffman (1975) placed the 1974 rooster harvest at 18.5 percent of the original
banded popUlation.
He added a crippling loss equal to 10.3 percent of the
retrieved bag, bringing the total estimate of mortality due to hunting to
20.4 percent.
The 1974 season was longer than in 1975 (29.5 days versus 12 days), pheasant
populations were much greater in 1974 and many more hunters were afield.
The
slightly higher percentage kill in 1975, if accurate, was believed due to snow
stress conditions which exposed and concentrated the pheasants where they were
accessible to hunters early in the season.
It is doubtful that half the
projected number of roosters would have been taken in 1975 if the snow condition had not accompanied the season.
The higher season bag per hunter in
1975 than in 1974 (2.50 compared to 1.67) reflects the greater hunting success
in 1975, even though pheasant popUlations were much lower.
Natural

Mortalities

The storm, which deposited about 10 inches of snow on the area at the start
of the 1975 pheasant season, also caused undetermined mortality.
One party
of hunters reported finding several dead pheasants a few miles south of the
study area. Two farmers reported observing several dead pheasants while
checking cattle in corn stubble fields.
At another location, a farm dog
reportedly brought in a couple pheasants killed by the storm.
Several pheasants were reported to have been buried in a shelterbelt near Holyoke and were
discovered during late winter snow melt.
No bands were recovered or reported
in these instances.
A lone bobwhite quail was observed in late February
within a study area tree planting site where November drifted snow had entombed
it.

Efforts to assess the impact of storm caused mortality were hampered by
reoccurring light snows which drifted over otherwise exposed birds.
Prior to
the March blizzard, roosters had dispersed to their territories along roadways where they were readily observed as mortalities after the storm.
In fall,
however, they were not dispersed in this manner, so mortalities if present,

�-19were more difficult to find. Birds that survived the fall snow did concentrate in weedy cover, old building sites, and other locations where they
were highly visible.
Searches on foot in late November and early December
were not effective.
Therefore, an accurate assessment of mortality could
not be made, but overall mortality was not considered to exceed twenty-five
percent of the existing fall population.
Annual Mortality

Rate

Funk (1966) illustrated a method for calculating annual mortality based on
only two years data. This procedure is illustrated in Table 10 and shows a
94.8 percent overall mortality rate for the 1974 fall banded rooster cohort.
Both reward and regular band recoveries were used in Table 10 to increase
sample size.
If only reward band data were used, as in computation of percent harvest, then the annual mortality rate in Table 10 would equal 96.5
percent.
However, this projection would be based on only one second year
recovery.
Obviously, band recovery sample sizes were too small to permit
accurate prediction of annual mortality.
Winter 1974-75 sex ratio counts,
March blizzard mortality evaluation, crowing counts, and triangulation
results all indicate that high mortality of the pheasant population did occur
within the projection interval.
Movement
A high degree of movement was observed with average movement approximating
4.6 miles from fall trapping to harvest.
The range in movement was from zero
to about 19.5 miles with only a few roosters moving great distances and the
majority moving less than three miles.
Hoffman (1975) reported an average movement during the fall of 1974 of 4.7
miles, with a range from less than one up to twelve miles.
Such movements
make it impossible to project fall population levels based on banded-unbanded
ratios within a four township study area.

Table 10. Relative recovery method of estimating mortality
roosters from the fall of 1974 to the fall of 1975.

Fall
Banded

Number
Marked

1974

233

36

1975

109

18

Harvest Recoveries
First Year
Second Year

Annual Survival

Rate

Annual Mortality

Rate

0.0086
0.1651

0.0086
0.1651

94. 79 percent

of

Recovery Rate
First Year
Second Year

2

5.21 percent

and survival

�-20LITERATURE

CITED

Funk, H. D. 1966. Review of duck literature relating to population dynamics,
and banding analysis techniques and findings. Colo. Dept. of Game,
Fish and Parks. Game Res. Rept. Oct. p. 77-108.
Hoffman, D. M. 1975. Pheasant mortality investigation.
Wildl. Game Research Rept. April. p. 5-35.

Prepared by
Warren D. Snyder
Wildlife Researcher

J

Colo. Div. of

�April 1976
-21JOB FINAL REPORT

State of

Colorado
~----~~~~~-----------

Project No.__~W~-~3~7_-~R~-~2~9
Work Plan No.

_

Game Bird Survey

3
Job No.
Sa
--------~~~~----~--------==----------------------Effects of Sagebrush Control

Job Title

on Distribution

and Abundance

of Sage Grouse

Period Covered:

April 1, 1963 to March 31, 1975

Personnel:
Thomas Beck, Don Benson, George Bock, Clait Braun, Gary Brown,
Lonnie Brown, Thomas Campbell III, Harold Carr, Don Clamp, Don Clamp, Jr.,
Steve Cook, Jack Corey, Courtney Crawford, David Croonquist, Mike Dorrance,
Earl Downer, Marcus Elkins, John Ellenberger, Dwayne Finch, Heather Flanagan,
Howard Funk, Fred Giese, Bruce Gill, Fred Glover, Don Gore, Jack Gustafson, John
Hobbs, Donald Hoffman, Richard Hoffman, Norm Hughes, Richard Imler, Jim Jackson,
Scott King, Edward Kochman, Russell Kozacek, Russell Mason, Terry May, Bruce
McCloskey, Mike McLain, Chet McCord, Ken Miller, John Monarch, Ken Morrison,
Thomas Morrow, Ron Oakleaf, Perry Olson, Sig Palm, Bruce Poley, Steve Porter,
Dale Reed, Mike Robertson, Glenn Rogers, Larry Roper, Kenneth Russell, Wayne
Russell, Ronald Ryder, Wayne Sandfort, Ray Schmidt, Jr., Bruce Sigler, Warren
Snyder, Howard Spear, Robert Streeter, Harold Swope, Michael Szymczak, Melvin
Taylor, John Wagner, Francis Waugh, Clayton Wetherill, Richard Wenger, Jerry
Whitaker, and Lee Yeager.
ABSTRACT
Distribution and abundance of sage grouse (Centrocercus urophasianus) in North
Park, Colorado were studied prior to and following experimental spraying of
4,000 acres of sagebrush (Artemisia spp ,) with 2,4-D.
Spraying resulted in
decreases in cover and relative frequencies of sagebrush and forbs.
Active
strutting grounds within the original study area declined from seven to six
from 1963 to 1974. Three strutting grounds were abandoned in the study
period while two apparently new grounds were located.
Maximum numbers of
male sage grouse counted on strutting grounds in the original study area
decreased significantly (P &lt; 0.05) from when counts were initiated (1959) to
1974. Apparent value of strutting ground counts of males in unknown due to
seasonal variations from 24 to 60 percent in maximum daily counts.
Adult
male sage grouse exhibited great fidelity to grounds where originally banded,
while those banded as subadults frequently were observed in years following
banding on grounds other than where initially marked.
Use of long handled
nets with a backpack power source was the most successful method of trapping
male sage grouse.
Weights of male sage grouse increased prior to the
strutting period and then decreased until late May.
Females gained weight
until time of nes t.Lng, Block spraying of sagebrush effectively eliminated
nesting by sage grouse and greatly reduced sage grouse use of treated areas.
While centers of sprayed strips were avoided by sage grouse, nesting and
other use of edges of sprayed strips did occur, apparently the result of low
percent sagebrush kill. Roadside censuses of broods were not effective at
assessing production or distribution probably because of the abundance of
brood habitat in the park. Harvest was variable and was not correlated with

�=22ABSTRACT

(Continued)

strutting ground counts or production estimates.
Observed sex ratios in
the harvest favored females in the older age classes and was not the result
of hunter selection.
Instead, sex ratios in the harvest approximated those
obtained during counts conducted in winter.
Harvest statistics were variable
and hunter success apparently decreased over the study period.
Hunter harvests
of grouse had little demonstrable effect on population levels as recoveries
of banded birds approximated 10 percent of those available.
Distribution of
grouse in winter was restricted due to snow depth, aspect, and slope.
Seven
important winter use areas were identified comprising less than 7 percent of
the sagebrush lands in North Park.
Sage grouse flocks were frequently
segregated by sex with females forming larger flocks which used denser stands
of sagebrush than did males.
Movements of sage grouse were more extensive
than expected.
Consequently, sage grouse in North Park should be considered
as one population.
Effects of sagebrush treatment on sage grouse in the
original study area and in North Park were difficult to assess as one third
of the area was in hay meadows and half of the sagebrush in the remaining
area had been treated.
Block treatment of sagebrush was especially detrimental to sage grouse use while effects of treatment in narrow strips could
not be ascertained.
Treatment of sagebrush in North Park has resulted in
shifts in sage grouse distribution and in apparent decreases in sage grouse
numbers.
If sage grouse popUlations are to stabilize in North Park no
additional treatment of sagebrush must occur.

"

�-23RECOMMENDATIONS

1.

Systematic searches for new or relocated strutting grounds should be
made annually throughout North Park.
Searches should be conducted from
the ground in April and May.

2.

All strutting grounds should be counted one-half hour before sunrise four
times each spring.
Two counts should be made between April 10 and 20,
at least 3 days apart, with the other two counts being made between
April 20 and 30 (one), and May 1 and 10 (one). Birds counted should be
classified as male and female.

3.

Seasonal patterns and lengths of male and female attendance on grounds
should be investigated in order to develop
a reliable index to spring
population levels.

4.

Two hundred individuals of each sex should be banded annually throughout
North Park through 1977 during the March through May period to assess
survival and turnover rates and population size.

5.

Techniques for capturing hens and chicks need to be investigated and
developed to further assess survival and turnover rates and population
size.

6.

Current brood routes should be discontinued.
Methods should be investigated and developed for obtaining 50 or more brood sightings by August
10. Emphasis on counting broods should be placed from 11 to 13 weeks
after peak hen attendance on strutting grounds.
All suitable areas in
North Park, especially native hay meadows, should be sampled for brood
utilization.

7.

Check stations should be operated annually both days of the opening
weekend of the hunting season at Gould, State Line, and Willow Creek
Pass to obtain estimates of juvenile, yearling and adult" age classes,
sex structure of the kill and distribution of the harvest.
A minimum
of 500 birds should be checked annually.

8.

Manipulation of season length and bag
effect, if any, of changes in harvest
levels.
Monitoring of harvest levels
permits and 100 percent questionnaire
a maximum of 2,000.

9.

Land management agencies should be provided with maps of sage grouse
winter use areas, strutting ground locations and important brood use
areas once the latter is available.
The Division of Wildlife should
vigorously oppose disturbance of the seven major winter use areas and
all strutting grounds through habitat alteration.

10.

If sagebrush is to be sprayed, only those areas where sagebrush is
completely snow covered in winter should be sprayed, and then only in
strips following these specifications:
(1) sprayed strips should not
exceed 50 yards in width or longer than two miles in length, (2)
unsprayed strips should be at least 150 yards wide, (3) sprayed strips
should be on slopes greater than 15 percent, and (4) percent kill of
sagebrush should not exceed 60 percent.

limit is desirable to learn the
regulations on spring population
should be through use of free
sampling of all permittees up to

�-24RECOMMENDATIONS

(Cont.)

11.

A management plan for sage grouse in North Park must consider
Park as one unit.

the entire

12.

Emphasis should be placed on investigating the biology and behavior of
female sage grouse.
Since wings are easily collected at check stations,
the relationship of primary feather molt and nesting success should
have highest priority.

13.

Investigations should be initiated into habitat improvement for sage
grouse with fertilization of sagebrush communities having highest
priori ty.

�-25-

EFFECTS
ON DISTRIBUTION

OF SAGEBRUSH CONTROL
AND ABUNDANCE OF SAGE GROUSE

Clait E. Braun and Thomas

D. I. Beck

Sage grouse inhabit areas dominated by sagebrush (Artemisia spp.) and
subdominant grass types with interspersed native and cultivated hay meadows.
Dependence on sagebrush for food and cover has been well documented in all
seasons (Griner 1939, Patterson 1952, Dalke et ale 1963, Autenrieth 1969,
Klebenow 1969, Eng and Schladweiler 1972).
As ;-result of agricultural
demands, livestock grazing, and encroachment of man-related developments,
there has been continual alteration of sage grouse habitat.
In the face of
changing land use patterns, sage grouse have continued to be highly
specialized in their habitat requirements and have not adapted to utilize
land dominated by plants other than sagebrush.
Plant species characteristics
of climax communities recover slowly after natural or man-caused
catastrophies (Daubenmire 1970).
Therefore, these plant species are
unreliable food resources where communities are frequently or massively
disturbed.
Intensive studies of sage grouse throughout the west have provided extensive
data on many phases of their life history.
Much research has been devoted to
studying the requirements for successful population maintenance or to their
behavior.
In the early 1950's when spraying of sagebrush with 2,4-D
(2,4-dichlorophenoxyacetic
acid) to kill the shrub overstory became an
integral part of western range management, there was little documentation
of
the effects of shrub reduction on wildlife.
In 1963 the Colorado Game and Fish Department (now Colorado Division of
Wildlife), in cooperation with the Bureau of Land Management, initiated a
study in North Park, Colorado to evaluate the effects of 2,4-D spraying of
sagebrush on abundance and distribution of sage grouse.
North Park (Jackson County) was chosen for study following a proposal in
1962 by the Bureau of Land Management to spray all BLM controlled sagebrush
lands in the park. This area included one of the larger strutting ground
complexes in Colorado (Rogers 1964).
The study was initially confined to
the northwest quarter (Lake John area) of North Park as it was believed this
area contained a large number of grouse throughout the year.
The area
initially studied (Fig. 1) was bordered on the north by Independence Mountain,
on the east by the North Platte River, on the south by the North Fork of the
North Platte, and on the west by Sheep Mountain and Boettcher Ridge.
This
encompasses approximately 50 mi2 of sagebrush lands and 21 mi2 of
associated meadows.
After review of results through 1973 the decision was
made in January 1974 to expand the area studied to all of North Park.
P. S. OBJECTIVES
To determine the effects of 2,4-D spraying of sagebrush on (1) sage grouse
abundance, (2) sage grouse distribution, and (3) vegetative composition and
density.

�-26-

N

i
Fig. 1. Original Lake John study area, North Park. Colorado. Areas
Geli~eated by heavy black lines are those experimentally treated by
spraying.

�-27DESCRIPTION

OF AREA

North Park is the northernmost of Colorado's four large montane parks and
is located immediately west of the Front Range of the Rocky Mountains.
It
lies within townships 5 to 11 north and ranges 77 to 82 west of the sixth
principal meridian.
North Park is completely enclosed by mountains, on
the west by the Park Range, on the east by the Medicine Bow Range, on the
south by the Rabbit Ears Range and on the north by Independence Mountain.
The area is approximately 50 miles wide and 56 miles long (Fig. 2).
North Park is relatively flat and is characterized by low undulating benches
and ridges separated by the main drainages.
The main streams form a
meandering pattern on low alluvial flood plains up to one-half mile wide.
Elevations of most sagebrush lands are between 7,870 and 8,500 ft above sea
level.
The principal exception is Owl Ridge-Peterson Ridge, two end-to-end
ridges running northwest from Owl Mountain on the east to Delaney Butte on
the west and rising 200-500 ft above the surrounding terrain.
Mountains
encircling the park rise steeply to elevations from 10,100 to 12,600 ft above
sea level.
Geology of the area has been described in numerous publications
(Beekly 1915, Finch 1957, Hail 1965).
Major drainage is to the north via
the North Platte River. Main tributaries to the North Platte include the
Roaring Fork, North Fork, Canadian, Michigan, and Illinois Rivers and Grizzly
Creek.
Over 721 mi2 of sage grouse habitat occur in this area as all sites not
timbered are considered potentially usable by sage grouse.
Of this, about
483 mi2 are dominated by sagebrush, while about 239 mi2 are irrigated
meadows of native sedges and grasses.
With the exception of hay meadows along watercourses,
the vegetation is
composed mainly of the shurb-bunchgrass
type dominated by five species of
sagebrush.
Artemisia tridentata vaseyana occupies approximately 90 percent
of the sagebrush type. Artemisia longiloba, !. cana viscidula, !. argilosa,
and A. -nova are dominant on limited areas where soil conditions are favorable
to these species (Beetle 1960, Smith 1966).
Other important shrubs in North
Park are greasewood (Sarcobatus vermiculatis),
rabbitbrush (Chrysothamnus
spp.), snakeweed (Gutierrezia spp.), willows (Salix spp.), and bitterbrush
(Purshia tridentata).
Herbaceous vegetation consists primarily of low
growing perennial forbs and perennial bunch grasses with few annual forbs
(Appendix A).
The climate of North Park is best described as cold and dry. U. S. Weather
Bureau precipitation and temperature data are available from Walden
(elevation 8,099 ft) in the center of the park and near Spicer (elevation
8,379 ft) at the southwest corner, where sagebrush merges with aspen
(Populus tremuloides).
The 30-year mean precipitation and temperature by
month from 1941-1970 are shown in Table 1. No wind measurements are available
but prevailing winds are from the southwest with frequent high velocities,
particularly in winter and spring.
Amount and distribution of precipitation
are variable from year to year. Much of the snow sublimates, thus providing
little moisture to the soil. Drifting is common throughout the park because
of constant winds and rolling terrain.
The average frost-free season in
Walden is 46 days, from mid-June to early August.
Growth patterns of native
vegetation are highly variable due to uncertain and often extreme environmental
conditions.

�-28-

.- 'c
.~

Fig. 2. North Park, Colorado with original Lake John study area delineated
by h~avy black line.

�-29-

Table 1.

Temperature and precipitation, North Park, 1941-1970.!I
Walden
Mean Monthly
Precipitation
Temperature
(In)
(oF)

Spicer
Mean Monthly
Precipitation
Temperature
(In)

(OF)

January

0.51

15.4

1.06

16.6

February

0.42

lS.l

0.94

20.2

March

0.50

23.7

1.10

25.2

April

0.72

35.3

1.25

35.3

May

1.02

44.S

1. 36

43.9

June

1.11

52.9

1.37

52.6

July

1.24

5S.7

1.22

59.0

August

1.29

56.3

1.69

57.0

September

1.01

4S.7

1.17

4S.3

October

0.71

39.1

0.91

39.4

November

0.54

26.7

1.01

27.3

December

0.54

lS.6

1.26

lS.7

Mean Annual
Precipitation

9.61

Temperature

Y

14.34
36.5

37.0

1/
- Data from U. S. Department of Commerce (1973). 30-year weather summary
for Colorado.
2/
- Temperature data not available from Spicer for 1941-1970, data presented
cover 1930-1965.

�-30-

METHODS

AND MATERIALS

Design of Sagebrush

Treatment

Following preliminary studies Gill (1965) recommended two patterns of
sagebrush treatment, strip and block sprayin~ with treatment to be conducted
in four areas.
Strips to be sprayed were to be from 50 to 250 yds wid~,
with 200 yd wide untreated strips between those treated.
Total acreage
to be sprayed in strips was 484 acres.
Two areas were recommended for block
spraying totaling 2,316 acres.
These areas were 515 and 1,801 acres in size
and were to be separated by 1,600 acres of un treated sagebrush.
The
remainder of the original study area was to serv~ as a control with no
sagebrush treatement.

Vegetation

Analysis

Twenty-one permanent macroplots
(each 100 ft2) were established in the Lake
John intensive study area with seven in each of the three height classes of
sagebrush.
Height classes were: less than 5 in, 5-10 in, and greater than
10 in. In 1964 all plots were in areas of no noticeable disturbance to the
sagebrush.
Plots were located so that four of the seven plots within each
height class would be in a sprayed area following spraying in June 1965.
Post-spray sampling was conducted in 1966 and 1973. Sampling techniques
were patterned after Hyder et al. (1963) and a detailed description is
presented by Gill (1965).
Each macroplot consisted of a 100 ft baseline at
right angles to the contour of the site. Four transect lines, one randomly
placed to the contour of the site. Four transect lines, one randomly placed
in each 25 ft segment of the baseline, were placed at right angles to the
baseline.
Beginning at the 2 ft marker on each transect line, sample points
were located at 4 ft intervals so that there were 25 sample points per
transect.
A one-foot square quadrat was placed at each sample point and
each plant species within the quadrat was recorded.
In addition to frequency
data, measurements of sagebrush crown intercept along the transect line were
recorded.
Density of sagebrush was determined by counting all sagebrush
plants within 6 inches of the transect line, thus obtaining plants per
100 ft2.
Location

of Strutting

Grounds

In April and May, 1963, strutting grounds were located by driving roads
bisecting the study area and stopping every mile to listen and look for
displaying males during the period one hour before and one hour after sunrise.
Surrounding
terrain was scanned with 7 X 35 and 7 X 50 binoculars.
In 1964
searches for strutting grounds were conducted from an airplane similar to
that described by Eng (1954). Aerial transects were flown along north-south
and east-west lines one-half mile apart and 100-200 it above the ground during
the two hours following sunrise.
Ground searches were not conducted in 1964.

�-31-

Searches for new or relocated grounds were not conducted in the 1965-1972
period.
Aerial and ground searches were utilized again in 1973, while
extensive ground searches were conducted in 1974.
Searches in 1973 and
1974 were conducted throughout North Park in contrast to the 1960's when
searching was limited to the Lake John area. Strutting ground locations
were plotted on U. S. Geological Survey topographic maps (7.5 minute series).
Strutting

Ground

Counts

Observations of sage grouse on strutting grounds were conducted from vehicles
as described by Patterson (1952) for use as an index to the number of breeding
birds.
Observations were made during the period 1/2 hour before sunrise to
1/2 hour after sunrise.
Each strutting ground was counted at least three
times during the strutting season, which usually lasted from late March to
late May.
Data collected were: (1) maximum number of males and females,
(2) age ratio of males, and (3) observation of marked birds.
The highest
of the three counts was considered to be the maximum number of males
attending a strutting ground.
The method is based on the assumption that all
males in the population regularly attend a strutting ground (Patterson 1952).
In 1974 counts were made with greater frequency on the larger grounds with
at least four counts conducted during the peak period of female attendance.
At least eight counts were made of each selected ground during the strutting
season.
Counts were normally made from distances of 10-100 yds, depending
on terrain, height of sagebrush, and observer preference.
Occasionally,
due
to weather conditions (deep snow), counts were made from distances up to 1 1/2
miles (B. Poley, personal communication).
This may have influenced counts
on some grounds in every year and particularly at Alkali Lake in 1968.

Trapping

and Banding

Five different trapping techniques were used during the study: (1) spotlight
trapping from a vehicle, (2) spotlight trapping with a backpack power source,
(3) ground-mounted
cannon nets, (4) bumper-mounted
cannon nets, and (5)
dri ve traps.
Spotlight trapping involved locating areas where sage grouse roosted at
night, primarily on strutting grounds in spring.
Grouse were located by
scanning possible roosting areas with a hand-held spotlight.
Individual sage
grouse selected for capture were blinded by holding the light beam on the
bird's head while a second person walked along the beam until close enough
to net the bird.
Initially, nets used consisted of a 39 in. diameter hoop
covered with 1.5 in. mesh cotton netting on a 10 ft metal pipe.
In 1974, 0.5
in. mesh nylon netting was used in an effort to reduce damage to primary
feathers.
In addition. hoop diameter was reduced to 28 in., with aluminum
tubing being used in net construction to reduce weight, thus providing
greater maneuverability.
Sealed beam spotlights wired to a vehicle's 12
volt electrical system were initially used for the light source.
However,
best results were obtained when using an aircraft landing light (250 watt
bulb) which was much brighter than sealed beam lights.
Captured birds were

�-32-

placed in burlap bags for holding until they could be banded.
A modification
of this procedure was developed during the spring of 1973. Instead of using
a motor vehicle electrical system for the power source, an automobile
battery (12 v) fastened to a pack frame was used.
This allowed greater
mobility and enabled workers to search areas inaccessible to motor vehicles.
When using the battery pack, tape recorded snowmobile noise was played to
cover the sounds of approach.
Cannon net trapping was first attempted in 1963 and subsequently used in
Nylon nets of 30 x 60 ft and 35 x 75 ft were used on strutting
grounds.
Nets were placed near primary mating centers set to fire with the
prevailing wind.
Nets were projected by three cannons and detonated
electrically.
Use of ground-mounted
cannon nets was limited to the period
of peak female attendance when bird concentrations were greatest.
A bumper
mounted cannon net (30 x 50 ft nylon net projected by two cannons) similar
to that described by Lacher and Lacher (1964) was used in 1965, 1968, and
1969. This method was used in spring on strutting grounds and in summer on
brood areas.

1965 and 1974.

Drive traps were used during the summers of 1963 and 1965 in attempts to
capture sage grouse chicks.
Traps consisted of a 3.5 x 8 ft catch pen of
variable depth covered with 0.25 in. nylon mesh and two 150 ft wings of 3 ft
tall wire deployed from the catch pen in a wide V. Drive traps were placed
to intersect travel routes to and from meadows.
All birds captured were banded with aluminum serially numbered size 16 (males)
and 14 (females) leg bands.
In addition, unnumbered colored aluminum leg
bands coded to capture location were placed on birds in 1963 and 1964.
Colored, numbered plastic bandettes were placed on all birds captured in
1965-1969 while colored~ unnumbered plastic bandettes were used in 1973 and

1974.
The following marking methods were also used in some years: back-tag harness
(Labinski and Mann 1962) 1963; jesse-knot neck collar (Craighead and Stocks tad
1956), 1963; poncho tags (Pyrah 1970), 1964-66 and 1974; dyeing with picric
acid solution, 1963; plastic patagial streamers, 1966-67; and metal patagium
tags for chicks, 1968-69. Very high frequency (mIF) radio transmitters
(Markus en brand) were placed on three hens and one cock in 1965, four cocks
and one hen in 1966. 10 hens in 1968, and 24 hens in 1969. The transmitter
harness was patterned after that described by Marshall (1963) in 1965 and
1966 and after Brander (1968) in 1968 and 1969.
Age and Sex Determination
Ages of males observed on strutting grounds were estimated using two methods.
Patterson's (1952) method, based on the width of the white band on the breast,
was utilized in 1964. 1965 and 1966. The accuracy of this method was
questioned by Dalke et ala (1963) on the basis that plumage differences
between adults and yearlings were not significant in years when the preceding
nesting season was relatively early.
Use of this method was discontinued
after 1966. In 1974 aging of males was based on the shape and length of

�-33-

rectrices as adults characteristically
have longer and more pointed rectrices
than yearlings (Lumsden 1968, Wiley 1973).
Since Dalke et ale (1963) also
questioned the accuracy of this technique, all males captured in 1974 were
examined to test if aging by rectrices
corresponded to the accepted technique
based on appearance of the outer primaries (Eng 1955).
All sage grouse captured in spring were classed as adults or yearlings based
on primary wear (Eng 1955).
This method is based on the retention of the
outer two primaries by juvenile birds until 15-16 months of age. Those
juvenile feathers are more pointed and frequently more frayed than adult
feathers.
Sage grouse wings collected during fall hunting seasons of 1963-73 were used
to estimate age and sex of harvested birds using the key developed by Crunden
(1963). This method differentiates
two age classes, juvenile and adult.
In 1974 Eng's (1955) method was utilized allowing differentiation
of three
age classes; juvenile, yearling, and adult.
Sex classification was based on
primary lengths.
Age (in weeks) and sex of juvenile birds were determined
by primary length (Pyrah 1963).
Suitable methodology for age and sex
classification of sage grouse wings to obtain maximum information have been
summarized by Beck et ala (1975).
Nest Searches
Forty-one la-acre nest search plots were established in 1964 (Gill 1965).
These plots were located at section and quarter-section corners, with one
plot established in each section of brushland in the Lake John intensive study
area. Two persons systematically searched each plot in 1964 in a grid fashion,
maintaining a l4-ft distance between themselves.
In 1965, 60, 5-acre plots
were established; 27 in sprayed areas and 33 in unsprayed areas (Carr 1967).
All plots selected in 1965 were within 1.25 mi of a strutting ground, with
12 within 0.25 mi. Systematic searches of these plots were conducted in 1965,
1966, and 1973. In 1968 and 1969 some hens fitted with VHF radio transmitters
were followed to their nests.
A few nests were found incidental to other
activities.
Upon locating nests, the following data were recorded: (I} age
of nest (nest of year or previous years), (2) clutch size, (3) fate of eggs,
and (4) vegetative cover in nest area.
Brood Census
Post-breeding season distribution and production estimates were investigated
in 1963 by classifying all birds by sex and age that were observed while
systematically traversing all major roads in the Lake John area. Upon sighting
a hen with chicks~ observers attempted to flush the brood to obtain complete
counts.
Based on the 1963 sampling, five trend routes (21.3, 22.8, l5~7, 26.6,
and 19.6 mi long) were established to provide a systematic method for deriving
production estimates.
Four routes sampled native hay meadows and adjoining
brushland, while the fifth sampled upland sagebrush areas.
Each route was
sampled once each week during July and August of 1964, 1965, and 1966.
Data

�-34obtained were: broods per mile, mean brood size, and percent of hens with
broods. All sage grouse observed were classified to age and sex when
possible.
In 1968 and 1969 broods observed while following radio equipped hens were
recorded but established transects were not counted. In 1973 and 1974
observers searched widely scattered meadows throughout North Park in an
attempt to observe greater numbers of broods. Three of the five established
routes were inaccessible in 1973 because of problems with landowners. Summer
distribution was inferred from sightings of sage grouse along established
routes as well as all sightings made during other field work.
Hunter Check Stations
During the period studied, the sage grouse hunting season opened on the second
Saturday of September, season length varied from one to three days, with
daily bag and possession limits of two and four. From 1963 through 1973
check stations designed to sample hunters from the Lake John area were
operated on the first two days of the season at Cowdrey and Walden. In 1974,
four stations were operated on the opening weekend to sample hunters from
throughout North Park. Stations were located at the state line on Colorado
125, Gould, Muddy Pass, and Willow Creek Pass. Data obtained each year were:
county of hunter origin, number of hunters, hours hunted, birds bagged, number
of banded birds and location of where each was obtained. One wing was obtained
from each bird that was checked except from birds which were completely dressed.
Hunter Questionnaire Survey
All North Park sage grouse hunters in 1974 were required to obtain and have
in their possession while hunting a free special permit. Purpose of the
permit was to provide a method for sampling hunters through questionnaires
to obtain accurate estimates of hunter numbers and total harvest. Permits
unlimited in number were available at all license agents in North Park, the
Fort Collins and Denver offices of the Division of Wildlife, and all Wildlife
Conservation Officers and research personnel working in North Park. All
permit holders were mailed a questionnaire within five days after the season
closed. Follow-up questionnaires were sent to non-respondents four weeks
after the initial mailing.
Winter Distribution
Aerial flights of the Lake John study area were conducted in February and
March, 1965, and January, February, and March, 1966; with one flight each
month (Carr 1967). Reports from Wildlife Conservation Officers were used to
summarize the distribution of sage grouse in the remainder of North Park
during the 1963-66 period. Extensive studies of sage grouse winter ecology
were conducted from 8 January to 8 March 1974 and 24 December 1974 to 1 March
1975 (Beck 1975). Distribution of wintering sage grouse in North Park was

�-35investigated by searching all sagebrush lands with all probable use areas
searched equally.
An area was considered potentially usable if any sagebrush
was visible above the snow.
Limited searches were also made in areas not
considered usable.
Areas inaccessible by four-wheel drive vehicle were
searched from snowmobile or snowshoes.
Aerial search (with two observers)
along random transects (covering 10 percent of the total area in the park)
were conducted on 21-22 February and 18 March 1974.
Data on grouse flocks
observed from the air were used in mapping distribution but were disregarded
when analyzing flock sizes because of inaccuracies involved in counting
flying grouse.
Once grouse were located from the ground they were first
observed through 7 X 50 binoculars to obtain a total count and to determine
sex composition.
Attempts to observe colored leg bands were made through
15-60 variable power scopes.
Written records were kept on both grouse
observations and grouse sign (tracks, droppings, roosts).
Observations
reported by reliable observers were followed by intensive searches of reported
locations within one day of the initial sighting.
All locations were plotted
on U. S. Geological Survey topographic maps (7.5 minute series).
A flock was considered to be any group (two or more) of birds
proximity to each other.
Sage grouse are sexually dimorphic,
sex of birds to be visually determined up to 200 yds. Flocks
greater than 50 percent of one sex were referred to as female
depending upon which sex predominated.

in close
thus allowing
comprised of
or male flocks,

The following physical and vegetal characteristics of each site where birds
were observed were recorded: aspect, slope, snow depth and condition, sagebrush
density, crown breadth, and height of sagebrush above snow.
Slope and aspect
were measured with an Abney level and compass, respectively.
Snow condition
was rated as crusted, powder, or melting.
Sagebrush density was measured
as number of plants per circular 0.01 ac. plot.
Crown breadth was calculated
as the mean of 10 representative shrubs arbitrarily chosen by the observer
from within the plot •. Height of sagebrush above snow was calculated as the
mean of three shrubs within the plot which the observer chose as being
representative of shrub height within the utilized area. The following weather
conditions were recorded at each site: wind direction, temperature, cloud
cover, and precipitation.
Movements
Data on movements of sage grouse from banding sites, primarily strutting
grounds, were available from observations of color marked birds, band
recoveries, recaptures, and radio tracking.
Movements of radio equipped birds
were monitored using a Markusen l2-channel VHF receiver with hand-held and
truck-mounted antennas.
Recoveries and reobservations of banded birds were
plotted on U. S. Geological Survey topographic maps (7.5 minute series).
Airline distances were then calculated for all recoveries and reobservations,
thus all movements represent minimum distances travelled.
RESULTS AND DISCUSSION
Sagebrush

Treatment

The ELM contracted to have 4,000 acres of sagebrush lands in the Lake John
area aerially sprayed with 2,4-D.
Rate of application was two pounds of

�-36iso-octyl ester of 2,4-D in 4.67 gallons of water per acre. Spraying was
accomplished during 2-5 June 1965 and followed the recommendations of Gill
(1965). Two basic patterns of spraying were conducted, block and strip,
with two modifications of each. The two block spray areas were about 515
.and 1,801 acres in size, with an intervening block of approximately 1,600
acres of undisturbed sagebrush. The largest strip spray area bordered the
block sprays on the west and encompassed about 1,180 acres. The sprayed
strips were 50 yds wide and aligned in an east-west direction with
intervening 200 yd wide strips of undisturbed sagebrush. The smaller strip
sprayed area, 484 acres, was sprayed so that strips of varying width with
intervening 200 yd wide control strips occurred. Seventeen strips were
sprayed: four 50 yards wide, four 100 yards wide, four 150 yards wide, three
200 yards wide, and two 250 yards wide.
Vegetation Analysis
Vegetation of the 21 permanent macroplots established in 1964 was described
in 1965, 1966 and 1973 (Tables 2 and 3). During pre-spray plot selection
and description of vegetation, none of the areas sampled appeared to be
altered. It is apparent from Table 2 that sagebrush and forb frequencies
were reduced following spraying with reductions continuing to 1973. As
expected, frequencies of grasses increased in all three sagebrush types
treated. Sagebrush crown intercept (Table 3) decreased in all height classes
following spraying and was still low in 1973. However, there was a marked
increase in crown intercept of seedling and young sagebrush in 1973 over 1965
and 1966. This indicates that the sagebrush in sprayed areas is in a
recuperative stage. It would appear that percentages of sagebrush killed
in strip sprayed areas were poor, while kill was excellent in the block spray
areas. However, this could not be adequately tested as 100 ft2 plots in
sprayed strips frequently were at the edge of the strip where spray application
was not uniform. Sagebrush kill did appear to be highest and similar to that
in block sprayed areas in the center of the strips. Thus, kill of sagebrush
within strips was a continuum with gradual but definite changes from the
outer edges to the center.
Observed decreases in sagebrush and forb frequency and crown intercept of
sagebrush are important in terms of sage grouse use of sprayed areas. Due
to sampling and plant identification problems, data were not available on
forb frequency by species for all years. It has been well documented that
total forb cover is not as important to sage grouse as cover of highly
preferred forb species (Klebenow and Gray 1968, Autenrieth 1970, Martin 1970,
Peterson 1970). Decreases in preferred forbs along with decreases in sagebrush frequency and cover are undoubtedly involved in the marked reduction
in sage grouse use of the treated areas.
It is important to note that statistical analyses of the vegetation data
could not be conducted. This was the result of inadequate sampling, failure
to meet treatment guidelines which caused some plots to be useless (one plot
had to be moved as 65-9 became 66-A), improper placement of plots into height
classes (four plots were discarded due to improper height class placement in
1965 and 1966), and lack of consistency in classifying some forbs, shrubs

�-37-

Table 2. Relative frequencies of vegetation in sprayed and control
plots, North Park.
Relative Freguency (%)
Growth
Form

&lt;5

in.

5-10 in.

&gt; 10 in.

Year and Treatment

Forbs

Grasses

Sagebrush

Other
Shrubs

1965

Control

48.8

33.7

8.9

8.6

1966

Control

49.8

33.4

7.1

9.6

1966

Sprayed

43.2

43.3

2.6

6.2

1973

Control

42.4

32.3

12.9

12.0

1973

Sprayed

35.3

43.0

9.6

12.0

1965

Control

41.8

35.9

10.2

12.1

1966

Control

46.9

33.2

9.6

9.9

1966

Sprayed

36.8

49.0

4.5

9.8

1973

Control

36.1

33.9

14.7

15.1

1973

Sprayed

31.0

46.8

9.3

12.8

1965

Control

39.9

48.3

11.4

1.3

1966

Control

28.8

51.6

16.4

3.2

1966

Sprayed

26.9

59.9

10.1

3.1

1973

Control

23.4

51.6

18.2

6.7

1973

Sprayed

19.2

53.3

14.5

12.9

�-38Table 3.

Sagebrush

Growth Form

&lt;5

in.

5-10 in.

&gt;10

in.

11
ND

No data.

intercept

in sprayed and control plots, North Park.

Year and Treatment

Length (ft.) InterceEted/lOO sg. ft.
Seedling-Young
Mature-Decadent
Dead

1965

Control

0

10.9

0.3

1966

Control

0

10.3

0.8

1966

Spray

0

3.9

6.4

1973

Control

0

8.0

ND .!.I

1973

Spray

0.3

3.8

ND .!./

1965

Control

0

14.0

0.5

1966

Control

0.1

15.1

1.2

1966

Spray

0

6.5

10.9

1973

Control

0.3

7.4

ND .!.I

1973

Spray

0.6

4.2

ND 1:/

1965

Control

0.9

34.8

3.9

1966

Control

0

40.5

5.1

1966

Spray

0.1

13.8

23.9

1973

Control

0.3

13.7

ND1/

1973

Spray

1.6

7.7

ND1/

�-39-

(Guterrezia spp.) and treatment type. Most of these problems resulted from
changes in personnel as no single individual was responsible for the project
throughout its duration.
While these problems prevent detailed statistical
analyses of the vegetation data, it is quite clear that spraying of sagebrush communities resulted in changes in species composition and cover.
These obvious changes in vegetative composition caused marked changes in
sage grouse distribution and use within the treated areas.

Distribution

of Strutting

Grounds

Eighteen known strutting grounds were active in North Park in 1963 of which
seven were located in the Lake John area.
Rogers (1964) reported 12 active
strutting grounds in the Lake John area in 1957-59.
None of the five inactive
grounds had been active during the 4 to 6 years prior to 1963 and it is
possible that these grounds were only temporarily used.
It is possible that
location of these grounds shifted and were not detected.
Gill (1965) reported
that the location of strutting ground No. 11 (Boettcher Flats) moved from
1963 to 1964.
Counts in 1963 were made at the original location, while those
in 1964 were from the new location.
No birds were counted on either of these
sites in 1965 or after 1970 (Table 4). It is probable that this ground moved
and has not been relocated by observers.
Nineteen strutting grounds (Appendix B) were known to be active in North Park
in 1974, six of which were in the original study area (Fig. 3). However, only
four of the seven grounds used in 1963 in this area were still in use. The
two previously unknown grounds within the original study area were located in
1973.
One additional ground was located in the northwest quarter of North
Park in 1973, about 4 miles east of the southeast corner of the original study
area. In 1974 searches in the north half of North Park resulted in two grounds
being located.
One was 4 miles east of the Lake John study area and less than
2 miles north of the new ground located in 1973. The second ground was approximately 3 miles southeast of Cowdrey, roughly 7 miles east of the original
study area. While number of known active strutting grounds in North Park
remained constant from 1963 to 1974, location of grounds used changed.
It is
unfortunate that total numbers of grounds used in a given year within North
Park are not definitely known.
It would be ideal if, as locations of grounds
change or new grounds are established, these grounds would be immediately
located,
However, this does not appear to be possible.
Because of the problems with location of all active strutting grounds, it is
difficult to relate numbers of strutting grounds in an area to vegetation
disturbance.
Considering only the original Lake John study area, number of
known active strutting grounds decreased from seven to six from 1963 to 1974,
with three being abandoned, while two new grounds were located.
It
is probable that abandonment of the three grounds was related to sagebrush
treatment in the area.
It is also probable that new grounds formed due to
changes in sage grouse distribution caused by spraying.

�-40-

Fig. 3. Location of strutting grounds in North Park active in 1975. Includes
one grolli~dlocated in May 1975, and one which was active all years from
1960-75 except 1974.

�-41-

The existence of strutting grounds, formerly unknown, in surrounding areas
underlines a weakness of the original study area designation. Even though
strutting grounds ceased to be active or decreased in number of birds
attending during the study, interpretation of changes is hampered by not
knowing when adjacent, new grounds first supported birds or if there are
other grounds currently in existence. Quite possibly the new grounds represent relocations of old grounds or distributional shifts in response to
changing land use. Considering all of North Park, the number of known active
strutting grounds was greater (by one ground) in 1974 than in any previous
year of the study. However, searching intensity in areas other than Lake
John was greater in 1974 than in other years.
The shift in strutting ground locations documented in North Park indicates
that although some strutting grounds are used for long periods (Patterson
1952), a strutting ground site cannot be considered to be permanent. Thus,
sampling procedures for spring breeding population numbers must acknowledge
that strutting ground complexes are dynamic and should incorporate procedures
to search for unknown grounds every year. Existence of strutting grounds of
any size and thus one or more mating sites per ground may be more important
in population maintenance than number of males on a given ground.
Breeding Activities
Strutting Ground Attendance
Counts of the number of males on all known strutting grounds were made from
1963 through 1974 and were available for a few grounds from 1959-1963. The
highest count on a ground was considered the maximum number of males attending
the ground and thus, an index to the breeding population. Trends in numbers of
males counted in North Park are presented in Fig. 4, while actual counts for
each strutting ground are presented in Table 4.
It is readily apparent that maximum counts of male sage grouse on strutting
grounds fluctuated from 1963 (and earlier) to 1974. Considering only counts
of males on strutting grounds on the original study area, sage grouse numbers
decreased from 372 in 1959 to 177 in 1973. This decrease is significant
(P &lt; 0.05). Counts were high in the 1959-1964 period (range 337 to 446), low
in 1965 (166), moderately high in 1967-1971 (range 261 to 326) and low in
1974(177). The low number in 1965 occurred prior to any spraying in the study
area. Initial inspection of all available data suggest that the decrease in
1965 was real and not due to observer inefficiency (R. B. Gill personal comm.).
Check station data for 1964 indicate that 41 percent of the fall harvest was
comprised of juveniles. Thus production in 1964 was lower than in 1963,
assuming juvenile vulnerability did not change. However, production was not
low enough to be responsible alone for the observed decrease in spring counts
in 1965. It is doubtful then, that the apparent decrease in 1965 was real.
The Monahan Draw strutting ground was located within the large block spray
area. Numbers of males counted on this ground remained stable from 1965 to
1971 (range 52 to 81 in 1965 and 1971, respectively), significantly (P &lt; 0.05)
decreased in 1972 (from 81 in 1971 to 10 in 1972) and remained low in 1973
and 1974 (11 and 3, respectively). This area was sprayed in 1965 and a

�Table 4.

Maximum number of males on strutting grounds, North Park, 1959-1974.

Strutting Ground

1959

1960

1961

1962

1963

1964

1965

1966

1967

1968

1969

1970

1971

1972

1973

1974

Alkali Lake 1./

214

130

96

219

216

120

53

48

63

57

82

66

88

96

81

72

Boettcher Lake
Jct. 1/

146

167

109

71

85

99

52

85

68

71

107

97

96

62

59

46

0

12

50

5

11

7

0

9

24

12

5

4

0

0

0

0

Boettcher Flats 1/
Canuck

29

Coalmont

10

6

17

9

23

17

6

10

11

15

17

62

46

57

47

27

Cowdrey In 1./

1

43

4

0

19

25

9

14

10

11

8

8

10

0

0

0

Cowdrey 112 1/

0

10

5

7

0

0

0

0

4

0

0

0

0

0

0

0

Cowdrey #3 1./

6

5

0

0

1

0

0

3

9

2

6

7

5

0

0

0

Cowdrey 114 ])

-

14

0

0

1

0

0

0

0

0

0

0

0

0

0

0

30

19

I

I

Cowdrey #5 1./
Deer Creek

23

35

0

71

65

58

12

83

-

84

63

53

44

39

37

11

Delaney Butte

7

13

25

23

12

14

3

5

11

0

0

9

22

9

4

13

75

52

52

69

-

-

-

66

57

53

67

62

Fish Hatchery

+'
I-.)

Hound

-------------------------------------------------------------------------------------------------------------------

69

�Table 4.

Maximum number of males on strutting grounds, North Park, 1959-1974 (continued).

Strutting Ground

1959

1960

1961

1962

1963

1964

1965

1966

1967

1968

1969

1970

1971

1972

1973

1974

Lost Cr.

ill

57

39

58

98

137

53

60

58

71

75

68

25

33

13

0

Los t Cr.

tI2

54

54

56

51

63

44

24

37

48

24

31

15

19

10

1

14

12

0

16

0

0

0

0

0

0

0

0

0

0

0

Lake John 112 1./

17

77

28

15

13

4

8

2

2

15

16

21

0

0

0

Monahan Draw 1./

19

2

52

22

58

52

64

74

53

80

57

81

10

11

3

184

52

129

120

108

71

45

36

44

71

127

76

64

36

33

1/
Lake John III -

Ridge Road

7

162

10

Roth

I

12

Riley 1./

15

i

19

30

36

31

39

26

21

-

27

53

49

35

24

46

33

Spring Cr. 112

25

38

55

60

61

24

46

-

74

163

69

39

34

39

15

Spring Cr. #3

18

0

0

1

0

0

0

-

0

0

0

0

0

0

0

Spring Cr. #4

14

38

12

16

13

5

15

-

16

0

23

15

21

9

10

Spring Cr. III

47

1/
Wat tenberg III -

0

0

7

5

3

5

0

2

0

1

2

0

0

0

0

0

1/
Wattenberg 112 -

0

5

1

8

7

22

0

3

7

52

36

19

25

4

24

22

38

37

Walden

1/

t;

Located in original Lake John study area.

�0
0
0

,
en

0
0

Fig. 4. Maximum
1959-1975.

eo

0
0

0
0
~

0::0

WW

z

m~

:::J::&gt;

z8

::&gt;w

~oo
:2!='
-ct
)(~
&lt;(LL

0

I

10

,
0
0
.q

10

I'~
rt)

f"-

I'-

(.\J

I'f"-

0

en
W

s

1'-0:::
(0)-

&lt;.O&lt;t
&lt;.OW

&lt;.0

U)

""CD
r6
(\f

W

&lt;D
0
&lt;.0

m

grounds in North Park,

0
0

0
0
&lt;D

numbers of males on strutting

�-45-

decrease was anticipated in 1966 or 1967. The decrease did not occur until
1972, long after most of the sagebrush in the area had been killed.
Reasons
for the delay are not clear, nor is it known if the decrease was related to
spraying of sagebrush.
However, it would appear that a cause and effect
relationship does exist between sagebrush spraying and the overall decrease
in numbers of males counted in the original study area.
Strutting

Ground

Count Analyses

Although counts of strutting males have long been used as an estimate of
yearly populations, scientific effort has not been directed toward evaluating
the precision and accuracy of each count. Analyses of the methods used in
this study may explain some of the fluctuations evident in Table 4 and Fig.
4. Personnel changes were frequent throughout the study, and little uniformity in observation methodology was maintained.
Observations were at times
made from distances of 0.25 to 1.0 mi distant.
Counts made from farther
than 100 yds are highly questionable because of inability to see all birds
well enough to distinguish between sexes. This would inflate male counts
if large numbers of females were present.
Another problem was the lack of
uniformity in spacing counts throughout the strutting season.
Frequently,
all three required counts would be made on an easily accessible ground
during early April, whereas, the earliest count on a ground which was
temporarily
inaccessible or snowbound would not be accomplished until midMay.
If the pattern of male attendance was constant, this would pose no
problem.
However, male attendance patterns, though highly variable from day
to day, follow a general trend (Stanton 1958, Patterson 1952, Eng 1963).
During late March and early April adult males predominate on strutting grounds.
Hens come to grounds from 1 April to 21 May, but peak numbers usually occur
in a 7 to 10 day period sometime during 7-25 April.
High numbers of adult
males coincide with peak numbers of females.
Shortly after hen numbers are
highest, yearling males appear in large numbers and maximum male counts are
obtained at this time (Eng 1963).
Adult male numbers steadily decline
following peak hen attendance (Fig. 5). Thus, it is conceivable that a
series of early or late counts would miss the peak of male attendance.
Further complications result from methods of reporting counts, primarily
during the 1959-62 period.
During these years, accurate distinction
was
not made between counts of males and total birds, the latter which
included females.
Most counts of females prior to 1974 were incidental to
male counts and thus are incomplete for analysis as a population index.
In
1974 greater effort was expended during the peak of hen attendance to obtain
accurate male counts as adult male attendance is supposedly most stable
during this period (Patterson 1952).
Five grounds in the northern half of
the park were counted from four to seven times during the 9-day period of
peak hen attendance (Table 5). Had males on each ground been counted on the
peak day during the week of maximum hen attendance. the difference from the
seasonal maximum of males would only have been 3.4 percent (246 vs 238).
However, had counts coincided with minimum male counts during the peak of hen
attendance, the difference from the seasonal maximum would have been 26.9
percent.
Variation during the 9-day period when numbers are considered most
stable was 24.3 percent.
These are minimum values as lower and/or higher
numbers of birds could have been present on days not counted.
In reality,
observers seldom obtain one count on each ground during the period of peak
hen attendance as one person often has responsibility
for counting 7 to 10

�-46-

ADULT MALES

SUBADULT MALES
FEMALES

240

..........

-----

220
200
180
160

o

NUMBERS

..
..

\

\
.
\
:
\~O

OF
BIRDS

o.

t
:\

:q
. \
\
\
~

MAR.

MAY

JUNE

Fig. 5. Sage grouse attendance on all Lake John area strutting grounds
combined, 1963.
Strutting ground attendance patterns in 1963 were typical,
but chronologically 2 weeks earlier than in most years (from Gill 1965).

�Table 5.

Variation in male numbers attending selected strutting grounds during April and May, 1974.

Strutting Ground

Max. Male
Count

Third Lowest l/Max. Male Count
in Hen Peak Week
Count

Possible
Seasonal Variation
Percent
- (2)
(1)
Min. Male Count
(1)
in Hen Peak Week

Possible Peak
Week Variation
Percent
(3) - (4)
(3)

Alkali Lake

72

33

69

48

54.2

36.4

Boettcher Lake
Jct.

46

19

44

42

58.7

4.5

Hound l:..l

69

59

69

57

14.5

17.4

Walden

37

19

37

19

48.6

48.6
I

.j::'-

Wattenberg /12

22

14

19

14

36.4

26.3

Totals

246

144

238

180

41. 5

24.3

1/

Theoretically this count could represent the maximum male count for the year if only three counts were made.

])
Ground first located during peak week of hen attendance, thus no counts during first 4 weeks of activity.

'-I
I

�-48-

grounds.
Also, hen attendance is surprisingly synchronous throughout North
Park.
Considering the spread in counting dates during the two month mating
season, variation in numbers of males from 35 to 60 percent (Table 5) can
be attributable to daily male attendance patterns.
The low variation
documented for the Hound strutting ground is likely due to all counts being
made during or shortly after « 8 days) the week of peak female attendance.
Since 1959 fluctuations between years in total male counts have exceeded 35
percent 6 of the 16 years possible and exceeded 60 percent twice.
It is
questionable if counts of strutting males can indicate changes in sage grouse
populations due to the high degree of variation in daily attendance.
Therefore, while numbers of males counted in North Park have decreased, and by
association, the entire population has decreased, the reliability of using
strutting male counts to document increases or decreases in population size
is unknown.
It is not known whether or not the apparent decrease was real.
Although male sage grouse have been intensively studied during the breeding
season throughout their range, little is known of attendance patterns of
individual males.
Hartzler (1972) documented 50+ days of consecutive
attendance on a strutting ground by three individually marked male sage grouse.
All of these males were on central territories and two occupied primary
mating spots.
No data on the regularity of attendance of surrounding males
were presented although this segment of the male population contributes most
to total counts of males.
In recent years intensive studies of other grouse
species have concentrated on the apparent non-breeding segment of the male
population.
Non-displaying males have been documented for other lek grouse;
black grouse (Lyrurus tetrix) (Robel 1969) and sharp-tailed grouse (Pedioecetes
phasianellus)
(Rippin and Boag 1974).
It is possible that variation between years in counts of male sage grouse on
strutting grounds in North Park could have reflected real population fluctuations (Fig. 4). However, considering all variables involved, it is also
probable that all changes in male counts were not caused by population fluctuation alone.
Patterson (1952) suggested that all males in a population
regularly visit strutting grounds.
Until data are available to support or
reject this hypothesis, counts of strutting males should be closely examined
to ascertain when counts were made in relation to period of peak hen attendance on grounds.
It is presently doubtful that two or three counts of males
per ground per breeding season have any management implications.
Movements

Between

Strutting

Grounds

Recaptures and observations of 228 marked male sage grouse indicated that
most (193 of 228, 84.6 percent) exhibited high fidelity to the ground where
originally trapped (Table 6). Thirty-five males were known to have changed
grounds between or within years with most (77.1 percent) inter-ground movements occurring between years.
If observations and recaptures of marked
birds within initial year of banding are excluded, then 27 of 101 (26.7
percent) marked birds changed strutting grounds in years after their initial
capture.
The remainder (73.3 percent) did not change grounds between years.
While sample sizes are small, adults exhibited greater fidelity to initial
banding location than did subadults.

�-49-

Table 6. Pooled
grounds. 1../

observations

and recaptures

of marked

males on strutting

Number Observed on Different
Ground on Which Banded in Year
x+4
x+3
x+l
x+2
x

Age

Number Observed on Same
Ground on Which Banded in Year
x+4
x+l
x+2
x+3
x

Adult

18

19

2

0

2

5

2

0

0

0

Subadu1t

5

6

1

0

1

3

7

3

1

0

Unknown

96

43

0

0

0

0

14

0

0

0

1/ Undoubtedly
not individually

some individuals were observed
color marked, identification

more than once.
Since birds were
of individuals was not possible.

Inter-ground movements documented in North Park emphasize the importance of
locating and counting all strutting grounds in an area. Decrease in attendance on one ground may be a result of inter-ground movements between years.
Such a decrease may be compensated by increases on neighboring grounds.
Interground movements within North Park could not be related to sagebrush treatment
within the original study area. Unfortunately,
few, if any, of the males on
the Monahan Draw strutting ground were marked prior to the decrease in 1972.
Whether the males from this ground were absorbed into existing grounds or
established a new ground is unknown.
Trapping
Numbers

Captured

and Banding

and Banded

Trapping efforts were not uniform in all years of the study.
Numbers of birds
banded by year, age and sex are presented in Table 7. About 73 percent (72.7)
of the birds banded for which sex was ascertained were males even though Beck
(1975) estimated that males comprised only 40 percent of the spring population.
This disparity was the result of greater vulnerability of males to capture
because of their tendency to roost at night on strutting grounds throughout
the 7 to 9 week strutting period.
Relatively few females roosted on strutting
grounds at night during the brief 7 to 10 day period that large numbers of
hens were present in early morning hours.
In addition, males were more noticeable at night because of larger size and consipicuousness
of their white
breast feathers.
Most females caught in 1969 and 1974 were captured with
cannon nets placed near primary mating spots on strutting grounds.
Over half
of the females caught in 1973 were spotlighted at night in areas where birds
were concentrated by late spring snows prior to strutting grounds becoming
snow-free.
Weather conditions as experienced in 1973 appear to be unusual
and methods need to be developed to capture females which are not dependent
upon unusual weather conditions.

�Table 7.

Numbers of sage grouse banded in North Park, 1963-1974.!1

Year

Adult Males

1963

44

1964

Subadult Males

Adult Females

Subadu1t Females

Chicks

Total

11

5

3

0

63

47

22

4

9

0

82

1965

41

15

9

3

0

68

1966

6

10

6

3

8

33

1967

50

27

7

5

0

89

1968

40

21

11

6

7

85

1969

22

19

13

20

1

75

1970-72

0

0

0

0

0

0

1973

89

80

54

41

0

2871.1

1974

89

54

29

20

0

192

Totals

428

259

138

110

16

974

I

VI

0
I

II
- Grouse banded but removed from the population by project personnel prior to the hunting season in year of
banding are not included.
21
Includes 9 males and 14 females of unknown age.

�-51-

Age ratios of males captured were undoubtedly not representative of the male
segment of the population because of differences in behavior and seasonal
attendance on strutting grounds.
Adult males appeared to be more wary than
yearling males, but this was a subjective evaluation.
The preponderance of
adults early in the strutting season and of subadults during the last three
weeks can noticeably affect the age composition of captured birds, depending
on when samples were obtained.
Capture efforts were not continuous throughout the strutting season during 1963-69.
In most years there was more effort
early in the strutting season, probably resulting in more adult than yearling
males being captured.
Also, if adult males attend grounds more regularly
than yearlings, the result might be a disproportionate
percentage of adults
being captured.
Analysis

of Capture Techniques

Over 90 percent of all sage grouse were captured by spotlighting and netting.
The use of the battery pack with hand-held nets in 1973 and 1974 greatly
increased efficiency of trapping crews and was the most successful method
employed.
Drive nets for trapping broods were unsuccessful in North Park,
probably due to few known summer concentrations of hens with broods.
Since
there is one acre of irrigated hay meadow for every two acres of sagebrush
in North Park, and since hay meadows are well distributed, the probability
of concentrations of grouse in one area is not high.
Use of cannon nets on
strutting grounds was partially successful.
However, activity sites frequently changed once nets were in place.
This problem may be reduced by
placing nets near primary mating spots prior to any strutting activities.
Primary mating spots occasionally shifted, but the apparent high degree of
permanence of these sites may allow early placement of nets to habituate
birds to their presence.
The major deficiency in trapping techniques was
the inability to capture hens and chicks in representative numbers.
Analysis

of Techniques

for Age and Sex Determination

Analysis of the rectrix shape method of aging indicated a high degree of
accuracy.
Using primary shape and wear as the standard, only 3 of 143 males
(2.1 percent) captured in 1974 were placed in the wrong age class based on
shape of rectrices.
Each of the three birds erroneously aged (classified
as juveniles) by rectrix shape were adults, based on primary appearance.
Although highly accurate with birds in hand, accuracy probably declined when
aging free ranging males at distances of 10 to 40 yds. Attempts to age all
males on strutting grounds by retrix shape were unsuccessful, because of the
great amount of movement and change of position.
However, the method was
useful for aging specific males at distances less than 40 yds.
Use of width of white band on the breast of males as an age criterion was
discontinued after 1966. This technique appeared to be accurate only when
the preceding year's hatch was early, but accuracy between years was not
consistent.
Use of Crunden's (1963) method for age and sex classification
of wings collected during the hunting season was discontinued in 1974 because
it did not distinguish between yearlings and adults, thus available data were

�-52-

not fully utilized. Eng's (1955) method based on primary wear was used in
1974. For comparative purposes, wings from 40 juveniles sexed by gonadal
inspection were collected from hunters in 1974 and lengths of primaries were
recorded. Measurements of these 23 males and 17 females fit within Eng's
(1955) limits. Wings of yearling and adult birds can be accurately classified to sex by size (subjective) and use of primary measurements (objective).
Aging and sexing techniques resulting from the 1974 analysis and review of
literature were summarized by Beck et al. (1975).
Analysis of Sage Grouse Weights
Weights of sage grouse captured were taken in most years (Table 8). While
sample sizes were variable between years, differences were slight and data
from all years were pooled. In general, weights of males increased immediately prior to initiation of strutting followed by gradual decline through
April and May. Yearling males weighed less than adults, but gained weight
until early May with weights declining in late May. This pattern closely
follows attendance of yearling males on strutting grounds (Fig. 5).
Females typically gained weight until early May with weights of yearling
females lagging behind those of adults. Weight gains by females in spring
were related to preparation of the reproductive organs for egg laying and
deposition.
While differences in changes of weights were not significant (P&gt; 0.05) for
the pooled samples, declines in weight for both age classes of males in 1974
were significant (p &lt; 0.05) for each two-week period. The importance of
weight declines through spring may be manifest in higher mortality for males
showing the greatest weight loss. Stress of strutting, coupled with weight
loss, may result in the lower survival of males as indicated by the unhalanced sex ratio in the harvest reported in this study and in winter (Beck 1975).
Due to inadequate samples, weight differences between birds trapped in the
original study area and elsewhere in North Park could not be tested. Effects
of sagebrush treatment on weights of sage grouse are unknown.

Reproduction
Nest Densities
Nest densities were estimated from searches of 10 (1964) or 5-acre (1965, 1966,
1973) plots. Results are presented in Table 9. Due to inconsistencies in
sampling and reporting, analyses are limited. Data from 1965, 1966 and 1973
are not comparable to those collected in 1964 as no attempt was made in the
1965-1973 period to sample all potential nesting habitat.
Densities of nests found in 1964 and 1965 (Table 9) were high and probably
were not representative of all sagebrush areas in North Park. It is unclear
why more nests were found in sprayed than unsprayed areas in 1966. Much of
the sagebrush was dead in the sprayed areas, but cover was considered good
as little deterioration of the brush was noted. While this may have influenced nest site selection, due to small samples) apparent differences could

�Table

Mean weights

8.

of captured

sage

grouse

by time period,

North Park,

all

years

Females

Males
1-

2+

combined (grams).
Chicks
Unclassified

Sex

Time Period

1-

2+

January

15-31

2554 (

2)

February

1-14

-

2724 (

1)

-

1419 ( 1)

February

15-28

2298 ( 4)

2724 (

3)

1291 ( 4)

1419 ( 5)

March 1-14

2610 ( 1)

3008 (

1)

1816 ( 1)

March 15-31

2426 ( 4)

3284 (

8)

-

1532 ( 1)

April

1-14

2936 (25)

3231 ( 81)

1507 (38)

1723 (45)

April

15-30

2812 (71)

3139 (103)

1603 (41)

1778 (40)

May 1-14

2834 (60)

3146 ( 71)

1682 ( 9)

1874 (10)

May 15-31

2796 (47)

3110 ( 57)

1725 ( 5)

1803 ( 7)

June 1-14

-

2546 (

-

1243 ( 1)

2399 (

1466 ( 3)

1358 ( 9)

321 (12)

-

1423 ( 7)

438 ( 2)

-

577 ( 2)

1178 ( 1)

878 ( 5)

June 15-30
July

1-14

2283 ( 3)

July

15-31

-

August 1-14
August 15-31
September

15-30

1)

-

3)

I

V1

w
I

1617 ( 4)

1244 ( 2)

�-54have been due to chance alone.
It would appear that block spraying effectively curtailed nesting while strip spraying did not. However, as
mentioned previously, kill of sagebrush in strip sprayed areas was highest
in the center and lowest along the edges. Established nest plots in strip
sprayed areas frequently encompassed a continuum of sagebrush ranging from
almost total kill to almost no kill.
In contrast, sagebrush within the
nest plots in the block sprayed areas was mostly dead. It is concluded that
block spraying resulting in a high percentage of the sagebrush being killed
is detrimental to sage grouse nesting through loss of cover.
This is especially true once the dead sagebrush has deteriorated.
Strip spraying with
partial kill of sagebrush appears to have little effect on nesting.

Table 9.

Results of nest searches

in the Lake John area, North Park.

Area

Year

Acres Searched

Nests Found-

Nest Densl y
(Nests/Mi Z )

Control

1964

410

21

32.0

Control

1965

300

23

49.1

SprayeCF

1966

135

8

37.9

Control

1966

165

2

7.8

Block sprayed

1973

70

0

0.0

Strip sprayed

1973

45

2

28.4

Unsprayed

1973

45

2

28.4

1973

120

5

26.7

2/

strip

Control

1/

1/ Only includes nests of the year.

2/ Includes

block sprayed

Nest Placement

area and sprayed

strips.

and Success

Sage grouse nests were found under sagebrush at 91.9 percent of the 198
nest sites located (including both nests of the year and nests of earlier
years).
The 16 nests not under sagebrush were under greasewood or rabbitbrush.
Nesting success from examination of nests located was estimated to range from
20 to 30 percent in 1964, 1965 and 1966. No estimate is available for 1973
due to lateness of nest search and conditions of egg fragments found. Nest

�-55-

predation was attributed primarily to badgers (Taxidea taxus) and ground
squirrels (Spermophilus richardsoni).
Nest searches in 1965 and 1966 were
conducted from 1 to 3 months after peak of hatching and thus, any nest
disturbed by a predator in the interim may have been recorded as unsuccessful.
Since some nests reported as being predated may have been successful, nesting
success estimated from discovery of nests must be interpreted with caution,
especially in view of small samples and higher visibility of egg remains in
disturbed nests.
Census of Broods
Broods were censused in the Lake John study area from 1963 through 1966 and
in 1968 and 1973 following routes described by Gill (1965). Mean brood sizes
for complete broods were 3.7 (N = 52) in 1963, 2.9 (N = 40) in 1964, 3.4
(N = 28) in 1965, 4.0 (N = 45) in 1966, 4.7 (N = 17) in 1968, 3.7 (N = 29)
in 1969, and 4.1 (N = 19) in 1973.
Basic problems with brood counts during
the study period related to recording of dates and how many different broods
were observed.
Interpretation is difficult as easily sampled areas such as
the accessible meadows in the Lake John area were sampled from four to twelve
times per summer from 1963 through 1966.
Thus, a relatively few number of
broods observed each trip could account for all of the sightings.
For example,
the 52 broods observed in 1963 could possibly have been only seven different
broods, as seven was the maximum number of broods seen on anyone
complete
set of the five routes.
Sampling of limited areas thus inflates the importance of a few broods and creates artificial concentration areas which may
be artifacts of methods used rather than bird selection.
The period of maximum brood observations was relatively constant each year.
Of 215 broods observed in all years, at least 86 (40 percent) were seen in
the 11th and 12th weeks after the estimated peak of female attendance on
strutting grounds.
Therefore, brood counts should be concentrated in the
period from 11 to 13 weeks after the peak of hen attendance on strutting
grounds.
Percentages of females accompanied by chicks varied from 18.2 to 34.4 from
1963 through 1973 (34.4 in 1963, 18.2 in 1964, 30.1 in 1965, 22.3 in 1966,
20.0 in 1969, 32.4 in 1973). These percentages appear to be low and are
probably more a reflection of sage grouse behavior than actual production
success.
Since unsuccessful hens form flocks in mid to late summer and
remain in sagebrush uplands, such groups are probably more conspicuous than
hens with broods which utilize heavy vegetation along meadows.
Primary feather molt patterns of harvested yearling and adult hens were
recorded in 1974.
Since successful hens initiate molt of primaries later than
unsuccessful hens, estimates of nesting success of females can be obtained
from examination of primary feather molt patterns.
Females unsuccessful in
hatching their clutch normally have primary feather molt patterns similar
to those of males harvested in the same time period.
Hens which are successful in hatching their clutches but lose all of their chicks within several

�-56weeks of hatching are intermediate in primary feather molt and thus
complicate use of this technique.
In 1974, 91 of 92 yearling and adult
males had molted through primary 8, while 160 of 254 (63 percent) yearling
and adult females had molted through primary 8. Thus, the minimum estimate
of nesting success in 1974 based on primary molt was 37 percent.
Patterson (1952) suggested that roadside trend counts were accurate indices
of production but presented little supportive data. Yet brood routes do
not appear to be a practical way to obtain adequate estimates of production
in North Park, principally because of the widespread abundance of excellent
brood habitat.
Obtaining maximum number of observations of broods throughout the park rather than on specified routes appears to be more suitable and
should be tested.
Brood routes conducted by Wildlife Conservation Officers
in North Park are only censused three times each summer and none of the
20-mile routes sample more than 2 miles of good meadow-type brood habitat.
These three routes (one per officer) seldom produce more than 1 to 3 broods
per year, and thus are of little quantitative value in estimating production.
Harvest

Analysis

Check Stations

Hunter check stations designed to obtain data on hunting pressure and success,
harvest trends, age and sex composition of the harvest and band recoveries
were operated during all years of the study (Table 10). From 1963 through
1973 stations were operated only at Walden and Cowdrey to sample hunters
from the northwest quarter (original study area) of North Park.
In 1974,
stations were operated at the four major exits from North Park to sample all
hunters from throughout the park.
During the l2-year period, hunting pressure was extremely variable.
While
reasons are not known, it is possible that hunter distribution changed.
In
1963 most of the hunting pressure and harvest in North Park was estimated
to have occurred in the Lake John area. Data collected from more extensive
check stations in 1974 indicated that hunting pressure was relatively evenly
dispersed throughout the park and the Lake John area was hunted by only 18
percent of the hunters who accounted for 16.4 percent of the checked harvest.
If in fact, hunter pressure and kill decreased in the original study area,
this would suggest that numbers of sage grouse available to hunters also
decreased.
Unfortunately,
data to support this suggestion are not available.
It is apparent from data in Table 10 that both birds per hunter and time
required to kill one bird were variable.
In general, the trend in hunter
success was down (except in 1974) while it progressively took more time to
harvest one bird.
These data support a hypothesis that sage grouse numbers
in North Park decreased from 1963 to 1974. This apparent decrease is
undoubtedly the result of increased habitat alteration.

�Table 10.

North Park sage grouse hunter checks, 1963-1974.

1/

Juvenile Percent
Juvenile
Hens

Unclassified
(Dressed) Total
Birds
Birds

Birds
per
Hunter

Hours
Hunted
per Bird

Year

Bag
Limit

Hunters
Checked

Hours
Hunted

Adult
Cocks

Adult Percent Juvenile
Hens Adults
Cocks

1963

2

592

2,460

62

150

42

113

181

58

- 3/

506

1.03

4.86

1964

2

217

624

25

81

59

28

45

41

- ]j

179

0.82

3.49

1965 ]j

2

150

626

27

30

49

26

26

51

- 1/

116

0.77

5.40

1966

2

306

1,227

31

116

56

45

71

44

- ]j

263

0.86

4.67

1967

2

300

1,177

50

127

67

42

46

33

- 1/

267

0.89

4.41

1968

2

546

2,604

80

135

42

156

141

58

- 3/

512

0.94

5.09

1969

2

662

2,936

79

180

70

37

74

30

199

569

0.86

5.16

1970

2

564

2,617

58

89

55

53

66

45

-

266

0.47

9.84

1971

2

357

1,802

28

67

51

54

39

49

76

264

0.74

6.83

1972

2

452

2,377

50

69

67

40

18

33

76

253

0.56

9.40

1973

2

366

1,599

24

54

52

29

42

48

44

193

0.53

8.28

1974

2

730

5,912

94

254

50

171

179

50

87

785

1.08

7.53

}j
Data from 1963 through 1973 are based on Walden and Cowdrey check stations operated during opening weekends.
(These stations checked harvested birds primarily from the northwest part of North Park) . Data from 1974 are based
on four check stations (State line, Gould, Muddy Pass and Willow Creek) operated during the opening weekend.
(These stations sampled all of North Park).
])
A one day season; hence only a one day check.

1/
Composition of kill projected from wing samples; hence no unclassified birds.

I

V1
-...j
I

�-58Hunter

Questionnaires

Prior to 1974 total harvest and hunter numbers in North Park were ascertained
through the annual state wide survey of small game hunters.
Data from prior
to 1968 are not available as surveys were not designed to estimate harvest
and hunter numbers in discrete units or areas. Available data on numbers
of hunters and harvest of sage grouse in North Park are presented in Table 11.

Table 11. Hunter pressure and sage grouse harvest
wide small game survey. 1/

Year

in North Park from state-

Number
Number

of Hunters

of Sage Grouse
Harvested

1968

1,029

1,861

1969

1,926

3,398

1970

1,606

2,172

1971

1,003

1,962

1972

1,566

2,846

1973

1,171

2,007

1974

1,759

2,509

1/ Not available

for years prior to 1968.

In 1974, all sage grouse hunters in North Park were required to obtain and
have in possession a free permit.
A total of 1,184 permits were issued.
Most hunters (43.3%) were from the metro-Denver area (Adams County 5.1%,
Arapahoe County 5.6%, Denver County 16.9%, Jefferson County 15.7%). Only
11.1 percent of the hunters originated from North Park, with 21.7 percent
from Larimer County, 9.5 percent from Boulder County, and 5.0 percent from
Weld County.
All other counties each contributed less than 2 percent of the
total hunters, and only 9.5 percent of all hunters.
Questionnaires were sent to all permittees immediately following the 1974
sage grouse season in North Park.
Responses were received from 886 hunters.
In mid-October a followup letter was sent to all non-respondents
and 175
additional responses were received.
In all, 1,061 permittees (89.7%)
responded (Table 12). Only 12 questionnaires were returned for insufficient
address.
Mean values calculated for hunters responding to the followup
letter were used to project for the 123 non-respondents.

�-59-

Table 12.

North Park sage grouse hunter questionnaire data, 1974.
Projected For
Non-Respondents

1,184

123

1,061

All
Permittees

No. in sample

886

175

Percent total of
permittees

74.9

14.8

89.7

10.3

100

No. hunters

749

124

873

87

960

Percent hunters

84.5

70.9

82.3

70.9

81.1

No. non-hunters

137

51

188

36

224

Percent non-hunters

15.5

29.1

17.7

29.1

18.9

No. successful
hunters

398

70

468

49

517

Percent hunters
successful

53.1

56.5

53.6

56.5

53.9

No. hunter days

199

1,217

Days/hunter
No. sage grouse
bagged

1.62

1.60

151

859

1,555

139

1,416
1.62

1.60

1,116

106

1,010

1.62

Grouse/permittee

.97

.86

.95

.86

.94

Grouse/hunter

1.1

1.2

1.2

1.2

1.2

Grouse/successful
hunter

2.2

2.2

2.2

2.2

2.2

No. grouse lost

48

6

54

4

58

Crippling loss/hunter
Total kill

.05

.06
906

157

1,064

.06

.05

.06
110

1,174

Percent crippling loss

5.3

3.8

5.1

3.8

4.9

Percent success of
permittees

44.9

40.0

44.1

40.0

43.7

Banded birds

28

3

31

2

33

Percent nondeliverable

12.0

0.0

12.0

1.01% (12)

�-60-

Eighty-one percent of all permittees went hunting and 53.9 percent of those
actually hunting were successful.
These hunters harvested 1,116 sage grouse
and reported crippling and losing an additional 58 birds.
The total calculated harvest in North Park in 1974 was 1,174. This is substantially less
than the number estimated from the small game hunter survey (2,509) (Table
11). Strong evidence exists that because of sampling problems, the harvest
in North Park, as projected from the Small Game Questionnaire Survey has
been overestimated.
Also, the Small Game Survey appears to over-estimate
the number of hunters by a factor of two. If the relationships evident in
1974 existed in previous years and if they exist statewide it is apparent
that the annual harvest is also over-estimated by 100 percent.
Age and Sex Composition

of Harvest

Prior to 1974 wings collected from hunters at check stations were classified
to sex and as juveniles or adults.
In 1974, wings were separated by sex and
age (juveniles, yearlings and adults).
Depending upon vulnerability, behavior and hunter selection, percentage of
juveniles in the harvest can serve as an index to annual production.
During
the 1963-1974 period, juveniles comprised from 30 to 58 percent of the
havest (Table 10). Apparent production of young was average or better in 9
of the 12 years for which data are available.
These data would suggest that
treatment of sagebrush in the Lake John area had little effect on hatching
success or survival of young to the hunting season.
Production estimates
from brood counts, counts of successful and unsuccessful hens, nest searches,
and climatic factors such as spring temperature and precipitation could not
be related to percentages of juveniles in the harvest.
Hunters had good
success in 1969, but percentage of juveniles in the harvest was low (30
percent).
In 1965, 1971 and 1973, hunters had poor success but percentages
of juveniles in the harvest were relatively high (48 to 51 percent).
Thus
total harvest was apparently not a function of production of that year.
Little is known about vulnerability of juvenile sage grouse.
All available
data (questionnaire survey, observations) indicate that hunters shoot the
first birds for which they have an opportunity.
While it is possible that
juveniles in a group of grouse flush first, or that they are more vulnerable
because of their affinity for edges of meadows, data to support these hypotheses are not available.
Molt data from juvenile sage grouse harvested in 1973 and 1974 indicated
that hatching peaked the third week of June. However, observations of radioequipped hens (based on 12 nests) indicated peak hatching in 1968 and 1969
occurred during the first week of June.
An early June peak of hatch is
supported by scattered observations throughout the period of study and by
timing of peak hen attendance (Patterson 1952) on strutting grounds.
Thus,
the reliability of the aging method (Pyrah 1963) must be questioned.
Data
collected, though limited, suggest the technique of back dating wings from
juveniles in the harvest may underestimate age by 10 to 14 days.
The consistently greater number of adult (yearling and older) hens than
males in the harvest has long caused concern among biologists and game
managers.
It is commonly thought that hunter selection toward smaller birds

�-61-

(females and immatures) was occurring, possibly having an adverse effect
on sage grouse populations.
However, data from the winters of 1973-74 and
1974-75 indicated the sex ratio in winter was 62 females:38males,
and 61
females:39 males, respectively (Beck 1975).
If hunter selectivity was having
a noticeable impact by harvesting smaller birds it should have skewed the
ratios in favor of males.
In 1974 three age classes were distinguished in
the harvest.
Classification of 698 wings by age and sex gave the following
results:
Immatures
Yearlings
Adults

(50.1%)
(19.8%)
(30.1%)

Males
Males
Males

= 171 (48.9%)
49 (35.5%)
45 (21.4%)

Females
179 (51.1%)
Females = 89 (64.5%)
Females = 165 (78.6%)

The sex ratio obtained from pooling these data is 63 females:38 males, similar to the observed winter sex ratio (61:39).
It is obvious from the 1974
harvest data that for each progressive age class, males constitute a smaller
percentage of the kill. More females than males are harvested, but it now
appears this is reflective of the actual sex composition of the population.
Once they are flying, yearling and adult birds cannot be aged and no selection
is possible.
Thus, the lower percentage of males in the adult class than
in the yearling class is indicative of higher mortality rates for males than
females.
The opposite should be true if hunter selection for females was a
reality and was of significant magnitude to affect the population.
Interviews
with hunters support the contention that hunter selectivity is negligible.
Hunters indicated they did not normally get a lot of shots so they shot at
all grouse flushed.
Because of segregation by habitat and sexes and the fact
that immatures and females are similar in size, flocks of birds of like size
are typically observed, making it difficult to select birds by relative size.
It would appear that a predominance of females in the harvest is a reflection
of the actual sex ratio of the population.
The natural sex ratio of a sage
grouse population appears to be skewed in favor of females by natural forces
and not by hunter selectivity.
Band Recoveries
Recoveries of banded birds were small throughout the study.
Direct recoveries
rarely exceeded 10 percent of those banded in a given year, while all recoveries of a given age and sex class did not exceed 15 percent.
Reasons for
the low recovery rate could involve movements of birds out of the area, nonreporting of harvested banded birds, high turnover rate, and annual harvests
of less than 10 percent of the fall population.
However, no bands have been
reported from any location other than Jackson County.
And, although nonreporting does undoubtedly occur, it is not thought to be of sufficient extent
to appreciably alter the low rate of return.
Thus, high turnover and low
annual harvests of fall populations are the likely causes of low band return.
More extensive banding could supply answers to both questions.
Band recoveries by age and sex class are presented in Tables 13 through 16.
Only three indirect recoveries were obtained from 16 birds banded as chicks,
one in year x+4 and two in year x+l. Data from 1974 include the largest

�-62Table 13.

Recoveries of males banded as adults.

Year

No. Banded

X

1963

44

4

4

1964

47

1

1

1965

41

3

1966

6

1967

50

5

1

1968

40

6

4

1969

22

1

1973

89

6

1974

89

8

Totals

428

34

Year of Recovery
X + 1
X + 2
X+ 3

X + 4

2

Total

5
0
6

1

11

1

2

4

10
8

o

2

Year of Recovery
X + 1
X + 2

X + 3

9

2

Table 14.

Recoveries of males banded as yearlings.

Year

No. Banded

1963

11

1964

22

1965

15

1966

10

1967

27

4

1

1968

21

2

1

1969

19

1973

80

7

1974

54

6

Totals

259

21

X

47

Total

0
1
2

1
2

1

2

1
5

1
1

4
3

4

11
6

8

1

3

33

�.\
I

-63Table 15.

Recoveries of females banded as adults.

Year

No. Banded

1963

5

1964

4

1965

9

1

1966

6

1

1

1967

7

1

1

2

1968

11

1

1969

13

4

1973

54

5

1974

29

2

Totals

138

14

Table 16.

Recoveries of females banded as yearlings.

Year

No. Banded

1963

3

0

1964

9

0

1965

3

0

1966

3

1

1967

5

1

1968

6

1

1

1969

20

5

5

1973

41

2

1974

20

2

Totals

110

12

X

X + 1

Year of Recovery
X + 3
X + 2

X + 4

Total

0
1

1

X

1

2

3

2

4
6

1

2

3

3

1

X+

0

Year of Recovery
X + 3
X + 2

1

21

X + 4

Total

2

1

2

1

4

2

2

2

1

1

o

16

�-64-

samples, 30 recoveries (22 males, 7 females, 1 unknown) from an estimated
317 (217 males, 100 females) banded birds in the population.
These limited
data indicate that hunters harvested 10.1 percent of the banded adult and
yearling male segment of the population, and 7 percent of the banded adult
and yearling female segment.
Thus, overall hunting mortality was approximately 9.5 percent of the fall population of banded sage grouse.
Best
present estimates suggest that up to 30 percent of the fall population may
be harvested on a sustained annual basis.
Winter

Activities

Distribution

During 8 January to 8 March 1974, 129 sage grouse flocks and 10 individual
birds were sighted, amounting to 2,753 birds.
Between 1 January and 1 March
1975, 70 flocks and 7 single birds were sighted, involving 2,327 birds.
Plotting of flock locations (Fig. 6) indicates three areas where no flocks
were observed, seven areas of high use, and scattered observations within
remaining areas of sagebrush.
The largest area of no observed use was the area south of Owl and Peterson
Ridges, which amounts to 36.0 percent (172 mi2) of the sagebrush lands in
North Park.
Only 1 of 216 sage grouse sightings was south of this ridge line.
Little sagebrush was available in this area due to higher snowfall and more
extensive snow cover even in the mild winter of 1974-75.
The area between
the Canadian River and Medicine Bow Range had ample available sagebrush
throughout winter but no observed grouse use. This area accounts for 9.1
percent (44 mi2) of the sagebrush lands in North Park.
The third area of
no observed grouse use was the broken ridges south of Independence Mountain
and north of Jackson County Road No.6,
encompassing 5.0 percent (28 mi2)
of the sagebrush lands.
Steep slopes probably limited grouse use of this
area as extensive stands of tall, dense sagebrush on relatively steep slopes
were common.
Thus, 50.1 percent of the sagebrush-dominated
lands in North
Park received no observed use by sage grouse during winter months in 1974
and 1975.
Seven areas of high use were evident and comprised 6.8 percent (32.8 mi2)
of sagebrush lands in North Park. These seven areas were not searched more
frequently than other less productive areas, thus eliminating the bias of
implied heavy use due to high frequency of search time. Searches for sage
grouse were conducted during 60 days each winter.
At least one or more of
the seven areas was searched on 30 days each year.
Areas subsequently
determined to be used lightly or not at all were searched on at least 20
days during each season.
Nearly 75 percent (74.4) of all flocks and 79.2 percent of all grouse
observed in 1974 were within these seven areas whereas only 37.1 percent of
all flocks and 51.2 percent of all grouse observed in 1975 were in these
areas (Table 17). During the mild winter of 1974-75 less sagebrush was

�-65-

,

COl(JHAt,)()

r.·

.,')~

.

..

- ,

1/ r :
--&amp;..:. - • ""- .

,

(.,
J

'IJOf'¥.~Ar..'~")V.·"'lIrr'

~D '.,~
(

•.
~'

•••••
~,,-

-: ...
-~.;.~..;."

,

.

.f

Fig. 6. Location of sage grouse flocks during winters of 1973-74 and
1974-75, North Park, Colorado. Each dot represents one flock. Numbered
areas correspond to high use areas described in Table 17.

�Table 17.
1974-75.

Location and size of areas of high use by wintering sage grouse, North Park, 1973-74 and

Approximate
Area
(mi2)

1974

1975

1974

1975

NE!!.;
S13, E~ S12, RSOW, T10N; NE!!.;
S19, SlS,
SW~ S17, NW~, S~S7, R79W, T10N

3.0

14

0

114

0

sE!!.;
S19, S~ S20, SW~ S27, S~, NW~ S2S, S29, E~
S32, S33, S34, R79W, T10N; N~ S3, N~ S4, NE!!.;
S5, R79W, T9N

6.5

16

1

214

60

SE~ Sl, E~ S12, NE~ Sl3, RS1W, T9N; W~, NE~
S7, NW~ S20, S21, RSOW, T9N

3.0

19

1

223

11

Location
1.

2.

3.
4.
5.

6.
7.

No. of Flocks
Observed

Total Birds
Observed

sE!!.;
S7, S~ SS, N~, SE~ S17, W~ S16, sE!!.;
S19,
S20, S21, S2S, NE~ S29, N~ S33, RSOW, T10N

5.5

S

10

179

44S

E~ S7, SW~ SS, SW~ S16, S17, E~ SlS, E~ S19,
S20, S21, SW~ S22 W~ S27, S2S, S29, N~ S30,
NE~ S33, R7SW, T9N; S13, W~ S24, E~ S23, R79W,
T9N

10.3

19

S

S66

521

S~ S31, S~ S32, R7SW, TSN; W~, NE~ S5, S6,
N~ S7, NW~ SS, R78W, T7N

3.5

13

6

371

152

S2, RS1W, T10N

1.0

7

0

213

0

32.S

96

26

2,lS0

1,192

Totals

I

0"1
0"1
I

�-67completely covered, and consequently, a larger area was available for
foraging.
More available sagebrush also accounts for the reduction in
occupany of the high-use areas delineated in 1973-74 and the increased
difficulty in locating flocks. As the winter of 1973-74 was more severe
than normal, these high-use areas possibly were the last suitable habitat
available.
Only three of the seven 1973-74 high-use areas sustained high use
in 1974-75, thus discounting the possibility of affinity for specific wintering grounds.
Wintering areas in Idaho and Wyoming are also determined by snow
accumulation rather than affinity for specific areas (Dalke et ale 1963,
Patterson 1952).
The high-use area near California Gulch sustained high
use throughout the winter of 1974-75, but only during the last three weeks
of winter in 1973-74.
During this three week period, snow was melting and
cover conditions were similar to those found throughout 1974-75.
The
California Gulch area was the first part of North Park to become snow free
in the springs of 1974 and 1975.
Possibly the birds observed in this area
in both years were associated with the five nearby strutting grounds.
The
other six high-use areas sustained heaviest utilization prior to any melting
and likely were important wintering areas.
Thirty-two percent (155 mi2) of the sagebrush lands in North Park have been
disturbed by 2,4-D spraying, plowing and seeding, and burning since 1957.
Only four flocks· were observed in areas that were still noticeably altered.
Flock Composition

and Size

In winter, sage grouse in North Park were gregarious, the flock being the
prevalent social unit (Table 18). Over 85 percent (85.4) (2,350) of the
2,753 birds observed in 1974 were classified as to sex. Males numbered 889
and females 1,461 for a pooled sex ratio of 62 females:38 males.
Approxi~
mately 85 percent (84.3) (1,984) of the 2,327 sage grouse observed in 1975
were classified as to sex with males numbering 773 and females 1,211.
The
pooled sex ratio for 1975 was 61 females:39 males.

Table 18. Number, frequency of encounter, and percentage of total sage
grouse observed as singles and flocks, North Park, winters of 1973-74
and 1974-75.

Number of encounters
Frequency

of encounter-

1/

Percent of all birds
observed

1/ In percent of total.

Size of Unit
26-50
51-100

1

2-25

17

153

21

16

9

7.9

70.8

9.7

7.4

4.2

0.3

32.5

14.5

22.0

30. 7

&gt;100

�-68Strong segregation by sexes was evident in sage grouse flocks in winter.
Of 176 flocks classified in 1974 and 1975, 69.9 percent (123) were composed
of at least 90 percent birds of like sex. Of the 123 groups, 81.3 percent
(100) were composed of birds of the same sex. Segregation was more pronounced in male than female flocks as 71.6 percent of the predominantly male and
39.5 percent of the predominantly female flocks were unisexual.
Only one
flock (six birds) was composed equally of males and females.
Segregation in sage grouse flocks in winter was as great or greater th~n that
reported for other species of grouse (Koskimies 1957, Seiskari 1962, Weeden
1964, Braun and Schmidt 1971). Reasons for strong sex segregation in sage
grouse are unknown but are of interest.
Review of literature indicates sex
segregation by grouse is accompanied by differential habitat utilization
(Koskimies 1957, Seiskari 1962, Weeden 1964, Braun and Schmidt 1971). Data
from the winter of 1973-74 indicated female flocks utilized denser stands
of sagebrush than males.
However, possibly due to s~ple
size, differences
in vegetal
characteristics between sites used by male and female flocks in
both winters were not significant (P&gt;O.lO)
(Table 20).
Flock size was 50 or less in 88.4 percent of all observations
(Table 19).
Mean size of flocks in 1974 was 12.2 for males, and 29.2 for females, whereas mean sizes in 1975 were 21.2 for males and 48.1 for females.
Differences
between male and female flock size during the two winters were significant
(P &lt; 0.05). Increase in size between years (1974 vs 1975) w~s significant
(P ~ 0.05) for male but not for female flocks.
Median size for male flocks
was 10 in 1974 and 15.5 in 1975, while corresponding values for female flocks
were 15 and 20.5. Evaluation of mean and median flock sizes indicate females
had a greater tendency to form large flocks (&gt;150), although most female
flocks were within the same size range as male flocks « 50). Possibly the
tendency toward larger flock size shown by females was associated with the
use of denser sagebrush stands.
Such an association can be attributed to
either large groupings selecting dense sagebrush for cover and food or the
use of dense stands allowing the congregation of large numbers.
The large increase in flock size for both sexes during 1974-75 can possibly
be attributed to good production the preceding summer and/or the mild winter
of 1974-75 which resulted in more available sagebrush and thus larger flocks.
In the relatively severe winter of 1973-74 flock size was small, probably
because of limited availability of sagebrush.
Areas suitable for use were
small, consequently such sites were suitable only for small groups of grouse.
It would appear that sage grouse control their flock size depending upon
availability of food and cover.
Habitat
Physical Characteristics.--Flocks
of sage grouse were typically located on
south, southwest, or west facing slopes of less than 5 percent (Figs. 7 and 8).
The average percent slope for areas used by female flocks was 4.9 (N=74) and
for male flocks was 6.0 (N = 82). Sixty-six percent of the areas had slopes
less than 5 percent, while only 12.8 percent had slopes greater than 10 percent,

�-69-

Table 19. Sex composition of winter flocks of sage grouse, North Park,
1973-74 and 1974-75. 1-/

Size of Flock

51-59

Percent in Flock
80-89
70-79
60-69

90-99

100

Total

36

42

Males
2-9
10-19

1

2

2

2

1

1

3

4

24

34

2

2

2

5

11

20-29
40-49

1

50-74

1

1
2

1

75-99
Sub-total

3

4

5

7

8

2

6

1

1

68

95

16

24

Females
2-9

5

1

2

10-19

1

1

5

5

11

23

20-29

2

3

1

2

8

30-39

1

1

1

1

5

40-49

1

3

1

5

1

2

50-74
2

75-99
&gt;100

1

2

4

1

3

1

4

2

1

9

Sub-total

1

10

6

17

15

32

81

Total

4

14

11

24

23

100

176

Percent of
All Flocks

2

8

6

14

13

57

1/
Excluding one flock of 50:50 males and females.

�Table 20.

Vegetal

characteristics

of winter

Mean Sagebrush
Density
(Plants/O.Ol
acre)

Sample
Size

use sites,

1974 and 1975.

Mean Height of
Sagebrush Above
Snow (inches)

Mean Average
Crown Bread th of
Sagebrush (inches)

F

M

F

M

F

M

F

641];)

Mean Cover
(Density x Height)
(in.-plant/0.01
acre)

Year

M

F ])

M

1974

58

42

46.0

68.22:./

7.8

9.4

14.8

17.3

359

1975

22

29

63.7].1

77 .5

11.61/

14.0 1/

16.1

17.5

7391/

flock

of &gt; 50 percent

1,0851/

])
M denotes

males,

F denotes

flock

of&gt; 50 percent

J.j

females.
I

Significant
].1
Significant

difference

between sexes

during

a year,

P ~0.05.

o
"
I

difference

between years

within

a sex,

P &lt; 0.05.

�-71-

N

E

FLOCK SIZE
• 2-9
o 10-19
o 20-29
A
~30
Fig. 7.
Aspect and percent slope of areas utilized by flocks of male
sage grouse in winter.
Each concentric circle represents 5 percent slope
with the center representing zero slope.
Symbols representing different
flock sizes are given in the legend.

�-72-

N

0

•
W

t 4 c9 •

~:

o .o~.
{:)

S

E

~

FLOCK SIZE

2-9
• 10-19
0

c
.6-

20-29
&gt;30

Fig. 8. Aspect and percent slope of areas utilized by flocks of female
sage grouse in winter.
Each concentric circle represents 5 percent slope
with the center representing zero slope.
Symbols representing different
flock sizes are given in the legend.

�-73-

On areas with slope greater than 5 percent, 66.7 percent of female and 93.1
percent of male flocks were observed on west, southwest, south or southeast
facing slopes.
Wind was present during 73.7 percent of 152 observations
for which data were collected.
Wind direction was from the southwest 60.7
percent of the time, west 17.0 percent, and south 12.5 percent.
No data on
wind velocity were collected.
Although snow cover was a major determinant in amount of available sagebrush,
slope and aspect appeared to further restrict the amount of sagebrush range
suitable for wintering sage grouse.
No data were available on relative abundance of varying slopes although numerous steep slopes dominated by sagebrush
exist.
No observed use occurred on steep slopes (&gt;30 percent) and most
observed use was on areas with less than 10 percent slope.
Steepness of
slope modified by aspect thus eliminated many sagebrush areas from suitable
habitat status.
Vegetal Characteristics.--Vegetal
measurements were taken at 100 use sites
in 1974 and 51 sites in 1975. Mean sagebrush density, mean height of sagebrush above snow, mean crown breadth, and mean cover (ht x density) were
used in comparing sites utilized by male and female flocks.
Significant
differences (P&lt;0.05) were documented in sagebrush density and cover in 1974,
when females used denser sagebrush stands than males.
Significant differences (P ~ 0.05) between years occurred for sagebrush density on sites used
by male flocks, height of sagebrush above snow and cover for sites used by
both male and female flocks (Table 20). Although the difference in mean
values for male and female flocks was less in 1974-75, variability in data
was also less. Density and cover values were much higher in 1974-75,
especially for male flocks.
This was probably a reflection of the low snowfall the second winter.
Selection for differing densities of sagebrush may
also have been tempered by the mild winter in 1974-75.
Density and cover
values did not increase as much for female as for male flocks in 1974-75,
suggesting that female flocks used sagebrush stands approaching the maximum
density available in North Park.
An unfortunate weakness in the data was
that measurements of sagebrush stands were not taken in areas of no observed
use. Thus, although female flocks used denser stands than males, there are
no data to indicate where densities used align on the density continuum of
available sagebrush.
Vegetal characteristics of each high-use area were compared to those of the
combined values for all locations not within a high-use area.
Only one highuse area had vegetative characters significantly different (P &lt; 0.05) from
other high-use areas and the scattered-use locations.
This area was located
in the McCallum oil field where earlier researchers also observed wintering
flocks of sage grouse.
Sagebrush in this area was less dense than in other
areas of flock sightings.

�t

Twenty*six of the 199 flock locatlons were used as roosting sites, the
remainder being feeding-loafing sites. Vegetative differences between the
two groupings were not significant (P&gt; 0.05). Roosting depressions in the
Snow at eight sites was crusted and
snovr occurred at 13 of the 26 sites.
at five sites was powder. These values were similar to the frequency of
crusted and por,rder condltions recorded at all sltes (64.9i1 crusted, 29.8''l
powder, 5.32 melting). Depressions to depths of. L2 in. were observed.
Seasonal Movements

l:{ost movement data were available for birds banded in the Lake John area of
Nort}r Fark. Movements deternr-ined from band recoveries represent the spring
ro fa11 period (Tab1e 2i). Since no significant (P&gt; 0.05) differences l,lere
found between movements resulting from direct and indirect recoveries, all
data rnere pooled. Significant (P 1 0.05) differences were found, only
betr^:een movements of adult males and yearling females as yearling females
noved greater distances. However, in general, females of both age
classes mor,,ed greater distances than males" It is not knornrn if these
differences relate to sagebrush treatmenE. Nineteen of the 100 recoveries
we::e from outside of the or:iginal T,ake John srudy area.
l4ovements could not be attributed sole1y to attraciion to wet sites as such
areas occurred in abundance r^rithin four miles of any banding site. Only 22
cf the 100 recoveries r,rere rvithin two miles of the banding site. Movements

from banding sites, usually strutting grounds' were significantly (P&lt; 0'05)
grearer tharL two mi1es. Thus, attention should be focused on seasonal
requirements of sage grouse instead of maintaining protected zones within a
two mile radius of strutti-ng grounds. Based on data cotrlected in this study
(Table 21) and assuming that all. sagebrush disturbance is harmful to sage
grouse, a six mile radius of undisturbed sagebrrrsh around strutting grounds
rrrould b+: most sriitable.
Movements of birds from spring banding sites to stmmer use areas were

considerably shorter than spring to fa11 movements. Dlstances traveled by
males (N = 18) were no greater than those of females (N = 4)r 3.7 to 3-4
miles, respectively. Sma1l samples and the fact that only 3 of the 22
sightings were after 30 July pose restrictions on inferred conclusions.
However, the shorter mid-sununer movements probabLy indicate that considerable
movement occurs in late summer just prior to the hunting season. Whether
this is associated with juvenile dispersal or changes in food supply is not
known.

Nine females in 1968 and 24 femaLes in 1969 were fitted with VHF transmitters
after being trapped on strutting grounds. In 1968 four hens were followed
to nests, while 8 of the 24 hens in 1969 were followed to nest sites. These
12 females traveled a mean distance of 3.12 mil-es from the strutting ground
where captured (range of 0.4-8.2 mi). No pattern ln direction of movement
was evident. Early nesters moved greater distances to nest sites than late
nesters though differences were not statistically significant (P&gt;0.05).
Relatively leve1 areas (&lt; l0% slope) were utilLzed by the 12 telemetered
females and no preference for aspect was noted.

,€

�-7 5-

Table 21. Spring to fa11 movements of sage grouse calculated from band
recoveries.
Dis tance

(ltiles )

0-

Adult Males

Yearling
Males

Adult Females

Yearling
Females

1.9

L2

4

3

3

2,0

3.9

9

t2

6

0

4.0

5.9

11

4

J

3

6.0

7.9

2

1

4

B.0

9.9

5

3

0

I
I

10.0

11 .9

2

1

0

1

L2.0

13.9

I

1

0

0

14.0

15. 9

0

I

1

0

16.0

L7.9

1

0

1

1

18.0 - 19.9

0

1

0

0

20.0 - 21.9

0

0

1

0

l,tean (i-)

4 9

5 3

6 0

6.25

Sample size

4J

2B

19

10

Movements of hens equipped with WIF radio transmiEters, based on sma11 sample
sizes, indicaEe the general trend of brood movement. Movements from nesting
sites to wet meadorus were highly variabl-e. Movements to meadows began on
days l, 4 and 30 after hatching for the three hens monitored. In addition,
three hens with young were located after reaching meadows. Data from these
six hens indicate that movements after reaching meadows were minimal, This
supports the hypothesis that frequent brood counts in a limited area prowide

data on only a few frequently reobserved broods.

Movements of sage grouse from spring to winter were calculated from 57
observations of banded birds ln the winter of L973-74 and 22 in 7974-75
(Tab7e 22). Average distance traveled from banding sites for males was 6.5
mi. (N = 50) in L973-V4, and 3.2 ml.. (N = 17) in 1974-75" Corresponding
values for females were 11.5 mi. (N = 5) and 9.5 mi. (N = 5).

�-76-

Table 22. Minimum distances traveled from spring to winter by male and
female sage grouse banded near: Lake John, Norrh park, t9l3-7h and t9l4-75.

Year

0-3

3. r-6

197 3-7 4

13

797 4-7 5

10

Distance Traveled (ltiles)
6. 1-9

9.L-tz

12.1-18

&gt;18

10

20

9

4

1

6

J

I

2

0

Sirorter distances traveled in 1974-75 probably resulted from the milder
tvinter when more sagebrush was available throughout the north half of the
park. Longer distances for females may have resulted from their apparent
preference for dense stands of sagebrush which are not evenly distributed
throughout the Park. A11 large concentrations of females were located in
the northeast quarter of the park rvhere most dense stands of sagebrush were
avai-lab1e. This is approximately 10 mlles fronn the center of the Lake John
study area.
DisEances traveled indicated saEIe grouse were moving freely throughout North
Park. Thus, all sage grouse in the park should be considered as one population rather than several localized, sedentary subpopulations. The long movements and concentrations of birds in the northeast quarter of North park
during winter indicate definite seasonal use patterns for different areas
wlthin North Park. Consequently, the park must be consj-dered as a single
management unlt.
Conclusions

Effects of sagebrush spraying upon distribution and abundance of sage grouse
in North Park were difficult to assess. L,Ieile problems arose wiEh size of
area studied and frequent changes in project personnel, a major problem also
occurred as more sagebrush within the original study area was treated Lhan
planned. Prior to the experimental spraying for study purposes, 3,610 acres
^
(5,0+ rr:i4
of sagebrush in the study area had been plowed and seeded to
grasses. Area sprayed during the study involved about 41000 acres (6.25 mi2).
Within 5 years of the origlnal spraying, an additional 1,250 acres (1.9S mi2)
ln the study area were sprayed wi-th 2,4-D. Thus, immediately prior to and
during the course of the study a minimum of 8,860 acres (13.84 mi2) or 27,1
percent of, tire sagebrush lands in the Lake John study area had been treated
by plowing or spraying with 2,4-D. These activitles could have easily
masked any effects of the 4,000 acres treated for ti-le study.
Block spraying r:f an area slrrround.ing a Large strutting ground resulted in
al.most compiete a}:andonment of the ground within 7 years of spraying. Block
spraying with nearly 100 percent sagebr-ush ki11 had an adverse effect on
r:se of the area for nesting, primarily due to removal of cover. Spraying of

?

�-77-

sagebrush reduced availability of forbs and sagebrush and reduced sage
grouse use of sprayed areas, especially those areas where sagebrush kill
was nearly complete.
Distribution and use of strutting grounds adjacent to
treated areas changed, probably as a result of spraying.
While problems
exist with interpretation of counts of grouse on strutting grounds, numbers
of male sage grouse on strutting grounds in the original study area decreased
significantly (P &lt; 0.05) from 1963 to 1974. This decrease can only be related
to sagebrush treatment.
Effects of sagebrush spraying on movements are
unclear.
Movements of adult and yearling grouse were greater than expected,
possibly indicating lack of suitable habitats within the original study area.
These larger than expected movements could be the result of sagebrush alteration. Nesting success and production did not appear to be affected by
sagebrush treatment except through reducing grouse use of large areas.
Reliable harvest data for North Park were not available until the last year
of the study. Hunter success declined throughout the study, possibly as a
result of fewer grouse being available.
Treatment of sagebrush by spraying
undoubtedly reduced availability of sagebrush in winter, further reducing
the capacity of the area to sustain numbers of sage grouse.
Strip spraying
as described in the report had almost no measurable effect on sage grouse.

LITERATURE

Autenrieth,
Rept.

CITED

R. E. 1969. Sage grouse investigations.
Idaho Fish and Game Dept. 25 p. Mimeo.

1970. Sage grouse investigations.
Fish and Game Dept.
21 p. Mimeo.

Third Annual Prog.

Fourth Annual Prog. Rept.

Idaho

Beck, T.D.I.
1975. Attributes of a wintering population of sage grouse,
North Park, Colorado.
M. S. Thesis.
Colorado State Univ., Ft.
Collins.
49 pp.
_____ , R. B. Gill, and C. E. Braun.
1975.
sage grouse from wing characteristics.
(revised).
Colorado Div. of Wildlife.

Sex and age determination of
Game Inf. Leaflet No. 49
4 pp.

Beekly, A. L. 1915. Geology and coal resources of North Park, Colorado.
U. S. Geol. Surv. Bull. 596. 121 pp.
The section Tridentate
Beetle, A. A. 1960. A study of sagebrush.
Wyoming Agr. Exp. Sta. Bull. 368. 83 pp.
Artemisia.
Brander, R. B. 1968. A radio-package
Manage. 32(3):630-632.

harness

for game birds.

of

J. Wildl.

Braun, C. E., and R. K. Schmidt, Jr. 1971. Effects of snow and wind on
wintering populations of white-tailed ptarmigan in Colorado
pp. 238250. In A. O. Haugen (ed). Proc. Snow and Ice Symposium, Ames, Iowa.

280 pp ,

�-78-

Carr, H. D. 1967.
Sage grouse and sagebrush control.
Colorado State Univ., Ft. Collins.
106 pp.

M. S. Thesis,

Craighead, J. J., and D. S. Stockstad.
marking birds.
J. Wildl. Manage.

neckband

1956.
A colored
20(3):331-332.

Crunden, C. W.
1963.
Age and sex of sage grouse
Manage. 27(4):846-849.

for

J. Wildl.

from wings.

Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F.
Schlatterer.
1963.
Ecology, productivity
and management of sage
grouse in Idaho.
J. Wildl. Manage. 27(4):811-841.
Daub enmire, R.
Sta. Tech.

1970.
Bull.

Steppe vegetation
62. 131 pp.

Eng, R. L. 1954.
Use of aerial
counts.
Proc. West. Assoc.

of Washington.

Agr.

Exp.

coverage in sage grouse strutting ground
State Game and Fish Comms. 34:231-233.

1955.
A method for obtaining sage grouse
wings.
J. Wildl. Manage. 19(2):267-272.
1963.

Washington

Observations
on the breeding
27(4):841-846.

age and sex ratios

biology

of male

from

sage grouse.

J. Wildl. Manage.

_____ , and P. Schladweiler.
habitat USe in central

1972.
Sage grouse winter movements and
Montana.
J. Wildl. Manage. 36 (1) :141-146.

Finch, W. C., ed. 1957.
Guidebook to the geology of North and Middle
Parks Basin, Colorado.
Rocky Mtn. Assoc. Geol., Denver.
Gill, R. B. 1965.
Distribution
and abundance of a population of sage grouse
in North Park, Colorado.
M. S. Thesis.
Colorado State Univ., Ft.
Collins.
187 pp.
Griner, L. A. 1939.
A study of the sage grouse (Centrocercus urophasianus),
with special reference to life history, habitat requirements,
and
numbers and distribution.
M. S. Thesis.
Utah State Univ., Logan.
83 pp.
Hail, W. J. 1965.
Geology of northwest
Surv. Bull. 1188.
133 pp.

North Park,

Colorado.

U. S. Geol.

Hartzler, J. E. 1972.
An ana~ysis of sage grouse lek behavior.
Dissertation.
Univ. Montana, Missoula.
200 pp.

Ph. D.

Hyder, D. N., C. G. Conrad, P. T. Tueller, L. D. Calvin, Co E. Poulton,
and F. A. Sneva.
1963.
Frequency sampling in sagebrush-bunchgrass
vegetation.
Ecol. 44(4):740-746.
Klebenow, D. A. 1969.
J. Wildl. Manage.

Sage grouse nesting
33(3):649-661.

and brood habitat

in Idaho.

�-79-

_____ , and G. M. Gray.
1968. Food habits
Range Manage. 21(2):80-83.

of juvenile

sage grouse.

J.

Koskimies, J. 1957. Flocking behavior in capercaillie, Tetrao urogallus
(L.), and blackgame, Lyrurus tetrix (L.). Finnish Papers Game Res. 18.
32 pp.
Labinski, R. F., and S. H. Mann.
1962.
J. ur rai . Manage. 26(4) :393-399.

Backtag markers

Lacher, J. R., and D. D. Lacher.
Manage. 28(3):595-597.

A mobile

1964.

for pheasants.

cannon net trap.

Lumsden, H. G. 1968. The displays of the sage grouse.
and Forests, Res. Rept. 83. 94 pp.

Ontario

J. Wildl.

Dept. Lands

Marshall, W. H. 1963. Studies of movements, behavior and activities of
ruffed grouse using radio' telemetry techniques.
Univ. Minn., Minneapolis.
Prog. Rept. 30 pp.
(Multilithed).
Martin, N. S. 1970. Sagebrush control related to habitat
occurrence.
J. Wildl. Manage. 34(2):313-320.
Patterson, R. L. 1952.
Denver.
341 pp.

The sage grouse in Wyoming.

and sage grouse

Sage Books, Inc.,

Peterson, J. G. 1970. The food habits and summer distribution of juvenile
sage grouse in central Montana.
J. Wildl. Manage. 34(1) :147-155.
Pyrah, D. G. 1963.
Sage grouse investigations.
Idaho Fish and Game
Dept., Wildlife Restoration Div., Job Compl. Rept., Fed. Aid Proj.
W-125-R.
71 pp.
___

1970.
466-467.

Poncho markers

for game birds.

J. Wildl. Manage.

Rippin, A. B., and D. A. Boag.
1974. Recruitment to populations
sharp-tailed grouse.
J. Wildl. Manage. 38(4):616-621.

34(2):

of male

Robel, R. J. 1969. Movements and flock stratification within a population
of blackcocks in Scotland.
J. Anim. Ecol. 38(5) :755-763.
Rogers, G. E. 1964. Sage grouse invetigations in Colorado.
Fish and Parks Dept. Tech. Bull. 16. 132 pp.

Colorado

Game,

Seiskari, P. 1962. On the winter ecology of the capercail1ie, Tetrao
urogallus, and the black grouse, Lyrurus tetrix, in Finland.
Finnish
Papers Game Res. 22. 119 pp.
Smith, E. L., Jr. 1966.
Soil-vegetation relationships of some Ar t emf.s La
types in North Park, Colorado.
Ph. D. Dissertation.
Colorado State
Univ., Ft. Collins.
203 pp.

�-80Stanton, D. C. 1958.
grouse population
Moscow.
87 pp.

A study of the breeding and reproduction in a sage
in southeastern Idaho. M. S. Thesis.
Univ. Idaho,

U. S. Department of Commerce.
1973. Monthly normals of temperature,
precipitation, and heating and cooling degree days, 1941-70.
Climatography of the U. S. No. 81 (Colorado).
12 pp.
Weeden, R. B.
ptarmigan

1964. Spatial separation of sexes in rock and willow
in winter.
Auk 81(4):534-541.

Wiley, R. H. 1973. Territoriality and non-random mating in sage grouse
Centrocercus urophasianus.
Anim. Behav. Mono. 6(2):87-169.

Prepared by:

L_:!_~_/-'--~-_~-&gt;_.~2~~4T~L~~_~_t_~_)
Clait E. Braun
Wildlife Researcher

Thomas D. 1. Beck
C.B
Graduate Research Assistant

_

�...
81-

APPENDIX

�-82APPENDIX A
Common species in sagebrush uplands of North Park, Colorado.-II
Scientific Name

Common Name
FORBS

Antennaria microphylla
Arenaria fendleri
Artemisia frigida
Aster leucanthemifolius
Astragalus purshii
1':... spatulatus
Chrysopsis villosa
Erigeron spp ,
Eriogonum umbellatum
Eurotia lanata
Mertensia bakeri
Penstemon spp.
Phlox bryoides
Sedum stenopetalum
Senecio amplectens
Trifolium sp.
Vicia americana

Pussy toes
Sandwort
Fringed sage
Aster
Vetch
Vetch
Hairy golden aster
Daisy
Sulphur flower
Winterfat
Bluebells
Beard tongue
Phlox
Stonecrop
Senecio
Goats beard
American vetch
GRASSES

Agropyron dasystachyum
A. riparium
A. smithii
1':... spicatum
Bouteloua gracilis
Bromus anomalus
B. tectorum
Calamagrostis montanensis
Distichlis stricta
Festuca idahoensis
F. ovina
Hordeum jubatum
Koeleria cristata
Poa canbyi
P. fendleriana
P. nevadensis
P. secunda
Sit anion hystrix
Stipa comata
Stipa lettermani

!/Adapted from Smith (1966) and Gill (1965).

Thickspike wheat grass
Streambank wheatgrass
Western wheatgrass
Bluebunch wheatgrass
Blue grama
Nodding brome
Cheatgrass
Plains reedgrass
Inland saltgrass
Idaho fescue
Sheep fescue
Foxtail barley
Junegrass
Canby bluegrass
Mutton bluegrass
Nevada bluegrass
Sandberg bluegrass
Squirrel tail
Needle and thread grass
Needlegrass

�-83APPENDIX B

Legal description of active strutting grounds, North Park, 1975.

Section

Quarter
Section

Range

Township

Alkali Lake

31

NW

80

10

Boettcher Lake Junction

13

SW

81

10

Canuck

16

NW

79

10

Coalmont

24

SW

81

7

Cowdrey 115

9

NE

80

10

Deer Creek

31

NE

78

7

Delaney Butte (Recently
inactive)

35

SE

81

9

Fish Hatchery

9

SW

80

8

Hound

13

NW

80

9

Lost Creek ill

9

SE

79

6

Lost Creek ffi2 (Recently
inactive)

36

NE

80

7

Monahan Draw

29

NW

80

10

Railroad 1/

2

SW

80

7

Ridge Road

22

SW

81

8

Riley

28

NW

80

10

Roth

33

SW

78

9

Spring Creek III

35

SE

79

8

Spring Creek 112

1

SW

79

7

Spring Creek #4

3

NE

79

7

Walden

23

SE

80

9

Wattenberg if2

8

NW

80

9

Strutting Ground

_V Located in 1975, all others located during study period 1963-1974.

�-84APPENDIX C

List of Publications and Theses
Beck, T.D.I. 1975. Attributes of a wintering population of sage grouse,
North Park, Colorado. M. S. Thesis. Colorado State Univ., Ft. Collins.
49 pp.
1975.
7(6):43.

Winter ecology of sage grouse.

_____ , R. B. Gill, and C. E. Braun. 1975.
sage grouse from wing characteristics.
Colorado Div. of Wildl. 4 pp.

Froc. Colo.-Wyo. Acad. Sci.

Sex and age determination of
Game Inf. Leaflet 49 (Revised).

Carr, H. D. 1966. Preliminary findings of effects of sagebrush control on
sage grouse. Colorado Coop. Wild1. Res. Unit Tech. Paper 5. 11 pp.
1967. Sage grouse and sagebrush control.
State Univ., Ft. Collins. 106 pp.

M. S. Thesis. Colorado

1968. A literature review on effects of herbicides on sage grouse
(Centrocercus urophasianus). Colorado Dept. Game, Fish and Parks Spec.
Rept. 13. 16 pp.
, and F. A. Glover. 1970. Effects of sagebrush control on sage grouse.
-----Trans.
No. Am. Wi1d1. and Nat. Res. Conf. 35:205-215.
Gill, R. B. 1965. Distribution and abundance of a population of sage grouse
in North Park, Colorado. M. S. Thesis. Colorado State Gniv., Ft.
Collins. 187 pp.
1966. A literature review on the sage grouse.
Game, Fish and Parks Spec. Rept. 6. 39 pp.

Colorado Dept.

, and F. A. Glover. 1965. Daily and seasonal movements of sage grouse.
~--Colorado Coop. Wi1dl. Res. Unit Tech. Paper 3. 6 pp.
May, T. A., and B. E. Poley. 1969. Spring and summer movements of female
sage grouse in North Park, Colorado. Proc. Biennial West. States Sage
Grouse Workshop 6:173-179.
Poley, B. E. 1969. Seasonal movements of sage grouse in Colorado.
Field Ornith. 5:1-4.

Colo.

�April 1976

-85-

State of
Project No.
Hork Plan No.
Job Title
Period Covered:

COLORADO
Game Bird Survey

ItJ-37-R-29
3

Job No.
9
Part I
Evaluation of the Effects of
Changes in Hunting Regulations on Sage Grouse Populations
April 1, 1975 through March 31, 1976

Personnel:
Fred Giese, U.S. Fish and Wildlife Service, Ron Ryder, Colorado
State Dniversity; Heather Flanagan Alexander, Tom Beck, Don Benson, Clait
Braun, Tom Campbell, Courtney Crawford, Daryl Crawford, Howard Funk, Ken
Giesen, Don Gore, John Hobbs, Richard Hoffman, Jim Jackson, J. Edward Kautz,
Fran Marcoux, Sue McElderry., Ken Miller, Sig Palm, Brett Petersen, Steve
Porter, Sue Quinlan, Wayne Russell, Howard Spear, and John \.;ragner,
Colorado
Division of Wildlife.
ABSTRACT
Investigations concerning the relationships of hunting to breeding population
levels, and mortality and survival rates of sage grouse (Centro cercus
urophasianus) in North Park, Colorado were initiated in 1975. Numbers of
strutting grounds remained stable, although two grounds active in 1974 were
inactive in 1975. One ground inactive in 1974 was active in 1975, while one
new ground was located.
Average number of cocks per ground increased from
28 in 1974 to 31 in 1975. Peak period of female attend~~ce in 1975 was
between April 15-23.
Total estimated spring population size was about 4,4004,500 birds.
In 1975, 422 (291 males, 130 females, 1 unknown sex) sage grouse
were banded.
A 9 day permit only hunting season was allowed in North Park
in 1975 with bag and possession limits of 2 and 4. Data from wings (N = 505)
collected at check stations and field checks revealed that the harvest was
comprised of 42 percent young of the year, 22 percent yearlings and 36 percent adults.
Total permits issued numbered 1,541, of which· only 77 percent
of the permittees went hunting.
Estimated total harvest including crippling
loss was 1,053 grouse.
The increase in total harvest resulting from the 6
day longer season in 1975 was 158 birds.
Sixty-two bands were recovered and
reported
by hunters, of which 40 were from 1975 bandings ,'&gt;lith
11 each from
1974 and 1973. Approximately 10 percent of the adult and yearling birds
banded in 1975 were harvested.
Harvest of banded birds was not uniform by
area of banding with two strutting grounds having harvests of more than 20
percent of the cocks counted in 1975. Estimated fall population size was
about 10,000 birds, a value considered to be higher than the actual population due to possible differential vulnerability of chicks.

�-86RECOMMENDATIONS

1.

Trapping efforts should be intensified with more effort being made to
capture females in the spring and females and chicks in July and August.

2.

Changes in hunting regulations should be promulgated
and bag and possession limits of 3 and 6 in 1976.

with a 9 day season

�-87-

EVALUATION OF THE EFFECTS OF CHANGES
IN HUNTING REGULATIONS ON SAGE GROUSE POPULATIONS
Clait E. Braun

Little is known concerning effects of hunting, if any, on populations of
sage grouse in Colorado or anywhere within their principal range.
It is
connnonly believed by field personnel and managers in Colorado and elsewhere
that sage grouse are easily hunted and harvested, and thus are exceedingly
vulnerable to hunting.
Evidence supporting these ideas is not available.
Most state management agencies base sage grouse hunting seasons lengths
and bag limits on changes in numbers of males counted on strutting grounds
and on ~hanges in brood size and number.
Unfortunately,
data documenting
relationships between hunting and counts of breeding males made the following spring are lacking.
Basically, problems exist in relating strutting
ground counts to size of populations and to hunting pressure and harvest
during the preceding hunting season.
Most work relating to sage grouse numbers has been concerned with counts of
males on strutting grounds as cocks are highly visible on these breeding
areas in April and early May.
Despite previous efforts, it is not know
whether all males in a population appear on strutting grounds.
Patterson
(1952) suggested that all males visited these communal breeding areas, but
presented no data to support this hypothesis.
Even less is known about
females, although they may outnumber males in the population by 2 to 1
(Beck 1975, Braun and Beck 1976).
Since the social structure of the sage
grouse breeding system allows only a few males to breed, many males are
excess.
Thus, emphasis should be placed on understanding the dynamics of
the female segment of the population and the relationship between harvest
and population size.
P.N.O. OBJECTIVE
The objectives of this study are to increase hunter
tional use of sage grouse in North Park, Colorado.
been developed are:

opportunity and recreaHypotheses which have

a.

Hunter harvest does not affect spring counts of strutting
peak spring counts of females under any regulations.

b.

Counts of females during the peak of attendance on strutting grounds
a reliable index to overall numbers of sage grouse in spring.

c.

Hunter

d.

Nesting success and brood size to 15 August, while important in determining age composition of the harvest, have no relationship to hunter
success and total harvest.

success

and total harvest

are a function

males or

are

of August precipitation.

�-88-

SEGMENT OBJECTIVES
La, Males will be trapped and marked with color-coded bands at night
while roosting on or near strutting grounds.
A sample of 200 will be
marked.
Trapping will be conducted from mid-March to late May by spotlighting and netting.
Bandings will be evenly distributed in four areas
of North Park.
lb. Females will be trapped during the early A.M. on strutting grounds
during April through use of drop, mist and cannon nets. A sample of 300
will be marked with color-coded leg bands and colored numbered patagium
tags. Bandings will be evenly distributed in four areas of North Park.
2a. Hunting will be by free permit only and all hunters will be mailed
a questionnaire within one week of the end of the hunting season.
One
followup letter will be sent two weeks later. A goal of 90 percent response
is desired.
2b. Check stations will be operated at three locations during the
first two weekends of the hunting season.
These stations will be at
Gould, State Line and Willow Creek Pass.
A goal of 500 wings is desired.
2c. Number and location of marked birds shot will be obtained through
use of check stations, field hunter checks and voluntary mail reporting.
A
goal of 50 first year recoveries is desired.
2d.
stations

Hunter success will be determined through
and from the mail questionnaire survey.

data collected

2e. Age composition of the harvest will be determined
tion of wings collected at check stations.
Classification
yearling, juvenile, and male and female.
2f. Total harvest will be estimated (90 percent
the mail questionnaire survey of all hunters.
3.

Promote

4.

Compile data, analyze
METHODS

results

through examinawill be as adult,

confidence

a 9 day season with a bag and possession

at check

level) from

limit of 2 and 4.

and prepare progress

report.

AND MATERIALS

Counts of male and female sage grouse present on strutting grounds were
periodically made by project personnel and Wildlife Conservation Officers
following proscribed procedures (Braun and Beck 1976).
Counts were made
from 19 April through 17 May at times within one-half hour of sunrise.
Searches for new or relocated grounds were made through ground and aerial
reconnaissance.
Male and female sage grouse were trapped at night along
roads and on strutting grounds where they roosted.
Some birds, primarily

�-89-

hens, were captured at dawn on strutting grounds through use of cannon
nets.
Most birds were captured through use of a spotlight with a backpack power source and long handled nets (Braun and Beck 1976). Sage
grouse banded and released were marked with serially numbered aluminum
bands (size 16 for males, 14 for females) and colored plastic and aluminum
bands coded to capture locations.
Age and sex classification of birds
captured followed Eng (1955).
Sage grouse hunters in North Park in 1975 were required to have in their
possession a free numbered permit.
Permits unlimited in number were available from Division of Wildlife offices in Fort Collins, Denver, and Colorado Springs, all license agents in North Park, and all project and management personnel assigned to work in North Park during the hunting season.
Questionnaires were sent to all permittees immediately following the sage
grouse season in North Park, with one followup letter being mailed in midOctober to all non-respondents.
Check stations were operated at three locations (State Line, Gould, Willow
Creek Pass) on the opening weekend (13 and 14 September) and the second
Sunday (21 September) of the season.
Each station was operated from about
1000 to 1800 MDT, depending upon traffic load. Data obtained per party
were:
county of origin, number of hunters, hours hunted (total of all
hunters), birds observed, birds bagged, birds lost, number of banded birds
and location of where each was obtained, and area hunted.
One wing was
obtained from each bird checked with ovaries being taken from a sample of
hens.
Efforts were made to ascertain sex by gonadal inspection for a
sample of birds.
These data were recorded on tags with wings being individually marked with corresponding information concerning actual sex of that
bird. Wings were frozen and stored for later analysis.
Collected wings were thawed and classified to age (chicks, yearlings, adults)
and sex following procedures outlined by Eng (1955), and Beck et al. (1975).
Hatching dates ""ere calculated for chicks of the year using data from
Pyrah (1963).

Description

~J Are~

North Park lies completely in Jackson County, Colorado.
This relatively
high altitude (7,800 to 8,500 ft) area is essentially a closed basin, as
it is encircled by mountain ranges.
Drainage is to the north, with lowest
elevations occurring north of Cowdrey.
Vegetation is principally a sagebrush (Artemisia spp.)-grassland
type with stream courses being dominated
by native grasses, sedges and deciduous shrubs.
Detailed geographic,
geologic, vegetational and climatic features of the area have been described
by Gill (1965), Carr (1967), Beck (1975). Braun and Beck (1976) and will be
treated in the final report.
RESULTS AND DISCUSSION
Strutting

Ground Counts

Counts of male and female sage grouse present on strutting grounds were
initiated on 19 March (Alkali Lake) and continued until 20 May (Fish

�-90-

Hatchery) and 24 May (Railroad).
During this interval 129 counts were
made of 19 active grounds.
Maximum male and female counts per ground are
presented in Table 1. The total of 588 cocks counted was somewhat higher
than the 527 counted in 1974. Average number of cocks per active ground
was 31 in 1975, versus 28 in 1974. Two grounds, Delaney Butte and Lost
Creek #2, which were active in 1974 were inactive in 1975, while one
ground, Lost Creek #1, inactive in 1974 was active in 1975. In addition,
one new ground was located in 1975. This ground, Railroad, was located
on 14 May, late in the strutting period.
The importance of the total count of 573 females in 1975 is unknown, as
comparable data are not available for earlier years.
It should also be
observed that the count of hens is low, as several grounds, Deer Creek,
Lost Creek #1, Railroad, Spring Creek #2, were not counted until late April
or early May after the peak period of female attendance.
The peak period
of hen attendance in 1975 was diffuse, but most grounds experienced highest
hen counts during the 15-23 April interval.
Estimation

of Spring Population

Size

Reliable estimates of total numbers of sage grouse in North Park during
the breeding period are difficult to derive.
Major problems involved are
those relating to how many males and females of the total population are
present on strutting grounds when peak counts are attained and how many
of the actual number of strutting grounds in North Park are known.
Studies
of black grouse (Lyrurus tetrix) (Robel 1969) and sharp-tailed grouse
(Pedioecetes phasianellus)
(Rippin and Boag 1974), two other species of
lek grouse, suggest that no more than 50 percent of the male segment of the
population is present on a lek at a given time.
If it is assumed that peak.
counts of male sage grouse represent 50 percent of the male population in
an area, then at least 1,176 (588 x 2) cocks occur in North Park.
It is
quite obvious that not all strutting ground locations are known, as new
grounds have been found every year during which searches have been made.
If it is assumed that 75 percent of the active grounds have been located,
then a minimum of 5 (.25 x 19) unknown grounds should occur somewhere in
North Park.
Assuming that these 5 grounds each have an average of 31 cocks
during the peak period of male attendance, they represent a minimum of 310
additional cocks (31 x 5 = 155 x 2 = 310). Thus, the total estimated
number of cocks in North Park in the spring of 1975 was 1,486.
Beck (1975) found a late winter sex ratio of 69 hens to 31 cocks, a value
remarkably similar to the sex ratio of adult and yearling birds in the fall
harvest (Braun and Beck 1976). If it is assumed that the spring sex ratio
of the North Park sage grouse popUlation is similar to that observed in
late winter and early fall, then 2 hens occur for every cock.
Thus, the
total spring hen population can be estimated, based on the estimated number
of cocks in the spring population.
Therefore, there should have been a
minimum of 2,972 hens in the spring population (1,486 x 2 = 2,972). The
estimated total spring population of sage grouse in North Park in 1975 was
4,458 birds.

�-91Table

1.

Sage grouse

ground counts,

Maximum
No. of
Males

Ground

Alkali

strutting

Lake

Date

68

April

50

Canuck

1975.

Maximum
No. of
Females

Date

64

April

16 and 21

May 5

84

April

19

22

April 14 and
May 7

22

April

11

Coalmont

29

April

8

35

April

22

Cowdrey 115

9

May 2 and 9

2

April

11 and 21

Deer Creek

27

May 17

1

May 13

Delaney Butte

0

Fish Hatchery

69

May 5

Hound

33

April

Lost Creek III

18

May 1

Creek 112

0

0

Monahan Draw

6

April

Railroad!/

86

Ridge Road

Boettcher

Junction

5

North Park,

0
47

April

29

66

April

13

14

April

24

1

April

21

May 14

3

May 24

27

April

33

April

25

Riley

12

May 9

11

April

21

Roth

14

April

22

April

11

Lost

14

21

29

11

Spring

Creek III

49

May 9

62

April

22

Spring

Creek tI2

14

May 2 and 9

1

April

26

Spring

Creek 114

3

April

23

2

April

23

Walden Reservoir

34

April

27

36

April

21

Wattenberg

18

April

26

67

April

5

Total

588

1/ Located

in 1975.

573

�-92-

Capture and Banding
Intensive efforts to capture and band sage grouse in 1975 were initiated on
1 April. Prior to this date, 11 birds (5 males and 6 females) were trapped
incidental to other activities in the 1975 calendar year. During the 1
April-26 May interval, 410 sage grouse were banded. One chick of the year
was banded in July for a total of 422 sage grouse banded in North Park in
1975 (Table 2).
Table 2.

Sex and age composition of sage grouse banded in North Park 1975.
Males

Females

Adult

153

67

Subadult

138

63

Chick

0

0

1

Totals

291

130

1

Unknown

Distribution of bandings within North Park in 1975 was not equal as only 24
grouse were banded in the northeast portion of the park, while 87 were banded
in the southeast part of the park. In contrast, 168 birds were banded in the
southwest quadrant, while 143 were banded in the northwest quadrant. Most
trapping was accomplished on strutting grounds, consequently, few birds were
trapped in the northeast part of the park, as only one known ground occurs in
this area. The same is true for the southeast portion of the park, as only
5 active grounds are known in this area. Thus, distribution of the banded
sample reflected known distribution of strutting grounds.
Sex and age composition of the trap sample does not reflect the actual compo-sition of the population. Based on the 1975 fall harvest sample, actual
composition of the spring population was 37 percent males and 63 percent females,
somewhat higher than the 31:69 male to female ratio found by ~eck (1975) in
late winter. Age composition in the spring of 1975 should have approximated
about 31 percent immatures (males and females) to 69 percent adults. Trap
samples portray a more even age ratio, primarily because yearlings habituate
strutting grounds longer than adults in spring and are probably less wary
than experienced, older birds. Trap samples also indicate that males outnumber females by more than 2:1 in the spring population. This is not the
actual situation, but rather reflects behavior patterns as males roost at
night on strutting grounds where they are vulnerable to trapping. Females
cannot be found at night on strutting grounds except during a short period
and are not as vulnerable to night lighting and trapping as males.

�-93-

Females were poorly represented in the trap sa~ple9 as only 130 were banded.
While females outnumber males in the population they are difficult to locate
at night as they do not commonly roost on strutting grounds.
Of the sample
of 130 females, 25 were trapped along roads at Qight. while the remainder
were trapped on strutting grounds, in early morning with cannon nets (53)
or at night (52).
In the course of trapping operations, 17 grouse banded in prior years were
recaptured.
All had been banded as sub adults in either 1973 (!~males, 2
females), or 1974- (9 males. 2 females).
The oldest birds in this sample
were 3 years of age. Trapping and banding were not accomplished in North
Park in the 1970-1972 period.
Little use can be made of the recapture data
other than longevity, as efforts were made not to recapture birds
that were obviously banded.
This selection could be made while trapping at
night9 as sage grouse frequently stand just prior to being netted.
Hunting

Season Data Collection

The sage grouse season in North Park (Unit 12) in 1975 opened at sunrise on
13 September and closed at sunset on 21 September.
This was the longest
season for sage grouse held in North Park in modern times.
The daily bag
limit was two and the possession limit was four. All hunters hunting sage
grouse in North Park were required to have in their possession a free special
permit.
Permits unlimited in number were available at all license agents in
North Park, the Fort Collins. Colorado Springs and Denver offices of the
Division of Wildlife, and all Wildlife Conservation Officers and research
personnel working in North Park.
Purpose of the permit was to obtain reliable estimates concerning hunter activities in North Park.
Check Stations
Three check stations were operated in 1975.
These stations were located at
the State Line, Gould and Willow Creek Pass.
The station at Muddy Pass was
not operated, as only 40 hunters who harvested 39 birds were checked at this
site in two days in 1974. As in 1974, each station was operated from about
1000 to 1800 MDT, depending upon traffic load.
Check stations were operated
on the 13th. 14th, and 21st. At least one research person and two Wildlife
Conservation Officers were assigned to each check station.
Data obtained per
party were:
County of origin, number of hunters, hours hunted (total of all
hunters). birds observed. birds bagged, birds lost. number of banded birds
and location of where each was obtained, and area hunted.
Most importantly.
one wing was obtained from each bird that was checked.
Few birds (67) were
completely wingless.
Ovaries were collected from over 100 females, and sex
by gonadal inspection was ascertained for about 210 birds.
During the three days of check station operation. 738 hunters ~","ith
551 sage
grouse were checked (Table 3). These hunters reported observing 5,735 sage
grouse.
Some duplications are undoubtedly present in the 5,735 observations.
Hunter efficiency was low (9.6 percent) (551 birds harvested 7 5,735 birds
observed), but only 39 birds (7.1 percent of those retrieved) were reported
crippled and Los t , Hunter efficiency and crippling loss were slightly
higher in 1975 than in 1974 (7.7 and 5.1 percent, respectively).

�Table 3.

Year

North Park sage grouse hunter checks, 1963-1975. -1/

Bag
Limit

1963
2
1964
2
1965 'l:.../ 2
1966
2
1967
2
1968
2
1969
2
1970
2
1971
2
1972
2
1973
2
1974
2
1975
2

Hunters
Checked

Hours
Hunted

Adult
Cocks

Adult
Hens

Percent
Adult

Juvenile
Cocks

Juvenile
Hens

Percent
Juvenile

592
217
150
306
300
546
662
564
357
452
366
730
738

2,460
624
626
1,227
1,177
2,604
2,936
2,617
1,802
2,377
1,599
5,912
5,596

62
25
27
31
50
80
79
58
28
50
24
94
98

150
81
30
116
127
135
180
89
67
69
54
254
179

42
59
49
56
67
42
70
55
51
67
52
50
57

113
28
26
45
42
156
37
53
54
40
29
171
96

181
45
26
71
46
141
74
66
39
18
42
179
111

58
41
51
44
33
58
30
45
49
33
48
50
43

Unclassified
(Dressed)
Birds

- 3/

--J/
- 3/
- 3/
- 3/
- 3/
199

76
76
44
87
67

Total
Birds

Birds
per
Hunter

Hours
Hunted
per Bird

506
179
116
263
267
512
569
266
264
253
193
785
551

1.03
0.82
0.77
0.86
0.89
0.94.
0.86
0.47
0.74
0.56
0.53
1.08
0.75

4.86
3.49
5.40
4.67
4.41
5.09
5.16
9.84
6.83
9.40
8.28
7.53
9.85

1/ Data from 1963 through 1973 are based on Walden and Cowdrey check stations operated during opening weekends. (These
stations checked ha~lested birds primarily from the northwest part of North Park). Data from 1974 are based on four check
stations (State Line, Gould. Muddy Pass and Wi1lmv Creek) operated during the opening weekend, (These stations sampled
all of North Park). Data from 1975 are based on three check stations (State Line, Gould, and Willow Creek) operated for
three days (Saturday, Sunday, Sunday) of a 9 day season.
~/ A one day season; hence only a one day check.

1/ Composition of kill projected from w.i.ng samples; hence no unclassified birds.

I

\0
.j:-o
I

�-95-

Distribution of the harvest and hunting pressure within North Park was
uneven in 1975. Leading harvest zones were the Lake John area (24.7 percent),
Walden Reservoir (16.7 percent), Ridge Road (15.6 percent), Peterson RidgeMcFarlane Reservoir (14.3 percent), and Eagle Hill (9.8 percent). All other
zones (Pole Mountain, Independence Mountain, Owl Ridge-Spring Creek, and
Michigan River Southeast) contributed only 15.8 percent of the total harvest.
Point of harvest could not be ascertained for 3.1 percent of the birds
checked. In 1975 percent of total harvest checked increased 8.3 percent from
the Lake John area and decreased 7.2 percent in the Owl Ridge-Spring Creek
area from levels determined in 1974. With minor exceptions, distribution of
hunters closely coincided with distribution of the harvest.
Most hunters contacted at check stations were asked whether or not they
normally hunted in North Park. Of the 666 hunters responding, 432 (64.9
percent) reported that they normally hunted in North Park, 187 (28.1 percent)
were first time sage grouse hunters, while 47 (7.0 percent) reported that
they normally hunted elsewhere. Percentage response to this question was
almost identical in 1974 (68.3, 24.3, and 7.4 percent, respectively) and 1975.
Two important points are apparent from these data. First, there is substantial annual turnover in sage grouse hunters and second, more retrictive
seasons west (1974 and 1975) and south (1975) of North Park did not result
in major shifts of hunters into North Park.
Wing Analyses
In 1975 wings were received from 505 sage grouse (484 from check stations and
21 from field checks). Classification of these wings by age and sex gave the
following results:
Immatures
212 (42.0%)
Yearlings = 111 (22.0%)
Adults
182 (36.0%)

Males = 101 (47.6%)
Males = 52 (46.8%)
Males = 55 (30.2%)

Females = 111 (52.4%)
Females
59 (53.2%)
Females = 127 (69.8%)

These data are somewhat dissimilar from those collected in 1974 in that chicks
comprised 42.0 percent of the harvest in 1975 (50.1 percent in 1974) and the
sex ratio of yearlings in the harvest in 1975 was nearly even (46.8 percent
males:53.2 percent females); whereas in 1974 it was distorted (35.5 percent
males:64.5 percent females). This reduction in sex ratio distortion between
1974 and 1975 was also evident in adults (30.2 percent males:69.8 percent
females in 1975 to 21.4 percent males:78.6 percent females in 1974). Thus,
production was less than excellent in 1975, overwinter survival of chicks
hatched in 1974 was slightly better than in 1973 to 1974, and overwinter
survival (1974 and 1975) of birds hatched prior to or in 1973 was good. All
of these factors contributed to an older and smaller fall population in 1975
than in 1974.
Evidence from the 1975 hunting season continues to support the hypotheses of
even sex ratios at hatching, and disproportionate survival in older age classes
favoring females.

�-96-

Examination of the molt of primary wing feathers of immature sage grouse
harvested in 1975 indicate that hatching started the week of June 1-7, peaked
about June 25 to July 5 with some broods still hatching as late as July 27.
This is about 7 to 10 days later than in 1974. Wings of 185 females (adult =
126, yearling = 59) were classified as to wing molt. An estimated 48.6
percent (adults = 53.2 percent, yearlings = 39.0 percent) of this sample
were successful in nesting. This is only average nesting success. The young
to adult hen (including yearling hens) ratio in the harvest was 1.1:1,
slightly less than the 1.4:1 ratio in 1974.
Hunter Questionnaire
A total of 1,541 permits were issued for hunting sage grouse in North Park
in 1975, an increase of 357 (30.2 percent) over 1974. As in 1974, most
hunters (47.4 percent) were from the metro Denver area (Adams 6.4 percent,
Arapahoe 8.5 percent, Denver 16.8 percent Jefferson 15.6 percent). Only 8.4
percent of the hunters originated from Jackson County, with 20.9 percent from
Larimer County, 8.1 percent from Boulder County, and 5.2 percent from Weld
County. All other counties each contributed less than 2 percent of the total
hunters and only 9.1 percent of all hunters.
Questionnaires were sent to all permittees immediately follOWing the sage
grouse season in North Park. Responses were received from 905 permittees.
In mid-October a followup letter was sent to all non-respondents and 204
additional responses were received. In all, 1,356 permittees (88.0 percent)
responded (Table 4). Only 19 questionnaires were undeliverable. Mean
values calculated for permittees responding to the followup letter were
used to project for the 12.0 percent (185) non-respondents.
From data in Table 4 it is apparent that 77.0 percent of all permittees went
.hunting, and only 38.3 percent of those actually hunting were successful.
These hunters harvested 979 sage grouse and reported crippling and losing an
additional 74 birds. The total calculated harvest in North Park in 1975 was
1,053 birds, down slightly (10.3 percent) from the 1,174 birds harvested in
1974.
Time period of harvest was received for 862 birds. Of this sample, 83.3
percent (718) was harvested on the opening weekend (13 and 14 September),
8.2 percent (71) during the week (15-19 September), while 8.5 percent (73)
was harvested during the last weekend (20 and 21 September). It is not known
how many birds are harvested on the last day of a three day season, consequently, direct comparisons cannot be made. However, if it is assumed that
85 percent of the harvest in 1975 occurred during the initial three days of
the season, then the longer season in 1975 resulted in an additional 158 sage
grouse being harvested, for a 15 percent increase in total harvest. Contrary
to impressions from field observations, more hunters were afield during the
opening weekend in 1975 than in 1974. In 1974, hunters had few choices when
they could hunt. Thus, all hunters (960 actual hunters) hunted sometime
during the three day season. In 1975, hunters had a choice of when they
wanted to hunt. Examination of questionnaire data reveal that 88.6 percent
of all hunters (1,052) hunted during the opening weekend. Only 5.2 percent
(62) hunted during the week, while 6.9 percent (82) hunted during the second
weekend. (The total of 1,196 hunters is more than the 1,187 calculated in
Table 4 because a few hunters participated in more than one time period).

�-97Table 4.

North Park sage grouse hunter questionnaire data, 1975.

No. in sample

1,356

204

1,152

88.0

Projected
for

Projected for
1,541

185

1,541

12.0

100.0

Percent of total
permittees

74.8

13.2

No. of hunters

905

148

Percent hunters

78.5

72 .5

77.7

72.5

77.0

No. of non-hunters

247

56

303

51

354

Percent non-hunters

21.5

27.5

22.3

27.5

23.0

No. of successful
hunters

373

43

416

39

455

Percent successful
hunters

41.2

29.1

39.5

29.1

38.3

No. of hunter days
Days/hunter
No. of sage grouse
bagged

1,594

257

1.76
806

91

2,084

233

1,851

1.74

1,187

134

1,053

1.76

1. 74

979

82

897

1. 76

Grouse/permittee

.70

.45

.66

.45

.64

Grouse/hunter

.89

.61

.85

.61

.82

Grouse/successful
hunter

2.2

2.1

2.2

2.1

2.2

No. of grouse lost

66

4

70

4

74

.073

Crippling loss/
hunter
Total kill

872

.027

95

967

.062

.027

.066
86

1,053

Percent crippling
loss

7.57

4.21

7.23

4.65

7.03

Percent success
of permittees

32.4

21.1

30.7

21.1

29.5

Banded birds

60

1

61

1

62

Non-deliverab1e
surveys

19

0

19

o

19

Percent nondeliverable

1.23

�-100percent of the cocks banded in 1975 harvested (Fish Hatchery,S of 36 = 13.9;
Alkali Lake, 2 of 14 = 14.3; Walden Reservoir, 2 of 14 = 14.3; Lost Creek No.
1, 2 of 13 = 15.4; Railroad,S of 27 = 18.5).
Considering all recoveries in 1975 of males banded from 1973 to 1975, a
mlnlmum of 7 of a high count of 33 males were harvested from Hound Strutting
Ground, 6 of 68 were harvested from Alkali Lake, 7 of 69 from Fish Hatchery,
3 of 34 from Walden Reservoir, 3 of 12 from Riley,S of 86 from Railroad,
with all other grounds having 2 or less banded cocks harvested. It is not
presently known at what level of harvest decreases will occur in peak numbers
of cocks counted on grounds the following spring. However, the 21 and 25
percent removal of the total cocks counted at Hound and Riley strutting
grounds may result in decreases in peak cock counts in 1976 at these grounds.
Estimation of Fall Population Size
Estimates of fall population size are difficult to derive, primarily because
of unknown vulnerability of chicks and inexact mortality rates for all age
classes. Provided that birds of all age and sex classes are equally vulnerable to hunters and that the annual mortality rate of all banded sage grouse
approxiamtes 50 percent, the 1975 fall population size can be estimated as
follows:
No. of
Birds Banded
1973 = 288
1974 = 191
1975

Alive in 1974

Alive in 1975

Assumptions

144

72
95

50 percent dying each year
50 percent dying from 1974
to 1975
10 percent dying by 1
September

422

380

Total banded birds alive on 1 Sept.

547

Thus, there were 547 banded birds possible in the fall population, of which
62 were harvested. If 58 percent of the 1,053 birds harvested (Table 4) were
adults and yearlings (from wing analysis) then 611 of the birds harvested
were older than one year. Of this number, 62 (10.1 percent) were banded.
Thus, there should have been about 6,100 adult and yearling birds alive in
North Park in the fall of 1975 (if 10.1 percent = 611 birds, then 100 percent = 6,100 birds). Immature birds comprised 42 percent (from wing analysis)
of the harvest, thus there should have been about 4,000 chicks in the 1975
;i.on· tpe fa ~
fall population. The total

rgr1JJ.~

u

ot

I nou

,~~~!
J + ,00.

Of the 293 adult and yearling birds for which wings were available, 186 (63.5
percent) were from females. Thus, of the approximately 6,110 adult and yearling birds in the preseason population, 3,880 (6,110 x 63.5 = 3,874) were
hens. Thus, total chicks in the fall population in 1975 based on 1.1 chicks
per hen in the harvest (1.1 x 3,874 = 4,261) approximated 4,261. This estimate is remarkably similar to the 4,000 chicks in the fall population estimated from overall age ratios and band recoveries win the 1975 harvest.
These data do not suggest that chicks are more vulnerable to hunting than are
adul ts.

�-101It is interesting to compare estimates of adults and yearling alive in the
spring and fall. Spring estimates based on strutting ground counts indicate
that only about 4,500 birds (4,458) were present, while fall estimates
based on age and sex ratios in the harvest and band recovery data indicate
that about 6,100 adult and yearling birds were present. This is a difference of about 1,600 birds. It is probable that spring estimates are low
because either (1) more strutting grounds occur than are known or suspected, or (2) peak strutting ground counts represent less than 50 percent of
the cocks present in an area. It is also possible that fall estimates based
on harvest age and sex ratios and band recoveries are somewhat high because
(1) hunters are not adequately sampling the actual sex and age composition
of the population, (2) banded birds are unevenly distributed in the population, (3) banded birds are more vulnerable than unbanded birds, or (4) large
numbers of bands recovered are not reported.
LITERATURE CITED
Beck, T.D.I. 1975. Attributes of a wintering population of sage grouse,
North Park, Colorado. M. S. Thesis, Colorado State Univ., Ft. Collins.
49 pp.
_____ , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of
sage grouse from wing characteristics. Game Inf. Leaflet No. 49
(revised). Colo. Div. Wildlife. 4 pp.
Braun, C. E., and T.D.I. Beck. 1976. Effects of sagebrush control on
distribution and abundance of sage grouse. Colorado Div. Wildlife,
Final Rept., Fed. Aid Project W-37-R, Work Plan 3, Job 8a. In Press.
Carr, H. D. 1967. Sage grouse and sagebrush control.
rado State Univ., Ft. Collins. 106 pp.

M. S. Thesis, Colo-

Eng. R. L. 1955. A method for obtaining sage grouse age and sex ratios
from wings. J. Wildl. Manage. 19(2):267-272.
Gill, R. B. 1965. Distribution and abundance of a population of sage grouse
in North Park, Colorado. M. S. Thesis. Colorado State Univ., Ft.
Collins. 187 pp.
Patterson, R. L. 1952.
Denver. 341 pp.

The sage grouse in Wyoming.

Sage Books, Inc.,

Pyrah, D. G. 1963. Sage grouse investigations. Idaho Fish and Game Dept.,
Wildlife Restoration Div., Job Compl. Rept., Fed. Aid Project W-125-R.
71 pp,
Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male
sharp-tailed grouse. J. Wildl. Manage. 38(4) :616-621.

�-102-

Robel, R. J. 1969. Movements and flock stratification within a population
of blackcocks in Scotland. J. Anim. Ecol. 38(5):755-763.

P

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Clait E. Braun
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Wildlife Researcher

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�April,

1976

-103JOB PROGRESS

COLORADO

State of
Project

REPORT

W-37-R-2,.::...9

No.

_

Game Bird Survey

Job No.
9 - Part 2
Hork Plan No. -----=3
Evaluation of the Effects of Changes in Hunting Regulations
Job Title __ •..•
--- ~-~ n'rOO ~ Invest;
S-"'8C1';g€.-Grouse· Nes..t.i.n.g-S.ucC=---..c1J.1..LL-U
gat; ons

on

Q

Period

Covered:

Personnel:

1 July 1975 to 31 March

1976

Carrol Donner and Fred Giese, U.S. Fish and Wildlife Service,
Heather Alexander, Tom Beck, Clait Braun, Howard Funk, Don Gore,
Ed Kautz, Sue McElderry, Steve Porter, Sue Quinlan, Steve Steinert,
John Wagner of Colorado Division of Wildlife.

ABSTRACT
Studies were initiated in 1975 to develop techniques to determine production
of sage grouse (Centrocercus urophasianus)
(nesting success and brood size)
prior to September 1. In all, 184 sage grouse were observed during July
and August.
Twelve single broods and 6 groups of multiple broods were
counted.
Young to mature hen ratio for all broods was 1.7:1.
Comparing
systematic searches on foot to roadside searches from a vehicie, broods
were more readily located and observed from the road. Responses to tape
recorded chick distress calls were immediate from hens with young broods
but varied as broods matured.
Early in July broods occupied medium to
dense sagebrush (Artemisia spp.) bordering areas of succulent vegetation.
In August broods moved to the more open areas frequented by broodless hens
and adult males.
P'recf.pLt a t Lon over the summer months was less than normal.
Although the Weather Bureau in North Park reported a marked difference in the
amount of rainfall north and south of Owl and Peterson Ridges in August,
measurements from rain gauges at selected sites over the park indicated a
relatively uniform precipitation distribution.
Estimated nesting success
from wings of 185 females was 49 percent (adult = 53 percent, yearling = 39
percent).
Only 75 percent of sexually mature hens appeared to have ovulated
(adult = 77 percent, yearling = 67 percent).
The sex ratio of juveniles in
the harvest was nearly even, however, evidence to conclude that vulnerability
of chicks to hunting is the same as for yearlings and adults is lacking.

�-104-

RECOMMENDATIONS

1.

Efforts should be increased to locate broods
systematic searches early in summer.

2.

Methods

3.

Ovarian material should continue
colorless fixitive is advised.

should be devised

through the use of

to capture chicks and hens in August.
to be collected

and the use of a

�-105-

NESTING

SUCCESS AND BROOD INVESTIGATIONS
Heather

F. Alexander

In 1975 an intensive la-year study to determine the effects of sagebrush
spraying on sage grouse in North Park, Colorado was completed.
During
the 1963-74 period, some population data were obtained through strutting
ground counts, trapping and banding operations and hunter check stations.
In 1975, intensive studies of the population dynamics and effects of
hunting on sage grouse in North Park were initiated.
As one part of the
overall study, investigations were made of techniques suitable for locating
broods, capturing chicks and estimating production levels.
Production is the key factor influencing population size prior to the hunting
season.
In order to accurately estimate production, improved techniques to
census broods and greater understanding of brood ecology are necessary.

P.N.O. OBJECTIVE
The objectives of this study are to increase hunter opportunity and recrea~
tional use of sage grouse in North Park, Colorado.
Hypotheses which have
been developed are:
(1) hens with broods readily respond to recorded chick
distress calls, (2) distribution of hens with broods is related to moisture
and vegetation patterns within 2 air miles of strutting grounds, (3) differences
exist between yearling and adult hens in percent ovulating, and nesting success
and (4) vulnerability to hunting of both sexes of chicks is the same as for
adults and yearlings.

SEGMENT OBJECTIVES
1.

Chicks will be located and trapped in brood areas during late July and
August through use of tape recorded calls and noosing.
Some chicks may
be caught through use of mist nets and drive traps. A sample of 200
chicks (100 males and 100 females) will be marked each year with colorcoded leg bands or wing tags. Bandings will be evenly distributed in
four areas of North Park.

2.

Nesting success will be estimated through counts of successful and unsuccessful hens in July and August.
Successful females will be located
through use of tape-recorded calls, while unsuccessful hens will be
located and counted in known summer use sites.
A sample of 200 hens
will be classified each year.

3.

Brood size will be determined by counts of chicks accompanied by hens
in July and August.
A sample of 100 broods will be counted each year.

�-106-

4.

August precipitation will be determined from 12 rain gauges placed
at selected sites throughout sage grouse habitat. Sites will be
selected that typify particular areas within North Park. Each rain
gauge will be sampled once a week. Data collected will be compared
to that collected by -t.he U. S. Weather Bureau in Walden.

5.

Check stations will be operated at three locations each year during
the first two weekends of the hunting season. These stations will
be at Gould, State Line and Willow Creek Pass. A goal of 500 wings
is desired each year.

6.

Age composition of the harvest will be determined through examination
of wings collected at check stations. Classification will be as adult,
yearling, juvenile, and male and female.

7.

Compile data, analyze results and prepare progress report.

REVIEW OF LITERATURE
Much of the published literature on sage grouse mentions various aspects
of brood ecology as one segment of a brooder research topic. Relatively
few research projects have been devoted to broods exclusively. Notable
among these are studies by Walles tad (1970) on dispersal and habitat use
and Peterson (1970) on distribution and food habits.
Patterson (1952) contributed much to the overall knowledge of sage grouse
life history. Census techniques for sage grouse broods have been discussed
by Rogers (1964), Eng (1952), Klebenow (1969) and Martin (1970). Eng (1955),
Pyrah (1963), and more recently Beck et al. (1975) described sex and age
determination from wing characteristics. Food habits were intensively
studied by Griner (1939) and Klebenow and Gray (1967); while Gill (1965),
Eng (1952), and Wallestad and pyrah (1974) discussed dispersal. Three.
sources (Stanton 1958, Pyrah 1958 and Dalke et ale 1963) described
collection and examination of sage grouse ovaries.
By far the majority of publications on sage grouse during the past 10 to
15 years have dealt with habitat requirements of all sex and age classes.
Important studies are those of Carr (1967), Klebenow (1972), Eng and
Schladweiler (1972) and Martin (1970).

METHODS AND MATERIALS

Brood Observations
Hens with broods were located during July and August by two general methods;
systematic searches of selected areas and observations along roadsides.
Systematically searching for broods involved walking moist areas and drainages
within 2 air miles of known strutting grounds with a tape recorded chick distress call (Braun 1969 and Walles tad 1970). Other areas within North Park
which appeared to be suitable brood habitat were selected to census in the
same manne r•

�-107Searches for broods were also made from a vehicle traveling previously
established brood census routes (Gill 1965) and other suitable appearing
areas.
These routes were traveled shortly after sunrise at a speed of
approximately 15 to 20 miles per hour.
For routes traveled successive
mornings, direction of travel was alternated.
Tests for response to tape recorded chick distress calls were conducted
of all grouse encountered.
Standardized observation cards were used to
record date, time, number of each sex and age of grouse observed, and
to describe data on location, weather, vegetation and brood behavior.
Vegetative characteristics of brood sites were described qualitatively.
Gills' (1965) classification of Artemisia tridentata was used to describe
sagebrush growth forms; dense, vigorous sage (A-3) , sagebrush in open
stands of medium height (A-2), and sparse sagebrush characteristic of
ridge tops and poor site conditions (A-I).

Trapping

and Banding

Once broods were located, attempts were made to capture chicks.
As all
birds were quite capable of flight, two methods of capture were used.
A 5 m
telescoping noose pole described by Braun (1969) equipped with 40.5 Kg
(90 lb) plastic coated steel wire and a short handled hoop net were tested.
The handle of the hoop net was approximately 1.5 m in length with a 0.6 m
diameter hoop.
Upon capture, chicks were banded with numbered aluminum patagium tags if
age was less than 6-8 weeks or numbered aluminum leg bands if over 8 weeks
of age. All birds captured were weighed with primary length and molt, and
possible age and sex being recorded.

Precipitation

Measurement

Twelve standard sunken or pit rain gauges were randomly established in sagebrush types over North Park to measure precipitation and describe precipitation
patterns prior to the hunting season.
Each gauge was constructed for temporary use from two 3.8 liter (1 gal) tin
cans soldered together.
Inside diameter was 15.2 em (6 in). Splash shields
were made from burlap approximately 1 m square.
Rodent screens of 0.1 em
(1/4 in) wire mesh were installed about 20 em below ground level.
Diesel
fuel was used to retard water evaporation.
All rain gauges were checked and weighed weekly

Wing and Ovarian

through August.

Examination

During the 1975 sage grouse hunting season, three check stations were operated
in North Park.
These stations were located at the State Line, Gould and Willow
Creek Pass on September 13, 14 and 21.

�-108One wing from most birds checked and ovaries from most mature females
retaining them were collected. Each ovary was placed in a numbered vial
containing Bouin's fixitive. The vial number was recorded on a tag
attached to the wing taken from the same bird.
All wings were examined and assigned to age and sex classes. Nesting
success was estimated for yearling and adult females from primary wing
molt (Beck et ale 1975). Ovaries were examined and classified as having
or not having ovulated following techniques described by Meyer et ale
(1947).

STUDY AREA
North Park, Colorado was chosen for field study due to past sage grouse
research and management efforts in this area.
North Park (Jackson County) borders Wyoming in north-central Colorado,
immediately west of the Front Range of the Rocky Mountains. The park is
essentially a semi-arid basin enclosed by mountains. The topography
is generally flat to rolling varying in elevation from 2,400 to 2,585 m
above sea level. Noted exceptions are two prominent ridges extending
northwest seemingly splitting the park into north and south segments.
About 64 percent of the land area lies to the north of Owl and Peterson
Ridges. The geography of North Park has been completely described by
Beck (1975).
Two main vegetative types are evident. Big sagebrush (Artemesia tridentata)
dominates the sagebrush type which occupies approximately 1,252 sq km, almost
one fifth of the area of North Park. Irrigated native hay meadows make up the
second prominent vegetative type, nearly 620 sq km or about one third of the
suitable sage grouse habitat (Beck 1975).
Climatically, North Park can be described as part of the Upper Sonoran life
zone receiving less than 65 em of precipitation annually, about half falling
as snow in autumn, winter and spring. During the summer months (June-August),
average preCipitation is normally 9.25 em north of Owl and Peterson Ridges
(U. S. Weather Bureau, Walden) and 10.87 em to the south (U. S. Weather
Bureau, Spicer).
Fairly uniform seasonal temperatures and extreme temperature fluctuations
between night and day are characteristic of montane regions (Trewartha 1968).
Constant winds are also a factor in North Park.

RESULTS AND DISCUSSION
Brood Observations
Systematic Searches
Five of 12 single sage grouse broods were observed by systematic searching
during July and August in North Park. Complete counts of 2 of these broods of
young chicks were not obtained due to concealment of the young (Table 1).
Two multiple broods were encountered with a young to female ratio of 1:2.

�Table 1.

Brood observations, July and August 1975, North Park.

Single Broods
Method
Systematic
Search

Sub-total
Roadside
Search

Sub-total

Females

youngll

1
1
1

2
2
3

3

7

1
1
1
1
1
1
1

3
5
5
4
1
3
1

7

22

y/F

Rati~1

2.3:1

3.1:1

Mu1tiE1e Broods
young·!.!
y/F Rati~1
Females
6

3

1.0:2

2

1

1.0:2

8

4

1.0:2

3
6
3

7
12
7

2.3:1
2.0:1
2.3:1

12

26

2.2:1

I
I-'

0

\D

Total

11
21

11

l(}~j

29

2.9:1

20

Counts of chicks are considered to be minimal.
Ratio = Young to Female.
Of 12 broods observed, counts of chicks were made on only 10.

y/F

30

1.5 :1

I

�-110Only 2 hens with broods responded verbally to the tape recorded chick
distress call. No hens with mUltiple broods responded.
Most broods were encountered in early morning between 0730 and 0930.
Although some broods were sighted in mid-day, none were observed in
the evening. Seventeen unsuccessful hens and 3 adult males were also
observed.
Roadside Searches
Seven of 12 single broods with a young to hen ratio of 3.1:1 were first
observed from a vehicle. Although the established Lake John brood census
route was traveled to a lesser extent than generally traveled throughout
North Park, broods were observed on only the Lake John route. This route
was traveled six times in July and August. Three broods were counted on
two mornings. Multiple broods were observed late in August. The young
to female ratio for all single broods observed was 2.2:1 (Table 1).
As with systematic searches, most broods were observed in early morning
between 0730 and 0900. Only one brood was sighted in the evening. Of
the broods observed, only one hen with young responded verbally to the
chick distress call. Fourteen broodless females and 68 males were
observed.
Considering all broods counted, the young to hen ratio was approximately
2.0:1, somewhat higher than the 1.1:1 ratio calculated from wings taken
during the 1975 sage grouse hunting season.
It would appear from these data that broods are more readily located
and counted from the road.
Response to Tape-recorded Chick Distress Calls
A variety of both verbal and behavioral responses to the tape recorded
chick distress call were encountered. Although some overlap exists,
response patterns associated with particular groups are apparent.
Hens with broods of young chicks (less than 5 weeks) reacted to the call
in much the same way as ptarmigan females (Braun et ale 1973). Depending
on the visibility and proximity of the observer, the hen either moved
slowly toward or away from the source of the call. In one instance, a
female flew about the head of the observer. Verbal responses were given
immediately. Hens continued to make low guttoral noises as the call was
repeated. They were reluctant to fly and did so only if approached too
quickly. Flights were short and the hens appeared reluctant to leave the
area where their broods were assumed to be.
Hens with older broods usually did not respond verbally to the call of a
young chick. In all instances the hen remained still, either alert or
squatting depending on the visibility of the observer. These hens took
flight readily, usually flying over 300 m.

�-111When the call was played to chicks, they would freeze or hide beneath
sagebrush where they were extremely difficult to locate.
Birds five
weeks and older would fly readily from sudden movement or noise.
Some
vocalized as they flushed.
Broodless females were highly variable in their responses.
Early in
July, unsuccessful hens became alert at the sound of the call but no
other response could be detected.
When the call was played to broodless
females observed late in July, the call produced a similar alert response.
With continued calling, two of the hens approached the vehicle within 15 m.
Both appeared agitated and clucked while moving about in one spot and
looking at the vehicle.
Neither noticed a badger (Taxidea taxus) which
emerged from a hole immediately behind the hens and then disappeared.
These two broodless hens may have lost their broods just prior to being
observed and tested.
Groups of males in general showed no apparent
reaction to the call.
From these observations it would appear that chick distress calls were
effective in eliciting responses from hens with young broods.
However,
responses decreased as broods matured.
It should be noted that the call
used was that of a young chick.
If recordings were available from older
juveniles, it is possible that most hens with broods may respond.

Trapping

and Banding

Only two chicks were captured and marked.
Both were caught in the southern
portion of North Park within two days of each other on July 25 and 27.
One was captured by hoop net in sagebrush of medium height and density (A-2).
The other was caught with the noose pole in sparse A-I sagebrush type.
Chicks hidden under sagebrush plants were well concealed.
They could not
be observed from the distance required for use of the noose pole.
The
short handled hoop net proved to be useful in this instance.
When chicks
were quite visible, capture was attempted with the noose pole.
Trapping by these means was difficult.
As brood organization broke down
in August and large groups of birds moved to more open areas (pastures
and hay meadows), capture using these methods was not possible.

Characterization

of Summer Use Sites

In July, hens with broods were generally found in medium A-2 sagebrush
adjacent to areas of succulent vegetation.
In many situations, free
water from irrigation ditches was available.
The importance of free
water for grouse broods in North Park is not known.
It is possible all
water requirements are met by eating succulent vegetation.
In August, more broods were observed in open pastures and hay meadows near
willow (Salix spp.).
Frequently free water was available from nearby
creeks and irrigation ditches.
By mid-August most ditches were dry.

�-112Broodless females and adult males, although segregated from hens with
chicks, were often found in similar sites throughout the summer. These
groups of grouse appeared to prefer open areas of sparse sagebrush cover
in July while mowed hay meadows and pastures near willow were used in
August.
It was not uncommon to find large segregated groups of males, and females
and young utilizing the same areas prior to the hunting season.

Precipitation Measurement
From climatological data obtained from the Weather Bureau, U. S. Department
of Commerce, summer precipitation north of Owl and Peterson Ridges was
17.1 percent below the 30 year average while precipitation in the south
over the three summer months was 13.2 percent below normal (Table 2).
August precipitation data obtained from the Weather Bureau Station at
Walden indicated that the northern portion of the park received 62.0
percent more rainfall than the southern portion, 13.1 percent above the
30-year average. During this period, precipitation in the southern portion
of the park was 46.6 percent below normal as derived from the Spicer Station.
Table 2.

Summer precipitation, North Park, Colorado (1975)!~

Station

June

PreciEitation (cm)
July
August

Total

Walden

1.47

2.49

3.71

7.67

Walden, 30-year average

2.82

3.15

3.28

9.25

Spicer

2.82

4.32

2.29

9.43

Spicer, 30-year average

3.48

3.10

4.29

10.87

!/

Climatological Data, U. s. Department of Commerce (1975).

Due to heat and wind, the diesel fuel used to prevent moisture loss in the
12 rain gauges began to evaporate within the first week in the field. Since
the apparent rate of evaporation was low, untreated measurement data obtained
from the gauges were used to examine distribution of rainfall over the park.
Average August precipitation from the rain gauges was obtained from both the
northern and southern portions of North Park (Table 3). From these data,
rainfall appeared to have been evenly distributed over the park. Although
rainfall south of Owl and Peterson Ridges was still below normal, the
difference was less than 20 percent. Rainfall north of Owl and Peterson
Ridges was slightly above average (1.8 percent).
It appeared that rainfall in August paralleled Owl and Peterson Ridges and
tracked NNW of Peterson Ridge.

�-113-

Table 3.

Rain gauge data, August 1975.

1/
Gauge-

Precipitation
(em)

X Precipitation
(em)
3.48

South
Manville Draw

5.85

McFarlane Reservoir

2.57

Deer Creek

2.02

Peterson Ridge

3.67

Lerand

3.28
3.34

North
ViIs Ranch

4.03

Roth

2.53

McCallum

2.56

Canadian River

4.14

California Gulch

2.39

Lake John

2.53

Walden Reservoir

5.19

1/ General location of rain gauges south and north of Owl and Peterson Ridges.
Wing and Ovarian Analysis
Nesting success was estimated from examination of primary molt of wings from
185 females (adults = 126, yearlings = 59) collected at check stations. Since
molt is delayed in females during the nesting and brooding cycle, those females
with primary wing molt resembling that of males (completed molt or retaining
old primaries 10 and 9) were probably unsuccessful in nesting or lost their
broods shortly after hatching. Those females retaining old primaries 10, 9,
and 8 or more were most likely successful nesters. Of all females, 48.6
percent (adults = 53.2 percent, yearlings = 39.0 percent) were estimated
to be successful in nesting (Table 4).
Independent estimates of nesting success were made from counts of brooded
and unbrooded hens observed throughout July and August. Seventeen unbrooded
hens were observed while only 12 females known to be successful were sighted.
Nesting success from this method was estimated to be 41.4 percent. However,
successful and unsuccessful hens could not be distinguished in groups of
multiple broods. Thus, these data may be unrealistic.

�-114-

Table 4.

Analysis of primary molt of hunter harvested female sage grouse, 1975.

Age Class

Primary Molt &lt; 7

Primary Molt &gt; 7

Total

Yearling

23 (39.0%)

36 (61.0%)

59

Adult

67 (53.2%)

59 (46.8%)

126

Adult &amp; Yearling

90 (48.6%)

95 (51.4%)

185

Percent ovulation was determined from examination of 135 ovaries (adults = 69,
yearlings = 39, juveniles = 13) collected at check stations. The appearance
of ovulated follicles on the surface of the ovary signified normal ovulation.
An estimated 24.6 percent of sexually mature hens (adults = 23.2 percent,
yearlings = 30.8 percent) did not appear to have ovulated though many showed
signs of stimulation (Table 5).
Table 5.

Ovarian analYSis of hunter harvested sage grouse, 1975.1/

Age

Unclassified

Non-ovulating

Ovulating

Total

Juvenile

0 (0.0)

13 (100.0)

0 (0.0)

13

Yearling

1 (2.5)

12 ( 30.8)

26 (66.7)

39

Adult

0 (0.0)

16 ( 23.2)

53 (76.8)

69

Unknown (&gt;1 Yr.)

0 (0.0)

2 ( 14.3)

12 (85.7)

14

Adult &amp; Yearling

1 (0.8)

30 ( 24.6)

91 (74.6)

121

1/

Percentages are given in parentheses.

These data do not agree with previous findings. In a study in Wyoming, Pyrah
(1958) presented no data but reported positive ovulation of mature hens greater
than 95 percent. Stanton (1958) reported all adult females (32) from a sample
in Idaho had ovulated while one fourth (5/20) of the yearlings had not.
A number of factors might have introduced error into the results. The use of
the yellow Bouin's solution appeared to mask the reddish pigment indicative
of ovulated follicles as involution progresses (Kabat et a1. 1948). When
ovulated follicles regress in size, some may become indistinguishable from
neighboring follicles. Thus an ovary from an early laying hen may not
appear to have ovulated.

�-115Kabat (1947) studied the resorption rate of ovulated follicles from hen
pheasants (Phasianus colchicus) and concluded that few, if any, had
disappeared by mid-September. It would seem unlikely that birds with
lower ovulation rates, such as sage grouse, would resorb follicles at a
faster rate.
Another source of error may lie in the condition of the ovaries. Although
complete ovaries were believed obtained in most cases, some ovaries may
have been only portions of the total organ. Since only a relatively small
area of the ovary may become active, should this portion be lost, the ovary
would appear non-ovulatory.
Data from both ovarian analysis and examination of wing molt indicate that
adult hens were more successful in laying and brooding than yearlings.

Chick Vulnerability to Hunting
To determine if chick vulnerability to hunting is the same as for adults and
yearlings, a spring population of 5,000 birds was estimated. Winter observations
(Beck 1975), indicate that sex ratios of males to females in early spring approach
30:70. Thus, 3,500 females would occur in North Park out of a total population
of 5,000 (.70 x 5000). For 3,500 females, if we assume 53.2 percent nesting
success from wing molt data in the harvest and a production estimate of 2.0
chicks per clutch (from brood observation data), the resulting fall population
of juveniles would have been 3,724 individuals at the minimum. Not allowing
for spring to fall mortality of adults and yearlings, the fall population
would have consisted of 1,500 cocks (.30 x 5000), 3,500 hens and 3,724 chicks
for a total of 8,724 birds. Of this total, 17.2 percent would be adult and
yearling males, 40.1 percent would be adult and yearling females, while 42.7
percent would be juveniles.
Of 505 sage grouse wings collected during the hunting season, 21.2 percent
(107/505) were adult and yearling males, 36.8 percent (186/505) were adult
and yearling females and 42.0 percent (212/505) were juveniles. Thus, it
would appear that chicks are harvested in the same proportion as they appear
in the population. No differential mortality in the harvest was evident for
juveniles as 47.6 percent were males and 52.4 percent were females. However,
there was some disparity between adult and yearling male and female ratios
in the harvest as all females comprised 63.5 percent of the adult and yearlings
harvested. This may be due to hunter selection for smaller birds. Interviews
of hunters at check stations suggest that most do not have the opportunity
to select birds when hunting since most hunters do not attain the bag limit.
Consequently, all birds flushed within shooting distance are fired upon.

LITERATURE CITED
Beck, T. D. I. 1975. Attributes of a wintering population of sage grouse,
North Park, Colorado. M.S. Thesis. Colo. State Univ., Ft. Collins 49 pp.
_____ , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of sage
grouse from wing characteristics. Colo. Div. Game, Fish and Parks, Game
Information Leaflet 49 (revised) 4 pp.

�-116Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado. Ph.D. Dissertation. Colo. State Univ.,
Ft. Collins. 189 pp.
_____ , R. K. Schmidt, and G. E. Rogers. 1973. Census of Colorado whitetailed ptarmigan with tape-recorded calls. J. Wildl. Manage. 37(1):90-93.
Carr, H. D. 1967. Sage grouse and sagebrush control.
State Univ., Ft. Collins. 106 pp.

M. S. Thesis.

Colo.

Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F. Schlatterer.
1963. Ecology, productivity and management of sage grouse in Idaho.
J. Wildl. Manage. 27(4):811-841.
Eng, R. L. 1952. Detailed studies of population trends and breeding potential.
Montana Fish and Game Corom.,Wildl. Restoration Div. Quarterly Progress
Rept. 3(4):56-62.
_____ . 1955. A method of obtaining sage grouse age and sex from wings.
Wildl. Manage. 19(2):267-272.

----- , and P. Schladweiler.

use in central Montana.

J.

1972. Sage grouse winter movements and habitat
J. Wildl. Manage. 36(1):141-146.

Gill, R. B. 1965. Distribution and abundance of a population of sage grouse
in North Park, Colorado. M. S. Thesis. Colo. State Univ., Ft. Collins.
187 pp.
Griner, L. A. 1939. A study of the sage grouse (Centrocercus urophasianus)
with special reference to life history, habitat requirements, and
numbers and distribution. M. S. Thesis. Utah State Agr. ColI.,
Logan. 83 pp.
Kabat, C. 1947. Reading breeding history from pheasant ovaries.
Univ. Wisconsin, Madison. 38 pp.

M. S. Thesis.

----- , I. O. Buss, and oR. K. Meyer.

1948. The use of ovulated follicles in
determining eggs laid by the ring-necked pheasant. J. Wildl. Manage.
12(4):399-416.

Klebenow, D. A. 1969. Sage grouse nesting and brood habitat in Idaho.
Wildl. Manage. 33(3):649-662.

J.

1972. The habitat requirements of sage grouse and the role of fire
in management. Proc. Tall Timbers Fire Ecology Conf. 12:305-315.

----- , and G. M. Gray.

1967.
Manage. 21(2):80-83.

Food habits of juvenile sage grouse.

J. Range

Martin, N. S. 1970. Sagebrush control related to habitat and sage grouse
occurrence. J. Wildl. Manage. 34(2):313-320.
Meyer, R. K., C. Kabat, and I. O. Buss. 1947. Early involutionary changes
in the post-ovulatory follicles of the ring-necked pheasant. J. Wildl.
Manage. 11(1):43-49.

�-117Patterson, R. L. 1952. The sage grouse in Wyoming. Wyo. Game and Fish
Comm., and Sage Books Inc., Denver, Colorado. 341 pp.
Peterson, J. C. 1970. The food habits and summer distribution of juvenile
sage g~ouse in central Montana. J. Wildl. Manage. 34(1):147-155.
Pyrah, D. B. 1958. Wyoming's sage grouse management program.
Assoc. State Game and Fish Comm. 38:184-186.

Proc. West.

1963. Sage grouse investigations. Idaho Fish and Game Dept.
Compl. Rept., Proj. W-12l-R-2. 71 pp.
Rogers, G. E. 1964. Sage grouse investigations in Colorado.
Fish and Parks Dept. Tech. Publ. 16. 132 pp.

Job

Colo. Game,

Stanton, D. C. 1958. A study of breeding and reproduction in a sage grouse
population in southeastern Idaho. M.S. Thesis. Univ. Idaho, Moscow.
87 pp.
Trewartha, G. T. 1968. An intruduction to climate, 4 ed. McGraw-Hill
Book Co., New York. 408 pp.
U. S. Department of Commerce.

1975.

Climatological data, Colorado. 80(6-8).

Wallestad, R. 1970. Summer movements and habitat use by sage grouse broods
in central Montana. M.S. Thesis. Mont. State Univ., Bozeman. 51 pp.
1975.
Montana.

Life history and habitat requirements of sage grouse in central
Mont. Fish and Game Dept., Game Manage. Div. 65 pp.

_____ , and D. B. Pyrah. 1974. Movements and nesting requirements of sage
grouse hens in central Montana. J. Wildl. Manage. 38(4):630-633.

Prepared by

;iJj~A/(, a~t.~"

Y-d ..z(h4.·
Heather F. Alexander " .;:.'
Graduate Research Assistant

.,1..
Approved by

/

J

~~~~~.~~~.~~~.
__~,c~~~.~/~~~~~~~~d~_~&lt;~~(~/~PL/~·
Clait E. Braun
,"","
Wildlife Researcher

��April 1976

-119JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

10
3
Investigations of the Distribution and Status
of Sagebrush and Sage Grouse in the Moffat County Area

Work Plan No.
Job Title
Period Covered:
Personnel:

Game Bird Survey

W-37-R-29

January 1, 1976 to March 31, 1976

Donald M. Hoffman.

ABSTRACT
A study was designed and implemented to ascertain the distribution and status
of sagebrush and the distribution, status and trends of sage grouse populations
in the Moffat County area. Field work was started on January 5, 1976 and
continued through March 26, 1976.
An inspection of past sage grouse strutting ground census data for the Moffat
County area gathered by Division of Wildlife's Research and Management Personnel indicates populations apparently peaked in 1969 and have apparently been
declining since then. Numbers of active grounds counted each spring have also
declined steadily since 1969, so caution is advised in using these trends
until more information is gathered.
At least eight sage grouse wintering flock locations were identified through
field surveys in the Moffat County area during January, February, and March
1976. Two isolated instances of sage grouse mortalities were found during
this period while checking for sage grouse and field signs.

�-120-

INVESTIGATIONS OF THE DISTRIBUTION AND STATUS
OF SAGEBRUSH AND SAGE GROUSE IN THE MOFFAT COUNTY AREA
Donald M. Hoffman

P. S. OBJECTIVES

Major objectives of this study are to (1) ascertain the distribution of sagebrush in the Moffat County area, (2) determine the status (i. e. principally
alterations) of sagebrush in this area, (3) delineate the distribution of
sage grouse in the Moffat County area, and (4) identify the past and present
trend of sage grouse population status in this area.

SEGMENT OBJECTIVES
1.

Review
Moffat

literature concerning
County area.

vegetative

descriptions

for studies

in the

2.

Ascertain the distribution of sagebrush through examination of vegetative
maps and aerial photographs obtained from public land management agencies.

3.

Examine records of public land management agencies to determine
information is available concerning alterations of sagebrush.

4.

Delineate sage grouse distribution through examination
and Federal Agencies, interviews of selected personnel
field reconnaissance.

5.

Record locations of sage grouse observed
vegetative type.

6.

Ascertain

7.

Compile

data into maps and/or overlays

8.

Prepare

progress

what

of records of State
and landowners, and

by date and time, location,

trends in sage grouse populations

from management

and

surveys.

and tables.

report.

METHODS

AND MATERIALS

A preliminary review of literature concerning vegetative descriptions from
studies in the Moffat County area was made in the Division of Wildlife's (DOW)
Research Center, and Colorado State University (CSU) libraries.
Major areas of sagebrush range were outlined on Colorado Division of Highway's
road maps for Moffat County, scale one-half inch per mile, using types outlined on U. S. Soil Conservation Service's (SCS) range resource maps, combined

�-121with field reconnaissances.
Progress was also made in determining the
distribution of sage grouse through these field reconnaissances.
Unfortunately,
many sage grouse areas in Moffat County could not be checked during the
period January 1 to March 31, 1976 because of impassable roads and trails.
Records of locations, amounts, and types of sagebrush range alterations by
Federal agencies in Moffat and western Routt counties were secured from
Wildlife Researcher Ron Kufeld (DOW).
A search for wintering concentrations of sage grouse was made as weather and
road conditions permitted during the months of January, February and March
1976. Locations of sage grouse observed or reported were recorded by date and
time, location and vegetative type.
Statistics on sage grouse populations using strutting ground counts from Rogers
(1964) the Wildlife Management Section, and the northwest Region for Moffat
and western Routt counties were compiled.
Statistics concerning agricultural activities, human populations, and vehicular
license sales in Moffat County were gathered and compiled since these have a
direct bearing on this study.
Data were compiled

into maps and tables and a Job Progress
RESULTS

Vegetative

Descriptions

Report was prepared.

AND DISCUSSION

from Studies in the Moffat

County Area

A preliminary search of the literature has revealed few vegetative studies
completed in the Moffat County area. Two studies list techniques of measuring
vegetation which may prove useful in this study in future years.
Powell (1968)
secured vegetative measurements of sagebrush species in Moffat County utilizing crown cover, height, and density.
He used a line intercept method of
sampling range vegetation described by Canfield (1941) to determine crown
cover and maximum plant height and procedures described in Weaver and Clements
(1938) to determine density using belt transects.
Vories (1974) inventoried
vegetation in the Piceance Basin and adjacent drainages as part of an extensive Oil Shale Study for the Thorne Ecological Institute.
He used six transect
lines with plots at 400 meter intervals and a nested quadrat system for studying vegetation communities.
The nested quadrat system is described by Phillips
(1959).
During the early 1950's the SCS Office in Denver maintained a file of county
map sheets, scale one-half inch per mile, showing sagebrush ranges.
A
request was made to use these, but they have apparently been misplaced or lost
in recent years.
Personnel of the SCS in Denver are in the process of compiling
a statewide range resource map using work maps prepared by their personnel
stationed in the various counties.
The County work maps were prepared using a
series of range site descriptions which include physiographic,
climatic, vegetative, and soil features.
Copies of the work maps for Moffat and Routt
counties were secured from the Denver office of the SCS. The expected date of
publication
of the Statewide map is not known.
Unfortunately,
for Project

�-122p~rposes, two or three range site types are often combined within the big
sagebrush and complimentary types.
These, however, have been quite useful
in mapping sage grouse ranges through the delineation of big sagebrush types.
Mapping

of Sagebrush

Ranges

Approximately one-half the potential sage grouse range of Moffat County west
of Highway 13 was cover type mapped, using one-half inch per mile scale work
maps.
A copy of the SCS's range resource map for Moffat County, SCS's range
site descriptions, and extensive field surveys were used in this work.
Major
vegetative cover types delineated include: (1) dense (high quality) big sagebrush-grassland,
(2) sparse (low quality) big sagebrush-grassland,
(3) grassland, (4) cultivated dryland farmland, (5) irrigated farmland, (6) pinonjuniper, (7) mountain shrub, (8) greasewood, and (9) steep and rocky.
This
survey of major cover types is needed as a basis for measuring alterations
of sagebrush communities in future comparisons.
A number of areas yet to be cover type mapped are inaccessible during the
winter period, so plans are to survey these during the summer of 1976.
Sagebrush

Alterations

Requests for information on alterations of sagebrush ranges in the Moffat
County area were made at the SCS and U. S. Bureau of Land Management (BLM)
Offices in Denver, DOW Office in Fort Collins, CSU Extension Service, and U. S.
Agriculture Stabilization and Conservation Service (ASC) Offices in Craig.
The best and only complete compilation of sagebrush range conversions on
Federal lands containing sage grouse populations (BLM lands) is maintained by
Ron Kufeld, DOW, Fort Collins.
Copies of all individual project forms and
map locations on all completed range conversions on BLM lands were obtained.
All of these range conversion projects were completed prior to 1970.
The ASC and CSU Extension Service in Craig recommended use of Colorado Agricultural Statistics, published annually by the Colorado Department of Agriculture
to obtain estimates of amounts of cropland and numbers of cattle and sheep
in Moffat County, since they do not maintain these records.
The SCS Office in Craig maintains a complete file of aerial photographs taken
approximately four years ago which may be of use in future comparisons of past
and present vegetative cover types on specific areas.
Sage Grouse Populations
Strutting ground census data obtained from counts made by Wildlife Conservation
Officers on individual grounds from 1969 through 1975 in Moffat County are
listed in Table 1. Similar information for western Routt County is shown in
Table 2. These data were compiled from Wildlife Management Section and
Northwest Region records.
These will be used as a basis for future strutting
ground counts to be made by Project as well as Management personnel.

�11

Table L

Sage grouse strutting ground count data for individual grounds. Moffat County. 1969-1975.

Counter(s)
L. Vidakovich

Name of
Ground
Karrl.!nRanch
Bear Creek
Escalante
S t a teline
(Stuntz)
Stock Pond
Haslim Cow Camp

P. Mason. F.
Rinella or C.
Brown

High
Cock
Count

Total
Birds

109
58
79
94

114
58
83
94

75
23
28
35

161
24
28
35

27

28

-

1972

1971
High
Total
Cock
Count Birds

High
Cock
Count

Total
Birds

87
13
54
63

18
0
2
64

18
0
2
67

87
13
54
63

1973
High
Total
Cock
Count Birds
45
12
33
36

45
12
33
36

1974
High
Cock. Total
Count Birds
78
14
31
75

1975
High
Total
Cock
Count Birds

78
14
31
76

37
12
22
0

37
12
22
0

57

57

53
124

59
130

367

377

161

248

217

217

84

87

126

126

255

256

43
130
132
142

49
137
154
160

33
92
78
0

36
98
86
0

42
52
94
0

42
55
97
0

8
48
45
75
34
20
6

10
56
62
110
46
30
8

0
57
0
0
30
0
16

0
73
0
0
38
0
16

14
18
58
0
19
0
16

14
21
62
0
27
0
16

0
8
110
0
0
0
0
13

0
8
141
0
0
0
0
14

447

500

203

220

188

194

236

322

103

127

125

140

131

163

Subtotals
Spring Creek /11
Cross Hountain III
Cross Hountain 112
Big Gulch 111
Big Gulch 112
Big Gulch 113
Big Gulch 114
Grassie
Clarida Ranch
Lay Creek III
Bord Gulch

30
12
26
35
57
72

30
12
28
39
60
83

18

33
27
0
9
9
12
0
0
0
80
37

35
32
0
10
16
15
0
0
0
93
39

0
0
0
0
20
6
8
0
0
65
0

0
0
0
0
27
9
12
0
0
76
0

0
27
0

0
33
0

0
0
0

0
0
0

71
2
167
10

70
18
32
35
74
20
50
19
0
54
4

18

65
1
167
10

70
17
29
29
50
20
50
19
0
51
4

0
0
0
0

0
0
0
0

19
0
44
10

31
0
51
10

Subtotals
H. Overmeyer
or R. Lowry

1970

1969
High
Total
Cock
Birds
Count

Sand Wash Ridge
Road
Hay Shed
Gee Flats
Beaver Basin
Sand Wash Rim
Goodman Draw
Snake River West
Powder Wash Hill

-

-

21

-3
24
54
36
34
67
44

26

-

3
27
57
41
46
71
44

------------------------------------------------------------------------------------------------------------------------------------------------

I

~

N
W
I

�Table 1.

Sage grouse strutting

1969
High
Cock
Total
Count
Birds

Name of
Ground

Counter(s)
P. Mason, F.
Rinella or C.
Brown (contd.)

H. Dobbs, D.
Morgan or C.
Woodward

Timberlake
Timberlake
Timberlake
Fan Rock

III
112
#3

Subtotals

26

1970
High
Cock
Count
3
131

County,

1971

Total
Birds

High
Cock
Count

Total
Birds

4
152

0
72

0
72

1969-1975

1/
- (continued).

1972
High
Cock
Total
Count Birds

1973
High
Cock
Total
Count Birds

1974
High
Cock
Total
Count Birds

0
0
37

0
42
0

0
42
0

-

25

25

0
0
42

1975
High
Cock
Total
Count Birds

83
0

117
0

-

40
25
0

44
37
0

43

45

15

27

473

532

373

405

244

282

166

191

153

195

153

200

158
190

158
208

66
32
0
142

89
37
0
78

93
37
0
162

169
0
73
130

196
0
79
130

70
12
8
158

70
12
8
281

0
0

96

57
29
0
129

0

96

52
27
155
212

0

-

46
24
131131
137

444

462

338

446

215

240

204

292

372

405

248

371

87
66
57
67
0
70

93
68
59
67
0
70

77
76
54
61
0
38

79
79
57
63
0
40

72
60 '
50
27
1
59

78
65
52
29
1
60

63
60
28
27
0
42
21

76
65
29
29
0
50
25

37
37
13
28

347

357

306

318

269

285

241

274

1,522

1,803

1,299

1,374

1,037

1,268

1,008

1,123

Region

data.

-

Subtotals

405
2,164

Management

Moffat

528

78
141
50
31
38
67

Wildlife

26

grounds,

501

Juniper
Axial Basin
High Mesa
Round Bottom
Dry Lake III
Dry Lake 112
Deception Creek

TOTALS

l/Source:

-

Greasewood 111
Upper 19 Road
Fortification #1
Lay Creek (new
ground)
N. Fork Big Gulch
Sub totals

W. Roland

ground count data for individual

Section

81
144
50
36
38
67

416
2,283 "

and Northwest

0
0

0

0

•....

0

0

+'
I

37
37
17
31

39
51
0
0

49
78
0
0

0
14

0
14

40
1

42
1

129

136

131

170

910 1,098

539

663

'"

�Table 2.

Sage grouse strutting

ground count data for individual
1969

H. Wixson, A.
Greer, J.
Ce rrans , or C.
Hector

Total
Birds

26

26

116

142

Twenty Mile
Upper 18 Road

Name of
Ground

Counter(s)

TOTALS

1970
High
Cock
Count

High
Cock
Count

Routt County,

Total
Birds

High
Cock
Count

21

23

129

100

142

121

Total
Birds

13

13

116

114

142

127

1969-1975.

1972

1971
High
Cock
Count

1)
Source:

grounds, Western

1973

1/
1974
High
Total
Cock
Birds
Count

1975
High
Cock
Count

Total
Birds

9

0

0

27

32

0

0

36

41

0

C

Total
Birds

High
Cock
Count

Total
Birds

20

20

21

23

9

102

68

74

63

73

125

88

94

84

96

•..
I

Wildlife

Management

Section

and Northwest

Region data.

N
VI
I

�-126-

Populations of sage grouse in Moffat County apparently peaked in 1969 and
have apparently been declining since 1969, based upon strutting ground counts
made by Wildlife Conservation Officers.
During the period of record highest
numbers of cocks (2,164) and highest numbers of total birds (2,283) were
recorded for the year of 1969. Less than 600 cocks were counted on strutting
grounds during 1958, 1960, 1964 to 1966, and 1975.
The number of active
strutting grounds counted by Wildlife Conservation Officers has also declined
steadily since 1969, so these data may not give a true picture of sage grouse
populations for Moffat County.
It is planned to secure counts on as many
active known strutting grounds as possible, check for populations at or near
apparently abandoned grounds, and locate additional new grounds during future
years as an important part of this investigation.
An overlay, classified as sensitive (not available for public viewing) showing
strutting ground locations in the Moffat County area is maintained by the BLM
Office in Craig.
Information for these locations and numbers of sage grouse
is supplied entirely by the DOW, since the BLM does not make independent
surveys on sage grouse.
Several variables. including weather factors, elevations, and changes in
counters, strutting ground locations, dates of counts and time of day of
counts are considered to be factors which bias sage grouse strutting ground
counts.
These are discussed separately.
Weather Factors - Access to most of the strutting grounds at lower elevations
in Moffat County is normally possible by mid-April, following normal or below
normal winters, as measured by amounts of snowfall.
Highest counts of sage
grouse on strutting grounds are often secured during early April.
Following
extreme winters, such as occurred during the first quarter of 1975, access
to even lower elevation strutting grounds may be delayed up to a month.
Strutting activity and numbers of birds normally declines after May 1 so late
counts may not give a true picture of spring populations.
Elevations - Access to strutting grounds at higher elevations such as those in
the Blue Mountain and Cold Springs Mountain areas is normally delayed from
three to four weeks, compared with access to strutting grounds at lower elevations in Moffat County, due to delays of melting snow.
Peak numbers of sage
grouse visiting strutting grounds may be missed when access to these higher
elevational grounds is not possible.
Changes in Counters - There has been numerous
personnel changes in Wildlife
Conservation Officer ranks in Moffat County during the period 1969-1975,
except for the Rangely and Craig-South Districts (Table 1). With this variation in personnel, and differences in interest and counting ability, considerable bias in strutting ground counts may have occurred.
Changes in Ground Locations - Through the years a numer of active strutting
grounds have either moved or been abandoned for various reasons.
While efforts
have been made by Wildlife Conservation Officers to keep the number of active
strutting grounds counted fairly consistent, records show that numbers of
active grounds counted in Moffat County have steadily declined from 29 in 1969
to only 17 in 1975. When a strutting ground becomes inactive, an intensive
effort to locate and count relocated or satellite grounds would undoubtedly
improve the population data collected.

�-127Changes in Dates of Counts - The date that a strutting ground is counted in
relation to the time of peak attendance by sage grouse is known to have a
definite effect on numbers of birds which may be counted.
Following a peak
period of attendance, numbers of birds visiting a strutting ground declines
so that counts made late in the spring may indicate lower populations than
is actually the case.
Changes in Time of Day of Counts - The time that a count is conducted
relation to daylight is known to have a definite effect on the number
(both sexes) which may be counted on a particular strutting ground.
Sage Grouse Observations

and Reported

in
of birds

Sightings

Observations of sage grouse, field signs, and reports secured during January,
February, and March 1976 are listed in Table 3. Locations of at least eight
wintering flocks were identified from these observations and reports.
Three
additional locations are classified as possible wintering flock locations.
Sage Grouse Mortalities
Only two isolated instances of sage grouse mortalities were found during the
period January through March 1976. These are listed in Table 3 and summarized
below.
Scattered feathers from one unclassified sage grouse were found on January 30,
1976 along Moffat County Road 9 at the Thornburgh Gulch Crossing.
These were
from a mortality from the current winter
period and were found in tall,
dense, big sagebrush cover near the draw bottom.
This bird had been eaten by
unknown predators.
From field signs observed in the vicinity it appeared
that a group of sage grouse had spent at least part of the winter period in
this locality, but no live birds or recent field signs were found.
A freshly killed mature male sage grouse, hit by a vehicle, was found on
March 23, 1976 along Moffat County Road 7 northwest of Craig, approximately
1.4 miles southeast of the road junction with Moffat County Road 17. This
location is approximately 2 airline miles southeast of Grassie's Reservoir
strutting ground and 1.6 miles northwest of a site where 13 male sage grouse
were observed on January 30, 1976.
Trends in Sage Grouse Populations
Progress was made in compiling sage grouse strutting ground census data for
Moffat County from Rogers (1964), Wildlife Management Section, and Northwest
Region for the period 1958 through 1975 to determine possible trends in populations.
Data is missing on the number of active strutting grounds counted
for the years 1962-1968 and on the number of strutting ground counts made for
the years 1973-1975 so completed summaries will be included in the Job
Progress Report for next segment.

�Table 3.

Sage grouse

Date

Time

1-7-76

2:00 PM

1-21-76

observations

and reported

sightings.

Moffat

County,

January-March,

No. of Sage Grouse
Males
Females
Unclass.

Location
Moffat Co. Road 19 at Red Wash Crossing
(1~ mi NW of Road 19 strutting ground)

4

11:00 AM

Sand Wash Ridge Road, vicinity of Sand
Wash Ridge Road strutting ground

1-22-76

1:45 PM

Sand Wash Ridge Road, 10 mi N. of Sand
Wash Ridge Road strutting ground

1-22-76

2:15 PM

Sand Wash Ridge Road, 12 mi N. of Sand
Wash Ridge Road strutting ground

1

Vegetative
Cover Type

Observed
By

Remarks

Dense big sagebrushgrassland

D. Hoffman

Flushed this group
of 4 but tracks of
many more in area

3 flocks Dense big sagebrush(smallest 30)grassland

R. Haskins

Reported wintering
flocks

3

Sparse big sagebrushgrassland

D. Hoffman

Tracks only

55 to 60

Dense big sagebrushgrassland with a few
scattered junipers

D. Hoffman

Flushed

Dense big sagebrushgrassland

D. Hoffman

Tracks only

13

Disturbed big sage~
brush-grassland
(probably disked)

D. Hoffman

Flock flushed from
cover to road

40

Dense big sagebrushgrassland

D. Hoffman

Flushed

Several
tracks

Dense big sagebrushgrassland

D. Hoffman

Field sign and
feathers of mortality found

5

Dense big sagebrushgrassland

D. Hoffman

Flushed flocks of
7, 5, and 34

Dense big sagebrushgrassland

D. Hoffman

Flushed

Deception Creek Road from Red Barn to
Price Creek Hill

1-22-76

1976.

24

flock

I
•....

'"

co

1-30-76

8:30 AM

Moffat Co. 7, 3 mi E. of RJ Moffat
with 17

1- 30-76

11:20 AM

Moffat Co. Road 9, 3.2 mi S. of RJ
with Moffat Co. 4

1-30-76

2:00 PM

Moffat Co. Road 9 at Thornburgh
Crossing

1:45 PM

Moffat Co. Road 55, 4 mi. SW of RJ
with Moffat Co. 17

2:30 PM

Moffat Co. Road 55, 2.5 mi. SW of RJ
with Moffat Co. 17

2-19-76

2-19-76

Co.

Gulch

41

13

I

flock

flock

------------------------------------------------------------------~-----------------------------------------------------------------------------

�Table 3.

Date

Sage grouse observations

Time

and reported

sightings,

Moffat

County,

January-March,

No. of Sage Grouse
Males
Females
Unclass.

Location

1976 (continued).

Vegetative
Cover Type

Observed
By

Remarks

3-20-76

10: 30 AM

Lookout Mountain Road, 5~ mi. SW of
RJ with Moffat Co. 4

-

-

80 to 100

Dense big ssgebrushgrassland

D. Hoffman

Mostly males,
flushed birds from
strutting ground

3-23-76

9:15 AM

Moffat Co. Road 7, 1.4 mi SE of RJ
with Moffat Co. 17

1

-

-

Dense big sagebrushgrassland

D. Hoffman

Roadkilled
male

3-23-76

10: 30 AM

Vicinity of Grassie's
strutting ground

Reservoir

-

-

Tracks and
droppings

Dense big sagebrushgrassland

D. Hoffman

Indicates recent
use of ground

3-23-76

11: 30 AM

Vicinity
ground

of Bard Gulch strutting

-

-

4

Dense big ssgebrushgrassland

D. Hoffman

Flushed birds and
field signs show
recent use of
ground

mature

•...
I

N

3-24-76

u.S. Highway
Crossing

3-30-76

Yellowjacket Road, approximately 6
to 8 mi N. Thornburgh Mtn. Monument

40 at Skull Creek

25 to 50

Several birds

Greasewood

wash

Sheepherder
and Highway
Maint. man

Flock observed
January, 1976

W. flcKean

Recent sightings
of birds.

in

,

so

�-130Agricultural Statistics
Annual estimates of winter wheat acreages planted, hay acreages harvested,
numbers of cattle and calves on farms, and nwnbers of stock sheep as of
January 1 each year for Moffat County, adapted from Colorado Agricultural
Statistics are listed in Table 4. These statistics are published annually
by the Colorado Crop and Livestock Reporting Service and were examined for
indicated trends as follows:
Winter Wheat Acreages Planted - The number of acres of winter wheat planted
in Moffat County has fluctuated considerably from 1960 through 1973 (range
25,500 in 1970 to 35,500 acres in 1967) but no clearcut trends are evident
from these data. The estimated number of acres planted to winter wheat in
1973 (30,400) was only slightly higher than that planted in 1960 (31,570).
Preliminary estimates for 1974 were considerably higher (38,400) but final
estimates may be lower, based upon a comparison of final estimates with
preliminary estimates for previous years.
Hay Acreages Harvested - Estimates of acreages of hay harvested in Moffat
County increased from 20,320 in 1960 to 29,300 in 1962, held relatively stable
from 1962 through 1965, dropped considerably during the period 1966 through
1970 (range 17,200 to 20,000), and again increased approximately to 1962
levels during the period 1971 through 1973 (range 27,500 to 31,000).
Number of Cattle and Calves on Farms - Trends shown by numbers of cattle and
calves on farms in Moffat County indicated a steady increase from 1961 (24,170)
to 1974 (37,000), consistent with general trends shown for the entire state
of Colorado. Estimates indicated there were approximately one and one-half
times as many cattle and calves on farms in Moffat County in 1973 and 1974
(37,000 both years), as in 1961 and 1962 (24,170 both years).
Number of Stock Sheep as of January 1 Each Year - Trends shown by numbers
of stock sheep in Moffat County indicated a steady decrease from 1961
(160,400) to 1974 (93,000). Moffat County has ranked first in numbers of
sheep by County for many years and has shown the same decline in numbers
as reported for the entire state of Colorado. The estimate of 93,000
stock sheep on hand as of January 1, 1974 was considerably lower than the
estimate of 160,400 on hand as of January 1, 1961.
Human Populations
Past, present, and future human population estimates for Moffat County and for
its county seat of Craig, Colorado are listed in Table 5. U. S. Bureau of
Census data from the World Almanac (World Telegram and Sun 1965-1976) and
information contained in the Craig Area Comprehensive Plan (Moffat County
Planning Department 1975) were used for these estimates. Human populations
had stabilized at approxiamtely 7,000 for Moffat County, and 4,000 for Craig
during the 1960's. Following the beginning of an energy boom it was estimated
the population of Moffat County had increased to 9,540 by 1975, with 7,227
living in the town of Craig.
Moffat County has long been known to contain vast areas of energy producing
minerals including oil shale, coal, oil and gas, and uranium. With these

�-131-

resources and an increasing demand for energy, future population projections
indicate 16,715 people will be living in Moffat County by 1985, with 13,174
living in the town of Craig. It is anticipated population growth will
continue through 1995 but at a reduced rate. By 1995, 17,320 people are
expected to be living in Moffat County, with 14,275 in Craig.
Vehicular Licenses
Numbers of vehicles licensed and amounts of vehicular traffic on the streets,
roads, and highways of Craig and Moffat County has and will undoubtedly
continue to keep pace with human population growth. Information on numbers
of passenger vehicle and state truck licenses sold in Moffat County from
records maintained in the Moffat County Clerk's office are presented in Table
6 for the years 1973, 1974, and 1975. Numbers of passenger vehicle licenses
sold in Moffat County increased from 3,593 in 1973 to 4,123 in 1975, and
numbers of state truck licenses increased from 799 in 1973 to 2,079 in 1975.
LITERATURE CITED
Canfield, R. H. 1941. Application of the line intercept method in sampling
range vegetation. J. Forestry 39:388-394.
Colorado Crop and Livestock Reporting Service. Yearly 1960-1975.
agricultural statistics. Colo. Dept. of Agric. Denver.

Colorado

Moffat County Planning Department. 1975. Craig area comprehensive plan.
Craig Planning Commission Members. Sept. 58 p.
Phillips, E. A. 1959. Methods of vegetation study.
Winston, Inc. New York. 107 p.
Powell, J.
brush.

Holt, Rinehart, and

1968. Site factor relationships with volatile oils in big sagePh. D. Thesis, Colo. State Univ., Ft. Collins. 82 p.

Rogers, G. E. 1964. Sage grouse investigations in Colorado.
16. Colo. Game, Fish and Parks Dept. Denver. 132 p.

Tech. Pub. No.

Vories, Kimery C. 1974. A vegetation inventory and analysis of the Piceance
Basin and adjacent ranges. M. A. Thesis. Western State College of Colo.,
Gunnison. 243 p.
Weaver, J. E., and F. E. Clements.
Co. New York. 601 p.

1938.

Plant ecology.

McGraw-Hill Book

World Telegram and Sun and Newspaper Enterprises Assoc., Inc.
1965-1976. World Almanac. New York and Cleveland.

m

Wildlife Researcher

Yearly,

�~132-

Table 4.

Agricultural statistics for Moffat County, Colorado, 1960-1974. !/

Year

Winter Wheat
Acreage Planted

Hay Acreage
Harvested

No. of Cattle
and Calves on Farms

No. of Stock
Sheep (Jan. 1)

1960

31,570

20,320

24,660

157,000

1961

30,100

22,200

24,170

160,400

1962

27,500

29,300

24,170

150,000

1963

28,000

28,500

24,600

146,000

1964

30,200

28,500

25,200

141,000

1965

33,000

24,770

27,000

123,000

1966

28,300

17;400

28,000

134,000

1967

35,500

17,700

29,000

127,500

1968

32,450

18,200

30,200

108,000

1969

32,000

20,100

34,000

106,000

1970

25,500

17,200

32,000

104,000

1971

34,500

31,000

34,000

104,000

1972

31,000

28,000

36,000

109,000

1973

30,400

27,500

37,000

97,000

1974 J)

38,400

37,000

93,000

1:/
Source: Colorado Agricultural Statistics, published yearly by Colorado
Crop and Livestock Reporting Service, Denver, Colorado.

Preliminary statistics only.

�Table 5.

Human population statistics for Moffat County and Craig, Co1orado~ 1950-1995.

11

Past Years 1960
1950

Item

1970

Present an dE.Future Years
1985
1980
1975

stlmates- 21
1995
1990

Number of people,
Moffat County, Colo.

5,946

7,061

6,521

9,540

11,820

16,715

16,011

17,320

Number of people,
Craig, Colorado

3,080

3,984

4,205

7,227

9,175

13,174

12,892

14,275

l/source:

World Almanac, using U.S. Bureau of Census information.

1/Source:

Craig Area Comprehensive Plan.

Table 6.

Numbers of vehicular licenses sold, Moffat County, Colorado, 1973-1975. !I

Item
Number of passenger vehicle
licenses sold
Number of state truck licenses
sold
Totals

I

I-'
VJ
VJ

1973

Year
1974

1975

3,593

3,835

4,123

799

1,874

2,079

4,392

5,709

6,202

l/source: Records in County Clerk's Office, Craig, Colorado.
licenses excluded).

I

Includes only new licenses issued (transfer

��April,

-135-

1976

JOB PROGRESS REPORT

State of

COLORADO
----....::.::.::::

9

\'iorkPlan No.
Job Title
Period

Game Bird Survey

W-37-R-29

Proj e c t No.

Population

Covered:

Job No.
Dynamics

and Habitat

5

Relationships

of Blue Grouse

April 1, 1975 to March 31, 1976

Personnel:
H. Alexander, A. Anderson, D. Benson, C. Braun, G. Brown, L.
Carpenter, R. Clippinger, M. Coghill, J. Corey, C. Crawford, D. Freddy, H. Funk,
D. Gore, L: Green, J. Gumber, D. Hoart, J. Hobbs, R. Hoffman, J. Jackson, J. E.
Kautz, R. Lowry, F. Marcoux, B. McCloskey, S. McElderry, S. McEllin, K. Miller,
R. Oakleaf, S. Palm, S. Porter, S. Quinlan, G. Smith, M. Smith, H. Spear, S.
Steinert, D. Todd, and J. Wagner.
ABSTRACT
Investigations of the population dynamics and habitat relationships of blue
grouse (Dendragapus obscurus) were initiated in Colorado in 1975. Work during
the first year was primarily devoted to (1) reviewing literature, (2) selecting
study locations, (3) developing and testing new and existing methods for censusing, capturing, and marking blue grouse, and (4) collecting harvest data.
The Green Mountain-Blue Ridge area in Middle Park was selected for intensive
study, while three other areas (Vail Pass, Eby Creek, and Deep Creek) are
tentative selections pending further investigation.
Only one of the three areas
will be selected.
Acoustical census was used to locate territorial males and
some females during the breeding season.
Systematic search with the aid of a
pointing dog was the most successful method for finding breeding hens, females
with chicks, and non-territorial males.
Limited success was obtained in using
chick distress calls to locate females with chicks.
Most attempts to capture
blue grouse with a noose pole or mist net were unsuccessful because of their
wildness.
Captured grouse were individually identified with color combinations
of anodized, aluminum, butt-end leg bands.
Check stations were operated at
various locations throughout the state in order to collect information on
hunter pressure, success, and efficiency, and to collect wings from hunter
harvested birds.
Volunteer wing collection stations were devised and utilized
to increase samples of wings collected .. Analysis of the wings provided data
on age and sex composition of the harvest, nesting success, hatching dates, and
production and survival rates.
Collected wings were also used for evaluating
and testing new and existing methods of aging and sexing blue grouse.

�-136-

RECOMMENDATIONS
1.

Continued efforts should be made in 1976 to develop successful
methods for capturing blue grouse.

2.

Further

3.

An experimental season for blue grouse should be held during the big
game season in selected management units.
Legal weapons should be
.22 caliber rimfire rifles and hand guns, and standard shotguns.
Check
stations should be operated on weekends with field checks being conducted
throughout the season in order to evaluate the experimental season.

testing of census techniques

should be conducted

in 1976.

�-137-

HABITAT

POPULATION DYNAMICS AND
RELATIONSHIPS OF BLUE GROUSE
Richard W. Hoffman

Sound management of game birds requires considerable knowledge concerning
the biology of each species and factors affecting their populations.
Blue
grouse, a relatively unexplored species in Colorado. have been hunted in
modern times in the state since the early 1950's. yet data upon which to
base management recommendations have been and still are inadequate.
Lacking
data, season lengths and possibly bag limits have been conservative and
based on impressions of field workers and administrators.
Blue grous.e occur in varying densities over more than 51,800 sq km (20, 000
sq mi) of diverse habitat and terrain in Colorado (Rogers 1968) and, in
terms of hunter harvest (18,000 average annual harvest), are second only to
the ring-necked pheasant (Phasianus colchicus) among resident upland game
birds.
Effects, if aDy, of present hunting seasons on blue grouse populations are not known, but it is doubtful if hunters take 5 percent of the fall
grouse population.
Data from other western states suggest that 25 to 30
percent of the fall population could be harvested with no adverse effects on
subsequent breeding populations (Zwickel 1958).
It appears this level has
not been attained anywhere with present hunting methods regardless of bag
limit and season length.
Before increased hunter opportunity and recreation
on this species can be justified in Colorado, basic data on population
levels, habitat relationships, and effects of present hunting seasons
must be obtained.

P. S. OBJECTIVE
Major objectives of this study are to (1) increase the harvest of blue grouse
in Colorado (double present harvest estimates) without harm to breeding
populations in subsequent years, (2) to identify differences in breeding
densities due to differing habitats, and (3) to document the stability of
breeding densities over time.

SEGMENT OBJECTIVES
1.

Review literature concerning (a) techniques for marking, capturing, and
censusing grouse, (b) methods of aging and sexing blue grouse, (c) population dynamics, behavior, and habitat requirements of blue grouse, and
(d) effects of hunting on grouse populations.

2.

Evaluate and test established marking, capturing and census procedures
used in other states and provinces and develop suitable methods for
breeding and production surveys in Colorado.

3.

Select study areas in two different habitat types.
Candidate
under consideration include the mixed conifer-aspen-sagebrush
the aspen-oakbrush-spruce-fir
type.

4.

Trap and individually mark 50 birds (including adults,
juveniles) on each of the selected study areas.

habitats
type and

subadults

and

�-1385.

Compile data, analyze

results,

LITERATURE

and prepare progress

report.

REVIEW

Although the literature abounds with information on blue grouse, primarily
resulting from studies conducted in Montana, Washington, British Columbia,
and Alberta, only limited research has been conducted on the particular
subspecies (~. ~. obscurus) inhabiting Colorado.
At present, the best
account of the ecology of this subspecies is from investigations in Wyoming
(Harju 1974). Except for work by Steinhoff (1956) and Rogers (1968), published information on blue grouse in Colorado is confined to short accounts
relative to occurrence, an observation or kill, a single crop.content, or a
nest seen and described.
Some additional data have been presented by Braun
(1971) on age and sex determination of blue grouse and by King and Braun
(1970) on fall food habits of blue grouse in subalpine areas of Colorado.
A more comprehensive literature review relevant to the topics discussed is
incorporated into the text of this report.

METHODS AND MATERIALS
The initial year of investigation was primarily devoted to selecting study
areas and developing and testing new or existing techniques for censusing,
capturing, and marking blue grouse.
Methods and materials pertaining to
these objectives are discussed in detail under Results and are only briefly
mentioned under Methods and Materials.
Grouse were located by systematic search with the aid of a pointing dog and
by acoustical census.
During acoustical census, a portable, lightweight,
battery powered Norelco Model 150 Carry Corder utilizing cassettes was used
to play back tape recorded calls of blue grouse.
Once birds were located
they were first observed through 7X50 binoculars to count numbers present
and ascertain sex. Attempts were made to capture grouse through use of
noose poles or mist nets whenever possible.
All grouse captured were weighed,
aged, sexed, and banded.
Scales accurate to within ± 5 grams were used to
weigh
grouse less than 1000 grams; grouse exceeding 1000 grams were weighed
with a scale accurate to within ± 10 grams.
Color combinations of anodized,
aluminum, butt-end, leg bands (size 14) were used to individually mark grouse.
Methods of determining sex and age of blue grouse have been described
(Caswell 1954; Bendell 1955; Smith and Buss 1963; Mussehl and Leik 1963; Boag
1965; Zwickel and Lance 1966; Schladweiler et ale 1970; and Braun 1971) with
most studies, except Braun (1971), being described for and tested on other
races of blue grouse.
The limited data collected in Colorado by Braun (1971)
on age and sex determination of blue grouse based on examination of wing
plumage characteristics
and primary replacement parallel that reported by
Mussehl and Leik (1963) and Schladweiler et ale (1970).
Check stations were operated in Middle Park, North Park (in conjunction with
sage grouse (Centrocercus urophasianus) check stations), Crown Point, and
Coffee Pot Springs during the 1975 blue grouse season.
Stations in Middle
Park and Coffee Pot Springs were operated the first two weekends of the 23 day
season (September 13 to October 5). The Crown Point station was operated

�-139-

opening weekend and stations in North Park were operated opening weekend
and the second Sunday.
Ten volunteer wing collection stations were constructed and tested for use in Middle Park during the blue grouse season
(Hoffman and Braun 1975).

RESULTS AND DISCUSSION

Selection

of Study Areas

Potential study areas were located by searching for concentrations of grouse.
To assist in this endeavor, a blue grouse observation form was mailed to all
Division of Wildlife employees working in mountainous districts (Appendix).
Useful information was also obtained by personal communication with Division
personnel and landowners.
Areas where breeding grouse and females accompanied
by chicks were consistently observed were investigated to ascertain (1) density of grouse utilizing the area, (2) variations in slope, aspect, cover
density, and vegetative composition within the area, (3) accessibility,
(4)
availability to hunters, and (5) hunter use.
In all, 14 areas were investigated as potential study locations.
One area
was selected for intensive study in 1975 while three others were tentatively
identified pending further investigation in 1976. Only one of the three
tentative areas will be selected.
All areas are west of the Continental
Divide and include: Green Mountain-Blue Ridge area in Middle Park, Eby Creek
north of Eagle, Deep Creek northwest of Dotsero, and the east side of Vail
Pass north of 1-70. Green Mountain-Blue Ridge represents the initial area
selected for intensive study.
The Green Mountain-Blue Ridge area is approximately 19 kilometers (12 mi)
south of Kremmling in Grand and Summit counties.
Area investigated is in
T2S, R79W, parts of sections 4,5,6,7,8,9,17,
and 18; and T2S, R80W, parts of
sections 3,10,11,15, and 14. While small portions of the area are under
private ownership or are natural resource lands administered by the Bureau
of Land Management, U. S. Department of Interior, the majority of the area is
national forest lands administered by the Kremmling District, Arapaho National
Forest, U. S. Forest Service, Department of Agriculture.
Initially, only Green Mountain (2.6 sq km or 1 sq mi) was selected for intensive
study. However, due to (1) the limited size of the area, (2) low hunting
pressure, and (3) early departure of grouse from Green Mountain, the study area
was expanded to include about 5 square kilometers (2 sq mi) of the adjacent
Blue Ridge area.
Exact study boundaries have not been established, but
breeding season investigations will be confined to Green Mountain with production surveys including both Green Mountain and Blue Ridge.
Approximately 4 kilometers (2.5 miles) of an open, sagebrush (Artemisia spp.)
dominated flat separates Green Mountain from Blue Ridge.
The study area lies
in the southwestern part of Middle Park, a large topographic depression that
is bounded on the south and east by the Front Range, on the north by the Rabbit
Ears Range, and on the west by the Park Range.
Unlike other intermountain
basins or parks (South and North Parks) that mainly have broad rolling grass

�-140covered gentle floors, Middle Park is mount:ainous and locally heavily
forested.
At the study location, coniferous forest predominates above
2700 meters (8800 ft) and extends downward to 2600 meters (8500 ft) on
Green Mountain.
Unforested areas below this elevation are vegetated predominantly with big sagebrush (Artemisia tridentata).
Intermediate areas
are best described as a mixed conifer, aspen (Populus tremuloides), sagebrush, serviceberry (Amelanchier alnifolia) type.
The coniferous coverage
includes Douglas fir (Pseudotsuga menziesii), Engelmann spruce (Picea
engelmannii), lodgepole pine (Pinus contorta), and a few scattered limber
pines (Pinus flexilis), alpine firs (Abies lasiocarpa), and Rocky Mountain
junipers (Juniperus scopulorum).
Thick stands of willows (Salix spp.)
predominate along water courses.
Topography of the Green Mountain-Blue Ridge area is irregular varying from
steep slopes and ridges to gently rolling expanses hundreds of hectares in
size.
Cliffs and ledges are common in the area as are small isolated knolls.
Maximum relief is about 900 meters (3000 ft); elevations range from 2400
meters (7850 f t ) to 3300 meters (10800 f t ) above sea level.
Geology of the
lower Blue River area which includes portions of the study area has been
described by Holt (1961).
Taggart (1962) presents a detailed discussion
of the geology of the entire Mt. Powell quadrangle.
Blue Ridge is drained by many small tributaries that flow west into the
north:-flowing Blue River.
Approximately 19 kilometers (12 mi) north of the
study area, the Blue River empties into the Colorado River.
No well delineated streams occur on Green Mountain where most of the runoff drains
directly into the Blue River.
While the majority of streams on Blue Ridge
maintain a year-round water flow, a similar situation does not exist on
Green Mountain where most streams are essentially dry by late June or early
July.
Climate of the area is typically continental, with frequent extremes in
wind velocity and temperature.
Prevailing winds are westerly.
Weather
data compiled at Frazer, Colorado, in the southeast portion of Middle Park,
show that the annual precipitation averages about 53 centimeters (21 in)
and occurs mainly in April and July. Precipitation from late October to early
April is usually in the form of snow or sleet, turning to rain in late April.
0
Average winter temperature is around l3 F and average summer temperature
o
about 51 F.
Once a second study area is selected, a thorough literature review will be
conducted and incorporated into a detailed description of the geology, soils,
vegetation, climate, topography and drainage patterns of the study areas.
This information will be supplemented with data collected during field investigations which will include intensive analysis of the vegetation at both study
locations.

Census

Breeding

Season

The most common breeding census method for blue grouse presently in use is the
acoustical census.
Acoustical census makes use of blue grouse behavior and

�-141-

involves the play back of a tape recorded hen whinney (precopulatory call)
which stimulates territorial males to respond by hooting (Stirling and
Bendell1966).
Hooting counts are used in California, Washington, Oregon,
and British Columbia where races of the sooty grouse occur (Rogers 1968).
Behaviorial differences between the sooty and dusky races, primarily relating to hooting volume, precludes the use of hooting counts in Colorado
or Wyoming (Rogers 1968; Harju 1974).
Hooting is the main long-range advertising display of sooty grouse, whereas,
dusky grouse perform elaborate flutter-flight displays to attract females
and proclaim territories.
Female dusky grouse in turn utter a loud cackling
call to advertise their presence to males.
By using tape recorded hen
cackle calls, Harju (1974) was able to elicit flutter-flights
from territorial
males.
Additionally, some females responded to the call by cackling, indicating perhaps the cackling of one hen stimulates nearby hens to respond
or else the cackling has some communicative function between hens.
Flutterflights of one male frequently initiate a chain reaction response from
surrounding males.
Ninety two males were located during the 1975 breeding season.
Seventy
seven percent (71) of the males located initially responded to the tape
recorded hen cackle.
Wing-flutters were the dominant response accounting for
56 of the 71 (79%) responses.
Only 15 males (21%) responded by hooting;
however, the 21 males failing to respond may have been hooting but were not
heard.
Seldom was the initial response in the form of hooting unless the
call was played within 50 meters (164 ft) of the territorial male.
Males
that initially wing-fluttered
could also be induced to hoot by approaching
close and playing the tape recorded precopulatory call.
Systematic search of the habitat with the aid of a pointing dog was the most
successful method for finding hens during the breeding season, but use of the
hen cackle call greatly increased numbers of females found. Thirty six percent of the 33 hens located initially responded to the call. Due to their
more secretive habits and lower response rate, fewer hens (33) than males (92)
were observed.
These two factors are probably responsible for the distorted
sex ratio (2.8 males per female) indicated by breeding season observations.
Use of the hen cackle call was most effective in the early morning and late
evening hours, but some males and females could be stimulated to respond
throughout the mid-day hours.
Distance at which the wing-flutter or cackling
could be heard varied from less than 100 meters (328 ft) to over 500 meters
(1640 ft) depending upon wind velocity.
Males responded to the call from
early April to mid-June with the frequency and intensity of responses reaching
a peak in mid-May.
Responses by both males and females declined rapidly in
June when most hens should have initiated nesting.

Brood Season
Both systematic search and acoustical census were employed to locate females
accompanied by chicks.
Acoustical census involved the play back of tape
recorded chick distress calls.
Of 16 hens found that were accompanied by
chicks, 4 (25%) responded to the chick distress call by uttering loud alarm
calls (3 females) or moving towards the call in an aggressive posture (1 female),
9 (56%) were located by systematic search using the dog, and 3 (19%) were

�-142found by systematic search without the dog. The 4 .hens obviously responding all had chicks less than 4 weeks of age. Apparently, hens with
older broods are less protective of their young and less likely to alarm
call in response to recorded sounds (Harju 1974). No unsuccessful females
were known to respond to the call.

Capture

Noose Pole
Successful methods for capturing blue grouse have been described including
noose poles (Zwickel and Bendell 1967), mist nets (Schladweiler and Mussehl
1969), drive traps (Tomlinson 1963), and funnel traps (Henderson 1960).
The
most effective and practical method is the noose pole.
The noosing apparatus
is a 7 meter (22 ft), telescoping, fiberlgass, fishing pole. Nylon covered,
stainless steel, leader wire is used to make the nooses.
Different strengths
of wire are used depending upon the size of the grouse to be captured.
Harju (1974) reported using a noose pole to successfully capture dusky grouse
in Wyoming.
Most attempts to capture blue grouse in Colorado with a noose
pole were futile.
Lack of experience on part of the person trying to capture
the birds may have been partially responsible for the poor success, but many
of the capture attempts failed because the birds were too wild to approach
within noosing distance.
Less than 2 percent of the 212 birds observed were
captured.
Usually the grouse would flush when approached within 12 meters
(40 ft) and would subsequently land in a tree out of reach of the noose pole.
Others would fly several hundred meters downhill and could not be relocated.

Mist Net
Some attempts were made to capture birds with a 4 ply, 10 cm, black, nylon
mist net measuring approximately 3 meters (10 ft) x 18 meters (60 ft). The
net was positioned 10 (33 ft) to 30 meters (98 ft) below the grouse.
Birds
were then approached from the uphill side and maneuvered towards the net.
Just before reaching the net, grouse were flushed.
Since only one man was
involved in this operation, considerable time was required to set up the net.
Oftentimes the bird would move away or flush while the net was being positioned
or else one man was not sufficient for moving the bird towards the net.
Consequently, all attempts to capture blue grouse with the mist net failed.

Territorial

Trap

A modified version of the trap developed by Bray et ale (1975) for capturing
territorial male red-winged blackbirds (Agelaius phoeniceus) was constructed
for similar use on territorial male blue grouse.
Unfortunately, the trap was
not completed until after the cessation of breeding activities in 1975 so
its usefulness will not be tested until the 1976 breeding season.
The
principle of this trap takes advantage of the behavior territorial males exhibit towards females that enter their territories.
When the trap, with a
mounted female (precopulatory position) in the center compartment, is placed
in a territory, the male should enter the trap in an effort to display to
and copulate with the female.
Tape recorded precopulatory calls will be used
to further 'entice the male into the trap.

�-143Marking
Captured grouse older than 6 weeks were individually marked following the
banding scheme described by Gullion (1965).
Eight colors of anodized,
aluminum bands are used allowing for more than 4000 birds to be individually
marked without repeating a combination by placing two colored bands on each
leg. The eight colors include green, red, copper, blue, orange, yellow,
lavender, and silver.
In this system, each of the 4 bands placed on a particular bird carries the same number which identifies that bird for record
keeping purposes.
Thus, any band which might be found among scattered bird
remains, or turned in by a hunter, can be referred to the proper bird; the
bird can also be identified in the field by "reading" the color combination.
The band on the lower right leg is called the base band.
All birds banded
in the same year have the same color base band.
The band opposite the base
band (upper left leg) is called the alternate band and is used to identify
the general locality where the bird was initially banded.
Anodized colors show little tendency to fade and the eight colors chosen
are the easiest to distinguish (personal communication, G. Gullion).
Similar anodized bands placed on ruffed grouse (Bonasa umbellus) were still
brightly colored 60 months after banding (Gullion 1965).
Since it is unlikely that a blue grouse will be recaptured, the lasting nature of anodized
bands is particularly useful for marking this species.

Harvest
Although harvest data may not exactly reflect the structure of the population
from which they came (Hickey 1955), nor does hunter success provide criterion
for determining grouse population levels (Zwickel 1958), this information is
useful for comparisons among areas and years when collected over an extended
period of time and in conjunction with intensive population studies.
During
the 1975 blue grouse season, check stations were operated in the Middle Park,
North Park, Crown Point, and Eagle areas.
Ten volunteer wing collection
stations were also available to hunters at various locations throughout
Middle Park. The primary objectives of check stations were to estimate hunter
pressure, success and efficiency, and to collect wings from hunter-harvested
birds.
From analyses of these wings, information was acquired on age and sex
composition of the harvest, nesting success of breeding females, hatching
dates of juveniles, and production and survival rates.
Age and sex composition of the harvest sample from each area are shown in Table 1.
Small samples from North Park, Crown Point, and the Eagle area prevent comparisons among areas.
Conclusions could not be formulated because of the
limited amount of data collected.
However, due to use of wing collection
stations, adequate samples of hunter-harvested
birds were checked in Middle
Park to allow analysis.
It was concluded that because of low hunting pressure,
scattered distribution of hunters, and extensive occupied range of blue grouse,
use of manned check stations was not economically feasible and was inefficient
for collecting sufficient samples of wings to satisfy management requirements.
Conversely, the volunteer wing collection stations were instrumental in inexpensively increasing the collection of harvest data (Hoffman and Braun 1975).
Volunteer wing collection stations accounted for 66 percent (120) of the 183
wings collected in Middle Park and 45 percent of the total wings (270)
obtained from all areas.

�-144Table 1. Age and sex composition
checked during the 1975 season.

Adults
Males
Females

Area

Middle

of hunter harvested

II

Subadults Males
Females

blue grouse

Juveniles
Males
Females

Unk.

Totals

Park

30

41

5

15

36

43

13

183

North Park

11

18

5

3

5

9

6

57

Eagle Area

6

8

0

0

4

3

2

23

Crown Point

0

1

1

3

0

2

0

7

Totals

47

68

11

21

45

57

21

270

llBecause some subadults had probably completed their primary molt and thus
could not be distinguished from adults, number of birds assigned to the subadult category was minimal.

Assuming subadults molt similarly to adults, approximately 40 percent of the
sub adult males and only 6 percent of the subadult females should have completed
their primary molt by the end of the hunting season and thus could not be
distinguished
from adults.
Consequently, number of birds assigned to the subadult category was minimal.
Males initiate their primary molt earlier than
most females so more subadult males than females are indistinguishable
from
adults in the fall. This is reflected in the harvest data from Middle Park
which shows males comprised 42 and only 25 percent of the adult and subadult
harvest, respectively.
Since some subadults were assigned to the adult category, both age classes are grouped together as adults for discussion purposes.
Adults (9V [includes subadults] and juveniles (92) made up equal portions of
the harvest sample from Middle Park, but a major difference occurred in the
proportion
of males harvested from each age class.
Based on 91 adult and 79
juvenile wings for which sex was determined, males comprised 46 percent (36)
of the juvenile harvest and only 38 percent (35) of the adult harvest.
Thus,
the sex ratio of juveniles did not deviate significantly from 1:1, whereas,
the adult segment of the harvest showed a deficiency of males.
The unbalanced sex ratio indicated for adults was not considered indicative of the
population but was probably caused by differences in habitat selection between
sexes at the time of the mid-September to early October season.
The lack of
adult cocks in the harvest is due to their early departure from the more
accessible lowlands to higher, more inaccessible areas. As a result, the bulk
of the harvest (81% for Middle Park) in 1975 was taken from the hen and young
segment of the population.
If this situation occurs normally, and sex ratios
of adults are evenly balanced as indicated for other blue grouse populations
where sufficient data are available (Bendell et al. 1972 and Swickel 1972),
then sex ratios of adults as indicated from hunter harvests warrant extreme
caution in their interpretation.

�-145Blue grouse hunting is mainly limited to range adjacent to roads or other
easily accessible locations with many areas receiving virtually no hunting
pressure.
After opening weekend, hunting pressure decreased markedly.
In
Middle Park, 99 hunters were contacted opening weekend compared to only 13
hunters located the following weekend.
Of 158 blue grouse observed by the
112 hunters, 45 were harvested for a hunter success (birds bagged per
hunter) and hunter efficiency (birds bagged per birds observed X 100) of
.4 blue grouse/hunter and 28.4 percent, respectively.
Two factors tending
to lower hunter success and efficiency were (1) many hunters were unfamiliar
with the regulations, and (2) they could not distinguish blue grouse from
sage grouse.
Consequently, they tended to harvest birds in accordance with
the more conservative 2-bird bag and possession limit for sage grouse instead of the 3 bird bag and 6 bird possession limit for blue grouse.
Using knowledge gained from intensive blue grouse investigations conducted
in Montana (Mussehl 1960), Washington (Zwickel 1958), British Columbia
(Bendell and Elliott 1967), Wyoming (Harju 1974), and the preliminary studies
in Colorado (Rogers 1968), the general consensus is that blue grouse are not
being adequately harvested for maximum utilization and seasons could be
liberalized until some effect of hunting can be demonstrated on the population.
In order to gather data on the potential impact of hunting blue grouse during
the big game season and to provide support for an extended season, a question
concerning blue grouse was included on the Middle Park deer hunter questionnaire.
The question simply stated "How many blue grouse did you see while hunting deer in
Middle Park".
Eighteen hundred and eight questionnaires were returned of
which 1459 (81%) hunters indicated they had participated in the Middle Park
deer season.
Of the 1459 hunters, only 356 (24%) reported seeing blue grouse
for a total of 1904 observations.
Projecting the estimates to the total deer
hunter population (4991), approximately 970 hunters should have seen a total
of about 5200 blue grouse.
Based on the 28.4 percent hunter efficiency calculated from grouse hunter check stations operated in Middle Park in 1975,
of the 5200 blue grouse observed potentially about 1500 would have been harvested.
However, many hunters probably would not take grouse during the big
game season for fear of spoiling their chances of taking a deer or elk. Even
so, a harvest of 1500 birds spread over an area the size of Middle Park
(5180 sq km [2000 sq mil) would have negligible effects on the grouse population (.29 grouse harvested per sq km, or .75 per sq mi). Obviously there are
drawbacks inherently associated with making such estimates, but their effect
on the overall estimate were probably minimal.
While reliability of the data
are questionable, a rough idea was obtained from a management and biological
standpoint on the feasibility of extending the blue grouse season.

Age and Sex Determination
Accurate methods of sex and age classification are requisite to intensive
popUlation studies and analysis of harvest data.
Sex and age criterion have
been described for Colorado blue grouse (Braun 1971).
Most birds can be sexed
by wing coloration pattern alone, but this technique is more reliable for
adults than juveniles and a supplemental means of determining sex of juveniles
has been needed.
Major age classes (adults, subadults, and juveniles) can be
distinguished by the shape and wear of primaries 9 and 10 (primaries consecutively numbered from proximal to distal) (Petrides 1942).
More refined
techniques for aging juveniles have been described (See Methods) and applied

�-146to other races of blue grouse.
Two of these techniques (Zwickel and Lance
1966 and Schladweiler et al. 1970) were tested for use in Colorado along
with developing an additional method of accurately ascertaining sex of
juveniles.
Supplemental data for evaluating existing age and sex determination methods
were obtained in this study by collecting wings from hunter-harvested
birds.
To substantiate plumage data, sex was determined by gonadal inspection whenever possible.
Plumage characteristics
of each wing were examined and
measurements were taken of all primaries present.
Differences in the length
of fully grown-in primaries were tested for use in sexing juveniles.
The
length of actively growing primaries was the basis for estimating age (in
weeks) of juveniles (Zwickel and Lance 1966; Schladweiler et al. 1970).
One hundred and three wings of hunter-harvested
juveniles (46 males and 57
females) w·ere aged using the tables presented by Zwickel and Lance (1966)
and Schladweiler et al. (1970).
Peak of hatch was then determined by backdating.
Since portions of the aging key presented by Schladweiler et al.
(1970) employs actual primary measurements,
it was easier to apply than
Zwickel and Lance's system which required estimates of the proportion of
growth of post juvenile primaries.
Zwickel and Lance's method almost invariably resulted in a higher estimate of age than indicated by Schladweiler
et al. (1970) for the same bird.
The difference did not vary more than 2
to 5 days and was attributed to the greater precision incorporated into the
aging key presented by Zwickel and Lance (1966).
One interesting factor which arose was the difference in peak of hatch
calculated separately for males and females; thus, creating doubt concerning
the reliability of the aging keys.
The peak of hatch calculated according
to Zwickel and Lance (1966) showed 95 percent of the females and only 67
percent of the males hatching between July 1 to 14. Proportionally more
males (28%) than females (5%) were indicated to have hatched after July 14.
Similar calculations using Schladweiler et al. (1970) gave the following
almost identical results: (1) 93 percent of the females and 63 percent of the
males hatched between July 1 to 14, and (2) 7 and 33 percent of the females
and males, respectively, hatched after July 14.
Perhaps the molt sequence and rate of growth of primary feathers is slower
for males than females as is the case with sage grouse (Pyrah 1963).
If so,
using the same key for both sexes would tend to underestimate the age of males
as suggested by the data.
Furthermore, definite differences in the length
of primaries exists between males and females (Hoffman, unpubl. data) which
should further necessitate a distinction between the sexes before estimating
their age. Neither Zwickel and Lance (1966) nor Schladweiler et al. (1970)
mentioned any substantial differences in the aging criterion for juvenile
males and females.
From the limited data currently available, the two
techniques are very similar in the results produced, but until more data can
be collected from juveniles of known age and sex, their applicability to
Colorado blue grouse remains questionable.
Differences in the length of individual primaries between males and females
were used as the basis for developing an additional sex criterion for
juveniles.
Sexes were separated according to wing coloration pattern (Braun
1971) and measurements were taken of all full-in primaries (no blood quill).
Although samples are small, the data presented in Table 2 show a marked

�-147difference in length of corresponding primaries for males and females,
especially P4 through 10 (excluding P7). Due to the fact that all birds
included in the sample were either in the process of molting P7 or else
P7 was not completely developed, no measurements were obtained for P7.
It should also be noted that PI through 6 are post juvenile feathers
while P8, 9 and 10 are juvenile feathers.

Table 2.
Primary1/
Number -

Primary measurement data for juvenile blue grouse.
No. Primaries
Measured

Ave. Length of
Completed Primary (mm)

Range in Length of
Completed Primary (mm)

Males
1

45

117

109-124

2

45

130

121-137

3

46

139

131-146

4

42

152

146-162

5

26

170

165-177

6

9

174

170-180

7

0

8

26

149

140-164

9

22

159

154-166

10

21

126

120-133

Females
1

54

109

100-116

2

53

120

111-127

3

54

126

117-134

4

52

138

128-148

5

38

154

149-163

6

13

159

154-165

8

24

137

129-141

9

42

144

135-152

10

40

114

106-120

7

1/
PI through 7 are post juvenile feathers and P8 through 10 are juvenile
feathers.

�-148When the entire sample of measurements for P4 through 10 are compared,
only 11 of 209 measurements (5%) for females fall within the range of
measurements for males. No overlap was recorded for P5, 6, and 9, and
only 2 (1%), 5(2%), and 4 (2%) overlapping measurements occurred for P4,
8, and 10, respectively. Also, just one juvenile female out of 57 had
more than one primary (P4 and 8) falling within the range of measurements for juvenile males. More data should be collected and the technique
must be tested using known sex birds (gonadal inspection). Refinement
of the technique and its subsequent use with other indicators of sex
(wing coloration) should greatly increase the accuracy in determining sex
of juveniles.

Nesting Success and Production
Successful hens have a delayed primary molt, usually retaining 2 or 3 old
primaries into late September, and can be distinguished from unsuccessful
hens which retain none or only one old primary into September (Braun 1971).
Examination of wing molts of 56 hens harvested in Middle Park revealed that
36 (64%) were successful nesters in 1975. Field observations indicated
somewhat dissimilar results as 78 percent of the females observed in Middle
Park were successful. However, hens with broods are easier to locate than
those without, and this factor results in overestimation of nesting success.
There were approximately 1.6 chicks per female and 2.6 chicks per successful female in the harvest sample. These data and the high percentage (50%)
of immatures in the harvest suggest that nesting success and production
were good in Middle Park in 1975.
Adult and subadult females differed greatly in their ability to bring off a
brood. Nesting success as determined for 41 adult and 15 subadult hens was
76 and 27 percent, respectively. From data collected in another study
(Bendell and Elliott 1967), the poor success of subadults can be attributed
to their lack of experience rather than their failure to mate. The potential
implication of this behavorial difference is that in a population which
sustains a heavy harvest of adult females,their subsequent replacement in the
breeding population by juveniles and subadults should result in lower production.

LITERATURE CITED
Bendell, J. F. 1955. Age, molt and weight characteristics of blue grouse.
Condor 57(6):354-361.
Bendell, J. F., D. G. King, and D. H. Mossop. 1972. Removal and repopulation
of blue grouse in a declining population. J. Wildl. Manage. 36(4):11531165.
Bendell, J. F., and P. W. Elliott. 1967. Behavior and the regulation of
numbers in blue grouse. Can. Wildl. Servo Rep. Ser. 4. 76 p.
Boag, D. A. 1965. Indicators of sex, age, and breeding phenology in blue
grouse. J. Wildl. Manage. 29(1):103-108.

�-149Braun, C. E. 1971. Determination of blue grouse sex and age from wing
characteristics.
Colo. Div. Wildl. Game Information Leaflet No.
86. 4 p.
Bray, O. E., J. L. Guarino, and W. C. Royall, Jr. 1975. A trap for
capturing territorial male red-winged blackbirds.
Western Bird
Bander 50(1):4-7.
Caswell, E. B. 1954. A preliminary study on the life history and ecology
of the blue grouse in west-central Idaho. M.S. Thesis.
Univ. Idaho,
Moscow.
105 p.
Gullion, G. W. 1965.
Improvements in methods for trapping
ruffed grouse.
J. Wildl. Manage. 29(1) :109-116.
Harju, H. J.
grouse.

and marking

1974. An analysis of some aspects of the ecology
Ph.D. Thesis.
Univ. Wyoming, Laramie.
142 p.

Henderson, V. B. 1960.
central Washington.
96 p.

of dusky

A study of the blue grouse on summer range, northM.S. Thesis.
Washington State Univ., Pullman.

Hickey, J. J. 1955. Some American population research on gallinaceous
birds.
Pp. 326-396 in A. Wolfson, ed. Recent studies in avian biology.
Univ. Illinois Press, Urbana.
Hoffman,R.
W., and C. E. Braun.
1975. A volunteer wing collection
Colo. Div. Wildl. Game Information Leaflet No. 101. 3 p.

station.

Holt, H. E. 1961. Geology of the lower Blue River area, Summit and Grand
counties, Colorado.
Ph.D. Thesis.
Colorado Univ., Boulder.
107 p.
King, J. S., and C. E. Braun.
1970.
Fall foods of subalpine populations
of blue grouse in Colorado.
J. Colo. Wyo. Acad. Sci. 6(2):52.
Mussehl, T. W. 1960. Blue grouse production, movements, and populations
the Bridger Mountains, Montana.
J. Wildl. Manage. 24(1):60-68.
Mussehl, T. W., and T. H. Leik.
1963.
J. Wildl. Manage. 27(1):102-106.
Petrides, G. A. 1942. Age determination
Trans. N. A. Wildl. Conf. 7:308-328.

Sexing wings

in

of adult blue grouse.

in American

gallinaceous

birds.

Pyrah, D. B. 1963. Sage grouse investigations.
Idaho Fish and Game Dept.
Wildl. Restoration Div. Job Compl. Rept.
Proj. W-125-R.
71 p.
Rogers, G. E. 1968. The blue grouse in Colorado.
Parks.
Tech. Publ. No. 21. 63 p.

Colo. Div. Game, Fish and

Schladweiler, P., and T. W. Mussehl.
1969. Use of mist nets for recapturing
radio-equipped blue grouse.
J. Wildl. Manage. 33(2):443-444.

�-150Schladweiler,
P., T. W. Musseh1, and R. J. Greene.
1970.
Age determination
of juvenile blue grouse by primary development.
J. Wild1. Manage.
37(3) :649-652.
Smith, N. D., and 1. O. Buss.
1963.
tions of blue grouse.
J. Wild1.

Age determination
and plumage
Manage. 27(4):566-578.

observa-

Steinhoff, H. W. 1956.
The dusky grouse and its ecology in Colorado.
Ph.D. Thesis.
Syracuse Univ., Syracuse, New York.
173 p.
Stirling, I., and J. F. Bendell.
1966.
Census of blue grouse with recorded
calls of a female.
J. Wild1. Manage. 30(1):184-187.
Taggart, J. N. 1962.
Geology of the Mount Powell quadrangle,
Ph.D. Thesis.
Harvard Univ., Boston, Mass.
239 p.
Tomlinson, R. E. 1963. A method
Manage. 27(4):563-566.
Zwickel, F. C.
Game Bull.

for drive-trapping

1958.
North-central
10(4):3-4.

Washington

dusky grouse.

grouse studies.

Zwickel, F. C.
1972.
Removal and repopulation of blue grouse
population.
J. Wildl. Manage. 36(4):1141-1152.
Zwickel, F. C., and J. F. Bendell.
1967.
J. Wildl. Manage. 31(1):202-204.

A snare

Zwickel, F. C., and A. N. Lance.
1966.
Determining
grouse.
J. Wildl. Manage. 30(4):712-717.

Prepared

by ~bb~~~~--~~~~~
Richard W. Hoffma
iii
Asst. Wildlife Researcher

_

Colorado.

for capturing

J. Wildl.

Wash.

State

in an increasing

blue grouse.

the age of young blue

�-151-

A P PEN

D I X

�BLUE GROUSE OBSERVATIONS

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_

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1/ Locat::i.on:
De s c r Lb e as completely
as possible
TO\1uship, Section,
forest,
dr ai.n agc , creek,
etc.

Observers

by County,

Nome:

Address:

Z..! VcJ]~_~..:..'-t_~:'!C'.
.l~"(1~0..: Nixed

conifers,
pine,
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bottoD, oak, pine-oak, aspen-pine, aspen-sagebrush,
etc.

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displaying,
etc.

Lf.s t added LnEormat Lon such aa birds
fceding,
resting
- female with chicks,

activity
nesting,

-

NOTE:

Please

return

this

form to:

Ri cha rd W. lIo ££111a11

Wildlife

Research

Center

P. O. Box 2287
fort Collins,
Colorado

80522

�April 1976

-153JOB PROGRESS

REPORT

State of

COLORADO
--------~~~.~~-----------

Project

No.

17

Work Plan No.
Job Title
Period

Continued

Job No.
Inventory

2

---------------------------------

of Selected Ptarmigan

Populations

------------------------------------------------------------------------

Covered:

Personnel:

Game Bird Survey

W-37-R-29

April 1, 1975 to March 31, 1976

W. John Arthur, Clait E. Braun, Ken Giesen, Richard W. Hoffman,
Tom Lytle, and Ron Oakleaf.

ABSTRACT
Investigations of white-tailed ptarmigan (Lagopus leucurus) populations at
four sites in Colorado initiated in 1965 were continued in 1975 with major
emphasis in ascertaining population levels and relationships of nesting
success and production to climatic conditions.
Breeding densities in 1975
increased in Rocky Mountain National Park and at Crown Point, remained stable
at Mt. Evans and decreased at Independence Pass. Observed increases at
Rocky Mountain National Park and Crown Point were the result of good production at these sites in 1974 and slightly better than average survival of
chicks to 1975. Stability was reached at Mt. Evans because of harvest
stabilization in 1974, while the population
decreased at Independence Pass
due to poor production and excessive harvest in 1974. Nesting success was
good at Rocky Mountain National Park, and Crown Point, average at Independence
Pass, and poor at Mt. Evans.
Nesting was delayed by cool wet weather in May
and June, but relationships between production and climate were unclear.
Hunting pressure was apparently low at Crown Point, the season was closed at
Mt. Evans, while about 63 percent of the fall population was harvested at
Independence Pass. Initial analysis of population data from Mt. Evans and
Crown Point indicate that males of both age classes (yearling and adult)
have higher survival rates than females.
Survival of chicks is poor each
year, averaging from 14 to 21 percent at the two areas. No real differences
were found in nesting success between yearling and adult females at either
area.

�-154-

RECOMMENDATIONS

Data from this project should be prepared for publication as a Wildlife
Monograph, with sho~ter technical papers being submitted to appropriate
Journals.

�-155-

CONTINUED

INVENTORY

OF SELECTED

PTARMIGAN

POPULATIONS

Clait E. Braun

Knowledge concerning population levels of white-tailed ptarmigan and factors
influencing observed densities is essential for proper management.
This
project was undertaken to examine long term population trends and effects of
climatic conditions on white-tailed ptarmigan in Colorado.
The initial 5
years of research on this grouse were previously reported (Braun and Rogers
1971).
Progress reports on work since 1969 have been prepared (Braun 1971a,
1972, 1973, 1974, 1975).

P. S. OBJECTIVES
To test the hypotheses that (1) populations of white-tailed ptarmigan in
Colorado are not cyclic, (2) production and fall population densities of this
species in Colorado are related to temperature and moisture conditions during
June and early July of the same year, and (3) harvest of over 50 percent of
the fall population adversely affects breeding densities the following spring.

SEGMENT OBJECTIVES
1.

To estimate breeding population levels of ptarmigan in 4 study areas (Mt.
Evans, Crown Point, Independence Pass, and Rocky Mountain National Park).

2.

To estimate

3.

To obtain weather data from a representative alpine area for use in
determining correlations between spring weather conditions and fall
ptarmigan populations.

4.

To initiate data compilation
final report.

and analyses

METHODS

AND MATERIALS

nesting

success

and production

in the above areas.

in preparation

for writing

the

Techniques used were essentially those developed under Work Plan 17, Job 1 and
reported in detail by Braun and Rogers (1971), and updated by Braun (197la).
In 1975, green (Rocky Mountain National Park), pink and white (Mt. Evans),
and yellow (Crown Point, Independence Pass, and Mt. Evans) bandettes with
black numerals were used to mark all newly banded birds and those whose older
bandettes had become too worn for individual recognition.
Weather data were
obtained from the Institute of Arctic and Alpine Research (INSTAAR), University of Colorado, for reasons previously explained (Braun 1971a).
Hunting
statistics were collected through return of bands from successful hunters
(Independence Pass and Crown Point).
A check station was operated opening
weekend on the only access road at Crown Point.

�-156DESCRIPTION

OF AREA

Areas intensively investigated have been described in detail by Braun and
Rogers (1971) and have been presented by Braun (1971a).
Braun (1971b) further
described vegetation and habitat usage of the study areas.

RESULTS AND DISCUSSION
Breeding

Densities

Censuses of breeding birds were initiated on 9 May at Rocky Mountain National
Park, 15 May at Mt. Evans, 26 May at Independence Pass, and 4 June at Crown
Point.
Formation of pairs occurred throughout May, with most pairing being
completed by 30 May. Timing of breeding activities was delayed in 1975,
apparently' the result of adverse weather conditions.
Densities observed are
presented in Table 1.

Table 1.

White-tailed

ptarmigan

breeding

densities,

all areas, 1975.

No. of
Breeding
Pairs

Unmated
Males

Total
Breeding
Population

Birds per
Square
Mile

1.25

9

5

23

18.4

Toll Memorial

.19

5

5

15

78.9

Fall River Pass

.70

9

5

23

32.9

Total - Rocky Mountain
National Park

2.14

23

15

61

28.5

Crown Point

1. 93

3

2

8

4.2

Mt. Evans

1.54

11

3

25

16.2

1.12

4

6

14

12.5

Study Area

Rocky Mountain

S.l.ze1/

National

Park

Tombstone Ridge-Sundance
Mountain

Independence

Pass

1) In square miles.
Breeding densities between 1974 and 1975 remained stable at Mt. Evans (no
change), decreased at Independence Pass (3.6/birds sq mi), and increased at
Crown Point (.6 bird/sq mi), and Rocky Mountain National Park (7.9 birds/sq mi).
Nesting success in 1974 was good (about 60 percent of all hens were successful)

�-157on all study areas (Braun 1975); consequently, increases in breeding densities
were expected in 1975. The expected increases did not materialize at Mt.
Evans, possibly the result of hunter harvest in 1974. However, only about 32
percent of the fall population at this area was harvested in 1974 (Braun 1975).
Possibly this is the approximate harvest level that can be attained without
adverse effects on the spring population the following year.
Previous data
collected on this study indicated that optimum harvest of white-tailed ptarmigan approached one-fourth (25 percent) to one third (33 percent) of the fall
population.
As in most years since 1969, the fall population at Independence
Pass was again over harvested in 1974, as at least 40 percent of the population
was removed.
Pairs did not re-establish on this area in usual numbers in 1975
and the population declined further.
This area is presently a sink for ptarmigan recruited from surrounding areas.
The population at Crown Point was expected to increase more than was documented.
Reasons for this are not known, but are undoubtedly related to poor recruitment of the few males produced on the area in 1974 (only one brood was present)
and low immigration to Crown Point from the alpine areas one to two miles
west.
Recruitment of yearling birds to the spring population in Rocky Mountain National Park in 1975 was excellent, as 30 different individuals were
identified.
The population at this area increased as the result of continued
good survival of adults and increased survival of young when compared to other
years (20 yearlings in 1974, 14 in 1973).
Breeding densities at all study areas from 1966 through 1975 are presented in
Table 2. Data presented in this table would indicate that unhunted populations
oscillate about every 7 to 10 years.
Data from the hunted areas show no clear
pattern, but clearly demonstrate that hunting is a major mortality factor
for white-tailed ptarmigan, as most if not all, major changes in breeding
densities are related to level of harvest.
Obviously, white-tailed ptarmigan
do not compensate for hunting mortality by having larger clutches, better
chick survival or increased survival of adults.
Nesting

Success

and Production

In 1975, the month of May was cool and wet (Table 3). Snow melt was retarded
and plant phenology was delayed.
June was also cooler and wetter than in
1974; consequently nesting activities were retarded on all areas.
Most laying
occurred in July and the estimated peak of hatching was 20-25 July.
Giesen
(1976) located 11 nests of ptarmigan in Rocky Mountain National Park, of
which 6 were successful.
Of the six nests hatching, four hatched between 2025 July, while two hatched between 1-7 August.
Nesting success was good at Rocky Mountain National Park (22 of 36 hens observed were successful) (Giesen 1976), Crown Point (2 of 2 hens observed were
successful), average at Independence Pass (1 of 2 hens observed were successful), and poor at Mt. Evans (3 of 10 hens observed were successful).
Seven
hens were observed during the brood periods off major study areas, of which
4 were accompanied by chicks.
Considering all areas, 32 of 57 hens (56.1
percent) were successful in 1975.
This is slightly down from the estimated
62 percent success in 1974.

�-158Table 2.

White-tailed ptarmigan breeding densities 1966-1975.

Year

Rocky Mountain
National Park

1966

29.0

1967

Study Area
Crown Point

Mt. Evans

Independence Pass

14.5

7.8

18.7

25.2

21.2

7.1

18.7

1968

29.4

18.1

7.1

18.7

1969

30.8

14.5

5.8

17.9

1970

24.8

10.4

5.2

10.7

1971

23.4

5.7

11.0

21.4

1972

22.4

1.0

19.5

18.7

1973

20.1

4.7

16.2

17.0

1974

20.6

3.6

16.2

16.1

1975

28.5

4.2

16.2

12.5

Table 3.

Weather data, Niwot Ridge, 12,300 ft, May-July 1975.

May

Month
June

July

Temperature
Maximum

53

F

60

F

62

F

Minimum

6 F
35.6 F

12

F

33

F

Mean daily maximum
Mean daily minimum
Days temperature

&lt;320

F

23.3 F

42.3 F
28.4 F

39.6 F

25

21

F

0

F

53.3 F

Precipi tation (inches)

4.58

2.42

Y

1.74

Days relative humidity &gt; 90%

18

91/

15

Wind speed (mph)

'l:./

'l:./

'l:./

1/ Only 17 days of data.
2/ Data not available.

�-159Initial brood size in 1975 was approximately 5.6 (Table 4) (Giesen 1976) and
steadily decreased to 15 September, after that time average brood size
increased slightly due to brood breakup and formation of gang broods.
These
data are similar to those collected in most years of the study.

Table 4.

Nmnber of broods

and average brood size, all areas, 1975. 1/ Y

Number of
Broods Observed

Average Number of
Chicks per Brood

July 16-31

5

5.6

August 1-15.

12

3.9

August 16-31

12

3.7

Month

September

1-15

15

3.5

September

16-30

8

4.2

1/

Only distinct broods

are included.

~/ Data in part from Giesen

(1976).

Fall Densities
Estimates of densities of white-tailed ptarmigan on September 1, while useful
in illustrating population gain through production, are difficult to accurately derive due to a number of variables.
Estimated fall densities for each
area studied are presented in Table 5. These densities were calculated
following the three basic assumptions discussed in detail by Braun and Rogers
(1971). Percent gain was adequate only at Rocky Mountain National Park and
Crown Point to maintain and increase populations the. following spring.
If
survival is stable, populations at these two areas should increase in 1976.
The population at Mt. Evans is not expected to decrease, but no real increase
is expected.
The population at Independence Pass should decrease in 1976,
but may not due to immigration from surrounding areas onto the study site.

Hunting Mortality
In 1975 all alpine areas of the state were open to ptarmigan hunting from 13
September through 5 October, except for Unit 70 (Pikes Peak) and the area in
Unit 52 within one-half mile on each side of State Highway 5 (Mt. Evans Road)
from the Echo Lake turnoff to the smnmit of Mt. Evans.
These areas were closed
to ptarmigan hunting.
As in recent years, no ptarmigan hunting was allowed
during any of the early big game seasons.
Bag and possession limits for
ptarmigan alone were 3 and 6.

�-160Table 5.

Estimated

fall densities

of white-tailed

ptarmigan

1975.

Rocky Mountain
National Park

Crown
Point

Mt. Evans

60

65

30.0

50.0

Average brood size
on September 1

3.9

4.0

3.5

2.0

Total production

55

8

12

4

Total bree?-ing population

61

8

25

14

Total population
September 1

110

15

35

17

Birds per square mile

51.4

7.8

22.7

15.2

Percent

50.0

53.3

34.3

23.5

Estimated
nesting

percent
success

gain

Independence
Pass

on

Since no bands have been reported from Crown Point since 1970 a check station
was operated on the only road into the area on both 13 and 14 September.
Purpose of the check station was to ascertain if ptarmigan were being harvested
on the study area as non-reporting
of bands was suspected due to the low
increase in this population since 1972.
During the two days of dawn to dusk
station operation, no ptarmigan were checked.
As in every year since 1970,
no bands were reported from the Crown Point area. Thus hunting is ruled out
as the cause for the continued low population at this site.
The hunting season at Mt. Evans in 1975 was effectively curtailed by the
management closure of the area within one-half mile of the paved highway.
Despite the closure, 4 banded ptarmigan were harvested on the study area by
two hunters, along with one unbanded bird (apparently a young of the year).
Thus at least 4 of at least 39 banded birds known to be alive (10.3 percent)
were harvested in 1975 with a closed season on the study area.
Of the five
birds known to have been harvested, three were adult males (3±, 3+, 3+), one
was an adult female (3+) and the other was a presumed chick, unknown sex.
Band recoveries reported by hunters from ptarmigan harvested at Independence
Pass are suspected to have decreased in recent years.
This trend may not
have continued in 1975, as bands from 12 of at least 19 banded birds known to
be alive (63.2 percent) were received.
Obviously, the population at this site
was over harvested in 1975, a situation that has occurred almost every year
since 1968. This population should decrease further in 1976 unless maintained
by immigration from birds produced in adjacent areas.

�-161Analysis

of Long Term Population

Data

Analysis of all data collected from 1966 through 1975 was initiated for
preparation of the final report.
Population and movement data were summarized
for Crown Point and Mt. Evans but not for the other two study areas. The
average annual survival rate based on reobservation and harvest data combined
within the same year for all birds banded was 31.4 percent (N = 74 7 236) at
Crown Point, and 36.5 percent (N = 127 7 348) at Mt. Evans.
Males had a
higher survival rate than females (41.7 to 29.3 percent for yearlings, and
57.1 to 29.3 percent for adults at Crown Point; 47.1 to 35.2 percent for
yearlings and 38.2 to 32.8 percent for adults at Mt. Evans).
Chicks survived
poorly at both areas, with average annual survival being 14.3 percent at
Crown Point and 21.4 percent at Mt. Evans.
Hunting decreased survival rates
at both areas.
Only three"movements
(one female to Rocky Mountain National Park and back, and
two (one female and one male) shot near Comanche Peak) were documented from
Crown Point to other areas, while one movement of a banded bird to Crown
Point was identified (female banded as a chick from Rocky Mountain National
Park).
Four birds banded at Mt. Evans (2+ male, 1+ female, chick female,
chick male) moved to Guanella Pass, while two birds (2+ female, yearling
female) moved to the Sugarloaf area northwest of Mt. Evans.
Nine movements
of birds banded at Guanella Pass (8) and Sugarloaf (1) were documented to Mt.
Evans.
Of these birds, 5 were females (four yearlings and one chick), while
4 were males (three yearlings and one adult).
Average success of hens in rearing young was compared at both sites.
At Mt.
Evans 18 of 47 yearling hens (38.3 percent) and 20 of 56 adult hens (35.7
percent) were successful in bringing off broods.
In contrast, 22 of 26
yearling hens (84.6 percent) and 17 of 18 adult hens (94.4 percent) were
known to be successful at Crown Point.
These data may be misleading, as
unsuccessful females apparently departed the Crown Point area and summered
elsewhere.
One known occurrence of this phenomena was documented at Mt. Evans,
unsuccessful hens from adjacent sites moved onto the study area in late summer,
thus the percentage of nesting success was lower than actual.
Of most
importance though is the fact that within areas, age of females had no
apparent relationship to probability of being successful or unsuccessful.
This is contrary to hypotheses developed from preliminary review of available
data.

�-162LITERATURE CITED

Braun, C. E. 1971a. Continued inventory of selected ptarmigan populations.
Colo. Div. of Game, Fish and Parks, Game Res. Rept. Fed. Aid Proj.
W-37-R. April. p. 91-106.
1971b. Habitat requirements of Colorado white-tailed ptarmigan.
Proc. Western Assoc. State Game and Fish Comms. 51:284-292.
1972. Continued inventory of selected ptarmigan populations. Colo.
Div. of Wildl. Game Res. Rept. Fed. Aid Proj. W-37-R. April. p. 91-98.
1973. Continued inventory of selected ptarmigan populations. Colo.
Div. of Wildl. Game Res. Rept. Fed. Aid Proj. W-37-R. April. p. 355361.
1974. Continued inventory of selected ptarmigan populations. Colo.
Div. of Wildl. Game Res. Rept. Fed. Aid Proj. W-37-R. April. p. 65-72.
1975. Continued inventory of selected ptarmigan populations. Colo.
Div. of Wildl. Game Res. Rept. Fed. Aid Proj. W-37-R. April. p. 81-89.
_____ , and G. E. Rogers. 1971. The white-tailed ptarmigan in Colo.
Game, Fish and Parks Tech. Publ. No. 27. 80 p.

Colo.

Giesen, K. M. 1976. Organization, mortality and dispersal of grouse broods.
Colo. Div. of Wildl. Game Res. Rept. Fed. Aid Proj. W-37-R. April. In
press.

Prepared by __~i~

/&lt;:l ~:~ t- /_/ /C:"

!&lt;)~~;~,. ~

~L2~-/~--~--~--~-~~~---_-_-~/~
~~_-~!~~~~~'~~_~-~'~~-~
__
__
'

__

Clait E. Braun "/
Wildlife Researcher

_

�April 1976
-163JOB l'ROG.R.ESS

State

of

Project

.

~COlORA=D~O

No.

Game Bird Survey

17

Organization,

Period Covered:
Personnel:

_

W-37-R-29

ivork Plan No.
Job Title

}~EPORl'

5

Job No.
Mortality

and Dispersal

April 1, 1975 through March

of Grouse Broods

31, 1976

Kenneth M. Giesen, William W. Mautz and Kenneth R. Russell, Colorado Cooperative Wildlife Research Unit; Clait E. Braun and
Richard W. Hoffman, Colorado Division of Wildlife; Ed Blekicki,
Colorado Mountain College.

ABSTRL'&gt;.CT

Investigations of the nesting ecology, organization, mortality and dispersal
of broods of white-tailed ptarmigan (Lagopus leucurus) in Colorado were
initiated in 1975 on approximately 16.9 km2 of alpine habitat in Rocky
Mountain National Park. Average clutch size determined from 11 nests
was 5.7 eggs. The peak of hatch occurred between 20 and 25 July,
with a smaller peak due to renesting occurring between 5 and 10
August.
Nesting success was approximately 60 percent, with observed egg
hatchability being 100 percent.
Twenty-two broods were located and observed a total of 134 times prior to 31 October.
Brood size decreased over
31 percent by 15 August with the only documented mortality due to predation.
Movements of 15 broods to summer areas averaged less than 600 m, while
movement within the summer ranges averaged 212 m between observations.
Size
of brood home range varied from 14 to 32 ha. Movements to fall concentration
areas averaged 1,405 m and were precipitated by a major snowstorm on 13
October.
Movement to wintering areas was gradual and documented movements
for 12 chicks from fall to winter sites averaged 3,310 m. Brood breakup
occurred gradually after chicks reached 8 weeks of age and most dispersal
occurred after the 13 October snowstorm.
At least 3 broods remained intact
until mid-November.

�-164RECOMMENDATIONS

1.

A minimum of 2 hens should be fitted with radio-transmitters
at time
of hatch in order to facilitate daily observation and monitoring of
movements and chick mortality.

2.

In order to facilitate individual chick identification, all chicks
should be marked with numbered bandettes in addition to aluminum state
bands.

3.

A minimum of 6 chicks should be fitted with radio transmitters prior
to dispersal in order to monitor dispersal movements and mortality.

4.

A minimum of 20 chicks from adjacent alpine areas should be marked in
order to document dispersal movements of chicks onto the study area.

5.

Intensive effort should be made in December to locate all wintering
areas surrounding the study area in order to locate wintering juveniles previously banded on the study area.

�-165-

ORGANIZATION,

MORTALITY

AND DISPERSAL

OF GROUSE BROODS

Kenneth M. Giesen

Grouse are important game birds in Colorado and four species [blue grouse
(Dendragopus obscurus), sage grouse (Centro cercus urophasianus), sharptailed grouse (Pedioecetes phasianellus), and white-tailed ptarmigan]
presently provide considerable hunting opportunity.
Little is known concerning broods of three of these species in Colorado (blue grouse, sharptailed grouse, and sage grouse) due to habitats occupied and lack of detailed
study. Production of young and survival to the hunting season and following
breeding season are important biological parameters as these factors affect
fall hunting populations and breeding populations the following year.
It is
thus important to understand organization and dispersal of broods as they
directly affect hunting success and overwinter survival.
It is also important to identify major mortality factors of juvenile grouse and to understand their relationship to fall and resulting spring population levels.
Intensive studies of white-tailed ptarmigan in Colorado (Schmidt 1969, Braun
and Rogers 1971, Hoffman and Braun 1975) support the conclusion that their
biology is similar to that of other grouse, especially in respect to brood
ecology and dispersal.
Preliminary observations of white-tailed ptarmigan
broods suggested they would be ideal study organisms as (1) they can be
easily and readily located, (2) they can be easily marked, (3) they occupy
small home ranges until time of dispersal, (4) they can be observed without
vegetative interference, and (5) requirements for wintering are known.
P. S. OBJECTIVES
The objective of this study is to increase the knowledge of factors affecting
numbers of young grouse in fall and recruitment to the breeding population
the following spring.
Hypotheses which have been developed are: (1) mortality of chicks is predator-dependent
to 1 October and weather-dependent
from 1 October to 1 November; (2) brood breakup and dispersal are functions
of chick aggression to brood mates; (3) timing of dispersal is age-dependent
with dispersal not occurring until chicks are older than eight weeks; and
(4) dispersal is non-random.
SEGMENT OBJECTIVES
1.

Locate a minimum

of three nests for documentation

2.

Locate and individually

3.

Document timing and causes of mortality
from hatching to dispersal of young.

mark a minimum

of hatching.

of three hens with broods.
for members

of three broods

�-168-

METHODS

AND MATERIALS

Territorial pairs in spring and females with broods in summer were located
with the use of tape-recorded calls (Braun et al. 1973).
Intensive search
on foot and frequent viewing with binoculars were necessary to locate nonterritorial males and unsuccessful females.
Once ptarmigan were located they were observed using 7 X 50 or 8 X 32
binoculars.
If the bird had been banded previously, its band color and
number were recorded along with the time and location, on standardized
observation cards. Unbanded birds were pursued until captured or flushed
out of sight.
Captures were made with a 60 pound test nylon covered wire
snare (40 pound test for chicks) attached to the end of a 5 or 7 m telescoping fiberglass fishing pole as described by Zwickel and Bendell (1967b)
for blue grouse and adapted for ptarmigan by Braun and Rogers (1971).
Each adult ptarmigan newly captured was banded on the right leg with a
gold aluminum serially numbered leg band (Size 8, National Band and Tag Co.,
Newport, Kentucky) and on both legs with colored, numbered (1-100) plastic
leg bandettes (Size 6, National Band and Tag Co., Newport, Kentucky).
Chicks captured after 2 weeks of age were banded on the left leg with a gold
aluminum serially numbered leg band.
Forty-five selected chicks were additionally leg banded with various combinations of colored plastic coil bandettes (Size 3, National Band and Tag Co., Newport, Kentucky).
All captured ptarmigan were weighed with a spring scale accurate to within
5 gros. Measurements of carpal length, individual primary lengths and length
of the outer rectrix were recorded using a transparent, flexible plastic
millimeter ruler.
Adults were sexed and classified as either adult or
yearling according to plumage characteristics
described by Braun and Rogers
(1971).
Chicks were aged by examination of various plumage characteristics
including lengths of carpal and primary feathers and by progression of primary molt.
Ages were assigned by comparing specific measurements with those
of known age chicks.
Sex of chicks was based on behavior, primary feather
and carpal lengths, pigmentation of the eye stripe
and by the amount of
pigmentation on post-juvenile P8, juvenile P9, PlO, and PClO (Braun, pers.
comm.) .
Nests were located by (1) observation of laying hens, (2) following incubating
hens back to nests after feeding, and (3) intensive search of known territories.
Additional nests were located incidental to other field activities.
After incubation began, eggs were measured using a vernier caliper.
RESULTS

AND DISCUSSION
Nesting

Description

of Nest Sites

Characteristics
of rock and vegetative cover, slope and aspect were determined for each nest site (Table 1). Six of 12 nest sites examined were in
a spruce (Picea engelmanni) or willow (Salix spp.) krummholz situation, while
5 were in rock or boulder fields and 1 was in an open alpine meadow site.

�-169Table 1.
in 1975.

Nest site characteristics

Rockl/
Cover-

for nests located within

the study area

Height of
2/
Vegetative Cover-

Nest

Aspect

1/
Slope-

BW 7

South

23

10

Salix-Picea

krummholz

15

BW14

North

12

50

Geum-Carex

rock field

8

BW16

North

26

50

Geum-Trifolium

BW17

North

43

5

Salix-Picea

krummholz

26

BW27

East

21

35

Picea-Salix

krummholz

150

BW36

East

11

5

Salix krummholz

G34

North

23

50

Geum-Trifolium

G37

South

38

30

Geum-Carex

rock field

14

G74

West

10

5

Salix-Picea

krummholz

30

G76

West

38

75

Geum-Trifolium

G84

South

35

10

Geum-Carex

meadow

Unk1/ West

21

5

Picea-Salix

krummholz

Vegetative

Cover Type

rock field

6

40
rock field

boulder

field

5

5
9
150

1.1 In percent.
]j Measured

in centimeters.

1/ Nest site 1974.
Slopes of the nest sites ranged from 10 to 43 percent and averaged 25 percent.
With the possible selection against steep slopes, no selection
for slope
was apparent.
The slopes recorded appeared to be typical of those within the
study area. Aspect did not appear to be important, as every direction was
represented.
Since nests are generally found within the male's territory
(Schmidt 1969, Braun and Rogers 1971), hens are generally limited on selection of aspect.
Height of vegetation surrounding the nests varied with site. Two nests found
under dwarf spruce were surrounded by spruce 100-150 em in height, while
the 4 nests found in willow clumps were surrounded by vegetation 15-40 cm
in height.
Vegetation heights at nest sites in the open alpine or in rock
or boulder fields averaged less than 10 cm.

�-170Five nests were located in areas having rock or boulder cover comprising
an average of 50 percent of the ground cover. Three of these nests were
adjacent to rocks while 2 were within 20 cm of rocks. Height of the rocks
ranged from 8-24 cm and may have served to protect the nests from wind, as
the nests were east or north of rocks, while the prevailing wind was
from the west.
Choate (1960) and Braun and Rogers (1971) mentioned that nests are characteristically located in snowfree sites.
Choate (1960) also observed that
nests were generally close to running water.
This characteristic may not
indicate selection as much of the alpine has an abundance of water in the
spring due to rapidly melting snow. These same areas may be extremely dry
in mid-July.
Two char act er t.st r cs were common to the 12 nest sites examined.
Nests were
generally protected from the wind by either rock or vegetative cover, and
all nests were in or immediately adjacent to open areas. These open areas
may have facilitated access for the hens and escape from nest predators.
Clutch Size and Egg Characteristics
Clutch size was determined from 10 complete clutches observed during incubation.
Average clutch size was 5.7 eggs, with clutches of 4 eggs (1),5
eggs (3), 6 eggs (4), and 7 eggs (2) recorded.
The clutch of 4 eggs
was thought to be the result of renesting.
Braun and Rogers (1971) reported
an average clutch size of 5.7 eggs determined from 20 complete nests located
from 1966 to 1969. Choate (1963) reported an average clutch size of 5.2
eggs for 10 nests located in Glacier National Park, Montana.
Measurements recorded for 56 eggs from 10 clutches averaged 44.1 X 29.8 mm
(range 39.2-47.8 mm and 28.5-30.9 mm) in size. Braun and Rogers (1971)
reported an average of 43.2 X 29.6 mm for 67 eggs examined.
Egg fertility was 100 percent as all eggs (32) in the 6 successful nests
hatched.
Braun and Rogers (1971) reported 81.1 percent of 37 eggs in 7
successful nests hatched, while in Montana Choate (1963) reported 86 percent
of the eggs in successful nests hatched.
Known dates of hatching for 6 successful nests were: 21 July (1), 22 July
(2), 23 July (1), 1 August (1) and 7 August (1). The clutch hatching on 7
August was thought to be a renest.
Behavior

of Females During Egg Laying

Activities associated with egg laying were observed for 7 females (Table 2).
All hens observed visited their nests for deposition of eggs between 0945
and 1430. Three females spent an average of 94 minutes at the nest while
depositing the second egg of their clutches.
Another female spent over 280
minutes at the nest when laying the fifth egg of a 6 egg clutch.
It was
assumed that incubation began at this time, as the hen remained on the nest.

�-171Table 2.

Time spent on nest during egg laying.

Hen
Time approached

nest

Time spent on nest
Time left nest site

G84

BW36

BW17

BW27

G37

BW7

G76

1120

1340

0945

&lt; 1430

1141

1231

1220

84.1/ &gt; 140

44

ND~j

ND

153

&gt;280

1029

ND

&gt;1400

1244

&gt;1600

1504 &gt; 1710

No. eggs in nest

2

2 or 3

2

1

ND

2

Time spent covering eggs

64

ND

40

ND

ND

34

5
ND

]) Time in minutes
]j No data.
A generalized description of the daily activities of a white-tailed ptarmigan
pair during egg laying is presented.
This description is based on 38 hours
of observation of 7 different pairs on the day of egg deposition.
Prior to approaching the nest the pair alternately feeds and rests within
their territory.
The female generally rests at the base of a rock or shrub
while the male selects a rock or hummock overlooking the female.
During
periods of activity the female feeds more intensively than does the male.
When the time for egg deposition nears, the female rapidly approaches the
nest site (Normally 100-200 m distant) with the male following 5-10 m behind.
During nest occupation by the female the male will select a site overlooking
the nest, approximately 10-20 m away_ During the first 10-20 minutes on the
nest the female moves around and often pokes her head underneath her breast
as if moving the eggs. Apprently the egg is dropped at this time as continued observation reveals little further activity by the hen until she prepares
to leave the nest.
Before leaving the nest the female becomes restless and
begins pecking at nearby vegetation and dropping the vegetation near her,
thus building up the rim of the nest bowl.
As the hen begins reaching
further and further for vegetation, she is forced to stand up. From the
standing position near the nest she begins covering the eggs by dropping bits
of vegetation on them at the rate of approximately 20 pieces of vegetation
per minute.
Once the eggs are no longer visible the stimulus for covering
them diminishes and the female suddenly leaves the nest and joins the male.
Once the female is away from the nest she begins feedings.
The average interval between deposition of successive eggs was estimated to
be between 30 and 36 hours.
Three females (BW7, BW17, G84) deposited 3 eggs
within a 4 day period, and another female (BW27) deposited 4 eggs within a
6 day period.
Approximately 9 days were required for a complete clutch, with
incubation beginning 1 or 2 eggs prior to clutch completion.
Behavior

of Incubating

Females

Females remain on the nest during the entire 22-23 day incubation period,
except for short feeding excursions at dawn and dusk. At these times, the

�-172-

female normally flies from the nest to the feeding area where she actively
feeds for 10-20 minutes.
The same feeding areas are used consistently.
Upon completion of feeding the hen walks or flies back to the nest.
During
these feeding periods the eggs are not covered.
Observations indicate that some hens feed during the day. Female BW14 was
observed feeding during the day on 3 occasions, always during foggy weather.
Another hen (G34) was twice observed feeding between 0800 and 0815 during
clear or partly cloudy weather.
Of 68 daytime observations made of nests
during incubation, females were absent on only 5 occasions (7.4 percent).
Disturbance of incubating females early in incubation often caused the
hen to flee the nest and perform distraction displays (Schmidt 1969).
During later stages of incubation hens became reluctant to leave the nest,
even when .t.o uched , and had to be physically forced off in order to examine
the eggs. Hens immediately returned to the nest after disturbance by the
observer.
Nesting

Success

Nesting success was estimated from (1) success of known nests, (2) numbers
of successful and unsuccessful hens (those hens with and without broods,
respectively) seen during the brood season and which had previously been
identified during the breeding census, and (3) total numbers of successful
and unsuccessful hens observed during the brood season.
Close agreement
among the 3 methods suggest that actual nesting success was approximately
60 percent.
Six of 11 active nests hatched successfully (55 percent) but this estimate
of nesting success may be high since only 2 of 7 nests located prior to
onset of incubation were successful.
Renesting compensated for this loss
as 2 of 5 hens whose nests were predated renested successfully.
Thus 57
percent (4 of 7) of the hens whose nests were located during egg laying
succeeded, either initially or through renesting, in bringing off broods.
All 4 nests found during various stages of incubation hatched successfully.
During the breeding census, 31 females were observed.
All females were seen
with a male and were thought to be paired.
Of these 31 females, 22 were
later seen during the brood season (13 successful, 9 unsuccessful) giving
an estimated nesting success of 59 percent.
This estimate may be biased
as success of the 9 females observed during the breeding census but not
during the brood season was not determined.
Braun and Rogers (1971) suggest
that all females attempt to nest.
Since the brood survey was thorough, it
is doubtful that many successful females escaped observation.
POSSibly some
of the 9 females died or nested elsewhere.
Another possibility is that most
of these hens were unsuccessful and were not observed during the brood
season because they summered off the study area (Braun pers. comm.).
Another method for estimating nesting success is to examine the total number
of successful and unsuccessful hens observed during the brood season.
From
mid-July until dispersal in mid-October 36 hens were observed (22 successful,
14 unsuccessful) for a calculated nesting success of 61 percent.
There are 2

�-173shortcomings of this method.
Unsuccessful hens are less likely to be
observed than successful hens (Braun and Rogers 1971) and there is possible
movement of hens onto or off the study area during the brood season.
These
factors may compensate for each other as the study area is on a ridge
almost completely surrounded by subalpine vegetation.
This would inhibit
the movement of hens with young broods but not broodless hens.
Most of
the successful hens on the study area were thought to have been observed.
It is probable that movement of unsuccessful females onto the study area
equalled movement of unsuccessful hens away.
Brood Relationships
Changes

in Brood Sizes

Initial brood size was estimated from average clutch size (5.7) and observed
egg hatchability
(100 percent).
From time of hatch until 31 October, 134
observations of brood sizes were recorded.
From these data, changes in
brood size were calculated for biweekly intervals during the summer (Table 3).
It is apparent that a large decrease in brood size occurred between 1
August and 15 August.
Part of this decrease was caused by the appearance of
broods resulting from renests which are normally composed of 4 or fewer
chicks (Braun and Rogers 1971).
The decrease in brood size was real, however, as 7 of the broods which hatched in July averaged 4.0 chicks by 15
August, a decrease of 32.1 percent from the original 5.9 chicks per brood
at time of hatching (first nesting only).
By 15 September, brood size had
decreased over 37 percent from time of hatch.
This early mortality is
similar to that reported by Zwickel and Bendell (1967a) who estimated a
40 percent loss of blue grouse chicks in the first two weeks of life.

Table 3.

Changes

in average brood size between

16 July and 31 October,

Number
Broods
Observed

Average
Brood
Size

Change From
Previous
Period

Change
From
.Hatch

Y

Date

1975.

16 July-3l

July

5

5.6

- 1. 8

- 1.8

1 August-IS

August

10

3.9

-30.4

-31.6

16 August-3l

August

11

3.9

0.0

-31.4

1 September-IS

September

9

3.6

- 7.7

-37.5

16 September-30

September

8

4.1

+13.9

-27.7

1 October-IS

October

10

4.2

+ 2.4

-26.3

1 October-3l

October

11

3.2

-22.0

-31.4

1/ Change from hatch.

�-174After mid-September,
average brood size began to increase.
At this time,
chicks were over 8 weeks of age and brood breakup had begun.
Chicks were
temporarily attracted to certain hens and large "gang" broods resulted.
One hen, G84, was seen with as many as 10-12 chicks during this period.
In October many chicks were seen apart from broods, most often in large
mixed flocks of adults.
Such observations were not included in brood
counts since marked chicks from several broods were often represented.
Mortality

Causes and Timing

Causes of mortality in free living animals if often difficult to document.
Remains of birds disappear rapidly and the predators, which are common on
the alpine, are capable of completely consuming ptarmigan chicks.
Any
remains may be buried or otherwise hidden from sight.
Mortality was documented in 2 instances involving a total of 7 chicks.
In
the first instance, a raptor, probably a prairie falcon. (Falco mexicanus),
killed a hen less than 24 hours after she hatched her clutch of 6 eggs.
The entire brood was assumed to have died, although only 1 chick was found.
The other case involved 1 chick approximately 4 weeks old. The remains
were found approximately a month later and death was attributed to a weasel
(Mus tela spp.).
Brood-Female

Interactions

Selected broods were intensively observed at irregular intervals between
time of hatch and brood breakup.
Observations were made using 8 x 32
binoculars from semi-concealed positions.
Occasionally, suitable concealment was lacking and broods were observed from an open position but this
did not appear to noticeably affect their behavior.
Distance between brood
and observer varied from less than 10 m to over 100 m but was generally less
than 50 m. Schmidt (1969) observed broods under similar conditions without
noticeably influencing their activities or behavior.
Lengths of observations
ranged from less than 30 minutes to over 10 consecutive hours.
Emphasis was
placed on recording brooding, feeding and maintenance activities.
Reactions
of the hen and brood were also recorded when the chicks were captured for
banding and measurement of growth parameters.
Brooding was observed most often in chicks less than a week old. Observations
of 3 broods on the day of nest abandonment indicated that an average of 69
percent of the day time (328 out of 470 minutes of observation) was devoted
to brooding.
Schmidt (1969) reported that day old chicks were brooded over
50 percent of the time.
Theberge and West (1973) observed that Alaskan
rock ptarmigan chicks were no longer brooded after 6 days of age. May (1975)
presented data suggesting that white-tailed ptarmigan chicks are able to
maintain body temperatures at 15-20 days of age after which time the need
for brooding decreases.
Brooding of older chicks sometimes occurs under
certain conditions.
On several occasions, female G84 was observed brooding
her 4 chicks when she was approached by the other 4 chicks which she had
adopted, in most instances all 8 chicks were successfully brooded.
Thes~
latter 4 chicks were over 2 weeks older than her own chicks and at the tlmes

�-175-

of observation were approximately 4 weeks old. Braun (pers. comm.)
observed a female attempting to brood 6-7 chicks during a snowstorm in
September 1967. Brooding was rarely observed in the day time after chicks
reached 2 weeks of age except during inclement weather.
Observations were recorded of routine or maintenance activities.
Most
periods of chick activity were devoted to feeding.
Brood hens appear to
feed only occasionally, but appeared to be more proficient and thus,
probably required less feeding time. Young chicks appear to be inefficient
feeders and close observation revealed that several pecks were often required
to obtain insect or vegetative food particles.
Although feeding behavior
of chicks may be learned by imitating their mother, May (1975) showed that
juveniles ingest a wider variety of food materials than do adults.
After chicks were 3-4 weeks of age, they became more independent and foraged
greater distances from the female.
It often appeared that foraging chicks
determined the direction of daily movement.
At certain times when the brood
hen appeared to have a certain goal, she rapidly led the brood while giving
the chick attraction call. These goal-orientated hen movements were generally
observed when the female led her brood to dusting locations and when the
female moved to a night roosting site. At times these movements were 200-300
m and were traversed in 10-20 minutes.
Brood-Brood

Interactions

Successful hens usually abandoned their breeding territories and moved to
more favorable brood rearing habitat within several days of hatching.
Since
only a small portion of the alpine tundra is preferred brood habitat, successful hens often concentrated in relatively small areas.
Schmidt (1969) reported 8 broods in 1967 and 10 broods in 1968 utilized an area of approximately
130 ha surrounding an old quarry at Toll Memorial.
Such concentrations
increase chances of contact between broods and thus provide opportunities
for brood mixing.
Chambers and Sharp (1958) reported that brood mixing in
ruffed grouse occurred more frequently during population highs when broods
tend to come into contact more often.
From hatching until 13 October when summer areas were abandoned, 116 observations were recorded of 21 b"roods. Over 84 percent of the observations (97)
were of single broods while combined broods were observed on 14 occasions.
In addition, there were 5 observations of unsuccessful hens associating
with successful hens and their broods.
Occasionally apparent bonds were
temporary and the recorded observation was likely a chance encounter between
the broods involved.
In most situations the association was of long duration.
Successful hens BW16 and G85 were observed together on 9 occasions (of 9
total observations) between 22 September and 7 October (16 days).
Successful hens BW5 and G86 were observed together on 3 occasions (of 4 total observations) between 11 August and 3 October (54 days).
On at least one occasion
(3 October), all 4 of the above hens with their broods were observed foraging together.

�-176-

In both instances of combined broods, the chicks and hens foraged together
with no observed distinction between broods.. Neither of the hens showed
aggression towards each other or towards any of the chicks.
Aggression or
avoidance behavior was not noti.ced among any of the chicks in the combined
broods.
In both situations,. the chicks were the same approximate age and
it was impossible for the observer to distinguish the chicks from each
brood unless the chicks were captured and their bands read.
Unsuccessful hens or rarely males were sometimes associated with successful
hens and their broods.
Female G96 with her brood was observed on 5 occasions
between 20 August and 21 September (33 days) with G70, an unsuccessful
female, always in attendance.
Some hens seemed to adopt chicks from other hens during the summer and fall,
especially during brood breakup and dispersal.
One instance was documented
of the adoption of orphan chicks when female G84 adopted the 4 chicks of
female BW14 when the latter disappeared and was assumed de ad , In this
instance, female G84 had 4 chicks from a renesting attempt and the 4 chicks
she adopted were approximately 16 days older.
Even though there was an
age and size difference between the 2 "broods" of chicks, there was little
aggression observed.
Movements
Movements

to Summer Areas

Braun (1969) described summer brood areas as being high, rocky and frequently
windswept ridges.
Rocks over 30 cm diameter often comprised over 50 percent of the ground cover while late lying snowfields were present in protected areas.
The vegetation was dominated by sedges (Carex spp.) with Geum
rossii, Polygonum spp. and Trifolium spp. present in varying amounts.
These 4 groups of plants provide the majority of the food utilized by both
chicks and hens (May and Braun 1972, May 1975).
During the first few days after hatching broods often remain within a few
hundred meters of the nest site. Within a few weeks, successful hens with
their broods begin showing up at traditional summering areas where they
remain until brood dispersal or inclement fall weather (Braun 1969).
Observations of 4 broods whose nest sites were known revealed that they
moved an average distance of 510 m to summer areas. No movements greater
than 600 m were recorded between breeding territories and summer areas for
11 other successful hens.
Braun (1969) also observed an average movement
of less than 600 m for successful hens, while Schmidt (1969) documented one
movement of 1400 m in 4 days for one successful hen.
Movements

Within

the Summer Area

Successful hens with their broods move to specific brood rearing habitat
within 3 weeks after hatching (Schmidt 1969).
Once on the brood area,

�-177-

movements of the brood became localized until fall dispersal or snowstorms
forced the broods to move downslope into more sheltered areas.
Four brood concentration areas were located within the study area.
The concentration area at Toll Memorial was studied most intensively with the
Medicine Bow Curve, Iceberg Lake, and Sundance Mountain concentration areas
being studied less intensively.
Movements within a summer concentration area were recorded using extensive
daytime observations
(up to 10 consecutive hours) and by plotting day-today observations on a map.
Of 22 broods observed during the 1975 summer, 16
were observed more than once between 1 August when broods began appearing at
the concentration areas and 13 October when the first severe snowstorm
caused abandonment of these areas.
Three broods were observed only once
each, while the remaining 3 broods were observed only before 1 August or
after 13 October.
The 16 broods were observed a total of 115 times on the summer areas.
Each
observation was plotted on a 7.5 minute (scale 1:24,000) map, and distances
between consecutive observations measured to the nearest 25 m. Average distance moved between observations was 212 m for all broods combined, and
ranged from 67 to 726 m for individual broods (Table 4). Only 3 of 99
movements were greater than 600 m. Only 3 broods (G84, G85 and BW16) were
observed more than 10 times during the 74 day observation period and these
broods averaged 117 m between observations.
Home ranges were calculated for the 3 broods which were observed at least
10 times during the summer period.
Observations were plotted on a 7.5
minute (scale 1:24,000) map and outside points were connected.
Odum and
Kuenzler (1955) noted that estimated size of a home range tended to increase
with the number of observations until a maximum was reached.
With the small
number of observations recorded for each brood, size of brood ranges are
minimum estimates.
Brood G84, which was observed 28 times, had a calculated
summer range of 32 ha. Broods G85 and BW16 (observed 11 and 12 times, respectively) were often observed together and had calculated summer ranges of
approximately 14 ha.
Daily movements recorded during intensive observations indicated that young
broods move little, while broods over 4 weeks of age move longer distances.
Daily movements often tend to be circular routes and these routes are usually
similar from one day to the next.
Individual broods varied in their daily
movements according to abundance and availability of resources, especially
food. Brood G84 often moved 800-1600 m per day, while broods G85, G86,
BW5 and BW16 seldom traveled over 400 m in a day.
Movements

to Fall Areas

After the first snowstorm on 13 October, ptarmigan were observed in widely
scattered flocks throughout the study area. Fall use areas are generally
lower than summering areas and often at the edges of krummholz areas (elevation 3440-3480 m). These areas are characterized by an abundance of willow
shrubs and numerous small snowfields.
Braun (1969) suggested that ptarmigan utilized the last remaining green vegetation in these areas, which are

�-178the last areas to become snowfree in summer. At this time the ptarmigan
are also actively molting into the white winter plumage and are seldom
observed in windswept snowfree areas.

Table 4.

Brood movements within a summer area.

Average Distance Movedll
Between Observations

Brood

No. of Observations

G76

5

726

G34

7

260

G85

11

98

G86

5

186

G51

2

288

G56

5

402

G49

5

300

G84

28

176

G96

5

366

G99

7

132

BW5

6

67

BW14

2

192

BW16

12

78

BW25

3

108

BW27

5

318

BW39

7

300

Total

99

Average

II Distance in meters.

212

�-179-

Although brood breakup was recorded prior to the snowstorm on 13 October,
incidence of intact broods decreased rapidly after movement to fall areas.
Brood breakup was most apparent in late October as observed flocks of
ptarmigan contained mixtures of adults (males and females) and chicks.
Flock composition varied from day to day and little interaction was observed
between hens and chicks.
The incidence of brood breakup is reflected in the banding records.
Prior
to 1 October, 59 juvenile ptarmigan had been banded.
At time of banding,
these juveniles were with hens known or assumed to be their parent.
Between
1 October and 1 November, 24 juvenile ptarmigan were banded, of which only
6 were associated with hens throught to be their parent.
During November
an additional 12 chicks were banded and none was associated with hens
assumed to be its parent.
Not all broods broke up, as at least 3 broods
(G34, G76. BW27) remained intact until early November.
Most of the chicks newly banded after 1 October were assumed to have dispersed onto the study area from adjacent alpine areas.
This assumption is
supported by calculations of chicks produced on the ~tudy area and surviving
until 1 October.
Observations of 12 broods in September showed an average
brood size of less than 3.2 chicks (eliminating observations of known gang
broods).
With 20 successful hens remaining on the study area (1 hen was
killed, another hen lost all of her chicks shortly after hatch), the estimated production was 64 chicks as of 1 October.
Since a total of 95 juveniles were banded on the study area with 30 being of unknown origin, the
most likely explanation is that they were dispersing onto the study area.
Braun (pers. corom.) suggests that juvenile females disperse the greatest
distances in fall. Banding records from 1 October to 30 November support
this hypothesis, as 61 percent (22) of the chicks banded during this period
were classified as females.
Movements of 42 chicks to fall use areas from summer brood areas were documented.
Although some broods remained intact, most chicks were associated
with mixed age and sex flocks.
Movements to fall use areas were abrupt and coincided with the first major
fall snowstorm on 13 October.
Prior to the snowstorm, most ptarmigan were
closely associated with the few remaining snowfields.
These old snowfields
provide water and green vegetation and serve as a protective background
for the ptarmigan during their molt into white winter plumage (Braun 1969).
Average movement documented for 42 chicks was 1405 m (range 50-3900 m).
Prior to 13 October, 33 chicks had been observed at Toll Memorial.
Of these,
26 were later located on fall use areas with 73 percent (19) moving to
Tundra Curves approximately 1000 m west.
One chick was documented moving
to Medicine Bow Curve (3900 m nothwest), while 4 chicks moved to Sundance
Mountain Basin (2900 m southeast).
Apparent dispersal of juveniles onto the study area occurred during October.
Although all portions of the study area were not searched equally, 2 fall
use areas, Tundra Curves and Fall River-Gore Turnout attracted dispersing
chicks.
Of 21 chicks banded after 12 October, 16 were captured at one of
these two locations.

�-180Movements

to Wintering

Areas

The vegetative and environmental factors characteristic of white-tailed
ptarmigan wintering areas have been described by Braun (1969), Braun (1971),
Braun and Schmidt (1971), and Hoffman and Braun (1975). Segregation of
sexes usually occurs in winter flocks of ptarmigan, with males utilizing
higher, more windswept areas, while females prefer lower areas at or below
timberline.
The abundance and availability of willow is a prime consideration, as the buds and twig tips of willow comprise over 90 percent of the
winter diet of ptarmigan in Colorado (May and Braun 1972).
This study was not designed to monitor wintering populations of ptarmigan
in Rocky Mountain National Park. Due to lack of intensive effort and
inaccessibility of the study area, relatively few observations of ptarmigan
were made "in winter.
Sixteen subadult ptarmigan were observed on wintering
areas, with 12 having been previously marked.
Eight of the 12 had been
banded as chicks on summer areas, while 4 were banded during dispersal in
the fall. Hoffman (1974) designed a study with one objective being to
document chick movements to wintering areas.
In two consecutive summers
he banded a total of 126 chicks.
Excluding 16 chicks which were harvested
by hunters, only 2 of a possible 110 banded chicks were observed on wintering
areas.
Average distance moved to Wintering sites from summering areas was 1.37 km.
Movements of 12 chicks from fall concentration areas to wintering sites
averaged 3.31 km. Few conclusions can be based on these data since all but
2 observations were made at the one wintering site (Sundance Mountain Basin)
most accessible in winter.
Also, 4 of the observations came from a single
brood (G76) which remained intact until late November.
It is likely that
most ptarmigan winter off the study area and these areas were not searched.
It is also possible that juvenile ptarmigan are more mobile in winter than
previously thought and may move among several Wintering sites, especially
early in the wintering season.

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189 pp.

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___

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migan in Colorado.

1972. Seasonal foods of adult white-tailed ptarJ. Wildl. Manage. 36(4):1180-1186.

Odum, E. P., and E. J. Kuenzler. 1955. Measurement of teritory and home
range size in birds. Auk 72(1) :128-137.
Quam, L. O. 1938. The morphology of landscape of the Estes Park area,
Colorado. Ph. D. Dissertation, Clark Univ., Worcester, Mass. 165 pp.
Robel, R. J., J. N. Briggs, J. J. Cebula, N. J. Silvy, C. E. Viers, and
P. G. Watt. 1970. Greater prairie chicken ranges, movements, and
habitat usage in Kansas. J. Wildl. Manage. 34 (2):286-306.
Schmidt, R. K., Jr. 1969. Behavior of white-tailed ptarmigan in Colorado.
M. S. Thesis. Colorado State Univ., Ft. Collins. 174 pp.
Stiven , A. E. 1961. Food energy available for and required by the blue
grouse chick. Ecology 42(3):547-553.
Theberge, J. B., and G. C. West. 1973. Significance of brooding to the
energy demands of Alaskan rock ptarmigan chicks. Arctic 26(2) :138-148.
Walles tad, R. O. 1971. Summer movements and habitat use by sage grouse in
central Montana. J. Wildl. Manage. 35(1):129-136.
Weimer, R. J., and J. D. Haun (eds.). 1960. Guide to the geology of Colorado. Geol. Soc. Am., Rocky Mountain Assoc. Geol., Colorado Sci. Soc.,
Denver. 310 pp.
Willard, B. E. 1963. Phytosociology of the alpine tundra of Trail Ridge,
Rocky Mountain National Park, Colorado. Ph. D. Dissertation. Univ.
Colorado, Boulder. 245 pp.
Wing, L., J. Beer, and W. Tidyman.
grouse. Auk 61(3):426-444.

1944.

Brood habits and growth of blue

Zwickel, F. C. 1967. Some observations of weather and brood behavior in
blue grouse. J. Wildl. Manage. 31(3) :563-568.

�-183Zwickel, F. C., and J. F. Bendell. 1967a. Early mortality and the regulation of numbers in blue grouse. Can. J. Zool. 45(4) :817-851.
_____ , and
1967b. A snare for capturing blue grouse.
Manage. 31(1):202-204.

J. Wildl.

_____ , I. O. Buss, and J. H. Brigham. 1968. Autumn movements of blue
grouse and their relevance to populations and management. J. Wildl.
Manage. 32(3):456-468.

Prepared by

~1(----'

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1, i
)
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...• "

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__ ,_:_·o,_:.._'t.._''''-_'-''&lt;."..'~--:'/-:-'i_';''''_/

__

Kenneth M. Giesen
"...:'
Graduate Research Assistant

&lt;:

Approved bY

~(_~'_1~~/~~'_,~{·~~'_':_·-_i_·~_/'_,~~'_,~~_~_-~'_)~~Z'-,{_&lt;_:_/_1l_/_'
_

Clait E. Braun
Wildlife Researcher

'"L

��April 1976

-185JOB I'ROG!!.ESS REPO]\T

State of

COLORADO-'-----W-37-R-29

Proj ect No.
\~ork Plan No.
Job Title
Period

Job No.
1
Investigation of Popui~a-t-l~'o-n-------------------Status of Bobwhite Quail in Eastern Colorado
20

--------.----------------~-------.-------------------------

Covered:

Personnel:

Game Bird Survey

-----.--------~-------------

April 1, 1975 through March 31, 1976

Lucien Brevard, William
and Warren D. Snyder.

T. Howard,

Steve Steinert,

Larry Crooks,

ABSTRACT
Bobwhite quail whistling counts obtained on the Tamarack Wildlife Area in
June and July, 1975 were again found to accurately indicate forthcoming fall
population level and harvest.
Based on five years data a correlation
coefficient of r = 0.97 was obtained between census and fall population
level.
A similar coefficient of r = 0.99 was obtained between census and
harvest data on the Tamarack.
A Tamarack breeding population, believed to.
contain approximately 137 birds, survived the March 27, 1975 snowstorm.
This population was able, with excellent reproduction, to provide a fall
population approximating 650 quail by early fall. Of these, 239 were banded
or retrapped.
A short quail season, November 19. to December 7, was accompanied by the only major snow stress period of the winter.
Hunter success
was generally poor and the 15 percent harvest was the lowest recorded in
any year of study. Winter trapping and covey search efforts indicated
the combined hunting-snow stress situation had reduced the population to
an estimated 250 quail which survived the remainder of the winter in good
shape.
Again as in previous years, winter weather and snow stress conditions were the primary limitations to bobwhite survival.
Management census
information is included for trend comparison.
Assistance was provided to
Southeast Region personnel in conducting the 1975 bobwhite census along
the Arkansas River.
'

�-186RECOMMENDATIONS

1.

That a final publication in technical bulletin form be prepared for
dissemination of the findings of this study. Game Information Leaflets
should be prepared covering census procedures and basic habitat needs
with manipulation recommendations.
An article for the Colorado Outdoors
magazine should be prepared covering the species, environmental limitations and hunting.
Consideration should be given to a Journal of
Wildlife Management article covering population dynamics and census
results on the Tamarack.
Specific recommendations should be prepared
in writing for use in managing the Tamarack and other Division properties for increased survival and harvest of bobwhite.

2.

Census should be continued through the smnmer of 1976 on the Tamarack
by research personnel to supplement study findings to date.

3.

Assistance should be provided to management personnel in mid-October
monitor banding of at least 100 bobwhite on the Tamarack.

for

�-187-

INVESTIGATION OF POPULATION
STATUS OF BOBWHITE QUAIL IN EASTERN COLORADO
Warren

D. Snyder

P. S. OBJECTIVE
To investigate (1) the distribution and relative density of bobwhite quail
in eastern Colorado, and (2) population structure and level, and rate of
harvest, and identify key factors that limit bobwhite on the Tamarack
Management Area.
SEGMENT OBJECTIVES
1.

To investigate
Colorado.

the distribution

2.

To investigate population structure and level, and rate of harvest
bobwhite quail on the Tamarack Management Area.

3.

To identify
Area.

key factors

limiting

METHODS

and density

bobwhite

of bobwhite

in eastern

on the Tamarack

of

Management

AND MATERIALS

Methods and materials used in this investigation were listed by Snyder
1973,1974,
and 1975) in previous segment progress reports.

(1972,

RESULTS AND DISCUSSION
Bobwhite

Distribution

and Density

in Eastern

Colorado

1975 Census
Whistling counts were again illustrated to be an effective method of bobwhite
census and prediction of fall population level in 1975.
Counts completed on
the East Tamarack indicated some population increase over the previous year,
whereas the Hest Tamarack population appeared stable (Table 1).
Wildlife Conservation Officers, Warren Rupke, Clayton Wetherill, Rodd
Richardson, Larry Budde and Brett Peterson completed counts elsewhere along
the South Platte (Table 1). In general, population declines were evident,
although upriver from Sterling two routes showed increases over the previous
year.
The impact of a major March snowstorm will be covered in a later
section of this report.

�-188Table 1. A five year comparison of bobwhite quail whistling counts completed
along the South Platte River.

Percent Change From
Previous
4 Year
Year
Average

Route

Year

Counts

Stops

Calling
Males

Stateline-Ovid
(Warren Rupke)

1971
1972
1973
1974
1975

2
2
4
3
3

20
20
40
33
33

78
65
93
115
14

3.90
3.25
2.33
3.48
0.42

-87.9%

-87.0%

1971
1972
1973
1974
1975

2
4
6
5
6

20
45
64
54
55

67
202
135
65
94

3.35
4.49
2.10
1.20
1.71

+42.5

-38.7

1971
1972
1973
1974
1975

6
7
5
5
6

75
79
57
64
82

219
312
109
53
64

2.92
3.95
1.91
0.83
0.78

- 6.0

-67.5

1972
1973
1974
1975

2
2
4
3

19
20
41
30

63
73
133
61

3.33
3.65
3.24
2.03

-37.3

-40.4

1971
1972
1973
1974
1975

2
2
3
2
3

20
20
30
28
27

108
182
246
77
150

5.40
9.16
8.20
2.75
5.55

+101.8

-l3.0

1971
1972
1973
1974
1975

2
2
3
2
3

20
20
30
20
45

77
163
60
72
l33

3.85
8.15
2.00
3.60
2.96

-17.8

-32.7

1971
1972
1973
1974
1975

2
2
1
1
4

20
23
11
12
36

23
32
7
19
91

'1.15
1.39
0.64
1.58
2.53

+58.2

+110.0

1971
1972
1973
1974
1975

16
21
24
22
28

175
226
252
252
308

572
1019
723
534
607

3.27
4.51
2.87
2.12
1.97

- 7.1

-38.2

East Tamarack

West Tamarack

Sterling-Iliff
(C. Wetherill)

Atwood-Merino
CR. Richardson)

Brush-Ft. Morgan
(Larry Budde)

Goodrich-Orchard
(B. Petersen)

South Platte
Average

Average

�-189-

Census had not been conducted in the Arkansas Valley in 1974. At the
request of management personnel there assistance was provided in affirming
and adjusting routes and stations as needed in 1975.
Censuses conducted
by Wildlife Conservation Officers Gordon East, Dale Coven, Dan Potts, Jim
Dennis and Glen Eyre are summarized in Table 2, along with comparable data
from 1972 and 1973. Data presented in Table 2 show that bobwhite have
declined in number on the lower Arkansas east of LaJunta, but routes from
LaJunta upriver to Fowler contained good populations of quail.
The rangeland route in southeast Baca County declined.
Whistling

Counts as Indicators

of Fall Population

Level and Harvest

Whistling counts on the Tamarack showed close correlation with subsequent
fall densities (r = 0.97) (Fig. 1) and also with harvest (r = 0.99) (Fig. 2)
based on five years data. Although management personnel do not usually
complete as many replicates as conducted on the study area, the available
information illustrates that the whistling count can be a reliable method
for predicting fall population level and harvest potential.
It should therefore be consldered as an essential tool to use in the management of bobwhite
in Colorado.
The fall densities in Fig. 1 are presented in quail-per-square-mile
form
since the approximate acreage of quail habitat on the Tamarack was known.
This would facilitate projecting approximate popUlation levels along river
bottoms elsewhere in Colorado on a birds-per-square-mile
basis.
This method
should not be used in rangelands where quail are heard from greater distances,
often in all directions, rather than from a semicircle of river bottom
habitat.
Population
Rate of Harvest
Past Segment Band Recoveries,

Structure, Level and
on the Tamarack Wildlife
Mortality

Area

Rates and Population

Estimates

Previous Segment Band Recoveries.--Band
recoveries from quail banded in
preceding trapping periods are illustrated in Table 3. These few recoveries
do not provide enough information to merit revision of fall and winter population estimates previously presented (Snyder 1975).
Recoveries over a
longer time span are also subject to more bias to quail movement.
The high
mortality and relatively short life span of bobwhite is indicated in Table 3
by the rapid depletion of recovery information.
Fall, 1974 to Fall 1975 Mortality and Population Level.--The early fall
population level on the Tamarack in 1974 was placed at 499 bobwhite.
One
year later, 17.86 percent of 650 bobwhite were observed to be adults, or
there were approxiamtely 118 survivors from the original 499 birds (Table 4).
The loss of 381 quail from fall to fall represents an annual mortality rate
of 76.35 percent.
This projected rate assumes there was no significant
migration into or out of the study area during the interval.
Band return
data, presented later, do not give any clue that significant movement did
occur.

�-190Table 2. A summary of bobwhite whistling counts completed in southeast
Colorado from 1971 through 1975.

Year

Number of
Counts

Number of
Stops

Natural
Tally

Mean
No. Calls

E. Prowers County
(Dan Potts)

1971
1972
1973
1975

1
3
2
3

10
30
26
33

8
33
45
11

0.8
1.1
1.7
0.3

Purgatqire
(Dale Coven)

1971
1972
1973
1975

1
2
3
4

10
24
40
56

52
194
158
57

5.2
8.1
3.9
2.5

Las Animas-La
Junta
(Dale Coven)

1972
1973
1975

2
4
4

27
47
40

152
128
57

5.6
2.7
1.4

La JuntaRocky Ford
(Gordon East)

1972
1973
1975

2
3
3

23
37
35

76
87
124

3.3
2.3
3.5

Rocky FordManzanola

1972
1973

2
2

18
17

8
12

0.4
0.7

Manzanola-Fowler
(Gordon Eas t )

1975

3

31

133

4.3

Fowler-Boone

1972
1973

3
1

34
:1,.4

9
4

0.3
0.3

1971
1972
1973
1975

1
2
2
2

10
28
25
24

38
145
49
37

3.8
5.2
2.0
1.5

Location and Route

Arkansas River Valley

Cimarron River Valley
Southeast Baca
County
(Glen Eyre and
Jim Dennis)

�-191-

400

350

300

250

200

Y'

150

= 97.28

y = -23.88

+ 94.22x

100

50

o

1

2

3

BOBWHITE WHISTLING

C\LL

4
INDEX

Fig. 1. Relationship of the bobwh i.t e whistling call index to subsequent
fall density per square mile on the Tamarack Wildlife Area.

5

�-192700

600

.::::

~

0::

-c

500

t.J

C::..
H

0-4
0

--l

H

3

~ 400
u
-&lt;
~
s;

&lt;

8
,. .,
::r::
.LJ

8

300

z

0
8

Y'

(;1

W

&gt;
x

y =

-&lt;

::r::

w

= 99.43
105.1 + 180.4 x

200

E-t
H

::r:
~s

rn
0

en

100

o

1

2
BOBVJHITE

\A/HISTLING

3
C,\LL

4
INDEX

Fig. 2. Relationship of the bobwhite whistling call index to subsequent
fall harvest on the Tamarack Wildlife Area.

5

�-193Table 3. Number
previous banding

of quail band recoveries and retrap information from
periods obtained during 1975 and the winter of 1976.

Period Obtained

Period Banded

Number
Banded

Late
Winter
1975

Fall 1971 through
Win ter 1973

Fall
1975

Harvest
1975

Late
Winter
1976

No Bands .Recovered

Fall 1973

300

4

1

0

1

Winter

12

2

1

0

0

Fall 1974

194

38

5

7

4

Winter

46

6

2

5

1974

1975

The 76 percent annual mortality is a marked drop from an observed 87 percent
rate the preceding year and an approximate 94 percent rate in 1972-73.
Reduced 1974-75 mortality permitted an upswing of the fall population to occur.
Unfortunately,
two factors make it difficult to precisely differentiate the
mortality into fall-winter and spring-summer segments.
First, band recovery
samples were relatively small, and therefore, are subject to substantial
error when used in projection.
Second, a major snow storm occurred in late
March 1975 after winter banding had terminated.
Some quail mortality apparently occurred, but no accurate assessment of the storms impact could be
obtained at the time.
Two methods were used to project fall-winter mortality.
The first utilized
proportional band recoveries from fall 1974, and winter 1975 that were
obtained during fall, harvest, and winter 1975-76.
Only 13 bands were
recovered from the original 197 placed on quail in fall 1974, and 17 were
recovered from winter 1975, when 90 quail were trapped.
Proportional projections place the fall-winter (1974-75) natural mortality rate at 46.7
percent.
Addition of 18.27 percent harvest mortality brings the total fallwinter mortality to 65 percent of the original 499 quail.
This method projects
175 quail alive at the end of winter trapping.
The second method is based on the proportion of winter banded quail among the
total adults in subsequent 1975-76 fall-winter samples.
Forty-nine adults
were handled, of which 17 had been marked during the previous winter.
Projection based on this ratio placed the surviving late winter population at
259 quail for a natural fall-winter mortality rate of 29.46 percent.
Combined
natural and harvest mortality would approximate 48.10 percent.

�-194-

Table 4. Projected mortality of bobwhite quail on the Tamarack Wildlife
Area from October 1974 through October 1975.

West
Tamarack

East
Tamarack

Total
Area

Total Fall 1974 population estimate

246

253

499

Hunting season removal

59

34

93

Remainder subject to natural mortality

187

219

406

Winter 1975 popUlation estimate
(To midpoint of trapping)

119

140

259

Fall-early Winter natural mortality

68

79

147

Percent natural mortality

27.64

31.23

29.46

Total Fall-early Winter mortality

127

113

240

Percent total mortality

51.63

44.66

48.10

March 27 storm mortality

64

39

103

Total natural mortality to early Spring

53.66

46.64

50.10

Percent natural + harvest mortality

74.39

70.75

72.54

Early Spring population estimate

63

74

137

Percent Spring-Summer mortality

3.65

3.95

Fall 1975 popUlation estimate (adults)

54

64

118

Annual mortality rate (percent)

78.05

74.70

76.35

Item

3.81

The two methods of projection yield rather widely differing estimates of
fall-winter mortality and late winter population level. These discrepancies
were either the result of (1) inadequate sample size, or (2) egress of banded
quail and ingress of unbanded quail. The latter factor would cause the 259
quail projection to be in excess of the actual, but if only a few more fall
banded quail had been recovered the lower estimate of 175 quail would have
been revised upward dramatically.
Comparison of percentage banded among fall-winter (1974-75) quail with
survivors one year later shows about 48.1 percent of the original 499 quail
were banded by the end of winter. The ratio of banded to unbanded was 23 to
49, or 46.9 percent in the following year. This indicates little movement

�-195-

into or out of the study area occurred through the spring and summer of
1975 and gives credence to the projection method based on the proportion
winter banded among surviving adults.
Emperical observations, based on past trapping experience, placed the winter
1975 population at around 200 to 250 quail (Snyder 1975).
The fact that
ninety were handled indicates in excess of 200 were present since it is
difficult to trap in excess of 50 percent of a previously trapped winter
population.
In addition, weather through the fall and winter was relatively
dry and mild, permitting good survival as indicated by large covey sizes
observed during the trapping period.
Based on information presented in preceding paragraphs, 259 quail will be
used as the projection of late winter 1975 population level (Table 4).
Unfortunately, the March 27 snowstorm came after trapping had terminated.
Accurate assessment of its impact on the resident quail could not be made.
A few surviving quail were observed but it was extremely difficult to find
mortalities of present.
Hundreds of passerines were observed lost to the
storm along the riverbottom, and pheasants, crows, hawks, waterfowl and
other wildlife and livestock were lost in surrounding localities.
June-July whistling counts showed the quail population had taken a considerable drop in number by that time. Most of that decline can be realistically
attributed to the March 27 storm, so that overall winter mortality was conSiderably higher than that observed up to termination of winter trapping.
The whistling counts were previously illustrated to be fairly accurate
indicators of population level.
In 1974 a call index of 1.00 was based on
an early spring population of approximately 119 quail.
In direct proportion
a call index of 1.15 in 1975 would be based on a spring population of about
137 quail.
Respective surviving numbers of adults for fall 1974 and fall
1975 were 91 and 118. Assuming these projections were approximately accurate,
then our best estimates of March storm mortality was 100 to 120 quail or
about 20 to 24 percent of the original fall population.
Only about four
percent of the original population was lost during the late spring through
late summer interval.
It should be understood that the above information
and that presented in Table 4 are based on considerable conjecture and
should be considered only as our best estimates of mortality trend.
Fall Banding

1975

Warm, dry weather and inexperienced trapping personnel combined to slow
early fall trapping success.
Success picked up with the onset of colder
weather in mid-October so that 226 quail were banded and 13 previously banded
adults were retrapped to bring the total marked sample to 239 birds (Table 5).
Better access and trapping weather skewed banding success disproportionately
to the East Tamarack where harvest recoveries projected that 45.6 percent of
the population was marked.
In contrast,
only 26.8 percent of those on the
West Tamarack were banded for an overall average of 36.8 percent.

�-196-

Table 5. A summary of banding, harvest and population projections from
the Tamarack based on fall 1975 and winter 1976 data.

West
Tamarack

East
Tamarack

Total
Area

100

126

226

1

12

13

101

138

239

Fall 1975 banded

11

19

30

PreViously banded adults

o

2

2

11

21

32

Crippling loss (projected total)

3

6

9

Known harvest

41

46

87

Projected total harvest mortality

44

52

96

Percent crippling loss (of harvest)

7.32%

13.04%

10.34%

Projected total band recoveries

12

24

36

Percent banded (Fall and previous)

26.83%

45.65%

36.78%

Percent harvest mortality (includes
cripples)
Winter 1976 Banded

11.88%

17.39%

15.06%

Retrapped from Fall 1975

5

30

35

Other quail banded or retrapped

13

23

36

Total quail handled

18

53

71

Percent Fall banded

27 78%

Fall population estimate

376 (360)

302 (290)

650.!!

Finite confidence limits

+110.5

+ 89.4

+169.9

Item

Fall 1975 Banding Information
Number of quail banded
Number of retrapped adults
Total marked
Harvest Band Recoveries

Total band recoveries
Harvest Statistics

56.60%
Population Projection for Fall 1975 Based on Harvest Recovery Data
0

49.30%

1/ Separate computation of the East and West Tamarack data yields a combined
total of 678 quail in contrast to a total area calculation of 650 quail.
The latter figure is considered most accurate.

�-197-

Fall Movement
As in 1974, there was no evidence of bobwhite movement off the Tamarack,
either upstream or downstream.
Relatively low quail densities and a shortened time lapse between trapping and quail season were believed responsible.
As a consequence, corrections to population estimates based on movement
were not applied in Table 5.
Fall Population

Estimate

Increased trapping success in fall, 1975 was indicative of an increase in
quail abundance over the previous fall. The fall population estimate
derived from the banded-unbanded
ratio in harvest confirms this increase.
This was also the first time the annual mortality rate had dropped below
80 percent since the 1971-72 interval.
Win ter 1976 trapping data were excluded in calculating the fall population
estimate of 650 quail (Table 5) because a large proportion of that sample
was obtained along the East Tamarack meadow where fall trapping also had
been concentrated.
Although sample size would have been increased, bias
due to poorly distributed sampling was believed an overrriding factor.
Mark and recapture samples were not sufficient to prevent errors greater
than 10 percent with 95 percent accuracy as recommended by Robson and
Regier (1964). Confidence limits for the populat ion estimate are provided
in Table 5.
The 1975 Hunting

Season

Drought conditions in 1974 followed by the devastating snowstorm of March
27, 1975 reduced pheasant populations in the Tablelands of extreme northeastern Colorado to near zero. As a result~ Division management personnel
delayed the opening of pheasant and quail season to sunrise on Wednesday,
November 19 in Small Game Management Unit 2, which includes the Tamarack
and lower South Platte River.
The season opened at noon on Saturday,
November 15 elsewhere in the state.
In Unit 2 the pheasant season terminated Sunday, November 30, whereas quail season continued through December 7,
1975. This season provided 19 days for quail hunting (Table 6).
A snowstorm enveloped the region during the season opening, depositing 8
to 10 inches of snow. Despite the snow, a few hunters went after quail on
the opening day of season, but most hunting occurred during the first weekend, November 22 and 23. The heavy snow conditions persisted through the
first two weeks of season.
Warming temperatures during the last week of
quail season reduced snow cover on the ground to partial coverage by season
end.
Runting Pressure and Hunting Success.--During the first eight days of the
1975 season, an average of 55 hunters per day checked into the Tamarack.
This compares to 86 hunters per day in 1974 and 103 hunters per day in 1973
(Table 7). Hunter use of the area held at a fairly high level so that the
average use per day for the season was not too far below hunter use rates
in previous years.

�-198-

Table 6.

Bobwhite

fall population and harvest
Area.

statistics

for 1971 through

1975 on the Tamarack Wildlife

1971

1972

1973

1974

1975

1,745

2,319

722

499

650

Item

Fall population

estimate

20.5

31.5

32.5

29. sl/

19

Known harvest

411

548

202

73

87

Projected total harvest
mortality '1:../

472

634

227

93

96

Percen t h arvest morta I"lty-2/

27 .0

26.7

31. 7

18.3

15.1

Season length

(days)

1/ Quail

and pheasant hunting terminated at 2:00 P.M. from November
through December 15, 1974 on the Tamarack.

29

2/
Figures

include projected

crippling

loss based on hunter

questioning.

Hunter success generally reflected quail abundance with 10.03 hunters using
the area per quail bagged (Table 7). It must be remembered not all were
quail hunters, and since most hunters were opportunists, there was no accurate way to segregate quail hunters from waterfowl and pheasant hunters.
The
heavy snow conditions made it difficult to walk long distances in pursuit
of quail, but it also drastically reduced the amount of survival and escape
cover available to the birds.
Harvest of 87 quail, plus reported crippling loss projected an approximate
total harvest mortality of 96 bobwhite.
This was only slightly above the
harvest level of 1974 where 73 quail were taken, with projected mortality
totaling 93 birds (Table 6). Harvest mortality, including projected crippling loss, represented only about 15 percent of the original fall popUlation
and represents the lowest harvest rate of any year of study (Table 6). A
high crippling loss (27 percent of reported harvest) was indicated by hunters
in 1974 when warm, dry conditions made it difficult for dogs and men to
locate downed birds.
The much reduced 1975 crippling loss was possibly due
to ease of locating quail in the snow.
Total species harvest is summarized in Table 8, and compared there with the
take of previous years.
Whereas, quail harvest was slightly higher, pheasant
take was approximately twice that of the preceding year.
Factors causing
this include better reproduction in 1975 and snow conditions during the season
which made it easier to find birds.
Pheasant survival was probably much
better in the heavy cover along the river bottom during the March storm, in
contrast to high storm mortality in the adjacent tablelands where cover is
sparse.

�-199Table 7. Hunter success ratios on bobwhite quail with the progression of
hunting seasons from 1971 through 1975.

Interval

Number of!/
Hunters

Average
Hunter Use

No. of Quail
Harvested

Hunters per
Quail Harvested

1971 Season
First 8 days

638

79.75

258

2.47

Second 8 days

363

45.38

100

3.63

Last 5 days

258

51.60

53

4.87

Season,Average

59.95

3.06

1972 Season
First 8 days

606

75.75

230

2.63

Second 8 days

386

48.25

167

2.31

Third 8 days

369

46.12

100

3.69

Fourth 8 days

392

49.00

36

10.89

Season Average

54.78

3.25

1973 Season
First 8 days

823

102.87

142

5.80

Second 8 days

339

42.38

36

9.42

Third 8 days

161

20.12

10

16.10

Fourth 9 days

303

33.67

l3

23.30

Season Average

49.27

8.09

1974 Season
First 8 days

686

85.75

43

15.95

Second 8 days

414

51. 75

17

24.35

Third 8 days

138

17.25

9

15.33

Last 6 days

176

29.33

4

44.00

Season Average

47.13

19.37

1975 Season
First 8 days

440

55.00

52

8.46

Second 8 days

310

38.75

17

18.23

Last 3 days

123

41.00

18

6.83

Season Average

45.95

11 Includes all hunters using the Tamarack.
hunters late in the season were after ducks.

10.03
During 1972 and 1973 most

�-200The snow conditions during the 1975 season also increased waterfowl and
cottontail hunting success, whereas the colder weather put fox squirrels
into their dens early. These variables are reflected in the harvest figures
in Table 8.

Table 8. A comparison of small game harvest on the Tamarack Wildlife Area
during the 1971 through 1975 quail and pheasant seasons.
Year

Season Interval

Quail

Pheasants

Ducks

Rabbits

Squirrels

1971

20.5 days

411

87

196

30

9

1st 20.5 days

480

l30

555

126

4

1972

31.5 days

548

143

704

156

4

1973

1st 20.5 days

184

53

283

40

3

1973

32.5 days

202

62

441

63

3

1974

1st 20.5 days

64

49

69

19

11

1974

29.5 days

73

141

23

15

1975

19 days

86

340

38

6

1972

1/ Pheasant season in 1975 was shortened to 12 days.

Hunter Use and Success by Hunting Area.--Table 9 summarizes hunter numbers,
hunter hours, quail banded in fall, and banded and total quail harvest by
hunting area. Banding effort was well distributed along the Tamarack, but
banded quail per area on the East Tamarack exceeded that on the west by
almost a 3 to 1 ratio. Harvest too, was well distributed among the hunting
areas with about 17 percent harvest mortality on the east section and about
12 percent harvest on the west. The east areas sustained 134 hours of
hunting pressure per area, compared to 102 hours per area on the West
Tamarack. It should be remembered that although quail harvest was usually
reported by hunters as the area to which they were assigned, actual harvest
may have occurred in some proximal area. However, check station personnel
frequently were able to assign harvest to the proper location by questioning
hunters.
Win ter Banding 1976
Bobwhite banding commenced in early February 1976, and during a brief interval
of cold weather, was quite successful. Trapping success slowed drastically
through the remainder of February under relatively warm dry weather conditions.
Thirty bobwhite were banded on the Tamarack and 41 were retrapped, bringing
the total to 71 quail handled (Table 5). Most, 53 of the total, were taken
on the East Tamarack where several coveys were found along the meadow. Only
18 birds were taken on the larger West Tamarack.

�-201Table 9. A summary by hunting area of fall banded bobwhite, hunter use, total
harvest and banded harvest recoveries on the Tamarack Wildlife Area in 1974.

Area

No. Quail
Banded

No. of!/
Hunters

Hunte~1
Hours

Total Quai1~1
Harvest

Banded Quai121
Harvest

West Tamarack
1

27

89

6

1

2
3
4

22
32

75
75

6

1

10

42

80

1
10

18
16
21
12

6
5

7
11

4

42

11

25
34
49

5
6
7

8
9

10
11
12
13
14
15
18 41
19
20
21
22
23

17

27

1

42
27

1
12

34
25
32
20
23
16
604

5

24

13

25
Subtotal

5

101

77

2
4
2

42
70
30
26
33
149
86
158
191
110
160
119
205
144
134
85
58
62
2,258

2
2
1
1

3
1

1

5
2
1

5
2

3

1

41

11

3
4

2

East Tamarack
Braddock
Pasture
1
2
3
4
5
6

7
8
9

2
9

14
10
13
5
8
22
16
23

41
28
22
21
31

98
122
90

29

97
38
58
43

78

14
19
18
28
28
31

57
66
75

10

1

11

Subtotal

15
138

269

Total

239

873

5
9

1
2
2

11
3
2

1
5
1

2
1
4
3
1
1

822

4
46

21

3,080

87

32

11 Includes only known hunters and hours reported through the check station.
II Includes only known harvest. Some hunting probably went unrecorded.
1/ Approximates the location where quail were taken. Exact area is unknown.
!!../ Braddock Pasture and areas 16 and 17 west were closed to public hunting.

�-202-

Considerable effort was directed toward covey searches since trapping efforts
produced marginal results.
Only a few coveys were found on the West Tararack
(Fig. 3), although harvest recoveries projected a population exceeding that
found on the east section in fall. Lack of tracking snow made covey
searching results uncertain.
It is also possible that quail on the west
part of the study area suffered higher combined hunter-snow stress mortality
during the fall hun~ing season.
Food plots were not available to the West
Tamarack quail, whereas, a majority of the East Tamarack quail were found
proximal to and using available food plots.
Summer whistling counts should
provide some evidence as to whether differential mortality occurred between
the two segments of the study area.
Age Ratios - Fall to Winter

1975-76

Age ratio information obtained during the work segment is summarized in Table
10. Based on the harvest sample, in excess of nine young per pair of adults
were present in the fall. The percentage adults, 17.86, was the second
lowest percentage recorded during the five year study, however, the proportion adults has not deviated significantly anyone
year.
From age ratio
information obtained for the past several years, reproduction capability
does not appear to be a limiting factor to bobwhite on the Tamarack.
Fall
population numbers have been in direct proportion to the number of breeding
quail present in spring.
Sex Ratios

According to information summarized in Table 11 males dominated the population
during the fall and winter of 1974-75 and this sex ratio continued relatively
stable through the summer.
Dominance of males in past years showed up after
the reproduction season (Snyder 1972), however, a dominance of males among
young in 1974 was observed.
Males and females were near equal in number
among young {n the fall 1975 popUlation (Table 11).
Factors
Fall and Winter

Limiting

Bobwhite

on the Tamarack

Mortality

Weather Related Mortality.--Winter
continued to appear as the primary limitation to bobwhite survival and increase in northeastern Colorado.
The snow
stress relationship to winter mortality for the first four years of study is
summarized in Table 12. A direct relationship between snow cover and winter
mortality was indicated.
But, all the variables of snow stress cannot be
condensed into tabular form for correlation.
To illustrate, above average
survival was evident through the winter of 1974-75 up to the end of quail
trapping, and the potential for a much improved breeding popUlation was present. The snowstorm of March 27, 1975 which followed, caused death to numerous quail by exposure due to accompanying high winds, rather than by prolonged snow cover occurrence on the ground.
Snow depths were not excessive
and most of the snow was melted within a few days.

�'rJ
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.
.

()Q

TAMA RACK MANAGEMENT

w

AREA

t;:I

a
0-

~
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n-

(1)

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a

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(1)
"&lt;:

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Ul

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f-'

a
o

OJ
r+
(1)
0..
0..
I::!
Ii

•....

tv

::J

()Q

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f-'

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:;::

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r+
(1)

EAST

Ii

AREAS

n-

o

Ii

~

'1:j

•....
::J

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(1)

LEGEND
1

HUNTING AREA

0

COVEY

LOCATION

t-n
t-n

~UNFARMED
RIVER BOTTOM ~ALFALFA
~
AND RIVER CHANNELS
~

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~GRASSLAND

a

r+

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Ul

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SC ALE
y.

Mi.

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�-204Table 10. Bobwhite age ratio data obtained
during the fall and winter of
1975-76 on the Tamarack Wildlife Area.

Sample

Number
Adults

Number
Young

Total

Percent
Adults

Adult:Young
Ratio

Fall Banded

26

213

239

10.88

1:8.19

Banded

2

30

32

6.25

Unbanded

l3

39

52

25.00

Total

15

69

84

17.86

1:4.60

Winter Banded

14

57

71

19.71

1:4.07

Hunting Season

Table 11. Sex ratios of bobwhite quail from fall-winter 1974-75 to fallwinter 1975-76 intervals.
Item

Male

Female

Total

Harvest and late winter 1974-75

88

68

156

Percent

56.41

43.59

Surviving adults, fall-winter 1975-76

28

21

Percent

57.14

42.86

Fall 1975 banded juveniles

106

106

212

Harvest 1975 juveniles

34

35

69

Winter 1976 juveniles

32

25

57

Combined total of juveniles

172

166

338

Percent of total juveniles

50.88

49.12

49

�-205-

Table 12. A comparison of fall-winter natural mortality,
and subsequent call index change from the previous year.

snow conditions,

Fall-Winter
1972-73

Interval
1973-74

1974-75

11

52

50

25

Days with snow cover

14

76

Days with 4" or more of snow

o

37

30

9

16.3

60.0

52.2

50.1!:,,/

+37.5

-51. 3

-50.5

Item

1971-72

Total snowfall

Fall-Wint~r

(inches)

mortality

ratell

Percent change in subsequent
call index from previous year

II Does not include

harvest

29

+15.0

mortality.

11 Includes

mortality resulting from late March blizzard.
up to that time was projected at 29 percent.

Natural mortality

Hunting as an Additive Stress.--A deep snow in November 1975 coincided with
the start of quail season so that resident populations were simultaneously
subject to prolonged deep snow, severe cold weather, and hunting stress.
The impact of the combined stresses is as yet unknown, but is believed to
be greater than if the limitations had occurred at separate intervals.
Survival of remaining quail appeared satisfactory with at least 250 quail
coming through the winter.
Forthcoming censuses will provide additional
evidence of population survival through the fall and winter of 1975-76.
Table 13 compares bobwhite census on the Tamarack with that in adjacent areas
through 1975.
Greater population decline on the Tamarack was observed to
follow years when hunting and snow stress were in unison.
With no combined
snow-hunting stress in fall 1974, the subsequent census index was proportionately higher on the Tamarack than in adjacent areas (Table 13).
Spring and Summer Mortality

and Reproduction

Lack of change in adult sex ratios (Table 11) is one index of excellent
survival of the breeding population through summer in 1975.
During both 1972
and 1973, sex ratios changed from spring to fall in favor of males, and these
were also the years when summer mortality of adults was highest.
Several factors probably combined to reduce summer stress on the breeding
population.
The population was low so competition for nesting and other
cover was not significant.
Severe thunderstorms, hail and flooding did not
occur on the study area.
It is also possible that extensive removal of
raccoons by trapping and hunting with dogs during the previous fall and
winter reduced predator stress on bobwhite through the summer months.

�-206-

Table 13. A comparison of bobwhite whistling counts on the Tamarack
Wildlife Area with counts located up and down river.

Year

Routes UE and Down River-1/
Number
X Count/
Percent
Stops
Stop
Change

1972

79

5.99

1973

120

3.93

1974

122

1975

135

Number
Stops

Routes on Tamarack
X Count/
Percent
Stop
Change

124

4.15

-34.4

121

2.02

-51.3

3.25

-17.3

118

1.00

-50.5

2.65

-18.5

137

1.15

+15.0

!/ Includes average of Brush-Ft. Morgan, Atwood-Merino, Sterling-Iliff and
Ovid-Stateline routes.
Projected Period and Peak of Hatch
A sample of 210 quail trapped in fall provided evidence that the primary
quail hatch occurred from mid-June to early July (Fig. 4). This was approximately two weeks later than in 1974, but ahead of the main hatching periods
of 1972 and 1973. Unlike the 1974 hatch, which declined rapidly after an
early peak, the 1975 hatch was sustained at a higher level from mid-July to
early September. Precipitation approximated 14 inches in 1975 compared to
only 7.18 in 1974 in the study area.
Quail Utilization of Cover
Vegetative cover on a large proportion of the Tamarack was rated by ocular
inspection during the winter of 1975-76. Shrub, grass and forb abundance
was sytematically sampled and rated on a scale of one to five. Comparison
of site results with covey occurrence in winter has not been completed.
Results of this evaluation, along with more detailed presentation of the
overall limiting factors to bobwhite on the Tamarack will be forthcoming in
the final publication during the next work segment.

�-207-

34
32 io
30
Sample Size

28

N =

26

= 288 in 1973
N = 177 in 1974
N = 210 in 1975
N

r:t
0\
r-i

24
22
'0

768 in 1972

20

aJ
u
+l
co 18 ••

.c

:r:

....•
co

.j.J

16

,

o
E-t
lH

o

14

+l

§ 12
u

~

0.

10
8

6

,,...--

4
2 I-

o If'
18

/

,,

l

/
(XX:
0\,

25
May

2

9

17
June

24

3
Weekly

11

19
July

27

4

12
20
August

28

Mid-Points

~ig. 4. Period and peak of bobwhite quail hatch projected from primary
feather growth for 1972 through 1975 on a percent of total basis.

7 14
Sept.

�-208-

LITERATURE CITED

Robson, D. S., and H. A. Regier. 1964. Sample sizes in Peterson markrecapture experiments. Trans. Amer. Fish Society 93(3) :215-226.
Snyder, W. D. 1972. Population status of bobwhite quail in eastern Colorado. Colo. Div. of Game, Fish and Parks. Game Res. Rept. April.
pp. 121-150.
1973. Population status of bobwhite quail in eastern Colorado.
Colo. Div. of Wildl., Game Res. Rept. April. pp. 253-296.
1974. Population status of bobwhite quail in eastern Colorado.
Colo. Div. ofWildl., Game Res. Rept. April. pp.119-l47.
1975. Population status of bobwhite quail in eastern Colorado.
Colo. Div. of Wildl., Game Res. Rept. April. pp. 115-137.

Prepared by

_~7d-=~=---::~~~'7---"~~~--~-~-)1&lt;---Warren D. SnYder~
Wildlife Researcher

�-209JOB PROGRESS

State of
Project No.

April 1976

nEPORT

COLORJ,DO
W-37-R-29

---------

110rk Plan No.

.---------

Game Bird Survey

Job Title

Job No.
1
Minimum Tillage Techniques
For Establishing Shrubs in Clump Plantings

Period Covered:

April 1, 1975 through March 31, 1976

Personnel:

21

David C. Bowden, Lucien Brevard, Daniel Walsworth,
William T. Howard, and Warren D. Snyder.

Larry Crooks,

ABSTRACT
Shrub plots were established on three additional sites during April, 1975
bringing the total sites to nine within three soil types. Precipitation
approximated 14 inches in 1975, or about double the 7.14 inches recorded in
1974. This additional moisture greatly increased shrub survival and growth.
Within the six sites planted in April 1974, 73.4 percent of the Hansen rose
and 85.0 percent of the wild plum remained alive in Spring 1975. Almost no
mortality occurred among these shrubs through the 1975 growing season.
Oversummer survival within the three sites planted in April 1975 exceeded 98
percent for both species, although some increased mortality was observed
following the fall-winter period.
None of the herbicides or mulch treatments
showed evidence of h av ing significant impact on survival through the 1975 growing
season.
Some rabbit and deer damage to wild plums occurred on loam and clay
loam sites during a fall 1975 snow stress period.
Among sites established
in 1974, spring and summer 1975 growth approximated 20 inches of height
increase on the Karg 1, Karg 2, and Tamarack Meadow sites. Less growth was
recorded on the two sandhill sites. A severe iron sclerosis condition
reduced growth and survival on the Smith Meadow site, and this and other
biases resulted in the exclusion of this site from growth analysis.
Greatest
growth increase occurred on the plastic mulch and organic mulch plots.
Herbicide treatments did not show evidence of decreasing shrub growth. The
plastic mulch treatment significantly increased growth of both shrub species
over the other three treatments within the three sites planted in April 1975.
Late May-early June sampling of annual weeds showed rather high weed densities
on the cultivated control plots within both the 1974 and 1975 established
sites. All herbicide and mulch treatments significantly reduced weed
competition in comparison to the cultivated controls, however, there was no
significant difference in weed control effect among these treatments.

��-211-

MINIMUM TILLAGE TECHNIQUES
FOR ESTABLISHING SHRUBS IN CLUMP PLANTINGS
Warren D. Snyder

P. S. OBJECTIVE
To evaluate ground cover techniques, pre-emergent herbicide techniques, a
combination of the two, and cultivation techniques for establishing shrubs
in clumpplantings
in eastern Colorado.
SEGMENT OBJECTIVES
1. To prepare and establish one additional site within each of the
three site types to supplement 1974 plantings that were adversely affected by
drought and other mortality factors.
2. To apply pre-emergent herbicide treatments, mulch tre-atments, and
a combination of the two, to six of the seven plots within each planting site.
The remaining plot within each site will be retained as a cultivated control.
3. To measure and evaluate shrub survival,
competition among treatments and sites.
METHODS

shrub growth and weed

AND MATERIALS

Initial methods and materials concerning planting, site location and other
procedures were summarized by Snyder (1975). Three additional sites were
planted in April 1975.
Site location and treatment sequence were illustrated
by Snyder (1975) and are listed in Table 1. In summary, three sites have
been selected in each of three soil types. Within each site, seven treatment plots were established, each of which initially contained 32 wild plums
and 32 Hansen roses spaced at three foot intervals.
Chemical
include:

applications

applied

in 1975 to the six sites established

in 1974

1. Simazine (Princep) was applied at the rate of 1.2 pounds per acre on
loamy sand soils and 2.0 pounds per acre on other soils.
Date of application
was April 5, 1975 to two plots within each site (Table 1). Less than one
inch of organic mulch in the form of wood chips and/or corn cobs was applied
on the Simazine mulch plots on May 15 and 16, 1975. No soil incorporation was
attempted.
Application was made using a hand sprayer modified to contain two
nozzles for even distribution of the chemical between shrub rows.
2. Granular G-4 Casoron was applied with a hand spreader to one plot within
each of the six sites (Table 1) on April 14, 1975. Application rate approximated 150 pounds per acre. No soil incorporation was conducted.

�-212Table 1. Reference sequence of test shrub clump plots within
1974 p1antings, and (2) 1975 plantings.

Plot
Number

Sandhills
Red Lion
Sharptail

River Bottom
Tamarack
Smith Meadow

sites for (1)

Tableland
Karg #1
Karg #2

1974
1

organlc-. 1/

Plastic

Control

Plastic

Ep-Tref

Sim-M

2

Plastic

Organic

Plastic

Organic

Simazine

Simazine

3

Control

Control

Organic

Control

Sim-M

Casoron

4

Ep-Tref

Simazine

Casoron

Casoron

Casoron

Ep-Tref

5

Simazine

Ep-Tref

Simazine

Ep-Tref

Organic

Plastic

6

Sim-M

Casoron

Sim-M

Sim-M

Plastic

Control

7

Casoron

Sim-M

Ep-Tref

Simazine

Control

Organic

E-W

NE-SW

NE-SW

E-W

N-S

NE-SW

Sequence
Direction

---------------------------------------------------------------------------------I-76

Sonnenburg

Karg #3

1975
1

Organic

Sim-M

Plastic

2

Plastic

Ep-Tref

Organic

3

Control

Simazine

Control

4

Simazine

Casoron

Casoron

5

Sim-M

Control

Ep-Tref

6

Ep-Tref

Plastic

Simazine

7

Casoron

Organic

Sim-M

E-W

NE-SW

NE-SW

Sequence
Direction

1/ Organic
herbicides

and Plastic are mulch treatments, Ep-Tref represents Eptam-Treflan
and Sim-M represents Simazine overlaid with Organic mulch.

�-2133. Eptam and Treflan pre-emergent herbicides were applied as a mixture
using the modified two nozzle hand sprayer on April 23 and 24, 1975.
Shallow soil incorporation with a garden rototiller was conducted immediately
to each site. Eptam was applied at the rate of two pounds per acre (active
ingredient) and Treflan was applied at the rate of four pounds per acre.
4. Iron sulfate was applied to all plots within the Smith Meadow site at
the rate of about 140 pounds per acre in efforts to alleviate a severe iron
sclerosis condition.
Application was made using granular iron sulfate on
June 25, 1975. A second treatment was made in March 1976.
5. Repeat applications of Casoron and Eptam-Treflan were completed in late
March 1976. Application rates were the same as those shown in numbers 2
and 3 above.
Organic mulch and plastic mulch, with its organic overlay were applied in
late May 1975 to the 1-76 site. The Sonnenburg and Karg 3 sites received
early June mulch treatments.
Competing vegetation was tallied as annual or perennial.
A sample scheme,
illustrated in Fig. 1, utilized a square foot sampling hoop placed systematically on the nine open strips adjacent to the eight shrub rows within each
plot. Four samples were obtained within each strip to yield a 36 square
foot, or approximate 5 percent sample within each plot. Annual and perennial
forbs and grasses were tallied by species within each sample.
After sampling
competing vegetation was removed by shallow hoeing as needed on the chemical
plots, by hand pulling on the mulch plots, and by use of a garden type rototiller and hoe on cultivated plots.
Mid- to late summer vegetative sampling was done by tally of individual plants
per plot rather than by use of a sampling method since forb densities were
sparse and most competing vegetation was perennial in growth form.
Height and survival measurements of shrubs were obtained in mid-May with a
second survival check in early July. Measurements were again obtained in
late summer after termination of growth.
Spot applic~tions of 2-4,D Amine based contact herbicide were used to control
bindweed, Canadian thistle,
and whitetop on the Tamarack and Smith Meadow
sites.
A hand sprayer and paint brush were used in December 1975 to apply rodent
and deer repellent to wild plums receiving damage from rabbits on the riverbottom and tableland soil sites.
Efforts to trap pocket gophers on the
sandy soil sites were generally unsuccessful but no damage to Hansen rose
shrubs was noted during the winter of 1975-76.
RESULTS AND DISCUSSION
Environmental

Measurements

Soil Characteristics
Soil characteris tics of the nine si tes containing shrub plots are summarized
in Table 2. Characteristics of the six sites established in 1974 were
repeated here for comparison with data on the three new sites.

�-214-

0+

+

+

o

0 +

+-

0+

0 ...
+ 0+
0 +

+

+

+

D t-

D +

+-

+

0+

o *"

+

0 +

+-

0+

+

+

1-

0

...
0+

0+

...0 ...
0 +

-t-

t-

0 +

...0 ...

+

+

+

0 +

0 ...

0+

+-

.,.0

0 .•.
+0

T

-t-

0+

+

+

D -+

T

0 +

0+

+-

0+

+

Dt-.
+0
0 +

Fig. 1
Method used in systematic sampling of forb and grass
vegetation the the shrub clump plots.

c=J Square Foot Sample

+ Planted Shrub

--

Plot Boundary

�-215Table 2.

Soil analyses

Location
Site

Year
Es tablished

Level

Red Lion

1974

6.2

1.7

2.3

76

Sharptail

1974

6.0

1.6

1.7

1-76

1975

6.4

0.5

Karg 1

1974

7.6

Karg 2

1974

Karg 3

completed

pH

on the nine shrub clump sites.

Analzsis
Sol Salts
Cnunhos/cm) Org. Mat.

1/

Percent
Sand Silt

Clay

Texture

16

8

Sandy
loam

82

11

7

Loamy
sand

0.9

80

13

7

Loamy
sand

1.1

3.2

48

30

22

Loam

6.9

1.5

5.4

46

35

19

Loam

1975

7.4

0.4

1.7

56

27

17

Sandy
loam

Tamarack

1974

7.7

3.4

6.3

22

41

37

Clay
loam

Sm. Meadow

1974

7.6

3.3

7.6

22

39

39

Clay
loam

Sonnenburg

1975

7.7

2.6

2.3

43

27

30

Clay
loam

Sandhills

Tableland

Riverbottom

1/ Soil analyses
University,

were completed by the Soils Laboratory
Fort Collins.

at Colorado

State

The three main locational groupings - sandhills, tableland and riverbottom
show distinctive soil differences.
Riverbottom soils all contained high Ph,
high soluble salts, and high clay levels.
The Sonnenburg site apparently
contained a higher sand level with less silt than the other sites and is
believed
to be closely underlaid by sand. In contrast, the Smith Meadow
site is underlaid by clay which is believed a primary factor in the iron
sclerosis condition among shrubs there.
Seepage from neighboring irrigation

�-216ditches apparently affects the west end of this site more than the east end.
This is shown by better summer growth among plants in the west part of the
site.
Although Red Lion and Karg 3 sites were both classed as having a sandy loam
texture (Table 2), the latter contained 20 percent less sand. All Karg sites
lie within a quarter mile radius and all have similarshortgrass
origin and
characteristics.
In future study reference, the sandhill sites will be
called loamy sand soils, the Karg sites will be considered predominately
loam in texture, and the riverbottoms sites will be referred to as clay loarns.
Precipitation
The severe drought year of 1974, which severely hampered shrub establishment,
was followed by a more normal year for precipitation in 1975 (Table 3). A
precipitation
recorder, located at the Tamarack headquarters, showed about
14.10 inches of moisture for 1975, compared to only 7.18 inches in 1974.
U. S. Weather stations near Sedgwick and Sterling, respectively recorded
16.91 and 13.64 inches of moisture in 1975.
Amounts received on the shrub
sites also varied to some degree, but recording devices could not be located
at all sites.
The increased precipitation
in 1975 significantly increased
survival and growth of original and newly established plots alike as shown
in the following section.
Shrub Establishment

Survival Within

Sites Established

and Survival

in 1974

First Year Survival 1974-75--An approximate 10 to 15 percent mortality of
the wild plum seedlings and about 20 to 25 percent mortality of Hansen rose
shrubs was sustained during the 1974 growing season, as reported by Snyder
(1975).
In addition, burrowing rodents destroyed about 31 percent of the
Hansen rose shrubs in the Red Lion site, and 14 percent of the roses in the
Sharptail site over winter in 1974-75.
Plums were not seriously damaged.
Rodent damage to shrubs was serious only in the two sandhill sites.
Efforts
to determine survival of remaining shrubs were hampered during
the spring
of 1975 because some plants resprouted from roots, so that a final assessment of annual survival was not completed until early July.
It showed that
overall, 73.4 percent of the Hansen rose and 85.0 percent of the wild plum
remained alive (Table 4). This was higher than expected, considering the
severe drought conditions which persisted throughout 1974 and early 1975
(Snyder 1975).
The two mulch plots within the Red Lion site were excluded
in the annual survival figures since nearly all plants had died. These two
plots were replanted in April 1975.
Shrub Survival during the 1975 Growing Season--The brief period from early
July to mid-September 1975 does not give a complete picture of mortality
during the growing season, but it does provide a general index of summer
stress.
Losses of only one to two percent were evident during this brief
interval.
Moisture conditions were much improved over those in the 1974
growing season.

�-217Table 3. Precipitation
during 1974 and 1975.

received

at the Tamarack

Management

Area headquarters

Month

1974

1975

January

0.25

0.05

February

0.67

0.15

March

0.91

1.25

April

0.45

0.65

May

0.70

5.55

June

2.05

2.00

July

1.15

0.55

August

0.40

1. 60

September

0.15

0.55

October

0.00

0.00

November

0.25

1.20

December

0.20

0.55

Annual Total

7.18

14.10

Treatment Impact on Survival of Plots Established in 1974--None of the
treatments, including the spring, 1975 applications of Simazine, Simazine
mulch, Casoron and Eptam-Treflan (2nd year) appeared to have a significant
impact on survival during the 1975 growing season.
All four chemical plots
per site had received a pre-planting treatment with Eptam-Treflan in 1974
without evidence of increased shrub mortality.
However, both the Simazine
and Casoron appear to present a greater threat since both are much more
persistent chemicals.
Neither is recommended on light sandy soils where
rapid penetration to the root zone of the shrubs is a threat.
One wild plum was believed killed by Simazine on the Karg 1 site, however,
all other shrubs in the plot showed no discernable treatment effect.
Whether this mortality was due to a spot chemical overdose or to other
factors remains unknown.
Some leaf burning of plums within the Simazine and Simazine-mulch plots on
the Sharptail site was observed, but no actual mortality was noted.
Additional chemical treatment effects, especially on the sandy sites, may show
up in the future.

�-218Table 4. Shrub survival among shrub clump treatments from Fall 1974
through September 1975.

Treatment

Fall
1974

Rose
Spring
1975

Fall
1975

Fall
1974

Plum
Spring
1975

Fall
1975

CuItivated

73.4

65.1

65.6

92.7

88.0

88.5

Organic mulch

68.1

68.7

66.9

83.1

80.0

80.0

Plastic mulch

87.5

87.5

82.5

93.8

91.9

92.5

S.lTIlaz
. 1/
lne--.

79.7

73.4

74.0

87.0

84.4

83.9

Simazine mulch

79.7

74.0

75.5

91.7

89.6

86.4

Ep tam-Tre flan

81.2

73.4

73.9

85.4

82.8

82.8

Casoron

84.4

72.9

71.7

87.5

79.2

78.6

Overall Mean

79.2

73.4

72.7

88.8

85.0

84.6

l/

All chemical plots were treated alike with an Eptam-Treflan mixture in 1974.

Both the organic mulch and plastic-organic mulch plots retained excellent
shrub survival in 1975. This was in direct contrast to the previous year
when increased mortality was expecially evident on the organic mulch sites
due to treatment over dry soil.
Shrub Survival of 1975 Established Plantings
Site Effect--Shrubs planted in April 1975 on the Karg 3, Sonnenburg and I-76
sites overcame planting shock and showed excellent survival during their
first growing season (Table 5). The only shrubs showing stress at the end
of summer were Hansen rose on unmulched plots within the sandy soil of the
I-76 site. High mortality was not evident, but some top growth die-back
occurred in late summer. An overall 98+ percent survival rate on the three
sites can only be considered as excellent.
Treatment Effect--There was no evidence of treatment effect during the first
growing season on survival. First year treatments included cultivation,
organic mulch, plastic mulch overlaid with organic mulch, and Eptam-Treflan
pre-emergent herbicide treatment of all four chemical plots per site. This
lack of treatment effect is in direct contrast to 1974 when organic mulch
plots contained higher shrub mortality than other treatments. The difference

�-219apparently lies in root establishment due to moisture condition.
In 1974
only light rains were received for nearly two months after planting (Table
3). The organic mulch tended to absorb much of this, preventing the new
root systems from becoming established.
Increased mortality resulted.
In
1975 mulching was delayed until root systems could become established.
In
addition, near normal precipitation prevailed in 1975.

Table 5. Shrub survival
established sites.

1/
Treatment-

from planting

Karg 3
Hansen
Wild
Rose
Plum

in April to Fall 1975 on three newly

Percent Survival
Sonnenburg
Hansen
Wild
Rose
Plum

1-76 2}
Hansen
Wild
Rose
Plum

Cultivated

100

100

96.9

100

100

93.8

Organic mulch

100

100

100

100

96.9

100

Plastic mulch

100

100

100

100

100

100

Simazine

100

100

100

93.8

96.9

96.9

Simazine mulch

100

100

93.8

100

100

93.8

Eptam-Treflan

100

100

93.8

100

100

100

Casoron

100

100

100

93.8

96.9

96.9

Mean

100

100

97.8

98.2

98.7

97.3

98.8

98.5

Overall Mean
1/
- All four chemical plots received
April 1975.
~/ Survival of several of the Hansen
the time of fall observation.

Shrub Survival

the same Eptam-Treflan

treatment

in

rose on the 1-76 site was uncertain

at

- All Sites During the Fall and Winter of 1975-76

All plots in all sites were believed to have retained good to excellent
survival through the fall and winter of 1975-76, although actual mortality
remains unknown at this time. A heavy snow deposited moisture on November 19
which melted in early December.
Several additional light snows deposited
small amounts of moisture through the winter.

�-220Rabbit damage occurred
Karg 3 sites, although
degree.
Repellent was
snows or rabbit damage

primarily on the newly established Sonnenburg and
Karg 1 and 2 sites also were damaged to a lesser
applied in mid- to late December.
No other deep
was observed through the remainder of the winter.

Hansen rose shrubs sustained little damage overwinter.
Those in the sandhills sites were closely observed following the rodent destruction of root
systems during the previous winter.
No evidence of damage reoccurrence
was noted.

Sh rub Growth
Growth on Sites Established

in 1974

The shrub plots established in April 1974 attained no significant growth
during th~ir first year of existence because of transplant shock and drought
conditions.
In addition, only four treatments, cultivation, organic mulch,
plastic mulch and Eptam-Treflan were used during the first growing season.
The latter was applied equally to all four chemical plots.
Much improved
moisture conditions in 1975 resulted in significant recovery and growth.
Overall growth probably would have been even better if the shrubs had not
come into the 1975 growing season in such poor condition.
Many had to
resprout from surviving roots or basal areas of the plants.
Exclusion

of the Smith Meadow

Site From Analysis

Soil tests showed the Smith Meadow site to be very similar to the Tamarack
Meadow in pH, salts level, and texture (Table 2). However, some basic
differences led to the exclusion of Smith Meadow shrub growth data from
analysis.
Iron sclerosis, which was evident there in 1974, reached a critical level in Spring 1975.
Little new growth was attained and some mortality
occurred on the site in spite of adequate spring and summer moisture conditions.
Granular iron sulfate was applied in late June 1975 to all plots
within the site, but treatment was too late to have much impact in 1975.
The heavy clay soils underlying the site may be a factor responsible for
the iron deficiency condition persisting there.
There was also evidence of a graduated increase in shrub growth on the Smith
Meadow site from east to west.
This bias was believed due to subsurface
seepage moisture which supplied the west end of the site with more soil
moisture than the east end received.
The latter is at a slightly higher
level and therefore seepage water does not approach the surface as readily.
Interpretation of Shrub Growth Data--Preceding
analysis showed that treatment effects were very similar for both shrub species.
Using the cultivated
treatment as the standard or control, both plastic and organic treatments
increased shrub growth.
Use of Simazine, Casoron, and the Eptam-Treflan
mixture had no discernable impact on growth.
Weeds were not permitted to become highly competitive for moisture within the
cultivated plots.
This would not always be the situation where large scale
shrub establishment was conducted or if private landowners were responsible
for weed control.
If weeds were permitted to become more competitive, treatment of these sites with herbicides might achieve greater growth in comparison to straight cultivation maintenance.

�-221Growth Analyses per Treatment Within Site Within Soil--Wild plums averaged
approximately 20 inches of growth during the spring and summer of 1975 on
the Karg sites and on the Tamarack Meadow site (Table 6). Hansen rose,
which sent out more basal stems, did not average as great a height increase,
but may have put out equal growth.
The Red Lion and Sharptail sites in the
sandhills averaged less growth (Table 6). This was potentially attributable
to poorer soil quality.
The same factor, soil quality, may also account for
the suppressed growth on the Sharptail site in relation to that on the Red
Lion site. According to soil analyses (Table 2) the soil in the Sharptail
site contained about 82 percent sand, whereas, that in the Red Lion site
was composed of 76 percent sand.
Dr. David C. Bowden conducted statistical analyses of 1975 growth per observation data among shrubs established in 1974. As previously mentioned, the
Smith Meadow site was excluded from this analysis.
As a consequence, the
Tamarack Meadow data, within the same soil type, were also excluded in
initial tests for interaction.
This reduced the number of soils to two, loam
and loamy sand, with two sites per soil and seven treatments per site.
Wild Plum--Preliminary
analysis indicated treatments did not significantly alter wild plum growth among the four sites (F6 10 = 3.06) at the 95
percent confidence level.
A second test, ignoring soiis effect and including
the Tamarack Meadow data, showed a significant difference among treatments
did exist (F6,22 = 3.44) at the 95 percent confidence level.
Fig. 2 indicates the only significant differences among treatment growth rate means
was between plastic mulch and Simazine treatments.
The latter was significantly lower.
Fig. 2 also shows that average growth per shrub was near
equal between the plastic and organic mulch sites, whereas, the chemical
treated plot means and the cultivated control mean was at a lower level.
Hansen Rose--Dr. Bowden found that Hansen rose growth per observation
data differed significantly depending on soil type (F6 ,10 = 4.88), however,
treatments averaged over differential soil effects were still found to be
significant (F6 ,10 = 23.90).
As illustrated in Fig. 3, both plastic and
organic mulch treatments yielded significantly greater Hansen rose growth.
Again, as with the plums, the chemical plots and the cultivated plot mean
growth .rates were closely grouped with no significant differences among
means.
Within loamy sand soils, the plastic mulch treatment yielded significantly
greater rose growth than other treatments with the exception of organic mulch
(F6,4 = 14.95) (Fig. 4). Organic and plastic mulch plots within the Red Lion
site had sustained very high mortality during the preceding year, and as a
consequence, had been replanted in April 1975. Prorated means based on
results in the Sharptail site were substituted in the analysis.
Therefore,
the data in Fig. 4 are considered weak and high confidence cannot be placed
in the results.
Hansen's rose, recelvlng the straight organic treatment, achieved significantly
greater growth than all other treatments, except plastic mulch within the
loam soil site (F6,6 = 11.01) (Fig. 5). Means of the chemical plots and the
cultivated plot were again closely grouped with no significant differences
evident.

�Table 6.

Average

Treatment

shrub growth

per plot and treatment

Loam
Karg 1

Karg 2

during

the 1975 growing

Sites Within Soils
Clay Loam
Tamarack
Smith
Neadow.!.i
Nanagement

season

on the six original

Loamy Sand
Red Lion
Sharptail

sites.

Treatment
He an

Wild Plum
Cultivated

22.55

17.55

17.80

4.00

15.94

3.58

14.37

Organic

mulch

21.84

22.10

24.22

6.61

17.58

mulch

19.47

18.59

26.42

6.56

-

10.14

Plastic

14.43

16.66

17.93

16.42

20.21

9.35

12.10

4.85

13.28

mulch

19.58

25.20

19.04

6.24

15.13

3.79

14.75

Ept am-Trefl an

19.27

20.85

20.50

4.04

13.07

5.00

13.34

Casoron

17.35

15.04

23.28

4.00

12.75

5.86

13.86

Site He an

19.73

19.38

21.58

5.96

13.84

6.79

14.74

Simazine
Simazine

Hansen

Rose

I

Cultivated

12.48

13.35

12.75

5.20

10.18

2.20

9.71

Organic mulch

20.04

17.50

17.13

0.27

-

8.21

14.57

Plastic

16.13

16.57

18.30

2.52

-

13.22

13.79

11.55

12.20

10.48

9.30

8.50

3.75

9.29

mulch

13.27

13.21

11.14

14.91

11.41

1.57

11. 72

Ep tam-Tref Ian

13.67

13.31

14.88

9.11

10.54

3.90

10.92

Casoron

12.74

9.55

12.61

3.50

11.92

6.00

9.86

Site He an

14.16

13.71

14.06

6.94

13.27

5.63

11.30

mulch

Simazine
Simazine

1./
Smith Neadow
biases.

data excluded

from statistical

analysis

I

N
N
N

due to persisting

iron sclerosis

condition

and other

�-223-

WILD PLUM--GROWTH PER OBSERVATION

p

o

I'\.)

o

IV
IV

n
c

I-'

IT

•

0

1"1

.o

·
I

:s:
c:

I-'
()

zr
-o

I-'

OJ
(Jl

•I

3:

c:

I-'
()

rr
1-3
::::l
(T]

»
1-3

:s:

t'l
Z
1-3

~.
3
Ul

OJ
N

~.
::s
(1)

~.
3
Ul

·
OJ

I
3:

c

I-'
()

zr

M
'd

IT

•I

1-3

1"1
(1)

'"h

•

o

OJ

(Jl

0

1"1

0

::s

Fig. 2. Average wild plum growth in 1975 per treatment (center mark)
with a standard deviation of 5.83 (vertical line) based on data from
five shrub sites established in 1974.

�-224-

HANSENS

U1

ROSE--GROWfH

PER OBSERVATION

•...
U1

n

c
•.....
rf"

•

0

'"1

&lt;0

•I

:s:

c
•.....
o
:J
'1J
•.....

OJ
III

•I

:s:
C

•.....

n

:J

1-3

::0

Ul

~~
M

:s:

M
Z
1-3

OJ

N

/-'-

:J

&lt;D

Ul
•....

3
OJ

•I
:s:

C
•.....

o
:J
M
'0
rf"

•I

1-3
'"1
&lt;D
'il

n
OJ

III

0
'"1
0

:J
Fig. 3. Average Hansen rose growth in 1975 per treatment (center
mark) with a standard deviation of 3.76 (vertical
line) based on
data from five shrub sites established
in 1974.

�-225-

HANSENS ROSE-LOAMY

I

w

I

~

SAND--GROWTH

PER OBSERVATION

N
W

n
c
~
IT

•

0

~
~

•I

3

C

~
0
~
~
~
~
~

·
I

8

~
M
~

8
3

~
~
0
~
~
~
3
~
N

~ ~.
Z ~
~ ro
~

~.
3

~

•I

3
C

~
~

0

M

~IT

•I

8

~
ro
~

•

n
~
~
0
~
0
~

Fig. 4. Average Hansen rose growth in 1975 per treatment (center
mark) with a standard deviation of 7.64 (vertical line) based on
data from loamy sand sites established in 1974.

�-226-

HANSENS ROSE-LOAM--GROWTH

~

w

PER OBSERVATION

~

w

()

c
•.....
IT

•

0

11
\Q

•I

3:

C
•.....

o
:r
'U
•.....
OJ
V'l

•I

3:

c
•.....
o
:r
t-'3
::&lt;:J

t'1

Ul
,..,-

3

~

OJ
N

Z

~

3:
t'1
t-'3

,..,rD

L,)

f-'.
3
OJ

•I

3:

C
•.....
()

zr
t'1

"0
r+

•I

t-'3
t;
(l)
H'\

•
()

OJ
V'l

0

11

0

~
Fig. 5. Average Hansen rose growth in 1975 per treatment (center
mark) with a standard deviation of 4.8 (vertical line) based on
data from loam sites established in 1974.

�-227Growth on Sites Established

in 1975

Shrubs in the Karg 3, Sonnenburg and 1-76 sites, all established in April 1975,
made excellent recovery from planting shock and most attained fair growth in
1975.
Average growth per plot and site for both species is illustrated in
Table 7. All four chemical plots received the same treatment, an Eptam-Treflan
mixture, prior to planting.
Growth data for 1975 from the three sites was combined, ignoring potential site
or soil effect, and tested by analysis of variance.
Four treatments, cultivation, organic mulch, plastic mulch and Eptam-Treflan were compared as to their
impact on growth.
For wild plum shrubs a significantly different growth rate
among treatments was obtained (F6,665 = 20.3).
A similar result (F6,658 =
12.24) was observed for growth of Hansen rose. Using Student-Newman-Keuls'
test (Steel and Torrie 1960) the plastic mulch treatment was observed to significantly increase growth over all other treatments.
There were no significant
differences in growth means among the remaining treatments.
As noted in Table
7, organic mulch growth rate of both shrubs was second to plastic mulch and not
too much greater than growth on the chemical and cultivated plots.
Late summer observations indicated plastic mulch treated shrubs were still
growing throughout the summer, whereas the other shrubs had stopped growth much
earlier.
Shrubs within the mulched plots on the 1-76 site were greener and
healthier in appearance than those on the bare soil treatments.
There was also
evidence on the 1-76 site that over winter 1975-76 survival was much better
within the organic and plastic treated sites.
Weed Competition
Sites Established

in 1974

Annual Weed Occurrence--Analysis
of variance tests were completed by David C.
Bowden on treatment effect on annual weeds.
Results indicated that all chemical and mulch treatments significantly reduced annual weed competition in comparison to the cultivated control (F6,18 = 6.62).
There was no significant
difference, however, among any of the chemical and mulch treatments (F5,15 =
1.44).
The average occurrence of annual weeds per square foot sample per treatment is illustrated in Table 8.
Ocular estimates of spring weed density on the plots was obtained and are
presented in Table 9. These show, as the previously presented analysis did,
that cultivated plots contained much higher annual weed occurrence than any
of the other treatments.
Initial May-early June sampling of weed competition, summarized in Tables 8 and
9, provided the most reliable data covering treatment effect on annual weed
occurrence.
A July 28, 1975 sample of total weed reoccurrence is presented
in Table 10. These mid-summer data are biased because rototilling, used on
the cultivated plots, was more effective in weed removal than hoeing or hand
pulling used on the other plots.
Shallow hoeing often removed only part of
the plant so that it resprouted and continued to grow. The data in Table 10
reflect this bias, but information is presented to illustrate the low weed
density on the plots in mid-summer.
No significant annual weed germination
was observed after the July 28 removal on any site.

�-228Table 7. Average first year shrub growth per site and treatment on the
three sites established in April 1975.

Treatment

Karg 3

Sonnenburg

I-76

Treatment
Mean

Wild Plum
Cultivated

6.34

1.66

4.53

4.17

Organic mulch

8.97

0.50

5.90

5.12

Plastic Mulch

13.25

8.06

10.13

10.47

S.a.mazi..n e+1/

5.44

4.33

5.26

Simazine mul ch

3.31

2.16

2.10

Ept am-Tref Ian

4.56

3.00

5.03

Casoron

6.38

4.80

2.06

Site Mean

6.89

3.48

5.04

5.15

4.04

Hansen Rose
Cultivated

10.00

0.00

2.09

4.75

Organic mulch

10.19

0.75

4.38

5.61

Plastic mulch

12.81

4.31

9.26

9.18

S.amazi.ne+
. 1/

7.06

2.94

0.94

Simazine mul ch

6.50

2.13

2.03

Eptam-Treflan

7.16

2.43

1.63

Casoron

8.38

3.84

2.00

Site Mean

8.87

2.35

3.18

3.95

4.82

1/ Simazine, Simazine mulch and Casoron plots all received the EptamTreflan pre-emergent herbicide treatment prior to their first growing
season. Listed treatments were applied prior to the second growing
season.

�Table 8. Average occurrence of annual weeds per square foot sample in 1975 within the six sites and
three soil types established in 1974.

Treatment

Sandhills
Red Lion
Sharptail

Riverbot tom
Tamarack
Smith
Management
Meadow

Tableland
Karg 1
Karg 2

Overall
Mean

Cultivated control

30.20

29.81

119.00

27.78

12.11

24.28

29.29

Organic mulch

1.28

0.64

2.69

0.17

0.06

0.39

0.87

Plastic mulch

0.03

0.72

1.00

0.33

0.08

0.11

0.38

Simazine

0.72

1. 31

8.58

0.00

0.52

1.47

2.10

mulch

2.36

1.55

22.80

0.00

0.61

5.39

5.45

Eptam-Treflan

0.33

0.58

0.42

1.08

2.58

2.78

1.15

Casoron

6.28

1.72

21.69

2.69

0.72

0.28

5.56

Site Mean

3.09

5.19

14.38

4.29

2.27

4.96

5.59

Sd.rnaz Ln-a

I

N
N

'-I'&gt;

�-233-

Perennial Weed Occurrence in the Shrub Plots--The chemical and mulch
treatrM:.ntsacted primarily to prevent weed seed f rom gerruinating and were
not highly e f f ec t Lvs. against perennials whose root systems were already
present.
Preferably, all sites should have been summer fallowed for one or
two years to destroy existing root systems, and sites should have been
placed where noxious perennials were not abundant.
Fall and winter site
selection made this impossible to determine.
Noxious perennials were confined to the riverbottom clay-loam sites where
Canadian thistle, white top, field bindweed, saltgrass, and Nebraska sedge
were the dominate species.
Salmon mallow and skeleton daisy were most prevalent on the loam sites.
Sandsage, perennial ragweed and several grasses were
most common on the loamy sand sites.
P~rennials were not a significant problem in either the loam or loamy sand sites in 1975, but did require some
control. ~umbers per square foot obtained during late spring 1975 sampling
are illustrated in Table 12. Data are from the six sites established in 1974.
Table 12. Average perennial weed occurrence per square
foot sample within
the three soil type on sites established in 1974.
Treatment

Loamy Sand

Loam

Clay Loam

Overall

Cultivated

1.28

0.33

6.83

2.42

Organic mulch

0.19

0.00

1.11

0.44

Plastic mulch

0.00

0.00

0.25

o. 08

Simazine

0.46

0.22

1. 76

0.81

Simazine mulch

0.54

0.26

2.76

1.19

Eptam-Treflan

0.32

0.24

2.99

1.13

Casoron

0.60

0.06

1.50

0.72

Sites Established

in 1975

Results of mid-June sampling of annual weed occurrence in plots established
in 1975 are summarized in Table 13. All three sites showed similar results
with no annual weeds on the recently treated organic and plastic mulch plots,
moderate weed densities on the Eptam-Treflan treated plots and much higher
densities on the cultivated control plots.
Use of the Eptam-Treflan herbicide mixture significantly reduced weed abundance as shown by simple Chi
square analysis of overall mean densities in Table 13 (Chi square = 5.70
with 1 d.f.). A meaningful comparison could not be made with the two mulch
plots but they did remain weed free through the remainder of the summer.

X

�-234-

Table 13. Annual weed occurrence
clump sites.

in mid-June

1975 on the three new shrub

Annual Weed Occurrence Eer Sguare
1-76
Sonnenb ur g

Treatment

Cultivation

12.33

10.92

Organic mulch

No weeds present

Plastic

No weeds present

mulch
1. 87

Eptam-Treflan
(Four plot mean)

4.53

Foot Sample
Karg 3

Overall
Mean

lO.44

11.23

3.33

3.24

Percent ground cover by annual weeds was obtained using ocular estimate and
is shown in Part 2 of Table 9. Although densities were not much greater on
the Sonnenburg site than on the Karg 3 and 1-76 sites (Table 13), more rapid
weed growth produced much higher percentage ground cover as shown in Table
9. Due to biased efforts at weed control, further mid- and late summer
sampling of weed depsities were not attempted.

LITERATURE

Snyder, W. D. 1975.
clump plantings.

CITED

Minimum tillage techniques for establishing shrubs in
Colorado Div. of Wildl. Game Res. Rept. April:141-153.

Steel, R.G.D., and J. H. Torrie.
1960. Principles
statistics.
McGraw-Hill, New York.
481 pp.

and procedures

of

...

-:1..
Prepared

by

.

)

i"

/}

:

__' _ ·J_!·~~~f~~~~V~~=;_/~
~(~(~-.~~~/~(~.(~'~:~i/~2~.t~;~1~:-;~~~.~.~.,~
Warren D. Snyder
Wildlife Researcher

I

/

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                  <text>July, 1976

-1-

JOB PROGRESS REPORT

State of

COLORADO
------~~~--------

Project No.

W-lOl-R-lB

Work Plan No.

4

Job Title

Game Range Investigations
Job No.

Inventory of Range Manipulation

la
-----------------------

Projects in Colorado

Period Covered: April 1, 1975 through March 31, 1976
Personnel:

Roland C. Kufeld and Regional Game Biologists.

ABSTRACT
An inventory was made of all range vegetation-modification
projects
completed during 1974 in the western half of Colorado, on lands administered by the u.s. Forest Service and Bureau of Land Management.
Acreages
treated were: Forest Service - 0 and Bureau of Land Management - 3,277.

��-3-

INVENTORY OF RANGE MANIPULATION PROJECTS IN COLORADO
Roland C. Kufeld

P. S. OBJECTIVE
To collect information which describes proposed and completed range
vegetation modification projects that are located in Colorado, and
their effects on the range, wildlife and livestock; and to provide
desired IBM listings of these data to cooperating agencies upon their
request.

SEGMENT OBJECTIVES
1.

To collect information which describes proposed and completed range
vegetation modification projects that are located in Colorado, and
their effects on the range, wildlife and livestock.

2.

To compile, codify, process and analyze inventory data.

3.

To provide desired IBM listings of inventory data to cooperating
agencies upon their request.

METHODS AND MATERIALS
An inventory was made of all range vegetation modification projects
completed through 1972, in Colorado, west of Interstate Highway 25 on
lands administered by the U.S. Forest Service and Bureau of Land Management (Kufeld 1968, 1970, 1971, 1973, 1974 and 1975). Kufeld (1968 and
1970) also covered projects completed on the Southern Ute and Ute Mountain
Indian Reservations through 1969. Indian Reservation projects were
dropped from the inventory after 1969. This report concerns vegetation
modification projects completed on U.S. Forest Service and U.S. Bureau
of Land Management lands west of Interstate Highway 25 during 1974.
Data were collected llsing procedures outlined in Colorado Division of
Wildlife Administrative Directive No. 204 entitled "Range Vegetation
Modification Projects", and by Kufeld (1968, 1970 and 1974). Inventory
data were transferred from original data sheets to Mark Page Reader Forms,
and then to IBM cards.
Procedures outlined in Administrative Directive No. 204, call for evaluations to be made on each vegetation modification project at the end of
the 2nd, 5th, and 10th years following treatment. During 1974, 2 year
post-treatment evaluations were made on projects completed in 1972, and
5 year post-treatment evaluations were made on projects completed in 1969.
Data were recorded on a form described by Kufeld (1971).

�-4-

Table 1. Acreages of range land treated during 1974
in Colorado by
the U.S. Bureau of Land Management 1/.
Vegetation
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Sagebrush

Plow

2

780

780

Pinyon-juniper

Chain

2

2,497

2,497

4

3,277

3,277

All Vegetative Types

l/All projects were located on the Montrose BLM District.

Prepared by

;e-h.t~cclc))d~~It:/'
; Roland C. Kufeld
Wildlife Researcher

c-

�-5-

RESULTS AND DISCUSSION
Acreages treated during 1974, on all lands administered by the U.S.
Bureau of Land Management west of Interstate Highway 25, are shown
by vegetation type and kind of treatment in Table 1. All of the
BLM projects completed in Colorado during 1974 were located on the
Montrose BLM District. No vegetation modification projects were
completed on U.S. Forest Service lands in Colorado during 1974.
A total of 3,277 acres were treated on BLM lands in 1974. To date
(through Dec. 31, 1974) a total of 587,759 acres have been treated
in Colorado west of 1-25 on lands administered by the U.S. Forest
Service and Bureau of Land Management. Of this 244,975 acres were
on Forest Service and 342,784 acres were on Bureau of Land Management lands.
Two and 5 year post-treatment evaluations have been made on some
vegetation modification projects since 1970. Data from these evaluations will be kept on file and covered in a future report when enough
2 and 5 year post-treatment evaluations have been made to provide a
basis for a meaningful data analysis.

LITERATURE CITED
Kufeld, Roland C. 1968. Inventory of range manipulation projects in
Colorado. Colo. Div. Game, Fish and Parks. Game Res. Rept.
P.R. Proj. W-lOl-R-lO, WP 4, Jla, July, Part 1. p. 1-121.
1970. Inventory of range manipulation projects in Colorado.
Colo. Div. Game, Fish and Parks. Game Res. Rept. P.R. Proj.
W-10l-R-12, WP 4, Jla, July, Part 1. p. 59-94.
1971. Inventory of range manipulation projects in Colorado.
Colo. Div. Game, Fish and Parks. Game Res. Rept. P.R. Proj.
W-10l-R-13, WP 4, Jla, July, Part 1. p. 1-15.
1973. Inventory of range manipulation projects in Colorado.
Colo. Div. Wildl. Game Res. Rept. P.R. Proj. W-10l-R-15, WP 4,
Jla, July, Part 1. p. 1-8.
1974. Inventory of range manipulation projects in Colorado.
Colo. Div. Wildl. Game Res. Rept. P.R. Proj. W-10l-R-16, WP 4,
Jla, July, Part 1. p. 1-11.
1975. Inventory of range manipulation projects in Colorado.
Colo. Div. Wildl. Game Res. Rept. P.R. Proj. W-10l-R-17, WP 4,
Jla, July, Part 1. p. 1-6.

��-7-

July, 1976

JOB FINAL REPORT

State of __
Project

C_O_L_O_RA_D_O

_
Game Range Investigations

No. W-lOl-R-18

Job No •.

4

Hork Plan No.

2

----------------------------------

Experimental.Impro~ement
ofnOakbrush
k
Job Title __0_n D e_e_r
__a_n
__d__~_1
__
k_W_1_n_t_e_r
__K_a_n~g~e_s
__- e_a_v_e_r
cr__e_e
Period

Covered:

Personnel:

April 1, 1975 through March 31, 1976

Roland C. Kufeld

P. S. OBJECTIVE
To determine if deer and elk forage production and game use can be increased on overage Gambel oakbrush winter game ranges by spraying with
2,4,5-TP to induce sprouting.

CURRENT STATUS
A manuscript containing the final results of the study was prepared during
the 18th project segment, and has been accepted tor publication in the
Journal of Range Management.

Prepared

by

;e~«tI C %'-el~
Roland C. Kufeld
Wildlife Researcher

_

��July, 19.76

-9JOB PROGRESS REPORT

State of

COLORADO
------~~~~---------W-lOl-R-18

Project No.

Game Range Investigations

4
3
Job No.
Experimental Improvement of Oa-~k~b~r-u~s~h-------------------------on Deer, Elk and Cattle Ranges - Hightower Mountain

Work Plan No.
Job Title

Period Covered:
Personnel:

April 1, 1975 through March 31, 1976

Roland C. Kufeld

P. S. OBJECTIVE
To determine the extent to which deer, elk and cattle forage production
and game use can be" increased and maintained by chaining, spraying and
controlled burning on overage Gambel oak winter game ranges.

METHODS AND MATERIALS
The study plan for this job calls for post-treatment evaluations to be
made after 2, 5 and 8 years following treatment. The2-year evaluation
was made during the summer of 1973. The 5-year evaluation will be made
during the summer of 1976. There was no activity on the study during
the 18th project segment.

Prepared

by

~,tud

C AfJb/

Roland C. Kufeld
Wildlife Researcher

��-11-

July, 1976

JOB PROGRESS REPORT

State of

.:::.C.:::,O=LORAD::::.=:::..;O=-_

Project No.

W-lOl-R-18

Work Plan No.

6

Job Title

Game Management

Period Covered:
Personnel:

Game Range Investigations
Job No.

l

Unit Inventory

April 1, 1975 - March 31, 1976

William T. McKean and Bruce D. Trindle

ABSTRACT
Broad descriptive and wildlife species management information were
compiled for Wildlife Management Units 11 (Strawberry Creek) and 12
(Williams Fork). Subjects discussed included:
Unit Description
(Boundaries, Size, Physical Features and Climate); Landownership Status;
Land Use Status; Human Populations (by County); Wildlife Species Checklists; Hunting Pressure, Harvests and Seasons; Narrative and Map Descriptions of Game Species Distribution and Abundance; Introductions of Game
Species; Census Areas and Results; Habitat Restoration Projects; Management Problems Checklist; and Pertinent Research References by Species
and Author.

_

�•

�-13-

GAME MANAGEMENT UNIT INVENTORY
William T. McKean
Bruce D. Trindle

P. S. OBJECTIVE
To compile broad descriptive and wildlife species management
tion by wildlife management unit.

informa-

SEGMENT OBJECTIVES
1.

Describe boundaries, areal extent, physical features, and climate
of Wildlife Management Units 11 and 12.

2.

Inventory by unit the landownership
human population.

3.

Inventory by unit the wildlife characteristics including species
lists, distribution and abundance, harvest data, introduction records,
census areas and routes, and prepare appropriate research and management bibliographies.

4.

Inventory habitat restoration projects from existing records.

5.

Inventory present and future management problems.

status, land use status, and

METHODS AND MATERIALS
To produce the desired information
were taken:

for Units 11 and 12 the following steps

1.

Working with sources of information now available in the Fort Collins
and Denver offices, as much data as possible concerning each item listed
under objectives above were assembled and condensed into the standard
format used for units previously completed. This included seasonal
distribution maps of game species.

2.

At a meeting in Meeker, W.C.O.'s Glen Smith and Ron Krager reviewed
the preliminary copy and made revisions to game species distribution
maps. W.C.O. Bill Roland reviewed the same data at a separate meeting
in Craig.

3.

After the above corrections were entered, copies were circulated among
management personnel in the Denver and Grand Junction offices for review.

4.

Final revisions to maps and narratives were made and all information
gathered into file binder folders, except for the 1/2" per mile distri-·
bution maps which were reproduced by hand and bound in a separate
folder. Three copies of all this information were prepared and personally delivered to: (a) the Northwest Regional Office, (b) the Denver

�-14-

Office, (c) the Federal Aid Section. Distribution maps at 1/4"
per mile scale were prepared on Forest Service base maps cut to
8" x 11" size. Xeroxed copies of these were inserted into the file
binder folders. Additional xeroxed copies of these maps and of the
narrative were made. These were delivered to the local W.C.O.'s,
Bureau of Land Management, and Forest Service personnel. A few
additional copies have been xeroxed for general distribution as
requested.

RESULTS AND DISCUSSION
Wildlife Management Units 11 (Strawberry) and 12 (Williams Fork) were
completed during the project year in spite of interruptions in the work
plans necessitated by my reassignment to part-time work on the proposed
reclamation project, Yellow Jacket, of the U. S. Bureau of Reclamation.
Much of the information compiled for Units 11 and 12 is directly
applicable to needs of the Yellow Jacket Project. Also, they should be
useful to persons currently writing impact statements concerning coal
developments in these areas.
During the work segment there have been 25 requests for copies of these
unit reports. These have come from: U. S. Forest Service, U. S. Bureau
of Land Management, U. S. Fish and Wildlife Service, U. S. Bureau of
Reclamation, Atlantic-Richfield eompany, Ecological Consultants Inc.,
and private individuals.

,

Prepared by

'ir~Tl-;f.

}/1'C/CeCl~'l./-

William T. McKean
Wildlife Researcher

�-15-

APPENDIX A

WILDLIFE MANAGEHENT UNIT 11 (STRAWBERRY)
Moffat and Rio Blanco Counties, Colorado
Information on Unit Description, Landownersh~p
Land Use, Human Population, Wildlife Species
Checklists, Harvests, Seasons, Narrative
and Hap Descriptions of Distribution and
Abundance, and other Allied Data.
To: December 1975.
Compiled by: W.T. McKean and B.D. Trindle

�-16-

�-17TABLE OF CONTENTS,
ITEM
UNIT DESCRIPTION
LANDOW~~RSHIP

(boundaries,

size, physical

features,

HUMAN POPULATION

_
~

(Rio Blanco County,

(Big game mammals,

OTHER AVIAN SPECIES

1960-1970,

("Varmint"

"varmint"

birds, nongame

_

and projections)

small game mammals

OTHER MANHALIAN SPECIES (Furbearers,
mammals) -------------

nongame

(Antelope,

AND ABUNDANCE OF S~MLL GAME ~lliLS
hare, red (pine) squirrel) --

HARVEST,

DEER HARVEST,

SEASONS,

AND HUNTING

SEASONS AND HUNTING

LINEAR REGRESSION

_

black bear,

(cottontail

PRI~rrTIVE DEER HARVEST
TOTAL DEER HARVEST,

ARCHERY

MOUNTAIN

DATA ANALYSIS

ELK HARVEST,

rabbits,

BY SEX

(1967-1974)

(1956-1974)

AND SEASONS

dove,
_
_
_

_

(1956-1974)
PRESSURE

_

_

UNIT 7 (1963-1974)
PRESSURE

_

_

(1970-1974)

SEASONS AND HUNTING

AND SEASONS

LION HARVEST

SMALL GAME
(Small
(Small
(Small

(1956-1974)

(1956-1973)

AND SEASONS

SEASONS AND HUNTING

BLACK BEAR HARVEST

(1956-1974)

OF EITHER SEX DEER SEASONS --

'ARCHERY DEER HARVEST AND SEASONS

ELK HARVEST,

PRESSURE

PRESSURE

_

birds, raptors)

DISTRIBUTION AND ABUNDANCE OF SMALL GAME BIRDS (Grouse, mourning
chukar, pheasant, turkey, waterfowl) ANTELOPE

_

and game birds)

mammals,

DISTRIBUTION AND ABUNDANCE OF BIG GAME MAMMALS
elk, mountain lion, mule deer) ------DISTRIBUTION
snowshoe

_

STATUS --------------

LAND USE STATUS -----------

GAME SPECIES

climate)

(1965-1974)

_
_
_
_
_

HARVEST AND HUNTING PRESSURE (1968-1973)
Game Management Unit 8) -------Game Management Unit 6) ----------Game Management Units 16 and 20 - 1974) -----

_
_
_

DUCK AND GOOSE HARVEST AND HUNTING PRESSURE
(Rio Blanco and Moffat counties, 1955-1973)
(Small Game Management Units 16 and 20, 1974) ------

_
_

�-18'-

TABLE OF CONTENTS (Continued)
ITEM
SMALL GAME MANAGEMENT UNITS MAP, 1974---------------------------------SMALL GAME MANAGEMENT UNITS MAP, 19M-1973 - --------------------------_
BIG GAME POPULATION DYNAMICS ---------------------------------

_

GA}IE SPECIES INTRODUCTIONS ----------------------------------------

_

HABITAT RESTORATION PROJECTS ---------------------------------

.

MANAGEMENT PROBLEMS CHECKLIST -------------------BIG GAME MAMMAL RESEARCH REFERENCES ------------------SMALL GAME MAMMAL RESEARCH REFERENCES -----------------------------GAME BIRD RESEARCH REFERENCES --------------------NONGAME MAMMAL RESEARCH REFERENCES ------------------- ---FURBEARER RESEARCH REFERENCES --------------------- ----GENERAL MAMMALIAN REFERENCES -----------------------.-.------GENERAL AVIAN REFERENCES -------------------------

_

�-l~-

DESCRIPTION

- WILDLIFE MANAGEHENT UNIT 11 (STRAl.JBERRY)2/

Boundaries ~/.--"That

portion of Hoffat

and Rio Blanco counties east of the

Coal Creek-Crooked

Wash Creek divide, south of U. S. Highway 40 from the

Crooked Wash-Coal

Creek divide, east to the Yampa River, south of the Yampa

River to the Williams Fork River, south and west of the Williams Fork River
from the Yampa River to State Highway 13, west of State Highway 13 from the
Williams Fork River to the White River, and north of the White River from its
junction with State Highway 13 to the Crooked Wash-Coal

Creek divide."

Size.--866 square miles (planimetered from 1/2" BLH, 1974 map).

Physical Features.--Host

of Wildlife Hanagement Unit 11 lies within the

Uintah Basin section of the Colorado Plateau physiographic
1966).

province

(U.S.D.A.

It is comprised of diverse topography including lesser plateaus,

sharp ridges, abrupt cliffs, and rugged badlands interspersed with open parks
and basins.

Those portions of Unit 11 draining into the Yampa River lie

within the extreme southern portion of the Wyoming Basin physiographic
province,

a huge area of plateaus underlain by deposits of relatively soft

sedimentary
mountain

rocks, bordered in part by abrupt mountain slopes and containing

ridges.

In the northeast portion of Unit 11 is a prominent topographic

feature,

quoting U.S.D.A. 1969, - - - "the Axial Basin, formed by an uplift whose
axis has been deeply eroded exposing underlying soft rocks and forming a

!/Legally termed "Big Game Management Unit" (see footnote 2/ below) but
tentatively called "Wildlife Management Unit here to incl~de broader aspects
of animal life present.
!/Co1orado Division Wildlife Laws and Regulations Hdbk., 1975 (p. 8, Chap.
2-Big Game).

�-20-

sharply outlined

trough.

Two isolated mountains

rise abruptly from the floor

of this trough, Juniper Mountain

and Cross Mountain.

through both of these mountains

in deep canyons.

The Yampa River cuts

Along the south side of

the Axial Basin is a low range of hills, the Danforth Hills, lying about 2,000
feet above the adjacent valleys."

Elevations within Unit 11 range from 5,588

feet on the White River near the mouth of Smizer Gulch, upward to 8,720 feet
in the Danforth Hills.

Rock underlying

the eastern third of Wildlife Unit 11 are of the Mesa Verde

Formation

(U.S.D.A. 1966).

Moving westward

across the unit, a band of

relatively

soft sedimentary

rock of the Wasatch Formation

(tertiary) occur.

These are replaced by extrusions of less erodible oil shale bearing rocks of
the Green River Formation

on the west.

The Green River Formation

extends in

an irregular band northward

from the White River to the divide separating

the White and Yampa Rivers.

It is replaced again in the southwest

thirds of the unit by more Wasatch Formation.
occurs in the northwest
Formation,

two-

One other notable rock exposure

portion of Unit 11, which is of the Brown's Park

being largely soft sandstone material.

Among numerous

intermittent

River on the south boundary
Gulch, Black's

drainages flowing from Unit 11 into the White
are: Sulphur Creek, Strawberry

Creek, Smith's

Gulch, Colorow Gulch, and Crooked Wash Creek.

creeks draining ncrthward

Principal

into the Yampa River include: Milk Creek, Morgan

Gulch, Temple Gulch, Deception

Creek, and Cedar Springs Draw.

Most of these

are intermittent.

Soils data for Unit 11 can be obtained by consulting U.S. Soil Conservation
Service Work Unit personnel located at Meeker.

Descriptions

and a map of

�-21-

broad soil classifications

also can be found in U.S.D.A. 1966 and 1969 cited

below.

Climate.--National

Weather

Station data concerning

climatic conditions within

Wildlife

Unit 11 are available from three stations--Maybell,

Meeker.

All of these stations lie on unit boundaries

south sides.

Table 1 indicates

that precipitation

Hamilton,

and

on the north, east, and

is less in the open

northern

portion than L~ the eastern or southern

river bottom sites.

sections

are probably driest of all.

annual mean precipitation

Long-time

all stations is 16.5 inches .•ranging from 11. 69 at Maybell
Hamilton.

When broken down by periods,

even more pronounced

(7.43-24.06).

the variation

Cloudburst

lllliform, bi:gbeTo~.ations

for

to 19.23 at

between stations was

storms of high intensity

short duration occur over small areas and result from convective
./is ilLadjDjDjngmrtts~seas.ona1

Western

di.stribution of precipitation

ze.cei:ving a greater proportion

and

conditions •

is fairly

in winter in the

form of snow •

.lfean-8llD.Ual1:empErat:ures:were qui1:e similar between stations, varying only
from 42.8 to 44.4, with a long-time mean for all stations of 43.5 degrees.
Pe:r:i.odicmmna] .raOges we:re::"SD1IIeWhat
wider

(41.6 to 44.7) during the 15 year

..per.;iD~~9.59~..I9.14.~
....Exos.t-£r.ee.da.ysaveraged88.7 for all years and all
!It:::ati.mm .•:.::the''mea:u:&amp;"-:rar.gfn:g~.:f:rom88.2
to' 89.6.

stations

and all years was 56

to 139.

Range by years for all

�-2:2-

Table 1. Precipitation,
temperature and frost-free days, Wildlife Management
Unit 11, by period, from National Weather Station records,
1891-1974.

Precl.Eitation
Mean
Range

TemEerature
Mean
Range

Frost-Free
Dals
Mean Range

43.8

88.4

Station

Period

Years

Meeker

1961-1974

(13)

15.71

1951-1960

( 9)

17.68

44.4

1931-1952

(21)

16.85

43.9

1891-1927

(36)

J.5.81

42.8

1961-1974

(13)

18.05

1951-1960

( 9)

17.22

1947-1952

C 5)

19.23

"..J.959~97.l.~ (5)

ll.69

Hamilton

Maybell

9.90-21.18

42.5-44.7

70-121

88.2

11.71-24.06

7.43-18.92

42.8

41.6-44.2

89.6

56-139

li:i:teratureCited

U. S. Department

o£ Agriculture.

1966.

Water and related land resources,

1lhiteJtiver Rasin h,L'nlorado.

Coop. Study by Colo. Water Conserv.

Board

..
~an.dD S 11•.A.Econ •..
~

Forest Service, and Soil Conserve

Serv.

liIovembE.:l:.
Denver.

u. S. ~partment

•.Service,

"&gt;2 pp,

of Agriculture.

(Processed).

1969.

Water and related land resources,

Yampa River Basin Colorado

and Wyoming.

Conserve

Board and D.S.D.A.

Econ. Res. Service,

Conserve

Service.

April 1969.

Denver.

Coop. Study by Colo. Water

164 pp.

Forest Service, and Soil
(Processed).

�-23-

U. S. Department
Monthly

of Commerce.

(Continuing).

and Annual Summaries.

Asheville,

Climatological

data.

Colorado.

National Climatic Center, Fed. Bldg.

N. C. 28801.

William T. McKean
August 1975

�-24LANDOWNERSHIP

STATUS - WILDLIFE MANAGEMENT

UNIT 11

Bureau of Land Management

271,745 Acres

Private lands
Municipal

259,265 Acres

and County lands

50 Acres

Other Federal lands including National Park
Service, Forest Service, Indian and Military
Reservations, and Bureau of Sport Fisheries
and Wildlife
Colorado Division

of Wildlife

State Land Board Administration

o Acres

lands (north of river)
Lands

(Moffat County)

Total

360 Acres
21,970 Acres

553,390 Acres*
(864.7 sq. miles)

*

Total land acreage di~fers from total in Unit Description Section (554,240
acres or 866 sq. miles).
Bureau of Land Management lands, private lands,
and a total unit acreage were derived from a 1/2" = 1 mile B.L.M. planning
unit map (1974). A planimeter and grid were used to compute acreages.
Municipal and County lands were derived from an estimate by the City
Manager of Meeker, Colorado (8-75).

B. D. Trindle
August 1975

�-25-

LAND USE AND VEGETATIVE

Irrigated

COVER STATUS - ~~LDLIFE MANAGEME1~

cropland

Non-irrigated
Grassland

UNIT 11

9,434 Acres

cropland

19,620 Acres

with half-shrub

mixtures

21,662 Acres

Sagebrush

222,727 Acres

Brush

147,036 Acres

(desert and mountain)

Woodland

(pinon-juniper

and oakbrush)

106,991 Acres

Aspen

14,152 Acres

Commercial

timber

Riverbottom

vegetation

0 Acres

!/

454 Acres

Urban Jj

500 Acres

Miscellaneous

1/

10,764 Acres
553,390 Acres

Total

(864.7 sq. miles)
Cover type categories ,were derived from 1/2" = 1 mile S.C.S. Land Use and
Cover Type }1aps, 1953; 195.5, for Moffat and Rio Blanco counties.

A plani-

meter and grid were used to derive and compute acreage.

!/
Includes

only woods or brush-woods

mix along the White River.

Urban - Those areas used for residential and municipal purposes as illustrated on the 1/2" = 1 mile B.L.M. planning unit base map (1974) for the
't own of Meeker.
l/Miscellaneous - Interpreted to consist mainly of bare ground with sparse,
scattered vegetation of various types.

B. D. Trindle
August 1975

�-26-

HUMAN POPULATION

Specific

human population

- WILDLIFE MAJ~AGEMENT UNIT 11

data for the unit are impossible

cause U. S. Census Bureau records are on a county-wide
cause the unit encompasses
counties.

only small portions

to obtain be-

basis only and be-

of Moffat and Rio Blanco

Tables 1 ane 2 indicate county totals, incorporated

rural populations

for Rio Blanco and Moffat counties,

the only unincorporateq
unofficial

estimate

Table 1.

Urban-rural

cities and

respectively.

city within Unit 11 had 50 residents

in 1970, by

(tabulated here as rural).

population,

Rio Blanco County, 1960-70.

11

1960

1970

Meeker

1,655

1,597

Rangely

1,464

1,591

Rural

2,031

1,654

Total

5,150

4,842

Human density per square mile in Rio Blanco County decreased
1960 and 1970 from 1.6 to 1.5.

One Projection

Maybell,

slightly between

~I

of human population

~/ for Rio Blanco County is: 1975 - 5,270;

1980 - 5,267; 1990 - 6,119; 2000 - 7,088.

For Moffat County: 1975 - 7,206;

1980 - 7,485; 1990 - 8,589; 2000 - 10,226.

Another

projection

20,000;

2000 - 186,000;

A third projection
duction

of h-wman population

3/ for the White River Basin is: 1980 _

2020 - 204,000.

if for the oil shale area, based upon cumulative

oil pro-

in barrels per day, is: 1974 - 6,631; 1980 - 46,738; 1985 - 100,000.

�-2Z-

);/
U. S. Bureau of the Census and Colorado Yearbook,

1962-64.

u
Colorado State Plrulning Office, advance population
personal communication 1975.

projections,

unpublished

3/
- U. S. Bureau of the Census, Colorado Water Conservation
Field Party.
In U.S.D.A. 1965 ~ cit.

Board and USDA

U. S. Dept. of Interior (1972). Draft environmental statement for the proposed prototype oil shale leasing program.
Vol. I. Washington, D. C. 465p.

Table 2.

Urban-rural

Craig
Dinosaur

Moffat County 1960-1970.

1/

1960

1970

3,984

4,205

318

247

2,759

2,073

7,061

6,525

population,

(Artesia)

Total

1/
- U. S. Bureau of the Census and Colorado Yearbook,

lbe trend of human population

1962-64.

in Moffat County closely resembles

that in

R;.) Blanco

County.

purposes.

Human density per square mile in Moffat County decreased

between

COtmty-wide

statistics

are included here for comparative
slightly

1960 and 1970 from 1.5 to 1.4. -2/

2:./
Colorado State Planning Office, advance population
personal communication 1975.

projections,

unpublished

�-28-

CIl
'0

5

p
cO

5,150

tIl

:l
0

..c::

4

~

Q
"ri

3,135

3

Q
0

4,842
2,980

OM

~

cO

2

;:l

c,
0
P-t

1

2,943

~32

r-i

1,690

1900

1910

Fig. 1.

Population

1920

1920

1940

1950

1960

trend of Rio Blanco County.

7,061
7

.~

5,129 . .

5,086

1920

1940

1

Fig. 2.

1930
Population

1950

trend of Moffat County.

1960

1970

1970

�-29-

GAME SPECIES - WILDLIFE MANAGEME:t-.'T
UNIT 11

Big game mammals

!/

Black bear (Ursus americanus) Uncommon.
Elk (Cervus canadensis) Uncommon.
Mountain lion (Felis concolor)
Common.
Nule deer (Odocoileus hemionus) Cormnon.
Pronghorn (Antilocapra americana) Uncommon;

Small game mammals

!/

Cottontail rabbit (Sylvilagus auduboni; ~. nuttallii) Uncommon.
Pine (red) squirrel (Tamiasciurus hudsonicus) Common.
Snowshoe hare (Lepu~ americanus) Common.

Small game birds
Migratory

II

waterfowl

and shorebirds

Great Basin Canada goose (Branta canadensis moffitti) Common.
Black brant (Branta nigricans) Possible rare migrant.
~~ite-fronted goose (Anser albifrons frontalis) Possible rare migrant.
Snow goose (Chen .caerulescens caerulescens) 11 Possible rare migrant.
~fuistling swan (Olor columbianus)
Uncommon
migrant.
}Iallard (Anas platyrhynchos platyrhynchos) Common resident.

l/Nomenclature according to Lechleitner, R. R. 1969.
Colorado.
Pruett Publishing Co., Boulder.
254 pp.

Wild mammals of

21

- Nomenclature from Bailey, A. M., and R. J. Niedrach.
1967. Pictorial
checklist of Coloracl.oBirds. Denver Mus. Nat. Rist. 168 pp. Information on
occurrence and status adapted from the above reference and from Cringan, A. T.,
and L. Carlson. 1973. Wildlife in the Piceance Creek Basin, In: An environmental reconnaissance of the Piceance Basin, Rio Blanco &amp;~d Garfield counties,
Colorado.
A report on the completion of Part 1, Phase One of the environmental
inventory, analysis and impact study portion of the Regional Oil Shale Study
being done for the State of Colorado by the Thorne Ecological Institute,
Boulder, Colorado, 144 pp. Additional information on occurrence, in employing
the term "possible", is adapted from the foregoing references and also from,
Davis, W. A. 1969. Birds in western Colorado.
Colo. Field Ornithologists.
61 pp.

31
- Unverified

in hunters' bag checks but legal game 1972 through 1975.

Definitions:
Common:-should be noted regularly in the normal habitat at the
proper season of the year. Uncommon:-recorded regularly but unlikely to be
seen on most field trips. Rare:-recorded a few times only, and very unlikely
to be seen. Resident:-any species living in the unit. Migrant:-any species
moving through the unit. Where the adjective "possible" is absent, actual
sightings have been reported by Division personnel or qualified observers.

�-30-

GAME SPECIES - W1LDLIFE

MANAGEMENT

UNIT 11 (continued)

Small game birds
Gadwall (Anas strepera) Common spring and fall migrant.
Pintail (Anas acuta) Common spring and fall migrant.
American green-winged teal (~as crecca carolinensis) !!.../
Common migrant
and uncommon yearlong resident.
Blue-winged teal (Anas discors discors) Common migrant.
Cinnamon teal (~I2..ascYanoptera septent,rionalium) Common migrant.
-American wigeon (Anas americana) 4/ Common migrant and rare winter resident.
Northern shoveler (Anas clypeata)-!!.../
Common migrant and uncommon summer
resident.
Redhead (Aythy~ americana) Uncommon migrant.
Ring-necked duck (!&gt;ythya collaris) Uncommon migrant.
Canvasback (~?:.
valisineria) Uncorrnnonto rare migrant.
Greater scaup (Aythya marila nearctica) Possible rare resident.
Lesser scaup (Aythya affinis) Uncommon migrant.
Common golden-eye (Eucephala clangula americana) Common migrant and uncommon winter resident.
Barrow's golden-eye (Bucephala islandica) 1/ Rare winter visitor.
Bufflehead (Bucephala albeola) Uncommon spring and fall migrant and
rare winter resident.
Ruddy duck (Oxyura jamaicensis rubida) Common migrant and occasional
summer resident.
Hooded merganser (Lophodytes cucullatus) Rare winter visitor on river.
Common merganser (Hergus merganser ame rLcanus ) Common winter resident,
uncommon summer resident.
Red-breasted merganser (~ergus serrator serrator) 1/ Possible rare migrant.
American coot (Fulica americana arnericana) Common migrant and summer resident.
Common Wilson's snip~ (Capella gallinago delicata) Common migrant and rare
winter resident.
Sandhill crane (Grus canadensis tabida) Regular migrant.
Virginia rail (Ralus limicola limicola) Possible uncommon summer resident.
Sora (Porzana carolina) Possible uncommon summer resident.

Upland game birds
Blue grouse (Dendragapus obscurus obscurus) Common.
Sage grouse (Centrocercus urophasia~us urophasianus) Uncommon to common.
Ring-necked pheasant (Phasianus colchicus) Uncommon.
Chukar (Alectoris chukar) 4/ Uncommon to rare.
Band-tailed pigeon (Columb~ fasciata fasciata) Possible uncommon summer
and fall migrant.
Mourning dove (Zenaida macroura marginella) !!.../
Common summer resident.
Wild turkey (Meleagris gallopavo merriami) Rare.
Sharp-tailed grouse (Pedioecetes phasianellus columbianus) Rare.

4/
- Changes in the nomenclature follow the thirty-second supplement to the
American Ornithologists Union check-list of North American birds published
in Auk 90:411-419, April, 1973.

W. T. McKean
September 1975

�-31-

OTHER MAMMAl,IAN SPECIES
Furbearers

1,/ - l.JILDLIFEHANAGEMENT

UNIT 11

~I

Short-haired
Beaver (Castor canadensis) Uncommon.
Nink (Hustela vison) Uncertain.
Muskrat (Ondatra zibethicus) Uncommon.
Ringtail ~~~i~'~Ut~s)
Rare.
Weasels (Hus t ela ern·ll·nea,·
frenata)
,',
n.
Uncorrnnon.
M.

H

ermfnea

Unc er-t
adn ,; M • frenata

Long-haired
Kit fox (Vulpes velox) Uncommon.
Gray fox (Urocyon cinereoargenteus) Uncommon.
American badger (Taxidea taxus) Common to uncommon.
Spotted skunk (Spil~~ius)
Uncommon.
Striped skunk (Mephitis mephitis) Common.
"VaDn:int" mannnals
Coyote (Canis latrans) Common.
Red ..
fox JV.ulpes fulva) Uncommon.
Raccoon (PTC'..c:yon lotor) Common ,
Porcup.lne.g.:etbi2 ~D. darsatum) Common.
Btibcat{w".J.ldcat)(Lynx rufus) Common ,
Black-tailed jack rabbit (Lepus californicus) Rare to uncommon.
White-tailed jack rabbit (Lepus townsendii) Common.

11
- These species, grouped separately as "Furbearers", "Varmints" and "Nongame mannnals" and outside of "game" categories, follow Chapter 62, Colo.
Rev. Statutes 1963.l,,,, Amended, in Colo. Game, Fish and Parks Div. Laws and
Regulations .Hdhk.~ 1'975 (Art. 1, Items 14, 20, 31, and 42, Definitions, p. 2).
-,

~.

: Nomem:.1:a:t:nr.':',0U1.
Lechleitner, R. R. 1969. Wild mammals of Colorado.
Pruett Publi.shing Co., Boulder.
254 pp. Information on occurrence and status
from the above reference and: Cringan, A. T., and L. Carlson.
1973. Wildlife
in the Piceance Creek Basin, In: An environmental reconnaissance of the
Piceance Basin, Rio Blanco and Garfield counties, Colorado.
A report on the
completion of Part 1, Phase One of the environmental inventory, analysis and
impact study portion of the Regional Oil Shale Study being done for the State
of Colorado by the Thorne Ecological Institute, Boulder, Colorado, 144 pp.
Also, Armstrong, D. M., 1972, Distribution of mammals in Colorado. Monograph
of the Museum of Natural History, the Univ. of Kansas, Number 3, 1972. 415 pp.

Definitions:
C{)r~p],,-:-should
be noted regularly i.n the normal habitat at the
proper season of th-,
· year. Uncommon:-recorded regularly but unlikely to be
seen on most field trips. Rare:-recorded a few times only, and very unlikely
to be seen. Resident:-any species living in the unit. Migrant:-any species
moving through the unit. Where the adjective "possible" is absent, actual
sightings have been reported by Division personnel or qualified observers.

�OTHER HAMMALIAN

-32SPECIES (continued)

"Varmint" mammals

(continued)

Yellow-bellied marmot (Hal~ota flaviventris) Common.
White-tailed prairie dog (Cynomys leucurus) Common.
Richardson's ground squirrel (Spermophilus richardsonii) Common.
Thirteen-lined ground squirrel (Spenuophilus tridecem1.ineatus) Common.
Rock squirrel (§permophilus variegatus) Common to uncommon.
Northern pocket gopher (Thomomys talpoides) Common.
Nongame mammals
Golden-mantled ground squirrel (Spermophilus lat~alis) Common.
White-tailed antelope squirrel (Ammospennophilus leucurus) Uncommon.
Least chipmunk (Eutamias minimus) Common.
Colorado chipmunk (Eutamias ,guadrivittatus) Common.
Uinta chipmunk (Eutamias umbrinus) Uncommon to uncertain.
Vagrant shrew (Sorex vagrans) Uncommon.
Merriams shrew (Sorex merriami) 3/ Uncertain.
Masked shrew (Sorex cinereus) Uncommon.
Townsend's big-eared bat (Plecotus townsendii) Common.
Silver-haired bat (Lasionycteris noctivagans) Uncommon.
Hoary bat (Lasiurus cine reus) Uncommon - common.
Big brown bat (Eptesicus fuscus) Common.
\~estern pipistrelle (Pipistrellus hesperus) Common.
Long-legged myotis (Mvotis volans) Uncommon.
California myotis (Myotis californicus) Uncommon.
Small-footed myotis (Myotis leibii) Common.
Long-eared myotis (Myotis evotis) Uncommon.
Little brown myotis (Myotis lucifugus) Uncommon.
Ord's kangaroo rat (Dipodomys ordii) Uncommon - uncertain.
Apache pocket mouse (Perognathus apache) Uncommon.
Western harvest mouse (Reithrodontomys nagalolis) Uncertain.
Canyon mouse (Peromyscus crinitus) Common - uncommon.
Deer mouse (Peromyscus maniculatus) Common.
Pinon mouse (Peromyscus truei) Common.
Bushy-tailed wood rat (Neotoma c.inerea) Common.
Desert woodrat (Neotoma lepida) Uncommon.
Gapper's red-backed vole (Clethrionomys gapperi) Uncommon.
Montana vole (Microtus montanus) Uncommon.
Long-tailed vole (Microtus longicaudus) Common.
Sagebrush vole (Lagurus curtatus) Common.
House mouse (Mus musculus) Uncommon.
Western jumping mouse (Zapus princeps) Uncommon.
110ccurrence listed by the Colo. Div. of Wildlife as extremely unusual very few documented records within the past decade. (1972 Status Evaluation
for Selected Colorado Species) appended to: 1973 Wildlife Operations Work
Plan, Field Order No. 4- 1973).
W. T. McKean
September 1975

�-33OTHER AVIAN SPECIES 1../ - \HLDLlFE MANAGEMENT UNIT 11
"Vannin t" birds
English sparrow (Passer domesticus domesticus) Common resident.
Black billed magpie (Pica .EJ:~ hudsonia) Common resident.
Starling (Sturnus vulgaris) Common resident.
Rock dove (Columbia livia) Uncommon resident.
Nongame birdJ.I
Common loon (Gavia immer) Possible rare migrant.
Horned grebe (Podic~ps auritus cornutus) Possible rare migrant.
Eared grebe (Podiceps migricollis californicus) Corrunonmigrant 1/.
Western grebe (Aechmophorus occidentalis) Rare migrant.
Pied-billed grebe (Podilymbus podiceps podiceps) Possible uncommon
migrant and uncommon summer resident.
Double-crested cormorant (Phalacrocorax auritus auritus) Possible rare
migrant.
Great blue heron (Ardea herodias treganzai) Common summer resident.
Snowy egret (Egretta thula brewsteri) Uncommon summer resident 1/.
Black-crowned night heron (Nycticorax nycticorax hoactli) Possible
common summer resident.
Least bittern (Ixobrychus exilis exilis) Possible rare summer migrant.
American bittern (Botaurus lentiginosus) Possible rare summer migrant.
~lite-faced ibis (Plegadis chihi) Possible uncorrunonmigrant.
Semipalmated plover (Charadrius semipalmatus) Possible rare migrant.
Killdeer (Charadius vociferus vociferus) Corrunonsurrunerresident and
rare winter resident.
Mountain plover (Charadrius montanus) Possible rare migrant 1/ .
.!/These species, grouped separate~y as "Varmint" birds, "Nongame birds"
and "Raptors" and outside of "game" categories, follow Chapter 62, Colo.
Rev. Statutes 1962 As Amended, in Colo. Game, Fish and Parks Div. Laws and
Regulations Hdbk., 1975.
(Art. 1, Items 20, 28, 31 and 42, Definitions,
p. 2).

21Nomenclature from Bailey, A. M., and R. J. Niedrach. 1967. Pictorial
checklist of Colorado Birds. Denver Mus. Nat. Hist. 168 pp. Information on
occurrence and status adapted from the above reference and Davis, W. A.
1969. Birds in western Colorado. Colorado Field Ornithologists.
61 pp.
l.1Changes in nomenclature follow the Thirty-second Supplement to the American
Ornithologists Union Checklist of North American Birds published in Auk
90(2):411-419.
April, 1973.
Definitions:
Common:-should be noted regularly in the normal habitat at the
proper season of the year. Uncommon:-recorded regularly but unlikely to be
seen on most field trips. Rare:-recorded a few times only, and very unlikely
to be seen. Resident:-any species living in the unit. Migrant:-any species
moving through the unit. Where the adjective "possible" is absent, actual
sightings have been reported by Division personnel or qualified observers.
Notations concerning seasonal abundance of raptors are by Gerald R. Craig,
Wildlife Biologist.

�-34OTHER AVIAN SPECIES

(continued)

Nongame birds
Black-bellied plover (Pluvial is sguatarola) Possible uncommon migrant 1/.
Long-billed curlew (Numenius americanus americanus) Possible rare migrant.
Spotted sandpiper (Arc t Lt Ls ~llac_ulari&lt;l)
Possible cornmon summer resident.
Solitary sandpiper (Tringa ~9l:i:.~ariacinnamomea) Possible common migrant.
and occasional summer resident.
Greater sandhill .:rane (Grus canadensis tabida) Regular migrant and
possible summer resident endangered.
\-1illet(CataptrophoTus semipalmatus inornatus) Rare migrant.
Greater yellowlegs (Tringa melanoleuca) Possible cornmon migrant
/.
Lesser yellowlegs (Tringa flavipes) Possible uncommon migrant l/~
Red knot (Calidris canutus rufa) Possible rare migrant 1/.
Pectoral sandpiper (Calidris melanotos) Possible rare migrant 3/.
Baird's sandpiper (Calidris bairdii) Possible common migrant 3/.
Least sandpiper (Calidris millutilla) Possible common migrant 3/.
Long-billed dowitcher (Limnodromus scolopaceus) Possible unco;mon migrant.
Stilt sandpiper (Micropalma himantopus) Possible rare migrant.
Semipalmated sandpiper (Calidris pusilla) Possible rare migrant 1/.
Western sandpiper (Calidris mauri) Possible uncommon migrant 3/.
Marbled godwit (Limosa fedoa) Possible rare spring migrant.
Sanderling (Calidris alba) Possible rare migrant 3/.
American avocet (Recu~ostra
americana) Rare migrant.
Black-necked stilt (Hamantopus mexicanus) Rare migrant.
Wilson's phalarope (Steganopus tricolor) Possible common migrant and
uncommon summer resident.
Northern phalarope (Lobipes lobatus) Possible uncommon migrant.
Herring gull (La~usargentatus
smithsonianus) Possible uncommon migrant.
California gull (Larus californicus) Possible rare migrant.
Ring-billed gull (Larus delawarensis) Possible uncommon migrant.
Franklin's gull (Larus pipixcan) Possible uncommon migrant.
Bonaparte's gull (Larus philadelphia) Possible rare migrant.
Sabine's gull (Xema sabini sabini) Possible rare migrant.
Forster's tern ~rna
forsteri) Possible rare migrant.
Common tern (Sterna'hirundo hirundo) Possible rare migrant.
Least tern (Sterna albafrons athalassos) Possible rare migrant.
Black tern (Chlidonfas niger surinamensis) Possible uncommon migrant.
Rock dove (Columba livia) Possible common resident.
Unprotected.
Yellow-billed cuckoo (Coccyzus americanus americanus) Possible rare
summer resident.
Poor-will (Phalaenoptilus nuttallii nuttallii) Uncommon summer resident.
Common nighthawk (Chordeiles minor hesperis; ~. ~. howeli) Common
summer resident.
White-throated swift (Aeronautes saxatalis sclateri) Common summer resident.
Black-chinned hummingbird (Archilochus alexandri) Uncommon summer resident.
Broad-tailed hummingbird (Selasphorus platycercus platycercus) Common
summer resident.
Rufous hummingbird (Selasphorus rufus) Possible common late summer migrant.
Calliope hummingbird (Stellula calliope) Possible common late summer
migrant.
Belted kingfisher (Megaceryle alcyon alcyon) Possible common resident.
Common flicker (Colaptes auratus cafer) Common resident 1/.
Lewis' woodpecker (Asyndesmus lewis) Possible uncommon summer resident.
Red-headed woodpecker (Melaner~rythrocephalus)
Possible rare resident.

�,

-3~-

OTHER AVIAN SPECIES

(continued)

Nongame birds
Pileated woodpecker (Dryocopus pileatus picinus) Possible rare summer
resident.
Yellow-bellied sapsucker (SphY~icus
varius nuchaliae) Common summer
and possible occasional winter resident.
Williamson's sapsucker (~rapicus
thyroideus nataliae) Possible
uncommon SUIDrrler
resident.
Hairy woodpecker (Dendrocopos villosus monticola) Common year-a-round
resident.
Downy woodpecker (Dendrocopos pubescens leucurus) Common year-a-round
resident.
Eastern kingbird (Tyrannus tyrannus) Uncommon summer resident.
Western kingbird (fyrannus verticalis) Common summer resident.
Ash-throated flycatcher (Hyiarchus cinerascens cinerascens) Common summer
resident.
Say's phoebe (Sayornis saya saya) Common summer and possible rare winter
resident.
Willow flycatcher (Empidonas traillii) Possible uncommon summer resident 1/.
Hammond's flycatcher (Empidonax hammondii) Possible migrant and uncommon
summer resident.
Dusky flycatcher (Empidonax oberholseri) Summer resident.
Gray flycatcher (Empidonax wrightii) Common summer resident.
Western flycatcher (Empidonax diffic:ilis hellmavri) Common summer resident.
Western wood peewee (Contopus sordidulus v~liei) Common summer resident.
Olive-sided flycatcher (Nuttallornis borealis) Possible common summer
resident.
Horned lark (Eremophila alpestris leucolaema) Common year-a-round resident
and migrant.
Violet-green swallow (Trachycineta thalassina lepida) Common summer resident.
Tree swallow (Iridoprocne bicolor) Common migrant and summer resident.
Bank swallow (Riparia.riparia riparia) Possible uncommon migrant and
summer resident.
Rough-winged swallow (Stelgidopteryx ruficoll:i.sserripennis) Uncommon
migrant and summer resident.
Barn swallow (Hirundo rustica erythrogaster) Common summer resident.
Cliff swallow (Petrochelidon pyrrhonota pyrrhonota) Common summer resident.
Gray jay (Perisoreus canadensis cap,italis) Possible uncommon resident.
Steller's jay (~citta
stelleri macrolopha) Common resident.
Scrub jay (Aphelocoma coerulescens woodhouseii) Common resident.
Common raven (Corvus corax sinuatus) Common resident.
Common crow (Corvus b~rhvnchos
brachyrhynchos) Uncommon resident.
Pinon jay (Gymnorhinus cyanocephalus) Common summer 1/ and winter resident.
Clark's nutcracker (Nucifraga columbiana) Common resident.
Black-capped chickadee (Parus articapillus garrinus) Common resident.
Mountain chickadee (Parus gambeli gambeli) Common resident.
Plain titmouse (Parus inornatus ridgwayi) Uncommon resident.
Bushtit (Psaltriparus minimus plumbeus) Possible common resident 1/.
White-breasted nuthatch (Sitta carolinensis nelson i) Possible uncommon
resident.
Red-breasted nuthatch (Sitta canadensis) Possible rare year-a-round resident.
Pygmy nuthatch (Sitta pygmaea melanotis) Possible uncommon resident.

�-36-

OTHER AVIAN SPECIES

(continued)

Nongame birds
Brown creeper (Certhia famili8ris~an~)
Possible uncommon year-around resident.
Dipper (Cinclus mexicanus unicolor) Common resident.
House wren (T;:-i8l;;dYJ-:.es
;;ec1on'p_arbnan~i)Common summer resident.
b ew i ck Lf _eremophilu~) Possible summer resident
Bewick's wren (ThrY012.ane~2.
and rare winter resident.
Canyon wr en (Salpinctes mexicanus consper~) Possible uncommon summer
resident.
Rock wren (Salpinctes obsoletus obsoletus) Common summer and possible
rare winter resident.
Mockingbird (Himus polyglottos leucopterus) Uncommon summer resident.
Gray catbird (Dumetel1a carolinensis) Possible rare summer resident 3/.
Sage thrasher (O~;Ptes
ruontanus) Possible common sumnler resident~
American robin (Turdus migratorius propinguus) Common summer and winter
resident 3/.
Hermit thrush (Catharus guttatus audubonii)
Common summer resident 1/.
Swainson's thrush (Catharus ustulatus almae) Uncommon migrant.
Veery (Catharus fucescens saliCi~
Uncommon migrant and summer resident 1/.
Western bluebird (8ialia mexicana bairdi) Common migrant and uncommon
summer resident.
Mountain bluebird (8ialia currucoides) Cownon migrant and summer resident
and possible occasional winter resident.
Townsend's solitaire (Nyadeste~ to,",7J.1sendi
townsendi) Uncommon resident.
Blue-gray gnatcatcher (Poliptila caerulea amoenissima) Common summer
resident.
~olden-crowned kinglet (Regulus satrapa amoenus) Possible uncommon
migrant and rare summer resident.
Ruby-crowned kinglet (Regulus calendula cineraceus) Possible uncommon
migrant and summer resident.
Bohemian waxwing (Bombycilla garrulus pallidiceos) Possible irregular
winter migrant 1/.
Cedar waxwing (Bombycilla cedrorum) Uncommon resident.
Northern shrike (Lanius excubitor invictus) COIDlllon
winter resident.
Loggerhead shrike-C~us
ludovicianus excubitorides) Possible uncommon
summer and uncommon winter resident.
Gray vireo (Vireo vicinior) Possible common summer resident.
Solitary vireo (Vireo solitarius plumbeus) Common summer resident.
Red-eyed vireo (Vireo olivaceus) Possible rare summer resident.
Warbling vireo (Vireo gilvus swainsonii) Common summer resident.
Tennessee warbler (Vermivora Eeregrina) Possible rare but regular migrant.
Orange-crowned warbler (Vermivora celata orestera) Common migrant and
summer resident.
Nashville warbler (Vermivora ruficapilla ridgwayi) Possible rare migrant.
Virginia's warbler (Vermivora virginiae) Common summer resident.
Yellow warbler (Dendroica petechia aestiva) Common summer resident.
Yellow-rumped warbler (Dendroica coronata memorabilis) Cownon summer
resident 3/; (D. c. coronata) Possible common migrant 3/.
Black-throat;d gr~y ;arbler (Dendroica nigrescens) Possible common summer
resident.
Townsend's warbler (Dendroica townsendi) Uncommon fall migrant.
MacGillivray's warb l.er (Oporornis tolmiei monticola) Common summer resident.

�-37OTHER AVIAN SPECIES (continued)

Nongame birds
Common yellowthroat (Geothylpi~ trichas occidentalis: £. !.. campicola)
Possible uncommon summer resident 3/.
Yellow-breasted chat (Icteria virer~ ;-uricollis) Possible common summer
resident.
Wilson's warbler (Wi!.~6ni~ l?usilla .£i}.eol
ata) Possible common migrant
and summer resident.
American redstart (Setopha?~ rutic~l]:~ tricolora) Possible rare migrant.
House spar row (Passer domesticus domestic.us) Common resident.
Bobolink (Dolicli~ny~oryzivorus) Possfble UnCOllh'TIOn
summer migrant.
WesteTIl meadowl ark (.§tun~ella_neglecta neglecta) Common summer and
possible uncommon winter resident.
Yellow-headed blackbird (Xanthocephalus xanthocephalus) Common summer
resident.
Red-w.Lnged blackbird (Agelaius phoenJceus. fortis) Common resident.
Northern oriole (~rus
galbula bullockii) Possible common summer
resident 1/.
Rusty blackbird (Euphagus carolinus carolinus) Possible rare winter migrant.
Brewer's blackbird (Euphagus cyanocephalus) Common resident.
Common grackle (Quiscalus quiscula) Possible rare summer resident.
Brovm-headed cowbird (Nolothrus ater artemisiae) Common summer resident.
WesteTIl tanager (Piranga ludovici;~)
Common migrant and summer resident.
Scarlet tanager (Piranga olivacea) Possible rare migrant.
Black-headed grosbeak (Pheucticus melanocephalus melanocephalus) Uncommon
summer resident.
Lazuli bunting (Passerina amoena) Common summer resident.
Evening grosbeak (Hesperiphona vespertina brooksi) Uncommon summer
resident and irregular winter resident.
Cassin's finch (Carpodacus. cassinii) Common resident.
House finch (Carpodacus mexi~anus frontalis) Common summer and possible
uncommon winter resident.
Pine grosbeak (Pinicola enucleator montana) Possible uncommon resident.
Gray-crowned rosy finch (Leucosticte tephrocotis tephrocotis; L. t.
littoralis) Possible common winter resident.
Black rosy finch (Leucosticte atrata) Possible common winter migrant.
Brown-capped rosy finch (Leucosticte australis) Possible common winter
migrant.
Common redpoll (Acanthis flammea flammea) Possible rare winter migrant.
Pine siskin (Spinus pinus pinus) Common resident.
American goldfinch (Spinus tristus tristus; ~. !.. pallidus) Possible
common summer and uncommon winter resident.
Lesser goldfinch (Spinus psaltria psaltria) Possible uncommon summer and
rare winter resident.
.
Green-tailed towhee (Chlorura chlorura) Common summer resident.
Rufous-sided towhee (Pipilo erythrophthalmus montanus) Common summer
and rare winter resident.
Lark bunting (Calamospiza melanocorys) Possible uncommon summer resident.
Savannah sparrow (Passerculus sandwichensis nevadensis; l. ~. anthinus)
Common migrant and summer resident.
Grasshopper sparrow (Ammodramus savannarum perpallidus) Possible uncommon
summer resident.
Vesper sparrow (Pooecetes gramineus confinis) Common migrant and
summer resident.

�-38OTHER AVIAN SPECIES

(continued)

Nongame birds
Lark sparrow (Chondestes·grammacus·strigatus)
Common migrant and
summer resident.
Black-throated sparrow (Amphispiza ?ilineata deserticola) Possible common
Sage sparrow (Amphispiza belli nevadensis) Common summer resident.
Dark-eyed junco (Junco .ki:emalisaikeni) Possible rare winter resident 1/;
(L· ~. hyemalis; L· ~. cismontanu~) Rare winter resident 1/;
(!. ~. oreganus) Common winter resident 1/; (L. ~. ~. var. mearnsi)
Common winter resident 3/.
Gray-headed junco (Junco c;niceps caniceps) Common summer and winter
resident.
Tree sparrow (Spizella arborea ochracea) Possible uncommon winter migrant.
Clay-colored sparrow (Spizella pallida) Possible common migrant.
Chipping sparrow (Spizella passerina boreophila) Common summer resident.
Brewer's sparrow (Spizella breweri breweri) Common summer resident.
Harris' sparrow (Zonotrichia querula) Possible uncommon migrant and rare
winter resident.
White-crowned sparrow (Zonotrichia leucrophrys) Common resident.
~~ite-throated sparrow (Zonotrichia albicollis) Possible rare migrant.
Fox sparrow (Passerella iliaca schistacea) Rare summer resident.
Lincoln's sparrow (Helospiza lincolnii alticola) Possible common migrant
and summer resident.
Song sparrow (Melospiza melodia) Common summer and uncommon winter resident.

Rapt or s

~/

Turkey vulture (Cathartes ~
meridionalis) Common summer and rare
winter resident.
Goshawk (Accipiter gentilis atricapillus) Uncommon summer and common
winter resident.
Sharp-shinned hawk (Accipiter striatus velox) Uncommon summer and common
winter resident.
Cooper's hawk (Accipiter cooperii) Common summer and winter resident.
Red-tailed hawk (Buteo jamaicensis calurus) Common resident.
Swainson's hawk (Buteo swainsoni) Uncommon summer resident.
Rough-legged hawk (Buteo lagopus ~. johannis) Common winter resident and
migrant.
Ferruginous hawk (~uteo regalis)
Common summer and common winter resident.
Golden eagle (Aquila chrysaetos canadenSis) Common resident.
Bald eagle (Hal Laae t u's leucocephalus alascanus) Common winter resident.

�-39-

OTHER AVIAN SPECIES

(continued)

Raptors

&lt;s!-.~

Marsh hawk
syaneus hudsonius) Common summer and
common
winter resident.
Osprey (Pandion haliaetus carolinen:=;}_s)
Uncommon migrant, rare
summer resident.
Prairie falcon (Falco mexicanus) Uncommon resident.
Peregrine falcon (Falc~~1us
anatum) Rare migrant, endangered.
Merlin (Falco columbarius) Uncommon winter migrant 3/.
American kestrel (Falc~~arverius
sparverius) Con®;n summer and
unconunon winter resident.
Screech owl (Otus asio) Common resident.
Flammulated owl (Otus flamrneolus flammeolus) Rare summer resident.
Great horned owl (Bubo virginianus) Connnon resident.
Pygmy owl (Glaucidium gnoma californicum) Rare resident.
Burrowing owl (Speotyto cunicularia hypugaea) Common summer and
possible rare winter resident.
Long-eared owl (Asio otus wilsonianus) Uncommon resident.
Short-eared owl. (Asi£.pamrneus flammeus) Common summer resident and
unc~mmon winter migrant.
Saw-whet 0107) (Aegolius acadicus acadicus) Uncommon resident.

ADDITIONAL

LITERATURE

REFERENCES

.Cringan, .A. T., and L. Carlson.
1973. Wildlife in the Piceance
£reek.:Basin, In: An environmental reconnaissance of the Piceance
Basin, Rio B1arco and Garfield counties, Colorado. A report on
the completion of Part 1, Phase one of the environmental inventory,
analysis and impact study portion of the Regional Oil Shale Study
being done for the State of Colorado by the Thorne Ecological
Inst::it:ut.f', Boulder, Colorado.
144 pp •
•---~:,&gt;:f:

'='., P. H. Baldwin, and E. B. Reed.

-nf 1riri!~from the Yampa Valley, northwestern
7f) (l) :ll3-116.

1974. Recent records
Colorado.
Condor

Smith, A. G. 1973 •. Avian environmental features at the Colony plant
site, Garfield Co'unty, Colorado.
Environmental inventory by
Thorne Ecological Institute for Colony Development Operation,
Atlantic-Richfield Company, Operator, October, 1974.

W. T. McKean and B. D. Trind1e
September 1975

�-40-

DISTRIBUTION

AND ABUNDANCE OF BIG GAME HAHMALS

WILDLIFE

HANAGEHEl\l1' UNIT

11

ANTELOPE

Antelope
Roland

utilize approximately

(1975 pers. comm.) reported

into two areas.

of Duffy Mountain,

approximately

1,000 antelope on it.

approximately

W.C.O.

that the 70 square miles are divided

One area south of Haybell,

miles, has approximately

on it.

70 square miles of Unit 11.

51 square

The other area southwest

19 square miles, has about 70 antelope

W.C.O. Roland feels that there is some interchange

two areas, but that the populations

remain relatively

between the

stable yearlong.

BLACK BEAR
&amp;lack bears .gene.ra.l.Ly
occupy

the southeast quarter of Unit 11.

incJ.~!deBtht'.J)&lt;;!'L;
i.,rtb Hills and CoLo row Mountain
provide suitable
.~

E,70D

habitat

£eet ,with

The Colorado

jj) ..a.ezrs

EstImated

'?'!

The elevation

ranges between

6,200

of Wildlife,

Wildlife

Inventory

Systems

(1974),

figure of .035 for the above range, which yields a total

"eveza.l L" population
-we.r£

regions both of which

the lands being timbered and brush covered.

Division

,~ts,a:.dJ!'nS:1ty

for bears.

This area

of 10 bears on Unit 11.

W.C.O. estimates

for 1975 also

Unit 11.

annual black bear harvest for the 19 year period 1956-1974

in-

clusive was 2.3 bears, based upon hunter report card surveys 1956-1971,
the random sample 1971-1974.

and

Annual harvest ranged from 0-11; there were

eight years out of the nineteen when no bears were reported

taken.

ELK
W.C.O. 's Krager, Ko1and, and Smith (1975 pers. comm.) estimate a total of

�-41-

515 elk are on Unit 11 year-round.
found in the Colorow Mountain

The majority

of these elk (450) are

and Danforth Hills regions.

During the

summer 90 percent of these elk are found east of Strawberry

Creek in the

Danforth Hills, with 10 percent summering on Color ow Mountain.
winter approximately

325 elk occur east of Strawberry

occur west of Strawberry
winters

Creek on Colorow Mountain.

on both areas and

of their range.

Three other smaller herds are located within Unit 11.
30 head yearlong
Mountain

on Duffy Mountain,

according

cluding archery.
presented

There are about

20 on Isle Mountain,

to W.C.O. Roland

}lean annual elk harvest

Creek and 125 elk
During severe

the elk will move off the higher elevations

occupy the lower elevations

In the

and 15 on Cross

(1975 pers. cOlmn.).

(1956-1974) was 96 with 6 closed seasons, in-

Annual harvest ranged from 6 to 258 (see harvest data)

in this report.

MULE DEER

Unit 11 is almost entirely mule deer winter range except for the highest
elevations

occurring

range migrate

in the Danforth

Hills.

The deer that occupy this

into the Unit from outlying areas.

estimated by the Colorado Division

of Wildlife,

Winter population,
Wildlife

Inventory

Systems

(1974), was 10,253 animals with a density of 12.7 deer per square mile.

Relatively
able.

few deer summer on Unit 11, with no data on abundance

avail-

Those deer that do remain during the summer can be found along the

White and Yampa rivers and in the highest elevations

Mean annual deer harvest

of Unit 11.

(1956-1974) was 2,674, including archery.

Annual

�-42-

harvest

ranged from 892 to 5,674 (see harvest data) present

in this

report.

MOUNTAIN

LION

Mountain

lions may be found in the southern half of Unit 11 and two

distinct

areas bordering

highest

concentration

Colorado

Armstrong

(1972), Dixon

(1969) make specific

citations

concerning

of Wildlife,

bution and abundance
W.C.O.

data estimated

estimates

Inventory

a total population

Hunter's

are as follows:

(1967), and

lions with Unit 11.

Systems

for 1975 were a maximum

mum of 8 lions on Unit 11.
period 1965-1974

Wildlife

(1974) distri-

of 14 lions within

of l5'lions

and a mini-

reports of lions killed during the

1965 - 2; 1966 - 9; 1967 - ND; 1968 - 0;

1969 - 2; 1970 - 0; 1971 - 0; 1972 - 3; 1973 - 1; 1974 - O.
harvest

The

with the high, rough terrain

region.

Division

Unit 11.

(see mylar map overlay).

of lions is associated

of the Colorow Mountain
Lechleitner

the Yampa River

Mean annual

for this period was 1.9 lions (no data for 1967; see accompanying

Harvest Data).

Armstrong,

D. M;

1972.

Kansas Printing

Colorado

Division

wide mammal

LITERATURE

CITED

Distribution

of mammals

Service, Lawrence.

of Wildlife,

Wildlife

and bird distribution

area supervi.sor, and regional
office, Colo. Div.Wi1dl.

in Colorado.

Univ. of

4l5pp.

Inventory

Systems.

1974.

maps and data compiled

biologist

Unpublished.

input.

Located

State-

from W.C.O.,
at Denver

�-43-

Dixon, K. R.

1967.

pp. 141-164.

Evaluation

In Game Res. Rep., July - Part II.

Game, Fish and Parks.

Lechleitner,
Boulder.

R. R.

of effects of mountain

1969.

lion predation,
Colo. Div.

pp. 73-310.

Wild mamnlals in Colorado.

Pruett Publ. Co.,

234pp.

B. D. Trindle and W. T. McKean
November 1975

�DISTRIBUTION AND ABUNDANCE OF SMALL GAME MAMMALS
WILDLIFE HANAGEMEi\TTUNIT 11

COTTONTAIL

RABBIT

Nearly every square mile of Unit 11 is considered
at least some cottontail

rabbits yearlong, however,

they are more abundant

at the lower elevations.

tail and the Nuttall's

cottontail

being common at lower elevations
tail being less abundant

(5,000-7,000)

1969, Armstrong

No specific

data is ,available on cottontail

for Small Game Management

that

Both the desert cotton-

and the Nuttall's

in the higher elevations

as reflected

it is believed

should occur; the desert cottontail

(Lechleitner

For abundance,

capable of supporting

cotton-

(6,000-8,000)

1972).

rabbit densities

for Colorado.

in harvest data, see tables on Hunter Harvest

Units 6 and 8, following.

SNOWSHOE HARE
Snowshoe

hares may exist in Unit 11 above 8,000 ft. in elevation

Douglas-fir,

Douglas-fir-aspen

mix.

Shepherd

8,000 ft. in elevation would be potential

No specific

data is available

(1965) suggested

in the

that above

range for much of Colorado.

for dis~ribution

or abundance

of snowshoe

hares in Unit 11.

RED (PINE) SQUIRREL
Armstrong

(1972) and Lechleitner

(1969) suggested

should be fairly common in the coniferous
fir, in the higher el~vations

No data on abundance

forests, principally

of Unit 11.

or harvest

that this squirrel

are available.

Douglas-

�-45-

LITERATURE
Armstrong,

D. M.

1972.

CITED

Distribution

of mammals

of Kansas Printing Service, Lawrence.

Lechleitner,

R. R.

Boulder.

Shepherd, H. R.

1969.

in Colorado.

Univ.

4l5pp.

Wild mammals of Colorado.

Pruett Publ. Co.,

254pp.

1965.

Colorado long range game species management

plans, 1965-1975,

for snowshoe hare.

Parks Dept. rept.

11pp.

Colorado Game, Fish and

VIimeo).

B. D. Trindle and W. T. McKean
November 1975

�-46-

DISTRIBUTION

AND ABUNDANCE

WILDLIFE

OF SMALL GAME BIRDS

NANAGEMENT

UNIT 11

BLUE GROUSE
Blue grouse distribution

includes the Danforth

Hills and Colorow

regions of Unit 11 starting at about 7,000 feet in elevation

Mountain

and co~tinuing

upwards.

This includes most of the timbered areas of

the unit which provides

suitable habitat for the grouse.

W.C.O. 's Krager and Smith (1975 pers. comm.) suggest that both the Danforth Hills and Colorow Nountain
They estimated
mile

J

£01:'

regions have low populations

of grouse.

that an average density woul.d be about 2 birds per square

the occupied

range.

SAGE GROUSE
Sage-gr(;1lSe inhabit

all of Unit 11 except for a small area surrounding

t:beCi.ty of Heeker,

according

to w.e.o.'s

Krager, Smith, and Roland

(1975

They also suggest a density of less than one bird per
square mile for all the potential

habitat

in Unit 11, except for the Axial

.Basin:where the suggested

density is 15 birds per square mile.

figur.es-axein

with Rogers'

accordance

SHARP-TAILED

w.e.D. Smith has received
eastern

edge of Unit 11.

reported

These

(1964 ) estimates.

GROUSE

sightings

of sharp-tailed

No specific data on distribution

grouse on the
or abundance

are

available.

MOURNING
Acco.rdLng to Grieb
mourning

doves.

DOVE

(1"965) the entire unit is potential

Mourning

doves are common throughout

breeding

range for'

the unit during the

�-47-

spring and summer months,
Mourning

promptly migrating

with the first frosts.

doves have been observed throughout

the unit but are more

abundant .at lower elevations

Abundance

of mourning

Wildlife,

Hildlife

and in association

with agricultural

land.

doves varies seasonally.

Colorado Division

of

Inventory Systems

(197/+) gives a range of densities

from 0.86 birds per square mile to 689.8 birds per square mile, the
latter being a migratory

density.

CHUKAR
According
Systems
11.

to the Colorado Division

of Wildlife, t.,rildlifeInventory

(1974) there are two small groups of chukar remaining

on Unit

The first group of 12 birds is located on a 1.6 square mile area

at the head of Morgan Gulch, the other group of 12 birds is located on
a 1 square mile area on Maudlin Gulch near Deer Canyon.

Both populations

are marginal.

PHEASANT
Pheasants

are found in close association

in Unit 11.

Swope

to very poor.
the pheasant

(1965) describes

with the agricultural

the pheasant range in Unit 11 as poor

W.C.C.'~ Krager, Smith and Roland
occurring'in

lands with-

two locations;

(1975 pers. comm.) show

along the White kiver from Crooked

Wash Creek to 3 miles south of Smith Gulch, and a small area surrounding
Axial.

These two areas contain only marginal

popUlations

of pheasants.

TURKEY
Wild turkeys are found just south of Juniper Mountain
mately 4 square miles, according
total population

to W.C.C. Roland

in an area approxi-

(1975 pers. comm.).

of turkeys in this area is marginal,

The

and very low at best.

�-48-

WATERFOtvr..
Waterfowl

can be found on Unit 11 where suitable habitat exists, mostly

on stock ponds.

Populations

and lVhite rivers.
cinnamon

are concentrated

around the bordering Yampa

The major breeding birds on these rivers are mallards,

teal, blue •.
Tinged teal, green winged teal, American mergansers,

and Canadian

geese (Boeker 1953).

The minor breeding species are pintails,

gadwalls,

goldeneyes,

and shovelers

(Boeker 1953).

W.C.G. 's Krager. Smith,

and Roland

(1975 pers. conun.) concurred with the species presented by

Boelre r ,

Winter populations

can be found wherever open water exists on the unit.

These winter populations
(Colorado DhYision

are very small compared to the sununer populations

of Wildlife, Wildlife

LITERATURE
Beekez., -,;li. 1~.

1953.

Waterfowl

~ste~s

thesis.

Inventory Systems 1974).

CITED

production

in the Yampa River Valley,

Colorado State University.

Colorado Division of ~i1dlife, Wildlife Inventory Systems.
·i.Ioi.ide-wfli1!M.;.&gt;J

and bird distribution

Grieb, J. R.

1965.

1974.

Located at Denver

Division of Wildl.

Unpublished.

Colorado long-range

game species management

1965-1975, for migratory birds.

State-

maps and data compiled from W.C.G.,

-%Ce.al!m~~-\ljsor, and regional director input.
.&gt; off"t~e;Colorado

117pp.

plans,

Colo. Dept. Game, Fish and Parks.

36pp.

Roger s , .J~. E,
No. 16.

1964.

Sage grouse investigations

Colo. Dept. Game, Fish and Parks.

in Colorado.
l32pp.

Tech. Pub.

�-49-

Swope, H. M.

1965.

1965-1975,
Parks.

Colorado long-range

for ring-necked

pheasant.

game species management

plans,

Colo. Dept. Game, Fish and

54pp.

B. D. Trind1e and W. T. McKean
November 1975

�-50-

ANTELOPE HARVEST , SEASONS ~ AND HUNTING P_RESSURE
WILDLIFE MANAGEMENT UNIT 11
(From Colorado Big Game Harvest 1956-1974)
YEAR

HUNTING
PRESSURE

BUCKS

HARVEST
DOES
FAWNS

TOTAL

HUNT AND SEASON

l./

1956

----------

1957

250

167

62

7

236

250ES, 9/21 - 9/23

1958

300

157

103

21

281

300ES, 9/20 - 9/22

1959

300

173

88

23

284

300ES, 9/26 - 9/28

1960

400

243

121

26

390

400ES, 9/17 - 9/19

1961

100

64

24

7

95

100ES, 9/23 - 9/25

1962

300

121

136

24

281

150BO, 150DFO, 9/22 - 9/24

1963

50

21

16

7

44

25BO, 25DO, 9/21 - 9/23

1964

100

25

49

18

92

25BO, 75DO, 9/19 - 9/21

1965

100

43

37

0

80

50LH, 50SH, 9/25 - 9/27

1966

150

74

58

4

136

75LH, 75SH, 9/17 - 9/19

1967

70

35

16

8

59

70ES, 9/16 - 9/18

1968

100

59

33

3

95

100ES, 9/28 - 9/30

1969

80

52

21

4

77

80ES, 9/27 - 9/29

1970

49

38

8

2

48

50ES, 9/26 - 9/28

1971

98

86

8

2

96

100ES, 9/25 - 9/27

1972

189

153

31

2

186

200ES, 9/23 - 9/25

1973

240

195

19

3

217

250ES, 9/22 - 9/24

1974

229

159

47

14

220

250ES, 9/28 - 9/30

NO DATA

---------------

250ES, 9/22 - 9/24

!/
Antelope management areas differ from Wildlife Management Units. The
following is a list of area descriptions for the above data which are
applicable to Wildlife Management Unit 11:
1956-1957

Area 2:

That portion of Moffat County lying south of U. S. Highway 40 from Deception
Creek to Maybell; and south and west of State Highway 318 from Maybell to

�-51-

the Little Snake River; and all lands south and east of the Little
Snake River from State Highway
River;

318 to its confluence

and all lands north and east of Cross Mountain

Hills; and all lands north and west of Deception
1968

with the Yampa
and the Danforth

Creek.

Area 2:

That portion

of Unit 3 south of State

11 west of Deception

Hf.ghway

318; that portion of Unit

Creek and east of the divides between Crooked Wash

and Coal Creek-\-lolf Creek.

1970

Area 2:

That portion of Unit 3 south of State
west of the Strawberry

Creek-Deception

Hf.ghway

318; that portion of Unit 11

Creek road, and north of the Moffat-

Rio Blanco County line.

1971

Area 2:

That portion

of Unit 3 south of State Highway

north of U. S. Highway

40, east of Middle Wolf Creek and Disappointment

Creek and south of Dinosaur
of Strawberry

318; that portion of Unit 10

Creek-Deception

National

Park; that portion of Unit 11 west

Creek road and south of the Moffat-Rio

Blanco

County line.

1972-1974

Area AS:

That portion

of Moffat County bounded on the north by State Highway

on the east by the Strawberry

Creek-Deception

318,

Creek road, on the south by

the Moffat-Rio

Blanco County line and on the west by Middle Wolf Creek and

Disappointment

Creek to the Yampa River, the Yampa River to the Little Snake

River and the Little Snake River to State Highway

318.

�-52Note:

Hunt symbols are explained as follows:

ES

Either sex.

BO

Bucks only.

DO

Does only

DFO

Does and fawns only.

LH

=

Horns longer thml ears.

SH

=

Horns shorter than ears.

B. D. Trindle
November 1975

�-53DEER HARVEST, SEASONS AND HUNTING PRESSURE
WILDLIFE MANAGEMENT UNIT 11
(From Colorado Big Game Harvest 1956-1974)

YEAR

l

HUNTING
PRESSURE 1/

BUCKS

DOES

HARVEST
FAWNS

TOTAL
--

HUNT AND SEASON

1956

4684

3733

2287

586

6606

2ES 10/15 - 10/31

1957

3870

2485

2059

723

5267

2DM 9/14 - 9/30, In Part
2DM 10/15 - 11/17

1958

1827

733

458

168

1359

ES 10/15 - 11/2

1959

1880

957

397

100

1454

ES 10/17 - 11/3

1960

2723

2107

982

338

3427

1DM 10/17 - 11/6
1DM 12/1 - 12/15, In Part

1961

3320

2886

2720

879

6485

3DA 10/21 - 11/8
1DM 12/2 - 12/15, In Part

1962

1715

855

332

59

1246

1DM 8/18 - 9/3, In Part
ES 10/20 - 11/4

1963

4074

2815

2230

546

5591

1DM 10/19 - 11/7
1DMA 11/23 - 12/1, In Part

1964

4367

2871

2329

466

5666

1DM 10/17 - 11/5, In Part
ES 10/17 - 11/5, In Part
1DM 11/26 - 12/13, In Part

1965

1690

446

320

126

892

ES 10/16 - 11/5
1DM 11/25 - 12/5, In Part

1966

1600

505

333

63

901

ES 10/15 - 10/31

1967

1791

751

320

53

1124

ES 10/21 - 11/9

1968

2010

1369

503

135

2007

ES 10/19 - 11/7, In Part
ES 10/19 - 10/27, In Part
1DM 10/28 - 11/7, In Part
ES 11/27 - 12/8

1969

3123

1226

669

210

2105

ES 10/18 - 11/6

1970

3615

2018

385

39

2442

1DB 10/17 - 11/6, In Part
ES 10/17 - 11/6, In Part

1971

3428

2101

2101

1DB 10/30 - 11/11

1972

4951

2702

1348

174

4224

ES 10/14 - 10/23
IDA 11/18 - 11/26, In Part

1973

5313

2039

1340

248

3627

ES 10/13 - 10/23
IDA 11/17 - 12/2, In Part

1974

2997

1556

1556

!DB 10/26 - 11/5

(Footnotes on following page)

�-54-

DEER HARVEST, SEASONS AND HUNTING PRESSURE (Continued)
WILDLIFE MANAGEMENT UNIT 11

1/
- Based upon combined total resident and non-resident license sales and
expressed in number of licenses, not including archery hunters.

Note:

Hunt symbols explained as follows:

ES

Either sex, one deer (one license).

2ES

Either sex, two deer (one license).

IDA

One deer, antlerless only (one license).

IDB

One deer, buck only (one license).

lDMA

One deer multiple, one must be antlerless
per individual).

lDM

One deer multiple, either sex (2 licenses and 2 deer per individual).

2DM

Two deer multiple, either sex (2 licenses and 4 deer per individual).

(2 licenses and 2 deer

3DA

=

Note:

The term IDM evolved into 2D, HC or two deer, hunters choice (2 licenses
and 2 deer per individual) and is synonymous.

Three deer, one must be antlerless
on 2nd license).

(2 licenses plus 3rd deer coupon

B. D. Trindle
August 1975

I

�-55LINEAR REGRESSION OF EITHER SEX DEER SEASONS 1:..1
WILDLIFE MANAGEMENT UNIT 11

3000

2500

2000

HARVEST

1500

•
1000

500

500

1000

1500

2000

2500

3000

HUNTING PRESSURE
Correlation
r

=

Coefficient

Intercept

0.9453

-45.8463

Slope

0.7017

.•

1/
- Either sex season data obtained
Pressure Section.

from Deer Harvest,

Seasons and Hunting

�-56ARCHERY DEER HARVEST AND SEASONS
lolILDLIFEHANAGEHENT UNIT 11
(From Colorado Big Game Harvest 1956-1973)

YEAR 1:./

BUCKS

DOES

FAWNS

TOTAL 1/

HUNT AND SEASON 1/

1956 i/

ES .10/1 - 10/111

1957

Regular Season Only

1958

ES 9/1 - 9/30

1959

4

2

o

6

ES 9il5 - 9/30

1960

o

1

o

1

ES 9/10 - 9/30

1961

2

2

o

4

3DA 8/26 - 9/10
3DA II/II - 11/26

1962

ES 8/25 - 9/23

1963

1

1

o

2

ES 8/17 - 9/8

1964

4

4

o

8

ES 8/15 - 9/13

1965

o

5

o

5

ES 8/21 - 9/12

1966

ES 8/20 - 9/18

1967

3

2

o

5

ES 8/19 - 9/17

1968

3

1

o

4

ES 8/17 - 9/15

1969

9

o

o

9

1DB 8/16 - 9/14

1970 ~/

8

5

o

13

1DB 8/15 - 8/31
ES 9/1 - 9/20

1971

7

o

o

7

1DB 8/21 - 9/12

1972

15

3

o

18

1DB 8/19 - 8/31
ES 9/1 - 9/20

1973

6

8

3

17

1DB 8/18 - 8/31
ES 9/1 - 9/23

1974

8

o

o

8

IDB 8/17 - 8/31
1DB 9/1 - 9/22

(Footnotes on following page)

�-5],-

ARCHERY DEER HARVEST AND SEASONS
WILDLIFE MA..1\lAGEHENT
UNIT 11

(Continued)

1:./
Year 1956 free p~rmits issued to hoIders of regular big game hunting
licenses by application only. Years 1957-1960. no issue of special permits;
archery hunting allowed to holders of valid deer licenses.
Years 1961-1974,
separate archery license regulations in effect.

JJ
Inconclusive harvest data omitted for years 1956-1958,
Does not include harvest by archers during gun seasons.

1962, and 1966.

3/
- Bow hunting permitted
Colorado deer license.

during all regular rifle seasons with regular

!i/
For years 1956-1969

no hunter pressure

data available

for Unit 11.

~/
Resident and non-resident archery hunters 1970-1974,
33; 1971 - 80; 1972 - 98; 1973 - 112; 1974 - 45.
Note:
ES
lDB
3DA

Hunt symbols are explained

=

=

by year were: 1970 _

as fo l l.ows :

Either sex, one deer (one license).
One deer, buck only (one license).
Three deer, one must be ant1erless (2 licenses plus 3rd deer coupon
on 2nd license).

B. D. Trindle and W. T. McKean
September

1975

�-58-

PRIMITIVE DEER HARVEST AND SEASONS
WILDLIFE MANAGEMENT UNIT 11
(From Colorado Big Game Harvest 1970-1974)

YEAR

1:./

HARVEST
FAWNS

BUCKS

DOES

1970

7

2

0

9

ES 9/12 - 9/20

1971

12

0

0

12

IDB 8/21 - 9/12

TOTAL

HUNT AND SEASON

1972

?::../

38

0

0

38

1DB 8/26 - 9/10

1973

1/

11

27

0

38

ES 9/8 - 9/16

1974

Not Open

1-'
Free permits issued to holders
app.l Lcat-fo.. only from 1970-1973.

of regular big ga~e hunting

2/
Eighty~'!f:f)H;resident

and non-resident

muzzle

loaders.

and non-resident

muzzle

loaders.

1./
N.inet:y~:i)';:: resident

Note!

Hunt symbols
Either

.._DE'

are explained

as follows:

sex, one deer (one license) •

.One deer,

buck only (one license).

B. D. Trind1e
September 1975

licenses

by

�-59TOTAL DEER HARVEST, DATA
ANALYSIS UNIT 7, 1963-1974, BY SEX 1/

YEAR

BUCKS

DOES

FAl\1NS

TOTAL

1963

16117

14999

5018

36134

1964

14736

12170

2946

29850

1965

8161

10127

2739

21027

1966

835'4

7558

2361

18313

1967

10582

8143

2150

20875

1968

11076

8468

2407

21951

1969

8108

5493

1676

15277

1970

10335

3577

400

14312

1971

8945

0

0

8945

1972

12655

4870

554

18079

1973

9323

3978

642

13943

1974

7380

220

33

7633

1.1
Deer DAU Unit 7 incorporates Wildlife Management Units 11, 12, 13, 22,
23, 24, 25, 26, 31, 32, 33, and 34.

B. D. Trind1e
August 1975

�-60ELK H}u~VEST, SEASONS, AND IDJNTING PRESSURE
WILDLIFE HANAGEHENT UNIT 11 (DAU 11)
(From Colorado Big Game Harvest 1956-1974)

YEAR

HUNTING
PRESSURE

1956

188

20

8

1957 '!:./

94

12

4

1:./

BULLS

HARVEST
CO\V-S CALVES

TOTAL

HUNT AND SEASON

2

30

AO + 100 ESP'!:./
10/15 - 10/31

o

16

AO + 100 ESP 10/15 - 10/31

1958

CLOSED

1959

23

6

o

o

6

AO 10/17 - 11/3

1960

62

10

o

o

10

AO 10/17 - 11/6

1961

18

17

o

o

17

AO 10/21 - 11/8

1962

CLOSED

1963

CLOSED

1964

CLOSED

1965

CLOSED

1966

CLOSED

+ 70 C 10/21 - 11/9, In

1967

120

10

32

7

49

50 B
Part

1968

157

47

31

5

83

50 B + 70 C 10/19 - 11/7, In
Part

1969

496

118

67

7

192

AO + 50 C 10/18 - 11/6

1970

494

141

34

4

179

AO + 50 C 10/17 - 11/6

1971

398

68

16

4

88

AO + 40 C 10/16 - 10/25

1972

427

127

33

5

165

AO + 50 C 10/28 - 11/12

1973

567

110

20

1

131

AO + 50 C 10/27 - 11/11

1974

900

211

41

6

258

AO + 75 C 10/12 - 10/22

1--/
Based upon combined resident and non-resident license sales and expressed in
number of licenses; not including archery hunters.
During the period 1956-l970~
when elk season coincided with deer season, the hunting pressure statistics were
very probably conservative.
Despite difficulties verifying that assumption, unknown substantial numbers of deer hunters may also have carried elk licenses hoping
for a bull elk under antlered-only regulations.
This latter happened regardless
of low elk popu1ati~ns.
Thus, because of low success and tendency (above non(footnotes

continued

on following

page)

�-61-

ELK HARVEST, SEASONS, AND HUNTING PRESSURE
WILDLIFE MANAGEHENT UNIT 11

(Continued)

1:..1 continued.
reporting statewide figures used in projections) for non-successful elk
license holders not to report by card, elk present in Unit 11 probably
received extra pressure over that indicated by card projection years of
antle~ed-only seasons.

JJ

Units 11, 22, and 32 combined.

Note:

Hunt symbols are explained as follows:

AO

Antlered

only.

AO + ESP

Antlered

only, plus either sex permits.

B .;- C

-

Bul:: (antlered) and cow (antlerless) permits only (specified).

= ..lmtlexed only plus specified

number of cow permits.

B. D. Trfudle
September 1975

�-62ARCHERY ELK HARVEST, SL\SONS AND HUNTING PRESSURE
WILDLIFE MANAGEMENT UNIT 11
(From Colorado Big Game Harvest 1967-1974)

YRAR

HUNTING
PRESSURE

1967 ]j

--------------------

1968

6

1969

BULLS

HARVEST 1/
COYJS CALVES

HUNT AND SEASON ]j

TOTAL

NO DATA --------

ES 9/2 - 9/17

0

0

0

0

ES 8/17 - 9/15

3

0

0

3

AO 8/16 - 9/14

1970

33

5

3

0

8

AO 8/15 - 8/31
ES 9/1 - 9/20

1971

12

3

0

0

3

AO 8/21 - 9/12

1972

23

0

2

0

2

AD 8/19 - 8/31
ES 9/1 - 9/20

1973

78

3

3

0

6

AO 8/18 - 8/31
ES 9/1 - 9/23

1974

26

0

0

0

0

AO 8/17 - 8/31
ES 9/1 - 9/22

1/
Includes harvest only during specified archery seasons.

~/
Elk bow hunting permitted during all regular seasons with regular Colorado
elk license.
3/
- First statewide elk archery season with archery elk license.
Notes:

Wildlife Management Unit 11 (Strawberry) and elk "Data Analysis Unit"
11 are identical.
Hunt symbols explained as follows:

ES
AO

Either sex, one elk (one license).

=

Antlered only, one elk (one license).

B. D. Trindle and W. T. McKean
September 1975

�-63BLACK BEAR HARVEST AND SEASONS
WILDLIFE HANp.GEMENT UNIT 11

1/

HUNT AJ'IDSEASONS

2;/, 1/, i/

YEAR

HARVEST

1956

11R

5/15 - 8/15; pre, post and regular deer and elk
seasons.

1957

3R

4/1 - 11/15; pre, post, and regular deer and elk
seasons.

1958

2S

4/1 - 11/15; pre, post, and regular deer and elk
seasons.
.'

1959

0

4/1 - 11/1; pre and regular deer and elk seasons
ending 11/1.

1960

5S

4/1 - 9/15; pre and regular deer, elk, and archery
seasons ending 11/6.

1961

5R

4/1 - 9/15; pre and regular deer and elk seasons
ending 11/8.

1962

0

4/1 - 9/15; Ere and regular deer seasons ending 11/4.

1963

I.R

4/1 - 9/15; Ere and regular deer seasons ending 11/7 .

1964

0

4/1 - 9/15; Ere and regular d.eer seasons ending 11/5.

1965

0

4/1 - 9/30; Ere and regular deer seasons ending 11/5.

1966

6S

4/1 - 9/3°2 Ere and regular deer seasons ending 10/31.

1967

0

4/1 - 9/30; regular deer and elk seasons ending 11/9.

1968

3S

4/1 - 9/30; regular deer and elk seasons ending 11/7.

1969

0

4/1 - 6/30; regular deer and elk seasons ending 11/6.

1970

IS

4/1 - 6/3°2 regular deer and elk seasons ending 11/6.

1971

0

4/1 - 6/3°2 regular deer and elk seasons ending 11/11.

1972

3R

4/1 - 6/30; regular deer and elk seasons ending 11/12.

1973

1R

4/1 - 6/30; regular deer and elk seasons ending l1/ll.

1974

0

·4/1

- 6/30; regular deer and elk seasons ending 11/5.

1/

R = Regular big game (deer and/or elk) licenses; S = Bear license for special
spring, summer, or spring-summer seasons; also see annual regulations for archery
hunting.

(Footnotes

continued

on following

page)

�-64-

BLACK BEAR HARVEST AND SEASONS (Continued)
WILDLIFE MANAGEMENT UNIT 11

From 1956-1959,.1 bear per special bear license and/or 1 bear per bear
coupon on either or both deer and elk licenses; - from 1960-1965, same as
1956-1959, except special bear license variously invalid after September
15 or 30, when bear coupon on either or both deer and elk license covered
bag and possession limits; - 1966, same as 1960-1965, except one bear,
hunter's choice, per person per calendar year; - from 1967 to 1975, bear
coupon removed from regular deer and elk licenses; thus 1 bear, hunter's
choice, per special bear license, or sportsman's license, per person per
calendar year on!z during special bear and regular deer and/or elk seasons.

11
Up to eight dogs permitted except when bear seasons concurrent with deer
and/or elk seasons. Beginning in 1970 to date adult bears with one or more
cubs were protected.

~/
Data taken from Colorado Big Game Harvest and Commission Meeting Minutes.

B. D. Trindle and W. T. McKean
September 1975

�-65MOUNTAIN LION HARVEST ~~ SF~~SONS
WILDLIFE K&lt;U\,AGEMENTUNIT 11
YEAR

1=-'

HARVEST

HUNT AI'm SEASON

1965

2F 2:/
(Rio Blanco Co.)

10/16/65 - 3/31/66 ~Jest Slope; 10/23/25
3/31/66 East Slope, 1 either sex.

1966

3M
(Moffat Co.)
3M', 3F 2:/
(Rio Blanco Co.)

_

1/1/66 - 12/31/66 in all counties west of the
Continental Divide and in Jackson, Conejos,
Alamosa, Mineral, Saguache, Rio Grande,
Costilla, Archuleta, Hinsdale, and San Juan
counties; 3 either sex.
10/22/66 - 2/28/67 in all counties east of
the Continental Divide except those specifically listed above; 3 either sex.

1967

No Data

1968

o 1/

9/1/68 - 3/31/69 stateWide;

1 either sex.

1969

1M; IF 1/

9/1/69 - 3/31/70 statewide;

1 either sex.

9/1/67 - 3/31/68 statewide; 9/1/67 - 5/21/68
west of State Hwy. #13 and north of U. S. Hwy.
#6' 1 either sex.

1970

0

No season.

1971

0

9/1/71 - 10/12/71; 11/12/71 - 12/31/71
signated areas; 1 either sex.

1972

2M; IF

2/1/72 - 4/30/72 designated areas; 1 male
only; 10/14/72 - 10/23/72 and 10/28/72 11/12/72 statewide; 1 either sex; 11/13/72 12/31/72 designated areas including Unit 11,
S either sex permits.

1973

IF

2/1/73 - 4/30/73, 5 either sex permits for
Unit 11; 10/13/7310/23/73 and 10/27/73 _
11/11/73 statewide; 1 either sex; 11/12/73 12/31/73 designated areas including Unit 11,
5 either sex permits.

1974

o

de-

2/1/74 - 4/30/74 designated areas including
Unit 11, 5 either sex permits; 10/12/74 _
10/22/74 and 10/26/74 - 11/5/74 statewide;
1 either sex; 11/9/74 - 4/30/75 designated
areas including Unit 11, 4 either sex permits.

!/
Bounty of $50 per lion paid during period May 7, 1929 - March 12, 1965,
hence no protection.
Protected by statute as of July 1, 1965 when mountain
lion license was established.
(Footnotes

continued

on following

page)

�-66-

MOUNTAIN
WILDLIFE

LION HARVEST AND SEASONS
MANAGEMENT UNIT 11

(Continued)

:?:./
Credited to "1965-1966" and "1966-1967" for Rio Blanco and Hoffat
-----counties as reported by Dixon, K. R. 1967. Evaluation of effects of
mountain lion predation, p. 147; In Game Res. Rep., July - Part II.
Colo. Div. G&amp;~e, Fish and Parks, ~- 73-310. No data available in reports from se~tion of Game Management for 1965-1967.
Dixon's information is probably conservative in that the damage law al Lows for lax
reporting of kills of marauders by stockmen.
For the years 1968-1974
data are from Colorado Big Game Harvest Surveys.

]/
Three lions taken and reported each year by BSF&amp;W from Rio Blanco
County in 1968 and 1969.

B. D. Trind1e
September

1975

�SMALL GAME HARVEST AND HU1~ING PRESSURE
SMALL GAME MANAGEMENT UNIT 8, 1968-1973

1/

SPECIES 2:./

1968

NUMBER OF HUNTERS
1969 1970
1971 1972

1973

Average 2/

Cottontail
rabbit

1984

1328

1319

1411

1502

1070

1435

Snowshoe
hare

101

30

73

95

244

253

133

134

78

Blue grouse

298

154

213

124

442

140

229

1172

73

368

Sage grouse

190

301

258

267

537

183

289

603

767

Chukar

122

453

151

223

NOS

NOS

238

76

Gambe1' s
quail

27

NOS i/ 61

NOS

NOS

NOS

44

Pheasant

313

621

250

256

71

274

129

1968

1969

HARVEST
1971
1972

1970

1973

Average

7139

13415

838

1575

678

78

-815

144

Lf42

724

597

1134

584

735

606

247

358

NOS

NOS

332

31

NOS

153

NOS

NOS

NOS

92

586

1040

127

246

446

212

443

24813 11606

9276 14148
5/

13511
5/

cr 1403 -

I

aN

I

Band-tailed
pigeon

Nosi/

NOS

NOS

NOS

NOS

NOS

--

NOS

NOS

NOS

NOS

NOS

NOS

Mourning dove

NOS

276

290

242

447

107

272

NOS

5828

2412

2091

4373

920

3125

1/
- During years 1968-1973 Small Game Management Unit 8 contained the White River drainage of Wildlife Management Unit
11. Small Game Units were changed in 1974. Thereafter, S.G.M.U. 20 contained White River drainage portion of
W.M.U. 11.
.
2/
.
- Game Management Section questions the ability of hunters to distinguish between spec~es of rabbits and hares, and
the grouse family, thus adding inconsistency to the above data.
3/
- Mean for years data shown.
4/
- NOS means no data or no open seasons.
5/
- Possible sample size error.
B. D. Trind1e
September 1975

�SMALL GAME HARVEST iL\~ HUNTING PRESSURE
SMALL GAME ~~AGEME~~ UNIT 6, 1968-1973 11
NUMBER OF HUNTERS
1970
1971 1972

];/

1968

1969

Cottontail
rabbit

1763

1417

1532

2233

3869

Snowshoe
hare

NOS !!.j

NOS

NOS

NOS

Blue grouse

119

106

61

Sage grouse

612

1351

Chukar

49

Gambel's
quail
Pheasant

SPECIES

HARVEST
1971
1972

Average 1/

1968

1969

1970

678

1915

16977

18514

20589 35786

NOS

36

36

NOS

NOS

NOS

59

110

NOS

91

164

200

1635

1131

2470

988

1364

1967

69

NOS

NOS

NOS

NOS

59

NOS

NOS

NOS

NOS

NOS

NOS

815

483

388

518

103

NOS

1973

1973

Average

43533

3904

23217

NOS

NOS

716

716

177

182

35

NOS

151

5656

3896

3857

6528

2153

4010

313

27

NOS

NOS

NOS

.NOS

170

--

NOS

NOS

NOS

NOS

NOS

NOS

461

1026

1337

380

480

781

NOS

801

I
0\

co
I

Band-tailed
.pigeon

NOS

NOS

NOS

NOS

NOS

NOS

--

NOS

NOS

NOS

NOS

NOS

NOS

Mourning
dove

NOS

628

290

948

485

107

491 .

NOS

9065

2412 10158

8268

601

6100

l/During years 1968-1973 Small Game Management Unit 6 contained the Yampa River drainage of Wildlife Management Unit
11. Small Game Units were changed in 1974. Thereafter, S.G.M.U. 16 contained the Yampa River drainage of
W.M.U.ll.
l/Game Management Section questions the ability of hunters to distinguish between species of rabbits and hares, and
the grouse family, thus adding inconsistency to the above data.
l/Mean for years data shown.
4/
- NOS means no data or no open season.
B. D. Trindle
September 1975

�-V;.1.-

SMALL GAHE HARVEST AND HUNTING PRESSURE Y
SMALL GAHE NANAGEHEl'."'T
UNITS 16 AND 20 1974
NO. OF HUNTERS ]/
UNIT 16
UNIT 20

SPECIES Jj
Cottontail

rabbit

HARVEST
UNIT 16
UNIT 20

814

98

4704

492

Snowshoe hare

30

0

30

0

Pine squirrel

0

0

0

0

Blue grouse

66

31

130

0

Sage grouse

955

125

1807

341

Chukar

NOS il

NOS

NOS

NOS

NOS

NOS

NOS

NOS

30

0

91

0

NOS

NOS

NOS

NOS

441

44

3697

720

64

0

158

0

Marmot

66

33

392

328

Badger

64

33

64

0

Red fox

142

0

426

0

Bobcat

33

0

64

0

Raccoon

33

0

33

0

Coyote

307

98

186

492

Prairie dog

75

178

8205

5166

Jackrabbit

370

30

2308

122

Crow

0

0

0

0

Magpie

130

0

1255

0

Gambe1's

quail

Pheasant
Band-tailed
Mourning

pigeon

dove

Sharp-tailed

grouse

11
- Preliminary

analysis by Game Management

Section.

21
- Game Management Section questions the ability of hunters to distinguish between species of rabbits and hares, and the grouse family, thus adding inconsistency to the above data.
(Footnotes

continued

on following page)

�-70-

SHALL GAHE HARVEST AND HUNTING PRESSURE
SHALL GAHE MANAGEl1ENT UNITS 16 and 20

(Continued)

1/
Three percent sample size; 33 projected hunters and 33 harvested
one actual hunter reporting a harvest of one animal.

4/
- NOS means no data or no open season.

B. D. Trindle
September 1975

equals

�-71DUCK AND GOOSE HARVEST AND HUNTING PRESSURE
RIO BLA!~COAND NOFFAT COUNTIES l 1955-1973 1:../
(From Colorado Waterfowl Kill Surveys)
YEAR

DUCKS
EST. NO HUNTERS

EST. HARVEST

GEESE
EST. NO HUNTERS
EST. HARVEST

1955

420

3158

361

154

1956

NO DATA

1281

NO DATA

57

1957

NO DATA

2750

NO DATA

NO DATA

1958

236

1236

NO DATA

NO DATA

1959

322

1610

NO DATA

NO DATA

1960

112

575

39

NO DATA

1961

96

553

34

34

1962

293

1313

99

132

1963

185

666

119

57

1964

256

760

105

78

1965

162

464

130

210

1966

271/

1027

173

150

1967

181

609

125

234

1968

239

2023

296

155

1969

145

951

157

203

1970

294

1937

200

198

1971

262

1332

172

131

1972

232

2649

-170

293

1973

142

1405

121

36

!/
Waterfowl kill data are available only by county, 1955-1973.
2/
- Obvious inaccuracy.

About 127 hunters would be more correct.

�-72DUCK AND GOOSE HARVEST A1~ HUNTING PRESSURE
SMALL GAME MANAGEMENT UNITS 16 &amp; 20

YEAR

DUCKS
EST. NO. HUNTERS

EST. HARVEST

1974

204

1497

1974 II

GEESE
EST. NO. HUNTEJ~S
22

II

EST. HARVEST

o

- Prior to 1974 waterfowl kill data were available only by county. 1974
is the first year hunting pressure and harvest are available by Small
Game Hanagement Unit.

B. D. Trindle
September 1975

��z

I-

::&gt;

z

2:

0"&gt;

I"-

ro
W

1

W

&lt;C

o

�-7~-

BIG GAME POPULATION DYNAMICS
WILDLIFE MlU~AGEMENT UNIT 11

Computer population simulations of elk and deer herds have become a
basic management

tool.

Population data, including the trend of the

herd, are input parameters of the computer model.

The simulated popula-

tion parameters represents the manager's hypothesis of the dynamics of
the herd.

The follo,ving graphs (1 and 2) in which the elk and deer herds of wildlife management Unit 11 are contained are based on data analysis units
(see Harvest Data).

The four graphs of each herd are explained as

follows:
A

Birth of young

June 1

B

Count one

September 20

C

Count two

December 11

D

Population at end -

May 31

B. D. Trindle and W. T. McKean
November 1975

�GRAPH

1

............ __ E:q\_lp1l.T~ ANALYSIS UNIT E-5)
WILDLIFE

-------- ..3000~C
- _ .. _.'
_._---

MA."JAGENENT UNIT 11

••••••=s•••••amaa.aa=.mD~.e••n=.m=mncm.D=Q~=;mc~DZ.naz~~aa~m~zaD=nm~$=~~a~~m$c~~mr.~~m~M~re~.~msnnG.m~s
l' ...---1·-··--1----1--l.
1
1
•

1

1

1

1
---~-

......•~.....

=---..

2700.0

1

1

1

1

1
1

1
1

1

"1

1
1

1

------

-

__ ao_:

-------------------.;; - "--'-----_.-'.

A
S
C

-

-

------------~.o~~~=---------·------------~,.---------------------------------------~----------------__~ -0

------------------;.
2100~O

C

:-_..

B

---------------------~----.=

-~
1\.------------

------------;.:---________________1800.0
_
_:

-------...,SllO;-a--

-,
E-or
••
-

A

::

.

i

A

-~----

~-

=

::::

~~

A

--

A

0

-

300.00

:=1 ~

0

-'--- :;;~ ~
~'j

::

I

...• ....,

C

.----=-

0

~

t.1

-~

~

H

0
Z

t::1

0

I)

~
- tr3
- o 0"d ---!.-.--.J
~

B

~

0
0

-- :::::~

=

H

~

::=

0"

--:
----;

-

;--------

:------_ .. - - _._---

-~-----------------------9.

S

E

9

~ueu,~O~---E----

------------------------:

A
610.

C'

H

-

A
A
•• ·13-----------:-=-

A

tIl

----:

ca
o

,.

--------------------1200.0
=------------_._._'-900.00
___________________________

._---::-

1
1
1
1
!
1
·1------·1----:---1-

111
111
-1-

'1

1---'

1
1
t
1
1----1-

1
1

-----

_
:-----

a.a ••••••• s~.ec.~~me~fi~~mmID~~aamneo~~me~==~~M20~~.nm=~D~mn~e~~~~~~m~=U~~~Ecm~mm=~~=nm~m~~R~~m~eoocsmn

------------'t9bOo~·--

.-1962.0

968.

196~o

1966.0

1970.0

972.o---- ..-----!976·o·0980.
1974.0
19?i3.0

•.••
0----

T"IMe::---------------------------------(A~ POP AT STA~T. &amp;~ POP AT CNTONE.

c- POP AT C~TT~~~ D. POP ~T [NO)

�GRAPH

DEER (DATA,ANALY~~LtrNIT ~

~

,.,.ar.,..:tamK~~H~=r~!~q.~~~!
•.w.IJ-]J3="1.t",.l~~
.•j~.?i2~t,,,,~J:f"?:!!*~~~,,~2~&gt;!2:~,,;z'J.~~,"~J:!::~"1,,,citJ.4a

" ••
u:mo ••

2.00COOE+OS

:

1

=

;

All

1

1

1

1

111

1

1

1

~

1

1

1

1

1

1

1

--;------

1

1

1

:

.1'

~

------

1.aOOOOE+OS :----

---

--------

-

-

B
I-;60000E·OS
___

.....;:....-

A

:----

-- ... =

A

~-

C
::

A

A

B

8

A
A

B

:----

B

A
A

B
1.40000E+OS

a

B

A

A:=

e

B
1.20000E·OS

_

..

=:---.

~
::

.--~=

_

:=

-

,

o

_

c

e

----:

---

.

'oz"

~
%

••
--r- ~
'-J

aocao.

~=

:---=

.

~

=
:---~
=
=

60000.

•

;0

I

~

e
t)

~

----:

~
~

:____
-

_

It

"

~--f.

3:
n

~
~
~
~

•
;;:

H

••

Vl ---

2

-~=

..
o

co~"
;;

-=

-;
--

-

_

--;----

20000.

-

:

=
:=
=

o.

=2=Cs.DaaaB.ca.a.==.==

-----------..."",=0.

1962.0

1
1

1

1

----2

~

:

:;

=

=

=

:=

••~•••• ~==.a.aa.DaC.C"ne.c==~eEa~aft~sa=~~m=~~=======2:~s=~ae~:mM£mnD=.~Da=~as==

1
Q 61+;'0

0

1
1

197z-;1)

66.
0

1966.0

1970.0

r97()";o
1974.0

f980.0

19ie.o

-----------------------'---~HfE------------------_

,."..•_

•

0

~
~~
~

_..•.._ J.~;;
..EQP A~~§TA.el!J.!..,!g.P~-~.,,~~.!q_~~~.~9~i\L£.::H~¥lQ.L...Q!.....e.9~L.~.t!.9)'__ _

x

~

~'

�GAME SPECIES INTRODUCTIONS - WILDLIFE MANAGEME~~ UNIT 11

Species

Date

Males

Females

Total

Release Area

Source of Stock

Elk

x-x-64

-- NO DATA --

15

T4N, R95W, Section No Data

Estes Park

Chukar

3-17-67

49

51

100

T4N, R93W, Sections 9 and 10

Little Hills

Hungarian
partridge

4-30-68

94

94

188

T6N, R92W, Section 29

F.C.W.R.S.

Hungarian
partridge

5-01-68

24

24

48

T6N, R92W, Section 29

F.C.W.R.S.

Hungarian
partridge

4-19-69

-- NO DATA --

389

T5N, R92W, Sections 24 and 25

F.C.W.R.S. (374)
C.S .U. ( 15)

Hungarian
partridge

4-19-69

Hungarian
partridge

4-24-70

117

4-24-70

135

Hungarian
partridge

-- NO DATA --

240

T5N, R92W, Section 16

F.C.W.R.S.

117

234

T5N, R93W, Section 35

F.C.W.R.S. and
Little Hills

135

270

T4N, R95W, Sections 30 and 31

F.C.W.R.S. and
Little Hills

Literature References
Colorado Division of Wildlife.

Denver office, Game Management Section, game specLe s introduction file.

Hoffman, D. M. 1973. Study of Hungarian partridge adaptability.
Federal Aid Project W-37-R, April, Part II. pp. 191-215.

Colo. Div. of Wildlife, Game Research Report,

I
'-I

00
I

�1

HABITAT RESTORATION PROJECTS - WILDLIFE MJu~AGEMENT1~IT 11 VEGETATIVE
TYPE

NO. ACRES
TREATED

PURPOSE OF
TREATMENT

DATE

LOCATION

1941

Strawberry Cr.
Scattered in TIN R94W, T2N
R95W, T3N R93, 94W

Sagebrush

Seeded with Agropy~on cristatum,
Agropyron smithii, Bromus inermis,
Melilotus officinalis, Melilot~s
alba.

360

Increase
forage

BLM

1942

Strawberry Cr.
T3N R93W, 94W

Sagebrush

Seeded ~"ithAgropyron cristatum,
Agropyron smithii, Melilotus
offieinalis.

500

Increase
forage

BLM

1942

Morgan Gulch - Yampa River
T4N R94W, TSN R91, 92, 93,
95W

Sage
brush

Seeded with Agropyron cristatum,
Agropyron smithii, Bromus inermis,
Melilotus offieinalis.

333

Increase
forage

BLM

1944

Dry Fork - Piceance Cr.

Sagebrush

Seeded with Agropyron eristatum,
Agropyron smithii, Bromus inermis,
Melilotus offieinalis, Melilotus
alba.

233

Improve
forage

BLM

TREA~~NT

AGENCY

I

"-J

\0

I

1944

Morgan Cr. - Yampa River

Sagebrush

Seeded with Agropyron cristatum,
Agropyron smithii, Bromus inermis,
Poa bulbosa, Melilotus offieinalis,
Meli10tus alba.

450

Increase
forage

BLM

1959

Coal Cr. - Crooked Wash Cr.,
White River
T4N R98W, T5N R98W

PinonJuniper

Chain one way.

607

PinonJuniper
control

BLM

1965

Yampa River
T6N R96, 97W

Sagebrush

Chain and seed (aerial broadcast)
with Agropyron cristatum.

52

BLM
Sagebrush
eradication

1/
If details are desired on any of these projects, make requests directly to Mr. Ron Kufeld, Colorado Division
of Wildlife, Research Center, Fort Collins, Colorado 80522.

�-80MANAGEMENT PROBLEMS

A.

CHECKLIST

-

WILDLIFE

MANAGEHENT UNIT

11

Present
1.

Winter grazing by domestic livestock.
A large portion of Unit
the western half).

11

has winter grazing permits

(primarily

Both c8ttle and sheep utilize the range

during the winter, thus reducing critical browse necessary

for

wildlife.

2.

Deer-auto

collisions

along

Hf.ghway s

13 and 64.

Deer from the Unit 22 herd cross both highways enroute to the
Williams

Fork summering area.

The Unit

11

wintering

cross Hf.ghway 13, coming from the Williams Fork.
result in collisions

between deer and automobiles.

number of collisions

occurs along

Hf.ghway

64.

herd must

Both migrations
The greater

In the fall and

spring many deer utilize the hay fields along the ~~ite River.
This results in a significant

number of Unit 11 deer getting

killed while crossing the highway to get to these fields.

3.

Lack of water at lower elevations.
There is very little running water, few stock ponds, or bodies
of still water on Unit 11 at lower elevations.

Available water

becomes critical during the hot, dry months of the summer.
additional

water developed

in Unit 11 would be beneficial

Any
for

wildlife.

B.

Future
1.

Coal development.
Coal development

will create many new problems for the wildlife

within Unit

The associated

11.

development will increase human

�-81-

and vehicular

activity in the unit.

as building,

fencing, and mining will also follow coal

development.

All these activities will restrict the natural

movement

Other activities

of wildlife and alter their migration

such

patterns and

life styles.

Many of the problems are still unknown.
affect wildlife?
industry?

How will strip mining

What are the water requirements

of the coal

What time table is involved in revegetation

All these unanswered
considerations

of spoils?

questions, and more will become management

in the future.

�•

-82--

...

•
BIG GN1E MAMXAL RESE/IRCH

REFERENCES

WILDLIFE MANAGDrEtIT UNIT t 1

•
•

BLACK BEAR
Denney, R •.N., and D. L. Gilbert. 1952. Fur resources and bear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 60 pp.
Gilbert. D. L. 1951. Fur resources study and bear investigati.ons. pp. 1-8.
~ Quart. Prog. Rep., July. Colo. Dept. Game and Fish. 102 pp.
• 1951. Fur resources study and bear investigations. pp. 1-9. In
---Quart.
Prog. Rep •• Oct. Colo. Dept. Game and Fish. 117 pp.
_____ • 1953. Fur resources and bear studies. pp. 59-60.
Rep, , Jan. Colo. Dept. Game and Fish. pp. I-HI.
1953. Deer-elk investigations.
April. Colo. Dept. Game and Fish.
Gilbert, P. F. 19q6. Bear studies.
Dept. Game and Fish. 4 pp.

In Quart. Prog.

p. 60. In Quart. Prog. Rep.,
pp. 1-99-.

In Quart. Prog. Rep., June.

1948. Deer-elk-bear investigations. pp. 30-34.
Rep., Oct. Colo. Dept. Game and Fish. 80 pp.
1952. Checking station survey. pp. 39-43.
Jan. Colo. Dept. Game and Fish. 60 pp.

Colo.

In Quart. Prog.

In Quart. Prog •.Rep.,

Gilbert, P. F., and G. E. Rogers. 1954. Deer-elk investigations. p. 127.
~ Quart. Prog. Rep , , July. Colo. Dept. Game and Fish. pp. 1-183.
Hammit, H. C. 1950. Deer-elk-bear investigations.
Proj. W~38-R, Oct. Colo. Dept. Game and Fish.

pp. 1-14.
21 pp.

Fed. Aid

lIay, K. G., C. N. Hunter, and L: Robbins. 1961. Big game manage~ent in
Colorado, 1949-1958. pp. 71-73. In Tech. Bull. No.8.
Colo. Dept.
Game and Fish. 112 pp.
Hunter, G. N. 1954. Deer-elk-bear seasons in Colo. 1948-1953.
Colo. Dept. Game and. Fish. 64 pp.
1966 •. Colo. big game harvest, 1959-1965. pp. 30-32.
Game, Fish and Parks. Spec. Hgmt. Rep. No. 1. 41 pp.

pp. 60-61.

Colo. Dept.

HcKean, W. T. 1965. Colorado long range game species management pbns,
1965-1975, for black bear. Colo. Dept. game, Fish and Parks. 6 pp.
Remington, J., et al. 1955. Fur resources and bear studies. Fed. Aid
Proj. W-61-R. Final Rep. April. Colo. Dept. Came and Fish. 85 pp.

�8
T!tncr, J. R., and D. L. Cilbcrt.A co~t;ibution toward a bIbliography
on the black bear. Tech. Pub1. No.6.
Colo. Dcp t , Game and Fd sh , 43 pp.

1960.

U. S. Dept. of Ar,r.

1965. Water and related land resources,
CoLo rado River
Basin in Colorado.
Coop. Study Rep. of Colo. Water Conserve Board and
U$S.1J.A. Econ. Res. Service, Forest Service, and Soil Conserve Service.
May. Denver. 183 pp. (looseleaf)

•
,11oycls R. Se 1970. Deer-elk investigations.
pp.133-213.
In Game Res. Re p ,,
July - Part IL
Colo. Div. Carae, Fish and Parks. pp. 127-286.

___ ~,_. 1970.

:Elk of the Hhite River Plateau, Colorado.
Colo. Divo Cane" Fish and Parks. 126 pp.

---,-

Tech. Publ. No. 25.

1972.
Elk, hlftite River experimental
management study.
pp.
In Game Res. Rep. , July - Part II.
Colo. Div. Wildl.
251 pp.

Res.

1973.
Rep.,

Experi.me;)tal elk harvest
regulations.
July - Part II.
Colo. Div. Wildl.

pp. 45-67.
230 pp.

'.
l35-16l.

'

In Game

Denney, R. N. 1965. Colorado long range ga~e species ~anage~ent plans,
1965-1975, for elk., Colo. Dpet. Gane, Fish and Parks. 61 pp.

Gilbert, P. F.o 1953.
RCp~t
4

Apri1~

1954.

-July.

Deer-elk investigations.
pp. 59-61.
Colo. Dept. Ga~e and Fish. pp. 1-99.

In Quart. Prog.

0

Deer-elk

investigations.

Co Lo , Dept. G&lt;:iDCand Fish.

pp. 125-128.
pp. 1-182 •

In Quart. Prog. Rep.,

•

Hammit, H. C. 1951. Deer-elk-bear investigations.
Rep., Jan. Colo. Dept. C~e and Fish. 33 pp.

pp. 16-22.

1!!. Quart. Prog.

llay', K. G., G. N. Hunter, and L. Robbins. 1961. Big game man2ge~ent in
~
Colorado, 1949-1958.
pp. 57-70. In Tech~ Pub1. No.8.
Colo. Dept.
Came and Fish. l12 pp.

lrunter, G. N.

19S~. Deer-elk-bear
Came and Fish. 64 pp.

·.0
seasons in ~olorado,

1948-1953.

---.,.. •

1966. Colo. big ga~e harvest, 1959-1965.
pp. 9-30.
Game, Fishoand Parks.
Spe,c. !-!gnt.Rep. No. 1. 41 pp.

Riord~n, L. E. ' 19~8.
Prog. Rep., Jan.
~

Colo. Dept.

Deer-e1k-bear investigations.
pp. 29-33.
Colo. Dept. Gaae and Fish. pp. 1-49.

,1948.
Deer-clk-bear investisations.
pp. 7-8.
Rep., April.
Colo.
Dept.
Gaoe
and
Fish.
pp.
1-54.
'

Colo. Dept.

In Quart.

In Quart. p!og.

.

• 1949. Dcer-elk-bcar investi~ations.
pp. 39-46.
----~Roep., July. Colo. Dcpt. Came and fish. pp. 1-98.

~

Quart. Pro~.

Rogers, G. E•. 1951. Checkinr, station survey. pp. 32-33.
)~ep•• Jan. Colo. Dcpt. Camc. and Fish. ~3 pp.

~

Quart. ProC.

tl1eston, J. v. 1962.
1957. pp. '70-78.

81 pp.

A resume of Color:tdo bir, ~:lmc research project!;, 1939In Tech. .I'ub l., No.9.
Colo. Dcp t , G;lmc and Fi!ih.

..

�-84-

V. S. Dept. of Ar,r. 1965.

Water ~nd related land resource~. Color~eo River
Basin 1n Colorado.
Coop. Study Rep. of Colo. Hater Con~erv. lloard and
V.S.D.A. Econ. Res. Service, fore5t Servicc, and.Soil Conscrv. Service.
May. Denver. 183 pp.
(looseleaf)

Uil1i~ms•.J.t et al. 1952. Location~d extent of winter ranges. pp. 57-62 •
.!!lQuart. Prog , Rcp , ; April.
Colo. Dept. Game and Fish. pp.1-62.
HODNT/,IN t.rox
~.----,--.--

Boyd , R. J .. 19(,5.
1965--1975, for

.

•.
.oJ.

Colorado long l:ange g ame species manag encn t plans,
predators.
Colo. Dc p t , Garnc , Fish and Parks.
21 pp •

Dixon, K. R., and R. J. Boyd. 1967. Evaluation of the effects of mountain
lion predation.
pp. 141-165.
In G~~e Res. Rep., July-II. Colo. Dept.
Games Ff.sb and Parks.
pp. 73--309.

u. S. Dept. of Agr.

1965. Water and related land resources, Colorado River
Basin in Colorado.
Coop. Study Rep. of Colo. Hate:::Co nserv , Board and
U. S. D.A. Econ •.R2S. Service, Fo res t Service, and Soil Ccns erv . Service .
. Hay. Denver. 183 pp.
(looseleaf)

HUJ,E DEER
e

Baker, B. D. 1955. Deer-elk investigations.
pp. 149-157.
In Quart. Prog.
Rep_, July - Part II. Colo. Dept. Game and Fish. pp. 117-175.
1956. Deer-elk investigations.
pp. 43-64. In Quart. Prog. Rep.,
July - .Par t 1. Colo. Dept. Cane and }'ish~ pp. 1-102.
1957. A directory of Coloraco big ga'":!ce~closures concerning th~ir
locations, studies and other gen~ral infor~ation. SFCC. Rep., Fed. Aid
Proj , U-38-?',
Ja.n. Colo. Dept. Came and Fish. 60 pp.
1959. D2er-elk investigations.
pp. 189-249.
Rep., July. Colo. Dept. Gane and Fish. 249 pp.

In Quart. Prog.

19EO. A detailed stuey of range forage by use of fenced exclosures.
pp. 1-10, 111-42. .!.!:. Quart. Rcp . , April. Co~o. Dept. Carae and Fish. 47 pp.
1961. A directory of Colo. big game exclosures concerning progra~
history, stuey techni~tles, locations, and other g2neral infor:::3tion.
Spec. Rep. Fed. Aid Proj. W-IOI-R, Work Plan I, Job No.2.
J.:lnuary.
Colo. Dept. Gauc and Fish. 51 pp.
1961. Carne range Lnvcs t Iga t Lons • pp , 71-72, 76-112.
July.
Colo. Dept. Game and Fish. 149 pp.
1962. G41me r'anr;c Lnvc s t Lga t Ions ,
·Co10. Dcpt. Came and Fish. 96 pp.

pp. 1-39.

In Quart. Rcp ,,

lE. Qua r t , Rcp ., AprIl.

�-85-

.

____

'

1962. Deer-elk investigations •. pp , 129-iJ8.
Colao Dept. Came and fish. pp. 129-249.
G

IE. Quart. Rcp ,, July •
•

Came range investigations.
pp. 3-6.
.------Colo.1964.
Dept. Camc~ FiGh and Pa~ks.· 121 pp.
o·

In Game Res. Rep., Jan •

1965. Came range investigations.
pp. 129-133.
Julyv Colo. Dept. Ga~e~ Fish and Parks. 249 pp.
e

-=-.
=~~__ 1965.. The history

In Game Res. Rep_,

t
function, and es t ab Lf shmerit of exclosures in
Colorado big game nanagemen t , Colo. Dept. Game, Fish and Parks. Outdoor
Fact.s No. 18. J pp.

pp • 53-67.
• 1966. Game r&lt;lnge investigations.
July - Part I. Colo. Dept. Gaoe, Fish and Parks.

In Came Res. Rep. ,
93 pp.

1970. Game range investigations.
pp. 15-58. In Carae Res. Rep. ,
Jul:r - Part I. ·Colo. Div. Garile,Fish and Parks. 126 pp.

~~."",--

Bart-mann, R. H. 1967~ Deer trapping and tagging results at the Little Hills
£xperi racnt; Statir,n.
Colo. Dept. Caae , Fish and Parks. Outdoor Facts
Uo&amp; ...5.~l.:

~~ .F'··'

1968 •. P'Lay Lng tag with the \,'hiteRiver deer herd.
-"·,.17(1):2D-22 e

Colo. Outdoors.

•
,,"_.,.1:965. ,. Results

. Colo.
o Comin.

from an 18 year deer tagging program in nor t hwe s t ern
pp. 161-166. 48th Ann. ConL t~est.Assoc. State Game and Fish
724 pp.

----.;&lt;ll~'~.

Mule deer migration at the Little Hills Game Exper-Iment
Station, Colo. p. 9. 13th Trans. Ann. SU~"':ler
Conf. C.X.P.S., The

llililli,£e

..$oc_:e'LY.

. \ '.

1972. 'Evaluation of the effects of spring-stl.;"':'ner
grazing by deer
on alfalfa. pp. 113-133. In Game Res. Rep., July - Part II. Colo.
Div. of Wildlife. pp. 71-251.
,'.,
___

1972. Pi'ceance deer study-population distribution.
pp. 317-337.
In Came Res. Rep., July - Part III. Colo. Div. of Hi!dlife.
252-377.
e

We

1972. Piceance deer study-population densit~. pp. 339-344.
Came tes. Rep., July - Part III. Colo. Div. of ~ildlifc •
.l&gt;p. 252-377.

•

.In

�-86-

1972. Piceance deer .study-productivity
In Came Res. Rep •• July - Part III.
Colo.

and ~ort31ity.
Div. of Wildlife.

pp.

345-350.

pp. 252-377.
• 1972.

Piceance
July - Part

-.fept.,

1973.
-'243-254.

deer
III.

study-food
habits.
pp. 351-355.
Tn Game Res.
Colo. Div. of Wildlife.
pp. 252-377 ~

Piceance deer study - p~1-;ulation density and structure, pp .
In Game Res. Rep , , July _.Part II. Colo. Div. Wildl.
230 pp ,

Piceance deer study - population distribution.
. 1973.
---242.
In Game Res. Rep., July - Part II. Colo. Div. Wildl • 230 pp.

pp. 211-

.

______ • '1973. Piceance deer study - productivity and mortality.
261. In Game Res. Rep., July - Part II. Colo. Div. Wildl.
~

pp. 255230 pp.

1974. Piceance deer study - population distribution.
pp. 325-362.
Game Res. Rep., July - Part II. Colo. Div. Wildl.
223 pp.

1974. Piceance deer study - productivity and mortality.
pp. 371-380.
In Game Res. Rep., July - Part II. Colo. Div. Wildl.
223 pp.
1974. Piceance deer study - population density and structure. pp.
363-370. In Game Res. Rep , , July - Part II. Co Lo , Div. Wildl. 223 pp.
1975. Piceance deer study - population density and structure. pp.
349-354. In Game Res. Rep., July - Part II. Colo. Div. Wildl. 314 pp.
1975. Piceance deer study - population distribution.
pp. 327-349.
In Game Res. Rep., July - Part II. Colo. Div. Wildl.
314 pp.
1975.
362.

»:lrto.lann,

179.

Piceance deer study - produc~ivity and mortality.
In Game Res. Rep ;, July - Part II. Colo. Div. Wildl.

pp. 355314 pp.

R. M., and W. T. McKean. 1968.' Game r anr;c invc!;tigations.
l!!. Game Res. Rep •• July -. Colo. Div. Game, Fd sh and Parks.

__----I nnd

1969.

Res. Rcp , , July.
Bockcr , U. M.

1959.

Summer ConL,

Game ran~e invcstigation5.
pp. 305-393.
Colo •. Div. Game, Fish and Parks.
393 pp.

C,olorado deer trapping.
pp , 52-55.
C.M.P.S -., TIle Wildlife
Society.

~th Trans.

pp. 167208 pp.
In Came

Ann.

�-87-

Boyd, R. J. ·1961. Study of deer losses OR .Co+orado highways.
pp. 75-82.
In Fed Aid Quart. Rep., April (Part 1). Colo. Dept. Game and Fish.
% pp. (processed) •

-.....---::- •

1962. Study of deer losses on Colorado highways. pp. 93-100.
Fed. Aid Quart. Rep., October. Colo. Dept Game and Fish. 137 pp.

In

(proc es sed) •

1964.

~ec1.

Study of deer losses on Colorado highways. pp. 36-41. In
Aid Qua r c , Rep., April (Part 1).
Colo. Dcp t , Game, Fish and Parks.
69 Pl'. (processed).
1970. Deer-elk investigations.
pp. 133-213.
July" Part II. Colo. Div. Game) Fish and Parks.

Burdick, H.

1963.

Deer tags tell a tale.

In Game Res. Rep.,

pp. 127-286.

·Colo. Outdoors 12(5):52-53.

Campbell, R. L. 1950. Little Hills Game Exper:i!:lent
Station. pp , 38--/f 3.
~ Quart. Prog. Rep., July •. Colo. Dept. Game and Fish. 80 pp.
1951. Experimental projects at Little Hills Experiment Station .
.-pp. 58-65. Spec. Rep. The Glenwood Springs Wildl. Conf. 85 pp.
______

a

Jan.

1951. Deer-elk investigations.
pp. 5-11.
Colo. Dept. Gane and Fish. 33 pp.-

1951. Deer-elk investigations.
April. Colo. Dept. Ga~e ana Fish.
___

In Quart. ~rog. Rep.,

pp. 50-53.
64 pp.

a . 1951.·
Deer-elk investigations.
pp . 71-77.
July. ~olo. Dept. Game and Fish. 101 pp.

1952.

Deer vs. livestock.

In Quart. Prog. Rep.,
~

Quart. Prog. Rep ,,

Colorado Outdoors 1(2):20-22.

1954. Deer-elk investigations.
pp. 149-154.
Ju~y. Colo. Dept. Game and Fish. 182 pp.
1956. Deer-elk investigations.
pp . 65-71.
July. Colo. Dept. Game and Fish. 102 pp. _

~

~

Quart. Prog. Rep.,

Quart. Pr og , Rcp ,,

. Campbell, R. L., and C. E. Rogers. 1950. Deer-elk surveys. pp. 20-21.
-- .!.!l Quart. Prog , Rcp ,, October. Colo. Dept. G~me and Fish. 21 pp.
Carh~rt. A. 11. 1940. Dcer-elk survey. Vol.~.
Colo. CW:1C and 'Fish Commission.
19 pp.

Fed. Aid Pro}, H-4-R.

�-88-

----:-Commission.
e.,1940.

Deer-elk aurvey, Colorado.

..

Vol. 5. 'Colo. Came anu Fish

50 pp.

____ ~.

1940. Deer-elk survey, Colorado.
Game and Fish Corrm Lss Lcn, 8 PP&gt;

.. Vol. 6. • Colo.

Tables XIV, XV.
.

.'

______ • 1941.

Deer-elk survey, Colorado. Vol. 5, Part I~ Deer food requirements in Colorado.
Colo. Came and Fish Comm. 28 pp.

______ • 1943.

The Piceance-White River deer ,herd. Deer-elk survey Supplemental Rep. Colo. Came and Fish Coraa i.ss Lon , 20 pp.

Denney, R. N. 1965. Colorado big game species as possible vectors of sooc
li.vestock.diseases.
Colo. Game, Fish and Parks Dept. Outdoor Facts
No. 20.
3 pp.

•
Denney, R. N., et a1. 1965. Came range surveys. pp , 17-23. Colo. Dept.
Game, Fish and Parks. Game Research Review, 1964. 35 pp.
Gilbert, P. F.

1948.

Prog , Rep, , April.

1948.
Rep., Juiy.

Deer-elk-bear investigations.
Colo. Dept. Game and Fdsh ,

Deer-elk-bear investigations.
Colo. Dept. Ga~e and Fish.

pp ,

pp. 14-18.
54 pp.

31-34.

In Quart.

In Quart. Prog.

83 pp.

• 1948. Deer-elk-bear investigations.
pp. 30-34. In Quart. Prog.
---Rep.,
October. Colo. Dept. Ga~e and Fish. 80 pp.
,

• 1949.

Deer-elk-bear investigations.
pp. 42-45.
Colo. Dept. Ga:ne and Fish. 67 pp. ,

In Quart. Prog.

-Rep., January.

pp. 47-51.
98 pp •

~

pp. 33-36.
84 pp.

In Quart. Prog.

1950. Deer-elk-bear investigations. pp. 41-~4.
Rep., January. Colo. Dept. G~e and Fish. 66 pp.

In Quart. Prog.

Deer-elk-bear investigations.
---,-.Rep.,1949.
April. Colo. Dept. Game and· Fish.
e

Quart. Prog.

.'

1949.

Deer-elk-bear investigations.
,Rep., July. Colo. Dept. Ga~e and Fish.

______

0·

1950.

Deer-elk-bear investigati?ns.
pp. 60-64.
Colo. Dept. G~ue and Fish. 101 pp.

Rep.
'. ______ •

In Quart. Prog.

1950. Deer-elk-bear investigations.
·1950. July. Colo. Dept. GaQC and Fish.

Three year Sum. Rep., '194721 pp.

1952~ Deer-elk investig~tions.
January. Colo. D~pt. Game andflsh.

39-43.

1952.
Quart.

pp.

.!..!!. Quart. Prog. Rep , ,

60 pp.

Deer-elk investir.~tions. pp. 41-64, 66-67, 69-82, 83-84.
Colo. Dept. Carne and Fish. 171 pp.

Pr og , Rcp , , July.

In

�-89-

__

--:-. 1953. Deer-elk Lnvcs t f ga t Lons, pp -.59-61.
April. Colo. Dept. Game and Fish. 99 pp.

____ ~.

.!.!l Quart. Prog , .Rcp, ,

1953. Deer-elk investigations.
pp. 51-63 • .!.!l Q~art. Prog. Rep.,
Colo. Dept. Cace and Fish. 129 pp.

July.

____~..
1954. Deer-elk investigations.
pp. 129-143.
July. Colo. Dept. Game and Fish. 182 pp. ,
Deer-elk investigations.
pp. 1-26.
Colo. Dept. Game and Fish. 111 pp •

.!.!l Quart. Prog. Rep.,

In Quart. Prog. Rep.,

....-,
.._. _" 1957. De er+e Lk investigations.
pp. 121-126~ 141, 145-1116. In
• Quart. Rep.» July. Colo. Dept. Game and Fish. 213 pp.

--_.

1958.

The deer's biggest problem.

. •...
,_.

'1960.

Where, to hunt-1960.

Colorado Outdoors 7(3):22-24.

Colorado Outdoors 9 (5): 5.

Gilbert~ P. F. et ale 1951. Deer-elk investigations.
pp. 109-110. In
Quart. Prog. Rep.~ October.
Colo. Dept. G~e and Fish. 117 pp.
,Gilbert, P. 'F.,

128. ~

and G. E. Rogers.
1954. Checking station survey.
Quart. Prog. ~ep., July. Colo. Dept. Gane and Fish.

Gilbert, P. F., et al. 1955.
Quart. Prog. Rep., July.

pp. 125182 pp.

Deer-elk investigations.
pp. 158-161.
Colo. Dept. Ga~e and Fish. 225 pp.

In

Gilbert, P. F., and R. L. Campbell.
1955. Deer-elk investigations. pp. 163170. ~ Quart. Prog. Rep., July. Colo. Dept. G~~e and Fish. 225 pp.
Gilbert, P. F., and J. Harris.
1958. Deer-elk investigations. pp. 131-138.
~ Quart. Rep.,. October.
Colo. Dept. Gane and Fish. 166 pp.
.• and
--'-=-8 (2): 18-20.

1959.

How far do deer travel?

Colorado Outdoors

Gordon, D. F., nnd R. }!. Bart~ann.
1971. Evaluation of the effects of
spring-su:::ncr\:razing by deer on alfalfa. pp. 99-112.
In Game Res.
Rep., July - Part II.
Colo. Div. Game, Fish and Parks.-pp.
87-224.
, lIammit, H. C. 1C)~,O. Deer-elk-bear investigations.
pp , 10-14.
Proj. U-38-R. October.
Colo. Dept. Ga~e and Fish. 21 pp.

1951. n~~r-cl~-bcar investigations. pp. 16-22.
W-38-R, Janll;\l"Y.Colo. Dept. Game and Fish. 33 pp.

Fed. Aid

Fed. Aid Proj.

1950. Dccr+clk-bca r investigations.
pp. 35-37 • ..!.!l Quart. Prog. Rep., July. Colo. Dept. Came and Fi5h.

Hammf t , H. C., and 1'. F. Gilbert.

Robbins. 1961. Dig gamc manng crncnt I n
Colorado, 19~~-1958. pp. 71-73.
In Tech. ruble No.8.
Colo. Dept.
G:llllC
and·rJ::h. 112 pp ,

Ilay , K. C. t C. N. Uunter,and

",

L.

�-90Dcer-clk-benr ~cosons 1n Coldrodo, 1948-1953.
Colo~ Dept. Came and Fish. 64 pp.

Uuntcr, C. N. 1954.
60.

pp. 22-

• 1966. Colorado big g~e harvest, 1959-1965. pp. 9-30. Colo. Dept.
---Came~Fish
and P~rks. Spec. Memt. Rep. No.1.
41 pp.,
Jeep, F. T., and L. E. Riordan. 1947. Deer-elk studies.
Rep., January - Part IV. Colo. Dept. Game and Fish.
Kufeld, R. C.

1968.

Game range jnvestigations.

1. 'Colo. Div. Game, Fish and Parks.

~art

In Quart. Prog.
7 pp.

In Ga~e Res. Rep., July pp. 1-121.

1969. Game ran8c investigations.
pp. 299-304. Tn Carne Res. Rep
Colo. Div. Garne, Fish and Parks. pp. 249-393.

c ,

-:--:ruly - Part III.
• 1970.

---~--Suly -Part
_,_"

Game range investigations.
pp.. 59-94. In Game Res. Rep.,
I. Colo. Div. G~e, Fish and Parks.p~
1-126.

1971. Inventory of raanf.pul.a
t f.on projects in Colorado.
pp. 7-15.
In Game Res. Rep ,, July - Part 1. Colo. Div. Game, Fish and Parks.
1·-86. .
.

Wo

Laur Ldson, D. G. 1948. Game and forage expe rImcnr station.
Conment s 10(6):9,-10, 22.
Lind, C. 1957. Deer-elk investigations.
Colo. Dcpt. Game and Fish. 213 pp ,

pp -.147-151.

Colo. Cons erv ,

In Quart. Rep., July.

pp. 1~23. In Quart. Rep, ,
W. 1'.
1958. Deer-elk investigations.
. , July - Part 1. Colo. Dept. Ga~e and Fish. pp. 1-103.

'lScKcan,

,

1959. Game range inv.estigations. pp. 193-212.
~July. Colo. Dept. Gace and Fish. 237 pp.

• 1960.
-October.

Ga~e range investigations.
Colo. Dept. Gaoe ana Fish.

pp. 3-23.
103 pp.

In Quart. Rep.,

In Quart. Rep. J

1965. Pellet group deposition rates for captive deer.
Game, F~sh and Parks. Outdoor Facts ~!o. 24. 3 pp.
• 1967. Game range f.nv es t Lgat i.ons pp , 155-165.
----~uly.
Colo. Dept. Gane, Fish and Parks. 208 pp.
c

Colo. Dept.

In Game Res. Rep.,

1970.

Game range investigations.
pp. 7-9. In G~~e Rcs. Rcp.)
Colo. Div. Game, Fish and Parks. 126 pp.

July.

McKean. U. T., and 11. E. Burdick. 1963. Game range invcstigations. pp. 89115 • .!E. Game Res. Rep •• January. Colo. Dcpt. Game and Fish. 115 pp.
_,

nnd
Rcs ,

}~CP'I

• 1964. Cat:1Crange invcsti.gations. pp. 27-67. In Game
January.
Colo. Dcpt. Game, Fish and Par ks , 121 pp.

______ ' nnd
Res. Rcp , , July.

1965.

Came range invcstig~tion~.
pp. 147-139.
Colo. Dcpt , Camc, Fbh and Parks.
249 pp.

In G.:lmc

�-91and R. N.

McKcan~ \.1. T.,
transplant!;

County, Co Lo rado
lb. 1,2. 3 pp~

-~~-

~ snd

RCBo

1~:lI
...
t.mann ,
1906.
Su rv Lva I of moun t a In mohor-nny
s i.t c s , Littlc
Hill!;
Expc r Lncn t St.n t i on , J~10 Blanco
1965.
Colo. Dcpt. Carnc , F'Ls h and Pa r ks , Outdoor- Facts

on tt.lO

1969. Game rangc invcstigations.

Rep.~ July.

Colo. Div. G~ruc. Fish and Parks.

pp. 7.53-260. 1n Ci.1L:le
J93 pp.

____ ~__ , and
1971.
Dc er+Ldv cstock r eLa t Lo ns on a p Lnon+jun Iper range
:.tn no r t hvc s t crr-Colorado.
Final Rcpo r t , Colo. Div. Game, Fish and Parks .

..132 pp.
lkl(!;2.n)

T~~ arid D. Go Smith.

lI.

4710
.

.•

and

t

-Rep.,

(processed).

.../.
1965.

In Gace Res. Rep.s July.

pp. 1123Colo. Dept. Ga~e) Fish and Parks. 546 pp.
Came range

Lnves t Lga t Ions ,

• 1966.
Came range investigations, pp. 5-1,9.
July-.- Co Lo , Dcp t, Carae , Fish 2Gd Parks. 93 pp.

In Carne Res.

He Kcan , W. T., et 21. 1968. Game range investigations.
pp , 155-165.
Came Res. Rcp ,, ~TUly. Colo. Div. GaI&gt;1e, Fish and Parks. 208 pp •

In

..

llediu 10 D. E. 1958. Deer-E;lk investigations.
pp. 63-68.
January •..Colo. Dept. Carae and Fish. 131 pp.

In Job Co~p. Rep.,

1960. Physical site factors influencing annual production of true
mountain Eahoga~y~ C~rcocarJus 2cntan~s.
Ecology 41(1):454-460.
1960. References on t:!ethodsof neasuring production and uti1izatioa
---of. range
and pasture
f or age ,
Colo. Dept. Game and Fd sh Tech. ·Publ. 1:0.

•.....

6"

43 pp.
Colorado long range garae species raanagccerrt
Colo. uepto Ga~e, Fish and Parks. 15 pp.

plans, 1965-1975 &gt;

l!ustard, E. n., Jr. 1958. Deer-elk investigations, pp. 9-34.
April. Colo. Dep.t, Carne and Fish. 113 pp.

In Quart. Rep.,
..

1958. Bro~se differences in ex?~ricent2l ~astures under controlled
ptocking with cattle, sheep, and ~ule deer in northwestern Colorado.
H. S. Thesis. Colo. State Dniv., Ft. Collins. 105 pp.
.
1959. Stucy of browse reproduction in relation to controlled grazing
in cxpe rI•...
ental pastures. 1A Quart. Prog. Rep, , January. Colo. Dept.
Game and Fish. 41 pp •
.·}~ers, C. T. 1969~ An investigation of deer-auto accidents.
pp. 147-178.
1n Fed.
Aid Cane Res. Rep., July
Part 2 •. Colo. Div •.Game. Fish and
Parks.
(processed).
pp. ".03-1,37.
---Fed. 1970.
. An investigation of deer-auto accidents.
Aid Ga~e Res. Rep., July - Part 3. Colo. Div. Game, Fish and

Parks.
Neil,

In

(processed).

P. H.
1973. Winter
nortality of cule deer
in Par~c}lute
Crcek Valley,
Garfield County, Color-ado. ..!.E: The Colony cnviron::lcntalstudy, P.1racilute
Creek. Ca r f Lc ld County, Co Lo rado , Prcpared for Colony Dcv cloprnon t;
Operation

by

).973.

Part Ill.

Tho r n c

Eco Log

ic n I

Iuc t i t ut o ,

Boulder,

Co l o r ado

,

Aug us

t

�-93-

Nielson, J. L. 1966. Soil survey of experimental pas t ur cs , Little ·1!il1sCame
Experiment Station.
Report by U. S. Soil Con~crv. Service to Colo. Dept.
Game, Fish and Parks.
Unpublished communication.
42 pp.

Faure L. F. 1972.

~fovcrnents of mule deer in the Parachute Creek basin,
u!nter 1971-72.
In The Colony environmental study, Parachute Creek,
Garfield County, Colorado.
Prepared for Colony Development Operation
• by Thorne Ecological Institutc Boulder, Colorado. August, 1973.
p

Pnrt XII.

.
.e ,

Pojar,· T. H. 197J... Dee r+aut.o accident investigations.
pp. 227-318.
In
Game Res. Rep, , July - Part IlL
Colo. Div Game, Fish and Parks.pp. 227-366.
&lt;

• 1971. Evaluatioc of devices to prevent deer-auto accidents.
pp. 333-340. In Ga~e Res. Rep.) July - Part III. Colo. Div. Games
Fish and Parks-.- pp. 227-366 •

.
• 1971. Research - 1971 dee r+h.i.ghway11z!1ting proj eet report.
pp. 355-366." In Gawe Res. Rep .• July - Part III.
Colo. Div. Game,
Fish and Parks-.- pp. 227-366.
____

-

• 1972. Honitor potentially critical, deer-vehicle accident areas
statewide.
pp. 269-275.
I~ G~e Res. Rep., July - P~rt III. Colo.
Div. of Wildlife.
pp. 252-377.

____ ~.
1972.
Effect of lighted deer crossing signs on number of deer
killed by vehicles.
pp. 300-304. In Gace Res. Rep., July - Part III.
Colo. Div. of {-j'ildlife. pp. 252-377":
"

..

1972. An evaluation of deer-proof fence length required to prevent
deer movements on or across high speed h i ghvavs , pp. 305-310. In Carne
"'Res. Rep , , July.- Part III. Colo. Div. or i~ild1ife. pp. 252-377.
Reed, D. r .. 1971~ Investigation of one-way de~r structures. pp. 321-330.
In Game Res. Rep , , July - Part III. Colo. Div. Carae , Fish and Parks.
227-366.

pp.

197~: Deer underpass evaluation.
pp. 343~353" In Game Res. Rep.,
July - Part III. Colo. Div. Gane, Fish and Parks. pp. 227-366.
________
• 1972.
Investigation of one-way deer structures.
pp. 278-282. In
Game Res. Rep., July - Part III. Colo. Div. of Wildlife.
pp. 252-377.
______

___

___

1972." Deer underpass evaluation.
pp. 285-290.
In Game Res. Rep.,
July - Part III. Colo. Div. of Wildlife.
pp. 252-377.
0"

1972.
Effects of h Lghvay lighting on number of deer killed by
vehicles.· pp. 292-298.
In Ga~e Res. Rep., July - Part III. Colo.
Divo of Wildlife. pp. 252-377.
0

1972.
Effects of 8-foot fence angle in d Ivor t Lng deer from their
es t ab lfshcd direction of movcncn t • pp.
312-3)./1. In Came Res. Rcp ••
July - l'ar t lII.
Colo. Df v , of WlldJ He.
pp. 252-377.
0

�-93lUonl.:Jnt,t.
E.
19/,7.
A proposed cxperiment
~t Little
St.at Lon ,
CoJ.~. Cons. Cornrne n t s 10(1); 20-21.
__

._

1948.

lIills

Dcc r+c Lk+b ea r Lnv c s t i.ga t fon s , pp. JJI-30.
Colo. Dept. Came and Fish. 49 pp.

Rep •• January.
~

.

• 1948.

Decr-elk-bear
investis~tiG:':;.
pp. 41-4G.
R(~p.~ October.
Colo. Dept. Carne and Fish.
,SO pp •

In Quar t , Pr og ,

....• ."'"' ..•.

•.._~~__ • 19f;9.,
Hcpc~

De.er+c Lk+b ea r :Lnvcstigations
.. pp . 36--40.
Janum:y.
CoI o , Dept. Came and Fish.
67 pp.

•,~______ 1949. 1)eE:t'-el1:-be.a r invcs Lisa t Jo ns .
~cp •• April.
Colo. Dept. Cawe and Fish.
_____

19tj9.

o"

Dr.er--elk-·bear

Rep.~ October.
•.

___

-...

investigations.

Came Expc r Iracnt;

In Ouar t , Pr og ,

pp. 7/:-82.
97 pp.

In. C;'u2T.L

pp.

In Quart.' Pr og ,

Colo. Dept. Game aDd Fish.

86-90.

hog .

141 pp .

" 1:}56.
SE'lf~ctj_v~_ty
and u t il.f za t+on of three k ey species
of ranGe:
forage by cattle,
sheep and deer in ~estcrn
Colorado. M.S. TIlesis.
Colo. State Vniv., Ft. Collins.
196 pp.
1956.
prog ra:a,
0,

'

The results
of the Little Hills expe r iznent a I r angc r e s ca r ch
Ls t; Trans. Arm. SU:-:-.8er
Conf. C,:·f.P.S.,
The Hildl.
Soc.
13 pp.

--

1957,

decr,

Differences
in range veg e t a t fon r esu l t fng f r orn ~~ra;.ir.g by
c ar t l e , and sheep.
Pz oc . Soc. Arne r , For c s t er s , 19.:;/:1/)7-J.51.

)957.-(.
Some results of a lO-ye3r study of deer-livestock
compctition for range forage.
Proc. 10th Ann. Neeting Araer , Soc.. Range Hg::-:L
·14 pp.
(m iraco , )

-Riordan, L. E., and R. L. Canpb elL,
Outdoors 5(1):1-6.
Robinette,
Rocky

1956.

Harmony on the rang e ,

W. L., et al. 1957. Notes on tooth deve1op~ent and wear for
Hountain tau le deer. J. Hildl. !-fgr.1t. 21(2):134-153.

Rogers, G. E. 1948. Deer-elk-bear
investigations.
Frog. Rep •• April.
Colo. Dept. Caoe and Fish.

1948.
Deer-elk-bear
Rep., July.
Colo. Dept.
•_--.
Rep.,

.

investigations.
Came and Fish •

pp. 8-9.
54 pp ..

pp.

1948: Deer-clk-o.:!ar
in'lC'stigatior.s.
pp , 27--30 • .!E. QU&lt;1rt. Pr og ,
October.
Colo. Dept. Cane and Fish. 80 pp.
Deer-elk-bear
investir,&lt;ltions.
pp , 110-42.
Janu~ry.
Colo. Dept •. Cane and Fish.
67 pp.

,_~._, __• 1949.
Rep.,

I~ Quart.

pp. 24-29. In Quart. Pr og ,
83

19~9.
Rep.,

Colorado

April.

1949.
Rcp , , July.

Decr-c1i~·-bc&lt;lr invcs tif..1 t Jo ns .
Colo. Dept. Came &lt;lnd fish.
Dccr-elk~bcar
inv~sti~~tinns.
Co Lo , Dcp t , C,II:1C LInd Fbh.

In Quart.

Pr og ,

p p • ~1-47.
97 pp.

pp. 26-29.
8ft

pp.

~

Qua r t , Pr og ,

�1.951.

Chcck:i.n~ statton
survey.
Colo. Dept. Gaoe and Fish.

January.
______

..

1952. Gane-livetock

c

235$
.

32nd Ann.

1953.
11ildl.

--~j~.

Conf.

33 pp.

forage conpetition
study in Colorado.
P? 228West. Assoc. State Game and Fish c~.
252 pp.

Func t Lon and operation
HgL"lt.

?p. 32-33.

of [.ig £ame check stati.ons
-'

_,_,

:in Col.orcdo .

17 (3): 256-267.
0

1965.
Sex and a ge ratios
of deer harvested
under a hun t cr s+cho Lce ,
lQl'lt:tpJ.eo-liccnse hunting s ecs on , Colo. Dept. Game, fish and Parks.
Outdoor Facts No. 34. 2 pp.
4

l::O~crs,

G.,

O. Ju1ander,
and W. J..•• Robinette.
1958.
Pe l Let+g roup counts
and range-use
index. J. Wild1. Mg~t. 22(2):193-199.

for

deer census

Sed~lcy. E. f., and TI. M. Boeker.
Conscrv. 26(8):177-178.

1961.

If500 cattle

deer.

and 100,000

Soil

'Shepherd,
R. OR., et a1. 1966. G~e range investigations.
pp. 91-92. In
Game Res. Rep." July - Part 1. Colo. Dept. Garae , Fish and Parks.
pp. 1-93.
, .et a.l,

1966.

and iPa rks ,

-Fish

Ga:.1erange

surveys.
pp. 14-18.
Colo.
Review 1965.
48 pp.

Dept.

Game,

Ga~e Research

. .

_____ ,.~' 1967.
G;:
range surveys.
pp.26-31.
Parks.
Gat:,C ",...:searcn Rev c iw 1966.
46 pp.

Colo. Dept.

Game, Fish

and

______
196B. Gawe range surveys.
pp. 25-28.
Par~. Game Research Review 1967. 44 pp.

Colo.

Game, Fish

and

L:n:itn,

G.

,J.

l!!. Quart.

1958.
Deer-elk
Rcp , , October.

~,964.. Factors

2~3-D. M.S. ~hesis.

investigations.
Colo. Dept.

affecting
Colo.

Dept.

"pp , 139-140, 157-159,
Came and Fish.
166 pp.

the response of rubber rabbitbrush
State Univ., Ft. Collins.
87 pp.

4!'-:J.:::6~, Controlling
rubber r abb t tbrush '-lith 2,4-D.
-----:-Fish
ana i'ar;.;.s, Outdoor Fac zs No. 19;
2 pp.
Stephens,
L. N.
Colo. State

1971.
Vniv.,

S\&gt;ope, H. H., ct a1.
Fish and Parks.

___

• 1970.
Parks.

Browse use by raul,e deer
Ft. Collins.
37 pp.
0

and livestock.

surveys.
pp.
Review 1969.

to

Dept.

Gane ,

H. S. Thesis.

'

1969. Ga~e range surveys.
Ga~e Research Review 1968.

Game range
Game Research

Colo.

162.

Colo.

pp. 18-23.
36 pp.

27-29.
35 pp.

Colo.

Div.

Div. Game,

Game, Fish

and
•
0

J. ~~. 1962.
A r csume of Colorado big game research
projects,
pp. 1.6-65. In Tech. Publ. No.9.
Colo. Dept. Game and
l-'ish.
81 pp •

'fHeston,

·1939-1957.
••

~o

••

.. .~.

•

"

...•.

�-9~-

Ut. 5e Dept.

of Agr.
1965.
\l.:lter and related
land resources.
Color~do River
in Colorado.
Coop. Study Rep. of Colo. ~atcr Con~erv. Board and
U.S.1&gt;.A. Econ. Res. Sc rv I c e , Forest Service,
and Soil Conserve Se rv Lcc ,
Basin

..

MaYA

183 pp.

Williams 5 J. E.
Rep_, July.

--

•. 19.56.

.

c.

(looseleaf)

Deer-elk investigations.
pp. 68-71.
Colo. Dept. Ga~e and Fish. 102 pp.

1951.

Deer-elk investigations.

TC::lluary. Colo.

l1J11i.&lt;1lUS, .J., et a1.
.In Quart. Prog.

Dept.

pp. l~lO.
119 pp ,

G.1.1!e
and Fish.

In Quart.

•

IE. Quart.
Prog.

Rep.,

.,

1952.
Loca t Jon and extent of -..;inter ranges.
pr.57-62
Rep. ~ April.
Colo. Dept. Gane and Fish.
62 pp •

•

•

•

-.

•

•

•

.',
'

...

..

Pr og ,

.

..

•

•

�-96-

SMALL GAME HAl-t·tAL RESEARCH REFERENCES

WILDLIFE MAl~AGEHENT UNIT 11

•

COTTONTAIL RABBIT
De.nncy , R. N. t and D. L. Gilbert.

ppe 1-31.

1952. Fur resources and bear studies.
In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 60 pp.

Sandfort, W. \-1. 1951. Game hird surveys. pp.45-47.
July. Colo. Dept. Game and Fish. 102 pp.

In Quart. Pr og , Rep , ,

Shepherd, H. R. 19653. Colorado long range game species management plans,
1965-197.)t for cottontail raLbits. Colo. Dept. Galle, Fish and Parks.
12 pp.
U. S. Dept. of Agr. 1965. Water and related land resources, Colorado River
Basin of Colorado. Coop. Study Rep. of Colo. Water Conserve Board and
U.S.D.A. Econ. Res. Service, Forest Service, and Soil Conserve Service.
May. Denver. 183 pp. (looseleaf)

SNOWSHOE HArtE
Dolbeer, R. A. 1972. Population dynamics of the snowshoe hare in Colorado.
Ph.D. Thesis. Colo. State Univ., Ft. Collins. 193 pp.

--:

Porter, K. 1959. Effects of subalpine timber cutting on wildlife :LnColorado.
" In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. pp. 151-185.
Shepherd, H. R. 1965b. Colorado long range game species manage8cnt plans,
1965-1975, for snowshoe hare. Colo. Dept. Game, Fish and P~rks. 11 pp.

U. S. Dept. of Agr.

1965. Water and related land resources, Colorado River
Basin, Colorado. COC'lp.Study Rep. of Colo. t~ater Conserve Board and
.U.S.D.A. Econ. Res. Serv-ice, Forest Service, and Soil Conserv.. Service.
}Iay·. Denve r , 183 PP ° (looseleaf)
."

0-

•

0

..
•

•
•
.

~....

�-97-

...

CAME BIRD RESEARCH REFERENCES
trrLDLIFE lW:AG81ENT UNIT 11

•

BAUD-TAILED PIGEON
Braun, C. E.

---~

Colorado Outdoors 19(5):26-29.

1970. Ki.gratory bird investigations,
Rep.,

__

1970. The band-tailed pieeon.

October.

pp. 151-171. In Game Res.
Colo. Di.v. Gat.le,Fish and P&lt;i:cks. 171 pp.

1971. Band-taiJ.ed pigeon investigations. pp. 209-236. In Game
Res. Rep., Oc tober. Colo. Di v , Game , Fish.and Parks. 236 pp.
c

1972. Dap.d-tailed pigeon investigations.
Res. Rep., October. Colo. Div. of Wil-life.

pp. 123-141.
168 pp.

In Game

Braun, C. E., and D. E. Benson. 1972. Band-tailed pigeon investigationsbreeding and nesting chronology studies. pp. 159-168. In Game Res.
Rep., October. Colo. Div. of Wi~dlife. 168 pp.
Grieb, J. R. 1965. Colorado long range gane species management plans,
1965-1975, for migratory birds. pp. 33-36. Colo. Dept. Game, Fish
and Parks. 36 pp.
Neff, J. A·s and J. C. Culbreath. 1946. Colorado band-tailed pigeon.
pp. 1-24. Fed. Aid Proj. 4-R. Colo. Dept. Game and Fish. 24 pp •

. U. S. D~pt. of Agr.

1965. Water and related land resources, Colorado River
Basin, Colorado. Coop. Study of Colo. Water Conserv. Board and U.S.D.A.
Rcon. Res. Service, Forest Service, and Soil Conserve Service. }~y.
Denver. 183 pp. (looseleaf)

White, J. A. 1972. Plumage studies of band-tailed pigeons. pp. 143-158 •
.!!!. Game Res. Rep., October. Colo. Div. of Wildlife. 168 pp.
BLt.;EGROUSE
Aldrich, J. W., and A. J. Duval. 1955. Distribution of American gallinaceous
game birds. U. S. Fish and Hildl. Circ. 34. 30 pp.
Anderson, A. E. 1959. Deer-elk investigations. pp. 118-128.
Rep., January. Colo. Dept. Game and Fish. 128 pp.

In Quart.

-----. 1960. Effects of sagebrush eradication by chemical means on deer
and related wildlife. Job CompI. Rep , , July. Colo. Dept. Game and
Fish. 72 pp.
I:
-----.. 1960. Distribution and abundance indices of selected biOla in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Univ., Fort Collins. 130 pp.

�-98___

• _19(;6.
An Lnv c s t f rta t Lon of 2t~-D npp l Icn t Lon to !i~r.cbrush v Lt h In a
mount aI n shrub-con if c r' comp l cx , Unc ompnhgr c U.lt Lonn I Fo r cs t.
In Carne
}?Cl&gt;. Rcp , , July - I'a r t; HI.
Colo.
Dept. Game, Fish and Parks'--IJP'
345-

~20.
• 1969. 2.4-D, sagebrush and mule deer-cattle
use in upper ~intcr
..._--_.--range.
Spec. Rcp. lio. 21.
July.
Colo. J?iv. Game, Fish and Parks.

21 pp.
Hyers»

c. T.

1965. Eva l ua t fon of a corab in ed b l uc g r ou s e+o lLd turkey hunt ing .
'Out doo r Facts No. 27.
Colo.' Dcp t , Carne , Fish and Parks.
1 p.

eeason ,

l-:.oger.sJ C. E.

1962.

Part 1.

Colo.

Canc bird survey.
Dept. Ga.':1C and Fish.

pp. 121-J.31.
pp. 1--160.

In

Quar t , Rcp , , July

-

1963.

Game bird survey.
pp. 157-168, 191.
In CaI:1e Res. Rep . ,
- Part 110 Colo. Dept. Cane, Fish. and Parks.
pp. 109-241.

.-.,-~. -October
",

~. 1963. Bluc'grouse
census and harvest
---Canada.
J. i-li1dl. ~·rg::lt. 27(4);579-585.

in the

United

States

and

• 1965. Coloraco long range g2we species uanagenent
plans,
1965---1975,
for blue grouse.
pp. 1-3.
Colo. Dept. Ga~e, Fish and Parks.

16 pp.
1968. The blue grouse in Co10!·a~o.
Par'ks , Tech. Pub l., ~;o. 21. 63 pp.

..

'Sandfort,
ll.
"October.
sEd.nhoff,

1950.

W.

Colo.

H. H.

Game bird surveys.
Dept. Came and Fish.

1956.

The dusky
Syracuse
univ.

Ph.D. Thesis.

Colo.

Dept.

p. 37.
44 pp.

Fed.

grouse and its
173 pp.

Came , .Fd s h and

Aid Proj.

W-37·-R,

'- in Colorado.

ecology

• 1958. A rating
scale
for blue grouse ih Colorado.
---I~er.
For.,
Wildlife
So~.
pp. 133-138.

Froc.

Soc.

u. S. Dept.

of Ag r ,
1965. ~]ater and related
land resources.
Colorado
. River Easin in Colorado.
Coop. St udy Rep. of Colo. ~'7ater Conserve
Board and U.S.D.~.
Econ. Res. Service,
Forest
Service,
and Soil Conserve
Service.
May. Denver.
183 pp.
(looseleaf)

-

CnUl~R

Aldrich,
J. U., and A. ,1. Duvall.
1955. Distribution
game birds.
U.S.D.!. Fish and \·!ildl. Ser., Circ.

nartnann, R. M.

1963.

Rep ,', October
1
---..--I'ar• t 196
II.

l.

Canc bird surveys.
- Part 1.
Colo. Dept.

Carne bird
Colo.

Dept.

pp.

of Aracr i c an gc l Li nac cous
xe , 34.
23 pp.

33-40, 45-69.

Tn Ga~e Res.

Cane, Fish and Parks .-pp. 1-108.

5urvey.
pp. 151-178.
Game, Fi.:;h and Pa r ks ,

Tn Carne Res. Rep.,
pp.

•

71-2':33.

April

-

�-92Evansf

R. L. , and W. '-1. Sand Io r t ,

Quart. Rep. , October.

1957. Came bird survey. pp. 55-67.
Colo. Dept. Came and Fish. 116 pp.

Ilurd, C. L. t and W. w. Sandfort.
Quart. Prog, Rcp ,, October.
e

'
and
Rep.) April.

In

1955. Came bird survey. pp. 85-88.
Colo. Dept. Came and Fish. 10~ pp.

1956. Carne bird survey.
Colo. Dept. Came and Fish.

pp. 75-81. ~
~Ol pp.

In

-

Quart. Prog.

lkrming, D. D. 1962. Came bird survey. pp. 9-16, 25-31. In Quart. Rep.,
J.uly - Part I.
Colo. Dept. Cazae and Fish. pp. 1-160.
lH.ller, J. 1959., Carae bird survey. pp. 99-107,115-124.
October. Colo. Dept. Game and Fish. 124 pp.

In Quart. Rep.,

Miller, J., and t-1. W. Sandfort.
1959. Gane bird survey. pp. 67-73.
~Jart. Rep., January. Colo. Dept. G@ae and Fish •. 128 pp.
Sandf'o r t , \07. 1I. 1950~ Game bird survey. pp , 51-52.
April.. Colo. Dept. Came and Fish. 101 pp.

In

In Quart. Prog. Rep. s

1950. Ga~e bird survey. pp. 37-39. Fed. Aid Proj. W-37-R, October.
Colo. Dept. Ga~e and Fish. 44 pp.
1951.
Uildl.

Chukar census. pp. 22-31. Spec. Rep., Glenwood Springs
Conf., January 16-19.
Colo. Dept •.Game and Fish. 85 pp.

• 1951. Game bird survey. pp • 37-41.
, Colo. Dept. Ga&lt;::eand Fish. 102 pp.

E Quart. Prog. Rep. , July ..
-:;::.

1951. Game bird survey. pp. 80-82.
"
Colo. Dept. Gawe and Fish. 117 pp.

In Quart. Prog. Rep. , October.

Ga!!lebird survey. pp. 40-56.
" 1952.
Colo. Dept. Gace and :Fish. 62 pp.

In Quart. Prog. Rep. , April.

• 1952. Game bird survey. pp • 93-95.
Colo. Dept. Cane and Fish. 105 pp.

In Quart. Prog. Rep , , October.

• 1953. Game bird surVEy. pp . 81-;-89.In Quart. Prog. Rep. , April.
Colo. Dept. Gane and Fish. 99 pp.

1953. Game bird survey. p. 107.
.Colo. Dept. Gane and Fish. 129 pp.

~

1954. Game bird survey. pp. 13-33.
Colo. Dept. Gaoe and Fish. 79 pp.
___

Quart. Prog. Rep., July •

In Quart. Prog. Rep., April.

a
1954.
Game bird survey. pp. 67-72• .!2l Quart. Prog , Rep , , July.
Colo. Dept. Game and Fish. 182 pp.

1955. G.1mc bird survey. pp. U-18 • .!2l QU.1rt. Proc. Rep., January.
Colorado Dept. Came and fish. 94 t' p ,
G:.me b Lrd nurvcy , pp. 82-M • .!.!! QU:lrt. Prog , Rcp , , July _
---'Par t 1955.
1.. Colo. Dcpt ,
and Ffsh , 1'1'. 1-115.
C:UIlC

�-100.: 1957.
Came b i rd
pp , 29-38, 1,5-51. In Quart. Prog , Rcp , ,
---July
- P~rt I. Colo. Dept. Came and Fish. pp. 1-119.
SU1-VC)·.

1958.
C3me bird su~cy.
Colo~ Dept. Game and Fish.

______ •

•

1959. GaClC bird survey.
pp , 53--65•
Dept. Gacc and Fish. 128 pp.

~-Colo.

--~

• 1959. G~e bird survey.
Colo. Dept. Ga~e and Fish.
19GO.

c

"~~j)~epto

Ga:lE:

Came arid

bird
Fish.

Cane bird

• .1.960.

-Dept.

Game and. Fish.

survey.
74 pp.

I~ Quart. Rep., July

1962.

for

II.

•

.!E.. Quart. Rcp ,, January.

pp. 1:9--5/1 •

91-94.

.!E.. Quart. Rep , , •.
TanU31.-Y.
In Quart.

\.]ith the chukar ,

Colorado Outdoors

G~~e bird survey.
pp. 17-23, 33-44.
Colo. Dept. Gc~e and Fish. pp. 1-160.

Colo.

Colo.

Rep , , April.

9(5):26-29.

" 1961.
Carae bird surveys.
pp. 119-126. 139-155.
I~ quart.
April - Part lIe Colo. Dept. Carae and Fish. pp. 97-208.
Part 1.

Part

pp. 33-36. ' In Quart. Rep., April - Part I.
pp. 1-116.

survey .. pp.
161 pp.

Your chances

1960.

___

pp. 129-13~.
105-181~

pp.

Rep , ,

In Quart. Rep ,, July

-

1965. Colorado 1cng range g~e species nanage~ent plans, 1965-1975,
chukar pa r t.r Ldg c , Colo. Dept. Came , Fish and Parks. 18 pp .

---• 1967. A decade of chukar hunting. Coloraco Outdoors 16(6):20-23.
Sandfort, l~. l-l., and D. Nolting.
Quart. Prog. R~p., October.
Sandfort,
July.

1952. Game bird survey. pp. 89-91.
Colo. Dept. 9affie and Fish. 105 pp.

\~. t-l., et ala 1962. Game bird surveys.
Colo. Dept. Ga3e and Fish. 159 pp.

pp. 9-44.

In Quart. Rep ,,

1963.
bird
pp. 33-44, 63-70. In
-~-.
October.
Colo. Dept. Gaffie, Fish and Parks.
pp.

Quart.

Tull~, R.

14(6):16.

G2~e

surveys.

In

~ep.,

107

1965.

Gawe birds for 1965.

Colorado Outdoors

"u•.S. Dept.

of ·Agr.
1966. !.;rater and related
land resources)
i~l1ite River
Basin in Co Lor c do , Coop. Study Rep. of Colo. ~,1ater Conserve Board and
V.S.D.A.
Econ . Res. Service,
For c s t Service
and Soil Cons e rv , Service.
}~ovc;nber.
Denver.
92 pp. (looseleaf)

and J. E. llillians.
1960.
Escalante.
pp. 21-23. Color.:ldo
Outdoors, No.5, Sept.-Oct •. Colo. Dept. Game and Fish. 32 pp •

\1hitc, C.,

•

.' .

.. .

�..

-101-

CAW~EL' S OITML

l==. QU0.rt.- Prog , Rcp ,,

w. W.

Snndfort,

July.

1950. Game bird surveys.
pp. 17-21.
Colo ..Dept. G&lt;lme &lt;lnd Fish. 80 pp.

• 1950.

Game

bird survey.

-----COlo. Dept. G0.QC and Fish.

pp. 37-39. Fed. Aid P~~j. W-37-R. October.
44 pp.

• 1951.0 Came bird survey. pp , 8082. ~ Quart. P~~g. Rep., Octobcr.
---Colo.
Dept. GP.me and Fish. 117 pp.
'

..

o

'··"'~Col0.
___

-

1952. Game b Lrd survey.
Dept. GfiOe aud Fish.

pp , 93--95.
105 pp.

}-n Quart.

" 1953. Came bird survey.
pp.8l-89.
Colo. Dept. Gace and Fish. 99 pp.
•
for

~'::og.

Rep. J October.

In Quart. ?:::-og. Rcp , , April.

1965.
Colorado Ions range g ane species manage:::e:::-::plans, 1965--1975.
Gat.bcl's
quail. Colo. Dept. Cane ;: Fish and Par ks, 8 pp ,

Sandfo r t , W. W. t and D. !iolting.
Quart. Prog. Rep., October.

1952. Came bird su rvevs , pp . 89-91.
Colo. Dept. Ga~e and r~~~. 105 pp.

In

Sandfort. W. Y., and H. ~. S~ope.
1954. Game bird surve+ , pp. 19-29.
-Quart. Prog. Rep.s 3anuary.
Colo. Dept. Ga:::.e
and F~~_':. ·113 pp.

In

•
150URNH,G DOVE

c

_

Braun, C. E. 1971. Trapping and band i.ngdov es , pp . 201-'::07. IT!. Quart.
'..
Res. Rcp ,, October.
Colo. Div. Came , Fish and Par ks , 236 pp.
1972. Hourning dove trapping and banding. pp , ::5-121.
Quart. Res. Rep.; October.
Colo. Div. of ·Hild1ife. ':~8 pp.
erieb, J •.R.

1964.

Dove hunting deluxe.

In

Colorado Outc~.:"~-$
13(4):32-39.

1965. Colorado long range gane species nanage~e~~ ?1ans, 1965-1975,
for migratory birds.
pp. 33-36. Colo. Dept. Ga~e, :~$~ and Parks.
36 pp.
Nolting, D., et a1.
Rep., October.

1952. Cane bird survey. pp. 99-101.
Colo. Dept. Ca~e and Fish. 105 pp.

Sandfort, W ..W. 1950. Cane bird survey.
October. Colo. Dept. Cane and Fish.

pp. 41-42.

•

1951. Cnmc bird survey. pp. 83-8~.
Dep~. Canc and Fish. 117 pp.

195". G~mc bird survey. pp. 23-24.
Colo. Dept. C~mc and Fi::;h.111 pp.

Fec: .. Aid Proj. H-37-R,

In.' Quart.

~

Qua r t ,

Tn Ouart.
•

.~

Quart. Prog .

~4 pp.

1951. Game bird survey. pp. ~2-~3.
Colo. Dept. Ca~e and Fish. 102 pp.

---~olo.

~

Rep., July.
;&gt;"',.~..,.,
-. - .•..
'0·

Rep., October.

Rep., October.

-,

�-102- .

______ " .1955. Citmc bird survey. pp , 91-92. In Quart.
Fnrt I. Colo ••Dept. Came and Fish. 115 pp.

Prog. Rcp ,, July -

Snndfort,
H. W., and D. Nolting.
1952. Came bird survey. pp. 103-105.
~ Quart. Prog. Rep., October. Colo. Dept. Game and Fish. 105 pp.

v. Se Dept. of Agr.

1965. Water and related land' resources. Colorado River
Basin 1n Colorado. Coop. Study Rep. of Colo. \-later Conserve Board and
. U.S.D.A. Econ. Res. Service, Forest Service and Soil Conserve Service.
May. Denver. 183 pp. (looseleaf)
,

Rn~G-NECKED ?l!~AS.~:T

.

JJdrich, J. W., and A. J. Duvall. 1955. Distribution of ~erican
gam~ birds. U.S.D.I.~ Fish and Wildl. Circ. Ko. 34. 23 pp.
Figge, H. 1952. Gane bird survey. pp. 45-60.
Colo. Dept. G~e and Fish. 60 pp.

gallinaceous

In Quart. Prog. Rep., January.
--..
'

Figge, H., et a1. 1953. G~e bird survey. pp. 75-86.
January. Colo. Dept. Game and Fish. 141 pp.

In Quart. Prog. Rep.,

Hoffman, D. M. 1971. Pheasant nest site selection study.' pp. 1-26. In
Quart. Res. Rep ,, April. Colo. Div. Cane , Fish and Parks • 125 pp.
_.

.

1972. Pheasant nest site selection st'l!dy. pp. 1..:19. In Quart.
Res. Rep ,, April. Colo. Div. Came , Fish and Parks. 150 pp.

Sandfort,

1950.
Game bird survey, pp , 17-21.
Colo. Dept. Ga~e and Fish. 80 pp.

W. W.

..July.

In Quart.

Prog.

Rep.,

• 1950. Carre bird survey. pp'.1-12.
Colo. Dept. Ga~e and Fish. 44 pp.

Fed. Aid Proj. 1-l-37-R, October.

1951. Game bird sur/eve pp. 1-7.
Colo. Dept. Ga~e and Fish. 35 pp.

Fed. Aid Proj. W-37-R, January.

I

______ • 1951." Ga~e bird survey. pp. 9-20.
Colo. ~ept. G~~e and Fish. 102 pp.

In Quart. Prog. Rep., July.

1952. G~~e bird survey. pp. 115-119.
Colo. Dcpt~ Game and Fish. 171 pp •.
1952. Game bird investigations.
October. Colo. Dept. Ga~e and Fish.
.,....
__ • 1955.· Game bird survey. pp. 1-9.
Colo. Dept. Gat:1Cand Fish. 94 pp.

In Quart. Prog. Rep., July.
-.-

pp , 37-41.
105 pp •

~

In-Quart. Prog. Rep., January.

1955. G~~c bird survey. pp. 49-63, 65-69. In
July - Pa r t 1. Co Lo Dcp t , Game and Fish. 115 pp •

----

1956.

.

Upland &amp;amc birds.

Quart. Prog. Rep.,

Quart. Prog. Rep.,

Colorado Outdoors 5(6):2.
•

-:'.

�.

-103-

-

• 1957.

Upl~nd &amp;~mc birds •

Colorado Outdoors '6(6):2.

-= .

1960. G.:l.OC
bird survey. pp. 37-55, 59-65.
Colo. Dept. Gaoe and fish. 161 pp.

______

___
. .

1961. Game bird survey. pp. 97-102.
Colo. Dept. Ga~e and fish. pp. 97-208.
e

l!!. Quart. Rep •• April..

•

In Quart. Rep., July - P~rt II.

• 1963. Came bird survey. pr. 213-223. In Came Res. Rep., October
Par t 110 Colo~ Dept. Ga~eJ Fish and Park~.
109-241.

:pp.

Sandfort,

. , 1£

w. ~l.s

and H. M. Svo pe , J.95!,.
Cane b Lrd survey, pp. 19..:.29 •
Quart. Pr og., Re p ,, January. Colo. Dcp t •.Carne and Fish. 113 pp.

Stie~~, H. A. 1950. Ca~e bird survey. pp. 1-4.
J~ly&lt;; Colo. Dept. Game and Fish. 80 pp.
Svope~ H. H.

1965.

In Quart; Prog. Rep.,
•

Colorado long range ga!:lespecies manage~ent plans)
Colo. Dept. G2!ile, Fish and Parks. 54 pp.

1965-1975, for pheasants.

U. S. Dept. of Agr.

1966. Water and related la~d resources) ~iliiteRiver
Basin in Colorado. Coop. Study Re? of Colo. Water Conserve Board and
U.S.D.A. Econ. Res. Service, ?orest Service, and Soil Conserve Service.
Hovember. Denver. 92 pp. (looseleaf)

Aldrich, J. i-T., and A. J.• Duvall. 1955. Distribution of A'-:lerican
gallinaceous
game birds.
U. S Fish and \-lildl.Circ. 34. 30 pp -.
co

Boeker, H. H. 1955. Game bird surveys. pp. 70-71.
,July. Colo. Dept. G~e and Fish. pp. 1-115.'

In Quart. Prog. Rep.,

Funk, H. D. 1971. Effects of sagebrush control on distrib~tion and abundance
of sage grouse. pp. 41-47. ~ Quart. Res. Rep., April. Colo. Div. Ga~e,
Fish and Parks. 125 pp.
1972.' Effects of sagebrush control en distribution and abundance of
sage grouse. pp. 3l-3i. .IE. Quart. Res. Rcp ,, April. Colo. Div. Carne ,
Fish and Parks. 149 pp.
Rogers, G. E. 1958.
July - Part II.

Ga~e bird surveys.' pp. 123~125. In Quart. Prog. Rep.,
Colo. Dept. Ga~e and Fish. pp. 105-181.

1959. Cane bird surveys. pp. 13-32.
Colo. Dept. Gaille and Fish. 116 pp.

--~.
•

1963.

Th~ sage grouse of Colorado •

In Quart. Prog. Rep., April.

Colorado Outdoors 12(3):22-24.

1963. Game bird survey •. pp , 13-32 •. Tn Came Res. Rcp ,, October Par t 1. Colo. Dept. Game, Fish and Parks. pp. 1-108 •

••

..

\

,

�-104-

"1964. Sa~e g rous c Invcs t Laa t Lons, 1n Colorado.
Fish und Park~. Tech. Publ. No. 16. 132 pp.

___

0

Colo. Dept." Game,

Colorado long range game species management p Lans , 1965-1975.
----:-for• '1965.
sage grouse.
pp. 4-7. Colo. Dept. Game, Fish and Parks. 18 pp •
. U. S. Dept. of Agr , 1965.
Water and related land resources, CoLo rad o River
Basin in Co Lo rado , Coop. Study Rep. of Colo. Water Conserve Bo ar d and
U.S.D.A.
Econ. Res. Service, For cs t Service, and Soil Con se rv , Serv Lcc ,
lhy. Denver. 183 ppo
(looseleaf)
",
",

tIA'fERFOt.-TL

Funk, H. D. 1971. Waterfm.rl kill survey. pp , 51-73. In Quart. Res. Rep. ,
"
October.
Colo. Div. Game, Fish and Parks. 236 pp.
Crieb~ J R. 1965. Colorado long range game species manageznent; plans, 19651975, for migratory birds.
pp. 1-32. Colo. Dept. Carne, Fish and Parks.
36 pp.
e

Hopper, R. M. 1963. Wetlands of Colorado.
Tech. Pub!. No. 22, }!arch. 89 pp •

Colo. Dept. G~e,

Fish and Parks.

•
IJ.urich, J. tol., and A. J. Duvall." 1955. Distribution of Anerican gallj:naceous
game birds. U.S.D.I., Fish and Wildi. Ser., Circ. No. 34. 23 pp.
Burget, M. L. 1946.
Fish. 21 pp •

Colorado wild turkey.

Vol. II.

Colo. Dept. Game and

..•

• 1.948. 'Hild turkey surveys and investigations.
pp. 47~62. In
Quart. Frog. Rcp , , October.
Colo. Dept. Game and Fish. 80 pp.
surveys and investigations.
pp. 91-106.
In
---••Quart.1949.Prog.WildRep.turkey
Colo. Dept. Gaue and Fish. 141 pp.

1949.
April.

Wild turkey develop~ent.
Colo. Dept. G~c an~ Fish.

pp. 3-18.

In Quart. Prog. Rep.,

9.8 pp.

1951. Wild turkey rehabilitation in Colorado. pp. 7-17. Spec.
Rep. Glen~ood Springs ~~ildl. Coni., Jan. 16-19. Colo. Dept. Game and
.Fish. 85 pp.
" 1957.
January.

The wild turkey in Colorado.

1960. Wild turkey development.
Colo. Dept. Game and: Fish. 110 pp.

--_.

1961.

October.

Colo. Dept. Came and Fish.

68 pp.
pp. 5-42.

In Quart. Rep., July.

Yild turkey deve~opmcnt.
pp. 23-43. 51-55.
Colo. Dept. Gane and FJsh. 103 pp.

-,

.!.!:!. QU.:lrt.Rcp , ,

�-10~1963.
Ylld turkey investic~tlons.
Colo. Dept. Came and fish.

. April.

Burcet, M. L.

t

and C. Ford.

1951.

Pr og , Rcp , , Ap r t l ,

Quarto

pp. 71-85.
109 pp.

Wild turkey

Colo.

Dept.

In Game Res.

Rep_,

pp. 1-7.

development.
Fish.

64 pp.

Gat;1C and

In

•.

.

.

Durget,
H. L., and D. M. Ho f fman,
1950. Uild turkey surveys and investi.gations.
Three Year Sura Re p , , 1947-1950. July.
Colo. Dept.
Carne and fish.

- 4 pp.

-r .
t
and
1950.
-"-~~-Ocl:ober-=--C-;;10. Dept.

----;::-

, aud

• l.!:. Quart.

_,

"
Prog.

1.951.
Rep.,

Hild t ur key investi{;ations.
Cane and Fish.
18 pp .

, and

1952~

H-39-Rt

m.ld turkey surveys and Lnve s t Lg a t f ons , pp • 11--29.
October.
Colo. Dept. Carae and Fish.
117 pp.

and
1952.
llild t.ur key investigations;
Pzog , Rep. ~ "-priL
Colo. Dept. Carie dud Fish.

,.

Fed. Aid Proj.

pp , 1-5.

In Quart.

62.pp.

'Hild turkey investigations.
pp. 1-9, 11-14, 21-24.
Colo. Dept. GCi:L:e
and Fish.
105 pp.

--.~ iE. Quart. Prog. Rep , , October.
----;::-

,and

Frog.
__

Rep.,

1953.
Wild turkey investigations.
April.
Colo. Dept. G~e and Fish.

--::-., and

1953.

.Prog •. Rep , , July.

__

pp. 19-33.

In Quart.

99 pp.

'l-7ild turkey investigations.
pp.
Game and Fish.
129 pp.

1-6.

~

Quart •

Co.Lo , lJept.

~~ ·nn:~.
-,·.:-·~.·,·;~·:::t~:5&lt;·:
lhJd. turkey investigations.
pp , 12-2l,.
Pzog.-·':Rep_ ~ J"uly .•·-&lt;...:.L:t.~J..o.~,.lJ.~pt.GaI!1C and Fish.
182 pp.

~

Quart.
_

, and
1955~. l-lild turkey investigations.
pp. 1-8, 13-19.
--·Quart.
Prog. Rep ",, Oc::::ober.
Colo. Dep
Gane and Fish.
108 pp.

In

t •.

.
. and
1957.~· 'Rild turkey investigations.
pp. 69-86.
--'
Rep , , October •. Colo. Dept. Game and Fish.
116 pp.

In Quart.

, and
1955.,11D.d
turkey inv~stig.:!tions.
pp. 147-169.
In
---Quart
.• 13ep., July - 'Part II.
Colo. Dept. Came and Fish.
pp. 105-181.

-l:vcins,R. ~1... ··1963~

l-lild c:m:-k;:::y investi£2..tions.
Colo. Dcp t; , G(.l.t1eand fish.

Rep , , :April.

Hofrean, D. X.
.1965-J.975,

1965.
Colorado
for v i.Ld turkey.

pp. 71-85.
109 pp.

G. T.
1963.
October - Part

Came bird
II.
Colo.

•

Res.

long range game species canagenent
plans,
Colo. Dcp t , Cx:1C, Fish and Po r ks , 11 pp.

Ilunt er , G. N.
1965.
Colorado big garnc harvest,
1959-1965.'
Rep. No.1.
Colo. Dept. Carne, Fish and P~rks.
41 pp •
lIyers,

In Cane

Spec.

H~mt.

survey.
pp , 173-178.
In Carne Res. Rcp , ,
Dcpt. G.1r.IC,Fish and Par ks , pp . 109-2/,1.
r.

..

�. -106-

Came bird survey. pp. 2~3-280. ~ Game Res. Rep., April Colo. Dept. Game, fish and Parks. pp. 235-397.

• '1964.
~art

III.

--_.

1965. Evaluation of a combincd blue grouse-wild turkey hunting
tlcason. Outdoor Facts No. 27. Colo. Dept. Gamc, Fish and Parks.
1 p.
•

--

1965. Reliability of turkey sex and age ratio data based on hunter
rcport card returns. Outdoor Facts No. 29. Colo. Dept. Ganc, Fish and
6

Parks.
c

2 pp.

1965.

I.
"-~~~"-Part
_____

e

pp. 91-17/,.
In Cane Res. Rcp , , April
Game, Fish and Parks. pp. 1-17/••

Carse b f.rd survey.

Colo ...Dept.

'1966. Cane bird survey.

Colo. Dept.

1967.
-

.:«.

April

Garae ,

pp. 69-84. In Gaoe Res. Rep., April.
Fish and Parks. 192 pp.-

..

Cane bird survey. pp. 167-182, 205-211.
Colo. Dept. Cace , Fish and Parks.

.4 Part II.

• 1968. ,Came bird survey. pp . 71, 99-110.
---,:-Colo.
Dept. Cane, Fish and Parks. 131 pp.,
• 1969. Came bird survey. pp. 107, 153.
Colo. Div. Cazie , Fish and Parks. 181 pp.
e&gt;

1969.

The topic is turkey.

Game bird survey. pp. 161, 165.
Colo. Div. Caine , Fish and Parks. 198 pp.
C. C. 1970.
.19 (3): 30-31 •

In Game Re s, Rep., April.

In Gaoe Res. Rep. , April.

Colorado Outdoors 18 (5): 30-34.

1970.

Sheats)

In Gar-e Res. Rep ••
pp. 107-2l11.

In Game Res. Re p , , April.

Hy experience \vit~ the turkey.

Colorado Outdoors

•
•
"

....
'

..

•

"

..

.

�•

..

:.

•

-107-

iWNGAME MA1-r-tAL RESEARCH

REFERENCES
o

WILDLIFE

MA.~AG21ENT

UNIT Jl

•

Boyd, R. J. 1965. Colorado long range game species management plans, 19651975, for predators. Colo. Dept. Came, Fish and Parks. 21 pp.
Remington, J. D •. ,1959. Furbearers of Colorado.
Colo. Dept. Gane and Fish. 16 pp.
Rutherford,

w. H.

1965.

Educational Pamphlet No.4.

Colorado long range game species management plans,
Coloo Dept. Game, Fish and Parks. 12 pp.

1965-1975, for furDearers.
COYOTE
Boyd, R. J.
1975,

1965. Colorado long range game species management plans, 1965for predators. Colo. Dep t-, Carae , Fish and Parks. 21 pp.

Craighead, F. C. 1951. A biological and economic evaluation of coyote
predation. New Yo rk Zoological Society and The Conservation Foundation.
23 pp.

..

Denney, R. N., and D. L. Gilbert. 1952. Fur resour~es and bear studies.
pp~ 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 60 pp .
Sperry, C. C. 1941. Food habits of the coyote.
Serv. Wildl. Res. Bull. 4. 70 pp ;:

U.S.D.I. Fish and Wildl.

1965. Colorado long range game species management plans, 19651975, for predators. Colo. Dept. Game, Fish and Parks. 21 pp.

.Boyd, R. J.

Denney, R. N., and D. L. Gilbert. 1952. Fur resources and ~ear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 60 pp.
Remington, J. D. 1959. Furbearers of Colorado.
Colo. Dept. Game and Fish. 16 pp.

Educational Pamphlet No.4.

Rutherford, t-l. H. 1965. Colorado long range game species management plans,
1965-1975, for furb~arers. Colo. Dept. Game, Fish and Parks. 12 pp •

.
'

RODENTS

..

Turner. G. T., R. H. Hansen, V. U. Reid, H. P. Tietjen, and A. L. Ward. 1973.
Pocket gophers and Colorado mountain rangeland. Colo. State Univ. Exper.
Sta. Bull. No. 55~S. 90 pp.

•
.S·' ~2'0

.. ::.!

�-108-

Remfng ton, J. D •. 1953. Fur resources and bco r s t ud Ics , pp , 17-23 •
.!!!. Quart. Pr og , Rcp., Oct.
Colo. Dcpt.' Game and Fl!ih.
60 pp.
_______ • 1959. Furbcarcrs of Colorado.
Colo. Dept. Game and Fish. 16 pp •

Educational Pamphlet No.4.

.Rutherford. U. H. 1965. Colorado long r~ngc game species management plans,
1965-1975, for f urbea rcrs , Colo. Dept. Came , Fish and Pa rks , 12 pp •
. -'.

troSKRAT

Denney) R. N. 195.0.
Fur resource survey. pp , 57-66.
• Jan. Colo. Dept. Ga~e and Fish. 66 pp.

In Quart. Frog. Rcp , ,

1950. Fur resource survey. pp. 93-l0l.
April. Colo. Dept. Game and Fish. 101 pp.

In Quart. Prog. Rep. ,

1950. Fur resource survey. pp. 75-80.
Colo. Dept. Cane and Fish. 80 pp.

In Quart. Prog , Rep. , July.

• 1951. Fur resource survey •
Dept. Game and Fish. 8 pp.

In Quart. Prog. Rep. , Jan.

Colo.

Denney, R. No, and D. L. Gilbert.
1952. Fur resources and bear studies.
Colo. Dept. Game and Fish. 60 pp.
np.
1-31. - In Quart. Prog. Rep., Jan.
¥'
0
. lteman5.c, 1-1. H. 1942.
.
t·:i..~h
COQD1issian.

Fur resource survey.
6 pp.

Vol. 2, Jan.

Remington, J. D. 1953. Fur resources and bear studies.
..
Quart. Prog. Rep ,, Oct. Colo. Dept. Cane 'and Fish.
_______ • 1959. Furbearers of Colorado.
Dept •.Game and Fish. 16 pp.

Colo. Game and

pp. 17-23.
60 pp.

Educational Pamphlet No. 4.

In

Colo.

Rutherford, w. H. 1965. Colorado long range game species ~anagenent plans,
1965-1975, for furbearers.
Colo. Dept. Ga~e, Fish and Parks. 12 pp.

".'EASEL

Denney, R. N. 1950. Fur resource survey. pp. 93-101.
.April. Colo. Dept. Ga~c and Fish. 101 pp.

July.

1950. Fur resource sur/eye pp. 75-80.
Colo. Dept. Game and Fish. 80 pp.

~

In Quart. Prog. Rep.,

Quart. Prog. Rep.,

Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bC3r studies.
pp. 1-31. ..!.!l Quart. Pr og . Rcp ,, Jan. Colo. Dcpt. Game and Fish. 60 pp.
Ncmanlc, W. M. 1942.
l'1sh Commission.

Fur resource survey.
6 pp.

Vol. 2, Jan.

Colo. Game and

�-,
-109:-

Rcmlncto~, J. D •..1953. Fur resources and bear -~tudl~s.
Quart. Prog. Rep., Oct. Colo. Dept. Came and fl~h •
•

• 1959.
~~lo.

Furbearers of Colorado.
Dept. Came and Fish. 16 pp.

pr. 17-23.

In

~O pp.

Educational Pamphlet No.4.

•

Rutherford. W. U. 1965. Colorado long range game species-m&lt;lnagcment plans,
lS65-1975, for furbearers.
Colo. Dept. C~~et Fish and Parks. 12 pp •

-.

•

..J.

..
•

•

••

.

•

'

..

•

.'

'0

•
•

0

•••

••

•

•

..

.- ' "'

,

�•

•

-110-

.- .

•

'

FURBEARER RESEARCH REFERENCES

•
WILDLIFE HANAGNEh'T UNIT 11

•

.!tRAVER

..Anonymous.
1937.
1(32):5, 7.

Beavers ~ flood control experts.

Natl. Nature News.

l?:tuley~ W. L.

1937.
The beaver··.·.conserver of soil and 'later. N. Amer.
Wildl. Conf.'Trans. 2:295-297.

•

Leighton, R. S., and J. A. Lee. 1952. A rpogress report on a technique to
control water levels in beaver impoundments. N. Ame r , 1-lild!.Con£.
Proc. 8:4 pp.
•

..

Rm.llngton, J. D. 1955. Fur resources and bear studies.
\-l-61-R. Colo. Dept. Cane and Fish. 85 pp.
• 1959. Furbearers of Colorado.
---.Dept.
Game and Fish. 16 pp.

Fed. Aid. Proj.

Educational Pamphlet No.4.

Colo.

Retzer, J. L. 1955. Physical environnental effects on beavers in the
Colorado Rockies. Western Assoc. Came and Fish Coma , ·.Proc.35.
Retzer, J. L., H. M. Swope, J. D. Remington, and W. H. Rutherford.
1956.
Beaver managecent in the Rocky Hountains of Colorado. Fed. Aid Proj.
W-83-R. Tech. Bull. ~o. 2. Colo. Dept. Cane and Fish. 33 pp.
Rutherford, W. H. 1953. Effects of a sumner flash flood upon a beaver
popUlation. J. HaUl. 34(2):261-262.

1964. The beaver in Colorado; its biology, ecology, nanagenent
'and economics. Colo. Came, Fish and Parks Tech. Pub. No. 17. 49 pp.

---:• 1965.

Colora~o long range game species canagement plans, 19651975, for furbearers. Colo. Dept. Game, Fish and Parks. 12 pp.

Scheffer, P. }1. 1938.

The beav~r as an upstream engineer.

Soil Cons.

3(7) :178-.181.

HINK
Denney, R. N: 1950. Fur resource surveys, pp. 75-80.
Rep., July. Colo. Dept. Game and Fish. 80 pp.

~

Nemanic, W. M·. 1942.
.Flsh Commission.

Jan.

Fur resource surveys.
6.pp •

Vol. 2.

•

..

~.....•...,

Quart. Prog.

Colo. Game and

�•

•

•
CENERfJ.

MA!-fY::\LIA.'t

REFERENCES

•
•
Ant.flOl1Y, H. E. 1928.
Son s , N'e'Y! York.

Field book of North
674 pp.

1.:n/!8tron~;:D. l-L 1972.
,
PrLn t f ug Servt_ce,

D:Lstrlhut:t0:'j, of :;;:\I;:;;TJ;.;,ls in. Co Lo r adc ,
Lavr en ce , 415 pp.

Bur t , W. H., and R. P. Grossenheider.
Houghton }liff1in
Co., Boston.

American
./

196(1.
284 pp.

mammals.

f. f ield

guide

G. P. Putn am+s

Un Iv , Kan s as

to the rcanraa.Ls ,

•

Cringan,
A. T.
1973.
Warn blood vertebrate
inventory
an a Ly s f.s and :tr~?act
s t udy of the Parachute
Creek area, Garfield
County, Colorado.
In TIH~
Colony cnv Lroricen tal s t.udy-e-Pa rachut e Cr eek , Garfield
County, Colorado.
Pxcpa re d for Colony Dcve Lo pnen r Operation by Thorne Ecological
Institute,
Boulder,
Colorado.
August, 1973.
Part II.
Vol. 2, Cahp t , VII.
pp. 1-137.
Balli

E. R., and K. R. Kelson.
1959.
The mamrna
Ls of North
Press Co., New York.
2 Vols.
1162 pp.

Lechleitner,

Boulder,

R. R. 1969.
254 pp.

Wild

mamua.Ls

of Colorado.

Pruett

Arae rd ca ,

Pub.

Co.,

Hiller,

G, S., and R. Kellogg.
1955.
List of North American recent
Smdt.hs cnd.an In s t , , t{ashington,
D. C. BulL No. 205.
954 pp.

Hurie,

O. J.
Boston.

Palmer,

1954.
A field
374 pp.

R. s. 1954.

City, N. Y.
Warren,

gu Lde to animal

R. R.

The manma L guide.
384 pp.

Doubleday

1942. The mamma l,s of Colorado.

330 pp~

..

..

'--

tracks.

mammal.s .

Houghton }lifflb

and Co.,

Univ. Okla.

Ronald

Co.,

Inc. , Garden

Press,

Norman ,

�-112-

GENERAL AVIAN REFERENCES
WILDLIFE }Lt,NAGEHENT UNIT n..

Bailey, A. M., and R. J. Niedrach. 1965. Birds of Colorado.
Mus. of Nat. Hist •• Denver, 2 Vols. 895 pp.
1967. Pictorial checklist of Colorado birds.
of Nat. Hi~t., Denver. 168 pp.
Davis, W. A.
61 pp.

1969.

Birds of western Colorado.

Denver

Denver Hus.

Colo. Field Ornithologists.

Kortright, F. A. 191.3. The ducks, geese,·and swans of North America.
The Amer. l-lildl.Institute, loJashington,C. D. 476 pp.
Linduska, J. P. 1964. Waterfowl tomorrow. U.S.D.I. Bur. Sport Fish. and
Wildl., Washington, D. C. 770 ~p.
Martin; G. S., P. H. Baldwin, and E. B. Reed. 1974. Recent records of
birds from the Yampa Valley, Northwestern Colorado. Condor. 76(1):
113-116.
Pearson, T. G. 1940.
N. Y. 289 pp.

Birds of America.

Garden City Publ. Co., Inc.,

Peterson, R. T. lS41. A field guide to western birds.
Co., Boston. 240 pp.
Robbins, C. D.,B. Brunn, and H. S. Zim.
Golden Press, N. Y. 340 pp.

1966.

Houghton Miffllll

Birds of North ,America.

Smith, A. G. 1973. Avian environmental inventory and impact study for
Colony Development Operation in Garfield County, Colorado. Part I.
Environmental inventory by Thorne Ecological Institute for Colony
Development Operation, Atlantic-Richfield Company, Operator, October,
1973.
Sprunt, A., IV., and H. S. Zim. 1961. Gamebirds--a guide to North American
spec~es and their habits. Golden Press, N. Y. 160 pp.
\.zooding,J. 1973. A census of the breeding birds of the Roaring Fork
watershed. Colorado Field Ornithologist. 9 pp.

�-113-

APPENDIX B

WILDLIFE 11A1~AGENENTUNIT 12 (WILLIAMS FORK)
Garfield,

Moffat,

Rio Blanco, and Routt Counties,

Colorado

Information on Unit Description,
Landovner-shi.p,
Land Use, HumanPopul.at i.on; ~Jildlife Species
Checklists,
Harvests, Seasons, Narrative
and Hap Descriptions of Distribution
and
Abundance; and other Allied Data.
To: December 1975.
Comp~~edby: W.T.

McKea.'1
and B.D. Trindle

�-114TABLE OF CONTENTS
ITEM

DESCRIPTION
LANDOWNERSHIP

(Boundaries,

size, physical

features,

climate)

.

STATUS ----------------

LAND USE AND VEGETATIVE
Hu}~N POPULATION

_

COVER STATUS --

_

---------------

_
~

LAND USE STATUS RELATING TO ELK SUMMER AND WINTER RANGE, WHITE
RIVER ELK HERD --------------GAME SPECIES

(Big game mammals,

small game mammals

OTHER MAMMALIAN SPECIES (Furbearers,
game mammals) ---------------OTHER AVIAN SPECIES

("Varmint"

"varmint"

and game birds)

mammals,

and non-

birds, non-game

birds and rap tors)

DISTRIBUTION AND ABUNDANCE OF BIG GA}ffi~t~~~S
mule deer, mountain lion) ---------

(Black bear,· elk,

DISTRIBUTION AND ABUNDANCE OF SMLL GANE ~IHALS
snowshoe hare, red (pine) squirrel) --DISTRIBUTION AND ABUNDANCE OF S~~LL Gt~lli BIRDS
pigeon, mourning dove, waterfowl, turkey)
DEER HARVEST,
PRIMITIVE
ARCHERY

SEASONS AND HUNTING

DEER HARVEST

LINFAR REGRESSION
ELK HARVEST,
ARCHERY

AND SEASONS

DEER HARVEST AND SEASONS

TOTAL DEER HARVEST,

ELK HARVEST,

DATA ANALYSIS

(1956-1974)

_
_
_

BY SEX

_
_

(1956-1974)

_

PRESSURE

(1967-1974)

_

BY SEX

_

UNIT 6, 1963-1972,
(1956-1974)

_

LION HARVEST AND SEASONS ----

_

SMALL GAME HARVEST AND HUNTING PRESSURE,
Management Unit 1Q) ----------------

1968-1973

S~~LL GAME HARVEST AND 'HUNTING PRESSURE
Unit 26, 1974) ----------------

(Small Game ~~nagement

i

rabbit,

_

UNIT 7, 1964-1964,

SEASONS AND HUNTING

_

(Grouse, band-tailed

(1973-1974)

PRESSURE

_

_

OF EITHER SEX DEER SEASONS -

BLACK BEAR Hl\RVEST AND SEASONS
MOUNTAIN

(Cottontail

(1956-1974)

DATA ANALYSIS

SEASONS AND HUNTING

TOTAL ELK HARVEST

PRESSURE

_

(Small Game
_

_

�-115TABLE OF CONTE~fS

(Continued)

ITEM
DUCK AND GOOSE HARVEST AND HUNTING PRESSURE (Small Game Management
Unit 26, 1974) ------------------------------------------------S~~LL GAME MANAGEMENT

UNITS MAP, 1968-1973

S~~L

UNITS MAP, 1974 -------------------------------

GAME MANAGEMENT

--------------------------

BIG GAME POPULATION

DYNAMICS ---------------------------------------_

HABITAT RESTORATION

PROJECTS

GAME SPECIES
MANAGEME~~

ItTRODUCTIONS

PROBLEMS

BIG GAME ~~~L

-------------------------------------

CHECKLIST

RESEARCH

NONGAME ~~

RESEARCH

---------------------------------

REFERENCES

REFERENCES

-------------------------------

----------.-----------------------.----

REFERENCES

__

---------------------------------_

FURBE.ARER RESEfu~CH REFERENCES

-----------------------_---------------

G'ENERAL t-Wfr~LIAN REFERENCES

--------------------------------------_

GENERAL AVIAN REFERENCES

_

---------------------------------------

REFERENCES

SMALL Gru~E MAMMAL RESEARCH
GAME BIRD RESEARCH

----------------------------------------

----------------------------------------

ii

_

�-116-

DESCRIPTION

- WILDLIFE HANAGEHENT

d .
2/
"
B
oun ar a.e s - .-- Those portions
counties

bounded

UNIT 12 (WILLIAHS FORK)

of Hoffat, Routt, Rio Blanco and Garfield

on the north by Colorado Highway

Road 29 from its junction

1.../

with Colorado Highway

317 and Routt County

317 to the Williams

Fork-

Yampa River divide, on the east by the Williams

Fork River-Yampa

divide,

on the south by the Yellow Jacket-Sleepy

Cat divide including

Williams

Fork drainage

Size.--491.l

Physical

and on the west by Colorado

square miles

(planimetered

Features.--Wildlife

Management

province.

is a southeastward

It also constitutes

Rocky Hountains

physiographic

from 1/2" BLH, 1974 base map).

Unit 12 is drained by the Williams

extension

of Craig.

of the Wyoming Basin physiographic

part of the northern
province

the

Highway 13."

Fork River which drains into the Yampa River a few miles southwest
This drainage

River

boundary

of the Southern

of which the White River Plateau is

a part.

In the west portion of Unit 12 is a prominent
U.S.D.A.

1969,

soft rocks and forming a sharply out-

•••Along the south side of the Axial Basin is a low range of

hills, the Danforth
Northeast

feature, quoting

••• " ••• the Axial Basin, formed by an uplift whose axis has

been deeply eroded exposing underlying
lined trough.

topographic

Hills, lying about 2,000 feet above the adjacent valleys.

of the Axial Basin, the Williams

back ridge formed by resistant

sandstone

Fork Mountains

layers."

are a high hog-

Elevations

within Unit 12

1/
- Legally termed "Big Game Hanagement Unit" (see footnote ~./ below) but
tentatively called "Wildlife Hanagement Unit" here to include broader aspects
of animal life present ;:

2/
- Colorado Div. Wildlife
Big Game).

Laws and Regulations

Hdbk., 1975 (p. 8, Chap. 2 __

�-117-

range from 6,200 feet on Milk Creek in Axial Basin upward to 12,110 feet
at Mount Orno on the, southeast boundary of the unit.

By far the most abundarrt rocks within Unit 12 are of the Mesa Verde Formation

(Mesozoic).

They occur throughout

sist mainly of sandstone with interbedded
These rocks are usually resistant
ridges, and cliffs

almost the entire unit, and conshale and several coal beds.

to erosion and often form prominent

(U.S.D.A. 1969).

Small intrusions

of Tertiary volcanics,

consisting

of basaltic

several hundred

feet deep, can be found along the south border of the unit.

They are rocks of the north flank of the White River uplift
Immediately
Formation

adjoining

them are small rock exposures

of late Tertiary

age.

tuffaceous

greenish

(U.S.D.A. 1969).

Among numerous
Williams

near-permanent

Fork River are:

(late Mesozoic).

of the Browns Park

These rocks consist of: " ••• chalky white

to light gray soft crossbedded
gray sandy mudstone"

lava flows

sandstone with. thin layers of

or permanent drainages

tributary

to the

Milk Creek, ¥~rapos Creek, Waddel Creek, South

Fork and East Fork of l-lilliamsFork.

Climate.--National

Weather

Station data concerning

climatic conditions

in Unit 12 are available only from Craig and Hamilton.
on the northwest
that precipitation
thousand

corner and the west boundary

feet higher.

These stations are

of Unit 12.

is less at Craig than at Hamilton,

Table 1 indicates

which is at least one

The eastern portions of Williams Fork, being still

higher, probably receive around ten inches more precipitation
Steamboat

with-

Springs 23.47).

Long-time mean precipitation

is 15.8 inches, ranging from 13.09 to 19.23.
the range was 8.91-24.06.

Cloudburst

(Marvine 20.1,

for the two stations

During the period 1961-1974

storms of high intensity and short

�-118-

duration

occur over small areas and result from convective

As in adjoining
uniform,

units, seasonal distribution

higher elevations

receiving

conditions.

of precipitation

a greater proportion

is fairly

in winter in

the form of snow.

Mean annual temperature
40.2 to 44.4.

(Craig only) is 42.5, with a range (1961-1974) of

Craig has the only pertinent

data concerning

frost-free

days,

where it averaged 103.5 days and ranged from 69-125 days (1961-1974).

Table 1. Precipitation, temperature and frost-free days, Wildlife Management Unit 12, by period from National Weather Station records, 1936-1974.

PreciEitation
Mean
Range

TemEerature
Mean
Range

Frost
Free-Da~s
Mean
Range

42.2

103.5

Station

Period

Years

Craig

1961-1974

(13)

13.18

1951-1960

( 9)

13.09

42.6

1936-1952

(16)

13.94

42.3

1961-1974

(13)

18.05 11.71-24.06

1951-1960

( 9)

17.22

1947-1952

( 5)

19.23

Hamilton

8.91-16.63

Literature
U. S. Department

of Agriculture.

Water and related land resources,

Yampa River Basin Colorado and Wyoming.
Board and U.S.D.A.

Soil Conserv.

Service.

U. S. Department
Monthly

ofConnnerce.

Asheville,

Coop. Study by Colo. Water

Econ. Res. Service, Forest Servf ce, and

April 1969.

Denver.

(Continuing).

and Annual Summaries.

69-125

Cited

1969.

Conservation

40.2-44.4

164p.

(processed).

Climatological

data.

Colorado

National Climatic Center, Fed. Bldg.,

N.C. 28801.
William T. HcKean
September 1975

�-119LANDOW1'ERSHIP STATUS - WILDLIFE MANAGEMENT
Bureau of Land Management
Private
Municipal

UNIT 12

21,965 Acres

lands

152,230 Acres

and County lands

15 Acres

Other Federal lands including National Park
Service, Forest Service, Indian and Military
Reservations, and Bureau of Sport Fisheries
and Wildlif e
Colorado Division

of Wildlife

State Land Board administration

133,515 Acres

lands

2,039 Acres

lands (Moffat Co.)

Total

4,540 Acres

314,304 Acres*
(491.1 sq. miles)

*

Bureau of Land Nanagement lands, priVate lands, Colorado Division of Wildlife
lands, State Land Board Administration lands, and a total unit acreage were
derived from a 1/2" = 1 mile B.L.N. planning unit map (1974). A planimeter
and grid were used to compute acreages.
Nunicipal and County lands were derived from an estimate by the Moffat County Assessors office.
Forest Service
lands were derived by subtracting the above lands from the total unit acreage.

B. D. Trindle
August 1975

�-120-

LAND USE AND VEGETATIVE

Irrigated

cropland

Non-irrigated
Grassland

COVER STATUS - WILDLIFE MANAGEMENT

12,642 Acres

cropland

2,785 Acres

with half-shrub

mixture

23,805 Acres

Sagebrush
Brush

47,048 Acres

(desert and mountain)

Woodland

(pinon-juniper

97,175 Acres

and oakbrush)

1,375 Acres

Aspen
Commercial

UNIT 12

86,137 Acres
timber

Miscellaneous

39,356 Acres

11

3,661 Acres

Total

313,984 Acres*
(490.6 sq. miles)

*Total land acreage differs from total in Unit Description

Section (314,304
acres or 491.1 sq. miles).
Cover type categories were derived from 1/2" =
1 mile S.C.S. Land Use and Cover Type Maps, 1953, 1955, for Moffat, Rio
Blanco, Routt, and Garfield counties.
A planimeter and grid were used to
derive and compute acreage.

!I
Miscellaneous - Interpreted to consist mainly of bare ground with sparse,
scattered vegetation of various types.

B. D. Trind1e
August 1975

�-121HUMAN POPULATION

- WILDLIFE MANAGEMENT

UNIT 12

This unit includes small portions of four counties: Moffat, Rio Blanco,
Routt and Garfield.

There are no incorporated

towns within its boundaries.

The hamlet of Hamilton contains an estimated

30 persons.

Rural population

estimates made by the Moffat County Assessors

office for the unit were:

Moffat County portion - 60; Rio Blanco County portion - 10; Routt County
portion - 50; Garfield
lation,

County portion - O.

including H~i1ton,

Human population

A total estimated

rural popu-

is 150.

iensities per square mile for all of Moffat, Rio Blanco

and Routt counties

from 1960 and 1970 were:
1960

1970 II

Moffat County

1.5

1.4

Rio Blanco County

1.6

1.5

Routt County

2.5

2.8

The small portions of Routt County included would probably have a population density similar to Moffat County.

Table 1.

Urban-rural

population,

Moffat County, 1960-1970.

II

1960

1970

3,984

4,270

318

247

Rural

2,759

2,073

Total

7,061

6,525

Craig
Dinosaur

(Artesia)

11
U. S. Bureau of the Census and Colorado Yearbook,

1962-64.

�-122-

Table 2.

Urban-rural

population,

Rio Blanco County, 1960-1970.

1960

1970

Meeker

II

1,597

Rangely

1,464

1,591

Rural

2,031

1,654

Total

5,150

4,842

One projection

of human populations

II for Rio Blanco County is: 1975 _

5,270; 1980 - 5,267; 1990 - 6,119; 2000 - 7,088.

For Moffat County: 1975 _

7,206; 1980 - 7,485; 1990 - 8,589; 2000 - 10,226.

Another

projection

of human population

11 for the White River Basin is:

1980 - 20,000; 2000 - 186,000; 2020 - 204,000.

A third projection
production

~

for the oil shale area, based upon cumulative

in barrels per day, is: 1974 - 6,631;1980

oil

- 46,738; 1985 - 100,000.

II

u. S. Bureau of the Census and Colorado Yearbook,

1962-64.

21
- Colorado State Planning Office, advance population
personal communication 1975.

projections,

unpublished

31
- U. S. Bureau of the Census, Colorado Water Conservation
Field Party.
In U.S.D.A. 1965 £R cit.

Board and USDA

U. S. Dept. of Interior (1972). Draft environmental statement for the
proposed prototype oil shale leasing program. Vol. I. Washington, D.C.
465p.

�-123III

-c

p

5
5,150

III
III

;:l
0
..c::

4

~

~

4,842
3,135

3

p
0
"ri

~

2,980

2

2,943

III

r-I

;:l

p..
0
p.,

1

1,690

1900

1910

Fig. 1.

Population

1920

1930

1940

1950

1960

1970

trend of Rio Blanco County.

11
10

9
~-----------

III

-c

a

7
6,525
6,050

III

;:l

0

..c::

Routt County

6

~

~
p

Moffat County
5

0
"ri

~III

r-I

;:l

e,

4

0
p.,

3,661

2

1

1900
Fig. 2.

1910 1920 1930 1940 1950 1960 1970
Population

trends for Routt and Moffat counties.

�-124-

LAND USE STATUS RELATING TO ELK SUMMER AND WINTER RANGES
WHITE RIVER ELK HERD !/

During the summer seasons of 1961, 1962 and 1963 the White River elk
winter range was surveyed by the Colorado Division of Wildlife

using

the Big Game Analysis

technique

The

following

and tables (1, 2, 3, 4 and 5) are taken from

information

that survey

(U. S. Forest Service 1958).

(Boyd 1970).

Summer Range

i

Summer range of the Rocky Mountain
consists

of all the land above 8,500 feet elevation.

characterized
spruce-fir

elk (Cervus canadensis

by large open parks surrounded

forest, and scattered

nelsoni

Bailey)

This land is

by dense stands of dead

to extensive aspen stands.

Winter Range
Four basic vegetative

types occur on the area surveyed.

. consist of aspen, mountain

browse, pinon-juniper

the four major claSSifications,

These types

and sagebrush.

18 sub-types occurred,

Within

as listed in Table

1.

!/
The White River elk herd study area consists of Wildlife Management
Units 12,13,
23,·24, 25, 26, 33, 34.

�-125LAND USE STATUS RELATING TO ELK SUMMER AND WINTER RANGES

Table 1.

Vegetative

types and associated

(continued)

acreages on elk winter range.

Type

Acres

1.

Sagebrush

type (4-Artr)

2.

Mountain

browse consisting

of oakbrush

(5-QUE)

71,620

3.

Mountain

browse consisting

of snowberry

(5-SYM)

2,296

4.

Mountain

browse consisting

of serviceberry

5.

Mountain

browse consisting

of mountain mahogany

6.

Mountain

browse consisting

of chokecherry

(5-Prvi)

349

7.

Mountain

browse consisting

of rabbitbrush

(5-Chvd)

738

8.

Pi~on-juniper

with sagebrush understory

(9-PJ-Artr)

3,709

9.

piDon-juniper

with mountain mahogany understory

10.

pillon-juniper with oakbrush understory

(9-PJ-QUE)

1,763

11.

Pinon-juniper

(9-PJ-SYM)

621

12.

Aspen with snowberry understory

13.

Aspen with serviceberry

understory

(lO-A-AME)

12,411

14.

Aspen with chokecherry

understory

(10-A-Prvi)

2,824

15.

Aspen with Oregon grape understory

(lO-A-Mare)

163

16.

Aspen with oakbrush understory

17.

Aspen with rose understory

18.

Aspen with buffaloberry

understory

In order to more completely

understand

analysis

findings

46,583

with snowberry understory

(5-AME)
(5-CEMO)

shown in Tables 2 and 3.

1,852

(9-PJ-Cemo)

(lO-A-SYM)

2,906

27,213

(lO-A-QUE)

40

(lO-A-ROS)

424

(10-A-Shca)

194

terms included in winter range

in Tables 4 and 5 for the above vegetative

summary of the browse condition

6,197

and soil stability

condition

types, a brief
scorecards

are

�-126LAND USE STATUS RELATING TO ELK SUHMER AND WL~TER RANGES (ccn t Inued)

Table 2.

Browse condition class scorecard.

Rating

Item

Composition
High:

Desirable

and intermediate

species

(must be two or more) making

up at least 75% or more of the composition,

with desirables

at

least 45% of the composition.
Medium:

Desirable

and intermediate

composition,
Low:

Desirable

species making up 50% or more of the

with desirables

at least 15% of the composition.

and intermediate making up at least 50% of the

composition.
Density
Very dense:

66% plus

High

36% to 65%

Medium

16% to 35%

Low

15% minus

High:

Hedging of key species mostly light or moderate with less
than 16% of the plants heavily hedges, and decadent minus
young less than 15% of the total number of plants.

Medium:

Hedging of key species mostly mod.erate, not more than 35%
heavily hedged and decadent minus young not more than 35%
of the total number of plants.

Low:

More than 35% of the plants of key species heavily hedged or
decadent minus young more than 35% of the total number of
plants.

�-127-

LAND USE STATUS RELATING TO ELK SUMMER AND WINTER RANGES (continued)
Table 3. Soil stability condition class scorecard.
Rating

Item

High:

Soil movement slight or none, usually 65 or more hits on
ground cover and rocks.

Medium:

Soil movement moderate, usually 35-65 hits on ground cover
and rocks.

Low:

Soil movement advanced, usually less than 35 hits on ground
cover and rocks.

Table 4. Summary of browse composition, density and vigor on elk winter
range as determined by 640 transects in the White River elk study area.
Vegetative
Type

Low

4-Artr

92

35

2

20

93

5-QUE

19

144

19

3

5-SYM

64

73

3

5-AME

0

10

5-Cemo

o

5-Prvi

Density
Hed. High

Low

Vigor
Med.

High

13

47

25

57

60

119

29

62

91

5

48

87

20

36

84

12

0

9

13

4

8

10

5

18

2

14

7

14

6

3

0

2

0

0

1

1

0

0

2

5-Chvd

6

1

1

2

5

0

2

2

4

9-pJ

2

7

8

14

3

0

12

1

4

10-A

57

49

11

51

54

12

3

8

106

Total

240

326

74

98

290

252

131

148

361

Percent

37.5 50.9

11.6

15.3

45.3

39.4

20.5

23.1

56.4

I

•

ComEosition
Med.
High

Low

�-128-

LAND USE STATUS RELATING TO ELK SUMMER AND WINTER RANGES (continued)

. Table 5. Summary of soil conditions on elk winter range as determined
by 640 transects in Wildlife Management Units 23 and 24.
Vegetative
Type

Low

Soil Condition
Medium

High

33

64

32

41

84

57

23

55

62

9

9

4

14

8

1

0

1

1

2

2

0

16

1

0

1

8

108

Total

139

232

269

Percent

21.7

36.3

42.0

4-Artr
5-QUE
5-SYM
5-AME
5-Cemo
5-Prvi
5-Chvd
9-PJ
10-A

LITERATURE CITED
Boyd, R. J. 1970. Elk of the White River Plateau, Colorado.
of Wildl. Tech. Publ. No. 25. l26p.
U. S. Forest Service. 1962. Big game range analysis.
Rocky Mtn. Reg. USDA Publ. 49p.

B. D. Trindle
September 1975

Colo. Div.

U. S. For. Ser.,

�-129GAME SPECIES - WILDLIFE ~~AGEMENT
Big game mammals

UNIT 12

1:./

Black bear CUrsus american~s) Uncommon.
Elk (Cervus canadensis) Common.
Mountain lion (Felis concolor) Uncommon.
Mule deer (Odoc~s
hei:lionus)COImilon.

Small game mammals

!!

Cottontail rabbit CSylvila2us auduboni; S. nuttallii)
Pine (red) squirrel (Tamiasciurus hudsonfcus) Common.
Snowshoe hare (Lepus ane rLcanus ) Common.

Small game birds
Migratory

Common.

~!

waterfowl

and shorebirds

Great Basin Canada goose CBranta canadensis moffitti) Uncommon.
Black brant (Branta nigricans) Possible rare migrant.
White-fronted goose (Anser albifrons frontalis) Possible rare migrant.
Snow goose (Chencaer~ens
caerulescens) 1/ Possible rare migrant.
hThistling swan (Olor colu12bianus) Possible rare migrant .
•mllard CAnas pl~h\~chos
DlatYrh\~chos) Common resident.

!/Nomenclature
Colorado.

according to Lechleitner, R. R. 1969.
Pruett Publishing Co., Boulder.
254 pp.

Hild maTJlll1als
of

2/
- Nomenclature from Bailey, A. M., and R. J. Niedrach.
1967. Pictorial
checklist of Colorado Birds. Denver Nus. Nat. Hist. 168 pp , Information on
occurrence IDld status aciapted from the above reference and from Cringan, A. T.,
and L. Carlson. 1973. h'ildlife in the Piceance Creek Basin, In: An environmental reconnaissance of the ~iceance Basin, Rio Bla~co and Garfield counties,
Colorado •. A report on the completion of Part 1, Phase One of the environmental
inventory, anclysis and impact study portion of the Regional Oil Shale Study
being done for the State of Colorado by the Thorne Ecological Institute,
Boulder, Colorado. 144 pp. Additional information on occurrence, in employing
the term "possible", is adapted froo the foregoing references and also from,
Davis, W. A. 1969. Birds in western Colorado.
Colo. Field Ornithologists.
61 pp.

3/
- Unverified

in hunters' bag checks but legal game 1972 through 1975.

Definitions:
Common:-should be noted regularly in the normal habitat at the
proper season of the year. Uncor.~on:-recorded regularly but unlikely to be
seen on most field trips. Rare:-recorded a few times only, and very unlikely
to be seen. Resident:-any species living in the unit. Migrant:-any species
moving through th~it.
Where the adjective "possible" is absent, actual
sightings have been reported by Division personnel or qualified observers.

�-130-

GAME SPECIES - WILDLIFE

HANAGEHENr

UNIT 12 (continued)

Small game birds
Gadwall (Anas strepera) Common spring and fall migrant.
Pintail (Anas acuta) Common spring and fall migrant.
American green-winged teal (Anas crecca carolinensfs) 41 Common migrant
and uncommon yearlong resident.
Blue-winged teal CAnas discors discors) Common migrant.
Cinnamon teal CAnascyanoptera
septentrionalium) Common migrant.
American wigeon (Anas ~~ericana) 4/ Common migrant and rare winter resident.
Northern shoveler (Anas clypeata)-i/ Common migrant and uncommon summer
resident.
Redhead (Aythya americana) Uncommon migrant.
Ring-necked duck (Aythva collaris) Uncommon migrant.
Canvasback (Aythya valisineria) Unccmmon to rare migrant.
Greater scaup (Avthya ~arila n~arctica) Possible rare resident.
Lesser scaup (Aythya affinis) Uncommon migrant.
Common golden-eye (Eucephala c1angula americana) Common migrant and uncommon winter resident.
Barrow's golden-eye (Bucephala ~slandica) 11 Rare winter visitor.
Bufflehead (Bucephala albeola) Uncommon spring and fall migrant and
rare winter resident.
Ruddy duck (O:0{ura jamaicensis rubida) Common migrant and occasional
summer resident.
Hooded merganser (Lophodytes cucullatus) Rare winter visitor on river.
Common merganser (~!ergu~ merganser ame r i canu s) Common winter resident,
uncommon summer resident.
Red-breasted merganser e·lergus serra tor serrator) 11 Possible rare migrant.
American coot (Fulica americana americana) Common migrant and summer resident.
Common Wilson's snipe (Capella gallinago delicata) Common migrant and rare
winter resident. .
Sandhill crane (Grus canadensis tabida) Regular migrant.
Virginia rail (Ralus limicola limicola) Possible uncommon summer resident.
Sora (Porzana carolina) Possible uncommon summer resident.

Upland game birds
Blue grouse (Dendragapus obscurus obscurus) Common.
Sage grouse (Centrocercus urophasianus uroohasianus) Un common to common.
Ring-necked pheasant (Phasianus colchicus) Uncommon.
Chukar (Alectoris chukar) 4/ Uncommon to rare.
Band-tailed pige;n (Columb; fasciata fasciata) Possible uncommon summer
and fall migrant.
Mourning dove (Zenaida macroura marginel1a) il Common summer resident.

4/

Sharp-tailed grouse (Pedioecetes phasianellus columbianus)
Wild turkey (Meleagris gallopavo merriami) Rare.

Uncommon.

- Changes in the nomenclature follow the thirty-second supplement to the
American Ornithologists Union check-list of North American birds published
in Auk 90:411-419, April, 1973.

W. T. McKean
September

1975

�-131OTHER MAMMALIAN SPECIES 1/_ WILDLIFE MANAGEMENT UNIT 12
Furbearers ];/
Short-haired
Beaver (Castor canadensis) Uncommon.
Mink (Mustela vision) Uncertain.
Muskrat (Ondatra zibethicus) Uncommon.
Ringtail (Bassariscus astutus) Rare.
Weasels (Hustela e.rminea, M. frenata) M. erminea Uncertain; M. frenata
Uncommon.
Long-haired
Kit fox (Vulpes velox) Uncommon.
Gray fox (Urocyon cinereoargenteus) Uncommon.
American badger (Taxidea taxus) Common to uncommon.
Spotted skunk (Spilogale ~ius)
Uncommon.
Striped skunk (Mephitis mephitis) Common.
"Varmint" mammals
Coyote (Canis latrans) Common.
Red fox (Vulpe~. !ulva) Uncommon.
Raccoon (Procvc1"'lotor) Uncommon.
Porcupine (Erethizon dorsatum) Common.
Bob.cat.(v., i ~-;-1'··)(Lynxrufus) Common.
Black-tailed ,;ck -;.iLbit(Lepus californicus) Rare to uncommon.
White-tailed j.:~ckrabbit (Lepus townsendii) Common.
Yellow-bellied marmot (Marmota flaviventris) Common.
lmite-tailed prairie dog (Cynomys leucurus) Uncommon.
Richardson's ground squirrel (Spermophilus richardsonii) Common.
Thirteen-lined ground squirrel (Spermophilus tridecemlineatus) Common.
Rock squirrel
(Spermophilus variegatus) Common to uncommon.
Northern pocket gopher (Thomomys talpoides) Common

1/

- These spe cfes , gr.ouped separately as "Furbearers", "Varmints" and "Nongame
mammals" and c.uts i.«- of "g ame" categories, follow Chapter 62, Colo. Rev.
Statutues 1963 As hfiel:.led,inColo. Game, Fish and Parks Div. Laws and
Regulations Hdbk., 1975 (Art. 1, Items 14, 20, 31, and 42, Definitions, p. 2).
IINomenclature from Lechleitner, R. R. 1969. Wild mammals of Colorado.
Pruett Publishing Co., Boulder. 254 pp. Information on occurrence and status
from the above reference and: Cringan, A. T., and L. Carlson. 1973.
Wildlife. in the Piceance Creek Basin, In: An environmental reconnaissance of
the Piceance Basin, Rio Blanco and Garfield counties, Colorado. A report
on the completion of Part 1, Phase One of the environmental inventory, analysis
and impact study portion of the Regional Oil Shale Study being done for the
State of Colorado by the Thorne Ecological Institute, Boulder, Colorado, l44pp.
Also, Arms t roig , D. N. 1972. Distribution of mammals in Colorado. Monograph
of the Ml!seum of Natural History, the Univ. of Kansas, Numb er 3, 1972. 4l5pp.

�-l32Definitions:
Connnon:-shouid be noted regularly in the normal habitat at the
proper season of the year. Uncommon:-recorded regularly but unlikely to be
seen on most field trips. Rare:-recorded a few times only, and very unlikely
to be seen. Resident:-any species living in the unit. Migrant:-any species
moving through the unit. Where the adjective "possible" is absent, actual
sightings have been reported by Division personnel or qualified observers.
Nongame mammals
Golden-mantled ground squirrel (Spermophilus lateralis) Common;
White-tailed antelope squirrel (Ammosoermophilus leucurus) Uncommon.
Least chipmunk (Eutamias minimus) Common.
Colorado Chipmun~ (Eutamias quadrivittatus) Connnon.
Unita ch f.pmun k (Eutamias umbrinus) Un conmori to uncertain.
Vagrant shrew (Sorex vagrans) Uncommon.
Merriams shrew (Sorex merriami) 3/ Uncertain.
Masked shrew (Sorex cinereus) Un~ommon.
Ta.vnsend's big-eared bat (Plecotus townsendii) Common.
Silver-haired bat (Lasionycteris noctivagans) Uncommon.
Hoary bat (Lasiurus cinereus) Uncommon - common.
Big brown bat (Eptesicus fuscus) Common.
Western pipistrelle (Pipistrellus hesperus) Common.
Long-legged myotis(Myotis
volans) Uncommon.
California myotis (Myotis californicus) Uncommon.
Small-footed myotis (Myotis leibii) Common.
Long-eared myotis (Myotis evotis) Uncommon.
Little brown myotis (Myotis lucifugus) Uncommon.
Ord's kangaroo rat (Dipodomys ordii) Un common - uncertain.
Apache pocket mouse (Perognathus apache) Uncommon.
Western harvest mouse (Reithrodontomys magalolis) Uncertain.
Canyon mouse (Peromyscus crinitus) Common - uncommon.
Deer mouse (Peromyscus maniculatus) Common.
Pinon mouse (Peromyscus truei) Common.
Bushy-tailed wood rat (Neotoma cinerea) Common.
Desert wood rat (Neotoma lepida) Rare.
Gapper's red-backed vole (Clethrionomys gapoeri) Uncommon.
Montane vole (Mi:rotus montanus) Uncommon.
Long-tailed vole (Microtus longicaudus) Common.
Sagebrush vole (Lagurus curtatus) Common.
House mouse (Hus musculus) Uncommon.
~estern jumping mouse (Z~pus princeps) Uncommon.

~/Occurrence listed by the Colo. Div. of Wildlife as extremely unusual - very
few documented records within the past decade. (1972 Status Evaluation for
Selected Colorado Species) appended to: 1973 Wildlife Operations Work Plan,
Field Order No. 4 - 1973.

W. T. McKean
November 1975

�-133OTHER AVIAN SPECIES II - WILDLIFE MANAGEMENT UNIT 12
"Varmint" birds
English sparrow (Passer domesticus domesticus) Common resident.
Black-billed magp~Pica
pica 11Udsonla) Common resident.
Starling (Sturnus vu1~s)
Co~on ~esident.
Rock dove (Columbia livia) Uncommon resident.
Nongame birds 11
Common loon (Gavia immer) Possible rare migrant.
Horned grebe (Podiceps auritus cornutus) Possible rare migrant.
Pied-billed grebe (Podi1ymbus podiceps podiceps) Possible uncommon
migrant and unco~on summer resident.
Great blue heron (Ardea herodias treganzai) Common summer resident.
Snowy egret (Egretta thula brewsteri) Uncommon summer resident 11.
Black-crowned night heron (Nycticorax nycticorax hoact1i) Common
swnmer resident.
White-faced ibis ~legadis chihi) Uncommon migrant.
Semipalmated plover (Charaarius semipa1matus) Possible rare migrant.
Killdeer (Charadiu~ vociferus vociferus) Common summer resident and
rare winter resident.
Mountain plover (Charadrius montanus) Possible rare migrant 11.

!I
These species, grouped separately as "Varmint" birds, "Nongame birds"
and "Raptores" and outside of "game" categories, follow Chapter 62, Colo.
Rev. Statutes 1962 As Amended, in Colo. Game, Fish and Parks Div. Laws and
Regulations Hdbk., 1975. (Art. 1, Items 20, 28, 31, and 42, Definitions,
p , 2).

1/
Nomenclature from Bailey, A. M., and R. J. Niedrach.
1967. Pictorial
checklist of Colorado Birds. Denver Mus. Nat. Hist. 168 pp. Information on
occurrence and status adapted from the above reference and Davis, W. A.
1969. Birds in western Colorado. Colorado Field Orrif.t hoLog Lst.s, 61 pp ,

3/
- Changes in nomenclature follow the Thirty-second Supplement to the American
Ornithologists Union Checklist of North American Birds published in Auk
90(2):411-419.
April, 1973.
Definitions:
Common:-should be noted regularly in the normal habitat at the
proper season of the year. Uncommon:-recorded regularly but unlikely to be
seen on most field trips. Rare:-recorded a few times only, and very unlikely
to be seen. Resident:-any species living in the unit. Higran~:-any species
moving through the' unit. 'iVherethe adj ective "possible" is absent, actual
sightings have been reported by Division personnel or qualified observers.
Notations concerning seasonal abundance of raptors are by Gerald R. Craig,
Wildlife Biologist.

�-134OTHERAVIAl'i

SPECIES (continued)

Nongame birds
Black-bellied
plover
(Pluvialis
squatarola)
Possible
uncornmon migrant
3/.
.Long-billed
cur-lew (Numenius americanllS americanus)
Possible
rare migr;nt.
Spotted
sandpiper
(Arctitis
Placularia)
Possible
common summer resident.
Solitary
sandpiper
(Trin&gt;'a solitaria
cinnamomea) Possible
COIrJl1on
migrant.
and occasional
summer resident.
Greater
sandhill
crane
(Crus c:madensis
possible
surr~er resident
endangered.

ta~Jida)

Regular

migrant

and

Willet
(CataptroDhorus
semipalmatus
inornatus)
Possible
rare migrant.
Greater
yellm-llegs
(Tringa rr:elanoleuca)
Possible
common migrant
3/.
Lesser
yelloH'legs
(Tl~Lng~ f1avipes)
Possible
uncommon migrant 1/:E'.~dknot (Calidris
C2~.ltus r uf a ) Possible
rare migrant ]).
Pectoral
sandpi.per
(C&lt;,l.lidris med.anotos) Possible
rare migrant
3/.
Baird's
sandpiper
(Caliclris
bairdii)
Possible
common migrant
Least sandpiper
(C81~~i~tili-;-)
Possible
common migrant 3/.
J.ong-billed
dowi t che r (Lii":moclromus scolopaceus)
Possible
unco;'Jl1on migrant.
Stilt
sandpiper
(Nicropalmn }!i:nantopus)
Possible
rare migrant.
Semipalmated
sandpiper
(Caliclris
~~ill~)
Possible
rare migrant ]./.
l-lestern sandpiper
(Calidris
maur i) Possible
UnCOITUllOn
migrant
3/.
Harbled
godwi t (Limosa fedoa) Possible
rare spring migrant.
Sanderling
(Calidris
alba)
Possible
rare migrant
3/.
American avocet
(Recu"i--V:;:-~ostraaii1erjc •.
ma) Rare migrant.
Black-necked
stilt
(!!;:n:lantopus me}:ic&lt;lnu~) Possible
rare migrant.
llilson's
phalarope
(Steganopus
tricolor)
Possible
common migrant and
uncommon summer- resident.

37.

Northern
phalarope
(Lob~
lobatus)
Possible
uncommon migrant.
Herring
gull
(Larus argentatus
S!"'1ithsonian~~) Possible
unCOIl1ffion
migrant.
California
gull
(Larus cali.fornicus)
Possible
rare migrant.
Ring-billed
gull
(Larus deLmarensis)
Possible
uncommon migrant.
Franklin's
gull
(La ru s p-ipixcan)
Possible
unCOTllITlon
migrant.
Bonaparte's
gull
(~~~il~~elphia)
Possible
rare migrant •
.Sabine's
gull
(Xem3 sabini
s2bini)
Possible
rare migrant.
Forster's
tern (St;rna
fors~~possible
rare migrant.
Common tern
(Sterna
hirundo hirundo)
Possible
rare migrant.
Least tern
(Sterna'albafrons
2tba12sso~)
Possible
rare migrant.
Black tern
(Chlidonias
niger
sur~namensis)
Possible
uncommon migrant.
Rock dove (CoLunb a Ltv I a ) Possible
common resident.
Unprotected.
YelloH-billed
cuckoo (CoccyZUS amer'iC;1llUS americanus)
Possible
rare
summer resident.
Poor-,vill
(Phalaenoptilus
nuttallii
Common n Lgh t hawk (Chorde i.Les minor
surrmer resident.

nuttallii)
besperis;

Uncommon summer resident.
C. m, howeI L) Common
-

White-throated
swt.f t (Acronautes
saxatalis
sc.Lat cr L) Common summer resident.
Black-chinned
hummingbird
(Archi.lochus
aLexand r I ) Uuconrnon summer resident.
Broad-tailed
hurruningblrcl (Selasphorus
platycercus
platycercus)
Common
summer resident.
Rufous hummingbird
(Sela~;phorus .£ufus) Possible
common late
summer migrant.
Calliope
hUllliningbJrd (Stel1ula
cal1iore)
Possible
common late
summer
migrant.
Leltt.,J kingfisher
(Hegaceryle
nlcyon nlcyon) Possible
common resident.
Common flicker
(Col~tcs
au r a t us carer)
Commonresident
Lewf.s" woodpecker
(Asvndo smun le\vis)
Possible
uncommon summer resJdent.
Red-headed
woodpecker
(!'!cl-;;l~rPc~rythrocephaltls)
Possible
rare resident.

1/.

�-135-

OTHER AVIAN SPECIES

(continued)

Nongame birds
Pileated woodpecker
resident.

(Dryocopus pileatus oicinus) Possible rare summer

Yellow-bellied sapsucker (Sphyrapicus varius nuchaliae) Common summer
and possible occasional vlinter resident.
Williamson's sapsucker (Sphyrapicus thyroideus nataliae) Possible
uncommon summer resident.
Hairy woodpecker (Dendrocopos villosus monticola) Common year-a-round
resident.
Downy \'loodpecker (Dendroc~")os l'ubescens leucurus) Common year-a-round
resident.
Eastern kingbjrd (Tyrannus tyrannus) Uncommon summer resident.
Western kingbird (Tyrili~nusverticalis) Common Su~mer resident.
Ash-throated flycatcher (?!riarchus_cinerascens cinerascens) Common summer
resident.
Say's phoebe
resident.

(Sayornis ~

saya) Common summer and possible rare winter

Willow flycatcher (Empidonas traillii) Possible uncommon summer resident )./.
Hammond's flycatcher (Emoidonax hammondii) Possible migrant and uncommon
summer resident.
Dusky flycatcher (Empidonax oberholseri) Summer resident.
Gray flycatcher (EmDidonax wrightii) Common summer resident.
Western flycatcher (Emoidonax dHficilis hellmavri) Common summer resident.
Western wood :)ee~"ee(Contopus ;-~~
~iei)
Common summer resident.
Olive-sided flycatcher (Nuttallornis borealis) Possible common summer
resident.
Horned lark (Erernophila alpestris leucolaerna) Common year-a-round resident
and migran t ,
Violet-green swallow '(Trachvcineta thalassina lepida) Common summer resident.
Tree swallow (Iridoorocne bicolor) Cornman migrant and sununer resident.
Bank swallow (~aria
riparia riparia) Possible uncommon migrant and
summer resident.
Rough-wf.nged
swa l Loo (Stelgidooteryx ruficollis serripennis) Uncommon
migrant and summer resident.
Barn swallow (Hirundo rustica erythrogaster) Cocrrnonsummer resident.
Cliff swallow (Petrochelidon ovrrhonota pvrrhonota) Conunon summer resident.
Gray jay (Perisoreus canadensis caDitalis) Possible uncommon resident.
Steller's jay (Cyanocitta stelleri macrolopha) Common resident.
Scrub jay (Aphelocoma coerulescens woodhouseii) Common resident.
Common raven (Corvus corax sinuatus) Common resident.
Common crow (Corvus b~rhynchos
brachyrhvnchos) Uncommon resident.
Pinon jay (Gymnorhjnus cvanoceohalus) Common SUllli"fier
)./ and winter resident.
Clark's nutcracker (Nucifraga columbiana) Common resident.
Black-capped chickadee (Paru~ articapillus garrinus) Common resident.
Hountain chickadee (Parus gambeli g&lt;1moeli) Common resident.
Plain titmouse (Parus_ inornatus ridg~"ayi) Uncommon resident.
Bushtit (Psaltriparus minimus olumbeus) Possible cornman resident 1/.
White-breasted nuthatch (Sitta carolinensis nelsoni) Possible' uncommon
resident.
Red-breasted nuthatch (Sitta canadensis) Possible rare year-a-round resident.
Pygmy nuthatch (Sitta pygmaea rnelanotis) Possible uncommon resident.

�-136-

OTHER AVIAN SPECIES (continued)
Nongame birds
Brown creeper (Certhia famlliarismorttana) Possible uncommon year-around resident.
Dipper (Cinclus mexicanus unico10r) Common resident.
House wren (Trog~~edon
parkmanii) Common summer resident.
Bewick's wren (Thrvooanes pe,.;ickiieremoohilus) Possible summe r resident
and rare winter resident.
Canyon wren (Sa1pinctes mexicanus consperus) Possible uncommon summer
resident.
Rock wr en (~.?lpinctes obso1etus obsoletu~) Common summer and possible
rare winter resident •
.Hock Ingb Lrd (!-fiouspol vz Lo t t os leucooterus) Uncommon summer resident.
Gray catbird (Dumete11a caroli~ensis) Possible rare summer resident 3/.
Sage thrasher (Oreoscops_es ::lontanus)Possible common summer resident-:American rob In (Turdus !2igratorius propinquus) Common summer and winter
resident 3/.
Hermit thrush (Catharus guttatus audubonii)
Common summer resident 1/.
Swainson's thrush (Catharus ustulatus almae) Uncommon migrant.
Veery (Catharus fucescens salicico1a) Uncommon migrant and summer resident 1/.
Western bluebird (5i211a oex1cana baird i) Common migrant and uncommon
summer resident.
Mountain bluebird (S1a1ia currucoides) Common migrant and summer resident
and possible occasional winter resident.
Townsend's solitaire C.fvadestes townsendi tovnsend L) Uncommon r-esLden t ,
Blue-gray gnatcatcher (Poliptila-C;~ru1ea amoenissima) Common summer
resident.
Golden-cro'-'11edkinglet (Regulus satrapa amoenus) Possible uncommon
migrant and rare sunner resident.
Ruby-crowned kinglet (Regulus calendula cineraceus) Possible uncommon
migrant and sumwer resident.
Bohemian waxwing (Bor::bvcillagarrulus pallidiceps) Possible irregular
winter migrant 1/.
Cedar waxwf.ng (Bornbvc i Ll.a cedrorum) Uncommon resident.
Northern shrike (Lanius excub1tor invictus) Common winter resident.
Loggerhead shrike (L2.:1iusluiovic1anus excubitorides) Possible uncommon
summer and unco~on winter resident.
Gray vireo (Vireo vicinioc) Possibl~ COlll..'!lon
summer resident.
Solitary vireo (Vireo soli-tarius pLumb eu s) Common summer resident.
Red-eyed vireo (Vireo oliv2c2us) Possible rare summer resident.
Warbling vireo (Vireo gi~ '/11S 5"ainsonii) Common summer resident.
Tennessee warbler (VelTIivora oere£rina) Possible rare but regular migrant.
Orange-crow'I1ed warbler (Vermivora celata orestera) Common migrant and
summer resident.
Nashville warbler (Vernivora ruficapilla ridgwayi) Possible rare migrant.
Virginia's warbler (Vermivora viYS!iniae) Common surnmer resident.
Yellow warbler (Dendroica petecnia aestiva) Common su~uer resident.
Ye.I Low-rrurupcd warbler (Dcndroica coronata memorabilis) Common summer
resident 3/; (D. c. co ronn t a) Possible common migrant 3/.
Black-throat;d gr-;;-y
~-;arble.r(DE:ndroica nigrescens) Possible common summer
resident.
Townsend's warbler (Dendroica towu·sendi) Uncommon fall migrant.
MacGillivray's warbler (Oporornis tolmiei monticola) Common summer resident.

�-137-

OTHER AVIAN SPECIES

(continued)

Nongame birds
Common yellowthroat (Ceothylpis trichas occidentalis: Q. ~. campicola)
Possible uncommon summer- resident 3/.
Yellow-breasted chat (Icteria virens auricollis) Possible common summer
resident.
..
Wilsonis warbler (Wilsonia pusilla pileolata) Possible common migrant
and summer resident.
American redstart (Setophaga ruticilla tricolora) Possible rare migrant.
House sparrow (Passerdomesticus
domesticus) Common resident.
Bobolink (Dolichonyx orvzivorus) Possible uncommon summer migrant.
Western meadowlark (Sturnella neglecta neglecta) Common summer and
possible uncommon winter resident.
Yellow-headed blackbird (Xanthocephalus xanthocephalus) Common summer
resident.
Red-winged blackbird (Agelaius phoeniceus fortis) Common resident.
Northern oriole (Icterus galbula bullockii) Possible common summer
resident 3/.
.
.
Rusty blackbird (Euphagus carolinus carolinus) Possible rare winter migrant.
Bre'ver's blackbird (Euphagus cvanoceDhalus) Common resident.
Common grackle (Quiscalus quiscula) Possible rare su~er resident.
Brown-headed cowbird (}lolothrus ater artemisiae) Common summer resident.
Western tanager (Piranga ludovic~)
Common migrant and summer resident.
Scarlet tanager (Piran~ olivacea) Possible rare migrant.
Black-headed grosbeak (Pheucticus melanocephalus melanoceuhalus) Uncommon
summer resident.
Lazuli bunting (Passerina amoena) Common summer resident.
Evening grosbeak (Hesperiphona vespertina brooksi) Uncommon summer
resident and irregular winter resident.
Cassin's finch (Carpodacus cassinii) Common resident.
House finch (Carpodacus mexicanus frontalis) Commonsummerand_possiple
uncommon winter resident.
Pine grosbeak (Pinicola enucleator montana) Possible uncommon resident.
Gray-cro,med rosy finch (Leucosticte teuhrocotis tephrocotis; L. t.
littoralis) Possible common winter resident.
Black rosy finch (Leucosticte atrata) Possible common winter migrant.
Brotvn-capped rosy finch (Leucosticte australis) Possible common winter
migrant.
Common redpoll (Acanthis flammea flammea) Possible rare winter migrant.
Pine siskin (Spinus pinus pinus) Common resident.
American goldfinch (Spinus tristus tristus; ~ •.~. pallidus) Possible
Common summer and uncommon winter resident.
Lesser goldfinch (Suinus usaltri.a psaltria) Possible uncommon summer and
rare winter resident.
Green-tailed towhee (Chiorura chlorura) Common summer resident.
Rufous-sided towhee (Pipilo erythrophthalmus montanus) Common summer
and rare winter resident.
Lark bunting (Calamospiza melanocorys) Possible uncommon summer resident.
Savannah sparrow (Passerculus sand\vichensis nevadensis;
anthinus)
Common migrant and Slli~er resident.
Grasshopper sparrow (Ammodramus ~avannarum perpallidus) Possible uncommon
summer resident.
Vesper sparrow (Pooecetes gramineus confinis) Common migrant and
summer resident.

.!:. ~.

�-138-

OTHER AVIAN SPECIES (continued)

Nongame birds
Lark sparrow (Chondestes grammacus strigatus) Common migrant and
summer resident.
Black-throated sparrow (Amphispiza bilineata deserticola) Possible common
resident.
Sage sparrow (Amphispiza belli nevadensis) Common summer resident.
Dark-eyed junco (Junco hyemalis aikeni) Possible rare winter resident 1/;
(~. ~. hyemalis; ~. ~. cismontanus) Rare winter resident 1/;
(J. h. oreganus) Common winter resident 3/; (J. h. o. var. mearnsi)
C~mm~n winter resident 3/.
- - Gray-headed junco (Junco c;niceps caniceps) Common summer and winter
resident.
Tree sparrow (Spizella arborea ochracea) Possible unco~~on winter migrant.
Clay-colored sparrow (Spizella pallida) Possible courrnonmigrant.
Chipping sparrow (Spizella passerina boreophila) Common summer resident.
Brewer's sparrow (Spizella breweri breweri) Common summer resident.
Harris' sparrow (Zonotrichia guerula) Possible uncourrnonmigrant and rare
winter resident.
White-crowned sparrow (Zonotrichia leucrophrys) Common resident.
White-throated sparrow (Zonotrichia albicollis) Possible rare migrant.
Fox sparrow (Passerella iliaca schistace~) Rare summer resident.
Lincoln's sparrow (~lelospiza lincolnii alticola) Possible common migrant
and summer resident.
Song sparrow: (Melospiza melodia) Common summer and uncommon winter resident.

Raptors

1/
;/

Turkey vulture (Cathartes ~
meridional is) Courrnonsummer and rare
winter resident.
Goshawk (Accipiter gentilis atricapillus) Common resident.
Sharp-shinned hawk (Accipiter striatus velox) Common resident.
Cooper's hawk (Accipiter cooperii) Comm~sident.
Red-tailed hawk (Buteo jamaicensis calurus) Common resident.
Swainson's hawk (Buteo swainsoni) Uncommon summer resident.
Rough-legged hawk (Buteo lagopus ~. johannis) Common winter resident and
migrant.
Ferruginous hawk (Buteo regalis) Uncommon resident.
Golden eagle (Aquila chrysaetos canadensis) Common resident 4/.
Bald eagle (Haliaeetus leucocephalus alascanus) Common. winter resident 4/.

�-139OTHER AVIAN SPECIES (continued)

Raptors
Marsh hawk (Circus syaneus hudsonius) Cornmon summer and uncommon
winter resident.
~
Osprey (Pandion haliaetus carolinensis) Uncommon migrant, rare
summer resident.
Prairie falcon (Falco mexicanus) Uncommon resident.
Peregrine falcon (Falco peregrinus anatum) Rare migrant, endangered.
Merlin (Falco colurnbarius) Uncommon winter migrant 3/.
American kestrel (Falco sparverius sparverius) Common summer and
uncommon winter resident.
Screech owl (Otus asio) Cornmon resident.
Flammulated owl (Otus flarnmeolus flarnrneolus)Rare summer resident.
Great horned owl (Bubo virginianus) Common resident.
Pygmy owl (Glaucidium gnoma californicum) Rare resident.
Burrowing owl (Speotyto cunicularia .bY.Eugaea)Rare resident.
Long-eared owl (Asio otus wilsonianus) Uncommon resident.
Short-eared owl (Asio~mmeus
flammeus) Common summer resident and
uncommon winter migrant.
Saw-whet owl (Aegolius acadicus acadicus) Uncommon resident.

ADDITIONAL LITERATURE REFERENCES
Cringan, A. T., and L. Carlson. 1973. Wildlife in the Piceance
Creek Basin, In: An environmental reconnaissance of the Piceance
Basin, Rio Blanco and Garfield counties, Colorado. A report on
the completion of Part 1, Phase one of the environmental inventory,
analysis and impact study portion of the Regional Oil Shale Study
being done for the State of Colorado by the Thorne Ecological
Institute, Boulder, Colorado. 144 pp.
Martin, G. S., P. H. Baldwin, and E. B. Reed. 1974. Recent records
of birds from the Yampa Valley, northwestern Colorado. Condor
76(1):113-116.
Smith, A. G. 1973. Avian environmental features at the Colony plant
site, Garfield County, Colorado. Environmental inventory by
Thorne Ecological Institute for Colony Development Operation,
Atlantic-Richfield Company, Operator, October, 1974.

W. T. McKean and B. D. Trindle
September 1975

�-140DISTRIBUTION

AND ABUNDANCE

OF BIG GAME MAMHALS

WILDLIFE MANAGEHENT

UNIT 12

BLACK BEAR
Almost

the entire unit is occupied by black bears.

section

in the extreme northwest

Only a small

part bordering Axial is excluded

from their range.

The Colorado
suggests

Division

a density

total occupied

of Wildlife,

figure ranging between

range~

(1974),

.056 and .035 mile2 for the

W.C.O. estimates for 1975 yielded a

of 15 bears and a minimum

Estimated

Inventory Systems

These density estimates yield an "overall" popu-

lation of 23 bears on Unit 12.
maximum

Wildlife

of 9 bears on Unit 12.

average annual black bear harvest for the 19 year period 1956-

1974, inclusive,

was 1.4 bears.

This is based upon hunter report card

surveys

for years 1956-1971

and a random sample for 1971-1974.

Annual

harvest

ranged from 0 to 6; there were ten years out of the nineteen

when no bears were reported taken.

ELK
The summer range of elk on Unit 12 is almost entirely restricted
two national
elevations

forests

(Routt and White).

of the unit, generally

distribution

w.e.o. Roland

the higher

above 8,000 feet in elevation.

the arrival of winter and increased
higher elevations

This area represents

to the

With

snow depths the el~ pull off of the

and move onto their winter range (see accompanying

map).

(1975 pers. comm.) estimates about 2,500 elk summer and
,
winter on Unit 12. He feels that there is some interchange between Units

�-14111, 23, and 24 but that the population

Mean annual elk harvest
Annual harvest

remains fairly constant.

(1956-1974) was 375, including archery.

ranged from 151 to 814 (see Harvest Data presented

in this report).

MULE DEER
Wildlife

Management

Unit 12 is basically divided

to mule deer winter and summer ranges.
contains higher elevations
tions in the northern
accompanying

The southern half of the unit

and is the summer range.

distribution

map).

Dividing

elevation between the two

7,500 feet.

Data provided by Colorado Division

of Wildlife,

(1974) indicated a winter population

density of 12.7 deer per square mile.
the same population

The lower eleva-

half of the unit make up the winter range (see

rIDlges is approximately

Systems

in half with respect

Wildlife

Inventory

of 2,416 animals with a

This source also assumed that

summers on the unit.

W.C.O. Roland

(1975 pers.

comm.) concurs with this estimate.

Mean annual deer harvest

(1956-1974) was 724, including archery.

harvest ranged from 303 to 1,764 (see Harvest Data presented

Annual

in this

report).

MOUNTAIN

LION

No mountain

lions are found on Unit 12, according

of Wildlife,

Wildlife

Inventory Systems

to Colorado Division

(1974), and W.C.O. 's connected

with the unit.

Since establishment

of the mountain

lion as a game species on July 1,

1965, no recorded kill exists for Wildlife Management

Unit 12.

�-142LITERATURE
Colorado

Division

Statewide

Wildlife

Inventory Systems.

mammal and bird distribution

from W.C.O.,
Located

of Wildlife,

CITED

area supervisor,

1974.

maps and data compiled

and regional biologist

at Denver office, Colorado Div. Wildlife.

input.

Unpublished.

B. D. Trind1e
December 1975

�-143DISTRIBUTION

AND ABUNDANCE OF SMALL GAME HAMMALS
WILDLIFE HANAGEHENT UNIT 12

COTTONTAIL

RABBIT

Nearly every square mile of Unit 12 below 8,000 feet elevation
sidered capable of supporting
Both the desert cottontail
the desert cottontail
and the Nuttall's

at least some cottontail

and the Nuttall's

cottontail

being common at lower elevations

cottontail

being less abundant

rabbits yearlong.
should occur;
(5,000-7,000

ft) (Lech1eitner

1969, Armstrong

1972).

No specific

data are available

on cottontail

rabbit densities

it should be kept in mind that the populations

are cyclic, presenting

the appearance

ft)

in the higher elevations

(6,000-8,000

However

is con-

for Colorado.

of these rabbits

of high and low densities

from

year to year.

SNOWSHOE HARE
Snowshoe hares are distributed

throughout

(see accompanying

map).

distribution

the eastern half of Unit 12

This range includes most of the

timbered

land above 8,000 feet in elevation.

Colorado

Division

of Wildlife,

a density

of L4

hares per square mile for the occupied

density

Wildlife

figure yields a year-round

Inventory

population

Systems

(1974) gives

range.

This

of 296 snowshoe hares in

Unit 12.

RED (PINE) SQUIRREL
Armstrong

(1972) and Lech1eitner

be fairly common in the coniferous

(1969) suggest that this squirrel
forests of the higher elevations

12.
No data on abundance

or harvest are available.

should
in Unit

�/

-144LITERATURE
Armstrong,

D. M.

1972.

of Kansas Printing

Colorado Division

CITED

Distribution

of mammals

Service, Lawrence.

of Wildlife, Wildlife

415 pp.

Inventory Systems.

Statewide mammal and bird distribution
from W.C.D., area supervisor,

in Colorado, Univ.

1974.

maps and data compiled

and regional biologist

Located at Denver office, Colo. Div. Wildlife.

input.

Unpublished.

B. D. Trindle
December 1975

�-145DISTRIBUTION AND ABUNDANCE OF S~~L GAME BIRDS
WILDLIFE MANAGE~lliNTUNIT 12

BLUE GROUSE
Blue grouse are distributed
southern

two-thirds

throughout

of Unit 12.

the upper elevations

of the

This includes most of the brush and

timbered areas of the unit which provide suitable habitat for the
grouse.

Colorado Division

(1974) estimated
the occupied

of Wildlife,

Wildlife

Inventory

Systems

blue grouse density at 4.7 birds per square mile for

range.

This estimate resulted

in a year-round

population

of 1,493 birds.

SAGE GROUSE
The distribution
the northwest

of sage grouse in Unit 12 consists of three areas in

corner according

to W.C.O. Roland

Two of the areas have relatively

high densities

(1975 pers. comm.).
of birds.

along Milk Creek and the other is at the base of Monument
Stinking Gulch.

The low density area surrounds

(see accompanying

distribution

map).

Colorado

of Wildlife,

Wildlife

Division

One is located
Butte along

the aforementioned

Inventory Systems

areas

(1974) estimate

the high density areas to be between 13 and 21 birds p~r square mile, and
the low density area at about 1 bird per square mile.

W.C.O. Roland

(1975

pers. corom.) agrees with these estimates.

SHARP-TAILED
Sharp-tailed

GROUSE

grouse are found in two areas within Unit 12.

area (82 square miles) surrounds Thornburgh
round popUlation
Wildlife

of 74 birds according

Inventory

Systems

(1974).

Mountain

The largest

and contains a year-

to Colorado Division

of Wildlife,

The other area (12 square miles)

�-146-

extends along the Williams
Fork of the Williams

Fork River from Waddle Creek to the East

Fork.

This area supports a total population

of

53 birds.

BAND-TAILED
The summer distribution
of Unit 12, generally
accompanying

of band-tailed

pigeons is in the southern half

being within the White River National

distribution

300 band-tai1e1

PIGEON

map).

Forest (see

This area is thought to contain up to

pigeons during the summer months

(C1ait Braun, pers.

comm.).

MOURNING
According

to Colorado Division

(1974) mourning

of Wildlife,

doves are distributed

edge of the unit.
agricultural

DOVE
Wildlife

Inventory Systems

in the northwest

and northcentral

This area takes up most of the lower elevations

lands of Unit 12.

Mourning

and

doves are fairly common in the

unit during the spring and summer months, promptly migrating

with the

first frosts.

Abundance

of mourning

doves on Unit 12 is estimated

at approximately

1,254

birds with a density of 8.1 birds per square mile of occupied range.

WATERFOWL
Waterfowl

can be found everywhere

Suitable

habitat 'includes lakes, rivers, streams, stockponds,

No data on abundance

in Unit 12 where suitable habitat exists.
and marshes.

are available.

PHEASANT
Pheasant

distribution

Unit 12, according

includes the suitable habitat in Axial Basin within

to W.C.O. Roland

(1975 pers. comm.).

The popUlation

is

�-147doing poorly in this area and is thought to be marginal

at best.

TURKEY
W.C.O. Roland
established
mately

(1975 pers. comm.) says that wild turkeys were well

in Unit 12 during the 1960's, with a population

200 birds.

population

When this report was compiled

had either migrated

of approxi-

(Dec. 1975) the turkey

out of the area or had dropped to such

low levels that they had become unobservable.

The turkey's original

distribution

of Sand Creek with the

is centered around the convergence

East Fork of the Williams

Fork.

LITERATURE
Colorado Division
Statewide

Wildlife

Inventory

mammal and bird distribution

from W.C.O.,
Located

of Wildlife,

CITED

area supervisor,

Systems.

1974.

maps and data compiled

and regional biologist

at Denver office, Colorado Division

input.

of Wildlife.

Unpublished.

B. D. Trindle
December 1975

�-148DEER HARVEST, SEASONS ~~ HUNTING PRESSURE
WILDLIFE HANAGEMEh"'TUNIT 12
(From Colorado Big Game Harvest 1956-1974)
YEAR

HUNTING
PRESSURE l/ ~

HARVEST
DOES
FAIVNS TOTAL

HUNT AND SEASON

1956

469

200

89

14

303

·IDH 10/15 - 10/30

1957

972

525

365

103

993

2DH 9/14 - 9/30, In Part
2DH 10/15 - 11/17

1958

750

307

138

54

499

ES 10/15 - 11/2

1959

605

290

114

48

452

ES 10/17 - 11/3

1960

771

376

235

70

693

ES 10/17 - 11/6
1DM 12/1 - 12/15, In Part

1961

924

543

260

100

903

IDM 10/21 - 11/8, In Part
3DA 10/28 - 11/3, In Part
1DM 12/2 - 12/152 In Part

1962

1312

519

340

65

924

IDM 8/18 - 9/3, In Part
ES 10/20 - 11/4

1963

1588

941

593

230

1764

2DM 10/19 - 11/7, In Part
2DMA 10/19 - 11/7, In Part.

1964

1451

485

261

67

813

ES 10/17 - 11/5

1965

1730

478

320

111

909

ES 10/16 - 11/5
1DM 11/25 - 12/52 In Part

°1966

1325

381

179

55

615

ES 10/15 - 10/31

1967

1191

422

214

33

669

ES 10/21 - 11/9

1968

1556

544

261

42

847

ES 10/19 - 11/7
ES 11/27 - 12/82 In Part

1969

1080

307

267

61

635

ES 10/18 - 11/6

1970

1265

502

5

0

507

1DB 10/17 - 11/6

1971

631

384

0

0

384

1DB 10/30 - II/II

1972

942

484

204

20

708

1DB 10/14 - 10/23, In Part
IDA 11/18 - 11/262 In Part

1973

1181

445

212

67

724

1DB 10/13 - 10/23, In Part
IDA 10/11 - 10/23. In Part
IDA 11/17 - 12/2, In Part

1974

616

290

0

0

290

1DB 10/26 - 11/5

(Footnotes on fo11oW1ng page)

�-149-

DEER HARVEST, SEASONS AND HUNTING PRESSURE
WILDLIFE MANAGEMENT UNIT 12

(Continued)

1/
- Based upon combined total resident and non-resident license sales and
expressed in number of licenses, not including archery hunters.
Note:
ES

Hunt symbols explained as follows:
=

Either sex, one deer (one license).

2ES

Either sex, two deer (one license).

lDA

One deer, antlerless

lDB

One deer, buck only (one license).

IDM

One deer multiple,

either sex (2 licenses and 2 deer per individual).

lDMA =

One deer multiple,
per individual) •

one must be antlerless

2DM

Two deer multiple,

either sex (2 licenses and 4 deer per individual).

2DMA

Two deer mUltiple,
per individual).

one must be antlerless

3DA

Three deer, one must be antlerless
on 2nd license).

Note:

The term lDM evolved into 2D, HC or two deer, hunters choice
licenses and 2 deer per individual) and is synonymous.

only (one license).

(2 licenses and 2 deer

(2 licenses and 4 deer

(2 licenses plus 3rd deer coupon

B. D. Trindle
August 1975

(2

�-150PRIMITIVE DEER HARVEST AND SEASONS
WILDLIFE MANAGEHENT UNIT 12
(From Colorado Big Game Harvest 1973-1974)

HARVEST
YEAR
1973 JJ

BUCKS
---

DOES

FAWNS
---

TOTAL

HUNT AND SEASON

4

9

4

17

ES 9/8 - 9/16, In Part

4

0

0

4

lDB 9/14 - 9/22, In Part

1./
1974

!/

Free permits issued to holders of regular big game hunting licenses by
application only.

Forty resident

and non-resident

muzzle-loaders.

1/
Thirty

Note:
ES
lDB

two resident

and non-resident

muzzle-loaders.

Hunt symbols as follows:

=

Either sex, one deer (one license).
One deer, buck only (one license).

B. D. Trindle
Septe.mbe:r1975

�-151ARCHERY DEER HARVEST AND SEASONS
WILDLIFE MANAGEHEt;T~IT 12
(From Colorado Big Game Harvest 1956-1974)
YEAR 1/

BUCKS

DOES

FAWNS

1956 !±/

ES 10/1 - 10/14

1957
1958

Regular Season Only
0

4

0

4

1959
1960

ES 9/1 - 9/30
ES 9/15 - 9/30

2

0

0

2

1961

1962

3/
HUNT AND SEASON -

TOTAL 2:./

ES 9/10 - 9/30
1DM 8/26 - 9/10, In Part
3DA 8/26 - 9/10, In Part
1DM 11/11 - 11/26, In Part
3DA 11/11 - 11/26, In Part

0

2

0

2

1963

ES 8/25 - 9/23
ES 8/17 - 9/8

1964

4

5

0

9

ES 8/15 - 9/13

1965

0

5

0

5

ES 8/21 - 9/12

1966

0

2

0

2

ES 8/20 - 9/18

1967

0

3

0

3

ES 8/19 - 9/17

1968

2

6

0

8

ES8/17

1969

2

0

0

2

lDB 8/16 - 9/14

1970 ~/

6

5

0

11

lOB 8/15 - 8/31
ES 9/1 - 9/20

1971

1

0

0

1

lOB 8/21 - 9/12

1972

11

7

0

18

lOB 8/19 - 8/31
ES 9/1 - 9/20

1973

13

13

0

26

lDB 8/18 - 8/31
ES 9/1 - 9/23

1974

0

0

0

0

lOB 8/17 - 8/31
1DB 9/1 - 9/22

(Footnotes on following page)

- 9/15

�-152ARCHERY DEER HARVEST AND SEASONS
WILDLIFE MANAGEMENT UNIT 12

(Continued)

1/
Year 1956 Free pe~its issued to holders of regular big game hunting
licenses by application only. Years 1957-1960, no issue of special permits; archery hunting allowed to holders of valid deer licenses. Years
1961-1974, separate archery license regulations in effect.

2/
- Does not include harvest by archers during gun seasons.
Inconclusive
harvest data omitted for years 1956, 1957, 1959, 1961, and 1963.

3/
- Bow hunting permitted during all regular rifle seasons with regular
Colorado deer license.

For years 1956--1969 no hunter pressure data available for Unit 12.
and non-resident archery hunters, 1970-74 by year were: 1970 _
65; 1971 - 92; 1972 - 88; 1973 - 105; 1974 - 55.

~/Resident

Note:

Hunt symbols are explained as follows:

ES

=

Either sex, one deer (one license).

IDB

=

One deer, buck only.

IDM
3DA

One deer multiple,

=

either sex (2 licenses and 2 deer per individual).

Three deer, one must be antlerless
on 2nd license.

(2 licenses plus 3rd deer coupon

B. D. Trindle and W. T. McKean
September 1975

�-153TOTAL DEER HARVEST,DATA
"ANALYSIS UNIT 7, 1963-1974, BY SEX !I

HARVEST
YEAR

"BUCKS

--

DOES

FAWNS
--

TOTAL
--

1963

16117

14999

5018

36134

1964

14734

12170

2946

29850

1965

8161

10127

2739

21027

1966

8394

7558

2361

18313

1967

10582

8143

2150

20875

1968

11076

8468

2407

21951

1969

8108

5493

1676

15277

1970

10335

3577

400

14312

1971

8945

0

0

8945

1972

12655

4870

554

18079

1973

9323

3978

642

13943

1974

7380

220

33

7633

1/
DAU Unit 7 incorporates Wildlife Management Units 11, 12, 13, 22, 23, .
24, 25, 26, 31, 32, 33, and 34.

B. D. Trind1e
August 1975

�-154LINEAR REGRESSION OF EITHER SEX DEER SEASONS 1/
WILDLIFE MANAGEMENT UNIT 12

3000

2500

2000

HARVEST

1500

1000 .

500 .

L-

~

500

1000

1500

~

~___~

2000

2500

3000

HUNTING PRESSURE
Correlation

Coefficient

r = 0.9209

Intercept

Slope

230.8780

0.3589

1.1
Either sex season data obtained
Pressure

Section.

from Deer Harvest,

Seasons and Hunting

�-155ELK HARVEST, SEASONS AND HUNTING PRESSURE
WILDLIFE HA.c\lAGEMENTUNIT 12
(From Colorado Big Game Harvest 1956-1974)

YEAR

HUNTING
PRESSURE

1956

HARVEST
CALVES

TOTAL

1/
HUNT A1TD SEASON -

10

151

AO + 150 ESP 10/15 - 10/31

26

2

137

AO + 100 ESP 10/15 - 10/31

179

20

2

201

AO + 50 ESP 10/15 - 11/2

669

179

25

2

206

AO + 100 ESP 10/17 - 11/3

1960

660

238

23

1

262

AO +

1961

874

226

226

AO 10/21 - 11/8

1962

885

202

202

AO 10/20 - 11/4

1963

954

210

210

AO 10/19 - 11/7

1964

1390

341

28

8

377

AO + 50 ESP 10/17 - 11/5

1965

1238

355

24

3

382

AO + 50 C 10/16 - 11/5

1966

1325

365

33

8

399

AO + 75 C 10/15 - 11/3

1967

1525

358

4.0

11

409

AO + 150 C 10/21 - 11/9

1968

1756

446

94

18

558

AO + 180 C 10/19 - 11/7

1969

1569

429

109

15

553

AO + 200 C 10/18 - 11/6

1970

2170

141

34

4

619

AO + 250 C 10/17 - 11/6

1971

1064

286

34

4

324

AO + 200 C 10/16 - 10/25

1972

1006

157

40

2

199

AO £/ + 200 C 10/28 - 11/12

1973

1981

645

160

31

836

AO + 400 C 10/27 - 11/11

1974

2687

616

164

19

799

AO ;- 600 C 10/12 - 10/22

BULLS

--

COWS

519

103

38

1957

554

109

1958

668

1959

50 ESP 10/17 - 11/6

1:../
Antlered

only, four points or better on one side.

~/
The hunt and season column reflects permits issued for Area B. In 1956
Area B consisted of Wildlife Management Units 12, 13 and 26. During the
years 1957-1974 Area B consisted of Units 12 and 13.
(Footnotes

continued

on following page).

�-156ELK HARVEST, SEASONS AND HUNTING PRESSURE (continued)
WILDLIFE MANAGEMENT UNIT 12

Note:

Hunt symbols are explained as follows:

AO
AO + ESP = Antlered only, plus either sex permits.
AO + C

= Antlered only, plus specified number of cow permits.

B. D. Trindle and W. T. McKean
September 1975

�-157ARCHERY ELK HARVEST, SEASONS A1~ HUNTING PRESSURE
WILDLIFE MANAGEHENT UNIT 12
(From Colorado Big Game Harvest 1967-1974)

YEAR

HUNTING
PRESSURE

BULLS

No Data
58

1969

TOTAL

----

3/
1967 1968

HARVEST 1/
cous CALVES

2/
·HUNT AND SEASON -

----

ES 9/2 - 9/17

8

8

0

16

ES 8/17 - 9/15

7

0

0

7

AD 8/16 - 9/14

1970

65

8

3

0

11

AD 8/15 - 8/31
ES 9/1 - 9/20

1971

109

11

0

0

11

AD 8/21

1972

104

0

4

2

6

AD 8/19 - 8/31
ES 9/1 - 9/20

1973

91

9

0

2

11

AD 8/18 - 8/31
ES 9/1 - 9/23

1974

109

0

15

0

15

AD 8/17 - 8/31
ES 9/1 - 9/22

9/12

1/
Includes harvest only during specified archery seasons.
!/Elk bow hunting permitted during all regular seasons with regular Colorado
elk license.
3/
- First statewide elk archery season with archery elk license.
Note:

Hunt symbols explained as follows:

ES = Either sex, one elk (one license).
AD = Antlered only, one elk (one license).
B. D. Trind1e and W. T. McKean
September 1975

�-158-

TOTAL ELK HARVEST DATA ANALYSIS
UNIT 6 (WHITE RIVER PLATEAU) 1963-1972, BY SEX 1/

YEAR

BULLS
-

COWS

CALVES
--

1783

TOTAL
--

377

45

2205

2501

881

120

3502

1993

673

92

2758

1873

942

200

3015

1737

907

188

2827

2184

1418

292

3894

1969

2262

1035

234

3531

1970

2517

801

91

3409

1971

1039

528

57

1624

1972

1800

743

107

2650

1963
1964
1965
1966
1967
1968

Elk DAU 6 includes Wildlife Management Units 12, 13, 23, 24, 25, 26,

33 and 34.

W. T. McKean
September 1975

�-159BLACK BEAR HARVEST AND SEASONS
WILDLIFE HAt'lAGE-1ENT UNIT 12

(From Colorado

1/

Big Game Harvest 1956-1974)

2/ , -3/ , -4/

YEAR

HARVEST -

1956

3R

1957

o

1958

o

1959

o

1960

6R

1961

o

1962

o

4/1 - 9/15; pre and regular deer, elk seasons
ending 11/4

1963

5R

4/1 - 9/15; pre and regular deer, elk seasons
ending 11/7

1964

2R

4/1 - 9/15; pre and regular deer, elk seasons
ending 11/5

1965

o

4/1 - 9/30; pre and regular deer, elk seasons
ending 11/5

1966

o

4/1 - 9/30; pre and regular deer, elk seasons
ending 11/3

1967

o

4/1 - 9/30; regular deer and elk seasons ending
11/9

1968

4S

4/1 - 9/30; regular deer and elk seasons ending
11/7

1969

o

4/1 - 6/30; regular deer and" elk seasons ending
11/6

1970

IS

4/1 - 6/30; regular deer and elk seasons ending
11/6

1971

o

lID1'T AND SEASON -

5/15 - 8/15; pre, post, and regular deer and
elk seasons
4/1 - 11/15; pre, post, and regular deer and
elk seasons
4/1 - 11/15; pre, post, and regular deer and
elk seasons
4/1 - 11/1; pre and regular deer and elk seasons
ending 11/1
4/1 - 9/15; pre and regular deer, elk and
archery seasons ending 11/6
4/1 - 9/15; pre and regular deer and elk seasons
ending 11/8

4/1 - 6/30; regular deer and elk seasons ending
11/11

----------------------------------------------------------------------------

�-160BLACK BEAR HARVEST AND SEASONS
WILDLIFE MANAGEMENT UNIT 12

HARVEST

(Continued)

1../

5:./, ~./,i/

YEAR

----

1972

2R

4/1 - 6/30; regular deer and elk seasons
ending 11/12

1973

2R

4/1 - 6/30; regular deer and elk seasons
ending 11/11

1974

2

4/1 - 6/30; regular deer and elk seasons
ending 11/5

HUNT AND SEASON

:!./
R = Regular big game (deer and/or elk) licenses; S = Bear license for
special spring, summer, or spring-summer seasons; also see annual regulations
for archery hunting.

2/
- From 1956-1959, 1 bear per special bear license and/or 1 bear per bear
coupon on either or both deer and elk licenses; - from 1960-1965, same as
1956-1959, except special bear license variously invalid after September 15
or 30, when bear coupon on either or both deer and elk license covered bag
and possession limits; - 1966, same as 1960-1965, except one bear, hunter's
choice, per person per calendar year; - from 1967 to 1975, bear coupon
removed from regular deer and elk licenses; thus 1 bear, hunter's choice,
per special bear license or sportsman's license per person per calendar
year only during special bear and regular deer and/or elk seasons.

3/

.

.

.

- Up to eight dogs permitted except when bear seasons concurrent with deer
and/or elk seasons.
Beginning in 1970 to date adult bears with one or more
cubs were protected.

4/
- Data taken from Colorado Big Game Harvest and Commission Meeting Minutes.

B. D. Trindle and W. T. McKean
September 1975

�-161-

MOUNTAIN LION HARVEST AND SEASONS
WILDLIFE HANAGE-IEm DiUT 12
YEAR

!I

HARVEST

Since the establishment
1, 1965 NO RECORDED

of the mountain

HUNT AND SE..£\'SON

lion as a game species on July

KILL EXISTS for Wildlife

Hanagement

Unit 12.

11
Bounty of $50 per lion paid within period Hay 7, 1929 - Harch 12, 1965;
protected by statute as of July 1, 1965 when mountain lion license was
established.

B. D. Trind1e
September

1975

�SMALL GAME HARVEST AND HU~7ING PRESSURE
SMALL GAME M}u~AGEMENT UNIT 10, 1968-1973 !/
1969

1970

1971

HARVEST
1972

1973

Aver age

2285

553

452

1387

1318

1231

1204

227

1190

428

2280

877

336

107

869

596

634

1172

930

1885

716

1381

2161

1374

411

402

372

733

899

594

769

1453

1095

924

98

158

NOS

209

444

515

0

129

295

NOS

345

NOS

NOS

NOS

--

NOS

NOS

NOS

NOS

NOS

NOS

--

SPECIES ];/

1968

1969

NUMBER OF HUNTERS
1970 1971
1972

1973

Cottontail
rabbit

397

133

213

196

263

285

248

Snowshoe
hare

126

158

473

299

163

144

Blue grouse

393

497

1380

353

585

Sage grouse

274

520

294

334

Sharp-tailed
grouse

293

352

148

Chukar

NOS i/ NOS

NOS

Average 1/ 1968

I
f-'

o-

N

I

Gambe1's
quail

NOS

NOS

NOS

NOS

NOS

NOS

--

NOS

NOS

NOS

NOS

NOS

NOS

Pheasant

313

207

194

46

102

NOS

172

586

148

507

89

335

NOS

Band-tailed
pigeon

NOS

NOS

NOS

NOS

NOS

NOS

--

NOS

NOS

NOS

NOS

NOS

NOS

Mourning
dove

NOS

276

193

132

149

107

171

NOS

3238

2111

1494

2242

1097

333

2036

1.../
Wildlife Management Unit 12 was wholly contained within former Small Game Management Unit 10, 1968-1973; Small
Game Units were changed in 1974. W.M.U. 12 is now wholly contained in S.G.M.U. 26.
2/
- Game Management Section questions the ability of some hunters to distinguish between species of jackrabbits and
snowshoe hares, and the grouse family, thus adding inconsistency to these data.
3/
- Mean for years data shown.
4/
- NOS means no data or no open seasons.

B. D. Trind1e
September 1975

�-163SMALL GAME HARVEST AND HUNTING PRESSURE };/
SMALL GAME HANAGEMENTUNIT
SPECIES!:../
Cottontail

. NO. ·OF HUNTERS

rabbit

1/

HARVEST

129

156

hare

292

257

Pine squirrel

33

66

Blue grouse

549

1170

Sage grouse

417

1001

162

294

Chukar

NOS !!.../

NOS

Gambe1' s quail

NOS

NOS

Pheasant

NOS

NOS

NOS

NOS

67

262

Marmot

98

262

Badger

31

o

Red fox

128

93

Bobcat

64

30

Raccoon

0

o

Coyote

66

o

Prairie dog

0

o

Jackrabbit

128

923

Crow

33

328

Magpie

97

743

Snowshoe

Sharp-tailed

Band-tailed
Mourning

,

26, 1974

grouse

pigeon

dove

l./

!/
Preliminary

ana.l.ys
i,s by Game Hanagement

Sec tion as of September

1975.

2/
- Game Hanagement Section questions the ability of hunters to distinguish
between species of jackrabbits and snowshoe hares, and the grouse family,
thus adding inconsistency to the above data.
l/Three percent sample size; 33 projected hunters and 33 harvested
one actual hunter reporting a harvest of one.

~/
NOS means no data or no open season.
B. D. Trind1e
September 1975

equals

�-164-

-DUCK AND GOOSE HARVEST A1~ HUNTING PRESSURE
SMALL GAl-iEMA....~AGENENT
mIlT 26 1974 1:/, 2:../

YEAR

DUCKS
EST. NO. HUNTERS

EST. HARVEST

GEESE
EST. NO. HUNTERS

EST. HARVEST

1974

189

1701

54

22

1/
- 1974 is the first year hunting pressure and harvest data available by
Small G&lt;UIleManagement Unit. Wildlife l-ianagement Unit 12 is contained
wholly within

S.G.M.U.

26.

2/
- Prior to 1974 waterfowl kill data available only by county. There are
no identifiable harvest or hunting pressure data for Wildlife Management
Unit 12, 1955-1973.

B. D. Trindle
September 1975

1

I

t

�r

:::&gt;

z

z

f-

W

:2!

w

(.!)

&lt;l:

Z
&lt;l:

:2:
W
~

c::(
(!)

.J
.J

ro
r--.
(j)

co
0'&gt;

&lt;D

�SMALL

GAME MANAGEMENT

1974

UNITS

�-167-

BIG GAME POPULATION DYNAHICS
WILDLIFE

Computer

population

basic management

MANAGENEh"'Till'llT

simulations

tool.

12

of elk and deer herds have become a

Population

data, including the trend of the

herd, are input parameters

of the computer model.

The simulated popula-

tion parameters

the manager's

of the dynamics of

represents

hypothesis

the herd.

The following

graphs (1 and 2) in which the elk and deer herds of wild-

life management

Unit 12 are contained are based on data analysis units

(see Harvest Data).

The four graphs of each herd are explained as

follows:
A =

Birth of young

June 1

B =

Count one

September 20

C =

Count two

December 11

Depopulation

at end -

May 31

B. D. Trindle and W. T. McKean
November 1975

I

.

�-----------.--.-----.---EuC"(bid'A
32000.

ANiiLYsisUlH'Y -E':':bY---·-· - -- --.- .. .. --.

.

-- ..

======;;=======~~If~=~~~~~Q~~1=PJtr~=1~~]J~~\1~~$S~~~~j~~J~==:=====~:;===============;
1
1
1
1·1
1
1
1:-----1-·· .-.-.--~----111

111

1

111

111

111

1

A

B

30000.

c

------

ZnOQ.

o

._----A
25000.

A

.-----------------------~~-------------------------------------------------------------..-----2ItOOQ.

B

-

C

-----:

-

t\

g

e

A

"220OJ.:--u

~

20000.
A
A

laooo.

AB _

_B

"

A--r--

:

-

C

8 .-_-_B-_

8

H

-

\

0
Z

-0.-

-

t:1

I

7

-- -- - ---

0

I

-- ~---..

-- - -

B

c::

~ ..- ~~ f;
~

--:

B
-

B

"

A

8

Q
H

:--Q~
-----:
~ -o

g

A

to

:.~--·0

A

B

A

_

-----

-

:----

; ..

----:

f-'

co

~
~

H
(")

-.-

CJ)

l~ODO.

~c~

--

o__

11t000.
1
1
1--

·12000.

----:-----------

C
0

0

1

1

1

1

. 1

0

1

1

1

1

1

1

1

- ..~

1
1

1

0

0

0

0

0

----------------------------------------------~---------.
1

-+----..--.--~.
-...-1

;

===;~==:==:==:==:===~==:~=====::======::==:=:======:=======~====.==~======z=================.z~ca~===
19&amp;2.0

1972.0

lY&amp;3.0

19&amp;4.0

1960.0

19&amp;&amp;.0

---1974.0

197-&amp;;0--1978.0 --·--198ii-.;·o~---

197Q.0
TIHE

_____

-'-( 4~£&gt;_O~ __AL_ST.A8TJ

_B~ _PQP

~T .. C.NT.9NELC"POp

.A:L.~NJllo(9.L.Q.¥

POP~L£_NgL_._._

.

�.....
::::~~M

DEER (DAl'A.AN,tlLYSIS_WlIL7)

'1

. ••YlILDLIFE
lIu\,NAG;&lt;:V;?::T
m;ITS 11
12 1". ::2:'.' ~3 'J.I 215 26 'l1 '':\2 "'3. and 'V
,= :.I= ' '" ="'= :=,,::::
== ;::.::: ="'=".,::;.;::,.
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z.000OOE. os =",e"':!=:&lt;'lllla"'''''lrft:!u,

.1

All

1

111

1

1

1

----------------------~~-----~-~,----~--------~----------..
B
1

:

1

1

1

1

1

1

1

1

1

'

1

1

1

,-

1

_

::

::

:---------

::

..---=

1.aooooE·OS :---A
B

1;~~~CJE·055----

A
o
v

A

A

A

B

1.40000E.OS

...--=

A
~
A

8

:..
A

B

5----

~
A

A

B

80

A

B

~
_

_

A
A

-----------------------------------~~---------------------------,'1.20000E·OS ~----

-----~~:'
------------

B

B
A

-

._--:

to

H
Q

C

---r-.-.

-

o

0"

\0
--.l..-

)

901)00.

--

:----

,0

D

-

60000.

.•...-:

-0---

=

-.-.

\

=:----

=

._--:

:;,

I-'
C"'l
t!)

~
4-00C;O.

~
~.
~

~

...•-..

... ----

=
20000.

o.

:----

::

-

1
1

:

1

1
1

1
•

1
I.

.:0

1

1

::

_

_

===a=s:=cnn=C=Ba~=======~~:=s~=:==sc=&amp;~:==e==a~c=~=~===.C=~~~e~==~~:=====c=~cs~=~=7.~~:es~~~~=~X:==K==
1964.0

T"'51).O

1962.0

1958.0

1972.0
1970.0

1966.0

1'976--;0
1974.0

r9S0.0
lQ7S·0

r rsr
____________________________________

(_A,,_!~P

AT

S~_~E!. 8"

P_OP AT CNT(~~~.

C~~Qr::.._AI~r-\i!~Q.:..._Q!I

POP A! __[~J_Q.!..)

_

�•

1/
HABITAT RESTORATION PROJECTS - WILDLIFE MANAGEMENT UNIT 12 -

DATE

LOCATION

VEGETATIVE
TYPE

1946

East Fork Williams Fork
T2N, R87, 8SW.

1947

TREATMENT

NO. ACRES
TREATED

Gr2ss

Seeded with Bromus sp., Poa sp.,
Ph1eum pratense.

160

Increase
forage

BLM

East Fork Williams Fork
T2N, R87W.

Grass

Seeded with Bromus carinatus,
Poa pratensis, Ph1eum sp.

160

Increase
forage

BLM

1954

Williams Fork
T2N, R88W.

Grass

Spray with 2,4-D

5

Increase
forage

BLM

1965

Beaver Cr.
T3N, R88, 89W.

Sagebrush

Aerial spray with 2,4-D

520

Control
sagebrush

GF&amp;P

1969

Beaver Cr.
T3N, R89W.

Oakbrush

Aerial spray with 2,4,5-TP

67

Control
oakbrush

GF&amp;P
USFS

Grass

Spray with 2,4-D

1971

Morapos Cr.
T2N, R9lW.

60

PURPOSE OF
TREATMENT AGENCY

Livestock
GF&amp;P
improvement USFS

1/
If details are desired on any of these projects, make requests directly to Mr. Ron Kufe1d, Colorado Division
of Wildlife, Research Center, Fort Collins, Colorado 80522.

I
I-'
"-J

0
I

�GAME SPECIES INTRODUCTIONS - WILDLIFE ~~AGEMENT

UNIT 12

SPECIES

DATE

MALES

FEMALES

TOTAL

RELEASE AREA

Wild Turkey

2-27-68

3

o

3

T5N, R90W, Section 26

Mesa, Colorado

Literature References
Colorado Division of Wildlife, Denver Office,Game

Management Section.

Game species introduction file.
I
I-'

...•.•
I-'

I

�-172-

MANAGEMENT

A.

PROBLEMS

CHECKLIST

- WILDLIFE 11ANAGEMENT UNIT 12

Present
1.

Logging

roads and associated

The primary management
velopment

secondary roads.

problem of Unit 12 to date is the de-

of logging and secondary

roads.

The problem is

there are just too many roads for the area that are unacceptable
for wildlife.

They create unneeded

access and disrupt the

animals daily activities.

B.

Future
1.

Coal development.
Coal development

will create many new problems for the wildlife

within Unit 12.

The associated

and vehicular
building,

activity

in the unit.

Other activities

such as

will restrict the natural movement

and alter their migration

affect wildlife?

How will strip mining

What are the water requirements

of the coal

Wnat time table is involved in revegetation

All these unanswered
considerations

questions,

in the future.

of

patterns and life styles.

Many of the problems are still unknown.

industry?

will increase ht~an

fencing, and mining will also follow coal development.

All these activities
wildlife

development

of spoils?

and more will become management

�..

•

-173- .

, '

.

•
BIG GAME HAM!1AL RESF..ARCHREFERENCES

•
•

WILDLIFE MANAGF.HENT m1IT 12
BLACK BEAR

Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 60 pp.
Gilbert, D. L. 1951. Fur resources study and bear investigations. pp. 1-8.
lA Quart. Prog. Rep., July. Colo. Dept. Game and Fish. 102 pp.
1951. Fur resources study and bear investigations. pp. 1-9.
Quart. Prog. Rep., Oct. Colo. Dept. Game and Fish. 117 pp.

.. .

1953. Fur resources and bear studies. pp. 59-60.
Rep., Jan. Colo. Dept. Game and Fish. pp. 1-141.
1953. Deer-elk investigations.
April. Colo. Dept. Game and Fish.
Gilbert, P. F. 1946. Bear studies.
Dept. Game and Fish. 4 pp.

In

In Quart. Prog.

p. 60. In Quart. Prog. Rep.,
pp. 1-99.

In Quart. Prog. Rep., June.

1948. Deer-elk-bear investigations. pp. 30-34.
Rep., Cct. Colo. Dept. Game and Fish. 80 pp.
1952. Checking station survey. pp. 39-43.
Jan. Colo. Dept. Game and Fish. 60 pp.

Colo.

In Quart. Prog.

In Quart. Prog.,Rep.,

Gilbert, P. F., and G. E. Rogers. 1954. Deer-elk investigations. p. 127.
~ Quart. Prog. Rep., July. Colo. Dept. Game and Fish. pp. 1-183.
liammit, H. C. 1950. Deer-elk-bear investigations.
Proj. W-38-R, Oct. Colo. Dept. Gareeand Fish.

pp. 1-14.
21 pp.

Fed. Aid

Hay, K. G., G. N. Hunter, and L. Robbins. 1961. Big game management in
Colo. Dept.
Colorado 1949-1958. pp. 71-73. In Tech. Bull. No.8.
Game and Fish. 112 pp.
J

Hunter, G. N. 1954. Deer-elk-bear seasons in Colo. 1948-1953.
Colo. Dept. Game and,Fish. 64 pp.
1966., Colo. big game harvest. 1959-1965. pp. 30-32.
41 pp.
Game, Fish and Parks. Spec. Hgtat , Rep. No.!.

pp. 60-61.
Colo. Dept.

HcKean, ll.T. 1965. Colorado long range game species management plans,
1965-1975, for black bear. Colo. Dept. game, Fish and Parks. 6 pp.
Remington, J .• ~~.
Proj. W-6l-R.

..

1955. Fur resources and bear s t ud Lc s , Fed. Aid
Final Rep. April. Colo. Dept. Game and Fish. 85 pp •

..

�-\?94

R., and D. r.. Gilbcr"t.
6"O. A co~t;lbutlon toward a bIbliography
on the black bear. Tech. Publ. No.6. Colo. Dept. Game and fish. 4J pp.

Titner,J.

U. S. Dept. of Agr.

~atcr and related land resources, Colorado River
Hoard and
V.S.D.A. Econ. Res. Service~ Forest Service, and Soil Conserve Service.
I1ny. Denver.
18J pp.
(looseleaf)

;...:

1965.

Bas Ln in Colorado.

Coo p, Study Rep. of Colo. Water Conserve

'-.

.Boyd, R. J.

1970.

Deer-elk

investigations.

pp. 133-213.
In Game Res. Rep.,
pp. 127-286.

July - Part II. Colo. Div. Gawe~ Fish and Parks.

• 1970. Elk of the Hhite River Plateau, Colorado.
---Colo.
Div. G~e,
Fish and Parks. 126 pp.

Tech. Publ. No. 25.
•.

1972. Elk, White River experimental management study. pp. 135-161.
In Game Res. Re:p.,July - Part II. Colo. Div. Wildl. 251 pp.
______

1973. Experimental elk harvest regulations. pp. 45-67.
Res. Rep.) July - Part II. Colo. Div. Wi1dl. 230 pp.

In Game

a

Denney, R. N. 1965.
Colorado long range g2J!lespecies canage~ent plans)
1965-1975, for" elk.- Colo. Dpet. Carae , Fish and Parks. 61 pp.
Cllbert, P. F •. 1953. Deer-elk investigations.
pp. 59-61.
Rep., April.
Colo. Dept. Cane and Fish. pp. 1-99.

~

Quart. Pr og ,

• 1954. Deer-elk investigations.
125-128.
In Quart. Prog. Rep , ,
_--July.
Colo. Dept. G~e and Fish. pp. 1-182.

•.•..

PPd

;:

•

UammitJ H. C. 1951.; Deer-e1k-bear investigations.
Rep •• Jan. Colo. Dept. Cane ~nd Fish. 33 pp •

pp. 16-22.

~

Quart. Prog.

.

K. G., G. N. Hunter, and L. Robbins. 1961. Big game oan2g~~ent in
Colorado, 1949-2958.
pp. 57-70. In Tech~ Publ. No.8.
Colo. Dept.
Came and Fish.
112 pp.
Dunter, G. N. 1954. Deer-elk-bear
Came and Fish.
64 pp.

..
seasons in ~olorado,

1948-1953.

--'-- • 1966.

Colo. big ga~e harvest, 1959-1965.
pp. 9-30.
Game, Fish arid Parks.
Spe.c. !-!got.Rep. No. 1. 41 pp.

Riordan, L. E •. 1948.
ProE. Rep., Jan.
~_A

Colo. Dept.

Deer-elk-bear investigations.
pp. 29-"33.
Colo~ Dept. C~~e and Fish. pp. 1-49.

----~

1948. Deer-e1k-bear investi~ations.
pp. 7-8.
Rep •• April.
Cblo. Dept. Gaoe and Fish. pp. 1-54.
•

~

Colo. Dept.

In Ouar
-'

t•

Quart. p!og.

• 1949. Deer:"elk-bear investig:ltio:1s. 'pp. 39-46.
---Rep.,
July. Colo. Dept. Game and Fish. pp. 1-98.

I~ Quart. Prog.

Rogers, G. E •. 1951. Checking station survey. pp. 32-33.
Rcp •• Jan. Colo. Dept. Game and Fish. 33 pp.

E!. Quart. Frog.

T:i1eston, J. ll. 1962.
1957. pp. 70-78.

VI pp.

A resume of Color.:tdobig g:lme rese3rch project!;, 1939In Te ch ..l'ub L, ~~o."9. Colo. Dept. Caruc and Fish.

�-175-

V. S. Dept. of Ar,r. 1965.

Water ~nd related l~nd resourcc~. Color~co River
Basin in ·Colorado. Coop. Study Rep. of Colo. Water Conserve Uoaruand
V.S.D.A. [con. Res. Service, Forest Service, and ,Soil Conserve Service.
May. Denver. 183 pp. (looseleaf)

Uil1iams, J., et a1. 1952. LocaticnjOd extent of winter ranges. pp. 57-62 •
.!!!. Quart. Prog. Rep., April. Colo. Dept. Game and Fish. pp. 1-62.

MOUNTAIN

../,

tIOl{

Boyd, R. J. 1965. Colorado long range gane species managenent plans,
1965-1975, for predators.
Colo. Dept. G~e, Fish and Parks. 21 pp.
Dixon, K. R., and R. J. Boyd. 1967. Evaluation of the effects of mountain
lion predation. pp , ll;1-165. In Came Res. Rep., July-II. Colo. Dept.
Came, Fish and Parks. pp. 73-309.

u. S. Dept. of Agr.

1965. Water and related land resources, Colorado River
Basin in Colorado.
Coop. Study Rep. of Colo. \-Jate~Conserve Board and
U. S.D.A. Econ •.Res. Serv Lc e , rarest Service, and Soil Coris erv , Service •
.Hay. Denver. 183 pp. (looseleaf)

HUJ.E DEER
o

·Baker, B. D. 1955. Deer-elk investigations.
pp. 149-157.
In Quart. Pr og .
Rep., July - Par~ II. Colo. De?t. Game and Fish. pp. 117-175.
1956. Deer-elk investigations.
pp. 43-64. In Quart. Prog. Rep.,
July - Part I. Colo. Dept. Gane and Fish~ pp. 1-102.
1957. A directory of Colorado big game exclosures concerning their
locations, studies and other gen~ral infor~ation. SFec. Rep., Fed. Aid
Proj. W-38-~,Jan. Colo. Dept. G~e and Fish. 60 pp.
1959. Deer-elk investigations.
pp. 189-249.
Rep., July. Colo. Dept. Ga~e and Fish. 249 pp.

In Quart. Prog.

1960. A detailed study of r2n~e forage by use of fenced exclosures.
pp. 1-10, 41-42. In Quart. Rep., April. Co~o. Dept. Ga~e and Fish. 47 pp.
1961. A directory of Colo. big g~~e exclosures cbncerning program
history, study techniques, 1oc.:1tions,and other general infor::;ation.
Spec. Rep. Fed. Aid Proj. W-IOl-~, Work Plan I, Job No.2.
January.
Colo. Dcp't, GaL:lC and Fish. 51 pp.
1961. Ga~c range investig~tions.
pp. 71-72, 76-112.
July. Colo. Dept. Game and Fish. lq9 pp.
1962. Game rang c investig&lt;1tions.
Colo. Dept. Game and Fish. 96 pp.

pp. 1-39.

~

Tn Quart. Rep.,

Qua r t , Rep., AprIl.

�-176-

e
1962. Deer-elk inve5tig~tions •. pp. 129-i38.
Colo. Dept. Came and fish. pp. 129-249.

______

~

------e·· 1964.

Came range investig~tions. pp. 3-6.
Colo. Dept. Game, Fizh and Parks.. 121 pp.

~

Quart. Rep., July.

•
In Came Res. Rep., Jan.

• 1965. Game range investigations. pp. 129-133. In Game Res. Rep.,
--. July. Colo. Dept. Game, Fish and Parks. 249 pp.
1965 •. The history, function. and cstablish~ent of exclosures in
Colorado big ba~e oanagement.
Colo. Dept. Ga~e, Fish and Parks. Outdoor
Facts No. 18. 3 pp.
1966. Car:Ierange investigations.
pp. 53-67.
July - Part 1. Colo. Dept. Gane, Fish
and Parks.

.

In Game Res. Rep.,
93 pp.

1970. Came range investigations.
pp. 15-58. In Gane Res. Rep. ,
July - Part I. ·Colo. Div. Cane ,
Fish and Parks. 126 pp.
Bartmann, R. }!. 1967. Deer trapping ar.d tagging results at the Little Hills
ExperL~ent Station. Colo. Dept. Ga~e, Fish and Parks. Outdoor Facts
No. 57. 3 pp.
1968.

Playing tag with the l,1f1ite
River deer hcr d , Colo. Outdoors •

. 17 (1) : 20- 22.

•
1968. Results from an 18 year deer tagging progran in northt,estern
Colo. pp. 161-166. 48th Ann. Conf. West. Assoc. State Game and Fish
, Comm ,
724 pp.
1968. }fule deer migration at the Little Hills Game Experinent
Station, Colo. p. 9. 13th Trans. Ann. Su~~er Conf. C.~.P.S., The
Wildlife Society.
~.
1972. Evaluation of the effects of spring-s~~er grazing by deer
on alfalfa. pp. 113-133. In Game Res. Rep., July - Part II. Colo.
Div. of Wildlife. pp. 71-251.
1972. Pi'ceance deer studY-population distribution. pp. 317-337.
Res. Rep .• July - Part III. Colo. Div. of Hildlife.
pp. 252-377.

~

Came

1972. Piceance de~r study-population densit~. pp. 339-344.
Game tes. Rep., July - Part III. Colo. Div. of ~ildlife.
pp. 252-377.
•

In

•

�-177-

1972.

Piceance deer.study-productivity and nortality.
pp. 345-350.
Colo. Div. of Hildlife.
pp. 252-377.

lE. Game Res. Rcp , , July - Part III.

1972. Piceacce deer study-food h~bits. pp. 351-355. In Came Res.
~ept •• July - Part III. Colo. Div. of Wildlife. pp. ~52-377.
1973. Piceance deer study -population density and structure. pp.
243-254. In Game Res. R~p., July - Part II. Colo. Div. Wildl. 230 pp.
1973. Pf.ce.ance deer study - population distribution. pp , 211242. ~ Game Res. Rep., July - Part II. Colo. Div. Wildl. 230 pp •

.

1973. Piceance deer study - productivity and mortality. pp.255261. ~ Game Res. Rep., July - Part II. Colo. Div. Wildl. 230 pp.
-,

1974. Piceance deer study - population distribution. pp. 325-362.
In Game Res. Rep., July - Part II. Colo. Div. Wildl. 223 pp.
~

1974. Piceance deer study - productivity and mortality. pp. 371-380.
Game Res. Rep., July - Part II. Colo. Div. Wildl. 223 pp.

1974. Piceance deer study - population density and structure. pp.
363-370. In G~e Res. Rep., July - Part II. Colo. Div. Wildl. 223 pp.
1975. Piceance deer study - population density and structure. pp.
349-354. In Game Res. Rep., July - Part II. Colo. Div. Wildl. 314 pp.
1975. Piceance deer study - population distribution. pp. 327-349.
In Game Res. Rep., July - Part II. Colo. Div. Wildl. 314 pp.
1975. Piceance deer study - productivity and mortality. pp. 355362. In Game Res. Rep., July - Part II. Colo. Div. Wildl • 314 pp.

•0

Bar traann , R. M., and Y. T. Md~can. 1968. Came r'ang e inve5tiRations.
pp. 167179. ~Came
Res. Rep •• July -. Colo. Div. Came, Fd sh and Parks. 208 pp.
0

______ , and
1969. Came range investigations.
pp. 305-393.
Res. Rcp ,, July. Colo •. Div. Game, Fish and Parks. 393 pp.
Boeker, H. M. 1959. Colorado deer trapping. pp. 52-55.
Sutmner Conf., C.M.P.S.ot The Wildlife Society.

In Game

4th Trans. Ann.

�-178-

£oyd, R. J. ·1961. Study of deer losses OR .Co~orado highways. pp. 75-82.
In Fed Aid Quart. Rep., April (Part l).Colo.
Dept. Game and Fish.
96 pp. (processed).
1962. Study of deer losses on Colorado highways. pp. 93-100.
Fed. Aid Quart. Rep., October. Colo. Dept Game and Fish. 137 pp.
(processed).

In

1964. Study of deer losses on Colorado high•..
'ays. pp , 36-41. In
Fed. Aid Quart. Rep., April (Part 1). Colo. Dept. Game, Fish and Parks~'
69 pp. (processed).
1970. Deer-elk investigations.
pp. 133-213.
July - Part II. Colo. Div. Gace, Fish and Parks.
Burdick, H.

1963.

Deer tags tell a tale.

In Gam~ Res. Rep.,
pp. 127-286.

Colo. Outdoors 12(5):52-53.

Campbell, R. L. 1950.· Little Hills G~e Exper~ent Station. pp. 38-43.
~ Quart. Prog. Rep., July. ·Colo. Dept. Game and Fish. 80 pp.
1951 • Experimental projects at Little Hills Expe r Irnen t Station.
..pp. 58-65. Spec. Rep. The Glenwood Springs Wildl. Conf. 85 pp.
1951. Deer-elk investigations.
pp. 5-11.
Jan. Colo. Dept. Gane and Fish. 33 pp.

In Quart. Prog. Rep.,

1951. Deer-elk investigations.
pp , 50-53.
April. Colo. Dept. Came ana Fish. 64 pp.

In Quart. Prog. Rep.,

1951.' Deer-elk investigations.
pp. 71-77.
July. ~olo. Dept. Game and Fish. 101 pp.

In Quart. Prog. Rep.,

1952.

Deer vs. livestock.

Colorado Outdoors 1(2):20-22.

1954. Dcer-elk investigations.
pp. 149-154.
July. Colo. Dept. Game and rish~ 182 pp.
1956. Deer-elk inves tiga t ions. pp , 65-71.
July. Colo. Dept. Game and Fish. 102 pp. -

In Quart. Prog. Rep.,

Tn Quart. Pr og , Rcp , ,

Campbell, R. L. ~nd G. E. Rogers. 1950. Deer-elk surveys. pp. 20-21.
~ Quart. Prog. Rep., October. Colo. Dcp t. Came and Fish. 21 pp.
t

Carhart, A. lie
19/,0. Deer-elk survey,
Vol.~.
Colo. GaHC and Fish Commission. 19 pp.

Fcd. Aid Proj. \-I-/I-R.

I

4

�.

-179"-

---

• .1940. Deer-elk survey, Colorado.
Commission.
50 pp.

Vol. 5. ·Colo. C~~C and Fish

1940. Deer-elk ~urvey, Colorado.
Game and Fish Corza Lss Lon , 8 pp.

Tables XIV,

Vol. 6.

"XV.

Colo.

•

1941. Deer-elk survey, Colorado. Vol. 5, Part I~ Dcer food requirefllcntsin Colorado.
Colo. Gar.lCand Fish Com...-n.
28 pp.
1943. The Piceance-~mite River deer ,herd. Deer-elk survey Supplemental Rep. Colo. G~e and Fish Co~ission.
20 pp.
Denney, R. N. 1965. Colorado big gane species as possible vectors of some
livestock diseases.
Colo. Game, Fish and Parks Dept. Outdoor Facts
No. 20. 3 pp~
Denney, R. N., et a1. 1965. Game range surveys. pp. 17-23. Colo. Dept.
Game, Fish and Parks. Gawe Research Review, 1964. 35 pp.
Cilbert, P. F. 1948. Deer-elk-bear investigations.
Prog. Rep., April. Colo. Dept. Ga.~e and Fish.

pp. 14-18.

In Quart.

54 pp.

pp. 31-34.
83 pp.

In Quart. Prog.

1948. Deer-e1k-bear investigations.
pp. 30-34.
Rep., October. Colo. Dept. Gallieand Fish. 80 pp.

In Quart. Prog.

1948. Deer-elk-~ear investigations.
Rep., July. Colo. Dept. Ga~e and Fish.

1949. Deer-elk-bear investigations.
pp. 42'-45. In Quart. Prog.
Rep., January. Colo. Dept. Ga~e and Fish. 67 pp. ,
pp , 47-51.
• 1949. Deer-e1k-bear investigations.
---Rep.,
April. Colo. Dept. Ga.':le
and' Fish. 98 pp.

In Quart. Prog.

1949. Deer-elk-bear ir.vestigatior.s. pp. 33-36.
.Rep., July. Colo. Dept. GaQc and Fish. 84 pp •

In Quart. Prog.

1950. Deer-elk-bear investigations.
pp. 41-44.
Rep., January. Colo. Dept. G~e and Fish. 66 pp.

In Quart. Prog.

_____

e·

Rep.
'.

1950. Deer-clk-bear investigations.
pp. 60-64.
Colo. Dept. G~ae and Fish. 101 pp.

1950. Dcer-elk-bcar investigations.
1950. July. Colo. Dept. GaQc nnd Fish.

~

Quart. Prog.

Three ycar Sun. Rep.,·194721 pp.

1952'. De er+c Lk Lnve s t Lga t Lons , pp , 39-43.
January. Colo. D~pt. Gamc and fIsh. 60 pp.

In Quart. Prog, Rep.,

1952! Decr-elk inve~tications.
pp. 41-64, 66-67, 69-82, 83-84.
Quart. Prog , Rcp ,, July. Colo. Dept. Game and Fish. 171 pp •
•

..

'•...

•

In

�-180-

• 1953. Decr-elk 1nvc5tif,atlons. pp., 59-61.
April. Colo. Dept. Game and Fish. 99 pp.

In Quart. Pr o g , Rcp ••

1953. Deer-elk investigations.
pp , 51-63.
July. Colo. Dept. Came and Fish. 129 pp.

In QU::lrt.Prog. Rep.,

____ ~.
1954. Deer-elk investigations.
pp. 129-143.
July. Colo. Dept. Came and Fish. 182 pp. .
1956.
Oct obe r,

Deer-elk investigations.
pp. 1-26.
Colo. Dept. Game and Fish. III pp.

In Quart. Prog. Rcp.~

In Quart. Prog. Rep.,

1957. Dee r+e Lk investigations.
pp. 121-126, 141, 145-146.
Quart. Rep.~ July. Colo. Dept. Carne and Fish. 213 pp.
1958.

The deer's biggest problen.

--_. 1960. ~~erc to hunt-1960.

In

Colorado Outdoors 7(3):22-24.

Colorado Outdoors 9(5):5.

Gi1bert~ P. F. et a1. 1951. Deer-elk investigations.
pp. 109-110. In
.......... Quart. Pr og , Rcp , , October.
Colo. Dept. Gazie and Fish. 117 pp •
.Gilbert, P. F., and G. E. Rogers.
1954. Checking station survey.
128.
Quart. Prog. aep., July. Colo. Dept. Ca~e and Fish.

In

Gilbert, P. F., et al. 1955.
Quart. Frog. Rep.~ July.

pp. 125182 pp.

Deer-elk investigations.
pp. 158-161.
Colo. Dept. Ga~e and Fish. 225 pp.

In

Gilbert, P. F., and R. I•• Campbe lL, 1955. Deer-elk investigations.
pp. 163170. In Quart. Prog. Rep , , July. Colo. Dept. Carne and Fish. 225 pp.
e'

pp. 131-138.
Gilbert, P. F., and J. Harris ~ 1958. Deer-elk investigations.
In Quart. Rep., October.
Colo. Dept. Gzoe and Fish. 166 pp.
--·~8(2):

, and
18-20.

1959.

How far d~ deer travel?

Colorado Outdoors

Gordon, D. F., nnd R. }!. Bartnann.
1971. Evaluation of the ef:ects of
sprine-su::::TIcr
~:razing by deer on alfalfa. pp. 99-112.
In Game Res.
Rep., July - Part II.
Colo. Div. Came, Fish and Parks.-pp.
87-224.
llammit, H. C. l&lt;)~,O. Dccr+e Lk-be ar investigations.
pp , 10-14.
Proj. W-38-R. October.
Colo. Dept. Ca~e and Fish. 21 pp.
1951. n~~r-c1k-bc3r investigations.
pp. 16-22.
W··38-R, .Janu.u-v , Colo. Dept. Gazae and Fish. 33 pp.

Fed. Aid

Fed. Aid Proj.

Ilammf t , H, C •• and P.. F.

pp. 35-37.
Hay,

~

Gilbert.
1950. Dce r+c l.k -bca r investigations.
Quart. Pr og . Rep., July. Colo. Dept. Came and Ff.sh ,

K. C., C. N.' Ilun rcr , and L. Robb Ln s , 1961.
nig g anc ma nng crncn t I n
Colorado, 19,'.')·d&lt;J58.
pp. 71-73. In Tech. Publ. No.8.
Colo. Dept.
Game and 1'1:;h. 112 pr.

�-181\

l!untcr, G. N. 1954.
60.

pp. 22-

Deer-elk-bear ~easong in Colorado, 1948-1953.
Colo. Dept. Game and Fish. 64 pp.

1966. Colorado big ~~le harvest, 1959-1965.
pp. 9-30.
Game, Fish and Parks. Spec. Hent. Rep. No.1.
41 pp •.
Jeep, Y. T., anu L. E. Riordan.
1947. Deer-elk studies.
Rep., January - Part IV. Colo. Dept. Game and Fish.

In

Colo. Dept.

7 pp.

Kuf'e ld , R. C.
1968. Carne r cng c Lnvc s t Lg a t Lon s , In Came Res. Rep , ,
Par t 1. "CoLo, Div. Came , Fish and Parks. pp. 1-121.

1969. G&lt;11:1C
range investigations.
pp.299-304.
July - Part III.
Colo. Div. Came , Fish and Parks.

Frog.

Quart.

July -

Tn Carne Res. Rep , ,
p-p.-249--393.

1970. Ga.~e range investigations.
pp. 59-94. In Gane Res. Rep.,
July - Part I. Colo. Div. Ga2e, Fish and Parks.pp.
1-126.
1971. Inventory of nanipulation projects in Colorado.
pp. 7-15.
In Game Res. Rep., July - Part I. Colo. Div. Came, Fish and Parks.
1-86 •.

pp.

Lauridson, D. C. 194$. Gace and forage experkcnt
Co~ents 10(6):9-10, 22.
Lind, C. 1957. Deer-elk investigations.
Colo. Dept. Ca:ne and Fish. 213 pp.

station.

pp ..147-l5l.

Colo. Conserve

In Quart. Rep., July.

pp. 1-23. In Quart. Rep.,
)kKean, W. T. 1958. Deer-elk investig2tions.
\. . , July - Part 1. Colo. Dept. Gace and Fish. pp. 1-103.
1959. G&lt;l1.!lC
range investigations.
pp. 193-212.
"July. Colo. Dept. GaQc and Fish. 237 ppo
___

1960. Ga~e rangc investigations.
pp , 3-23.
October. Colo. Dept. Carie ana ,"ish. 103 pp.
0.

In Quart. Re p ,,

In Quart. Rcp , ,

1965 •. Pellet group deposition rates for captive deer.
Carae, F~sh and Parks. Outdoor Facts ~!o. 24.
3 pp.
. -July.

1967. Game range investigations ...pp. 155-165.
Colo. Dept. Caric , Fish and Parks. 203 pp.

Colo. Dept.

In Came Res. Rep.,

Res. Rep.)
1970. Gili~erange investigations.
pp. 7-9. In Cc:lr:c
July. Colo. Div. Ga~e, Fish and Parks. 126 pp.
HcKean, P. 1'., and H. E. Burdick. 1963. Carne range investiGations. pp. 89115 • .!E. C3C1C Res. Rcp ,, January. Colo. Dept. Came and Fish. 115 pp.
___

___

, and

196/1•

Itcs , Rcp , ,

January.

, nnd
Res. Rcp , ,

July.

Gar:1Cr ang e Lnvo s t Iga t i ons . pp. 27-67.
In Came
Colo. Dcpt. Camc , Fish and Park". 121 pp.

1965. Came r an gc Lnvc s t Lga t Lcn s , pr.1/,7-189.
Co10. Dept. Camc , Fbh and Parks. 2/,9 ppo

In Game

.\

�-182-

McKean, \-1. T.,

and R. M. Ba r tmann , 19fi6 •. Su rv Iva I or mountain maho aany
t ransp l an t s on t~o s J.t cs , Little Hill!; Expc r Imcn t Station,
Rio IHal1co
County, Colorado 1965. Colo. Dept. Game, Fish and Parks. Outdoor Facts
no. 42.. 3 pp.

-Res.

_

~ llnd
1969. Gar.1C range invcstigattons.
pp. 253-260 •. In Game
Rep •• .Ju l.y , Colo. Div. Game, Fish and Parks. 393 pp •

1971. Deer-livestock relations on a pinon-juniper range
in northwestern Colorado.
Final Report. Colo. Div. Game, Fish and Parks •
. 132 pp. (processed) •
-'.

.••...--:-'• and

McKean, \-1.T., and D. G. Smith. 1965. Game range investigations.
pp. 423471. In Game Res. Rep.~ July. Colo. Dept. Game, Fish and Parks. 546pp.
• • and
1966. Game range investigations, p~. 5-49.
--"i~ep.,
J-u-l-y-.
- Colo. Dept.Game,
Fish and Parks. 93 pp.

In Game Res.

Hcl;ean, \-1. T., et a1. 1968. Game range investigations.
pp , 155-165.
Game Res. Rep., July. Colo. Div. Game, Fish and Parks. 208 pp.
Medin, D. E. 1958. Deer-elk investigations.
pp. 63-68.
January •. Colo. Dept. GaQe a~d Fish. 131 pp.

In

In Job Compo Rep.,

______ " 1960. Physical site factors influencing annual production of true
mountain mahogany, Cercocar~us Lonta~us.
Ecology 41(1):454-460.

1960. References on ~ethods of measuring production and utilization
range and pasture forage. Colo. Dept. Came and Fish Tech. Publ. No.
6. 43 pp.

of

.

,
.

.

1965. Colorado long range gane species ~nage~ent
for deer • Colo. Dept. Game, Fish and Parks. 15 pp.

Hustard, E. U., Jr. 1958. Deer-elk investigations,
April. Colo. Dept. Game and Fish. 113 pp.

pp ,

plans, 1965-1975,

9-34. l!!. Quart. Rep.,

• 1958. Browse differences in ex?erioental pastures under controlled
---stocking
yith cattle, sheep, and cule deer in northwestern Colorado.

M. S. Thesis.

Colo. State Univ., Ft. Collins.

105 pp.

1959. Study of browse reproduction in r~lation to controlled grazing
experimental pastures.
In Quart. Prog. Rep , , January. Colo. Dept.
Game and Fish. 41 pp.
in

.·Myers, G. T. 1969. An investigation of deer-auto accidents. pp. 147-178•
In Fed. Aid Ga~c Res. Rep., July - Part 2•. Colo. Div. Game, Fish and
(processed) •
Parks.

1970 .. An investigation of deer-auto accidents. pp. 403-437. In
Fed. Aid Game Res. Rep., July - Part 3. Colo. Div. Game, Fish and
Parks.
(processed).
Neil, P. H.

1973. Winter mortality of mule deer in Par~chute Creek Valley,
Garfield County, Co lo rndo . Tn The Co l.o
ny cnv Lr onmcn t a l s t ud y , Par achu t c
Creek, Garfield County, Color~do. Prepared for Colony Development
Opcr:ltion by Thorne EcoloCic:l1 In:.;titute,Boulder, Co!orndo. '\ucust
1973. Part III.

�-183-

•

Niclt&gt;on, J. L.
1966.
So11 su rvcv of cxpc r Irncrrt a I pas t ur e s , Little
l!ills
Expcr Imcn t St .•c Io n ,
Report
by U. S. Soil Con c c rv , Service
to Colo.
(;a~:jc, Fish and Pa r ks ,
Unpublished c o.amunLca t Lon , 1,2 pp.

Came
Dept.

Paur , t. F.

1972. }rOVe:;Jcnts of mule deer in the Parachute Creek ba s Ln,
uintcr
1971-72.
In The Colony cnvt r ormcnt al ts cudy , Parachute Creek,
Garfield County, (;olor.:1co.
Pr epa r cd for Colony Dcve Lopocn t Opcr a t Lon
• by Thorne Ecological
Institutc,
Boulder, Colorado.
August, 1973.
Part III.
"
.'.

Pojar,- T. }~. 1971.
Came Res.
"

Deer-auto
accident
invcstigations.
pp , 227-318.
In
Rcp , , July - Part III.
Colo. Div. Came , Fish and Parks.

pp. 227-366.

.

____

1971.
Eva Lua t Lon of devices
pp. 333-340.
In Ga~e Res. Rc?,
fish and Parks-.- pp. 227-366.

to prevent deer-auto
accidents.
July - Part III.
Colo. Div. Game,

1971.
Research - 1971 deer-high~ay
lighting
project
pp. 355-366"
In Cane Res. Rep , , July - Part III.
Colo.
1'1.5h and Parks-.- pp. 227··366.

-

1972.

report.
Div. Came ,

Z-ronitor potentially
c r Lt i ca L, deer-vehicle
accident
s t a t cv..•.
ide.
p p , 2G9-275.
I:1 Cane Res. Rcp , , July - Part III.
Div. of hilcilife.
pp. 252-377.

__

0

1972.

areas
Colo.

Effect

of lighted
deer crossing
signs on nu:nber of deer
pp. 300-304.
In Gace Res. Rep., July - Part III.
Co10. Div. of ~";ildlife.
pp. 252-377:

killed by vehicles.

An eV31uation of deer-proof fence length required to prevent
novencnt s on or across high speed h i ghvavs , pp. 305-310.
In Carae
Rep., July.Part III. Colo. Div. of Wildlife.
pp. 252-377.

1972.
deer
\Res.
Reed,

D.

r.

1971~
Investigation
of orre-way deer structures.
pp. 321-330 .
.!.!! Came Res. Rep , , July - Part III.
Colo. Div. Carae , Fish and Parks.
pp. 227-366.

19n~
July

- Part

Deer underpass
evaluation.
pp. 343'-353." In Ga8e Res. Rep.,
III. Colo. Div. Game, Fis,h and Pa r ks , pp. 227-366.

1972. Investigation
of one-way deer structures.
pp.
Game Res. Rep., July - Part III. Colo. Div. of Wildlife.
___

278-282.
In
pp. 252-377.

" i972.
Deer underpass
evaluation.
pp , 285-290.
In Game Res. Rcp , ,
July - Part III. Colo. Div. of Wildlife.
pp. 252-377.
1972.
Ef f cc t s of h I ghwa y light ing on nurnbc r of deer killed
by
vehicles.
pp. 292-298.
In Ga~c Res. Rep., July - Part III.
Colo.
Div. of Wildlife.
pp. 252-377.
1972.
Effects
of 8-foot fence &lt;lngle in diverting deer from their
csl"bUshcd
direction
of movcmcnt , pr.
312-31,/ r , ~
G:lme Res , Rcp , ,
July - Part III.
Colo. Div. of WIldlife.
pp. 252-377.

�-184-

•• E.

n1ord~n"I

Station.

A pro poscd cxpc rtrncnt at Little Hills Carne Expcr Iracrrt
Col~. Cons. Comments 10(1):20-21.
191.7.

1948. Deer-elk-bear invcstigations.
pp. 1~-30.
Rep., January. Colo. Dept. CaQc and Fish. 49 pp •
.•••
_.~_

19ti3.

Decr-elk--be::ll"

Rep., October.

pp.

Lnve s t Lg a t i on s ,

Colo. Dept. Game and Fish.

~

111-!}6.

In Quar t , 1':(08·

'ao pp.

Deer-elk-bear investi~ati,ons. 'pp. 36-40.
Rep., January. Colo. Dept. Game and Fish. 67 pp.
1949.

_______

1949. Deer-elk-bear investibations.
Rep., April. Colo. Dept. Game and Fish.

0_____
Rep.,

Quart. Prog ,

In Quart. Prog.

In Quart.

1'1" 74-82.
97 pp.

1949. Deer-elk-bear investiz:tt Ions , pp. 86-90.
October. Colo. Dept.Game
and Ffsh • lLd. pp.

Prog.

In Quart.' PrC?g.

1956.

Sel~ctivity and utilization of tllree key species of range
cattle, sheep and declo in '.,'estern CoLorado , }1.S. Thesis.
Colo. State Univ., Ft. Collins. 196 pp.
f orag e by

1956. The results of the Little Eills expcrbcntal
range research
----_.,
protran. 1st Trans. Ann. Sirtmer Conf. C.:-r.P.S., The \-lildl. Soc. 13 pp.

---

1957. Differences in ran~e vegetation
reSUlting fro~ grazine by
deer, cattle,
and sheep. Proe. Soc. A~er. Foresters. 1957:147--151.
1957a.
So~e results of a 10-ye2r study of deer-livestock con:petition for range f orag e , Proc. 10th Ann. Heeting Arae r , Soc. Range r!g!:lt.
,Ill pp.
(m.iraco , )

,Riordan, L. E., and R. L. Caopbell.
Outdoors 5(1):1-6.

1956.

Harmony on the range.

Colorado

Robinette, W. L., et al. 1957. Notes cn tooth develop~ent and wear for
Rocky Hountain t;lule deer.
J. vn,n , !-!g:;lt. 21(2);134-153.
Rogers, G. E.

1948.

Deer-elk-bear investigations.
Colo. Dept. Gane and Fish.

Frog. Rcip., April.

.

1948. Deer-elk-bear investigations.
Rep., July. Colo. Dept. Game and Fish.
19/,8.

pp. 8-9.
54 pp •

pp. 24-29.
83 pp.

In Quart.

Prog.

invest i!j3 tions. pp. 27-30.
Colo. Dept. Ga~e and Fish. 80 pp.

In Quart.

Prog ,

~

Quart.

Prog.

P-rog.

Deer-elk-bear

Rep., October.

In Quart.

1949. Deer-e1k-bear investir,ations. pp. 40-42.
Rep •• January. Colo. Dcp t Cane and Fish. 67 PP&gt;
v

R~p't

1949. Deer-elk-bear invc~tir3tions.
April. Colo. Dept. Game and fish.

pp. 41-47.
97 pp.

Tn Quart.

1949. Decr-clk~bear inv~~ti~~tinns.
JUly. Colo. Dcp t , Cnrnc ;1I1U Fbh.

pp. 26-29.
8f, pp.

~

Rcp , J

Quart. Pr og ,

�-185-

1951. Checking station survey. pp. 32-33.
January.
Colo. Dept. Game and Fish. 33 pp.

235.

..

~

Quart. Prog. Rep.,

1952. Ca~e-livetock forage competition study in Colorado. P? 22832nd Ann. Conf. West. Assoc. State Game and Fish Comm. 252 pp.
1953.

Function and operation of big game check stations in Colorado.

.r, Wi1dl. Hgtat;
, 17(3):256-267.

-'0

1965. Sex and age ratios of deer harvested under a hunters-choice,
multiple-license hunting season. Colo. Dept. Game, Fish and Parks.
Outdoor Facts Xo. 34. 2 pp.
Rogers, G., O. Julander, and W. L. Robinette. 1958. Pellet-group counts for
deer census and range-use index. J. Wildl. Hgrnt. 22(2):193-199.
Sedgley, E. F., and H. M. Boeker.
C9nserv. 26(8):177-178.

1961.

1,500 cattle and 100,000 deer.

Soil

Shepherd, R. OR., et ale 1966. Game range investigations.
pp. 91-92. In
Game Res. Rep., July - Part I. Colo. Dept. Game, Fish and Parks.-pp. 1-93~
.
______ , et ale 1966. Game range surveys. pp. 14-18. Colo. Dept. Game,
Fish and Parks. GaQe Research Review 1965. 48 pp.

I

• 1967. Game range surveys. pp. 26-31.
PaI;ks. GaQe Research Reveiw 1966. 46 pp.

Colo. Dept. Game, Fish and.

• 1968. Game range surveys. pp • 25-28.
Parks. Game ReSearch Review 1967. 44 yp.

Colo. Dept. Game, Fish and.

:

Smith, D. G. 1958. Deer-elk investigations. 'pp. 139-140, 157-159, 162.
~ Quart. Rep., October •. Colo. Dept. Game and Fish. 166 pp.
• 1964. Factors affecting the response of rubber rabbitbrush to
---2,3-D.
H.S. Thesis. Colo. State Univ., Ft. Collins. 87 pp.
1965. Controlling r~bber rabbitbrush with 2,4-D.
Fish and Parks, Outdoor Fa~ts No. 19. 2 pp.

Colo. Dept. Gane,

0

Stephens, L. H. 1971. Bro •...
se use by oule deer and livestock.
Colo. State Univ., Ft. Collins. 37 pp ,
o

0

S\O;ope,II. H., et al.
Fish and Parks.

1969. Gane range surveys.
Game Research Review 1968.

1970. Game range surveys. pp. 27-29.
Parks. Game Research Rev Lew 1969. 35 pp.

H.S. Thesis.

0

pp. 18-23.
36 pp.

Colo. Div. Game,

Colo. Div. Game, Fish and
•

Tilcston, J. ~-1. 1962. A r csume of Colorado big game research projects,
'1939-1957. pp. 116-65. In Tech. Publ. No.9.
Colo. Dept. Game and
J.'ish.
81 pp •
• 00

0

�-186-

.

V. S. Dept. of ACr• 1965. Uatcr and related land resources, Color~do River
Basin in Color~do. Coop. Study Rep. of Colo. Water Con:.crv. Bo~rd and
U.S.D.A. Econ. Res. Sc~'icc. Forest Service) and Soil Conserve Service.
i-L3Y., 183 pp. (looseleaf)
~l
"

t!illi&lt;lm~, J. r.
Deer-elk Investi~3tlons. pp. 68-71.
Rep., July. 1951.
~
Colo. Dept. Ga~c and Fish. 102 pp.

1956.
----"
.January.

Deer-elk investigations.
pp. 1-10.
Colo. Dept. G~~e and Fish. 119 pp •

~

•
Quart. Prog.

Quart. Prog. Rep~,

Yillia~s, J., et a1. 1952. Location and extent of vintcr ranges. .: pp. 57-62.
]A Quart. Prog. Rep., April. Colo. Dept. G2nC and Fish. 62 pp •
•

•
•

-.

•

•

,

-.

.. ,

.

",

�-187-

SMALL GAHE H;\..M:-L\L RESf~\RCH REFERENCES

• r

WILDL IFE HANAGP.-1EN! UNIT 12

•

COTTmITAIL RABBIT
Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies.
pp. 1-31. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. 60 pp.
Sand for t , W. W.

July.

1951. Carae bird surveys. pp. 45-47.
Colo. Dept. Game and Fish. 102 pp.

In Quart. Prog. Rep , ,

Shepherd, H. R. 1965a. Colorado long range gzme species ~nagement plans,
1965-1975, for cottontail rabbits. Colo. Dept. Game, Fish and Parks.
12 pp.
U. S. Dept. of Agr. 1965. Water and related land resources, Colorado River
Basin of Colorado. Coop. Study Rep. of Colo. t,'aterConserv. Board and
V.S.D.A. Econ. Res, Service, Forest Service, and Soil Conserv. Service.
May. Denver. 183 pp. (looseleaf)

.:

SNmiSHOE HARE
Dolbeer, R. A. 1972. Populatio~ dynamics of th~ snowshoe hare in Colorado.
Ph.D. Thesis. Colo. State Un:i.v.,Ft. Collins. 193 pp.

,..Porter, K. 1959. Effects of subalpine ti~ber cutting on ~~ld1ife in Colorado •
. In Quart. Prog. Rep., Jan. Colo. Dept. Game and Fish. pp. 151-185.
Shepherd, H. R. 1965b. Colorado long range game species management plans,
1965-1975, for snovshoe hare. Colo. Dept. Game, Fish and Parks. 11 pp.
U. S. Dept. of Agr. 1965. Water and related land resources, Colorado River
Basin, Colorado, C00p. Study Rep. of Colo. t~ater Conserv. Boa~d and
U.S.D.A. Econ. Res. Service, Forest Service, and Soil Conserv.. Service.
Hay. J?enver. 183 pp. (lcoseleaf)

'-

..
.'

..

�...

'-188~

C/.HE BIRD RESE.AJ.~CHREFERENCES

•
. BAh1&gt;-TAILED

Braun, C. E.
___

PIGEON

1970.

The band-tailed pigeon.

Colorado Outdoors 19(5):26-29.

• 1970. Higratory bird Lnve s t Lgc t Lons . pp , 151-171. In Game Res.
Rcp., October. Colo. Div. GaI:.e,Fish and Parks. 171 pp.
1971. Band-tailed piEcon investigations. pp. 209-236. In Game
Res. Rep., October. Colo. Div. Ga=e, Fish and Parks. 236 pp.
1972. Band-tailed pigeon investigations.
Res. Rep., October. Colo. Div. of Wil-life.

pp. 123-14l.
168 pp.

In Game

Braun, C. E., and D. E. Benson. 1972. Band-tailed pigeon investigationsbreeding and nesting chronology studies. pp. 159-168. In Game Res •.
Rep., October. Colo. Div. of hi1dlife. 168 pp.
Grieb, J. R. 1965. Colorado long range ga::lespecies management plans,
1965-1975, for migratory birds. pp. 33-36. Colo. Dept. Game, Fish
and Parks. 36 pp.
lIeff,

J. A., and J. C. Culbreath. 1946.. Co1oracio band-tailed pigeon •.
pp. 1-24. Fed. Aid Proj. '.-R.
Colo. Dep t s Carae and Fish. 24 pp •

U. S. Dap t , of Agr.
1965. '~ater and related land resources, Colorado River
Basin, Colorado. Coop. Study of Colo. \o!aterConserve Board and U.S.D.A.
Econ. Res. Service, Forest Service, and Soil Conserve Service. }~y.
Denver. 183 pp. (loosel~af)
~~ite, J. A. 1972. Plumage studies of band-tailed pigeons. pp. 143-158.
In Game Res. Rep., October. Colo. Div. of Wildlife. 168 pp.

BLUE C:::'OUSE
Aldrich, J. W., and A. J. Duval. 1955. Distribution of American gallinaceous
. game birds. V. S. Fish and Hildl. Circ. 34. 30 pp ,
Anderson, A. E. ·1959. Deer-elk investigations. pp. 118-128.
Rep ; , January. Colo. Dept. Caraeand Fish. 128 pp.

In Quart.

1960. Effects of sagebrush eradication by chemical means on deer
and related wildlife. Job Coopl. Rep., July. Colo. Dept. Game and
Fish. 72 pp.
~:
1960. Distribution and abundance indices of selected biota in
western Colorado before and two months after 2,4-D application. M.S.
Thesis. Colo. State Univ., Fort Collins. 130 pp.

�-189-

---

• )9G6.

.

An 1nve~tf~3tion

of 2,4-D ~ppli~~tion' to ~~~ebrush within a
mount aIn shrub-conifer
complex,
Unco(;",p;1I1r,rc.:
Na t Lo na L Forest.
In Came
Res. Rcp , , July - Pa r t; III.
Colo. Dcp t , Camc, Fish and Parks .-pp.
345-

420.
• 1969. 2,4-D, sagcbrJsh and mule deer-cattle use in upper winter
---range.
Spec. Rep. No. 21. July. Colo. Div. Game, Fi~h and Parks.
21 pp.
Myers, G. T. 1965. Evalu3tion of a combined qlue grouse-wild turkey hunting.
aeason , 'Outdoor Facts No. 27. Colo. Dept. Ga:ne, Fish and Parks. 1 p.
Rogers, G.E.
Part I.

1962. Gane bird survey. pp. 121-l3l.
Colo. Dept. GaLle and Fish. pp. 1-160.

In Quart. Rep., July -

•
1963. Gane bird survey. pp. 157-168, 191. In Gane Res. Rep.,
-----c)ctober - Part II. Colo. Dept. Game, Fish and Parks. pp. 109-241.

1963. Blue "grouse census and harvest in the United States and
Canada.

J. ~ildl. l·rg::lt. 27 (4); 579-585.

1965. Colorado long ra~ge gawe species managecent plans, 19651975, for blue grouse. pp. 1-3. Colo. Dept. Ga~e, Fish and Parks.
16 pp.
1968. The blue grouse in Co1oraQo.
Parks. Tech. Publ. No. 21. 63 pp. ~

Colo. Dept. qame, .Fish and

'S311dfort,\1. H. 1950. Game bird surveys. p. 37.
" ~ October. Colo. Dept. Cazie and Fdsh , 4/. pp.

Fed. Aid Proj. W-37-R,

'-

Steinhoff, H. w. 1956. The dusky grouse and its ecology in Colorado.
Ph.D. Thesis. Syracuse Vnivo 173 pp.
1958. A rating scale for blue grouse ih Colorado.
Amer. For., Wildlife So~. pp. 133-133.

Proc. Soc.

U. S. Dept. of Agr.
1965. ~ater and related land resources. Colorado
River Easin in Co Lo rado , Coop. Study Rep. of Colo. ~':ater Conserv.
Board and U.S.D .•
\. Econ , Res. Service, Forest Service, and Soil Conserve
Service. }~ay Denver. 183 pp. (looseleaf)
0

CHUY-AR
Aldrich, J. H., and A. J. Duvall. 1955. Distribution of Anc r i can ga l l Lnac eous
garce birds. U.S.D.1. .Fish and \!ildl. Sc r ,, Circ. No , 34. 23 pp.
Bartnann, R. M. 1963. Came bird surveys. pp. 33-40, 45-69. In Carne Res.
Rep., October ~ Part I. Colo. Dept. Cane~ Fish and Parks. pp. 1-103.

1964. Cane bird ~urvcy.
Pa r t II.

pp. 151-178. In C.Jme Rcs. Rep., April Colo. Dcp t , Camc , Fi~h and Parks. pp. 71-233.

�-190- .

Evans, R. L., and Y. U. S~ndrort.
1957. Came bird survey. pp. 55-67.
Quart. Rep., October.
Colo. Dept. Came and Fish. 116 pp.

In
-

Hurd, C. L., and W. Y. Sandfort.
Quart. Prog. Rcp.~ October.

In

1955. Carne bird survey. pp. 85-88.
Colo. Dept. Game and fish. lO~ pp.

, and
1956. Ca~e bird survey. pp. 75-81.
---Rep.,
April •. Colo. Dept. GaQc and Fish. 101 pp.

In Quart. Prog.

l~nning, D. D. 1962. Game bird survey. pp. 9-16, 25-31. In Quart. Rep.,.
J.uly - Part 1.
Colo. Dept. Came and Fish. pp. 1-160.
Hiller, J. 1959. Came bird survey. pp . 99-107, 115-124.
October.
Colo. Dept. Ga~e and Fish. 124 pp.

~

Rep.,

Quart.

l111lcr, ~T., and H. Y. Sandf'o r t , 1959. Carre bird survey. pp. 67-73.
Quart. R2p., January.
Colo. Dept. G~~e and Fish •. 128 pp.
Sandfort, H. U. 1950. G2l:JCbird survey. pp , 51-52.
A~ril.
Colo. Dept. Ga~e arid Fish. 101 pp.
1950. Ga~e bird survey. pp. 37-39.
Colo. Dept. Cane and Fish. 44 pp.

In

In Quart. Prog. Rep, ,

Fed. Aid Proj. W-37-R, October.

1951. Chukar census.
pp. 22-31. Spec. Rep., Glenwood Springs
Wildl. Coni., January 16-19. Colo. Dept •.Gane and Fish. 85 pp.
1951. Gane bird survey. pp • 37- a.
, Colo. Dept. Ga;::eand Fish. 102 pp.

In.Quart. Prog. Rep. , July.

1951. Carne bird survey. pp. 80-82.
Colo. Dept. Ga-we and Fish. 117 pp.

In Quart. Prog. Rep. , October.

1952. Gat::ebird survey. pp. 40-56.
Colo. Dept. Ga;::eand Fish. 62 pp.

In Quart. Prog. Rep. , April.

1952~ Cane bird su~vey. pp. 93-95.
Colo. Dept. Ga~e and Fish. 105 pp.

In Quart. Prog. Rep., October.

1953. Came bird survey. pp. 81-89.
Colo. Dept. Gaoe and Fish. 99 pp.

In Quart. Prcg. Rep., April.

l

1953. Carne bird survey. p , 107.
Colo. Dept. Gaoc and Fish. 129 pp.

In Quart. Prog. Rcp , , July.

195~. GaQC bird survey. pp. 13-33.
Colo. Dcpt. Gane and Fish. 79 pp.

In Quart. Prog. Rcp , , April.

1954. Came bird survey. pp. 67-72.
Colo. Dept. Ga~e and Fish. 182 pp.

12::. Quart. Prog. Rep. , July •.

1955. Came bird survey. pp. :1.7-18.
Colorado Dept. Game and Fish. 94 l' p ,

~

Quart. ProC. Rep., January.

1955. Cnmc- b Lr d !;urvcy. pp. 82-811. ~ QU3rt.
Par t 1. Colo. Dept. Carne and Fir-h.
PP&gt; 1-115.

Pr og , Rcp , ,

July-

�-191-

.

---

• 1957. Came bird survey. pp. 29-38, ~5-51. In Quart. Prog. Rep.,
July - Part I. Colo. Dept. Came and Fish. pp. 1-119.

b Quart. Rep , , July - Part II.

Carie bird survey.
pp. 129-131,.
Dept. Carae and Fish. pp. 105-18L

• '1953.

-----Colo.

• 1959. C~~e bird survey. pp. 53-65.
-----C~10. Dept. Gc1~C and Fish. 128 pp.

1n Quart. Rep., Janu~ry.

• 1959. Ca::;e bird survey. pp • 33-36. • In Quart. Rep. , April - Part I.
Colo. Dept. Gane and Fish. pp. 1-116.

1960.

Cane bird survey.
Dept. Gnrae and Fish.
74 pp.
e

___

Colo.
Colo.

Game and Fish.' 161 pp.

1960.

---"

l!!. Quart. Rep.', January.

.1960. Carie bird survey .. pp. 91-9!,. l!!. Quart. Rep., April.

---Pept.
I •

pp. 49-54.

Your chances with the chukar.

Colorado Outdoors

9(5):26-29.

1961. Cane bird surveys. pp. 119-126, 139-155. In Quart. Rep.,
April - Part II. Colo. Dept. C~e and Fish. pp. 97-203.
0

1962.
Part I.

Cane bird survey. pp. 17-23, 33-44.
Colo. Dept. Game and Fish. pp. 1-160.

In Quart. Rep., July

-~-

• 1965. Colorado lcng range gane species manag ccent; plans, 1965-1975)
for chukar part.ricige. Colo. Dept. CaGle, Fish and Parks. 18 pp.

--- .19670

A decade of chukar hunting.

Colorado Outdoors 16(6):20-23.

Sandfort, l-7.l-l.,and D. l\olt:ing. 1952. Ga~e bird survey. pp. 89-91.
Quart. Prog. Rep., October.
Colo. Dept. 9ame and Fish. 105 pp.
Sandf or t , 1-7. W., et ale
1962. Game bird surveys.
July. Colo. Dept. Case and Fish. 159 pp.

1963.
October.

Tully, R.

pp. 9-44.

In

In Quart. Rep ,,

GaBe bird surveys.
pp. 33-44, 63-70. In Quart. ~ep.,
Colo. Dept. Game, Fish and Parks. 107 pp.

1965.

Colorado Outdoors 14(6):16.

Cane birds for 1965.

U e , S. Dept. of ·Agr.
1966. IJater and related land resources, l~'hiteRiver
Basin in Col.orado , Co op . Study Rc p of Colo. !-!aterConserve Board and
U.S.D.A. Econ. Res, Service, Forest Service and Soil Conserve Service.
November.
Denver.
92 pp. (looseleaf)
'~'hite,C., and J. E. l-lillians. 1960. Escalante.
pp. 21-23.' Colorado
•
Outdoors, 1~0. 5. Sept.-Oct •. Colo. Dept. Came and Fish. 32 pp •

-

..

�C,..ttI~ELI S Oil" It

~192-

t~ndfort, W. w. 1950.
July.
Colo. Dept.
•...
·~olo.

..

.

C~mc bird surveys. pp. 17-21.
Came and Fish.
80 pp.

1950. Came bird survey. pp. 37-39 •. Fed.
Dept. G.1taCand Fish. 44 pp.

Aid

-

r~ Quart.·Prog. Rep.,

P-=-::&gt;j.
W-37-R, October •

1951. Came bird survey. pp. 8082.
Colo. Dept. G(~c and Fish. 117 pp.

In Quart. P~~g. Rep., October.
.'

1952. Gane bird survey. pp. 93-95.
Colo. Dept. G~c ar.d Fish. 105 pp.

In Quart. 27og. Rep., October.

1953. Cane bird survey. pp. 31-89.
Colo. Dept. Gazie and Fish. 99 pp.

In Quart. ?:-og. Rep, , April.

1965. Colorado long range gane species manage=e~~ plans, 1965-1975,
for G~bel's quail. Colo. Dept. Gane, Fish and Par:~~. 8 pp.
Sandfort, U. W., and D. !iolting. 1952. Carae bird surveys ,
pp. 89-91.
Quart. Prog. Rep., October. Colo. Dept. GaQe and F~~~.
105 pp.

In

Sandfort, !{. ~.J., and H. ~I. S~"ope. 1954. Game bird SUn"2::-.
pp. 19-29.
·Quart. Proge Rep., January. Colo. Dept. Caae and 1::':::'.::.
113 pp •

In

•
HOUR.,'UXG DOVE

Braun, C. E. 1971. Trapping and banding dov~s. pp. 201-~07.
In Quart.
Res. Rep., October. Colo. Div. Ga~e, Fish and Park~. 236:pp.
1972. Mourning dove trapping and banding. pp. ::5-121.
Quart. Res. Rep , , October.
Colo. Div. of ·Hild1ife. :'~8 pp.
Grieb, J. R.

1964.

Dove hunting deluxe.

• t'

In

--

Colorado Out~~~=s 13(4):32-39.

1965. Colorado long range gane s?ecies nanage~e~~ ~lans, 1965-1975,
for migratory birds. pp. 33-36.
Colo. Dept. Cazie, : ':..$n and Parks.
36 pp.
Nolting, D., et a1.
Rep., October.

1952. Cane bird survcy. pp. 99-101~
Colo. Dept. Ca~e and Fish. 105 pp.

Sandfort, W ..W. 1950. Gane bird survey.
October. Colo. Dcpt. Gaoc and Fish.

pp. 41-42.
pp.

Aid Proj. W-37-R,

411

1951. G;U:1Cbird survey. pp. "2-43.
Colo. Dept. Ga~c and Fish. 102 pp.
1951. Came bird survey. pp.83-8ft
Colo. Dept. Ca~c and Fish. 117 pp.

Fe~,

In Quart. Prog.

In'Quart. ?-;:'-,:,g .• Rep ,, July.

• .!!!.Qu.nt.?,,;::~Z. Rcp ,, October.

195ft.
Carne bird survey.
pp , 23-24.
Colo. Dcpt. G:lOlCand FLsh , 111 pp •

Tn Oua r t , P";::,"IC.

Rep., Oc t obcr ,

....
...

-- ..
.

"

�-193-

c

Tn Q'J.:Irt. Prog , Rcp ,, July .19550 Came bird survey. pp , 91-92.
Colo •.Dept. Came and Fish. 115 pp.

Sand Eor t , ~l. \I., and D. Nolting.
1952. Came bird sur-vey , pp.103-105.
1A Quart. Prog. Rep., October. Colo. Dept. Game and Fish. 105 pp.
U. S. Dept. of Agr. 1965. Water and related lando resources. Colorado River
Basin in Coloraco.
Coop. Study Rep. of Colo. t-latcr Conserve Eoard and
- UoS.D.A.
Econ , Res. Service, Forest Service and Soil Conserve Senricc.
Hay.
Denver. 183 ppo
(looseleaf)

.

Aldrich, J. i-:., and A. J. Duvall.
1955.
Distribution
of Aner Lc an gallinaceous
gcU3~ birds.
u.S.D.L~ Fish and Hile!. Cire. !:o. 34. 23 pp.

-

I .
1

0

In Qu~:~. Prog. Rep.» January.

Figge, H. 1952. Ca2e bird survey. pp. 45-60.
Colo. Dept. C.&lt;',;::C and Fish.
60 pp.

II

Figges H.~ et a1.
1953.
Ca~e bird survey. pp. 75-86.
January. Colo. Dept. Cane and Fish. 141 pp.

In Quart. Prog. Repos

Uoffman, D. ~!. 1971. Pheasant nest site selection study. pp. 1-26. In __
QU3rt. Res. Rep. ~ April.
Colo. Div. Ca~:·o, Fish and Parks • 125 pp.
0

~___

1972.
Res.

Rep.,

.

Pheasant nest site selection study. pp. 1~19. In Quart.
April. Colo. Div. Gane, Fish and Parks. 150 pp.

Sandf c r t , 't}'~ u. 1950.
Ca::!C! bird survey.
pp , 17--21.
. _"July.
Colo. Dept. G.:u::!C and Fish. 80 pp.

Cane bird survey.
pp , 1 -12.
Dept. Ca.aeand Fish. 44 PI"

1950.

-Colo.

In Quarto Pr og , Rep. 1)

Fed.

Aid Proj. H-37-Rg

Fed.

Aid Proj. '-:-37-R,

0

October.

I

1951. Ca~e bir~ survey.
~--Colo.

Dept. Carae and Fish.

pp. 1-7.
35 pp.

______
o

~_

1951: Ca~e bird survey. pp. 9-20.
Colo. Dept. C~~e and Fish.
102 pp.

~ __ • 1955. C2.;:tC bird survey.
pp.I-9.
Colo. Dept. Canc and Fish.
94 pp.
~

In Quart. Prog. Rep., July.

1952. C3.r71C bird survey. pp , 115-119.
Colo. Dcpt~ Ca~e and Fish.
171 pp.o
1952. Gane bird investigations.
October. Colo. Dc?t_ Ga~e and Fish.

•

1955.

C.:L":lC

July - Part 1.

----. 1956.

bird

s ur vcy,

pp.

In Quart. Prog.

.

Rep., July.

37-41.

105 pp.
In-Quart',

Prog.

/19-63,65-69.
In
Colo. Dcp t , Came .:1Od Fish. J.l5 pp •

Upland g~nc birds.

January.

pro

Color~do Outdoors

Rcp ,, January;

Qua r t , Pr og , Rcp , , .

5(6):2 •

-.:- ••

�-194-

___

•

.

1957.

Upland game birds.

Colorado Outdoors "6(G):2.

1960.

Garac bird survey.

59-65.

Colo. Dept. G~e

and Fish.

pp. 37-55,
161 pp.

•
In Quart. Rep., July - Part II.

G2.~e bird survey. pp. 97-102.
Colo. Dept. Gane and Fish. pp. 97-208.
•

.!.!!. Quart. Rep. , April..

1961.

Gz~e bird survey. pp. 213-223.
In Game Res. Rep.~ October
---Par• 1963.
II.
Colo. Dept. Ga~e, Fish and Park~. -PP. 109-241.

.

t

Sandfort, W. W., and E. M. Swope. 1954. Gane bird survey. pp. 19~29.
" l!!. Quart. Prog. Rep ,, January. Colo. Dept. .Game and Fish. 113 pp.
Stie1-L'";l)

H. A.

July.

1950. Cane bird survey. pp. 1-4.
Colo. Dept. Gane and Fish. 80 pp.

In Quart; Prog. Rep.,

Swope, H. ?-f. 1965. Colorado long z ange game species manag emen t plans,
J.965-1975, fo::pheasants. Colo. Dept. Game, Fish and PzrKs. 54 pp.

U. S. Dept. of Agr.

1966. Water and related la~d resources, wnite River
Basin in Colorado. Coop. Study Re? of Colo. Hater Conserve Board and
U.S.D.A. Econ. Res. Service, ?orest Service, and Soil Conserve Service.
HovelI!ber. Denver. 92 pp. (looseleaf)

SAGE G!\OVSE
Aldrich, J . H., and A.J.
Duvall. 1955. Distribution of Ame rLcan gallinaceous
game birds. U. s.· Fish and HildI. Circ. 34. 30 pp -,
Boeker, H. }of. 1955. Cane bird surveys. pp , 70-71.
..Ju1y.
Colo. Dept. Ga::leand Fish. pp. 1-115.
Funk,

.!E. Quart. Pr'og . Rep , ,

H. D. 1971.
Effects of sagebrush control on distribution and abundance
of sage grouse. pp. 41-47.
~ Quart. Res. Rep., April. Colo. Div. G&amp;~e,

Fish and Parks.

125 pp.

1972." Effects of sagebrush cont::ol en distribution and abundance of
sage grouse. pp. 31-37.
~ Quart. Res. Rep , , April. Colo. Div. Carae ,
Fish and Parks. 149 pp.
Rogers

1958.

J

art II.

Ga~e bird surveys." pp. 123~125.
In Quart. Prog. Rep.,
Colo. Dept. Gene and Fish. pp. 105-181.

1959.
Cane hire! surveys. pp. 13-32.
Colo. Dept. Came and Fish.
116 pp.

---• ,1963.
l'art

The s ag e grouse of Colorado.

In Quart. Prog. Rcp , , April.

Colorado Outdoors 12 (3): 22-24.

1963.
Came bird survey •. pp. 13-32.·
Tn Carne Res. Rcp , , October1. Colo. De p t , C.:lC1C, Fish and Parks. pp. J.-I08.

..

�-~6

-1964.

-195-

S~~e crouse invcsti~~tions
Tech. Publ. No. 16.

in Colorado.
132 pp •

Fish and Parks.

Colo. Dept. Come,

• ·1965. Colorado lonf, range g&lt;lr:lC species ruanagc:ncntplans. 1965-1975.
-----f-or sage grouse. pp. 4-7. Colo. Dept. Gacc, Fish and Parks. 18 pp •
.Ue S. Dept. of Agr.

1965.

Yater and related land resources, Colorado River
Basin in Colorado. Coop. Study Rep. of Colo. ~later Conserve Board and
U.S.D.A. Eeon. Res. Service, Forest Service, and Soil Conserve Service.
l~y. Denver. 183 pp. (looseleaf)

HATERFOWL
Funk, H. D. 1971. Haterf~·",lkill survey. pp , 51-73. In Quart. Res. Rep.,
October. Colo. Div. Cane , Fish and Parks. 236 pp.
Grieb, J. R. 1965. Colorado long range ga~e species ~anag~ent
plans. 1965-.
1975, for migratory birds. pp. 1-32. Colo. Dept. G~ue, Fish and Parks.
36 pp.
Uopper, R. M. 1963. ~7etl~m(lsof Colorado.
Tecll. Publ. !~o. 22, ~f.3.rch.89 pp.
hTILD TU~1.(EY

Colo. Dept. G~c,

Fish and Parks.

..

.Aldrich, J. ~-l., and A. J. Duva l.L,.' 1955. Distribution of Aner Lcan gallinaceous
gaoe birds. U.S.D.I., Fish and Wildl. Ser., Cire. No. 34. 23 pp.
Burget, }L L. 1946.
Fish. 21 pp.

Colorado wild turkey.

Vol. II.

Colo. Dept. Game and

19!,8.
HUd turkey surveys and investigations.
pp. 47-62. In
Quart. Prog. Rcp;, October.
Colo. Dept. Cane and Fish. 80 pp.

_____ " 19!t9. Uildturkey
surveys and investigations .. pp , 91--106. In
Quart. Prog. Rep. Colo. Dept. Gac,e and Fish. 141 pp.
1949. Uild turkey develop~ent.
April. Colo. Dept. G~e an~ Fish.

pp. 3-18.
9.3 pp.

In Quart. Prog. Rep.,

1951. Ui1d turkey rehabilitation in Colorado. pp. 7-17. Spec.
Rep. Glcn~ood Springs Wildl. Coni., Jan. 16-19. Colo. Dept. Game and
·Fisll."
85 pp.
. ..
1957. The ~~ild turkey in Colorado.
January. 68 pp.
1960. Uild turkey dcvc lo pcicn t ,
Colo. Dept. Camc and Fish. 110 pp.

Colo. Dept. Game and Fish.

pp , 5-42.

..!E.. Quart. Rcp ,, July.

1961. Yild turkey devc~opmcnt.
pp. 23-43, 51-55 •
October. Colo. Dept. Carne and Ffsh , 103 pp.

.!Eo Quart. Rep.,

�-196• 1963.
Ulld turkey investiG~tion9.
pp. 71-85.
---I.pril.
Colo. Dept.
and Fish.
109 pp.

In Came Rc~. Rep.,

Carac

Burget,
M. L •• and C. Ford.
1951.
Hlld turkey dcvc l o pmcnt , pp. 1-7. '.!.!l
Quar t , Pr og , Rcp, , April.
Colo. Dept.' Came and Fish.
M pp.
.0
o

Burget,

•

H. L., and D. H. Ilo f fraan ,
1950.
Uild turkey
su rvcys and investigations.
Three Year SL.'7.1 Rcp , , 1947-1950, July.
Colo. Dcpt. Carne and Fish.

~ pp.

.."".

,2nd
~---October:-Colo.

.'

~ and
~~·-In

Quart.

Prog.

1950.
Dept.

tUld
Carae

turkey investigations.
and Fish.
18 pp.

___

, and

1952.

.!E. Quart.
,---'

Prog.

and

---'

Rep , ,

pp ,

1-5.

In. Quart.

62.pp.

Wild turkey investigations.
pp. 1-9, 11-14, 21-24.
October.
Colo. Dept. Gwue and Fish.
105 pp •
pp.
99 pp.

19-33.

~

and
1953.
Hild turkey investigations.
pp. 1-6.
Pr og , Re p , , July. Colo. Dept. Carae arid Fish.
129 Pl'.

, and
---Prog
, Rep.
---.'

&gt;

___

, and

1955. llild tur:~ey i:lvestigations.
P?
Rep:, October.
Colo. bept. GaGe and Fish.

_

Rep , , October.

---'

and
Quart.

1957.

t~ild

Colo. Dept.

turkey

investiga

G2.we and Fish.

In Quart.

_

1-8, 13-19.
108 pp.

pp . 69-86.

In

.!E. Quart.

1953.
Hild turkey i~'lcsti22.tions.
pp. 147-169.
In
~e~.) July - Part II. Colo. Dept. Gane and Fish.
pp. 105-181.

Evans, R. L. 1963~ t-lild turkey invcstir;2..tions.
Rcp , , April.
Colo. Dept. G&lt;.W.eand Fish.

Roff~3n, D. X.
.1965~1975,
llunter,
G. N.
Rep. 1\0.
Hyers,

tions.
116 pp.

Quart •.

In Quart.

1954. Wild turkey investigations.
pp. 12-24.
July. Colo. Dept. Carne and Fish.
182 pp.

and
·Quart. Prog.

H-39-R~

surveys and investigations.
pp • 11-29.
Colo. Dept. GaCle and Fish.
117 pp.

October.

1953.
l-lild turkey investigations.
April.
Colo. Dept. G~e and Fish.

Prog. Rep.,

Aid Proj.

1951. "Jl1d turkey

Rep.,

and
1952. llild turkey investigations;
Prog , Rep , , Ap:::-iJ.. Colo. Dept. Carne and Fish.

-~.-'

Fed.

1965.
Colorado
for ui1d turkey.
1965.

pp. 71-85.
109 pp.

long range game species nanagenent plans,
Colo. Dc?t. G2~e) Fish 2nd Parks.
11 pp.

Colorado big g~~e harvest,
1959-1965.
Gane, Fish and Parks.
41 pp •

1. Colo. Dept.

C. T.
1963.
G'.Jmc bird
October - Part II. Colo.

•

In Ga~e Res.

survey.
Dept.

Spec.

Hgmt.

pp. 173-178.
In G.1::1C Res. Rcp ,,
Camc, Fish and Parks.
pp. 109-2/,1.

..

�-197-

• ·-19M,.

---rm.-t III.

-

Came bird survey. pp. 243-280.
In G.:lL1C Res. Rep. t April Colo. Dcpt. Carnc, Fish and Parks .-pp. 235-397.

1965. Evaluation of a co~bincd blue crouse-wild tu~key hunting '---_.scason.
Outdoor Facts No. 27. Colo. Dept. Game, Fish and Parks.

1 v.

•

1965. Reliability of turkey sex and a~c ratio data based on hunter
report card returns. Outdoor racts No. 29. Colo. Dept. Gili~e,Fish and
Parks. 2 pp.
.J.

1965. Ga~ce bird sU~ley. pp. 91-174. In Cane Res. Rep. 9 April
Part I • Cole. Dept. Gaue, Fish and Parks. pp. r-iz«.
• '1966. C~e bird survey. P? 69-84. In Ga~e Res. Rep.)
---C-·olo. Dept. Gazie, rish and Parks. 192 pp.-1967. Ga=e bird survey. pp. 167-182, 205-211.
April - Part II. Colo. Dept. Ga=e, Fish and Parks •
I'

• 1968. G.?::lC
bird survey. pp , 71, 99-110.
---Colo.
Dep t. Caae , Fish and Parks. 131 pp.
1969. Game bird survey. pp. 107, 153.
Colo. Div. G2::le, Fish auJ Parks. IS1 pp.

1969. The topic is turkey.

April.

..

In Ga~e Res. Rep.,
pp. l07-2l.1.

In Carne Res. Rep ., April.

In Gace Res. Rep., April.

Colorado Outdoors 18(5): 30-34.

1970. Carre bird survey.
pp , 161, 165.
Colo. Div. G2.L~C, Fish and Parks. 193 pp.
Sheats, C. C. 1970.
.19 (3): 30-31. .

-

In Game Res. Rep. J April.

By experience \lith the turkey.

Colorado Outdoors

. ...
'.

"

�•

-198-

•

•
•
•
BOBCAT
Boyd, R. J. 1965. Colorado long r ange garcc species tcanagement; plans, 19651975, for predators. Colo. Dept. Game, Fish and Parks. 21 pp.
:tellinf,tol1,
J. D. 1959.
Fuxbc arers of Colorado.
Colo. Dept. Ga:.:eand Fish. 16 pp.

Educational Pamphlet No. '••

Rutherford, H. H. 1965. Co1ori1do long ri1nge game species nanagp.ment plans,
1965-19jS~ for f urbea rers , Colo. Dept. Game, Fdsh and Parks. 12 pp.
COYOTE
noyd, R. J. 1965. Colorado long range game species management plans, 196519/5, for p reda t o rs , Colo. Dept. Cane , Fish and Parks. 21 pp.
Craighead, F. C. 1951. A biological and econo~ic evaluation of coyote
pred&lt;ltion. Sew York Zoological Society and The Conservation Foundation.
23 pp.
Denney, R. N., and D. L. Gilbert. 1952. Fur resources and bear studies.
pp. 1-31. In Quart. Pr og , Rcp .• .Ian , Colo. Dept. G&lt;1i::e
and Fish. 60 pp.
Sperry, C. C. 19~1. Foed habits of the coyote.
Servo Hi1dl. Re's , Bull. 4. 70 pp."

U.S.D.I. Fish and Wild1.

SKU:-''"K

Boyd, R. J. 1965. Co lorado long range game species management plans, 196519/5, for p redc t ors, Colo. Dept. Cane , Fish and Parks. 21 pp.
Denney. R. N., and D. L. Gilbert. 1952. Fur resources and bear studies.
pp. 1-31. l_~~Quart. Proe. Rep , , Jan. Colo. Dept. Game and Fish. 60 pp.
Remington, J. D. 1959. Furbearers of Colorado.
Colo. De p t , Game and Fish. 16 pp.

Educational P3I:lphletNo.4.

Rutherford, W. II. 1965. Colorado long range game species management plans,
1965-1975J; for furb~arers. Colo. Dept. Game, Fish and Parks. 12 pp •

.
'

RODENTS
Turncr, G. T., R. H. Hansen, V. H. Reid, 11. P. Tietjen, and A. L. Hard. 1973.
Pocket gophcrs ~nd Colorado mountain rangeland. Colo. State Unlv. Expcr.
Sta. Bull. No. 55~S. 90 pp.
•

�FURJ3EARER RESEARCH

REFERENCES

•
.BEAVER
AnODYDOUS. 1937.
1(32):5, 7.

Beavers, flood control experts.

Natl. Nature News.

Finley) t.•••L. 1937. The beavnr-v-conse rve r of soil and water
~ildl. Conf. Tr~ns. 2:295-2~7.
•

0

N. Amer.

J.•eighton, R. S., and J. A. Lee.

1952. A rpogress report on a technique to
control water levels in beaver impoundments. N. A:ree.r.
v!ild1. Conf.
Proc. 8:4 pp.
•

..

Remington, J. D. 1955. Fur resources and bear studies.
W-6l-R. Colo. Dept. Gane and Fish. 85 pp.
1959. Furbc3rers of Colorado.
Dept. Game aud Fish. 16 pp.

Fed. Aid. Proj.

Educational Pamphlet No.4.

Colo.

Retzer, J. L. 1955. Physical environ~ental effects on beavers in the
Colorado Rockies. Western Assoc. Game and Fish COm@ •.Proc. 35.

."

Retzer, J. L., H. ~I. Swope, J. D. Remington, and W. H. Rutherford. 1956.
Beaver manageI::entin the Rocky Hountains of Colorado. Fed. Aid Proj •
W-S3-R. Tech. B~ll~ ~o. 2. Colo. Dept. Game and Fish. 33 pp.
Rutherford, W. H. 1953. Effects of a sumner flash flood upon a beaver
population. J. Har::m.3[. (2) : 261-262.
1964. The beaver in Colorado; its biology, ecology, canageoent
-and econoQics. Colo. Ga~e, Fish and Parks Tech. Pub. No. 17. 49 pp.
1965. Colorado long range game species canagement plans) 19651975, for furbe3rers. Colo. Dept. Ga~e, Fish and Parks. 12 pp.
Scheffer, P. M. 1938.
3(7) :178-.131.

The beav2r as an upstream engineer.

Soil Cons.

}fINK

Denney, R. N: 1950. Fur resource surveys, pp. 75-80.
Rep., July. Colo. Dept. Game and Fish. 80 pp.

In Quart. Prog.

Nemanic, W.}~

Jan.

Fish

1942~

Commfs s Lon,

Fur resource surveys.
6 pp.

Vol. 2.

Colo. Game and

�-200-

Rcminr,ton, J. D. 1953. Fur resources and be3r ~tudics.· pp. 17-23.
1E. Quart. Pr og , Rcp , , Oct. Colo. Dept.' Carne and Ff sh , 60 pp.
• 1959.
---Colo.

Educ a t Lo naL Pamphlet No.4.

Fur-bearers of Colorado.
and Fish. 16 pp.

'Dcp t , Carne,

Rutherford, Y. H. 1965.
Colorado long range game species management plans,
1965-1975, for fu=bearcrs.
Colo. Dept. Game, Fish and Parks. 12 pp •

.-" .

MUSKRAT

Denne)', R. N. 1950. Fur resource survey. pp. 57-66.
Jan. Colo. Dept. Cane and Fish. 66 pp.
1950. Fur resource survey.pp.
93-101.
April. Colo. Dept. G3~e and Fish. 101 pp.
1950. Fur resource survey. pp. 75-80.
Colo. Dept. G~e and Fish. 80 pp.
1951. Fur reso~rce survey.
Dept. Ca~e aud Fish. 8 pp.

~

Quart. Prog. Rep.,

In Quart. Prog. Rep , ,

..!E. Quart. Prog. Rep ,, July:

In Quart. Prog. Rep., Jan.

Colo.

Denney, R. N., and D. L. Gilbert.
1952. Fur resources and bear studies.
pp. 1-31 •. In Quart. Prog. Rep., Jan. Colo. Dept. Gane and Fish. 60 pp.
0
- Nenanic, H. H. 1942.
}:'ishCorcu ss Lon.•

Fur resource survey.
6 pp.

Vol. 2, Jan.

Remington, J. D. 1953. fur resources and bear studies.
Quart. Prog. Rep., Oct. Colo. Dept. Gane·and Fish.

Colo. Ga:ne and

pp. 17-23.
60 pp.

In

.

,

j

;

1959. Furb ear ers of Colorado.
Dept •.C~e an~ Fish. 16 pp.

Educational Panphl.et No. II.

Colo.

Rutherford, ~. H. 1965. Colorado long range ga~e species ~anagencnt plans,
1965-1975, for furbe~rers.
Colo. Dept. Ga~cJ Fish and Parks. 12 pp.

\:EASEL

Denney, R. N. 1950. Fur resource survey. pp. 93-10l.
April. Colo. Dept. Ga~e and Fish. 101 pp.

July.

1950. Fur resource sur/cy. pp. 75-80.
Colo. De~t. Game and Fish. SO pp.

~

..!E. Quart. Prog , Rep.,

Quart. Prog. Rep.,

Denney, R. N., and D. L. Gilbert. 1952. Fur resourccs and bC3r studies.
pp. 1-31. .!E. Quart. -Pr og , Rcp ,, .Jnn, Colo. Dcpt. Carne and Fi~h. 60 pp.
Ncrnanic, W. M. 19~2. Fur rcsource survey.
}':ish Cor.-:nlssion.6 pp.

Vol. 2, Jan.

Colo. C:lmc and

I

�-201RC::1in;:;to!,), J. D•. 195J.

Fur

Quart. Frog. Rep_v Oct.

resources
and bca r 's tud l c s .
Colo. Dept. Came and fl~h •

•

1959.
~10.

Furbeare~s
of Colorado.
Dept. Gtinc and Fish.
16 pp.

~utllcrford, W. H.
1965-1975,

for

In

pp. 17-23.
~O pp.

Pa.mphlet No. ~.

Educational

•

1965.
Colorado long ran~e g~~e spccics-nanagcmcnt plans,
f ur b ca r er s ,
Colo. Dept. Cara e , Fish and Parks.
12 pp •

.-' .

..
•

•

•

•

.:

•

. -..
.

..

...

,

�•

-202GEHERP~ EA~r1ALIA:rREFERENCES

"

IDl.DLUE ~{A.NAGE1E{TUNIT 12;
•
Antnony, H. E. 1928. Field book of North American mammals.
Sons, New York. 674 pp ,
.'

G. P. Putnam's

Armstrong, D. M. 1972. Distribution of mammals in Colorado.
Priating Service; Law~ence. 415 pp.

Univ. r~sas

Bur t , W. H.,

and R. P. Grossenheider.
1964.
Houghton ~li£flin Co., Boston. 284 pp.

A field guide to the mammals.
•

Cringan, A. T. 1973. \.,Tarm
blood vaz t ebra t a inventory analysis and impact
study of the ParaG~ute Creek area, Garfield County, Colorado. In The
Colo:1Y enviror.=ental study--Para~hute Creek, Garfield Co~~ty, Colorado.
Prepared for Colony Develop=e~t Operation by Tno~e Ecological Institute,
Boulder, Colorado. August, 1973. Part II.
Vol. 2, Cahpt. VII. pp. 1-137.
Hall, E. R., and K. R. Kelson. 1959. The m~~~ls
Press Co., New York. 2 Vols. 1162 pp.
Lechleitner, R. R. 1969.
Boulder, 254 pp.

of North America.

Wild mamoals of Colorado.

Ronald

Pruett Pub. Co.,

lliller, G. S., and R. Kellogg. 1955. List of North ft~erican recent mammals.
Sm.Lt hson i an Inst., ~.[ashi11gton,
D. C. Bull. No. 205. 954 pp.
Huric, O. J.
Boston.

1954. A field guide to anjmal tracks.
pp.

374

Palmer, R. S. 1954. r.~e ma~a1
'City, N. Y. 384 pp.
Harren, R. R.
330 pp.

1942.

guide.

Houghton !-lifflinCo.,

Doubleday and Co., Inc., Garden

The mamaa Ls of Colorado.

Univ. Okla. Press, No rman ,

..

...•.

�-203GENERAL AVIAN REFERENCES
WILDLIFE }W~AGEHENT UNIT 12

Bailey, A. M., and R. J. Niedrach.
1965. Birds of Colorado.
Mus. of Nat. Hist., Denver, 2 Vols. 895 pp.
1967. Pictorial checklist of Colorado birds.
---of Nat.
Hist., Denver. 168 pp.
Davis, W. A.
61 pp ,

1969.

Birds of western Colorado.

Denver

Denver Hus.

Colo. Field Ornithologists.

Kortright, F. A. 1943. The ducks, geese, and swans of North America.
The AIDer. Wi1dl. Institute, Washington, C. D. 476 pp.
Linduska, J. P. 1964. Waterfmv1 tomorrow.
Wildl., Washington, D. C. 770 pp.

U.S.D.I. Bur. Sport Fish. and

Martin~ G. S., P. H. Baldwin, and E. B. Reed.
birds from the Yampa Valley, Northwestern
113-116.
Pearson, T. G. 1940.
N. Y.
289 pp.

Birds of America.

1974. Recent records of
Colorado. Condor. 76(1):

Garden City Publ. Co., Inc.,

Peterson, R. T. 1941. A field guide to western birds.
Co., Boston. 240 pp.
Robbins, G. D., B. Brunn, and H. S. Zim.
Golden Press, N. Y. 340 pp.

1966.

Houghton Mifflin

Birds of North America.

Smith, A •.G. 1973. Avian environmental inventory and impact study for
Colony Development Operation in Garfield County, Colorado. Part I.
Environmental inventory by Thorne Ecological Institute for Colony
D~velopment Operation, Atlantic-Richfield Company, Operator, October,
1973.
Sprunt, A., IV., and H. S. Zim. 1961. Gamebirds--a
speci~s 'and their habits. Golden Press, N. Y.

guide to North American
160 pp.

Wooding, J. 1973. A census of the breeding birds of the Roaring Fork
watershed.
Colorado Field Ornithologist.
9 pp.

��July, 1976

-205JOB FINAL REPORT

State of

cO_L_O_RAD
__ O

Project No.

W-lOl-R-lB

Work Plan No.

_
Game Range Investigations

7
Job No.
-------Reprint Book: Winter Guide

1

Job Title _~t_o_N~a~t_i_v._e~S_h_r~ub~s~o~f~t~h~e~C_e_n~t_r_a~l~R~o~c~k~y~MO~u~n~t~a~i~n~s~
_
Period Covered:
Personnel:

April 1, 1975 to March 31, 1976

William T. McKean and Paul H. Neil

ABSTRACT
Concluding work for this segment on the manuscript included minor reVl.Sl.ons
Final printing
to keys and text. Details of proof reading were completed.
and distribution duties will be finished in the early months of the next
segment. Printing cost per copy was approximately $1.70 for 10,000 copies.

��-207REPRINT BOOK: WINTER GUIDE
TO NATIVE SHRUBS OF THE CENTRAL ROCKY MOUNTAINS
William T. McKean
Paul H. Neil

P. S. OBJECTIVE
To revise and reprint the book: Winter Guide to Native Shrubs of the
Central Rocky Mountains.

SEGMENT OBJECTIVES
1.

Continue work on key to genera and descriptions
and usability.

to improve accuracy

2.

Continue revision of all introductory printed page matter.

3.

Arrange printing details with publications
printer.

chief and contract

METHODS AND MATERIALS
1.

Suggestions for improvements in the key and descriptions have been
solicited and received from several cooperators.
These were incorporated.
In a few cases verification of characters were necessary,
using herbarium specimens largely, but freshly collected material
where necessary.

2.

A simple matter of updating and correcting all pages of the preface,
introduction, glossary, index and cover sheets was needed.

3.

Cooperation with Charles Hjelte, publications chief, in the Denver
office was obtained. Printing bids were solicited; a close liaison
with the printer was maintained; and care will be taken to retrieve
and store all original copy when the job is completed. Distribution
of the new book to Division personnel and cooperators will be
supervised.

RESULTS AND DISCUSSION
As of this segment ending, final proofs of all printed material and
illustrations have been checked and returned to the printer. Most of
the detail of printing has been supervised by Charles Hjelte, publications
chief.
There remain only the chores of: (1) Retrieving copy from the printer
(especially the illustrations), (2) Storing it, (3) Distributing the new
book to Division personnel and cooperators, (4) Arranging safe storage
for remaining copies of the book.

�-208-

The printing contract was awarded to Eastwood Printing Company, 2854
Larimer, Denver, Colorado for 10,000 copies (hard bound) costing
$16,879.00 or approximately $1.70 apiece. A sale price of $3.50 each
was established for the general public.

Prepared by

11'

~~~7-1-

%~1c::;t;-CL4t···/

-----------------------------------William T. McKean
Wildlife Researcher

�July, 1976

-209JOB PROGRESS REPORT

State of

COLORADO
------~~~~--------.
Game Range Investigations

W-101-R-18

Project No.

1
8
Job No.
Determination of Digestible Nutrients of
Important Deer and Elk Forage Plants - A Feasibility

Work Plan No.
Job Title

Period Covered:
Personnel:

Study

April 1, 1975 through March 31, 1976.

Roland C. Kufe1d, Marilyn Stevens

ABSTRACT
Five 70 to 125 gram, green weight composite samples of big sage (Artemisia
tridentata) and Gambe1 oak (Quercus gambe11ii) were collected from each
of 9 widely separated areas throughout western Colorado during January,
1976, to determine the average digestible nutrient and degree of variation
in digestible nutrient content of those species during winter.
To date no results are available as samples are currently being prepared
for chemical analysis.

��-211-

DETERMINATION OF DIGESTIBLE NUTRIENTS OF
IMPORTANT DEER AND ELK FORAGE PLANTS - A FEASIBILITY STUDY
Roland C. Kufeld

INTRODUCTION
Availability of winter range is one of the main factors limiting the
size of big game herds on most western ranges. During summer deer
and elk are usually distributed over vast high mountain areas where
the forage supply is adequate. During winter, however, deep snow
forces animals to migrate to the lower valleys where they are concentrated on limited winter range areas. Gilbert, Wallmo and Gill
(1970) found in Middle Park, Colorado that snow over 18 inches deep
essentially precludes deer use. In two of the three winters of their
study over 90 percent of the winter range was excluded from use because
of snow. This type of situation causes heavy concentrations of deer on
critical winter ranges where forage is in extremely short supply, and
where vegetation frequently becomes so heavily used it has little
opportunity for recovery.
Robinette et al. (1952) observed winter deer herd losses of 9, 26, and
42 percent respectively, for three winter range areas in Utah due to
concentration of deer with resulting food shortages. They concluded
that moderate stocking of deer is necessary to maintain the forage
yielding capacity of the ranges, soil and watershed values, and in
addition, the health and well-being of the herds.
Elk die-offs also occur due to excessive concentrations of large numbers
of animals on limited winter range. Jackson Hole, Wyoming is one of the
better known areas where limited winter range has resulted in extensive
elk wint8r losses (Anderson 1958).
Most estimates of carrying capacity or ability of range to support
wintering big game animals are based on observations of animal condition
and population trend records, in belief that carrying capacity has been
reached or exceeded when animal condition begins to decline and dieoffs
begin. Unfortunately, by the time this occurs the range has often
suffered serious damage from which it may take years to recover. Because
of a lack of information and complexity of carrying capacity determination,
wildlife managers have little 'idea of carrying capacity from the range
standpoint.
Some of the primary questions which need to be answered through research
before carry~ng capacity can be determined are listed below:
1.

What plant species do deer and elk eat and what are their relative
preference ratings?

2.

What is the digestible nutrient content of preferred deer and elk
forage species?

�-212-

3.

What are the nutritional requirements of deer and elk?
a.

How much protein, carbohydrate, fats and minerals do deer and
elk require during each season of the year?

b.

How much total volume of plant material do deer and elk eat
per day?

4.

How much forage and digestible energy is produced on the range and
how do you measure this on large areas?

5.

How much of the forage produced on the range can be consumed without damaging the range?

6.

How many deer and elk are on the range, and how can the number of
deer and elk be determined in terms of animal unit months?

7.

Why are there large areas with apparently good forage which are
not used to any appreciable extent by deer and elk?

It will be the objective of this study to answer question number 2,
"what is the digestible nutrient content of preferred deer and elk
forage species"? Deer and elk food habits studies reported in the
literature have been summarized by Kufeld, Wallmo and Feddema (1973)
and Kufeld (1973). These summaries will be used to provide a basis
for selection of the preferred deer and elk forage species to be
evaluated.
Prior to widespread collection of plants for nutrient analysis it will
be necessary to determine the extent of variation in digestible nutrient
content among plants of the same species during winter due to various
environmental factors. It is well known that such variation does occur,
but it mhst be determined if the variation is relatively small, or so
great as to preclude the use of average nutrient values for plant species
in computation of carrying capacity estimates without some sort of data
stratification.
Laycock and Price (1970) have published an excellent review of literature
on environmental influences on nutritional value of forage plants. Their
review summarizes 85 references on the subject. These influences include
season of the year, precipitation, temperature, amount of light, and soil
properties including moisture content, depth, nutrient content, parent
material and texture. Their discussion of factors affecting plant nutrient
content can best be summarized by the statement by Cook and Harris (1950)
who concluded that "the nutrient content of forage is influenced by many
interdependent factors and the result is the additive or mass effect of
all factors operating Simultaneously". If plant nutrient content data are
to be used by wildlife managers in carrying capacity estimates, it is
therefore apparent that these will have to be based on average digestible
nutrient content values for each plant species during each season of the
year. Use of average digestible nutrient content data will be feasible
only if variation due to all affecting factors is reasonably small. Many

�-213-

of the studies have shown the influence of certain factors resulted in
nutrient content differences that were statistically significant. Even
though statistical significance was shown many of the differences were
not particularly large numerically. Thus, the first phase of this
study will be directed toward determining whether the degree of nutrient
variation will permit use of average nutrient values in carrying capacity
estimates.
If the variation in digestible nutrient content among plants of the same
species is found ~o be quite large, and it is ascertained that digestible
nutrient content cannot be sampled adequately for management purposes
even through stratification the project may be terminated at this point.

P. S. OBJECTIVE
To determine the average digestible nutrient content values and the degree
of variation in digestible nutrient content of selected range forage
plants during winter.
SEGMENT OBJECTIVE
To determin2 the average digestible nutrient content values and the degree
of variation in digestible nutrient content of Artemisia tridentata and
Quercus gambellii during winter.

METHODS AND MATERIALS
Five 70 to 125 gram, green weight, composite samples of big sage (Artemisia
tridentata) and Gambel oak (Quercus gambellii) were collected from each
of 9 geographically widely separated areas within the normal winter range
of deer and elk. The 5 samples were taken from widely distributed sites
within each major area with all 5 samples collected from the same vegetation type. For sagebrush collections 3 of the major collection areas
represented the sagebrush type, 3 represented the pinyon-juniper type
and 3 represented the Gambel oak type. Major collection areas for Gambel
oak represented the oak, pinyon-juniper and ponderosa pine-oak types and
3 major areas were selected representing each type.
Ten composite big sagebrush samples of at least 200 grams each were
collected from 10 widely separated areas within the normal winter range
of deer and elk. These samples will be used to determine the percentage
of volatile oils lost due to freeze drying and grinding sagebrush samples,
-which is a necessary step in preparing samples for chemical analysis.
Each 200+ gtam sample was divided in half resulting in 10 paired samples.
A reconnaissance trip was made throughout the western half of Colorado
during the summer of 1975 to establish locations where the desired species
could be collected. Plant collections were made during January, 1976.
Oak collections consisted of current annual growth stems with leaves and
acorns excluded. For big sage current annual growth stems with leaves

�-214-

attached wer e collected.

Sage seed stalks were not collected.

Information recorded on each composite plant sample included the
following: 1) date, 2) exact location, 3) vegetation type, 4) elevation, and 5) green weight of plant samples to the nearest gram when
collected.
Samples of big sage were placed in sealed plastic bags, packed in a
snow filled ice chest immediately upon collection and frozen as soon
thereafter as possible to prevent vaporization of volatile oils which
might result in unrealistically high digestion coefficients.

RESULTS
To date no results are available.
for chemical analysis.

Samples are currently being prepared

LITERATURE CITED
Anderson, Chester C. 1958. The elk of Jackson Hole. A review of Jackson
Hole elk studies. Wyo. Game and Fish Corom. Bull. No. 10. 184 pp.
Cook, C. Wayne, and Lorin E. Harris. 1950. The nutritive value of range
forage as affected by vegetation type, site and state of maturity.
Utah State Agric. Expt. Sta. Tech. Bull. No. 344. 45 pp.
Gilbert, Paul F., Olof C. Wallmo, and R. Bruce Gill. 1970. Effect of
snow depth on mule deer in Middle Park, Colorado. J. Wildl. Manage.
34(1):15-23.
Kufeld, Roland C. 1973. Foods eaten by the Rocky Mountain elk.
Manage. 26(2):106-113.

J. Range

, O. C. Wallmo, and Charles Feddema. 1973. Foods of the Rocky Mountain
mule deer. U.S.D.A. For. Ser. Rocky Mtn. For. and Range Exp. Sta. Res.
Pap. RM-lil. 31 pp.
Laycock, William A., and Donald A. Price. 1970. Factors influencing
forage quality. In: Range and wildlife habitat evaluation - a research
symposium. U.S.D-:-A.For Servo Misc. Pub. No. 1147. pp. 37-47.
Robinette, W. Leslie, Odell Julander, Jay S. Gashwiler, and Justin G. Smith.
1952. Winte~ mortality of mule deer in Utah in relation to range
condition. J. Wildl. Manage. 16(3):289-299.

Prepared by

~«LdC,~
Roland C. Kufeld
Wildlife Researcher

�July 1976
-215JOB PROGRESS REPORT

State of

COIDRADO
--------~~~~-----------

Project No.

W-125-R-2

Work Plan No.

15

Job Title

Monitor Potentially

Deer-Auto Accident
Job No.

Investigations

l_a

Critical Deer-Vehicle

Accident Areas

_
Statewide

Period Covered: April 1, 1975 through March 31, 1976
Personnel:

All Area Supervisors and Wildlife Conservation Officers in
Colorado, James D. Fleming, Edward E. Kochman, Sharon L.
McDonnell, Dale F. Reed, and Thomas N. Woodward.

ABSTRACT
A total of 1,527 big game-vehicle accidents were documented in Colorado in
1975. Two hundred thirty-five deer were killed by vehicles in seven
specific areas in western Colorado. Recommendations for installation of
devices to reduce deer-vehicle accidents were submitted during this segment for three of cheae areas.

�-216-

RECOMMENDATIONS
Recommendations were developed from this study and made to the Colorado
Division of Highways for installation of the eight-foot fences and
associated structures referred to in this report.

�-217-

MONITOR POTENTIALLY CRITICAL
DEF,R-VEHICLE ACCIDENT AREAS STATEWIDE
Thomas N. ~..]oodard

P. S. OBJECTIVE
Locate an~ examine potentially critical deer-vehicle accident areas in
Colorado and recommend methods or devices which may reduce the problem
for these critical accident areas.

SEGMENT OBJECTIVES
1.

Locate and examine potentially
in Colorado.

critical deer-vehicle

accident areas

2.

Recommend methods or devices which may reduce the problem for these
critical accident areas.

3.

Select recommended underpasses, overpasses, 2.44-m fence lengths, or
other devices or methods for evaluation when sample size considerations
can be met.

DESCRIPTION OF AREAS
Highway 6-24 (1-70) Rifle T..]est
Pojar (1972) and T..]oodard
(1973a, 1975) described this area.
Highwav 1-70 Eagle East
Pojar (1972) and Woodard

(1973a, lq74) described this area.
Highway 1-70 Avon-Wolcott

Pojar (1972) and Woodard

(1973b) described this area.
Highway 82 - Glenwood-Basalt

Myers (1969) and Woodard

(1975) described this area.

Highway 6-24 (1-70) Eagle-Glenwood
Woo~ard

(1975) described this area.

Canyon

�-218-

Highway 550 Colona-Ridgwav

.

This area extends from the Uncompahgre River bridge 7.0 km south of Colona
to Owl Creek Road 18.6 km south of Colona, a distance of 11.6 km. The
present h Lghwav alignment is ad[acen t to and east of the Uncompahgre River.
Pinvon-;uriiper (Pinus spp.-Juniperus sPP.) connnunities predominate on
adjacent west faCing slopes. Approximatelv 8.0 km of the present alignment
is scheduled for realignment to the east to bypass Ridgway Reservoir, part
of the U.S. Bureau of Reclamation Dallas Creek water development nroject.
The realigned section of highwav will be Q.6 km long. The present alignment
will remain for approximate Iv 2.3 and 1.3 km at the north and south ends
of this section of highway, respectively.
Highway 50 Salida East
This area extends from Howard, Colorado, to the east approximatelv 8.4 km.
The highway is adjacent to and southwest of the Arkansas River. Pinyonjuniper connnunities predominate on adjacent northeast facing slopes. This
segment of highway is scheduled for improvement construction.

METHODS AND MATERIALS
Methods and materials have been described by Pojar (1972).
Highway 550 Colona-Ridgway
The numbers and locations of deer-vehicle accidents during 1974 and 1975
were plotted on U.S.G.S. quadrangle maps (scale 1:24,000). One evening
ground inventorv was made on 17 April 1975 along the present alignment
and proposed realignment.
Cost:benefit analvsis for 2.44-m fencing was
completed for the 3.6 km of remaining highwav and 9.6 km of realignment
using the technique described by Woodard and Reed (lQ74).
Deer-vehicle,accidents
on the realignment were projected bv extrapolating
from the present alignment. The assumption was made that deer near the
present alignment will be diSPlaced to the east along the proposed
realignment.
Accidents on the proposed realignment without fencing were
projected by multiplying the number occurring on the present alignment
by the percent increase of deer reSUlting from displacement, using data
collected on the 17 April 1975 ground inventory.
Accidents on the
realignment with fencing were calculated by multiplving the projected
accidents without fenCing by 0.22, the converse of the effectiveness of
five 2.44-m fences installed to date in Colorado (Woodard 1976).
Highway 50 Salida East
The number and locations of deer-vehicle accidents during 1975 were
plotted on a U.S.G.S. quadrangle map (scale 1:62,500).

�-219-

Statewide
All Area Supervisors and Wildlife Conservation Officers in Colorado were
requested to document big game-vehicle accidents and send records to
"Records, Big Game-Vehicle Accidents" in Glenwood Springs for filing and
review. Implementation of this request was I Julv 1975.

RESULTS AND DISCUSSION
Highway 6-24 (1-70) Rifle West
Twenty-five vehicle-killed deer were documented on 29 km of highway in
the Rifle West study area; a decrease of 19.4 percent from the preceding
segment.
~iftv-two percent of the deer were killed during April.
Highwav 1-70 Eagle East
Twenty-four vehicle-killed deer were documented on 13.7 km of highway in
the Eagle East study area; an increase of 4.3 percent over the preceding
segment. Deer kill continued to be reduced on the portions of highway
opposite the 7.7-km barrier fence (Woodard 1976).
Highway 1-70 Avon-Wolcott
Fifty-three vehicle-killed deer were documented on 22.5 km of highwav in
the Avon-Wolcott studv area; a decrease of 11.7 percent from the preceding
segment. Deer kill continued to be reduced on the portions of highwav
opposite two sections of 2.44-m fence O-Joodard 1(76).
Highway 82 - Glenwood-Basalt
Seventy vehicle-killed deer were documented on 29 km of highwav in the
Glenwood-Basalt study ,area; a decrease of 27.8 percent from the preceding
segment.
Twenty-nine percent were killed during February and March.
Thirty-nine percent were killed during April. Deer-vehicle accidents
continued to be reduced on the portions of highway opposite two 1.77-km
lengths of 2. 44-m fencing (Woodard 1n6) .
Highway 6-24 (1-70) Eagle-Glenwood

Canyon

Eight vehicle-killed deer were documented on Highway 6-24 from Eagle to
the west 9.9 km to Gypsum.
Sixteen vehicle-killed deer were documented
on Highway 6-24 f roia Gypsum to the west 15.0 km to the beginning of
Glenwood Canyon. Recommendations for structures to reduce deer-vehicle
accidents when Interstate 70 is constructed between Eagle and Gypsum
were submitted to the Colorado Division of Highways.
Recommendations
included two open bridge structures and 8.6 and 6.8 km of 2.44-m fencing
adjacent the north and south sides of Interstate 70, respectively.

�-220-

Highway 550 Colona-Ridgway
An annual mean of 31.0 deer-vehicle accidents were recorded on the present
Highway 550 alignment during 1974 and 1975. There were 26.0 accidents
along the 8.0 km scheduled for relocation.
Two hundred twenty-seven deer were observed during the evening of 17 April
1975 adjacent the segment of Highway 550 scheduled for realignment. The
same evening 95 deer were observed along the proposed realignment. Based
on these data, 42 percent more deer would be located along the realignment
if deer are displaced to the east of the reservoir. A corresponding 42
percent increase in deer-vehicle accidents would result in an annual mean
of 36.9 accidents on the 9.6 km of realignment. There would be 41.9
projected accidents annually for the entire 13.2 km of highway.
Recommendations for 2.44-m fenCing on this segment of highway have been
submitted to the U.S. Bureau of Reclamation. Recommendations were for
13.2 and 14.5 km of fence on the east and west side, respectively, and
20 one-way gates. The cost:benefit ratio for this fencing after 20 years
will be 1:0.8
Estimates of projected deer-vehicle accidents on the realignment are
conservative because "documented deer-vehicle accidents on the present
alignment are'considered minimal. If this is the case, then the cost:
benefit ratio for the 2.44-m fencing would be improved. The cost:benefit
ratio would also be improved if a monetary value was placed on the loss of
deer. Using a value of $709 (Norman et al. 1975) for each deer the 20-year
cost:benefit ratio for the fencing plan would be 1:2.0.

Highway 50 Salida East
Fourteen deer-vehicle accidents were documented on this section of highway
during 1975. Some 2.44-m fencing has been recommended for both sides of
the highway and will tie into two existing open bridge underpasses. Both
of these underpasses were monitored for deer use in 1975. Results were reported by Reed (1976).

Statewide
A total of 1,527 big game-vehicle accidents were documented in Colorado
in 1975. The number of accidents documented in the northeast, southeast,
southwest, and northwest regions were 206, 256, 656, and 409, respectively.

LITERATURE CITED
Myers, G. T. 1969. An investigation of deer-auto accidents. Pp. 147-178
in Game Research Report, W-38-R, July, Part 2. Colo. Game, Fish and
Parks Div. Fed. Aid.

�-221-

Norman, R.L., L.A. Roper, ~.D. Olson, and R.L. Evans. 1975. Using wildlife
values in benefit/cost analysis and mitigation of wildlife losses.
Colorado Div. Wildl. DOW-M-D-3. l8pp.
Pojar, T.M. 1972. Monitor potentially critical deer-vehicle accident areas
statewide. Pages 267-275 in Game Research Report, July, Part 3.
Colorado Div. Wildl. ~ed.-Xid.
Reed, D. F. 1976. Responses of deer to highway underpasses.
Report. Colorado Div. Wildl. ~ed. Aid. In Progress.

In (;ame Research

Woodard, T.N. 1973a. Monitor potentially critical deer-vehicle accident
areas statewide. Pages 163-170 in Game Research Report. July, Part 2.
Colorado Div. Wildl. Fed. Aid.
1973b. An evaluation of deer-proof fence length required to
prevent deer movements on or across high speed highways. Pages 197202 in Game Research Report, July, Part 3. Colorado Div. lVildI. Fed .
Aid.
. 1974: Monitor potentiallv critical deer-vehicle accident
------areas statewide. Pages 293-298 in Game Research Report, July, Part 2.
Colorado Div. loJildI.Fed. Aid.
, and D.P. Reed. 1974. Economic considerations in reduction
of deer-vehicle accidents. Central Mtn. and Plains Sect. TVildI. Soc.
Conf. Bpn . Typescript.

--------

________
. 1975. Monitor potentially critical deer-vehicle accident
areas statewide. Pages 295-303 in Game Research Report, Julv, Part 2.
Colorado Div. Wildl. Fed. Aid.
1976. An evaluation of deer-proof fence length required to
prevent deer movements on or across high sneed highways. In r.ame
Research Report. Colorado Div. Wildl. Fed. Aid. In Progress.

Prepared by

:--()j~~-&amp;J--+---

-::::f-fl---Lh--=.L6fn-,---o. 0_· ~A/--:.
Thomas N. Woodard
Assistant Wildlife Researcher

��July 1976
-223JOB PROGRESS REPORT

State of

COLORADO
--------~~~~------------

Project No.

W-125-R-2

Deer-Auto Accident Investigations

Work Plan No.

15

Job No •

Job Title

6a

-------------------------------

Evaluation of Deer Underpasses

Period Covered:
Personnel:

~------------------

.

April 1, 1975 through March 31, 1976.

James D. Fleming, Arthur J. Gresh, Willie Travnicek, Don Masden,
Thomas N. Woodard, and Dale F. Reed.

ABSTRACT
Nine underpasses were monitored for deer use. Moderate to high numbers of
deer passages (n=139 and n=124) occurred through two of the underpasses,
one located east of Avon and the other southeast of Salida. Extremes of
openness or tunnel effect (0.03 and 5.17) were represented by these two
structures in the study sample. Early morning observations were made of
deer behavior at the Vail underpass exit. No flight 'reaction was observed
during 92.9 percent of westbound traffic (n=449) when deer were present
at the Vail deer underpass. The behavior modes of 108 deer exits from the
underpass were 47, 41, and 12 percent, trotting, walking, and bounding,
respectively.

��-225-

EVALUATION OF DEER UNDERPASSES
Dale F. Reed
Mule deer response to an underpass 3.05 m bv 3.05 m and 30.48 m long under
Interstate 70 has been reported (Reed et al. 1971)). This studv purports
to evaluate underpasses of different sizes and design and to further
evaluate deer behavioral responses.

P. S. OBJECTIVE
Determine the use of underpasses by deer in critical highway deer kill
areas and improve the effectiveness of these structures.

SEGMENT OBJECTIVE
Measure deer use of underpasses
associated with their use.

and the extent of behavioral

reluctance

METHODS ~~D MATERIALS
Nine underpasses which were potentially used bv deer were monitored for
use by track-count methods previously described (Reed 1971). A small
structure under 1-70 east of Avon, three concrete box underpasses under
1-70 east of Eagle, a single-span structure at Chaffee r,ulch north of
Ridgwav, the arch (structural plate pipe arch) deer undernass west of
Durango, and three underpasses, two of which were large onen structures
southeast of Salida, were evaluated.
Observations were made in the early
morning hours at the Vail deer underpass. A 2flX spotting scone was used
to observe deer at the underpass from about 0.5 km away. Vegetational
cover and camouflaging were utilized.
The observations were made during
morning periods for approximate Iv two hours from about 053() to ()73()DST
for 30-35 days during the June-July migration period.
Flight reaction
or escape behavior (Hediger 195fl, Scott 1958) was recorded for every
animal or group of animals exhibiting such behavior in a confined area
north of the underpass entrance when vehicles passed to the west. Only
westbound vehicles were considered since they are closer (within 15-33 m
of the animals) and generate greater auditory and visual stimuli to the
animals than eastbound vehicles.
The number of animals of each groun
that exhibited flight reaction by leaving the area during vehicular
passage or immediately thereafter were noted in three categories (zero,
I-SO, or 51-100 percent).
Exit behavior (walk, trot, or bound) and
additional notes relating to the animals apparent wariness was recorded
for each animal observed exiting the underpass.
Use of and behavioral
response to this underpass during a previous 4-year studv has been
reported (Reed et al. 1975).

�-226-

RESULTS ~~D DISCUSSI0N
The underoass east of Avon received 139 passages (Table 1) between 24
October and 17 December 1975. ~iftv passages occurred to the north and
89 to the south. Use on this small structure, 1.06- X lOB.7-m (radius
X length), was discovered during a spotlight count (Reed 1969) when a
doe and two fawns moved to the south highwav right-of-wav and "disappeared."
A doe and two fruYilswere subsequently (within 10-15 min.) counted on the
north side of the interstate and 2.44-m fence. It was hvpothesized that
the animals had crossed the highway and passed through one of several
one-way gates. An immediate check of these structures indicated this did
not occur. A check of the underpass approach, inside-floor-surface,
and
exit, yielded deer tracks directed to the north. Regular checks of the
underpass were initiated promptly.
Two of three underpasses east of Eagle received passages.
Six nassages
have occurred through underpass No.1
(previous No. 7.), three to the south
and three to the north, and three have occurred through undernass ~o. 2
(previous No.3),
all to the south. The passages that occurred to the
north are of special interest since they took nlace ,vithout the effect of a
2.44-m fence along the south side of the interstate.
Only the north side
of the interstate is fenced with 2.44-m fencing in the Eagle East area.
The Chaffee Gulch underpass was checked (n=16) mostly by southwest Region
personnel.
These checks yielded 35 passages, 15 east and 20 west. The
sub-standard bridge had been replaced with an equal Iv open single-snan
bridge.
The arch deer underpass located west of Durango and the Cherrv Creek
campground, has received an average of 66.5 passages ner year. The deer
activity adjacent the underpass has been light. Considerable human and
cattle use has possibly affected deer movements in this area. A cost:benefit
ratio, calculated bv methods previously described (Woodard and Reed 1974), for the
area involving the underpass was unfavorable (1:&lt;0.0).
The two large open underpasses southeast of Salida received most of the
passages in this area (Table 1). It was hypothesized that deer use of
these structures was related to the availability of water in the Arkansas
River, located within 200 m north of the structures.
Track counts yielded
79 and 45 passages to the north and south respectivelv for underpass No.1,
and 20 and 15 to the north and south respectively for underpass No.2.
Both of these structures have the highest openness or tunnel effects (5.17
and 5.39) so far encountered in this study. Plans call for evaluating the
behavioral response of deer to the Salida East No. 1 underpass in the next
segment.
Daily observations (n=3l) were made at the Vail deer underoass from 4 June
to 4 July 1975. Of 449 instances of vehicles traveling west when deer were
near the underpass entrance and the west bound lanes, 417 did not elicit

�-227-

flight reaction from 104 different individuals or groups (ran~e 1-11
deer). Seven vehicles elicited flight reaction from 1-50 percent of the
individuals from eight different groups ranging in size from one to eight
deer (Table 2). Twenty-five vehicles elicited flight reaction from 5l-l0~
percent of the individuals from seven different groups ranging in size
from one to six. Previous to thes'e observations it was hypothesized that
deer at proximal distances of 10 to 20 m from moving traffic were frequently
caused to leave the area. Other observations indicated that when vehicles
stopped at or when bicyclists traveled by the underpass, most deer elicited
immediate and intense flight reaction.
The behavior modes of 108 deer exits
(leaving the underpass to the south) were 47, 41, and 12 percent, trotting,
walking, and bounding, respec tively. These combined data from 1974 and
1975 yielded (n=108 deer exits) a walk:trot:bound ratio of 3.4:3.9:l.~.
Wariness appeared to be exhibited irrespective of the mode of exit (walk,
trot, or bound) in many instances.
The predominant exit behavior of
trotting supports the renorted reluctance (Reed et al. lQ75) of deer to
use a structure of this size and character.

LITERATURE

CITED

Hediger, H. 1950. Wild animals in cantivity. Butterworth,

London. P. 19.

Reed, D.F. 1969. Techniques for determining notential1v critical deer
highway crossings. Game Infor. Leaf1. 73. Colorado Div. r.ame, ~ish
and Parks. 3pp.
________ ~. 1971. Deer underpass evaluation. Pages 341-351 in r.ame Research
Report, July, Part 3. Colorado Game, Fish and Parks Div. ~ed. Aid.
_________ ., T.N. Woodard, and T.M. Pojar. 1975. Behavioral resnonse of
mule deer to a highway underpass. J. Wild1. Manage. 39(2):361-367.
Scott, J.P. 1958. Animal behavior.
Chicago. 349pp.

2nd ed. University

of Chicago Press,

Woodard, T.N., and D.F. Reed. 1974. Economic considerations in reduction
of deer-vehicle accidents. Central Mtn. Plains Sect. Wild1. Soc.
Conf. 8pp. Typescript.

"Prepared by

~Ai&lt;i&amp;:~k

Dale F. Reed
Wildlife Researcher

�-228-

Table 1. Highway underpasses, height and width dimensions or open-end surface
area, number of deer passages, openness or tunnel effect, and deer activity
adjacent to the underpasses.
Only the first and last structures were intended
for regular deer use.

Height X width
or open-end
surface area

Underpass
(Location)

}j

(m)

Arch deer
(W. Durango)

3/
3.05 X 6.10 4/
n(1.06)2 -

Avon 1-70 No. 1
Chaffee Gulch
(N. Ridgway)

20.46

'2/

Deer
passages
per year

Openness
or tunnel
effect 2/
(m)

-

Adjacent deer
activity/barrier
fencing adjacent
structure

66.5

0.61

Light/both sides

139.0

0.03

Light/one side

35.'l

1.51

Light/none

Eagle 1-70 No. 1

4.27 X 4.27

3.0

0.42

Moderate/one

side

Eagle 1-70 No. 2

2.44 X 2.44

1.5

n.lO

~oderate pressure
close to entrance/
one side

Salida East
No. 1

6/
5.49 X 14.63 -

124.0

5.17

Concentrated during
April-May/none

Salida East
No. 2

6/
7.32 X 14.63 -

35.0

5.39

Moderate/none

Salida East
No. 3

2.44 X 3.05

1.0

0.23

Moderate/none

Vail deer 1-70

3.05 X 3.05

345.11)

0.31

Concentrated, highly
motivated migration/
both sides

l/ Width measured

parallel to direction of traffic flow.

2/ An expression

of openness is ca1cuted as follows:

height X width (or open-end surface area) =
length

1/ Arch-type

structure.

~/ Circular structure.

'if Single-span
i/ Adjusted

bridge with fill-slope on each side.

for uneven inside topography.

2/ Seasonal mean resulting from a 4-year study (Reed et ala 1975), presented for
comparison.

�-229-

Table 2. The number of westbound vehicles which elicited flight reaction
or escaue behavior (Hediger 1950, Scott 1958) at the Vail deer Underpass
entrance for 1974 and 1975.

Flight Reaction
(percent of group)

Year
1974

1975

Total

193

224

417

1-50

4

3

7

51-1()()

1

24

25

~
()

I

•

I

�I

4

I

�,

-231-

July, 1976

JOB PROr.RESS REPORT

State of

COLORADO
------------------------

Project No. ~W_-~1~2~5_-~R~-~2
Work Plan No.
Job Title

15

Deer-Auto Accident

Investigations

Job No.

7

Effects of Highway Lighting on Number of Deer Killed by Vehicles

Period Covered:
Personnel:

_

April 1, 1975 through March 31, 1976.

Dennis Donnelly (Division of Highways), Claudia A. Doose,
James D. Fleming, Larrv L. r.reen, Rich Griffin (Division of
Highways), Sharon L. McDonnell, John W. Seidel, Dale 1&lt;'. Reed,
and Thomas N. 1..Joodard.

ABSTRACT
The effect of highway lighting on the rate of deer-vehicle accidents
was evaluated for the third year. Estimated deer crossings per kill
was q.7 percent higher with the lights on compared to lights off.
There was no significant difference in the crossings per kill ratios
(X2 = 0.073, P&gt;0.75).
Calculated mean horizontal illumination and the
horizontal uniformity ratio for the l.l-km lighting system were 1.02
foot candle and 18.55:1, respectively.

��I

-233-

EFFECTS OF HIGHWAY LIGHTING
ON NUMBER OF DEER KILLED BY VEHICLES
Thomas N. Woodard

P. S. OBJECTIVE
Determine if deer-vehicle accidents are affected by the fixed highway
illumination on a l.l-km segment of Colorado Highway 82, and investigate
and compare responses of motorists to deer on the highway and deer
responses to the motorists, with and without fixed illumination.

SEGMENT OBJECTIVES
1.

Measure and describe the light availability
segment of Colorado Highway 82.

on an illuminated

2.

Determine if deer-vehicle accidents are affected by the fixed
illumination on the l.l-km segment of Colorado Highway 82.

3.

Investigate and compare responses of motorists
with and without fixed illumination.

4.

Investigate and compare deer responses
fixed illumination.

l.l-km

to deer on the highway,

to motorists,

with and without

METHODS AND MATERIALS
Methods

and materials

have been described by Woodard

(1974).

The effect of lighting was evaluated from 1 April to 23 April 1975 and
2 January to 25 March 1976 during this segment.
Data collected during
1974 and from 2 January to 31 March 1975 are also included in this report.
An anemometer, hygrothermograph,
recording rain gauge, snow stake, and
maximum and minimum thermometers located approximately 10 m from the
edge of the highway were used to collect weather data.
Horizontal illumination levels for the Highway 82 lighting system were
measured in footcandles (fc) with a General Electric SL480A light meter
on 10 April 1975. The photo-sensitive cell was placed parallel to the
roadway surface on the hood of a vehicle traveling approximately 16 kmph
in the driving and passing lanes and along the shoulder of the highway
in both directions.
Mean horizontal levels for the 1.1- and O.4-km
segments of illuminated highway (Woodard 1974) were calculated by averaging
the illumination levels estimated from a continuous trace at each 7.6-m
distance.
Horizontal uniformity levels were calculated bv dividing the
mean horizontal levels (X) with the mean minimum illumination levels (minX).

�-234-

RESULTS AND DISCUSSION
Estimated deer crossings per kill was 9.7 percent higher with the lights
on compared to lights off (Table 1). There was no significant difference
in the ratios (X2 = 0.073, P&gt;0.75).

Table 1. Estimated deer crossings and total kill in the Highwav 82
lighting studv area during 1974 and 1975.

Lights OFF

Lights ON

1581

1637

Total Kill

18

17

Crossings/Kill

87.8

96.3

Estimated

Crossings

There were no estimated crossings or kill during the 1976 portion of this
segment.
Few deer were observed adjacent the study area (Table 2). The
probable reason for the low deer numbers in 1976 was the relative mild
weather conditions (Table 3).

Table 2. Mean deer observations during spotlight counts and total kill
in the Highway 82 lighting studv area from 1974 to 1976.

1
Year

Mean count

1974

27.1
(n=71)

13

1975

40.3
(n=98)

22

1976

0.3
(n=65)

0

Total kill

�-235-

Table 3. Mean daily ambient temperatures and snow depth in the Highway
82 lighting study area during January-March periods from 1974 to 1976.

1/

Year

Mean temperatures
(OC)

1974

-3.5

20.8

1975

-2.8

22.8

1976

-1. 7

4.7

1/ The January-March

Mean snow depth
(em)

mean of the daily means of maximum
ambient temperatures.

and minimum

The Roadway Lighting Committee (1972) recommends lighting standards
based on mean horizontal illumination and illumination uniformity ratios
for different roadway and area cLass LfLcat.Lons.. Recommendations for the
Highway 82 roadway and area classification are 1.0 fc mean horizontal
illumination and an illumination uniformity ratio of 3:1. Calculated mean
horizontal illumination and illumination uniformity ratios for the 0.4and l.l-km segments of Highway 82 lighting were 1.47 fc and 15.47:1 and
1.02 fc and 18.55:1, respectively.
These uniformity ratios support
visual observations that dark areas were present in the study area during
lights on nights.
Since the lighting system is not uniform and because contrast ratios
are the most important elements for motorist visibility (Gallagher and
Meguire 1974), a spotmeter (Spectra Model UBA, Photo Research, 3000 N.
Hollywood Way, Burbank, California) was purchased during this segment.
This instrument will be used during future segments to make target (deer)
and background luminance measurements from which contrast ratio calculations
can be made at each kill location. Deer-vehicle accidents which do not
occur under sufficient visibility can then be deleted from the sample.
An observation tower which will be used to investigate motorist and deer
responses was constructed during this segment.
Specific methodology
which will be used was described by Woodard (1975).

LITERATURE

CITED

Gallagher, V.P., and P.G. Meguire. 1974. Contrast requirements of urban
drivers. Fed. Highway Admin. Rep. Number PHWA-RD-74-76. 72pp.

�-236-

Roadwav Lighting Committe. 1977.. American national standard practice
for roadway lighting. J. Illuminating Eng. Soc. 1(4):334-371.
Woodard, Thomas N. 1Q74. Effects of highway lighting on number of deer
killed bv vehicles. Pages 307-312 in Game Research Report, Ju1v,
Part 2. Colorado Df.v , I.Jildl.Fe d , Aid.
----~----~-----.
1975. Effects of highway lighting on number of deer
killed by vehicles. Colorado Div. Wi1d1. Fed. Aid. Program Narrative
Outline. 7pp.

Prepared by

/k.a s J/ 0-f)~
Thomas N. Woodard
Assistant Wildlife

Researcher

�July, 1976

-237JOB FINAL REPORT
State of

COLORADO

---------------------

Project No.

W-125-R-2

Work Plan No.

15

Job Title

Job No. .

8

-----------------------------------

Crossing Signs ~~fNS~~e~fotifl~~fdK~II~d by Vehicles

Period Covered:
Personnel:

Deer-Auto Accident Investigations

April 1, 1975 to March 31, 1976

D. F. Reed and T. N. Woodard

ABSTRACT
No work was accomplished on this job during the past segment because a
suitable study area for an independent test of the effects of lighted
deer crossing signs could not be located. This job will not be reactivated
in the next segment because of severe time limitations of project personnel.

��-239-

July, 1976

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No.

W-125-R-2

Work Plan No.

15

Deer-Auto Accident

Investigations

Job No.
10
An Evaluation of Deer-Proof Fence Length
Job Title Required to Prevent Deer Movement on or Across High Speed Highways

---------------------------------

Period Covered:
Personnel:

,

April 1, 1975 through March 31, 1976.

Claudia A. Doose, James D. Fleming, William R. Heicher, David C.
Hoart •.Kenneth R. Kincaid, Sharon L. McDonnell, Dale F. Reed, and
Thomas N. Woodard.

I

ABSTRACT
Annual or seasonal deer kill caused by vehicles was reduced by 57, 78, 85,
62, and 81 percent along 2.44-m fence lengths of 1.8, 1.8, 3.6, 3.6, and
7.7 km, respectively.
Sixty-four, five, and 14 deer escaped the highway
rights-of-way adjacent to the two 3.6- and the 7.7-km long 2.44-m fences,
respectively, through 16 one-way gates. Collared does (N=20) and bucks
(N=7) moved laterally or parallel to the 2.44-m fence for mean distances
of 0.66 and 0.76 km, respectively.
The mean daily linear movements on
winter range for five radio-telemetered deer behind the Eagle 2.44-m fence
was 0.65 km (N=95, where N=the number of radio-locations and sightings).

�f

�-241-

AN EVALUATION OF DEER-PROOF FENCE LENGTH
REQUIRED TO PREVENT DEER MOVEMENT ON OR ACROSS HIGH SPEED HIGHWAYS
Thomas N. Woodard

D

P. S. OBJECTIVE
Measure effects of various lengths of 2.44-m fences on numbers of deer-vehicle
accidents on highwavs and deer movements in relation to fence length.
Effectiveness of fences will be evaluated on the basis of benefit:cost
ratios.

s.Er;MENTOBJECTIVES

,

1.

Determine the reductions in deer-vehicle accidents on highwavs,
the installation of various lengths of ?.44-m fence.

t

2.

Measure deer movements

after

in relation to 2.44-m fences.

DESCRIPTION

OF AREAS

Highway 82 - Diamond S
The Highway

82 study area has been described by Poi ar (1972).
Highway 82 - Carbondale

I

~

l

This studv area has been described by Reed (lQ75).
Interstate

70 - Avon, Edwards, Vail, and Eagle

The Interstate 70 Avon and Edwards studv areas have been described bv
Woodard (lQ73), the Vail study area by Reed (lQ7l), and the Eagle studv
area by Reed (1974).

METHODS AND MATERIALS
Methods and materials have been described by Poiar (1972).
Radio-tracking transmitter collars (Model MK 3, Telonics, 1048 E. Norwood,
Mesa, Arizona) were placed on five yearling or mature does trapped in
clover traps (Clover 1956) behind the Interstate 70 Eagle 7.7-km fence.
Attempts were made daily to locate these animals by establishing bearings
from three to nine permanent observation points. These bearings were plotted
on copies of a U.S. Geological Survey quadrangle map (scale 1:24,000).
Telemetered animal locations were estimated by determining the area where
three or more bearings converged.

�-242-

RESULTS AND DISCUSSION
Highway

82 - Diamond S

Deer concentrate in the crested wheatgrass fields northeast of the highway
in late winter and early spring (Table 1). The mean number of deer crossings
during March-May periods in 1971 before the 2.44-m fence was installed was
11.7 (N=32, where N-number of 24 hour periods) per day. After the fence
was installed the means were 2.1 (N=38) in 1972, 5.5 (N=34) in 1973, 10.3
(N=38) in 1974, and 17.8 (N=73) in 1975. This represents an 82, 53, and
12 percent decrease in crossings in 1972, 1973, and 1974, respectively,
and a 52 percent increase in 1975. The mean deer kill per year before
the 2.44-m fence was installed (October, 1968 - September, 1971) was 10.0.
The mean deer kill per year after fence installation (October, 1971 _
September, 1975) was 2.25; a reduction of 78 percent.

Table 1. The March-May mean number of deer observed on spotlight counts
between quarter-mile section markers 25 to 30 for 1968 through 1975 on
Highway 82 (n=number of counts).

Mean count
1971
1972

Month

1968

1969

1970

March

134.8
(n=4)

151.2
(n=4)

104.5
(n=4)

66.8
(n=4)

April

73.0
(n=4)

34.0
(n=5)

56.0
(n=S)

51.4
(n=5)

6.0
(n=5_

0.0

1.5

(n=L)

(n=Z)

0.0
(n=2)

66.2
(n=13)

77 .5
(n=10)

63.7
(n=ll)

47.6
(n=Ll )

May

MEAN
TOTAL

1973

1974

102.2
(n=4)

137.4
(n=5)

143.5
(n=4)

126.8
(n=4)

4.5
(n=4)

47 J)
(n=4)

52.3
(n=3)

93.0
(n=4)

53.4

97.2
(n=9)

104.4
(n=7)

109.9

(n=B)

(n=B)

Highwav 82 - Carbondale
The mean number of deer observed on spotlight counts during January - April
periods of 1968 through 1973 was 15.7 (n=99). Seventy deer were killed
from January, 1968 to April, 1973 (14 deer per year), before the fence was
installed.
Six were killed from 17 October 1974 to 17 October 1975 after
the 2.44-m fence was installed: a reduction of 57 percent.

�-243-

Interstate

70 - Avon

The mean number of deer observed on spotlight counts during the August-December
periods was 15.5 (n=17) and 13.4 (n=2l) in 1971 and 1972, respective Iv ,
10.8 (n=ll) in 1973 for October-December, and 7.3 (n=7) and 4.2 (n=R) for
November-December in 1974 and 1975, respectivelv.
Sixty-four deer made
passages to the north through six one-wav gates (Reed et al. 1974) located
in the 2.44-m fence during this segment. Twentv-eight deer were killed
from 5 October 1971 to 5 October 1972 before the fence was installed.
Thirteen (4.33 per year) were killed from 5 October 1972 to 5 October 1975
after the fence was installed: a reduction of 85 percent.
Interstate

70 - Edwards

Twenty-seven deer were killed on this segment of highwav from 5 October
1971 to 5 October 1972 before the fence was installed.
Thirty-one (10.37
per vear) were killed after fence installation from 5 October 1972 to
5 October 1975; a reduction of 62 percent.
~ive passages occurred through
four of six one-way gates located in the 2.44-m fence during this segment
Interstate
No additional
length.

70 - Vail

data were collected in this study area in regard to fence

Interstate

70 - Eagle

One hundred sixty-seven deer were killed from 5 October 1972 to 5 October
1973 before the fence was installed.
Sixty-three (31.5 per year) were
killed from 5 October 1973 to 5 October 1975 after the fence was installed;
a reduction of 81 percent. An estimated 70-80 deer, including several collared
animals, came across the highwav from the south and either 1) passed through
the one-way gates or an openi~g in the fence, 2) went around the ends of
the fence, 3) went back to the south to or across the Eagle River, or 4) were
killed on the highway.
~ourteen deer made passages to the north through
six of ten one-way gates located in the 2.44-m fence during this segment.
According to 92 observations of collared deer north of the 2.44-m fence,
does (N=20) and bucks (N=7) have moved laterally or parallel to the fence
for mean distances of 0.66 and 0.76 km, respectively.
~ive does (RT-l, RT-3, RT-4, RT-5, and RT-6) , banded with radio-transmitter
(RT) collars, were radio-tracked for 63, 56, 8, 35, and 5 days, respectively.
Plotted bearings indicated mean daily linear movements on the l.,interrange
of 0.70 (n=44), 0.75 (n=2l), 1.03 (n=3), 0.46 (n=24), and 0.40 km (n=3)
for RT-l, RT-3, RT-4, RT-5, and RT-6, respectively.
Doe RT-5 has been
within 60-70 m of the 2.44-m fence on at least four occasions.

�-244-

LITERATURE
Clover, M.R. 1956. Single-gate

CITED

trap. California

~ish and r,ame. 4~:lqq-~0l.

Pojar, T.M. 1972. An evaluation of deer-proof fence length required to
prevent deer movements on or across high sneed highways. Pages
305-310 in Game Research Report, July, Part 3. Colorado Div. Wildl.
Fed. Aid.
Reed, D.F. 1971. Deer underpass evaluation. Pages 341-351 in r,ame Research
Report, July, Part 3. Colorado Div. Wildl. ~ed. Aid.

----:,.---- . 1974. An evaluation of 8-foot fence length required to prevent
deer movements on or across high sneed highwavs. Pages 3l3-3~0 in
Game Research Report, Julv, Part 2. Colorado Div. Wildl. ~ed. Aid.
_____

, T.M. Poiar, and T.N. Woodard. lQ74. Use of one-wav gates bv
mule deer. J. Wildl. Manage. 38(1):9-15.

_____

. 1975. An evaluation of 8-foot (2.44-m) fence length required to
prevent deer movements on or across high speed high~.:ravs.Pages 315-320
in Game Research Report, Julv, Part 2. Colorado Div. Wildl. ~ed. Aid.

Woodard, T.N. 1973. An evaluation of deer-proof fence length required to
prevent deer movements on or across high sneed highwavs. Pages 197-202
in Game Research Report, July, Part 2. Colorado Dfv , TATildl.Fed, Aid.

/1

prepared by

~~a~

.1/ tlJJa£

Thomas N. Woodard
Assistant Wildlife

Researcher

I

•

�-245-

July, 1976

JOB FINAL REPORT
State of

COLORADO
----------~---------

Project No.

W-125-R-2

Work Plan No.

15

Period Covered:

April 1975 - March 1976

Deer-Auto Accident

Investigations

11
Job No.
Effects of a Simulated 8-Foot Fence
Job Title Angle in Diverting Deer from Their Direction of Movement

Personnel:

D. F. Reed and T. N. Woodard

ABSTRACT
This job was terminated one year earlier than anticipated, and therefore,
the 1974-75 Job Progress Report was written as a Job Final Report and
summarized all activities for all years this job was active.

�J

�-247-

July, 1976

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~-----------

Project No.

lJ-125-R-2

Work Plan No.

15

Job Title

Job No.

~1~4

Evaluation of Deer Overpasses

Period Covered:
Personnel:

Deer-Auto Accident Inyestigations

April 1, 1975 through March 31, 1976

Claudia A. Doose, James D. Fleming, Thomas N. Woodard, and
Dale F. Reed.

ABSTRACT
Ninety-nine, 93, and 77 deer crossings over a sub-standard 3.2- X 4.9- X
l3.4-m, height (under-bridge clearance) X width X length (direction of
traffic), bridge overpass were recorded by a video system during the fall
of 1974 and spring-summer and fall of 1975, respectively.
Video replay
of 84, 67, and 61 crossing modes yielded walk:trot:bound ratios of
7.8:1.8:1.0, 7.6:4.8:1.0, and 4.6:2.0:1.0 for the fall of 1974 and springsummer and fall of 1975 migration periods, respectively.
No look-up or tailup responses were detected.

_

��-249-

EVALUATION

OF DEER OVERPASSES
Dale F. Reed

There are areas in Colorado where migratory deer cross highways.
When
these highways carry high-speed, high-volume traffic, the potential
danger to deer and motorists becomes especiallv great. Therefore, it is
essential to keep the animals off the highway.
Overpasses with specified
dimensions and characteristics located at strategic points within migration
routes may provide adequate means for these necessary deer movements.
This
study purports to evaluate overpasses of different sizes and design and
to further evaluate deer behavioral responses.

P. S. OBJECTIVE
To test the effectiveness of deer overpasses in reducing deer-vehicle
collisions on high-speed high-traffic volume highwav systems.

SEr.ID.:NT
OBJECTIVE
1.

Determine
areas.

the use of overpasses bv deer in critical highwav deer kill

2.

Compare detailed deer behavioral responses to overpasses after moderate
to high use of the structures or immediate areas has been documented.

DESCRIPTION

O~ AREA

The Dowd deer overpass is located in Eagle County 7.5 km west of Vail,
Colorado, adjacent Interstate 70. The structure spans Gore Creek and was
used by vehicles before the completion of Interstate 70 (1-70) .. It is
just north of the 1-70 right-of-way on the Dale Mikkelson property.
Approximately 3.0 miles of associated 2.44-m fenCing generally parallels
the highway in both directions east of the overpass.
Deer apparentlv
move around the west ends of this fenCing and cross the bridge overpass
rather than Gore Creek.

METHODS AND MATERIALS
The study utilized a sub-standard bridge 3.2- X 4.9- X l3.4-m, height
(under-bridge clearance) X width X length (direction of traffic), over
Gore Creek. Approaches to the overpass were checked for deer tracks
during spring-summer (Jun-Jul) and fall (Oct-Nov) migration periods.
A
video time-lapse surveillance system (Reed et al. 1973) was used to record
imagery of crossings and overt behavioral responses.
It was activated
for 30 to 35 davs from approximatelv 2ln0 to 0730 and 1900 to 0730 for
the Jun-Jul and Oct-Nov periods, respectively.
The number of video-recorded

�-250-

1/
2/
3/
deer approaches~ entrances~ exits~ and behavioral responses including
muzzle-to-ground (Reed et al. 1975:366), hesitation (cessation of forward
movement for 1.0 second or more), and crossing mode (walk, trot, or
bound) were tallied during video tane reu Lav ,

R~SULTS AND DISCUSSION
Ninety-nine, 93, and 77 deer crossings Over the deer overpass were recorded
by video system for the fall of 1974 and spring-summer and fall of 1975,
respectively.
Replay of 84, 67, and 61 crossing modes vie1ded wa1k:trot:bound
ratios of 7.8:1.8:1.0, 7.6:4.8:1.0, and 4.6:2.0:1.0 for the fall of 1974
and spring-summer and fall of 1975 migration periods, respectively (Table 1).
An additional 27 animals changed their behavior (i.e. walk to trot, walk to
bound, trot to bound, trot to walk, and trot to walk to trot) during the
process of crossing during the spring-summer period. One deer wal.kad half-way
across (entered) the structure, turned around and bounded (exited) out the
entrance. No look-up or tail-up responses were detected for the ~69
crossings. This is of importance when compared to the response frequencies
observed at the Vail deer underpass (Reed et al. 1975). Although preliminary,
these results indicate deer exhibit only slight reluctance in crossing an
overpass of this size and design.

LITERATURE

CITED

Reed, D.F., T.M. Pojar, and T.N. Woodard. 1973. A video time-lapse system
for wildlife surveillance. Colorado Div. Wild1., r,ame Inf. Leafl. 94. 3pp.

----,--- , T.N. Hoodard, and T.11. l'o;ar. 1975. Behavioral response of mule
deer to a highway underpass. J. Wildl. 11anage. 39(7.):361-376.

~(24-l~~~\-_""'_

Prepared bv _~-L.J,.:::L.J.&lt;L.::lo.r.I!.-&lt;-::o-_- =L-4-~
Dale F. Reed
Wildlife Researcher

1/ Approach

denotes a deer coming into the overnass entrance area covered
by the surveillance camera and either going onto or turning and
leaving the structure.

~/ Entrance denotes a deer coming onto the overpass at the entrance.

1/ Exit denotes a deer leaving the overpass entrance, having either turned
around on the structure after having made an entrance, or made a passage
in the opposite direction of the migration.

•

�-251-

Table 1. The number of muzzles-to-ground, hesitations, and crossings at
the Dowd deer overpass for the fall migration of 1974 and spring-summer
and fall migrations of 1975.
1974
Fall

Spring-Summer

",all

25

38

26

38

64

38

No. deer

19

26

47

Frequency

26

38

90

Time (sec ,)

24.2

11. 2

12.0

walk

62

38

37

trot

14

24

16

bound

8

5

8

14.8

18.7

93

77

~uzzle-to-ground

1975

~I

No. deer
Frequency

'E...I
Hesitations

Crossing
Mode

Time
Total crossings

99

~I
~uzz1e-to-ground denotes a deer lowering its muzzle to the ground. It
has been surmised this response was an olfactory examination of the surface
environment including a possible scent left by the interdigital glands
of other deer (Reed et a1. 1975:365).

'£.1
Cessation of forward movement for 1.0 second or more.

�· ~

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                  <text>-253-

July, 1976

JOB FINAL REPORT

State of

Colorado
-----------------

Project No.

Deer-Elk Investigations

W-38-R-3l

vlork Plan No. __1_4

_

5

Job No.

Job Title Middle Park Cooperative Deer Study-Experimental
Period Covered:
Personnel:

Range Fertilization

April 1. 1969 through March 31, 1976

L. H. Carpenter, O. C. Wallmo, D. W. Reichert, W. L. Regelin, R. B.
Gill, D. L. Baker, P. F. Gilbert, L. A. Roper, D. Benson, W.
Paintner,R. Davies, C. Grand Pre, R. Glaze, L. Brown, R. Hoffman,
and J. Wassink.
ABSTRACT

Three areas located on big sagebrush range in Middle Park, Colorado were
studied to evaluate the effect of ammonium nitrate fertilizer applied with and
without 2,4-D herbicide on herbage yields and botanical composition.
Nitrogen
was applied at five levels: 0, 33.6, 67.2, 100.8, 134.4 kg/ha (0, 30, 60, 90,
120 lb/ac) and 2,4-D herbicide at two levels: 0, 2.2 kg/ha (0, 2lb/ac).
Estimates of botanical composition were made in July and herbage yield in August
each year for three consecutive years following treatment. Measurements of
grazing use by tame, trained deer were made each winter on the various treatments. A fourth study area was added to evaluate deer preferences for fertilizer
treated plants and to test the effects of ammonium nitrate fertilizer 134.4 kg
N/ha (120 lb/ac) and magnesium sulfate at a rate of 56 kg Mg/ha (50 lb/ac) on
total volatile oil content of big sagebrush. Yields of total herbage and shrub
herbage increased significantly with increasing levels of nitrogen fertilizer
each year. Nitrogen fertilizer did not consistently increase forb and grass
yields. The highest nitrogen level, 134.4 kg N/ha (120 lb/ac) was most efficient
in terms of increasing the kg yield of herbage per kg N applied. Nitrogen did
not affect the numbers of forb, shrub, or grass plants. The herbicide reduced
shrub herbage, total herbage, and forb yields in all years. Grass yields increased in all years on the herbicide treated plots.. Herbicide treatments also
significantly reduced numbers of big sagebrush plants, total shrubs, phlox,
bluebell, and total forb plants. Nitrogen-herbicide treatments increased yields
of total herbage each successive year, even though shrub herbage remained about
the same each year. Ammonium nitrate and magnesium sulfate, each increased
volatile oil content of big sagebrush plants; Big sagebrush plants that were
heavily grazed by tame mule deer had a significantly lower total oil content
than other big sagebrush plants sampled. The deer did not show a significant
preference for plants in anyone individual treatment,
Contrary to general belife, grasses consistently made up over 60 percent of deer diets during winter
grazing trials.
A disseration entitled "Nitrogen Herbicide Effects on Sagebrush Deer Range"
was written and published for the purpos~ of acquiring a Ph.D and will serve
as the final report for this job. Copies of thesis are on file at Colorado
State University Library and the Library of the Colorado Division of Wildlife
Research Center, Fort Collins. Appropriate portions are being printed in part
for publication.
Sufficient copies of published materials will be forwarded
to the Fish and Wildlife Service when available.
\

.

�-254-

RECOMMENDATIONS
1.

On areas critical to mule deer in winter, where it is desirable to
increase forage yields, a single application of nitrogen fertilizer
should be applied.
To increase longevity of the yield response,
nitrogen should be applied at the rate of 120 pounds actual nitrogen
per acre (133.4 kg N/ha).

2.

On areas dominated by sagebrush (Artemisia sp.) nitrogen fertilizer
should be applied with a treatment of 2,4-D herbicide to increase
nitrogen availability to plants other than sagebrush.
Based on the
estimations of food preferences of mule deer made in this study increased grass production resulting from the fertilizer-herbicide
combination treatment will be beneficial to mule deer.

3.

The possibility that nitrogen fertilizer may concentrate grazing
animals requires consideration of alternative application procedures.
If large areas (over 20 acres; 8.1 ha) are treated the fertilizer
should be applied in strips or in a "checkerboard" pattern.

4.

Nitrogen should be applied in the fall (late October).
Winter snows
will cover the fertilizer and insure proper soil infiltration.

5.

Nitrogen should be applied with a fixed-wing aircraft where possible,
to minimize application costs.

Prepared by __

,{..:..~--=-.;::..-:../t_·_C~'a~,./fl,_
..
cf;:--·

Len H. Carp~
Asst. Wildlife Researcher

_

�July •.1976

-255JOB FINAL REPORT

State of

--------- COLORP~O
--------------

Project

No.

W-38-R-31

Deer-Elk

14

Work Plan No.

Job No.

Job Title Middle Park Cooperative
Period Covered:

5a

---------------------------

Deer Study - Experimental

Range Fertilization

April 1, 1973 through March 31, 1976

Len H. Carpenter

Personnel:

Investigations

and D. L. Baker

ABSTRACT
A draft of a manus crLp't entitled "Effects of Varying Applicate Dates of
2,4-D on Mule Deer Winter Range" is in preparation and will be submitted
to the Journal of Range Management for publication.

Prepared

by

L//cov-~

---L-e-n--H-.~C~a~r-p-e~n-t-~~r----------Asst. Wildlife

Researcher

��July, .1976

-257JOB FINAL REPORT

State of

Colorado

Project No.

W-38-R-3l

Work Plan No.

14
--------.---------

Deer-Elk Investigations
Job No.

8

Job Title Middle Park Deer Study - Experimental Harvest Regulations
Period Covered:
Personnel:

April 1, 1975 - March 31, 1976

D. Freddy, L. Carpenter, D. Bowden, K. Karrow, S. Koster, P.
Powell, W. Adrian, H. Riffel, D. Clippinger, B. McCloskey, D.
Luce, J. Wolfe, D. Bartmann, and D. Mueller.

ABSTRACT
Estimated deer density in Middle Park in January, 1976 was 7.958 + 1.207
per square mile of winter range yielding a projected population estimate of
4677 ± 1181 (P~.lO).
Pre-natal productivity was 156 fawns:lOO does. Net
productivity based on post-hunting season classification counts was 70 fawns:
100 does. Winter mortality was 149 deer or 2.5 percent of the January, 1975
population estimate. Total harvest during 1975, including a 20 percent
wounding loss was 1114 deer, 30 percent below the harvest objective of 1600
deer. Success of all hunters in 1975 was 15.1 percent. An estimated 82.7
percent of the permitted hunters participated in the 1975 regular season.
Restricting antlered-only permittees to Middle Park appeared to increase
success ratios. Percent of antlered-only permittees participating during
regular seasons averaged 80.6 while percent of either-sex permittees participating averaged 86.6. Participation by late-season either-sex permittees
averaged 75.4 percent. Either-sex permittees harvested 63.4 and 75.4 percent
antlerless animals during regular seasons and 38.6 and 50.6 percent antlerless
during late-seasons.
There was no significant difference between composition
of deer passing through check station and composition of deer reported harvested on returned hunter surveys (X2.05,3). Failure to adjust the Middle
Park deer population to a desired density on the winter range can be attributed to the inability to consistently estimate and predict population size
and inconsistent policies governing issuance of permits and restrictions on
permits.

�-258-

RECOMMENDATIONS
1.

Prerequisite to evaluating whether 'a specific number of deer can be
harvested is the ability to consistently estimate and predict population size. The objective of harvesting deer to maintain a herd at a
specific density is futile if the population size cannot be estimated.

2.

Research involved with evaluating sport-hunting must remain insulated
from the political arena of wildlife management. Once procedures are
established, political whims must not be allowed to alter the research
process, because to do so obviates meaningful between year comparisons.
Research must maintain total control over permit allocation, distribution, and issuance to insure consistency between years of a study.

3.

Research must maintain total control over hunter survey procedures,
especially sample sizes, permit listings, timing, data summary, and
analysis. To be comparable, methods for estimating yearly harvests
must be consistent for all years.

4.

Both antlered-only and either-sex permits should be utilized in harvest
studies to induce sufficient hunter participation and to improve precision
limits of harvest estimates.

5.

Permits for late-season hunts should restrict hunters to one Game Management Unit to control distribution of the harvest.

6.

An average of approximately 1000 deer were harvested in Middle Park
during regular seasons with an either-sex hunting season. If more than
1000 deer need to be harvested to attain a desired population density,
then a late-season will probably be needed. Probably not more than 2-3000
deer can be harvested in Middle Park without the use of multiple-deer
licenses. Population size should not be allowed to increase above the
level of potential harvest.

�-259-

MIDDLE PARK DEER STUDY EXPERIMENTAL HARVEST REGULATIONS
Dave Freddy

P. S. OBJECTIVE
To adjust a mule deer population to a specified density on a winter range.

METHODS AND MATERIALS
Methods for estimating deer density, composition, productivity
mortality have been ~eported by Gill (1969a, b, 1971).

and winter

Methods for estimating harvest, success of hunters, and hunter participation
have been reporten by Roper (1973,1974) and Freddy (1975). Exceptions are:
1.

A mail questionnaire was sent to 60 percent of the regular season
antlered-only permit holders to estimate the antlered-only harvest
in 1975.

2.

A mail questionnaire was sent to all regular season either-sex permit holders to estimate the either-sex harvest during 1975. This
survey was part of the state-wide survey of either-sex hunters conducted by the Denver Office, Colorado Division of Wildlife (DOW).
Research personnel did not conduct this survey because of difficulties in determining the total listing of Middle Park either-sex
permittees arising from issuing part of the either-sex permits on a
first-come basis to persons having antlered-only licenses.

Methods for determining if specified permits are capable of achieving predetermined harvest goals have been reported by Roper (1973,1974).
Methods for scheduling and publicizing season recommendations
ported by Roper (1973,1974).

have been re-

RESULTS AND DISCUSSION

Population Characteristics

1975

Population Density and Size
The size of the Middle Park deer population was estimated from helicopter
counts of deer on square-mile sample quadrats from January 7-9,1976
(Table 1).
The estimated deer density of 7,958 + 1.207 per square mile yielded a projected
population estimate of 4677 ± 1181 (P&lt;.lO).
The 1976 population estimate was
not significantly different than the January, 1975 estimate of 6005 + 1714
(P &lt; .10) (Table 2) when they were tested with a paired t-test.
-

�Table 1. Deer counted per quadrat within eight sampling strata in four deer population sub-units, Middle Park, Colorado, January
1972-1976.
Quadrat
No.

1972

MuddI Creek High DensitI Strata
1973
1974
1975
1976

Quadrat
No.

1972

Muddl Creek Low DensitI Strata
1973
1974
1975

1976

1M

0

0

0

0

0

11M

0

0

0

0

2M

0

0

0

0

0

14M

2

0

4

0

0

3M

0

2

0

0

0

20M

0

0

0

0

0

4M

0

0

0

0

0

23M

0

6

0

0

0

5M

0

0

0

0

0

24M

0

0

0

0

0

6M

1

0

0

0

10

25M

0

0

0

0

0

7M

8

40

13

1

7

26M

0

0

0

0

0

8M

11

60

25

47

9

27M

0

0

0

0

0

9M

24

132

125

86

23

28M

0

0

0

0

1

I
IV

10M

39

97

52

77

14

29M

0

0

1

0

3

0
I

12M

0

52

54

5

0

30M

0

0

7

0

2

13M

5

46

66

0

0

31M

6

0

0

0

5

15M

35

42

32

4

0

16M

6

0

0

0

16
19

8

6

12

0

18

0.7

0.5

1.0

0.0

1.5

17M

1

0

0

0

IBM

1

0

0

0

0

19M

0

0

0

0

0

0

0

0

21M

0

3

22M

0

3

3

0

17

32M

27

3

0

0

9

158

480

370

220

124

24.0

18.5

11.0

I:yi

y

c

=

7.9

6.2

------------------------------------------------------------------------------------------------------------------------------------------

'"

�Table 1.
1972-1976

Deer counted per quadrat within eight sampling strata in four deer population

sub-units, Middle Park, Colorado, January

(continued).
Blue River High Density Strata
1975
1974
1973
1972

Quaurat
No.

1976

Quadrat
No.

1972

Blue River Low Densit~ Strata
1975
1974
1973

1976

44

2B

0

0

0

14

5

0

9

25

0

1B

0

3B

0

9

0

0

4

0

4

0

4

4B

0

0

5B

0

0

1

63

36

6

6

3

7B

48

19

6B

0

0

10

65

78

19

50

90

8B

47

12B

30

66

128

52

24

156

87

82

19

9B

0

16B

0

0

0

0

0

28

0

41

0

lOB

0

0

0

0

0

29

11B

9

I

12

6

0

17B

47

19B

2

24

91

27

13

21B

0

3

0

0

1

13B

15

78

48

14B

22

74

50

48

0

15B

64

50

79

31

13

18B

52

102

65

38

21

20B

0

2

2

11

12

256

203

32

50

275

100

637

499

139

339

21.3

16.9

3.6

5.6

30.6

11.1

53.2

41.6

15.4

28.3

y

=

a•...
I

17

l:yi =

N

-------------------------------------------------------------------------------------------------------------------------------------------

�Table 1. Deer counter per quadrat within eight sampling strata in four deer population sub-units, Middle Park, Colorado, January
1972-1976 (continued).
Wms. Fork River High Densitl Strata
1975
1974
1972
1973

Quadrat
No.

Quadrat
No.

1972

6W

0

0

3

0

0

7W

0

0

0

5

0

8W

0

0

0

0

3

9W

0

0

0

0

0

lOW

0

0

0

0

0

1W

9

0

12

0

0

2W

0

0

0

0

0

3W

52

45

60

35

45

4W

71

54

17

43

12

0

5W

0

15

1

Wms. ~ork River Low Densitl Strata
1975
1974
1973

1976

1976

8
I
N

aN

I

l:yi =
y

=

132

99

104

79

65

0

0

3

5

3

26.4

19..8

20.8

15.8

13.0

0.0

0.0

0.6

1.0

0.6

------------------------------------------------------------------------------------------------------------------------------------------

�Table 1. Deer counted per quadrat within eight sampling strata in four deer population sub-units, Middle Park, Colorado, January
1972-1976 (continued).
Granb~ Low Densit~ Strata
1975
1974
1973

1976

0

0

0

0

0

0

0

1

11

20

0

0

4G

0

0

0

0

10

8

5G

0

0

0

0

0

17

51

6G

0

0

0

11

0

14

0

50

7G

0

0

0

0

0

31

41

21

50

0

21

1

0

0

12

33

16

17

5

1:yi =

219

365

296

270

227

2

14

20

11

10

-y

21.9

36.5

29.6

27.0

22.7

0.3

2.0

2.9

1.6

1.4

Troublesome Cr. High Densit~ Strata
1975
1973
1974

1976

Quadrat
No.

1972

15

21

0

1G

1

3

40

41

46

6

2G

0

2

2

0

0

0

3G

4T

70

70

54

63

57

5T

20

55

30

85

6T

40

61

84

7T

27

31

8T

21

9T
lOT

Quadrat
No.

1972

IT

3

21

2T

24

3T

I
N

a-

w

=

I

1972

1973

1974

1975

1976

=

9.24

17.14

16.455

10.217

7.958

2S (yst) =

1.96

5.24

6.93

3.086

1.456

sy

1.40

2.29

2.632

1. 757

1.207

yst

=

�-264-

Table 2. Deer population
January 1972-1976.

estimates by sub-unit

POEulation Sub-Unit
Wms. Fork
Troub les ome
River
Creek

Year

Muddy
Creek

Blue
River

1972

1,002

2,259

742

1973

2,856

4,186

1974

2,272

1975
1976

in Middle Park, Colorado,

Granby

Total
Population

1,404

22

5,429 ± 1,913 l/

556

2,340

149

10,087

4,663

622

1,897

216

9,670 ± 2,566

1,281

2,370

506

1,729

119

6,005 ± 1,714

893

1,824

402

1,455

103

4,677

± 2,239

± 1,181

1./
(P~

.10)

Sex and Age ComEosition
No pre-season sex and age classification counts were conducted.
The 1969-74
average of .83 fawns/doe for pre-season counts was utilized for life-table
purposes (Table 3). Post-season classification counts were conducted by helicopter from December 5-7, 1975. Two observers classified 1415 deer resulting
in a buck:doe:fawn ratio of 44:100:70.
Post-season classification counts
were also conducted from the ground, yielding a buck:doe:fawn ratio of
55:100:47 based on 723 deer (Table 4). The helicopter estimate was considered
more reliable because subjective evidence indicated deer may have been incorrectly classified during the latter portion of ground surveys.

Table 3. Pre-hunting season deer sex and age clas,ification counts obtained
from helicopter surveys during October 1969-1974 1, Middle Park, Colorado.

Year

Bucks

Does

Fawns

Total
Classified

1969

195

451

419

1,065

43

93

1970

152

361

258

771

42

72

1971

20

41

30

91

49

73

1972

73

101

91

265

72

90

1973

174

314

273

761

55

87

1974

58

76

91

225

76

120

l/No counts in 1975.

Ratios Eer 100 Does
Bucks
Fawns

�-265Table 4. Post-hunting season deer sex and age classification counts
obtained from helicopter and ground surveys during November and/or December,
1967-1975, Middle Park, Colorado.

Year

Type of
Count

Bucks

Does

Fawns

Total
Classified

1967

Aerial

316

558

501

1,375

57

90

Ground

147

396

363

906

37

92

Aerial

261

490

418

1,169

53

85

Ground

319

582

505

1,406

55

87

Aerial

211

471

363

1,045

45

77

Ground

104

231

178

513

45

77

Aerial

-------------------

Ground

356

783

321

1,460

45

41

Aerial

218

398

244

860

55

61

Ground

276

390

261

927

71

67

Aerial

362

693

540

1,595

52

78

Ground

818

1,281

1,047

3,146

64

82

Aerial

396

674

594

1,664

59

88

Ground

-------------------

Aerial

309

632

476

1,417

49

75

Ground

194

363

236

793

53

65

Aerial

291

663

461

1,415

44

70

Ground

197

357

169

723

55

47

1968

1969

1970

1971

1972

1973

1974

1975

Ratios Eer 100 Does
Bucks
Fawns

No Counts -----------------------

No Counts -----------------------

Productivity
Pre-natal productivity was estimated as 156 fawns:100 does during 1975. This
estimate was based on 14 fetuses present in 9 mature road-killed does collected
from January-May, 1975 (Table 5). Net productivity was estimated to be 70 fawns:
100 does as of December, 1975.

Winter Mortality
Estimated over-winter mortality
deer (Table 6). This projected
mortality transects.

for 1975 was 149 deer or 2.5 percent of 6,005
estimate was based on 6 dead deer found on 93

�-266There was also record of 44 deer dying from deer-auto collisons and 28
deer dying from deer-train collisons from January-May, 1975. These are
minimal estimates of deer dying from these causes.

Table 5. Fetuses per 100 mature female deer in Middle Park, Colorado,
January-May 1969-1975.
Deer collected by selective shooting, from highway
accidents, and from trapping accidents.
Number Collected
Highway

Trapping

Total
Deer

Fetuses Per
100 Females

8

0

41

193

33

9

0

42

164

1971

33

8

10

51

161

1972

32

7

0

39

187

1973

0

26

0

26

162

1974

0

7

0

7

157

1975

0

9

0

9

156

Year

Shooting

1969

33

1970

Table 6. Projected winter mortality for deer in Middle Park, Colorado 19721975 based on dead deer found on 93 randomly located 1 x .10 mile mortality
transects.

Year

Number Dead
Deer Found

Projected Number
Dead Deer

Percent of
Population Estimate

1972

2

50

0.9

1973

23

647

6.4

1974

15

420

4.3

1975

6

149

2.5

�-267Harvest Mortality

Harvest Objective and Permit Numbers--The harvest objective for 1975 was
1600 deer, inclusive of an arbitrary 20 percent inflation factor for
wounding loss (Table 7). This objective was based on the January, 1975
estimate of 6005 deer, a calculated life-table pre-season population estimate of 8539 deer and the desire to increase the population by 1000 to
approach the objective of 10,000 deer post-season.
Archery, high-country
bucks, and regular deer season hunters were required to obtain a permit
prior to hunting in Middle Park. Only regular season either-sex permittees
were restricted to hunting exclusively in Middle Park. Either-sex permits
were allocated by Game Management Unit (GMU) and either-sex hunters were
restricted to one unit. DOW management personnel initiated a state-wide
policy in 1975 requiring all either-sex permits to be allocated by GMU.
Prior to this decision, either-sex permits issued for Middle Park in previous years were valid in any of 4 GMU's.

Table 7. Deer pre-season population sizes and harvest objectives, Middle
Park, Colorado, 1972-1975. Harvest includes regular, late, archery and
high-country buck seasons.

Year

October PreJanuary
Season Pop.
Census Est. (Life-table)

Harvest
Harvest
Objective 1/ Estimate
w/20% W.L.- w/20% W.L.

Percent Harvest
Objective
Attained

1972

5,429

8,057

Small Harvest

427

1973

10,087

l4,l3l

4,000

2,376

59.4

1974

9,670

13,497

3,400

2,319

68.2

1975

6,005

8,539

1,600

1,114

69.6

JJ

W.L.

Wounding loss.

Of the permits issued, 586 were for archery, 10 for high-country buck, and
7076 for the regular season. Of the regular season permits, 4991 were
antlered-only and 2085 were for either-sex.
A total of 2750 either-sex permits were allocated for Middle Park: 1000 for Unit 18, 886 issued; 500 for
Unit 27, 260 issued; 500 for Unit 28, 351 issued; and 750 for Unit 37, 588
issued. A total of 1260 either-sex permits were issued by a random drawing
held in June at the Denver Office, DOW, while 825 were issued on a first-come
basis at the Grand Junction Regional Office, DOW, from August 1 - October 24,
1975.

�-268Harvest Size and Composition--Harvest size and composition were estimated
from survey questionnaires returned by hunters. A total of 1732, or 83
percent, of 2085 season either-sex questionnaires sent were returned. A
total of 1799, or 60 percent, of 2992 antlered-only questionnaires sent
were returned. Archery and high-country buck hunters were surveyed by
state-wide hunter surveys and harvest estimates 'were 44 and 0 deer, respectively.
Archery hunters accounted for 4.7 percent of the total harvest
(Table 8).

Table 8.

Summary of deer harvested

Type of
Season

Bucks

Does

Buck
Fawns

fue
Fawns

Total

Percent
Total Harvest

Archery

29

15

0

0

44

4.7

0

0

0

0

0

0.0

485

228

80

91

884

95.3

High-country
Regular Deer

Totals

Buck

514
243
(55.4%) (26.2%)

in Middle Park, Colorado, 1975.

80
(8.6%)

91
928
(9.8%) (100.0%)

100.0

The projected 1975 regular season harvest based on returned questionnaires
was 884 deer, or 95.3 percent of the total harvest (Table 8). Either-sex
permit holders harvested 509 + 18 while antlered-only hunters harvested 375 +
60 deer (P&lt; .05) (Tables 9, 10). The projected composition of the total
harvest was-54.9 pe,~cent bucks, 25.8 percent does, 9.0 percent buck fawns
and 10.3 percent doe fawns (Table 11).
During the regular season, 478 deer harvested in Middle Park were checked at
Idaho Springs check station west of Denver. The composition of these deer
was 49.8 percent bucks, 32.0 percent does, 6.7 percent buck fawns, 8.6 percent doe fawns, and 2.9 percent unknown (Table 12).
An age pyramid derived from check station returns and mail-in tooth envelopes
(sent with either-sex permits) indicated a young, productive population (Odum
1959:173, Kormondy 1969:81), assuming the harvest reflects the age distribution of the live deer population. Yearlings comprised 35.9 percent, 2 yearolds 18.9 percent, and fawns 19.3 percent of the harvest (Fig. 1). The lower
percentage of fawns as compared to 1 year-olds is assumed to reflect hunter
selection against fawns. The fawn harvest was considerably higher than in
1974 when fawns comprised 7.3 percent of the harvest. This increase may have
been due to the increased number of either-sex permits, high fawn production
or vulnerability, lack of adult animals, or lack of hunter discretion.

�-269Table 9. Composition of the projected regular season either-sex permit
deer harvest, Middle Park, Colorado, 1974-1975, based on returned hunter
survey questionnaires.
Unit
28

37

13

9

17

66

42

13

18

24

96 ( 53.4%)

B. Fawns

5

1

1

4

12

(

6.2%)

D. Fawns

4

2

3

0

9

(

4.8%)

Unknown

1

0

0

0

1 (

0.7%)

Totals

79
(42.8%)

29
(15.8%)

31
(16.6%)

45
(24.8%)

1975

18

Management Unit
28
27

37

Totals

Bucks

37

24

12

52

125 ( 24.6%)

Does

96

34

32

60

222

B. Fawns

21

17

8

25

71 ( 14.0%)

D. Fawns

34

17

8

32

91 ( 17.8%)

Totals

188
(37.0%)

92
(18.0%)

60
(11.8%)

169
(33.2%)

509 (100.0%)
(100%)

1974

18

Management
27

Bucks

27

Does

Totals
(

35.9%)

184 (100.0%)
(100%)

(

43.6%)

Table 10. Composition of the projected regular season antlered-only permit
deer harvest, Middle Park, Colorado, 1975, based on returned hunter survey
questionnaires.
1975

18

Management
27

Unit
28

37

Totals

Bucks

94

102

47

117

361 ( 96.3%)

Does

3

3

0

0

6 (

1.5%)

B. Fawns

3

3

3

0

8 (

2.2%)

Totals

100
(26.7%)

50
(13.3%)

117
(31.1%)

108
(28.9%)

375 (100.0%)
(100%)

�-270Table' 11. Composition of the
total projected regular season deer harvest,
Middle Park, Colorado, 1972-1975 based on returned hunter survey
questionnaires.
Management
1972

18

27

Unit
28

37

Totals

Bucks

127

84

86

59

356 (100.0%)

Totals

127
(35.6%)

84
(23.6%)

86
(24.2%)

59
(16.6%)

356 (100.0%)
(100%)

Management
1973

Unit

18

27

28

37

Bucks

159

155

36

204

555

(

56.3%)

Does

91

71

50

149

360

(

36.6%)

B. Fawns

8

12

6

6

32 (

3.2%)

D. Fawns

18

6

0

14

38 (

3.9%)

276
(28.0%)

244
(24.8%)

92
(9.3%)

373
(37.9%)

Totals

Totals

985 (100.0%)
(100%)

18

Management
27

Uriit

1974

28

37

Totals

Bucks

139

77

116

81

413 ( 76.3%)

Does

42

'13

18

29

102 ( 18.9%)

B. Fawns

5

6

1

4

16 (

2.9%)

D. Fawns

4

2

3

0

9

(

1.7%)

Unknown

1

0

0

0

1 (

0.2%)

191
(35.3%)

98
(18.1%)

138
(25.5%)

ll4
(21.1%)

Totals

Management

541 (100.0%)
(100%)

Unit

1975

18

27

28

37

Totals

Bucks

131

126

59

169

485 ( 54.9%)

Does

99

37

32

60

228 ( 25.8%)

B. Fawns

24

20

II

25

80 (

D. Fawns

34

17

8

32

91 ( 10.3%)

Totals

288
(32.6%)

200
(22.6%)

llO
(12.4%)

286
(32.4%)

884 (100.0%)
(100%)

9.0%)

�-271-

Table 12. Composition of deer harvested in Middle Park, Colorado, and
checked at Idaho Springs check station, 1973-1975.

1973

18

27

Bucks

53

36

Does

24

B. Fawns

Management Unit
28

37

Totals

20

50

159 ( 51.8%)

35

15

35

109

(

35.5%)

4

3

2

10

19

(

6.2%)

D. Fawns

5

5

4

4

18

(

5.9%)

Unknown

0

0

0

2

2 (

0.6%)

Totals

86
(28.0%)

79
(25.7%)

41
(13 .4%)

1974

18

27

Bucks

60

63

26

56

205

(

75.1%)

Does

19

4

9

17

49

(

18.0%)

B. Fawns

6

1

0

1

8 (

2.9%)

D. Fawns

4

2

2

0

8

(

2.9%)

Unknown

3

0

0

0

3 (

1.1%)

Totals

92
(33.7%)

70
(25.6%)

37
(13.6%)

74
(27.1%)

Management Unit
28

101
(32.9%)

307 (100.0%)
(100%)

37

Totals

273 (l00 .0%)
(100%)

Management Unit
1975

18

27

28

37

Totals

Bucks

57

61

34

86

238 ( 49.8%)

Does

56

38

20

39

153

(

32.0%)

B. Fawns

13

6

2

11

32 (

6.7%)

D. Fawns

18

8

2

13

41 (

8.6%)

Unknown

7

4

1

2

14

(

2.9%)

Totals

151
(31.6%)

117
(24.5%)

59
(12.3%)

151
(31.6%)

478 (100.0%)
(100%)

�-272BUCKS n = 271

DOES

n = 189

o.~

0••
41&gt;

0.4%
0.41&gt;

o.~
2.4%

1------------------- 18.9%
------------ 35.9%
1------------ 190)%
percent

50

40

30

20

10

o

10

20

)0

40

50 percent

Fig. 1. Age distribution
of deer harvested in Middle park, Colorado during
the regular season, October 25-November4, 1975. year classes 2 through 1)
were aged by dental cementum,while 82 percent of the fawns and 88 percent of
the yearlings were aged by tooth replacement and wear. Teeth obtained from
hunter mail-in tooth envelopes and check stations.

�-273Hunter Participation and Success--Hunter participation and success were
estimated from ~eturned hunter survey questionnaires.
Of 2085 potential
either-sex hunters, 1791 ± 27 (P~ .05), or 85.9 percent, hunted during
the regular season. The success rate for all either-sex permittees was
24.4 percent and for those that hunted, 28.4 percent (Table 13).
Of the 4991 potential antlered-only hunters, 4058 ± 90 (P~ .05), or 81.3
percent, hunted during the regular season in Middle Park. All ant1eredonly permit holders had a success rate of 7.5 percent while the success
of those that hunted was 9.2 percent (Table 13).

Fou~ding Loss--Hunters having antlered-only permits were asked to specify
on survey questionnaires the number of deer they wounded. Based on hunter
responses, wounding loss was 27.4 percent of the reported harvest of 135
deer. This estimate is considerably higher than the 6.0 percent reported
by antlered-only hunter.s in 1974. No estimate of wounding loss was available for either-sex permit holders because the state-wide questionnaire
used for this survey did not request wounding loss information.

Evaluation of the Study 1972-1975
The objective of adjusting the Middle Park deer population to a specified
density on the winter range was prompted by severe winter die-offs during
the 1950's and more recently in 1964-65. Hunting was considered a potential
tool to maintain the Middle Park herd at a density that would minimize large
die-offs. In 1971 a population level of 10,000 deer post-season was established as a herd-size objective. This study, begun in 1972, was aimed at
evaluating whether the population could be held at this level by sporthunting.
Prerequisite to the study was the ability to measure population parameters
of herd size, composition, mortality, and reproduction in order to predict
pre-hunting season population levels (life-table calculations) so that
harvest objectives and permit numbers could be established.
Secondly, a
post-hunting season deer population at or above 10,000 was needed to evaluate
the ability of sport-hunting to adjust the population size. Once the yearly
harvest objective was defined, permits and multiple seasons would be used to
harvest the required number of deer.
Life-tables were calculated each year on a January-December time frame. Each
January a new life table was started, utilizing the most recent census estimate as a base population. Adjusting the life-table each year to the census
lessened the chance of perpetuating incorrect estimates of herd size, reproduction and mortality from year to year.
Population Parameters
Estimating the size of the Middle Park deer population was fundamental to
establishing harvest quotas. A stratified random quadrat census technique
has provided quantifiable estimates of population size within known precision
limits since 1968 (Freddy 1975). The census did however, underestimate the
post-season life-table population estimate in January, 1972 by 1700 deer, in
January, 1975 by 5000 deer and in January, 1976 by 1800 deer while overestimating in January, 1973 by 2400 deer (Table 14).

�Table 13. Harvest and hunter
Colorado, 1972-1975.

participation

and success

during regular

and late deer seasons

in Middle Park,

Year and Season
1972
Reg. 1.1
Total Permits Issued
Antlered-Only
Permits
Either-Sex Permits

4,317
4,317

Total Projected Hunters
Antlered-Only
Permits
Either-Sex Permits

3,336
3,336

-

-

% Success All Permittees
Antlered-Only
Permittees
Either-Sex Permittees
% Success Those Hunting
Antlered-Only Permittees
Either-Sex Permittees
Permits Required-Restricted
Antlered-Only Permits
Either-Sex Permits
Projected Harvest
Antlered-Only Permits
Either-Sex Permits

Late 2:..1 Reg.

-

-

% Hunters Participating
Antlered-Only Permits
Either-Sex Permits

1973

77 .3
77 .3

4,200
2,277
1,923

-

28.9
16.8
41.3

-

-

Yes-Yes
Yes-Yes

356
356

-

985
289 + 53
696 "+ 65

-

~I

Yes-No

-

-

l/Regular season held in October
2/
- Late season held in December.

and November.

l/All

at (P &lt; .05)

confidence

~/Restricted

limits expressed

to hunting

only in Middle

Reg.

2,157

3,543
2,861
682

5,199

7,076
4,991
2,085

-

2,157

81.1
75.6
87.6

-

10.7
10.7

Late

-

23.5
12.7
36.2

-

Reg.

3,406
1,664
1,722 ± 52
1,684 ± 47 1,664

-

1975

Late

-

-

8.2
8.2

1974

-

5,199

± 6811 3,113
2,525 ± 79
± 68
588 + 10
77 .1

-

77 .1

44.7

-

44.7
58.0

58.0

Yes-Yes
965 + 87

-

965 + 87

87.9
88.3
86.2

Late

3,830 ± 81 5,849
4,058 ± 90
3,830 ± 81 1,791 ± 27
73.7

73.7

15.3
12.5
27.0

26.2

-

17.4
14.1
31.3

-

26.2
35.5
35.5

-

Yes-Yes
Yes-Yes

Yes-Yes

541
357 + 79
184 + 14

± 80
1,362 ± 80
1,362

-

82.7
81.3
85.9
12.5
7.5
24.4

-

15.1
9.2
28.4

Yes-No
51
Yes-Yes 884
375 + 60
509 + 18

P:rk.

l/Either-sex
permittees were restricted to one Game Management Unit within
all other years they could hunt in anyone
of four management units within

Middle Park only during 1975, in
Middle Park.

I

N
-....I

~
I

�Table If•. Life equation

for deer 'in Middle Park, Colorado,

January 1?68-January

1976.
Wounding Loss
(20% of

Post-Hunt

POE'
Census Percent
2/
Est. DifferenceExpe c t ed Ij'

Pre-Hunt
Pop.

Harvest
Percent PreHunt. Pop.
No.

43.35

12,807

2,909

23

582

9,316

9,112

+ 2.2

4,550

53.70

13,023

4,503

35

901

7,619

7,206

+ 5.4

6,071

2,636

43.42

8,707

2,436

28

487

5,784

5,730

+ 0.9

8.5

5,241

2,497

47.64

7,738

516

7

103

7,119

5,429

+23.7

50

0.9

5,379

2,678

49.78

8,057

356

4

71

7,630

10,087

-32.2

10,087

647

6.4

9,440

4,691

49.69

14,131

1,980

14

396

11,755

9,670

+17.7

1974

9,670

420

4.3

9,250

4,247

45.91

13 ,497

1,933

14

386

11,178

6,005

+46.3

N

1975

6,005

149

2.5

5,856

2,683

45.82

8,539

928

+37.0

"
'"I

1976

4,677

Year

January
Census
Pop. Est.

Winter Loss
Percent
No. Jan. Est.

Pre-Fawn
Pop.

Fawn ReEroduction
Percent Increase
No. of Spring Pop.

1968

10,640

1,706

16.0

8,934

3,873

1969

9,112

639

7.0

3,473

1970

7,206

1,135

15.8

1971

5,730

489

1972

5,429

1973

}j
Formula used to calculate expected population

11

Ha rves t )

186

indicates

4,677

level was: P2 = PI - ~ + bn - ~, where P2 = expected population level; PI = census populab = net productivity derived from October pre-hunting season doe:fawn ratios; ~ =
n

tion estimate from previous year; M = winter mortality;
w
harvest mortality.
2/
- Percent difference

7,425

the amount the expected population

estimate differs from the quadrat census estimate.

I

�-276Unfortunately, these years of discrepant censuses-life-table estimates
occurred during this study. Because harvest quotas were based on predicted
pre-season population levels calculated from life tables, and census estimates strongly disagreed with post-season life-table estimates, the decision
to adjust the population size was not judicious considering the apparent inability to consistently estimate and predict population size.
Using a paired t-test (P~ .10), the 1972 census estimate was significantly
different than 1973. 1973 was not different than 1974, 1974 differed
significantly from 1975, and 1975 did not differ from 1976 (Table 2). Insufficient snowfall in December 1974 and 1975 could have reduced census
estimates in January 1975 and 1976 (Fig. 2, Table 2). If deer were not
forced onto the winter range (census sample area) by snowfall, then the
primary assumption of the census, that all deer were present on the sampled
area, was not valid.
Measurements of other population parameters, although not subject to
statistical evaluation, did not suggest similar fluctuations in population
size. Estimated pre-natal productivity varied from 156-187 fawns:lOO does
between 1972 and 1975 (Table 5). Post-natal productivity for the 4 years
as measured in October ranged from 87:120 fawns:lOO does while December
ratios varied from 70-88 fawns:lOO does (Tables 3,4).
Also, the relatively
stable percent yearling male, yearling female and total yearling in the
harvest from 1972 through 1975 suggests a yearly recruitment of 30 percent
(Table 15). Winter mortality was less than 7 percent during all 4 years
(Table 6). Precision limits about the estimated harvest were acceptable,
being + 10 percent of the mean for either-sex harvests and + 22 percent for
antler;d-only harvests (Table 13). Unless a large number of persons hunted
in Middle Park without a permit or a large number of deer were harvested
illegally, information suggests no large unknown loss of deer through harvest.
Although these measurements of population characteristics may be in error,
they do not suggest large fluctuations in population size.

Table 15. Percent yearling in the regular season harvest, Middle Park,
Colorado 1972-1975.
Percent Male Yearl.
of Total Male
in Harvest

Percent Female Yearl.
Percent
of Total Female
Yearl. of
Harvest
Total Harvest

Year

Season
Type

1972

Antlered-Only

50.3

1973

Either-Sex

39.7

23.1

32.6

1974

Either-Sex

42.7

36.2

41.2

1975

Either-Sex

42.8

25.9

35.9

50.3

�-277-

)0

,.....
U)

~

0

~
'-'
~

ffi

20

0

1704. inches

~
~
0

~

r:LI
0

~

10

~

z

U)

~

g

2

YEAR

Fig. 2. Total inches of snow received in Decemberat Hot Sulphur Springs
weather station, Middle park, Colorado, 1967 through 1975. Kremmling
weather station measurements substituted in 1974 for missing data.

�-278Harvest Objectives
Fluctuations in deer numbers indicated by the census resulted in attempts
to increase the herd during 1972 and 1975 and reduce the herd to 10,000
in 1973 and 1974 (Table 7). The harvest objective in 1972 was a minimum
harvest to increase the herd to 10,000 post-season. This objective was
accomplished based on the January, 1973 census of 10,087. The 1973 harvest objective was 4000 deer. Only 59.4 percent of the desired harvest
was attained. In 1974 the harvest objective was 3400 deer but only 68.2
percent was harvested. Late-seasons were held in both 1973 and 1974 to help
attain desired harvests. In 1975, 1114 deer or 69.6 percent of the desired
harvest was attained.
Failure to achieve harvest objectives may be traced to: 1) inadequate
numbers of permits issued, 2) inadequate measures of harvest, or 3) inadequate measures of the number of deer available for harvest. Also, at
high hunter densities associated with large numbers of permits issued to
harvest several thousand deer, problems of hunter interactions may alter
the ability of hunters to harvest deer. If the Middle Park deer population
is to be held in-check by sport-hunting, small harvest objectives are needed.
The population should not be allowed to expand more than 2-3000 deer above
the desired post-season level.

Permit Numbers, Distribution, and Restrictions
Persons hunting in Middle Park from 1972-1975 were required to obtain a
Middle Park permit so a list of hunters for Middle Park could be developed
for hunter surveys to improve the precision of harvest estimates (Table 13).
Antlered-only permits were unlimited in number and available from license
agents within Middle Park and at offices of the DOW. Either-sex permits
were limited in number and available by drawing.
The primary problem with antlered-only permits was maintaining control and
records of numbers issued. Permits were lost or issued without record. An
unknown number of permits at-large negated efforts to obtain a list of
hunters for surveys and reduced the accuracy of survey results. This was a
problem primarily during the first 2 years of the study.
Problems with regular-season either-sex permits involved restrictions on
drawings for permits. In 1973, three drawings were held for permits and
only 25 percent of the allotted 2000 permits were issued. Remaining permits
were issued on a first-come basis at DOW offices in Kremmling and Denver.
In 1974, only one drawing was conducted and few permits were issued. Eithersex permits in 1975 were available by drawing in June and then unsolicited
permits were again issued on a first-come basis at the Regional Office, DOW
prior to the season. This latter decision hindered efforts to maintain an
accurate listing of the number of permits issued and to whom. The result
of these decisions was an inconsistent number of either-sex permits issued
even though the number allotted for issue was fairly consistent. The inconsistent either-sex hunting pressure influenced harvest size and composition
and hunter success.
A further problem with regular-season permits involved decisions regarding
whether hunters were restricted to Middle Park. In 1972 and 1975 antleredonly permittees were not restricted to Middle Park while in 1973 and 1974 they

�-279were. During 1973 and 1974 either-sex permittees were restricted to
Middle Park (any of 4 GMU's) while in 1975 they were restricted to one
GMU within Middle Park. These inconsistent decisions (made in contradiction
to Research recommendations) affected hunter distribution, hunting intensity,
harvest size and possibly harvest composition, but the magnitude of this
effect could not be measured.
Late-season permits restricted hunters to Middle Park. Hunters were allowed
to hunt in any of 4 GMU's. In 1974, 54.8 percent of the late-season harvest
was in GMU 18 (Table 16). Subjective evidence indicated much of this harvest
occurred in a restricted locale called "Gunsight Pass". To prevent a massive
shift of hunters and possible over-harvest in a specific area, late-season
permits should restrict hunters to one GMU.

Table 16. Composition of projected late-season either-sex permit deer
harvest, Middle Park, Colorado, 1973-1974, based on returned hunter survey
questionnaires.

1973

18

Management Unit
27
28

37

Totals

Bucks

227

75

66

224

592 ( 61.4%)

Does

124

35

44

123

326

(

33.7%)

B. Fawns

17

5

8

5

35

(

3.6%)

D. Fawns

0

4

0

8

12

(

1.3%)

Totals

368
(38.2%)

119
(12.3%)

118
(12.2%)

360
(37.3%)

1974

18

27

Bucks

357

72

92

149

3

673

(

49.4%)

Does

330

72

65

112

3

582

(

42.7%)

B. Fawns

37

8

15

8

2

70

(

5.1%)

D. Fawns

20

5

2

8

0

35

(

2.6%)

Unknown

2

0

0

0

0

2 (

0.2%)

Totals

Management Unit
28
37

746
157
174
277
(54.8%) (11.5%) (12.8%) (20.3%)

965 (100.0%)
(100%)

Unknown

8
(0.6%)

Totals

1,362 (100.0%)
(100%)

�-280Hunter Success and Participation
Restricting antlered-only permittees to Middle Park appeared to increase
success ratios. Success ratios of antlered-only permittees were 10.7
and 9.2 percent in 1972 and 1975 when permit holders were not restricted
to Middle Park. In 1973 and 1974 when antlered-only hunters were restricted to Middle Park, success ratios were 16.8 and 14.1 percent (Table
13). Restrictions may cause hunters to become more familiar with an area
or to hunt more intensively.
The percentage of antlered-only permittees who hunted in Middle Park was
no greater when permits were restrictive (Table 13). However, during nonrestrictive seasons hunters may have hunted a short time and if immediately
unsuccessful proceeded to another hunting area. Thus, on surveys these
hunters would report hunting in Middle Park but their actual effort in
Middle Park was probably negligible. Percent participation would be inflated and success ratios lowered.
Success of either-sex permittees during regular seasons varied from 41.3
to 28.4 percent from 1973 through 1975 (Table 13). The decreasing success
could indicate a declining availability of deer (Holsworth 1973). Opening
day of 1975 was immediately preceded by 9-12 inches of snow. This should
have aided hunters in finding deer and improved success but also could
have prevented hunters from getting to areas where deer were located.
Percent of either-sex permittees participating during regular seasons
varied from 87.6 to 85.9 percent from 1973 through 1975 (Table 13). This
higher participation rate as compared to antlered-only permittees suggests
that persons who draw for a permit for the regular season are more inclined
to hunt.
Hunter participation was 77.1 and 73.7 percent during late-seasons in 1973
and 1974 (Table 13). Late-season permits were for either-sex and available
only by drawing. Hunter participation would be expected to increase as
compared to the regular season because deer are more vulnerable in December.
However, weather may have decreased hunter participation because late-seasons
were subject to cold temperatures and deep snow.
Success ratios during late-seasons were 58.0 and 35.5 percent in 1973 and
1974 (Table 13). These ratios were higher than the regular season. Success
ratios would be expected to increase during late-seasons when deer are on
winter ranges and more concentrated.

Composition of the Harvest
Composition of the total harvest during the regular seasons appeared related
to the number of either-sex permits issued. In 1973 and 1975 when about 2000
either-sex permits were issued, percent antlerless in the harvest was 43.7
and 45.1 percent. In 1974 when only 682 permits were issued, 23.7 percent
of the total harvest was antlerless (Table 11). Composition of the total
harvest resulting from a combined antlered-only - either-sex season may reflect little about composition of the herd, being more influenced by the
number of either-sex permits issued.

�-281-

Either-sex permittees harvested 63.4 and 75.4 percent antlerless animals
during regular seasons in 1974 and 1975 (Table 9). This suggests that
during regular seasons either-sex hunters are inclined to harvest any deer
regardless of sex. Also, as the number of either-sex permits increased
from 1974 to 1975 the percent antler1ess in the harvest increased (Table
11) .
However, during late-seasons the percent antlerless in the harvest was
only 38.6 and 50.6 percent (Table 16). There is an apparent consensus
among hunters that late-seasons offer the opportunity to harvest "big bucks".
This apparent selectivity towards bucks raises the possibility of altering
sex ratios of deer by repeated antlered-only late seasons.
Composition of deer checked at check stations is often assumed to accurately
reflect the composition of the harvest. Middle Park serves as a hunting
area for the Denver metropolitan region. Both access routes from Middle
Park to Denver merge west of Denver. The Idaho Springs check station located
east of this merger affords a comparison of check station results with returned hunter survey questionnaires.
From 1973-1975 estimates of the composition of the harvest from check station
and returned surveys compared favorably (Tables 11, 12). A chi-square test
was used to test differences between check station and surveys; all deer
classified as unknowns were eliminated from calculations. Expected composition
frequencies were survey values because surveys were a random sample of all
Middle Park hunters. Composition did not differ significantly between check
station and surveys (X2.05,3) from 1973 through 1975. The percentage of
bucks sampled at check station as compared to the survey, was consistently
lower. People living within Middle Park, and not passing through check
station, may primarily harvest bucks which would inflate the survey buck percentages.
There were no significant differences in the sex and age composition of the
harvest samples from check station and the survey in 1973 and 1975 between
GMU's (X2.05,3) but there was a significant difference in 1974 (X2.05,3).
The problem in 1974 pertained to GMU 28 which may have been hunted primarily
by local persons who did not check through check station.

Political Influence and Maintaining Control of the Study
The primary deterrent to conducting this type of study is the inevitable
conflicts which arise from the need for research personnel to establish
hunting season regulations which is normally a management function. Hunting
seasons also are profoundly influenced by political considerations.
Political
decisions can influence the research process such that there is no longer
control necessary to achieve consistency in permit numbers, permit restrictions,
and harvest objectives. Secondly, research places itself in a delicate
position by establishing numerical objectives based on numerical input. If
measurements fail to agree with predictions then a credibility gap develops.
Few persons outside the research establishment are cognizant of the difficulty
of measuring highly variable biological parameters, and the discrepancies
reflect negatively on research in general rather than on temporary design
inadequacies of a particular research project. The ideal situation for an
experimental harvest study would exist when research had complete control

�-282over an area containing a discrete deer herd. Then rigid controls could
be imposed or experimental design so factors such as hunter numbers,
hunter distribution, hunter effort, season length and timing, and bag
limits could be varied as the design required. Such a situation is not
readily available in Colorado at this time.

LITERATURE CITED
Freddy, D. J. 1975. Middle Park Deer Study - experimental harvest
regulations.
Colo. Div. Wildl., Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rep., July, Part 2, p. 209-240.
Gill, R. B. 1969a. Middle Park Deer Study - population density and
structure.
Colo. Game, Fish and Parks, Game Res. Sec., Fed. Aid Proj.
W-38-R.
Game Res. Rep., July, Part 1, p. 105-122.
1969b. Middle Park Deer Study - productivity and mortality.
Colo.
Game, Fish and Parks, Game Res. Sec., Fed. Aid Proj. W-38-R. Game Res.
Rep., July, Part 1, p. 123-140.
1971. Middle Park Deer Study - population density and structure.
Colo. Game, Fish and Parks, Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rep., July, Part 2, p. 170-188.
Holsworth, W. N. 1973. Hunting efficiency and white-tailed
J. Wildl. Manage. 37(3) :336-342.

deer density.

Kormondy, E. J. 1969. Concepts of ecology.
Cliffs, N.J. 209 p.

Prentice-Hall

Inc., Englewood

Odum, E. P.
546 p.

W. B. Saunders Co., Philadelphia.

1959.

Fundamentals of ecology.

Roper, L. A. 1973. Middle Park Deer Study - experimental harvest regulations. Colo. Div. Wildl., Game Res. Sec., Fed. Aid Proj. W-38-R.
Game Res. Rep., July, Part 2, p. 147-162.
1974. Middle Park Deer Study - experimental harvest regulations.
Colo. Div. Wildl., Game Res. Sec., Fed. Aid Proj. W-38-R. Game Res.
Rep., July, Part 2, p. 197-210.

Prepared by

C21v..e-/ ~d,J,

/

Q

Dave Freddy
Asst. Wildlife Researcner

�-283July, 1976
JOB rI~OGRESS REPORT

State of ___

COLORAD_O

Project No.

W-38-R-31

Deer-Elk

Work Plan No.

14

Job No.

Job Title

Middle Park Cooperative

Period Covered:
Personnel:

_
Investigations

9
---------------------

Deer Study - Deer Habitat Evaluation

April 1, 1975 through March 31, 1976

L. H. Carpenter, D. L. Baker, M. A. Kautz, D. J. Freddy, D. W.
Reichert, W. L. Regelin, P. F. Gilbert, and R. B. Gill.

ABSTRACT
---Work during the 1975-76 segment centered on estimating energy expenditure
and forage intake of free-ranging mule deer during the winter period.
Four
deer were observed in a 4 ha pasture from an observation tower for fifteen
24-hour periods during January and February.
Deer were active 42.7 'and bedded
57.3 percent of total time during this period.
Deer were more active during day
than night. Diets, bite sizes, and number of bites per second of grazing were
estimated by accompanying the deer as they grazed. Estimated daily dry matter
intake for these four deer averaged 787 g. The four deer lost an average of
12.2 percent of initial body weight during 47 days they were at pasture.
Nutritional quality of ingested forage was evaluated by an in vivo digestion
trial. Summaries of horizontal and vertical components of deer-mQvements,
and laboratory analyses of diets of deer in the in vivo digestion trial' are
not yet completed, consequently no comparison of-energy intake to energy
expenditure of the deer is made in this report.

��-285-

MIDDLE PARK COOPERATIVE
DEER STUDY - DEER HABITAT EVALUATION
L. H. Carpenter

P. S. OBJECTIVE
Develop procedures for quantifying the capacity of Middle Park winter ranges
to support deer.

SEGMENT OBJECTIVES
1.

Develop and test a method for estimating energy expenditures of freeranging mule deer.

2.

Estimate forage intake and energy expenditures of deer during "critical"
winter months (January-March).

3.

Estimate nutritional quality of ingested forage in terms of in vivo digestible and metabolizable energy.

METHODS AND MATERIALS
Methods and materials employed in this study have been previously described
(Gill and Baker 1974; Carpenter and Baker 1975). Certain changes in methodology
have occurred; these are described below.

Method for Estimating Energy
Expenditures of Free-Ranging Mule Deer
This portion of the study is being conducted by M. A. Kautz, Colorado State
University.
Detailed methodology is in her study plan (Appendix A).

Estimate of Forage Intake and Energy Expenditure
A thesis entitled "Energy Requirements of Mule Deer Fawns in Winter" has been
prepared by Dan L. Baker. This thesis covers certain objectives of the study
and is presented in Appendix B.
Four mule deer, two, 18 months and two, 30 months old, were used during the
1975-76 segment. The deer were placed in a 2.4 ha conditioning pasture on
January 1, 1976, and left until January 7 when they were moved to a 4 ha
energy budget pasture and remained there until February 15. Deer received no
artificial supplement during this period except for the first four days when
each deer received 0.45 kg concentrate daily.
Two types of measurements were made during the study and these were alternated
from week to week. One measurement estimated forage intake. Deer were
accompanied by two observers as they grazed and bite sizes, total number of

�-286-

bites eaten per second, and total number of bites of each plant species
were recorded. The other measurement recorded observations from an
observation tower of deer activity and travel over 24-hour time periods.

Forage Intake
Estimates of forage intake were .made during the week of January 19-22
following procedures previously outlined (Carpenter and Baker 1975). Fourteen separate trials averaging 63 minutes each were conducted during the
4-day period. Size (weight) of an average bite was determined by two
observers each "plucking" what they considered to be an average bite grazed
by deer. Each observer plucked 20 "bites" of various plant species consumed
by each deer and placed them in a paper bag. Fifty of these 20 ''bite''
samples were collected by each observer.

Nutritional Quality of Ingested Forage
To determine nutritional quality of forage ingested by deer in the pasture
a unique in vivo digestion trial was conducted. At the conclusion of the
studies in the energy budget pasture on February 15, two deer (numbers 9 and
18) were maintained on native forage in the conditioning pasture with minimal
daily supplements of alfalfa hay and concentrate. At 6 P.M. February 27,
each deer was placed into a digestion cage (Carpenter and Baker 1975) for two
days of adjustment. The deer remained in the cages until 8 A.M. March 1
without any feed. Snow was the only source of free-water and was provided
ad libitum throughout the trial. Beginning at 8 A.M. March 1, and continuing
daily for 5 days, the two deer were taken out of the cages during daylight
hours and placed in the energy budget pasture to graze. The deer were returned to the cages for about 2 hours (11 A.M. to 1 P.M.) each day to permit
summarization of A.M. trial data.
During trials each deer was accompanied by an observer and information on
number of bites of each species and plant parts eaten was recorded. Each
time a deer defecated or urinated the total excretions were collected with
special long-handled containers. A third observer was available to quickly
label and place the excreted materials in separate containers. At night deer
were returned to the digestion cages and received no supplemental food. Feces
and urine were collected from the cages daily at 8 A.M. At the conclusion
of the grazing trials at 5 P.M. on March 5, the deer were returned to the
cages and remained until 8 A.M. March 7, when they were removed. The deer
received no feed during these two days and urine and feces were collected both
days.
Daily intake was estimated by multiplying total number of bites for each deer
times the average bite size. Daily fecal and urinary output was determined
from total collections while deer were at pasture and from the cages for the
7-day period March 1 to March 7. Aliquots of feces and urine were saved for
determinations of moisture, gross energy and protein as described by Baker
(1976). A diet for each deer for the total grazing period was determined,
and the top 10 species were identified and ranked. These 10 species made up
94 percent of the diet for deer number 9, and 97 percent of the diet for deer
number 18. Each of the 10 species was then collected in amounts proportional
to its occurrence in the diet. Plant parts similar to those grazed by deer

�-287were collected. Collected samples were taken to the laboratory, ground in
a Wiley mill and precise percentages of each species (by weight) composited
to simulate a diet. Analyses of gross energy, protein, and in vitro digestibility will be made on these diets to compare with in vivo determinations.

Energy Expenditure
Observations of 24-hour activity patterns of deer in the pasture were made
during 3 five-day periods. The dates for these periods were January 11 to
16, January 25 to 30, and February 8 to 13, 1976.

RESULTS AND DISCUSSION

Method for Estimating Energy
Expenditures of Free-Ranging Mule Deer
This portion of the study is in planning stages (Appendix A).
be presented next segment.

Results will

Estimate of Forage Intake and Energy Expenditure

Forage Intake
A total of 878 minutes (14.6 hours) was spent tallying bites of forage ingested by deer during 14 grazing trials. During this period 4,396 bites were
recorded (including bites of snow). These bites were consumed during 12,179
seconds of deer grazing activity. This resulted in a value of 0.36 bites!
second or 1 bite each 2.8 seconds. This is similar to the 0.35 value determined last segment (Carpenter and Baker 1975). The combined diet of the four
deer for the period January 19-22 is presented in Table 1. The total number
of bites differs from the 4,396 previously given as a result of deletion of
bites of snow, hair and feces. Muttongrass (Poa fendleriana) was the major
plant species in ~he diet. This contrasted with 1975 when vetch (Astragalus
convallarius) was the major food item, making up over 50 percent of the diet
of the deer in the pasture (Carpenter and Baker 1975). In 1976 vetch comprised
only 9.8 percent of the diet. The reasons for this difference are not known.
It is suspected that differences in snow depths (less snow in 1976) was a
major factor in this discrepancy. Big sagebrush (Artemisia tridentata) made
up a similar portion of the diet each year.
Bite sizes taken by the deer averaged 0.148 g (se + 0.004) wet weight and
0.109 g (se
0.004) dry weight. These bite sizes-are 33.1 and 17.4 percent
larger respectively than those determined last segment (Carpenter and Baker
1975). Average moisture content of the 100 samples was 26.2 percent, considerably greater than the 9.2 percent moisture content of bite samples in
1975. This moisture difference may account for the difference in bite sizes
between the two years.

±

�-288-

Table 1. Diet of four mule deer in a 4 ha energy budget pasture for period
January 19-23, 1976.
Plant Species

Total Bites

% Total Bites

Muttongrass (Poa fendleriana)

1,167

28.7

Rabbitbrush (Chrysothamnus vaseyi)

711

17.5

Big sagebrush (Artemisia tridentata tridentata)

653

16.0

Vetch (Astragalus convallarius)

400

9.8

Phlox (Phlox bryoides)

286

7.0

Pine needlegrass (Stipa pinetorum)

236

5.8

Serviceberry (Amelanchier alnifolia)

181

4.4

Mat penstemon (Penstemon caespitosus)

155

3.8

Buckwheat (Eriogonum umbellatum)

76

1.9

Snowberry (Symphoricarpos oreophilus)

51

1.2

Bluebunch wheatgrass (Agropyron spicatum)

41

1.0

Penstemon (Penstemon cyath6ph6:tus)

23

0.6

Western wheatgrass (Agropyron smithii)

19

0.5

Fringed sage (Artemisia frigida)

15

0.4

Needle-and-thread grass (Stipa comata)

10

0.2

Blue grama grass (Bouteloua gracilis)

9

0.2

Parry's rabbitbrush (Chrysothamnus parry i)

7

0.2

Bottlebrush squirreltail grass (Sitanion hystrix)

7

0.2

Ryegrass (Elymus spp.)

6

0.2
0.2

Unknown grasses

5

0.1

Bluebell (Mertensia la~ceolata)

3

0.1

Lichens (species unknown)

2

T

4,069

100.0

Totals

�-289Tower observations indicated deer spent 23.2 percent (se ± 2.14) of their
time grazing. On a 24-hour basis this would total 20,045 seconds grazing.
This total multiplied by the 0.36 bite/second value results in an estimated
total of 7,216 bites ingested by deer each day. Multiplying this total by
the average bite sizes of 0.148 g (wet weight) and 0.109 g (dry weight),
daily forage intake was estimated to be 1,068 and 787 g respectively.
On
a dry-weight basis this compared closely to the 765 g value determined
last segment (Carpenter and Baker 1975).
All deer lost weight during the 47-day period they were in the pastures
(Table 2). The average weight loss was 12.2 percent of initial body weight.
This represents a weight loss of 169 g/day. At this rate of weight loss
and at an average initial weight of 65.6 kg, the deer would experience an
average body loss of 25 percent at the end of 97 days. Weight loss greater
than 25 percent has been shown to cause mortality in mule deer fawns (Baker
1976).

Table 2. Weights (kg) of four mule deer at pasture during period December
30, 1975 to February 15, 1976.

9

Deer Number
18
20

39

Average

30

64.1

70.0

72.7

55.5

65.6

January 11

62.3

60.0

73.6

54.1

62.5

January 16

60.9

64.5

71.4

53.2

62.5

January 25

56.8

60.0

69.5

51.4

59.4

January 30

57.3

60.9

69.5

51.4

59.8

February 9

55.9

60.0

65.9

51.4

58.3

February 15

54.1

60.5

65.9

50.0

57.6

Total Loss

10.0

9.5

6.8

5.5

8.0

Percent Loss

15.6

13.6

9.4

9.9

12.2

Grams Per Day Loss

213

202

145

117

169

Dates

December

Nutritional

Quality of Ingested Forage

A total of 1277 minutes (21.3 hours) and 1322 minutes (22.0 hours) were spent
during the 5-day trial tallying bite counts of forage for deer numbers 9 and
18 respectively.
A total of 14,867 bites were recorded for deer number 9 while

�-2909,638 were recorded for deer number 18. Total bites taken daily by each
deer is presented in Table 3. Diet for the two deer during this period
was similar with respect to the top 10 plant species but varied considerably
in ranking of these 10 species (Table 4). The largest difference between
diets of the two deer was in amount of big sagebrush. Big sagebrush made
up only 5.1 percent of the diet of deer number 9 but comprised 30.6 percent of the diet of deer number 18 (Table 4).

Table 3. Number of bites taken daily by deer numbers 9 and 18 for 5-day
in vivo digestion trial conducted in a 4 ha energy budget pasture March 1-5,
1976.
Deer Number
Date

9

18

March 1

2,331

1,467

March 2

2,187

854

March 3

3,747

2,662

March 4

2,252

1,381

March 5

4,350

3,274

Totals

14,867

9,638

Laboratory work has not been completed, therefore no results are available
on digestibility, gross energy, or protein content of the diets. Results of
this work will be presented next segment and will be used to determine energy
intake for deer in the pasture.

Energy Expenditure
Fifteen 24-hour days were spent in the observation tower during 3 separate
weeks of observations.
Using a sampling strategy of four 5 minute observation periods each hour (Carpenter and Baker 1975), 7200 minutes (120 hours)
of observations were made.
Summaries of 5 minute samples for all trials indicated deer spent 42.7 percent
(se + 3.6) of their time grazing, standing, or moving, and 57.3 percent (se +
3.6)-of their time bedded. A summary of total activity observations for theentire 24-hour period (no sampling involved) indicated that deer were active
43.0 percent (se ± 1.1) and bedded 57.0 percent (se ± 1.1). In contrast to
last segment where the sampling strategy overestimated active periods, sampling
estimates this segment were very close to the actual activities.
It would
appear that the longer sampling periods employed this segment obviated the
discrepancies previously noted.

�Table 4. Diets of two mule deer in energy budget pasture during in vivo digestion trial March 1 to
March 5, 1976.

Food Item

Total Bites
Deer No. 9
Deer No. 18

Muttongrass (Poa fend1eriana)
Phlox (Phlox b!Yoides)
Vetch (Astragalus conva11arius)
B1uebunch wheatgrass (Agropyron spicatum)
Serviceberry (Ame1anchier a1nifo1ia)
Bott1ebrush squirre1tai1 grass (Sitanion hystrix)
Big sagebrush (Artemisia tridentata tridentata)
Rabbitbrush (Chrysothamnus vaseyi)
Snowberry (Symphoricarpos oreophi1us)
Buckwheat (Eriogonum umbe11atum)
Blue grama grass (Boute1oua gracilis)
Fringed sage (Artemisia frigida)
Currant (Ribes cereum)
June grass (Koe1eria cristata)
Unknown grasses
Unknown forbs
Penstemon (Penstemon cyathophorus)
Pusseytoes (Antennaria parvif1ora)
Pine need1egrass (Stipa pinetorum)
Lichen (species unknown)
Dry bark
Need1e-and-thread grass (Stipa comata)
Mat penstemon (Penstemon caespitosus)
Green rabbitbrush (Chrysothamnus viscidiflorus)
Indian rice grass (Oryzopsis hymenoides)
Astragalus (Astragalus shortianus)
Bluebell (Mertensia 1anceo1ata)
Alfalfa (Medicago sativa)
Western wheatgrass (Agropyron smithii)
Moss (species unknown)
Parry's rabbitbrush (Chrysothamnus parryi)

5,643
1,984
1,649
1,342
930
810
765
358
321
166
140
137
131
120
90
44
43
39
36
25
22
16
13
12
10
7
6
4
3

Totals

14,867

1

o

2,130
3,006
224
117
476
41
2,944
231
22
87
3

63
3
15
64
17
29

o
10
1

47

o
67
6

o
o
12
5
15

Percent Total Bites
Deer No.9
Deer No. 18
38.0
13.3
11.1
9.0
6.3
5.4
5.1
2.4
2.2
1.1
0.9
0.9
0.9
0.8
0.6
0.3
0.3
0.3
0.2
0.2
0.1
0.1
0.1
0.1
0.1
0.1
0.1

22.1
31.2
2.3
1.2
5.0
0.5
30.6
2.4
0.2
0.9
T

0.7
T

0.1
0.7
0.2
0.3
0.0
0.1
T

0.5
0.0
0.7
T

0.0
0.0
0.1

T

T

T

o

T

0.2
0.0

3

0.0

T

9,638

100.0

100.0

I
N
\D

I-'
I

�-292-

Deer were less active at night than during day. A summary of observations
for shifts 1 and 4 (night) indicated that deer were bedded 65.3 percent
(se + 3.73) of the time, while during shifts 2 and 3 (day) deer were bedded
on1y-49.5 percent (se ± 5.21) of the time.
Activity was consistent between trials and between individual deer (Table
5). These values were taken from total activity observations and represent
actual activity of each deer. A malfunction in the collar of deer number
20 in trial 1 required that certain observations be made on deer number 4
for only a portion of thjs trial. This combination may explain the larger
difference between trials for deer number 20 than for the other three deer.

Table 5. Comparison of percent total time up for four mule deer during 3
separate periods during January and February, 1976.
Deer Number
Trial Number

9

18

20

1

44.9

44.3

47.8 !/

42.2

2

44.1

42.1

37.2

46.3

3

44.6

41.6

33.6

47.0

44.5

42.7

39.5

45.2

Average

39

1/
Portion of observations
as deer number 20.

in this trial were made on deer number 4 as well

Twenty-four hour activity patterns of deer are graphed in Figure 1. Maximum activity (not bedded) occurred during hour 18 (6:00 P.M. MST). Maximum
bedding activity occurred during hour 15 (3:00 P.M.). A summary of continuous
activity records involving 523 individual active periods for all deer and all
trials indicated the average activity period of deer was 71 minutes (se ± 2.67)
[range 1-273] and for 519 bed periods the average was 95 minutes (se ± 2.92)
[range 1-350]. Average number of bedding periods per day was 8.7 (se ± 0.48).
A frequency distribution of 519 bed periods showed that the greatest number
were from 1-20 minutes long (Fig. 2).
Total number, average length, and number per day of both active and bedding
periods reveal certain differences between deer and between trials (Tables 6
and 7). Average time per active period for deer numbers 18, 20, and 39 were
similar. Deer number 9 consistently was active for shorter periods of time
(Table 6). Deer were not as consistent in lengths of bedding periods as with
active periods, but the same pattern was apparent. Deer number 9 consistently
bedded for shorter periods than other deer (Table 5).

�-293-

MIDNIGHT

24
80%

""

,.

"
p----,O

12

•

•

• PERCENT

ACTIVE

NOON
0- - 0--0

PERCENT

GRAZING

Fig. 1. Hourly (Mountain Standard Time) activity and grazing patterns of four
mule deer in 4 ha energy budget pasture during January and February 1976.

�-294-

1

N :519

20

60

100

. MINUTES

140

180

220

260

300+

BEDDED

Fig. 2. Frequency distribution of lengths of 519 bedding periods for 4 mule
deer at pasture during January and February 1976, expressed in 20-minute units.

�Table 6. Number of active periods, average length in minutes of each, and average number per day for each
deer, for 3 trials during January and February, 1976.

9

Trial 1
Deer Number
18
20 1/ 39

9

Trial 2
Deer Number
18
20

39

9

Trial 3
Deer Number
18
20

39

Total All Trials
Deer Number
18
20
9

39

Number

53.0

3~.0

34.0

50.0

46.0

32.0

39.0

35.0

57.0

52.0

42.0

44.0

156.0

123.0

115.0

129.0

Average
length

60.9

81. 7

101. 3

60.8

69.0

94.8

68.7

95.2

56.3

57.6

57.6

77 .0

61.6

74.9

74.3

75.6

Standard
error
(length)

7.9

10.2

12.5

8.9

8.4

12.8

8.3

12.4

6.5

7.3

6.3

9.3

4.4

5.7

5.5

5.9

Number
per day

I

10.6

1/Observations

7.8

6.8

10.0

9.2

6.4

include both deer numbers 4 and 20.

8.0

7.0

11.4

10.4

8.4

8.8

10.4

8.2

7.7

8.6

N
\0
V'I

I

�Table 7. Number of bedding periods, average length in minutes of each, and average number per day for each
deer, for 3 trials during January and February, 1976.
Total All Trials

Trial 3

Trial 2

Trial 1
9

Deer Number
18
20 1/

39

9

Deer Number
18
20

39

9

Deer Number
20
18

39

9

Deer Number
20
18

Number

52.0

38.0

33.0

49.0

45.0

31.0

34.0

58.0

53.0

43.0

44.0

155.0

122.0

115.0

127.0

Average
length

76.3 105.6 113.8

84.9

89.4 134.4

115.9 113.7

68.8

79.3

111.1

86.7

77 .3

101.5

113.5

93.2

Standard
error
(length)

7.9

12.5

8.6

10.6

11. 7

6.6

6.8

9.4

8.9

4.8

6.2

6.4

5.7

Number
per day

1/

12.5

39.0

13.1

12.4

39

I
N

10.4

Observations

7.6

6.6

9.8

9.0

6.2

include both deer numbers 4 and 20.

7.8

6.8

11.6

10.6

8.6

8.8

10.3

8.1

7.7

8.5

1.0
0\

I

�-297-

These activity values differ somewhat from those reported for mule deer
and white-tailed deer. Linsdale and Tomich (1953:355) reported that the
average length for 300 completed rest periods for mule deer was 47 minutes.
Moulton (1967), working with three radio-tagged white-tailed deer during
winter, found the average duration of 348 bedding periods to be 110
minutes and average duration of 346 active periods to be 124 minutes.
Montgomery (1963) reported a mean bedding period throughout the year of
106 minutes for white-tailed deer in Pennsylvania.
Twenty-four hour grazing pattern of the deer is also graphed in Figure 1.
Grazing activity parallels general activity.
Grazing activity was maximum
during hour 18 (6:00 P.M.) with smaller peaks at hours 8 (8:00 A.M.), hour
17 (5:00 P.M.), and 24 (12:00 P.M.). Percentage of time spent each hour in
various activities is summarized in Table 8.

Table 8. Percentage of time spent each hour in various activities by four
mule deer in 4 ha energy budget pasture during January and February 11976.

Hour

Percent Total
Time Up

Percent Total
Time Bedded

Percent Total
Time Grazing

Percent Active
Time Grazing

1 (1 AM)
2
3

39.9

60.1

22.1

55.4

37.6
18.4

62.4
81.6

18.8
8.4

50.1
45.5

4

36.5

63.5

29.3

80.4

5

30.4

69.6

13.1

43.2

6

30.2

69.8

8.8

29.2

7

55.3

44.7

16.1

29.1

8

60.2

39.8

32.7

54.3

9

56.7

43.3

21.6

38.1

10

43.3
38.3

56.7
61.7

24.4

11

23.5

56.3
61.4

12

59.5

40.5

30.2

50.7

13

49.3

50.7

19.1

38.7

14

34.4

65.6

17.0

49.3

15

13.1

86.9

9.5

72.5

16

38.9

61.1

22.0

56.7

17

74.0

26.0

38.8

52.4

18

16.0

51.1

60.8

19

84.0
61. 7

38.3

33.2

53.9

20

16.3

83.7

13.0

79.6

21

80.7

14.9

77.1

22

19.3
31.3

68.7

19.5

62.1

23

46.0

54.0

30.2

65.6

24

49.3

50.7

39.6

80.3

�-298-

Data on horizontal and vertical components of travel during each observation have not been sunnnarized. These values will be incorporated with
estimates of energy expenditure by activity to provide estimates of total
energy expenditure of deer in the pasture. Energy expenditure estimates
will then be compared to estimates of energy intake to develop energy
budgets for deer in the pasture. Energy budget comparisons will be reported next segment.

LITERATURE CITED
Baker, D. L. 1976.
M.S. Thesis.

Energy requirements of mule deer fawns in winter.
Colo. State Univ., Fort Collins. 76 p.

Carpenter, L. H., and D. L. Baker. 1975. Middle Park deer study - deer
habitat evaluation.
Colo. Div. Wildl., Game Res. Div., Fed. Aid
Proj. W-38-R-29.
Game Res. Rep., July, Part 2. p. 241-263.
Gill, R. B., and D. L. Baker. 1974: Middle Park deer study - deer habitat
evaluation.
Colo. Div. Wildl., Game Res. Div., Fed. Aid Proj.
W-38-R-28.
Game Res. Rep ,, July, Part 2. p. 211-225.
Linsdale, J. M., and P. Q. Tomich. 1953. A herd of mule deer.
California Press, Berkeley and Los Angeles. 567 p.

Univ.

Montgomery, G. G. 1963. Nocturnal movements and activity rhythms of whitetailed deer. J. Wildl. Manage. 27(3) :422-427.
Moulton, J. C. 1967. Movement and activity of three white-tailed deer
during the winter of 1964-65 in East-Central Minnesota determined
by telemetry. M.S. Thesis, Univ. of Minnesota. 82 p.

Prepared by --:.c-&lt;",-/~~,,---,,~!I.!....-'..o.U"'-'~-"-'#I-jJ;_"'t._-t;
Len H. Carpenter
Asst. Wildlife Researcher

_

�-299-

A P PEN

D I X

A

�-300-

STUDY PLAN
STUDY PLAN NO.

THE USE

3

OF HEART RATE TO ESTll1ATE

THE ENERGY EXPENDITURE

DECEMBER,

Marie

OF UNRESTRAINED

1975

A. Kautz

DEER

�-301-

Study Plan
The Use of Heart Rate to Estimate the Energy Expenditure
of Unrestrained Deer
A.

Need
To estimate the number of deer a range can support, it is necessary
to evaluate the capacity of the range to meet the requirements of individual
animals, one of wh i.chis energy (noon 1973).

An inadequate supply of

energy on deer wi.nt er ranges may result in Hinter mortal ity and limit deer
populations.

A certain amount of a deer's energy requirement must be

obtained from nutrients supplied by the range.
The energy expenditure of an animal can be determined by either direct
or indirect calorimetry (Brody 19:15). Direct calorime t ry i nvo lve s measur-ev

ment of the actual heat :~pr6dnc~d ," by the subject.

The direct measur-ement

of heat production requires elaborate equipment and is not a technique presently v i ewed as feasible wi th unrestrained animals.

Indirect calorimetry

involves the estimation of heat production (energy expenditure) through
the mcasur emen t of rates of related body functions.
Traditionally oxygen consumption and carbon dioxide production have
been measured and related to an equivalent heat production in order to calculate energy expenditure (Brody 1945).

Such experiments are often con ...

ducted in respiration chamber s . While such chambers are Lmpor t ant in the
study of haseline conditions, they do not permit consideration of the
physiological and behavioral responses evoked by a normal range of env i ron-..
mental conditions.

The energy expenditure of animals in the field might

be estimated by measurement of gas exchange, by relation of energy cxpenditurc to heart rate, respiration ~ate) or pulmonary ventilation rate,
by the use of radioisotope techniClues,or by the use of energy balance studies.

�-302-

!'I(!asurin~
oxygen antIcarhon dioxicl~~xc]:angc in the field
Gas collection from animals in the field is a prohlem.

Matt£eld

(1974) trained white-tailed deer to the use of face masks and restraint
by leashes from a very young age.

Of 13 fawns, two Here suitable for res-

piratory collections, and two others could be used with tranquilization.
Other workers have used face masks with reindeer (White and Yousef 1974),
burros (Yousef and Dill 1969 ~,~), and gazelles (Dan l3aharov,pel's. corom.),
anc.lnone reportec.lsuch serious problems.
Expired air can be ciollected without ~le use of face masks from animals
wh i ch have been prepared w i th tracheotomies.
COl'PI.)

Fair and Naut z (Mautz, pel'S.

used this method on wh i.t
e=tu i led deer and reported no problems related

to the surgical procedures.

However tracheotomy may disrupt heat exchange

in the nasopharynx and can affect respiration and met aboli sm (!31axterand
.Ioyce 1963).

Other components of the system for measuring gas exchange in the field
include either 1) an apparatus for st ori.ngexp i red air for later volume determi.nat i.onand gas :',alys i s , or 2) a portable f lovneter for measuring volume
of the expired air in the field/\'litha device capable of taking a sample to be
stored for later analysis.

In either case the apparatus is too cumbersome

to be car r i ed by a deer;

One small portable f lowme t er, the I'lrightrespiro-

meter, could be carried by a deer, but it is not rugged enough to meet specifications (~Iautz,pel's. comm.).

A small amount of expired air could be

collected in bags carried by the animal, but there are obvious limitations
to this method.
Thus measuring the gas exchange of deer in the .f i.eld wou'ld necessitate
control of activity by a leash or similar means, and the presence of a person to carry apparatus.

The technique would require \'lell-trainedanimals,

as nervousness could result in spurious estimates of energy expenditure.

�-303-

Heart rate
Oxygen consumption by the tissues is a function of three parameters:
heart rate, stroke volume, and the difference in the oxygen saturation of
arterial and venous blood (lvyndham,et ale 1959).

While stroke volume and

oxygen saturation cannot by determined readily in the field (Mike Wells,
pel's. comm.), heart rate can he monitored by currently available telemetry
systems.

The relation between heart rate and energy expenditure requires

careful determination before this parameter can be used as a predictive tool.
A nwnber of studies have investigated the heart rates of humans engaged in different activities, and some have related heart rate to energy
expenditure.

Ma Iho t ra et al. (1963) found linear correlations between

heart rates and energy expenditures of humans exercising ~m bicycle ergometers ,d th pulse rates ranging from 95 to 150 beats per minute.

The

relation could be used to predict the energy cos t s of other activities
performed by the same individual with less than 10 percent error.
Webster (1967) found a close relationship between heart rate and
energy expenditure in three of four sheep tested under cold stress ancI at
ditferent levels of feeding in thermoneutral conditions, standing and

IJ ing, and w i t h heart rates up to 90 beats per minute.

lieconcluded that

two of these sheep ,::ere:
sui t able for prediction of energy exnend i ture from
heart rate with errors of less than 10 percent.

Brockway and McEwan (1969)

found no relation between heart rate and energy expenditure in four of
five sheep tested.

They attributed this to a poor relationship between

heart rate and stroke volume, possibly clue to i.nadequate training and nervousness of the animals.
Silver (1973) found a linear relation with a correlation coefficient
of .805 be tween the heart rate and energy expenditure of fasted and fed
wh i t.e-ct a i Led deer in respiration chambers.

However , t he effects of different

activities and psychological conJitions on the relation have not heen demon-

�-304-

strated.

Boforo heart rate can he used to predict tIle energy expenditure

of unrestrained animals it is necessary to test the correlation over a range
of conditions more closely approximating those encountered by free roaming
deer.

BOiVcver such tests are not easily made.
Fair and t·lautz(Maut z, per s . comm.) investigated this relation recently

using a deer which had been tracheotomized and carried a small beg permitting
ga~ collections for short periods \~lile the animal was unrestrained.

They

obtained ruloverall correlation coefficient of about .5 for a wide range of
activities and stimuli.

This low observed corrnlation coefficient could be

due to one or more different factors, including nervousness due to the
experimental apparatus.

Indeed any apparatus currently available for

cstimat i ng energy expenditure of unrestrained animals by gas collection
(for use as a standard) would he likely to produce nervousness in deer, with
possible effects on the relationship being investigated.
To summar i ze , in humans there appears to be a strong relation between
heart rate and energy expend i ture wh i ch ho Ids for individuals over different
types of activit ..!s. Studies with sheep have shown a similar relation for
some subjects but md i.cs.t
e that other individuals are unsuitable for use as
experimental subjects, possibly due to nervousness associatcd with the experimental conditions.

Pooled data from white-tailed deer in respiration cham-

bers show a high correlation between the two variables.

A similar relation

has not yet been demonstrated for unrestrained decr and efforts to do so may
be complicated by the bul ky apparatus available for determination of energy
expenditure.

Denons t rati on of a high correlation between the heart rate

and energy expenditure of unrestrained deer \vould provide a strong empirical
basis for prcMcting

energy expenditure from heart rate.

�-305-

Other

techniques
Other work er s have suggested

from respiration

rate

can be estimated

(Ramanathan 1964) or pulmonary ventilation

(Datta

and

1974).

In addition,

techniques,

including

to

entry

(Yo,ung et al.

radioisotope
carbon

dioxide

have been suggested.
fo r study

rate

the use of carbon-14

1969, White and Long

et a l . 1971) and the use of doub l y-Tabe l ed water

carborid i ox i de production

(Lifson

et al.

Each of these

than the relation

mined by gas collection

1955, Lifson

alternatives

be tween heart

through

and }IcClintock

was judged

rate

to estimate

less

1966)

appropriate

and energy expenditure

deter-

a face mask.

Objectives
The objectives
face mask apparatus
investigate

of this

the relation

Expected

1) evaluate

collections

activities,

and,

associated

and time factors,

energ), expenditure

are to:

to heart

if

the usefulness

f'rom mule deer

between the hcar t rate

3) examine the variability
conditions

study

for respirator),

fmYns engaged in several

and energy

a relation

is

of a

fawns , 2)

expend i t.ur e of
demonstrated,

to

w i.th a wi der range of environmental

and 4) provide

predictive

equations

relating

rate.

Results:

Completion

of the proposed

study

wi l l provide

information

use of a face mask apparatus

and heart

rate

ture

equations

for predicting

of deer,

from heart
D.

expenditure

et a l , 1966, t-ial}lOtra et a l , 1962, }.lattfeld

1968, Corbett

C.

energy

Ramanathan 1969, Sharkey

determine

B.

that

and may provide

on the

to measure the energy
energy

expendi-

expenditure

rate.

Approach

for

Animals:

Two mule deer

Fawns and one whi.t c=t a i Lcd deer

earl)'

of equipment

and procedures.

tests

fawn w i l l be us cd

These fm~lS Nere bottle-fed

�-306-

and accus t.oned
experinental
h[:.rness,

to the experimenter

procedures.

leash,

from an ear l y age, hut Here not trained

In 1976, fawns will

and face mask and to enter

with the goal of having

four well-trained

be trained

to

to the use of a

crates

or a vehicle

for transport,

animals

accustomed to experimental

personnel.
!tespirationc:::pyaratus:
equipped

w i t h valves

Gas collection

for

inspiration

or a qu i ck hook-up device
SODe time after

or -for

Lnt errni t t errt sampling
lead

the mask is put. on.

backpack

air

bag,

further

analysis.

of the expired

as;

air

rubber

"ill

to carr)'

pass

w i l l be directed

to

Douglas bag) for storage;

into

the atmosphere.

It may

the Douglas bag and flowneter

on a

freedom for controlling

the subject

and manipulat-

necessary.

of a trial

samples may

On completion

from the Douzl as baa to a sample pump
(;:)

valves,

b

.•

Fl owmeter s , and collection

however no company known to

t iie

use on an animal.

(1974) gives

}~ttfeld

designed

and used successfully

for this

study will
on design,

Accuracy:

probably

estimates

using

wr i t cr manufactures

be similar.

will

to construct

of cattle

particular

of the apparatus

of a face mask he

be conSUlted for

face masks have been used routinely

0 fl· t 11S

for

The mask constructed

be hired

of the energy expenditure

the effect

description

deer.

Hattfeld

commercially;

a face mask suitable

an explicit

person will

iowever , tl !C accuracy

system,

J,..

bags are available

on \'Ihite-tailed

and a suitable
Syste~s

(Brody I S.tS) .}T
collection

The aliquot

pr obab Ly a neoprene-coated

for the experimenter

be transferred

satisfactory

a1101. for a calming period

volume of expired

in the mask and valves

advice

to

to a ~Iax Planck f Iowmeter for measuring

to a l l ow greater

Hoses,

This_will

collection

open-

whil e the remainder
be p0ssihle

hose ",HI permit

valve

to the expiration

fer

periods.

A three-way

of a face mask

1\ hose attached

and tak i ng an aliquot
a collection

and expiration.

for the expiration

hegin

ing will

appar atus wi l I consist

the mask.
to obtain

and otller

animals

face mask design

on tIle energy expenditure

andof a wild

�-307-

species,

and

the) c f f cc t of t.h« appcr a tus

on the r c l a Lion of hcar t ra t e to

expcn.l i t ur e uer it cons ide r a t i on fo r t h i s study.

criD'

be evaluated

using

and

heart

of

t h:

r e sp i r ati on chanhe r s at the llet abo l i c Lab.

rate

bench tests

of equipment,

anina l s in response

to

considerations

These

observatiollS

the apparatus,

enwi 11

of the behavior
and po s s i b l y by using

Spcc i.f i c que s t i ons and pr occdur cs to he used arc as FoLl ows:
1) Accur-acy of the apparatus
tested

for

air

The valves

mat cd a ir
arise

may provide

et a l , (1969)

varied

consider~tion.

cqu i p.nent

t6 tcst

the respiration

that

lLa.sk

~!ilX Planck

and suggested

and calibration

f Lovmetcr-

that

from an upr i ght, position.

unuoresti-

fur rhcr errors

I:light

These prob l crns r equ i r e

of the meter against

on the components of the sys t emd

operation

of the who}e system by conducting

chamher.

Further

thougLt and discussion

and feasibility

of the face ma~k and other
some ~ffect

dead air

other

·f CO2 in the dead air

,
I
CO-,'_LG.

1
'L.SO

t

may be

trials

in

arc reqUired

to deter-

of such trials.
apparatus

on energy expenditure

on the suhject)s

space or resistance
space may result

compoundi np the .i mpor t anc e of these

face

the

tests

of increased

Buildup

stated

Testing

The face mask may exert
result

Immersing the

to these

mine the importance
2) Effect

for

Fade

r.ay be necessary.

In addition
possible

arc specifically

a means of chec k i ng this.

volumes at Low flow rates

if the meters

further

(l"larren E. Collins)

and can be assumed to perform satisfactorily.

apparatus
Corbett

w i l l he

ti?,:lt.ne~s underwat.er .i n a tank as cons t ruc t ion procedcs.

and DOuglas bags

gas collection
entire

- The face mask and hose connections

factors

'
.
)
prOG1}Ce
e. ps:rc I,D..1 O[:J.cc:..

_

energy expenditure

as a

of the valves

and f l owraetcr .

in an increase

in rcspiration)

C',bttfcld

1974).

These factors

r-~r
....;) in "!:.h~sub.iect,
wiri cn
,-",,) or:•...,-,.
_

�-308-

overall effect of the face mask on energy expenditure (the sum of the

.The

three above-mentioned

factors and possibly others) n~y be evaluated in the

respiration chamber using trials with and without the face mask.
Characteristics

of the experimental situation other than the face mask

might also cause nervousness reflected in increased energy expenditure by
the sub j ec t s, Amoung these are the presence of the experimenter \·:i
th the
additional B?parattls,and restraint by a leash.

The effects of these factors

are not appropriate for study in the respiration chamber ; however some
indication

of their

heart

of subjects.

to

rates

these

Lmpor t ance can be gained
If

from exami.uat Lon of t.e.Lerne t e r a d

significant

changes
at

factors

are observed.

they

changes

in energy

expenditure;

however such information

into

effects

of specific

the

Another

of the

this

rate

point

expe r Lment a.I procedures

physiological-psychological

3) Effect

cannot

in heart

in response

be re Lat.ed to
can provide

insight

on the subject's

state.

apparatus

on the heart

rate-energy

expenditure

relation

-

r at i
.
1y mentl.one
.
d· 1S tht"
consi de
eratl.O!l
prevl.ous
a

nervousness

". aSSOCl-

ated \-liththe apparatus or situation may result in irregularities in the
The above-ment i oned

relation between heart rate and energy expenditure.
procedures

(estimating energy expenditure Hith and Hithout the face mask in

the

chamber,

and examination

of heart

rate

will

provide

s orne information

on this

top Le.

is

relation.

it

correlation

If

in

response

to the

the

apparatus

does af f e ct; the

expected

that

the effect

\ViII be a non-significant

or

be tween heart

rate

and energy

expenditure

a

cys t emat i.c error
would be ccrse

changes

('ivebster

rather

1967, Br-ockway and }!cE\van 1969).

app a re n t in field

trials

to be described

further

than

2pparatus)

10'.0]

Such a result
on.

�-309These sections have described the equipment for field gas collection,
possible sources of error related to the equipment and the responses of
deer to it, and some means for evaluating the effects of the equipment
on subject per formance , Further information and examples of f'ace mask
systems for field gas collection are described by Nattfeld (1974), Mlite
and Yousef (1974), and Yousef and Dill (1969a).
Gas analysis:

Aliquots of expired air collected in Douglas bags will

be analyzed to determine percent composition of oxygen, carbon dioxide, and
possibly nitrogen, by one of the following:
a) using a paramagnetic oxygen analyzer and infrared CO2 analyzer available at the Hetabolic Lab.

Since this equipment is not calibrated in the

appropriate range for expired air obtained by these means, the oxygen analyzer will have to be recalibrated using an appropriate gas standard and
another CO2 analyzer us ed , or air samples w i ll have to be diluted in some
known proportion \.;i
t11atmospheric air.
b) similar equipment adjusted to the appropriate range may be located
elsewhere on campus,
c) it may be possible ,to gain access to suitable gas chromatogtaphy
equipment, or
d) another fairly inexpensive ($1000 - 2000) instrument of suitable
accur~cy could be purchased if necessary.. Calibration and maintenance of
such an instrument ",ould involve some time and effort, hm,ever; and this
alternative does not seem appropriate for a study to be conducted close to
the CSU campus where other equipment is available.

�-310-

lleart rate:
transmitters

External heart rate transmitters were chosen over implanted

for use in this study because no surgery is required.

Thus a

single transmitter can be used on different deer and equipment repairs can
be done without surgery.

Problems associated with external transmitters

include the necessity for a well-designed harness to hold equipment in place
and protect it from being chewed or otherw i se damaged, and the possibility
of interference of cardiac signals by electrical potentials from muscle
movements

(.Jacobsen, 1973).

Other
affixed

equipment

to the

physiograph

for

monitoring

animal's

body,

heart

rate

an PM receiver,

"lith

at

least

means of marking

off

time. (Narco Biosystems).

Respiratory rate:

a channel

for

includes
a coupling

recording

surface

electrodes

device,

and a

the EKGtrace

and a

Respiratory rate may be deternine~ during trials.

Equip-

mffiltfor monitoring respiratory rate concurrently with other measurements will
be investigated if time permits.
Trials:

It is expected that the problems related to equipment outlined

in the previous section can be solved.

If this is true, the face mask and

associated apparatus w i Ll be used to investigate t:1erelation between the
heart rate and energy expenditure of fawns engaged in activities as fo1101-1s:
1) bedded
2) limited actiyity - standing and ",alking
3) increased activity - wa lk i.ng more and trotting
Four or more trials with each subject at each activity level Idll be
conducted during an intensive study period of three of four wceks , A set
of observations will be taken for all deer at all activity levels and then
another complete set taken and so on. to minimize confounding of the effects
of different activity levels or other variables w i th any training effect
(E.E. Remmenga. pel's. comm.).

�-311-

Length of trials

~lll

tolerate

the apparatus,

EKGtrace

will

also:

be determined by the anount of time deer will

bct .•
vcen 20 and 60 minutes if possible.

b~ recorded

continuously

be rccor~ed using telemetry~

specified
larly

t;me period

throughout

at suitable

to heart

trials.

These variables

rate

include

or auditory

Feed intake

not be restricted

'l11ese and other
state

factors

Subjects

or by an observer.

converted

stimuli,

level

,,;hich might affect

the 51lbject 1 s physiological-

by the experimenter

barometric

deficit

whet.her these stimuli

elicit

a response in the subject

stimuli

subject.

The next trial

Douglas bag.

Expired air

pressure,

apparent

variable

the face mask apparatus

wi l I use a different

tempe.rature,
relative

subject

deer nrnlber,

to the observer,

or not,

the observer uill

a buildino-D to avoid introducincr
.
D an additional
completion of a trial

using a tape recore

(Hoen 1973), cloud cover,

and auditory

If possible

exposure

to or during trials.

Date, time of day, length of trial,

to vapor pressure

its

during these

of feeding,

at any timeprevious

near the subject,

w.LLl, be recorded.

or affect

and time (age of animal and season).

visual

On

be weighed regu-

as much as possible

beh.av.io r of subject,

behavior

will

energy expenditure

thermal factors,

\~ill be recorded

wind speed and direction
humidity

intervals.

will be controlled

to unusual visual

recorder

may

or by counting valve movements for a

which may increase

relation

psychological

and respiration

the study pericd.

Other variables

"ill

during a trial,

The subject's

and experimenter

be stationed

inside

into the experiment.
w i I l be r-emovedfrom the

and a different

may be removed from the Douglas bags to sample pumps

�-312-

for temporary storage or transport, pcrmittinc reuse of the Douglas bags.

A

third person may be required to transport expired samples to a technician for
analysis.
Analysis:

Energy expenditure (heat production) in kilocalories \·;ill

be calculated from oxygen consumption and carbon dioxide production as outlined
by Brody(l94S) or by another method.

These values \\'i11be converted to a

-''
. mctalo
1 I'1C bO(y
1 wei~th
rate 0f energy cxp~n~1ture
per unIt

(keal / kg 0.75 -hr)

The EKG trace Hill be divided into one-minute intervals; the numb er
of 'QRS peaks (heats) will be counted for a random samp le of these interval
and variances will'be calculated.
Rates of energy expenditure \Yillbe plotted as dependent variable against
mean heart rates for each deer over all activity levels, and correlation
coefficients will be calculated.

A similar plot obtained by Silver (1973) for

pooled data from deer in respiration chambers is shown on page 313.
At this point a decision w i Ll he made about pooling the dat a from different
suhjects.
The usefulness of hear-t rate to predict energy expenditure w i H be tested
in appropriate models using standard regression techniques (Draper and Smith
19(6).

Step\Yise regression procedures will be used to evaluate the effects

of other variables entering the model in.addition to heart rate, and interactions
of these variables with hear t rate wi Ll be tested for significance.

Variables

to be considered include subject, activity level, temperature, nervousness
(using numbered levels or dummy variables), and any other of the factors
measured "hich appear to be meaningful.
If time permits, pulmonary ventilation rate (data obtained

2.S

par t of

th.e gas collection procedures) and respiration rate (if data \Vere collected)
\dl1 be used in separate analyses similar to those made for heart rate, and

�...', -.

,

( .'

t
"",

"

•

...•.

••••
_.

•.....

'.!

-.__

.••••
~._ •••••. _ .•..•..••.•••.•••.
_.••.••.•
__••
_ .•.•
~ _0., •....•.•...• _ •. ".&lt;.

.'."

.,"

...
-,

_- .•...~- .'_. :.:'.:'

, . .:'" -, ~...
....

,. •.•••.

__._

_.'"

•..

'.

..

. ..•••
::;;

,"

' . .-,_

-313-

•..••
'_

:. ' •. ,

••.•

., .:.:

~.

•.. :.-.;---.:.

;-

• i-

~I

v

"

.;"'1...

~- .
',-

......

.-

\_ ....-

•••• ..: ••••• ; •

,
,

....
"

',.
.

,.-

'.'"

,- ....

\~

\

.-

.,,::

-

I

I

I
I
i

\

I

i

j

i,

,j
I

j

I
,!

I
i

i

I
!

1

i
1

I

I
;

,
i

I

i

oj

,.

\..

s;

r-

!'in

;

, vi
I

i .t~
&lt;t

I ....,.~

II

j

\

.'\

�-314-

in a multiple regression with heart rate.
The usefulness of these relations for predictive purposes will be evaluated
and confidence intervals will be calculated where appropriate.
Additional experiments:
heart rate ~ight

be

If the expcr iments outlined above su~gest that

a useful

estimator

of energy expenditure in un-

restrained deer f'awns , and if time allOlvs}add i t i.onal experiments relevant
to the use of the technique may be conducted to assess the importanc.e of

f actors

other than acti vity as fo l.Lows:

l) Conduct several trials using one or more subjects in very cold thermal
environments, \Vith other conditions as in previous trials and at a single
activity level.

Determine

\Vhether these data points fall into the appro-

priate confidence intervals for an individual energy expenJiture at a given
heart rate.
Introduce v i.sual and auditory stimuli .wh i ch elicit a behavioral or

2)

heart rat e response in the subj ec t , w i th other cond i tions as ou t lined before
and at a single level of activit~
3) Conduct sevcrn l trials at nighttime wi th other conditions as before

and at a single activity level.
4) Webster

(1967) suggested that the relation between heart ratc and

energy expenditure of iridividual sheep could he used for predictive purposes
over a period of at least three wceks .

However, Silver (1973) pooled data

obtained from a number of deer year round, to obtain a relation between
energy expenditure and heart rate with a fairly high correlation coefficient
of .805.
Significant differences attrihuted to season have been demonstrated in
fasting metabolic rate (Silver et al. 1%9),
activities
deer.

in heart rates over different

(Jacobsen 1973), and in the heart size (Anderson et aJ.. 1974) of

It scems reasonable that such changes, as we lI as growth, might alter

the relations amon~ rates of hody functions.

Thus there is reason to doubt

�-315-

that the relation between heart rate and energy expenditure remains constant
over long periods even in one animal.
To investigate the usefulness of a heart rate - energy expenditure
relation over time, weckly trials will be run on one or more subjects over a
period of three months including the intensive study period, with other conditions as outlined before and at a single activity level., If time permits) trials
will also be conducted

twf.ce

a month throughout the fawns '

first year.

Target dates:
30 November 1975 - Study plan submitted
31 December 1975 - Study plan evaluated and modified if necessary
31 Barch 1976 - Equipment and procedures tested
31 Octoher 1976 - Fawns weaned and trained
31 January 1977·- Inteilsive study period completed
31 Nar ch 1977 - Data analysis comp let ed
31 .Iu Iy 1977 --Thesis completed

Responsibility:
person-days
Investigator

180

w.w. Mautz - Consultation

30

H.A.

L.Il.

Kaut z -

Ca rperrter

- Consultation

10

Cost estinates:
Total segr.:entcost (1 January 1976 - 31 Harch 1976) - $4845
Pe r s on a I se rv l ce s

H.A. Kautz - GrE-duate Res. Asst.
Face nask design and construction
Fl owme t er calibration
Eguil)ill("r;
t_
Face mask materials, hoses
Valves
Dougl.2.sbags (4)
Backpack apparatus'
Fl owme

t.o

r

Sample pU8pS (4)

12 months

$325

~ 975
800
200

$1975
$ 360
250

720
1;0

900
600
---~$2870

�Total

segment cost

-3161976 - 31 Harch 1977) -

(1 April

Personal services
N.A. Kautz - Graduate Res. Asst.
Student Assistant
- l"'aising and training
St udcn t As s i.s t an t s (2) - trials
His cellaneous technical help

$1'5 &gt;9,15"

12 months
3 months
1 month/

f awns

$325
560
560

$3900
1650
l120
~OO

$7270
Qperating supplies.and
services
Respiration
charnbe r trials
Gas analysis
Veterinary
supplies
Milk
Feed, hay, bedding
Computer time
Vehicle (Co-op Unit vehicle - gasoline
Photographic supplies
Shipping expenses

$ 200
600
50
150
200
200
250
100
200
$1950

only)

E ~1.~:~!J"!m2n
~
Heart rate transmitters
(5)
Receiver
Physiograph
Time and event channel
Recording channel
DC-ACcoupler
Surface electrode kits (5)
Ha rnes se,s.•
Le ashes
Valve replacements
Respiration
rate apparatus

Total
Personal

segment cost

$1625
780
895
195
680

2/10
90
50

500
1600
--$6655

.(1 April

i977 - 31 July

services
- G:::aduate Ras , Asst.

~1.A. Kautz

1977) - $1500

4 months

$325

Operating supplies ;lnd services
Thesis typing, graphics and duplication
E.

$ 200

Location
All work \'1ill be conducted at the Foothills
or at ano therjappropr i.at e Division

F.

$1300

Rel a t cd federal

of Wildlife

projects

intlude

wi nt er activities

of deer confined

which ,,'ill

to use heart

attempt

will

facility

near Ft. Collins.

a study directed

toward quantifying

proj ects:

R.elated federal

studies

Deer Pens, CSUFoothills

in large pens,

rate

and a proposed study

of deer to indicate

be conducted in Middlo Park, Colorado.

stress.

Both

Campus

�-317Literature Cited
Anderson, A.E., D.E. Ned i n , and D.C. Bowden,

1974.

Growth and morpho-

metry of the carcass, selected bones, organs, and glands of mule deer.
Wild!. Nono g , 39.

The Wildlife Society, \'iashington,D.C.

Blaxter, K.L., and J.P. Joyce.

1963.

l22pp.

The accuracy and ease with which

measur ements of respiratory metabolism can be made w i t.htracheosto-

mized sheep.
Brockway, J .~I.,

and E.H. Mcfiwan ,
in the sheep.

fornance

Brody, S.

Brit. J. Nutr. 17:523-537.

1945.

New York.

1969.

Oxygen uptake and cardiac per-

J. Physiol. 202:661-669.

Bioenergetics and growth .

Reinhold Publishing Co.,

l023pp.

Corbett, .J.L., D.J. Farrell, R.A. Leng, G.L. HcClymont, and B.A. Young.
1971.

Determination of the energy expenditure of penned and grazing

sheop from estimates of carbon dioxide entry rato.

Br. J. Nutr. 26:277-

291.

Corbett, J.L., R.A. Leng, and B.A. Young.

1969. Measurement of energy

expenditure by grazing sheep and the amount of energy supplied by
volatile fatty acids produced in the rumen.
Blaxter, J. Ki.elanowsk i , and G. Tho rbek , eds ,
farm animals.
Datta, S.R.,

Pages 177-186 in K.L.
Energy expenditure in

Oriel Press, Ncwcas t Le -upon-Tyne ,

and N.L. Ramanathan.

1969.

Energy expenditure in wor-k pre-

dicted f rom heart rate and pulmonary ventilation.

J. Appl. Physiol.

26(3):297-302.
Draper, ~.n.J and H. Smith.
Wiley &amp; Sons, New York.
Jacobsen, N.K.

1973.

white-tailed deer.

1966.

Applied regression analysis.

John

407pp.

Physiology, behavior, and thermal transactions of
Ph.D. Dissert.

Cornell Univ.

346pp.

�-318-

Lifson,

H., G.B. Gordon, and R. ~!CClintock.

.
carbon

dioxide

19S5.

~1easurement of total

°18 J. Appl. Phys ol , 7 (6) :

by means of O2

production

i

704-710.
Lifson,

!L, and R. ~1c.Clintock.

1966.

of body wat er for measuring
BioI.

Theory of use of the turnover

energy and material

balance.

J.

rates

Theoret.

12:46-74.

Halhotra,

}l.S.,

S.S.

S.N. Ray, and T.N. Shrivastav.

Ramaswamy,

Minute ventilation

as a measure of energy expenditure

J.

17(S):77S-777.

App l • Physiol.

Ma Ihot ra , H. S. ,J.

of energy

Sen.Gupta,

and R.M. Rai.

expenditure.

Mat t fe Id , G.r.

1974.

J.

1963.

Appl. Physiol.

The energetics

Pulse

1962.

during

count as a measure

18(5):994-996.

of wmt.or

foraging

by whi t e-f.a.i.Led

deer

- A perspective

on wi nt er concentration.

Hniv.

of !J.Y.,

ColI.

of Environm. Sci , and 1'01'., Syracuse.

1973.

Wildlife

N. L.

1964.

Mocn , A.!L

Ecology.

exercise.

Ph.D. Thesis.

St.
306 pp.

W.I1. Freeman and Co , , San Francisco.

458pp.
Ramanathan,

from respiratory
Sharkey,

B.J.,

pulmonary

J.F.

Reliability

frequency.

of estimation
J.

of metabolic

Appl. Phys i.ol , 19(3):497-S02.

HcDonald, and L.G. Corbridge.

ventilation

levels

as predictors

1966.

Pulse rate

of human energy cost.

and

Ergonomics

9(3):223-227.
Silver,

II.

Prog.

1973.

Laboratory

Rep, , Proj.

H.,

Webster,
eator

A.J.F.

of whi to-tailed
1967..

of the energy

P-R •.
lob

New Hampshire Fish and

18pp.

N.F. Col ovos , J.B.

metabolism

of game and game foods.

W-S1-R-6, Jobs 7 and 8.

Game Dept , , Concord.
Silver,

analyses

Holter,

and H.H. Hayes.

deer.

J.

Continuous
expenditure

Wildl.

Fasting

Manage, 33 (3).190-493.

measurement of heart
of sheep.

1969.

Br.

rate

as an indi-

J. Nutr. 21:769-78S.

�-319-

White,

R.G.,

ano. R.A. Leng.

1968.

of ener-gy

expenditure

in lambs.

R.G., ano. M.K. Yousef.

\fuite,

Carbon dioxide
Proc.

of reinc1eer-caribou

in nutritional

and environmental

AT(45-J)-2229,

Un i v . Alaska,

\'Iync1ham, C.lI.,

during

strenuous

B.A.,

Voting,

in Studies

adaptation.

AEC Prog.

Haximum oxygen intake

wor k ,

J.

App1. Phys i o l.

rate.

Pages 435-436

eels.

Energy expenditure

J.

interest

Rep.,

Contr:

Peter,

and

and maximum heart

rate

l4(6):927-9:~6.

G.L. };cClYJi1ont, and J.L.

expenditure

entry

7:335-341.

on the nutrition

in Alaska w i t.h special

1959.

of energy

as an index

Anim. Prod.

Mari t z , .1.F. \Iorrison,

R.A. Leng , R.G. \'lhite,

Estimation

rate

Fairbanks.

t~.B. St rydom , J.S.

Potgicter.

Z.ll.

Aus t , Soc.

1974.Page38-42

and metabolism

entry

from measurements

Corbett.

of carbon

1969.

dioxide

in )~.L. Bl axt.cr , "J. Ki e l anowsk i., and G.Thorbek,

in farm animals.

Oriel

Press,

Newca s t I e-

upon-Tync.
Yousef,

~I.K., and D.H. Dill.

respiratory

activity

1969a.

in the hurro

Resting

energy metabolism

Eqnus asim;~.

J.

Appl.

and cardioPhys i ol , 27 (2):

229-232.
Yousef,

M.K.,

and D.n.

Dill.

1969b.

man and bur ro !:quus asinus.

J.

Energy expenditure
Appl.

in desert

Phys i ol . 27(5):681-633.

walks:

�-320-

A P PEN

D I X

B

�-321-

THESIS

ENERGY REQUIREMENTS

OF

MULE DEER FAWNS IN WINTER

Submitted by
Dan L. Baker

In partial

fulfillment, of the requirements

for the Degree of Master

of Science

Colorado State University
Fort Collins,

Colorado

Spring, 1976

�-322-

COLORADO STATE UNIVERSITY

Spring,

WE HEREBY

RECOMMEND

THAT THE THESIS PREPARED

UNDER OUR SUPER VISION BY DAN L. BAKER
REQUIREMENTS

OF

MASTER

ENTITLED

OF MULE DEER FAWNS IN WINTER

TED AS FULFILLING

IN PART REQUIREMENTS

OF SCIENCE.

Committee

Advisor

Head of Department

1976

on Graduate

Work

ENERGY

BE ACCEP-

FOR THE DEGREE

�-323-

ABSTRACT OF THESIS

ENERGY REQUIREMENTS OF
MCLE DEER FAWNS IN WINTER

Two winter feeding trials

were conducted in Middle Park,

Colo-

rado in an attempt to quantify the energy demands of mule deer fawns
(Odocoileus hemionus
prevailing

tum,

hand-reared

of ad libitum,

pelleted 'concentrate
trials

mately 72 percent

timating

of ad libi-

of ad libitum) and fed a

digestible

Twenty-four

and metabolizable

Apparent dry matter

digestibility

and not affected by level of intake,

or phys iological

in vivo

condition .. A linear

Colo-

nutrient

was approxiambient temp-

reg res s ion of feed intake

daily body we ight gain or loss provided the bas is for es-

maintenance

weight measurements
these trials

25 percent

75 percent

ration (3.0 ME kcal/gm).

the apparent

content of the diet.

and average

(ad libitum,

were conducted in Fort Collins and Kremmling,

rado to estimate

e rature

pens exposed to the

mule deer fawns were randomly

to one of four treatments

50 percent

digestion

held in isolation

winte r environment.

Twenty-five
assigned

hemionus)

require

energy requirements.

Feed intake and body

from two winter trials

suggest that the fawns m

a metabolizable

WOo75/day to maintain

energy intake of 157 kcal/kg

body weight homeostasis.

�-324-

Body weight change was linearly
ly energy)

intake.

cody weight.

All ad libitum deer either

starvation

pr ior to death (1- 3 days),

(blood urea nitrogen,
to energy

Weather
ments.

non-esterified

Mean

(P

the blood energy metabolites
free fatty acid) showed no r e la-

status.
were similar

for both winter experi-

Ambient temperature

ranged from

-30

from

0.05)

Except for BUN concentrations

measurements

speed varied
exceeded

body weight.

for these fawns was significantly

than those of ad libitum deer.

tionship

gained or maintained

after los ing 32 percent

level of blood urea nitrogen

collected

to feed (and consequent-

Four of six fawns on the lowest level of energy intake

died of apparent

higher

related

0 to

13.5

meters

to

+11 C.

Wind

per second (oops) but seldom

4.5 mps.

Dan L. Baker
Department of Fishery and Wildlife
Biology
Colorado State University
Fort Collins, Colorado 80523
Spring, 1976

�-325-

ACKNOWLEDGEMENTS

Funding,
search

pen facilities

were provided

Colorado

Federal

equipment,

by the Colorado

Aid Project

computer

service

the Rocky Mountain Forest
Forest

animals

for this re-

Division of Wildlife through

W-38-R.

The laboratories,

and office facilities

and Range Experiments

vehicles,

were provided
Station,

by

USDA,

Service.
Appreciation

tee:

and experimental

Dr.

is expressed

to members

of my graduate

J. G. Nagy, major pr ofe s s o.r; Dr. O.C.

D. E. Johnson for their guidance,

commit-

Wallmo and Dr.

advice and critical

review of this

manuscript.
In addition,
persons

appreciation

for the ir contribution

L. H. Carpenter,
P.

sincere

is extended to the following

to this study:

Mr. D. W. Reichert,

Mr.

Mr. R. B. Gill,

Dr.

W. L. Regelin and Mr.

F. Gilbert.
Last but foremost

express
wife,

my gratitude,

Julie.

and without the words available
I acknowledge

to adequately

the many contributions

of my

�-326-

TABLE OF CONTENTS

ABSTRACT OF THESIS

323

ACKNOWLEDGEMENTS

325

LIST OF TABLES

328

.

LIST OF FIGURES

330

Chapter
1

INTRODUCTION

331

II

LITERATURE

3%

Energy

REVIEW.

metabolism.

Maintenance

e ne rgy

338

Thermoregulation

341

Blood metabolites

343

Weather-severity
III

334

index.

. • . .

346

METHODS AND MATERIALS

349

Study location

349

.

• .

Phys i ca l facilities

350

Acquisition

of fawns

350

Re ar ing and training

353

Experimental

design - 1974

356

Experimental

design - 1975

360

Body weight changes
Blood metabolites
Weather

362
.

363

measurements.

. 364

�-327-

Chapter
IV

Page
RESULTS

AND DISCUSSION.

Growth rates

365

Intake and food utilization.
Body weight changes

V

. .

368
.

376

Maintenance

require ments

377

Blood energy

metabolites

389

Environmental

conditions

391

SUMMARY

LITERATURE
APPENDIX

365

CITED
.....

AND CONCLUSIONS

395
398
403

�-328-

l.JST OF TABLES

Table

Page
Compos ition of milk formula fed to fawns
in 1974. . . . . . . . . . . . . . . .

354

2

Composition

355

3

Results of 12 digestion trials conducted in Fort
Collins, Colorado to evaluate the basal ration in
the 1974-1975 winter trials
.

359

Age, sex and initial body weights of the fawns
for each treatment in 1974 . . .. . . . . .

369

Linear regression
equations and correlation
statistics
relating mean daily energy intake (x)
and average daily body weight gain or loss (y)
for the 1O-week experiment of 1974 . . . . .

371

Age, sex and initial body weights of the fawns
for each treatment in 1975 . . . . . . • . .

373

Linear regression
equations and correlation
statistics
relating mean daily energy intake (x)
and average daily body weight gain or loss (y)
for the 7 -week experiment of 1975. . . . . . .

375

Linear regressions,
correlations,
ME maintenance requirements
and mean wind-chi.ll for the
first six weeks of the 1975 winter trial
....

383

Maximum, minimum and mean daily temperature,
humidity, wind speed and associated wind-chill
index for the period of January 14 to March I,
1975 . . . . . . . . . . . . . . . . . . . .

385

Means and standard er ror of blood urea nitrogen
concentrations
for fawns on three different levels
of dietary energy for each week of the 1974
ex pe r iment.
. . . . . . . . . . .

390

1

4

5

6

7

8

9

10

analysis

of pelleted

basal ration

�-329-

Table
11

12

13

14

Page
Means and standard error of blood urea nitrogen
concentrations
for fawns on two different levels
of dietary e ne rgy for each week of the 1975
experiment
. . . . . . ..

390

Means and standard error of plasma free fatty
acid concentrations
from fawns on three different
levels of dietary ene rgy, for each week of the
1974 winter trial
. . . . . . . . . . . .

392

Means and standard error of plasma free fatty
acid concentrations
from fawns on two different
levels of dietary energy, for each week of the
1975 winter trial
. . . . . . . . . . . .

392

Maximum, minimum and mean daily temperature,
humidity, wind s pe ed and as s oc Iate d wind-chill
index for the per iods of January 8 to March 28,
1974.
. . . . . . . . . • . . . . . . . . .

393

�-330-

LIST OF FIGURES

Figure
1

Page
Pen facilities,
Center

Junction Butte Deer Research
. . . . . . . . . . . . • . .

351

ofLso lat ion pens and handling facilities

352

2

Details

3

Mean weekly weight changes (percent) compared to pre-trial
weights of fawns on three
different levels of dietary energy, 1974
experiment.

4

5

6

7

8

. . . .

378

Mean weekly weight changes [per cent] compared to .pre-trial
weights of fawns on two
different levels of dietary energy, 1975
experiment.
. . . .

379

Regression of average daily body weight gain
or loss on average daily metabolizable energy
intake, 1974 experiment
.

381

Regress ion of average daily body weight gain
or loss on average daily metabolizable energy
intake, 1975 experiment
.....

382

Effect of wind-chill
energy maintenance
experiment.

386

index on metabolizable
requirements,
1975
. . . . . ...

Regression of average daily body weight gain
or loss on average daily metabolizable energy
intake, 1974 and 1975 experiments
combined.

388

�-331-

CHAPTER I

INTRODUCTION

Energy

is required

by all living organis ms to maintain

input of ene rgy tcran animal is through the metabolic
ingested

food.

The plant constituents

life.

The

transformation

of

consumed by an animal are

broken down, releas ing energy which is used by the animal for bas ic
body functions

(growth,

fetal development,

under adverse

climatic

conditions).

energy is therefore
value.

of great

in determining

the caloric

is of equal ecological

requirement

under different

standing the demands

importance.
natural

and heat

its nutritive

contribution

co mrnun ity , knowledge of the energy requirements
mers

activity

The ability of a plant to supply

importance

In addition to understanding

lactation,

of the plant consu-

The determination

conditions

of the plant

is a first

of this

step in under-

an animal makes upon its environment

and vice-

versa.
Little is known regarding

the energy requirements

nants or how well the environment
Quantitative
dependent

on understanding

deer nutrient
forage.

determination

requirements

supplies

of wild rumi-

those requirements.

of range "carrying

capacity"

the complex interaction
and the fluctuating

for deer is

of the ever-changing

nutrient

supply of range

�-332-

A major problem

confronting

hemionus)

dur ing winter

adjustment

of caloric

meet total energy

1S

mule deer (Odocoileus hemionus

heat conse rvation and the compens atory

intake in relation

requirements

to cold stress.

continues

mortality

If failure

to

must ultimately

occur.
Mule deer fawns are probably the age class
under-nutrition

and mortality

during winter.

most vulnerable

During this period the

fawn can no longe r depend on the doe s milk as a major source
I

energy.

It must meet its physiological

primarily
mulated

from the available
body tissue

stores,

knowledge of the postweaning

to

and homeothermic

of

demands

energy supply' of range forage and a ccuwhich are meager
maintenance

at this age.

Therefore,

energy requirements

of

fawns during winter would provide a portion of the basic biological
information

requisite

for an assessment

of range carrying

capacity

for

mule deer.
There

is little quantified,

getic relationship
their

information

of mule deer fawns to the climatic

winter ranges.

the digestible

phys iological

The purpose

and metab olizable

conditions

of this study was to:

energy requirements

of mule deer fawns during the winter months (January

on the enerof

(1) estimate
for maintenance 1
to March) by

1For the purpose of this experiment,
maintenance is defined as
the amount of energy intake necessary for body weight equilibrium
and includes energy cost for basal metabolism,
activity, and thermo-'
regulation.

�-333-

us ing bo/dy weight changes as an index of energy status,

(2) determine

the minimum dietary

energy

of mule deer

fawns during winter,

(3) estimate

requirements

time to moribundity

given levels of deficit energy intake,
bolites

(non-esterified

if they reveal

physiological

and (5) measure
humidity,
balance.

free-fatty

weather

states

for survival

(4) measure

for fawns on

certain

blood meta-

acids and blood urea nitrogen) to see
resulting

parameters

from energy intake levels,

(ambient temperature,

relative

wind s peed and chill index) to see if they influence ene rgy

�-334-

CHAPTER II

UTERATURE

Energy

REVIEW

Metabolis m
The minimal

absorptive

energy demand by an animal at rest

condition with complete

bient climate

has been described

The basal metabolic

rate

inactivity

gradients

circulation,

osmotic

the energy

spent as basal metabolism

(Crampton

basal metabolism

and Harris,

activity)

and production

gether

these

animal

(Gasaway and Coady,
The measurement

activity

and i s therefore

with domestic

energy

species.
expended

requirements

gestation,

of basal metabolic
a difficult

metabolic

in standing

body
above

(thermoregulalactation).

the total energy requirements

rate requires

parameter
Therefore,

used as the base-line,

Fasting

muscle tone

Toof an

1974).

and wild animals.

(FMR) is normally
these

Energy

(body growth,

constitute

excretion,

being spent on normal

include energy cost for maintenance

tion,

1945).

The major portion of

goes to maintaining

1969).

am-

of the ca lo -

respiration,

and muscle tone.

with the remainder

processes

(Brody,

(BMR) of an animal is a measure

secretion,

functions

in a "comfortable"

as basal metabolism

ric r equ ir e me nt needed to maintain

and body temperature

in a post-

to measure

minimal energy

postural

in-

when dealing

fasting metabolic

rate exceeds

and small

complete

rate

cost for

BMR by the amount of
movements

during

�-335-

measurement
ferred

(Silver et al.,

1969).

over BMR by Kleiber

provided

a more accurate

quirements

Fasting

(1961) because

measurement

that the difference

small among domestic
to stress

fasting metabolic

voluntarily

but greater

imposed by confinement.

Kleiber

BMR of adult homeotherms

equation 70 WOo 75 where

rate

inactive.

reB'Iaxte r'

between BMR and FMR may be

ruminants

the average

rate was pre-

of the mini murn energy

of an animal unable to remain

(1962) postulated

metabolic

among wild species
(1961) demonstrated

could be calculated

BMR = kcal/day

due
that

from the

and W = body weight in

kilograms.
Several

studies

have shown that wild ruminants

FMRs above this suggested

inters pecies

FMRs of eland (Taurotragus

oryx) and wildebeest

to be 104.3 and 111. 2 kcal/kg

mean.

generally

Rogerson

(1968) fcund

(Connochaetes

WO. 75/ day, respectively.

have

sp.)

Silver et al.

(1969) found fasting heat production

(a measure

tailed

to be cyclic with the average

FMR

WO.75/day

and

deer

(Odocoileus

of adults in summer

virginianus)

coat to be 143 (se ± 5.43) kcal/kg

in winter coat to be 96 (se ± 3.66) kcal/kg
variation

was attributed

to the insulative

winter and pos sible evolutionary
scarcity.

of FMR) of adult white-

WO. 75/ day.

quality of the hair coat in

metabolic

The FMR of red deer (Cervus

This seasonal

adaptation

to winter food

elaphus) was measured

every

two months for one year by Brockway and MaIo iy (1967) and was calcu1ated to be 90 kcal/kg
in heat loss.

0.75
W
/day

with no evidence

Hammel (1962) calculated

of an annual rhythm

the fasting heat production

�-336-

(FHP) of resting
day.

(Rangifer

McEwen (1970) reported

caribou
day,

reindeer

(Rangifer

tarandus

tarandus)

the metabolic

rates

WO.751

of fasted female

to be 91and 1)2 kcal/kg wO·751

groenlandicus)

which was 20 to 30 percent

at 132.8 kcal/kg

higher than the quoted inters pecific

mean of 70 kcal 1 kg W0.751 day.
In general,
measure

these

observations

ments of wild ruminants

and within animals
tend to be higher

of the same species,
than those reported

80 kc al (Blaxter,

.
of 70 kcal/kg

0.75
W
Iday (Kleiber,

is not entirely

buted to psychological
state,

insulation,

confinement
reported

effects

metabolic

(2) FMR of wild ungulates

for sheep,

1961).

55-60 k ca l , steers,

for these dif-

Some of the differences

may be attri-

stres s, activity,

reproductive

and age of the animal be ing measured.

The

on an animal are difficult to quantify,

but the higher

rates

of domestic

between these two groups

of wild ruminants
an inherent

over the rates
difference

(Whitt ow , 1971; Moen,

in cell metabolism

1973).

The age of an animal has been shown to increase
Ritzman

tion of subadult

mean value

The reason

and phys iological

ones should not be considered

production.

metabolic

between species

1962), or the inters pecific

known.

nutrition

that (1) fasting

vary cons iderably

approximately

ferences

suggest

and Benedict

(1930) measured

sheep and found FMR to decrease

from

birth until the animals

cattle

appeared

to reach

were six months old.

relatively

eight months of age (Ritzman

stable

and Colovos,

the heat produclinearly

with age

Young, growing

FMR after
1943).

its fasting heat

approximately

A similar

response

�-337-

was reported

for barren-ground

caribou

that the FMR of a 9-month"'old
.
/
0.75/
102 .5 kcal kg W
day.

by McEwan (1970) who found

female after six days of fasting was

Fasting heat production

antelope

(Antilocapra

averaged

9 2 and ranged from 6 1 to 110 kcal / kg W0.75/ day (Wesley et

al.,

1970).

(Odocoileus

americana)

of young, pronghorn

The resting
hemionus

heat production

columbianus)

ment but not in a post-absorptive
(1970).

ranging in age from 108 to 182 days,

The mean resting

of eight black-tailed

fawns in a thermoneutral

state was measured

heat production
/

heat production

was increased

and nutrient

metabolism.

Silver et al. (1969) reported

white-tailed

deer fawns in winter coat to be 90 (se

WOo75 /day.

Thompson et al.

due to heat of fermentation
the FHP of

± 4.49) kcal/kg

(1973) showed the FHP of white-tailed

deer fawns to vary with season.

The values for October,

January,

May and August were 112, 77, 104, 145 and 166 kcal/kg

0.75
W
/day,

respectively.

'The higher

FMR reported

could be a significant
maintenance

factor

requirements

energy

is a composite

activity

and heat increment,

ments would therefore
animal

by Nordon et al.

.
Smce
t h ese fawns were

not fasted,

March,

environ-

of these fawns was calcu-

lated at 147.00 (se ~ 1.52) k cal kg W0.75/ day.
..l

deer

for young animals

contributing

r e lat i.ve to adults

to the relatively

for this age class.

higher total

Since maintenance

of basal metabolis m plus thermoregulation,
any increase

increase

such as a deer fawn.

in basal metabolic

the total energy demands

This higher

maintenance

require-

of the young

requirement

in

�-338-

combination
contribute

with low environmental
to high fawn mortality

ity of winter

forage

Maintenance

Ene rgy

Forage

temperatures

in areas

could theoretically

where the quantity and qual-

is low.

consumed

must be converted

to a metabolically

useful

form before it can be of value to the animal for various

body functions .

Knowledge of the subsequent

would provide

an understanding

of the chemical

and its range (Moen,
The First
neither
fore,

conversion

of food material

communication

1973).

Law of Thermodynamics

be created

between the animal

nor destroyed,

when a plant substance

states

that "energy

but only changed in form."

is burned completely

chemical

There-

to CO , H 0 and
2
2

other gases,

the stored

heat energy,

and the amount of heat given off is cons ide red the gross

energy

content of t hat substance.

stance

is usually

kilocalories

measured

per gram.

mon food substance
be available

The gross
apparent

in the plant is converted

The gross

the gross

does. not, in itself,

content of a sub-

and is expressed

energy

because the efficiency

ener gy consumed

the food minus the heat of combustion

in

content of a com-

of its use by an ani-

(Church,

1971).

by an animal can be partitioned

and metabolizable

to

indicate the energy that would

and less than 100 percent

digestible

energy

in a bomb calorimeter

However,

to the animal

mal is variable

energy

can

energy.

The gross

into

energy of

of the feces is called apparent

�-339-

digestible

energy

(DE).

It represents

efficiency

of food digestion.

a first

approximation

of the

A portion of the gross energy consumed

is also lost in the urine and in gaseous

products

the heat of combustion

is accounted for and subtracted

from gross

energy intake,

lizable energy
available

of these losses

(ME).

the calculation

Metabolizable

of digestion.

is called apparent

energy is a measure

to or needed by an animal at a particular

tabolic requirements

tissue

and its use for the above body functions requires

The conversion

of a portion of this ME called the heat of nutrient
Metabolizable

and various

of ME to body
an expenditure

energy les s heat increment

forms of production

syn-

metabolis m or heat

true net e ne r gy (NE) and is that portion available
maintenance

of the energy

and may be used for work, heat and tissue

(Maynard and Loos l i , 1969).

(HI).

metabo-

time for all me-

thesis

increment

When

is termed

to an animal for

and activity

(Moen, 1973;

Maynard and Loos l i , 1969).
The concept of heat increment
therms

subjected

is important

to low environmental

to ruminant homeo-

temperatures.

The heat ex-

pended for nutr ient metabolis m comes from two sources
ruminant:

(l) heat produced in the digestive

tract

as a result

of mi-

(2) heat produced via intermediary

metabolism

as a

crobial

action,

result

of absorbed

energy is dissipated
hypothermal

in the

nutrients

(Crampton

and Harris,

1969).

This

without use unless the animal is in a critical

environment.

Under these conditions,

the heat increment

�-340 ....-

is used to maintain
energy

body temperature

for maintenance

Maintenance

and becomes

part of the net

(Moen, 1968).

requirements

for wild and domestic

species

have

been estimated.

Short and Golley (1968) concluded that maintenance

energy demands

fo r cattle

and perhaps
percent

(Bos taurus )in

for wild herbivores

above FMR.

quirements

Blaxter

of ruminants

Brody (1945) suggested

mated metabolizable
reindeer

under "usual"

36 percent

that maintenance

the maintenance
0.75/d
Way,

Ullrey et ale (1970) calculated

first year of life.
estimated

requirement

kg WO.75/day.

respiration

ion analysis

Maintenance

deer to be 146

Thompson et ale

calorimetry

of white-tailed

The mean apparent

by regress

wh ite-taile d does.

d 15 2 kcal / kg W0.75/ day

did not differ significantly.

from indirect

energy

to be 131 and 158
kcal/kg
,

of six adult white-tailed

.
. h t wmd
.
sm
k ca l / kg W0.75/ day with
a shg
Clan

maintenance

of a 70 kg

below 10 C, Holter et al. (1975) found ME

requirements

These values

The esti-

per day or about 200

1y. f or wi.,nte r ing , pregnant

At ambient temperatures

(1973) determined

energy require-

for maintenance

ME and DE requirements

.
respective

maintenance

1970).

re-

than FMR.

twice BMR.

in winter amounted to 5.5 megacalories

kcal / kg W0.75/ day (McEwan,

without.

greater

was approximately

is 15

that maintenance

digestible

energy requirement

range activities,

range conditions

(1962) generalized

averaged

ments for large herbivores

"normal"

the

deer fawns during the

maintenance

ME requirement

over all time periods

requirements

trials

were highest

was 154 k ca l j".
during periods

�-341-

of growth (173 kcal/kg

WO.75 / day) and lowest during the winter

WO.75 /day).

months (125 kcal/kg

ture of the maintenance

It was theorized

requirements

that the cyclic na-

and voluntary

decrease

matter

intake during winter was part of a complex adaptation

winter

survival.

in dry
for

The rmoregulation
The living mammal
constant

is cons idered to be capable of maintaining

body temperature

by adjusting

(Scholander

et al.,

1950).

production

(HP) or reduce

These are regulated
mental conditions
determine

insulation

An animal can either

and metabolism
increase

its heat loss (H-L)to maintain homeothermy.

both behaviorally

and physiologically.

and the amount of heat being produced

the extent to which heat must be conserved

(Irving and Krog,

1955).

in which metabolic

rate remains

All homeotherms
relatively

and an upper limit which,

in body temperature,

unless

The range of environmental
limit has been referred

conditions

constsnt.

There

controlled

conditions,

temperatures

is a lower

results

in a rise
response.

between this upper and lower

to as the "zone of thermal

Physiologists

range

a drop in body

for by a metabolic

nard and Loosli (1969), or the "thermoneutral
Ta r ne r (1961).

by the animal

have a thermoneutral

if exceeded,

compensated

Environ-

or dissipated

limit to this range below which the animal experiences
temperature

its heat

comfort"

by May-

zone" by King and

working with laboratory

animals

under

define "the r mone ut r al zone" as the range of

which does not cause a metabolic

res ponse to maintain

�-342-

homeothermy

during basal metabolic

considered

unrealistic

this term

in its natural

for ecologists

habitat and suggested

gical metabolic

rate"

measurements.

Moen (1973)

dealing with an animal

that an animal lives at an "ecolo-

and may spend varying amounts of time outs ide

the physiologically

defined thermoneutral

range.

Moen (1968) sugges-

ted that a "critical

thermal

exists

when the animal must

environment"

make a response

in order

duction is greater

than heat loss,

thermic

environment."

the animal

to maintain homeothermy.

hypothermic

The changing environmental

homeothermy
ments.

is in a "critical

If heat loss is greater

is in a "critical

s itate an increase

the animal

When heat prohyper-

than heat production

environment."

conditions

during winter may neces-

in HP of mule deer -Iawris in order to maintain

and as a result

Quantitative

increase

information

hypothermal

environment

white-tailed

deer in Western

maintenance

is sparse

of mule deer.
Minnesota

energy require-

relative

to the critical

Moen (1967) working with
suggested

that smaller

deer

(30 kg) would reach a negative energy balance at air temperatures
approaching
starvation

0 C and wind velocities, of
diet.

A full-fed,

± to 1 mph if they were on a

30 kg deer would reach a negative ther-

mal balance at winds le s s than 4 mph and 0 C.
diet could theoretically
wind velocities

of 8 mph.

tage to a homeotherm
ratio,

withstand exposure

which decreases

Larger

A 70 kg dee r on full

to -40 C temperatures

body size appears

to be an advan-

in cold weather due to the surface
as the animal increases

and

area to weight

in weight.

Total heat

�-343- _

production

increases

as the animal's

weight increases,

duction per unit of weight decreases

(Moen, 1968).

(1959) hypothesized

temperature

white-tailed

that the critical

adult white-tailed
wa s at 12 C.
northern
tion,

of fas te d , adult

deer in a temperature

controlled

deer in winter

metabolic

chamber

is between 5 and 20 C.

times

BMR in summer

and two times

in fall was only 9 pe r ce nt higher

(middle of "t.her morie ut r al z one ") whereas,
perature

of penned,

These data suggest that the "the r moneut r a l z one " for

white-tailed

expenditure

Holter et al.

that the minimum energy expenditure

the deer in this study were able to increase

to three

Silver et a1.

deer in winter coat may be below 0 C.

(1975) demonstrated

but heat pro-

by only 10 degrees

metabolic

In addirates

BMR in winter.

up

Energy

at -20 C than at 8 C
in spring a drop in tem-

(10 C-O C) increased

energy expenditure

by 8 percent.
These data suggest that fawns could be most vulnerable
thermia

and mortality

relatively

higher

in late winter and early spring

metabolic

weight ratio and presumably

rate,

less favorable

depleted

to hypo-

due to their

surface

area to

body fat reserves.

Blood Metabolites
During conditions

of submaintenance

catabolize

body tissues

functions.

Depending upon the tissue

result

for maintenance

in cat abo Iis m of adipose tissue

intake,

the.animal

must

of body temperature

reserves

available,

and body proteins

and vital

this may
with minimal

�•

-344-

catabolis m from the limited
long term

calorimetric

demonstrated
increased

studies

that metabolism

heat production

pose tissues

supply of carbohydrates.

of body fat accounted

proteins

more of the energy needed.

serve

available

during periods

Energy
animal

for catabolism

by gluconeogensis
tissue

processes

involve transport

possibility

of relating

balance

For wild ruminants

of inadequate

fatty acids from adipose

for most of the

energy

could be an important

of these

re-

intake.

from protein
(Annison,

more and

in winter the amount

in the severely

unde'r nour ls he d

and by mobilization

1960).

metabolites

levels of certain

has been suggested

As adi-

were found to provide

balance can be maintained

largely

Graham et a l . (1959)

of sheep exposed to cold stress.

were depleted,

of adipose tissue

with sheep,

In relatively

of

These physiological
by the blood.

blood constituents

The

to energy

by work with sheep and cattle as discussed

by Bowden (1971).
Levels
variation

of nutrient

intake have been shown to be associated

in the metabolism

of fat in the body, especially

zation of depot fat during low energy
acids (NEFA) are released

to supply the metabolic

Winkler,

1970).

ewes,

wethers

Reid and Hinks,
and cattle,

concentrations

Non-esterified

needs of the animal

has been shown to increase

and cows (Karihaloo
1962).

the mobilifatty

in blood plasma when adipose tissue

mobilized

Fasting

intakes.

with

Patterson

et al.,

(Jackson

is
and

NEFA levels

1970; Patterson,

in

1963;

(1963) concluded that in both sheep

of .1000 iJ.eq/ml or more of plasma

NEFA

�-345-

indicated

increased

mobilization

of depot fat.

reported

that mild cold (3-8 C) caused

little

Slee and Halliday

(1968)

change in NEFA in sheep

that were well fed or in those that were fasted for short

periods.

However, exposure

a marked

rise

to acute cold for 2-10 hours

in blood NEFA (up to .2000 u e q Zrn l},

with prolonged

fasting, levels

of plasma

produced

Russel

et al.

(1967) found

NEFA wouldincreareto

a

maxi-

mum of .1500- .2500 f.leq/ml.
Little

information

tional status
of baseline
animals.

is available

of wild herbivores.
"normal"

excitement

dealing with wild species.

Excitement
(Holmes

NEFA levels to the nutri-

This may be due in part to the lack

values which complicate

In addition,

mental animals

relating

blood studies

could be more pronounced

Blood sampling

and handling of the experi-

NEFA levels

and Lambourne,

in blood of both sheep and cattle

1970; Slee and Halliday,

1968).

Reid and Hinks (1962) found that plasma

NEFA decreased

peated handling and sampling,

as a result

de Calesta

found that NEFA levels

apparently

for non-starved

starvation

does (.240

However,
with re-

of reduced

(1973) working with semi-tame

(.260 u e q Zm l) were not significantly
of

when

have been shown to affect levels of NEFA in the blood.

increased

of excitement.

of these

level

mule deer,

f.leq/ml) and fawns

(P&gt; 0.1) different.

After 36 days

mean levels of NEFA for does (.610 f.leq/ml) were

Significantly

(P&lt; 0.05) higher

Presumably,

adults were able to maintain

acids longer due to their

than those for fawns (.290f.leq/ml).

proportionately

high blood levels
greater

of fatty

quantity of adipose

�-346-

tissue.

Mean levels

of NEFA for fawns starved

iJ.eq/ml) was significantly

higher

(P

33-36 days (.290

&lt; 0 .1) than that for non-starved

fawns (.260 iJ.eq/m.l).
As carbohydrate

and adipose tis sue are depleted, proteins

gluconeogensis

provide

ric homeostasis

(LeResche

more and more of the energy needed for caloet al.,

1974).

amino acids were rapidly deaminated
pose tissues

were depleted

appearance

of protein

to perform

normal

In tests

(Coulson and Hernandez,

from the cells decreased

cellular

functions

levels

in sheep (Leibholz,

tailed

deer that BUN levels we re relatively
and excitement.

1968).

(26.5 mg percent).

stable and unaffected

(1973) reported

BUN levels were relatively

(20.5 mg percent),

percent)

and we re significantly

non-starved

values

Weathe r-Severity

(42.3 mg percent)
fawns

stable up to 24 days of

after- which they increased

(34.4 mg

(P &lt; 0.05) highe r than the initial,

(17.2 mg percent).

Index

Single weather
representing

by

BUN levels of

higher than those for surviving

starvation

(BUN)

Seal et al , (1972) found with white-

deCalesta

(P&lt;0.05)

The dis-

Restricted

blood urea nitrogen

mule deer fawns dying after 33 -36 days of starvation
were significantly

rats,

the ability of the cells

and death followed.

intake has been shown to increase
1970).

with starved

and oxidized for energy as adi-

energy

handling

through

thermal

measurements
conditions

are crude estimates
of environments.

at best for

Heat exchange

�-347- .

between an animal and its environment
dients rather

is dependent on several

than on a single environmental

tempe rature

1968).

Complex models have been formulated

exchange

between an animal and its environment

ster,

1971).

Siple and Passel

based on research
a ir temperature

to describe

This index combines

is defined as the cooling power of the atmosphere
from the surface

Their formula

combinations

where H is thec;ooling
energy removed

of a human or animal.

power of the atmosphere

per square

the value 330 centigrade
wind conditions.

meter

This for-

cylinder

of

and wind speed.

in kilocalories

of

per hour; V is wind velocity

in

= ~

(330 - H/22

in degrees

is skin temperature

The value calculated

The value 22.066 reflects
minimum

(91.4°F)

still air wind-chill
of

heat

+ 10.45 - V) (330 - ta)

per second; ta is air temperature

equivalent,

Wind-chill

and indicates

of air temperature

of

is:

H = ( vt100 V

meters

index

the effects

mula is based on the rate of cooling of a small plastic
water in different

the heat

(Moen, 1968; Web-

and air movement during cold weather.

loss by convection

value (Moen,

(1945) developed a wind-chill

in Antartica.

gra-

centigrade;
under calm

for H can be converted

temperature

and

to an

by the formula:

.066) + 32

wind s peed of 4.0295 miles per hour.

wind speed is a measure

This

of the amount of heat lost from the

skin while walking at a rate of 4.0295 mph on a cold, calm day.
Petritz

(1972) suggested

that the chill- index developed by Siple and

�••
-348- .

Passel

(1945) provided a better

the wind-chill

1971).

production

of infant caribou

creased

of chill stress

on cattle than

index which was developed solely for use with cattle

(Webster,

values

measure

Data from Hart et al. (1961) suggested

of Siple and Passel.
linearly

could be predicted

that heat

from the wind chill

It was found that heat production

,with wind chill.

Results

values

reach or exceed approximately

calves

would perish

from prolonged,

in-

showed that when wind chill

1100 kcal/m

2

/hour,

continued exposure.

caribou

�-349-

CHAPTER III

METHODS AND MATERIALS

Study Location
The study was conducted
Research

Center,

two miles

principally

at the Junction

south of Kremmling,

portion

of the Radium Wildlife Management

Division

of Wildlife.

measurements

where wild deer periodically
The climate,
the general

topography,
area,

Colorado.

to winter

experience

vegetation,

Middle Park,

It is a

Area of the Colorado

This location was chosen in order

Iro m deer subjected

Butte Deer

climate

to obtain

typical of that

severe

nutritional

stress.

and winter

mortality

of deer

are discussed

by Gilbert

in

et al. (1970)

and Wallmo and Gill (1971).
The first
facility

digestibility

maintained

on the Foothills
Colorado.

were conducted at the deer

by the Department

Also all fawns were raised

mitted adjusting

of Fishery

at this facility.

chutes similar

the deer to the necessary

during the feeding experiments.

pen

and Wildlife Biology

Campus of Colorado State Univers ity , Fort

weighing and restraining

Training')

trials

Collins,

Alleyways

to those at Junction

and

Butte per-

handling (see 'Rearing

and

�-350-

Physical

Facilities

The general
of isolation

pens and associated

shown in Figure
and consisted
tion.

layout of the Iac ility is shown in Figure

2.

1.

weighing and restraining

Each isolation

Details

facilities

are

pen measured

3.04 x 9.12 meters

of woven wire sides and dirt floors

denuded of vegeta-

A 3.04 x 3.04 meter

Weighing and restraining

wooden roof partially
chutes and alleyways

covered

each pen.

from the isolation

permitted

the taking of weights and blood samples

with minimal

turbance

to the deer (Figure

chute,

deer while drawing
by Mautz et al.

2).

The restraining

blood, was constructed

similar

pens
dis-

used to hold

to that described

(1974).

Acquis ition of Fawns
In order
hand-reared

to accomplish
and trained

the objectives

of this study,

during the summers

fawns were

of 1973 and 1974.

The

fawns used in this study we re obtained from three

sources:

28 were

taken from captive.

winter

in Middle

Park

wild does trapped

the previous

and held until pa rt u r it ion ; 39 :vere obtained from semi-tame

does held at the Foothills
Collins,

Colorado;

various

areas

Office rs .

Deer Pens,

25 we re acquired

in Colorado

Colorado State University,
as orphans

and collected

by Division of Wildlife Conservation

Fort

from

�-351-

Drift fence
~~--e-

eil--e~

-Runway

;.
Isolation pen---i r-

~--~

lsolction pen
exercise area

(

Bp Holding pen

;--e---I

~~

~

-

~

~
(

B-,Metabolism
tJ cages

Loading
chute

\

./

~-6--f7~.

.••.....

[-

~

~

Work area

. j ~-~
I

~
~

P I

I

I __ Weigh

:l

building

~'

(,...0 0 ooooo~

Lob Qnd
storage
bldg

"

Holding pen

o

10

20

Scale - meters
Fi gu r e 1.

Pen facilities,
Junction Butte Deer Research
(Diagram by R. B. Gill)

Center.

�-352-

::0

c:
~:3
~

-

-

0

IT~
I

Wate~-fI
e
@

-

FOO/~

•

IX

I

0

7-

I

rl

-:

0

•

~~V

;-1

•
•

~!&gt;

•
•

"/

«" ,

"

,.........

I
'"

C.- Work areas

/~

L-

Walkway

I

II

b
\

02345
Scali - meters
Figure

2.

Details of isolation pens and handling
(Diagram by R. B. Gill)

A. - Weigh chute
B. - Restraining
chute

facilities.

�-353Rearing

and Training

Fawns were removed
Birth weight,
corded

from the dam 12-24 hours

date of birth,

sex and weekly growth rates

cedures

Fawn feeding schedules

outlined

by Reichert

fawns during the summer
Reichert

(1972).

(1972).

were re-

of 1973 was formulated

third-cutting

according

to that of
has not

fed to the fawns in 1974

to a pelleted

alfalfa hay.

fed to the

of this formula

from Robbins (1973).

fawns were given free access

of the

fed followed pro-

The milk formula

The milk composition

1) was obtained

90 days.

and amounts

The .ch e m ic a l composition

been determined.

leafy,

birth.

for each fawn from day of birth until the termination

experiment.

(Table

after

At 10 days of age,

concentrate

all

(Table 2) and

Fawns were weaned at approximately

Male fawns to be used in the study were castrated

at 5-8

weeks of age.
In order

to accurately

lites,

and digestion

accept

varying

a day inside

this,

After weaning,

maintained

each individual

that all animals

with a minimum

periods

of excitement.

fawn was bottle ied at least once

cage similar

to that of Cowan et a l , (1969)
sampling

each fawn was placed in a digestion

Acceptance
after

it was essential

and the weighing-blood

extended

seven days.

weekly weigitchanges,blocdmetabo-

of confinement

both a digestion

with modifications,

gradually

parameters,

degrees

To accomplish

measure

from 15 minutes

at first,

chute (Figure

2).

cage each day for
to a max imum of

of the weighing and blood sampling

chute was

weaning by weighing all fawns weekly in the weighing

�-354-

Table

1.

Composition

of milk formula

fed to fawns in 1974.

Nutrient

Content Percent

Water

75.65

Protein

10.14

Fat

5.83

Lactose

7.14

Ash

1.24

1

1Formulated
by adding to each can of evaporated milk 29.5 g
cas e in , 121.6 g tap water and 2.0 g of Agway Legumin mineral and
vitamin mixture.

�-355-

Table 2.

Compos ition a na ly s is of pelleted

Ingredient
Barley,

basal ration.
Percent

Weight

10.0

pulverized

Corn,

pulverized

30.0

Milo,

pulverized

5.0

Oats,

pulverized

7.5

Wheat middlings

6.5

Beet pulp,

2.5

Brewers

shredded

35.0

grain

Di c a l c iurn phosphat.e

1.0

Molas s e s , cane

2.5

Vitamin A, D, E premix

0.002 (Tr)

Trace

mineral package
(Mg, Zn, I, Fe, Co, Cu , Ca)

0.005 (Tr)
Percent

Analys is (Dry Matte r Bas is)
Dry matter

90.59

Crude

protein

19.34

Ether

extract

2.84

Ash

7.86

Cell wall constituents

52.82

Acid-detergent

13.58

fiber

3.58

Lignin
Apparent
Gross

lE.. vivo dry matter

digestibility

72.07
4.60

energy kcal/g

Apparent

digestible

Apparent

metabolizable

kcal/ g

3.45

ene rgy kcal/ g

3.05

energy

= 100.0%

�-356-

apparatus.
continued

This training

schedule

began at two weeks of age and

to the end of the experiment.

were transported
Junction

from the Foothills

Butte Deer Research

At six months of age the fawns
Deer Pens in Fort Collins to the

Center to be used in the feeding experi-

ment.

Experimental

Design--l974

Eighteen

fawns were acclimated

ment 13 days prior

to the Junction

to the initiation

which time they received,

of actual feeding trials

ad libitum,

the pelleted

they had been r e ce iving for five months prior
During this period

Butte environ-

mean dry matter

during

diet (Table 2) which

to this experiment.

intake was determined

for the

ensuing

experiment.

The 18 fawns were ranked from lightest

heaviest

and allotted

in six successive

deer within a trio were as similar
trio was random.ly selected
(treatment

A), a second

B),

the

and

to receive

random

These

measurements

groups

(dry matter

1105.18

(se

basis):

so that the weights of the

as possible.

to receive

C) percent

resulted
treatment

One member

ad libitum amounts

member

thir_d 50 (Treatment

intake.

trios

to

of each

of this feed

75 (Treatment

of the mean pre-trial

in the following treatment
A deer removed

an average

of

± 29.55) g/day but were offered 2000 g/day which was

more than they could consume
C were offered

in a 24-hour

period.

829 and 552 g/day respectively.

followed for ten weeks (January

18-March

Treatments

Band

This feeding plan was

28, 1974).

By setting

a

�-357-

constant

restricted

intake for treatments

body weight gain or loss,
of weather

in blood metabolites

data could be compared

to a known energ;

design accommodated

intake levels) with four replicate

of

and influence
int ake level.

three treatments

deer per treatment.

(energy

Differences

in

means for intake and body weight change for each week of

the 1974 trial
energy

C, relationships

changes

This experimental

treatment

Band

were analyzed

intake associated

by multivariate

with the three treatments

of digestible

and metabolizable

Regression

analysis

maintenance

maintenance

by solving for X (average

was used to estimate

y./J·75;day)w}en
Y (average

analysis.

The spread

in

allowed an estimate

energy requirements.
feed requirements

for

daily DE and ME intake/kg

daily body weight gain or loss,

kg) equals

zero.
Daily intake for each deer was calculated
nearest

gram.

As a water

source,

each deer.

Because

metabolism

cages were unavailable

complete

balance trials

energy

snow was provided

fecal collection

of the digestion

matter

digestibility

feed consumed.
chemical

and

the first year at Junction

Butte,

were conducted

at the Foothills

of the 1974 experiment.

was to determine

and digestible

quantitatively

and metabolizable

of the experimental

assess

the maintenance

Deer
The
the dry

energy value of

These values along with the ingredients

constituents

To quantitatively

trial

ad libitum to

bags were not successful,

Pens in Fort Collins at the conclusion
purpose

by weighing orts to the

and certain

feed are shown in Table 2.
ME and DE energy

�-358-

requirements
matter

of fawns in winter,

intake to digestible

(DE) and metabolizable

The DE and ME coefficients
digestion

trials

Colorado

(Table 3).

it was necessary

were determined

used in the energy
meter

dry

energy (ME) intake.

from 1:: conventional

conducted May through June 1974 in Fort Collins,

Four fawns from each treatment

x 3.04

to convert

balance trials.

metabolism

were randomly

selected

to be

Each fawn was placed in a 3.04

cage for a 10-day collection

trial.

The

deer were allowed to adjust to the cage for three days,

after which

their

Total feces

intakes

were measured

ove r a seven-day

period.

and urine were collected

and weighed once daily for seven days and

a l iquot s were preserved

for analysis.

moisture-proof

bags and frozen.

The feces were stored

After the collection

period,

in
the

feces were dried at 60 C in a forced air oven for 48 hours and then
allowed to equilibrate

at room temperature

equilibration,

the fecal pellets

I-mm

and sampled

screen

for 24 hours.

were ground in a Wiley mill through a

for analysis.

Duplicate

and feces were dried at 100 C for the correction
to 100 percent
collection.

dry matter.

Total

predetermined

day's

collection

of feed

of all determinations

The urine was acidified

we ight in grams

Aliquots

samples

with H S0 during
2 4

daily urine output for each deer was diluted to a

was then mixed', filtered
sampled.

After

by the addition of water.

through two layers

measuring

10 percent

were composited

of cheesecloth,

The urine
and

of the total volume of each

in a screw-cap

jar.

�Table

3.

Deer
Number
41

Results of 12 digestion trials conducted
in the 1974-1975 winter trials.

A
A

4
1
24

A

13
55
8
52

B
B

43

C
C
C

B
B

44
7
20

C

Mean
se
Gross energy:
Mean apparent
Mean apparent

DE3
kcaVkg

4

Colorado

5

to evaluate

6

the basal

7

r at ion used

ME8
9
kcaVkg
ME/DE
WO.: 7 / day
%
131.36
88.50
155.16
88.90
156.01
89.83
157.0089.31

DE/GE

%

%

%

u%.

%

%

74.86
75.90
77.46
75.60

76.31
77.58
78.90
77.37

148.42
174.47
173.67
175.78

3.25
3.09
2.53
2.81

23.68
22.41
21.09
22.26

5.49
5.49
5.49
5.49

67.54
68.99
70.88
69.43

69.32
65.44
77.82
71.33

71.23
68.16
79.52
73.84

152.41
136.58
158.30
230.01

2.93
4.81
2.86
2.17

28.76
31.83
20.47
26.15

5.49
5.49
5.49
5.49

62.81
57.86
71.17
66.18

134.39
115.93
141.66
156.27

88.17
84.87
89.49
89.62

68.83
64.18.
71.25
72.88

73.64
69.70
75.64
78.33

114.82
138.90
124.21
133.87

2.92
3.05
6.67
2.50

26.35
30 ..29
24.35
21.66

5.49
5.49
5.49
5.49

65.23
61.16
63.48
70.33

101.70
121. 87
104.24
120.20

88.57
87.74
83.92
89.78

72.07
1.30

75.01
1.07

155.12
8.90

3.29
.35

24.94
1.08

5.49
.00

66.25
1.22

132.98
5.91

88.22
.55

as a percent

of gross

WO .7

UE/GE

/ day

FE/GE

CH4/GE

ME/GE

4.60 kcal/g.
digestible
energy: 3.45 kcal/ g.
metabolizable
energy:
3.05 kcal/g.
6Metnane

1Digestible

dry matter.

2Digestible

energy

3Dige stible

e ne rgy intake per metabolic

4Urinary

Collins,

DDM1
Treatment
.h.

2

in Fort

energy

5 Fe ca l energy

as a percent

as a percent

as a percent

of gross

of gross
of gross

energy.
body size.

energy.

energy.

energy

7Metabolizable

energy

as a percent

8Metabolizable
size.

energy

intake

9Metabolizable
energy.

energy

as a percent

energy.

of gros s energy.

per metabolic

body

of diges tible

I
W
VI
\C

I

�-360-

Feed and feces were analyzed
ether

extract

and ash.

Cell wall constituents,

and lignin were determined
rized

by Goering

and Van Soest (1970).

The gross

energy

a manifold

style freeze

measured

acid detergent

Gross

with a Parr

energy
Adiabatic

of the urine was determined

lost in methane

crude protein,
fiber,

for the feed using the pr oce du re s summa-

feces and ur ine were determined

Energy

for dry matter,

dryer

(Vitrus

production

for white-tailed

of the feed.
Calorimeter.

after freeze-drying

Co. Inc .• Gardiner,

was estimated

in

N. Y .).

based on values

deer fawns on a similar

diet (Thompson et

a l , , 1973).

Exeerimental

Des ign- -1975

The primary
as those stated
partially

objectives
for 1974.

answered

of the winter trial
However,

restrict

energy

consumption

for survival

winter.

In order

of 1975.
1974.

treatment

energy

to adequately

was repeated

Treatment

intake to ascertain

levels

minimum

for a given period of time during
satisfy

these objectives.

during the winter

only two treatments

twins were used in the experiment

was similar

were tested

A (seven deer) were fed ad libitum,

ment B (six deer) were fed 25 percent

the entire

months (January-March)

Methodology for the 1975 winter trial

However,

(2) and (3) were only

during the winter of 1973-1974.

did not sufficiently

experiment

objectives

of 1975 were the same

in 1975.

to that of
Deer in

while those in treat-

of this amount.

Four sets of

and when weights of twins were

�-361-

similar,

one fawn from each set was as signed to a different

ment.

Fawns were randomly

In 1974, the trials
ments

and digestible
ration

at Junction

and metabolizable

were determined

The validity

is questionable.

apparent

dry matter

winter.

energy

values

results

responded

summer

of digestion

trials,

values

digestibility

to similar

requirements

Therefore,

changes

obtained

at Fort
conducted

from winter

of sheep and cattle

declined.

coefficients

of the experimental

Young and Christopherson

temperatures

maintenance

but the digestion

to early

pens.

Cl-:.,::~gy
r~quire-

Butte,

with performance

studies

efficients

maintenance

in late spring

of comparing

warm temperatures

as ambient

ass igned to one of 18 isolation

for estimating

were conducted

treat-

Collins.
in

feeding

(1974) reported
diets

decreased

if deer digestion

co-

in arribie nt temperature,

then

for mule deer fawns during the

but based upon DE and ME coeffic ients obtained

in summer,

would be in error.
To avoid this possible
constructed

source

in 1975 similar

These

cages were transported

Center

to (l) calculate

metabolizable

intake and physiological

to the Junction
coefficients

are influenced

cages were

Butte Deer Research
and the digestible

and

of the feed,

(2) measure

the effect of level of

condition

on apparent

dry matter

(ADMD), and (3) test the hypothesis
cients

two digestion

in size to those of Cowan et al. (1969).

digestion

energy

of error,

by ambient

that for deer,

temperature

digestibility

digestion

by comparing

coeffidigestion

�-362-

coeffic ient s calculated

in Fort Collins in late spring to those calcu-

lated in Middle Park during winter.
Fawns of similar
digestion
ment.

trials

age,

The same pelleted

used in the

of the 1975 experi-

ration was fed in 1974 and 1975.
were taken from the isolation

into one of two digestion

five-day

fecal collection

January

21 to March 4, 1975.

lyzed according

to those

of 1974 were used in this segment

one from each treatment,
placed

weight and training

cages.

pens and

A two-day adjustment

was conducted

to procedures

Two fawns,

and

on each of 12 fawns from

Feed and feces were handled and anadescribed

previously.

Body Weight Changes
For both the 1974 and 1975 experiments,
used as a criterion

of energy

tion of maintenance

energy

status

Average

in the body, and for the calcula-

requirements.

the beginning of the feeding trials

All fawns were weighed at

and at weekly intervals

dai.ly change in body weight over the entire

calculated

for each deer by subtracting

of a platform

weights were recorded
kilograms.

In order

scale situated
to the nearest

to minimize

was

The

des igned holding chute

beneath a wooden floor.

Body

t pound and then converted to

possible

fawns were weighed at approximately

expe riment

of days on the trial.

fawns were walked through a specially

consisting

thereafter.

the final weight from pre-trial

weight and dividing by the total number
trained

body weight change was·

error

due to gut fill,

the

the same time of day each week.

�-363

Blood Metabolites
In 1974 and 1975, blood samples
intervals

for additional

information

cedures

of handling and collecting

across

all treatment

groups.

into the weighing chute,

were taken from fawns at weekly
regarding

energy

were kept as constant

Previously

we re weighed,

trained

as pos sible

then were moved into a padded

chute des igned for taking blood samples

Ideally,

10 ml of blood were taken from the jugular

Samples

were collected

with a I-inch,

Pro-

fawns were walked

squeeze

vac ut a ine r tube containing

s t at us .

(Mautz et al.,

1974).

vein of each deer.

20-gauge

needle and 12-ml

Na EDTA or hepar in.
2

Whole blood was

kept chilled and centr ifuged within two hours after taking the first
sample.

Blood plasma

ately for later

was separated,

analysis.

pi petted and frozen

An effort was made to keep the fawns as

calm as pos sible prior to blood sampling.
each individual

immedi-

was classified

In the 1975 winter trial

each week into one of three

excitability

(not excited,

moderately

to sampling

so that the influence

excited

classes

and highly excited)

of excitability

on blood values

of
prior

might

be dete cted if it occur red.
Blood samples

taken from the fawns during the winters

and 1975 were analyzed
nitrogen.

Duplicate

for urea nitrogen
mg/lOO ml plasma
(u e q) of FFA/ml

for non-esterified

samples

of plasma

(Sigma Chemical

of 1974

fatty acids and blood urea
from trial

deer were analyzed

Company 1974) to the nearest

and for FFA to the nearest

0.1

0.01 microequivalent

(Mosinger, 1965; Dole and Meinertz,

1960).

�-364-'

Weather

Measurements

Weather

conditions

of 1974 and 1975.
recorded

were monitored

Daily temperature,

continuously

with seven-day

throughout

the study periods

humidity and wind speed were
recording

hygrothermograph

and anemometer.

Daily and weekly means were calculated

sampling

conditions

weather

over a 24~hour period.
ments relative

from these instruments

In order

to estimate

to changing environmental

require-

conditions during winter,

was made to establish

a multiple

would relate

weekly maintenance

requirements

severity.

every two hours

maintenance

an attempt

weather

by

regression

model that

to a weekly index of

The chill index developed by Siple and Passel

was used in 1974 and 1975.

Wind-chill

values were determined

mean weekly measurements

of wind and temperature.

(1945)
from

�-365-

CHAPTER

RESULTS

IV

AND DISCUSSION

Gr owth Rate s
1973-1974 Experiment
Daily growth rates
female)

during

an approximate

month post-weaning
period

were calculated

period.

for 18 fawns (9 male; 9

three-month

pre-weaning

Mean gain during the pre-weaning

was 155.08 (se ± 11.87) g/day

for males.

and 135.34 (se

11.06) g/day for females.

Mean daily gain did not differ

(P&gt; 0.05) between sexes.

Daily gain for both sexes

was 145.20 (se
sexes

± 11. 63) g/ day.

was 14.12 (se

pe riod.

fawns during this period.
Post-weaning

accelerated

during the three- month

183.41 (se ± 9.24) g/day.

(P &lt; 0 .05) greater

greater

(P&lt; 0.05)

for both sexes

than pre-weaning
increase

Male

rate than female

Daily gain for both sexes

growth rates

26 percent

showed a significant

during this pe riod

Male fawns showed a mean daily growth of

fawns gained at a significantly

proximately

significantly

± 0.48) kg.

213 .15 (se ~ 8.65) g/day and females

7.14).

±

Mean body weight at weaning for both

Body growth for both sexes
post-weaning

and three-

was 197.78 (se ~

combined
rates.

in growth after

were ap-

Both sexes
weaning.

�-366-

The mean daily growth rates
were 27 percent
mule deer,

of the pre-weaned

lower than those reported

Columbian black-tailed

over a similar

period.

1973; Robinette

et al.,

for captive

for captive dam-reared

to be approximately

1973).

dam- reared

deer and wh ite= t a i Ied deer fawns

Growth rates

deer fawns were reported

fawns in 1973

mule

213 g/day (Halford,

Cowan and Wood (1955) showed a 200

g/day growth rate for Columbian

black-tailed

deer fawns.

Robbins

and Moen (1975) found a mean daily gain of 243.7 (se ~ 19.2) g/day
by three

captive

maternal-nursed

100 days of life.

By duplicating

white-tailed
doe's

milk in energy

closely

as poss ible,

(249.7

± 5.2 g/day) for nine bottle-reared

they were able to approximate

Gastrointestinal
one factor
strated
diarrhea.

upsets

contributing

in phys iological

condition

norm of the species.
been inadequate

in 1973 was not chemically

as

this rate of growth

diarrhea

may have been

growth rates

All fawns showed varying
as suggested

that when growth rate

is likely to retard

A second factor

nutritional

meet the pre-weaning

and associated

from other species

and Hammond (1940) indicates

and protein

fawns.

to the lower pre-weaning

by the fawns in 1973.
Evidence

deer fawns for the first

of

by McMeeka n
is high a setback
to the

growth rate may have

intake.

The milk formula

analyzed.

It is possible

growth requirements

degrees

growth compared

reducing

demon-

fed to the fawns
that it did not

for mule deer fawns.

�-361--

1974-1975 Experiment
The mean birth weight of 23 fawns born between May 30 and
July 21,1974,

was 3.33 (se ~ 0.11) kg.

Average

bi r Lh weight of

male fawns was 3.50 (se ~ 0.15) kg while female fawns average
(se ~ 0.16) kg.

Birth weight did not differ significantly

3.22

(P&gt;0.05)

between sexes.
During the 1974 fawn-rearing
to more closely

simulate

fawns by adding casein
(Table 1).

operation,

the growth rates

as a protein

of dam-reared,

mule deer

pre-weaning

of 168.93 (se ± 11.65) g/day while female

± 10.90) g/day.

between sexes.

for 12 fawns (6
and three-month

Mean daily gain during the pre-weaning

was 159.25 (se ~ 8.13) g/day for both sexes.

(se

to the milk formula

were calculated

male; 6 female) during a three-month
p-eriod.

was made

supplement

Mean daily growth rates

post-weaning

an attempt

period

Males gained at a rate

fawns averaged

Rate of gain did not differ significantly

149·58
(P&gt; 0.05)

Mean body weight at weaning for both sexes was

17 . 10 (s e ± .83) kg .
In the pos t-weaning
rate of 184.26 (se
11.76) g/day.
(se

period of 1974, male fawns gained at a daily

± 10.46) g/day while females gained 187.68 (se ±

The average

weight gain for both sexes was 185.97

± 7.50) g/day and was similar

all post-weaning

to that of 1973 (197.78).

growth rate for both sexes

The over-

combined was significantly

(P &lt; 0.05) greater

than pre-weaning

growth.

to the significant

(P&lt; 0.05) increase

in the mean growth rates

This was primarily
of

due

�-368-

female

fawns relative

to male fawns during this period.

of male fawns was not significantly
pre-

and post-weaning
Pre-weaning

were greater

so better

growth rates

calculated

supplement

milk formulations

for the fawns born in 1974

(P&gt; 0.05) greater

sex or for both sexes

The casein

between the

period.

but not significantly

1973 for either

(P&gt; 0.05) different

Growth rates

combined

apparently
probably

than those in

(159.25 vs 145.20).

did not improve

growth rate,

are needed.

Food Intake and Utilization

1974 Experiment
During the 13-day acclimatization
by predators,
three

so the experimental

treatments

period four fawns were killed

design was modified to accomodate

with four deer per treatment.

age and sex of the fawns for each treatment
The 1974 experiment

began January

Initial body weights,

are shown in Table 4.
5 and terminated

March 28.

Mean weight and age of the 12 fawns at the beginning of the experiment
was 39.37

(se

Apparent

± 1.63) kg and 210.91 (se ~ 2.54

days,

in vivo dry matter

of the basal ration

72.07 (se ~ 1.30) percent,

digestibility

for metabolism

DE and ME coefficients
levels

was

while the percent DE and ME was 75.01

(se ~ 1.07) and 66.25 (se ~ 1.22) percent,
of DE available

respectively.

respectively.

was 88.22 (se ~ 0.55) percent.

did not differ significantly

of intake .. Correlation

The percent
The

(P&lt; 0.05) between

(r = .94) of dry matter

digestibility

�Table

Deer

4.

No.

1
4
41
24

Age,

sex

Mean
se

of the fawns for each treatment
Metabolic

Treatment

Age
{days ~

Sex

A
A
A
A

220
220
211
212

Male
Male
Male
Female

46.36
35.68
42.95
34.77

17.76
14.59
L6.77
14.31

39.94
2.81

15.85
.83

39.31
44.77
43.40
34.54

15.69
17.30
16.90
14.24

40.50
2.09

16.03
.62

40.45
36.13
46.13
27.95

16.03
14.73
17.70
12.15

37.~6
3.83

15.15
1. 17

215
2.46
B
B
B
B

215
200
218
191

Female
Male
Male
Male

206
6.36

Mean
se
7
20
43
44

body weights

Init ial
Body Weight
(kg)

Mean
se
13
55
8
52

and initial

C

c
C
C

219
212
207
206
211
2.97

Female
Male
Male
Female

Initial

in 1974.

Weight {kg WO. 75)
Average
for Trial1

se

17.35
14.52
16.95
14.71

.06
. 11
.21
.18

15.37
16.93
16.66
13.93

.10
.07
.08
.09

15.28
14.36
16.59
11. 78

.09
.07
.09
.07

IThe average
metabolic
weight for each deer was calculated
ir o m 10 weekly
during the trial and was used to calculate
e ne r g y intake per rne t ab ol ic body size.

measurements

taken

I

w

0\
1.0
I

�-370-

and apparent

digestible

energy

was significant

(P&lt; 0.05)

and

pos It ive .
During the winter trial,
fawns in treatment

mean daily dry matter

was 998 (se ± 22.02)

A, for 10 weeks,

Intake for this group did not differ significantly
deer or between weeks during the tr ial.
and C consumed

all feed provided

day of the trial.

Daily gross

plying daily dry matter

energy

Since the mean dietary

1.07) percent

digestible

were 3443.56

(se

digested

mean metabolizable

energy

dige s t e d kilocalories

was available

fawn over the entire

trial

88.22

kcal/day

each

in kcal/
(se ±
per day

for treatment

B

(se ± 0.55) pe r-

for metabolism
(se ± 44.38)

intake of 3037.91

kcal/ day for group C.

content,

A, 2860.43

B

by multi-

gross. energy was 75.01

Approximately

for the fawns in group A, 2523.47
1610.27

the caloric

± 51.67) for treatment

cent of the energy

between

intake was calculated

the apparently

C.

(P&gt; 0.05)

The fawns in treatments

intake by 4.60,

for treatment

g/day.

(829 and 529 g, respectively)

gm of feed.

and 1825.29

int ake for the

yielding a
kcal/day

for those in group Band

Mean daily DE and ME intake for each

is shown in Table 5.

body weight (Table 4) was used to express

Average

energy

metabol ic

intake per metabolic

body size.

1975 Experiment
Thirteen
to the Junction

fawns were transported

from Fort Collins,

Colorado,

Butte Deer Research

Center and acclimatized

to the

�Table

S.

Linear
i
equations and correlation
(or the 10-week experimcnt
of 1974.
r

eg

r

e

e

s

on

statistics

relatlnll

mea.n daily energy-intake

(x)

and

average

daily body

we

lg

ht

gain or loss

(y)

Deer
Number

DE

DE
Treatment
A
A
A
A
B
B
B
B
C
C
C
C

4
I
41
24
8
13
55
52
7
20
43
44

sc

3303.54
3329.03
3219.57
3326.36
2860.05
2860.05
2860.05
2860.05
1906.68
1906.68
19U6.68
l?C~. 68

158.81
180.45
213.37
124.90
.00
.00
.00
.00
.00
.00
.OU
.00

Intake

DE

kcal/day

DE

kcal/kg

ME

kcal/day

ME

kcal/kg

equals

lMaintenance
zero.

kcal/kg

kcal/day

WO. 75/day

wO.75/day
requirements

were

WO. 75 /day

227.51
191.87
lR~ .94
226.12
171 .67
lR6.0B
1(,8.93
20, .31
124.78
132.77
114.92
io;. .8:&gt;

kcal!day

.e

2'Jl4.38
2936.87
2R40.30
2934.51
2523.13
25l3.13
2523.13
2523.13
1682.07
1682.07
1682.07

200.70
159.56
125.~7
110.18
.00
.00
.00
.00
.00
.00
.00
.00

16H2.vl

Body Weight
Gain or Los Si
k!(/day

ME

ME
kcal/kg

WO.75/day

ZOO.;!
169.27
167.56
10.9.49
151.44
164.15
149.03
181.12
110.ns
117. i3
10l.3?
! tz , 79

1'.0210
+. 022~
~.0390
~.0569
.0000
+.0066
-.0081
- .0033
-.0406
-.0179
- .0Ri I

-.ooal

RegresBion
Eguation
~ = .000049(x)- .133392

Correlation
(r)
.82

Ma inte n an c e Requi rementl
2742.28

I
W

9 = .000R48(x)-.149659

.87

176.48

I

y: .000056(x)- .133392

. rl2

2382.00

9"

.87

153.73

calculated

.000Q6I(xl-

by solving

.1496~1

for x (average

da.ily feed intake) when y (average

daily body weight

gain or loss)

'-I
I-'

�-372-

winter

range environment

On the following 10 days,
measured.

Pre-trial

deer was 962.98

percent

January

December

21 to January

3 to 12, ad libitum

intake (dry matter

intake was

basis) of the seven ad libitum

period the seven deer in group A were fed

and the six deer in group B were offered 240 g/day (25

of pre-trial

seven weeks,

ad libitum

intake).

J'anua ry 3 to March 4.

sex of the fawns for each treatment

They received

this ration
age and

are shown in Table 6.

Mean

± 1.71) kg and 200.5 (se ± 5.49) days,
Digestion

search

trials

were conducted at the Junction

digestibility

1.50) percent

Butte Deer Re-

The apparent

of the experimental

and not significantly

was 38.00

respectively.

Cente r during the feeding experiment.

dry matter

in vivo

ration was 69.28 (se ~

(P &gt; 0.05) different

from the appa-

rent in vivo DMD of the same ration

calculated

Fort Collins.

did not differ significantly

Di ge s t ion coefficients

0.05) between levels

of intake.

tions did not appear

to impair

basal r at ion..

Dete riorating

from similar

phys iological

of its pre-trial

weight before entering

the digestion

matter

Another fawn died of apparent

day after
cent.

leaving the digestion

trials

in

(P &gt;

condi-

the ability of the fawns to digest the

One fawn had lost 25 percent

digestibility.

for

Initial body weights,

weight and age of the 12 fawns at the beginning of the trial
(se

2.

i" 22.38) g/day.

(se

During the treatment
2000 g/day,

for 13 days,

body

cage but showed a 73 percent

cage but its apparent

Body we ight loss of this fawn was 28 percent.

starvation

dry

one

DMD was 71 per-

�6. Age, sex and initial

Table

Deer

No.

34
2
18
39
38
6
17

Treatment

Age
{days)

A
A
A
A
A
A
A

186
214
221
193
168
208
205

B
B
B
B
B
B

Mean
se

during

Sex
Female
Female
Male
Male
Female
Male
Male

200
6.82

Mean
se

9
19
15
29
202
33

body weights

210
210
187
221
215
168
201
8.27

Female
Male
Male
Male
Female
Female

of the fawns for each treatment
Initial
Body Weight
{kg~

..

0.75

Me t a bo l ic Welght (kg W
Average
Initial
for Triall

37.27
44.77
42.27
33.18
29.77
42.27
40.45

15.08
17.30
16.57
13.82
12.74
16.57
16.04

38.56
2.05

15.44
.62

33.86
47.04
35.68
42.95
38.18
28.86

14.03
17.96
14.59
16.77
15.35
12.45

37.76

15.19
.80

2.6b

in 1975.

15,35
17.41
16.96
14.46
13.25
17.31
16.87

~
se

.05
.07
.13
.13
.09
.16
.17

I

w
....•
w
I

t2.81
16.36
13.07
15.46
14.28
10.99

1The average m:etabolic we ight for each deer was calculated
from eight weekly
the trial and was used to calculate
ene rgy intake pe r metabolic bod r s ize .

.45
.38
.69
.47
.26
.70

meas ur e me nt s t a.ke n

�-374-

Mean daily dry matter

intake for the deer in treatment

the entire

trial

was 987.35

(se ~ 15.46) g/day.

all of their

240 g each day of the trial.

Treatment

A for
B deer ate

Intake did not di ffe r signifi-

cantly (P &gt; 0 .05) between deer or between weeks during the trial,
one exception.

In the digestion

group A was significantly
The confinement

effects

cage,

the daily intake of all fawns in

(P &lt; 0.05) lower than in the isolation
of the digestion

in this study,

reduction

in daily intake and an increased
pens.

Most fawns in the digestion

In order

pens.

cage could not be adequately

evaluated

to the isolation

with

cage exhibited

a

rate of weight loss relative

to reduce this possible

cage effect,

the

daily intake and body weight change for each deer while in the digestion cage was not included in the weekly or total calculation

of main-

te na nce requirements.
Since the gross

energy

(4.60 kcal/gm)

percent

dige st ib le , the apparently

treatment

A and B were 3251.48

respectively.

Urinary

from the previous
calculated

energy

spring.

by Thompson

et al.

ment B.
trial

Mean

± 1.39)

per day for

(se ± 83.08) and 790.35 kcal/day,

loss was based on values

(1973).

The percent

was estimated

The mean metabolizable

(se ~ 72.26) kcal/day

kilocalories

Methane loss was estimated

energy that was metabolizable
percent.

digested

was 71.59 (se

for treatment

energy

calculated
from values

of digestible

to be 86.98 (se :t 0.28)

intake became

A and 687.45 kcal/day

2828.14
for treat-

daily DE and ME intake for each fawn over the entire

is given in Table 7.

used to expres s energy

Average

metabolic

intake per metabolic

weight (Table 6) was
body size.

�Table 7.

Llne a r regression
(y) for the 7-week

Deer
Number

Treatment

34
2
18
39
38
6
17
9
19
15
29
202
33

A
A
A
A
A
A
A
B
B·
B
B
B
B

equations and correlation
experiment
of 1974.

relating

mean daily energy

DE

DE
kcal/day

se

3052.84
3065.33
3037.82
3350.88
3218.31
3703.90
3305.00
790.35
790.35
790.35
790.35
790.35
790.35

44.92
110.95
118.57
68.50
67.27
43.49
60.50
.00
.00
.00
.00
.00
.00

Intake

statistics

kcal/kg

WO. 75 Iday

198.88
176.06
179.11
231 .73
242.89
213.97
195.90
61.69
48.30
50.47
51.12
55.34
71·91

Regres s ion Eguation

intake (x) and ave r age daily body weight gain or loss

Body Weight
Gain or Loss
kg/day

ME

ME
keall day

se

2655.36
2666.ll
26·12.29
2914.59
2799.28
3221.65
2874.1,8
687.44
687.44
687.44
687.44
687.44
687.44

39.07
96 .50
103.13
59.58
58.51
37.82
52.62
.00
.00
.00
.00
.00
.00

Correlation
( r)

kcal/kg

wO.75/day

172 .98
153.14
155.79
201.56
211. 26
186.11
170.40
53.66
42.01
52.59
44.40
48.14
62.55

+.0277
+.0230
+.0740
+.0695
+ .0649
+.1020
+.1067
- .2435
- .1993
- .4303
-.2759
- . 1391
- .3896

Maintenance

Reguirement

DE

kcal/day

~ = .000140(x)-

.389455

.92

DE

kcal/kg

WO. 75 Iday

9 = .002195(xl-

.3948~0

.89

179.89

ME

~ = .000160(x)-

.389450

.92

2434.06

ME

kcal/day
.
0.75
keal/kg
W
/day

~ = .002523(x)-

.394870

.89

1Sf .. ;0

equals

IMaintenancc
ze r o .

requirements

were calculated

by solving

{or x (average

daily feed intake) when y (average

2781.78

daily body weight gain or loss)

1

1
W
'-I
VI

.1

�-376-

Body Weight Change

Weight records

of individual deer for the 1974 and 1975 experi-

ments are shown in Appendix A and B, respectively.
culations, daily rates
number

of weight change were derived

For later

cal-

by dividing by the

of days on trial.

1974 Experiment
All fawns lost weight during the first
(Figure

3).

a mean rate

From two to ten weeks,

two weeks of the trial

groups A and B gained weight at

of 70 (se ~ .004) and 28 (se ± :006) g/day,

though one deer (#52) in group B lost slightly

respectively.

from week four to six.

Group C fawns continued to lose weight at a mean rate of 10 (se ~
.015) g/day.

with minor weekly fluctuations.

At the end of the tenth

week mean change in body weight compared

to initial weight was sig-

(P &lt; 0.01) different

nificantly

between treatments.

change for treatments

A. Band

0.58) (6.30 percent),

-0.08 kg (se ± 0.21) (-0.66

kg (se

± 1.13) (-6.20

gain by anyone
12 percent.

percent),

C after ten weeks was +2.43 kg (se ~

respectively.

fawn was 11 percent

The warmer

in preventing

C deer.

resulted

percent)

and -2.58

The maximum

weight

while maximum weight loss was

daily temperatures

may have been a' factor
No mortality

Mean weight

further

near the end of the trial
weight loss in treatment

from starvation

in 1974.

�-377-

1975 Experiment
All fawns in treatment
trial.

A gained weight during the seven-week

Mean gain was 3.25 kg (se ~ 0.60) (8.36

gain was 68 (se

± 0.01) g/day.

Tre atment B fawns exhibited
over time.
(Figure

Greatest

a linear

weight loss occurred

4) when wind chill was greatest

est estimated

pe r ce nt}, and daily

maintenance

of any week of the trial.

during the fourth week

(-23 C) resulting

in the high-

(192.65 kcal/kg

O
W .75 /day)

requirements

By the end of the seven-week

the six fawns had died of apparent
28 days and two after

decre ase in body weight

starvation.

trial

four of

Two fawns died after

42 days on the experimental

level of feeding.

The mean weight loss of the fawns that died was 11.19 (se ~ 0.48) kg
(32.19 percent),

and the average

g/day.

The two surviving

weight,

respectively.

weight for treatment

daily weight loss was 334 (se

fawns lost 16.03 and 13.73 percent

Figure

4 depicts

an apparent

increase

± 0.04)
body

in body

B during the fourth and sixth week of the trial.

This is due to a reduction

in s ample size as a re sult of the death of

two fawns during each of these weeks and not an actual weight gain.
The means for the fifth and seventh week are based on the mean weight
loss of the three

and one surviving

fawns.

Maintenance

Requirements

Feed requirements
for x (average

for maintenance

respectively.

were calculated

by solving

daily feed intake) when y (ave rage daily body weight

�-378-

8
6

-

4

.s:::.

0\
Q)

2

~

&gt;.
"'0

0
.0

0

C
Q)

0\

c
.s:::.
0

-2

&lt;.)

-4
-6
-8

2
Figure

3.

3

456
Weeks

7

9

8

Mean weekly weight change (percent) compared
initial weight of fawns on three different levels
dietary energy,
1974 experiment.

to
of

10

�-379-

10

5

-

s:

0\

Ad Libitum

-5

Q)

~

&gt;.

'"'0

0
..c
c -10
(1)

0\

c
0
s:

(,)

-15

-20
I)

B.
.25xAd Libitum

-25

(3)

2

3

4

5

6

Weeks
F'i gu re 4.

Mean weekly weight change 'pel'cent)
compared
to
initial weight of fawns on two different
levels of
dietary energy.
1975 expr- r irn ent,

7

�-380gain or loss) equals zero.

The regress

maintenance

(Table 5. Table 7) were based on average

requirements

daily DE and ME intake (kcal/day

ion equations

for estimating

O
W .75 /uay) and the

and kcal/kg

corres ponding daily weight change of the fawns over the entire
ment.

The DE and ME intake at which the resulting

zero (energy equilibrium)

is the apparent

experi-

curve crosses

maintenance

requirement.

1974 Experiment
The DE intake required
timated

to maintain
75

to be 176.48 kcal/kg

requirement

WO.

was 155.73 kcal/kg

energy equilibrium

/day while the ME maintenance
75

WO.

/day (Figure

tions of daily DE and ME intake with average
highly significant

(r

was es-

5).

The correla-

daily weight change were

= .87; P &lt; 0.01).

1975 Expe riment
The DE and ME requirements
seven-week

experiment

kg WO.75 /day.

were estimated

respectively.

cant (r = .89; P&lt; 0.01).
gain or loss on average
Figure

for maintenance

to be 179.88 and 156.50 kcal/

T he correlations

Regression

for the entire

of average

daily metabolizable

were highly signifidaily body weight

energy

intake is shown in

6.
Regression

equations

and ME maintenance

estimated

for each of the first

adequate

sample

calculation

requirements

six weeks of the trial

(Table 8).

size due to the death of four fawns prevented

of maintenance

requirements

were
Inthe

for the seventh week of the

�-381-

.OS
.06

- .04
0'

~

Cf)
Cf)

0

.02

1\

Y=.000961(x}-.I49661
r=.S7
se ± .005
Maintenance:
75
155.73 keal/kg WO.

•
•

~

0

c:
.-

-

0
0'

0

•

s:

0'
(1)

•

-.02

~
~
"0

0
(D

-.04

-06

-os
100

50

150

200

250

ME intake (kcal/ kg W 0.75 )
Figure

5.

Regre ssion of average daily body weight gain or 108.8
on average daily metabolizable
energy intake,
1974
experiment.

�-382-

.30
A

.20

- .10
0'

~

U&gt;
U&gt;

Y=.002523 (X) -.394870
r=.89
se±.02
Maintenance:
75
156.50 kcall kg WO.

0

'0
c:
.-

•

•

0

0
0'

-

-'=0' -.10

•
•

Q)

3:

~-.20
0
(I)

-.30
-.40

•
50

100

150

200

250

ME intake (kcal/kg W 0.75)
Figure

6.

Regression
on average
experiment.

of average daily body weight gain or loss
daily metabolizable
energy intake,
1975

�Table

8.

Linear
r e gr e s s ions, c o r r e lations,
first six weeks of the 1975 winter

ME maintenance
trial.l

requirements

arid mean

wind- chill

for the

Equation

C o r r e 1a t LOn
(r)

n

se

Maintenance
Requirement
0 75
kcal/kg
W .

.002134(x)-

.240609

·91

6

.005

112.75

-12

2

.002307(x)-

.360668

.90

5

.004

156.33

-18

3

.002611(x)-

.409340

.74

5

.016

156.77

-19

4

.002176(x)-.419214

.61

5

.049

192.65

-23

5

. 002494(x)-.

428089

.83

3

.016

171. 64

-17

6

.002745(x)-

.460853

.76

3

.030

167.88

-19

'

Week
Number

Regression

1

1The regression
equation
and maintenance
requirement
the trial due to an inadequate
sample
size in treatment
B.
died of apparent
starvation
I:.y the end of the seventh week.

Mean
Wind-Chill

°c

were not calculated
for the seventh
week of
Four of the six fawns in this treatment
had

I
W
00
W
I

�-384-

trial.

The s lopes

did not differ
quirements

of the equations

relatively

When weekly
wind-chill

values

wind speeds
(y) increased
35.9875
size

constant

maintenance
calculated

(Table

with wind-chill
(r

z;

suggests

were

= 5.57)

requirements
Y =

to the equation

(Figure

7).

is calculated

The small

precludes

the extra-

of this experiment.

coefficient

and small

hypothesis

However,

standard

for investigating

conditions

and energy

Maintenance

Requirements

sample

equilibrium

error

of

the reof mule

in winter.

Expe riments:

The overall
equilibrium

DE and ME intake

(maintenance)

The DE maintenance
179.89

were

155.73

the same

used

ration

and 156.50.

(P&gt; 0.05)

similar

required

for body weight

for both winter

experiments.

for 1974 and 1975 were

respectively,

for both winter

under

levels

similar

requirements

not significantly
were

were

75
WO. /day,

kcal/kg

requirements

weight

and

a workable

of environmental

Combined

were

trial.

from the mean daily temperatures

data beyond the scope

this function

re-

with

·93; se

high correlation

lationship

six weeks

compared

(x) according

the relatively

deer

requirements

(n = 6) on which this equation
of these

for the entire

9) it was found that maintenance

- 6.871(x)

polation

for the first

(P&gt; 0. 05) ind ic at ing that maintenance

significantly
were

calculated

and

while the ME maintenance
The slopes

of these

equations

Fawns

of similar

age and

different.
experiments

conditions.

176.48

and fed approximately
The refore,

a regress

ion

�Table

9.

Date

Maximum,
mlnionum
and mean dally te mpe r at u re , humidity,
were derived from readings
at Z-hour intervals.

wind speed

and a •• oclated

Tem~erature

Humidit~

!%)

Week
Number

(oC)

wind-chill

index

{or the

period

Wind Spc e d
(meters
[?:er so!l()nd~

of January

14 to March

Mean

1,1975.

Wind-Chilli
Still Air
Equivalent

Maximum

Minimum

Daily
Mean

0

-20

- 9

100

2S

62

15

0

2.7

1015

-12

1/14-1/20

Maximum

Daily
Mean

Minimum

Max i rnu rn

Mi ni mu m

Dally
Mean

kcal/mZ/hr

1/20-1/27

2

+1

-30

-10

100

30

68

20

2.5

4.5

1167

-18

1/27-2/3

3

-5

-26

-14

100

50

76

30

2.5

3.2

I In

-19

2/3-2/9

4

-4

-28

-17

96

:'l

75

30

2.5

3.2

1249

- 23

-1

-23

-10

98

26

60

30

2

3.6

1110

-17

2/9-2/16

Means

°c

I
W

ex&gt;
VI

2/16-2/23

6

-8

·-30

-18

100

42

78

30

0

2.2

1177

- 1q

2/23-3/1

7

+2

-27

-10

100

40

75

30

1

3.6

1110

-17

7 Week Ave rage

~

-'!.L

Temperat

ure

-13

.588

Relative

Humidity

73

.883

3.6

.369

Wind Velocity

(oC)
(%)

(mps)

Wind-Chill:
2
kcal/m /hr
Still

lSiple

and Passel,

1945.

At r Equivalent

1187

0c

-20

I

�-386-

-

200

&gt;.

0
"0

10'-

,....

0

3:

i75

Ol

~
&lt;,
0

o
~

-

150

en
c:
C1&gt;

E

•...

125

C1&gt;

;:,
0C1&gt;

/\

•...

C1&gt;

u
c:

y= 35.9875- 6.871{x)
r=.93

100

se± 5.57

0

c:
C1&gt;

c:

0

75

E

w
~

-30

Figure

-25

7.

-20

-15
-10
-5
Mean weekly wind -chill index (Oc)

Effect of wind -chill index on metabolizable
ene rgy
maintenance
r c qui r e rne nts , 1975 experiment.

o

�-387-

equation

based

on mean

corresponding
.was used

body weight

to estimate

maintenance
k ca l / kg W
these

0.75/

(Silver

68 percent

These

higher

for animals

both winters

The e s t i rnat e d

experiments

was

157.69

of maintenance

for

metabolism

value

deer

in a the r mo nc ut r a l

fawns

that for s he e p and cattle

of 90 kcal/

a roughage

eating

diet (2.

a concentrate

ales

°

ration

that ME maintenance

of energy

in a the rmo-

of us ing ME for maintenance

consuming

consuming
loss

and

1969).

the efficiency

data suggest

due to the greater

ME kcal/ g m)
(3.0

ME kcal/

requirements

could

s m et a bo l i z a ble ration,

as heat during

was

be

presumably

the digestion

of a high

diet.
The estimated

maintenance

exper irrie nt was 157.69
metabolizable

kcal/kg

concentrate

ration

efficiency

values

suggested

consuming

native

forage

mately

the fasting

for white-tailed

for animals

gm).

fiber

The estimate

reported

for animals

and 75 percent

8).

et al.,

Blaxte r (1962)

requirements.

combined

times

reported

environment

for the 25 fawns from

from

day (Figure

0.75
W
/day)

(ME kcal/kg

ME maintenance

fawns was 1.75

environment

intake

change

requirements

0.75
kg W
/day

neutral

daily

(2.

maintenance

wO. 75 /day.

°

requirement

for the fawns

WO..75 / day and was based
(3.

°

by Blaxter

ME kcal/ gm).
(1962)

with a metabolizable

are

value

ME kcal/ gm) (Gill and Wallmo,

1973),

requirement

to 173.92

would be increased

on a highly

Assuming

applicable

energy

in this

the

to deer
of approxi-

the equivalent
kcal/kg

ME

�-388-

.40

.30

_ .20
•..

0\

~

'-'
(f)
(f)

0

.10

to...

0

c
.0

0'1

-

1\

Y =.OO2292(x)-.361527
r=.87
se± .015
Maintenance;
75
157.69 kcal/kg WO.

Q

0
Et

.0

.c:.

•

0'1

.~ -.10
&gt;.

0

0

•

"0

LD -.20

-.30

-.40

50

100

150

ME intake (kcal/kg

Figure

8.

200

250

WO.

75

)

Regression of average daily body weight gain or loss
on average daily metabolizable energy intake, 1974
and 1975 experiments combined.

�-389-

However,
low their
tation
part

for animals

critical

exposed

temperature

present

situation

energy

balance

body temperature

(NE) for maintenance.

ME equals

fermen-

Accordingly,

in the

needed

to maintain

does not increase.

The blood energy

Metabolites

metabolites

or no measurable

for the fawns in this

relationship

Mean weekly blood urea

nitrogen

to energy

study exhi-

status.

(BUN) concentrations

from

each treatment

for each week of the 1974 and 1975 experiment

shown in Table

10 and Table

of BUN varied

greatly

response

to energy

the mean pre-trial
1.93)

mg percent

treatments.

In 1974, weekly

In 1975

for all fawns was 32.83

a mean BUN concentration
they were

levels

and showed no apparent

and was not significantly

(P&gt; 0.05)

of 75.48

different

starvation

(se

!

between
in treat-

(se ~ 4.10) mg

sampled

(1-3 days before

death),

(P&lt; 0.01) higher

than the means

for the fawns

A and the survivors

Non-esterified

are

or body weight change over time.

BUN concentration

which was significantly

treatment

within f:reatments

intake

the last time

in treatment

11, res pe ct ive ly .

The four fawns that died of apparent

ment B showed
percent

be-

ar.d thus become

NE and the ME intake

Blood Energy

bited little

conditions

the heat loss due to microbial

would be used to maintain
of the net energy

to environmental

free

in treatment

B.

fatty acid (NEFA) concentrations

for each week of the 1974 and 1975 experiment

for each
are given in

�-39010. Means

Table

trations
energy

Week
Number

Mean

0
1
2
3
4
5
6
7
8
9
10

46.00
55.00
25.25
38.16
39.33
30.40
23.00
37.10
32.66
44·90
22.37

1

mg urea

Table

and standard
error
of blood urea nitrogen concen1 for fawns on three different levels of dietary
for each week of the 1974 experiment.

A (high)
se

2.79
3.75
1.75
7.67
6.30
3.27
2.01
4.97
4.67
1.65
2.37

nitrogen

n

Treatment
B (medium~
Mean
se

n

Mean

4
4
2
3
3
3
3
3
3
3
3

50.25
40.83
24.83
32.83
44.58
26.20
34.75
41.12
36.66
37.25
22.90

2
3
3
3
4
4
4
4
3
4
4

30.25
34.50
22.00
38.50
45.00
25.60
27.50
32.62
41.83
35.87
30.25

5.26
9.68
2.91
2.66
7.10
2.47
1.25
4.26
1.91
2.49
1.75

C (low)
se

n

12.28
1.32
1.04
9.00
3.50
2.08
2.65
2.96
11.16
5.24
4.75

2
3
3
4
4
4
4
4
3
4
2

per 100 .ml plas ma .

11. Means

and standard
error
of blood urea nitrogen concentrations I for fawns on two different
levels of dietary
energy for each week of the 1975 experiment.
Treatment

Week
Number

Mean

0
1
2
3
4
5
6
7

33.62
34.73
34.73
29.84
39.82
37.08
35.77
34.36

1

mg urea

per

A {high~
se

2.99
2.75
3.03
2.35
3.05
2.88
1.86
4.41

100 ml plasma.

n

Mean

7
3
7
5
5
7
6
3

31.90
29.12
32.70
37.58
56.30
39.52
53.66
20.66

B (low)
se

2.57
3.45
3.58
11.09
7.69
2.97
10.93

n

6
5
6
5
6
4
4
1

�-391-

Table

12 and Table

centration

13, respectively.

(P &gt; 0.05)

was little

variation

from this

The mean pre-trial
(se

i" 0 .03)

different

f.l. eq/ml for

between

of apparent

the fawns in treatment
addition,
bility

lower

different

between

treatments.

NEF A concentration

for 1975 was

.448

all fawns and was not significantly
The NEFA levels

A or the surviving

fawns in treatment

B.

into three

of excita-

of individuals
excited

strained

mule deer.

sistently

sample

Because

all animals

an adequate

in 1975 compared

these

sample

Environmental

wind-chill

minimum

variability

these

treatment

9 and Table

humidity

and wind speed

and mean daily temperature,

14, respectively.
were

in

from

re-

to condata cannot

responses.

Conditions

humidity

for each week of the 1974 and 1975 experiments

in Table

The

to 1974 are

size and the inability

of the potential

In

pr ior to

blood metabolites

each week of the trial,
test

classes

from

blood parameters.

but may be due to the inherent

colle cting and analyzing

different

and highly excited)

to the sampled

NEFA concentration

Maximum,

of the fawns that died

(P&gt; 0.05)

handling,

be considered

(P&gt; 0 .05)

were not significantly

mode rately

to interpret

There
pe riod.

showed no relationship

pre-trial

difficult

± 0.004) f.l. eq/ 1 and

the sampling

the classification

(not excited,

handling

.978 (se

mean throughout

treatments.

starvation

NEFA con-

was

for the 1974 experiment

was not significantly

The mean pre-trial

derived

Means
from

are given

for temperature,

readings

and

at two-hour

�-392-

Table

12.

Week
Number
1

2
3
4
5
6
7
8
9
10

micro-equivalents

Table

Treatment
B (medium}
Mean
se
.978
.007
.915
.083
.935
.006
.003
.989
.450
.034
.973
.003
.001
.991
.021
.898
.955
.007
.001
.978
.011
.973

A (high}
Mean
. 978
.861
.954
.987
.522
.980
.997
·937
.969
.976
.986

0

1

Means and standard
error
of plasma free fatty acid concentrations 1 from fawns on three different
levels of dietary
energy,
for each week of the 1974 winter trial.

13.

SE:

.005
.007
.006
.003
.047
.003
.001
.012
.001
.004
.002

n
4
3
2
4
4
3
4
4
2
3
4

of free

fatty

acids

per

n
2
2
3
3
4
4
4
4
2
4
4

Mean
.978
.828
·947
·990
.436
.970
.996
.881
.971
·927
.993

C (low)
se
.008
.025
.003
.001
.020
.004
.001
.027
.009
.022
.003

n.
2
2
3
3
4
2
3
4
2
4
2

ml plas ma .

Means

and standard
error
of plasma free fatty acid concenfawns on two different
levels of dietary
energy.
for each week of the 1975 winter trial.

t r at lons+ from

Treatment
Week
Number

Mean

A (high}
se

n

Mean

0
1
2
3
4
5
6
7

.400
.455
.470
.610
.623
.52;4
.579
.651

.02
.04
.02
.03
.02
.03
.03
.04

7
3
6
5
5
7
6
3

.503
.451
.611
.598
.557
.516
.551
.562

1

micro-equivalents

of free

fatty

acid

B (low)
se

pe r ml pIa s ma .

.05
.09
.10
.05
.02
.05
.05

n
6
5
6
5
6
4
3
1

�Table

14.

Maximum,
minimum
Means were derived

and mean daily temperature,
humidity,
from readings
at Z...hour intervals.

Temperatu~c
Date

Week
Number

1/24-1/31

2

1/31-2/7
2/7

-2/14

4

2/14-2/21

(DC)

Humidity

and a •• ociated

wind-chill

(~,)

{me

index

tc

for the period

Wind Speed
r a per s e c orid)

of January

8 to March

Mean

18,

Wind-Chilli
Still Air
Equ iva le nt °c

Minimum

Maximum

Minimum

Mean

Max i rnu m

+ 2

-29

- 9

100

39

77

is

3.2

IOS6

- 15

+ I

-25

-12

88

24

62

20

1.8

992

-12

+ 1

-20

- 9

94

24

65

30

4.0

1110

-17

- 7

-27

-18

96

48

76

IU

1.3

1056

-IS

- 7

-24

- IS

80

SO

63

·9

892

- 7

2.2

846

- 5

Daily
Mrrn mu m

z

Daily
Mean

2

kcal/m /hr

2/21-2/28

6

+ 5

-12

- 2

92

20

35

IS

2/28-3/7

0'

+ 6

-11

- 3

100

40

70

40

3

S.4

1009

-12

3/7

8

+11

- 7

+ I

100

40

67

20

2

302

798

- 3

3/14-3/21

9

+ 9

- II

-

I

100

36

59

20

4. S

848

- S

3/21-3/28

10

+ 7

- 4

+ I

78

36

56

20

4.9

885

- 7

-3/14

10 Week

Average

Temperature
Relative
Wind

Humidity

Velocity

~

(DC)
(%)

(mps)

kcal/ m2 /hr
Still Air Equivalent
and Passel,

19°45.

=c

3

__

s~

-s

.G28

70

1. 46

3.6

1. 40

Wind-Chill:

ISiplc

1974.

Daily
Mean

Maximum

1/18-1/24

wind speed

1058
-i5

I
W
1.0
W
I

�-394-

intervals.

Wind-chill

measurements

winters.
but seldom

determined

for the 1974 experiment

in 1975 from

was - 8 C compared
tive humidity

were

from

mean weekly

ranged

from

of wind and temperature.

Temperatures
C while those

values

-30 to +2 C.

to a mean temperature

Mean temperature

for 1974

of -13 C for 1975.

ranged

between

24 and 100 percent

Wind speed

varied

from

exceeded

4.5 mps.

-15 and -20 C, res pectively.

-29 to +11

and was similar

0 to 13.5 meters

Mean wind-chill

Rela-

per second

both
{mps}

for 1974 and 1975 was

�-395-

CHAPTER

V

SUMMAR Y AND CONCLUSIONS

Two winter
Colorado

feeding

in an attempt

trials

to quantify

fawns held in pens exposed
Feed
fawns

intake

in these

value

average

an energy

fawn in these

maintaining

mined

from

ambient

gained
level

change

intake

32 percent

1-2 days before

measurements

of mule deer

environment.
indicated

that the

157 ME kcal/kg

during

winter.

of energy

WOo 75 /

Assuming

requirements,

this

the

2423 kcal/day

for

balance.
of the experimental
trials

was linearly
All deer

body we ight.

ration

was apparently

related

as deter-

not affected

Four

by

de ath was significantly
B.

intake

levels

conseeither

of the six fawns on the lowest

starvation

Mean level

in treatment

to feed (and,

on ad libitum

died of apparent

body weight.

and the survivors

Park.

level of intake or phys i.olo gic a l condition.

ene rgy) intake.

of energy

mately

digestibility

or maintained

winter

(39 kg) would require

24 in vivo digestion

Body weight
quently,

estimate

in Middle

demands

approximately

energy

tempe rature,

the energy

equilibrium

trials

a constant

The apparent

deer

required

to be an acceptable

conducted

to the prevailing

and body weight

trials

day to maintain

were

after

losing

of BUN for these

highe r than those
Except

approxifawns

of control

for BUN concentrations

�-396-

collected
in this

prior

to death,

study exhibited

However,
tently

because

the blood energy

no measurable

of small

sample

metabolites

relationship
size

all animals

each week of the trial,

be considered

an adequate

test

measurements

ments.

In 1974, no relationship

energy

requirements

quirements
creased

were

linearly

the small

sample

working

hypothesis

of the potential
were

similar

between

was established.

to energy

and the inability

sample

Weather

for the fawns
status.

to consis-

these

data cannot

treatment

responses.

for both winterexperiweather

conditions

and

In 1975, ME maintenance

correlated

with (r = 0.93)

wind-chill

as wind-chill

temperature

became

colder.

this

function

suggests

more

controlled

size used to establish
to be tested

under

re-

index and inHowever,
only a

experimental

conditions.
When comparing
with those
tors

calculated

the maintenance
from

mus t be cons ide red.

calorimetry

minimize

contribute

significantly

requirements.
the present

indirect
First.

animal

trial

respiration
values

activity

to the overall

The relatively

large

allow considerable

have been a significant

portion

Blaxter

that activity

(1962) reported

and in a thermoneutral

total

expenditure

from these

calorimetry

calculated

from

measurement
size

of maintenance

maintenance

environment

by 11 to 15 percent.

pens used

of movement

by sheep

two fac-

it does not

of the isolation

freedom

trials

ind ir e ct

to the point where

of the total

aric e intake
energy

requirement

and cattle

in

and may

require

ment .

at ma int eri-

would increase

their

�-397-

A second
neutrality.
based

factor

affecting

The energy

on controlled

in metabolic

requirements

ambient

rate.

with cold tolerance

Holter

et al..

jected

to a "critical

As a result.

a significant

factor

are usually

that do not cause

an increase

measurements

environment"

the increased

in increas

deer

to which the results

of these

rate

can be extrapo-

are probably

range

animals

and digestion

variables

such as the effects

tween an animal
logical

of forage

the energy

to adverse
investigation

requirements

weather

in maintenance

exchange

suggest

to adequately

in winter.

to
with

Unmeasured

and poss ible adaptive
conditions

derived

associated

on the energy

is required

of mule deer

Values

when applied

of low digestibility.

and its environment,

mechanisms

deal of additional

underestimates

of "cover"

could also be

trials

penned

activity

sub-

ments .

from

of the increase

were

require

mule deer fawns is speculative.

because

1959;

for much of the winter

lated to free-ranging
animals

(Silver.

trials

metabolic

ing total energy

1974 to 1975 to-

from

that the fawns in these

hypothermal

is thermo-

indoors

data from white-tailed

1975) suggest

experiment.

requirements

of animals

conditions

The weather

gether

The extent

maintenance

be-

phys iothat a great

determine

�-398-

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1945.
Measurements
of dry atmospheric cooling in subfreezing
ternpe r at ur e s . Proc.
Amer.
Phil. Soc. 89 :177.

Slee,

J. and R. Halliday.
1968. Some effects of cold exposure,
nutrition and experimental
handling on serum free fatty acid levels
in sheep.
Anim. Prod.
10:67-76.

m

1959.
Basal metaboJ. Wildl. Manage.

and H. H. Hayes.
deer.
J. Wildl.

1969.
Manage.

Thompson,
C. B., J. B. Holter,
H. H. Hayes, H. Siber and W. E.
Urban, Jr.
1973.
Nutrition of white-tailed
deer.
1. Energy
requirements
of fawns.
J. Wildl. Manage.
37 (3):301-311.
Ullrey,

D. E., H. E. Johnson,
W. G. Youatt, L. D. Fay. B. L.
Schoepke and W. T. Magee. "1970.
Digestible
energy requirements for winter maintenance
of Michigan white-tailed
does.
J. Wildl. Manage.
33(3):482-490.

Wallmo,
O. C. and R. B. Gill.
1971. Snow, winter distribution
and
population dynamics
of mule deer in the Central Rocky Moun··
t a in s , Snow and Ice in Relation
to Wildlife and Recreation.
Symp. Proc.
1-15.
Wallmo,
O. C. and R. B. Gill.
1973. Middle Park Deer Study-physical characteristics
and food habits.
Colo. Di v . Game,
Fish and Parks.
Game Res. Div. Fed. Aid Proj.
W-38-R- 27.
Game Res. Rept.
July. Part 2. pp. 81-103.

�-403-

APPENDIX

�Appendix

Weight r e c o r d s (kg) of individual
1974 experiment.

A.

Treatment
Deer

deer

and mean

55

5l

43.40

39.31

44.54

34.54

.48

41.59

36.13

44.31

33.86

-1.47

- I. 51

.42

42.27

37.04

42.Q;

32.

rz

-1. 28

.65

41.59

47.50

42.95

.80

.40

41. 81

37.84

.74

.75

42.27

.26

1.05

36.93

.50

45.45

38.06

44.65

47.04

45.68

Change

[or e a ch

--

_. _______
i

along

..vit h their

e r r o r te r m s

v

-

TE':~{!'1e~
In,d

40.45

3 h. 13

-16.13

.65

40.00

35.66

43.86

27.04

- 1.02

.43

- I .70

.23

37.95

35. !S

·13.18

25.yO

2.04

.49

33.40

-I. 58

.15

j7.95

35.Zl

42.72

26.36

-2.10

.54

43.86

33.75

- I . 13

.23

37.50

35.56

42.15

27.84

-I. 90

.Q:'

38.86

42.04

33. II!

- I. 36

.42

37.72

34. ~3

41. 81

27.38

- 2.33

. i9

41. 81

38.75

44.31

31.59

- I. 33

.61

36. IJ

34.54

41.36

25.9C

- 3. 18

.79

.64

~2.B.j

37.~5

oil. s s

33'.52

-,t.13

.22

3i .84

34. oj

011.5"

26.3b

- 2. ~,

.70

.96

.93

43 '.29

38.29

43.52

33.40

.82

.23

36.81

34.06

41.13

26.13

- 3.13

.i4

38.06

1.61

.62

44.31

39.43

43.06

34.88

.02

.51

37.61

33.52

41 .25

26.81

-2.86

.76

48.29

38.75

2.43

.58

43.40

39.77

43.97

34.31

.08

.21

37.61

34.88

40.45

27.38

- 2. 58

1.13

in Body

Weight.

46.72

34.77

35.22

42.95

44.54

33.40

-1.00

2

33.63

42.04

44.31

34.09

3

33.86

42.50

43.86

34.77

4

34.65

42.27

45.00

35.00

35.56

41.81

44.09

35.68

6

35.45

41.93

44.31

37.38

7

35.79

43.63

45.79

8

36.36

44.09

9

36.81

10

37.15

sc

sc

treatment

~3

42.95

Number

we-i g ht

20

35.68

Week

BWI

13

24

Weight:

r e d to initial

i;

1

Initial

c o mpa

Tr c at rnc nt B
BWI

41

r :

cha nge s in body wc l g ht

A

4

Numbe

weekly

4~

--

sc

27.95

I

.j::ooo

lBW

~ Mean

Weekly

-

0
.j::ooo
I

-

�Appe nd ix B.
Weight
record.
1975 experiment.

(kg) of individual

dcer

and mean

Treat

mem A
38

weekly

changes

in body weight

compared

to i n it i a l we ight for each

treatment

along

with their

error

terms-I

'Deer

Nurnbe r :

Initial

Weight:

Week

Nu~

34
37.27

2

6

33.18

29.77

6
42.27

BWI

17

40.45

19

15

29

mo nt B
202

33

33.86

47.04

35.68

42.95

38.18

28.86

33.86

45.45

33.86

42.04

37.27

40.90

Treat

-----

BW

~('

27.95

-I .02

.26

34.09

31.36

43.18

41.59

1.41

38.40

44.54a

43.18

34.54

31.

,q

41. O~

42.50

1.52

.18

31.81

4J.40

32.50a

35.90

26.13

-Z.55

.30

45.00

43.40

l.HI

.45

30.90

42.04

28.86

39.5-1

35.00

Zl.81a

- 4.27

.73

.6,

29.31

41.36

23.63

3 a.: Sa

34.5·!

17.95

-7.36

1.72

3!1. 86

b

35.22

32.95

b

-7.04

.91

J

., ..• 'J •.•
I._I

h

-' ...•

.5.~ . Z t

b

-9.24

l. , 1

37.27

b

31.36 a

b

-9.77

44.77

44.09

3l.13a

35.45

.29

38.IS

44.54

42.27a.

35.68

31.36

38.ISa

44.77

43.86

36.13

30 90

45.~0

44.09

Z .15

.6'0

27.04a

44.54

36.59

3 I . 5~

45.00a

41. 31

2.08

.0;7

23.63

3.25

.71

b

Weekly

45.45
45·90

Change

45.90

in Body

a

36.59

32.95

45·90

n.27

44.31

45.68

2.04

We ig ht ,

a
Confined

9

se

42.72

38.63

I BW = Mean

42.27

39

46.59

37.72

7

44.77

18

38.63

38.40
4

2

to d i g e s t io n cage ..-weight

bDied of apo a r e nt starvation.

change

not included

in calculation

of mean.

a

b

't..'

I

~

0
VI
I

��-407July, 1976
JOB PROGRESS

COLOR..\!)O

State of
Project

REPORT

"-------

No.

Deer-Elk

W-38-R-31
----

Investigations

14
Job No.
--- 10._-------------Middle Park Cooperative Deer Study Junction Butte Wildlife Habitat Improvement Project

Work Plan No.
Job Title
Period

Covered:

Personnel:

July 1, 1975 - June 30, 1976

Wayne 1. Regelin,

O. C. Wa1lmo,

L. H. Carpenter,

and D. L. Baker.

ABSTRACT
Results of snow n~nipu1ation studies regarding their effects on deer use and
vegetation responses are being summarized in a dissertation which will become
the final report for most of the objectives of this job. This dissertation
will be on file at the Colorado Division of Wildlife Research Center Library
and copies will be forWarded to appropriate Fish and Wildlife Service Federal
Aid personnel.
The experimental design was completed for the winter range seeding experiment and the various treatments and replications were planted.
Three methods
of planting seed are being tested on each of 5 different species of vegetation.
Planting methods are drilling, broadcasting, broadcasting and harrowing.
Plant species tested are crested wheatgrass, yellow sweetclover, Ladak alfalfa,
arrow1eaf ba1samroot, and Palmer's beardtongue.

��-409MIDDLE PARK COOPERATIVE DEER STUDY
JUNCTION BUTTE WILDLIFE HABITAT IMPROVEMENT PROJECT
Wayne L. Regelin

P. S. OBJECTIVE
In Middle Park. Colorado and other similar mule deer winter ranges in
mountainous sagebrush habitat native forage does not produce sufficient
supplies of chemical protein and energy to maintain deer in the demanding
winter months. The objective of this study is to develop revegetation
procedures to establish stands of species with high contents of digestible
protein and chemical energy to supplement native vegetation in meeting the
nutritional requirements of mule deer in winter.

SEGMENT OBJECTIVES
1.

Measure snow depth and density and configuration of snowdrifts in relation
to presence or absence of snowfences and in relation to shrub stands
protected and unprotected from the accumulation of snowdrifts.

2.

Measure occupation by deer of shrub stands protected and unprotected from
the accumulation of snowdrifts.

3.

Measure the relative use of browse by deer in shrub stands protected and
unprotected from snowdrifts.

4.

Measure the response of native vegetation (composition and density) in
relation to increased or decreased snow depth.

5.

Evaluate the success of forage species planted on sites of created snowdrifts and on control areas not covered with drifts.

6.

Measure soil mo Iature with relation to increased or decreased snow
accumulation.

7.

Extend the plantings of the most successful forage species on sites where
they were most productive in preliminary trials.

METHODS AND MATERIALS
The methods used to meet all segment objectives are reported by Regelin (1974,
1975) .

RESULTS AND DISCUSSION
Snow Drift Manipulation
A dissertation (Regelin 1976) is being prepared based on information obtained
from this study and is near completion. It will be on file at the Colorado

�-410-

Division of Wildlife Research Center Library and will become the final
report for Segment Objectives 1 through 6.

Seeding Deer Winter Ranges
Four blocks of land on Junction Butte with different slopes and aspects but
similar vegetation composition and density were selected as study sites during
2
June, 1975. Each block was divided into 4 plots, each 15 m.
Three treatments
and a control were randomly assigned to plots within each block. The layout
of each study site is shown in Figure 1.
Pre-treatment data on plant cover and herbage yield were collected during
July. The herbage yield in each plot was estimated using the Neal Electronics
herbage meter. However, correlation coefficients between herbage weight and
meter readings were very low due to rocky conditions and steep slopes. In
future years herbage yield will be estimated from clipped quadrants only.
The variance obtained from the clipped quadrants used to calculate the sample
size necessary to estimate biomass within 10 percent of the mean with a 95
percent confidence level. An n of 38 clipped quadrants per treatment plot was
calculated, so 50 quadrants per plot will be clipped in future years.
Plots were seede~ during September, 1975 with a seed mixture containing 5
species. These species and percent mixture by weight were: Agropyron desertorum
60 percent, Melilotus officinalis 15 percent, Medicago sativa 15 percent,
Balsamorhiza sagittata 5 percent, and Penstemon palmeri 5 percent. The broadcast and broadcast plus harrow treatments were seeded at a rate of 33 kg/ha and
the drill treatment at a rate of 11.2 kg/ha. In the drill treatment rows were
planted 30 cm apart with a cone seeder. Rows containing the grass and balsamroot were alternated with rows containing alfalfa, yellow sweetclover and
penstemon.

LITERATURE CITED

Regelin, W. L. 1974. Middle Park Cooperative Deer Study - Junction Butte
Habitat Improvement Project. Colo. Div. Wildl. Fed. Aid W-38-R-28,
Work Plan 14, Job 10, Job Prog. Rep., Game Res. Rep. July, Part 2.
p. 227-292. (proc.)
Regelin, W. P. 1975. Middle Park Cooperative Deer Study.- Junction Butte
Habitat Improvement Project. Colo. Div. Wildl. Fed. Aid W-38-R-29,
Work Plan 14, Job 10, Job Prog. Rep., Game Res. Rep. July, Part 2.
p , 265-294. (proc.)

�-411-

15 m
Drill

} 3 m

Oontro1

Broadcast

Broadcast
and
Harrow

Fig. 1.

Layout of site 1

�-412-

Control

Broadcast

Broadcast
and
Harrow

Drill

Fig. 1 cont.

Layout of site 2

�-413-

Broadcast

Control

Drill

Broadcast
and
Harrow

Fig. 1 cont.

Layout of site 3

�-414-

Broadcast
and
Harrow

Control

Broadcast

Drill

Fig. 1. cont.

Layout of site 4

�-415JOB PROGRESS

State of

.

July, 1976
REPORT

..::C.::.O=L..:..ORAJ-=:)..:::.O
__

Project No.

W-38-R-31

Deer-Elk

Work Plan No.

16

Job No.

Job Title

Piceance Deer Study - Population

Investigations
1

---------------------------

Distribution

Period Covered: April 1, 1975 through March 31, 1976
Personnel:

R. M. Bartmann,

J. J. Klein, Jr., and R. Hancock

ABSTRACT
One hundred sixty-seven deer were neckbanded and/or eartagged from December
3, 1975 through February 13, 1976. Eleven other deer were eartagged incidental to other trapping activities in Area 4 from February 15-25, 1976.
Poor trapping success and a late winter thaw caused an early ending to the
year's trapping even though about 70 deer remained to be banded in Area 13.
The composition of the total catch was 42 male fawns , 39 female fawns, 17
mature males and 80 mature females. This brings the total deer marked in
the White River drainage the past five winters to 1,923. Two hundred
thirteen sightings and 55 recoveries of banded deer were received during
the past year. The 1975-76 winter marks the end of trapping and banding
for this study, although two more years of data collection are scheduled.

��-417-

PICEANCE DEER STUDY - POPULATION DISTRIBUTION
Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.

SEGMENT OBJECTIVES
1.

To delineate deer sub-population boundaries and concentration areas on
the Piceance winter range.
a.

Define deer sub-population boundaries.

METHODS AND MATERIALS
See Bartmann (1972).

RESULTS AND DISCUSSION

Deer Trapping
Deer trapping and banding began December 3, 1975 and terminated February 13,
1976. The
winter
was miL~and
open...__ and a 4-wheel-drive pickup was used the
---------..._._'.... ::~-:.,-:-.-----~--·--~-_~_w~_~
entire winter. -Trapping success was extremely poor all winter and banded
deer quotas were reached in only two of the three areas trapped. Work in
'~r_ea_}3
__(lIammon~Draw - Gillam Draw) was curtailed after only 28 deer had been
marked over a 36-day period. Melting snow and muddy roads ruled out the
possibility of catchLng another 70 deer in this area to reach the original
quota. Eleven deer were eartagged in Area 4 incidental to other deer trapping
activities. The 178 deer neckbanded and/or eartagged included 42 male fawns,
39 female fawns, 17 mature males and 80 mature females (Table 1). This brings
the total number of deer marked during the past five winters in the White
River drainage to 1,923.
Two hundred thirteen sightings and 55 recoveries of banded deer were received
during the past year (Tables 2 and 3). A considerable number of sightings
were unusable this year because observers confused neckband colors. Unfortunately, this will probably be an even greater problem in the future because of the different neckband colors used in new areas.
The 1975-76 winter marks the end of deer trapping and banding for this study.
Banded deer sightings and recoveries will be recorded through the winter of
1977-78. This will allow two years of data collection for deer banded this
past winter.

�-418-

LITERATURE CITED
Bartmann, R. M. 1972. Piceance deer study - population distribution.
P. 315-337. In Game Research Report. Colo. Div. of Wildlife,
Denver. 3(Part 3):253-377.

).~
.

'\-.~,

/
I

/"

~~)

.

I

Prepa red by _-'-_---,·,L/-'·IC-·-'-·--'..::~/~· -4'\'~:'.i...~'-L·~V"':':"~V'-·:"'" ~'r',-,' .::cc'r':....~/
R. M. Bartmann
Wildlife Researcher

_

�Table 1. Record of deer trapped and marked on the w Lnt er range in Game }fanagement Units 11, 22, and 23,
winter 1975-1976.
Date
(1976)

Sex

Age

Eartag
No.

Area 4 - Green W/White
2-15
2-15
2-18
2-19
2-21
2-21
2-22
2-23
2-23
2-25
2-25

Female
Male
Male
Female
Female
Female
Male
Male
Male
Male
Male

Fawn
Fawn
Fawn
Fawn
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Fawn

Neckband
No.

Stripe Neckband

L-3275
L-3270
L-3272
L-3273
L-3276
L-3277
L-3278
L-3279
L-3280
L-3282
L-3283

Location
Twp. Range

(All Deer Eartagged
IN
IS
IS
IN
IN
IS
IN
IS
IS
IN
IS

96W
96W
96W
96W
96W
96W
96W
96W
96W
96W
96W

Remarks

Sec.

Only)

21
3

11
28
34
3
21
3
11
21
3

I

~

I-'

\0

I

Summary:
Date
(1975)
12-6
12-6
12-7
12-7
12-7
.12-7
12-7
12-8
12-8
12-8
12-8
12-8
12-8
12-8
12-8

5 Male Fawns; 4 Female Fawns; 2 Mature Males
Area 10 - Yellow Neckband

Male
Female
Female
Female
Male
Female
Female
Female
Female
Female
Male
Female
Female
Male
Female

Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Mature
Mature

L-3l02
L-3l03
L-3l04
L-3l05
L-3l06
L-3l07
L-3l08
L-3l09
L-3ll0
L-311l
L-3112
L-3113
L-3114
L-3115
L-3116

67
25
24
22
69
10
23
16
15
17
66
19
20
68
28

IS
1S
IS
1S
1S
IS
IS
1S
1S
IS
IS
IS
IS
IS
IS

92W
92W
92W
92W
92W
92W
92W
92W
92W
92W
92W
92W
92W
92W
93W

21
21
14
15
15
21
21
14
14
14
15
21
21
23
2

Only one eartag.

3" antler in velvet, left.

�Table 1. Record
winter 1975-1976

0 f

deer trapped and marked on the winter
(continued).

Date
(1975)

Sex

Age

Area 10 - Yellow Neckband
12-8
" 12-8
12-9
12-9
12-10
12-10
12-10
12-10
12-10
12-10
12-10
12-10
12-11
12-11
12-11
12-12
12-12
12-12
12-12
12-12
12-12
12-12
12-13
12-13
12-15
12-15
12-16
12-16
12-17
12-17
12-17
12-18

Female
Male
Female
Female
Female
Female
Male
Male
Female
Female
Female
Female
Male
Female
Female
Male
Male
Female
Male
Female
Male
Female
Female
Male
Female
Female
Female
Male
Female
Female
Female
Male

Eartag
No.

range in Game Management

Ne ckb and

No.

Twp.

18
65
14
11
21
12
59
64
27
30
29
73
36
26
31
71
40
74
44
75
43
76
77
38
78
79
80
37
82
83
86
41

1N
1N
1S
1S
1S
1S
1S
1S
1N
1N
1N
1N
1S
1N
1N
1S
1N
1N
1N
1N
1N
1N
1N
1N
1S
1S
1N
1S
1N
1S
1S
1S

Location
Range Sec.

Units 11, 22, and 23,

Remarks

(Continued)

Fawn
Fawn
Mature
Fawn
Mature
Fawn
Mature
Fawn
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Fawn
Fawn
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn

L-3117
L-3118
L-3119
L-3120
L-3121
L-3122
L-3123
L-3124
L-3125
L-3126
L-3127
L-3128
L-3129
L-3130
L-3131
L-3132
L-3133
L-3134
L-3135
L-3136
L-3137
L-3138
L-3139
L-3140
L-3141
L-3142
L-3143
L-3144
L-3145
L-3146
L-3147
L-3148

92W
92~v
92W
93W
92W
92W
92W
93W
92W
92W
92W
92W
93W
92W
92W
92W
92W
92W
92W
92W
92W
92W
92W
92W
92W
93W
92W
93W
92W
93W
93W
93W

26
26
23
2
14
21
23
2
26
26
26
26
2
34
26
21
34
34
34
33
26
26
34
33
23
2
34
2
33
2
2
2

-------------------------------------------------------------------------------------------------------------

I
.po.
N
0
I

�Table 1. Record of deer trapped and marked on the winter
(continued).
winter 1975-1976
Date
(1975)

Sex

Area 10 - Yellow Neckband
12-18
• 12-18
12-19
12-19
12-19
12-19
12-19
12-20
12-20
12-20
12-20
12-21
12-21
12-21
12-21
12-21
12-21
12-22
12-22
12-22
12-22
.12-22
12-22
12-23
12-23
12-23
12-23
12-29
12-29
12-29
12-30
12-30

Female
Female
Female
Male
Female
Female
Female
Female
Female
Male
Male
Male
Male
Male
Female
Female
Male
Male
Female
Female
Female
Male
Female
Female
Female
Female
Female
Male
Female
Female
Female
Female

Eartag
No.

Age

range in Game Management

Neckband
No.

Twp.

93
84
85
46
92
88
87
90
94
42
45
52
53
51
103
104
49
47
95
96
97
50
106
100
102
101
105
70
48
107
56
55

1S
1S
1N
1S
1S
1S
1S
1N
IN
1S
1S
1N
1N
1S
1S
1S
1N
1N
1N
1N
IS
1S
1S
1N
IN
1S
1S
1S
1S
IS
1N
1N

Location
Range Sec.

Units 11, 22, and 23,

Remarks

(Continued)

Mature
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature

L-3149
L-3150
L-3151
L-3152
L-3153
L-3154
L-3155
L-3156
L-3157
L-3158
L-3159
L-3160
L-3161
L-3162
L-3163
L-3164
L-3165
L-3166
L-3167
L-3168
L-3169
L-3170
L-3171
L-3172
L-3173
L-3174
L-3175
L-3176
L-3177
L-3178
L-3179
L-3180

93W
92W
93W
93W
93W
92W
92W
92W
93W
93W
92W
93W
93W
92W
92W
92W
92W
92W
92W
93W
92W
92W
92W
92W
93W
92W
92W
93W
92W
92W
92W
92W

2
23
34
2
2
23
23
33
34
2
23
34
34
14
22
23
34
34
33
34
23
14
15
34
34
22
22
2
21
23
33
33

One eartag.
May have injured neck.

-------------------------------------------------------------------------------------------------------------

I

"'"

N
•.....

I

�Table 1. Record 0 f deer trapped and marked on the winter
winter 1975-1976. (continued).
Date
(1975)

Sex

Area 10 - Yellow Neckband
12-30
" 12-30
12-30
12-30
12-31
12-31

Female
Female
Female
Fema1e_
Female
Male
Summary:

Eartag
No.

Age

Neckband
No.

range in Game Management

Location
Twp. Range

Sec.

92W
93W
92W
92W
92W
92W

33
34
22
22
15
22

Units 11, 22, and 23,

Remarks

(ContinuedL

Fawn
Mature
Mature
Fawn
Fawn
Fawn

L-3181
L-3182
L-3183
L-3184
L-3185
L-3186

54
57
72
58
63
62

IN
IN
IS
IS
IS
IS

21 Male Fawns; 20 Female Fawns; 5 Mature Males; 39 Mature Females
I

Date
(1976)
1-4
1-5
1-5
1-5
1-6
1-6
1-7
1-7
1-7
1-7
1-7
1-8
1-8
1-9
1-9
1-9
1-11
1-11
1-20
1-20

~

Area 12 - White with Red StriEe Neckband

N
N

I

Female
Male
Female
Female
Female
Female
Male
Female
Female
Female
Female
Male
Female
Female
Male
Female
Male
Female
Female
Female

Fawn
Fawn
Mature
Mature
Fawn
Mature
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Mature
Fawn

L-3187
L-3188
L-3189
L-3190
L-3191
L-3192
L-3193
L-3194
L-3195
L-3196
L-3197
L-3198
L-3199
L-3200
L-3201
L-3202
L-3206
L-3207
L-3209
L-3210

39
69
40
41
42
43
71
146
147
46
148
65
149
150
70
151
64
87
152
77

3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N

98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W

34
34
27
27
34
14
35
35
35
35
34
35
27
27
35
27
35
35
35
27

Only one eartag.

-------------------------------------------------------------------------------------------------------------

�Table 1. Record 0 f deer trapped and marked on the w Lnt e r range in Game Management
winter 1975-1976 (continued).
Date
(1976)

Sex

Age

Eartag
No.

Area 12 - White with Red StriEe Neckband
1-21
1-21
1-21
1-21
1-22
1-22
1-23
1-23
1-24
1-24
1-24
1-24
1-24
1-24
1-24
1-25
1-25
1-26
1-26
1-26
.1-26
1-27
1-27
1-27
1-27
1-29
1-30
1-30
1-30
1-31
2-1

Female
Female
Male
Male
Female
Female
Female
Male
Female
Female
Female
Male
Male
Female
Male
Male
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Female
Male
Female
Male
Male

Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Mature
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn

L-3211
L-3212
L-3213
L-3214
L-3215
L-3216
L-3218
L-3219
L-3221
L-3222
L-3224
L-3223
L-3225
L-3226
L-3227
L-3230
L-3231
L-3233
L-3234
L-3135
L-3236
L-3139
L-3240
L-3241
L-3242
L-3246
L-3248
L-3249
L-3250
L-3253
L-3256

Neckband
No.

Location
Twp. Range

Sec.

3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N
3N

35
34
34
27
36
27
36
35
35
35
35
35
27
27
27
35
34
35
35
27
27
35
34
27
35
35
35
34
34
35
27

Units 11, 22, and 23,

Remarks

(Continued)
153
154
99
68
156
155
45
67
83
74
79
103
101
76
102
95
73
85
84
72
75
98
78
92
88
44
80
96
82
97
104

98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98\.[
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W
98W

I
.p..
N
l.V

May have hurt neck.

-------------------------------------------------------------------------------------------------------------

I

�fable 1. Record of deer trapped and marked on the winter
(continued).
winter 1975-1976
Date
(1976)

Sex

Age

Eartag
No.

Area 12 - White with Red Stripe Neckband
2-1
~ 2-2
2-4

Male
Female
Female
Summary:

Fawn
Mature
Mature

L-3257
L-3258
L-3260

Neckband
No.

range in Game Management

Location
1';.:p.Range

Sec.

3N
3N
3N

27
35
35

Units 11, 22, and 23,

Remarks

(Continued)
100
90
93

98W
98W
98W

10 Male Fawns; 9 Female Fawns; 6 Mature Males; 29 Mature Females

Area 13 - Green and Yellow Block Neckband
1-9
1-10
1-10
1-11
1-22
1-23
1-24
1-24
1-25
1-26
1-26
1-28
·1-28
1-28
1-30
1-30
1-31
1-31
2-2
2-5
2-5
2-7
2-7

Female
Female
Female
Female
Male
Male
Male
Female
Male
Male
Female
Male
Male
Female
Male
Female
Female
Male
Female
Female
Female
Female
Female

Mature
Mature
Mature
Mature
Fawn
Fawn
Fawn
Fawn
Mature
Mature
Fawn
Mature
Mature
Mature
Fawn
Mature
Mature
Fawn
Fawn
Mature
Mature
Fawn
Mature

L-3203
L-3204
L-3205
L-3208
L-3217
L-3220
L-3229
L-3228
L-3232
L-3237
L-3238
L-3243
L-3244
L-3245
L-3251
L-3252
L-3254
L-3255
L-3259
L-3261
L-3262
L-3263
L-3264

1
7
9
3
2
55
53
11
None
51
10
57
69
5
59
12
14
65
8
15
17
16
13

2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N
2N

lOOW
lOOW
lOOW
lOOW
lOOW
100W
lOOW
lOOW
100W
lOOW
lOOW
100W
lOOW
lOOW
lOOW
lOOW
lOOW
lOOW
lOOW
100W
lOOW
lOOW
lOOW

22
15
23
17
15
15
17
17
22
17
22
17
22
22
17
17
15
23
22
18
18
31
31

I
.pN
.pI

One eartag.

-------------------------------------------------------------------------------------------------------------

�Table 1. Record of deer trapped and marked on the winter
winter 1975-1976 (continued).
Date
(1976)

Sex

Age

Eartag
No.

Neckband
No.

range in Game Management

Twp.

Range

Sec.

2N
2N
2N
2N
2N

lOOW
100W
lOOW
lOOW
100W

17
17
30
30
30

Units 11, 22, and 23,

Remarks

Area 13 - Green and Yellow Block Neckband
2-8
2-8
2:-9
2-9
2-9

Female
Female
Male
Female
Female
Summary:

GRAND SUMMARY:

Fawn
Mature
Fawn
Fawn
Mature

L-3265
L-3266
L-3267
L-3268
L-3269

(Continued)
24
25
67
27
28

6 Male Fawns; 6 Female Fawns; 4 Mature Males; 12 Mature Females

42 Male Fawns; 39 Female Fawns; 17 Mature Males; 80 Mature Females

I

.&amp;:'N

VI
I

Corrections to Previous Years' TraEEing
(Corrections are Underlined)

Records

12-31-72 Male
1-8-73
Female

Mature
Mature

L-18l7
L-1863

46
32

3S
3S

97W
96W

9
16

1-8-73

Female

Mature

L-1777

15

3S

97W

2

3-5-73

Female

Mature

L-2313

25

IS
96W
15
Yellow wired stripe neckband add - neckband mal be no. 27

12-19-73 Male

Mature

L-2232

25

IS
96W
16
Yellow wired stripe neckband
add - neckband mal be no. 27

Male

Mature

L-1787

59

1-9-74

3S

96W

16

Pink neckband.
Pink neckband - removed note
about guestionable eartag no.
Pink neckband - removed note
about guestionable eartag no.

Pink neckband.

-

�-426Table 2. Sightings of deer marked on the winter range in Game Management
Units 11, 22, and 23, received from March 1975 through February 1976.
Date

Neckband

(1974 &amp; 1975) Color

5-20-74
5-27-74
5-27-74
7-15-74
10-20-74
11-17-74
12-3-74
12-3-74
12-3-74
2-16-75
2-16
2-16
2-16
2-24
2-24
3-29
4-6
4-9
4-9
4-9
4-12
4-12
4-13
4-13
4-13
4-13
4-13
4-14
4-14
4-15
4-15
4-15
4-15
4-16
4-16
.4-16
4-16
4-16
4-16
4-16
4-16
4-16
4-16
4-16
4-16
4-16
4-16
4-16

Blue
Red wlwhite stripe
Red wlwhite stripe
Blue
Red wlwhite stripe
Red wlwhite stripe
Orange &amp; white blocks
Orange &amp; white blocks
Orange &amp; white blocks
Blue &amp; yellow blocks
Blue &amp; yellow blocks
Blue &amp; yellow blocks
Yellow
Blue &amp; ye Ll.ow blocks
Yellow
Yellow
Yellow wired stripe
Pink
Pink
Pink
Red &amp; blue blocks
Red &amp; blue blocks
White
White
White
White
Blue
White
White
Red &amp; blue blocks
White
Blue &amp; white
Yellow wired. stripe
Yellow wired stripe
Yellow wired stripe
Yellow wired stripe
Yellow wired stripe
Yellow wired stripe
Yellm,T wired stripe
Yellow wired stripe
Yellow wired stripe
Yellow wired stripe
Green w/white stripe
Green w Iwh ite stripe
White
Green w/whit.e stripe
Green wlwhite stripe
Green wlwhite stripe

Number

t
?

92
t

55
55
150
t
?

69
92
74
39
1
3
5
82
66
73
37
51
59
t
?

2
14
120
?
t

48
?

26
70
82
38
23
52
62
100
71
74
118
237
250
?

33
93
9

Twp.

Location
Range

Sec.

IN
2S
2S
2N
2S
2S
IS
IS
IS
IN
IN
IN
IS
IN
IS
IS
IS
3S
3S
3S
IN
IN
2S
2N
2N
2N
2N
IS
IS
2N
2S
IS
IS
IS
IS
IS
IN
IN
IN
IN
IN
IN
IN
IN
2N
IN
IN
IN

92W
94W
94W
88W
94W
94W
93W
93W
93W
93W
93W
93W
93W
93W
92W
92W
96W
96W
96W
96W
95W
95W
98W
98W
98W
98W
98W
98W
98W
96W
98W
97W
97W
96W
96W
96W
96W
96W
96W
97W.
97W
97W
97W
97W
97W
94W
93W
93W

17
23
23
29
14
14
36
36
36
19
20
20
24
29
22
19
10
5
5
1
32
32
6
4
4
4
4
15
15
28
19
28
21
10
13
13
32
32
32
36
36
36
36
36
34
24
17
11

------------------------------------------------------------------------------------

�-427-

Tabie 2. Sightings of deer marked on the winter range in Game Management
Units 11, 22, and 23, received from March 1975 through February 1976.(continued).
Date
(1975)
4-18
4-18
4-18
4-18
4-19
4-19
4-19
4-19
4-19
4-19
4-19
4-19
4-20
4-20
4-20
4-20
4-20
4-20
4-20
4-20 _
4-21
4-22
4-23
4-23
4-24
4-24
4-24
4-24
4-24
4-24
4-24
4-24
4-24
4-24
4-24
4-24
"4-27
4-27
4-27
4-29
4-29
4-29
4-29
4-29
4-29
4-29
4-29
4-29
5-1
5-1

Neckband
Color

Blue &amp; yellow blocks
Blue &amp; yellow blocks
Red &amp; blue blocks
Red &amp; blue blocks
Yellow wired stripe
Blue &amp; white
Yellow wired stripe
Blue &amp; white
Blue &amp; white
Blue &amp; white
Blue &amp; white
Blue &amp; white
White wired stripe
White wired stripe
Blue &amp; red blocks
White wired stripe
White wired stripe
White wired stripe
White wired stripe
White
Blue &amp; yellow blocks
Blue &amp; red blocks
Yellow wired stripe
Yellow wired stripe
Blue &amp; white
Green wlwhite stripe
Blue &amp; yellow blocks
Red &amp; blue' blocks
Red &amp; blue blocks
Red &amp; blue blocks
Red &amp; blue blocks
Red &amp; blue blocks
Red &amp; blue blocks
Red &amp; blue blocks
Yeliow wired stripe
Yellow wired stripe
Green wlwhite stripe
Blue &amp; white
Blue &amp; white
White
White
Green w/white stripe
Green wlwhite stripe
Blue &amp; white
Blue &amp; white
Blue &amp; white
Blue &amp; white
Yellow wired stripe
White
White

Number

Twp.

45
47
25

78

IN
IN
2N
IN
IS
IS
IS
IS
IS
IS
IS
IS
3N
3N
2N
2N
2N
2N
2N
2N
IN
IN
IS
IS
2S
IS
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IN
IS
IS
2S
2S
IN
IN
IS
IS
IS
IS
IS

?
?

2S

?

1
?

109
131
22
3
42
68
143
35
190
123
3
38
145
?
?

68
39
32
?

107
72
10
38
23
88
35
2
37
2
4
194
22
131
223
?

63
254
90
31
?
?

2S

Location
Range

Sec.

94W
94W
95W
95W
96W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
97W
93W
96W
96W
96W
96W
94W
94W
94W
94W
94W
94W
94W
94W
94W
94W
94W
97W
97W
97W
98W
98W
96W
97W
97W
97W
97W
97W
97W
98W
99W

30
30
1
23
5
11
2
11
11
11
11
11
36
36
2
2
2
18
18
20
18
15
10
10
5
5
3
18
18
18
18
18
18
18
18
18
1
11
11
10
10
31
36
21
21 .
28
28
21
19
24

----------------------~-----------------------------------------------------------

�-428-

Table 2. Sightings of deer marked on the winter range
in Game Management
Units ll~ 22, and 23~ received from March 1975 through February 1976 (continued).
Date
(1975)
5-2
5-12
5-12
6-26
7-?
7-?
7-2
7-9
7-18
8-?
8-?
8-?
8-1
8-4
8-21
8-21
8-26
8-29
8-30
9-?
9-2
9-21
9-22
9-23
9-26
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
"10-?
10-9
10-12
10-13
10-13
10-13
10-14
10-16
10-17
10-25
10-25
10-25
10-25
10-25

Neckband
Color

Blue &amp; yellow blocks
Yellow wIred stripe
Yellow wIred stripe
Orange &amp; white blocks
Blue &amp; red blocks
Blue &amp; red blocks
White
Blue &amp; yellow blocks
Blue &amp; red blocks
White
Blue &amp;,red blocks
Orange &amp; white blocks
Orange &amp; white blocks
Blue &amp; red blocks
Orange &amp; white blocks
Orange &amp; white blocks
White wIred stripe
Red &amp; blue blocks
Yellow
Yellow
White
Blue
Orange &amp; white blocks
Orange &amp; white blocks
Red &amp; blue blocks
Blue
White wIred stripe
White wIred stripe
Orange &amp; white blocks
Green wlwhite stripe
Blue
White
White
Red &amp; blue blocks
Green wlwhite stripe
Green wlwhite stripe
Orange &amp; white blocks
Blue
Green wlwhite stripe
White
Blue &amp; white
White wIred stripe
Green wlwhite stripe
Red wlwhite stripe
White wIred stripe
Blue &amp; white
Red &amp; blue blocks
Blue &amp; yellow blocks
Red wlwhite stripe
White

Number

Twp.

Location
Range

Sec.

10
?
27
?
200
?
?
?
?
?
?
57
14
8
123
"50
119
32
4
?
"?
?
15
50
?
?
?
?
12
?
?
?
?
?
?
162
8
?
?
?
?
?
?
71
?
?
?
83
?
?

IN
2S
IS
IS
IN
IN
3N
2N
2N
IS
IS
IN
IS
IN
IS
IS
4N
3N
IS
IS
IN
IN
IS
IS
3N
4S
3N
3N
IN
2S
IS
IS
2N
IN
IS
IN
IS
IN
2N
IS
IN
IN
IN
2S
2N
IS
IN
IN
3S
2S

93W
96W
96W
89W
92W
92W
93W
94W
92W
91W
93W
90W
94W
94W
94W
94w
95W
95W
93W
92W
89W
93W
94W
94W
95W
96W
93W
93W
93W
9lW
92W
92W
96W
95W
95W
96W
94W
90W
90W
99W
97W
97W
9lW
93W
92W
93W
9lW
92W
93W
94W

18
2
4
17
26
26
3
17
16
13
24
14
35
29
35
35
18
16
14
30
20
25
35
35
16
36
26
26
32
5
9
19
31
29
5
15
34
21
23
30
36
1
20
22
9
14
1
26
28
13

------------------------------------------------------------------------------------

�-429-

Table 2. Sightings of deer marked on the winter range
in Game Management
Units 11_ 22_ and 23_ received from March 1975 through
February 1976 (continued).
Date
(1975-76)

10-26
10-26
10-27
10-28
10-30
11-?
11-?
11-?
11-?
11-?
11-9
11-12
11-22
11-22
11-22
11-26
11-26
12-10
12-8
12-12
12-13
12-13
12-13
12-13
12-13
12-13
12-13
12-15
12-15
12-15
12-15
12-15
12-15
12-15
12-16
12-16
12-16
12-16
12-16
12-16
12-16
12-16
12-16
1-3-76
1-4
1-6
1-15
1-15

Neckband
Color

Blue
Blue &amp; yellow blocks
Red &amp; blue blocks
Red &amp; blue blocks
Blue
Green w/white stripe
Yellow wired stripe
Green w/white stripe
Green w/white stripe
White
Blue &amp; 'yellow blocks
White wired stripe
Orange &amp; white blocks
Orange &amp; white blocks
Orange &amp; white blocks
Green w/white stripe
Green w/white stripe
Blue &amp; yellow blocks
Yellow
Red w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
White
White
White wired ·stripe
White wired stripe
Blue
Green w/white stripe
Blue
Yellow wired stripe
Blue &amp; white
Yellow wired stripe
Yellow wired stripe
Yellow wired stripe
Pink
Green w/white stripe
Red w/white stripe
Red w/white stripe
Red w/white stripe
Green w/white stripe
Orange &amp; white blocks
Blue &amp; yellcw blocks
Blue &amp; yellow blocks
Yellow
Yellow

Number

Twp.

Location
Range

Sec.

?
?
200
?
?
?
?
?
?
?
12
62
50
42
12
?
?
11
3
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
68
?
44
77

3N
IN
IN
3N
IN
IS
IN
IS
IS
IS
IN
IN
IS
IS
IS
IN
IN
IN
IS
3S
IS
IN
IN
IN
IN
2N
2N
IN
IN
2N
IS
IS
IS
2S
IS
IS
IS
2S
IS
2S
2S
2S
2S
IS
2N
IN
IS
IS

95W
92W
92W
92W
92W
95W
94W
95W
95W
95W
94W
96W
94W
94W
94W
95W
95W
93W
92W
96W
95W
96W
96W
96W
96w
98W
98W
99W
99W
98W
95W
95W
96W
95W
96W
96W
96W
97W
94W
94W
94W
95W
94W
94W
93W
93W
96W
96W

27
26
25
24
25
19
7
1
1
1
'22
20
1
35
35
36
36
29
15
36
6
16
16
7
7
24
14
31
31
6
23
35
34
14
15
15
28
32
6
31
31
31
6
15
32 .
17
12
12

----------------------------------------~-----------------------------------------

�-430-

Table 2. Sightings of deer marked on the winter range in Game Management
Units 11, 22, and 23, received from }1arch 1975 through February 1976. (continued).
Date
(1976)

Color

1-20
1-26
1-27
1-27
1-27
1-27
1-27
1-27
1-29
1-31
2-?
2-?
2-14
2-17
2-17
2-19
2-23

Pink
Blue
Blue &amp; yellow
Yellow
Yellow
Yellow
Yellow
Blue &amp; yellow
Yellow
Green w/white
Yellow
Yellow
Blue &amp; yellow
Pink
White
Green wiwhite
Green w/white

Neckband

blocks

blocks
stripe

blocks

stripe
stripe

Number

Twp.

Location
Range

Sec.

66
2
103
37
53
71
8
?
83
?
76
84
?
15
155
33
9

3S
2N
IN
IS
IS
IS
IS
IN
IS
IN
IN
IS
IN
3S
IN
IS
IN

96W
lOOW
93W
93W
93W
93W
92W
93W
93W
95W
94W
92W
94W
97W
97W
96W
96W

5
23
18
3
2
2
23
35
3
31
5
23
29
2
29
3
21

�-431-

Table 3. Recoveries of deer marked on the winter range in Game Management
Units 11, 22 and 23, March 1975 through February 1976.
Date of
Recovery
(1975)
3-?
3-20
4-?
4-4
4-21
4-23
5-?
5-?
5-13
6-17
6-18
9-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-?
10-4
10-22
10-24
10-25
10-25
10-25
10-26
10-26
10-27
10-28
10-28
10-31
11-?
ll-?
ll-?
11-?
ll-?
11-2
ll-2
11-4

Neckband
Color
Yellow
Red w/white stripe
Blue &amp; yellow blocks
Blue &amp; red blocks
White wired stripe
Blue &amp; white
Red &amp; blue blocks
Red &amp; blue blocks
Green w/white stripe
Red w/white stripe
Green w/white stripe
Red and blue blocks
Blue &amp; yellow blocks
Orange &amp; white blocks
Red &amp; blue blocks
Pink
White
Pink
Green w/white stripe
Red &amp; blue blocks
Red &amp; blue blocks
Red &amp; blue blocks
Green w/white stripe
Green w/white stripe
White wired stripe
Blue
Red &amp; blue blocks
White
Yellow wired stripe
Orange &amp; white blocks
Blue
Orange &amp; white blocks
Red &amp; blue blocks
Blue &amp; white
Orange &amp; white blocks
Red &amp; blue blocks
Green w/white stripe
Blue
Red w/white stripe
Blue
Green w/white stripe
Orange &amp; white blocks
Blue &amp; yellow blocks
Red &amp; b Lue blocks
Red &amp; blue blocks
Red &amp; blue blocks
Red w/white stripe
Blue
Yellow

Number
32
63
48
106
48
69
100
109
126
97
90
140
22
117
122
35
Worn
32
142
146
107
90
93
92
60
62
123
33
113
94
103
14
42
45
105
142
202
110
115
30
81
148
23
130
117
139
49
150
34

Location
Twp. Range Sec.
IS
2S
IN
IN
2N
2S
2N
2N
2N
2S
IN
4N
IN
IS
2S
3S
5S
IS
IN
IN
IN
lS
6N
4N
2N
2N
3N
IN
2N
IS
IS
lS
2N
lS
2N
2S
2S
4S
lS
lS
3N
3N
2N
IS
IS

93W 11
97W 21
93W 20
93W 25
97W 18
98W 10
95W 25
95W 25
95W 19
97W 36
93W 11
88W 17
92W 34
93W 13
Unit ll?
97W 20
98W 15
96W
1
95W
5
95W
2
95W 29
94W
4
?
92W
9
96W 36
95W 21
95W
9
98W
4
88W 32
93W 16
93W 14
94W 36
?
91W 31
9lW
7
95W
6
93W 28
99W 26
95W 33
98W
8
95W 19
93W 24
93W 36
92W 30
94W 13
95W 26
?
95W
5
92W
5

Remarks
Found dead along road.
Road kill.
Road kill.
Road kill.

Leg in neckband.
Found dead.
Found dead.
Archery.

Probable road kill.
Eartags missing.
Found shot.
Found dead.
Died after tagging.
Wounding loss.
Found dead.

Found dead.
Found dead.
Caught in fence.

Dead a long time.
Hunter kill.

Only collar found.

Found dead.
Found dead.

-------------------------------------------------------------------------------------

�-432-

Table 3. Recoveries of deer marked on the winter range in Game Management
Units 11, 22 and 23, March 1975 through February 1976.
Date of
Recovery
(1975)

Color

11-12
12-24
l2-?
1-4-76
1-5-76
1-19-76

Orange &amp; white blocks
Blue &amp; yellow blocks
Yellow
Blue &amp; yellow blocks
Red &amp; blue blocks
Blue

Neckband

Location
Number

Twp.

Range

Sec.

115
12
39
97
182
11

IS
IN
IS
IN
IN

92W
94W
93W
93W
96W
100W

34
27
23

2N

Remarks

Found dead.
Died in stackyard.

9
9

29

Recatch later killed by
coyotes.

Corrections to Previous Recovery Records
10-14-73 Red w/white stripe
11
10-15-73 White
24
10-18-73 No neckband-eartag L-2065
10-21-73 White
226

IS
4S

IS
2S

94W
95W
96W
98W

17
36
31
7

Eliminate these four, recoveries from the recovery list.

Died soon after marking.
Found dead.
Died soon after marking.
Died soon after marking.

�July, 1976

-433JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-3l

Work Plan: No.

16

j
Deer~Elk Investigations
I

Job Title

Job No.
I

-------

-------

2

----

Piceance Deer Study - Population (Density and Structure
)

Period Covered:
Personnel:

April 1, 1975 through March' 31, 1976

R. M. Bartmann and J. J. Klein, Jr.

ABSTPACT
The 1975-76 winter deer quadrat count yielded 9.1 deer per quadrat or 36.4
deer per mile2. The population estimate for the Piceance Basin is 24,206
3,724 deer at the 90 percent confidence level. The estimated population is
4,430 deer fewer than the predicted level of 28,636. The 2,113 deer classified during the post-season survey in Game Management Unit 22 gave a buck:doe:
fawn ratio of 16:100:80. This is two bucks and five fawns per 100 does fewer
than in 1974.

±

��-435-

PICEANCE DEER STUDY - POPULATION DENSITY AND STRUCTURE
Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use in
making annual harvest recommendations.

SEGMENT OBJECTIVES
1.

To establish mule deer population density and sex and age structure
estimate techniques for the Piceance winter range.
a.

Estimate the size of the total wintering deer population in Game
Management Unit 22.

b.

Estimate the sex and age structure of the wintering deer population
in Game Management Unit 22.

METHODS AND MATERIALS

Deer Density
See Bartmann (1974).

Population Structure
See Bartmann (1974).

RESULTS AND DISCUSSION

Deer Density
The 1975-76 winter quadrat deer count in the Piceance Basin was made January
11-14. The winter was quite mild and open and deer distribution was scattered.
Snow depths ranged from 6-10 inches at lower elevations to 12-15 inches on the
higher winter range areas. In a few locations, particularly the far western
and eastern ends of the basin, deer could still be found above the winter range
bounds designated for the count. The effect of this on the total deer population estimate is unknown but is believed minor. The total area involved is
relatively small and the western portion of the basin is considered a low deer
density area. After the count snow c.over conditions deteriorated steadily
through the rest of the winter.
Counting conditions were considered good although high wind caused a departure
from the normal quadrat counting sequence one day. A Bell 47G3B2 helicopter
was used and the pilot did an exceptional job.

�-436-

The 1,089 total deer counted convert to 9.1 deer per quadrat or 36.4 deer
per mile2. The population estimate for the basin is 24,206 + 3,724 deer
at the 90 percent confidence level (Table 1).
Support for the observation that deer were well scattered is that the
number of plots with no deer declined from 42 and 53 in 1973-74 and 197475, respectively, to only 27 this year. The precision of the mean density
estimator also improved from± 20 and ± 18 percent of the mean the previous
two years to ± 15 percent this year which was the desired level set at the
start of the study.
Comparison of the estimated winter deer population to the predicted population is complicated by' the lack of an estimated 1974-75 starting base.
The 1974-75 population estimate was considered low for reasons previously
indicated (Bartmann 1975), so the predicted winter deer population figure
was used instead. The 1975-76 winter deer population estimate is 4,430 deer
lower than the predicted population of 28,636 (Table 2). Several reasons can
be offered concerning the difference between the two values, but there is
currently no means of verifying any of them.

Population Structure
Post-season deer sex and age classifications were made from a Hughes 500C
helicopter December 12-16, 1975 in Game Management Unit 22. A fresh snowfall after the first day improved counting conditions from fair to good.
The 2,113 total deer classified included 177 adult males, 1,074 adult females
and 862 fawns for 3 buck:doe:fawn ratio of 16:100:80. This is two bucks and
five fawns per 100 does fewer than in 1974. If groups containing unknowns
are eliminated, the result is 141 adult males, 819 adult females and 634
fawns for a 17:100:79 buck:doe:fawn ratio. When applied to the total estimated deer population, these figures project to 2,028 bucks, 12,203 does and
9,875 fawns (Table 3).
The present method of collecting classification data (Le. no true sampling
strategy) does not justify statistical evaluation of the results. Efforts
will be made to remedy this situation by next winter.

LITERATURE CITED
Bartmann, R. M. 1974. Piceance deer study - population density and structure.
Colo. Div. Wildl. Fed. Aid W-38-R-28, Work Plan 16, Job 2, Job Prog. Rep.,
Game Res. Rep. July. pp. 363-370. (proc.).
Bartmann, R. M. 1975. Piceance deer study - population density and structure.
Colo. Div. Wildl. Fed. Aid W-38-R-28, Work Plan 16, Job 2, Job Prog. Rep.,
Game Res. Rep., July. pp.349-354.
(proc.).

Wildlife Researcher

�-437-

Table 1. Number of deer counted on 120 ~-mi1e2 quadrats on the Piceance
winter range in Game Management Unit 22, January 11-14, 1976.
Quad.

Deer

Quad.

Deer

Quad.

Deer

Quad.

Deer

1
2
3
4
5
6
7
8
9
10

2

31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60

18
14
0
8
16

61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90

II

13
4
15
10
22
9
5
0

36
32
12
0
9
7
3
8
9
0
11
20
0
3
0
0
0
3
0
2
0
8
37
22

91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
llO

15
4
0
0
9
5
10
0
0
5
1
4
0
13
2
6
0
0
9
0
15
8
25
8
4
14
29
24
0
14

80.66

90% Confidence Interval

8.98

9.1 ± (1.658) (.82) = 9.1 + 1.4

s-

0.82

Total Population Estimate

C. V.

99%

II

II
9

12
l3
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30

5
20
7
3
16
10
19
11
19
4
l3
5
14
17
2
6
0
0
24

n
X
x

=

II

5
0
1
15
9
16
5
5
16
21
6
10
3
5
0
15
5
5
15
26
0
1
0
0

1120

s

1,089

s

9.1

EX2

19,481

(EX)2

1,185,92l

2

x

0
39
31
5
II

III

ll2
ll3
ll4
ll5
116
ll7
118
ll9
120

24,206 ± 3,724

�-438-

Table 2. Calculations
Management Unit 22.

to predict the 1975-76 deer population in Game

Item

Bucks

Does

Fawns

Total

Predicted 1974-75 winter pop.

1,340

12,437

10,571

24,348

Corrected for 1974 post-season
buck:doe:fawn ratio (18:100:85)

2,167

11,979

10,202

24,348

843

983

2,856

4,682

1975 winter mortality est.
1975 Summer population

1,324

Fawns apportioned

3,673

3,673~--

4,997

14,669

50:50

Adjusted 1975 summer pop.
1975 fawn productivity
(80 fawns:100 does)

19,666

19,666

est.
11,735

1975 pre-season pop.
1975 harvest estimate

~7,346

4,997

!/

14,669

11,735

31,401

2,765

Predicted 1975-76 winter pop.

2,232

14,669

11,735

28,636

Estimated 1975-76 winter pop.

1,985

12,345

9,876

24,206

1/
- Harvest estimate is based on archery and rifle season random survey estimates
inflated by 25 percent to acknowledge wounding loss and illegal kill.

Table 3. Estimated deer population of the Piceance Basin (GMU 22) by sex and
age classes, 1972-73 to 1975-76.
Sex and Age Classes

1972-73

1973-74

1974-75

1975-76

Bucks

2,756

2,596

1,700

2,028

Does

9,743

11,778

9,254

12,303

Fawns

7,185

7,438

7,932

9,875

19,684 ±
3,990

21,812 ±
4,256

18,886 +
3,458

24,206 +
3,724

Total Deer

�-439-

July,· 1976

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-3l

Work Plan No.

16

Job Title

Job No.

Piceance Deer Study - Productivity

Period Covered:
Personnel:

Deer-Elk Investigations
3

_

and Mortality

April 1, 1975 through March 31, 1976

R. M. Bartmann, J. J. Klein, Jr., J. Shepherdson, K. Karrow,
S. Koster, P. Powell, P. Neil, D. Freddy and B. Gill.

ABSTRACT
A fawn:doe ratio of 80:100 resulted from classification of 862 fawns and 1,074
does during the post-season survey. This is five fawns per 100 does fewer
than in 1974. However, the calculated fawn:mother ratio is higher in 1975
due to the greater proportion of non-productive yearling does believed present.
The total pre-season fawn production was estimated at 11,735. The 1975 deer
seasons, both archery and rifle, were antlered-only in all the White River
,area units. The estimated harvests were 21 and 2,191 deer during archery and
rifle seasons, respectively.
The 1975 deer winter mortality survey showed a
mean 0.88 dead deer per plot for a total loss of 4,682 ± 1,224 deer at the 90
percent confidence level. Composition of the loss was 1,428 male fawns,
1,428 female fawns, 983 adult females and 843 adult males.

��-441-

PICEANCE DEER STUDY - PRODUCTIVITY AND MORTALITY
Richard M. Bartmann

P. S. OBJECTIVE
To establish methodology for the systematic collection of mule deer population density and structure data on pinyon-juniper winter range for use
in making annual harvest recommendations.

SEGMENT OBJECTIVES
1.

To estimate increments and losses to the Piceance deer population to
allow evaluation of density estimate techniques.
a.
b.
c.

Estimate productivity rates.
Estimate hunter harvest rates.
Estimate winter mortality rates.

METHODS AND MATERIALS

Post-Season Classifications
See Bartmann (1975).

Hunter Harvest
See Bartmann (1974).

Winter Mortality
See Bartmann (1974 and 1975).

RESULTS AND DISCUSSION

Post-Season Classifications
Post-season classifications were made from a Hughes 500C helicopter December
12-16, 1975 in Game Management Unit 22. The 862 fawns and 1,074 does classified yielded 80 fawns:lOO does. Elimination of groups containing unknowns left
634 fawns and 819 does, or 79 fawns:lOO does.
The 1975 ratio was five fawns less than in 1974. This may be misleading in
terms of over-all production as there was a presumably greater proportion of
non-productive yearling does in the 1975 population. While there are no harvest
data to confirm this hypothesis, some insight is gained from available population

�-442-

data. In 1974, the adult female segment contained an estimated 11 percent yearlings.
The fawn:mother ratio (Anderson 1965) calculated after
removal of the yearling component was 96:100. In 1975, yearlings comprised
an estimated 25 percent of the adult female population and a fawn:mother
ratio of 107:100 resulted, thus suggesting higher production despite a lower
fawn:doe ratio. However, in both cases the differences are small enough to
easily be attributable to chance.
As the hunting season was again bucks only, wherein no significant antlerless loss is assumed, the post-season fawn:doe ratio was used without
adjustment to give a pre-season production of 11,735 fawns.

Hunter Harvest
The 1975 regular deer season in the White River area units was antlered only
and extended from October 25 through November 4. The archery season, also
antlered only, ran from August 16 through September 21. The total harvest
estimate for Unit 22 ,from the Game Management Section's random surveys was
2,191 deer during the regular season and only 21 during the archery season.
These figures are in~lated by 25 percent to acknowledge wounding loss and
illegal kill.
Six hundred fifty-nine antlered deer from Unit 22 were recorded through
three Division check stations during the first nine days of the season.
Of these, 365, or 55 percent, were field aged as yearlings.
This compares to
30 percent yearlings in the 1974 check; another indication of better fawn
survival during the 1974-75 winter.

Winter Mortality
The 1975 deer winter mortality survey was made April 21 through June 4. The
desired crew size of seven people was achieved the majority of the time with
the minimum never being below six. Forty-nine deer carcasses judged to have
died in the 1975-76 winter were found for a mean 0.88 deer per plot, or 4,682
± 1,224 total dead deer (90% c.l.) on the winter range (Table 1). This is a
19 percent loss based on the 1974-75 predicted population (the estimated population based on the quadrat census was considered low and was not used). The
loss was only six percent lower than in the preceding year despite an apparently
much milder winter.
Composition of the estimated loss with unknowns apportioned among the various
classes was 1,428 male fawns, 1,428 female fawns, 983 adult females and 843
adult males.
Fawn losses are apportioned between the sexes on a 50:50 basis.

LITERATURE

CITED

Anderson, A. E. 1965. Population density and structure.
Colo. Game, Fish
and Parks Dept: Fed. Aid W-I05-R-4, Work Plan 4, Job 1, Job Compl, Rep.,
Game Res. Rep. Jan. pp. 47-74. (proc.).

�-443-

Bartmann, R. M. 1976.. Piceance deer study - productivity and mortality.
Colo. Div. Wildl. Fed. Aid W-38-R-28, Work Plan 16, Job 3, Job Prog.
Rep., Game Res. Rep., July. pp. 371-380. (proc.).
Bartmann, R. M. 1975. Piceance deer study - productivity and mortality.
Colo. Div. Wildl. Fed. Aid W-38-R-29, Work Plan 16, Job 3, Job Prog.
Rep., Game Res. Rep., July. pp. 355-362. (proc.).

A

/ '-);'

/1/'

/--;

)'1

/ /c.&lt;,J

z:

-r:»

Prepared by __-+(/_I~_-~~'~,_·
~·_"~~~~~-~~.~.~~'~.~'~"~L1~/?~.~_/'~,~,~~,,~.~~
Richard M. Bartmann
Wildlife Researcher

�Table 1. Results of dead deer searches on 56 1/8-mi1e2 quadrats on the Piceance winter range in Game
Management Unit 22 during spring, 1975.

Quad.

Male

2
7
9
10
11
12
13
16
17
21
23
24
25
26
27
32
33
34
35
36
37
40
42
50
52
53
59
62

1

-

-

1

-

-

-

Adults
Female

2

-

-

Unk.

Male

Fawns
Female

Unk.

Quad.

Male

Adults
Female

Unk.

Male

Fawns
Female

-

-

-

-

-

-

1

1

1

1

65
68
70
73
74
75
76
78
81
84
86
87
91
93
95
96
99
101
107
109
110
111
112
114
115
116
118
119

-

2

-

-

-

-

-

1

-

-

1

-

-

-

-

-

-

-

-

-

-

-

1

-

1

1
2

8

13

-

-

-

1
1

1

1

-

-

-

-

-

1

2

1

-

-

-

-

1

1
1

1

-

1

1

1

1
1

-

-

-

2

1

1

-

1

1

1

1

-

-

1

-

-

-

1

-

-

-

-

1

-

-

-

1

5

5

9

9

1
1

Unk.

-

1

Totals

I

~
~
~
I

�July, 1976

-445JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~-----

Project No.

W-38-R-3l

Work Plan No.

16

Deer-Elk Investigations
Job No.

5

--------~----------.-----------

Job Title Piceance Deer Study - Food Habits Technique Development
Period Covered:
Personnel:

April 1, 1975 through March 31, 1976

R. M. Bartmann, W. R. Regelin, P. H. Neil, D. R. Reichert and
J. J. Klein, Jr.

ABSTRACT
Predator problems caused cancellation of all deer grazing comparison trials
except for those scheduled in early October. A U.S. Forest Service Research
Note entitled "The influence of Supplemental Feed on Tame Deer Food Habits"
is being prepared with Forest Service cooperators covering the results of
that work. Token grazing trials were conducted in December, January and
February to observe tame deer performance on pinyon-juniper winter range.
The 44 trials averaged 66 minutes each. The mean number of bites per trial.
declined from 301 to 123 to 49 from December through February. General
observation of the deer fawns indicate a general disinterest in grazing,
curiosity about their surroundings and a desire to play, and a reluctance
to move about in deep snow.

��-447-

PICEANCE DEER STUDY - FOOD HABITS TECHNIQUE DEVELOPMENT
Richard M. Bartmann

P. S. OBJECTIVE
To determine if forage selections by "artificially" maintained tame deer
are similar to those of "naturally" maintained tame deer.

SEGMENT OBJECTIVES
1.

Construct pen facilities for raising and handling tame deer.

2.

Raise and maintain two groups of eight tame deer each for technique
evaluation.

3.

Measure forage selections of "artifically" and "naturally" maintained
tame deer.

4.

Evaluate forage selection data to determine similarities between
"artificially" and "naturally" maintained tame deer.

5.

Make recommendations concerning the use of "artificially" maintained
tame deer for use in forage use investigations.

METHODS AND MATERIALS
A series of three 125 x l50-foot pens was constructed immediately west of the
Little Hills headquarters. A 10 x 20-foot shelter was built in each pen and
the entire outsid.e perimeter was dog and coyote-proofed with a poultry netting
apron.
Fawn raising was done cooperatively by now and Rocky Mountain Forest and Range
Experiment Station personnel at facilities in Fort Collins. Disease problems
reduced the number of available fawns to nine by time the grazing field work
was to begin. The fawns were all brought to Little Hills September 11, 1975.
Five of them were in~ediately placed in a 98-acre pasture and the other four
in a pen at the Station headquarters. Three days later two of the pasture
fawns were killed by predator(s) and the remaining three were brought back to
the holding pen.
After discussion with cooperators, it was decided to go ahead and conduct the
first trials as planned, but in another area with fewer observers. All later
trials were cancelled. Accordingly, on September 23, three fawns were taken
to a four-acre pen in the paddock complex on the ridge about two miles north
of the Station. Hay and concentrate feed were withdrawn over a five-day
period until fawns in the paddock were totally dependent upon native paddock
forage. Grazing trials were started September 30 and ran for 10 consecutive
days.

�-448-

Because considerable amounts of several peoples' time were scheduled for
the four cancelled grazing periods, it was decided that several token
trials throughout the winter might provide insight to tame deer performance
on pinyon-juniper winter range. Therefore, three grazing periods were
scheduled; one each in December, January and February. The December and
January trials lasted four days, with two days each spent on pinyonjuniper-mixed browse and pinyon-juniper-sagebrush winter range types.
The February trials lasted only three days and all were conducted on a
pinyon-juniper-mixed browse area. Grazing areas were shifted from higher
to lower winter ranges as the winter progressed. Two observers and four
deer were used during all periods except the last when a third observer
was present one of the three days. Two trials were usually run in the morning
and two in the afternoon each day. A trial consisted of an observer following
one deer for 1 to 1-1/2 hours and recording the number of bites by species.
Portable tape recorders were used in the field and the data later transcribed
onto forms.

RESULTS AND DISCUSSION
A U.S. Forest Service Research Note entitled "The Influence of Supplemental
Feed on Tame Deer Food Habits" is being prepared together with Forest Service
cooperators covering the methodology and results of the one planned series
of grazing comparison trials that was conducted. Continuation of the comparisons is planned for Segment 32 during winter and spring stress periods.
The same 98-acre pasture will be used after efforts to block predator entry
points are completed. This, together with the use of adult deer, will hopefully eliminate most problems.
Results of the preliminary winter grazing trials showed a steady decrease
in the mean number of bites per trial during December, January and February
from 301 to 123 to 49. This occurred even though hay and concentrate feed
in the holding pen were restricted the last two periods. Total natural
forage availability was probably lowest in January when 12 to 16 inches of
snow were present on the grazing area and highest in February when snow cover
was spotty and bare ground prevalent. Only 4 to 6 inches of snow were present
in Decemb er .
The 44 total trials occupied 2,925 minutes with a mean of 66 minutes per trial
and a range of 45 to 100 minutes (Table 1). Ingestion of 48 species were recorded including 16 shrubs, 24 forbs and 8 grasses. However, only 17 species
had 10 or more bites recorded for them during anyone trial and only these are
listed in Table 1.
Generalizations of deer feeding performance included: (1) A general disinterest
in grazing, particularly as the winter progressed; (2) A display of curiosity
about their surroundings often accompanied by playfulness; and (3) A reluctance
to move about in deeper snow (all deer were fawns).
More frequent exposure to the natural environment may help overcome the first
two situations and, in the third instance, older and larger deer may be better
able to cope with deep snow.

�-449-

Based on the above observations, it is cautioned that the data not be
construed to represent wild deer diets. While most of the species taken
by tame deer were probably eaten by wild deer, the relative proportions
may be misleading. Also, the small areas grazed are not representative
of the entire winter range as not all species were present in each area.

Prepared by
Richard M. Bartmann
Wildlife Researcher

�Table 1. Summary of total bites by species taken by tame deer during three monthly grazing periods on
two winter range types. 11

Species

Pinyon-Juniper-Mixed Browse
Dec.
Jan.
Feb.
All

Pinyon-Juniper-Sagebrush
Dec.
Jan.
All

881
798
377
86
92
2
2

1,846
561

Shrubs
Purshia tridentata
Cercocarpus montanus
Ame1anchier utahensis
Symphoricarpos oreophi1us
Quercus gambelli
Artemisia tridentata
Gutierrezia sarothrae
Atrip1ex confertifo1ia

95
93
327
206
19
29

13
9

68
13

12
12

989
900
772
305
111
31
14
12

499
385

2,345
946

3

"3

18

18

2

2

32

32

Forbs

I

~
\Jl

Lupinus caudatus
Cryptantha sericea
Hap10pappus nutta11ii
Aster 1eucanthemifo1ius
Eriogonum 10nchophy11um
Cirsium sp.
Hap10pappus acau1is

6
10
13
7

26
35
130
5

184
63

32
45
143
12
184
63

o
I

28
127

28
127
10

25

25

10

Grass
Sitanion hystrix
Poa sp.
Total Bi tes Jj
Number of Trials
Total Minutes

1/Species

11

4

1

16

6
28

6

28

2,327

821

583

3,731

2,485

1,140

3,625

8

8

12

28

8

8

16

610

510

715

1,835

580

510

1,090

listed are those for which 10 or more bites were recorded during at least one grazing trial.

liTota1s include bites on all species recorded during grazing trials.

�.,
-451-

July, 1976

JOB PROGRESS REPORT

St are of

COLORJ...DO
--------~~~~~----

Pr oj ect

No.

H-38-R-31

Deer-Elk Investigations

18

Wo:r:k.Flan No.

Job No.

1

Deer and Elk Management Study
Period Covered:
Personnel:

April 1, 1975 - March 31, 1976

Raymond J. Boyd and Thomas M. Pojar

ABSTRACT
Workshops designed to familiarize field management personnel with the
population simulation approach to game management were conducted. Of
120 WCO's and Area Supervisors in the State, 110 of them attended at least
one all-day workshop. The workshop program stressed extensive review of
the mechanics of the ONEPOP population model, input data and output interpretation.
Data collection and investigations are recommended based on my evaluation
of their potential for accomplishing the most toward enhancing the realism
of the population simulations.

��-453-

DEER AND ELK MANAGEMENT STDDY
Thomas M. Pojar

In addition to the stated segment objectives it became obvious during this
segment that understanding and acceptance of population modeling by all
levels of management was needed before the full benefits of this management tool could be realized. Therefore, all-day modeling workshops designed
to familiarize management personnel with the ONEPOP program and its output
were conducted. Nearly half of this segment was spent in preparation,
conducting, and fo11ow--up work associated with the workshops.
Consequently,
work towards the segment objectives was not fully completed.

P. S. OBJECTIVE
Devise and test a statewide deer and elk management system.

SEGMENT OBJECTIVES
1.

Identify data that are most useful for improving estimates of population
parameters and recommend procedures for collecting such data by DAD.

2.

Project average population parameters on a long-term basis for those
alternative management plans that are feasible for each DAD.

3.

Make final recommendations for data gathering and analysis procedures
for each DAD in a manual for the appropriate operations section.
METHODS AND MATERIALS

The following discussion on data evaluation partially fulfills the first
segment objective. The data analysis and utility is common to all deer and
elk Data Analysis Dnits (DAD's) in the State except when specifically excluded.
The data collection and investigations that are recommended are based on my
evaluation of their potential for accomplishing the most toward enhancing the
realism of the population simulations.

Harvest
Estimates of harvest were available for all deer and elk data analysis units
(DAD's) in the State. The precision of these estimates become a point of
interest because the outcome of a simulation for a particular DAD is greatly
affected by the number of animals harvested from the population. The 95 percent confidence intervals for the 1974 deer and elk harvest by DAD are listed
in Tables 1, 2. 3, and 4. The variances for resident (S~) and nonresident
(S~) harvest was calculated by:

�-454-

n)

N (N -

where

N
n
y

Total hunters
Total usable returns
No. of hunters reporting success, by DAU

The confidence interval is then calculated:
Estimated harves.t by DAU +

2.V/~~

There are only two ways of improving the preclslon of the deer (both sexes)
and elk (bull) harvest estimates. One is to increase the number of hunters
surveyed. This would result in more usable returns per DAU, however, the
improvement in the estimate would be in proportion to the number of hunters
in each DAU. Only slight narrowing of the confidence limits would result
for DAU's with low harvest.
The relation between the estimated deer harvest (X) andc!.~s~ + S~R (y) is:
y

=

85.29 + .0438X.

Under the present hunt Lng system of bucks only or either sex seasons, DAU's
with a harvest of 1000 deer will result in a confidence interval of about
13 percent of the estimated harvest.

±

The relation between

t he

estimated bull elk harvest (X) and 2.

J +
S~

S~R (y)

is: y = 50.56 + .1049X. With no control over where the purchaser of a
Regular Bull license will hunt, the confidence interval is related to the
number of bulls harvested by DAU. A harvest of 500 bulls from a DAU will
result in a confidence interval of about + 20 percent.
The second method of improving the harvest estimate is to issue only specified
deer and elk licenses statewide. This would permit sampling each DAU
at the level necessary to obtain the desired precision in the harvest estimates.
This is possible now for estimating the harvest of cow elk (Table 2).

Age Structure of the Harvest
Of the 4,307 elk aged from the top ten ranking DAU's, 2119 (49.2%) of the ages
were obtained from teeth collected by hunters. The Division provided tooth
envelopes to persons with Specified or Cow licenses. A total of 10,113 tooth
envelopes were sent and 21 percent (2,119) were returned with usable teeth.
The success rate of hunters with Specified and Cow permits is around 50 percent.
Therefore, of the estimat~d 5,056 successful hunters 41.9 percent returned the
tooth envelope. This occurred without any public information effort to encourage
hunters to comply with the tooth envelope request.

�-455-

Table 1.

Confidence intervals (95%) for 1974 deer harvest by DAU.

+ Percent

Lower
C.L.

Upper
C.L.

DAU

Harvest

+ No.

7

7370

364

4.9

7006

7734

12

6745

357

5.3

6388

7102

6

1366

160

11.7

1206

1527

19

4745

300

6.3

4446

5045

30

1822

187

10.3

1635

2010

2

958

137

14.3

821

1095

24

2890

233

8.1

2657

3124

8

2912·

238

8.2

2673

3150

16

1833

190

10.4

1643

2024

25

1934

194

10.0

1740

2128

20

2784

231

8.3

2553

3015

14

1684

182

10.8

1502

1867

13

516

101

19.6

415

618

29

1404

166

11.8

1239

1570

18

2042

199

9.7

1843

2241

27

915

135

14.7

781

1050

33

1014

140

13.8

874

1154

4

992

140

14.2

852

1132

11

241

69

28.4

173

310

1

766

121

15.8

646

887

32

924

135

14.6

789

1060

17

560

106

18.9

454

666

3

96

44

45.6

52

140

31

328

81

24.6

248

409

21

389

87

22.4

302

476

10

709

119

16.8

590

828

22

348

83

23.7

266

431

15

318

80

25.1

238

398

26

359

84

23.5

274

444

23

129

50

38.8

79

179

�-456-

Table 2.

Confidence intervals (95%) for 1974 cow elk harvest by DAU.

DAU

Harvest

+ No.

+ Percent

Lower
C.L.

Upper
C.L.

E 2

866

57

6.6

808

923

E 4

56

16

28.7

40

72

E 5

72

17

24.0

55

89

E 6

1,046

61

5.8

985

1,107

E 7

100

21

21.2

79

122

E 8

136

24

17.8

112

161

E10

7

6

86.1

1

l3

El3

80

19

23.5

61

99

E 9

9

7

74.6

2

16

Ell

4'

4

78.4

1

8

E12

35

13

36.2

22

48

E14

544

46

8.4

498

590

E15

101

21

20.3

81

122

E16

147

23

15.8

124

171

E21

432

42

9.7

390

474

E20

63

16

25.2

47

79

E25

408

40

9.8

368

448

E26

120

24

19.8

96

143

E24

215

29

13.5

186

244

E29

10

6

58.9

4

16

E30

56

16

27.6

41

72

E31

465

40

8.5

425

505

E32

153

26

16.9

127

179

E33

20

10

49.7

10

30

E27

74

19

25.5

55

93

�-457-

Table 3.
by DAU.

Confidence intervals (95%) for 1974 regular bull elk harvest

+ No.

+ Percent

Lower
C.L.

Upper
C.L.

DAU

Harvest

1

Specified

2

1,164

171

14.69

993

1,335

3

528

117

22.23

411

645

4

211

74

35.20

137

285

5

207

72

34.49

135

279

6

Specified and Regular

7

368

98

26.59

270

466

8

464

109

23.42

355

573

10

203

72

35.15

131

275

11

44

34

76.04

10

78

12

114

55

48.12

59

169

13

322

91

28.39

231

413

14

1,456

192

13.19

1,264

1,648

17

Specified

19

Specified

20

285

84

29.34

201

369

21

1,363

184

13.48

1,179

1,547

24

700

130

18.58

570

830

25

846

145

17.13

701

991

26

280

86

30.61

194

366

27

128

58

45.19

70

186

28

Specified

29

64

40

61.96

24

104

30

285

84

29.34

201

369

31

967

151

15.59

816

1,118

32

756

138

18.19

618

894

33

320

90

28.03

230

410

�-458-

Table 4.
by DAU.

Confidence intervals (95%) for 1974 specified elk hunt areas,

Harvest 1/

+ No.

+ Percent

Lower
C.L.

Upper
C.L.

Bull

114

12

10.53

102

126

Cow

185

22

12.00

163

207

Bull

25

2

9.14

23

27

Cow

20

2

12.50

18

22

Bull

32

11

35.07

21

43

Cow

12

2

17.80

10

14

DAU
No.

17

19

28

1/
- These harvest figures are not in exact agreement with those found in the
1974 Harvest Book. This is because these figures are not adjusted for
"unknowns" (Page 16 in 1974 Harvest Book).

To hunt in deer units 20 and 29 each hunter was required to first obtain a
permit in addition to a valid license. This permit was free of charge and
could be obtained at the Division's Northeast Regional Office. Upon obtaining
a permit, the hunter was given a tooth envelope and encouraged to return it if
he harvested a deer. A total of 504 deer were harvested from units 20 and 29,
and 287 usable teeth were returned. Of the successful hunters, 57 percent
complied with the tooth collection request.
The difference between deer and elk hunters' rate of compliance to the tooth
collection request may be due to the personal contact with the deer hunters.
It is obvious that large numbers of teeth can be collected using the hunter
tooth envelope approach. Probably a much larger proportion of hunters would
comply with the tooth collection request with an extensive public information
campaign.
There is a serious discrepancy in the percent of young in the harvest when
comparing the tooth envelope method and check station method of collecting
aging data (Figs. 1 and 2). This discrepancy appears to be linked in some
way to hunters' response to the tooth envelope request and/or to check
stations. The possibility of misidentifying teeth from young animals that
are mailed to the laboratory is minimal (Adrian, pers. comm.).

�50

•\
\
\
\

40
Mail-in from Units 20 and 29
_______

n = 287

Check Station Ages from Units 58 and 581 n = 185

30
I

E-&lt;
Z

~

U

&lt;o

~

\J1

I

po::

~
p..

20

10

01

I

I

I

I

1

2

3

4

5

6

7

8

k

=1

9

10+

AGE

Fig. 1.

Age distribution

of antlered deer harvested

in 1974 from two front range areas.

�30

25

Mail-in

n = 849

t
/

20

\

'" \
-------

/

V

Collected

at Check Station

n = 247

15
I
.p..
0\

o
I

~
~
J;;:l

\
\

p..,

~
\

10

\
\.
\

\

5

•... ...

.....
t-

"

Q

''-,
"

o

~

to

8

9

-

C

-

-

-

---~
0 ••••••
_......

.//

........._

•

I

C

1

2

3

4

5

6

7

12

13

in 1974 from the White River

elk

10

11

14

15+

AGE
Fig.

2.

Age distribution

of antlerless

elk harvested

data

analysis

unit.

�-461-

An investigation to determine which of the two methods is biased and the
extent of the bias is needed so that the aging data can be interpreted
properly.

Sex and Age Ratios, and Trend Counts
Fawn:doe and buck:doe ratios are still collected in a designated manner in
only a few DAU's. The general opinion of most Division personnel seems to
be that the only feasible means of collecting these data is by helicopter.
This opinion is not supported by the helicopter and ground classification
comparisons conducted by Gill (1972). The amount of time needed to classify
a certain prescribed number of animals from the ground compared with classifying
from a helicopter has been used to justify the use of helicopters.
In the
past this may have been true, however, recently Bowden and Anderson (1975)
have developed a sampling method which, in my opinion, makes ground classification a feasible method of obtaining herd structure data. These authors
suggest that further refinement of this method is possible. The fact remains
however, that this method provides an estimate of the fawn:doe ratio based on
statistically sou~d design and analysis procedures. In very general terms,
it would take approximately 20 to 40 person days to estimate the fawn:doe
ratio in a DAU within 10 percent at the 90 percent confidence level.
Application of this method to estimate elk cow:calf ratios has not been tri.ed.
Confidence intervals should be calculated for the age ratio data that are
currently collected by helicopter. This can be done if the data are summarized
in terms of "groups" of animals classified.
The validity of sex ratio classification has not been established. It is
suspected that sex ratio data collected either from the ground or air is
biased and may not accurately depict the sex ratio in the population (Gill
1976). It is common for the variance of sex ratio data to be quite large.
Even with these shortcomings, sex ratio data should be collected since it can
be done in conjunction with age ratio data collection, and if the biases in
these data can eventually be quantified then historic sex ratio data could be
corrected for bias.
Population trend counts are made in several of the deer and elk DAU's. Trend
counts, especially on deer, are plagued with problems. Not the least of which
are fluctuating deer distribution with varying weather conditions and unstable
counting conditions. At least in one case, there was actually a significant
(P&lt;.05) inverse relationship between the winter trend count and the following
season's antlered harvp-st (Baker 1975). Meaningful interpretation of trend
count data that is severely affected by either unstable animal distribution
or counting conditions would seem to be impossible.
Trend counts that could detect a 20 percent change in the population would be
useful. However, until census methods are adequate to detect this amount of
change in a population there will be no way of testing the precision or accuracy
of a trend count. At this point in time it seems most feasible to concentrate
on applying existing census methods and continue efforts to improve them
rather than continue trend counts.

�-462-

Currently, there are three Federal Aid projects (Bowden &amp; Anderson 1975,
Bartmann 1975, Freddy 1975) to census mule deer populations in various habitat
types. To apply any of these methods, approximate winter distribution needs
to be defined for each population. Knowledge of relative animal density
on the winter range allows for sampling stratification which can result in
considerably increased precision in estimates of population size. The
status of our knowledge of relative animal density and winter distribution
needs to be evaluated and census efforts initiated.
Tested, designed elk census methods are lacking. This is probably because
elk tend to form large groups in winter and managers may feel that an intensive effort to count all conspicuous groups approaches a total count of
the herd. However, the number of groups missed and the proportion of the
total animals counted remains unknown. A statistically designed sampling
scheme would minimize these problems and provide a means to calculate the
variance associated with the population estimate.

Population Simulation Workshops
Population modeling workshops were conducted for field management personnel.
Attendance at each all-day workshop varied from two to seven persons. A
total of 110 wca's and Area Supervisors attended at least one workshop.
The workshop program included extensive review of the mechanics of the
model, input data and interpretations of output.

LITERATURE CITED
Baker, B. D. 1975. Evaluation of interagency browse utilization and pellet
group count methodology. In Boyd, R. J., T. M. Pojar, B. D. Baker.
1975, Deer and Elk Management Study. Part II. Fed. Aid Game Res. Rep.,
July, Part 2. p. 365-412.
Bartmann, R. M. 1975. Piceance deer study - population density and structure.
Fed. Aid Game Res. Rep., July, Part 2, p. 349-354.
Bowden, D. C., and A. E. Anderson. 1975. Evaluation of herd structure
methodology.
Fed. Aid Game Res. Rep., July, Part 2. p. 475-499.
Freddy, D. J. 1975. Middle Park deer study - experimental harvest regulations.
Fed. Aid Game Res. Rep., Part 2. p. 209-240.
Gill, R. B. 1972. Middle Park deer study population density and structure.
Fed. Aid Game Res. Rep., July, Part 2. p. 169-177.
Gill, R. B. 1976. Mule deer management myths and the mule deer population
decline. Mule Deer Decline in the West Symp., Proc. l:(In Press).

Prepared

':;t;L;:;;;~/.UJ;ft'2
Thomas M. Pojar
.
Wildlife Researcher

�July 1976

-463-

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-38-R-3l

Work Plan No.
Job Ti.tle
Period Covered:

19

Deer-Elk Investigations
Job No.

l

_

Evaluation of Radio Telemetry
April 1, 1975 - March 31, 1976

Personnel: Mark Alan, James Albert, Allen E. Anderson, Allison Broida,
Charles DeYoung, Howard Geduldig, Jennie Goldberg, Wendy Haas, Dan Kimes,
Beth MacConnell, Arnold Olson, Chris Ribic, Dick Rubin, James Shepherds on ,
Carol Smith, Laverne Stelter, George Stewart, Colorado Division of Wildlife;
G. L. Terrell, Colo., Dept. of Agriculture; J. C. DeMartini, C. P. Hibler,
S. Henry, J. G. Wegrzyn, Colorado State University.
ABSTRACT
Thirty newborn mule deer fawns and nine coyotes were captured, aged, weighed,
measured, radio-collared, ear-tagged, and released. Twenty-four fawns and
nine coyotes were radio-tracked with ground and air high frequency (157 MHz)
telemetry equipment and six fawns with low frequency (46.5 MHz) telemetry
equipment within and adjacent to a 12,045 acre study area about 30 miles
northwest of Fort Collins. Based on 24 fawns the mean
SD number of days
from capture day to day found dead was 76.2
104.1. Five fawns died from
unknown causes, 12 from probable predation, four from probable abandonment
by doe, two from disease, and one from an accidental injury. One fawn disappeared, three lost their collars and two were presumed alive. Fawns were
relocated 1,600 times and there was great individual variation in the daily
distances moved from their capture site even among twins. All nine coyotes
were presumably still alive. Coyotes were relocated 385 times and the mean
distance moved from their capture sites to relocation sites was about one
mile, with a range of 0.2 to 4.4 miles from October through March. A
summary of data from 53 fawns radio collared since June 1974 shows that the
fate of 25 fawns is not known and includes those that disappeared, lost
collars, and those found dead from unknown causes. A similar summary for
coyotes showed that 7 of 23 have died mainly by shooting.

±

±

�-464-

EVALUATION OF RADIO TELEMETRY
Allen E. Anderson

The use of radio telemetry to assess neonatal mortality in deer (Odocoileus
spp.) has been briefly reviewed (Anderson 1975). It was pointed out that
relatively little has been published on deer-coyote (Canis latrans) interactions in Rocky Mountain mule deer (Odocoileus hemio~emionus)
and that
the use of radio telemetry on such studies has been quite limited, probably
because of the mountainous terrain typical of most mule deer habitats. To
our knowledge, the present study is the first to radio collar free-ranging
Rocky Mountain mule deer fawns and coyotes and conduct a near-daily telemetric surveillance of both species on a limited area. This report presents
results of the second year of field work mainly in tabular form. A more
complete analysis and interpretation of results will be presented in a subsequent report.
P. S. OBJECTIVE
To test and improve the field and statistical methodology used to estimate
selected population parameters of mule deer and coyotes.

SEGMENT OBJECTIVES
1.

Test the feasibility of using and perhaps improving radio telemetry to
estimate fawn mortality rates and coyote activity in mountainous habitats.

2.

Analyze data and submit reports for publications.

METHODS AND MATERIALS
In general, procedures and equipment used during 1975-76 were similar to those
described previously (Anderson 1975) and are not described herein. In that
report, recommendations were made for certain equipment and procedural
changes. Those incorporated are described as follows.

Radio Telemetry Eguipment
We used both high (157.0 MHz) and low (46.5 MHz) frequency equipment for radio
tracking fawns and high (157.0 MHz) frequency equipment for radio tracking
coyotes. The high frequency equipment was made by Davtron, Minneapolis, Minnesota and the low frequency equipment by Albertronics, Inc., College Station,
Texas. The Davtron, 24 channel receivers were used in conjunction with four
element, portable Yagi antennae. Both aerial and ground tracking were done

�-465-

with the same equipment. Aerial tracking was conducted with one antenna
mounted on each wing. A relay switch permitted more effective searches from
either side of the plane. Unfortunately, Albertronics, Inc. could not
provide new receivers so all low frequency work was done with very well
used Johnson 350 DF and Johnson Messenger III receivers. See Anderson (1975)
for a description of radio tracking with the Albertronic equipment. For
both high and low frequencies the transmitter-battery package used on fawns
was sealed in acrylic and secured to collars fabricated by us with several
wrappings of heavy plastic tape.
For fawns, the Geduldig .(1975:9) modification of the molded plastic, expandable collar of Fashingbauer (1962) was used again. In order to accommodate
the larger, heavier, high frequency, transmitter-battery package however,
collar width was increased from l~ inches to 2 inches and collar thickness
from 0.0625 to 0.09375 inches. In addition, the fawn transmitters were of
three types: (1) mercury switch, wherein fawn movement increased the signal
pulse rate; (2) temperature, wherein changes in pelage-skin interface temperatures change the signal pulse rate; and (3) stable, wherein the signal
pulse rate remained at a relatively constant rate. In type (2) the sensor
was attached to a flexible plastic extension of the collar and impinged on
the dorsal portion of the neck.
For coyotes, the transmitter-battery package was molded and bonded with
acrylic to a one-half inch width collar of heavy two-layered, nylon webbing.
The transmitter antenna was sewn between the two layers of nylon webbing .
The radio collar was secured to the neck of the coyote by two bolts through
pre-punched holes. The entire unit was designed and fabricated by Davtron.
Capturing and Radio Tracking Fawns
The 1974-75 work with t~ree persons had indicated that the flight response of
individual fawns to humans tended to be predictable on specific sites. Since
some sites also attenuated radio signals in a predictable manner, it seemed
likely that familiarity with individual fawns and their habitats should
increase the efficiency of radio tracking. Since the 1975 goal was 40 captured fawns, four people were assigned to the north, three to the southwest,
and four on the southeast portions of the 12,045 acre study area. Those
persons involved in capturing a particular fawn were primarily responsible
for radio tracking that fawn on a daily basis. An attempt was made to
visually examine each fawn located telemetrically.
In addition to daily
diurnal telemetric locations, the movements of up to three selected fawns
were monitored telemetrically for about 16, 24, 48, and 72 consecutive
hours during August 1975. All telemetric and visual observations of collared
fawns were plotted on a topographic map and pertinent information recorded
for each observation (Appendix I).
Assessing Fawn Mortality
On locating a fawn carcass, the observer(s) made a detailed written record
of carcass appearance and damage, if any, and searched the immediate area
for tracks and scat. Photographs or sketches were usually made of each fawn

�-466-

carcass. Fawn carcasses in good condition were taken immediately to the
Wild Animal Disease Research Center at Colorado State University for diagnosis.
As an aid in possible future diagnosis of disease in collared fawns, nasal
swabs were taken from 19 fawns at capture. In addition, eleven collared
fawns were recaptured and about 25 cc of blood was aspirated from the juglar
veins of each fawn by Dr. J. C. De Martini and S. Henry. Analyses of some
disease dia~lostic components of these blood samples was done at the Veterinary Clinic at Colorado State University, Fort Collins.
Ear-Tagging Fawns
In an effort to more readily identify individual collared fawns, it was
decided to increase the size of the ear tag. Black and white plastic ear
tags with numbers almost two inches in height seemed to meet identification
requirements.
RESULTS
Mule Deer Fawns
Thirty mule deer fawns were captured, weighed, measured, examined (Table 1)
and released adjacent to the study area (Fig. 1). Only four fawns (43, 52,
6lA, 62A) were ear+t agged because the plastic ear tags used seemed so large
as to possibly handicap the fawn or attract predators. Some temporal and
behavioral aspects of fawn capture are detailed in Table 2.
The Davtron transmitters were not delivered as requested and at the peak of
fawning none were available. Thus, when finally delivered, about one week
later, the fawning season was essentially over and we were 10 fawns short
of our goal of 40. Details on all transmitter-collar units used during 1975
are given in Table 3.
Blood values for 10 mule deer fawns are listed in Table 4 and results of the
nasal swabs from 20 mule deer fawns in Table 5. As compared to the blood
cellular values from 8 captive Rocky Mountain mule deer fawns 23 to 68 days
of age given by C0wan and Bandy (1969), the values in Table 4 are much lower
for hemoglobin and packed cell volume, but similar for leukocyte values.
The etiological significance of both the blood or nasal swab values is not
clear. Unfortunately, these data are not available for the one fawn (no. 75)
with a definite diagnosis of disease or the other (no. 62C) with a probable
diagnosis of disease (Table 6). Another fawn (no. 56) with a very high total
leukocyte count of 12.4 x 103 could have been a disease victim, but was last
seen on August 23, 1975.
How long did the radio collared fawns survive? Using the "day of capture to
day found dead" column (Table 7) as one approximation it seems that 24 fawns
survived (X + SD) 76.2+ 104.1 days with a range from 2 to 333 days. Significantly, there were very large differences among the north, southwest, and
southeast areas of fawn captu~e. There are obvious differences in habitat
with the longest survival period being in the north area, which also has the
steepest and most forested (shaded) habitat (Fig. 1). None of these presumed
differences have been tested for statistical significance.

�Table 1.

Some morphological

attributes

of 14 male and 16 female mule deer fawns radio-collared
Lengths
Time

Body Wt
(kg)

Hind
Foot

0835
1320
1010
1544
1835
1800
2010
2034
1327
1155
1155
1201
1030
1028
1343
0940
1005
1043
1145
0935
0906
1035
0954
1839
1036
1953
1957
1937
1555
1055

3.9
2.6
3.5
3.1
3.9
4.0
4.8
5.7
3.6
4.4
3.5
5.2
3.3
2.3
3.7
4.7
3.4
4.1
3.0
4.8
4.6
3.9
5.4
3.5
3.8
5.3
4.0
3.0
4.3
4.9

29.0
23.5
25.0
24.5
26.0
26.7
27.0
29.0
25.5
26.5
25.0
27.2
24.5
23.0
27.0
27.0
24.5
27.5
24.0
26.9
27.3
26.0
27.8
25.8
26.0
28.0
24.0
25.0
26.5
27.0

CaEtured

Collar
No.a

Sex

Date

61A
4362A
52
61B
38A
62B
63
53
39
64
40
55
67
44
65
62C
56
57
45
48
47
54
41
42
51
58
75
38B
49

M
F
F
F
M
M
F
M
M
F
F
M
M
M
F
F
F
F
F
M
F
M
M
F
M
M
F
M
F
F

6-9
6-10
6-11
6-12
6-12
6-13
6-14
6-14
6-14
6-15
6-15
6-16
6-16
6-16
6-16
6-17
6-17
6-17
6-17
6-19
6-19
6-20
6-21
6-21
6-25
6-27
6-27
6-28
6-28
6-29

aFawns 43-52, 62B-63, 55-67, 44-56, 42-38B and 45-48 were twins.

(em)
Hoof
Growth

-

0.37

0.39
0.43

-

-

during 1975 and listed chronologically.

Ear

~

-

-

-

-

-

0.34
0.53

-

0.49

-

-

-

-

-

-

0.59

11.5

-

-

-

-

-

-

0.55
0.50

12.0
11.0

0.44
0.48
0.41
0.49
0.37
0.31
0.36
0.39

11.0
10.8
10.8

-

-

-

11.3
10.3
10.3
11.1

Neck
Circumference (cm)
Max
Min

b Fawns probably 1-2 days of age at capture based on the criteria of ~~ite et al. 1972:904.
more days.

16.8

16.5
16.8
15.0
14.7
18.0
15.3
16.0
19.0
16.9
15.5
15.0
17.6
16.0
13.5
15.0
15.2
15.5
16.5
14.5
18.5
17.5
17.0
20.5
15.0
17.0
15.2
17.5
11.4
16.5
18.0

Fawn 47 may have been a twin of an unco11ared

Remarks

-

16.0
16.8
20.0
17.6
17.0
20.0
18.5
17.0
16.0
19.5
18.0
14.5
17.0
16.5
16.5
17.0
15.0
19.0
19.0
18.0
21.0
16.5
18.0
16.5
18.0
12.7
17.5
19.0

b

b

b
b
b

I

.l&gt;-

o....,
I

b

b

fawn.

The ages of other fawns were likely three or

�;

C)

,.c

s
p:j

M
M

..~~I

;'///t' i I,

I
'.~

j
1
i

I
1

1

Q)

;:l
I-)

(1)

i

(1)

(1)

~

I

\0

::;::

+-I

,.c
s::
o
00

~

s::

I-&lt;

4-;

(1)
(1)

(1)

'0
M

H

o

C)

co I-)

oj §

§

000'\

o

4-;

C)

§"s::

+-I

;:IN

1-&lt;0'\

(1)

4-;M

1-&lt; •.•...•
00'\

o

~

..

C")

!/

J
I'

1:J

i

I
!

b\

�Table 2.

Some temporal and behavorial aspects of capturing 30 mule deer fawns.
Nursing
Periods (min)

Time (hr)

Fawn Behavior
During and
After Release

Other Deer Nearby

Docile, cried
when eartagged.
Struggled,
bleated.

Laid quietly.

None seen.

Not recorded.

With twin 1143

Not recorded.

Not recorded.

Laid quietly.

None. Doe returned
1438, bedded or
wa l.kedaround until
1600 when observers
left.

Attempting to nurse.

Struggled,
bleated.

Laid quietly
Doe disappeared on
for about 5 min, approach.
then jumped up
bleating,tripped,
then laid quietly
10 ft from release.

Released

Last
Sighting

N

X

Min

Max

When First Observed

When Handled

1320

1420

1420

7

6.6

1

32

Attempting to nurse.

1543

1544

1626

1626

None

6-14

0954

1327

1352

6-17

0800

0940

1024

Area

Collar
No.

Capture
Date

First
Observed

Captured

North

43a

6-10

1028

52

6-12

53

65

1024

Not recorded.

3

1

1

2

1

1

I
.pO'

""I

45

6-19

0750

0906

0953

0953

1

1

Nursing.

Lay quietly.

Lay quietly.

2 does, 1 fawn
(twin)

48

6-19

0750

0906

0932

0932

1 1

Nursing.

Struggled,
bleated.

Lay still a
few seconds
then ran off.

2 does, 1 fawn (1145)

54

6-21

0748

0954

1052

Nursing.

Struggled
a lot, bleated
loudly.

Jumped from sack
1 Doe.
and took off 60 m
to top of ridge to
E. Kicked.

51

6-27

1813

1953

2014

Doe feeding in
vicinity for an hr.
wa lke d to fawn.

Extremely
active and
strong, cried
constantly.

Legs collapsed. 2 does. 1 fawn. 2
Ran down slope LInk.
100', fell down,
trotted up steep
slope, fell down ,
trotting H. on
trail.

2

2016

2.5

1

2

3

----------------------------------------------------------------------------------------------------------------------------------------------------------

�Table 2.

Some temporal and behavorial

aspects of capturing 30 mule deer fawns (continued).
Nursing
Periods (min)

Time (hr)
Area
North
(cont)

Released

Last
Sighting

N

X

Min

Max

1957

2010

2010

3

1

&lt;1

1

1937

1954

1954

Collar
No.

Capture
Date

First
Observed

Captured

58

6-27

1853

75

6-28-

1755

1,"1

maybe
more.

49

Southwest

Central

6-29

0900

1055

1120

1120

Not
observed.

Fawn Behavior
During and
After Release

Other Deer Nearby

When First Observed

When Handled

Not recorded.

Struggled
Ran over
hard, bleated
ridgetop to
loudly, panted. N.

Fawn walking about
near doe, followed
doe.

Bleated once,
did not
struggle.
Curled up when
found.

Did not move,
ears up.

2 does

Not recorded.

Doe kept
approaching to
within 30 yds.

Struggled, bleated
loudly

1 buck, 2 does.

Doe ran off,
fawn silent.

Fawn moved off and
could not be
found while we were
removing equipment
from capture site.

1 doe.

1 doe.

38A

6-13

1531

1800

1818

1818

5

4.6

1

12

Nursing.

39

6-15

0800

1155

1226

1231

2

2

2

2

Nursing.

Doe ran off,
fawn struggled
hard near completion of
processing.

Fawn wobbly, walked
up and over W-facing
slope immediately.

1 doe.

40

6-16

1135

1201

1210

1213

Followed doe and
bedded.

Calm and
quiet.

Lay down and got up
and walked quickly
uphill and out of
sight in rain.

None seen.

41

6-21

1125

1839

1857

1859

Nursing.

Doe ran off,
fawn made a
few attempts
to get out of
bag.

1 doe - what
Lay calm, but not
visible due to slope appeared to
exposure, 2 min after be blood in
ano-genital
release.
region.

2

2

1

3

I
.,.
....•

o
I

------------------------------------------------------------------------------------------------------------------------------------------------------------

�Table 2.

Some temporal and behavorial aspects of capturing 30 mule deer fawns.
Nursing
Periods (min)

Time (hr)
Collar
No.

Capture
.Date

First
Observed

Captured

Released

Last
Sighting

42

6-25

0707

1036

1052

1052

38B
(No collar
available)
38B

6-25

0707

1056

1106

1106

6-28

1553

1555

1613

1620

6-9

0653

0835

0921

0925

Area
Central
(contd)

South

61A

N

1

X

Min

Max

8

Fawn Behavior
When Handled

Following doe.

Struggled,
Remained where
then lay still.
placed.
No reaction.
Remained where
placed.

Nursing.

Bedded.

1

Not timed.

Doe alert; fawn
walking around.

62A

6-11

0730

1010

1035

1039

Fawn following doe;
doe was walking.

6lB

6-12

1530

1835

1845

1846

Both walking.

62B

6-14

1845

2010

2017

2017

63

6-14

2008

2034

2040

2040

64

6-15

Not recorded

1155

1205

1205

55

6-16

0729

1030

1045

1055

5

Nursing at 1835.

2

3

1

2

Fawn lying under
Cemo; doe running
away.
Doe grooming fawn;
fawn bedded.
Walking with doe.

1

3

3

DlIr.ingand
After Release

When First Observed

Other Deer Nearby

Some struggle.

Bedded for 2 min
and ran uphill in
direction of departing doe.
Doe ran when
Remained bedded
fawn approached,
at least few
still when
min.
handled.

Fawn bleated
frequently,
struggled, doe
appeared, then
ran away after
capture.
Calm; some
bleating.
Doe ran off
to East.
Quiet, some
kicking.
Doe ran off S,
fawn placid.
Bleated
several times.

Fawn walked away
on release; last
seen going over
hill 60 yds away.

2 does.
2 does.

1 doe, 1
fawn (#42)

At 1530 a doe
was observed
apparently
searching for
fawn near capture sight.
None seen.

Fawn stayed in previous bed.
Very quiet, few
days old?
At release, ran off
downhill quickly.

6 unk.

Remained in bed.

None seen.

Remained in bed.

None seen.

2 unk.
2 unk.

I

-I'-

"I
I-'

�Table 2.

Some temporal and behavorial

aspects of capturing

30 mule deer fawns (continued).
Nursing
Periods ~min2

Time (hr~
Area

South
(contd)

Collar
No.

Capture
Date

67

First
Observed

Captured

6-16

1028

44

6-16

62C

Released

N

X

hln

Max

When First Observed

When Handled

1028

1039

1055

-

-

-

-

Fawn bedded, no doe.

Very quiet
and still.

Remained in bed.

None seen

0739

1343

1348

1350

-

-

-

-

Fawn walking by
itself.

Very quiet.

Stayed in bed.

None seen.

6-17

0713

1005

1010

1011

- - -

-

Walking with doe;
covered about 1/4 mi
walking.

Very quiet,
no bleating.

Ran off upslope
rapidly.

None seen.

56

6-17

1043

1043

1103

1104

- - -

-

Bedded.

Very quiet.

Very still - remained in bed.

None seen.

57

6-17

0800

1145

1203

1204

-

-

-

Bedded with doe; doe
licked fawn frequently.

Very quiet.

Remained

None seen.

Nursing.

Bedded 'vith
doe 40 yds
away.

47

~ursing

Fawn Behavior

Last
Sighting

6-20

0930

1035

1040

1041

2

1.5

1

3

could not be detected with certainty but fawns were in a nursing position.

During and
After Release

in bed.

Other Deer Nearby

I

'""
N

I

Docile - didn't
run or bleat.

None seen.

�Table 3.

Summary of transmitter-collar
Transmitter

Collar
No.a

Receiver
Channel

43
52
53
65
45
48
54
51
58
75
49
38A
39
40
41
42
38B
61A
62A
61B
62B
63
64
55
67
44

9
19
4
1
7
16
11
3
14
22
18
C
B
G

E

D
C
5
6
5
6
18
3
13
15
12

Type
M
C
M
T
C
T
C
T
M

b

unit data and function as used on 30 mule deer fawns, 1975-76.
Data
AEErox. Pulse/Minute
On Fawn
Off Fawn

-

Ambient
Temperature

-

240
2/10
256
188
360
176
336

88

63
66
52

72

52

-

260

-

-

-

C

-

T
T
T
T
T
T
T
M
M
M
M
C
C
C
C
M

260
74
75
61
108
56
98

-

-

196

165
160
240
92
98

-

-

-

-

-

-

-

-

-

-

-

64
75
66
60
77
70

-

-

-

(F)

SummaEY Transmitter Function
Transmitter-Collar Unit
No. Days
Percent of Body
Last Date
Total
Functional
Functional
Wt. at Capture
Wt (g)
260

10.0

266
290
259
290
280
275
266
253
273
178
172
178
175
166
178
269
262
269
262
262
255

7.4
6.2
5.4
7.2
5.2
5.2
6.6
8.4
5.6
4.4
3.9
3.4
5.0
4.4
4.1
6.9
7.5
6.9
5.5
4.6
7.3

-

-

-

286

-

-

-

7.7

10-18-75
9-27-75
7-11-75
6-18-75
8-24-75
6-20-75
1-30-76
7-01-75
4-26-76
8-07-75
7-05-75
6-27-75
7-22-75
7-12-75
7-23-75
7-10-75
8-06-75
6-11-75
6-13-75
8-06-75
6-16-75
10-02-75
7-01-75
9-04-75
9-04-75
10-07-75

-----------------------------------------------------------------------------------------------------------------------------------------------

130
107
27
1
66
1
223
I,

303
40
6
14
37
26
32
15
44
2
2
55
2
110
16
80
80
113

..,..I
-.J

w
I

�Table 3.

Summary of transmitter-collar
Transmitter

Collar
No.
62C
56
57
47

Receiver
Channel

Type

6

M

17

T
T
T

6

14

b

unit data and function as used on 30 mule deer fawns, 1975-76
Data
AEprox. Pulse/Minute
Off Fawn
On Fawn

-

-

120
156

80
96

-

-

Ambient
Temperature

(F)

-

(continued).

Summa~ Transmitter Function
Transmitter-Collar Unit
No. Days
Last Date
Percent of Body
Total
Functional
Functional
Wt. at Capture
Wt (g)
7.62
292
276
255

7.7
7.1
9.2
6.5

7-29-75
8-15-75
6-25-75
7-24-75

42
65
8
34

a

Because transmitter-collar units were not delivered on schedule, units permanently lettered 38, 61, and 62 were removed from dead fawns
and placed on newly-captured fawns , These f awns are recorded on project forms as 38A, 38B, 6lA, 61B, and 62A, 62B and 62C in chronological
sequence of capture. Fawns 38A, 39, 40, 41, 42 and 38B we re fitted 'vith low frequency transmitters.
b
M
T
T

C

Hg switch, movement increases pulse rate.
Temperature; collars 38 to 42, low to high temperature decreases pulse rate.
Temperature; all other T collars, low to high temperature increases pulse rate.
Pulse rate is relatively stable.

I
.p.

'"

+I

�Table 4. Blood values from 10 radio-collared
aspirated from the juglar.
Variables
Date of Capture
Date of Blood Aspiration
Body Wt (kg) at First Capture

Erythrocytic

mule deer fawns.

Fawns were recaptured two to thirteen days after collaring.

Blood

47
M
6-20
6-22
3.9

52
F
6-12
6-25
3.1

53
M
6-14
6-25
3.6

Fawn No. and Sex
55
M
56
F
57
F
6-16
6-17
6-17
6-22
6-22
6-22
3.3
4.1
3.0

61B
M
6-12
6-22
3.9

63
M
6-14
6-22
5.7

64
F
6-15
6-22
3.5

67
M
6-16
6-22
2.3

3.52
23
7.2

37
12.2

4.97
29
9.6

4.98
29
9.2

39
13.4

41
14.0

31
10.2

4.84
24
7.6

Moderate
Moderate

Slight
Slight

Marked
Marked

Series

Erythrocyte (106/mm3)
Packed cell vol (%)
Hemoglobin (g/lOO m1)
Nucleated erythrocyte (106/mm3)
Polychromasia
Anisocytosis
Poikilocytosis

4

2

Slight
Marked

30
10.2

4.88
23
7.8
2

2

Marked
Moderate Moderate Moderate Moderate
Marked
Marked
Moderate Moderate Moderate
Moderate Marked
Moderate

I

...,
"'"
\.n

Leukocytic Series (Per mm3)

I

Leukocyte (103)
Segmented neutrophil (%)
Band neutrophil (%)
Lymphocyte (%)
Monocyte (%)
Eosinophil (%)
Basophil (%)
Platelets

2.2
88

27

5.6
88
1
10

4
1

6
2
1

o
o

o

Possibly low

Normal

Normal

High

104

88
6.75
142
240

88

210

o
12

o
o
o

5.3
62
3
27
3

4.5
64

o

12.4
71

3.7
81

5.5
73

6.8
58

7.4
58

3.2
80

1

1

o

o

o

1

22

16

27

29

42

18

2
4

1
1

1

12

o

o
o
o

o

o
o
o

o
o

Normal

Normal

Normal

Normal

Normal

Normal

100

108

104

90

104

101

95

285

178

250

270

240

305

135

1

1

Chemistry
Chloride
Potassium
Sodium
Alkaline phosphotase

105

�Table 4.
aspirated

Blood values from 10 radio-collared
from the juglar (continued).

Variables
Date of Capture
Date of Blood Aspiration
Body Wt (kg) at First. Capture

Chemist~

52
F
6-12
6-25
3.1

Fawns were recaptured

M
53
6-14
6-25
3.6

two to thirteen days after collaring.

Fawn No. and Sex
F
51
M
56
F
55
6-17
6-17
6-16
6-22
6-22
6-22
4.1
3.0
3.3

F
64
6-15
6-22
3.5

M
6lB
6-12
6-22
3.9

M
63
6-14
6-22
5.7

64
12.0

92
11.0

9.5

-

-

-

-

-

-

5.1
200

6.4
"-

6.3
200

5.5

Blood

-67

M
6-16
6-22
2.3

(continued)

CPK
Phosphorus
LDH
Calcium
Total serum protein
Fibrinogen
Remarks

47
M
6-20
6-22
3.9

mule deer fawns.

35

-

5.5

EDTA
slide
read
one
day
after
aspiration.

89
12.0
300

40
15.0

42
11.0
390

48

-

-

5.2
300

5.7
400

6.7
500

Small
reactive
lymphs,
some
with
densely
clumped
chromatin.

Probably
One
born on
giant
6-17.
platelet,
large
amount
of basophilic
stippling.

-

-

5.6
200
Marked
basophilic
stippling
in
RBC.

-

?

164
9.5
300

-

109
9.5
370

-

5.1
400
I

...,
""
aI

�-477-

Table 5. The presence of bacteria in the nasal swabs of 20 mule deer fawns
at the time of capture June, 1975.

.

I

r

.

(J)

£:1

.

~i'

,,-1

.

Co.
(:":

:-.

-

o,

.

P,

c:
E

(1.

.
.

~;

o,

&lt;lJ

[f;

.•..

'

U

.

CC

0
U

~

.
m

38

+

+

+

+

39

+

40

+

+

+

+

42

+

+

+

+

+

+

+

44

+

+

+

+

+

+

+

45

+

+

+

+

+

+

47

+

+

.+

+

a

+

+

l~8

+

u:

()

.D

~.

E

4-1

U

0:-1
.-l

c;j

(J)

h
0

r-'

.,-1

;J
Z

(J)

+

c;j

,D
cJ

co
0
r"
r-!

~
H

C;
h

0
U

Q)

;.t""

o.
(J)

0.

(J)

01

~I

u:

+

+

4-

+

+

+

+

+

+

+

+

+

54

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

.......

58

--_. 1--'
E,l

I +

--;1-

I

+

+

+

EI
c;j

~I ~I

I

.:-

+

+

- 1,,_-

__,

a Not radio-collared,

..

0

E

0
H

Q)

~~
p.,

&lt;iJ

«:

+

+

.I

+
+

+
++

I

+

+
-l-

+

I±
_;I_L_-1 -_.+

+

+

+

+

+

I +

I
--I':-=- ---,--- --~-j
r--·-r~~r---··
+

&gt;

+

I

Co'!..

~

E
0
-o
::J

+

,'-;:-:-.(--r-----·.--,,-------------I
61R

")

co

0

+

+

57

('"0

+

52

--

.

o,
(J)

Co

~I

f-.

55

.

o.

-i

I+ I

I'r

+

I

captured when transmitters were not available.

I

�Table 6.

Collar
No.

The history of 30 mule deer fawns equipped with radio-collars
Dates and Times (hr)
Last
Captured
Seen Alive

and located with telemetry on a near-daily- basis, 1975-76.

Circumstances
Found Dead

Field Exam.

Pathologists

of Death
Diagnosis

Probable
Cause of Death

Remarks

Habitat

43

6-10-75, 1320

2-.06-76, 1240

52

6-12-75, 1544

12-27-75, 1012

5-11-76, 1010

Scavanged
carcass

None

Unknown

Riparian

Portion hide lower jaw, dried
stomach contents.

53

6-14-75, l327

7-10-75, 1027

7-11-75, 0830

Recent
predation

Multiple lacerations

Predation, predator not
identified

Open pine

Orbital bones crushed,
puncture wounds.

65

6-17-75, 0940

5-03-76, 0920

Open pine

With 13 does, 9 fawns.

45

6-19-75, 0906

8-19-75, 1015

Transmitter

8-24-75, l334

Unknown
cause(s)

Massive subcutaneous
hemorrhage - bite
wounds

non functional.

Predation, presume Irrigation Only the head, neck and front
coyote
meadowlegs remained.
dense pine
ecotone

48

6-19-75, 0906

8-18-75, 0924

4-12-76, 1140

Scavanged

None

Unknown

-

54

6-21-75, 0954

1-26-75, 1045

1-30-75, 1030

Scavanged
carcass

Not examined

Unknown

Shrub

51

6-27-75, 1953

6-30-75, 1410

7-01-75, 1334

Intact
carcass

Starvation, dehydration

58

6-27-75, 1957

5-18-76, 1410

5-11-76, 1055

Scavanged
carcass

None

Predation - unknown agent

Riparian

Head, spine, portion of rib
cage, hide still intact.

75

6-28-75, 1937

8-06-75, 0834

8-07-75, 0845

Unknown
cause(s)

Acute viral and
parasitic enteritis
and necrotizing
cecitis and
colitis.

Acute enteritis,
cecitis, and
colitis disease
complex

Irrigated
meadow

Weighed 8.4 kg at death. Thus,
gained min of 0.12 kg per day.

Starvation, perhaps Open pine
abandoned by doe

Last transmitter
1439, 6-20-75.

signal heard

Noted limping on 12-28-75.

GI tract devoid of ingesta.

I
::-

"
Cf'

�Table 6.

The history of 30 mule deer fawns equipped with radio~collars

Collar
No.

Dates and Times (hr)
Last
Seen Alive
Captured

1.9

6-29-75, 1055

18A

39

and located with telemetry on a near-daily basis, 1975-76 (continued).

Circumstances

of Death

Found Dead

Field Exam.

Pathologists

7-02-75, 09·50

7-05-75, 0830

Starvation

Probable starvation,
dehydration

6-13-75, 1800

6-13-75, 1818

Collar
1635
only found
6-27-75 .

6-15-75, 1155

7-14-75, 0905

2-22-75, 1812

Scavanged
carcass in
advanced
autolysis

Diagnosis

Unknown

Probable
Cause of Death

Starvation, perhaps abandoned
by doe

Predation, predator not
identified

Habitat

Remarks

Open pine

No milk in rumen or abomasum.

Open pine

No evidence of fawn in
vicinity of collar.

Open pine

Intact skull with deep bites,
ribs and forelegs remaining.

J,..

40

6-16-75, 1201

7-11-75, 1300

7-12-75, 0800

Predation

Multiple traumatic
lesions about head
and neck

Predation, presume coyote

Riparian

Buried under debris and tree
limbs.

41

6-21-75, 1839

7-22-75, 1709

7-23-75, 0854

Recent
predation

Multiple traumatic
lacerations

Predation
(tentative)

Shrub

An acute disease, process
cannot be excluded.

42

6-25-75, 1036

7-08-75, 0944

7-10-75, 1413

Recent
predation

Recent presumed
predation

Predation, presume Felis spp .

Shrub

Unidentified
carcass.

38B

6-28-75, 1555

8-09-75, 0722

8-11-75, 0636

Recent
predation

Multiple trauma
(predator)

Predation, presume coyote

Dry wash

Doe protected carcass from
5 coyotes.

61A

6-09-75, 0835

6-10-75, 1132

6-11-75, 1136

Not predation

Starvation, dehydration

Starvation, dehydration, perhaps abandoned
by doe

Shrub

No food material in any portion of GI tract.

62A

6-11-75, 1010

6-12-75, 1130

6-13-75, 1140

Presumed
abandoned

Starvation, dehydration

Starvation, dehydration, perhaps abandoned
by doe

Shrub-dry
wash
ecotone

No food material in any portion of GI tract.

cat seen near

--------------------~--------------------------------------------------------------------------------------------------------------------------------~----

...•
\0
I

�Table 6.

The history of 30 mule deer fawns equipped with radio-collars

Collar
No.

Dates and Times (hr)
Last
Captured
Seen Alive

61B

6-12-75, 1845

62B

and located with telemetry on a near-daily basis, 1975-76 (continued).

Circumstances

of Death

Found Dead

Field Exam.

Pathologists

8-05-75, 0757

8-06-75, 0815

Intact carcass with
wounds

Multiple trauma, predator kill

6-14-75, 2010

6-15-75, 1030

6-16-75, 1500

Recent predation by
eagle

Not examined

63

6-14-75, 2034

1-20-76, 0935

Collar
1645
only found
5-10-75

64

6-15-75, 1155

6-30-75, 1040

7-01-75, 1045

55

6-16-75, 1030

9-04-75, 0822

9-09-75, 1250

67

6-16-75, 1028

9-04-75, 1055

44

6-16-75, 1343

10-07-75, 0744

No further
contact
10-30-75, -

Diagnosis

Probable
Cause of Death

Habitat

Remarks

Pred~tion, coyote
or lion

Shrub

Approached within one meter
on 8-5. Possibly sick.

Predation, presume golden
eagle

Shrub

All viscera, except one kidney
and left shoulder eaten.

Shrub

Transmitter non functional.

Recent predation

Multiple trauma, predation

Predation, presume Felis spp.

Dry wash

Carcass under rock covered
with debris.

I
~
ce

Well scavanged, predation unlikely .

Not examined

Unknown, dead
about 48 hr

Shrub

Located with aerial telemetry.

I

Extreme decay

Not seen

Unknown

Shrub

Found by hunter

"Presumed to be an"
undiscovered
disease process

Undiagnosed
disease process

Shrub

Abomasum contained much
ingesta, including milk curd.

62C

6-17-75, 1005

7-28-75, 0951

7-29-75, 1025

Unknown
cause(s)

56

6-17-75, 1043

8-23-75, 0822

Collar only
found 5-7-76

57

6-17-75, 1145

6-24-75, 0910

6-25-75, 1350

Recent injury

Traumatic injury to
vital organs

47

6-20-75, 1035

7-23-75, 0806

7-24-75, 0829

Recent predation

Not examined

Shrub
Injured by either
Shrub
a falling rock or
caught under a rock
Predation, presume Open pine
coyote(s)

Transmitter non functional.
Released from rock on 6-23;
seemed weak.
Only head, neck, and front
leg remaining.

o

�-481-

Table 7. The history of survival of 30 mule deer fawns captured,
radiocollared and tracked on a near-daily basis during 1975 and 1976.

Area
North

Southwest

Southeast

Fawn
No.

Day of Capture to
Last Day Seen Alive

43
52
53
65
45
48
54
51
58
75
49

241
199
26
320
61
60
219
3
315
39
3
X = 135.1 SD = 124.9

38A
39
40
41
42
38B
X = 23.3

0
29
25
31
13
42
SD = 14.8

X = 53.5

1
1
54
1
200
15
80
80
114
41
68
7
33
SD = 57.2

61A
62A
61B
62B
63
64
55
67
44
62C
56
57
47

Number of Days
Day of Capture to
Day Found Dead

Day to Last Seen Alive
to Day Found Dead

333
27

X=

134
1

66
297
223
4
318
40
6
146.0 SD = 143.5

X = 30.8

X = 36.1

37
26
32
15
44
Sf)

11.0

5
237
4
1
3
1
3
X = 43.2

X = 2.8

SD

8
1
1
2
2
SD = 3.0

2
2
55
2

1
1
1
1

16
85

1
5

115
42

23
1

8
34
SD = 39.0

1
1
SD = 6.9

X = 3.6

X = 77.4

X = 76.2

X = 18.3

SD = 94.3

SD = 104.1

SD = 170.4

84.7

�-482-

The circumstances of death of those fawns radio collared in 1975 are detailed,
as noted.previously, in Table 6. A general summary through May 28,1976 of
the history of fawns radio collared during 1974 and 1975 is given in Table 8.
Insofar as assessing mortality rates are concerned, these data are ambiguous
because of lost radiocollars and fawns which simply disappeared.
If the
number of fawns presumed alive are divided by the total documented deaths
the minimum percentage of fawns surviving would approximate 8 percent over 11
months during 1975-76 and 16 percent for 19 months (last sighting of 1974
fawn, January 1976) during 1974-75.
I hypothesized that survival of fawns might be link~d to activity level;
i. e., high levels of activity would increase the probability of predation or
accidental death. The index employed was the distance of each relocation
site for each fawn from its capture site. The 1,600 relocations are described
statistically by month in Table 9, and from capture to March 31, 1976 in
Table 10. Analyses to test this hypothesis will be presented in subsequent
reports. Of interest, however, are the grossly different mean distances
traveled by twins during one month. For example, during September 1975 fawn
number 43 had a mean distance of 2,158 ft, while fawn number 52 had a mean
distance of 865 ft (Table 9). Also of interest are the apparent distances
traveled by some very young fawns. Thus, although fawn number 39 was relocated only 8 times during its one approximate month of life, these distances
averaged 13,662.5 ft (Table 10) and inspection of the raw data showed that
she was about 15,550 ft from her capture site on both July 9 and July 22.
The percentages and ratios of radio collared fawns to unmarked fawns are
related to the search effort expended by month from June 1975 through March
1976 in Table 11. Both marked and unmarked fawns recorded per person hour
increased from June through September, then decreased through November and
fluctuated from December through March. As noted previously (Anderson 1975),
fawns recorded per person hour increased from June through September when
an actual decrease in absolute numbers of fawns must have been taking place.
This illustrates the very cryptic nature of mule deer fawns and the difficulty of assessing the status of fawns during the first three to four months
of life.
In Table 12, monthly tabulations of 14,430 deer observed in 7,666 person
hours show that more deer per person hour can be seen during September than
any other month, fawns were more readily seen than bucks and does were more
readily seen than fawns, and bucks were seldom seen except during October
and November. A comparison of the 1974-75 and 1975-76 indices of mule deer
recorded per person hour (Table 13) shows that even though several times more
person hours were expended during each month except September during 1975-76
more deer per person hour were seen during each month of 1974-75 except July.
These differences are believed to reflect the restricted area traveled by
most people during 1975-76. The much greater number of bucks seen per person
hour during 1974-75 may reflect an actual difference in the buck population
since the 1974-75 and 1975-76 indexes differ by a factor of 3 to 8.
Coyotes
Nine coyotes were caught by contract trapper George E. Stewart from October
1975 to January 1976 (Table 14). Some of their physical characteristics are

�-483-

given in Table 15. Trap injuries and the effects of drug immobilization
on each coyote are described in Table 16. Apparently only one coyote (no.
39) was released without broken skin, bones, or tendons. The beneficial
effects of the tranquilizer in preventing injury appears to be minimal at
best. Vertebrates other than coyotes captured, killed, escaped or released
are listed in Table 17. A two year comparison of abundance indices of cottontail rabbits (Sylvilagus sp.), a presumably important food of coyotes, is
made on a monthly basis in Table l7A.
Relevant details o.n capture sites, ear tags, and transmitters fitted to 9
coyotes are listed in Table 18. A comparison with the channels employed for
coyotes with those for fawns in Table 3 shows inadvertent duplicaton by
Davtron of the following channels: channel 11, fawn 54, coyote 40; channel 4,
fawn 53, coyote 41; channel 14, fawn 58, coyote 42; channel 18, fawn 49,
fawn 63, coyote 43; channelS, fawn 6lA, coyote 44; channel 9, fawn 43, coyote
45; channel 16, fawn 48, coyote 47. Some confusion may have resulted from the
duplications of channel 14 and channel 9 during the winter. However, by the
time the coyotes were radio collared most of the fawns with duplicate channels
had died.
Statistical descriptions of the distances coyotes traveled from their capture
to relocation sites are shown on a monthly basis for individual coyotes in
Table 19, and for all coyotes in Table 20. These values, based on 385
relocations may reflect either actual coyote activity or a stereotyped
pattern of radio tracking a widely ranging animal on a limited, familiar
area. For the four months involved, coyote travel seems quite limited, with
little change over time.
This impression of limited travel by the coyotes under study belies the data
for three coyotes in Table 21. The information therein also suggests a high
rate of mortality among radio collared coyotes since it is likely that others
had died and were not found and reported. Only two of the seven dead
coyotes were found by project personnel.
DISCUSSION
One aspect of telemetry that we wished to evaluate was the relative efficiency of high and low frequency equipment in radio tracking fawns in mountainous habitat. Because the low frequency receivers were well used, often
malfunctioned, and the high and low frequency transmitters differed in voltage, this evaluation could not be made.
The Davtron equipment was generally satisfactory, however, and once a signal
was received the fawn could usually be located in less than one-half hour.
The distances of signal reception ranged from less than 100 yards to over
three miles under ideal line-of-sight conditions from elevated sites. In
most topography, signals were commonly received at distances of about onequarter to three quarters of a mile. A high ridge or even small ridgetops
usually eliminated an effective signal regardless of range. The transmitters
indicating fawn movement were often helpful in tracking fawns in dense cover,
but those indicating a change in external body temperature were seldom helpful. This was probably due to the sensor not in contact with the animal.
Fabricating a device to constantly hold the sensor against the fawn neck from
an expandable collar proved to be a most intractable problem.

�-484-

Table 8. The history of 53 radio-collared mule deer fawns captured during
June, 1974 and 1975. Based on field work through May 28, 1976.
1975

1974

Total

4

Presumed alive

2

Disappe ared

1

5

6

Radio collar only found on ground

3

4

7

5

7

12

Documented deaths
Cause unknown
Probable predation
d

4

4

Presume golden eagle

1

1

Presume Felis spp.d

2

2

Predator unknownd

5

1

6

4

4

8

Presume coyote

b

Probable abandonment by doe '

c

Probable diseasec

2

2

Accidental injuryC

1

1

Total

30

23

53

a A male and female seen infrequently but always within one-half mile of
their capture sites. Not seen since February 1976.
b

Starvation and dehydration were the ultimate causes of death.

c Based on laboratory autopsies of 19 fawns.
d Based on field observations and laboratory autopsies.

�Table 9. The approximate distances (ft)a traveled by 29 radio-collared mule deer fawns from their capture sites to relocation sites,
10 June 1975 to 31 March 1976. An additional fawn (No. 38A) was never relocated.
Fawnb

43

Statistic

N
X

SD
Min
Max
52

N

X
SD
Min
Max
45

N
X

SD
Min
Max
48

49

N

June

July

August

September

October

November

December

January

10
1,485
833
600
2,800

31
1,782
791
400
3,400

30
1,960
818
300
3,200

30
2,158
635
600
3,200

15
2,030
776
700
3,900

-

6
1,408
592
600
2,100

5
1,510
976
800
3,100

15
2,653
1,047
950
4,200

30
1,683
1,098
200
3,300

30
1,055
902
100
3,350

30
865
748
0
2,600

14
1,346
1,226
250
4,800

31
894
800
4,600

23
2,133
1,246
800
4,700
5
3,240
1,026
1,800
4,500

11
2,636
1,246
1,600
5,450

1,947

X

2
3,750

SD
Min
Max

2,500
5,000

11
2,091
971
800
4,200

2
1,100
141
1,000
1,200

5
1,860
860
1,000
2,950

N
X

SD
Min
Max

1,768

-

1

-

-

450
450

February

March

1
1,700
1,700

5
2,150
1,598
400
3,400

---------------------------------------------------------------------------------------------------------------------------------------------

I

"'"
cc

VI
I

�Table 9. The approximate distances (ft)a traveled by 29 radio-collared mule deer fawns from their capture sites to relocation
10 June 1975 to 31 March 1976. An additional fawn (No. 38A) was never relocated (continued).
Fawnb

51

53

December

January

30
1,915
933
400
3,700

29
1,445
914
200
3,750

29
1,326
740
400
3,850

23
1,065
463
350
2,000

25
1,632
974
200
4,300

25
1,658
490
300
3,050

25
2,034
770
600
3,400

2
1,000
778
450
1,550

30
1,782
1,139
200
4,400

26
1,165
562
200
200

27
1,322
761
250
2,500

23
1,374
706
350
2,750

24
1,562
905
400
4,100

26
1,833
682
600
3,200

29
2,505
898
1,000
4,350

27
2,237
1,449
450
5,300

27
2,561
1,605
350
5,300

2
2,400
1,131
1,600
3,200

-

2
1,250
919
600
1,900

3
1,633
153
1,500
1,800

6
1,542
856
600
2,700

12
1,004
343
650
1,700

21
1,350
549
400
2,500

24
1,519
835
0
3,200

20
1,250
677
0
2,400

15
2,183
1,247
650
4,050

3
700
173
5'00
800

1
·800
800

14
1,514
1,053
150
3,400

11
1,514
460
550
2,200

10
2,645
1,045
700
3,500

N

N
X

SD
Min
Max
N
X

SD
Min
Max
65

November

N

SD
Min
Max

58

October

July

X

54

September

June

SD
Min
Max

N
j{

SD
Min
Max

March

February

August

Statistic

X

sites,

."..I

00

'"

I

---------------------------------------------------------------------------------------------------------------------------------------------

�Table 9. The approximate distances (ft)a traveled by 29 radio-collared mule deer fawns from their capture sites to relocation
10 June 1975 to 31 March 1976. An additional fawn (No. 38A) was never relocated
(continued).
Fawnb

38B

Statistic

June

N

X
SD
Min
Max

~

39

40

N

X
SD
Min
Max

o

1,400

August

23
6,015
1,862
200
7,200

7,325
402
6,700
7,800

N

1

7

11,700

SD
Min
Max

11,700
11,700

13,943
136
11,550
15,550

o

N

4

X

1,438
25
1,400
1,450

N

3

X

900
557
300
1,400

SD
Min
Max

September

October

November

December

January

February

March

6

6
4,892
2,648
1,400
7,600

X

SD
Min
Max
41

2

700
1,990

July

sites,

8
2,4l3
1,191
500
4,000
12
1,783
1,148
700
4,150

---------------------------------------------------------------------------------------------------------------------------------------------

I
~
00

-..J
I

�Table 9. The approximate distances (ft)a traveled by 29 radio-collared mule deer fawns from their capture sites to relocation sites,
10 June 1975 to 31 March 1976. An additional fawn (No. 38A) was never relocated (continued).
Fawnb

44

July

August

SD
Min
Hax

13
1,954
869
0
3,150

30
2,138
718
600
4,100

17
3,350
2,019
400
3,60d

N
if
SD
Min
Hax

13
1,942
906
0
3,200

30
2,253
707
600
4,200

22
2,307
682
450
3,050

N

10
3,505
3,519
0
11,500

24
8,623
1,196
6,300
12,150

13
2,261
1,419
600
4,850

31
1,543
902
500
4,400

31
1,497
633
300
2,550

5
2,710
1,370
1,000
4,500

13
2,485
1,334
1,000
5,000

31
1,589
898
500
4,450

31
1,393
684
100
2,550

4
2,762
1,617
1,000
4,600

N

ir

2Q.

47

if
SD
Min
Max
55

N

if
SD
Min
Hax

E

September

June

Statistic

N

if
SD
Min
Hax

October

November

December

January

February

March

..,..I
00
00

I

---------------------------------------------------------------------------------------------------------------------------------------------

�Table 9. The approximate distances (ft)a traveled by 29 radio-collared mule deer fawns from their capture sites to relocation
10 June 1975 to 31 March 1976. An additional fawn (No. 38A) was never relocated (continued).
Fawnb

57

Statistic

N

X
SD
Min
Max
61A

N

X
SD
Min
Max
61B

N

X"
SD
Min
Max
62A

N

X
SD
Min
Max

.ill

N

X
SD
Min
Max

June

July

August

September

October

November

December

January

February

sites,

March

8
1,212
752

o
2,600
3
133
231

o
400
16
1,456
867

o
3,050

30
2,543
2,588
300
10,400

6

1,700
708
700
2,650

3
100
132

o
250
2

900
990
200
1,600

-----------------------------------------------------------------------------------------------------------------------------------------------

.,.I
ex&gt;

'"

I

�Table 9. The approximate distances (ft)a traveled by 29 radio-collared mule deer fawns from their capture sites to relocation sites,
10 June 1975 to 31 March 1976. An additional fawn (No. 38A) was never relocated (continued).
Fawnb

Statistic

June

July

August

September

October

November

December

January

February

March

------

&amp;l

N

x
SD
Min
Hax

62C

N

X
SD
Hin
Hax
64

N

X
SD
Min
Max

a Based on measurements
each day.

16
2,769
1,312
100
4,400

31
3,198
1,092
1,200
5,650

13
1,319
1,029
0
3,800

29
1,912
1,167
300
4,200

13
6,762
4,538
0
17,500

31
2,935
1,468
800
5,550

27
3,170
1,231
600
5,000

1

I
.0-

-

'"'0

-

I

9,400
9,400

made on U.S. G.S. Quadrangles

b Underlined adjacent fawn numbers indicate twins.

(scale 1:24,000), contour interval -40 ft.

An attempt was made to re1oc.ate each fawn

�-491-

Table 10. The distances (ft) mule deer fawns moved from their capture sites
9 June 1975 to 31 March 1976 based on relocations with radio telemetry.
Eawn No. and
Dates of Relocations

No.
Relocations

Mean

SD

Min

43
6/15/75-2/6/76

124

1914.9

774.0

300

3900

45
6/20/75-8/23/75

65

2129.2

1l00.7

800

5450

48
6/20/75-8/18/75

18

2594.4

1195.9

800

5000

49
6/29/75-7/5/75

7

1642.9

796.0

1000

2950

51
6/27/75-7/1/75

4

725.0

150.0

500

800

52
6/15/75-12/27/75

125

1423.6

1142.0

100

4800

53
6/17/75-7/11/75

25

1514.0

830.2

150

3400

54
6/21/75-1/ 30/76

196

1642.4

874.1

200

4300

58
6/28/75-3/31/76

241

1827.6

1127.8

200

530

65
6/18/75-3/3/76

102

1495.1

848.1

0

4300

75
6/30/75-8/7 /75

35

4861. 4

1465.7

1000

6650

38B
7/1/75-8/9/75

29

6286.2

1744.9

200

8900

39
6/27/75-7/22/75

8

13662.5

1492.3

11550

15550

40
6/18/75-7/12/75

12

2087.5

1064.5

500

4000

Max

North Area

Southwest Area

------------------------------------------------------------------------------

�-492-

Table 10. The distances (ft) mule deer fawns moved from their capture sites
9 June 1975 to 31 March 1976 based on relocations with radio telemetry
(continued) .

--_._------------------Fawn No.
Date of Relocations

No.
Relocations

Mean

SD

Min

41
6/23/75-7/23/75

IS

1606.7

1101. 8

300

4150

42
6/25/75-7/10/75

8

3843.8

2985.5

o

7600

44
6/17/75-10/7/75

94

2402.1

1368.15

o

9400

47

34

7117.6

3158.82

o

12150

80

1715.0

1001. 97

300

4850

65

2193.8

746.53

o

4200

57
6/ 17 /75-6/25/75

8

1212.5

752.50

o

2600

6LA

2

200.0

282.89

o

400

52

2111.5

2084.01

o

10400

3

100.0

l32.29

o

250

62B
6/14/75-6/15/75

2

900.0

989.95

200

1600

62e

42

1728.5

ll47.75

o

4200

111

3183.3

1499.47

100

9900

64
6/15/75-7/1/75

14

6950.0

4417.40

o

17500

67
6/17/75 --9/4/75

79

1719.0

1032.93

100

5000

Max

Southwest Area (cont.)

Southeast Area

6/20/75-7/24/75
5S

6/17/75-9/9/75
56

6;17175-8/23/75

6/9:

75-6/11/75

61.3
6':'0/7'5-8,6/15
62A

6/11/75-6/13/75

6/17/75-7/29/75
63

6/14/75-

------_._----------

�Table 11. Percentages and ratios of marked (radio-collared)
expended by month within the study area, 1975-76.

Year
1975

Marked
No.
Percent

Unmarked
No.
Percent

June

107

32.4

223

July

352

46.3

August

286

September
October

Month

to unmarked mule deer fawns recorded and the person hours of search effort

Ratio-Marked

Fawns Recorded
Marked Per
Unmarked Per
Person Hour
Person Hour

l:

to 11nmarked

Total Person
Hours

67.6

330

.480

2,756

.039

.081

466

53.7

813

.755

1,406

.250

.331

39.8

433

60.2

719

.661

858

.333

.505

167

30.8

376

69.2

543

.444

315

.530

1.194

90

23.6

292

76.4

382

.308

361

.249

.809

273

.230

445

.115

.499

400

.246

469

.168

.684

November

51

18.7

222

81. 3

December

79

19.8

321

80.2

I
.J&gt;-

-o

w
I

1976

January

78

20.1

310

79.9

388

.252

429

.182

.722

February

47

14.9

269

85.1

316

.175

336

.140

.801

March

42

13.3

274

86.7

316

.153

291

.144

.941

�Table 12.

Year
1975a

Total numbers and indices of mule deer recorded per person hour by sex and age category within the study area, 1975-76.

Month

M

Total No. Mule Deer Recorded
Fawn
UncI.
F

l:

Total Person
Hours

M

F

Deer Per Person Hour
Uncl.
Fawn

l:

June

109'

1,277

330

1,543

3 ,259

2,756

.040

.463

.052

.569

1.188

July

68

810

818

161

1-.857 _

1,406

.048

.576

.331

.115

1.321

August

19

656

719

147

1,541

858

.022

.765

.505

.171

1. 796

September

24

627

543

62

1.256

315

.076

1.990

1.194

.197

3.987

October

40

485

382

68

975

361

.111

1.343

1.058

.188

2.700

November

38

478

273

447

1,236

445

.086

1.074

.613

1.004

2.777

December

25

414

400

373

1,212

469

.053

.883

.853

.795

2.584

-

-

-

.967

2.655

-

1.155

2.821

-

1.351

3.460

.044

.732

.585

.521

1.882

I
.I&gt;\0

1976b

l:

January

8

328

388

415

1,139

429

February

3

241

316

388

948

336

March

0

298

316

393

1,007

291

334

5,614

4,485

3,997

14,430

7,666

a Deer recorded on the randomly-selected,

X

1/4-mi1e2 quadrats (see 1976 Progress Rept., Work Plan 19, Job 2) are excluded.

b Antler loss and fffivngrowth precluded accurate identification of bucks, does, and fawns during January, February and March.
ratios, however, include the deer tentatively assigned to the three categories during those months.

The mean

.I&gt;I

�Table l3.

A comparison

of the 1974-75 and 1975-76 indices of mule deer recorded per person hour within the study area.
Male

Female

Total Deer

Fawn

Total Person Hours

Month

1974-75

1975-76

1974-75

1975-76

1974-75

1975-76

1974-75

1975-76

1974-75

1975-76

June

.l32

.040'

1.019

.463

.162

.052

2;027

1.188

593

2,756

July

.101

.048

.642

.576

.236

.331

1.114

1.321

288

1,406

August

.195

.022

1.234

.765

.532

.505

2.216

1.796

231

858

September

.485

.076

2.576

1.190

1.809

1.194

5.700

3.987

340

315

October

.354

.111

2.836

1.343

2.175

1.058

6.429

2.700

189

361

November

.333

.086

2.942

1.074

1.667

.613

5.717

2.777

120

445

.348

.053

3.485

.883

2.182

.853

6.212

2.584

66

469

.,.
I

December

'"I

IJ&gt;

�Table 14. The coyote trapping effort by month and individual coyotes caught within Sections 3, 4, 17, 22, 28, T11N, R70W and Section
34, T12N, R70W, 9 October 1975 to 25 January 1976.
No. Days
Traps Were Set

No. TrapsaSet
Per Day

Total
Trap Days

Total
Coyotes Captured

October
November
December
January ..

22
31
31
25

8-18
5-17
15-20
12-26

348
379
748
452

2
4
3
0

39
40
41
42

- 114
257
477
520

E

109

5-26

1,927

9

43
44
45
46
47

587
650
996
1,050
1,217

Month

Cumulative TraE Days Per CaEture
No. Days
Coyote No.

aTrap numbers were changed according to relative numbers of trap visitations by coyotes, disturbance by other animals, discovery of
more desirable trap sites or apparent trap malfunction.
I
-I&gt;-

'"a-I
Table 15. Some physical characteristics of four male and five female coyotes captured with steel traps, radio-collared,
and released on the study area, 9 October to 27 December, 1975.
Ear
Tag No.

Sex

Date
Captured

Body
Wt. (kg)

Hind Foot

39
40
41
42
43
44
45
46
47

M
M
F
F*
F
F*
M*
F*
M*

10-9
11-17
11-14
11-17
11-21
11-25
12-14
12-17
12-27

9.7
9.3
9.3
10.4
9.5
10.2
13.4
8.6
11.3

19.4
19.0
19.0
19.0
18.5
19.0
19.5
18.5
20.5

*Coyotes more than one year of age.
criteria of Gier (1957:50, 51, 55).

Lengths (cm)
Tail
Ear
33.9
40.0
31.5
39.0
29.0
38.0
39.0
35.0
32.0

11.4
11.5
10.0
13.5
11.5
11.5
11.0
11.0
11.0

ear-tagged

Total Body

Chest

Circumference (cm)
Min Neck

Max Neck

112.0
123.0
114.5
114.0
107.0
114.0
124.0
115.0
114.0

47.1
45.0
44.0
45.0
45.0
49.0
53.0
52.0
54.0

27.0
24.0
24.5
26.0
24.5
28.0
33.0
31.5
31.0

28.0
27.0
28.0
28.5
27.0
30.0
35.0
33.0
33.0

All others estimated to have been born during 1975 according to the growth curves and dentition

�Table 16. Description of steel trap injuries and drug immobilization to 9 coyotes, 9 October to 27 December 1975.
Montgomery No.4 w Ith padded, offset jaws and anchored by drag hooks attached to an 8 ft chain.
Coyote
No.

Tranquilizer Taba
Usage and Effects

Foot Caught and
Location of Trap Jaws

Traps were

Description of Damage

0
Left front, 1/2 way through pad, 45
angle.

No bleeding, no swelling, nc broken
skin, no broken bones.

Definitely tranquilized but ran
15 meters on approach.

Left front, above pad - upper side.

Lots of swelling, broken skin.

41

Chewed up but did not ingest
tab, docile.

Left front, above first joint.

Broken skin, tendon cut, swo l.l.en ,

42

Could not find tab, no apparent
tranquilizing effect.

Left front, above pad.

Swelling, cut skin on top, no broken
bones.

43

Ch ewed , but did not inges t drug,

Left front, by three toes on the
outside.

Very little swelling, minimal bleeding
which stopped during handling.

39

Tab chewed but not completely
eaten, no apparent effect.

40

docile on handling.
44

Could not find tab, no
tranquilizing effect.

Right front, above pad.

Feet frozen, skin broken and bones
visible. Bleeding evident, no broken
bones.

45

Did not chew or ingest tab.

Right hind, by 3 toes.

Some bleeding, no broken bones.

46

Could not find tab, docile on
handling.

Left front, inside 2 toes.

Small 1/2" cut on top, no broken bones.

47

Tab apparently ingested but
pulled drag about 1/2 mile;
docile on handling.

Left front, 2 center toes.

Some swelling, both toes broken at first
joint and cut.

a

600 mg of propomazine hydrochloride per tab.

Tab construction modified from Balser 1965.

J,.

'"

-..J
I

�Table 17. Vertebrates which were killed, released, or escaped but not radio-collared
1975 to 25 January 1976.
Species
Cottontail (Sylvilagus sp.)
Porcupine (Erethizon dorsatum)
Bobcat (Lynx rufus)
Coyote (Canis latrans) caught but escaped
1:
a

October

November

la

3b
la

during the coyote trapping effort, 9 October

December

January

Total

0
0
0

5
2
1
1

1

9

0
0
0

0
0

1
0
1
la

1

4

3

1

Released alive.

b

One rabbit released alive.

J,.
Table l7A. Cottontail rabbits recorded per person hour within the study area for each of 11 months 1975-76 compared to similar values
obtained during 1974-75.
Month
May
June
July
August
September
October
November
December
January
February
March

Total Number
Person Hours

Total Number
Rabbits

105
2,756
1,406
858
315
361
445
469
429
336
291

16
399
440
388
143
124
119
75
62
33
29

Rabbits Per Person Hour
1975-76
1974-75
.152
.145
.313
.452
.454
.343
.267
.160
.145
.098
.100

.413
.104
.601
.454
.374
.312
.467
.364
.107
.126
.213

'"

00
I

�Table 18.

Details on capture sites, ear tags and transmitters

fitted to nine coyotes 9 October 1975 to 27 December 1975.

CaEture Site
Elevation
Legal DescriEtion
Twp. R
1/4 S
(ft)

Ear Tags
Left
Color Posi.

a

Ear Tag
No.

Date of
Capture

Sex

Age
(Year)

39

10-9

M

&lt;

1

7,040

SE

34

12N

70W

Y

F

R

40

10-17

M

&lt;

1

6,700

NE

28

UN

70W

Y

F

41

11-14

F

&lt;

1

7,100

SE

4

11N

70W

R

42

11-17

F

&gt;

1

6,800

SE

3

UN

70W

43

11-21

F

&lt;

1

6,960

SE

22

11N

44

11-25

F

&gt;

1

6,720

NW

22

UN

Right
Color Posi.

No.

Transmitter
Channel

Wt (g)

R

4540

20

149

R

R

4539

11

150

R

Y

F

4496

4

R

F

Y

R

4538

14

70W

R

F

Y

R

4501

18

70W

R

R

Y

F

4497

5

148

157

~I

co
co

45

12-14

M

&gt;

1

7,000

SE

22

11N

70W

Y

R

R

F

4541

9

46

12-17

F

&gt;

1

6,920

SE

3

UN

70W

R

F

Y

R

4494

2

47

12-27

M

&gt;

1

7,000

NE

17

UN

70W

Y

R

R

F

4507

16

146

a
Identically numbered metal ear tags. Black numbers one inch in height on yellow (Y) or red (R) backgrounds.
of tags on ear; indicating visibility of numbers from front (F) or rear (R) of coyote.

Posi. denotes (position)

I

�Table 19. Approximate mean distances (miles) traveled by individual radio-collared coyotes from their capture sites to their
relocation sites, 10 October 1975 to 31 March 1976.
C0:i0te Number
Month

Statistic

39

40

October

N

12
.89
.33
.40
1. 70

6
1.17
.72
.32
2.20

18
1.08
.10
.86
2.08

13
1.62
.10
1.20
2.08

22
.89
.07
.46
1.25

-

X

SD
Min
Max
November

N
X

SD
Min
Max
December

N
X

SD
Min
Max
January

N
X

SD
Hin
Max
February

!'!
X

SD
Min
Max
March

N
X

SD
Min
Max

18
.81
.40
.20
1. 75
10
.78
.57
.06
1.81
9
1.45
.47
.52
2.03

41

42

43

44

7
.89
.87
.46
2.86

4
.52
.06
.48
.60

45

46

47

-

29
.68
.34
.30
1.90

9
.89
.52
.32
1.90

17
2.82
1.00
.75
4.40

9
.64
.27
.30
1. 20

6
1.50
.84
.90
2.80

5
.98
.62
.30
1.50

3
.13
.06
1.10
1.60

13
.61
.24
.40
1.30

4
2.18
0.63
1.60
2.90

5
3.18
.54
2.50
3.20

4
.78
.50
.60
1.20

5
.85
.29
.50
1.30

9
1.25
.46
.50
1.60

2
1.65
.78
1.10
2.20

1

7
.81
.53
.31
1.87

17
1.63
.80
1.25
1.84

-

16
1.01
.68
.31
2.06

22
.64
.17
.31
1.06

20
.54
.87
.16
1.06

1
1.00

5
.44
.07
.40
.56

16
1.63
.45
.59
1.84

13
1.00
.61
.53
2.12

14
.54
.09
.46
.84

10
.29
.25
.16
.98

3
.90
.97
.53
1.23

-

-

-

-

.81
.81
1

-

-

.25
.25

-

-

-

-

I

lJ1

0
0
I

1.00
1.00

�Table 20. Approximate mean distances (miles) traveled by all radio-collared
month, 10 October 1975 to 31 March 1976.

No. Coyotes

Month

i

as
42
97
63
94
71

9
8
8

Details on 7 of 23 radio-collared

Ear Tag
No.

Sex

Dates
Death
Capture

24

M*

7-31-74

1-21-75

26
28

M
F*

8-24-74
9-2-74

3-7-75
1-7-76

31
32
34
37

F*
M
M
F

9-6-74
9-15-74
11-11-74
12-15-74

3-7-75
10-7-75
10-7-75
4-12-75

*

No. Relocations

2
4
7'

October
November
'December
January
February
March

Table 21.
1976.

,.

coyotes from capture sites to relocation sites by

Mean

SD.

Min

Max

.98
1.16
1.18
1.16

.49
.52
.95
.81
.54
.64

.32
.46
.30
.20
.06
.16

2.20
2.86
4.40
3.20,
2.06
2.03

0.89
0.95

coyotes known to have died following capture with steel traps, 31 July 1974 to 31 March

Cause
of Death
Trapped and
choked
Shot (ground)
Shot (helicopter)
Shot (ground)
Shot (ground7)
Shot (ground)
Trapped &amp; shot

Legal DescriEtion of Sites
Death
CaEture
1/4
S
T
R 1/4
S T

R

Approx. Distance
(miles) Between Capture and Kill Sites

SW

29

llN 70W

SE

32 llN 70W

1.2

SW
NE

29
27

UN 70W
llN 70W

NE

20 llN 70W
32 ION 68W

1.4
9.5

NE
NW
SW
SW

29
27
29
26

llN
UN
llN
llN

NW
SE
SE

22 llN 70W
15 llN 70W
10 9N 72W
32 13N 7lW

1.3
1.3
16.8
13.0

70W
70W
70W
70W

-

-

Coyotes more than one year of age as estimated by the criteria of Gier (1957:50-51, 55).

I
IJ1

o
f-'

Remarks
Most of left front forepaw
gone.
Reportedly "molesting" cattle.
No foot injury, no collar, one
ear tag missing.
Reportedly "molesting" cattle.
Dead perhaps a week or more.
Found by landowner.
Some toes missing on one foot.
Upper Dale Creek, Wyoming.

I

�-502-

While radio telemetry is the only possible way to directly assess mortality
in free ranging fawns, radio tracking as described herein may in itself
constitute a source of mortality.
I speculate that the presence of the radio
collar and the daily tracking as employed in this study may attract predators
and that the daily disturbance and flight may place critical stress on the
fawn or cause the doe and fawn to separate, perhaps increasing the vulnerability of the fawn to predation or accident. Capture, handling, and
collaring almost certainly caused some does to abandon their fawns in this
study. White et al. (1972:897) reported that in Texas, white-tailed deer
(Odocoileus virginianus), "capture, and particularly the marking of young
fawns increased mortality." The disappearance of collared fawns and lost
collars accounted for 13 of the 53 fawns captured and equals the documented
loss to predators (Table 8). An additional 12 fawns were found dead, but
the cause of death was not determined (Table 8). Therefore, the history
of 25 of 53 fawns is essentially unknown. It is clear that any statistic
of fawn mortality presented herein must be regarded as a gross est.imate. I
hypothesize that appreciably more precise estimates of fawn mortality may
be very difficult and certainly more expensive to obtain.

LITERATURE CITED
Anderson, A. E. 1975. Evaluation of radio telemetry. Pp. 415-474. In
Colorado Div. Wildl. Game Research Report, July. Part 2. 191-504 p.
Balser, D. S. 1965. Tranquilizer tabs for capturing wild carnivores.
Wildl. Manage. 29(3):438-442.

J.

Cowan, I. McT., and P. J. Bandy. 1969. Observations on the haematology of
several races of black-·tailed deer (Odocoileus hemionus).
Canadian J.
Zool. 47(5):1021-1024.
Fashingbauer, B. A. 1962. Expanding plastic collar and aluminum collar
for deer. J. Wildl. Manage. 26(2):211-213.
Geduldig, H. L. 1975. Self-expanding plastic collar for radio-tagging deer.
Coyote Research Newsletter 3(1):9. (Abstr.).
Gier, H. T. 1957. Coyotes in Kansas. Kansas State Univ. Agric. Expt. Sta.
Bull. 393. (Revised 1968). 118 p.
White, M., F. F. Knowlton, and W. C. Glazener. 1972. Effects of dam-newborn
fawn behavior on capture and mortality. J. Wildl. Manage. 36(3) :897-906.

Prepared by __ ~_--:--:-----:£
__ ~
Allen E. Anderson
Wildlife Researcher

_

�APPENDIX I
-503COLLARED FAWN LOCATION RECORD, W-38-R, WP 19, J 1, Southern Portfon
Date(s)
,'Observer(s) and Hours of Search

~----------------.

, Total Man-Hr Searched

------''---='--

TIME

Fawn
No.

Comments:

Began
Search

First
Signal

Location
Telemetric
Visual

Observer(s)

Comments

------------------------------------------------,-------

��July 1976
-505-

JOB PROGRESS

State of
Project

REPORT

COLORADO
W-38-R-31

No.

Deer~Elk

Work Plan No.

19

Job Title

Evaluation

Period

April 1, 1975 to March

Covered:

Personnel:

Investigations

Job No.
of Herd Structure

2

_

Methodology

31, 1976

Allen E. Anderson, Dan Baker, David C. Bowden, Allison Broida,
Eugene G. Johnson, Beth MacConnell, Dean E. Medin, David L.
Otis, Tom H. Pojar, Elmer Remmenga, Laverne Stelter, Charles
Wallmo.

ABSTRACT
The validity of three models for sample observations from'each of two sampling
plans for estimating fawn:doe and buck:doe ratios of Rocky Mountain mule deer
(Odocoileus hemionus hemionus) are examined and the results of data analysis
under the most appropriate model are presented.
During 1962-65, the first
sampling plan (route counts) consisted of dawn-to.,.dusk circular routes
designed to classify the maximum number of deer on representative portions
of the Cache la Poudre winter range.
The second sampling plan (~uadrat counts)
consisted of classifying deer within randomly-selected,
l/4-mile
quadrats
stratified by subjective estimates of low and high deer densities and proportionally allocated wi-thin each of eight topographic quadrangles on Cache
la Poudre winter range and two quadrangles within and adjacent to the Maxwell
ranch during 1974-75.
In 1975-76, the two sampling plans were compared by
running each of 12. routes through as many randomly located quadrats as could
be traversed in one day. Thus randomized, the route and quadrat counts obtained
on consecutive days could be used to compare their respective fawn:doe
and buck:doe ratios and estimate sample sizes.
Time limitations, however,
reduced the number of quadrats on Cache la Poudre range from 106 to 58.
Because of the statistical models examined, fawn:doe and buck:doe ratios are
stated in terms of PF = F/(F + D) where PF = the proportion of fawns,
F = the number of fawns, and D = the number of does and PB = B/(B + D) where
PB = the proportion of bucks, B = the number of bucks, and D = the number
of does. Tests indicated a significant (P&lt;O.Ol) departure of the observations
from the binomial distribution, a biased SE in the linear regression through
the origin model, and no serious flaws in the standard ratio model.
The
following findings are based on the latter model .•
Route counts indicated a significant decrease (P&lt;O.Ol) in PF from 1962-65
during both fall and winter.
PF also decreased significantly
(P &lt; 0.01) from
fall to winter during 1962-63, 1963-64, and 1964-65.
PB increased significantly

�-506-

ABSTRACT

(Continued)

(P&lt;O.Ol) in successive winters 1962-65, but decreased significantly (P&lt;0.025)
from fall to winter when proportions were pooled over years.
In general,
the 1975-76 fall and winter PF and PB were significantly (P&lt;0.05) less than
their counterparts, 1962-65.
The significant annual changes in PF and PB during
1962-65 were concurrent with documented environmental changes and certain
physiological changes in Cache la Poudre mule deer.
Quadrat counts during 1974-75 revealed that significantly higher PF (P&lt;O.lO)
and PB (P&lt;O.Ol) were found in the low density strata.
During 1975-76, the
same pattern occurred but the differences were not significant.
The route and quadrat sampling plans yielded nearly identical PF and PB values
during 1975-76.
Based on the 1975-76 deer classifications,
it was found
that to be within 10 percent of the true ratio at the 95 percent confidence
level, route counts would require 6 routes for PF and 127 routes for PB while
quadrat counts would require 103 quadrats for PF and 635 quadrats for PB.
Minimum mean densities (±SE) of mule deer estimated on Cache la Poudre
winter range were l6.40±3.42 deer/mile2 based on quadrat counts and 43.l7±11.93
deer/mile2 based on route counts.
Possible errors in the measurement of the
route length and width may be factors in this discrepancy.

�-507EVALUATION OF HERD STRUCTURE METHODOLOGY
David C. Bowden and Allen E. Andersona

In a previous report (Bowden and Anderson 1975), we briefly reviewed the
literature on obtaining sample ratios of sex and age and their various
uses in North Anlerican deer. It was noted that there was little information
on either the statistical attributes or sampling plans needed to obtain the
necessary data. We pointed out that although Caughley (1974:562) had
concluded that; "age ratios cannot be interpreted without a knowledge of
the rate of increase and if we have an estimate of this we do not need age
ratios," there was some evidence (Smith ~ al. 1969, Swank 1958:78-79,
Taber and Dasmann 1958:129) that fawn:doe ratios may reflect the dynamics
of some deer populations. The present report covers much of the same material
as the 1975 report but with corrections and a major change in conclusions.
The Cache la Poudre data were used to develop models and compare sampling
plans since it represents the only available long-term source of fawn:doe
and buck:doe information obtained by ground counts according to a definite
sampling plan. A new model is presented which permits valid estimations
of the standard errors of both fawn:doe and buck:doe ratios but at a
practicable level of sampling intensity for fawn:doe ratios only.

P. S. OBJECTIVE
To test and impr~ve the field and statistical methodology used to estimate
selected population parameters of mule deer and coyotes.

SEGMENT OBJECTIVES
1.

Study existing herd structure data to provide an improved statistical
rationale for analyses of fawn:doe and buck:doe ratios.

2.

Analyze data and submit reports for publication.

METHODS AND MATERIALS
An ecological study of the Rocky Mountain mule deer population within a
1376.5 km2 portion of the Cache la Poudre River drainage in Roosevelt National
Forest in Larimer County, northcentral Colorado was conducted from 1961 to
aDr. Bowden, Dept. Statistics, Colorado State University, wrote most of the
METHODS AND MATERIALS and the RESULTS sections and with the assistance of
Graduate Students Gene G. Johnson and David L. Otis developed the statistical
rationale described herein. A. E. Anderson wrote the ABSTRACT, INTRODUCTION,
RECOMMENDATIONS AND DISCUSSION and CONCLUSION sections. Field work was performed by A. E. Anderson and D. E. Medin during 1962-65, by A. E. Anderson
and C. Wallmo during 1974-75, and by A. E. Anderson, D. Baker, A. Broida,
B. MacConnell, T. H. Pojar, L. Stelter during 1975-76.

�-508-

1965. In one facet of the study it was of interest to examine the fawn, doe,
buck components of population at specified time periods (Anderson and Medin
1965). Since it was not feasible to estimate the actual number of deer in
each category, the fawn to doe ratio of population sizes and the buck to doe
ratio of population sizes were studied.
Deer were classified into fawn, doe, buck categories in late November through
December and again in January through early February for 3 winter seasons
beginning in 1962: 11-29-62 to 12-18-62, 1-3-63 to 1-27-63, 11-21-63 to 12-14-63,
1-9-64 to 1-31-64, 12-3-64 to 12-17-64 and 1-13-65 to 1-27-65. The winter
range of the deer population in the study area was sampled by selecting 10
representative sections of the winter range (Fig. 1). A route to be traversed
by walking was outlined on aerial photographs of each representative section.
The routes were located to classify the maximum number of deer. The length
of the route was determined by the distance which could be covered in the
given terrain by walking from dawn to dusk. The routes were covered by two
observers working together, often with one observer maintaining a vantage
point while the other observer attempted to bring hidden deer into view.
Classifications of deer were made by using 8X40 or 9X35 binoculars and 20X
spotting scopes. Deer were classified as fawn, doe, buck or unidentified by
groups on the basis of relative body size, presence or absence of antlers
and the applicable criteria listed by Dasmann and Taber (1956:80). The time
to the nearest minute was recorded for each observation to avoid duplication
between observers. A group was identified by observing the joint behavior and
spatial distribution of the deer. Only those groups in which all deer were
classified (1962-65) were used in this analysis. We refer to this sampling
plan as the route counts.
In 1974 a new sampling plan was applied to the study area. The deer winter
range was divided on 1:24000 scale U.S. Geological Survey Topographic
Quadrangles into sampling units (1/4 mile2 or one-half mile per side)
and the sampling units placed into strata. Each sampling unit was classified
subjectively into high and low deer density strata by eight Topographic
Quadrangles (16 strata). Counts started 22 November and were concluded 31
December except for two sampling units covered 9 January, 1975. Sampling
effort was proportionally allocated according to the total number of sampling
units per stratum (Topographic Quadrangle) except that at least two sampling
units wer taken from every stratum. Two walking observers classified deer
on each selected sampling unit. The sampling unit was approached and traversed to classify the maximum number of deer. Four sampling units were
usually counted beginning at dawn and finishing at dusk each day. The
counting sequence of the sampling units was arranged so that adjacent or
nearby sampling units were counted on the same day to minimize duplicate
classification on the same deer and to allow for efficient use of available
time. Deer were recorded by groups as before. We refer to this sampling plan
as the quadrat counts.
In 1975 the two previous sampling plans were applied to the study area from
28 November 1975 to 6 January 1976. The locations of 12 routes were randomized
to permit valid; (1) comparison of the fawn:doe ratios and buck:doe ratios
obtained by the two procedures, and (2) estimation of sample sizes for the route
counts. The route locations were randomized on the Topographic Quadrangles

�-509-

by running each route through as many randomly located quadrats as could be
traversed by walking from dawn to dusk (Figs. 1-12). To allow sufficient time
for counting deer under both sampling plans, the 106 quadrats covered under the
1974 quadrat count was reduced to 58. The deer on each route were counted the
day prior to the corresponding quadrat count. During the route counts an
attempt to estimate the density of deer was made by separately recording
each group of deer observed within a 500 ft wide strip centered on the route
location as drawn on the topographic quadrangle. The 500 ft strip was
measured initially by pacing and by noting prominent topographic features
on the topographic quadrangle. Strip width was not checked once the count was
underway. Strip length was measured on the topographic quadrangle to the
nearest 0.01 miles with a cyclometer. In addition, if all deer with a group
were identified by buck, doe, fawn categories the group will be called completely
classified and if at least one deer was not identified by buck, doe, fawn
category the group will be said to be incompletely classified.
In this report we present these data and examine the validity of assumptions
or models for the sample observations from each sampling plan which permit
the development of measures of reliability of estimates of ratios of the deer
population size components. In addition, the results of the data analysis
under a model not contradictive to the data will be presented.

DESCRIPTION OF AREA
That portion of the Cache la Poudre winter ~ange sample has been described
by Anderson ~ al. (1970:390). The Maxwell ranch area is described in the
Work Plan 19, Job 1 Progress Report accompanying this report.

RESULTS

Statistical Models for the Observations - 'Route 'Counts
In the following presentation, the discussion will be on the fawn:doe ratio.
The assumptions and analyses developed also apply to the buck:doe ratio.
The results of analyses for both the fawn:doe and buck:doe ratios are presented.
The following notation permits an easy definition of the equivalent parameters.
Consider the population of deer at a specified point in time and define F, D
as the number of fawns, does, respectively, in the population at this time.
The stated ratio of interest is RF = F/D. Consider instead PF = F/(F + D)
Then clearly, RF = PF/(l - PF). The statistical models given here will be
stated in terms of PF so that the estimation problem will be to determine
the proportion of fawris (relative to does and fawns). Estimates of PF can
be converted into estimates of RF'
Much of the difficulty in objectively sampling natural wildlife populations
arises in determining a sampling frame (a list of sampling units) which can
be sampled by using statistical sampling plans while at the same time providing sampling units which when measured provide data on the target population

�-510-

j
/

."
'''"',

\

(
\

c

,

~

\

\

/------

-~-~

\

-...-,..-----;-~...,
.r)

ll..

I

,-'-'T-'"
,

..

" \\ .

Fig. 1. All-day walking route (about 7.4 miles in length) and randomly
selected, 1/4-mile2 quadrats, used to classify deer in the Horne area,
Cache la Pcudre drainage.

�-511-

fig. 2. All-day walking route (about 6.0 miles in length) and randomly
selected, 1/4-mi1e2 quadrats used to classify deer in the Sevenmi1e Creek
area, Cache 1a Poudre drainage.

�-512-

Fig. 3. All-day walking route (about 9.2 miles in length) and randomly
selected 1/4-mile2 quadrats used to classify deer in the Indian Meadows
area, Cache la Poudre drainage.

�-513-

Fig. 4. All-day walking route (about 6.2 miles in length) and randomly
selected 1/4-mile2 quadrats used to classify deer in the Bennett Creek
area, Cache la Poudre drainage.

�-514-

e

l~/ .:':(j
~V~.-

;.

(T)

o

./

Fig. 5. All-day walking :route (about 9.4 miles in length) and randomly
selected 1/4-mile2 quadrats used to classify deer in the Eggers area, Cache
la Poudre drainage.

�-515-

~ig. 6.

All-day walking route (about 6.6 miles in length) and randomly
selected 1/4-mile2 quadrats used to classify deer in the Sheep Mountain
area, Cache la Poudre drainage.

�-516-

./"

:;'")(~o

..

'

r·

)

Fig. 7. All-day walking route (about 5.0 miles in length) and randomly
selected 1/4-mile2 quadrats used to classify deer in the Red Mountain area,
Cache la Poudre drainage.

�-517-

Fig. 8. All-day walking route (about 5.8 miles in length) and randomly
selected 1/4-mile2 quadrats used to classify deer west of Seaman Reservoir,
Cache la Poudre drainage.

�Fig. 9. All-day walking route (about 7.6 miles in length) and randomlyselected 1/4-mile2 quadrats used to classify deer in the Livermore Mountain
area, Cache la Poudre drainage.

�-519-

Fig. 10. All-day walking route (about 5.6 miles in length) and randomlyselected 1/4-mile2 quadrats used to classify deer in the Greyrock Mountain
area, Cache la Poudre drainage.

�-520-

Fig. 11. All-day walking route (about 6.4 miles in length) and randomlyselected 1/4-mile2 quadrats used to classify deer in the west portion of
Hewlett Gulch, Cache la Poudre drainage.

�v

.~:
I

\

I
J

I
I

I
!

Fig. 12. All-day walking route (about 9.2 miles in length) and randomlyselected 1/4-mile2 quadrats used to classify deer east of Seaman Reservoir,
Cache la Poudre drainage.

�-522-

of interest. A statistical sampling plan is a scheme of sampling unit
selection such that the probability of obtaining each individual sampling
unit in a sample of a given size is known or can be calculated. Conceptually,
the use of the animals themselves as a sampling unit is an appealing sampling
frame. In its simplest form each animal is a sampling unit. A desirable
property of having the animals as sampling units is that conceptually the
value of PF could be determined without error if all the sampling units were
measured. The practical problem with such sampling frames is that once a
specified sample of animals was determined, the animals have to be located
and classified. Such a sampling frame also implies that the population size
is known and that the animals or groups of animals (sampling units) can be
listed.
As an alternative to forming sampling units from or of the animals themselves,
sampling frames are constructed by considering a given area or region, perhaps volume, that the population is associated with, restricted to, or contained in. As in the·1974 sampling of the Poudre drainage, the region is
subdivided into parts which are considered as the sampling units. The construction of such a sampling frame and the actually selection of the sampling
units to be measured is not difficult. However because of the movement of the
animals among the sampling units and because the animals'behavior limits their
detection by human observers, measurement errors on such sampling units can be
quite important. The explicit or implicit use of these two types of sampling
frames will now be examined relative to the data of the present study.
First, statistical models or sets of assumptions which give rise to the use of
a sampling frame dependent on the animals themselves will be examined. Thus,
let n represent the number of animals identified as either does or fawns while
d and f will be the numbers of does and fawns in the sample of n.
Consider the following assumptions:
1. The classifi~ation category for each animal is independent of the
other animals in the sample. Alternatively the probability that an individual
animal is a fawn does not depend on the other animals in the sample.
2.
is PF'

The probablility that an individual animal in the sample is a fawn

It is well known that these assumptions imply that the probability distribution
of f is given by the binomial distribution where n is the number of trials,
PF is the probability of success on an individual trial and f is the number of
successes out of n. In the sampling frame terminology this same binomial distribution is arrived at by assuming that a simple random sample with replacement
is taken from a sampling frame which lists the individual animals. The sampling
procedure without replacement implies the distribution of f is given by the
hypergeometric distribution. But in most situations this hypergeometric
distribution can be adequately approximated by the binomial distribution given
above.
In general, the practicality of actually constructing and using such a sampling
frame is nil, but it is common practice to assume the collected data, however
gathered, satisfy assumptions 1. and 2. and in essence claim that the

�-523-

collection procedure operates as if a simple random sample of the individual deer was taken. It remains to establish that the data are not in
strong disagreement with the implications of these assumptions.
Since the
alternatives to accepting assumptions 1. and 2. are complicated, the disagreement of the actual data with assumptions 1. and 2. will be explored
next.
The assumptions will be examined for the data collected on the route counts
from 1962 to 1965 and also for the 1975 route counts. On each route the deer
were observed in natural occurring groups, that is, classified into groups
by their behavior and the proximity of individuals to each others.
A group
in the present case would be one or more does or fawns. Now if assumptions
1. and 2. are true, then the distribution of the number of fawns within groups
of fixed size should be binomial.
If adequate numbers of groups were
observed at each given group size, then a chi square goodness of fit test
could be applied at each group size and pooled.
In the actual data, the
limited frequency of groups of a given size prohibited the use of this test.
However, Cochr&amp;n (1936) presented the index of dispersion test for such
cases of small expected frequencies.
The index of dispersion test can be
written as:

x2

m
~

(m-l)
i

= 1

(fi - ni p)2
ni

P (1 - p)

where m is the number of groups observed, fi is the number of fawns in the
ith group, ni is the number of fawns and does in the ith group and p = L fil
~ ni' The distribution of the index of dispersion is approximated by the
chi square distribution with m-l degrees of freedom, if assumptions 1. and 2.
are
2 true. Note that this is a two tailed test, that is, small values of the
X statistic indicate more regularity than expected from a binomial distribution and hence should lead to rejection of the binomial hypothesis as well
as large values of the X2 statistic.
Cochran (1954) suggests when m &gt; 15
that the mean and variance of the index of dispersion test should be used
to form a standard normal test statistic (~test).
Rejection of a common
binomial distribution for groups of all sizes can occur in the use of the
statistic even if the distribution of the fawns at a given group size is
binomial if the proportion of fawns changes over or among the group sizes.
The index of dispersion can be written in two steps to examine the alternative
just raised. First calculate:

K
~

2

X

(m-K)
k

mk

~
1

i = 1

K
k = 1 ~

Where K is the number of different group sizes, nk is the size of the groups
indexed by k, mk is the number of groups of size nk' fki is the number of

�-524-

fawns in the ith group of size nk, Pk is given as Pk = ~ fk·/mknk and Sk2 is
the sample variance of the number of fawns in the sampl~ oflmk groups of
size nk. This criterion, tests whether the distribution of the number of
fawns at a fixed group size is binomial. The significance levels of the
criterion can be approximated by using a chi square distribution with m-K
degrees of freedom. Since the K components of this test are independent
the lack of fit of the binomial distribution can be made at each group size
where each individual component has a chi square distribution (approximately)
with mk-l degrees of freedom.
Secondly calculate:
K

x2 (K-l)

L

nkmk (Pk-p)2
p

(l-p)

If one does not reje~t the binomial distribution at each group size, then
this criterion tests the equality of the proportion of fawns among groups
of different size. If at each group size the number of fawns is binomially
distributed, then the criterion above is approximately distributed as a chi
square distribution with K-l degrees of freedom. In this test only the
upper tail of the X2 distribution should be used for rejection. If in the
first step the denominator of x2m-K is changed from nkPk (l-Pk) to
nkP(l-p), then

The values of the index of dispersion test for the does and fawns recorded
on individual routes are given in Table 1. The calculations were performed
only for the 1962 and 1963 counts, since the uniformity of the rejections of
the binomial distribution clearly indicated an alternative model or set of
assumptions was needed. All significant X2 values were smaller than the
lower tail percentage point indicating less variation in the number of fawns
than expected from a binomial distribution. The component of the index of
dispersion x2m-K, was calculated for three route counts, which were selected
only because they had a fair range in group sizes. Thus, we examined only the
binomial distribution assumption by eliminating the possible influence of
changing proportions of fawns with group size (Table 2). Again the binomial
distribution is clearly in disagreement with the data.
The next set of assumptions will serve as a basis for the analysis of the
present data set. Therefore, some practical aspects of route sampling will
be examined here, which were not mentioned when the binomial distribution
was discussed. These aspects would also be important if one had accepted the
binomial distribution for a description of the number of fawns given the
group sizes. The areaof winter range covered by walking along a route for
one day is obviously limited. Hence, the area sampled has various relationships to the home range of the deer being classified. A few animals may
have their home range entirely within the area sampled while some animals
may have most of their ho~e range within the sampled area and some may have
only a limited part of their home range inside the sampled area. The
probablility of classifying a particular animal may depend on the proportion
of his home range within the area sampled and the proportion of time the
animal occupies this (included) part of the home range.

�Table 1. Values for the index of dispersion
on Cache la Poudre winter range, 1962-65.

CX2m_l)

test for does and fruvns recorded on 10, all-day walking

routes

Ivashout
Gulch

Sevenmile
Creek

Bennett
Creek

Kelly
.Flats

E. Elkhorn
Creek

E. Hewlett
Gulch

W. Hewlett
Gulch

Youngs
Gulch

Livermore
Mountain

Index

8.36

2.22

1. 09

8.14

18.92

11.90

8.13

6.79

11.99

12.81

df

13

2

3

26

43

27

19

14

25

25

3.94

4.56

11.21

3.13

5.06

4.00

7.53

4.23

3

16

33

11

7

1

19

13

Seaman
Reservoir

1962 November

1963 November
Index
df

.44
1

0
1

I
IJ1

N
IJ1

I

�-526Table 2. The component of the index of dispersion, X2 m-K, for three routes
selected because of their wide range of group sizes of does and fawns.
Sampling
Period

Route

X

2

Sig.
Level

d.£.

m-K

1962 November

E. Elkhorn Creek

18.06

34

P

= .01

1964 December

E. Elkhorn Creek

18.74

36

P

= .01

1963 January

Livermore Mountain

10.62

24

P -

.01

Now the primary interest in this study is with PF' Define F(t) and D(t) to
be the number of fawns and does, respectively, present in the sampled area
at time (t). Also, define PF(t) = F(t)!(F(t) + D(t)). It will be assumed
that although individual animals move into and out of the sampled area, PF(t)
will remain constant at the value PF. In addition, given that an animal is
present in the area when the classification is performed, its behavior may
be such that the chance of classifying the animals is greater or less than the
mean probability of classifying an animal given it is present. Actual knowledge of the individual probabilities of classification are not necessary for
unbiased estimation of PF, but knowledge of the difference in the mean probability of classifying a doe versus a fawn would be necessary.
The present data does not allow estimation of the difference in the mean classification probabilities. Hence, it will be tacitly assumed that the mean probability of classifying a fawn is equal to the mean probability of classifying a doe.
It may be reasonable to assume that
f.
r,

Sn. + E:i
~
i

1, 2,

...

,

m

where fi is the number of fawns in the ith group with ni total does and fawns,
13 is equal to PF and E:iis a random residual component and m is the number of
groups completely classified. Further, assume that the expected value of E:i
is zero with the variance of E:i = 02 and the covariance between E:iand Ei~'
ifi~ is zero. With the ni considered as fixed, the model is that of a linear
regression through the origin. If the distribution of fi given ni was binomial,
then the variance of Ei would be ni PF (l-PF) or proportional to ni' Examination
of plots of fi versus hi' indicate that the variance of E:iis constant, say
02 thus providing alternative evidence against the binomial distribution of fi
given ni.
An earlier examination of the data was performed with the linear regression
through the origin approach (Anderson and Bowden 1975). The least squares
estimators of the parameters of interest are ~ = ~ fi ni!~~n~2, ~
__
d_fi2
~ ~ finD! (ra+L) and an estimator of the standard error of S a.s v 02!~ni 2

�-527-

However, in a theoretical evaluation of the ratio estimation in finite
populations Otis and Bowden (to be published) found that the.above standard
error formula underestimates the true standard error of S if expected value
of € is not zero. It was found that when the proportion of fawns at each
group sizes varies slightly from S then a considerable bias may exist in
the standard error formula. Since the sampled deer population is finite it
would be unreasonable to expect the proportion of fawns at each group size
to be exactly the same.
Thus, the standard ratio estimator A
m f /~
R = L
i "ni
will be used in this report to estimate PF and PB. In the study by Otis and
Bowden it was found that
could be considered as an unbiased estimator of PF
or PB in sample sizes as small as 10. It was also found that the estimator
A
2
of the variance of AR,
1\
,,1:.
L (fj - Rnj)
variance (R) = n2
n(n-l)
is essentially an unbiased estimator of the variance of R whether the expected
value of ei was zero or not. It follows that the critical assumption required
by the regression estimator of equal proportions of fawns and bucks for the
population irregardless of group size if avoided by using R. In using the
ratio estimator instead of the regression estimator, there is a loss of
preC1Slon only if that assumption holds exactly which seems unreasonable,
again, in a finite population.

R

A

A

Now the parameter of interest is PF for the entire winter range from which the
routes were selected. The problem of determining how to combine the 1962-1965
individual route estimates of PF into an estimate of PF for the winter range
is considered next. It was noted that routes were not randomly selected from
the winter range. Essentially, the winter range was stratified into 10 areas
and a route was located within each strata. In combining the estimates of PF
by route or by strata it would be desirable to weight each route estimate by
the proportion of the deer population in the corresponding strata. Since this
weighting factor is unknown, this weighting is not possible.
If the route
estimates (R = PF) are not significantly different, the weighting problem is
not important. A test for significant differences among routes was performed
for each of the six sampling times. Significant differences (P&lt; .025) were
noted for the 1962 fall and 1963 fall counts, while for th~ other four times
no significant differences (p&gt; .10) were found. Thus, for at least the two
fall counts the weights given to each route would affect the value of combined
estimate.
The sampling procedure using routes as the sampling unit is a two stage
sampling procedure. First stage selection is made to determine a route location. Counting a route on a particular day is a second stage selection of m
groups of deer classified from the M groups which have a nonzero probability
of being classified on that route. We developed the assumption above that
the m groups give an unbiased estimate of PF for that route. The variability
of counting conditions from route to route indicate that the proportion of
groups classified (m/M) varies considerably and is also greatly influenced
by the weather conditions associated with the count time. Thus, an unweighted
ratio of total counts over routes of fawns to total does and fawns over
routes may have considerable bias as an estimator of PF for the winter range.
However, since appropriate weighting factors are not known, the unweighted
ratio estimator willAbe used. In addition, since route locations were stratified the formula SE(R) as previously given should over estimate the true

�-528-

A

standard error of R. The 1975-76 study, using a randomly located route
within a stratum, indicate that it is not unreasonable to consider the
general location of the 1962-1965 routes as randomly located within their
corresponding stratum. However, search techniques within the route locations
in 1962-65 tended to emphasize area coverage to find bucks.
The frequencies of the total groups of doe and fawn and buck and doe mule
deer are shown for each of six count periods, 1962-65, in Tables 3 and 4,
respectively. A similar tabulation for the 1975-76 total group frequencies is
shown in Table 5 (doe and fawn) and Table 6 (buck and doe). The raw data,
variances and covariances, fall versus winter differences, and sample size
requirements are shown for the doe and fawn, and buck and doe data by individual route for six count periods, 1962-65 in Tables 7 and 8, respectively. A
test for differences among fall fawn proportions indicated a significant
decrease (P&lt;0.05), 1962-65. Similarly, a significant decrease (P&lt;O.Ol) was
found among the winter fawn proportions. A test of equality of the Fall PF
with the following Winter PF pooled over the three pairs was also significant
(P&lt; 0.025). Similar tests were performed for the buck proportions. No significant differences were found among the fall proportions (P&gt;.05). However,
significant differences (P &lt; .01) were found among the winter proportions
and among fall versus winter proportions pooled over years (P &lt; .025). It
should be noted that the main contributor to these differences if the low
proportion observed in Winter 1963.
In general, a comparison of the proportion of fawns (PF) recorded on the
route counts of 1975-76, to the route counts of 1962-65 shows that the 1975-76
ratio is significantly less (P&lt; 0.05) than the earlier ratios. Exceptions
are the winter 1964 and winter 1965 proportions where no significant difference
(P&gt; 0.05) is noted.
Based on the 1975-76 counts, sample sizes (number of randomly determined
routes) for PF and PB on Cache la Poudre winter range are listed in Table 11.
It appears that adequate sampling of PB requires impractically large numbers
of routes.
Statistical Models for the Observations-Quadrat

Counts

For the quadrat sampling plan the proportion of fawns PF and the proportion
of bucks PB will be estimated by the use of the ratio estimator. As noted
under the route sampling plan this procedure is conservative if the true
model could be writ ten as
fi = Sni + Ei
i = 1, 2, ... , m
where m is number of sampling united observed, fi the total fawns out of ni
total does and fawns observed on the ith sampling unit and Ei has mean of 0
and variance 02. The ratio estimator is robust to departures of population
from the assumptions of mean Ei equal zero and variance of E equal 02 while
the regression estimator is not.

�Table
3 . Frequency of total groups of doe and f awn mule deer recorded
Poudre winter range during each of six sampling periods,
1962-65.
Sample Dates:

Nov. 29, 1962 to Dec. 18, 1962

on 10, all-day,

Sample Dates:

Jan.

1
2
3
4
CI)
5
~ 6
~
7
+ 8
(fJ
Q)
9
10
" 11
~112
13
14

1

44
4
2

10
47
15

3
1

2
5
20
19
7
1
1
1

3

1
2
3
4
2

8/

4

5

6

2
1
1
2

7

4

1
2

15
L

50

76

54

12

7

E

5

1

on Cache La

27, 1963

No. Fawns

54
56
37
23
9

1
3

routes

3, 1963 to Jan.

No. Fawns
0

walking

3

1
1

7
4
1
2
1
5
2
0
1
207

0

1

1
2
3
4
5
6
CI)
7
8
~
~
9
+ 10
a'l 11
o 12
~ 13
o 14
z 15
16
17
18
19
20

21
9
.8
1

2
28
33
19
6
1

E

40

1

1

2

14
12
22
6
4
1
1
1

3

2
7
6
6
3

4

6

7

8

E

23
37
55
32
30
16
11
10
5
2
6
4
0
0
0
0

1
1
2
1
2
2

2

5

1
4
2

0

1
90

63

24

9

5

1

0

1
1

1
0
1
233

------------------------------------------------------------------------------------------------------------~------------

I
\JI
N
1.0

I

�Frequency of total groups of doe and f awn mule deer recorded on 10, all-day,
3,
Table
Poudre ~vinter range during each of six sampling periods,
1962-65 (continued).

Sam121eDates:

Sam}21eDates:

Nov. 21, 1963 to Dec.
14, 1963

Jan.

1

1
2
3
rJl
4
~ 5
•....
+ 6
7
rJl
8
0
p
9
10
z11
12

18
2

8
27
16
3
1
1

E

21

1

2

10
16
2
1
1

(!)

3

No. Fawns

56

30

4

5

1
2
1
3

o

7

on Cache 1a

31, 1964

9, 1964 to Jan.

No. Fawns

0

wa.l.kf.n g routes

1
1
1
1
3

1

0

1

1
2
3
4
5
6
7
8
9
10
~ 11
•....
12
+ 13
~ 14
p 15
16
z 17
18
19
20
21
22
23
24

24
7
3

4
28
22
8

E

35

E

26
29
26
21
5
2
1
5
0
1
1
1
118

1

2

3

4

5

6

7

8

9

L

28
35
19
17
14
5
6

[J)

4L.

3
3
5
7
6
2
1

1
9
4
2
2
1

28
17
11
14
15
7
4
3
3
1
0
2
0
2
0
0
0
0
0

1
1
1
2
1

[J)

1

1

o

1

1

0

62

61

27

21

2

4

2

0

1
1

--------------------------------------------------_._------------------------------------------------------------------

1
215

I
IJ1

w
0
I

�3. Frequency of total groups of doe and fawn mule deer recorded on 10, all-day,
Poudre winter range during each of six sampling periods,
1962-65 (continued).

TLlhlc

Sample Dates:

Dec. 3, 1964 to Dec. 17, 1964

Sample Dates:

Jan.

1

1
2
3
4
rJl
5
~
cu 6
I-"&lt;
7
+ 8
rJl
9
g 10
°11
12
z13
14
15
16

43
12
6
2

5
47
29
11
3
2

L

63

1

2

19
23
13
6
2
1

3

4
4
3
5
1

4

5

6

7

8

I;

1
1

48
59
54
36
20
13
5
8
4
4
0
0
0
0
0
1
252

2
1

01

98

64

17

6

3

0

on Cache La

27, 1965

No. Fawns

1
1
1
3

routes

13, 1965 to Jan.

No. Fawns
0

walking

0

1
2
3
4
5
6
en 7
~ 8
•••• 9
+ 10
~11
12
°13
ci 14
z 15
16
17
18
19
20
I;

0

1

18
6
3
1

3
27
26
7
5
1
1

2

12
14
4
4
2
5
1
1

°

3

1
4
5
3
1
2
1

1

4

5

6

7 .

2
1
1
2

1
1
1
1

1
1
28

71

43

17

7

1
3

1

2

L

21
33
41
22
10
9
8
10
3
3
3
4
0
1
1
1
0
1
0
1
172

I
\J1

w
•....•
I

�Table 4. Frequency of total groups of buck and doe mule deer recorded on 10 all-day, walking
Cache 1a Poudre winter range during each of six sampling periods, 1962-1965.

November 21, 1963 to December 14,
1963

Sample Dates:

Sample Dates: November 29, 1962 to December
18, 1962

No. Bucks

No. Bucks

CIl

~

()

::I
I'CI

+
CIl
QJ

0

.

0
0

Z

0

1

1
2
3
4
5
6
7
8
9
10
11

86
26
9
2
2
1
2
2

38
20
9
6
2
1
2

z

130

2

3

6
6
5
2
2
2
2
1

4

z

1

124
52
25
13
6
4
6
4
1
1
1

1

237

1

1

Sample Dates:

26

78

December

routes on

2

CIl

~

::I

I'CI

+
CIl
QJ

0

0

0

z

0

1

1
2
3
4
5
6
7
8
9
10
11
12
13

40
24
4
2
1

29
10
10
1
1
1

~

71

52

2

1
4
2
1
1
1
2
1

13

3

4

5

z

1

69
35
19
7
3
2
1
2
2
0
0
0
1

1

141

1
2

1

3

1

3, 1964 to Dec. 17, 1964

No. Bucks

CIl

~

()

::I
I'CI

+
CIl
QJ

0

0

.

0

z

0

1

1
2
3
4
5
6
7
8
9
10

58
45
21
4
2

46
38
16
8
5
4

z

l30

117

2
10
7
6
4
6
3
2

3

1

1
4
2
k
1
2
3
1

39

15

4

2

1

3

~
104
93
45
22
15
11
4
5
3
2
304

-----------------------------------------------------------------------------------------------------------------

I
VI

w

N

I

�Table 4. Frequency of total groups of buck and doe mule deer recorded on 10 all-day, walking routes on Cache
1a Poudre winter range during each of six sampling periods, 1962-1965 (continued).

Sample Dates:

January 3, 1963 to January 27, 1963

Sample Dates:

January 9, 1964 to January 31, 1964

No. Bucks
0

1

1
2
3
4
5
6
7
CIl
Ql
8
0
Q
9
10
+
CIl
11
~u
12
=' 13
p::j
• 14
~ 15
16
17
18
19
20

64
34
25
17
12
4
2

9
5
10
4
2
5
2
1

L

1

0

0

2 .3

5
2
2
1

5

6

7

8

9

2
2
1
1

1
1

4

No. Bucks

2
1
3

0

0

1

1
1

1
1

0

0

0

1

0

0

0

0

0

1

158

39

13

8

7

1

0

1

0

1

L

73
44
39
25
16
11
7
4
4
3
0
0
1
1
0
1
0
0
0
1

+

1
2
3
4
5
6
7
8

CIl

9

CIl
Ql

0
Q

~ 10
p::j
=' 11
12
0
z 13
14
15
16
17

0

1

2

41
30
10
10
2
3
1
3

31
8
10
5
4
1
3
2
1

14
4
2
2
2
1
2

1
3
1
1
1
1

4

5

6

7

8

2
1
1
1

1
1
1

1
1

.

L

3

1
1

1

1

1
1
1
1

101

65

30

8

4

6

1

2

228

------------------------------------------------------------------------------------------_.--------------------------

1

L

72
52
25
22
10
9
8
8
2
4
2
0
1
0
1
1
1

218

I
VI

w
w
I

�-534-

Table 4. Frequency of total groups of buck and doe mule deer recorded on 10
all-day, walking routes on Cache 1a Poudre winter range during each of six
sampling periods, 1962-1965 (continued).

Sample Dates:

January 13, 1965 to January 27, 1965
No. Bucks

rJl
Q)

0
Q

+
rJl

.!G
()

;:l
I'Q

.

0

z

0

1

2

1
2
3
4
5
6
7
8
9
10
11
12
13
14

51
47
44
10
8
6
3
1

22
7
7
7
8
2
2
2
1

23
5
2
2
1
1
1

L

170

3

4

5

6

7

7
5
2
1

1
1
1
1

1
1
1

1

1
2
58'

38

10

7

2

3

1

L

73
77
63
24
18
12
8
6
3
2
0
0
1
2
289

Table 5. Frequency of total groups of doe and fawn mule deer recorded on 12
all-day walking routes on Cache 1a Poudre winter range, November 28, 1975 to
January 6, 1976.
No. Fawns

rJl

r..~
+
rJl
Q)

0
Q
0

z

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20

0

1

31
7
1
1

5
21
15
17
1
1

1

2

1
10
8
11
2
1
2
1
1

3

1
2
3
2
2
1
2
1
1

4

5

7

1

1
1

1
1

1
1

1

2

1

1

0
0
0
11
0
1
0
2

2

3

166

1
1

41

61

37

15

5

2

L

36
29
27
29
12
6
5
4
2
4
4
1
1

1

27
L

6

�-535-

Table 6. Frequency of total groups of buck and doe mule deer recorded on
12, all-day walking routes on Cache 1a Poudre winter range, November 28,
1975 to January 6, 1976.
No. Bucks
0

1

2

1

54

14

2

23

8

2

3

26

6

2

4

8

4

1

13

5

4

1

2

7

6

3

1

7

2

2

8

1

1

9

1

1

3

4

E

68
33
1

35

4
1

5
3

Ul
Q)

0
t=I

+
Ul

10

1

3

1

1

~

::J

r::Q

11

. 12

1

1

0

0

:z;

13

1

14

1

1

2
1
0

23

1

1

24

1

1

123

42

8

4

1

178

�Table 7. The fawn and doe data by individual
1a Poudre winter range, 1962-65.

Fall
1962
fi a
nia

route and sex count periods and their relevant statistics

Winter
1963
fi
ni

fi

ni

Winter
1964
fi
ni

1
0
3
19
47
6
9
13
35
21

·4
2
7
45
88
12
27
33
87
58

43
7
11
55
48
10
18
22
44
13

Fall
1963

on Cache

fi

ni

Winter
1965
ni
fi

44
18
6
50
61
14
18
15
56
42

119
52
25
130
151
37
52
44
128
97

41
49
11
72
52
15
27
26
50
26

Fall
1964

Routes:
Washout
Sevenmi1e
Bennett
Kelly
Elkhorn
West Hewlett
Eas t Hew1et t
Young's
Livermore
Seaman

24
5
19
83
168
63
100
47
83
74

9
2
7
40
76
25
44
17
33
40

110
55
16
153
156
62
136
39
161
96

41
22
5
66
66
29
52
17
67
41

136
23
30
148
117
25
51
67
120
40

113
147
33
t88
134
48
82
75
121
67

I
VI

w

'"

I

1\

R
1\

1\

SE R
Variance

666

293

Total

984

406

363

154

757

271

841

324

1,008

369

.2729

.1927

.311+8

.3165

.3277

.2621

.0206

.0199

.0213

.0291

.0244

.0303

Fall 1963

Winter 1964

Fall 1964

Winter 1965

10
79
1,005

43
64
284
162

55
43
54
35
150

39
32
69
54
92
105

and Covariance Matrix for ~ =

Count Period

Fall 1962

Fall 1962
Winter 1963
Fall 1963
Winter 1964
Fall 1964
Winter 1965

274

PF (X 10-6) b

Winter
31
74

1963

------------------------------------------------------------------------------------------------------------------

�Table 7. The fawn and doe data by individual
la Poudre winter range, 1962-65 (continued).

Intervals

route and sex count periods and their relevant statistics on Cache

95% Confidence
for Differences in PE Fall Versus Winter

Year

Sample Sizes
(No. Routes) Averaged for 6 Count Periods

if

Difference

Percent
1962-63
1963-64
1964-65

a
b
c
d

+
-

.0273
.0662
.0192

fi = number of f awn s ,
Procedures

.0333
.0480
.0165

+
+

c

05
10
20

95

Confidence Level (I-X)
90
80

29
8
2

21
6
2

13
4
1

ni = total does + fawns.

according to Cochran (1963:182).
I

Deviation from true ratio.

\J1

Add and subtract to difference

for confidence interval.

Table 8. The buck and doe data by individual
1a Poudre winter range, 1962-65.

Routes

~oJashout
Sevenmi1e
Bennett
Kelly
Elkhorn
\-JestHewlett
Eas t 1-lew1et
t

W
-..J

I

route and six count periods and their relevant statistics on Cache

Fall
1962
bi a

ni a

bi

ni

bi

ni

bi

ni

bi

ni

bi

ni

10
1
1
20
32
14
12

25
4
13
63
124
52
68

16
16
0
30
21
13
0

85
49
11
117
111
97
22

3

6
4
6
32
62
25
26

25

2
2
6
21
7
6

118
24
34
128
96
59
24

43
33
14
36
33
14
9

ll8
67
33
116
129
43
32

12
46
14
34
35
9

84
144
36
150
117
58
44

Winter
1963

Fall
1963

Winter
1964

8

15
35
27
14
9

Fall
1964

Winter
1965

II

�Table 8. The buck and doe data by individual
1a Poudre winter range, 1962-65 (continued).

route and six count periods and their relevant statistics

Winter
1965

Fall
1964

Winter
1964

Fall
1963

Winter
1963

on Cache

Routes

Fall
1962
bi a

ni a

bi

ni

bi

ni

bi

ni

bi

ni

bi

ni

Young's
Livermore
Seaman

10
20
20

40
70
54

2
24
16

35
118
71

3
24
22

9
76
59

26
40
26

59
116
53

16
23
31

50
95
86

3
30
33

58
101
74

R

.2729

.1927

.3148

.3165

.3277

.2621

SE R

.0206

.0199

.021~

.0291

.0244

.0303

Fall 1963

Winter 1964

Fall 1964

Winter 1965

A

Variance

A

and covariance matrix for R = RB (X 10-6)
Fall 1962

Winter 1963

b

I

V1

w

Fall 1962
Winter 1963
Fall 1963
Winter 1964
Fall 1964
Winter 1965

Intervals

95% Confidence
for Differences in PB Fall Versus Winter

Year
1962-1963
1963-1964
1964-1965

il

Difference
.0802
-.0017
.0656

+
+
+

Percent c

95

Confidence Level (%)
90

80

05
10

118
30
8

84
21
6

51
13
4

20

bi = number of bucks, ni = total bucks + fawns.

b

Procedures

according

c

Deviation

from true ratio.

d

Add and subtract to obtain confidence

to Cochran

Sample Sizes
(No. Routes) Averaged for 6_Count Periods

.039
.059
.072

a

(1963:182).

interval.

59
248
316
428
81
921

95
116
95
4
596

-69
-65
198
848

152
-27
453

217
397

423

00
I

�Table 9.

Total deer classified

into buck, fawn and doe categories and related statistics (1975-76) route counts.

All Deer in GrouE Classified
Outside
Inside
500'
500'
Combined
Width
Width

At Least One Unclassified
Deer Within GrouE
Outside
Inside
500'
500'
Combined
Width
Width

Inside
500'
Width

All GrouEs Combined
Outside
500'
Combined
Width

Bucks

28

28

50

10

14

24

32

42

74

Fawn

69

122

191

18

37

55

87

159

246

Doe

138

216

354

36

80

116

174

296

470

.333

.361

.351

.333

.316

.322

.333

.349

.3436

.0181

.0136

.0115

.1023

.0247

.0400

-

-

.0120

A

PF
1\ /\
SE RF

I

1..11

.537

.523

.500

.565

.540

.500

.463

.484

.500

PB

.1375

.1148

.1238

.2174

.1489

.1714

-

-

.1360

A "
SE PB

.0307

.0235

.0218

.1115

.0270

.0406

-

-

.0225

Buck/doe

.159

.130

.141

.278

.175

.207

.184

.142

.157

Fawn/doe
A

w

\0

I

�-540-

Table 10. Comparisons of the proportion of fawns (PF) and bucks (PB) among
groups of mule deer completely or incompletely classified and located
inside or outside the 500 ft strip on the route counts, 1975-1976.

Difference
(Category 1Category 2)

SE
(Difference)

Groups - all deer classified
(inside 500' versus outside 500')

-.0276

.0208

-1.33

Groups - at least one deer
unclassified (inside 500'
versus outside 500')

.0171

.0989

.17

Groups - all deer classified
versus groups - at leas t
one deer unclassified

.0286

.0435

.66

Groups - all deer classified
(inside 500' versus
outside 500')

.0227

.0304

.75

Groups - at least one deer
unclassified (inside 500'
versus outside 500')

.0685

.1163

.59

Groups - all deer classified
versus groups - at least
one deer unc1assfied)

-.0122

.0156

.78

Comparison

�-541-

Table 11. Number of randomly determined routes to be within X percent of
the proportion of fawns (PF) or proportion of bucks (PB) at (I - y) confidence level, Cache la Poudre drainage winter range.
PF 1975-76

PB 1975-76

Confidence Level

Confidence Level

x

95

90

80

x

95

90

80

05

23

16

10

05

505

356

217

10

6

4

3

10

127

89

55

20

2

1

1

20

32

23

14

The individual group counts are summarized in Table 12 for both the 1974-75
and 1975-76 quadrat counts. The raw data and ratio computations of PF mule
deer doe and fawn and buck and doe groups are tabulated as follows: doe and
fawn groups within low density quadrats, 1974-75 (Table 13); doe and fawn
groups with high density quadrats 1974-75 (Table 14); buck and doe groups,
1974-75 (Table 15); doe and fawn groups, 1975-76 (Table 16); and buck and doe
groups, 1975-76 (Table 17).
In a previous analysis (Anderson and Bowden 1975) using the regression approach
on the 1974-1975 data a significant difference was found in the PF values
between the high and low deer density strata (P &lt; 0.01). The larger standard
errors associated with the ratio estimates indicate a significant difference
in the PF values for 1974-75 only at the 10 percent significance level. In
the 1975-76 data the PF value was again higher in the low density quadrats
than the high density quadrats but with the sample size reduced nearly in
half over 1974 no significant difference is indicated. For the PB values, a
significantly higher (P&lt; 0.01) proportion of bucks was found on the low density
strata in 1974-75 using the ratio estimators. Again although in 1975-76 the
proportion of bucks on the low density quadrats was higher than the proportion
obtained on the high density quadrats (nearly twice the high density PF), the
difference was not significant. Since the quadrat sampling was proportional
to the number of quadrats a pooled data estimate is still reasonable even with
the observed significant differences.
Comparisons of Estimates Obtained by the Route and Quadrat Sampling Plans
Comparisons of the 1975-76 estimates of PF and PB between the route and quadrat sampling plans do not indicate any significant differences. The route
estimates (±SE) for PF and PB were respectively,
.3436 (+ .012) and .1360
(+ .023), while the quadrat estimates were respectively, -.3607 (± .027) and
.1818 (± .035).

�Table 12. Frequency of total groups counted on 1/4-mi1e2 randomly-selected quadrats subjectively assigned
to high and low deer density strata, Cache 1a Poudre winter range.

Sample Dates: November 22, 1974 to January 9, 1975
Low Deer Density Quadrats

High Deer Densit~ Quadrats

0
1
2

3
4

19
16
3

5
6
7

1

8
9
+ 10
~ 11
8 12
. 13
0
z 14
15
16
17
18
19

1

L

40

(I)

rz..~

1

2

18
14
3
3
3

9
11
3
1

1

2

1

N = 50

N = 56

No: Fawns

No. Fawns
4
2
3

3

4

5

I:
19
34
23
17

3
1
1
2

1

1

1

1

1
1

1
2

(I)

+

4
3
3
1
2
1

C1J

1

11
4

7
8

3
4
5

rz..~

6
9

0

4
3

7

0
A

I:

2
1
3

11
12
11
6
4
3
2
1
0
2
1
53

1
3
2

1
1
1

6

(I)

5

. 98

2

1
1

0

z 10
11
L

15

22

8

5

0

0

1

1

2

0
0
0
0

43

28

9

3

1
2

1
125

--------------------------------------------------------------------------------------------------------

I
V1

~

N
I

�Table 12. Frequency of total groups counted on 1/4-mi1e2 randomly-selected quadrats subjectively
to high and low deer density strata, Cache la Poudre winter range (continued).

Sample Dates:

Hi~e~r

November 28, 1975 to January 6, 1976

p~nsity Quadrats

Low Deer D8nsity Quadrats

No. Bucks

CIl

.-'"

o

::l

P=l

+
CIl
Q)

0
Q

.

0

z

0

1

1
2
3
4
5
6
7
8
9

15
9
2
1

3
2
1
1

L:

31

2

No. Bucks
3

L:

121
tJ
P=l

1
1

18
11
4
3
0
2
2
1
2

2

43

1
1

2
2
1
8

2

assigned

::l

+
CIl

Q)

0
Q

01

z

0

1

1
2
3
12

2
1
1

3
1
1

E

4

5

2

0

3

L:

1

5
2
2
1

1

10

-----------------------------------------------------------------------------------------------------------

I
.j::Ln

w
I

�Table 12. Frequency of total groups counted on 1/4-mi1e2 randomly-selected quadrats subjectively
to high and low deer density strata, Cache 1a Poudre winter range (continued).

assigned

Sample Dates: November 22, 1974 to January 9, 1975

Low Deer Density Quadrats

High Deer Density Quadrats

tIl

~(J

;:l

~

+
tIl

Q)

0

A

.

0

z

N = 56

N = 50

No. Bucks

No. Bucks

0

1

2

1
2
3
4
5
6
7
8
9
10
11
12
13
14

40
36
9
3
6
2
1
2
1

9
5
5

2

E

103

1

3

1

4
2

1
1
1

1

2

1
26

6

1

0

1

1
2
3
4
5
6
7

17
12
3
3
2

10
4
2
3

E

103

E

49
43
14
5
10
3
4
3
2
0
2
0
0
1

]
(J

;:l

~

+
tIl
Q)

8

.

2

3

E

1
11

1
1

1

27
17
5
6
3
2
3

26

6

1

136

1

1

0

Z

136

----------------------------------------------------------------------------------------------------------

I
V1
.po.
.po.
I

�Table 12. Frequency of total groups counted on 1/4-mi1e2 randomly-selected quadrats subjectively
to high and low deer density strata, Cache la Poudre winter range (continued).

Sample Dates:

00

~

::l
J:J:I

+
00
Q)

0
Q

.

0

Z

November 28, 1975 to January 6, 1976

High Deer Density Quadrats

Low Deer Density Quadrats

No. Bucks

No. Bucks

0

1

1
2
3
4
5
6
7
8
9

15
9
2
1

3
2
1
1

~

31

2

3

2
2
1

8

2

~

0

1

1
2
3
12

2
1
1

3
1
1

~

4

001

~

::l
J:J:I

18
11
4
3
0
2
2

1
1

+
00
Q)

0
Q

~I

5

1

2
3
00
4
~
~ 5
+ 6
7
00
Q)
8
Q
9
10
z 11
12

1

9
2

4
8
6
2
1

2

1
2
1

2

2

43

4

5

1
1

5

10

I

I

~

2
0
1
1

1

5

1

\J1

13
10
8
4
3

1
1

22

1

55
2
2
1.

No. Fawns

1

12

~

.j:'-

1

1

0

3

1

1
1

1

°
01

~
--.

3

2

\J1

No. Fawns
0

assigned

0

1

1
1

1

45

0

00'

~
~

+
00
Q)

0
Q

.

~I

1
2
3
4
14

1

~

1

1

2

1
1
1

1
1
1

3

4

5

~
1

3

3

2
2
2

0

0

1

1

1

8

�-546Table 13. Raw data and ratio computations of PF for mule deer doe and fawn
groups counted on 56 randomly-selected, 1/4 mi1e2 (one-half mile per side)
low deer density quadrats on Cache 1a Poudre winter range, November 22, 1974
to January 9, 1975.
No. Does + Fawns
D + F

No. Fawns
F

No. Does + Fawns
D + F

No. Fawns
F

5
10
0
0
0
0
11
12
0
3
0
0
0
22
2
0
0
8
17
3
7
5
0
0
0
0
10
3
0
0
0
0
8
0
2
15
6
3
1
0
0
0
0
0
0
0
0
11

1
4
0
0
0
0
5
3
0
1
0
0
0
8
1
0
0
3
7
1
3
3
0
0
0
0
5
1
0
0
0
0
2
0
0
2
3
2
0
0
0
0
0
0
0
0
0
4

5
4
1
1
8
0
0
0
183

3
2
0
0
4
0
0
0
68

Computat ions
N

56

E«D
+ F)
183
E (D + F)2
1,999
E (F)
68
E (F)2
= 286
E Fi(D + F) =
721

EF
E(D

+ F)
.0282

.3716

�-547Table 14. Raw data and ratio computations of PF for mule deer doe and fawn
groups counted on 50 randomly-selected, 1/4-mile2 (one-half mile per side)
high deer density quadrats on Cache la Poudre winter range, November 22,
1974 to December 31, 1975.
No. Does + Fawns
D + F

No. Fawns
F

No. Does + Fawns
D + F

No. Fawns
F

28
37
5
11
2
8
6
5
8
0
6
2
4
19
0
3
2
0
1
8
28
5
0
8
9
1
21
21
8
11
41
19
12
9
0
2
2
6
4
5
12
32
17
5
11

11
12
2
2
0
2
3
2
4
0
2
0
1
8
0
2
1
0
0

16
0
0
478

2
0
0
148

2

16

1

11
2
0
4
2
0
5
4
2
3
12
6
4
2
0
0
0
3
2
2
4
5
6
1
5
1
7

Computat ions

N

50
478
9,480
148
954
2,874

(D + F)
(D + F)2
(F)
(F)2
H(D + F)

L:
L:
L:
L:

1\

PF
1\

.3096
1\

SE (PF)

.0193

Pooled Low and High Density Quadrats
1\

PF

=

.3268

.0164

�-548Table 15. Raw data and ratio computations of PB (the proportions of bucks
to buck and does) on randomly-selected, 1/4 mile2 (one-half mile per side)
quadrats on Cache la Poudre winter range, November 22, 1974 to January 9,
1975.

B + D

7
12
0
0
0
0
7
12
0
3
0
0
0
17
1
0
0
8
12
0
2
4
3
0
0
0
0
7
2
0
0

Low Densit~
B
B + D

3
6
0
0
0
0
1
3
0
1
0
0
0
3
0
0
0
3
2
0
0
0
1
0
0
0
0
2
0
0
0

0
0
8
3
15
3
1
2
0
0
0
0
0
0
0
0
8
2
2
1
1
5
0
0
0
2

High Densit~
B
B + D

B

B + D

0
0
2
1
2
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
1

17
29
3
10
2
7
6
3
4
0
4
3
7
11
0
1
1
0
1
7
20
3
0
4
8

0
4
0
1
0
1
3
0
0
0
0
1
4
0
0
0
0
0
0
0
3
0
0
0
1

16
17
7
10
31

0
0
1
2
2

16
10
7
0
2
2
3
0
0
3
4
9
29
13
5
7
2
10
15

B

Computat ions

3
2
0
0
0
0

0
0
0
1
1
1
2
2
1
1
1
1
1

Low Density
N
L: (B + D)
L: (B + D)2
L: B
L: B2
L: B(B + D)

1

56
148
1,386
33
95
319

/\

.2230

PB
/\

/\

SE (PB)

.0317

High Density
N
L: (B + D)
L: (B + D)2
L: B
L: B2
L: B(B + D)
/\

PB

50
371
5,689
41
93
548

.1105

/\

SE (~B)

.0175

Pooled Data
A

PB

SE (PB)

.1426
.0178

�-549-

Table 16. Raw data and ratio computations of PF (the proportion of fawn
to fawns + does) on 58 randomly selected, 1/4-mi1e2 (one half mile per side)
quadrats on Cache 1a Poudre winter range, November 28, 1975 to January 6,
1976.

Low Densit:l
F + D
F

High Densit:l
F + D
F

1
0
4
2
21
7
5
3
2
2
Plus 20 Quadrats
with
0
0

1
0
5
1
4
2
1
0
1
1
4
1
7
0
10
5
3
1
5
2
21
6
10
3
3
2
2
1
4
2
5
1
2
1
14
5
6
2
5
2
19
6
12
4
3
3
3
1
Plus 9 Quadrats
with
0
0

Computations

Low Densit:l
N
L: (F + D)
L: (F + D)2
2: F
L: F2
L: FD

25
33
487
14
66
174

1\

.4242

PF
1\

1\

SE (PF)

.0759

High Density
N
L: (F + D)
L: (F ± D)2
L: F
L: F2
L: F (F + D)

33
150
1,622
52
188
526

/I.

.3467

PF
1\

"
SE (PF)

.0289

Pooled Data
A

.3607

PF
/\

/\

SE (PF)

.0267

�-550-

Table 17. Raw data and ratio computations of PB (the population of bucks
to bucks and does) on 58 randomly-selected 1/4-mi1e2 Cone-half mile per side)
quadrats on Cache la Poudre winter range, November 28, 1975 to January 6,
1916.

Low Density
B + D

High Density
B

1
1
2
1
3
1
17
3
2
0
1
1
1
1
Plus 18 Quadrats
with
0
0

B

+D

B

2
6
2

1
2
0
0
0

1

3
7
0
6
1
2
0
4
1
21
6
7
0
3
2
1
0
2
0
4
0
1
0
9
0
5
0
3
0
14
1
8
0
4
2
1
1
Plus 10 Quadrats
with
0
0

Computations

Low Density
N

25
(B + D)
(B + D)
(B)
(B)2
B (B + D)=

L
L
L
L
L

8
14
27
309
59

1\

.2963

PB

tE (PB)
"

.0939

High Density
N
L
L
L
L
L

(B + D)
(B + D)2
(B)
(B)2
B (B + D)=

P
"B

.1552

1\

33
18
54
116
1,072
184

1\

SE (PB)

.0417

Pooled Data
A

PB

.1818

1\

"
SE (PB)

.0355

�-551-

Sample size calculations can be made from each of the corresponding sets of
data. The sample sizes or number of quadrats required to be within X percent
of PF or PB at (l-y) confidence level is given in Table 18. Only the 1974-75
calculations are reported due to the larger sample sizes involved in estimating
the variance needed in the sample size calculation.

Table 18. Number of randomly determined quadrats to be within X percent of
the proportion of fawns (PF) or proportion of bucks (PB) at (l-y) confidence
level, Cache la Poudre drainage winter range. (Finite population correction
factor was not applied).

PF 1974-75

PB 1974-75

Confidence Level

Confidence Level

X

95

90

80

X

95

90

80

05

410

289

175

05

2,538

1,787

1,083

10

103

73

44

10

635

447

271

20

26

18

11

20

159

112

68

One of the advantages of the quadrat sample design is that ffilnlmumbound estimates of the population total may be made from the total deer on each sampling
unit using the stratified sampling estimate of a population total. The estimates along with their standard errors for the Poudre and the Colorado State
University Research Foundation Maxwell property and vicinity are given in Table
19.
In order to incorporate this advantage of the quadrat sample plan into the
route plan, the estimate of density is obtained from the counts of total deer
within the 500 foot width centered on the route transect. A ratio estimator
is used to obtain the density since the areas sampled by routes varied. The
density obtained in the 1975-76 route census is 43.17 deer/sq. mi., with a
standard error of 11.93 for the Cache la Poudre winter range.
RESULTS AND DISCUSSION
Of the three models applicable to fawn:doe and buck:doe ratios (binomial,
regression estimator, and ratio estimator) the latter model was most satisfactory. The ratio estimator
is an unbiased estimator of PF and PBAwith an
estimab*e if larger. standard error than for regression estimator S. Moreover,
unlike S, ~ does not require the critical assumption that fawns and bucks occ~r
in equal proportions regardless of group size. The utility of the estimator R

R

�-552-

Table 19. Estimated a~n1mum densities of mule deer on the Cache la Poudre
winter range and the C.S.U.R.F. Maxwell Ranch property and vicinity based
on ground counts of deer on randomly-selected, proportionally allocated,
and subjectively stratified, 1/4-mile2 quadrats during the winter.

Strata

Total No.
Units
(1/4 mi2)

Population
Estimate

SE
(Pop. Est.)

Density/
Sq Mile

SE
Density

Cache La Poudre Winter Range
November 22, 1974 to January 9, 1975a
Low deer density

229

958.66

161.66

16.75

2.82

High deer density

199

2,168.7

244.6

43.59

4.92

Total

428

312.36

293.2

29.23

2.74

...
b
Maxwell Ranch and V 1c1n1ty
November 18, 1974 to November 21, 1974
Low deer density

30

171.9

94.8

22.92

12.64

High deer density

31

237.5

160.6

30.65

20.73

Total

61

409.4

186.5

26.85

12.23

Cache la Poudre Winter Range
November 28, 1975 to January 6, 1976c
Low deer density

229

476.3

267.9

8.32

4.68

High deer density

199

1,278.6

249.7

25.70

5.02

Total

428

1,754.9

366.3

16.40

3.42

Maxwell Ranch and Vieinitx

d

November 21, 1975 to November 26, 1975
Low deer density

30'

94.3

59.0

12.57

7.86

High deer density

31

276.4

125.8

35.66

16.23

Total

61

370.7

138.9

24.31

9.11

a
b

c

Estimates based on 106 quadrats.
Estimates based on 15 quadrats.

Estimates based on 58 quadrats.
d Estimates based on 14 quadrats.

�-553-

is shown by: (1) the significant (P &lt; 0.01) decrease in PF from 1962 to 1965
detected by R was not detected (P &lt; 0.05) when analyzed by chi square (Anderson 1965); (2) sampling intensities can be practically achieved at adequate
levels of precision by the use of R.
The 293 deer recorded within the 500 ft strip width of the 12 routes compared
to 258 deer within the quadrats during 1975-76 was projected to 43.17 deer/
mile2 on the routes and 16.40 deer/mile2 on the quadrats. This discrepancy
suggests that the dimensions of the strip were inaccurate. By relying primarily on visual estimation, there is a good possibility of gross error in estimating the 500 ft wide strip, particularly under conditions of bright sun
and snow. It may be possible to improve the accuracy of the strip width by
using a range finder. Further tests of the route and quadrat sampling plans
relative to the discrepancy in estimated densities are recommended.
Even though the route counts and quadrat counts yielded 780 and 258 deer,
respectively, their PF and FE were nearly identical, but the standard errors
of the quadrat estimates were somewhat larger than the route estimates. Of
the two sampling plans, randomly determined route counts appears superior because PF can be estimated at fairly high levels of precision in far less time.
In addition, it has been shown that neither distance from the strip center or
the inclusion of groups containing one or more unclassified deer had Significant effects on either the PB or PF obtained on the route counts.
The significant (P &lt; O.01) trend in decreasLng PF, 1962-65, occurred concurrently with approximately a 40 percent reduction in annual precipitation and
mean yields of major browse species and significant (P &lt;0.01) changes in such
indices of stress as leaner deer, reduced mean erythrocyte counts (anemia),
and increasing mean neutrophil:lymphocyte ratios (Anderson et al. 1972:590).
While PF decreased there was a significantly increasing trend in PB during the
winter, but the contributor to these differences was the extremely low PB
value in 1963. Annual blood cellular values changed most drastically during
1963 (Anderson et al. 1970:400). We speculate that there is the possibility
that drouth reduced food supply, hence nutritional levels and fawn production
or survival. No similar speculative pathway can be suggested for the low PB
during 1963.
Route counts revealed that, in general, the 1975-76 PF was significantly less
than during 1962-65. Because of the lack of information from 1965 to 1974,
possible causal factors for this decrease will not be dealt with here.
Both PF and PB were higher on low density quadrats during 1974-75 and 1975-76.
Relative to PF, this finding seems to conform to the hypothesis that in deer
the "highest net numbe r of young per unit of breeding stock" are produced at
low population densities (Gross 1969:385). However, it seems unlikely that,
in a largely migratory population on winter range, such a phenomenon could
occur on only 1/4 mile2, particularly since low and high density quadrats
were frequently adjacent. Relative to PB, we speculate that during breeding
season observations such as ours, bucks might be more readily perceived on
low density areas because of the presumably greater activity bucks must expend
to locate does within areas of relatively low densities.

�-554-

LITERATURE CITED
Anderson, A. E., and D. E. Medin. 1965. Population density and structure.
Work Plan 4, Job 1, Job Completion Report. Pp. 405-427. In Colorado
Game, Fish and Parks Dept. Game Research Report, January. Part 3.
327-501 p , (Processed).
Anderson, A. E., D. E. Medin, and D. C. Bowden. 1970. Erythrocytes and
leukocytes in a Colorado mule deer population.
J. Wildl. Manage. 34(2):
389-406.
Anderson, A. E, D. E. Hedin, and D. C. Bowden. 1972. Indices of carcass fat
in a Colorado mule deer population.
J. Wildl. Manage. 36(2):569-574.
Anderson, A. E., and D. C. Bowden. 1975. Evaluation of herd structure
methodology.
Pp. 475-499. In Colorado Div. Wildl. Game Research Report,
July. Part 2. 191-504 p.
Caughley, G. 1974.
557-562.

Interpretation

of age ratios.

J. Wildl. Manage. 38(3):

Cochran, W. G. 1936. 'The chi square distribution for binomial and Poisson
series with small expectations.
Annals Eugenics 7:207-217.
1954. Some methods for strengthening
Biometrics 10:417.

the common chi square tests.

Dasmann, R. F., and R. D. Taber. 1956. Determining structure in Columbian
black-tailed deer populations.
J. Wildl. Manage. 29(1):78-80.
Gross, J. 1969. Optimum yield in deer and elk populations.
Wildl. Nat. Resou. Conf. 34:372-387.

Trans. N. Amer.

Smith, R. H., T. J. McMichael, and H. G. Shaw. 1969. Decline of a desert
deer population.
Arizona Game and Fish Dept. Wildl. Digest Abstr. 3:1-8.
Swank, W. G. 1958. The mule deer in Arizona and chaparral and an analysis
of other important deer herds. Arizona Game and Fish Department Wildl.
Bull. No. 3:1-109.
Taber, R. D., and R. F. Dasmann. 1958. The black-tailed deer of the chaparral:its life history and management in the north coast range of California. California Dept. Fish and Game, Game Bull. No. 8:1-163.

Prepa red by _--4,.cZLeb...a::;:.:=::.oa../,-:-~e_. ~t2twt=....:::w."""'M4~:caL,","""","
Allen E. Anderson
Wildlife Researcher

_

�July, 1976

-555JOB PI:OGl{ESSREPORT

State of
Project No.

~CO~LO=RADO

_

w-ss-a-ai

Work Plan No.

Deer-Elk Investigations

------------

------

20

Job No.

Job Title Brucellosis-Leptospirosis

1
-------------------------------

Survey of Big Game Animals in Colorado

Period Covered: April 1, 1975 through March 31, 1976
Personnel:

William J. Adrian, Marilyn Stevens, Sandra Pool, Carol Ann Weinland,
Richard Ford and Gene Grenerd.

ABSTRACT

During the 1975 season we checked deer for brucellosis and five species of
leptospirosis (leptospira canicola, grippotyphosa, hardjo, icterohemorrhagiae,
and pomona). A total of 7,501 blood kits were mailed to deer hunters.
Returns
with blood were received at the Cooperative Brucellosis Laboratory in Denver from
17 percent of the hunters who were mailed kits. Of those kits received, 61
percent were not usable. Thus the total sample of blood kits analyzed was 508.
All samples analyzed were negative for brucellosis and leptospirosis (all
species). The Center for Disease Control (Dept. HEW) is analyzing the samples
for exposure to Colorado tick fever and Colorado State University, Department
of Microbiology is checking for bluetongue.
The results of Colorado tick
fever and bluetongue are forthcoming.

��-557-

BRUCELLOSIS-LEPTOSPIROSIS SURVEY
OF BIG GAME ANIMALS IN COLORADO
William J. Adrian

P. S. OBJECTIVE
To survey big game populations within the State of Colorado for the presence or absence of brucellosis and leptospirosis.

SEGMENT OBJECTIVE
1.

Survey Colorado's deer populations for brucellosis and leptospirosis
during the annual hunting season.

METHODS AND MATERIALS
Blood sample kits were mailed to deer hunters with the request they draw
a blood sample from their kill and return the kit to the Colorado Cooperative
Brucellosis Laboratory in Denver, Colorado for processing. A maximum of 300
sampling kits were sent to hunters in each deer unit. If more than 300
permits were issued for one unit the names were randomly selected.
The kit consists of a cardboard mailer, blood tube and stopper, plastic bag,
rubber band, paper towel and instructions for obtaining the sample and
mailing the kits.

RESULTS AND DISCUSSION
The Wildlife Research Laboratory in cooperation with the Cooperative Brucellosis Laboratory in Denver has, in 1975, completed its seventh annual survey
for brucellosis and leptospirosis in Colorado's big game populations.
Brucellosis is a specific contagious disease which primarily affects cattle,
swine, and goats. It can, however, affect many other animals including elk,
deer, antelope and man (undulant fever, Malta fever and Bang's Disease). It
is caused by bacteria of the Brucella group and is characterized by abortion
in the female, and to a lesser extent, inflammation of the testes and infection of the accessory sex glands in the male and infertility in both sexes.
Leptospirosis is a febrile (feverish) disease caused by certain species of
leptospirae. The disease occurs with various manifestions in dogs, cattle,
pigs, sheep, goats, horses, and wildlife and is transmissable to man.
During the 1975 season we checked deer for brucellosis and five species of
leptospirosis (leptospira canicola, grippotyphosa, hardjo, icterohemorrhagiae,
and pomona).
A total of 7,501 blood kits were sent to deer hunters and we received 1,301
blood samples (17%) of which 793 (61%) were badly hemolyzed and were not

�-558-

usable. We also received 592 (8%) empty kits.
tested were negative.

Again in 1975, all samples

A summary of hunter response to this survey since 1968 is presented in
Table 1. The number of returned kits with blood is excellent when the percent return is adjusted for hunter success - thus showing a successful hunter
is a cooperative hunter.

Table 1. Summary of hunter response to the brucellosis-leptospirosis
from 1968-1975.

survey

Year

Species

Kit
Mailed

Blood SamEles
Number Percent

1968

Antelope

3019

1463

1969

--------_ ... _----------

1970

Elk

9930

2338

23.5

2021

20.4

4359

43.9

1971

Antelope

4590

2721

59.3

763

16.6

3484

75.9

1972

Deer

2500

774

30.9

201

8.0

975

39.9

1973

Elk

7502

1231

16.4

Deer

804

153

18.2

Antelope

630

273

43.3

1974

Antelope

3995

2044

51.0

214

5.4

2258

56.5

1975

Deer

7501

1301

17.3

592

7.9

1893

25.2

40507

12298

30.36

4043

9.98

16341

40.3

Totals

48.5

Returned EmEty
Number Percent

252

8.3

Totals
Number Percent

1715

56.8

NO MAIL SURVEY ---------------------------

The sera titer data are summarized in Table 2. All results, including the
1/50 titer for brucellosis and 1/40 titer for leptospirosis, are considered
negative due to the large sample size and its corresponding low incidence of
the suspected titers.
The antigen for the leptospirosis
wise indicated.

test was LeEtospira

Eomona unless other-

The discrepancy between blood samples received and those tested for either
brucellosis or leptospirosis is due to the condition of the samples when
received at the laboratory; this is a highly variable factor from year to

�-559-

year dependin8 on weather and the lag time from kill to laboratory.
Generally, however, approximately one-third of all samples received
are very good to excellent quality and approximately one-fifth are completely hemolyzed and unusable.
In 1975 we also began cooperative studies with the Center for Disease
Control (Dept. HEW) analyzing the remaining serum for exposure to Colorado
tick fever and with Colorado State University, Department of Microbiology
for exposure to bluetongue. We have attempted to check for bluetongue in
the past, but there was some difficulty with the antisera. Dr. Lloyd
Lauerman (Colorado State University), however, believes he has developed
a new antisera which is more effective for detecting the bluetongue virus.
These data along with the Center for Disease Control data will be reported
in the 1977 Progress Report for Work Plan 20, Job 1.

�-560-

Table 2.

Sununary of sera titer data from 1967-1975.
Brucellosis
.Negative
1/25

LeEtosEirosis
Negative
1/10

1/

Species

Year

Deer

1967

17

0

0

17

0

0

1967

107

0

0

0

0

0

1970

128

0

0

110

0

0

1971

774

0

0

53

3

1

1973

116

0

0

115

1

0

1975

508

0

0

508 JJ

0

0

. 1650

0

0

803

4

1

1967

28

0

0

28

0

0

1967

42

0

0

0

0

0

1968

125

0

0

125

0

0

1970

2338

5

1

1087

4

0

1973

693

5

0

692

6

0

3226

10

1

1932

10

0

1967

126

0

0

0

0

0

1967

264

0

0

0

0

0

1969

1463

0

0

229

51

2

1971

2486

0

0

263

8

0

1973

272

0

0

267

4

1974

660

0

1

646 ~/

12 1/

0
3 !!/

Totals

5271

0

1

1405

75

5

GRAND TOTALS

10147

10

2

4140

89

6

Totals

Elk

Totals

Antelope

1/50

1/
Antigen unless otherwise specified was Leptospira
2/
Antigen Leptospira
icterohemorrhagiae.

canico1a, grippotyphosa,

1/
Six icterohemorrhagiae,; 5 hardjo, 1 pomona.
4/
Two hardjo, 1 pomona.

pomona.

hardjo, pomona, and

1/40

�-561-

July, 19'76

JOB FINAL REPORT

State of

~C~o~l~o.r=~a~d~o _

Project No. W-38-R-3l

Deer-Elk Investigations

Work Plan No.

Job No.

Job Title

21
---------

Effects of Coal Strip Mining on Deer - Preparation

Period Covered:
Personnel:

1
of Study Plan

April 1, 1975-1976

Bertram D. Baker

ABSTRACT
No work was accomplished on this job in the past segment because the Project
Leader was reassigned to non-Federal Aid duties for most of the segment.
This job is being discontinued if or until the vacated position is returned
to the Colorado Division of Wildlife Research Section.

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                  <text>October;

-1-

1976

JOB PROGRESS REPORT

State of

---------COLO~\DO
~~-----------

Project No.

W-88-R-2l

Migratory

Work Plan No.

1

Job No.

Job Title

Production

Waterfowl

Period Covered:

Bird Investigations

---- 1

Surveys

April 14, 1975 to June 30, 1975

Personnel: C. Bryant and staff, Monte Vista National Wildlife Refuge; F. N.
Folks, Utah State Division of Wildlife Resources; R. Blumberg, R. Clark,
W. Dolezal, H. Funk, B. Goforth, R. Hopper, J. Lorentzson, R. Lowry, J.
Rauch, W ..Russell, G. Saville, R. Velarde, R. Weldon, S. Steinert, and M.
Szymczak.
ABSTRACT
Water conditions for waterfowl production were considered good in North Park,
Brown's Park and the Cache 1a Poudre and Yampa River Valleys, average in the
South Platte Valley and poor in the San Luis Valley. The number of duck
breeding pairs reached a record number 78,345 in 1975; about 22 percent
above the 1974 level and 38 percent above the long term average.
Increases
in the number of breeding pairs were recorded in all the areas except the
San Luis Valley and Yampa River Valley. Blue-winged teal, cinnamon teal,
green-winged teal, shoveler, wigeon and redhead all showed significant increases in comparison to 1974 levels. The post-nesting season population
of Canada geese in northwest Colorado is estimated to be 33 percent above
the long term average. Canada goose production in north-central Colorado
is up nearly 17 percent from 1974, but down 8 percent from the 1969-74
period, with the total number of adult geese observed up about 23 percent
from 1974 but down 10 percent from the 1969-74 period.

��-3-

WATERFOWL PRODUCTION SURVEYS
Michael R. Szymczak

P. S. OBJECTIVE
To formulate waterfowl harvest regulations for Colorado.

SEGMENT OBJECTIVE
To estimate the number of duck and goose breeding pairs, by species, in
each of Colorado's major waterfowl breeding areas.

METHODS AND MATERIALS
Present duck breeding pair and production surveys consist of a breeding
pair inventory of only major production areas.
The 1975 duck breeding pair surveys were conducted during the period of
May 19 to June 30. Surveys in North Park and the Cache la Poudre and
South Platte Valleys were conducted exclusively from the air. Ground
counts were made in the Yampa Valley. In the San Luis Valley intensive
aerial counts were adjusted for visibility by air-ground comparison
studies. Pair estimates for the Monte Vista National Wildlife Refuge
in the San Luis Valley were obtained from nesting transects. All survey
methods and sample areas for ducks remained the same as in previous years.
No duck breeding pair counts were made in Brown's Park, therefore, figures
presented are based on averages for the 1971-74 period.
Surveys of Canada goose production were conducted within the period of
April 14 to June 13. Production estimates for Moffat County were made
from as complete a count as possible of hatched or active nests and brood
size. Population estimates of north-central Colorado. were obtained from
counts of goslings and adults conducted from the ground during the period
in which the birds were flightless.
All flying was accomplished with a Cessna 185 aircraft. Areas sampled by
section or block were flown with one observer, while two observers were
used in sampling by transect.
On the basis of these studies, a report is submitted to the Bureau of Sport
Fisheries and Wildlife, which constitutes Colorado's part in the annual
continental cooperative breeding ground survey.

RESULTS AND DISCUSSION
Water conditions were considered good for duck production in all areas
except the San Luis Valley. Spring rains filled most small marshes and
drainage basins in the South Platte and Cache la Poudre Valleys. A heavy

�-4snowpack in addition to a considerable amount of spring moisture will
insure good production and brood survival in North Park, although the
hatch will be later than normal. Although there is very heavy snowpack
in the mountains to the west, the San Luis Valley will be very dry during
the waterfowl production period for the second consecutive year. The
retarded runoff in northwest Colorado will insure that flooding will
have a negative effect on goose production in that area. A heavy freeze
coupled with a snowfall during the peak of egg laying in north-central
Colorado resulted in a ccnsiderable amount of nest abandonment.
The number of estimated duck breeding pairs in Colorado's production
trend areas reached a record high 78,345, nearly 6,000 pairs higher
than the previous record set in 1968. The 1975 total is 22 percent
above the 1974 level and 38 percent above the long term average (Table
1). Major increases above 1974 levels were recorded in North Park and
the South Platte Valley. Numbers in the Cache la Poudre Valley continued at a high level, nearly double the long-term average (Table 1).
The majority of the increase in North Park was composed of green-winged
teal, a phenomenon that also occurred in 1969 and may be related to
retarded dissipation of the snowpack in the mountains surrounding North
Park. The South Platte increase was mainly composed of shovelers and
redheads.

Table 1. Summary of Colorado's duck breeding ground population estimates
in selected areas, 1975.

Area

Total Est. Breeding Pairs
Long Term
1975
1974
Average 1/

San Luis Valley

26,801

29,694

27,332

- 9.7

1.9

North Park 1/

27,134

16,657

15,970

+62.9

+ 69.9

South Platte Valley

14,152

8,457

6,264

+67.3

+125.9

Cache la Poudre
Valley

6,732

5,713

3,361

+17.8

+100.3

Yampa Valley

2,354

2,834

2,803

-16.9

- 16.0

Brown's Park

1,1721/ 1,128

1,029

Totals

78,345

56,759

+21.5

+ 38.0

64,483

Percent Change
From
From Long1974
Term Average

1/
-San Luis Valley and North Park averages are based on results of 1964 through
1974 and 1968 through 1974 surveys, respectively, because of changes in survey
methods utilized prior to those dates. Figures for other areas are 21 year
averages.

2/
-Aerial counts corrected by species from visibility ratios obtained in the
San Luis Valley.

]/
No count made in 1975.

Figure presented is average of 1971-1974 period.

�-5-

It is interesting to note that in a year in which total estimated
breeding populations reached a record high in Colorado, the mallard
population dropped in the San Luis Valley. This resulted in a state
wide reduction in the mallard population of 30 percent from the longterm average (Table 2).

Table 2. Species composition of Colorado's 1975 duck breeding pair
population.
No. of Breeding Pairs
1954-74
1975
1974
Average

Percent Species Composition
1954-74
1975
1974
Average

Mallard

19,090

28,427

27,690

'24.4

44.1

54.7

Bl ue-winged
and Cinnamon
Teal

15,078

11,441

5,238

19.2

17.7

10.4

Gadwall

3,984

8,464

5,396

5.1

13.1

10.7

Pintail

5,213

4,344

3,560

6.7

6.7

7.0

Green-winged
Teal

8,859

565

2,147

11.3

.9

4.2

Shoveler

8,005

3,918

2,378

10.2

6.1

4.7

American
Wigeon

4,868

1,087

884

6.2

1.7

1.7

Redhead

8,048

716

1,978

10.3

1.1

3.9

Other Divers

5,200

1,076

1,346

6.6

8.6

2.7

Totals

78,345

64,484

50,536 l/

Species

1/

Species composition computed from data from all areas for the 20 year
period regardless of changes in survey method.

The total number of geese observed in Moffat County, in northwest Colorado,
1,549 (Table 3) was essentially the same as in 1973. All areas were not
surveyed in 1974. In 1975, estimates for the Green River were up about
200 birds from 1973, while the Yampa estimate declined by about the same
number. The total number of geese observed in Moffat County was 33 percent above the long term average while estimated gosling production was up
25 percent (Tables 4 and 5).

�-6-

Table 3. Number of Canada geese, observed and estimated production,
Moffat County, Colorado, 1975.
Nesting
Pairs

Non -Nes ting
Birds

Total
Adults

Est. No.
Goslings })

Total
Birds

16
16
10

89
54
27

121
86
47

53
86
8

174
172
55

42

170

254

147

401

53
26

166
248

272
300

176
106

448
406

Sub-total

79

414

572

282

854

Little Snake River

19

188

226

68

294

Grand Total

140

772

1,052

497

1,549

Area

Yampa
Craig-Juniper
Juniper-Cross
Lily Park

Spgs.
Mtn.

Sub-total
G reen R"l.ver-21
Brown's Park
Dinosaur Nat1. Monum.

II

- Calculated using average brood size observed and number of successful nests.

1lData supplied by F. Neil Folks, Utah State Division of Wildlife Resources.

Table 4.

Total Canada geese observed, Moffat County, Colorado, 1975.

Area

No. Geese Counted
1956-1974
1975
1974
Average

Yampa River II

401

Percent Change
From 1956-1974
From 1974
Average

424

5.4

Green River
Brown's Park

448

Dinosaur Nat1.
Monum. 1./, 11

406

334

+ 21.6

Little Snake
River 1/ , ]./

294

247

+ 19.0

1,549

1,161

+ 33.4

Totals

326

II

156

- Surveys not conducted in these areas during 1974.

1lNot included in survey until 1970.
llNot included in survey until 1962.

+37.4

+187.2

�-7-

Table 5. Estimated number of Canada goose goslings, Moffat County,
Colorado, 1975.

Area

No. of Goslings
1956-1974
Average
1975
1974

Yampa R'1ver -II

147

Percent Change
From 1956-1974
Average
From 1974

+

136

8.1

Green River
119

+47.9

62

+183.9

Brown's Park

176

Dinosaur Natl.
Monum. 1), 2:..1

106

120

- 11. 7

Little Snake
River

68

79

- 13.9

Totals

497

397

+ 25.2

u. 11

l/Surveys not conducted in these areas during 1974.

l/Not included in survey. until 1970.
1/Not included in survey until 1962.

The results of the 1975 Canada goose production census in north-central
Colorado are presented by individual areas in Table 6. In north-central
Colorado, gosling production was up 17 percent from 1974, but down 8 percent from the long term average (Table 7). A major increase in production
occurred in the Denver area with minor increases over the 1974 level
occurring in the other trend areas (Table 7). The number of adult geese
observed on north-central trend areas was 23 percent-above the 1974 level
but 10 percent below the long term average (Table 8).

Fall Flight Prediction
Duck nesting in all trend areas in Colorado with the exception of the San
Luis Valley should result in excellent production and above average brood
survival. However, production in the San Luis Valley will be extremely
poor and the resulting fall flight from that area will be far below normal.
The Canada goose flight from northwest Colorado will be about average while
the north-central flight will be larger than 1974 but still below average.

-ffl&amp;Jl I) --1.4

Prepared

Prepared by ..
~hl'&lt;',4l
Michael R. Szymce;k:)
Wildlife Researcher
I

Research Tech. IB

~i

�Table 6.

Results of the north-central

Production
Area

Water Area

Fort Collins
(continued)

Fossil Creek Reservoir
Schuelke Reservoir
Wolaver Ponds

Colorado

No.
Broods

5
2
2

Total
Denver

Ketring Pond
Centennial Pond
Columbine Country Club
Chatfield Country Club
Bowles Lake
Kings Lake
Blackmer
Mohn Estates
Grant Lake
Colo. Bldv. &amp; Quincy
Marston Reservoir
Pinehurst Country Club
Bit-a-Sea
Reservoir If3
Ward Reservoir
Kendrick Lake
Federal Center
Sloan's Lake
Standley
Denver City Park
Total

GRAND TOTAL

!/Two geese still nesting.
°
-2/0 ne goose st~on nest~ng.

goose census, June, 1975 (continued).

10
4
2
3
6
10
1
4
2
5

2
9
1
1
1
4
13
2
1

Total
No. Goslings

Total No.
Adults &amp; Yearlings

Total
Birds

16
9
7

11
6
4

27
15
11

234

667

901

37
11
8
13
17
36
3
15
8
12
10
45
2
5
2
14
44
4
2

-

111
32
18
22
147
124
8
36
145
52
54
56
107
16
7
23
36
187
8
139

148
43
26
35
164
160
11
51
153
64
54
66
152
18
12
25
50
231
12
141

288

1,328

1,616

1,049

3,349

4,398

I
•.....
0
I

�-11-

Table 7. Number of Canada goose goslings produced in north-central
Colorado production trend areas, 1975.

Area

Number of Goslings
1969-1974
1975
1974
Average

Wellington

207

187

268

+10.7

-22.8

Fort Collins

234

229

274

+ 2.2

-14.6

Loveland

103

90

83

+14.4

+24.1

Boulder

217

206

237

+ 5.3

- 8.4

Denver

288

185

280

+55.7

+ 2.9

Totals

1,049

897

1,142

+16.9

- 8.1

Percent Change
From
From 1974
1969-1974

Table 8. Number )f adult Canada geese observed in north-central
production trend areas, 1975.

Colorado

Area

Number of Geese
1969-1974
1975
1974
Average

Wellington

570

635

812

-10.2

- 29.8

Fort Collins

667

477

717

+39.8

- 6.9

Loveland

318

239

204

+33.0

+55.9

Boulder

466

365

676

+27.7

-31.1

Denver

1,328

1,004

1,299

+32.3

+ 2.2

Totals

3,349

2,720

3,708

+23.1

- 9.7

Percent Change
From
From 1974
1969-1974

��October 1976
-13-

JOB FINAL REPORT

State of

COLORADO
------~~~~------------

Project No.

W-88-R-2l

Work Plan No.

1

Job Title

Job No.

2

T~r~a~p~p_i~n~g~a~n~d~B_a~n~d~in~g
__D~u~c_k~s~a~n~d_.~G~e~e~s_e~

Period Covered:
Personnel:

Migratory Bird Investigations

_

April 1, 1975 to March 31, 1976

C. Bryant and Staff of Alamosa and M~nte Vista National Wildlife
Refuges, S. Bissell, L. Budde, J. Carsella, G. Claassen, J. Corey,
D. Coven, M. Conner, C. Crawford, M. DePra, K. Dillinger,
W. Dolzal, G. East, G. Eyre, H. Funk, J. Gray, J. Gumber,
A. Hemmert, R. Holder, R. Leasure, J. Lorentzson, R. Mason,
J. Miller, K. Miller, W. Mink, K. Moser, R. Oakleaf, B. Peterson,
J. Pierce, S. Porter, R. Richardson, F. Rinella, Jr., W. Rupke,
H. Spear, S. Steinert, M. Stone, M. Szymczak, C. Wagner, J. Wagner,
K. Wagner, C. Wetherill, R. Velarde and R. Hopper.

ABSTRACT

Trapping efforts in Segment 21 resulted in the banding of 18,349 ducks of ten
species at eight locations in Colorado. The major species in the banded
sample was the mallard which contributed 11,209 or about 61 percent of the
total. Pre-season banding in the high country yielded 10,149, including
3,599 in North Park, 3,472 in South Park and 3,078 in the San Luis Valley.
Mallards and pintails were the dominant species in the sample in these three
areas. Post-season banding produced a sample of 8,200 mallards including
6,700 in eastern Colorado (Central Flyway) and 1,500 in west central Colorado
(Pacific Flyway). Goose trapping activities resulted in a banded sample
of 1,605 geese in Segment 21. Included in this sample were 65 goslings
transplanted to a production site in the South Platte River Valley. Summer
banding on established production areas in the San Luis Valley and in north
central Colorado accounted for 620. The remaining 920 were banded postseason in the Arkansas Valley.

�-16-

RESULTS AND DISCUSSION
Ducks
Over 18,300 ducks of 10 species were banded in Segment 21 (Table 1).
The South Platte Valley produced the largest sample (3,824) of the
eight general banding locations in Colorado, followed by North Park
(3,599), South Park (3,472), the San Luis Valley (3,078) and the
Grand Junction-Delta area of the west slope (1,500).
The mallard continued to dominate the banded sample, comprlslng 11,209,
or about 61 percent of the total. Pintails and green-winged teals were
the next most important contributors to the banded sample with 3,712 and
1,947, respectively.
High Country Study
Objectives of the high country study of duck populations are presented by
Hopper (1972). Species compositon by age and sex for 11,881 ducks banded
in the three high country areas during 1975 is shown in Tables 2-5. San
Luis Valley bandings were accomplished by both State and Federal personnel
(Table 4). The U. S. Fish and Wildlife Service continued to cooperate in
this study by banding a portion of the Valley quota on the Monte Vista and
Alamosa National Wildlife Refuges.
Mallards and pintails were again the main species in the banded samples
(Tables 2-4). They contributed 4,129 and 4,048 birds, respectively, for
all three areas combined (Table 5). The only other species for which
significant banded samples were obtained included green-winged teals
(2,084) and blue-winged and cinnamon teals (984). Practically all gadwalls
banded in North Park were a result of night-lighting efforts.
All phases of the High Country Duck Study will be continued in future segments under Work Plan 3, Job 9.

Winter Banding
Post-season banding of terminal wintering populations of mallards was continued
in eastern Colorado for the thirteenth consecutive year under Work Plan 3,
Job 6. The total banded sample amounted to 6,700 mallards for this period
in Segment 21 (Table 1). A detailed breakdown of this sample by age and
sex is shown in Table 6. Future banding work related to these wintering
populations will be reported as part of a new management oriented job (Work
Plan 3, Job 9).
An additional 1,500 mallards were banded on the Western Slope of Colorado
during Segment 21 (Table 1). This was the third consecutive year of banding
in this area as part of a study described by Hopper (1974) in a previous
Job Progress Report for Work Plan 1, Job 2. Age and sex ratios obtained
in ·this year's banded sample are shown in Table 7 by location. This study
will continue as a new and separate job of its own (Work Plan 3, Job 7)
in the ensuing segments.

�Table l.

Number of ducks banded by species, location and period of year 1975-76.

Number of Ducks Banded
Pre-season-

1/

Post-season-

2/

Species

North
Park

South
Park

San Luis
Valley

Cache 1a
Poudre
Valley

South
Platte
Valley

Bonny
Res.

Arkansas
Valley

West3/
Slope-

Total

Mallard

1,171

657

1,181

1,059

3,824

845

97'1.

1,500

11,209

Gadwall

484

--

2

--

--

--

--

--

486

--

--

--

--

63

--

--

--

--

1,947

--

--

--

897

American

Wigeon

Green-winged

Teal

Blue-winged and
Cinnamon Teal

55

4

4

372

1,274

301

---

16

567

314

--

--

I

I

Pintail

1,495

962

1,255

--

--

--

--

--

3,712

Shoveler

--

1

--

--

--

--

--

--

1

7

19

--

--

--

--

--

31

Redhead

5

Canvasback

1

--

2

--

--

--

--

--

3

3,599

3,472

3,078

1,059

3,824

845

972

1,500

18,349

Total
1/
- August-September.
2/
- January-February.

1/Co1orado-Gunnison-Uncompahgre

River Complex.

I-'
--.J

�-18Table 2.

Age and sex composition of ducks banded pre-season in North Park, 1975.

Species

AM

Number of Ducks Banded
Age and Sex
1M
AF
IF

Mallard

318

332

291

230

1,171

Gadwall

395

0

85

4

484

American Wigeon

28

12

6

9

55

Green-winged

176

99

35

62

372

3

3

1

9

16

Pintail

423

363

346

363

1,495

Redhead

2

0

3

0

5

Canvasback

0

0

1

0

1

1,345

809

768

677

3,599

Teal

Blue-winged and
Cinnamon Teal

Total

Table 3.

Total

Age and sex composition of ducks banded pre-season in South Park, 1975.

Species

AM

Number of Ducks Banded
Age and Sex
1M
AF
IF

Mallard

229

131

205

92

657

2

0

2

0

4

399

484

105

286

1,274

Blue-winged and
Cinnamon Teal

77

252

36

202

567

Shoveler

0

1

0

0

1

Pintail

306

296

147

213

962

Redhead

2

0

5

0

7

1,015

1,164

500

793

3,472

American Wigeon
Green-winged

Total

Teal

Total

�19

Table 4. Age and sex composition of ducks banded pre-season in the San Luis
Valley, 1975.Y.

Species

AM

Number of Ducks Banded
Age and Sex
1M
AF
IF

Mallard

494

744

411

652

2,301

Gadwall

o

12

1

21

34

American Wigeon

o

2

2

1

5

186

115

50

87

438

Blue-winged and
Cinnamon Teal

30

213

20

138

401

Pintail

289

557

205

540

1,591

Redhead

3

12

8

9

381:./

Canvasback

1

o

1

o

2

1,003

1,655

698

1,448

Green-winged

Total

Teal

Total

4 ,8l0Y

!/Includes 1,732 ducks banded by Federal personnel on Alamosa and Monte Vista
National Wildlife Refuges as part of the High Country Duck Study.
1:./Includessix "HY" birds unidentified as to sex.

�-20-

Table 5. Age and sex composition qf ducks banded pre-season in the three
high-country areas combined, 197sl/.

Number of Ducks Banded
Age and Sex
1M
AF
IF

Species

AM

Mallard

1,041

1,207

907

974

4,129

Gadwall

395

12

86

25

518

American Wigeon

30

14

10

10

64

Green-winged

761

698

190

435

2,084

110

468

57

349

984

Shoveler

0

1

0

0

1

Pintail

1,018

1,216

698

1,116

4,048

Redhead

7

12

16

9

50

Canvasback

1

0

2

0

3

3,363

3,628

1,966

2,918

Teal

Blue-winged and
Cinnamon Teal

Total

Total

Y

ll,881Y

!/Includes 1,732 ducks banded by Federal personnel on Alamosa and Monte Vista
National Wildlife Refuges as part of the High-Country Duck Study.
]j Includes six "HY" birds unidentified as to sex.

�-21-

Table 6. Mallards banded post-season by age and sex in the seven east slope
banding locations.

Location

AM

Number of Ducks Banded
Age and Sex
1M
AF
IF

Bonny Reservoir

250

250

148

197

845

Sterling-Julesburg

250

236

145

l32

763

Fort Morgan-Sterling

250

242

216

234

942

Greeley-Fort Morgan

354

290

197

205

1,046

Denver-Greeley

298

262

244

269

1,073

Fort Collins-LovelandWindsor

292

304

227

236

1,059

Arkansas Valley

367

234

146

225

972

2,061

1,818

1,323

1,498

6,700

Total

Table 7. Age and sex composition of mallards banded post-season
Colorado, 1975-76.

Total

in west central

Location

AM

Number of Ducks Banded
Age and Sex
1M
AF
IF

Grand Junction-Mack

156

91

98

90

435

Delta

241

274

265

285

1,065

Total

397

365

363

375

1,500

Total

�-22-

Geese
Over 1,600 Canada geese were banded during Segment 21 (Table 8). The Arkansas
Valley accounted for the largest number (920), while the Cache la PoudreSouth Platte Valleys, and the San Luis Valley followed with 666 and 19,
respectively.

Summer Transplants
Sixty-five Canada goose goslings (28 males, 37 females) were banded and
transported from the Front Range production areas and released on Prewitt
Reservoir in the South Platte River Valley (Table 8). The Canada goose
transplant program is no longer a part of Federal Aid Project W-88-R.

Table 8.
1974-75.

Number of Canada geese banded by location and period of year,

Number Banded
Summer 1/
Transplants-

1/
Pre-season-

2/
Post-season-

Total

Arkansas Valley

0

0

920

920

Cache la Poudre and South
Platte Valleys

65

601

0

666

San Luis Valley

0

19

0

19

Total

65

620

920

1,605

Location

!/June-July.
2/
- January-February.

Pre-season Bandiag on Colorado Production Areas
A study was initiated in 1974 to investigate the status of breeding populations
of Canada geese in the San Luis Valley and the Fort Collins-Boulder-Denver
area of north central Colorado. Pre-season banding is a major component of
this study. Details of the overall investigation are discussed under Work
Plan 2, Job 6, and future bandings will be reported under that job.

�-23-

Trapping efforts proved unsuccessful in the San Luis Valley, with only 19
geese being banded (Table 8). The banding crew was too small and attempts
to drive the birds proved futile. More time and man power will be allocated
for this endeavor next segment.
Pre-season trapping operations were highly successful in north central Colorado
and resulted in the banding of 601 adult and young geese (Table 9).
The sample consisted of approximately equal numbers of each sex by age class.

Pre-season Banding on Alberta Production Areas
The Colorado Division of Wildlife assisted in a Central Flyway and Pacific
Flyway cooperative banding study of Canada geese·on Canadian production areas
in southern Alberta for the second consecutive year. The major purpose of this
study is to define the breeding areas of the Great Basin and Hi-line Canada
Goose Populations in Alberta. This cooperative venture, involving personnel
from Colorado, Idaho, Utah, Wyoming, Alberta, the U.S. Fish and Wildlife Service
and the Canadian Wildlife Service resulted in the banding of 1,442 Canada
geese in a 10-day period in June and July. These bandings are not included
in any totals or tables in this report, since the study is the primary
responsibility of the Province of Alberta.

Table 9. Age and sex composition of Canada geese banded pre-season in north
central Colorado, 1975.

Age and Sex

Number Banded

Adult Male

213

Local Male

95

Adult Female

193

Local Female

100

Total

601

Winter Banding
Post-season activities produced 920 banded Canada geese in the Arkansas
Valley (Table 10). This banded sample of Shortgrass Prairie Canada geese
produced good numbers of birds by age and sex. Future post-season bandings
of this population of geese in Colorado will be reported under the newly
developed Work Plan 2, Job 9.

�-24-

Table 10. Age and sex composition of Canada geese banded post-season in
eastern Colorado, 1975-76.

Location

AM

Number Banded
Age and Sex
1M
AF

Arkansas Valley

223

213

182

IF

Total

300

920

LITERATURE CITED
Hopper, R. M. 1972. Trapping and banding ducks and geese.
Fed. Aid Game Res. Rept. , Oct. pp. l3-2l.

Colo. Div. Wildl. ,

Hopper, R. M. 1974. Trapping and banding ducks and geese.
Fed. Aid Game Res. Rept., Oct. pp. l3-22.

Colo. Div. Wildl. ,

Prepared by

D

-/ ~

"'~
if ,Ae/W:Z~~'
,

J
'~
&gt;'

Richard M. HopperT
Wildlife Researcher

'

Ll
.

�-25-

October 1976

JOB FINAL REPORT

State of

COLORADO

----------------------

Project No.

W-88-R-21
1

Work Plan No.
Job Title

Job No •.

3

---------------------------

Analysis of Waterfowl Banding Data

Period Covered:
Personnel:

Migratory Bird Investigations

1955-56 through 1975-76

Michael R. Szymczak

ABSTRACT
Work under this job has entailed a number of independent analyses of data
resulting from banding of various populations.
Each analyses has been
essentially a final report evolving from a certain set or sets of segment
objectives and procedures. Many times the report was in the form of a publication. Although each set of objectives was independent, they all were
formulated to help meet the P. S. Objective "To formulate waterfowl harvest
regulations for Colorado". In the future, each specific set of objectives
will be attached to a PROGRAM NARRATIVE OUTLINE with a certain job number.
Therefore, there is no longer a need for Work Plan 1, Job 3.
The most recent objectives, analysis of migration, mortality, recovery
distribution and relationships among populations of ducks banded in North
Park, South Park and the San Luis Valley was scheduled for completion March
1976. However, the analysis was postponed to allow the compilation of at
least five concurrent consecutive years of recovery data from bandings in
the three areas. This specific study has been expanded and as of April 1,
1976 is Work Plan 3, Job 9 entitled "Migration and Mortality Characteristics
of Duck Populations in the Inter-mountain Valleys of Colorado".

Wildlife Researcher

��October, 1976

-27JOB FINAL REPORT

State of

COLORADO

----------------------

Project No.

W-88-R-2l

Work Plan No.

1

Migratory Bird Inyestigations

Job No. 14
Determination of Methods for Developing and
Job Title Managing Waterfowl Habitat - Ammonium Nitrate Pothole Blasting Study
Period Covered:
Personnel:

---------------------------

April 1, 1967 - March 31, 1976

Richard M. Hopper

ABSTRACT
Field work was cOm?leted for this Job during Segment 21. The final report,
covering all phases not previously reported upon, is in the process of
preparation.
Further work on this report will continue under \vork Plan 6,
Job 1. The resulting report will be submitted to The Wildlife Society
for possible publication in the Journal or Bulletin, or will be
presented as a Division Special Report. Two previous publications
helped fulfill reporting responsibility for this Job.

��-29-

AMMONIUM NITRATE POTHOLE BLASTING STUDY
Richard M. Hopper

P. S. OBJECTIVE
To evaluate various size potholes blasted with ammonium nitrate in terms
of (a) life expectancy, (b) plant succession, (c) soil and water characters,
(d) waterfowl use and hunting potential, and (e) cost.

METHODS AND MATERIALS
Compile and analyze all data collected as part of this Job that hasn't been
reported upon earlier.

RESULTS AND DISCUSSION
Field work for this job was terminated on May 15, 1975. Since that time
office work has involved the tabulation and analysis of data collected in
Segment 21. These data are currently being incorporated with data from
all previous segments in preparation of the final report. All remaining
reporting responsibility for this Job will be completed under Work Plan 6,
Job 1. This final report will be prepared for publication in The Journal
of Wildlife Management or the Wildlife Society Bulletin, or as a Division
Special Report.
The papers listed below were published previously in partial fulfillment of
reporting responsibilities for Work Plan 1, Job 14:
Hopper, R. M. 1971. Use of ammonium nitrate-fuel oil mixtures in
blasting potholes for wildlife.
Colo. Div. Wildl., Game Inf.
Leaflet No. 85, 4pp.
Hopper, R. M. 1972. Waterfowl use in relation to size and cost of
. potholes. J. Wildl. Manage. 36(2):459-468.

Prepared by

£~~~~

;:;~
/,
/4t..-L..
.' Richard M. Hbpp~r
~
Wildlife Researcher

��October 1976

-31JOB PIWGRESS RE?ORT

State of

COLORADO
W-88-R-21

Project No.
Work Plan No.

Migratory Bird Investigations

2

Job Title

Job No.
~
2----__ -- __ ---------Experimental Studies on Improving
The Status of Canada Goose Populations

Period Covered:

April 29, 1975 through July 23, 1975

Personnel:

J. Lorentzson, S. Porter, W. Russell, S. Steinert, K. Wagner,
J. Wagner and M. Szymczak.

ABSTRACT
A total of 127 Canada geese (Branta canadensis) were observed during an
aerial flight in the South Platte River Valley on April 29, 1975. An estimated
forty-six indicated nesting pairs were sighted. Seventy-five of the 127
geese and 34 of the indicated nesting pairs were located in the Sterling to
Julesburg area. Seventeen nesting pairs were associated with the Jumbo Reservoir Annex release site. Ground observations indicated that most nests were
located in nesting structures. A total of 71 Canada geese were observed
during an aerial flight in North Park on May 4, 1975. An estimated 17 indicated nesting pairs were sighted. Twenty-four of the 71 geese and ten of the
16 indicated ne:stingpairs were located at Walden Reservoir. Occasional observation on some production areas resulted in observation of two goslings (one
brood) at MacFarlane Reservoir, three goslings (one brood) at Pole Mountain
Reservoir, three goslings (two broods) at Lake John Annex, and 56 goslings at
Walden Reservoir. A total of four known nests were established at Antero
Reservoir (two successful and two destroyed). The high count of geese at
Antero occurred on July 23, 1975 when 115 birds were observed.

��-33-

EXPERIMENTAL STUDIES ON IMPROVING
THE STATUS OF CANADA GOOSE POPULATIONS
Michael R. Szymczak

P. S. OBJECTIVE
To investigate the success of attempts to establish breeding populations of
Canada geese in,suitable habitat where they do not presently exist in Colorado.
SEGMENT OBJECTIVE
To examine the breeding population status of Crulada geese in the Prewitt
Reservoir and Jumbo Reservoir area in the South Platte Valley, Antero Reservoir in South Park, and in North Park.
METHODS AND MATERIALS
An aerial survey of the South Platte River was conducted on April 29, 1975
from the confluence of the Poudre and South Platte Rivers, located about
five miles east of Greeley, to the Colorado-Nebraska state line. All irrigation reservoirs, including Riverside, Empire, Jackson, Bijou, Prewitt, and
Jumbo were surveyed, as well as smaller ponds near the river such as Johnson's
Pond. All geese observed were recorded as birds on nest, singles, pairs or
groups. During the first week in May, ground observations were conducted in
most of the areas in which geese were sighted. Where applicable, all observations were recorded by township, range and section to facilitate locating
the area on the ground.
On May 4, 1975 an aerial survey of all lakes, reservoirs, ponds and major
streams, including Grizzly Creek, the Michigan, Illinois and North Platte
rivers, was conducted in North Park. Geese were classified in the same
manner as described above. Production counts were conducted on the larger
water areas through ~ay and early June. A nest search was conducted on Walden
Reservoir in early August. Periodic obse~ations of Antero Reservoir were
conducted from the ground.
RESULTS AND DISCUSSION
South Platte Valley
Aerial Survey
A total of 127 geese were observed and classified during the aerial flight
in the South Platte Valley (Table 1). Forty-one birds were observed as being
actually sitting on nests. An additional five birds observed as singles were
thought to be members of nesting pairs, bringing the number of estimated nesting pairs to 46. Twenty-three pairs considered to be sub-adults were also
observed.

�Table 1. Location and classification of Canada geese observed from the air in the South Platte Valley
from the confluence of the South Platte River and the Cache la Poudre River to the Colorado-Nebraska state
line on April 29, 1975.

Location

Nesting

Number of Geese Observed
Singles
Pairs
Groups

Total

Greeley to Fort Morgan
South Platte River
2 mi west of Kersey (5N, 64W, SB)
Bijou Inlet Ditch (4N, 63W, S12)
2 mi west of K-4 Ranch (4N, 62W, S17)
1 mi west of K-4 Ranch (4N, 62W, S16)
K-4 Ranch (4N, 62W, S23)

o
o
o
o
2

o

o
o

Gallagers

5

1

4

(4N, 6lW, SIB)

2

o
o
1

o
2

2

o
o
o
o
o

2
2
2

1
2

10

I
W
.p..

Empire Reservoir
Fort Morgan

(3N, BlW, Sl)

o

1

4

o

5

1

o

o

3

4

o

2

o

o
o
o

2

o
o

2

o

2
2
2

o

o

o

4

4

o
o
o
o

o
o
o
o

o

4

4

2

4

4
4

to Sterling

Fort Morgan Gravel Pits (4N, 57W, S32)
South Platte River
North of Brush (4N, 56W, S15)
~ md east of Snyder Bridge (4N, 56W, S12)
~ mi east of Bridge SW of Merino (6N, 54W,
S26)
Sterling (BN, 52W, S33)

o

Prewitt Reservoir
5N, 53W,
5N, 53W,
5N, 54W,
5N, 54W,

S5
S6
S12
S14

o
4

o
o

-----------------------------------------------------------------------------------------------------------

2

I

�Table 1. Location and classification of Canada geese observed from the air in the South Platte Valley
from the confluence of the South Platte River and the Cache la Poudre River to the Colorado-Nebraska state
line on April 29, 1975 (continued).

Location

Sterling

Nesting

Number of Geese Observed
Groups
Singles
Pairs

Total

to Julesburg

South Platte River
9N, 51W, S21
Tamarack Ranch (lON, 48W, S9)
Tamarack
Ranch (lON, 49W, S14)
Tamarack Ranch (lON, SOW, S26)
2 mi east of Sedgwick Bar (llN, 48W, S7)
North of Johnson's Pond (llN, 45W, SlO)

0

1
1
0
0
0

2

0

1

0
0

0

2

0
0
0
0
0

0
0

1

0

0

2

2
1
2
2
1
2

I
W

\J1

I

Jumbo Annex
llN, 48W, S24, S25
llN, 47W, S19
Ponds West (llN, 48W, S22)
Pond north of Jumbo Reservoir
Johnson's

Total

Pond

(llN, 48W, Sl)

14
1
1

6

6

0
1

0

0
0

6

4

26
1
12

1

0

4

0

5

14

3

0

4

21

41

17

46

23

127

�-36-

Most birds were sighted in the Sterling to Julesburg area, primarily around
Johnson's Pond and Jumbo Annex (Table 2). From Greeley to Fort Morgan a
total of 24 birds were observed including 11 indicated nesting pairs. Only
one-half of the birds observed were sighted on the K-4 Ranch-Gallager release
site, but 7 of the 11 indicated nesting pairs were on those areas. Twentyeight birds were observed in the Fort Morgan to Sterling area, with only one
indicated nesting pair. One-half the geese in the Fort Morgan to Sterling
area were observed near the Prewitt Reservoir release site. Thirty-four
of the 46 total South Platte River area indicated nesting pairs were observed in the Sterling to .Ju Lesburg area. Fourteen of these indicated pairs
were observed on Johnson's Pond, a Canada goose nesting area prior to
restoration activities at Jumbo Annex (Table 2). Seventeen of the remaining
19 indicated nesting pairs were located in the Jumbo Reservoir vicinity.
Ground Survey
Nest searches were conducted from the ground on certain areas. Searches in
river bottom areas in which indicated nesting pairs were observed from the
air were generally fruitless, when the birds were not associated with structures. At the K-4 Ranch three nests were found, one which had been destroyed
and two which were successful. Six nests were found at Gallagers, two on
floating structures, three on standing structures and one on the ground. The
one nest on a structure at the Fort Morgan Gravel Pit was located. One
nest, not observed from the air, was located on the south side of Prewitt
Reservoir. Two of the three probable nest sites observed from the air along
the South Platte between Sterling and Julesburg were located but no nests
were found. At Jumbo Reservoir Annex, thirteen nests were observed, one of
which had hatched. Six of the nests were located on standing structures
and seven on small man-made islands.
A comparison of the results of aerial counts of indicated breeding pairs
with ground counts of nests on selected areas is presented in Table 3.
Fifteen of the 24 nests located during the ground survey were positioned
on artificial nesting structures. An additional seven nests were located
on man-made islands constructed specifically as goose nesting sites.
North Park
Aerial Survey
A total of 71 Canada geese were observed during the aerial goose breeding
pair survey on May 4, 1975 in North Park (Table 4). Four geese were observed on nests. An additional 13 single birds were observed bringing the
indicated nesting pair total to 17. Twenty non-nesting pairs were also
sighted.
About one-third of the total birds observed and ten of the 17 indicated
nesting pairs were located on Walden Reservoir, the secondary release site
in North Park (Szymczak 1973, 1974). Two indicated nesting pairs and 3 nonnesting pairs were sighted at Lake John Annex, the primary release site
(Szymczak 1973). Other indicated nesting pairs were observed on Grizzly
Creek west of Hebron Ponds, at Boettcher Lakes, at Case Flats, and on the
North Platte River, southwest of Walden and west of Cowdrey.

�-37-

Table 2. Sunnnarization of goose observations along the South Platte
River area on April 29, 1975 from Greeley to Julesburg.

Area

On Nests

Number of Geese Observed
Pairs
Singles
Groups

Total

Greele~ to Fort Morgan
South Platte River
(including K-4Ranch)

2

3

4

0

9

Gallagers

5

1 1./

4

0

10

Empire Reservoir

0

1

4

0

5

Subtotal

7

5

12

0

24

South Platte River

0

0

6

4

10

Fort Morgan Gravel Pits

1

0

0

3

4

Prewitt Reservoir

0

0

6

8

14

Subtotal

1

0

12

15

28

6

0

10

6

0

27

10

4

17

0

4

21

Fort Morgan to Sterling

Sterling to Julesburg
2 ];./

South Platte River

2

Jumbo Annex

15

Jumbo Annex Area

2

Johnson's Pond

14

Subtotal

33

12

22

8

75

Grand Total

41

17

46

23

127

1./

Associated with goose observed on nest.

Jj One associated with goose observed on nest.

Y

6

1 1./
3 1./

�-38-

Table 3. Comparison of the number of indicated nesting pairs observed
from the air with ground count information on selected sites.

Number
Indicated Nesting Pairs
(Aerial)

Nests Observed
(Ground)

South Platte River (K-4 Ranch)

5

3

Gallagers

5

6

Subtotal

10

9

South Platte River

0

0

Fort Morgan Gravel Pits

1

1

PreWitt

0

1

1

2

South Platte River

2

0

Jumbo Annex Area

16

13

Subtotal

18

13

Grand Total

29

24

Area

Greeley to Fort Morgan

Fort Morgan to Sterling

Reservoir

Subtotal

Sterling to Julesburg

�Table 4.

Locations

and classifications

of Canada geese observed from the air in North Park on May 4, 1975.

Location

Nesting

Number of Geese Observed
Singles
Pairs
Groups

o
o

o
o

o
o

Total

2

o

2

o

4

4

o

2
6

o

7

10
6

1

11/

o

1

o

4
2

1

2

o

2

2

6

Case Flats (8N, 80W, S13)

1

o

2

Walden Reservoir

2

10'1:/

12

o
o
o
o
o
o
o
o

2
5

Lake John Annex (9N, 8lW, S2)

o
o
o
o
o

24

4

16

40

11

71

MacFarlane

Reservoir

Pole Mountain

(7N, 79W, S29)

Reservoir

(7N, 8lW, S22)

",.

Grizzly Creek

West of Hebron Ponds (7N, 80W, S33)
Fox Ranch (8N, 8lW, S24)

1

North Platte River
Southwest of Walden
West of Cowdrey (ION, 8W, SlO)
Southeast of Lake John
BoettcherLakes

(ION, 8lW, S20)

Furnow Pond (ION, 8lW, S6)

(9N, 79W, S19, S20)

Total

];./Associated with goose observed on nest.
~/ Two associated with goose observed on nest.

2

3
2
8

3

I

\..oJ
1.0

I

�-40-

Ground Surveys
No nest site checks were conducted in North Park immediately after the
survey. Occasional observations were made on some of the potential production areas during the brood rearing period. Broods were observed at
MacFarlane Reservoir (2 goslings), Pole Mountain Reservoir (3 goslings),
Lake John Annex (2 goslings, 1 gosling) and Walden Reservoir (56 goslings
total).
Indicated breeding pairs were observed at Walden and Lake John Annex during
the aerial survey but not at MacFarlane or Pole Mountain. The size of
goslings observed at MacFarlane in August indicated the adult pair that
produced the goslings weren't nesting at the time of the aerial flight. The
birds that produced the goslings at Pole Mountain nested along heavily
willowed Little Grizzly Creek adjacent to the Reservoir.
A nest search of the islands and structures at Walden Reservoir located a
total of 10 nests, 6 of which were judged to have been successful, one that
had been destroyed, one that was abandoned, and two that the fate was
unknown. All of the nests were located on islands. The advanced stage of
the vegetation on the island made locating nests difficult.
Antero Reservoir-South Park
Nesting pair surveys were not conducted at Antero Reservoir. Periodic counts
through late spring and early summer resulted in two broods being found. Two
destroyed nests were also located. The high count of geese at Antero occurred
on July 23, 1975 when 115 birds were observed.
One pair of geese, whose actions indicated they were accompanied by a brood,
were observed in late spring on Eleven Mile Reservoir, located about 20 miles
east of Antero Reservoir.
LITERATURE CITED
Szymczak, M. R. 1973. Experimental studies on improving status of Canada
goose populations. Colo. Div. of Wildl., Game Res. Rept., Fed. Aid
Project W-88-R. October. p. 33-42.
1974. Experimental studies on improving status of Canada goose
populations. Colo. Div. of Wildl., Game Res. Rept., Fed. Aid Project
W-88-R. October. p. 47-53.

Prepared

J.~~

bY_'-J.I,-n~A&lt;:L.:-M=· =--_.~~:&gt;.....:=-.. ~~~l
Michael R. szymcza~~
Wildlife Researcher

�October 1976

-41-

JOB PROGRESS r~PORT

State of ____

-'-'-~c..;;...:;;_'

COLORADO

Project No.

W-88-R-2..=1

Work Plan No.
Job Title

_

2

Migratory Bird Investigations
6
-------------------------

Job No.

Studies of Canada Goose Populations in Colorado Transplant Areas

Period Covered:

April 1, 1975 to March 31, 1976

Personnel: C. Bryant and Staff, Monte Vista National Wildlife Refuge; M.
Babler, D. Benson, S. Bissell, G. Brown, L. Budde, J. Corey,
G. Claassen, R. Clark, C. Crawford, M. DePra, R. Desilet, J~
Ellenberger, J. Frothingham, B. Goetze, V. Graham, J. Gumber,
S. Heath, T. Henry, J. Hobbs, R. Hopper, J. Jackson, R. Leasure,
C. Leonard, J. Lorentzson, T. Lynch, K. Miller, K. Moser, D. Owens,
C. Pabst, S. Palm, J. Pierce, B. Peterson, F. Rinella, C. Roberts,
L. Rottman, G. Saville, H. Spear, S. Steinert, E. Wagner, and M.
Szymczak, Colorado Division of Wildlife.

ABSTRACT
Banded Canada geese (Branta canadensis) from Dowling Lake Alberta, the
Cypress Hills area of Saskatchewan, Phillips County, Montana and the Big
Horn-Treasure County areas of Montana, all documented Hi-Line nesting areas
were well representei in the goose harvest in northcentral Colorado during
the 1974-1975 hunting season. First year or direct recoveries from Schuler,
Many Islands, Seven Persons, and Murray Lakes, all areas east of 111 longitude
in southern Alberta were also reported taken in northcentral Colorado. Southern Alberta bandings west of 111 longitude were not represented in the northcentral Colorado harvest. Various methods of estimating the number of "small"
geese harvested in northcentral Colorado based on tail fan collections are
discussed. Since the inception of post-season banding in northcentral Colorado
in 1971-1972, Alberta has been the major recovery area for birds considered
"small" Canada geese when banded. Analysis of band recovery distribution and
weight data indicate that geese classified as "small" birds banded post-season
in northcentral Colorado may not be a simple cross-section of "small" geese
from the short grass prairie Canada geese population. A record 68.8 percent
of the large geese banded post-season in northcentral Colorado and harvested
during the 1974-1975 hunting were reported taken in northcentral Colorado.
0

0

The January 1976 goose census in the Hi-Line population wintering range resulted
in a record 58,436 birds being counted. Ninty-three percent of the population
was located in Colorado during the count.
.
Canada goose counts in the San Luis Valley totaled 1,230 on November 20, 1975,
959 on December 18, 1975, and 827 on January 8, 1976. An estimated total of
256 active permit holders hunted an average of 4.5 days and harvested 86 geese
for a success rate of 0.34. An estimated 61 of the geese were harvested in
Rio Grande County.
(Abstract continued on next page)

�-42-

ABSTRACT

(Continued)

In Mesa County an estimated 256 active permit holders hunted an average
of 4.1 days, taking an estimated 93 geese. In Garfield County, an estimated
80 active permit holders hunted an average of 3.6 days harvesting 20 geese.
In the first year of goose hunting in Delta County an estimated 254 permit
holders hunted an average of 4.2 days and harvested 68 geese.

�•
-43-

STUDIES OF CANADA GOOSE
POPULATIONS IN COLORADO TRANSPLANT AREAS

Michael R. Szymczak

P. S. OBJECTIVE
To investigate the status of resident and migrant Canada goose flocks
and their interrelationships in areas in which populations have been
established through transplant programs in Colorado.

SEGMENT OBJECTIVES
la.

To determine breeding areas of Canada geese wintering in northcentral
Colorado.

lb.

To examine migration routes and/or harvest patterns and distribution
of Canada g~ese wintering in northcentral Colorado.

2.

To estimate hunting pressure on, and hunter harvest of Canada geese
in the San Luis Valley and westcentral Colorado.

3.

To estimate annual mortality of Canada geese wintering in northcentral
Colorado.

4.

To make recommendations for continuing Canada goose hunting seasons in
westcentral Colorado, northcentral Colorado and the San Luis Valley.

METHODS AND MATERIALS
Band recoveries listed on computer printouts provided periodically by the
Migratory Bird Populations Station were sorted and plotted as to banding and
recovery location, sex, and age at time of banding, and estimated breeding
area for birds associated with northcentral Colorado.
Tail fans reported from Canada geese harvested in northcentral Colorado and
collected through the U. S. Fish and Wildlife Service's Waterfowl Parts
Collection Survey were examined and classified as to whether the fans
originated from large or small Canada geese.
Canada geese in northcentral Colorado were counted on December 18, 1975 and
January 9, 1976. In the San Luis Valley geese were counted on November 20,
1975, December 18, 1975, and January 8, 1976.
All hunters obtaining special permits to hunt geese in the San Luis Valley
and westcentral Colorado were mailed questionnaires inquiring about their
hunting activity and success.

�-44Recoveries of Canada geese banded post-season in northcentral Colorado were
sorted by age and recovery year and mortality rates calculated by the composite dynamic. Adequate banding and recovery data for Hi-Line population
breeding areas were not obtained. Therefore comparisons of distribution
of harvest and of mortality estimates for the same population from two different banding operations, pre-season and post-season, could not be made.
All pertinent data collected concerning the status of the northcentral,
westcentral and San Luis Valley Canada goose populations were presented to
management personnel in Colorado for use in establishing hunting seasons.

RESULTS AND DISCUSSION

Hi-Line Population
Foreign Recoveries
During the 1974-75 hunting season 155 Canada geese that were banded outside
of Colorado were reported recovered in northcentral Colorado (Table 1).

Table 1. Banding areas outside Colorado of Canada geese recovered in northcentral Colorado during the 1974-75 hunting season.

Area

Direct

Indirect

Alberta
Edmonton-Camrose Area
Dowling Lake Area
Brooks Area
Schuler Lake
Many Islands Lake
Seven Persons Lake
Murray Lake

1

1

7

5
3

2
2

5
3

Saskatchewan
Manito Lake Area
Cypress Hills Area
Crane Lake
Val Marie-Frenchman River (Masefield)
Ponteix-Notukeu Creek Area

3
8

5

3

1
1

1

o

3

Mackenzie-N.W. Terr~tory
Thelon River Area

1

---------------------------------------

�-45Table 1. Banding areas outside Colorado of Canada geese recovered in northcentral Colorado during the 1974-75 hunting season. (continued)

Area

Direct

Indirect

1
25

1
31
5

CENTRAL FLYWAY
Montana
Bighorn County
Phillips County
Treasure County
Garfield County
Roosevelt County
Valley County
North Dakota
Ward County
Wzoming
Albany County
Goshen County
Crook County
Nebraska
Brown County

1

6

1
1

1

4

1
1

1

2

Kansas
Phillips County

1

Texas
Randall County

4

New Mexico
Socorro County
Chaves County
San Miguel County

5
2
1

2

PACIFIC FLYWAY
Utah
Sanpete
Boxelder County

1
1

Idaho
Fremont County

1

Totals

73

82

�-46An additional three foreign banded birds (Table 2) were recaptured during
an essentially unsu~cessful post-season trapping and banding operation.
Eighty-seven of the foreign recoveries or recaptures were classified as wildtrapped locals when banded (Table 3).
The 1974-75 hunting season was the first recovery year for birds banded during
the post-nesting season by the cooperative Canada goose banding crew operating
in southern Alberta. The direct recoveries listed in Tables 1 and 3 from Schuler, Many Islands, Seven Persons and Murray Lakes are from these bandings. All
of the mentioned areas are located east of 111 longitude in southern Alberta.
A total of 22 direct recoveries were reported from 262 bandings on those four
areas (Weaver, unpublished data). Twelve of those banded birds were reported
taken in northcentral Colorado while the remaining ten were taken either in
Alberta or in other areas within the Hi-Line Canada goose range. The only
other banding area east of 111 longitude, Cavan' Lake, was represented only
by recoveries from Alberta. West of 111 longitude, a total of 1,084 birds
were banded on 11 areas. These bandings resulted in 98 recoveries of which
none were reported from Hi-Line wintering areas.
0

0

0

Geese released in the restoration program at Manito Lake east of North Battleford, Saskatchewan were represented in the northcentral Colorado harvest by
both direct and indirect recoveries. These birds were also represented during
the 1973-74 season (Szymczak and Staffon 1975).
No new areas were represented by substantial numbers of recoveries in northcentral Colorado. Recoveries from populations nesting at Dowling Lake, Alberta,
the Cypress Hills areas of Saskatchewan, Phillip County, Montana and the Big
Horn-Treasure County areas in Montana, all documented Hi-Line nesting areas,
were well represented in the harvest.

Table 2. Banding areas outside Colorado of Canada geese recaptured postseason 1974-75 in northcentral Colorado.

Area

Saskatchewan
Val Marie-Frenchman River Area
Cypress Hills Area

Direct

Indirect

1
1

CENTRAL FLYWAY
Montana
Phillips County

Totals

1

1

2

�-47Tail Fans
A total of 129 of the 167 tail fans collected through the "Waterfowl Parts
Collection Survey" from geese bagged in northcentral Colorado were used to
establish the comparative rate of harvest of "small" Canada geese in northcentral Colorado during the 1975-76 hunting season. An estimated seven,
eight and 22 percent of the geese harvested in the counties of Larimer,
Boulder, and Weld, respectively, were considered to be "small" Canada geese,
(Table 4) most likely members of the short grass prairie Canada goose population. Sample size in the other three counties listed were too small to yield
valid results.
Over the six year collection period the percent of "small" birds in the harvest has been extremely consistent in Larimer County, varying from 7 to 14
percent with a year to year average of 9 percent. Larimer is the only county
that has had seemingly adequate samples for each year throughout the six year
period. In Boulder County, in the 4 years represented by more than 20 tail
fans, the percent of "small" birds in the harvest varied from eight to 16,
averaging 13 percent. In Weld County also, only four of the six years are
represented by more than 20 tail fans. The number of "small" birds in the
harvest in those years has varied from 18 to 41 percent with a year to year
average of 29 percent.
Since the rate of tail fans collected is not proportional to the.rate of harvest by county, the tail fan data can't be pooled to obtain an estimate for
all birds harvested in northcentral Colorado. However, to obtain an estimate,
by county, a simple proportion involving the tail fans and harvest can be constructed:
No. Tail Fans for County
No. of "small" Tail Fans

County Harvest
X (No. small fans in harvest)

Solving the equation will give the estimated total number of "small" birds
harvested by the County. The numerical values are valid only to the degree that
harvest estimates are valid and sample tail fans are representative of the harvest. If harvest estimates aren't reliable, the percent of harvest composed of
"small" geese can be obtained by dividing the total estimated number of "small"
geese harvested in the total area as obtained from County calculations, by the
total estimated harvest. If harvest estimates errors are equally proportional
by county, the estimate of the percent of harvest composed of "small" geese
should be valid.
In 1974, the harvest for geese in northcentral Colorado was estimated through
the use of a state-wide waterfowl harvest survey for the first time since 1963.
The estimated harvest from that survey was a record 17,000 plus during a season
that was generally considered poor for goose hunting, indicating the harvest
figures might be in error. Therefore, valid numerical estimates of the harvest
of "small" birds can't be obtained. But a reliable estimate of the percent of
harvest composed of "small" birds can be obtained by calculating numerical estimates by county using the formula, then dividing the pooled county estimates
by the pooled county harvest. Calculating the estimate as described above resUlting in an estimate that 22.2 percent of the harvest was composed of "small"
birds in 1974. The percents calculated in the same manner for 1970, 1971 and
1973 using the results of the special harvest survey were 10.4 percent, 19.8
percent, and 12.9 percent. Data for 1972 was not used because of poor sample
sizes in two of the three major harvest counties.

•

�-48-

Table 3. Banding locations of Canada geese classified as wild-trapped locals
which were recovered or recaptured in northcentral Colorado during the winter
of 1974-75.

Year of Banding
Before
1967 1967 1968 1969 1970 1971 1972 1973 1974

Area -

Alberta
Edmonton-Camrose Area
Dowling Lake Area
Brooks Area
Schuler Lake
Seven Persons Lake
Murray Lake
Saskatchewan
Val Marie-Frencnman
(Masefield)
Cypress Hills Area
Pontiex-Notukeu R.
Crane Lake

1

1
6

1
1

1

1
2
3
2

R. Area

3
1

1

2

1
2

8
1

3
5

18

1

CENTRAL FLYWAY
Montana
Bighorn County
Phillips County
Treasure County
Garfield County

1

3

1

1

5

1
5
1
1

PACIFIC FLYWAY
Utah
Sanpete County
Boxelder County
Idaho
Fremont County

1
1

1

In summary, nearly one-quarter of the harvest in northcentral Colorado was
compc sed of "small" birds in 1974, indicating that a record number of "small"
geese w0re in northcentral Colorado during the 1974-75 season.

�Table 4. Classification of Canada goose tail fans collected through the Bureau of Sport Fisheries
Collection Survey, from birds harvested in northcentral Colorado, 1970-1975 hunting seasons.

1970

1971

Year
1972
Percent
Percent
Large
Small

County of
Harvest

Percent
Large

Percent
Small

Percent
Large

Percent
Small

Larimer

92 (36).!/

8 (3)

91(41)

9(4)

92 (22)

Boulder

100(1)

0(0)

86(18)

14 (3)

Arapahoe

-

-

100(2)

Adams

100(2)

0(0)

Weld

82(18)

18(4)

Morgan

-

-

1973

Parts

1974

Percent
Large

Percent
Small

8(2)

93 (28)

7 (2)

100(10)

0(0)

84(42)

0(0)

-

-

100(4)

0(0)

100(1)

59(13)

41(9)

-

-

and Wildlife's

Percent
Large

1975
Percent
Small

Percent
Large

Percent
Small

86 (31)

14(5)

93 (27)

7(2)

16(8)

88(21)

12 (3)

92(36)

8 (3)

-

-

-

-

100(1)

0(0)

0(0)

-

-

100(1)

0(0)

100(3)

0(0)

67(2)

33 (1)

79(11)

21(3)

67 (16)

33(8)

78(43)

22(12)

-

-

0(0)

100(4)

-

-

100(2)

0(0)

1/ Sample size in parentheses.
I
.j:&gt;\C

I

�-50-

Winter Banding - Northcentral Colorado
A total of 2,446 Canada geese were banded post-season in January and February
in northcentral Colorado from 1971 through 1975. Two hundred forty-one of
those birds, 187 "large" geese wearing size 8 bands and 54 "small" geese with
size 7 bands, were banded in January 1975. Percentages of "small" geese in
the annual banded samples varied considerably over the 1971-75 period. However, of the total banded, 769 or 31.4 percent were considered "small" Canada
geese.
Through each successive recovery year for each banding cohort, Alberta has
generally been the major recovery location in Canada for birds classified
when banded as "small" Canada geese. Alberta is also the major recovery area
in Canada for short grass prairie birds banded in southeastern Colorado
(Szymczak 1974). Unfortunately the majority of the "small" birds, 148 and
463, were banded during two banding periods in 1971 and 1972, respectively,
in northcentral Colorado. In order to test the hypothesis that "small" geese
in the wintering population in northcentral Colorado are not composed of a
simple cross-section of geese of the short grass prairie population, the
recovery distribution of two banding cohorts, 1971 and 1972, was composed by
banding location, northcentral and southeastern Colorado. The results as
presented in Table 5 indicate some obvious variations in the two populations
as reflected by recovery distribution.
The distribution of direct recoveries
between the two Provinces from the two banding areas was quite different in
1971 with Alberta accounting for 36 percent of the recoveries from southeast
Colorado bands, but 86 percent of the northcentral bandings. A closer percentage distribution was found in comparing indirect recoveries with Alberta
taking 70 percent of the southeast bandings and 50 percent of the northcentral bandings. Unfortunately 1971 comparisons are hampered by a small
number of recoveries in the Provinces from northcentral Colorado bandings.
In 1971, with better sample sizes from the northcentral area, the data indicates a similar distribution from both banding areas when considering direct recoveries with Alberta accounting for 52 percent and 57 percent of the
southeast and northcentral Colorado bandings, respectively.
The difference
is greater in 1972 when comparing indirect recoveries. According to goose
weights recorded during the post-season banding period, geese judged to be
"small" in the banded population in northcentral Colorado are physically
larger than birds banded in southeastern Colorado (Table 6). However the
differences by age class, although generally consistent, are small. Unfortunately complete taxonomic measurements were not taken to further comparisons.
The weight differences cJuld simply be an indication of variation in body
condition between banded birds of the two sample areas.
The annual interchange 0etween the two wintering areas, southeast and northcentral Colorado, by "small" birds continued during the 1974-75 hunting season. Two "small" birds (size 7), 12.5 percent of the total recoveries, in
northcentral Colorado since 1971-72 were taken in southeast Colorado during
the 1974-75 season (Table 7). A record 7.3 percent of the recoveries during
the 1974-75 season of southeast Colorado banded birds was reported taken in
northcentral Colorado (Szymczak, unpublished data).

�-51-

Table 5. Percentage comparison of the recovery distribution between Alberta
and Saskatchewan of "small" Canada geese banded post-season in 1971 and 1972
in southeast Colorado with those banded in the same years in northcentral
Colorado and recovered through the 1974 hunting season.

Banding Location
Year of Banding
and Recovery Location

Southeast Co1orado(1032)11
Direct
Indirect Total

Northcentral Co1orado(463)Direct
Indirect
Total

II

1971
Alberta
Saskatchewan

36(9)])
64(13)

70(19)
30(8)

57(28)
43(21)

86(6)
14(1)

50(3)
50(3)

69(9)
31(4)

Total

100(22)

100(27)

100(49)

100 (7)

100(6)

100(13)

Alberta
Saskatchewan

52 (11)
48 (10)

55 (17)
45(14)

54(28)
46(14)

57(8)
43(6)

76(13)
24(4)

68(21)
32(10)

Total

100(21)

100(31)

100(52)

100(14)

100 (17)

100(31)

1972

I

1./
J./

.

Banded sample size.
Number of recoveries

in parentheses.

Table 6. Comparison of weights in pounds of "small" Canada geese banded postseason in 1971 and 1972 in southeast Colorado with those banded in the same years
in northcentral Colorado, by age and sex class.

Year and
Location of Banding

Adult Male

Weights bl Age and Sex (lbs)
Adult Female Immature Male Immature Female

Southeast
Northcentral

5.75(265)
6.49(43)

5.20(282)
6.05(59)

5.35(151)
6.28(19)

4.70(169)
5.76(24)

Southeast
Northcentral

5.75(390)
6.13(151)

5.16(305)
5.77(164)

5.68(179)
5.67(79)

5.03(164)
5.24(61)

1971

1972

�Table 7. Percent distribution of recoveries through the 1974-75 hunting season of Canada geese banded post-season in northcentral Colorado during
the four year period 1971 through 1974.

1971-72
Area

1972-73

Hunting Season
1973-74
Size 8
Size 7

1974-75
SOize 8

All Years
Size 8
Size 7

Size 8

Size 7

Size 8

Alberta

13.8 (11)

40.0 (6)

12.6 (13)

34.5 (10)

10.2

(9)

37.5

(9)

11.3

(9)

25.0 (4)

12.0

(42)

35.8 (29)

Saskatchewan

18.8 (15)

6.7 (1)

27.6

17.0 (15)

12.5

(3)

10.0

(8)

12.5 (2)

18.5

(65)

17.3 (14)

Manitoba

7.5

(6)

13.3 (2)

26.2 (27)
1.0 (1)

(0)

(0)

2.0

(7)

2.5

(2)

(0)

(0)

3.4

(2)

4.2

(1)

6.3

(5)

(0)

3.4

(12)

2.5

(2)

(6)

4.2

Montana
Wyoming

(2)

(0)

(0)

1.0

(1)

(0)

(0) .

(0)

(2)

6.7 (1)

1.0

(1)

(0)

(0)

(0)

(0)

(0)

(0)

(2)

(0)

(2)

(0)

(0)

(0)

(0)

(0)

2.3
1.1

46.3 (37)
(0)
(0)

20.0 (3)
13.3 (2)
(0)

10.3
20.7

(3)
(6)
(0)

52.3 (46)
2.3 (2)
1.1 (1)

(0)

(0)

2.3

3.4

2.5

2.5

1.9

(1)

2.3
6.8

(0)

2.8

(10)

(0)

(1)

(0)

0.6

(2)

(0)

\Jl

(0)

(0)

(0)

0.9

(3)

1.2

(1)

'"

(0)

(0)

(0)

6.3 (1)

(0)

1.2

(1)

(2)

(0)

(1)

12.5 (2)

2.0

(7)

2.5

~2)

(1)

(0)

(0)

(0)

0.3

(1)

(0)

(4)
(5)
(0)

68.8 (55)
(0)
(0)

12.5 (2)
12.5 (2)
(0)

(2)

(0)

(0)

6.3 (1)

49.0 (172)
3.4 (12)
(2)
0.6
1.1
(4)

14.8 (12)
16.0 (13)
(0)
1.2 (1)

(1)

(0)

(0)

(0)

(0)

0.6

(2)

1.2

(0)

(0)

(1)

(0)

0.3

(1)

(1)

(0)

12.5 (2)

(0)

(0)

(0)

0.3

(1)

(0)

(1)

(0)

1.0

(0)

(0)

(0)

(0)

Texas

(0)

(0)

(0)

(0)

Oklahoma

(0)

(0)

(0)

(0)

New Mexico

Utah

1.3

(1)

(0)
1.1

(1)

16.7
20.8

4.2

(1)

Size 7

(0)

33.0 (34)
9.7 (10)
1.0 (1)
1.9 (2)

Arizona

(0)

(0)

Kansas
Colorado
Northcentral
Southeast
Other

(0)

(2)

North Dakota
Nebraska

(0)

(0)

2.5

(5)

(8)

4.9
1.9

Idaho
South Dakota

Size 7

1.3

1.3

1.3

(0)

(1)
(0)

3.7

----------------------------------------------------------------------------------------------------------------------------------------------------------

(3)
(0)

I

I

�Table 7. Percent distribution of recoveries through the 1974-75 hunting season of Canada geese banded post-season
the four year period 1971 through 1974 (continued).

1971-72
Area

Size 8

Size 7

1972-73 .
Size 8
Size 7

Hunting Season
1973-74
Size 8
Size 7
1.1

(0)

(0)

1.0

(1)

(0)

(2)

(0)

1.9

(2)

(0)

Pennsylvania

2.5
1.3

(1)

(0)

(0)

Unknown

1.3

(1)

(0)

(0)

Nevada
California

in northcentral

Colorado during

Size 7

All Years
Size 8
Size 7

1974-75
Size 8

(1)

(0)

(0)

(0)

0.6

(2)

(0)

(0)

(0)

(0)

(0)

1.1

(4)

(0)

(0)

(0)

(0)

(0)

(0)

0.3

(1)

(0)

(0)

(0)

(0)

(0)

(0)

0.3

(1)

(0)
I

V1

IN
I

Total

80

15

103

29

88

24

80

16

351

81

�-54Harvest Patterns
The distribution of harvest through the 1974-75 hunting season of Canada geese
banded post-season in northcentral Colorado during the four year period 1971
through 1974 is presented in Table 7. A record 68.8 percent of the 80 reported
recoveries of large geese (size 8) were harvested in Colorado all in the northcentral area. Alberta and Saskatchewan combined for an additional 21.3 percent
of the large geese recovered with the distribution of recoveries by Province
essentially equal. Twenty-five percent of the 16 small (size 7) geese reported
recovered were taken in Alberta while 12.5 percent or two recoveries each came
from Saskatchewan, Nebraska, northcentral Colorado, southeast Colorado and Texas.
Population Distribution
The December 1975 coordinated inventory in the Hi-Line wintering area resulted
in a total of 59,638 geese being recorded, about 4 percent below the 1973 level
(Table 8, Szymczak 1975). The January count resulted in the highest total recorded during January since coordinated counts began in 1970-71, or 58,436 birds.
In four of the previous five years, the reduction in population size between the
two inventories has variei from 23 percent in 1970-71 to 62 percent in 1973.
The reduction was always directly related to declining numbers in Colorado, and
could not be attributed to harvest alone. These reductions indicate a movement
out of Colorado, normally during the mid-to-late December period. Analysis of
weather data indicates that these movements are not directly related to specific
weather conditions. The impetus for these movements and the area(s) to which
the birds moved are still unknown.
Eighty-seven percent of the population was in Colorado during the December inventory and 93 percent during the January count. Nearly 55 percent of the birds
were located on four reservoirs inside closed areas during the December count:
College Lake, New Windsor Reservoir, Valmont Reservoir and Terry Lake (Longmont)
(Table 9). In January, those same areas held nearly 47 percent of the geese.
A record number of geese for a January count were observed in the BrightonGreeley-Fort Morgan area (Table 10).
Mortality Estimates
Population mortality estimates for "large" Canada geese, as calculated from hunting season recoveries of post-season banded birds in northcentral Colorado, did
not change significantly with an additional year of banding and subsequent recoveries. Through the 1973-74 hunting season estimated mortality rates ~alculated by the
composite dynamic method were .279 and .326 for first year and all year recoveries, respectively.
For recoveries through the 1974-75 season, those same estimates
were .287 and .316.
Table 8.

Location

Results of Hi-Line Canada goose population inventories, 1975-76.

December 18

January 9

Montana

891

131

Wyoming

4,668

2,227

Colorado

51,891

54,444

New Mexico

2,188

1,634

Totals

59,638

58,436

�-55Table 9.

Results of northcentral Colorado goose surveys, 1975-76.

Area

December 18

January 9

1,880
265
604
225
1,360
1,207
420
657
750
10,636
1,864
450
400
7,850
275
3,000
60
420
175
300
180
61

2,671

o

215

34,039

29,118

5,600
188
25
900
2

4,900
347

Fort Collins - Loveland
Lindenmeier Lake
Longs Pond
Richards Lake
Reservoir 116
Terry Lake
Watson Lake
Claymore Lake
Sterling Ponds
Ft. Collins City Park
College Lake
Fossil Creek Reservoir
Boyd Lake
Hollister Lake (Old Windsor Res.)
New Windsor Reservoir
Tinmath Reservoir
Woods Lake
Cobb Lake
Douglas Reservoir
Reservoir 1115
Millers Pond
Horsetooth Reservoir
Reservoir #8 Annex
Reservoir #5

Sub-total

o
o
o
454
395
132
1,500
675
5,732
1,722
300

o
5,427
750
8,900

o
15

o

o

230

o

Boulder - Longmont Area
Terry Lake
McCall Reservoir (Stamp)
Baller Reservoir
Union Reservoir
Gaynor Lake
Baseline Reservoir
Valmont Reservoir
Faivre Ponds

Sub-total

o

o
o

4

o

4,285

o

8,793
40

11,004

14,080

�-56Table 9.

Results of northcentral Colorado goose surveys, 1975-76 (continued).

Area

December 18

January 9

Denver Area
Bowles Lake Area
Cherry Creek Reservoir
Cooley Sand &amp; Gravel
Denver City Park
Green Gables Country Club
Kettring Park
Sloans Lake
Standley Lake
Washington Park
Youngfield Lake (Maple Grove)
Lowry AFB
Rocky Mtn. Arsenal
S. Colo. Blvd./Quincy
Gravel Pit (60th Rnd Lowell)
Quincey Lake

Sub-total

975
1,000+
225
llO
250
250
98
170
180
5

o
o

1,850
550
325
250
350
215
37
360
161
156
450
32
30
238
12

3,263

5,016

o
o
o

Brighton-Greeley·-Ft. Morgan Area
Barr Lake
Horse Creek Reservoir
Milton Reservoir
Latham Reservoir
Riverside Reservoir
Empire Reservoir
Jackson Reservoir
S. Platte e/Orchard
Seeley Lake
Prospect Reservoir

400
60
300
800
22
1,610
275
33
50
35

3,80o-!/

Sub-total

3,585

6,230

51,891

54,444

GRAND TOTAL

1/

Aerial count.

225
45
200
5001/
607-

o
1,4001/

o
o

�-57Table 10.

Winter inventories of the Colorado Hi-Line Canada goose population.

Count Date

Fort Co1linsLoveland

Longmont-Boulder
Denver

Brighton-GreeleyFort Morgan

Total

1967
January 9

9,739

2,883

991

l3 ,613

1968
January 10

12,217

4,029

678

16,924

1968-69
November 20
December 19
January 2 &amp; l3

15,848
20,905
19,693

3,461
4,236
4,874

2,667
1,170
775

21,976
26,3ll
25,342

1969-70
November 5
November 28
December 23
January 6

8,737
31,350
18,522
30,650

2,255
3,782
5,668
5,060

390
1,374
1,259
1,914

ll,382
36,506
25,499
37,624

1970-71
November 4
November 23
December 22
January 6

12,612
29,970
36,034
19,879

1,690
16,710
12,664
15,566

348
1,370
3,055
2,425

14,650
48,050
51,753
37,870

1971-72
November 4
November 23
December 21
January 10

25,699
31,072
31.516
19,1l7

1,815 1./
9,181
19,525
10,742

6,905
l3,525
4,476
3,661

34,419
53,788
55,517
33,520

1972-73
November 6
November 22
December 20
January 10

15,230
21, us
l3,841
l3,595

1,240 1/
5,764
19,290
18,709

1,050
4,981
1,225
1,405

17,520
31,863
34,356
33,709

1973-74
December 13
January 9

31,558
14,854

10,336
6,802

10,380
1,809

52,274
23,465

1974-75
December 11
January 8

28,491
17,986

15,076
12,980

4,104
1,645

47,671
32,6ll

1975-76
December 18
January 9

34,039
29,118

14,267
19,096

3,585
6,230

51,891
54,444

}j Denver area not included.

�-58San Luis Valley
Population Distribution
Inventories of Canada geese in the San Luis Valley during 1975-76 indicated a
wide spread distribution of birds during late November, a concentration of
birds on the Monte Vista National Wildlife Refuge in mid-December and a dispersion of geese from the refuge area in early January directly after the huntins season (Ta01e 11). The total numbers of birds by period were similar to
what was recorded during 1974-75 (Table 12).
Harvest
Three hundred permits were issued authorizing the harvest of one goose each in
the San Luis Valley during the 1975 hunting season. An estimated total of 256
active permit holders hunted an average of 4.5 days and harvested 86 geese for
a success rate of 0.34 (Tables 13 and 14). Rio Grande County was the major
harvest area followed by Alamosa County, a return to the normal harvest distribution which was reversed in 1974 (Table 14). In the entire permit area, the
number of hunters and hunter activity in 1975 was three percent and eight percent, respectively, above the 1974 level, while harvest increased an estimated
25 percent (Table 15).

Westcentral Colorado
Harvest
A total of 700 permits were issued authorizing the harvest of one goose each
during the 1975 season in westcentral Colorado, 100 for Garfield County and
300 each in Mesa and Delta Counties. In Mesa County, an estimated 256 active
permit holders hunted an average of 4.1 days taking an estimated 93 geese for
a success rate of 0.36 (Tables 16 and 17). In Garfield County, an estimated
80 permit holders hunted an average of 3.6 days harvesting an estimated 20
geese. In Delta County, which was opened to goose hunting for the first time
since 1966, an estimated 254 permit holders hunted an average of 4.2 days and
harvested an estimated 68 geese.
About 42 percent of the birds taken in Mesa County was reported harvested along
the Colorado River below Grand Junction and 36 percent within five miles of
Highline Lake. In Garfield County, 44 percent of the birds were reported harvested along the Colorado River between Rifle and Rulison and 31 percent along
the Colorado River within three miles downstream from Grand Valley. In Delta
County 63 percent of the harvest was reported taken along the Gunnison River
between Current Creek and Escalante Canyon.
The January, 1976 aerial inventory of Canada geese in westcentral Colorado indicated that 44 percent of the 1,356 geese were located in Mesa County, with
the remaining birds similarly divided between Garfield and Delta Counties
(Table 18). The total of 1,356 geese was about 10 percent below the January
1975 total (Table 19).

�-59Table 11. Results of Canada goose surveys during the 1975-76 hunting season in
the San Luis Valley. l/
Area

November 20

Monte Vista National Wildlife Refuge
Rio Grande River
Conejos River to State Line
Del Norte to Monte Vista
Monte Vista to Alamosa
Alamosa to Conejos River
Alamosa to State Line
Conejos River
San Luis Lake
Sanchez Reservoir
McIntire Springs
Russell Lakes
Sego Springs Area 2/

84

December 18

January 8

868

550

301
75
309
52

8
1

8
375
10

91

50
10
208

16

Total

1,230

959

827

l/

Survey conducted by C. Bryant and Staff, Monte Vista National Wildlife Refuge;
R. Clark and G. Saville, Colorado Division of Wildlife, and compiled by M.
Szymczak, Colorado Division of Wildlife.

2/

Warm spring or well due north of Sego Springs Ponds on north side of river.

Table 12.

Winter inventories of Canada geese in the San Luis Valley of Colorado.

Year
1970-71
1971-72
1972-73
1973-74
1974-75
1975-76

October

1,570
1,006
289

l/ Compared monthly inventories

Number of Geese ..l/
November
December

January

1,490
913
1,196
736
1,0l3
1,230

1,261
1,286
1,621
905
709 2/
827

for the five years generally taken within five

days of each other.

2/

Geese not concentrated.

1,050
1,255
1,079
774
934
959

Poor count.

�-60-

Table 13.

Hunter actiyit¥ and success in the San Luis Valley goose permit

area.

Year

Number
Permits
Issued

Est. Number
Active
Hunters

Average
Days Hunted

Average
Seasonal
Bag/Hunter

1970-71

200

164

3.4

0.65

1971-72

350

296

4.2

0.60

1972-73

400

311

4.6

0.82

1973-74

300

233

4.2

0.!+8

1974-75

300

249

4.3

0.28

1975-76

300

256

4.5

0.34

Table 14. Distribution of harvest, by county, in the San Luis Valley goose
permit area.

Year

Rio Grande

Alamosa

Saguache

Conejos

Costilla

Total

1970-71

74

31

2

0

0

107

1971-72

121

50

1

5

0

177

1972-73

195

51

2

1

6

255

1973

62

40

3

1

6

112

1974

27

37

2

3

0

69

1975

61

19

1

3

2

86

�-61-

Table 15. Hunting pressure and harvest in the San Luis Valley goose permit
area 1974 and 1975.

Estimated Number
1974
1975
Total individual hunters

Percent Change
from 1974

249 (± 5)

256(±. 5)

+ 2.8

Total hunter trips

1,068(±40)

1,156(±42)

+ 8.2

Total geese bagged

69(± 5)

86(± 6)

+24.6

Average hunter trips/hunter

4.3

4.5

Average bag/hunter

0.28

0.34

Average bag/hun t er trip

0.07

0.08

Table 16. Hunter activity and success in the west central Colorado special
goose hunting permit area.

Year

Number
Permits Issued

Est. Number
Active Hunters

Average
Days Hunted

Average Seasonal
Bag/Hunter

1971-72

150

112

2.9

0.29

1972-73

250

182

3.7

0.32

1973
Mesa Co.
300
Garfield Co. 66
Entire Area 366

254
46
300

3.9
3.7
3.8

0.30
0.20
0.28

1974
Mesa Co.
300
Garfield Co. 100
Entire Area 400

235
67
303

4.3
3.6
4.1

0.28
0.28
0.28

1975
Mesa Co.
300
Garfield Co. 100
Delta Co.
300
Entire Area 700

256
80
254
590

4.4
3.6
4.2
4.1

0.36
0.25
0.26
0.30

�-62-

Table 17. Distribution of harvest by year in the west central Colorado special
goose hunting permit area;

Year

Delta County

Mesa County

Garfield County

Total

1971

33

1972

58

1973

75

9

84

1974

66

19

85

93

20

181

1975

68

Table 18.

Canada goose inventory, west central Colorado, January 7 and 8, 1976.

Area

Number of Geese

Delta County

385

Mesa County

601

Garfield County

370

Total

1,356

�-63-

Table 19.

Winter inventories of Canada geese in west central Colorado.

Date

Number of Canada Geese

1967

January 17

71

1968

January 16

92

1969

January 2

307

1970

January 19

339

1971

January 4

879

1971-72

November 24
January 5

485
989

1972-73

November 21
January 11

329
1,183

1974

January 11

965

1975

January 10, 11

1,515

1976

January 7, 8

1,356

LITERATURE CITED

Szymczak, M. R. 1974. Arkansas Valley Canada goose flock management studies.
Colo. Div. of Wildl., Game Res. Rept., Fed. Aid Project W-88-R. October.
p. 55-87.
Szymczak, M. R., and R. Staffon. 1975. Studies of Canada goose populations
in Colorado transplant areas. Colo. Div. of Wildl., Game Res. Rept., Fed.
Aid Project W-88-R. October. p. 53-117.

Prepared by

'/ /i(&lt;:'-A~t!-L, IC! ~f:;;1-)J)CJd
.Michael R. SzymczaiP
Wildlife Researcher

Z&gt;

__k~
&lt;1.~

��October 1976

-65JOB FINAL REPORT

State of

COLORADO

---------------------

Project No.

W-88-R-21

Migratory
---..:;.::

Bird Investigations

2

Job Title

7
Job No.
Non-hunting Mortality Investigations
of Canada Geese in Southeastern Colorado

Period Covered:

October 3, 1972 through March 31, 1975

Work Plan No.

Personnel:

Michael R. Szymczak

ABSTRACT
All data has been collected and essentially analyzed. Portions of a draft
of a publication entitled "Lead Poisoning of Canada Geese in Southeast Colorado" have been completed. A second publication concerning the implementation
of a ban on the use of lead shot as a means of reducing or eliminating lead
poisoning of Canada geese at Turk's Pond has not yet been initiated. Formulation and publication of these manuscripts have been written as segment
objectives under Work ,Plan 6, Job 1, Migratory Bird Publications for the
period April 1, 1976 through March 31, 1977.

��October 1976

-67JOB FINAL REPORT

State of

COLORADO

Project No.

W-88-R-21

Work Plan No.
Job Title

2

Migratory Bird Investigations
Job No.

8

Investigations of Canada Geese in the San Luis Valley

Period Covered: April 1, 1973 to March 31, 1976
Personnel:
C. Bryant and Staff, Monte Vista National Wildlife Refuge; M.
Strong and R. Tragstaad, Bureau of Land Management; D. Lowry, Colo. State
University; R. Clark, R. Desilet, N. Forbes, J. Frothingham, H. Funk, B.
Goforth, J. Grieb, J. Gustafson, R. Holder, J. Rauch, W. Rutherford, G.
Saville, W. Schuett, S. Steinert, R. Velarde, R. Weldon, B.Widhalm, M.
Zgainer and M. Szymczak, Colorado Division of Wildlife.

ABSTRACT
In 1974 an estimated 76 and 39 breeding pairs of Canada geese (Branta
canadensis) were located on river habitat and lake and pond habitat, respectively, in the San Luis Valley, excluding the Monte Vista National Wildlife
Refuge. In 1975 using a different method of sampling and calculation than
in 1974, only 40 breeding pairs were estimated for river bottom areas and 19
pairs for lake and pond areas. About 89 percent of the encounters with
geese transported and released on, or wild trapped and banded at the Monte
Vista National Wildlife Refuge occurred in the San Luis Valley. Only 39
percent of the reported encounters with geese released in conjunction with
restoration progr'ams at San Luis Lakes or on the Alamosa National Wildlife
Refuge have occurred in the San Luis Valley. Most of the encounters outside
the San Luis Valley with geese released either at San Luis Lakes or on the
Alamosa National Wildlife Refuge have occurred in the Bitter Lake National
Wildlife Refuge area near Roswell, New Mexico. From 1969 through 1974
there were 19 encounters in the San Luis Valley with birds banded at Wheatland Reservoir, a moulting area for Canada geese located north of Laramie,
Wyoming, indicating birds of the San Luis Valley nesting population are
using Wheatland Reservoir for moulting. Encounters with Canada geese banded
on Hi-Line population production areas were first recorded in the San Luis
Valley in 1960 and were reported sporadically from 1960 until 1972. No
encounters with geese banded on Hi-Line production areas have been reported
for the 1972 through 1974 period, indicating reduced presence of Hi-Line
geese in the San Luis Valley. Encounters in the San Luis Valley with geese
banded as moulting adults in the Denver area indicate ponds in the Denver
area may have been used as moulting locations for non-nesting geese of the
San Luis Valley population at least during the late 1960's. January inventories in the San Luis Valley indicated a progressive buildup in the number
of wintering geese from 45 in 1955 to 1,621 in 1973. During the period from
1962 through 1970, 86 percent of the geese counted on the January inventory
in the San Luis Valley were on the Monte Vista National Wildlife Refuge in

�-68-

ABSTRACT (continued)

Rio Grande County. However, during that same 1962 through 1970 period
essentially all the harvest occurred in Conejos and Costilla counties,
indicating most of the harvest was being extracted from the migrant population. The decline in the size of the goose harvest in the San Luis Valley
from an estimated 794 in 1962 to 194 in 1969 generally paralleled the decline
in the number of Hi--Line Canada geese wintering in New Mexico. Annual
periodic counts beginning with the initiation of the special Valley-wide
goose season in 1970 indicated population fluctuations throughout the fall
and winter period. Beginning in 1974, counts of geese began to show a
definite decline in the population through the fall and winter, indicating
the movement of geese out of the San Luis Valley. The goose hunting success
rate since special seasons were initiated in 1970 seem to be directly
related to the average number of geese present in the San Luis Valley during
the hunting season. Since 1970 Rio Grande and Alamosa counties have been
the major harvest areas, while very few geese are taken in Conejos or
Costilla counties. In 1970 and 1971, 35 to 40 percent of the goose harvest
in the San Luis Valley occurred in November during a season beginning
November 1 and ending in mid-January.
In 1972, with the season during the
same time period, 20 percent of the harvest was taken in November.
In 1973,
1974 and 1975 only 21 percent, 20 percent and 29 percent, respectively, of
the harvest was taken in the month of November, with a November-December
season. Aerial counts indicate geese are present in Conejos and Costilla
counties in November; but according to harvest surveys they are subject to
very little hunting pressure. A direct relationship exists between the
distribution of geese late in the hunting season and the distribution of
harvest during the entire hunting season.

�-69RECOMMENDATIONS
1.

Assuming that the size of the current nesting Canada goose population
in the San Luis Valley is not adequate to meet the recreational demand,
the following steps should be employed to increase the population.
a.

Erect and maintain nesting structures on open lake and pond areas
on both public and private lands where adequate natural nest sites
are not available. San Luis and Head Lakes are excellent examples
of areas lacking natural nest sites.

b.

Erect and maintain nesting structures on sloughs and oxbows adjacent
to the Rio Grande River where adequate natural nest sites are not
available.

c.

Encourage the development through any means available of permanent
water areas at Mishak Lakes.

2.

Continue experimental Canada goose breeding population surveys under
the Waterfowl Production Surveys job.

3.

Continue post-nesting season Canada goose banding concentrating on
capturing groups of adults with goslings throughout the Valley, banding
at least 200 birds a year for three years.

4.

Encourage the preservation of the Wheatland Reservoir mOUlting area by
expressing to the Wyoming Game and Fish Commission the importance of
that reservoir to the San Luis Valley Canada goose population.

5.

Conduct annual counts of Canada geese in the San Luis Valley in mid- to
late October, mid-November and mid-December.
Schedule the counts so
they occur on approximately the same day each year.

6.

Until more information is obtained concerning emigration of the local
population and possihle differential harvest, maintain basically the
same hunting season regulations; 300 permits authorizing harvest of one
goose each throughout the San Luis Valley, November 1 to December 31
season framework.

7.

Continue the special San Luis Valley goose season harvest survey.

��-71-

INVESTIGATION OF CANADA GEESE IN THE SAN LUIS VALLEY
Michael R. Szymczak

This investigation was initiated primarily to document the size, distribution, and productivity of the Canada goose flock in the San Luis Valley.
The flock developed through restoration efforts by federal and state personnel which began in 1953. The general development of the nesting population
was not monitored except on the Monte Vista National Wildlife Refuge.
Wintering population numbers, which were thought to be directly related to
the size and productivity of the nesting population were determined through
counts conducted throughout the winter period. In 1970, in response to
increasing numbers of wintering geese, a limited goose hunting season was
initiated in the San Luis Valley (Szymczak 1971). Winter population distribution, hunter activity, and harvest were monitored under another job.
In 1973 this study was initiated with a three phase approach: (1) develop
a reliable technique to measure the breeding population annually, (2) document the distribution and productivity of the population and (3) attempt to
establish the relationship, if any, between the size and productivity of
the breeding popUlation and the size of the fall and winter population, and
if possible, develop a harvest formula based on the relationship.
A sampling method for estimating the breeding population utilizing aerial surveys was designed and tested in 1974 (Lowry 1974, 1975). Testing of the aerial
methods, helicopter vs fixed wing counts, were continued in 1975, and are
planned again in 1976. Investigations to document the distribution and
productivity of the population were conducted in spring and summer of 1973
and 1974, and reported on in final form by Lowry (1974, 1975). Lowry's
findings will be referred to, but not duplicated in this report. The final
phase of this study, which will be the subject of this report, was to tie
all available information together from all sources and develope an initial
management plan.
P. S. OBJECTIVE
To investigate the size, distribution, and productivity of the Canada goose
population in the San Luis Valley during the spring and summer period and
develop a management. plan.

METHODS AND MATERIALS
Aerial Breeding Pair Counts
Lowry(1975) divided the occupied and potential Canada goose nesting habitat,
with the exceptior. of the Monte Vista National Wildlife Refuge, into sample
sections. River sections were 1 x 3 miles with their lengths approximately

�-72-

perpendicular to the flow of the stream. Sections covering pond and marsh
areas were 1 mile wide and varied in length depending on the size of the
wetland.
In 1974 Lowry conducted breeding pair counts on selected sample
sections using both fixed wing and helicopter aircraft in order to test ,the
observability rate of breeding pairs from both types of aircraft. In the
fixed wing aircraft four passes were flown perpendicular to the longitudinal
axis of the sample area at 100 to 150 feet above the ground with two observers
scanning 1/8 of a mile on each side of the aircraft. In the helicoptor
Lowry made 8 to 12 passes across the sample area in the same direction as in
the fixed-wing at from 20 to 150 feet in altitude with only one observer.
In 1974 in order to obtain an estimate of breeding pairs, sections 3, 6, 13,
20 through 36, 42 through 45, 48, 50 through 53, 57, 61, 65, and 66 were
counted on the river (Lowry 1974). In addition, all pond and marsh area
sections were counted with the exception of T. All counts were made from
a helicopter.
Comparative fixed wing counts were flown over areas 3, 6, 13,
25 through 27, 30, 42 through 45 B, C, D, and P.
In 1975 two additional sampling areas were added along the Rio Grand River
downstream from La Sauses, creating the sampling grid as it appears in Fig. 1.
Also, for economy reasons, only four helicopter passes were flown across each
sample area in the same manner as with the fixed wing. Sample areas 2, 4, 6,
8, 10, 12, 14, 16, 18, 20, 22 through 32, 34, 36, 38, 40 through 48, 50, 52,
54, 56, 58, 60, 62, 64 through 69 and all lettered sections were flown with
the helicopter.
Comparative fixed wing counts were conducted on all the
above numbered river sections except 31, 32, 58, 60, 62, and 64 through 69.
Only lettered sections Q and T were flown with the fixed wing. As in 1974,
sample sections on the rivers were grouped by area in analysis (Table 1).
On all flights geese observed were recorded in one of the following categories:
singles on nests, pairs w/nests, singles not on nests, pairs or groups. In
analysis, the total number of singles and pairs with nests and singles not
on nests in excess of the number of singles with nests were recorded as
indicated breeding pairs. Totals were recorded by sample unit.
As recommended by Lowry (1974) flights were conducted just prior to the estimated peak of hatch (April 23 and 24, 1975) in order to record the greatest
number of expected productive breeding pairs.
All reported band encounters of Canada geese banded in the San Luis Valley
and encountered anywhere, or those banded elsewhere and encountered in the
San Luis Valley were examined. Inferences concerning migrational behavior
were drawn as a result of analysis of the age, status and time period of
banding and location and means of recovery.
Because of a lack of breeding population data breeding populations were not
considered in any population relationships. Winter inventories, harvest
estimates, harvest success and harvest distribution were compared both within
and between years in an attempt to establish relationships between winter
population and harvest. However, because of the lack of sufficient data, a
harvest formula for the San Luis Valley Canada goose population was not
developed.

�-73-

N

D~SlloA

'--"'---Co A 8S

f
SCALf

IN

W8l~':6on

KILOMETERS

o
J1
iE3E?"'"1E3E3
SCALf

IN

Y E3 H

r

COo

\.

Ie

~
~

MILES
10

I

o Center
Dol/or

o

°1

046

Head

Loire

E z
I"

(;.

lot

Son Luis
Lolr8

"ONTE

VISTA

NATIONAL
WILDLIFE

..

"
'"...

REFUGE

".•

..

•...
R.4E.

Fig. 1. Geographic location of sample sections utilized in San Luis Valley
Canada goose breeding pair counts.

�Table 1.

Comparative results of fixed-wing and helicopter estimated breeding pair counts of Canada geese in the San Luis Valley, 1975.

Sample
Sections

SamEle Size
Helicopter
Fixed-Wing

Indicated Breeding Pair
Helicopter
Fixed-Wing

Non-Nesting Pair
Helicopter Fixed Wing

GrouEs
Helicopter
Fixed-Wing

Total Geese
Helicopter Fixed-Wing

River
1-10
11-21
22-32
33-39
~O-4849-55
56-64
65-69

5
5
11
3
9
3
5
5

5
5
9
3
9
3

-

2
7
20
2
8
0
0
1

4"0

Sub-total

2
4
12
.5
9
2

-

-

Incomplete

2
9
23
2
9
0
4
0

2
6
34
0
23
0

-

6
33
23
0
13
7
13
0

l;9

Incomplete

95

-

0
13
0
0
3
0

-

Incomplete

14
65
109
8
47
7
21
2

8
33
92
10
67
4

273

Incomplete

Lake and Pond
A
B-C

I
....•

Entire Areas
II

D

II

E-H
I-K
L

II
II
II
II

0

II

P
Q
R
S
T

II
II
II
II

Sub-total

-

-

-

-

-

-

Entire Area

-

-

Entire Area

0
11
1
4
0
1
0
0
0
0
2
0

-19

-

-

-

-

-

0

-

0

Incomplete

0
9
0
1
0
0
0
0
4
0
0
0

-14

-

-

-

-

-

1

0

Incomplete

0
5
0
3
0
4
0
0
0
0
3
0

-15

-

-

-

-

0

-

0

Incomplete

J:I

0
45
2
13
0
6
0
0
8
0
7
0

0

81

Incomplete

--

2

�-75-

RESULTS AND DISCUSSION
Aerial Breeding Pair Counts
1974
Lowry (1974) compared the results of helicopter and fixed wing counts.
He found generally, that helicopter counts of breeding pairs with himself
as the observer were in closer agreement with ground counts than were
fixed wing counts with two observers. However, fixed wing counts were
consistent and thus could be corrected using a visibility ratio.
Because of sampling problems, Lowry (1974) used only helicopter counts as
a basis for estimating the expected productive 1974 Canada goose breeding
pair population in the San Luis Valley, excluding the Monte Vista National
Wildlife Refuge. He estimated 76 breeding pairs in the river habitat
(numbered sections) and 39 pairs in lake and pond habitat (lettered sections) for a total of 105 expected productive breeding pairs. These
estimates resulted from a projection based not only on areas sampled, but
also on a visibility ratio developed by Lowry for his aerial observations.
The correction factors were 1.00 in areas designated as wooded river bottom,
1.75 in open river bottom areas, and 1.16 in open lake and pond areas.
Because of Lowry's familiarty with the nesting population from ground
observations these correction factors are considered to be valid only for
his observations.
1975
Comparative results of fixed wing and helicopter estimated breeding pair
counts for 1975 are presented in Table 1. These counts have been subjected
to area correction only. Unfortunately, weather conditions did not allow
fixed wing counts on all sample sections. Non-nesting pairs and groups were
recorded but the results should not be considered indicative of the size of
the non-nesting segment of the population.
Using Lowry's 1974 data, a correction factor of 1.70 for wooded river bottom
areas and 1.45 for lake and pond areas had to be applied to fixed winged
data to obtain the same number of breeding pairs as with helicopter.
Because
of sampling problems, a correction factor for open river bottom areas (Section
40-48) could not be calculated. For 1975, the correction factor was 1.56
for wooded river bottom and 0.89 for open river bottom areas. This time a
correction factor could not be calculated for lake and pond areas.
The total estimated breeding pairs via helicopter counts for 1975 was 40 for
river bottom areas and 19 for lake and pond areas (Table 1). These estimates
are not comparable to Lowry's 1974 figures of 76 and 39, respectively,
because of his personal correction factor and his generally more intense
coverage of each sample section.
Experimental goose breeding population surveys in the San Luis Valley will
be continued by Migratory Bird Project, at least through 1976 as a portion
of the Waterfowl Production Surveys job, and therefore no specific recommendations concerning documenting the size of the nesting population will
be offered at this time.

�-76-

Migrational Behavior
Nesting Population-Winter

Migration

Nearly all Canada geese banded in the San Luis Valley have been marked in
conjunction with the goose restoration program in that area. Banding of
wild trapped geese, mostly locally raised goslings, has taken place periodically since 1960 on the Monte Vista National Wildlife Refuge. In the summer
of 1974 six wild trapped goslings were banded at Russell Lakes in the northern portion of the Valley, but these bandings didn't result in any recoveries through the 1974-75 hunting season. All other bandings and subsequent
recoveries have resulted from restoration releases. Unfortunately, the
recovery distribution of Canada geese, flightless goslings or adults, which
are transported into any area and released cannot be considered indicative
of the migrational behavior of the developing nesting population. However,
after analyzing the distribution of all recoveries of both restoration and
wild trapped geese along with recoveries in the San Luis Valley of geese
banded outside the Valley, some life history factors seem evident.
First, the population of Canada geese associated with the Monte Vista
National Wildlife Refuge during the spring and summer period seem to be year
round residents of the Valley except for possible moult migrations by nonnesting birds. Of 106 encounters (observations of previously banded birds)
of all Canada geese transported, banded, released, or wild trapped and
banded on the Monte Vista National Wildlife Refuge during the summer period,
only 12, or 11.3 percent occurred outside the San Luis Valley. Fifty-two
of the above encounters were of wild trapped birds banded during the 1969
through 1973 period on the Monte Vista Refuge. Only 6 of those encounters,
or 11.5 percent occurred outside the San Luis Valley. Four encounters,
two released birds and two wild trapped birds occurred at Wheatland Reservoir, north of Laramie in Albany County, Wyoming, a suspected moulting area
for sub adults of the San Luis Valley nesting population. One of the released
birds and two of the wild trapped birds were reported encountered through
recapture in north central Colorado.
In conjunction with the restoration program in the San Luis Valley, flightless goslings were banded and released at Russell Lakes in 1955, on the
Alamosa National Wildlife Refuge in 1967 and 1968, and at San Luis Lakes
in 1969, 1970, and 1971. The only recoveries (7) reported of the geese
released at Russell Lakes were taken in west central Saskatchewan.
Twentyeight recoveries have resulted from the birds released on the Alamosa Refuge
of which 11, or 39 percent, were reported taken in the San Luis Valley.
Forty-four reported recoveries have resulted from birds released at San Luis
Lakes, of which 17 or 39 percent were taken in the San Luis Valley.
Some of the birds released on the Alamosa Refuge and San Luis Lake since
1968 have been reported either harvested or recaptured during the winter in
the Roswell, New Mexico area at Bitter Lake National Wildlife Refuge. However, through July of 1975 only one bird reported recaptured at Bitter Lakes
has subsequently been encountered in the San Luis Valley. Another goose
recaptured at Bitter Lake was again encountered at Wheatland Reservoir the
following summer. Many of the birds recaptured at Bitter Lake have been
caught in more than one winter indicating an annual return to Bitter Lake.

�-77-

In short summary, it is obvious that the birds released in areas outside
the Monte Vista Refuge did not have the same affinity to the San Luis Valley
in terms of year round residency as did both the wild trapped and released
birds on the Refuge.
Analysis of encounters in the San Luis Valley of geese banded outside the
Valley provides very little data for substantiating the migratory behavior of
the local nesting population. The recovery data includes scattered reports
of geese banded in the winter in the Rio Grande Valley of New Mexico being
taken in following years in the San Luis Valley. Some of the reports are
of first year recoveries, definitely indicating the birds may have been
members of the San Luis Valley breeding population. However, in reviewing
the banding areas outside the Valley of geese taken in the Valley, it is
obvious that at least a portion of the Hi-Line Canada goose population
(Rutherford 1965) either migrated through the San Luis Valley on their way
to the lower Rio Grande River wintering area in New Mexico and/or terminated
migration in the San Luis Valley. Therefore, birds banded along the lower
Rio Grande may have simply been members of the Hi-Line population.
Nesting Population-Moult Migration
The possibility of Wheatland Reservoir serving as a moulting area for subadults of the San Luis Valley nesting population was mentioned in discussing
recoveries of San Luis Valley banded birds. Analyzing recoveries of foreign
banded birds definitely confirms Wheatland as a moulting area for the San
Luis Valley population. From 1969 through 1974 there were 19 encounters in
the San Luis Valley with birds banded at Wheatland.
Seven of these recoveries have been direct. Since 1969 Wheatland banded birds have accounted
for nearly 50 percent of all foreign banded birds encountered in the San
Luis Valley.
Winter Population-Ingress of Migrants
Analysis of reports of Canada geese that were encountered in the San Luis
Valley but banded elsewhere confirm that at least a portion of the Hi-Line
Canada goose population migrated through the San Luis Valley enroute to
wintering areas along the Rio Grande River in New Mexico. Encounters of
Hi-Line geese were first recorded in the fall of 1960 when two birds banded
in north central Montana were encountered in the San Luis Valley. Another
3 north central Montana banded birds were encountered in the fall of 1961,
and six in 1962. During that three year period, the Hi-Line population
increased from about 7,500 to 9,000 and about 65 percent of the population
was wintering in New Mexico, while the San Luis Valley terminal wintering
population numbered less than 250 geese (Table 2). In 1963 the southwestern
portion of the San Luis Valley was closed to goose hunting, thereby reducing
the opportunity for encount'ers with migrant Hi-Line geese. No geese from
Hi-Line production areas were reported encountered in 1963 or 1964. In
1965 and 1966 two and one birds, respectively, that were banded in north
central Montana were reported encountered in the San Luis Valley. In 1967
only one bird from a northern Hi-Line production area, Southern Alberta,
was encountered, but five birds, including four adults, released in restoration efforts in north central Colorado were encountered in the San Luis
Valley. Four of the five birds from north central Colorado reported encountered were victims of a fowl cholera (Pasturella multocida) outbreak on
the Monte Vista National Wildlife Refuge.

�-78-

Table 2. The results of winter Canada goose inventories
Valley, January 1955-January 1976. 1/

Year

October

Number of Geese
November

December

in the San Luis

January

1955

45

1956

102

1957

o

1958
1959

97

1960

114

1961

164

1962

50

1963

250

1964

465

1965

304

1966

260

1967

388

1968

400

1969

830

1970

1,225

1970-71

1,490

1,050

1,261

1971-72

1,570

913

1,255

1,286

1972-73

1,006

1,196

1,079

1,621

736

774

905

1974-75

1,013

934

709 ~/

1975-76

1,230

959

827

1973-74

289 ~/

l/Compared monthly inventories for 1970-71 through 1975-76 generally
within five days of each other.
2/
.
- Geese not concentrated.

Poor count.

censused

�-79-

In 1968 three birds from north central Montana and four transplanted adults
from north central Colorado were encountered, with some of the birds being
cholera victims. In 1969 the entire western portion of the Valley was
closed to goose hunting, the size of the migrant Hi-Line population wintering in New Mexico had declined to about 3,500 birds and no Hi-Line encounters
were reported in the San Luis Valley. In 1970 a special limited goose
season was initiated, which permitted the harvest of geese throughout the
San Luis Valley. Three birds from northern Hi-Line production areas and
two adult transplants from north central Colorado were encountered in 1970.
In 1971 two north central Montana birds were encountered followed by no
Hi-Line recoveries in 1972, 1973, or 1974. By 1974 the number of Hi-Line
geese wintering in New Mexico had declined to approximately 1,100 birds.
Analyzing further the presence of the north central Colorado adult transplants in the San Luis Valley results in some interesting findings. The
adult transplants, with the assumption they were members of the local population, were removed primarily from the Denver City Zoo and Park during
their moult in an attempt to alleviate an over crowded situation (Szymczak
1975). Most of the birds were released at New Windsor and Reservoir No.8
east of Fort Collins, but a few were placed at Valmont Reservoir near
Boulder. According to the encounters a number of these birds migrated into
the San Luis Valley. One explanation for the movement of these birds to
the Valley would be that they simply joined the migrant Hi-Line birds and
moved down to the San Luis Valley. This explanation, although possible, is
not realistic when considering all the data. First, birds from all three
release sites were reported encountered in the San Luis Valley and were
present there for at least three consecutive years. Second, if the birds
had been members of the non-migratory Denver area nesting population, or
even of the north central Colorado restoration population, which according
to Szymczak (1975) is non-migratory, they probably would have returned to
their natal areas and would not have been present in the San Luis Valley
at least for three consecutive years. Third, representatives of geese
released both on and off the Monte Vista Refuge for restoration purposes
and those banded as wild trapped goslings on the Monte Vista Refuge have
been reported recaptured during their moult in the Denver area. Szymczak
(1967) mentioned the possibility of immigration of Canadas to Denver for
the purpose of moulting. Considering all these factors, it seems logical
to conclude th-at at least a portion of the non-nesting adults and subadults
of the San Luis Valley population were using Denver area ponds for moulting
during the late 1960's.
The first Wheatland Reservoir banded bird was encountered in the San Luis
Valley in 1969, and according to subsequent encounters, Wheatland has been
increasing in importance as a moulting area since that time. Currently the
importance of the Denver area as a moulting area for San Luis Valley geese
is unknown. So apparently the Denver area was used for moulting by nonnesting geese of the San Luis Valley population. But now the moulting
migrants may be using Wheatland predominantly.
SUMMARY
According to analysis of all available encounters of banded birds associated
with the San Luis Valley, the following statements can be made.

�-80-

1.
Some members of the San Luis Valley nesting population migrate out of
the Valley into New Mexico in late fall and winter. However, the geese
associated with the Monte Vista National Wildlife Refuge stay in the Valley.

2. Some subadults and non-nesting adults of the San Luis Valley nesting
population utilize Wheatland Reservoir, north of Laramie, in Albany County,
Wyoming, and possibly reservoirs and ponds in the Denver area for moulting.
3. Portions of members of the Hi-Line Canada goose population have been
documented as migrating through the San Luis Valley enroute to wintering
areas along the lower Rio Grande River in New Mexico. However, with the
decline in the numbers of Hi-Line geese migrating into New Mexico, the
number of Hi-Line geese no longer have a significant influence on the size
of the Canada goose population in the San Luis Valley throughout the fall
and winter period.
4. Presently, the size of the fall and winter population in the San Luis
Valley in anyone year is almost completely dependent upon the size and
production of the local nesting population and the number of birds of that
population which migrate south to other wintering areas.
Relationship Between Breeding
Population-Wintering Population and Harvest
Breeding Population
Unfortunately, any past relationships involving the breeding population of
Canada geese in the San Luis Valley are hampered by the lack of breeding
population estimates. Estimates are available only for 1974 and 1975, but
they were not determined by the same method. Additional years of breeding
population estimates are needed before any meaningful relationships, if one
exists, can be established. As mentioned before, breeding population
surveys will be continued in future years.
Wintering Populations and Harvest
The size of the wintering population of Canada geese in the San Luis Valley
has generally been monitored since 1955. Until special limited Candada
goose season was initiated in 1970, counts were conducted only in early
January (Table 2). The January counts showed a progressive build up in the
number of Canada geese wintering in the San Luis Valley. During the period
from January 1962 through January 1970, 86 percent of the birds counted
were found on the Monte Vista National Wildlife Refuge located in Rio Grande
County. Harvest estimates indicate that no harvest occurred in Rio Grande
County until the special season was initiated in 1970, even though the
County was open to goose hunting until 1963 (Table 3). Essentially, all of
the reported harvest occurred in Conejos and Costilla counties southeast of
Rio Grande County. Harvest estimates, although probably inflated because of
sampling procedures, generally exceeded the size of the terminal wintering
population (Tables 2 and 3). Both the size and distribution of the harvest
indicated the majority of the harvest was being extracted from migrant geese,
most likely of the Hi-Line population. The decline in the goose harvest

�-81-

Table 3. Harvest estimates of Canada geese, by county, in the San Luis
Valley, 1955-1975. l/
Hunting
Season

Rio Grande

Alamosa

Number Harvested
Conejos
Costilla

Saguache

Total

1955

o

1956

o

1957

o

1958

o
541

1959

101

1960

61

1961

34

66

100

440

61

1962

26

199

569

794

1963

20

411

176

607

335

24

359

543

106

675

1966

120

240

1967

231

162

1964
1965

26

1968

39

58

1969

35

88

74 (80)

31 (80)

1971

121

50

5

1972

195 (248)

51 (63)

1

1973

62

40

1

1974

27

37

3

1975

61

19

3

390
393
97

71
(80) 2:)

1970

30

194
2 (39)

107 (279)

1

177

6

2

255 (311)

6

3

112

2

69

1

86

2

Estimates through 1969 and those in parentheses in 1970 and 1972 obtained
from statewide small game harvest survey. Other estimates from 1970 through
1975 from special survey.

uCostilla County not included in Special Goose season area in 1970.

�-~-

from its peak in 1962, which also was the year the largest number Hi-Line
production area banded birds were encountered in the San Luis Valley,
until the special Valley wide season was initiated in 1970 generally
followed the decline in the number of Hi-Line Canada geese wintering in
New Mexico (Table 3) (Szymczak 1975).
Periodic counts throughout the fall and winter period were initiated in
conjunction with the special season in 1970. These surveys revealed population fluctuations throughout the fall period indicating movement into or
through the Valley by the remnant migrant population and emigration by the
local nesting population. Still the terminal wintering population continued to increase until January 1973 (Table 2). Beginning in 1973 the fall
and winter population began to show a definite decline. Census results in
1974-75 and 1975-76 indicated movement out of the San Luis Valley as the
season progressed. According to January inventories, wintering numbers of
ducks have shown a similar annual population fluctuation reaching a peak of
64,000 in January 1971 and declining to 20,000 by January 1976 (Szymczak et
al. 1971; Funk 1976). Fall plowing of barley fields in the San Luis Valley,
which reduces the waterfowl food supply, is suspected as being instrumental
in the decline of the winter duck population and may be affecting the geese
as well. The number of wintering ducks have declined in spite of a fairly
stable breeding population in the San Luis Valley.
The harvest of Canada geese has been closely monitored in the San Luis Valley
since the special season was initiated in 1970. The regulations specified
the issuance of a number of permits, one to a hunter, authorizing the harvest
of one goose. The number of permits issued and hunter activity throughout
the six year period are shown in Table 4. The harvest survey has indicated
that the success rate seems to be related to the average number of geese
present in the San Luis Valley during the hunting season in the November to
January period (Table 5).
Since initiation of the special season, a dramatic shift in the distribution
of goose harvest in the San Luis Valley has occurred. Harvest in Conejos
and Costilla counties, which begau declining in the late 1960's, has been
essentially non-existent since 1970 (Table 3). Rio Grande and Alamosa
counties are now the major harvest areas. The special season shifted
hunting pressure and harvest to the previously closed areas of the Monte
Vista Refuge vicinity and the Rio Grande River between the towns of Monte
Vista and Alamosa. In the first two years of the special season with a
November 1 opening date, the harvest was well distributed by time period,
with 35 to 40 percent of the harvest occurring in November, 30 to 40 percent
in December, and the remainder in the first half of January.
In 1972, with the season during the same time period, only about 20 percent
of the harvest was taken in November. In 1973, 1974, and 1975 only 21 percent, 20 percent, and 29 percent, respectively, of the harvest was taken
in the month of November with only a two month November-December season.
So hunting pressure and harvest has evolved from a wider distribution by
time period in the first two years of the special season, to a concentrated
late season hunt primarily in the Monte Vista Refuge area in the 1972
through 1975 period. At the same time, when hunting pressure and harvest

�-83-

Table 4. Hunter activity during the San Luis Valley Special Canada goose
seasons, 1970-71 through 1975-76.
Number
Permits
Issued

Est. Number
Active
Hunters

Average
Days Hunted

1970-71

200

164

3.4

1971-72

350

296

4.2

1972-73

400

311

4.6

1973-74

300

233

4.2

1974-75

300

249

4.3

1975-76

300

256

4.5

Year

Table 5. Comparison of the average number of Canada geese present in the
San Luis Valley, by year, during the November to January hunting period
with the harvest success rate during the Special Canada goose seasons 1970-71
through 1975-76.
Average Number of
Geese November-January

Seasonal
Success Rate

1970-71

1,267

0.65

1971-72

1,151

0.60

1972-73

1,299

0.82

1973-74

805

0.48

1974-75

974 1/

0.28

Year

1975-76

1,005

v Does not include incomplete count conducted in January.

0.34

�-84-

were well distributed by time period in 1970 and 1971, very few geese,
according to aerial counts, were located in Costilla or Conejos counties
(Table 6). Then beginning in 1972 as the special season evolved into
primarily, a late season hunt, the percent of birds recorded in Costilla
and Conejos counties on November surveys increased (Table 6). So even
though birds are available in Costilla and Conejos counties sometime during
the special season, they are not hunted, and most pressure is applied to
and harvest extracted from, those birds associated with the Monte Vista
Refuge late in the hunting season. According to counts, many of the geese
in the Costilla-Conejos area in November must move onto the Refuge in
December and January, but a portion of the birds probably move out of the
Valley and therefore are not subject to much hunting pressure during the
special season.

Table 6. The comparative percents of Canada geese recorded in Costilla
or Conejos counties during counts in the San Luis Valley 1970-71 through
1975-76.

Geese in Costilla or Conejos Counties by Month of Count
November
December
January
No.
Percent
No.
Percent
No.
Percent

Year

1970-71

80(1,490)

5

3(1,050)

&lt;1

195(1,241)

16

1971-72

30(

9l3)

3

10(1,255)

1

55(1,286)

4

1972-73

240(1,196)

20

309(1,079)

29

40(1,621)

2

1973-74

334(

736)

45

283(

774)

37

O(

905)

0

1974-75

776(1,0l3)

76

l47(

934)

16

48(

709)

7

1975-76

371 (1,230)

30

9l(

959)

9

259(

827)

31

Summary
The lack of annual consistent estimates of the breeding population prohibits
the development of examining the relationship between the breeding population and the wintering population of geese in the San Luis Valley. In
addition, data indicating emigration by at least a portion of the nesting
population of geese during the fall and winter period hampers the development of valid relationships.
There seems to be a somewhat direct relationship between the size of the
goose population during the season and harvest. Also, a direct relationship

�-85-

exists between the distribution of geese late in the hunting season and the
distribution of harvest during the entire hunting season. Since the harvest
is concentrated in certain areas, it may very well be affecting only
certain groups of birds, while others remain essentially unharvested.
Harvest Formula
Unfortunately, not enough information is available at this time to develop a
harvest formula for geese in the San Luis Valley. Most information indicates that migrants of the Hi-Line population no longer influence significantly the number of Canada geese in the Valley during the fall and winter
period. Apparently, most harvest will be extracted from birds of the local
nesting population. But emigration of portions of the local population
resulting in possible harvest outside the Valley must also be considered,
as must possible differential harvest of the local population resulting
from movement of a segment(s) of the nesting population out of the Valley.
In order to document and/or measure both emigration and differential harvest,
the banding of the local population was initiated in the summer of 1974.
Unfortunately, banding efforts have been essentially unsuccessful, with
10 and 19 birds marked in 1974 and 1975, respectively.
Publication
A draft of a short publication concerning the use of a helicopter to census
breeding pairs of Canada geese in the San Luis Valley was prepared. After
reviewing the draft the decision was made to incorporate additional years
of data. Future work on the publication will be funded through another
federal aid job.

LITERATURE CITED
Funk, H. D. 1976. The 1976 January Waterfowl Inventory, Colorado.
Div. of Wildl. 5 p. proc.

Colo.

Lowry, D. G. 1974. Canada geese in the San Luis Valley, Colorado.
Thesis, Colo. State Univ., Fort Collins. 76 p.

M. S.

1975. Investigations of Canada geese in the San Luis Valley.
Div. of Wildl. Fed. Aid Game Res. Rep. Oct. pp. 135-173.

Colo.

Rutherford, W. H. (ed.). 1965. Description of Canada goose populations
common to the Central Flyway. Central Flyway Waterfowl Council Tech.
Comm. 20 p.
Szymczak, M. R. 1967. Breeding biology of Canada geese of the MetroDenver area. M. S. Thesis, Colo. State Univ., Fort Collins. 83 p.

�-86-

Szymczak, M. R., C. R. Hayes, and H. D. Funk. 1971. The 1971 January
WatErfowl Inventory, Colorado. Colo. Div. Wildl. 7 p , Proc.
1971. Studies of Canada goose populations in Colorado transplant
areas. Colo. Game, Fish and Parks Dept., Fed. Aid Game Res. Rep.
Oct. pp. 165-189.
1975. Canada goose restoration along the foothills of Colorado.
Colo. Div. of Wildl. Tech. Pub 1. No. 31. 64 p .

Prepared by

.t7JiL~X.

i~hael R. szymcz~
Wildlife Researcher

J

,

�October,

-87-

1976

JOB FINAL REPORT
State of

CO_L_O_RAD
__ O

Project No.

W-88-R-2l

Work Plan No.
Job Title

Personnel:

Migratory Bird Investigations
Job No.

3

Investigation

Period Covered:

_

of Mallard Management

6
Units of Eastern Colorado

April 1, 1965 - March 31, 1976

Richard M. Hopper

ABSTRACT
Eleven papers have already been prepared that partially fulfill the reporting
responsibility of this job. Additional papers are currently being prepared
for final fulfillment of this obligation.
At least one paper will be
submitted to The Journal of Wildlife Management, while others will become
a part of a Division Special Report or Technical Report. Completion of
these papers will be accomplished under Work Plan 6, Job 1.

��-89INVESTIGATION OF MALLARD
MANAGEMENT UNITS OF EASTERN COLORADO
Richard M. Hopper

P. S. OBJECTIVE
To develop a harvest formula for eastern Colorado management units.

METHODS AND MATERIALS
Compile and analyze all data collected as part of this Job that hasn't
previously been reported upon.

RESULTS AND DISCUSSION
Field work for this Job was completed as of March 31, 1976. Following
Segment 17 (1971-72), field work for this job involved only trapping
and banding of mallards on a monitoring basis while in the process of turning
over the operation to the Northeast and Southeast Regions. Numbers banded
in Segment 21 are reported by Hopper (1976) under Work Plan 1, Job 2, as
was the case in all previous years. Therefore, the final report for Work
Plan 3, Job 6 is limited to data collected prior to Segment 17. Office
work since then has been largely that of tabulating and analyzing data
for the final report. A periodic update of banding data for all years
of banding will be conducted under a new W-88-R Job (Work Plan 3, Job 8).
Many of the findings of this investigation have already been reported upon
in one fashion or another; some as special reports or publications, and
others as direct contributions to reports of another agency. Thus, the
following reports represent partial fulfillment of reporting responsibilities
for Work Plan 3, Job 6:
1.

Funk, H. D., J. R. Grieb, G. F. Wrakestraw, D. Witt, and G. W. Merrill.
1967. The Central Flyway High Plains Mallard Management Unit.
Unpubl. Central Flyway Tech. Corom. Rept. (Suppl. Rept.), 20pp.

2.

Funk, H. D., J. R. Grieb, G. F. Wrakestraw, and D. Witt. 1968. A
proposed mallard drake season in Central Flyway Montana, Wyoming,
and Colorado. Unpubl. Central Flyway Tech. Corom. Rept., l2pp.

3.

Funk, H. D., J. R. Grieb, D. Witt, G. Schildman, G. W. Merrill,
and G. F. Wrakestraw.
1969. A proposed experimental pointsystem duck season for the western portion of the High Plains
Mallard Management Unit. Unpubl. Central Flyway Tech. Corom.
Rept., llpp.

�-904.

Funk, D. H., R. M. Hopper, J. R. Grieb, D. Witt, G. F. Wrakestraw,
T. Kuck, D. E. Timm, and G. W. Merrill. 1970. Preliminary
evaluation of the 1969-70 experimental point-system duck
season within the High Plains Mallard Management Unit of the
Central Flyway. Unpubl. Central Flyway Tech. Comm. Rept.,
36pp.

5.

Funk, H. D., J. R. Grieb, D. Witt, G. F. Wrakestraw, G. W. Merrill,
J. Sands, T. Kuck, D. Timm, T. Logan, and C. D. Stutzenbaker.
1971. Justification of the Central Flyway High Plains Mallard
Management Unit. Unpubl. Central Flyway Tech. Comm. Rept.,
48pp.

6.

Grieb, J., H. Funk, D. Witt, G. Wrakestraw, and L. Serdiuk.
1966. A proposed mallard management unit for the Central
Flyway. Unpubl. Central Flyway Tech. Comm. Rept., 32pp.

7.

Grieb, J. R., H. D. Funk, R. M. Hopper, G. F. Wrakestraw, and
D. Witt. 1969. Preliminary evaluation of the 1968-69
experimental mallard drake season in the Central Flyway
portions of Montana, Wyoming and Colorado. Unpubl. Central
Flyway Tech. Comm. Rept., 26pp.

8.

Grieb, J. R .. H. D. Funk, R. M. Hopper, G. F. Wrakestraw, and
D. Witt. 1971. Evaluation of the 1968-1969 experimental
mallard drake season in Montana, Wyoming and Colorado. Trans.
N. Am. Wildl. Conf. 35:336-348.

9.

Hopper, R. M., and H. D. Funk. 1970. Reliability of the mallard
wing age-determination technique for field use. J. Wildl.
Manage. 34(2):333-339.

10.

Kimball, C. F., R. A. Bishop, C. D. Crider, J. H. Dunks, R. M. Hopper,
and D. D. ~ennedy. 1971. Analysis of the l2-state point
regulation test, 1970, based on the hunter performance survey.
U. S. Fish and Wildl. Serv., Migratory Bird Pop. Sta. Admin.
Rept. No. 206, 22pp.

11.

Kimball, C. F. 1972. Results in the hunter performance survey in
the point regulation test states, 1971. U. S. Fish and Wildl.
Serv., Migratory Bird Pop. Sta. Admin. Rept. No. 215, 2lpp.

Other papers are in the process of being prepared under Work Plan 6, Job 1.
These will serve to finalize the reporting responsibility for Work Plan 3,.
Job 6. At least one paper will be submitted to The Journal of Wildlife
Management for publication, hopefully by mid-summer of 1976. The bulk of
the final report will be published as a Division Special Report or Technical
Bulletin.

�-91-

LITERATURE CITED

Hopper, R. M. 1976. Trapping and banding ducks and geese.
Wi1d1., Fed. Aid Game Res. Rept., Oct. (in press).

Prepared by

~~/
71'. ~.&lt;-c.=Chard M. Hopper ~.
Wildlife Resea~cher

Colo. Div.

��October, 1976
-93JOB FINAL REPORT

State of

Colorado

---------------------

Project No.

W-88-R-2l

\-lorkPlan No.

4

Job Title

Migratory Bird Investigations
Job No.

3

Trapping and Banding Doves

Period Covered:

April 1, 1964 through March 31, 1976

Personnel: Ken Baer, Jack Frost, Charles Hayes, Dale Horne and Jack Randall
of the U.S. Fish and Wildlife Service; Personnel of Ft. Carson, U.S. Dept.
of Army; W. John Arthur, Scott Bessire, William Borden, William Brandes,
Clait Braun, William Carpenter, Jack Corey, Dwayne Finch, Howard Funk, James
Goodyear, Larry Green, Jack Grieb, Al Heins, Al Hemmert, Gl~n Hinshaw, Richard
Hoffman, Richard Hopper, J. Edward Kautz, Marie Vendeville Kautz, Thomas
Kitzmiller, Wilbur Ladd, Charles Loeffler, Chet McCord, Ron Oakleaf, Brett
Petersen, Gary Robinson, Michael Robinson, Brent Smith, Marvin Smith, Norwin
Smith, Steve Steinert, Mark Stromberg, Darryl Todd, Willie Travnicek, Jack
Vayhinger, Ken Wagner, Mike Watkins, Bert Widhalm, J. Allen White, and Gary
Will of the Colorado Division of Wildlife.

ABSTRACT
Mourning doves (Zenaida macroura) were trapped and banded in Colorado through
a cooperative state and federal program from 1964 through 1974. Primary
objectives were to investigate migration patterns and mortality rates. A total
of 31,523 doves, mostly free flying, were trapped and banded in the study interval whereas only 2,702 had been banded prior to 1964 (1913-1963).
Distribution of bandings in the 1964-1974 interval was about 6,800 on the eastern
prairies, 6,000 west of the Continental Divide and about 18,700 along the
foothills and intermountain valleys east of the Continental Divide. Adult
males outnumbered females (57 to 43 percent) in trap samples while age ratios
of adults and immatures were similar (51 to 49 percent respectively).
Recaptures of previously banded birds were few and contributed little to overall
understanding of migration paths and mortality rates. Chronology of hatching
was compared between those immatures trapped and banded (14,560) and those for
which wings were collected from hunters (4,792). These data support the
hypothesis that most early hatched young were not available to hunters during
the hunting season in Colorado. Early migration of early hatched young was
definitely documented. Age ratios in the harvest approximated 1.4 immatures per
adult or 2.8 young per adult female. Hunter pressure and harvest greatly increased during the 1964-1974 period although probably not as much as indicated
by the annual survey of small game license purchasers.
Analysis of 683 recoveries of doves banded in Colorado (53 from bandings in the 1913-1963 interval, 630 from bandings in the 1964-1974 period) revealed that most (449)
were from within the Central Management Unit with smaller numbers (146) from
other countries, and other management units (W.M.U. = 33, E.M.U. = 2). Mexico,

�-94ABSTRACT

(Continued)

primarily the states of Ja1isco and Michoacan, was the major recovery
area for non United States recoveries (141 of 146). Within the United
States, Colorado was the leading recovery state (351) followed by Texas
(48), New Mexico (43), Arizona (19), and California (10). No more than
three recoveries were reported from any other state or country. Thus
migration of Colorado banded doves was primarily south-southwest.
Doves
banded outside of Colorado comprised about 20 percent of the annual number
of recoveries reported from within Colorado.
Most of these doves originated
north and northeast of Colorado with Montana being a major state of origin.
Survival rates for all doves banded in Colorado in the 1964-1974 period
averaged .523 annually.

RECOMMENDATIONS
1.

Reanalyze recovery data and calculate survival and mortality
by age, sex and geographic area of banding.

2.

Prepare manuscript
Report Series.

for the Colorado Division of Wildlife

rates

Special

�-95-

TRAPPING AND BANDING DOVES
Clait E. Braun

MOurning doves are important game birds throughout much of North America
as they are widely distributed and occur in huntable numbers throughout
most of their range (Peters 1961, Keeler 1976). In Colorado, this migratory
species nests throughout the state, primarily below 8,500 ft elevation
(Bailey and Niedrach 1965) and can be found from above timberline (Braun,
unpublished data) to river bottoms, prairies, and sagebrush (Artemisia spp.)
semi-deserts during spring and fall migration. Prior to advent of this study
in 1964, little was known of the timing of nesting, migration paths, wintering
areas, harvest patterns, and survival and mortality rates of doves seasonally
resident in Colorado. Previous studies in Colorado (Funk 1965) emphasized
timing of late summer and fall migration of doves through the state. Due
to the obvious need to more fully understand the biology of this migratory
species, this study was initiated in 1964. Cooperative banding studies
throughout the Central Management Unit as delineated by Kiel (1959) did not
get underway until after 1967. This report summarizes all banding and recovery data in two periods, 1913-1963 and 1964-1975, trapping and banding
data from 1964-1974, and derived hatching dates from bandings and harvest
collections from 1964-1975. Detailed survival and mortality studies of
doves banded in three geographic areas of Colorado will be presented in a
cooperative report which will assimilate banding and recovery data from the
entire Central Management Unit.

P. S. OBJECTIVE
To investigate migration patterns and mortality rates of mourning doves
banded in Colorado by age, sex and area.

METHODS AND MATERIALS
Mourning doves were trapped in suitable locations such as feedlots, fallow
fields, edges of little used roads, harvested grain fields, near watering
sites and adjacent to roosting areas. Most success was experienced at livestock feeding sites, fallow fields and near roosts. Most doves were captured
through use of 3 ft square cage type, wire funnel traps similar to the modified Kniffin dove trap described by Reeves et al. (1968). A few doves were
captured through use of cannon nets (Braun 1976) incidental to capture of
band-tailed pigeons (Columba fasciata). Prior to 1967, some doves were
captured in mist net~ following procedures described by Reeves et al. (1968)
Most doves captured were free flying but prior to 1967, a few nestlings were
banded. Bait used to attract doves varied with site but wheat (Triticum
aestivum), millet (Panicum spp.) and milo (Sorghum vulgare) were most frequently used. Wheat appeared to attract doves more readily in most areas of
the state.
All doves trapped were banded with U.S. Fish and Wildlife Service bands, size
3A, except in 1974 when some size 3 were used. Rewards bands (N=122) were

�-96-

placed on a sample of immature (RY) doves banded in 1971 in conjunction
with a Central Management Unit-wide study of reporting rates. Doves banded
were classified to age (HY=immature, AHY=adult) and sex (adult male and
female and a few immatures molting past primary 3) (Reeves et al. 1968).
A few adults were classified as unknown sex each year. The primary molt
of most immature doves banded was recorded for estimation of hatching
dates (Allen 1963). Each year doves were trapped only in the June through
mid August period with goals of 1,000 birds in the eastern prairies, 2,000
along the foothills and intermountain valleys east of the Continental
Divide and 1,000 west of the Continental Divide (Ruos 1968). Prior to
1969, Colorado had a suggested quota of 2,000 birds per year (Wight 1963).
Collections of wings of hunter harvested doves were made in some years to
ascertain age ratios in the harvest and hatching dates of immature birds.
Distribution of recoveries and recaptures from all Colorado bandings for
the 1913-1963 and 1964-1975 periods was ascertained from a recovery deck
obtained from the Bird Banding Laboratory, U.S. Fish and Wildlife Service.
Similarly, recoveries and recaptures from doves banded outside of Colorado
but recovered within the state were obtained from the Bird Banding Laboratory.
Distribution of recoveries and recaptures was grouped by Dove Management
Unit and State. Preliminary analysis of survival and mortality rates for
all doves banded in Colorado was by the relative recovery rate method (Geis
1972).

RESULTS AND DISCUSSION

Trapping
Trapping and banding of mourning doves in Colorado apparently was initiated
in 1913 when one bird was reported banded. Through 1974, 34,225 individual
doves had been banded with 2,702 being banded from 1913 through 1963 and
31,523 in the 1964-1974 period (Table 1).

Table 1.

Interval

Numbers of mourning doves banded in Colorado, 1913-1974.
Prior to 1964
No. Banded

Year

1964-1974
No Banded

1913-1920
1921-1930

1
156

1964
1965

295
808

1931-1940

17

1966

220

1941-1950

75

1967

2,232

1951-1960
1961
1962

2,096

1968

2,501

308
31

1969
1970

3,390
3,159

18

1971
1972
1973
1974

5,356
4,633
4,409
4,520

1963

Totals

2,702

31,523

�•

-97-

Most trapping and banding was accomplished by personnel of the Colorado
Division of Wildlife (24,408) and U.S. Fish and Wildlife Service (8,723)
with more than 100 birds being banded by only two other permittees, R. A.
Ryder (236) and D. Hein (212) both of Colorado State University.
The remaining 646 doves were banded by 42 different permittees over a 63 yr period.

Geographic Area of Banding
Trapping and banding efforts for mourning doves were not uniform by geographic area or by time intervals. With suggested quotas after 1968 of
1,000 doves west of the Continental Divide, 2,000 along the foothills and
valleys east of the Continental Divide, and 1,000 on the eastern prairies,
it is obvious from Table 2 that quotas were not met prior to 1971.

Table 2. Number of mourning doves banded by geographic area and year,
1964-1974. 1/
Foothills and Valleys 1/
West of
East of Continental Divide Continental Divid~/
No. Banded
No. Banded

Year

Eastern Prairie~/
No. Banded

1964

226

48

0

1965

105

679

0

1966

o

199

0

1967

761

1,373

0

1968

1,217

1,119

0

1969

477

2,114

703

1970

480

1,876

527

1971

1,000

2,756

1,477

1972

944

2,465

1,146

1973

780

2,619

1,000

1974

900

2,399

1,200

Totals

6,890

17,647

6,053

1/
- D.O.W. and U.S.F. &amp; W.S. bandings only.

2/
- Includes sites at Vineland, Fort Morgan, Rocky Ford, Springfield,
Fort Lyons, and Bonny Reservoir.

Pritchett,

3/

- Includes sites at Denver, Fort Collins, Monte Vista, Fort Garland, Longmont,
Salida, Fort Carson, LaVeta, Evergreen, Aspen Park, Woodland Park, Estes Park
and Rye.
~/Includes sites at Durango, Arboles, Montrose, Sanborn Park, Craig, Lay, Hayden,
Meeker, Buford, Unaweep Canyon, Carbondale, and Stoner.

�-98-

While doves were banded at many sites in the 1964-1974 interval, a relatively small number of sites produced the majority of doves banded.
Major sites trapped, number of doves banded, and number of years of banding
are presented in Table 3 by geographic area.

Table 3.

Major mourning dove trapping areas, Colorado, 1964-1974.1/,11

Eastern Prairies
No.
Location
Banded Years

Foothills and Valleys
E. of Continental Divide
No.
Location Banded
Years

West of
Continental Divide
No.
Location Banded Years

Vineland

3,481

8

Denver

5,984

10

Durango -

Ft. Morgan

2,810

7

Fort
Collins

5,052

8

Springfield
Area 11

354

3

Fort
Garland

2,787

5

Rocky Ford

171

4

Monte
Vista

2,680

Fort
Carson

41

2,435
Montrose2-1 1,632

4

Craig 61
Area -

871

4

6

Meeker I!

733

4

468

2

Unaweep
Canyon

358

2

Longmont

318

4

Salida

303

5

6

11

- D.O.W. and U.S.F.&amp; W.S. bandings only.

21

- Areas with less than 100 total bandings are not included.

31

- Includes bandings near Pritchett and Fort Lyons
~/Includes

3 birds banded at Arboles.

i/Includes 1 bird banded at Sanborn Park.
~/Includes bandings at Lay (522) and Hayden (101).
l/Includes

2 birds banded at Buford.

Age and Sex of Doves Banded
Once trapping and banding efforts were intensified, some attention was given
to attaining even age (AHY and HY) and sex (AHY only) ratios in the samples
banded. However, due to time and personnel constraints, banded samples were
frequently biased towards adults at some sites (trapped in June) and immatures
at others (trapped in late July and August). Age and sex composition of
banded samples combined for all years at each site and geographic area are
presented in Table 4.

�-99-

Table 4. Age and sex composition of banded samples by geographic
and location, 1964-1974.
1/
,
AHY

Geographic
Area and Location

area

HY

Males
No.
%

Females
No.
%

Vineland

804

50.4

792

49.6

1,596

Fort Morgan

335

43.0

444

57.0

37

67.3

18

Rocky Ford

7

87.5

Bonny Reservoir

8

88.9

Total
No.
%

%

Grand
Total 21

47.0 1,797

53.0

3,393

779

27.7 2,031

72.3

2,810

32.7

55

15.5

299

84.5

354

1

12.5

8

4.7

163

95.3

171

1

11.1

9

12.2

65

87.8

74

1,191

48.7 1,256

51.3

2,447

36.0 4,355

64.0

6,802

Denver

1,102

55.9

869

44.1

1,971

32.9 4,Ol3

67.1

5,984

Fort Collins

2,239

60.3 1,476

39.7

3,715

73.5 1,337

26.5

5,052

Fort Garland

671

63.3

389

36.7

1,060

38.0 1,726

62.0

2,786

Monte Vista

801

54.9

658

45.1

1,459

54.4 1,221

45.6

2,680

Fort Carson

182

56.9

138

43.1

320

69.4

141

30.6

461

Longmont

l38

60.0

92

40.0

230

72.3

88

27.7

318

Salida

152

58.7

107

41.3

259

87.2

38

12.8

297

Other 1/

37

74.0

13

26.0

50

90.9

5

9.1

55

5,322

58.7 3,742

41.3

9,064

51.4 8,569

48.6

17,633

Durango

1,118

56.3

867

43.7

1,985

81.6

449

18.4

2,434

Montrose

473

52.7

424

47.3

897

55.0

735

45.0

1,632

Craig Area

263

58.4

187

41.6

450

51.7

421

48.3

871

Meeker

334

59.3

229

40.7

563

76.8

170

23.2

733

Unaweep Canyon
41
Other -

151

66.2

77

33.8

228

63.7

l30

36.3

358

12

52.2

11

47.8

23

95.8

1

4.2

24

Subtotals

2,351

56.7 1,795

43.3

4,146

68.5 1,906

31.5

6,052

Totals

8,864

56.6 6,793

43.4 15,657

51.4 14,830

48.6

30,487

No.

Eastern Prairies

Springfield

Area

Subtotals
Eastern Foothills

Subtotals
West Sl°Ee

II

- D.O.W. and U.S.F. &amp; W.S. bandings only.
21
- Totals by area may not correspond to those in Table 3 as birds unclassified
as to sex (adults only) and age are not included.
l1Inc1udes birds banded at Evergreen-Aspen
Park (8), Estes Park (9) and Rye (1).
~/Inc1udes birds banded at Carbondale

Park (24), LaVeta (13), Woodland

(9) and Stoner (15).

�-100-

Differences between locations within geographic areas were occasionally
significant (P &lt; 0.05). It is doubtful that differences in sex composition
between locations within geographic areas were real. It is more probable
that observed differences in percentages of males and females trapped were
related to time of day of trapping effort and bird behavior patterns. Percentages of adults versus immatures in trap samples were most probably related to timing of the trapping effort in relation to the breeding and nesting
period. Higher percentages of immatures were banded at sites trapped in
August (Denver, Fort Garland) than at sites trapped in June (Fort Collins,
Durango). In a study in Missouri, Henry et al. (1976) demonstrated that
placement of traps could clearly bias age and sex composition of doves
captured while Lewis and Morrison (1973) presented data documenting biases
in trap results resulting from use of different types of traps and time of day.
All of these factors, except time of day and type of trap, were undoubtedly
involved in affecting the age and sex composition shown in Table 4. Trapping
in Colorado during the 1964-1974 interval was usually consistent with traps
being checked three times per day at approximately the same times. Type of
trap used did not change throughout the study

Recaptures
All previously banded mourning doves recaptured were recorded each year with
few exceptions. Known exceptions primarily involved returns of birds at
particular sites within a few days of original banding. Surprisingly few
banded doves were recaptured more than 90 days after initial capture (less
than 300 from 1964 through 1974) with only 79 individuals being recaptured
out of original degree block of banding since inception of banding in Colorado
in 1913. Origin of mourning doves recaptured in Colorado that had moved at
least one degree block is shown in Table 5.

Table 5.

Origin of mourning doves recaptured in Colorado, 19l3-1974.!1

Eastern Management Unit
State

No.

Alabama

1

Central Management Unit
State
Colorado
New Mexico
Texas

Western Management Unit

No.

State

No.

72
3
1

Arizona
Utah

1
1

llNo out of degree block of banding recaptures were reported prior to 1961.

Due to the limited recapture data available, detailed analyses outside of orlgln
could not be made. The longest intervals between banding and recapture were
six (one) and five years (two) with most being of a duration of 1 to 3 years.

�-101-

Hatching Chronology
During the 1964-1974 interval, wing molt data were recorded for 14,560
immature mourning doves newly banded and 4,792 hunter harvested immatures.
Hatching dates were calculated following procedures and data published by
Allen (1963) even though limited data from free-flying immature doves
banded and recaptured in Colorado suggest that Allen's molt schedules may
be 3 to 4 days slower than that actually occurring in the wild (Petersen
and Braun 1974). Hatching chronologies calculated by year for wild trapped
and harvested mourning doves in Colorado are presented in Tables 6 and 7.
Upon close inspection of data presented in Table 7, it is apparent that
doves produced prior to 12 June are poorly represented « 1.0 percent of the
total) in the harvest whereas doves produced in this early period comprise
about 54 percent of those trapped and banded (Table 6). Also, few doves
produced after 30 July are represented in trap samples as trapping normally
ceased by 15-20 August. However, doves produced after 30 July comprised
about 38 percent of the immature doves shot each year. These differences
are significant (P&lt;0.05) and suggest that real differences occur in availability of early and late produced immature doves to hunters in Colorado.
In an earlier study, Funk (1965) clearly documented that the peak of dove
migration through Colorado occurred prior to 1 September. Data from wings
of immature doves examined in the present study support Funk's earlier work
and it can reasonably be concluded that most early produced young have
migrated from Colorado prior to the hunting season.

Harvest Age Ratios
Wings from hunter harvested mourning doves were not routinely collected in
most years. Consequently, sample sizes ranged from few in 1968 to over 2,900
in 1973 when efforts to collect wings were made. Intensive efforts to
collect wings were made from 1972 through 1974. Data available are presented
in Table 8.
Data in Table 8 should be viewed with caution because of small sample sizes
in most years and uneven geographic distribution of the sample from within
Colorado. Most samples were from eastern Colorado, either primarily southeast (i.e. Springfield) in 1964-1971 or southcentral (i.e. Fort Carson) in
1972-1974. Neither of these areas may be representative of the total harvest
in the state. Assuming that larger sample sizes most nearly approximate
actual age composition of the harvest, it is interesting to note that age ratios
in the four ye ars (1967, 1972-1974) when over 1,000 wings were available each
year, ranged from 1.0:1 (two years) to 1.5:1 to 2.2:1. These ratios would
appear to be lower than those expected. In reviewing chronology of age ratios
within a hunting season, data collected at Fort Carson from 1972-1974 reveals
a low immature to adult ratio during the opening week of the hunting season
(usually from 0.5 to 0.8:1) with immatures predominating as the season progressed. Reasons for this phenomenon are unclear but may relate to the early
migration of immatures previously documented. If one assumes that the average
of 1.4:1 immatures to adult age ratio in the harvest (Table 8) is realistic
and 50 percent of the adult harvest is comprised of females, then 2.8 young
per female were produced. Thus it would appear that production is sufficient
to maintain the population.

�Table

6.

Estimated

hatching

dates

for

wild

trapped

immature

Prior to
May 1

May 1-7

May 8-14

May 1521

May 2228

1964

3(1.4)

1(.5)

7(3.3)

19(9.0)

1965

3(.4)

2 (.3)

3(.4)

1966

6(4.0)

mourning

doves,

Colorado,

1964-1974.

May 29June 4

June 511

June 1218

June 1925

33(15.7)

30 (14.3)

55(26.2)

59(28.1)

3(1. 4)

26(13.1)

92(13.1)

102(14.6)

150 (21. 4)

177(25.3)

92 (13.1)

12(7.9)

26(17.2)

47(31.1)

25(16.6)

16(10.6)

8(5.3)

11(7.3)

55(4.7)

113(9.7)

153(13.2)

158(13.6)

211(18.2)

156 (13.4)

57(4.9)

June 26July 2

July 39

July 1016

July 1723

July 2430

Total
No.

210
14(2.0)

18(2.6)

21(3.0)

700
151

1967

9 (.8)

20(1. 7)

1968

2 (.1)

12 (.8)

64(4.5)

93(6.5)

143 (10 .1)

177 (12. 5)

165 (11. 6)

213(15.0)

153(10.8)

170(12.0)

1969

13(1.2)

12(1.1)

69(6.6)

166 (15.9)

106(10.1)

127 (12 .1)

87(8.3)

126(12.0)

92(8.8)

91(8.7)

1970

3(.2)

5(.4)

14(1.1)

77(6.1)

124(9.9)

178(14.2)

86(6.9)

98(7.8)

137(10.9)

1971 '

3 (.1)

20 (.8)

57(2.3)

155(6.3)

337(13.7)

281(11.4)

313(12.7)

292 (11.6)

286 (ll.

125(5.4)

283(12.2)

340(14.6)

344(14.8)

288(12.4)

326(14.0)

51(4.4)

76(6.5)

66(4.6)

24 (1. 7)

1421

67(6.4)

66(6.3)

25(2.4)

1047

187(14.9)

191 (15.2)

93(7.4)

59(4.7)

308(12.5)

202(8.2)

193(7.8)

20 (.8)

2467

196(8.4)

139(6.0)

135(5.8)

78(3.4)

22 (1.0)

2323

40(3.4)

49(4.2)
139(9.8)

13(1.1)

1161

,...I
0
N

I

6)

1972

3(.1)

44(1.9)

1973

4 (. 2)

14(.7)

53(2.9)

126(6.9)

292 (16.0)

260(14.3)

220(12.1)

239 (13.1)

293(16.1)

237(13.0)

77(4.2)

1974

12 (.6)

31(1. 5)

101(5.0)

281(14.0)

344(17.1)

272(13.5)

229(11.4)

236(11.7)

205 (10. 2)

144(7.2)

139(6.9)

18(.9)

Totals

55(0.4)

167(1.1)

560(3.8)

1365(9.4)

2011(13.8)

1954(13.4)

1820(12.5)

1930 (13.3)

1525(10.5)

1330 (9 .1)

1017(7.0)

586(4.0)

1 (.1)

1253

1815
2012

226(1.6)

14(0.1)

14560

�Table

7.

Estimated

hatching

Prior to
June 12

dates

June 1218

for

hunter

harvested

June 1925

June
July

immature mourning

262

July 39

1964

doves,

Colorado,

1964-1974.

July 1016

July 1723

July 2430

July 31August 6

2(10.5)

5(26.3)

5(26.3)

1(5.3)

6 (31. 6)

August
7-13

1965

6(2.4)

17(6.7)

50(19.7)

7(2.8)

38(15.0)

38(15.0)

33(13.0)

29(11.4)

26(10.2)

9(3.5)

1966

6(2.2)

27(10.0)

31(11.5)

3 (1.1)

28(10.4)

40(14.8)

24(8.9)

37(13.7)

67(24.8)

7(2.6)

1967

1(.2)

2(.4)

11(2.0)

4(.7)

30(5.5)

52(9.5)

102(18.6)

123(22.4)

176 (32 .1)

39(7.1)

6(4.2)

15(10.6)

18(12.7)

25(17.6)

20(14.1)

24(16.9)

34(23.9)

9(7.4)

19(15.7)

18(14.9)

18(14.9)

15(12.4)

17(14.0)

23(19 .0)

1968
1969
1970

2(1. 7)
3 (1. 5)

15(7.8)

18(9.3)

7 (3. 6)

27(14.0)

34(17.6)

24(12.4)

26(13.5)

August
14-20

August
21-27

August 28Sept. 3

Sept. 410

Total
No.

19
254

1(.4)

270
4 (.7)

2(.4)

2 (. 4)

548
142
121

I

I-'

193

1(.5)

38(19.7)

286

1971

2 (. 7)

24(8.4)

18(6.3)

26(9.1)

48(16.8)

38(13.3)

31(10.8)

49(17.1)

36(12.6)

14(4.9)

1972

9 (1. 4)

4(.6)

102 (15 .6)

15(2.3)

48(7.3)

109(16.7)

79(12.1)

46(7.0)

111(17.0)

90(13.8)

20(3.1)

20(3.1)

1973

7 (. 5)

50(3.3)

90(6.0)

28(1.9)

131(8.7)

163(10.8)

102(6.8)

226(15.0)

206 (13. 7)

137(9.1)

118(7.8)

194 (12.9)

1~74

4 (.1)

28(3.5)

41(5.2)

23(3.0)

89(11.2)

70(8.9)

75(9.4)

95(11.9)

131(16.5)

71(8.9)

84(10.6)

Totals

40(0.8)

182 (3. 8)

395(8.2)

113(2.4)

475(9.9)

589U2.3)

510(10.6)

677(14.1)

849(17.7)

373(7.8)

225(4.7)

653
57(3.8)

1509

42(5.3)

44(5.5)

797

256(5.3)

106(2.2)

2(0.04)

4792

0

w
I

�-104-

Table 8.

Age ratios of hunter harvested mourning doves, Colorado, 1964-1974.

Year

AHY

HY

Total Wings

Immature:Adult

1964

62

19

81

0.3:1

1965

128

254

382

2.0: 1

1966

148

344

492

2.3:1

1967

569

582

1,151

1.0:1

1968

ND 1/

ND·U

ND 1/

ND )j

1969

51

121

172

2.4:1

1970

37

192

229

5.2:1

1971

24

286

310

11.9:1

1972 ]j

446

654

1,100

1.5 :1

1973

1,451

1,509

2,960

1.0:1

1974

368

797

1,165

2.2:1

Totals

3,284

4,758

8,042

Average

Ratio

1.4:1

1/
ND = No data.
2/
- Efforts to collect wings from hunter harvested

doves were intensified.

Hunter Pressure and Harvest
Harvest surveys were not included as part of the study reported here. Through
courtesy of the surveys group, Wildlife Management Section, they are included
here for completeness.
Estimated number of hunters and total harvest from
1964 through 1974 as derived from the annual survey of small game hunters are
presented in Table 9. During this interval, length of season was 60 days
(1 September through 30 October), daily bag and possession limits were either
12 and 24 (1964-1968) or 10 and 20 (1969-1974) with shooting hours of one half
hour before sunrise to sunset (1964-1969) or sunrise to sunset (1970-1974).

�-105-

Table 9.

Mourning dove harvest statistics, Colorado, 1964-1974.

II

Year

Estimated
Number of Hunters

Estimated
Total Harvest

1964

22,477

130,367

1965

19,545

137,401

1966

25,227

174,066

1967

24,593

140,180

1968

25,186

308,881

1969

25,119

323,773

1970

24,149

301,562

1971

22,033

298,767

1972

33,299

513,898 ]J

1973

31,688

484,660

1974

41,518

688,312 ~I

21

-liE·s t tmate d from annual survey of small game license holders.
.
-2/R eportlng
bias correction for multiples of the bag and multiples of 5
deleted.

llprocedural change in survey to go to multiple followup surveys instead
of original survey letter and one followup as in earlier years.

41
- Procedural change in survey due to stratification of hunters by license
type and splitting of waterfowl and small game surveys.

Due to several procedural changes in annual small game hunter surveys from
1964 to 1974, it is difficult to accurately discuss changes in hunter numbers
and annual harvests. Obviously hunter numbers and annual harvests have increased during the 11 year period, hunters by possibly 85 percent and harvest
by possibly 428 percent. While these increases may be feasible, they are not
overly plausible. Independent surveys of all hunters of certain minor game
species suggest that the annual survey of small game hunters may overestimate
hunter numbers by as much as 30 percent and harvest by 100 percent. It is
possible and quite probable that numbers of mourning dove hunters are slightly
overestimated (about 10 percent) with actual numbers of doves harvested being
moderately overestimated (as much as 50 percent). Thus, best estimates of
numbers of dove hunters and annual harvest in 1974 are about 35,000 + 3,000
and 500,000 + 50,000, respectively.
-

�-106-

Mourning Dove Recoveries
Mourning doves banded in Colorado from 1913 through 1974 totaled 34,225
(Table 1). Only one major period of intensive banding occurred as 31,523
primarily free flying doves were banded from 1964-1974. Some intensive
banding of primarily nestlings was done between 1951-1952 and 1956-1961
but numbers exceeded 400 only in 1960 when 557 were banded. During the
1951-1961 period, 2,404 of the 2,702 doves banded prior to 1965 were
captured and marked. Through 4 March 1976, 683 recoveries of doves banded
in Colorado had been reported. Of this number, only 53 were from the 2,702
bandings prior to 1964 (2.0 percent of the banded population). All but 8
were hunting season recoveries. Interestingly enough, recovery rates did not
change between the 1913-1963 and 1964-1975 periods as 630 recoveries were reported from 31,523 bandings (2.0 percent of the banded sample). Of the 630
total, 59 were no~ hunting season recoveries. This is a substantial decrease
from the non hunting mortalities reported in the 1913-1963 period (9.4 percent of all recoveries in 1964-1975 versus 15.1 percent in 1913-1963). This
decrease could be the result of larger samples, less nestling banding or
lower reporting rates.

Origin of Banded Mourning Doves Harvested in Colorado
In the 1913-1963 interval, 9 mourning doves banded outside of Colorado and 26
doves banded in Colorado were reported recovered within the state. Thus of
35 banded doves reported, 25.7 percent originated elsewhere. Point of origin
(banding location) for these 9 birds was two each from New Mexico and North
Dakota and one each from Arizona, Kansas, Nebraska, South Dakota and Texas.
Only one bird was banded outside of the Central Management Unit (Arizona).
Recoveries of birds banded elsewhere in the 1913-1963 period were too small
to make firm statements about origin of doves banded elsewhere and recovered
in Colorado.
During the 1964-1975 interval, 87 doves banded outside of Colorado and 351
doves banded in Colorado were reported recovered in the state. Thus, of 438
banded doves reported, 19.9 percent originated elsewhere. This would suggest
that about 20 percent of the doves harvested each year in Colorado originated
in other areas. Points of origin for the 87 out-of-state of banding recoveries are presented in Table 10.
Only 12 recoveries from bandings out of the Central Management Unit were
reported in the 1964-1975 period. Two originated in the Eastern Management
Unit, while 10 originated in the Western Management Unit. Patterns of recoveries from bandings in these two management units could not be ascertained.
Over 86 percent (86.2) of the "foreign" recoveries in Colorado were from other
states in the Central Management Unit. Over one-half (52.0 percent) originated in areas directly north of Colorado (Montana and Wyoming), with lesser
numbers (14.7 percent) originating in areas directly south of Colorado (New
Mexico and Texas).

�-107-

•
Table 10. Origin of banded mourning doves recovered in Colorado from outof-state bandings, 1964-1975.

Eastern Management Unit
State

Central Management Unit
Number

State

Number

Louisiana

1

Iowa

3

Wisconsin

1

Kansas

1

Total

2

Minnesota

1

Montana

32

Nebraska

7

Western Management Unit
State

Number

New Mexico

6

Arizona

3

North Dakota

2

California

1

Oklahoma

1

Nevada

2

South Dakota

10

Utah

1

Texas

5

Washington

3

Wyoming

7

Total

10

Total

75

Distribution

of Harvest

Through 4 March 1976, 683 recoveries of doves banded in Colorado had been
reported, of which 53 were from bandings in the 1913-1963 period and 630 were
from bandings from 1 June 1964 through 30 August 1974. Considering only the
53 recoveries from the 19l3~1963 bandings, 27 (50.9 percent) were from Colorado, 18 (34.0 percent) were from Mexico, 4 (7.5 percent) were from Texas,
2 (3.8 percent) were from Arizona, while 1 each were from New Mexico and
Louisiana. Only one recovery was reported from the Eastern Management Unit,
while two were reported from the Western Management Unit. Distribution of
recoveries from doves banded from 1964 through 1974 is shown in Table 11.
Recoveries were clumped in the Central Management Unit (449 of 630, 71.3 percent) and Mexico (141 or 22.4 percent) with lesser numbers from the Western
Management Unit (33 or 5.2 percent) and Eastern Management Unit (2 or 0.3 percent). Central American countries (4) and Canada (1) contributed little to
the overall reported harvest. Colorado was the leading harvest area (55.7
percent of all recoveries) followed by Mexico (22.4 percent), Texas (7.6 percent)
and New Mexico (6.8 percent). No other state or country provided more than 3
percent of the total reported recoveries.

�-108-

Table 11. Distribution
rado, 1964-1974. 1/

of mourning dove recoveries from bandings in Colo-

Canada
Quebec

, 1

Central America
E1 Salvador

2

Guatama1a

1

Nicaragua

1

Mexico

141

Total

146

1/ Through 4 March 1976.

�-109-

Considering only those recoveries in Mexico for which report to bander cards
were available (primarily after 1968), recovery locations of 84 doves were
approximated. This sample was reported from 11 states, with a majority (49)
being reported from Jalisco (28) and Michoacan (21) in west central Mexico.
More than five recoveries were reported in only two other States, Guanajuato
(10) and Sinaloa (9). Thus, mourning doves banded in Colorado are not exceptionally vulnerable to hunters in Mexico until they reach presumed wintering
areas. These data are similar to those reported from an earlier analysis of
all mourning dove recoveries in Mexico (Blankenship and Reeves 1970).
Particular attention was given to recoveries reSUlting from bandings west of
the Continental Divide since 1960. Only 51 recoveries from bandings in this
geographic area have been reported to date, with 16 (31.4 percent) from Mexico,
10 (19.6 percent) from New Mexico, 9 (17.6 percent) from Arizona, 6 (11.8
percent) from both California and Colorado, 3 (5.9 percent) from Texas, and 1
(2.0 percent) from Utah. This pattern is markedly different from that of all
recoveries of doves banded in Colorado in that recoveries in Colorado and Texas
are minimal.
It would appear that doves banded west of the Continental Divide
move southwest from Colorado and have some affinity with Western Management
Unit states (16 versus 19 recoveries in New Mexico, Colorado and Texas). When
exact locations of recoveries are examined, most of the 19 Central Management
Unit recoveries are from west of the Continental Divide in the three states
involved.
Mortality and Survival Rates
It is not the purpose of this report to present detailed analyses of mortality
and survival rates by age and sex class and geographic area of banding. These
analyses will be presented in a final cooperative report of all bandings in
the Central Management Unit in the 1967-1974 period. This report will be
completed in 1977, which allows most recoveries from doves banded in 1974 to
be included. Presented herein are analyses of all pooled recoveries from
1964 to 1974 with no separation of sex or age. Methodology is that of the
relative recovery rate method described by Geis (1972). Actual data are
given,in Tables 12 and 13.
Calculated annual survival rates varied from .0033 to 1.728 and averaged .523
with an average annual mortality rate of .477. The average survival rate
calculated for all doves banded in Colorado is substantially higher than
those calculated for various age and sex classes of doves banded in the Eastern
Management Unit (Hayne 1975) but similar to those calculated for doves banded
in South Dakota (Rice and Lovrien 1974).

�-110-

Table 12. Banding and recovery data from all mourning
doves banded in
Colorado, 1964-1975.

Year

No.
Banded

1

Hunting Season of Recoverl
2
3
4
5
6

1964

295

5

2

1

1

1965

808

15

12

5

1

1966

220

2

1

1967

2,232

45

19

7

4

2

1968

2,501

35

30

9

6

1

4

1969

3,390

57

23

9

6

3

1

1970

3,159

56

9

10

7

1

83

1971

5,356

27

27

16

3

5

78

1972

4,633

36

21

7

4

1973

4,409

44

11

5

1974

4,520

18

15

Totals

31,523

340

170

7

Total

9
0

1

34
3
77
85
1

100

68
60
33

69

32

J.2

6

1

630

�-111-

Table 13. Relative recovery rate estimates, all mourning doves banded in
Colorado, 1964-1974.

NU!1J.ber
of
Recoveries
1-n
2-n

Year

Number
Banded

1964

295

Can't Use

4

1965

808

34

19

.0421

.0235

1

.0136

.0045

Recovery Rates
1-n
2-n

.0136

1966

220

1967

2,232

77

1968

2,501

85

50

.0340

1969

3,390

100

43

.0295

3

32

.0345

.0143
.0200
.0127

Survival
Rate

Mortality
Rate

0.323

0.677

1.728
0.130

0.987

0.421

0.579

0.678

0.322

0.483

0.517

0.582

0.418

0.646

0.354

0.507

0.493

0.493

0.507

1970

3,159

1971

5,356

78

51

1972

4,633

68

32

.0147

.0069

16

.0136

.0036

.0073

.0033

.2302

.1204

5.991

4.854

Average survival rate:

.523

.599

.401

Average mortality

.477

1973

4,409

1974

4,520

Total

31,523

83

60
33

27

15

.0263
.0146

rate

.0085
.0095

LITERATURE CITED
Allen, J. M. 1963. Primary feather molt rate of wild immature doves in
Indiana. Ind. Dept. Conserv., Game Res. Sect. Circ. No.4.
4 p.
Bailey, A. M., and R. J. Niedrach.
Nat. Rist., Vol. 1. 895 p.

1965.

Birds of Colorado.

Denver Museum

Blakenship, L. R., and R. M. Reeves. 1970. Mourning dove recoveries from
Mexico. U. S. Fish and Wi1d1. Serv., Spec. Sci. Rept. Wi1d1. No. 135.
25 p.
Braun, C. E. 1976. Methods for locating, trapping and banding band-tailed
pigeons in Colorado. Colo. Div. Wi1d1. Spec. Rept. No. 39. In Press.

�-112LITERATURE CITED (Continued)
Funk, H. D. 1965. Mourning dove migration in Colorado.
Fish and Parks, Game Info. Leaflet No. 26. 2 p.

Colo. Div. Game,

Geis, A. D. 1972. Use of banding data in migratory game bird research and
management.
U. S. Fish and Wildl. Serv., Spec. Sci. Rept. Wildl. No.
154. 47 p ,
Hayne, D. W. 1975. Experimental increase of mourning dove bag limit in
Eastern Management Unit, 1965-1972.
Southeastern Assn. Game and Fish
Comms., Bull. No.2.
56 p.
Henry, D. L., T. S. Baskett, K. C. Sadler, and W. R. Goforth. 1976. Age and
sex selectively of trapping procedures for mourning doves. J. Wildl.
Manage. 40(1):122-125.
Keeler, J. (Chairman). 1976. The mourning dove. In Sanderson, G. C. (Ed.).
Management of migratory webless game birds in North America.
Intnl.
Assn. Game and Fish Comms. and U. S. Fish and Wildl. Servo In Press.
Kiel, W. H., Jr. 1959. Mourning dove management units, a progress report.
U. S. Fish and Wildl. Servo Spec. Sci. Rept. Wildl. No. 42. 24 p.
Lewis, J. C., a~d J. A. Morrison.
1973. Efficiency of traps and baits for
capturing mourning doves. Wildl. Soc. Bull. 1(3):131-138.
Peters, H. S. 1961. The past status and management of the mourning dove.
Trans. No. Am. Wildl. and Nat. Resources Conf. 26:371-374.
Petersen, B. E., and C. E. Braun. 1974. Observations on the primary feather
molt of imnature mourning doves in Colorado. J. Colo.-Wyo. Acad. Sci.
7(5):58.
Reeves, H. M., A. D. Geis, and F. C. Kniffin. 1968. Mourning dove capture
and banding. U. S. Fish and Wildl. Servo Spec. Sci. Rept. Wildl. No. 117.
63 p.
Rice, L. A., and H. Lovrien. 1974. Analysis of mourning dove banding in
South Dakota. J. Wildl. Manage. 38(4):743-750.
Ruos, J. L. 1968. Mourning dove banding needs, 1969-1971.
Wildl. Serv., Unpubl. Rept. 13 p.

U. S. Fish and

Wight, H. M. 1963. The role of pre- and post-season banding in mourning
dove research. Bur. of Sport Fisheries and Wildl., Migratory Bird Pop.
Sta. Admin. Rept. No. 19. Feb. 28. 8 pp. + 6 Tables.

Prepared by
Clait E. Braun
Wildlife Researcher

�October 1976

-113JOB FINAL REPORT

State of

Colorado

---------------------

Project No.

W-88-R-21

i-lorkPlan No.

4

Job Title

Migratory Bird Investigations
4

Job No •.

--------------------.----------------

Band-tailed Pigeon Investigations

Period Covered:

--~--------------------------

April 1, 1969 through March 31, 1975

Personnel: S. Amend, K. Baer and J. Randall, U.S. Fish and Wildlife Service;
J. Cruse, W. Paintner, C. E. Williamson and other personnel of the U.S. Forest
Service; R. Stabler and N. Kitzmiller, Colorado College; O. W. Olsen, Colorado State Univ.; W. J. Arthur, D. Benson, C. E. Braun, W. Carpenter, A. Cowan,
H. D. Funk, J. E. Kautz, M. V. Kautz, W. Ladd, Jr., S. Mathews, C. McCord, R.
Oakleaf, B. Petersen, M. Robinson, L. Seger, B. Smith, N. Smith, M. Stromberg,
M. Watkins, J. A. White, Wildlife Conservation Officers, Area Supervisors,
Conservation Aides, ane other personnel of the Colorado Division of Wildlife.

ABSTRACT
The objectives of this study have been fully achieved. Reports covering
the various objectives have been published, prepared and submitted, and in
some instances are in progress. Those in progress will be published under
Work Plan 6, Job 1 and are listed in the project documents for W-88-R-22.
Those published and submitted are listed below:
Braun, C. E. 1972. Movements and hunting mortality of Colorado band-tailed
pigeons. Trans. No. Am. Wildl. and Nat. Resour. Conf. 37:326-334.
1973. Distribution and habitats of band-tailed pigeons in Colorado.
Proc. West. Assoc. State Game and Fish Comms. 53:336-344.
1973. Studies of band-tailed pigeons in wintering areas of Mexico,
December 1972 tv April 1973. Mimeo Rept., Contract No. 14-16-0008-626.
Colorado Div. Wildl. 11 pp.
1974. Investigations of band-tailed
Proc. Wilson Ornithol. Soc. 55:19.

pigeons in Mexico.

Abstract.

1976. Methods for locating, trapping and banding band-tailed
in Colorado. Colo. Div. Wildl. Special Rept. No. 39:In Press.

pigeons

_____ , W. J. Adrian, and R. E. Keiss. 1977. Mercury residues in Colorado
band-tailed pigeons. Submitted to J. Wildl. Manage.
_____ , D. E. Brown, J. C. Pederson, and T. P. Zapatka. 1975. Results of
the Four Corners Cooperative Band-tailed Pigeon Investigation.
U. S.
Fish and Wildl. Serv., Resource Publ. 126. 20 pp.

�-114ABSTRACT (Cont.)

Gutierrez, R. J. 1973. Reproductive biology of the band-tailed pigeon
(Columba fasciata). M. S. Thesis, Univ. New Mexico, Albuquerque.
28 pp.
, C. E. Braun, and T. P. Zapatka. Reproductive biology of the band-----tailed pigeon in Colorado and New Mexico. Auk 92(4):665-677.
Olson, o. W., and C. E. Braun. 1977. Three new species of Splendidofilariinae (Filarioidea:Nematoda) from the band-tailed pigeon (Columba
fasciata fasciata [Say]) in Colorado. Submitted to Proc. Helminth
Soc. Washington.
Stabler, R. M., and C. E. Braun. 1975. Effect of virulent Trichomonas
gallinae on the band-tailed pigeon. J. Wildl. Diseases 11:482-483.
, N. J. Kitzmiller, and C. E. Braun. Blood parasites from band-tailed
pigeons. Submitted to J. Wildl. Manage.
White, J. A. 1973. Molt of Colorado band-tailed pigeons.
Colorado State Univ., Ft. Collins. 34 pp.

M. S. Thesis,

_____ , and C. E. Braun. 1972. Plumage investigations of Colorado bandtailed pigeons. J. Colo.-Wyo. Acad. Sci. 7(3):70.

Prepared by __

-,~~~~:-=-L,::-,:-+~~:;=.::.;,-,-Clait E. Brauh
Wildlife Researcher

_

�October 1976

-115JOB FINAL REPORT

State of

COLORADO
------~~~~-------------

Project No.

~W_-~8~8-~R~-~2~1~

Work Plan No.

_

Migratory
Job No.

5

Bird Investigations
~l~

Job Title

Investigations

Period Covered:

September I, 1973 through March 31, 1976

Personnel:

_

of Snipe in Colorado

C. Bryant and C. Donner, U. S. Fish and Wildlife Service; R.
Ryder, D. Hein, and R. Walter, Colorado State University, C.
Braun, H. Funk, and B. R. Johnson, Colorado Division of Wildlife.

ABSTRACT
Abundance, times of occurrence, and harvest of common snipe (Capella gallinago
delicata) in Colorado were investigated from September 1973 through June 1975.
Areas studied were near Fort Collins, and in North Park in north central
Colorado; the Yampa Valley in northwestern Colorado; and the San Luis Valley
in south central Colorado.
Estimated densities of breeding snipe based on
censuses conducted during May 1974 and 1975 were, by region: 1.5 snipe/ha
near Fort Collins; 0.6 snipe/ha in North Park; 0.6 snipe/ha in the Yampa
Valley; and 0.5 snipe/ha in the San Luis Valley. Mean density for all regions
was 0.6 snipe/ha on a total area of 245~'1 ha. On individual study areas
densities of snipe ranged from 0.3 to 1.9 snipe/ha. Areas with shallow,
stable water levels, sparse, short vegetation, and soft organic soils had the
highest densities of snipe. Twenty-eight nests were located having a mean
clutch size of 3.9 eggs. Estimated onset of incubation ranged from 2 May
through 4 July, with most occurring in May. Spring migration extended from
late March through early May. Maximum densities were recorded in all regions
during 10-23 April. Fall migration was underway by early September and was
completed by mid-October, with maximum snipe densities occurring the third
week in September.
High numbers of snipe, noted in early August, may have
been early migrants or juveniles concentrating on favorable feeding areas.
Snipe wintering near Fort Collins preferred sparsely vegetated shorelines,
with soft, water-saturated soils. Snipe utilized such areas from October
through April in densities up to 16.5 snipe/km.
Results of the 1972 and
1973 survey of small game license holders indicated that successful respondents (n = 22) harvested a mean of 3.7 snipe per hunter and spent 4.0 days
afield. Results of questionnaires to snipe hunters in 1973 and 1974 indicated
that successful respondents (n = 8) harvested a mean of 10.8 snipe per hunter
during 4.9 days af~eld. The lrnN number of responses indicate the harvest of
snipe in Colorado is small.

�116

RECOMMENDATIONS

1.

On areas managed for snipe, shallow, discontinuous water
coverage should be maintained through water manipulation
practices.
Depths should not .exceed approximately 50 to
60 mm and short vegetative cover should be encouraged.

2.

Vegetation on areas managed for snipe should be manipulated
through grazing by livestock, burning and/or mowing to
maintain low cover (less than 200 mm) suitable for snipe.

3.

Use of suitable habitats by snipe may be increased through
application of animal waste products that increase number
and availability of larval insects.

4.

Further research on the relationship of snipe numbers and
use to soils of varying productivity may be beneficial in
future management of habitats specifically for snipe.

�-117TABLE OF CONTENTS

Introduction
Review of Literature
Taxonomy
Distribution
Ecology
Harvest
Methods .••
Selection of Study Areas
Census Techniques • .
Physical Dimensions • •
Soil Characteristics
Vegetation Descriptions
Water Analysis
• • • •
Harves t Survey s • • • . • • • • •

•

Location and Description of Study Areas
Location
Climate ..
Topography
Soils . • • •
Vegetation
• • • •
Water Characteristics
Results and Discussion
Estimates of Breeding Densities •
Nesting Activities
Spring Migration
Fall Migration
Estimates of Winter Densities •
Harvest Assessment
Literature Cited
Appendix

LIST OF TABLES
Table
1

2

Descriptive and quantitative data for major soil types
on snipe breeding areas 3-14 in Colorado • • • • .
Results of water analyses on snipe study areas 3-14
in Colorado, June 1975
•••••••••••••••.•.•

••.

•

�-118LIST OF TABLES (Cont.)
Table

Page

3

Estimated densities of snipe breeding on study areas
3-14 in Colorado, 1974 and 1975
. . • .

4

Snipe nesting data in Colorado, 1974 and 1975 .

5

Maximum densities of snipe with date of occurrence
for areas 3-14 during spring (1975) and fall (1974)
migration in Colorado • . • • . . . • . . • . .

6

Results of the 1972 and 1973 small game surveys and
the 1973 and 1974 snipe hunter questionnaires for
Colorado

...........................
LIST OF FIGURES

Figure
1

Location of snipe study areas in Colorado • . • •

2

Densities of snipe on areas 3, 4, and 5 near Fort
Collins, Colorado (May 1974-June 1975) • . • • •

3

Densities of snipe on areas 6, 7, and 8, North Park,
Colorado (May 1974-June 1975) . •
• •....

4

Densities of snipe on areas 9, 10, and 11, Yampa
Valley, Colorado (May 1974-June 1975) •

5

Densities of snipe on areas 12, 13, and 14, San Luis
Valley, Colorado (May 1974-June 1975) •

6

Densities of snipe on area 1 (January 1974-July 1975)
and area 2 (January 1974-May 1975) near Fort Collins
Colorado
. • • • . • • • • . . . . • • • • . • • • .

�-119-

ACKNOWLEDGEMENTS

The financial support of this investigation was provided by the Accelerated
Research Program of the U. S. Fish and Wildlife Service funded through the
Colorado Division of Wildlife and through Colorado Federal Aid in Wildlife
Restoration Project W-88-R. I gratefully acknowledge the substantial assistance of these agencies.
Dr. Ronald A. Ryder, Professor, Department of Fishery and Wildlife Biology,
Colorado State University, was paramount in the initiation of this project
and served as my major advisor. For his advice, counseling, and exemplary
professionalism, I am most grateful. I also express my thanks to the
following individuals who served on my graduate committee. Dr. Clait Braun,
Wildlife Researcher, Colorado Division of Wildlife, served both as research
advisor and coordinator. His astute observations and pointed recommendations
helped eliminate much ambiguity from this investigation.
Dr. Dale Hein,
Associate Professor, Department of Fishery and Wildlife Biology, Colorado
State University, offered valuable suggestions and insights which contributed
greatly to the project. Richard G. Walter, Assistant Professor, Department
of Botany and Plant Pathology, Colorado State University, provided considerable assistance in the botanical portion of this investigation.
My thanks are also extended to Richard M. Hopper and Michael R. Szymczak,
Wildlife Researchers, Colorado Division of Wildlife, for their sound
recommendations and assistance in locating suitable study sites. Likewise,
I wish to thank Charles Bryant and Carroll Donner, refuge managers of the
Monte Vista and Arapahoe National Wildlife Refuges, respectively, and their
staff for their hel~ful consultation and permission to conduct investigations
on the refuges.
To the Wildlife Conservation Officers I contacted, I am most grateful for
their assistance, observations, and recommendations.
The many landowners,
who without exception, allowed me access to their property deserve my special
thanks. Without their generosity, this investigation would not have been
possible. I am also g:rateful for the help and information provided by the
Soil Conservation Service and Agricultural Stabilization and Conservation
Service.
These acknowledgements would be incomplete i.f I did not thank my wife,
Peggy. I will always value her understanding and perseverance throughout
this investigation.

��-121-

INVESTIGATIONS

OF COMMON SNIPE IN COLORADO

Bruce R. Johnson

The 12 species of the genus Capella have a cosmopolitan distribution
(Blair et al. 1968), but the common snipe (Capella gallirtago delicata)
is the only representative indigenous to North America and occurs
seasonally over most of the continent (Bent 1927). Unlike most members
of Family Scolopacidae, common snipe frequent bogs, wet meadows and
pastures, and marsh and stream habitats.
Furthermore, snipe are generally
solitary and crepuscular.
Comprehensive life histories of common snipe include Bent (1927) and Tuck
(1972); less inclusive studies include Erickson (1945, 1953) and Stout
(1967). Snipe in Colorado were mentioned by Cook (1897, 1898, 1900,
1909), Sclater (1912), Aiken and Warren (19l4~ Niedrach and Rockwell
(1939), Packard (1945), Boeker (1954), Beidleman (1955), Bailey and
Niedrach (1965), and Martin et al. (1974).
This 2-year investigation
mary objectives to:

of common snipe was initiated in 1973 with pri-

1.

Estimate densities of breeding and migrating common snipe on
representative habitat types throughout Colorado.

2.

Document

3.

Evaluate the importance of open water areas to common snipe
wintering in Colorado.

4.

Ascertain hunting pressure, harvest size, and hunter success
through hunter questionnaires.

spring and fall migration

periods.

REVIEW OF LITERATURE

Taxonomy
Common snipe belong to the genus Capella (Family Scolopacidae, Order
Charadriiformes, Class Aves) which is divided into 12 species. The
species f. gallinago is further separated into eight subspecies of which
f. ~. delicata is the only subspecies indigenous to North America (Tuck
1972). Each species and subspecies were described by Tuck (1972).

Distribution
Common snipe (hereafter referred to as snipe) normally breed from northwestern Alaska, northern Yukon, and east across Canada to central Labrador

�-122-

and Newfoundland south through the northern United States from West
Virginia to California and eastern Arizona.
Their winter range extends
from Alaska, western Canada, and the western and southern United States
south to Central and northern South America (Bailey and Niedrach 1965).
More detailed winter and breeding ranges are given by Bent (1927),
American Ornithologists' Union (1957), Stout (1967), and Tuck (1972).
In Colorado, snipe are common summer residents from the upper Sonoran
through Canadian zone (Niedrach and Rockwell 1939). Scattered populations spend winter months from low elevations (Bailey and Niedrach 1965)
to, at least, 2900 m (Anon. 1886). Snipe are most numerous throughout
the state during spring and fall migrations.

Ecology
The principal spring migration period for snipe is from March through midApril, with males preceding females by approximately 10 days (Tuck 1972).
Arrival dates for snipe in Colorado include 10 March for Denver, 19 March
for Boulder, 26 March for Sweetwater Lake (Bent 1927), 17 April for Rocky
Mountain National Park (Packard 1945), and mid-April for the Yampa Valley
(Boeker 1954). Numbers of snipe arriving in Colorado are relatively large
by late April (Niedrach and Rockwell 1939).
Breeding is restricted largely to areas of organic soils, primarily peatlands, mostly within northern forest regions. On the southern limits of
the breeding range, snipe occupy areas along ponds, meandering streams and
rivers, and similar marshy localities having organic soil and sparse
vegetation (Tuck 1972). Stewart and Kantrud (1968) described breeding
habitat for snipe in North Dakota as calcareous bogs or fens that were
frequently grazed, with soils normally saturated by groundwater, and
often with hummocks or quagmires of floating surface mats of emergent
vegetation.
Characteristic plants were aquatic mosses, scattered shrubs,
grasses, sedges, and forbs.
Nesting follows mating activities and selection of nest sites and has
been described by Bent (1927), Tuck (1955, 1972), and Stout (1967).
Clutch size is normally 4 eggs, occaSionally only 3 (Wolfe 1931) and
rarely 5 eggs are laid (Bent 1927). Only females incubate the clutch
(Tuck 1972). Length of incubation varies from 18 to 21 days (Bent 1927,
Erickson 1953) and is usually 19 days (Stout 1967). Nest records for
Colorado are limited, ranging from 1 May to 1 July with most nests located
in May and early June (Niedrach and Rockwell 1939, Bailey and Niedrach
1965). In.Utah, nesting dates range from 29 April to 24 July (Johnson
1899, Bent 1927) while, in California, a late nest was located on 1
September (Bryant 1915).
Young snipe are precocial (Bent 1927) and double their weight within 1
week after hatching (Tuck 1972). Flight is attained between 15 and 18
days of age (Stout 1967) and occasionally earlier (Tuck 1972). By 3 weeks
of age, young have fledged and are as heavy (90 g) as their molting parents
(Tuck 1972).

�-123-

Fall migration in southern Canada and the northern half of the conterminous United States occurs primarily from September to early October
(Stout 1967) during which time numbers of snipe in Colorado are probably
maximal (Bailey and Niedrach 1965). Late dates of departure for snipe
in Colorado are 3 November for Greeley and 24 December for Boulder (Bent
1927) .
Principal winter habitats utilized include coastal marshes, rice fields,
and wet cattle pastures which contain less decomposed organic soil than
characteristic breeding habitats (Tuck 1972).
In regions with continental climates such as Colorado, scattered populations of snipe winter
wherever marshes and borders of sloughs remain unfrozen (Bailey and
Niedrach 1965).
Food habits of snipe have been investigated by Whitehead (1965), White
and Harris (1966), Van Owens (1967), and Booth (1968) and have been
summarized by Tuck (1972).
Erickson (1941) examined 76 snipe and found 80 percent infected with
parasites.
Other accounts of ectoparasites and endoparasites of snipe
include Schmidt (1962), Whitehead (1965), Booth (1968), Tuck (1972), and
Naney (1973).

Harvest
The harvest of snipe in the United States in recent years has been estimated at 500,000 to 750,000 (Arnold 1975) and in Canada, at 80,000 (Tuck
1972). Estimates by Clark (1971) indicated that snipe harvested by waterfowl hunters increased from 263,900 in 1965-66 to 524,600 in 1970-71.
Recovery rates of banded snipe of 2.4 percent in the United States
(Arnold 1975) and 1.1 percent in Canada (Tuck 1972) indicate a relatively
small harvest.
In Colorado, harvest of snipe appears to be incidental to hunting more
popular game species. Consequently, numbers of snipe hunters and snipe
harvest are small. Based on annual surveys of small game license holders
in Colorado from 1955 through 1971, the Colorado Division of Wildlife
(1973 unpublished report) estimated 455 hunters harvested 1,177 snipe
annually during that period.

METHODS

Selection of Study Areas
Potential study areas for breeding surveys were examined in Fort Collins,
North Park, the Yampa Valley, and the San Luis Valley from September
through November 1973 and mid-March through early May 1974. Criteria
for selecting breeding areas were: (1) presence of snipe performing
mating and territorial displays; (2) water depths and permanence; (3)
similarity of the area to general habitat types in each region; (4)
diverseness of the area compared to other selected areas; and (5) sufficient

�-124-

area to provide adequate numbers of snipe.
areas were selected, 3 in each region.

A total of 12 breeding

Wintering areas were investigated near Fort Collins during December
1973 and January 1974.
Two contrasting sites were selected in midJanuary 1974 on which densities of wintering snipe were estimated.

Census Techniques
A systematic strip census was employed to estimate numbers of snipe
occurring on each area. This technique provided reliable estimates
regardless of time of year and proved more dependable than either a
winnowing index (Burleigh 1952, Robbins 1954, Solman 1954, Tuck 1972)
or the territorial response method (Tuck 1972).
Strip censuses consisted of walking linear transects at 20 to 25 m
intervals. Transects corresponded to cardinal directions and paralleled
the width rather than length of the areas whenever possible. Field maps
incorporating cenSU3 routes and their relation to conspicuous landmarks
were used to achieve comparative censuses. Approximate locations of
all snipe flushed, alighting, or displaying within the area were recorded.
Biweekly censuses were conducted in North Park, Yampa Valley, and San
Luis Valley with two censuses per area per trip. Single, weekly censuses
were conducted on areas near Fort Collins.
Nests encountered during censuses were inconspicuously marked for subsequent relocation to record nesting progress. Dates of the onset of
incubation were estimated using a water flotation technique (Westerskov
1950) based on a 19-day incubation period. Widths and lengths of eggs
were measured with a vernier caliper.
Censuses on winter study areas paralleled the streams from 1 to 5 m
from the shoreline. Censuses were conducted biweekly except for monthly
censuses in July and August.

Physical Dimensions
Physical dimensions of each area were obtained directly from U. S.
Geological Survey 7.5-minute series topographic maps (scale 1:24000).
The size of each breeding area was calculated using a Dietzgen planimeter.
Lenghts of winter study areas were calculated with a Keuffel and Esser
cyclometer.
Elevations, latitudes, and longitudes were determined from
topographic maps.

Soil Characteristics
Soil descriptions were limited to breeding areas 3 through 14 and were
obtained from the U.S. Soil Conservation Service. Information obtained
included location and range of soil types, descriptions of soil profiles,
and physical and chemical properties including texture, slope, water table
depth, permeability, available water capacity, and soil reaction. Texture

�-125-

is based on relative amounts of sand, silt, and clay in a soil. Permeability is the rate of downward movement of water in major soil layers
when saturated but allowed to drain freely. Available water capacity,
measured as the amount of water in a specific volume of soil, is related to particle size and arrangement and size of soil pores. Fine
textured soils have high water retention because of small pores. Soil
reaction is the intensity of soil acidity or alkalinity expressed in pH
units (Soil Conservation Service 1970).
Measurements of ground compaction were made on each area with a Soiltest
penetrometer.
The unconfined strength of the soil was measured by
forcing a 16 mm diameter shaft into the soil to a prescribed depth of 2
15 mm and reading the resultant pressure on a scale from 0 to 4.5 kg/cm

Vegetation Descriptions
Dominant plant species based on frequency of occurrence were described
for breeding areas 3 through 14. Initially, each area was divided into
major communities from aerial photographs and checked through on-site
observations.
Percent coverage for each species was then estimated by
an ocular survey of each major community. Mean height and range for
each dominant species were determined from measurements of representative
plants. Specimens vf dominant species were collected for subsequent
identification.
Mounted specimens of these species were filed in the
Colorado State University herbarium as the voucher collection for this
investigation.

Water Analysis
Free acidity, phenolphthalein alkalinity, total alkalinity, hardness,
and pH were measured on breeding areas 3 through 14 using a Hach Model
Al-36-B Test Kit. Free acidity measures the concentration of strong
mineral acids such as hydrochloric and sulfuric acids in water.
Alkalinity, defined as the capability to neutralize acids, is expressed
as phenolphthalein alkalinity which registers total hydroxide and one
half the carbonate or total alkalinity which includes all carbonate,
bicarbonate, and hydroxide alkalinity.
Total hardness primarily measures
concentrations of calcium and magnesium present. The pH of water is an
index of the hydrogen ion activity ranging from 0 (very acidic) to 14
(very basic) with pure water as 7 (neutral). The pH value represents the
instantaneous hydrogen ion activity rather than the buffering capacity
or total reserve as in acidity or alkalinity tests (Hach Chemical Company
1973).
Three to four water samples were collected from each area. Sampling sites
varied from flowing water if present to stagnant water. All analyses
of water samples were completed at the sampling site.

Harvest Surveys
Data consisting of total snipe bagged, total days hunted, and area most
frequently hunted were obtained from the 1972 and 1973 small game surveys

�-126-

conducted by the Colorado Division of Wildlife.
In 1973 and 1974,
questionnaires were sent to those respondents to the small game survey
who reported hunting snipe and to Colorado Division of Wildlife personnel for distribution to additional snipe hunters.
The special snipe
hunter survey requested information concerning area most frequently
hunted, total days hunted, number of hunters in party, total snipe seen,
total snipe bagged, and number of days when only snipe were hunted.

LOCATION AND DESCRIPTION

OF STUDY AREAS

Location
Four regions in Colorado were selected for investigations of snipe.
These regions were Fort Collins and North Park, both in north central
Colorado; the Yampa Valley in northwestern Colorado; and the San Luis
Valley in south central Colorado (Fig. 1).
Three study areas were selected in each region on which densities of
breeding snipe were estimated and habitat conditions influencing these
densities were evaluated.
In Fort Collins, the three areas (No.3, 4,
and S) varied from 10SoOO' to 10So04' W long. and 40°23' to 40°44' N
lat. with a total area studies of 2S.2 ha. Two additional areas (No. 1
and 2) were selected near Fort Collins on which numbers of wintering
snipe were estimated.
These areas ranged from 10SoOO' to 10So07' W long.
and 40°32' to 40°34' N. lat. and had a combined length of 6.4 km.
In North Park, locations of the three areas (No.6, 7, and 8) varied from
106°16' to 106024' W long. and 40°36' to 40°44' N lat. with a total area
of 92.8 ha. Locations of areas 9, 10, and 11 in the Yampa Valley ranged
from 1060S7' to 107020' W long. and 40029' to 40048' N lat. with a total
area of 63.S ha. In the San Luis Valley, areas 12, 13, and 14 varied in
location from 10S043' to 106006'W long. and 37°18' to 37034' N lat. and
had a total area of 63.6 ha. Total area for all regions was 24S.l ha
and varied in location from 10SoOO' to 107°20' W long. and 37°18' to
40048' N. lat.

Climate
Climatic conditions in Colorado are influenced mainly by its inland,
continental location and high elevation (mean altitude of 207S m above
sea level). The inland location results in wide daily and seasonal
temperature variations and low precipitation.
The high elevation affects
all climatic phases including increased precipitation in the mountains
and temperature modifications throughout the state (Colorado State
Planning Division 1964, Berry 1974).
Climates in local regions are greatly influenced by differences in
elevations and the orientation of mountain ranges. The climate of the
Fort Collins area is comparatively mild with long, warm summers (139-day
frost-free season) and open winters.
Precipitation is low (2S to SO cm)

�M~'Ar

wup

Ir()U"

SLD6WICI(

L06AN

""",n

II 10

MOil"""
"'ASltIW;TDN

rUMA

/f10 #LAN(O

GAII"ILD
Iflr CAlf SOli

""COUt

MISA
J0N
-...J

J
,,,trl,,111

1_

H

~I"'CA.O"W;-;LI:;,::-_"",, __

IJ KIOWA

'110 WIllS

UIIT

SAIIMI6UlL

DOLOlflS

MOlfrlZUIIAA
MCA

AIfClfIlUTA

Fig. 1.

Location

of snipe

study areas

in Colorado.

�-128-

and infrequent of which about 75 percent falls during the growing season (Robbins 1917). At Fort Collins, the mean annual temperature and
precipitation rate are 8.9 C and 35.8 cm (Colorado State Planning
Division 1964).
In North Park, the climate is typically continental and freezing temperatures may occur every month (56-day frost-free season).
Summer mean
temperatures are normally between 13 and 16 C and March and April
usually have the greatest precipitation (Robbins 1917). Walden, centrally
located in North Park, has a mean annual temperature and precipitation
rate of 2.3 C and 33.3 cm (Colorado State Planning Division 1964).
Climatic conditions in the Yampa Valley vary with elevation.
At lower
elevations, near Craig, the climate is mild (81-day frost-free season)
with comparatively long winters.
At higher elevations, near Steamboat
Springs, the climate is more severe with short, cool summers (75-day
frost-free season) and long winters with heavy snowfall (Colorado State
Planning Division 1964). July and August often have the greatest
precipitation although wide variations are common (Robbins 1917). Mean
annual temperatures and precipitation rates are 6.1 C and 31.8 em for
Craig and 4.1 C and 58.7 cm for Steamboat Springs (Colorado State Planning
Division 1964).
In the San Luis Valley, the climate is arid with less than 18 cm of
precipitation in central portions of the Valley and 25 to 30 cm along
the foothills (Bean 1962) with most precipitation normally occurring
from June through August. Mean annual temperatures vary from 2 to 6 C
with mean summer temperatures near 16 C (Colorado State Planning Division
1964). The frost-free season is approximately 103 days (Robbins 1917).

Topography
Topography is diverse among the four regions but similar within each
study area investigated.
The Fort Collins region, at an elevation of
about 1525 m, is part of the Colorado Piedmont.
This region is typified
by level or rolling prairie broken by streams, valleys, and occasional
sandhills (Colorado State Planning Division 1964). The five wintering
and breeding areas studies varied in elevation from 1480 to 1615 m.
Breeding areas were generally level with slopes of less than 3 percent.
Wintering area 1 was characterized by a narrow (1.5 to 3 m wide) shallow,
flowing stream with steep to perpendicular banks from 1.5 to 3 m high.
Area 2 also contained a shallow, flowing stream approximately 2 to 5 m
wide bordered by gentle to steep-sloping banks.
North Park has an area of about 1500 km2 and a mean elevation of 2475 m.
The Park floor is predominately rolling ane encompassed by mountains
(Fenneman 1931). Areas 6, 7, and 8 ranged in elevation from 2455 to 2510
m. Slopes were usually less than 3 percent or less but increased to 6
percent near ridges.

�-129-

The Yampa Valley is a long narrow floodplain varying in width from
0.4 to 16 km (Boek~r 1954). The Valley floor between Steamboat Springs
and Craig is gently sloping, declining in elevation from 1980 to 1890 m.
Breeding areas 10 and 11 are located in this region at elevations of
1975 m and 1915 m, respectively.
Area 9 is located at 2450 m near
Steamboat Lake. All areas in the Yampa Valley have slopes of 3 percent
or less.
The San Luis Valley is approximately 21,000 km2 in area (Colorado State
Planning Division 1964) and is bounded by mountains on all but the south
side. The basin floor is relatively flat with elevations ranging from
about 2300 to 2450 m (Fenneman 1931). Elevations for areas 12, 13, and
14 ranged from 2285 to 2320 m with slopes of less than 3 percent.

Soils
Data obtained from the U.S. Soil Conservation Service showed similarities
in major soil types on breeding areas 3 through 14. Major soil types
which characterized breeding areas near Fort Collins, in North Park, and
the San Luis Valley were deep, poorly drained, often alluvial with
excellent moisture holding capacity and slow to fair air, water, and root
penetration.
Surface layers ranged from clay loam to fine, sandy loam
often covered with a shallow, organic mat. Subsoils were generally clay
loam or sand, gravel, or shale.
In the Yampa Valley, soil types were more variable, ranging from deep,
well-drained loam or clay loam characteristic of mountain meadows (areas
9 and 10) to saline-alkali soils with a silty, clay loam surface and subsoil normally found in swales and valleys (area 11).
All areas in the Yampa Valley and area 8 in North Park contained a second
major soil type. This type is described as wet, alluvial land consisting
of a variety of continually wet soils of various depths and textures often
bordering streams and rivers. The surface of this soil is moderately
high in organic matter and mostly salt and alkali free. Descriptive and
quantitative data for major soil types on areas 3 through 14 are listed in
Table 1.
Snipe normally probe for food and prefer to feed in soft, organic soils.
To evaluate this preference quantitatively, measurements of ground compaction were taken on breeding areas 3 through 14 in late May and early
June 1974. The ground compaction was variable, influenced prinCipally
by soil texture, soil moisture, and vegetation densities. Locations with
the highest densities of snipe generally had ground compaction of less
than 0.1 to 0.75 kg/cm2, occasionally to 1.5 kg/cm2 in areas of dense
vegetation.
Areas with ground compactions of 2.5 kg/cm2 and greater were
slightly moist or dry and seldom provided feeding sites for snipe.
No soil descriptions or compaction measurements were collected for wintering
areas 1 and 2. The stream channel in area 1 was primarily gravel with few
protruding mud flats whereas area 2 contained many exposed, sparsely
vegetated mud flats which attracted snipe and were used as feeding areas.

�-130-

Table 1.

Area

Descriptive and quantitative data for m.ajor soil types on
snipe breeding areas 3 - 14 in Colorado.

Soil
texture

%

Water
table
depth
(cm)

Permeability
(cm/hr)
2.0 - 3.8

o. 16 -0. 21 7.5 -8.4

0.13-0.5

0.18 -0.21

8.4-9.0

Slope

Available
water
capacity
(cml cm)

Soil
reaction
(pH)

3

loam

0-3

50-130

4

clay

0-3

&lt; 30

5

clay loam

0-1

30 -100

0.5 - 2.0

0.18-0.20

7.7-8.4

6

clay loam

0-6

0-

30

0.5

- 1. 5

0.19 -0.21

7.9-8.4

7

coarse,
loamy -mixed

0-5

0-

15

1.5 - 5. 1

O. 13 -0. 18 7.9-8.4

8Y

loam; fine
sandy loam

0-2

15

5.1 -15.3

O. 13 -0. 18 6.6-7.3

91J

loam

0-3

60

1.5 - 5. 1

0.12-0.20

6.6-7.3

101) loam

0-3

120

1.5 - 5. 1

O. 12 -0.20

6.6-7.3

111) very fine
sandy loam

0-3

&gt; 180

1.5 - 5. 1

O. 12 -0.20

8.5-9.0

12

loam and
clay loam

0-3

50 - 90

0.5-5.1

0.16 -0.21

7.9 -8. 4

13

loam

0-1

30-100

1. 5 - 5. 1

0.16-2.0

7.9 -8.6

142)

In addition to soil data listed in Table 1, these areas contained extensive areas of wet, alluvial land with continually wet soils
of various depths and textures for which no quantitative data are
available.
No soil survey completed on this unit.

�-131-

Vegetation
Regional variations in vegetation reflect differences in such factors
as geographic location, elevation, and climate. The dominant natural
vegetation in the Fort Collins area is grass-steppe or short grass
formation (Robbins 1917). Currently, most land is used as pasture or
cropland. In North Park, sagebrush (Artemisia spp.) dominates, occurring
over approximately 65 percent of the Park. The remaining 35 percent
consists mostly of wetlands such as sloughs, oxbows, streams. lakes,
and flooded meadows and pastures (Beck 1976). Between Steamboat Springs
and Craig in the Yampa Valley, high meadows and farmland dominate. The
high water table has resulted in extensive sedge (Carex spp.) and cattail
(Typha spp.) marshes (Boeker 1954). In the San Luis Valley, a high water
table and alkaline soils have resulted in extensive black greasewood
(Sarcobatus vermiculatus) and rabbitbrush (Chrysothamnus spp.) associations
with sagebrush occurring on well drained areas. The region also retains
extensive coverage of hardy grasses such as gramma grass (Bouteloua spp.)
(Bean 1962).
Descriptions of vegetation on breeding areas were completed in late July
and early August 1974. Of the 12 areas, 7 (3-7, 11 and 12) were primarily
wet pasture partially flooded from early spring through at least late
summer often containing associations of Typha and Scirpus. Water sources
included irrigation, irrigation runoff from nearby croplands, springs,
and seepage. Species of Carex were the dominant plants on all seven areas.
Other plant species which occurred on a majority of the areas were ~
spp., Scirpus americanus, Hordeum jubatum, Trifolium spp., Scirpus
lacustris, Lemna spp., and Taraxacum spp. Major plant species for these
and subsequent areas are listed in decreasing frequency of occurrence.
Most areas were grazed by livestock beginning usually from late March
through early June and ending from early September into late October.
The remaining five areas had distinctly different habitats. Area 8 was
an irrigated riverbottom interspersed with wet depressions and oxbows.
Major plant species were Carex spp., Phleum pratense, Taraxacum spp.,
Trifolium spp., and Bromus inermis. Grazing was limited, occurring from
early July to early October.
Area 9 was ungrazed highland meadow with corridors of Salix spp. along
stream channels. Major plants were Carex spp ., Taraxacum spp ,, Salix
spp., Trifolium spp., and Poa pratensis.
Area 10 consisted of irrigated hayland and a flooded oxbow with Typha
marsh. Major plant species were Carex spp., Trifolium spp., Bromus intermis,
Typha spp., Salix spp., Phalaris ~inacea,
and Sporobolus airoides.
The area was ungrazed but the vegetation was mowed for hay in late summer.
Area 13 was marshland dominated by Eleocharis spp., Juncus arcticus, Carex
spp., Typha spp., and Scirpus lacustris. The area was bordered by
Sarcobatus vermiculatus in alkali soil.
Area 14 contained two large ponds (5 ha total area) and associated wet
meadows bordered by Sarcobatus vermiculatus.
Major plant species were
Eleocharis spp., Juncus arcticus, Hordeum jubatum, Scirpus americanus,

�-132-

Carex spp., and ~.pallidus.
Limited grazing occurred on both areas
13 and 14 from early July through at least September. Major plant
species for each breeding area are listed in the Appendix.
Vegetation on wintering areas 1 and 2 was primarily grasses (Graminae)
but no plant collections were made. On area 1, vegetation varied from
ungrazed near the stream channel to well grazed on the crests of the
banks. Vegetation on area 2 was more diverse and generally grazed or
mowed.

Water Characteristics
Water samples on all breeding areas were analyzed in June 1975. Results
of the analyses are summarized in Table 2. Variations between areas were
generally small. No free acidity was measured on any area. Acidity of
any natural water is normally low unless strongly acidic industrial waters
are introduced (Hach Chemical Company 1973).
Phenolphthalien alkalinity was detected on only one unit and total
alkalinity varied from 50 to 510 mg/l (as CaC03). The presence of carbonates, bicarbonates, mid hydroxides of ~alcium, magnesium, and sodium are
the most common causes of alkalinity in natural waters (Hach Chemical
Company 1973). Alkalinity in natural waters may have a wide range but
rarely exceeds 400 to 500 mg/l (as CaC03)' Generally, waters with reasonably
high bicarbonate alkalinity result in increased aquatic plant growth
(National Academy of Sciences 1973).
Hardness was the most variable characteristic ranging from 30 to 2100
mg/l (as CaC03).
Generally, the biological productivity of water is
directly related to its hardness.
However, while calcium and magnesium
contribute to hardness, other contributing elements in high concentration
may be toxic thereby reducing productivity (National Academy of Science
1973).' It should be noted that acidity, alkalinity, and hardness are
normally reported as equivalents of specific values of CaC03 solutions,
although actual constiLuents measured in each test may be entirely different (Hach Chemical Company personal communication).
Values for pH ranged from 7.2 to 9.5, thus all areas were slightly to
moderately basic. Most natural waters range from pH 4 to pH 9 and are
often somewhat basic because of the presence of carbonates and bicarbonates
(Hach Chemical Company 1973).

�-133-

Table 2.

Area

Results of water analyses on snipe study areas
Colorado, June 1975.

Free
Phenolphthatln
acidity V
alkalinity 1

3 - 14 in

Total
-alkalini ty V

HardnessV

pH

3

0

0

340-410

500-550

8.0 -8.2

4

0

0

140-310

1680-2100

7.6 -9.0

5

0

0

310 -360

550-800

7.8-8.8

6

0

0

240-510

270-740

7.8 -8.8

7

0

0

120-190

120-190

7.8-8.2

8

0

0

90-120

100-150

7.5-7.6

9

0

0

50 -90

30-90

7.2 -8.3

10

0

0

150-310

170-410

7.7 -8.9

11

0

0

290-460

260-460

8.5 -8.8

12

0

0

70 -170

70 -140

7.2-7.7

13

0

0

190 -290

120-170

7.7-8.8

14

0

0-30

150 -260

50-70

8.8 -9.5

1) Expressed

.a s rng/l

as CaC0 .
3

�-134-

RESULTS AND DISCUSSION

Estimates of Breedjng Densities
The relation between numbers of snipe recorded during spring and
summer censuses and actual numbers of snipe breeding on each area
proved difficult to evaluate. Results of censuses prior to and during
early May could possibly overestimate densities of breeding snipe by
including spring migrants.
Conversely, census data after early May
could underestimate these densities by excluding nesting snipe which
were difficult to flush. Consequently, estimated breeding densities
for each area were based on censuses conducted throughout May of 1974
and 1975 for Fort Collins, North Park, and the Yampa Valley. In the
San Luis Valley, breeding densities were based on censuses during May
1974 and late April through early June 1975 because only two May censuses were conducted in 1975. This procedure minimized overestimating
densities because of migration and underestimating densities because
of nesting snipe.
Densities of snipe utilizing areas 3 through 14 for breeding are tabulated in Table 3 and depicted in Figs. 2 - 5. These data show an
overall variation in breeding densities from 0.2 to 2.1 snipe/haG Of
the four regions, Fort Collins had the highest mean density of 1.5
snipe/ha for 1974-75. This high density was a reflection of the small
size and uniformly suitable habitat for snipe on each of the three Fort
Collins breeding areas. Study areas in the other three regions were
larger and contained more diverse habitats of various suitability for
snipe. Consequently, regional variations in snipe densities are less
pronounced than indicated in Table 3. The lower densities in 1974
probably resulted from late censuses in May.
Numbers of snipe varied with habitat factors, primarily water depths
and coverage, vegetation heights and densities, and soil conditions.
Those areas providing the most suitable habitat for snipe contained
shallow, stable, discontinuous water levels. Vegetation was low
(10 - 30 cm in late May), often grazed or mowed, and sparse. Soils
were moist to water saturated and frequently characterized by hummocks.
Areas 3 through 7, 11, and 12 were partially flooded pastures. These
areas had densities for 1974-75 ranging from 0.4 to 1.9 snipe/ha with
a mean of 0.8 snipe/ha on a combined area of 138 haG Of the seven
areas, those having characteristics described above had typically high
densities whereas areas with tracts of dry soil or extensively overgrazed or ungrazed vegetation had low snipe densities.
Of the remaining five areas, areas 9 and 10 had relatively high densities
of 1.2 and 1.0 snipe/ha, respectively.
Water levels on both units were
stable at least through May and were maintained by beaver dams on area 9
and irrigation water on area 10. Both areas were ungrazed; however,
vegetation heights on area 9 were normally low and area 10 was mowed in
late summer. Vegetation heights on both areas in late May ranged from
5 to 30 cm.

�-135-

Table 3.

Estimated densities of snipe breeding
14 in Colorado, 1974 and 1975.

on study areas

Region

Size

and Area

(ha)

1974

1975

3

4.5

1.7

2.1

4

7.7

0.8

1.2

5

13.0

1.5

1.8

Subtotal

25.2

1.3

1.7

6

17.4

0.9

1.2

7

38.9

0.4

0.6

8

36.5

0.6

0.3

Subtotal

92.8

0.6

0.6

9

10. 1

1.1

1.2

10

19.0

0.8

1.1

11

34.4

0.4

0.5

Subtotal

63.5

0.5

0.7

12

21. 9

0.7

0.8

13

17.4

0.5

0.5

14

24.3

0.2

0.3

Subtotal

63.6

0.5

0.5

245. 1

0.6

0.7

Estimated

Number of Breeding

Fort Collins

North Park

Yampa Valley

San Luis Valley

Total,

All Areas

3 _

SnipejHa

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JUNEI3-14

JUNE6-7

MAY 30-31

0
0

MAY 22-23

MAY 15

MAYS-9

MAY 1-3

~

Ul

P

,....

I-'
I-'

-.j

(0

0

e-t-

'i

0

f-rJ

APR25-26

APR17-19

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(1)

APR5-6

MAR29-30

OCT 10-17

OCTI-3

APR10-13

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SEPT 17

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SEPT 24-26

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JULY 9-10

JUNE30-JULY 2

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JUNE 9-10

JUNE 1-3

MAY 19-20

MAY 12-14

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NUMBER OF SNIPE PER HECTARE

MAY 26-27

MAY 6-7

MAY 5

APR22-23

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APR8

OCT 18

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SEPT 21

SEPT 13

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JULY 3-4

JULY 2-3

JUNEI2-13

JUNE I 1-12

MAY 24-25

MAY 23-24

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•....•

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JUNEI2-13

JUNE 11-12

MAY 29-30

MAY2S-29

MAY 7

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APR23

APR22

APRS

APR7

SEPT 14

JULY 22-23

JULY 5-6

JULY 4-5

JUNE 14-15

JUNE 13-14

MAY2S-30

MAY 27-29

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NUMBER OF SNIPE PER HECTARE

�-140-

Areas 8, 13, and 14 provided few suitable habitats for snipe and had
mean breeding densities of 0.4, 0.5, and 0.3 snipe/ha, respectively.
All three areas were lightly grazed from early July through September.
Consequently, vegetation was ungrazed during May and June and vegetation heights in early June were generally from 20 to 70 cm. Water
levels were relatively stable on areas 8 and 13. Both received
irrigation water although irrigation was delayed by 3 weeks on area 8
in 1975. This may have been partially responsible for the decrease
in snipe densities at this site in 1975. Water levels on area 14
declined through May and June in 1974 and 1975 making the area increasingly unfavorable as snipe habitat.
Breeding densities for snipe reported in the literature are limited.
Tuck (1972) tabulated breeding densities on three habitat types in
the peatlands of Canada. Densities ranged from 5.5 to 13.2 pair/lOO
ha (0.11 to 0.26 snipe/ha) in sedge bogs; from 3.5 to 7.7 pair/lOO ha
(0.07 to 0.15 snipe/ha) in fens; and from 9.5 to 12.7 pairs/lOO ha
(0.19 to 0.34 snipe/ha) in willow and alder swamps.
In addition to the 12 breeding study areas, 44 areas in the San Luis
Valley, North Park, Yampa Valley, Fort Collins region, South Park, and
along Tomichi Creek near Gunnison were investigated from late April
through early June 1975. The areas consisted of pastures and haylands
that were irrigated after late April and early May. Brief censuses
were conducted on each area and measurements of water depths, vegetation
heights, and ground compaction and estimates of water coverage were made.
No snipe were found on 82 percent of the 44 areas. On the remaining 18
percent, about three snipe were heard or seen per census. Water depths
and vegetation heights on these areas were usually 5 cm or less, rarely
exceeding 10 cm. Areas were typically level and water coverage was
extensive.
Ground compaction generally exceeded 1.5/kg2 and seldom was
below 1.0 kg/cm2•
By contrast, on habitats with substantially more snipe,
vegetation heights ranged from 10 to 30 cm and ground compaction from
less than 0.1 to 0.5 kg/cm2•
Although snipe utilized pastures and hayland, these areas have limited
suitability as breeding habitat for snipe for several possible reasons.
Irrigation begins in late April and May, probably after most snipe have
selected nest sites in other locations.
Furthermore, vegetation heights
of 5 cm or less provide poor cover, particularly for nesting.
Soils,
unless flooded for a considerable time, may remain too firm for easy
probing.
Finally, because of the generally level terrain, water coverage
was often continuous, ?roviding few unflooded sites for nesting.

Nesting Activities
All nest searches were done in conjunction with preliminary reconnaissance
and censuses of breeding study areas. These were conducted from late
March through mid-October 1974 and late March through mid-June 1975.
During these periods, 28 snipe nests were located.

�-141-

Nest sites were typically in grasses or sedges 20 to 40 cm high and
on moist but unflooded ground near water. An atypical nest site was
found in the base of a 3-m high willow (Salix spp.) which was part
of an elongated willow stand about 30 m wide.
Estimated onsets of incubation for all nests ranged from 2 May through
4 July (Table 4) with the bulk of nesting activity occurring in May.
Based on 1974 nest data, over 60 percent of the nesting snipe located
had begun incubatiun during May. Nesting may have extended into
August as indicated by a snipe flushed on 7 August 1974. The snipe
displayed a distress posture associated with distraction and brood
or nest protection, but neither a brood nor a nest was located.
Clutches of four eggs were found in 26 nests (93 percent) and clutches
of three eggs in two nests (7 percent). Forty-nine eggs were measured
having a mean length of 38.5 mm and a mean width of 28.4 mm with
standard deviations of 1.3 mm and 0.7 mm, respectively.
Measurements
ranged from 36.6 to 41.6 mm in length and 27.2 to 29.7 mm in width.
Spring Migration
From data collected on 12 breeding areas, the extent and peak of the
spring migration appeared indistinct. During censuses initiated in late
March and early April 1975, low numbers of snipe, probably migrants,
were observed on most breeding areas which were snow- and ice-free.
By mid-April, maximum numbers of snipe were recorded in all regions with
Fort Collins having a peak of 5.6 snipe/ha on 10 - 13 April, North Park
with 1.2 snipe/ha on 21 - 23 April, Yampa Valley with 1.1 snipe/ha on
22 - 23 April (omitting area 9 which was snow-covered until late May),
and San Luis Valley with 1.0 snipe/ha on 14 - 16 April (Table 5).
Numbers of snipe declined slightly but generally remained high through
early May, particularly in the San Luis and Yampa Valleys. This limited
decline was probably the result of continued northward migration.
A
second decline occurred in all regions from early May to late May which
may have been caused, in part, by a continuing northward migration.
Although migrating snipe begin arriving in Canada during April and early
May (Tuck 1972), breeding populations in Canada may not peak until late
May (Arnold 1975). Local movements to areas such as irrigated haylands
and pastures which had been previously dry and the onset of nesting and
incubation which made snipe more difficult to flush probably added to
the apparent decrease of snipe. After late May, numbers of snipe
stabilized on nlost study areas until the termination of censuses in midJune (Figs. 2 - 5).
From these data, spring migration in 1974 apparently began by late March
or early April. Peak numbers of snipe occurred in all regions and on
all open study areas except one from April 10 to 23. Migration continued
thereafter through at least early May and possibly into late May.
Fall Migration
Numbers of snipe were monitored during September and October 1974 on 8
of the 12 breeding areas in order to document timing of the fall migration.

�-142-

Table 4.

Snipe nesting

data in Colorado,

Date

1974 and 1975.

Estimated

Onset

Year

Area

Located

of IncubationU

Clutch Size

1974

Yampa Valley
3
5
San Luis Valley
8
8
8
7
10
9
9
9
5
12
9
10
10
5

May 1
May 12
May 14
May 15
May 23
May 23
May 23
May 25
May 28
May 29
May 30
June 13
June 16
June 20
July
4
July
5
July
5
July
9

May 4
Not Estimated
Not Estimated
Not Estimated
May 19
May 21
May 16
May 23
May 21
May 15
May 12
June 12
May 28
June 19
July
2
June 27
July
3
July
4

4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4

1975

3
5
9
10
5
8
9
10
9
12

May 8
May 9
May 28
May 29
May 31
June 10
June 11
June 12
June 12
June 15

May 7
May 2
May 23
May 20
May 17
June
1
May 28
May 31
June 7
June
5

4
4
3
4
4
4
4
4
3
4

lj

Based on a 19 -day incubation period
technique described
by Westerskov
(1950).

using the water

flotation

�Table 5,

Maximum densities of snipe with date of occurrence
and fall (1974) migration in Colorado.

Region
and area

Spring Migration,
1975
Maximum Density
(snipe/ha)

Date

for areas

3 - 14 during spring

Fall Migration,
1974U
Maximum Density
(snipe/ha)

(1975)

Date

Fort

Collins

5.6

April

10-13

2.9

Sept,

17

3

7.3

April

10

6.0

Sept,

7

4

5.8

April

10

6.0

Sept.

17

5

5.8

April 19

2.9

Sept.

17

North Park

1.2

April 22 -23

3.4

Sept.

21

6

1.9

April 22

7.9

Sept.

21

7

1.6

April 22

2.3

Sept.

13

8

0.4

April 23

1.1

April 22

Yampa Valley
9

Y

10

1.5

April 22, May 6

11

0.9

April 22

I
•...

+:w
I

�Table 5 Cont ' d.
Region
and area

Spring Migration,
1975
Maximum Density
(snipe/ha)

Date

Sept.

15-16

Aprll 16

0.6

Sept.

16

0.9

April 16

0.04)

0.7

April 14

0.4

Sept.

27

April

12

1.4

13
14

1) Censuses

Date

0.33)

1.0

San Luis Valley

Fall Migr ation, 19741)
Maximum Density
(snipe/ha)

not conducted on areas

15-16

~
~
~

8 - 11.

I

2j Snow-covered

3j Censuses
densities.

through mid-May,

1975.

during early August showed densities

~/ No snipe recorded

through September,

1974.

(0.9 snipe/ha)

which exceeded September

�-145-

Three areas in the Yampa Valley and one area in North Park were eliminated because of unfavorable water conditions and/or dense, ungrazed
vegetation.
Areas near Fort Collins and in North Park offered a
suitable habitat for snipe through the conclusion of censuses in midOctober. Censuses in the San Luis Valley were terminated in late
September because of declining numbers of snipe.
Data from September and October censuses indicated fall migration was
in progress by early September. During these censuses, maximum
numbers of snipe were observed in the San Luis Valley with 0.3 snipe/ha
on 15 - 16 September, in Fort Collins with 2.9 snipe/ha on 17 September,
and in North Park with 2.1 snipe/ha on 21 September (Table 5). These
data indicated that the peak fall migration period for 1974 apparently
occurred during the third week in September. This conclusion is based
on total numbers of snipe by region. This period coincided with peak
numbers on four of the eight study areas. Two areas had peak numbers
the second week of September, one area the fourth week, and one area
had no snipe through September (Table 5). Numbers of snipe declined
considerably after late September and by mid-October the fall migration
was near completion and numbers were too low to warrant continued
censuses (Figs. 2, 3, and 5).
Although the majority of fall migrants evidently passed through Colorado
during September, substantial increases in numbers of snipe were observed in August. Censuses conducted in early August on Fort Collins
and San Luis Valley study areas indicated such increases on four areas
(Figs. 2 and 5) whereas censuses in late July in North Park and the
Yampa Valley revealed no similar increases (Figs. 3 and 4). Densities
of snipe in the San Luis Valley reached 0.9 snipe/ha on August 9 - 10,
exceeding the September maximum of 0.3 snipe/ha. This increase may
indicate that the fall migration was underway by early August, or
juvenile snipe had concentrated on favorable feeding grounds. Tuck
(1972) reported th~t in Newfoundland, juveniles aggregated in groups
of a few birds in late July to flocks of a hundred or more in mid-August
and were likely to migrate together ahead of adults.

Estimates of Winter Densities
Twelve censuses were conducted on area 1 from late January 1974 through
June 1974. Thereafter, censuses were discontinued because of low numbers
of snipe utilizing the area during winter months. On area 2, 30 censuses were conducted from late January 1974 through early May 1975.
Because census routes followed streams with limited widths, densities
of snipe were translated into snipe/km rather than snipe/ha.
Densities on area 1 were low with a mean of 0.8 snipe/km from January
through April 1974 and a maximum density of 1.7 snipe/km recorded in
mid-April.
From mid-May to the termination of censuses in July, no snipe
were flushed from the route (Fig. 6).

�E E

..:II: ..:II:

-(\IV

~~

-146-

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Ol~d~

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t--

I
~
.....•
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f-:l CIl

::J

~u

~ 0

.....•

.....•
0

0'

t--. I'd

"&lt;!''"(j

I'd
_0

"8

I'd

I-&lt;

Q)

I'd

u,

~
o

f3

o

en
w
en
:J
en
z
w
o

8a~~W
8~~W
aa83.:1
683.:1
9aN~r
alN~r
17a:&gt;30
£ :&gt;30
17aI\ON
OII\ON
Oa.1:&gt;o
£1.1:&gt;0

619n~

aa.1d3S
J...1d3S

c:tc:t

-(\I

-0::0::

ww

za-oaser

01-£ 83.:1

6-£~~W
17a-J..183.:1

oa-91~d~
01~d~-6a~~w
ga-J..I~~w

17A~W-8a A~W

61-£IA~W

01-173Nnr

171-1Alnr
J..a-~13Nnr

c:tc:t

II
omOO~~~V~N-omOO~~~V~(\I-O
(\1----------

~3.13W011)l ~3d 3dlNS .:10 ~38WnN

�-147-

Densities on area 2 were considerably higher than those on area 1
with a mean density of 7.5 snipe/km from January through mid-April
1974. Thereafter, numbers of snipe declined precipitously and snipe
were absent from the area from May through September with the exception of one snipe flushed in August. Snipe returned in early
October in low numbers (1.8 snipe/km) and reached a maximum density
of 16.5 snipe/km in late December. Densities from January through
mid-April 1975 declined to an average of 11.2 snipe/km and were more
variable than the previous spring. As in 1974, a substantial decrease
was noted in late April and all snipe had departed by early May
(Fig. 6).
It is apparent th.~t snipe near Fort Collins occupy winter habitat
from October through April; however, factors responsible for the
arrival and departure of snipe to and from these areas are unknown.
Presumably, stream habitat of this type does not provide suitable
sites for breeding unless associated with sloughs, marshes, or other
flooded areas.
Large increases in numbers of snipe during November did not appear to
be correlated with any weather phenomena. Weather conditions were
mild during this time and surface water in many breeding areas remained unfrozen. Consequently, increased utilization of winter habitat
was apparently not a result of decreased suitability of some breeding
habitats.
Snow and ice accumulations on winter areas influenced the
distribution rather than total number of snipe. During such periods,
snipe concentrated in ice- and snow-free sites.
Overall, the suitability of winter habitat appeared to be primarily
influenced by topography, vegetation densities and heights, and water
levels relative to the shoreline. Snipe were flushed most frequently
in areas bordering stream channels which had gently sloping banks or
exposed mud flats with organic soil and low, sparse vegetation.
Area
1 had few of these areas whereas area 2 had many such sites resulting
in relatively high numbers of wintering snipe.

Harvest Assessment
The harvest of snipe in Colorado has been estimated from a survey of
small game license holders conducted annually by the Colorado Division
of Wildlife.
In 1972 and 1973, a total of 22 respondents to this survey reported hunting snipe. Results indicated these hunters harvested
a mean of 3.7 snipe per hunter and spent 4.0 days afield during the
1972 and 1973 hunting season.
In order to obtain more detailed information concerning snipe harvest,
attempts were made to identify and contact snipe hunters. Questionnaires
(38 in 1973 and 35 in 1974) were sent to the 22 respondents to the small
game survey and to Colorado Division of Wildlife personnel for further
distribution to snipe hunters. Of these 73 questionnaires, 19 (26 percent)
were returned. Of those hunters responding, eight (42 percent) reported
that they had hunted snipe in 1973 or 1974. These hunters harvested a

�-148-

a mean of 10.8 snipe per hunter and spent 4.9 days afield (Table 6).
Additional information from the questionnaires indicated that the
size of the hunting party varied from 1 to 3 persons and the respondents encountered from 1 to 350 snipe during the season with an
average sighting of 110 snipe.

Table 6.

Results of the 1972 and 1973 small game surveys and
the 1973 and 1974 snipe hunter questionnaires for
Colorado.

1972 and 1973
Small Game
Surveys 1../

1973 and 1974
Snipe Hunter
Questionnaires !/

Number of respondents
hunting snipe

22

8

Snipe harvested

3.7

10.8

Days afield per hunter

4.0

4.9

Snipe harvested per day

0.9

2.2

per hunter

Data from both years are combined.

The harvest of snipe based on the results of the special snipe hunter
questionnaires was considerably higher than the harvest indicated by
the small game surveys (Table 6). Possibly, respondents to the special
snipe hunter questionnaires represented persons who hunted snipe
specifically whereas respondents to small game surveys represented those
who hunted snipe incidentally to other game species. Of the respondents
to the small game survey who reported hunting snipe in 1972 and 1973,
only 33 percent hunted snipe the following one or two years. It is
obvious that snipe are a minor game species in Colorado and that the
annual harvest is small.

Prepared by

�-149-

LITEEATUEE

Aiken,

C. E.,
County,

and E.

R.

Colorado.

CITED

Warren.

1914.

Colorado

Co l l , Publ.

The birds

of E1 Paso

Sc i, Ser.

12:455 -603.

American
Ornithologists'
Union.
1957.
Checklist
of North
birds.
Lord Baltimore
Press,
Inc.
Baltimore,
Md.
Anonymous.
26(1):5.

1886.

Winter

snipe

in Colorado.

Forest

American
691pp.

and Stream

Arnold,

K., (Chairman).
1975.
Snipe (Capella
gallinago)
in G. C.
Sanderson,
e d , Management
of migratory
shore and upland
game birds in North America.
Int. Assoc.
Game,
Fish,
and Conserve
Corrirn, , Washington,
D. C. In press.

Bailey,

A. M., and E. J. Ni e d r a ch ,
2 Vol.
Denver Mus. Nat. Bist.

1965.
Birds of Colorado.
Denver,
Colo. 893pp.

Bean,

L.
J.,

sky people.

Beck,

T. D. 1. 1976.
Effects
of sagebrush
control
on distribution
and abundance
of sage grouse.
Final Report.
Colo rado Di v,
Wildl. Game Res. Rept.,
Fed. Aid Proj.
W -37 -R, April.
In press.

Beidleman,
Univ.

E.
1962.
Land of the blue
Monte Vista,
Colo. 119pp.

R. G.
1955.
Guide to the winter
Colorado
Mus. LeaH.
12.
6lpp.

birds

The Monte

Vista

of Colorado.

Bent,

A. C.
1927.
Life histories
U. S. Natl. Mus. Bull. 142.
D. C. 420pp.

Berry,

J. W. 1974.
The climate
of Colorado.
Pages 595-613 in
Offices
of the National. Oceanic and Atmospheric
Administration,
U. S. Department
of Commerce.
Clim.ate of the
States.
Vol. 2 -- We s t e r n States.
Water Information
Center,
Inc.,
Port Washington,
N. Y. 975pp.

Blair,

F. W., A. P. Blair,
P. Bradkorb,
F. R. Cagle,
and G. A.
Moore.
1968.
Vertebrates
of the United States.
McGrawHill Book Co. , Inc , , New York. 616pp.

of North American
shorebirds.
Smithson.
In s t ; , Washington,

�-150-

Boeker,
H. 1\1. 1954.
A census
and sora rail in the Yarrrpa
56(2):105-106.

of populations
River Valley,

of the Wilson snipe
Colorado.
Condor

Booth,

T. W., Jr.
1968.
The availability
and utilization
of the
food s of the common snipe (Capella
gallinago
delicata)
in
the rice growing regions
of southwestern
Louisana.
M. S.
Thesis.
Louisiana
State Univ.,
Baton Rouge.
160pp.

Bryant,

H.
snipe

C.
1915.
Two records
of the nesting of the Wilson
in California.
Calif.
Fish and Game 1(2):76-77.

Burleigh,
T. D. 1952.
Wilson's
snipe population
at Gray's
Lake,
Idaho.
Pages 53-55 in Investigations
of woodcock,
snipe,
and rails in 1951.
U-:-S. Fish Wildl. Servo Spec. Sci. Rep. Wildl.
14. Washington,
D~C. 58pp.
Clark,

E. R.
1971.
Measurements
of hunter kill of woodcock,
snipe,
and rails.
Page 31 in U. S. Bur. Sport Fish and Wildl.,
Migratory
Bird Populations
Stat. Ann. prog.
rep.,
fiscal
year--1971.
Laurel,
Md. 71pp.

Colorado
State Planning
Division.
1962-1964.
Denver.
1064pp.
Cooke,

W. W. 1897.
The birds
Agric.
ceu.. Fort Collins.
1898.
44:148-176.
No.4.

Agric.

1964.

Colorado

of Colorado.
239pp.

Bull.

year

book:

37.

State

Further
notes on the birds of Colorado.
State Agric.
Co Ll ; , Fort Collins,
Tech.

1900.
The birds of Colorado.
Agric.
Exp.
ColI.,
Fort Collins,
Bull. 56:179-239.

1909.
The bird s of Colorado
26( 4): 400 -422.
Erickson,
A. B.
53(1):62.

1941.

A study

- -third

of Wilson's

1945.

The Wilson's

snipe.

1953.

The j a ck s n ip e ,

Stn.

supplement.

snipe.

Conserv.

Nat.

State

Auk

Wilson

Bull.

Volunteer

8(44):

39 -43.
Minnesota

Bull.
Series

3:39-41.

�-151-

F'o

rria
M.
McGraw-Hill

rme

n

Hach

,

N.

1931.
Physiography
of the
Book Co.,
In c ; , New York.

Chemical
Company.
1973.
e d , Ames,
Iowa. n.p.

Johnson,
H. C.
1(1):26.

1899.

Nesting

Water

We

Analysis

of Wilson's

snipe

r
534pp.
st

e

n

United

Handbook.

in Utah.

States.

First

Condor

B. Reed.
1974.
Recent
Valley,
northwestern

Martin,

S. G., P. H. Baldwin,
and E.
records
of birds from the Yampa
Colorado.
Condor 76(1):113-116.

Nancy,

R. H.
1973.
Development
of external
sexing and aging
techniques
and trapping
method s for the common
snipe in
Louisiana
with notes on habitat
and behavior.
M. S. Thesis.
Louisiana
State Urriv , , Baton Rouge.
55pp.

National

Academy

of Sciences.

Environmental

Studies

Board.

Committee
on Water Quality
Criteria.
1973.
Water Quality
Criteria
1972.
U. S. Environ.
Protection
Agency.
Ecol.
Res. Sere E;PA-R3-73-033.
Washington,
D.C.
594pp.

Ni ed r a ch,

R. J. and R. B. Rockwell.
and mountain
parks.
Colorado
S e r , No.5.
19 6pp.

Packard,
F. M.
Colorado.

1939.
The birds of Denver
Mus. Nat. Hi st , Denver,
Pop.

1945.
The birds of Rocky
Auk 62( 3): 371 -394.

Mountain

National

Park,

Robbins,
C. S.
1954.
Further
investigations
on winnowing
method
of measuring
Wilson's
snipe populations.
Pages
61-65 in
Investigations
of woodcock,
snipe,
and rails
in 1953.
U. S.
Fish Wildl. Servo Spec. Sci. Rep. - Wildl.
24.
Washington,
D. C. 68pp.
Robbins,
F. W.
1917.
Native vegetation
and climate
of Colorado
in their relation
to agriculture.
Agric.
Exp. St.n; , Colorado
Agric.
CoIl.,
Fort Collins.
56pp.
Schmidt,
G. D. 1962.
Endoparasites
of Wilson's
snipe,
gallinago
d e l ic a ta ) Or d , in northeastern
Colorado.
Thesis.
Colorado
State Un iv ; , Fort Collins.
65pp.
Sclater,

W. L.
1912.
A history
and Co. Middle Row Place,

of the birds of Colorado.
London.
576pp.

(Capella
M.S.

Witherby

�-152-

Soil Conservation
Service.
1970.
Use and explanation
pretation
sheets.
Denver.
7pp.

of soil inter-

Solman,
V. E. F.
1954.
Wilson's
snipe "winnowing"
counts in
eastern
Canada,
1953.
Page s 57 -59 in Investigations
of
woodcock,
snipe,
and rails in 1953. U. S. Fish Wild l. Servo
Spec. Sci. Rep. -Wildl.
24.
Washington,
D. C. 68pp.
Stewart,
R. E. and H. A. Ka nt r ud ,
snipe in North Dakota.
Prairie

1968.
Nesting
Na t, 1(1):3-5.

Stout,

G. D. (ed.).
Viking Press,

Tuck,

L. M.
1955.
Some observations
on nocturnal
activities
of Wilson's
snipe -- Newfoundland
(1953).
Pages 41-50 in
Investigations
of woodcock,
snipe,
and rails in 1954.
U:s.
Fish Wildl. Servo Spec. Sci. Rep. -Wildl. 28.
Washington,
D. C. 62pp.

Wildl.

1967.
The shorebirds
N. Y. 270pp.

of North

of the common

America.

1972.
The snipes:
a study of the genus Capella.
Se r v , Monogr.
Ser. No.5.
Ottawa.
429pp.

Can.

Van Owens,
J. ·1967.
Food habits of the common snipe (Capella
gallinago
delicata)
in the pastures
of south central
Louisiana.
M. S. Thesis.
Louisiana
State Urri v; , Baton Rouge.
108pp.
Westerskov,
K.
bird eggs.
White,

1950.
Methods of determining
J. Wildl. Manage.
14(1):56-67.

M. and S. W. Harris.
and habitat use of snipe
Manage.
30( 1): 23 -24.

Whitehead,
C., Jr.
snipe (Capella
with ecological
and aging.
200pp.
Wolfe,

1966.
Winter occurrence,
in northwest
California.
J.

food,
Wildl.

1965.
Foods and feeding habitats
of the common
gallinago
delicata)
in Cameron
Parish,
Louisiana,
notes and a discussion
of methods
of sexing

M. S. Thesis.

L. R.
1931.
33( 2):49 -59.

the age of game

The breeding

Louisiana

State

Limicolae

Univ.,

Baton

of Utah.

Condor

Rouge,

�-153-

APPENDIX

�APPENDIX

Vegetative

species

and presence

Vegetative
Agrostis

Species

pa1ustris

1 = less than 10 cm
2 = 10 - 30 ern
3 = 30 - 60 ern
4 = 60 - 100 cm
5 = more than 100 cm

(30% or more)
(10 - 30%)
( 5 - 10%)
( 1 - 5%)
(1% or less)

3

4

5

03

02

02

c=:»

Study Areas
10
8
9

--

Asce1epias

in ca r na ta

Aster

13

14

S4
04
13

spp.

Beckmannia

12

13

A rtemi s ia ca na
tridentata

11

02

spp.

Artemisia

I
I-'
VI
.j:'I

05

Al ope cu r u s spp.
Artemisia

3 - 14.

Height, measured
in late
July and early August

Frequency of occurrence
with
approximate
percent coverage
A = abundant
C = COmlTIOn
S = scattered
o = occasional
I = infrequent

on area

syzigachnc

03

14

�Vegetative
Bromus

Species

inermis

---

3

4

5

6

7

02

Study Areas
8
10
9
03,4

14

11

C2

Carex festivella
Ce r a s t iu rn spp.

11

Chenopodium

13,4

spp.

Chrysothamnus

nauseosus

r

s

iurn
--

a

r

Deschampsia

ve

n

A3

A3

s

A3,4

A3

A4

03

03

13

C4

03

14
14,5

14

C2

Di s t i chl i s stricta

spp.

Equisetum

spp.

\J1
\J1

03

caespitosa

Eleocharis

I
I-'

I

e

angustifolia

S3

02

14

E1eagnus

A2,3

03

Cicuta doug1asii
C'i

A3

S3

spp.

C4

03
A2

Chrysothamnus

14

04

Caltha 1epto s epa1a
spp.

13

C5

Ca l.arriag r o s t i s inexpansa

Carex

12

02
15

02

12

A3,4

A3

15
S2
13

13

�Vegetative

Species

3

Evo1vu1us nuttallianus

11

4

5

6

7

Study Areas
10
8
9

11

12

13

14

04

Festuca

rubra

Fragaria

spp.

Gaillardia

spp.

Glyceria

maxima

Hordeum

spp.

Hordeum

brachyantherum.

Hordeum

ju ba tu rn

11
12
IS

IS

14
04

I
I-'

C2
C2.3 C3

52

I3

VI

53

I3

14

C3

A4

C3

13

I3

Iris rn i s s ou r i c n s i s

I3

--

C3

Juncus

a rct i cu s

Juncus

nodosus

03

Juncus

tenuis

C3

I3
01

Kochia spp.
51

Le rrma spp.
Melilotus

officinalis

Muhlenbergia

rni nut i s s irna

01

13

11
11

01
04
I3

'"
I

�Vegetative

Species

Pedicillaris

3

4

5

6

Stud y Areas
10
8
9

groenlandica

Pha1aris

arundinacea

Phleum

spp.

14,5

04

13

Poa pratens is

02

I
I-'
In

spp.

12, 3

Po Iyg onurn pennsylvanica

03

Polygonum

01

rurivagum

Polygopon monspeliensis
spp.
anserina

Potentilla

gracilis

Puccinellia

spp.

13

'-l

I

12
15

Potentilla

14

02,3

Poa neva den s i s

Populus

13

C4

--

Polanisia

12

05
13

criopoda

11

13

Phleum pratense
Plantago

7

02
13
03

13

�Vegetative
Puccinellia

Species

4

5

6

7

Study Areas
10
8
9

11

13

Sagittaria

01

Il

01

13

I5

Sarcobatus

05

I5

ve r rn i cu Ia tu s

14,5

I4
13

Scirpus

americanus

C3

C4

C4

03

Scirpus

lacustris

05

05

C5

05

Scirpus

pallidus

Taraxacum

spp.

Trifolium

spp.

Trifolium

hybridum

Triglochin

maritima

Typha spp.

S5

05
14,5

04,5
13

03

04

S3

05

05

14
03,4

C4
13

ne orn ex i ca na
airoides

S5

13,4

Sc ir pu s palludosus

Sporobolus

I2

I2

spp.

Salix spp.

14
14

nasturtium-aquaticum

Sidalcea

12

airoides

Ra nu ncu Iu s spp.
Rorippa

3

S4,5

01,2
Il
01

11, 2 Sl

C2

S2

S2

02

01

03

02

S5

15

C2,3

04
01

Il

S2

01

C1

C5

S5

S5

13,4

04
C5

C5

S4.5

C5

I5

I
I-'
VI
00
I

�October 1976

-159JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-88-R-21
5

Work Plan No.
Job Title

Job No.

2

Investigations of Rails in Colorado

Period Covered:
Personnel:

Migratory Bird Investigations

April 1, 1975 to March 31, 1976

Herman J. Griese and Ronald A. Ryder, Colorado State University;
Clait E. Braun, Howard D. Funk, and Wildlife Conservation Officers,
Colorado Division of Wildlife; Charles R. Bryant, Monte Vista
National Wildlife Refuge.

ABSTRACT

Investigations concerning distribution, breeding densities, habitat utilization, and timing of migration of rails (Rallinae) in Colorado were initiated
in May, 1975. Study areas were selected in north central, southeastern, south
central, and northwestern Colorado. Methods for ascertaining rail presence
included responses to playbacks of tape recorded calls of rails, observations,
and trapping. Sora (Porzana carolina) were more widely distributed in western
Colorado than Virginia rails (Rallus 1. limicola). A black rail (Laterallus
i. jamaicensis) was recorded in southeastern Colorado. Virginia rails were
found in winter in sm~ll numbers in the South Platte and Arkansas River
Valleys. Soras were found in winter in the Arkansas River Valley in lower
numbers than Virginia rails. Breeding densities of rails were highest in
southeastern and south central Colorado, moderate in north central Colorado
and lowest in northwestern Colorado. Mean breeding densities for Virginia
rails and soras were 2.47 and 0.9 rails/ha respectively. Highest sora and
Virginia rail breeding densities were located in south central and southeastern Colorado, respectively. Rails were most frequently recorded in
cattail (Typha spp.) marshes with shallow water. Virginia rails appeared to
be restricted by distribution of cattail marshes in western Colorado. Spring
migration apparently peaked in mid-May with peak fall concentrations of
rails occurring in early to mid-September.
In 1975, 54 sora and 42 Virginia
rails were newly banded.

�-160-

RECOMMENDATIONS

1.

Study areas should continue to be censused with emphasis on migrational
periods.

2.

Intensive nest searches need to be conducted at regional study areas.

3.

State wide reconnaissance for rail occurrence should be continued with
emphasis on drainages not yet investigated.

4.

Microc1imatic data should be obtained from marshes with contrasting
densities of Virginia rails and soras.

5.

Methodology needs to be developed to ascertain sex of Virginia rails.

�-161-

INVESTIGATIONS OF RAILS IN COLORADO
Herman J. Griese

The family Rallidae is the largest and most diverse family in the order
Gruiformes. Members cf this family may be found throughout the world except
in polar regions. Several flightless members of the family occurring on
oceanic islands have become extinct in recent times due to the introduction
of non-native predators (Austin 1961).
North American rails of the subfamily Rallinae reported in Colorado are the
Virginia rail, sora, yellow rail (Coturnicops ~. noveboracensis) and black
rail. The Virginia rail commonly nests and occasionally remains throughout
winter, soras are also common nesting birds in suitable habitats, while
black and yellow rails are rare summer residents (Bailey and Niedrach 1965).
Current regulations (1975-1976) allow an aggregate bag limit of 25 Virginia
rails and soras daily in the Central Flyway portion of Colorado, while in
the Pacific Flyway portion of the state, hunting of rails is not permitted.
Thus hunting regulations in Colorado differ markedly on either side of the
Continental Divide. Holliman (1976) indicated the harvest: of rails in Colorado was insignificant.
Previous investigations of Virginia and king rails and soras have been
conducted primarily in the upper Mississippi River Valley and in the New
England states. To improve knowledge of rails seasonally resident in Colorado and to provide a basis for sound management, this study was initiated
in 1975. In this report, unless otherwise specified, "rails" refers to soras
and Virginia rails.
P. S. OBJECTIVES
To obtain data necessary for development of a management plan for rails
breeding in and migrating through Colorado, including: distribution and
breeding densities in four different regions, botanical and physical characteristics of habitats utilized, and documentation of timing of spring and
fall migration.
Further, hypotheses have been developed that:
a.

Distribution of rails in Colorado is determined by the distribution
of cattail (11Eb~spp.) marshes.

b.

Densities of breeding rails differ significantly by species between
occupied habitats in the eastern plains and the high mountain valleys.

c.

Timing of spring and fall migration differs significantly between
areas east of the mountains and intermountain valleys.

�_162-

SEGMENT OBJECTIVES
1.

Review literature concerning records of rail occurrence in Colorado
and procedures for censusing rails.

lb. Circulate observation questionnaires to field personnel of the Colorado
Division of Wildlife, selected University and privately employed wildlife
scientists and amateur ornithologists.
lc. Investigate selected habitats for rail occurrence in Colorado.
2.

Select study areas approximating at least 20 hectares in size that
represent habitats typical of those on the eastern plains and in high
mountain valleys.

3.

Periodically census each study area through use of flush counts and tape
recorded ~alls from early May through mid-October.

4.

Describe vegetatively (through cover mapping) and measure site characteristics (such as water depth, pH, nature of substrate, etc.) each area
studied.

5.

Compile data, analyze results and prepare progress report.
REVIEW OF LITERATURE

Numerous accounts of observations of rails or occasional nests occur in the
literature.
Observations and known distribution of rails prior to the mid1950's have been summarized and reported by the American Ornithologists
Union (1957). Observations and known distribution of rails in Colorado prior
to the mid-1960's have been summarized and reported by Bailey and Niedrach
(1965). Audubon sponsored Christmas bird counts in Colorado (1961-1975)
listed in American Birds contain several sightings of wintering rails.
Life history and behavior of all North American rails have been described
by Bent (1926). Walkinshaw (1937, 1940). Pospichal (1952), Tanner (1953),
and Kaufmann (1971) have described sora and Virginia rail ecology. King rails
(Rallus~. elegans) have been studied by Meanley (1969). Batemann (1965)
and Roth et al. (1972) have described the ecology of clapper rails (Rallus
longirostrus saturatus). Habitat utilized by rails has been described by
Billard (1947), Tanner (1953), Horak (1964), Meanley (1969), Kaufmann (1971),
and Andrews (1973). Descriptions of foods utilized have been presented by
Horak (1964) and Meanley (1965, 1969). Hunting mortality of rails has been
summarized by MacDonald and Martin (1971), while Artmann and Martin
(1975)
and Van Velzen and Kreitzer (1975) documented sources of non-hunting mortality.
Boeker (1954), Smith (1955), and Post and Enders (1970) used responses of
rails to est~mate densities.
Tomlinson and Todd (1973) and Glahn (1974)
described census techniques using tape recorded calls. Pospichal (1952),
Tanner (1953), Batemann (1965), and Andrews (1973) compared trapping techniques
for rails. Literature of rails has been compiled by Reeves (1975) and
partially reviewed by Smyth (1933), Glahn (1968), Wilbur (1974), and Wilbur
and Tomlinson (1976).

�-163-

METHODS AND MATERIALS
Four primary study areas, one in each of four regions of Colorado, were
selected for their similarity to regional wetland types, size, presence of
rails, and accessibility. Study areas were censused by playback of tape
recorded calls of rails at 25 to 28 listening sites per area. Listening
sites were 100 to 130 m apart. Locations of responding rails were approximated and recorded on field maps (Appendix A). All observations of rails
and nest sites located were also recorded on the field maps.
Presence of rails was determined primarily by playing tape recorded advertising calls of sora and Virginia rails on a portable Nore1co cassette
recorder. Playbacks per listening site numbered 6 to 25 calls in aggregate,
equally distributed among sora and Virginia rail calls. Calls were played
at intervals of ~pproximate1y one minute. Calls of black and yellow rail
were played on occasion. Flushing, track searches and trapping were also
used in ascertaining rail presence with most wetlands ecnountered in traveling within the state being examined. A questionnaire relating to distribution, habitat, and hunting was developed and distributed to all Colorado
Division of Wildlife field personnel (Appendix B).
Dominant vegetative cover was sketched on field maps of each area (scientific
plant names follow Harrington 1964). Water depths and fluctuations were
documented. Water was chemically analyzed for acidity, alkalinity, hardness,
and pH using a Hach Chemical Company Water Ecology Kit, Model AL-36B.
To document timing of migration, relative densities of rails were estimated
for each visit to a study area between May and October. Study areas were
visited until presence of rails was no longer documented. Several marshes
were visited throughout the winter to ascertain presence or absence of
wintering rails.
Trapping was conducted to aid in ascertaining composition of rail populations.
Four to six funnel box traps, 25 x 31 x 61 em of 1.4 em welded wire mesh with
a 25 x 2 em funnel opening at one end, were used in trapping. Drift fences
3 to 15 m long of 2.5 em chicken wire, 30 cm in height were used in conjunction with the traps.
DESCRIPTION OF STUDY AREAS
Rails were censused at marshes in four regions of the state. Areas studied
were: (1) Lower Latham Reservoir, South Platte River drainage, Weld County,
north central Colorado; (2) John Martin Reservoir, Arkansas River drainage,
Bent County, southeastern Colorado; (3) Monte Vista National Wildlife Refuge,
Rio Grande River drainage, Rio Grande County, south central Colorado; and
(4) the Cary Ranch, Yampa River drainage, Routt County, northwestern Colorado.
The Lower Latham Reservoir study area (15.6 ha in size) occurs at an elevation of 1,422 m on the south shore of a "Type 5" wetland as described by
Hopper (1968). Narrow-leafed cattail (Typha fuLgustifolia) covers about 63

�-164-

percent of the study area. Cover types also include mixtures of spike rush
(Eleocharis spp.), rushes (Juncus spp.), and common three-square (Scirpus
americanus) (12 percent), bullrush (Scirpus validus) (6 percent), and
seasonally flooded grasses (Gramineae) (2 percent). Duckweed (Lemna minor)
was locally abundant in the remaining open water, 16 percent of~study
area. Average water depth was 10 cm with seasonal fluctuations greater than
60 em.
The study area at John Martin Reservoir is a "Type 3" wetland at 1,171 m elevation. Narrow-leaved cattail (89 percent), spike rush-rush-common three-square
mixture (4 percent), and open water (6 percent) are the major components of
this 15.1 ha study are.a. Duckweed and watercress (Rorippa nasturtiumaquaticum) cover much of the water surface. Average water depth is 7 cm,
with seasonal fluctuations of 10 cm.
The Monte Vista study area is a "Type I-A" wetland at 2,326 m elevation. A
mixture of rush-spike rush-alkali bullrush (Scirpus paludosus) (46 percent),
broad-leafed cattail (Typha latifolia) (31 percent) and open water (21 percent) dominate the 14.3 ha area. Water depths average 5 cm with 13 em fluctuations during the field season.
The Cary Ranch study area is composed of wetland "Types I-A, 3, and 4" at an
elevation of 1,914 m. The dominant cover types of this 12.3 ha area are
sedges (Carex spp ,) and grasses in periodically flooded fields (37 percent),
broad-leafed cattail (28 percent), open water much of which is covered by
pondweed
(Potamageton spp.) and duckweed (15 percent), rush-spikerush (11
percent), and bullrush (9 percent). Average water depth is 31 cm with greater
than 150 cm seasonal fluctuations.

RESULTS AND DISCUSSION
Distribution
Observations of rails reported by Bailey and Niedrach (1965) are concentrated
along the foothills between Denver and Fort Collins, probably because of
access and behavior patterns of observers. Elsewhere in the state, records
are scarce. Prior to the investigation reported here, only general information on rails was available. Bailey and Niedrach summarized the available
information and reported that Virginia rails were common breeding birds in
the "upper sonoran zone" (1,070-1,680 m), while soras were common breeding
birds on the plains and in the mountains as high as 3,050 m. Their summary
indicates that yellow and black rails have been recorded from the state only
one and two times, respectively.
Data obtained from the initial field season of state wide investigations,
responses to the questionnaires, and previous published records were compiled
to prepare a composite map of the summer distribution of soras and Virginia
rails in Colorado (Fig. 1). Presence of rails anywhere in a river drainage
system was assumed to indicate that other favorable habitats in the drainage
support rails. Discontinuous patterns of favorable habitat probably restrict
distribution more than is indicated in Fig. 1.

�COLORADO
MOFFAT

ItOUTT

I
I-'
0\
\JI

cHtreNNI!

I

DOLORES

MONTEZUMA

~

I

VZJ Virginia Rail
Fig. 1.

Preliminary breeding distribution

~
of rails in Colorado, 1975.

Sora

�-166-

Factors limiting distribution of rails appear to be presence of water between
April and July, sufficient emergent plant cover, and elevation. Soras have
been recorded nesting at elevations up to 3,231 m (Bailey and Niedrach 1965),
and Virginias to 2,325 m (this study). Frequency of occurrence of Virginias
and soras in investigated marshes in eastern Colorado in 1975 was 82 and 47
percent, respectively. Investigations in western Colorado show respective
frequencies of 26 and 50 percent. Soras appear to be more tolerant of higher
elevations, or wetlands associated with high altitudinal climates than
Virginia rails.
Distribution of the remalnlng reported species of rails in Colorado is not
indicated in Fig. 1. Yellow and black rails have been recorded near Denver
(Bailey and Niedrach 1965), while black rails were recorded during this study
at John Martin Reservoir. Although king rails have not been recorded officially in Colorado, a possible sighting of this species was made during field
investigations at Bonny Reservoir in Yuma County on 24 May. King rails breed
commonly at Cheyenne Bottoms in central Kansas, approximately 350 km east of
Bonny and John Martin Reservoirs (M. Schwilling, pers. comm.).
Distribution of wintering rails (time interval of 1 November-31 March) was
restricted primarily to the South Platte and Arkansas River drainages in
eastern Colorado. Virginia rails were the most abundant rail wintering in
Colorado. Soras have been reported near Denver on Audubon sponsored Christmas
counts in American Birds 1961-1975. During this investigation, soras were
found wintering with Virginia rails in the Arkansas River Valley. In western
Colorado rails have been reported in winter on Christmas counts at Grand
Junction and the Monte Vista National Wildlife Refuge. Wildlife Conservation
Officer responses indicate wintering rails near Hotchkiss and in North Park.
Rails reported from these areas in winter were only found in ice-free marshes.
Breeding Densities
Glahn (1974) has shown that responses to playbacks of tape-recorded rail
calls produce an index to breeding densities of rails during April and May
(1.6 sora responses/territory, 1.2 Virginia rail responses/territory).
Kaufmann (1971) indicated the breeding period of rails is characterized by
advertising calls with such calls being given before, during, and after
nesting. By analyzing responses of rails to playbacks of tape recorded calls
between May and July, indices of breeding densities of rails in Colorado were
obtained (Table 1).
Results presented in Table 1 indicate that densities of rails decrease from
south to north in Colorado. Densities of Virginia rails were highest in the
eastern plains, while densities of soras were highest in the inter-mountain valleys and west of the Continental Divide. Statewide densities for Virginia rails
(2.47 + 0.76 rails/ha, a = 0.05) were higher than soras (0.91 + 0.35 rails/
ha, a ~ 0.05). These findings would indicate that soras prefer~ed higher elevations
than did Virginia rails with selection against marshes in the eaStern plains.
These data coincide with those utilized in the map of rail distribution in
Colorado.

�· ".

Table 1.

Breeding

densities of rails on four study areas in Colorado, 1975.
Areas
Monte Vista National
Wildlife Refuge

John Martin
Reservoir

Lower
Latham Reservoir

Cary Ranch

Dates
20 June - 3 July

31 May - 25 June

1 June - 23 June

21 June-4July

x

1.89

4.06

3.10

0.09

S.D. ]j

0.83

1.06

0.25

0.16

x

2.14

0.13

0.75

0.94

S.D.

0.36

0.08

0.32

0.41

3

4

4

3

x

4.04

4.19

3.85

1.03

S.D.

1.18

1.14

0.57

0.57

Virginia Rail/ha 1/
I

t-'

0\

Sora/ha

To tal Rails /ha

3/

n-

1/0ne rail/ha
l/S.D.

=

0.4 rails/acre.

= Standard

deviation.

3/
- n = Number of censuses taken.

.....•
I

�-168-

Study areas in northern Colorado, having lowest densities of rails, also
had the largest fluctuations in water levels. The Cary Ranch was subject
to seasonal flooding of the Yampa River, while Lower Latham Reservoir
received a great influx of water during periods of heavy rainfall in May.
In both study areas, water depths increased during the period of nest
initiation. Billard (1947) found highest densities of rail nests in
marshes with slowly decreasing water levels. Therefore, in areas of
fluctuating water levels, breeding densities may appear unstable due to
movements to areas with shallow water.
Differences in vegetative composition between northern and southern study
areas did not appear to be correlated with differences in total densities.
Differences in vegetative composition between plains and mountain study
areas may indicate a species preference for certain vegetative cover.
Cattails covered from 63 to 91 percent of the plains study areas where
Virginia rails had highest densities. Inter-mountain study areas with high
sora densities were 29 to 34 percent covered with cattails. It is possible
that vegetation composition of marshes may influence breeding densities of
rails in eastern and western Colorado.
Total area suitable for rails in different regions of Colorado is dependent
upon quantity and quality of wetlands. Hectares of suitable wetlands for
breeding rails (Table 2) were estimated from data presented by Hopper (1968).
Wetlands Types I-A, 3, and 4 were assumed to be highly suitable. Portions
of Type 5 wetlands and small portions of ditch and canal mileage were also
considered. Estimates for the Yampa Valley were obtained during the course
of this investigation.

Table 2. Estimated potential rail breeding habitat in four regions of
Colorado (approximated from Hopper 1968).

Hectares

Arkansas River
Valley

San Luis
Valley

Region
South Platte
River Valley

400

4,300

2,000

Yampa River 1./
Valley

500

l/ Hectares estimated during this study.
Considering recorded breeding densities and available habitat, the highest
breeding potential for rails in Colorado occurs in the San Luis Valley (Rio
Grande River drainage). The South Platte, Arkansas and Yampa River Valleys
follow in order of potential. Vegetative composition was not considered and
may influence the final analysis.

�-169-

Habitat Utilization
Periods of importance when considering habitat utilization are breeding,
nesting, brood development, and fall migration.
In all periods, quality
of cover is of most importance. During brood development and fall migration, food quantity is probably also important.
Breeding
Responses during the breeding period, May through June, at the Monte Vista
National Wildlife Refuge study area, indicated a preference for cattail
cover types by both species of rails. Dominant vegetation at this site was
34 percent cattail, 45 percent spikerush-rush, 6 percent alkali bullrush,
and &lt;1 percent bullrush. Soras responding in cattails comprised 76 percent
of total soras recorded, while 87 percent of the Virginia rail responses
were in cattails.. The remaining calls came from fine (grass like) vegetation.
At the Cary Ranch, 20 percent of all rail responses during the breeding
season were from areas dominated by flooded grasses and sedges. Almost all
remaining rail responses in the state during this period were recorded in
cattail or cattail-bullrush cover.
Water depths in areas of responses were not documented.
appeared to prefer deeper water.

However, soras

Nesting
Intensive nest searches were not conducted during 1975, and only 17 nests
were located. Of this total, one nest was of a sora, while four were of
Virginia rails, with the remainder being unidentified. All nests were
located in cattails, except one Virginia rail nest found in rushes at Monte
Vista National Wildlife Refuge. This nest was apparently destroyed by avian
predators.
Nesting habitat of sora and Virginia rails has been studied in the midwest
and northeast United States. Kaufmann (1971) summarized these data and
Andrews (1973) presented additional data. These studies found that 71 percent of all Virginia rail nests (162) and 60 percent of all sora nests (220)
were in cover of robust vegetation. Thirty-eight percent of all Virginia
rail nests and 36 percent of all sora nests were in robust vegetation
of cattails. Fine vegetation produced 18 percent of the Virginia rail
nests, and 32 percent of the sora nests. Twenty-one percent of all sora
nests were in blue-joint (Calamagrostis canadensis) in these studies.
Eight percent of all nests were round in mixed robust and fine vegetation.
Kaufmann (1971) suggested that mixed vegetative types may be under estimated
in preference to dominant vegetation.
In Colorado, Glahn (1974) found 9 Virginia rail nests and 5 sora nests in
cattail dominated vegetation. Bailey and Niedrach (1965) indicated that
rails nest in robust and fine vegetation with nests of soras being frequently
located in fine vegetation. Glahn also indicated that the edge vegetation
of territory borders may be important in determining rail nest placement.

�-170-

Nests found in Colorado were in mean water depths of 12.8 and 7.8 em for
soras and Virginia rails, respectively. Water depths at nests in studies
prior to 1963 are summarized by Horak (1964). He reported that sora and
Virginia rails chose vaciable water depths in which to construct nests.
Soras appeared to prefer water deeper than Virginia rails; however, ranges
overlap. Nests of soras were found in &lt; 15 to 48 em of water. Virginia
rail nests were in &lt;15 to 38 em of water.

Brood Period
Responses of chicks during the brood development period appeared to be
totally from cattail vegetation. Andrews (1973) trapped most rails during
July in cattail dominated vegetation.
In this study immature rails about
one month of age or less were trapped in cattail (7 Virginia rails and 10
soras). Cattails apparently afford necessary cover and food for chicks.
Migration
Bent (1926) reported that in late summer rails moved to more stable wet
marshes prior to migration. On all study areas, movement of rails corresponding to receding shore lines was noted. On the Monte Vista National
Wildlife Refuge rails moved into alkali bullrush, while on other areas
studied, cattails were present on all stable marshes.
During fall migration, rails, primarily sora, were flushed and trapped in
alkali bullrush when sufficient water depths were present. At this time,
numbers of responding rails in cattails also increased. Water depths
preferred during migration varied from none (muddy) to 60 em in areas of
responses and sightings. Soras were most often flushed in shallow water.
Andrews (1973) indicated soras pref erred reflooded areas of nodding smartweed (Polygonum lapthifolium), Walter millet (Echinichola walteri), and
redroot cyperus (Cyperus erythrorhizos) during fall concentrations, probably because of the abundance of available food.

Water Chemistry
Chemical characteristics of water on the four study areas differed only
slightly in degree of alkalinity. Correlations of chemical characteristics
and rail presence have not been completed. It is believed that vegetation
is affected by soil and water characteristics which in turn, may affect
rail densities.
Timing of Migration
Investigations to date indicate that migration of Virginia rails into Colorado begins in mid-April. During this study soras were recorded in large
numbers on 2 May at Jonn Martin Reservoir. In western Colorado, migration

�-171-

is suspected to begin in late April. Bailey and Niedrach (1965) indicated
that spring arrival of Virginia rails in Colorado was in late April or
early May. They also found soras reached north central Colorado as early
as 3 May and that soras nested in northwestern Colorado at a time indicating
a mid-to late April arrival. In previous extensive rail studies, spring
migration was documented to occur from mid-April to early May. Pospichal
(1952) found rail migration in spring to be controlled by the progression
of spring.
Peak concentrations of Virginia rails during spring migration occurred in
mid-May at Lower Latham Reservoir and late May at John Martin Reservoir.
However, poor weather during the mid-May census at John Martin may have
biased results.
Peaks in spring concentrations were not recorded for areas in western
Colorado as the first censuses occurred in June. It is of interest that
Virginia rails were not recorded at Cary Ranch until 4 July.
Peak spring concentrations of soras in southeastern Colorado occurred in
early May, while in north central Colorado peak concentrations occurred
in mid-May. In northwestern Colorado, soras apparently peaked in late May.
Lack of census data prior to 1 May prevents a clear understanding of spring
migration in 1975.
At Lower Latham Reservoir highest numbers of Virginia rails (4.17 rails/ha)
and soras (2.50 rails/ha) occurred in mid-August and mid-September, respectively. At John Martin Reservoir, both species peaked in mid-September
(4.04 Virginia rails and 3.25 soras/ha, respectively). At Monte Vista
National Wildlife Refuge highest numbers occurred prior to mid-August, probably the result of drying of the study area. A similar situation occurred
at Cary Ranch in mid-August.
In North Park, Jackson County, in north central
Colorado, soras appeared to peak in mid-August (Wagner pers. comm.)
Trapping
Trapping was conducted to aid in ascertaining composition of rail populations. Trapping results during the 1975 field season are presented in Table 3.
Ninety-six rails were newly banded, of which 23 were adults (14 Virginias
and 9 soras), while 73 were immatures (28 Virginias and 45 soras). Trapping
success was .064 rails trapped per trap hour for 1,617 trap hours. Success
was highest when traps were checked at 3 to 6 hour intervals, and moved
every third day. Predation by raccoons (Procyon lotor) occurred at three
traps causing a loss of at least three rails.
Table 3 indicates that immature rails were 82, 73, and 64 percent of banded
rails during August, September and November, respectively. Of soras banded,
immatures accounted for 86, 82, and 50 percent respectively. Immature
Virginia rails accounted for 73, 60, and 66 percent respectively. Trapping
time and methods may introduce biases causing production estimates to be
inappropriate.

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Table 3.

Distribution of bandings of rails, by species, Colorado, 1975.

Number Banded
Aug.

Soras
Sept. Nov.

Totals

Aug.

Virginia Rails
Sept. Nov. Totals

Milk Creek
Adult

0

0

1

1

Immature

16

16

1

1

Adult

4

4

0

0

Immature

4

4

0

0

Adult

0

0

1

1

Immature

6

6

10

10

Walden Reservoir

Monte Vista NWR

Wellington Mgmt. Area
Adult

0

0

1

1

Immature

2

2

0

0

Adult

2

2

2

2

Immature

10

10

3

3

Adult

2

2

3

3

Immature

6

6

6

6

Fossil Creek Reservoir

Lower Latham Reservoir

John Martin Reservoir
Adult

0

1

1

3

3

6

Immature

0

1

1

2

6

8

Totals
Adult

4

4

1

9

5

6

3

14

Immature

26

18

1

45

13

9

6

28

�-173-

In an effort to determine the relation between induced rail responses and
drive trapping success, two .5 ha marshes at Wellington Wildlife Management area in north central Colorado were censused and immediately drive
trapped in September. Orgffilizeddrives with drivers one meter apart
resulted in 3 rails being trapped or flushed. Eight rails had previously
responded to tape-recorded rail calls. Thus, 37 percent of the rails
present were accounted for by drive trapping under the marsh and seasonal
conditions in effect. This value could be lower if more than 8 rails, as
determined by calls, were present.

LITERATURE CITED

American Ornithologist's Union. 1957. Check-list of North American birds.
5th Ed. Lord Baltimore Press, Inc., Baltimore. 691 pp.
Andrews, D. A. 1973. Habitat utilization by soras, Virginia rails, and
king rails near southwestern Lake Erie. M. S. Thesis, Ohio State Univ.,
Columbus. 112 pp.
Artmann, J. W., and E. M. Martin. 1975. Incidence of ingested lead shot
in sora rails. J. Wildl. Manage. 39(3):514-519.
Austin, O. L.

1961.

Birds of the world.

Golden Press, New York.

Bailey, A. M., and R. J. Niedrach. 1965. Birds of Colorado.
Nat. Hist., Denver. Vol. I. 454 pp.

316 pp.

Denver Mus.

Batemann , H. A. 1965. Clapper rail (Rallu~ longirostrus) studies on
Grand Terre Island, Jefferson Parish, Louisiana. M. S. Thesis,
Louisiana State Univ., Baton Rouge. 145 pp.
Bent, A. C. 1926. Life histories of North American marsh birds.
Natl. Mus. Bull. 135. 490 pp.

U. S.

Billard, R. S. 1947. An ecological study of the Virginia rail (Rallus
limicola) and sora (Porzana carolina) in some Connecticut swamps.
M. S. Thesis, Iowa State Univ., Ames. 84 pp.
Boeker, H. M. 1954. A census of populations of the Wilson's snipe ffild
sora rail in the Yampa River Valley, Colorado. Condor 56(2):105-106.
Glahn, J. F. 1968. Literature related to the Virginia and sora rails.
Unpubl. Manuscript, Colo. State Univ., Fort Collins. 30 pp.
1974. Study of breeding rails with recorded calls in north
central Colorado. Wilson Bull. 86(3):206-214.
Harrington, H. D. 1964. Manual of the p1&amp;ltS of Colorado.
Swallow Press, Inc., Chicago. 666 pp.

2nd Ed.

The

�-174-

Holliman, D. C. (Chairman). 1976. Rails (Rallidae) in Glen C. Sanderson,
ed. Management of migratory shore and upland ga;e birds in North
America. Int. Assoc. Game, Fish and Cons. Comms., Washington, D. C.
(In press).
Hopper, R. M. 1968. Wetlands of Colorado.
Dept. Tech. Publ. No. 22. 88 pp.

Colorado Game, Fish and Parks

Horak, G. J. 1964. A comparative study of Virginia and sora rails with
emphasis on foods. M. S. Thesis, Iowa State Univ., Ames. 73 pp.
Kaufmann, G. W. 1971. Behavior and ecology of the sora, (Porzana carolina)
and Virginia rail (Rallus limicola). Ph. D. Thesis, Univ. Minn.,
Minneapolis.
114 pp.
MacDonald, D., and E. Martin. 1971. Trends in harvest of migratory game
birds other than waterfowl, 1964-65 to 1968-69. U. S. Fish and Wildl.
Servo Spec. Sci. Rept. Wildl. 142. 29 pp.
Meanley, B. 1965. Early fall food and habitat of the sora in the Patuxent
River Marsh, Maryland.
Chesapeake Sci. 6(4):226-227.
1969.
108 pp ,

Natural history of the king rail.

N. Amer. Fauna No. 67.

Pospichal, L. B. 1952. A field study of sora rail (Porzana carolina)
and Virginia raiJ. (Rallus .limicola) populations in central Minnesota.
M. S. Thesis, Univ. Minn., Minneapolis.
80 pp.
Post, W., and F. Enders. 1970. Notes on a salt marsh Virginia rail population. Kfn gbLrd 29(2) :61-67.
Reeves, H. M. (Compiler). 1975. A contribution to an annotated bibliography
of North American cranes, rails, woodcock, snipe, doves, and pigeons.
U. S. Fish and Wildl. Servo Publ. 240999. 527 pp.
Roth, R. R., J. D. Newsom, T. Joanen, and L. L. McNease. 1972. The daily
and seasonal behavior patterns of the clapper rail (Rallus longirostris)
in Louisiana coastal marshes. Proc. S. E. Assoc. State Game and Fish
Comms. 26:136-159.
Smith, A. B. 1955. Sora rail populations in Alberta, 1953-54. In
Investigations of woodcock, snipe, and rails in 1954. u. S. Fish and
Wildl. Servo Sci. P£pt. Wildl. 28. pp. 59-62.
Smyth, J. A. 1933. The literature of the Virginia rail.
Cornell Univ., Ithaca. 122 pp.

M. S. Thesis,

Tanner, W. D. 1953. Ecology of the Virginia and king rails and sora in
Clay County, Iowa. Ph. D. Thesis, Iowa State Univ., Ames. 154 pp.

�-175-

Tomlinson, R. E., and R. L. Todd. 1973. Distribution of two western
clapper rail races as determined by responses to taped calls. Condor
75 (2) :177-183.
Van Velzen, A., and J. F. Kreitzer.
1975. The toxicity of p,p'-DDT
the clapper rail. J. Wildl. Manage. 39(2):305-309.
Walkinshaw,

L. H.

1940.

1937.

The Virginia

rail in Michigan.

Summer life of the sora rail.

to

Auk 54(4):464-475.

Auk 57(2):153-168.

Wilbur, S. R. 1974. The literature of the California black rail. U. S.
Fish and Wildl. Servo Spec. Sci. Rept. Wildl. No. 179. 17 pp.
__

, and R. E. Tomlinson.
1976. The literature of the Western clapper
rails. U. S. Fish and Wi1d1. Servo Spec. Sci. Rept. Wi1d1. No. 194.
31 pp.

Prepared

BY_--,L=..:==.:.."I-4J~/--"t0~::~~_
..t:-:..;,i~=-"_/_'

_

Herman J. iG'riese

Ill'

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Approved BY_-"u:.~V'a::l:·~~_·
-="-:;:..&gt;-_'--Io:(.:::;5_-t..,~~
__
)
C1ait E. Braun

_

�-176APPENDIX
MONTE VISTA NATIONAL

A

WILDLIFE

REFUGE

STUDY AREA

r»

"/~-'

--'-----()

COMMENTS:
DATE:
END:

TIME BEGIN:
TEMP. :
WIND:
SKY:
%
l.

6.

11.

16.

21.

2.

7.

12.

17.

22.

3.

8.

13.

18.

23.

4.

9.

14.

19.

24.

5.

10.

15.

20.

25.

�-177APPENDIX B

STUDIES OF RAILS IN COLORADO
1.

District

_

2.

Do rails occur to your knowledge in your present district?
If no, answer question No. 5 only.

3.

If rails are known or suspected to occur in your district, please list all
locations.

Yes

No

Generalizations can be made:

_

4. What vegetation types occur where you hear or see rails?

_

S.

Please list major cattail marshes in your district not mentioned above

6.

What time(s) of the year do you most commonly hear or see rails?

7.

Have you encountered rails during the 1 November through 31 March period?
Yes

8.

No

If yes, where?

Have you ever encountered
any rail hunters in your district? Yes
,
If yes, where?

_

_

No
_

��-179-

October 1976

JOB PROGRESS REPORT

State of

COLORADO
---------------------------W-88-R-21

Project No.

Migratory

6

Work Plan No.

Job No.

Job Title

Migratory Bird Publications

Period Covered:

April 1, 1975 through March 31, 1976

Personnel:

Bird Investigations
1

_

Richard M. Ropper, Michael R. Szymczak, John F. Corey, and Jack
R. Grieb.

ABSTRACT
Publications
follows:

planned for and completed under this job for Segment 21 are as

1.

Szymczak, M. R. 1975. Canada goose restoration along the foothills of
Colorado. State of Colorado, Division of Wildlife Technical Bulletin
Number 31. 'September.
64 pp.

2.

Hopper, R. M., A. D. Geis, J. R. Grieb, and L. Nelson, Jr. 1975.
Experimental duck hunting seasons, San Luis Valley, Colorado, 19631970. Wildlife Monographs No. 46. November.
68 pp.

Also completed but not yet published is a special report as follows:
1.

Szymczak, M. R., and J. F. Corey. 1976. Construction and use of the
Salt Plains duck trap in Colorado. Colorado Division of Wildlife,
Division Report No.6.
In Press.

Prepared by__ --=_··_/_~J_I
,-=,h::::/f:",'i.::;;.J.4":,I,_, __ L_,_.~_·~_~--,:;,:::,,?_:~-=-.
-.,.
Howard D. Funk
Section Chief,
Small Game Research
[,-:-"'-::'U:",'

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                  <text>-1JOB FINAl,

State of

W-4l-R-26

\-lorkPlan No.

1

Bighorn Sheep and Mountain Goat Investigations
Job No.

Evaluation of the Nutritional

Period Covered:
Personnel:

REPORT

Colorado

Project No.

Job Title

January, 1977

16

.---------------------------Requirements

of Bighorn Sheep

June 1, 1971 through May 31, 1976

Robert E. Keiss and Gene G. Schoonveld

ABSTRACT
An evaluation of the nutritional status of bighorn sheep on existing ranges
was undertaken to measure the adequacy or inadequacy of these ranges to meet
the nutritional requirements of bighorn sheep at all seasons of the year.
The general study outline consisted of selecting study areas, developing
analytical procedures for the chemical partitioning of forage and fecal
samples, evaluating micro techniques for plant fragment composition of
fecal material, comparative digestibility trials, comparative forage quality
analysis at all seasons of the year, and finally bighorn sheep tissue analysis
for trace mineral levels.

��-3-

EVALUATION

OF THE NUTRITIONAL

REQUIREMENTS

OF BIGHORN SHEEP

Robert E. Keiss

P. S. OBJECTIVE
To develop methods: (a) for the evaluation of nutritional requirements
of bighorn sheep under natural conditions, and (b) to measure the adequacy
of selected ranges to meet these requirements.

METHODS AND MATERIALS
Range forage, unlike most cultivated crop feeds is harvested by the grazing
animal in an assortment of different plant species and plant parts as determined by the selectivity of the animal. In order to evaluate the nutritive
intake by these grazing animals it is necessary to determine the digestibility
of the ingested plant parts and thereby develop a useful index to the nutritional quality of range herbage available to game animals in any given area.
There have been a number of studies devoted to the digestibility of the
nutrients in range forage by grazing animals. These free ranging animals
need protein for growth and repair of tissues, fats and carbohydrates for
production of heat and energy, vitamins for important physiological processes,
and minerals for general body function and bone building.
It is not feasible to make recommendations on supplementing animals until
the specific nutrient requirement of the animal, as well as the nutrient
content of the range forage is known. If any nutritive discrepancy exists
between what is required by the animal and what is present in the forage,
then it would be necessary to investigate methods to supplement the nutrient
in order to give the animal a balanced ration.
With the above general considerations in mind a study was designed to examine
the nutritive quality of range forage available to bighorn sheep. In order
to make comparisons between different bighorn sheep ranges and ass.eas
differences between animal vitality, three study areas were selected and
all comparisons made to include the yearly cycle of forage growth and maturity.
The general study outline then consisted of selecting study areas, developing
analytical procedures for the chemical partitioning of forage and fecal
samples, using and testing microelement techniques for plant fragment comr
position of fecal material, comparative forage digestibility trials, comr
parative forage quality analysis on the study areas at various seasons of
the year and finally bighorn sheep tissue analysis for trace minerals.
These different categories will be discussed separately in this report by
first outlining the methods and materials, followed by an assessment of
the results of these comparisons.

�-4DESCRIPTION OF STUDY AREAS
Three bighorn sheep ranges were selected for nutritional investigations.
The study areas were selected on the basis of their differing quality
of bighorn sheep herd success based upon past history of population
trends and estimated bighorn sheep numbers. Differing types of vegetative
cover as well as year-around accessibility were also considered (Fig. 1).
Pikes Peak study area, located in El Paso and Teller counties, Colorado
is both alpine and sub-alpine (10,200-14,100 ft) bighorn sheep range.
This area experienced a major die-off during 1953-54 and the population
has experienced a steady increase which is somewhat associated with
mineral supplementation which was carried out by Frank Colley of the
Colorado Division of Wildlife.
Buffalo Peaks study area, located in Park and Chaffee counties, Colorado
is primarily alpine (11,600-12,900 ft) bighorn sheep range. Quality of
this range is considered marginal as the bighorn sheep population had been
declining for several years. This area is not too accessible during
winter months as the snow is deep and over-snow vehicles are required.
Trickle Mountain study area, located in Saguache County, Colorado is a
low altitude (8,400-12,000 ft) sheep range primarily of a pinyon-juniper
type. Quality of this range is considered good as the bighorn sheep
populations in this area have steadily increased over the past years and
many animals are trapped from the area for transplant purposes.

Forage and Fecal Analytical Procedures
In the early years chemists attempted to define the nutritive quality of
feeds (Thaer 1809; Boussingault 1843; Henneberg and Stohmann 1860) and
their chemical methods were ultimately adopted and developed into the
total digestible nutrient (TDN) system of feed analysis as proposed by
Henry and Morrison (1910). The Weende method of partitioning forages is
based upon the TDN system (crude protein, crude fat, crude fiber, and
nitrogen free extract) but has several major flaws and assumptions which
are very questionable by today's standards of feed analysis.
Other modifications and improvements have been made by a number of workers
in recent years (Crampton and Maynard 1938; Van Soest 1964; Colburn and
Evans 1967; Van Soest and Wine 1967). Fonnesbeck and Harris (1970a, 1970b,
1971) and Harris (1970) have developed a procedure for examining the cell
wall constituents of all types of feeds which have been adopted by our
laboratory and were used throughout this experiment. The plant cell is
baSically separated into two main components, the cell walls and the cell
contents. The cell contents include the majority of the nutritive matter
that is digested by enzymatic action within the digestive system and subsequently absorbed and utilized by the animal (protein, ether extract,
soluble ash, and soluble carbohydrate).
Cell walls include both partially
nutritive matter and non-nutritive matter (lignin, acid insoluble ash,
cellulose and hemicellulose).
Partially nutritive matter include cell wall
carbohydrates, cellulose and hemicellulose.
These chemically complex
compounds are partially broken down by the action of bacterial enzymes

�COLORADO
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I

Pikes Peak
Study Area
MONTROSE
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o

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Trickle Mountain
Study Area

I)OLOIl£S

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Fig. 1.
Location of three study areas (Pikes Peak, Buffalo Peaks,
evaluation
of nutritive
quality
of forage for bighorn sheep.

and Trickle

Mountain)

selected

for

�-6-

within the gastro-intestinal tract and the total digestibility of the
cellulose and hemicellulose can be combined and reported as holocellulose as the differences for most animals is not significantly
different to report separately. The non-nutritive matter within the
cell walls includes the lignin and acid insoluble ash. These constituents
have no known nutritive value.
The system used for analysis of the nutritive value of feeds used in this
experiment requires the determinations of protein, ether extract, ash,
cell walls, cellulose, hemicellulose, lignin, acid insoluble ash, soluble
ash, and soluble carbohydrate.
Mineral analysis for calcium, phosphorus, magnesium, manganese, zinc,
copper, iron, potassium were made on selected forages as a part of this
study.

Protein
The micro-Kjeldahl method for the determination of crude protein (total
nitrogen x 6.25) was used. Two hundred fifty milligrams of plant material
was placed in a 100 ml Kjeldahl flask with 1 gm of catalytic mixture and
3 ml of concentrated H S0 . The material was predigested overnight then
2 4
placed upon a micro-Kjeldahl digester and heated until the sample was
completely clear. After cooling, the digested sample was transferred to
a micro-Kjeldahl distillation chamber.
Ten ml of 12 N.N OH is added and
the sample distilled with steam into a boric acid solutio~ containing 2
drops each of bromocresol green and 2,4, dinitro-phenol indicator solutions.
The boric acid solution was titrated with 0.05715 N. H S0 .
2 4
Calculation:
Percent Protein

ml titrant x 2

Reagents:
l.

2.
3.

4.
5.
6.

Catalytic Mixture
100 gm CUS04
Boric Acid
Bromocresol Green Indicator
2,4 Dinitro-phenol Indicator
l2N. NaOH
0.05715 N. H2S04

Ether Extract
A one gram (oven dry) sample is weighed into a previously tared, fat-free
extraction thimble. The thimble was placed in a Saxhlet fat extraction
apparatus and extracted for eight hours with anhydrous petroleum ether.
After the extraction period the thimbles were removed, allowed to dry overnight, cooled in a desiccator and weighed. The loss in weight of the sample
was calculated as percent ether extract on a dry weight basis.

Ash
A one gram (oven dry) sample was weighed into a previously tared crucible
and placed in a cold muffle furnace. The heat was allowed to build up

�-7-

slowly to 5500C and held overnight. The following day the heat was shnt
off, allowing the crucibles to cool. After the crucibles were cooled to
approximately 2000C they were transferred to a desiccator where they were
allowed to cool to room temperature and weighed. Percent ash was calculated
on a dry weight basis. The ash was then dissolved in 2 ml of concentrated
NH03 over a steam bath and quantitatively transferred to a 50 ml volumetric
flask, made to volume, and the solution kept for mineral analysis.

Cell Walls
A one gram (oven dry) sample was added to a 600 m1 Berzelius beaker and 75
m1 of citrate buffer and 1 ml decalin added. The beakers were placed on a
refluxing apparatus consisting of six individually controlled hot plates and
500 m1 round bottom water cooled flasks on top of the beakers. The samples
were brought to a boil and allowed to reflux for 30 minutes from the onset
of boiling. The beakers were removed, cooled and 25 m1 of pepsin solution
added. The beakers were placed in a water bath at 450C. Since we had no
means to keep the flasks in constant motion as recommended, we allowed the
digestion to proceed for approximately 48 hours with occasional stirring.
After the pepsin digestion the liquids were removed from the beakers and
the residue rinsed with hot water, followed by a thorough acetone washing.
After the acetone washing, 100 m1 of detergent solution was added and the
beakers refluxed for 60 minutes on the refluxing apparatus. The detergent
samples are then poured into a previously tared filter crucible while drawing
off the liquid with suction. The cell walls were then thoroughly washed
with large volumes of hot water, followed by thorough rinsing with acetone.
The filter crucibles were dried at 1050C overnight, placed in a desiccator
to cool, and weighed.
Calculation:
Weight of Cells Walls
Percent Cells Walls

Weight of Sample

X 100

Reagents:
1.

Citrate Buffer

Citric Acid
42 gm
Tween 40
10 gm
Water
1,000 ml
(Adjust pH to 3.0 with NH40H)

2.

Pepsin Solution
Pepsin
40 gm
Citric Acid
40 gm
Water
1,000 ml
(Make fresh as needed - 1 gm pepsin and 1 gm
citric acid per 25 m1 - which is required for
each sample)

3.

Detergent Solution
42 gm
Citric Acid
Dodecyl Sodium
20 gm
Sulfate
1,000 ml
Water
(Adjust to pH 3.0 with NH40H)

4.

Decalin (Decahydronaphthalene)

5.

Acetone

�-8-

Hemicellulose
The residue from the cell wall determination was removed from the filter
crucible and placed in the Berzelius beaker, taking great care to remove
all the material.
Seventy-five m1 of 8 percent H2S04 was added to the
beaker and they were refluxed on the refluxing apparatus for 1 hour.
After refluxing, each sample was washed back into the same filter crucible
from which it was removed, the residue washed in large volumes of hot
water followed by a thorough acetone rinse. The filter crucibles were
placed in the drying oven overnight, removed to a desiccator until cool
and weighed. Hemicellulose was computed as the loss of weight from the
cell wall residue.
Calculation:
% Hemicellulose

Wt. Cell Wall Residue Wt. Extracted Cell Wall Residue X 100
Sample Weight

1.

8% Sulfuric Acid (Specific gravity of 1.052 at 200C)

Lignin and Acid Insoluble Ash
The residue from the hemicellulose determination is digested with 72 percent
H2S04 by placing the filter crucible into a beaker containing the acid.
The acid is allowed to soak into the residue from the bottom through the
ft!°itteddisc so that the residue is wetted from the bottom up. The residue
was left in the acid for 3 hours after which the acid was filtered off,
followed by a thorough washing with large volumes of hot water. It is
important to remove all traces of acid from the residue and filter crucible.
The residue was then dried overnight, cooled in a desiccator and weighed.
The filter crucibles were then placed in the muffle furnace at SOOoC overnight, removed, cooled in a desiccator and weighed.
Calculation:
Wt. Residue Before Ashing - Wt. Residue After Ashing
% Lignin = ---------------------------------------------X 100
Sample Weigh t
% Acid Insoluble Ash

Wt. Residue After Ashing - Wt. Tared Crucible
---------------------------------------X 100
Sample Weight

Reagents:
1.

72% Sulfuric Acid (Specific gravity of 1.63 at 200C)

Cellulose, by Calculation
Calculation:
% Cellulose

= % Cell Walls - (% Hemicellulose + % Acid Insoluble Ash)

�-9-

Soluble Ash, by Calculation
Calculation:

% Ash - % Acid Insoluble Ash

% Soluble Ash

Cell Contents, by Calculation
Calculation:
% Cell Contents

100 - % Cell Walls

Soluble Carbohydrate, by Calculation
Calculation:
% Soluble Carbohydrate
Ash + % Ether Extract)

% Cell Contents - (% Protein

+ % Soluble

Summary of Procedures:
The cellular components are chemically separated and evaluated by these
methods as described by Fonnesbeck and Harris (1970a, 1970b, 1971) and
Harris (1970) with a few modifications to better adapt the procedures to
our particular laboratory and needs. The basic data produced by these
procedures are as follows:

% Cell Content
% Protein
% Lipid - Ether Extract
% Soluble Ash
% Soluble Carbohydrate
% Cell Walls
% Hemicellulose ) % HIll
% Cellulose
)
0 oce
% Acid Insoluble Ash
% Lignin
0

1
u ose

Sample preparations for mineral determinations were done by two different
methods during the course of this investigation.
Dry ashing was initially
used but some minerals are lost by volatilization by this process, so a
method of wet ashing as described by Adrian (1973) was adopted.
After dry ashing a one gram sample for percent ash determination, the ash
was dissolved in 2 ml concentrated HN03 and transferred to a 50 m1 volumetric
flash. This solution was the stock solution for running mineral analysis.
Phosphorus was determined by the molybdophosphoric acid spectrophotometrically
using a B &amp; L Spectronic 20. All other minerals were determined using a

�-10-

Perkin-Elmer 303 Atomic Absorption Spectrophotometer using standard procedures. The following minerals were analyzed for, in selected samples:
Phosphorus
Calcium
Magnesium
Manganese

Zinc
Copper
Iron
Potassium

Wet ashing was accomplished by placing one gram of the sample in a 4 oz
Nalge bottle. Two milliliters of perchloric acid-sulfuric acid (7+1)
plus four milliliters of nitric acid were added to each bottle and allowed
to pre-digest for twenty-four hours at room temperature. The digestion
bottles were stoppered, placed in a water bath at 700c for 3 hours. It
is important that the bottles are tightly stoppered as the pressure within
the bottle is essential to the procedure. After the digestion was completed
each sample was made up to 100 ml volumetrically with distilled water.

Food Habits Analysis
Bighorn sheep feca.lsamples were collected from the three study areas beginning in June 1971. Collections were made monthly June - September and
thereafter during November, January and March.
Estimates of forage species used by bighorn sheep were made using microanalysis techniques to identify residues of plant tissues in fecal samples.
These samples were collected from areas where bighorn sheep were observed
grazing so to be reasonably sure that feces were from bighorn sheep and
were fresh and to reflected the current season of the year. A minimum of
ten pellet groups were collected from each study area during each collection
period. Individual groups were placed in envelopes and labeled. Efforts
were made to collect the entire pellet groups, however, this was not always
possible due to snow conditions or defecation while the animal was moving.
Prior to microanalysis the pellets were placed in a Tyler screen (4mm) and
agitated to remove foreign matter such as dirt, plant parts, etc. If soil
or dust remained on the pellets they were rinsed with small amounts of water
and then allowed to air dry. Sub-samples of 6-8 pellets from each pellet
group were selected at random and ground in a Wiley mill using a 2mm screen.
Estimations of food habits were subsequently made by the Composition
Analysis Laboratory, Department of Range Science, Colorado State University,
using1microanalysis techniques. Procedures used for preparing microscope
slides, identifying tissue residues and estimating percent relative density
of forage species comprising sheep diets were those of Sparks and Malechek
(1968). Twenty systematically located microscopic fields (lOOX) were
observed on each of ten prepared slides. Residues of plant species were
identified and the percent relative density of different forages estimated.
A plant collection from the three study areas was made by the laboratory and
sets of ne fezience slides were prepared. These reference slides were employed
as comparative aids to laboratory technicians during microanalysis of samples.

�-11-

Representative forage samples were also collected from the same areas
where sheep were observed grazing and the pellet groups collected.
Initially, all forage plants in a given area were collected, however,
as food habits became known only those forages used by the sheep were
collected. Grasses and grasslike plants were collected by clipping the
entire plant as close to the ground as possible. Browse plants were
collected by clipping current annual growth. Clipped materials were
placed in paper bags, labeled and returned to the laboratory where they
were allowed to air dry.
The collected forages were identified as to species using Bighorn Sheep
Herbarium Reference Manuals (Shepherd 1971) or Manual of Plants of
Colorado (Harrington 1954). After the forage samples were dry they were
ground in a Wiley mill using a lmm screen. Samples were stored in 4 oz
sample jars until chemically analyzed.
After the first year of this study it became evident that the microanalysis
technique for fecal analysis was not accurately reflecting the food habits
of the bighorn sheep on their ranges. As a result an experiment was
designed to test the accuracy of the method as follows:
Two consecutive l2-day feeding trials using a fistulated domestic goat
were conducted. The animal was put into an isolation pen (6.1 X 1.8 m,
with cement floor) 2 weeks prior to the first trial to accustom it to
confinement and measure·its normal daily food intake. Daily intake of its
regular feed was used as a basis for the amount of test diet fed during the
trials. Ad libitum intake was measured by offering a weighed amount of
feed and weighing back uneaten amounts each following day. Prior to feeding
a test diet, the animal was fasted 48 hours to aid in voiding its digestive
tract of previously ingested food. During the l2-day trials, test diets
were force-fed twice daily by introducing 250g of the diet through the
fistula directly into the rumen. No other food was available to the animal.
After each feeding the most recent fecal group was collected, placed in a
paper sack and labeled as to date and time of collection. The pen was then
swept clean of all remaining fecal pellets.
Test diets fed were formulated with true mountain mahogany (Cercocarpus
montanus), bitterbrush (Purshia tridentata), big sagebrush (Artemisia
tridentata),fringed sagebrush (Artemisia frigida), rubber rabbitbrush
(Chrysothamnus nauseosus), snow willow (Salix nivalis) and alfalfa (Medicago
sativa). Plant species, except Salix and Medicago, were collected during
late winter by hand-clipping the previous year's growth from plants along the
front range near Fort Collins. Salix was collected from the Pikes Peak range
during June when the species was in full leaf with mature catkins present.
Alfalfa used was an exceptionally leafy, baled, third-cutting from the
previous season. Each plant species was ground individually in a hammer
mill with a 3mm screen resulting in particles varying from extremely small
to 3cm in length. After grinding, material was stored in plastic sacks at
-lOoC.
Two test diets were formulated with varying percentages of individual plant
species (Table 1). Species used were sampled to determine moisture content,
compounded, thoroughly mixed and stored in plastic bags at OoC until fed.

�-12-

Table 1. Composition (percent dry weight basis) of two formulated diets
fed during feeding trials.
Composition (Percent)
Diet I
Diet II

Plant Species

Artemisia tridentata

9.3

5.0

Artemisia frigida

14.1

10.0

Cercocarpus montanus

9.7

25.0

0

9.9

Medicago sativa

35.5

10.7

Purshia tridentata

9.3

24.9

Salix nivalis

22.1

14.5

Chrysothamnus nauseosus

Microanalysis

of Fecal Samples

Fecal samples were analyzed by the Composition Analysis Laboratory, Department of Range Science, Colorado State University, using micro-techniques
as previously described.
The laboratory technician who prepared and read the slides had extensive
training and experience in micro-techniques.
She did not have prior
knowledge of the diet-compositions but was furnished a complete list of
all browse species that could be expected but were not necessarily included
in the diets.

Digestibility Experiments
The digestibility of a particular type of feed or combination of feeds by
specific animals can be determined by feeding trials. These feeding trials
are carried out under very carefully controlled conditions where all feed
consumed by the animal is precisely measured and all feces from the animal
is collected. Chemical partitioning of both the feed being tested and the
feces being eliminated are made and the difference between those values
being consumed and those values being eliminated is the digestibility of
that forage.
The "classical" methods employed in the conduct of digestion trials are
not applicable to the free ranging big game animals so several modifications
and innovations have been employed in this experiment. These methods allow
digestion trials to be analyzed on bighorn sheep under natural conditions
on their natural ranges. It should be emphasized that these digestion

�-l3-

trials are experimental and are not to be interpreted as being without
problems and shortcomings.
This segment of the project was devoted to
trial runs on these digestion evaluation methods and the results are reported here.
The methods employed depend upon two main factors - one being the use of
an indigestible indicator occurring naturally in the forage and the other
is the accuracy of the identification of plant fragments in the bighorn
sheep feces by the Food Habits Laboratory at Colorado State University.
The "indicator" used in these studies was lignin. The plant constituent
is relatively undigestible and is easily assayed for by the methods of
Fonnesbeck and Harris (1970a, 1970b, 1971) as discussed previously.
The theory of this procedure then involves the analysis of lignin as indicator and using the fecal analysis as inference on the food intake of
the animal. By chemically partitioning the forages making up the diet of
the animal and the chemical partitioning of the feces we can then estimate
digestibility by using the following formula:
% Indicator in Feed
Digestibility = 100 - (100 % lndlcator ln Feces X

% Nutrient in Feces)
% Nutrient ln Feed

In order to compute the total digestible nutrients (TDN) value of a feed,
the digestion coefficient of each nutritive constituent is computed by
multiplying the amount of that nutrient in 100 pounds of total feed by
the digestibility for that constituent.
(The digestibility of fat is
multiplied by 2.25 because it has more energy value than the other
nutrients) •
The nutritive ratio is that ratio between the digestible protein, expressed
as unity, to the sum of digestible ether extract, digestible carbohydrate,
and digestible holocellulose (cellulose + hemicellulose) .
Two separate experiments were designed to test the feasibility of the procedure to estimate digestibility.
Each experiment consisted of collecting
feeds and feces from three study areas (Trickle Mountain, Buffalo Peaks,
and Pikes Peak) at three seasons of the year (summer, fall, winter).
In the first experiment forages were grouped into two categories (grass
and browse) where a composite sample from each grass and each browse species
collected were "pooled", thoroughly mixed and analyzed for the nutrient
constituents.
Fecal samples from each area at each season of the year were
"pooled" by taking equal quantities of at least ten separate pellet groups.
These "pooled" fecal samples were thoroughly mixed and one portion sent to
the Food Habits Laboratory at Colorado State University where they were
examined for plant composition by microanalysis of plant fragments, and the
other portion was analyzed chemically to partition the nutritive constituents.
The second experiment was conducted using specially compounded diets based
upon the percent food composition found in the feces. The compounded diet
was based upon those genera of plants which were found to be greater than
five percent in the feces. These compounded diets were made to represent
each of the three study areas at three seasons of the year (summer, fall and
winter). For specific details of these diets see Federal Aid Report w-4l-R23 "Bighorn Sheep and Mountain Goat Investigations", Work Plan 1, Job 16,
January 1974.

�-14-

Fecal samples were divided into two parts, one part was sent to the
Food Habits Laboratory for analysis and the other part was kept for
chemical analysis.
After the laboratory analyses were completed, the total digestible
nutrient (TDN) and the nutritive ratio (NR) were computed for each
combination of grass-browse and feces from each study area at each
time of the year.
Composite diet values for protein, ether extract, soluble carbohydrate,
holocellulose, and lignin were derived by mUltiplying each value of the
nutrient as determined by laboratory analysis by the percent of that
feed component in the total diet and then adding these together. For
example, if grass was analyzed as having 6.20 percent protein and grass
made up 84.58 percent of the diet then grass contributed 5.24 percent
protein (6.20 X .8458) and if browse was analyzed as having 9.80 percent
protein and made up 15.42 percent of the diet, then browse contributed
1.51 percent of the total protein (9.80 X .1542). Adding 5.24 percent
and 1.51 percent protein gives an adjusted value of 6.75 percent protein
in the composite diet.

Forage Quality Analysis Between Study Areas
An experiment was designed to test for differences in forage quality between the three study areas at three seasons of the year. This was
necessary in order to evaluate the changes in nutritional quality of bighorn sheep forages as the growing season progresses from summer into fall
and winter and the plant species grow, mature, and go into dormancy.
Because of the complexity of the composition of different plant species on
the three study areas this study was designed to limit the nutritional
comparisons to general plant types (grass and browse) rather than attempt
the complex comparisons based upon individual plant species. The main
purpose of this phase of the study was to compare the nutritive quality
of bighorn sheep forages on the three study areas at three seasons of the
year. For this reason samples were designated as "grass" and ''browse''
and no further attempt was made to differentiate plant species composition.
Five transects were selected on each of the three study areas. These
transects were laid out to cover the known winter range areas of each study
area. A wooden frame measuring one meter on each side was used to aid in
plant collections. The frame was placed on the ground at one end of the
transect and all the grass within the one square meter was clipped and
placed in a paper sack. After the grass within the frame was collected we
moved one hundred paces along the transect and at this point collected
another one square meter grass sample. A total of ten samples over a one
thousand pace transect were obtained and returned to the laboratory for
analysis.
Browse samples were collected more or less at random along "modified"
transects. Because of the distribution of browse plants on the bighorn
sheep winter ranges it was necessary to select groups of plants to sample

�-15-

and a rigid sampling scheme similar to that described for grass was not
possible. For the most part browse was collected from plants closest to
points where grass samples were collected. Current annual growth was
clipped from the plants for chemical analysis.
All forage samples were returned to the laboratory and allowed to air
dry. They were then ground in a Wiley mill (equipped with stainless
0
steel screens) and placed in glass jars after being oven dried at 60 C
for 24 hours.
Because of the large number of individual samples it was not practical to
do chemical analysis on each one so each transect was considered as a
separate entity for comparison purposes so that all samples collected
along one transect were composited into a single sample. This was
accomplished by thoroughly mixing each individual forage sample and removing a ten gram subsamp1e. These subsamp1es were then composited and
the composite sample was then thoroughly mixed and submitted for chemical
analysis.
The above described sampling scheme then resulted in five grass samples
and five browse samples representing five transects from each of the
three study areas at three seasons of the year, so that the entire study
design resulted in a total of ninety samples for complete laboratory
analysis.

Bighorn Sheep Tissue Analysis
Tissue samples from bighorn sheep lambs collected during the course of
this study were analyzed for trace mineral levels. At the time these
lambs were necropsied tissues were collected and frozen for this purpose.
Seven lambs were collected during the summer of 1972, nine lambs collected
during the summer of 1973, and seven lambs collected during the summer of
1974. The following tissues were analyzed from each lamb: hair, liver,
spleen, kidney, skeletal muscle, and heart muscle.
The tissues were removed from the freezer, allowed to thaw and placed in a
drying oven at 600C until dry. Low drying temperatures were used because
of the reported volatility of selenium from tissues when dried at high
temperatures.
A one gram sample of dried tissue was weighed into a digestion vessel and
digested by the wet pressure method previously described (Adrian 1973) and
mineral levels were determined by procedures previously described.

RESULTS AND DISCUSSION
Nutrition involves various chemical and physiological activities which
transform food elements into body elements, therefore a brief consideration
of the chemical composition of the animal body in relation to the composition
to its food is useful to give a general picture of the nutrition process.
Food must supply nutrients which can be used to build and renew the comr
ponents of the body and to produce growth, hair, milk, etc. and it must

�-16-

furnish energy for all these processes as well. While there are certain
animal species which are entirely carnivorous, the plant kingdom is the
original and essential source of all animal life because plants are able
to utilize the energy of the sun and build substances which will nourish
the animal. Plants make use of carbon dioxide, water, nitrates and other
mineral salts to form carbohydrates, fats and proteins which the animal
must have to build and maintain their bodies. Thus plants store and
animals dissipate energy.
Plants contain the same substances found in animal bodies, but the relative
amounts are very different. Plants generally show larger variations in
composition from one species to another than do animals.
The principal constituent of both plants and animals is water. This
water content in plants decreases as the plants mature and in animals the
water content will vary somewhat with age. The newborn animal has a much
higher percentage of water than does the mature fat animal. It is vital
to the life of the organism that the moisture level be maintained. Water
functions as a solvent in which nutrients are transported and in which
waste products are excreted.
The main difference in the composition of plants and animals lies in the
fact that dry matter of plants consists principally of carbohydrate. This
constituent in plants serves as both structural and reserve material
while in animals protein comprises the structure of the soft tissues and
fat is the reserve. Although carbohydrate is a minor constituent of the
animal body it is the principal food of most animals, especially ruminants.
Research workers conducting experiments on the nutritive requirements of
animals normally base their conclusions on the average gain in weight or
feed efficiency for groups of animals. Where deficiency symptoms are the
criteria for determining nutrient adequacy or inadequacy, freedom from
the deficiency syndrome in all animals in experimental groups is usually
the basis for conclusions. When dealing with wild-free ranging ungulates
the problems of studying the nutritional requirements presents many problems not experienced by those workers dealing with domestic livestock.
The approach to any given problem must by necessity be different than what
might be used in more conventional circumstances.
For these reasons the
experimental designs and procedures outlined in this study do not necessarily
follow well developed and generally accepted procedures of nutritionists
and the results and interpretations are in most cases compared to what is
known about domestic livestock.

Study Areas
Since the start of this project several years ago the bighorn sheep herd
on Pikes Peak has experienced a number of changes. During the summer of
1971 it was noted that several lambs were not doing well and this was the
start of a rather severe lamb mortality. Since that time extensive effort
has been expended on this bighorn sheep herd by the College of Veterinary
Medicine at Colorado State University and the results of these efforts are
reported under Federal Aid Reports W-41-R, Work Plan I Contract - "Spontaneous
Diseases of Bighorn Sheep".

�-17The Buffalo Peaks and Trickle Mountain study areas are unchanged from
the original concepts at the outset of the study.

Forage and Feces Analytical Procedures
The methods of Fonnesbeck and Harris (1970a) have developed a method for
cell wall analysis which more accurately partitions the cell wall constituents and may be used to analyze all types of feeds. The plant cell
wall constituents include the partially nutritive matter and the nonnutritive matter. The partially nutritive matter includes the cell wall
carbohydrates, cellulose and hemicellulose, and is digested only by
enzymes produced by microorganisms within the digestive tract. The
nutritive value and digestibility of cellulose and hemicellulose may not
vary enough to warrant their separation. This supposition was followed
for most parts of this study as the analytical procedure to separate the
two are time consuming and not warranted so all results are reported as
"holocellulose". The non-nutritive matter includes lignin and acid insoluble ash.
The nutritive matter is found in the cell contents which is digested by
enzymatic action in the digestive system or is otherwise soluble enough
for absorption. This includes soluble carbohydrates, proteins, ether
extract and soluble ash.
The chemical methods of Fonnesbeck and Harris were developed from Van
Soest's acid detergent lignin (Van Soest 1963), and cell wall constituents
(Van Soest and Wine 1967) analysis.
According to Harris (1970) the main deviations from Van Soest's methods
are:
1.

Cell walls are determined at pH of 3.0 in a citrate buffered
detergent solution. At this pH, there is a recovery of cell
wall constituents, particularly acid insoluble ash.

2.

A pepsin digestion of the cell wall sample followed by detergent
extraction removes 90 to 98 percent of the protein. Less than
1.0 percent protein of a sample containing 10 percent protein
would be retained in the cell wall residue. This is less than
the amount remaining in the acid detergent fiber in preparation
for the acid detergent lignin analysis. Therefore, 72 percent
sulfuric acid lignin can be determined directly on the cell
wall residue.

3.

In Van Soest's analysis, cellulose is determined as the difference between acid detergent fiber and acid detergent lignin or
as the organic residue from the determination of permanganate
lignin. Hemicellulose is calculated as the difference between
cell wall constituents and acid detergent fiber. Fonnesbeck
and Harris separate the cellulose and hemicellulose of the cell
wall with a hemicellulose and lignin analysis.

�-18-

4.

Lignin values determined on the cell wall residue or on a
hemicellulose extracted cell wall residue are considerably
higher than acid detergent lignin values. Subsequent investigations showed that the acid detergent solution was
dissolving part of the lignin.

s.

The techniques of the 72 percent H2S04 lignin method of
Fonnesbeck and Harris make it unnecessary to add asbestos
to the crucible to aid in filtering. Therefore, the ash
remaining after the lignin determination is entirely from
the original sample and can be reported as acid insoluble
ash, or the acid insoluble ash can be extracted with 48
percent hydrobromic acid for a presumptive analysis for
silica.

Food Habits Analysis
A knowledge of food habits by bighorn sheep in any given area is a necessary
prerequisite for studying forage competition with other ungulates or
studying the nutritional adequacy of these ranges to meet the requirements
of the bighorn sheep. Past food habit studies in Colorado were primarily
made on rumen samples collected from hunter harvested bighorn sheep during
late summer or early fall seasons. These studies have been supplemented
by a few observations and rumen samples collected at opportune times (Moser
1962) • The food habits of bighorn sheep on a year-around basLs is not well
documented.
Collecting animals from already declining sheep herds was not
practical, therefore the microscopic technique of examining plant fragments
in feces was used. These methods have been used extensively by other
workers for studying food habits of small rodents, rabbits, and grasshoppers (Baumgartner and Martin 1939; Dusi 1949; Myers 1962; Bear and
Hansen 1966; Sparks 1968; Sparks and Malechek 1968). More recently the
technique has been used to examine the feces of large herbivores (Storr
1961). Cooperrider (1969) used the microscopic technique in studying the
food habits of bighorn sheep in Montana.
An extensive study was conducted by Todd (1972) which was conducted on the
Trickle Mountain study area.
Primary forage species identified in bighorn sheep feces from the three
study areas are summarized in Table 2. Comparisons of data among the three
areas indicate sheep using the Trickle Mountain range utilize a wider range
of plant species than the sheep on either Pikes Peak or Buffalo Peaks.
Grasses and grass-like forage comprise the major portion of bighorn sheep
diets with increasing utilization of browse during early fall and into
winter (Fig. lA). Use of forbs by sheep on all three study areas was found
to be relatively small being less than four percent of any diet examined.
The key to the digestibility trials as originally designed into this study
relied heavily upon the accuracy of the micro element fecal analysis to
differentiate plant components to species of origin. It became evident
that the technique was not accurately breaking the fecal material into
precise components so an experimental feeding trial was designed to test
the procedure.

�-19-

Table 2. Primary forage species identified in fecal samples of bighorn
sheep collected from the Buffalo Peaks, Pikes Peak and Trickle Mountain
ranges, June 1971 - September 1972.

Plant Species

Buffalo Peaks

Agropyron spp.

0.6 -

7.4

Percent (Range)
Trickle Mountain
Pikes Peak

0.8 - 10.3
1.1-

Bouteloua gracilis
2.7 - 53.6

Carex spp.

4.2 - 19.2

7.8-91.8

0.1 - 6.7

Danthonia parryi
6.8

Deschampsia caespitosa

6.0 -

Festuca spp.

l3.0 - 66.8

Muhlenbergia

spp.

29.9

0.1 -

0.5 - 57.6

0.7 - 52.9
0.1 - 56.6

5.6

0.2 - 11.0

Oryzopsis hymenoides
0.5 - 14.7

3.5 - 15.2

Stipa spp.

2.5 - 10.7

0.1 -

Artemisia frigida

0.1 -

7.7

0.2 - 75.4

0.2 -

7.4

0.1 - 14.1

Poa spp.

2.4 - 10.5

12.3

Cercocarpus .montanus

0.7 - 42.0

Holodiscus dumosus
Pinus spp.
Salix spp.

0.4 -

6.8

0.1 - 28.8

27.6

35.1

0.5 -

9.9

0.3 - 15.7

Yucca spp.
Shrubs (not identified)

5.4

8.4 - 53.4

0.2 - 68.3

0.7 - 40.4

Force feeding test diets directly into the rumen through a fistula proved
to be an effective means of feeding a diet of known composition to an
animal. This method eliminated any selectivity of food intake by the animal
which might have occurred if the diets were offered free-choice or in a
cafeteria type feeding trial. During the trials rumination appeared normal,
the animal remained in tood flesh and quite often was observed contentedly
chewing cud. Feces eliminated were a normal size and consistency.

�-20-

Sununer

Fall

Winter

100
PIKES
PEAK

50

a
100

BUFFALO
PEAKS

50

a
100

TRICKLE
MOUNTAIN

50

a

GRASS

BROWSE

Fig. lA. Percent relative density of grass and browse use by bighorn sheep
on Pikes Peak, Buffalo Peaks and Trickle Mountain at three seasons of the
year - sununer, fall and winter.

�-21-

Fecal samples collected after the animal had been fed the test diet for
10 and 12 days during each trial, contained little identifiable plantresidues - a few Artemisia trichomes and a few unattached trichomes which
may have been from Salix, Cercocarpus or Purshia. The bulk of material
was unidentifiable woody-stem material (Hanson, letter of 20 Sept. 1972).
Similar results were reported by Zyznar and Urness (1969), who fed 17
tree, shrub and herbaceous plant species separately to captive mule and
white-tailed deer. They found only low percentages of recognizable plant
residues in fecal samples using microanalysis techniques.
Unfed samples of the test diets were also analyzed by microanalysis techniques as an additional test of the method's accuracy in identifying plant
species in the diets and estimating percent composition. All species in
the diets were identified. However, microanalysis results include 2.1
percent Chrysothamnus sp. and 0.2 percent Pinus sp. in Diets I and II,
respectively. These species were not formulated in the respective diets.
Estimates of the relative percentages of some plant species in the diets
were also highly over or underestimated (Table 3). Stewart (1967) questions
validity of the microanalysis technique for estimating relative percent of
plant species in an animal's diet due to some forage plants fragmenting
more easily than others. We found, after feeding a number of perennial
grasses to African ungulates, that some plants easily fragmented and consequently appeared more important in their diets although intake was the
same or less than that of other species.

Table 3. Composition (percent dry weight basis) of two test diets compared
to the estimated relative composition determined by microanalysis.
(y/x is
ratio of known percent (y) to estimated percent (x); values&gt; 1 are overestimates and CL underestimates.

Plant Species

Compo

Diet I
Estimated

y/x

Artemisia tridentata

9.3

6.4

0.7

5.0

10.7

2.1

Artemisia frigida

14.1

42.1

3.0

10.0

41.6

4.2

Cercocarpus montanus

9.7

1.3

0.1

25.0

5.8

0.2

0

2.1

9.9

8.6

0.9

Medicago sativa

35.6

25.6

0.7

10.7

11. 7

1.1

Purshia tridentata

9.3

2.1

0.2

24.9

10.0

0.4

Salix nivalis

22.1

20.4

0.9

14.5

11.4

0.8

0

0

0

0.2

Chrysothamnus nauseosus

Pinus sp.

Diet II
Compo Estimated

y/x

�-22-

These results question usefulness and accuracy of fecal microanalysis
techniques for determining food-habits of ruminants. Reasons why no
identifiable plant residues were found in fecal samples from these trials
are unknown. Other investigators (Hegg 1961; Kiley 1966; Lay 1965;
Stewart 1967; Zyznar and Urness 1969) have used the method, with varying
success, to identify plant residues in feces from other ruminants. Differences in digestive capabilities among various species of ruminants studied
during these investigations and the domestic goat used during these trials
may account for differences of results obtained. Digestion may be more
complete in ruminants possessing relatively large capacity digestive organs
such as the goat, compared to ruminants with small organs. Easily digestible
plants or plant parts may have been completely digested by the goats digestive system leaving only undigestible wood-stem material to be eliminated
in feces.
Accurate food-habits data are essential for nutritional studies involving
any species of animal. Based on results of these trials, application of
fecal microanalysis techniques for determining food-habits of bighorn sheep
in our current studies cannot be reliably used.
However, determining food
habits of free-ranging wild animals is extremely difficult and to date fecal
microanalysis offers the best approach to the problem. It is recommended
therefore, that use of the technique be continued and further investigations
be made to clearly define its application and limitations. Most of the
digestibility work therefore has been limited to two broad classes of
plants - "browse" and "grass" with no attempt to break the diets into more
precise elements.

Digestibility Trials
During the initial segment of this study attempts were made to estimate
digestibility of range forages by free ranging animals, but as described
previously the technique relied heavily on the accuracy of the microelement
fecal analysis to identify plant parts as to species of origin. Because
of the problems associated with the microelement analysis technique this
phase of the study was not persued after the initial trials.
Two separate experiments were designed to test the feasibility of estimating
digestibility.
Each experiment consisted of collecting forages and feces
from the three study areas as previously described and subjecting them to
the following experimental procedures.
The first experiment consisted of grouping forages into the two categories
of "grass" and ''browse'',where a portion of each grass and each browse was
pooled, thoroughly mixed and analyzed for their nutrient contents. Fecal
samples from each area at each season of the year were pooled by taking
equal quantities of at least ten separate pellet groups. These pooled
fecal samples were thoroughly mixed and one portion was sent to the Food
Habits Laboratory for microanalysis and the other was analyzed chemically
to partition the nutritive constituents.
After the laboratory analyses were complete, the total digestible nutrients
(TDN) and the nutrient ratio (NR) were computed for each combination of
grass-browse and feces from each study area at each time of the year (detailed
data on this experiment was presented in Federal Aid Progress Report W-4l-R-23

�-23-

June 1, 1972). Composite diet values for protein, ether extract,
soluble carbohydrate, ho1oce11u1ose and lignin were derived by multiplying each value of the nutrient by the percent of that feed component
in the total diet and then adding these together. For example, if grass
was analyzed as having 6.20 percent protein, and grass made up 84.58 percent of the diet then grass contributed 5.24 percent of the protein (6.20 x
.8458) and if browse was analyzed as having 9.80 percent protein and browse
made up 15.42 percent of the diet, then browse contributed 1.51 percent of
the total protein (9.80 x .1542). Adding 5.24 percent and 1.51 percent
protein gives an adjusted value of 6.75 percent protein in the composite diet.
The second experiment was conducted using specially compounded diets based
upon the percent food composition found in the feces. The compounded diet
was based upon those genera of plants which were found to be greater than
5 percent in the feces. These plants were then added to the compounded
diet in these same percentages, thoroughly mixed and then analyzed chemically along with the feces from which the plant composition analysis was
made. These comparisons again were made on samples from the three study
areas at three seasons of the year.
Tables 4 and 5 show the results of the two experiments previously described.
The total digestible nutrients and the nutritive ratio resulting from the
two methods are comparable, but a great deal of additional work must be
done before either technique can be used to evaluate range quality for bighorn sheep. As stated previously the whole technique is based upon the
accuracy of the microanalysis techniques for fecal composition of plant
fragments. For these reasons the data obtained in this phase of the study
is presented here, but any conclusions must be carefully considered in light
of the results of the feeding trial experiments previously described.

Table 4. Results of nutritive quality of bighorn sheep ranges by the
technique of compounding forage samples by percent grass and percent browse.
Buffalo Peaks
NR
TDN

Pikes Peak

Season

Trickle Mountain
NR
TDN

Summer

51.34

1:10.80

63.30

1:11.08

65.77

1:12.05

Fall

64.12

1:16.33

73.03

1:19.40

67.88

1:19.08

Winter

68.27

1:16.73

52.54

1:22.88

62.10

1:20.27

TDN

NR

�-24Table 5. Results of nutritive quality of bighorn sheep ranges by the
technique of compounding forage samples by percent plant composition.

Season

Trickle Mountain
TDN
NR

Buffalo Peaks
TDN
NR

Summer

72.85

1:11.22

69.89

1:14.06

68.54

1: 6.84

Fall

59.94

1:11.78

56.02

1:15.14

72.13

1:20.60

Winter

50.52

1:19.50

69.49

1:29.88

62.00

1:29.24

Pikes Peak
TDN
NR

With domestic livestock the most common problem in nutrition is the deficiency of energy. This may result from a lack of sufficient feed or from
too little net energy available to the animal from the feed consumed. Poor
quality roughages are not only poorly digested but are usually consumed in
smaller amounts. Snow cover, drought or low dry-matter content of lush feeds
may also contribute to a shortage of total nutrients consumed by an animal.
Digestibility decreases in a plant as it matures due to increase in lignification. Total intake decreases with decreased digestibility, slower
passage through the digestive tract and with reduction of the palatability
of the feeds.
These energy shortages may be complicated by other deficiencies such as
protein, minerals and vitamins with the reSUlting effect of slowing growth,
loss of weight, reproductive failure and increased mortality. Internal
parasite infestations are more severe in animals suffering from malnutrition
because of their lowered resistance.
The requirements for any given animal will vary according to its sex and
stage of maturity. Those requirements for a mature ram will be different
from a ewe for example and the requirements for the ewe will vary from time
to time depending on the period of gestation, lactation, etc. In the case
of wild ungulates it would appear that as the animal requirements for nutritional factors goes up it corresponds to the season of the year when forage
quality decreases.

Forage Quality Analysis Between Study Areas
In range investigations the worker is interested in the nutritive value of
the herbage available to the animal and the nutritive value of the forage
actually consumed. The most reliable measurement of the nutritive value of
range herbage is obtained by conducting a well-designed grazing trial, but
because of the difficulties in carrying out these studies with wild ungulates,
the relative nutritive value of forage may be estimated from chemical
analysis provided the data is properly interpreted.
Chemical determinations
are used to measure specific nutrients and are of value in revealing deficiencies or lack of deficiencies.
The presence or absence of toxic materials
can also be determined by chemical analysis.

�-25-

Factors such as stage of growth, botanical composition, soil type and
topography have been found to influence the chemical composition of forage,
and it is essential that these variables be considered (Cook and Harris
1950a, 1950b; Crampton and Jackson 1944; Garrigus and Rusk 1939; Martin
1952; Reid and Kennedy 1959b; Reid et al. 1950, 1959a).
Under open range conditions it is difficult to determine the diet of animals
because there are many species of plants. Topography influences animal
distribution and forage is harvested by grazing animals in a highly selective
manner (Cook et al. 1948, Harris et al. 1959). As has already been discussed
the material selected by the animal should be sampled and the diet must be
known so that individual analysis may be properly used to calculate the composition of ingested forage. In cases where free ranging wild ungulates are
under investigation it is not feasible to use the more sophisticated methods
employed with domestic livestock so the investigator is forced to rely on
indicator methods or chemical analyses of the herbage available for grazing
to characterize the nutritive value o,f the forage.
Sampling methods as described for this phase of the study was designed somewhat after that used in studies at Cornell University where they used an
iron circle with an area of two square feet for harvesting forage for
botanical and chemical composition. This ring was tossed at random on the
range under investigation and the forage within the ring was harvested by
clipping to a uniform height.
In using chemical analysis to predict quality, the investigator must consider two serious limitations: (1) the forage samples used in chemical analysis
may not be strictly representative of the forage consumed by the grazing
animal, and (2) the feeding value and chemical composition are not necessarily
closely related.
The investigator should understand the limitations of interpretation of
chemical determinations, but if done carefully it is possible to characterize
differences in forage quality. With these considerations in mind the following
data is presented and discussed to compare the nutritive differences between
the three study areas at three seasons of the year and will be treated on an
individual component basis.
In an attempt to correlate total diet values of both grass and browse used by
sheep at any given period of time and on a specific study area the total
nutrient value of any single component was computed and totaled. For example
if Pikes Peak summer range forage contained 8.56 percent protein in the grass
and 8.64 percent protein in the browse and the fecal composition was found to
be 65 percent grass and 35 percent browse then the total protein would be
(8.65 x .65) + (8.64 x .35) = (5.56 + 3.02) = 8.58 percent protein. Values
used for these computations are found in Figure 1.

Cell Contents
The cell contents are the total of the protein, ether extract, soluble carbohydrate and soluble ash and make up the majority of the digestible portion of
the edible plant. The total cell walls are the reciprocal of cell contents
therefore the two divisions of plant parts can be considered together.

�-26Grasses exhibited the greatest differences between areas during the summer
season (sig. 99%). This can be attributed to a large part of the differences in plant growth and maturity due to a large part by the differences in
altitude between the three study areas and the differences in growth and
maturity of the plants. The Trickle Mountain area is lower in altitude
and the growing season is at least three to four weeks earlier than the other
two.
Browse plants also show significant differences (99%) during the summer season.
This again could be attributed to differences in the growing season between
the different study areas. Figure 2 shows the values of total cell contents
from the three study areas at three seasons of the year for grass, browse and
a composite diet.

Protein
Figure 3 shows the results of crude protein analysis on grass and browse
samples from the three study areas at three seasons of the year and a comr
posite diet protein level based upon the percent use of grass and browse by
bighorn sheep. Protein is the principle constituent of the organs and soft
tissues of the animal body and a liberal, continuous supply is needed in the
food throughout life for growth, and repair. The term protein is a
collective one which embraces an enormous group of closely related but
physiologically distinct members. From the standpoint of nutrition the
important aspect is the amino acid make-up of proteins. The quality of protein which is influenced by the amino acid make up or balance is not a critical
factor for ruminants because of the activity of the microorganisms within
the rumen. The quantity of protein becomes more important than the source,
and in general the quantity of digestible protein will vary widely with
different feeds, particularly between coarse, mature, weathered, or poor
quality roughages.
Insufficient protein intake results in reduced appetite, lowered feed intake
and poor feed efficiency. This lowered feed intake results in poor growth
and muscular development, reduced reproductive efficiency, as well as anemia
and problems with gestation and lactation in ewes.
Significant differences (99%) were found between the three study areas at
all seasons of the year but more importantly is the level of protein in the
diet when compared to the animal needs. During the summer months it appears
that there would be adequate levels of crude protein, but this supply falls
off sharply during the fall and winter in all areas. As grass matures and
leaves fall from the browse plants the protein levels change considerably
and bighorn sheep are obligated to subsist on these levels throughout the
balance of the year. Protein levels in excess of 12 percent are considered
excellent, 10 percent to 12 percent levels are considered good, 7 percent to
10 percent are considered fair and below 7 percent protein is considered poor
for ruminants.
It would appear that crude protein levels could be a problem for bighorn
sheep on these study areas during the winter months.

�-27-

CELL CONTENTS

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COMPOSITE
DIET

o Pikes Peak
o Buffalo Peak
L:::.. Trickle Mtn.

Fig. 2. Total cell contents of forage from three study areas at three seasons
of the year for gr~ss, browse and a composite diet based upon percentage of
grass and browse used by bighorn sheep.

�-28-

PROTEIN

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o Pikes Peak
o Buffalo Peak
D.. Trickle Mtn.

Fig. 3. Percent crude protein in forage from three study areas at three seasons
of the year for grass, browse and a composite diet based upon percentage of
browse and grass used by bighorn sheep.

�-29-

Ether Extract
Figure 4 illustrates the amounts of ether extract in forage from the three
study areas at three seasons· of the year as well as a composite diet based
upon percentages of grass and browse in the feces of bighorn sheep. The
ether extract contains a complex of materials including the fats, oils and
some waxes as well as fat soluble vitamins, phospholipids, and chlorophyll.
The fats in food serves as a source of energy the same as for carbohydrates.
Fat is much concentrated than carbohydrate and furnishes 2.25 times as much
energy· per pound of feed. For this reason a diet high in fat will have a
higher percentage of total digestible nutrients than a diet low in fat.
This could have some effect· on bighorn sheep diets as the winter food values
drop considerably and this is the season of the year when energy requirements for maintaining body heat will be higher.
There was a significant difference (95%) in the amount of ether extract
found in grass during the summer and fall. This difference could be attributed to differences in stages of plant maturity between the study areas.

Carbohydrates
The broad group of foodstuffs classed as carbohydrates includes the sugars,
starches, celluloses, hemicelluloses, gums and related substances such as
lignin. The soluble carbohydrate (Fig. 5), holocellulose (Fig. 6) and lignin
(Fig. 7) will be discussed together under the general heading of "carbohydrate".
Carbohydrates form the largest part of the animals diet as carbohydrates form
about seventy-five percent of the dry weight of the plant on which all animals
ultimately depend for sustaining life.
The soluble carbohydrates which include the sugars and starches are easily
digested by the animal and have a high feeding value. The more complex
carbohydrates such as cellulose and hemicellulose form the ''woody''fiber of
the plant and are less completely digested. The highly complex lignin is not
utilized by the animal and passes through the digestive system unaltered.
As plant material grows and matures the relative percentage of carbohydrates
change from the more desirable soluble sugars to the more complex celluloses
and hemicellulose (holocellulose) and lignin and this again contributes to
the fact that the total digestible nutrients in any given forage decreases
as the growing season progresses. One of the more drastic changes occur in
browse plants as the leaves falloff the stems after the killing frosts.
The digestion of carbohydrates in the rumen is dependent upon microorganisms
which break down the more complex forms to products which can be assimilated
by the body. This process by microorganisms requires the expenditure of
energy and the more complex the carbohydrate the more energy required to
break them down and the less net energy available to the animal. This fact
in part accounts for the differences in digestibility of forages between
the summer and winter forages. The total energy available to the animals on
poor winter forages therefore is considerably less than that for summer
forages. The significance of these differences in bighorn sheep is not
known but warrants further investigation.

�-30-

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COMPOSITE
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Pikes Peak

o Buffalo Peak
D.. Trickle Mtn.

Fig. 4. Percent ether extract in forage from three study areas at three seasons
of the year for grass, browse and a composite diet based upon percentages of
grass and browse used by bighorn sheep.

�-31-

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0 Pikes Peak
0 Buffalo Peak
6. Trickle Mtn.

Fig. 5. Percent soluble carbohydrate in forage from three study areas at three
seasons of the year for grass. browse and a composite diet based upon percentage
of grass and browse used by bighorn sheep.

�-32-

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o Pikes Peak
[] Buffalo Peaks

D. Trickle Mtn.
Fig. 6. Percent holocellulose in forages from three study areas at three seasons
of the year for grass, browse and a composite diet based upon the percentages of
grass and browse used by bighorn sheep.

�-33-

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o Buffalo Peak
6. Trickle Mtn.

Fig. 7. Percent lignin in forages from three study areas at three seasons of the
year for grass, browse and a composite diet based upon the percentages of grass
and browse used by bighorn sheep.

�-34-

Ash
One of the advantages of the methods of Fonnesbeck and Harris in the
partitioning of nutritional constituents is that the procedure allows
for the separation of the ash fraction into a soluble ash and an acid
insoluble ash. Figure 8 shows the percentages of soluble ash and Figure
9 shows the percentages of acid insoluble ash in the forages from the
three study areas at three seasons of the year for grass, browse and a
composite diet based upon the percentages of grass and browse used by bighorn sheep.
Total ash is present in all plant material and consists of the mineral
matters of the plant tissue. Mineral nutrients are essential for growth
of both the plant and animal and consist of phosphorous, potassium, sulfur,
calcium, iron, magnesium, copper, zinc, and many other minerals. The acid
insoluble ash fraction is primarily the silicates and have no nutritional
value as far as the animal is concerned.
Each of the essential minerals will be considered separately and their
importance to the animal can not be underestimated.

Calcium and Phosphorus
Forages which are deficient in either calcium or phosphorus or both result
in abnormal bone development causing rickets or osteomalacia.
Calcium is not generally a problem in the western United States where soils
are derived from calcium carbonate or dolomite. The forage may be too high
in calcium and actually interfere with phosphorus metabolism. The calcium
levels found in the forages on the three study areas do not appear to be
causing any real problem except in association with phosphorus.
Phosphorus is utilized in the metabolism of almost all nutrients through its
role in vitamin and enzyme activity.
Aphosphorosis causes osteomalacia,
production of weak lambs and decreased milk production. Low phosphorus intake
affects energy utilization resulting in slow growth, depraved appetite,unthrifty appearance and general lack of body fat. Phosphorus deficiency in
ranges is rather widespread throughout the west with forage becoming deficient after it reaches maturity.
Figures 10, 11 and 12 show the levels of calcium, phosphorus and the calcium
phosphorus ratio in forages from the three study areas at various seasons of
the year. In any discussion of calcium and phosphorus it is important to
consider the ratio of calcium to phosphorus.
If there is an excess of calcium
or of phosphorus, detrimental effects may be produced. In examining the
levels of calcium and phosphorus in forage from the three study areas the
amounts of calcium seem adequate while phosphorus is in short supply
especially during the winter months. This results in a Ca:P ratio which is
not desirable and can be a significant factor in the management of bighorn
sheep.

�-35-

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COMPOSITE
DIET

0 Pikes Peak
0 Buffalo Peak
6. Trickle Mtn.

Fig. 8. Percent soluble ash in forages from three study areas at three seasons
of the year for grass, browse and a composite diet based upon the percentages
of grass and browse used by bighorn sheep.

�-36-

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DIET

0 Pikes Peak
0 Buffalo Peak
b. Trickle Mtn.

Fig. 9. Percent acid insoluble ash in forages from three study areas at three
seasons of the year for grass, browse and a composite diet based upon the
percentages of grass and browse used by bighorn sheep.

�-37-

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o Pikes Peak

o Buffalo Peak
6. Trickle Mtn.

Fig. 10. Percent calcium in forage from three study areas at three seasons of
the year for grass, browse and a composite diet based upon the percentages of
grass and browse used by bighorn sheep.

�-38-

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COMPOSITE
DIET

Buf f a.Lo Peak

6. Trickle Mtn.

Fig. 11. Percent phosphorus in forage from three study areas at three seasons
of the year for grass, browse and a composite diet based upon the percentages
of grass and browse used by bighorn sheep.

�-39-

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o Pikes Peak

o Buffalo Peak
6Trickle Mtn.

Fig. 12. The calcium to phosphorus ratio of forages from three study areas at
three seasons of the year for browse, grass and a composite diet based upon
percentages of grass and browse used by bighorn sheep.

�-40-

Potassium
The levels of potassium in forages from the three study areas are shown
in Figure 13. The amount of potassium necessary in the diet is about
0.30 percent and all values in forage analyzed show levels higher than
this level. From the results of these data a potassium deficiency in
bighorn sheep on these study areas is not likely.

Magnesium
The minimum requirement for magnesium in the diet has been set at approximately 0.08 percent and all values in forages on the three study areas
exceed this amount (Fig. 14). It is unlikely that deficiency problems
for magnesium could exist on the study areas for bighorn sheep.

Iron
Figure 15 shows the levels of iron in forages consumed by bighorn sheep.
These levels are somewhat low in all three study areas at all seasons of
the year. Additional work should be done on iron levels in these forages
to further evaluate the effects, if any, of these forage levels on the
efficiency of bighorn sheep production.
Iron plays an important function in the body as a major constituent of
hemoglobin so iron is vital to every tissue in the body. It occurs as an
iron-porphyrin nucleus, known as heme, and is found not only in hemogloblin
but also in proteins that are components of cytochrome c, peroxidase, catalase
and other enzymes.
The most notable effect of an iron deficiency is anemia. The effects of
anemia are severe on both the adult animal and any offspring which suffer
from the lack of iron in the diet. The animal has the ability to conserve
or recycle its iron reserves and there would be some question of how severe
a borderline deficiency of iron in forage would be.
Iron in the soil is not always available to the plants. The iron can exist
in complex forms which are not easily broken out in the soils and the soil
pH will influence the availability of iron to the plants.

Zinc
Figure 16 shows the levels of zinc in forages from the three study areas.
Investigations by Smith (1962) showed that zinc is essential to domestic
sheep. Sheep on a zinc deficient diet for a period of ten weeks showed
slipping wool, swelling and lesions around the hooves and eyes, excessive
salivation, anorexia, eating wool, general listlessness, and reduction in
growth.
Due to the low levels of zinc found in the fall and winter diets and especially
in the Buffalo Peaks and Trickle Mountain areas it would indicate possible
problems with this essential mineral. Additional work should be done in
this area to further investigate these potential problems.

�-41-

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W

GRASS

BROWSE

COMPOSITE
DIET

o Pikes Peak
o Buffalo Peak
f::.. Trickle Mtn.

Fig. 13. Percent potassium in forages from three study areas at three
sepsons of the year for browse, grass and a composite diet based upon
percentages of grass and browse used by bighorn sheep.

�-42-

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BROWSE

COMPOSITE
DIET

o Pikes Peak
D Buffalo Peak
b. Trickle Mtn.

Fig. 14. Percent magnesium in forages from three study areas at three seasons of
the year for browse, grass and a composite diet based upon the percent grass and
browse used by bighorn sheep.

�-43-

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o Pikes Peak
o Buffalo Peak
b.

Trickle Mtn.

Fig. 15. Parts per million iron in forages from three study areas at three
seasons of the year for browse, grass and a composite diet based upon percentage
of grass and browse used by bighorn sheep.

�-44-

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~
p..,

14

G)

p..,

30 .1-

~

e

e
EJ
~
&amp;.

i

e

Iii
s.

20 -l-

E&gt;
I&gt;

8

m

i

10 .1I

J

S

••
I

F

I

I

I

I

I

I

-1

I
I

W

S

F

W

S

GRASS

BROWSE

I

;

I

T

W

COMPOSITE
DIET

o Pikes Peak
D Buffalo Peak
~

Trickle Mtn.

Fig. 16. Parts per million zinc in forage samples from three study areas at
three seasons of the year for browse, grass and a composite diet based upon
percentage of grass and browse used by bighorn sheep.

�-45Copper
Copper as a micronutrient is very complex and can be associated with
other mineral problems. Figure 17 shows the copper levels found in the
forages from the three study areas and was found to be well below recommended
levels. Copper deficiency may manifest itself as a primary deficiency or
in combination with cobalt and possibly iron. Animals suffering copper
deficiency suffer from anemia as the animal appears to have trouble absorbing iron at a normal rate. Symptoms of copper deficiency are seen in
young lambs (domestic) with muscular incoordination accompanied by partial
paralysis of the hindquarters and degeneration of the myelin sheath of the
nerve fibers. Copper deficiency also affects the wool or hair coat causing
a rough-lackluster appearance.
To further complicate the copper metabolism picture it has been found that
a copper-molybdenum sulfur complex exists where excess molybdenum in the
presence of sulfur can tie up the copper causing deficiency problems. In
view of the fact that many areas of Colorado alpine are high in molybdenum
this complex should be investigated further to see if this could cause
problems with bighorn sheep.

Selenium
The relationship of selenium with vitamin E metabolism and the importance
in preventing muscular dystrophy in lambs has been established by Muth et
al. (1958). Muscular dystrophy has been diagnosed in bighorn sheep from
Pikes Peak and the levels of selenium, as shown in Figure 18, indicate that
selenium levels are below recommended levels of 0.10 ppm in forage from Pikes
Peak. Those forage levels from Trickle Mountain would indicate adequate
levels with Buffalo Peaks offering an intermediate position.
These preliminary results justify further work in this area especially in
view of recent work showing correlations of selenium with immune response
by the body to disease organisms.

Bighorn Sheep Tissue Analysis
The mean concentration and standard deviation of seven elements (calcium,
phosphorus, sodium, iron, zinc, magnesium and copper) on six tissues (hair,
liver, spleen, kidney, skeletal muscle and heart muscle) from bighorn sheep
lambs collected on Pikes Peak over a three year period are shown in Tables
6 to 11. These comparisons were made to test the tissue levels of these
lambs in an attempt to see if a supplemental feeding program associated with
baiting animals to drop net traps had any noticeable effect on the lambs.
Due to a small sample size during each year (1972 - 4 lambs; 1973 - 7 lambs;
1974 - 6 lambs) it is difficult to draw any conclusions from the data obtained.
Additional problems in analyzing this dat.aare due to the fact that during the
time between collections supplemental feeding was carried out on the Pikes
Peak herd and it is very difficult to determine if the lambs were from ewes
on supplemental feed or not.

�-46-

.1-

2'
Po.
Po.

•I-

'-'
p:;
~
Po.
Po.
0

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u
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8

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p:;

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4.0

.

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C3

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a

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.-l

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,
I

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--'-

_1

S

F

~

J

S'

F

'w

BROWSE

COMPOSITE
DIET·

o Pikes Peak

o

Buffalo Peak

6 Trickle Mtn.

Fig. 17. Parts per million copper in forages from three study areas at
three seasons of the year for browse, grass and a composite diet based
upon percentage of grass and browse used by bighorn sheep.

�-47-

.16 .15

-

A

•.140~

2'
§::

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e.

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.08° r0.07°r-

e

e

e

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e

a

e

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e

c:)

.02-r.0I-I-

,
S

I

i

•

I

I

I

F
GRASS

W

S

o Pikes Peak

I

F
BROWSE

-.l

I

I

I

I

I

I

I

W

S

F
W
COMPOSITE
DIET

o Buffalo Peak
~

Trickle Mtn.

Fig. 18. Parts per million selenium in forages from three study areas at three
seasons of the year for browse, grass and a composite diet based upon percentages
of grass and browse used by bighorn sheep.

�Table 6. Mean concentration and standard
sheep lambs collected on Pikes Peak.

1972
(4)

1973
(7)

of trace mineral levels in hair samples from bighorn

Percent
Calcium

Percent
Phosphorus

Percent
Sodium

Ppm
Iron

Ppm
Zinc

Ppm
Magnesium

Ppm
Copper

x

.16

.11

.04

287.2

112.5

381

80.7

Range

.15-17

.08-.14

.03-.05

172.4-615.4

91. 7-158.0

299-470

63.2-92.6

SD

.012

.025

.012

218.88

30.64

70.02

12.54

.20

.10

.05

269.4

105.2

323

89.1

.16-.29

.08- .13

.03-.10

190.5-470.9

94.3-121.3

254-390

69.3-122.~

SD

.052

.022

.024

99.36

9.88

44.20

19.30

x

.22

.11

.03

272.6

105.1

675

97.0

.16-.39

.08-.15

.02-.04

195.5-453.6

93.4-120.7

291-1600

74.7-148.5

.087

.024

.008

97.12

12.35

360.30

27.80

x

Range

197L1
(6)

deviation

Range
SD

I
.I:00
I

�Table 7. Mean concentration and standard deviation of trace mineral levels in liver samples from bighorn
sheep lambs collected on Pikes Peak.

1972
(4)

Percent
Calcium

Percent
Phosphorus

Percent
Sodium

Ppm
Iron

Ppm
Zinc

Ppm
Magnesium

Pp1!1
Copper

.08

.17

.24

333.3

162.3

474

15.9

.07-.09

.12-.24

.19-.29

302.8-378.4

134.5-190.9

458-490

9.4-23.6

SD

.008

.053

.055

33.88

23.39

17.1

5.95

x

.09

.16

.29

353.8

229.8

452

35.3

.08-.10

.10-.25

.20-.35

249.5-426.3

119.3-376.8

337-600

8.1-66.3

.009

.051

.057

64.74

102.79

105.0

27.15

.09

.17

.27

471.3

162.85

534

50.8

.07-.10

.10-.24

.20-.39

332.2-548.6

81.0-252.6

482-590

6.5-95.1

.012

.061

.067

74.59

73.8

35.20

x
Range

1973
(7)

Range
SD

1974
( 6)

x

Range
SD

146.08

I

~

'"
I

�Table 8. Mean concentration and standard
sheep lambs collected on Pikes Peak.

1972
(4)

deviation

of trace mineral levels in spleen samples

from bighorn

Percent
Calcium

Percent
Phosphorus

Percent
Sodium

Ppm
Iron

Ppm
Zinc

Ppm
Magnesium

Ppm
Copper

.09

.17

.24

333.3

99.1

546

5.4

.09-.09

.13-.20

.19-.29

302.8-378.4

74.9-120.3

495-592

4.9-5.5

SD

0.0

.030

.055

33.87

21.22

53.24

.300

x

.10

.32

.29

353.8

103.9

649.3

7.1

x
Range

1973
(7)

I
\J1

Range

1974
(6)

.08- .19

.24-.41

.20-.35

249.5-426.3

89.5-115.6

578-702

4.8-13.0 :

SD

.039

.061

.057

64.74

10.01

41.90

2.74

x

.11

.28

.27

471. 3

105.1

773.5

6.3

.08-.15

.20-.35

.20-.39

332.2-548.6

98.5-107.1

549-1127

4.9-8.5

.030

.065

.067

146.08

6.05

225.46

1.35

Range
SD

0
I

�Table 9. Mean concentration and standard deviation
sheep lambs collected on Pikes Peak.

of trace mineral levels in kidney samples from bighorn

Percent
Calcium

Percent
Phosphorus

Percent
Sodium

Ppm
Iron

l'pm
Zinc

Ppm
Magnesium

Ppm
Copper

.10

.25

.57

356.5

105.8

565

19.5

.08-.11

.11-.42

.30-.78

302.4-396.1

88.0-112.5

511-633

17.9-20.8

SD

.015

.l36

.202

39.81 .

11.89

54.2

1.21

x

.11

.32

.72

359.3

116.5

606

20.6

1972
(4)

x

Range

1973
(7)

Range

1974
(6)

.10-.12

.24-.40

.59-.99

297.6-403.2

76.1-168.9

571-673

15.3-22.7

SD

.008

.053

.148

42.91

33.45

37.2

2.75

x

.11

.33

.61

356.1

123.2

615

21.3

.09- .15

.22-.42

.41-.70

283.7-449.4

89.9-222.5

519-866

19.7-23.3

.023

.076

.104

66.16

50.97

128.9

1.50

Range
SD

I
\Jl
f-'

I

�Table 10. Mean concentration and standard deviation
from bighorn sheep lambs collected on Pikes Peak.

1972
(4)

Percent
Calcium

Percent
Phosphorus

Percent
Sodium

Ppm
Iron

Ppm
Zinc

Ppm
Magnesium

Ppm
Copper

.07

.18

.38

232.6

118.3

756

7.9

.06-.08

.16-.19

.17-.22

226.6- 241. 9

105.9-131. 8

716-775

6.4-9.5

SD

.010

.015

.243

10.24

13.60

27.09

1.68

x

.09

.21

.26

245.1

149.8

724.6

9.5

x
Range

1973
(7)

Range

1974
(6)

of trace mineral levels in skeletal muscle samples

.07- .12

.18-.28

.18-.46

221.3-288.6

SD

.016

.036

.104

x

.11

.26

.09-.17
.032

Range
SD

91.7-193.7

632-844

24.30

38.81

91.61

1.66

.22

271.3

124.8

9l7.7

9.5

.21-.30

.17-.29

242.1-311. 2

58.7-193.3

704-1419

.033

.044

25.37

48.36

255.54

7.5-12.2

7.3-15.0
3.01

I
\J1

N
I

�Table 11. Mean concentration and standard deviation
from bighorn sheep lambs collected on Pikes Peak.

1972
(4)

of trace mineral

levels in heart muscle samples

Percent
Calcium

Percent
Phosphorus

Percent
Sodium

Ppm
Iron

Ppm
Zinc

Ppm
Magnesium

Ppm
Copper

.09

.24

.37

323.6

74.0

747

19.5

.08- .10

.21-.29

.30- .44

256.6-385.3

71.7-79.8

692 -860

17.9-20.8

SD

.008

.036

.058

52.80

3.88

76.58

1.21

x

.10

.29

.39

323.9

80.5

772

20.6

.09-.12

.25-.36

.36-.44

281.6-360.2

76.9-90.9

635-958

15.3-22.7

SD

.011

.041

.037

27.98

5.35

131.38

2.75

x

.10

.28

.35

342.3

85.38

862.8

21.3

.09-.12

.23-.32

.26-.44

319.0-379.4

72.2-129.1

653-1023

19.7-23.3

.013

.036

.070

23.01

21. 76

210.82

x
Range

1973
(7)

Range

1974
(6)

Range

SD

1.50

I
VI
LV
I

�-54-

LITERATURE

CITED

Adrian, W. J. 1973. A comparison of a wet pressure digestion method
with other commonly used wet and dry-ashing methods. Analyst
98:213.
Baker, E. W., and B. W. Wharten.
1952.
MacMillan Co., New York. 465 p.

An introduction

to acarology.

Baumgartner, L. L., and A. C. Martin. 1939. Plant histology as an aid
in squirrel food habit studies. J. Wildl. Manage.
3:266-268.
Bear, G. D., and R. M. Hansen. 1966. Food habits, growth and reproduction
of white-tailed jackrabbits in southern Colorado. Colo. Agri. Exp.
StaG Tech. Bull. No. 90. 59 p.
Boussingault, Jean-Baptiste.
1843. Economic rurale considderee dans ses
rapports avec la chimie, la physique et la meteorologie.
Bechet
Jeunne. Paris.
(In 2 volumes). Translated by Sir George Laws.
1845. Rural economic in its relation with chemistry, physics and
meterology.
D. Appleton and Company, New York.
Colburn, M. W., and J. L. Evans. 1967. Chemical composition of the cell
wall constituents and acid detergent fiber fraction of forages. J.
Dairy Science 50:1130.
Cook, C. W., and L. E. Harris, 1950a. The nutritive content of the
grazing sheep's diet on summer and winter ranges of Utah. Utah Exp.
StaG Bull. 340.
Cook, C. W., and L. E. Harris. 1950b. The nutritive value of range forage
as affected by vegetation type, site, and stage of maturity.
Utah
Tech~ Bull. 344.
Cook, C. W., L. E. Harris, and L. A. Stoddart. 1948. Measuring the
nutritive content of a foraging sheep's diet under range conditions.
J. Anim. Sci. 7:170.
Cooperrider, A. Y. 1969. The biology and management of the bighorn sheep
of Rock Creek, Montana. Montana Game and Fish Dept. Fed. Aid Rept.
92 p.
Crampton, E. W., and I.R.C. Jackson. 1944. Seasonal variation in chemical
composition of pasture herbage and the relation to its digestibility
by steers and sheep. J. Animal Sci. 3:333.
Crampton, E. W., and L. A. Maynard.
1938. The relation of cellulose
and lignin content to the nutritive value of animal feeds. J.
Nutrition 15:383.
Dusi, J. L. 1949. Methods for the determination of food habits by
plant microtechniques and histology and their application to cottontail rabbit food habits. J. Wildl. Manage. 13:295-298.
Fonnesbeck, P. V., and L. E. Harris.
1971. Determining hemicellulose
in plant cell walls. Proc. Western Sec., Amer. Soc. Animal Sci.
22:77.

�-55-

Fonnesbeck, P. V., and L. E. Harris. 1970a. Determination of plant cell
walls in feeds. Proc. Western Sec., Amer. Soc. Animal Sci. 21:153.
Fonnesbeck, P. V., and L. E. Harris. 1970b. Determination of holocellulose, lignin and silica of plant cell walls. Proc. Western
Sec., Amer. Soc. Animal Sci. 21:162.
Garrigus, W. P., and H. P. Rusk. 1939. Some effects of the species and
stage of maturity of plants on the forage consumption of grazing
steers of various weights. Ill. Agr. Exp. Sta. Bull. 454.
Harrington, H. D. 1954.
Denver. 666 p.

Manual of the plants of Colorado.

Sage Books,

Harris, L. E. 1970. Nutrition techniques for domestic and wild animals.
Vol. 1. An international record system and procedures for analyzing
samples. L. E. Harris, Logan, Utah.
Harris, L. E., C. W. Cook, and J. E. Butcher. 1959. Symposium on forage
evaluation. V. Intake and digestibility techniques and supplemental
feeding in range forage evaluation. Agron. J. 51:226.
Hegg,

o. 1961. Analyses of big-game droppings to determine their dietary
composition in Swiss National Park. Revue suisse de 800logie 68(12):
156-165. (Translated by J. J. Stansky, U.S.D.A. Forest Service,
Southern Forest Experiment Station. 9 p.).

Henneberg, W., and F. Stohmann.
fritterung der weiderkur.

1860.

Henry, W. A., and F. B. Morrison.
1910.
W. A. Henry, Madison, Wisconsin.

Berundung einer rationellen

Feeds and Feeding.

10th ed.

Kiley, M. 1966. A preliminary investigation into the feeding habits of
the waterbuck by faecal analysis. Journ. E. Africa Wildl. 4:153-157.
Lay, D. W. 1965. Fruit utilization by deer in southern forests.
Wildl. Manage. 29(2):370-375.

J.

Martin, C. M. 1952. Seasonal changes in the physical and chemical
composition of pasture herbage and their effects upon the responses
of grazing and hand-fed steers. Ph.D. Thesis, Cornell University.
Moser, C. A. 1962. The bighorn sheep of Colorado.
Dept. Tech. Bull. No. 10. 49 p.

Colo. Game and Fish

Muth, O. H., J. E. Oldfield, L. F. Remmert, and J. R. Schubert.
Effects of selenium and vitamin E on white muscle disease.
128:1090.

1958.
Science

Myers, G. T. 1962. Food habits of the plains pocket gopher in the sandhills of eastern Colorado. M.S. Thesis, Colorado State Univ., Fort
Collins. 73 p.
Reid, J. T., W. K. Kennedy, K. L. Turk, S. T. Slack, G. W. Trimberger, and
R. P. Murphy. 1959a. Effect of growth stage, chemical composition,
and physical properties upon the nutritive value of forages. J.
Dairy Sci. 42:567.

�-56Reid, J. T., and W. K. Kennedy.
1959b. Symposium on forage evaluation.
I. What is forage quality from the animal standpoint?
Agron. J.
51:213.
Reid, J. T., P. G. Woolfolk, C. R. Richards, R. W. Kaufman, J. K. Loosli,
K. L. Turk, J. I. Miller, and R. E. Blaser. 1950. A new indicator
method for the determination of digestibility and consumption by
ruminants.
J. Dairy Sci. 33:60.
Shepherd, H. R.

1961.

Colorado Div. of Wildlife,

Smith, E. L. 1962. In Mineral Metabolism,
Press. pp. 349-369.

personal communication.

Vol. 2, Part B.

Academic

Sparks, D. R., and J. C. Malechek.
1968. Estimating percentage dry
weight in diets using a microscope technique.
J. Range Manage.
21(4):261-265.
Sparks, D. R. 1968. Diet of black-tailed jackrabbits
land. J. Range Manage. 21(4):203-208.

on sandhill range-

Stewart, D. R. M. 1967. Analysis of plant epidermis in faeces: a technique
for studying the food preference of grazing herbivores.
J. Appl.
Exol. 4(1):83-111.
Storr, G. M. 1961. Microscopic analysis of faeces, a technique for
ascertaining the diet of herbivorous mammals. Austral. J. BioI. Sci.
14:157-164.
Thaer, A. 1809. Grundsatze der rationelle landwirtschaft.
275, Die Realschulbuchhandlung.
Berlin.

Vol. I, Sec.

Todd, J. W. 1972. Food habits of the Rocky Mountain bighorn sheep.
Thesis. Colorado State Univ., Fort Collins.
Van Soest, P. J., and R. H. Wine. 1967. Use of the detergents in the
analysis of feeds. IV. Determination of plant cell-wall constituents.
J. Assoc. Off. Anal. Chem. 50:50.
Van Soest, P. J. 1964. Symposium on nutrition and forage and pastures:
New chemical procedures for evaluating forages. J. Animal Sci.
28:838.
Van Soest, P. J. 1963. Use of detergents in the analysis of fibrous
feeds. II. A rapid method for the determination of fiber and
lignin. J. Assoc. Off. Agr. Chem. 46:829.
Zyznar, E., and P. J. Urness. 1969.
forage ruminants in deer feces.

Prepared by:
Robert E. Keiss
Wildlife Researcher

Qualitative identification of
J. Wildl. Manage. 33(3):506-510.

�-57-

JOB PROGRESS

State

0f

WOLk Plan No.

Period

REPORT

W-41"":R-26

Bighorn

1

Job No.

----------------

Manipulation

Coyered:

Personnel:

1977

C.:...O.:...I.:....O:c;:RAD=:...:O'-_

.Proj ect No.

Job Title

January

of Vegetation

Sheep

&amp;

Mourita·in Goat Investigations

17

-------------------------------

on Bighorn

Sheep Ranges

June 1, 1975 to May 31, 1976

George D. Bear, Lynn Stevens,
Stan Koster.

Karl Karrow,

Patrick

Powell,

and

ABSTRACT
Study plots near Cathedral, Colorado were treated with varied applications of
nitrogen and phosphorus fertilizers, and2,4-D
herbicide in 1971-72.
Two areas
were selected for study sites: (a) an alpine range, and (b) a ponderosa pinebunchgrass range.
The effects of these treatments were monitored during the
period 1972-1976.
Data are being analyzed and a final report prepared.

��-59-

MANIPULATION OF VEGETATION ON BIGHORN SHEEP RANGES
George D. Bear

P. S. OBJECTIVE
Improve the herbage yield, vegetative density and vegetative composition
on selected bighorn sheep ranges in Colorado.

SEGMENT OBJECTIVES
1.

Determine the effect of each treatment on the composition, production
and chemical content of plant species.

2.

Determine the grazing preferences of wild bighorns on the area in
respect to each treatment.

PROCEDURES
The study area is located about 35 miles south of Gunnison, Colorado.
This bighorn herd winters in two areas; in a low (8,000 feet elevation)
bunchgrass and ponderosa pine type, and in a high (11,000-14,000 feet
elevation) alpine vegetative type. Study plots were established on these
two winter ranges; and the plots were treated with phosphorus fertilizer,
nitrogen fertilizer, and 2,4-D herbicide (16 treatment-levels) in 1971-72.
Effects of these treatments we~e monitored from June, 1972 to June, 1976.
Data are being analyzed (multiple analysis of variance) at Colorado State
University Statistical Center, and a final report is being prepared.
Permanent transects were established on each of the plots to determine
change in vegetative composition and production which may result from the
various treatments. A frequency sampling technique was used to monitor
vegetative composition on the plots. Quantity of vegetation produced on
the treatment plots was determined by using a Neal Electronics Herbage
Meter.
Vegetative samples were collected from each treated area and analyzed for
their chemical content. Only samples of the most common plant species were
collected. Since the study areas are bighorn winter ranges, the vegetation
was sampled at the completion of the growing season, in mid-winter, and
late winter.
Animal reaction to the treatments or selectivity was determined by direct
observation of grazing animals and visual reconnaissance of utilization
on the vegetation.

Prepared by

~..ft#;. cfI

~-L~.

Ceor s D. Bear
Wildlife Researcher

��January, 1977
-61JOB PROGRESS

State of

REPORT

COLORADO
--------~~~~----~-----W-41-R-26

Proj'ect No.
Work Plan No •
.Joh Title

Job No.~.__ ~_2_1

~

_

Trapping, Marking and Collecting Bighorn Sheep

Period Covered:
Personnel:

1

Bighorn Sheep &amp; Mountain Goat Investigations

June 1, 1975 - May 31, 1976

G. D. Bear, F. CoLl.ey , M. P. Elkins, C. P. Hfb Ler , R. E. Keiss,
E.Latson,
J. Morris, P. H. Neil, R. L. Schmidt, T. Spraker, and
J. Wegryzn.

ABSTRACT ..
Fifty-two bighorn sheep were trapped during the winter of 1976 by the use of
a drop net trap. Most of· these captured animals were treated for Protostrongylus
Several bighorn sheep were transported
sp. infestation, marked rand released.
to holding facilities for more intensive research investigations,

��-63-

TRAPPING, MARKING AND COLLECTING BIGHORN SHEEP
Robert L. Schmidt

P. S. OBJECTIVE
To trap or collect bighorn sheep for adding to the captive flocks at
Little Hills Experiment Station; for obtaining nasal swabs and blood
samples; for marking and treatment with therapeutic drugs, or (in case
of dead animal collection) for necropsy examination.

SEGMENT OBJECTIVE
1.

Collect bighorn lambs at intervals and locations prescribed
tractual agreement.

by con-

2.

Trap an indefinite number (as many as can reasonably be caught and/or
handled) of bighorn sheep of all age classes.

METHODS AND MATERIALS
Trapping techniques used during this segment are the same as those previously
described (Erickson 1970, Schmidt 1976). A drop net, triggered by remote
control, is utilized to capture the bighorn sheep which are attracted under
the net by apple pulp and alfalfa.
The bighorn sheep were treated with chemotherapeutic drugs showing efficacy
for Protostrongylus sp. After treatment they were marked with either eartags on lambs or neck bands on adults and released. Drugs used in this treatment scheme were Cambendazole (8.4 gms/dose) and Diethylcarbamazine
(3.2 gms/
dose) •
Bighorn sheep were trapped at two locations - the Cache la Poudre Canyon west
of Fort Collins, and the Pikes Peak area west of Colorado Springs, Colorado.
Tables 1 and 2 list the animals trapped, treated and released from the two
areas.

LITERATURE

CITED

Erickson, ~. A. 1970. Use of drop net and collars in a study of Dall sheep.
North Wild Sheep Council, Trans. pp. 20-21.
Schmidt, R. L. 1976. Trapping, marking and collecting bighorn sheep. Colo.
Div. Wildl. Fed.
1, Job 21, Job Prog. Rep.,
Game Res. Re

Wildlife Tech II

�-64-

�-65-

Table
Study

2.
Bighorn
Area.

sheep

trapped,

Age

No.

Date

Sex

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35

3/11/76
3/11/76
3/11/76
3/11/76
3/11/76
3/11/76
3/11/76
3/11/76
3/11/76
3/11/76
3/11/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
3/22/76
4/1/76
4/1/76
4/1/76
4/1/76
4/1/76
4/1/76
4/1/76
4/1/76
4/1/76

F
F
F
F
F
F
F
F
F
M

F
F
F
F
F
F
F
M

F
F
M

F
M
M
M

F
F
F
F
F
F
F
M
M

F

treated

New
Neck Band

and released

Old
Neck Band

from the

Pikes

Eartag

Blue R 1
7
Blue R 2
4
Blue R 3
8
White. 3
Blue R 4
8
Blue R 7
5
Blue R 8
8
Yellow. 11
10
Blue R 10
Lamb
Yellow 1
Lamb
Yellow 2
Lamb
Yellow 3
8 (Lost in Trapping)
7
Blue Rll
9
Blue R 12
Yellow. 9
8
Blue Rl3
Blue. 3
9
Blue R 14
Yellow. l3
8
Blue R 15
7
Blue R- 1
Lamb
Yellow 4
Lamb
Yellow 5
Lamb
Yellow 6
Lamb
Yellow 7
Lamb
Yellow 8
Lamb
Yellow 9
Lamb
Yellow 10
1
Yellow 12
Lamb
Yellow 1 (E. T.)
9
Blue R 17
8
Blue R 18
9
Blue R 20
8
Blue R 21
Blue. 1
8
Blue R 22
7
Blue R 23
Lamb
Yellow 11
Lamb
Yellow 12
(Lost in Trapping)
7

Peak

Weight

1751/
1651/
1601/
107fl

1751/

��January 1977

-67JOB PROGRESS

State of
Project

REPORT

_ COLORADO
W-4l-R-26

No.

1

Work Plan No.
Job Title

Monitoring

Period Covered:
Personnel:

Bighorn Sheep &amp; Mountain
Job

t;ro.

Goat Investigations

22

-~--------------------------

Bighorn Sheep Populations

.--------------------------

June 1, 1975 - May 31, 1976

F. Colley, M. P. Elkins, C. P. Hibler, J. Morris, P. Neil, and
R. 1. Schmidt.

ABSTRACT
Bighorn sheep in the Pikes Peak Study Area, the Cache la Poudre Canyon Study
Area, and the Rampart Range Study area were treated with chemotherapeutic drugs
showing efficacy for Protostrongylus sp. during the study year.
Evaluations of the effects of these treatments have been made and indicate
that lamb survival has been enhanced by drug treatments as indicated by
improved ewe-lamb ratios on Pikes Peak.

��-69-

MONITORING BIGHORN SHEEP POPULATIONS
Robert L. Schmidt

P. S. OBJECTIVE
To monitor bighorn sheep by visual surveillance
characteristics of lamb mortality.

to further elucidate the

SEGMENT OBJECTIVES
1.

Monitor bighorn sheep numbers, age and sex composition,
appearance, behavior and activity.

general body

2.

Monitor numbers, distribution, association with other sheep, and lamb
births and survival, of bighorn sheep which have been trapped, treated
with therapeutic drugs, neck collared, and released at the trap site.

METHODS AND MATERIALS
All of the information collected concerning numbers, age and sex composition,
and lamb survival of bighorn sheep populations in general was gathered on
populations which were being monitored for evaluation of drug treatment effects.
These results will be presented along with the results of drug treatment evaluations. In addition a simple Lincoln Index was used to estimate size of the
current Pikes Peak bighorn sheep population.

Chemotherapeutic

Drug Treatments

(1976)

Pikes Peak
Bait treatment stations established and used in drug treatments on Pikes Peak
are shown in Figure 1. This figure shows the locations of several different
bait-treatment stations where bighorn sheep were treated with chemotherapeutic
drugs during the winter of 1976. Chemotherapeutic drugs were incorporated into
apple mash as previously reported (Schmidt 1976) except that the amount of drug
for a single sheep dose was mixed with three pounds of mash instead of the five
pounds utilized in the past.
Table 1 lists the bait-treatment stations, the dates chemotherapeutic drugs
were administered, the number and identity of bighorn sheep consuming drug
impregnated apple mash. Chemotherapeutic drugs used during this segment
included Cambendazole (8.4 gm/dose) and Diethylcarbamazine (3.2 gm/dose).

�Manitou Springs

Dome Rock
7.~

#
Sentinel
Point
'4 •

5.~
Knob

,.,.Sheep
Mtn.
I

-....J

o
I

Bait Site
Fig. 1.

Locations of drug treatment bait stations, Pikes Peak Area, 1976.

#1 - Saddle; #2 - Gravel Pit; #3 Peak; #4 Tunnel; #5 East Beaver; #6 Four Mile Road; #7 Four Mile Bait.

�I

-71-

Table 1. Bait treatment stations on the Pikes Peak Study Area; treatment
and bighorn sheep receiving drug treatment.

Bait Treatment
Station

Date
Treated

Ifl "Saddle"

2/19/76

3/9/76

112 "Gravel Pit"

2/19/76

3/9/76

113 "Peak"

2/19/76

3/9/76

114 "Tunnel"

3/13/76

Bighorn Sheep
Treated

No.

dates,

Identification

Lambs

4

Yearlings

2

Uncollared

4

Collared

5

Lambs

3

Yearlings

1

Uncollared

12

Collared

10

Lambs

3

Yearlings

1

Uncollared

6

Collared

2

Lambs

11

Yearlings

3

Uncollared

9

Collared

4

Lambs

11

Yearlings

1

Un collared

1

Collared

9

Lambs

4

Yearlings

0

Uncollared

1

Collared

2

(Bd12; 09)

Lambs

3

(et Y4; et Y8; et Y11)

Yearlings

0

Uncollared

2

Collared

8

(Y-3; Yd5; Yd6; Wd3;
Bdl)

(Wd2; Wd3; Wd6; Yd5;
Yd15; Ydll; Bdl;
Bd4; Rdl; 0-10)

(Bd2; Wd2)

(Ydl4; Yd9; Bd3; Od6)

(Yd11; Yd13; Yd9; Yd14;
Od6; 09; Rdl; Wd6;
Bd3)

(0 10; 09; Wd4; Yd15;
Yd16; Bd12; BRl2*;
BRl3**)

------------------------------------------------------------------------------

�-72-

Table 1. Bait treatment stations on the Pkes Peak Study Area; treatment
dates, and bighorn sheep receiving drug treatment (Cont.).

Bait Treat men t
Station

Date
Treated

Bighorn Sheep
Treated

115 "East Beaver"

3/13/76

Lambs

No.

Twelve treatments were
placed on the bait
station. no observations
were made due to a
snowstorm

Yearlings
Uncollared
Collared
116 "Four Mile
Road"

4/7 /76

4/15/76

4/16/76

117 "Four Mile
Bait"

4/6/76

4/15/76

Identification

Lambs

0

Yearlings

4

Uncollared

0

Collared

4

Lambs

2

Yearlings

4

Uncollared

1

Collared

3

Lambs

2

Yearlings

2

Un collared

1

Collared

4

Lambs

2

Yearlings

3

Uncollared

3

Collared

6

Lambs

0

Yearlings

2

Uncollared

1

Collared

4

(Yd17; Yd20; Yd21; B4)

(Yd17; Yd20; M)

(Yd17; Yd21; Rd8; B-?)

(Rd7; Rd8; B5; W16;
W17; Yd19)

(B5 ; B2; B d5; W15)

B - Blue; Y - Yellow; W - White; R - Red; d - Dot; et - Ear Tag; 0 - Orange

* BR12 was previously

Yd9.

**BRl3 was previously Bd3.

�-73-

Poudre Canyon
Bait-treatment stations established and used in drug treatments in the
Cache la Poudre Canyon are shown in Figure 2. This figure shows the
location of four different bait-treatment stations where bighorn sheep
were treated with chemotherapeutic drugs. All treatments are the same
as described for Pikes Peak Study Area.
Table 2 lists the bait-treatment stations, the dates chemotherapeutic
drugs were administered, the number and identity of bighorn sheep consuming drug impregnated apple mash.

Rampart Range
One bait-treatment station was established at the Rampart Range gravel
pit located west of Colorado Springs, Colorado. Baiting techniques were
the same as described for Pikes Peak and Poudre Canyon Study Areas except
that a different drug was used. Fenbendazole was administered in apple
mash at several dosage rates. Table 3 lists the treatment schedule
followed on the Rampart Range during the winter of 1976.

Pikes Peak Bighorn
Sheep Population Estimates 1975-1976
Population estimates made from data in this segment reflect the drug treatment results from the winter of 1974-1975 as reported in the previous segment
of this project (Schmidt 1976).
Composited observational data on the proportion of collared vs uncollared
bighorn sheep observed over the period June 6, 1975 to April 6, 1976 were
used to estimate the size of the Pikes Peak bighorn sheep population. A
simple Lincoln Index was used to project these data to population estimates.
Since all sheep which were collared were adults, population projections
were pertinent only to adults. Proportions of yearlings and lambs in classification data were used to project to final estimates of total populations.

RESULTS AND DISCUSSION
Chemotherapeutic

Drug Treatments

(1975)

Pikes Peak
There are two parameters which can be measured in a wild bighorn sheep
population to evaluate the efficacy of chemotherapeutic drugs on bighorn
sheep lamb survival. One is to measure total herd recruitment and hope for
an increase in total numbers of animals on a sustained year to year basis,
and second to measure the number of first stage Protostrongylus larvae
deposited in the fecal pellet groups. If chemotherapeutic drugs are to be
effective in controlling adult lungworms in the air passages of the animal,
then it should be reflected in the number of larvae these adults worms will
produce. During the present segment, pellet groups are being collected and
Baermannized to count first stage larvae and these data will be presented
during the next segment of the project.

�~
~

N. BALD MTN.
MIDDLE BALP MTN.

i) BALD MTN.

f1'if

3.

LONETREE MTN.
-...JI

-'I"

§
~

e

'g

••••

,,0
•......

-- •..•.

CROWN POINT

• = Bait Site
Fig. 2. Location of bait-treatment stations in the Poudre River Canyon, Colorado.
#1 - Meadows; #2 - Poudre Ponds; #3 - New Arrow Head; #4 - Indian Meadows.

�-75-

Table 2. Bait treatment stations in the Poudre River Canyon study area,
treatment dates, and bighorn sheep receiving drug treatment.

Bait Treatment
Station

Date
Treated

Bighorn Sheep
Treated

No.

III "Meadows"

3/1/76

Lambs

3

Yearlings

3

Uncollared

3

Collared

11

(Yl; Y2; Y3; Y4; Rl;
R3; B4; B16; B20; WR2)

Lambs

3

(et Rl; et Bl)

Yearlings

3

Uncollared

2

Collared

8

Lambs

Uncollared

o
o
o

Collared

3

(WG; OB; R5)

Lambs

4

(et Y2; et Y3; et Y9)

Yearlings

1

(et Y12)

Uncollared

o

Collared

3

Lambs

1

Yearlings

o

Uncollared

6

Collared

2

(Bl3; Bll)

Lambs

5

(et Y5)

Yearlings

o

Uncollared

13

Collared

3

Rams

9

3/14/76

112 "Poudre Ponds" 3/3/76

Yearlings

3/29/76

113 "New Arrow
Head"

3/18/76

3/29/76

114 "Indian
Meadows"

3/3/76

Identification

(et Rl)

(Y2; Y3; Y4; R3; R8;
B20; WR2; RW2)

(WG; OB; RS)

(Bl; B10; Bl3)

�-76-

Table 3. Bait treatment station on Rampart Range study area, treatment
and dosage rates of Fenbendazole for bighorn sheep.

Treatment
Date

Fenbendazole
Dosage

3/31/76

5 mg.

4/6/76

5 mg.

4/15/76

Note:

20 mg.

Bighorn Sheep
Treated

No.

Lambs

1

Yearlings

2

Unco Ll.a re d

2

Collared

7

Lambs

1

Yearlings

2

Un collared

2

Collared

7

Lambs

1

Yearlings

2

Uncollared

2

Collared

7

On 4/14/76 eight rams and one uncollared

Cambendazole.

dates,

Identification

(et R7; et R9)

(Y6; Y7; YB; Y9;
Yll; Yl4; R2)

(et R7; et R9)

(Y6; Y7; YB; Y9;
Yll; Yl4; R2)

(et R7; et R9)

(Y6; Y7; YB; Y9;
Yl1; Y14; R2)

ewe were treated with 8.1 gm.

�-77-

Bighorn sheep population estimates for the Pikes Peak Study Area are based
upon the following observation counts made between May 1975 and April
1976 :
Collared Ewes
Uncollared Ewes
Collared Rams
Uncollared Rams

329
231
23
31

By direct observations it was determined that a total of 46 collared ewes
and 8 collared rams were known to be on Pikes Peak, therefore it is possible
to use a simple Lincoln Index to estimate the adult bighorn population.

Ewes (X)

46 x 560
329

78

8 x 54
Rams (Y) = --'-";;';;2-'3:""";""19
By using population classification data derived by observations made between
September 1975 and April 15, 1976, it was estimated that the ewe-lamb ratio
was 61 lambs per 100 ewes. Thus if we estimate the ewe population to be
78 then it is reasonable to estimate the lamb population at 48.
Two different estimates of the number of yearlings in the Pikes Peak population can be calculated. First, yearling numbers can be calculated by
assuming that the ratio yearling observations to total adult ewe observations reflect the true population ratio of yearlings to adult ewes (i.e.
there are no cumulative observability or identifiability biases which favor
the detectability of one group over the other). Then yearling numbers can
be estimated by multiplying the yearling to adult ewe ratio (130) by the

3W

estimated number of adult ewes (78), thus, number of yearlings
78 x

=

~~g= 18.

Secondly, yearling numbers can be calculated by assuming that the ratio of
yearling observations to adult ram observations (130/54) reflect the true
population ratio of yearlings to adult rams. An estimate of yearling
numbers is then obtained by multiplying the estimate of adult rams (19) by
the yearling to adult ratio. Thus, number of yearlings = 19 x 130 = 46.
54
Another comparison made on lamb survival is the ewe-lamb ratio between
treated and untreated ewes within' the Pikes Peak Study Area population.
Table 4 shows the data comparison between the ratios of ewes to lambs
during three periods in the study area.

�Table 4. Comparison
three time intervals

of ewe-lamb ratios between collared-treated
during the 1975-1976 study year.

Time Period

June-August

1975

September-December
January-March

1976

1975

ewes and collared-untreated

ewes during

Untreated
Ewes

Lambs

Ewe/Lamb
Ratio

Treated
Ewes

Lambs

Ewe/Lamb
Ratio

30

16

53:100

73

59

81:100

30

15

50:100

88

64

73:100

9

2

22:100

38

28

74:100

I

--.J
00
I

�-79-

LITERATURE

CITED

schmidt, R. L. 1976. Trapping, marking and collecting bighorn sheep.
Colo. Div. Wildl. Fed. Aid W-4l-R-25, Work Plan 1, Job 21, Job
Prog. Rep., Game Res. Rep. January. pp. 49-54.

Prepared

by

~LijP
Rob rt L. chmidt
Wildlife Tech II

��-81-

JOB PROGRESS

State of
Project

1977

REPORT

---.,;C::.;O:.:LO:;.::..;::RAD=..;;.O
_
W-4l-R-26

No.

Work Plan No.
Job Title
Period

January,

1

Sheep

Job No.
Bighorn Sheep Lungworm Control
By Biological Control of Snail Intermediate

Covered:

Personnel:

Bighorn

Harold

&amp; Mountain

Goat Investigations

~23~
Hosts

june 1, 1975 - May 31, 1976
R. Shepherd

ABSTRACT
Preliminary plans were made with the United States Forest Service and
Colorado State University to conduct and monitor prescribed burning of
aspen patches in a portion of the Poudre River bighorn sheep range.
Subsequent to tentative Forest Service approval of the proposed study,
a detailed plan, the Program Narrative Outline; was prepared to conduct
a study to learn th~ effects of prescribed burning of aspen on landsnail abundance.

_

��BIGHORN SHEEP LUNGWORM CONTROL
BY BIOLOGICAL ,CONTROL OF SNAIL INTERMEDIATE HOSTS
Harold R. Shepherd

P. S. OBJECTIVE
To evaluate and develop methods for field application of pesticides
and/or biological control mechanisms that have been laboratory tested
and found to be effective in controlling snails and/or lungworm larvae.

SEGMENT OBJECTIVES
1.

Prepare a literature review of land snails, including information
about their predators, parasites, diseases, life cycles and natural
or imposed biological controls.

2.

Conduct suitable reconnaissance field and/or laboratory surveys preliminary to the search for means of biologically controlling land
snails.

3.

Plan a study of the life cycles and the predators, parasites, diseases,
and habitat relationships of the species of snails which are intermediate hosts of lungworms in bighorn sheep.

RESULTS AND DISCUSSION
Arrangements were made with Biological Information Service Company of Los
Angeles for computerized retrospective literature searches under the main
subjects of Bighorn Sheep, Lungworms, and Snails. Upon receipt of titles
and some abstracts generated by the literature searches, pertinent literature was selected, obtained, read and abstracted.
Abstracts were placed
on Unisort Analysis cards and filed according to appropriate subject
headings.
A review of the literature suggests that land-snail control using natural
snail enemies or molluscicides is presently impractical.
However, snail
and prescribed burning literature and exploratory snail habitat sampling
of Poudre River bighorn sheep range provides reasons to believe that concentrations of snails occurring in aspen patches on sheep range may be
effectively reduced by prescribed burning of aspen patches. Such habitat
manipulation, if done periodically, might reduce the incidence of lungworm
infection in sheep.
Because the prescribed burning study envisaged would be on U.S. Forest
Service lands, the permission of the Forest Service was desirable before
further effort and expense was expended in planning.
Consequently, several
conferences and field trips were arranged with appropriate Forest Service
personnel to acquaint them with the general objectives of the proposed

�-84study and the sites where it would be undertaken.
As a result of the
meetings, the Forest Service gave tentative approval of the study
contingent upon approval of the final plans.
Personnel of the Colorado State University Fire Ecology Department have
agreed to cooperate in the study to the extent of preparing prescriptions
for the bu-rning procedures and pre- and post-burn monitoring of burn
parameters.
A Program Narrative Outline was prepared and submitted to my immediate
supervisor for approval. It constitutes a review of literature pertinent
to the study discussed above, together with detailed procedures, work
schedules and budget estimates sufficient to permit others, if necessary,
to successfully conduct the study.

Prepared by

t?~kM

-: i,~
Harold R. Shepher
.Wildlife Researcher

�-85-

January, 1977
JOB PROGRESS REPORT

State of

COLORADO
--------~~---------------

Project No.

W-4l-R-26

Work. Plan No.

1

Bighorn Sheep &amp; Mountain .Goat Investigations
Job No.

~2~4~ __ ~

~----------

Job Title __~E=x~t~e=n=d=i=n~g~B=i~gh~o~rn=-~Sh=e~e~·
p~R~a=n~g~e~s~
Period Covered:
Personnel:

_

June 1, 1975 to May 31, 1976

George D. Bear, George Jones, Cliff Coghill, Robert E. Keiss.

ABSTRACT
Twenty-five bighorn sheep were transplanted to the lower portion of the
Cache la Poudre River drainage during the last work segment.
It appeared
that sixteen sheep had settled on .the Sheep Mt. release site. They remained
in the area until November, when deer hunters apparently caused them to disperse. Seven.reappeared on the area. only to leave again ::tnlate December.
These bighorns are very nomadic now, wandering around in the lower portion of
the canyon.
Nine bighorns (6 ewes, 1 lamb, and 2 rams) were transplanted to the Cebolla
Creek Wildlife Area last winter.
These b Lghorns remained at the release-site
until the snow melted, then they started exploring the surrounding area.
This band of sheep is also very noniadic,but are still within six miles of
the release-site.

��-87-

EXTENDING BIGHORN SHEEP RANGES
George D. Bear

P. S. OBJECTIVE
Determine if the distribution and population of bighorn sheep herds can
be increased by transplanting sheep into suitable areas adjacent to the
present concentrations.

SEGMENT OBJECTIVES
1.

Trap and transplant sheep onto the Cebolla Creek Wildlife Area.

2.

Monitor survival, movements, distribution and reproduction of transplanted sheep in Cache la Poudre Canyon and Cebolla Creek Wildlife
Area.

METHODS AND MATERIALS
Cache la Poudre Canyon
During the previous segment (1974-75) twenty-five bighorn sheep were
trapped in the upper Cache la Poudre River drainage and transplanted to
the lower portion of the canyon (Sheep Mt.). The sheep moved around a
great deal during the first thirty days after their release. Some individuals remained together, while others moved between groups. By spring
(April-May, 1975) it appeared that sixteen sheep had settled on the Sheep
Mt. release-site.
Continuing efforts were made during this segment to
record movements of the transplanted bighorns. This primarily consists of
walking through the areas occupied by the sheep and recording the location
of the individuals observed.

Cebolla Creek
A similar transplant was made in the LaGarita Mountains. The LaGarita
bighorn herd winters in two distinct areas; in the alpine range above
12,000 feet elevation, and on the Cebolla Creek Wildlife Area at approximately 8,000-9,000 feet elevation. Information gathered from local residents
indicates that in the early 1900's it was common for these sheep to migrate
from the alpine range down to Cebolla Creek for the winter. However, since
the die-off in the 1950's, only a few rams have migrated to the lower range
during the winter months, while the lambs and ewes have remained in the high
country. During the past two winters this migratory segment of the ram
population has been reduced to a few animals.

�-88-

The alpine zone in the LaGarita Mountains is very extensive and offers
an abundance of summer range; however, suitable wintering areas are
limited to the windswept ridges. Cebolla Creek area offers an abundance
of winter range. Since this herd has been increasing during the past ten
years, increased demands have been placed upon the limited alpine wintering
areas. Further population increases in this herd will be precluded by
limited alpine winter range unless historic migration to winter ranges at
lower elevations can be reestablished through transplanting operations.
A small group of bighorns were trapped on Saguache Creek in southcentral
Colorado during January and February, 1976, then transplanted onto the
Cebolla Creek Wildlife Area to encourage bighorn use in the lower country.
The sheep were fitted with collars made from 4-inch wide canvas webbing
with an outer layer of red vinyl sewn on the webbing.
A white numeral
was painted on each collar so individual animals could be identified.
Nine sheep (6 ewes, 1 lamb, and 2 rams) were trapped, marked with collars,
and transported to the release-site in a large livestock trailer (Table 1).

Table 1. Bighorn sheep released at the Cebolla Creek Wildlife Area
during January-February, 1976.
Sex

Age

Collar Number

Ram

3

3

Ram

3

4

Ewe

Adult

Telemetry

collar 112

black

Ewe

Adult

Telemetry

collar 113

amber

Ewe

Adult

5

Ewe

Adult

6

Ewe

Adult

7

Ewe

Adult

9

Ram

Lamb

Red eartags -- /14

Two of the ewes were fitted with telemetry collars obtained from Te10nics,
Mesa, Arizona.
The transmitters are powered by a "D" size lithium battery
(40-month life expectancy) and transmit at 148 MHz. Transmitter components
were attached to a two-inch wide leather collar with nylon thread and encapsulated in ure.thane. The entire unit weighed appoximate1y 450 grams;
however, newer units preassembled at Te10nics will only 270 grams.

�-89-

RESULTS AND DISCUSSION

Cache la Poudre Canyon
The transplanted bighorns exhibited very erratic movement patterns after
their release. Only 11 sheep (8 ewes, 2 lambs, and 1 ram) were left in
the release area at the beginning of this work segment. Three lambs were
born in June. This group of bighorns remained in the Sheep Mountain area
throughout the summer until deer hunting season in early November (Table
2). Then the heavy concentration of hunters apparently caused them to
disperse out of the area. Four of these sheep were later found in the
upper portion of the canyon, from which they had been transplanted.
The
remaining seven sheep reappeared on Sheep Mountain. In late December
some unknown factor again caused the bighorns to leave the Sheep Mountain
area. There were a few reports of motorists seeing blue-collared sheep
5-10 miles downstream from the release-site.
In late January three of
these sheep were observed 3 miles above the relase-site; two were seen on
Sheep Mountain in March. Then on June 2, 1976 five of the transplanted
sheep were seen approximately five miles below Sheep Mountain; however,
they were not relocated during the next few days.
An aerial survey, using a super-charged Bell helicopter, was attempted
in December. The flight was unsuccessful.
The sheep hid under the trees
when they heard the helicopter approaching, therefore, we observed only a
few bighorns even in areas normally frequented by them.
Due to the rough and extensive terrain observation data on these transplanted sheep has been too meager to make decisive conclusions.
Apparently
the normal behavior patterns of these sheep were disrupted when they were
transplanted from their established territories. Normally bighorn ewebands tend to remain in established areas; however, this group is very nomadic.
They readily abandoned the Sheep Mountain area after they had been there for
several months. Factors and intensity of the disruption necessary to cause
bighorns to abandon an area is relatively unknown. People-use on the Sheep
Mountain area occurs throughout the year, with peak usage in the summer
and fall periods. Tracks of predators (coyote, bear, bobcat, and mountain
lion) were observed on the area at all times of the year. More information will have to be gathered to evaluate the overall results of this .
transplant.

Cebolla Creek
Seven bighorns (6 ewes and 1 ram) remained in the general vicinity of the
release-site.
These sheep were released in February, when there was 1-2
feet of snow on the ground. They remained on the snow-free slopes in the
immediate area until late April. Then as the snow melted, the sheep started
exploring the surrounding area. They were observed crossing the river,
barbed-wire fences, and the roadway with little hesitation.
These explorations were limited to a three-mile radius of the release-site.
The sheep
seemed to be constantly moving, but returned to the release-site every two
to three days. In late May this group moved to Rock Creek, approximately
six miles from the release-site.
This is a small canyon with rough cliffs
along it. This area is away from the main road and not readily accessible
to the general public.

�Table 2. Location of bighorn sheep following release at Sheep Mountain.
Numbers in table represent
miles from release-site; positive numbers being mile:s (airline distance) upstream and negative numbers
being miles downstream.

Date

1

9

10

11

June 3

- - - - 11 - - - - - - - - - - - - 1 - - - - - - - 1
1
- - - - - - - 1 - - 14
- - - - - -

-

-

June 5
June 10
Aug. 9
Sept. 17
Sept. 18
Nov. 14
Nov. 15
Nov. 26
Nov. 28

2

4

5

6

7

-

-

-

15

-

-

1
1

-

16

-

-

Feb. 27

- 13
- 1 - - - 18 18 - - - - 16 - 3 - - - - 18 - 16 - - 16 17 - - 16 16

Mar. 10

-

Dec. 6
Dec. 15
Jan. 19
Jan. 23
Feb. 25

l/These

-

-

Collar Number
12 13 14

15

16

20

18

19

R-1

R-2

Lambs -

-

-

- - -

-

1

- - - 1
- - - - - - - - - - - -

-

1

-

1

1

1

- - - 16 - - 1
- - - - - -

-

-

1

1

1

1 'l:/

-

-

17

-

- 11
- 11
- 13

1

1

-

-

-

1

-

1

1

1

1

1
1

-

14

-

- - -

-

14

-

15

1

-

1

-

-

1

9

1

-

-

1

-

1

1

12

12

- - - - - - - - - - - 12 - - 23

-

-

3

16

-

-

16

-

9

-

1

-

-

-

-

1

11

17

1

-

-

-

-

-

-

sheep were lambs when they were transplanted,

l/Two ear tagged sheep were seen at the same time.

-

-

14

-

3

-

- - 17 -

-

-

-

18
17

-

but are yearlings now.

-

-

-

-

16
1

I

\D

0
i

�The other two bighorns (adult ram and a ram lamb) are unaccounted
for. They were released on the Cebolla Creek Wildlife Area in January,
and immediately joined-up with a ram native to the area. These animals
left the area before the next release was made in February. It was
speculated that the native ram moved back toward the alpine range,
taking the two transplanted sheep with him. However, this has not been
confirmed.
The original plans specified twenty-five animals were to be transplanted
for this study. Due to prior commitments only nine were available.
Another 12-15 animals will be transplanted to the Cebolla Creek Wildlife
Area during the next work segment.
The telemetry equipment has worked very well and been extremely beneficial
in locating and tracking these sheep. Contacts are limited to "line-ofsight"; therefore, due to the rough terrain initial contacts from the
ground are generally less than two miles. The receiving range is extended
considerably by using a fixed-wing aircraft, thus sheep are easily located
by griddingthe area. Radio-telemetry equipment would have been a great
asset in the Cache la Poudre study.

Prepared by ~.~/_~
__.__.~~~~ __~_.~_~
__~_~_~
George D. Bear
Wildlife Researcher

~

.__

��January

-93,..

JOB PROGRESS

State of
.Project

1977

REPORT

:::..CO=-I:=40::..;RAD=:...:O:..__
W~4l-R-26

No.

Work Plan No.
.Joh Ti tle

Personnel:

1

Controlled

¥.eriod Covered:

.

Burning

Bighorn

Sheep. &amp; MOuntain

Job No.

2_5

of Bighorn

Goat Investigations
~

.

Sheep Ranges

June 1, 1975 - May 31, 1976

H. R. Shepherd.

ABSTRACT
No work was accomplished ori this job during the past segment except to begin
a review of the literature.
Work on this project was purposefully delayed
because Harold R. Shepherd, the scientist currently in charge of this program,
will retire before the program gets underway.
It was decided that it was
more appropriate to let his successor plan and conduct this experiment
rather than have to undergo a lengthy orientation and training program for
the new researcher in order to familiarize him with a program which was incompletely initiated.
Mr. Shepherd's replacement will begin his tenure on
this program August 1, 1976 and progress on this job will commence at that
time.

��January, 1977

-95JOB PROGRESS REPORT

State of
Project No.

W-41-R-26

Work Plan No.

1

Job Tjtle

Job No.

26

(CONTRACT)

Investigation of Spontaneous Diseases of Bighorn Sheep._----

Period Covered:
Personnel:

Bighorn Sheep &amp; Mountain Goat Invest~gations

June 1, 1975 - May 31, 1976

Robert Schmidt and Paul Neil, Colorado Division of Wildlife;
C. P. Hibler, T.Spraker, R. Davies, J. DeMartini, Ed Latson,
J. Wegryzn, L. C1um andJ. Hurlock, Colorado State University.

ABSTRACT
Experimental treatment of bighorn sheep to control lungworm (Protostrongylus
sp.) was continued during this segment. Treatments included field trials
of administering drugs through feed supplements, as well as trapping,
treating and releasing. Attempts to monitor treatment effects were made
by direct observation of treated vs unt:reated sheep and measuring larvae
output in feces collected from treated bighorn sheep. Controlled pen
studies on captive bighorn sheep and mouflon/bighorn cross sheep were also
made.

��-97- .

INVESTIGATION OF SPONTANEOUS DISEASES OF BIGHORN SHEEP
C. P. Hibler

P. S. OBJECTIVE
To determine causes, nature and effects of spontaneous diseases in
Colorado's bighorn sheep with particular reference to mortality in
lambs, and to evaluate various means of promoting lamb survival ..

SEGMENT OBJECTIVE NO.1
Experimentally induce transplacental transmission of Protostrongylus
stilesi in bighorn sheep and determine the immune response of lambs to
infection by this route.

METHODS AND MATERIALS
The study of immunodepression (= immune tolerance or immune unresponsiveness)
for parasitic forms of life is in its infancy; immunologists are still
experimenting with extremely simple, single antigens, and the parasite has
a multitude of complex antigens. Moreover, most investigators in the field
cannot agree on the most acceptable procedure to unequivocally answer the
question about unresponsiveness.
The procedure decided upon for this pilot
study is "Lymphocyte Stimulation", one of the procedures considered acceptable.
Eight bighorn or bighorn hybrid ewes will be experimentally infected orally
with third stage larvae of Protostrongylus stilesi during gestation. Lambs
from these ewes and from four non-infected ewes will be collected at birth
and reared on an artificial diet. Heparinized blood will be collected from
each lamb at weekly intervals until death of the infected lambs (expected to
occur within three months of birth). Lymphocytes are separated from the
blood by Ficolhypaque centrifugation and oultured at 2,000,000 cells/ml in
RPMI 1640 media in 5 m1 tubes. Phytohemaglutinin (PHA) will be. used as a
T lymphocyte stimulant and the lipopolysaccharide (LPS) of !. coli will be
used as a B lymphocyte stimulant. DNA synthesis (blastogenesis) will be
determined by uptake of tritium-labelled thymidine by the cells and subsequent
quantitation of Beta emissions in a liquid scintillation counter. Results can
be compared between infected and non-infected lambs by means of computing a
"stimulation index" for each lamb at each time interval.

Stimulation Index

CMP in cultures with stimulant-CPM in control cultures
CPH in control cultures

Depression of the stimulation index of T or B lymphocyte responses to the
appropriate chemical in lungworm infected lambs compared to control lambs
would suggest that immunodepression by the parasite is an important factor
predisposing the lambs to infection by other disease agents.

�-98-

RESULTS AND DISCUSSION
This objective was to be done by Dr. J. DeMartini, Department of Pathology. While he made a concerted effort to accomplish this objective,
problems with support, personnel and equipment made accomplishment of
this objective impossible.
These (equipment) problems currently have
not been corrected.

SEGMENT OBJECTIVE NO.2
Treat bighorn sheep in the Pikes Peak study area with antihelmintics
and evaluate the results.

METHODS AND MATERIALS
Sheep in the Pikes Peak study area currently are being treated with antihelmintics which appear to act against the infective larvae stored in the
ewe's lungs. Sheep on Pikes Peak and at 4-Mile are being treated with a
combination of Cambendazole and Diethylcarbamazine.
Sheep on the Rampart
Range are being treated with Dichlorvos.
Since all sheep (or most) will be
treated, the only means by which efficacy can be evaluated is: (1) Lamb
survival, and (2) Lungworm larval output per gram of feces.
Lamb mortality has been essentially 100 percent in these areas for five
to six years. Lambs produced by ewes generally die in mid- to late summer.
Lungworm larval output is generally very high (2,000 to 10,000 L/g). Evaluation of efficacy will necessitate collection of lamb feces from July, 1975,
through February, 1976, and counting the larvae/gram.
For this data to be
meaningful or valid, it must be compared with data obtained for the last
several years in these herds. It must also be compared with similar data
from known healthy herds. If lambs survive, and the larval output/gram of
feces is low, the treatment can be considered effective.

RESULTS AND DISCUSSION
Much of the results of this effort are included in the report by Robert
Schmidt; therefore, the actual counts will not be duplicated in this report.
Numerous trips were made to evaluate lamb survival by visual means, and to
collect feces from lambs born of treated ewes. The results of this segment
will include only the fecal evaluations.
Forty-six lamb samples collected from Pikes Peak in August, 1975, ranged
from 0-50 larvae per gram of feces; the mean larval output was 13 larvae per
gram of feces. Five lamb samples collected from ewes on Sheep Mountain
ranged from 212-1427 larvae per gram of feces. Four lamb samples collected
from Oil Creek ranged from 365-835 larvae per gram.
Twenty samples obtained from the summer range of the Rampart herd in August,
1975, ranged from 0-382 larvae per gram, average 45 larvae per gram. This
sample includes lambs, yearlings and adult samples. Very few lambs were
available because, as indicated in Robert Schmidt's report, most of the ewes
treated with Dichlorvos aborted and few lambs were available to evaluate.

�-99A total of 48 lamb, yearling and adult samples were collected at 4-Mile.
Larval output in lambs (8 lambs) ranged from 4-933 larvae per gram,
average 126 larvae per gram; however, only one lamb was high. Excluding
this lamb, the range was 4-185, average 46 larvae per gram. Adult and
yearling samples (40 samples) ranged from 3-922, average 214 larvae per
gram.
Samples were collected from Pikes Peak (main herd) in February and March,
1976, for evaluation.
Forty lamb samples collected ranged from 0-1342
larvae per gram, average 160 larvae per gram, a significant increase from
the previous (August) collection.
Adult and yearling samples collected during this period (34 samples)
ranged from 16-687, average 260 larvae per gram. The great majority were
passing 150-600 per gram.
Fenbendazole was evaluated on the Rampart Range herd this year and the
results are given in Table 1. Fenbendazole, as is indicated by the results,
requires longer to eliminate the parasites than other compounds attempted.
Very few fecal samples were obtained from the Poudre herd this year. Ten
adult samples collected in January ranged from 65-928, average 182 larvae
per gram. Only one lamb sample was obtained and it had 48 larvae per gram.

SEGMENT OBJECTIVE NO.3
Evaluate antihelmintics for efficacy against Protostrongylus
captive bighorn sheep and bighorn/mouflon hybrids.

stilesi using

METHODS AND MATERIALS
A total of four adult bighorn sheep ewes and 16 bighorn/mouflon hybrids are
maintained in captivity. All four of the bighorn sheep are infected with
Protostrongylus stilesi and five of the hybrids are infected. Snails
currently are being infected to expose an additional three ewes to infection
with this parasite, bringing the total number of infected hybrid ewes to
eight. Presently, there are four rams in the herd and all of these will also
be infected. Four ewes will be kept as uninfected controls. Lambs from the
infected ewes, and the uninfected ewes and their lambs will be used for
immunity experiments through mid-August, 1975; thereafter, they will be used
in chemotherapy.trials.
A total of 16 infected adult sheep will be available for chemotherapy trials
and (hopefully) 16 lambs (eight infected transplacentally and eight uninfected).
Both age groups can be used in chemotherapy trials, but adult ewes with stored
third-stage larvae will be the most important animals.
Two drugs are presently scheduled for evaluation:
Diethylcarbamazine
Hygromycin B. A third drug (HOE-88l) may be included.

and

�-100-

The type of experiments planned for evaluation of these drugs are referred to as "Modified Critical Evaluations" in that the worm burden in
individuals is measured before and after treatment using larval counts/
gram of feces as the basis for efficacy. Control animals which have not
been treated are also examined.
The 12 infected sheep will be examined daily for two weeks to determine
the level of lungworm larval output/gram of feces. Four will be treated
with Diethylcarbamazine, four with Hygromycin B and four kept as untreated
controls.
Fecal exams will be performed three times per week on each
animal for at least three weeks post-treatment.
If either (both) compound(s) show promise (reduction in larvel output).
then the (both) compound(s) will be evaluated in ewes supporting stored
third-stage larvae. Evaluation must then await evaluation of the lambs
in July and August, 1976.
In the event that neither drug shows promise, an experimental Imidazole
derivative, HOE-88l, will be evaluated. Possibly some additional promising
drugs will be marketed during the trials.
Should a drug show efficacy, it will then be offered to sheep in apple
pulp; a drug can be effective, but to be useful it must be palatable.

RESULTS AND DISCUSSION
Plans to evaluate specific anthelmintics for efficacy against Protostrongylus
stilesi did not materialize as expected. Sheep were experimentally infected,
and sheep that were naturally infected were organized for drug trials. However, Division personnel (R. Schmidt) requested a trial to evaluate
palatability of Camvet (liquid Cambendazole formulated for horses) and the
100 percent powdered Cambendazole.
There also was a suggestion that treatment with Cambendazole on a five day high dose level regimen would remove
all adult lungworm; therefore, much of the planned work was postponed for
these two important experiments.
Before any five day high level dose regimen
could be planned, a toxicity experiment had to be initiated.
In the first experiment (palatability), all adult sheep, both mouflon/bighorn
and bighorn, were offered Camvet in apple pulp, powdered Cambendazole in
apple pulp and straight apple pulp, in three piles. Sheep ate straight
apple pulp and the powdered Cambendazole in apple pulp, but would not
touch the Camvet mixture; it was finally removed two days later.
In the toxicity trials for Cambendazole on a five day regimen, infected
sheep had to be used because, in addition to evaluation for toxicity, the
efficacy of the regimen also had to be evaluated. Four infected sheep were
administered Cambendazole orally at dose levels of 25 mg/lb, 50 mg/lb, 75 mg/
lb and 100 mg/lb. Each sheep was treated daily. The three highest dose
levels caused toxicity in the sheep and each died, despite treatment.
Toxicity was observed on the first· day of trial. The only surviving sheep was
given the dose level currently in use in field situations (see Table lA;
also Tables 2 and 3.

�-101-

Use of these sheep practically eliminated all previous plans; however,
HOE-88l (Fenbendazole) was evaluated on one infected sheep (the only
sheep remaining). This drug, at a dose level of 20 mg/lb of body weight,
effectively eliminated all lungworm. As is indicated in Table 1, it
also has been used in a field trial. More research is currently underway
with this drug.

SEGMENT OBJECTIVE NO.4
Determine the health status of the Poudre River bighorn sheep herd.

METHODS AND MATERIALS
The status of the bighorn sheep herd in the Poudre Canyon will be evaluated
by:
1. Determining
the level of lungworm infection. Fecal samples will
be collected and examined to determine the level of lungworm
larvae/gram of feces throughout the year. This will furnish
information relative to the level of infection in the population.
This can be used as a basis for evaluation at present (by comparison with other herds; e.g. Pikes Peak)or for a future evaluation of the Poudre herd.
2.

Determining relative abundance, distribution and prev~lence of
infection with lungworm as is directly related to the prevalence
of infected snails. Areas of concentrated sheep use will be
sampled by an acceptable and repeatable method to evaluate the
snails.

3.

Information from 1 and 2 above will be combined with lamb production and survival as a basis for evaluating the health status
of the herd. Lamb survival will be evaluated by observation of
the herd, especially the marked animals. Lambs in the herd will
be judged healthy (admittedly subjective) if they do not show
signs of pneumonia.

RESULTS AND DISCUSSION
Due to the press of other work, the maximum efforts on the Poudre herd was
treatment. However, considerable physiological baseline data was obtained
(Tables 4 and 5) along with some pretreatment fecal examinations (Table 6)
and bacteriological results (Table 7).

�-102Table l. Pre- and post-treatment
Fenbendazole.

data on Rampart Range herd using

Rampart Range Pretreatment
April 12, 1976

Sample

Feces
Weight(g)

Total L1

11.L.s

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
.23

3.0
2.6
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0

89
48
260
366
412
48
292
96
1184
300
243
175
255
138
498
186
90
234
252
120
654
54
414

29.7
18.5
86.7
122.0
137.3
16.0
97.3
32.0
394.7
100.0
81.0
58.3
85.0
46.0
166.0
62.0
30.0
78.0
84.0
40.0
218.0
18.0
138.0

Rampart Range Post-Treatment
April 19, 1976
1
2
3
4
5
6
7
8
9
10
11

3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0

109
71
3
54
378
108
126
168
132
96
1170

36.3
23.7
1.0
18.0
126.0
36.0
42.0
56.0
44.0
32.0
390.0

-----------------------------------------------------------------------------

�-103Table 1. Pre- and post-treatment data on Rampart Range herd using
Fenbendazo1e (continued).
Rampart Range Post-Treatment
April 28 &amp; 30, 1976

Sample

Date

Feces
Weight(g)

Total L1

11L.&amp;

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40

4/30
4/30
4/30
4/28
4/28
4/28
4/28
4/28
4/28
4/28
4/28
4/30
4/30
4/30
4/30
4/30
4/28
4/28
4/28
4/28
4/30
4/28
4/28
4/28
4/28
4/30
4/30
4/28
4/28
4/28
4/28
4/28
4/28
4/28
4/30
4/28
4/30
4/28
4/28
4/28

2.8
2.1
3.9
4.5
5.0
3.1
4.9
5.0
4.1
5.0
3.1
2.5
3.7
3.3
5.1
5.0
4.4
3.9
2.5
4.9
3.0
5.0
5.0
5.0
4.6
2.5
4.5
3.6
3.8
3.4
5.0
5.0
5.0
5.0
2.7
5.0
5.0
5.0
5.0
3.5

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
10
0
0
0
0
0
257
0
0
50
0
0
0
0
0
0
0

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2.0
0
0
0
0
0
75.6*
0
0
10.0*
0
0
0
0
0
0
0

*An untreated ewe was observed on the range on these dates.

�-104Table lA.

Fenbendazole

treatment of ram lamb from Trickle Mountain.

*

Baermann Results
Date

Treated

Treated

17 March
19 March
20 March
22 March
24 March
25 March
26 March
27 March
28 March
30 March
31 March
2 April
6 April
8 April
9 April
10 April
11 April
14 April
16 April
17 April
18 April
19 April
20 April
29 April

Feces
Weight(g)

Total L1

11Lz

5.5
3.0
2.3
7.3
6.7
5.1
4.0
4.0
1.8
2.6
7.3
7.8
4.3
6.7
8.3

142
51
126
90
61
42
76
83
27
51
25
21
1
2
4

28.00
17 00
54.78
12.33
9.10
8.20
12.96
20.75
15.00
19.62
3.40
2.60
0 23
0.30
0.48

5.1
4.6
1.8
7.5
5.4
6.6
5.6

2
2
21
3
0
0
0

0.39
0.43
11.67
0.40
0
0
0

0

0.5 gr. Fenbendazo1e

0

20 mg./1b.

*This ram died in late summer and did not have lungworm.

(Old feces)

Neg. on 2 floats

�Table 2. Results of fecal sample evaluations
prior to drugadmiriis tratTon.*

Animal

1

2

3

4

5

made on experimental

sheep 13 days

Day
(Larvae/ gram)
6
7
8
9

10

11

12

13

M

-

21

-

7

2

3

1

2

+

1. "Grandma"

-

23

168

- 32

2. "Brandy"

254

373

668

78

44

347

205

541

64

28

216

60

117

+

3. "Minnie"

1

19

42

54

58

25

23

7

49

1

12

3

28

+

4. "Willy"

16

63

68

43

45

3

17

10

11

0

4

7

12

+

1.
2.
3.
4.

x = 20
x = 232
x = 25
x = 23

*Cambendazo1e:

3% Suspension, Merch-Sharpe-Dome.

I

b
Ut
I

Table 3.

Toxicity

trials.

'X-Profostrong~Tus
Larvae/gram

Animal

sp. 'IC

MuellerIus sp ,
Larvae

Dosage

Days

1. "Grandma"

20
(range: 0-168)

+

25 mg/1b

7

2. "Brandy"

232
(range: 44-668)
25
(range: 1-58)
23
(range: 0-68)

+

100 mg/lb

6

+

75 mg/1b

3

+

50 mg/1b

5

3. "Minnie"
4. "Willy"

*Resu1ts

listed on Table 2:

13 days consecutive

pre-experimental

Duration
Negative

Histologic Changes
Noted in Lungs

Still alive; L1/g =
12 ProtostrongIlus,
2 Muellerius.
Until death: Larvae and ova are
11/14
dead.
Until death: Larvae and ova are
11/11
dead.
Until death: Larvae and ova are
11/17
dead.
14 weeks
(11/17-3/2)

fecals.

�Table 4.

Co 11a r Co lor
end Number
Blue 1
- Blue 2
- Blue 4
Blue 5
Blue b--Blue 7
- Blue
9
Blue
10
Bl
ue
11
-Blue 12
Blue 13
Blue
14
Blue 17
Blue 18
--B 1ue 19
Blue no #
Red 1
- Red 3
Red 4
5
- Red
Yell ow 1
Yell ow 2
-Yellow 4
- Yell ow 5
- t'lean
Ranqe
-Standard
Deviation

Sex

Blood values of Poudre Canyon bighorn

Age in
Years

PCV
%

Hemoglobin
gr/l00ml

Ewe
4.5
62
22.2
Ewe
5.5
55.5
19.4
Ewe
5.5
54
18.6
Ewe
6.5
50
17.6
Ewe
5.5
51
18.1
Ewe
3.5
59
20.0
Ewe
6.5
57
19.8
Ewe
2.5
50
17.8
E\oJe
6.5
53
18.6
Ewe
4.5
50
18.0
Ewe
2.5
48
16.8
21.1
Ewe
4.5
49
Ewe
3.5
54
19.1
1.5
Ram
45
15.8
Ewe
2.5
57
26.4
1.5
Ram
53
18.4
Ewe
4.5
53
19.3
Ewe
3.5
50
17.6
Ewe
2.5
54
20.0
Ewe
8.5
50
17.8
Ewe
6.5
57
20.2
Ewe ~.5 or 4.5&gt;i 49
17.6
Ewe
3.5
56
19.4
Ewe
5.5
56
19.7
4.5
53.02
19.14
1.5-8.5
145-59 15.8-26.4
3.97

2.09

I :~')tal
Protein
gr/l00ml

sheep.

Fibrinogen
mg/l00ml

White
Blood Cell s
per ml

Neutrophils

Differel t ia l (%)
Lympho.
Mono.

8.1
7.1
7.1
6.5 .
7.5
6.6
7.4
5,9
6,5
6.0
5.7
7.3
6.6
5.7
6.6
7.0
6.1
7.2
6.0
6.5
6.6
5.3
6.7
6.9
6.66
5.7-8.1

300
200
100
300
300
300
100
100
300
200
200
200
300
200
600
300
300
500
200
200
100
100
200
200
241.67
100-600

4700
2600
3400
3700
4600
5100
3200
5000
1600
4200
4400
7700
5200
3900
5300
9900
3700
9100
4700
4700
4300
7600
6100
4200
4954.17
1600-9900

54
55
43
73
58
43
72
75
71
87
86
69
29
76
21
63
85
81
78
76
61
53
56
63
63.67
21-87

30
39
39
21
33
40
24
21
24
10
11
27
54
20
71
33
13
10
18
21
35
40
41
29
29.33
10-71

0.6

121.285

1938.619

17.362

14.435

5
3
8
2
3
3
2
2
3
3
2
2
4
2
2
2
1
3
2
2
1

Eosin.
10
3
10
3
7
13
2
2
2

-

Baso.
1

-

-

1

-

1

-

--

2.46
0-8

1
2
12
1
5
1
1
5
2
1
3
7
1
8
4.25
0-12

.33
0-1

1.61

3.87

.48

-

2

-

1
1
1
1

-

1

-

-

* 4.5 Years used in calculations
I

I-'

o
0\

I

�Table 5.

Serum chemistry values.

See key on attached page for numbers 1-17.
-------

·Collar Color
and Number
=
Blue 1
Blue 2
Blue 4
Blue 5
Blue 6
-Blue 7
B1ue 9
81ue 10
81ue 11
Blue 12
81ue 13
B1ue 14
Blue 17
Blue 18

1
176
85
75
95
125
109
78
202
80
176
182
65
117
142

Blue 19
-Blue Ram
Red 1
Red 3
Red 4
.
Red 5
Ye11ow 1
Yellow 2
Yellow 4
Yellow 5

118
130
182
186
208
240
160
137
157
127

t~ean
Range

139.67 13.98 2.89 6.45 1.62 09.67 4.94 143.17 .54
1.1-:
2.15 13.65 - 141 - 2.45
;65-240 c...-0-19 3.8
51-85 i 6.65
148
.4-.8

S-ianClard
k\~yjQ t i on

2
14

10.5
11.5
10
15
12
11
11
19
14
15
17
12
13
18
16
11.5
11
13
17
15
19
18
12

-I-~
47.65 2.9

3
3
3.5
3.1
3
3.8
2.45
3.7
2.25
3.45
2.6
2.15
3.25
2.4
2.3

4
6.1
7.1
7
6.3
7.2
6.4
7.1
5.8
7.7
6.0
5.7
7.2
6.3
5.6

2.8
3.3
2.5
3.4
2.4
3.2
2.6
2.55
2.8
2.75

6.3
6.7
5.8
6.5
5.7
6.3
6.3
6.3
6.7
6.7

5
1.4
1.4
1.6
1;75
1.75
1.45
2.45
1.50
1.65
1.6
1.75
1.8
1.1
1.49
1.6
2.0
1.4
1.8
1.55
1.5
1.45
1.6
1.45
1.75

6
73
85
80
80
65
75
65
70
82
62
59
79
68
58

7
4.95
5.35
.4.45
4.95
4.1
5.4
5.4
6.65
6.4
3.65
4.12
5.98
5.32
4.25

8
142
143
144
142
142
144
143
142
147
142
142
148
144
142

9
.5
.5
.6
.6
.8
.7
.5
.5
.6
.5
.5
.7
.4
.5

10
95
45.5
50
37
43
95
61.5
122.5
68
85
115
59
95
185

11
330
300
235
167
195
142
175
212
145
165
208
115
195
150

12
105
75
118
125
130
89
80
99
'95
130
118
105
105
95

13
172
133
120
115
149
127
49
121
130
54
95
121
130
93

14
5.3
7.4
6.8
6.6
7.1
6.9
6.2
4.7
8.4
2.9
3.2
7.9
6.7
4.9

15
350
345
300
250
375
275
375
425
275
300
340
250
200
360

16
10.6
11
11.6
11.9
11.6

55
61
~5
51
70
~9
~O
72
p8
~O

4.9
5.2
3.75
5.5
4.5
4.32
4.58
5.42
4.8
4.65

145
142
142
141
142
142
142
144
142
142

.5
.7
.5
.5
.5
.5
.4
.5
.5

135
115
51
117
105
35.5
335
82.5
95
32.5

178
128
195
525
325
325
160
195
195
128

135
89
100
112
135
115
112
95
145
105

190
89
119
385
234
229

8.1
6.1
3.6
4.9
4.5
5.4
4.8
6.2
6
5.9

335
325
360
555
475
530
275
275
345
370

11
3
11.7 2.7
10.7 2.3
12
2.6
11.1 3
11
/3
10.2 3
10.9 2.3
16
2.4
11.6 2.7

.

r-

;)

,:~

.48

.56

1.02

9.2

2.40

3.13 .308

88
96
120
85

10.6
10.7
11.6
11.9
10.9
11.1
14
10.7
11.4

17
2.3
2.3
2.5
2.5
2.8
2.7
2.4
2.1
2.2
2.9
2.8 I
3.1 ~
2.5
2.4

81.60 212.00'110.21 135.17 5.85 344.38 11 .49 2.61
200- 10.2- 3.:..L.....
2.1 14
fss5 - iH - 75-145 4j85 ~:4-' 555
38 82

91.69

50 34

70.10

I

1.48

85.41

3.55

2.23

�-108-

~or
l.

Attached Table

15.
16.
17.

Glucose mg/100 ml
Blood Urea Nitrogen mg/100 m1
Globulin gr./100 ml
Total Serum Protein gr/100 ml
Uric Acid mg/100 ml
Cholesterol mg/100 m1
Potassium mg/l
Sodium mg/l
Total Bilirubin mg/100 ml
Alkaline Phosphatase IU/la
..
b
S.G.O. T.
IU/l
c
S.G.P.T.
IU/l
d
C.P.K.
IU/l
Inorganic Phosphorus mg/100 m1
lDHe IU/l
Calcium mg/100 m1
Creatinine mg/100 m1

a.
b.
c.
d.
e.

International Units/liter
Serum Glutanic Oxaloacetic Transaminase
Serum. Glutanic Pyruvic Transaminase
Creatine Phosphokinase
lactic Dehydroginase

2.
3.
4.
5.
6.
7.
8.
9.
10.
1l.
12.
13.
14.

�-109-

Table 6. Pretreatment
January 7, 1976.

fecal Baermanns

- Poudre Canyon,

Location

Age

Weight (g)

Total L1

11.Ls

A

3.0
3.0
3.0
3.0
3.1
3.0
2.8
3.0
3.0
3.0
3.0

114
858
624
871
767
792
2104
1416
688
448
16

39
286
208
290
255
264
751
472
229
150
5

3.0
3.0
3.0
3.1
3.1
3.1
3.2
3.0
2.7
3.0
3.0
3.0
3.0
3.0
3.0
3.1
3.0
3.0
3.0
3.0

32
704
272
408
679
24
64
192
336
240
1445
196
247
230
688
315
119
406
546
2786

11
235
91
136
226
8
21
64
l12
80
482
65
82
77
230
105
40
135
182
930

Meadows

A

A
A
A
A

A
A

A
A
Y
Ponds

A
A
A
A
A
A
A
A

L
A
L
A
A
A
A
L/Y
A

A
A

Y/A

�-110-

Table 7.
results.

Poudre trapping operation - January 21, 1975, nasal bacteriological

en

~

~
~
..-l

E

r-l

&lt;Il

0

~
&lt;:

75PR-3
75PR-4
75PR-S

j:!l

+
+
+
+

75PY-l
75PY-2
75PY-lf

+

--

75PB- 6

75PB-4
75PB-5
7SPB-7
-i.J.l.
~pp.
'.0
.u-..L
--_.
75PB-9
75PB-IO
7SPB-ll
75PB-12
iSPB-13
75PB-14
.75PB-15
75PB-17
75PB-19
75PB-Ram

o

('j

,.0

0

+

-

0

'.-1

:r::

~
~
~
14

('j

~

::&lt;!

+
+
+
+
+
+
+
+
+
+

14

::s

~Or-l61

'"::s

(I)

14
p.,

Ul

(I)

::s

::-

p.,

(I)

~

-e
aJ
..-l

0
(.J
0

4-1

?

~ 0

'.-1

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~

(1)4-1

co

g
•...

aJ
14

'.-1

CJ)

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CJ)

CJ)

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r-l

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r-l
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(I)

en

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(.J
0
r-l

&lt;Il
r-l
r-l

~
0
~
0

tI)
Ul

Z

+
+

14

Ul
tI) '.-1

..-l

..-l

~

&lt;Il

&lt;Il

(I)

aJ

::s
(.J

en

14
OJ

Ul

(I)

+
++
+

75PB-l
75PB-2

B

co

+
+
+
+

75PR-l

Y
R

::s

r-l
r-l
''-;
(.J

r-l

&lt;:

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en

co

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14
aJ

Number

CJ
r-l
r-l

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+
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+

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+

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+

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!

--1------

+
+

+---1

+
+
+

+
+

+
+

+
+

+

Blue collar
Yellow collar
Red collar
Pasteurella species, though seldom isolated from routine nasal swabs taken during winter
months, are apparently still avabilable to cause disease complications, e.g. a ewe lamb
picked up in Poudre Canyon January 24, 1975 by Game and Fish personnel contained a
septicemic Pasteurellosis, apparently induced by heavy burdens of Protostrongylus 3?..

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                  <text>-1-

April 1977

JOB FINAL REPORT
State of

COLORADO
--~~~~~------

Pr.oject No.

W-37-R-30

Work Plan No.
Job Title
Period

Job No.

21

Pheasant Mortality Investigation

Covered:

Personnel:

1

Game Bird Survey

April 1, 1974 through March 31, 1977

Donald M. Hoffman, John Corey, Howard D. Funk, Steven Steinert,
Lucien Brevard, Larry Crooks, Bruce Trindle, Daniel Walsworth,
W. Thomas Howard, Michael Fedorko and Warren D. Snyder.

ABSTRACT
This study was initiated to investigate mortality rates of ring-necked
pheasants on the Tablelands of northeastern Colorado. Poor reproduction
in 1974 followed by high mortality from a March, 1975 snowstorm drastically
reduced populations on the study area. Additional mortalities were recorded
during a November, 1975 snow and a devastating March, 1977 snowstorm, so
that only a remnant population of pheasants remained within the region at
termination of the study. Crowing indices had dropped by nearly 50 percent
from the mid-1960's to commencement of this study in 1974. An additional
68 percent decline was noted from 1974 to 1975. The crowing index showed
little change from 1975 to 1976. Production census showed similar trends
averaging 2.69 birds observed per mile from 1966-1968, 1.88 in 1974,
0.38 in 1975 and only 0.29 in 1976. Nightlighting techniques were used
to trap and band 233 roosters in the late summer-early fall interval of
1974, 109 in 1975, and 142 in 1976. Hens trapped and marked included
301 in 1974, 150 in 1975, and 187 in 1976. Reward bands, used to correct
for failure of some hunters to return bands, showed that a majority of
hunters failed to mail in regular bands obtained during hunting seasons.
~eward band information was used to project harvest rates of 18.5 percent
in 1974, 22.4 percent in 1975, and 16.0 percent in 1976. Even when crippling
loss was included, total harvest mortality never exceeded 25 percent of
the available roosters. Projections of fall-winter rooster mortality based
on sex ratio changes were much higher than those derived from band return
information for unknown reasons. Based on very limited band return data,
average annual mortality rates approximated 97 percent for roosters and
75 percent for hens during the period of study. Environmental observations
indicate dry weather apparently reduced reproduction and potentially
decreased wheat stubble height. Wheat stubble height was considered of
potential importance to pheasant survival during winter snowstorms. Windbreaks and shelterbelts, although few in number, were of little value
to pheasants during these winter storms and high pheasant mortality would
still have occurred if many more such cover areas had been present.

�-2-

RECOMMENDATIONS
Recommendations
this report.

have been incorporated

with conclusions

at the end of

�-3-

PHEASANT MORTALITY INVESTIGATION
Warren D. Snyder

This mortality study was initiated to gain better insight into the dynamics
of pheasant survival and mortality.
Efforts were designed to determine
what proportion of annual mortality was attributed to natural causes and
what proportion of the roosters were taken each year by hunters.
It
attempted to also ascertain annual turnover rates of both sexes within
the povulation. Regrettably, our efforts in attaining these objectives
were severely hampered by a series of unforeseen environmental limitations
which, by drastically reducing populations, have resulted in early termination of this study.
P.N.O. OBJECTIVE
To investigate mortality

rates of a population of ring-necked pheasants.

METHODS AND MATERIALS
Field surveys and surveys of census information were used in selection
of a study area in the primary pheasant range in the Tablelands of northeastern Colorado. A study area, encompassing the four southwest townships
of Phillips County (Townships 6 and 7 north and Ranges 46 and 47 west)
was selected during the summer of 1974 on the basis of population densities.
landowner cooperation, and habitat types. Some extension of study area
boundaries was made during fall trapping operations of the second and
third years of study.
Pheasant populations within the selected study area were monitored throughout
the study. One crowing route was established and censused in 1974 and two
routes (Fig. 1) were censused in subsequent years using the standardized
Colorado method (Sandfort, 1960a). In 1975 the starting and ending locations of these routes were varied to partially correct for call variance
relative to time within counts. Efforts were made to tally total calling
males on 40 square miles within the study area in conjunction with crowing
census in 1975. Four square miles surrounding crowing stations numbers 1,
3, 5, 7 and 9 on each of the two routes were checked using triangulation
methods. This sample of 40 square miles represented 27.8 percent of the
study area (Fig. 2). Two men in separate vehicles with radio communication
pin-pointed each male during early A.M. calling periods. Repeat counts
were made on different mornings to increase accuracy.
One trend production route was censused in 1974 and four routes were
established and censused in subsequent years of study by standard brood
census methods used in Colorado (Sandfort, 1960b). Two of these utilized
previously mentioned crowing routes (Fig. 1). Pheasant sightings were
segmented at two-mile intervals along the routes to pinpoint potential
trapping sites in 1975 and 1976.
Spring sex ratio counts were attempted, but study area pheasant populations
were too sparse to permit attaining adequate samples so efforts were

�&gt;....
2:

:::&gt;

o
u

17

YUMA
COUNTY
Fig. 1. Pheasant crowing count routes with listening stations (cross checked
lines). which were also utilized for production censuses. and additional census
routes (dotted lines) within the mortality study area.

�-5-

L 7N

25

30

i621 •

~~~O

"II

c

II

o

lG

"

II

Z

&lt;2:

c

:3.

C)

o

5

\

c

\

J

b.

.I
;\
)

).

.J
R.47W

11

COUNTY
YUMA
Fig. 2. Triangulation blocks censused within the study area during the spring
of 1975. Numbers represent the total roosters recorded per four square mile
block.

o

!

i

�-6-

discontinued.
Late winter sex ratio tallies were obtained within the study
area and in adjoining areas in February and March of 1975 and 1976.
Pheasants were captured using nightlighting techniques developed in Illinois
(Labisky 1968). Two crews of two men each and two 4-wheel drive vehicles
were used through most of the study. Trapping operations were confined to
wheat stubble fields after obtaining access permission from landowners.
Because of low populations, pheasant trapping was extended to locations
outside the study area during the falls of 1975 and 1976. Late winter
trapping, conducted in February, 1975, was not resumed in subsequent
winters due to depleted populations, evidence of egress from the study
area and low numbers of roosters present.
Bait trapping efforts were
abandoned in winter, 1975 after preliminary efforts failed to attract
birds to the traps.
All trapped pheasants were banded with consecutively numbered aluminum
leg bands. Bands were placed on all birds in the six to seven week age
class and older. Approximately one half were also banded with gold colored
aluminum reward bands during fall, 1974 and winter, 1975. Reward bands
were not placed on hens in subsequent years and reward bands were placed
on about one half of the roosters in fall, 1975 and on all of the roosters
trapped in fall, 1976. Trapping efforts in fall were concentrated from
mid-August to mid-October.
Pheasants were aged in the field when possible
and the first (proximal) primary was taken for later shaft measurement and
age verification (Wishart 1969).
Hunting mortality data were collected during field contacts of hunters, by
hunter questionnaires, and in 1974, by contacts of hunters at the Fleming
check station.
Hunters reporting banded birds were surveyed to obtain
hunter success, crippling loss, location of kill and other pertinent
information.
Their hunting companions, too, were queried to obtain additional
information.
Those hunters reporting a gold "reward" band were given a 144
page book, Cooking the Sportsmen's Harvest, published by the South Dakota
Department of Game, Fish and Parks, as a reward.
DESCRIPTION

OF AREA

This study was initiated in the northeast corner of Colorado and was
contained for the most part in southwest Phillips County (Fig. 1).
This location over previous years and at the time the study was initiated
contained some of the higher pheasant densities within the region and
state. Some study efforts were extended into adjoining areas of Yuma
and Logan counties and also to the east in Phillips County as the study
progressed.
This area is essentially high plains tableland, predominately flat and
containing loam and sandy loam soils. A few intermittent drainages
flow from west to east across the study area and provide occasional small
areas of unfarmed native grass and weed cover. Some willow patches also
are distributed along these drainages (Fig. 3). Elevations approximate
3,700 to around 4,000 feet. Few farmers live in the study area, although
some who farm there reside in the town of Haxtun to the north. Reference

�-7-

Fig. 3. Pheasants surviving the snowstorms of March 1975 and 1977
congregated in sparse willow and cottonwood cover along small intermittent
drainages like this. (Photo by Warren D. Snyder)

�-8-

is made to Snyder (1970) for a breakdown of cover and crop types present
in the mid-1960's.
Winter wheat, which is summer fallowed on a biennial
basis, is still the predominate crop, however, deep well irrigation has
increased in recent years increasing the quantity of corn within the region.
The primary irrigated areas lie to the south and east of the main study
area. Precipitation approximates 17 inches per year on the average with
most of this falling during the 147 day growing season. Average annual
snowfall approximates 34 inches. Winter wheat stubble provides the predominant cover for pheasants in the area through fall, winter, and early
spring periods.
Green wheat replaces stubble as cover in late spring and
early suwmer. Only a few windbreaks and shelterbelts are present within
the study area and most are marginal in value during snow stress periods.
RESULTS AND DISCUSSION
Pheasant Population

Monitoring

Pheasant Crowing Counts
Crowing counts, when compared from year to year and also with years previous
to the study, probably most accurately depict breeding population level and
trend. Results summarized in Table 1 and illustrated in Figure 4 show the
marked downward trend in crowing indices. No significant difference was
noted between count totals in 1975 and 1976. Based on observations of
pheasants present in late March, 1977, the subsequent call index, if conducted,
would have been even lower than in the two previous years.
Four replicates were conducted on one route in the western part of the study
area in 1974, and this information was combined with that from a management
census route also conducted in the region of study. Thirteen counts were
completed on two routes in 1975 and eight replicates were completed on the
two routes in 1976. Starting points of counts were rotated among stations
probably resulting in some bias upward in high count per station indices in
1975 compared with other years. This is due to the fact that roosters
usually peak in crowing activity about 20 to 30 minutes before sunrise.
Distribution of this activity peak among more stations would increase the
overall average.
The high number of replicates probably also increased
census in 1975 since chances of hitting peak counts would be increased
when more counts were used.
Table 1. Pheasant
pheasant mortality

Interval

1963-68
1974

crowing counts as indices of population
study region.

Census Route

Central control
Wages-Haxtun

1975

East

1976

East

&amp;

&amp;

study rte.

west study routes

s west study routes

change in the.

High Count/Station

Percent Change

45.3}

_490Z}
-83.9

23.0}
-68.3
7.3}
+8.2
7.9

�-9-

.5
3.0.

1
.5

o

n

1963-68

1974

1975

1976

Fig. '4. Comparative trends of crowing count; indices .and pheasants obae.rved
per mile during production counts in the mortality study area region.

�-10-

Triangulation

Inventory

Triangulation methods were used in conjunction with crowing census in 1975
to relate crowing indices to actual pheasant densities.
Rooster densities
were very low which facilitated pinpointing of individual roosters.
As a
consequence, results were believed to be fairly accurate.
Projections based on data in Figure 2 indicate that if 63 roosters were
present in the 40 square miles sampled, then approximately 227 roosters
were resident in the total 144 square mile study area. This represented
1.58 roosters per square mile. Very few birds were located in the northwest part of the study area in spring 1975 as shown in Figure 2. Much
higher densities were located to the southeast.
This differential was
primarily due to greater severity of the preceding March storm which caused
higher mortality in the north and west parts of the study area.
Pheasant Production

Counts

Table 2 summarizes August production counts completed within the study
area from 1974 through 1976 and compares indices with those obtained in
the late 1960's (Fig. 1). The average birds-per-mile observed in the
late 1960's was 2.69. This index dropped to 1.88 in 1974, to 0.38 in
1975 and to 0.29 in 1976. The latter index was only slightly more than
one-tenth of that obtained in the late 1960's. Data were based on seven
replicates in 1974, 15 in 1975, and 18 in 1976.
Pheasant

Sex Ratios

Sex ratios derived from trapping efforts and flushing observations were
obtained in early fall, 1974 through early fall, 1976 and are assumed to
be fairly accurate (Table 3). Biases undoubtedly existed in both trapped
and some observed samples, but samples observed in April and November of
1975 and in late winter 1976 were obtained during snow cover periods which
was believed to increase accuracy.
Low densities were the primary variables
reducing sample size and accuracy throughout the study. Sparse densities
prevented collection of spring sex ratio counts befo~e and during crowing
count census except for limited samples obtained under snow conditions in
April 1975. Trapped samples were predominantly juveniles.
Adult hens
were slightly easier to trap than roosters in fall, however roosters were
more readily spotted during night lighting so ratios are considered near
actual.
Changes in sex ratios from fall trapping to late winter favored hens
and only part of that change could be attributed to hunter harvest as
will be discussed later in the text. Sex ratio changes from late
winter to fall favored rooster survival over that of hens (Table 3).
Based on 3.44 hens per rooster surviving the March 1975 snowstorm and
triangulation projections of 1.58 roosters per square mile, there were
5.4 hens per square mile present in spring, 1975. Crowing indices averaged
about 8 percent higher in 1976 than in 1975 (Table 1) and the proportion
of hens per rooster was slightly lower, so it is doubtful that hen density
changed much from one year to the next.

�-11Table 2. A comparison of pheasants observed
counts in the mortality study area.

per mile during production

Interval

Miles

Birds Observed

Birds-per-Mi1e

1966-68

515

1,383

2.69

1974

189

355

1.88

1975

390

149

0.38

1976

457

l34

0.29

Table 3. Pheasant sex ratios obtained from early fall, 1974 through
early fall, 1976 on the mortality study area.

Roosters

Hens

Total

Percent
Males

Percent
Females

Ratio

Adults

48

143

191

25.1

74.9

1:2.98

Young

185

158

343

53.9

46.1

1:0.85

Total sample

233

301

534

43.6

56.4

1: 1.29

Observed

36

179

215

16.7

83.3

1:4.97

Trapped

32

141

173

18.5

81.5

1:4.41

34

117

151

22.5

77 .5

1:3.44

Adults

19

63

82

23.2

76.5

1:3.16

Young

94

94

188

50.0

50.0

1:1.00

Total sample

113

157

270

41.9

58.1

1: 1. 39

71

104

175

40.6

59.4

1:1. 46

185

261

446

41.5

58.5

1 :1.41

119

373

492

24.2

75.8

1:3.l3

Adults

30

74

104

28.8

71.2

1:2.47

Young

115

120

235

48.9

51.1

1:1.04

Total

145

194

339

42.8

57.2

1: 1. 34

Sample

Trapped sample,
fall, 1974

Late winter,

AEril

1975

'75 observed

Trapped sample,
fall, 1975

Nov.

'75 observed

Combined

fall total

Late winter
observed

'76

Trapped sample,
fall, 1976

�-12-

Pheasant Trapping

and Banding

Trapping success peaked in fall 1974, in spite of lack of experience and
equipment problems.
That fall, 534 birds were captured, banded and released
in wheat stubble fields using night-lighting methods (Fig. 5). The sample
dropped to 259 pheasants marked in fall 1975 plus 11 recaptures of previously
marked birds. Fall samples in 1976 included 329 banded plus 10 retrapped
recoveries.
These figures actually encompass two distinct populations
since roosters were hunted and females were not, and mortality data must
be computed separately.
Samples for the three years are displayed by age
and sex in Table 4. Hoffman (1975) also trapped 173 pheasants during
the late winter of 1975 of which 149 were banded and 24 were retrapped
from fall.
Gold leg bands stamped "Reward" were placed on slightly more than half of
all pheasants banded during fall 1974 and winter 1975 intervals.
Approximately half the roosters were again reward banded in fall 1975 but regular
bands alone were used on all hens in fall peri9ds of 1975 and 1976. All
roosters trapped in fall 1976 were marked with reward bands to increase
sample size of recoveries.
A summation of those banded with regular and
reward bands is presented in Table 5.
Age Ratios of Trapped Samples
The number of young per adult and per hen are presented in Table 6 for the
three years of study. Ratios obtained during production counts and fall
trapping are provided for comparison.
These figures suggest the probability
that estimates of proportion of young present during production counts
were higher than were actually present in the population, mainly because
unsuccessful hens and moulting roosters were more difficult to observe.
Young were as readily observed as adults during fall trapping but were
more easily captured, which has some effect on fall trapping ratios as
well. Production census immediately preceded the primary period of
trapping, but some differentially greater mortality of young than adults
probably occurred through the late summer-early fall interval.
These
factors probably acted in combination to account for age ratio differences
between production count and trapping data.
Both sets of data in Table 6 suggest that reproductive success in 1974 was
exceptionally poor. Young per adult ratios increased in 1975 and were still
higher in 1976 as indicated by both indices.
In comparison, young per hen
ranged from 2.45 to 5.75 during production counts in the same general area
from 1963 through 1968 (Snyder 1970). The average young per hen index
during that interval was 4.38. Thus, production comparisons between the
60's and 70's data indicate 1974 was below average and 1975 and 1976 were
above average.

�-13-

Fig. 5. Pheasants were netted by hand in wheat stubble fields by two
2-man crews working with spotlights. (Photo by Donald M. Hoffman)

�-14-

Table 4. Age ratios by sex in fall and winter banded and retrapped
samples from fall 1974 through fall 1976.

Adult

Male
Juv. Total

Adult

Female
Juv. Total

Combined
Adult Juv. Total

Number

48

185

143

158

191

343

Percent

20.6

79.4

47.5

52.5

35.8

64.2

Number

12

20

80

61

92

81

Percent

37.5

62.5

56.7

43.3

53.2

46.8

Number

19

94

63

94

82

188

Percent

16.8

83.2

40.1

59.9

30.4

69.6

Number

30

115

74

120

104

235

Percent

20.7

79.3

38.1

61.9

30.7

69.3

Interval
Fall 1974

Winter

233

301

534

1974-75
32

141

173

Fall 1975
113

157

270

Fall 1976
145

194

Table 5. Pheasants banded within and adjacent
area from fall 1974 through fall 1976.

Sample

to the mortality

339

study

Fall 1974
Winter 1975
Fall 1975
Fall 1976
Reg. Reward.!! Reg. Reward.!! Reg. Reward!! Reg. Reward.!!

Male
Adult

15

33

1

9

5

10

0

27

Juvenile

92

93

8

10

47

-47

0

115

Total

107

126

9

19

52

57

0

142

Combined

233

28

109

142

Female
Adult

77

66

35

33

56

0

67

0

Juvenile

94

64

30

23

94

0

120

0

Total

171

130

65

56

150

0

187

0

Combined

301

121

150

187

0

24

11

10

Retrap recoveries

1/ Both regular aluminum
on reward-banded

bands and gold bands marked
pheasants.

"reward" were placed

�-15-

Table 6. Number of young per adult and young per hen found during late
summer production counts and fall trapping intervals.

Sample

1974

1975

1976

Number of young per adult

2.87

3.87

4.15

Number of young per hen

3.83

4.46

6.00

Number of young per adult

1.80

2.29

2.26

Number of young per hen

2.40

2.98

3.18

Production data

Fall trapped sample

Trapping Success per Unit Area
Despite indicated poor production in 1974, higher pheasant densities
permitted concentration of trapping efforts in a smaller quantity of
land while obtaining more birds than in subsequent years of study.
Ninety-one pheasants were trapped in three fields within one section in
the southwest part of the study area in 1974 while sixty-three birds
were captured in three fields in another section in the southeast portion
of the study area. Trapping efforts seldom yielded over 20 pheasants per
section in subsequent years of study.
The two 2-man trapping crews, working in unison, searched approximately
10,340 acres of wheat stubble during the mid-August to mid-October interval of 1975. Approximately 8,910 acres of this was in the study area
proper with much effort concentrated in the southeast portion. An
additional 1,430 acres in locations adjacent to the primary study area
were searched. This total acreage represented about 123 fields and search
efforts covered the equivalent of all stubble fields in about 46 square
miles. Only about 2.46 roosters and 3.41 hens were banded per section in
1975, and an average of less than one rooster was obtained per field
searched. Data for 1976 were comparable, however, more effort was concentrated adjacent to irrigated cornfields to the east and south of the
main study area.
Most landowners granted trapping access permission although denial increased
as the study progressed and pheasant populations deteriorated.
A few
farmers believed trapping caused movements away from their farms, most
voiced resentment of hunting seasons; and a few were afraid of fire danger.
Fortunately, all three fall periods were abnormally dry, permitting routine
trapping to progress from late summer to early fall. Heavy fall rains would
have halted operations and increased landowner objections to trapping
because of the effect of vehicles on fields. Trapping success per bird
observed was high in late summer and tended to begin deterioriating rapidly
in early October. This decrease in success was partly due to weed growth

�-16-

in the fields which dried and became noisy underfoot and under vehicles.
Pheasants also became more wary and difficult to catch as they matured in
early fall, a factor which remained or increased in late winter.
Age Determinations
Nearly all old hens could be distinguished from juveniles during August
and early September trapping because the adults had not yet lost all of
their old primaries.
Likewise, adult roosters could readily be distinguished
from juveniles, 15 weeks or less in age, by moult and spur characteristics.
However, late in fall trapping intervals, ages could not always be accurately
determined in the field. Therefore, the first (proximal) primary was collected from all trapped pheasants in 1974 and 1976 and from a majority of
those banded in fall, 1975. Subsequent measurements provided confirmation
or occasional correction of uncertain field aging based on measurement data
explained in Wishart (1969) and previously plotted by Snyder (unpublished
data) based on feather-spur comparisons for roosters in the early 1970's.
Feather shaft diameter and length data were plotted on scatter diagrams
as illustrated in Figures 6 and 7. Further statistical analyses of these
data will be completed providing confidence limits for aging pheasants
in eastern Colorado in future years.
Hunting Seasons
Seasons and Weather Variables
Hunting began at noon on Saturday, November 16, 1974 and continued through
December 15 in the region of study. In the following year, efforts were
made by the Division to divert hunting pressure away from the March, 1975
storm impacted area in the northeastern corner of Colorado.
Whereas,
seasons elsewhere in the state opened at noon, Saturday, November 15, 1975,
the opening was delayed to sunrise on Wednesday, November 19 in the region
of this study. The 1975 season continued for 12 days and terminated at the
end of the month. Hunting of pheasants began at noon on Saturday, November
20, 1976 and continued for 11 days to the end of the month. Bag and possession limits were, respectively, three and six roosters throughout the
period of study.
Weather conditions were dry in 1974 and 1976, however, cold winds severely
hampered opening weekend hunting efforts in the latter season. A major
snowstorm with considerable wind hit the northeast part of the state on
November 18, 1975 during the night before hunting was to commence.
It
persisted through the first day of season and deposited approximately
10 inches of snow. Nearly all roads in and surrounding the study area
were closed by snow drifts.
County roads were opened slowly and with
difficulty so that no significant hunting occurred until the first Saturday
of season on November 22, 1975.

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�-19-

Table 7. Hunting pressure and hunter success on first and second weekends
of hunting seasons through the period of this mortality study in comparison
with indices from 1965 through 1968 in the same study region.

Year

Miles per Hunter
1st Weekend
2nd Weekend

Hours!
Pheasants
Pheasants Bagged/Hunter
Bagged 1)
Field Checks
Season x!/

1974

0.77

30.00

0.42

1.67

4.68

1975

8.60

22.50

0.82

2.50

3.94

1976

1.25

5.95

0.53

2.19

5.69

1965

0.65

3.29

1966

0.35

2.62

1968

0.56

2.72

0.58

1/ Some potential bias above expected averages possibly existed because
the majority of hunters in questionnaire surveys were those who were
successful or within successful hunting parties.

Hunting Pressure and Success
Table 7 compares first and second weekend hunting pressure among the three
seasons of this study and contrasts these indices of pressure with those
obtained in the region of study in 1965, 1966, and 1968 (Snyder 1970).
First weekend hunter density in 1974 (Hoffman 1975) approached that of the
1960' s. Hunter density dropped drastically in 1975 because of the delayed
opening and snow conditions.
Increased pressure was again noted in 1976.
Second weekend hunting pressure during this study was much below that
which occurred in the mid- to late 1960's when seasons were longer and
more pheasants were present. Second weekend pressure dropped to very low
levels in 1974 and 1975, but was much higher in 1976, possibly because
the shortness of the season induced hunters to increased "early" effort.
Pheasants, while low in number in 1975, had been pushed out of their
normal wheat stubble habitat by deep snow and were concentrated and
exposed along roadways at old building sites and in similar meager cover.
As a result, they were highly vulnerable to hunting by vehicle. One
party of three hunters was checked on the second day of season with 12
birds. All had been shot along roads south of the town of Haxtun. A
majority of hunters checked on Saturday, November 22, 1975 had good
success. Most simply drove the roads and removed exposed roosters as
they found them. A few attempted walking, but deep snow drifts made

�-20-

hunting afoot difficult.
An estimate would place at least 90 percent of
the harvest in 1975 adjacent to roadways in the study area.
Cold winds raked the region of study during the opening of the 1976
pheasant season and cut sharply into field pursuit of pheasants.
Lack
of snow, cold winds and low pheasant densities made even road hunting,
which many hunters attempted, difficult.
This points out one significant
problem with opening hunting of pheasants in late November.
Cold winds
and blustery winter weather often severely curtail hunting effort, hunter
opportunity and hunter enjoyment of the sport. Milder weather, earlier
in the month usually provides much better pheasant hunting conditions.
Hunter Band Return and Harvest Mortality

Projections

Proportional Band Recovery -- Band return information was obtained by
field contacts, follow-up questionnaires, and by mail-in reports.
Hoffman
(1975) obtained a considerable amount of information at the Fleming check
station, however, this management station was closed during subsequent
years of study. Band return information is summarized in Table 8. In
1974, 36 bands were obtained from hunters of which 23 were reward bands
and 13 were regular bands.
In 1975, 13 of 17 returned bands were marked
reward and four were regular bands from immediate fall trapping.
In
addition, one regular band was obtained from the 1974 fall trapping
effort. All roosters were reward banded in 1976 and 22 reward bands were
recovered.
In addition, one reward banded rooster was reported from 1974
banding efforts.
This bird also had been retrapped in fall, 1976.
Hunters did not report or return regular bands at even half the rate which
they returned reward bands unless contacted in the field. Hoffman (1975)
noted that nine of 13 bands obtained in field or check station contacts
were regular bands and four were reward bands. In contrast, of 23 bands
voluntarily reported by hunters in 1974, 19 were reward bands and only
four were regular bands. He found a significant difference (X2 = 10.63;
P &lt; 0.01) between return rates of reward and regular bands from contacts
and voluntary mail-ins.
Only 66.7 percent of the expected number of
regular bands were obtained in 1974 when field check and mail-in returns
were combined.
The same situation existed in fall 1975. Originally 57 reward bands
were placed on roosters that fall of which 13 (22.8 percent) were
recovered.
Fifty-two regular bands were placed on roosters of which only
four (7.7 percent) were recovered.
Twelve regular bands rather than four
should have been reported when ratios were compared.
This represented
a 33.3 percent reporting rate on regular bands. Had field contacts not
been used, mail in reporting rates of regular bands would have been much
lower in both 1974 and 1975.
Sample sizes in both 1974 and 1975 were small, but they clearly illustrated
that hunters failed to return most pheasant bands by mail unless a reward
was offered.
Therefore, all roosters trapped in fall 1976 were reward
banded to increase recovery sample size. There is no reliable way of
knowing whether all reward bands were reported, however, that assumption

�-21-

was made in projecting harvest estimates.
If reward bands had not been used
all three years and field contacts had not been made, then very little
information would have been recovered and there would have been serious
biases in projecting percent harvest or annual mortality rates.

Table 8. Numbers of banded roosters and hens recovered or retrapped from 1974
through 1976 within the mortality study region.

1974
Reward Reg. Total

1975
Reward Reg. Total

1976
Reward Reg. Total

ROOSTERS
126

107

233

57

52

109

142

o

142

Immed. harvest

23

13

36

13

4

17

22

o

22

2nd fall retrap.

1

1

2

1

o

1

2nd fall harvest

o

1

1

• 3rd fall retrap.

1

1

2

3rd fall harvest

1

o

1

o

187

187

Number marked
Recoveries

HENS
171

130

301

o

150

150

2nd fall retrap.

3

2

5

o

4

4

3rd fall retrap.

1

1

2

~umber marked
Recoveries

Harvest Estimates -- Correcting for non-return of regular bands, Hoffman
(1975) placed the estimate of total banded birds harvested and retrieved
by hunters in 1974 at 43 rather than the 36 reported.
Thus, 1974 fall
harv.est was placed at 18.45 percent of the original fall population (Table
9). In 1975, 13 reward bands were recovered from the 57 placed on roosters
earlier in the fall and 1 reward banded rooster from 1974 recaptured in
fall of 1975. This represented 22.41 percent harvest of the original early
fall population.
Projections would place the total banded harvest in 1975
at 13 reward bands and 12 regular bands. Twenty-three reward bands were
recovered by field check and mail-in in 1976 from 144 roosters previously
reward marked (including two 1976 retrapped reward banded roosters) in

�-22-

fall. This represented 15.97 percent bagged harvest.
These harvest
percentages are based on the number of roosters originally fall banded
and those retrapped in a specific fall banding period. The actual
number of banded or retrapped roosters remaining alive at the beginning
of subsequent hunting seasons remained unknown.

Table 9. Comparative harvest and crippling loss indices for the three
years of study on the mortality study areas.

Sample

1974

1975

1976

Retrieved harvest based on
reward band recoveries

18.5%

22.4%

16.0%

Crippling loss as a percent
of retrieved harvest

10.3%

8.2%

28.7%

Projected

20.4%

24.2%

20.6%

1.67

2.50

2.16

total mortality

Average season bag per hunter-1/

l/ The majority

of hunters queried as to success in post-season surveys
were those who were successful or were in successful parties so the
average season bag per hunter was biased upward.

Crippling Loss and Total Harvest Mortality -- Table 9 summarizes
estimates of bagged harvest, crippling loss and total harvest mortality
through the three years of study. Crippling loss was based on information
from hunter questionnaires which totaled 52 in 1974, 34 in 1975, and
43 in 1976. Loss was reported as 10.3 percent of the retrieved bag in
1974, 8.2 percent in 1975, and 28.7 percent in 1976 .. These figures
contained considerable potential for error. The much higher rate of
loss reported in 1976 was partially due to information from one hunter.
He reported 10 cripples for 12 birds bagged.
Even without his data,
the loss rate approximated 21 percent, well above that for previous
years.
Crippling loss, as a function of fall density, approximated
1.9 percent in 1974, 1.8 percent in 1975 and 4.6 percent in 1976.
Snow conditions, prevailing during the 1975 season, were believed
responsible for the reduced crippling loss that year in spite of
slightly increased harvest.
Dry, cold, windy conditions potentially
increased crippling loss in 1976. Similar conditions were reported
for bobwhite on the Tamarack Wildlife Area by Snyder (1977).
The rate of harvest, including crippling loss, never exceeded 25
percent of the original early fall population during any of the three
years of study based on reward band return projections.
This was in
part due to low hunting pressure, but was more a function of low pheasant

�-23-

densities.
Studies in other locations show that harvest estimates ln
areas of high pheasant density and high hunter use can approach 70 to
90 percent or more of available roosters. These references and projected
rates of harvest are summarized in Table 10.
There was no evidence in any study listed in Table 10 that high rooster
mortality in any way impeded reproductive success the following year.
The low rate of harvest achieved in eastern Colorado undoubtedly is of
no significance to subsequent reproduction.
Vast fields, low pheasant
densities, the abilities of roosters to learn to avoid hunters with
season progression, and the rapid decline of hunter effort with season
progression preclude achieving a much greater percent harvest than is
currently obtained. Increased pheasant density, however, is the one
factor which could increase percent harvest, but then only to a limited
degree.

Table 10. Percentage harvest of the rooster segment of pheasant populations
as reported in the literature from several states.ll

Reference

State

Percent Rooster Harvest

This study

Colorado

15-25

Allen (1947)

Michigan

49-76

Bizeau (1964)

Idaho

50

Edwards (1963)

Ohio

76

Farris (1973)

Iowa

71

Harper (1960)

California

49-56

California

53

Hart (1954)

California

48

Hartman and Sheffer (1971)

Pennsylvania

88-93

Robertson

Illinois

60

Stokes (1954)

Ontario

78-93

__

Utah

76-88

Wisconsin

59-84

, Hart &amp; Shaffer (1951)

(1958)

(1968)

Wagner, Besadney and Kabat (1965)

II

- Most researchers used change-in-sex-ratio
harvest.

data to calculate percent

�-24-

Braun (1975) stated that most estimates of harvest in past studies
(Table 10) were based on change-in-ratio methods between the sexes
rather than on banding information.
Results from this study provide
evidence leading one to question this basis of derivation of harvest
estimates.
To illustrate, trapped ratios projected 77.5 roosters per
100 hens in the fall 1974 population.
Hunting, including crippling
loss, removed 20.4 percent of the roosters, which would change the
post harvest ratio to 61.7 roosters per 100 hens (assuming no hens were shot).
Late winter trapping showed 22.7 roosters per 100 hens indicating 39
additional roosters per 100 hens were unaccounted for due to unknown
causes. Observed winter sex ratios showed even fewer roosters present in
late winter than trapped samples did (Table 3). Loss per 100 hens totaled
54.8 roosters or 70.7 percent of the original fall rooster population.
Projections of rooster harvest based on reward banded data approximated
less than a third of that which would have been indicated by change-inratio methods alone.
Findings for the second segment of the study followed a pattern similar
to that of the first year. There were 71.8 roosters per 100 hens in the
fall 1975 trapped sample but only 31.9 roosters per 100 hens in the observed late winter sample. Total harvest mortality based on return of
reward bands was placed at 24.7 percent of the trapped population which
would reduce the number of roosters in post season populations to 54.1
per 100 hens. Twenty-two additional roosters per 100 hens were unaccounted
for in relation to hen loss for the fall winter period of 1975-76.
Again, as in the previous year, banding data indicated a much lower
harvest than change-in-ratio data showed.
The reason or reasons for these apparent greater unaccounted for losses
of roosters based on banded data remain unknown.
Illegal poaching, nonreporting of reward bands, potential for movement (at least in 1974-75),
inaccurate sex ratio data, or differentially greater predation of males
than females are all possibilities, singularly or in combination.
Whatever the case, change-in-ratio methods used alone, would probably have
assigned this additional rooster mortality or unaccounted for loss to
harvest. More intensive study using telemetry methods, controlled access
hunting units or other modifications would be needed to derive answers
to this unknown variable.
Differential Harvest of Adults and Juveniles -- Hoffman (1975) showed
evidence that adults were less susceptible to hunter harvest than juveniles. In 1974, it was suggested that 19.5 percent of the immature
roosters were bagged and only 14.6 percent of the adults were retrieved.
Figures in 1975 projected a 20.0 percent harvest of adults and a 23.4
percent harvest of juveniles.
In contrast, in 1976, 16.7 percent of
the adult roosters were taken compared to a 15.7 percent harvest of
juveniles.
Sample sizes were not large anyone year, especially for
adults, and definite conclusions cannot be drawn on this limited amount
of data.

�-25-

Hunter and Landowner Comments Regarding Hunting Seasons
Questionnaires were sent to hunters contacted in the field and those who
mailed in bands. A number responded with unsolicited comments during
the three years and the general results are summarized in Table 11.
Very few comments were made in 1975 when snow conditions exposed available
birds. A majority of those returning questionnaires in 1976 made comments.
Lack of pheasants and lack of cover were the most frequently mentioned
items shown in Table 11. Several commented on increased hunting pressure
and lack of birds, however, hunting pressure actually was declining.
Several requested that the hunting season be closed or shortened to let
the population build up, and several felt the Division had "taken'! them
for a license when no birds were present. Other less frequently mentioned
items included predator control, hunter access difficulties, and stocking.
Farmers in the study region, contacted prior to and during fall trapping,
usually also expressed unsolicited opinions. They were nearly unanimous
in requesting hunting season closure. A few wanted the season shortened
and a curtailment of hunting pressure.

Table 11. Unsolicited comments provided by hunters on questionnaires
following the 1974 through 1976 hunting seasons.

General Commen t

Number of hunters making statement

Lack of pheasants and poor hunting

35

Saw lot of birds or hunting was good

4

Lack of cover noted

18

Wanted hunting season closed

12

Hunting season was too long

2

Had difficulty

4

finding place to hunt

Wanted pheasant stocking

4

Wanted predator control (bounty on coyotes)

4

Wanted license increase to develop cover

2

Wanted Division to do something to
increase birds

2

Other comments included develop roadside cover, acquire more public
land, and reduce poaching by local farmers.

�-26-

Movements
Average movements from the location of fall trapping to location of
harvest among recovered roosters was 4.7 miles in 1974, 4.6 miles in
1975 and 1.3 miles in 1976. The midpoint of fall trapping came in
mid-September, approximately two months prior to the beginning of
hunting seasons.
Some errors could have resulted from incorrect
reporting by hunters, but some high movements were evident.
Three
movements in 1975 ranged from 15 to 20 miles, however, all other
movements from banding to harvest were less than four miles in
distance that fall and most were less than two miles.
Similar results
were evident in 1974. Movements tended to be directed toward the east
and south in 1974 and 1975. Potentially, these were from areas containing mostly wheat fields to areas containing some irrigated corn.
This is supported by lack of excessive movement in 1976 where most of
the trapping effort was concentrated adjacent to cornfields because
pheasants were lacking in locations containing only wheat.
In addition,
there was no evidence of a distinct south and east direction of movements in 1976.
Some movements might have resulted from trapping disturbance, however,
it is more probable that such movements were a natural occurrence.
Directional movements in 1974 and 1975 tend to support this hypothesis.
Environmental
Environmental

Variables

Impacts and Mortalities

of Production

The four-township area originally established for this study was nearly
all flat to gently-rolling wheatland.
Pheasant densities in the region
appeared, by census, to be higher than in most other parts of northeast
Colorado.
First year (1974) late summer-fall trapping efforts revealed
that most birds were concentrated, if not produced, in association with
the dry to intermittent drainages traversing the region. Sections
lacking drainages and associated unfarmed waste land usually contained
few if any pheasants.
Areas containing draws, especially if corn and
wheat fields were adjacent, usually contained higher concentrations of
birds. Potentially, these unfarmed drainages were an important variable
for nesting in a region almost entirely devoted to dryland wheat.
Rainfall in 1974 was deficient in quantity with approximately 11 inches
received through the year. Nearly four inches of this was received
during one day, June 8, and most of the remainder of that summer was
dry. Quantity of precipitation is not the only variable to be considered
when nesting and brood survival is involved.
Distribution through the
growing season and whether moisture comes as gentle rain or violent
thunderstorms are additional variables to be considered.
Information
is lacking on the impact of these and other variables as to how they
affect reproduction.
However, both production census and fall trapping
indicated 1974 was a poor summer for pheasant production because large
numbers of hens without broods were evident in late summer. Data from

�-27-

Snyder (1970) show that the number of young in fall was primarily a
function of the percentage of successfully nesting hens, and this
seemed to be the situation in 1974. Much additional research along
these lines of study is needed before we can gain insight into pheasant
nesting and brood survival as affected directly by precipitation and
indirectly through precipitation impact on vegetation and insect abundance.
Fewer hens without broods were found in late summers of 1975 and 1976
than in 1974 so reduced numbers of young in the latter years was attributed
directly to inadequate breeding populations.
Age ratios indicate nesting
success was better in both 1975 and 1976 than in 1974 (Table 6). Below
average rainfall persisted through the summer and fall and nearly all
pheasants found during fall 1976 trapping were proximal to irrigated
cornfields. Whether these birds were produced in wheatfields adjacent
to cornfields or migrated to these adjacent fields after hatching is
unknown. One can speculate that free water there permitted better
nesting success and/or brood survival, but this is uncertain.
Hens
may have also concentrated there after movements to wintering sites
during the previous fall.
Environmental

Variables of Survival

Three major storms, on March 1975, November 1975, and March 1977, hit
the region during the period of study. The two March storms were
especially devastating to pheasants. The first, in March 1975, began with
an evening rain which soaked and chilled exposed birds. The rain quickly
changed to snow and severe high winds and snow persisted through the next
day accompanied by lowering temperatures.
Thousands of livestock were
lost in the storm impacted area along with a majority of the pheasants
and other wildlife in the region.
The storm of March 1977 began in a manner somewhat similar to that of
the storm two years earlier. Winds were even higher and much more snow
was received, however temperatures did not drop much below freezing.
This difference in temperature was the primary factor reducing mortality
impact on livestock and wildlife during the second storm. Post storm
searches revealed few dead pheasants, but failure to find many live birds
was probably more indicative that the storm had caused considerable
mortality to pheasants. Dead birds were both difficult to find and
quickly consumed by predators while access into ~he study region was
delayed and difficult during both storms.
The snow storm of November 1975 reportedly caused some pheasant mortalities
as indicated by both farmers and hunters, however dead birds were not
observed during field searches. Persistent drifting snows made field
searches difficult.
The percentage mortality from that storm was
believed much lower than the mortality resulting from the two March
storms in the study region.
Under normal winter conditions and following winter storms, pheasants
concentrate and use windbreaks and shelterbelts for protective cover.

�-28-

These tree and shrub plantings are considered a vital element in wintering
pheasant habitat and Lyon (1958) noted heavy winter use by pheasants.
Storms such as those of March 1975 and 1977 apparently caught the pheasants
in wheat stubble fields, where they died of exposure or suffocation (Fig. 8).
However, had the birds been concentrated in shelterbelts during either
storm, nearly all would have been buried under huge drifts. Lyon (1958)
noted that pheasants apparently abandoned shelterbelts during the blizzard
of 1949 when such cover was needed most.
Severe winter storms did not
reoccur during subsequent years of his evaluations so he was not able
to make further assessments of the value of windbreaks during blizzards.
Observations following the March storms of 1975 and 1977 indicate additional
windbreaks probably would not have increased pheasant survival.
Much
taller wheat stubble, especially in 1975, probably would have had greater
impact on survival by providing increased protection from wind and snow.
Figure 8 illustrates the way many of the pheasants were found after the
March storms.
They apparently climbed onto the snow to prevent being
buried and died of exposure and/or suffocation.
Some walked or attempted
to fly into. the wind and ended up in bare fallow fields along the north
edge of stubble fields.
Annual Mortality

Rates

The relative recovery method was used in assessing annual mortality rates
based on band recoveries obtained during harvests (Funk 1966). The
calculated average annual mortality rate was 97.23 percent for the interval
sampled. Unfortunately, sample sizes were small and little confidence
could be placed in results.
However, whether or not the projected mortality
rate approximated the real situation, rooster mortality was high during
the period of study. Most of this mortality was due to natural causes.
As hens were not legal game, only retrap information during second and
third fall intervals (Table 8) was available for assessing annual mortality.
A procedure substituting retraps for shot recoveries (Table 13) was used
in estimating the annual mortality rate of 75.28 percent for hens. As
with roosters, samples were inadequate and little confidence can be placed
in proj ections.
High annual mortality rates obtained during this study do not typify
those of a stable pheasant population.
Annual mortality and increment
of young must be near identical over a period of years if population
stability is maintained.
For example, if the annual mortality rate averaged
70 percent from one September to the next, then 70 percent of the September
popUlation would have to be young. Assuming fall trapped samples were
accurate indicators of adult to young ratios, then the percent young
ranged from 64.3 to 69.6 percent during the three year period. To maintain a stable population, the annual mortality from fall to fall should
have been within this general range. Since hens were not hunted and
roosters were, differences in survival and recruitment between sexes
would be expected and were evident.
Young roosters comprised from 79
to 83 percent of the male segment in fall trapped samples while young
hens comprised from 44 to 62 percent of the female segment. Neither

�-29-

Fig. 8. A mortality from the March 1975 blizzard.
Numerous pheasants were
found throughout the study region like this in wheat stubble fields.
(Photo
by Warren D. Snyder)

�-30group of samples approached mortality levels derived in Tables 12 and 13.
Mortality at the 70 percent level from fall to fall would have established
near stable populations in 1975 and 1976 on this study area. Thus, since
indicated mortality rates were much higher than recruitment rates, the
population declined over the 1974-76 period. Past studies have placed
annual mortality of stable pheasant populations from 70 to 80 percent
(Mallette and Harper 1964; Leopold et al. 1943; Buzz 1946; Dumke and
Pils 1973; Harper 1960; and others).

Table 12. Relative recovery rate method for estimating annual mortality
and survival of roosters within the mortality study area.

Fall
Banded

Number
Marked

Recoveriesl/
l-n
2-n

Recovery Rate
l-n
2-n

1974

233

2

.0086

1975

109

17

1976

142

22

0

.1560

survival rate

Average mortality

rate

Mortality
Rate

.0051

.9449

.0000

1.0000

.0000

.1549
.3109

Average

Survival
Rate

.0086

= 2.77 percent
= 97.23 percent

1/ l-n = all recoveries
r~coveries

from the segment banded in that year; 2-n = all
from the segment banded that year except 1st year recoveries.

Table 13. An estimation of annual mortality and survival of hens
substituting retraps for recovered mortalities based on two years of retrap.

Fall
Banded

Number
Marked

1974

301

5

1975

150

4

1976

187

Retraps
1st yr. 2nd yr.

Retrap Rate
1st yr. 2nd yr.

2

rate

= 24.72 percent

Mortality

rate

= 75.28 percent

Mortality
Rate

.2472

.7528

.0066
.0267

Survival

Survival
Rate

�-31-

CONCLUSIONS AND RECOMMENDATIONS
1. Until an improved technique is developed, nightlighting appears to
be the only feasible and practical method of capturing significant
numbers of pheasants in the Tablelands of eastern Colorado. Use of an
auxilIary flood light system is essential in applying this technique.
Trained crews can capture most observed pheasants in late summer and a
majority of observed birds in fall and winter. Deficient population
levels were the primary deterents to trapping success during this study.
2. Reward bands placed on a portion of the roosters clearly indicated
that hunters failed to report a majority of regular bands on pheasants.
The same situation would undoubtedly occur with quail and other upland
game bird species. Therefore, reward bands are an essential part of
any study where mail-in reporting of bands is used as a basis for
deriving harvest, annual mortality, or other information.
3. Poor reproduction and subsequent devastating storms curtailed obtaining
both adequate banding and recovery samples during this period of study.
A study using telemetry methods is about the only way additional mortality
information can be obtained under existing very low pheasant density
levels. Such a study should be directed toward determining when, how
and to what degree hen mortality occurs and its impact on reproductive
success.
4. Study findings show that less than a fourth of the roosters initially
fall banded were taken by hunters during any of the three years of study.
The Division should utilize this information, contrast it with harvest
data from other states, and attempt to (1) educate the farming, hunting,
and general publics that hunting of roosters has no impact on subsequent
reproduction and populations; and, (2) point out the real elements, weather,
farming practices and other environmental variables which have reduced
populations to near zero in much of the State. This education effort
must be long term and repetitious. However, it is doubtful that percentage
harvest can be increased significantly while pheasant populations remain
at current very low levels. The reason why change~in-ratio methods
projected much higher unaccounted for rooster loss than harvest band
return information is uncertain. Additional research is needed, again
by use of telemetry.
5. Results from this study clearly indicate that both the hen and rooster
segments of northeastern Colorado pheasant populations sustain high annual
mortality turnover rates. High productivity and high mortality go hand in
hand. Weather variables in accompaniment with short wheat stubble and
generally deficient cover were the primary factors in the drastic population declines which occurred during the period of study. Additional windbreak and shelterbelt plantings would have done little to increase survival
during this period because of the severe storms which occurred. However,
they would be valuable cover in more normal winter periods.
6. Many hunters, faced with lack of birds to hunt and resentment of their
presence by local farmers, voiced opinions that the Division should have
closed the hunting season. Nearly all farmers in the region voiced the

�-32-

same opinion.
It is the Division's duty to inform these publics as to
what pheasant population levels can be expected and that they should not
anticipate quality hunting conditions.
At the same time the Division
must stress to both the hunting and farming publics that removal of
roosters has no impact what-so-ever on subsequent populations and reproduction.
Basic simple explanation and understanding is needed. At the
same time, marginal pheasant populations can provide limited recreational
opportunity for those persevering enough to try.
LITERATURE

CITED

Allen, D. L. 1947. Hunting as a limitation to Michigan pheasant populations. J. Wildl. Manage.
11(3):232-243.
Bizeau, E. G. 1964. Results of hen pheasant seasons in Idaho.
Western Assoc. State Game and Fish Comms. 44:211-215.

Proc.

Braun, C. E. 1975. Mortality, survival and effects of hunting on grouse,
partridge, pheasants and quail - An annotated bibliography.
Colorado
Division of Wildlife Rept. No.3.
39 p.
Buzz, I. o. 1946. Wisconsin
Publ. 326. 184 p.

pheasant populations.

Wisc. Cons. Dept.

Dumke, R. T., and C. M. Pils. 1973. Mortality of radio-tagged pheasants
on the Waterloo Wildlife Area. Wisc. Dept. Nat. Resources, Tech.
Bull. No. 72. 52 p.
Edwards, W. R. 1963. Proportionate harvest of ring-necked
cocks and hens. Game Res. in Ohio 2:11-22.

pheasant

Farris, A. L. 1973. The ring-necked pheasant in Iowa - 1972. Iowa
Conservation Commission.
Wildl. Research Bull. #7. 38 p.
Funk, H. D. 1966. Review of duck literature relating to population
dynamics and banding analysis techniques and findings.
Colo. Dept.
of Game, Fish and Parks. Game Res. Rept. Oct. p. 77-108.
Harper, H. T. 1960. The effects of a three year limited hen season on
pheasant populations in California, 1955-1957.
Proc. Western
Assoc. State Game and Fish Comms. 40:168-176.
, C. M. Hart, and D. E. Shaffer.
1951. Effects of hunting pressure
-----and game farm stocking on pheasant populations in the Sacramento
Valley, California, 1946-1949.
Calif. Fish and Game 37(2):141-176.
Hart, C. M. 1954. Pheasant hunting pressures in California.
Western Assoc. State Game and Fish Comms. 34:219-226.

Proc.

Hartman, F. E., and D. E. Sheffer.
1971. Population dynamics and hunter
harvest of ring-necked pheasant populations in Pennsylvania's primary
range. Trans. N. E. Section of the Wildl. .Soc . 28:179-205.
Hoffman, D. M. 1975. Pheasant mortality investigation.
Wildl. Game Research Rept. April p. 5-35.

Colo. Div. of

�-33-

Labisky, R. F. 1968. Nightlighting: its use in capturing pheasants,
prairie chickens, bobwhites, and cottontails.
BioI. Notes No. 62.
Illinois Natural History Survey, Urbana.
12 p.
Lyon, L. J. 1958. Evaluation of effects of habitat impr6vement on
wildlife.
Colorado Dept. of Game and Fish. Pittman-Robertson Qtly.
Progress. Rept., April. pp. 75-113.
Mallette, R. D., and H. T. Harper.
1964. Population studies of ringnecked pheasants in California.
Calif. Fish and Game. 50(4):292-304.
Robertson, W. B., Jr. 1958. Investigations of ring-necked pheasants in
Illinois.
Ill. Dept. Cons., Div. Game Manage. Tech. Bull. No.1.
137 p.
Sandfort, W. W. 1960a. Pre-nesting studies. Colorado Game and Fish Dept.
Quart. Res. Prog. Rept., April. p. 37-47.

--- , 1,960b. Pheasant brood survey.
Res. Prog. Rept., April.

Colorado Game and Fish Dept.
pp. 49-55.

Quart.

Snyder, W. D. 1970. Pheasant hen harvest investigation.
Colorado Div.
of Game, Fish and Parks. Game Res. Rept., April. pp. 3-81.
Stokes, A. W. 1954. Population studies of the ring-necked pheasants on
Pelee Island, Ontario. Ontario Dept. Lands and Forests Tech. Bull.
Wildl. Ser. No.4.
154 p.
___

1968. An eight-year study of a northern Utah pheasant population.
J. Wildl. Manage. 32(4):867-874.

Wagner, F. H., C. D. Besadny, and C. Kabat.
and management of Wisconsin pheasants.
No. 34. 168 p.

1965. Population ecology
Wisc. Cons. Dept. Tech. Bull.

Wishart, W. 1969. Age determination of pheasants by measurement
proximal primaries.
J. Wildl. Manage.
33(3):714-717.

Prepared by ~'~_/.~'_~_~~~~~~~
Warren D. Snyder
Wildlife Researcher

_
C

of

��April 1977

-35-

JOB PROGRESS REPORT

State of

COLORADO
--~----~~~~~----------

Project No.
Work Plan No.

W-37-R-30

Game Bird Survey

3

Job Title

Job NOo
9
_
Evaluation of the Effects of Changes
~l~·n~H~u~n~t=i~n~g~R~e~g~u~l~a~t~l~·o~n~s~o~n~S~a~g~e~G~r~o~u~s~e~P~o~p~u~l~
_

Period Covered:

April 1, 1976 through March 31, 1977

Personnel:
Fred Giese, U. S. Fish and Wildlife Service; Heather F.· Alexander,
Don Benson, Clait Braun, Deborah Covic, Courtney Crawford, Sue Erickson,
Howard Funk, Ken Giesen, Don Gore, Paul Gorenzel, John Hobbs, Richard
Hoffman, Jim Jackson, Fran Marcoux, Sig Palm, Brett Petersen, Steve Porter,
Brent Renfrow,Wayne Russell, Howard Spear, Robert Stark, and John Wagner,
Colorado Division of Wildlife.

ABSTRACf
Investigations concerning the relationships of hunting to breeding population
levels, and mortality and survival rates of sage grouse (Centrocercus
urophasianus) in North Park, Colorado initiated in 1975 continued in 1976.
Numbers of known strutting grounds increased as two new grounds were located.
Average number of cocks per ground increased from 31 in 19.75 to 32 in 1976.
Peak period of female attendance in 1976 was between April 5-15, about 10
days earlier than in 1975. Total estimated spring population size was about
3,600-4,200 birds. In 1976, 384 (239 males, 145 females) sage grouse, including 5 chicks, were banded. Ninety-two broods were observed with a composite
average of 2.0 chicks per successful hen. Estimated nesting success based
on 184 females seen during the brood period was 50.0%. A 9 day permit only
hunting season was allowed in North Park in 1976 with bag and possession
limits of 3 and 6. Data from wings (N = 504) collected at check stations,
wing barrels and field checks revealed that the harvest was comprised of 42
percent young of the year, 24 percent yearlings and 34 percent adults.
Calculated hatching dates for 211 hunter harvested juveniles ranged from late
May to 19 July with a peak between 8-21 June: Estimated nesting success
from examination of wings of adult and yearling females was 43 percent with
more adults (53%) than yearlings (27%) being successful.
Ovarian analysis
(N = 118) indicated that 75% of the yearling and 88% of the adult females
had ovulated.
Total permits issued numbered 1,258, and only 77 percent of
the permittees actually hunted. Estimated total harvest including crippling
loss was 829 grouse. This was a decrease of approximately 21 percent from
1975 harvest levels. Fifty-eight bands were recovered and reported by hunters
of which 37 were from 1976 bandings, 14 were from 1975, 4 were from 1974,
while 3 were from 1973 bandings.
Females (11) were poorly represented in the
harvested sample of banded birds. Direct recovery rates were 13.3% for
yearling males, 14.0% for adult males, 1.7% for yearling females and 1.8%
for adult females. Harvest of banded males was not uniform by area of banding
with percentage of cocks harvested vs total cocks counted varying from 0 to
16% per strutting ground. Survival and mortality rates for each age and sex
class were calculated with males having lower survival rates than females.
Estimated fall population size was about 7,000 birds and the harvest rate
was about 12%.

�-36-

RECOMMENDATIONS

1.

Trapping efforts should be intensified with more effort being made to
capture females in spring and females and chicks in August and early
September.

2. Changes in hunting regulations should be promulgated with a 16 day
season and bag and possession limits of 3 and 6 in 1977.

�-37-

EVALUATION OF THE EFFECTS OF CHANGES
IN HUNTING REGULATIONS ON SAGE GROUSE POPULATIONS
C1ait E. Braun
Sage grouse are widely distributed in the western United Stat~s, occupying
sagebrush dominated rangelands.
Throughout most of their occupied range,
state wildlife agencies routinely collect different types of population
data for the species and regulate hunter opportunity and harvest.
Management of sage grouse has primarily been based on counts of cocks on strutting
grounds in April and estimates of nesting success and brood size obtained
in July and August.
Unfortunately, evaluation of existing data gathering
procedures and knowledge of the impacts of hunting have been and are limited.
Data presented in this report concern these problems and represent the
period of mid-March 1975 through mid-March 1977.
P.N.O. OBJECTIVE
The objectives of this study are to increase hunter opportunity and recreational use of sage grouse in North Park, Colorado.
Hypotheses which are
being tested are:
a.

Hunter harvest does not affect spring counts of strutting males or peak
spring counts of females under any regulations.

b.

Counts of females during the peak of attendance on strutting grounds
are a reliable index to overall numbers of sage grouse in spring.

c.

Hunter success and total harvest are a function of August precipitation.

d.

Nesting success and brood size to 15 August, while important in determining age composition of the harvest, have no relationship to hunter
success and total harvest.

e.

Differences exist between yearling
and nesting success.

and adult heps in percent ovulating,

SEGMENT OBJECTIVES
1a.

Counts of males will be made on 10 selected strutting grounds three
times each week between 5:00 to 7:30 A.M. during the month of April.

lb.

Counts of females will be made on 10 selected strutting
day for 10 days in April (period of peak attendance).

2a.

Males will be trapped and marked with color-coded bands at night while
roosting on or near strutting grounds.
A sample of 200 will be marked.
Trapping will be conducted from mid-March to late May by spotlighting
and netting.
Bandings will be evenly distributed in four areas of
North Park.

grounds every

�-38-

2b.

Females will be trapped during the early A.M. on strutting grounds
during April through use of drop, mist and cannon nets. A sample of
300 will be marked with color coded leg bands. Bandings will be evenly
distributed in four areas of North Park.

2c.

Chicks will be located and trapped in brood areas during late July and
August through use of tape recorded calls and noosing.
Some chicks may
be caught through use of mist nets and drive traps. A sample of 200
chicks (100 males and 100 females) will be marked with color coded leg
bands or wing tags. Bandings will be evenly distributed in four areas
of North Park.

3a.

Nesting success will be estimated through counts of successful and
unsuccessful hens in July and August.
Successful females will be located
through use of tape recorded calls, while unsuccessful hens will be
located and counted in known summer use sites. A sample of 200 hens
will be classified.

3b.

Brood size will be determined by counts of chicks accompanied
in July and August.
A sample of 100 broods will be counted.

4.

August precipitation will be determined from 12 rain gauges placed at
selected sites throughout sage grouse habitat.
Sites will be selected
that typify particular areas within North Park. Each rain gauge will
be sampled once a week. Data collected will be compared to that collected by the U. S. Weather Bureau at Walden and at Spicer.

5.

Population size will be estimated in spring and fall. Spring population
size will be estimated through use of ratios of males to females on
strutting grounds, observation and recaptures of marked birds and ratios
of males to females and juvenile to yearling to adult age classes in
the harvest the preceding fall. Fall (pre-hunting) population size will
be estimated through examination of data on nesting success, brood size
to 1 September and spring population data. Independent fall population
estimates will be made through use of data collected at check stations
on age composition of the harvest and number of marked birds shot.

6a.

Hunting will be by free permit only, and all hunters will be mailed a
questionnaire within one week of end of hunting season. One follow up
letter will be sent two weeks later. A goal of 90 percent response
is desired.

6b.

Check stations will be operated at three locations each year during
the first two weekends of the hunting season. These stations will be
at Gould, State Line, and Willow Creek Pass. A goal of 500 wings is
desired.

6c.

Number and location of marked birds shot will be obtained through use
of check stations, field hunter checks and voluntary mail reporting.
A
goal of 50 first year recoveries is desired.

6d.

Hunter success will be determined through data collected
stations and from the mail questionnaire survey.

by hens

at check

�-39-

6e.

Age composition of the harvest will be determined through examination
of wings collected at check stations.
Classification will be as adult,
yearling, juvenile, and male and female.

6f.

Total harvest will be estimated (90 percent confidence
mail questionnaire survey of all hunters.

7.

Different regulations will be promoted each year, with 9 days and a
bag and possession limit of 3 and 6 in 1976.

8.

Compile data, analyze results, and prepare progress

level) from the

report.

METHODS AND MATERIALS
Counts of male and female sage grouse present on strutting grounds were
periodically made by project personnel and Wildlife Conservation Officers
following proscribed procedures (Braun and Beck 1976). Counts were made
from 23 March through 19 May at times within one-half hour of sunrise.
Searches for new or relocated grounds were made through ground and aerial
reconnaissance.
Male and female sage grouse were trapped at night along
roads and on strutting grounds where they roosted.
Some birds, primarily
hens, were captured at dawn on strutting grounds through use of cannon nets.
Most birds were captured through use of a spotlight with a back-pack power
source and long handled nets (Brau~ and Beck 1976). Sage grouse banded and
released were marked with serially numbered aluminum bands (size 16 for
males, 14 for females) and colored plastic and aluminum bands coded to
capture locations.
Age and sex classification of birds captured followed
Eng (1955). Some chicks were captured in August through use of a truck
mounted cannon net (Lacher and Lacher 1964).
Hens with broods were located during July and August through systematic
searches of selected areas and observations along roads and trails.
Systematic searches for broods involved walking moist areas and playback of tape
recorded chick distress calls (Braun et al. 1973, Wallestad 1970).
Searches
for broods along roads and trails were made from a vehicle following established routes (Gill 1965) and other suitable appearing areas. Routes were
traversed shortly after sunrise at a speed of 15 to 20 miles per hour.
Twelve
sunken or pit rain gauges which were checked weekly through early
September were randomly established throughout the sagebrush range in North
Park. Each gauge was constructed from two one gallon (3.8 liter) cans
soldered together.
Splash shields were made from 1 yard square (approx. 1 m2)
burlap.
Rodent -and insect screens of 0.1 cm (1/4 in) wire mesh were installed
about 20 cm below the surface of the ground in each 15.2 cm (6 in) diameter
can. Cetyl alcohol was used instead of diesel fuel to retard water evaporation.
Sage grouse hunters: in North Park in 1976 were required to have in their
possession a free numbered permit. Permits unlimited in number were available from Division of Wildlife offices in Fort Collins and Denver, all
license agents in North Park, and all project and management personnel
assigned to work in North Park during the hunting season. Questionnaires
were sent to all permittees immediately following the sage grouse season in
North Park, with one followup letter being mailed in mid-October to all
non-respondents.

�-40-

Check stations were operated at three locations (State Line, Gould, Willow
Creek Pass) on the opening weekend (II and 12 September) and the second
Sunday (19 September) at Willow Creek Pass. Each station was operated from
about 1000 to 1800 MDT, depending upon traffic load. Data obtained per party
were: county of origin, number of hunters, hours hunted (total of all hunters), birds observed, birds bagged, birds lost, number of banded birds and
location of where each was obtained, and area hunted.
One wing was obtained
from each bird checked with ovaries being taken from a sample of hens.
Efforts were made to ascertain sex by gonadal inspection for a sample of
birds.
These data were recorded on tags with wings being individually marked
with corresponding information concerning actual sex of that bird. Wings
were frozen and stored for later analysis.
In addition to the three check stations, a wing barrel and sign (Hoffman and
Braun 1975) were placed along Colorado 14 near Muddy Pass on 11 and 12
September, and near Walden Reservoir on Jackson County Road 12 from 13 through
19 September.
Collected wings were thawed and classified to age (chicks, yearlings, adults)
and sex following procedures outlined by Eng (1955), and Beck et al. (1975).
Hatching dates were calculated for chicks of the year using data from Pyrah

(1963).
Ovaries collected were stored in an alcohol, formalin and acetic acid solution
and later examined for presence of ovulated follicles as described by Kabat
et al. (1948).
Description

of Area

North Park lies completely in Jackson County, Colorado.
This relatively high
altitude (7,800 to 8,500 ft) area is essentially a closed basin, as it
is encircled by mountain ranges.
Drainage is to the north, with lowest
elevations occurring north of Cowdrey.
Vegetation is principally a sagebrush
(Artemisia spp.)-grassland type with stream courses being dominated by
native grasses, sedges and deciduous shrubs.
Detailed geographic, geologic,
vegetational and climatic features of the area have been described by Gill
(1965), Carr (1967), Beck (1975), Braun and Beck (1976) and will be treated
in the final report.
RESULTS AND DISCUSSION
Strutting

Ground Counts

Counts of male and female sage grouse present on strutting grounds were
initiated on 23 March (Alkali Lake, Boettcher Lake Junction, Railroad,
Walden, and Wattenburg #2) and continued until 19 May (Owl Creek).
During
this interval, 252 counts were made of 22 active grounds.
Maximum counts
of males and females and dates obtained are presented in Table 1. The total
of 712 males counted is higher than the 588 counted in 1975 but represents
two new grounds (Bighorn and Owl Creek) and one ground (Delaney Butte) which

�-41-

was active for a short time in 1976. Average number of cocks per ground
in 1976 was 32.4 while this average was 30.9 in 1975 and 27.7 in 1974.
These data suggest a slight increase in breeding population from 1974
through 1976. One ground inactive in 1975, Delaney Butte, was active briefly
in 1976, while two new grounds (Bighorn and Owl Creek) were located in 1976.
It is probable that other grounds exist, especially near Delaney Butte.
Two small grounds were located near Baker Draw and Case Flats in 1976. Due
to the small size «10 cocks each) and sporadic use of these grounds, they
have not been listed as strutting grounds.
The importance of the high counts of females (total = 599) in 1976 is unknown.
This number is slightly above the total of high counts of hens (573) in
1975. Due to logistic problems in counting all strutting grounds on the
same days during the peak of mating, it is probable that counts of hens will
not be useful for other than estimating onset of nesting and expected appearance of broods.
The peak period of female attendance in 1976 was between
5 and 15 April.
Three or more counts of males in a seven day period were made at 15 different
grounds in 1976. Number of different seven day periods represented per
ground varied from 1 to 4. Counts of males at four grounds did not change
in a seven day period while counts at the remaining 11 grounds varied from
-66 to +100 percent within a seven day interval.
Time of counts in relation
to peak of female attendance and size of ground in terms of numbers of cocks
did not appear to be of major importance in controlling numbers of cocks
present on individual grounds.
It is suspected that predator activity prior
to sunrise near the grounds and intrinsic factors affected by weather may be
most important in controlling numbers of cocks on a given ground on any
given day.
Estimation

of Spring Population

Size

Reliable estimates of total numbers of sage grouse in North Park during the
breeding period are difficult to derive. Major problems involved are those
relating to how many males and females of the total population are present
on strutting grounds when peak counts are attained and how many of the actual
number of active strutting grounds are known and counted.
There is some
evidence to suggest that only 50 percent of the male segment of the population
of lekking species is present on a ground when peak counts of males are
attained (Robel 1969, Rippin and Bogg 1974). If it is assumed that peak
counts of male sage grouse represent 50 percent of the male population in an
area, then at least 1,424 (712 x 2) cocks occurred in North Park in the
spring of 1976. It is quite obvious that not all strutting ground locations
are known in a given year. If it is assumed that 80 percent of the active
grounds have been located, then a minimum of 4 (.20 x 22) unknown grounds
should occur somewhere in North Park. Assuming that these 4 grounds have
an average of 32 cocks during the peak period of male attendance, they
represent a minimum of 256 additional cocks (32 x 4 = 128 x 2 = 256). Thus,
the total estimated number of cocks in North Park in the spring of 1976 was
1,682. This is higher than the 1,486 males estimated to have been in North
Park in the spring of 1975. Realistically, these figures may be higher than
actual numbers of birds present. A better estimate for both 1975 and 1976
might be 1,200 ~ 200 males in the spring population.

�-42-

Table 1.

Sage grouse strutting

ground counts, North Park, 1976.

Maximum
No. of
Males

Ground

Date(s)

Maximum
No. of
Females

Date(s)

Alkali Lake

39

23 March

7

3 April

Bighorn

46

24 April

14

15 April

62

20 April

96

10 April

Canuck

30

11 April

20

11 April

Coalmont

28

28 April

34

16 April

Cowdrey 115

5

5 and 13 April

6

5 April

Deer Creek

36

25 and 30 April

18

14 April

Delaney

Butte

43

29 March

0

24 March
through 4 May

Fish Hatchery

81

24 April

44

14 April

Hound

27

10 April

60

9 April

Lost Creek 111

33

22 April

6

21 April

Monahan

1

5 April

0

1 through 28
April

Owl Creek

14

11 May

1

11 May

Railroad

87

31 March

69

14 April

Ridge Road

32

29 April

46

12 April

Riley

18

8 April

39

8 April

Roth

9

4 April and 4 May

7

12 and 16 April

Spring Creek 111

49

28 April

78

8 April

Spring Creek 112

11

10 April

14

15 April

Spring Creek 114

8

10 April

6

11 April

Walden Reservoir

37

5 April

20

5 April

16

23 March

14

5 April

Boettcher

Draw

Wattenberg

Totals

Lake Jct.

112

712

599

�-43Beck (1977) found a late winter sex ratio of 69 hens to 31 cocks, a value
remarkably similar to the sex ratio of adult and yearling birds in the fall
harvest (Braun and Beck 1976). If it is assumed that the spring sex ratio
of the North Park sage grouse population is similar to that observed in late
winter and early fall, then 2 hens occur for every cock in the population.
Thus, the total spring female population can be estimated based on the estimated number of cocks in the spring population.
Therefore, there should
have been a minimum of 3,364 hens in the 1976 spring population (1,682 x 2 =
3,364). If the figure of 1,200
200 cocks is used, then the spring female
population should have been 2,400 ± 400 birds. The estimated total spring
population of sage grouse in North Park in 1976 was 5,046 birds if actual
projections are used. If the figure of 1,200 + 200 cocks is used, the
estimated 1976 spring population is 3,600 + 600 birds.
It is reasonably
safe to assume that the 1976 spring population of sage grouse in North Park
was between 3,000 and 5,000 birds.

±

Capture and Banding
Intensive efforts to capture and band sage grouse in the 1976 field period
were initiated on 22 March. During the 22 March through 19 May interval,
376 birds were newly banded. An additional 8 birds were newly banded in
the 12 through 26 August period. Age and sex data for the banded sample are
given in Table 2.

Table 2. Sex and age composition
North Park, Colorado 1976.

of newly banded sage grouse by month,

1+

Males
2+

March

4

2

6

1

1

2

April

80

91

171

54

59

113

May

35

21

56

15

13

28

Month

Chick

Total

Chick

Females
1+
2+

Total

August

5

1

0

6

0

1

1

2

Totals

5

120

114

239

0

71

74

145

The total of 384 birds banded in 1976 was less than the 422 birds banded in
1975. As in 1975, the goal of 200 males (100 of both age classes) was
achieved while the goal of 300 females was not. The 145 females banded in
1976 was the largest number ever banded in one year in North Park.
Distribution of bandings within North Park in 1976 was not equal as only 25
grouse were banded in the northeast portion of the park, 50 were banded in
the southeast, while 116 were banded in the northwest and 193 were banded in
the southwest quadrant. Most trapping was associated with strutting grounds,

�-44-

consequently, few birds were trapped in the northeast part of the park, as
only two known strutting grounds occur in this area. The same is true for
the southeast portion of the park, as only 5 active grounds are known in
this area. Distribution of the banded sample in 1976 as in 1975 reflected
known distribution of strutting grounds.
Distribution of strutting grounds
may reflect quality of available habitat but may also be a reflection of
access.
Sex and age composition of the trap sample does not reflect the actual
compostion of the population.
Males are more easily trapped than females
as they are associated with strutting grounds at night for up to two months.
Of the sample of 145 females banded, 56 were trapped along roads at night
with the remainder (91) being trapped on or near strutting grounds (28
with cannon nets).
The even age ratios of males and females apparent~n
the 1976 trapping results are more a reflection of grouse behavior patterns
than the actual age compositon of the population.
In the course of trapping operations, 58 different birds (50 males, 8
females)'banded in previous years were recaptured.
Six (5 males, 1 females)
had been banded in 1973, 9 (8 males, 1 female) had been banded in 1974,
while 43 (37 males, 6 females) had been banded in 1975. The oldest males
recaptured were at least 5 years of age (2) as was the oldest female (1).
It was not possible for birds to be known to be older than 5+ years as no
banding of sage grouse was accomplished in North Park in the 1970-1972
interval.
Little use can be made of recapture data.outside of longevity
and movements as efforts were made not to recapture banded birds.
This was
done to reduce possible mortalities of banded birds.
Summer Observations
Systematic examination for brood patches (area of bare skin at the posterior
end of the abdomen)
on females captured was conducted from 23 April through
19 May. The first indication of a brood patch was observed on 27 April on
1 of 3 females checked.
After 1 May, only 1 female of 32 examined did not
have a partial or full brood patch. These data suggest that some hens
initiated incubation the last week of April, while all nesting hens had
initiated laying and incubation prior to 19 May 1976. 'Consequently, the
earliest nest could have hatched (assuming a 25 day incubation period) on
about 20-25 May.
Searches for hens with chicks were made by walking selected transects and
using playbacks of tape recorded chick distress calls and by slowly driving
roads in preselected areas. The first brood was observed on 25 June and
consisted of 4 (3-4 weeks of age) chicks. Groups containing progeny from
more than one brood were first observed on 22 July. During the 1 June-30
August period, 551 sage grouse were observed including 177 young of the
year. Classification of birds observed by time period is presented in
Table 3.

�-45-

Table 3.
1976.

Composition

of observed

sage grouse, North Park, 1 June-3l August,

Category

1-15
June

16-30
June

l-l~
July

16-31
July

1-15
August

16-31
August

Total

Adult males

8

29

37

24

3

89

190

Adult females

5

12

12

55

38

62

184

Chicks

0

16

26

36

17

82

177

3.2(5)

3.7(7)

2.6(14)

4.2(4)

1.3(62)

1.9(92)

Chicks/successful
female 1/

1/ Hens accompanied
parenthesis

by chicks were assumed to be successful.
represent number of assumed successful hens.

Numbers

in

Sage grouse younger than 3 to 4 weeks of age are difficult to locate because
of dense cover used and behaviours which cause them to freeze if danger is
apparent.
Several (3) females which appeared to be broody were observed in
early June. These hens responded to the tape recorded chick distress call
but no chicks could be located. All were assumed to have had broods.
Apparent
brood sizes throughout the summer appeared to be lower than expected if sage
grouse lay from 7 to 9 egg clutches.
It is probable that broods observed
before mid-July are under counted because of dense cover and associated
behavior patterns.
Brood breakup after mid-July and size similar to females
may result in inaccurate counts of young later in the summer.
It presently
appears that brood counts of sage grouse give unreliable estimates of production because of problems associated with visibility, behavior patterns and
size.
Because of the above problems it is difficult to accurately assess nesting
success and production from counts of hens with and without broods.
Data
presented in Table 3 suggest that 92 of 184 females (50%) were successful,
with each successful hen rearing two chicks to age of independence (8 weeks+).
It is exceedingly doubtful that counts of sage grouse after mid-August and
possibly after 31 July have any merit.
It appears that moisture patterns and
dryness of vegetation after these dates, expecially mid-August, have more
effect on numbers of birds observed than does production success.
Precipitation
Twelve standard pit rain gauges as described by Alexander (1976) were again
placed at randomly located sites in North Park in 1976. Gauges were checked
weekly from 6 August to 13 September.
Unfortunately, few useful data were
obtained because of changes in observers (some gauges were not properly
located), animal disturbance, and evaporation of the medium used to supposedly

�-46-

prevent evaporation.
All 12 gauges gained and lost moisture and were
disturbed at least once in the study period. Consequently, data from this
source were not used.
Data from the Walden and Spicer stations maintained by the Weather Bureau,
U. S. Department of Commerce are presented in Table 4.

Table 4.

Weather data, North Park, Colorado,

1976.

SEicer

Walden

June

July

August

20

31

80

No. days 32 F
or below

1-10
Sept.

June

July

August

25

22

34

25

84

63

80

86

81

11

1

12

7

1

7

Days of ppt.

8

8

17

5

7

15

Total ppt. (in)

1.15

1.20

1.23

30 year average ppt. (in)

1.37

1.22

1.69

0

Min. temp. F
0

Max. temp. F

1-10 1/
Sept.-

0

2
.33

.67

.95

1.37

1.11

1.24

1.29

1/ Observer died, no data.
From data in Table 4 it is readily apparent that the June through August 1976
period in North Park was somewhat drier than the 30 year average.
Total precipitation was down about 16 percent (16.4) in the southwestern part of North
Park (Spicer) and about 18 percent (17.9) in the central area of the park
(Walden). However, these decreases were offset by frequent showers (15 and 17
days out of 31 possible) in August 1976. As a consequence, desiccation of
vegetation was not noticeable in late summer and large concentrations of birds
around moist sites did not occur.
When the 200 square miles of hay meadows in North Park are considered (Braun
and Beck 1976) the edges of which are all potential grouse use areas, it
would appear that summer precipitation patterns may have little impact on
grouse distribution in fall. Since most hay meadows are on private land and
much of the private area is closed to hunting, it is logical reasoning that
a sizeable portion of the sage grouse population in North Park is not available to hunters in the fall. Thus amount of private land, and behavior
patterns of grouse may be more important than summer precipitation in affecting
harvest of grouse in North Park.

�-47-

Hunting

Season Data Collection

The sage grouse season in North Park (Unit 12) in 1976 opened at sunrise on
11 September and closed at sunset on 19 September.
Season length was identical to that in 1975 (9 days) but longer than those seasons prior to 1975
(normally 3 days).
The daily bag limit was three with a possession limit of
six. This was more liberal than the two and four daily bag and possession
limits allowed through 1975. As in 1974 and 1975 all hunters hunting sage
grouse in North Park were required to have in their possession a free special
permit.
Permits unlimited in number were available at all license agents in
North Park; the Fort Collins and Denver offices of the Division of Wildlife,
and all Wildlife Conservation Officers and research personnel working in North
Park. Purpose of the permit was to obtain reliable estimates concerning hunter
activities in North Park.
Check Stations
Three check stations were operated in 1976. These stations were located at
the State Line, Gould, and Willow Creek Pass. As in previous years of
operation, each station was open from about 1000 to 1800 MDT, depending upon
traffic load. All check stations were operated on the 11th and 12th, with
Willow Creek also being operated on the 19th. At least one research person
and two Wildlife Conservation Officers were assigned to each check station.
Data obtained per party were: county of origin, number of hunters, hours
hunted (total all hunters), birds observed, birds bagged,. birds lost, number
of banded birds and location where each was obtained, and area hunted within
North Park. Most importantly, one wing was obtained from each bird that was
checked.
Few birds (9) were completely wingless.
Ovaries were collected
from 121 females, and sex by gonadal inspection was ascertained for 304 birds.
In addition to the three check stations, a wing barrel and sign were placed
along Colorado 14 near Muddy Pass on September 11 and 12, and near Walden
Reservoir on Jackson County Road 12 from September 13 through 19. Twentytwo wings were received in the wing barrel on September 11 (10), and 12 (12)
while 2 were received from September 13 through 19.
During the three days of check station operation, 595 hunters with 459 sage
grouse were checked.
These hunters reported observing 3,393 sage grouse.
Some duplications are undoubtedly present in the 3,393 observations.
Hunter
efficiency was low (13.5 percent) (459 birds harvested 7 3,393 birds observed)
but only 26 birds (5.7 percent of those retrieved) were reported crippled
and lost. Hunter efficiency was higher than in 1974 (7.7%) and 1975 (9.6%)
while crippling loss was less than in 1975 (7.1%) but similar to that calculated in 1974 (5.1%).
Distribution of the harvest and hunting pressure within North Park was uneven
in 1976. Leading harvest zones were Lake John, Ridge Road, and Peterson RidgeMacFarlane Reservoir areas each with about 21 percent of the total harvest.
Next in importance were Walden Reservoir (12.0%), Owl Ridge-Spring Creek (6.3%),
Michigan River Southeast (5.9%), Eagle Hill (5.0%), Pole Mountain (4.5%), and
Independence Mountain (3.0%). Point of harvest could not be ascertained for
3.9 percent of the birds checked.
Considering the three years for which

�-48-

location of harvest within North Park has been determined, harvest trends
'are downward at Walden Reservoir, Eagle Hill, Owl Ridge-Spring Creek and
essentially stable in the other zones. With the exception of the Ridge Road
area (13.0% of the hunters harvested 21.1% of the birds checked), distribution of hunters in 1976 coincided with distribution of the harvest.
Most hunters contacted at check stations were asked whether or not they
normally hunted in North Park. Of the 565 hunters responding, 366 (63.8%)
reported that they normally hunted in North Par~ 159 (28.1%) were first
time sage grouse hunters, while 40 (7.1%) reported they normally hunted
elsewhere.
Percentage response to this question has not varied significantly
since inception of the intensive study in 1974 (Table 5).

Table 5. Previous sage grouse hunting experience
North Park, Colorado 1974-1976.

of sage grouse hunters,

Year

Normally Hunt
In North Park
Number
Percent

1974

183

68.3

20

7.4

65

24.3

1975

432

64.9

47

7.0

187

28.1

1976

366

64.8

40

7.1

159

28.1

Normally Hunt
Elsewhere
Number
Percent

First Time
Sage Grouse Hunters
Number
Percent

Two important points are apparent from these data. First, there is a substantial annual turnover in sage grouse hunters and second, more liberal
seasons in North Park (1975 and 1976) have not resulted in major shifts of
hunters into North Park from other areas.
Wing Analyses
In 1976 wings were received from 504 sage grouse (450 from check stations,
24 from wing barrels, 20 from field checks and 10 from the wing envelope
survey). Age and sex structure of the harvest as estimated from wings from
1974 through 1976 is presented in Table 6.
As can be readily ascertained from data in Table 6, sex and age compostion
of the harvest in 1976 was almost identical to that in 1975 and was not
greatly different from that determined in 1974. Contrary to expectations
derived from observations of broods in July and August, production of young
did not increase in 1976. It is obvious that no selection by hunters for
smaller birds (females) occurs in the young-of-the-year age class even though
immature males are larger by 400-500 grams (1 pound) by time of the hunting
season.
Selection for smaller birds (hens) in the older age classes is
possible, but is highly unlikely considering hunter opportunity, hunter
efficiency and the late winter sex ratio of 62 females:38 males.
The sex

�"

Table 6.

Age and sex composition of the sage grouse harvest, North Park, Colorado 1974-1976.

Year

Males
No.
%

Immatures
Females
No.
%

Total
No.
%

Males
No.
%

Yearlings
Females
No.
%

Total
No.
%

Males
No.
%

Adults
Females
No.
%

Total
No.
%

1974

171

48.9

179

51.1

350

50.1

49

35.5

89

64.5

138

19.8

45

21.4

165

78.6

210

30.1

1975

101

47.6

111

52.4

212

42.0

52

46.8

59

53.2

111

22.0

55

30.2

127

69.8

182

36.0

1976

104

49.5

108

50.9

212

42.1

46

38.7

73

61.3

119

23.6

49

28.3

124

71.7

173

34.3

I
.po
\0

I

�-50-

ratio in the harvest for all three age classes combined was 62 fema1es:38
males, 59 fema1es:41 males, and 60 fema1es:40 males for 1974-1976, respectively.
From examination of data in Table 6 it is evident that overwinter
survival of immatures to yearling and yearling to adult classes has not
changed markedly in the three year period, nor has the overwinter survival
of males or females. What small changes occurred were probably the result
of numbers of chicks produced the preceding year. It is also evident that
survival is disproportionate favoring females.
Consequently, turnover is
fastest in the male segment of the population, with complete turnover probably
occurring every 5 years for males and 7 years for females.
Examination of the molt of primary wing feathers of immature sage grouse (N =
211) harvested in 1976 indicated hatching started in late May, peaked between
8-21 June and continued as late as the week of 13-19 July. This is a .somewhat
earlier schedule (7-10 days) than that of 1975, but similar to the schedule
observed in 1974. Of interest is the fact that a major storm occurred in
North Park in the 13-18 June period, with up to 5 inches of snow. This storm
undoubtedly affected nesting success.
Wings of 192 females (adult = 121, yearling = 71) were classified as to wing
molt. An estimated 43.2 percent (adults = 52.9%, yearling = 26.8%) of the
hens in this sample were successful in nesting.
These data are similar to
those obtained in 1975 but lower than in 1974. The young to adult hen
(including yearling hens) ratio in the harvest was 1.1:1, identical to that
in 1975, but lower than the 1.4:1 ratio in 1974. The young per successful
hen ratio in 1976 was 2.5:1.
Ovarian Analysis
Ovaries were collected from 118 hunter harvested sage grouse in 1976 and were
analyzed following procedures described by Meyer et ale (1947), Kabat et a1.
(1948), and Buss et ale (1951). Data for 1976 and for the 99 ovaries from
1975 that were reanalyzed are shown in Table 7.

Table 7.

Analysis

of sage grouse ovaries, North Park, Colorado

1975 and 1976.

Age Class

Percent Ovulating
1975 !/
1976

Number Ovulating/Examined
1975 !/
1976

Yearling

96.9

75.0

32/33

33/44

Adult

96.8

87.8

61/63

65/74

!/ A series of ovaries classified as not ovulating or questionable was sent
to I. o. Buss for examination.
The information for these ovaries is not
available

causing the 1975 data to be biased upward.

�-51-

Considering only the data from 1976, it is apparent that differences exist
between yearlings and adult sage grouse in percent ovulating.
Adults are
more likely to ovulate than yearlings suggesting that some sage grouse
females are not sexually mature before one year of age. Considering some
of the difficulties in evaluating ovaries two or more months after egg laying,
the accuracy of the ovulated follicle technique may be minimal.
Hunter Questionnaire
A total of 1,258 permits was issued for hunting sage grouse in North Park in
1976, a decrease of 283 (18.4%) from 1975. As in previous years, most hunters
(46.7%) were from the metro Denver area (Adams 6.7%, Arapahoe 7.6%, Denver
17.1%, Jefferson 16.3%). Only 8.4 percent of the hunters originated from
Jackson County, with 21.5 percent from Larimer County, 7.6 percent from
Boulder County, and 3.8 percent from Weld County. All other counties each
contributed less than 2 percent of the total hunters and only 12 percent of
all hunters.
Questionnaires were sent to all permittees immediately following the sage
grouse season in North Park (24 September).
Responses were received from
861 permittees.
On 15 October a fo1lowup letter was sent to all nonrespondents and 222 additional responses were received.
In all, 1,083 permittees
(86.1%) responded (Table 8). Fourteen questionnaires were undeliverable,
the lowest number since 1974 (12). Mean values calculated for permittees
responding to the followup letter were used to project for the 13.9 percent
(175) non-respondents.
From data in Table 8 it is apparent that 77.5 percent of all permittees went
hunting, similar to the 77 percent who hunted in 1975, but lower than the 81
percent that hunted in 1974. Actual numbers of hunters were similar in 1974
(960) and 1976 (975) but lower than in 1975 (1,187). Only 33.4 percent of
the actual hunters were successful in 1976. This is somewhat lower than in
1975 (38.3%) but substantially less than in 1974 (53.9%). Total harvest in
1976 was 829 birds (783 bagged, plus 46 crippled and lost). This is a substantial decline from the 1,053 birds harvested in 1975, and 1,174 birds
bagged in 1974.
Time period of hunting was received for 961 hunters ~ho harvested 677 birds.
Based on this sample, 83.9 percent of harvest occurred during the opening
weekend, 5.0 percent occurred during the week, while 11.1 percent occurred
during the second weekend.
These data are similar to those obtained in 1975,
the only other year with a 9 day season.
Since total harvest decreased
approximately twenty percent (21.3) in 1976 it is not possible to calculate
how many more birds were harvested under a 3 and 6 season than a 2 and 4
season. However, in 1976 only 47 hunters (28 = 1 day, 19 = 2 days) achieved
the bag limit. This is in contrast to 1975 when 106 hunters (62 = 1 day,
39 = 2 days, 1 = 3 days, 4 = 4 days) achieved the bag limit. Obviously, it
was more difficult to get 3 birds than it was 2. Also, birds per successful
hunter did not change markedly in 1976 (2.4) from levels calculated in 1974
and 1975 (2.2 each year). These data do not suggest that a more liberal
bag limit coupled with a 9 day season increases the harvest.

�-52Table 8~ North Park sage grouse hunter questionnaire data, 1976.
Projected
For

Projected
For 1,258

175

1,258

No. in sample

861

222

Percent of total
permittees

68.4

17.6

86.1

13.9

100.0

No. of hunters

701

153

854

121

975

Percent hunters

81.4

68.9

78.9

68.9

77.5

No. of non-hunters

160

69

229

54

283

Percent non-hunters

18.6

31.1

21.1

31.1

22.5

No. of successful
hunters

238

49

287

39

326

Percent successful
hunters

34.0

32.0

33.6

32.0

33.4

No. of hunter days
Days/hunter
No. of sage grouse
bagged

1,192

1,083

269

1.70

564

1,461

1.76

123

213

1.71

687

1,674

1.76

96

1.72

783

Grouse/permittee

.66

.55

.63

.55

.62

Grouse/hunter

.80

.80

.80

.80

.80

Grouse/successful
hunter

2.4

2.5

2.4

2.5

2.4

No. of grouse lost

40

4

44

2

46

Crippling loss/
hunter
Total kill

.06
604

.03
127

.05
731

.03
98

.05
829

Percent crippling
loss

6.62

3.15

6.02

3.15

5.5

Percent success of
permittees

27.6

22.1

26.5

22.1

25.9

Banded birds

43

9

52

7

59

Non-deliverable
surveys

14

o

o

o

14

Percent nondeliverable

1.11

�-53-

Hunter pressure and sage grouse harvest in North Park as derived from the
Small Game Questionnaire Survey are not presently available for 1976.
Results of this survey in 1974-75 are presented in Table 9. This survey
over-estimated hunter numbers by 83 percent and total kill by 114 percent
in 1974 and hunters by 35 percent and kill by 87 percent in 1975.

Table 9. Comparative data, number of sage grouse hunters and total harvest,
North Park, 1974-1976.

Year

Number of Hunters
Statewide Survey
This Study

Number of Sage Grouse Harvested
Statewide Survey
This Study

1974

960

1,759

1,174

2,509

1975

1,187

1,600

1,053

1,973

1976

975

829

Band Recoveries
Fifty-eight bands were reported from birds harvested or found freshly dead
during the 1976 hunting season. Of this number, 11 were from females
(1973 = 2, 1974 = 1, 1975 = 4, 1976 = 4), while 47 were males (1973 = 1,
1974 = 3, 1975 = 10, 1976 = 33). Banding and recovery data are presented
in Table 10, while survival and mortality rates are shown in Table 11.
Data presented in Tables 10 and 11 indicate that survival rates are different
between age class and sex. These differences are most pronounced for males.
The direct or first year recovery rate in 1976 was 13.3 percent for yearling
males, and 14.0 percent for adult males. Direct recovery rates for yearling
males were similar in 1975 and 1976 but higher than those calculated in
1973 (7.5%) and 1974 (11.0%). Direct recovery rate~ for adult males have
varied from 6.5 percent (1975) to 14.0 percent (1976). Direct recovery rates
for adult (1.8%) and yearling females (1.7%) were similar in 1976. Direct
recovery rates for females are lower than males each year, possibly a result
of small sample sizes. However, it is quite obvious that males are more
readily harvested than females.
This supports the hypothesis that hunters
do not select for smaller birds.
In fact, some selection may occur for
larger birds.
Harvest of males banded on various strutting grounds in 1976 varied from 0
(Canuck, Cowdrey #5, Owl Creek, Roth. Spring Creek #1, and Spring Creek #4)
to over 20 percent (Alkali Lake, 5 of 11 = 45.5; Hound, 1 of 4 = 25.0;
Riley, 1 of 3 = 33.3; Wattenburg #2, 2 of 5 = 40.0; Bighorn, 5 of 19 = 26.3;
Spring Creek #2, 1 of 1 = 100.0). Five other grounds had from 11 to 17
percent of the cocks banded in 1976 harvested (Boettcher Jet., 1 of 9 = 11.1;
Fish Hatchery, 3 of 25 = 12.0; Coalmont, 1 of 7 = 14.3; Railroad, 7 of 45 =
15.6; Lost Creek #1, 2 of 12 = 16.7). Six known grounds (Canuck, Cowdrey #5,
Owl Creek, Roth, Spring Creek #1, and Spring Creek #4) had no banded birds
reported shot in 1976. With the exception of Canuck (30 males) and Spring
Creek #1 (49 males), these six grounds had peak male counts of less than 15
birds.

�-54-

Table 10.

Sage grouse banding and recovery data, North Park, 1973-1976.

Year

Age

No.
Banded

1973

1+

80

6

4

6

1

41

2

2

0

1

2+

91

7

3

0

0

54

5

1

3

1

179!/

13

82:./ 72:./

1

10~/

7

3

3

2

Total

1974

Total

]j
]j

Females
Number Recovered
1973 1974 1975 1976

54

6

5

2

22

2

1

1

2+

88

8

5

1

27

2

0

0

142

1r}:/

10

3

49

4

1

1

1+

138

18

5

62

6

2

2+

153

10

5

68

6

2

291

28

10

l30

13!!./

4

Total
1976

No.
Banded

1+

Total

1975

Males
Number Recovered
1973 1974 1975 1976

1+

120

16

71

2

2+

114

16

74

2

234

32

145

4

Includes 8 unknown age males.
Includes 1 unknown age male at time of banding.

1/ Includes 14 unknown age females.
!!)

Includes one unknown age female (aluminum band
not present when recovered).

�-55-

Table 11. Survival and mortality rate estimates
North Park by age and sex, 1973-1976.

Average

Survival Rates

for sage grouse banded

Average

Mortality

Yearling males

.561

.439

Adult males

.337

.663

All males

.463

.567

.579

.421

Adult females

.558

.442

All females

.5l~/

.484

Yearling

females

in

Rates

!/ The calculated

survival rate for all females was lower than for either
yearling or adult females because of inadequate banded samples in 1974
and small numbers of recoveries from those samples.

Considering all recoveries in 1976 of males banded from 1973 to 1976, a
minimum of 6 (15.4%) of a high count of 39 males was harvested from Alkali
Lake strutting ground, 8 (9.9%) of a high count of 81 from Fish Hatchery,
3 (11.1%) of a high count of 27 from Hound, 4 (12.1%) of a high count of 33
from Lost Creek #1, 8 (9.2%) of a high count of 87 from Railroad, and 5
(10.9%) of a high count of 46 at Bighorn.
All other grounds had two (3
grounds) or one (5 grounds) banded cocks reported shot. It is not presently
known at what level of harvest decreases will occur in peak numbers of cocks
counted on grounds the following spring.
Percentages of harvest from 3 to
15 would not appear to be excessive.
If only recoveries from 1976 bandings are examined, it is possible that the
45.5% harvest of cocks from Alkali Lake strutting ground may affect counts
in the spring of 1977 at that ground.
The high recovery of cocks banded on
smaller strutting rounds, i. e. Hound (25.0%), Riley (33.3%), and Wattenburg
112 (40.0%) may depress counts at those grounds in 1977.
In 1975 an estimated 21 and 25 percent of the cocks counted at Hound and
Riley strutting grounds were harvested (Braun 1976). The high count of
cocks at Hound in 1976 decreased by 18.2% from 1975 levels (27 vs 33) while
at Riley the high count increased by 50% (18 vs 12). These data are
inconclusive.
The percentage of banded birds in the adult and yearling male and female
segments of the fall population was calculated from check station data (sex
and age compostion), harvest questionnaires (total harvest), and band
recoveries.
The calculated percentage of adult and yearling males banded in
the fall population was 29.5 (95 7 504 = 18.8% x 829
156, 46 7 156 = 29.5%)
while for adult and yearling females it was 3.4 (197 7 504 = 39.1 x 829 = 324,
11 7 324 = 3.4).

�-56-

Estimation

of Fall Population

Size

Estimates of fall population size are difficult to derive, primarily
because of unknown vulnerability of chicks and inexact survival estimates
for all age classes, especially females. Provided that all birds of each
age and sex class are equally available and vulnerable to hunters and using
the average mortality rates for all males and all females listed in Table 11
the 1976 fall population size in North Park was calculated as follows:
Number of Birds Banded in
1973
1974
1975
1976

Total Banded Birds Alive:

Alive in 1976
Males
Males
Males
Males

18
30
135
234

Females
Females
Females
Females

.15
.13
67
145

Males

= 417

Females

= 240

Of the 417 banded males alive in 1976, 46 were harvested, while 11 of the
240 banded hens were harvested.
If 58 percent (check station data) of the
829 birds harvested were adults and yearlings, then 481 of the birds
harvested were older than one year. Of this number, 57 (11.9%) were banded,
thus there should have been about 4,040 adult and yearling birds alive in
North Park in the fall of 1976 (if 11.9% = 481 birds, then 100% = 4,040 birds).
Immature birds comprised 42 percent (check station data) of the harvest, thus there should have been about 2,927 chicks in the 1976 fall population.
The total population in the fall of 1976 should have been about
6,967 or 7,000 birds. This estimate is realistic but may be low.
The overall harvest rate was about 12% (829 7 7,000) with adult and yearling
males having a higher harvest rate (11.0% estimated, 13.7% direct recovery
rate) than adult and yearling females (4.6% estimated, 2.8% direct recovery
rate).
Of the 292 adult and yearling birds for which wings were available, 197
(67.5%) were from females.
Thus, of the approximately-4,040
adult and yearling birds in the preseason population, 2,727 (4,040 x 67.5 = 2,727) were
hens. Thus, total chicks in the fall population in 1976 based on 1.1 chicks
per hen in the harvest (1.1 x 2,727 = 3,000) approximated 3,000. This number
is only slightly higher than the 2,927 chicks estimated from overall age
ratios and band recoveries in the 1976 harvest to have been in the fall population in 1976. These data do not suggest that chicks are more vulnerable to
hunting than are adults and yearlings.
It is interesting to compare estimates of adults and yearlings alive in the
spring and fall. Spring estimates based on strutting ground counts indicate
that about 5,000 (5,046) birds were present, while fall estimates based on
age and sex ratios in the harvest and band recovery data indicate that about
4,000 (4,040) adult and yearling birds were present.
This is a difference
of about 1,000 birds.
It is possible that spring estimates are incorrect
as either (1) more or fewer strutting grounds are unknown than estimated, or
(2) peak strutting ground counts represent more than 50 percent of the cocks

�-57-

present in an area. It is also possible that fall estimates based on
harvest age and sex ratios and band recoveries are low. It does appear
that spring populations levels were somewhere between 3,000 and 5,000 birds
in North Park in the spring of 1976.

LITERATURE

CITED

Alexander, H. F. 1976. Evaluation of the effects of changes in hunting
regulations on sage grouse:nesting success and brood investigations.
Colorado Div. Wildlife, Prog. Rept., Fed. Aid Proj. W-37-R, Work Plan 3,
Job 9, Part 2. pp. 103-117.
Beck, T.D.I. 1975. Attributes of a wintering population of sage grouse,
North Park, Colorado.
M. S. Thesis. Colorado State Univ., Ft. Collins.
49 pp.
1977. Sage grouse flock characteristics
winter.
J. Wildl. Manage. 41(1):18-26.

and habitat selection

in

_____ , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of
sage grouse from wing characteristics.
Game Inf. Leaflet No. 49
(revised). Colo. Div. Wildlife.
4 pp.
Braun, C. E. 1976. Evaluation of the effects of changes in hunting regulations on sage grouse populations.
Colorado Div. Wildlife, Prog. Rept.,
Fed. Aid Proj. W-37-R, Work Plan 3, Job 9, Part 1. pp. 85-102.
_____ , and T.D.I. Beck. 1976. Effects of sagebrush control on distribution
and abundance of sage grouse. Colorado Div. Wildlife, Final Rept., Fed.
Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84.
_____ , R. K. Schmidt, and G. E. Rogers.
tailed ptarmigan with tape-recorded
90-93.

1973. Census of Colorado whitecalls. J. Wildl. Manage. 37(1):

Buss, I. 0., R. K. Meyer, and C. Kabat. 1951. Wisconsin pheasant reproduction studies based on ovulated follicle technique.
J. Wildl.
Manage. 15(1):32-46.
Carr, H. D. 1967. Sage grouse and sagebrush control.
rado State Univ., Fort Collins.
106 pp.

M. S. Thesis.

Colo-

Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios from
wings. J. Wildl. Manage. 19(2):267-272.
Gill, R. B. 1965. Distribution and abundance of a population of sage grouse
in North Park, Colorado.
M. S. Thesis. Colorado State Univ., Fort
Collins.
187 pp.

�-58-

1975. A volunteer wing collection
Hoffman, R. W., and C. E. Braun.
3 pp.
Game Inf. Leaflet No. 101. Colo. Div. Wildlife.

station.

Kabat, C., I. O. Buss, and R. K. Meyer.
1948. The use of ovulated follicles
in determining eggs laid by the ring-necked pheasant.
J. Wildl. Manage.
12(4):399-416.
Lacher, J. R., and D. D. Lacher.
Manage. 28(3):595-597.

cannon net trap.

J. Wildl.

Meyer, R. K., C. Kabat, and I. O. Buss. 1947. Early involutionary
in the post-ovulatory follicles in the ring-necked pheasant.
Manage. 11(1):43-49.

changes
J. Wildl.

pyrah, D. G.
Wildlife
71 pp.

1964.

A mobile

1963. Sage grouse investigations.
Idaho Fish and Game Dept.,
Restoration Div., Job Compl. Rept., Fed. Aid Proj. W-125-R.

Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations
sharp-tailed grouse.
J. Wildl. Manage. 38(4):616-621.
Robel,
R. J. 1969. Movements
of blackcocks in Scotland.

and flock stratification within
J. Anim. Ecol. 38(5):755-763.

of male

a population

Wallestad, R. O. 1970. Summer movements and habitat use by sage grouse
broods in central Montana.
M. S. Thesis. Montana State Univ., Bozeman.
51 pp.

Prepared

by__

.L..&lt;a",,-,~~_·~_.

-"~=:::::.::..-

Clait E. Braun
Wildlife Researcher

_

�April 1977

-59-

JOB PROGRESS REPORT

State of

COLORADO
----------~~~~-----------

Project No.

W-37-R-30

Game Bird Survey

Job Title

Job NO.__~~~~l~O~~
Investigations of the Distribution and Status
of Sagebrush and Sage Grouse in the Moffat County Area

Period Covered:

April 1, 1976 to March 31, 1977

Work Plan No.

Personnel:

3

_

Clait E. Braun, Charles E. Brown, .Iohn F. Corey, Howard D. Funk,
Richard W. Hoffman, William T. Howard, Roland C. Kufeld,
Annabell Leckler,- Robert L. Mangus, Deirdre W. Plummer, William
I. Roland, Louis D. Vidakovich, Claude E. White, Charles W.
Woodward, and Donald M. Hoffman.

ABSTRACT
Approximately three percent of the Federally administered lands in Moffat
County has been altered by intentional habitat manipulations.
This has been
done almost exclusively on BLM lands. Most range conversions -have been
through spraying (39.2%), compared with plowing and seeding (25.0%), chaining
(19.5%), reseeding (14.6%), burning and r~seeding (1.2%), and rotobeating
(0.5%). A sample of 20 landowners within prime sage grouse ranges, owning
10.3 percent of the total deeded land in Moffat County indicated they have
altered approximately 25.2 percent of their deeded lands in order to bring
land under cultivation or to improve pastures for grazing purposes.
These
individuals plan to alter an additional 1.7 percent of their land within the
next ten year period, if economic conditions are favorable.
A total of 965
sage grouse were counted on 27 active strutting grounds in Moffat County
during the spring of 1976. An average of 31.92 cocks per ground was counted
on 24 grounds where sexes were determined.
Three new grounds were located
and counted and six old grounds, which no longer were counted by Division of
Wildlife personnel, were relocated.
A total- of 564 sage grouse were counted along ten trial brood routes in Moffat and western Routt counties in August
1976. A total of 412.6 miles (663.9 km) were driven on these routes for
an average of 1.37 birds per mile (.85/km).
Seventeen broods counted along
these routes averaged 5.29 young per brood. A young-to-adult ratio of 2.37:1
indicated that sage -grouse experienced good reproductive success in 1976 in
the Moffat County area. A total of 748 sage grouse wings were collected
from successful hunters through the use of 16 wing barrel collection stations
during September 11-13, 1976. Wing molt patterns of breeding-age females
revealed that 97 of 132 (73.5%) adults and 49 of 74 (66.2%) subadult hens
were probably successful nesters in 1976. Thus, nesting success and production were also determined to be good in the Moffat County area in 1976
through use of the wing collection data. A minimum of 1,223 sage grouse
were counted on 18 different wintering areas during the period December, 1976
through March, 1977. At least ten of these wintering areas are considered
to be major wintering areas within the study area. An average of 67.9 sage
grouse were counted per wintering area.

��-61-

INVESTIGATIONS OF THE DISTRIBUTION AND STATUS
OF SAGEBRUSH AND SAGE GROUSE IN THE MOFFAT COUNTY AREA
Donald M. Hoffman

P.N.O. OBJECTIVES
Major objectives of this study are to: (1) ascertain the distribution of
sagebrush in the Moffat County area, (2) determine the status (i.e. principally alterations) of sagebrush in this area, (3) delineate the distribution of sage grouse in the Moffat County area, and (4) identify-~the past
and present trend of sage grouse population status in this area.
SEGMENT OBJECTIVES
1.

Review literature concerning
the Moffat County area.

vegetative

descriptions

from studies

in

2.

Ascertain the distribution of sagebrush through examination of vegetative maps and aerial photographs from public_ land management agencies.
Check delineations through field reconnaissance and assemble composite
vegetative maps.

3.

Ascertain
agencies,

4.

Interview selected landowners to determine extent of past, present,
and planned alterations of sagebrush ranges.

5.

Delineate distribution of sage grouse through examination of records of
various agencies, interviews of landowners, and field reconnaissance.

6.

Maintain records of sage grouse observed by date and time, locations,
and vegetative types.

7.

Collect and store materials from road killed sage grouse and others
found dead for detailed laboratory analyses.

8.

Ascertain trends in sage grouse populations through examination of data
from management surveys.
Give areas where populations have seemingly
disappeared particular attention.

9.

Systematically count sage grouse on known strutting
rearing areas, and winter concentration areas.

10.

Locate new strutting
aerial searches.

alterations in sagebrush communities from records of various
aerial photographs, and field reconnaissance.

grounds and wintering

grounds,

brood

areas through ground and

�-62-

11.

Obtain harvest data through use of wing collection
cally located throughout the area.

stations,

strategi-

12.

Compile data into maps and/or overlays
report.

Prepare

progress

and tables.

METHODS AND MATERIALS
Searches to locate literature concerning
in the Moffat County area were continued
Fort Collins.

vegetative descriptions from studies
in the DOW and CSU libraries in

Major areas of sagebrush range within the Moffat County area were mapped on
0.5-in-per-mi county highway maps utilizing Soil Conservation Servic~ Range
Resource Maps, traverse and sketches of range vegetative types in the field,
and sketches of range vegetative types from aerial photographs and mosaics.
Records of locations, amounts, and types of past range alterations by federal agencies were secured from Division of Wildlife personnel.
Locations
were plotted on a map of Moffat County and data were summarized in table
form by type of manipulation, date(s) treated and number of acres treated.
A sample of 20 landowners in Moffat County were interviewed to estimate
percentages of native habitat altered to bring land under cultivation or to
improve pastures for grazing purposes.
Unsuccessful inquiries were made at BLM and DOW offices to Ipcate and summarize information on the distribution of sage grouse within the study area.
Available information on strutting grounds and brood count routes censused
and check stations manned by DOW personnel were gathered and summarized.
Records of all summer and winter period sage grouse observations made by
Project personnel were tabulated and work maps were prepared showing locations and numbers of birds.
Records of all road killed sage grouse and other mortalities were tabulated
by date, time, location, and cause of death if known. Wings from all
mortalities were dried and saved for future reference.
Sage grouse were counted on all known active strutting grounds in Moffat
County during the spring of 1976. Attempts were made to locate and count
as many old and new grounds as possible rather than attempting replicate
counts on fewer grounds.
A crew of three men, using three 4-wheel drive
vehicles, searched various areas of Moffat and Western Routt counties from
mid-April through May 1976.
Sage grouse broods were counted along ten trial routes in Moffat and western
Routt counties during August 1976. Sage grouse wings were collected through
the use of 16 wing barrel collection stations placed in Moffat and western
Routt counties.
Age and sex composition of the wing samples were determined
by Clait E. Braun and data were summarized.
Ground searches for wintering concentrations of sage grouse were made from
December 1976 through March 1977. Locations of sage grouse observed or
reported were .recorded by date and time, location, and vegetative type.
Locations of wintering concentrations were used to locate new strutting
grounds during the spring of 1977.

�-63Sage grouse population data gathered in management surveys were analyzed
in order to attempt to ascertain any trends in sage grouse populations.
RESULTS AND DISCUSSION
Vegetative

Descriptions

Additional references concerning specific vegetative studies within the
Moffat County area have not been located.
General references to descriptions
of vegetative types and methods of vegetative analyses which may be useful
in future studies are summarized as follows.
Harrington (1954) described the semi-desert vegetative zone in no~thwestern
Colorado of which the sagebrush plant community is an integral part. This
plant community supports dense stands of big sagebrush (Artemisia tridentata)
with varying mixtures of herbaceous plants in the understory.
He states
the type is best developed below 7,000 feet in middle western Colorado, but
may extend to more than 10,000 feet in a few localities.
At its lower
border, the type merges with the saltbush (Atriplex spp.) and greasewood
(Sarcobatus vermiculatus) communities, while at its upper border, the
boundary between sagebrush and mountain shrub is usually clear cut. Its
contact with pinon-juniper (Pinus edulis-Juniperus spp.) is frequently
diffusive.
Extensive methods of mapping range types including traverse and sketch and
random sketch from aerial photographs are described by Stoddart and Smith
(1943). Costello and Schwan (1946) discussed methods of judging conditions
and trends on ponderosa pine ranges in Colorado using the criteria of vegetative composition, vigor, and density, soils, and erosion.
Principles
outlined have been adapted and applied to other ranges.
Kuchler (1967) describes
tation studies including
niques.

in detail various methods used in intensive vegeuse of transects, quadrats, and step-point tech-

Personnel of the BSF &amp; W in Fort Collins are mapping vegetative types within
five test areas of the western United States as part of the Western Energy
and Land Use Team. Remote sensing, including infra~ed photography and
satellite photography (ERT) combined with computer mapping are being
utilized in this mapping.
While purchasing satellite photographs and computer print outs for an area as large as Moffat County is prohibitive in
cost, copies of their vegetative maps covering a portion of the study area
may be possi~le to secure through purchase or loan.
Distribution

of Big Sagebrush

Major areas of big sagebrush plant communities within Moffat County have
been delineated on work maps (scale 0.5-in-per-mi).
Extensive field
surveys involving traverse and sketch, random sketch from SCS aerial photographs taken in 1973, and interpretation of types shown on SCS range
resource work maps for Moffat County were utilized to map areas of big sagebrush and complementary range types. Areas of big sagebrush are in the
process of being transferred to a county map (scale 0.25-in-per-mi).
The
final map should be completed during the next segment.

�-64Alteration

of Big Sagebrush

Table 1 lists amounts of land in Moffat County by ownership as determined
from information secured from various Federal and State Agencies.
More than
one-half of the total land area of 3,035,530 acres (1,228,479 hectares) in
Moffat County is administered by Federal Agencies (51.3%), compared with
42.1 percent in private ownership, and 6.6 in State ownership.

Table 1.

Land ownership

Ownership

in Moffat County, Colorado

1976.

Acres

Percent of Total

u. S. Government
BLM

1,354,552

44.62

N.P.S.

149,444

4.92

U.S.F.S.

41,763

1.38

BSF&amp;W

12,380

0.41

Subtotal

1,558,139

51.33

State of Colorado
State Land Board
(as of 5/1/76)
DOW

191,525

6.31

9,120

0.30

Subtotal
Private Land

200,645
1,276,746

6.61
42.06

Subtotal

1,276,746

42.06

Total for Moffat County

3,035,530

100.0

Locations of all known range conversions on Federal lands in Moffat County
were secured from Division of Wildlife Researcher Roland C. Kufeld and recorded
on a map (Fig. 1). Table 2 summarizes range conversions on Federal lands by
type of habitat manipulation and acreages.
The largest amount of range conversions on Federal lands in Moffat County has been through spraying (39.2%),
followed by plowing and seeding (25.0%), chaining (19.5%), reseeding (14.6%),
burning and reseeding (1.2%), and rotobeating (0.5%). Acreages and percentages by major vegetative type have not as yet been determined.

�I ~...:

.... :.: ... f~·

\

I

.J~

, ~m'ff'%h~t}
t-ui

'\

II

iv-

1
'?r:'

7t

-.;cp

-

~-~~./
'~

E

_

~

1C

._

~

I
GENERAl.

ROAD

1'.1A P

MOFFAT COUNTY
COLORADO

M~ttnt Count,..

(V.l.p b7 ~1rGro

I:i,

Pl~&lt;)r)

to,.:
~

Location

12-01

Oder

(1)

--_ ..

seea

OnlJ'

(L) Ch4in or WI

(8) Spmy

(M) Plou

(C) Chun

(11) Burn (, Seed

(Ii) Rot-oheat

,~&lt;h;;'1'

\'08

.:.

ot Project

ot Project

&amp;

Soed

6: So4:I.

0"I.JI

I

�-66-

Table 2. Summary of habitat manipulation
Moffat County. 1/

projects on Federal lands in

Percent
of Total

Number
Acres

Map
Symbol

Type of
Manipulation

(A)

Seed (not
fire reseedings)

4,597

9.5

Fire reseedings

2,455

5.1

(A)

Subtotal

7,052

14.6

(B)

Spray

18,919

39.2

(C)

Chain

2,607

5.4

(C-M)

Chain, plow and seed

1,500

3.1

(L)

Chain or rail and seed

5,302

11.0

Subtotals

9,409

19.5

(H)

Rotobeat

240

0.5

(M)

Plow and seed

12,056

25.0

(N)

Burn and seed

570

1.2

48,246

100.0

Totals

1/ Includes

only projects on BLM and F.S. lands.

It was determined that approximately 3.1 percent of the Federal lands in
Moffat County has been altered intentionally through various habitat manipulations.
Except for one small area of 60 acres (24.3 hectares) on the Routt
National Forest; all intentional habitat manipulations on Federal lands in
Moffat County have been on BLM lands.
Numerous wild fires have resulted in range conversions over undetermined
additional areas of Federal, State, and private lands.
Landowner

Interviews

Twenty private landowners within prime sage grouse ranges in Moffat County
were interviewed to determine the past and future extent of habitat alterations on native sage grouse ranges on private lands. Table 3 summarizes
information gathered in this survey.

�Tllbt..

,.

L.ulbwner

SUJruIary of habitat
Type

of

manipulations-

c::_

OpeJ:Kion

cr•• aect

1_76

Sheep &amp;
Cattle
Ranching

1,969

Sheep

16,.000

.-76

&amp;

within

prime

sage

grouse

Number Acres Controlled
S--tat.
L••••
Sch •. Landa-.

Pvt.

920

-

3,OOa

-

C!attlw
Ranchln9

n,ooo

4-76

Cattl.
Ranching

980

5-76

Sheep &amp;
Cattle
Ranching

340

6-'1.

Landowner
Only

520

7_76'

Ca.t.tle a
SlI_p
Ranching

37,000

8-76

cattle
Ranching

1,.500

-

9-16

Cattle
Ranching

1:7,,500

150

Sheep, CattIe &amp; Dry
land Farmlng

No.' Ac..r.es

.lpprox.
Total

640

-

3,529

AJ.'tered

3,.O-cro

2'4,000

800

4,200

-

1,920

29,080

1,365

-

-

%

Burn,
&amp;

IS

Approx.

Acres
plow,

No.

325

Rail,

(Deeded)

seed

0.0

or

--

0

-

0.0

Disk or brush_
cutter
&amp; seed

9.0

100

Brushcutter

2.8

Brushcutter
seed

88.2

20

Plow

seed

5.9

-

--

0

-

0.0

Brushcutter,
cUltivate
&amp; seed

1.3

-

0.0

Unknown

Plow &amp; seed
spray

320
,Oeedec1 &amp;
Pvt. Lease)
300

-

-

-

520

2,000

2,500

12,,000

-

53,500

8t-000
(Deeded &amp;
Pvt. Lease)

Brushcutter,
cUltivate
&amp;. seed

20.5

500

1,280

-

2,940

90
(Deeded)

Disk or plow
(horses)
&amp; seed

6.0

0

59,.840'"

10,.160

90,.150

4,000
(Deeded)
3,400
(Deeded)

Aerial

22.9

Unknown

Aerial

33.7

160

Disc,
seed"

0

240

320

160
2,500

-

800

-

960

(Deeded)

-

1,.840

-

5,480

11.2

-

&amp;

-

340

&amp;

0

harrow,

seed

-

3",.540

plow,

0.1

Plow,

(Deeded")

-

2',560

~

Method

Planned

47.2

&amp;

seed

Lease)

46 rOOO

42,000

Method

(ht.)

(Deeded

Pvt.

3••
76

11-76

County.ill

BLM

Clitti.

Cattle
Ranching
&amp; Dryland"
Far.ing

Moffat
NPS-

Ranching

10-76

range,

Unknown

&amp;

sprayed

Plow &amp; seed

&amp;

spray

19.4

350
(Deeded")

&amp;

4,520
(Deeded &amp;
Pvt. Lease)

Spec. tiller,
seed or aerial
spray or burn

100.0

240
(Deeded)

Hand &amp; team&amp;

75.0

0

-

0.0

-

O.~

Disc, burn,
seed

plow

plow

&amp;

15.4

0.0

-...J
I

12-76

Cattle
Ranching
&amp; DryIand
Fuming

320

-

-

-

-

320

r3-76

lJt'yland"
Farming

720

3,040-

-

-

-

3,760

3,"560
(Oeoded &amp;
Pvt. Lease)

Brushcutter
t
plow, &amp; seed

94.7

0

Cattle
Ranching
Dryland
"arming

1,980

-

-

-

-

1,980

900
(Deeded)

Hand &amp; teams

45.5

200

Sheep &amp;
Cattle
Ranching

1,280

-

-

138,240

-

139,520

Z60
(Deeded)

Brushcutter
t
burn,. plow,. &amp;
seed

20.3

0

Cattle
Ranching
ory1and
Farming

1,.531

It 780

-

1,920

-

5,231

I,880
(Oaeded a.
Pvt. r.eeae)

Brushcutter
or
disk &amp; burn,.
plow &amp; seed"

56.8

80

17-76

Cattle
Ranching
Ie lJt'yland
Farming

320

3217

-

-

-

640

160
(Deeded)

Plow or brushcutter

25.0

0

-

0.0

18-76

Sheep
Ranching

12,.000

-

12,800

80,640

-

105,.440

21)0
(Deeded)

TeaJrl &amp; burn

1.7

0

-

0.0

19-76

Cattl.
Ranching

5,000

2,000

1,.280

9,000

-

17,.280

8.6

600

Brushcuttu
seed

Cattle
Ranching

160

5,120

9,410

23.8

70

DiSk

Total.

115,.120

14-76

Brushcutter
,.
plow, &amp; seed

10.1

&amp;

tS-76

r~76

&amp;

20-76

50

1:6,.380

!J Inforllation

from landowner

intarviwa

aecured

Brushcutter
seed

50
(Deeded)

Team} burn
plow

600

-

4,080

24,.160

362,.480

15,280

533,420

30,210
out of 119,980
(Unk.-out)

I2.6"

26.8JI

10.2"

17.6%

25.2%

131.500
Percentage.,
10.31'
of Total.
sampled

(peeded)

1n 1976-1977.

-

&amp;

&amp;

~

--

2,055
out of 119.980
(Unk
c-ou t )
s

-

I
0\

--

1.7%

-

0.0

Brushcutter
,.
plow &amp; seed"

&amp;

seed

&amp;

2..4

8.6

33.3

�-68-

Landowners interviewed owned and/or controlled grazing rights on lands
averaging 26,671 acres (10,794 hectares) (range 320 to 139,520 acres or 130
to 56,464 hectares).
Private lands in these 20 holdings amounted to 10.3
percent of the total deeded land within Moffat County.
It was determined that approximately 25.2 percent of the deeded land has been
altered in the past in order to bring land under cultivation or to improve
pastures for grazing purposes.
These individuals plan to alter an additional
1.7 percent of their land within the next ten year period, if economic conditions are favorable to do so.
Table 4 lists amounts and percentages of croplands on 16,535 acres (6,692
hectares) owned or leased by the twenty landowners interviewed within Moffat
County during 1976. By use, 79.4 percent of the croplands were used-to grow
dryland small grains (wheat or barley), 13.8 percent to grow drylandhay,
and 6.8 percent to grow irrigated hay.
Plans are to continue this survey of landowners within prime sage grouse
ranges and also compare the extent of habitat manipulation on lands of selected landowners within former prime sage grouse ranges where habitat and population are only remnants of former levels.
Included in the category of
former prime sage grouse ranges are the areas of extensive dryland farming
northeast of the town of Craig, in Moffat County, and northwest, southeast,
and southwest of the town of Hayden in western Routt County.
Distribution

of Sage Grouse

Complete records of all field observations of sage grouse secured during
both summer and winter periods are being maintained by date and time, specific locations, numbers of birds, and vegetative cover types.
Separate maps,
(scale 0.25-in~per-mi) showing summer and winter distribution of sage grouse
were prepared but plans are to reproduce final maps following one more year's
observations.
Collections
Wings from nine sage grouse collected from road kills and other mortalities
since January 1, 1976 have been dried and stored for future reference.
Cold
storage facilities for long term storage of whole or other parts of sage
grouse are not available in Moffat County.
A collection of sample wings of
different sexes and ages from hunter collections were mounted on a large
board for comparisons with collected materials.
Sage Grouse Populations
Strutting

Ground Counts

A minimum of 965 sage grouse of both sexes were counted on 27 active strutting
grounds in Moffat County during the spring of 1976 (Table 5). Seven-hundred
sixty-six males were observed on 24 of these grounds where sexes were determined for an average of 31.92 cocks per ground.
During this first spring,
attempts were made to locate as many old and new grounds as possible, rather

�-69Table 4.

Landowner
Code

Summary of cropland

use within Moffat County 1976.1/

Dryland
Hay

Acres of Croplands by Type
Dryland
Wheat or Barley

Irrigated
Hay

1-76

0

300

65

2-76

0

0

0

3-76

400

0

100

4-76

80

60

0

5-76

0

0

100

6-76

0

0

0

7-76

1,000

5,000

0

8-76

0

0

90

9-76

0

0

200

10-76

80

270

0

11-76

0

1,560

0

12-76

40

200

0

13-76

15

3,545

0

14-76

400

500

0

15-76

0

0

260

16-76

200

1,600

80

17-76

70

90

0

18-76

0

0

200

19-76

0

0

0

20-76

0

0

30

Totals

2,285

l3,125

1,125

Percentages

13.8

79.4

6.8

1/ Basis - a sample of 20 landowners

(same individuals

listed in Table 3).

�Table 5.

Summary of sage grouse strutting ground counts, Moffat County, spring 1976.

W.C.O.
District

Date

Counter(s)

Name of Ground

Time of
High Count

5/14/76
5/13/76
5/13/76
5/13/76

D. Hoffman and J. Corey
D. Hoffman and J. Corey
D. Hoffman and J. Corey
D. Hoffman and J. Corey

Karren Ranch
Escalante
Stateline
Haslim Cow Camp
Subtotals of high counts

6:15 AM
6:50 AM
5:40 AM
6:30 AM

R. Lowry

4/23/76

D. Hoffman

Powder Wash Hill
Subtotals of high counts

7:00 AM

C. Brown

4/15/76
4/22/76
4/16/76
4/21/76
4/20/76
5/9/76
5/9/76
4/13/76
4/13/76
4/20/76
4/22/76
4/14/76
4/24/76
4/15/76
4/24/76
4/15/76
5/6/76

D. Hoffman
D. Hoffman and J. Corey
D. Hoffman
T. Howard
D. Hoffman and T. Howard
T. Howard
T. Howard
D. Hoffman and J. Corey
D. Hoffman and J. Corey
D. Hoffman and T. Howard
D. Hoffman and J. Corey
D. Hoffman
D. Hoffman and'T. Howard
J. Corey
T. Howard
J. Corey
D. Hoffman

Spring Creek 111
Spring Creek 111
Cross Mountain #1
Big Gulch #1 (Beckett's)
N. Fork Big Gulch #2
N. Fork Big Gulch #2
Grassie Reservoir
Lay Creek
Bord Gulch
Bord Gulch
Greasewood #1
Upper 19 Road
Upper 19 Road
Bighole Gulch #1
Bighole Gulch #1
Lester Cox Ranch
Bighole Butte (new grounds)
Subtotals of high counts

6:05 AM
5:15 AM
5:30 AM
5:26 AM
5:25 AM
6:10 AM
6:53 AM
5:30 AM
6:50 AM
6:25 AM
6:10 AM
5:45 AM
8:00 AM
6:00 AM
5:27 AM
6:50 AM
7:45 AM

L. Vidakovich

Number of
Cocks

Total
Birds

26

30
45
45 to 50
65 to 70
185 to 195

--26
26

26

26
26

25
23
18
3
31
13
21
32
32
30
30
32
23
17
19
20
17
280

28
23
19
4
32
13
21
32
32
33
30
42
23
22
19
21
17
301

------------------------------------------------------------------------------------------------------------------------

I

"-.J
0
I

�Table 5.

Summary of sage grouse strutting ground counts, Moffat County, spring 1976 (cont.)

W.C.O.
District

Date

Counter(s)

Name of Ground

Time of
High Count

Number of
Cocks

Total
Birds

C. Woodward

5/5/76
5/7/76
5/5/76
5/7/76
5/4/76
4/22/76

D. Hoffman
D. Hoffman
D. Hoffman
D. Hoffman
D. Hoffman and J. Corey
T. Howard

Timberlake 111
Timberlake 111
Timberlake 112
Timberlake 112
Fan Rock
Dressler Gulch (new ground)
Subtotals of high counts

7:05 AM
6:50 AM
6:02 AM
6:40 AM
6:10 AM
6:28 AM

29
12
59
60
113
36
238

31
12
62
62
117
38
248

W. Roland

4/16/76
4/16/76
4/20/76
4/25/76
4/21/76
5/11/76
4/21/76
4/20/76

J. Corey
J. Corey
J. Corey
J. Corey
J. Corey
T. Howard
J. Corey
J. Corey

Juniper
Axial Basin
Axial Basin
Round Bottom
Drylake 112
Drylake 112
Deception Creek
Yellow Jacket (new ground)
Subtotals of high counts

6:40 AM
6:00 AM
6:00 AM
6:05 AM
6:00 AM
5:55 AM
7:00 AM
5:10 AM

24
25
28
55
57
19
14
18
196

24
28
31
57
58
19
14
21
205

Totals of High Counts

766

Summary:
965 to 975 total birds counted on 27 grounds (average 35.74 to 36.11 birds per ground)
766 cocks counted on 24 grounds (average 31.92 cocks per ground)

I
......•
t-'
I

965 to 975

�-72-

than make replicate counts on fewer grounds.
The 3 new grounds were located
and counted and six old grounds, which no longer were counted by Division of
Wildlife Regional personnel were relocated.
Table 6 lists the status of
all known sage grouse strutting grounds in Moffat County, and Fig. 2 shows
locations of these strutting grounds.
No effort was made to count strutting
grounds in western Routt County during 1976, but plans are to expand searches
to this area during 1977.
Brood Counts
Sage grouse were counted along ten routes (Fig. 2) in Moffat and western
Routt counties in August 1976 (Table 7). A total of 412.6 mi (663.9 km)
were driven on these routes and 564 birds were counted, for an avera~e of
1.37 birds per mi (.85/km). All routes were run in the early morning,
commencing as soon as it became light enough to see well.
Seventeen separate broods were counted along these routes before the juveniles became too large to distinguish from the adults.
These 17 broods
averaged 5.29 young per brood. A young-to-adult ratio of 2.37:1 indicated
that sage grouse in Moffat and western Routt counties experienced good
reproductive success in 1976. Seven additional sage grouse broods not
counted on established routes averaged 3.86 young per brood, bringing the
total number of broods tallied in 1976 to 24.
Harvest Data from Wing Collections
Sage grouse wings were obtained from the Moffat County area during 1976
through use of 16 wing barrel collection stations (Fig. 2) placed in this
area, wing envelopes returned to the Wildlife Management Section, and miscellaneous hunter contacts; number of wings collected were: Great Divide, 129;
Cedar Mountain, 127; Dinosaur Headquarters, 105; Timberlake, 90; Cold Springs
Mountain, 52; Deception Creek, 35; Moffat County Road 4, 31; Maybell City
Park, 24; Axial Basin, 19; California Park Road, 16; Juniper Springs, 13;
Wolf Creek, 7; Sand Wash, 7; Slater Creek, 6; Powder Wash, 0; Sand Wash
Ridge, 0, wing envelope collections, 44; and miscellaneous hunter contacts,
43. In all, 748 wings of sage grouse were collected during September 11-13,
1976. Age and sex composition of the wing samples are listed in Table 8.
Females comprised 84.1 percent of the adult segment of the harvest, 64.3
percent of the subadult or yearling segment of the harVest, and 57.4 percent
of the immature or juvenile segment of the population, suggesting differential mortality is occurring in the Moffat County area. These data further
suggest that segregation of sexes had started in the Moffat County area
prior to the hunting season.
Analyses of wing molt patterns of breeding age females revealed that 97 of
132 (73.5%) adults, and 49 of 74 (66.2%) sub adult hens were probably successful nesters in 1976. These data and the high percentage of immatures in the
harvest sample (63.7%) indicate that nesting success and production were good
in the Moffat County area in 1976. The percentage of subadults or yearlings
in the harvest (15.4%) indicates reproduction during 1975 was at least fair.
In 1976, 2.3 chicks per hen (3.3 per successful hen) were present in the
harvest.
No comparable data are available for the Moffat County area for
previous years.

�Table 6.

Status of sage grouse strutting grounds, Moffat County 1976.

Status

C. Brown

Active

N. Fork Big Gulch #2
Grassie Reservoir
Lay Creek
Bord Gulch
Greasewood #1
Upper 19 Road

Relocated

Spring Creek If1
Cross Mountain #1
Bigho1e Gulch #1
Lester Cox Ranch
Big Gulch #1 (Beckett's)

Wildlife Conservation Officer District
W. Roland
C. Woodward
R. Lowry

Juniper
Axial Basin
Dry Lake 112
Deception Creek
Round Bottom

Timberlake 112
Fan Rock

Powder Wash Hill
Gee Flats 11

L. Vidakovich

Karren Ranch
Escalante
Stateline
Has1im Cow Camp
Bear Creek ])

Timberlake If1
I

'-l
W
I

New

Bigho1e Butte

Yellow Jacket

Dressler Gulch

Old Grounds

Cross Mountain #2
Clarida Ranch
Fortification
Big Gulch 112
Big Gulch 113
Big Gulch #4

Dry Lake If1
High Mesa il

Timberlake #3

-11 Counted by R. Lowry.

11 Counted by L. Vidakovich.
-3/ Not as yet relocated.

il Report of 5 to 7 cocks received by W. Roland from Power Plant personnel.

Sand Wash Ridge Road
Hay Shed
Beaver Basin
Sand Wash Rim
Goodman Draw
Snake River West

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L

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t\1
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\
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, •. I: -7,' '" ..
'qC
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•••

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-~':~r.'&lt;

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II.

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I

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•

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•

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Rolocated
.ow

Brood count

, ' *
W1n&amp;ba

route

•••••
,"ll

••

••
"m.t&amp;"m

•
1

, ',
I

'---

_
~

....,I
.j::I

�Table 7.

Summary of sage grouse brood counts, Moffat and Routt counties 1976.

Sage Grouse Counted
Adults
Young
Unc1ass.
Cocks
Hens

Total

Number
Broods

Young/
Brood

Number
Brood1ess
Hens

Birds/
Mile

Birds/
Minute

31

3

5.33

2

1.10

0.30

0.49

0.14

1

8.00

1

0.46

0.12

1

0.24

0.06

County

Date

Name of
Route

Moffat

8/3

Juniper-Axial

A.M.

28.1

105

5

o

16

10

8/27

Juniper-Axial

A.M.

28.3

100

o

8

o

6

14

8/4

Co. Road 7 to 4

A.M.

28.4

110

2

3

8

13

o
o
o

4

o
o

o

12

12

9

17

Routt

Time

Miles

Minutes

1

4.00

0.39

0.10

28

2

4.50

1.04

0.23

1

16

1

9.00

0.59

0.15

30

52

3·

6.33

2.43

0.50

0.83

0.18

1.37

0.32

6

8/5

Timberlake

A.M.

24.7

100

2

8/24

Timberlake (Revised) A.M.

31.1

120

o

8/6

Spring Creek

A.M.

26.8

120

2

8/25

Spring Creek

A.M.

26.9

110

1

5

9

8/10

Blue Mountain

A.M.

21.4

105

3

19

8/11

Powder Wash

A.M.

24.0

110

1

o
o

19

20

41

41

11

236

250

o
o
76

8/19

Great Divide

A.M.

23.6

110

8/26

Great Divide
(Revised)

A.M.

22.7

130

o
o

o
o
o
o
o
o

Subtotals

331.2

1,510

19

16

Average (Moffat County)

25.48

116.15

30.1

115

1

28.8

110

o

22.5

110

2

Subtotals

81.4

3

Average (Routt County)

27.13

335
111.67

8/31 Powder Wash (Revised)

A.M.

30.0

130

o

8/14

A.M.

15.2

160

3

8/18

Cold Springs Mtn.

E1khead

A.M.

8/30

E1khead (Revised)

A.M.

8/20

Slater Park

A.M.

Totals
,
.Average (Moffat and Routt Counties)

412.6
25.79

1,845
115.31

22

o
o
o
o

16

4

1

16.45

1.56

8

8

0.34

0.07

28

28

1. 23

0.22

408

519

5.43

1. 57

0.34

4.00

0.37

0.10

0.35

0.09

5.00

1. 07

0.22

4.67

0.55

0.13

1. 37

0.31

6

11

3

14

1

o

10

10

12

24

2

14

28

45

3

90

436

3.66

5

10

564

17

5

5.29

....,I

~I

�-76-

Table 8. Numbers and percentages of sage grouse in the harvest sample
(N = 748) by age class and sex, 1976.

Sex

Number

Adult
Percent

Subadult
Number
Percent

Immature
Percent
Number

Males

25

3.3

41

5.5

203

27.1

Females

l32

17.6

74

9.9

273

36.6

Totals

157

20.9

115

15.4

476

.63.7

Approximate dates of hatching, determined by backdating, using stages of
primary molt, were estimated for all but one chick in the harvest sample
collected within the Moffat County area (N = 439). Wings collected by the
Wildlife Management Section, using wing envelopes, were not included in these
age data. Estimated hatching dates ranged from May 20 to July 15, with the
overall average peak of the hatch occurring from June 10 to 14. This overall
peak of hatch was similar to that determined for sage grouse ranges at intermediate elevations, approximately 6,500-7,500 ft (1,981-2,286 m), 'such as
the Great Divide-Timberlake
area. Peak of hatch was estimated to be approximately 10 days later than intermediate-elevation
ranges for Cold Springs
Mountain at elevations from 8,400 to 8,600 ft (2,560-2,621 m) and approximately 7-9 days earlier than intermediate-elevation
ranges for the Axial
Basin-Juniper-Deception
Creek area at elevations varying from 6,200 to 6,600
ft (1,890-2,012 m). Peak numbers of hens on strutting grounds located at
intermediate elevations, such as in the Great Divide area, probably occurred
during the week of April 11-17, 1976.
Winter Concentration

Counts

A minimum of 1,223 sage grouse were counted on 18 different wintering areas
during the period December 1976 through March 1977 (Table 9). An average
of 67.9 sage grouse were counted per wintering area during this period. At
least ten of these areas are considered to be major wintering areas within
the Moffat County area.
Sixty separate flock locations were recorded for these 18 wintering ranges,
for an average of 3.3 wintering locations per wintering range. An average of
20.4 birds were counted per wintering flock location.
At least one strutting ground has been located for each of the known 18
wintering ranges, indicating the wintering areas are important for reproduction as well as winter survival.
Many of the wintering areas are also
known to be important as sage grouse brood rearing ranges.
Although
Mountain

small wintering flocks of sage grouse were located on Cold Springs
and Blue Mountain during the exceptionally mild, open winter of

�Table 9.

Sage grouse wintering

areas, Moffat County, December 1976 to March 1977.

Minimum
Number
Birds

Minimum
Number
Flock
Locations

Wintering
Birds
Previous
Winter?

Birds
Stayed
Entire
Winter?

Strutting Ground(s)
In Area?

50
50
230
122
131
48

3
2
8
5
7
4

Yes
Yes
Yes
Yes
Yes
Yes

Yes
Yes
Yes
Yes
Yes
Yes

Yes
Yes
Yes
Yes
Yes
Yes

141
48
71
15
18
29
73
12
13
27

3
7
2
1
1
1
3
1
1
3

Unknown
Unknown
Unknown
Unknown
Unknown
Unknown
Unknown
Unknown
Unknown
Unknown

Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes

39

2

Unknown

Yes
Yes
Yes
Yes
No
Yes
Probably
Unknown
Unknown
Probably
(2 flocks)
Yes
(1 flock)

106

6

Unknown

1,223

60

Name of Area

Red Wash 1/
Deception-Creek 1/
Sand Wash Rim-Lo~kout Mountain
North Fork Big Gulch 1/
Thornburgh Gulch-Dry Gulch 1/
Jesse Gulch-Morgan Gulch l/Greasewood Draw-Moffat Count
Road 6 1/
Bigho1e Gulch 1/
Spring Creek II
Round Bottom
Fan Rock
Axial Basin
Four Mile Creek 1/
Moffat County Roads 18 and 29
Lay Creek
Cold Springs Mountain
Duffy Mountain
Blue Mountain-Moffat
Road 16

Total
Summary:

l! Location

1/

County

Number of known wintering areas = 18
Number of wintering areas considered to be major wintering
Average = 20.4 birds per wintering flock location
Average = 67.9 birds per wintering area
Average = 3.3 flock locations per wintering area

considered

to be major wintering

area.

Yes
(1 flock)

areas

10

Yes

Yes

I

"
"
I

�-78-

1976-77 flocks may not normally be able to overwinter in these higher
elevational ranges. Flocks, however, probably spend most winter periods
within the other 16 areas listed in Table 9.
Trends in Sage Grouse Populations
Brood counts of sage grouse within the Moffat County area have been conducted
sporadically for management purposes with very few exceptions during recent
years because their reliability has been questioned.
The one checking station
used to collect management harvest data on sage grouse in the Moffat County
area (Cedar Mountain) has not been manned for the past three hunting seasons
(1974-1976).
Any possible indicated trends in sage grouse populations during
recent years from management information must, therefore, be based upon
strutting ground count information, for the most part.
Numbers of male and total sage grouse counted within Moffat County by Wildlife Conservation Officers (Fig. 3) indicate populations declined steadily
during the period 1969 through 1975 increased markedly during 1976. Numbers
of strutting grounds counted by Wildlife Conservation Officers (Fig. 4) also
decreased steadily from 1969 through 1975 (except for 1972) and also increased
markedly during 1976. Trends in average numbers of male sage grouse and total
birds counted on strutting grounds by Wildlife Conservation Officers in Moffat
County from 1969 through 1976 (Fig. 5) tended to substantiate claims that
populations had indeed experienced gradual but steady declines from 1969
through 1975, and improved during 1976.
It is known that reproductive success of sage grouse within Moffat County was
good during the summer of 1976, based upon brood counts completed and hunter
harvested wing samples collected.
The strutting ground counts for 1976,
completed during the spring period, would not be influenced by any increase
in populations through good reproductive success in 1976. Also, most Wildlife Conservation Officers in the Moffat County area considered 1975 to be a
year of relatively poor reproduction for sage grouse, while hunter harvested
wing samples collected during 1976 indicated 1975 was a year of fair reproduction.
This investigation was started during January 1976 resulting in
renewed interest in counting sage grouse on known strutting grounds by WCO's
within the Moffat County area. This fact probably best accounts for increased
numbers of strutting grounds counted during 1976 in comparison with 1975 and
at least partially accounts for the increased numbers of males and total sage
grouse counted. Any real decline in sage grouse populations within the Moffat
County area, if this has been the case, has been accented because fewer and
fewer strutting grounds were counted annually.
The number of new and relocated strutting grounds found in this study indicates the overall number of
grounds censused annually could have been maintained with additional effort.

�2,500

o
III
E-t
~

o

u

2-,000

III

-, .

U1

g
t:t::
~

r..l
~

-..JI
\0

I

b.,

1,500

U1

r...

&lt;,

o
Ul
t:t::

r..l
III
:t
::&gt;

1,000

Z

'o-----o-

~ig. 3 • Trends in'numbers of
male and total sage grouse counted
on sage grouse strutting grounds by
W.C.O •.'s in Moffat County, 1969-1976.
Key::;-. - Ma:tes,

&lt;,

p

v

/
,

'-·if

Total Birds

lH;;""

o "0.&gt;t

~ ::3

"'+I
QlU1

~

500

+I
Cf)

~
~
,r...
~

1969

1970

1971

1972
YEAR

1973

1974

1975

1976

�30
Q
tr.1
E-4

8u

25

tIl
Q

§
o

0::
t!)
t!)

20

Z

H

E-4
E-4

I
00

~
tIl

rz..
o

o
I

rs

tIl

.•....•

C3

.&gt;t

m

~

::s
+'
tI)

~ 10

4-l

o

5

'"'

Fig. 4.Trends in numbers ot active sage grouse
strutting grounds counted by W.C.O.'s in Mottat
County, 1969-1976.

Itl

Q)

~

+'
III
;H
'"'
.rz..

9

1970

1971

1972
YEAR

1973

1974

1975

1976

�80

•
•\

.

70

\

~

§

\\.~.

60

8
rz:I
~
rz:I
~

.'-- .~.\
~

50

CI)

\

Ii&lt;.

o

&amp;1

i
i

.

»&gt;:

40

Fig. 5. Trends in average numbers of \
sage grouse counted on strutting ground~
/
by W.C.O. 's in Moffat County,I969-1976. \ ./'

/

........•.•

•

,

-,

-,

""'/

-, ./

•

Key:
30
Average No. Birds
Average No. Males

,

/

/

•

-g~
~
CI)
~

0

~
~
Q)
~
~U)
..-I~
Ii&lt;.

'-'

1969

1970

1971

1972
YEAR

1973

I
00

•.....

"&lt;,

•

~

&lt;

/'

. -.
.,

1974

1975

1976

I

�-82-

LITERATURE CITED

Costello, D. F., and H. E. Schwan. 1946. Conditions and trends on ponderosa
pine ranges. Rocky Mountain Forest and Range Exper. Sta. 33 p.
Harrington, H. D. 1954.
Denver. 666 p.
Kuchler, A. W.

1967.

Manual of the plants of Colorado.

Vegetation mapping.

Stoddart, L. A., and A. D. Smith.
Book Company. N. Y. 547 p.

Prepared by

1943.

The Ronald Press. N. Y.
Range management.

j)~'JYl)r:/?If"~
Donald M. Hoffman
Wildlife Researcher C

Sage Books.

475 p.

McGraw-Hill

�April

-83-

1977

JOB PRCG~lESS REPORT

State of

COLORADO
------~==~~~--------

Project

No.

'\.J-37-R-30

Plan No.

9

Work

Job Title
Period

Job No.

5

-------------------------------Population Dynamics and

Habitat Relationships of Blue Grouse
----~~==~==~~~~~~~~~---------------

Covered:

Personnel:

Game Bird Survey

April

1, 1976 to March

31, 1977

D. Benson, C. Braun, J. Claassen, R. C'Li.ppLnge r , J. Corey, D. Covic,
C. Crawford, S. Erickson, H. Funk, J. Gerrans, K. Giesen, D. Gore,
P. Gorenzel, J. Hobbs, D. Hoart, D. Hoffman, R. Hoffman, J. Jackson,
J. Kitzmiller, W. Larrick, D. Luce, F. Marcoux, T. Marinelli,
B. McCloskey, S. McEllin, R. Oakleaf, S. Palm, S. Porter, B.
Sigler, M. Smith, R. Stark, S. Steinert, J. Wagner and W. Woodard.
ABSTRACT

Investigations concerning the effects of hunting on blue grouse (Dendragapus
obscurus) populations, stability of breeding population levels, and habitat
relationships of blue grouse were initiated in 1975.
Studies were conducted
in northwestern Colorado at two locations,Green
Mountain-Blue
Ridge and Eby
Creek.
Acoustical census and systematic search were used to locate birds.
Playing of the hen cackle call greatly enhanced the observers efficiency in
locating territorial males, but only limited success was obtained in using
chick distress calls.
Most attempts to capture grouse, regardless of the
method employed, were unsuccessful.
Capture success (birds captured/birds
observed) amounted to slightly less than 5 percent.
Based on ecological
densities, Eby Creek (20 birds/km2) sup~orted higher 'densities of blue
grouse than Green Mountain (14 birds/km).
The population on Green Mountain
remained stable between 1975 and 1976.
Overall nesting success (62%) was
good and was similar to nesting success in 1975.
However, differences were
noted in the percent of subadult females that were successful nesters in
1975 (27%) and 1976 (61%). Peak of hatch occurred about two weeks earlier
in 1976 (16 to 30 June) than in 1975 (26 June to 10 July).
Even though a
40 to 50 percent reduction in brood size occurred production still contributed to nearly a doubling of the spring breeding population by late September.
Data from wings (N = 341) collected in Middle Park revealed that
the harvest was comprised of 39 percent adults, 12 percent subadults, 49
percent juveniles.
Volunteer wing collection stations accounted for 86
percent of the wings obtained in }1iddle Park.
About 14 percent of the fall
population on Green Mountain was harvested.
The total harvest during the
experimental season in Middle Park was 270 grouse, or about 1 percent of
the estimated total population.

�-84-

RECOMMENDATIONS

1.

Trapping efforts should be decreased with more effort being made to
collect population and harvest data.

2.

Another experimental season should be held in 1977, at least in Middle
Park.
The blue grouse season should extend through the big game
season with a bag and possession limit of 3 and 6 birds.

�-85-

POPULATION DYNAMICS AND
HABITAT RELATIONSHIPS OF BLUE GROUSE
Richard W. Hoffman

Sound management of game birds requires considerable knowledge concerning
the biology of each species and factors affecting their populations.
Blue
grouse are the most widespread member of the upland game birds in Colorado.
They occur in varying densities over more than 51,800 km2 (20,000 mi2) of
diverse habitat and terrain in Colorado (Rogers 1968). Breeding populations
have been found in essentially every habitat type from 1,830 m (6,000 ft)
to over 3,050 m (10,000 ft) in elevation.
Furthermore, blue grouse are
second and possibly first among the resident upland game birds in terms
of numbers harvested (18,000 average annual harvest).
Yet, little is known
about the population dynamics, habitat requirements, or the effects of
hunting on populations of blue grouse in Colorado.
Colorado has had a long history of conservative seasons on blue grouse.
This conservatism has mainly resulted from lack of sufficient data upon
which to base management recommendations and the misconception that hunting
can be a major mortality factor on blue grouse.
Based on intensive studies
conducted in other western states and provinces (Mussehl 1960, Zwickel 1958,
Harju 1974, Bendell and Elliott 1967, and Boag 1966) the blue grouse season
in Colorado could be liberalized without harm to breeding populations in
subsequent years.
Thus, through intensive population studies, emphasis
should be placed on collecting basic data on population dynamics and habitat
requirements of blue grouse in Colorado in order to understand the relationship between harvest and population size.

P.N.O. OBJECTIVE
Major objectives of this study are to (1) increase the harvest of blue
grouse in Colorado (double present harvest estimates) without harm to breeding
populations in subsequent years, (2) to identify differences in breeding
densities due to differing habitats, and (3) to document the stability of
breeding densities over time.
SEGMENT OBJECTIVES
1.

Review literature concerning (a) techniques for marking, capturing, and
censusing grouse, (b) methods of aging and sexing blue grouse, (c) population dynamics, behavior, and habitat requirements of blue grouse, and
(d) effects of hunting on grouse populations.

2.

Trap and individually mark 50 birds (including adults, subadults,
juveniles) on each of the two selected study areas.

3.

Estimate breeding densities on the study areas through the use of
acoustical census and systematic search.

and

�-86-

4.

Estimate

nesting

success and production

5.

Vegetatively describe areas studied and correlate blue grouse densities
with features of the environment.

6.

Manipulate

7.

Compile data, analyze results, and prepare progress

hunting

on the study areas.

season and obtain harvest data.
reports.

METHODS AND MATERIALS
Reference
materials
discussed

is made of Hoffman (1976) for a detailed discussion of methods and
used in this study. Any deviations or additions to these are
in the Results and Discussion section.

DESCRIPTION

OF STUDY AREAS

Intensive studies were conducted on the Green Mountain-Blue Ridge (GM-BR)
area in 1975 and 1976 (Fig. 1). The geographic, geologic, vegetational and
climatic features of the area have been described in a previous report
(Hoffman 1976). Detailed studies on the Eby Creek area were not initiated
until the spring of 1976. The following discussion relates to this area.
The Eby Creek area is approximately 10 km (6 mi) north of Eagle, Colorado in
Eagle County (Fig. 2). Area investigated is in T35, R84W, parts of Sections
31 and 32; and T4S, R84W, parts of Sections 4, 5, and 6 (Fig. 2). The study
area encompasses about 5 km2 (2 mi2). While a small portion of the area is
under jurisdiction of the Bureau of Land Management, U. S. Department of
the Interior, over 95 percent of the area is under private ownership.
The study area lies at the head of a small U-shaped valley that is bounded
on the southwest by Greenhorn Mountain and on the northeast by Castle Peak.
Vegetation of the study area can best be described as a patchwork of aspen
(Populus tremuloides) stands varying in size from less than 2 ha (5 A) to
over 100 ha (250 A) and separated by open sagebrush (Artemisia spp.)
dominated meadows.
Portions of the sagebrush meadows have been treated with
herbicide.
Such areas have been invaded by bunchgrasses and a variety of
perennial
forbs. Dense stands of serviceberry (Amelanchior alnifolia),
snowberry (Symphoricarpos occidentalis), chokecherry (Prunus virginiana),
and currant (Ribes spp.) commonly occur along the edges of the aspen or
bordering streams.
Mixed conifer-aspen stands predominate at the higher
elevations, whil~ the lower elevations support principally a sagebrushjuniper (Juniperus spp.) type. Coniferous types are almost totally absent
on the study area.
Topography of the Eby Creek area is characterized by a relatively flat valley
floor that rises abruptly on the southwest and northwest sides. Cliffs and
ledges are common along the valley rim. Above the rims the terrain quickly
changes into a gently rolling upland.
When viewed from the air, the area
appears to be a large plateau that has been bisected by a U-shaped valley.
Maximum relief is about 400 m (1,300 ft); elevations range from 2,500 m
(8,200 ft) to 2,900 m (9,500 ft) above sea level. Geology of the Wolcott

�\

~,

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�-89-

region which includes most of the study area has been described by Stauffer
(1953). Major drainage is to the south via Eby Creek •. Numerous smaller
streams occur throughout the study area and empty into Eby Creek, which
flows into the Eagle River about 10 km (6 mi) south of the study area. Most
of the streams maintain a year round water flow.
U. S. Weather Bureau precipitation and temperature data were available from
Green Mountain Dam (elevation 2,360 m) approximately 1 km southwest of the
GM study area. Weather records for the Eby Creek area were from the Eagle
County Airport (elevation 1,980 m) about 12 km southwest of the study area.
These data are summarized in Table 1. No wind measurements were available,
but prevailing winds are primarily westerly with frequent high velocities
in winter and spring.
The Eby Creek area supports slightly higher temperatures but lower annual precipitation than the Green Mountain area. Climate
of both areas is typically continental with seasonal and sometimes daily
variations in temperature, precipitation, and wind velocities.
RESULTS AND DISCUSSION
Analysis

of Capture Techniques

Various methods for capturing blue grouse have been developed and tested for
use in Colorado (Hoffman 1976). As in 1975, efforts were continued in 1976
to capture blue grouse. Unfortunately, most attempts to capture grouse,
regardless of method employed, were unsuccessful.
The major problem in
trapping grouse was the inability to approach birds close enough to catch
before they flushed.
In over 80 percent of the encounters with grouse, the
birds flushed or else the situation did not permit any capture attempts to
be made.
Of 451 birds observed in 1976, only 76 (17%) capture attempts were
possible; 22 (29%) were successful and 54 (71%) were unsuccessful.
Based on
the total birds observed (451), capture success amounted
to 5 percent.
Capture success was slightly higher in 1976 (5%) than in 1975 (2%).
All birds were captured with either a noose pole (17 birds) or a hoop net
(5 birds).
Nine chicks « 2 wks old) were caught by hand, but these birds
were too small to carry a band. Older chicks were captured with the hoop
net. However, the chicks were very difficult to find after the brood was
flushed.
No mortalities occurred as a result of trapping operations.
It is
believed that with present manpower (2 people) and time available for trapping,
an adequate sample (50+ birds/study area) of blue grouse cannot be captured
using the existing techniques.
Analysis
Breeding

of Census Techniques

Season

Blue grouse were censused during the breeding and brood periods through the
use of acoustical census and by systematic search (Hoffman 1976). Of 109
males located during the 1976 breeding season, only 37 (34%) initially responded to the tape recorded hen cackle call. Wing flutters were the dominant
response accounting for 28 of the 37 (77%) responses.
In most cases, when a
male responded by wing fluttering this induced a chain reaction of responses

�-90-

from nearby territorial males. Thus, 65 males (60%) were located either by
initially responding to the call (37 males) or as the result of a chain
reaction response (28). The remaining 44 males were found by systematic
search with the aid of a dog.
Table 1. Temperature and precipitation, Eagle and Green Mountain, 10 year
averages from 1966 to 1976. 1/

Temperature

Eagle
Precipitation
(cm)

Green Mountain
Precipitation
Temperature
(OC)
(cm)

Month

(OC)

January

7.2

2.4

7.1

2.9

February

- 4.0

1.2

- 6.7

2.2

March

- 1.5

1.9

- 1.9

3.8

April

5.2

2.1

3.3

3.6

May

10.9

1.1

9.3

3.2

June

15.2

2.7

l3.3

4.4

July

19.2

3.6

16.8

3.8

August

l3.9

1.8

16.2

3.3

September

12.9

2.6

11.7

3.2

October

3.7

3.2

2.6

3.8

November

- 1.0

1.2

- 1.3

2.9

December

- 8.0

2.7

- 7.0

3.0

4.9

26.5

4.1

40.1

Mean Annual

1/
- Data from: u.. S. Department of Commerce, Weather Bureau.
Data, Colorado Annual Summaries.

Climatological

�-91-

Systematic search of the habitat with the aid of a dog was the most successful method for finding hens during the breeding season, but some hens did
respond to the cackle call. Due to their more secretive habits and lower
response rate, only 43 hens were located; 4 (9%) responded to the call and
39 (91%) were found by systematic search. As indicated by the distorted
sex ratio (2.5 males per female), males were considerably easier to find
during the breeding season than females.
A higher percentage of males (77%) and females (36%) initially responded to
the call in 1975 compared to 1976. The difference was attributed more so to
the observer's increased ability to find birds in 1976 without the call
rather than an actual change in response rates between years.
Tests of the
reliability of the hen cackle call in censusing breeding blue grouse could
not be effectively conducted since the exact number of females and territorial
males was not completely determined for either study area. However, playing
of the call greatly enhanced the observer's efficiency in locating blue
grouse, especially territorial males.
All known territorial males responded
to the call at least once during the breeding season.
Limited success was obtained in using chick distress calls as only 8 percent
(4) of the 50 successful hens observed in 1976 responded to the call. This
compares to a 25 percent response rate in 1975. All documented responses
were by hens with chicks less than 4 weeks old. Most broods were found by
systematic search with the dog. Playing the call after the brood was
disturbed elicited a response from 74 percent of the hens.
Thus, the call
was useful in relocating successful hens after the brood was flushed.
Occasionally, males would respond, but those that did so appeared to be
attracted by the calling of the hen to her chicks.
Breeding Densities
Censuses of breeding birds were initiated on both study areas during the first
week of April in 1975 and 1976. Early arrivals were primarily males with
numbers of females increasing towards mid- to late April.
Activities
associated with breeding and density of breeding grouse peaked in early to
mid-May.
The total number of territorial males recorded during the peak of breeding
events was used as an index to the population size. An accurate estimate of
the number of breeding hens and non-territorial yearling birds was not
obtained because of their secretive habits, extensive movements, and poor
response rate to the recorded calls.
Since data collected in other studies
suggest there is a 1:1 sex ratio among breeding birds (Zwickel 1972, Bendell
et al. 1972), the breeding population was estimated by doubling the count
of territorial males.
Breeding densities in 1976 were slightly higher on Green Mountain (6 males/
km2, 16 males/mi2) than at Eby Creek (5 males/km2, 13 males/mi2).
However,
based on ecological densities, Eby Creek (50% suitable habitat) supported
a density of 10 males/km2 (26 males/mi2), whereas the ecological density

�-92-

of territorial males on Green Mountain (90% suitable habitat) was 7 males/
The population on Green Mountain remained stable
km2 (18 males/mi2).
between 1975 and 1976 (no change).
A similar comparison for Eby Creek was
not possible because only one year of intensive work has been conducted on
this area.
It was impossible to traverse the Eby Creek area to a sufficiently intensive
degree in one day in order to estimate the total number of females on the
breeding area. Two such attempts were conducted on the Green Mountain
area in May 1975 and 1976. The resulting ecological densities as determined
from the direct counts were 6 females/km2 (16 females/mi2) in 1975 and 8
females/km2 (19 females/mi2) in 1976. While the accuracy of these counts
are not absolute, they do reflect an essentially 1:1 sex ratio in the
breeding population at Green Mountain.
Assuming there is a 1:1 sex ratio,
the estimated ecological densities of the spring breeding population at
Green Mountain and Eby Creek were 14 and 20 blue grouse per km2, respectively
(36 and 52 birds/mi2).
Nesting

Success and Hatching Dates

Since successful and unsuccessful hens do not have the same probability of
being observed, it was difficult to estimate nesting success from field
observations.
Such estimates were considered over-inflated, but to what
extent is uncertain.
Analyses of wing molts from hunter harvested birds was
used as the best estimate of nesting success; however, an inadequate sample
of hunter harvested birds were examined from the Eby Creek area to reliably
estimate nesting success.
Due to the lack of information for the Eby Creek
area, the following discussion relates primarily to data collected in Middle
Park. Estimated nesting success for both areas is presented in Tables 2
and 3.

Table 2.

Estimated

nesting success based on field observations.!/

Study Area

Green Mountain
Eby Creek

!/ Estimates of nesting
be over-inflated.
'.!:.../ No data.

Nesting Success
1975

(Percent)
1976

78

78

m/;-/

83

success based on field observations

are believed

to

�-93-

Table 3. Estimated nesting
hunter harvested birds.

success based on analyses

Nesting

Success

of wing molts

(Percent)

1975

1976

Green Mountain

64

62

Eby Creek

~/

3~/

Study Area

from

1/ No data.
2/ An inadequate sample (15 total birds) of hunter harvested
examined to reliably estimate nesting success.

birds was

Examination of wing molt patterns of 41 adult and 15 subadult hens harvested
in Middle Park revealed that 78 percent of the adults and only 27 percent
of the subadults were successful nesters in 1975. Comparative figures for
1976 show 63 and 61 percent nesting success for adult (N = 60) and subadult
(N = 28) hens, respectively.
Overall nesting success was good in both years
averaging 64 percent in 1975 and 62 percent in 1976.
From data collected in another study (Bendell and Elliott 1967), the poor
nesting success of subadults can be attributed to their lack of experience
rather than their failure to mate.
Reasons for the large variation in
nesting success of subadults between years is uncertain, but may be related
to the earlier onset of nesting events and longer nesting season in 1976,
conditions that might have been favorable for better nesting success of
subadults.
Hatching dates for 1975 ranged from 21 June to 24 July with 79
percent of the hatch occurring between 26 June and 10 July.
Similar estimates for 1976 show a wider range of hatching dates (1 June to 22 July) and
earlier peak as 68 percent of the hatched occurred during the period 16 to
30 June.
Climatic conditions at Green Mountain were slightly warmer and wetter in 1976
than in 1975. However, a heavy snowfall of 56 cm (22 in) in late April of
1975 delayed the onset of nesting events. As a result of the warmer spring
temperatures in 1976 accompanied by less snowfall (16 cm, 6 in), most of
the snow had melted approximately
2 weeks earlier in 1976 than in 1975.
This apparently allowed the birds to start nesting earlier in 1976. Warm,
dry conditions prevailed throughout June of both years, thus, the weather
was favorable for good nesting success in 1975 and 1976.
It was anticipated that because of the warmer, drier conditions and earlier
snow melt at Eby Creek, the hatch would be earlier than in Middle Park. However, this was not the case. Hatching dates in 1976 as determined from 32
chicks harvested in the Eby Creek area ranged from 1 June to 24 July. The
peak of hatch occurred during the same period as estimated for Middle Park
(16 to 30 June).

�-94-

Production
From late June until late August 1976, observations were made on 32 distinct
broods, including 19 at Eby Creek and 13 at GM-BR. During the same period
in 1975, 11 broods were observed, all in the GM-BR area. The increase in
broods observed from 1975 to 1976 was related to improved observer efficiency
and larger area searched.
Average brood size, range of brood sizes, and
number of broods observed are given by months in Table 4.

Table 4. Average brood size, range, and number of broods observed
intervals, 1975 and 1976. 1../

Month

Green Mountain
1975
1976

by monthly

2/
Eby Creek:=1976

June
Mean

~/

5.3

5.0

Range

ND

3-7

4-6

Sample Size

ND

3

2

Mean

5.2

4.1

3.7

Range

2-7

2-6

1-7

Sample Size

4

7

9

Mean

3.0

3.0

2.2

Range

1-5

Sample Size

6

July

August

1-4
1

5

1../ Only distinct broods with full counts of brood size are included.
2/ Studies at Eby Creek were not initiated

until 1976.

1/ No data, most nests did not hatch until

the last week of June through the

first week of July.

�-95-

Average brood sizes recorded for the GM-BR area were 3.9 and 4.4 chicks per
brood in 1975 and 1976, respectively.
Brood size at Eby Creek averaged 3.4
chicks in 1976. The apparently lower average brood size at Eby Creek may
be attributed to a larger sample of broods counted in late July-early August,
whereas, primarily early to mid-July counts were made at GM-BR.
Rogers (1968)
found average brood sizes of 4.1, 2.6, and 3.9 in 3 years of study in western
Colorado.
A 4-year average of 4.2 chicks per brood was reported for blue
grouse in southeastern Wyoming (Harju 1974).
The sex ratio of chicks was determined from hunter harvested birds for which
sex was ascertained.
Based on the examination of 245 juvenile wings in
1975 (79) and 1976 (166) the sex ratio did not deviate significantly from
1:1. However, in both years, the sex ratio showed a slight imbalance in
favor of females, averaging 1 male: 1. 2 females.
Average brood size in late August or early September may be misleading as
field observations indicated some brood breakup and shuffling had begun by
this time. Broods observed after the second week in August are not included
in Table 4. While harvest may not accurately reflect popUlation composition
due to hunter and bird behavior patterns, it is believed that the young to
successful female ratio in the harvest did give a good measure of brood size
and survival of chicks through September.
The sample of hunter harvested
birds from Eby Creek was too small to estimate brood size. However, adequate
samples were obtained from Middle Park to permit analysis.
Average brood size for September was calculated to be 2.6 chicks in 1975 and
3.0 chicks in 1976. These estimates are based on the exmination of 128 hunter
harvested birds (36 successful females, 92 juveniles) in 1975 and 221 hunter
harvested birds (55 successful females, 166 juveniles) in 1976. Thus,
average brood size from the time of hatch until late September-early October
decreased by 2.6 chicks (50% reduction) in 1975 and 2.0 chicks (40 %) in 1976.
Fall Densities
Estimated fall densities for the Green Mountain area, 1975 and 1976, are
presented in Table 5. The calculations are based on the following assumptions:
(1) the estimated percent nesting success was correct, (2) immigration equalled
emigation, (3) a 1:1 sex ratio existed in the breeding population, and (4)
the estimated average brood size by September 30 was accurate.
Mortality of
the breeding population from April 1 to September is not figured into the
calculations, but is probably minimal «10%)
(Bendell and Elliott 1967).
Obviously none of the assumptions, nor the accuracy of the data, are absolutely correct, but it is believed that Table 5 does reflect the general
situation on the Green Mountain study area.
Even though nearly a 50 percent reduction in brood size occurred over a 3
month span, production still contributed to nearly a doubling of the spring
breeding population at the time of the fall hunting season.
For the two
year period, production was remarkably uniform between years, indicating
that regardless of size or age of the breeding population, nesting success
or clutch size, survival of chicks to late September is stable.
Considering
there is only a 25 to 30 percent turnover in the breeding population,
production was more than adequate to replace natural losses.

�-96-

Table 5. Fall densities of blue grouse on the Green Mountain-Blue
study area, 1975 and 1976.

Ridge

Year

Total
Breeding
Population

Nesting
Success
(Percent)

Average
Brood
Size by
Sept. 30

Total
Production
by
Sept. 30

Total
Population
by
Sept. 30

Birds
per
km2

Percent
Gain

1975

32

64

2.6

27

59

23

46

1976

32

62

3.0

30

62

24

48

Harvest
Hunting

Season

The 1975 blue grouse season opened statewide at sunrise on September 13 and
closed at sunset on October 5. In 1976 the regular season opened on September
11 and closed on October 10, except in Unit 28 (Middle Park) where the season
was extended to October 26 in conjunction with the regular elk season (October
16 to 26). Daily bag limit in both years was 3 and the possession limit was 6.
Volunteer

Wing Collection

Stations

Ten volunteer wing collection stations were constructed and tested for use
in Middle Park in 1975. The volunteer stations were instrumental in inexpensively increasing samples of wings collected (Hoffman and Braun 1975,
Hoffman 1976); thus, their use was expanded in 1976. Thirteen stations were
set up in Middle Park, 2 in the Poudre Canyon, 2 in the Eby Creek area, 1 in
North Park, and 16 in Moffat and Routt counties.
Stations in North Park,
Moffat, and Routt counties were primarily for obtaini~g wings of hunter
harvested sage grouse, but some blue grouse wings were collected.
The stations remained available to hunters throughout the blue grouse season
in Middle Park. Due to low hunter pressure, stations were not operated the
entire season in the Poudre Canyon and Eby Creek areas.
Stations in North
Park and Moffat and Routt counties were operated during the 9 and 3 day sage
grouse seasons, respectively.
Each station was checked at least twice a
week, the wings were removed and tagged according to the date and location
of the station.
A summary of the use of volunteer wing collection stations
is presented in Table 6.
Check Stations
In 1976, check stations were operated in the Middle Park, North Park, Poudre
Canyon and Eagle areas. During the opening weekend and second Saturday,
stations in Middle Park, North Park, and Eagle were open from about 1000 to

�-971800 MDT, depending upon traffic load. Due to low hunter pressure, the
Poudre Canyon station was only operated opening day. After the opening weekend, hunters were contacted opportunistically in conjunction with the checking
of the volunteer stations.
Data obtained per hunting party included: county
of origin, number of hunters, hours hunted (total all hunters), location,
birds observed, birds bagged, and birds lost. Whenever possible, the crop,
gizzard, ovary (if a female), and one wing were obtained from each bird
checked.
Sex by gonadal inspection was ascertained when possible.
Table 6.

Volunteer wing collection

station summary,

1975 and 1976.
1/

No. of
Stations

No. Days
Available

No. Wings
Collected

Percent ofTotal Wings
Collected

Middle Park

10

23

120

65.6

Middle Park

13

14

292

85.6

Eagle

2

16

19

28.4

North Park

1

9

0

0

2

16

29

80.6

16

3

33

41.8

44

81

493

68.6

Year

Area

1975
1976

2/

Poudre Canyon-

3/

Moffat and Routt-

Total

1/ Percent of total wings collected
l/ Only one station was available
1/ Of the 16 stations,

from manned

stations and volunteer

stations.

to hunters after 9pening weekend.

only 3 were set up near potential

blue grouse hunting

areas.

Harvest data obtained from check station operations are summarized in Table
7. It is concluded from these data that hunting pressure is low and has no
adverse effects on the population.
Only 8 birds were reported killed on the
Green Mountain area. Based on the estimated fall population of 59 birds,
less than 14 percent of the fall population on Green Mountain was harvested
in 1976. Data from other western states suggest that 25 to 30 percent of
the fall population can be safely harvested with no adverse effects on
subsequent spring breeding population (Zwickel 1958, Bendell and Elliott

1967).

�Table 7.

Blue grouse hunter checks, 1975 and 1976.

Hours
Hunted

Birds
Observed

Birds
Bagged

Birds
Lost

1/
HunterSuccess

2/
HunterEfficiency
(Percent)

Hours Hunted
per Bird

Crippling
Loss
(Percent)

Year

Area

Hunters
Checked

1975

Middle Park

112

578

158

45

2

.40

28.4

12.8

4.4

Eagle

57

227

83

22

1

.38

26.5

10.3

4.5

North Park

28

219

55

21

0

.75

38.2

10.4

0.0

Poudre Canyon

33

113.5

15

10

0

.30

66.7

11.3

0.0

Middle Park

138

610.5

141

61

5

.44

43.3

10.0

8.2

1976

I
\0

Eagle

73

494.5

159

76

3

1.0

47.8

6.5

3.9

North Park

49

422

171

54

0

1.1

31.6

7.8

0.0

Poudre Canyon

11

39

10

3

0

.27

30.0

13.0

0.0

1/ Birds harvested per hunter.
~/ Birds bagged/birds observed x 100.

ex&gt;
I

�-99-

Blue grouse hunting was mainly limited to range adjacent to roads or other
easily accessible locations, with many areas receiving virtually no hunting
pressure.
After opening weekend, hunting pressure decreased markedly.
Opening weekend accounted for over 60 percent of the wings collected and 75
percent of the hunters checked.
Of all the hunters checked in 1976 (271),
59 percent were unsuccessful, 21 percent harvested 1 bird, 10 percent harvested 2 birds, and 10 percent harvested the daily bag limit of 3 birds.
Wing Analyses
In 1976 wings were received from 542 blue grouse (373 from volunteer stations,
113 from check stations, 52 from the mail wing survey, and 4 from field
checks).
Five hundred and thirty six wings were classified to age and sex.
Age and sex composition of the harvest sample in 1975 and 1976 are presented
in Table 8. Small samples from all areas except Middle Park prevent comparisons among areas. The following discussion relates primarily to data
collected in Middle Park, but the general situation is believed to be applicable to the other areas.
Assuming subadults molt similarly to adults, approximately 73 percent of the
adult and subadult males and 22 percent of the adult and subadult females
had completed their molt by the end of the 1976 hunting season.
Thus, a
large portion of the subadults, especially males, could not be distinguished
from adults.
Males begin their primary molt before females so more subadult
males than females are indistinguishable
from adults in the fall. Similar
percentages estimated in 1975 for adult and subadult males and females were
40 and 6 percent, respectively.
The higher percentage of birds completing
their molt by late September in 1976 reflects the earlier nesting and peak
of hatch; consequently, the birds initiated their primary molt earlier in
1976 than in 1975. Since some subadults were assigned to the adult category
both age classes are grouped together as adults.
The 1976 data for Middle Park are slightly dissimilar from data collected in
1975 in that chicks comprised 49 percent of the harvest in 1976 (46 percent in
1975), and the sex ratio of adults (includes subadults) in the harvest was
nearly even in 1976 (1 male:1.l females); whereas, in 1975 it was distorted
(1 ma1e:l.6 females).
Adults and juveniles made up almost equal portions
of the harvest in 1975 and 1976, indicating production was good in both years
and possibly slightly better in 1976 than in 1975. The deficiency of cocks
in the adult harvest in 1975 was because their migration to higher, inaccessible areas, less frequented by hunters, occurred prior to the hunting season.
The females and chicks still remained on the lower areas where the bulk of
the hunting pressure occurred.
Conversely, with the earlier hatch in 1976,
the females and chicks had also initiated their upward migration by the
opening of the fall hunting season so all the birds were using essentially
the same habitat.
Hunters therefore had a better chance of encountering
and harvesting males.

�Table 8.

Year

1975

Age and sex composition of harvest sample, 1975 and 1976.

Adult
Female

Total
Adults

Subadu1t
Male

Subadu1t
Female

Tota11/ Juvenile
Subadu1ts
Male

Juvenile
Female

Total
Juveniles

Total
Birds

Middle Park

30 (42)l/ 41 (58)

71 (42)

5 (25)

15 (75)

20 .(12)

36 (46)

43 (54)

79 (46)

170

Eagle

6 (43)

8 (57)

14 (67)

0 (0)

0 ( 0)

o ( 0)

4 (57)

3 (43)

7 (33)

21

North Park

11 (38)

18 (62)

29 (57)

5 (62)

3 (38)

8 (16)

5 (36)

9 (64)

14 (27)

51

Poudre Canyon

0 (0)

1(100)

1 (14)

1 (25)

3 (75)

4 (57)

o ( 0)

2 (100)

2 (29)

7

47 (41)

68 (59) 115 (46)

11 (34)

21 (66)

32 (13)

45 (44)

57 (56)

102 (41)

249

68 (51)

64 (49)

15 (35)

28 (65)

43 (12)

74 (45)

92 (55)

166 (49)

341

Area

Totals

1976

Middle Park

Adult
Male

132 (39)

I

t-'

o

Totals

Eagle

7 (50)

7 (50)

14 (21)

0 ( 0)

8(100)

8 (12)

15 (33)

30 (67)

45 (67)

67 l'

North Park~/

6 (67)

3 (33)

9 (69)

0 ( 0)

0 ( 0)

o ( 0)

1 (25)

3 (75)

4 (31)

13

Poudre Canyon

5 (42)

7 (58)

12 (33)

1 (25)

3 (75)

4 (11)

6 (30)

14 (70)

20 (56)

36

Routt

20 (62)

12 (38)

32 (51)

4 (67)

2 (33)

6 ( 9)

11 (44)

14 (56)

25 (40)

63

Moffat

1 (33)

2 (67)

3 (19)

0 ( 0)

2(100)

2 (12)

7 (64)

4 (36)

11 (69)

16

107 (53)

95 (47)

202 (38)

20 (32)

43 (68)

63 (12)

114 (42)

157 (58)

271 (50)

536

1/ Because some subadu1ts had completed their primary molt and thus could not be distinguished from adults, number of
birds assigned to the subadult category was minimal.

1/ Number in parentheses represents percent.
1/ Most birds checked in North Park in 1976 were harvested in Routt County.

�-101-

Exp_erimental Season
Based on information collected in this study, along with knowledge gained
from intensive studies in other western states and provinces (Bendell and
Elliott 1967, Mussehl 1960, Harju 1974, Zwickel 1958, among others) the blue
grouse season in Colorado could be liberalized without harm to breeding
populations in subsequent years.
Length of seasons in Utah (69 days),
Nevada (44 days), Montana (80-90 days) and Oregon (32 days) have had no
measurable effect on their grouse populations.
Bendell and Elliott (1967)
reported that after replacements for mortality in the breeding population
are set aside, approximately 30 percent of the autumn population may be
harvested.
In view of the strong justification for longer seasons, an experimental blue
grouse season was held in Middle Park (Small Game Unit 28) in 1976. The
regular season opened statewide on September 11 and closed October 10 except
in Unit 28 where the season was extended to October 26 in conjunction with
the regular elk season (October 16 to 26). In order to evaluate the season,
a questionnaire was sent to all holders of cow elk validations (1,450) for
Big Game Units in Middle Park. The total population of elk hunters (regular
bull elk and cow elk) (6,719) was obtained from estimates based on the statewide survey of big game hunters (pers. comm. Harlan Riffle, Division of Wildlife). These estimates tend to be over inflated, but are the best available.
About 22 percent of the elk hunters in Middle Park received a questionnaire.
Questionnaires
(1,450) were sent immediately following the elk season.
Responses were received from 1,017 hunters (70%). In late November a follow
up letter was sent to all non-respondents
(433) and 246 additional responses
were received for a total return of 1,263 surveys (87% return) (Table 9).
Only 8 questionnaires were non-deliverable.
Mean values calculated for
hunters responding to the follow up survey were used to project for the 13
percent (187) non-respondents.
It is apparent from data in Table 9 that few elk hunters (23.0%) were even
aware of the experimental season, and only 18.6 percent (4.8% of total sample)
of those aware of the season actually participated.
Nearly half (48.4%)
of the elk hunters that hunted grouse were successful in harvesting 54 blue
grouse, for a hunter success of 1.8 birds per successful hunter (.87 birds/
elk hunter hunting grouse).
Applying these estimates to the total elk
hunter population gives the following estimate of the total harvest of blue
grouse in Middle Park during the experimental season:
Total elk hunter population
23 percent aware of season
18.6 percent participation
48.4 percent successful
1.8 grouse per successful hunter
Plus 8.2 percent crippling loss

6,719
1,545
287
139
250
270 Total Blue Grouse
Harvested

Crippling loss is based on the regular season estimate of 8.2 percent as
determined from data collected on hunter check stations (Table 7). The
calculations do not take into account those individuals that hunted blue
grouse, but were not elk hunters.
However, during field checks no such
individuals were contacted and it is believed their numbers were minimal.

�-102-

Table 9. Middle Park experimental
data, 1976.

blue grouse season, hunter questionnaire

1,017
No. in sample
70.1
Percent return
Number elk hunters aware
215
of season
Percent elk hunters aware
21.1
of season
No. inactive elk huntersl/
81
Percent inactive elk
8.0
hunters
No. elk hunters hunting
55
grouse
Percent elk hunters
5.9
hunting grouse l/
No. elk hunters not
881
hunting grouse
Percent elk hunters not
94.1
hunting grouse
No. of successful grouse
23
hunters
Percent successful grouse
41.8
hunters 1/
Percent of elk hunters
2.5
harvesting grouse l/
42
No. of grouse bagged
.04
Grouse/elk hunter
Grouse/elk hunter
.76
hunting grouse
Grouse/successful
grouse
1.8
hunter
No. elk hunters observing
366
grouse
Percent elk hunters
39.1
observing grouse l/
Total blue grouse observed 2,216
8
Non-deliverable
surveys
Percent non-deliverable

l/ Inactive

elk hunters

Projected
For

Projected
For 1,450

246
17.0

1,263
87.1

187

12.9

1,450
100.0

67

282

51

333

27.2
45

22.3

126

27.2
34

23.0
160

18.3

10.0

18.2

11.0

4

59

3

62

2.0

5.2

2.0

4.8

197

150

1,078

1,228

98.0

94.8

98.0

95.2

4

27

3

30

100.0

45.8

100.0

48.4

2.0
7
.03

2.4
49
.04

2.0
5
.03

2.3
54
.04

1.7

.83

1.7

.87

1.7

1.8

1.7

1.8

74

440

56

496

36.8
465

38.7
2,681
8

36.8

352

38.4
3,033
8
.5

o

are those individuals

o

that had a license but did not

hunt.

l/ Percentage

calculations exclude
in the field to participate.

inactive

3/ Based on number elk hunters hunting

elk hunters

grouse.

since they were not

�-103-

Middle Park encompasses about 5,960 km2 (2,000 mi2) of which about 3,370 km2
(1,300 mi2) is estimated to be forest (potential blue grouse breeding
habitat) (Wallmo et ale 1977; pers. comm. R. B. Gill, Division of Wildlife).
Assuming an average minimum fall density of about 8 birds per km2 (20 birds
per mi2), the total estimated blue grouse population in Middle Park is
26,960 birds. From this population, only 270 birds or approximately 1
percent were harvested during the experimental season.
Examination of Table 10 reveals that: (1) elk hunters hunting grouse observed
more birds per hunter (3.3) than grouse hunters during the regular season
(1.0); (2) hunter success (birds bagged/hunter) was higher in the experimental season (.9) than in the regular season (.4); and (3) more hunters
were successful in the experimental season (48.4%) than in the regular
season (41.0%). Thus, the potential for increasing the harvest of blue
grouse by extending the season into the big game season exists provided more
hunters would participate.
Few hunters (23%) were aware of the experimental
season, consequently, participation was low. However, based on information
collected in Washington, many hunters will not harvest grouse during the big
game season for fear of spoiling their chances of taking a big game animal.
Therefore, even if more elk hunters had been aware of the season, only a
small percentage would have participated with the increase in harvest having
negligible
effects on the grouse population.

Table 10. Comparison of regular season and experimental
for blue grouse in Middle Park 1976.

Regular-~j
Season

Harvest Data

season harvest data

Experlmenta
.
12/
Season

Number of hunters

l38

62

Number birds observed

141

205

Birds observed/hunter

1.0

3.3

Number birds bagged

61

54

Birds bagged/hunter

.4

.9

Percent successful

hunters

41.0

48.4

Hunter efficiency

(percent)

43.3

26.3

l/ Based on information
]j

presented

in Table 7.

Based only on those elk hunters surveyed.

�LITERATURE

CITED

Bendell, J. F, D. G. King, and D. H. Mossop.
1972. Removal and repopulation
of blue grouse in a declining population.
J. Wildl. Manage. 36(4):
1153-1165.
_____ , and P. W. Elliott.
1967. Behavior and the regulation
blue grouse.
Can. Wildl. Servo Rep. Ser. 4. 76 p.

of numbers

Boag, D. A. 1965. Indicators of sex, age, and breeding phenology
grouse.
J. Wildl. Manage. 29(1):103-108.
Harju, H. J.
grouse.

in

in blue

1974. An analysis of some aspects of the ecology of dusky
Ph. D. Thesis. Univ. Wyoming, Laramie.
142 p.

Hoffman, R. W. 1976. Population dynamics and habitat relationships of blue
grouse. Colorado Div. Wildl. Job Prog. Rept. Fed. Aid Proj. W-37-R.
April 1976. p. 135-152.
, and C. E. Braun.

1975.

A volunteer wing collection
3 p.

-----Div. Wildl. Game Information Leaflet No. 101.

station.

Colo.

Mussehl,
T. W. 1960. Blue grouse production, movements, and populations
in the Bridger Mountains, Montana.
J. Wildl. Manage. 24(1):60-68.
Rogers, G. E. 1968. The blue grouse in Colorado.
and Parks. Tech. Publ. No. 21. 63 p.

Colo. Div. Game, Fish

Stauffer, J. E. 1953. Geology of an area west of Wolcott,
M. S. Thesis. Univ. of Colorado, Boulder.
Colorado.

Eagle County,
62 p.

U. S. Department Commerce, Weather Bureau.
1966-1976.
Climatological data.
Annual summaries, Colorado.
U. S. Government Printing Office, Washington, D. C.
Wallmo, o. C., L. H. Carpenter, W. L. Regelin, R. B. Gill, and D. L. Baker.
1977. Nutritional basis for evaluating deer habitat.
J. Range Manage.
30(2):In Press.
Zwickel, F. C. 1958. North-central
State Game Bull. 10(4):3-4.

Washington

grouse studies.

Wash.

1972. Removal and repopulation of blue grouse in an increasing
population.
J. Wildl. Manage. 36(4):1141-1152.

�-105-

April

1977

JOB FINAL REPORT

State

of _-..:::C~O:=L:.::::O~RAD=:!··~'
'~O
_

Project

No.

W-37-R-30
i7

Work Plan No.
Job Title
Period

Game Bird Survey
job No..

2

-------------------------------

Cont.Lnued Irivent.or y of Se.Lec t ed Pi:armigan Popul.at Lons
--~~~~~~~~~~~~~~~~~~~~==~~~----------

Covered:

April

l,

197d to 31 March i977

w.

Per sonne'l i

t~rry A~.Mciy ~ University
of coio:bido;
.Johri Ai-thur, Clait E.
Braun, iIot.r.i:rdD. Furik; Kenneth M.Gieseri, .RichardW.
Hoffman,
T?iil;.i.:i1:+~,. Rart Ociideaf, and charles w&lt;igri~r, Colorado Division
of Wildlife.

The objecHvesof
tliis EltudY ii~ve been ftiliy achieved.
Reports covering the
various objectives
ha\tebeen
p~blished,
prepared,arid
submitted,
and in some
Lns tarices , are in .i):hjgtess~ .THo~eiriprogr~sswili
be published under Work
Plan 22, Job iand are listed
ih the project
documents for W-37~R-3l.
Those
pUblished are iisted
beiow.

c. E.~~rid

May.

Brauri,
T.A.
1972.
co16radoaipine
tundra avifaunal
gations,i§g677l
.. ~~~i~3-i~g
in Proc , 1972 n. s. Turidra Biome
uriiv. wash:l.ngtori.

investi-

SyInpo~{Jin;

___

, R. K.

tailed
90-93.

Schmidt; JL j arid G. E. Rogers. 1973. Census of Colorado whitewith tape-recorded cails.
J. Wildi. Manage. 37(1):

ptarmigan

____ , andi.
G~Biun.ib~rg.
Condor 75(3):345-346.
1975.
Seasonal
Wilson brnii:hoi.
(Abstract).
___

197:3.

Hahitat

An albinistic

selection

blue

grouse

by white-taiied

Sac. and Cooper Orriithol.

Soc .. Joint

from Colorado.

ptarmigan.
Proc.
Mtg. 2: 14.

, R. w. Hoffman, and G. E. Rogers.
1976. Wintering areas and winter
ecology of whii:~-taii~d
ptarmigan in Colorado.
Coiorado Div. Wildlife
Spec. Rept.
pp.

No. 38. 38

:. I . ':,,'

-~.

.....,.
t

-'.

.

Hoffman, R. W., arid C. E. Braun.
1973. Migration of white-tailed
to and froina majot iviritering area .. Proc. Aim. Conf , Central
and Plains Sec. ~ The Wildl. Soc., 18:i6-l7.
(Abstract).
___

, and
flocks
68-69.

1974.
Composition,
of whih~-tailed
ptarmigan.
(Abstract).

ptarmigan
Mtns.

d i s t r Lbut Lon, and siz~ of winter
J. Colo.-Wyo. Acad , ScL 7(5):

�-106-

Hoffman, R. W., and C. E. Braun. 1975. Migration of white-tailed
in Colorado.
J. Colo.-Wyo. Acad Sci. 7(6):45.
(Abstract).

ptarmigan

_____ , and
1975. Migration of a wintering population of whitetailed ptarmigan in Colorado. J. Wildl. Manage. 39(3):485-490.
_____ , and
1977. Characteristics of a wintering population of whitetailed ptarmigan in Colorado. Wilson Bull. 89 (1):107-115.
Marti, C. D., and C. E. Braun. 1975. Use of tundra habitats by prairie
falcons in Colorado.
Condor. 77(2):213-214.
May, T. A. 1975. Physiological ecology of white-tailed ptarmigan in
Colorado.
Ph. D. Thesis. Univ. of Colorado. Boulder. 311 pp.
1975. A population bioenergetics model for Colorado white-tailed
ptarmigan.
U. S. Tundra Biome Rept. 75-1. 89 pp.
_____ , and C. E. Braun. 1972. Weight dynamics of free-living white-tailed
ptarmigan in Colorado.
J. Colo.-Wyo. Acad. Sci. 7(3):70-71.
(Abstract).
_____ , and
_
in Colorado.

1972. Seasonal foods of adult white-tailed
J. Wild1. Manage. 36(4):1180-1186.

_____ , and
_
in Colorado.

1973. Gizzard stones from adult white-tailed
Arctic and Alpine Res. 5(1):49-57.

ptarmigan

ptarmigan

Stabler, R. M., N. J. Kitzmiller, and C. E. Braun. 1974. Hematozoa from
Colorado birds.
IV. Galliformes.
J. Parasitol. 60(3):536-537.

Prepared by

.!.&lt;ttaI~=-::....___2~.
-.:~~
Clait E. Braun
Wildlife Researcher

_

�-107-

April 1977

INTERIM JOB PINAL REPORT
State of

COLORADO
--~~~~~----------

Project

No.

Work Plan No.
Job Title
Period

17

Organization,

Covered:

Personnel:

Game Bird Survey

W-37-R-30

Job No.
Mortality,

5

and Dispersal

of Grouse Broods

1 April 1976 to 31 March 1977

William W. Mautz, Colorado State University; Clait Braun,
Deborah Covic, Ken Giesen, Richard Hoffman, and Jim Kitzmiller,
Coloradq Division of Wildlife.

ABSTRACT
Investigations of mortality and movement.s of juvenile ~hite-tailed
ptarmigan (Lagopus le~curus) were c01-ld~ctedon 16.9 km of alpine
habitat along Trail Ridge Road in Rocky Mountain National Park, Colorado
from May 1975 through Japuary ~977. Emphasis was placed on q.etermination
of causes and tirp.ingof juvenile mortality and fall dispersal movements.
Nesting success varieq frotp.60 percent (1975) to 54 percent (1976) ~ith
all nest loss attribqteq. to predation.
Two instance~ of renesting were
documented in 1975. Clutch size averaged 5.6 eggs with 89 percent of
the eggs in s~ccessful 1-lestshatchi1-lg. Infertility and embryonic
mortality were responsible for failut-e of eggs in successful nests to
hatch. Movements to pj..lJWller
habitat i3.veraged350 m for seven broods
with known nest sites .. Repeateq observations of nine broods indicated
summer movements were restt-icteq.withi1-la1-laverage home range of 15.4
hectares.
ObservatiQ1-ls of 49 marked broods indicated chick mortality
approached 34 percent wiJ:hi1-l
two weeks posthatch with predation being
a primary cause. Once chicks reached two weeks of age, additional
mortality was less tha1-l10 Percent until brood breakup.
Brood breakup
preceded dispersal anq. began gradually once ch~cks reached eight weeks
of age. Brood break~p and dispersal were facilitated by early fall
snowstorms which causeq. broods to abandon sqmmer habitat and move downhill
into protected areas ~here willow (Salix spp.) was abundant.
Fall
movements were docurp.ented for 42 j~venile females and 32 juvenile males.
Fall dispersal movementS (&gt; 1.0 km) were significantly greater (P &lt; 0.05)
in juvenile females tha1-lj~venile male~ with movements up to 8.4 km
being recorded.
DiSPersing individ~als moved along continuous alpine
habitat rather than f1-lrandom directions.
Movements between brood
range and wintering sites were documented for nine juvenile females and
six juvenile males.
J~venile females tended. to migrate greater distances
than juvenile males with one movement of 9.2 km being recorded.

�-108-

TABLE OF CONTENTS

Page
INTRODUCTION

.••••.••••••••.•••.....•••.•.••••..••.••••.....••••••

REVIEW OF LITERATURE

•..•...••.....••••...•••••••.•...•..••..••••.

LOCATION AND DESCRIPTION
METHODS AND MATERIALS

OF ~HE STUDY AREA .•••••.•••••.•..••••••.

••••••.••.••••••••••••••••••••••••.••••.•••.

III

112
112
114

RESULTS •••..••..••.•..•....••....••.••..•.•.•..••.•••.•••.•.••.••
Nesting
•..•••.•.••.••••.•••••.•••••..•••••..•.••••.•••.•..•••.
Clutch
Size
.•.•••.••••..••..•.•••••..•••••.••••.•.•..•.•...
Ha:tchability
•...••..•.•...••••.••••••...•••.•••.••••.••.••.
Nes ting
Success
•..•••••...••.••••••.....••••.••••••......•.
Mortality
•.•.•.••..••..•.•..•.•.•.••.••••.•..••.•••••.•.••.•.•
Timing
••..•.••••.••••.••..•.......•••••.•.•••••.•••••..••••
Causes
.••••.•.•••••.••••..•.••.•••••..•.•••..•.••••..••••••
Dispersal
•.•••••••.••...•••.•..•..•...•••.....•.•...•••••...•.
Movements
to Summer Habitat
.•••••.••.•••••..•....••...•.•.•
Movements
Within
Brood Home Range
•••.•••••....•••..•..••••..
Brood Breakup
•.•••••••••••••.•••••••....••.••...••.••.•.•••
Fall
Dispersal
•••...••••.••••.•.•••.•••..•...•••..••....•.•
Summer to Winter
Movements
•••..••••••...•.••..••••••..•.•..

" 115

DISCUSSION
Mortality
Dispersal

132
132
135

•...•.•••.••••••••••.•••••.•••••••..•••..•••••..•••..••
••.•••••.••••••••••••.•••••••••••.•••••••••••..••..•.
•••••.••..•.••.•.••••.•.•••••...•..•..•••••.•.•.•••••

SUMMARY .•.••••..•••.••••.•••••.•••••••••••.•••••••.•••••••••..•..
LITERATURE

CITED

.................................................

115
115
116
116
117
117
119
121
121
122
122
123
129

136
137

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LIST OF TABLES

Table
1

2

3

4

5

6

Number of white-tailed ptarmigan broods and average
size, 1975 and 1976 ••...•....•..•.....................•.•.•

ll8

Radio tracking data for nine juvenile white-tailed
ptarmigan in 1976 .•..•... ,.......•••.........•.•..••....•.•

124

Fall movements
Rocky Mountain

125

of juvenile white-tailed ptarmigan in
National Park, 1976 .••.....•......•...••....

Hypothetical production of white-tailed ptarmigan chicks
per 100 females .•......••.......•..•.......................

133

Nesting success of white-tailed ptarmigan hens observed
both years (1975 and 1976) .••.•..•...•......•.....•....••.•

134

Comparative brood size of white-tailed ptarmigan
hens observed both years (1975 and 1976) •.•...•..•••.•.....

135

LIST OF FIGURES

Figure
1

Trail Ridge Study Area, Rocky Mountain

2

Individual average fall movements of juvenile whitetailed ptarmigan .••..••••..............•..••.•....•••....

126

Individual average dispersal movements of juvenile
white-:-tailed ptarmigan .........•.•...•.•..........••..•.•

127

Dispersal movements of male and female juvenile whitetailed ptarmigan •..•••...•..........•....•........•..••••

128

Distance
juvenile

and direction of dispersal movements of
white-tailed ptarmigan •.•..•..•••.••............

130

Movements
summering

of juvenile white-tailed ptarmigan from
areas to wintering sites .......•...•••...••..••

131

3

4

5

6

National

Park

113

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ACKNOWLEDGMENTS

Financial support was provided with a graduate research assistantship
through the Colorado Cooperative Wildlife Research Unit and the Colorado
Division of Wildlife, Federal Aid in Wildlife Restoration Project W-37-R.
The support of these agencies is gratefully acknowledged.
The U.S. Department of Interior, National Park Service, Rocky Mountain
National Park allowed access to the study area and provided necessary
permits.
The cooperation of Dave Stevens, Park Biologist, is sincerely
appreciated.
Dr. William W. Mautz, former Assistant Leader of the Cooperative Wildlife
Research Unit, served as my advisor and critically reviewed this thesis.
His encouragement and interest are appreciated.
Other members of my
graduate committee, Dr. Ronald A. Ryder, Department of Fishery and
Wildlife Biology, and Dr. Philip N. Lehner, Department of Zoology and
Entomology reviewed this thesis and were available for consultation
throughout the study.
Dr. Clait E. Braun, Wildlife Researcher, Colorado Division of Wildlife,
served on my graduate committee and was responsible for initiating the
study.
In addition to expert assistance in the field he provided administrative support which made my work easier.
His knowledge, patience
and enthusiasm for research constantly stimulated my research efforts.
His interest in my progress as a student and development as a biologist
is sincerely appreciated.
The U.S. Fish and Wildlife Service, Denver Research Center, provided
radio transmitters which facilitated my work and provided data not
otherwise easily obtainable.
The expertise and generous help of Gene
Bourassa and Larry Kolz is greatly appreciated.
Others instrumental in assisting in various aspects of this study
included Howard Funk, Colorado Division of Wildlife, for administrative
support and review of the thesis; Ed Blekicke, Nik~ Goodson, and Jim
Kitzmiller provided assistance in the field at critical times. The
assistance of these people is sincerely appreciated.

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ORGANIZATION,

MORTALITY,

AND DISPERSAL

OF GROUSE BROODS

INTRODUCTION

Grouse are important game birds in Colorado and four species, blue grouse
(Dendragapus obscurus), sage grouse (Centrocercus urophasianus), sharptailed grouse (Pedioecetes phasianellus), and white-tailed ptarmigan
(Lagopus leucurus) presently provide considerable hunting opportunity.
Little is known concerning broods of three of these species (blue grouse,
sage grouse, and sharp-tailed grouse) due to habitats occupied and lack
of detailed study. Production of young and survival to the hunting
season and following breeding season are important biological parameters
as these factors affect fall hunting populations and subsequent spring
breeding populations.
It is thus important to understand behavior and
dispersal of broods as they may directly affect hunting success and overwinter survival.
It is also important to identify major mortality
factors of juvenile grouse and to understand their relationship to fall
and subsequent spring population levels.
Intensive studies of white-tailed ptarmigan in Colorado (Schmidt 1969,
Braun and Rogers 1971), Hoffman and Braun 1975) support the conclusion
that the biology of this grouse is similar to that of other tetraonids,
especially in respect to brood ecology and dispersal.
Preliminary
observations of white-tailed ptarmigan broods suggested they would be
ideal study subjects as (1) they can be easily and readily located,
(2) they can be easily marked, (3) they occupy small home ranges until
time of dispersal, (4) they can be observed without vegetative interference,
and (5) seasonal habitat requirements are known.

P.N.O. OBJECTIVE
The objective of this study was to increase the knowledge of factors
affecting numbers of young grouse in fall and recruitment to the breeding
population the following spring.
It was expected that data concerning
time of and factors affecting dispersal and mortality of young grouse
could markedly affect timing and duration of fall hunting seasons.
Hunting seasons could be established to coincide with periods of dispersal
and high natural mortality in order to more fully utilize birds that are
being lost due to natural factors.
Hypotheses which were tested were:
(1) mortality of chicks is predator-dependent
to 1 October and weatherdependent from 1 October to 1 November; (2) brood breakup and dispersal
are functions of chick aggression to brood mates; (3) timing of dispersal
is age-dependent with dispersal not occuring until chicks are older than
eight weeks; and (4) direction of dispersal is non-random.

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REVIEW OF LITERATURE
Little detailed information on the ecology of white-tailed ptarmigan
had been published prior to 1960. Most early studies concerned distribution or notes on life history; these have been summarized by Braun
(1969). Recently, intensive studies have occurred in Montana and Colorado.
Choate (1960) investigated breeding biology of ptarmigan in Glacier
National Park, Montana and later (1963) studied population dynamics and
ecology of the same unhunted population.
Since 1966 intensive research
efforts have been directed at various aspects of white-tailed ptarmigan
ecology in Colorado.
Braun (1969) and Braun and Rogers (1971) investigated population dynamics on hunted and unhunted areas, life history and
distribution in Colorado.
Other studies have investigated parasites of
ptarmigan (Haskins 1969, Stabler et al. 1974), food habits, physiology
and energetics (May and Braun 1972, May 1975), breeding behavior (Schmidt
1969), habitat requirements (Braun 1971, Braun and Schmidt 1971, Braun
et al. 1976), and winter ecology (Hoffman 1974, Hoffman and Braun 1975).
Except for limited work by Schmidt (1969) little data concerning brood
ecology, especially mortality and dispersal, has been published.
Various aspects of brood mortality, movements and behavior have been
investigated in several grouse species in areas other than Colorado.
Studies of ruffed grouse (Bonasa umbellus) (Bump et al. 1947, Gullion
and Marshall 1968), blue grouse (Stiven 1961, Zwickel 1967, Zwickel and
Bendell 1967a), red grouse (Lagopus lagopus scoticus) (Jenkins et al.
1963), and rock ptarmigan (L. mutus) (Theberge 1971, Theberge and West
1973) have discussed causes-and timing of mortality in juvenile grouse.
Brood movements including dispersal have been investigated for ruffed
grouse (Chambers and Sharp 1958, Hale and Dorney 1963, Godfrey and
Marshall 1969, Godfrey 1975), blue grouse (Wing et al. 1944, Zwickel
et al. 1968), spruce grouse (Canachites canadensis) (Ellison 1973),
greater prairie chickens (Typanuchus cupido) (Robel et al. 1970, Bowman
and Robel 1977), and sage grouse (Wallestad 1971). Zwickel (1965)
reviewed much of the literature concerning mortality in grouse and other
animals and Keppie (1975) reviewed some of the literature on grouse
movements, including dispersal, and also discussed the relation of
dispersal to animal populations.

LOCATION

AND DESCRIPTION

OF THE STUDY AREA

Field investigations were conducted in Rocky Mountain National Park
(RMNP) along Trail Ridge Road (Fig. 1) between 1 May 1975 and 31 January
1977. Previous studies by Schmidt (1969) and Braun and Rogers (1971)
indicated a relatively high population of white-tailed ptarmigan utilized
this area from April to October.
Also, previous investigations indicated
that in late summer, female ptarmigan with broods were attracted to a
few concentration areas (Braun pers. comm.) which facilitated location
and observation of broods.
The study area was in Larimer County, Colorado, at approximately 105°44' W.
longitude, and 40°25' N. latitude.
Included were T5N, R74W, parts of

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Sections 6, 7, 8, 9, 15, 16, 17, 18, 21, 22, and 27; T6N, R74W, parts
of Sections 30 and 31; T5N, R75W, parts of Sections 1, 2, and 3; T6N,
R75W, parts of Sections 35 and 36. Total area was approximately 16.9
km2 (6.5 mi2).
Elevation ranged from 3,414 m to over 3,799 m. In
1976 some chicks were marked on Mt. Chiquita, Specimen Mtn. and the
Continental Divide between Mt. Ida and Sheep Rock. These areas lie
east, northwest and west of Trail Ridge, respectively, with all less
than six km distant.
The climate of the Front Range has been described by Judson (1965) and
Marr (1961, 1967). Weimer and Haun (1960) and Quam (1938) described
the geology and physiography of the Front Range, while Willard (1963)
and Eddleman (1967) characterized the vegetation.
A more detailed
physical description of the Trail Ridge area is provided by Braun
(1969) and Braun and Rogers (1971).

METHODS AND MATERIALS
Territorial pairs in spring and females in summer were located with
tape-recorded calls (Braun et ale 1973). Intensive search on foot
and frequent viewing with binoculars were necessary to locate nonterritorial males and unsuccessful females.
Once ptarmigan were lcoated they were observed using 7 x 50 or 8 x 36
binoculars.
If the bird had been banded previously its band color and
number were recorded, along with the time and location, on standardized
observation cards. Unbanded birds were pursued until captured or flushed
out of sight. Captures were made with a 27.3 kg (60 pound) test nylon
covered wire snare (9.1 kg for chicks) attached to the end of a five
or seven meter telescoping fiberglass fishing pole as described by
Zwickel and Bendell (1967b) for blue grouse, and adapted for ptarmigan
by Braun and Rogers (1971). All locations were later plotted on 7.5
minute (scale 1:24,000) U.S. Geological Survey topographic maps.
Each adult ptarmigan newly captured was banded on the right leg with a
gold aluminum serially numbered band (size 8, National Band and Tag
Co., Newport, Kentucky) and on both legs with colored, numbered (1-100)
plastic bandettes (size 6, National Band and Tag Co., Newport, Kentucky).
Green bandettes were used in 1975 and yellow bandettes in 1976. Chicks
captured after two weeks of age were banded on the left leg with a gold
aluminum serially numbered band.
Selected chicks (45 in 1975, 46 in
1976) were additionally banded with various combinations of colored
plastic coil bandettes (size 3, National Band and Tag Co., Newport,
Kentucky).
In 1976 all chicks captured or recaptured after 1 September
were banded with colored (pink) numbered plastic bandettes similar to
those used on adults.
Captured ptarmigan were weighed with a spring scale accurate to within
five grams for adults and two grams for chicks. Measurements of carpal
length, individual primary lengths and length of the outer rectrix were

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recorded using a transparent, flexible plastic millimeter ruler. Adults
were sexed and classified as either adult or yearling according to plumage
characteristics described by Braun and Rogers (1971). Chicks were aged by
examination of various plumage characteristics including lengths of
carpal and primary feathers and by progression of primary molt. Ages
were assigned by comparing specific measurements with those of known-age
chicks.
Sex of chicks was based on behavior (vocalizations and aggression),
primary feather and carpal lengths, pigmentation of the eye-stripe and by
the amount of pigmentation on post-juvenile primary (P) 8, juvenile P9,
PIO and primary covert 10 (Braun, pers. comm.).
Nests were located by (1) observation of laying hens, (2) following
incubating hens back to nests after feeding, and (3) intensive search
of known territories.
Additional nests were located incidental to other
field activities.
In 1976, 174 MHz radio transmitters (U.S. Fish and Wildlife Service,
Denver Research Center) were placed on three females with broods and
on nine chicks at 12 to 16 weeks of age. Transmitter packages weighed
12-13 gms and were attached to the central rectrices of the birds with
a bolt and clamp device similar to the tail clip described by Bray and
Corner (1972). A model LA 12 receiver (AVM Company, Champaign, Illinois)
and a hand-held three element yagi antenna (Model 23 Hy-Gain, Wildlife
Materials, Inc., Carbondale, Illinois) were utilized in radio location
of transmitter-marked birds.
Differences in clutch sizes, brood sizes and dispersal movements were
statistically tested using the Students t-test unless otherwise noted.

RESULTS
Nesting
Clutch Size
Clutch size was determined from 18 complete clutches examined in 1975
and 1976. Three additional nests were located but the clutches were
destroyed by predators before completion or before examination.
Average
clutch size was 5.6 and ranged from four to seven. Clutches of five or
six were most common and comprised 83 percent (15 of 18) of the complete
clutches examined.
Clutches larger than seven are thought to be uncommon,
while clutches of four or less are thought to represent renesting efforts
(Braun and Rogers 1971). Renesting was documented in 1975 when two marked
hens were observed on summer range with broods after their initial nests
were known to have been destroyed.
There were no significant differences
(P &gt; 0.05) in clutch size between years or between age classes (yearlings
versus adults).
Braun and Rogers (1971) examined 21 complete clutches
and computed an average clutch size of 5.7 eggs. Choate (1963) calculated
an average clutch size of 5.2 eggs based on ten complete clutches in
Montana.

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Hatchability
Hatchability was based on the percentage of eggs hatching in successful
nests (those nests incubated to completion).
In 1975 hatchability and
egg fertility were 100 percent as all eggs (n = 32) in six successful
nests hatched.
Hatchability declined in 1976 to 74 percent (17 of 23
eggs) in four successful nests.
This decrease was due to one nest (Y58)
in which only four of seven eggs hatched.
One fertile egg cracked and
was removed by the observer.
Two other fertile eggs chilled after the
hen apparently dislodged them when flushing from the nest. Another
two eggs, possibly infertile, were buried under the nest cup and were
not incubated.
The remaining egg was found to be infertile.
Thus
the minimum egg fertility in 1976 decreased to 87 percent.
Braun (1969)
reported that 81.1 percent of the eggs (n = 37) in successful nests
hatched while Choate (1963) reported 81.2 percent (26 of 32 eggs) in
seven successful nests hatched.
Neither Braun nor Choate calculated
egg fertility.
Nesting

Success

Nesting success was estimated from (1) success of known nests, (2)
numbers of successful and unsuccessful hens (those hens observed with
and without broods, respectively) seen during the brood season and
which had been identified during the breeding season, and (3) total
numbers of successful and unsuccessful hens observed during the brood
season.
Six of 11 active nests (55 percent) located in 1975 hatched successfully,
but this estimate of nesting success may be inflated since only two of
seven nests (29 percent) located prior to onset of incubation were successful. In 1976 only four of ten active nests located hatched successfully
(40 percent), with only one of four nests located prior to incubation
hatching successfully (25 percent).
Renesting compensated for some
nest loss in 1975 as two of five hens who lost their initial nests renested successfully.
None of the five hens (excluding one which was
killed on the nest) who lost their initial nests in 1976 was known to
renest successfully.
During the 1975 breeding census 31 females were identified.
All females
were observed with territorial males and were assumed to be paired.
Of
the 31 females, 22 were later observed during the brood season (13
successful, 9 unsuccessful), resulting in an estimated nesting success
of 59 percent.
This estimate may be biased as success of nine females
identified on breeding territories was not ascertained.
Braun (unpublished data) examined the ovaries of females collected during the
breeding season and determined that all females ovulated.
Field observations of mated pairs also indicated that all females attempted to nest.
Since the brood survey was thorough, it is unlikely that many successful
females escaped detection.
Possibly some females died prior to or during
nesting or nested elsewhere.
Another possibility is that the majority
of these hens were unsuccessful and were not observed during the brood

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season because

they summered off the study area (Braun pers. comm.).

During the 1976 breeding season, 42 females were identified of which
30 were observed during the brood season (16 successful, 14 unsuccessful) for an estimated nesting success of 53 percent.
Again a large
number of hens (11) disappeared between the breeding and brood seasons
(one hen was killed on the nest).
It is not known what proportion of
these 11 hens was successful.
Another method of estimating nesting success was employed which
eliminated the need for using the number of hens observed during the
breeding season.
This method was not biased by any hens missed during
the breeding census. The technique is to estimate nesting success using
the proportion of successful hens to unsuccessful hens observed during
the brood season, regardless of any identification during the breeding
season.
The assumption is that successful hens remain on the area while
immigration and emigration of unsuccessful hens is equal. Since the
study area is large (16.9 km2) and somewhat isolated from other alpine
areas this assumption may be true.
During the 1975 brood season (20 Ju1y-13 October) 36 hens were identified
on the Trail Ridge area (22 successful, 14 unsuccessful).
This estimate
of nesting success (61 percent) appears realistic since the complete
study area was searched weekly, even though successful hens are more
likely to be located than unsuccessful hens (Braun and Rogers 1971).
In 1976, 50 hens (27 successful, 23 unsuccessful) were observed during
the brood season (10 Ju1y-30 September) for an estimated nesting success
of 54 percent.
The last two techniques arrive at similar estimates (±2 percent) for
each year. Nesting success in 1975 was slightly higher than in 1976
(60 percent versus 54 percent).
Mortality
Timing
Timing of mortality was based on observed decreases in average brood
size and calculated for bi-week1y intervals (Table 1). From 20 July
to 15 October 1975, 128 observations of broods were recorded compared
to 157 brood observations between 10 July and 30 September 1976. The
different interval sampled between years reflected the earlier peak of
hatch in 1976 and corresponding earlier brood breakup.
The difference
in number of observations was due to a higher number of broods on the
study area in 1976 (27 versus 22), radio location of three hens in
1976 and observer experience.
It is apparent that a large decrease in brood size occurred within
two weeks of hatch (Table 1). The peak of hatch in 1975 was between
20 and 25 July and broods located between 1 and 15 August (n = 10)
had decreased in average size by 30.4 percent.
In 1976 the peak of

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hatch occurred between 10 and 20 July and average brood size on 31 July
was 3.7, a decrease of 33.9 percent from hatch.
Some of the observed
decrease in brood size was attributed to renesting which usually resulted
in smaller initial brood size (four or fewer chicks).

Table 1. Numbers of white-tailed
1975 and 1976.

No. of
Broods
Time Interval

ptarmigan

broods and average size,

Brood
Observations

Avg. No. of
Chicks/Brood

1975

1976

1975

1976

1975

1976

1-31 July

5

15

13

42

5.6

3.71,2/

1-15 August

10

12

23

41

3.9])

4.1

16-31 August

11

9

27

24

3.9

3.8

1-15 September

9

9

18

26

3.6

3.2

16-30 September

8

11

23

24

4.1

3.6

1-15 October

10

!/Significant

difference

~/Significant

decrease

24

4.2

between years (P &lt; 0.05).

from hatch or previous

time interval

(P &lt; 0.05).

After the initial mortality in the first two weeks post-hatch, there
was little additional mortality (as reflected in average brood size)
to September.
After chicks reached eight weeks of age, broods began
to break up. At this time some hens lost all their chicks, while other
hens were observed with as many as 12 chicks from several broods.
"Gang"
broods were responsible for the September increase in average brood size
shown in Table 1.
Timing of mortality was closely monitored for members of three broods
in 1976 in which the hens were fitted with radio transmitters.
The first
hen (Y90) was radio marked on 12 July when her clutch of six eggs had
pipped.
She left the nest on 13 July and was located on 14 July with
all six chicks.
The transmitter then failed, but I relocated the hen
on 17 July at which time only one chick remained in her brood. No sign
was found of the other chicks which had been banded at hatch.
The remaining
chick was observed periodically until 11 November.

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A second female (G72) was radio marked on 22 July when her brood of
four chicks was approximately 15 days old. It can be assumed that
if her clutch size was average she had lost approximately 30 percent
of her brood. Later observations show that one chick was last observed
on 11 August, while the remaining three chicks were observed periodically
until brood breakup in late September and early October.
A third female, G97, was radio marked on 3 August while she brooded
five downy chicks which had just hatched.
On 4 August the female was
located 50 meters from the nest with four chicks. A 30-minute search
failed to locate the fifth chick. The brood was located once or twice
daily until 13 August with the remaining four marked chicks identified
at each observation.
I was absent from the study area on 14 and 15
August and when I located the brood on 16 August only two chicks remained.
These two chicks survived until 21 September at which time
one chick appeared to have sustained a leg injury. When the brood was
located the following day the injured chick was missing.
It was later
observed on a wintering site 4.7 km away on 4 December.
Choate (1963) estimated chick mortality between 35 and 44 percent during
the brood period and suggested the largest losses of chicks occurred
in the first two weeks after hatch.
Braun and Rogers (1971) presented
data showing decreases in average brood size during the two weeks following
hatch of 20 to 30 percent which is less than the 32 percent in this
study. Their data represented broods observed throughout Colorado between
1966 and 1969. For blue grouse Zwickel and Bendell (1967a) reported a
40 percent loss of chicks during the two weeks following hatch, while
Keppie (1975) reported a 36 percent loss of spruce grouse chicks in the
first 40 days following hatch.
It would appear that early mortality of
chicks is similar in all species of grouse studied.
Causes
Specific causes of mortality in most free-living animals, especially
birds, are difficult to document.
Most predators which inhabit the
alpine are capable of consuming an entire ptarmigan' chick and any
remains disappear rapidly.
Ptarmigan chicks less than two weeks of
age (when most mortality occurred) are small (less than 50 grams)
and possess a downy plumage which blends into the surroundings.
Mortality was documented for seven chicks in 1975. A raptor, most
likely a prairie falcon (Falco mexicanus) killed hen G74 within 24
hours after she hatched a clutch of six eggs. The entire brood was
assumed to have died, although only one dead chick was located.
The
hen apparently had been dusting on a pocket gopher (Thomomys talpoides)
mound when the raptor attacked.
The other instance of chick mortality
in 1975 involved an unbanded chick approximately three weeks of age.
The remains were found nearly a month after death and examination indicated mortality was likely caused by weasel (Mustela spp.) predation.
In 1976 five mortalities
were killed by vehicular

of juvenile ptarmigan were located.
Two chicks
traffic on Trail Ridge Road which traverses the

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length of the study area. They were located together on 29 August and
were approximately six weeks old.
One chick was found dead in the nest of female Y73 after she led her
brood from the nest on 18 July. At least four chicks had hatched prior
to 19:30 p.m. the previous evening, but it was not known if the dead
chick was one of these. Examination of the chick indicated its weight
and carpal measurement were comparable to other chicks examined on day
of hatch.
It is possible that death occurred by suffocation when the
chick was brooded.
In nine other nests examined after hatch no mortality
was observed.
One juvenile ptarmigan (P64) was found dead less than four days after a
radio transmitter package had been attached.
Although death was due to
raptor predation it may have been influenced by the observer.
P64 was
captured at dusk on 8 October approximately one kilometer east of Tundra
Curves.
It was carried, along with two other juvenile ptarmigan, to
a vehicle at Tundra Curves where transmitters were attached.
The birds
were released singly from the vehicle.
Two birds flew towards the point
of capture while P64 flew in the opposite direction.
On 9 October P64
was located 300 m west from the point of release.
He was alone, but
was observed feeding and seemed to be in good condition.
I next searched
the area on 12 October and found the remains of P64. Apparently P64 had
attempted to rejoin flock as he had moved east to within 500 m of where
he had been captured.
It is conceivable that if he hadn't been separated
from his original flock he would have survived.
None of the other eight
juveniles fitted with transmitters died while I tracked them in October
and November.
The remains of another juvenile (unbanded) was found during a search
of alpine habitat three kilometers west of the study area. Death was
attributed to raptor predation and may have occurred several days prior
to its discovery on 8 November.
Examination of primary feathers indicated
that it was probably a female.
Excluding the chick which failed to leave the nest and .the two road kills,
all mortalities (both years) of juveniles were attributed to predation.
All but one were due to raptor predation.
Although raptors, especially
the prairie falcon, are common on the alpine during the summer and fall
(Braun and Rogers 1971, Marti and Braun 1975) the proportion of predation
attributed to them may be biased.
Fresh raptor kills are often easy to
locate as feathers are widely scattered at the kill site. Mammalian predation is more difficult to detect, thus its occurrence may be under represented.
Possible causes of mortality were documented in two other chicks.
On
14 July 1976 I located female G84 with a brood of six chicks approximately
six days old. All chicks were captured, banded, weighed and measured
(carpal length).
Five chicks had weights between 26 and 28 grams with the
remaining chick weighing 18 grams. The carpal measurements of the five
larger chicks ranged from 45 to 47 millimeters; the carpal measurement
of the smaller chick was 39 mm. It is possible that the chick was adopted
from another hen and was younger, but this was not likely based on the age

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of the brood. No adoption was observed in broods less than two weeks of
age. After release the smaller chick was observed and it appeared to have
difficulty in keeping up with the remainder of the brood.
Even though
female G84 was never sighted again (presumably depredated) the five larger
chicks were later observed as parts of two other broods.
The smaller chick
was never seen again. Although the smaller chick did not appear to be
injured, it appeared to be physically defective.
It may have had internal
injuries or genetic deformities which were not detected.
As previously mentioned, one chick of brood G97 appeared to have a leg
injury when seen on 21 September.
Both chicks in the brood had been
captured on 18 September and were in good condition.
The injured chick
was captured on 21 September and examined.
No injuries were noted, even
though the legs were carefully examined.
When released the chick continued
to limp but was able to follow the hen. I relocated the brood the next
day (22 September) but the injured chick was not seen. However, it was
later observed on a wintering area.
Weather may be a possible mortality factor, especially immediately after
hatch and again in fall and winter.
Observations of ptarmigan chicks
immediately after hatch indicated that inclement weather did not appear
to bother them. Although they were brooded a large proportion of the time,
they did feed periodically.
Theberge and West (1973) did not feel that
inclement weather decreased survival of young rock ptarmigan chicks.
Zwickel (1967) observed blue grouse chicks after hatch and also concluded
that weather did not seem to increase mortality.
Dispersal
Movements

to Summer Habitat

During the first few days following hatch broods often remained within a
few hundred meters of their nest site. Within a few weeks successful hens
with their broods began arriving at traditional summering areas where they
remained until brood dispersal or fall snowstorms.
Braun (1969) described
summer brood areas as being high, rocky and frequently windswept ridges.
Rocks over 30 cm diameter often comprised over 50 percent of the ground
cover, while late lying snowfields were present in protected areas.
The
vegetation was dominated by sedges (farex spp.) with Geum rossii, Polygonum
spp. and Trifolium spp. present in varying amounts.
These four groups of
plants provided the majority of food utilized by both hens and chicks
(May and Braun 1972, May 1975).
Movements between nest site and center of brood ranges for seven hens whose
nest sites were located (4 in 1975, 3 in 1976) averaged 350 m. Movements
between breeding territories and brood ranges for 22 other successful hens
(11 in 1975, 11 in 1976) ranged from zero to 3,300 m. No movements greater
than 600 m were observed in 1975 while the average movement to brood range
in 1976 was 1,052 m. Possibly the drier conditions in 1976 were responsible
for longer movements to suitable summer habitat.
Braun (1969) observed an
average movement of less than 600 m for successful hens, while Schmidt (1969)
documented one movement of 1,400 m in four days for one ~uccessfu1 hen.

�-122-

Movements

Within Brood Home Range

Once broods became established on suitable summer habitat their movements
became localized until fall dispersal or early snowstorms caused downslope
movements into more protected areas. Occasionally brood habitat became
unsuitable due to drying which then caused broods to seek areas where
moisture was present.
Four major brood concentration areas were located within the study area.
The concentration area at Toll Memorial was studied most intensively while
brood concentration areas at Medicine Bow Curve, Iceberg Lake and Sundance
Mountain were studied less. Movements within a brood concentration area
were recorded using extensive daytime observations (up to 10 consecutive
hours) and by plotting consecutive observations on a topographic map for
later measurement.
Radio telemetry locations were not used unless the
brood was sighted, since topography of the area prevented exact location
by triangulation of radio signals.
Individual broods were located an average of 7.2 times during the summer
in 1975 and 5.5 times in 1976 (6.1 overall).
Some broods were only observed once (many of these hens lost their broods early) while others were
observed up to 34 times. Brood home ranges were calculated for broods
observed a minimum of ten days since home range size tends to increase
with the number of observations (Odum and Kuenzler 1955). Brood home
range was calculated for 9 broods (3 in 1975, 6 in 1976) and averaged
15.4 hectares (range 3-32 ha). It is interesting to note that both
extremes in home range size occurred at the same area (Toll Memorial) and
that both hens involved adopted chicks from other hens by early August.
Movements within summering areas were estimated by measuring distances
between consecutive observations of marked broods.
In 1975 broods were
located an average of 252 m from the previous observation while in 1976
this distance was 211 m. For all broods observed more than ten times
during the summer (3 broods in 1975, 6 broods in 1976) the average distance
between consecutive locations averaged 144 m in 1975 and 161 m in 1976.
In order to obtain information on daily movements, selected broods were
intensively observed periodically throughout the summer.
These observations
indicated that young broods moved little, while broods over four weeks of
age moved longer distances.
Daily movements were often circular or elliptical routes which were similar from one day to the next. These routes
were traveled several times each day. Individual broods varied in their
daily movements, most likely in response to weather and the distribution
and availability of resources, especially food. Brood G84 often moved
800 to 1,600 m per day while broods BW5, BW16, G85 and G86 seldom traveled
over 400 m per day.
Brood Breakup
Brood breakup began gradually after chicks reached eight weeks of age.
Brood m~x~ng or brood shuffling occurred earlier due to death of brood
hens or because of close association of two hens on brood areas. Brood

�-123-

breakup was gradual until the first severe fall snowstorm which appeared to
increase its incidence.
Not all broods broke up in fall as at least three
hens (G34, G76 and BW27) maintained intact broods until late November 1975.
The incidence of brood breakup was reflected in banding records.
Prior to
1 October 1975, 59 juvenile ptarmigan had been banded and at time of banding
were associated with hens known or assumed to be their parent.
Between
1 October and 1 November 1975, 24 juveniles were banded of which 6 (25 percent)
were with hens assumed to be their parent.
During November an additional
12 chicks were banded and none was associated with a hen assumed to be its
parent.
Banding data for 1976 were similar.
Prior to 1 October, 88 chicks were
banded and only three were with a hen not assumed to be its parent.
Between
1 October and 18 December 1976, 33 chicks were banded and only one was with
a hen assumed to be its parent.
Braun (pers. comm.) suggested that juvenile
females dispersed greatest distances in fall. This was supported by banding
data from this study. Combined data from both years show that 51.3 percent
(60 of 117) of juveniles banded were females.
Sixty-one percent (22) of
the chicks banded between 1 October and 30 November 1975 were females and
67 percent (22) of the chicks banded between 1 October and 18 December 1976
were females.
Since most chicks which hatched on the study area were banded
prior to 1 October, chicks banded after that date were assumed. to have
dispersed onto the study area.
No specific causes of brood breakup were discovered.
Individual broods were
observed up to ten consecutive hours in September and early October 1975 in
an attempt to understand factors responsible for breakup.
Aggressive behaviors between hen and brood were not observed, nor was aggression shown
between chicks of the same brood or among chicks of different broods.
Many
hens continued to respond to chick distress calls until early October, long
after brood breakup began.
Since timing of brood breakup was variable among
broods there may be other behavioral or genetic factors responsible.
These
factors may be associated with weather (snowstorms) or other stimuli in
the environment.
Fall Dispersal
Movements of juvenile ptarmigan between brood breakup and establishment on
a wintering area may be one of three types:
(1) daily movements associated
with feeding and other maintenance activities, (2) dispersal movements,
and (3) migration to a wintering site. The latter two types of movement
are difficult to distinguish and it is likely that a series of dispersal
movements result in migration to the wintering area. Hoffman and Braun
(1975) recorded movements between breeding or summering areas and wintering
sites but did not examine movements within the fall period.
Also, their
data were on adults and yearlings, mainly females, and represented migration
rather than true dispersal.
In this study dispersal movements were defined as any movement of at
least one kilometer occurring during or after brood breakup.
The distance

�-124-

(1 km) is arbitrary but was selected since most fall use areas were separated
by a minimum of one kilometer.
Movements of less than one kilometer were
common and occurred during daily maintenance activities.
Also, movements
greater than one kilometer usually resulted in the individual abandoning
one flock and associating with another.

Few data concerning dispersal movements were obtained in 1975. Although
most chicks which hatched on the study area were banded it was difficult
to identify individuals without recapturing them. Recapture was difficult
since fall flocks often flushed before a capture attempt could be made.
Other birds, especially those which had been captured several times previously, were wary of the noose and were difficult to capture.
Limited
data indicated juveniles dispersed up to 3,900 m but all observations were
on the study area and no attempt was made to locate juveniles moving off
the study area.
Fall dispersal data were less biased in 1976 as most juveniles were identifiable without capture and alpine areas adjacent to Trail Ridge were searched.
Also, in mid-August 31 chicks outside the study area were banded.
Some
of these individuals later passed through the study area and provided important data on timing, direction and distance of dispersal.
Nine juveniles
were marked with miniature radio transmitters in October 1976 (Table 2)
which facilitated location of individuals dispersing off the study area.

Table 2.
in 1976.

Radio tracking

data for nine juvenile white-tailed

ptarmigan

Band
Number

Sex

Capture
Date

Days
Transmitting

P98

Male

8 October

21 days

transmitter

failure

P88

Female

8 October

31 days

transmitter

failure

P64

Male

8 October

4 days

killed by raptor

P67

Female

8 October

1/
31 days-

moved out of range

P82

Male

8 October

4 days

lost transmitter

P65

Female

9 October

42 days

transmitter

failure

P56

Female

9 October

30 days

transmitter

failure

P58

Female

22 October

1/
18 days-

moved out of range

P73

Male

23 October

15 days

transmitter

Cause of Transmission
Termination

.
-l/M"1n1mum
number of days before bird moved out of range.

failure

�-125-

Fall movements were recorded for 74 juvenile ptarmigan (32 males, 42
females) in 1976 (Table 3). There was no significant difference (P &gt; 0.05)
between males and females in average distance moved between consecutive
observations although females tended to move greater distances (Figure 2).
Fall dispersal movements (~1.0 km) were recorded for 60 juvenile ptarmigan
(25 males, 35 females) in 1976 with 53.3 percent having more than one
dispersal movement.
Dispersal movements (n = 68) of juvenile females
were significantly (P &lt; 0.05) longer than dispersal movements (n = 53)
of juvenile males.
The mean dispersal movement of juvenile females was
3,025 m while it was 2,306 m for juvenile males (Figure 3). When an
average dispersal distance was calculated for each individual with more
than one dispersal movement, juvenile females had significantly (P &lt; 0.05)
longer average dispersal movements than males (3,067 m and 2,354 m,
respectively) (Figure 4).
Distance of dispersal movements did not appear to be normally distributed;
this may reflect a genetic variability for dispersal or it may be due to
biased search efforts and uneven distribution of suitable habitats.
Females
comprised 56.8 percent (42 of 74) of all juveniles identified in the fall
and accounted for 56.2 percent (68 of 121) of all dispersal movements.
Thus the average number of dispersal movements between sexes was approximately equal with juvenile females dispersing greater distances than males.
Table 3. Fall movements of juvenile white-tailed
Mountain National Park, 1976.

ptarmigan

in Rocky

S.D.17

Number

Female
Mean!?

S.D

1519

1147

42

1792

1571

53

2306

1349

68

3025

1948

Individual average
dispersal

25

2354

1125

35

3067

1377

Summer to winter

6

3467

2039

9

5089

1999

Number

Male
Meanl}

32

Total dispersal

Movement

Individual

-lID'lstance

fall average

!}

e •

. .me ters.
ln

Most dispersal movements were along the southeast-northwest
axis of the
Trail Ridge study area (Figure 5) indicating that ptarmigan preferred
to move along areas of continuous alpine habitat.
Several exceptions
were noted. Most movements to the southwest were by individuals moving
from Fall River to the Divide Range between Mt. Ida and Sheep Rock.
This
route is characterized by short Krummholz or alpine.
Several juveniles

�-126-

-1--1--1-

(42 )

(32)

Males

Figure

2.

Females

Individual average fall movements of juvenile white -tailed
ptarmigan.
Vertical line,· range; horizontal line, mean;
vertical bar, standard deviation.
Sample sizes are in
parenthesis.

�-127-

10

9

4(68)
(53)

2

o~----------------------------------------Males

Figure

3.

Females

Dispersal movements of male and female juvenile white. tailed ptarmigan.
Vertical line, range; horizontal line,
mean; vertical bar, standard deviation.
Sample sizes are
in parenthesis.

�-128-

10

98
7

6
5
4
( 35)

3

2
1

o~--------------------------------------Males

Figure

4.

Females

Individual average dispersal movements
tailed ptarmigan.
Vertical line, range;
mean; vertical bar, standard deviation.
in parenthesis.

of juvenile whitehorizontal line,
Sample sizes are

�-129-

apparently flew north-northeast between Toll Memorial and Mt. Chapin and
three radio-marked ptarmigan flew southwest from Tombstone Ridge to Sprague
Pass. These latter birds had to fly three to four kilometers across Forest
Canyon to reach the nearest alpine. Although search efforts may be biased
it appeared that ptarmigan usually dispersed along areas of continuous
alpine habitat although they were capable of flying across several kilometers
of subalpine forest to reach other suitable alpine areas.
Summer to Winter Movements
The vegetative and environmental factors characteristic of white-tailed
ptarmigan wintering areas in Colorado have been described by Braun (1969,
1971), Braun and Schmidt (1971), Hoffman and Braun (1975), and Braun et al.
(1976). Segregation of sexes usually occurs in winter flocks of ptarmigan,
with males utilizing higher, more windswept areas, while females prefer
lower areas at or below treeline.
Abundance and availability of willow
is a prime consideration, as the buds and twig tips of willow comprise over
90 percent of the winter diet of ptarmigan in Colorado (May and Braun 1972).
This study was not designed to monitor winter populations of ptarmigan.
Due to lack of intensive effort and inaccessibility of the study area,
relatively few observations of ptarmigan were made in winter.
In 1975,
16 juvenile ptarmigan were identified on wintering areas with 12 having
been previously marked (8 in summer, 4 in fall). Average distance moved
between summer habitat and winter sites for these 8 ptarmigan was 1.37 km.
Few conclusions can be based on these data since all but two observations
were made at the one wintering site (Sundance Mountain Basin) most accessible
in winter.
Also, four of the individuals observed belonged to the same
brood (G76) which remained intact until late November.
In 1976 movements between summer areas and wintering sites were documented for
15 marked juveniles (6 males, 9 females).
There was no significant difference
between sexes (Mann-Whitney U-test, T ~ 0.05) in distances moved, but this
may reflect the small sample and uneven search effort.
Juvenile males
migrated an average of 3,467 m to wintering areas, while juvenile females
averaged 5,089 m (Figure 6). The combined mean movement was 4,440 m (median
4,700 m). Only one male (P87) migrated over 5,000 m, whereas six of nine
females exceeded that distance.
One possible source of error lies in the accuracy in sexing juveniles.
No
field technique has been devised to sex juvenile ptarmigan externally with
100 percent accuracy.
The best technique relies on measurement of carpal
and primary feather length along with pigmentation of the eyestripe.
Once
primaries are completely grown (18 weeks) this technique is approximately
95 percent reliable.
Most chicks were sexed at initial capture and with
every recapture, however, few marked chicks were captured in winter after
primaries were fully grown.
It is suspected that migration distances of juveniles, especially juvenile
females, were grossly underestimated.
Alpine areas greater than 10 km
from the study area were not searched although fall movements exceeding
10 km have been documented for juvenile females (Hoffman 1974). Little is

�-l30-

N

s
Figure 5.

Distance and direction of dispersal rnovernents of juvenile
white-tailed pta rrnigan, Open circles represent rrra Ies;
closed circles. Ierna le s , Each concentric circle represents 1.0 KIn.

�-131-

o~------~-----------------------------Females
Males

Figure

6. Movements

of juvenile white-tailed
ptarmigan from
summering areas to wintering sites.
Vertical line, range;
horizontal line, mean; vertical bar, standard deviation.
Sample sizes

are in parenthesis.

�-132-

known concerning movements of juveniles in winter and it is possible that
they may move between several winter sites. This was indicated by one
juvenile female, P8S, which was observed below Gore Turnout on 4 December
1976, and reobserved on 31 January 1977 in the Sundance Mountain Basin,
7.7 km east-southeast.
The winter of 1976-77 was extremely dry and lack
of adequate snow may have been a factor in this movement.

DISCUSSION
Mortality
Mortality is often described as loss to a population.
Mortality is defined
here to include prehatch mortality (nest predation) as well as posthatch
mortality, since both are important as potential factors influencing recruitment to a population.
Since all female ptarmigan (yearlings and adults) are assumed to nest, the
initial loss of potential recruitment occurs when a female fails to incubate
a clutch to completion.
The major factor is predation, as 52.4 percent of
all nests located during this study were destroyed by predators.
Renesting
compensates for some of this loss as up to 30 percent of hens unsuccessful
in their initial nesting attempt will renest. Adverse weather may affect
nesting success by causing desertion, but it is more likely that late
snowstorms increase nest predation.
Another loss to production occurs when some eggs in successful nests fail
to hatch.
Fertility was found to be quite high (94.5 percent) as expected
since white-tailed ptarmigan are monogamous and the male remains on the
territory with the female until late in incubation.
Not all fertile eggs
hatch but observer interference may have caused some eggs to chill or
become cracked.
Overall, 80 to 90 percent of the eggs in successful nests
hatch. Hypothetical production is summarized in Table 4.
Specific causes of mortality in chicks during their initial two weeks of
life were difficult to document.
Either extrinsic or intrinsic factors
may be responsible.
Some possible extrinsic factors are weather, predation,
food shortage or diseases (including parasites).
Observation of young broods
indicate they are not directly affected by inclement weather, nor did
excessive brooding times appear to cause starvation.
It does not appear
that food shortage is a problem due to low densities of ptarmigan and the
fact that broods are dispersed after hatching.
There are few data concerning food habits of young (less than two weeks of age) grouse, but
possible food items on the alpine (insects and plant material) do not
appear to be lacking.
Ptarmigan generally appear to be free of deleterious diseases or parasites (Haskins 1969, Stabler et al. 1974) although
young chicks have not been sufficiently tested. Predators, both mammalian
(weasels and coyotes, Canis latrans) and avian (prairie falcon), are
common on the alpine but there is an array of prey items. Zwickel (1965)
hatched blue grouse eggs and raised the chicks in captivity and found
that mortality of aviary chicks was equal to that of wild chicks which
indicated that extrinsic mortality factors may be unimportant.
Theberge
(1971) noted high mortality (&gt; 40 percent) in rock ptarmigan chicks in

�-133-

both wild and various aviary situations.
Since successful hens move their broods to suitable summer habitat within
two weeks of hatch it is possible that mortality occurs during this movement
as young chicks may have difficulty in traveling long distances.
However,
when proportion of chicks lost was compared to distance moved no correlation
was detected.
If intrinsic factors are responsible they should be passed on through the
hen. These factors are likely behavioral rather than physical attributes
as only one chick of over 50 handled soon after hatch was physically inferior
(using weight and size as criteria).
If mortality is due to behavioral
differences (responses to the hen) it seems likely that most mortality
would be clumped within the first two weeks rather than being distributed
over the entire summer.

Table 4. Hypothetical
100 females.

production

of white-tailed

ptarmigan

chicks per

Calculations

Change

Net Result

Initial laying

100 hens x 5.6 eggs

+560

560

Nest predation

100 hens x -60%·!/ x 5.6 eggs

-336

224

Renesting

60 hens x 30%~1 x 4.0 eggs

+72

296

Egg infertility

296 x -2%

-6

290

Embryonic mortality

290 x -6%

-17

273

Chick mortality
1 September

273 x -32%

-87

186

Situation

to

I
...
1 nests.
-lIE· stlmate d pre dat
atl0n
rate on lnltla

21

- Percent of unsuccessful

females renesting

successfully.

Individual hens occasionally lost their entire brood but the usual occurrence was for each hen to lose approximately one-third of her brood.
If
intrinsic factors are responsible it would be expected that unfit chicks
would quickly be lost from the population and each hen would possess a
characteristic brood survival rate. Twenty-four hens identified in 1975
were reobserved in 1976 with 75 percent (18) being observed during the
brood season each year. Five were successful both years, five were unsuccessful each year and eight were successful only one year (Table 5).
Data for the five hens which were successful both years show that brood
size for each hen differed between years (Table 6), although differences

�-134-

Table 5. Nesting success of white-tailed
both years (1975 and 1976).

Hen

1975

ptarmigan

hens observed

1976

BWl

Successful

Successful

BW5

Successful

Successful

BW26

Unknown

Unsuccessful

BW27

Successful

Unsuccessful

BW36

Unsuccessful

Unsuccessful

BW46

Unknown

Unknown

BW53

Successful

Unsuccessful

BW54

Unsuccessful

Successful

G3

Successful

Unsuccessful

G16

Unsuccessful

Unsuccessful

G30

Unsuccessful

Successful

G37

Unsuccessful

Unsuccessful

G39

Unsuccessful

Unsuccessful

G49

Successful

Successful

G56

Successful

Unkown

G72

Unknown

Successful

G76

Successful

Unsuccessful

G84

Successful

Successful

G86

Successful

Unknown

G96

Successful

Unsuccessful

G97

Unsuccessful

Successful

G99

Successful

Successful

Yll

Unknown

Unknown

Y25

Unsuccessful

Unsuccessful

�-135-

Table 6. Comparative brood size of white-tailed
observed both years (1975 and 1976).

ptarmigan

hens

Brood Siz~/
Hen

1975

1976

BWI

6

1

BW5

2

3

G49

4

2

G84

4

5

G99

2

5

1/
- Brood size in late August.

were great in only two hens (BWl, G99). Intrinsic factors affecting
juvenile mortality may be caused by annual variations in quality of vegetation
consumed by females prior to laying.
Quality of vegetation depends on
several factors including weather.
Thus it is possible that annual variations
in winter and spring weather may ultimately determine chick survival (Jenkins
et al. 1963).
Mortality in older chicks was uncommon until dispersal.
During dispersal
at least two juveniles died due to predation.
No other juvenile mortality
was documented so it is difficult to estimate rate or causes of mortality
during fall dispersal.
However, my first hypothesis, that mortality of
juveniles is predator-dependent
to 1 October and weather-dependent
from
1 October to 1 November, is not supported by data cQllected during this study.
Dispersal
Although animal movements are relatively easy to document, little is known
about the effects of movement, especially dispersal, on populations, and
less is known about mechanisms responsible for dispersal movements.
Dispersal
is important for understanding population regulation since it is a major
factor, along with recruitment and mortality, in determining population size.
Keppie (1975) studied a non-migratory population of spruce grouse and determined that dispersal occurred both in fall and in spring, and that spring
dispersal was a proximate factor regulating breeding densities.
Dispersal
also functions as a mechanism of gene flow in a population and dispersal of
juvenile females may be responsible for retaining genetic variability within
a population.

�-136-

No specific factors causing brood breakup or dispersal were identified in
this study. Brood breakup occurred gradually once chicks reached eight
weeks of age and preceded dispersal.
This same sequence of fall behavior
has been observed in juvenile ruffed grouse (Godfrey and Marshall 1969).
Intensive observations of broods during brood breakup and dispersal failed
to support my second hypothesis that agression among chicks or between hen
and brood was responsible for brood breakup.
Weather may facilitate brood
breakup and dispersal since fall snowstorms cause flocking in protected
areas.
Snow cover also affords protection to dispersing individuals.
Other behavioral factors may be involved in brood breakup since all chicks
within a brood usually abandon the hen (or vice versa) wihtin a few days.
However, there was great variation in timing of brood breakup among different
hens on the same area. Although age of chick was one factor in brood breakup,
other factors, thought to be intrinsic, may have been involved and are not
understood.
The incidence of dispersal as reflected in banding records and observations
of juvenile ptarmigan indicates that all dispersing individuals (n = 89)
were a minimum of eight weeks of age with most being 10 to 16 weeks of age.
Thus I accept my third hypothesis that timing of dispersal is age dependent
and does not occur until chicks are at least eight weeks of age.
I also fail to reject my fourth hypothesis, that direction of dispersal
is non-random.
Data presented indicate that dispersing juveniles moved
along continuous alpine habitat regardless of compass direction.
Data
presented in Figure 5 indicate dispersal movements exceeding 3.0 km
were more random with respect to direction.
Dispersal movements were
limited in the northeast quadrat but this may reflect uneven search effort.

SUMMARY
Intensive studies of timing and causes of mortality and fall dispersal
in juvenile white-tailed ptarmigan were carried out along Trail Ridge
Road and adjacent alpine areas in Rocky Mountain National Park, Colorado.
Data were collected from May 1975 to January 1977.
Observations of 49 individual broods showed a 32 percent loss of chicks
during the initial two weeks following hatch with little additional mortality
observed until brood breakup.
Observed mortalities were primarily due to
predation, most often raptors.
Road kills and injuries accounted for
additional known mortalities.
Two hundred eighty-five brood observations made between hatch and brood
breakup indicated broods were relatively sedentary on summer range. Daily
movements of broods ranged from 400 to 1,600 m within a three to 32 hectare
home range. Drying of alpine vegetation in late summer resulted in increased
brood movements.
Intensive observations of individual broods during brood breakup in September
and October indicated that brood breakup was gradual once chicks reached

�-137-

eight weeks of age. There was no evidence that aggression between hen and
chicks or among chicks caused breakup of broods.
Dispersal followed brood
breakup and was documented for 35 juvenile females and 25 juvenile males.
Juvenile females dispersed significantly (P &lt; 0.05) greater distances than
juvenile males.
Dispersing individuals moved along continuous alpine habitat
rather than dispersing in random directions.
Mortality of dispersing juveniles was light with two instances of raptor mortality being documented.
Movements between brood range and wintering areas were documented for nine
juvenile females and six juvenile males in 1976. Juvenile females averaged
longer movements to wintering sites than juvenile males (3,467 versus 5,089 m).
One juvenile female moved 7.7 km between December 1976 and January 1977
indicating that some juvenile ptarmigan are mobile throughout winter.

LITERATURE

CITED

Bowman, T. J., and R. J. Robel.
1977. Brood break-up, dispersal, mobility,
and mortality of juvenile prairie chickens.
J. Wildl. Manage. 41(1):
27-34.
Braun, C. E. 1969. Population dynamics, habitat,
tailed ptarmigan in Colorado.
Ph.D. Thesis.
Fort Collins.
189 pp.

and movements of whiteColorado State University,

1971. Habitat requirements of Colorado white-tailed
West. Assoc. State Game and Fish Comms. 51:284-292.

ptarmigan.

Proc.

_____ , and G. E. Rogers.
1971. The white-tailed ptarmigan in Colorado.
Colorado Div. Game, Fish and Parks Tech. Publ. 27. 80 pp.
_____ , and R. K. Schmidt, Jr. 1971. Effects of snow and wind on wintering
populations of white-tailed ptarmigan in Colorado.
Pages 238-250 in
A. O. Haugen, ed. Proc. Snow and Ice Symposium.
Iowa Coop. Wildl-.Res. Unit, Iowa State University, Ames.
280 pp.
_____ , R. W. Hoffman, and G. E. Rogers.
1976. Wintering areas and winter
ecology of white-tailed ptarmigan in Colorado.
Colorado Div. Wildl.
Spec. Rept. No. 38. 38 pp.
_____ , R. K. Schmidt, and G. E. Rogers.
tailed ptarmigan with tape recorded

1973. Census of Colorado
J. Wildl. Manage.
calls.

Bray, O. E., and G. W. Corner.
1972. A tail clip for attaching
to birds.
J. Wildl. Manage. 36(2):640-642.
Bump, G., R. W. Darrow, F. C. Edminster, and W. P. Crissey.
ruffed grouse;
life history, propagation, management.
Dept., Hilling Press, Inc., Buffalo.
915 pp.
Chambers, R. E., and W. M. Sharp.
population of ruffed grouse.

white37(1):90-93.

transmitters

1947. The
New York Conserve

1958. Movement and dispersal within
J. Wildl. Manage. 22(3):231-239.

a

�-138-

Choate, T. S. 1960. Observations on the reproductive activities of
white-tailed ptarmigan (Lagopus leucurus) in Glacier Park, Montana.
M.S. Thesis, University of Montana, Missoula.
113 pp.
1963. Ecology and population dynamics of white-tailed ptarmigan
(Lagopus leucurus) in Glacier Park, Montana.
Ph.D. Thesis. University
of Montana, Missoula.
205 pp.
Eddleman, L. E. 1967. A study of the phyto-edaphic relationships in
alpine tundra of northern Colorado.
Ph.D. Thesis.
Colorado State
University, Fort Collins.
148 pp.
Ellison, L. N. 1973. Seasonal social organization
spruce grouse.
Condor 75(4):375-385.

and movements

of

Godfrey, G. A. 1975. Home range characteristics of ruffed grouse broods
in Minnesota.
J. Wildl. Manage.
39(2):287-298.

----- , and W. H. Marshall.
grouse.

1969. Brood-breakup
J. Wildl. Manage. 33(3):609-620.

and dispersal of ruffed

Gullion, G. W., and W. H. Marshall.
1968. Survival of ruffed grouse in
a boreal forest. Living Bird 7:117-167.
Hale, J. B., and R. S. Dorney.
1963. Seasonal movements of ruffed
grouse in Wisconsin.
J. Wildl. Manage. 27(4):648-656.
Haskins, A. G. 1969. Endoparasites of white-tailed ptarmigan (Lagopus
leucurus) from Colorado.
M.S. Thesis. Colorado State University,
Fort Collins.
147 pp.
Hoffman, R. W. 1974. Characteristics and migration of wintering populations
of Colorado white-tailed ptarmigan.
M.S. Thesis.
Colorado State
University, Fort Collins.
58 pp.

----- , and C. E. Braun.
white-tailed

1975. Migration
ptarmigan in Colorado.

of a wintering population of
J. Wildl. Manage. 39(3):485-490.

Jenkins, D., A. Watson, and G. R. Miller.
1963. Population studies of
red grouse in northeast Scotland.
J. Animal Ecol. 32(2):317-376.
Judson, A. 1965. The weather and climate of a high mountain pass in
the Colorado Rockies. U.S. Forest Servo Res. Paper RM-16.
28 pp.
Keppie, D. M. 1975. Dispersal, overwinter mortality, and population
size of spruce grouse (Canachites canadensis franklinii).
Ph.D.
Thesis.
University of Alberta, Edmonton.
121 pp.
Marr, J. W. 1961. Ecosystems of the east slope of the Front Range in
Colorado.
University of Colorado Studies, Ser. BioI. 8. 134 pp.
1967. Data on mountain environments.
I. Front Range, Colorado,
sixteen sites, 1952-53. University of Colorado Studies, Ser. BioI. 27.
110 pp.

�-139-

Marti, C. D., and C. E. Braun.
1975. Use of tundra habitats by prairie
falcons in Colorado.
Condor 77(2):213-214.
May, T. A. 1975. Physiological ecology of white-tailed ptarmigan in
Colorado.
Ph.D. Thesis, University of Colorado, Boulder.
311 pp.
_____ , and C. E. Braun.
1972. Seasonal foods of adult white-tailed
ptarmigan in Colorado.
J. Wildl. Manage. 36(4):1180-1186.
Odum, E. P., and E. J. Kuenzler.
1955. Measurement
home range size in birds. Auk 72(1):128-137.

of territory and

Quam, L. O. 1938. The morphology of landscape of the Estes Park area,
Colorado.
Ph.D. Thesis, Clark University, Worcester, Massachusetts.
Robel, R. J., J. N. Briggs, J. J. Cebula, N. J. Silvy, C. E. Viers, and
P. G. Watt.
1970. Greater prairie chicken ranges, movements, and
habitat usage in Kansas.
J. Wildl. Manage. 34(2):286-306.
Schmidt, R. K., Jr. 1969. Behavior of white-tailed ptarmigan in Colorado.
M.S. Thesis.
Colorado State University, Fort Collins.
174 pp.
Stabler, R. M., N. J. Kitzmiller, and C. E. Braun.
1974. Hematozoa
Colorado birds.
IV. Galliformes.
J. Parasitol. 60(3):536-537.
Stiven, A. E. 1969. Food energy available
grouse chick. Ecology 42(3):547-553.

from

for and required by the blue

Theberge, J. B. 1971. Population fluctuations and changes in the quality
of rock ptarmigan in Alaska. Ph.D. Thesis. University of British
Columbia, Vancouver.
186 pp.
_____ , and G. C. West. 1973. Significance of brooding to the energy
demands of Alaskan rock ptarmigan chicks. Arctic 26(2}:138-l48.
Wallestad, R. O. 1971.
in central Montana.

Summer movements and habitat use by sage grouse
J. Wildl. Manage. 35(1) :129-136.

Weimer, R. J., and J. D. Haun (eds). 1960. Guide to the geology of
Colorado.
Geol. Soc. Am., Rocky Mountain Assoc. Geol., Colorado
Sci. Soc., Denver.
310 pp.
Willard, B. E. 1963. Phytosociology of the alpine tundra of Trail
Ph.D. Thesis.
Ridge, Rocky Mountain National Park, Colorado.
University of Colorado, Boulder.
245 pp.
W'1ng, L ., J • B eer, an d W . T'd
1 yman. 1944 .
of blue grouse. Auk 61(3):426-444.

Brood habits and growth

Zwickel, F. C. 1965. Early mortality and the numbers of blue grouse.
Ph.D. Thesis. University of British Columbia, Vancouver.
153 pp.

165 pp.

�-140-

1967. Some observations of weather and brood behavior
grouse.
J. Wildl. Manage. 31(3):563-568.

in blue

--- , and J. F. Bendell.
of numbers

1967a. Early mortality and the regulation
in blue grouse.
Can. J. Zool. 45(4):817-851.

____

, and
Manage.

1967b. A snare for capturing
31(1):202-204.

blue grouse.

J. Wildl.

___

, I. O. Buss, and J. H. Brigham.
1968. Autumn movements
grouse and their relevance to populations and management.
Manage. 32(3):456-468.

of blue
J. Wildl.

Prepared

by _~~_~~~~~~.~~~~~.~~~~~
_
Kenneth M. Giesen
Graduate Research Assistant

Approved

by

_:---!:a:::::..::...:::~~·_J._.---,-g=~=
Clait E. Braun
Wildlife Researcher

_

�April,

-141-

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Game Bird Survey

W-37-R-30

Work Plan No.

17

Job Title Evaluation

of White-tailed

Period

Covered:

Personnel:

1977

Job No.
Ptarmigan

6

---------------------------Introductions

to Pikes Peak

April 1, 1976 to March 31, 1977

George Bock, Clait Braun, Frank Colley, Marcus Elkins, Howard
Funk, Kenneth Giesen, Richard Hoffman, James Kitzmiller, Thomas
Lytle, James Morris, and Walter Larrick.

ABSTRACT
Division of Wildlife personnel concluded after 3 days of intensive searching
in May 1974 that no breeding population of white-tailed ptarmigan (Lagopus
leucurus) ,was present on Pikes Peak, but preliminary habitat investigations
suggested the area could potentially support a small population.
The initial
work of trapping and transplanting was completed in 1975~ Six pairs were
released in June followed by another release of 9 males, 7 females, and 12
chicks in September.
Both a spring breeding census (May) and summer brood
survey (August) were conducted in 1976. The breeding population consisted
of 7 pairs and 4 unmated males.
One bird (I-female) was unbanded indicating
this bird either moved to the area from elsewhere or represents production
from one of the spring released hens.
Spring released birds exhibited a
high fidelity to their release sites (maximum movement = 1.0 km) while fall
released birds dispersed the greatest distances (maximum movement = 3.4 km).
Overall survival was 50 percent.
As expected, chicks had the lowest survival
rate (25%) and males the highest (80%), but fewer ~emales survived (38%)
than expected.
Because of the greater mobility of females and chicks and
the limited size of the Pikes Peak area, most losses of females and chicks
were contributed to dispersal off the area rather than mortality resulting
on the area. Only 3 of 7 hens (2 unsuc~essful and 1 successful hen with
3 chicks) observed on territories were accounted for during the brood census.
The other 4 hens were probably unsuccessful and went undetected.
Based on
the hens found, nesting success was 33 percent.
However, if the latter
assumption is true, then nesting success was only 14 percent.
The poor production was attributed to a lack of suitable brood habitat.
Unless some
successful hens escaped detection, it is anticipated that recruitment into
the population will be inadequate to replace natural losses.

�-142-

RECOMMENDATIONS
1.

Continued efforts should be made in 1977 to conduct breeding
production surveys on Pikes Peak.

and

2.

It is recommended that no further transplants
to the Pikes Peak area.

3.

It is recommended that the data obtained in this study be turned
over to management after the 1977 field season for future monitoring
of the population.

4.

Information concerning the introduction of ptarmigan to Pikes Peak
should be made available in form of a popular article and special
report.

of ptarmigan be made

�-143-

EVALUATION OF WHITE-TAILED
PTARMIGAN INTRODUCTIONS TO PIKES PEAK
Richard W. Hoffman

Guidelines for introdu~ing
ptarmigan into unoccupied habitats have been
developed, but virtually nothing is known concerning what happens following introduction or whether the guidelines are adequate.
Removal studies
have provided information on the natural repopulation of previously
occupied areas (Braun 1975). However, because prior transplants were made
to other states (Oregon, California, and Utah), circumstances have not
permitted evaluation of introductions into new habitat.
The Pikes Peak
introduction represents the first opportunity within Colorado to conduct
followup studies.
Whether or not white-tailed ptarmigan ever occurred on Pikes Peak is uncertain.
Aiken and Warren (1914) and Knorr (1959), in studying the avifauna
of El Paso County, noted the absence of ptarmigan on Pikes Peak and reported
there is no authentic record of this alpine grouse occurring in the county.
No reference is made to ptarmigan on Pikes Peak in early distributional
records summarized by Sclater (1912) and more recently by Bailey and
Niedrach (1965). Occasional sightings of ptarmigan on Pikes Peak have been
reported, but most observations were by tourist types or novice bird watchers
and the validity of such sightings are questionable and/or unconfirmed.
Many of the sightings that were followed-up turned out to be blue grouse
(Dendragapus obscurus).
Although the total absence of this species on Pikes Peak cannot be completely
ruled out, there is strong evidence suggesting the absence of a breeding
population and only circumstantial evidence that ptarmigan might be a rare
visitor to the region.
This is not difficult to understand, as the Pikes
Peak area is a geographically and ecologically isolated alpine range
approximately 60 to 80 km (40 to 50 mi) from the nearest occupied range.
Even during periods of glaciation, access routes to the Pikes Peak region
probably never existed.

P.N.O. OBJECTIVES
Major objectives of this study are to: (1) evaluate the success or ,failure
of transplanted ptarmigan to establish a breeding population on Pikes Peak,
(2) ascertain if the distance of Pikes Peak from other occupied habitats
prevented males (but not females) from reaching the area and consequently
establishing a breeding population, (3) identify critical deficiencies in
the habitat (including minimum size) whigh might contribute to a failure
of ptarmigan to establish a breeding population, (4) evaluate the adequacy
of existing guidelines for ptarmigan introductions, possibly improve upon
them and/or develop additional guidelines, and (5) increase knowledge
available on the pioneering abilities of ptarmigan.

�-144-

SEGMENT OBJECTIVES
la.

Review pertinent literature and interview selected individuals
order to learn the past status of ptarmigan on Pikes Peak.

lb.

Ascertain present status of ptarmigan on Pikes Peak by field reconnaissance using developed techniques for locating and/or determining
the presence of ptarmigan during the peak period of breeding activity.

2.

Review and compare weather data and vegetative descriptions of the
Pikes Peak area with other selected alpine areas in Colorado inhabited
by ptarmigan.

3.

Search all suitable breeding, brood rearing, and summering habitat,
and keep records by date, location and vegetative type where ptarmigan
are observed.
Trap and band all unmarked birds.

4.

Estimate breeding population levels of ptarmigan
established census techniques.

5.

Estimate

nesting

6.

Compile

data, analyze

success and production
results,

METHODS

in

on Pikes Peak through

of ptarmigan

and prepare progress

on Pikes Peak.
report.

AND MATERIALS

Status of ptarmigan on Pikes Peak prior to the introduction was ascertained
by (1) field reconnaissance,
(2) review of avifauna records, and (3) personal
communications with Division of Wildlife personnel and other persons
acquainted with the area. Evaluation of the habitat in terms of the suitability
for ptarmigan was based on the habitat requirements of the species as presented by Braun (1971). Using data from earlier studies (Whitfield 1933;
Buechner, unpubl.) along with information collected in this study, vegetative
types were described according to procedures outlined by Braun (1969).
Plant nomenclature follows Weber (1972). Transplant procedures follow
established guidelines set forth by Braun (1975). Cl·imatological data were
obtained from Whitfield (1933), U.S. Department of Commerce, Weather Bureau
(1920-1934), Marr (1961), and Marr et ale (1968a and b).
Both a spring breeding census (May-June) and summer brood survey (AugustSeptember) were conducted in 1976. The techniques employed for locating
ptarmigan have already been described (Braun et ale 1973), along with efficient
trapping and identification procedures (Braun and Rogers 1971). Attempts
were made to identify all marked birds and to capture and band any unmarked
individuals.
Each sighting was recorded at the time on standardized observation cards, and locations were plotted on 7.5 minute U.S. Geological Survey
topographic maps. Most of the Pikes Peak region was traversed in order to
determine the distribution of the introduced birds and to delineate potential
seasonal use sites. All areas appearing suitable for use by ptarmigan were
outlined on U.S.G.S. topographic maps and subsequently planimetered to
estimate the amount of habitat available.

�-145-

DESCRIPTION

OF STUDY AREA

The Pikes Peak region lies in the east-central part of Colorado along
the Front Range of the Rocky Mountains at approximately 38030" north
0
latitude and 106 west longitude.
It is included within the Pike
National Forest in portions of El Paso and Teller counties (T14S, R68
and 69W, and T15S, R69W) and represents the eastern most extension of
alpine range in the ~tate. 2Total area that can be classified as alpine
includes about 392km (l~ mi ); however, the area under investigation
encompasses 23 km (9 mi ) (Fig. 1). Location of the Pikes Peak area in
relation to ptarmigan distribution and alpine range in Colorado is shown
in Figure 2.
Geologically, Pikes Peak is similar to much of the alpine area in Colorado.
The core of the front range in this area consists mainly of the Pikes Peak
granite, a natural, intrusive, igneous rock that was forced through the
overlying sedimentary rock during the Laramide Revolution.
Most of the
sedimentary rock has since been eroded away, leaving the granite exposed
(Koschmann 1956). Although the major rock type is granite, metamorphic
rock such as schist, quartzite, and gneiss are present.
With the exception
of quartzite, soils developing from these rocks are productive and can be
classified as coarse, loamy, and deep. However, some are occasionally
shallow to bedrock.
Soils termed by Nimlos and McConnell (1965) as alpine
meadow and alpine bog types occur frequently in wet, poorly drained sites.
Topography of the area is irregular, with elevations ranging from 3500 m
(11,500 ft) at treeline to 4300 m (14,109 ft) on the summit of Pikes Peak.
The study area lies on the southwest side of a ridge about 10 km (6 mi)
long. The southwest side of the ridge has a moderate slope, and merges
into a gently rolling upland.
This is in striking contrast to the northeast side where glacial action has created a rugged precipice with steep
walled cirques, rock heaps, and U-shaped valleys (Pearl 1964).

Vegetation
For descriptive purposes, the alpine region was divided into the low and
high alpine.
The low alpine contains many of the woody plants of the
alpine zone, and lies approximately between timberline and 3811 m (12,500
ft). The conspicuous dominant of the low alpine is Salix spp., but
krummholz communities alternately dominated by clumps of Picea engelmannii
or bushes of Salix spp. are major vegetative components of the low alpine.
Many of the drainage systems originating in the alpine have been blocked
by glacial action resulting in extensive wet areas dominated by dense
stands of Salix spp. These areas frequently extend from above treeline
into the subalpine.
Other plants locally abundant in the low alpine include: Pinus flexilis, Pinus aristata, Dryas octapetala, Pentaphylloides
floribunda, Kobresia bellardii, and Carex spp.
Further upslope, between the upper limits of the low alpine and the higher
talus slopes and ridges, are the alpine meadows which are transitory sites
between the low and high alpine.
Salix spp. extend into the lower portions
of the meadows, while rocky areas predominate along the upper portions.

�I

CO,_!

-~

'"I"':~f'

,_., ~~__ J.

NI,I"h.

•

I'
/

KEY
o Foil Release Site
t:: Spring Release Area .
x TI rritory ~Spring Released Male, Mated
Territory - Spring Released Male, Unmated
• Territory - Fall Released Mole, Mated
•••Territory~ Fall Released Mole, Unmated
c Summer Flock
'
Brood

*

iON

i

.~
I·

!
j,
'1

i

i
i~

'1'

r

A L

"""f~'
, ,

\

,

"

"

'

'-:.".", .'.~'.

,

~ "
~

'.,'

Sechett

Mountai(\

,:
"

-t
N

'\

.::.:.. -~ - --:-{:;,",'~~~

,?g

PIKES PEAK AREA
Study Area Boundary
Scale in Kilometers
o
112
1

~'~'~~/~····~~~~~-,.~j
./

~/

j'
~;

32

.•••

to'

/l1".'7T(rIir

./

-\ Nu.s

:\.

Fig. 1. Pikes Peak study area showing release sites and locations of
breeding territories, broods, and summer flocks.

/1"",1U:/:c:

'33,'

�-FORT COLLINS

~

-CRAIG

Ji

-DENVER

I

-GRAND
JUNCTION

I-'

+:""-l
I

~

o

- COLORADO SPRINGS
.PIKES PEAK
N

GUNNISON -

1)
SCALE IN KILOMETERS

o,

b
-DURANGO

fie?

TRINIDAD

Fig. 2. Location of the Pikes Peak area in relation to alpine range and
ptarmigan distribution in Colorado.

50

100

I

J

�-148Most of the area is characterized by low growing vegetation classified
as Kobresia-Carex-Geum and Kobresia-Carex alpine meadows. Deschampsia,
Polygonum, Potentilla, Pedicularis and Hymenoxys are some of the other
plants found throughout the meadows.
The high alpine includes the steep slopes and ridge tops above 3,963 m
(13,000 ft). The most obvious features of the high alpine are the abundance
of rocky areas and the exceedingly low growing vegetation. Vegetative
communities consist primarily of ~arex rock, Carex poa rock, and CarexGeum rock meadows, but Trifolium, Senecio, Kobresia, and Sedum are also
present in certain areas. Large boulder fields are common throughout the
area, especially at the higher elevations. Some have little or no vegetative cover. Similarly, many of the precipitous slopes in the northeast
portion of the area are devoid of vegetation.
Climate
Recent weather data were lacking for the study area. Some recent data were
available for subalpine sites near the study area, but the information was
incomplete. Consequently, reference is made to data collected from 1920 to
1934 by the U. S. Forest Service, Fremont Experiment Station, at a treeline
site approximately 4 km (2.5 mi) east of the study area (Table 1). Wind
velocities are excluded from Table 1 because the data were recorded in mph
of wind per day with no maximum, minimum or mean figures. Comparative data
were obtained from long term climate investigations conducted by Marr et al.
(1968a, 1968b) at Niwot Ridge (40003' N lat. and 105037'W long.) (Table 2).
This site lies 120 km (75 mi) north of the study area along the Front Range.
Although climatic data for the alpine are available from the studies by Marr
et al. 1968a, 1968b) these data are not directly comparable with the treeline data collected near the study area. Therefore, the data presented in
Table 2 also represent climatic conditions at treeline. It should be noted
that alpine air temperatures are usually 2 to 70C (4 to l30F) lower than
temperatures in the adjacent subalpine (Marr 1961). The amount of variation
is dependent upon elevational extremes, with minimum temperatures showing
the least difference (Marr 1961). Conversely, precipitation increases from
the subalpine to the alpine, but the precise increment is unknown (Marr 1961).
A detailed review of climatic studies in Colorado alpine areas and general
descriptive information is presented by Braun and Rogers (1971).
Climatic conditions in the alpine of Pikes Peak are similar to other alpine
regions in the state, and can best be described as continental. Frequent
extremes in wind velocities and temperature are common. Prevailing winds
are westerly. The alpine spring is short, cool, and wet. Upslope easterly
winds typically occur in the spring, and when accompanied by moisture, result
in heavy, wet snowfalls (Marr 1961). Summers are short and cool to warm,
with most precipitation occurring in the form of thunderstorms. Autumn is
short, clear, dry, and mostly cold; however, extended periods of warm weather
are not unusual. The winter is long, very cold, dry, and windy. Blizzards
are frequent, but much of the moisture occurring in winter is ineffective
because constant winds blow most of what fall as snow from exposed sites.

�Table 1. Climatological
data,
Weather Bureau. Climatological

Pikes Peak, 15-year average from 1920 to 1934 (U.S. Dept. Commerce,
data, Colorado annual summaries).

Month

Temperature,
Maximum

Air °c (oF)
Minimum

January

15 (58)

-24 (-11)

- 4 (24)

3.22 (1.27)

.05 ( .02)

1.01 ( .40)

February

16 (60)

-34 (-29)

- 4 (25)

5.28 (2.08)

.28 ( .11)

1.85 ( .73)

March

16 (61)

-26 (-14)

- 3 (27)

11.81 (4.65)

.99 ( .39)

4.19 (1.65)

April

18 (65)

-20 (- 3)

1 (34)

18.34 (7.22)

2.06 ( .81)

5.69 (2.24)

May

23 (73)

- 8 ( 17)

6 (43)

14 . 88 (5. 86)

2.26 ( .89)

7.14 (2.81)

June

27 (80)

- 2 ( 29)

12 (53)

20.75 (8.17)

.96 ( .38)

6.35 (2.50)

July

29 (83)

o ( 32)

15 (58)

19.63 (7.73)

4.29 (1.69)

10.18 (4.01)

Mean

Precipitati~~
cm Water (in)
Maximum
Minimum

Mean

I
I-'

~
~
I

August

27 (81)

- 1 ( 31)

13 (56)

18.77 (7.39)

2.82 (1.11)

9.96 (3.92)

September

26 (78)

-12 ( 11)

11 (51)

6.45 (2.54)

1.17 ( .46)

3.17 (1. 25)

October

23 (73)

-15 (

5)

5 (41)

12.70 (5.00)

.05 ( .02)

3.02 (1.19)

November

17 (62)

-26 (-14)

o (32)

8.69 (3.42)

.33 ( .13)

2.46 ( .97)

December

15 (58)

-30 (-22)

- 4 (25)

3.94 (1.55)

.10 ( .04)

1. 55 ( ~61)

Average

21 (69)

-16 (

4 (39)

12.04 (4.74)

1.28 (.50)

56.57 (22.28)

3)

�Table 2.
and b).

Climatological

data,

Niwot

Ridge,

11-year

average

from 1953 to 1964 (Marr

et

a1.

1968a

Month

TemEerature,
Maximum

o
0
Air
C (
Minimum

Mean

PreciEitation,
cm Water
Maximum
Minimum

January

12 (54)

-37

(-34)

- 7 (19)

8.38

(3.30)

1.27

( .50)

4.39

( 1. 73)

February

12 (53)

-32

(-25)

- 7 (19)

6.86

(2.70)

1.02

( .40)

4.42

( 1. 74)

March

12 (54)

-29

(-19)

- 6 (21)

9.02

(3.55)

1.27

( .50)

5.69

( 2.24)

April

17 (62)

-18

(-

1)

- 3 (27)

22.48

(8.85)

1.40

( .55)

7.77

( 3.06)

May

21 (69)

-16

(

4)

4 (39)

18.54

(7.30)

3.68

(1.45)

8.43

( 3.32)

June

25 (77)

- 7 ( 19)

9 (48)

9.40

(3.70)

.25

( .10)

5.46

( 2.15)

July

25 (77)

- 2 ( 28)

12 (53)

9.75

(3.84)

3.61

(1.42)

6.70

( 2.64)

6.83

( 2.69

F2

(in)
Mean

I
•.....
V1

0

August

25 (77)

- 3 ( 27)

11 (52)

18.11

(7.13)

September

25 (76)

-14

(

7)

8 (46)

13.16

(5.18)

.35

( .14)

5.03

( 1.98)

October

18 (65)

-16

(

4)

4 (39)

6.35

(2.50)

.51

( .20)

2.79

( 1.10)

November

13 (55)

-27

(-17)

- 3 (27)

7.75

(3.05)

2.41

( .95)

3.96

( 1. 56)

December

13 (55)

-26

(-15)

- 6 (22)

9.14

(3.60)

.38

( .15)

4.19

( 1.65)

Average

18 (64)

-19

(-

1 (34)

11.58

(4.56)

1.48

( .58)

65.66

(25.86)

2)

1. 65 ( .65)

I

�-151-

Upon examination of Tables 1 and 2, it is apparent that warmer, drier
conditions prevailed in the alpine regions of Pikes Peak as compared to
Niwot Ridge. Monthly maximum, minimum, and mean temperatures near the
study area averaged 30C (50F) warmer, while the average annual precipitation was 9.1 cm (3.6 in) less than the same estimate for Niwot Ridge.
Forty seven percent of the annual precipitation (56.57 cm, 22.28 in)
in the Pikes Peak area occurred from June to August (26.49 cm, 10.43 in)
with fall through late winter (Sept. to Feb.) being the driest period
(13.08 cm, 5.15 in). Only 29 percent of the annual precipitation
(65.66 cm, 25.86 in) at Niwot Ridge fell from June to August (19.00 cm,
7.48 in) whereas over 10 cm (4 in) more precipitation accumulated from
September through February (24.79 cm, 9.76 in). Spring was the wettest
season at Niwot Ridge.

RESULTS AND DISCUSSION
The initial work of trapping and transplanting ptarmigan to Pikes Peak
was completed in 1975. Ptarmigan were trapped from 5 different areas
in Colorado (Guanella Pass, Hagerman
Pass, Weston Pass, Loveland Pass,
and MOsquito Pass) to insure genetic diversity in the stock of transplanted birds.
Two transplants (spring and fall) involving a total of
40 birds were made. Six mated pairs were released in June followed by
another release of 9 males, 7 females, and 12 chicks in September.
All
birds were turned loose within 1-1/2 days of when they were trapped.
No
mortalities occurred as a result of transplant operations.
Release sites were selected based on (1) the spring and fall habitat requirements of the species (Braun 1971), (2) the proximity of other
seasonal use areas (fall and winter habitat) in relation to the release
sites, and (3) the accessibility to the release sites. Both the spring
and fall release sites were in the same general locality, but the fall
release site was at a slightly higher elevation.
During the spring
transplant, birds were released as mated pairs at approximately 180 m
(200 yd) intervals apart. The female was released first, immediately
followed by the male. No special precautions were taken in the fall
release except for releasing chicks in close proximity to a hen. The
remaining birds were turned loose in groups of 4 to 5 birds.
All birds
were marked prior to release and pertinent information about each bird
was recorded.

Breeding

Survey

Density of ptarmigan during the breeding season (late April to early July)
was ascertained by acoustical census (Braun et al. 1973).
It is believed
that none of the breeding pairs and less than 5 percent of the unmated
birds escaped detection.
All potential breeding habitat was searched,
but efforts were primarily concentrated around the release sites.
Results of the 1976 breeding survey are summarized in Table 3. The
breeding population consisted of 18 birds including 7 pairs and 4 unmated

�-152-

males.
Three additional males were located during the summer census that
went undetected in the spring. These males were probably unmated and possibly non~territorial.
No unmated hens were located, but one unbanded juvenile hen was captured in the spring of 1976. This female was not an introduced bird and either moved to the area from elsewhere or represents production in 1975 from one of the spring released hens.
Spring released birds set up territories during the first year of introduction (1975). There were no extensive movements away from the release
site to other potential breeding areas. Soon after release, the birds
spaced themselves out, established territories, and resumed breeding
activities.
No further attempt was made to monitor the population until
1976.
Since ptarmigan are monogamous, mate for life, and return to the same
territory year after year (Schmidt 1969), it was not unusual that the
spring released birds were breeding in the same area (release site) in
1976 where they had established territories in 1975. Of the 6 spring
released males, 5 were situated on territories within 1 km (.6 mi) of
their release site. The other male was non-territorial and his exact location during the breeding season was uncertain.
However, he was located
later in the summer .3 km (.2 mi) from where he was released.
Two spring
released hens survived until 1976 and both paired with their original
mates.
One of the spring released males paired with a fall released
female.
Even though spring and fall releases were made in the same general locality,
fall released birds did not exhibit the fidelity to the release site as
demonstrated by the spring released birds.
The nearest territory of a
fall released male in relation to its release site was 2.1 km (1.3 mi)
away. The outermost territorial males were over 3 km (2 mi) from the fall
release site. These birds were released after the breeding season, therefore, they did not establish territories until 1976. Because the spring
released birds had already occupied territories immediately surrounding
the release area, the fall released birds were forced to search elsewhere
for breeding habitat.
Figure 1 shows the location of breeding territories.
It is evident that
most territories (and release sites) were in the southern portion of the
study area, and that males established territories in close proximity to
another male.
There was just one territorial male isolated from any other
males. As a result of this attraction between males, and because the
spring released males had the first opportunity to establish territories,
the location of their territories appeared to at least partially influence
where the fall released males set up territories.
This is suggested by
the fact that some fall released males occupied territories in marginal
habitat near another male; whereas, seemingly better quality habitat,
absent of other males, went unoccupied.

�-153-

Table 3.

Breeding

season survey, Pikes Peak area, 1976.~/

Movement

No.

Age

Sex

Breeding

Status

Time of
Release

2321

2+

M

Unmated, non-territorial

Spring

2322

2+

F

Paired w/original

mate

Spring

.62

2323

2+

M

Territorial, paired w/
original mate

Spring

.62

2324

1+

M

Territorial, paired w/
original mate

Spring

.72

2325

1+

F

Paired w/original

Spring

.72

2028

2+

M

Unmated,

territorial

Spring

.99

2327

1+

M

Unmated,

territorial

Spring

.91

2330

1+

M

Territorial, paired w/fall
released female

Spring

.41

2476

2+

M

Territorial, paired w/fall
released female

Fall

2.09

2478

1+

F

Paired w/fall

Fall

2.09

2479

2+

M

Unmated,

territorial

Fall

2.55

2480

2+

F

Paired w/fall released male

Fall

2.14

2484

2+

M

Territorial, paired w/fall
released female

Fall

2.14

2489

1-

F

Paired w/fall released male

Fall

3.36

2490

1-

M

Unmated, non-territorial

Fall

2492

2+

M

Unmated, non-territorial

Fall

2493

1+

M

Territorial, paired w/fall
released female

Fall

3.36

2496

1-

M

Unmated,

Fall

3.20

2497

1+

F

Paired w/spring

Fall

.41

2499

1+

M

Territorial,
female l:../

paired w/new

Fall

3.20

2701

1-

F

Paired w/fall

released male

1/

Band

mate

released male

territorial
released male

Y

(km)

1/
- Excludes birds that were not located and believed

2/

- Movement

from release site to territory,

lost from the population.

excludes non-territorial

males.

3/
- This female was not introduced to the area and either represents production
in 1975 from one of the hens released in the spring, or else moved to the
area from elsewhere.

�-154-

As expected, juveniles and subadults moved the greatest distances.
Adult males are the most sedentary portion of the population (Hoffman
and Braun 1975). The location of their territories in relation to the
release site indicated they traversed the minimum distance necessary to
find a vacant slot where they could become established.
The habitat they
selected was not always the best available, but it was the closest, unoccupied, suitable area where two or more males could establish territories.
All potential breeding habitat was delineated on 7.5 minute U.S.G.S. topo2
2
graphic maps. Whereas th~ entire area encompassed 23.4 km (9.1 mi ),
only 27.6 percent (6.52km ) consisted of suitable breeding habitat.
Eighty percent (5.2 km ) of the habitat and consequently, 95 percent (20)
of the birds were in the sout~ern portion o~ the study area. At an
average density of 8 birds/km
(20 birds/mi ) (Braun 1969), this area
could presumably support a breeding population of about 25 pairs.

Survival
Survival data are based on the reobservation of banded birds following
one year of introduction.
All the introduced birds were banded prior to
release.
It was assumed in the calculations that any birds not reobserved
were dead. Nevertheless, some fraction of the population undoubtedly
escaped detection, but the portion was believed to be small «5%).
Survival rates for the spring and fall released birds are presented in
Tables 4 and 5. Comparative survival rates of the introduced population
and an unhunted population of white-tailed ptarmigan in Rocky Mountain
National Park (RMNP) are shown in Table 6. The RMNP data are from Braun
(1969). Data for the introduced population are based on combined survival
rates of the spring and fall released birds.
Except for adult females,
survival rates of the transplanted birds were similar to survival in a
natural population.

Table 4. Survival
to Pikes Peak.
Release

Period

Spring

Fall

1/1- birds

rates by age class of white-tailed

ptarmigan

introduced

Age 1/

No. Released

1-

7

57.1

2+

5

80.0

1-

12

25.0

2+

16

56.2

Survival

(%)

in the spring represent chicks hatched the previous summer (10+
months), whereas 1- birds in the fall are chicks of that year (7-8 weeks).
Adults (2+) and subadu1ts (1+) are grouped as adults (2+).

�-155-

Table 5. Survival
to Pikes Peak.

rates by sex of white-tailed

ptarmigan

introduced

Release Period

Sex 1./

Spring

M

6

100.0

F

6

33.3

M

9

66.6

F

7

43.0

Fall

(%)

Survival

No. Released

-1/ Sex of fall released birds represent adults (1+, 2+) only because
chicks (1-) could not be accurately sexed.

the

Table 6. Comparative survival rates of the introduced population and a
natural population of white-tailed ptarmigan in Rocky Mountain National
Park.
Survival Rates (%) 1/
Pikes Peak

Age and Sex

RMNP

Adults

67

62

Males

75

80

Females

65

38

65

57

Males

75

100

Females

55

25

Ch'1Ck s -3/

23

25

Annual Survival - All Birds

56

50

Subadults

~/

l/Survival rates for adults in the introduced population
survival of the spring and fall released birds.

represent

l/Spring released juveniles
years of age).

as subadults

3/

- Fall released juveniles
of age).

(10+ months)

are classified

(7-8 weeks) are classified

combined

(1-2

as chicks (0-1 year

�-156Males typically have the highest survival rate and juveniles the lowest
(Braun 1969). However, survival of adult females (38%) appeared to be
abnormally low compared to their survival in the natural population
(65).
Since the survival rates of the other age and sex classes COEr
pared favorably with the natural population (Table 6), the excessive
loss of females could not be entirely contributed to mortality on the
area.
It is believed that due to the high mobility of females (Hoffman
and Braun 1975) and the limited size of the area, more females were lost
from the introduced population as a result of dispersal off the area
than would normally occur in a natural population.
Females frequently
move long distances to seek out suitable wintering habitat and return
to breeding areas the following spring (Hoffman and Braun 1975). In
natural situations where the habitat is mostly continuous, only a minimum amount of unsuitable habitat needs to be traversed to reach wintering
grounds.
Any movement away from the Pikes Peak area almost certainly resulted in death as there was no other habitat within 60+ km (40+ mi).
Juvenile birds were divided into two categories based on their age: (1)
birds released in the spring were chicks hatched in 1974 (10+ months old
when released), and (2) those released in the fall were chicks hatched in
1975 (7-8 weeks old when released).
Mortality of ptarmigan varies with
age class and is highest in the first year of life (0-1 years) and
diminishes substantially during the second year (1-2 years) to a constant
level that is maintained in the older age classes (Braun 1969). These
differences are reflected in the survival rates of spring and fall released
juveniles which represent survival from 1 to 2 years of age (57%) and from
o to 1 year of age (25%), respectively. Braun (1969) reported similar
estimates on survival of subadults to adults (60%) and chicks to subadults (23%) in RMNP.
Further examination of the data showed that juvenile hens (10+ months old)
released in the spring suffered exceedingly high mortality as previously
described for adult females.
Survival of 3 juvenile males released in the
spring was 100 percent; whereas, only 1 of 4 (25%) juvenile hens released
in the spring of 1975 was reobserved in 1976. The same factor responsible
for the unnaturally high mortality of adult females was also believed to
affect survival of juvenile hens released in the spring.

Summer Survey
Summer brood investigations resulted in the location of 17 ptarmigan (11
males, 2 unsuccessful females, and 1 successful female with 3 chicks).
Only 3 of 7 hens observed on territories were accounted for during the
brood census. Status of the other 4 hens is uncertain, but it is doubtful
if they succumbed to mortality and more likely they were unsuccessful and
went undetected.
Hens with broods are easy to locate using tape recorded
calls and tend to remain sedentary on brooding areas, while unsuccessful
hens are more difficult to find because they wander widely and seldom
respond to recorded calls (Braun and Rogers 1971). Just 2 males observed
on territories could not be located during the summer census.

�-157-

Based on the actual number of hens observed (3), nesting success was
33 percent, but may be as low as 14 percent assuming those hens not
located were present but unsuccessful.
There was obviously a substantial decline in the introduced population the first year. Mortality and emigration contributed to a 50 percent reduction in the
population; whereas, production amounted to only a 12 percent increase
by late August.
There was essentially no immigration (possibly 1 bird)
into the population to offset emigration.
Unless some successful hens
escaped detection, it is anticipated that recruitment into the population
will be essentially zero.
The poor production was attributed to a lack of suitable brood habitat
rather than the failure of hens to successfully hatch a clutch of eggs.
Brood areas are characterized by an abundance of rock and low growing
vegetation, and generally are near late lying snowfields or wet areas
(Braun 1971). Such areas were limited in size and far between on the
study area with most areas searched in late summer being in a state of
desiccation.
Snow patches were absent on areas suitable for use by
ptarmigan.
Moist sites were mainly limited to areas adjacent to streams,
especially along the willow bottoms.
These areas are seldom used by
ptarmigan in summer.
Summering flocks of ptarmigan were found at two sites, but 88 percent
(15 birds) of the birds were occupying the rocky areas near the fall re- 2
lease zite (Fig. 1). The total area used by the birds was less than .3 km
(.1 mi).
Wet areas at this site were limited to a few small swales
scattered among the rocks. The swales were the only rem~ining source of
green vegetation, except for lower areas bordering streams.
The main
summer flock was composed of 10 males and 1 unsuccessful female. Although
the brood was using the same area, the hen and her chicks remained separate
from the main flock. Two additional birds (male and unsuccessful female)
were found on Sheep Mountain, about 2.8 km (1.7 mi) south of the fall
release site.
Changes in seasonal use patterns by ptarmigan as effected by grazing
(Braun 1971), human encroachment (Hoffman 1974), and weather (Braun 1969,
Braun and Schmidt 1971) have been documented.
If for some reason summer
use sites at higher elevations are not adequate, the birds may move downhill to rocky wet places within the krummholz.
Summering sites at Pikes
Peak were situated along the upper fringe of breeding areas. Thus,
summer flocks were formed primarily by the lateral movement of birds
from their territories to the summer concentration areas. Movements in
excess of 3 km (1.9 mi) were made by both males and females to reach the
main concentration area, but most birds (including the brood) traveled
less than 1 km (.62 mi). There were no signs of ptarmigan use in the
rocky krummholz areas or the higher rock fields, however, the krummho1z
areas could possibly be used as brood habitat.

Conclusions
Even though many of the environmental and physical features of the Pikes
Peak area are similar to other occupied ranges, it is questionable if the

�-158-

area can support a population of ptarmigan on a sustained basis. Possible
causes for the eventual failure of the introduction are (1) the limited
amount of available habitat, and (2) the isolation of the area from other
ptarmigan populations.
The cause-effect relationship is threefold.
First, due to the isolated
nature of the area, little if any immigration occurred into the population to replace losses occurring from emigration.
This was especially
noted for females and chicks.
Secondly, as a result of the limited amount
of breeding habitat (6.5 km2) the area can onlysuppotta
small population.
Braun (1975) reported that areas with less that 8 km2 of breeding habitat
probably cannot support a population on a sustained basis. Preliminary
results of this study indicated production in a small population without
immigration of females was inadequate to replace natural losses. Finally,
the lack of suitable brood areas in combination with the excessive loss of
females further contributed to poor production and subsequently, low
recruitment into the population.
If white-tailed ptarmigan are to be released in isolated mountain ranges
where they do not occur, it is imperative these areas be large enough to
maintain a self-sustaining population.
The minimum size of an area necessary to support a population is difficult to determine.
Braun (1975)
reported a minimum area of 8 km2 (3 mi2), but unless the habitat is ideal,
it is believed that such a small area cannot support a population when
emigration is occurring without immigration.
Prior introductions of ptarmigan to the Sierra Nevada's (Calif.) and the
Cascade Mountains (Oregon) have apparently been successful, while a recent
introduction has been made to the Uinta Mountains (Utah). These mountain
ranges are extensive, being in excess of 260 km2 (100 mi2); consequently,
emigration can occur to other portions of the same range. Eventually,
several, sUb-populations may become established with immigration
and emigration occurring between these smaller populations.
Individuals
unable to find vacant territories in the established areas then disperse
to adjacent unoccupied areas, thus allowing for expansion of the population.
To further insure a successful introduction, results of this study indicate
a spring release of mated pairs is better than a fall release.
If the
spring release is made early in the breeding season (early to mid-May)
there is a chance the introduced birds will successfully breed and produce
young the first year. This would eliminate having to introduce chicks in
the fall. Fall releases should include primarily females (1+, 2+) and a
few males (1+, 2+). It may be necessary to release additional females the
following year to insure adequate production.

�-159-

LITERATURE

CITED

Aiken, C. E. H., and E. R. Warren.
1914. The birds of E1 Paso County,
Colorado I. Colorado College Pub1. General Sere No. 74. 12(13):
455-496.
Bailey, A. M., and R. J. Niedrach.
1965. Birds of Colorado.
Museum Natural History, Vol. 1. 895 p ,

Denver

Braun, C. E. 1969. Population dynamics, habitat, and movements of
white-tailed ptarmigan in Colorado.
Ph.D. Dissertation.
Colorado
State Univ., Fort Collins.
189 p.
1971. Habitat requirements of Colorado white-tailed ptarmigan.
Proc. Western Assoc. State Game and Fish Coroms. 51:284-292.
1975. Experimental removal of a breeding population of whitetailed ptarmigan.
Colo. Div. Wi1d1., Game Res. Rept., Fed. Aid
Proj. W-37-R.
April. pp. 91-109.
______ , and G. E. Rogers.
1971. The white-tailed ptarmigan in Colorado.
Colo. Div. Game, Fish and Parks Tech. Pub1. No. 27. 80 p.
____ --,-,and R. K. Schmidt, Jr. 1971. Effects of snow and wind on
wintering populations of white-tailed ptarmigan in Colorado.
Pp.
238-250.
In A. O. Haugen, ed. Proceedings snow and ice symposium.
Iowa Cooperative Wi1d1. Res. Unit, Iowa State Univ., Ames.
280 p.
~
, and G. E. Rogers.
1973. Census of Colorado whitetailed ptarmigan with tape-recorded calls. J. Wi1d1. Manage.
37(1) :90-93.
Hoffman, R. W. 1974. Characteristics and migration of wintering populations of Colorado white-tailed ptarmigan.
M.S. Thesis.
Colorado
State Univ., Fort Collins.
58 p.
______ , and C. E. Braun.
1975. Migration of a wintering population of
white-tailed ptarmigan in Colorado.
J. Wild1. Manage. 39(3) :485-490.
Knorr, o. A. 1959. The birds of E1 Paso County, Colorado.
Studies, Sere BioI. 5. 48 p.

Univ. Colo.

Koschmann, A. H. 1956. Topographic map of Pikes Peak and vicinity,
Colorado.
U.S. Geo1. Survey map. U.S. Government Printing Office,
Wash., D.C.
Marr, J. W. 1961. Ecosystems of the east slope of the Front Range in
Colorado.
Univ. Colo. Studies, Sere Bio1. 8. 134 p.

�-160-

Marr, J. W., J. M. Clark, W. S. Osborn, and M. W. Paddock.
1968a.
Data on mountain environments III. Front Range, Colorado, four
climax regions, 1959-1964.
Univ. Colo. Studies, Sere BioI. 29.
179 p.
______ , A. W. Johnson, W. S. Osborn, and O. A. Knorr.
1968b. Data
on mountain environments. II. Front Range, Colorado, four climax
regions, 1953-1958.
Univ. Colo. Studies, Sere BioI. 28. 169 p.
u.S. Department of Commerce, Weather Bureau.
1920-1934.
Climatological
data. Annual Summaries, Colorado.
U.S. Government Printing
Office, Wash., D.C.
Nimlos, T. J., and R. C. McConnell.
Soil Sci. 99(5):310-321.
Pearl, R. M. 1964.
Colo.
36 p.

1965.

America's mountain.

Alpine soils in Montana.

Alan Swallow Publ. Co., Denver,

Sclater, W. L. 1912. A history of the birds of Colorado.
and Co., London.
576 p.

Witherby

Schmidt, R. K., Jr. 1969. Behavior of white-tailed ptarmigan in
Colorado.
M.S. Thesis.
Colorado State Univ., Fort Collins.
Weber, W. A.
Boulder.

1972. Rocky mountain
438 p.

flora.

Whitfield, C. J. 1933. The vegetation
Monographs 3(1):76-105.

Colorado Assoc.

174 p.

Univ. Press,

of the Pikes Peak region.

Ecol.

�April 1977

-161JOB FINAL REPORT

COLORADO

State of
Project

---------------------No.

W-37-R-30
20

Work Plan No.

1
Job No •.
Investigation of Population
S_t_a_t_u_s
__o_f
__B_o_b_w_h_l_·t_e
__Q~u_a_l_·l
__ l_·n
__ E_a
__s_t_e_r_n
__C_o_l_o_r_a_d_o

Job Title
Period

Covered:

Personnel:

Game Bird Survey

~~~--~---------------------

April 1, 1976 through March

Larry Crooks, William
Warren Snyder

T. Howard,

31, 1977
Michael

Fedorko

and

ABSTRACT

Field data were collected as·in previous years, analyses were completed
on the information obtained during this and the five previous segments
of study, and reports were written as follows:
Snyder, W. D. 1976. Bobwhite.
Colorado Outdoors 25(6):11-15.
(A semi-popular article covering the species, its habitat, distribution,
limiting factors, hunting and management in Colorado.)
Snyder, W. D. 1977. The bobwhite quail in Colorado.
A detailed summation
of study findings to be published as a technical bulletin under W-37-R-3l,
Work Plan 22, Job 1. The first draft has been completed and editing and
revisions are in progress.
Snyder, W. D. 1977. Census of bobwhite in Colorado.
A summation of
census findings and recommendations
for management to be published
as a Game Information leaflet under W-37-R-31, Work Plan 22, Job 1.
The first draft has been completed and editing and revisions are in
progress.

Prepared

by __~~
~
~~~~~-,~I~
__ ~~
Warren D.· sn~
Wildlife Researcher

_

_

��-163-

April 1977

JOB PROGRESS REPORT

State of

COLORADO
------------~--------

Project No.
Work Plan No.
Job Title

21

Job No.

1

Minimum Tillage Techniques for Establishing Shrubs in Clump Plantings

Period Covered:
Personnel:

Game Bird Survey

W-37-R-30

April 1, 1976 through March 31, 1977.

David C. Bowden, William T. Howard, Larry Crooks, John Corey,
and Warren Snyder.

ABSTRACT
Lack of adequate precipitation during the growing season on 1976 and throughout the year severely curtailed shrub growth but had no significant impact
on survival to fall. However, considerable die-back and winter kill was
noted during the winter of 1976-77 on the loam-soil Karg sites where
moisture conditions were especially deficient in 1976. Treatments had no
significant impact on shrub growth in 1976, but early spring 1977 observations indicated winter survival was better and die-back was less severe on
mulch treated sites. Some rabbit damage was inflicted to plums on the
Sonnenburg site and the three Karg sites in late fall 1976 before repellents
could be applied. All treatments, again, as in 1975, significantly reduced
annual weed occurrence in comparison to that on the cultivated controls.
Simazine, in the second year after application of this chemical herbicide,
lost some of its weed control effectiveness.
Some accumulative effect of
a second spring application of the chemical, casoron, was noted. This
herbicide and plastic mulch treatments were most effective in weed control
in 1976. Plant spacing, as a factor in survival and competition for
moisture was discussed.

��-165-

MINIMUM TILLAGE TECHNIQUES FOR
ESTABLISHING SHRUBS IN CLUMP PLANTINGS
Warren D. Snyder

P.N.O. OBJECTIVE
To evaluate ground cover techniques, pre-emergent herbicide techniques,
a combination of the two, and cultivation techniques for establishing shrubs
in clump plantings in eastern Colorado.

SEGMENT OBJECTIVES
1.

To apply pre-emergent herbicide treatments, mulch treatments, and a
combination of the two, to six of the seven plots within each planting
site. The remaining plot within each site will be retained as a
cultivated control.

2.

To measure and evaluate shrub survival, shrub growth and weed competition among treatments and soils in relation to weather variables.

METHODS AND MATERIALS
Methods and materials used in this study were summarized by Snyder (1975)
and Snyder (1976). Procedures listed there were followed very closely
during this work segment.
Rabbit repellent was placed only on wild plum on the loam and clay loam
sites in November 1976 to reduce further rabbit damage.
The Smith Meadow
site, within the clay loam riverbottom remained excluded from evaluations
and analyses because of biases which were discussed in Snyder (1976).

RESULTS AND DISCUSSION

Environmental

Measurements

Precipitation
A recording precipitation measuring gauge located at the Tamarack headquarters indicated 10.23 inches of precipitation was received during 1976
at that location.
A rain gauge was not located at the Karg site to the
northwest, however, observations indicated rainfall through the primary
growing season was up to several inches below the amount received at the
site of the rain gauge. Very little growth was observed at the Karg sites
as a consequence.
As shown in Table 1, 1976 precipitation was above that
received in 1974, below that recorded in 1975, and amounts received all
three years were below the average annual amount received in the study
region.
This lack of moisture was a primary factor affecting study evaluation to date.

�-166-

Table 1. Precipitation received at the Tamarack Management
quarters during 1974, 1975 and 1976.

Area head-

Month

1974

1975

1976

January

0.25

0.05

0.83

February

0.67

0.15

0.20

March

0.91

1.25

0.40

April

0.45

0.65

1.60

May

0.70

5.55

1.90

June

2.05

2.00

1.10

July

1.15

0.55

1.10

August

0.40

1.60

1.10

September

0.15

0.55

1.60

October

0.00

0.00

0.25

November

0.25

1.20

0.15

December

0.20

0.55

0.00

Annual Total

7.18

14.10

10.23

Average annual precipitation

in the study region approximates

15 to 16 inches.

Rabbit Damage
A mid-November 1975 snow induced considerable rabbit pruning and girdling
of wild plum on the Sonnenburg site and the three Karg sites. Follow up
December treatments with rabbit repellent prevented further damage.
Injuries inflicted in accompaniment with subsequent lack of rainfall had
significant impact on shrub survival and growth in 1976. Considerable
pruning of plums was also noted in early fallon the Sonnenburg site in
1976 with minor damage on the Karg locations.
Treatment with repellents
curtailed damage through the remainder of the winter.
Shrub Survival
Shrub Survival Within Sites Established

in 1974

Wild plums sustained approximately 14 percent mortality during their first
year after planting.
Hansen rose sustained about 25 percent mortality
during this interval, most of which was attributed to rodent damage in

�-167-

winter on the loamy sand sites. Little mortality was apparent from fall
1975 through fall 1976 as shown in Table 2. However, precipitation
deficiencies since late 1975 have greatly reduced plant vigor on most
locations.
Spring 1977 inspections, prior to writing this report, indicated considerable winter mortality due to desiccation on several sites,
but final late spring survival assessment will be forthcoming.
Root
sprouting was also occurring on numerous shrubs.

Table 2. Percentage shrub survival from fall 1975 through
the five sites established in April 1974.

Treatment

Fall 1975

fall 1976 on

Percentage Alive In
Spring 1976 ~I

Fall 1976

Wild Plum
Cultivated

91.88

91.88

90.63

Organic mulch

82.03

82.81

82.03

Plastic mulch

90.67

90.63

90.63

Simazine

84.38

83.75

83.13

Simazine-mulch

85.63

87.50

87.50

Eptam-Treflan

83.13

86.25

85.63

Casoron

83.13

84.37

81.88

Overall Average

85.80

86.74

85.89

Hansen Rose
Cultivated

66.25

71.88

71.88

Organic mulch

75.00

82.03

81.25

Plastic mulch

85.16

86.72

85.94

Simazine

74.38

79.38

77 .50

Simazine-Mulch

76.25

76.25

76.25

Eptam-Treflan

76.88

81.25

81.25

Casoron

73.13

75.63

74.63

Overall Average

75.00

78.69

78.03

II
- New root sprouts replaced previous mortalities to slightly
percentage survival on some plots during the spring of 1976.

increase

the

�-168-

Root sprouts from adjoining plants have replaced some of the original
mortalities, especially among Hansen rose, by early fall 1976. The slight
increase in plant survival in Table 2 from fall 1975 to fall 1976 reflects
this replacement by root sprouting.
Moisture deficiencies were most obvious in 1976 on the Karg sites which
had sustained the best growth in 1975. This, in combination with winter
rabbit damage and drought desiccation, severely impacted survival at the
time of this report.
Some leaf burning during the summer of 1976 was observed on the loam and
sandy loam sites. This was believed to have resulted from a combination
of moisture deficiency and pre-emergent herbicides, however, as yet the
impact of the herbicides could not be measured in reduced plant survival.
Shrub Survival Within Sites Established

in 1975

Survival among shrubs planted in April 1975 was better than that for shrubs
planted during 1974. One exception occurred on the loamy sand of the 1-76
site where Hansen rose sustained about 32 percent mortality by spring 1976
(Table 3). This primarily occurred in late summer and fall in 1975 due to
excessive drying of the light sandy soil. Dramatic treatment effect was
noted on that site as the plastic and organic mulch plots sustained much
higher survival than the other bare plots did. Survival was good to
excellent during the 1976 growing season on all sites in spite of lack of
moisture.
The Karg 3 site, where precipitation was especially deficient
through 1976, suffered considerable die-back and some winter mortality
through the fall and winter of 1976-77. Wild plums appeared to suffer the
most. Herbicide treatments had no discernible impact on survival through
the 1976 growing season.

Shrub Growth
Shrub Growth on Sites Established

in 1974

Treatment effect among sites and oils did not significantly influence
growth of either plum or Hansen rose in 1976 (Table 4) based on analyses
by David C. Bowden, Statistician.
Lack of discernible treatment effect
was believed to result primarily because little growth was sustained due
to lack of moisture.
Table 5 contrasts growth of shrubs in 1975 with that
in 1976 showing the much greater growth in 1975. Most significant was the
reduction in growth on the Karg 1 and Karg 2 sites in the loam soil type.
Shrubs on the Tamarack Meadow site attained much better growth in 1976 than
those on the other sites (Table 4). Appearance would indicate that Tamarack
Meadow shrubs had attained enough growth by 1976 to tap the relatively high
water table underlying this site. This additional moisture sustained them
through the dry summer and fall and little winter die-back or mortality was
evident.

�-169-

Table 3. Percentage shrub survival from fall 1975 through fall 1976 on
the three sites established in April 1975.

Treatment

Fall 1975

Percentage Alive In
Spring 1976 1./

Fall 1976

Wild Plum
Cultivated

97.92

95.83

93.38

Organic mulch

100.00

95.83

93.38

Plastic mulch

100.00

98.96

98.96

Simazine

96.88

94.79

93.38

Simazine mulch

97.92

95.83

93.38

Eptam-Treflan

100.00

93.38

92.71

Casoron

96.88

86.46

86.46

Overall Average

98.51

94.49

93.30

Hansen Rose
Cultivated

98.96

86.46

85.42

Organic mulch

100.00

92.71

91.67

Plastic mulch

98.96

98.96

98.96

Simazine

98.96

95.42

85.42

Simazine mulch

97.92

96.46

86.46

Eptam-Treflan

97.92

87.50

84.38

Casoron

98.96

81.25

80.21

Overall Average

98.81

88.39

87.50

1/

- New root sprouts replaced previous mortalities to slightly
percentage survival on some plots during the spring of 1976.

increase

the

�Table 4. Average shrub growth in inches per site and treatment during the 1976 growing season on the five sites
established in 1974 and the three sites eastab1ished in 1975. 1/

-Soil Type
Site

Cultivated

Organic
Mulch

Plastic
Mulch

Treatment
Simazine
Mulch
Simazine

EptamTref1an

Casoron

Site
Mean

Wild Plum
Loam Soil
Karg 1
Karg 2
Karg 3

3.27
9.57
8.91

4.48
5.24
13.66

3.63
4.06
17.39

6.29
7.13
15.31

7.48
9.65
17.38

10.93
6.46
15.09

8.58
11.92
16.28

6.35
7.62
14.85

Clay Loam
Tamarack Meadow
Sonnenburg

22.37
12.53

14.08
12.59

20.67
19.72

7.28
21.07

11.21
16.17

12.81
20.27

7.44
20.86

14.14
17.60

I

f-'
-..J

Loamy Sand
Red Lion
Sharp tail
I-76

7.16
7.21
10.62

13.88
2.32
12.61

12.12
11.21
16.25

11.21
5.67
10.10

9.35
4.50
10.96

8.07
7.03
9.77

Treatment Mean

10.27

11.00

12.90

10.76

11.01

11.39

9.48
6.9l
8.96
11.57

9.01
7.63
11.53
11.28

Hansen Rose
Loam
Karg 1
Karg 2
Karg 3

3.35
4.21
3.34

1.48
3.23
5.69

2.39
3.41
7.44

2.10
5.93
3.22

3.84
5.80
7.09

4.41
4.43
4.25

2.00
5.07
3.47

2.81
4.60
4.93

Clay Loam
Tamarack Meadow
Sonnenburg

9.92
3.50

7.32
4.66

7.18
4.84

5.52
4.06

6.25
7.29

4.86
5.73

7.32
5.06

6.89
4.99

4.46
5.53
8.77

5.40
3;53
8.79

0.83
6.14
9.50

4.68
4.52
7.57

6.08

4.98

4.55

5.06

Loamy Sand
7.69
4.97
5.91
7.67
Red Lion
1.92
6.08
5.62
4.48
Sharp tail
5.67
6.68
7.08
7.40
1-76
4.24
5.37
5.09
5.10
Treatment Mean
l/ The Karg 3, Sonnenburg and I-76 sites were established in 1975.

0
I

�-171-

Table 5. Comparisons of shrub growth in inches during the growing seasons
of 1975 and 1976 on the shrub clump sites.

Mean Growth-Plum

Mean Growth-Rose

1975

1976

1975

1976

Karg 1

19.73

6.35

14.16

2.81

Karg 2

19.38

7.62

13.71

4.60

Karg 3 1:./

6.89

14.85

8.87

4.93

21.58

14.14

14.06

6.89

3.48

17.60

2.35

4.99

Red Lion

13.84

9.01

13.27

4.68

Sharptail

6.79

7.63

5.-63

4.52

1-7J:./

5.04

11.53

3.18

7.57

Soil Type
Site

Loam Soil

Clay Loam
Tamarack Meadow

II

SonnenburgLoamy Sand

!I Sites established

in April 1975 contrasted

Shrub Growth on Sites Established

to those established

in 1974.

in 1975

Shrubs on these three sites, after recovering from planting shock in 1975,
made fair growth in 1976 (Table 4). They averaged much better growth than
sites established a year earlier.
This was probably because they were
younger, smaller, and therefore, in less competition for deficient moisture
through the 1976 growing season.
Treatments on these sites did not produce significantly different growth
effects in 1976 on either plum or rose species.
As shown in Table 4 considerable growth variance occurred among treatments and sites within soils.
Dr. David Bowden, Statistician, by analysis, found that these sites had
growth patterns differing from those of the 1974 established sites so the
two groups could not be incorporated for a more thorough analysis of treatment effect.
Winter Die-back
Winter die-back and mortality of shrubs has been previously mentioned.
During
the winter of 1975-76 this loss amounted to approximately half the growth
achieved during the preceding summer as shown in Figs. 1 and 2. Data from
sites established in both 1974 and 1975 were combined here for these comparisons. As shown in Fig. 2 Hansen rose growth in 1976 hardly compensated
for previous winter attrition on most treatments.

�46
44
42
40
38
36

~

34

Height,

Fall, 1976

32
30

Fjall, 1975

Ul

q) 28

~

26

,s 24
+l

..c: 22
01

'@ 20

:

',:.'::.':

::~'::.~

I'.:.~::'~:'

,'::: .'

:'00::':,

:::'::

:I:

18

.\\:\

~::::.~
.....
',0- ••,

Die-back

.;~:\;

Spring,

:::::

':: :0,:

::::':'.:

//!,~ Height, Spring" 1975

:':.:
....

16
14
12

o
Cultivated

Organic
Mulch

1976

Plastic
Mulch

Simazine

Sim.Mulch

EptamTreflan

Casoron

Fig. 1. Average shrub clump height and growth of wild plum per treatment
eight sites from spring, 1975 through fall, 1976.

on the

l

I
I-'

-...J

N
I

�34
32

l

30

m

28
26
24

~

~."l'

l

$::~
.,,~.(.

VJ
(lJ

:. ,0,

g 16
.j.J

rw

{U(
~

H

.~

:.:::
.....
.:.::',
'''','
......

:-:
":.
......
......

18

14

/ Height, Fall, 1975
~ Winter Die-Back

rt""l
.':.:.

22
20

.-"Height, Fall, 1976

~~~,

.:.:::
. .

. .

)(
....

Spring, 1976

::-:.:.::'
,

....
.....

:'::':':';,

~:

': ~:::

Height, Spring, 1975

12' J.

-&amp; 10',

I
•.....
-...J

•.-j

l.;.&gt;
I

(lJ

::x:

I1~

8
6
4
2
0

Cultivated

Organic
Mulch

Plastic Simazine
Mulch

Simazine Eptam
Mulch
Treflan

Casoron

Fig. 2. Average shrub clump height and growth of Hansen rose per treatment on
the eight sites from Spring, 1975 through Fall, 1976.

�-174-

Shrub Spacing
Shrubs were planted at three foot intervals in this study design in efforts
to achieve a closed canopy of thicket type cover within as short a time as
possible.
The more rapid the canopy closure the sooner maintenance and weed
competition could be eliminated.
This canopy closure could potentially
have been achieved by the end of the third or fourth growing season if rainfall had been average or above during the first three years of study. Severe
drought impact, as it affected the Karg sites through 1976 points out the
fact that such close spacing increases competition among the shrubs themselves when rainfall is deficient.
Severe dessication and die-back over
winter may occur as they grow larger. However, in naturally occurring old
thickets of plum in eastern Colorado the plants and their root sprouts become
closely spaced and competitive for moisture in much the same manner.
Spacing
of shrubs greater distances apart in initial planting so that they may reach
greater maturity before becoming highly competitive for moisture is the
primary alternative for use on dryland sites where supplemental moisture is
not available.
This, of course, increases maintenance time and costs of
establishment.
Wider spacing may not be warranted where mulch treatments
are used because available moisture is better conserved and utilized by the
shrubs and growth and survival are better.
Costs in mulching wider spaced
shrubs would also be increased tremendously.
Where chemicals are applied or
cultivation maintenance alone is used, wider spacing at four to six foot
distances may be the most practical compromise between growth-survival
variables and years needed in maintenance.

Weed Competition
All chemical and mulch treatments significantly reduced numbers of annual
weeds present in spring, 1976 when compared with cultivated controls
(F6,36 = 5.67 P &lt;0.001) (Table 6). Samples analyzed here by Dr. Bowden
included those from the five original sites and the three sites established
in 1975. Analyses within treatments, omitting the control, showed that weeds
were reduced to a significantly greater degree on plastic mulch and Casoron
treated plots than on Simazine treated plots (F5 ,30 = 2.83, P &lt;0.05).
Increased weed density apparently occurred on the Simazine plots over that of
1975 because this chemical was less effective in its second year after application (on the five original sites). Table 7 shows that average weed densities per square foot sample increased from 2.10 in 1975 to 2.66 in 1976,
whereas weed densities declined on most other treatments, including the controls because of dry weather conditions in 1976. A second application of
Simazine at a much reduced rate should be applied prior to the third growing
season after application to supplement the original application.
Reduced
weed density was indicated in 1976 over 1975 on the Simazine mulch plots
(Table 7) where weed control appeared to be slightly more effective than
where Simazine was the only treatment applied.
Casoron, a pre-emergent herbicide applied each early spring interval after
shrub establishment, was highly effective in controlling annual forbs in
1976 (Table 6). As Table 7 illustrates control in 1976 was much more effective than in 1975 indicating the probability of chemical retention from the
previous year and build up with the second year application.
Because this
residual carryover
from year to year was indicated, subsequent annual
applications should be at reduced rates to prevent excessive shrub mortality.

�-175Table 6. Average occurrence of annual weeds per square foot sample in May,
1976 within nine sites within three soil types.

Soil Type
Site

Treatment
Plast.
Mulch Sim.

Cult.

Organic
Mulch

Karg 1

20.28

0.36

0.11

Karg 2

3.00

0.19

Karg 3 }./

14.33

Tamarack Meadow

'1:./

Sim.
Mulch

Eptam
Treflan

Casoron

Site
Mean

7.31

2.50

2.81

0.11

4.78

0.17

6.03

12.75

0.64

0.08

3.27

0.03

0.08

1.28

0.25

1.83

0.25

2.58

9.80

0.19

0.33

0.19

0.06

0.81

0.14

1.65

62.30

0.75

1.25

1.67

0.14

5.50

0.08

10.24

16.28

2.14

0.00

1.14

0.44

2.22

0.28

3.21

Red Lion

0.19

0.03

0.00

0.50

0.03

0.06

0.03

0.12

Sharp tail

3.05

0.03

0.00

3.63

0.08

0.28

0.08

1.02

1-76

6.11

0.11

0.00

2.22

0.47

0.47

0.06

1.35

Overall Mean

15.04

0.43

0.22

2.66

1.86

1.62

0.12

3.14

Loam

Clay Loam

Smith Meadow
Sonnenburg
Loamy Sand

1/ The Karg 3, Sonnenburg

and 1-76 sites were establ~shed
The other six sites were planted in April 1974.

in April 1975.

l/

The Smith Meadow site was excluded from survival and growth analyses because
of unforeseen biases inherent within the site.

Application of the Eptam-Treflan pre-emergent herbicide chemical mixture
were moderately effective in weed control in both 1975 and 1976 as indicated
in Table 7. These chemicals break down during the growing season and no
evidence of a chemical build up was indicated.
Plastic mulch, with its organic mulch overlay, was highly effective in weed
control during both 1975 and 1976 and plots so treated remained virtually
weed free except where the plastic became exposed and torn (Table 7).
Exposure and tearing of the plastic was minimized by retaining an adequate
coverage of organic mulch or the ridges between rows. Plots treated singularly

�-176-

with organic mulch usually contained few weeds in either year (Table 7).
however, some progressive settling and deterioration of the mulch overlay
was noted.
Some increase in weed occurrence might have occurred if the 1976
growing season had not been excessively dry.

Table 7. A comparison of average annual weed densities
sample within treatments in 1975 and 1976.

per square foot

1975

1976

29.29

15.04

Organic mulch

0.87

0.43

Plastic mulch with organic overlay

0.38

0.22

Simazine

2.10

2.66

Simazine with organic mulch overlay

5.45

1.86

mixture

1.15

1.62

5.56

0.12

Treatment

Cultivated

control

Eptam-Tref1an
Casoron

Annual weed densities were sampled in mid- to late May 1976, and removed from
all sites by mid-June.
Lack of rain to stimulate new sprouting retained the
sites in a near maintenance free, weed free situation throughout the remainder
of the summer. A thundershower stimuated some late summer weed germination
and growth in the loamy sand (Sandhi11s) sites. These weeds were removed in
August and none had attained significant growth to be competitive with the
shrubs. As with earlier spring samples, simazine and cultivated control
plots possessed the higher weed densities and Casoron and plastic mulch plots
retained the lowest weed occurrence.
Perennials, including noxious perennials,
were less a problem in 1976 than in previous years.
LITERATURE
Snyder, W. D. 1975.
clump p1antings.

CITED

Minimum tillage techniques for establishing shrubs in
Colo. Div. of Wi1d1.
Game Res. Rept. Apri1:141-153.

1976. Minimum tillage techniques for establishing shrubs in clump
p1antings.
Colo. Div. of Wi1d1.
Game Res. Rept. April: 209-234.

Prepared

by

9JJ'~
~
Warren D. Sny er
Wildlife

"'.L

Researcher

C

�-177-

JOB PROGRESS REPORT

State of

Colorado

Project No.

W-124-R-4
1

Work Plan No.
Job Title

Raptor Investigations
Job No. 1

Statewide Raptor Populations

Period Covered:

and Characteristics

March 27, 1975 through February

Studies

28, 1976

ABSTRACT
One hundred and thirty-four

golden eagle nests were surveyed,

of which

51 were occupied and 35 produced

and fledged an average of 1.3 young per nest.

The number of occupied peregrine

falcon eyries continued

1974 to 8 in 1975 which represents
27%.

Only 5 young peregrine

was 0.8 young/successful
young Jledged

a drop in percent. occupancy

falcons were fledged

pair).

to decline from 9 in

Prairie

from 29% to

(average fledging success

falcon productivity

per site for 99 of 105 active nest sites.

averaged

3.37

Osprey productivity

continued

to remain minimal with 6 of 14 being active and only one egg being

produced

and no young fledged.

The aerial trend count in northeastern

yielded an average of 6.7 golden eagles/lOO
golden eagles/lOO

square miles was calculated

count in northwestern
Luis Valley
Wintering
viduals

square miles.

Colorado.

An additional

resulted in an average

Colorado

A density of 14.6

for a newly added aerial trend
aerial trend count in the San

of 11.2 golden eagles/lOO

square miles.

bald eagles along the South Platte count area declined from 42 indiin 1975 to 29 in 1976, but the percent of juveniles

The San Luis Valley aerial trend count yielded
with juveniles

accounting

remained at 35%.

16.0 bald eagles/lOO

for 30% of the population.

square miles

��-179-

S~TEWIDE

RAPTOR POPULATIONS

AND CHARACTERISTICS

STUDIES

Gerald R. Craig

To investigate
including

status of current populations,

distribution,

Eagles, Osprey,

migration,

Peregrine

patterns

characteristics

and chronology

and Prairie Falcons

and trends

of Bald and Golden

and Burrowing

Owls in or through

Colorado.
SEGMENT
1.

Develop

and refine methods

Golden

Eagles, Osprey,

OBJECTIVES

for determining

Peregrine

breeding

numbers

and Prairie Falcons

of Bald and

and Burrowing

Owls

winter trend information

for

in Colorado.
2.

Develop

and refine methods

for determining

Bald and Golden Eagles, Peregrine

and Prairie Falcons

and Burrowing

Owls

in Colorado.
3.

4.

Locate and categorize

major and important

and Golden

Eagle, Osprey,

populations

in Colorado.

Establish

and maintain

Eagles, Osprey,
Colorado

Peregrine

from inventory

inventories

and Prairie Falcons

methods

and associated

Bald

and Prairie Falcon and Burrowing

annual or periodic

Peregrine

habitats

selected

Owl

of Bald and Golden

and Burrowing

Owls in

from Project Statement

Procedures

1 and 2.
5.

Monitor

mortalities

brought

to the Division

Eagles, Osprey,

6.

and inspect

to determine

Peregrine

species

that are obtained

Initiate

and maintain

tion, chronology,
Eagles, Osprey,

a lO-year raptor banding

Peregrine

reported

or

factors of Bald and Golden
and Burrowing

Owls and other

incidentally.

movement,

program

life tables and mortalities

and Prairie Falcons

to determine

migra-

of Bald and Golden

and Burrowing

Owls and other

accidentally.

Prepare a series of comprehensive
guides at intervals

mortality

and Prairie Falcons

species that are encountered
7.

injured and dead raptors

information

when the information

leaflets and recreational

becomes

available.

�-180RESULTS
Only 38 percent
occupied

(51 sites) of 134 golden eagle nests were determined

in 1975 which is a continued

decline

at that time).

(35 sites) producing

young for an average brood size of 1034 young per productive

site and an average

fledging

lagomorph

productivity

from 1974 (65

sites were occupied

tinued decline

However,

in site occupancy

to be

increased with 69%

success of 1.30 young per productive

in the number of active nest sites continues

populations

which are the predominant

site.

The con-

to reflect the low

prey of golden eagles throughout

the region.
The bald eagle nest that was discovered
in 1974 was not occupied

in 1975.

at Electra Lake in La Plata County

The osprey that had traditionally

the nest failed to return to the site in 1975.
discovered

nesting

occupied

falcon production

nest site was discovered,

1974 to 8 in 1975.

to the land owner, they

occupied

remained poor.

Despite the fact that one previously

the number of occupied

sites declined

between

in any given breeding

80 and 90 percent of the eyrie sites were

season.

Young production

also continued

to decline and only five young were known to have fledged successfully
in 1975.

This represents

to occur with shell thickness averaging

shells considered

to be "normal"

(prior to the DDT era).

Ninety-nine

of 105 prairie

falcon eyrie sites were determined

The number of active sites which produced

60% in 1974 to 71% in 1975) as did fledging success
1975).

into the wild

an average of 0.8 young fledged per adult pair.

thinning continues

in 1975.

from 9 in

At this time only 31 percent of the known eyries are occupied.

Prior to the DDT era (mid-1940's)
normally

According

The pair suc-

the nest in 1973 and reared at least one young, but failed to nest in 1974.

Peregrine
unknown

A second pair of bald eagles were

along the Little Snake River in Moffat County.

ceeded in raising a single young in 1975.

occupied

The average brood size increased

somewhat

to 4.22 young per brood in 1975), and average
the same (from 3.00 young per successful

20 percent

eggs increased

Eggshell

less than

to be occupied
slightly

(from

(from 53% in 1974 to 56% in

(3.70 young per brood in 1974

fledging

success remained nearly

site in 1974 to 3.37 young per success-

ful site in 1975).
The osprey nesting

success continued

nest sites were occupied

to be disastrous.

in 1975 and only 1 of those sites produced

young were known to have been hatched or fledged.
artificial

nest platforms

Only 6 of 14 known

were constructed

As an experiment

at two locations

eggs.

No

in 1974,

on Grandby Lake (Grand

�-181-

County)

and two were placed on Shadow Mountain

of the artificial

structures

Reservoir

(Grand County).

One

was occupied by a pair in 1975 and an adult was

observed near a second structure.
The number of wintering

golden eagles observed

northwestern

Colorado

declined

from previous

years is consistent

and cottontail

rabbit populations.

during the aerial flights

to 6.7 eagles per 100 square miles.
with the region-wide
Preliminary

reduction

information

to adults is also increasing,

nesting

The aerial transects which were traditionally

success.

U. S. Fish and Wildlife

Service

of jackrabbit
that the

which may be indicative

of poor

flown by the

since 1972 were taken over by the Division

1976 and the data for all past flights is provided.

Consistent

northeastern

decline

Colorado,

This reduction

indicated

ratio of juveniles

there has been a significant

in

in

with trends in

in wintering

eagles.

In 1972 an average of 91.6 eagles per 100 square miles was observed.

The average

has declined until only 14.6 eagles per 100 square miles was counted

in 1976.

third aerial census route was delineated
flight yielded an estimated
area.

in the San Luis Valley

being juveniles.

The

11.2 golden eagles per 100 square miles of the study

Unlike the golden eagles, the wintering
Platte River has remained

in 1976.

A

bald eagle population

fairly stable with approximately

along the South

35% of the population

The aerial flight in the San Luis Valley yielded an estimated

16.0 bald eagles per 100 square miles.

�-182-

TABLE 1
ACTIVITY AND PRODUCTIVITY OF COLORADO GOLDEN EAGLES
Site No.

Activit~

No. of Young

No. Fledged

CH 4

A

2

2

CH 6

A

1+

1+

CN 1

IA

CN 2

IA

CN 3

IA

CN 4

IA

CO 1

IA

CU 2

A

2

2

CU 3

IA

DO 4

A

EA 1

IA

EA 2

A

2

2

EA 4

IA

EA 5

A

FA 7

IA

FA8

Adults observed courting
2

2

A

1

1

FA 9

A

1

1

FA 10

A

1

1

FA 12

A

FA 13

IA

EB 4

A

1

1

EP 1

IA

EP 2

IA

EP 3

IA

EP 4

IA

EP 5

IA

EP 6

IA

FR 1

IA

GA 4

IA

GA5

IA
1

1

2

2

GA

6

A

GR 1

IA

GR 2

A

GR 4

IA

GR 6

IA

Comments

2 eggs

-----------------------------------------------------------------------------------

�-183Table 1

Site No.

(continued) Activity

and Productivity

of Colorado

Activity

No. of Young

GR 7

A

1

GR 8

A

2

GR 9

IA

GR 11

A

1

1

GR 12

A

1

1

GR 13

IA

HU 1

IA

HU 2

IA

HU 3

A

2

2

HU4

IA

HU 6

A

o

o

HU 7

IA

JE 2

IA

JE 3

IA

JE 4

A

2

2

JE 6

IA

JE 7

IA

JE 8

A

JE 9

A

o
o

o
o

JE 10

A

2

2

IA 1

IA

LP 1

IA

LP 2

IA

LP 3

IA

LP 4

IA

LPS

IA

LS 1

IA

LS 2

IA

4

IA

LS 6

A

2

2

LS 7

IA

LS 8

IA

LS 9

IA

ME 6

IA

LS

No. Fledged

Golden Eagles

Comments

2

One adult observed

2 addled eggs

Both adults present
Adult

on nest

--------------------------------------------------------------------------------

on nest

�-184Table

1

Site

No.

(continued)

Activity

Activity

and

No.

of

Productivity

Young

of

No.

ME 9

A

ME 10

IA

ME 11

IA

MI 1

IA

MI 2

IA

MI 4

IA

MI 5

IA

MF 2

A

MF 3

IA

MF 4

IA

MF 5

IA

MF 6

IA

MF 10

A

0

0

MF 11

A

1

1

MF 13

IA

MF 14

A

1

1

MF 15

IA

MF 16

IA

MF 17

IA

MF 18

IA

MF 19

A

1

1

MF 21

IA

MF 22

IA

MF 23

IA

MF 24

IA

MF 25

IA

MF 29

IA

MF 31

IA

MF 35

IA

MF 37

A

1

1

MF 38

IA

MF 43

IA

MF 44

IA

MF 46

IA

MF 48

IA

2

Colorado

Fledged

Golden

Eagles

Comments

2

2 adults

present

1 adult

present

----------------------------------------------------------------------------------

�-185Table 1

Site No.

(continued) Activity and Productivity of Colorado Golden Eagles

Activit:l

ME' 49

A

ME' 52

IA

ME' 54

IA

MF 55

IA

MF 57

A

ME' 58

IA

ME' 61

IA

MF 62

No. of Young

No. Fledged

Comments
1 egg in nest

1 adult present
0

0

A

0

0

MF 63

A

1

1

ME' 64

IA

OU 1

A

1

1

OU 2

A

1

1

OU 3

A

2

2

OU 4

A

0

0

OU 5

A

1+

PA 2

A

PA 3

A

PI 1

A

PU 2

A

PU 3

IA

2 adults present

PU 5

A

2 eggs in nest

PU 6

A

1 whole addled egg &amp; fragments

PU 7

IA

RO 1

A

RO 2

IA

RO 3

IA

RO 4

0

2

2

1

1

A

1

1

RO 5

A

1

1

RO 6

A

1

1

RO 7

A

1

1

-------------------------------------------------------------------------------------

�-186-

Table

1

(continued)

Site No.

Activity

Activity

WE 1

IA

WE 2

A

WE 3

IA

WE 4

and Productivity

of Colorado

Golden Eagles

No. of Young

No. Fledged

Comments

1

1

A

0

0

WE 5

A

2

2

WE 6

A

1+

1

WE 7

A

1+

1

WE 8

A

2

0

Both chicks killed by hail

WE 9

A

0

0

Egg fragments

WE 10

A

2

2

WE 11

A

1+

1

WE 12

IA

WE 13

IA

S~RY

OF ESTIMATED

PRODUCTIVITY

OF COLORADO

60 of 147 (38%) sites were determined

Adult

GOLDEN ~GLES

to be active.

4S (70%) of the active sites were known to have produced.
young ,
39 (65%) of the active

sites were known to have fledged

yOtttlg.

A total of 57 young were observed
fledging,

thus average brood size was estimated

young per productive

thus average

young per productive

at 1.30

site.

A total of 54 young were observed
nests,

in 42 nests prior to

fledging
site.

to have fledged from 41

success was estimated

at 1.32

incubating

in nest

�OCCUPANCY OF COLORADO PEREGRINE FALCON EYRIES 1964-1975
TABLE 2
Site
1
2
3
4
5
6
7
8
9

-io
*11
12
13
14
15
16
17
18
19
20
21
22
23
24
25

Pre1964

+
+
+

1964

1965

P
P

P
P
A
A
M

P

+

1960

1967

1968

1969

1970

1971

1972

1973

1974

1975

P
P

P

P
P
P
P
P

M
P
P

P
P

P
P

P
P
P
A
P
P
P
P
P

P
P
P
P
P
P
P
P
P
P

M
P
M
V
P
P
P
V
P
V
P
V
V
V
P
V
V
V
V
V
V
V
V
V

V
V
Ml/
VP
P2/

P
P
A
P

V

P
A
F

+
+
+
+
+
+
+
+
+

V
V

+

V

P
V

P
M
P

A

V
V
V
V

V
V
V
V

V
V
A

!:J:./

V
V
V
V
A

V

V
A

P
V
A
V
V
V
V
V
V
V
V
V
V

r=

V
P
V3/

r=

V
V
Vp=.
5/
V
V
V
V
V
V
V
V
V
P

P = Pair, M = Male, F = Female, A = Lone Adult, V = Site was Vacant, Blank Spaces = No Data Available.

* These are neighboring sites (approximately 1 mile apart) and this could represent a shift of one pair from
site 10 to ll.

1/ Late reports indicate the pair may have relocated 2-3 miles upstream of the historic site.•
~/ Lone adult male observed throughout most of season, attracted juvenile female in June.

1/ Adult male was replaced by a juvenile male midway through incubation.
!:J:.I An adult female was found dead in the vicinity of the eyrie.
11 Adurt female disappeared shortly after hatching. Male succeeded in rearing two young.

I
I-'

co
-...J
I

�-188~BLE

3

RESULTS

OF NESTING ATTEMPTS

Eyrie

Visits

Eggs

Young

6

9

1+, 4

o , ~f

5

7

4

?

7

5

o

o

11

4

3

o

15

6

4

4, 2

Pair present March 30; adult female in
addition of pair observed April 25; 4
eggs were discovered May 4; 4 young
were found on May 17 and an additional
chick from eyrie 6 was added; female
disappeared June 20; 3 dead chicks
were discovered July 1 - 2 chicks were
alive; 2 young fledged mid-July.

25

2

?

3, 3

Pair observed mid-May; 3 half-grown
young observed early July; all young
assumed to have fledged successfully.

!f First

figure refers to the number
fledged.

OF COLORADO

PEREGRINES

IN 1975

Remarks
Pair observed copulating March 26;
incubating April 13; ceased incubating
April 25; 1+ broken eggs found
April 28; 4 new eggs were found May 19
and 2 were removed for artificial
incubation; pair failed and 1 broken
egg was found June 16; artificially
incubated eggs hatched and 1 chick was
placed with young in eyrie 5 and 1
chick in eyrie 15.
Pair present at site March 30; observed
copulating April 25 when they should
have been incubating - had also moved
to a new portion of the cliff; not
incubating May 2; incubating May 22;
4 eggs were found May 27; 2 young were
found June 30 and 1 artificially incubated and hatched young from eyrie 6
was added; only adult male was observed
August 10 and 11 - assume the nesting
attempt failed as the female and young
could not be found.
Lone adult male observed on visits
from March 28 through May 1; adult male
and yearling female observed June 6.
Pair observed March 29; 3 spoiled eggs
discovered May 31 and the adult male
had been replaced by a yearling male.

in the brood,

second figure refers to the number

�-189TABLE 4
OCCUPANCY AND PRODUCTIVITY OF COLORADO PEREGRINE EYRIES 1972-75
Year

1972

1973

1974

1975

4F

of eyries visited

15

23

25

26

4ft

of occupied sites

11

12

9

8

4F

of adult pairs

8

11

7

6

4F

of lone adults

3

1

2

1

4F

of young fledged

2

11

51/

4F

of young/adult pai~/

0.2

1.6

0.8

. ell/
% of sites occup1.e

73%

52%

38%

31%

% of sites w/adu1t pairs

53%

49%

29%

27%

1/ One site contained 3 young, but adult female and young could not be found
2 to 3 weeks after they should have fledged.
is reduced to 5.

Thus, probable fledging success

1/ An average of at least 1.4 young fledged per pair is considered necessary
to sustain the wild population.

1/ Between 10% and 20% of the eyrie sites would normally be expected to be
unoccupied in any particular year.

�-190TABLE 5
EGGSHELL CONDITION OF COLORADO PEREGRINES

Eyrie

Year

1
2

1973
1973
1974

"
"
"

Thickness (mm) 11
W/Membrane
vto Membrane

"

"

"

6

1974
1975
1971

"
"

1975

5

"

"

"

7
8
10
11

1974
1973
1974
1975

"

"

"

II.

"

II

15

1974
1975

."

.319
.299

.325
.305
.287
.282
.259
.259
.259
.284
.294
.300
.246
.292
.272
.254
.297
.305
.297
.284
.305
.203

Ratcliff's
Index
1.62
1.52
1.43
1.39

Remarks

Hatched
1.49
Hatched

.231
.278
.288
.244
.203
.236

Broken
Hatched
Hatched

.248

1.45
1.33

.244
.234

1.64
.1.61
1.61

Hatched
Spoiled
Spoiled
Spoiled

.252

Means
1975
1974
1973
Pre 19402.1

.273
.280
.295
.395

(n=8)
(n=7)
(n=3)

.235

(n=7)

.289

(n=3)

11 Measurements taken around waist of egg
11 Eggshells fromA1ta.,

Sask., Montana; Anderson and Hickey, 1972.
of XVth Int. Orith. Congr.

In ,Proceedings

�-191-

TABLE 6
ACTIVITY AND PRODUCTIVITY OF COLORADO PRAIRIE FALCONS
Site No.

Activity

No. of
Eggs

No. of
Young

No.
Fledged

3+

3+

BE 1

A

BO 1

A

BO 2

A

CN 1

A

3+

3+

CN 2

A

3+

3+

CN 3

A

1+

1+

Commerrt s

CN 4
DO 1

A

DO 2

A

DO 3

A

3+

3+

DO 4

A

3

3

DO 5

A

DO 6

A

0

0

DO 7

A

DO 8

A

DO 9

A

2+

2+

DO 10

A

5

5

DO 11

A

5

5

DO 12

A

5

5

DO 13

A

DO 14

A

4+

4+

DO 15

A

3+

3+

DO 16

A

DO 17

A

1+

1+

DO 20

A

0

0

DO 21

A

3+

3

DO 22

A

4

DO 23

A

1

5
Adult incubating

2

Adult incubating

EP 1
EP 2

A

EP 3

A

EP 4

A

EP 5

A

EP 6

IA

EP 7

IA

-----------------------------------------------------------------------------------

�-192-

Table 6

Site No.

(continued) Activity and Productivity of Colorado Prairie Falcons

Activity

No. of
Eggs

No. of
Young

No.
Fledged

Comments

EP 8

A

EB 1

A

4

4

EB 2

A

4

4

EB 3

A

0

0

EB 6

A

4

4

EB 7

A

5

5

GR 1

A

4

4

GR 2

A

0

0

GR 3

A

5

4

4

GR 4

A

HU

1

A

HU

2

A

2

0

0

HU 4

A

JA 1

A

LS 4

A

4

4

LS 5

A

5

5

JE 1

A

1+

1+

JE 2

A

2+

2+

3

A

PA 2

A

PA 3

A

1+

1+

PU 2

IA

PU 7

A

0

0

Adult incubating

PU 10

IA

PU 11

A

PU 12

IA

PU 13

A

RG 1

A

0

0

Adult incubating

RG 2

A

RO 1

A

3

3

RO 2

A

5

5

SA 1

A

SA 2

A

MF

5

Adult incubating

2 addled eggs

-----------------------------------------------------------------------------------

�-193-

Table 6

(continued) Activity and Productivity of Colorado Prairie Falcons

Site No. Activity

No. of
Eggs

No. of
Young

No.
Fledged

Comments

LR 1

A

4

4

LR 2

A

4

4

LR 3

A

4

4

LR 5

A

4

4

LR 6

A

3+

3

WE 1

A

3

3

WE 2

A

5

1

WE 3

A

1+

1

WE 4

A

1+

1+

WE 5

A

0

0

WE 6

A

5

5

WE 7

A

3

3

WE 8

A

2+

2

WE 9

A

4

0

all young appeared to be blind

WE 10

A

2+

1

one dead below nest

WE 11

A

5

5

WE 12

A

5

5

WE 13

A

3

1

WE 14

A

4

4

WE 15

A

2+

2

WE 16

A

5

2+

WE 17

A

?

0

bobcat scat in nest

WE 18

A

0

0

adult incubating

WE 19

A

0

0

WE 20

A

2+

2

WE 21

A

2

2

WE 22

IA

WE 23

A

WE 24

A

5

5

WE 25

A

WE 26

A

3+

3

WE 27

A

2

5

2 dead young in nest, 2 dead below

2 eggs addled

2 young dead below nest

3 eggs were addled

adult female dead below nest

------------------------------------------------------------------------------------

�-194-

Table

6

Site No.

(continued) Activity and Productivity of Colorado Prairie Falcons

Activity

WE 28

A

WE 29

A

WE 30

A

WE 31
WE 32

No. of
Eggs

No. of
Young

No.
Fledged
0

1+

1

0

0

A

2+

2

A

5

5

2

Comments
all young were taken
2 addled eggs

TABLE 7

SUMMARY OF ACTUAL AND ESTIMATED PRODUCTIVITY OF COLORADO PRAIRIE FALCONS
99 of 105 (94%) sites were determined to be active
70 (71%) of the active sites produced eggs.
58 (59%) of the active sites hatched young.
55 (56%) of the active sites fledged young.
Total number of young observed prior to fledging = 135
Average brood size of 32 sites with known production = 4.22
Adjusted young production for all sites (58) hatching young
Total number known to have fledged from 41 sites = 138
Average known fledging success = 3.37
Adjusted fledging success of all productive sites (56) = 188

244

�-195TABLE

8

1975

Site No.

Activity

OSPREY ACTIVITY

AND PRODUCTIVITY

No. of Eggs

No. of Young

No. Fledged

Connnents

GR 1

IA

GR 2

A

GR 3

A

GR 4

A

GR 5

A

?

?

GR 6

A

0

0

GR 6a

IA

Artificial nest sticks blown off platform. Adult nearby.

GR 7

IA

Artificial

nest

GR 8

IA

Artificial

nest

JA 1

A

JA 2

IA

JA 3

IA

IA 1

IA

LP 1

IA

SUMMARY

1

Egg on ground,
abandoned
0

0

0

Artificial nest,
adult incubating
empty nest

0

OF ACTUAL AND ESTIMATED

PRODUCTIVITY

6 or 14 (43%) sites were determined
1 of 6 (17%) sites produced

o of the active

nest

0

Adult

OF COLORADO

OSPREYS

to be active.

eggs.

sites produced

Sticks added to nest,
adult nearby

or fledged young.

present

�-196TABLE 9
RESULTS OF AERIAL FLIGHTS FOR GOLDEN ~GLES
Northeastern Colorado 1972-1976 (10% sample of 3,000 square miles)
Date

Adults

Juveniles

Unknown

Eagles per
100 sq. mi.

Estimate of
Total Eagles

Jan. 24, 1973

16

8

0

8.0

240

Jan. 16, 1974

19

3

0

7.3

220

Jan. 22, 1975

22

5

0

9.0

270

Feb. 19, 1976

17

3

0

6.7

200

Northwestern Colorado 1972-1976 (7% sample of 4,100 square miles) Jj
Date
Adults
Juveniles
Unknown
Eagles per
Estimate of
100 sq. mi.
Total Eagles
Jan. 25 and
Feb. 26, 1972
91.6
86
80
97
3,757
Jan. 23, 1973
41.5
35
14
70
1,700
Jan. 22 and
Jan. 29, 1974

6

8

47

21.2

871

Jan. 29, 1975

11

8

17

12.5

514

Jan. 26, 1976

29

3

10

14.6

600

San Luis Valley, Colorado 1976 (10% sample of 2,500 square miles)
Date
Jan. 29, 1976

Adults

Juveniles

Unknown

Eagles per
100 sq. mi.

Estimate
Total Eagles

17

9

2

11.2

280

1/ Flights from 1972 through 1975 were conducted by the U. S. Fish and Wildlife Service

�-197TABLE 10
RESULTS OF AER~L

FLIGHTS FOR BALD EAGLES

Northeastern Colorado 1972-1976 (exact count)
Date

Adults

Juveniles

Unknown

Total

% Juveniles

Jan. 24, 1973

18

13

0

31

42%

Jan. 16, 1974

16

15

0

31

48%

Jan. 22, 1975

28

14

0

42

33%

Feb. 19, 1976

19

10

0

29

35%

Northwestern Colorado 1972-1976 {7% same1e of 42100 sguare miles 2
Date

Adults

Juveniles

Total

Eagles per
100 sg. mi.

Estimate of
Total Eagles

Jan. 25 and
Feb. 26, 1972

1/

35

23, 1973

1/

4

12.2
1.4

500

.Jan ,

1/
1/

Jan. 22 and
Jan. 29, 1974

1/

1/

5

1.7

71

Jan. 29, 1975

1

0

1

0.4

14

Jan. 26, 1976

7

4

11

3.8

157

57

San Luis Valle~2 Colorado 1976 {10% same1e of 22500 sguare miles2 .
Date
Jan. 29, 1976

Adults

Juveniles

Total

Eagles per
100 sg. mi.

28

12

40

16.0

Estimate of
Total Eagles

1/ No distinction was made between adult and juveniles on these flights.

400

��-199-

JOB PROGRESS

State

of

REPORT

~C~o~l~o~r~a~d~o~

_

Pro j ec t No. -..:.:.W_-..=1.::2..:,4_R:.:--4-=-

_

Raptor

Work

_

Job No. 2

Plan No.

Job Title:
Period

.::.1

Raptor

Covered:

Population

March

and Productivity

27, 1975 through

Studies

February

Investigations

in Northeastern

28, 1976

ABSTRACT

Nine golden

eagle,

nests were occupied
Colorado.

31 prairie

on the 2,000

Productivity

of golden

to 1973 and 1974, but the number
remained

stable.

falcon,

and 12 ferruginous

square mile
eagles

study area in Northeastern

continued

of productive

,hawk

to be low compared

prairie

falcon

nests

has

Colorado

��-201-

RAPTOR

POPULATION

AND PRODUCTIVITY
COLORADO

STUDIES IN NORTHEASTERN

Gerald R. Craig

To intensively
chronology,
raptors

habitat

document nesting
requirements,

on the Pawnee National

and wintering

productivity

Grasslands
SEGMENT

densities,

migration

and prey preferences

and adjacent

prairie

of

lands.

OBJECTIVES

1.

Document exact nesting
on the Pawnee National

densities and productivity of raptors present
Grasslands and adjacent prairie lands.

2.

Document

3.

While at nests, record disturbance factors,
habitat type and nest site features.

prey items at nest sites.
land-use practices,

RESULTS
While

fewer active golden eagle nests were located in 1975 (22 sites

were occupied
fledged

in 1974, while 9 were active

(5 young in 1974 and 10 in 1975).

is blamed

on the continued

population

in 1975), more young were
The low number

decline of rabbits

of active nests
in Northeastern

Colorado.
The number of active prairie

falcon nests increased

in 1974 to 97% in 1975) and the average
young/site

in 1974 to 4.15 young/site

decreased

slightly

brood size increased

(from 2.94 young/site

(53 young fledged

Ferruginous

in 1975).

to have fledged remained

nearly

in 1974 compared with 58 young fledged

hawk nesting

success improved

considerably

12 of 14 sites

in 1975 while only 6 of 18 sites (33%) were occupied
sites increased

1974 while 9 sites produced
fledged

success

in 1974 to 2.79 young/site

not all the known nest sites were checked,

of productive

(from 3.92

in 1975), but the fledging

Thus, the total number of young estimated
constant

in 1975 (from 79%

significantly

in 1975).

in 1975.

While

(86%) were active

in 1974.

The number

(only 1 site was productive

young in 1975) and the total number of young

in 1974 was 35 while only 3 young fledged

in 1974.

in

�-202-

ACTIVITY AND PRODUCTIVITY OF GOLDEN EAGLES IN NORTHEASTERN COLORADO

Site No.

Activity

No. of Young

No. Fledged

1

1

Comments

WE 1

IA

WE 2

A

WE 3

IA

WE 4

A

0

0

WE 5

A

2

2

WE 6

A

1+

1

WE 7

A

1+

1

WE 8

A

2

0

Both chicks killed by hail

WE 9

A

0

0

Egg fragments in nest

WE 10

A

2

2

WE 11

A

1+

1

WE 12

IA

WE 13

IA

S~RY

Adult incubating

OF ACTUAL PRODUCTIVITY OF GOLDEN EAGLES IN NORTHEASTERN COLORADO

9 of 13 sites (69%) were determined to be active.
7 (78%) of the active sites produced young.
6 (67%) of the active sites fledged young.

A total of 10 young were produced by 7 sites (average
brood size
1.43 young).
A total of 8 young were fledged from 6 sites (average
fledging success = 1.34 young).

�-203-

PREY OBSERVED AT GOLDEN EAGLE NESTS IN NORTHEASTERN COLORADO

Prey Item

Number

Great Horned Owl

1

Mallard

1

Mule Deer (leg of fawn and hindquarter adult)

2

White-tailed Jackrabbit

1

Black-tailed Jackrabbit

5

Cottontail Rabbit

8

�-204-

ACTIVITY AND PRODUCTIVITY OF PRAIRIE FALCONS.
IN NORTH~STERN

Site No.

Activity

No. of
Eggs

COLORADO

No. of
Young

No. Fledged

Comments

WE 1

A

3

3

WE 2

A

5

1

WE 3

A

1+

1

WE 4

A

1+

1+

WE 5

A

0

0

WE 6

A

5

5

WE 7

A

3

3

WE 8

A

2+

2

WE 9

A

4

0

all young appeared to be blind

WE 10

A

2+

1

one dead below nest

WE 11

A

5

5

WE 12

A

5

5

WE 13

A

3

1

WE 14

A

4

4

WE 15

A

2+

2

WE 16

A

5

2+

WE 17

A

?

0

bobcat scat in nest

WE 18

A

0

0

adult incubating

WE 19

A

0

0

WE 20

A

2+

2

WE 21

A

2

2

WE 22

IA

WE 23

A

WE 24

A

5

5

WE 25

A

WE 26

A

3+

3

adult female dead below nest

WE 27

A

WE 28

A

0

all young were taken

WE 29

A

WE 30

A

WE 31
WE 32

2

5

2 dead young in nest, 2 dead below

2 eggs addled

2 young dead below nest

3 eggs were addled

.--

1+

1

0

0

A

2+

2

A

5

5

2

2 addled eggs

�-205-

SUMMARY OF ACTUAL AND ESTIMATED PRODUCTIVITY
OF PRAIRIE FALCONS IN NORTH~STERN COLORADO
31 of 32 sites (97%) were determined to be active
22 (71%) of the active sites were known to have produced young.
Total number of young known to have been produced by 13 productive sites = 54 (4.15 young/site)
Total adjusted productivity for 22 sites = 91 young
Total number of young known to have fledged from 19 productive
sites = 53 (2.79 young/site)
Total adjusted fledging success for 21 productive sites

58 young

�-206-

ACTIVITY AND PRODUCTIVITY OF FERRUGINOUS HAWKS IN NORTH~STERN

COLORADO

Site No.

Comments

Activity

No. of Young

No. Fledged

WE 1

A

4

4

WE 2

A

1+

1

WE 3

A

2+

2

WE 4

A

4

4

WE 5

A

3

3

WE 6

A

3

3

WE 7

A

3

3

WE 8

A

3

3

WE 9

IA

WE 10

IA

WE 11

A

3

3

WE 12

A

3

3

WE 13

A

4

4

WE 14

A

2+

2

S~RY

1 addled egg in nest

artificial nest structure

OF PRODUCTIVITY OF FERRUGINOUS HAWKS IN NORTH~STERN

COLORADO

12 of 14 (86%) of the sites were active.
12 of 12 (100%) of the active sites produced and fledged young.
Total number of young known produced by 9 productive sites = 30
(3.34 young/site)
Total adjusted productivity of 12 sites = 40 young
Total number of young known fledged by 12 sites = 35
(2.92 young/site)

�-207-

JOB PROGRESS REPORT
State of

~C~o~l~o~r=a~d~o

_

Project No.

W~-~1~24~-~R~-_4~

_

Raptor Investigations

_

Job 3

Work Plan No.

~l~
Raptor population

Job Title:

Period Covered:

and Productivity

Studies

March 27, 1975 through February

in Southeastern

Colorado

28, 1976

ABSTRACT
In Southeastern

Colorado,

three out of nine golden eagle nests fledged

a total of 5 young.

Twenty-six

of 38 ferruginous

young and 60 of 91 Swainson's
of productive

114 young.

The number

nests and total number of young of golden eagles and ferrugi-

nous hawks increased
Swainson's

hawk nests produced

hawk nests fledged 74

significantly

hawks declined.

in 1975 while the nesting success of

��-209-

RAPTOR POPUL&amp;TION

AND PRODUCTIVITY

William
To intensively
chronology,

habitat

C. Andersen

document

National

COLORADO

and Gerald R. Craig

nesting

requirements,

raptors on the Comanche

STUDIES IN SOUTHEASTERN

and wintering

productivity
Grasslands

densities,

migration

and prey preferences

and adjacent

prairie

of

lands.

SEGMENT OBJECTIVES
1.

To continue
golden

eagles, prairie

red-tailed
Comanche
2.

documentation

of nesting

falcons,

densities

ferruginous

and productivity

hawks, Swainson's

hawks, great horned owls, and burrowing

National

To continue

Grasslands

documentation

den eagles, prairie

and adjacent

prairie

ferruginous

hawks, great horned owls and burrowing

3.

To continue

documentation

4.

To investigate
turbance

factors, habitat

relief,

etc.

To continue

documentation

6.

To continue

to gather

To categorize

as dis-

type, human use, dominant vegetation,

of migration

trend information

important

hawks,

owls.

nest sites by such features

chronology.
on wintering

tions.
7.

cycles of gol-

of prey items at nest sites.

and categorize

5.

lands.

hawks, Swainson's

red-tailed

hawks,

owls on the

of the timing of breeding

falcons,

of

wintering

habitats.

raptor popula",

�-210-

RESULTS

Three of nine golden eagle nests were successful
of 5 young which averages
2 successfully
fledged
attempt.

fledged 3 young.

Twenty-six

In 1974 only

site.

of 38 ferruginous

hawk nests

74 young for an average of 2.85 young fledged per successful nesting
Thus, the average

to increase

fledging

(from 1.47 young/site

2.85 young/site
dropped

1.67 young per productive

in fledging a total

in 1975).

success for ferruginous

in 1973 to 1.89 young/site

in 1974 to

The number of occupied Swainson's

hawk nests

to 91 (176 sites were occupied

in fledging

in 1974) and only 60 sites succeeded

114 young (1.90 young/nesting

Due to mechanical

difficulties,

attempt).

weather

conditions

the migration

and winter trend counts were discontinued.

was available

to establish

that the trend information
overall population

hawks continues

chronology

of movements

Sufficient

data

and it was determined

could not yield satisfactory

densities.

and lack of funding,

information

about

�-211-

ACTIVITY AND NESTING SUCCESS - 1975
SOUTH~STERN

COLORADO
No.Successful
Nests

Young
Fledged

Young Fledged!
Succ , Nest

Species

County

Nest
Attempts

Golden Eagle

Baca

2

1

2

2.00

Bent

2

0

0

0.00

Las Animas

3

1

2

2.00

Otero

1

1

1

1.00

Lincoln

1

0

0

9

3

5

0.00
1.67

Baca

5

4

14

3.50

Bent

3

2

3

1.50

Cheyenne

3

1

3

3.00

Crowley

3

2

5

2.50

Kiowa

3

1

3

3.00

Las Animas

4

4

12

3.00

Lincoln

9

6

16

2.67

Otero

8

6

18

3.00

38

26

74

2.85

Baca

24

18

42

2.34

Bent

4

2

2

Crowley

9

5

8

1.00
1.60

Elbert

4

2

5

2.50

Kiowa

5

2

4

2.00

Las Animas

8

6

14

Lincoln

12

7

12

2.33
1.71

Otero

25

18

27

1.50

91

60

114

1.90

Ferrug. Hawk

Swainson's
Hawk

��-213-

JOB PROGRESS REPORT

State of

~C~o~l~o~r~a~d~o~

_

Project No. __~W_-~1~2~4~-~R~-_4~

~R~a~p~t~o~r~I~n~v~e~s~t==ig~a~t~i~o~n~s~
_

Work Plan No.

~J~ob~N~o~.~2~

~2~

_

Job Title: __~P~e~r~e~g~r~i~n~e~F~a~lc~on~~P~r~o~p~a~g~a~t~~~·o~n~a~n~d~R~e~i~n~t~r~o~d~u~c~t~~~·o~n~
_
Period Covered:

March 27, 1975 through February

28, 1976

ABSTRACT
Four pair of peregrine

falcons laid 39 eggs of which 10 were fertile, 10

of the eggs were broken by breeding adults.
cially incubated and five young hatched

The ten fertile eggs were artifi-

successfully.

Poor hatching

success

may have been due to excessive relative humidity which caused abnormal
weight

egg

loss.

Two of four eggs were removed from a wild peregrine
artificially

incubated.

falcon eyrie and were

The chicks were then returned to other wild eyries to

be reared with the wild broods.

In both cases the chicks were readily adopted.

��-215-

PEREGRINE

FALCON PROPAGATION

AND REINTRODUCTION

STUDIES

James H. Enderson and Gerald R. Craig
To captively

produce

eggs and/or young from peregrine

into the nests of wild peregrine

falcons

for reintroduction

falcons that have low or no natural

SEGMENT

productivity.

OBJECTIVES

1.

To establish

the cause for reproductive

2.

To produce young or eggs from peregrine falcons being held in captivity
at the facilities of Dr. James H. Enderson.

~.

To visit wild peregrine falcon eyries where wild adults are present
replace spoiled eggs with captively produced eggs or young.

4.

To keep those eyries containing captively produced eggs or young under
observation to ascertain the success of the endeavor.

S~RY
Attached
breeding

In addition

at Colorado

The particular

successfully

incubate

the peregrine

efforts which were reported

falcon nest was manipulated

production.

site in question

by Enderson,

in an effort to increase wild
has had a history

of failing to

and hatch the first clutch of eggs that are laid.

1972, the wild pair have experienced
has been laid.

which summarized

College.

to the captive breeding

one wild peregrine

and

OF RESULTS

is a report by James H. Enderson

project

It is suspected

cause for the egg breakage.
'recycled'

failure of wild peregrines.

egg breakage

that pesticide

However,

sometime

induced

Since

after the first clutch

~ggshell

thinning

is the

each time the first clutch failed,

and laid a second clutch which they

successfully

hatched.

the pair

In 1975,

an effort was made to visit the nest just after the first clutch was laid so the
eggs could be removed and artificially

incubated

so they wouldn't

abuse they experience

adults.

The thought was that the adults

under incubating

would then recycle and lay a second clutch as they had in previous
tunately,

when the site was visited,

adults had already
had completed

abandoned

the site and moved

the second clutch.

in hatching

had already

years.

occurred

to another nesting

disturbed

to the

Unfor-

and the

location and

The second nest location was adjacent

trail and the adults were frequently
they would not succeed

egg breakage

be exposed

to a

by hikers and it was feared that

the second clutch.

Therefore,

it was decided

that two of the four eggs in the clutch would be removed and artificially

incubated

�-216-

and the adults would be permitted

to incubate

adults were successful

the eggs, the two chicks that were hatched

in captivity

in hatching

would be returned

to the site to be naturally

If the wild pair failed to successfully

and two two-week

to care for them.

old prairie

at the site and left there for most of the day.
recognize

young.

Unfortunately,

young were placed on the nest ledge for

the adults were not inclined

nest ledge was revisited

they were returned

and the wild adults had failed to hatch their eggs

the two captive hatched

several hours,

raised by the adults.

in captivity,

to the wild eyrie site to be placed with the wild hatched

and although

If the

and held in captivity.

days after the two young had hatched

the worst had been realized

two eggs.

hatch the eggs, not all would be lost

since at least two would have been hatched
Several

the remaining

them and all efforts

to attract

The next day, the

falcon chicks were placed

Again,

the wild pair would not

the wild adults back to the nest ledge

were abandoned.
After

the two peregrine

falcon chicks were about two weeks of age, one of

them was placed

in another wild peregrine

Unfortunately,

the adult female disappeared

falcon eyrie with a brood of four young.
within

several days and the adult

male was forced to attempt

to raise the entire brood.

This he could not do and

three of the young died.

He was successful

two of the young.

in rearing

the orphan chick was not marked when it was placed
of identifying

falcon nest site which contained

While
proven

two similarly

Thus it has to be assumed

in another wild peregrine

aged young.

but neither

The site was revisited

they nor the adult female

that this site also experienced

dif-

and no young were fledged.
it is difficult

to assess

that eggs could be removed

incubated

falcon chick was placed

the young should have fledged,

could be located.
ficulties

there was no way

it, so its fate is unknown.

The second orphan peregrine

shortly after

in the brood,

Since

and hatched.

the success of the endeavor,

from wild pairs of peregrines

it was conclusively

and artificially

The young could then be placed with wild broods at other

sites and would be readily adopted by the wild adults.

�-2171975
Falcon Breeding

Peregrine

James H. Enderson,

This is a report on the results
Colorado

College

Zoological

in 1975.

Society

work is to produce
Rocky Mountain

The significance

in producing

many of these were produced

the release

of this

indigenous

to the

from the region

official planning

is derived

for augmentation

In 1975, Enderson
in the region,down

of

and J. Craig
from 14 in

pair was found in 1975; it was successful
Presently

from the western U.S. in captivity,

in breeding

being drawn up by the Rocky Mountain
emphasize

The purpose

young along with only two of the other seven pairs.

are fewer than 20 pairs of peregrines

at

jointly by the New York

of peregrines

found 7 pairs of adults

One additional

project

and falconry purposes.

with birds bred in captivity.

1973 and 10 in 1974.

falcon breeding

of Wildlife.

restocking

decline and increased

(Colo. Div. of Wildlife)

College

stock of the peregrine

of captive breeding

from their continuing
the wild population

was supported

Division

a captive breeding

region for eventual

The Colorado

of the peregrine

The project

and the Colorado

Project Report

projects.

The Recovery

Region recovery

there

and

Plan, currently

team will almost certainly

of captive bred birds as one means of affecting

a recovery

of the population.
Summary
five prairie

of Past Results.

This project was started

falcons and eight peregrines

Of the peregrines,

pairs remain at the Colorado
Results

in 1975.

College

Peregrine

Through

1974,

(3 in 1973, 5 in 1974) were produced.

one pair was sent to the breeding

and a second pair to the Cornell University

in 1968.

facilities

facilities

of E. Freienmuth,

in Colorado.

The other two

project.

falcons only were held in this project

including:
3 pairs adult anatum peregrines
1 adult pair, anatum female and male of unknown

origin

in 1975,

�-218-

1 pair of adult pea1ei peregrines
1 adult anatum male
2 pairs F1 captive bred anatum peregrines
1 adult anatum male

(on loan to R. Berry)

The three pairs of anatum peregrines
broken.
nation

Of the 22 intact eggs, 10 were fertile,
(AI).

yolk sac.

Six of the ten hatched,

The remaining

The Peale's
fertile

laid 31 eggs including

falcons

including

9 that were later

4 by artificial

insemi-

but one chick died with a large external

five (2 females,

3 males) are normal.

laid eight eggs in three clutches,

none of which were

despite AI.

The low productivity
copulating

relates

to three unresolved

difficulties.

1) The only

pair broke seven of the 14 eggs they laid, these were mainly

by the female from a high perch.

2) Of 19 eggs laid by the three non-copulating

pairs after AI, only four (21%) were fertile.
eggs, all were artificially
Information

on eggshell

dropped

3) Of the total of ten fertile

incubated,

but only five successfully

condition

was collected

so.

and awaits analysis

along

with that from former years.
The AI technique
be readily placed
elsewhere,

was improved with the help of R. Graham, and semen can now

in the oviduct.

this procedure

requires

Although

other t echnfques

the use of a speculum

have been worked out

to spread the vent open-

ing.
Proper water
obtained
incubation

loss from some artificially

incubated

since two eggs had lots of fluid remaining

eggs was apparently

at death near the end of

and three dead chicks near full term had large unretracted

The following

table shows the weight

that hatched

or went nearly

loss data collected

full term.

Humidity

1.10 g per 5 days as a target, and measured

not

yolk sacs.

on nine fertile

was varied

eggs

to yield a loss of

by a human hair hygrometer

of dubious

�-219-

accuracy.

Although firm conclusions are impossible, eggs from female A (13 yrs.

old) tended to lose water more rapidly than eggs from the other female.

Yet her

eggs had unretracted yolk sacs and remaining fluid, even with considerable weight
loss.

There is a possibility her eggs are initially abnormal.

If so, and if the

eggs of pair B are a guide, then a weight loss of 1.0 to 1.2 g per five days (2.02.5% loss of initial weight per five days) may be in the proper range.

1975

**

% loss
per 5 days

Result

Period
measured
(days)

Wt. loss
(g)

Wt. loss
per 5 days
(g)

47.49

23

5.17

1.10

2.4

Normal

46.26

23

6.03

1.30

2.7

Dead-yolk
sac &amp; fluid

44.99

21

5.83

1.40

3.1

Normal

40.36

21

5.95

1.40

3.5

Dead-yolk
sac

45.90

27

5.83

1.10

2.4

Dead-yolk
sac

47.63

23

4.49

0.95

2.0

Normal

47.41

23

3.93

0.85

1.8

Dead-fluid

45.77

21

4.59

1.10

2.4

Normal

45.71

21

4.26

1.00

2.2

Normal

Pair

Initial wt.
(g)

A ***

B

-- Weight of Artificially Incubated Peregrine Eggs*

*Ro11ex incubator - 98.50F.
**Usual1y measured 7 days after last egg in clutch was laid.
***Copu1ating pair.

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                  <text>July, 1977
-1-

JOB PROGRESS

State of
Project

REPORT

COLORADO
W-10l-R-19

No.

Game Range Investigations

Work Plan No.

4

Job Title

of Range Manipulation

Inventory

Period Covered:
Personnel:

Job No.

la

------------------------------

Projects

in Colorado

April 1, 1976 through March 31, 1977

Roland C. Kufeld and Regional

Game Biologists

ABSTRACT
An inventory was made of all range vegetation-modification
projects completed during 1975 in the western half of Colorado, on lands administered
by the u.S. Forest Service and Bureau of Land Management.
Acreages
treated were: Forest Service - 1208, and Bureau of Land Management - O.

��-3-

INVENTORY OF RANGE MANIPULATION PROJECTS IN COLORADO
Roland C. Kufeld
P.N.O. OBJECTIVE
To collect information which describes proposed and completed range
vegetation modification projects that are located in Colorado, and
their effects on the range, wildlife and livestock; and to provide
desired IBM listings of these data to cooperating agencies upon their
request.

SEGMENT OBJECTIVES
1.

To collect information which describes proposed and completed range
vegetation modification projects that are located in Colorado, and
their effects on the range, wildlife and livestock.

2.

To compile, codify, process and analyze inventory data.

3.

To provide desired IBM listings of inventory data to cooperating
agencies upon their request.
METHODS AND MATERIALS

An inventory was made of all range vegetation modification projects cornr
pleted through 1972, in Colorado, west of Interstate Highway 25 on lands
administered by the U.S. Forest Service and Bureau of Land Management
(Kufeld 1968, 1970,1971, 1973, 1974, 1975 and 1976). Kufeld (1968 and
1970) also covered projects completed on the Southern Ute and Ute Mountain
Indian Reservations through 1969. This report concerns vegetation modification projects completed on U.S. Forest Service and U.S. Bureau of
Land Management lands west of Interstate Highway 25 during 1975.
Data were collected using procedures outlined in Colorado Division of
Wildlife Administrative Directive No. 204 entitled "Range Vegetation
Modification Projects", and by Kufeld (1968, 1970 and 1974). Inventory
data were transferred from original data sheets to Mark Page Reader Forms,
and then to IBM cards.

RESULTS AND DISCUSSION
Acreages treated during 1975, on all lands administered by the U.S. Bureau
of Land Management west of Interstate Highway 25, are shown by vegetation
type and kind of treatment in Table 1. All of the U.S. Forest Service
projects completed in Colorado during 1975 were located on the San Juan
National Forest. No vegetation modification projects were completed on
U.s. Bureau of Land Management lands in Colorado during 1975.

�-4Table 1. Acreages of range land treated during 1975 in Colorado by
the U.S. Forest Service. 1/
Vegetation
Type

Grass

Sagebrush

2/
Browse -

Kind of
Treatment

No. of
Projects

Acres
Treated

Plow

1

66

66

1

66

66

1

250

250

1

250

250

Chaal.ni
3/

1

175

0

Burn

2

517

0

3

692

0

1

200

200

1

200

200

6

1,208

516

Plow

Pinyon-Juniper
Juniper
Burn

All Vegetation
Types GRAND TOTAL

Acres Seeded as
Part of Treatment

1./
All vegetation modification projects completed by the U.S.
Forest Service in 1975 were located on the San Juan National Forest.

All vegetation modification projects conducted during 1975 in the ''browse''
vegetation type were in the "oakbrush " subtype.

1/
Chaining includes chaining, cabling, railing and bulldozing.

�-5-

A total of 1,208 acres were treated on Forest Service lands in 1975.
To date (through Dec. 31, 1975) a total of 588,967 acres have been
treated in Colorado west of 1-25 on lands administered by the U.S.
Forest Service and Bureau of Land Management.
Of this, 246,183 acres
were on Forest Service and 342,784 acres were on Bureau of Land Management lands.
Procedures outlined in Administrative Directive No. 204, call for
evaluations to be made on each vegetation modification project at the
end of the 2nd, 5th, and 10th years following treatment.
Two and 5-year
post-treatment evaluations have been made on some vegetation projects
since 1970 by regional personnel.
Data from these evaluations have been
kept on file until enough 2 and 5 year post-treatment evaluations have
been made to provide a basis for a meaningful data analysis.
Due to
the workload and limited available manpower within the DOW Regions some
scheduled evaluations have been omitted or delayed.
This has caused a
breakdown in continuity of data which makes it virtually impossible to
derive any useful information from the relatively limited data that has
already been collected.
It is recommended that the post-treatment
evaluation phase of this project be discontinued at the end of the 19th
project segment.
It is also recommended that Division of Wildlife Management personnel be polled to determine their feelings concerning whether
there is a need for any part of the inventory of range manipulation
projects to be continued in the future.
If it is decided that the inventory should be continued, it is recommended that it only be concerned
with collecting data which describes proposed and completed range vegetation modification projects, with no emphasis placed on evaluating their
effects on the range, wildlife and livestock.

LITERATURE

CITED

Kufeld, R. C. 1968. Inventory of range manipulation projects in
Colorado.
Colo. Div. Game, Fish and Parks.
Game Res. Rept.
P.R.Proj. W-lOl-R-lO, WP 4, Jla, July, Part 1. p. 1-121.
1970. Inventory of range manipulation projects in Colorado.
Colo. Div. Game, Fish and Parks.
Game Res. Rep·. P ••
R P rOJ.
.
W-lOl-R-12, WP 4, Jla, July, Part 1. p. 59-94.
1971. Inventory of range manipulation projects in Colorado.
Colo. Div. Game, Fish and Parlffi. Game Res. Rep. P.R. Proj.
W-lOl-R-13, WP 4, Jla, July, Part 1. p. 1-15.
1973. Inventory of range man Lpu.La t i.on projects in Colorado.
Colo. Div. Wildl.
Game Res. Rept. P.R. Proj. W-lOl-R-lS, WP 4,
Jla, July, Part 1. p. 1-8.
1974. Inventory of range manipulation projects in Colorado.
Colo. Div. Wildl.
Game Res. Rept. P.R. Proj. W-lOl-R-16, WP 4,
Jla, July, Part 1. p. 1-11.

�-6-

Kufeld, R. C. 1975. Inventory of range manipulation projects in
Colorado.
Colo. Div. Wildl.
Game Res. Rept. P.R. Proj. W-10l-R-17,
WP 4, Jla, July, Part 1. p. 1-6.
1976. Inventory of range manipulation projects in Colorado.
Colo. Div. Wildl.
Game Res. Rept. P.R. Proj. W-10l-R-18, WP 4,
Jla, July, Part 1. p. 1-5.

Prepared

by

1(~·4~1
c, ){u/,t~d
Roland C. Kufeld
Wildlife Researcher

C

�-7-

July, 1977

Jon FIKt.&gt;.L REPORT

Colorado

State of
Project

No.

Game Range Investigations

W-10l-R-19

2
4
Job No.
Experimental Improvement of Oakbrush
on Deer and Elk Winter Ranges - Beaver Creek

Hork Plan No.
Job Title
Period

--------------------------~~~----------------------------------

Covered:

Personnel:

April 1, 1976 through March 31, 1977

Roland

C. Kufeld

P.N.O. OBJECTIVE
To determine if deer and elk forage production and game use can be increased on overage Gambel oakbrush winter game ranges by spraying with
2,4,5-TP to induce sprouting.

CURRENT STATUS
Results

of the study have been published

under the following

citation:

Kufeld, Roland C. 1977. Improving Garnbel oak ranges for elk and mule
deer by spraying with 2,4,5-TP.
J. Range Manage. 30(1) :53-57.

Prepared

by

12~ c, 2/J,tu/
Roland C. Kufeld
Wildlife Researcher

C

��July, 1977

-9JOB PROGRESS

REPORT

State of

--------~~~~~-----------

COLOR.4DO

Project

No.

W-10l-R-19

Game Range Investigations

3
4
Job No.
Experimental Improvement--o?f-nO~aTk~b-r-us-'hr--------------------on Deer, Elk and Cattle Ranges - Hightower Mountain

Work Plan No.
Job Title
Period

Covered:

Personnel:

April 1, 1976 through March 31, 1977

Roland C. Kufeld,
John Scotese.

Steve Brown, John Erickson,

Gordon Pepperd,

ABSTRACT
Five-year post-treatment vegetation measurements were made in 600 permanent
meter square plots on the Hightower Mountain oak study area. Data analysis
is underway and results will be presented in a future report.

��-11-

EXPERIMENTAL IMPROVEMENT OF OAKBRUSH
ON DEER, ELK AND CATTLE RANGES - HIGHTOWER MOUNTAIN
Roland C. Kufeld

P.N.O. OBJECTIVE
To determine the extent to which deer, elk and cattle forage production,
forage quality and game use can be increased and maintained by chaining,
spraying and controlled burning of overage Gambel oak winter game ranges.

SEGMENT

OBJECTIVE

To determine the extent of vegetative composition and forage production
changes, and the extent of deer and elk use changes which have resulted
from implementation of each habitat improvement method tested.
Determine
if cattle exhibit a preference toward certain habitat improvement units.

METHODS AND MATERIALS

Post-Treatment

Vegetation

Measurements

Five year post-treatment vegetation measurements were made in 600 permanent
meter square plots on the Hightower Mountain oak study area using the same
procedures employed during pre-treatment evaluations (Kufeld 1971).
The
measurement period was July 6, 1976 through August 6, 1976. Prior to beginning vegetation measurements observers were trained in the use of the
vegetation measurement technique, and in identification of all plants on
the study area.

Determination

of Nutrient

Quality

Changes

Samples of plant species designated by Kufeld (1971) as "moisture indicator
plants" and used to reflect changes in vegetation moisture content during
the measurement period were collected in Units 5, 6, 7 and 8 between July
19, and 22, 1976, inclusive.
Samples consisted of approximately 100 grams
green weight of each species and were collected throughout each habitat
improvement unit. In Units 5 and 7 samples were collected from those areas
where it was certain burning and spraying had been severe.
Samples from treated areas will be compared with those of the same species
collected in Unit 6, a control area, to determine changes in nutrient
quality of forage resulting from chaining, burning and spraying.
Samples
will be analyzed by the Division of Wildlife Research Laboratory to determine changes in content of dry matter, protein, energy and trace mineral
values 5 years after treatments were applied.
Laboratory analysis has not
been completed at the end of the 19th project segment, so results will be
presented in a later report.

�-12-

Photo Point Photographs
Post-treatment photos were taken at permanent photo points established
in 1970, prior to treatment.
These show effects of spraying, burning
and chaining during the fifth summer following treatment.
A black and
white photo, and a 35 mm color slide was taken at each photo point with
the same cameras used to take pre-treatment photos.

Post-Treatment

Deer, Elk and Cattle Use Measurements

Accumulated deer and elk pellet groups were removed from all pellet plots
on August 31, 1976.
Groups deposited during the winter of 1976-77 will
be counted in May 1977.
When pellet plots on the study area are cleared in the fall a record is
always kept of the number of cow chips in each plot in order to provide
an index to cattle use. The area is grazed on a rest-rotation system
and was "rested" in 1976 so cattle use data were not available.
This
information will be collected in early September, 1977.

RESULTS

Post-Treatment

AND DISCUSSION

Vegetation

Measurements

Data have been summarized and are currently undergoing computer analysis
by the Colorado State University Statistics Department.
Data are being
compared with results of the 1970 pre-treatment evaluation and 1973
2-year post-treatment evaluation, which were described by Kufeld (1975).
Results of the 5-year post~treatment evaluation will be presented in a
later report upon completion of the data analysis.

LITERATURE

CITED

Kufeld, R. C. 1971. Experimental improvement of oakbrush on deer, elk
and cattle ranges - Hightower Mountain.
Colo. Div. of Game, Fish
and Parks.
P-R Project W-lOl-R-13, WP 4, J 3, Game Research Rept.
July.
Part One. pp. 23-86.
1975.
Experimental improvement of oakbrush on deer, elk and
cattle ranges - Hightower Mountain.
Colo. Div. of Wildlife.
P-R
Project W-lOl-R-17, WP 4, J 3, Game Research Rept. July. Part One.
pp. 25-92.

Prepared

by

j(~ad C, 9L~/dd
Roland C. Kufeld
Wildlife Researcher

C

�July, 1977

-13JOB FINAL REPORT

Colorado

State of

-----------------------

Project

No.

W-lOl-R-19
6

vlork Plan No.
Job Title

Game Management

Job No.

1

Unit Inventory

-------------------------------------------------------------

Period Covered:
Personnel:

Game Range Investigations

April 1, 1976 through March 31, 1977

W. T. McKean,

Craig Jump, and Wally Jobman

ABSTRACT
Only 1/2 of the Program Leader's time was spent on W-lOl-R activities
during the Segment (19). The remainder was occupied in preparing an
Environmental Impact Statement for the Yellow Jacket Project.
During
time spent on W-lOl-R, a Big Game Analysis Report was completed for
Unit 12, the Williams Fork Unit.
With the completion of Unit 12, eight Big Game Unit Analyses have been
written.
These are: Unit 11, Strawberry Creek; Unit 12, Williams Fork;
Unit 21, Douglas; Unit 22, Piceance; Unit 31, Roan Creek, and Unit 32,
Rifle. All reports are on file in the Research Library at the Colorado
Division of Wildlife Office in Fort Collins, Colorado.

��-15-

JOB PROGRESS

State of

July, 1977

REPORT

COLORADO

------------------------------

Proj ect No.

W-10l-R-19

Work Plan No.

8

Job Title

of Preferred

Period Covered:
Personnel:

Game Range Investigations

1
Job No.
Digestible Nutrient~~-----------------------------Content
Deer and Elk Forage Plants During Winter

April 1, 1976 through March 31, 1977

Roland C. Kufeld, Marilyn

Stevens.

ABSTRACf
Crude protein content was determined for 45 big sagebrush and 45 Garnbel
oak samples collected during January, 1976. Each species was collected
from 45 sites throughout the western half of Colorado.
Collection sites
for each species represented 9 major areas of the state and 3 vegetation
types.
For sage the median crude protein level from the 9 areas is 10.1
percent and the 95 percent confidence level ranges from 8.8 to 10.6 percent. For oak the median crude level is 5.2 percent and the 95 percent
confidence interval ranges from 4.6 to 5.3 percent.
A 14.7 percent loss of essential oils in big sagebrush was found to occur
as a result of freeze drying and grinding sagebrush samples in preparation
for in-vitro digestibility trials.
This loss was not significant at the
5 percent significance level.

��-17-

DIGESTIBLE NUTRIENT CONTENT
OF DEER AND ELK FORAGE PLANTS DURING WINTER
Roland

C. Kufeld

P.N.O.

OBJECTIVE

To estimate the average digestible nutrient
of variation in digestible nutrient content
plants during winter.

SEGMENT

content values, and degree
of selected range forage

OBJECTIVE

To determine the degree of variation in digestible
selected range forage plants during winter.

METHODS

nutrient

content

of

AND MATERIALS

During January, 1976, five 70 to 100 gm, green weight, composite samples
of each of two species, big sagebrush (Artemisia tridentata) and Gambel
oak (Quercus garnbellii), were collected from 9 widely separated areas
throughout the western half of Colorado, but within the normal winter
range of deer and elk. The 5 collection sites within each major area
were widely separated geographically, but were situated within the same
vegetation type.
For sagebrush collections, 3 of the major areas represented each of the following vegetation types: sagebrush, pinyonjuniper, and Garnbel oak. Gambel oak samples were collected in the pinyonjuniper, Garnbel oak, and ponderosa pine-oak types.
Oak collections consisted of current annual growth stems with leaves and
acorns excluded.
Sage collections consisted of current annual growth
stems with leaves attached but seed stalks excluded.
Samples of big sage
were sealed in plastic bags, refrigerated immediately upon collection and
frozen as soon thereafter as possible to prevent vaporization of essential
oils which may result in unrealistically high digestion coefficients.
A portion of each sample was air and oven dried, ground in a Wiley mill
and subjected to proximate analysis in the Division of Wildlife Research
Laboratory.
Samples were analyzed to determine content of the following
nutrients and minerals:
Total cell contents, crude protein, ether extract,
soluble ash, soluble carbohydrate, total cell walls, acid insoluble ash,
lignin, hemicellulose,
cellulose, calcium, copper, iron, potassium,
magnesium, manganese, sodium. phosphorus, and zinc (Harris 1970; Adrian 1973).
The remaining portion of each sample was retained for determination
of
digestible dry matter, gross energy and digestible energy.
This analysis
has not yet been completed.
In preparation for this analysis the remaining
portion of each oak sample was air dried and ground in a Wiley mill.
Sagebrush samples were freeze dried for 7 days by the Colorado State University

�-18Natural Resource Ecology Laboratory in order to minimize the loss of
essential sagebrush oils which occurs during the drying process.
A study was made to determine the magnitude of essential oil loss
which occurs in sagebrush during the freeze drying and grinding process.
Ten composite big sagebrush samples of at least 200 grams each were
collected from 10 geographically widely separated areas within the
normal winter range of deer and elk during February, 1976.
Each 200+
gram sample was divided in half.
Each half was weighed to the nearest
gram, sealed in a separate plastic bag, refrigerated immediately upon
collection and frozen as soon thereafter as possible.
The result of
this collection was 10 paired samples.
One half of each pair of sagebrush samples was freeze dried and ground
at a temperature just above freezing exactly as prescribed for all other
sagebrush samples destined for in vitro digestibility trials.
Each was
weighed after freeze drying to determine moisture content.
The green
half of each sample was adjusted to dry weight on the basis of moisture
content in the freeze dried samples.
Both halves of each sample were
then distilled to determine percent of essential oils lost from freeze
dried and ground samples as compared to green ones.

RESULTS AND DISCUSSION

Nutrient Content Variation Among
Vegetation Types and Among Major Areas
To date only the results of the crude protein determination are available.
These data for big sage and Gambel oak from the 45 collection sites are
shown in Tables 1 and 2. Analysis of variance shows significant differences
among major areas and among vegetation types at the I-percent significance
level for both sage and oak. There are also significant differences among
areas within vegetation types at the 5-percent level for oak, but not for
sage.
Even though these significant differences occur, the differences are relatively small, biologically,
as evidenced from the mean and median values
in Tables 1 and 2. For sage the median crude protein level from the 9
major areas is 10.1 percent, and the 95 percent confidence interval ranges
from 8.8 to 10.6 percent.
For oak the median crude protein level is 5.2
percent and the 95 percent confidence interval ranges from 4.6 to 5.3 percent.
From the practical standpoint the data reflect relatively little
variation in crude protein content of big sage and Gambel oak from one
locality to another during mid-winter.
Thus~ for management purposes median
protein values may be used to estimate, with a reasonable degree of accuracy.
crude protein production of big sagebrush and Gambel oak during winter on
widely separated ranges.
Non-parametric
tolerance levels can be applied to further minimize the
error incurred in projecting crude protein production based on average
values (Dixon and Massey 1969).
It is desirable that carrying capacity be
underestimated
rather than overestimated to avoid overpopUlation
of the

�-19-

range. Non-parametric
tolerance levels permit determination of the portion of the overall population of nutrient values that could possibly be
encountered for a given species, which, with a 95-percent confidence,
exceeds the smallest sample value.
For example, using the sample means,
which range from 4.6 to 5.4 percent crude protein, from the 9 areas where
oak was collected, we can be 95 percent sure that at least 70 percent of
the area crude protein values encountered in oak in Colorado during
January would exceed the minimum value of 4.6 percent.
For big sage we
can be 95 percent sure that at least 70 percent of the area crude protein
values encountered in Colorado during January would exceed the minimum
value of 8.6 percent.
By using the minimum value a manager could, therefore, be reasonably sure his estimate of protein production is on the
conservative side.

Sagebrush

Essential Oil Loss in Big
Due to Freeze Drying and Grinding

Oil content averaged 1.3036 percent in green samples and 1.1123 percent in
those which were freeze dried and ground (Table 3). These data indicate
a 14.7 percent loss of essential oils due to the freeze drying and grinding
process.
A paired t test, however, indicated the loss was not significant
at the 5 percent significance level.

LITERATURE

CITED

Adrian, W. J. 1973. A comparison of a wet pressure digestion method
with other commonly used wet and dry-ashing methods.
Analyst 98:213-216.
Dixon, W. J., and F. J. Massey, Jr.
analysis.
McGraw-Hill, 3rd ed.

1969. Introduction
638 pp.

to statistical

Harris, L. E. 1970. Nutrition research techniques for domestic and
wild animals.
Pub. by Lorin E. Harris, 1408 Highland Dr., Logan,
Utah. Vol. 1.

Prepared

by

-&amp;fc.a~

C'J?BM

oland C. Kufeld
Wildlife Researcher

C

�Table 1.

Big sage crude protein content (%) from three vegetation types during January.
Oak

Pinyon-JuniEer

Ponderosa Pine-Oak
Steamboat
SiltB. CanyonPaoniaSpgs.-Hayden
Carbondale
Cedaredge
Meeker
Eagle

Craig

Middle
Park

Gunnison

PiceanceStrawberry

Naturita

CortezDurango

12.2

12.4

9.0

9.2

8.6

9.4

10.4

9.6

10.2

8.6

12.4

12.4

8.6

11.8

10.4

9.2

9.4

9.4

10.0

11.4

10.8

10.8

9.8

10.4

8.8

8.2

7.6

12.2

11.2

9.6

10.4

10.0

9.8

8.8

8.2

7.6

8.8

9.2

11.4

8.8

10.4

11.6

9.2

8.8

8.0

11. 3

10.6

9.6

10.1

10.3

9.3

8.8

8.6

X (Area)
10.4

X (Vegetation Type)
10.0

10.8

=

9.9

Total Median =

10.1

Total Mean

8.9

I
N
0
I

�Table 2.

Gambel oak crude protein content (%) from three vegetation types during January.
Pinyon-JuniEer

Ponderosa Pine-Oak
DurangoLaVetaPagosa
Denver
Rye
Springs
Castle rock

Steamboat
Spgs .-Hayden

Oak
SiltCarbondaleEagle

DoloresMancos

MeekerRifle

Naturita

Colo. Spgs.Canon City

5.4

5.0

5.2

4.4

4.4

5.0

5.6

5.6

5.2

5.0

4.8

5.6

4.4

4.4

5.2

4.8

5.2

5.6

4.8

4.6

4.8

4.6

5.0

5.0

5.4

4.4

5.2

5.6

4.8

6.0

5.0

4.2

5.2

5.0

4.6

5.6

5.6

4.6

5.6

4.8

4.8

5.0

5.0

6.0

5.0

I
N
I-'

I

X (Area)
4.8

5.3

5.4

4.6

4.6

5.1

5.2

5.2

X (Vegetation Type)
5.2

Total Mean

= 5.0

Total Median

= 5.2

4.8

5.2

5.3

�-22-

Table 3. Essential oil loss due to freeze drying and grinding
paired big sagebrush samples collected during January.
Percent

of 10

Oil Content

Paired
Sample
Number

Distilled
Green

Distilled After
Freeze Drying
And Grinding

1

1.5803

0.8933

-0.0687

2

1.8600

1. 3017

-0.0543

3

1.5100

1. 3288

-0.0122

4

1.0900

0.8789

-0.2111

5

1.0495

0.9307

-0.1188

6

1.2300

1.5508

+0 .3208

7

0.9700

1.0119

+0.0419

8

1.0711

1.1492

+0.0781

9

1.5048

1.0521

-0.4527

10

1.1700

1.0252

-0.1448

x

1. 3036

1.1123

-0.1913

Loss or Gain
Due to Drying
And Grinding

1/

This represents a 14.7 percent loss of essential oils due to freeze
drying and grinding.
It is not significant at the 5 percent significance
level.

�July, 1977

-23JOB PROGRESS

State of

COLORADO
------------------------------

Project

No.

REPORT

Deer-Vehicle

W-125-R-3

Accident

Investigations

Work Plan No.
Job Title
Period

1
Job No. 1, 2, 3, 4, 5 and 6
Deer Vehicle Accidents Statewide
and Methods and Devices to Reduce Them

Covered:

Personnel:

April 1, 1976 through March 31, 1977

Thomas D. I. Beck, Claudia A. Doose, James D. Fleming, Sharon
L. McDonnell, Robert W. Morse, Kenneth R. Kincaid, Dale F. Reed,
Thomas N. Woodard, Area Supervisors and Wildlife Conservation
Officers.

ABSTRAcr
Five methods, devices, or structures related to reducing the number of
deer-vehicle accidents were evaluated or experimentally tested.
These
were highway lighting, underpasses, overpasses, 2.44-m fences and one-way
deer gates, and deer guards. No additional data on the effect of highway
lighting on the rate of deer-vehicle accidents were obtained because of
mild winter conditions.
No major changes occurred with underpass use.
The experimental procedure for manipulating the Dowd deer overpass width
was implemented for the fall season.
Twenty-five and 19 crossings,
occurring during control (4.93 m) and variable (2.48 m) widths, respectively,
were reviewed on the video surveillance system. Although preliminary, no
major behavioral differences were detected.
The average reduction of deervehicle accidents adjacent six 2.44-m fences was 75.8 percent.
No data
were obtained on deer guard prototypes V and VI.

��-25-

DEER-VEHICLE ACCIDENTS STATEWIDE
AND METHODS AND DEVICES TO REDUCE THEM
Dale F. Reed and Thomas

P.N.O.

D. I. Beck

OBJECTIVE

Locate and examine potentially critical deer-vehicle accident areas in
Colorado and recommend methods or structures which may reduce deer-vehicle
accidents in these areas.
Subsequently, measure the effects of methods
recommended and investigate deer responses to various experimental
structures.

SEGMENT
Job 1 - Monitoring
statewide.
1.
2.
3.

potentially

2.
3.
4.

2.

accident

of highway

lighting

on number

responses

of deer to highway

areas

of deer

underpasses.

Measure deer use of underpasses and the extent
reluctance associated with their use.

Job 4 - Evaluation
1.

deer-vehicle

Measure and describe the luminance on an illuminated 1.I-km
segment of Colorado Highway 82.
Determine if deer-vehicle accidents are affected by the fixed
illumination on the l.l-km segment of Colorado Highway 82.
Investigate and compare responses of motorists to deer on the
highway, with and without fixed illumination.
Investigate and compare deer responses to motorists, with and
without fixed illumination.

Job 3 - Evaluate
1.

critical

Locate and examine potentially critical deer-vehicle accident
areas in Colorado.
Recommend methods or devices which may reduce the problem for
these critical accident areas.
Select recommended underpasses, overpasses, 2.44-m fence
lengths, or other devices or methods for evaluation when sample
size considerations
can be met.

Job 2 - Evaluating the effects
killed by vehicles.
1.

OBJECTIVES

of deer responses

to highway

of behavioral

overpasses.

Determine the use of overpasses by deer in critical highway
deer kill areas.
Compare detailed deer behavioral responses to overpasses after
moderate to high use of the structures or immediate areas has
been documented.

�-26-

Job 5 - Evaluate deer responses
to highways.
1.
2.

(8-ft) fencing adjacent

Determine the reductions in deer-vehicle accidents on highways after the installation of various lengths of 2.44-m fence.
Measure deer movements in relation to 2.44-m fences.

Job 6 - Evaluating
1.

to 2.44-m

responses

of deer to guards.

Compare deer responses to a black-and-white
scintillation
and roll-bar (VI) deer guard prototypes under controlled
conditions.

DESCRIPTION
Job 1 - Monitoring
wide.

potentially

Highway
Pojar

OF AREAS

critical

(1972a) and Woodard

Woodard

(1975) described

this area.

(1975) described

this area.

Job 2 - Evaluating the effects
by vehicles.

Canyon

this area.

this area.

50 - Salida East

(1976a) described

~ighway

this area.

550 - Colona-Ridgway

(1976a) described
Highway

Woodard

(1973b) described

6-24 (1-70) - Eagle-Glenwood

Highway
Woodard

(1973a, 1974) described

82 - Glenwood~Basa1t

(1969) and Woodard
Highway

this area.

70 - Avon-Wolcott

(1972a) and Woodard

Myers

areas state-

West

(1973a, 1975) described

(1972a) and Woodard

Highway

accident

70 - Eagle East

Interstate
Pojar

deer-vehicle

6-24 (1-70) -Rifle

Interstate
Pojar

(V)

this area.

of highway

lighting

82 - Jammeron

Curve

Pojar et al. (1975) described

this area.

on number

of deer killed

�-27-

Job 3 - Evaluate

responses

of deer to highway

underpasses.

A small structure under 1-70 east of Avon, three concrete box
underpasses under 1-70 east of Eagle, a single-span structure
at Chaffee Gulch north of Ridgway, the arch (structural plate
pipe arch) deer underpass west of Durango, three underpasses
(two were large open structures) southeast of Salida, two
bridge-span underpasses east of Rifle, and a concrete box underpass west of Rifle were monitored.
Job 4 - Evaluation

of deer responses

to highway

Dowd Junction

1-70

Reed (1976a) described

this area.

Job 5 - Evaluate deer responses
highways.

to 2.44-m

Highway
The Highway

overpasses.

(8-ft) fencing adjacent

82 - Diamond

S

82 study area has been described
Highway

by Pojar

(1972b).

82 - Carbondale

This study area was described
Interstate

to

by Reed (1975).

70 - Avon, Edwards,

Vail, and Eagle

The Interstate 70 Avon and Edwards study areas have been described
by Woodard (1973b), the Vail study area by Reed (1971), and the
Eagle study area by Reed (1974).
Cherry Creek-Hesperus
The Cherry Creek fence is 1.6 km long and adjacent to both sides of
Colorado 160. The east end of the fence is 9.8 km west of Hesperus
and an underpass goes beneath the highway 0.8 km from either end of
the fence.
Job 6 - Evaluating

responses

of deer to guards.

Trail Gulch-Dotsero
Reed et al. (1974) described

this area.

METHODS AND MATERIALS

Job 1 - Monitoring
wide.

potentially

critical

deer-vehicle

accident

areas state-

Methods have been described by Pojar (1972a) and Woodard (1976a).
Because no new critical areas were delineated, methodology was
limited to tabulation of statewide Wildlife Conservation Officer
kill reports.

�-28-

Job 2 - Evaluating the effects
killed by motorists.

of highway

lighting

on number

of deer

Methods were described by Woodard (1976b).
Luminance measurements were taken at kill sites from earlier years.
Luminance
values (foot-lamberts) were recorded with a spotmeter, Spectra
Model DBA, and were taken on a target at the kill site and on
the background as viewed by an approaching motorist.
The target
was a taxidermy mount of the left half (transverse section) of
a female mule deer and the actual spotmeter recording area was
a 26-cm diameter circle midway between the shoulder and hip.
Spotmeter readings were taken from a height of 1.3 m and a
distance of 15 m. For background luminance readings the spotmeter was placed 60 m from the kill site, resulting in a target
area diameter of 102 cm. The taxidermy mount was removed during
this background measurement.
Measurements were taken between
0400 and 0600 to minimize traffic interference.
Background
luminance (Lb) and target luminance (Lt) measurements were transformed into a visibility index (VI) by the following equation
(Gallagher and Meguire 1974):
C(RCS Lb) (DGF)
VI

=

5.74

where C

Lb-Lt
Lb

Relative contrast
sensitivity for the
recorded background
luminance
DGF

Disability glare
factor = 1.0

RCS values were obtained from standard tables (Gallagher and Meguire
1974).
VI values can then be compared to standards established for
urban lighting requirements.
These values also provide a better
description of the visibility of deer in the lighted portion of the
study area; which, in light of the poor horizontal uniformity ratio
in the lighted section (&gt;15.0:1), should allow a more refined
analysis of the effectiveness of highway lighting.
Job 3 - Evaluate

responses

of deer to highway

underpasses.

Trackbeds were established at 12 underpasses and monitored for deer
use as described by Reed (1971).
Observations at the Vail deer
underpass were continued as described by Reed (1976b).
Job 4 - Evaluation

of deer responses

to highway

overpasses.

Methods were previously described by Reed (1976a).
In addition, the
bridge (Dowd deer overpass) was modified (deck removed and rebuilt)
so the width of the structure could be varied.
It was designed to
be readily changed from its original width of 4.93 m (control) to
2.48 m (variable) every few days. The width was changed every three
days during most of the fall migration from 30 October to 1
December 1976.

�-29-

Job 5 - Evaluate deer responses
to highways.

to 2.44-m

(8-ft) fencing adjacent

Methods and materials have been described by Pojar (1972)
and Woodard (1976c).
Benefit-cost analysis followed procedures described by Howe (1971).
Costs for which values
were available include initial cost of 2.44-m fence and
associated passage structures.
Estimates of maintenance costs
were not available.
Total cost was reduced by the cost of the
appropriate length of 1.07-m (42-inch) standard right-of-way
fence which was not built because of the 2.44-m fence. Benefits
for which a value was available was the potential savings accrued
by a reduction in deer-vehicle accidents and related vehicle
repairs.
Savings by reduction in personal injury and death were
omitted because of difficulties in obtaining reliable values.
No value was assigned to deer. Therefore, benefit-cost analysis
is essentially from a highway department accounting stance.
All
future benefits were discounted at an annual rate of 5.5 percent.
No monetary values have been assigned for aesthetic considerations,
either positive or negative.
Job 6 - Evaluating

responses

of deer to guards.

Methods were described

by Reed et ale (1974).

RESULTS AND DISCUSSION
Job 1 - Monitoring
statewide.

potentially

Highway

critical

deer-vehicle

accident

areas

6-24 (1-70) - Rifle West

Thirty vehicle-killed deer were documented on 29 km of highway
the Rifle West study area; an increase of 2 over the preceding
3-year mean but markedly less than the 1972-73 kill of 92.
December accidents accounted for 46.7 percent of the total.
Interstate

in

70 - Eagle East

Only 4 vehicle-killed deer were documented along 13.7 km of highway in the Eagle East study area; a decrease of 83.3 percent over
the preceding segment.
This low kill is likely a reflection of
the mild winters of 1975-76 and 1976-77 where few deer were forced
to winter along the highway.
Deer kill on portions of the highway
adjacent to the 7.7-km barrier fence was correspondingly
low.
Interstate

70 - Avon-Wolcott

Twenty-eight vehicle-killed deer were documented on 22.5 km of
highway in the Avon-Wolcott study area; a decrease of 55.3 percent
from the preceding 3-year mean (62.7).
Deer killed continued to be
reduced on the portions of highway opposite two sections of 2.44-m
fence.

�-30-

Highway

82 - Glenwood-Basalt

Twenty-six vehicle-killed
deer were documented on 29 km of highway in the Glenwood-Basalt
study area; a decrease of 66.9 percent from the preceding 3-year mean (78.7).
Thirty-one percent
were killed during April.
Deer-vehicle accidents continued to
be reduced on the portions of highway opposite two 1.77-km lengths
of 2.44-m fencing.
Highway

6-24 (1-70) - Eag1e~Glenwood

Canyon

Eight vehicle-killed deer were documented on Highway 6-24 from
Eagle to the west 9.9 km to Gypsum.
The preceding segment kill
was also 8. Fifteen vehicle-killed
deer were documented on
Highway 6-24 from Gypsum to the west 15.0 km to the east end of
Glenwood Canyon.
The preceding segment kill on this stretch was
16. Recommendations
for structures to reduce deer-vehicle accidents
when Interstate 70 is constructed between Gypsum and Glenwood
Canyon were submitted to the Division of Highways.
Recommendations
called for 2.4 km of 2.44-m fence on the north side of the alignment west of Dotsero.
Highway

550 - Colona-Ridgway

Twenty-two vehicle-killed
deer were documented along 11.5 km of
Highway 550 between the Owl Creek road south of Dallas and the
Uncompahgre River bridge south of Colona, a decrease of 29 percent
from the 1974 and 1975 mean of 31.0. No highway construction was
initiated in this study area during this segment.
Highway

50 - Salida

East

Ten deer-vehicle accidents were documented on this 8.4 km section
highway during 1976; a decrease of 4 from the previous segment.

of

Statewide
A total of 1,859 deer-vehicle accidents were documented in Colorado
in 1976; an increase of 21.7 percent over the preceding segment.
Regional totals were Northeast 308, Southeast 206, Northwest 564,
Southwest 781.
Job 2 - Evaluating the effects
by vehicles.

of highway

lighting

on number

of deer killed

There were no estimated crossings or kills during this segment.
No deer were observed adjacent the study area during January-March
1977. The probable reason for the lack of deer in 1977 was the mild
weather conditions.
Although mean daily temperatures for this
period (-2.30C) were slightly higher than the previous 3-year mean
(-2.70C), the major difference was the lack of snow. Snow depth

�-31-

measurements at three sites immediately northeast of the study
area averaged 46.4 cm, 27.5 cm, and 8.0 cm for the JanuaryMarch periods of 1975, 1976, and 1977, respectively.
Luminance measurements
1975) (Table 1).

were

taken at two kill sites

Table 1. Visibility index measurements
lighting study area.

(from March

from kill sites, Highway

82

Contrast

Relative
Contrast
Sensitivity

Visibility
Index

No.

Background
Luminance (£1)

Target
Luminance

897

0.043

0.0325

0.244

4.1

0.17

899

0.33

0.00325

0.990

14.4

2.484

(£1)

Both visibility index values are lower than mlnlmum levels needed
for a roadway with a 50 mph speed limit; although the 2.484 value
is approaching the level where 85 percent of the motorists can see
a target at satisfactory separation distance (Gallagher and Meguire
1974) •
Job 3 - Evaluate

responses

of deer to highway

underpasses.

Eleven underpasses were monitored for deer use. Moderate to high
numbers of deer passages occurred through three of the underpasses,
one located west of Vail, one east of Avon, and the other southeast
of Salida.
Extremes of openness or tunnel effect were represented
by two of these structures in the study sample.
No important changes
were noted in deer use at eight of the unde rpas ses reported previously
(Reed 1976b:228).
The number of passages at the Salida East No.1
underpass decreased to a total of 88 during this segment.
This decrease was noted in particular during April when the video surveillance
system (Reed et a1. 1973) was installed and activated.
No deer
passages occurred while the system was activated and consequently
no behavioral response data were collected.
A segment of newly completed Interstate 70 between Silt and Rifle
was opened to traffic on 13 November 1976. Two underpasses,
adjoining 2.44-m fencing at Mamm Creek and Dry Creek, have received
6 and 9 deer passages. respectively.
Daily observations (n=29) were made at the Vail deer underpass from
1 June to 29 June 1976. With 1974, 1975, and 1976 combined, 77.5
percent of 182 groups of deer did not exhibit flight reaction when
westbound vehicles traveled by the area (Table 2). Likewise, of

�-32-

692 instances of vehicles traveling west when deer were near
the underpass entrance and the west bound lanes, 635 did not
elicit flight reaction from 182 different individuals or groups
(range 1-11 deer).
Fourteen vehicles elicited flight reaction
from 1-50 percent of the individuals from thirteen different
groups ranging in size from one to eight deer.
Forty-three
vehicles elicited flight reaction from 51-100 percent of the
individuals from twenty different groups ranging in size from
one to six. Previous to these observations it was hypothesized
that deer at distances of 10 to 20 m from moving traffic were
frequently caused to leave the area. Other observations indicated
that when vehicles stopped at or when bicyclists traveled by the
underpass, most deer elicited immediate and intense flight reaction.
On three occasions during 1976, several groups of deer exhibited
moderate to intense flight reaction to unknown stimuli (where
observer was unable to determine a potential cause).

Table 2. The number and percent of deer groups not exhibiting flight
reaction (Hediger 1950, Scott 1958), and the number and percent of westbound vehicles not eliciting flight reaction at the Vail deer underpass
west of Vail.

Year

GrouEs of Deer
Percent Not
Number of Individuals
Exhibiting
or Groups 1/ Near
Flight
Westbound Traffic
Reaction

1974
1975
1976

Vehicles
Percent Not
Number Traveling
Eliciting
West When Deer
Flight
Present
Reaction

37

86.5

198

97.5

67

71.6

251

89.2

78.2

2432:,/

89.7

77 .5

692

91.8

78

Totals/Avg.

Westbound

182

1/
Number

of deer in groups range from 2 to 11 animals.

2/
Six semi-trucks

elicited

flight reaction while seven did not.

�-33-

The behavior modes of 163 deer exits (leaving the underpass to
the south) were 54.6, 33.7, and 11.7 percent, trotting, walking,
and bounding, respectively.
These combined data from 1974, 1975,
and 1976 yielded a walk:trot:bound
ratio of 2.9:4.7:1.0 (Table 3).
Wariness, as illustrated by the change in exit mode (trot to
bound) and by the tension stance behavior (front legs spread),
appeared to be exhibited irrespective of the mode of exit in many
instances.
The predominant exit behavior of trotting supports
the continued reluctance (Reed et al. 1975) of deer to use a
structure of this size and character.

Table 3.
underpass

The number of exits observed at the south end of the Vail deer
and the walk:trot:bound
ratio of exit behavior.
Walk:Trot:Bound
Ratio During Exits

Year

No. of Exits
Observed

1974

35

4.0:12.5:1.0

1975

73

3.3: 2.4:1.0

1976

55

1.8: 6.3:1.0

Totals/Avg.

163

2.9: 4.7:1.0

Job 4 - Evaluation

of deer responses

to highway

overpasses.

Deer crossings over the deer overpass were recorded by the video
system for the fall of 1974, spring-summer and fall of 1975, and
the spring-summer of 1976. Replay of 84, 67, 61, and 26 crossing
modes yielded walk:trot:bound
ratios of 7.8:1.8:1.0, 7.6:4.8:1.0,
4.6:2.0:1.0, and 4.0:8.0:1.0 for the fall of 1974, spring-summer
and fall of 1975, and the spring-summer of 1976, respectively.
A mean of 25.3 animals changed their behavior (i.e. walk to trot,
walk to bound, trot to bound, trot to walk, and trot to walk to
trot) during the process of crossing during the spring-summer and
fall of 1975, and the spring-summer of 1976.
The number of selected behavioral responses exhibited by deer and
the calculated behavioral response ratios are summarized in Table
4. There were no significant differences (P&gt;0.50) between the fall
and spring ratios of the number of animals exhibiting behavioral responses (BR ) to the number of crossings, or of the number of instances
of behavior~l responses (BR.) to the number of crossings, for both
muzzle-to-ground
and hesitaEion behaviors.
It was not understood
why few animals exhibited the muzzle-to-ground
behavior during the
spring of 1976.

�-34-

Table 4. The number of selected behavioral responses exhibited by deer
crossing the Dowd deer overpass during four migration seasons, 19741976, and calculated behavioral response ratios.

Behavioral
Response (BR)

Muzzle-to-Ground

No. of
Animals
Exhibiting
BR (BR )
a

No. of
Instances
of BR (BR.)
1

No. of
Crossings Per
BR :C BR. :C BR. :BR
Season (C)
a

1

1

a

11

Fall 1974

25

38

99

0.25:1

0.38:1

1.52:1

Fall 1975

26

38

77

0.34:1

0.49:1

1.46: 1

Spring 1975

38

64

93

0.41:1

0.69:1

1.68:1

Spring 1976

3

9

60

0.05:1

0.15:1

3.00:1

10

23

60

0.17:1

0.83:1

2.30: 1

Fall 1974

19

26

99

0.19:1

0.26:1

1.37: 1

Fall 1975

47

90

77

0.61:1

1.17:1

1.91:1

Spring 1975

26

38

93

0.28:1

0.41:1

1.46 :1

Spring 1976

17

31

60

0.28:1

0.52:1

1.82:1

Low-Head -21
Spring 1976
H'eSltatlon
. -31

11
- Muzzle-to-ground denotes a deer lowering its muzzle to the ground.
2/
- Low-head denotes a lowering of the head where the axis of the neck declines
(posterior to anterior) below the ho r Lzon t a.I .
llCessation of forward movement for 1.0 second or more.
The number of selected behavioral responses exhibited by deer at the
experimental-variable-width overpass and the calculated behavioral
response ratios are summarized in Table 5. No major differences
appear between the ratios of the control and variable for each behavioral response listed. Although preliminary, these results indicate that there may not be an observable difference in the response
of deer to a 2.48-m wide overpass of this size and design characteristic.
Further analysis will be made during the next segment when additional
sample size will be obtained.

�-35Table 5. The number of selected behavioral responses exhibited by deer
crossing an experimental-variable-width
overpass during the fall migration
of 1976 and calculated behavioral response ratios.
Behavioral
Responses (BR)

No. of Animals
Exhibiting BR (BR )
a

No. of Instances
of BR (BR.)
~

BR

:c1/ BR.:
C
~

BR. :BR

a

~

a

2/

Muzzle-to-Ground

-

Control

6

8

0.24:1

o .32: 1

1. 33 :1

Variable ]j

6

7

0.32:1

0.37:1

1.17:1

Control

9

10

0.36:1

0.40: 1

loll: 1

Variable

5

5

0.26:1

o .26 :1

1.00 :1

Control

10

16

0.40:1

0.64:1

1.60:1

Variable

10

17

0.53:1

o .89:1

1. 70 :1

Control

6

10

0.24:1

o . 40 :1

1. 67: 1

Variable

10

14

0.53:1

o . 74 :1

1.40 :1

Control

9

14

0.36:1

0.56:11.56:1

Variable

15

26

0.79: 1

0.37:1

Muzzle-to-Structure

~/

5/

Low-Head -

6/

.
.
H es~tat~on

-

Alert Stance ]j

1.73:1

1/
C denotes the number of crossings during the season.
occurred during control and 19 during variable.
l/Muzzle-to-ground
l/Control

denotes a deer lowering

its muzzle

Twenty-five

crossings

to the ground.

and variable widths were 4.93 m and 2.48 m, respectively.

4/

- Muzzle-to-structure
or raising its muzzle

5/

- Low-head
(posterior
.
-6/C essat~on

denotes a deer lowering
to the bridge railing.

its muzzle

denotes a lowering of the head where
to anterior) below the horizontal.
0f

forwar d movement

to the bridge

deck

the axis of the neck declines

for 1 .0 secon d or more.

l/Alert stance denotes a position where the head and neck are above horizontal
and the ears are erect with the vertical axis of the ear either perpendicular
to horizontal or inclined forward.

�-36-

Job 5 - Evaluate deer responses to 2.44-m (8-ft) fencing adjacent to
highways.
Highway 82 - Diamond S
Deer normally concentrate in crested wheatgrass fields northeast
of the highway in late winter and early spring. Mean number of
deer crossings during March-May periods continued to increase
(Table 6) in spite of the decrease in numbers of deer seen during
spotlight counts. No vehicle-killed deer were documented in the
study area during this period. A summary of fence effectiveness
and benefit-cost analysis data is provided in Table 7.
Table 6. Mean number of deer observed on spotlight counts and mean number
of deer crossings between quarter-mile section markers 25 to 30 on Highway
82 during March-May for 1968 through 1976 (n = number of counts or number
of 24-hour periods).
Mean Mar.-May
Crossings

Year

March

April

Mean Total

1968

134.8
(n=4)

73.0
(n=4)

103.9

1969

151.2
(n=4)

34.0
(n=5)

86.1
(n=9)

1970

104.5
(n=4)

56.0
(n=5)

77 .6
(n=9)

1971

66.8
(n=4)

51.4
(n=5)

58.4
(n-9)

11.7
(n=32)

1972

102.2
(n=4)

4.5
(n=4)

53.4
(n=8)

2.1 J)
(n=38)

1973

137.4
(n=5)

47.0
(n=4)

97.2
(n=9)

5.5
(n=34)

1974

143.5
(n=4)

52.3
(n=3)

104.4
(n=7)

10.3
(n=38)

1975

126.8
(n=4 )

93.0
(n=4)

109·9
(n=S)

17.8
(n=73)

78.0
(n=4)

61.8
(n=4)

69.9
(n=8)

21.0
(n=28)

1976

!/

(n=B)

1.77 km of 2.44-m fence constructed during summer of 1971.

�Table 7.

Benefit-cost analysis of 2.44-m fencing and associated structures.
(2)

Area
(km)

(1)
Accident
Cost in
Year of
Construction

(3)

Mean PreFence
Mortality

Vail
(4.83)

$ 315

36

68.3%

20

Avon
(3.62)

362

28

83.9%

Edwards
(3.62)

362

27

Eagle
(7.72)

383

167

(4)
Expected
Mean
Usable
Annual
Life
Reduction
(Yrs)

(5)

(6)

(7)

(8)

(9)

Actual
Discount
2.44-m
PVB 1/
Rate
Fence
Factor (6=lX2X3X4X5) Costs

Associated
Structure
Cost

Alternate
42"-Fence
Costs

Net Cost
B:C
(10=7+8-9) (6:10)

0.63067

$ 97,694

$ 39,600

$ 21,934

$ 7,633

$ 53,901

1.81:1

20

0.63067

107,266

37,000

0

6,731

30,269

3.54:1

66.7%

20

0.63067

82,230

43,000

0

4,079

38,921

2.11:1

85.8%

20

0.63067

692,205

100,002

0

20,364

79,638

8.69:1

(10)

(11)

pvc'l:../

I

w
"-J
I

Diamond S
(1.77)

342

10

82.0%

20

0.63067

35,373

13,257

0

0

l3 ,257

2.67:1

Carb 82
(1.93)

396

14

67.9%

20

0.63067

47,482

26,914

0

0

26,914

1.76:1

Cherry
Creek
(3.22)

383

2

-316.7%

20

0.63067

-30,599

41,637

26,665

5,120

63,182

-0.48:1

1:./PVB = Present value of benefits.
])
PVC = Present value of costs.

�-38-

Highway

82 - Carbondale

The mean number of deer observed on spotlight counts during
January-April periods of 1968 through 1974 was 15.3 (n=114).
Seventy deer were killed from January, 1968 to April, 1973
(14 deer per year), before the 2.44-m fence was installed.
Nine deer were killed from 17 October 1974 to 17 October
1976 after the fence was installed - a reduction of 67.9 percent. Post-fence installation spotlight counts averaged 17.9
(n=32) deer per count for the January-April periods.
Benefitcost analysis data is presented in Table 7.
Interstate

70 - Avon

The mean number of deer observed on spotlight counts during the
August-December
periods was 15.5 (n=17) and 13.4 (n=21) in 1971
and 1972, respectively; 10.8 (n=ll) in 1973 for October-December;
and 7.3 (n=7), 4.2 (n=8), and 7.4 (n=5) for November-December
in
1974, 1975, and 1976, respectively.
Deer passages to the north
through six one-way gates in the 2.44-m fence totaled 167, a
marked increased from the preceding segment (Table 8) and the
previous 3-year mean (91.3).
Eighteen deer were killed from 5
October 1972 to 5 October 1976 (4.5 per year) after the fence
was installed; a reduction of 83.9 percent from pre-fence installation kills of 28 deer per year.
Benefit-cost analysis
data is presented in Table 7.
Interstate

70 - Edwards

Five vehicle-killed
deer were documented in this study area during
this segment; thus bringing the mean annual kill since 2.44-m fence
installation down to 9.0 (a reduction of 66.7 percent).
Use of
one-way gates continued to be very low with only 2 passages during
this segment (Table 8). Benefit-cost analysis data is presented
in
Table 7.
Interstate

70 - Vail

Mean annual kill in this area since fence installation was 11.4
(n=7); a 68.3 percent reduction from the pre-fence annual kill of
36. Eight vehicle-killed
deer were documented in the area during
this segment.
Benefit-cost analysis data is presented in Table 7.
Interstate

70 - Eagle

Eight vehicle-killed
deer were documented along 7.7 km of 2.44-m
fence during this segment.
Mean annual kill since fence installation is 23.7 (n=3); a reduction of 85.8 percent from the 167 deer
killed in the year preceding fence installation.
Only 10 passages
north through one-way gates were recorded (Table 8), with only 4
to 10 gates receiving any use. Benefit-cost analysis data is
presented in Table 7.

�-39-

Table B. The number of passages through one-way deer gates for 1975
and 1976, and mean annual number of passages through one-way gates
located in four 2.44-m fences.
2.44 Fencing
Location

1975

1976

Mean One-Way
Gate Passages

Vail - both sides
Interstate 70 (1-70)

9

B

44.9 (7,B) !/

Avon - one side
1-70

64

167

7

2

9.0 (4,7) ~/

13

10

21.0 (3,10)

110.3 (4,6)

Edwards - one side

~m
Eagle - one side
1-70

!/
(n,N) n denotes number of years of post-installation data, and N number
of gates. In this case, N was reduced to seven in fall of 1973 and six
in the spring of 1976.

N was reduced to six in spring of 1975.

Based on 119 observations of collared deer north of the 2.44-m
fence, does (n=B5) and bucks (n=34) have moved lateral or parallel
to the fence for mean distances of 0.59 and 0.76 km, respectively.
Based on 19 estimated locations of three radio-collared does north
of the fence between 24 March and 4 May 1976, mean lateral movements were 0.54 km.
Cherry Creek - Hesperus
Twenty-five deer have been recorded killed in this study area since
fence installation (B.33 per year). Compared to the documented
pre-fence kill of 2 deer per year it is apparent that either the
pre-fence installation documentation was poor or that more deer are
now being killed in this area, presumably because of the fence.
Better documentation should be required before proposing future
fencing projects. Benefit-cost analysis data is presented in Table
7.
A word of caution is in order regarding the fence evaluations.
Although substantial reductions in deer losses were realized at all
but the Cherry Creek study area such reductions can not summarily

�-40-

be attributed to fence installation.
The recorded reductions
may also be a function of changes in deer populations, weather
conditions, deer response to the fence, and traffic conditions.
The same can be said for the recorded increase in kill at
Cherry Creek. Additionally, one should bear in mind the weakness of data comparisons separated by time and thus attribute
cause-and-effect cautiously.
Job 6 - Response

of deer to guards.
Trail Gulch - Dotsero

No deer were tested on prototypes V and VI during this segment
because of a lack of test animals.
Trapping conditions were very
poor during this segment.

LITERATURE

CITED

Gallagher, V. P., and P. G. Meguire.
1974. Contrast requi~ements of urban
drivers.
Fed. Highway Admin. Rep. ~umber FIDvA-RD-74-76. 72pp.
Hediger,

H.1950.

vlild animals

in captivity.

Howe, C. W. 1971. Benefit-cost analysis
Geophysical Union.
H"ashington, D. C.

Butterworth,

London.

19pp.

for water system planning.
l44pp.

Amer.

Myers, G. T. 1969. An investigation of deer-auto accidents.
Pages 147-178
in Game Res. Rep., July, Part 2. Colorado Game, Fish and Parks Div. Fed. Aid.
Pojar, T. M. 1972a. Monitor potentiallv critical deer-vehicle accident
areas statewide.
Pages 267-275 in Game Res. Ren., July Part 3. Colorado
Div. Wildl. Fed. Aid.
1972b.
An evaluation of deer-proof fence length required to
prevent deer movements on or across high speed highways.
Pages 305-310
in Game Res. Rep., July, Part 3. Colorado Div. Wildl. Fed. Aid.
, R. A. Prosence, D. F. Reed, and T. N. Woodard.
1975. Effectiveness
of a lighted, animated deer crossing sign. J. Wildl. Manage. 39(1):87-91.

---::---:--

Reed, D. F. 1971. Deer underpass evaluation.
Pages 341-351 in Game Res.
Rep., July, Part 3. Colorado Div. Wildl. Fed. Aid.
1974. An evaluation of 8-foot fence length required to prevent
deer movements on or across high speed highways.
Pages 313-320 in Game
Res. Rep., July, Part 2. Colorado Div. l-Jildl.Fed. Aid.

�-41-

1975. An evaluation of 8-foot (2.44-m) fence length required
to prevent deer movements on or across high speed highways.
Pages
315-320 in Game Res. Rep., July, Part 2. Colorado Div. Wildl. Fed. Aid.
1976a. Evaluation of deer overpasses.
Pages 247-251
Res. Rep., July, Part 2. Colorado Div. Wildl. Fed. Aid.

in Game

1976b. Evaluation of deer underpasses.
Pages 223-229 in
Game Res. Rep., July, Part 2. Colorado Div. Wildl. l&lt;'ed.Aid.

------- , T. M. Pojar, and T. N. Hoodard.
system for wildlife
Leafl. 94. 3p.

deer guards.

surveillance.

1973. A video time-lapse
Colorado Div. Wildl., Game Infor.

____________ ~. 1974. Mule deer responses
J. Range Manage. 27(2):111-113.

to

______
, T. N. Woodard, and T. "M. Pojar.
1975. Behavioral response
mule deer to a highway underpass.
J. Wildl. Manage. 39(2):361-367.
Scott, J. P. 1958.
Chicago.
349pp.

Animal behavior.

2nd ed. University

of

of Chicago Press,

Woodard, T. W. 1973a. Honitor potentially critical deer-vehicle accident
areas statewide.
Pages 163-170 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. l&lt;'ed.Aid.
1973b. An evaluation of deer-proof fence length required to
prevent deer movements on or across high speed highways.
Pages 197-202
in Game Res. Rep., July, Part 2. Colorado Div. Wi1d1. Fed. Aid.
1974. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 293-298 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. l&lt;'ed.Aid.
1975. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 295-303 in Game Res. Rep~, July, Part 2. Colorado
Div. Wildl. Fed. Aid.
1976a. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 215-221 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. Fed. Aid.
1976b. Effects of highway lighting on number of deer killed
by vehicles.
Pages 231-236 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. Fed. Aid.
1976c. An evaluation of deer-proof fence length required to
prevent deer movements on or across high speed highways.
Pages 239-244
in Game Res. Rep., July, Part 2. Colorado Div. Wildl. Fed. Aid.

e-"?"

Prepared

by

I:-..
ct-CZ~

--··-;~(lfu

- -./

Dale F. Reed

.'

Thomas D. I. Beck

l.HjY1

��July, 1977

-43-

JOB FINAL REPORT

State of
Project

~C~o~l~o~r~a~d~o~ _
No.

W-38-R-32

Work Plan No.
Job Title

14

Job No.

Middle Park Cooperative

Period Covered:
Personnel:

Deer-Elk

Sa

Deer Study - Experimental

April 1, 1976 to March

Len Carpenter, Patrick
Patrick Firlick.

Investigations

Range Fertilization

31, 1977

Powell,

Glenn Crouch, Wayne Regelin,

and

ABSTRACT
This study evaluated residual effects of nitrogen and 2,4-D herbicide on
deer forage species within big sagebrush communities 6 years after the
initial treatment year.
No significant (P&lt; 0.05) yield responses were
measured as a result of nitrogen fertilizer treatment after 6 years.
However, significant (P&lt; 0.01) responses were measured with yields of total
herbage, shrub herbage and grass herbage as a result of herbicide treatments.
Over the 6-year period the highest level of nitrogen fertilizer (134.4 kg
N/ha) was the most efficient in terms of kilograms of herbage produced per
kilogram of nitrogen applied.
Largest yields were obtained on plots treated
with the highest level of nitrogen in combination tiith 2,4-D herbicide.
Plots
treated with 33.6, 67.2, and 100.8 kg N/ha without herbicide produced lower
yields of vegetation than did control plots.
It is recommended that this
depression of yield on nitrogen treated plots be evaluated further.

�-44-

RECOMMENDATIONS
1.

On sagebrush winter
applied for maximum
ment.

ranges, large rates (134 kg N/ha) should be
results of a single nitrogen fertilizer treat-

2.

Combination treatments of 2,4-D herbicide and nitrogen fertilizer are
superior to single applications of either chemical in improving sagebrush-dominated winter ranges.

3.

Long-term (10 year) evaluations are needed to fully evaluate
effects (both good and bad) of these chemical treatments.

lasting

�-45-

EXPERIMENTAL

RANGE FERTILIZATION

Len H. Carpenter

With the tremendous increase in cost of nitrogen fertilizers today as
compared to 5 years ago, it becomes increasingly important to know how
long the residual yield effect of a single application of nitrogen fertilizer will be. If a significant yield response can be obtained over a
period of years (5-7), the initial cost of the fertilization project can
be prorated over that period of time resulting in a much better cost-benefit
ratio.
Various workers have reported on the residual yield responses of vegetation
to a single application of nitrogen fertilizer on rangelands (Hubbard and
Mason 1967; Bowns 1972; Baldwin et al. 1974).
In general, it appears that
the number of growing seasons that vegetation will respond to a single application of nitrogen is mostly dependent upon the rate of the initial application.
Lower rates result in shorter response periods with higher rates giving yield
responses for longer periods of time.
The evaluation period following an application of nitrogen is usually for 2-3
years (Kilcher et al. 1965; Johnston et al. 1968; Mason and Miltimore 1959).
However, Mason and Miltimore (1972), working in southern British Columbia,
reported on 10 year yield responses of beardless wheatgrass (Agropyron inerme)
to a single application of nitrogen.
Their results showed that considerable
increases in yields of native grassland could be obtained for period of ten
years with all rates of application (28,56 ••• to 504 kilograms actual nitrogen per hectare).
Even in the tenth year after application of the fertilizer,
there was still a large yield increase with the highest nitrogen application
rate producing 2.2 times the zero rate.
These results encourage the investigation of long term responses to fertilizer treatments.
In October 1969, a single application of ammonium nitrate
fertilizer at the rate of 33.6, 67.2, 100.8, and 134.4 kilograms actual
nitrogen per hectare was applied with and without 2,4-D herbicide (2.2 kg/ha)
to three permanently located study areas in northcentral Colorado (Carpenter
1970).
The dominant vegetation on these areas was big sagebrush (Artemisia
tridentata).
The yield response of vegetation to these treatments were
evaluated for 3 years after treatment during the growing seasons of 1970, 1971,
and 1972 (Carpenter 1970, 1971, and 1972).
Yields of. vegetation responded in a linear fashion to increased levels of
nitrogen in all evaluation years.
In all years, this yield response was
Significant (P&lt; 0.05).
Certain yield responses to combination treatments
of nitrogen fertilizer and 2,4-D herbicide were also interesting with regard
to improving mule deer ranges.
To better evaluate the management potential of these chemical treatments,
longer term yield measurements were needed.
Yield measurements were taken
during the 1976 growing season, to aid in the evaluation of the long-term
effects of a single application of nitrogen fertilizer applied with and

�-46-

without 2,4-D herbicide on native sagebrush rangeland.
This study evaluated
the residual effects of nitrogen and 2,4-D on deer forage species with big
sagebrush communities 6 years after the initial treatment year.
Descriptions
of all areas, methods, and procedures pertinent to this study are detailed
by Carpenter (1976).

P.N.O. OBJECTIVE
To measure the effects of a single application of nitrogen fertilizer and
2,4-D herbicide (applied singly and together) on the production and composition of mule deer forages on sagebrush winter range sites.

METHODS

AND MATERIALS

This study evaluated residual effects of nitrogen and 2,4-D on deer forage
species within big sagebrush communities 6 years after the initial treatment
year.
Estimates of herbage yield for the 10 treatments at each of 3 study areas
were made with an electronic capacitance meter.
This instrument, when placed
in vegetation, measures the capacitance or ability of vegetation to absorb
energy from the head-produced electromagnetic
field. As the mass of vegetation increases, the ability to absorb the produced energy increases in a
linear relationship.
Therefore, capacitance is closely related to mass or
weight of the vegetation monitored (Currie et al. 1969). Dimensions of this
meter were 30.5 x 61.0 x 45.7 cm, width, length, and height, respectively.
All vegetation within this hexahedron was sensed.
A more detailed presentation of this instrument is given in Neal and Neal (1973).
A double sampling system was employed where one 30 x 61 cm quadrat was
estimated with the meter and clipped for every five that was meter-estimated
only.
Past experience (Carpenter 1972) has shown that this 5:1 ratio is most
efficient in terms of obtaining a yield estimate that is within 10 percent of
the true population mean, 95 percent of the time.
In late July and early August 1976, 75 quadrats were estimated on each 30 x 30 m
treatment of which 15 were both meter-read and clipped.
The location of these
quadrats was chosen from a grid system in a random fashion to insure total
coverage of each treatment plot.
The first quadrat to be clipped on each
treatment plot was randomly chosen from the numbers one through five, and
thereafter every fifth quadrat was systematically clipped after it was read
by the meter.
The procedure for clipping vegetation within a quadrat was as follows:
first,
vegetation immediately outside the probes of the meter that was not part of
the quadrat, and not sensed, was clipped and removed.
A meter reading was
taken and recorded of the total vegetation on the quadrat.
The meter then was
removed and taken to the next quadrat.
The leaves and stems of all shrubs
were clipped and sacked separately by species.
Current annual growth of this
material was separated in the laboratory from the older, woody portion.
Once
the shrubs were clipped, all forbs and grasses on the quadrat were clippeg
and sacked separately.
Yield estimates were computed on an oven-d,ry (100 C
for 24 hours) basis.
All samples were weighed to the nearest one-tenth gram.

�-47-

Clipped weights of total herbage and shrub herbage were regressed against
the meter readings.
A computer program for double sampling (DUB SAM 2) l/
to estimate weight for all meter-estimated
quadrats for both total yield
and shrub herbage yield was developed.
Past experience (Carpenter 1972)
has shown that at each study area regressions are very similar among liketreated plots so that it is possible to combine them into larger samples
to provide more accurate predictive equations.
Thus, data were combined
for the four fertilizer plots (plus a control) and the five herbicide plots
to give a small sample of 75 and a large sample of 375 for each five treatment combination.
The resulting predictive equations were then used to
estimate total herbage and shrub herbage on each treatment at each study
area.
The ten yield estimates for each weight variable were analyzed by
covariance where each value was adjusted to 1969 pre-treatment conditions.
In previous work, meter readings did not correlate with weight of grasses
and forbs (Carpenter et al. 1973). Apparently variability of understory
components, coupled with the influence of woody material in the overs tory
on each quadrat, render these correlations impossible.
Therefore, yield
estimates for forbs and grasses were obtained from only the 15 clipped
quadrat samples on each treatment.
Analysis of covariance was used where
the 1966 values for both forbs and grasses were adjusted to 1969 pre-treatment conditions.

Data Analysis
The experimental design for this study was a randomized block replicated
times with a 5x2 factorial of fertilizer and herbicide treatments.
The
source table for analysis of covariance of herbage yield data for this
experiment was as follows:
Source

Combination

3

df

Replications (~
Nitrogen (A)
Herbicide (B)
RA
RB
AB
RAB

2
4
1
8
2
4
8

Total

29

of sources used for error term RA + RB + RAB

= 17.

Values for nitrogen fertilizer efficiency were also determined.
These values
were calculated for both total herbage and shrub herbage by adding the yields
obtained on each nitrogen treatment without herbicide for the 4 evaluation
years (1970, 1971, 1972, and 1976) and dividing by the proper nitrogen rate,
i.e. 33.6, 67.2, 100.8, or 134.4.
These values provided a measure of the
residual yield responses for each nitrogen treatment.

l/This program, developed by Dr. J. K. Kovner, Principal Biometrician, is
on file at the Rocky Mountain Forest and Range Experiment Station, Fort
Collins, Colorado.

�-48-

DESCRIPTION

OF AREA

Field work was done in Middle Park in north central Colorado approximately
161 km west of Denver.
Vegetation in Middle Park is predominantly big
sagebrush (ArtemiSia tridentata tridentata) with bunchgrasses as understory
at elevations below 2,743 m and aspen-conifer forests above that elevation.
The climate of Middle Park is characterized by cool summers and cold winters.
Izett (1968) reported an average annual precipitation in Middle Park of 53 cm.
Three areas situated on big sagebrush range were chosen for this study.
The areas were designated by the following names: Junction Butte (Area 1),
Flightline (Area 2) and Corral Creek (Area 3). The Junction Butte area was
on land owned and administered by the State of Colorado, Division of Wildlife.
The Flightline and Corral Creek study areas were on public land administered
by the Bureau of Land Management.
Junction

Butte Study Area

This area represents critical mule deer range in that it is in the zone of
most concentrated use in late winter.
It is located approximatly 6 km southeast of Kremmling, Grand County, Colorado (NE 1/4, Sec. 28, TIN, R80W) on the
toe of an eight percent slope that extends out below steeper slopes with
shallow soils and rock outcrops.
The aspect is southwesterly, and, as a
result, the study area is often free of snow during winter periods.
Elevation
is approximately 2,380 m.
The soil profile of the area is identified as a Tigiwon sandy loam, a member
of the fine loamy, mixed-over, sandy skeletal family of Corollic Haplargids.
The soil has an A horizon of less than 15 cm, a B horizon of clay accumulation,
and horizons of lime accumulation below the B-2-t horizon.
It is well drained
with permeability of between 2 and 3 cm per hour. Annual precipitation is
about 28 cm of which one-half occurs during the growing season.
The dominant plant species on the area is big sagebrush.
Common forb species
present are bluebell (Mertensia spp.) and phlox (Phlox spp.).
The major grass
species present are bluebunch wheatgrass (Agropyron spicatum), blue grama
(Bouteloua gracilis), needle-and-thread
grass (Stipa comata), and pine needlegrass (Stipa pinetorum).
Vegetation is sparse, probably due to prior management and poor soil and moisture conditions.

Flightline

Study Area

This area is located approximately 27 km northeast of Kremmling, Grand County,
Colorado (SW 1/4, Sec. 36, T2N, R79W).
It represents early winter range for
mule deer.
The study area is located on gently sloping uplands at an elevation
of 2,380 m. Snow depths exceeding 46 cm occur at this area and snow cover
often remains until late April or early May.
The soil profile at this area is identified as an unnamed fine, sandy loam.
This soil is a member of the fine Montmorillonitic
family of Argic Cryoborolls.
It has a dark-colored A horizon up to 25 cm in thickness and a distinct B
horizon of clay accumulation.
Definite horizons of lime accumulation exist
below the B-2 horizon.
The soil is well drained with a permeability between
0.5 and 1.5 cm per hour.
Annual precipitation is approximately 36 cm with
about one-half occurring during the growing season.

�-49-

Big sagebrush is the dominant plant species on the area. Other shrub
species important to this area are green rabbitbrush (Chrysothamnus
viscidiflorus) and bitterbrush (Purshia tridentata).
Important forb species
are bluebell, phlox spp., and buckwheat (Eriogonum umbellatum).
The major
grass species are bluebunch wheatgrass, squirreltail (Sitanion hystrix), pine
needlegrass, and mutton grass (Poa fendleriana).
Prolific plant growth
characterizes this area.

Corral Creek Study Area
This area is located approximately 29 km northeast of Kremmling, Grand
County, Colorado (NE 1/4, Sec. 36, T2N, R79W) at an elevation of 2,500 m.
This area is considered representative of intermediate winter range for mule
deer. The study area is on a mountainside of westerly aspect and average
slope of about six percent.
Snow depths are intermediate to the Junction
Butte and Flightline areas.
The soil profile at this area was identified as an unnamed loam.
This soil
is a member of the fine Montmorillonitic
family of Vertic Cryoborolls.
It
has a dark-colored A horizon up to 31 cm in thickness with weak B-2 horizons.
The soil is well drained with permeability of between 0.2 and 0.5 cm per hour.
Annual precipitation is about 36 cm with about one-half of this amount received during the growing season.
Big sagebrush is the dominant plant species on this area.
Dominant forb
species are bluebell, phlox spp., and buckwheat.
However~ other forb species
such as paintbrush (Castilleja spp.), clover (Trifolium gymnocarpon), and
vetch (Astragalus drummondii) are common.
Major grass species are bluebunch
wheatgrass, western wheatgrass (Agropyron smithii), mutton grass, and
squirreltail.
Vegetaton is more diverse than on the other two areas.

RESULTS AND DISCUSSION

Main Effects of Nitrogen

Fertilizer

No significant (P&lt; 0.05) reponses in yield were measured in 1976 for forbs,
grasses, shrubs or total herbage in response to nitrogen fertilizer.
This
contrasted to previous results at these study areas (Carpenter 1976) where
nitrogen fertilizer produced significant yield increases for three years after
treatment.
Unadjusted yields of shrub and total herbage for 1969 (pre-treatment),
and adjusted values for all evaluation years as a function of increasing levels
of nitrogen are graphed in Figure 1. The small yield response to fertilizer
in the 1976 measurements, as contrasted to previous measurements, is obvious.
It would appear from this graph that 1976 was a poor year for plant growth.
Yields on control plots (no treatment) in 1976 were less than yields on control
plots in all previous years (Fig. 1). No explanation for this is known.
A
summary of precipitation records for 1976 did not differ significantly from
previous measurement years or from l5-year averages in amounts or in time-ofyear the precipitation was received.

�-50-

TOTAL

HERBAGE
/.1970
/
/

.....e-- __

--./

./"

_.r-"

.
.
..
.
....
-. - -: .,..a...:.~':-»:"

".

*"""

•••

.""

-

••0 •• '/'
• • •

~

_--0

• ••

••••••• .••

/

/

1972
./.:~~1971

/

. .,1"

0

... .,....,- .- ..P----.A1969

1976

SHRUB

HERBAGE
.1970
/

.... ..01971

//
/

.
-

•

/

,...--... __
---0'

..
/

.".,0-- -_

.•
•••
----.----- .-•....
-~ 50~------~------------~
./

.",.",......•...:-...0 ..•'

Q

••••

-

o

.&gt;:

»&lt;&gt;:

/

".,

~,

--.

/

.:&gt;

,,-1972
~~

./

'

33.6

67.2

NITROGEN

1969
1976

100.8

134.4

LEVELS (kgl ha)

Figure 1. Total herbage yields and shrub herbage yields for 1969 (pretreatment) and for 4 separate years after treatment in response to increasing levels of nitrogen. Means for 1970, 1971, 1972, and 1976 are
adjusted by covariance to 1969 pre-treatment conditions.

�-51-

For the three-year period (1970-1972) the average total herbage yield of
untreated plots for all three study areas was 1,278 kg/ha and 1,805 kg/ha
on the 134.4 kg N/ha treatments (Table 1), an average increase of 41 percent.
In 1976 the average total herbage yield on untreated plots was 816
kg/ha and 881 kg/ha on the 134.4 kg N/ha treatments (Table 2), an average
increase of only 8 percent.
Plots receiving 33.6, 67.2, and 100.8 kg N/ha produced less total herbage
in 1976 than did the control treatments (Table 2). This contrasted to
measurements made for the 1970-1972 period where a linear increase in total
herbage yield was obtained with each increasing level of nitrogen fertilizer
(Table I). The small response to nitrogen fertilizer in 1976 is further
demonstrated by comparing the adjusted fertilizer effects in Table 2 to the
adjusted fertilizer effects in Table 1.
Yields of shrub herbage in 1976 responded to nitrogen fertilizer in a manner
similar to total yields (Fig. 1). The average yield on untreated plots for
all three study areas combined was 398 kg/ha and 491 kg/ha on 134.4 kg N/ha
treatments (Table 3), an average increase of 23 percent.
This contrasted to
shrub yields measured during 1970-1972 that were 74 percent greater on 134.4
kg N/ha treatments as compared to control plots (Table 4). However, as
with total yields, the intermediate levels of nitrogen fertilizer (33.6, 67.2,
and 100.8 kg N/ha) produced less shrub herbage than did the controls.
Yields of forbs were not significantly (P&lt; 0.05) changed by the addition of
nitrogen fertilizer in any of the previous measurements years or in 1976.
This is clearly shown by the adjusted fertilizer effects in Tables 5 and 6.
Forb yields in 1976 were about one-half of average forb yields measured during
the 1970-1972 period, another indication of poor vegetative growth in 1976.
Grass yields measured in 1976 (Table 7) were not significantly
(P&lt; 0.05)
changed by the addition of nitrogen fertilizer.
This contrasted to previous
measurements (Table 8) where significant inereases in grass yields were
obtained with nitrogen fertilizer.
All grass yields in 1976 were lower than
those of previous years, but the decrease was not as great as that measured
with shrubs and forbs.

Main Effects

of 2,4-D Herbicide

Significant (P&lt; 0.01) yield responses were measured for total herbage in
1976 as a result of herbicide application.
In contrast to previous years
(Table 1) where the response to herbicide was a decrease in total yield, in
1976, the response was an increase in total yield (Table 2). A large portion of this increase was the result of grass responses to the herbicide
(Table 7). Herbicide treated plots produced 18.6 percent more total herbage
than did control plots in 1976 (Table 2). Six years after treatment the
combination treatments of nitrogen fertilizer and 2,4-D herbicide produced
the greatest yields of any treatments, with the largest yields being produced
on the plots receiving the highest level of nitrogen with herbicide.

�Table 1. Average yield of total herbage (kg/ha) for three-year period 1970-1972. Values are means
for three study areas as adjusted by covariance for 1969 pre-treatment conditions.

2,4-D Herbicide

(kg/ha)

o

Nitrogen Fer~i=l=i=z=e~r~(k~g~/h~a~)~
_
33.6
67.2
100.8
134.4

Adjusted
Herbicide
Effects 1/

o

1,278

1,388

1,482

1,528

1,805

1,496

2.2

1,164

1,272

1,398

1,298

1,411

1,309

Adjusted Fertilizer
Effects 1/

1,221

1,330

1,440

1,413

1,608

l/Due to covariance analysis, adjusted effects are not means of the given cell values.
I

Table 2. Average yield of total herbage (kg/ha) for 1976.
adjusted by covariance for 1969 pre-treatment conditions.

Values are means for three study areas as

Adjusted
Herbicide
Effects 1/

o

Ni trogen Ferti1izer~(&gt;..:.k:.c:gu./~h..:::aL.)
_
33.6
67.2
100.8
134.4

o

816

741

791

776

881

801

2.2

969

886

954

1,047

984

968

Adjusted Fertilizer
Effects 1/

892

814

872

911

932

2,4-D Herbicide (kg/ha)

l/Due to covariance analysis, adjusted effects are not means of the given cell values.

V1
N
I

�Table 3. Average yield of shrub herbage (kg/ha) for 1976. Values are means for three study areas
as adjusted by covariance for 1969 pre-treatment conditions.

_
134.4

Adjusted
Herbicide
Effects 1/

o

Nitrogen Fertiliz_e_r
__(~k~g~/_h_a~)
33.6
67.2
100.8

o

398

327

384

385

491

397

2.2

312

234

317

354

303

304

Adjusted Fertilizer
Effects ]j

355

280

350

369

397

2,4-D Herbicide (kg/ha)

l/Due to covariance analysis, adjusted effects are not means of the given cell values.
I
\J1
W

I

Table 4. Average yield of shrub herbage (kg/ha) for three-year period 1970-1972.
three study areas as adjusted by covariance for 1969 pre-treatment conditions.

Values are means for

0

Nitrogen Fertilizer (kg/ha)
100.8
67.2
33.6

l34.4

Adjusted
Herbicide
Effects 1/

o

618

673

805

820

1,072

798

2.2

330

385

504

410

511

428

Adjusted Fertilizer
Effects 1/

474

529

655

615

791

2,4-D Herbicide (kg/ha)

l/Due to covariance analysis, adjusted effects are not means of the given cell values.

�Table 5. Average yield of forb herbage (kg/ha) for 1976.
adjusted by covariance for 1969 pre-treatment conditions.

Values are means for three study areas as

_
134.4

Adjusted
Herbicide
Effects ))

84

115

114

98

54

124

83

105

69

120

o

Nitrogen Fertiliz~~r~(~k~g~/h==aL)
33.6
67.2
100.8

o

129

128

112

2.2

78

61

Adjusted Fertilizer
Effects 1./

103

95

2,4-D Herbicide (kg/ha)

l/Due to covariance analysis, adjusted effects are not means of the given cell values.
I
VI
.l&gt;I

Table 6. Average yield of forb herbage (kg/ha) for three-year period 1970-1972.
three study areas as adjusted by covariance for 1969 pre-treatment conditions.

Values are means for

Adjusted
Herbicide
Effects 1/

o

Ni trogen FertHi zer (.=.:k£2g~/h=a)c-_
33.6
67.2
100.8
134.4

o

267

249

253

207

305

256

2.2

169

146

205

155

117

158

Adjusted Fertilizer
Effects 1/

218

197

229

181

211

2,4-D Herbicide (kg/ha)

l/Due to covariance analysis, adjusted effects are not means of the given cell values.

�Table 7. Average yield of grass herbage (kg/ha) for 1976. Values are means for three study areas
as adjusted by covariance for 1969 pre-treatment conditions.

134.4

Adjusted
Herbicide
Effects 1./

279

255

288

438

614

645

582

365

446

450

0

Nitrogen Fertilizer (kg/ha)
100.8
67.2
33.6

o

312

304

291

2.2

521

693

Adjusted Fertilizer
Effects 1./

416

499

2,4-D Herbicide (kg/ha)

!/Due to covariance analysis, adjusted effects are not means of the given cell values.
I
V1
V1
I

Table 8. Average yield of grass herbage (kg/ha) for three-year period 1970-1972.
three study areas as adjusted by covariance for 1969 pre-treatment conditions.

Values are means for

Adjusted
Herbicide
Effects 1./

o

Nitrogen Fertilizer (kg/ha)
33.6
67.2
100.8

o

361

544

430

562

467

473

2.2

547

717

703

835

901

740

Adjusted Fertilizer
Effects !/

454

630

567

698

684

2,4-D Herbicide (kg/ha)

134.4

!/Due to covariance analysis, adjusted effects are not means of the given cell values.

�-56-

Shrub herbage as measured in 1976 continued to be significantly
(P&lt; 0.01)
reduced by herbicide.
This compared favorably to previous measurements.
The reduction was not as great as previously measured.
For the three-year
period 1970-1972 there was a 46 percent reduction in shrub yields on herbicide treated plots as compared to plots receiving no herbicide (Table 4).
In 1976 there was only a 23.4 percent reduction in shrub herbage on herbicide treated plots (Table 3). This may indicate partial recovery of
herbicide treated plants over the six year period.
There was no significant (P&lt; 0.05) response in yields of forbs to herbicide
treatments in 1976. This contrasted to previous measurements where forb
yields were significantly reduced by herbicide.
However, there was still
a 27 percent reduction in forb yields on herbicide treated plots as compared
to plots receiving no herbicide (Table 5). For the three-year period 19701972 there was a 38 percent reduction.
Apparently, sample sizes were not
adequate in 1976 to detect this difference.
However, this may be an indication that forb species are recovering on herbicide treated plots.
Grass yields in 1976 were significantly
(P&lt; 0.01) increased as a result of
herbicide treatments.
The increase in grass yields on herbicide treated
plots exceeded grass yields on plots receiving no herbicide by 102 percent
in 1976 (Table 7). This compared to a 56 percent increase in grass yields
on herbicide treated plots over yields on plots receiving no herbicide for
the three-year period 1970-1972 (Table 8). Even with the decrease in total
grass yields in 1976 as compared to grass yields in previous years, it
appeared that in relative terms, grass yields continued to increase on
herbicide treated plots as compared to plots receiving no herbicide.
Apparently, whatever the reasons for low vegetative production in 1976 were,
the effect was less on grass species than on other plant species.

Interactions

Between

Nitrogen

Fertilizer

and 2,4-D Herbicide

There were no significant (P&lt; 0.05) interactions between 2,4-D and nitrogen
fertilizer for any of the weight variables in 1976. This was also true for
the three-year evaluation period (Carpenter 1976).
These results suggest
that all groups of plants (forbs, grasses, shrubs) responded to the chemical
treatments at the three study areas in a similar manner.

Nitrogen

Fertilizer

Efficiency

Ratios

Adjusted values for shrub herbage and total herbage for each nitrogen treatment without herbicide over the four evaluation years were used to calculate
efficiency ratios for nitrogen fertilizer.
The adjusted increases in yields
obtained at each nitrogen level over the control treatments (no nitrogen) for
each year were added together and this total divided by the level of nitrogen
applied to obtain this ratio.
It is apparent that for both shrub herbage
(Table 9) and total herbage (Table 10) the highest level of nitrogen (134.4
kb N/ha) was the most efficient in terms of kilogram of herbage produced per
kilogram of nitrogen applied.
For total herbage yield, the second most efficient
level was the second rate of nitrogen (67.2 kg N/ha) followed by 33.6 and 100.8
kg N/ha levels, respectively.
For shrub herbage, 67.2 kg N/ha was second most

�-57-

efficient, followed by 100.8 and 33.6 kg N/ha, respectively.
As shown in
Tables 9 and 10, benefits from the lower nitrogen rates became negligible
by the third year, while increases in yields at the highest level were
obtained even in the sixth year.

Table 9. Adjusted increases in shrub herbage yields (kg/ha) over control
plots for years 1970, 1971, 1972, and 1976 and the corresponding efficiency
ratio for each level of nitrogen.
Rates of Nitrogen
67.2

Fertilizer (kg/ha)
100.8

134.4

288

190

569

116

289

306

463

1972

-51

-16

109

328

1976

-71

-14

-13

93

Totals

94

547

592

1,453

2.80

8.14

5.87

10.81

Year

33.6

1970

100

1971

Efficiency

Ratio

Table 10. Adjusted increases in total herbage yierds (kg/ha) over control
plots for years 1970, 1971, 1972, and 1976 and the corresponding efficiency
ratio for each level of nitrogen.
Fertilizer (kg/ha)
100.8

134.4

361

262

647

175

261

339

546

1972

7

-8

152

388

1976

-75

-25

-40

65

Totals

255

589

713

1,646

7.59

8.76

7.07

12.25

Year

33.6

1970

148

1971

Efficiency

Ratio

Rates of Nitrogen
67.2

�-58-

It appears from these results that most of the beneficial effects of one
application of nitrogen fertilizer on yields of plant species common to
sagebrush rangeland is obtained in the first 3 years after treatment.
Six years after treatment, very small responses in vegetation yield were
obtained.
As evidenced by yields on control treatments, 1976 was a poor
year for plant growth.
Reasons for this are unclear.
As a result of this
"year effect", one is limited as to the conclusions that can be drawn regarding lasting effects of nitrogen fertilizer.
It does appear however,
that six years after treatment only the high rate of fertilizer application
results in increased yields.
The decrease in yields on the 33.6, 67.2, and
100.8 kg N/ha treatments relative to the control treatments is puzzling and
somewhat disturbing.
If these decreases are real they may indicate detrimental effects to plant growth as a result of nitrogen fertilizer.
Perhaps
larger root systems (as a result of fertilizer treatments) utilize moisture
less efficiently than root systems not stimulated by fertilizer and as a result yields are decreased.
Increased competition (between plant species)
as a result of nitrogen fertilizer may also be involved.
It is evident
that the main plant species responding to nitrogen is big sagebrush.
Perhaps stimulation of growth of this species with nitrogen results in an inbalance of available nutrients to other plant species.
This should be
evaluated further, as long-term effects of nitrogen fertilizer on native
vegetation are as important as immediate effects.
Herbicide treatments appeared to be longer lasting than fertilizer treatments, although larger vegetation yields were obtained with combination
treatments of nitrogen and 2,4-D than with either treatment applied alone.
Much of this increased yield was due to responses of grass species at the
expense of forb and shrub species.
Dominance of grass species would not
be as great if treated areas were subject to grazing by domestic livestock.
With the single exception of decreasing forb yields on herbicide treatments,
results of this study suggest that fertilizer-herbicide
combination treatments are superior to either treatment applied separately, in improving
deer winter ranges.
This conclusion is based primarily on increased vegetation yields and altered composition of forage classes making up these
yields, and on measured food habits of mule deer on these winter ranges
(Carpenter 1976).

LITERATURE

CITED

Baldwin, D. M., N. W. Hawkinson, and E. W. Anderson.
1974. High-rate
fertilization of native rangeland in Oregon.
J. Range Manage.
27:214-216.
Bowns, J. E. 1972. Low level nitrogen and phosphorus fertilization
high elevation ranges.
J. Range Manage. 25:273-276.

on

Carpenter, L. H. 1970. Middle Park deer study - range fertilization.
Colo.
Div. Game, Fish and Parks.
Game Res. Div. Fed. Aid Proj. W-38-R-24.
Game Res. Rep., July, Part 3. p. 371-391.
1971. Middle Park deer study - range fertilization.
Colo. Div.
Game, Fish and Parks.
Game Res. Div. Fed. Aid Proj. W-38-R-25.
Game
Res. Rep., July, Part 3. p. 225-253.

�-59-

Carpenter, L. H. 1972. Middle Park deer study - range fertilization.
Colo. Div. Wildlife.
Game Res. Div. Fed. Aid Proj. W-38-R-26.
Game
Res. Rep., July, Part 2. p. 213-251.
______ , O. C. Wallmo, and M. J. Morris.
1973. Effect of woody stems on
estimating herbage weights with a capacitance meter.
J. Range Manage.
26:151-152.
_______
1976. Nitrogen-herbicide effects on sagebrush deer range.
Dissertation.
Colo. State Univ .• Fort Collins.
l59p.

Ph.D.

Currie, P.O., D. L. Neal, and M. J. Morris.
1969. Development of the
heterodyne meter and partial test results on some range and wildlife
vegetation types in the western U.S.A.
In-service report.
USDA For.
Ser., Rocky Mtn. For. and Range Exp. Sta. 7p. (unpublished).
Hubbard, W. A., and J. L. Mason.
1967. Residual effects of ammonium
nitrate and ammonium phosphate on native ranges of British Columbia.
J. Range Manage. 20:1-5.
Johnston, A., A. D. Smith, L. E. Lutwick, and S. Smoliak.
1968. Fertilizer response of native and seeded ranges.
Can. J. Plant Sci. 48:467-472.
Kilcher, M. R., S. Smoliak, W. A. Hubbard, A. Johnston, A. T. H. Gross, and
E. V. McCurdy.
1965. Effects of inorganic nitrogen and phosphorus
fertilizers on selected sites of native grassland in Western Canada.
Can. J. Plant Sci. 45:228-237.
Mason, J. L., and J. E. Miltimore.
1959.
content of native bluebunch wheatgrass
Can. J. Plant Sci. 39:501-504.

Increase in yield and protein
from nitrogen fertilization.

______ , and
1972. Ten year yield response of beardless wheatgrass
from a single nitrogen application.
J. Range Manage. 25:269-272.
Neal, D. L., and J. L. Neal. 1973. Uses and capabilities of electronic
capacitance instruments for estimating standing herbage.
J. Br. Grassl.
Soc. 28:81-89.

P repa red by __ -,-:k~.L,-"=-\\.-,--"::_._C."-&lt;:.d\u..c),'"-'~-N"-'g •.••
-"",,,,,J;
...•...•.
~,,,.....),---Len H. Carpenter~
Wildlife Researcher C

�-65-

July, 1977

Jon PRC(~~ESS kEPO!n'

COLOPJ'.DO

State of
Project

Deer-Elk Investigations

W-38-R-32

No.

Hork Plan No.

14

Job No.

9

------------------------------------Period

Cove.red:

Personnel:

April 1, 1976 through March 31, 1977

Len H. Carpenter, M. A. Kautz, D. J. Freddy, L. L. Strong, D. G.
Milchunas, and R. B. Gill

ABSTRACT
Diets of two deer were evaluated by both in vivo and in vitro digestion
techniques.
Resulting values were very similar for one diet but differed
by more than 20 percent for the other.
Chemical analyses of the two diets
were similar.
In a separate study, daily food intake of two pregnant mule
deer was influenced by both level and type of food. Energy intake for the
two deer was estimated to be less than that previously measured for growing
fawns.

��-67-

MIDDLE PARK COOPERATIVE
DEER STUDY - DEER HABITAT EVALUATION
Len H. Carpenter

P.N.O. OBJECTIVE
Develop procedures
to support deer.

for quantifying

the capacity

of Middle

Park winter

ranges

SEGMENT OBJECTIVES
1.

Develop and test a candidate
of free-ranging mule deer.

method

2.

Estimate energy requirements
deer at pasture.

for activity

3.

Estimate nutritional quality of ingested
digestible and metabolizable energy.

4.

Estimate

protein

requirements

METHODS

for estimating

energy expenditures

and develop

energy budgets

for

forage in terms of in vivo

of fawns for maintenance

and growth.

AND MATERIALS

Methods and materials employed in this study have been previously described
(Gill and Baker 1974, Carpenter and Baker 1975, Carpenter 1976). Poor success
in raising deer fawns in 1976 prevented detailed nutritional experimentation
and completion of Segment Objective No.4
during the winter of 1976-77.
Summary
of previous data and review of literature occupied much of the work effort this
segment.

Method for Estimating Energy
Expenditures of Free-Ranging Mule Deer
This portion of the study is being conducted by M. A. Kautz, Colorado State
University.
A Progress Report by M. A. Kautz, covering a portion of this work
is presented in Appendix B. A thesis covering this work is in progress and will
be presented in the next segment report of Work Plan 14, Job 9.

Estimating
and Developing

Energy Requirements for Activity
Energy Budgets for Deer at Pasture

Data on energy requirements by various activities obtained by M. A. Kautz and
data on activity patterns of deer at pasture (Carpenter 1976) is being incorporated into an overall energy budget for mule deer.

�-68-

A simulation model developed by D. M. Swift, Natural Resources Ecology
Laboratory at Colorado State University is being used as a first approximation of carrying capacity of Middle Park winter ranges.
To measure energy costs of pregnancy, food intake measurements were taken
for two tame mule deer during the period January 11 to March 24, 1977
(3-5 months gestation).
Ad libitum intakes were monitored daily during
this period.
Three food types and three different levels of food were
offered and varied between trials to determine influence on intake.
The
three food types offered were: 1) pawnee special (alfalfa pellets), 2)
dairy ration (cereal grains, no alfalfa pellets), 3) mixed ration (cereal
grains and alfalfa pellets).
Levels offered were: 1) 1,000 grams, 2) 1,500
grams, and 3) 2,000 grams.
The different foods were tested because all
are or have been used in maintenance of the experimental deer herd and
differences have been noted with deer in acceptance of the various food
types in pen situations.

Estimating

Nutritional

Quality

of Digested

Forage

A thesis on file at Colorado State University entitled "In vivo-In vitro
relationships of Colorado mule deer forages" (Daniel G. Milchunas 1977.
M.S. Thesis, 133 p.) covers portions of this objective.
The abstract of
this thesis is presented in Appendix A.
Data obtained in the in vivo digestion trial described last segment
(Carpenter 1976) wereanalyzed
this segment.
In vitro digestion trials
were conducted on the overall diet of each deer and on the top ten plant
species in each diet using the method of Tilley and Terry (1963) and
modified by Pearson (1970).
In vitro values were compared to in vivo
values for the two diets.
Determinations were also made of drY-matter,
ash, cell wall constituents, acid detergent fiber, lignin, cellulose,
hemicellulose,
and gross energy for each diet. Triplicate samples were
used for determinations
of dry matter, cell wall constituents and ash,
while duplicate samples were read for all other variables.
Dry matter and
gross energy were also determined for feces and urine samples collected from
each deer. Dry matter was determined at 1000C for 24 hours.

RESULTS AND DISCUSSION
Method
Expenditure

for Estimating Energy
of Free-Ranging Mule Deer

A thesis covering this work is in progress and will be presented in the next
segment report.
A progress report covering this work is presented in Appendix

B.
Estimating Energy Requirements for Activity
and Developing Energy Budgets for Deer at Pasture
Results of the simulation modeling
will be reported on next segment.
incorporated into this model.

approach to estimate carrying capacity
Data from previous segment work will be

�-69-

Daily ad libitum intakes for the two pregnant does are presented in Table
1. Type of food and level offered appeared to affect intake.
Within a
food type, an increase in amount offered increased intake.
However, when
both type of feed and level were varied, the two deer did not always respond in the same manner (Table 1). More deer and additional levels of
food offerings are needed to adequately test these relationships.
These
data do however, point out the importance of standardizing food type and
level of food offered before drawing conclusions on feeding experiments.
Energy intakes of the pregnant does were compared to energy intakes of
growing fawns (Baker 1976).
The same ration (pawnee special) was offered
to both groups in the same amounts and under similar conditions.
Baker
(1976) reported an average daily intake of 987 grams for 7 fawns offered
2,000 grams per day.
Gross energy of this food was determined to be 4.60
kcal/gm with 71.6 percent of the gross energy being digested (Baker 1976).
It was further estimated by Baker that 87.0 percent of digestible energy
was metabolizable.
Consequently, the mean metabolizable energy intake for
the 7 fawns was determined to be 2,828 kcal/day.
Using average metabolic
weight (kg W 0.75) of the fawns (15.94) it was determined that average
energy intake per metabolic body size was 177.4 kcal/kg W 0.75/day.
Assuming digestibility and metabolizability
of the food fed adult does was
the same as that measured with growing fawns, the following comparisons on
daily energy intake are possible.
Average daily intake for the two does was
929.1 grams (Table 1). Using the same digestibility and metabolizability
values discussed above it follows that the mean metabolizable energy intake
for the does was 2,661 kcal/day.
Using average metabolic weight (kg W 0.75)
of 21.8 (61 kg) for the two does, average energy intake per metabolic body
size was 122.1 kcal/kg W 0.75/day, about 55 kca1 less than the intake determined for growing fawns. However, the assumption that both age classes
of deer utilize food in a similar manner may be tenuous, and as a result
this comparison may not be valid. Hopefully, complete digestion and energy
balance trials on pregnant does can be conducted in the future.

Estimating

Nutritional

Quality

of Ingested

Forage

In vitro digestion values for the top 10 species and overall diet for deer
numbers 9 and 18 as measured in the 5-day grazing trial in March 1976 are
presented in Tables 2 and 3. Digestible dry matter for each individual plant
species was weighted according to its percentage in the diet.
The sum of all
weighted values represents the weighted digestible dry matter of the diet.
The digestibility of the entire diet as a whole is also presented in Tables
2 and 3. Weighted values in both cases are less than the value determined
for the diet as a whole.
In vitro values for the two deer diets are very similar even though composition of the two diets are dissimilar.
The ten top species in the diet of
deer number 9 were 55.7 percent grasses, 27.2 percent forbs, and 17.1 percent
shrubs.
The ten top species in the diet of deer number 18 were 24.0 percent
grasses, 36.2 percent forbs, and 39.8 percent shrubs.
A complete forage
analysis of each diet is presented in Table 4. Chemically, the two diets are
similar.

�Table 1. Daily intake (grams) of two pregnant mule deer as a function of varying types and levels
of offered (ad libitum) foods during period January 11, to March 24, 1977.

Food Type

Daily Amount
Offered (grams)

Days
Offered

Deer No. 7

Daily Intake
(grams
se)
Deer No. 50

Both
Deer Combined

±

-Mixed Ration*

1,000

25

527.8 + 52.4

604.2 + 23.4

566.0 + 29.2

Mixed Ration

1,500

7

935.4 + 27.1

798.7 + 17.3

867.1 + 24.3

Dairy Ration**

2,000

7

1,037.0 ±. 36.8

673.3 + 85.3

855.1 + 67.3

Pawnee Specia1***

1,000

10

934.7 + 25.3

855.7 + 22.1

895.2 + 19.0

Pawnee Special

2,000

10

± 47.5

831.0 + 44.4

929.1 + 39.2

1,027.1

I
-..J

0
I

*Mixed ration = cereal grains and alfalfa pellets.
**Dairy ration = cereal grains only.

*** Pawnee special = alfalfa pellets only.

�-71-

Table 2. In vitro digestible dry matter of 10 top species and overall
diet for deer 9 during period March 1-5, 1976. Grazing trials conducted
in 4 ha energy budget pasture, Junction Butte Research Center, Colorado.

Plant Species

Percent
of Diet

Percent
In Vitro DDM

Weighted*
Percent DDM

Poa fendleriana

40.3

38.75

15.61

Phlox bryoides

14.2

25.40

3.60

Astragalus

convallarius

11.8

39.52

4.66

Amelanchier

alnifolia

6.7

31.94

2.13

Sitanion hystrix

5.8

46.24

2.68

Artemisia

5.5

55.10

3.03

tridentata

Chrysothamnus

vaseyi

2.6

43.62

1.13

Symphoricarpos

oreophilus

2.3

27.68

0.63

Eriogonum

umbellatum

1.2

20.38

0.24

Agropyron

sEicatum

9.6

46.26

4.44

38.15

*Weighted
**

by percentage

Diet percent

DDM

of diet i.e. 38.75 X .403

= analysis on entire diet.

15.61

Diet**
Percent DDM

44.39

�-72-

Table 3. In vitro digestible dry matter of 10 top species and overall
diet for deer number 18 during period March 1-5, 1976. Grazing trials
conducted in 4 ha energy budget pasture, Junction Butte Research Center,
Colorado.

Plant Species

Percent
of Diet

Percent
In Vitro DDM

Weighted*
Percent DDM

Phlox bryoides

32.2

25.40

8.17

Artemisia tridentata

31.5

55.10

17.35

Poa fendleriana

22.8

38.75

8.83

Amelanchier alnifolia

5.1

31.94

1.62

Astragalus convallarius

2.4

35.92

0.94

Chrysothamnus vaseyi

2.5

43.62

1.09

Agropyron spicatum

1.2

46.26

0.55

Eriogonum umbellatum

0.9

20.38

0.18

Penstemon caespitosus

0.7

30.64

0.21

Artemisia frigida

0.7

48.45

0.33

39.27

*Weighted by percentage of diet i.e. 25.40 X .322
**
Diet percent DDM = analysis on entire diet.

8.17.

Diet**
Percent DDM

42.35

�-73-

Table 4. Forage analyses of diets for deer numbers 9 and 18 during period
March 1-5, 1976.
Grazing trials conducted in 4 ha energy budget pasture,
Junction Butte Research Center, Colorado.

Deer
No.

%
Moisture

9
18

%

%
Hemicellulose

Gross
Energy
kcal/g

Cell Wall

%
Ash

%
ADF

%
Lignin

%
Cellulose

5.55

55.83

13.13

51.85

8.46

43.39

3.98

3.73

5.11

42.97

19.01

49.52

10.87

38.65

0

3.59

The two diets were also evaluated in vivo. Table 5 presents daily intake and
feces and urine output for the two deer during the 5-day trial.
In vivo
digestibility
for the diet of deer number 9 was 43.6 percent, very comparable
to the in vitro value of this diet. However, the in vivo digestibility of the
diet of deer number 18 was 68.1 percent, or about 20 units higher than the in
vitro value.
Reasons for this wide discrepancy are not fully understood.
Deer number 18 consistently had a higher daily intake and a lower fecal output
than did deer number 9 (Table 5). Intuitively, the diet of deer number 9 should
have had the higher digestion coefficient because of the greater percentage of
grasses which were higher in digestibility (Tables 2 and 3). The problem of
soil contamination in plant samples is greater with low-growing plants such as
phlox (Phlox bryoides) than with other plant species.
Perhaps the large amount
of phlox and the corresponding large amount of ash in the diet of deer number
18 (Table 4) resulted in a portion of this error.
It is recommended in future
analyses that soil contamination be accounted for. In addition, errors in bite
counting and quantifying bite sizes were perhaps larger with deer number 18.
The higher percentage of phlox in the diet of deer number 18 may have been
responsible for a portion of this error.
Counts of bites from this forb is
difficult.
However, it does not seem reasonable that these factors alone could
account for such a large difference.
Hopefully, deer fitted with esophageal
fistulas will be available soon which will facilitate the quantification of
intake of free-grazing mule deer.
It is suggested that for future in vivo
trials, data be collected for 7 days in place of 5 days. This should give a
better measure of intake and output.
Comparisons of urine output (grams per day) (Table 5) for the two deer are also
interesting.
Urine output for deer number 18 was greater than for deer number 9.
Gross energy of the urine of deer number 18 was 3.04 kcal/g while for deer
number 9 it was 2.63 kcal/g.
For the 5-day trial this projected to a total
urinary energy loss of 568 kcal for deer number 18 and 279 kcal for deer number
9. Big sagebrush (Artemisia tridentata) made up 31.5 percent of the diet of
deer number 18 but made up only 5.5 percent of the diet of deer number 9. It
is known (Cook 1970) that many of the volatile oils (high in gross energy) of
sagebrush are not metabolized by ruminants but are excreted in the urine.
These
data support that finding.
Gross energy of the feces of deer number 9 was
3.58 kcal/g while for deer number 18 it was 3.28 kcal/g, '3:gainindicating
similarities in digestibility of the two diets.

�-74-

Table 5. Daily intake (grams) and feces and urine output (grams) for
deer numbers 9 and 18 durin§ in vivo digestion trial March 1-5, 1976.
All weights oven-dry at 100 C for 24 hours.

Day

Intake
Deer 9
Deer 18

Deer 9

Feces
Deer 18

Deer 9

Deer 18

1

152

242

142

81

33

55

2

88

227

107

69

12

26

3

275

389

142

77

23

42

4

143

234

96

128

19

37

5

339

452

75

138

19

27

Totals

997

1,544

562

493

106

187

Urine

LITERATURE CITED
Baker, D. L.
Thesis.

1976. Energy requirements of mule deer fawns in winter.
Colorado State Univ., Fort Collins. 76p.

M.S.

Carpenter, L. H. 1976. Middle Park Deer Study - deer habitat evaluation.
Colo. Div. Wildl., Game Res. Div., Fed. Aid Proj. W-38-R-3l. Game
Res. Rep., July, Part 2. p. 283-298.

--- , and D. L. Baker.

1975. Middle Park Deer Study - deer habitat evaluation. Colo. Div. Wildl., Game Res. Div., Fed. Aid Proj. W-38-R-29. Game
Res. Rep., July, Part 2. p. 241-263.

Cook, C. W.
TBl09.

1970. Energy budget of the range and range livestock.
Colo. State Univ. Expt. Station, Fort Collins. 28p.

Bull.

Gill, R. B., and D. L. Baker. 1954. Middle Park Deer Study - deer habitat
evaluation. Colo. Div. Wildl., Game Res. Div., Fed. Aid Proj. W-38-R-28.
Game Res. Rep., July, Part 2. p. 211-225.
Pearson, H. A. 1970. Digestibility trials: In vitro techniques. p. 85-93.
In Range and Wildlife Habitat Evaluation. A research symposium. U.S.D.A.
Forest Service Misc. Publ. 1147. 220p.
Tilley, J. M., and R. A. Terry.
digestion of forage crops.

1963. A two-state technique for in vitro
J. Brit. Grassl. Soc. 18(2):104-111.

Prepa red by __ ~-=--L'='-' -,~,-",--_,---,&gt;C,""".d::U~'-'6.:J-~~=~c..::.=::=.==-_
Len H. Carpenter~
Wildlife Researcher C

�-75-

A P PEN

D I X

A

(Abstract of Daniel Milchunas

Thesis)

�-76-

ABSTRACT

IN VIVO - IN VITRO RELATIONSHIPS

Morphological
digestibility
fir summer

and phenological

were observed

OF COLORADO

differences

winter

Twelve summer and thirteen winter

forages examined

constituents,

crude protein,

and 55.3 and 38.1% in vitro DDM.

between

content.

Significant

native

multiple

possible

negative

and multi-species

that nitrogen

experiments

and

stimulates

micro-

effects are not observed

carbon structure

exploitation

effects

unless the

potential.

opposing

positive

caution must be used when relating

combinations

From regressing

or diets.

the equation

Y = 1.377

+ 0.935x (r 2 = 0.95) is the

of in vivo DDM from in vitro DDM.

best predicted

from in vitro DDM "by weighted

mean" rather

vitro digestion

of the mixes.

energy as a percent

Metabolizable

Mixed

forage diets are
than from in
of digestible

from 65.5% to 77.0%.

Total nutritive
digestibility.

in vitro

in vitro to in vivo DDM data of this and other studies

combinations,

ranged

With

in vitro to in vivo

best predictor

energy

were

effects were observed between

forages,

suggests

in vivo associative

effects,

DDM for species

Large differences

nitrogen supplementation

but that associative

also increases

in various

two native

analysis

mixture

associative

27.2 and

6.3 and 10.9% lignin, 11.8 and 5.6

in vitro associative

Non protein

regression

organisms,

averaged

in vitro DDM and in vitro DOM in forages of high ash

forage and alfalfa,

combinations.

from mule deer spruce-

range in Middle Park, Colorado.

46.5% cell wall

observed

in forage in vitro

in species examined

range and sagebrush

MULE DEER FORAGE

input to an animal

is a function

Intake under bulk limiting

conditions

of both intake and
is a function

of gut

�-7:]-

capacity of forage turnover
consistently

time.

related to chemical

due to the importance

Further,

rate of passage

constituents

of particle

of a forage or digestibility

size reduction

in passage

appears

that in certain cases lignin can provide brittleness

physical

characteristics

rumination-mastication
by simulating

whereby
occurs.

meals whereby

is not

rates.

It

to forage

particle

size reduction

is enhanced

Relative

forage intakes were calculated

an animal eats to a constant

when

"fill" and these

meals "decay" at a rate based on forage passage

functions.

Relative

at steady state and DE or ME are then combined

to produce

a digestible

energy intake index (DElI) and metabolizable
Across

forage comparisons

reordering

in DElI or MEII may provide
observations

energy intake index (MEII).

of mean DElI or MEII to DE or ME indicates

of forage value.

It is suggested
explanations

where deer are utilizing

intake

that turnover

to seemingly

large quantities

a

rate as applied

paradoxical
of a forage of high

fiber, low diges tibility.

Daniel George Milchunas
Natural Resource Ecology Laboratory
Colorado State University
Fort Collins, Colorado 80523
Winter, 1977

�-78-

A P PEN

D I X

B

PROGRESS REPORT

Project:

The Use of Heart Rate to Estimate the Energy Expenditure
of Mule Deer Fawns.

Colorado Cooperative Wildlife Research Unit
Funded by Colorado Division of Wildlife Contract No. C102064
Marie A. Kautz, Graduate Research Assistant
Period Covered: April 1, 1976 -March 31, 1977

�-79-

THE USE OF HEART RATE
TO ESTIMATE

THE ENERGY EXPENDITURE

OF MULE DEER FAWNS

OBJECTIVES
1.

Evaluate the usefulness of a face mask apparatus
collections from mule deer fawns.

for making

respiratory

2.

Investigate the relation between the heart
of fawns engaged in several activities.

rate and energy

expenditure

SEGMENT OBJECTIVES
1.

Complete
air.

development

of a system

for measuring

2.

Rear and train four fawns to experimental

procedures.

3.

Perform

measurements

and heart rate.

4.

Analyze

data.

of energy expenditure

METHODS

and collecting

expired

AND MATERIALS

The methods and materials used to meet 'Objectives 1, 3, and 4 are reported
by Kautz, in Carpenter (1976).
Fawns were obtained from captive does or acquired as orphans.
reared using procedures similar to those described by Reichert

They were
(1972).

A training program was developed as the fawns grew. At 2-3 weeks of age
open-ended styrofoam cups with elastic straps were placed on their muzzles
during bottle feeding.
Gradually the cups were made more restrictive and
heavier by leaving the distal end partially closed in and taping on weights.
At about 1 month the fawns were introduced to plastic cups used in the same
manner but held on by webbing in the same way as the real mask. Weights were
added gradually to the plastic cups until the fawns accepted about fourteen
ounces and weighed about 50 pounds.
At this time they were judged ready for
the real mask without valves.
After this became acceptable, the face-cuff of
the mask was tightened, the valves put in, and the hoses attached.
The fawns
were given raisins as a distraction and reinforcement, both through the partially open-ended cups and after removing the real mask. Harness and leash
training was done in conjunction with the mask training as described by
Reichert (1972). Each fawn was trained in this manner daily for 10-30 minutes
until 3-4 months of age, when the intensity was decreased to 3 or 4 times a
week.
Four fawns were brought through the training program.

�-80-

RESULTS AND DISCUSSION
Success of Training
The training procedures were highly effective. Of the four deer trained
intensively three were usable as experimental subjects. The fourth (deer
54) accepted the mask throughout the study period, but would not accept
handling from a technician who joined the study when the deer were over
five months of age. Since hooking up the apparatus required the technician's
assistance, deer 54 was dropped from the study. The deer's position as
dominant female of the group may have been related to her intolerance of
being handled by a new person. No such problems arose with the other three
deer (two males and one female), all of which accepted handling by the new
technician within two weeks.
Of the three deer used in the study the two males could be outfitted readily.
The female (deer 55) accepted the face mask only by being coaxed or tricked
into it. Often several attempts were necessary but it was always possible
to get the mask on her. The history of deer 55 was different from those of
the others in that she was about a month younger. She was raised for her
first three weeks at the home of a researcher, and she was trained less than
the other three. In general she was more active, independent, and aggressive
than the others. Thus the poorer acceptance of handling procedures by the
females is not clearly a sex difference.
At certain times all the deer were excited and showed a high level of activity.
Trials were not conducted at these times because the deer were difficult to
handle and restrain. If the deer were bedded, standing around, or quietly
feeding, outfitting could be readily accomplished and trials completed with
minimal restraint.

Measurement of Energy Expenditure
One hundred-ten simultaneous measurements of heart rate and energy expenditure
were completed when the deer were between six and nine months of age. Of
these, eleven trials were done on deer fasted for 48 or more hours. Thirteen
trials included some bedding; other activities included standing and walking
and some trotting (less than 10 percent of any trial). The ranges of oxygen
consumption, heart rate, and percent of time spent bedding and standing combined for each deer are presented in Table 1. For all deer oxygen consumption
ranged from 14.2 to 63.4 ml/kg 0.75 min and heart rate from 49.3 to 96.0 beats
per minute. Percent of time spent bedding and standing combined ranged from
20 to 100.

�-81-

Table 1.
trials.

Ranges

of measured

parameters

for individual

deer during all

51 d"

Deer Number
53 c3"

Oxygen consumption
ml/kg 0.75 min.

15.6-46.6

18.6-63.4

14.2-48.8

Heart

50.5-93.1

49.3-92.7

53.0-96

28-98

20-99

21-100

Parameter

rate beats/min

Percent time spent
bedding and standing
combined

55 ~

Data analyses are incomplete and some heart rate counts need to be rechecked.
Therefore the following results which include heart rates are tentative.
Measurements of oxygen consumption for fasting trials are presented in Table
2. In general fasted animals were more likely to bed and had lower oxygen
consumptions than fed animals.

Table 2. Oxygen consumption for fasting trials by deer and activity,
order of increasing activity.

in

(h Cons.
ml/kgO.75

Deer No.

Trial No.

Percent Bedding

51

30

64

34

15.62

26

54

35

18.42

28

0

73

27.85

25

0

64

26.23

27

0

58

29.95

29

0

54

39.80

111

81

16

14.17

112

63

36

16.61

110

10

82

19.39

109

0

89

35.94

108

0

36

45.36

55

Percent

Standing

min

�-82-

In trials in which bedding occurred, percent of time spent bedding appeared
to be related to oxygen consumption.
In trials without bedding, percent
standing was clearly correlated with oxygen consumption (r =-.672).
Other variables tested in a stepwise mUltiple regression with oxygen consumption as dependent variable were Julian day, temperature, and pulmonary
volume, and heart rate. The correlation for pulmonary volume and oxygen
consumption (r = .549) was somewhat lower than expected, probably due to the
effects of the face mask and the common occurrence of panting during warm
or active trials.
Heart rate was most strongly correlated with oxygen consumption.
Correlation
coefficients were .819, .913, and .914 for data from deer 51, 53, and 55,
respectively and .692 for all deer. A plot of oxygen consumption vs heart
rate for all deer is presented in Figure 1. The data from deer 55 are distinct from those of deer 51 and 53. This may be related to sex and/or age
and size. Deer 55 was a female, approximately one month younger and 13 kg
lighter than the mean weight of deer 51 and 53.

RECOMMENDATIONS
All segment objectives were met except No.4.
Further checking of some
heart rates, involving spot checks of counts from electrocardiographs
and
checks of the calculation of mean heart rates, is needed before analysis is
continued.
Analyses should also include consideration of the effects of
time of day; a more complete analysis of activity data; possibly consideration of data on human activity and subject nervousness; and further examination of relationships among the different variables measured.
Estimated
completion date for further analyses and a final report is September 30.

LITERATURE

CITED

Kautz, M. A. 1976. The use of heart rate to estimate the energy expenditure
of unrestrained deer. Appendix A, pp. 299-319 in Carpenter, L. H.
Middle Park deer study - deer habitat evaluation.
Colo. Div. Wildl. Fed.
Aid Proj. W-38-R-3l, WP 14, J9, Job Prog. Rep., Game Res. Rep. July.
Reichert, D. W. 1972. Rearing and training deer for food habits studies.
USDA For. Servo Res. Note RM 208, 7 p. Rocky Mt. For. and Range Exp.
Sta., Fort Collins, Colo.

�-83-

?O.II...----,.---r--.----,----.---.,---r--.---r---'----;.---I

1 - Deer 51
60.0

3 - Deer 53
5 - Deer 55

:z

!S1.e

r
I
to
r-

5 5

3

&lt;:)
(,.;:)

~
&lt;,

-'
~

I
3
40.0

3 3

~

0
l-

1 ~3

'!'

'~l"

3

JO.O

=c
:::l

.

33

:z

1

5 %
55

3

5

~
S
5

0

u

5

5

5
5
5 5
5
5

5
5

5

3

3t

5

5

5

33

333

13~

Vl

5

33

I
3
1 .; I 1
13

I

::z
Q..

,

3

5

5

5

:z

w
l..:&gt;
&gt;-

3

10.0

5

3:

X

5 5

0

5 5
5

5

5
10.00

~

.•

0

.,;

~

co.

Co

;:

~

co
co

••

"!

""~

HEART RATE CBEATS/H I NJ

Fig. 1. The relationship
deer fawns.

between oxygen consumption

and heart rate for three

��-85-

July, -1977

JOB FINAL REl'ORT

Colorado

State of

------------------------

Project

No.

W-38-R-32

Hork Plan No. -

14

Job Title Middle Park Cooperative
Period Covered:
Personnel:

Deer-Elk

Investigations

Job No.

10

Deer Study - Junction Butte Habitat

Study

April 1, 1976 through March 31, 1977

Wayne L. Regelin, Donald W. Reichert,
L. Firlick, and Len H. Carpenter

Glenn L. Crouch, Patrick

ABSTRACT
Effects of snow manipulation on deer use and vegetation growth were
summarized in a dissertation which serves as a final report for most
objectives of this job. The dissertation is on file at the Colorado
Division of Wildlife Research Center Library and the Colorado State
University Library and copies have been forwarded to appropriate Fish
and Wildlife Service Federal Aid personnel.
The second four blocks of treatment plots for the winter range seeding
experiment were installed and the first post-treatment plant cover and
composition measurements were made on plots established in 1975.

��-87-

MIDDLE PARK COOPERATIVE
DEER STUDY - JUNCTION BUTTE HABITAT

STUDY

Glenn L. Crouch

A dissertation entitled "Effects of snow drifts on mountain shrub
communities" (Regelin 1976) is on file at the Colorado Division of Wildlife Research Center Library and at the Colorado State University Library.
This dissertation presents information regarding effects of snow manipulation on vegetation growth and deer use and serves as a final report for the
snow manipulation portion of this study.

PeN.O. OBJECTIVE
In Middle Park, Colorado and other similar mule deer winter ranges in
mountainous sagebrush habitat, native forage does not produce sufficient
supplies of chemical protein and energy to maintain deer in the demanding
winter months.
The objective of this study is to develop revegetation procedures to establish stands of species with high contents of digestible
protein and chemical energy to supplement native vegetation in meeting the
nutritional requirements of mule deer in winter.

SEGMENT OBJECTIVES
1.

To compare the establishment and spread of planted species using three
methods of planging (drill rows, broadcast and harrowed, and broadcast
only).

2.

To compare herbage yield on three seedbed

treatments

and control areas.

METHODS AND MATERIALS
Methods employed in this study have been previously
(1974, 1975 and 1976).

detailed by Regelin

RESULTS AND DISCUSSION

Seeding. Deer Winter Ranges
The first post-treatment measurements of plant cover and composition were
made in July 1976 on plots established in 1975. Plants of all seeded species
were found, but amounts of each were minimal after only one growing season.
In addition, the second set of four treatment blocks were selected, and pretreatment plant cover, composition, and biomass were determined.
These sites
were prepared for seeding and seed sown in September as described by Regelin
(1975) •

�-88-

LITERATURE CITED
Regelin, W. L. 1974. Middle Park Cooperative Deer Study - Junction
Butte Habitat Improvement Study. Colo. Div. Wildl., Fed. Aid
W-38-R-28, Work Plan 14, Job 10, Job Prog. Rep., Game Res. Rep.
July, Part 2. p. 227-292.
1975. Middle Park Cooperative Deer Study - Junction Butte
Habitat Improvement Study. Colo. Div. Wildl., Fed. Aid W-38-R-29,
Work Plan 14, Job 10, Job Prog. Rep., Game Res. Rep. July,Part 2.
p. 265-294.
1976. Effects of snow drifts on mountain shrub communities.
Ph.D. Dissertation. Colorado State University, Fort Collins. 106 p.

�July, 1977

-89JOB PROGRESS

State of

COLORADO
------~~~~~---------

Project

No.

W-38-R-32

Deer-Elk

Investigations

14

Job No.

11

Work Plan No.
Job Title
Period

Snowmobile

Covered:

Personnel:

REPORT

April

Harassment

--------------------~-----------

of Mule Deer on Cold Winter

1, 1976 - March

Ranges

31, 1977

D. Freddy, L. Carpenter, T. Spraker, L. Strong, P. Neil, B.
McCloskey, J. Sweeting, Colorado Division of Wildlife; L. Ward
and J. Cupal, U.S. Forest Service.

ABSTRACT
An implanted heart-rate telemetry system was placed in 2 male, semi-tame
mule deer. Heart-rates of these deer were monitored during various daily
activities while deer were at pasture.
The telemetry system generally
performed well but artifact problems prevented obtaining accurate heartrates for walking, running, and some grazing activities.
While at pasture
these deer were harassed by 1 person, 2 persons, person plus a dog, and a
snowmobile moving on prescribed routes at varying distances from the pasture.
Deer reacted to harassment and heart-rate was found to be a sensitive measure
of deer reactions to harassment.

��-91-

SNOWMOBILE HARASSMENT OF
MULE DEER ON COLD WINTER RANGES
David J. Freddy

The expanding human population is continually encroaching upon wildlife
populations.
There appears to be two primary effects of this encroachment; direct loss of wildlife habitat to development and harassment of
wildlife populations.
Geist (1971) describes harassment as any activity of man that influences
animals, such that animals increase their energy demands (calories needed
to survive and reproduce) above normal.
Repeated often enough, harassment may cause animals to draw abnormally upon body energy reserves, or
consume increased amounts of limited forage. This increased "cost of
living" would be innnediately detrimental to the individual and in the
long run, detrimental to the population.
Pregnant females may abort or
resorb young or die because of resulting nutritional deficiencies or
physiological stresses.
Populations may respond to harassment by shifting
their distribution or eventually being eliminated.
Harassment may be more critical at certain times of the year. Winter is
considered the critical season for survival of mule deer (Odocoi1eus
hemionus) in the central Rocky Mountains.
Limited winter ranges and
limited amounts of quality forage place nutritional burdens upon deer.
Any activity that forces deer to abandon their normal routines of behavior,
movement and range utilization may increase the caloric energy needed for
deer to survive.
The snowmobile is a potential source of harassment to mule deer in Colorado.
Few deer winter ranges are exempt from snowmobile activity.
These ranges
are often associated with heavily traveled highways or adjacent to human
population centers which increases the susceptibility of deer to snowmobile
harassment.
Previous studies on the effects of snowmobiles on deer are limited in number.
Eckstein and Rongstad (1973) radio-collared white-tailed deer (Odocoi1eus
virginianus) and monitored deer responses to snowmobile traffic.
Home ranges
of deer did not significantly vary when snowmobiles were present nor did
snowmobiles appear to upset the daily movement patterns of deer. However,
deer did move away from snowmobile trails in response to the disturbance.
Dorrance et al. (1975) also studied white-tailed deer and utilized radio
telemetry to monitor deer responses to snowmobiles.
Home ranges of individual deer tended to increase in size, although not significantly, in
response to snowmobiles.
Deer appeared to shift their activities away
from snowmobile trails and increase their movements in response to snowmobiles.
The authors felt that deer become somewhat habituated to snoumobile activity but that in critical winters any disturbance by snowmobiles
eliciting a response by deer could seriously upset the energy budgets of
individual animals.
Although these studies using telemetered deer give insight into deer behavior resulting from contact with snowmobiles, no quantitative estimate

�-92-

of the energy cost of harassment is possible from this type of study.
An estimate of the energy expenditure of the reactions of deer to
snowmobiles is needed to quantitatively describe the effects of harassment.
Several approaches have been utilized to estimate energy expenditure in
wild and domestic ruminants.
Respiratory gas exchange methods have been
used to estimate energy expenditure in grazing animals (Young and Corbett
1968, Corbett et a1. 1969). However, this technique requires the animal
being studied carry awkward equipment under restrained conditions and
therefore difficult to use in a free ranging situation.
Also, the use of
radio isotopes, primarily a form of labeled CO , in injection or infusion
2
techniques appears to give reliable estimates of energy expenditure
(Young et a1. 1969, White and Leng 1968, Corbett et a1. 1971). However,
the infusion process may not be reliable during cold winter temperatures
common to deer winter ranges.
A technique to estimate energy expenditure
that appears favorable is heart-rate.
Recent advances in micro-biotelemetry
have provided easily obtainable external and internal heart-rate radio
transmitters.
These transmitter systems place minimal restraints on free
ranging animals.
Heart-rate has been monitored in man and wild animals and related to
metabolic rate and heat production, or energy expenditure (Wyndham et a1.
1959, Booyens and Hervey 1960, Webster 1967, Datta and Ramanathan 1969,
Bradfield et a1. 1969, Mbrhardt and Morhardt 1969, Owen 1969).
Jacobsen
(1973) summarizes published estimates of heart rate-energy expenditure
relationships for various animals and presents an equation, based on these
estimates, for predicting energy expenditure in deer using heart-rate.
The equation:

+ 0.09371

Y =

-1.0653

(Y)=

rate of energy transaction

and, (X)=

heart rate in beats/minute

where

could be used to estimate
mobile harassment.

(X)

the energy expenditure

in kcal/kg·75/hour

imposed

on deer by snow-

With the availability of telemetry equipment and the apparent relationship
between heart-rate and energy expenditure, the effect of harassment in
terms of energy cost might be estimated.
Responses of deer to snowmobile
activity can be monitored and concurring energy costs estimated.
This study is being conducted in cooperation with the u.S. Forest Service,
Rocky Mountain Forest and Range Experiment Station, Laramie, Wyoming under
Cooperative Agreement No. l6-67l-CA.

P.N.O.

OBJECTIVE

Evaluate whether snowmobile activity on winter ranges inhabited by deer
decreases the ability of deer to survive winter by modifying the activities
of deer so as to significantly increase energy expended.

�-93-

SEGMENT OBJECTIVES
1.

Monitor heart-rates of deer during various daily activities to
quantify the energy costs of these activities during winter.

2.

Ascertain activities
harassment.

3.

Evaluate observed reactions of deer to snowmobiles in terms of
appropriate heart-rate measurements and corresponding energy
costs.

4.

Evaluate the energy cost of snowmobile harassment to deer in relation to the total estimated energy budget of deer (concurring
study).

s.

Examine alternatives to minimize
snowmobiles on deer.

of deer responding

to prescribed

potential

detrimental

snowmobile

effects of

METHODS AND MATERIALS
Two semi-tame, male castrate mule deer, age 2-1/2 years, were equipped
with implanted heart-rate monitoring transmitters.
The telemetry system
was essentially the same as that reported by Cupal et al. (1974) (Figs.
1, 2).
Surgery for implanting transmitters proceeded as follows: 1) animal was
anesthetized with Rompum, intramuscularly, and Ketamine, intravenously,
2) hair was shaved at the base of neck and top and bottom of sternum,
3) hair and skin were aseptically cleansed, 4) subcutaneous incisions
were made at the base of neck and sternum areas, 5) blunt dissection was
performed within the neck incision to prepare a pouch for transmitter
implant, 6) a catheter (large gauge hypodermic needle) was rammed subcutaneously from the incision at the top of the sternum to the neck incision, 7) electrodes were fed into the catheter and the implant placed
in the pouch, 8) catheter removed (electrodes now surfaced at the top
sternum incision, and rammed subcutaneously from the bottom to the top
sternum incision, 9) electrodes fed into catheter, catheter removed,
electrodes at bottom sternum incision (Fig. 3), 10) antibiotic salve
placed into incisions, incisions sutured, and animal given large dose of
penicillin.
Surgery resulted in one electrode within the tissue at both
the top and bottom of the sternum.
Electrodes were not affixed to tissue
but simply pushed into sternum tissue. Aseptic procedure was followed.
These deer were placed in a 10 acre pasture of native sagebrush winter
range located at Junction Butte Research Center, Middle Park, Colorado.
Animals were visually and telemetrically monitored from a glass enclosed
tower adjacent to the pasture (Fig. 4). From 7:00 AM to 5:30 PM for two,
5 consecutive day trials in February, heart-rates and corresponding activities
were monitored.
Heart-rates and activities were sampled during 3, systematically spaced 10 minute intervals per hour. Deer sampled were alternated
every other 10 minute interval, with the first deer sampled each day chosen
at random.
Activities were classified as bedded, bedded plus rumination,

�-94-

standing, standing plus grazing, walking, walking plus grazing, running,
alert reaction bedded, and alert reaction while up. This approach will
allow examination of the variability in heart-rate due to deer, activity,
trial, and time-of-day.
Deer fed primarily on native forage but were
occasionally given grain supplement.
While at pasture, deer were harassed by a snowmobile, people, and a
person and dog moving on prescribed routes at varying distances from the
pasture.
Deer activities and heart-rates were monitored for 5 minutes
prior to, during, and for 5 minutes after each harassment activity.
Heart-rates were determined directly from strip-chart recordings
a Hewlett-Packard
MOdel 9820A digitizer-calculator
system.

using

RESULTS AND DISCUSSION
Heart-rates
determined.

for deer activities

and harassment

reactions

are still being

The telemetry system generally performed satisfactorily.
However, problems exist and the system needs refinement.
Initially, heart-pulse
signal was being lost when the animals were in a head-down feeding position.
The transmitting range of the implant was about 12-14 inches and, when the
transceiver neck-collar slid down the neck beyond the implant's transmitting
range, the signal was lost. A sustaining collar about 7 inches wide made
of vinyl-coated cotton webbing lined with poly-sponge was buckled around
the neck in front of the transceiver collar.
This kept the transceiver
collar close to the implant so signal was not lost during feeding (Fig. 5).
The extra collar did not appear to bother the deer.
To receive continuous
heart-rate data from wild deer, either a sustaining collar must be employed
or the implant's transmitting distance increased.
The latter option would
increase battery power needed and possibly the implant's size.
The second problem with the telemetry system was more severe and complex.
During bedded, "quiet" standing, and "quiet" grazing activities ("quiet"
meaning little body movement), heart-rate signal was received well.
However, during "active" feeding, such as head-up, head+down motion in quick
succession, or walking and running activities, heart-rate signal was
erratic (Fig. 6). This indicated artifact (spurious electrical noise) was
being interpreted as heart-pulses
(R-waves) by the telemetry system and
leading to erroneous heart-rate recordings.
Further evidence of this problem occurred during bedded activities.
Heart-rate signal was constant
until the animal moved its head or foreleg which seemed to cause an instantaneous rise in heart-rate (Fig. 7). Because of the artifact problem,
heart-rates for walking, running, and most grazing activities could not
be ob tained .
Causes of this artifact problem are unknown.
Possible causes are electrical
impulses created by: 1) muscle movement near the sternum, 2) slight movement of the implant and electrodes within the animal as the animal moves,
and 3) electrode movement resulting in minute changes in distances between
electrodes.
Further developmental research must be undertaken to solve this
problem or else accurate heart-rates for walking, running, and grazing
activities cannot be obtafned.

�-95-

Fig. 3. Surgery to implant transmitter package, Top: electrodes
initially fed into catheter, Center: electrodes ready to feed into
catheter again, Bottom: suturing implant incision. (Photos byF. Waugh).

�-96-

fig. 1. 'I'r
ansc ei.ver- neck collar, left, and heart-pulse
transmitter, right, used to monitor heart-rates of deer.
Two stainless steel electrodes exit from transmitter
within a protective teflon plastic tube and surface outside the tube (points A &amp; B) to contact animal tissue.
(Photo by F. Waugh).

Fig. 2. Receiver
rate signals.

and strip-chart

recorder

used to receive

heart-

�-97-

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�-98-

Fig. 5. Sustaining collar
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bottom picture.

used to keep transceiver neck-collar
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�-100-

To measure the magnitude of artifact, an external ECG telemetry system
(Kautz, unpublished data, Colo. State Univ. 1977) was affixed to one
deer and both the external and internal heart-rate systems were monitored
simultaneously.
However, the external system also produced artifact
during walking and grazing activities, preventing measurement of heartrates for these activities.
When the deer was standing quietly, the
external system produced an excellent ECG waveform trace (Fog. 8). Through
simultaneous monitoring the internal transmitter was found to be accurately
pulsing for every "R wave" produced during a quiet stand activity.
Harassment trials were conducted in February and March.
Forms of harassment tested were one person, two persons, and one person plus a German
Shepherd dog walking, and a person riding a snowmobile over 6 routes ranging
from 0 to 400 yards from the pasture.
Successive routes were traveled on
one day with a minimum of 10 minutes between routes.
"Harassers" were out
of sight of the deer while waiting to travel the next route.
Duration of
harassment ranged from 8-26 minutes.
Animals were not actively pursued
during harassment trials.
Little or no snow was present during the trials.
Deer reacted to harassment with some degree of overt behavior during all
trials except two (Table 1). Two persons walking 100 yards above the
pasture elicited the strongest response from the deer. Deer reacted by
running to the farthest corner of the pasture away from this harassment,
and remained alert for at least 5 minutes after '~arassers" were out of
sight.
A person walking with a dog appeared to have little effect on the
deer (Table 1). Snowmobile activity brought varied responses from deer
with severity of deer response apparently increasing with decreasing distance from the snowmobile (Table 1).
Increases in heart-rate could not be accurately measured if deer began to
walk or run. However, on 3 occasions increases in heart-rate were apparent
while deer were either standing still or bedded (Table 1, Fig. 9). These
increases in heart-rate suggested that although deer may not elicit strong
overt behavioral reactions to harassment, "flight" response mechanisms may
be active, indicative of some degree of stress.
Visual contact by deer with forms of harassment preceded overt responses.
These deer appeared to rely on sight more than smell or hearing as a defense
mechanism.
This segment's work revealed problems with measuring heart-rate during
certain deer activities, demonstrated that semi-tame deer at pasture will
react to harassment, and that heart-rate may be a sensitive measure of nonovert reactions of deer to harassment.

�-101-

Table l. Reactions of deer to various types of harassment, Hidd1e Park,
Colorado, 1977 .

Harassment
One
Person
Walking

Date

Deer
No.

Distance
From
Pasture

Overt
Behavior
Reaction

Apparent
Increase
In Heart
Rate

2-21-77
2-21-77
2-21-77
2-21-77

39
18
39
18

300-400 yds
200 yds
100 yds
0 yds

a
1
1
1
1

Yes
Yes
Yes
No

Change
In Deer
Activity

Duration
of
Harassment
(Hinutes)

No
Yes
No
No

13c
11
c
9
11

--------------------------------------------------------------------------------1
1
1
1
0

No
Yes
No
Yes
No

No
Yes
No
Yes
No

9
10
9
13
8

3

Yes

Yes

26

1
1

No
Yes
No

No
No
No

300-400 yds
100-300 yds
0-1700 yds

1
2
1

No
Yes
No

No
Yes
Yes

12

300-400 yds
0-400 yds

o

No
Yes

No
Yes

12

3-07-77
3-07-77
3-07-77
3-07-77
3-07-77

18
39
18
39
18

300-400 yds
200 yds
100 yds
0 yds
b
0 yds

3-08-77

39

100 yds

Person
Plus Dog
Walking

3-09-77
3-09-77
3-09-77

18
39
18

300-400 yds
200 yds
100 yds

1

Person
On
Snowmobile

3-18-77
3-18-77
3-18-77

39
18
39

Person
On
Snowmobile

3-19-77
3-19-77

18
39

Two
Persons
Walking
Two
Persons
Walking

b

2

7

23

11

a

o = no overt reaction; 1 = ears-up, alert appearance, no movement away from
harassment; 2 = ears-up, alert appearance, moderate movement; 3 = ears-up, alert
appearance, strong movement.
b

Route located e1evationa11y above pasture.
c

Increases in heart-rate possibly indicate of "flight" response.

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~p 0;,},"'=":.&amp;r:'0 ! ~:
v ~,
-----....-.:..../ \"..~,,-,

Y

,

.20 BPM~
t""'

-~

!

~ ~~~,

I

j;:!,

1

r:

...

: ~

: . , "
.

'

,. ~.

,-.-!"

"

DEER

..

I ~,

.·'r~~i---f--·-~~-'! i,

c.

(",

..~

,

speed

"

.

1.6/min

o ~ ~ ~ ~ n ~ ~ ~ 0 0

~

-v
.,

"11j

Fig. 9. Heart-rate trace of deer No. 39 while 2 persons were walking 200 yards from
pasture. Long delay in heart-rate decrease indicative of increased heart-rate while
initially alert.

�-103-

LITERATURE CITED
Booyens, J., and G. R. Hervey. 1960.
measuring metabolic rate in man.
38:1301-1309.

The pulse rate as a means of
Can. J. Biochem. Physiol.

Bradfield, R. B., P. B. Huntzicker, and G. J. Fruehan. 1969. Simultaneous
comparison of respirometer and heart-rate telemetry techniques as
measures of human energy expenditure. Am. J. Clin. Nutr. 22:696-700.
Corbett, J. L., D. J. Farrell, R. A. Leng, G. L. McClymont, and B. A.
Young. 1971. Determination of the energy expenditure of penned and
grazing sheep from estimates of carbon dioxide entry rate. Br. J.
Nutr. 26:277-291.
______ , R. H. Leng, and B. A. Young. 1969. Measurement of energy expenditure by grazing sheep and the amount of energy supplied by volatile
fatty acids produced in the rumen. Pp. 177-186 In K. L. Blaxter,
J. Kielanowski, and G. Thorbek (eds.). Energy metabolism of farm
animals. Oriel Press, Newcastle upon Tyne.
Cupal, J. J., A. L. Ward, and R. W. Weeks. 1974. A repeater type biotelemetry system for use on wild big game animals. Bio.Med. Sci.
Instrumentation 10:145-152.
Datta, S. R., and N. L. Ramanathan. 1969. Energy expenditure in work
predicted from heart rate and pulmonary ventilation. J. Appl.
Physiol. 26:297-302.
Dorrance, M. J., P. J. Savage, and D. E. Huff. 1975. Effect of snowmobiles on white-tailed deer. J. Wildl. Manage. 39(3):563-569.
Eckstein, R. G., and o. J. Rongstad. 1973. Effects of snowmobiles on
the movements of white-tailed deer in northern Wisconsin. Midwest
Fish and Wildl. Conf. Proc. 35:39.
Geist, V. 1971.
14:12-13.

Is big game harassment harmful?

Oil Week.

June.

Jacobsen, N. K. 1973. Physiology, behavior, and thermal transactions of
white-tailed deer. Ph.D. Thesis. Cornell University. 346p.
Morhardt, J. E., and S. S. Mbrhardt. 1969. Field measurements of mammalian
metabolic rates using radiotelemetry of heart rates. Am. Zool. 9:1093.
Owen, R. B., Jr. 1969. Heart rate, a measure of metabolism in bluewinged teal. Compo Biochem. Physiol. 31:431-436.
Webster, A. J. F. 1967. Continuous measurement of heart rate as an indicator of the energy expenditure in sheep. Br. Jr. Nutr. 21:769-785.
White, R. G., and R. A. Leng. 1968. Carbon dioxide entry rate as an
index of energy expenditure in lambs. Proc. Aust. Soc. Anim. Prod.
7:335-341.

�-104-

Wyndham, C. H.,N. B. Strydom, J. S. Maritz, J. F. Morrison, J. Peter,
and Z. U. Potgieter. 1959. Maximum oxygen intake and maximum
heart rate during strenuous work. J. Appl. Physiol. 14(6) :927-936.
Young, B. A., and J. L. Corbett. 1968. Energy requirements for maintenance of grazing sheep measured by calorimetric techniques.
Proc. Aust. Soc. Anim. Prod. 7:327-334.
______ , R. H. Leng, R. G. White, G. L. McClymont, and J. L. Corbett.
1969. Estimation of energy expenditure from measurements of carbon
dioxide entry rate. Pp. 435-436 In K. L. Blaxter, J. Kielanowski,
and G. Thorbek (eds.). Energy metabolism of farm animals. Oriel
Press, Newcastle upon Tyne.

�July, 1977

-105-

JOD PROGRESS

COLOR..;\.D0

State of
No.

W-38-R-32

h'orK

Plan No.

16

Job

Title

Project

EEPO!ZT

Estimating

Pe r Iod Covered:
Personnel:

Deer-Elk

Investigations

1-2-3
Job No. ---------_._---_
Piceance Deer Study Parameters of Population Dynamics

..

_--- .-

April 1, 1976 through March 31, 1977.

R. M. Bartmann, J. J. Klein, Jr., D. J. Freddy, L. H. Carpenter,
R. B. Gill, W. S. Brown, D. Casado, J. D. Erickson, G. L. Pepperd,
and J. M. Scotese.

ABSTRACT
Seventy sightings and 32 recoveries of banded deer were recorded from March
1976 through February 1977. The winter quadrat deer census in Game Management Unit 22 was cancelled due to a lack of snow cover. Therefore, the predicted population of 29,820 will be the basis for the 1977 population
calculations.
A candidate sampling design based on randomly located squaremile quadrats was implemented for both ground and air post-season deer sex
and age classifications.
This work was done cooperatively with the Experimental Deer Inventory Project (Work Plan 22, Job 1) and results are reported
under that project.
The previous non-random post-season survey was continued
and yielded a buck:doe:fawn ratio of 22:100:67 based on 1,030 deer classified.
Application to the predicted population figure gives 3,459 bucks, 15,775 does,
and 10,586 fawns. The 1975-76 winter mortality estimate of 1,915 deer was
eight percent of the estimated 1975-76 winter population.
The combined
archery and rifle deer harvested in Unit 22 was estimated at 3,249 bucks
which is a 47 percent increase over 1975's harvest.

��-107-

ESTIMATING

PICEANCE DEER STUDY PARAMETERS OF POPULATION

DYNAMICS

R. M. Bartmann

P.N.O. OBJECTIVE
To develop and test a method for estimating
deer populations in the Piceance Basin.

density

of over-winter

mule

SEGMENT OBJECTIVES
1.

To collect

and tabulate deer band sighting

and recovery

2.

To estimate the size of the wintering
Management Unit 22.

deer population

in Game

3.

To estimate the sex and age structure
in Game Management Unit 22.

of the wintering

deer population

4.

To estimate

the annual productivity

5.

To estimate

the annual winter

6.

To estimate

the annual hunter harvest

of the Piceance

mortality

data.

deer population.

of the Piceance

of the Piceance

deer population.

deer population.

METHODS AND MATERIALS

Deer Banding
See Bartmann

(1972).

Deer Density
See Bartmann

(1974a).

Population

Structure

The previous method for obtaining deer sex and age structure data involved
flying in a somewhat systematic manner over most of the occupied deer areas
within Game Management Unit 22 (Bartmann 1974a).
Although many animals were
classified, the lack of a valid sampling scheme precluded statistical analysis
of the data.
This past winter, in cooperation with the Experimental Deer Inventory Project (Work Plan 22, Job 1), sampling procedures based on randomly located
square-mile quadrats were implemented for both ground and air classifications.
Methodology and results of these new procedures are reported under Work Plan
22, Job 1 in this Game Research Report.
For comparative purposes, the
previous non-random method was repeated.

�-108-

Productivity
See above (Population Structure) and also Bartmann (1975).
Winter Mortality
See Bartmann (1974b and 1975).

Hunter Harvest
See Bartmann (1974b).
RESULTS AND DISCUSSION
Deer Banding
Seventy sightings and 32 recoveries of banded deer were recorded fromMarch
1976 through February 1977 (Tables 1 and 2).
Deer Density
Insufficient snow cover to force deer down onto the "normal" wintering areas
and to provide a good counting background forced cancellation of the 1976-77
winter count. By mid-February, only 10 inches of total snowfall was recorded
at Little Hills. Total snowfall for the same period the previous five
winters ranged from 36 to 56 inches and averaged 45 inches. While several
snowfalls did occur after mid-February, they were poorly distributed over the
winter range and melted quickly in the warm weather.
The predicted 1976-77 winter population in Game Management Unit 22 is 29,820
(Table 3). Application of the 1976 post-season buck:doe:fawn ratio of 22:100:67
gives 3,459 bucks, 15,775 does and 10,586 fawns. In lieu of a census estimate,
these figures will provide the bases for the 1977 popuiation calculations.
Population Structure
Results of the experimental post-season classification sampling procedures
generated more questions than they answered. For this reason, results from
the non-random procedure will continue to be used. This year, classifications
were made December 11 by two observers from a Bell 47G3B helicopter. Snow
cover was spotty at lower elevations and fair at higher ones. Estimated
ratios for each observer were similar. Both buck:doe ratios were up slightly
from previous years while fawn:doe ratios were lower. Again to preserve

�-109-

continuity,

only my data are used.

Observer

Bucks

Does

Fawns

Total

Buck:Doe: Fawn

R. Bartmann
D. Freddy

120
129

545
608

365
396

1,030
1,133

22:100:67
21:100:65

Productivity
The 1976 hunting season was bucks only, so the post-season fawn:doe ratio
of 67:100 was used without adjustment to give a pre-season production of
10,778 fawns.
The adult doe population contained an estimated 27 percent
yearlings.
Removal of these from the doe population resulted in a fawn:
mother ratio of 92:100.
While slightly lower than in the two previous
years, 107:100 in 1975 and 96:100 in 1974, the differences are difficult
to interpret without additional information.

Winter Mortality
The 1976 deer winter mortality survey was made April 16 through May 27.
Twenty carcasses judged to be of the current year were found for a mean
0.36 deer per plot or 1,915 + 692 total dead deer (90 percent C.L.) on the
winter range (Table 4). This is an eight percent loss based on the estimated 1975-76 winter population.
This is the lowest loss recorded in the
four years the survey has been conducted and the 1975-76 winter was also
the mildest of the past four.
Composition of the estimated loss with unknowns apportioned among the
various classes was 527 male fawns, 526 female fawns, 670 mature females
and 192 mature males.

Hunter Harvest
The 1976 regular deer season in Game Management Unit 22 was antlered only
and ran from October 30 through November 9. The archery season, also
antlered only, ran from August 21 through September 23. The total harvest
estimate from the Game Management Section's random survey was 3,221 and 28
for the regular and archery seasons, respectively.
This is a total increase
of 1,037 deer, 47 percent. over the 1975 estimated harvests.
All figures
are inflated by 25 percent to acknowledge wounding loss and illegal kill.
A change in the days and hours of operation of check stations, plus the
addition of two "satellite!' stations at Dinosaur and Rabbit Valley, disrupted the continuity of some data. All stations, except Idaho Springs,
were only operated during the third through the seventh days of the season
for 10 to 14 hours per day. Idaho Springs was operated the first nine days
as in the past but was only open 14 hours per day instead of 24.

�-1l0-

At Idaho Springs, 516 bucks were checked compared to 562 in 1975. Additional deer passing the station after it was closed may have boosted the
check to near what it was last year, but no data are available that might
indicate what adjustment would be appropriate.
On the basis of the above
figures, an eight percent decrease is indicated compared to the 47 percent estimated harvest increase between the two years.
The two "satellite"
stations contributed 318 bucks, or one-third of the statewide check of
936 for Unit 22.
According to the Division statistician in charge of surveys, it is not
possible to retrieve from the computer a breakdown of deer field-ages by
individual check station even though they are entered this way.
Therefore,
a between-years comparison of yearlings in the Idaho Springs check cannot
be made.
The percent yearlings field-aged at the three regular stations
combined and the two "satellite" stations were 62 and 57 percent, respectively.
The predicted yearling percentage in the adult buck population
was 71.

LITERATURE

CITED

Bartmann, R. M. 1972. Piceance deer study - population distribution.
315-337.
In Game Research Report.
Colo. Div. of Wildl., Denver.
3(Part 3):253-377.
(proc.)

P.

1974a.
Piceance deer study - population density and structure.
P. 363-370.
In Game Research Report.
Colo. Div. of Wildl., Denver.
3(Part 2) :185-398.
(proc.)
1974b.
Piceance deer study - productivity and mortality.
380. In Game Research Report.
Colo. Div. of Wildl., Denver.
185-398.
(proc.) •

P. 3713(Part 2):

1975. Piceance deer study - productivity and mortality.
362. In Game Research Report.
Colo. Div. of Wildl., Denver.
195-504.
(proc.).

P. 3553(Part 2):

Prepared

bYKJ1~
R. M. Bartmann
Wildlife Researcher

C

�-111-

Table 1. Sightings of deer marked on the winter range in Game Management Units 11, 21, 22 and 23, received from March 1976 through February
1977 .
Date
(1975-76)

7- ?-75
10- ?
10-12
10-12
10-12
10-12
10-13
11- ?
11- 5
2-16-76
3-13
3-14
3-19
3-20
3-22
3-22
3-22
3-26
3-26
3-29
3-30
3- ?
4- 9

4- 9
4-14
4-14
4-17
4-17
4-20
4-20
4-20
4-20
4-24
4-29
5- 1
5- 1
5- 1

5- 1
5- 7
5-24
9-11
10-30
10-30
11- 1

Neckband
Color

Number

Twp.

White
Orange &amp; white blocks
Yellow
Blue
Yellow
White
Orange &amp; white blocks
Green wlwhite stripe
White
Blue &amp; yellow blocks
Blue
Blue &amp; yellow blocks
Blue &amp; yellow blocks
Yellow
White wIred stripe
White wIred stripe
White wIred stripe
Yellow
Yellow
White
Yellow wIred stripe
White wIred stripe
Yellow wIred stripe
Yellow wIred stripe
White wIred stripe
Red &amp; blue blocks
Pink
White
Green wlwhite stripe
Yellow wIred stripe
Blue &amp; white
Blue &amp; white
Green wlwhite stripe
Yellow
Yellow
White wIred stripe
White wIred stripe
Yellow wIred stripe
Yellow wIred stripe
Orange &amp; white blocks
Blue
White
Yellow &amp; green blocks
Pink

135
12
?
9
?
?
13
?
?
?
2
40
?
54
14
120
133
12
23
?
110
39
82
15
34
190
75
157
237
109
5
106
237
79
53
38
145
27
101
39
?
?
?
4

4N
3N
2N
6N
6N
4N
3N
2N
4N

IN
IN
IN
IS
IS
2N
2N
2N

IS
IS
IS
IN
IN
IS
IS
2N
2N
2S

IS
IN
IS
IS
IS
IN
IS
IS
IN
IN
IN
IN
IS
4S
3N

IN
3S

Location
Range

Sec.

91W
89W
88W
90W
90W
93W
89W
94W
92W
93W
94W
92W
92W
92W
96W
96W
96W
93W
93W
98W
97W
94W
96W
96W
97W
97W
96W
97W
96W
97W
97W
97W
97W
93W
93W
96W
96W
95W
96W
93W
100W
95W
lOOW
96W

---------------------------------------------------------------------------

29
16
12
22
22
22
16
6
9
20
22
14
29
30
20
20
20
14
14
18
36
35
4
4
2
2

29
15
31
11
11
21
36
3
2
36
36
28
31
7

25
17
21
17

�-112-

Table 1. Sightings of deer marked on the winter range in Game Management Units 11, 21, 22 and 23, received from March 1976 through February
1977 (continued).
Date
(1976-77)

Color

11- 2-76
11- 2
11- 4
11-15
11- ?
11- ?
12- 5
12- 5
12- 5
12- 5
12- 5
12- 5
12- 5
12- 5
12- 5
12- 5
12- 5
12- 5
12-7
12-10
12-10
12-10
12-10
1- 8-77
2-20-77
2-27-77

Red &amp; blue blocks
Yellow &amp; green blocks
Red &amp; blue blocks
White wired stripe
Red &amp; blue blocks
Yellow
Yellow &amp; green blocks
Blue
White
White
Yellow wired stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Green w/white stripe
Blue
Green w/white stripe
Green w/white stripe
White
Yellow
Blue &amp; white
White
Yellow wired stripe
Yellow wired stripe
Yellow
Yellow

Neckband
Number

Twp.

Location
Range

Sec.

?
?

3N
2N
IN
IN
2S
2S
IN
IN
2N
2N
2S
2S
IS
IS
IN
2N
IS
IS
2S
IS
IS
2S
2S
IS
IN
IN

94W
100W
94W
97W
92W
92W
99W
99W
98W
98W
94W
95W
95W
95W
96W
99W
95W
95W
94W
96W
97W
99W
96W
96W
96W
96W

34
21
2
1
7
7
6
5
11
4
6
12
6
6
7
29
12
12
18
5
24
7
4
15
23
23

6
?
?
?
?
?

12
14
?
?
?
?
?
?
?

?
?

26
?
?
?
?

80
77

Correction to Banding Data
1- 7-72

Blue neckband #30, eartags L-1158 and L-1159 trapped T2N,
R100W, Sec. 24, should be recorded as a Male Fawn.

�-113-

Table 2. Recoveries of deer marked on winter range in Game Management
Units 11, 21, 22 and 23, received from March 1976 through February 1977.
Date
(1975-76)

Color

Number

Twp.

Location
Range

Sec.

10- ?-75
1-15-76
4- 7
4-15
4- ?
3- ?
3-23
5- ?
5- ?
5- ?
10-17
10-29
10-30
10-30
10-30
10-30
10-31
11- 1
11- 1
11- 1
11- 2
11- 2
11- 2
11- 2
11- 4
11- 4
11- 6
11- ?
11- ?
11- ?
1- ?-77
2-21-77

Blue
Blue
Blue
Yellow wired stripe
Blue &amp; yellow blocks
Yellow
Red &amp; blue blocks
White wired stripe
Yellow wired stripe
Red &amp; blue blocks
Green &amp; yellow blocks
Yellow wired stripe
Green &amp; yellow blocks
Yellow wired stripe
Green w/white stripe
Red w/white stripe
Green &amp; yellow blocks
Green w/white stripe
White wired stripe
Red w/white stripe
Green w/white stripe
Yellow
Yellow
Green w/white stripe
White
Red &amp; blue blocks
White wired stripe
Blue &amp; yellow blocks
White wired stripe
Red w/white stripe
White wired stripe
Orange &amp; white blocks

7
39
120
39
7
49
163
58
66
31
53
31
55
75
L-3272
65
51
L-3270
65
L-1964
208
45
46
L-3282
174
70
59
33
43
71
115
86

2N
3N
3N
IS
IN
IN
IN
2N
IS
IN
2N
2S
IN
IN
IN
2S
3N
IN
2N
IS
IS
2S
IS
IS
IN
IN
IS
IN
8N
2S
2N
IS

97W
93W
99W
96W
94W
94W
96W
97W
97W
94W
98W
96W
95W
96W
96W
97W
97W
96W
98W
92W
95W
87W
93W
96W
98W
95W
97W
92W
90W
96W
97W
93W

24
2
35
12
29
32
5
36
13
18
12
2
28
31
31
9
14
27
4
26
1
33
2
12
36
29
1
18
15
25
28
5

Neckband

�-114-

Table 3. Calculations to predict the 1976-77 winter deer population in
Game Management Unit 22.

Bucks

Does

Fawns

Total

Est. 1975-76 winter pop.

1,985

12,345

9,876

24,206

Est. 1976 winter mortality

192

670

1,053

1,915

1976 summer population

1,793

11,675

8,823

22,291

Fawns apportioned 50:50

4,412

4,411

Adjusted 1976 summer pop.

6,205

16,086

1976 fawn production est.
(67 fawns:l00 does)

10,778

1976 pre-season pop.

6,205

1976 harvest estimate

3,249

Predicted 1976-77 winter
population

Predicted population
corrected for 1976
post-season buck: doe: fawn
ratio (22:100:67)

16,086

10,778

33,069

2,956

16,086

10,778

29,820

3,459

15,775

10,586

29,820

�-115-

Table 4. Results of dead deer searches on 56 1/8-mi1e square quadrats
on winter range in Game Management Unit 22 during spring, 1976.
The
39 quadrats with no dead deer are omitted.

Quadrat

2

Male

Adults
Female

Unknown

Male

Fawns
Female

1

13

1

21
24

1

1

1

1

35

1

40

1

42

1

65

1

75

2

76

1

107
110

1

1

25
26

Unknown

1
1

III

1

114

1

115

1

Totals

5

4

1

2

8

��July, 1977

-117JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

W-38-R-32

Deer-Elk

Work Plan No.

16

Job No.

Job Title Piceance

Deer Study - Food Habits Technique

Period Covered:
Personnel:

Investigations
5

----------------------------Development

April 1, 1976 through March 31, 1977

R. M. Bartmann, J. J. Klein, Jr., L. H. Carpenter,
P. H. Neil, D. R. Reichert and W. R. Regelin.

R. B. Gill,

ABSTRACT
Observations of "naturally" versus "artificially" maintained tame deer
forage selections were made only in January.
The February trials were
cut short and the March o~es cancelled because the artificially maintained
tame deer became untractable.
During the January 4-12 trials, the pasture
deer took 24,250 bites of 56 different species while pen deer took 9,422
bites of 40 species.
Because of the above-mentioned deer handling problems
and a paucity of snow cover, the series of three grazing trial comparisons
will be repeated next winter.
Procedures will be revised as much as
practical based on results of statistical analyses which are not yet completed.

��-119-

PICEANCE

DEER STUDY - FOOD HABITS

TECHNIQUE

DEVELOPMENT

R. M. Bartmann

P.N.O. OBJECTIVE
To determine if forage selections by "artificially" maintained
are similar to those of "naturally" maintained tame deer.

tame deer

SEGMENT OBJECTIVES
1.

To raise tame deer for grazing comparisons.

2.

To develop an herbarium

3.

To maintain

pasture

4.

To conduct

grazing comparison

of plants in the study area.

fences.

METHODS
Fences around the 98-acre
repairs made and predator
Injury to personnel

trials.

AND MATERIALS

pasture were checked
entry holes plugged.

resulted

in incomplete

in the spring

plant collections

and fall and

in the pasture.

Six of the eight yearling deer used in the 1976-77 winter grazing comparisons
had been used as fawns in the October, 1975 trials.
These animals were
maintained at the Fort Collins pen facilities.
On December 10, 1976 the
four least tractable of the eight, one female and three male castrates, were
brought to Little Hills and placed in a 98-acre pasture.
After one week,
all supplemental feed was removed and the deer were forced to subsist on
native forage.
The other four deer, two females and two male castrates, were
brought to Little Hills January 3, 1977 and placed in the pens by the headquarters where they were provided hay and concentrate feed ad libitum.
Grazing trials were started January 4 and completed January 12. The original
10-day schedule was shortened to nine by the addition of two mid-day trials.
Each day during early morning and again in mid-afternoon,
two pen deer were
transported and grazed with the pasture deer.
Four observers watched two
deer from each group for a total of 40 trials per treatment.
A trial is
defined as one observer following one deer for one to two hours and recording the number of bites by species.
Observations of each deer were
split as equally as practical between morning and afternoon periods and
observers.
Handling problems forced substitution of a 3-1/2-year-01d male
castrate for one of the pen deer for three trials.

�-120-

After the January trials, the pasture deer were left in the pasture and
the pen deer were maintained in the pens at Little Hills until the next
trials which began February 8. The same procedures were followed except
that two pastures were used instead of one. Six deer and four observers
in one area proved too cumbersome and the pen deer were flighty which
affected each group's performance.
The two groups were grazed in separate
pastures with the intent of switching pastures halfway through the trials.
However, the pen deer became harder to handle and caused cancellation of
the trials after the second day. The March trials were also cancelled.

RESULTS AND DISCUSSION
Snow cover during the January trials ranged from bare ground to about four
inches on steeper north slopes.
Minimum temperatures measured at the Little
0
Hills Station ranged from -16 to 220F and maximums 220 to 38°F. Light
snow occurred on three of the nine days. Minimums and maximums for February
0
o
were near 0 and 50 F, respectively, with no precipitation.
Snow depths
were similar to those in January except for a smaller proportion of bare
ground.
In January, pasture deer took 24,250 bites of 56 different species, exclusive
of unknowns, while pen deer took 9,422 bites of 40 species (Table 1). The
range in number of bites per trial was 85 to 1,599 for pasture deer and 7
to 605 for pen deer.
Twenty-three species eaten by pasture deer were not
eaten by pen deer. Most comprised less than one percent of the total bites.
All seven species used by pen but not pasture deer comprised 0.2 percent
or less of the total bites.
Mean trial lengths were similar, 78 and 77
minutes for pasture and pen deer, respectively.
In February, the pasture deer took 5,858 total bites in eight trials compared to 870 for the pen deer (Table 2). The pasture deer ate 37 species,
exclusive of unknowns, and the pen deer 16 species.
Comparison of forage
selections between the two periods is cautioned against due to the large
differences in total bites and the use of different pastures.
Statistical analyses of the January grazing comparison data are not completed.
When the three series of comparisons are rescheduled for the 1977-78 winter,
procedures will be revised as much as practical on the basis of these results.

Prepared

by /~

-rJ~~-----R. M. Bartmann
Wildlife Researcher

C

�-12l-

Table 1. Number of bites, by species, taken by pasture
during grazing comparison trials, January 4-12, 1977.
Species

Purshia

tridentata

Chrysothamnus

viscidif10rus

Poa species

and pen deer

Pasture

Pen

15,041

5,476

2,278

576

1,021

21

Tetradymia

canescens

915

140

Gutierrezia

sarothrae

801

140

Ame1anchier

utahensis

683

1,420

636

4

332

623

Commandra

umbe11ata

Cercocarpus
Berberis

montanus

312

repens
grass

259

26

1udoviciana

240

116

Unidentified
Artemisia

Bromus inermis

162

Linum lewisii

159

43

Hap10pappus

154

108

142

366

141

12

Quercus

nutta11ii

gambe11ii

Unidentified

forb

Stipa comata

128

Lepidium

montanum

116

17

Artemisia

tridentata

116

21

Chrysothamnus

nauseosus

107

42

Symphoricarpos

oreophi1us

72

100

1anata

61

1

Circium species

50

12

Hedysarum

borea1e

47

Oenothera

coronopifo1ia

44

2

Oryzopsis

hymenoides

22

10

Juniperus

scopu1orum

22

Tragopogon

dubious

21

Chrysopsis

vi110sa

18

Eurotia

5

Pinus edu1is

12

2

Cryptantha

sericea

11

1

Koe1eria

cristata

10

3

Physaria

species

9

------------------------------------------------------------------------------

�-122-

Table 1. Number of bites, by species, taken by pasture and pen deer
during grazing comparison trials, January 4-12, 1977 (continued).

Pasture

Pen

9

11

Aster species

8

3

Artemisia

frigida

8

Agropyron

species

7

15

Juniperus

osteosperma

6

1

Erigeron

species

6

Species
Euphorbia

robusta

Chaenactis

douglasii

6

Penstemon

fremontii

5

Cilia aggregata

5

Bouteloua

5

gracilis

Rosa woodsii

4

Penstemon

4

Lupinus

caespitosa
caudatus

4

9

18

Lithospermum

ruderale

4

Descurainia

species

4

B romus anomal us

4

Aster leucanthemifolius

4

Balsamorhiza

3

Artemisia

sagittata

dracunculus

3

3

Phlox hoodii

2

15

Helianthus

2

6

eatoni

2

tectorum

1

Erigeron
Bromus

annuus

Arabis holboellii

1

Antennaria

1

Sitanion

species

5

hystrix

Ribes species

7

Polygonum

sawatchense

7

Eriogonum

urnbellatum

11

Elymus

3

cinereus

Agropyron

spicatum

17

Achillea

lanulosa

4

Total Bites

24,250

9,422

�-123-

Table 2. Number of bites, by species, taken by pasture
during grazing comparison trials, February 8-9, 1977.
Species

and pen deer

Pasture

Pen

Purshia

tridentata

4,077

374

Quercus

gambe11ii

583

223

Artemisia

tridentata

438

5

Tetradymia

canescens

213

14

Artemisia

frigida

130

1

112

7

Chrysothamnus

viscidif10rus

Artemisia

1udoviciana

108

6

Cercocarpus

montanus

38

73

Gutierrezia

sarothrae

36

Linum lewisii

22

Ame1anchier

utahensis

12

137

Unidentified

grass

10

2

Aster 1eucanthemifo1ius

9

Chrysothamnus

8

4

8

5

Eriogonum

nauseosus

umbe11atum

Circium

species

7

Lepidium

montanum

7

Hap10pappus
Berberis

nutta11ii

repens

7
5

Cryptai1.thasericea

5

Juniperus

4

scopu1orum

Unidentified
Physaria

browse

species

4
3

Aster species
Lithospermum rudera1e

2

Symphoricarpos

2

Unidentified

oreophi1us

2

forb

2

Eriogonum

lonchophyl1um

1

Oryzopsis

hymenoides

1

Pinus edulis

4

1

3

2

Ephedra viridis

4

Juniperus

6

osteosperma

Total Bites

5,857

870

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                  <text>-1-

October 1977
JOB PROGRESS

State of

COLOPJWO
------------~--~-----------

Project

No.

W-88-R-22

-------------------------

Work Plan No.

I

Job Title

Waterfowl

Period Covered:
Personnel:

REPORT

Migratory
Job No.

Production

Bird Investigations
1

----------------------------------

Surveys

March 31, 1976 to June 30, 1976

C. Bryant and staff, Monte Vista National Wildlife Refuge;
F. N. Folks, Utah State Division of Wildlife Resources;
R. Clark, J. Corey, W. Dolezal, B. Goforth, G. Hinshaw,
R. Hopper, J. Lorentzson, R. Lowry, W. Russell, G. Saville,
R. Velarde, R. Weldon, M. Zgainer, S. Steinert, and M.
Szymczak.
ABSTRACT

Water conditions for waterfowl production were below average in all survey
areas. The total number of duck breeding pairs was estimated at 71,021. This
is a 9.3 percent reduction from the record high in 1975, but still about
22 percent above the long term average.
Increases in the number of estimated
breeding pairs were recorded in the San Luis and Cache 1a Poudre valleys.
Major declines occurred in North Park and the South Platte Valley.
Gadwall
CAnas strepera), green-winged teal (Anas crecca), and shoveler CAnas c1ypeata)
showed increases from 1975. All other species were down from 1975 levels.
The
post-nesting season population of Canada geese (Brant a canadensis) in northwest Colorado was estimated to be similar in size to that of 1975 but nearly
31 percent above the long-term average.
Canada goose production in northcentral Colorado was up 23 percent from 1975 and 14 percent above the longterm average, however the total number of adult geese observed was do~m 13
percent from 1975 and 20 percent from the long-term average.
Two new surveys
were added in lQ7n. The first was a helicopter survey of Canada goose breeding
pairs in the San Luis Valley, from which we estimated a total 92 breeding pairs.
The second was a Canada goose breeding pair survey conducted in west central
Colorado with a total of 172 birds observed.

��-3-

WATERFOWL

PRODUCTION

SURVEYS

Michael R. Szymczak
Steven F. Steinert

P.N.O. OBJECTIVES
1.

To estimate the number of duck breeding pairs, by species,
major waterfowl nesting areas in Colorado.

on selected

2.

To estimate the number of goose breeding pairs, and in some cases,
obtain production data on selected goose nesting areas in Colorado.

3.

Compile
federal

data and submit reports to appropriate state personnel and
agencies for use in establishing hunting season recommendations.

SEGMENT

OBJECTIVES

1.

To estimate the number of duck breeding pairs, by species, in the San
Luis, Cache la Poudre, South Platte and Yampa Valleys, and in North
Park and Brown's Park.

2.

To estimate the number of goose breeding pairs in the San Luis Valley
obtain goose breeding pair and production data on the Yampa, Little
Snake and Green Rivers in northwest Colorado and in north central
Colorado.

3.

Compile

data and submit appropriate

METHODS

and

reports.

AND MATERIALS

Present duck breeding pair and production
inventory of only major production areas.

surveys

consist

of a breeding

pair

The 1976 duck breeding pair surveys were conducted during the period of May 3
to June 30. Surveys in North Park and the Cache la Poudre and South Platte
valleys were conducted exclusively from the air. Ground counts were made in
the Yampa Valley and Brown's Park.
In the San Luis Valley intensive aerial
counts were adjusted for visibility by air-ground comparison studies.
Pair
estimates for the Monte Vista National Wildlife Refuge in the San Luis Valley
were obtained from nesting transects.
All survey methods and sample areas
for ducks remained the same as in previous years.
Surveys of Canada goose production

were conducted

within

the period

of March

31

�-4-

to June 15. Production estimates for Moffat County were made from as complete
a count as possible of hatched or active nests and brood size. Population estimates of north central Colorado were obtained from counts of goslings and adults
conducted from the ground during the period in which the birds were flightless.
Population estimates for the Colorado River drainage were estimated from aerial
surveys.
In the San Luis Valley a survey of expected productive breeding pairs was
conducted by flying transects in defined sample blocks as described by Lowry
(1974). Four evenly-spaced transects were flown in each 1 x 3 mile selected
sample block on the Rio Grande and Conejos rivers, while a simple total count
was conducted in sample blocks containing ponds or lakes, only.
Each selected
section was flown with both fixed-wing and helicopter aircraft and the results
compared.
All fixed-wing flying was accomplished with a Cessna 185 aircraft and the helicopter flying with a Hughes 500. Two observers were used in sampling by
transects with the fixed-wing aircraft.
One observer was used for the helicopter flight and some of the block or section sampling with the fixed-wing
aircraft.

RESULTS AND DISCUSSION
Water conditions were considered below average for duck production in all
survey areas.
The San Luis Valley was very dry for the third consecutive
year, although water in the north end of the Valley showed some improvement
over 1975. As a result of a dry winter, water conditions were poor in both
North Park and the Yampa Valley, with very little additional runoff predicted
for the remainder of the spring and summer.
Surveys of the Cache la Poudre
and South Platte Valley areas were conducted prior to the initiation of the
irrigation season.
Water conditions in the Poudre and Platte valleys, although
poor during the early spring nesting season should improve for the late nesting
and brood rearing period.
The small and steady runoff in northwest Colorado
kept flooding loss of goose nests in Moffat County to a minimum.
A generally
mild spring with no severe storms provided excellent conditions for goose
production in north central Colorado.
The number of estimated duck breeding pairs in Colorado's production trend
areas declined in 1976 from the record high 1975 level.
The total of 71,021 is
9.3 percent below the 1975 level, but 21.3 percent above the long-term average
(Table 1). The number of estimated nesting pairs increased in the San Luis
and Cache la Poudre valleys.
Major numerical declines were recorded in both
North Park and the South Platte Valley (Table 1). Over 60 percent of the San
Luis Valley breeding population in 1976 was composed of shovelers and greenwinged teal. Normally green-wings are not prominent in the San Luis Valley
breeding population.
In North Park, no green-winged teal were observed on
aerial transects, therefore, the green-winged teal population which was estimated to be over 7,000 in North Park in 1975 was reported as non-existent in
1976. Obviously, the reliability of estimating green-winged teal populations
using these methods is suspect.

�-5Table 1. Summary of Colorado's duck breeding ground population estimates
in selected areas, 1976.

Total Estimated Breeding Pairs
Long Term
1976
1975
Average 1/

Percent Change
From
From Long
1975
Term Average

San Luis Valley

34,759

26,801

27,280

+29.7

+27.4

2/
North Park-

16,535

27,134

17,163

-39.1

- 3.7

South Platte Valley 10,062

14,152

6,623

-28.9

+51.9

Cache La Poudre
Valley

6,927

6,732

3,514

+ 2.9

+97.1

Yampa Valley

2,145

2,354

2,783

- 8.9

-22.9

Brown's Park

593

1 ,1721/

1,172

78,345

58,535

Area

Total

71,021

-49.4
- 9.3

+21.3

1/ San Luis Valley and North Park averages are based on results of 1964 through

,
b

1975 and 1968 through 1975 surveys, respectively, because of changes in survey
methods utilized prior to those dates. Figures for other areas are 22-year
averages.

1/ Aerial counts corrected by species from visibility ratios obtained in the
San Luis Valley.

1/ No count made in 1975. Figure presented is average of 1971-1974 period.
The percent of the estimated breeding population composed of mallards remained
at about 24 percent (Table 2). Because of the large estimated breeding population of shovelers in the San Luis Valley, that species was numerically the
largest breeding population in Colorado in 1976.
A total of 1,561 Canada geese were observed in Moffat County in northwest
Colorado in 1976 (Table 3). The total is essentially the same as was recorded
in 1975 (Table 4). Total birds observed on the Yampa River increased substantially over the 1975 level as a result of an increase in both gosling production
(Table 5) as did the number of adults. Gosling production in 1976 decliaed below
1975 levels on the Green River (Table 5) as did the number of adults. Gosling
production in 1976 increased over the 1975 level on the Little Snake River
(Table 5), but the number of adults observed declined. Overall, in northwest
Colorado in 1976 the production increased, but the size of the total population
remained comparable to 1975 levels.

�-6-

Table 2. Species composition of Colorado's 1976 duck breeding pair
population.

Number of Breeding Pairs
1954-75
1976
1975
Average

Percent S;eecies Com;eosition
1954-75
1976
1975
Average

Mallard

17,324

19,090

27,299

24.4

24.4

52.6

Blue-winged and
Cinnamon Teal

7,432

15,078

5,685

10.5

19.2

11.0

Gadwall

4,205

3,984

5,332

5.9

5.1

10.3

Pintail

2,980

5,213

3,635

4.2

6.7

7.0

Green-winged Teal

9,136

8,859

2,452

12.9

11.3

4.7

Shoveler

21,145

8,005

2,634

29.8

10.2

5.1

American Wigeon

2,750

4,868

1,065

3.9

6.2

2.1

Redhead

3,623

8,048

2,254

5.1

10.3

4.3

Other Divers

2,426

5,200

1,521

3.4

6.6

2.9

Totals

71,021

78,345

51,8771/

Species

J

~
~

1/ Species composition computed from data from all areas for the 22-year
period regardless of changes in survey method.

�-7-

Table 3. Number of Canada geese observed,
Moffat County, Colorado, 1976.

and estimated

production,

Nesting
Pairs

Non-Nesting
Birds

Total
Adults

Est. NO.
ll
Goslings-

Total
Birds

Craig-Juniper
Springs

22

l32

176

87

263

Juniper SpringsCross Mtn.

32

101

165

122

287

Lily Park

13

66

92

48

140

67

299

433

257

690

Brown's Park

36

200

272

133

405

Dinosaur Nat'l.
Monument

22

63

107

104

211

Sub-total

58

263

379

237

616

22

120

164

91

255

147

682

976

585

1,561

and number

of successful

Area
Yampa

Sub-total
Green River

Little

Snake River

GRAND TOTAL
~

II Calculated

using average brood size observed

nests.

~

Results of the 1976 Canada goose production census in north central Colorado
are presented by individual areas in Table 6. Gosling production increased
over 1975 levels on all trend areas except Boulder (Table 7). Yet the total
number of geese observed declined because of the reduction in the number of
adult geese observed (Table 8). Production in 1976 was 14 percent above the
1969-1975 average, but the total number of geese was down 12 percent during
the same period.
Projections resulting from breeding pair counts from a helicopter on sample
areas in the San Luis Valley indicated there were 92 active nesting pairs of
Canada geese in the San Luis Valley during the count period (Table 9), an
increase of 56 percent over the 1975 level.
Twenty-six percent more breeding
pairs were estimated with the helicopter than with the fixed-wing aircraft
(92 vs. 73), while the same number of sub-adult pairs were estimated from each
craft (Table 9). On comparable sample areas in 1976, 226 geese were observed
with the fixed-wing aircraft compared to 261 with the helicopter.
The results of a Canada goose breeding pair survey, begun in 1976 in west
central Colorado are presented in Table 10.

�-8-

Table 4.

Total Canada

geese observed,

Moffat

County,

1976.

Percent Chan8e
From 1956-1975
From 1975
Average

Area

1976

1975

1956-1975
Average

Yampa River

690

401

423

+72.1

+63.1

405

448

171

-9.6

+136.8

2ll

406

348

-48.0

-39.4

255

294

251

-l3.3

+1.6

1,561

1,549

1,193

+0.1

+30.8

Green River
Brown's

Park

Dinosaur Nat'l.
Monument.V
Little Snake
Riverll
Total

])

Not surveyed

II Not surveyed

Table 5.
1976.

until

1970.

until 1962.

Estimated

number

of Canada goose goslings,

Moffat

County,

Colorado,

Percent Change
From 1956-1975
From 1975
Average

1976

1975

1956-1975
Average

257

147

137

+74.8

+87.6

l33

176

67

-24.4

+98.5

104

106

117

-1.9

-ll.l

Little Snake
Riverll

91

68

78

+33.8

+16.7

Total

585

497

399

+17.7

+46.6

Area
Yampa River
Green River
Brown's

Park

Dinosaur Nat'l.
Monument.!'!

II

- Not surveyed

until 1970.

21
- Not surveyed

until 1962.

�Table 6.

Production
Area
Wellington

Results of the north central Colorado goose census, June, 1976.

Water Area
Terry Lake
Douglas Reservoir
Stewart Pond
1
Dry Creek Reservoir-I
North Poudre 111
North Poudre 112
North Poudre 115
Bureau of Standards
Pond 111
Divide 118
Elder Reservoir
Annex 118]:.1
Dale Pond
Van Sant Pond
Cobb Lake
Watson Lake
Curtis Lake
Water Supply and Storage 114

No.
Broods

Total
No. Goslings

Total No.
Adults &amp; Yearlings

Total
Birds

7
0
3
8
0
0
6

27
0
11
35
0
0
22

47
3
6
19
23
3
13

74
3
17
54
23
3
35

3
5
0
5
2
2
0
0
0
6

10
20
0
25
12
6
49
33
0
28

19
36
17
17
21
19
102
134
63
13

29
56
17
42
33
25
151
167
63
41

278

555

833

39
7
14
17
67

92
8
150
16
25

131
15
164
33
92

144

291

435

TOTAL
Loveland

Boedecker ~eservoir
Flatiron Gravel Pits
McNeil Reservoir
Reservoir 1112
Welch Reservoir
TOTAL

6
3
3
4
13

------------------------------------------------------------------------------------------------

I
1.0
I

�Table 6.

Results of the north central Colorado goose census, June, 1976 (continued).

Production
Area
Boulder

Water Area
Ish Lake
Haden Lake
Terry Lake
Faivre Ponds
Rest Home Pond
Valmont Reservoir
Boulder Valley Farm Pond
King Pond
Eddy Pond
Angus Ranch Pond

No.
Broods

Total
No. Goslings

Total No.
Adults &amp; Yearlings

Total
Birds

0

18

0
0
0
0
0
0
0

0

10
2
134
21
39
189
11
6
7
11

28
2
167
44
65
228
15
8
13
22

0
0

33
23
26
39
4
2
6
11

I
I-'

0
I

TOTAL
Fort Collins

Herring Lake
College Lake
Dean Acres
Claymore Lake
Sterling Gravel Pits
Lindenmeier Lake
Grey Lakes
Novak Reservoir
Winick Pond
Anderson's Pond
Parkwood Lake
Kitchel Lake
Timnath Reservoir
Romily Gravel Pit
Flatiron Gravel Pits

14
3
9
0
0
0
5
2
5
4
4
4
3
5
2

162

430

592

60
11
32
24
72
26
23
7
15
17
19
18
9
28
5

17
111
65
51
77
44
23
4
10
26
16
14
127
24
4

77
122
97
75
149
70
46
11
25
43
35
32
136
52
9

------------------------------------------------------------------------------------------------------

�~

Table 6.

.••....•. ~-

Results of the north central Colorado goose census, June, 1976 (continued).

Production
Area
Fort Collins
(cont.)

Water Area
Fossil Creek Reservoir
Schuelke Reservoir
Wolaver Ponds
Peterson Ponds
Maxwell Pond

No.
Broods

Total
No. Goslings

Total No.
Adults &amp; Yearlings

Total
Birds

0
2
1
1
4

0
7
6
3
19

14
2
3
18

14
9
9
6
37

401

653

1,054

64
13
21
19
17
7
4
0
6
0
4
5
58
0
6
10
2
47
17
0
2
2

41
20
15
12
143
127
2
64
51
40
44
46
31
4
9
17
84
7
169
32
28

105
33
36
31
160
134
6
9
70
51
44
49
104
31
10
19
19
131
24
169
34
30

304

995

1,299

1,289

2,924

TOTAL
Denver

Ketring Pond
Centennial Pond
Columbine Country Club
Chatfield Country Club
Bowles Lake
Kings Lake
Blackmer
Mohn Estates
Grant Lake
Colo. Blvd. &amp; Quincy
Marston Reservoir
Pinehurst Country Club
Bit-O-Sea
Bowles Reservoir #1
Ward Reservoir
Kendrick Lake
Federal Center
Sloan's Lake
Standley
Denver City Park
Reservoir #3
Tule Lake
TOTAL
GRAND TOTAL

15
3
5
5
3
2
2
0
2
0
2
1
12
0
2
3
1
9
4
0
1
1

3

9

-

l/ Birds actually found on a pond about 1 mile N.E. of Dry Creek in the S.W. 1/4 of Sec. 14.
1/ Birds actually found on pond located 1/2 mi. N. of the Annex and 1/2 mi. W. of No.8.

4,213

I
I-'
I-'
I

�-12Table 7. Number of Canada goose goslings produced in north central Colorado
production trend areas, 1976.

No. of Goslings
1969-1975
1975
Average

Percent Change
From
From
1975
1969-1975

Area

1976

Wellington

278

207

259

+34.3

+7.3

Ft. Collins

401

234

268

+71.4

+49.6

Loveland

144

103

86

+39.8

+67.4

Boulder

162

217

234

-25.3

-30.8

Denver

304

288

281

+5.6

+8.2

Total

1,289

1,049

1,128

+22.9

+14.3

Table 8. Number of adult Canada geese observed in north central Colorado
production trend areas, 1976.

No. of Geese

Percent Change
From
From
1975
1969-1975

Area

1976

1975

1969-1975
Average

Wellington

555

570

777

-2.6

-28.6

Ft. Collins

653

667

710

-2.1

-8.0

Loveland

291

318

220

-8.5

+32.3

Boulder

430

466

646

-7.7

-33.4

Denver

995

1,328

1,303

-25.1

-23.6

Total

2,924

3,349

3,656

-12.7

-20.0

�-13-

Table 9. Estimated number of breeding pairs of Canada geese in the San Luis
Valley, as determined from surveys with fixed-wing and helicopter aircraft,
1975 and 1976.

No. Indicated

Nesting

Pairs

No. Sub-adult

Pairs

Aircraft

1975

1976

1975

1976

Fixed-wing

2sl/

731/

65

74

Helicopter

59

92

59

74

1/ Incomplete

Table 10.

counts, not comparable

West Slope goose breeding

Area

to helicopter

data.

pair survey, March 31, 1976.

Singles

Geese Observed
Pairs
Groups

Total

Roaring Fork River from Basalt
to Glenwood Springs

3

2

18

25

Colorado River from Glenwood
Springs to Grand Junction

13

26

69

134

Gunnison River from Grand
Junction to the Black Canyon

2

4

6

7

7

98

172

Uncompahgre River from
Delta to Colona

Total

18

LITERATURE
Lowry, D. G.
Thesis.

Prepared

by

28 pro

CITED

1974. Canada geese in the San Luis Valley,
Colorado State Univ.
76 p.

~J2.~
Michael R. Szymc
Wildlife

Researcher

s£eillert

Steven F.
Wildlife Tech. II

Colorado.

M. S.

��-15-

October

1977

JOB FINAL REPORT

State of
Project

COLORADO

2

Hork Plan No.
Job Title
Period

Covered:

Personnel:

Migratory

W-88-R-22

No.

Bird Investigations

Job No. ·~
~~
__ ~2~
Experimental Studies on Improving
T_h
__e__S_t_a_t_u_s
__o_f
__C_a_n_a_d_a
__G_o_o_s_e
__P_o_p~u
__l_a_t_i_o_n_s

_
_

April 1, 1976 through March 31, 1977

Michael

R. Szymczak

ABSTRACT
All data for this project have been collected and essentially analyzed.
Some work has been accomplished on a Special Report concerning the establishment of Canada goose breeding po pu La t.Lons in Colorado.
This report will
summarize the entire effort and present information on current status of
each population.
Formulation and publication of this manuscript is included
as a segment objective under Work Plan 6, Job 1, Higratory Bird Publications
for the period April 1, 1977 through March 31, 1978. One major publication,
listed below, has resulted from this job.
Canada goose restoration along the foothills
Szymczak, M. R. 1975.
Colorado Div. of Wildl. Tech. Publ. 31. 64 p.
Colorado.

Prepared

by

-m.J..J 12 ~
Michael R. szymc~
Wildlife Researcher

a

d.

of

��October 1977

-17-

JOB PROGRESS

State of

REPORT

COLORADO
--------~~~~~-----------

Work Plan No.
Job Title

2

Job No.

Studies of Canada Goose Populations

-Period Covered:
Personnel:

Migratory

W-88-R-22

Project No.

Bird Investigations
6

---------------------------in Colorado

Transplant

Areas

April 1, 1976 to March 31, 1977

S. Brock, M. Conner, J. Corey, W. Dolezal, V. Graham, R. Hopper,
R. Leasure, J. Lorentzson, D. Nelson, R. Oakleaf, J. Tischbein,
R. Velarde, and M. Szymczak

ABSTRACT

All areas in which Canada geese (Branta canadensis) were banded in southern
Alberta east of 1110 longitude continued to contribute geese to the north
central Colorado harvest during the 1975-76 hunting season.
Birds from
Dowling Lake and the Edmonton-Camrose area in Alberta, the Cypress Hills
area of Saskatchewan, Phillips County and the Yellowstone-Big Horn rivers
in Montana, all documented Hi-Line nesting areas, were well represented in
the north central Colorado harvest.
A total of 825 Canada geese were banded
pre-season on production areas in northcentral
Colorado while 204 were
banded in the San Luis Valley.

��-19-

STUDIES OF CANADA GOOSE
POPULATIONS IN COLORADO TRANSPLANT
Michael

AREAS

R. Szymczak

P.N.O. OBJECTIVES
1.

To document the breeding
central Colorado.

range of Canada geese wintering

2.

To document the distribution of harvest and estimate the survival
rate of the resident foothills Canada goose population.

3.

To document the distribution
the San Luis Valley.

of harvest

in north

of Canada geese nesting

in

SEGMENT OBJECTIVES
la. Plot the banding locations of Canada geese (Branta canadensis)
banded outside northcentral
Colorado and reported recovered in
north central Colorado.
b. Participate in the cooperative
southern Alberta.

Canada goose banding

operation

2a. Trap and band 500 Canada geese on production
along the foothills north of Denver.

and moulting

b. Complete,
cards.

schedules

submit and file appropriate

banding

3a. Trap and band at least 200 Canada geese on production
in the San Luis Valley.
b. Complete,

submit and file appropriate

METHODS

banding

in

areas

and recovery

and moulting

schedules

areas

and recovery

cards.

AND MATERIALS

Band recoveries listed on computer printouts provided periodically by the
Migratory Bird Populations Station were sorted and plotted as to banding
and recovery location, sex, and age at time of banding, and estimated
breeding area for birds associated with northcentral
Colorado.
Banding of flightless adults and goslings was accomplished through drive
trapping operations in southern Alberta, north central Colorado and the San
Luis Valley of Colorado.
The banding in Alberta was accomplished by representatives of state, provincial and federal conservation agencies including
Colorado.

�-20-

Banding schedules and recovery reports pertaining to the San Luis Valley
and north central Colorado were completed and submitted to the Bird Banding
Laboratory.
Record keeping for the Alberta operation was the responsibility
of the Alberta Fish and Wildlife Division.

RESULTS

AND DISCUSSION

Hi-Line

Population

Foreign Recoveries
During the 1975-76 hunting season 210 Canada geese banded outside of Colorado
were reported recovered in north central Colorado (Table 1). One hundred ten
of the foreign recoveries were classified as wild-trapped locals when banded
(Table 2).
In Alberta direct recoveries from the second year of banding in southern
Alberta indicated Cavan Lake 14 miles S.E. of Medicine Hat as a production
area for Hi-Line Canada geese.
All other areas on which banding has occurred
east of 1110 longitude in southern Alberta were represented by recoveries in
north central Colorado.
Those areas are Schuler, Many Islands, Cavan, Seven
Persons and Murray lakes. According to all recoveries birds from these areas
are oriented toward the Central Flyway in general and Hi-Line Population specifically (Weaver, unpublished data).
In addition, recoveries were reported
from Grassy Lake (direct), Bantry #2 (indirect), Scope (indirect) and C.P.R.
(indirect); all located west of 1110 longitude.
All but Bantry #2, which
seems to be a transition area, contribute birds primarily to the Pacific
Flyway.
Dowling Lake and the Edmonton-Camrose
area also have been represented
by recoveries in north central Colorado in previous years.
The Cypress Hills area in Saskatchewan and Phillips County in north central
Montana continue to contribute birds to Colorado's Hi-Line harvest area.
Recoveries of wild-trapped locals from Prairie, Treasure and Rosebud counties
in Montana present further evidence that the birds from the Big Horn and
Yellowstone rivers in Montana are members of the Hi-Line. Population.
Birds from newly developed production areas in northeast Wyoming and from
Glendo Reservoir on the Platte River between Casper and Torrington also
were represented in the north central Colorado harvest.
No other production areas outside Colorado were well represented
birds in the north central Colorado harvest.

by banded

Two moulting areas, the Liverpool Bay area in the Northwest Territories of
Canada and Wheatland Reservoir in southeast Wyoming, were represented by
banded birds in the harvest.
Although the birds from the Liverpool Bay
area were wearing "size 8" bands, it is suspected that they were actually
small geese of the shortgrass prairie population and not large moulters from
Hi-Line production areas.

�-21-

Banding continued for the third year on production areas in north central
Colorado.
A total of 825 birds were banded bringing the combined total to
2,493 (Table 3). Geese were captured at 15 locations with 56.7 percent of
the birds banded being goslings (Table 4).

San Luis Valley Population
For the first time in three years the pre-season Canada goose banding
operation in the San Luis Valley was a success with the number of birds
banded totaling 204 (Table 5). Birds were captured at 6 different locations
(Table 6). About 73 percent of the geese banded were goslings.

Prepared by

~rYh-c;,e~-&lt;jP ~ctJ.
Michael R. Szymczak
Wildlife Researcher

Table 1. Banding areas outside Colorado of Canada geese
recovered in north central Colorado during the 1975-76
hunting season.

Area

Direct

Indirect

Alberta
Edmonton-Camrose
Area
Dowling Lake Area
Brooks Area
Bantry 112
Scope Reservoir
Grassy Lake
C. P.R. Reservoir
Murray Lake
Seven Persons Lake
Cavan Lake
Many Islands Lake
Schuler Lake

3
3
0
0
0
1
0
7
5
1
1
1

4
12
2
1
2
3
1
2
0
1
6
0

Saskatchewan
Cypress Hills (Maple Creek)
Cypress Hills Area
Pontiex-Notukeu R.

0
0
0

1
10
2

�-22-

Table 1. Banding areas outside Colorado of Canada geese
recovered in north central Colorado during the 1975-76
hunting season. (continued).

Area

Direct

Saskatchewan (cont'd.)
Vidora Area
Pelican Lake
Buffalo Pound Lake
Battleford Area
Manito Lake Area
Mackenzie - N.W. Territory
Liverpool Bay Area
Smoke River Delta
Harrowby Bay
Central Flyway
Montana
Lake County
Phillips County
Treasure County
Garfield County
Roosevelt County
Valley County
Rosebud County
Prairie County
Custer County

o

Indirect

3

o
o

1
1

4

o

1

2

4
4

o

0
19
3
0
1
0

1
28
8
2

o

1

0
3
0
0
0

North Dakota
Adams County
Dunn County
Williams County

0
0
0

1
1
1

South Dakota
Bennett County
Stanley County

0
0

1

Wyoming
Platte County
Albany County
Goshen County
Crook County
Weston County
Big Horn County
Sheridan County

2
2

2

8

1

7

8

2

0

0
0
0

5

2

4
1
0

--------------------------------------------------------------

�-23-

Table 1. Banding areas outside Colorado of Canada geese
recovered in north central Colorado during the 1975-76
hunting season (continued).

Direct

Indirect

0

1

0

1

0

1

0

1

Utah
Salt Lake County
Rich County

0
0

1
1

Oregon
Harney

0

1

82

128

Area
Central Flyway (cont'd.)
Nebraska
Garden County
Kansas
Phillips

County

MississiEEi Flyway
Missouri
Chariton County,
Pacific Flyway
Washington
Pacific County

Total

l
r

County

Swan Lake

�-24-

Table 2. Banding locations of Canada geese classified as wild-trapped
locals which were recovered in north central Colorado during the 1975-76
hunting season.

Area

1967

1968

Alberta
Edmonton-Camrose Area
Dowling Lake Area
Brooks Area
Bantry 112
Scope Reservoir
Grassy Lake
C.P.R.
Murray Lake
Seven Persons Lake
Cavan Lake
Many Islands

Wyoming
Weston County
Crook County
Platte County

Year of Bandin~
1970 1971 1972 1973

2
2
1

1

1

1974

1975

2
8

2
4

1
2
1
1
1
4

Saskatchewan
Cypress Hills Area
Pontiex-Notukeu R.
Vidora Area
Buffalo Pound Lake
Pelican Lake
Manito Lake
Central Flyway
Montana
Lake County
Phillips County
Treasure County
Valley County
Roosevelt County
Prairie County
Rosebud County

1969

7
5
1
1

9
2
3

1
1

,,

2

I

1
3
1

7
1

2
1

5
2
1

3

1
2
2
3
2
1

Nebraska
Garden County

1

North Dakota
Williams County
Dunn County
Adams County

1
1
1

-----------------------------------------------------------------------------

�-25-

Table 2. Banding locations of Canada geese classified as wild-trapped
locals which were recovered in north central Colorado during the 1975-76
hunting season (continued).

1967

1968

1969

Year of Banding
1970 1971 1972

1973

1974

1975

Area
Pacific Flyway
Utah
Salt Lake County
Rich County

1
1

Oregon
Harney County

1

Table 3. Numbers of-Canada geese banded pre-season on production
areas in north central Colorado, 1974-1976.

Year

Adults

Goslings

Total

1974

751

315

1,067

1975

406

195

601

1976

357

468

825

Table 4. Location and number, by age and sex, of Canada geese banded
in north central Colorado, pre-season 1976.

Location
Spitzer Lake
Rocky Ridge Res.
Watson Lake
Sterling Ponds
Dean Acres Lake
Cobb Lake

Adult§
Females
Males
5
2
1
9
3

15

5
7
1
5
2
20

Goslings
Females
Males
12
20
10
8
5

28

Total

13
20
7
12
12
23

-------------------------------------------------------------------------

35
49
19
34
22
86

�-26-

Table 4. Location and number, by age and sex, of Canada geese banded
in north central Colorado, pre-season 1976 (continued).

Location

Males

Adults
Females

Goslings
Males
Females

Total

Dixon Res.
Anderson's Pond
Timnath Res.
McNeil Res.
Welch Res.
Faivre Ponds
Sawhill Ponds
Valmont Res.
Henderson
(Denver Hatchery)

0

0

11
14
35
11
6
25
30

12
25
51
6
8
19
29

11
44
0
4
36
11
25
23

2
26
10
24
28

26
94
39
92
79
35
93
110

0

0

5

7

12

Total

167

190

242

226

825

15
27
0

Table 5. Numbers of Canada geese banded pre-season on production and
moulting areas in the San Luis Valley, 1974-1976.

�-27-

Table 6. Location and number, by age and sex, of Canada geese banded
in the San Luis Valley of Colorado, pre-season 1976.

Location

Males

Adults
Females

Goslings
Females
Males

-Total

10

10

39

24

83

Head Lake

2

2

0

0

4

Alamosa N.W.R.

3

3

10

5

21

Rio Grande Mgt.
Area

5

7

14

16

42

Monte Vista N.W.R.

4

2

15

7

28

Sheep Pond

4

3

8

11

26

Total

28

27

86

63

204

Russell

Lakes

��-29October

JOB PROGRESS

P~PORT

State of

COLORADO
--------~~~==~-----------

Project

No.

Work Plan No.
Job Title
Period Covered:
Personnel:

Migratory

W-88-R-22

1977

Bird Investigations

2
Job No. ----~~~~--------------------9
Monitor Banding of the Shortgrass Prairie
Canada Goose Population in Southeastern Colorado
January 23, 1977 to February

Cathy Craig, Jennifer
R. Hopper, D. Madsen,
and M. Szymczak.

25, 1977

Slater, J. Carsella, D. Coven, G. Eyre,
S. Steinert, J. Stevenson, K. Wagner,

ABSTRACT

The trapping and banding operation of Canada geese (Branta canadensis) in
southeast Colorado resulted in bands being placed on 354 small Canada geese
for the purpose of monitoring population status.

��-31MONITOR BANDING OF SHORTGRASS PRAIRIE
CANADA GOOSE POPULATION IN SOUTHEASTERN COLORADO
Michael R. Szymczak
This job was initiated in April 1976 as a continuation of a trapping and
banding operation being carried out under the former Work Plan 1, Job 2,
Trapping and Banding Ducks and Geese. The job involves annual post season
banding of Canada geese in southeast Colorado, primarily by personnel of
the southeast region, and periodic analysis of resulting recovery data by
research personnel. The banding operation in 1977, as in the past few years
was under the direction of Kenneth Wagner of the southea t region.

g

P.N.O. OBJECTIVE
To continually document, through monitor banding and analysis of recovery
data, the annual and long-term status of the southeastern Colorado (Arkansas
Valley) segment of the shortgrass prairie Canada goose population.
SEGMENT OBJECTIVES
1.

Band a m1n1mum of 1,000 Canada geese in southeastern Colorado during the
post-season period.

2.

Prepare and submit banding schedules, band recovery and return reports,
and progress reports.
METHODS AND MATERIALS

All birds banded in southeast Colorado in 1977 were captured with baited
cannon nets during the period from January 26 through February 25. With few
exceptions, the age and sex of each bird was determined through cloacal and
tail feather examination.
All banding schedules, including recapture information, were submitted to
the U. S. Fish and Wildlife Service's Bird Banding Laboratory, Patuxent,
Maryland.
RESULTS AND DISCUSSION
A total of 354 small Canada geese were banded post-season in southeast
Colorado. All trapping attempts occurred at Two Buttes Reservoir. The age
and sex composition of the birds banded was as follows: 113 adult males, 124
adult females, 47 immature ma1es, 67 immature. females, 2 unknown.
Extreme difficulty in trapping was encountered because of low water levels
at the primary trap site, Two Buttes Reservoir. In addition, the majority of
the geese in the area were roosting on Turk's Pond and permission to trap
there could not be obtained.

��-33-

October

JOB PROGRESS

State of

REPORT

COLORADO
------~~~~~--------Migratory

W-88-R-22

Proj ect No.
Work Plan No.

3

Job No.

Job Title Popul'ation Characteristics
Period

1977

Covered:

Personnel:

Bird Investigations
7

---------------------------------

of Mallards

Wintering

in West Central

Colorado

April 1, 1976 to March 31, 1977

H. Cox, J. Ellenberger, J. Frothingham, J. Gray, J. Gumber,
J. Leslie, J. Miller, R. Oakleaf, D. Owens, S. Steinert,
M. Stone, H. Wilson, and R. Hopper

ABSTRACT

The study of the wintering population of mallards in west central Colorado
continued during segment 22 for the fourth consecutive year and is progressing
on schedule.
Procedures in segment 22 included: (1) conducting a mid-winter
inventory to obtain an estimate of population size, (2) making ground counts
to estimate the sex ratio in the population, and (3) banding adequate samples
of birds after the hunting season to provide a basis for estimating other
population parameters.
The 1977 January inventory, conducted by Northwest
Regional personnel, yielded an estimate of 16,280 birds on the west central
Colorado wintering grounds at that time. This figure is below the 1976
estimate (23,125), but above the 19-year average (13,474).
A total of
2,654 birds were sexed by ground counts in January and February of 1977,
producing a sex ratio of 107.5 males:lOO females.
This compares to a ratio
of 99.2 males:lOO females for almost 3,500 birds counted in January of 1976.
Postseason trapping in January and February of 1977 resulted in the banding
of 1,498 birds, with a generally good distribution of the sample by location
and age and sex class. This brings the total banded sample to 5,956 for the
four years of effort.

��-35-

POPULATION CHARACTERISTICS OF
MALLARDS WINTERING IN WEST CENTRAL COLORADO
Richard M. Hopper

This report presents results of the fourth consecutive year of study of the
population of mallards wintering in west central Colorado.
In addition,
this report summarizes all previous data collected since initiation of
this investigation four years ago. Past results of this study were reported
annually as part of the job concerning the overall waterfowl banding program
(Work Plan 1, Job 2 - Trapping and Banding Ducks and Geese) (Hopper 1974,
1975,1976).
Work Plan 1, Job 2 was dropped at the end of segment 21 and
specific banding studies were assigned as separate jobs with the beginning
of segment 22, including Work Plan 3, Job 7. As a result, all phases of
a particular population study will now be covered under one job, including
the actual banding work and analysis of band recovery data.
P.N.O. OBJECTIVE
1.

To estimate population parameters of mallards wintering in west central
Colorado by age and sex category; specifically (1) population size,
(2) recovery rates, (3) survival rates, (4) mean life span, (5) sex
ratio, and (6) geographic distribution of the harvest.

2.

To develop a management plan for the population
in west central Colorado.

of mallards

wintering

SEGMENT OBJECTIVES
1.

Conduct an aerial count of wintering numbers of mallards during the
first and second week of January as part of the annual mid-winter survey
conducted throughout the United States.

2.

Conduct a minimum of two ground counts (sex ratio) of 500 mallards each
in each of the following two concentration areas: (1). Grand Junction Highline Lake area, and (2) Montrose - Delta area.

3.

Trap and band 1,500 mallards during the postseason period, including 750
in the Grand Junction - Highline Lake area, and 750 in the Montrose Delta area. Distribute the sample equally among the four age and sex
classes.

4.

Preparation and submission of banding
reports, and progress reports.
METHODS

schedules,

band recovery

and return

AND MATERIALS

An aerial count was made in the Uncompahgre - Gunnison - Colorado River Complex
in west central Colorado on January 10-11, 1977. This was simply a continuation
of the annual count made in major waterfowl concentration areas in Colorado for
many years as the State's contribution to the coordinated, nationwide survey
each year.

�-36-

Personnel in the Northwest Region conducted the aerial count as they have
done in past years.
They utilized a Cessna 185 aircraft and a crew consisting
of a pilot and two observers.
Sex ratio counts were made from the ground in the two general mallard concentration areas, the Grand Junction - Highline Lake area and the Montrose _
Delta area. Specifically, this year's counts were made at either Highline
Lake near Lorna or at Sweitzer Lake near Delta.
These counts were conducted
in as unbiased a manner as possible.
All birds in a group were counted, with
care taken not to concentrate on the more colorful drakes of the species.
The counts were made with the aid of binoculars and spotting scope.
Trapping methods involved the use of baited, Salt Plains type traps and
cannon net traps. Whole and cracked corn were used to attract the birds
to the traps. All birds were aged by the wing-aging technique (Carney and
Geis 1960, Carney 1964, Hopper and Funk 1970). Recapture information was
maintained for each banding location.
Records were kept of the number of
mallards released unbanded by age and sex class. Banding schedules and
recovery reports were prepared and submitted to the Bird Banding Laboratory.
RESULTS AND DISCUSSION
Aerial Census
Results of the 1977 January count of wintering mallards are shown in Table 1
by general area. Also, the previous three years' counts are included for
comparison.
The 1977 estimate was 16,280 birds, substantially less than the
1976 figure (23,125), but above the 1974 and 1975 estimates.
The long-term
(1959-1977) average annual estimated size of this population is 13,474.

Table 1. January inventory of mallards wintering
1974-1977.

Location

in west central Colorado,

1974

1975

1976

1977

Highline

Lake

2,400

4,700

10,200

6,900

Colorado

River

3,110

1,830

1,545

3,390

Gunnison

RO1ver-1/

700

1,345

9,100

2,215

0

485

270

225

0

0

1,900

1,800

2,975

3,400

110

1,750

9,185

11,760

23,125

16,280

Uncompahgre

River

Sweitzer Lake
Burlingame
Total

1/

Pond

Includes North Fork of Gunnison

River to Hotchkiss.

�-37-

The areas covered in this inventory include: (1) High1ine Lake northwest
of Grand Junction, (2) the Colorado River from Silt to the Utah state line,
(3) the Gunnison River from the Black Canyon to its confluence with the
Colorado River near Grand Junction, (4) the North Fork of the Gunnison
River from Hotchkiss to its confluence with the Gunnison River, (5) the
Uncompahgre River from Montrose to Delta, (6) Sweitzer Lake southeast of
Delta, and (7) Burlingame Pond south of Delta.
The importance of each of
these areas as wintering locations can vary from year to year, as can
readily be determined by referring to Table 1. Figure 1 shows a map of the
general area covered by the aerial survey, which also fairly well delineates
the boundaries of the overall study area.
Sex Ratio Counts
Winter sex ratio counts of mallards were initiated in west central Colorado
in January of 1976. They were again conducted in January and February of
1977. The results of these counts are presented in Table 2 by date and
location.
The counts in 1976 involved the sexing of nearly 3,500 birds
and yielded an evenly balanced sex ratio (99.2 ma1es:100 females).
In
1977, 2,654 birds were counted, producing a sex ratio slightly in favor of
males (107.5 ma1es:lOO females).
It is interesting to compare these preliminary results with those obtained from eastern Colorado wintering populations of mallards.
In the latter area, over 73,000 birds were counted during
a 9-year period (1963-64 through 1971-72), resulting in an average sex ratio
of 147 males:lOO females.
Trapping

and Banding

Trapping efforts in January and February of 1977 resulted in the banding
of 1,498 mallards in west central Colorado (Table 3). This was the fourth
consecutive year of postseason banding in this area, bringing the total
banded sample for the four years (1974-1977) to 5,956 birds.
The distribution
of the banded sample between the two general areas (Grand Junction and Delta)
remains fairly good.
Subadult birds of both sexes were well represented in the banded sample in
1977, just as they were each of the previous three years (Table 3). Overall,
the adult female has contributed the fewest number of birds for banding;
either because this age and sex class is not as well represented in the
population as the other three classes, or because adult females are less
vulnerable to capture by the techniques we employ.

�-38-

Table 2. Sex ratio counts of mallards in west central Colorado during
the winters of 1976 and 1977.

Location

Date

No.
Males

No.
Females

Total

Males:
100 Females

Burlingame Pond

1-16-76

244

282

526

86.5

Sweitzer Lake

1-16-76

264

243

507

108.6

1-17-76

239

237

476

100.8

1-20-76

252

249

501

101.2

1-23-76

238

236

474

100.8

Subtotal (Sweitzer Lake)

993

965

1,958

102.9

1-17-76

252

248

500

101.6

1-18-76

252

260

512

96.9

Subtotal (Highline Lake)

504

508

1,012

99.2

1,741

1,755

3,496

99.2

High1ine Lake

TOTAL (1976)
Sweitzer Lake

1-26-77

270

235

505

114.9

High1ine Lake

1-27-77

262

246

508

106.5

1-28-77

288

278

566

103.6

1-31-77

282

266

548

106.0

2-1-77

273

254

527

51.8

Subtotal (High1ine Lake)

1,105

1,044

2,149

105.8

1,375

1,279

2,654

107.5

TOTAL (1977)

�-39-

Table 3. Number and age and sex composition of mallards banded postseason
in west central Colorado, 1977 and totals for the period 1974-1977.

AM

Age and Sex
SM
AF

SF

Total

253

167

179

182

781

Delta Area

165

313

62

177

717

Total

418

480

241

359

1,498

799

636

549

801

2,78?

Delta Area

713

1,024

528

906

3,171

Total

1,512

1,660

1,077

1,707

5,956

Year and Location

1977
Grand Junction

Area

1974-1977
Grand Junction

Area

LITERATURE

CITED

Carney, S. M., and A. D. Geis.
1960. Mallard age and sex determination
from wings.
J. Wildl. Manage. 24(4):372-381.
1964. Preliminary keys to waterfowl age and sex identification by
means of wing plumage.
U.S. Fish and Wildl. Serv., Spec. Sci. Rept.,
Wildl. No. 82. 47 pp.
Hopper, R. M., and H. D. Funk.
1970. Reliability of the mallard wing
age-determination
technique for field use. J. Wildl. Manage. 34(2):
333-339.
1974. Trapping and banding ducks and geese.
Fed. Aid Game Res. Rept., Oct. pp. 13-22.

Colo. Div. of Wildl.,

1975. Trapping and banding ducks and geese.
Fed. Aid Game Res. Rept. , Oct. pp. 13-23.

Colo. Div. of Wildl.,

1976. Trapping and banding ducks and geese.
Fed. Aid Game Res. Rept. , Oct. pp. 13-24.

Colo. Div. of Wildl .•

Prepared

by

~'k.

df',$",ee

Richard M. Hopper
Wildlife Researcher

C

��October

-41JOB PROGRESS

State of
Project

REPORT

COLORADO
--------~~~~~-----------

Job Title

3

Monitor Banding

Covered:

Personnel:

Migratory

W-88-R-22

No.

Work Plan No.

Period

1977

Job No.

Bird Investigations
8

of Eastern Colorado Wintering

_
Mallard

Populations

April 1, 1976 to March 31, 1977

M. Babler, G. Brown, L. Budde, J. Carsella, D. Coven, J. Corey,
C. Craig,C.
Crawford, M. DePra, K. Dillinger, M. Gardner,
C. Grand Pre, R. Hancock, J. Jackson, B. Leasure, C. Like, R. Lopez,
G. Lorentzson, T. Lynch, F. Marcoux, D. Masden, R. Oehlkers,
C. Pabst, J. Pogorelz, F. Rinella, C. Roberts, W. Rupke, J. Slater,
H. Spear, S. Steinert, J. Sweeting, K. Wagner, T. Washington,
R. Zaccagnini, R. Hopper.

ABSTRACT
Segment 22 was the first year for the gradual transfer of the field operations
of the monitor banding of eastern Colorado wintering mallard populations
from the Research to the Management section.
Project personnel assisted
management personnel through orientation, instruction, and field training,
which included the preparation and issuance of an instruction manual to each
crew leader.
The first year of this transfer was successful, but additional
training in record keeping and the wing-aging technique are needed.
Nearly
5,500 mallards were banded during the January-February period, about 1,500
birds over the minimum established quota of 4,000 birds~
The age and sex
composition of the banded sample remained about the same as in previous
years.

��-43-

MONITOR BANDING OF EASTERN
COLORADO WINTERING MALLARD POPULATIONS
Richard M. Hopper

This report summarizes results of the postseason mallard banding program
in eastern Colorado during Segment 22 and describes the orderly transfer of
this monitoring activity from a research to a management function.
This
is the first year of a new job recommended as an ongoing program to provide
periodic population information upon which to base sound management proposals.
Banding relative to the populations of mallards in eastern Colorado began
an intensive research study during the winter of 1963-64 and developed into
the "Investigation of Mallard Management Units of Eastern Colorado"
(Work
Plan 3, Job 6) under Federal Aid Project W-88-R.
This research was initiated
to gain knowledge regarding the population characteristics of these wintering
birds.
The information obtained, plus that from other states, was responsible
for the establishment of the current High Plains Mallard Management Unit of
the Central Flyway (Funk et al. 1971).
Since this accomplishment,
the job
developed into almost strictly a monitor banding program of a management
nature.
Thus, it was decided to terminate Work Plan 3, Job 6 and transfer
the actual banding work to the Northeast and Southeast Regions as a management function.
Work Plan 3, Job 8 was then initiated simply to cover project
personnel's time in (1) instructing management personnel in the field conduct
of the banding program, (2) issuing bands and preparing and submitting banding
schedules and recovery reports, and (3) conducting periodic, updated analyses
of band recovery data resulting from this program.
The final job report for Work Plan 3, Job 6 will appear as one or more publications prepared under Work Plan 6, Job 1. A number of published and unpublished
papers have already resulted from Work Plan 3, Job 6, and one other paper is
currently being reviewed for publication by "The Journal of Wildlife Management".
A list of these papers, excluding the latter one, is presented by Hopper (1976).
P.N.O. OBJECTIVE
1.

To establish monitor banding of wintering mallard
Colorado as an annual management function.

populations

in eastern

2.

To continually document, through monitor banding and analysis of recovery
data, the annual and long-term status of eastern Colorado wintering
mallards to provide a basis for annual hunting season recommendations.
SEGMENT OBJECTIVES

1.

Instruct management personnel in the field conduct of a postseason mallard
banding program by (a) preparing an instructions manual covering all phases
of the operation, (b) holding wing-aging classes, and (c) assisting and
training management personnel in the field.

�-44-

2.

Band a mlnlmum of 4,000 mallards during the postseason period, including
a minimum of 500 birds in each of the following general areas of the
South Platte Valley and Arkansas Valley:
(1) Denver-Greeley,
(2) Fort
Collins-Loveland-Windsor,
(3) Greeley-Fort Morgan, (4) Fort MorganSterling, (5) Sterling-Julesburg,
(6) Bonny Reservoir, (7) Manzanola-Lamar,
and (8) Two Buttes Reservoir area. Divide the banded sample in each area
equally among the four age and sex classes.

3.

Prepare and submit banding
progress reports.

schedules,

METHODS

band recovery and return, and

AND MATERIALS

The Regional Managers and Area Supervisors selected individuals in the
regions to participate in the banding program.
These individuals were assigned
to crews, with each crew consisting of four members, including an alternate.
Each crew was also assigned specific banding areas in which to operate.
All
crew members met in Fort Collins prior to the banding season for orientation,
instructions and preliminary training in aging techniques.
An instructions
manual was prepared and distributed at this meeting.
Two persons from Project
W-88-R spent nearly full time immediately prior to and during the January
and February banding period in instructing management personnel, both in
the office and in the field.
Salt Plains type traps were employed to capture mallards on the banding site,
with whole corn used as bait (Szymczak and Corey 1976). Birds were classified
by sex into either the adult or subadult age categories according to the
wing-aging technique (Carney and Geis 1960, Carney 1964, Hopper and Funk 1970).
Recapture information was maintained at each banding location.
RESULTS AND DISCUSSION
Instruction

of Management

Personnel

Eight banding crews made up of management personnel were formed as a result
of assignments by Regional Managers and Area Supervisors, .seven in the
Northeast Region and two in the Southeast Region.
This number was a few more
than necessary or desirable in the Northeast Region, but it seemed as though
everyone wanted to become involved the first year. Hopefully, this is an
indication of the management personnel's interest in and support for the
postseason mallard banding program.
One crew was assigned to each of the following areas or units:
1, 2, 3, 4,
11, 12-13 (Fig. 1). Two crews split the banding responsibilities in unit 6.
Banding in unit 9 (Bonny Reservoir) will remain the responsibility of the
Research Section because of more intensive banding studies currently being
conducted in association with the population.
A crew leader and an assistant crew leader were designated for each crew.
The crew leader was responsible for the bands and trapping equipment issued
to the crew and for the preparation and submission of field records.
Each
crew leader received an instructions manual outlining all procedures necessary

�WATERFOWL

MANAGEMENT

UNITS

,

WYOMING

-------'
"L":"
, ~?--&gt;;;.
'.=""=-"~-----jl·
'A"~ "\,4 .' .·.;·
J '1

7·

t'\,-;:;'

I,~

WI \ .. "

-rrsrr--r
'. i -;

N'

R

ASK

A

l

---~.-Ii

-.-

'~"r

•••.••
~~

NEB

\,

,,:~,7.~~
.~~:ih,' t!..
I \'
.",.&gt;",~c!".1+-cT
j.'t\,(
, .. , ~):.\,:!d".;!-

",'

+~

N

I

"

OKLAHOMA

GFP-R-M-l

Fig.

1.

in

�-46-

in conducting the field operation.
This manual, contained
binder for field use, included the following information:

in a three-ring

1.

Background information--need
as expected benefits.

for and obj ectives of the program,

2.

General
banding

"do's" and "don'ts"
program.

in conducting

3.

Banding

quotas by area and age and sex class.

4.

Issuance

of bands.

5.

Proper procedures for recording data.
a. Sequence of band usage.
b. Example of properly completed "Field Banding Form".
c. Replacement of old bands on recaptured birds.
d. Example of properly completed "Recapture Field Form".

6.

List of all past banding

7.

Age determination.
a.
b.
c.

locations

the field portion

as well

of any

by area.

Detailed description and key of wing-aging technique from Carney
Diagram of wing showing location and names of feathers.
Reprint of field reliability of the mallard wing-aging technique
during the postseason banding period by Hopper and Funk (1970).

8.

Booklet

by Szymczak

9.

Submitting

and Corey

(1976) describing

(1964).

use of the Salt Plains

trap.

field records and unused bands.

The first year in the transformation of banding duties from the research to
the management section was considered a success.
Management personnel were
cooperative and showed an active interest in the program.
With few exceptions,
their field records were legible and completed in the proper manner.
A number
of individuals need additional training and experience with the wing-aging
technique, and this should come as the more dedicated and interested personnel
continue their participation in the next few years.
Banding
The total banded sample amounted to 5,486 mallards for the January-February
period of Segment 22 (Table 1). This exceeded the 4,000 minimum quota by
almost 1,500 birds.
However, most of this excess was provided by the Bonny
Reservoir location where a more intensive banding study is being conducted
by Project W-88-R.
All eight banding areas contributed good samples, except
for the Manzanola-Lamar
area in the Arkansas Valley.
The age and sex composition of the banded sample was typical of most years,
with adult males being the most represented, while adult females contributed
least to the sample (Table 1). There was also the usual amount of fluctuation
in the age and sex composition among the various banding areas.

�-47Table 1. Mallards banded postseason by age and sex in the eight eastern
Colorado banding areas, January-February 1977.

Banding
Area

AM

Number of Ducks Banded
Age and Sex
SF
AF
SM

Bonny Reservoir

528

498

203

315

1,544

Sterling-Julesburg

152

210

127

158

647

Fort Morgan-Sterling

150

196

125

179

650

Greeley-Fort

184

194

81

126

585

Denver-Greeley

156

140

159

l35

590

Ft. Collins-Loveland-Windsor

212

136

155

124

627

Manzanola-Lamar

12

6

3

10

31

Two Buttes Area

242

122

198

250

812

1,636

1,502

1,051

1,297

5,486

Literature

Cited

Total

Morgan

Carney, S. M., and A. D. Geis.
wings.
J. Wildl. Manage.

Total

1960. Mallard age and sex determination
24(4) :372-381.

from

1964. Preliminary keys to waterfowl age and sex identification by
means of wing plumage.
U. S. Fish and Wildl. Serv., Spec. Sci. Rept.,
Wildl. No. 82. 47 p.
Funk, H. D., J. R. Greib, D. Witt, G. F. Wrakestraw, G. W. Merrill, J. Sands,
T. Kuck, D. Timm, T. Logan, and C. D. Stutzenbaker .. 1971. Justification of the Central Flyway High Plains Mallard Management Unit.
Unpubl. Central Flyway Tech. Comm. Rept.
48 p.
Hopper, R. M., and H. D. Funk.
1970. Reliability of the mallard wing agedetermination technique for field use. J. Wildl. Manage. 34(2):333-339.
1976. Investigation of mallard management units of eastern Colorado.
Colo. Div. of Wi1dl. Fed. Aid Game Res. Rept., Oct. pp. 87-91.
Szymczak, M. R., and J. F. Corey.
Plains duck trap in Colorado.
l3 p.

Prepared

1976. Construction and use of the Salt
Colo. Div. of Wildl., Div. Rept. No.6.

by _""-~~~~-==-.:::::.:
..",*=---~-f---=+L:.f'HfUJ&amp;t~~--Richard M. H~
Wildlife Researcher

��-49-

JOB PROGRESS

State of

COLORADO
------------------------------

Project

No.

W-88-R-22

Migratory
3

Bird Investigations

----------------------------

---------------=~~~~~~~~~~~~-------------

Period Covered:
Personnel:

REPORT

9
Job No.
Migration ani Mortality Characteristics. of Duck Populations
in the Inter-mountain Valleys of Colorado

Work Plan No.
Job Title

October 1977

July 15, 1976 to March 31, 1977

Pete Bryant and staff, Monte Vista National Wildlife Refuge;
M. Conner, J. Corey, W. Dolezal, D. Flenthrope, R. Hopper,
J. Lorentzson, D. Nelson, R. Oakleaf, S. Porter, J. Wagner,
K. Wagner, R~ Velarde and M. Szymczak, Colorado Division of
Wildlife.
ABSTRACT

Totals of 3,285, 3,624 and 4,979 ducks were banded in North Park, South Park
and the San Luis Valley, respectively, in 1976. A total of 374 duck wings
~ere collected during the early duck season in North Park of which 28.1
percent were gadwall, 21.4 percent wigeon and 11.2 percent redheads.
The
species composition of the harvest in North Park in 1976 was similar to what
was found in 1975. As the season progressed in North Park, a definite
decline was noted in the percent of the harvest composed of blue-winged
teal. Of the 92 wings collected at Antero Reservoir in South Park, 28.3
percent were wigeon, 25.0 percent were green-winged teal and 23.9 percent
were blue-winged or cinnamon teal.

��-51-

MIGRATION AND MORTALITY CHARACTERISTICS
OF DUCK POPULATIONS IN THE INTER-MOUNTAIN
VALLEYS OF COLORADO
Michael

R. Szymczak

This job was initiated in order to combine all facets of this investigation
under one program.
In previous years the banding operations were carried
out under the former Work Plan 1, Job 2, Trapping and Banding Ducks and
Geese, while the recovery analysis was scheduled to be accomplished under
the former Work Plan 1, Job 3, Analysis of Waterfowl Banding Data.
Therefore, even though this job is in the first year, a considerable
amount of information has accumulated in past years.

P.N.O.

OBJECTIVES

1.

To investigate migration, mortality, recovery distribution and
relationships
among populations of selected species of ducks present
in North Park, South Park and the San Luis Valley during the midJuly through mid-September
period.

2.

To document the species composition of ducks harvested during early
October seasons, should seasons occur in North Park, South Park
and the San Luis Valley.

3.

To examine the feasibility of establishing early special duck seasons
in September in the high mountain park areas or to examine the
feasibility of special early October seasons in South and North
Park similar to what has been recommended for the San Luis Valley.

4.

To establish procedures to monitor the effect of early special duck
seasons on duck populations in the high mountain park areas if granted.

SEGMENT

OBJECTIVES

1.

Trap and band ducks in North Park, South Park and the San Luis Valley
during the mid-July through mid-September
period as designated in
Program Narrative Outline.
Complete, submit and file appropriate
banding schedules and recovery cards.

2.

Collect and analyze data concerning the species composition of the
harvest during early October seasons in North Park, South Park and
the San Luis Valley utilizing wing collection barrels and results of
the U.S. Fish and Wildlife Service's Parts Collection Survey.

3.

Analyze

band recovery

data through

the 1976 recovery

year for mallards,

�-52-

pintails and green-winged teal banded during the pre-season period
in North Park, South Park and the San Luis Valley to determine
feasibility of requesting early seasons in the high mountain areas.
4.

Prepare

progress

report.

METHODS

AND MATERIALS

Ducks were captured from August 7-September 10 in South Park, August 9September 10 in the San Luis Valley and July 26-September 19 in North
Park.
Primarily "salt plains" type bait traps were used (Szymczak and
Corey 1976).
In North Park night lighting from airboats was used to
capture most gadwall and wigeon.
The age and the sex of all birds captured
and banded were determined.
Wings from ducks bagged during the October 2, 1976 through October 17, 1976
period were collected in North and South Parks through the use of voluntary
collection barrels (Hoffman and Braun 1975).
Barrels were placed at
Walden Reservoir (2 barrels) and Lake John Annex (1 barrel) in North
Park and at Antero Reservoir (2 barrels) in South Park.
Some additional
wings were collected at ~cFarlane
Reservoir.
All wings collected were
classified by species, age, sex and location, and in some cases periods
of harvest.
The species composition of the harvest in the San Luis Valley
was scheduled to be obtained at the Central Flyway, U.S. Fish and Wildlife
Service's Parts Collection "wing bee". However that information wasn't
gathered and now must be requested from the U.S. Fish and Wildlife Service
and thus will not be presented in this report.
Banding and recovery tapes plus recapture and return information covering
all pre-season banding and resulting recoveries beginning in 1961 in the
San Luis Valley, 1968 in South Park and 1954 in North Park through the
1975-76 recovery year were requested from the Bird Banding Laboratory
for mallards, pintails and the three species of teal.
In addition information
concerning all species of ducks recovered or recaptured in the above three
areas but banded elsewhere was also requested.
That information has been
received but has not been analyzed due to problems in ADP program conversions.

RESULTS AND DISCUSSION
Trapping

and Banding

Totals of 3,285, 3,624, and 4,979 ducks were banded in North Park, South
Park and the San Luis Valley, respectively, in 1976. The species composition
of the birds banded in these areas are presented in Tables 1, 2 and 3,
respectively.
Established quotas of 300 mallards of each age and sex were
not met for females in North Park, for any classification in South Park and
for adult males in the S~n Luis Valley.
Green-winged teal quotas of 500 were
not met in North Park while gadwall quotas of 300 of each sex in North Park
were achieved.
In total nearly 12,000 ducks were banded (Table 4).

�-53-

Table 1. Number of ducks banded, by species, in North Park during the
pre-season period, 1976.
Age and Sex
AM

1M

AF

IF

Total

Mallard

316

343

189

298

1,146

Gadwall

493

50

250

54

847

American Wigeon

83

9

19

7

118

Green-winged Teal

124

28

77

47

276

Blue-winged and
Cinnamon Teal

3

10

2

10

25

Pintail

195

245

180

235

855

Redhead

4

4

6

4

18

1,218

689

723

655

3,285

Total

Table 2. Number of ducks banded, by species, in South Park during the
pre-season period, 1976.

AM

Age and Sex
1M
AF

IF

Total

Mallard

187

199

95

170

651

Gadwall

0

3

0

0

3

Green-winged Teal

635

518

155

238

1,546

Blue-winged and
Cinnamon Teal

67

142

29

85

323

Pintail

385

356

101

240

1,082

Redhead

12

0

7

0

19

1,286

1,218

387

733

3,624

Total

�-54-

Table 3. Number of ducks banded, by species, in the San Luis Valley during
the pre-season period, 1976. 11

AM

Age and Sex
1M
AF

IF

Total

Mallard

287

526

379

457

1,649

Gadwall

3

12

3

20

38

American Wigeon

3

3

0

4

10

Green-winged Teal

658

397

117

191

1,363

Blue-winged and
Cinnamon Teal

140

264

57

165

626

Pintail

255

349

254

373

1,231

Redhead

11

19

12

20

62

1,357

1,570

822

1,230

4,979

Total

11 Includes 1,965 ducks banded by the Monte Vista National Wildlife Refuge and
Alamosa National Wildlife Refuge.

Table 4. Number of ducks banded, by species, in North Park, South Park, and
_th~SanL4is _Yal,1eyduring the pre+season perLod 1976.

Mallard
Gadwall
•••

~

•.••

~

.••••••

-&lt;

-.-

AM

Age and Sex
1M
AF

IF

Total

790

1,068

925

3,446

496
••

--.-.--

•.••••

--~-------'"--.--

American Wigeon

•• -,.--------,-'--.--~.-.

663

65
-

•• ~.---'-.~.,

.•

'-.----,

••••

-

•••••

253
' •••

~-

.-,

-

-&lt;

.,•••~-" ••,--.-

••-.-

'.",-",

74
~.- ~'----.'-"-'

,.• ,- .•• "." ""-,,.

"'-'--"--.

888
--"--.~-. -""---~

86

12

19

11

128

Green-winged Teal

1,417

943

349

476

3,185

Blue-winged and
Cinnamon Teal

210

416

88

260

974

Pintail

835

950

535

848

3,168

Redhead

27

23

25

24

99

3,861

3,477

1,932

2,618

Total

.....

11,888 11

II Includes
1,965 ducks banded by Monte Vista N.W.R. and Alamosa N.W.R.

�-55-

Species Composition

of the Harvest

A total of 374 duck wings were collected during the early duck hunting
season in North Park.
Over 70 percent of the wings came from Walden
Reservoir.
Nearly 50 percent of the harvest was comprised of gadwall
(28.1%) and wigeon (21.4%) (Table 5). The redhead (11.2%) was the only
other species which amounted to more than ten percent of the harvest.
The composition varied somewhat by location of harvest.
The species composition of the 1976 early season harvest in North Park
was very similar to the composition in 1975.
In 1975, the percent of
the harvest composed of gadwall and wigeon was the same (49.5), however
the wigeon was the dominant species (Table 6). The only other significant
changes between the two years were the comparative reduction in the harvest
of blue-winged or cinnamon teal and the increase in the redhead.
The composition of the harvest in North Park through the season varied
only slightly according to the two time periods indicated in Table 7.
The only significant change was the reduction in the percent harvest of
blue-winged teal. Plotting the distribution by time period for Walden
and Lake John Annex separately indicated slightly more species variation
by individual areas through time than did the pooled results (Tables 8
and 9).
Ninety-two wings were collected at Antero Reservoir in South Park.
Over
75 percent of the harvest was composed of wigeon, blue-winged teal or
cinnamon teal and green-winged teal in nearly equal proportions (Table 5).
An insignificant number of gadwall were harvested at Antero.
The age and sex composition of the harvest in North Park and South Park
are presented in Tables 10 and 11, respectively.

�Table 5. Species composition of the harvest in North Park and South Park during the October 2-17, 1976
early duck hunting season according to wings voluntarily placed in collection barrels.

Walden Res.

North Park
L. John Annex
MacFarlane Res. 1./

Gadwall

88 (33.3)~./

12 (14.1)

A. Wigeon

60 (22.7)

Redhead

Total

South Park
Antero Res.

5 (20.0)

105 (28.1)

2 (2.2)

13 (15.3)

7 (28.0)

80 (21.4)

26 (28.3)

29 (11.0)

13 (15.3)

o (0.0)

42 (11.2)

5 (5.4)

Blue-winged or
Cinnamon Teal

21 (8.0)

8 (9.4)

4 (16.0)

33 (8.8)

22 (23.9)

Green-winged Teal

15 (7.6)

6 (7.1)

6 (24.0)

27 (7.2)

23 (25.0)

1. Scaup

20 (5.7)

6 (7.1)

Mallard

13 (4.9)

Pintail

26 (7.0)

o (0.0)

10 (11.8)

o (0.0)
o (0.0)

23 (6.1)

8 (8.7)

9 (3.4)

8 (9.4)

1 (4.0)

18 (4.8)

3 (3.3)

N. Shoveler

8 (3.0)

1 (1.2)

1 (4.0)

10 (2.7)

2 (2.2)

Ruddy

1 (0.4)

5 (5.9)

o (0.0)

6 (1.6)

Ring-necked Duck

o (0.0)

3 (3.5)

1 (4.0)

4 (1.1)

o (0.0)
o (0.0)

C. Merganser

o (0.0)

o (0.0)

o (0.0)

o (0.0)

1 (1.1)

85

25

Total

264

1/ Wings collected only on October 2-3, 1976.
~/ Percent of total in parentheses.

374

92

I
U1
(J\

I

�-57-

Table 6. Species composition of the harvest in North Park, Colorado during
early duck hunting seasons in 1975 (October 4-14) compared to 1976 (October
2-17) according to wings voluntarily placed in collection barrels.
1975

1976

Species

No.

Percent

No.

Percent

American Wigeon

143

28.9

80

21.4

Gadwall

102

20.6

105

28.1

Blue-winged or
Cinnamon Teal

76

15.4

33

8.8

Green-winged Teal

38

7.7

27

7.2

Mallard

34

6.9

23

6.1

Pintail

25

5.1

18

4.8

Lesser Scaup

25

5.1

26

7.0

Redhead

21

4.3

42

11.2

N. Shoveler

17

3.4

10

2.7

Ruddy

8

1.6

6

1.6

Ring-necked Duck

1

0.2

4

1.1

Unknown

4

0.8

o

0.0

Total

494

374

�-58-

Table 7. Species compositions of the harvest by time period at Walden
Reservoir and Lake John Annex, North Park, Colorado during the early October
1976 duck hunting season according to wings voluntarily placed in collection
barrels.

October 2-3
No.
Percent

Species

Gadwall

Time Period
October 4-17
No.
Percent

No.

Entire
Percent

74

28.9

26

28.0

100

28.7

50

19.5

23

24.7

73

20.9

Redhead

31

12.1

11

11.8

42

12.0

Blue-winged or
Cinnamon Teal

27

10.5

2

2.2

29

8.3

Lesser Scaup

19

7.4

7

7.5

26

7.4

Mallard

17

6.6

6

6.5

23

6.6

16

6.5

5

5.4

21

6.0

Pintail

10

3.9

7

7.5

17

4.9

N. Shoveler

7

2.7

2

2.6

9

2.6

Ruddy

5

2.0

1

1.1

6

1.7

o

0.0

3

3.2

3

0.9

American

Wigeon

Green-winged

Ring-necked

Total

Teal

Duck

256

93

349

�-59-

Table 8. Species compositions of the harvest by time period at Walden
Reservoir, North Park, Colorado during the early October 1976 duck hunting
season according to wings voluntarily placed in collection barrels.

Time Period
October 2-3
October 4-17
No.
Percent
No.
Percent

No.

Entire
Percent

Gadwall

72

33.3

16

33.3

88

33.3

American Wigeon

43

19.9

17

35.4

60

22.7

Redhead

24

11.1

5

10.4

29

11.0

Blue-winged or
Cinnamon Teal

21

9.7

0

0.0

21

8.0

Lesser Scaup

17

7.9

3

6.3

20

7.6

Green-winged Teal

13

6.0

2

4.2

15

5.7

Mallard

10

4.6

3

6.3

13

4.9

Pintail

8

3.7

1

2.1

9

3.4

N. Shoveler

7

3.2

1

2.1

8

3.0

Ruddy

1

0.5

0

0.0

1

0.4

Total

216

Species

t

~
~

48

264

�-60-

Table 9. Species composition of the harvest by time period at Lake John
Annex, North Park, Colorado during the early October 1976 duck hunting
season according to wings voluntarily placed in collection barrels.

Species

Oct. 2-3
No.
Percent

Time Period
Oct. 11-17
Oct. 4-10
No.
Percent
No.
Percent

American Wigeon

7

17.5

6

16.7

a

0.0

13

15.3

Redhead

7

17.5

6

16.7

a

0.0

13

15.3

Gadwall

2

5.0

7

19.4

3

33.3

12

14.1

Mallard

7

17.5

1

2.8

2

22.2

10

11.8

Blue-winged or
Cinnamon Teal

6

15.0

2

5.6

a

0.0

8

9.4

Pintail

2

5.0

6

16.7

0

0.0

8

9.4

Green-winged Teal

3

7.5

a

0.0

3

33.3

6

7.1

Lesser Scaup

2

5.0

3

8.3

1

11.1

6

7.1

Ruddy

4

10.0

1

2.8

a

0.0

5

5.9

Ring-necked Duck

a

0.0

3

8.3

0

0.0

3

3.5

N. Shoveler

0

0.0

1

2.8

0

0.0

1

1.2

Total

40

36

9

Entire
No. Percent

85

t

~

~

�-61-

Table 10. Age and sex composition of the harvest by species in North Park
at Walden Reservoir, Lake John Annex and MacFarlane Reservoir during the
October 2-17, 1976 early duck hunting season according to wings voluntarily
placed in collection barrels.

Ad.

Males
Inun.

Unk.

Ad.

Gadwall

21

37

0

7

21

American Wigeon

37

13

0

13

Redhead

20

7

0

Blue-winged or
Cinnamon Teal

2

15

Green-winged Teal

1

Lesser Scaup

Species

r

~

Females
Imm.
Unk.

Unknown

Total

0

19

105

15

0

2

80

8

3

0

4

42

I

5

7

1

2

33

6

0

3

9

1

7

27

7

3

0

8

5

0

3

26

Mallard

7

sll

0

5

6

0

0

23

Pintail

6

8

0

I

3

0

0

18

N. Shoveler

6

3

0

1

0

0

0

10

6l:-1

Ruddy

6

~

Ring-necked Duck

I

Jj Includes one local male.
]j

All adult unknowns.

0

0

I

1

0

I

4

�-62-

Table 11. Age and sex composition of the h~rvest by species in South Park
at Anteror Reservoir during the October 2-17, 1976 early duck hunting season
according to wings voluntarily placed in collection barrels.

Species

Prepared

Ad.

by

Males
Imm.

Unk.

J7'h;Ll72~
Michael R. S

Ad.

za
Wildlife Researcher

Females
Imm.
Unk.

Unknown

Total

�-63October

1977

JOB FIN.A.LREPORT

State of

COLORADO
----~~~==~-----

Project

No.

W-88-R-22

Work Plan No.
Job Title
Period

Investigations

Covered:

Personnel:

5

Migratory

Bird Investigations

Job No.

2

of Rails in Colorado

1 May 1975 through 31 March 1977

C. Braun, H.Funk,
and H. Griese, Colorado Division of Wildlife,
R. Ryder, G. Swanson, and R. Walter, Colorado State University,
and C. Bryant, U. S. Fish and Wildlife Service.

ABSTRACT
Distribution, breeding densities, peaks in densities during migrations, and
habitat utilized by rails were investigated in Colorado between May 1975
and October 1976. Five major study areas located in the Arkansas, South
Platte, Rio Grand-e, and Yampa river drainages were censused by playback of
tape recorded rail calls.
Breeding distributions of soras (Porzana carolina)
and Virginia rails (Rallus limicola) were delineated.
Soras were found to
breed up to 3,140 m elevation, while Virginia rails were found to to 2,730 m.
Other species of rails were not found to breed in Colorado.
State-wide,
Virginia rails exhibited higher breeding density indices (2.6 rails/ha)
than soras (1.3 rails/ha).
Indices for Virginia rails ranged from 4.7 in
southeastern Colorado to 1.0 in northwestern Colorado.
Indices for soras
ranged from 2.5 in the south central region to 0.2 in the southeastern region
of Colorado.
Distribution of Typha marshes was not found to determine
distribution of rails. Mean April temperatures, a function of topography,
appeared to regulate species composition in 1975 and 1976. Soras were more
abundant than Virginia rails when mean April temperatures were 5.6oC and
below.
Typha appeared to be the most abundant emergent vegetation in Colorado, it was also the most preferred cover.
Seed producing emergents
attracted soras during fall migration.
Water depths and water level
fluctuations appeared to influence breeding densities.
Clutch initiation
dates (116) calculated from nest searches, literature, and estimated ages
of trapped chicks revealed mean clutch initiation dates of 14 June and 10
June for soras and Virginia rails in Colorado, respectively.
Initiation
dates ranged from 22 April to 4 August.
Peak concentrations of migrating
rails appeared to occur in late April to early May in the spring, and midAugust to mid-September in the fall. Migration did not appear to differ
substantially between plains and inter-mountain regions of Colorado.
Wintering rail populations were estimated not to exceed 200 individuals of
which 75 to 90 percent were Virginia rails.
During this study, 294 rails
(142 soras, 152 Virginia rails) were captured and banded.
Late summer adult
to juvenile ratios for soras and Virginia rails in trap samples were 1:4.4
and 1:3.2, respectively.

�-64-

RECOMMENDATIONS

1.

Marshes specifically managed for breeding rails should contain primarily
(&gt;60 percent) robust vegetation (e.g. Typha) interspersed with small
openings containing short-thin emergent species (e.g. Scirpus americanus)
or mud flats. Water levels should be mainipulated to average 15 cm in
depth during spring migration.
A gradual decrease in water levels through
summer months can be tolerated provided sufficient levels are present in
late summer.
Reflooding in late August should enhance the attractiveness
of marshe~ to rqils.

2.

Periodic breeding censuses, every three to five years, should be conducted.
An average of two censuses on consecutive days at selected marshes in
late May to late June sh.ould produce comparable indices.
Habitat losses
should be monitored regionally.

3.

Ideally, hunting $easons in Colorado should begin no earlier than the
second weekend in September, ending in mid-October.
Consideration should
be given to in~reasing hunter interest in rails, possibly through more
attainable bag and possession limits.
Harvest data should be routinely
collected.

4.

Rail hunting shoU~4 be allowed on the west slope of Colorado.
should $tqrt concurrently state-wide.

5.

Research. should be conducted on: 1) determination of reliable field technique$ for det~rrnin:i,ngsex and age of rails during all seasons; 2)
implicat:i,qns qf lqte $ummer nesting; 3) motivation for migration and
delineatiqn of IllJg1."ati.,c;mal
routes; and 4) estimation of recovery and
survival rates through band:i..ng
programs.

Seasons

�-65-

TABLE OF CONTENTS
Page

........

INTRODUClfION

69

P.N.O. OBJECTIVE

70

LITERATURE REVIEW
Taxonomy
Distribution
Ecology . • •
Census Techniques

70
70
70

71
71

•

METHODS • • • . • . . .
Study area Selection
Census Techniques . • •
Physical and Botanical Dimensions
Water and Substrate Descriptions
Microclimatic Data
Trapping
• . • . .

.........

LOCATION AND DESCRIPTION OF STUDY AREAS
Location
Description . • . • • • . •
Water Chemistry • • • .
Climate and Topography
Microclimates
• . • •
RESULTS AND DISCUSSION
Distribution
Breeding Densities
Habitat Utilization
Spring Migration
Nesting . . • • .
Brood Development
Fall Migration
Timing of Migration . •
Spring
Late Summer Movement
Fall
Trapping
SUMMARY

•

75
75
75
76
76
78

..........
.........

...•

LITERATURE
APPENDICES

72
72
72
74
74
74
74

'. .

78
78
84
91
91
91
93

94
94
94
94
100
100
102

CITED

104

..........

107

�-66-

LIST OF TABLES

Table
1

2

3

4

5

6

7

Results of water analysis at regional study areas
in Colorado, August 1975 and April 1976
....•

77

Initiation dates of clutches of soras and Virginia
rails in Colorado by region as estimated from nest
searches, literature, and estimated ages of trapped
chicks • • . • • • • • • . • . . • . ••
.•.•

81

Chi-square analyses testing hypothesis that
distribution of Typha-dominated marshes
determine distribution of rails in Colorado,
state-wide, by region, and elevation •

82

Indices of breeding densities of rails in four
regions of Colorado, 1975 and 1976 . .

85

Results of testing hypotheses concerning regional
and yearly differences in breeding densities of
rails for Colorado
•....••..•.

86

Breeding densities of rails at five regional study
areas in Colorado in 1976 as determined by spotmapping territories

...........

87

Distribution of bandings
Colorado, 1975-1976

of rails, by species,
. • . • • • . . . . •

101

�-67-

LIST OF FIGURES
Figure
1

2

3

4

Study areas, trapping sites, and wetlands censused
during investigations of rails in Colorado, 19751976
..•••.......••..••.•.
Breeding
township

distribution
in Colorado,

of Virginia rails by
1975 and 1976

73

........

79

......

80

.......

88

......

89

.....

90

.....

92

Breeding distribution of soras by township
in Colorado, 1975 and 1976 ••
•...
Breeding densities of all rails as determined
by spot mapping territories correlated to
averaged spring censuses
= 0.20 + 0.90x;
r = 0.97; P&lt;0.05)
.....••••..••

(Y

5

Breeding densities of soras as determined
by spot-mapping territories correlated to
averaged spring censuses
= 0.16 + 0.95x;
r = 0.97; P&lt;0.05)
•••..•
• •.••

(Y

6

Breeding densities of Virginia rails as determined by spot-mapping territories correlated
to averaged spring censuses
-0.36 + l.llx;
r = 0.93; P&lt;O.lO)
..•••.••....

(Y

7

t
8

9

10

11

12

Relationship between mean April temperatures
(OC) near study areas and percent composition
of the breeding population~, 95 percent confidence intervals depicted (Y = 7.25e-l.96x;
r = -0.94; P&lt;O.Ol) ...••••
Densities of rails responding to taped calls at
John Martin Reservoir, April through
September, 1975 and 1976 •••.
Densities of rails responding to taped calls at
Lower Latham Reservoir, April through
September, 1975 and 1976 ..••

95

96

Densities of rails responding to taped calls at
Monte Vista NWR, April through September 1975 and 1976

97

Densities of rails responding to taped calls at Cary
Ranch, April through September 1975 and 1976 . . . .

98

Densities of rails responding to taped calls at Milk
Creek I and II, April through September 1976 . . . .

99

��-69-

INVESTIGATIONS

OF RAILS IN COLORADO

ACKNOWLEDGMENTS
This project was funded by The Accelerated Research Program for Migratory
Shore and Upland Game Birds, U. S. Fish and Wildlife Service, Contract No.
14-16-008-2000 administered through Colorado Division of Wildlife Federal
Aid in Wildlife Restoration Project W-88-R.
I extend my thanks to these
agencies for their support.
I thank the project coordinator, Dr. C. E. Braun, Colorado Division of
Wildlife, and my academic adviser, Dr. R. A. Ryder, Department of Fishery
and Wildlife Biology, Colorado State University, who were responsible for
the initiation of this project.
Their criticisms and suggestions throughout the term of this project were vital to its successful completion.
I
also thank my graduate committee members, Mr. R. G. Walter, Department of
Botany and Plant Pathology, and Dr. G. A. Swanson, Department of Fishery
and Wildlife Biology, Colorado State University, for their helpful advice.
Thanks go to C. Bryant, Manager, Monte Vista National Wildlife Refuge, U. S.
Fish and Wildlife Service, and the numerous
landowners in Colorado for
allowing me to readily utilize their marshes.
I extend my appreciation
to those Colorado Division of Wildlife personnel who aided my search for
rails.
I especially thank Wildlife Conservation Officers D. Coven, D.
Kenvin, D. Miller, and J. Wagner and the numerous work study students at
Colorado State University who accompanied me into marshes with an exuberance
that was appreciated.
Appreciation is expressed to the Soil
Conservation Service, U. S. Department of Agriculture, for the use of
unpublished data.
My special thanks go to bird watchers in Colorado whose observations
invaluable in completing this project.

were

INTRODUCTION
Five species of rails have been reported in Colorado (Bailey and Niedrach
1965, Colorado Field Ornithologists Records Committee, pers. comm.).
The
two most common rails, Virginia rail and sora, can be hunted east of the
Continental Divide.
Black (Lateral Ius jamaicensus), yellow (Coturnicops
noveboracensis) and king rails (Rallus elegans) are not legally hunted in
Colorado.
There is no open hunting season for rails west of the Continental
Divide (i.e. in the Pacific Flyway).
To meet an increasing necessity for management plans for rails, research
in recent years has been directed at developing techniques for ascertaining
their status.
Tomlinson and Todd (1973) and Glahn (1974) were successful
in developing census techniques for rails.
Subsequently, use of taperecorded calls has proven practical for censusing most species of rails
(Mangold 1974, Baird 1974, Tacha 1975, Holliman 1976).
Limited data exist concerning the status of rails in Colorado.
No more than
museum specimens at state universities and at the Denver Museum of Natural
History and field notes exist for the state.
These field notes and data

�-70-

from museum specimens, primarily from the east slope, were compiled by
Bailey and Niedrach (1965) and Lane and Holt (1975). By counting flushed
or vocal soras, Boeker (1954) was able to approximate territorial pairs
(120) in the Yampa Valley of northwestern Colorado in 1953. Glahn (1974)
used taped recorded calls do delimit distribution of soras and Virginia
rails in selected marshes in north central Colorado, and subsequently, was
able to estimate breeding densities.
P.N.O. OBJECTIVE
The major objective of this study was to obtain data necessary for development of a management plan for rails breeding in and migrating through Colorado. Data considered necessary were those relating to distribution,
breeding densities in at least four different areas, descriptions of habitats
utilized according to vegetation, water depths and other selected physical
characteristics,
and documentation of timing of spring and fall migration.
Hypotheses

tested included:

a.

Distribution
of Typha.

of rails in Colorado

b.

Densities of breeding rails differ significantly by species between
occupied habitats in the eastern plains and the high mountain valleys.

c.

Timing of spring and fall migration differs significantly
areas east of the mountains and inter~mountain valleys.
LITERATURE

is determined

by the distribution

between

REVIEW

Taxonomy
The five species and three subspecies of rails recorded from Colorado are
Rallus elegans Audubon, Porzana carolina (Linnaeus), Rallus!.
limicola
Vieillot, Laterallus i. jamaicensis (Gmelin), and Coturnicops ~.
noveboracensis
(Gmelin).
They represent the sub-family Rallinae, Family
Rallidae, Order Gruiformes, and Class Aves and regularly occur in North
America north of Mexico (AOU 1957).
Distribution
In North America north of Mexico, inland species of rails (i.e. excluding
clapper rails (Rallus longirostrus»
breed from Nova Scotia and central
Canada to southern California east to the Atlantic coast in favorable marsh
habitat.
They are generally migratory, wintering primarily in Atlantic and
Pacific coastal marshes and southern states to northern South America.
Largest concentrations winter in coastal marshes of southern and southeastern
United States and Mexico (AOU 1957).
In Colorado, Virginia rails and soras (hereafter referred to as "rails"
unless otherwise specified) are common breeders.
Virginia rails commonly
nest in suitable habitat in the upper sonoran zone (1,070-1,680 m) and

�-71-

rarely remain throughout the winter.
Soras nest commonaly in marsh habitat
in the plains and occasionally in the mountains to 3,050 m. Black and king
rails (recent records) are rare summer residents.
Yellow rails are rare
migrants.
These three species are not known to breed in Colorado (Bailey
and Niedrach 1965).
Ecology
Life history and behavior of all North American rails have been described by
Bent (1926) and Ripley (1977). Allen (1934), Walkinshaw (1937, 1940),
Pospichal and Marshall (1954), Tanner and Hendrickson (1954, 1956), and
Kaufmann (1971) have described the ecology of rails.
King rails have been
studied by Meanley (1969). Descriptions of foods utilized by various rails
have been presented by Pospichal and Marshall (1954), Webster (1964),
Meanley (1965), and Horak (1970). Habitat utilized has been described by
Billard (1947), Tanner and Hendrickson (1954, 1956), Horak (1970), Andrews
(1973), Baird (1974), and Tacha (1975).
Vocalizations of rails have been described by Bent (1926), Kellogg, Allen,
and Peterson (1959), and Kaufmann (1971). Descending calls of rails are
given throughout the breeding season in response to other rail calls and
loud noises.
The hollow sounding descending call of Virginia rails evokes
responses from other territorial males but not from mates (Kaufmann 1971).
Census Techniques
Early attempts to census rails produced little success.
Boeker (1954) used
flush counts and counted vocalizations to estimate numbers of breeding soras
in the Yampa Valley, Colorado.
Smith (1955) censused soras in Canada by
counting distress calls evoked by tossing stones into potholes.
Bateman
(1965) had erratic success when attempting to flush clapper rails in Louisiana. Meanly (1969) and Post and Enders (1970) found that by mapping
approximate locations of vocalizations (spot-mapping), locations and number
of breeding territories could be approximated for king and Virginia rails,
respectively.
Nest surveys have been utilized to estimate rail populations
(Pospichal and Marshall 1954; Tanner and Hendrickson 1954, 1956), but rail
nests are difficult to locate and identify.
Playback of tape-recorded calls to evoke responses has proven acceptable.
Tomlinson and Todd (1973) utilized recorded
clapper rail calls to delineate
distribution of two races of this species in southwestern United States and
Mexico.
Mangold (1974) and Holliman (1976) presented auditory indices for
breeding populations of clapper rails in New Jersey and Alabama, respectively.
Dow (1970) found that playing taped calls to obtain auditory indices produced biased results in areas of high densities of birds.
Responsiveness
increased for individual birds and responses by close birds masked distant
responses.
Glahn (1974) found this effect applicable to rail censuses.
Rail breeding populations were estimated by Glahn (1974) by spot-mapping
responses to taped calls.
Recent field studies in Kansas indicated that
estimates of numbers of rails responding to taped calls yielded acceptable
indices of king and Virginia rail populations.
Responses of soras were more
variable and the technique could not be applied to that species (Baird 1974,
Tacha 1975).

�-72-

METHODS
Study Area Selection
One study area was selected to represent typical wetlands in each of four
major regions in Colorado (Fig. 1). Initial selection of study areas was
conducted during May and June 1975.
In April 1976 a study area was added
in northwestern Colorado to better ascertain the status of rails in that
portion of the state.
Criteria for selection included: 1) similarity to
regional wetlands, 2) diversity of emergent types, 3) size approaching 20 ha,
and 4) presence of rails.
Census Techniques
Study areas were censused by playback of tape-recorded calls of rails at
sunrise or sunset May through October 1975 and April through October 1976.
Tape-recorded advertising calls of Virginia rails and soras, recorded from
Peterson's record of western bird songs, were played on a portable Norelco
cassette recorder at 12 to 26 listening sites per study area.
Listening
sites were 80 to 130 m apart.
Amplitude of the taped calls when played
ranged from 80.5 (behind recorder and observer) to 96 db (directly away
from speaker) at one meter and 48 to 65 db at 50 meters (Simpson Model 885
sound level recorder) under favorable weather conditions.
Playbacks per
listening site averaged nine (range 6-15) calls played in a series of
alternating groups of three repetitions of either Virginia rail or sora
calls. A listening period of one minute was allowed between calls.
Calls
of black, king, or yellow rails (also from the Peterson series) were
occasionally played.
Approximate locations and nature of responses by
rails were recorded on field maps (spot-mapped).
Relative breeding densities were estimated for 1975 and 1976 utilizing
results from spot-mapping responses to taped calls. An index of breeding
densities, at each study area for 1975 and 1976, expressed as audibly responding rails/ha (Baird 1974, Glahn 1974, Tacha 1975) was obtained by
averaging results from censuses immediately following peak concentrations
through the first week of July. A second method, employed in 1976 only,
followed the mapping method utilized in breeding bird censuses (Svensson
1970) and used by Glahn (1974). Results from this method were expressed
as territories/ha.
Breeding and wintering distributions of rails were delineated from censuses
of marshes encountered while traveling between study areas, responses to
questionnaires,
and previously published records.
Marshes, encountered
while traveling, were censused utilizing call sequences of 12 to 24 calls
at one to seven sites.
Flushing transects, track searches, and trapping
were utilized when response results were unsatisfactory.
If marshes did
not appear to possess rails during censuses or responses to questionnaires
described marshes and did not specify whether they had breeding rails,
these marshes were judged to be suitable or non-suitable habitat based on
marsh characteristics
and location.
Marsh descriptions and response results

�- ..-

MO"Ar

••

•••
•
••••

••
"""'OOA . "

ItIO'lANCO

SC06WICII

tOGAIt

WUD

11011"

IWlttln

•

:~:.~ ••
MOIf(iAIt

YIIMA

WA$ltIIt~TOIt

•

•

•

•

•••

GA""nD
tlNCDUt

MESA

Klr CARlON

•

•

•

•

,

Clftr~lIlIf

•

.
-...J

,

•

W

KIOWA
Cltowur

•

•
:

IANM/~IIEL

,."OWE'"

•••

1I0LOItfir

•

~CA

MOllrEZIlMA

r

•••

o
Fig.

1.

Study areas

• Trapping sites

• Censused wetlands

Study areas, trapping sites, and wetlands censused during investigations
rails in Colorado, 1975-1976.

of

�-74-

were recorded on field forms (Appendix A). Vegetative dominance was
described as plant species producing Cover judged to be sufficient for
rails to easily hide.
Certain marshes were censused throughout the winter
of 1975-76 to ascertain presence or absence of wintering rails.
Questionnaires concerning distribution and hunter participation
(Appendix B) were
distributed to Colorado Division of Wildlife field personnel in December
1975.
Physical

and Botanical

Dimensions

Field maps of study areas were derived from U. S. Geological Survey 7.5
minute topographic maps and aerial photographs.
Chain (or pace) and compass
and plane table and alidade were utilized to refine details.
Dominant
vegetative cover, based on frequency of occurrence, was sketched on field
maps during a thorough inspection of each study area.
Plant specimens
collected at study areas were filed in the Colorado State University herbarium.
Sizes of study areas and percent composition of major vegetative
types (Appendix C) were calculated from field maps utilizing a planimeter.
Open water, mud flats and dry land areas were subtracted
from total study
area sizes as these areas are less frequently used by rails.
Water and Substrate

Descriptions

Characteristics
of substrates and waters were described for each study
area.
Substrate type was derived from Soil Conservation Service soil surveys adjacent to marshes as marshes are seldom investigated.
Average depths
of organic matter were estimated from footing depths (i. e. depth to solid
footing for the investigator).
Fluctuations of water depths were recorded
during censuses.
Water was chemically analyzed for total acidity, alkalinity
(methyl orange and phenolphthaline),
hardness, and pH using a Hach Chemical
Company Water Ecology Kit, Model AL-36B.
Sources of water were documented.
Microclimatic

Data

Microclimatic data were obtained for May and June 1976 in two marshes with
contrasting species composition.
Temperature and humidity were recorded
on weekly oriented thermo-hygrographs
(Model 3083-AC) made by C. F. Casella
of London.
Thermo-hygrographs
were placed on fixed, sheltered, floating
platforms in locations typical of the respective marshes.
Trapping
Trapping was conducted to aid in ascertaining composition of rail populations
at selected marshes (Fig. 1). Four to nine funnel box traps (adapted from
Bateman 1965), 25 x 31 x 61 cm of 1.4 cm hardware cloth with a 25 x 2-3 cm
funnel opening at one end were used in trapping.
Prior to trapping in 1976,
25 x 25 cm hardware cloth ramps were installed in six traps to more effectively retain rails.
Drift fences 3 to 33 m long of 2.5 cm mesh poultry
netting 30 to 45 cm in height were staked at appropriate intervals with 1
cm diameter x 90 cm wood dowels.
Physical characteristics of trapped and
banded rails were recorded on field forms (Appendix D). Rails were banded
with U. S. Fish and Wildlife Service size 3 aluminum bands.

�-75-

LOCATION

AND DESCRIPTION

OF STUDY AREAS

Location
Study areas were located at: 1) Lower Latham Reservoir (104038' W long.,
40020' N lat.), South Platte River drainage, Weld County, north central
Colorado; 2) John Martin Reservoir (103007' W, 38005' N), Arkansas River
drainage, Bent County, southeastern Colorado; 3) Monte Vista National Wildlife Refuge (NWR) (106004' W, 37028' N), Rio Grande River drainage, Rio
Grande County, south central Colorado; 4) Cary Ranch (107020' W, 40029' N)
and Milk Creek (107044' W, 40017' N), Yampa River drainage, Routt and
Moffat counties, respectively, northwestern Colorado.
Description
The Lower Latham Reservoir study area (13.1 ha in size) occurred at an
elevation of 1,422 m as a wide emergent zone of a large irrigation reservoir.
Typha angustifolia covered 76 percent of the study site with the remaining
area covered by a mixture of Eleocharis macrostachya-Juncus
ba1ticus-Scirpus
americanus (14 percent), Scirpus acutus (7 percent), and seasonally flooded
Poaceae (2 percent).
Lemna minor was locally abundant on the study area.
Average water depths from May through July, 1975 and 1976 were 10 and 2 cm,
respectively.
Water levels were recorded at 0-63 and 0-17 cm in 1975 and
1976, respectively.
Floating vegetative debris (dead Typha) capable of
supporting rails was uncommon in emergent vegetation between May and July.
Organic matter ranged from &lt;2 to 15 cm in depth on a sand to loam substrate.
The John Martin Reservoir study area was an extensive deep emergent marsh
at 1,171 m elevation.
Typha angustifolia-Scirpus
acutus (94 percent) and
E1eocharis macrostachya-Juncus
ba1ticus-Scirpus
americanus (5 percent)
dominated the 14.5 hectares.
Lemna minor and Rorripa nasturtium-aquaticum
were locally abundant within the emergents.
Average water depths were 7
(2-17) and 4 (0-7) cm for 1975 and 1976, respectively.
Recumbent dead Typha
covered 90 percent of the study area.
Open water was present as a result
of muskrat (Ondatra zibethicus) and beaver (Castor canadensis) activities.
Organic matter was estimated to range from 15 to 85 em in depth on a sandy
loam to fine gravel substrate.
At Monte Vista NWR the study area was a shallow emergent marsh at 2,326 m
elevation.
Eleocharis macrostachya-Juncus
balticus-Scirpus
paludosus (59
percent) and Typha 1atifolia-Scirpus
validus (40 percent) dominated this
14.3 ha area. Numerous 1-3 m wide areas of open water were present in the
non-Typha cover (10 percent) but were judged not to warrant separation.
Water depths averaged 5 cm for both years and fluctuated 0-12 and 0-15 cm,
respectively.
Floating debris was uncommon, but recumbent Juncus ba1ticus
was abundant.
Organic matter ranged in depth from 3 to 18 cm on a sandy
loam substrate.
The Cary Ranch study area was a deep open water marsh existing as oxbows
with adjacent seasonally flooded hay fields at an elevation of 1,914 m.
Dominant cover types of this 6.1 ha area were Typha latifo1ia (46 percent),
Carex spp.-Poaceae seasonally flooded (20 percent), Juncus balticus-

�-76-

Eleocharis macrostachya-Triglochin
maritima (16 percent), and Scirpus
validus (16 percent).
Average water depths were 31 (5-130) and 20 (4-91)
cm for 1975 and 1976, respectively.
Mats of dead Typha were locally abundant in water up to 10 cm in depth.
In late summer, mats of Potomogeton
pectinatus, capable of supporting rails, was abundant in open water between
irrigation periods.
Organic matter varied from 0 to 40 cm in depth on a
river gravel substrate.
The Milk Creek study area (4.8 ha) at 1,903 m elevation was separated into
a deep open water marsh (2.4 ha), and a deep emergent marsh (2.4 ha), hereafter referred to as Milk Creek I and II, respectively.
Typha latifolia
dominated both marshes (83 and 71 percent).
Respective minor vegetative
cover was Scirpus validus-Scirpus
acutus (5 and 15 percent), Carex spp.
(7 and 14 percent), and Salix sp. (4 and 0 percent).
Average water depths
in 1976 were 10 and 4 cm, respectively.
Water fluctuated at Milk Creek I
a total of 50 cm due to water level regulation of adjacent Axial Lake.
Milk Creek II, located below the Axial Lake dam had 4 cm of water level
fluctuation.
Recumbent Typha was locally common at both marshes.
Organic
matter depths were estimated at 2-18 cm and 15-60 cm, respectively, on
sandy loam substrates.

Water Chemistry
At regional study areas, water was uniformly basic (pH&gt;7.0) (Table 1). In
Colorado, chemical characteristics of water are usually attributed to the
water's source and the organic make-up of the wetlands.
Subterranean water
sources are moderately baSic, but high organic content tends to neutralize
alkaline waters.
In addition to rainfall, sources of water for study areas
were: 1) Lower Latham Reservoir - primarily irrigation water with some seepage, 2) John Martin Reservoir - springs from a calcareous substrate, seasonally present irrigation water, and septic seepage, 3) Monte Vista NWR _
artesian wells and irrigation water from snow melt, 4) Cary Ranch - irrigation water from snow melt and alkaline shale seeps, and 5) Milk Creek _
snow melt and shale seeps (Soil Conservation Service, pers. comm.).
Climate and Topography
Geographical characteristics of Colorado directly influence regional climatic
conditions which determine quantity and quality of wetlands.
Lower elevations
receive less precipitation than mountain ranges, thus wetlands are restricted
to narrow flood plains of river systems and their associated irrigation
districts.
Wet meadows and hay fields are common in mountain parks and
small basins at high elevations.
Recording stations in the South Platte and
Arkansas River drainages indicate ranges of mean annual precipitation of 18
and 56 and 15 to 63 em, respectively.
In addition, west slope water is
directed by transmountain diversion into these river systems for irrigation
t o: domestic purposes.
Mean annual precipitation for the Rio Grande and
Yampa River drainages ranges from 18 to 43 and 33 to 61 em, respectively
(U. S. Department of Commerce 1975, 1976).
Temperatures normally reflect
elevation and local topography.
Monthly average and average daily high and
low temperatures from April through September 1975 and 1976 for recording
stations nearest to study areas are presented in Appendix E.

,

�Table 1.

Results

of water analysis

at regional

study areas

in Colorado,

August 1975 and April 1976.

Study area

Total 1
acidity-/

Pheno1phthalien
a1kalini ty .!}

Total
1
a1kalinity!J

Hardness.!}

pH

John Martin
Reservoir

0-41

0 •.239

291-2172

239-1419

8.3-9.2

Lowe r Latham
Reservoir

26-48

0 •.34

239-479

393-581

8.2 -9. 1

Monte Vista
NWR

0-9

0-68

68-222

68-120

7.9-9.7

Cary Ranch

0-14

0-342

239-2668

188-889

8.5-9.2

Milk CreekY

38-43

256-308

1710-2309

1265-1402

8.4 -8.6

Y Expressed

as mg/1 of CaC0 •
3

Y Analyzed in April 1976 only.

I
......•
......•
I

�-78-

Microclimate
Temperature and relative humidity in marshes were regulated by macroclimatic conditions, water temperature and height, density, and thickness
of emergents.
High humidity readings (40-100 percent) reflected closeness
of the water's surface.
Differences between average macro- and microenvironmental temperatures at John Martin Reservoir (2l.40C, l4.30C, respectively) were 2.3 times greater than differences at Monte Vista NWR (13.30C,
10.20C) in June 1976. Differences were attributed to 2.7 m tall Typha
growth at the former, and O.S m of standing dead Typha at the latter.
Respective average temperatures in May were (15.90C, l5.70C) and (10.90C,
9.S0C).
Typha averaged 1.4 m in height at John Martin Reservoir and 0.8 m
at Monte Vista NWR in May.
RESULTS AND DISCUSSION
Distribution
Breeding distributions of Virginia rails and soras by townships are presented
in Figs. 2 and 3. Rails recorded in a marsh, May through July were assumed
to be breeding there.
This interval was based on estimated dates of nest
initiation obtained through nest searches and trapping (Table 2). The entire
state was not surveyed (Fig. 1), thus if a species bred in a marsh within a
river drainage it probably bred inuninvestigatedfavorable
habitat throughout that drainage.
High altitude sedge marshes and meadows were noticeably
under-represented.
Altitudinal extremes of breeding records obtained during
this field study were 1,120-3,140 m and 1,120-2,730 m for soras and Virginia
rails, respectively.
Examination of 174 marshes indicated that distribution of Typha-dominated
marshes did not determine the breeding distribution of rails (Table 3).
When marshes with or without Typha as a dominant species and not exhibiting
the presence of breeding rails were compared by chi-square analysis to Typha
or non-Typha marshes possessing breeding rails, a highly significant (P&lt;O.Ol)
difference was apparent.
Further comparisons by regions east and west of
the Continental Divide and above and below 2,100 m elevation also yielded
significant (P&lt;0.05) differences.
Although not included in the comparisons,
stands of Typha latifolia were occasionally present on damp slopes (no
surface water) in western Colorado.
Rails were not observed in those habitats.
Although no positive breeding records are available for king, black, and
yellow rails in Colorado, king and black rails were recorded during this
investigation.
Single king rails were recorded at Bonny Reservoir, Yuma
County, on 25 May 1975 (this study), and near Pueblo, Pueblo County on 12
June through 3 July 1976 (Griffiths 1976). During this investigation a
black rail was recorded at the John Martin study area, Bent County, 11, IS
and 25 June 1975. Black rails were also reported near Red Lion, Logan County,
April 1976(T. Marsh pers. comm.), and at the Arapahoe National Wildlife
Refuge, Jackson County, August 1976 (M. Lindvall pers. comm.).
Breeding
activities (courtship vocalizations) were exhibited by male black and king
rails at Ft. Lyons and Pueblo, respectively.
King and black rails breed
at Cheyenne Bottoms, Barton County, Kansas, 122 km east of Colorado (Baird

�fZj
lZl
GAI",nD

I'
'-.I
\0
1

ID
SANM'~UEL

MO"TI!ZUMA

•• Rails recorded

~

£) Suitable habitat

~
Fig. 2.

Breeding

distribution

of Virginia

rails by townships in Colorado,

1975 and 1976.

t

�rrzy

liNCOLN

~.

o
J

I _BU. _

i.:

~

Fig.

3.

Breeding

distribution

L__
~.._

i

,KIOWA

Suitable

of soras

by townships in Colorado,

1975 and 1976.

habitat

�Initiation dates of clutches of soras and Virginia rails in Colorado by region as estimated
from nest searches,
literature.!../, and estimated ages of trapped chicks.

Table 2.

Virginia

Sora

rail
Number

Range

Region

Mean

Range

Number

Mean

North central

13 June

10 May-l August

15

13 June

22 April-27

Southeast

16 May

10 May-21 May

2

27 May

22 May-2 July

14

Northwest

23 June

22 April-4

13

28 June

20 May-2 August

4

South central

11 June

18 May -20 July

22

13 June

19 May-18 July

23

State -wi de

14 June

22 April-4

52

10 June

22 April-2

64

August

July

23

I

!/

(Bailey and Niedrach

August

1965, Stabler and Kitzmiller

1971)

August

co
f-'
I

�Table 3.

Chi-square analyses testing hypothesis that distribution
determine distribution of rails in Colorado, state-wide,

of Typha-dominated marshes
by region, and elevation.

Number of Marshes
With Tr:eha
Without Tr:eha
With
rails

Without
rails

Total

With
rails

Without
rails

Total

2
X (df = 2)

p

Either species
state -wide

91

34

125

29

20

49

26.41

&lt; 0.01

Soras state -wi de

64

61

125

24

25

49

41.52

&lt; 0.01

Conditions

I

ex&gt;

Virginia rails
state -wide

68

57

125

12

37

49

28.93

&lt; 0.01

Either species East of
Continental Divide

60

18

78

21

10

31

18.38

&lt; 0.01

Either species West of
Continental Divide

29

18

47

8

10

18

10.45

&lt; 0.01

Soras East of
Continental Divide

36

42

78

16

15

31

30.87

&lt; 0.01

Soras West of
Continental Divide

28

19

47

8

10

18

11.24

&lt; 0.01

N

I

�Table 3 (Continued)

Conditions

With
rails

Number of Marshes
Without .!r.E.ha
With TYEha
Without
With
Without
rails
Total
rails
Total
rails

X2 (df = 2)

P

Virginia rails East of
Continental Divide

52

26

78

12

19

31

13.31

&lt; 0.01

Virginia rails West of
Continental Divide

16

31

47

0

18

18

20.45

&lt; 0.01
I
00

Either

w

species

&lt; 2100 rn
species
&gt;2100 m

I

72

31

103

13

4

17

19.27

&lt; 0.01

19

3

22

16

16

32

8.41

&lt; 0.01

Either

Soras

&lt; 2100 rn

46

57

103

11

6

17

38.66

&lt; 0.01

Soras

-&gt;2100 m

18

4

22

13

19

32

6.01

&lt; 0.05

59

44

103

9

8

17

23.56

&lt; 0.01

9

13

22

3

29

32

7.96

&lt; 0.01

Virginia

rails
&lt; 2100 rn

Virginia

rails

&gt; 2100 rn

�-84-

1974, Tacha 1975, Parmelee, Schwilling, and Stephens 1970). Although two
yellow rails have been recorded in Colorado (Bailey and Niedrach 1965,
Colorado Field Ornithologist Records Committee pers. comm.) breeding is
unlikely.
Distribution of rails wintering (30 November-3l March) in Colorado appears
to be restricted primarily to the South Platte and Arkansas River drainages
in eastern Colorado.
Virginia rails were the most common rail recorded,
although soras have been found in winter near Denver (Christmas bird counts,
American Birds 1961-1976), and in the Arkansas River Valley (this study).
In western Colorado, Virginia rails have been reported in winter near
Hotchkiss, Delta County (D. Miller pers. comm.) and on Christmas bird counts
at Grand Junction, Mesa County, and Monte Vista NWR, Rio Grande County in
south central Colorado.
In Colorado, winter rail presence appears to be
dependent upon distribution of warm water sloughs with ample emergent vegetation.
Typha was the dominant emergent in 88 percent (7) of marshes
possessing wintering rails during the winter 1975-76.
The total rail population for the winter 1976-76 was estimated to not exceed 200 individuals
of which 75 to 90 percent were Virginia rails.
Breeding

Densities

Indices of breeding densities for 1975 and 1976 are presented by region in
Table 4. Breeding densities of soras were highest in the south central
region in 1976. Virginia rails had highest breeding densities in 1976 in
southeastern Colorado and appeared to be more common state-wide that year.
Data for the northwestern region were derived from Cary Ranch only. Milk
Creek exhibited breeding densities of 2.01 ± 0.84 S.D., 2.01 ± 0.12 S.D.,
and 4.02 + 0.79 S.D. per ha for soras, Virginia rails and all rails,
respectively for 1976.
By combining the Milk Creek and Cary Ranch study
areas breeding densities for northwestern Colorado in 1976 were 0.80 ±
0.51 S.D., 0.98 + 0.14 S.D., and 1.78 + 0.61 S.D. per ha for soras, Virginia
rails and all rails.
These values are-believed to better estimate breeding
densities for this region.
Hypotheses concerning regional and yearly differences of breeding densities
are tested in Table 5. Indices from study areas for each species were first
compared by F-tests and subsequently by parametric t-tests if F-statistics
were not highly significant (P&gt;O.Ol) (Snedecor and Cochran 1967). Yearly
results were treated as individual samples.
For purposes of consistency,
Milk Creek values were not utilized during testing.
Highly significant (P&lt;O.Ol) t-statistics were obtained for differences in
breeding densities, as exhibited by Virginia rails, between eastern and
western Colorado and differences between breeding densities of Virginia
rails in eastern Colorado and breeding densities of soras in western Colorado
(Table 5). State-wide breeding densities of Virginia rails (2.61 ± 1.66 S.D.
responding rails/ha) were higher than breeding densities of soras (1.26 ±
0.78 S.D.), but differences were not significant (P&gt;0.05).
Breeding densities
of soras in plains study areas (0.73 + 0.34 S.D.) were not significantly
(P&gt;0.05) less than in inter-mountain valley study areas (1.64 + 1.02 S.D.).
There was no indication of a change in breeding densities between 1975 and
1976. However, Virginia rails appeared to occur more frequently at higher
elevations in 1976.

�Table 4.

Indices of breeding densities

of rails in four regions of Colorado,

1975 and 1976.

Density (Responding rails per hectare).!.!
1975
Region

Sora

Virginia
rail

Southeast

0.46+0.57

4.21 + 1. 12

North central

1.13±.0.82

South central
NorthwestY

1976
All rails

Number of
censuses (n)

Sora

Virginia
rail

All rails

Number of
censuses (n)

4.66+

1.58

7

0.44 ±. 0.26

4.67 ±. 0.72

5.11±.0.91

4

3.50 ±. 0.76

4.62+

1.53

5

0.88±.0.62

3.84±.0.96

4.73 + 1.52

3

2.20 ±. 0.19

1. 88 + 0.64

4.08 + 0.83

3

2.51±.1.16

1. 60 + 0.24

4.11 + 1.38

3

1.64+0.71

0.16 + 0.28

1.80 ±. 0.91

3

0.21 ±. 0.25

0.41 ±. 0.21

0.62 + 0.34

3

11
Mean of n censuses

~/
Cary Ranch only.

+ one standard deviation.

-

I
00
VI
I

�Table 5.

Results of testing hypotheses concerning
densities of rails for Colorado.

regional

and annual differences

t Test-

F TestY
df

F

FY

State -wi de , Vi rginia rail
breeding densities equal
sora breeding densities.

7,7

3.79,

6.99

Virginia rail breeding
densities of eastern
Colorado equal sora
breeding densities of
western Colorado.

3,3

9.28,29.5

Hypothesis

c

in breeding

II

Y

. df

t

4.19)~11

14

2.14,

2.98

1. 96

No

4.12

6

2.45,

3.71

4.25**1.1

Yes

F

s

t

c

Reject?

s

I

co
0\

I

Virginia. rail breeding dens it ie s of eastern and western Colorado are equal.

3,3

9.28,29.5

2.89

6

2.45,3.71

6.14**11

Yes

Sora breeding densities
of eastern and western
Colorado are equal.

3,3

9.28,

29.5

9.18

6

2.45,

1. 70

No

Virginia rail breeding
densities are equal each
year.

3,3

9.28,

29.5

1. 19

6

2.45,3.71

0.14

No

Sora breeding densities
are equal each year.

3,3

9.28,

29.5

1. 96

6

2.45,3.71

0.54

No

!l df = degrees
. .
t-stahshc:

t

of freedom: F = critical
C
..
= ca 1cu 1ate d t-stahstlC.
s

F-statistic:

F
s

= calculated

21 a. = 0.05 and 0.01.

-

11 * = significant

(P&lt;O.OS):

~(*= highly significant

(P&lt;O.Ol).

3.71

F-statistic;

t

c

= critical

�-87-

Breeding densities obtained in other studies are generally higher than
those reported in this study.
In Iowa, breeding densities of both soras
and Virginia rails approached eight pairs per hectare of marsh habitat
(Tanner and Hendrickson 1954, 1956).
In Minnesota, sora breeding densities
were as dense, but Virginia rails exhibited highest densities of 3.7 pairs
per hectare (Pospichal and Marshall 1954).
Glahn (1974) found breeding
densities in north central Colorado to be as high as 1.0 and 2.3 pairs per
hectare for soras and Virginia rails, respectively.
When breeding densities of all rails for 1976 as determined by spotmapping territories (Table 6) and averaged spring censuses (Table 4) were
cQmpared by linear regression (Fig. 4), a significant (P&lt;0.05) correlation
(Y = 0.20 + 0.89x; r = 0.97) was obtained.
Like comparisons were applied
to individual species (Figs. 5 and 6).A Methods for estimating breeding
densities of soras were more erratic (Y = -0.36 + 1.12x; r = 0.93; P&lt;O.lO)
than were the same methods for estimating breeding densities of Virginia
rails
= 0.16 + 0.95x; r = 0.25; P&lt;0.05). However, both regression lines
approached a theoretical line (Y = 0.0 + 1.Ox; r = 1.0) which would describe
identifical results for two censusing methods.
Thus, assuming that sample
sizes were adequate, utilizing only averaged spring censuses for analyzing
differences between breeding densities by region and years was justified.

(Y

Table 6.
~

Breeding densities of rails at five regional study areas
in Colorado in 1976 as determined by spot-mapping
territories.

Study area

Size in
hectares

Dens it:x:{ter ritorie 5 /ha)
All
Virginia
rails
Soras
rails

Latham Reservoir

13.1

4.73

1,53

3.21

John Martin Reservoir

14.5

5.86

0.76

5.10

Monte Vista NWR

14.3

3.99

2.38

1. 61

Cary Ranch

6.1

0.82

0.33

0.49

Milk C'reek

4.8

3.55

1.67

1. 88

I

2.4

0.84

0.42

0.42

II

2.4

6.25

2.92

3.33

�-88-

Q)

J.l
C1I

•

5

~
o
Q)

::r:
J.l
Q)

p..

4

III

.....•

...•
C1I

IX:
co
J::

....

3

A

"tj

Y=O.20+0.89x

0
0..

r = 0.97

J::

~

III
Q)

IX:

2

'+-I

•

0
J.l
Q)

S::s 1
Z
Q)

co
C1I
J.l
Q)

&gt;

&lt;

0
0

Fig. 4.

1
Estimated

2
3
4
5
Density of Breeding Territories

Breeding densities of all rails as determined by
spot-mapping ter~tories
co r re lated to averaged
spring censuses (Y = 0.20 + 0.89x; r = 0.97;
P&lt; 0.05).

6

�-89-

5

4
Ul

....•
ro
cr:;
.-I

~
ij
s::
o
0..
Ul
CLl

cr:;

1\

Y = -0. 36 + 1. 12 x

3

r

= 0.93

2

'H

o
~
CLl

S 1
i

•
•
o
Fig.

•

1
Estimated
5.

2
3
4
5
Density of Breeding Territories

Breeding densiti e s of soras as determined by
spot-mapping territories
correlated
to averaged
A
spring censuses (Y = .0.36 + 1. 12 x; r = 0.93;
P&lt;O.lO).

6

�-90-

•
•

"

+ 0.95x

Y = 0.16
r = 0.97

o

1
Estimated

Fig.

6.

2
Density

3
of Breeding

4

5

6

Territories

Breeding densities of Virginia rails as determined
by spot-mapping
territories
correlated
to averaged
1\
spring censuses (Y = 0.16 + 0.95 x; r :: 0.97;
P&lt;0.05).

�-91-

Further analysis revealed that composition of breeding populations was
dependent upon regional temperatures during early phases of migration.
Spring migration began in April and probably peaked in late April to early
May. Random migration throughout Colorado by both species and like attractant forces (i.e., cover, food, etc.) at all marshes were assumed.
When
mean temperatures for April near study areas were compared to percentage
of species composition (Fig. 7), a highly significant (P&lt;O.Ol) relationship
= 7.25e-l•96x; r = -0.94) was apparent. As temperatures decreased, sora
populations increased.
Soras began to compose 50 percent of the breeding
population as mean April temperatures approached 5.60C. Temperatures in
April 1976 at all study areas were higher than in 1975. At all study areas
except Monte Vista NWR, 1975 breeding populations had higher percentages
of soras. Other factors may more strongly regulate composition in other
states, but indications were that the breeding range of the sora or Virginia
rail in Colorado was dependent upon temperature.

(Y

Habitat Utilization
Periods of importance when considering habitat utilization were spring
migration, nesting, brood development, and fall migration.
In all periods
quality of cover was most important.
During brood development and fall
migration food quantity was assumed to be important.
Spring Migration
Spring migrants utilized dead, standing emergents, primarily Typha.
Although
up to 90 percent of the John Martin Reservoir study area had been burned
during the winters of 1975 and 1976, remaining old growth and new growth
«30 cm) Typha attracted substantial numbers of rails.
In marshes not containing Typha, rails utilized any emergent cover available.
In western Colorado soras were observed in Salix spp. during spring floods.
At other times
herbaceous
emergent vegetation was commonly utilized by rails when hiding
from the investigator.
There was often snow or ice in emergent vegetation,
particularly Typha.
Nesting
In Colorado rails preferred nesting in cover of robust emergent vegetation
(i.e., Typha, Scirpus), particularly Typha.
Eighty percent of rail nests
(25) found during this investigation were in Typha spp. Responses by both
species of rails during the nesting period (May through June) at the Monte
Vista NWR study area indicated that when a choice was present Typha (40
percent coverage) was chosen over short fine vegetation (i.e., Eleocharis,
Juncus).
Soras responding in Typha comprised 78 percent of the total soras
recorded, while 91 percent of the Virginia rail responses were in Typha.
Small percentages of sora (21 percent) and Virginia rail (4 percent)
territories as determined for Monte Vista in 1976 were in the short fine
vegetation.
Deviations from the trend to utilize Typha were apparent at
higher elevations where Carex was the common emergent or when high water
made Typha unavailable.

�-92-

5

4-

ro,..
0
U)

tl.O

.S
Q)

It:)

~
0

:;;

ro
...•
3-

~

0..
0

fl.

.•.~
Q)

U
I--&lt;
Q)

fl.

j

2-

1
1

Fig.

7.

Relationship
between m.ean April tem.peratures
(oe) near
study areas and percent com.position of the breeding populations,
95 percent confidence intervals
depicted (~ = 7.25 e -1.96 x
r = -0.94;

P&lt; 0.01).

�-93-

Nests located during this investigation were combined with nesting data
compiled by Kaufmann (1971). These data indicated that a higher percentage
of Virginia rail nests were found in robust vegetation than were sora
nests.
Seventy-one percent of all Virginia rail nests (167) and 60 percent
of all sora nests (223) were in robust vegetation.
Approximately 50 percent of those nests in robust vegetation were in Typha.
Results from nest
searches are often biased as nests in short emergents are probably more
easily discovered.
Pospichal and Marshall (1954) and Glahn (1974) suggested
that rails locate nests adjacent to or near openings in emergents.
Too few
nests were located in this study to test this hypothesis.
Water depth at initiation of nesting may affect breeding densities of rails.
Marshes at Cary Ranch were dry in 1976 until after peak migration periods
and subsequently harbored a small rail population (Table 2). There appeared
to be a correlation between density of all rails and percentage of suitable
emergents with water less than 15 cm at initiation of spring censuses.
Previous studies have satisfactorily proven that rails prefer water 0-8 cm in
depth (Baird 1974, Tacha 1975).
Implications are that since rails prefer
not to swim (Kaufmann 1971) they utilize areas of marshes with water depths
not exceeding the length of their legs (7-8 cm). Actual water depths may
exceed 8 em, but floating dead or living vegetation is often sufficient to
support the weight of rails.
The floating mat formed by dead recumbent
Typha often persisted through the summer months in marshes in Colorado.
Degree of water level fluctuations appeared to affect nesting activities.
In 1976 Cary Ranch had fluctuations of water levels totalling 87 cm between
extremes and only one Virginia rail territory of five possible rail territories remained intact.
Milk Creek I had fluctuations of water levels
totalling 50 cm between extremes and again only one Virginia rail territory
remained intact.
A sora nest was found abandoned with seven eggs at Milk
Creek I following a rise in the water level of 30 cm in a one week period.
Initially, five sora and five Virginia rail territories were present at
Milk Creek I. In cases where water fluctuated less than 20 cm or if water
levels decreased gradually densities did not appear to be affected.
Nests located in Colorado in 1975 and 1976 were in average water depths of
8.2 cm. Of those nests positively identified, sora nests (4) were in water
depths averaging 10.7 cm of water, while Virginia rail nests (9) were built
over 7.1 cm of water.
In most situations a floating vegetative mat was
present.
Brood Development
Responses (peeps) of chicks during the brood development period appeared
to originate totally from robust emergent vegetation.
Immature rails less
than one month in age were trapped consistently in Typha in 1975 and 1976.
Andrews (1973) trapped most rails during July (late brooding period) in
Typha dominated vegetation.
Typha apparently furnished the necessary cover
and favorable food resources for developing chicks.
Chick mortality was noted (2 Virginia rails, 1 sora) at Monte Vista NWR in
1976 after receding water levels left 90 percent of the study area without
water.
All chicks were found farther than 100 meters from the nearest water.

�-94-

Estimated age of chicks was less than four days in all cases.
There was
no indication as to cause of death, but two dead chicks (Virginia rails)
were found near dried mud holes, perhaps indicating separation from the
brood and subsequently dying for lack of water.
Fall Migration
Rails, primarily sora, were flushed from short emergents during fall migrations.
Number of vocalizing rails elicited by taped calls also increased
in Typha marshes with sufficient water.
Water depths preferred during
migration varied from &lt;1 (muddy) to 60 cm in areas of responses and sightings.
Andrews (1973) indicated soras preferred reflooded areas of Polygonum
lapthifolia, Echinichola walteri and Cyperus erythrorhizos during fall concentrations, probably because of the abundance of food.
In Colorado,
Virginia rails appeared to remain more in Typha as indicated by responses.
Timing of Migration
Spring
During this investigation rails were first observed in previously rail free
marshes as early as the second week of April.
Soras arrived at the John
Martin Reservoir study area on or prior to 10 April 1976.
In north central
Colorado near Ft. Collins, Virginia rails were first observed on 11 April
1976.
Bailey and Niedrach (1965) indicated that spring arrival of rails
in north central Colorado was in early May, but arrival of soras in northwestern Colorado began as early as late April.
Results from this investigation suggested that Virginia rails arrived in eastern Colorado prior to
the arrival of soras.
Densities of rails responding to taped calls as depicted in Figs. 8-12
indicate peak concentrations state-wide occurred within the first and
second weeks of May.
In other studies (Glahn 1974, Baird 1974) a two to
three week period of no vocalizations occurred after the arrival of rails.
A silent period after arrival might therefore disgui~e an earlier peak
concentration.
Taking this potential silent period into consideration,
actual peaks in spring migration should have occurred in late April to
early May.
In either case, spring migration did not appear to differ between plains and inter-mountain valleys.
Soras appeared to reach peak concentrations one to two weeks prior to Virginia rails in southeastern and
northwestern Colorado.
In north central and south central Colorado, peak
concentrations occurred simultaneously.
Late Summer Movement
Marsh conditions, specifically water levels, obviously caused early desertion of marshes by rails. At Monte Vista NWR in 1975 and 1976 and at
Latham Reservoir in 1976 decreasing water levels reduced amounts of suitable
habitat by 95 percent in July or early August.
At Monte Vista many rails
were concentrated in sparse cover near remaining water.
Rails were easily

�~

-...".

Soras

Virginia

o

1975

mill··

~

1976

~

10

Rails

* No Census

d)

~

•..uro
d)

::r:
~

o,d)

(J)

...•
ro

I
\D

VI
I

cr:;
d)

:&gt; 5
...•
(J)

~

0
0..

(J)

d)

cr:;

....
0

~
d)

.0

S::l
Z
:::&lt;

0

ft.

1

4
April

Fig.

2

3
May

8.

4

1

2
3
June

Densities
of rails responding
September,
1975 and 1976.

4

":, (:T'
"

pm ':'
tt.

3412341
July
Week of Census

to taped

l2

calls

at John Martin

2
3
September

August
Reservoir,

April

through

4

�Soras

10

.,

Virginia

D

1975

~

1976

Rails

fill.
~

(1)

~

*

n!

No Census

()
(1)

:r:
~

o,
(1)

(/)
.....•
......

ro

r::r;

~

(1)

&gt;
......

I
\0
0'
I

5

(/)

r::
0
0..
(/)

0
I-&lt;

I

(1)

~

!

:::I

:::i
.:::1'
~...
C·"
~:::

'+-&lt;

S

...
:::1
...
...
...
...
...
...
...
...
...
...
,..
...
...
...
...
...
...
...
...
...
...
...
...,.,
...
...
...
...
.. ,
g~':&lt;

~
!:::I

(1)

r::r;

f::

i

.]

~

r'

:;~

0 ~
2

3

1W':,

1111,:&lt;

4

1

2
3
June

4
April

1

Fig.

Densities
of rails responding
September,
1975 and 1976.

9.

May

'...."........
.,.
'..
,..
...
,..
,..
...

""

.'".....
...
...
.......
........
...
...
.....
....
'"
...
...
...
...
...
.....
.....
.....
...

:::1
mi
... ,
:::i
:::1

;;;!

g~~:&lt; ~g ~:&lt; 1:: ,:&lt;
1

2
1
2
2
1
4
3
3
3
4
August
September
July
Week of Census
to taped calls at Lower Latham Reservoir,
April through
4

1

::::1':&lt;
4

�Soras

10 l

I

a&gt;
~
.....,

()

a&gt;

n

1975

~

1976

Rails

mm
lliill
~

,:~ No Census

1

(Ij

Virginia

::r::

'"'

c,a&gt;
Ul

.-&lt;

.....•
(Ij

~
a&gt;

:&gt;
.....•

I
\0

5

'-l

Ul

r+:

~

r--

0
0..
Ul

a&gt;
~

....

I

f:mi

0

r:

S~

z

0

r

l -'-r-

r

4
April
Fig.

1

2
May

10.

3

4

1

2
3
June

Densities
of rails responding
September,
1975 and 1976.

"
4

I
I!

::::/

I~

'"'

a&gt;
,..0

I

gm •...
......

....j'
"
W
..../
....
....
.... ,

-..
...

1111! 111
-'-J'ml':~
-r- :~
m:/ ,:~ H~'''I

t.v.1

I

1

2
3
July
Week of Census

to taped

calls

4

1

at Monte

Vista

I

1""IXXld

2
3
August
NWR,

I

I

4

1

April

through

I

I

2
3
September

I

4

�Soras

10

Virginia

o

1975

~
IiliiJ

§§

1976

~

Rails

(1)

I-t

..,I'd

No Census

Yoc

u
(1)

::c
I-t

c,
(1)

CIl
....•
....•
I'd

~

I
\0

(1)

::- 5
....•
CIl

0:&gt;

-1

I

I

~

I

0
0..

l

CIl
(1)

~

"'0"
!-I
(1)

,.0

S
~
0

,

,
1

4

April

Fig.

11.

':' 1::::1'
.... :'
2
3
May

4

Densities
of rails
1975 and 1976.

1

2
3
June

responding

4

1

I:::;
::: ':'

2

3

4

1

July
Week of Census
to taped

calls

at Ca r y Ranch.

2
3
August

April

4

through

1

2
3
September

September,

4

�Soras
10

Virginia

o

MC!

~

MC II

mm
llilli
~

Q)

I-f

•..

No Census

&gt;:&lt;

('(j

Rails

()
Q)

::r::
I-f
Q)

Ili

.-&lt;Ul
.....
('(j

p:;
I

Q)

~
~

.....&gt; 5

I

Ul

~

0
0..
Ul
Q)

~
'+-I

0
I-f
Q)

..0

S::s

Z

0

I * ~:~
4
April

Fig.

~~&lt; &gt;!:

1

2

3
May

12.

4

1

2
3
June

Densities
of rails responding
September,
1976.

4

1

2

3
July
Week of Census

to taped

calls

4

at Milk Creek

1

2
3
August

4

I and II, April

1

2
3
September

through

4

�-100-

observed as well as trapped, and on successive visits noticeable reductions
were apparent. Tracks on mud banks of ditches indicated that rails left
by foot as well as by flight. Bent (1926) mentioned a late summer movement
of rails to more stable marshes. Peaks in densities in late July and early
August at study areas may have resulted from movement from nearby drying
marshes. Irrigation practices in Colorado promoted midsummer draw downs of
wetlands.
Fall
Although responses were difficult to evoke in August and September, indications of peak concentrations were obtained in mid-August to mid-September.
Local movement in August may mask actual migration. Soras appeared to have
a more extensive migration than Virginia rails, beginning earlier, peaking
at higher concentrations, and ending later. Extent of migration for Virginia rails was less distinct, but appeared to generally coincide with sora
migration.
Peak fall concentrations of rails in inter-mountain valleys were not pronounced possibly indicating that small numbers of rails migrate through
western Colorado. Observations of migrating soras (2 individuals) at Milk
Creek on 18 September 1976 may have indicated similar migrational patterns
on both sides of the Continental Divide. However, fall migrational peaks
were usually indistinct, making comparisons difficult.
Trapping
During this study, 294 rails (142 soras, 152 Virginia rails) were banded at
nine trapping sites primarily during July, August, and September 1975-76
(Table 7). Two Virginia rails accounted for two returns at Wellington
Wildlife Area, Larimer County. One Virginia rail was caught on three
occasions over a 213 day period. Trap mortality was less than two percent.
In 1975 trapping success was 0.06 rails per trap hour for 1,617 hours.
Trapping success approached 0.20 rails per hour in 1976 for 1,100 trap hours.
Although trapping methods and time periods may bias results, adult to juvenile ratios for soras and Virginia rails were 1:4.7 and 1:2.9, respectively.
A major portion of clutch initiation dates utilized in Table 2 was obtained
as a result of trapping. Approximate clutch initiation dates were obtained
by estimating ages of chicks less than six weeks of age and allowing 19 days
for incubation for both species (Walkinshaw 1937, 1940; Wood 1937; Pospichal
and Marshall 1954; Tanner and Hendrickson 1954, 1956). A staggered hatch
of 7.8 days (Kaufmann 1971) for soras was taken into consideration. Measurements of tarsus, culmen and weight of developing chicks obtained in Minnesota (Pospichal and Marshall 1954) were used to estimate age. Relative
lengths of outer primaries and capability of flight were also used to approximate age of chicks between three and six weeks of age. Chicks older than
six weeks of age were not aged. Since trapping was conducted intensively
in late July through early September, mean clutch initiation dates were
weighted toward late hatching broods as early hatched young could have left
the areas prior to trapping. Nonetheless, results showed that flightless
young and adult rails were present in Colorado in early September and as
late as mid-September.

�Table 7.

Distribution

of bandings

of rails,

County and Location

by species,

Colorado,

1975-1976.

Soras
Aug. Sept.

Number

Banded

Total

Feb.

April

I

I
I

-

-

I
16

Virsinla rails
July Aug. Sept.

Nov.

Total

3
2

-

3
6

7
24

0
0

I
2

-

-

I
2

-

-

I
5

-

-

I
5

-

-

-

2
3

11
9

I
0

2
0

-

10
19

4
11

-

-

J
18

-

-

-

I
1

I
1

-

2
2

-

-

0
0

-

-

0
0

-

-

-

0
0

.-

-

I
55

-

-

5
31

I
10

-

-

6
41

2
6

-

2
6

-

-

-

-

3
6

-

3
6

11
22

I
I

27
liS

I
0

2
0

6
47

17
38

10
21

3
6

39
113

Feb.

April

July

Adult
Juvenile

-

-

0
0

0
0

-

Adult
Juvenile

-

-

3
2

0
4

-

-

3
6

-

-

Adult
Juvenile

-

-

-

6
10

-

-

6
10

-

Adult
Juvenile

-

-

-

2
10

0
0

0
0

5
5

6
4

-

2
10

Adult
Juvenile

-

Adult
Juvenile

-

-

0
16

I
2

Adult
Juvenile

-

-

0
0

-

Adult
Juvenile

-

-

1
49

0
6

Adult
Juvenile

-

-

-

-

4
51

11
41

Nov.

~
John Martin

Reservoir

I

~
Hotc hkl s a
Jackson
North Park
Larimer
Fossil

Creek

Re s e rvot r

Wellington Wildlife
Management Area

~
Milk Creek

11

-

-

~
Pueblo

CF&amp;I Reservoira

Rio Grande
Monte Vista NWR

-

Weld
Latham

Reservoir

Totals

Adult
Juvenile

2
3
17
30

I

I-'

0

I-'

I

�-102-

SUMMARY
Investigations of rails were conducted in Colorado between May 1975 and
October 1976.
Investigations were conducted state-wide with major study
areas in the Arkansas, South Platte. Rio Grande, and Yampa River drainages.
Objectives were to determine distribution, breeding densities, timing of
migration and habitats utilized.
Breeding distributions of soras and Virginia rails were delineated by townships. As rails were found to nest between 22 April and 4 August in Colorado, rails recorded in marshes during that period were assumed to be
nesting.
Soras were found to breed to 3,140 m elevation, while Virginia
rails were found to 2,730 m. King and black rails were not recorded to
breed in Colorado, although they were recorded two and one times, respectively, during early summer in eastern Colorado.
Examination of 174 marshes indicated that the distribution of Typha dominated marshes did not determine the breeding distribution of rails in Colorado. A highly significant (P&lt;O.Ol) number of marshes not having Typha
had breeding rails, and as many Typha-dominated marshes appeared to have
no rails.
Wintering rails were found in warm water marshes in the South Platte,
Arkansas, and Gunnison river valleys.
Total rail populations for the winter
of 1975-76 were estimated not to exceed 200 individuals of which 75 to 90
percent were Virginia rails.
Indices of breeding densities were obtained by averaging census results
between mid-May and early July.
State-wide, Virginia rails exhibited a
breeding dnesity of 2.61 + 1.66 (S.D.)/ha which was higher (P&lt;O.lO) than
the density of soras (1.26 + 0.78).
Virginia rails had higher breeding
densities in the eastern plains (4.06) than in inter-mountain valleys (1.16).
Breeding densities did not change substantially between 1975 and 1976,
although Virginia rails occurred more frequently at higher elevations in
1976.
Indices of breeding densities, obtained by averaging census results
were correlated significantly (P&lt;0.05) to estimates of densities of territories obtained by spot-mapping responses.
A highly significant (P&lt;O.Ol) relationship was obtained between mean April
temperatures and percent composition by species.
The equation Y = 7.25e-l. 96x
where x equals mean April temperature (OC) gave the approximate percentage
of soras expected to breed in a particular marsh.
Thus, temperatures during
early migration appear to affect composition of breeding rails in Colorado
marshes.
As the majority of suitable marsh habitat in Colorado was vegetatively
dominated by Typha, rails appeared to utilize cover of Typha more often
than other species of emergents.
Eighty percent of the 25 rail nests found
during this investigation were in Typha.
During fall migration soras preferred shorter emergent vegetation possibly due to greater food availability,
while Virginia rails generally remained in tall, dense vegetation (e.g.
Typha).

�~

I

-103-

Rapid water level fluctuations disrupted breeding activities.
Desertion of
established territories and one sora nest with seven eggs was observed when
water levels rose more than 20 cm in northwestern Colorado.
Decreasing
water levels in south central Colorado prevented renesting and apparently
caused chick mortality in an extreme case.
Spring migration into Colorado by soras and Virginia rails appeared to coincide beginning during the first and second week of April, peaking the last
week of April or the first week of May.
There was no indication that migration differed notably in plains and inter-mountain valleys, but a week
delay in peak migration may occur in the mountains.
Late summer movement of rails from drying marshes was pronounced.
Irrigation
practices and water laws in Colorado contribute to decreasing water levels
in late July. Rails appeared to leave drying marshes by flight and afoot
along ditches.
Fall migration began in mid-August and peaked in early September on the
plains.
Migration was less pronounced in inter-mountain valleys possibly
indicating that few rails migrate through western Colorado.
Soras appeared
to outnumber Virginia rails during peak migration periods in plains study
areas.
The Rocky Mountains may serve to funnel more northerly breeding
rails (primarily soras) through the plains of Colorado.
Trapping was conducted utilizing funnel box traps with drift fences.
During
this study 294 rails (142 soras, 152 Virginia rails) were banded.
There
were 63 recaptures.
One Virginia rail was captured on three occasions over
a 213 day period.
Ages of trapped chicks were utilized to estimate mean
clutch initiation dates.
Late summer adult to juvenile ratios for soras and
Virginia rails in trap samples were 1:4.4 and 1:3.2, respectively.

�-104-

LITERATURE CITED

Allen, A. A. 1934. The Virginia rail and the sora.
36'(3):196-204.

Birdlore

American Ornithologist's Union. 1957. Check-list of North American birds.
5th Ed. Lord Baltimore Press, Inc, , Baltimore.
691 pp.
Andrews, D. A. 1973. Habitat utilization by s or as , Virginia rails,
and king rails near southwestern Lake Erie. M. S. Thesis,
Ohio State Univ., Columbus. 112 pp.
Bailey, A. M., and R. J. Niedrach.
1965. Birds of Colorado.
Denver Mus. Nat. Hist., Denver. Vol. I. 454 pp.
Baird, K. E. 1974. A field study of the king, sora and Virginia
rails at Cheyenne Bottoms in west central Kansas. M. S.
Thesis, Fort Hays Kansas State CoIl., Fort Hays. 37 pp.
Batemann, H. A. 1965. Clapper rail (RaIlus longirostrus) studies
on Grand Terre Island, Jefferson Parish, Louisiana. M. S.
Thesis, Louisiana State Univ•• Baton Rouge. 145 pp.
Bent, A. C. 1926. Life histories of North American marsh birds.
U. S. Natl. Mus. Bull. 135. 490 pp.
Billard, R. S. 1947. An ecological study of the Virginia rail (Rallus
limicola) and sora (Porzana carolina) in some Connecticut
swamps. M.S. Thesis, Iowa State Univ., Ames. 84 pp.
Boeker, H. M. 1954. A census of populations of the Wilson's snipe
and sora rail in the Yampa River Valley, Colorado. Condor
56(2):105-106.
Dow, D. D. 1970. Indexing population densities of the cardinal with
tape-recorded song. Wilson Bull. 82(2):83-91.
Glahn, J. F. 1974. Study of breeding rails with recorded calls in
. north central Colorado. Wilson Bull. 86(3):206-214.
Griffiths, D. 1976. King rail - first Colorado record.
Ornithol. J. 28:17•

Colo. Field

�-105-

Holliman, D. C. 1976. Clapper rail (Rallus longirostrus) studies in
Alabama. U.S. Fish and Wild!. Servo Accel. Res. Program.
Project Final Rep. Contract No. 14-16 -0008 -703. 50 pp.
Horak, G. J. 1970. A comparative study of the sora and Virginia
rail. Wilson Bull. 82(2):206-213.
Kaufmann, G. W. 1971. Behavior and ecology of the sora, (Porzana
carolina) and Virginia rail (Rallus limicola). Ph. D. Thesis,
Univ. Mi nn, , Minneapolis. 114 pp,
Kellogg, P. P., A. A. Allen, and R. T. Peterson.
1962. A field
guide to western bi r d songs. Houghton Mifflin Co , , Boston.
Phonodisc.
Lane, J. A., and H. R. Holt. 1975. A birder's guide to eastern
Colorado. L. and P. Press, .Denver. 136 pp.
Mangold, R. E. 1974. 1974 Final Report: Research on shore and
upland migratory birds in New Jersey, Clapper rail studies.
U.S. Fish and Wildl. Servo Accel. Res. Program.
Contract
No. 14-16-008-937.
17 pp.
Meanley t B. 1965. Early fall food and habitat of the sora in the
Patuxent River Marsh, Maryland. Chesapeake Sc i , 6(4):
226-227.
I

1969. Natural history of the king rail.
No. 67. 108 pp.

t

N. Amer.

Fauna

Parmeleeo D. F •• M. D. SchwiHing, and H. A. Stephens. 1970.
Gruiform birds of Cheyenne Bottoms. Kans as Ornithol. Soc.
Bull. 21(4):25-27.
Pospichal, L. B., and W. H. Mar sha ll., 1954. A field study of the
sora rail and Virginia rail in central Minnesota. Flicker
26(1):2_32.
Post,

W., and F. Enders.
1970. Notes on a salt marsh Virginia
rail population. Kingbird 29(2):61-67.

Ripley, S. D.
. In press.

1977.

Rails of the world.

Godine Press,

Boston .

Smith, A. B. 1955. Sora rail populat ions in Alberta, 1953-54. In
Investigations of woodcock, snipe, and rails in 1954. U.S. Fish
and Wildl. Servo Sci. Rept. Wildl. 28. pp. 59-62.

�-106-

Snedecor, G. W., and W. G. Cochran. 1967. Statistical methods.
Iowa State Univ. Press, Ames. 593 pp.
Stabler, R. M., and N. J. Kitzmiller.
1971. A second Virginia
rail nesting for El Paso County, Colorado. Colo. Field
Ornithol. J. 9:21.
Svensson, S. 1970. An international standard ·for a rnappirig method in
bird census work recommended by the international bird
census committee. Audubon Field Notes 24(4}:722 -726.
Tacha, R. W. 1975. A survey of rail populations in Kansas, with
emphas is on Cheyenne Bottoms. M. S. Thesis, Fort Hays
Kansas State ceu., Fort Hays. 54 pp.
Tanner, W. D., and G. O. Hendrickson.
Virginia rail in Clay County, Iowa.

1954. Ecology of the
Iowa Bird Life 24(4}:65-70.

1956. Ecology of the sora in Clay County, Iowa.
Bird Life 26(4):78 -81.

Iowa

Tomlinson, R. E., and R. L. Todd. 1973. Distribution of two
western clapper rail races as determined by responses to
taped calls. Condor 75(2):177 -183.
U. S. Dept. of Com.m.erce. 1975. Climatological data: Colorado.
National Oceanic and Atmospheric Admin. 80(13):1-13.
• 1976. Climatological data: Colorado. National Oceanic
---and Atmospheric Admin. 81(13):1-14.

I

t
Walkinshaw, L. H. 1937.
54(4):464 -47 5.

The Virginia rail in Michigan.

Auk

I

4
1940.

Sum.m.erlife of the sora rail.

Auk 57(2):153-168.

Webster, C. G. 1964. Fall foods of soras from two habitats in
Connecticut. J. Wildl. Manage. 28(1):163 .•165.
Wood, H. B. 1937.
54(4):535 -536.

Approved

by_~{!Ia~U~'

Incubation period of the Virginia rail.

----!.~='_'_~~~~=.!::.:....=..._

Clait E. Braun
Wildlife Researcher

Auk

�-107-

APPENDICES

�-108-

APPENDIX A
Marsh Investigation

DATE- __

-

_______

__

1

- Rail Observations

9--= TIME-_to_hrs.;

LOCATION-

County, U.S. -Colo, -_HWY.

MAJOR RIVER DRAINAGE- ..•......

, __

~miles __

---:: DISTANCE TO NEAR-

EST PERMANENT STREAM___________
SIZE__

-Stream-

---': ELEVATION -__
~ha: Continuous

--.-

- Disjunct;

RAILS RECORDED-

% WETLAND TYPES-~
---..

em,

---"{Sora),

{

---"(Virginia Rail),

; % MARSH CENSUSED-

CHARACTERISTICS OF R~L

WEATHER-

...;m-.: MARSH

: VEGETATION COMPo -

AVERAGE WATER DEPTH-

_____

of

_

OCCURRENCE-

.....•..

, T emp.

_

_

°c.

�-109-

APPENDIX B
Studies of Rails in Colorado

1. District

--------------------------~------------~--------

2.

Do rails occur to your knowledge in your present district?
Yes_
NO_If
no, answer question No. 5 only.

3.

If rails are known or suspected to occur in your district,
list all locations. Generalizations can be made:

please

---------------

4.

What vegetation types occur where you hear or see rails?

5.

Please list major cattail marshes in your district not mentioned above.

6.

What time(s) of the year do you most commonly hear or see
rails?

7.

Have you encountered rails dur ing the 1 November through 31
March period? Yes
No·
If yes, where?

-

8.

-

Have you ever encountered any rail hunters in your district?
Yes_
No_If
yes, where?

_

�APPENDIX C
Percent

Dominance of Vegetative

Species at Regional Study Areas

Stud}: Area
Monte Vista
NWR

Cary
Ranch

Milk
Creek

L. Latham
Reservoir

John Martin
Reservoir

76

86

TI..eh! latifolia

-

Trace

35

46

77

Juncus balticus

5

3

39

5

Trace

Scirpus

acutus

1

8

1

3

Scirpus validus

Trace

-

-

4

16

7

3

Trace

8

1

10

9

Trace

10

Trace

-

1

2

+99

+99

+99

Species
Typha angustifolia

Scirpus

americanus

5

1

Scirpus

paludosus

1

-

10

4

1

10

-

-

-

Eleocharis

macrostach.!!.

Triglochin

maritima

Carex nebraskensis
Poaceae
Salix spp.
Total

,

.

1

-

Trace

Trace

+99

+99

I

I-'
I-'
0

I

�-111-

APPENDIX D
Rail Banding Form
Species

_
~ Band #

_

Location; County

_

Wetland Type

; Vegetation,

_

Date

Time

--!

Weight

--'gm; estimate

Lengths:

Culmen,

(-c1aw)
_____

--..;mm; Middle toe

--..;mm; Wing chord,

--..;mm; Radius -ulna

~mm;

8
7
6
5
4

3
Z

1
Plumage

wing

Retricies -right side

mm - Outer
mm
mm
mm
mm
mm
mm
mm
mm
mm

9

_

rnrn; Tarsus

Primaries-right
10

of time in trap,

Description:

6

mm - Outer
mm
mm
mm
mm
mm

5
4
3
Z
1

---I Back,

Breast,

F~ank

J

_

Sora only- Brown auricular

patch (joined to nape) (separ.ate)

Gray supracilliary

line (broken by black in front)

(unbroken)
Soft part coloration:

Legs
Iris

Specimen fI's:

--'
------

#4 primarY
white flank feather

REMARKS:

Bill

--'

_
---I most anterior

black 8.

,
_

�Appendix E.

Average,
September,

Region
of
ColoradoY

Ave.
Dev.

deviation

from normal!!,

average

1975 and 1976, at regional

Aeril
Ave. Max.
Ave. Min.

Ave.
Dev.

dally maximum,

study areas

Max
Ave. Max.
Ave.• Min.

and average

dally minimum

(U.S.

Dept. Commerce

1975, 1976).

Ave.
nev.

June
Ave. Max.
Ave. Min.

Ave.
Dev.

temperatures

JulX
Ave. Max.
Ave. Min.

(oF) for April through

Ave.
nev.

August
Ave. Max.
Ave. Min.

Ave.
Dev.

Seetember
Ave. Max.
Ave. Min.

!.ill.
46.Z
-1. Z

Southeastern

50.5
-3.4

68.5

South
central

40.Z

Northwestern

66.1
.0.5

70.6

56.4
-0.9

60.7
31.6

North
central

4Z.Z

8Z.1
50.1

76.6

+Z.6

89.8
56.7

89.4
53.7

61. 5
-0.7

78.3

+0.9

65.6

8Z.4

.Z.O

48.8

n.8

4Z.9

-0.3

90.0
55.1

77.8
-0.3

93.9
61.7

78.0
+1. 9

95.5
60.5

75.9
39.7

65.3
+1. 8

81. 5
49.0

6Z.3

79.Z

+0.6

45.3

54.3
-0.6

70.8
37.8

n.9

57.8
-0.3

63.9

56.7

6Z.8

8Z.5

55.Z

-Z.O

f.7.3
+1. 4

.85.Z

34.Z

73.5
39.9

49.3

-1. 4

43.1

-0.5

74.1
36.3

90. 1
57.4

70.3
-1. 0

87.3
53.0

6Z.Z

95.9
6Z.1

76.1
+0.0

93.6
58.6

66.4
-0.7

8Z.3

60.3
-1.4

76.Z

55.5
+0.6

71.4
39.5

64.4

8Z.8

57.8

+O.Z

46.0

+Z.l

75.4
40.0

79.9
44.4

n.6

3Z.5

6Z.Z
-1. Z
49.6
-1. 0

66.3

-1. 7

56.5
Z3.7

38.5
-3.4

5Z.3
Z4.7

49.Z
-Z.l

!.ill.
49.4

North
central

+Z.O
55.Z

Southeastern

+1. 3

South
central

43. Z

Northwestern

43.5
+1.6

!J

Y

+1. 3

Deviation
Stations

64.6
34.1

n.1
38.Z
59.3
Z7.1
58.8

Z8.Z

from normal
in respecti

average

ve regions

58.1
+0.8

73.8
43.0

66.0
-0.6

8Z.6
49.4

73.8
+0.5

6Z.0

78.0
45.9

n.z

90.8
53.5

79.0
+0.9

-1. 4
51. 8

59.1
+1.0

68.0
35.6

+1. Z

59.4
+0.7

69.0
36.0

5Z.5
+1. Z

temperature
are:

+0.5

for month,

1) Greeley

UNC;

obtained

77.5
40.6
77.4
41.4

66.1

83.Z

+Z.6

49.0

69.5
+3.6

88.7
50.3

from previous years.
3) Monte Vista NWR; 4) Hayden.

Z) Las Animas;

44.4

-0.3

76.7
45.7
51. 5

...
I

I-'

N

I

�October

-113JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Migratory

W-88-R-22

Work Plan No.

6

Job No.

Bird Investigations

1
----------~----------------

Job Title

Migratory

Period Covered:

April 1, 1976 through March 31, 1977

Personnel:

1977

Bird Publications

William J. Adrian, Clait E. Braun, John F. Corey, Howard D. Funk,
Richard M. Hopper, Bruce R. Johnson, Robert E. Keiss, N. J.
Kitzmiller, O. W. Olsen, Ronald A. Ryder, R. M. Stabler, Michael
R. Szymczak.

ABSTRACT
Publications
as follows:

planned for and accomplished

under this job for Segment

22 are

Braun, C. E. 1976. Methods for locating~ trapping and banding band-tailed
pigeons in Colorado.
Colo. Div. Wildl. Spec. Rept. No. 39. 20 pp.
Braun, C. E., W. J. Adrian, and R. E. Keiss.
1977. Mercury residues
Colorado band-tailed pigeons.
J. Wildl. Manage. 41(1):131-134.

in

Hopper, R. M., H. D. Funk, and D. R. Anderson.
Age specificity in mallards
banded post-season in eastern Colorado.
Submitted to J. Wildl. Manage.
in 1976.
1977. Breeding densities and migration
Johnson, B. R., and R. A. Ryder.
Wilson Bull. 89(1):116-121.
periods of common snipe in Colorado.
Olsen, O. W., and C. E. Braun.
1976. New species of Splenidofilaria and
Chandlerella (Filarioidea:Nematoda),
with keys to the species from
the band-tailed pigeon (Columba fasciata fasciata) in the Rocky Mountain
region.
Great Basin Nat. 36(4):Pagination not available due to printing
delays.
1977. Blood parasites
Stabler, R. M., N. J. Kitzmiller, and C. E. Braun.
4l(1):128-l30.
from band-tailed pigeons.
J. Wildl. Manage.
Szymczak, M. R. 1975. Canada goose restoration along the foothills
Colo. Div. of Wildl. Tech. Pub. 31. 64 p.
Colorado.

of

�-114-

Szymczak, M. R. Steel shot use on a goose hunting area in Colorado.
Manuscript completed and is being finalized for submission to The
Wildlife Society Bulletin in 1977.
Szymczak, M. R., and J. F. Corey.
plains duck trap in Colorado.
6. 13 p.

1976.
Construction and use of the salt
Colo. Div. of Wildl. Div. Report No.

Szymczak, M. R., and W. J. Adrian.
southeast Colorado.
Submitted
in 1977.

Lead poisoning
to The Journal

Prepared

by

in Canada geese in
of Wildl. Manage.

k~~

Howard D. Funk
Section Chief
Small Game Research

~.

�-115October 1977
JOB PROGRESS REPORT

State of

COLORADO

------------------------------

Proj ect No.

W-88-R-22

Migratory

Work Plan No.
Job Title
Period

Investigation

Covered:

Personnel:

Joh No.

Bird Investigations
1

----------~=--------------------

of American Coots in Colorado

August 1, 1976 to March 31, 1977

W. P. Gorenzel, and R. A. Ryder, Colorado State University,
C. E. Braun and H. D. Funk, Colorado Division of Wildlife.

ABSTRACT
Investigations concerning distribution, breeding denSities, reproductive
success, habitat utilization, timing of migration, and level of hunting
mortality of coots (Fulica americana) in Colorado were initiated in August
1976. Three study areas were selected, one in the eastern plains, and two
in high mountain valleys.
Coots were distributed throughout most of Colorado,
breeding in most areas and resident year round in low numbers along the
western boundary of the plains.
Preliminary data indicated fall migration
extends from late summer to mid-November.
Spring migration began in March.
Review of harvest data indicated coots are of minor importance in Colorado

�-116-

RECOMMENDATIONS

1.

One additional study area should be selected
the Continental Divide.

2.

Migration

3.

Vegetative descriptions and measurement
be undertaken on all study areas.

4.

Data concerning

5.

Conduct additional
ence in Colorado.

6.

Methodology needs to be developed
hunter's bag.

counts should be conducted

breeding

densities

investigations

to represent areas west of

on all study areas.
of site characteristics

and productivity

must be collected.

of selected habitats

to ascertain

should

for coot occur-

age of coots in the

�-117-

INVESTIGATION

OF AMERICAN

COOTS IN COLORADO

Warner P. Gorenzel

The American coot (Fulica americana Gmelin) is a member of the order
Gruiformes, family Rallidae.
Two subspecies occur in the United States.
Fulica americana americana is widespread in North America, nesting locally
throughout much of continent and wintering mainly along the Gulf coast and
in Mexico and California (Ryder 1963).
The Hawaiian coot (!. a. alai Peale)
is restricted to the Hawaiian Islands.

,•

Current state and federal regulations list the coot, hereafter used in
reference to F. a. americana, as a migratory game bird. As such it is classed
with ducks and geese as "waterfowl".
In certain areas of the United States
coots can constitute a large percentage of the waterfowl harvest.
Kiel and
Hawkins (1953), Burton (1959), and Jahn and Hunt (1964) have indicated the
significance of the coot harvest in the Mississippi Flyway.
Nationwide,
overall hunting interest in coots is low (Fredrickson 1977).
In Colorado, Bailey and Niedrach (1965) list the coot as a common nesting
species, but only occasionally as a wintering species.
They also summarize
general information on nest observations, early and late occurrences, and
distribution.
No previous investigations on the status of coots in Colorado have been
conducted.
Consequently, little is know about habitat preferences, timing
of habitat use, productivity, and hunter utilization.
In order to improve the
knowledge and management of coots in Colorado this study was initiated in
1976.
P.N.O. OBJECTIVES
The major objective of this study is to obtain data necessary for the development of a management plan for coots breeding in and-migrating through
Colorado.
Data considered necessary are those relating to distribution,
breeding densities and production in the eastern plains, high mountain
valleys and west of the Continental Divide, descriptions of habitats utilized
according to vegetation, water characteristics, documentation of timing of
spring and fall migration, and present levels of reported harvest.
Hypotheses that have been developed are:
1.

Distribution of coots in Colorado is determined by the occurrence of
cattail (Typha spp.) and/or bulrush (Scirpus spp.) dominated marshes.

2.

Densities of breeding coots, nesting success and productivity differ
significantly between occupied habitats in the eastern plains, high
mountain valleys, and west of the Continental Divide.

3.

Timing of spring and fall migrations
east of the mountains, high mountain
Divide.

differs significantly between areas
valleys, and west of the Continental

�-118-

4.

Hunting in Colorado is the most important source of mortality for locally
produced coots.
SEGMENT OBJECTIVES

1.

Review literature concerning records of coot occurrence, distribution,
nesting and harvest in Colorado.

2.

Circulate questionnaires to field personnel of the Division of Wildlife,
selected University and privately employed wildlife scientists and
amateur ornithologists concerning occurrences, distribution, nesting
and harvest in Colorado.

3.

Investigate selected habitats for coot occurrence in all major topographic regions in Colorado (i.e, eastern plains, inter-mountain valleys,
and west of the Continental Divide).

4.

Compile data, analyze results and prepare progress report.
REVIEW OF LITERATURE

The general life history of the coot has been described by Bent (1926),
Forbush (1912), and Fredrickson (1977). Records on distribution have been
noted by Armistead (1970), Bailey (1928), Burleigh (1972), Jones (1940), and
within Colorado by Bailey and Niedrach (1965) and Sclater (1912).
Numerous studies have been conducted on the breeding biology and reproductive
success of the coot. Major studies have centered in four areas, California
(Anderson 1956, 1957; Gullion 1954, 1956; Hunt and Naylor 1955; Miller and
Collins 1954), Utah (Ryder 1958, 1961), Iowa (Crawford 1975; Fredrickson
1967, 1969a, 1970; Friley et ale 1938; Provost 1947; Weller 1971), and
Wisconsin (Jahn and Hunt 1964). Kiel (1955) examined reproduction in
Manitoba and Vaa et ale (1974) in South Dakota.
Various aspects of territorial and intraspecific behavior have been documented
by Burger (1973), Gullion (1951, 1952c, 1953b), Joyner (1973), McNicholl
(1975), Munro (1939), Ryder (1957, 1958, 1959), Sooter (1945), and Wick and
Penstila (1957).
The effects of changing habitat conditions on coot numbers and production
have been studied by Harris and Marshall (1957), Hendrickson (1936),
Johnsgard (1956), Smith (1961), Weller (1975), Weller et ale (1958), and
Wolf (1955).
Food habits have been studied by Eley (1975), Eley and Harris (1976), Jones
(1940), and Stollberg (1949).
Sex and age determination in relation to physical measurements is discussed
by Boss (1973), and Crawford (1975), Fredrickson (1968), and Gullion (1950,
1952a, 1954). Gullion (1953a) also described the molt of the coot.

�--~ --~
IOU

HIJ

l-L:-.------ ..

&lt;I

I·

·

I•
I

i

r
i

40

-..

--~---\
101

,-----

w.
'--_._e..-,

106
~Q_C!l_.

104

----n"

~._a_.~ . .-.
103

-~\

r

W

~

4Q

I

,

r--

-

'I

~

~-

,

e

I•

v-~

I

r

Jli

I,
I•

I

\

,.-

f\

• \8£&lt;

~

"h

'--~

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�-120-

Disease,
(1952b),
Trautman

parasites, and other mortality factors have been examined by Gullion
Rausch (1947), Roudabush (1942), Trainer and Fischer (1963), and
et ale (1939).

Migration and other movements of coots are discussed by Alley and Boyd (1949),
Burton (1959), Fredrickson
(1969b), Heit (1948), Hochbaum (1955), Jung (1924),
Prill (1931), and Ryder (1963).
Dispersal of members of the Rallidae
in
general was discussed by Ripley (1976).
Harvest data for coots in the United States are presented by Boeker (1953),
Fredrickson
(1977), and the Pacific Flyway Council (1976), and in Canada by
Cooch (1976). Aspects of population dynamics and status have been analyzed
by Burton (1959), Kiel and Hawkins (1953), and Ryder (1963).
METHODS

AND MATERIALS

Three methods
the selection

were used to investigate
of study areas:

1.

Extensive

field trips as part of planned

2.

Field

3.

Questionnaires
sent to field personnel
and amateur ornithologists
(Appendix).

trips in conjunction

coot occurrence

with graduate

in Colorado

and aid in

research.
courses.

of the Colorado

Division

of Wildlife

Responses from the questionnaire,
personal observations,
and results from the
Study of Colorado Bird Distribution
(Kingery 1976) were combined to plot coot
distribution and status according to the latilong system.
In this system, the
state was divided into distinct areas based on latitude and longitude (Fig. 1).
Coot status was then determined within each area.
Three study areas were selected

based on the following

1.

Sufficient area and cover to provide
breeding coots.

2.

Stable water

3.

Similarity

4.

Accessibility.

5.

Freedom

adequate

criterea:

numbers

(25 pairs)

of

levels.

of the area to general

from excessive

external

wetlands

in the region.

interference.

To investigate levels of hunting mortality, coot harvest data from the Central
and Pacific Flyways, and especially Colorado, were examined.
In addition,
a questionnaire
(Appendix) was designed to determine harvest levels and hunter
attitudes.
Students from the Colorado State University Wildlife Techniques
class were assigned to check stations to question hunters leaving the Wellington Wildlife Area on 9 and 10 October 1976.

�-121-

Migration counts were started in late October 1976 at the Beebe Draw study
area on a biweekly basis.
A field sheet (Appendix) was devised to aid in
the collection of data. Migration counts continued at Beebe Draw until
start of the goose season on 6 November when access was denied.
Thereafter,
a selection of reservoirs and lakes northeast and south of Fort Collins were
investigated for suitable habitat and coot occurrence.
Coot numbers were
monitored at College and Lindenmeier lakes, both in the Fort Collins area,
on a weekly basis starting mid-January 1977. With the first indications of
migrational movements, counts were again initiated at Beebe Draw starting
15 March.
RESULTS

AND DISCUSSION

Distribution

and Status

The coot is listed by Bailey and Niedrach (1965) as resident in Colorado;
common as a nester and uncommon as a wintering species.
Both Bailey and
Niedrach (1965) and Sclater (1912) note that the coot breeds mainly on the
plains, and in mountain parks up to 3,048 m. Sclater (1912) mentions that
it is not so abundant on the western slope, and that a few winter around
Denver.
Data obtained from questionnaires,
field investigations,
and published records
were compiled to determine coot distribution and status according to the latilong system (Table 1).

Table 1. Coot distribution and status in Colorado according to the latilong
system.
M = migrant only, R = resident year round, B = breeding, b.= v~ry likely
breeding, bird found in suitable habitat.
Columns are those shown ln Flg. 1.

B

B

M

B

B

R

R

B

B

B

R

R

B

B

b

R

R

M

R

B

B

B

B

B

B

R

Coots are resident year round along the eastern boundary of the Rocky Mountains
from near Fort Collins south to Pueblo.
The presence of wintering coots in
other areas, notably the southwestern corner of Colorado may be dependent on
the severity of winter weather.
Coots breed throughout most of the state; in
all cases breeding coots were associated with cattail or bulrush marshes, and
occasionally willows (Salix spp.).
Future investigations will further delimit
coot distribution and status.

�-122-

Selection

of Study Areas

Three study areas were selected, one representing the eastern plains region,
and two representing high mountain valleys.
Detailed habitat descriptions
have not yet been undertaken.
The Beebe Draw study area is in Weld County, T4N, R65W, Section 34, at an
elevation of 1,446 m. The area is a Type 5 wetland (Shaw and Fredine 1956)
of approximately 11 hectares.
Emergent vegetation consists primarily of
cattails and bulrush.
The Lake John Annex study area is in Jackson County, T9N, R8lW, Section 2,
at an elevation of 2,500 m. Total area is approximately 16 hectares.
Ocular
examination indicated the emergent vegetation was dominated by bulrush.
Ice Pond, near Buena Vista, is in Chaffee County, T13S, R78W, Sections 5, 6,
7, 8, at an elevation of 2,425 m. Ice Pond is a Type 5 wetland approximately
12 hectares in size with emergent and shoreline vegetation of mixed stands
of cattail, bulrush, and sedges (Carex spp).
Questionnaire respondents mentioned several areas west of the Continental
Divide where at least 10-15 or more breeding pairs have been observed
(Table 2). The suitability of each location as a potential study a;ea will
be investigated in the spring of 1977.
Harvest
Coot harvest data in the Central and Pacific Flyways, and especially Colorado
were examined.
Cooch (1976) reported in Canada that the kill of coots for
the period 1969-1974 showed a significant drop on a national basis.
However,
the kill in western Canada remained relatively stable, the decline being
restricted to Ontario and Quebec.
Cooch indicated the harvest of this
relatively abundant species is low in relation to numbers present.
Fredrickson (1977) reported on coot harvest for the period 1952-72.
The
majority of coots shot in the Central and Pacific Flyways were taken in
Texas and California, respectively.
A review of Boeker (1953) indicated
that few western states had listed coot data in their waterfowl bag records
prior to 1953.
The Pacific Flyway Council (1976) reported 123,577 coots
harvested in 1975-76, but this figure included data only from Nevada (270),
Washington (12,620), Oregon (70,184), Idaho (30,300), and California (10,203).
The data from California included only coots taken on state and federal
shooting areas.
In Colorado waterfowl harvest records are derived from mail surveys sent to
approximately three percent of the small game license buyers.
Colorado Small
Game Hunter Harvest surveys from 1954 through 1972 did not give specific
harvest figures for coots, but instead grouped them in the category of
"Others and Unknown".
This group included coots, mergansers, goldeneyes,
buffleheads
and unknowns.
Data from more recent small game harvest surveys
are presented in Table 3 (H. Riffel, pers. comm.).

I

!

�-123-

Table 2. Locations
questionnaires.

of major western

slope coot breeding

areas based on

Location

County

Echo Canyon Reservoir

Archuleta

Crawford

Delta

Reservoir

Sweitzer Lake

Delta

Ground Hog Reservoir

Dolores

Narriguinnep

Dolores

Sweetwater

Reservoir
Lake

Garfield

Beaver Lake

Gunnison

Lilly Lake

Gunnison

Lake San Cristobal

Hinsdale

Chapman Lake

La Plata

Harper Pond

La Plata

Highline

Mesa

Reservoir

Denny Lake

Montezuma

Totten Reservoir

Montezuma

Table 3.

Year

Coot harvest

in Colorado

1973-75.

Estimated

Harvest

Percent

of Duck Harvest

1973

4,579

1.50

1974

5,054

1.41

1975

2,924

1.16

Fredrickson

(1977) estimated

the 1971-72 Colorado

coot harvest

to be 6,700.

�-124-

Despite liberal daily bag limits of 15 and 25 in the Central and Pacific
Flyways, respectively, coot harvest is low in Colorado.
Out of 67 questionnaires returned by Division of Wildlife personnel, only seven indicated
having encountered any hunters who retained coots in the bag.
Of 54 hunters polled by students at the Wellington Wildlife area in 1976
only 11 indicated they normally hunt coots.
The majority of non-coot
hunters cited bad taste and unsporting behavior as reasons for not hunting
coots.
Other hunters either preferred ducks or were unaware that coots were
legal game birds.
Seven hunters harvested 28 coots.
Timing of Migration
Bailey and Niedrach (1965) state that coots band into large flocks in late
September and that the majority have migrated south by the last of October.
Observations in August 1976 (Table 4) indicated that coots had already
begun to gather in large flocks.

Table 4.

Locations

and approximate

numbers

of coots observed,

August

1976.

Location

County

San Luis Lake

Alamosa

250

Cucharas

Huerfano

200

Jackson

150

Jackson

600

MacFarlane

Reservoir
Reservoir

Walden Reservoir

Number

Observed

Nesting cover in the above mentioned areas was either absent or too sparse
to support such large breeding populations.
Migration counts were started too late in the fall of 1976 to determine peak
periods.
However, 2,000 to 2,500 coots were still present at Lake John
Annex by 15 October.
Counts at Beebe Draw indicated heavy but declining use
of the area through October into early November.
When access to Beebe Draw
was denied, investigations of lakes or reservoirs in the Fort Collins area
indicated coots were absent from most areas by mid-November.
Sclater (1912)
states that coots arrive from the south during the second half of March.
Counts at Beebe Draw indicated coot numbers had increased from one on 1 March
to over 100 by 29 March 1977.

�-125-

LITERATURE CITED

Alley, R., and H. Boyd. 1949. Effects of the cold spell of 1947 on the
coot in North Somerset. Brit. Birds 42(7):225-232.
Anderson, W. 1956. A waterfowl nesting study on the grasslands, Merced
County, California. Calif. Fish and Game 42(2):117-130.
1957. A waterfowl nesting study in the Sacramento Valley, California,
1955. Calif. Fish and Game 43(1):71-90.
Armistead, H. T. 1970. First Maryland breeding of American coot at Deal
Island. Maryland Birdlife 26(3):79-81.

I

Bailey, A. M., and R. J. Niedrach.
Nat. Hist. Vol. I. 454 pp.

I

Bailey, F. M.
807 pp.

t
t

1928.

1965.

Birds of New Mexico.

Birds of Colorado.

Denver Mus.

New Mexico Dept. Game and Fish.

Bent, A, C, 1926. Life histories of North American marsh birds.
Nat. Mus. Bull. 135:358-371.

U. S.

Boeker, H. M. 1953. A summary of North American waterfowl hunting statistics.
Colorado Coop. Wildl. Res. Unit Qtr. Rept. 6(3):14-62.
Boss, A. S. 1963. Aging the nests and young of the American coot.
Thesis. Univ. Minnesota, St. Paul. 62 pp.

M. S.

Burger, J. 1973. Competition between American coots and Franklin's gulls
for nest sites and egg predation by the coots. Wilson Bull. 85(4):
449-451.
Burleigh, T. D. 1972.
Idaho. 467 pp.

Birds of Idaho.

The Caton

Printers, Ltd.

Caldwell,

Burton, J. H., II. 1959. Some population mechanics of the American coot.,
J. Wildl. Manage. 23(2):203-210.
Cooch, F. G. 1976. Kill of migratory game birds other than waterfowl by
hunters in Canada 1975. Can. Wildl. Servo Prog. Notes 68. 7 pp.
Crawford, R. D. 1975. Breeding biology of American coots in relation to
age. Ph. D. Thesis. Iowa State Univ., Ames. 42 pp.
Eley, T. J., Jr. 1975. Winter ecology of the American coot along the lower
Colorado River. M. S. Thesis. Humboldt State Univ., Arcata, Calif.
45 pp.

----- , and S. W. Harris.

1976. Fall and winter foods of American coots
along the lower Colorado River. Calif. Fish and Game 62(3):225-227.

�-126-

Forbush, F. M. 1912. A history of the game birds, wild-fowl and shorebirds of Massachusetts and adjacent states. Massachusetts Bd. of
Agric., Boston. 622 pp.
Fredrickson, L. H. 1967. Some aspects of reproductive behavior of American
coots (Fulica americana). Ph. D. Thesis. Iowa State Univ., Ames.
no pp.
1968. Measurements of coots related to sex and age.
Manage. 32(2):409-411.

J. Wildl.

1969a. An experimental study of the clutch size of the American coot.
Auk 86(3):541-550.
1969b. Mortality of coots during severe spring weather.
Bull. 81(4):450-453.
1970. Breeding biology of American coots in Iowa.
82(4):445-457.

Wilson

I

Wilson Bull.

I

(Chairman). 1977. Coots. In Sanderson, G. C. (ed.). Management
of migratory shore and upland game birds in North America. Int. Assoc.
Game, Fish and Cons. Commsi, Washington, D. C. In Press.
Friley, C. E., L. J. Bennett, and G. O. Hendrickson.
coot in Iowa. Wilson Bull. 50(2):81-86.

1938.

The American

Gullion, G. W. 1950. Voice difference between sexes in the American coot.
Condor 52(6):272-273.
1951.
157-166.

The frontal shield of the American coot.

Wilson Bull. 63(3):

1952a. Sex and age detennination in the American coot.
Manage. 16(2):191-197.

J. Wildl.

1952b. Some diseases and parasites of American coots.
and Game 38(3):421-423.

Calif. Fish

1952c. The displays and calls of the American coot.
64(2):83-97.

Wilson Bull.

1953a.
102-103.

Observations on molting of the American coot.

Condor 55(2):

1953b.
169-186.

Territorial behavior of the-:.American
coot.

Condor 55(4):

1954. The reproductive cycle of American coots in California.
Auk 71(4):366-412.
1956. An observation concerning the validity of coot brood counts.
J. Wildl. Manage. 20(4):465-466.

t

�-127-

Harris, S. W., and W. H. Marshall. 1957. Some effects of a severe windstorm
on coot nests. J. Wildl. Manage. 21(4):471-473.
Heit, W. S. 1948. Texas coastal waterfowl concentration areas and their
1947-48 wintering population. Trans. N. Am. Wildl. Conf. 13:323-338.
Hendrickson, G. o. 1936. Observations on nests and young of the coot.
Wilson Bull. 48(3):216-218.
Hochbaum, H. A. 1955. Travels and traditions of waterfowl.
Minnesota Press, Minneapolis. 301 pp.

Univ. of

Hunt, E. G., and A. E. Naylor. 1955. Nesting studies of ducks and coots in
Honey Lake Valley. Calif. Fish and Game 41(4):295-314.
Jahn, L. R., and R. Hunt. 1964. Duck and coot ecology and management in
Wisconsin. Wisc. Cons. Dept. Tech. Bull. 33. 212 pp.
Johnsgard, P. A. 1956. Effects of water fluctuation and vegetation change
on bird populations, particularly waterfowl. Ecology 37(4):689-701.
Jones, J. C.
bution.

1940. Food habits of the American coot with notes on distriU. S. Fish and Wildl. Serv., Wildl. Res. Bull. 2. 52 pp.

Joyner, D. E. 1973. Interspecific nest parasitism by ducks and coots in
Utah. Auk 90(3):692-693.
Jung, C. S.
29-30.

1924.

Actions of coots during freeze-up.

Wilson Bull. 36(1):

Kiel, W. H. 1955. Nesting studies of the coot in southwestern Manitoba.
J. Wildl. Manage. 19(2):189-198.

----- , and A. S. Hawkins.
Flyway.

1953. Status of the coot in the Mississippi
Trans. N. Am. Wildl. Conf. 18:311-322.

Kingery, H. E., ed. 1976. Study of Colorado bird distribution.
Wildl. Unpubl. Rept. 42 pp.

Colo. Div.

McNicholl, M. K. 1975. Interactions between Forster's terns and American
coots. Wilson Bull. 87(1):109-110.
Miller, A. W., and B. D. Collins. 1954. A study of ducks and coots on
Tule Lake and Lower Klamath National Wildlife Refuges. Calif. Fish and
Game 40(1):17-37.
Munro, J. A. 1939. The relations of loons, Holboell's grebes, and coots
to duck populations. J. Wildl. Manage. 3(4):339-344.
Pacific Flyway Council. 1976.
90 pp.
Unpubl. Rept.

Pacific waterfowl flyway report No. 75.

�-128-

Prill, A. G.

1931.

A land migration of coots.

Wilson Bull. 43(2):148-149.

Provost, M. W. 1947. Nesting of birds in the marshes of northwest Iowa.
Amer. MidI. Nat. 38(2):485-503.
Rausch, R.
189.

1947.

Ripley, S. D.

Pullorum disease in the coot.

1976.

Rails of the world.

J. Wildl. Manage. 11(2):

Amer. Scientist 64(6):628-635.

Roudabush, R. L. 1942. Parasites of the American coot (Fulica americana)
in central Iowa. Iowa State ColI. Journ. Sci. 16(4):437-441.
Ryder, R. A. 1957. Coot-waterfowl relations on some northern Utah marshes.
Proc. Utah Acad. Sci. 34:65-68.
1958. Coot-waterfowl relationships in northern Utah.
Utah State Univ., Logan. 219 pp.

Ph. D. Thesis.

1959. Interspecific intolerance of the American coot in Utah.
Auk 76(4):424-442.
1961. Coot and duck productivity in northern Utah.
Wildl. Nat. Res. Conf. 26:134-137.

Trans. N. Am.

1963. Migration and population dynamics of American coots in
western North America. Proc. Internatl. Ornith. Congr. 13:441-453.
Sclater, W. L. 1912.
London. 567 pp.

A history of the birds of Colorado.

Witherby and Co.,

Shaw, S. P., and C. G. Fredine. 1956. Wetlands of the United States.
Fish and Wildl. Servo Circ. 39. 67 pp.

U. S.

Smith, A. G. 1961. Some ecological factors and censusing problems associated with coots in the parklands of Alberta, Canada. U. S. Fish and
Wildlife Serv., Wildl. Res. Lab. Denver. 46 pp.
Sooter, C. A. 1945. Relations of the American coot with other waterfowl.
J. Wildl. Manage. 9(2):96-99.
Stollberg, B. P. 1949. Competition of American coots and shoal-water ducks
for food. J. Wildl. Manage. 13(4):423-424.
Trainer, D. 0:, and G. W. Fischer. 1963.
J. Wildl. Manage. 27(3):483-486.

Fatal trematodiasis of coots.

Trautman, M. B., W. E. Bills, and E. L. Wickliff. 1939. Winter losses
from starvation and exposure of waterfowl and upland game birds in
Ohio and other northern states. Wilson Bull. 52(2):86-104.

�-129-

Vaa, S. J., K. L. Cool, and R. L. Linder. 1974. Nesting by American coots
in South Dakota. Proc. South Dakota Acad. Sci. 53:153-156.
Weller, M. W. 1971. Experimental parasitism of American coot nests.
Auk 88 (1):108-115.
1975. Studies of cattail in relation to management for marsh wildlife.
Iowa State J. Res. 49(4):383-412.
-----, B. H. Wingfield, and J. B. Low. 1958. Effects of habitat deterioration on bird populations of a small Utah marsh. Condor 60(4):
220-226.
Wick, W. Q., and H. E. Penstila. 1957. Observations on a coot-muskrat
relationship. Condor 59(5):341-342.
Wolf, K. 1955. Some effects of fluctuating and falling water levels on
waterfowl production. J. Wildl. Manage. 19(1):13-23.

Prepared bY__ ~~~~~~·z~~_~~~
__ ~~~
_
Warner P. Gorenzel
Graduate Research Assistant

Approved by__

--=a~u:..::...:..., _2._.--J.....:~=--Clait E. Braun
Wildlife Researcher

_

�-130-

APPENDIX

�-l31-

SURVEY OF COOTS IN COLORADO

1. District

-------------------------------------------------------------------

2.

To your knowledge, do coots occur in your present district? Yes
If no, please answer number 9 only and return survey form.

No

3.

If coots are known to occur in your district, please list major locations
where they have been observed

-----------------------------------------------

4.

Please list wetlands where you have observed at least 10-15 or more breeding
pairs of coots
~--------------------------------

5.

Which wetland vegetation types occur where you see coots during June and
July (i.e. cattails, bulrush, willow, etc.?}
_

6.

Which month(s) of the year are coots most commonly present in your district?

7.

Have you encountered coots in your district during 1 December through
31 March? Yes
No.
If yes, where?

---------------------------------

8.

Have you encountered any hunters who retain coots -in the bag in your
district? Yes
No
If yes, where?

-----------------------------------

9.

List any major we t Lands in your district where you do not normally observe
coots
_

�-132-

Surveyor Coots in Colorado

1.

County (or counties)

2.

'Ib your knowledge,do coots occur in the area you listed above?
Yes

_

No

----

rr no, please answer number8 only and return survey rorm.

3.

Do coots breed in your area?
rr yes, please list

Yes

No

_

wetlands where you have observed at least

10-15 or more breeding pairs of coots.

Please be speciric.

4.

Please list

5.

Whichwetland vegetation types occur where you see coots during
June and July (Le. cattails, bulrush, willow, etc.)

6.

Whichmonth(s) of the year are coots mst conmonlypresent in your
area?

other major areas where coots have been observed.

----------------------------------------------

7.

Are coots present in your area during most or all of the 1 December
through 31 Marchperiod? Yes
No
IT yes, where?

8.

List any major wetlands in your area where you do not normally observe
coots.

----------------------------------------------

�Location
Date

Check Station #
T~e

Stqrt

Name of Observer

Time Stop

-----------------ASK ALL HUNTERS LEAVING WITHOUT COOTS:

ASK ALL HUNTERS LEAVING WITH COOTS:
Do you
normally
hunt coots?

If no,
why!
Comments

------------------------------

/I Hit
II
Bagged

Not
Retrieved

Hrs.
Spent
Hunting

Do you
ever hunt
coots?

If no,
why!
Comments

Hrs.
Spent
Hunting

I
I-'

W
W
I

-----

i

�-l34-

Observer

Date

Location

Time

Estimated Cloud Cover

%

Wind:
Precip:

None

med

strong

gusty

rain

mist

snow

fog

intermit

Hrs.
FOor CO

Temp

light

None

to

direction

steady

Number Observed:
Coots

Adults

Ducks:

Imma

by species

Mal

Pin

G-W-T

B-W-T

Gad

Wid

Shov

Geese

Red

Scaup

Other

TOTAL Observed
Other Species: (i.e., grebes, shore birds)

Other Conditions or Changes:

_

Comments:

_

10/76

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                  <text>-1-

January 1978
JOB PROGRESS REPORT

COLORADO

State of

Bighorn Sheep andMt.

Goat Investigations

Project No.

W-41-R-27

Work Plan No.

:. Job No.
------------------------------------------------------Evaluation of Trace Mineral Availability to Bighorn Sheep

16

1

Job Title

~

Period Covered:

June 1, 1976 to May 31, 1977

Personnel:

~o~n~W~i~n~t~e~r~R~a~n~g~e~s~i~n~C~o~l~oLr~a~d~o~

_

Robert E. Keiss, Bruce Goforth, Willie Travnicek, Robert Schmidt,
Paul Neil.

ABSTRACT

Trace mineral formulations were developed and tested for-animal preference.
As a result of these tests a trace mineral mix for bighorn sheep was developed
and is in use throughout many areas in Colorado as a supplement to the
nutritional requirements of animals for essential trace minerals.
An attempt
was made to measure the physiological response of bighorn sheep to trace
mineral supplementation by analytical testing of fecal material collected
from study areas where animals were giv~n trace mineral free choice.

��-3-

EVALUATION OF TRACE MINERAL AVAILABILITY
TO BIGHORN SHEEP ON WINTER RANGES IN COLORADO
Robert E. Keiss
P.N.O. OBJECTIVE
The objective of this study is primarily directed towards evaluating certain
nutritional supplements on bighorn sheep herds in an effort to improve lamb
survival and viability of bighorn sheep to bring about an overall increase
in the total numbers of animals within the State of Colorado.
SEGMENT OBJECTIVES
1.

2.
3.
4.

Formulate a trace mineral mix which can be placed on bighorn sheep
winter ranges to supplement the nutritional trace mineral requirements
of bighorn sheep.
Measure the amounts of trace mineral mix consumed by bighorn sheep and
determine seasonal preferences.
Measure trace mineral content of bighorn sheep fecal material and relate
fecal mineral levels to potential trace mineral deficiencies.
Deternline the effects of trace mineral mix consumption on the physiology
of bighorn sheep.
PROCEDURES

Experimental trace mineral mixes were developed and tested on selected bighorn
sheep ranges in Colorado to measure animal preference for specific formulations.
The basic ingredients were purchased from Feed Products, Inc.,
1000 West 47th Avenue, Denver, Colorado and included the following:
1.
2.

3.
4.

SP-4 Feed Supplement - manufactured by FMC Corporation, 2000 Market
Street, Philadelphia, Penn.
(This product is 100% sodium phosphate).
Ruminant Trace Mineral Premix - manufactured by Feed Products, Inc.,
Denver, Colorado.
(This product contains 4.40% manganese, .30% iodine,
.20% cobalt, 6.60% iron, 1.30% copper, 12.00% zinc, and 20% magnesium.
The following ingredients are utilized to give the above listed element
concentrations: Manganese oxide, calcium iodate, calcium stearate, cobalt
carbonate, iron oxide, copper oxide, magnesium oxide, iron carbonate, zinc
oxide, and calcium carbonate).
The oxide and carbonate compounds are
water insoluble, whereas, the sulfates and chlorides are water soluble.
As a general rule the water insoluble compounds are preferred for supplements because they are less likely to be leached out of the mix by moisture from weather elements.
Sodium chloride - Feed grade.
Sodium bentonite - "Vo1c1ay", a product of American Colloid Company,
5100 Suffield Ct., Skokie, Illinois 60076.

Four different foru1ations were selected for testing on bighorn sheep ranges
and the percent ingredients are listed in Table 1. The ingredients were
weighed and mixed in a portable concrete mixer.
Each mix was tumbled for

�-4-

at least 30 minutes

to assume a uniform mixture.

Boxes to hold the trace mineral mixes were constructed so that there were
four separate compartments.
These boxes were placed on the bighorn sheep
ranges and a different formulation was placed in each compartment.
The
animals could then select, by free choice, the formulations they preferred.
The mixtures were put out on bighorn sheep ranges in the Medano Creek,
Tarryall Mountains, Trickle Mountain, South Platte River Canyon, Collegiate
Range (Love Ranch), Collegiate Range (Jump Steady), Mount Evans (Grant),
and Poudre River.
The boxes were put out in January 1976 with 11.36 kg
(25 lb) of each mixture added to the boxes.
Each study area was checked
every other month (March 1976, May 1976, July 1976, September 1976, and
November 1976). The amount of trace mineral mix remaining in the box was
weighed and returned to the box. (A 2-3 g sample was taken of each mix,
placed in a closed vial, and returned to the laboratory for moisture
analysis.
All weights were then corrected to a dry weight basis for
comparative purposes.)
Tables 2, 3, 4, 5, 6, 7, 8, and 9 show the results of use preferences,
by study area, on the amounts of trace mineral consumed by animals on
the bighorn sheep areas.
Table 10 summarizes the total amount of trace mineral consumed by animals
on the study areas by trace mineral mix formulations.
These data would
indicate a preference for the TMIII and TMIV formulations.
Table 11 summarizes the total amount of trace mineral consumed by animals
on the study areas by season of the year. There is an indication that the
total consumption increases during the spring and early summer periods.

Table 1. Trace mineral mix formulations
bighorn sheep winter ranges.

tested for animal preference

TMI

TMII

TMIII

TMIV

Sodium chloride

10%

30%

50%

70%

Bentonite

65%

45%

25%

5%

Trace mineral mix

5%

5%

5%

5%

Phosphorus

20%

20%

20%

20%

(XP-4)

on

�-5-

Table 2. Kilograms of trace mineral mix formulations
range in the Tarrya11 Mountains study area.

TMl

January

1976

TMII

used on bighorn

TMIll

TMlV

Start of test period - 11.36 kg of each TM used.

March 1976
Kg TM used

..29

3.87

4.62

3.79

% of total TM used

2.31

30.79

36.75

30.15

Kg TM used

.17

4.44

6.82

5.27

% of total TM used

1.02

26.59

40.84

31. 56

Kg TM used

.06

3.29

8.86

6.93

% of total TM used

.31

17.19

46.29

36.21

Kg TM used

.05

2.28

3.87

2.95

% of total TM used

.55

24.92

42.30

32.24

Kg TM used

.05

1.01

1.95

1.06

% of total TM used

1.23

24.82

47.91

26.04

Total kg - test period

.62

14.89

26.12

20.00

% of total TM used

1.01

24.16

42.38

32.45

May 1976

July 1976

September

November

sheep

1976

1976

�-6-

Table 3.

Kilograms

of trace mineral

mix formulations

range in the Trickle Mountain

study area.

TMl

TMIl

January
March

1976

used on bighorn

TMIll

sheep

TMlV

Start of test period - 11.36 kg of each TM used.

1976

Kg TM used

.03

.37

.16

.33

% of total TM used

3.37

41.57

17.98

37.08

Kg TM used

.10

1. 76

2.38

1.86

% of total TM used

1.71

30.14

40.61

31.85

Kg TM used

.05

3.75

5.72

4.89

% of total TM used

.35

26.02

39.69

33.93

Kg TM used

.01

.46

1.29

.97

% of total TM used

.37

16.85

47.25

35.53

.05

.07

.19

.l3

11.36

15.91

43.18

29.55

Total kg - test period

.19

6.41

9.74

8.18

% of total TM used

.77

26.14

39.72

33.36

May 1976

July 1976

Se:etember 1976

November

1976

Kg TM used

% of total TM used

�-7-

Table 4.
ranges

Kilograms

of trace mineral

in the South Platte River Canyon

TMI

January
March

mix formulations

1976

used on bighorn

sheep

study area.

TMII

Start of test period

TMIII

TMIV

- 11.36 kg of each TM used.

1976

Kg TM used

.32

2.38

1.65

3.17

% of total TM used

4.26

31.65

21.94

42.15

Kg TM used

.15

3.89

4.79

2.17

% of total TM used

1.36

35.36

43.55

19.73

Kg TM used

..05

3.19

4.17

3.86

% of total TM used

.44

28.31

37.00

34.25

Kg TM used

.08

1. 38

3.77

2.90

% Of total TM used

.98

16.97

46.37

35.67

Kg TM used

.11

.86

1.58

1.03

% of total TM used

3.07

24.02

44.13

28.77

Total kg - test period

.71

12.12

15.96

13.13

% of total TM used

1.69

28.91

38.07

31.32

May 1976

July 1976

SeEtember

November

1976

1976

�-8-

Table 5.

Kilograms of trace mineral mix formulations used on bighorn sheep

ranges in the Collegiate Range (Love Ranch) study area.

TMI
January 1976

TMll

TMIll

TMIV

Start of test period - 11.36 kg of each TM used.

March 1976
.10
1.08

2.33

3.87

2.98

25.11

41.70

32.11

Kg TM used

.05

3.68

5.91

6.03

% of total TM used

.32

23.48

37.72

38.48

Kg TM used

2.70

5.32

1.53

1.59

% of total TM used

24.24

47.76

13.73

14.27

Kg TM used

.10

3.21

4.63

4.22

% of total TM used

.82

26.46

38.08

34.70

Kg TM used

.05

2.08

1.72

3.61

% of total TM used

.67

27.88

23.06

48.39

Total kg - test period

3.00

16.62

17.66

21.27

% of total TM used

5.12

28.39

30.16

36.33

Kg TM used
% of total TM used
May 1976

July 1976

Se:etember 1976

November 1976

�-9-

Table 6.

Kilograms

of trace mineral mix formulations

ranges in the Collegiate

Range

TMl

January

1976

used on bighorn

(Jump Steady) study area.

TMIl

TMIll

TMlV

Start of test period - 11.36 kg of each TM used.

March 1976
Kg TM used

.02

.76

.93

1.33

% of total TM used

.66

25.00

30.59

43.75

Kg TM used

.05

1.37

2.81

2.03

% of total TM used

.80

21.88

44.89

32.43

.12

1.87

3.62

3.21

1. 36

21.20

41.04

36.39

Kg TM used

.01

.80

2.79

1. 68

% of total TM used

.19

15.15

52.84

31. 82

Kg TM used

.03

.75

2.61

1. 85

% of total TM used

.57

14.31

49.81

35.31

Total kg - test period

.23

5.55

12.76

10.10

% of total TM used

.80

19.38

44.55

35.27

May 1976

July 1976
Kg TM used
% of total TM used
SeEtember

November

sheep

1976

1976

�-10-

Table 7.

Kilograms of trace mineral mix formulations used on bighorn sheep

ranges in the Mount Evans (Grant) study area.

TMI
January 1976

TMIl

TMIIl

TMIV

Start of test period - 11.36 kg of each TM used.

March 1976
Kg TM used

.54

2.86

3.86

2.36

% of total TM used

5.60

29.72

40.12

24.53

.18
1.60

3.27

4.36

3.41

19.14

38.86

30.39

Kg TM used

.05

.82

1.45

.68

% of total TM used

1.70

27.33

48.33

22.66

Kg TM used

.06

1.63

2.36

2.01

% of total TM used

.10

26.89

38.94

33.16

Kg TM used

.03

1.87

2.93

1.82

% of total TM used

.45

28.12

44.06

27.36

Total kg - test period

.86

10.45

14.96

10.28

% of total TM used

2.35

28.59

40.93

28.13

Ma~ 1976
Kg TM used
% of total TM used
Ju1~ 1976

Se.etember 1976

November 1976

�-11-

Table 8.

Kilograms

of trace mineral mix formulations

used on bighorn

sheep

ranges in the Poudre River study area.

TMI

January

1976

TMII

TMUI

TMIV

Start of test period - 11.36 kg of each TM used.

March 1976
Kg TM used

.37

4.17

3.62

3.81

% of total TM used

3.09

34.83

30.24

31.82

Kg TM used

.05

3.72

4.52

3.61

% of total TM used

.42

31.26

37.98

30.33

Kg TM used

.04

2.60 .

4.82

3.20

% of total TM used

.37

24.39

42.21

30.01

Kg TM used

.01

1.61

1.82

1.32

% of total TM used

.21

33.82

25.21

27.73

Kg TM used

.05

.27

1.06

1.41

% of total TM. used

1. 82

9.85

38.68

51.45

Total kg - test period

.52

12.37

15.84

13.35

1.24

29.40

37.64

31. 72

Mal 1976

Ju1l 1976

SeEtember

November

1976

1976

% .of total TM used

�-12-

Table 9. Kilograms of trace mineral mix formulations
ranges in the Medano Creek study area.

TMI

April

TMIll

sheep

TMIV

Start of test period - 11.36 kg of each TM used.

1976

October

TMll

used on bighorn

1976

Kg TM used

4.26

8.37

10.18

9.36

% of total TM used

l3.24

26.02

31.64

29.10

Table 10. Kilograms of trace mineral mix formulations used on all bighorn
sheep range study areas during the duration of the study.

TMI

TMll

TMIII

TMIV

1.67

16.74

18.71

17.77

May 1976

.75

22.l3

31.59

24.38

July 1976

3.07

20.84

30.17

24.36

Date

March

1976

September

1976

.32

11.37

20.53

16.05

November

1976

.37

6.9l

12.04

10.91

6.18

77 .99

113.04

93.47

Total

�-13-

Table 11. Kilograms of all trace mineral mixes consumed by season
of the year for all study areas during the duration of the study.

Date

Kg

March 1976

54.89

May 1976

79.03

July 1976

78.44

September

1976

48.27

November

1976

30.23

As a result of the trace mineral mix preference measurements it was generally
concluded that the animal preferred TMIII formulation.
In November 1976
the following trace mineral mix was established as the preferred formula
and all subsequent trace mineral work has been based upon this formula:
Sodium chloride
50%
Ruminant Trace Mineral Premix
5%
XP-4
20%
Sodium bentonite
19.75%
Sulfur
.25%
Molasses
Vitamin A 135,000 units~
5%
35,000 units/lb
./
Vitamin D
1,000 units /lb./
Vitamin E
0

The trace mineral formulation is mixed, blended and dispensed in 25 lb sacks
by Feed Products, Inc., 1000 West 47th Avenue, Denver, Colorado
802ll,at
a cost of $250.00 per ton.
Fecal samples were collected from each study area whenever the trace mineral
boxes were serviced.
Both chemical analysis and "Baermanization" procedures
were carried out on the samples in an attempt to get some measure on the
physiological response of the animals to trace mineral supplementation.
The fecal samples were collected into paper sacks and allowed to air
dry in the laboratory before any analytical measurements were made. After
drying the fecal material was screened to remove any soil and/or vegetation
which might have been picked up in the process of collecting.
The samples

�-14-

were ground in a Wiley Mill and thoroughly mixed.
One portion of the sample
was "Baermanized" to measure the number of Protostrongylus sp. larvae
present and the remainder was used in the chemical analysis.
For specific
details see Federal Aid Document - Program Narrative Outline W-4l-R-27; Work
Plan 1, Job 16.
Tables 12, 13, 14, 15, 16, 17, 18 and 19 show the results of the chemical
analysis of the pellet groups for total ash, acid insoluble ash, acid
soluble ash, calcium, phosphorus, the calcium-to-phosphorus
ratio, magnesium,
potassium, sodium, copper, zinc, manganese, and iron.

Prepared

by

~-+/-=~_J--_.L.:::.~=-=-';::'=::!=='

Robert E. Keiss
Wildlife Researcher

C

_

�Table 12. Chemical analysis of composited bighorn sheep fecal pellet groups collected from the
Tarrya11 Mountain study area.
Collection
Date
Treatment

%

% Acid

% Acid

Total
Ash

lnso1.
Ash

Sol.
Ash

%

%

%

P

Ca:P
Ratio

%

CA

Mg

K

ppm
Na

ppm
Cu

ppm
Zn

ppm
Mn

ppm
Fe

Jan. '76

25.76

19.94

5.82

1.62

.57

2.84

.10

.25

152

10

50

ll5

375

Mar. '76

27.62

21.31

6.31

1.73

.65

2.66

.15

.32

ll3

9

48

125

275

May '76

18.32

14.ll

4.21

1.18

.73

1.61

.ll

.30

l37

II

59

175

350

July '76

15.73

ll.88

3.85

.97

.75

1.29

.12

.40

126

12

62

190

475

Sept. '76

16.38

ll.53

4.85

.84

.62

1.35

.l3

.35

ll5

9

55

180

400

Nov. '76

17.21

11.95

5.26

1.15

.73

1.57

.15

.40

145

8

64

ll5

375

Feb. '77

18.91

14.65

4.26

.87

.72

1.20

.15

.45

106

13

55

150

500

Mar. '77

18.58

12.90

5.68

.92

.77

1.19

.10

.42

121

l3

65

215

550

I
I-'
VI
I

�Table 13. Chemical analysis of composited bighorn sheep fecal pellet groups collected fr6m the
Trickle Mountain study area.
Collection
Date
Treatment

%

% Acid

% Acid

Total
Ash

Inso1.
Ash

Sol.
Ash

%

CA

%
P

Ca:P
Ratio

Mg

%
K

ppm
Na

ppm
Cu

ppm
Zn

ppm
Mn

ppm
Fe

August '75

18.76

11.50

7.26

1.37

.63

2.17

.19

,28

111

9

21

150

170

Jan. '76

28.72

21.89

6.83

1.47

.52

2.82

.26

.29

162

7

27

175

181

Mar. '76

21.82

14.07

7.75

1.36

.68

2.00

.27

.36

273

12

26

180

135

May '76

18.31

12.02

6.29

1.29

.73

1.76

.22

.41

385

15

34

170

176

July '76

17.63

10.31

7.32

1.18

.87

1.35

.29

.32

722

18

39

165

181

Sept. '76

15.28

8.81

6.40

1.76

.62

2.83

.37

.28

650

14

21

180

172

Nov. '76

18.73

13.53

5.20

1.65

.89

1.85

.18

.19

730

11

27

170

ll2

Jan. 'n

22.60

15.34

7.26

1.27

.73

1.73

.19

.35

290

13

18

160

ll3

Mar. 'n

22.72

17.38

5.34

1.25

.n

1.61

.22

.33

721

10

28

175

217

Apr. 'n

18.32

11.45

6.87

1.18

.94

1.25

.28

.41

890

14

34

180

218

%

I
I-'
0'\
I

�Table 14. Chemical analysis of composited bighorn sheep fecal pellet groups collected from the
South Platte River Canyon study area,
Collection
Date
Treatment

~;Acid
Insol.
Ash

% Acid

Total
Ash

Jan. '76

29.76

Mar. '76

%

Sol.

%

%

%

CA

P

Ca:P
Ratio

%

Ash

Mg

K

ppm
Na

ppm
Cu

ppm
Zn

ppm
Mn

ppm
Fe

22,37

7,39

1,87

.97

1..93

.11

,32

50

lJ

51

125

112

28.73

21.25

7.48

1,93

.81

2.38

,23

,28

50.

16

52

135

163

May '76

19.16

12.34

6.82

1,37

1.09

1.25

,29

,39

10

16

67.

110

212

July '76

18.37

13,27

5.51

1,42

1.19

1.19

,37

,47

173

13

6'3

115

229

Sept, '76

17,29

9.38

7,91

1,32

.9.4

1,40

,44

,51

1]5

11

77

120

235

Nov, '76

24.57

17.85

6.72

1.68

1,11

1 •.
51

,3]

,43

191

12

76.

130

172

Jan~ '77

19,11

12.43

6,68

1.62

1,25

1,29

,30

,50

150

16

68

16(1

187

Mar. '77

18.79

11.27

7.52

1.47

1.37

1,07

,46

.47

144

24

74

175

219

May '77

16.37

10.99

5.38

1.29

1.46

.88

.42

.56

168

27

83

185

235

,....I

-,..J
I

�Table 15. Chemical analysis of composited bighorn sheep fecal pellet groups collected from the
Collegiate Range (Love Ranch) study area.

Collection
Date
Treatment

Acid
Insol.
Ash

%

Total
Ash

Acid
Sol.
Ash

%

%

CA

P

Ca:P
Ratio

Mg

%
K

ppm
Na

ppm
Cu

ppm
Zn

ppm
Mn

ppm
Fe

Jan. '76

32.37

23.65

8.72

1.75

.65

2.69

.14

.35

175

10

25

150

120

Mar. '76

28.76

22.41

6.35

1.47

.63

2.33

.19

.47

175

12

36

175

115

May '76

19.27

14.06

5.21

1. 65

.70

2.35

.25

.63

250

14

48

150

146

July '76

15.26

10.11

5.15

1.55

.77

2.01

.23

.72

276

15

35

125

135

Sept. '76

19.51

11.22

8.29

1.50

.77

1.92

.23

.63

251

18

63

175

115

Nov. '76

20.33

12.57

7.76

1.50

.87

1.71

.23

.63

227

18

55

175

143

Jan. '77

29.18

23.84

5.34

1.13

1.08

1.04

.15

.45

208

28

175

200

325

Mar. '77

24.37

19.99

4.38

1.10

1.32

.83

.25

.47

276

21

125

200

350

May '77

19.76

15.04

4.72

1.62

.77

2.10

.16

,35

380

18

25

175

143

%

%

%

I
I-'
00

I

�Table 16. Chemical analysis of composited bighorn sheep fecal pellet groups collected from the
Collegiate Range (Jump Steady) study area.

Collection
Date
Treatment

%

% Acid

% Acid

Total
Ash

Inso1.
Ash

Sol.
Ash

%
P

Ca:P
Ratio

%

%

CA

Mg

K

ppm
Na

ppm
Cu

ppm
Zn

ppm
Mn

ppm
Fe

Jan. '76

29.76

23.41

6.35

1.68

.86

1.95

.16

.10

20

55

135

387

425

Mar. '76

31.28

24.53

6.75

1.73

.97

1.78

.19

.11

45

63

187

396

470

May '76

28.76

21.48

7.28

1.87

1.08

.79

.24

.16

172

48

221

425

480

July '76

24.73

17.10

7.63

1.93

1.27

.66

.21

.19

143

68

286

525

550

Sept. '76

37.62

29.90

7.72

1.75

1.12

.63

.25

.15

208

75

225

538

500

Nov. '76

34.62

26.89

7.73

1.27

1.16

1.09

.19

.21

118

81

151

681

475

%

I

I-'
\0

I

Jan. '77

38.72

32.14

6.58

1.09

.99

1.10

.20

.17

117

72

165

427

400

Mar. '77

31.28

23.99

7.29

1.73

1.38

1.25

.24

.26

146

65

186

480

475

May '77

27.63

21.09

6.54

1.12

1.18

.95

.17

.37

210

75

225

520

525

�Table 17. Chemical analysis of composited bighorn sheep fecal pellet groups collected from the
Mount Evans (Grant) study area.

Collection
Date
Treatment

%
Total
Ash

% Acid
Inso1.
Ash

% Acid
Sol.
Ash

%

CA

Jan. '76

29.62

23.41

6.21

Mar. '76

23.81

16.43

May' 76

18.62

July '76

%

%

P

Ca:P
Ratio

Mg

K

ppm
Na

ppm
Cu

ppm
Zn

ppm
Mn

ppm
Fe

1.27

.63

2.01

.10

.40

280

12

50

165

275

7.38

1.32

.69

1.74

.15

.47

460

11

49

187

300

13.51

5.11

.95

.71

1.26

.18

.58

935

13

62

193

375

17.63

13.01

4.62

.92

.93

.99

.26

.56

872

16

73

190

450

Sept. '76

20.28

15.47

4.81

.97

.95

1.02

.27

.41

621

17

65

181

425

Nov. '76

21.72

17.00

4.72

.92

.88

1.04

.31

.40

933

12

73

165

375

Jan. '77

20.35

16.17

4.18

.95

.89

1.07

.28

.46

1120

15

82

173

475

Mar. '77

18.81

14.88

3.93

.88

.92

.96

.21

.43

918

14

63

163

500

May' 77

18.62

14.77

3.85

.89

.91

.97

.30

.58

1151

16

85

182

550

%

I
N

0
I

�Table 18. Chemical analysis of composited bighorn sheep fecal pellet groups collected from the
Poudre River study area.
Collection
Date
Treatment

% Acid
Insol.
Ash

% Acid

Total
Ash

Sol.
Ash

%

%

%

Mg

K

ppm
Na

ppm
Cu

ppm

P

Ca:P
Ratio

%

CA

Zn

ppm
Mn

ppm
Fe

Jan. '76

18.72

13.66

5.09

1.16

.75

1.55

.10

.37

181

9

30

128

125

Mar. '76

15.79

9.97

5.82

1.71

.63

2.71

.12

.37

107

1

30

131

130

May '76

14.38

10.07

4.31

1.51

.81

1.86

.15

.40

111

3

40

137

125

Aug. '76

12.92

9.20

3.72

1.15

.76

1.51

.17

.48

152

5

35

118

128

Oct. '76

15.72

11.54

4.18

1.00

.85

1.17

.15

.57

163

7

30

129

140

Jan. 6, '77 13.54

10.10

3.44

.87

.90

.96

.15

.90

181

9

40

138

125

Jan. 14, '77 13.65

9.89

3.76

.87

1.02

.85

.12

.37

105

8

42

138

142

Jan. 21, '77 17 ..
22

12.15

5.07

1.12

.87

1.28

.17

.40

115

14

80

238

187

Feb. 3, '77 14.75

10.57

4.18

1.00

1.02

.98

.12

.35

148

13

55

138

157

Mar. 3, '77 12.55

9.26

3.29

.87

1.02

.85

.12

.47

129

8

65

163

102

%

I
N

t-'
I

�Table 19. Chemical analysis of composited bighorn sheep fecal pellet groups collected from the
Medano Creek study area.
Collection
Date
Treatment

%

% Acid

% Acid

)Total
Ash

Insol.
Ash

Sol.
Ash

%

%

%

P

Ca:P
Ratio

%

CA

Mg

K

ppm
Na

ppm
Cu

ppm
Zn

ppm
Mn

ppm
Fe

Aug. '75

24.52

16.84

9.33

2.20

.83

2.73

.30

1.15

253

15

64

200

235

Apr. '76

27.96

18.69

8.91

2.53

.58

4.47

.32

1.07

273

11

53

201

195

Oct. '76

21.41

14.12

7.29

2.04

1.28

1.67

.46

1.10

764

21

72

262

307

May '77

19.79

12.38

7.41

1.15

.84

1.30

.65

1.25

893

24

101

296

405

I

I
N
N

I

�January,

-23JOB PROGRESS

REPORT

State of

COLORADO
------~==~~~--------

Project

No.

Job Title

1

Manipulation

Period Covered:
Personnel:

Bighorn She~p

W-4l-R-27

Work Plan No.

Job No.
of Vegetation

1978

&amp; Mountain Goat Investigations
17

----------------------------

on Bighorn

Sheep Ranges

June 1, 1976 to May 31, 1977

George D. Bear

ABSTRAcr
Study plots near Cathedral, Colorado were treated with varied applications
of nitrogen and phosphorus fertilizers, and 2,4-D herbicide in 1971-72.
Two areas were selected for study sites: (a) an alpine range, and (b) a
ponderosa pine-bunchgrass range. The effects of these treatments were
monitored during the period 1972-1976.
The data has been analyzed and the
first draft of the report prepared.

��-25-

MANIPULATION

OF VEGETATION

ON BIGHORN

SHEEP RANGES

George D. Bear

P.N.O. OBJECTIVE
Improve the herbage yield, vegetative density
on selected bighorn sheep ranges in Colorado.

and vegetative

composition

SEGMENT OBJECTIVE
To analyze

data and prepare a manuscript.

PROCEDURES
The study area is located about 35 miles south of Gunnison, Colorado.
This
bighorn herd winters in two areas; in a low (8,000 feet elevation) bunchgrass
and ponderosa pine type, and in a high (11,000-14,000 feet elevation) alpine
vegetative type. Study plots were established on these two winter ranges,
and the plots were treated with phosphorus fertilizer, nitrogen fertilizer,
and 2,4-Dherbicide
(16 treatment levels) in 1971-72.
Effects of these
treatments were monitored from June, 1972 to June, 1976. The data was analyzed
at the Colorado State University Statistical Center ?nd the first draft of the
manuscript has been completed •

Prepared

by

. /f.~~~'~

~__ ~or~
George D. Bear
Wildlife Researcher

_

��January,

-27JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~----------

Project

No.

W-4l-R-27

1978

Bighorn

Sheep

&amp; Mountain

Goat Investigations

1

23
~,-,;----::::-_:
Job No.
Bighorn Sheep LungwormC ~-o-n~t-r-o~l------------------------Job Ti tIe __B_y
__B_i_o_l_o_g_l._·
c_a_l
__C_o_n_t_r_o_l
o_f Sn_a_i_l
__I_n_t_e_rm
__e_d_i_a_t_e
__H_o_s_t_s
_

Work

Plan No. ____

Period

Covered:

Personnel:

Chester

June 1, 1976 - May 31, 1977
Ng, Harold

R. Shepherd,

and Thomas

N. Woodard.

ABSTRACT
Preliminary
sampling to determine sample size requirements
was completed.
Snail distribution
is apparently clumped to the extent that the maximum
number of samples which c6uld be collected and analyzed would result in
a poor estimation of the snail population.
The decision was therefore
made to cancel the original objectives of the study.
A new objective is
to compile and publish a literature review ori land snail ecology~

��-29-

BIGHORN SHEEP LUNGWORM CONTROL
BY BIOLOGICAL CONTROL OF SNAIL INTERMEDIATE

HOSTS

Thomas N. Woodard

P.N.O. OBJECTIVE
To test the effects of burning treatments as a tool to reduce populations
of the snail intermediate hosts for bighorn sheep lungworm parasites.

SEGMENT OBJECTIVES
1.

Select two aspen patches within a portion of the Poudre River bighorn
sheep range selected for a prescribed burning experiment to improve
range for bighorn sheep.

2.

In each aspen patch, establish
lanes between blocks.

3.

Construct
patch.

with numbered

4.

Test-sample

variables

5.

Analyze

4 treatment blocks and construct

steel stakes a sampling

fire

grid in each aspen

in each aspen patch.

snail and soil samples in laboratory

and calculate

sample size.

METHODS AND MATERIALS
Methods

and materials

have been described

by Shepherd

(1976).

RESULTS AND DISCUSSION
Preliminary sampling to determine sample size requirements for estimating
snail populations was conducted during August and September, 1976. Random
and stratified sampling schemes were used. Based on random sampling, there
was a mean of 11.88 + 2.14 (SE) live snails in each I-liter sample of soil.
A sample size requir;ment of approximately 25 was computed for each replicate
to estimate the population within 30 percent of the true mean with 95 percent
confidence.
Stratified sampling did not improve precision.
The decision was made to cancel the original objectives of this study because of poor precision and time limitations which prevented ,increasing sample
sizes to improve precision.
The new objective is to compile and publish a
literature review on land snail ecology.
Such a publication would be beneficial to future investigations involving snail ecology and control.

�-30-

LITERATURE CITED
Shepherd, Harold R. 1976. Bighorn sheep lungworm control by biological
control of snail intermediate hosts. Colorado Div. Wildl., Fed. Aid
Program Narrative Outline. l5p.

Prepared by

~~A&lt;w~
N.

Thomas
Woodard
Wildlife Researcher

'

�January,

-31JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~----------

Project

No.

Work Plan No.
Job Title
Period

W-4l-R-27

Bighorn

1

Job NO.

Sheep &amp; Mountain

Goat Investigations

24

E_x_t_·e_n_d_1_·n_g~B_1_·g~h_o_r_n
__ S_h
__e_e~p
__R_a_n~g_e_s

Covered:

Personnel:

1978

June 1, 1976 to May 31, 1977

George D. Bear

ABSTRACT
Bighorn sheep transplants were monitored in Cache la Poudre Canyon in northcentral Colorado and on the Cebolla Creek Wildlife Area in southcentral
Colorado.
Twenty-five bighorns were transplanted to the lower portion of
Cache la Poudre Canyon on January 21, 1975. This herd became very nomadic
in behavior and many migrated to the upper portion of the canyon.
Two years
following the transplant (spring 1977) a small herd (15 head) has become
established at the release area and there is some migration between their
original range and the new site.
A small group (9 head) of sheep were transplanted onto the Cebolla Creek
Wildlife Area during February, 1976, and an additional nine head in March
1977. The ewe-lamb segment of the herd summers in the vicinity of the wildlife area, while the rams migrate to the alpine to summer with bighorns
native to the area. Both segments of the transplanted herd winters on the
wildlife area where they were released.

_
_

��-33-

EXTENDING

BIGHORN

SHEEP RANGES

George D. Bear

P.N.O. OBJECTIVE
Determine if the distribution and population of bighorn sheep herds can be
increased by transplanting sheep into suitable areas adjacent to the present
concentrations.

SEGMENT OBJECTIVES
1.

Monitor the transplanted
Cebolla Creek (Fig. 1).

2.

Transplant

additional

herds in Cache la Poudre River Canyon and on

animals

from Saguache

Creek to Cebolla Creek.

METHODS AND MATERIALS
Information was gathered on the transplanted herds by observing from roads,
hiking, or aerial surveillance, whichever was most appropriate.
The
following information was recorded for each individual observed: date,
location (also plotted on a map), and collar number.
Additional bighorns were secured from the Trickle Mountain herd and released
on Cebolla Creek. An unsuccessful attempt was made to capture sheep using
the permanent trap located on Trickle Mountain.
Due to the lack of snow the
sheep did not move to the bait stations at the trap. Nine bighorn sheep (7
ewes and 2 lambs) were captured on Middle Creek, approximately Smiles
away,
using a portable drop-net trap. Five of the ewes were marked with the commonly
used 4-inch collars (white with black numerals), two ewes with radio-telemetry
collars, and the lambs were marked with red 2-inch Richie eartags.
The sheep
were transported to the release site in a large livestock trailer, and held
overnight in a temporary corral constructed with 8-foot wooden elk panels.

RESULTS AND DISCUSSION
Cache la Poudre
The herd (25 sheep) being evaluated was transplanted on January 21, 1975.
Reactions of these displaced bighorns were extremely variable.
They moved
around a great deal during the first 30 days after their release.
Some individuals remained together, while others moved between groups.
Eighty days
after the transplant it appeared that 16 sheep had settled in the vicinity of
the Sheep Mountain release site, while the others returned to the upper portion of the canyon (Fig. 2). Several of the transplanted bighorns traveled
up the canyon, mingled with other bighorns, and then went back down the
canyon and joined the Sheep Mountain group.

�COLORADO
MO"AT

SEDGWICI(

LOGAN

WELD

ROUTT

PHILliPS

MONGAN
WASHINGTON

I YUMA

/lI08LANCO

EAGLE

GARFIELD
DOUGLAS

ELBERT

LINCOLN

MESA

I

I

~

CHEYENNE

(

,

MONTIIOSE

I"

~
PUEBLO

SAGUACHE

SANMIGUEL

~

~
~

I

L......
,at Cebolla

Creek
~Study Area

"

{"lJSTER

I

I

I
~

ICROWUY

I

II

IPliO WEllS

SENT

"DOLORES
RIOGHANDE
MONTEZUMA

F\ /
.,

ARCHULETA

Fig. 1 • Location

CONEJOS

..,

LAS ANIMAS

of the Cache la Poudre and Cebolla Greek study areas in Colorado.

ISACA

I

w
.l:I

�I

w

U'1

I

Co.

~.J.

lo.~e
flO~O

e

$

llJ
-.J

&amp;

.Stove

Prairie

N

t
o

3

Scole:Miles

Fig. 2.

6

Distribution of transplanted bighorn sheep in Cache la Poudre River Canyon.

�-36-

By mid-summer (1975) only seven ewes, one yearling, and three new lambs
were found in the Sheep Mountain area. Hunting activity during the deer
season caused the sheep to abandon the area. Two of them returned to the
trap site, while the others assumed a nomadic behavior and wandered around
in the lower portion of the canyon throughout the winter and the summer of
1976. Two young rams returned from the upper canyon area and remained with
the transplanted group during the winter, then moved back up the canyon.
Also, one ewe returned from the upper area to rejoin this small band during
the summer of 1976. Dogs were reported to have been chasing the sheep in
the vicinity of Poudre City on July 29, 1976. One lamb was separated from
the group; its fate is unknown.
Also, two collared ewes were reportedly
killed on Red Mountain by deer hunters; this was not confirmed.
This group of transplanted sheep has been very difficult to follow, and many
of the collars are now torn which makes individual identification nearly
impossible.
It appears there are now at least fifteen sheep in the transplant
area - seven adult ewes from the original transplant, two young rams (lambs
when transplanted), two 1975 lambs (both males), and four 1976 lambs. The
two young rams from the original transplant move between the herds in the
upper part of Poudre Canyon and this transplanted herd.
Therefore, a small
herd was established in the lower portion of the canyon, and there is some
migration between the established bighorn range and the new site. However,
there is a need to supplement this small herd to make it a viable population.
There is some speculation as to how the transplanting procedures could have
been altered to improve the success of this transplant.
Young animals, 1-2
years old, may be less home-oriented and more adaptable than adult sheep,
therefore, the young sheep may have established at the release site more
readily.
Also, it is believed that animals from another locality would have
been more appropriate for the transplant.
They likely would become established
at the release site before making contact with the native herd in the upper
portion of the canyon.
At the release site the sheep left the transplanting truck in a frantic haste,
and several were observed several miles away in just a few hours.
This may
have been prevented if the sheep were held in a pen and allowed to calm down
before release.
Another factor influencing the transplant was human activity in the area,
primarily during hunting seasons.
This activity disturbed the sheep and
dislocated them. It is speculated that a temporary "land closure" would have
allowed the transplanted bighorns to become more firmly established on Sheep
Mountain.
An additional recommendation would be to equip several individuals
in a transplanted herd with radio-telemetry collars.
A large percentage of
the days (approximately 85%) were unsuccessfully spent searching for the
sheep. Whereas, 90 percent of the attempts to relocate sheep on the Cebolla
Creek transplant site were successful.
It is desirable to have more than
one individual telemetry-marked,
since the transplanted herd often splits-up,
an individual is injured, or a transmitter becomes nonfunctional.

�-37-

Cebolla Creek
A small group of bighorns were trapped on Saguache Creek in southcentral
Colorado during February 1976 and transplanted onto the Cebolla Creek
Wildlife Area to encourage bighorn sheep use in the lower ranges.
Nine
(6 ewes, 1 lamb, and 2 rams) were trapped with a large group-trap, marked
with collars, and transported to the release site in a large livestock
trailer.
Two of the ewes were fitted with telemetry collars obtained from
Telonics, Mesa, Arizona.
These collars have been very beneficial in
locating this group of transplanted sheep.
The sheep remained at the release site on Cebolla Creek Wildlife Area until
mid-March, when the snow started melting.
Two rams native to the area joined
the group. In March these rams, the two transplanted rams, and the lamb
migrated to the alpine range. The transplanted rams migrated from the alpine
range back to the release site in mid-September, while the lamb did not appear
on the wildlife area until December.
During the spring the transplanted ewes explored the canyon up to 3 miles in
each direction, but returned to the release site every 1-2 days. By June they
had settled along Rock Creek, approximately 2-4 miles from the release site,
and remained there for the summer (Fig. 3). These six ewes had five lambs.
Two ewes and two lambs migrated back to the Cebolla Creek Wildlife Area in
September and joined the rams.
Activity during the elk hunting season caused the sheep to leave the wildlife
area. One of the radio-collared ewes was shot on Fish Creek, approximately
5 miles from the release site. The carcass had been skinned and hung in a
tree; the head and collar buried under debris approximately 100 feet away.
There was 6 inches of snow on the ground, yet the collar and head were easily
found by using the telemetry equipment.
By December all the other sheep,
except three of the new lambs, were back on the wildlife area.
On March 31, 1977, nine bighorns (7 ewes and 2 lambs) were trapped on Saguache
Creek and transported to the Cebolla Creek Wildlife Area.
The trailer was
backed-up to the holding corral and the trailer door left open. It took the
sheep 10-15 minutes to leave the trailer.
They ran along the corral fence,
but did not strike it or attempt to jump over it. The sheep were left in the
pen overnight.
When we arrived the next morning, they were very calm, some
were feeding on the hay left in the pen, and others were bedded down. The main
gate to the holding pen was then opened.
It took approximately 20 minutes for
the first sheep to find the open gate and walk out of the pen. It started
feeding in the immediate area. It took 45 minutes for all of the sheep to
exit from the pen. They were very calm and fed around the holding pen for some
time, then gradually moved up the slope feeding as they went.
Therefore,
holding the sheep overnight in a corral proved to be very useful in calming
them down and releasing them as one group.

�-38-

..x
Q)
Q)

'u

EWE - RAM
N"'WINTER
RANGE

r

~,.'~ San Luis Pk.

N

1

o
Fig. 3.
Wildlife

Distribution
Area.

of bighorn

sheep transplanted

2
Scale. Miles

4

to the Cebolla Creek

�-39-

The new transplants remained in the area for approximately three weeks.
There was very little snow at the release area, and the lamb-ewe group
of the previous years transplant had returned to their summer range on
Rock Creek. Thus, the new group joined up with the ram group. By late
April the rams migrated to the alpine range on Baldy Chato, accompanied
by some of the ewes. This was determined by aerial surveillance or tracking
of one radio-collared ewe; contact wasn't made with the animals on the ground
to identify individuals.
The remainder of the sheep remained on the wildlife
area, but were exploring the surrounding area in groups of two's and three's.
The telemetry collar in this group ceased to function.
By the end of the
work segment (June 1977) these new sheep had joined up with the lamb-ewe group
on Rock Creek. Two ewes are unaccounted for and it is assumed they are on the
alpine range with the radio-collared ewe.

Prepared by

~....,_-..;...._._._.VJJ_. _'_~
George D. Bear
Wildlife Researcher

.__

��January, 1978
-41JOB PROGRESS REPORT

State of

COLORADO
------~~~~~--------

Project No.

W-4l-R-27

Work Plan No.

1

Job Title

Job No.

~2~5

Prescribed Burning to Improve imd Enlarge Bighorn Sheep Ranges

Period Covered:
Personnel:

Bighorn Sheep &amp; Mountain Goat Investigations

June 1, 1976 through May 31, 1977

Thomas N. Woodard

ABSTRACT
A detailed Program Narrative Outline was completed during this seglnent.
Field work, consisting of pretreatment data collection. will begin during
Segment 28.
.

_

��January,

-43-

1978

JOB FINAL REPORT

State of
Project

C_o_1_0_r_a_d_o-----W-4l-R-27

No.

Work Plan No.
Job Title
Period

Investigation

Covered:

Personnel:

1

Bighorn

Sheep

Job No. ·__
of Spontaneous

&amp; Mountain

Goat Investigations

...:C..:.O_N_TRA_C
__
T---...:{~2..:.6~)

Diseases

of Bighorn

_

Sheep

June 1, 1975 - May 31, 1977

Dr. C. P. Hibler, R. E. Keiss,
Spraker, and J. Wegryzn.

R. L. Schmidt,

B. Davies,

T.

ABSTRACT
The drug cambendazole was tested in a form marketed as CamVet for its
efficacy in eliminating both lungworm adults and somatically stored larvae.
Results of this test indicated that the preservative in CamVet (scorbic
acid) was toxic to bighorn sheep rumen microbiota and five sheep died
after three days of oral administrations
of CamVet.
Clearly~ the drug in
its marketed form is not suitable for use on wild bighorns as an anthelmintic.
In addition CamVet was not palatable to sheep when offered in apple mash.
If the scorbic acid preservative was removed then the remaining pure cambendazole was palatable, but its efficacy against lungworm adults and larvae was
not determined.
The drug fenbendazole was evaluated for its toxicity to sheep and for its
palatability in apple mash.
It was found to be non-toxic and palatable.
Its efficacy against lungworm ·also was not tested.

��-45-

SPONTANEOUS

DISEASES

OF BIGHORN

SHEEP

C. P. Hibler

OBJECTIVES
The specific

objectives

of this study are:

1.

Evaluate two new anthelmintics, fenbendazole and albendazole
for efficacy against lungworms, both adult and somatically
stored larvae. Also to evaluate these drugs, in the event
that they show efficacy, for toxicity and palatability.

2.

Evaluate the efficacy of cambendazole and thiabendazole against
lungworms, both adult and somatically stored larvae, on a 3 to
7 day therapeutic regimen; in addition, evaluate both fenbendazole
and albendazole using the same regimen.

3.

Evaluate
Rampart,
lungworm

4.

Experimentally infect ewes with lungworm to induce transplacental
transmission of lungworm, and infect lambs and yearlings with
lungworm for use in drug trials.

5.

Perform postmortem examinations to determine
any and all bighorn sheep in Colorado.

6.

Raise bighorn and mouflon lambs for use in drug trials and for
use by personnel with the Colorado Division of Wildlife for other
studies.

7.

Support Division of Wildlife trapping and transplanting
bighorn sheep herd evaluations throughout Colorado.

the efficacy of treatment of bighorn ewes on Pikes Peak,
4-Mile and Poudre Canyon by monitoring fecal output of
larvae in lambs, adults and yearlings.

cause of death of

efforts and

PROCEDURES
The evaluation of various anthelmintics at different dose levels the past
two years has essentially depleted infections in the bighorn sheep and bighorn/mouflon sheep.
Consequently, considerable effort must be expended this
year to culture and infect snails with lungworm larvae and then to infect
sheep with their larvae.
The snails currently in use (Vallorria spp.) will
be used but, in addition, a different species of snail (Triodopsis albolabris)
will be evaluated.
This rather large snail (2.5-3.5 cm) has proven to be an
excellent experimental host for other species of lungworm; hopefully, it will
work for bighorn sheep lungworm.
Moreover, it propagates rather well in
captivity.
Currently, we have available for drug evaluation studies a total of 32 sheep;
lambs we expect this year are 16-18, making a total of 48-50 sheep available
for anthelmintic trials.
Some of these sheep are infected; their infection

�-46will be "bolstered" with additional larvae and the uninfected animals
will be infected as soon as possible (some are currently being infected) .
1.

Two experimental anthelmintics: fenbendazole and albendazole must
be evaluated for efficacy.
This will be done with a single
therapeutic dose administered orally.
Lambs and yearlings will
be used for this purpose.
Three groups of 5 animals each will be
used; each group will be given a drug and then fecal larval output
evaluated for the next 8-12 weeks.
Any drug showing efficacy will
be evaluated for palatability by mlxlng in apple pulp and for
toxicity by oral administration.
The same sheep will be used
for these two purposes.
Drugs showing efficacy against adult lungworm will then be evaluated for efficacy against somatically stored larvae in adult bighorn
and bighorn/mouflon sheep ewes. A total of 19 ewes are available.
All will be infected with lungworm for transplacental transmission
and 12 will be administered (orally) one (or more) of the drugs; remaining sheep (seven) will be used as controls.

2.

Both cambendazole and thiabendazole have high efficacy against
somatically-stored
larvae.
Preliminary studies indicate these two
compounds have considerable efficacy against adult lungworm if
administered daily for 3 to 7 days. Therefore one group of 5 sheep
(lambs and yearlings) will be administered cambendazole daily for
5 days; the other group given thiabendazole daily for 5 days. Lungworm larval output will be monitored for 8-12 weeks (or as long as
necessary) to determine efficacy.

3.

Since treatment of bighorn sheep on Pikes Peak, Rampart, 4-Mile and
the Poudre Canyon herd will be continued indefinitely, these efforts
must be monitored by examination of fecal samples for lungworm larval
output.
Lambs, yearlings and adults must be monitored, but efforts
should be concentrated on lambs. At least 50 fecal samples (from
each of the.above areas) per month in July and August, October and
November, and January, February.
These samples will be dried,
weighed, examined by the Baermann apparatus and evaluated quantitatively for larvae/gram feces. Data will be compared with previous
years samples from Pikes Peak and other herds to evaluate efficacy.

4.

See under procedures

5.

Bighorn sheep sick and/or dead from any cause will be given a
thorough post-mortem examination, including histopathology,
to determine the cause and nature of the illness and/or death. Moreover, all
animals will be examined for somatically stored larvae of lungworm.
The examination will include complete microbiologic, virologic,
parasitologic and toxicologic possibilities.

6.

Currently we have 9 adult female mouflon and mouflon/bighorn hybrids
and 10 adult female pure bighorn sheep in captivity.
Lambs will be
removed from these sheep and bottle-raised as imprinted animals.
Lambs are removed at 24-48 hours following birth and put on a bovine

- first paragraph,

and procedures

numbered

1 and 2.

�-47-

milk-syrup-egg diet with human vitams added when necessary.
Rearing lambs (successfully) in captive conditions necessitates
that they be separated into 2 or 3 groups. Personnel rearing
one group have no contact with other groups.
This is, of course,
essentially a full-time job for at least two months, requiring
almost constant attention and veterinary care.
We anticipate about 24 lambs this coming year.
lambs will be used in procedures 1, 2, and 4.
7.

Eventually,

these

Proposed studies by the Division of Wildlife on trapping and transplanting, and herd evaluation throughout Colorado will necessitate
some support from the personnel with the Wild Animal Disease Center.
This will take the form of: (1) Training, and (2) Service.
We
anticipate training personnel involved in the project to (a) Drench
sheepwith anthelmintics currently in use, (b) Take nasal swabs for
bacteriologic and virologic examination, (c) Take fecal samples for
parasitologic examination, and (d) Remove blood for physiologic
baseline data and virologic titers.
Fecal, bacteriologic and
virologic examinations will be performed on each animal to determine
its health status.
Population evaluations are, fo"r the most part, based on census and
fecal examination.
Where herd evaluations are conducted the center
will provide support to examine fecals brought in by Division personnel.
Any sick and/or dead animal resulting
examined as in procedure 5 (above).

from these efforts will be

RESULTS
1.

Objective No. 1 was not completed during this fiscal year because
of difficulty encountered in the rearing and infecting snails with
the larval stages of Protostrongylus.
Snail cultures became infected with fungi and mites, resulting in a high degree of mortality.
Moreover, because feces containing first stage larvae of
Protostrongylus were not made available by Division of Wildlife personnel, considerable difficulty was encountered in obtaining sufficient
fecal matter to infect the snails that survived.
Therefore, only five
lambs were infected, and these are still infected.
Currently, snails
are growing quite well, and considerable numbers have been infected.
This segment objective will be completed during this coming fiscal
year.

2.

Cambendazole has been used for treatment of sheep to eliminate the
somatically stored third stage larvae of Protostrongylus at a dose
level of 8.4 grams per adult sheep (based on a 150 pound sheep).
Studies conducted by Dr. Terry Spraker at the San Diego Zoo indicated
that if this compound was fed to lungworm infected animals on a 5
to 7 day regimen then all lungworm would be eliminated.
Consequently,
five bighorn/mouflon sheep were placed in a pen to determine if a

�-48-

5 to 7 day treatment regimen would remove all their lungworm.
The drug was administered orally at the above dose level once
a day. On the third day of oral dosing, all five sheep went
off feed and became depressed.
Later that same day, a bloody
diarrhea developed.
All animals continued to become progressively weaker despite supportive therapy and all died.
Postmortem examination revealed considerable hemorrhagic enteritis. Since we did not know if this was the result of cambendazole
or the preservative in the cambendazole, the proposed trial with
thiabendazole (5 to 7 day regimen) was cancelled.
Subsequent
research revealed that sorbic acid, the preservative added to
CamVet was a fungicidal and bacteriostatic agent. The large
amount of CamVet (240 cc) necessary to insure that each sheep
received a therapeutic dose apparently caused the problem.
Because the powdered form of cambendazole (100% cambendazole) was
becoming increasingly difficult to obtain, and because the results
of the 5 to 7 day regimen indicated toxicity from the active ingredient or from the preservative, a considerable amount of subsequent research plans were set aside so that we could evaluate
CamVet prior to the bighorn sheep trapping and transplanting season.
It was necessary to determine if CamVet was palatable in apple
pulp; and to do so necessitated mixing a therapeutic dose (in three
pounds of apple pulp) for each sheep and observing their reaction.
At the same time, personnel involved in the trapping-transplanting
operation requested that we evaluate fenbendazole for palatability
and toxicity.
As a consequence of this type of research (which was absolutely
necessary to the entire operation) our attempts to evaluate fenbendazole were then set aside. The procedure used was to mix CamVet
in one batch of apple pulp. fenbendazole in another batch of apple
pulp; the third batch was pure apple pulp. This was placed in the
pens and offered to all of the mouflon sheep available to determine
selectivity.
The results indicated that CamVet in apple pulp was
not palatable.
Since research in the early and late fall of 1976 indicated that
either cambendazole or the sorbic acid was toxic to sheep and since
CamVet was not palatable to sheep, considerable time and effort was
then initiated to remove the sorbic acid from the CamVet, leaving
pure cambendazole.
It was later determined that if the CamVet was
chilled, centrifuged and then the sediment washed once with water,
virtually all the sorbic acid was removed. When this was mixed with
apple pulp and offered to the sheep, they found it to be quite
palatable.
Since pure cambendazole was not available in quantities
sufficient to treat all of the bighorn sheep the Division proposed
to treat in 1976-77, most of the personnel assigned to work with
snails and bighorn sheep and larvae were assigned the job of removing
the sorbic acid from the CamVet to make it palatable and, of course,
non-toxic to the sheep. As a result of this effort and as a result
of evaluating the cambendazole, CamVet and fenbendazole for toxicity,
all of the sheep that were either naturally infected or experimentally
infected with lungworm for the fenbendazole trials were at that point
unusable.

�-49-

Evaluation of fenbendazole in five bighorn sheep, at dose
levels of four times the dose currently in use did not cause
any toxic side effects that were observable.
Unfortunately,
the animals on which fenbendazole was evaluated (for toxicity)
were not infected with lungworm.
3.

Fecal samples that were to be collected from bighorn sheep from
Pikes Peak, Rampart, 4-Mile and the Poudre Canyon herd were, for
the most part, given to Robert Keiss for evaluation.
These data
must be gleaned from his report.
The only fecal samples obtained
for evaluation on a pre-treatment and post-treatment basis have
been analyzed and the results are attached.
Unfortunately,
fenbendazole was used indiscriminately
throughout the course of
the year, and this was done without any significant information
regarding its efficacy at killing all the adult lungworm and/or
the somatically stored third stage larvae. When used in the field,
no pre-treatment and very few post-treatment samples were obtained.
Many of the post-treatment samples were evaluated by Robert Keiss.

4.

Considerable difficulty was experienced in rearing snails, obtaining
first stage larvae, infecting snails, and keeping snails alive a
sufficient period of time to experimentally infect the mouflon/
bighorn sheep for evaluation of fenbendazole and albendazole.
As
a result of the request by Colorado Division of Wildlife to evaluate
fenbendazole for palatability and toxicity, and the results of
Objective No. 2 (evaluation of the 5 to 7 day regimen) most of the
bighorn/mouflon
sheep scheduled for the drug evaluation trials had
to be utilized for this purpose.
Since that time, snail culture
and infection has improved immensely and about 1/3 (seven sheep)
have been infected; however, this is not as yet a sufficient number
to conclusively evaluate the efficacy of the above mentioned drugs.

5.

The sheep dying as a result of the CamVet sorbic acid trials are
reported above.
One lamb captured in the net up the Poudre River
died as a result of heat prostration in the net. The only other
bighorn sheep examined at postmortem during the last fiscal year
were four cases of Johne's disease from the Mount Evans-Grant herd.
The postmortem results of these four animals is appended to this
report.

6.

The bighorn ram used for breeding the mouflon/bighorn
cross ewes
this last year apparently was not a satisfactory ram, because only
four of the ewes were settled (producing six lambs) •

.-~

Prepared

by

C: . \-'

'" . 0 0

""~.":~~~Q6.J

C. P. Hibler

�-50-

Bighorn Fecal Samples
Sample it

Wt (g)

Tot Ll

'L,/g feces

Saguache - Collected 3/24/77
Sl
S2
S3
S4
S5
S6 lamb
S7 lamb
S8 lamb
1 lamb
2
3
4 lamb
5
6
7 lamb
8
9
10
11
12
13
14
21
22
23
24
25 Ram
26
27 Ram
28 lamb
29
30
31
32 Ram
33
41 lamb
42
43 Ram
44
45
46 lamb
47 lamb

5.2
18
4.3
10
9.3 (not fully dried) 313
5.85
90
4.2
20
4.1
36
1.1
4
1.4
115
6.5
44
236
9
14
1030
2300
7
350
5
6.5
550
1075
7.5
700
4.5
23
4
575
4
2
5
9
5.5
850
8.5
125
6
300
4
300
3.75
130
6
562
4.5
6
345
645
8
1S
3
600
4.5
525
4
150
3
150
3.5
93
6
150
4.5
2.5
75
5
375
6.5
173
6.5
750
3
150
3.5
470
2.5
150

3.5
2.3
37
15.4
4.76
8.8
4
82
7
26
74
329
70
85
143
156
6
144
0.5
0.6
100
21
83
80
22
125
58
81
25
l33
l31
50
43
16
33
30
.75
27
115
50
135
60

�-51-

Sample 4f

Wt (g)

4-Mile - Teller Co - Collected Winter 76-77

Tot L1

L]/g feces

(Pre-treatment)

3
3
1.5
1
2.5
2.5
2.5
4
2.5
3

500
200
50
50
50
100
400
150
250
150

167
67
33
50
25
40
160
37.5
100
50

4-Mile - Collected 3/24/77
3.5
1
4
2
6.5
3
6
4
7.5
5
6
6
7
3.5
7.5
8
4.5
9
4.5
10
11
3
12
5
5.5
13
12.5
14
5
15
5.5
16
5
17
6.5
18
3.5
19

7
56
0
1125
1
2850
64
15
6
5
0
4
46
975
1
1800
1
750
83

2
14
0
188
0.13
475
18
2
1.3
1.1
0
0.8
8.4
78
0.2
327.3
0.2
115.4
24

1
2
3
4
5
6
7
8
9
10

Rampart Range - Collected Winter 76-77 (pre-treatment)
1

2
3
4
5
6
7
8
9
10

1.5
2
4.5
3.5
3
4
4
3
3
3.5

450
700
250
200
250
150
350
600
350
350

300
350
56
57
83.3
38
87.5
200
116.7
100

�-52-

Sample 4F

Wt (g)

Tot L1

L1/g feces

11
12
13
14
15
16
18
19
20
21
22
23
24
25
26
27
28
29
30

3
4.5
3.5
3
4
4
4.5
2.5
4
4
5
4
5
3
2
2.5
4
4.5
5

700
500
800
400
100
0
250
500
0
0
0
350
150
100
200
250
100
300
0

233
111
229
133
25
0
55.5
200
0
0
0
87.5
30
33
100
100
25
67
0

Rampart Range - Collected Winter 76-77 (Post-treatment with Fenbendazole)
1
2
3
4
5
6
7
8
9
10

4
3.5
5
4
2.5
3
7
3
2
3

0
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
0
0
0
0

Rampart Range - Collected Winter 76-77 (Post-treatment with Cambendazo1e)
1
2
3
4
5
6
7
8

9
10
11
12
13

2
4
3
3

1.5
2
2
1.5
1.5
2
2

2.5
2

0
0
50
50
100
100
0
0
0
0
0
0
0

0
0
17
17
67
50
0
0
0
0
0
0
0

�-53-

Sample iF

Wt (g)

Tot L1

LlJg feces

Pike's Peak - Collected Winter 76-77 (pre-treatment)
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40

2.5
2.5
2
2
2
2.5
2.5
2.5
2
1.5
2
3
2
2.5
3
2.5
1.5
4.5
2
3.5
1.5
2
2
2
2
2
1.5
3
2.5
1.5
1.5
4
1
2
1.5
3
4
9
2.5
1.5

2025
350
375
1025
1250
150
800
675
162.5
1212.5
475
700
300
3000
900
1850
50
900
1050
800
300
200
300
750
1300
450
450
250
1600
250
0
1350
600
100
500
300
400
0
100
225

810
140
188
512.5
625
60
320
270
81.3
808
238
233
150
1200
300
740
33
200
350
229
200
100
150
375
650
225
300
83
640
167
0
338
600
50
333
100
100
0
40
150

0
0
0
1
0
0
1
0
0
0

0
0
0
1
0
0
0.7
0
0
0

Pike's Peak - (Post-treatment)
1 (yellow ear tag 2)1
1
2 (12)
1
3
1
4
1.5
5 (31)
1.5
6 (E33)
7 (collar tag 28) 1.5
8 (collar tag 27) 1
3.5
9 (30)
1
10 (old yellow 2)

�-54-

Sample 1t

Wt (g)

Pike's Peak - (Post-treatment
11 (lamb 39)
12
11
6 (ear tag)
32
47
64
65
70
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40

1
5.5
2
2.5
4
1.5
1.5
3
5
2
2
2
2
3
3
1.5
2
2
1.5
2
2.5
1.5
2.5
2
1.5
1
2
2
2
2.5
3
3
1
1.5
3.5
2
3.5
2
2.5
2
1.5
3
2
- 2.5
2
2.5
2
1.5
2

Tot L1

Lljg feces

0
0
0
0
0
0
0
79
0
39
210
50
0
0
0
50
100
0
50
100
3300
250
350
150
0
0
0
0
2050
150
200
0
0
0
0
0
0
250
150
50
100
2100
500
200
450
0
0
0
50

0
0
0
0
0
0
0
26
0
20
105
25
0
0
0
33
50
0
33
50
1320
167
140
75
0
0
0
0
1025
60
67
0
0
0
0
0
0
125
60
25
67
700
250
80
- 225
0
0
0
25

continued)

�-55-

Bighorn Fecal Samples

Co~ Creek - 3une 9, 1977 - Set up August 2, 1977
Samp,le ~.
Wt (g)
'tot Ll
1
tot feces 1.3
15
2
tot!~feces 1.2
0

3.4
tot feces 1.8

0

5 y. ole sam~le
1.1 4 pellets
6
tot feces 2.3

0

3

4

Cross ~ountain
1

2
3
4
5

6
7

a

9

10
11

12
13
14

15
16

17
1a
19

2.8
2;4
4.6
4.3
1.1
4.6
2.7
2.3
3.1
7.0
4.8

o

o
o

- 3une 6, 1977 - Set up August 4, 1977
0
3.6
tot feces 1.7
3

4.1
tot feces 2.5
4.3
tot feces 2.1
tot feces 3.7
tot feces 2.1

feces

11.5
1.1

2

1
larval output not reflected - (sample wts too small).

T~e

~/g

2
1
1

22
3
12
Set up August 8, 1977
16
1

0

0
18

0.4

0

1.8
0.5
0.4
0.2
10..
5
0.8
5.1

'.1
0.4
0
0

16.4

0
6

0

5

2.2

5
2
19

1.6
2.0
4.0

•

2.2

�-56-

Basalt CaEtive SheeE Herd
Pre-treatment 2-15-77
g Feces Used

L1 Count

L/g

Lamb 1)

3.3g

56

16.97

Lamb 2)

3.3g

66

20

Lamb 3)

3.6g

156

43.33

4)

3.9g

0

0

5)

3.9g

88

22.56

6)

3.9g

0

0

7)

4.0g

0

0

8)

3.1g

0

0

9)

3.9g

0

0

10)

3.9g

33

8.46

11)

3.6g

67

18.61

12)

3.1g

65

20.97

13)

3.5g

0

0

14)

3.8g

180

47.31

15)

3.8g

0

0

16)

3.8g

16

4.21

17)

3.9g

79

20.26

18)

3.9g

31

7.95

19)

3.9g

10

2.56

20)

4.0g

4

1

�-57January,
JOB PROGRESS

REPORT

State of

COLORADO
------~~~~~--------

Project

No.

W-4l-R-27

Bighorn Sheep &amp; Mountain

Work Plan No.

3

Job No.

Job Title

and Translocating

Bighorn Sheep

Period

Trapping

Covered:

Personnel:

1978

Goat Investigations

1

--------------------------------

June 1, 1976 through May 31, 1977

Robert L. Schmidt, Paul H. Neil, V. Feuerstein, M. Connors,
M. Middleton, P. Yates, J. Morris, F. Colley, R. Mason, and
B. Widhalm.

ABSTRACT
Bighorn ewes on Pikes Peak and the Poudre Canyon which were treated with
therapeutic drugs (Carnbendazole - Merck &amp; Co.) in apple pulp had 94 lambs
per 100 ewes. Rampart ewes treated with the therapeutic drug Fenbendazole
(Hoechst-Rossel) had 60 lambs per 100 ewes, but larval output in the feces
remained negative ·for 8 months.
A total of 112 bighorn sheep were trapped
and transplanted from 5 herds and implanted into 6 new areas. A total of
978 sheep were treated with therapeutic drugs from 9 different herds.

��-59-

TRAPPING

AND TRANSLOCATING

BIGHORN SHEEP

Robert L. Schmidt

P.N.O. OBJECTIVE
The principle objective of this study will be to increase the total number
of bighorn sheep within the State of Colorado by either introducing bighorn
into areas which were historically bighorn sheep ranges and where no animals
are in evidence today, and by adding additional animals to small remnant herds.
By expanding bighorn sheep from crowded areas into other areas it is hoped that
they will be better able to utilize the range and increase their numbers.
Also
the introduction of new animals, especially rams, into remnant populations
should improve the reproductive capacity and livability of the lambs by improving
the gene pool and lowering the percentage of inbreeding.

SEGMENT OBJECTIVES
1.

Trap and translocate

between

150 and 200 bighorn

sheep.

2.

Treat all captured bighorn sheep with chemotherapeutic
trol of Protostrongylus sp. lungworm parasitism.

3.

Monitor all translocated bighorn sheep for the degree of parasitism by
collecting fecal pellet groups and checking them by standard laboratory
procedures.

drugs for the con-

METHODS AND MATERIALS

Baiting Procedures
Baiting sites should be in the areas that sheep are using during the time of
year that you intend to trap or treat sheep. The best areas appear to be close
to bedding grounds, preferably upwind.
If you intend to trap, a location fairly
flat and free of rocks must be found. Snow depths should also be considered in
snow country and tops of ridges should be looked at.
Three or four months prior to trapping or treating, white salt blocks should be
placed at all likely trap sites. Then at the time baiting is started, bait
only the sites where salt has had heavy use.
Apple pulp has been an excellent bait. It is the part of the apple which is
left after the juice has been pressed out for apple cider. Some companies
add sawdust, wood fiber, and rice hulls to the cider making process.
This type
of apple pulp has not been very successful.

�-60-

Apple pulp must be picked up fresh from the cider mill, placed on concrete
or plastic to retain the juice. All air must be tramped out as you would
in corn silage.
The deeper it is piled the better.
Before using the pulp
for bait it should ferment for at least two weeks, with the supply being
close to the baiting area. In cold winter months it will keep about a week
on a pickup after being removed from the pile, however, in warm weather only
about two days.
If pulp is to be held for a longer period after removing
from the pile it should be bagged and frozen.
When starting to bait an area, three types of bait are used; apple pulp,
white block salt, and good green alfalfa hay. Place some of the pulp on
and around salt bloc-ks as well as on the alfalfa.
Sheep do not know what
apple pulp is and must come in contact with it to get them "hooked".
This
should take about three weeks.
If large numbers of sheep are in the area,
put out about 2 lbs of apple pulp per sheep per day to hook them.
A good example is: If a hundred sheep are in the area and you bait with
40 lbs of apple pulp you will only hook about twenty sheep. When starting
to bait, 40 lbs is about the .right amount, then as more sheep come, increase
the bait in proportion to the additional numbers of sheep coming.
For every
pound of pulp used, use a lb of alfalfa at the bait station.
Sheep that are
on apple pulp will also eat big sage and other vegetation surrounding the bait
station.
Alfalfa will supplement the apple pulp and their requirement for
roughage.
If permitted an adult sheep will eat about 5 lbs of apple pulp per
day; about 3-1/4 lbs in 30 minutes in the forenoon, then return in the afternoon and consume another 1-3/4 lbs.
Sheep that are hooked will come to the bait station every morning,
the run. At this time you are ready to trap or treat the sheep.

Treating

usually on

Procedures

Treatment of bighorn sheep has been accomplished by two separate methods.
Drugs may be administered either by the addition of drugs to apple pulp or by
orally drenching individual animals after trapping.
Drugs were administered to the sheep at the trap after they had been removed
from the net and tied. Both drugs were kept in a heated vehicle until usage
to prevent freezing, as freezing is known to severely reduce effectiveness.
The drug suspension was drawn up into a 50 cc syringe, a test tube attached
and the solution was expressed into the throat, after inserting the test tube
along the cheek past the molars.
Gentle stroking of the throad aids in swallowing
and helped prevent animals from holding the drugs in the mouth and later spitting
them out.
Administraiton of drugs without trapping was accomplished by mlxlng drugs with
apple pulp at the bait station.
Pre-mixing of drugs and apple mash was not
practiced due to wastage.
The amount of pulp is reduced when treating to help
insure the consumption of the entire dose of drug, 3 lbs/adult, 2 lbs/yearling,
lIb/lamb.
A small top loading spring scale (graduated in lbs) and a trash can

�-61-

lid are used to weigh the pulp. The drug is mixed thoroughly with the
weighed pulp (in the lid) by hand. using rubber gloves, and put out on the
bait station.
Putting the medicated pulp in 1-2 dose piles helps prevent
bunching and uneven distribution of drugs.
Since it is known that sheep
commonly feed in the morning and afternoon it was shown best to complete
treatment in the mornings to reduce double treatment.
Sheep were monitored
during treatment at a distance to insure that more drug was put out for newcomers.
Two drugs have been successfully utilized for treatment, Cambendazole (Merck
&amp; Co.) and Fenbendazole (Hoechst-Rossel).
Cambendazole has been proven
effective against transplacentally transmitted larvae in the fetus, when
administered in pregnancy.
Cambendazole is marketed as Cam-Vet, a 3 percent
active solution which due to the addition of a preservative sorbic acid has
proven unpalatable to sheep. We have centrifuged the 3 percent solution to
yield the concentrated drug + water and found this to be highly palatable.
The adult dose of this concentrated solution is 30 cc (8.4 grams drug), 20 ccl
yearling (5.6 grams drug), 10 cc/lamb (2.8 grams drug).
Fenbendazole (Hoechst-Rossel) has been shown to be effective against adult
Protostrongylus Spa to a greater extent than Cambendazole (Rampart Range 1976).
Fenbendazole is available in a 10 percent active solution.
The dosage is 20
mg/lb body weight or 30 cc/adult sheep, 20 cc/yearling, 10 cc/lamb.
Two separate treatments have proven much more effective, however, both drugs
have not been used simultaneously.
If both drugs were used Fenbendazole
was given first, followed in 10 days by treatment with Cambendazole.
Two
treatments with Fenbendazole were also given 10 days apart. Two treatments
with Cambendazole were given 3 weeks apart due to a resistance shown by sheep
to the second treatment.
Also, it was not uncommon for sheep not to return
to bait the day following treatment.

Trapping

Procedures

The trap net was set up over the preferred bait site at least a week prior to
trapping operations to allow sheep to become accustomed to going under the net.
The initial reaction of the sheep was variable, but in those herds which were
solidly hooked on apple pulp reluctance soon disappeared.
After sheep come to bait regularly under the net we use a 70 ft square drop
net. The trap weighs approximately 280 lbs for the net plus the poles. With
an experienced crew the trap will handle 15 to 25 sheep. Start with 5 to 8
sheep. The net is in four 35 ft squares that snap together making it easier
to handle, which is important in getting sheep out of the trap.
We use a radio-controlled detonator to drop the trap, which gives considerable
flexibility and does not require a man at one point at all times. The detonator now being used has been quite successful, not having had a fake drop
in 3 years. The range is over 2 miles, however you should stay as close as
possible as it is important to get to the dropped trap immediately.
The trap is designed to be set up on frozen ground and can be set up in
approximately an hour and a half.
It withstands extremely high winds, provi ding all steel anchor posts are wrapped with burlap at points where anchor

�-62-

ropes are to be tied. Failure to wrap posts results in broken
the collapse of the net.

ropes and

The four corner posts must be set first and squared at 76 feet. Next the
four anchor posts are set and wrapped with burlap.
The anchor post must
be set in exact line with the corner post and the opposite corner post.
Now snap the net together and tie in four corner blasting ropes. Blasting
rope should have a Bolen loop on one end, which can be run through net
corner ring and blasting rope run back through it. The other end of the
blasting rope can be half hitched to loop in end of pulley rope. A total
of 8 blasting cap ropes are used on the trap and all attached in the same
manner.
When corner blasting rope is attached, wire the blasting cap wires
to the detonator wire and pull corners tight. There should be about one to
two-foot of rope between pulley and blasting cap rope. Now tie in side
anchor ropes with blasting cap ropes. One side anchor rope, the up-hill
one, does not have a blasting cap. The side anchor post should be 20 ft
from line between corner posts. Corner anchor post should be at least 15
ft from corner post. The further all anchor posts are away from the net,
the better they hold.
Set side anchor post in line with snap line, then
tie in center blasting rope. Put up side braces, making sure side brace
without blasting rope is not standing straight up. Put up center pole and
tie slack rope to corner post. They should be tight for up to 7 sheep and
have 6 to 8 ft of slack for 20 to 25 sheep.
After trap is up, bait should be placed in a circle one-half way between
center pole and side of net with bait line from center bait to a point 10
ft outside of net. Two or 3 lines should lead out the way you think sheep
will come in from; one of these should lead out the down wind side. It is
best to allow sheep to come under the net for 2 or 3 days in large sheep
populations before trapping as the less wary sheep will encourage others to
come under the net.
When you are ready to trap and the sheep begin coming into the net they will
tend to group causing problems as the sheep should be evenly distributed
under the net. Before they can be evenly distributed you may find you have
more sheep than the trap will handle.
If this occurs, move in so that some
or all will leave. If they have not been there more than 10 to 15 minutes
some will come right back under. This time stay closer to the net so some
sheep will stay back.
The total time the sheep will be under the net is 30
minutes and it is not a good idea to drop the net in the sheep's face if you
intend to trap them later.
When taking sheep out of the trap a 5-man crew works best; four men to remove the sheep from net and 1 man assigned to observing sheep in the net.
One of the important things to watch for is a sheep with its head turned
back along the body so tight that it is unable to breathe or is lying on top
of its head.
These sheep must be straightened out at once. The observer
can also tell the crew which sheep should be removed from the net first.
Snaps in the net are extremely helpful in getting the sheep out. Sheep should
be removed by pulling them out by the back legs, not by the horns or head as
t hf,s could cause neck damage.
Hog-tieing works well when collaring and treating sheep for release, and temperature is not too cold. When hog-tied a burlap
bag should be placed over the head.
Sheep that are to be transported can be
removed from the net and walked to a truck by one man holding up one back leg
and one man walking along side the neck with a hand on each horn so as to guide
the sheep.

�-63-

In order to facilitate the larger number of trap sites three complete
nets were assembled, and two detonators purchased.
The additional nets
allowed more than one preferred bait station or more than one area to be
worked at one time.

RESULTS AND DISCUSSION

Poudre·Canyon

and Pikes Peak

All of the bighorn ewes collared in the Poudre treatment area (both this
year and years previous) were treated at least once with drugs in apple
pulp. Observations conducted between November 15, 1976 and December 15,
1976, reported 94 lambs per 100 ewes. Eighty-seven percent of the total
ewes ever collared on Pikes Peak were also treated at least once with drugs
in apple pulp. A similar percent of the unco11ared ewes in the Pikes Peak
area were treated along with the collared ewes. Observations conducted
from November 15, 1976 to January 15, 1977, reported more than 100 lambs
per 100 ewes (Morris 1977). The large 1amb:ewe ratio is thought to be to
old age mortality of ewes between September and January.

Rampart Range
Fenbendazo1e

Trial Results

Bighorn ewes on Rampart Range were treated three times with the therapeutic
drug Fenbendazo1e in April 1976. Subsequent monitoring of larvae per gram
output in the ewes reported larval output to decline to negative and remain
negative for 8 months.
Field observations conducted from November 1, 1976
to January 15, 1977 reported 60 lambs per 100 ewes.

LITERATURE
Morris,

J.

1977.

Personal

CITED

communication.

Colo. Div. Wildlife,

S.E. Region.

�Table 1.

Trapping, treating, transplant and re-re1eased data of bighorn sheep, June 1976 to June 1977.

Pulp

Net

Treated
Cambendaz ole

Fenbendazo1e

Trapped

Released

Poudre Canyon

187

29

104

108

30

4

25 11

1

Georgetown

16

-

16

Mt. Evans

84

38

84

37

38

17

20

u

1

Grant

83

23

106

-

23

23

Tarrya11

115

20

115

20

41

21

20 11

Pikes Peak

182

29

117

94

51

31

19 !il

1

Trickle Mtn.

64

48

83

29

49

20

19 :2./

1

Basalt

23

-

23

-

-

-

Rampart

38

-

19

19

Area

11

Apishapa Canyon.

]j
Cross Mountain.

11
Cow Creek.

!il
Soap Creek.

il
Lamb property.
6/

Cebolla Creek.

Transplanted

9 2..1

Fatality

I

.,..

0\

I

�-65-

Table 2. Identification numbers of bighorn
River, February 9, 1977.
Collar No. 1/

])

sheep transplanted

* to Apishapa

Sex

Age

Trap Site

30

F

Lamb

Poudre Canyon

31

M

1-1/2

Poudre Canyon

32

M

Lamb

Poudre Canyon

33

M

Lamb

Poudre Canyon

34

F

Lamb

Poudre Canyon

35

M

1-1/2

Poudre Canyon

39

F

Lamb

Poudre Canyon

M

1-1/2

Poudre Canyon

20

F

10+

Poudre Canyon

21

F

1-1/2

Poudre Canyon

22

F

4

Poudre Canyon

23

F

10+

Poudre Canyon

24

F

3

Poudre Canyon

25

F

1-1/2

Poudre Canyon

26

F

1-1/2

Poudre Canyon

27

F

2-1/2

Poudre Canyon

28

F

6

Poudre Canyon

29

F

7+

Poudre Canyon

30

F

5

Poudre Canyon

31

F

2-1/2

Poudre Canyon

?

F

1-1/2

Poudre Canyon

2

F

5

Poudre Canyon

3

F

6

Poudre Canyon

Ear Tag No.

21/

*Total number of sheep released was 25; eartags of two sheep were not recorded at the trap site, they were believed

to be 37, 38.

~/Yellow

collar with black numbers.

l/White

eartag with black numbers,

placed in right ear.

l/Yellow

eartag with black numbers

placed in right ear.

�-66-

Table 3. Identification numbers of bighorn
in Poudre Canyon, February 9, 1977.

};/

l/White

Site

Age

M

6

Meadows

F

7+

Meadows

36

M

3-1/2

Meadows

?

M

3

Meadows

Red 8
Orange/black

Release

Sex

Ear Tag No.

Collar

sheep trapped and released

stripe

ear tag with black numbers placed in right ear.

Table 4. Identification
Creek, March 31, 1977.

numbers

of bighorn

]j

Sex

Age

5

M

Lamb

Trickle Mountain

7

M

Lamb

Trickle Mountain

35

F

1

Trickle Mountain

36

F

1

Trickle Mountain

37

F

3

Trickle Mountain

38

F

4

Trickle Mountain

39

F

4

Trickle Mountain

R a d·10-3/

F

3

Trickle Mountain

Ra d·10-4/

F

3

Trickle Mountain

Collar No. 1/

l/White

Ear Tag No.

Sheep transplanted

collars with black numbers.

2:.../ Red ear tag with black numbers placed in right ear.
l/Telemetry

4/

- Telemetry

collar, yellow with red stripe.
collar, black.

to Cebolla

Trap Site

�-67-

Table 5. Identification numbers of bighorn sheep transplanted to Cow
Creek, March 17, 1977.
Collar No. l/

l:/

Sex

Age

Trap Site

49

M

1

Tarryall

50

F

Lamb

Tarrya11

51

M

Lamb

Tarryall

52

M

Lamb

Tarryall

53

M

Lamb

Tarryall

4

F

1

Tarryall

5

F

3

Tarryall

6

F

2

Tarryall

7

F

5

Tarryall

8

F

4

Tarryall

9

F

6

Tarryall

10

F

5

Tarryall

11

F

4

Tarryall

12

F

2

Tarrya11

l3

F

3

Tarryall

14

M

2

Tarryall

15

F

6

Tarryall

16

F

5

Tarrya11

17

F

5

Tarrya11

18

F

4

Tarryall

l/

Ear Tag No.

White collar with black numbers.

2/
White eartag with black numbers placed in right ear.

�-68-

Table 6.
Mountain,

Identification numbers
January 26, 1977.

Collar No. 1./

Ear Tag No.

of bighorn

2/

Sex

sheep transplanted

Age

to Cross

Trap Site

7

F

Lamb

Mt. Evans

8

M

Lamb

Mt. Evans

9

F

Lamb

Mt. Evans

10

,.
M

Lamb

Mt. Evans

11

M

Lamb

Mt. Evans

12

M

l~

Mt. Evans

13

M

l~

Mt. Evans

14

M

l~

Mt. Evans

4

F

4

Mt. Evans

5

F

5

Mt. Evans

6

F

6+

Mt. Evans

7

F

3

Mt. Evans

8

F

5+

Mt. Evans

9

F

3

Mt. Evans

10

F

3+

Mt. Evans

11

F

2

Mt. Evans

12

F

7

Mt. Evans

13

F

3

Mt. Evans

14

F

3

Mt. Evans

15

F

4

Mt. Evans

1/ Yellow collars with black numbers.

2/ White ear tags with black numbers, placed in right ear.

�-69-

Table 7· Identification numbers
property March 31, 1977 •
Collar No. 1/

Ear Tag No.

of bighorn sheep transplanted

]:./

to Lamb

Sex

Age

Trap Site

68

M

Lamb

Trickle Mountain

69

F

Lamb

Trickle Mountain

72

M

Lamb

Trickle Mountain

73

M

Lamb

Trickle Mountain

74

M

1

Trickle Mountain

75

M

1

Trickle Mountain

11

F

6

Trickle Mountain

12

F

6

Trickle Mountain

14

F

6

Trickle Mountain

15

F

1

Trickle Mountain

16

F

1

Trickle Mountain

17

F

2

Trickle Mountain

81

F

5

Trickle Mountain

19

F

5

Trickle Mountain

20

F

5

Trickle Mountain

21

F

5

Trickle Mountain

23

F

6

Trickle Mountain

25

F

3

Trickle Mountain

Radio

F

2

Trickle Mountain

1./
Orange collars with black numbers.

2/
White eartags with black numbers, placed in right ear.

�-70-

Table 8. Identification numbers of bighorn
at Grant, Colorado, April 14 and 20, 1977 •
1/
Collar No. -

Ear Tag No.

sheep trapped and released

Sex

Age

Release Site

3 2:/

M

Lamb

Grant

4 Jj

F

Lamb

Grant

5 J:j

F

Lamb

Grant

M

Lamb

Grant

M

Lamb

Grant

F

Lamb

Grant

M

Lamb

Grant

M

Lamb

Grant

M

Lamb

Grant

M

Lamb

Grant

16l/

F

Lamb

Grant

17 1/

M

Lamb

Grant

181/

M

Lamb

Grant

19 1/

F

Lamb

Grant

38

F

4

Grant

39

F

1

Grant;

42

F

3

Grant

43

F

4

Grant

44

F

1

Grant

45

F

4

Grant

46

F

1

Grant

47

F

3

Grant

R ad I
10-4/

F

5

Grant

6 J:j

7 2:../
8 2:../
9 2:../
lO 2:../
11 2:../
12 2:../

1/
- Orange collar with black numbers.
2/
- Orange eartag with black numbers placed
3/
- Yellow

in right ear.

eartag with black numbers placed in right ear.

4/
- Telemetry

collar,

green with orange stripe.

�-71-

Table 9. Identification numbers of bighorn sheep trapped and released
Mt. Evans, January 25 and February 1, 1977.
Collar No. 1/

Ear Tag No. l:./

on

Sex

Age

Release Site

1

M

l~

Ht. Evans

2

M

l~

Mt. Evans

3

F

Lamb

Mt. Evans

4

F

Lamb

Mt. Evans

5

M

3

Mt. Evans

6

M

2

Mt. Evans

15

M

5

Mt. Evans

16

F

Lamb

Mt. Evans

18

F

Lamb

Mt. Evans

19

F

2

Mt. Evans

20

F

Lamb

Mt. Evans

1

F

3

Mt. Evans

2

F

3

Mt. Evans

3

F

3

Mt. Evans

16

F

l~

Mt. Evans

17

F

5

Mt. Evans

19

F

5

Mt. Evans

II Yellow collar with black numbers.
~I White ear tag with black numbers, placed in right ear.

�-72-

Table 10. Identification
Creek, March 8, 1977.

Collar No. ]j

numbers of bighorn

Ear Tag No. ]j

sheep transplanted

Sex

Age

Trap Site

40

M

Lamb

Pikes Peak

41

F

Lamb

Pikes Peak

42

M

Lamb

Pikes Peak

43

M

Lamb

Pikes Peak

44

M

Lamb

Pikes Peak

45

F

Lamb

Pikes Peak

M

1

Pikes Peak

32

F

1

Pikes Peak

33

F

1

Pikes Peak

34

F

Pikes Peak

35

F

Pikes Peak

36

F

Pikes Peak

37

F

Pikes Peak

38

F

Pikes Peak

39

F

Pikes Peak

40

F

Pikes Peak

41

F

Pikes Peak

42

F

Pikes Peak

43

F

11

]j

2

1/ Yellow collar with black numbers.

2/

to Soap

White ear tag with black numbers,

1/ Yellow

ear tag with black numbers,

placed in right ear.
placed in right ear.

Pikes Peak

�-73-

Table 11. Identification numbers of bighorn sheep trapped and released
at Tarrya11, March 17, 1977 .
Collar No. II

Ear Tag No. Jj

Sex

Age

Release Site

49

M

2

Tarrya11

50

F

Lamb

Tarrya11

51

M

Lamb

Tarrya11

52

M

Lamb

Tarryall

53

M

Lamb

Tarrya11

54

M

4

Tarryal1

55

M

3

Tarrya11

56

M

3

Tarrya1l

57

M

3

Tarrya11

58

F

Lamb

Tarrya11

59

M

Lamb

Tarrya11

60

M

2

Tarryall

61

M

1

Tarryall

62

M

4

Tarryall

63

M

5

Tarryall

4

F

1

Tarryall

5

F

3

Tarryall

6

F

2

Tarryall

8

F

4

Tarryall

9

F

6

Tarryall

10

F

5

Tarryall

11

F

4

Tarryall

12

F

2

Tarrya1l

-------------------------------------------------------------------------

�-74-

Table II. Identification numbers of bighorn sheep trapped and released
at Tarryall, March 17,1977 (continued).
Collar No. 1/

Ear Tag No. ]j

Sex

Age

Release Site

13

F

3

Tarryall

14

M

2

Tarryall

15

F

6

Tarryall

16

F

5

Tarryall

17

F

5

Tarryall

18

F

4

Tarryall

19

F

4

Tarryall

20

F

5

Tarryall

21

F

5

Tarryall

22

F

5

Tarryall

23

F

5

Tarryall

24

F

5

Tarryall

25

F

6

Tarryall

26

F

3

Tarryall

27

F

5

Tarryall

28

F

4

Tarryall

29

F

7

Tarryall

1/

White collar with black numbers.

]j
White eartag with black numbers placed in right ear.

�-75-

Table 12. Identification numbers of bighorn sheep trapped and released
on Trickle Mountain, April 4 and 8, 1977 .
Collar No. II

J:./

Sex

Age

Release Site

76

M

Lamb

Trickle Mountain

77

F

Lamb

Trickle Mountain

78

M

Lamb

Trickle Mountain

79

M

Lamb

Trickle Mountain

80

M

Lamb

Trickle Mountain

13

F

6

Trickle Mountain

22

F

5

Trickle Mountain

24

F

3

Trickle Mountain

26

F

2

Trickle Mountain

27

F

1

Trickle Mountain

28

F

4

Trickle Mountain

29

F

5

Trickle Mountain

30

F

7

Trickle Mountain

31

F

5

Trickle Mountain

32

F

4

Trickle Mountain

33

F

3

Trickle Mountain

34

F

3

Trickle Mountain

35

F

3

Trickle Mountain

36

F

4

Trickle Mountain

37

F

4

Trickle Mountain

Ear Tag No.

l/orange collar with black numbers.
J:./Whiteeartag with black numbers placed in right ear.

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                  <text>April 1978

JOB PROGRESS

State of

COLORADO
--------------~~-----------

Project

No.

Work Plan No.
Job Title
Period Covered:

REPORT

Game Bird Survey

W-37-R-3l

9
Job No.
Evaluation of the Effects of Changes
~i~n~H~u~n~t=i~n~g~R~e~g~u~l~a~t==i~o~n~s~o~n~S~a~g~e=_~G~r~o~u=s~e~P~o~p~u==l=a~t
_

3

---------------------------------

April I, 1977 through March 31, 1978

Personnel:
Fred Giese, U. S. Fish and Wildlife Service; Layne Adams, Don
Benson, Clait Braun, Deborah Covic, Courtney Crawford, Steve Emmons, Sue
Erickson, Howard Funk, Ken Giesen, Don Gore, John Hobbs, Jim Jackson, Bob
Leasure, Tom Lynch, Sig Palm, Brett Petersen, Steve Porter, Barbara Quinlan,
Howard Spear, John Wagner, Ron Zaccagnini, Colorado Division of Wildlife.
ABSTRAcr
Investigations concerning the relationships of hunting to breeding population
levels, and mortality and survival rates of sage grouse (Centrocercus
urophasianus) in North Park, Colorado initiated in 1975, continued in 1977.
Numbers of known leks increased as five new leks were located.
Average
number of cocks per lek remained constant.
Total estimated spring population
$ize was about 5,780 birds.
In 1977, 560 (313 males, 245 females) sage grouse
were banded.
Seventy-five broods were observed with a composite average of
3.2 chicks per successful hen. Estimated nesting success based on 127 females
seen during the brood period was 59 percent.
A 16 day permit only hunting
season was allowed in North Park in 1977 with bag and possession limits of 3
and 6. Data from wings (N = 632) collected at check stations, wing barrels
and field checks revealed that the harvest was comprised of 46 percent young
of the year, 19 percent yearlings and 35 percent adults.
Calculated hatching
dates for 290 hunter harvested juveniles ranged from late May to 12 July
with a peak between 10-15 June. Estimated nesting success from examination
of wings of adult and yearling females was 50 percent with more adults (59%)
than yearlings (33%) being successful.
Ovarian analysis (N = 89) indicated
that 96 (yearlings) to 98 percent (adults) of all females had ovulated.
Total permits issued numbered 1,093, and only 77 percent of the permittees
actually hunted.
Estimated total harvest including crippling loss was 1,070
grouse.
This was an increase of approximately 20 percent from 1976 harvest
levels.
Eighty-four bands were recovered and reported by hunters.
Survival
and mortality rates were calculated with adult males having lower survival
estimates than yearling males and all females.
Estimated fall population
size was about 7,600 birds with a harvest rate of about 14 percent.

�-2RECOMMENDATIONS

1.

Trapping and banding of sage grouse should be terminated in North Park
except to achieve objectives relating to limited areas (groups of leks)
and specialized studies.

2.

Counts of male
sage grouse present on leks in April and early May
should be continued indefinitely into the future. However, revision of
present procedures (4 counts during specified intervals of all leks) may
be warranted upon completion of this study.

3.

Counts of hens with broods along roadsides from vehicles and walking
brood transects in North Park have little merit and should be terminated.

4.

Collection of harvest data (i.e. wings) and proper analysis
continued indefinitely for management purposes.
Collection
be through manned check stations or volunteer wing barrels.

5.

The basic sage grouse season in North Park should be a minimum
days with bag and possession limits of 3 and 6.

6.

The special hunting

should be
of wings can

permit in North Park should be discontinued.

of 16

f'.

�-3-

EVALUATION OF THE EFFECTS OF CHANGES
IN HUNTING REGULATIONS ON SAGE GROUSE POPULATIONS
C1ait E. Braun

Sage grouse are widely distributed in western North America, occuping sagebrush dominated rangelands.
Presently they are hunted on a regular basis
in 9 western states and on an occasional basis in two other states.
States
which hunt sage grouse routinely collect different types of population data
and regulate hunter opportunity and harvest.
Management of sage grouse has
primarily been based on counts of cocks on leks in April and estimates of
nesting success and brood size obtained in July and August.
Unfortunately.
evaluation of existing data gathering procedures and knowledge of the impacts of hunting have been and are limited.
Data presented in this report
concern these problems and represent the period of mid-March 1977 through
mid-March 1978.
P. N. OBJECTIVE
The objectives of this study are to increase hunter opportunity and recreational use of sage grouse in North Park, Colorado.
Hypotheses which are
being tested are:
a.

Hunter harvest does not affect spring counts of strutting
spring counts of females under any regulations.

b.

Counts of females during the peak of attendance on strutting grounds
are a reliable index to overall numbers of sage grouse in spring.

c.

Hunter

d.

Nesting success and brood size to 15 August. while important in determining age composition of the harvest, have no relationship to hunter
success and total harvest.

e.

Differences
and nesting

success

and total harvest

exist between
success.

are a function

yearling

of August

males

or peak

precipitation.

and adult hens in percent

ovulating,

SEGMENT OBJECTIVES
1a.

Counts of males will be made on 10 selected leks three times each week
from 5:00 to 7:30 A.M. during the month of April.

lb.

Counts of females will be made on 10 selected
days in April (period of peak attendance).

2a.

Males will be trapped and marked with color-coded bands at night while
roosting on or near leks. A sample of 200 is desired.
Trapping will
be conducted from mid-March to late May by spotlighting and netting.
Bandings will be evenly distributed in four areas of North Park.

leks every day for 10

�-4-

2b.

Females will be trapped during the early A.M. on leks during April
through use of walk in, mist and cannon nets. A sample of 300 is
desired.
Bandings will be evenly distributed in four areas of North
Park.

2c.

Chicks will be located and trapped in brood areas during late July and
August through use of tape recorded calls and noosing.
Some chicks
may be caught through use of cannon nets and drive traps. A sample of
200 chicks (100 males and 100 females) is desired.
Bandings will be
evenly distributed in four areas of North Park.

3a.

Nesting success will be estimated through counts of successful and
unsuccessful hens in July and August.
Successful females will be
located through use of tape recorded calls, while unsuccessful hens
will be located and counted in known summer use sites. A sample of
200 hens is desired.

3b.

Brood size will be determined by counts of chicks accompanied
in July and August.
A sample of 100 broods is desired.

4.

August precipitation will be determined from 12 rain gauges placed at
selected sites throughout sage grouse habitat.
Sites will be selected
that typify particular areas within North Park. Each rain gauge will
be sampled once a week. Data collected will be compared to that
collected by the U. S. Weather Bureau in Walden and at Spicer.

5.

Population size will be estimated in spring and fall. Spring population size will be estimated through use of ratios of males to females
on leks, observation and recapture of marked birds and ratios of males
to females and juvenile to yearling to adult age classes in the harvest
the preceding fall. Fall (pre-hunting) population size will be estimated through examination of data on nesting success, brood size to 1
September and spring population data. Independent fall population
estimates will be made through use of data collected at check stations
on age composition of the harvest and number of marked birds shot.

6a.

Hunting will be by free permit only, and all hunters will be mailed a
questionnaire within one week of the end of the hunting season.
One
followup letter will be sent to all non-respondents three weeks after
the initial mailing.
A goal of 90 percent response is desired.

6b.

Check stations will be operated during the opening weekend at State
Line and Willow Creek Pass. Wing collection barrels will be placed at
Muddy Pass, Gould, Saratoga Hf.ghway , and Independence Mountain during
the entire season.
Collection barrels will also be available near
State Line and Willow Creek Pass when manned check stations are not in
operation.
A goal of 500 wings is desired.

6c.

Number and location of marked birds shot will be obtained through use
of check stations, field hunter checks and voluntary mail reporting.
A goal of 50 first year recoveries is desired.

by hens

�-5-

6d.

6e.

Hunter success will be determined through data collected
stations and from the mail questionnaire survey.

at check

Age composition of the harvest will be determined through examination
Classification will be as adult,
of wings collected at check stations.
yearling, juvenile, male and female.

6f.

Total harvest will be estimated (90 percent
mail questionnaire survey of all hunters.

7.

Regulations of up to three and six birds, bag and possession,
days (season length) will be requested in 1977.

8.

Compile data, analyze results,
METHODS

and prepare

confidence

progress

level)

from the

and 21

report.

AND MATERIALS

Counts of male and female sage grouse present on leks were periodically made
by project personnel and Wildlife Conservation Officers following proscribed
procedures (Braun and Beck 1976).
Counts were made from 22 March through 24
May at times within one-half hour of sunrise.
Searches for new or relocated
grounds were made through ground reconnaissance.
Male and female sage grouse
were trapped at night along roads and on leks where they roosted.
Some birds,
primarily hens, were captured at dawn on leks through use of cannon nets.
Chicks were captured at night or in the day where they roosted or hid. Most
birds were captured through use of a spotlight with a back pack or hand held
power source and long handled nets (Braun and Beck 1976).
Sage grouse banded
and released were marked with serially numbered aluminum bands (size 16 for
males, 14 for females) and colored plastic and aluminum bands coded to
capture locations and year. Age and sex classification of birds captured
followed Eng (1955).
Hens with broods were located during July and August through systematic
searches of selected areas and observations along roads and trails.
Systematic searches for broods involved walking moist areas and playback of tape
recorded chick distress calls (Braun et al. 1973, Wallestad 1970).
Searches
for broods along roads and trails were made from a vehcile following established routes (Gill 1965) and other suitable appearing areas.
Routes were
traversed shortly after sunrise at a speed of 15 to 20 miles per hour.
Sage grouse hunters in North Park in 1977 were required to have in their
possession a free numbered permit.
Permits unlimited in number were available from Division of Wildlife offices in Fort Collins and Denver, all
license agents in North Park, and all project and management personnel
assigned to work in North Park during the hunting season.
Questionnaires
were sent to all permittees immediately following the sage grouse season in
North Park, with one fo1lowup letter being mailed in mid-October to all
non-respondents.
Check stations were operated at two locations (State Line and Willow Creek
Pass) on the opening weekend (10 and 11 September) and the second Sunday (18
September).
Each station was operated from about 1000 to 1800 MDT, depending

�-6-

upon traffic load. Data obtained per party were: county of or1g1n, number
of hunters, hours hunted (total of all hunters), birds observed, birds
bagged, birds lost, number of banded birds and location of where each was
obtained, and area hunted.
One wing was obtained from each bird checked
with ovaries being taken or examined from a sample of hens.
Efforts were
made to ascertain sex by gonadal inspection for a sample of birds.
These
data were recorded on tags with wings being individually marked with corresponding information concerning actual sex of that bird. Wings were
frozen and stored for later analysis.
In addition to the two check stations, wing barrels and signs (Hoffman and
Braun 1975) were placed along Colorado 14 near Muddy Pass, near the State
Line along Colorado 125, along Colorado 14 near Gould, at the junction of
Jackson County roads 6 and 34, north of the junction of Colorado 125 and
127 along Colorado 127 and at Willow Creek Pass along Colorado 125.
Volunteer stations were in operation the entire season near Muddy Pass
and Gould, on all days that manned check stations were not operating on
Colorado 127 north of its junction with Colorado 125 and along Colorado 125
at Willow Creek Pass and for the opening weekend only at the junction of
Jackson County roads 6 and 34 and on the Saratoga Highway (Colorado 125).
Collected wings were thawed and classified to age (chicks, yearlings, adults)
and sex following procedures outlined by Eng (1955), and Beck et al. (1975).
Hatching dates were calculated for chicks of the year using data from pyrah
(1963).
Ovaries collected were stored in AFA (alcohol, formalin and acetic acid)
and later examined for presence of ovulated follicles as described by Kabat
et al. (1948).
Description

of Area

North Park lies completely in Jackson County, Colorado.
This relatively
high altitude (7,800 to 8,500 ft) area is essentially a closed basin, as
it is encircled by mountain ranges.
Drainage is to the north, with lowest
elevations occurring north of Cowdrey.
Vegetation is principally a sagebrush (Artemisia spp.)-grassland
type with stream courses being dominated
by native grasses, sedges and deciduous shrubs.
Detailed geographic,
geologic, vegetational and climatic features of the area have been described
by Gill (1965), Carr (1967), Beck (1975), Braun and Beck (1976) and will be
treated in the final report.
RESULTS

AND DISCUSSION

Counts of Sage Grouse on Display

Areas

Counts of male and female sage grouse present on leks were initiated on 22
March (Alkali Lake, Bighorn, Boettcher Lake Junction, Hound, Walden and
Wattenburg #2) and continued until 24 May (Bighorn and Boettcher Lake
Junction).
During this interval 357 counts were made of 26 active leks.
Maximum counts of males and females and dates obtained are presented in
Table 1. The total of 809 males counted is higher than the 669 counted in

�-7-

Table 1.

Peak lek counts of sage grouse, North Park, 1977.

Lek

Males

Date(s)

Females

Date(s)

Alkali lake

36

25 April

49

8 April

Arapahoe

46

10 May

3

23 April

Aspen

21

17 May

1

26 April, 17 May

Bighorn

52

20 April,
6 May

54

23 April

Boettcher Lake
Junction

76

23 April

35

13 April

Canuck

27

22 April

15

7 April

Coalmont

21

15 April

29

13 April

Cowdrey #5

5

8 April

0

All Dates

Deer Creek

31

22 &amp; 28 April

17

8 April

Denmark

58

3 May

10

23 &amp; 24 April

Fish Hatchery

64

17 May

59

7 April

Hound

21

6 April

28

13 April

Lost Creek III

47

6 May

30

12 April

Monahan Draw

1

6 &amp; 22 April

2

6 April

Owl Creek

16

8 April

11

15 April

Pronghorn

15

24 May

0

24 May

Railroad

41

17 April

52

7 April

Raven

73

23 May

5

23 May

Ridge Road

32

6 April

34

12 April

Riley

9

16 April

10

12 April

Roth

8

6 April

0

All Dates.

Spring Creek III

45

10 May

42

13 April

Spring Creek 112

23

20 &amp; 26 April

15

7 April

Spring Creek 114

3

21 &amp; 27 April

1

15 April

Walden
Reservoir

26

2 April

55

6 April

Wattenburg #2

12

29 April,
9 May

7

4 April

Total

809

564

�-8-

1976 but represents 5 new leks (Arapahoe, Aspen, Denmark, Pronghorn, and
Raven).
Two other small display areas (Case Flats = 3 males and Badger
9 males) were not listed as permanent leks even though they have been
sporadically active for several years.
Trends in numbers of male sage grouse counted from 1973 through 1977 are
presented in Table 2. Average number of cocks counted per lek has remained
consistent through the 5 year period although number of known leks has
increased from 17 to 26. During this same interval only 2 known leks have
been abandoned by sage grouse.
These data strongly suggest that either
numbers of sage grouse in North Park are increasing or project personnel
are becoming more knowledgeable and efficient.
It is probable that additional grounds remain to be located.
In the fall of 1976 a coal haul road was constructed adjacent to the Roth
lek. During the 1977 breeding period traffic by the ground started at
5:30 A.M. MST with an average of 11 large trucks per hour traversing the
road during the breeding interval.
It is expected that this ground will
be abandoned before 1978.
The importance of the counts of females on leks (total = 564) in 1977 is
unknown.
This number is less than the counts of 573 in 1975 and 599 in
1976. Due to logistic problems of counting all leks on the same days
during peak of mating and proper interpretation, it is probable that counts
of hens will not be useful for other than estimating onset of nesting and
expected appearance of broods.
The peak period of female attendance in
1977 was between 6 and 13 April.
These dates are almost identical to those
observed in 1976 (5-15 April).
Analysis of daily and weekly changes in counts of male sage grouse on leks
for the 1973 -1977 period have been initiated and pertinent data will be
prepared for publication.
These data will be published in a technical
journal article.
Estimation

of Spring Population

Size

Reliable estimates of total numbers of sage grouse in North Park during the
breeding period are difficult to derive.
Major problems involved are those
relating to how many males and females of the total population are present
on leks when peak counts are attained and how many of the actual number of
active leks are known and counted.
There is some evidence to suggest that
only 50 percent of the male segment of the population of lekking species
is present on a ground when peak counts of males are attained (Robel 1969,
Rippin and Boag 1974).
If it is assumed that peak counts of male sage
grouse represent 50 percent of the male population in an area, then at
least 1,618 (809 x 2) cocks occurred in North Park in the spring of 1977.
It is quite obvious that all lek locations are not known in a given year.
If it is assumed that 80 percent of the active leks have been located, then
a minimum of 5 (.20 x 26) unknown leks should occur somewhere within North
Park. Assuming that these 5 grounds have an average of 31 cocks during

�-9-

Table 2.

Peak counts of male sage grouse on leks, North Park 1973-1977.

AVERAGE
GROUND
ALKALI LAKE
ARAPAHOE (1977)*
ASPEN (1977)
BIGHORN (1976)
BoETTCHER LAKE JUNCTION
BoETTCHER FLATS
&lt;1963-1970)
CANUCK (1974)
COAU1ONT
COWDREY #1
&lt;1968-1971&gt;
COWDREY 113
&lt;1963-1971&gt;
COWDREY #5 (1973)
DEER CREEK
DELANEY BUTTE
(1970-1972)
DENMARK &lt;1977&gt;
(1970-1972)
FISH HATCHERY
HOUND (1974)
LAKE JOHN #2
&lt;1968-1971)
LOST CREEK #1
LOST CREEK #2
MONAHAN DRAW
OwL CREEK (1976)
PRONGHORN (1977)
RAILROAD (1975)
RAVEN &lt;1977&gt;
RIDGE ROAD
RILEY (1973)
ROTH &lt;197LI)
SPRING CREEK #1
SPRING CREEK #2
(1968, 1970-1972)
SPRING CREEK #4
WALDEN RESERVOIR (1973)
&lt;1963-1969)
WATTENBURG #1
WATTENBURG #2

1973

1974

1975

1976

1977

77.3

31

72

68

39

36.6
7.0

59
0

39.4
9.2
5.0
56.6
13.3

117
0
0
30
37
4

46
0
29
27
0
0
19
11
13

50
0
22
29
0
0
9
27
0

46
62
0
30
28
0
0
5
36
0

53.7

67

13.5
54.4
27.4
56.2

0
13
10
11

62
69
0
0
1
3

69
33
0
13
0
6

81
27
0
33
0
1
14

36

37

27
12
14
49
14
3
34
0
18

32
18
9
49
11
3
37
0
16

36
46
21
52
76
0
27
21
0
0
5
31
0
58
64
21
0
47
0
1
16
15
41
73
32
9
8
45
23
3
26

76.4

36
12

37.6
75.8
13.7

46
39
9
38

1.5
27.2

0

24

33
15
10
33
15
10
37
0
22

0

12

- (19) 563&lt;17&gt; 527(19) 588(19) 669(21) 809(26)

TOTALS (H)
AVERAGE/AcTIVE

1968-1972

GROUND

39.1

33.1

27.7

30.9

31.9

* Year of initial location is given next to name of all recently
located grounds.

31.1

�-10-

the peak period of male attendance, they represent a minimum of 310
additional cocks (31 x 5 = 155 x 2 = 310). Thus the total estimated
number of cocks in North Park in spring 1977 was 1928. This is higher than
the 1,486 cocks estimated to have been in North Park in 1975 and the 1,682
cocks estimated in 1976. All available data suggest that these estimates
are realistic and that the apparent increase in the population is real.
Beck (1977) found a late winter sex ratio of 69 hens to 31 cocks, a value
remarkably similar to the sex ratio of adult and yearling birds in the fall
harvest (Braun and Beck 1976).
If it is assumed that the spring sex ratio
of the North Park sage grouse population is similar to that observed in
late wiriter and early fall, then 2 hens occur for every cock in the population.
Thus, the total spring female population can be estimated based
on the estimated number of cocks in the spring population.
Therefore,
there should have been a minimum of 3,856 hens in the 1977 spring population (1,928 x 2 = 3,856).
Thus, the estimated total spring population
of sage grouse within North Park in 1977 was 5,784 birds if actual projections are used.
This is substantially higher than the 5,046 birds estimated in 1976.
It is reasonable to assume that the 1977 spring sage grouse
population in North Park was between 5,000 to 6,000 birds.
Capture and Banding
Intensive efforts to capture and band sage grouse in the 1977 field period
were initiated on 21 March and continued through 24 May. Efforts to locate,
trap and band young of the year were initiated in early July and continued
until 9 September.
Two additional hens were banded in December.
Age and
sex data for the banded sample are given in Table 3.
The total of 560 birds banded in 1977 was the highest number since intensive
trapping efforts were initiated in 1973. This included the highest number
of females and chicks in the 5 year period.
Greater success in 1977 was
related to excellent access (the winter of 1976-77 was mild with little
snow), improvement in a power source for trapping along roads, use of two
different crews each night during a 10 day period, and trapping effort each
night from mid-March to mid-May.
As in earlier years, distribution of bandings within North Park was not
uniform as trapping effort paralleled distribution of known leks.
Since
relatively few leks were known to occur in the northeast quarter of the
Park until mid-May, only 66 of the 529 birds banded during the breeding
period were in this area. The same was partially true for the southeast
quarter of the Park where 95 birds were banded in the March-May interval.
During this same interval 174 birds were banded in the northwest quarter
while 194 were banded in the southwest quarter of the Park.
Composition of the banded sample of adults and yearlings does not reflect
age and sex composition of the population.
Adult males are underrepresented in the sample as many on known leks were banded in previous
years.
Males are easier to locate than females due to their tendency to
roost at night on leks. Thus females are also under-represented
in trap

�-11-

samples.
The age composition of adult and yearling females may approach
reality as harvest data suggest that adult hens comprise about 60 percent
of the fall population of hens older than one year. No selection for age
class of hens or differences in overall behavior between age classes is
known to occur provided that trapping effort is uniform throughout the
breeding period.
Sample sizes of chicks were too small to evaluate
although both trapping and harvest data suggest there are slightly more
chick hens in the fall population than chick males.

Table 3. Sex and age composition
North Park, Colorado 1977.

of newly banded sage grouse by month,

1+

Males
2+

Total

March

11

16

April

134

~y

Month

Chick

1+

Females
2+

Total

27

6

13

19

66

200

80

102

182

37

36

73

14

14

28

Chick

July

4

0

3

7

1

0

0

1

August

2

1

2

5

9

1

4

14

September

1

0

0

1

1

0

0

1

December

0

0

0

0

0

0

2

0

Totals

7

183

123

313

11

101

135

245

During the course of trapping operations in North Park in 1977, 72 different
sage grouse were recaptured.
This included 20 birds banded in 1977, 28
(23 males, 5 females) banded in 1976, 19 (18 males, 1 female) banded in
1975, 3 (2 males, 1 female) banded in 1974 and 2 (1 male, 1 female) banded
in 1973. The oldest males were at least five years of age, while the
oldest female was at least six years of age, as it was banded as an adult
in 1973.
It was not possible for birds to be known to be older than 6+
years as no banding of sage grouse was accomplished in North Park in the
1970-72 interval.
Little use can be made of recapture data outside of
longevity and movements as efforts were made not to recapture banded birds.
This was done to reduce possible trap mortalities of banded birds.
Despite
this precaution, one male banded in 1975 was accidentally killed in trapping
operations.
Total known trapping mortality in 1977 was 5 birds (4 males,
1 female).

�-12-

Summer Observations
Systematic examination for brood patches (area of bare skin at the posterior end of the abdomen) on females captured was conducted throughout
the breeding period.
The first indication of a brood patch was observed
on 20 April.
After 4 May no adult females failed to have a brood patch
while no yearling females failed to have a brood patch after 10 May.
These data suggest that some hens initiated incubation in mid-April, while
all nesting hens had initiated laying and incubation prior to 10 May.
Yearlings were about one week behind adult hens in appearance of a brood
patch.
Assuming a 2S day incubation period, the earliest nest could have
hatched about lS-20 May.
Searches for hens with chicks were made by walking selected transects and
using playbacks of tape recorded chick distress calls and by slowly driving
roads in preselected areas. While intensive field work was not initiated
until early July (because of inability to locate chicks prior to 3-4 weeks
post hatch), a newly hatched brood was observed on 28 May. During the 1
July-3l August period, 720 sage grouse were classified including 241 young
of the year.
Classification of birds observed by time period is presented
in Table 4.

Table 4. Composition
August 1977.

of observed

sage grouse, North Park, 1 July-3l

August

July
Category

I-IS

16-31

I-IS

16-31

Total

Adult males

lS4

131

67

0

352

Adult females

39

43

42

3

127

Chicks

79

82

73

7

241

Chicks/successful
female 1/

3.2(25)

3.2(26)

3.5(21)

2.3(3)

l/Hens accompanied by chicks were assumed to be successful.
parenthesis represent number of assumed successful hens.

Numbers

3.2(7S)

in

June precipitation
(next section) was quite low in North Park in 1977 and
as a result, broods were starting to concentrate near meadows by early July.
This allowed good samples of broods to be located in early July. However,
July and August were quite wet in North Park and as a consequence, broods
dispersed into upland sagebrush areas and became difficult to locate by
early August.
Because of problems delineated in 1976 (Braun 1977) and
dispersal of broods from meadows, essentially no searches for broods were
conducted after mid-August 1977. Data presented in Table 4 suggest average

�-l3-

chick survival in 1977 with an estimated 59 percent (75 of 127) oJ the
hens being successful.
Each successful hen appeared to raise 3 chicks to
age of independence.
Precipitation
Because of problems encountered in 1975 and 1976 with use of pit rain
gauges (Braun 1977) precipitation data were obtained only from standard
weather stations operated for the Weather Bureau, U. S. Department of
Commerce at Spicer and Walden.
These data are presented in Table 5.
Table 5.

Weather

data, North Park, Colorado

1977.

Spicer

Walden
June

July

Aug.

Sept.
1-9

June

July

Aug.

Sept.
1-9

Min. temp. of

27

30

28

22

34

34

30

24

Ma x. T emp. of

78

78

78

78

82

87

81

81

No. days 32 F
or below

9

3

8

3

o

o

3

2

Days of ppt.

5

16

14

1

3

14

12

1

.17

2.63

3.10

.11

1.54

2.42

1.37

1.22

1.69

1.11

1.24

1.29

o

Total ppt. (in.)
30 yr. average
ppt. (in.)

.16

.13

From data presented in Table 5 it is readily apparent that June in North Park
was warm and exceedingly dry. Total precipitation was down 87 percent in
the southwestern part of North Park (Spicer) and 90 percent in the central
area of the Park (Walden).
In contrast, precipitation in July and August
was almost 200 percent of normal near Spicer and 150 percent near Walden.
As a consequence, dessication of vegetation was not noticeable in late
summer and early concentrations
(late June) of sage grouse dispersed in July
and did not reform in late August or early September.
These data support
observations that sage grouse were well dispersed in small groups throughout
North Park going into and during the 1977 hunting season.
Hunting

Season Data Collection

The sage grouse season in North Park (Unit 12) in 1977 opened at sunrise on
10 September and closed at sunset on 25 September.
Season length was 16
days, the longest in modern times and 7 days longer than in 1976.
The daily
bag limit was three, with a possession limit of 6, identical to bag limits

�-14-

in this area in 1976. As in the 1974-76 period all hunters hunting sage
grouse in North Park were required to have in their possession a free special
permit.
Permits unlimited in number were available at all license agents
in North Park, the Fort Collins and Denver offices of the Division of Wildlife, and all Wildlife Conservation Officers and research personnel working
in North Park.
Purpose of the permit was to obtain reliable estimates
concerning hunter activities in North Park.
Check Stations
Two check stations were operated in 1977. These stations were located at
the State Line and Willow Creek Pass. As in previous years of operation,
each station was open from about 1000 to 1800 MDT, depending upon traffic
load. Both check stations were operated on the 10th, 11th and 18th. Except
for the 18th (one research person and one Wildlife Conservation officer) at
least two research persons and two Wildlife Conservation Officers were
assigned to each check station.
Data obtained per partv were: Countv of
origin. number of hunters. hours hunted (total of all hunters). birds
observed. birds bagged. birds lost. number of banded birds and location
where each was obtained. and area hunted within North Park. Most importantly, one wing was obtained from each bird that was checked.
Few birds
(51) were completely wingless.
Ovaries were classified as to having ovulated
in 1977 or not, for 41 females, with an additional 48 being collected for
analysis.
Sex by gonadal inspection was ascertained for 98 young of the year.
In addition to the two check stations, wing barrels and signs were placed
along Colorado 14 near Muddy Pass, near the State Line on the highway to
Saratoga, Wyoming, on Colorado 14 near Gould, at the junction of Jackson
County 6 and 34, north of Three Way and at Willow Creek Pass. Volunteer
stations were in operation the entire season near Muddy Pass and Gould, on
all days that manned check stations were not operating north of Three Way
and Willow Creek Pass and for the opening weekend only at Independence
Mountain (JC 6 and 34) and on the Saratoga Highway.
Research and management personnel field checked hunters whenever possible.
During the three days of check station operation, 353 hunters with 385 sage
grouse (1.1 birds per hunter) were checked.
These hunters reported observing
3,303 sage grouse.
Some duplications are undoubtedly present in the 3,303
birds observed.
Hunter efficiency was low at 11.7 percent (385 birds harvested + 3,303 birds observed) and 41 birds (10.6 percent of those retrieved)
were reported crippled and lost. Comparative data from 1974 through 1977
are presented in Table 6.
As in other years in the 1974-1976 period, distribution of harvest and
hunting pressure was not uniform within North Park. Lake John (23.9 percent)
and Ridge Road (20.8 percent) were the leading harvest areas, with the
Peterson Ridge-MacFarlane
Reservoir (12.7 percent), Eagle Hill (13.2 percent)
and Walden Reservoir (12.2 percent) zones being important harvest areas.
Comparative data for the 1974-1977 period are presented in Table 7.

�-15-

Table 6.

Sage grouse harvest

statistics,

North Park 1974-1977.1/

No.
Birds
Observed

No.
Birds
Harvested

Hunter
Efficiency

Crippling
Loss

Year

No.
Hunters
Checked

%

%

Birds
per
Hunter

1974

730

6,062

785

7.7

5.1

1.1

1975

738

5,735

551

9.6

7.1

0.7

1976

595

3,393

459

13.5

5.7

0.8

1977

353

3,303

385

11.7

10.6

1.1

1/ Only those statistics

collected

at check stations.

Upon examination of data presented in Table 7 it is apparent that hunter
pressure and harvest have declined at Independence Mountain and Spring
Creek-Owl Ridge, remained fairly constant at Walden Reservoir, Pole Mountain,
Peterson Ridge-MacFarlane
Reservoir, Michigan River Southeast, and Eagle
Hill, and increased substantially at Lake John and Ridge Road.
These
apparent changes may relate in part to changes in distribution of sage
grouse within North Park. Leks adjacent to Independence Mountain (Cowdrey
grounds) have been abandoned or have decreased in numbers of birds counted.
This is only partly true for known leks in the Spring Creek-Deer Creek-Owl
Creek area. Lek counts of males in the Walden Reservoir area have been
decreasing but percent of total harvest has remained fairly constant.
Counts
of sage grouse on leks in the Lake John and Ridge Road areas have increased
in recent years and new leks have been found in the Eagle Hill area.
These
data suggest that a relationship exists between numbers of active leks,
trends in counts of males on leks, area of harvest and related hunting
pressure.
Most hunters contacted at check stations were asked whether or not they
normally hunted sage grouse in North Park.
Of the 325 hunters responding,
211 (64.9 percent) reported they normally hunted sage grouse in North Park,
82 (25.2 percent) were first time sage grouse hunters, while 32 (9.8 percent)
reported they normally hunted sage grouse elsewhere.
Comparative data for
the 1974-1977 period are presented in Table 8.
It is obvious from data presented in Table 8 that more liberal bag limits
and season lengths in North Park in the 1975-1977 interval did not attract
hunters who normally hunted elsewhere.
Apparently sage grouse hunting in
North Park is traditional for many hunters and hunters who normally hunt
elsewhere were not attracted to North Park in large numbers.
It is of
interest to note that about one fourth of all sage grouse hunters contacted
each year were hunting sage grouse for the initial time. This suggests an
annual turnover of 25 percent of all sage grouse hunters afield.
Reasons
for this turnover are not known but may be related to non-success the
previous year(s), death, interest changes, etc.

�Sage grouse harvest and hunting pressure within North Park, 1974-1977):'/ 2:./

Table 7.

Independence
Mountain

Walden
Reservoir

Lake John

Ridge Road

Pole
Mountain
%

%

Spring
Peterson
Creek-Owl
Ridge-MacRidge
Far1ane Res.
%

%

%

Michigan
River
SE
%

%

%

%

%

%

%

%

%

%

4.7

6.0

18.1

16.4

12.4

15.4

9.6

11.3

4.7

3.8

20.3

19.6

17.9

13.4

24.7

14.2

16.7

11.1

15.6

1.8

2.0

17.2

14.3

13.6

%

Eagle
Hill
%

%

Harv. Hunt. Harv. Hunt. Harv. Hunt. Harv.
Hunt. Harv. Hunt. Harv. Hunt. Harv. Hunt. Harv. Hunt. Harv. Hunt.

1974

2.2

1.3

10.0

12.8

6.2

2.6

3.8

10.4

9.8

J

1975

3.1

3.8

24.3

1976

2.3

3.0

28.2

21.3

13.0

12.0

13.0

21. 1

2.9

4.5

18.5

21.3

10.9

6.3

3.1

5.9

8.0

5.0

1977

1.4

0.5

28.9

23.9

15.8

12.2

16.9

20.8

3.2

6.5

16.3

12.7

8.0

8.1

2.0

2.1

7.4

13.2

-1/Data from check stations only.
_2/Totals may not approximate 100 percent as each year some hunters and birds harvested could not be allocated
to a particular zone.

I
I-'
0\
I

�-17-

Table 8. Previous sage grouse hunting
North Park, Colorado 1974-1977.

experience

of sage grouse hunters,

Normally Hunt
Elsewhere
Percent
No.

First Time Sage
Grouse Hunter
Percent
No.

Year

Normally Hunt
in North Park
Percent
No.

1974

183

68.3

20

7.4

65

24.3

1975

432

64.9

47

7.0

187

28.1

1976

366

64.9

40

7.1

159

28.1

1977

211

64.9

32

9.8

82

25.2

Analysis

of Wings

A total of 632 sage grouse wings was collected in North Park during the 1977
hunting season.
Of this total, 334 were obtained at the two check stations
in North Park, 14 were collected at the Green Mountain Reservoir check
station in Middle Park, 185 were obtained in wing barrels (Gould = 129,
Muddy Pass = 21, Three Way = 21, Willow Creek Pass = 14, Independence Mountain = 0, Saratoga Highway = 0), 38 were received in mail envelopes, while
the remainder (61) were obtained through field contacts.
Distribution of
the wings collected by time of collection is presented in Table 9.

Table 9. Time distribution
Colorado 1977.

Number
Percent

of total

of sage grouse wings received,

10-11

12-16

425

39

67.3

6.2

North Park,

September
17-18
19-23

24-25

Total

83

33

52

632

l3.l

5.2

8.2

100.0

Data in Table 9 strongly suggest that the objectives of increasing hunter
opportunity and spreading the harvest over time were achieved in 1977.
Obviously, most harvest (88.6 percent of the wings received) occurred on the
three weekends but approximately 10 percent (11.4) of the harvest was
achieved during the weeks of 12-16 and 19-23 September.
Age and sex structure of the harvest as ascertained from examination
is presented in Table 10 along with comparative data from 1974-1976.

of wings

�Table 10.

Age and sex composition of the sage grouse harvest, North Park, Colorado 1974-1977.1/

Immatures
Females
No.
%

Year

Males
No.
%

1974

171

48.9

179

1975

101

47.6

1976

104

1977

136

Total

Males
No.
%

Yearlings
Females
No.
%

Total
No.
%

Males
No.
%

Adults
Females
No.
%

Total
No.
%

No.

%

51.1

350

50.1

49

35.5

89

64.5

138

19.8

45

21.4

165

78.6

210

30.1

111

52.4

212

42.0

52

46.8

59

53.2

111

22.0

55

30.2

127

69.8

182

36.0

49.5

106

50.5

210

42.3

46

39.3

71

60.7

117

23.5

49

28.8

121

71.2

170

34.2

46.9

154

53.1

290

45.9

47

38.2

76

61.8

123

19.5

48

21.9

171

78.1

219

34.6

~--------

-1/Data from wing collections only.

I
I-'
00
I

�-19-

As can be readily ascertained from data in Table 10, sex and age composition of the harvest in 1977 differed only slightly from the composition
in 1976. Percent of young in the harvest increased slightly in 1977
while percentage of yearlings decreased slightly.
Data in Table 10
suggests the following conclusions:
1.

Sex ratio at hatching

approximates

1:1.

2.

Differential survival favoring females starts before young grouse are
four months of age and is most pronounced in the older age classes.

3.

Production of young in the 1974-1977 interval has ranged from average
to good but has not been excellent in anyone
year.

4.

Overwinter survival of young grouse to the yearling class has been
average to good and has been adequate to maintain the population.

5.

The annual turnover rate for females is approximately
for males it is approximately 50 percent.

6.

Females comprise
the population.

34 percent

about 69 percent of the adult and yearling

and

segment of

Wings from 247 females (adult = 171, yearling = 76) were classified as to
primary feather molt. Because of the early nesting year, many hens that
had been successful in hatching their clutches had molted more old primaries
than they would have in "normal" hatching years.
Thus only 68 (51 adults
and 17 yearlings) (27.5 percent) females were classified as successful from
wing examination.
However, upon calculating the percentages of yearling
and adult males in each primary molt stage, and comparing those percentages
with the percentage of yearling and adult hens in each molt stage, it became
apparent that more hens were successful than first estimated.
Using this
method, an estimated 59.3 percent of the adult hens were successful nesters
while 32.7 percent of the yearling hens were successful.
This gave an
overall success rate of 50.3 percent.
These data are slightly higher than
those obtained in 1976 but are similar to estimated nesting success based
upon field observations (59%). The young to adult hen (including yearling
hens) ratio in the harvest was 1.2:1, while the young per successful hen
ratio was 2.0:1.
Comparative data for 1974-1977 are presented in Table 11.
Hatching dates were calculated for 290 chick sage grouse for which wings
were available.
These data are presented in Table 12. Because of obvious
limitations with the juvenile sage grouse aging technique, it is suspected
that calculated hatching dates are probably one week later than actual
hatch dates.
Hatching in 1977 started in late May. peaked about 10-15 June
with a few nests hatching in early July. Young hatching after about 20 June
were probably progeny of hens who lost their first clutch and renested.
Below normal precipitation in June and above average precipitation in July
and August would appear to have no measurable effect on nesting success or
brood survival.
Comparative hatching data for the 1974-1977 period are
presented in Table 13.

�-20-

Table 11. Sage grouse nesting success and production rates, North Park,
Colorado 1974-1977.

Young per Hen

Young per
Successful Hen

58.7

1.4:1

2.3:1

39.0

48.6

1.1:1

2.3:1

52.9

26.8

43.2

1.1:1

2.5:1

1977

59.3

32.7

50.3

1.2:1

2.0:1

Table 12.

Sage grouse hatching dates, North Park, Colorado 1977.

Year

Estimated Nesting Success
Total
Yearlings
Adults

1974

65.4

46.1

1975

53.2

1976

Time Interval

Males
No. Hatching
% of Total

Females
No. Hatching
% of Total

May 25-31

2

1.5

4

2.6

June 1- 7

37

27.2

25

16.2

8-14

43

31.6

45

29.2

15-21

29

21.3

50

32.5

22-28

11

8.1

17

11.0

June 29-July 5

11

8.1

11

7.2

July 6-12

3

2.2

2

1.3

136

100.0

154

100.0

Total

Ovarian Analysis
Ovaries were collected from 48 hunter harvested sage grouse in 1977 and were
analyzed following procedures described by Meyer et al. (1947), Kabat et
al. (1948), and Buss et al. (1951). An additional 41 ovaries were classified in situ at check stations. Data for 1977 are presented with comparative data from 1975 and 1976 in Table 14.

�-21-

Table 13.

1/
Sage grouse hatching dates, North Park, Colorado 1974-1977.-

Time
Interval

Sexes Combinedl/
1974

1975

Males
1976

1977

1975

Females
1976

1977

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

1.5

0.0

0.9

2.6

3.0

13.7

27.2

0.0

10.1

16.2

June

8.9

38.2

31.6

0.0

24.8

29.2

June

23.8

23.5

21.3

18.0

24.8

32.5

June

22.8

11.8

8.1

17.1

16.5

11.0

20.8

9.8

8.1

21.6

11.9

7.2

8.9

2.0

2.2

20.7

10.1

1.3

6.9

1.0

0.0

12.6

0.9

0.0

5.0

0.0

0.0

9.9

0.0

0.0

May 24 )

3.4
May 25-31
June 1-7
8.3

June
5.7
July
1.7
July
1.7
July

1/ Percent of total.
2/

Data from 1974 are not strictly comparable.

Table 14.

Sage grouse ovarian analysis, North Park, Colorado 1975-1977.
Percent Ovulating
1976
1977

Age Class

1975

Yearling

84.6

Adult

92.8

75.0
87.8

96.6
98.3

Number Ovulating/Examined
1975
1976
1977

33/39

33/

44

28/

64/69

65/

74

59/

29

60

�-22-

Data in Table 14 indicate that most females ovulated in 1977, possibly the
result of the early spring and low apparent stresses during the winter and
spring periods.
It would appear that not all females lay in a given year
with adults being more consistent layers than yearlings.
Differences
appear to be both age and year related and may be dependent upon late winter
and early spring weather conditions which conceivably affect nutrition.
Ovulation and timing may be nutritionally controlled by plant substances
that affects hormone release.
A similar hypothesis has been tested and
supported by data for California quail (Lophortyx californicus) (Leopold
et al. 1976).
Hunter Questionnaire
A total of 1,093 permits was issued for hunting sage grouse in North Park
in 1977, a decrease of 165 (13.1%) from 1976. As in previous years, most
hunters (37.7%) originated from the Denver area (Adams 7.2%, Arapahoe 5.9%,
Denver 12.4%, Jefferson 12.2%).
Only 10.8 percent of the hunters originated
from Jackson County, with 24.1 percent from Larimer County, 12.4 percent
from Boulder County, and 4.6 percent from Weld County.
All other counties
and states other than Colorado each contributed less than 2 percent of
the total hunters and less than 11 percent of all hunters.
Questionnaires were sent to all permittees immediately following the sage
grouse season in North Park (28 September).
Responses were received from
734 permittees.
On 19 October a followup letter was sent to all nonrespondents and 195 additional responses were received.
In all, 929
permittees (85.0%) responded (Table 15). Fourteen questionnaires were
undeliverable.
Mean values calculated for permittees responding to the
followup letter were used to project for the 15 percent (164) nonrespondents.
Comparative data for all four years of the permit season in North Park are
presented in Table 16. Upon examination of the data, it is apparent that
number of hunters fluctuated from 845 (most liberal season) to 1,187 (first
liberalizing of season length).
Hunter success decreased from 53 percent
in 1974 to 33 percent in 1976 and increased to about 46 percent in 1977.
Total harvest varied from 1,193 grouse (1974) in the most conservative
season to 847 grouse (1976). These variations were associated with sage
grouse nesting success and production of young.
Days per hunter and grouse
per successful hunter increased during the experimental seasons.
It would
appear that some hunters, especially successful hunters increased their
sage grouse hunting activities as a result of longer seasons and more
liberal bag and possession limits.
Despite the small increase in certain
hunter characteristics, overall hunter activity and success and total harvest
did not change with liberalizing of season length and bag and possession
limits.

�-23-

Table 15.

North Park sage grouse hunter questionnaire data 1977.
Projected
For

Projected
For 1,093
1,093

No. in sample

734

195

929

164

Percent of total
permittees

67.1

17.8

85.0

15.0

100.0

No. of hunters

569

150

719

126

845

Percent of hunters

77.5

76.9

77.4

76.9

77.3

No. of non-hunters

165

45

210

38

248

Percent non-hunters

22.5

23.1

22.6

23.1

22.7

No. of successful
hunters

291

55

346

46

392

Percent successful
hunters

51.1

36.7

48.1

36.7

46.4

Percent success of
permittees

39.6

28.2

37.2

28.2

35.9

No. of hunter days
Days/hunter
No. of sage grouse
bagged

1,115
2.0
810

297

1,412

2.0
112

2.0
922

252
2.0

88

1,664
2.0

1,010

Grouse/permitee

1.1

0.6

1.0

0.6

0.9

Grouse/hunter

1.4

0.7

1.3

0.7

1.2

Grouse/successful
hunter

2.8

2.0

2.7

2.0

2.6

No. of grouse lost

43

9

52

8

60

Crippling loss/
hunter

0.08

0.06

0.07

0.06

0.07

Total kill
Percent crippling loss

853
5.0

121
7.4

974
5.3

96
7.4

1,070
5.6

-----------------------------------------------------------------------------

�-24-

Table 15.

North Park sage grouse hunter questionnaire data 1977 (cont.).

Nl

N2

Banded birds

57

14

Non-deliverable surveys

14

0

Percent nondeliverable

1.3

Projected
For

Projected
For 1,093

71

11

82

0

0

14

Table 16. Comparative data, questionnaire survey, North Park sage grouse
seasons 1974-1977.
Category
Number of permits issued

1974

1975

1976

1977

1,184

1,541

1,258

1,093

87.0

85.0

979

845

77 .0

77 .8

77 .3

53.0

38.8

33.5

46.4

43.2

29.9

26.1

35.9

2,116
1.8

1,682
1.7

1,664

Days/hunter

1,587
1.6

Number of sage grouse
retrieved

1,121

1,000

797

1,010

Percent response

90.7

Number of hunters

966

Percent hunters

81.6

Percent successful hunters
Percent successful permittees
Number of hunter days

88.3
1,187

2.0

0.6

'0.6

Grouse/hunter

0.9
1.2

0.8

0.8

0.9
1.2

Grouse/successful hunter

2.2

2.2

2.4

2.6

Unretrieved grouse

72

75

50

60

1,193

1,075

847

1,070

Grouse/permittee

Total harvest
Percent crippling loss

6.0

7.0

5.9

5.6

Season length (days)

3

9

9

16

Bag and possession limit

2/4

2/4

3/6

3/6

�-25-

Time period of hunting was received for 867 hunters who harvest 914 birds.
Based on this sample, 67 percent of the hunters harvested 66 percent of
the total harvest during the opening weekend.
The initial week of the
season attracted 8.5 percent of the hunters who bagged 6.1 percent of
the harvest, eleven percent of the hunters harvested 13 percent of the
birds during the second weekend, five percent of the hunters harvested 6
percent of the grouse harvested during the second week, while 8.1 percent
of the hunters harvested 9.1 percent of the grouse during the third and last
weekend of the season.
Obviously, a longer season has resulted in a
reduction of hunter pressure during the initial weekend without increasing
overall hunter numbers or total harvest.
Despite the longer seasons and higher bag and possession limits in 1976 and
1977, percentage of successful hunters attaining a full bag limit (17.9% in
1976 and 19.1% in 1977) has remained fairly constant.
Obviously, as
indicated from estimates of hunter efficiency, it is not easy to harvest
sage grouse.
Hunting as presently allowed in North Park and other areas of
Colorado is not a threat to long term stability of sage grouse populations.
Hunter pressure and sage grouse harvest in North Park as derived from the
Statewide Small Game Questionnaire Survey are not presently available for
1977. Results of this survey in 1974-1976 are presented in Table 17. This
survey continues to overestimate hunter numbers and total harvest.

Table 17. Comparative
North Park 1974-1977.

Year

data, number of sage grouse hunters

Number of Hunters
This Study
Statewide

Survey

and total harvest,

Number of Sage Grouse Harvested
Statewide Survey
This Study

1974

960

1,759

1,174

2,509

1975

1,187

1,600

1,053

1,973

1976

975

877

829

1,402

1977

842

1,069

Band Recoveries
Eighty-four bands were reported from birds harvested during the 1977 hunting
season.
Four additional bands were reported in 1977 from road casualties
(2), and predator kills (2). Of the 88 total recoveries, 36 were from
females (1973 = 3, 1974 = 1, 1975 = 5, 1976 = 6, 1977 = 21), while 52 were
from males (1973 = 1, 1974 = 1, 1975 = 9, 1976 = 8, 1977 = 33). Four (3
shot recoveries and 1 road casualty) were from the 18 chicks banded in 1977.
Excluding the four chicks, banding and recovery data are presented in Tables
18 and 19, while survival and mortality estimates are shown in Table 20.

�-26-

Table 18.

Year

Male sage grouse banding and recovery data, North Park 1973-1977.

Number Banded

1973

1974

Number Recovered
1976
1975

1977

Yearlings
6

1973

80

1974

54

1975

139

1976

120

1977

183

4

6

1

0

6

5

3

1

18

6

7

16

5
19

Adults
1973

99

1974

88

1975

152

1976

114

1977

123

7

4

1

0

1

8

5

1

0

10

4

2

16

3
12

Data presented in Tables 18, 19 and 20 indicate that survival rates are
different between age classes and sexes. Direct recovery rates for males
continue to be about 10 percent, while those for females continue to vary
from 5 to 10 percent. These data suggest that males are slightly more
vulnerable to hunting than are females. This further supports available
data that females predominate in the population and are not selected for by
hunters.
Estimation of Fall Population Size
Estimates of fall population size are difficult to derive, primarily because
of unknown vulnerability of chicks and inexact survival estimates. Provided
that all birds of each age and sex class are equally available and vulnerable
to hunters and using the average mortality rates given in Table 20 the 1977
fall population size in North Park was calculated as follows:

�-27-

Number of Birds Banded In:

Alive in 1977:

1973

Males

9

Females

27

1974

Males

7

Females

18

1975

Males

78

Females

65

1976

Males

148

Females

1977

Males

306

Females

234

Total Banded Birds Alive

Males

548

Females

446

=

102

Table 19.

Female sage grouse banding and recovery data, North Park 1973-1977.

Year

Number Banded

1973

1974

Number Recovered
1976
1975

1977

Yearlings_
1973

41

1974

22

1975

62

1976

71

1977

101

2

2

4

1

2

2

1

1

0

6

2

1

2

4
6

Adults
1973

68

1974

27

1975

68

1976

74

1977

133

5

1

0

1

1

2

0

0

1

6

2

4

2

2
13

�-28-

Table 20. Survival and mortality
Park by age and sex 1973-1977.

estimates

for sage grouse banded

Average
Survival Rate

Category

in North

Average
Mortality Rate

.813

.187

Adult males

.436

.564

All females

.705

.295

Yearling

males

Of the 548 banded males potentially alive in 1977, bands were received from
50, while bands were received from 34 of the 446 hens potentially alive.
If
54 percent (wing survey data) of the 1,070 birds harvested (questionnaire
data) were adults and yearlings, then 578 of the birds harvested were
older than one year.
Of this number, 81 (84-3 chick recoveries)
(14.0%) were banded, thus there should have been about 4,121 adult and
yearling sage grouse alive in North Park in the fall of 1977 (if 14.0%
578 birds, then 100% = 4,121 birds).
This is remarkably similar to the
4,040 adults and yearlings estimated to have been alive in the fall of 1976
(Braun 1977).
In 1977, immatures comprised 46 percent (wing survey data)
of the harvest, thus there should have been about 3,496 chicks in the 1977
fall population.
Thus the total population in the fall of 1977 should have
been about 7,617 or 7,600 birds.
I believe this estimate is conservative.
Problems still exist with small samples of banded hens and no valid estiamtes
of chick vulnerability.
Of interest are the three shot recoveries from 18
chicks banded in 1977. This is a direct recovery rate of 17 percent and
suggests that chicks may be more vulnerable than adults.
However, the sample
is not large and was not well distributed throughout North Park.
The overall harvest rate was about 14 percent (1,070 + 7,600) with adult and
yearling males having a somewhat higher harvest rate (10.1% direct recovery
rate) than adult and yearling females (8.1% direct recovery rate).
Of the
342 adult and yearling birds for which wings were available, 247 (72.2%)
were from females.
Thus, of the approximately 4,121 adult and yearling birds
in the pre-season population, 2,975 (4,121 x 72.2 = 2,975) were hens.
Total
chicks in the fall population in 1977 based on 1.2 chicks per hen in the
harvest (1.2 x 2,975 = 3,570) approximated 3,570.
This number is only slightly
higher than the 3,496 chicks estimated from overall age ratios and band
recoveries in the 1977 harvest to have been in the fall population.
These
data do not suggest that chicks are more vulnerable to hunting than are
adults and yearlings.
It is interesting to compare estimates of adults and yearlings alive in the
spring and fall.
Spring estimates based on counts of males on leks indicated
that about 5,784 birds were present.
Fall estimates based on age and sex

�-29-

ratios in the harvest and band recovery data indicated that about 4,121
adult and yearling birds were present.
This is a difference of about 1,600
birds.
It is possible that spring estimates are too high. as either (1)
more or fewer leks are unknown than estimated, or (2) peak counts of
males on leks represent more than 50 percent of the cocks present
in a given area.
It is also possible that fall estimates based on harvest
age and sex ratios and band recoveries are low. It would appear that the
1977 spring population in North Park was at least 4,000 and probably not
more than 6,000 birds.

LITERATURE

CITED

Beck, T.D.I.
1975. Attributes of a wintering population of sage grouse,
North Park, Colorado.
M. S. Thesis.
Colorado State Univ., Ft. Collins.
49 pp.
1977. Sage grouse flock characteristics
winter.
J. Wildl. Manage. 41(1):18-26.
_____ , R. B. Gill, and C. E. Braun.
1975.
sage grouse from wing characteristics.
(Revised).
Colo. Div. Wildl.
4 pp.

and habitat

selection

in

Sex and age determination of
Game Inf. Leaflet No. 49

Braun, C. E. 1977. Evaluation of the effects of changes in hunting regulations on sage grouse populations.
Colorado Div. Wildlife, Prog. Rept.,
Fed. Aid Proj. W-37-R, Work Plan 3, Job 9. pp. 35-58.
_____ , and T.D.I. Beck. 1976. Effects of sagebrush control on distribution
and abundance of sage grouse.
Colorado Div. Wildlife, Final Rept.,
Fed. Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84.
_____ , R. K. Schmidt, and G. E. Rogers.
tailed ptarmigan with tape-recorded
90-93.

1973. Census of Colorado
J. Wildl. Manage.
calls.

white37(1):

Buss, I. 0., R. K. Meyer, and C. Kabat.
1951. Wisconsin pheasant reproduction studies based on ovulated follicle technique.
J. Wildl.
Manage. 15(1):32-46.
Carr, H. D. 1967. Sage grouse and sagebrush control.
rado State Univ., Fort Collins.
106 pp.

M. S. Thesis.

Colo-

Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios
from wings.
J. Wildl. Manage. 19(2):267-272.
Gill, R. B. 1965. Distribution and abundance of a population of sage
grouse in North Park, Colorado.
M. S. Thesis.
Colorado State Univ.,
Fort Collins.
187 pp.
1975. A volunteer wing collection
Hoffman, R. W., and C. E. Braun.
Colo.
Div. Wildlife.
3 pp.
Game Inf. Leaflet No. 101.
Kabat, C., I. O. Bu~s, and R. K. Meyer.
1948. The use of ovulated
follicles in determining eggs laid by the ring-necked pheasant.
Wildl. Manage. 12(4):399-416.

station.

J.

�-30-

Leopold, A. S., M. Erwin, J. Oh, and B. Browning.
adverse effects on reproduction in California
191(4222):98-99.

1976. Phytoestrogens:
Science
quail.

Meyer, R. K., C. Kabat, and I. O. Buss. 1947. Early involutionary changes
in the post-ovulatory follicles in the ring-necked pheasant.
J.
Wildl. Manage. 11(1):43-49.
Pyrah, D. G.
Wildlife
71 pp.

1963. Sage grouse investigations.
Idaho Fish and Game Dept.,
Restoration Div., Job Compl. Rept., Fed. Aid Proj. W-125-R.

Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations
sharp-tailed grouse.
J. Wildl. Manage. 38(4):616-621.

of male

Robel, R. J. 1969. Movements and flock stratification within a population
of blackcocks in Scotland.
J. Anim. Ecol. 38(5):755-763.
Wallestad, R. O. 1970. Summer movements and habitat use by sage grouse
broods in central Montana.
M. S. Thesis.
Montana State Univ.,
Bozeman.
51 pp.

Prepared

by __

-..!::~~{a/::::::..:...-'_s._.---L~-="":::"=::~

Clait E. Braun
Wildlife Researcher

_

�April 1978

-31-

JOB FI~

REPORT

COLORADO

State of

----------------------

Project

No.

Game Bird Survey

W-37-R-31

Job No. ·
1_0
_
3
Investigations of the Distribution and Status
Job Title __~o~f~S~a~g~e~b~r~u~s~h~a~n~d~S~a~g~e~G~r~o~u~s~e~l~·n~~th~e~M~o~f~f~a~t~C~o~u~n~t~y~
_

Work Plan No.

Period Covered:
Personnel:

January 1, 1976 through March 31, 1978

Donald M. Hoffman.

ABSTRACT
A draft of a manuscript entitled "Investigations of the Distribution and
Status of Sagebrush and Sage Grouse in the Moffat County Area" has been
completed and submitted for review and subsequent publication as a Division
of Wildlife Special Report.
This is scheduled for completion in the 1978-79
Segment.

Prepared

by
Donald M. Hoffman
Wildlife Researcher

��April 1978

-33-

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Game Bird Survey

W-37-R-31

Work Plan No.

9

Job Title

Dynamics

Population

Period Covered:

Job No.

5

----------------------------

and Habitat Relationships

of Blue Grouse

April I, 1977 to March 31, 1978

Personnel:
T. Beck, D. Benson, R. Binford, C. Bonney, C. Braun, R. Clippinger,
J. Corey, D. Covic, K. Duncan, H. Funk, J. Gerrans, K. Giesen, D. Hoart,
D. Hoffman, R. Hoffman, D. Luce, B. McCloskey, R. Oakleaf, B. Sigler, M.
Smith, S. Steinert, and W. Woodard.

Investigations concerning the effects. of hunting on blue grouse (Dendragapus
obscurus) populations, stability of breeding population levels, and habitat
relationships of blue grouse were initiated in 1975 and continued in 1976
and 1977. Studies were conducted on two selected locations of differing
habitat types in northwestern Colorado.
Density of territorial males
appeared to be slightly higher on Green Mountain (6.6 males/km2) than at Eby
Creek (5.8 males/km2).
However, based on ecological density, Eby Creek
supported 9.7 males/km2, whereas, Green Mountain maintained about 7.4 males/
km2 of suitable habitat.
Changes in breeding densities from 1975 through 1977
were not significant.
Nesting activities were advanced in 1977 due to low
overwinter snowfall.
Estimated peak of hatch for both study areas was
between 11 and 24 June; almost one week earlier than in 1976 and over two
weeks earlier than in 1975. Overall nesting success in 1977 was somewhat
higher on Green Mountain (64.1%) than at Eby Creek (57.1%) and compared
favorably with 1975 (64%) and 1976 (62%) estimates for Green Mountain.
Adult
and yearling hens were equally successful in bringing off broods in 1977.
Observations were made on 71 broods, including 37 at Eby Creek and 34 in
Middle Park, with a composite average brood size of 3.8 and 3.9 chicks per
brood, respectively.
Estimated fall densities of approximately 26 birds/km2
for both study areas suggested good production and survival of young in 1977.
Data from wings (N = 948) collected at check stations (114), wing barrels
(798), and from the mail wing survey (36) revealed that the harvest was
composed of 51 percent immatures, 3 percent yearlings, and 46 percent adults.
Due to an early onset of primary molt, most yearlings were indistinguishable
from adults and thus, were under represented in the harvest sample.
For
Middle Park, hunter efficiency was high (47.1%) and only 9 birds (4.0%) were
reported crippled and lost. Hunter success was calculated at .99 birds/
hunter.
About 6 percent of the population on Green Mountain was harvested in
1977. Estimates of hunter efficiency (39.1%), crippling loss (6.7%), and
hunter success (.63 grouse/hunter) for the Eagle area were slightly different

�-34ABSTRACT

(continued)

but compared favorably with data from Middle Park. Overall, hunter pressure
was down from previous years, but hunter success was up. A total of 1,502
big game hunters were surveyed in 1977 in order to evaluate the combined
blue grouse-big game season.
Few big game hunters (6.3%) took advantage of
the longer grouse season, but their general response to the combined blue
grouse-big game season was highly favorable (52.1%). Few hunters «1%)
were in the field during the big game seasons strictly to hunt grouse.
The
additional harvest of grouse resulting from the longer season amounted to
only about a37 percent increase in the total kill.

�-35-

HABITAT

POPULATION DYNAMICS AND
RELATIONSHIPS OF BLUE GROUSE
Richard W. Hoffman

Blue grouse are the most widespread member of the upland game birds in
Colorado, occurring in varying densities over more than 51,800 km2 (20,000
mi2) of diverse habitats and terrain throughout the state (Rogers 1968).
Furthermore, blue grouse rank first in terms of population numbers (250,000+)
and annual harvest (14,000) among resident grouse species.
Yet, Colorado
has had a long history of conservative seasons on blue grouse, in spite of
its abundance and wide distribution.
This conservatism has mainly resulted
from: (1) lack of data upon which to base management recommendations,
and
(2) the misconception that hunting can be a major mortality factor on blue
grouse.
Data presented in this report concern these problems and represent
results of population studies initiated in 1975.
P. N. OBJECTIVE
Major objectives of this study are to (1) increase the harvest of blue grouse
in Colorado (double present harvest estimates) without harm to breeding populations in subsequent years, (2) to identify differences in breeding densities
due to differing habitats, and (3) to document the stability of breeding
densities over time.
SEGMENT OBJECTIVES
1.

Review literature concerning (a) techniques for marking, capturing, and
censusing grouse, (b) methods of aging and sexing blue grouse, and (c)
population dynamics, behavior, and habitat requirements of blue grouse,
and (d) effects of hunting on grouse populations.

2.

Trap and individually mark 50 birds (including adults,
juveniles) on each of the two selected study areas.

subadu1ts,

3.

Estimate breeding densities on the study areas through
acoustical census and systematic search.

the use of

4.

Estimate

5.

Vegetatively describe areas studied and correlate
with features of the environment.

6.

Manipulate

7.

Compile data, analyze

Reference
materials
discussed

nesting

success and production

hunting

and

on the study area.

season and obtain harvest

blue grouse densities

data.

results,

and prepare progress

METHODS

AND MATERIALS

reports.

is made to Hoffman (1976) for a detailed discussion of methods
used in this study. Any deviations or additions to these are
in the Results and Discussion section.

and

�-36-

DESCRIPTION

OF STUDY AREAS

Research is presently being conducted on two selected study areas of differing
habitat types in northwestern Colorado.
These areas are described in detail
in previous reports (Hoffman 1976 and 1977) and are depicted in Figs. 1 and 2.
RESULTS AND DISCUSSION
Breeding

Densities

Grouse arrived on the breeding range at both study areas in early April.
Early
arrivals were mostly males, with numbers of females observed increasing toward
mid- to late April.
Activities associated with breeding and density of
breeding grouse peaked in late April-early May. Timing of breeding events
was similar to that in previous years.
Total number of territorial males recorded during peak of breeding activities
was used as an index to population size. Number of breeding hens and nonterritorial yearling males could not be accurately ascertained because of
their secretive habits, extensive movements, and poor response rate to recorded
calls.
Since data collected in other studies suggest there is a 1:1 sex ratio
among breeding birds (Zwickel 1972, Bendell et al. 1972), breeding population
levels were estimated by doubling the count of territorial males.
Density of territorial males appeared to be slightly higher on Green Mountain
(6.6 males/km2, 16.7 males/mi2) than at Eby Creek (5.8 males/km2, 14.9 males/
mi2).
However, based on ecological density, Eby Creek (60% suitable habitat)
supported 9.7 males/km2 (24.9 males/mi2) and Green Mountain (90% suitable
habitat) maintained about 7.4 males/km2 (19 males/mi2).
Total estimated
breeding populations in 1977 were: Green Mountain - 34 birds, 13.1 birds/km2;
Eby Creek - 58 birds, 11.6 birds/km2.
Changes in breeding densities from 1975
through 1977 were not significant (Table 1).

Table 1.

Blue grouse breeding

Study Area

Size
(km2)

Total
1/
Breeding Population1975
1976
1977

Birds per km2
1975
1976
1977

Green Mountain

2.6

32

32

34

12.3

12.3

13.1

Eby Creek

5.0

Nnl/

58

58

NO

11.6

11.6

1/ Excludes

1/ No data.

non-territorial

densities,

males.

1975

to 1977.

�. i,

/

-37-

\

(

"\

,

"

\

.'

:\
\.

':..

,.,/1
: .

,. /.:.

• i : ::
•

, t··
'/

MOUNTAIN
AREA
C LORADO

'_,,':
I
~I

./~ ..~ ....:-: :,.&lt;'

. \

GREEN

.1

- STUDY

AREA

BOUNDARY

,
"

..

·

~
~

·
"

,
}

l

.,..

i...

-,

\_--

'

.••••

60

.
••

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'\

,-

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•••••••.• ":10.

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&lt;,

~-

.
.f'I,

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..•.

,

'

,

.\

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.•..-

Ilr--l

__

S_C_A_L_E_'N_O-L-KI_LO_M_E_T_E_R_S_
Fig,

1.

Green Mountain

study area,

,. \.;

.~ ,
~'

,

�.,

j~2

lij.:"'-'m E.

j.j •

-4.)

-

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,
r-r+ . -"--

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. -~..:

03 '

~

j.

WHITE

liJVEI'

;\AT

}.'()j{

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1079·.

~:1&gt;~

".::--~

r..-l
~/6 1/ -~

o~',."

./

~~\i
o6/1"~ _ -2,-,
9042

.+

V,&gt;-

'\,

"'1/

-j;

i

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/

'-

.•

.~\

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&lt;,

I
W

00
I

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,~

v~&lt;:,&lt;;&gt;

EBY CREEK AREA
COLORADO

'\
;-4

l::t
N'_'''";~

\r

I

----STUDy AREA BOUNDARY
SCALE IN KILOMETERS
I

"
Fig~ 2.

Eby Creek study area.

0

~J!n II!!

,
x I" /

�-39-

Hatching

Dates and Nesting

Success

Since hens with broods are easier to locate
than those without, this factor
tends to produce an over-inflated estimate on nesting success when calculated
from field observations.
Consequently, analyses of the wing molt pattern
of hunter harvested birds was used as the best estimate of nesting success
(Braun 1971). This technique provides a minimum estimate, as some successful
hens will have completed their molt by mid- to late September and cannot
be distinguished from unsuccessful hens. Due to an early onset of primary
molt in 1977, many yearling birds had completed their molt by the fall
hunting season and could not be distinguished from adult grouse.
Consequently,
sample sizes for yearlings were small. Blue grouse nesting succes, production rates, and hatching dates for 1977 are given in Table 2.

Table 2.
1977 .

Blue grouse nesting

success, production

rates, and hatching

dates,

Nesting
Success

Percent
Immatures
In Harvest

Young
Per Hen
Ratio

Young per
Successful
Hen Ratio

Peak of
Hatch

Middle Park

64.1

50.9

1. 9:1

3.0:1

11-24 June

Eagle

57.1

50.7

2.3:1

4.5:1

11-24 June

Area

In 1977, preparation for nesting by females was advanced due to low overwinter snowfall resulting in early plant phenology and abundance of bare
ground.
Some females started laying in early May and most had initiated
incubation by late May. Estimated peak of hatch in 1977 for 414 chicks
examined from Middle Park (265) and Eagle (149) was between 11 and 24 June,
almost one week earlier than in 1976 and over two weeks earlier than in 1975.
Hatching dates ranged from 31 May to 23 July. Seven percent of the chicks
hatched in July. These birds were believed to be the result of renesting.
Timing of nesting events was similar between study areas.
Middle Park
Wings from 159 females (adults = 143, yearlings = 16) were classified as to
primary molt. Examination of the molt pattern revealed 91 of 143 (63.9%)
adult hens and 11 of 16 (68.7%) yearling hens were successful nesters in
1977. Overall nesting success was 64.1 percent (Table 2). Samples of
yearlings were small and may not be representative of this age class; however,
the data indicate equally good nesting success for yearling and adult hens
in 1977. Comparative estimates of nesting success in 1975 (64%) and 1976
(62%) show little variation among years.
Adult and yearling hens were equally successful in bringing off brdods in
1976 and 1977, but not so in 1975 (Table 3). Reasons for this variation

�-40-

are uncertain, but may be related to the earlier onset of nesting and
longer nesting seasons in 1976 and 1977, conditions that were apparently
favorable for better nesting success of yearlings.

Table 3. Estimated nesting
Park area, 1975 to 1977.

success

for adult and subadult

hens, Middle

Age

1975

(N)

1976

(N)

1977

(N)

Adult

78

(41)

63

(60)

64

(143)

Subadult

27

(15)

61

(28)

69

( 16)

Total

64

(56)

62

(88)

64

(159)

Eagle
For the Eagle data, adults and yearlings were grouped together for analyses.
In all, wings from 67 females (adults = 63, yearlings = 4) were classified
as to primary molt of which 38 (57.1%) were considered successful nesters
in 1977.
Inadequate samples of wings were obtained from the Eagle area in
1975 and 1976 to reliably estimate nesting success.
Production
From mid-June until late August 1977, complete counts were made on 71 broods,
including 37 at Eby Creek and 34 in Middle Park.
During the same period in
1975 and 1976 complete counts were made on 10 (1975) and 27 (1976) broods
observed on the two areas.
The increase in brood observations from 1975
to 1977 was related to improved observer efficiency and larger area searched.
Results of brood surveys from 1975 to 1977 are given in Table 4.
Yearly average brood sizes recorded were: Green Mountain, 1975 = 3.9,
1976 = 4.4, and 1977 = 3.8; Eby Creek 1975 = No data, 1976 = 3.4, and
1977 = 3.9. Average brood size was similar between areas in 1977.
The
apparently lower brood size at Eby Creek in 1976 may be attributed to a
larger sample of broods counted in late July-early August, whereas, primarily
early to mid-July counts were made at Green Mountain.
Rogers (1968) found
average brood sizes of 4.1, 2.6, and 3.9 in 3 years of study in western
Colorado.
Sex ratio of chicks was determined from hunter harvested birds for which sex
was ascertained.
Based on the examination of 723 juvenile wings collected in
1975 (79), 1976 (166), and 1977 (478), sex ratios did not deviate significantly
from 1:1.

�-41-

Table 4. Average brood size, range, and number of broods observed
monthly intervals, 1975 to 1977. 1/

by

Eby Creek

Green Mountain

1975

1976

1977

1976

1977

Mean

ND'!:/

5.3

5.1

5.0

4.6

Range

ND

3-7

4-7

4-6

3-6

Sample Size

ND

3

11

2

7

Mean

5.2

4.1

4.0

3.7

4.1

Range

2-7

2-6

1-7

1-7

2-9

Sample Size

4

7

10

9

10

Mean

3.0

3.0

2.5

2.2

3.6

Range

1-5

1-4

1-4

1-10

Sample Size

6

13

5

20

Month

June

July

August

1/ Only distinct
2/

1

broods with full counts are included.

No data, late hatching

year.

Average brood size by September 1 provides some indication about survival of
chicks.
However, average brood size determined from counts in late August
or early September may be misleading due to brood breakup and dispersal.
While harvest may not accurately reflect population composition, it is
believed that the young to successful female ratio in the harvest provided
the best measure of brood size and survival of chicks through September.
These data are summarized in Table 2 along with estimated nesting success
and hatching dates.
For both study areas, production and survival of
young in 1977 was good.
Fall Densities
Estimated 1977 fall densities for both study areas are presented in Table 5.
Calculations are based on the following assumptions: (1) estimated percent
nesting success is correct, (2) immigration equals emigration, (3) a 1:1
sex ratio exists in the breeding population, and (4) estimated average brood

�-42-

size by September 30 is accurate.
Mortality of the breeding population
from April 1 to September 30 is not figured into the calculations, but is
probably minimal «10%) (Bendell and Elliott 1967). Obviously none of the
assumptions, nor the accuracy of the data, are absolutely correct, but it
is belived data presented in Table 5 reflects the general situation on the
study areas.
Even though brood size decreased over the summer months, production still
contributed to nearly a doubling of the spring population by late September.
Percent gain was more than adequate to maintain the breeding population.
Furthermore, data from Middle Park indicate production was remarkably
uniform between years, and regardless of size or age of the breeding population, nesting success, or clutch size, survival of chicks until late
September was very similar (Table 6).
Harvest
Hunting

Season

The regular blue grouse hunting season in 1977 opened on 10 September and
closed at sunset on 9 October.
An extended season was held from 15 October
until 15 November in conjunction with big game seasons.
During this period,
hunters could take blue grouse in areas west of Interstate 25 and in Unit 80
whenever these areas were open to hunting of deer and elk. Daily bag and
possession limits were 3 and 6 grouse, respectively.
Middle Park
A check station was operated at the Prairie. Point Campground along Highway 9
on the south end of Green Mountain Reservoir in Middle Park. The check
station was operated opening weekend from 1000 to 1800 MST. All vehicles
were stopped on the highway, but only grouse hunters were directed to pull
off at the check station.
Data obtained per party included: County of origin,
number of hunters, hours hunted, birds observed, birds bagged per hunter,
area hunted, and location where each bird was harvested.
One wing was
removed from each bird checked provided no wings had been deposited in wing
barrels.
Whenever a bird was missing one or both wings, the hunter was
questioned as to what he did with the wings.
Thus, the status of all wings
was recorded as follows: (1) hunter disposed of wings, (2) hunter deposited
wings in wing barrel, and (3) wing collected at check station.
Sex by
gonadal inspection was ascertained for all young of the year whenever
possible.
Ovaries were collected from adult and yearling hens and crops
and gizzards were collected from all birds when present.
In addition to the check station, 15 volunteer wing collection stations were
available to hunters throughout the entire season at various locations in
Middle Park. Hunters were also contacted opportunistically
in the field
whenever possible.
Wing envelopes distributed by the management section also
provided wings from the Middle Park area.

�Table 5.

Estimated fall densities of blue grouse, 1977.

Total
Breeding
Population

Nesting
Success
(%)

Average
Brood Size
by Sept. 30

Total
Production
by Sept. 30

Total
Population
by Sept. 30

Birds
per
km2

Percent
Gain

Green mountain

34

64

3.0

33

67

26

49

Eby Creek

58

57

4.5

74

132

26

56

Area

Table 6.

Comparative fall densities of blue grouse on Green Mountain, 1975 to 1977.

Year

Total
Breeding
Population

1975

..,..I

Nesting
Success

Average
Brood Size
by Sept; 30

Total
Production
by Sept. 30

Total
Population
by Sept. 30

Birds
per
km2

Percent
Gain

32

64

2.6

27

59

23

46

1976

32

62

3.0

30

62

24

48

1977

34

64

3.0

33

67

26

49

w
I

�-44-

A total of 606 blue grouse wings were collected from the Middle Park area
in 1977 (Table 7). Fifteen wing collection stations accounted for 501 of
606 wings collected.
Number of wings collected from each barrel were as
follows: Beaver Creek 8, Williams Fork 0, Corral Creek 53, Troublesome 18,
City Reservoir 17, Gore Pass 69, Pinto Creek 5, Chimney Rock 85, Kremmling 14,
Trough Road 74, Lawson Ridge 2, Spring Creek 62, Williams Peak Road 63, Ute
Pass 26, and Rock Creek 5. The remaining wings were collected from the check
station (87 wings) and mail wing survey (18 wings).
Because of adverse
weather conditions opening weekend, hunter pressure was low. Otherwise, it
is believed that more wings would have been collected at the check station.
Hunters from areas outside of Middle Park were also contacted at the check
station, but their hu~ting activities are excluded from this report.

Table 7.
season.

Number and source of blue grouse wings collected

Wing Barrels

Area

Check Stations

during the 1977

Mail Survey

Total

Middle Park

501

87

18

606

Eagle

297

27

18

342

Total

798 (84.2%)

114 (12.0%)

36 (3.8%)

948

During the two days of check station operations, 228 hunters with 226 blue
grouse were checked.
Total birds reported observed by these hunters was 480.
Some duplications are undoubtedly present in the 480 observations.
Hunter
efficiency was high (47.1%) (226birdsharvested
; 480 birds observed) and
only 9 birds (4.0%) were reported crippled and lost. Hunter efficiency was
substantially higher in 1977 (47.1%) than in 1975 (28.4%) and just slightly
higher than in 1976 (43.3%). Crippling loss was less than 1976 (8.2%) but
similar to that calculated in 1975 (4.4%). Indications were that hunter
pressure was down, but due to the good production year, hunter success in
1977 (.99 birds/hunter) was up from 1975 (.40) and 1976 (.44) estimates.
Of the 228 regular season hunters checked in 1977, 64 percent were unsuccessful, 13 percent harvested 1 bird, 10 percent harvested 2 birds, and 13
percent harvested the daily bag limit of 3 birds.
After opening weekend, hunting pressure decreased markedly.
Although the
entire blue grouse season lasted 57 days, 42 percent of the wings collected
from volunteer stations were obtained opening weekend.
In order to evaluate
the effectiveness of wing barrels for sampling, Middle Park hunters were
questioned as to what they did with the wings from the birds they harvested.
This information is summarized as follows:
Percent
Percent
Percent
Percent

wings
wings
wings
wings

deposited in barrels
collected that should have been deposited
disposed
of unknown status

60.7
18.9
17.4
3.0

�-45-

Only 4 birds were reported killed in the Green Mountain study area. Based
on the estimated fall population of 67 birds, about 6 percent of this population was harvested in 1977. Furthermore, only 1 of 20 (5%) birds banded
in 1977 on the area was recovered.
Data from other studies suggest 25 to
30 percent of the fall population can be safely harvested with no adverse
effects on the subsequent spring breeding population (Zwickel 1958, Bendell
and Elliott 1967).
Age and sex composition of the 1977 harvest sample is presented in Table 8
along with 1975 and 1976 data. As can be readily ascertained from data in
Table 8, sex and age composition of the harvest in 1977 was not greatly
different from that determined in 1975 and 1976. Few yearlings were identifiable in 1977 due to the early onset of primary molt.
Since males initiate
their molt before females, there was a pronounced deficiency of males in
the yearling age class in all years, but especially so for 1977. Otherwise,
sex ratios for the other two age classes did not deviate significantly from
1:1. There appeared to be no selection for certain age or sex classes.
It
is also apparent that production and survival did not change markedly during
the 3 year period, but were best in 1977.

A total of 342 blue grouse wings were collected in the Eagle area during the
1977 season
(Table 7). Ten wing collection stations accounted for 297
of 342 wings collected.
Number of wings collected from each barrel were as
follows: Red Sandstone Road 57, Squaw,Creek 5, Lake Creek 5, Muddy Pass 63,
Milk Creek 23, Eby Creek 19, Brush Creek 27, Gypsum Creek 8, Cottonwood
Pass 11, and Coffee Pot Springs 79. The remaining wings were obtained
through the operation of a check station on Muddy Pass (27) and from the
mail wing survey (18). Adverse weather conditions opening weekend
resulted in low hunter pressure; consequently, the check station was only
operated opening day and few wings were collected.
Seventy-one hunters, with 45 grouse, were contacted at the Eagle check
station.
These hunters reported observing 115 blue grouse.
Hunter efficiency
(39.1%) and crippling loss (6.7%) compared favorably with data from Middle
Park. Crippling loss was estimated to be lower in 1975 (4.5%) and 1976
(3.9%), while hunter efficiency varied from 26.5 percent in 1975 to 47.8
percent in 1976. Hunter success in 1977 (.63 grouse/hunter) was up from 1975
(.38) but down from 1976 (1.0). Most of the pressure occurred opening
weekend as 48 percent of the wings collected from volunteer stations were
obtained during the first 3 days of the season.
Forty-five of 71 hunters
(64%) checked were unsuccessful, 13 (18%) had one bird, 10 (14%) shot 2
birds, and only 3 hunters (4%) succeeded in harvesting the bag limit of 3
birds.
Exact number of birds harvested from the study area is uncertain.
However,
only 22 wings were obtained from the Eby Creek area. The Eby Creek road is
approximately 26 km (16 mi) long of which about 3 kID traverses the study
area. Thus, it is highly unlikely all 22 birds were shot on the study area.
Even so, this would only represent about 17 percent of the estimated fall
population (132 birds).
No band recoveries were reported from the Eby
Creek area. Age and sex composition of harvest samples from 1975 to 1977
is given in Table 9. Inadequate samples of wings were collected in 1975 and

�Table 8.

Age and sex composition of harvest samples from Middle Park, 1975 to 1977.

Male

1/
Subadu1tsFemale

Total

Male

Immatures
Female

Total

Total
Birds

Year

Male

Adults
Female

1975

30(42)1/

41(58)

71 (42)

5(25)

15(75)

20(12)

36(46)

43(54)

79(46)

170

1976

68(51)

64 (49)

132(39)

15 (35)

28(65)

43(12)

74(45)

92(55)

166(49)

341

1977

133(48)

144(52)

277(46)

3(16)

16(84)

19(3)

164(53)

143(47)

307(51)

603

Total

1/ Because some subadu1ts had completed their molt by mid- to late September and thus could not be distinguished
from adults, number of birds assigned to this age class was minimal.
..,..I

1/ Number in parentheses represents percent.
Table 9.

0\
I

Age and sex composition of harvest samples from the Eagle area, 1975 to 1977.

Male

1/
Subadu1tsFemale

Total

Male

Immatures
Female

Total

Total
Birds

Year

Male

Adults
Female

1975

6 (43)1/

8(57)

14(67)

0

0

0

4(57)

3(43)

7(33)

21

1976

7(50)

7 (50)

14(21)

0

8(100)

8(12)

15 (33)

30(67)

45(67)

67

1977

87 (57)

65(43)

152(45)

10(71)

4(29)

14(4)

81(47)

90(53)

171(51)

337

Total

1/ Because some subadu1ts had completed their molt by mid- to late September and thus could not be distinguished
from adults, number of birds assigned to this age class was minimal.
1/ Number in parenthese represents percent.

�-47-

1976 to permit reliable analyses or comparisons among years.
As with the
Middle Park data, few yearlings were identifiable and thus were underrepresented in the samples.
Age and sex composition of the harvest in 1977
was similar between the Eagle and Middle Park areas.
Sex ratios for adults
and immatures did not deviate significantly from 1:1. Production and survival of young in 1977 was good.
Hunter Questionnaire
A total of 1,502 big game hunters were surveyed at the Idaho Springs check
station during the 1977 deer (500), elk (500), and combined deer-elk (502)
seasons.
This survey was conducted in order to evaluate the combined blue
grouse-big game season. Results of the survey are summarized in Table 10.
Hunters were contacted opportunistically at the check station.
Only one
hunter from each party was surveyed.
To insure a diversity of hunters were
questioned, sampling periods included days at the beginning, middle, and end
of each big game season. Few non-residents «5%) were included in the sample.
From data in Table 10, it is apparent that few big game hunters (6.3%)
actually took advantage of the longer grouse season.
Even through participation was low, general response of hunters to the combined blue grouse-big
game season was highly favorable.
The primary complaint of those hunters
opposed to the season (11%) was their concern about the potential increase
in number of hunters (big game + grouse hunters).
However, of the 1,502
hunters contacted, only 2 (.13%) were strictly grouse hunters.
Most grouse
were observed during the deer and elk seasons with a substantial decline in
sightings and harvest occurring during the combined season.
By opening of
the combined season, most grouse had reverted to aboreal habits; consequently,
they were harder to find.
Surveys in 1976 (Hoffman 1977) and 1977 produced similar results.
It is
interesting even though more hunters were aware of the longer grouse season
in 1977 (48%) than in 1976 (23 percent), participation remained relatively
constant from 1976 (4.8%) to 1977 (6.3%). Therefore, only a small percentage
of hunters were interested in hunting grouse during the big game season
regardless of awareness.
It is apparent from data in Table 10 that about half the big game hunters
(48.1%) were aware of the combined blue grouse-big game season, and only
13.1 percent of those aware actually hunted grouse.
Of" those hunting grouse,
27.4 percent were successful in harvesting 53 grouse, for a hunter success
of 2.? birds per successful hunter (.6 birds/big game hunter hunting grouse).
ApplYl~g these estimates to the total big game hunter population gives the
followlng crude estimate of total harvest of blue grouse during the big
game seasons:
Total big game hunter population
48.1 percent aware of grouse season
13.1 percent participation
27.4 percent successful
2.0 grouse per successful hunter
Plus 5 percent crippling loss

226,000
108,706
14,240
3,902
7,804
8,194 Total blue grouse
harvested.

�Table 10.

Combined blue grouse-big game season, hunter questionnaire data, Idaho Springs 1977.

No. in sample
No. big game hunters aware of season
Percent big game hunters aware of season
No. big game hunters unaware of season
Percent big game hunters unaware of season
No. big game hunters unaware that would have hunted
Percent big game hunters unaware that would have hunted
No. big game hunters unaware that would not have hunted
Percent big game hunters unaware that would not have hunted
No. big game hunters hunting grouse
Percent big game hunters hunting grouse
Percent big game hunters aware of season that hunted grouse
No. big game hunters not hunting grouse
Percent big game hunters not hunting grouse
Percent big game hunters aware of season that did not hunt
grouse
Total grouse observed
Percent big game hunters observing grouse
Grouse observed/big game hunter
Total grouse harvested
Percent successful grouse hunters
Grouse/big game hunter hunting grouse
Grouse/successful grouse hunter
Percent hunters favorable of combined blue grouse-big game
season
Percent negative
Percent indifferent

Deer
Season

Elk
Season

Combined
Season

Total All
Seasons

500
253
50.6
247
49.4
78
31.6
169
68.4
40
8.0
15.8
460
92.0

500
260
52.0
240
48.0
90
37.5
150
62.5
25
5.0
9.6
475
95.0

502
210
41.8
292
58.2
66
22.6
226
77 .4
30
6.0
14.3
472
94.0

1,502
723
48.1
779
51.9
234
30.0
545
70.0
95
6.3
13.1
1,407
93.7

84.2
693
24.8
1.4
13
20.0
.3
1.6

90.4
2,102
41.8
4.2
28
52.0
1.1
2.1

85.7
620
18.5
1.2
12
16.7
.4
2.4

86.9
3,415
28.4
2.3
53
27.4
.6
2.0

52.0
13.4
34.6

52.2
9.2
38.6

52.0
10.4
37.6

52.1
11.0
36.9

I
.j:-.

co
I

�-49-

An estimate of the total number of big game hunters as derived from the
Big Game Questionnaire Survey is not presently available for 1977. Therefore, the estimate derived from the 1976 survey was used. Crippling loss
is based on the regular season estimate of 5.0 percent as determined from
data collected on hunter check stations.
Calculations do not take into
account those individuals that hunted blue grouse, but were not big game
hunters.
However, few such individuals were contacted and it is believed
their numbers were minimal.
Based on an 11 year (1966-1976) average annual harvest of 22,100 blue grouse,
the additional harvest (8,194 birds) resulting from the longer season
amounted to about 37 percent increase in the total kill. Estimated harvest
of blue grouse in 1977 was 30,300 birds (pers. comm. H. Riffle) or approximately 11 percent of the estimated total population of 281,000 grouse.
Zwickel (1958) and Bendell and Elliott (1967) suggest that 25 to 30 percent
of the fall population can be safely harvested with no adverse effects on
the subsequent spring breeding population.
Pre- and post-seasons estimates
of breeding densities in 1976 and 1977 have shown no significant fluctuations
as a result of the longer grouse seasons.
It is therefore concluded that
from a biological standpoint, a more liberal blue grouse season is a sound
management practice.

LITERATURE

CITED

Bendell, J. F., D. G. King, and D. H. Mossop.
1972.
lation of blue grouse in a declining population.
36(4):1153-1165.

Removal and repopuJ. Hildl. Manage.

_____ , and P. H. Elliott.
1967. Behavior and the regulation
blue grouse.
Can. Hildl. Servo Rep. Ser. 4. 76 p.
Braun, C. E. 1971.
characteristics.
No. 86. I} pp.

of numbers

in

Determination of blue grouse sex and age from wing
Colorado Div. Game, Fish and Parks.
Game Info. Leaflet

Hoffman~ R. H. 1976. Population dynamics and habitat relationships of
blue grouse.
Colorado Div. Hildl. Job Prog. Rept. Fed. Aid Proj. H-37-R.
April 1976. p. 135-152.
1977. Population dynamics and habitat relationships of blue grouse.
Colorado Div. Hildl. Job Prog. Rept. Fed. Aid Proj. H-37-R.
April,
1977. p. 83-104.
Rogers, G. E. 1968. The blue grouse in Colorado.
and Parks.
Tech. Publ. No. 21. 63 p.
Zwickel, F. C. 1958. North-central
Game Bull. 10(4):3-4.

Hashington

Colo. Div. Game, Fish

grouse studies.

Hash. State

in an increasing

Hildlife

Researcher

��April

-51-

JOB FINAL

State

of

C_O_L_O_RAD
__ O

Project No.

Job Title
Period

_
Game Bird Survey

Organization,

5

Job No •.

17

Covered:

Personnel:

REPORT

W-37-R-3l

Work Plan No.

1978

-------------------------------------

Mortality,

1 April

and Dispersal

1975 to 31 March

of Grouse

Broods

1978

William W. Mautz, Colorado State University; Clait E. Braun,
Deborah Covic, Kenneth Giesen, Richard Hoffman, and James
Kitzmiller, Colorado Division of Wildlife.

ABSTRACT
The objectives of this study have been fully achieved.
Reports covering
the various objectives have been published, prepared and submitted, and
in some instances are in progress.
Those in progress will be published
under Work Plan 22, Job 1 and are listed in the project documents for
W-37-R-32.
Manuscripts published, submitted or in progress are listed below.
1977.
Mortality
Giesen, K. M.
M
..S.
Thesis.
ptarmigan.
1978.

Egg retrieval

and dispersal of juvenile white-tailed
Colorado State Univ., Fort Collins.
55 pp ,

by incubating

white-tailed

ptarmigan.

Auk

(In Press).
_____ , and C. E. Braun.
1976.
Renesting of white-tailed
ptarmigan
Colorado.
J. Colo.-Wyo. Acad. Sci. 8(1):71-72.
(Abstract).
_____ , and1978.

Renesting

of white-tailed

ptarmigan

in

in Colorado.

(Manuscript

submitted

, and

1978.
A technique for age determination
of juvenile
ptarmigan in Colorado.
(Manuscript submitted to J. Wildl.

-----white-tailed

to Condor).

Manage.).
_____ , and
Colorado.
_____ , and
Colorado

1978.
Dispersal of juvenile
(Manuscript in preparation).
1978.
(Manuscript

Mortality

of juvenile

white-tailed

ptarmigan

in

white-tailed

ptarmigan

in

in preparation).

_____ , and T. A. May.
ptarmigan in Colorado.

1978.
Nesting ecology of white-tailed
(Manuscript submitted to Wilson
Bull.).

�-52-

_____ , and
1978. Growth and development of juvenile whitetailed ptarmigan in Colorado. (Manuscript in preparation).
_____ , and R. K. Schmidt, Jr. 1978. Nesting behavior of female
white-tailed ptarmigan in Colorado. (Manuscript submitted to Condor).
, and
1978. Summer and fall movements of juvenile
----white-tailed ptarmigan in Colorado. (Manuscript in preparation).

Prepared by

~1f1

~

Kenneth M. Giesen
Graduate Research Asst.

Approved

by_~tk~~J-L!.:_:?_''---Li--=:~:....::-:..--=__
Clait E. Braun
Wildlife Researcher

�April 1978
-53-

JOB FINAL REPORT

State of

COLORADO

----------------------

Project No.

W-37-R-3l

Work Plan No.

17

Game Bird Survey

Job No.

Job Title

Evaluations of White-tailed
Ptarmigan Introductions to Pikes Peak

Period Covered:

April 1, 1976 to March 31, 1978

Personnel:

6

George Bock, Clait Braun, Frank Colley, Kellan Duncan, Marcus
Elkins, Howard Funk, Kenneth Giesen, Richard Hoffman, James
Kitzmiller, Thomas Lytle, Walter Larrick, and James Morris.

ABSTRACT
Division of Wildlife personnel concluded after 3 days of intensive searching
in May 1974 that no breeding population of white-tailed ptarmigan (Lagopus
leucurus) was present on Pikes Peak, but preliminary habitat investigations
suggested the area could potentially support ptarmigan in small numbers.
In 1975, 6 pair were released in June, followed by another release of 9
males, 7 females, and 12 chicks in September.
Breeding and production
surveys were conducted in 1976 and 1977. Reproduction occurred the first
year (1975), but was insufficient to replace natural losses, consequently,
the population declined by 42 percent from 1975 to 1976. Production and
survival of young in 1976 and 1977 were good. However, rate of population
increase was only 8.7 percent from 1976 to 1977. Overall survival increased
from 55 percent (1975 to 1976) to 69 percent (1976 to 1977). While
ptarmigan have been successfully introduced on Pikes Peak, population
increase and expansion into new areas has been slow. At the present rate
of increase, 10 to 12 years will be required for the population to attain
maximum supportable densities.
Guidelines for successful establishment of
ptarmigan into new or unoccupied habitats are discussed.

�-54RECOMMENDATIONS

1.

It is recommended
to Pikes Peak.

that no further transplants

of ptarmigan

be made

2.

It is recommended that data obtained in this study be turned over to
management personnel in 1978 for future monitoring of the population
for at least 2 or 3 more years.
Some assistance should be provided
by research in 1978 to ensure census procedures are properly understood and utilized.

3.

Guidelines for successful introduction of white-tailed
new or unoccupied habitats will be published.

4.

Data from Work Plan 17, Job 6 should be incorporated with data from
Work Plan 17, Jobs 1, 2, and 3 and studies by Braun et al. (1978)
into a manuscript entitled "Experimental Removal and Tranplant Studies
of White-tailed Ptarmigan."

ptarmigan

into

�-55-

EVALUATIONS OF WHITE-TAILED
PTARMIGAN INTRODUCTIONS TO PIKES PEAK
Richard W. Hoffman

Unlike the erratic distribution of white-tailed ptarmigan in most states
and provinces within its range, this grouse inhabits practically all alpine
areas in Colorado, with the exception of Spanish Peaks and Pikes Peak.
Absence of ptarmigan On Spanish Peaks can be attributed to an almost
complete deficit of vegetation above treeline; whereas, Pikes Peak, with
its seemingly suitable habitat, apparently has never supported a breeding
population of ptarmigan.
Aiken and Warren (1914) and Knorr (1959), in
studying the avifauna of El Paso County noted the absence of ptarmigan on
Pikes Peak and reported no authentic records of this alpine grouse occurring
in the County.
No reports were mentioned of ptarmigan on Pikes Peak in
early distributional records summarized by Schlater (1912) and more recently
by Bailey and Neidrach (1965). Using developed techniques for locating
ptarmigan, Division of Wildlife personnel similarly confirmed after 3 days
of intensive searching in May 1974 that no ptarmigan were present on Pikes
Peak (Hoffman unpubl. data).
Furthermore, due to the geographically
isolated nature of Pikes Peak from other occupied ranges, it is highly
unlikely ptarmigan ever reached the area in sufficient numbers to establish
a population.
Intensive studies of white-tailed ptarmigan in Colorado have focused on
nearly every biological aspect of this species (Schmidt 1969, Braun and
Rogers 1971, Braun and Schmidt 1971, Haskins 1969, May 1975, May and Braun
1972, Hoffman and Braun 1975, and Giesen 1977). While removal studies
have provided information on natural repopulation of previously occupied
habitats (Braun 1975), little is known concerning characteristics
of a
population when introduced into new areas.
Prior transplants have been
made to other states including Oregon, California, and Utah following
established guidelines set forth by Braun (1975). With the exception of
the Utah transplant, (Braun et al. 1978) circumstances have not permitted
evaluation of the introductions or adequacy of the guidelines.
The Pikes
Peak introduction represents the first opportunity within Colorado to
conduct fol1owup studies.
This report describes the Pikes Peak transplant.
P. N. OBJECTIVES
Major objectives of this study are to: (1) evaluate the success or failure
of transplanted ptarmigan to establish a breeding population on Pikes Peak,
(2) ascertain if distance of Pikes Peak from other occupied habitats
prevented males (but not females) from reaching the area and consequently
establishing a breeding population, (3) identify critical deficiencies
in the habitat (including minimum size) which might contribute to a failure
of ptarmigan to establish a breeding population, (4) evaluate adequacy of
existing guidelines for ptarmigan introductions, possibly improve upon
them and/or develop additional guidelines, and (5) increase knowledge
available on the pioneering abilities of ptarmigan.

�-56-

METHODS AND MATERIALS
Status of ptarmigan on Pikes Peak prior to the introduction was ascertained
by (1) field reconnaissance,
(2) review of avifauna records, and (3)
personal communications with Division of Wildlife personnel and other
persons acquainted with the area. Vegetative communities were visually
described and classified as to their suitability for ptarmigan based on
habitat requirements of the species as presented by Braun (1971). Using
data from earlier studies (Whitfield 1933, Buechner, unpubl.) along with
information collected in this study, vegetative types were described
according to procedures outlined by Braun (1969). Plant nomenclature
follows Weber (1972). Transplant procedures follow established guidelines
set forth by Braun (1975). Climatological data were obtained from Whitfield
(1933), U. S. Department of Commerce, Weather Bureau (1920-1934), Marr
(1961), and Marr et al. (1968a and b).
Both a spring breeding census (May-June) and summer brood survey (AugustSeptember) were conducted in 1976 and 1977. Techniques employed for locating
ptarmigan have already been described (Braun et al. 1973), along with
efficient trapping and identification procedures (Braun and Rogers 1971).
Attempts were made to identify all marked birds and to capture and band any
unmarked individuals.
Each sighting was recorded at the time on standardized observation cards, and locations were plotted on 7.5 minute U. S.
Geological Survey topographic maps. Most of the Pikes Peak region was
traversed in order to determine distribution of introduced birds and to
delineate potential seasonal use sites. All areas appearing suitable for
use by ptarmigan were outlined on U.S.G.S. topographic maps and subsequently
planimetered to estimate amount of habitat available.
DESCRIPTION

OF STUDY AREA

The Pikes Peak region lies in the east central part of Colorado along the
Front Range of the Rocky Mounains at approximately 38030' north latitude
and 1060 west longitude.
It is included within the Pike National Forest
in portions of El Paso and Teller counties (T14S, R68 and 69 W, and T15S,
R69W) and represents the eastern most extension of alpine range in Colorado.
Total area classified as alpine includes about 39 km2(15 mi2); however,
the area under investigation encompasses 23 km2(9 mi2) (Fig. 1). Location
of Pikes Peak in relation to ptarmigan distribution and alpine range in
Colorado is shown in Fig. 2.
Geologically, Pikes Peak is similar to much of the alpine area in Colorado.
The core of the front range in this area consists mainly of Pikes Peak
granite, a natural, intrusive, igneous rock that was forced through the
overlying sedimentary rock during the Laramide Revolution.
Most of the
sedimentary rock has since been eroded away, leaving the granite exposed
(Koschmann 1956). Although the major rock type is granite, metamorphic
rocks such as schist, quartzite, and gneiss are present.
With the exception
of quartzite, soils developing from these rocks are productive and can be
classified as coarse, loamy, and deep. However, some are occasionally
shallow to bedrock.
Soils termed by Nimlos and McConnell (1965) as alpine
meadow and alpine bog types occur frequently in wet, poorly drained sites.

�ORE

~ T

KEY

o

o

'\el Potnt

o

X

ION

p

o
e

A

Fall Release Site
Spring Release Area
Territory 1976
Territory 1977
Territory 1976 and 1977

-t
N

/

PIKES PEAK AREA
Study Area Boundary
Scale in Kilometers

o

1/2

\
\

i

Fig. 1.
breeding

Pikes Peak study area showing
territories.

release

sites and location

of

1

�~

-CRAIG

•FORT COLLINS

J}
-DENVER

I
VI
00
I

-GRAND
JUNCTION
~

o

• COLORADO SPRINGS
.PIKES PEAK

GUNNISON -

N

f)

-DURANGO
Fig. 2.

Location

of Pikes Peak in relation

b
flcP

SCALE IN KILOMETERS
50
tOO

o

I

I

J

-TRINIDAD

to alpine range and ptarmigan

distribution

in Colorado.

�-59-

Topography is irregular, with elevations ranging from 3,500 m (11,500 ft)
at treeline to 4,300 m (14,109 ft) on the summit of Pikes Peak. The study
area lies on the southwest side of a ridge about 10 km (6 mi) long. The
southwest side of this ridge has a moderate slope, and merges into a
gently rolling upland.
This is in striking contrast to the northeast side
where glacial action has created a rugged precipice with steep walled
cirques, rock heaps and U-shaped valleys (Pearl 1964).
Vegetation
Examination of vegetative communities and available habitats prior to the
transplant indicated that alpine areas on Pikes Peak and adjacent mountains
were suitable for white-tailed ptarmigan.
A description of these areas is
presented below.
For descriptive purposes, the alpine region was divided into the low and
high alpine.
The low alpine contains many of the woody plants of the
alpine zone, and lies approximately between timberline and 3,811 m (12,500
ft). The most conspicuous dominant of the low alpine is Salix spp., but
krummholz communities alternately dominated by clumps of Picea engelmannii
or bushes of Salix spp. are major vegetative components of the low alpine.
Many of the drainage systems originating in the alpine have been blocked by
glacial action resulting in extensive wet areas dominated by dense stands
of Salix spp. These areas frequently extend from above treeline into the
subalpine.
Other plants locally abundant in the low alpine include: Pinus
flexilis, Pinus aristata, Dryas octapetala, Pentaphylloides floribunda,
Kobresia bellardii, and Carex spp.
Further upslope, between the upper limits of the low alpine and the higher
talus slopes and ridges, are the alpine meadows which are transitory sites
between the low and high alpine.
Salix spp. extend into the lower portions
of the meadows, while rocky areas predominate along the upper portions.
Most of the area is characterized by low growing vegetation classified as
Kobresia- Carex-Geum and Kobresia-Carex alpine meadows.
Deschampsia,
Polygonum, Potentilla, Pedicularis and Hymenoxys are some of the other
plants found throughout the meadows.
The high alpine includes the steep slopes and ridge tops above 3,963 m
(13,000 ft). Obvious features of the high alpine are the abundance of
rocky areas and exceedingly low growing vegetation.
Vegetative communities
consist primarily of Carex rock, Carex Poa rock, and Carex-Geum rock
meadows, but Trifolium, Senecio, Kobres~
and Sedum are also present in
certain areas. Large boulder fields are common throughout the area,
especially at higher elevations.
Some have little or no vegetative cover.
Similarly, many of the precipitous slopes in the northeast portion of the
area are devoid of vegetation.
Climate
Recent weather data were lacking for the study area. Some recent data were
available for subalpine sites near the study area, but the information was
incomplete.
Consequently, reference is made to data collected from 1920 to

�-60-

1934 by the U. S. Forest Service, Fremont Experiment Station, at a treeline site approximately 4 km (2.5 mi) east of the study area (Table 1).
Wind volocities are excluded from Table 1 because data were recorded in
mph of wind per day with no maximum, minimum or mean figures.
Comparative
data were obtained from long term climate investigations conducted by Marr
et al. (1968a, 1968b) at Niwot Ridge (40003' N lat. and 105037' W long.)
(Table 2). This site lies 120 km (75 mi) north of the study area along
the Front Range.
Although climatic data for alpine sites are available from studies by Marr
et al. (1968a, 1968b) these data are not directly comparable with treeline
data collected near the study area. Therefore, data presented in Table 2
also represent climatic conditions at treeline.
It should be noted that
alpine air temperatures are usually 2 to 70C (4 to l30F) lower than
temperatures in adjacent subalpine areas (Marr 1961). Amount of variation
is dependent upon elevational extremes, with minimum temperatures showing
the least difference (Marr 1961). Conversely, precipitation increases from
subalpine to alpine areas, but the precise increment is unknown (Marr 1961).
A detailed review of climatic studies in Colorado alpine areas and general
descriptive information is presented by Braun and Rogers (1971).
Climatic conditions on Pikes Peak are similar to other alpine regions in
the state, and can best be described as continental.
Frequent extremes
in wind velocities and temperature are common.
Prevailing winds are
westerly.
The alpine spring is short, cool, and wet. Upslope easterly
winds typically occur in spring, and when accompanied by moisture, result
in heavy, wet snowfalls (Marr 1961).
Summers are short, and cool to warm,
with most precipitation occurring in form of thunderstorms.
Autumn is
short, clear, dry, and mostly cold; however, extended periods of warm
weather are not unusual.
The winter is long, very cold, dry, and windy.
Blizzards are frequent, but much of the moisture occurring in winter is
ineffective because constant winds blow most of what falls as snow from
exposed sites.
Upon examination of Tables 1 and 2 it is apparent warmer, drier conditions
prevailed on Pikes Peak as compared to Niwot Ridge. Monthly maximum,
minimum, and mean temperatures near the study area averaged 30C (50F)
warmer, while average annual precipitation was 9.1 cm (3.6 in) less than
the same estimate for Niwot Ridge.
Forty seven percent of the annual
precipitation (56.57 cm, 22.28 in) on Pikes Peak occurred from June to
August (26.49 cm, 10.43 in), with fall through late winter (September to
February) being the driest period (13.08 cm, 5.15 in). Only 29 percent of
the annual precipitation (65.66 cm, 25.86 in) at Niwot Ridge fell from June
to August (19.00 cm, 7.48 in), whereas, over 10 cm (4 in) more preC1p1tation accumulated from September through February (24.79 cm, 9.76 in).
Spring was the wettest season at Niwot Ridge.
RESULTS AND DISCUSSION
Releases
The initial work of trapping and transplanting ptarmigan to Pikes Peak was
completed in 1975. Ptarmigan were trapped from 5 different areas in

�Table 1. Climatological data, Pikes Peak, IS-year average from 1920 to 1934 (U. S. Dept. Commerce, Weather
Bureau. Climatological data, Colorado annual summaries).

Month

TemEerature, Air °c (OF)
Mean
Minimum
Maximum

Maximum

Precipitation, cm Water (in)
Minimum

Mean

January

15 (58)

-24 (-11)

- 4 (24)

3.22 (1.27)

.05 ( .02)

1.01 ( .40)

February

16 (60)

-34 (-29)

- 4 (25)

5.28 (2.08)

.28 ( .11)

1.85 ( .73)

March

16 (61)

-26 (-14)

- 3 (27)

11.81 (4.65)

.99 ( .39)

4.19 (1.65)

April

18 (65)

-20 (- 3)

1 (34)

18.34 (7.22)

2.06 ( .81)

5.69 (2.24)
I
(J\

May

23 (73)

- 8 ( 17)

6 (43)

14.88 (5.86)

2.26 ( .89)

7.14 (2.81)

June

27 (80)

- 2 ( 29)

12 (53)

20.75 (8.17)

.96 ( .38)

6.35 (2.50)

July

29 (83)

o ( 32)

15 (58)

19.63 (7.73)

4.29 (1.69)

10.18 (4.01)

August

27 (81)

- 1 ( 31)

13 (56)

18.77 (7.39)

2.82 (1.11)

9.96 (3.92)

September

26 (78)

-12 ( 11)

11 (51)

6.45 (2.54)

1.17 ( .46)

3.17 (1.25)

October

23 (73)

-15 ( 5)

5 (41)

12.70 (5.00)

.05 ( .02)

3.02 (1.19)

November

17 (62)

-26 (-14)

o (32)

8.69 (3.42)

.33 ( .13)

2.46 ( .97)

December

15 (58)

-30 (-22)

- 4 (25)

3.94 (1.55)

.10 ( .04)

1.55 ( .61)

Average

21 (69)

-16 (

3)

4 (39)

12.04 (4.74)

1.28 ( .50)

56.57 (22.28)

f-'

I

�Table 2.

Climatological data, Niwot Ridge, 11-year average from 1953 to 1964 (Marr et a1. 1968 a and b).

Month

Temperature, Air °c (OF)
Mean
Minimum
Maximum

January

12 (54)

-37 (-34)

- 7 (19)

8.38 (3.30)

.1.27 ( .50)

4.39 (1.73)

February

12 (53)

-32 (-25)

- 7 (19)

6.86 (2.70)

1.02 ( .40)

4.42 (1.74)

March

12 (54)

-29 (-19)

- 6 (21)

9.02 (3.55)

1.27 ( .50)

5.69 (2.24)

April

17 (62)

-18 (- 1)

- 3 (27)

22.48 (8.85)

1.40 ( .55)

7.77 (3.06)

May

21 (69)

-16 ( 4)

4 (39)

18.54 (7.30)

3.68 (1.45)

8.43 (3.32)

Maximum

Precipitation, cm Water (in)
Mean
Minimum

I

June

25 (77)

- 7 ( 19)

9 (48)

9.40 (3.70)

.25 ( .10)

5.46 (2.15)

July

25 (77)

- 2 ( 28)

..
12 (53)

9.75 (3.84)

3.61 (1.42)

6.70 (2.64)

August

25 (77)

- 3 ( 27)

11 (52)

18.11 (7.13)

1.65 ( .65)

6.83 (2.69)

September

25 (76)

-14 (

7)

8 (46)

13.16 (5.18)

.35 ( .14)

5.03 (1.98)

October

18 (65)

-16 ( 4)

4 (39)

6.35 (2.50)

.51 ( .20)

2.79 (1.10)

November

13 (55)

-27 (-17)

- 3 (27)

7.75 (3.05)

2.41 ( .95)

3.96 (1.56)

December

13 (55)

-26 (-15)

- 6 (22)

9.14 (3.60)

.38 ( .15)

4.19 (1.65)

Average

18 (64)

-19 (- 2)

1 (34)

11.58 (4.56)

1.48 ( .58)

65.66(25.86)

'"

N
I

�-63-

Colorado (Guanella Pass, Hagerman Pass, Weston Pass, Loveland Pass, and
Mosquito Pass) to insure genetic diversity in the stock of transplanted
birds.
Two transplants (spring and fall) involving a total of 40 birds
were made (Table 3). Six mated pairs were released in June followed by
another release of 9 males, 7 females, and 12 chicks in September.
All
birds were released within 1 1/2 days of when they were trapped.
No
mortalities occurred as a result of transplant operations.

Table 3.

Age and sex composition

Release
Period

of spring and fall releases

on Pikes Peak.

Sex

Adults

Yearlings

Immatures

Total

Males

3

3

0

6

Females

2

4

0

6

Males

6

3

9

18

Females

3

4

3

10

14

14

12

40

June 1975

September

Totals

1975

Release sites were selected based on (1) spring and fall habitat requirements of the species (Braun 1971), (2) proximity of other seasonal use areas
(fall and winter habitat) in relation to the release sites, and (3) accessibility of release sites. Both spring and fall release sites were in the
same general locality, but the fall release site was at a slightly higher
elevation.
During the spring transplant, birds were released as mated
pairs at approximately 180 m (200 yds) intervals apart.
The female was
released first, immediately followed by the male. No special precautions
were taken in the fall release except for releasing chicks in close proximity to a hen. The remaining birds were turned loose in groups of 4 to 5
birds. All birds were marked prior to release and pertinent information
about each bird was recorded.
Within a few days following their introduction, spring released cocks spaced
themselves out, established territories, and resumed breeding activities
with their original mate.
Other than normal spacing of territories, no
extensive movements occurred away from the release site. Fall released
grouse exhibited no unusual behavior when released.
Some birds slowly

�-64-

wandered off into surrounding rocky areas.
Others flew short distances
«100 m), while a few birds flew several hundred meters and disappeared
from view.
No further attempt was made to monitor the population until 1976.
Establishment

and Dispersal

Observations of sign indicated many birds wintered in stands of willow
(Salix spp.) near the release sites, but at lower elevations.
Distances
from release sites of ptarmigan observations in 1976 and 1977 are shown in
Table 4. No birds were observed more than 3.4 km (2.1 mi) from the release
areas, but sign was found up to 6 km (4 mi) away.

Table 4. Distance from release
Pikes Peak area. 11

Distance
(km)

Spring
Release

sites of ptarmigan

1976 Observations
Fall
Release
Progeny

Total

Spring
Release

observations,

1976-77,

1977 Observations
Fall
Release
Progeny

Total

&lt;0.5

2

1

0

3

1

0

0

1

0.5-0.9

5

3

0

8

4

2

1

7

1. 0-1.4

1

0

0

1

1

0

2

3

1. 5-1. 9

0

0

0

0

0

0

1

1

2.0-2.4

0

4

0

4

0

2

0

2

2.5-2.9

0

1

0

1

0

0

3

3

3.0-3.4

0

5

1

6

0

5

3

8

1lExc1udes duplicate

observations.

Release sites and location of breeding territories are depicted in Fig. 1.
Whereas, the entire study area encompasses 23.4 km2 (9.1 mi2), only 27.6
percent (6.5 km2) constitutes suitable breeding habitat.
Eighty percent of
the breeding habitat and consequently, 91 percent of the birds were in
the southern portion of the study area in 1976. This decreased to 80 percent
in 1977 as more birds moved into the northern part of the study area.
Even
so,dispersa1 was slow as average distance of territories from release sites
increased by only .3 km from 1976 to 1977. Also, progeny of the 1976 population did not pioneer into new areas in 1977 which further reflects the
slow rate of dispersal.

�-65-

Observations of transplant birds during 1976 revealed that spring released
birds were clustered around the release sites while fall released birds
had spread out into surrounding breeding areas.
Innately, ptarmigan form
the same pair bond and return to the same territory year after year (Schmidt
1969).
Therefore, it was not unusual for surviving spring released birds
to be breeding on the same area (release site) in 1976 where they had
established territories in 1975. As a result of this behavior, and because
fall released birds did not establish territories until spring 1976, they
were forced to search elsewhere for breeding habitat.
Mean distance of
territories from release sites for fall released males was 2.48 km (range
.79 - 3.36 km) compared to a mean distance of only .73 km (range = .41 - .99
km) for spring release males.
As expected, fall released juveniles and subadults dispersed the greatest
distances from release sites. Adult males, especially those released in
spring, were the most sedentary portion of the population.
Locations of
their territories in relation to release sites indicate they traversed
the minimum distance necessary to find vacant breeding habitat.
With one
exception, all males established territories within calling distance of
another male.
Movements of spring and fall released females were similar to males
released at the same time. No females were located outside the bounds of
occupied habitat; thus, their movements were at least in part controlled
by where the males established territories.
Due to their high mobility
(Hoffman and Braun 1975) some females may have dispersed off the study
area. Evidence supporting this hypothesis is limited to unconfirmed
slightings of ptarmigan several kilometers from the study area.
Population

Characteristics

Ptarmigan were located during the breeding and brood seasons through use
of tape recorded male challenge and chick distress calls as described by
Braun et al. (1973). Two censuses were conducted each year during both
the breeding (May-June) and brood (July-early September) seasons.
All
potential habitat was systematically searched.
Nevertheless, some
fraction of the population undoubtedly escaped detection, but it is
believed most breeding birds were located.
Results of breeding surveys in 1976 and 1977 are summarized in Table 5.
Estimates of nesting success and production could not be accurately ascertained based on summer field observations because of difficulties encountered in finding hens with chicks.
However, number of juveniles located
during spring breeding surveys provided some indication of the previous
year's production and survival of young.
In 1976 the spring population consisted of 23 birds, including 8 pairs,
4 unmated
territorial males, and 3 non-territorial males.
No unmated hens
were found. Of 23 grouse observed, 22 were already marked and were remaining
individuals from the introduced population.
The unbanded bird was a juvenile hen which paired with a fall released male.
Obviously this hen was
not an introduced bird and most likely represents surviving offspring of
the 1975 spring released birds.

�-66-

Table 5. White-tailed ptarmigan spring population
status on Pikes Peak 1976 and 1977.

densities

and breeding

Year

Total
Spring
Population

Breeding
Pairs

Unmated
Territorial
Males

NonTerritorial
Males

Unmated
Females

1976

23

8

4

3

o

1977

25

9

5

2

o

Percent
Rate of
Increase

8.7

Annual survival of the introduced population from 1975 to 1976 was similar
to survival in a natural population (Table 6). However, production and
survival of young in 1975 was insufficient to replace natural losses from
the population, consequently, from 1975 to 1976, there was an initial
population decline of 42.5 percent.

Table 6. Comparative survival rates of the introduced population and a
natural population of white-tailed ptarmigan in Rocky Mountain National Park.

1-./

Survival Rates

(%)

Age and Sex

RMNP

Adults

67

64

Males

75

78

Females

65

40

Subadults

65

64

Males

75

100

Females

55

37

J uvena·1es-3/

23

33

56

55

Annual

Survival

All Birds

1/ Data from Braun

1/ Survival

(1969).

of spring and fall released

1/ Sex of juveniles

Pikes Peak ])

birds from 1975 to spring 1976.

could not be accurately

ascertained

at time of release.

�-67-

Survival of ptarmigan varies with age class and is lowest in the first year
of life (0-1 years) and increases substantially during the second year
(1-2 years) to a constant level that is maintained in the older age classes
(Braun 1969). Also, males typically have a higher survival rate than
females because of their more sedentary habits (Braun 1969).
These characteristics are suggested by the data presented in Table 6 with the exception
that survival of adult and subadult females appeared to be abnormally low.
Since survival rates of the other age and sex classes compared favorably
with the natural population, the excessive loss of females could not be
entirely contributed to mortality on the study area. Due to the high
mobility of females (Hoffman and Braun 1975) and limited amount of suitable
habitat on Pikes Peak, more females were probably lost from the introduced
population as a result of dispersal off the area without replacement than
would normally occur in a natural population.
Any movement of birds away
from the Pkes Peak area almost certainly resulted in their death as there
is no other habitat within 65+ km (40+ mi).
A total of 17 ptarmigan (11 males, 2 unsuccessful females, and 1 successful
female with 3 chicks) were located during summer brood investigations in
1976. Only 3 of 8 hens observed on territories were accounted for during
the brood census.
Status of the other 5 hens is uncertain, but it is doubtful if they succumbed to mortality and most likely they went undetected.
Based on the actual number of hens observed (3), nesting success for 1976
was only 33 percent.
These data suggest the conclusion that production and
survival of young in 1976 and subsequent recruitment into the population
was low.
However, results of breeding surveys in 1977 were not in agreement with this
conclusion.
Nine juveniles were identified in spring 1977, whereas, only 3
juveniles were originally located during summer brood investigations in 1976.
While just 3 chicks were banded in 1976, 2 (66%) were recaptured in 1977,
which implies excellent survival of chicks.
Assuming a similar survival
rate for unbanded chicks, then at least 14 chicks were produced in 1976.
Since brood size by September 1 does not change significantly between years
and averages 3.5 chicks per brood (Braun 1969), then the 14 chicks produced
in 1976 represent 4 broods and estimated nesting success for 1976 is 50
percent instead of 33 percent.
The population not only benefited from good production jn 1976, but also
was enhanced by excellent survival from 1976 to 1977. Eighteen of 26 birds
(including chicks) identified in 1976 were relocated in 1977 for an estimated
annual survival rate of 69 percent (Table 7). From 1976 to 1977, total
population size increased from 23 to 25 birds (Table 5). Rate of increase
in the population was 8.7 percent.
Mated pairs surviving from the 1976
population formed the same pair bond in 1977. In addition, 10 of 12 territories occupied in 1976 were re-established in 1977 along with 4 new
territories.
The abandoned territories were situated in marginal habitat
and were probably temporary in nature.
Four broods were observed on the Pikes Peak area during August 1977. Brood
sizes were 6, 5, 4, and 4 chicks (average = 4.7 chicks/brood).
Nine chicks
were captured, including one trapping mortality, and based on primary wing
molt, they all hatched between 23 June and 4 July. Except for 2 males, no
other birds were located.
Observation of sign suggested the birds were
scattered in small flocks throughout the area.

�-68-

Table 7.
1977.

Survival

rates of white-tailed

ptarmigan

on Pikes Peak, 1976 to

Survival

Age and Sex

(%)

67

Adults
Males

77

Females

40

Subadults

80

Males

50

Females

100

Juveniles

6~/

Annual

Survival All Birds

69

l/Based

on survival

of 3 chicks banded

in 1976.

Although data are limited, a minimum of 4 of 9 hens (44%), and probably more,
were successful nesters in 1977. An average brood size of 4.7 chicks by
August 1 further indicates good production and survival of young in 1977.
Therefore, it is anticipated that the breeding population will increase in
1978.
CONCLUSIONS
Published references regarding successful introductions of ruffed grouse
(Bonasa umbellus) (Ammann and Palmer 1958; Moran and Palmer 1963), bobwhite
quail (Colinus virginianus)
(Cookingham and Ripley 1964), and ring-necked
pheasant (Phasianus colchicus) (Einarsen 1942, Stokes 1954) all describe
the same basic population trend; rapid dispersal throughout the range and
erruptive buildup of populations followed by a leveling off to more normal
numbers.
While white-tailed ptarmigan have been successfully introduced
to Pikes Peak, events succeeding the release did not conform to the general
pattern described in the lit,erature. By the first breeding season after
introduction, population numbers had declined by 42 percent and since then
the population has only been slowly increasing and expanding into new areas.
In Michigan, only 4 breeding seasons were needed for an introduced population
of ruffed grouse to reach saturation levels (Moran and Palmer 1963).
Estimated suitable breeding habitat on Pikes Peak is 6.5 km2 (2.5 mi2).
At
2
an average breeding density of 8 birds/km2 (20 birds/mi ) (Braun and Rogers
1971), this area can presumably support about 25 pairs.
Assuming the present

�-69-

rate of population increase (8.7%) remains stable, 10 to 12 years will
be required for the population on Pikes Peak to attain maximum supportable
densities.
Due to time constraints and problems inherent with holding wild ptarmigan in
captivity, the initial trapping of birds for release was sometimes opportunistic and age and sex composition of the released stock was not ideal.
As a consequence, a deficiency of adult and yearling hens and an excess of
chicks were included in the fall release.
In addition, only 6 pair were
released in spring 1975 which limited potential production the first year.
The problem was further amplified by abnormally low survival of females,
creating lopsided sex ratios favoring males (1976, 1.9 males:l female;
1977, 1.8 males:lfemale).
Most certainly, the uneven sex ratios coupled
with the lack of experienced birds (especially adult hens) and poor production
in 1975 were not conducive to maximum reproductive output from the population.
Even so, characteristically,
ptarmigan are not prolific breeders (Braun 1969)
and even if conditions were ideal, the limited reproductive capabilities of
the species would prevent rapid, short term increases in the population.
Conversely, species such as ruffed grouse, pheasants, and bobwhites have
tremendous reproductive capabilities under ideal conditions with the potential
of doubling or trippling population numbers in one year.
Failure of birds to rapidly disperse throughout the habitat was a manifestation of the sedentary nature of males (Hoffman and Braun 1975).
Fall
released juveniles were expected to disperse from release sites and extend
the range of the introduced population.
However, few survived until the
following spring and the majority of birds remained concentrated around the
release areas.
Since males banded as chicks return to their natal areas in
search of vacant territories (Braun 1969), it was not surprising that progeny
of the introduced birds also established territories near the release sites.
Some did move into adjacent habitats, but dispersing males did not have to
travel very far before they encountered unoccupied breeding habitat.
Thus,
expansion into new areas was slow, and controlled by the movements of males.
Braun (1975) similarly describes the important role played by males in
extension of range and re-establishment
of ptarmigan populations.
Guidelines
In considering transplants of white-tailed ptarmigan and possibly other
grouse into new or unoccupied habitats, the following guidelines for successful establishment are recommended.
1.

Pre-release evaluation and selection of habitats should be conducted
personnel experienced with the species in native habitats.

by

2.

Release sites should be selected based on the immediate requirements of
the birds when they are released.
Preferably,
other seasonal use sites
should be in close proximity to the release area.

3.

Unless the habitat is ideal, areas smaller than 5 km2 (3 mi2) will
probably not support huntable populations on a sustained basis without
immigration from other occupied ranges.
Otherwise, excessive losses may
occur due to emmigration without replacement.

�-70-

4.

Releases should be made with wild trapped stock of an appropriate
subspecies or geographical
race. Birds trapped from several different
areas will ensure genetic diversity in the released stock.

5.

Large numbers of animals are not necessary for successful establishment
provided (1) the habitat is suitable, (2) all birds are relased in the
same area, and (3) released stock includes mostly adult and yearling
birds.

6.

Spring releases should only include mated pairs.
Ten to 15 pairs should
be adequate.
Timing of the release is very important.
By early June
pair bonds are strongly formed and soon after, ovulation is initiated.
This information, coupled with the knowledge that the pair bond is not
easily broken, and once initiated, ovulation and egg laying cannot be
readily terminated, suggest that spring releases in early June will
ensure cohesiveness of the pair bond, re-establishment of territories,
and completion of the laying cycle.

7.

A combination of yearling and adult birds with sex ratios favoring
females (2:1) should compose the fall (September) release.
Few, if any,
juveniles should be included.
Birds should be released in small groups
of 4 or 5 individuals near the same locality as the spring transplant.

8.

It may be necessary to release additional
supplement losses and enhance production.

9.

Annual breeding and production surveys for 3 to 5 years after introduction are desirable in order to ascertain success or failure of the
transplant and factors responsible.

LITERATURE

females the following year to

CITED

Aiken, C.E.H., and E. R. Warren.
1914. The birds of El Paso County, Colorado I. Colorado College Publ. General Ser. No. 74. 12(13):455-496.
Ammann, G. A., and W. L. Palmer.
1958.
Michigan Islands.
J. Wildl. Manage.
Bailey. A. M •. and R. J. Niedrach.
1965.
Natural History_ Vol. 1. 895 pp.

Ruffed grouse introductions
22(3):322-325.
Birds of Colorado.

on

Denver Museum

Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado.
Ph. D. Dissertation.
Colorado State Univ.,
Fort Collins.
189 pp.
1971. Habitat requirements of Colorado white-tailed ptarmigan.
Proc. Western Assoc. State Game and Fish Comms. 51:284-292.
1975. Experimental removal of a breeding population of white-tailed
ptarmigan.
Colo. Div. Wi1dl., Game Res. Rept., Fed. Aid Proj. W-37-R.
April.
pp. 91-109.

�-71-

Braun, C. E., and G. E. Rogers.
1971. The white-tailed ptarmigan in
Colorado.
Colo. Div. Game, Fish and Parks Tech. Publ. No. 27. 80 pp.
_____ , and R. K. Schmidt, Jr. 1971. Effects of snow and wind on wintering
populations of white-tailed ptarmigan in Colorado.
Pp. 238-250.
In
A. O. Haugen, ed. Proceedings of snow and ice symposium.
Iowa Cooperative Wildl. Res. Unit. Iowa State Univ., Ames.
280 pp.
_____ , and G. E. Rogers.
ptarmigan with tape-recorded

1973. Census of Colorado white-tailed
calls. J. Wildl. Manage. 37(1):90-93.

____ ~' D. H. Nish, and K. M. Giesen.
1978. Release and establishment of
white-tailed ptarmigan in Utah.
Southwest Naturalist.
(In press).
Cookingham, R. A., and T. H. Ripley.
1964. Vital characteristics
insular bobwhite population.
J. Wildl. Manage. 28(4):855-857.

of an

Einarsen, A. S. 1942. Specific results from ring-necked pheasant studies
in the Pacific Northwest.
Trans. N. Am. Wildl. Conf. 7:130-146.
Giesen, K. M.
ptarmigan.

1977. Mortality and dispersal of juvenile white-tailed
M. S. Thesis.
Colorado State Univ., Fort Collins.
55 pp.

Haskins, A. G. 1969. Endoparasites of white-tailed ptarmigan (Lagopus
leucurus) from Colorado.
M. S. Thesis.
Colorado State Univ., Fort
Collins.
147 pp.
Hoffman, R. W., and C. E. Braun.
1975. Migration of a wintering population
of white-tailed ptarmigan in Colorado.
J. Wild1. Manage. 39(3):485-490.
Knorr, O. A. 1959. The birds of El Paso County, Colorado.
Studies, Ser. Biol. 5. 48 pp.

Univ. Colo.

Koschmann, A. H. 1956. Topographic map of Pikes Peak and vicinity, Colorado.
U. S. Geol. Survey map. U. S. Government Printing Office, Washington, DC.
Marr, J. W. 1961. Ecosystems of the east slope of the Front Range in Colorado. Univ. Colo. Studies, Ser. Biol. 8. 134 pp.
1968a. Data on mountain
----- , J. M. Clark, W. S. Osborn, and M. W. Paddock.
environments III. Front Range, Colorado, four climax regions, 1959-1964.
Univ. Colo. Studies, Ser. Biol. 29. 179 pp.

----- , A. W. Johnson~ W. S. Osborn, and o. A. Knorr.

1968b. Data on mountain environments II. Front Range, Colorado, four climax regions, 19531958. Univ. Colo. Studies, Ser. Biol. 28. 169 pp.

May, T. A. 1975. Physiological ecology of white-tailed ptarmigan in Co1orado. Ph. D. Thesis, University of Colorado, Boulder.
311 pp.

----- , and C. E. Braun.
ptarmigan

1972. Seasonal foods of adult white-tailed
in Colorado.
J. Wildl. Manage. 36(4):1180-1186.

�-72-

Moran, R. J., and W. L. Palmer.
1963. Ruffed grouse introductions and
population trends on Michigan Islands.
J. Wildl. Manage. 27(4):606614.
Nimlos, T. J., and R. C. McConnell.
Sci. 99(5):310-321.
Pearl, R. M. 1964.
Colo. 36 pp.

1965.

America's mountain.

Alpine soils in Montana.

Soil

Alan Swallow Publ. Co., Denver,

Sclater, W. L. 1912. A history of the birds of Colorado.
Co., London.
576 pp.

Witherby

and

Schmidt, R. K., Jr. 1969. Behavior of white-tailed ptarmigan in Colorado.
M. S. Thesis.
Colorado State Univ., Fort Collins.
174 pp.
Stokes, A. W. 1954. Population studies of the ring-necked pheasant on
Pellee Island, Ontario.
Ontario Dept. Lands and Forests.
Wildl. Sere
No.4.
154 pp.
U. S. Department of Commerce, Weather Bureau.
1920-1934.
Climatological
data. Annual Summaries, Colorado.
U. S. Government Printing Office,
Washington, D. C.
Weber, W. A.
Boulder.

1972. Rocky Mountain
438 pp.

flora.

Whitfield, C. J. 1933. The vegetation
Monographs 3(1):76-105.

Wildlife

Researcher

Colorado Assoc. Univ. Press,

of the Pikes Peak region.

Ecol.

�-73-

JOB PROGRESS

Aprii 1978

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Game Bird Survey

W-37-R-3l

Job Title

1
Job No.
21
Minimum Tillage Techniques for
~E~s~t~a~b~l~i~s~h~l~·n~g~S~h~r~u~b~s_=i~n~C~l~u~m~p~P~l~a~n~t~l~·n~g~s~
_

Period' Covered:

April 1, 1977 through March 31, 1978

Work Plan No.

Personnel:

----~---------------------------

Warren D. Snyder

ABSTRACT
The shrub sites received near normal precipitation in 1977. A severe hail
storm in late May 1977 caused extensive injury and defoliation to plums
on the three Karg sites, resulting in reduced growth and subsequent
increased winter die-back and mortality.
Mulch treatments increased shrub
vigor, reduced winter die-back and reduced mortality during the winter of
1976-77.
A slight increase in mortality of plums was evident in 1977.
Wide variability in shrub growth in 1977 resulted from previous die-back
and elongated root sprouts. However, overall height remained greatest
among plastic-mulch treated plots.
Plastic mulch also retained the highest
capacity to prevent weed competition.
Organic mulch lost some weed control
effectiveness due to its decomposition and simazine showed similar results.
The herbicide, casoron, which was applied in late winter each year,
continued to be most effective in weed control. All treatments reduced
annual weed occurrence extensively, as in previous years.

��-75-

MINIMUM TILLAGE TECHNIQUES FOR
ESTABLISHING SHRUBS IN CLUMP PLANTINGS
Warren D. Snyder

P. N. OBJECTIVE
To evaluate ground cover techniques, pre-emergent herbicide techniques, a
combination of the two, and cultivation techniques for establishing shrubs
in clump p1antings in eastern Colorado.
SEGMENT OBJECTIVES
1.

To apply pre-emergent herbicide treatments to the Eptam-Treflan
Casoron plots and to apply cultivation to the control.

2.

To measure and evaluate
tition among treatments

and

shrub survival, shrub growth and weed compeand soils in relation to weather variables.

METHODS AND MATERIALS
Methods and materials used in this study were summarized by Snyder (1975)
and Snyder (1976). Procedures listed there were followed very closely
during this work segment.
Use of rabbit repellent duplicated that of the
previous segment (Snyder 1977).
RESULTS AND DISCUSSION
Environmental

Measurements

Precipitation
The rain gauge located at the Tamarack Headquarters measured 15.68 inches
of precipitation for 1977. Most of this precipitation was general over the
region and distributed to all test sites.
The amount received was considerably above that for 1976 and near the average expected in the region
(Table 1).
A moderately severe hail storm pounded the three Karg sites north of Crook
on May 29, 1977 with considerable detrimental impact.
These shrubs, many
of which had winter killed and were starting new root sprouts, were
severely defoliated and sustained extensive injury to their cambium.
The
upper branches were bruised and debarked which affected both older survivors
and new growth so that high mortality and winter die-back was evident in
spring 1978.

�-76-

Table 1. Precipitation received at the Tamarack Management
quarters from 1974 through 1977. 1/

Area head-

Month

1974

1975

1965

1977

January

0.25

0.05

0.83

0.36

February

0.67

0.15

0.20

0.07

March

0.91

1. 25

0.40

1.50

April

0.45

0.65

1.60

3.50

May

0.70

5.55

1.90

3.35

June

2.05

2.00

1.10

1.40

July

1.15

0.55

1.10

0.95

August

0.40

1.60

1.10

2.65

September

0.15

0.55

1.60

1.20

October

0.00

0.00

0.25

T

November

0.25

1.20

0.15

0.50

December

0.20

0.55

0.00

0.20

Annual

7.18

14.10

10.23

15.68

total

1/Precipitation

for winter months could not be accurately measured so much
of the snowfall information was taken from the weather stations five miles
south of Sedgwick and at Sterling.
August precipitation was a major asset to all shrubs in 1977 permitting
them to enter the fall and winter period with increased vigor, however,
fall-winter 1977-78 conditions remained dry. Moisture was deficient going
into the 1978 growing season.
Rabbit Damage
Wild plum on the Sonnenburg site received considerable pruning and girdling
loss in fall 1976 prior to application of a chemical repellent.
The Karg
sites received minor damage from rabbits and other sites were not affected
to a noticeable degree.
The rabbit damage had considerable impact on 1977
growth on the Sonnenburg site. Earlier application of repellent in fall
1977 and lack of appreciable fall-winter snowfall reduced rabbit impact
going into the 1978 growing season.

�-77-

Insect Damage
Heavy infestations of grasshopper
hills during the summer of 1977.
in reduced growth there; however,

occurred on the 1-76 site in the sandThey were considered an important factor
no control treatments were applied.

Shrub Survival
Shrub Survival Within

Sites Established

in 1974

The mulch treatments, even the thin layer of mulch applied to the simazinemulch plots, brought the wild plums on those sites through the winter of
1976-77 with observably better vigor, less winter die-back, and less winter
mortality.
Root sprouting replaced previous mortalities to increase the
number of shrubs present on organic mulch, plastic mulch and simazine
mulch plots from spring to fall, 1977; whereas, shrub percentages declined
on all other sites (Table 2). Although a statistical evaluation of survival
has not been completed, the information in Tables 2 and 3 indicates the
chemicals have had some impact in reducing wild plum survival over cultivated control treatments.
The effects of chemicals on Hansen rose, if any, are not apparent in Table
2. Cultivation of the control with a garden rototiller destroyed many new
root sprouts, preventing replacement of mortalities and therefore, held the
survival down in relation to the chemical treatments.
Organic and plastic
mulch both sustained increased survival over other treatments.
Considerable
replacement of previous mortalities was noted on the mulched sites in 1977
following the increased rainfall.
The only appreciable summer 1977 mortality
occurred on the chemically treated and cultivated plots of the Karg 1 and
Karg 2 sites where winter die-back had been severe and the root sprouts
were in a weak condition when the hail occurred.
Shrub Survival on Sites Established

in 1975

Almost no mortality has occurred among wild plum on the plastic mulch
treated plots during the last three years (Table 3). All other treatments
show survival rates in fall 1977 from 83 to 90 percent, with cultivated
controls running slightly above the rest. Shrubs on all four chemical
treatments were similar in survival having suffered about 15 percent loss
during the last three years.
Hansen rose survival was again highest on the plastic mulch treatment and
organic mulch ranked second (Table 3). Cultivated and chemical plots
averaged about 80 percent survival and nearly all of this had occurred prior
to the spring of 1977.
When the tabular data from the sites established in 1974 and 1975 are compared (Tables 2 and 3) the one similarity that stands out is the higher
survival on the plastic mulch treatment for both species for both sets of
data. Considerable variation occurred in survival among the other treatments, sites, years, and species.

�-78-

Table 2. Percentage shrub survival from fall 1975 through fall 1977 on
the five sites established in April 1974. II

Treatment

Fall
1975

Spring
1976

Percentage Alive In
Fall
Spring
1976
1977

Fall
1977

Wild Plum
Cultivated

91.88

91.88

90.63

87.50

84.38

Organic mulch

82.03

82.81

82.03

85.16

89.06

Plastic mulch

90.67

90.63

90.63

92.19

94.53

Simazine

84.38

83.75

83.13

78.13

76.88

Simazine-mulch

85.63

87.50

87.50

87.50

90.63

Eptam-treflan

83.13

86.25

85.63

76.88

70.63

Casoron

83.13

84.37

81.88

72.50

70.63

Overall average

85.80

86.74

85.89

82.48

81.82

Hansen Rose
Cultivated

66.25

71.88

71.88

70.63

71.25

Organic mulch

75.00

82.03

81.25

85.94

92.19

Plastic mulch

85.16

86.72

85.94

89.06

92.19

Simazine

74.38

79.38

77 .50

79.38

81.25

Simazine-mulch

76.25

76.25

76.25

78.75

84.38

Eptam-treflan

76.88

81.25

81.25

80.63

85.63

Casoron

73.13

75.63

74.63

70.63

75.63

Overall average

75.00

78.69

78.03

78.79

82.67

II New roots sprouts replaced previous mortalities to slightly increase the
percentage survival on some plots during the spring of 1976 and 1977.

�-79-

Table 3. Percentage shrub survival from fall 1975 through fall 1977 on
the three sites established in April 1975. !/

Treatment

Fall
1975

Spring
1976

Percentage Alive In
Fall
Spring
1976
1977

Fall
1977

Wild Plum
Cultivated

97.92

95.83

93.38

90.63

90.63

Organic mulch

100.00

95.83

93.38

91.67

88.54

Plastic mulch

100.00

98.96

98.96

97.92

97.92

Simazine

96.88

94.79

93.38

86.46

86.46

Simazine mulch

97.92

95.83

93.38

85.42

83.33

Eptarn-treflan

100.00

93.38

92.71

83.33

84.38

Casoron

96.88

86.46

86.46

82.29

84.38

Overall average

98.51

94.49

93.30

88.24

87.95

Hansen Rose
Cultivated

98.96

86.46

85.42

80.21

81.25

Organic mulch

100.00

92.71

91.67

88.54

88.54

Plastic mulch

98.96

98.96

98.96

97.92

97.92

Simazine

98.96

95.42

85.42

82.29

80.21

Simazine mulch

97.92

96.46

86.46

82.29

81.25

Eptarn-treflan

97.92

87.50

84.38

80.21

79.17

Casoron

98.96

81.25

80.21

75.00

78.13

Overall average

98.81

88.39

87.50

83.78

83.78

!/ New root sprouts replaced previous mortalities to slightly increase
the percentage survival on some plots during the 1976 and 1977 growing
seasons.

�-80Shrub Growth
Wild Plum
Table 4 shows considerable variability in shrub growth among sites and
treatments in 1977. Figs. 1 and 2, respectively, illustrate growth and
winter loss variables per treatment for plum and rose. The loam soil
sites (Karg sites 1, 2 and 3) all showed a similar pattern of limited plum
growth on the organic mulch and plastic mulch treatments in comparison to
considerable growth on the cultivated and chemically treated plots (Table
4). Observations indicated this greater growth was primarily comprised of
elongated root sprouts following severe drought dessication and die-back
during the previous winter (Fig. 3). Since the mulch plots did not winter
kill and die back as severely, their growth was added to existing branches
and was understandably less.
This comparison is best illustrated in Figs.
1 and 3 for plum, and Fig. 2 for Hansen rose. The shorter growth on the
organic and plastic mulch plots increased 1977 height over that of 1976,
whereas, the greater growth on the cultivated and chemically treated plots
only approximately compensated for the previous winter's loss. The hail,
that was previously discussed, also was a primary factor reducing growth
and survival on the larger shrubs present in the mulched plots of the Karg
sites.
The pattern of growth among plum shrubs on the other clay-loam and sandyloam sites was not clear cut. No analysis of variance test were attempted
on 1977 growth because of the above mentioned variables.
Hansen Rose
Table 4 shows the comparative growth characteristics of rose shrubs in
1977, which like plum, did not show a clear cut pattern among treatments.
Analysis of variance was not attempted on the 1977 data. Fig. 2 illustrates
the' comparative growth loss variables from 1975 through 1977. Casoron
treated plots made the greatest growth in 1977 placing them a close second
to plastic mulched shrubs in total fall 1977 height.
Some form of blight
or other disease caused considerable loss of large portions of most roses
on nearly all sites during the summer of 1977.
Wild plums established in 1975, when contrasted to those established in
1974, attained greater 1977 growth (Table 5). Hansen rose did not show

this growth difference among samples.
Site differences have become more noticeable with progression of the study.
The plums on the Red Lion site have consistently made much better growth
than those on the more sandy Sharp tail site and are presently in a near
closed canopy status.
The Tamarack Meadow plums on the cultivated, organicmulch, and plastic-mulch treatments have achieved heights up to seven feet
or more and have formed a dense thicket of cover. Chemically treated
plums there are shorter and more open, but still of considerable cover
value.
In contrast, plums on the loam soil sites, which achieved excellent
growth during their first two years, became over competitive for moisture
during severe drought conditions in 1976 and as a consequence, are in poor
shape and appearance at present.

�Table 4. Average shrub growth in inches per site and treatment during the 1977 growing season on the five
sites established in 1974 and the three sites established in 1975. Jj
Treatment
Soil Type
Site

Cultivated

Organic
Mulch

Plastic
Mulch

Simazine

Simazine
Mulch

EptamTref1an

Casoron

Site
Mean

Wild Plum
Loam Soil
Karg 1
Karg 2
Karg 3

11.21
10.54
13.50

7.76
9.00
2.78

4.38
7.03
3.48

8.81
12.29
15.52

6.31
10.19
15.39

7.71
10.76
15.62

14.95
10.33
14.50

8.39
9.77
11.26

Clay Loam
Tamarack Meadow
Sonnenburg

11.77
18.80

7.76
13.09

10.68
12.28

7.75
11.15

5.19
18.90

4.82
15.60

4.04
8.58

7.57
14.17

Sandy Loam
Red Lion
Sharp tail
1-76

6.81
1.88
4.72

15.28
6.33
7.57

9.66
4.72
6.32

8.77
4.69
11.00

8.29
3.39
10.07

6.35
2.31
9.38

8.04
5.86
13.96

9.06
4.12
8.89

Treatment Mean

10.15

8.61

7.32

10.03

9.64

9.06

9.99

9.20

Hansen Rose
Loam Soil
Karg 1
Karg 2
Karg 3

11.15
10.15
11.00

9.10
9.28
9.41

8.75
15.35
7.66

12.67
14.59
9.94

10.26
11.97
11.69

7.22
11.79
10.91

10.59
16.78
10.55

9.98
13.81
10.14

Clay Loam
Tamarack Meadow
Sonnenburg

10.93
7.19

10.10
5.47

1.68
8.91

6.03
7.69

6.71
6.57

4.48
6.30

4.38
7.13

6.38
7.05

Sandy Loam
Red Lion
Sharp tail
1-76

9.08
5.80
4.52

11.94
6.67
8.00

11.84
8.22
6.40

16.41
1.96
8.75

12.46
7.44
9.00

14.78
7.19
11.82

l3.04
5.08
12.69

12.72
l3.00
8.33

Treatment Mean

9.04

8.84

8.70

9.60

9.71

8.88

10.17

9.25

-1./ The Karg 3, Sonnenburg, and 1-76 sites were established in 1975.

I
00
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I

�-82-

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20
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Fig. 1. Comparative wild plum height,
characteristics per treatment averaged
1975 through fall 1977.

II

1

growth, and winter die-back
for the eight sites from spring

�-83-

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20

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14
12
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F~g, 2, Cpmparqtive Hqnsen rose height, growth, and winter die-back
characteristics per treatment averaged for the eight sites from spring
1975 through fall 1977.

L

�-84-

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28

~
o-

1M
\

~

r

~

J

26

g) 24

8

22

~

I

~

a

20

~

18

;
CI)

16
14
12
10
8

0-'=

a
~

i:f

Eo4

6

~~
~

i~

Fig. 3. Comparative wild plum height,
characteristics per treatment averaged
from spring 1975 through fall 1977.

growth, and winter die-back
for the Karg 1 and Karg 2 sites

�-85Table 5. Comparisons of shrub growth in inches during
of 1975 through 1977 on the shrub clump sites.

Mean Growth - vli1d Plum
1975
1976
1977

Soil Type
Site

the growing

seasons

Mean Growth - Hansen Rose
1975
1976
1977

Loam Soil
Karg 1

19.73

6.35

8.39

14.16

2.81

9.98

Karg 2

19.38

7.62

9.77

13.71

4.60

13.81

Karg 31)

6.89

14.85

11.26

8.87

4.93

10.14

21.58

14.14

7.57

14.06

6.89

6.38

3.48

17 .60

14.17

2.35

4.99

7.05

Red Lion

13.84

9.01

7.63

13.27

4.68

13.25

Sharp tail

6.79

7.63

4.12

5.63

4.52

13.00

1-761=.1

5.04

11.53

8.89

3.18

7.57

8.33

to those established

in April

Clay Loam
Tamarack
Meadow
Sonnenburg-

1/

Sandy Loam

!/Sites
1974.

established

in April 1975 in contrast

The Hansen rose, when contrasted to the plums, do not appear capable of
attaining comparable heights at present in these dense thicket plantings.
The rose, however, shows a greater tendency to root sprout and produce more
basal branches.
Weed Competition
The plastic mulch treatment retained its capability as the most effective
annual weed control treatment for the third consecutive year (Table 6), in
addition to its effectiveness in increasing shrub survival and growth.
As
in previous years, all treatments (chemical and mulch) greatly reduced
annual weeds in comparison with densities on the cultivated control.
Annual
weed densities were higher on all treatments in 1977 because of increased
spring moisture (Table 6). In addition, there was some deterioration in
the effectiveness of the organic mulch treatments resulting from compaction
of mulch and rodent disturbance.
Casoron, as in 1976, continued to be
highly effective in weed control, although the rate of application was
reduced in 1977. Simazine and Eptam-treflan treatments showed weed control
effectiveness comparable with that of previous years.

�-86-

Table 6. A comparison of average annual weeds per square foot sample
within treatments on the eight sites from 1975 through 1977.

1975

1976

1977

29.29

15.04

64.54

Organic mulch

0.87

0.43

6.57

Plastic mulch

0.38

0.22

1.04

Simazine

2.10

2.66

7.59

Simazine mulch

5.45

1.86

9.88

Eptam-treflan

1.15

1.62

6.13

Casoron

5.56

0.12

1.93

Treatment

Cultivated

control

Table 7 compares the average number of annual weeds per square foot
sample within treatments per site for May 1977. The Sonnenburg site,
within clay-loam soil, contained very high weed densities in the cultivated
control and higher than average densities in all other treatments.
It
was primarily responsible for the dramatic weed density increase in the
organic mulch treatment.
Simazine was applied at a reduced rate in late winter 1977 to treatments
within the five original sites. Weed control there was noticeably more
effective than on the three sites established in 1975 where it was not
re-applied, confirming its deterioration in effectiveness by the second
year after application.
Statistical analyses were not conducted on this set of data because of
similar trends shown with previous years.
However, a complete analysis
all years data will be forthcoming at study termination.

of

Annual weeds were removed in mid-May.
Removal operations then and in June
were not 100 percent effective so no attempt was made at further evaluation.
However, there was no major re-sprouting of weeds after the initial removal
operations and all plots remained near completely weed free from early
July through the remainder of the summer, even though increased precipitation was received.

�Table 7. Average occurrence of annual weeds per square foot sample in May 1977 within eight sites in
three soil types. 1/

Cultivated

Organic
Mulch

Plastic
Mulch

Karg 1

30.39

7.31

0.39

Karg 2

30.39

15.58

Karg 3

28.22

Soil Type mean

Soil Type
Site

Treatment
Simazine
Mulch
Simazine

EptamTref1an

Casoron

Site Mean

8.83
0.44

8.32
9.23

Loam
2.19

0.39
1.36

3.50

8.06
11.17

0.72

0.86

22.03

23.33

2.75

0.36

11.18

29.67

7.87

1.15

7.93

9.90

7.32

3.12

9.58

13.92

0.00

0.11

0.00

0.60

4.50

0.58

2.74

4.33

52.08

2.86

Clay Loam
Tamarack Meadow

I

Sonnenburg

284.89

22.44

3.22

16.56

26.94

6.19

Soil type mean

149.40

11.22

1.67

8.28

13.50

5.35

2.46

27.41

Red Lion

30.44

1.39

0.11

1.56

0.86

7.22

0.83

6.06

Sharptail

50.78

0.97

0.89

8.75

0.61

3.08

0.03

9.30

I-76

47.28

4.17

0.56

10.11

20.83

6.06

0.31

12.76

Soil type mean

42.83

2.18

0.52

6.81

7.44

5.45

0.39

9.37

Overall Treatment Mean

64.54

6.57

1.04

7.59

9.88

6.13

1.93

13.96

I

Sandy Loam

-

1/ Averages include data from the five sites established in 1974 and the Karg 3, Sonnenburg, and I-76
sites established in 1975.

co
-....J

�-88-

LITERATURE CITED

Snyder, W. D. 1975. Minimum tillage techniques for establishing shrubs
in clump p1antings. Colorado Div. of Wi1d1. Game Res. Rept. April:
141-153.
1976. Minimum tillage techniques for establishing shrubs in
clump p1antings. Colo. Div. of Wi1d1. Game Res. Rept. April: 209234.
1977. Minimum tillage techniques for establishing shrubs in
clump p1antings. Colo. Div. of Wi1d1. Game Res. Rept. Apri1:163176.

Prepared by.

~~..
f/...I&lt;:/~/-=·~=:,'~~).:::.~=:-4--=J~I,:::.~?-=W=---:...:'t~:..::..::":.....)J_"
Warren D. Snyder
Wildlife Researcher C

�April 1978
-89-

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Job Title

1
---------------------------------

Upland Game Publications

Covered:

Personnel:

Job No.

22

Work Plan No.

Period

Game Bird Survey

W-37-R-3l

April 1, 1977 through March 31, 1978

T.D.I. Beck, C. E. Braun, T. Britt, A. G. Haskins, R. W. Hoffman,
E. O. Hohn, O. W. Olsen, W. D. Snyder, R. M. Stabler, and R. O.
'\.Jalles
tad.

ABSTRACT
Pulbications

planned for and accomplished

under this job for Segment

31 are

as follows:
Braun, C. E. 1978. Weight dynamics
To be published in 1978.

of sage grouse.

Accepted

by Condor.

Braun, C. E., and T.D.I. Beck.
1977. Population dynamics of sage grouse,
North Park. Colorado.
Proc. Ann. Conf. Central Mtns. and Plains Section,
The Wildlife Society.
22:Abstract.
Braun, C. E., T. Britt, and R. O. Wa11estad.
1977. Guidelines for maintenance of ~age grouse habitats.
Wi1d1. Soc. Bull. 5:99-106.
Hoffman, R. W. 1977.
. 26(5) :30-32.

Blue grouse - an overlooked

resource.

Colo. Outdoors

Hohn, E. 0., and C. E. Braun.
1977. Seasonal thyroid gland histophysiology
and weights in white-tailed ptarmigan.
Auk 94:544-551.
Olsen, O. W., A. G. Haskins, and C. E. Braun.
1978. Rhabdometra alpinensis
n. sp. (Cestoda:Paruterinidae:Dilepididea)
from southern white-tailed
ptarmigan (Lagopus leucurus altipetens Osgood) in Colorado, U.S.A.,
with key to the species of Rhabdometra Cholodkowsky, 1906. Canad. J.
Zool. 56:446-450.
Snyder, W. D. 1978. The bobwhite
Tech. Publ. No. 32. 88 pp.

in eastern Colorado.

Stabler, R. M., C. E. Braun, and T.D.I. Beck. 1977.
from Colorado.
J. Wildl. Diseases 13:414-417.

Howard D. Funk
Section Chief, Small Game Research

Colo. Div. of Wildl.

Hematozoa

in sage grouse

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                  <text>-1-

JOB PROGRESS

State of
Project

1978

REPORT

..::C.:::.O=.LO.:::.RAD:.::.::::..O=_
No.

W-125-R-4

Work Plan No.
Job Title
Period

July,

Covered:

Personnel:
R. Kincaid,

Deer-Vehicle

Accident

Investigations

1
Job No. 1, 2, 3, 4, 5, and 6
Deer Vehicle Accidents Statewide
and Methods and Devices to Reduce Them

April 1, 1977 - June 30, 1978

Thomas D. I. Beck, James D. Fleming, Sharon L. McDonnell, Kenneth
Dale F. Reed, Area Supervisors and Wildlife Conservation Officers.

ABSTRACT

Methods, devices, or structures related to reducing the number of deer-vehicle
accidents were evaluated or experimentally tested.
These were highway
lighting, underpasses, overpasses, 2.44-m fences and one-way deer gates, and
deer guards.
Nine deer were killed in the highway lighting study area, five
and four with lights off and on, respectively.
Background and target luminance
measurements were taken at 10 additional kill sites. The overall visibility
index mean was 1.366 ± 0.732 (SD). No major changes occurred with underpass
use. The experimental procedure for manipulating the Dowd deer overpass
width was continued for three seasons.
A combined total of 123 and 65 crossings
occurring during control (4.93 m) and variable (2.48 m) widths, respectively,
were reviewed during video tape replay.
Although selected behavioral differences
were noted during these seasons, the combined data for four seasons has not
shown significant differences at the 95 percent confidence level. The average
reduction of deer-vehicle accidents adjacent to six 2.44-m fences was 76.4
percent.
Limited data were obtained on deer guard prototype V.

��-3-

DEER-VEHICLE ACCIDENTS STATEWIDE
AND METHODS AND DEVICES TO REDUCE THEM
Dale F. Reed and Thomas D. I. Beck

P. N. OBJECTIVE
Locate and examine potentially critical deer-vehicle accident areas in
Colorado and recommend methods or structures which may reduce deer-vehicle
accidents in these areas.
Subsequently, measure the effects of methods
recommended and investigate deer responses to various experimental
structures.

SEGMENT OBJECTIVES
Job 1 - Monitoring
statewide.

potentially

critical

deer-vehicle

areas

1.

Locate and examine potentially
accident areas in Colorado.

2.

Recommend methods or devices which may reduce the problem
for these critical accident areas.

3.

Select recommended underpasses, overpasses, 2.44-m fence
lengths, or other devices or methods for evaluation when
sample size considerations can be met.

Job 2 - Evaluating the effects of highway
killed by vehicles.

critical

accident

deer-vehicle

lighting on number of deer

1.

Measure and describe the luminance
segment of Colorado Highway 82.

on an illuminated

l.l-km

2.

Determine if deer-vehicle accidents are affected by the
fixed illumination on the l.l-km segment of Colorado Highway

3.

Investigate and compare responses of motorists to deer on
the highway, with and without fixed illumination.

4.

Investigate
and without

Job 3 - Evaluate
1.

responses

to motorists,

with

of deer to highway underpasses.

Measure deer use of underpasses and the extent of behavioral
reluctance associated with their use.

Job 4 - Evaluation
1.

and compare deer responses
fixed illumination.

of deer responses

to highway

Determine the use of overpasses
deer kill areas.

overpasses.

by deer in critical

highway

82.

�-4-

2.

Compare detailed deer behavioral responses to overpasses
after moderate to high use of the structures or immediate
areas has been documented.

Job 5 - Evaluate deer responses
to highways.

to 2.44-m

(8-ft) fencing adjacent

1.

Determine the reductions in deer-vehicle accidents on highways after the installation of various lengths of 2.44-m fence.

2.

Measure

Job 6 - Evaluating
1.

deer movements
responses

in relation

to 2.44-m fences.

of deer to guards.

Compare deer responses to a black-and-white
(V) and roll-bar (VI) deer guard prototypes
conditions.

DESCRIPTION
Job 1 - Monitoring
wide.

pontentially

scintillation
under controlled

OF AREAS

critical

deer-vehicle

accident

areas state-

Generally, Myers (1969), Pojar (1972a), and Woodard (1973a, 1973b,
1974, 1975, 1976a) described the areas most intensively monitored.
The completion of deer-vehicle accident reports (includes location)
has been required statewide since 1974. Hence, the potential
existed for the detection of new problem areas statewide.
Job 2 - Evaluating the effects of highway
killed by vehicles.

Highway

82 - Jammeron

Pojar et al. (1975) described
Job 3 - Evaluate

responses

lighting

on number of deer

Curve

this area.

of deer to highway

underpasses.

The Vail deer underpass (Reed et al. 1975), a small structure
under 1-70 east of Avon, two concrete box underpasses under 1-70
east of Eagle, twin bridge structures under 1-70 west of Eagle,
and twin bridge structures east of Rifle were monitored for deer
use or behavioral responses.
Job 4 - Evaluation

of deer responses

to highway

Dowd Junction
Reed

(1976a) described

1-70

this area.

overpasses.

�-5-

Job 5 - Evaluate deer responses
highways.

Highway
The Highway

to 2.44-m

(8-ft) fencing adjacent

82 - Diamond

S

82 study area has been described

Highway

by Pojar

(1972b).

82 - Carbondale

This study area was described

Interstate

to

by Reed (1975).

70 - Avon, Edwards,

Vail, and Eagle

The Interstate 70 Avon and Edwards study areas have been described
by Woodard (1973b), the Vail study area by Reed (1971), and the
Eagle study area by Reed (1974).
Job 6 - Evaluating

respones

of deer to guards.

Trail Gulch-Dotsero
Reed et al. (1974) described

this area.

METHODS AND MATERIALS
Job 1 - Monitoring
statewide.

potentially

critical

deer-vehicle

accident

areas

Methods have been described by Pojar (1972a) and Woodard (1976a).
Because no new critical areas were delineated, methodology was
limited to tabulation of statewide Wildlife Conservation Officer
kill reports.
Job 2 - Evaluating the effects of highway
killed by motorists.

lighting

on number of deer

Methods were described by Woodard (1976b). Luminance measurements
were taken at kill sites from earlier years.
Luminance values
(foot-lamberts) were recorded with a spotmeter, Spectra Model UBA,
and were taken on a target at the kill site and on the background
as viewed by an approaching motorist.
The target was a taxidermy
mount of the left half (transverse section) of a female mule deer
and the actual spotmeter recording area was a 26-cm diameter circle
midway between the shoulder and hip. Spotmeter readings were ·taken
from a height of 1.3 m and a distance of 15 m. For background

�-6-

luminance readings the spotmeter was placed 60 m from the kill
site, resulting in a target area diameter of 102 cm. The
taxidermy mount was removed during this background measurement.
Measurements were taken between 0400 and 0600 to minimize traffic
interference.
Background luminance (Lb) and target luminance (Lt)
measurements were transformed into a visibility index (VI) by the
following equation (Gallagher and Meguire 1974):
C(RCS Lb)
VI

(DGF)

=

where C
5.74

and where DGF

= Disability

Lb-Lt
Lb

glare factor

Relative contrast
sensitivity for
the recorded background luminance

= 1.0.

RCS values were obtained from standard tables (Gallagher and Meguire
1974). VI values can then be compared to standards established for
urban lighting requirements.
These values also provide a better
description of the visibility of deer in the lighted portion of the
study area; which, in light of the poor horizontal uniformity ratio
in the lighted section (&gt;15.0:1), should allow a more refined
analysis of the effectiveness of highway lighting.
Job 3 - Evaluate

responses

of deer to highway

underpasses.

Seven underpasses were monitored for deer use as described by
Reed (1971). Observations at the Vail deer underpass were
continued as described by Reed (1976b).
Job 4 - Evaluation

of deer responses

to highway

overpasses.

Methods were previously described by Reed (1976a) and Reed and
Beck (1977). The width (control = 4.93 m, variable = 2.48 m)
of the Dowd deer overpass was changed every three days during most
of the spring migration of 1977 and 1978, and the fall migration
of 1977. In addition during the fall migration of 1977 and the
spring migration of 1978, the video housing was moved to cover the
distant approach areas (approximately 20 m beyond the bridge entrance
and exit).
Job 5 - Evaluate deer responses
highways.

to 2.44-m

(8-ft) fencing adjacent

to

Methods and materials have been described by Pojar (1972b) and
Woodard (1976c).
Benefit-cost analysis followed procedures
described by Howe (1971). Costs for which values were available
include initial cost of 2.44-m fence and associated passage structures.
Estimates of maintenance costs were not available.
Total cost was
reduced by the cost of the appropriate length of 1.07-m (42-inch)
standard right-of-way fence which was not built because of the 2.44-m
fence.
Benefits for which a value was available was the potential
savings accrued by a reduction in deer-vehicle accidents and related

�-7-

vehicle repairs.
Savings by reduction in personal injury and death
were omitted because of difficulties in obtaining reliable values.
No value was assigned to deer. Therefore, benefit-cost analysis
is essentially from a highway department accounting stance. All
future benefits were discounted at an annual rate of 5.5 percent.
No monetary values have been assigned for aesthetic considerations,
either positive or negative.
Job 6 - Evaluating

responses

of deer to guards.

Methods were described by Reed et ale (1974).

RESULTS AND DISCUSSION
Job 1 - Monitoring
statewide.

potentially

critical deer-vehicle

accident areas

Two Division of Wildlife Regions, Southwest and Northeast, have
taken over the responsibilities of maintaining their deer-vehicle
accident files. Area analyses of deer-vehicle kills will not be
generated unless requested because of manpower requirements.
Generally no new critical deer-vehicle accident areas were detected.
A total of 2,174 deer-vehicle accident kills were reported in
Colorado in 1977; an increase of 16.9 percent over 1976. Regional
totals were Southwest 887, Northwest 625, Northeast 416, and Southeast 246. In early 1978 some potentially critical areas had a
higher than average number of kills, possibly because of above
average snow accumulation.
Job 2 - Evaluating the effects of highway
killed by vehicles.

lighting .on number of deer

Deer occupied the highway lighting study area for eight weeks beginning
4 February.
Four weeks of lights off and four weeks of lights on
data (lights off and on for alternate weekly periods) were collected
when deer were in the study area. Although a high number of deer_
were in the general study area (highest count and second highest X
count in the 11 year period) only a moderate number of deer we re
killed by vehicles.
Snowfall adjacent to the study area at higher
elevations was considerably greater than the previous two years and
snow accumulation at three snow stakes above the study area was
comparable to 1975. Minimum temperatures dropped below OOF on only
two days (Feb. 12 and 13) and maximum temperatures exceeded 320F on
all but five days during the January-March period.
The warm temperatures, coupled with rain, in March, opened up considerable range,
as evidenced by low numbers of deer counted in the study area during
the last week of March, so the duration of stay on the study area
was not extended.
For the last two winters we were short on snow;
then this winter long on snow, short on cold weather.
The vagaries
of weather make the study of wild populations a high-risk venture.

�-8-

Nine deer were killed during an estimated total of 436 crossings
(Table 1). The deer crossings to deer kill ratio with lights
off was less than the ratio with the lights on 36.4 and 63.5,
respectively.
However, by chi-square analysis, there was no
significant difference (P &gt; 0.25) between these ratios.
In
addition, the ratios are less than the combined ratio data
obtained during 1974-75 (i.e. 87.8 and 96.3, off and on,
respectively).
The differences, however, are not significant
(P &gt; 0.05 and P &gt; 0.25 off and on, respectively) according to
chi-square analyses.
It is reasonable to postulate that crossings
per kill ratios may change over extended periods of time because
of the annual increases in traffic volume.

Table 1. Estimated deer crossings
lighting study area during 1978.

and total kill in the Highway

82

Off

On

Total

182

254

436

Total kill

5

4

9

Crossing/kill

36.4

63.5

Estimated

crossings

Background and target luminance measurements were taken at 10
additional sites. The visibility index mean (1.366 ± 0.732 [SD])
value is lower than minimum levels needed for a roadway with an
80.5 km (50 mile) per hour speed limit. It approaches the level
where 60-65 percent of the motorists can see a target at satisfactory separation distance (Gallagher and Meguire 1974). To
attain a level where 85-95 percent of the motorists can see a
target at satisfactory separation distance, a visibility index
value of 2.6 - 3.6 is required (Gallagher and Meguire 1974:51).
The 2.484 value (Table 2) approaches the 85 percent level.
Job 3 - Evaluate

responses

of deer to highway

underpasses.

Seven underpasses were regularly checked for deer use during this
segment.
Moderate (30-90) to high (&gt;90) numbers of deer passages
occurred through two of the underpasses, one located west of Vail
and one east of Avon. Extremes of openness or tunnel effect were
represented by these two structures.
The primary stimulus of a
given underpass structure to approaching deer may be termed the
"openness effect."
Calculated as follows:
height x width

(or open-end
length

surface area)

�-9-

the openness effects of the Vail and Avon underpasses were 0.3
and 0.03 (metric measurements), respectively.
There are several
factors to consider before relating openness effect to deer
behavioral response.
In addition, any reasonable attempt to
relate openness effect to deer use must consider deer density
and motivation at the structure.

Table 2. Visibility index measurements
lighting study area.

from kill sites, Highway

82

Contrast

Relative
Contrast
Sensitivity
(Percent)

Visibility
Index

0.0325

0.244

4.10

0.174

0.330

0.0032

0.990

14.40

2.484

12

0.061

0.0094

0.846

5.25

0.774

13

0.235

0.0255

0.891

12.05

1.870

14

0.060

0.0048

0.920

5.20

0.833

15

0.285

0.0040

0.986

13.325

2.289

16

0.080

0.0165

0.794

6.20

0.858

17

0.240

0.4700

-0.958

12.20

-2.036

18

0.086

0.1300

-0.512

6.50

-0.580

19

0.180

0.01l0

0.939

10.40

1.701

20

0.197

0.0690

0.650

10.95

1.240

21

0.180

0.0260

0.856

10.40

1.551

No.

Background
Luminance
(Lb)

Target
Luminance
(Lt)

9

0.043

11

Daily observations (n = 33) were made at the Vail deer underpass
in June of 1977 and 1978. With 1974 through 1978 combined, 78.5
percent of 303 groups of deer did not exhibit flight reaction
when westbound vehicles traveled by the area (Table 3). Likewise,
of 990 instances of vehicles traveling west when deer were near
the underpass
entrance and the west bound lanes, 897 did not

�-10-

elicit flight reaction from 303 different individuals or
groups (1-14 deer).
Twenty-one vehicles elicited flight
reaction from 1-50 percent of the individuals from fifteen
different groups ranging in size from one to ten deer.
Seventy-two vehicles elicited flight reaction from 51-100
percent of the individuals from fifty different groups
ranging in size from one to eight. Previous to these
observations it was hypothesized that deer at distances of
15 to 30 m from moving traffic were frequently caused to
leave the area. Observations indicate that when vehicles
stop at or when bicyclists travel by the underpass, most deer
elicit immediate and intense flight reaction.
Five groups
of deer exhibited moderate to intense flight reaction to
unknown stimuli (where observer was unable to determine
potential cause).
During the 1977 and 1978 seasons moderate
flight reaction (trotting from stimulus) occurred more
frequently (n = 42) than intense flight reaction (bounding
from stimulus) (n = 29).
Table 3. The number and percent of deer groups not exhibiting flight
reaction (Hediger 1950, Scott 1958), and the number and percent of westbound vehicles not eliciting flight reaction at the Vail deer underpass
west of Vail.

Westbound

Groups of Deer

Year

Numbers of Individuals
or Groups 1/ Near
Westbound Traffic

Vehicles

Percent Not
Exhibiting
Flight
Reaction

Number Traveling
West When Deer
Present

Percent Not
Eliciting
Flight
Reaction

1974

37

86.5

198

97.5

1975

67

71.6

251

89.2

1976

78

78.2

1977

59

86.4

14s1/

92.7

1978

62

74.2

ISO'!)

83.3

Totals/Avg.

303

78.5

990

90.6

1/ Number

24-}:./

of deer in groups ranged from 2 to 14.

l/ Six semi-trucks

elicited

flight reaction,

seven did not.

3/ One semi-truck

elicited

flight reaction,

eight did not.

elicited

flight reaction,

three did not.

4/

Ten semi-trucks

89.7

�-11-

The behavior modes of 279 deer exists (leaving the underpass to
the south) were 49.1, 33.7, and 17.2 percent, trotting, walking,
and bounding, respectively.
Combined data 1974 through 1978
yielded a walk:trot:bound
ratio of 2.0:2.9:1.0 (Table 4).
Wariness had been noted irrespective of the mode of exit in many
instances.
The predominant exit behavior of trotting supports
the continued reluctance of deer to use a structure of this
size and character (Reed et al. 1975).

Table 4.
underpass

The number of exits observed at the south end of the Vail deer
and the walk:trot:bound
ratio of exit behavior.

Year

No. of Exits
Observed

1974

35

4.0:12.5:1.0

1975

73

3.3: 2.4:1.0

1976

55

1.8: 6.3:1.0

1977

46

4.0: 2.7:1.0

1978

70

0.7: 1.4:1.0

Totals/Avg.

279

2.0: 2.9:1.0

Job 4 - Evaluation

of deer responses

to highway

Walk:Trot:Bound
Ratio During Exits

overpasses.

The video time-lapse surveillance system was operated at the
Dowd experimental-variable-width
overpass during three seasons,
spring and fall of 1977 and spring of 1978, for 24, 36, and 24
consecutive nights, respectively, except for a six night period
(3 nights control and 3 nights variable) in 1978 when the operation
of the video system deteriorated and equipment was repaired or
replaced.
When data from the spring of 1976 (last segment) are
included, a total of 188 crossings has been examined during video
replay (Table 5).
Number

of Crossings

More crossings occurred under control than variable width during
each of the four seasons (Table 5). A test for independence
shows that the difference between the number of control and
variable crossings during the alternate three-day periods is
nearly significant (X2 = 7.964, df = 4, P &gt; 0.05). However,

�Table 5. The number of deer crossings and approaches examined on video replay at the experimentalvariable-width
overpas~ and the crossings per approach (C:a) ratios for four migration seasons (Fall
1976 - Spring 1978).

Crossings

Control
Distant 1/
2/
Approac~
Approac~

C:a

Crossings

Variable
Distant
Approach

Approach

C:a

Total
Crossings

1976
Fall

25

5

5:1

19

5

3.8:1

44

41

4

10.2:1

19

4

4.8:1

60

1977
Spring

1977
Fall

31

2

o

15.5:1

13

4

7

1.2:1

44

26

o

4

6.5:1

14

7

8

0.9:1

40

123

2

13

8.2:1

65

11

24

1. 9: 1

188

1978
Spring
Totals/Average

1/ Denotes

a deer that entered the overpass area not encompassed by an entrance and exit trackbed and that
was oriented toward the structure.
The overpass area was that area covered by the video surveillance system
during the fall of 1977 and the spring of 1978.

~/ Denotes

a deer that entered

either

the entrance

or exit trackbed

area without

crossing

the overpass.

I
•....
N
I

�-l3-

differences, as determined from a similar analysis (X2 = 19.599,
df = 4, P &lt; 0.005), in pre-experimental data (three-day periods
during four seasons:
fall 74-75 and spring 75-76) are sufficiently
great that this statistical method is inapplicable.
C:a Ratios
Fewer approaches of both types (Table 5) occurred during control
(15) than during variable (35). The control crossings per
approach ratios are significantly (P &lt; 0.025) greater than that
for variable.
Generally, this would be expected if there were a
greater reluctance of deer to cross a narrower structure (i.e.
more reluctant animals either approached and failed to cross, or
made more than one approach before crossing).
The significance
of the ratio difference, however, is diminished if, as was discussed
previously, the difference in the number of crossings is considered
inapplicable.
Duration

of Hesitations

and Crossings

The means of the combined seasons (Table 6) of both the duration
of hesitations and the duration of crossings are not significantly
different (P &gt; 0.20 and P &gt; 0.10, respectively).
Generally, if
the narrower width was expected to result in more reluctance to
cross, then longer hesitations would be expected.
The means, except
for the spring of 1977, support this trend. Reasons for the
exception in the spring of 1977 are unknown.
Likewise, all of the
seasons except one (fall of 1976) have longer mean crossing times,
where animals being more reluctant in crossing the narrower width
(variable) would be expected to do so more hurriedly (more and/or
faster trotting and bounding).
BR:C Ratios
There were more animals exhibiting behavioral responses (BR) and
more instances of behavioral responses per crossings (C) during
variable than control for each of the responses studied (Table 7).
Although the ratios for instances of behavioral response per
crossing are larger for variable width in each case, they are not
significantly larger (P &gt; 0.20). Therefore, differences in these
behavioral responses may be due to chance and not to the reluctance
of animals in crossing a narrow bridge width.
Conclusions
Based upon the above analyses, it does not appear that an important
level of reluctance of deer to bridge width has been reached.
The
statistical analyses used in this job will be reviewed and possibly
expanded during the next segment.
Further experimentation on the
parameter of width is not considered practical since physical constraints do not allow a narrower «2.48m) width to be readily
constructed and subsequently changed to a "control".

�Table 6. The mean duration (seconds) of hesitations and crossings during control and variable
widths at the experimental-variable-width overpass.

sr=:
1976F-

1977S

1977F

1978S

TOTAL

Control

3/
4.5±1.5(n=10)

13. 2±14. 2 (n=29)

8.3±7.7(n=34)

9.2±6.9(n=30)

9.6±9.8(n=103)

Variable

7.4±4.8(n=14)

4.9±3.5(n=6)

11.7±11.1(n=18)

l5.7±14.7(n=26)

11.8±11.8(n=64)

7. 4±4. 9 (n=25)

12. l± 15.7 (n=4l)

8.4±5.3(n=27)

8.9±5.8(n=26)

9.6±10.3(n=119)

6. 8±5. 7 (n=14)

7.5±6.3(n=63)

--

2/

p-

Hesitations
p

&gt;

0.20

p

&gt;

0.10

Crossings
Control
Variable

8.3±3.8(n=19)

8. 8±8. 9 (n=19)

4. 8±4. 9 (n=ll)

I
I-'

~
I

1/

- FaIlor

spring (F or S) migration periods.

2/

- Independent t statistic.

3/

- Standard deviation.

�-15Table 7. The number of selected behavioral responses exhibited by deer
crossing an experimenta1-variable-width
overpass during four migration seasons
Ciall 1976, spring and fall 1977, and spring 1978) and calculated behavioral
response ratios.
No. of Animals
Exhibiting
BR (BRa)

Behavioral
Responses (BR)

Muzzle-to·-Ground

No. of
No. of
Instances
Crossings
(C)
of BR (BR )
i

:cl/

BR

BR. :BR
1
a

a

2/
32

55

123

0.26:1

o .45 :1 1.72 :1

18

33

65

0.28:1

0.51:1

1.74:1

Control

20

24

123

0.16:1

0.20:1

1.20:1

Variable

15

22

65

0.23:1

0.34:1

1.47:1

Control

60

103

123

0.49:1

o . 84 :1 1.72 :1

Variable

40

95

65

0.62:1

1.46:12.38:1

Control

62

103

123

0.50:1

0.84:11.66:1

Variable

42

66

65

0.65:1

1.02:1

1.5"1:1

Control

33

63

123

0.27:1

0.51:1

1.91:1

Variable

34

57

65

0.52:1

0.88:1

1.68:1

during

the four seasons.

Control

1/

Variable

Muzzle-to-Structure

if

Low-Head ~f

6/
-

.
.
HeSltatlon

Alert Stance

llc denotes

7/

the number

~!Muzzle-to~ground
l!Control

of crossings

denotes

and variable

a deer lowering

widths were

its muzzle

to the ground.

4.93 m and 2.48 m, respectively.

~/Muzzle-to-structure
denotes a deer lowering its muzzle
or raising its muzzle to the bridge railing.

to the bridge

deck

~/Low-head denotes a lowering of the head where
(posterior to anterior) below the horizontal.

the axis of the neck declines

~!Cessation

or more.

of forward movement

for 1.0 second

l/Alert stance denotes a position where the head and neck are above horizontal
and the ears are erect with the vertical axis of the ear ei.ther perpendi.cular
to horizontal or inclined forward.

�-16-

Job 5 - Evaluate deer responses
highways.

to 2.44-m

(8-ft) fencing adjacent

to

Six 2.44-m (8-ft) fences were evaluated as to the reduction
of highway deer kills after installation.
The range of
reduction was 61.9 to 88.9 percent with a cumulative average
of 76.4 percent (Table 8). Since this evaluation is based
upon year-to-year comparisons the cumulative average changes
each segment.
The reduction in kills adjacent to the Avon,
Edwards, Eagle, and Diamond S fences increased, whereas, it
decreased for the Vail and Carbondale fences.
The JanuaryMarch winter period of 1977 was exceptionally mild and may
have resulted in fewer kills at some of the fences. Data
from severe winters would not be discarded from the sample,
similarly, data from mild winters probably should not be
discarded either.
No important changes occurred in the use of
28 one-way deer gates located in the Vail, Avon, Edwards, and
Eagle fences.
Only minor changes would be shown in benefit :cost ratios
previously described (Reed and Beck 1977), if they were
recalculated for this segment.
They will be presented in a
final analysis after the next and final segment.
A manuscript
draft titled "Methods of reducing deer-vehicle accidents:
benefit-cost analysis" was prepared during this segment.
Table 8. The mean annual number or pre- and post-installation
deer
highway kills and percent reduction for six 2.44-m fences adjacent to
Interstate 70 and Highway 82.
Mean annual
preinstallation
mortality

Mean annual
postinstallation
mortality

Percent
reduction
mortality

Fence

(Hwy)

Length of
hwy "fenced
km (Miles)

Vail

(1-70)

2.4 (1.5)

36 (3)

11.8 (8)

67.4

Avon

(1-70)

3.6 (2.3)

28 (1)

4.4 (5)

84.3

3.6 (2.3)

27 (1)

7.4 (5)

72.6

7.7 (4.8)

167 (1)

18.5 (4)

88.9

1.8 (1.1)

10 (3)

1. 6 (6)

83.3

1.8 (1.1)

14 (5)

5.3 (3)

61.9

Edwards
Eagle
Diamond

(1-70)

(1-70)

S (82)

Carbondale

(82)

Cumulative

Avg.

76.4

1/ (n) denotes the number of years of pre- and post-installation

data.

�-17-

Deer movements lateral or parallel to 2.44-m fences have been
further documented at the Vail, Avon, and Eagle fences (Table 9).
Based on 141 lateral movements of radio-collared or neckbanded
deer north of the Eagle 2.44-m fence, females and males have moved
lateral to the fence for mean distances of 0.588 and 0.762 km,
respectively.

Table 9. Female and male deer mean lateral movements (km) along or adjacent
to the field side or highway side of 2.44-m fences at Vail, Avon, and Eagle
as determined under several conditions.
Vail
Side of Fence

Female

Avon
Male

Female

Eagle
Female

Male

Male

Field Side

II

Apparent-

0.900(n=3)110.SOO(n=2)

31

Observed-

O.141(n=5)

0.588 (n=107) 0.762(n=34)
O.280(n=1)

Hwy Side
Apparent

O.617(n=3)

Observed

0.385(n=4)

O.506(n=4)

Observed &amp;
harassed!!..!

O.480(n=4)

--

O.975(n=4)

O.646(n=5)

IIMovements estimated by noting the locations of radio-collared or neckbanded
deer at selected points along the fences at widely separated time periods.

21

- (n) denotes number of lateral distances.

31

_...
Movements

observed

during

their entire duration.

~/Animals were usually chased with a vehicle at night until they either
escaped from the fenced side of the highway or "broke" back against the
vehicle.

Job 6 - Response

of deer to guards.

Four deer were tested on deer guard prototype V (black and white
scintillation or rotary motion pattern described by MacKay [1957]).
All deer crossed the prototype, three walking and one bounding,
without exhibiting behavioral responses indicative of much reluctance.

�-18-

Due to the difficultly of preparing this prototype, it is
probable that minimal conditions for the perception of rotary
motion (Braunstein 1977) had not been met at a low oblique
angle (about I m in height to match visual level of deer).
This prototype will not be tested further.

LITERATURE

CITED

Braunstein, M. L. 1977. Minimal conditions for the perception
rotary motion.
Scand. J. Psychol. 18:216-223.

of

Gallagher, V. P., and P. G. Meguire.
1974. Contrast requirements of
urban drivers.
Fed. Highway Admin. Rep. Number FHWA-RD-74-76.
72pp.
Hediger,

H.

London.

19pp.

Howe, C. W. 1971. Benefit-cost analysis for water system planning.
Geophysical Union.
Washington, D. C. 144pp.

Amer.

MacKay, D. M.
patterns.

1950.

Wild animals in captivity.

Butterworth,

1957. Moving visual images produced by regular stationary
Nature 180(4591):849-850.

Myers, G. T. 1969. An investigation of deer-auto accidents.
Pages 147178 in Game Res. Rep., July, Part 2. Colorado Game, Fish and Parks
Div. Fed. Aid.
Pojar, T. M. 1972a. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 267-275 in Game Res. Rep., July Part 3.
Colorado Div. Wildl. Fed. Aid.
Pojar, T. M. 1972b. An evaluation of deer-proof fence length required
to prevent deer movements on or across high speed highways.
Pages
305-310 in Game Res. Rep., July, Part 3. Colorado Div. Wildl. Fed.
Aid.
Pojar, T. M., R. A. Prosence, D. F. Reed, and T. N. Woodard.
1975.
Effectiveness of a lighted, animated deer crossing sign. J. Wildl.
Manage 39(1):87-91.
Reed, D. F. 1971. Deer underpass evaluation.
Pages 341-351 in Game
Res. Rep., July, Part 3. Colorado Div. Wildl. Fed. Aid.
Reed, D. F. 1974. An evaluation of 8-foot fence length required to
prevent deer movements on or across high speed highways.
Pages
313-320 in Game Res. Rep., July, Part 2. Colorado Div. Wildl. Fed. Aid.
Reed, D. F. 1975. An evaluation of 8-foot (2.44-m) fence length required
to prevent deer movements on or across high speed highways.
Pages
315-320 in Game Res. Rep., July, Part 2. Colorado Div. Wildl. Fed. Aid.

�-19-

Reed, D. F. 1976a. Evaluation of deer overpasses.
Pages 247-251 in
Game Res. Rep., July, Part 2. Colorado Div. Wildl. Fed. Aid.
Reed, D. F. 1976b. Evaluation of deer underpasses.
Pages 223-229 in
Game Res. Rep., July, Part 2. Colorado Div. Wildl. Fed. Aid.
Reed, D. F., T. M. Pojar, and T. N. Woodard.
1974. Mule deer responses
to deer guards.
J. Range Manage. 27(2):111-113.
Reed, D. F., T. N. Woodard, and T. M. Pojar.
1975. Behavioral response
of mule deer to a highway underpass.
J. Wildl. Manage. 39(2):361-367.
Reed, D. F., and T. D. I. Beck.
1977. Deer-vehicle accidents statewide
and methods and devices to reduce them. Pages 23-41 in Game Res.
Rep., July, Part 1. Colorado Div. Wildl. Fed. Aid.
Scott, J. P.
Chicago.

1958. Animal behavior.
349pp.

2nd ed.

University

of Chicago Press,

Woodard, T. W. 1973a. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 163-170 in Game Res. Rep., July, Part 2.
Colorado Div. Wildl. Fed. Aid.
Woodard, T. W. 1973b. An evaluation of deer-proof fence length required
to prevent deer movements on or across high speed highways.
Pages
197-202 in Game Res. Rep., July, Part 2. Colorado Div. Wildl.
Fed. Aid.
Woodard, T. W. 1974. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 293-298 in Game Res. Rep., July, Part 2.
Colorado Div. Wildl. Fed. Aid.
Woodard, T. W. 1975. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 295-303 in Game Res. Rep., July, Part 2.
Colorado Div. Wildl. Fed. Aid.
Woodard, T. W. 1976a. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 215-221 in Game Res. Rep., July, Part 2.
Colorado Div. Wildl. Fed. Aid.
Woodard, T. W. 1976b. Effects of highway lighting on number of deer killed
by vehicles.
Pages 231-236 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. Fed. Aid.
Woodard, T. W. 1976c. An evaluation of deer-proof fence length required
to prevent deer movements on or across high speed highways.
Pages
239-244 in Game Res. Rep., July, Part 2. Colorado Div. Wildl. Fed.
Aid.

Prepared by ~~~~
~_.__~~_f
__
~
Dale F. Reed
Wildlife Researcher

_

��-21-

JOB PROGRESS

July,

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Job Title
Period

1

Guidelines

Covered:

Personnel:

Big Game Investigations

W-126-R-1

Work Plan No.

Job No.
for Wildlife

September

1978

1

---------------------------------

Habitat Manipulation

1, 1977 - June 30, 1978

W. H. Rutherford,

W. D. Snyder

ABSTRACT
Literature review and interviews have provided a background of information,
from which a manuscript rough-draft is being prepared, on techniques and
procedures for manipulation of wildlife habitat.

��-23-

GUIDELINES

FOR WILDLIFE

William H. Rutherford

HABITAT MANIPULATION

and Warren D. Snyder

P. N. OBJECTIVE
To publish a comprehensive manual of directional habitat management guidelines
for sustaining and enhancing wildlife populations for consumptive and nonconsumptive uses on Division of Wildlife properties.
SEGMENT OBJECTIVE
To review literature and collect background information on habitat management
techniques for all wildlife species listed in Program Narrative.
METHODS AND HATERIALS
Approaches toward the literature review included use of the Fish and Wildlife
Service Reference Library, The Colorado State University Library, The U.S.
Forest Service Rocky Mountain Forest and Range Experiment Station Library, and
the Division of Wildlife Research Center Library.
In addition to the review
of literature, other approaches included interviews with Division of Wildlife
Regional Biologists, and Denver Office game and non-game management biologists.
Literature references were catalogued and abstracted on 3x5 file cards, and
on edge-punch cards.
Interviews were conducted by using a previously established
set of questions, plus informal and random conversation, with notes being taken
by the interviewer(s).
RESULTS
Card reference files of 130 pieces of literature applicable to big game and
furbearer habitat manipulation were established.
These were read and abstracted.
Many more pieces of literature were read, and rej ected as being not applicable.
Several hundred pieces of literature applicable to small game and non-game
habitat manipulation were reviewed, and many of these indexed on edge-punch cards.
Priority was placed on references conta~ning habitat manipulation and management
recommendations.
In addition, basic ecological information was collected for
most wildlife species.
At the end of this segment (June 30), a considerable portion of the manuscript
has been rough-drafted.
It is anticipated that only limited additional literature
review will be required during the next segment, and that most effort will be
directed toward finishing the manuscipt, submitting it to referees for critique,
revising and rewriting, and setting up plans for printing and publishing.
Also,
selected wildlife research personnel with backgrounds in wildlife habitat modification research will be interviewed before the final draft of the manuscript
is completed.

��July,

1978

-25-

JOB PROGRESS

State of

COLORADO
----------------------------

Project

No.

Work Plan No.
Job Title
Period

Covered:

Personnel:

W-126-R-1

REPORT

Big Game Investigations

7

Job No.

1

---------------------------------

Black Bear Investigations
April 1, 1978 - June 30, 1978
Thomas D. I. Beck

ABSTRACT
A PROGRAM NARRATIVE for black bear studies to be conducted during 1 July 197830 June 1987 was completed and approved.
Field surveys of possible study
areas based on recommendations from DOW personnel were conducted although a
final decision on study area selection was not made.
A considerable volume of
literature on black bear and forest ecology was reviewed.

��-27-

BLACK BEAR INVESTIGATIONS
Thomas D. I. Beck
P. N. OBJECTIVES
1.

Develop techniques
levels.

to accurately

2.

Determine

preferences

habitat

and precisely

estimate

bear population

of selected bear populations.

SEGMENT OBJECTIVES
1.

Select a study area.

2.

Develop

a detailed

3.

Habitat

type analysis

PROGRAM NARRATIVE

for Black Bear Investigations.

of study area.

RESULTS Pu~D DISCUSSION
A PROGRAM NARRATIVE
and accepted.

covering

the period

1 July 1978 - 30 June 1987 was completed

Numerous field personnel with the Colorado Division of Wildlife have been
contacted regarding possible study areas.
Field surveys have been conducted
in most of the recommended areas in western Colorado and are being continued.
Field work has been restricted by the heavy snowpack and subsequent inaccessibility of many areas until mid-June.
Consequently, selection of a final study
area has not been completed.
Considerable time was spent on reviewing literature on forest ecology and forest
habitat-type mapping.
Projects by other research and management organizations
have been reviewed to assess techniques and prevent unnecessary duplication of
work.
No field studies were conducted pending final selection of a study area.

Prepared

$_A_•. _WV_"'-

by -,-1!-,-~~·__
_£_Thomas D. I. Beck
Wildlife Researcher

_

��July 1978
-29-

JOB FINAL REPORT

State of

COLORADO

Project No.

W-38-R-33

Deer-Elk

Investigations

8
Job No.
14
Management Implementation of Quadrat Census Technique
to Estimate the Size of the Middle Park Deer Population

Work Plan No.
Job Title

Period Covered:
Personnel:

April

1, 1977 to June 30, 1978

D. J. Freddy, B. McCloskey,

L. H. Carpenter.

ABSTRACT

Estimated deer density in Middle Park in January, 1978 was 15.6 ~ 4.0 deer
per square mile of winter range (P ~ .10) (6.0 ± 1.5 deer/km2).
This
density projected to a total population estimate of 9,182
2,351 deer.
Management personnel have been sufficiently trained to continue the
census. The census should be continued as it provides a quantified estimate
of the size of an important deer population over a lengthy
time period.

±

�-30-

RECOMMENDATIONS
1.

The census should be continued by management personnel with
consideration given to dropping three relatively unimportant
to save time and expense.

strata

�-31-

MANAGEMENT
TO ESTIMATE

IMPLEMENTATION OF QUADRAT CENSUS TECHNIQUE
THE SIZE OF THE MIDDLE PARK DEER POPULATION
David J. Freddy

PROGRAM NARRATIVE

OBJECTIVE

Instruct management personnel on how to conduct the quadrat
deer in Middle Park, Colorado.

census of

SEGMENT OBJECTIVES
1.

Explain the theory and design of the census technique
personnel.

2.

Fly the census with management

3.

Assist management

personnel

to management

personnel.

in analyzing

census data.

METHODS AND MATERIALS
During the past 8 years, a stratified random quadrat census technique using
a helicopter has been employed to estimate the size of the Middle Park deer
population (Gill 1969; 1971). This technique has provided quantifiable
estimates of population size within known precision limits.
Deer are counted on 80, square·-mile permanently marked quadrats selected at
random from 8 sampling strata containing a total of 588 square-miles.
The
census is usually conducted in January using a Bell G47B-l or similar
helicopter.

RESULTS AllD DISCUSSION
Estimated deer density in Middle Park in January, 1978 was 15.6 + 4.0 deer
per square mile of sampled winter range (6.0 + 1.5 deer/km2).
This density
yielded a projected population estimate of 9,182 + 2,351 deer (P &lt; .10)
(Table 1). Snow depth was sufficient to force deer onto the sample area,
unlike the winters of 1976 and 1977. From the standpoint of snow and deer
distribution, this was the best census since January, 1975.
WCO, Bruce McCloskey flew the census as a training exercise and also
participated in statistical calculations of the population estimate (Table 2).
He has been sufficiently trained to continue the census without further
involvement of research.
The census should be continued as it provides a
quantified estimate of the size of an important deer population over time.

�Table 1. Numbers of deer counted per quadrat within eight sampling strata in four deer population sub-units, Middle Park, Colorado, January,
1978. Data entries are in numeric sequence for each strata.
Granby
Low Density

Williams Fork R.
Low Density

Troublesome R.
High Density

23

14

14

0

0

0

0

13

0

0

0

88

0

0

0

0

70

0

17

0

38

0

0

0

116

19

24

0

22

0

0

0

61

0

28

0

24

0

44

0

25

0

62

0

43

47

75

104

0

53

0

2

0

12

187

0

4

127

0

0

0

Muddy Creek
High Density

Muddy Creek
Low Density

Blue River
High Density

Blue River
Low Density

0

0

103

0

0

0

0

0

0

0

Williams Fork R.
High Density

0

34
0

0

----------------------------------------------------------------------------------------------------------------------------------------------

I
W
tv
I

�Table 1.
January,

Numbers

of deer counted

per quadrat within eight sampling strata in four deer population
sequence for each strata.
(Cont'd).

sub-units,

Middle

Park, Colorado

1978. Data entries are in numeric

Muddy Creek
High Density

Muddy Creek
Low Density

Blue River
High Density

Blue River
Low Density

Williams Fork R.
High Density

Williams Fork R.
Low Density

Troublesome R.
High Density

14

217

Granby
Low Density

o
o
o
o
5
I

l:yh 229

16

804

66

152

w
w

0

I

nh

20

12

12

9

5

5

10

7

yh

11.5

1.3

67.0

7.3

30.4

2.8

21.7

0

Nh 116.6

115.0

73.1

53.0

28.1

63.3

64.1

74.5

Wh

.198

.196

.124

.090

.048

.108

.109

.127

(W )2
h

.039

.038

.015

.008

.002

.012

.012

.016

Sh

2

l:yh
nh
yh

726.1

10.7

3319.1

1129.3

260.8

Total deer counted in each stratum
Number of sample units (square miles) per stratum
Mean number of deer counted per sample unit

39.2

Nh

Total square miles per stratum

W
h

Ratio of Nh
l:Nh

513.6

S~

= stratum

0
variance

2
2
= Yh(Yh)

~
h - 1

n

�-34-

Table 2. Summary of statistical calculations for Middle Park deer
census, 1978.
.2
587.7 ml.

N

2
Mean deer/mi

-

Variance of yst

Standard Error of yst
JVAR yst

-

Confidence limits for yst

=

2.410 = Sy

(yst ± T [Sy])

where T

1.66 at alpha = .10 and 71 degrees freedom; thus the C.L.

for yst

15.624 ± 1.66 (2.410) = 15.624 + 4.001

TOTAL POPULATION ESTIMATE WITH CONFIDENCE LIMITS
N(yst) ± N (T[Sy])
(587.7) (15.624) ± (587.7) (4.001)
9,182 ± 2,351 deer @ P&lt; .10

�-35-

About 3~ days are needed to complete the census as presently flown. To
save time and expense, consideration should be given to dropping the Muddy
Low, Williams Fork Low, and Granby Low strata. Since 1968, these 3 strata
have contributed less than 5 percent of the total population estimate,
(Freddy 1975, 1976). One drawback of dropping these strata is that at
population levels higher than have occurred since 1968, these strata may
become important if deer are distributed differently when densities are
higher.

LITERATURE

CITED

Freddy, D. J. 1975. Middle Park deer study-experimental harvest
regulations.
Colo. Div. Wildl. Game Res. Rep. July, Part 2:

209-240.

Freddy, D. J. 1976. Middle Park deer study-experimental harvest
regulations.
Colo. Div. Wildl. Game Res. Rep. July, Part 2:

257-282.

Gill, R. B. 1969. Middle Park deer study-population density and structure.
Colo. Game, Fish and Parks Dept. Game Res. Rep. July Part 1: 105-122.
Gill, R. B. 1971. Middle Park deer study-population density and
structure.
Colo. Game, Fish and Parks Dep. Game Res. Rep. July,
Part 2: 170-188.

��July,

-37-

1978

JOB PROGRESS REPORT

State of

COLORADO

-----------------------------

Project No.
Work Plan

W~38-R-33

Deer-Elk

Investigations

14
---------------------------

Job Title
Period Covered:

9
Job No.
Nutritional Basis for Quantifying Capacity
of Winter Ranges to Support Deer

April 1, 1977 to June 30, 1978

Personnel: Len H. Carpenter, M. A. Kautz, L. L. Strong, D. J. Freddy,
Cathy A. Moore, Linda E. Sanders, Paul H. Neil, Bruce D. Trindle, Kate
Trindle, and Steve C. Torbit.

ABSTRACT

Eighteen mule deer fawns were successfully raised in Fort Collins and at
Junction Butte Research Center near Kremmling for studies of protein and
energy relationships of mule deer in winter.
Birth weights and growth
rates for these fawns are presented.
All fawns were trained to use
digestion cages and weigh chutes. Additionally, four of the most tractable
fawns were trained for food habit determinations and to wear a face mask
for oxygen consumption measurements.
A thesis summarizing the relationship
between heart-rate and energy expenditure for three mule deer fawns is
presented.

��-39-

NUTRITIONAL BASIS FOR QUANTIFYING
CAPACITY OF WINTER RANGES TO SUPPORT DEER

Len H. Carpenter

PROGRAM NARRATIVE
Develop procedures
to support deer.

for quantifying

OBJECTIVE

the capacity

of Middle Park winter

ranges

SEGMENT OBJECTIVES
1.

Estimate

protein requirements

of mule deer fawns in winter.

2.

Compare forage intake estimates of free-ranging
esophageal fistulated deer in pastures.

mule deer with

3.

Estimate

activities

4.

Develop sampling methodology for estimating
availability of deer forage in winter.

energy costs associated

with selected

of mu~e deer.

annual production

and

METHODS AND MATERIALS
Methods and materials employed in this study have been previously described
(Gill and Baker 1974, Carpenter and Baker 1975, Carpenter 1976, and 1977,
Baker 1976). To secure adequate numbers and properly trained deer to meet
objectives in this study the major work effort this segment was dedicated to
fawn rearing and training.
Sufficient deer were not available to adequately
complete certain segment objectives.
Estimating Protein
Requirements of Mule Deer in Winter
This portion of the study is being conducted by Stephen Torbit, graduate
student at Colorado State University.. This study is in initial planning
stages and results will be presented next segment.
Method for Estimating Energy
Expenditures of Free-Ranging Mule Deer
A method for measuring energy expenditures of free-ranging mule deer was
developed by Marie A. Kautz, Colorado State University.
A thesis covering
this work is presented in Appendix A.

�-40-

Comparisons of Forage Intake With
Intact vs. Esophageal Fistulated Deer
This phase of the study was inactive during the past segment because no
surplus deer were available for fistulation.
This portion of th2 investigation
will be initiated in later segments.
Estimating
and Developing

Energy Requirements for Activity
Energy Budgets for Deer at Pasture

Data on energy requirements by various activities obtained by Kautz (1978)
and data on activity patterns of deer at pasture (Carpenter 1976) are being
incorporated into a simulation model for carrying capacity at Colorado State
University, (Hobbs and Baker 1977). This model will serve as a first approximation of carrying capacity of Middle Park winter ranges.
Estimating Annual Production
and Availability of Deer Forage in Winter
This portion of the study is being funded by U.S. Department of Agriculture,
U.S. Forest Service, Rocky Mountain Forest and Range Experiment Station
through its studies concerning evaluation of wildlife habitat.
The work is
closely tied to determination of carrying capacity of Middle Park deer ranges.
The study is in initial planning stages and pertinent results will be
summarized next segment.
Fawn Rearing

and Training

Fourteen wild does were trapped and transported to the Junction Butte Research
Facility to maintain until fawning.
Five does were trapped at the Little Hills
Experiment Station with the remaining nine being trapped in the Middle Park
area. All trapping was done during the period February 12 to 28, 1977. Fawns
were also obtained from two tame does maintained at the Junction Butte facility.
All wild does were released from the pen on June 24, 1977.
Fawns were removed from the dam between 24-48 hours after birth.
They were
immediately weighed, sexed, and checked for general health and abnormalities.
Each fawn was assigned a number and identified by a necktag.
Fawns of similar
age were housed in individual isolation pens, usually in groups of three. They
were trained to nurse from plastic baby bottles.
The formula was two parts whole milk to one part evaporated milk with buttermilk making up 1/10 of total daily volume.
Baby vitamins were supplemented
at the rate of I ml daily up to 4 weeks of age, I ml every other day up to
8 weeks, and I ml every 3 days from 8-12 weeks. The following schedule for
feeding was followed.
with dam
Day I

0-2 weeks

2 oz feeding, five times/day
from 6 AM - 10 PM

�-41-

Feeding Schedule

(cont'd)
2-4 weeks

3 oz feeding,

4 times/day

from

6 AM - 10 PM
4-6 weeks

3-5 oz feeding, 3 times/day
(morning, mid-day, evening)

6-8 weeks

6 oz feeding, 2 times/day,
morning and evening

8-11 weeks

6-8 oz feeding, I time/day

Fawns were offered alfalfa hay and pelleted grain ration ad libitum after
two weeks.
After weanin&amp; all fawns were maintained on alfalfa hay and
pellets were offered ad libitum.
Fawns were weighed weekly from birth to 14 weeks.
They were then weighed
at bi-weekly intervals until 39 weeks of age when they were weighed at
tri-weekly intervals.
Four of the most tractable fawns were trained to
wear face masks for oxygen consumption measurements (Kautz 1978) and to
participate in forage selection trials. All fawns were trained to enter
and accept weigh chutes, digestion cages, and transportation cages. A
whistle was used at all feeding sessions to train fawns to respond to the
sound. This facilitated moving fawns and commanded their attention during
training sessions.
Training continues daily.
Diarrhea was encountered throughout the first 7 weeks of life. In most cases
it was treated and controlled by combinations of isolation, decreases in
amounts formula, kaopectate, "life saver" solutions, chloramphenicol, or
sulfonamide drugs.
Of those fawns that died selected individuals were sent
to the Animal Disease Center at Colorado State University for necropsy.
RESULTS AND DISCUSSION
Method for Estimating Energy
Expenditure of Free-Ranging Mule Deer
A thesis covering

this work is presented

in Appendix

A.

Fawn Rearing and Training
Seventeen fawns were obtained from the 14 wild and two tame does at Junction
Butte. Twelve were males, 5 were females.
There were 7 sets of twins and
3 singles.
Three fawns died within 30 hours of birth, two to a tame doe
suffering from chronic wasting, which died five days after giving birth, and
one single fawn born to a wild doe.
Female fawns averaged about 70 grams heavier than male fawns however, two of
the males (numbers I and 2) were born to the chronic wasting doe and were

�-42-

abnormally underweight (Table 1). Birth weights of male fawns that died
within 86 hours of birth averaged 984.2 grams, less than birth weights
of surviving male fawns (Table 1) of 11 fawns that died during the first
year of life, death in six fawns was attributed directly or indirectly to
chronic diarrhea (Table 2). The last fawn to die from diarrhea (fawn #104)
was diagnosed as having Giardia.
One other fawn (#125) was also diagnosed
as having Giardia and was successfully treated.
It is suspected that other
fawns suffering and dying from diarrhea also had Giardia but no diagnostic
tests were conducted to confirm the SUsplClon.
Apparently Giardia in mule
deer can be cyclic, appearing for a few days and then disappearing, only to
reappear a few days later (personal communication, Robert Davies, Wild
Animal Disease Center, Colorado State University).
Later tests showed that
diarrhea could be present for 2-3 days before any Giardia cysts would be
found in the feces.
Continued testing of healthy fawns has shown no presence
of Giardia cysts.

Table 1. Birth weights (grams) by sex of 17 fawns born at Junction Butte
Research Center during June, 1977.

Fawn Number

Male

150

3493

151
152

Female

3796
4388
3931

153
154

3401

155

3898

156*

2795

157

2810

158

3480

159

3600

160

3675

161

3313

162

2824

�-43-

Table 1. Birth weights (grams) by sex of 17 fawns born at Junction
Research Center during June, 1977.
(Cont'd).

Female

Fawn Number

Male

163

3244

1*

2012

2*

2117

3*

3645

+ std. dev.

3298.4

Mean

Overall mean

+ std.dev.

Butte

+ 691. 2

3368.2

+ 529.2

3318.9 + 632.1

* died within 86 hours of birth
Mean + std. dev. of surviving male fawns 3626.5 + 139.6
Mean + std. dev. early postnatal male fawn mortalities
hours of birth)
2642.3 + 753.2

Table 2.

(died within 86

History of fawns that died during fawn-rearing

Probable

Cause

Fawn No.

Single/Twin

Dam Source

Time to Death

1

twin

tame

27 hours

weak at birth

2

twin

tame

27 hours

weak at birth

3

single

wild

18 hours

unknown

156

twin

wild

86 hours

diarrhea

�-44-

Table 2.

History

of fawns that died during fawn-rearing

(Cont'd).

Single/Twin

Dam Source

Time to Death

Probable

161

twin

wild·

13 days

diarrhea

152

single

wild

86 days

urinary bladder

158

twin

wild

104 days

lepto-meningitis,
brain swelling

163

twin

wild

128 days

diarrhea

157

twin

wild

134 days

diarrhea

160

twin

wild

141 days

diarrhea-Giardia

104

twin

wild

248 days

diarrhea-Giardia

Fawn No.

Cause

rupture

Deaths of fawn numbers 152 and 158 were attributed to rare causes not related
to husbandry practices.
The causes of death in fawn numbers 1, 2, and 3 were
unknown but attributed to weakened conditions at birth.
On September 13, 1977, fourteen fawns were transported from Fort Collins to
the Junction Butte facility.
Shortly afterwards a form of blue tongue disease
was isolated from the tame deer herd in Fort Collins.
About October 1, several
fawns from both groups (Fort Collins and Junction Butte) began to show various
symptoms of poor health.
These symptoms varied from arthritic stiffness to
inappetence and lethargy.
These symptoms, with the presence of diarrhea,
continued until late November.
No definite causes of the poor health were
diagnosed in the fawns but it is suspected that blue tongue may have been a
factor.
To control diarrhea in these fawns, food (pelleted ration) was restricted.
Alfalfa hay continued to be offered ad libitum.
As a consequence of food
restriction and poor health, fawns did not gain weight during this period and
many lost weight.
The affects of ration restriction on body size (as compared
to healthy fawns maintained at Little Hills) is still visible.
Fawns raised in Fort Collins apparently gained weight more rapidly than did
fawns raised at Junction Butte based upon comparisons of available weight data
(Table 3 and 4, respectively).
Growth rates will be maintained on these deer
during the next year. Growth rates of the 18 Junction Butte fawns rearing
one year of age (353 days) averaged about 120 grams per day (Table 5).

�Table 3. Weights (grams) at birth and through eight weeks of age for 12 mule deer fawns born at Junction
Butte Research Center during June, 1977.
Fawn Number

Birth

1st week

2nd week

3rd week

4th week

5th week

6th week

7th we ek

8th week

150

3493

3585

3640

4210

5058

6160

6670

7037

8399

151

3796

3815

3760

4220

4830

5657

6210

6356

7718

152

4388

4130

4013

4335

4715

5200

6120

7037

9080

153

3931

3708

3690

3980

4920

6172

6920

8399

9534

154

3401

3381

3710

4250

5540

6645

8500

10442

12258

155

3898

3538

3680

3800

4620

5535

7070

8172

10215

157

2810

2860

2665

2900

3655

4615

5965

7491

9194

158

3480

3493

3590

4205

4860

5785

6350

7718

9080

159

3600

3487

3567

4J.70

4725

5750

6990

7718

9332

160

3675

3490

3420

3625

3800

4185

4450

4947

5570

162

2824

3020

3125

3905

5030

5720

7200

7945

9307

163

3244

3270

3140

3915

4245

5255

6630

7605

9307

Mean (grams)

3545.0

3481.4

3500.

3959.6

4666.5

5556.6

6589.6

7572.3

9082.8

Std. dev.(g)

451.8

337.3

363.0

396.1

534.0

676.9

949.7

1293.8

1549.6

Mean (kg)

3.5

3.5

3.5

4.0

4.7

5.6

6.6

7.6

9.1

Mean (Lbs )

7.8

7.7

7.7

8.7

10.3

12.2

14.6

16.8

20.1

I

.jO-

V1
I

�Table 4. Weights (grams) at birth and through eight weeks of age for 12 mule deer fawns born in Fort
Collins during June, 1977 and maintained at Junction Butte Research Area.

Fawn Number

Birth

1st week

2nd week

3rd week

4th week

5th week

6th week

7th week

8th week

103

2974

2729

3065

3632

4199

4881

5902

6470

7151

104

3341

3405

4200

5221

5789

7151

8286

9307

10329

105

3178

3632

4427

5108

6016

6924

7945

10102

10669

107

3450

3405

3973

4313

5335

6016

7151

7718

8967

109

3314

3292

3973

4540

5448

6583

8172

8967

10669

111

4041

3519

4200

5108

5902

7151

8513

9648

10896

4654

5448

6243

6583

7605

-9761

I

~

115

3441

3519

4313

118

3519

3519

4200

4994

5675

6810

7945

8399

119

3632

3746

4200

4767

5675

6583

7491

8172

120

3405

3405

3859

4540

5562

7151

8059

8967

121

4654

--

--

--

8172

9534

10669

123

3473

--

---

--

--

7151

9080

10669

Mean

3535.2

3416.6

4041.0

4687.7

5504.9

6734.7

7888.4

8891. 0

9777.4

SD

435.2

275.3

382.8

475.8

505.7

800.1

1006.5

1286.2

1335.4

Mean (kg)

3.5

3.4

4.0

4.7

5.5

6.7

7.9

8.9

9.8

Mean (Lbs)

7.7

7.5

8.8

10.4

12.1

14.8

17.4

19.6

21.6

0\
I

�-47-

Table 5. Growth rates (grams gained/day) for 18 fawns maintained
Butte Research Area for first year of life.

at Junction

Fawn Number

Age (Days)

103

364

37.6

103.3

105

359

44.9

125.1

107

356

37.2

104.5

109

356

39.5

111.0

III

360

39.9

110.8

115

349

44.0

126.1

118

349

41.3

118.3

119

345

46.3

l34.2

120

346

42.6

123.1

121

341

41.7

122.3

123

340

35.4

104.1

150

360

46.3

128.6

151

360

44.0

122.2

153

357

41.5

116.2

154

353

45.8

129.7

155

353

44.0

124.6

159

351

50.3

143.3

162

353

37.2

105.4

Mean

352.9

42.2

119.6

SD

6.7

3.8

11.0

Weight

(Kilograms)

Gain (Grams/day)

�-48-

LITERATURE

CITED

Bake r, D. L. 1976 • Energy requirements of mule deer fawns in winter.
Colorado State University, Fort Collins.
76p.
M.S. Thesis.
Carpenter, L. H., and D. L. Baker.
1975. Middle Park deer study - deer
habitat evaluation.
Colo. Div. Wildl., Game Res. Div., Fed. Aid
Proj. W-38-R-29 Game Res. Rep., July, Part 2. p. 241-263.
Carpenter, L. H. 1976. Middle Park deer study - deer habitat evaluation.
Colo. Div. Wildl., Game Res. Div., Fed. Aid Proj. W-38-R-31.
Game
Res. Rep., July, Part 2. p. 283-298.
Carpenter, L. H. 1977. Middle Park deer study - deer habitat evaluation.
Colo. Div., Wildl., Game Res. Div., Fed. Aid Proj. W-38-R-32.
Game
Res. Rep., July, Part 1. p. 65-83.
Gill, R. B., and D. L. Baker.
1974. Middle Park deer study - deer
habitat evaluation.
Colo. Div., Wildl., Game Res. Div., Fed. Aid
Proj. W-38-R-28.
Game Res. Rep., July, Part 2. p. 211-225.
Hobbs, N. T., and D. L. Baker.
1977. Systems modeling big game
populations - simulations of the carrying capacity of the Rocky
Mountain National Park elk winter range.
Colo. Div. Wildl, Game
Res. Div., Fed. Aid Proj. W-38-R-32.
Game Res. Rep., July, Part 2.
p. 125-200.
Kautz, M. A. 1978. Energy expenditure and heart rate of active mule
deer fawns. M.S. Thesis.
Colorado State University, Fort Collins.

Prepared

~A~,
_A-.----tc:~-

by ....:.~...L•••
~ ••••••• G"--~-=-t-=-.ilLen H.
Wildlife Researcher

_

76p.

�-49-

APPENDIX A

Marie Kautz Thesis

�/

-50-

THESIS

ENERGY EXPENDITURE

AND HEART RATE

OF ACTIVE MULE DEER FAWNS

Submitted
Marie

Vendeville

Fishery-and

In partial

Wildlife

fulfillment

for the Degree
Colorado
Fort

by
Kautz
Biology

of the requirements

of Master

of Science

State University

Collins,

Colorado

Summer,

1978

�-51-

COLORADO STATE UNIVERSITY

Summer

1978

WE HEREBY RECOMMEND THAT .THE THESIS PREPARED
UNDER OUR SUPERVISION
ENTITLED

BY MARIE VENDEVILLE

ENERGY EXPENDITURE

MULE DEER FAWNS
REQUIREMENTS

BE ACCEPTED

KAUTZ

AND HEART RATE OF ACTIVE
AS FULFILLING

FOR THE DEGREE OF

IN PART

MASTER OF SCIENCE.

Comrrlittee on Graduate Work

/ 7:1/!/ &lt;,.' /

Adviser

u.! - IJ!J'i' ~;Jj

-;

c-:

{J

I

Ll.Jd~ ~

/;,~."

.;'

ID

dla7

191'6

�-52-

ABSTRACT

OF THESIS

ENERGY EXPENDITURE

AND HEART RATE

OF ACTIVE MULE DEER FAWNS

The relationship
studied

in three

lowed to bed,
to evaluate

between

mule deer

stand,

walk,

the usefulness

energy

hemionus

and trot

outdoors

during

of heart

rate

for active

deer.

penditure

of humans

and domestic

The deer

collection.

of the electrocardiogram.
kcal kg

. -1
rmn

Correlation

ranged

statistically
of similar
weight.

Energy

coefficients

from 0.81 to 0.92
significant
weight,

There

due to activity

Heart

al-

rates

ex-

for use with
determinaby telemetry
from 0.065

from 49 to 96 beats
expenditure
deer.

from

and heart

There

in the relationships
did differ

ex-

energy

was measured
ranged

in order

of energy

expenditure

to be any difference

level or feed level.

hemionus)

was adapted

expenditures

for the three

but the males

did not appear

rate

for energy

difference

was

used to measure

animals

-0 75
-1
.
min
and heart

to 0.31

rate

Methodology

rate

winter,

as a predictor

wore a face mask for energy

tion by respiratory

and heart

fawns (Odocoileus

penditure

deer.

expenditure

was no

for two males

a female

of lighter

in the relationship

Although data were not adequate

�-53-

to provide
,

a precise

predictive

equation

for estimating

penditure

from heart

rate for active deer,

heart

rate

energy
appears

expromis-

ing for this use.

Marie Vendeville Kautz
Department
of Fishery and
Wildlife Biology
Colorado State University
Fort Collins, Colorado 80523
Summer,
1978

�ACKNOWLEDGEMENTS

Funding
ative Wildlife
through

for this research
Research

Colorado

experimental
life.

Office space

ative

Wildlife

mitters

were provided

Research

at Colorado

Aid Project

and vehicle

Gas analysis
tory

Unit by the Colorado

Federal

animals

equipment

Division

W- 38-R.

Cooper-

of Wildlife

Pen facilities

by the Colorado

were provided

was provided

State University.

Additional

to the Colorado

and

Division

of Wild-

by the Colorado

Cooper-

Unit.

and set up telemetry

Laboratory.

was provided

R. J.

equipment

suggestions

by the Metabolic
Scott built heart

Labora-

rate

trans-

loaned by the CSU Surgical

on telemetry

were given by

J. J. Cupa l , .
I thank my committee
penter,

D. E. Johnson,

guidance

on all aspects

provided

additional

Gilbert

shared

of the study.

Remmenga

L. H. Carand

and R. B. Gill

support,

and D. L.

on the thesis.

built the face mask,

p. H. Neil,

responsibilities

K. R, Russell

and administrative

gave he lpful suggestions

collection.

W. W. Mautz,

and G. M, Van Dyne for suggestions

guidance

E. O. Crispe
data

members,

and C. A. Moore assisted

D. W. Reichert

for fawn raising

and B. D. Trindle

and deer

and D. C. Bowden gave statistical

care.

advice.

E. E.
Photographs

in

�-55-

to L. J.

were taken by W. K. Seitz.

Appreciation

Niemann,

D. M. Roche and D. L. Baker.

D. G. Milchunas,

My husband

Ed Kautz

study.

providing

constant

ranged

from fawn care

was a friend

encouragement

to computer

is also expressed

and helper

programming.

scientists

at Colorado

their

openness,

these

pursuit

and their

of this goal.

the

and help on jobs which

thank a group of female
example

throughout

Finally,

I want to

State University.

women sustained

By

me in my

�-56-

TABLE

OF CONTENTS

Chapter
I

11

III

INTRODUCTION
Literature
Review
Methods
•.••
Energy Expenditure
Wild Ungulates

1

........

2
2

of
5

8
8
11

METHODS AND MATERIALS
Animals
Physical
Facilities
Apparatus
.•..••.
Heart Rate Apparatus
Respiration
Apparatus
and
Gas Analysis
..•••
Face mask
.•.•••
Flowmeter
and collection bags
Gas analysis
.••.
Trial Procedures
Weather Parameters
•.•.
Calculation
of Energy Expenditure
•••..
Activity Data ••..•..•••
Contro 1 of Activity
••
. . • • • • • •
Trials on Fasted Deer
. . • • • • . •••
Statistical
Analysis
Individual Deer
All Deer • . . • . • • • •
Confidence Intervals
for Predicting
Energy Expenditure
. • . • . • • ••.

12
12

14
14
16

19
19
21
21
23
23

27
27
27
30

31

RESU LTS •.•••••.•.•
Effectiveness
of Method .
Problems
with Respiration
Equipment
. • • . . . • . . • • • .
Relationship
between Energy Expenditure
and Heart Rate
. . . • • • .
. • • •
Individual Deer
All Deer •..•
Effect of body weight .
Confidence Intervals
for Predicting
Energy Expenditure
.•..••.••

...

32
32
33

34
34
39
41
41

�-57-

TABLE

OF CONTENTS

(continued)

Chapter
45
45

DISCU SSION . . . . • • . • • . • . . . •
Comparisons
to Literature
Values
Effects of Activity Leve land
Feed Level
.
Comparison
of Regression
with
Literature
Results
.•••
Individual Differences
in
Regression
Equations
•...
Confidence of Predicting
Energy Expenditure
from Heart Rate
Application
of Results
...

IV

51
51
54

56
57

0

..........
... ..................

SUMMARY AND CONCLUSIONS

V

LITERATURE
APPENDIX
A

B

C

CITED
•

•

•

•

•

•

•

•

•

III

•

•

•

•

•

0

•

•

•

•

•

•

•

•

•

•

•

58
61

67

Data collection
form modified from
Consolazio
et al , (1963) for measuring
mule deer energy expenditure

68

Derivation
of formula to calculate
oxygen
consumption
and energy expenditure
from
volume and fractional
concentration
of
oxygen in expired air, assuming
volume of
methane produced
is insignificant
. • . • .

69

1
Weights (kg), oxygen consumptions
(rnl miri- ),
energy
expenditures
(kcal kg-O. 75 min-1),
heart rates (beats min - 1), and activities
(percent
bedding,
standing,
walking,
trotting)
for trials using three fasting and fed
mule deer fawns in winter
....•.

71

�-58-

LIST OF TABLES

Table
1

2

3

4

5

Sexes, ages (days), and weights (kg) of
mule deer subjects
at times of weaning
and experiment,
and peak weights of
fawns which were fasted
• . . • •••

9

Range of energy expenditure,
heart rate,
and activity measured
for three mule
deer fawns during January and February

25

Number of simultaneous
energy expenditure
and heart rate measurements
at different
levels of activity and feed for three mule
deer fawns in winter,
and comparisons
for effects of activity level and feed level

28

Mean square residuals
for regression
of
energy expenditure
on heart rate for three
mule deer fawns at four activity levels
and two feed levels
•.....•••..•

•

•

•

•

0

•

40

Heart rates of mule deer and white-tailed
deer for different
activities

46

6

Measurements
of fasting and resting
production
for young wild ungulates

48

7

Measurements
of energy cost of activities
for wild ungulates
•.•...•••••.•

8

Regression
heart rate

9

.............
heat

.

49

equations of energy expenditure
on
for white-tailed
deer and mule deer

53

1
Weights(kg),
oxygen consumptions
(ml min- ),
energy expenditures
(kcal kg-O. 75 min-1),
heart rates (beats min-1),
and activities
(percent bedding,
standing,
walking, trotting) for
trials using three fasting and fed mule deer
fawns in winter
....•.•..••..•••

71

.

�-59-

LIST OF FIGURES

Page

Figure
1

Face ma sk for rne a s ur ing energy expenditure
of mule deer fawns
..•

15

0

2

3

4

Measuring
energy expenditure
rate of mu le deer fawn

and heart
0

•••

0

Schexnatic breakdown of trials into four
activity levels on the basis of percent of
t irne spent at specific activities

26

Relationship
of energy expenditure
and
heart rate for deer no. 51 by activity
level and feed level
.•••

35

Relationship
of energy expenditure
and
heart rate for deer no. 53 by activity
level and feed level
•...•••..••••••••

36

Relationship
of energy expenditure
and
heart rate for deer no. 55 by activity
level and feed level
.........•

37

Regressions
of energy expenditure
on
heart rate for three rnu le deer fawns
in winter
. . . . . . .

38

Regressions
of energy expenditure
on
heart rate and 95 percent confidence intervals
for predicting
me an energy expenditure
fr om heart rate for two 40 kg
rria Ie mu le deer and one 30 kg fexnale
mul e deer in winter

42

Regression
of energy expenditure
on
heart rated and 95 percent confidence interval for predicting
rne an energy expenditure
f r orn heart rate based on data
f'r om three rnul e deer fawns in winter

44

0

5

6

7

••

................

8

9

18

�-60-

CHAPTER

I

INTRODUCTION

Throughout

the history

of wildlife

been concerned

with maintaining

levels

to provide

sufficient

so high as to cause

range

trends,

trends

duction

and use of key browse

heavily

on the knowledge

in aerial

basis
indices

species.

approach

of weather

to deer

(973).

and nutrient

The possibility

of functional

nutrient

is intuitively

harvest

and trends

in pro-

techniques

Interpretation

poor

statistical

is a product

of managing

ungulate

rather

and computer

of Moen's

between

supply of the range,

nutrient

elaborated
herds

technology

information.

Although information

needs

supply is still

incomplete

by

on the

than empirical

of the necessary
and range

depend

(Gill 1976}.

as the relation

relationships

appealing,

conditions,

management

capacity

but not

utilizing

and skill of the field worker.

of carrying

of the animals

These

at

have been de-

condition,

of counts,

and the "myth of the browser"

(1973) concept

Robbins

types

have

species

for harvest

Techniques

and range

or other

by variability

A recent

needs

condition

managers

of big game

of animals

deterioration.

to evaluate

is complicated

populations

an excess

vised

techniques,

herd

management,

trend

enables

on both deer

(Moen 1973), numerous

use

�-61-

studies

have provided

of carrying

capacity

Part
rado,

sufficient

data for making basic

(Wallmo et al , 1977).

of an ongoing r e s aar-ch program

is aimed

(Odocoileus

at quantifying

hemionus

vious studies

the energy

hemionus),

(Milchunas

quality

tenance

of fawns in winter

(Baker

budgets

of deer

(Carpenter

portance

using data on energy
activity
of energy

costs

cedures

difficult

uring

costs

requirements

of an attempt

energy

mentioned
activities

was to investigate

and heart

rate

for active

Literature

Review

Pre-

1976). nufor main-

and time

1976).
to assess

budget of deer

above.

requires

useful

the relationship

in winter,

deer.

and pro-

Thus the primary

methodology
deer.

between

with

measurement

equipment

of active hand -reared

the im-

in conjunction

However,

to use even on hand -reared

expenditure

(Carpenter

of sp ec if ic activities

for specific

Colo-

of mule deer

1976), and activity

of this study was to provide

energy

jective

in the total

and time budgets

objective

1977), energy

study was part

of activity

needs

utilization

tritional

in pastures

in Middle Park,

for use in such calculations.

have dealt with forage

The present

calculations

for meas-

A related

ob-

energy

expenditure

of measuring

energy

expenditure

This method,

referred

to as

deer.

Methods
The most
requires

widely used method

collection

of expired

air.

�-62-

indirect

respiration

calorimetry,

tion chamber s with confined
highly artificial
of activity,

conditions

thermal

Indirect
tory provided
measured

is employed

animals.

Respiration

and restrict

subjects

conditions,

respiration

and sensory

calorimetry

that air expired

and sampled.

success

from white-tailed

deer

a tracheotomized

whitetail

Other researchers
expenditure
isotope

energy

employing

carbon-14

with domestic

1968,1972;

1971), domestic

reindeer

and caribou

Although respiratory
quires

the subject

urine

must be sampled

to estimate

dioxide

cattle

collection

is unnecessary,

An animal

air.

A radio-

entry

rate)

(Young 1970), and

the technique
infusion,
trained

pump can move about and forage

without direct

during

the measurement

Carbon dioxide entry

carbon

dioxide production.

Each liter

energy

(Young and McEwan 1975).

a pump for isotope

period.

(1978) used

sheep (Young and Corbett

tarandus

regularly.

were used
collection

of expired

Rangifer

to carry

air using a

masks

or

expenditure.

(carbon

has been used successfully
Corbettetal.

expired

and Fair

methods

collection

the laboracollected

for respiratory

have developed

range

is either

Face

virginianus,

to estimate

which do not require

technique

to collect

(1974),

impose

environment.

subjects.

Odocoileus

chambers

can be used outside

It is possible

by Mattfeld

respira-

from a normal

from the subjects

face mask or from tracheotomized
with limited

in laboratory

of carbon

re-

and blood or

to carry

the

hum.an interference
rate

reflects

dioxide produced

is

�-63-

equivalent

food substance

oxidized

dioxide

entry

rate

is related

to

expenditure.
A second

18

radioisotope

water

(D 0
2

energy

expenditure

).

for long-term

studies

to energy

and pulmonary

heart

rate

and energy

1969), rodents
Morhardt

with respect

include

rate.

heart

to their

rate,

'!'he relationship

between

and Hervey

1960, Datta and Ramanathan
1967. Brockway

sheep (Webster
and Hart

in humans

rela-

respira-

was studied

(Segrem

(Malhotra

1969,

and McEwan

1967,§!:.b, Lund and Folk 1976,

1971), and white-tailed

of humans

between

heart

Some of the rodents

and rodents

indicated

rate

and energy

studied

by Morhardt

to have non-linear

conducted

suitable

deer

(Holter

et at.

1978).

Studies
tionship

These

ventilation

and Morhardt

1976, Fair

it is most

by Lifson

expenditure

et a l , 1966),

Sharkey

the

As described

(1966),

have been studied

rate

use of doubly-labelled

animals.

expenditure.

Booyens

animals.

and McClintock

of small

tory

et ale 1963,

involves

can also be used to estimate

of unrestrained

Other parameters
tionship

technique

The technique

et a l , (1955) and Lifson

peared

on the

1945, citing Lusk 1928 and Zuntz and

(Brody

1901), and thus carbon

Schumberg
energy

5.05 and 6.69 kcal of energy depending

to between

with animals

relationships.

a strong

expenditure

linear

rela-

in most cases.

and Morhardt

(1971) ap-

The studies

of sheep were

that were tracheotomized,

wore face masks,

�-64-

or were in ventilated

hoods.

lationship

between

heart

strated,

while strong

weak relationship
penditure

were

oratory

(Holter

correlation

For six sheep in the two studies

rate

and energy

relationships

expenditure

correlated

et al , 1976).

Heart

rate

for white-tailed

However,'

in these parameters

was demon-

were found for two sheep and a

was found for a third.
strongly

no re-

and energy
deer

(I978),

Fair

for a white-tailed

in the lab-

reported

deer

ex-

a weak

studied

out-

side the laboratory.
Respiratory

rate

and pulmonary

than heart

rate

as pred ictors

useful

(1973),

and Gessaman
and energy
useful

predictor

monary
ture

expenditure,

Ramanathan
metering

Energy

is more

rate

Expenditure

rate

for large

mammals.

Pul-

related

to energy

expendi-

closely

rate

or heart

rate

(Datta and
for tele-

is not available.

of Wild Ungulates

metabolic

of life-sustaining

rate

is the energy

activities

as the sum of basal

rate

post-absorptive

ment) and increments

for functions

cost to a free-ranging

and processes

can be estimated
of a resting,

Hargrove

rate was not a

1969, Sharkey et al , 1966), but methodology

this parameter

less

on respiratory

that respiratory

respiratory

appear

expenditure.

literature

expenditure

than either

Ecological
animal

rate

of energy

reviewing

concluded

of energy

ventilation

in humans

after

ventilation

metabolic

animal

(Moen 1973).
rate

(the metabolic

in a thermoneutral

such as digestion,

It

environ-

�-65-

thermoregulation

and activity.

have been measured

Energy

for a number

costs

of wild ungulate

measurements

include

fasting

post-absorptive

animal

in a thermoneutral

metabolic

(that of a fed animal

effects

rate

of fluctuations

of fasting

amount of energy
retention

been studied
Fasting
(Silver

required

equals

zero).

less

ured

species.

Types

(the metabolic

environment),

temperature

to carry

components

in a thermoneutral

rate

of a

resting

on the metabolic
energy

of

environm.ent),
rate

requirements

out body functions

(the

when energy

cost of activity

in wild ungulates

has

rate has been measured

for white-tailed

deer

frequently.

metabolic

et ale 1969, Thompson

1970), red deer

Cervus

elaphus

1968, Maloiy et ale 1968), and pronghorn
(Wesley

rate

and maintenance

Energy

et a l . 1959, Silver

bou (McEwan

metabolic

in ambient

and fed anim.als,

of these

et a l , 1970,

for black-tailed

1973).

Resting

deer

Odocoileus

et a l , 1973), cari-

(Brockway

Antilocapra

metabolic

and Maloiy

am.ericana

rate has been meas-

hem.ionus columbianus

(Nordan

et a l , 1970), caribou

(McEwan

1970, McEwan and Whitehead

red deer

(Brockway

and Maloiy

1968, Maloiy et al , 1968), pronghorn

(Wesley

et a.l, 1973), wildebeest

1966,

1968), and eland Taurotragus

Rogerson
effects

1968).

deer,

by Hart

taurinus

oryx (Taylor

Siver et ale (1971) and Holter

of am.bient tem.perature

white-tailed
caribou

Connochaetes

on m.etabolic

(Rogerson

and Lym.an 1967,
et ale (1975) studied

rate

of fasting

while the sam.e problem. was investigated

et aI , (1961),

for pronghorn

1970),

by Wesley

and fed
for

et ale (1973),

�-66-

and for mountain
(1954).

Maintenance

long term
(Ullrey
al.

energy

for young caribou

tailed

were

of bedding,

the energy

deer

and pulling

Wesley
horns

standing,

made by Mattfeld

et al.

energy

for pregnant

(McEwan and Whitehead

while Hammel

measured

studies

have been estimated

1970), for white-tailed

et ale (1961) measured

standing

requirements

by Krog and Monson

expenditure

in a respiration

the energy

chamber.

(1978).

of

whiteHart

caribou,

expenditure

of rein-

and White and Yousef (1974)

of standing

et ale (1973) estimated

Measurements

of standing

et

1976), and

walking and running

expenditure

sleds,

deer

fawns (Baker

1970).

using

fawns (Thompson

(1974) and by Fair

(1962) measured
loaded

white-tailed

deer

et ale 1977), for mule deer

expenditure
deer

energy

balance

et ale 1969,

1973, Holter

energy

goats Oreamnos americanus

the energy

and walking reindeer.
cost of standing

for prong-

�-67-

CHAPTER

II

METHODS AND MATERIALS

Based on information
energy

expenditure

respiratory
feeding,

by indirect

collection.

jects,

accurate

serve

as a standard

entry

rate

necessitated

for evaluating

heart

the carbon

the advantage

and indirect

using

could be expected,

I rejected

of less

and indirect

a face mask for

more

results.

restricted

some restraint

rate

as a predictor

entry

rate

restraint.

required

calorimetry

to measure

and the method

dioxide

calorimetry

and apparatus,
accurate

calorimetry

apparatus

results

though it offered

I decided

Although use of the face mask

and associated

expenditure.

in the literature

of sub-

would
of energy

technique

Both carbon

aldioxide

sophisticated

training

could be expected

to give

Animals
Subjects
orphans

were

turned

obtained

either

over to the Division

as fawns from
of Wildlife.

left with does for approximately

24 hours

reared

by Reichert

using

methods

of fawns at weaning,
peak weights

described
weights

at start

captive

Newborn

and then were
(1972).

deer

fawns were

removed

and

Ages and weights

and end of the experiment,

of fawns which were fasted

or as

are given in Table

1.

and

�Table

1.

Sexes, ages (days), and weights (kg) of experimental
ment, and peak weights of fawns which were fasted.

subjects

at times

of weaning and experi-

Experiment
Weaning
Deer

Start

Sex
Age

Weight
Age

Weight

Peak
Weight

Final

co

Weight

d'

6-1-76*

81*

16.2

217*

44.3

47.2

44.8

51

d'

6-5-76

77

11.3

213

38.0

--

42.4

53

59

9.9

187

29.4

30.4

&lt;;&gt;

7-1-76

31.5

55

* Orphan fawn, birth date and age estimated.

I
0'
I

�-69-

Fawn training
trained

to wear

scribed

by Reichert

styrene

plastic

elastic

straps

more

restrictive

began at the age of 10 days.

a harness

and accept

(1972).

bottle

on their

feeding.

and heavier

by leaving

each day in conjunction

with harness

month,

replaced

cups were

with nylon webbing.

weeks until the fawns accepted

face cuff and outlet
valve inserted,
throughout
Four

ready

fawns were trained

each day for 90 days.

From

At this time

and later

factorily.

However

one fawn would not accept

the deer

from the study.

of 8

first

with the

were used

for

10 to 30 minutes
trained

three

the mask and restraint

the study when the deer

was dropped

cups held on

Raisins

that time on they were

required

a

with the cuff tightened,

in this manner

All fawns accepted

Since hooking up the apparatus

closed

and as a distraction.

a week.

who joined

made

the fawns weighed

mask,

hose attached.

four times

nician

by

After

were added over a period

the study for reinforcement

poly-

end partially

plastic

to wear the actual

and collection

open-ended

and leash training.

400 g.

valve removed,

as de-

on the fawns for 20 minutes

with sturdier

Again weights

about 20 kg and were

a leash

initially

the cups were

the distal

Cups were placed

were

muzz les and attached

Gradually

and taping on weights.

these

from

At 2 to 3 weeks of age,

cups were placed
during

restraint

Fawns

were

handling

from

or

satisa tech-

150 days of age.

the technician's

asssistance,

�-70-

Of the three
fitted

readily.

deer used for trials,

The female

(deer

by being coaxed or tricked.
necessary,
straint.

no. 55) accepted

Although several

it was always possible
This deer

the two males

the face mask only

attempts

were often

to get the mask on her without re-

was about 30 days younger than the others.

was initially

kept at the home of a researcher,

days of age,

and was hurried

tion,

could be out-

through

She

began training

the training

process.

at 21
In addi-

she was the most active,

independent,

and inquisitive

of the

At certain

were excited

and showed a high level

deer.

of activity.

times

all deer

I did not conduct trials

were difficult

to handle

and restrain.

standing,

or quietly

feeding,

and trials

completed

with minimal

Except
diet of alfalfa
forage

ration.

Physical

during

fasting

Supplemental

times

because

the deer

If the deer were bedded,

outfitting

could be readily

accomplished

restraint.

periods

hay and a commercially
sodium

deer

were fed ad libitum

available
chloride

mixed grain

on a
and

was also provided.

Facilities

The experiment
1977, at the Colorado

was conducted

during

Division

of Wildlife
Elevation

pus,

Colorado

State University.

m.

The deer

were housed

square.

at these

outdoors

January

Deer

Pens,

and February
Foothills

was approximately

in two group pens roughly

Cam1,585
30 rn

�-71-

Apparatus
Heart

Rate Apparatus
Radiotelemetry

trials.

Transmitters

Colorado

(Krohn-hite),

I tried
tained

an FM receiver

several

different

during

Laboratory

on a harness
A television

(Eddystone),

sites

by attaching

a band -pass

filter

recorder

(MFE

electrodes

and ob-

chart

for attaching

one electrode

to the right

of the

and the second on the left side near the lower posterior
Electrode

electrode attachment,

and alligator

hide of the fawns.
fawns'

hides

tures

no longer

for electrode

nal was obtained
in contact

Initially,

signals

detected

and reattached.

trodes

attached

in this manner.

were problems

from active

deer.

steel

sutures

directly

sec-

of
to the

were made in the

After about a week the su-

adequately,
alone,

but a satisfactory
provided

sig-

they remained

clips which pulled off were
Deer appeared

associated
Clear

at the sites

clips were attached

clips

Alligator

readily

There

was applied

stainless

with alligator

with the skin.

paste

attachment.

conducted

at

system.

tion of the rib cage.

signals

by deer

and a strip

the telemetry

best results

sternum

and were carried

a signal processor,

completed

by the Surgical

webbing and spandex bandages.

(Winegard),

Corp.)

were constructed

State University

made of upholstery
antenna

was used to obtain electrocardiographs

signals

oblivious

with obtaining

to elec-

satisfactory

were obtained

for bedding,

�-72-

standing,

and usually

accompanied
sociated

for walking,

by deterioration

with movement

to one another

cation

from R. J. Scott,

Poor

signals

by adjusting
signals

rate

of the signal.

of the lead wires

spect

rather

obtained

Surgical

the alligator

rates

for trials

15 seconds
artifact

periods

and counts

to a IS-second
On several

At times

were impossible;

Samples

the tracing
however.

the strip-chart

Beats

of 15 seconds

by

could be

they were converted

recorder

from the receiver

was nonwas used to take

were counted from the audio signal for

during

as many minutes

the same way as samples

as a supplement
samples

were

for the first

if counts

as samples.

occasions

heart

was obliterated

and included

were treated

cardiograph

the trial.

equivalent

samples.

ings and used

to obtain better

by sampling

period

heart

samples

efforts

of the sample

and the audio signal

any period

be improved

or more

functional,
rate

could usually

were calculated

throughout

with re-

(oral communi-

R waves from the electrocardiograph

of each minute.

made for a third

levels

as-

State Univ.).

failed.

Mean heart

was usually

Colorado

but persistent

levels

taken by counting

artifact

clips,

activity

I-minute

and transmitter

Laboratory,

at low activity

activity

This was probably

than with muscle

at higher

during

but increased

to them.

were available

If less

for a trial,

as possible.

Audio

from the EKG tracthan four electroand a greater

�-73-

number

of audio samples

ples.

In trials

with four or more

used audio samples
were

illegible

minutes

Face
cated

(if they were available)

for the minutes
Trials

mask-A

fiberglass

for use on deer.

with fewer than four

face mask was designed

mache antelope

with modelling

necessary

to provide

clay.

of valves

to the right

muzzle

E.

(Warren

E.

Collins

Collins)

let valve to be removed

for training.

entirely

is shown in Figure
circular

to a section

The mask with outlet

1.
face cuff was made by cementing

tire

tube (2.54 cm diameter)

bicycle

tire

valve.

and installing

a Presta

to the inside

of the mask as a seal to fit around the muzzle
through

of a

allowed the out-

of bicycle

The valve protruded

The in-

The outlet valve was con-

This arrangement

ject.

clearance,

"V" valve).

PVC valve glued onto the mask.

an inflatable

on a

Two holes were cut in the fiberglass

to the mask.

connector

the end s of a section

and fabri-

Clay was added to the head where

(Warren

let valve was glued directly
nected by a rubber

head fashioned

space for valve functioning,

of a face seal.

for insertion

Initially

which

The shell of the mask was constructed

dimensions

valve inserted

I

and Gas Analysis

form made from a paper

shell

samples.

using a single method were discarded.

Apparatus

and installation

I used only the audio sam-

electrocardiograph

on the electrocardiograph.

sampled

Respiration

were available,

together

This cuff was cemented
of the sub-

a hole cut in the fiberglass

so that

�-74-

,~

FACE

.•

Figure

1.

'/&gt;

10.

II

11

I~

Face mask for measuring energy expenditure of mule
deer fawns.
Ma s k is mounted on a plaster muzzle
form and viewed from above.

�-75-

the cuff could be inflated

with the hose and bulb from a sphygmo-

manometer.

even when inflated,

However,

form to the deer's
was modified

muzzle

and allowed passage

before

data were

collected.

of 1.27 cm thickness

neoprene

rubber

fill the space

the deer's

between

7 c rn cross-section

motocycle

When the mask was on the

to the head by three

Straps

were fastened

around

and unfastened

3 brackets

by spring

on the mask.

clamps

to allow rapid

for leaks by applying

a soap solution

and adjustments.

The face seal was checked
to the muzzle

of one fawn,

with the respiration

from the mask were detected.

to the face seal was checked
age such as moisture
were detected

during

intact.

of the muzzle
trials

adjacent

for signs of leak-

at the corner

by wetness

No

of the mouth

of the hair

and by

such leaks to the mask not being put on tightly

enough and discontinued
Flowmeter

Leaks

occasions

apparatus

The area

routinely

or frost.

on several

I attributed

meter

tube was put

The mask was secured

of 2.54 cm webbing riveted

ration

A

down over the deer's

straps

sound.

tire

shell.

of the tube was turned

to form

leaks

in the mask to

muzzle" and the fiberglass

muzzle

attachment

a seal.

so this design

sponge installed

of 10.8 ern diameter

the folded section

of air,

to con-

The final seal was a ring

over the mask and folded back on itself.
deer,

the cuff failed

the trial

and collection

(Instrumentation

when this occurred.
bags-I

Associates)

carried

a Max Planck

to measure

respi-

and sample

�-76-

expired

air (Figure

2.68 cm. plastic
face mask.
through

2).

The meter

tubing (Warren

E. Collins)

The m.eter delivered

a 30 cm section

polyethylene

was connected

by a section

to the outlet valve of the

a 0.6 percent

aliquot of expired

of 4.8 m.m. ID Tygon tubing

bag for storage.

Metallized

of

to a metallized

bags were m.ade as de-

scribed

by Johnson

et al , (1967) in a 25 x 35 cm. size.

several

metallized

bags in a pack sack on my back to allow rapid

switching

of bags between

Flowmeter

meter

the two meters

at varying

periment

was checked by connecting

(American

rates

ranged

ture

and pressure

saturated

from

10.2 to 46.2

liters

Initial

and final calibration
factor

factors
derived

Metallized

gas bags were

mersing

them. in water

nections

between

through

This was done before
were completed.

the exFlow

per m.inute at body tempera-

(BTPS) for the initial

per minute

it in series

Co.) and breathing

all trials

from 4. 1 to 103.2 liters

obtain the volum.e m.easurement

soap solution.

Meter

flow rates.

began and again after

I used the calibration

I carried

trials.

calibration

with a wet test

air

calibration,

and

BTPS for the final calibration.
were

1. Oland

0.99

from the initial

respectively.

calibration

to

for all trials.
routinely

and applying

checked

for leaks by im-

light pressure.

the face mask and flowmeter

were

Hoses

and con-

checked

with a

�-77-

Figure

2.

Measuring energy expenditure
of mule deer fawn.

and heart

rate

�-78-

Gas analysis

-Oxygen

man paramagnetic
(Servomax).

oxygen analyzer

Endpoints

as the lower point,
percent

carbon

mainder

nitrogen)

fore and after
(determined
standards

Trial

analysis

were

by running
were

and,

recorder

with 100 percent

nitrogen

oxygen,

trace

argon,

These

standards

several

times

1.21
re-

were run be-

if the instrument

the standards

run more

chart

methane,

point.

samples,

using a Beck-

gas (19.64 percent

O. 11 percent

as the upper

running

wired to a strip

established

and a standard

dioxide,

was performed

was unstable

initially),

the

frequently.

Procedures
To prepare

fawn,

for a trial,

the experimenter

stand the fawn if necessary,

An assistant

would place

the electrodes
leading

harness

placed

over the heart

was fed raisins

rate harness.

After

neck by the back straps,

muzzle

while the strap

around

The forehead

strap

that the mask

was secure,

and hook up

a distraction.

loosely

and then quickly

A

1972) was then

a satisfactory

the face mask was placed

deer's

Deer

as

a

rate harness.

in the harness

of 2.54 cm nylon webbing (Reichert

had been obtained,

muzzle.

and put on the heart

the transmitter

while the deer

would approach

EKG signal

around

slipped

the

over the

the back of the head was tightened.

was then fastened,

and after

a check to ensure

the face seal was unfolded

were not restrained

at any time during

down over the
this procedure.

�-79-

Next,

a 4 m rope leash

the subject

was restrained

be threaded

through

outlet

was attached

slightly

so that the collection

loops on the leading harness

valve of the face ma sk (Figure

into a separate

yard where trials

from the gas collection

system.

and by the assistant

watch,

switching

human

activity,

gas collection

throughout

the EKG signal

at the beginning

provide
were

bedding trials

an adequate

shorter

sample,

from

of the deer,

were

recorded

The assistant

of the trial

and several

checked

where

needed

were

up to 30 minutes

on

usually

and helped

Most of the trials

the stop

and opening the

Activities

in the area

lasted

Consolazio

10 minutes

in order

of the mo st active

to

trials

than 10 minutes.

Following
aliquot

bag.

air

in my

was begun by starting

the trial.

for the r erna ind e r of the trial.

was led

and data were

carried

and flowmeter,

and any disturbances

long; however,

remarks

tape recorder

The trial

the tape recorder

to the

was allowed to flush outside

on the transmitter

valve to the metallized

and attached

on a data fo r rn adapted

et ale (I 963), (Appendix A).

hose could

Then the subject

Introductory

at this time on a casette

pocket,

2).

and

were conducted.

A rrrirrirnurn time of 2 minutes

recorded

to the leading harness

a trial,

connection

the meter

was switched

backpack.

Subsequent

to flush.

If all

trials

reading

was recorded,

and the

to another

gas collection

bag in the

were begun after

equ ipme nt continued

allowing

to function

the systetn

properly,

up to

�-80-

four

trials

were

run

In sequence

without removing

apparatus

from

was used to determine

tem-

the deer.

Weather

Parameters

A thermometer
perature

on the flowmeter

of expired

temperature

air at time of volume

was measured

measurement.

using a thermometer

side of a building

about 1.5 m above the ground.

tures

-4.5

varied

from

I obtained

barometric

microbarograph

(Julian

Friez

P,

State Univer s ity Atmospheric

pressure

elevation,

on the Colorado

State University

converted

a standard

correction

readings

Calculation

of Energy

Expenditure

were

quires

knowledge

of oxygen consumption

(RQ).

RQ is the ratio
under

type of nutrient

normal

expenditure

conditions

being metabolized

Zuntz and Schumberg

1901).

instrument

readings

for elevation

made at one-half

of energy

It varies

from a

at the Colorado

Campus

factor

Calculation

of carbon

readings

from another

Campus.

pressure

at the warmer

Building about 0.5 km distant

or occasionally

Barometric

on the north

Ambient tempera-

and Sons) located

Science

and at a similar

using

located

to 16.0 C, with most trials

temperatures.

Ambient

were

change.

hour intervals.

from oxygen consumption
and the respiratory

dioxide produced

quotient

to oxygen consumed.

from O. 7 to 1.0 depending
(Brody

re-

1945, citing

Since I did not measure

on the

Lusk 1928 and
,
the carbon

�-81-

dioxide

concentration

air I assumed

of expired

animals,

as suggested

animals.

Each liter

o. 70 for fasted

by Brody (1945), and an RQ of
of oxygen consumed

4.686 kcal of heat at an RQ of 0.82,

results

in the production

Oxygen consumption
ments:

the volume

is strictly
carbon

correct

dioxide

methane

production

to standard

RQ's for fed and fasting
zero.

Derivation

with a fed deer
expenditure

animals,

are included

resulting

error

and pressure

assuming

volumes

dry,

deer,

in the assumption

(both

or STPD) are

by using the assumed

methane

B.

from this calculation
for fasted

are equal (and

production

and a s arnpl e calculation

in Appendix

This

it is only when

and expired

volume to inspired

of the formula

and -7 percent

considering

that inspired

temperature

I co r r e ct ed expired

equal.

air and its oxygen concentration.

and oxygen consumption

is zero)

1901).

from only two measure-

only at an RQ equal to 1. 0, because

production

converted

was calculated

of expired

of

or 5.047 kcal at an RQ of 0.70

1945, citing Lusk 1928 and Zuntz and Schumberg

(Brody

deer

an RQ of 0.82 for fed

Correction

for a trial
in energy

was -4 percent

but was probably
that methane

was

for fed

not significant

production

was

zero.
E nergy

expe nditure

pressing

energy

weight)

reduces

was expresse

expenditure
variation

. ca 1k g -0. 75 mrn
. -1
d In

on this basis

related

(in units

to size differences

of metabolic

Exbody

among animals.

�-82-

Activity

Data

Sporadic
changes

activities

in activity

recordings
Percent

of the deer

for later

sampling.

by timing activit ies during

of time

made it necessary

spent at bedding,

Sampling

the first

standing,

this sampling

scheme

of sampling

for 16 trials

with results

obtained

trial.

Sample percentages

determined

by complete

variation

for an activity

Control

by comparing
by timing

were within

for all but three

by the two methods

plans

activity

became

called

levels

apparent

tent required

results

activities

10 percent

trials.

for

of values

Max irnurn

was 14 percent.

for three

replicates

on each deer.

of one trial

As the experiment

that it was not possible

to control

to achieve

this design.

Because

to lead and would accept

only limited

restraint,

consisted

of a mixture

of standing

through

the experimental

period,

and walking.

Measurements

could be made by attaching

using care

made at higher

not to arouse

activity

were unaccustomed.

levels

the subject.

progressed

activity

deer

it

to the ex-

were not trained

most early

trials

About half-way

deer

rate

equip-

would often bed.

the collection

hose at this

Measurements

using a second assistant

Male deer

at each of

I found that if the heart

ment and face mask were left on about an hour,

time,

were

of Activity

Initial
three

timing

minute.

walking and trotting

I evalutated

all

was done from tape

15 sec of every

calculated.

the entire

to record

were

to whom deer

would follow this person

around

�-83-

trying

to rub their

them to maintain
discouraged

heads

on her.

By walking rapidly

a fast walk with some trotting.

because

heart

rate

signals

and I could not keep up.

ture,

and activity

sented

heart

rate

in Table 2.

ing, walking and trotting.
trial.

Only two bedding trials

single

activity.

All other

trials

equipment
expendi-

for each deer

are pre-

included

did not exceed

for deer

were

in energy

which occurred

Trotting

paces

obscure,

Ranges

measurements

Activiti.es

Faster

became

often detached,

she could induce

bedding,

10 percent

stand-

of any

no. 55 were made up of a

were composites

of two or more

of

the four activities.
To determine
expenditure

if the relationship

was the same over the entire

data into four activity
ages of time
were

between heart

levels

on the basis

spent walking and trotting.

less than 45, the trial

percent-

If the combined

percentages

were between 45 and 70, the trial

the combined

percentages

time

Finally

spent bedding:

bedding,

the trial

went into Level 2.
levels

Levels

if greater

A diagram

distribution

3.

or 2; if the combined
went into Level

than 70. the trial
split on the basis

than 10 percent

went into Levell;

is given in Figure

well-balanced

1 and 2 were

went into
of percent

if less than 10 percent,

This breakdown

3; if

of

of the time was spent

of the breakdown

of trials

I divided

of the combined

percentages

Level 4.

and energy

rar.ge of activity,

went into Levell

were greater

rate

scheme

resulted

for all subjects.

the trial

for activity

in a fairly
Analysis

of

�-84-

Table

2.

Range of energy expenditure,
heart rate, and activity
measured
for three mule deer fawns during January and
February.
Deer

Number

Parameter

cr

51 if

53

Energy expenditure
(kcal kg-O. 75 min-l)

.073-.214

.076-.306

.066-.235

Heart rate
(beats min -1)

54.3-81.6

48.7-92.7

54.5-96.0

28-98

20-99

21-100

Time spent bedding
and standing
(percent)

55 ~

�-85-

All trials

Less than
45

Percent
walking/
trotting:

Greater
than 10
Levell

Percent
bedding:

Figure

3.

45-70
Level 3

Greater than
or equal to 70
Level 4

Less than
or equal
to 10
Level 2

Schematic breakdown of trials into four activity levels
the basis of percent of time spent at specific activities.
Levels are numbered in order of increasing
activity.

on

�-86-

variance

for heart

in mean heart
jects.

rate

rates

showed that there

at different

activity

suggests

that activity

This evidence

used to test the effect of activity
rate

and energy

expenditure.

scribed

below.

Trials

on Fasted

deer.

activity

Analysis

Individual

Deer

Regression
squares

penditure
able.

repetition

can be

between heart

for this effect are de-

in obtaining

with fasted

analysis

of the study with
trials

animals

using deer

at different

until the end of

no.

51 and six using

48 or more hours

of fasting.

of factors

level and feed level)

are given in Table

Statistical

least

required

Five measurements

at each combination

for each deer

tests

trials

no. 55 were made after

of trials

level grouping

on the relationship

of difficulties

I delayed

the experiment.
deer

differences

for each of the sub:'

Statistical

design

Because

levels,

levels

significant

Deer

The experimental
fasted

were

(activity

Numbers

3.

was done on the data for each deer using

analysis

(Draper

and Smith 1966), with energy

as dependent

variable

and heart

To evaluate

effects

tionship

between heart

variance

was used.

rate

of levels

of activity

and energy

Analysis

rate

as independent

vari-

and feed on the rela-

expenditure,

of covariance

ex-

analysis

for comparing

of co-

regression

�-87-

Table

Deer

3.

Number of simultaneous
energy expenditure
and heart rate
measurements
at different levels of activity and feed for
three mule deer fawns in winter,
and comparisons
for effects of activity level and feed level.
(See Figure 3 for
activity level descriptions.)

Feed

51

53

55

Level
.•..
-r-

Totals

1

2

3

4

None

Fast

3

a
3

0

0

0

6

Fed

0

19

6

1

2

28

Totals

3

22

6

1

2

34

Fast

0

0

0

0

0

0

Fed

4

d
12

13

Sd

1

35

Totals

4

12

13

5

1

35

Fast

2

2

0

0

1

Sf

Fed

Se

8

17

2

1

3l

Totals

7

10

17

2

2

38

.•..
..•.
Activity
a, b, fC
c, d,e

Activity
Level

ac

d

not measured

c

d

e

e

or did not fit into any level.

.
o mpa r i s ons for effect

.
Co mpa r i s on s for effect

of activity

level.

of feed level.

b

b

�-88-

lines

is described

by Snedecor

Used in this way,
cepts

and Cochran

it is a procedure

for comparing

of reg res sion lines obtained

in some attribute.
different

from groups

I used the procedure

combinations

of activity

(l967) and Li (1964).
slopes

and inter-

of data which differ

to compare

regressions

and feed level for individual

at
deer

(cells).
Since sample
cific comparisons
pare

deer

activity

were small

could be made,

the relationship

different
levels

sizes

as illustrated

between heart

levels,

in some cells,

rate

regressions

no.

53 at levels

and trials

on deer

in Table

and energy

from trials

Fed 2 and Fed 3 were compared;

only a few spe-

on deer

regressions

no. 55 at levels

Fed 1, Fed 2, and Fed 3 were

expenditure

at different

levels

of feed,

on deer

no. 51 were compared;

and all fed trials

and fed trials

for fasted

on

Fed 1, Fed 2, Fed 3 and Fed 4 were compared;

the relationship

gressions

at

no. 51 at

from trials

To compare

for all fasted

To com-

expenditure

compared.

trials

3.

on deer

between heart
regressions

rate

and energy

from all fasted
regressions

no. 55 were compared;

and re-

and fed trials

on deer

level 2

for analysis

of covariance

no. 51 at activity

were compared.
An assumption
residuals

for the different

sumption'

I used Bartlett's

binations

of activity

populations

are equal.

test of homogeneity,

is that mean square
To test this astesting

level and feed level with the largest

the four comsample

size.

�-89-

All Deer
Analysis

of covariance

was conducted

differences

and heart

rate

the relationship

between

deer.

test was done to test the assumption

Bartlett's

energy

to examine

expenditure

In

among

of homogeneity

I

i

of inean square

residuals

for data from the three

Two characteristics

which differed

among individuals

body weight and sex (see Table

1).

nos.

characteristics.

51 and 53 in both of these

tionship

between heart

individuals
ships,

rate

Deer no.

and energy

would be of greater

55 differed

and h ea r t rate

expenditure

As mentioned

earlier,

when energy

the basis

of metabolic

body size,

different

individuals

are corrected.

species

of different

the exact

body weight,

relationship.

vary directly

and Teissier
there

is an inherent

length of the muscle.
versely

proportional

rate

to many

relationenergy

ex-

further.
expenditure

Heart
but there

rate

IS expressed

also varies

on

that heart

the problem

citing

which varies

that heart

to the length of the heart

rate

should

Lambert

from the standpoint

contraction

They suggested

regarding

or inver sely with

and Rodahl (1970),

of muscle

among

is disagreement

per unit weight,

o
Astrand

approached

rela-

d iffe r enc e s due to body weight of

with heat production

(1927),

applicable

between

Brody (1945) suggested

the cube root of weight.

from deer

use than many individual

was investigated

were

Since a single

effect of body weight on the relationship

penditure

deer.

rate

muscle,

that

with the

should be inthat is,

to the

�-90-

one -third

power

of body weight.

lected

by Rihl (1927),

ferent

size was inversely

Kleiber

found that heart

rate

proportional

weight,

a r e lat ion sh ip theoretically

tissues

(hence basal

fourths

power

considerations,

between

energy

possible

power of body

to the three-

Correlations

were determined

expenditure

per unit of metabolic

ations

of heart

which adjusted

ships .. The transformations

body weight raised

1) heart

by body

power,

Energy

mean energy

energy

expenditure

on heart

sions

for grouped

individuals

30 kg).

multiplied
rate

multiplied

by

Expenditure
were calculated

and plotted

from the regression

for all deer,

(males

A formula

and Smith (1966).

intervals

expenditures
rate

rate

and 2) heart

for Predicting
confidence

between

relation-

power.

The 95 percent

in the

body size and two transform-

to the one-fourth

Intervals

rate

for each of the proposed

were:

to the one -third

of body size on

and heart

energy

rate

effects

expenditure

examined.

Draper

of dif-

with blood flow to the

being proportional

study were

approximately

species

to the one-fourth

present

for predicting

data col-

of body weight.

the relationship

Confidence

examining

of several

consistent

metabolism)

Based on these

weight raised

(l961),

of

and from the regres-

approximately

40 kg and female

for this calculation

is given by

�-91-

CHAPTER III

RESULTS

Effectiveness

of Method

Behavior

of the deer

was affected

in several

ways.

Standing the deer

increased

activity

and energy

ing procedures
procedures
logical

altered

began,

factors

deer

rate harness

subsequent

The consistency
behavior.

altered

their

Once harnessing

and physical

energy

of train-

and psycho-

expenditure.

After

they would pace by the gate of the pen until taken out.

no. 55 was more

active

than the others

while pacing

creased

could be minimized

activity

least

to put on the heart

expenditure.

head movements

were

procedures

knew the routine,

probably

being outfitted
Deer

their

by experimental

active,

the fences

and would often make

during

trials.

by working

or by waiting until one-half

Such in-

with deer
hour after

when they
outfitting

to

begin trials.
A second
ing trials
spaces

alteration

subjects

or climbing

they accepted
emotional

resulted

often had to be restrained
on objects

this restraint

factors

of behavior

from restraint.
from

in the experimental

without physical

may have affected

exploring
area.

struggle,

measurements.

Durnarrow

Although
associated

�-92-

With the methodology
avoid.

Fear

rare

occasions

frightened
mental

such effects

was an emotion that appeared
when equipment

the deer,

procedures

Problems

used,

to be avoided.

was dropped

no fright behavior

were impossible

Except

in connection

with the experi-

was observed.

with Respiration

Equipment

with respiration

equipment

often resulted

cessful

trials.

The most serious

of these

was an inoperative

valve,

possibly

caused

structure.

by the rubber

This occurred

53.

An audible wheezing

later

by the deer

the deer

mask before

struggling

peared

to breathe

secreted

large

mouth.

trotted;

at other times

due to incorrect

about one minute

later

saliva

trials

deer

On
the

no. 53 ap-

each day and

when the mask was on.
at the corners

of the

hose would often detach from the flow meter

pull the hose off the face mask.
tained,

no.

for mask removal.

Unlike the others,

of mucous

with deer

in the

the sound and removed

failed due to mask leaks

The collection

when deer

I recognized

began.

inlet

Twice when this happened

and was restrained

more deeply during

amounts

Some trials

four to six times

not being able to inhale.

occurrences

in unsuc-

disk of the value catching

sound was accompanied

began to struggle

the remaining

on

or a sudden movement

Problems

support

to

extensive

rubbing by the deer

For many trials

adjustment

would

no aliquot was ob-

of the flow meter.

Finally,

the

�-93-

oxygen analyzer
creasing

would sometimes

precision

Relationship
Individual

Energy

Expenditure

was a high correlation

rate

0.92

and 0.92

for all three

of energy

deer

between

thus de-

Rate

The correlation

are plotted

in Figures

on heart

levels

by activity

were 0.81,

4, 5 and 6.

for the three

Regres-

deer

in slopes

for any deer

(p = 0.86,

deer

51, 53 and 55 r e spe ct ive lyl ,
in intercepts

at different

53 and 55 (p = 0.21,

0.19

nificant

51 (p = 0.05).

no.

over all activity

slopes

of regression

There

activity

respectively),

of covariance

deer

level for trials

differences

activity

Analysis

coefficients

and

are

7.

of covariance

for deer

expenditure

51, 53 and 55 respectively.

rate

at different

differences

energy

nos.

showed no significant

nos.

and Heart

from deer

expenditure

in Figure
Analysis

deer.

for trials

Data for the three

deer

analysis,

Deer

heart

plotted

during

of analysis.

between

There

sions

drift

levels
lines

of regression
0.42,

levels

showed no significant

was no significant

0.47

for deer

but the difference

nos.

lines
for

were no significant

by feed level for trials

for deer

of individual

nos.

was sig-

of individual
differences

in

51 or 55 (p = O. 19 and 0.98

respectively).

There

difference

the regressions

for deer

no. 55 (p =: 0.69),

51 (p = 0'102).

Analysis

of covariance

but there

in intercept

for

was for deer

by feed level for trials

no.

of deer

�-94-

350

z
~

•...

I

It)

o 250
(!)

~

"-

&lt;i

o

3

x

:5

W200
~
~

x

(5

z

W

Q.

X

w 150

&gt;

(!)

a::

w
z
w

2
A
A

50L---L---L---L- __~ __~ __~ __~ __~ __~ __~ __ -L
40

50

60

70

80

90

_

100

HEART RATE (BEATS/MIN)
n= 34, r = 0.81
2/

Fed, activity level 2 (less than 45% walking and trotting,
than 10% bedding), n = 19.

3/

Fed, activity
Fed, activity
n = 1.

4/
A/
B/
X/

Figure

4.

level 3 (45-70%
level 4 (greater

less

walking and trotting),
n = 6.
than 70% walking and trotting),

=

Fasted,
aCtivity level 1 (greater
than 10% bedding), n
3.
Fasted,
activity level 2 (less than 45% walking and trotting,
than 10% bedding), n = 3.
Fed, no activity measurement,
n = 2.

Relationship
of energy expenditure
by activity level and feed level.

and heart

rate

for deer

less

no.

51

�-95-

350

z
~
I

m
•....
0 lZ50
C)
:tC

"..J

4

&lt;tlt

3

(.)

3

r

234 3

0

z
lLJ

2

I
2

~

2

Q..

X
lLJ

343X
3
4

3

150

&gt;-

(!)

a:

3
3

12

lLJ

I

z

2

I.U

'00

2

50L---~---L---L--~--~~
40
50
60

__~ __~ __~ __-L__~
70

~

90

80

_

100

HEART RATE (BEATS/MIN)
n

= 35, r = 0.92

11 Fed, activity level 1 (less than 45% walking and trotting,
greater
than 1010 bedding), n = 4.
21 Fed, activity level 2 (less than 45% walking and trotting, less
than 10% bedd ing), n = 12.
31 Fed, activity level 3 (45-7010 walking and trotting), n = 13.
41 Fed, activity level 4 (greater than 70% walking and trotting),
n = 5.
XI Fed, no activity measurement,
n = 1.

Figure

5.

Relationship
of energy expenditure
by activity level and feed level.

and heart

rate

for deer

no.

53

�-96-

350

z
~
I

It)

ci 250
(!)

~

"
«
o
.J

2
3

~200
:::&gt;
~

3

3

3

3
22

o
z

Ul
Q.

B~3

3

x

Ull50

3

&gt;(!)

a:

2

2

w
z
w

3

3

2
2

3

2
100

1,8,
I A

,

A
50L---L---L-~~~~~--~
40

__ ~ __ ~ __ ~ __ ~ __ ~

50

60

10

80

_

90

100

HEART RATE CBEATS/MIN)
n

= 38, r = 0.92

1 I Fed, activity level 1 (less than 45% walking
than 10 % bedding), n = 5.

21

Fed, activity level
than 10% bedding),

31 Fed,

41

activity
Fed, activity
n
2.

=

2 (less
n
8.

=

and trotting,

than 45% walking and trotting,

greater
less

level 3 (45-70 % walking and trotting),
n = 17.
level 4 (greater
than 70% walking and trotting),

AI Fasted,

BI

XI
YI

Figure

6.

activity level 1 (less than 45% walking and trotting,
greater
than 10% bedding), n = 2.
Fasted,
activity level 2 (less than 45% walking and trotting,
than 10% bedding), n = 2.
Fed, no activity measurement,
n
1.
Fasted,
doesn't fit in any activity level, n = 1.

less

=

Relationship
of energy expenditure
by activity level and feed level.

and heart

rate

for deer

no.

55

�-97-

350

3 ..··

300

DEER 5~.···
3

250
55

3

3
5
I
5
5 5

lJJ

0:

::&gt;

200

3

~

z

3

UJ

850

/

&gt;-

/

DEER 55

tr

./

I

100

5

//~f55
5

5

5
5

"Y

UJ

Z

;,

5 /

3

(!)

UJ

5 /5
5
5:.."

~/S

o
0..
X
UJ

5

/'

5

5

/

5~Y5
.-

3155,6
A
/

50~~---L--~--L-~~~--~--~--L-~---L--40

50

60

70

HEART RATE

Deer

51:

80

90

100

(BEATS/MIN)

n .:: 34, r:-:0.81

y = -102.09 + 4.09 (xl
Deer 53: n = 35, r = 0.92
" = - 130.44 + 4. 60 (x)
y
Deer 55: n = 38, r = 0.92
3. 91 (x)
Y = -151.59+
J\

Figure

7.

Regressions
of energy expenditure
on heart rate
three mule deer fawns in winter.
Data points
Iabe Ie Ll , 3, 5, are from deer nos. 51, 53 and
55, respectively.

for

�-98-

no.

51 at activity

level 2 indicated

or intercept,

but probability

0.10

respectively).

and 0.09

Mean square

square

residuals

differences

levels

residuals

of feed level and activity

no significant
were

necessary

level are

appeared

to significance

(p =

lines

at combinations

shown in Table 4.

Although mean

to increase

(p = o. 10).

for analysis

in slope

for the regression

as activity

for the four combinations

not significant

close

differences

tested

Therefore,

level increased,

using Bartlett's

I accepted

of covariance,

test

were

the assumption

that variance

was homogeneous

for aU ce Us.

AU Deer
The correlation
heart

rate

equation

coefficient

for 107 trials

obtained

ENEX equals

equals

heart

rate

on all three

by least

where

between

squares

was

energy

expenditure

was 0.47;

energy

was associated

ciated

analysis

2.53

(R2) for this equation
expenditure

was 0.69.

=

min -1.

The coefficient

min -1 and HR

of determination

thus about half of the variation
with heart

with individual

Bartlett's

significant

The regression

in cal kg -0.75

in the regression

did not reveal

differences

and

HR - 14.79,

Much of the variability
deer.

expenditure

deer

ENEX

in beats

energy

in

rate.
equation

was asso-

test for homogeneous

variance

among the mean square

residuals

�-99-

Table 4.

Mean square residuals
for regression
of energy expenditure on heart rate for three mule deer fawns at four activity
levels and two feed levels.
(See Table 3 for activity level
de s c r irrt i.on s ~)
Activity

Deer

51

Totals

Feed Level

Fasted

1

2

10.35

113.85

Fed

53

563.03

..•.
234.98 ....

209.65

*

499.66*

251. 77

11. 66

167.54

359.27

47.44

Fasted
Fed

19.46

* Mean square residuals
test.

4

3

Fasted
Fed

55

Level

*

296.34

591. 68

tested

for homogeneity

475.934

using Bartlett's

�-100-

of the regression
were

for individual

deer

(p = 0.75),

compared

using analysis

of covariance.

cant differences

among slopes

of the regression

deer

but there

(p = 0.30),

(p &lt; 0.0001).

The intercepts

fered

significantly

cepts

for deer

nos.

51 and 53, t=

from deer

were

nos.

-1.09

for

no.

about 30 kg).

energy

the one-third

to correct

in the relationship

no. 53).

55 dif-

The interno.

55 v s .

Onthisbasisdata

and separate

regressions

are plotted

of heart

in Figure

coefficient
rate

8.

between

and body weight to

as compared

to 0.69 between

energy

heart

The correlation

coeffi-

rate.

and the product

power

was 0.78.

for body size removed

between

no.

= 13.73 for

correlation

expenditure

body weight to the one-fourth
rate

for deer

51 and 53.

Regressions

was 0.81,

energy

in intercepts

51 and 53 (male's about 40 kg) and deer

nos.

and untransformed

cient between

nos.

51vs.

and the product

power

expenditure

differences

combined,

of body weight-The

expenditure

of heart

no.

51 and 53 were
for deer

Effect

of deer

were no signifi-

lines for the three

51 and 53 did not differ(t

were obtained
55 (female

There

of the regression

from those

nos.

significant

so the regressions

energy

expenditure

of heart

Thus,

rate

and

transformation

some of the variability
and heart

rate

among

the deer.

Confidence

Intervals

The 95 percent
energy

expenditure

for Predicting
confidence
from heart

Energy

interval
rate

Expenditure

for predicting

using the regression

the mean
determined

�-101-

350

-z
~

I

10

": 250
o(!)

~
"-.J
&lt;l

o

lJ.J 200

0::

::&gt;

~

o
Z
lJ.J

a..

x
lJ.J

---- --- ---

-

150

:&gt;
(!)
0::

ui
Z

//

l.!.J

/

~

//
//

--

--

/

/

/

/

/

V
/
50~--~--~--~--~~--~--~--~--~--~----~--~-100
90
80
50
60
10
40
HEART

Deer

Deer

51 and 53 (males,

55 (female,

30 kg):

RATE (BEATS/MIN)

40 kg):

=

=

n
69, r
0.89
1\
y = -125.33 + 4.50

(x)

n = 69, r = 0.92
Y -151. 59 + 3. 91 (x)

=

Figure

8.

Regressions
of energy expenditure
on heart rate and 95 percent
confidence intervals
for predicting
mean energy expenditure
from heart rate for two 40 kg male mule deer and one 30 kg
female mule deer in winter.

�-102-

f'rom all deer
tervals
basis

is plotted

9.

The 95 percent

confidence

in-

equations

for individuals

grouped

on the

in Figure

using the regression

8.

of sex and body weight are plotted

in Figure

Using the regression

the 95 percent

terval

for mean energy

for all deer,

expenditure

at the mean heart

confidence
rate

in-

(69.22

OO 75
. -1 ; at h eart
. -1). IS 154
b eats rrnn
.• 11 &lt; 1-1-&lt; 166.43 cal kgrrnn
-

of 60 and 80 beats

rates

x-

rrrin -1,

the confidence

are 87.42

intervals

&lt;

1-1~ 186.48 and 129.67 &lt; 1-1:s.2450 39 respectively.
The 95 percent

confidence

intervals

termined

from

grouped

of heart

rates.

Using the equation

confidence
rate

interval

(63.87

beats

60 and 80 beats

individuals

1

rnin -1,

and 153.88

min

-1

expenditure

cal kg

for the female,

vals at the mean heart

60 and 80 beats

for the two males,

the confidence

intervals

or decreases.

of

126. 19 &lt; 1-1:s.

-1 .
) and at heart

rates

150.68 &lt; 1-1-&lt;
164.09,
x-

creases

are

rate

inter-

are

rate

At heart

confidence

(78.94 beats rnin

the mean heart

the 95 percent

-0 75
1
.
min - respectively.

rate

row near

range

at the mean heart

the 95 percent

:s. 1-1s. 169.17 respectively.

de-

over a larger

is 157.43S.1-1~S.166.19.

162.67 and 160.31 &lt; 1-1
&lt; 308.39
Using the equation

are narrower

for mean energy
min- )

about the regressions

17.28 &lt; 1-1&lt; 149.21,

Confidence

but widen rapidly

of

intervals

as heart

rate

are narin-

�-103-

Deer

51, 53 and 55 (2 males,
n
107, r
0.69
v= -14.79 + 2.53

=

Figure

9.

40 kg and 1 female,

30 kg):

=

(x)

Regression
of energy expenditure
on heart rate and 95 percent confidence
interval
for predicting
mean energy expenditure
from heart rate based on data from three mule
deer fawns in winter.

�-104-

CHAPTER

IV

DISCUSSION

Comparisons

to Literature

To compare

measurements

trials

in which the subject

Trials

with more

as bedding trials;

ing were

categorized

category

was broken

fed trials,

into trial.s

six standing

Heart
mule deer
walking are

ering

trials

rates

trials;

trials,

rates

for specific

to minima

female

(pe r s , co mrn , ) monitored

with greater

as walking trials.

on fasting

deer

standthan

The bedding

and trials

on fed deer.

trails,

six bedding-

and eight walking trials.

5.

Jacobsen

activities,

in February.

in February.

and trials

deer for bedding,

shown in Table

activity.

than 80 percent

in this study and other

and white-tailed

I used

spent bedding were cate-

were four bedding-fasted

measured

a nine -month-old

trates

of time

with greater

as standing

there

with my data

spent most of the time at a single

walking were categorized

Using the scheme

in heart

in the literature

than 50 percent

gorized

70 percent

Values

studies

standing,

on penned

foraging,

and

(1973) found a seasonal

trend

with heart

Jacobsen's

rates

gradually

data in Table

5 are from

in February.

Freddy

white -tailed

deer

heart

of two adult mule deer

rates

low-

cas-

�Table

5,

Heart

rates

of mule

deer

and white-tailed

deer

Cor different

activities.

Activity
Source

Species
Bedding

Standing

Foraging

Walking

Deer 39
Trial
1

49

64

78

91

Deer 39
Trial 2

44

65

69

79

Deer 18
Trial
I

35

50

55

62

Mule deer
Freddy,
personal

2
comm,

Mule deer
Deer

2

3

This

61

51

51

study

4
72

Deer

53

Deer

55

565

Deer

55

596

•....•
Jacobsen

62

White-tailed
deer

52

57

White -tailed
deer

65

687

1913

o

\J1

I

IData

in files,

2Mean

for trials

pers,

788

comm.

from

D. Freddy,

oCa fasted

male;

percent

Colo.

Div , of Wildlife,

= 54 to 63,

bedding

3Mean

for trials

of a fed male;

percent

standing

= 81 to 86,

for trials

of a fed male;

percent

walking

= 73 to 77, remainder

5Mean

for trials

of a fasted

6Mean

for trials

of a fed female;

7perctlnt

standing&gt;

50.

8percent

walking&gt;

50.

percent

bedding

p e r c e nt bedding

remainder

= 63 to 81,

= BO to lOa,

Kremmling,
mostly

9Energy
e xp enrl it u r e and heart
rate for several
act iv it ie s of white-tailed
tained from W, W, Mautz,
Un iv , New Hampshire,
Durham.

standing.

walking,
mostly

remainder
remainder

Colo,

~

remainder

4Mean

female;

Mautz and Fair,
unpublished
ms , 9

standing,

standing
standing

and walking,
and walking.

deer--unpublished

ms , ob-

�-106-

Heart

rates

measured

in this

with those

from the two studies

rates

were

slightly

pected

due to the mixture

higher

study were

mentioned

above.

than in the other
of activities

in general

agreement

Bedding heart

studies,

but this is ex-

including

standing

white-tailed

deer

and walking

as well as bedding.
Heart

rates

for an adult female

(Mautz and Fair,
the other

unpublished

three

studies.

with a seasonal

effect.

Mean energy
for the same
literature

The difference

expenditures

categories

values

for fasting

cost of activities

81 percent

of the time

reported

was 109

for young mule deer
et a l , 1969),

Higher

values

6) and with values
(Table

which resulted
input for these

chambers.

to

heat produ&lt;:-

for the energy

7).
deer

bedding

54 to

=- 6 kcal kg -0 •75 day -1 (4.5 ±. O. 3 kcal
higher

than previously

(Knox et al , 1974), white-tailed

in this

in

were calculated

and resting

study for fasted

and pronghorn

observed

mask procedure,
ity and sensory

errors

This value is 18 to 49 percent

(Silver

piration

in this

rates

to make comparisons

heat production

for wild ungulates

expenditure

kg -0. 75 hr -1).

in order

(Table

than heart

may have been associated

and standard

of trials

tion of young wild ungulates

Energy

rn s , ) were higher

in summer

(Wesley

et a l , 1970,

study may be related
in a somewhat
animals

wider

than for those

deer

1973).

to use of the face
range

of act iv-

studied

in res-

�Table

6.

Measurements

Species

Mule deer

of fasting

and resting

Level of
Feed

Method

Fast

Face-mask

(fed) heat

production

Sub iect aConditions

(or young wild

ungulates.

Energy

Expenditure

Activity
kcal

As Reported

Source

kg-O• 75 hr-1

109

k c a Ik g -0.75d

ay -1

4.6

This

--

89

k ca Ik g -0.75d

ay -I

3.7

Knox et al ,
1974

Fawnswinter

~~

90.2

kcal

kg

3.8

Silver et a l ,
1969

Fawns
2d. 29

--

92

kcal

kg-O• 75 day-I

3.8

Wesley et al ,
1970

--

73

kcal

kg-O• 75 day~1

3.0

Wesley et a l ,
1973

121

kcal

kg-O, 75 day~1

5.0

This

154.0

kcal

kg

6.4

Nordan et al ,
1970

149.0

kcal

kg

114

kc a l kg-a'

Fawns

let.

Bedding &gt;50'1.
X= 65'1.

19

study

8 mo
Mule deer

Wh ite ta il ed
deer
s

Pronghorn

Fast

Fa&amp;t

Fast

Chamber

Chamber

Chamber

Fawns
9-12

mo

108-182
Fast

Chamber

Mule deer

Fed

Face-mask

d ay

-I

e;

mo

Fawns
let,

Bedding&gt;
50%
X = 88%

19

study

8 mo
Black-tailed
deer

Fed

Chamber

d-153 days
9-166

Pronghorn

~-

Fawns

Fed

Chamber

days

Fawns
9-7.5

mo

---

-0.75
-0,75

day
day

-I
-I

75 day-I

•....
0

days

Fawns
7.5

-0.75

6.2
4.8

Wesley et a l ,
1973

"
I

�Table

7.

Measurements

of energy

cost

of activities

for wild

ungulates.

Energy
Species

Expenditure

Activity

k

As Reported
Wh ite -ta i led
deer

White -tailed
deer

Mule

deer

Reindeer

W.

318.2

kc a l kg

Walkingno snow

650.9

kcal

Walkingsnow

898.6

k ca Ik g -0.75d

Running

14fl8,8

kcal

Plunging
in snow

\035,4

k ca I k g -0. 75 d ay - I

Stand ing (&gt;50 %)
remainder
walking

0,093

kcal

kg

kg -0,75

kg

Walking
(&gt;50%)
remainder
standing

0,100

kc a l kg

Running

0.145

kcal

or bounding

k

-0.75

kg

c a Ik g

-0,75

-0

,

75

-0 75
.

day

5,6

B.O

12,7

304

k ca 1.-"g - 0, 7 5 d ay - I

Standing

175

kr a l hr

fl8B

kcalhr-I

Stanrl ing

45

kc a l kg

-I

as average

d if Ie r e nce

in heat

5,5

This

study

Hammel

et a l , 1962

Wesley

et ai,

1973

obtained

from

21.8
-0.75

day

_12

of wh i t e

p rod uct io n [o r lying

•.....
00
I

B,7

-I

Walking
172-77%)
(20 -28'7. standing,
0-70/0 trotting)

I

6,0

-I

193

Mautz and Fair,
unpublished
me,

o

-1

Standing
180-95'7.)
(5-20 '7. walking)

load

1974

43,1

,-\

ay

Mattfeld

61,2

rrn n

min

Source
r

37.4

day-I

min

h

27. I

-I

ay

-0.75 -l
13.3

-1

day

-O,75d

IEnergy
ex p e nd it u r e and heart
rate for s e ve r a l activities
VI, Maut z , Un iv , New Harnp s h i r e , Du rh a rn ,
2H.cported

-\

Stand ing

Pulling
Pronghorn

-0.75

c a II "g

6.9

s

t a i le d

and standing

deer

- u npub l i s h ed ms.

animals.

�-109-

Resting

energy

expenditure

of fed deer

cent of the time was 121 + 3 kcal kg -0.75
k g -0.75h
ments

r, -1)

slightly

higher

deer

day -1 (5.0 + 0.1 kcal

Iower t h an preVIOUS
"
measure-

wh iICh" IS 26 percent

for fed black-tailed

bedding 73 to 100 per-

fawns (Nordan

than measurements

et a l , 1970), but

for young pronghorn

(Wesley

et

a l , 1973).
The energy
time was 193+

expenditure

17kcalkg

This value is intermediate
white-tailed

deer

of deer

standing

80 to 95 percent

-0 75
-1
•
day
(8.0+0.7kcalkg
between

values

by Mautz and Fair

reported

(unpublished

of the

-0 75
-1
•
hr
).
for standing

ms.)

and Mattfeld

et at.

(1974) respectively.

Values for adult reindeer

and young pronghorn

(calculated

from Wesley

et at.

1973) were

My measurement

is probably

higher

than a true

slightly

lower.

value for standing
small

amount

mule deer

of walking

Mean energy
304 + 16 kcal kg-·

walking

what greater

inc luded in these

greater

cost of the

in this

study was

trials.

for walking trials

by Mautz and Fair
than 50 percent

whitetails

in the present

if trials

reported

(unpublished
of the time,

by Mattfeld

entirely

ms.)

This

is

for

but about half
(1974).

study could be expected

had consisted

1962)

energy

o 75 day - 1 (12.7.±. o. 7 kcal kg -0 •75 hr -1 ).

the value for walking
value measured

due to the additional

expenditure

twice the value reported
whitetails

(Hammel

of walking.

The

to be some-

�-110-

Effects

of Activity
From

penditure

Level and Feed Level

the high correlations

for individuals

effect

of activity

level

energy

expenditure

This conclusion

it appears

and heart

cepts

of the regressions

Based

on the limited

expenditure

levels,

that the same

rate

applies

level.

expenditure

-heart

relationship

the activity

and feed levels

tested.

Comparison

of Regression

with Literature

In their
were

rate

reported

study

by Holter

165 energy

made using
rates

different

equations

trials

of energy

were

by varying

obtained

it appears

between

that

energy
differ-

wh en the comparison
that a single

was

energy
over

Results
expenditure

on heart

et ala (1976) for white-tailed

expenditure

were varied

no. 51.

would hold for an individual

six adult white-tailed

and heart

of deer

in inter-

and fed data from all

Thus it appears

the regression

for

between

was a significant

no. 51 for fasting
disappeared

of covariance

difference

levels

Although there

the difference

ex-

over a l l vac t iv ity levels.

does not affect the relationship
rate.

and energy

relationship

due to the significant

m.ade within an activity

with results

rate

of analysis

for the two activity

for deer

I compared

heart

amount of data for fasting

and heart

ence in intercept
activity

and the results

is tentative

feed level likewise

between

and heart
deer.
ambient

rate

deer.

rate d et e r rrrin at iorrs
Energy

expenditures

temperature.

for the four seasons

Four

of the year.

�-111-

Equations
deer

are given in Table 8, along with my equations

converted

to predict

Equations

from the present

those

obtained

grouped

2.90,

in the whitetail

individuals

the female,

energy

expenditure

in my study were 6.47 for males

with the winter

species

or age difference.

partially

explain

like well-conditioned

energy

expenditures

ones.

It is also possible

energy

at a given heart

possible

and heart

rates

rate

so different

a possible

wholly to a

condition

could

would be expected
rate

than poorly
rates

ani-

to have higher

conditioned
in the present

sometimes

being

is that the relationship

between

is different

are active

cold stress.

at a given level of energy

•

basis

of a study by Segrem
Peromyscus

expenditure

While conclusions

in size cannot be generalized

theoretical

when deer

Results

such a difference.

posed to cold than when exercising.
cies

unlikely

since well-conditioned

due to the EKG signal

(I 967b) suggested

heart

1. 10 to

It seems

in physical

that some of the heart

explanation

than when they are under

higher

from

activity.

expenditure

and Hart

at 1.10.

in slope,

humans,

underestimated

A third

varied

from

and 5.64 for

could be attributed

Difference

the difference

. mals,

during

deer

slope being lowest

that a differ ence of this magnitude

obscured

slope than

Slopes of the equations

while those for white-tailed

study were

in the same units.

study have a greater

study.

for mule

to deer,

for such an explanation.

had
when ex-

from

a spe-

there

is also

Heart

rate

and

�-112-

Table

B;

Regression
equations
for white-tailed
deer

\vl1ite-tailed

EE

= energy

.
HR = heart

deer

et ale 1976)

Summer:

MR = 2.90

(HR) - 15.2

Fall:

MR = 2.40

(HR) + 1. 6

Winter:

MR = 1. 10 (HR) + 42.4

Spring:

MR = 2.79

(This

study

(HR) - 26.6

- winter)

*

Males:

EE

= 6.47

(HR) - 180.48

Female:

EE = 5.64

(HR) - 218.29

Overall:

EE = 3.64

(HR) - 21.30

rate

(kcal kg

expenditure
rate

on heart

(Holter

Mule deer

MR = metabolic

of energy expenditure
and mule deer.

(beats

-0 75
-1
•
day
)

(kcal kg
min

-1

-0 75
.

day

-1

)

)

.•.
-r- Units

of equations

converted

for comparisons.

o

r at e '

�-113-

stroke

volume together

tissues.

determine

the amount of blood supplied

to the

in exercising

due

Stroke volume increases

to the effect of muscle

activity

to the heart (Mountcastle
resulted

from activity

in increasing

1974).
in deer,

If similar

these

Differences

Energy
equivalent

expendi-

slope of the regression

in the present

in Regression

expenditure

of deer

study appears

of energy

consistent

with

in having consistently

planations

variance

for the two males

higher

but this deer differed
heart

rates

about the regression

for this difference

expenditure

Equations

no. 55 for a given activity

to that of the other deer,

without increased

energy

rate

considerations.

Individual

others

volume increases

animals- at the same energy

The greater

rate

of venous blood

animals

ture

on heart

stroke

partly

for active

inactive

expenditure

return

a lower heart

than for cold-stressed,
could be expected.

humans,

on heart

in the intercept
rate

for deer

include nervousness,

was

from the

for given activities
line.

Possible

of the regression

exof

no. 55 from the equation

level of conditioning,

sex and

body weight.
In studies
lower correlations

of sheep,
between

animals
energy

way and McEwan 1969, Webster
55 was high; therefore,
did not manifest

itself

which were more nervous
expenditure

1967).

and heart

The correlation

if a high level of nervousness
as in the sheep.

rate

had
(Brock-

for deer

no.

was a factor

it

�-114-

Consideration
better

of differences

explanation

of individual

differences.

efficients

between

rate

on body size supported

based

sions based

energy

on a sample

size be considered

in body size seemed

expenditure

of three

to explain

Higher

to provide

correlation

and transformations

this explanation.
are tentative,

individual

co-

of heart

While conclu-

I suggest

variation

a

that body

in similar

investi-

gations.
Sex differences
among individuals.
not provide
dividual

may explain another
Because

a test

of sex differences

differences.

Holter

influence

of sex on metabolic

stress.

Coefficients

and spring

trials

However,

Astrand

using both sexes

(1960) presented

heart

rate

rate

studies

of human males

were

consistently

higher

(although

ceding paragraph

sexes

observed

this experiment

deer

for regressions

a composite

and females.

respectively.

based on two separates

of females

at any given energy

of body weight differences
Thus there

which tends to corroborate

on fall

graph of the relation-

Heart

than those of males

in this study.

of in-

exposed to cold
based

were O. 79 and 0.88

and oxygen consumption

effects

explanation

did

1976) found no significant

of white-tailed

were not considered).

dence from humans

samples,

et a l , (1975,
rate

of the variability

as a possible

of determination

ship between

penditure

of limited

part

ex-

noted in the preis limited

differences

evi-

among

�-115-

In summary,
plain significant
tion of deer
number

differences

differences

in the intercept

no. 55 from that of deer

of subjects

Confidence

prevented

of Predicting

ments

the low variance
at the center

provides

wide range
pattern

in heart

most confidence
of heart

most closely,

the widest

range

confidence

could be placed

heart

rate.

and heart

rates.

tribute

measurements

more

of measure-

The data pattern

no.

Because

of the data patterns,
of energy

based

design.

over a

intervals

between
individuals

data.

energy

from

This is

on the

not of the biological

evenly over a wider range

over

expenditure

grouped

studies

this

little

expenditure

on these

Future

that

55 approximated

confidence

and is a result

but of the experimental

expenditure

is evenly distributed

were high for data from

of sex and body size,

Rate

in narrow

equations

tionship

rates.

of heart

of the fact that correlations

rate

energy

rate due to clustering

in predictions

rate using regression

true in spite

basis

of heart

equa-

The limited

from Heart

for predicting

Data from deer

resulting

ex-

conclusions.

in prediction

rates.

51 and 53.

Expenditure

inter-vals

of the range

of the regression

nos.

further

Energy

Shape of confidence
reflects

in body size or sex could partly

rela-

should disof heart

�-116-

Uses of the Results
High correlations
individual

deer

between

indicate

energy

that individual

expenditure
deer

ship was known could be used to measure
using heart
measured

rate

alone.

To do this,

relationship

for a deer

used to estimate

was known, heart

the energy

Results

the energy

should be

rate

Once the

could theoretically
and conditions

that precise

estimation

expenditure

from the heart

not possible

with present

knowledge.

However,

of heart

and energy

expenditure

suggests

heart

can be used as an ind ex of energy

heart

rate

rates,

rate

be

other

the relationship.

of this study indicate
rate

cost of activities

and activities.

cost of activities

than those used for determining

rate for

for which this relation-

the relationship

over a wide range of conditions

and heart

of deer not individually

of energy

calibrated

is

the high correlation
that in this range of
expenditure.

�-117-

CHAPTER

SUMMARY

A method
domestic

collection

cardiograph

measurements

as a predictor
Measurements

two fasted

for other

expenditure

Energy

Eleven

wild ungulates.

values

Heart
(r

= 0.81,

heart

energy

0.92,

rates,

of energy

rate

heart

conditions.

of bedding,

stand-

were made using
rates

for similar

Energy

Electro-

under these

and heart

of measurements

under

for evaluating

measurements

expenditures

and

a face mask for

fawns.

information

expenditure

bedding most of the tim.e was up to 50 percent
reported

mule deer

were made for combinations

were within the range
ported

8 -month-old

provided

and trotting.

deer.

of humans

were made using

from three

of energy

expenditure

for use on active

Measurements

respiratory

ing, walking,

energy

was adapted

penned conditions.

rate

AND CONCLUSIONS

for measuring

animals

V

higher

of fed deer

activities

re-

of fasted

deer

than previously

for young wild ungulates.
was correlated
0.92

expenditure

expenditure

for all deer

respectively),

but due to limited

intervals

were too wide for precise

confidence

expenditure

with energy

from heart
on heart

while that for the female

rate

differed,

rate.

Regression

for the two males
possibly

distribution

prediction

equations
were

of

of

similar,

due to differences

in sex

�-118-

and body weight.

Regression

slope from those

of Holter

possible
tures

explanation

and heart

stressed

rate

IS

study,

expenditure

However,

variation

deer.

A

between

energy

for active

animals

heart

shows promise

is limited,

heart

Main contributions
collection

system

ditioned

to accept

present

rate

expendi-

than it is for cold-

energy

costs

of this

of certain
between

mule deer

over the range

Future

study were:

heart

studies

rate

and energy

should attempt
cost,

ing energy

expenditure

for use under

the range

and distribution

an increase

in heart

to evaluate

the energy

fawns conthe

and demonstrating

a

for individual

rates

without

activities

for de-

methods

of estimat-

conditions,

3) spread

other

different

expenditure,
rate

deer

tested.

2) investigate

of heart

of

a respiratory

measuring

expenditure

to 1) isolate

of energy

estimates

calibrated.

adapting

for mule deer,

of activities

energy

precise

for use on active

termination

in predicting

of intraspecific

without tranquilization,

activities

correlation

knowledge

not individually

with a face mask

as an

which have been individually

cannot provide

deer

the system

rate

for deer

because

exp e nd i.tur e for active

in order

in

et ale (1976) for white-tailed

different

of this

of energy

calibrated.

elicit

differed

animals.

estimator

fidence

for both groups

is that the relationship

On the basis

energy

equations

in order

to increase

4) introduce
an overt

expenditure-heart

stimuli

increase

conwhich

in activity,

rate.relationship

in

�-119-

this

situation,

and 5) evaluate

weight diff~rences
expenditure

hypotheses

among individuals

and heart

rate.

regarding

effects

on the relationship

of sex and

of energy

�-120-

LITERATU RE CITED
o
Astrand,
1. 1960. Work capacity in m.en and wom.en with special reference to age.
Acta Physiologica
Scandinavica
49. Supplem.entum. 169. 92 pp.
o
Astrand,
P.O.,
and K. Rodahl.
McGraw Hill, New York.
Baker,

1970. Textbook
669 pp.

D. 1976. Energy requirem.ents
of m.ule deer
M. S. Thesis.
Colorado St. Urriv , 76 pp.

Booyens, J., and G. R. Hervey.
m.easuring m.etabolic rate
38: 130 1- 1309.

Brockway,
J. M., and E. H. McEwan.
cardiac perform.ance
in the sheep.
S. 1945.
New York.

Bioenergetics
1023 pp.

fawns in winter.

1960. The pulse rate as a m.eans of
in rnan , Can. J. Biochem.. Physiol.

Brockway,
J. M., and G. M. O. Maloiy.
1968.
of the red deer.
J. Physiol.
194:22-23.

Brody,

of Work Physiology.

Energy

m.etabolism.

1969. Oxygen uptake and
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202: 661-669.

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Reinhold

Publishing

cs.,

Carpenter,
L. H. 1976. Middle Park deer study - deer habitat evaluation.
Colo. Div. Wildl.,
Gam.e Res. Div., Fed. Aid Proj.
W-38-R-31.
Gam.e Res. Rep., July, Part 2. p.283-405.
Consolazio,
C. F.,
Physiological
McGraw-Hill,

R. E. Johnson,
and L. J. Pecora.
1963.
m.easurem.ents
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505 pp.

Corbett,
J. L., D. J. Farrell,
R. A. Le ng , G. L. McClym.ont, and
B. A. Young.
1971. Determ.ination
of the energy expenditure
of penned and grazing
sheep from. estim.ates of carbon dioxide
entry rate.
Br. J. Nut r , 26:277-291.
Datta,

S. R., and N. L. Ram.anathan.
1969. Energy expenditure
work predicted
from. heart rate and pulm.onary ventilation.
App l , Physiol.
26(3):297-302.

in

J.

�-121-

Draper,
N. R., and H. Smith.
1966.
John Wiley &amp; Sons, New York.

Applied regression
407 pp.

analysis.

Fair,

J.
1978. An analysis of the relationship
between energy expenditure and heart rate of unrestrained
white-tailed
deer.
M.S. Thesis.
Univ. of New Hampshire.
61 pp.

Gill,

R. B. 1976. Mule deer management
myths and the population
decline.
Pr oc , Deer Decline in the West Symposium.
(In
press) •

Hammel,
H. T., T. R. Houpt, K. Lange Anders n, and S. Sjenneberg.
1962. Thermal and metabolic measurements
of a reindeer
at
rest and in exercise.
Tech. Doc. Rep.
AAL-TDR 61-54.
Arctic Aeromedical
Lab, Fort Wainwright,
Alaska.
34 pp.
Hargrove,
J. L., and J. A. Gessaman.
1973. An evaluation of
respiratory
rate as an indirect monitor of free-living
metabolism.
Pages 77 -85 in J. A. Ges saman, ed , , Ecological
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of homeotherms.
Utah State University
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1967. A versatile
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oxygen
consumption
in man.
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Kleiber,

M.

1961.

0':: life.

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454 pp.
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Theory
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G. F., and G. E. Folk.
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G.

1928.

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994-996.
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Comparative
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McEwan, E. H. 1970. Energy metabolism
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McEwan, E. H. and P, E. Whitehead.
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458 pp.

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Morhardt,
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1971. Correlations
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Nordan, H. C., 1. McT. Cowan, and A. J. Wood.
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Can. J. Zool.
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D. W. 1972. Rearing and training deer for food habits
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J.

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Original

Robbins,
C. T.
1973.
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The biological basis for the determination
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A. 1966. The utilization
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u,

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Rogerson,
A.
1968. Energy utilization
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Comparative
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Oxygen supply and per-(
formance
in Peromyscus.
Metabolic and circulatory
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Pharm.
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Silver,

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Fasting metabolism
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J. Wildl. Manage.
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Effect of falling temperature
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white-tailed
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1967.
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Statistical
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the
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1973. Nutrition of white-tailed
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D. E., W. G. Youatt, H. E. Johnson,
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1969. Digestible
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J. Wildl. Manage.
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Ullrey,

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0

Wesley, D. E.,
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physiologie

1.

�-126-

APPENDIX

�-127-

APPENDIX A: Data collection form modified from Consolazio et a l ,
(1963) for measuring mule deer energy expenditure.

Experiment

No.

Date

-------------------Location
------------------

Deer No.
Wt. ____

Ibs , x 0.4536

Time Hookup

= wt , kg. __

= wt. kgO•75

_

Time Start

----------------

Time Stop

_

Time Elapsed
Meter Final

---------

Initial

-----------

°

x CF 1.

Diff.
Start

----------- °c

Finish

----------- °c

Avg.

°c

Meter Temp.

-

1

Outside Temp.

S

°c

F

°c

=

mb x 0.750

Bare Pres.

14

_

mm

PH 0------2

Volu me STPD

Bag No.

=

1 [760 ~~: :.::;~ TJ
1

°

0

=

----------------

2 ---------

1.

�-128-

APPENDIX

Symbols:

B:

Derivation
of formula to calculate
oxygen consumption
and energy expenditure
from volume and fractional
concentr ation of oxygen in expired air, as suming
volume of methane produced is insignificant.

VI

=

volume

of air

=

volume

of air expired

= fractional

inspired

concentration

of oxygen in inspired

air

= fractional
concentration
(% concentration/
100)

of oxygen in expired

air

(% concentration/IOO)

Vo

=

2

Veo

2

RQ

volume

= volume

of oxygen consumed
of carbon

= respiratory

dioxide

quotient

produced

= Veo

IV 0
2/

Known:

F I = • 20946
VE'

FE measured

VI is unknown but can be found for specific
methane production
is assumed to equal zero.

Since

=

VE
VI =

At

2

VI - V

02

+ Veo

VE + V

02 - Veo

RQ = 0.82
VI =
=

VE + Vo

2

+ 0.18
E

V

- 0.82

"o

2

Vo
2

2
2

values

of RQ if

�-129-

APPENDIX

B (continued).

Substituting

for

VI in equation

for Vo

,
2

[(V E + O. 18 Vo ) F

=

Vo

2

2

= [(VE + 0.18
=

1

V

) 0.209]

"o - 0.038

Vo

0.962

Vo

- [VE

02

VE) + (0.038

(0.209

11 - [ VE FE]

V

)

FE]

- [ VE FE]

02

=

0.209

=

VE (0.209

- FE)

=

VE (0.209

- FE) 10. 962

1 1..02 results

in 4.825

2

2

VE - (VE FE)

2
Vo

At

RQ

= 0.82

so

EE

=

J

4.82

(1)

2
kcal

of heat

produced,

x Vo
2

Substituting

for V0 '
2
EE

Sample

= 4.82

[V

0.209

=

12.0 1. Imin

E

(2)

- FE)]/O.962

calculation:
V

E

at STP

FE is . 171

= [12.0
EE

= 4.82

- 0.171)1/0.962

(0.209

(0.474)

=

2.?'9kcal/min

= 0.4741/min

�APPENDIX

C (Table

9):
Weights (kg). oxygen consumptions
(ml min -1), energy expend itur e s (kc a l
l
kg-C.75 min- ),
heart rates (beats min-1), and activities
(percent bedding,
standing, walking, trotting) for trials using three fasting and fed mule deer
fawns in winter.

Trial
No.

1

Feed
Level

Deer

Weight
(kg)

Oxygen
Consump.
(ml/min)

Energy
Expend.
MI3W rn in]

Heart
Rate
(beats /
min)

Percent
Bedding

Percent
Standing

Percent
Walking

Percent
Trotting

(kc a l/

2
3
4
5

Fed
Fed
Fed
Fed
Fed

51
51
51
51
51

44.34
44.34
44.34
44.34
44.34

483
550
548
415
524

Fed
Fed
Fed
Fed
Fed

57.7
62.4
59. 3
54. 3
56. 7

51
51
51
51
51

0
0
0
0
0

44.68
44.68
44.68
44.68
44.68

69
50
80
76
63

598
566
589
529
629

31
50
20
24
37

.. 0

6
7
8
9
10

0.136
0.154
0.154
O. 117
O. 147
O. 167
O. 158

0.164
0.148
O. 175

11
12
13
14
15

Fed
Fed
Fed
Fed
Fed

62.0
63.5
65.6
59.0
62.3

51
51
51
51
51

0
0
0
0
0

44.68
44.68
44.68
45.93
45.93

72
70
69 .
73
51

627
559
606
573
600

28
30
31
27
49

16
17
18
19
20

0.175
0.156
0.169
0.157
O. 164

0
0
0
0
0

Fed
Fed
Fed
Fed
Fed

64.0
58.1
57.3
67.0
60.4

51
51
51
51
51

0
0
0
0
0

46.61
46.61
46.61
46.61
47. 17

64
57
52
81
81

681
531
503
567
800

35
43
48
19
19

O. 184
0.144
O. 136
0.153
0.214

1
0
0
0
0

66.2
61.8
63. 1
62.3
73.8

0
0
0
0
0

28
79
67
46
53

72
21
33
54
47

0
0
0
0
0

0
0
0
0
I
t-'

w

0
I

�APPENDIX

C (Table 9) (continued).
Energy
Expend.
(kcal/
MBW min)

Heart
Rate
(beatsI
min}

Percent
Bedding

61.3
64.4
61.6
65.6
51.8

0
0
0
0
54

74
61
75
64
35

26
39
25
36
11

0
0
0
0
0

0
0
13
63
0

58
73
47
35
77

42
27
40
2
23

0
0
0
0
0

Trial
No.

Feed
Level

Deer

Weight
(kg)

Oxygen
Consump.
(ml/min)

21
22
24
25
26

Fed
Fed
Fed
Fast
Fast

51
51
51
51
51

47. 17
47. 17
47.17
44.91
44.91

528
626
578
455
320

O. 141

27
28
29
30
31

Fast
Fast
Fast
Fast
Fed

51
51
51
51
51

44.91
44.91
44.91
44.91
44.79

520
483
691
271
436

O. 140
O. 130
O. 186

0.073
O. 121

60.7
57. 1
66.2
50.6
56.4

32
33
34
35

51
51
51
51

73.3
81.6
70.5
58.5
66.0

53

43

4

0

568

0.207
0.205
0.180
O. 100
O. 179

0

53

44.79
44.79
44.79
44.79
37.99

744
735
647
357

36

Fed
Fed
Fed
Fed
Fed

0

86
48

14
52

0
0

37
38
39
40
41

Fed
Fed
Fed
Fed
Fed

53
53
53
53
53

37.99
37.99
37.99
37.99
38.44

566
547
595
537
633

0.178
O. 173
O. 188
0.169
O. 198

66.7
66.7
71.3
75.0
72.0

0
0
0
0

57
61
50
38

43
38
49
57

0

0.168
0.155
O. 123
0.086

Percent Percent
Standing Walking

Percent
Trotting

1
1
5

I
&gt;-'

w
•....•
I

�APPENDIX

C (Table 9) (continued).

Weight
(kg)

Oxygen
Consump.
(ml/min)

Energy
Expend.
(kcal!
MBW min)

Heart
Rate
(beats/
min)

53
53
53
53
53

38.44
38.44
38.44
38.44
39.35

639
625
656
634
508

0.200
0.195
0.205
0.198
o. 156

62.4
71.0

Fed
Fed
Fed
Fed
Fed

53
53
53
53
53

39.35
39.35
39.35
40.25
40.25

580
592
601
500
543

O. 178

0.182
O. 184
O. 151
O. 164

53
54
55
56
57

Fed
Fed
Fed
Fed
Fed

53
53
53
53
53

40.26
40.26
40.26
40.26
40.71

618
537
446
518
474

O. 162
O. 135
O. 156
O. 142

58
59
60
61
62

Fed
Fed
Fed
Fed
Fed

53
53
53
53
53

40.71
40.71
40.71
40.71
40.71

624
478
909
741
771

Trial
No.

Feed
Level

Deer

42
43
44
45
46

Fed
Fed
Fed
Fed
Fed

47
48
49
51
52

0.186

O. 186
O. 143

0.272
0.222
0.231

Percent
Bedding

Percent
Standing

Percent
Walking

Percent
Trotting

68.4
66.7

0
0
0
0
0

50
54
59
51
54

48
45
41
48
46

2
1
0
1
0

62.0
67.6
72.0
63.0
60.9

0
0
0
0
0

67
36
23
58
33

33
64
77
42
67

0
0
0
0
0

62.7
63.0
57.5
59.7
59.6

0
0
0
0
0

51
55
74
68
59

47
45
26
29
41

2
0
0
3
0

64.4
63.1
81.6
83. 1
74.9

0
0
0
0
0

24
67
20
44
22

76
33
73
47
74

0
0
7
9
4

--

I
•....•

w

N
I

�APPENDIX C (Table 9) (continued).

D

eer.

Weight
(kg)

Oxygen
Consump.
(ml/min)

Energy
Expend.
(kcal/
'MBW min)

Heart
Rate
(beats/
'min)

Percent
Bedding

Percent
Standing

Percent
Walking

Percent
Trotting

Trial
No.

Feed
Leve l

63
64
65
66
67

Fed
Fed
Fed
Fed
Fed

53
53
53
53
53

41.50
41. SO
41.50
41, SO
41.50

658
564
420
448
305

O. 194
0.166
O. 124
O. 132
0.090

69.3
63.4
56,0
56,8
51.6

0
0
42
18
73

27
64
42
47
26

73
36
16
35
1

0
0
0
0
0

68
69
70
71
72

Fed
Fed
Fed
Fed
Fed

53
53
53
53
53

41,50
42.41
42,41
42,41
39.35

427
1054 '
335
495
246

O. 126
0.306
0.097
O. 144
0.075

61.0
92.7
49.3
60.2
48.7

0
0
0
0
80

74
34
95
63
10

26
65
5
35
10

0
1
0
3

,-0 a

74
75
76
77
78

Fed
Fed
Fed
Fed
Fed

55
55
55
55
55

29.37
29.37
"z-q:37
29.37
29.37

611
517
541
655
486

0.233
0.197
0,207
0.250
0, 186

89.0
82,0
90.4

0
0
0
0
0

21
35
28
40
46

75
65
72
50
54

4
0
0
10
0

Fed
Fed
Fed
Fed
Fed

55
55
55
55
55

-2q::-37
28.80
28.80
28.80
28.80

426
607
565
511
459

O. 163
0,235
0.219
O. 198
O. 178

85.0
87.2
91.0
85.0
86.9

0
0

44
38

56
58

0

79
80
81
82
83

0
0

50
57

48
43

2
0

-r-

--

86.2

4

I
I-'

w
w
I

�APPENDIX

C (table 9) (continued).

Deer

Weight
(kg)

Oxygen
Consump.
(ml/min)

Energy
Expend.
(kcall
'MBW rnin)

Heart
Rate
(beats I
min)

Percent
Bedding

Percent
Standing

Percent
Walking

Fed
Fed
Fed
Fed
Fed

55
55
55
55
55

28.80
28.80
28.80
28.80
28.80

603
435.
429
295
552

0.234
O. 169
O. 166
O. 114

92.9
84.0
84.0
71.5
82.4

0
0
0
0
0

43
46
46
43
53

55
55
55
55
55

30.84
30.84
30.84
30.96
30.96

6
1
1
0
2

601
457
317
441
392

96.0
86.3
76.0
78.7
80.6

0
0
0
0
0

94
95
96
97
98

63
52
52
55
75

Fed
Fed
Fed
Fed
Fed

35
48
48
42
25

2

91
92
93

Fed
Fed
Fed
Fed
Fed

51
53
53
57
45

55
55
55
55
55

0
0
3
0

30.96
31. 53
31. 53
31. 53
31. 53

484
414
441
364
376

86.2
85.8
84.4
84.9
77,8

0
0
0
0
0

63
53
44
80
77

99
100
101
102
103

Fed
Fed
Fed
Fed
Fed

37
47
56
18
23

55
55
55
55
55

31. 53
31. 53
31. 53
31. 53
31. 53

0
0
0
2
0

518
564
239
236
409

89.8
86.7
58.2
63.3
77.0

0
0
80
85
0

36
39
10
11
50

62
54
10
4
50

2
7
0
0
0

Trial
No.

Feed
Level

84
85
86
87
88
89
l.9U

- 0.214
0.221
O. 168
O. 117
O. 162

0, 144
O. 178
O. 150

0.160
O. 132
O. 136
O. 188

0.204
0.086
0.086
O. 148

Percent
Trotting

I
I-'

w

~
I

�APPENDIX C (Table 9) (continued).

Deer

Weight
(kg)

Oxygen
Consump.
(mlfmin)

Energy
Expend.
(kcal!
'MEW min)

Heart
Rate
(beat sf
'min)

Percent
Bedding

Percent
Standing

Percent
Walking

Fed
Fed
Fed
Fed
Fast

55
55
55
55
55

31.53
31.35
31.53
31.53
30.39

360
235
211
218
587

0.130
0.085
0.076
0.079
0.213

70.7
59.8
58.7
54.5
91. 1

0
90
100
100
10

76
9
0
0
36

24
1
0
0'
53

Fast
Fast
Fast
Fast

55
55
55
55

30.39
30.39
30.39
30.39

465
251
184
215

0.168
0.091
0.066
0.078

81.8
61.5
56.7
55.8

0
10
"81
63

89
82
16
36

11
8
3
1

of bedding and trotting

activities.

Trial
No.

Feed
Level

104
105
106
107
108
109
110
111

112

aE stimated

activity breakdown.

bAtypical trial

due to combination

Percent
Trotting

0
0
0
°b

1

0
0
0
0

I
I-'

w

VI
I

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                  <text>July,

-137-

JOB PROGRESS

REPORT

State of

COLORADO
------~~~~~---------

Project

No.

W-38-R-33

14

Deer-Elk

J~ ~.

Investigations

11

~rkPI~

~.

Job Title

Snowmobile Harassment of Mule Deer on Cold Winter Ranges
--------------------------------------------------~--

Period

Covered:

April

1978

-------------------------------

1, 1977 - June 30, 1978

Personnel:
D. Freddy, L. Carpenter, B. Lance, B. Williams, L. Strong, C.
Moore, B. McCloskey, P. Neil, Colorado Division of Wildlife; A. L. Ward, U.S.
Forest Service; E. Tom Thorne, Wyoming Game and Fish Commission.

Heart
Heart

rates associated with activities of 2 semi-tame mule deer are presented.
rate varied by deer and activity and ranged from 34.87 ± 0.28 (SE) to
78.74 ± 4.90 (SE) (BPM) for bedded and walking activities, respectively.
Heart
rate data verify their potential to calculate energy costs of overt reactions
of deer to harassment.
Changes in heart rate, both positive and negative, also
indicated deer responded to harassment stimuli in the absence of strong overt
reactions.
Heart rate changed from -21 to 154 percent when compared to preharassment measurements for similar activities.
Elevated heart rates during
post-harassment
sampling period also suggested that physiological
responses
may linger for several minutes after stimuli have passed.
Three semi-tame
female deer, age 1~ years, and two wild deer, a yearling buck and a mature doe,
were equipped with implanted heart-rate monitoring systems during this segment.
Heart-rate data from these animals will be presented in next segment's report.

��-139-

SNOWMOBILE HARASSMENT OF MULE DEER
ON COLD WINTER RANGES
David J. Freddy
P. N. OBJECTIVE
Evaluate whether snowmobile activity on winter ranges inhabited by deer
decreases the ability of deer to survive winter by modifying the activities
of deer so as to significantly increase energy expended.
SEGMENT OBJECTIVES
1.

Monitor heart rates of deer during various daily activities
the energy costs of these activities during winter.

2.

Ascertain

3.

Evaluate observed reactions of deer to harassment in terms of appropriate
heart rate measurements and corresponding energy costs.

4.

Evaluate the energy cost of harassment to deer in relation
estimated energy budget of deer (concurring study).

5.

Examine alternatives
harassment on deer.

activities

of deer responding

to minimize

to prescribed

potential

detrimental

to quantify

harassment.

to a total

effects of

METHODS AND MATERIALS
1977
Heart rate for activities of 2 semi-tame, castrate mule deer, age 2~ years,
telemetered during winter of 1976-77 were analyzed during this segment.
Heart
rates for each activity (Tables 1, 2) were determined as follows.
Deer were
sampled from 7:00 AM to 5:30 PM during 2 trials in February.
Each trial
consisted of 5 consecutive days. Heart rates and activities were sampled during
3, systematically spaced 10 minute intervals per hour. One deer was sampled
per 10 minute interval.
Deer were alternately sampled, with the first deer
sampled each day selected at random.
Heart rate was monitored continuously
during the 10 minute interval and output was recorded on a strip-chart recorder
(Freddy 1977). Activities of deer were noted directly on the strip-chart.
Heart rates were then determined for each activity using a Hewlett-Packard
9820A digitizer-calculator
system. An activity had to occur continuously for
1 minute to sample heart rate. Within a I minute interval (approx. 4 cm of
strip-chart), 6-8 systematic hea.rt rate data points were digitized and an
average value determined for that 1 minute.
Within a 10 minute sample period,
up to 10, one minute samples of heart rate could be obtained.
Heart-rate for
each activity for each deer was based on the mean of several one minute samples
taken throughout each trial. T-tests were used for all tests of significance
at a probability level of P &lt; .05.

�-140-

Table 1. Definitions
at pasture.

of activities

for measuring

heart-rate

of mule deer

Definition

Activity

Abbreviation

Bedded

(B)

Lying down, not active

Bedded plus
rumination

(BR)

Lying down plus actively chewing and
possibly belching, enough of head must
be seen to discern jaw movement

Stand

(S)

Up, but not moving

Stand plus graze

(SG)

Standing still and actively grazing, or
occasionally walking very slowly, a few
steps at a time, and grazing

Walk

(W)

Moderately fast ambulatory movement,
all 4 feet touching ground alternately

Bedded-alert

(BA)

Alert appearance while bedded; head up,
ears forward, possible tail-flick,
apparently watching or listening to some
disturbance

Table 2. Heart-rates (BPM) for activities of 2 semi-tame mule deer castrates,
age 2Yz years, February, 1977 . Trial 1 was conducted from February 7-12 and
trial 2 from February 14-19.
Trial 2
Deer 39
Deer 18
HeartHeartrate + SE
rate + SE
n

Abbrev.

Trial
I
Deer 18
Deer 39
HeartHeartrate + SE
n
rate + SE

Bedded

(B)

48.80+0.23

485

34.87+0.28

197 44.06+0.44

216

Bedded plus
rumination

(BR)

50.53+0.32

94

37.62+0.58

55 47.55+0.71

60

Stand

(S)

64.05+1.49

49

49.84+1. 86

16 65.35+3.93

16

Stand
plus graze

(SG)

78.37+1.20

99

55.15+0.84

64 68.59+2.18

31

Walkb

(W)

91.37+3.79

8

62.35+5.41

4

Bedded-alert

(BA)

53.21+0.95

48

33.55+0.98

7 78.74+4.90
12 50.60+1. 66

Activity

a
b

equals number of one minute samples
t-tests were not conducted

n

a

used to compute heart-rate

for the walk activity because

n
--

27

for each activirv.

of small sample sizes·

�-141-

Harassment trials utilizing these same castrate deer were conducted subsequent
to unharassed heart rate trials.
Six routes were traversed by one person,
two persons, person plus a dog, and a snowmobile.
Routes were 0-360 m from
the pasture fence. Up to 4 routes were traversed in one day. Heart rate
and overt behavior were monitored continuously for one deer while each route
was traversed and also for 5 minutes prior to (pre-route) and after (postroute) completion of each route. Harassment was considered to begin and
end when harassing stimuli were either within sight or audible to the observer.
Deer monitored were alternated with each route, with the first deer to be
sampled each day selected at random.
Procedure for determining harassed
heart-rate was the same as for unharassed heart-rate trials.
1978
During winter of 1977-78, 3 semi-tame female deer, age 1~ years, were equipped
with implanted heart rate transmitters.
Transmitters were essentially the
same as used in 1977 but were modified in packaging and circuitry to reduce
problems of artifact (Freddy 1977).
Surgery for implanting the transmitters was the same as in 1977 (Freddy 1977).
However, the drugs M-99 and Rompun were combined for anesthesia instead of
the Rompun-Ketamine combination used previously.
Approximately 4 mg of M-99
combined with 30 mg of Rompun were given intramuscularly for deer averaging 60
kg (133 lbs).
These semi-tame deer were placed in a 4 ha pasture of native sagebrush winter
range and visually and telemetrically monitored from a glass enclosed tower
adjacent to the pasture (Freddy 1977). Heart rates and corresponding activities
were monitored in January, February, March, April, and June. Numbers of deer
per trial and days per trial varied between months according to the number
of telemetry systems functioning.
Heart rate for each activity was determined
using the same procedure as in 1977. While at pasture these deer were harassed
by a snowmobile, person, and a person and a dog moving on prescribed routes
at varying distances from the pasture.
Deer activities and heart rates were
monitored using previous procedures.
Two wild mule deer, one yearling male and one mature female, were also equipped
with implanted heart rate transmitters.
Deer were baited with alfalfa hay
and trapped in modified Clover traps. Handling procedures were as follows:
1) deer were physically restrained in the trap by a trap tender,
2) M-99 and
Rompun were injected into the rump of the first deer (male) while only Rompun
was injected into the second deer (female). This was done because surgery was
sequential and we did not want the second deer under the effects of M-99 for
about 1 hour prior to surgery and then again for 45 minutes during surgery,
3) deer were placed in small carrying boxes to minimize their struggle,
4)
after drugs had quieted deer, they were transported 360 m to research facility
for surgery,
5) the male was immediately taken to surgery while the female
was left in the carrying box under the effect of Rompun,
6) surgery was
conducted as previously described.
M-99 was given to the female just before
her surgery.
These deer were released and their heart rates monitored for
approximately 30 days.

�-142-

RESULTS AND DISCUSSION

1977
Heart rate (beats per minute - BPM) increased with increasing levels of
activity for both deer (Table 2). Within trial 1, significant differences
in heart rate occurred between all activities for deer 39. No data were
obtained from deer 18 because of implant failure. Within trial 2, significant differences in heart rate occurred between all activities for deer 18
except for activities (B) vs (BA) and (S) vs (SG) and for deer 39 except for
activities (S) vs (SG) (See Table 1 for activity symbol definitions).
Between trials, there were significant differences in heart rate for all
activities for deer 39 except for activities (S) and (BA). Heart rates for
deer 18 and 39 were significantly different between deer for all activities
during trial 2. Deer 39 weighed about 60 kg during trials 1 and 2 while
deer 18 weighed about 63 kg during trial 2. Weights of both deer declined
from January through March.
Kautz (1978) also found heart rate increased by increasing activity level
and varied between individual deer. Furthermore Kautz (1978) demonstrated
a positive relationship between heart rate and energy expended.
Data from
our 2 semi-tame deer verify the potential to use heart rate for estimating
energy expended for different activities and therefore the potential for
placing energy costs on overt reactions of deer to harassment.
Changes in heart rate also indicated deer responded to harassment stimuli
even in the absence of strong overt reactions (Table 3). Changes in heart
rate, both positive and negative, indicated animals were stressed to some
degree.
The extent of heart rate change varied considerably with stimulus,
route, and deer. Heart rate changed from -21 to 154 percent when compared
to pre-harassment measurements for the same activities.
Elevated heart-rates
were usually of short duration.
The negative heart rate response of deer
18 (Table 3) would suggest alarm bradycardia.
Moen et al. (1978) also detected
alarm bradycardia responses in white-tailed deer fawns when there was little
or no change in overt behavior.
Elevated heart rates during post-route sampling
periods (Table 3) also suggests that physiological responses may linger for
several minutes after stimuli have passed.
Selye (1976) commented that the
cardiovascular system is particularly sensitive to stress and usually responds
with rising blood pressure and pulse rate. Data from these trials support the
contention that heart rate can indicate deer react to stress without changes
in overt behavior.

1978
Heart rate data obtained during

1978 are being processed.

During 1978, the telemetry system performed satisfactorily.
Artifact (as
reported by Freddy 1977) was reduced and heart rates for walking and running
activities were obtained to a greater extent than in 1977. The one problem

�Table 3.

Heart-rate

(HR) of semi-tame

Distance (m)
to
Stimulus

deer for similar

Deer

One person
walking

18

0

Standing

39

78

Bedded

39

366

Bedded

before,

during,

and after harassment,

Sam]2ling Period

Deer
Reaction
To stimulus

Stimulus

activities

Deer
Ac t Lv Lt y=

Pre-route
Route
Post-route
HR(BPM) (n)b HR(BPM) (n) HR(BPM) (n)

February

- March,

1977.

% Change HR
Pre-route Pre-route
to
to
Post route
route

Duration
Of HR Change (min}
Pre-route
Pre-route
to
to
post-route
route

S
SG

43.5 (2)
58.1 (2)

88.0 (1)
73.4 (2)

56.0 (1)
59.8 (4)

102
26

29
3

1
2

1
4

alert

B

37.0 (5)

94.1 (1)

39.3 (6)

154

6

1

6

alert

B

36.5 (5)

42.0 (4)

42.2 (5)

15

16

4

5

alert

---------------------------------------------------------------------------------------------------------------------------------------------------Two persons
walking

39

o

Bedded

alert

B

46.3 (5)

83.5 (1)

80.6 (1)

80

74

1

1

18

78

Bedded

alert

B

33.7 (5)

45.8 (2)

37.3 (5)

36

11

2

5

39c

156

Standing alert
and bedded alert

S
B

18

366

Standing

S

48.2 (2)

41.7 (2)

18

78

Bedded

B

38.1 (3)

30.1 (5)

alert

_______________________________________________________
~~
22~~_i~2
Dog plus
person
walking

156

39

366

18

Bedded
Bedded

alert
alert
alert

a See Table I for definitions

of codes.

b Number of one minute

used to compute

c Heart-rate

values

samples

during harassment

:

-

-13

-

2

-

*

32.3 (5)

-21

-15

5

5

*

5
5

~2~~_i~2

:-------:!~-------------:----------~----

B

43.2 (4)

53.1 (5)

42.3 (5)

23

-2

5

B

32.2 (4)

38.1 (2)

29.9 (5)

18

7

2

value •

are compared

to normal

heart-rate

values

I

5

32

58.1 (5)

~
v.&gt;

7

22

79.5 (7)

r
•.....

in Table 2 to compute

% changes

in heart-rate.

�-144-

with the implant was life expectancy.
Implants functioned for 18 to 182
days with a mean of 58 days (n = 8). The short duration of the implants
has been attributed to faulty manufacturing of one electrical component
and not with the design or assembly of the implant.
The M-99 - Rompun combination was excellent for anesthesia.
Deer succumbed
to the drugs usually within 10 minutes and remained in deep anesthesia
until an antidote, M-50-50, was administered.
Deer regained motor control
within 5 minutes after M-50-50 was given. Rompun continued to slightly
sedate the deer for up to 2 hours after surgery but animals were either up
and moving or bedded with their heads up and alert during this time.
LITERATURE

CITED

Freddy, D. J. 1977. Snowmobile harassment of mule deer on cold winter
ranges.
Colo. Div. Wildl., Game Res. Rept., July, Part 1, p. 89-104.
Kautz, M. V. 1978. Energy expenditure and heart rate of active mule
deer fawns. M.S. Thesis.
Colo. St. Univ. 76pp.
Moen, A. N., M. A. Della Fera, A. L. Hiller, and B. A. Buxton.
1978. Heart
rates of white-tailed deer fawns in response to recorded wolf howls.
Canad. J. Zool. 56(5):1207-1210.
Selye, H.

1976.

Stress in health and disease.

Butterworths,

Boston.

1256p.

�July,

1978

-145-

JOB PROGRESS

State of
Project

COLORADO
No.

W-38-R-33

Work Plan No.

16

Job Title

Deer Study-Estimating

Period

REPORT

Piceance

Covered:

Deer-Elk

Investigation

Job No.

1-3

Parameters

of Population

Dynamics

April I, 1977 - June 30, 1978

Personnel: R. M. Bartmann, R. S. Bliss, L. H. Garpenter, C. A. Carron, J. M.
Cribari, J. E. Enote, D. J. Freddy, A. L. Fricke, D. Goddard, R. B. Gill, W.
D. Hadden, H. Hansell, J. J. Klein, Jr., L. M. Lofgren, C. A. Moore, M. C.
Odum, J. L. Sexton, P. F. White, and L. S. Zeise.

ABSTRACT
Forty-six sightings and 27 recoveries of banded deer were recorded during
Segment 33. The 1977-78 winter deer census estimate in Unit 22 was 34,846
± 7,980 deer (90 percent confidence level) compared to the predicted population of 35,915 deer. The buck:doe:fawn ratio, based on 1,791 deer classified
in December, 1977, was 17:100:67.
The post-season ratio is the same as in
1976, but adjustment for antlerless harvest in 1977 yields a slightly lower
pre-season ratio of 64 fawns:100 does. The 1976-77 winter was one of the
mildest on record and only 1,064 ± 638 deer (90 percent confidence level) were
estimated lost over winter which is about 4 percent of the predicted winter
population.
The following winter, 1977-78, was also relatively mild and a
5 percent winter loss was estimated (1,809 + 851 deer at the 90 percent
confidence level). A total deer harvest fo~ all seasons in 1977 was 6,096
deer. This is an 88 percent increase over 1976's estimated harvest due to a
combination of factors including a larger deer population, higher hunter
success, and the insurance of 1,901 either-sex permits.

��-147-

ESTIMATING

PICEANCE DEER STUDYPARAMETERS OF POPULATION

DYNAMICS

R. M. Bartmann

P. N. OBJECTIVE
To develop and test a method for estimating
deer populations in the Piceance Basin.

density of over-winter

mule

SEGMENT OBJECTIVES
1.

To collect and tabulate deer band sighting and recovery data.

2.

To estimate
Unit 22.

the size of the wintering

deer population

in Game Management

3.

To estimate the sex and age structure
in Game Management Unit 22.

of the wintering

deer population

4.

To estimate

the annual productivity

5.

To estimate

the annual winter mortality

6.

To estimate annual hunter harvest

of the Piceance

of the Piceance

of the Piceance

METHODS AND MATERIALS
Deer Banding
See Bartmann

(1972).
Deer Density

See Bartmann

(1974a).
Population

See Bartmann

Structure

(1974a).
Productivity

See Bartmann

(1975).
Winter Mortality

See Bartmann

(1974a and 1975).

deer population.
deer population.

deer population.

�-148-

Hunter Harvest
See Bartmann

(1974b).
RESULTS AND DISCUSSION
Deer Banding

Forty-six sightings and 27 recoveries of banded deer were recorded from
March 1977 through June 1978 (Tables I and 2). In Segment 34, it is planned
to begin the analysis and write-up of the deer distribution and banding data
to be published as a Division Special Report.
Deer Density
The 1977-78 winter deer census occurred February 18-22 which was about four
weeks later than desired due to previously scheduled work that could not be
changed.
As a result, the deer were concentrated on south slopes, which were
mostly snow-free, to avoid the deeper (18-24 inches; 46-61 cm) denser and
partially crusted snow on protected north exposures.
A Hiller 12EJ3 helicopter with a Soloy turbine conversion was used in lieu
of the Bell 47G3B because of the extra power and similar visibility.
Also,
the pilot sat in the middle which allowed both observers to sit on the outside
for maximum visibility, although communication between the two was difficult.
The 1,577 deer counted converts to 13.1 deer per quadrat, 52.4 deer per mile2
(20.2 deer per km2), or 34,846 ± 7,980 total deer (90 percent confidence level)
on the winter range in Unit 22 (Table 3). The precision of the mean estimate
(± 21 percent) is the poorest to date and is attributed largely to the concentrated deer distribution.
This population estimate compares favorably with
the predicted population of 35,915 (Table 4).
Population

Structure

Post-season sex and age classifications were made in Unit 22 December 10-12, 1977.
The 1,791 deer classified consisted of 169 bucks, 970 does, and 652 fawns for a
buck:doe:fawn ratio of 17:100:67.
The fawn:doe ratio is the same as last year's
and the buck:doe ratio is 5 bucks: 100 does lower which is in line with the
predicted ratio of 15:100.
Productivity
The 1977 deer season in Unit 22 was modified from antlered-only to include the
limited taking of antlerless deer through the availability or 3,500 either-sex
permits, of which only 1,901 were issued.
The Game Management Section's random
survey results (inflated arbitrarily to include a 25 percent increase to account
for wounding loss and illegal kill) show a total antlerless harvest for all
seasons of 903 does and 33 fawns. Thus, the post-season fawn:doe ratio of 67:100
converts to a pre-season ratio of 64:100 for a pre-season fawn production of
13,255 animals.

�-149-

Table l. 8ightings of deer marked on the winter range in Game Management
Units 11, 21, 22, and 23, received from March 1977 through June 1978.
Neckband
Twp.

Location
8ection
Range

Yellow wired stripe

IN

93W

12

10-30

White wired stripe

IN

96W

25

12-15

White

18

96W

32

12-17

Red and blue blocks

IN

96W

15

3-?-77

Red and blue blocks

IN

94W

18

3-1

Green and yellow blocks

IN

96W

32

4-?

Red w/white

stripe

28

96W

32

4-?

Red wlwhite

stripe

28

96W

32

4-12

White

216

18

98W

34

4-12

White

229

18

98W

29

4-12

White

240

18

98W

29

4-22

White

93

18

98W

29

5-10

Yellow wired stripe

IN

96W

9

10-17

White

18

lOOW

25

10-19

Green wlwhite

stripe

28

95W

12

10-20

Red and blue blocks

IN

93W

20

10-22

Orange and white blocks

4

28

94W

2

10-22

Orange and white blocks

139

2S

94W

2

10-24

Yellow

101

IS

92W

23

10-28

White

2N

98W

3

11-12

Red and blue blocks

2N

94W

19

11-17

Yellow wired stripe

IS

96W

10

11-28

Green wlwhite

IN

96W

27

12-1

Pink

3S

97W

2

12-10

Green wlwhite

IN

95W

31

12-11

White

IN

97W

21

12-11

Blue and white

28

96W

4

12-12

Pink

2S

96W

5

12-12

White

2N

98W

l3

Date

Color

7-?-76

Number

46

stripe
72
stripe

------------------------------------------------------------------------------

�-150-

Table l. Sightings of deer marked on the winter range in Game Management Units
ll, 21, 22, and 23, received from March 1977 through June 1978. (Continued).
Neckband
Twp.

Location
Range
Section

Green and yellow blocks

lN

99W

17

12-12

Green w/white

stripe

2S

95W

2

12-12

Green w/white

stripe

2S

95W

2

12-12

Green w/white

stripe

lS

95W

2

12-12

Green w/white

stripe

IS

95W

13

12-12

Green w/white

stripe

lS

95W

12

12-12

Green w/white

stripe

lN

95W

33

12-12

Green w/white

stripe

lS

95W

6

12-12

Red w/white

3S

96W

23

12-16

Yellow

27

lN

92W

26

12-16

Red and blue blocks

178

lN

92W

26

2-19-78

Yellow wired

stripe

lN

96W

36

2-20

Red and blue blocks

lN

94W

30

2-22

White

lS

98W

21

2-23

Yellow vT/red stripe

lN

96W

36

2-25

Blue and yellow blocks

12

lN

94W

34

6-4

Yellow

55

lS

91W

3

Date

Color

12-12

Number

stripe

Table 2. Recoveries of deer marked on winter range in Game Management
ll, 21, 22, and 23, received from March 1977 through June 1978.
Neckband

Units

Date

Color

Number

Twp.

Location
Section
Range

4-10-77

Blue

57

2N

9/W

20

4-20

Red and blue blocks

39

2N

94W

18

8-23

White wired

42

3N

98W

34

10-?

Blue

31

2N

98W

5

10-?

Yellow wired

stripe

llO

IS

96W

10

10-?

Red and blue blocks

144

4N

89W

8

stripe

�-151-

Table 2. Recoveries of deer marked on winter range in Game Management Units
11, 21, 22, and 23, received from March 1977 through June 1978.
(Continued).
Neckband
Number

Twp.

Location
Range
Section

7

IS

95W

32

68

4S

97W

21

L-3283

IS

92W

12

Yellow wired stripe

102

4S

95W

15

10-15

Green w/white

156

IN

96W

32

10-15

White wired stripe

69

2N

98W

24

10-15

Red and blue blocks

27

3N

95W

25

10-16

Red and blue blocks

96

IN

95W

14

10-16

Orange and white blocks

102

IS

91W

13

10-17

Pink

75

2S

96W

32

10-24

Blue

107

3N

99W

34

11-?

Blue

109

2N

99W

8

11-?

White

?

IN

98W

19

11-9

Green and yellow blocks

2

2N

99W

9

11-9

Red w/white

48

2S

96W

15

12-?

Green w/white

168

IS

94W

19

12-9

Blue and White

51

28

97W

25

3-30-78

Blue

103

2N

98\-1

5

4-28

Blue and white

101

IS

96W

18

5-?

Blue

97

IS

97W

12

5-1

Yellow wired stripe

46

2N

93W

15

Date

Color

10-?

Green w/white

10-?

Red w/white

10-15

Green w/white

10-15

stripe

stripe
stripe

stripe

stripe
stripe

2
quadrats
Table 3. Number of deer counted on 120 ~-mile
range in Game Management Unit 22, February 18-22, 1978.

on the Piceance

Quad.

Quad.

Deer

Quad.

Deer

Quad.

Deer

winter

Deer
- .. -.- ..~.-.

1

15

31

22

61

0

91

0

2

9

32

65

62

20

92

0

3

3

33

81

63

31

93

27

4

9

34

7

64

19

94

0

5

87

35

41

65

14

95

13

------------------------------------------------------._------------------------

�-152-

Table 3. Number of deer counted on 120 ~-mile2 quadrats
range in Game Management Unit 22, February 18-22, 1978.

on the Piceance winter

Quad.

Deer

Quad.

Deer

Quad.

Deer

Quad.

Deer

6

27

36

30

66

71

96

0

7

6

37

10

67

8

97

0

8

2

38

40

68

28

98

0

9

0

39

0

69

11

99

4

10

0

40

0

70

4

100

0

11

0

41

0

71

0

101

0

12

0

42

0

72

0

102

2

13

0

43

40

73

13

103

10

14

0

44

0

74

27

104

3

15

19

45

8

75

0

105

3

16

42

46

21

76

3

106

23

17

29

47

0

77

0

107

3

18

31

48

3

78

28

108

63

19

0

49

0

79

0

109

6

20

12

50

8

80

0

110

0

21

16

51

0

81

0

III

56

22

28

52

34

82

17

112

17

23

1

53

0

83

0

113

8

24

72

54

0

84

0

114

3

25

39

55

0

85

13

115

0

26

87

56

0

86

0

116

0

27

19

57

0

87

9

117

4

28

0

58

0

88

30

118

23

29

0

59

0

89

0

119

0

30

0

60

0

90

0

120

0

90% Confidence

Interval

n =

120

L:X= 1,577

13.1 + (1.658) (1.82) = 13.1 + 3.0

x =

13.1

s =
sx =

19.96

Total POEulation

1.82

34,846

C.V.=

152%

± 7,980

Estimate

�-153-

Table 4. Calculations to predict
Game Management Unit 22.

the 1977-78 winter

deer population

Bucks

Does

Fawns

Total

3,459

15,775

10,586

29,820

489

287

288

1,064

1977 summer population

2,970

15,488

10,298

28,756

Fawns apportioned

5,149

5,149

8,119

20,637

Est. 1976-77 winter

pop.

Est. 1976-77 winter mortality

Adjusted

50:50

1977 summer pop.

1977 fawn prod. est.
(64 fawns: 100 does)

1977 pre-season
1977 harvest

20,637

13,255

42,011

estimate

5,160

903

33

6,096

2,959

19,734

13,222

35,915

3,304

19,538

13,073

35,915

Predicted

population

post-season
ratio

13,255

8,119

1977-78 winter

corrected

28,756

pop.

Predicted

in

pop.

for 1977
buck:doe:fawn

(17:100:67)

Winter Mortality
The 1977 deer winter mortality survey in Unit 22 was made April 19 through May
23. Only 11 carcasses were found for a mean 0.20 dead deer per plot or 1,064
638 dead deer (90 percent confidence level) on the winter range.
This is
nearly a 4 percent loss of the predicted 1976-77 winter population and is the
lowest mortality percentage of the study.

±

The 1976-77 winter was the mildest
1973. Because of this, an unknown

since the mortality work was started in
proportion of the deer population wintered

�-154-

above the area sampled for mortality which could have contributed to a
lower estimate.
However, it is equally logical to assume the decreased
mortality estimate is real and reflects the milder winter conditions of
1976-77.
The few carcasses found (1 male and 2 fawns of unknown sex, 4
adult males, 2 adult females and 2 adults of unknown sex) makes the
projection to total mortality by sex and age class even more hazardous than
usual.
Nevertheless, the estimated composition of the total loss is 144
male fawns, 144 female fawns, (fawn mortality was assumed to be 50 percent
male fawns and 50 percent female fawns because of the extremely small sample
of dead fawns encountered) 489 adult males and 287 adult females.
The 1977-78 winter was also relatively mild although there was more snow
than in the previous winter.
The mortality survey, conducted April 17 through
May 18, 1978, yielded 19 carcasses comprised of 1 male, 6 female and 6 fawns
of unknown sex, 4 adult females and 2 adults of unknown sex. The mean number
of carcasses per quadrat, 0.34, projects to 2.72 per mile2 (1.05 per km2) or
1,809 ± 851 dead deer (90 percent confidence level) on the Piceance winter
range which is about a 5 percent loss of the estimated winter population.
Projected composition of the dead deer is 619 male fawns, 618 female fawns,
96 adult males and 476 adult females.
Hunter Harvest
Restructuring of the 1977 big game seasons resulted in a deer-only hunt
October 14-19 and a combined deer and elk hunt November 3-15. The basic
season was antlered-only with an additional 3,500 either-sex permits
available in Unit 22. Only 1,901 of these permits were issued.
The archery season ran September 3-25 and was either-sex.
A muzzle-loader
season was also held September 10-18 and was antlered-only.
The total buck harvest, according to random survey results, was up nearly
60 percent over last year's (Table 5). This is partly due to the extra season
but also reflects slightly higher hunting success, 59 percent in 1976 and
70 percent in 1977 for antlered-only hunters.
Addition of either-sex permit
holders only raises the 1977 success one percent.
Permit holders accounted for 918 antlerless deer of which 33 were fawns.
The estimated fawn take is half of that checked at Idaho Springs suggesting
reporting bias, sampling error or both in survey results, although confidence
intervals were not provided for harvest data.
The percentage of yearlings in the check station-aged buck harvest, 59 is
similar to the 62 percent in1976 from three "regular" stations (Table 6).
The female yearling percentage, 32, cannot be compared since there was no
female harvest in 1976. Predicted yearling percentages were 63 and 25 for
bucks and does, respectively.

�-155-

Table 5.

Summary of the 1977 deer harvest

Season

Bucks
Adult
Fawns

Archery

50

Muzzle-loader

1/

in Unit 22 for all seasons.Does
Adult

Fawns

Total

0

18

0

68

3

0

0

0

3

2,455

0

0

0

2,455

596

19

589

14

1,218

1,858

0

0

0

1,858

198

0

296

0

494

5,160

19

903

14

6,096

Split
regular
either-sex
Combined
Regular
either-sex
All

l./All figures include and arbitrary
wounding loss and illegal kill.
LITERATURE

inflation

of 25 percent

to acknowledge

CITED

Bartmann, R. M. 1972. Piceance deer study-population distribution.
P. 315337. In Game Research Report.
Colo. Div. of Wildl., Denver.
3(Part 3):
253-377.
(Proc.).
Bartmann, R. M. 1974a. Piceance deer study-population density and structure.
P. 363-370.
In Game Research Report. Colo. Div. of Wildl., Denver.
3(Part 2):185-398.
(Proc.).
Bartmann, R. M. 1974b. Piceance deer study-productivity
and mortality.
P. 371-380.
In Game Research Report.
Colo. Div. of Wildl., Denver.
3(Part 2):185-398. (Proc.).
Bartmann, R. M. 1975. Piceance deer study-productivity
and mortality.
371-380.
In Game Research Report.
Colo. Div. of vlildl., Denver.
3(Part 2):195-504.
(Proc.).

P.

�Table 6. Deer checked at Idaho Springs check station from the 1977 separate and combined seasons
in Unit 22.

Season

Mature

Bucks
Yrlg.

Fawn

Unk.

Mature

1/
Separate--

166

298

28

84

128

Combined11

98

82

7

49

49

Does
Yr1g. Fawn

Unk.

Unk.
Unk.

Total

64

23

28

54

873

19

8

21

15

348

11 An additional 56 deer, checked during the separate elk season and presumed killed during the
separate deer season, are omitted as an age breakdown is not available.

1/ Age data for deer harvested in the combined season are not available. The figures used are
the differences between deer checked in the separate season and total deer checked for all seasons.
I
•.....
l.r1
C"'

I

Prepared

by&amp;d~tn~~
Richard M. Bartmann
Wildlife Researcher

�July, 1978
-157-

JOB PROGRESS

State of

REPORT

COLORADO
--------------~~~---------

Project No.

W-38-R-33

Deer-Elk

Investigations

Work Plan No.

16

Job No.

5

--------------------------------Job Title
Piceance Deer Study-Food Habits Technique Development
------------------~--------~~~~~~~~~~~------Period Covered: April 1, 1977 - June 30, 1978

Personnel: A. W. Alldredge, R. M. Bartmann, J. M. Cribari, A. L. Fricke, J. J.
Klein, Jr., L. M. Lofgren, P. H. Neil, M. L. Stevens, and L. S. Zeise.

ABSTRACT
Fourteen fawns were reared at Little Hills in 1977 and 2 more were picked
up as orphans later in the summer. Four did not adapt to a change in
handlers and were sent to Fort Collins for use in other studies.
Another
was killed by coyotes in April, 1978. There were 260 different plant species
collected and identified taxonomically in the Piceance Basin in 1977. Two
8-day tame deer grazing evaluations were run in 1978. The "pen" treatment
group took noticeably more bites of grass than the "pasture" group both times.
This is presumed a prior-conditioning response as they had been previously
used in predominantly grassland situations.
Otherwise, with few exceptions,
the major species eaten by one group were similar to those eaten by the other
group.

��-159-

PICEANCE

DEER STUDY-FORAGE

PREFERENCE

R. M. Bartmann
P. N. OBJECTIVE
To estimate forage preferences
in the Piceance Basin.

of mule deer on pinyon-juniper

winter

range

SEGMENT OBJECTIVES
1.

Analyze data from food habits
pertinent results.

technique

development

2.

Prepare a detailed study plan for estimating
preferences of deer in the Piceance Basin.

3.

Maintain

4.

Select specific

5.

Prepare

6.

Conduct deer foraging

7.

Rate forage availability

8.

Test necessity

9.

Tabulate

work and publish

fall-winter-spring

forage

and train tame deer.
sites for food habits

an herbarium

sampling.

of plants from the Piceance

Basin.

trials.
during deer foraging

of converting

data and prepare

trials.

bite data to a weight basis.

annual progress

report.

The tame deer grazing evaluations schedule in Segment 32 were only partially
completed due to intractable deer.
The lack of snow cover also created
extremely atypical winter range conditions.
Therefore, in lieu of the food
habits work planned, it was deemed necessary to repeat the evaluations in
Segment 33. In addition, the lack of enough suitably trained deer for the
scheduled food habits work prompted the raising of fawns in 1977, and they
will not be ready for use under Piceance winter range conditions until fall,
1978. In view of the preceding, Segment 32 objectives (Bartmann 1977) are also
applicable in Segment 33.
METHODS

AND MATERIALS

Fawn Rearing

and Training

Ten fawns were obtained from wild does kept in pens at Little Hills, 2 were
obtained from does in the Fort Collins pens, 2 were picked up as orphans
shortly after birth, and 2 were picked up in late August after they were
hand-reared by other people.
Procedures for hand-rearing followed Halford
(1974) with various modifications as deemed necessary for individual deer.

�-160-

Plant Collecting
A major effort was made to develop an herbarium of plant species in the
Piceance Basin with emphasis on the proposed deer food habits study areas.
A temporary employee worked from May 23 thru October 14 on this job. All
specimens were submitted to Dr. W. A. Weber, Curator of the University of
Colorado Herbarium in Boulder for confirmations of taxonomic identifications.
Tame Deer Grazing Evaluations
Evaluations of "tame" versus "wild" deer forage selections were made in late
January and mid-March, 1978. Several procedure changes were made based on
assessment of the 1977 grazing work (Bartmann 1977).
Five deer, one a spare, were brought from the Fort Collins pens November 15,
1977 and placed in Pasture 116 to serve as the "pasture" treatment group.
The deer were all 2~-years old and had been used in grazing work at Little
Hills the previous 2 winters.
Supplemental feed was provided the first week
but was largely ignored.
Afterwards, a few handsful of grain were provided
several times a week to make the deer easier to locate and to check on their
condition.
Four tame deer, borrowed from the Department of Radiology and Radiation
Biology, Colorado State University, served as the "pen" treatment group. Three
were 2~-years old and one l~-years old. All had been used previously in grazing
work at the Rocky Flats Nuclear Weapons Plant, which is predominately a grassland plant community.
About a week prior to both the January and March grazing evaluations, the
"pasture" deer were moved to Pasture 115 to become familiar with vegetation and
physiography.
The "pen" deer were allowed one day to roam freely in Pasture
116, and thereafter were kept in a small pen in the pasture when they were not
being grazed.
The 2 groups were grazed in their respective pastures for 4
consecutive and coincidental day grazing periods.
Pastures were then switched
and each group allowed another day of familiarization followed by 4 more days
of grazing observations.
As in Pasture 116, a small pen in Pasture 115 served to
contain the "pen" deer when they were not being grazed.
"Pen" deer were fed token
daily rations of grain supplement.
Four observers
recorded bite-count data during morning and afternoon periods
each day. Two watched 2 deer from the "pen" group while the other 2 watched
2 deer from the "pasture" group for a total of 4 trials per group each day, or
32 total trials per group for each 8-day evaluation period.
Individual deer were grazed an equal number of morning and afternoon periods
in each pasture and observers were switched among deer both within and between
treatment groups to minimize biases from these sources.

�-161-

RESULTS
Fawn Rearing

and Training

Fourteen fawns were hand-reared of which 4 did not adapt to a change in
handlers.
They were taken to Fort Collins for use in other studies.
Training for food habits work has proceeded with 12 deer, including 2 orphans
picked up in late August.
Training included frequent trips to nearby
winter range to acquaint the deer with native vegetation.
In April, 1978,
all the deer were moved to Pasture #5 where they can graze native vegetation
yearlong.
They have been kept in a small pen in the pasture at night after
one was killed by a coyote.
Trips to summer range areas were started in
the summer, 1978.

Plant

Collecting

The 260 plant species collected during the summer of 1977 are listed in
Table 1. Plant collections will be continued on a time-permitting
basis
future Segments.

Tame Deer Grazing

in

Evaluations

Snow depths during the January, 1978 grazing evaluations ranged from 12 to
15 inches, with morning and afternoon temperatures ranging from 20 to 300F
(-6.7 to 1.10C), respectively.
In March, the ground was mostly bare but
snow accumulations up to 12 inches occurred in protected areas.
Morning and
afternoon temperatures ranged from 30 to 400F (-1.1 to 4.4°C). respectively.
A major difference in forage selections of the 2 deer treatment groups
occurred in the graminoid category (Tables 2 and 3). In January, graminoids
comprised 32 percent of the "pen" group's total bites and almost none of the
"pasture" group's.
In March, graminoid percentages were 26 and 15 for "pen"
and "pasture" deer, respectively.
In spite of deep snow in January, the "pen"
deer would seek exposed grass under trees and shrubs.
These deer had been
used exclusively in grassland types at Rocky Flats where grasses averaged 5S
percent of forage consumption, by weight, in winter and 41 percent in spring
(Arthur 1977).
Apparently, this conditioning to grass forage carried over tc
the Little Hills area which suggests a potential bias in the use of tame deer
to collect forage preference data in areas with noticeably different forage
compositions from those areas where the deer were first allowed to forage.
Within the browse category, differences were apparent in proportions of total
bites by species, but as in the past, major species eaten by one group w\:'r.essentially the same as those eaten by the other group.
The one exc.ep t i on I.V&lt;l!='
oakbrush which was taken in considerably greater quantities by "pen" than
"pasture" deer in January and, to a lesser extent, in March.
Any conclusions,
however, will have to await completion of statistical anlayses of the data.

�-162-

Table 1.
1977 .

Plant species

collected

in the Piceance

Basin ,during summer,

Trees and Shrubs
Acer negundo
Amelanchier utahensis
Artemisia tridentata
Atriplex canescens
Atriplex confertifolia
Atriplex obovata
Ceanothus martinii
Ceratoides lanata
Cercocarpus intricatus
Cercocarpus montanus
Chrysothamnus
depressus
Chrysothamnus
linifolius
Chrysothamnus nauseosus
Chrysothamnus
viscidiflorus
Cratageus saligna
Ephedra viridis
Holodiscus dumosus
Juniperus scopulorum
Juniperus osteosperma

Mahonia repens
Pachystima myrsinites
Pinus edulis
Pinus ponderosa
Populus angustifolia
Prunus virginiana
Pseudotsuga menziesii
Purshia tridentata
Quercus gambellii
Ribes aureum
Ribes cereum
Rhamnus smithii
Rhus trilobata
Rosa woodsii
Sarcobatus vermiculatus
Shepherdia argentea
Symphoricarpos
oreophilus
Tamarix pentandra
Tetradymia canescens

Graminoids
Agropyron dasystachyum
Agropyron desertorum
Agropyron elongatum
Agropyron inerme
Agropyron repens
Agropyron riparian
Agropyron smithii
Agropyron spicatum
Bouteloua gracilis
Bromopsis ciliata
Bromopsis inermis
Bromus tectorum
Carex pityophila
Ceratochloa marginata

Elymus cinereus
Elymus salinus
Hordeum jubatum
Koeleria macrantha
Oryzopsis hymenoides
Oryzopsis micrantha
Phragmites australus
Poa agassizensis
Poa fendleriana
Poa nemoralis interior
sitanion hysterix
sitanion longifolium
Stipa comata
Stipa occidentalis

Forbs
Achillea lanulosa
Agoseris aurantiaca
Allium acuminatum
Alyssum desertorum
Amaranthus albus

Amaranthus retroflexus
Androsace septentrionalis
Antennaria parvifolia
Antennaria pulcherrima
Antennaria rosea

�-163-

Table 1. Plant species collected
1977 . (Cont' d) .

in the Piceance

Basin during summer,

Forbs
Aquilegia barnebyi
Arabis divaricarpa
Arabis drummondii
Arabis lignifera
Areneria eastwoodiae
Artemisia biennis
Artemisia dracunculus
Artemisia frigida
Artemisia ludoviciana
Asclepias speciosa
Asclepias subverticilliata
Aster adscendens
Astragalus chamaeleuce
Astragalus diversifolius
Astragalus lutosus
Astragalus oopho:us
Astragalus purshii
Astragalus spatulatus
Astragalus tenellus
Atriplex argentea
Atriplex rosea
Balsamorhiza sagittata
Barbarea orthoceras
Brickellia grandiflora
Calochortus nuttalli
Campanula rotundifolia
Capsella bursa-pastoris
Castilleja chromosa
Castilleja linariaefolia
Caulanthus crassicaulis
Chaenactis douglassi
Chamaesyce fendleri
Chenopodium fremontii
Chenopodium leptophyllum
Cirsium arvense
Cirsium undulatum
Clematis hirsutissima
Clematis ligusticifolia
Cleome serrulata
Collinsia parviflora
Collomia linearis
Comandra umbellata
Conioselinum scopulorum
Conyza canadensis
Corydalis aurea

Crepis acuminata
Cryptantha bakeri
Cryptantha sericea
Cymopterus fendleri
Cynoglossum officinale
Delphinium melsonii
Descurainia californica
Descurainia richardsonii
Descurainia sophia
Epilobium adenocalllon
Erigeron eatonii
Erigeron englemannii
Erigeron speciosus macranthus
Eriogonum alatum
Eriogonum brevicaule
Eriogonum cernuum
Eriogonum effusum
Eriogonum lonchophyllum
Eriogonum ovalifolium
Eriogonum umbellatum
Erysimum asperum
Euphorbia esula
Euphorbia robusta
Frasera speciosa
Galium coloradoensis
Gayophytum diffusum
Gilia pinnatifida calcarea
Grindelia squarrosa
Hackelia floribunda
Halogeton glomeratus
Hedeoma drummondii
Hedysarum boreale
Helianthella uniflora
Helianthus annuus
Heliomeris multiflora
Heracleum sphondylium montanum
Heterotheca villosa
Heuchera parviflora
Hymenopappus filifolius megacephalus
Hymenoxys acaulis
Hymenoxys richardsonii
Ipomopsis aggregata
Ipomopsis congesta
Iva xanthifolia
Kochia iranica

�-164-

Table 1. Plant species
1977 • (Cont' d) •

collected

in the Piceance

Basin during

summer,

Forbs
Kuhnia rosmarinifolia
chlorolepis
Lactuca serriola
Lactuca tatarica pulchella
Lappula redowskii
Lepidium densiflorum
Lepidium montanum
Lepidium perfoliatum
Leptodactylon pungens
Lesquerella montana
Lesquerella subumbellata
Leucelene ericoides
Linaria vulgaris
Linum lewisii
Lithospermum ruderale
Lomatium grayii
Lomatium orientale
Lomatium simplex
Lupinus caudatus
Machaeranthera
grindelioides
Marrubium vulgare
Melandrium dioicum
Melilotus officinalis
Mentzelia albicaulis
Mentzelia dispersa
Mentzelia humilis
Mentzelia multiflora
Mentzelia rushbyi
Mertensia ciliata
Mertensia fusiformis
Microseris nutans
Microsteris gracilis
Moldavicia parviflora
Nicotiana attenuata
Oenothera caespitosa
Oenothera coronopifolia
Oenothera lavandulaefolia
glandulosa
Oenothera pallida
Opuntia polycantha
Orobanche fasciculata
Oxybaphus linearis
Oxytropis lambertii
Pediocactus simpsonii
Penstemon caespitosus
Penstemon fremontii
Penstemon osterhoutii
Penstemon strictus
Penstemon teucrioides

Penstemon watsoni
Phacelia glandulosa
Phacelia heterophylla
Phlox hoodii
Phlox longifolia
Phlox multiflora
Physaria acutifolia
Polanisia dodecandra
Polemonium caeruleum
Polygonum aviculare
Polygonum douglasii
Portulaca oleracea
Pulsatilla patens
Ranunculus cymbalaria
Rollinsia mUlligan
Rumex crispus
Rumex salicifolius
Salsola iberica
Schoenocrambe
linifolia
Senecio eremophilus
Senecio integerrimus
Senecio multilobatus
Senecio spartoides
Senecio wootonii
Sisymbrium altissimum
Sisymbrium loeseli
Smilacina stellata
Solanum triflorum
Solidago sparsifolia
Sphaeralcea coccinea
Stanleya pinnata
Stellaria jamesiana
Stenotus acaulis
Stephanomeria
tenuifolia
Streptanthus cordatus
Taraxicum officinale
Thelypodiopsis elegans
Thlaspi montanum
Tragopogon dubius
Trifolium gymnocarpon
Urtica dioica gracilis
Verbascum thapsus
Vicia americana
Viola adunca
xanthium strumarium
Xanthocephalum
sarothrae
Zygadenus venenosus gramineus

�-165-

Table 2. Summary of bite-count
January, 23-31, 1978.

data from tame deer grazing

Total Bites
Pasture

Species

Pen

evaluations,

Percent of Total Bites
Pasture
Pen

Browse
Amelanchier utahensis
Purshia tridentata
Cercocarpus montanus
Artemisia tridentata
Symphoricarpos
oreophilus
Pinus edulis
Chrysothamnus
viscidiflorus
Ceratoides lanata
Quercus gambellii
Tetradymia canescens
Juniperus osteosperma
Juniperus scopulorum
Chrysothamnus nauseosus
Ribes cereum
Prunus virginiana

Total Browse

25,887
13,836
5,474
2,169
522
191
130
125
112
105
49
28
23
5

5,867
2,279
1,724
1,890
337
600

o
79
3,138
39
91
4
34

o

o

48,656

53.1
28.4
11.2
4.4
1.1
0.4
0.3
0.3
0.2
0.2
0.1
0.1
tr.
tr.

24.5
9.5
7.2
7.9
1.4
2.5
0.3
13.1
0.2
0.4
tr.
0.1
tr.

5

16,087

99.7

o

tr.
tr.
tr.
tr.
tr.
tr.

67.2

Forbs
Artemisia

Unknown

frigida

forb

Linum lewisii
Cryptantha sericea
Artemisia ludoviciana
Eriogonum sp.
Chenopodium sp.

21
13
7
2
2
1

22

o
o

0.1

o

68
6
8

46

104

0.1

0.4

Oryzopsis .hymenoides /
1
and St~pa comata Agropyron inerme
Poa sp.
Sitanion hysterix
Koeleria macrantha

27

3,264

0.1

13.6

30
21
3

0.1
tr.
tr.

Unknown

13

2,620
1,551
170
107
31

o

5

10.9
6.5
0.7
0.4
0.1
tr.

82

7,748

Total Forbs

0.3
tr.
tr.

Graminoids

grass
Bromopsis sp.
Total Graminoids

l/ Oryzopsis
identification

o

and Stipa comata are combined
of the two species.

hymenoides

tr.

0.2

as errors occurred

32.4

in

�-166-

Table 3. Summary of bite-count data from tame deer grazing evaluations,
March 13.21. 1978.

Species

Total Bites
Pasture

Pen

Percent of Total Bites
Pasture
Pen

Browse
Purshia tridentata
Amelanchier
utahensis
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus
viscidiflorus
Symphoricarpos
oreophilus
Juniperus osteosperma
Pinus edulis
Ceratoides lanata
Mahonia repens
Ribes cereum
Quercus gambellii
Tetradymia canescens
Juniperus scopulerum

Total Browse

23,803
10,447
1,439
1,366
363
136
102
58
55
47
45
42

o
o

2,337
9,228
1,355
526
2

212
64
50
15

o
o
868
84

50.3
22.1
3.0
2.9
0.8
0.3
0.2
0.1
0.1
0.1
0.1
0.1

11.2
44.2
6.5
2.5
tr.
1.0
0.3
0.2
0.1

2

4.2
0.4
tr.

37,903

14,743

80.0

70.6

865
641
439
205
61
53
53
40
38
36
28
27
23
11

61
15
304
59

1.8
1.4
0.9
0.4
0.1
0.1
0.1
0.1
0.1
0.1
0.1
0.1
tr.
tr.
tr.
tr.

0.3
0.1
1.5
0.3

Forbs
Artemisia frigida
Xanthocephalum
sarothrae
Phlox hoodii
Machaeranthera
grindelioides
Penstemon caespitosa

Moss
Liverwort
Unknown forb
Eriogonum sp.
Artemisia ludoviciana
Cryptantha sericea
Stenotis acaulis
Heterotheca villosa
Linum lewisii
Cirsium sp.
Physaria acuminata
Senecio multilobatus

Total Forbs

2
2

o
2,524

o
150

o
10
2

42

o
27

o
o
o
o

tr.
tr.
0.2
0.1

tr.

6

676

0.7

5.3

3.2

�-167Table 3. Summary of bite-count
March 13, 21, 1978.
(Cont'd).

data from tame deer grazing evaluations,

Total Bites
Pasture

Species

Pen

Percent of Total Bites
Pasture
Pen

Graminoids

11

4,320
1.235
824

2.641
857
1,793

9.1
2.6
1.7

12.6
4.1
8.6

116
52
0
10
0

0.6
0.3
0.1
0.1
tr.

0.6
0.2

gracilis.

280
146
70
63
4

Total Graminoids

6,942

5,469

14.7

26.2

Poa sp. Agropyron inerme
Oryzopsis hymenoides
21
and Stipa comata Sitanion hysterix
Koeleria macrantha
Bromus tectorum

Unknown

grass

Bouteloua

II Some Carex
11 Oryzopsis
identification

tr.

sp. is included with Poa sp. due to errors in identification.
and Stipa comata are combined as errors occurred
of the two species.

hymenoides

LITERATURE

in

CITED

Arthur, W. J., III. 1977. Plutonium intake by mule deer at Rocky Flats,
Colorado.
M.S. Thesis.
Colo. State Univ., Ft. Collins.
123pp.
Bartmann, R. M. 1977. Piceance deer study - food habits technique development.
P. 117-123.
In Game Research Report.
Colo. Div. of Wildl., Denver.
3(Part 1):1-123.
(Proc.).
Halford, D. K. 1974. A method for artificially ralslng mule deer fawns.
M.S. Thesis.
Colo. State Univ., Fort Collins.
21pp.

Prepared by

~Ck4J
hJ· ~.~
ichard M. Bartmann
Wildlife

Researcher

��July,

1978

-169JOB PROGRESS

REPORT

State of

Colorado
--------~~~~~~----------------

Project

No.

W-38-R-33

Deer-Elk

Investigations

Work Plan No.
17
2
Systems Modeling Big Game Populations:
Simulations of the Carrying
Job T it Le
C=a=_p~a_=c::cic::t'_Ly~o:..:f'___'t:::.:h~e:=:.._R=o_=c~k:.Ly~M::::o:::u:::n~t::.:
n~:::N_=a:.::t-=i:.:o:..:n::::a:::l~P::..:a=_r=_k=
__
Period

Covered:

Personnel:

April

1, 1977 - June 30, 1978

D. L. Baker, N. T. Hobbs, D. G. Milchunas, D. W. Reichert, G. D
Bear, M. Campion, R. B. Gill, P. H. Neil, D. R. Stevens, G. Wagner,
D. M. Swift, J. E. Ellis, L. Stevens.

ABSTRACT
Composition and quality of diets of tame elk as well as relative abundance of
major food items were examined on winter and summer ranges in Rocky Mountain
National Park during the 1977-78 segment.
As reported in Hobbs and Baker (1977),
during the 1976-77 diet sampling period, graminoids were the most frequently
chosen forages during both winter and summer, accounting for over 60 percent of
observed bites.
Mean browse consumption was greater during winter (~± 90 percent
confidence interval = 32.7 ± 7.9) than summer (18.0 percent ± 7.5) and increased
from 17.3 (± 10.7) percent in November to 32 (±7.9) percent in March.
Mean crude
protein content of summer diets was 12.6 (±3.8) percent, while winter diets contained an average of 5.4 (±0.31) percent protein.
Mean in vitro digestible dry
matter (IVDDM) of winter and summer diets were 42.7 (±2.8) and 51.5 (±7.4) percent,
respectively.
Crude protein content and IVDDM of the diet declined acutely with
advancing season from a high in midsummer to a low in late winter.
During winter,
decline in protein content of grass was correlated with an increased proportion
of browse (leaves and stem) in the diet. As browse increased in the diet IVDDM
decreased.
Results of the 1977-78 winter sampling indicate nearly equal percentages of
graminoids and browse throughout winter.
Mean graminoid and browse ingestion were
49.81 C±8.51) and 44.36 C±8.82) percent, respectively.
Eleven plant species
accounted for 81.43 percent of the 132,152 bites observed in the composite diet
totaled across animals, months and plant communities.
Summer food habits sampling
is currently being conducted.
During September, 1977, total herbage biomass was estimated for seven vegetation
communities comprising the Rocky Mountain National Park winter range.
Biomass
was estimated within 15 percent of the mean with 90 percent confidence.
Estimates
from all plant communities ranged from a low of 33.11 (+4.38) grams per square
meter (333.1 + 43.8 kg/ha) in ponderosa pine-shrub to a-high of 179.07 (+19.71)
(1,790.7 ± 197.1 kg/ha) in willow vegetation type. Reasonably precise e~timates
of the more COTIllil0n
species within each vegetation type were also calculated.
Evaluation of elk forage quality for both the 1976-77 and 1977-78 sampling
periods is currently in progress.

��-171SYSTEMS MODELING BIG GAME POPULATIONS:
SIMULATIONS OF THE CARRYING CAPACITY OF THE
ROCKY MOUNTAIN NATIONAL PARK ELK WINTER RANGE
D. L. Baker and N. T. Hobbs
PROGRAM

NARRATIVE

OBJECTIVE

To develop and test a model simulating carrying capacity of elk winter
using Rocky Mountain National Park as a study population.
SEGMENT

OBJECTIVES

1.

To determine what plant species on Rocky Mountain National
elk winter and summer ranges constitute elk food.

Park

2.

To estimate relative quantities
winter diets of elk.

in the

3.

To examine
preference

the relationship between
for forage plants.

4.

To assess
available

the quantity, chemical content,
to elk wintering in RMNP.

5.

To develop

and test an elk carrying
METHODS

ranges

of individual

plant

nutritional

species

quality

and elk

and digestibility

capacity

simulation

(RMNP)

of foods

model.

AND MATERIALS

Methods and materials employed in the winter carrying capacity study have been
previously described (Hobbs and Baker 1977). Detailed methodology and study
area description for the summer experiment are presented in the enclosed study
plan (Appendix 1) "Quantitative and Nutritional Description of the Summer Diet
Selection of Elk in Rocky Mountain National Park" prepared by D. L. Baker.
RESULTS
Food Habits

AND DISCUSSION
Sampling

1976-1977

Winter diet sampling was conducted for 10 days each month from November through
March.
Fifteen plant species accounted for 86.2 percent of the 155,250 bites
observed in the composite diet totaled across animals, months, and plant communities (Table 1). Summer diet sampling was done 3 days each month from July
to September.
Ten species made up 87.3 percent of the 57,846 total bites (Table 2).
Winter

diets during this sampling period were dominated by bluegrass (Poa
willow leaves (Salix spp.), mountain muhly (Muhlenbergia montana)
aspen leaves (Populus tremuloides) and timothy (Phleum pratense).
Important
summer forages included sedges (Carex spp.), leaves, stems and inflorescences
of willow (Salix brachycarpa, Salix planifolia), tufted hairgrass (Deschampsia

pratensis),

�-172Table 1. Mean percentage composition with 90 percent confidence intervals
(C.l.) across 5 animals and all vegetation types during winter 1976-77
in Rocky Mountain National Park (RMNP). Species listed are those which
contribute 2 percent or more of total bites.
Mean
Percent

90 Percent
C. r ,

Poa pratensis

24.5

+4.5

Salix spp . (leaves)

8.6

+3.4

MUhZenbergia

8.5

+5.3

7.2

+3.2

Carex spp.

5.7

+1.9

phZeum pratense

5.1

+1. 7

PotentiZZa

5.0

+2.3

4.2

+0.8

Bromus inermis

3.3

+1.1

Salix spp. (stems)

3.3

+1.7

Eriogonum

3.2

+1.1

Stipa comata

2.6

+1.7

Purshia tridentata

2.6

+1.3

Species

montana

PopuZus tremuZoides

(leaves)

fruticosa

CaZamagrostis

canadensis

umbeZZatum

Table 2. Mean percentage composition with 90 percent confidence
intervals (C.r.) for 1977 summer diets of elk in Rocky Mountain
National Park.
Species listed are those which contribute 2 percent
or more of total bites.
MEAN
PERCENT

90 PERCENT
C. t ,

Caretc spp.

43.2

+20.1

Salix pl.ani fol-ia

8.1

+4.5

SaZix brachycarpa

6.4

+4.9

SPECIES

-------------------------------------------------------------------------

�-173-

Table 2. Mean percentage composition with 90 percent confidence intervals
(C.l.) for 1977 summer diets of elk in Rocky Mountain National Park. Species
listed are those which contribute 2 percent or more of total bites (Continued).
Mean
Percent

90 Percent

6.0

+6.7

6.0

+4.5

Geum rossii

4.2

+3.2

Kobresia

3.9

+2.0

3.8

+2.9

spp.

2.9

+1.7

leptosepala

2.8

+3.1

Species

Juncus

balticus

Deschampsia

caespitosa

myosuroides

Calamagrostis
Vaccinium
Caltha

canadensis

C.1.

and alpine avens (Geum rossii).
Mean diet percentage and confidence
intervals for individual species in all summer vegetation communities have not
been calculated at this time. Percentage shown in Tables 3-5 are calculated
from the total bites observed in each vegetation type.
caespitosa),

Table 3. Percentage composition of elk diet acroSs all animals in willowpark vegetation type during summer 1977 in Rocky Mountain National Park.

July

August

September

spp.

51.72

42.58

46.28

Salix planifolia

17.03

10.39

14.88

Juncus balticus

2.72

13.05

11.72

spp.

9.19

6.93

8.91

Salix brachycarpa

4.61

8.87

1.26

Deschampsia

4.87

1.91

4.44

2.90

2.82

2.83

Species

Carex

Vaccinium

Caltha

caespitosa

leptosepala

�-174-

Table 3. Percentage composition of elk diet across all animals in willowpark vegetation type during summer 1977 in Rocky Mountain National Park.
(Continued).
Species

July

August

September

.75

3.48

1.85

o

o

3.70

spp.

.10

2.84

.32

% Grass

60.97

65.62

66.93

Forb

8.02

7.94

5.61

% Shrub

31.01

26.43

27.46

TOTAL BITES

7245

4544

6262

Calamagrostis
Bromus
Poa

%

canadensis

inermis

Table 4. Percentage composition of elk diet across all animals in krummholzecotone vegetation type during summer 1977 in Rocky Mountain National Park.

Species

July

August

September

spp.

50.64

37.57

46.55

Salix brachycarpa

12.46

16.73

19.27

Kobresia

11.33

8.78

2.15

Salix planifolia

6.15

5.83

7.94

Deschampsia

4.11

3.88

5.95

.14

5.54

3.46

1.91

6.45

.21

.10

.38

6.76

3.85

2.11

0

Carex

Juncus

myosuroides

caespitosa

balticus

Geum rossii
Calamagrostis
Trifolium

canadensis

dasyphyllum

�-175-

Table 4. Percentage composition of elk diet across all animals in krummholzecotone vegetation type during summer 1977 in Rocky Mountain National Park.
(Continued).

July

Species

August

September

Poa artica

.85

3.57

.70

Oxyria

o

o

2.28

% Grass

70.37

61.34

66.93

% Forb

9.61

14.26

5.61

% Shrub

20.03

24.40

27.46

TOTAL BITES

6965

5595

6180

digyna

Table 5. Percentage composition of elk diet across all animals in alpine
tundra vegetation type during summer 1977 in Rocky Mountain National Park.

July

August

September

25.17

38.49

49.94

14.82

10.33

4.35

1. 79

13.77

12.67

Geum rossii

15.82

5.93

2.05

Caltha

leptosepala

4.07

8.09

5.08

Juncus

balticus

4.36

1.62

10.32

Salix planifolia

8.61

2.90

2.67

Poa artica

3.05

5.10

3.54

10.39

.62

.63

Species

Carex

spp.

Deschampsia
Trifolium

Trifolium

caespitosa
parryi

nanum

�-176-

Table 5. Percentage composition of elk diet across all animals in alpine
tundra vegetation type during summer 1977 in Rocky Mountain National Park
(Continued).

August

September

4.31

3.06

.03

Salix bra ch ycarpa

3.32

1.89

0

Salix spp.

0

1.07

3.79

Anemone

1.08

.58

2.27

.69

2.29

.68

% Grass

52.05

58.66

68.64

% Forb

35.89

35.47

24.52

% Shrub

12.06

5.87

6.83

TOTAL BITES

7941

6340

6774

July

Species

myos uroi des

Kobresia

canadensis

Polygonum

bistortoides

Elk diets were similar with respect to rankings of forage classes during winter
and summer (Fig 1). During both seasons graminoids were the most frequently
chosen food item. Mean percentage of graminoids in the winter diet was 59.9
(+8.3); summer diets averaged 65.5 (+5.1) percent graminoids.
Browse (leaves
a~d stems of shrubs) consumption was-greater in winter
= 32.7 + 7.9) than
summer
= 18.0 + 7.5) and increased significantly (P = 0.0003) through the
winter period from a mean of 17.3 (± 10.7) percent in November to 42.9 (± 10.0)
percent in March (Fig 2).

(x

(x

Diet Quality
Nutritional
manner:

quality of winter and summer diets were determined

in the following

1.

The proportion of each plant species in the diet based on frequency
of bites observed was calculated.

2.

The proportion by weight in the diet was calculated
for which bite weights were estimated.

for those species

�Diet

Composition

330/0
Shrub s

Forbs
17 %
I
,--...J

-...J
I

600/0

65%

Grass and
Grasslike

Gra ss and
Grasslike

Winter

Figure 1.

Mean percentages of forage classes
Rocky Mountain National Park.

Summer

in elk diets during winter

and summer,

1976-1977,

�•

I

60

I

•

•
••

•

50,

...-

~

~
0

---lW
0

•

•

1

40

•

•

I

•

z
-

•

30
w
(j)

~

•

•

•
•

3:
0
a:
en 20

•
I

"-

'-I

P =0.0003

•

•

IOl

•

R2 =0.43

•
•

•
Nov

Figure 2.

Dee

Jon

Feb

Percentage of browse in 1976-1977 winter diets of five elk.

Mar

CXl
I

�-179-

3.

Since weights were not estimated for all species, it was necessary
normalize these proportions to sum to 100 percent.

Therefore,
by:

diet in vitro digestible
DIGjp

= L

(IVDDM.

lp

i=l, kp

dry matter

to

(IVDDM was calculated

x CW .. )

lJP

where, DIGjp is digestibility of diet of animal j during grazing trial p,
IVDDMiP is digestibility of plant species i in diet of animal j during
grazing trial p, and kp is number of plant species contributing 2 percent
or more of total bites during grazing trial p. CWijp is the proportion by
weight of plant species i in the diet on animal j during grazing trial p.
Similarly,

crude protein
CPo

JP

content of diets were calculated

[E
i=l, kp

(N.

~

x CW .. )]
~p

by:

x 6.25

where CPjp is crude protein content of diet of animal j during grazing trial
p, Nip is nitrogen content of plant species i during grazing trial p, and
CWijp and kp are as defined above.
Confidence intervals on diet percentages were calculated as simple + intervals
on the means of untransformed percentages across five animals (Simpson et al.
1960:156).
Although rankings of forage class composition of elk diets was similar between
seasons, diet quality differed markedly between winter and summer.
Crude protein
content of summer diets averaged 12.6 (+ 1.4) percent while winter diets contained
on the average 5.4 (± 0.3) percent crud; protein (Fig 3). Differences in
digestibilities were less pronounced; mean summer IVDDM coefficients (51.5 ± 7.4)
were 17 percent higher than winter values (42.7 ± 2.8) (Fig 3). However, the
similarity of digestibilities of the two diets may be an artifact of inoculum
source.
Summer IVDDM coefficients were based on rumen inoculum collected from
domestic cows fed native grass hay while winter coefficients were determined
using inoculum obtained from wild elk grazing on winter range. We suspect coefficients for summer forages which were run on cow rumen fluid may be depressed.
This bias is currently being investigated.
Diet Quality and Taxonomic Composition--Taxonomic
mix of winter diets appeared
to respond to seasonal shifts in overwinter forage quality.
Crude protein
content of the seven most frequently consumed grasses declined significantly
(P = 0.0001) during winter; mean protein content of grasses in March (4.0 ~
0.6 percent) was 30 percent less than the average crude protein in grass in
November (5.7 ± 0.8) (Fig 4).
Declines were also observed in dietary protein (Fig 5). Crude protein in diets
fell from a mean of 5.9 (± 0.3) percent in November to 4.8 (± 0.5) percent in
March, an 18 percent decrement.
Slope of the regression line of crude protein

�-180-

12-

50 -

...~

-0

..-..

-- 40
~
0

c:

f0-

..--

&lt;l&gt;

6 -

0

..--

Q)

-

'0....

..c

(/)

30 r-

'"0

:J

01

'-

0

0

&lt;l&gt;

&lt;l&gt;
0

- 20 -

..--

0

&lt;l&gt;

Q)

01
0

0

'-

(l)

&gt;
&lt;!

l-

8 f-

(l)

&gt;.

10-

4 -

01

'-

10 ~

(l)

&gt;
&lt;!

Summer

Fig.

3.

Winter

2I

Summer

+
W· In.er

Mean dietary in vitro digestible dry matter (IVDDM) and
mean dietary crude protein of winter and summer elk diets,
1976-77 .

�-181-

•
•

7

•

•
6

~
0

--

•
•

5

en
en
0
~

(9

c

4

c

&lt;V

0

~

0...

3

•
•

&lt;V
"D

::J

~

u

•

•

•

•

•

R2 =0.37
2

P =0.0001

Nov

Fig.

4.

•

Dee

Jon

Feb

Crude protein content of grasses most frequently
consumed by elk during winter, 1976-77.

Mar

�-182-

-

7

~

0
......••.

•

-

&lt;l&gt;
0

•
•

6

c
C

&lt;l&gt;

0+-

0

"0....

5

&lt;l&gt;

•
R2 =0.77
P =0.00001

-0

::J

"0

•
•
•

4
Nov

Fig.

Dec

Jon

Feb

Mar

5. Crude protein content of elk diets during winter,
1976-77 .

�-183-

in grass against time was significantly greater (P = 0.07) than for the
corresponding regression of dietary protein with time (Fig 6). This
difference in rate of decline resulted in a 1.6 percentage point difference
between the two lines in late March (Fig 6). Such a difference in protein
would likely be of substantial significance to the animal.
Robbins et al.
(1974) reported metabolic fecal nitrogen excretion for white-tailed deer
fed a large variety of foods was 4.72 g of crude protein per 100 g dry matter
intake.
If this value is applied to elk, it is apparent that observed March
diets would likely provide protein close to equilibrium requirements, while
hypothetical diets composed of grass only would certainly fall below
nitrogen requirements (Fig 6).
The observed changes in diet mix mitigated the decline in forage protein.
As the percentage of protein in grass declined, there was a concomitant increase in percentage of browse leaves and stems taken in the diet (Fig 7).
Thus, it appears that elk compensated for decreased protein in grass by
shifting intake to include more browse with a higher protein content.
However,
if this tactic successfully increased protein intake in late winter, why did
the elk not consistently eat shrubs and maximize dietary protein during all
months?
There are two plausible explanations for the gradual change in diet
mix. First, the difference between protein content of browse and grass
increased during winter so that the relative benefit of consuming browse
leaves and stems was greater in March than in November.
Second, browse intake
was not without penalty; as the percentage of browse in the diet increased,
dietary IVDDM declined (Fig 8).
These observed diet shifts are consistent with optimum fcod selection tactics.
When grass and browse were equal in protein content, animals consumed primarily
grass and thereby maintained relatively high protein intake and diet digestibility.
When grass protein fell to critical levels, the elk switched to browse
at expense of forage digestibility.
CONCLUSIONS
Like other herbivores in temperate environments, elk are confronted with
seasonal changes in the quality of food resources (Fig 9). Diet quality is
at a highpoint in spring or early. summer and declines sharply as the quantity
of forage nutrients is diluted by elaboration of structural carbohydrate during
plant growth.
During winter, diet quality continues to decline although at a
reduced rate. Loss of nutrients over winter is attributable to leaching of
cell solubles from plant tissue.
Elk mitigate loss of forage quality through
alteration of diet mix.
Seasonal shifts in diet quality necessitate grazing strategies which replenish
endogenous nutrient reserves during periods when food resources are abundant
and conserve those reserves when food is less available.
Thus, winter and
summer diets interact to determine year-round animal condition.
Food Habits

Sampling

1977-78

A second winter of elk diet sampling was conducted for 10 days each month from
November through March, 1977-78.
Eleven plant species accounted for 81.43

�-184-

7

-

6

~

0
.•.....•.

c

- 5
Q)

0

\0...

Q)

-0

::J

\. E? %

\-

0

4
Ho: B D &lt; BG

P = 0.069
3
Nov

Fig.

6.

Dee

Jon

Crude protein in grass compared
winter, 1976-77.

Mar

Feb

to dietary

protein

during

�-185-

60

•
•

50

---~

0

•

• •
•
•

40

Q)

0

c

30

Q)

sn
3

0
\-

CO

20

R2 =0.43
10

•

P =0.0004

•
•

0

3
Average

Fig.

7.

Crude

6

5

4
Protein

in

Gra 55

(%)

Percentages of browse (leaves and stems)
in elk winter diets as a function of mean
crude protein in grasses.

�-186-

55

•

•
•

50

-~

~

°

"--"

~
0
0

•

•

•

•

•

45

.0-

&lt;1.&gt;

•

0

40

R2

=0.57
P =0.00002

•
•••

•

•

35

I

0

Fig.

10

8.

20

30

Browse

in Diet

40

50

60

(%)

Dietary in vitro digestible dry matter as a function
of percentage of browse (leaves and stems) in winter
diets of elk, 1976-77.

�-187-

80

16

~
\

Protein

\

\

70

14

\
\
0

•.......
~
0

•....•. 60

0

\

&gt;.

c

\

...a

-

12 ----

\

+-

If)

.--.
~

\

Q)

+-

50

10

Q)

01
.-

Q)

u
::J

0
+Q)

0
~
0...

~

U

40

8

,,

0

30

0

""-

0--

_0

6

&lt;, &lt;,
&lt;,

'0- - -0&lt;,

20

&lt;,

&lt;,

......•
4

Nov

Fig.

9.

Seasonal
means

Jon

Mar

course of changes in crude protein content and
dry matter of elk diets.
Points are
for diets of five animals.

in vitro digestible

Q)

�-188-

percent of the 132,152 bites observed in the composite diet totaled across
animals, months, and plant communities (Table 6). Mean diet percentage
and 90 percent confidence intervals have not been calculated for all
individual vegetation types at this time. Percentages shown in Tables 7-14
were calculated from total bites observed in each vegetation type sampled.

Table 6. Mean percentage composition with 90 percent confidence intervals
(C.l.) across 5 animals and all vegetation types during winter 1977-78 in
Rocky Mountain National Park.
Species listed are those which contribute
2 percent or more of total bites.

Species

Mean Percent

Muhlenbergia

montana

90 Percent C.l.

14.34

+2.42

Salix

spp. (stems)

10.46

+1.62

Salix

spp. (leaves)

9.53

+3.35

Carex

spp.

7.97

+2.54

Poa pratensis

7.51

+2.23

Potentilla

7.06

+3.72

6.59

+ .85

5.99

+4.98

5.08

+1.30

4.22

+2.42

2.68

+2.18

% Grass

49.81

+8.51

% Forb

5.82

+3.47

% Shrub

44.36

+8.82

fruticosa

Calamagrostis

canadensis

Purshia

tridentata

Populus

tremuloides

(leaves)
Phleum

pratense

Eriogonum

umbellatum

�-189-

Table 7. Percentage composition of elk diet across all animals in mesic
meadow vegetation type during winter 1977-78 in Rocky Mountain National
Park.

Species

November

December

January

February

March

Phleum

pratense

15.34

30.22

24.12

17.41

2.50

Carex

spp.

9.42

8.74

16.82

23.39

25.58

4.40

7.25

13.63

12.27

21.95

18.79

16.06

8.95

8.17

2.38

5.21

7.30

9.99

5.74

8.32

Potentilla

fruticosa

Poa pratensis
Calamagrostis

canadensis

Salix

spp. (leaves)

4.45

5.09

3.99

9.04

7.88

Bromus

inermis

22.l3

.57

.58

.56

1.75

Salix

spp. (stems)

3.35

2.71

6.94

6.15

3.81

Juncus

balticus

.62

5.66

7.57

4.28

3.69

o

11.95

3.99

1.88

0

.05

o

0

3.61

12.38

1.48

1.37

1.05

4.48

2.31

2.39

o

o

o

o

.10

o

.06

o

2.00

% Grass

80.31

82.03

73.13

66.46

49.53

% Forb

3.97

1.62

1.43

2.19

1.63

% Shrub

15.73

16.35

25.44

31.35

48.84

TOTAL BITES

2,092

4,384

3,632

2,877

1,599

Muhlenbergia
Rosa

richardsoni

spp.

Deschampsia
Alnus

caespitosa

tenuifolia

Muhlenbergia

(leaves)

montana

�-190-

Table 8. Percentage composition of elk diet across all animals in wet
shrub-meadow vegetation type during winter 1977-78 in Rocky Mountain
National Park.
Species

Potentilla

fruticosa

November

December

January

February

March

11.81

15.71

39.77

20.93

33.04

Carex

spp.

8.61

12.38

20.92

18.99

27.08

Salix

spp. (leaves)

25.05

13.89

.52

22.58

15.14

Salix

spp. (stems)

19.88

19.10

14.70

6.82

4.82

2.70

12.26

10.16

18.55

9.14

Poa pratensis

11.43

5.40

3.79

2.96

1.04

Phleum

pratense

7.08

6.29

1.81

2.32

2.08

Juncus

balticus

1.72

1.51

2.96

2.05

2.54

Calamagrostis

canadensis

Deschampsia

caespitosa

2.43

.92

0

.51

1.70

Muhlenbergia

richardsoni

0

4.40

0

0

0

.22

.60

0

1.39

2.10

.45

2.47

0

.80

0

.12

0

2.11

0

.75

% Grass

38.57

47.38

40.53

46.66

43.97

% Forb

2.54

1.64

2.18

.32

.09

% Shrub

58.89

50.98

57.28

53.02

55.94

TOTAL BITES

7330

5838

5539

4734

3468

Lonicera
Bromus
Rosa

involucrata

inermis

spp.

�-191Table 9. Percentage composition of elk diet across all animals in wet
meadow vegetation type during winter 1977-78 in Rocky Mountain National
Park.

Species

Carex

spp.

Potentilla

fruticosa

Poa pratensis
Salix

spp. (stems)

Calamagrostis

canadensis

November

December

January

February

7.54

l3.29

20.89

3.55

61.07

0

37.08

15.50

20.92

10.81

13.46

10.87

15.44

28.55

1.21

28.77

6.23

10.54

2.48

6.88

1.62

22.49

8.93

14.36

5.14

l3.52

3.90

0

15.96

4.38

March

Salix

spp. (leaves)

Bromus

inermis

.17

.46

15.00

7.27

0

Phleum

pratense

12.76

1.21

5.95

.53

1.28

14.85

0

0

0

0

.35

1.95

.74

2.84

.83

Muhlenbergia
Juncus

montana

balticus

Lonicera

involucrata

0

0

0

0

4.08

Ceanothus

fendleri

0

0

0

3.37

0

0

0

2.54

0

0

% Grass

52.09

50.37

69.50

57.09

69.61

% Forb

3.77

1.86

.25

.18

.60

% Shrub

44.14

47.77

30.25

42.73

29.78

TOTAL BITES

1724

1076

1613

564

l323

Muhlenbergia

richardsoni

�-192-

Table 10. Percen~agecompositionof elk diet across all animals in
ponderosapine/shrub vegetationtype during winter, 1977-78in Rocky
Mountain National Park.
November December January February March

Species

Muhlenbergia
Purshia

montana

tridentata

15.17

53.26

43.53

49.56

48.13

6.86

12.37

29.42

25.41

16.12

Eriogonum

umbellatum

14.82

4.38

8.47

5.57

4.40

Bouteloua

gracilis

8.66

16.06

3.23

5.29

2.35

13.42

.38

2.27

.61

5.13

7.13

.20

.38

.22

3.42

parryi

2.04

2.86

0

.47

5.26

Carex heliophila

7.42

.60

.57

.36

1.55

Poa pratensis

9.53

0

.45

.19

.11

.31

4.38

1.09

1.85

1.52

Stipa

comata

Bromus

inermis

Danthonia

Pinus ponderosa

(needles)

Ceanothus

fendleri

3.47

0

.09

.85

.67

Astragalus

spp.

2.53

.23

.03

.41

1.47

Agropyron

spicatum

3.13

.28

.13

.08

.91

o

0

0

0

3.31

o

0

3.04

0

0

2.53

.23

.03

0

0

% Grass

67.53

73.70

51.84

57.08

67.70

% Forbs

17.08

8.59

11.07

10.31

9.05

% Shrub

15.39

17.71

37.09

32.61

23.25

TOTAL BITES

4502

3992

9815

3628

3746

Rosa

spp.

PrunuS

virginiana

(leaves)
Populus

tremuloides

(leaves)

�-193-

Table 11. Percentage composition of elk diet across all animals in sagebrush
vegetation type during winter 1977-78 in Rocky Mountain National Park.
Species

Muhlenbergia
Purshia

montana

tridentata

November

December

January

February

March

52.55

57.65

24.21

61.63

49.39

0

5.07

38.83

23.36

25.64

Eriogonum

umbellatum

4.56

5.43

14.91

3.42

3.33

Ceanothus

fendleri

6.30

3.95

4.77

1.59

7.23

Boute.loua gracilis

11.39

5.57

1.57

.93

1.27

Stipa

8.45

10.15

.74

.14

.26

comata

Agropyron

spicatum

7.77

2.18

2.36

.41

.04

Potentilla

spp.

2.28

.85

3.49

2.72

2.54

% Grass

83.11

77 .87

30.61

66.46

53.37

% Forb

7.64

7.96

21.16

2.19

8.02

% Shrub

9.25

14.16

48.23

31.35

38.61

TOTAL BITES

746

1419

2032

3451

2282

Table 12. Percentage composition of elk diet across all animals in willow
vegetation type during winter 1977-78 in Rocky Mountain National Park.
Species

November

December

January

February

March

Salix

spp. (stems)

27.80

37.17

45.13

25.79

31.39

Salix

spp. (leaves)

28.71

24.33

12.25

31.76

38.62

2.64

8.91

13.75

11.07

20.38

Calamagrostis

canadensis

-----------------------------------------------------------------------------

�-194-

Table 12. Percentage composition of elk diet across all animals in willow
vegetation type during winter 1977-78 in Rocky Mountain National Park. (Cont'd).
November

December

January

February

March

5.33

9.17

19.35

8.81

4.72

Poa pratensis

20.27

12.79

3.85

7.64

2.42

Phleum

6.17

1.34

.43

3.06

.12

0

0

2.42

1.74

0

.01

.09

0

3.36

0

0

0

0

2.44

0

% Grass

36.83

33.46

39.49

32.98

28.10

Forb

2.10

.47

.31

.10

0

% Shrub

61.07

66.06

60.20

66.92

71.90

TOTAL BITES

6921

2322

4181

5746

2434

Species

spp.

Carex

pratense

Potentilla
Betula
Acer

01

/0

fruticosa

occidentalis

glabrum

Table 13. Percentage composition of elk diet across all animals in willow
vegetation type during winter 1977-78 in Rocky Mountain National Park.
Species

Muhlenbergia

montana

Prunus

virginiana

Stipa

comata

Prunus

virginiana

Eriogonum
Potentilla

(leaves)

(stems)

umbellatum

spp.

November

December

January

February

March

25.80

43.02

36.23

34.11

28.31

0

11.84

3.67

39.90

4.95

32.46

7.59

6.25

1.84

1.22

0

11.30

4.17

8.83

20.33

14.78

6.09

5.32

8.86

7.34

10.72

3.46

8.05

1.06

1.06

-----------------------------------------------------------------------------

�-195-

Table 13. Percentage composition of elk diet across all animals in willow
vegetation type during winter 1977-78 in Rocky Mountain National Park. (Cont'd).

November

December

January

February

March

.58

5.92

6.25

1.29

2.13

o

0

15.46

0

0

o

0

0

0

12.67

Poa pratensis

6.38

1.04

4.82

.16

0

Potentilla

2.32

0

4.82

.91

.53

.58

.83

0

.29

6.76

o

1.71

2.66

.32

1.81

o

0

0

0

4.90

o

1.54

0

0

2.55

% Grass

66.96

60.82

51.98

38.60

34.06

% Forb

28.70

12.17

19.70

10.67

16.18

% Shrub

4.35

27.01

28.32

50.73

49.76

TOTAL BITES

345

2399

1391

3093

1879

Species

Bouteloua

gracilis

Purshia

tridentata

Symphoricarpus

oreophilus

fruticosa

Lichen
Juncus

balticus

Acer glabrum
Rosa

spp.

Table 14. Percentage composition of elk diet across all animals in aspen
vegetation type during winter 1977-78 in Rocky Mountain National Park.

Species

Populus

tremuloides

November

December

January

February

March

(leaves) 32.23

34.27

35.39

33.36

53.20

28.97

17.47

15.75

5.87

16.91

.98

16.54

16.58

27.59

7.66

.63

.44

11.36

12.00

7.89

Poa pratensis
Calamagrostis
Salix

canadensis

spp. (leaves)

�-196-

Table 14. Percentage composition of elk diet across all animals in aspen
vegetation type during winter 1977-78 in Rocky Mountain National Park. (Cont'd).

Species

November

December

January

February

March

Salix

spp. (stems)

8.17

3.44

6.10

6.84

2.21

Phleum

pratense

3.05

12.04

3.98

3.25

.52

Carex

spp.

1.19

7.23

3.93

2.18

3.99

(stem)

1.12

4.01

2.29

2.25

2.79

(leaves)

7.38

0

0

0

0

6.10

0

0

0

0

.95

.84

0

3.83

0

.19

.04

3.60

1.23

0

Populus

tremuloides

Prunus

virginiana

Ceanothus

fendleri

Potentilla

fruticosa

Juncus

balticus

Danthonia

parryi

2.14

2.60

0

0

0

Bouteloua

gracilis

0

0

0

0

3.12

% Grass

38.59

56.73

43.85

40.33

32.20

% Forb

7.89

.04

.02

.21

15.81

% Shrub

53.52

43.23

56.13

59.46

52.00

TOTAL BITES

4294

2267

4577

3816

3081

Graminoid and browse (leaves and stem of shrubs) consumption was relatively
constant throughout the winter. Mean percentage of grasses and grasslikes
in the winter diet was 49.81 (± 8.51) percent. This forage class was dominated
by mountain muhly (Muhlenbergia montana), sedges (Carex spp.), bluegrass
(Poa pratensis) and canadian reedgrass (Calamagrostis canadensis).
Browse
species accounted for 44.36 (+ 8.82) percent of the winter diet. Willow stems
and leaves (Salix spp.), shrubby cinquefoil (Potentilla fruticosa), bitterbrush
(Purshia tridentata), and aspen leaves (Populus tremuloides) were outstanding
in this forage class. Forbs contributed 5.82 (± 3.47) percent to the winter
diet and were generally insignificant with the exception of sulphur flower
(Eriogonum

umbellatum).

�--197-

Differences in diet choices between months and between
been statistically tested.
Sampling Herbage

Biomass

communities

have not

1977-78

During September 1977, 960 quarter meter quadrats were ground level clipped
of all herbaceous vegetation.
Twenty-seven different species were separated,
sacked, dried, and weighted.
Estimates of willow and aspen leaves were made in early fall when the
majority of leaves had fallen.
In addition, current annual growth of aspen,
willow, shrubby cinquefoil, and bitterbrush twigs was collected from 200 two
meter plots.
Total herbage biomass was estimated within 15 percent of the
mean with 90 percent confidence.
Reasonably precise estimates of the more
common species within each vegetation type were obtained (Tables 15-22).

Table 15. Biomass estimates and percentage composition for standing herbage
in ponderosa pine-shrub vegetation type. Biomass estimates are mean grams
per meter squared with 90 percent confidence intervals.
Stem estimates are
for current growth only.

Species

Biomass

Forbs

7 .51 (±1.9 1)

22.68

5.73 (±0.97)

17.30

5.46 (±1.09)

16.49

2.17 (±0.45)

6.55

1. 83 (±O.60)

5.52

Carex heliophila

1.62 (±O.42)

4.89

Purshia

1.51 (±1. 14)

4.56

1.46 (±O. 50)

4.41

1.36 (±0.30)

4.11

Misc. shrub

1.10 (±0.91)

3.32

Agropyron

smithii

1. 0 1

(.:to. 36)

3.05

Danthonia

parryi

0.84 (.:to. 54)

2.54

Agropyron

spicatum

Muhlenbergia
Stipa

montana

comata

Bouteloua

tridentata

Eriogonum
Koleria

gracilis

umbellatum

cristata

(g/m2)

% Composition

�-198-

Table 15. Biomass estimates and percentage composition for standing herbage
in ponderosa pine-shrub vegetation type. Biomass estimates are mean grams
per meter squared with 90 percent confidence intervals.
Stem estimates are
for current growth only.
(Continued).

Biomass

Species

Chrysopsis

villosa

Misc. grass
spp.

Potentilla

2

(g/m )

% Composition

0.80 (±0.60)

2.42

0.43 (±0.32)

1.29

0.25 (±0.22)

0.76

33.11 (±4.38)

TOTAL

Table 16. Biomass and percentage composition for standing herbage in grassland
vegetation type. Biomass entries are mean grams per meter squared with 90
percent confidence intervals.
Shrub estimates are for current growth only.

(g/m

2

)

% Composition

Species

Biomass

Forbs

14.76 (±1.73)

21.87

8.74 (±l.76)

12.95

montana

Muhlenbergia
Danthonia

parryi

6.94 (±l.76)

10.28

Bouteloua

gracilis

6.84 (±1.38)

10.14

6.50 (±0.94)

9,63

Stipa

comata

Potentilla

spp.

5.17 (±1.38)

7.66

Agropyron

smithii

4.62 (±1.28)

6.85

Agropyron

spicatum

4.l3 (±1.05)

6.12

Koeleria

cristata

3.12 (±0.48)

4.62

2.46 (±0.49)

3.65

2.39 (±l.21)

3.54

Carex

heliophila

Eriogonum

umbellatum

�-199-

Table 16. Biomass and percentage composition for standing herbage in grassland
vegetation type. Biomass entries are mean grams per meter squared with 90
percent confidence intervals.
Shrub estimates are for current growth only.
(Continued) .

Biomass

Species

Chrysapsis

(g/m2)

% Composition

villosa

(stems and leaves)
Paa spp.

0.79 (±0.62)

1.17

0.48 (±0.20)

0.71

Carex

spp.

0.31 (±0.11)

0.46

Juncus

balticus

0.06 (±0.06)

0.09

0.02 (±0.03)

0.03

.02 (±0.03)

0.03

Misc. grass
Festuca

idahaensis

TOTAL

67.48 (±3.39)

Table 17. Biomass and percentage composition for standing herbage in wet
shrub-meadow vegetation type. Biomass entries are mean grams per meter
squared with 90 percent confidence intervals.
Stem estimates are for current
growth only. Leaf estimates are for leaves on the ground in late November.

Species

Carex

spp.

Calamagrastis

canadensis

Forbs

Biomass (gm/m2)

% Composition

54.02 (±12.18)

39.20

35.23 (±7.91)

25.57

18.98 (±5.12)

l3.77

Phleum

pratense

6.72 (±2.64)

4.87

Salix

spp. (stems)

4.67 (±2.10)

3.39

3.47 (±1.09)

2.52

3.22 (±1.46)

2.34

Paa spp.
Patentilla

(stems)

fruticasa

�-200-

Table 17. Biomass and percentage composition for standing herbage in wet
shrub-meadow vegetation type. Biomass entries are mean grams per meter
squared with 90 percent confidence intervals.
Stem estimates are for
current growth only. Leaf estimates are for leaves on the ground in late
November.
(Continued).

Biomass

Species

Salix

spp.

Glyceria
Juncus

(leaves)

grandis

balticus

(gm/m2)

% Composition

2.71 (±1.49)

1.20

2.24 (±1.26)

1.76

2.22 (±1. 42)

1.61

Sporabolis

crytandrus

1.39 (±1.0l)

1.01

Deschampsia

caespitosa

1.01 (±.689)

0.73

.99 (+.75)

0.72

Misc.

grass

137.79

TOTAL

(±17. 47)

Table 18. Biomass and percentage composition for standing herbage in wet
meadow type. Biomass entries are mean grams per meter squared with 90
percent confidence intervals.

Biomass

2
(g/m )

% Composition

46.41

(±12.24)

41.44

16.87

(±4.30)

15.06

13.40 (±4.80)

11.96

10.06 (±3.49)

8.98

8.40 (±5.12)

7.50

7.59 (±3.36)

6.78

3.09 (±1.88)

2.76

Misc. grass

2.96 (±1.47)

2.64

Poa spp.

1.48 (±0.56)

1.32

Species

Carex

spp.

Forbs
Glyceria
Juncus

grandis

balticus

Deschampsia
Calamagrostis
Phleum

caespitosa
canadensis

pratense

---------------------------.------------------------------------------------

�-201Table 18. Biomass and percentage composition for standing herbage in wet
meadow type.
Biomass entries are mean grams per meter squared with 90
percent confidence intervals.
(Continued).

Species

2
(g/m )

Biomass

Muhlenbergia

richardsoni

% Composition

1.07 (.±.76)

0.95

Sporabolis

crytandrus

0.49

(.±.72)

0.44

Potentilla

spp.

0.l3

(.±0.15)

0.12

lll. 99

TOTAL

Table 19. Biomass and percentage composition for standing herbage in the
mesic meadow vegetation type.
Biomass entries are mean grams per meter
squared with 90 percent confidence intervals.

Species

Carex

(g/m2)

% Composition

(.±12.70)

29.51

Biomass

spp.

33.82

22 .57 (.±8.l3)

19.69

14.28 (.±5.64)

12.46

11. 56 (.±3.36)

10.09

8.35 (.±4.05)

7.29

6.49 (.±2.40)

5.66

6.33

(.±2.99)

5.52

5.79

(.±2.51)

5.05

Poa spp.

3.25 (.±1.28)

2.83

Misc.

grass

0.79

(.±0.49)

0.69

Bromus

inermis

0.47

(.±0.32)

0.41

Glyceria

grandis

Muhlenbergia
Juncus

richardsoni

balticus

Koeleria

cristata

Forbs
Agropyron

spp.

Deschampsia

caespitosa

�-202-

Table 19. Biomass and peycentage composition for standing herbage in the
mesic meadow vegetation type. Biomass entries are mean grams per meter
squared with 90 percent confidence intervals. (Continued).

Species

Biomass

% Composition

0.40 (±0.49)

0.35

pratense

0.37 (±0.45)

0.32

spp.

0.12 (±0.18)

0.10

Calamagrostis
Phleum

2
(g/m )

Potentilla

canadensis

114.60 (±14.53)

TOTAL

Table 20. Biomass and percentage composition for standing herbage in
sagebrush vegetation type. Biomass entries are mean grams per meter squared
with 90 percent confidence intervals.
Shrub estimates are for current growth
only.

% Composition

Species

Artemisia

tridentata

(stems and leaves)

13.90 (±3.43)

28.76

8.02 (±1.69)

16.59

5.16 (±0.96)

10.67

4.98 (±2.15)

10.30

3.08 (±0.78)

6.37

3.07 (±0.93)

6.35

2.05 (±0.81)

4.24

Carex heliophila

1.79 (±0.46)

3.70

Eriogonum

umbellatum

1.39 (±0.73)

2.87

Agropyron

smithii

1.28 (±0.68)

2.65

Muhlenbergia

montana

Forbs
Chrysopsis

villosa

(stems and leaves)
Agropyron
Stipa

spicatum

comata

Bouteloua

gracilis

�--203-

Table 20. Biomass and percentage composition for standing herbage in
sagebrush vegetation type. Biomass entries are mean grams per meter squared
with 90 percent confidence intervals.
Shrub estimates are for current growth
only.
(Continued).

Species

Biomass

(g/m

2

)

% Composition

Purshia

tridentata

0.99 (±0.56)

2.04

Koeleria

cristata

0.97 (±0.31)

2.01

Danthonia

parryi

0.45 (±0.31)

0.93

Potentilla

spp.

0.36 (±0.26)

0.74

0.22 (±0.18)

0.45

0.16 (±0.10)

0.31

0.01 (±0.01)

0.02

Poa spp.
Festuca
Carex

idahoensis

spp.

TOTAL

48.33 (±4.08)

Table 21. Biomass and percentage composition for standiIlg herbage in willow
vegetation type. Biomass entries are mean grams per meter squared with 90
percent confidence intervals.
Stem estimates are for current growth only.
Leaf estimates are for leaves on the ground during late November.

Species

Carex

spp.

Calamagrostis
Salix

Biomass

canadensis

spp. (stems)

Forbs
Salix

spp. (leaves)

Poa spp.

(g/m

2

)

% Composition

92.00 (±15.99)

51.38

59.09 (±16.42)

33.00

10.67 (±4.27)

5.96

6.54 (±2.97)

3.65

6.23 (±3.68)

3.48

1.94 (±0.84)

1.08

�-204-

Table 21. Biomass and percentage composition for standing herbage in willow
vegetation type. Biomass entries are mean grams per meter squared with 90
percent confidence intervals.
Stem estimates are for current growth only.
Leaf estimates are for leaves on the ground during late November.
(Continued).

Biomass

Species

Phleum

spp.

Glyceria

grandis

Misc. grass

2
(g/m )

% Composition

1.08 (±1.0l)

0.60

0.52 (±0.79)

0.29

0.42 (±0.37)

0.23

179 .07 (±19 .71)

TOTAL

Table 22. Biomass and percentage composition for standing herbage in the
aspen vegetation type. Biomass entries are mean grams per meter squared
with 90 percent confidence intervals.
Stem estimates are for current growth
only. Leaf estimates are for leaves on the ground in late November.

Biomass

Species

Populus

(g/m2)

% Composition

tremuloides

27.28 (±7.39)

27.74

Forbs

26.51 (±4.43)

26.70

Poa spp.

10.64 (±2.03)

10.71

Phleum

8.66 (±2.03)

8.72

8.65 (±5.78)

8.71

inermis

4.53 (.±3.02)

4.56

spp. (stems and
leaves)

3.06 (.±1.43)

3.08

2.10 (.±0.94)

2.11

0.98 (.±0.71)

0.98

(leaves)

pratense

Calamagrostis
Bromus

canadensis

Rosa

Populus

tremuloides

(stems)
Agrostis

album

----------------------------------------------------------------------------

�-205-

Table 22. Biomass and percentage composition for standing herbage in the
aspen vegetation type. Biomass entries are mean grams per meter squared
with 90 percent confidence intervals.
Stem estimates are for current growth
only. Leaf estimates are for leaves on the ground in late November.
(Continued).

Biomass

Species

(g/m

2

)

% Composition

0.97 (±0.71)

0.98

0.42 (±0.35)

0.27

0.27 (±0.33)

0.42

0.26 (±0.18)

0.26

Misc. grass

0.34 (±0.30)

0.34

Misc. shrub

0.22 (±0.29)

0.22

TOTAL

99.30 (±8.97)

Juncus

balticus

Glyceria

grandis

Deschampsia
Agropyron

Nutritional

caespitosa

spp.

Analysis

and Diet Quality

Chemical analysis on the major species in the 1977-78 elk diet are currently
in progress.

LITERATURE

CITED

Hobbs, N. T., and D. L. Baker.
1977. Systems Modeling Big Game Populations:
Simulations of the Carrying Capacity of the Rocky Mountain National Park
Elk Winter Range.
Colo. Div. Wildl., Game Res. Div., Fed. Aid Proj.
W-38-R-32.
Game Research Report., July, Part 2. p. 125-200.
Robbins, C. T., R. L. Prior, A. N. Moen, and W. J. Visek.
1974. Nitrogen
metabolism of white-tailed deer. J. Anim. Sci. 38(1):186-191.
Simpson, G. L., A. Roe, and R. C. Lewontin.
1960.
Harcourt, Brace, and Work, New York.
440pp.

Prepared by _---"'-LL-=--c·~·:.~"",,--,-_--,gliiii!:...!f:~=~.1i~'
Dan L. Baker
Wildlife Researcher

_

Quantitative

zoology.

�-206-

STUDY PLA..'&gt;iI

STUDY PLAN NO. 2

QUANTITATIVE AND NUTRITIONAL
DESCRIPTION OF THE SUMMER DIET
SELECTION OF ELK IN ROCKY MOUNTAIN NATIONAL PARK

June 17. 1977

Dan L. Baker

�-207-

Study Number

A.

2.

Quantitative and Nutritional Description of the
Summer Diet Selection of Elk in Rocky Mountain
National Park.

NEED
Nutritional evaluation of wild ungulate diets in winter (Wallmo et
ale 1977, Urness et ale 1975, Dietz et ale 1959, Short et ale 1966,
Robbins 1973, Amman et ale 1973, Mautz et ale 1973, Ullrey et ale
1972, Ward 1970) together with the physiological adaptations and requirements of these herbivores (Silver et ale 1969, Silver et ale
1971, Holter et ale 1975, Ullrey et ale 1969, Thompson et ale 1973,
Milchunas 1977, McEwan 1970, McEwan and Whitehead 1970) suggest that
winter habitat in general is inadequate to meet total energy requirements and survival is dependent in part on energy stored as fat during
summer and fall. Thus summer forage and its ability to meet nutritional
requirements for lactation, reproduction, growth and replenishment of
endogenous energy reserves may be significant to survival on nutritionally
restricted winter habitat.
Food habitat information is a first step in understanding animal-environment relationships.
Food is consumed, digested and used metabolically
to meet nutritional needs which are constantly changing (Moen 1973).
Selection of forage species is a complex phenomena which can be influenced
by availability, season, nutritional requirements and geographical location. Little quantitative information is available regarding the summer
food habits of Rocky Mountain elk (Cervus canadensis) in specific vegetation communities.
At the same time little attention has been given to
the nutritional evaluation of these forages and their potential role in
meeting the physiological requirements of elk throughout the year.
Food habits of Rocky Mountain elk in the western U.S. have been extensively
reviewed by Kufeld (1973). Of pertinence to this study is that literature
which deals with forage preferences and forage quality relevant to the
area of Rocky Mountain National Park.
In Rocky Mountain National Park
(RMNP), Stevens (1969) observed elk grazing and reported elk diets in
summer (July-Sept.) consisted of 82 percent forbs (Trifolium dasyphy11um,
Trifolium nanum). 32 percent shrubs (Salix sp., Vaccinium sp.) and 11
percent grass (Carex sp., Phleum sp.). Tileston (1962) collected 16
stomach samples from elk grazing in RMNP in summer.
Grasses comprised
68 percent of the diet while shrubs and forbs contributed 17 and 15 percent, respectively.
Unfortunately, the majority of studies describing diet selection of elk
have been done by observing foraging animals from a distance, stomach
analysis or by fecal microhistology.
The inherent bias and shortcomings
of these techniques have been reported (Bergerud and Russel 1964,
Buechner 1950, Wallmo et al. 1973, Westoby et al. 1976, Adams et al. 1962,
Keiss and Schoonveld 1976).

�-208-

One of the main objectives of this study is to quantify diet selection
of elk for individual plant species in specific vegetation types, so
diet observation of tame elk will be used to quantify ingested foods in
RMNP in summer.
This technique is described by Wallmo and Neff (1970).
It has been found effective for describing food habits of other wild
ungulates (McMahan 1964, Watts 1964, Healy 1967, Neff 1967b, Wallmo 1964,
Regelin et al. 1974, Schwartz and Nagy 1976, Hobbs and Baker 1977). M~jor
criticism of this method is directed toward the artificiality of using
pen-reared animals to estimate the diet preference of their wild counterparts.
Empirical evidence suggest that both tame and wild species select
similar forage plants (Wallmo and Neff 1970, Schwartz and Nagy 1974,
McMahan 1964, Healy 1967).
Nutritional evaluation of summer elk forage has not been documented.
Johnson et al. (1968) reported the chemical composition and in vitro digestibility of 19 alpine plants estimated to be important to bighorn sheep.
Results of this study showed that crude protein and phosphorus content of
all species decreased with advancing maturity while calcium and cellulose
content increased.
In vitro digestibility coefficients decreased with
advancing maturity.
Regelin et ale (1974) evaluated mule deer forage
throughout summer (2867-3202 m) from logged and unlogged areas in Colorado.
Crude protein and moisture content declined significantly between dates
but in vitro digestibility did not change as the plants matured.
B.

OBJECTIVES
The specific

C.

objectives

of this study are:

1.

To describe and quantify the summer (June-Sept.)
elk in Rocky Mountain National Park.

2.

To assess the chemical constitution
selected by elk in summer in RMNP.

EXPECTED

RESULTS

diet selection

and digestibility

of

of foods

OR BENEFITS

Successful completion of the objectives of this study should provide basic
ecological knowledge relevant to the quality of summer elk forage. More
importantly, this information together with qlmntitative estimates of the
quantity and quality of forage resources available to elk in winter will
contribute to a more thorough understanding of the nutritive ecology of
elk in Rocky Mountain National Park.

D.

APPROACH
Food Habits
Five tame elk (13 mo. old) previously reared and trained for food habits
experiments (Hobbs and Baker 1977) will be used to estimate summer forage
preferences in RMNP.
Feeding observations will be conducted in three

�-209-

general plant communities comprlslng the sub-alpine-tundra
region.
These three communities (willow-park, krumrnholz-ecotone, alpine tundra)
accounted for 97 percent of the 1,176 elk observed on summer range by
Stevens (1969).
1.

Willow-park (elevation 3050-3355 m). This community is characterized
by mesic swales, wet marshes and xeric upland sites.
Salix sp.,
Bet.!:!}.a.
~ndu.losa,
_~~Eex sp ,, Calamo..8..!:.ostis
canadensis, Deschampsia
caespitosa, and Caltha leptosepala are the dominant species.
Twentyeight percent of the summer elk observations were recorded in this
type (Stevens 1969).

2.

Krumrnholz-ecotone (elevation 3355-3660 m). This unit is the ecotone
between the subalpine forest below and the alpine tundra above.
Species of Salix occur in small patches and dominate the area. Small
islands of Abies lasiocarna and Picea engelmanni are interspersed
throughout.
Deschampsia caespitosa is characteristic of the understory vegetation (Braun 1969).
Stevens (1969) recorded 41 percent use
by elk in summer for this unit.

3.

Alpine tundra (3660 + m). This ecological region varies from sharp
peaks and ridges to gently rolling expanses.
Some soils are thin and
rocky while others are stable with well developed rich soil. Kobresia
meadow is the climax vegetation on the level winter snow-free sites.
Other common species are: Deschampsia caespitosa, Silene acaulis,
Trifoliu~ sp., Carex sp. (Marr 1962).
Twenty-eight percent of the elk
observed were in this community (Stevens 1969).

The vegetation in this study area is complex and difficult to characterize.
The boundaries between types are seldom discrete, but gradually intergrade
into each other creating a mosaic of many small communities.
The delineation of elk summer range into three major vegetation types was based on the
best ecological data available as well as the logistical and statistical
constraints inherer.t in sampling according to this study design.
Experimental animals will be transported from the Division of Wildlife
Research Center in Fort Collins to holding pens maintained at Rocky
Mountain National Park. Animals will not be fed during the sampling period
except for small amounts of grain used as rewards for loading into the
transporting vehicle and leading to the study sites.
The first day of each
sampling period, the animals and observers will adjust to new surroundings
and exposure to available forage. After the adjustment period a morning
grazing trial will be conducted each day for six consecutive days. Due to
the intensive recreational use of the study area and likely interference
with grazing animals each trial will begin at sunrise and end in early
morning.
Since a maximum of three animals can be sampled in this time
period, two days will be required to sample each vegetation type. Vegetation types will be randomly chosen with replacement for each day of the
trial until each type has been sampled by each of the five animals.

�-210-

Two observers will lead two or three tame elk to a randomly chosen
point within each vegetation type and allow the animals to graze.
Animals will not be restricted to one vegetation type. When three
animals are sampled in one trial each animal will be observed for
one hour by each observer.
Observers will alternate animals at
hourly intervals until each animal has been observed for two hours.
When two animals are sampled each observer will observe each elk for
one hour. This design will allow ten hours of grazing per vegetation
type with five replicate animals per type. Within each grazing trial
elk will be randomly assigned to observer and grazing rotation.
Each observer will count the number of bites by species consumed by
elk. Forage choices, vegetation type, elk number and weather conditions
will be recorded on tape recorders and later transcribed to data forms.

Forage Evaluation
Following each morning trial, observers will hand pluck forage samples
of species grazed by elk. Parts of several plants will be collected to
approximate the same proportion of leaves, stems and flowers selected.
Locations of feeding sites and/or plants grazed will be marked by flagging
during each trial. Approximately 100 g (wet wt.) of each species comprising one percent or more of the diet will be collected from marked
locations, stored in moisture-proof bags, labeled by date, vegetation
type, animal, plant species and frozen for later analysis.
In addition 25 simulated bites will be collected in paper bags from
marked locations for each species comprising one percent or more of the
diet in order to obtain an average weight per bite. Estimates of diet
composition based on these weights will be compared to diet composition
derived from bite counts.
Nutritional analysis of summer elk forage will include: Dry matter, ash,
in vitro digestibility, gross energy, cell wall constituents, lignin,
acid detergent fiber, and nitrogen.

Statistical

Analysis

The principle questions asked in this study are what plants in summer
constitute elk food, and what is the relative quality of this forage
throughout summer and early fall.
Statistical treatment of diet data (percentages) has been reviewed by
Hobbs and Baker (1977). Similar analysis using t intervals will be used
to make inferences on the precision of diet estimates.
Multivariate analysis of variance
hypothesis testing for:

(Harris 1975, Peden 1972) will permit

(1) differences in forage choices across animals between
periods and between vegetation types.

sampling

�-211-

(2) differences in forage quality between
vegetation types.

sampling

periods

and

Stepwise regression analysis will be used in an attempt to relate elk
forage preference to several measured attributes of vegetation (Hobbs
and Baker 1977).
A statistical weakness in this experimental design is lack of independence of observations.
Due to logistical considerations, two or three
animals will be grazed simultaneously, thus introducing possible bias
in forage selection due to the influence of one animal upon another.
Hopefully, this bias will be partially overcome by the homogeneity and
relatively large size of each defined vegetation type.

Scheduling
Define study area and vegetation
Pretrial

June 1, 1977

types

June 3, 1977

grazing with elk

First sampling

period:

1.

Take animals

2.

Pretrial

3.

Begin grazing

facility

in RMNP

grazing and plant collections

End grazing
Second sampling

to holding

June 23, 1977
June 24 through June
25, 1977
June 26, 1977

trial

July I, 1977

trial

period:

1.

Take animals to RMNP

August

2.

Pretrial

August 6, 1977

3.

Begin grazing trial

August

7, 1977

End grazing trial

August

12, 1977

Third sampling

grazing and plant collections

5, 1977

period:

1.

Take animals to RMNP

Sept. 6, 1977

2.

Pretrial

Sept. 7, 1977

3.

Begin grazing trial

Sept. 8, 1977

End grazing trial

Sept. 13, 1977

and plant collection

Analysis of data and writing
Completion date

manuscript

Sept .-Dec. 1977
Feb. 1978.

�-212-

Responsibility
D. L. Baker (Colo. Div. Wildl.)
N. T. Hobbs (Nat. Res. Ecol. Lab-CSU)
Lyn Stevens (Colo. Div. Wildl.)
)
Don Reichert (U.S. Forest Service)
)

Principal Investigator
Principal Investigator
Laboratory Analysis

Estimated Cost
Field work
Laboratory analysis
Equipment and supplies
Computer time
Mileage and per diem

man-days
42
40

cost ($)
1,000.00
200.00
100.00
300.00
1,600.00

E.

LOCATION
The study area is bounded by Fall River Road on the north and Trail Ridge
Road on the south. Fall River Pass delineates the western edge while
Willow Park patrol cabin is the eastern boundary. This area lies partly
in T6N R7SW, and partly in TSN, R7SW and is approximately 10So4S'W long.
o
and 40 26'N. lat. The general area is Rocky Mountain National Park,
Larimer County, Colorado (see map) •

F.

RELATED FEDERAL PROJECTS
This study is part of the Rocky Mountain National Park Cooperative Elk
Project involving Rocky Mountain National Park, U.S. Forest Service
(Rocky Mt. Forest and Range Experiment Station), Natural Resource Ecology
Lab (NREL), Colorado State University, and Colorado Division of Wildlife.

��-214-

LITERATURE

CITED

Adams, L., W. G. O'Rega, and D. J. Dunaway.
1962. Analysis of forage
consumption by fecal examination.
J. Wildl. Manage. 26(1) :108-111.
Amman, A. P., R. L. Cowan, C. L. Mothershead, B. R. Baumgardt.
1973. Dry
matter and energy intake in relation to digestibility in white-tailed
deer. J. Wildl. Manage. 37(2):195-201.
Bergerud, A. T., and L. Russel.
for Newfoundland caribou.

1964. Evaluation of rumen food analysis
J. Wildl. Manage. 28:809-814.

Braun, C. E. 1969. Population dynamics, habitat,
tailed ptarmigan in Colorado.
Ph.D. Thesis.
Fort Collins.
189 p.

and movements of whiteColorado State University.

Buechner, H. K. 1950. Life history, ecology, and range use of the pronghorn antelope in Trans-Pecos, Texas.
Am. MidI. Nat. 43(2) :257-354.
Dietz, D. R., R. H. Udall, H. R. Shepherd, and L. E. Yeager.
1959. Seasonal
progression in chemical content of five key browse species in Colorado.
Proc. Soc. Am. Foresters Meeting 58:117-122.
Harris, R. J.
New York.

1975. A primer of multivariate
332 p.

statistics.

Academic

Press,

Healy, W. M. 1967. Forage preference of captive deer while free ranging in
the Allegheny National Forest.
M.S. Thesis.
Pennsylvania State Univ.,
Unive:rsity Park.
93 p ,
Hobbs, N. T., and D. L. Baker.
1977. Simulations of the carrying capacity
of the Rocky Mountain National Park elk winter range.
Colo. Div. Wildl.
Fed. Aid W-38-R-32, WP 17, Job 2. Prog. Rep., Game Res. Rep. July (In Press).
Holter, J. B., W. E. Urban, Jr., H. H. Hayes, H. Silver, and H. R. Skutt.
1975.
Ambient temperature effects on physiological traits of white-tailed deer.
Can. J. Zool. 53:679-685.
Johnson, A., L. M. Bezeau, and S. Somoliak.
1968. Chemical composition and
in vitro digestibility of alpine tundra plants.
J. Wildl. Manage. 32:(4):
773-777.
Keiss, R. E., and G. G. Schoonveld.
1976. Evaluation of the nutritional
quirements of bighorn sheep.
Colo. Div. Wildl., Fed. Aid W-4l-R-26,
1, Job 16. Job Prog. Rep., Game Res. Rept. January.
p.1-56.
Kufeld, R. C. 1973.
26(2) :106-113.

Foods eaten by the Rocky Mountain

elk.

reWP

J. Range Manage.

Mautz, W. W., H. Silver, J. B. Holter, H. H. Hayes, and W. E. Urban, Jr. 1973.
Digestibility and related nutritional data for seven northern deer browse
species.
J. Wildl. Manage. 40:630-638.
Marr, J. W. 1962. Utilization of the front range tundra, Colorado.
In:
Grazing in terrestrial and marine environments.
D. J. Crep (Ed.) Br.
Ecol. Soc. Symp. 4:109-118.

�- 21')-

McEwan, E. H. 1970. Energy metabolism of barren ground caribou
tarandus).
Can. J. Zool. 48:391-392.

(Rangifer

McEwan, E. H., and P. E. Whitehead.
1970. Seasonal changes in the energy
and nitrogen intake in reindeer and caribou.
Can. J. Zool. 48:905-913.
McMahan, C. A.
livestock.

1964. Comparative
J. Wildl. Manage.

food habits of deer and three classes of
28:798-808.

Milchunas, D. G. 1977. In vivo-in vitro relationships of Colorado mule
deer forages. M. S. Thesis. Colorado State University, Ft. Collins.
l33 p ,
Moen, A. N.
458 p.

1973.

Wildlife Ecology.

W. H. Freeman Co., San Francisco.

Neff, D. J. 1967b. Field feeding studies with tame deer.
Ann. Meeting New Mex., Ariz. Sect. Proc. 6:68-77.
Peden, D. G.
plains.

Wildl.

Soc.

1972. The tropic relations of Bison Bison to the shortgrass
Ph. D. Thesis, Colorado State University, Ft. Collins.
134 p.

Regelin, \-1. L., O. C. Wallmo, J. Nagy, and D. R. Dietz.
1974. Effect of
logging on forage values for deer in Colorado.
J. Forestry 72:282-285.
Robbins, C. T. 1973. The biological basis for the determination of
carrying capacity.
Ph. D. Thesis, Cornell Univ., Ithaca.
239 p.
Schwartz, C. C., and J. G. Nagy .. 1974. Use of trained pronghorn to
measure forage consumption.
Proc. Sixth Antelope State Workshop
6: 58-73.
Schwartz, C. C., and J. G. Nagy. 1976. Pronghorn dietary quality relative
to forage availability and other ruminants in Colorado.
J. Wildl.
t1anage. (In Press).
Short, H. L., D. R. Dietz, and E. E. Remmenga.
1966.
mule deer browse plants. Ecology 47(2):222-229.

Selected nutrients

in

Silver, H., N. F. Colovos, J. B. Holter, and H. H. Hayes.
1969. Fasting
metabolism of white-tailed deer. J. Wildl. Manage. 33(3):490-498.
Silver, H., J. B. Holter, N. F. Colovos, and H. H. Hayes.
1971. Effect of
falling temperatures on heat production of fasting white-tailed deer.
J. Wildl. Manage. 35(1):37-46.
Stevens, D. R. 1969. Ungulate ecology studies in Rocky Mountain Park.
National Park Service Progress Rep. on file, Park Headquarters, Rocky
Mountain National Park.
(Mimeo). 74 p.
Thompson, C. B., J. B. Holter, H. H. Hayes, H. Silver, and W. E. Urban.
1973.
Nutrition of white-tailed deer. I. Energy requirements of fawns. J.
Wildl. Manage. 37(3):301-311.

�-216-

Tileston, J. V. 1962. Big game research projects.
Technical
Fed. Aid Project W-38-R.
Colo. Div. Wildl.
79 p.

Bull. No.9.

Ullrey, D. E., W. G. Youatt, H. E. Johnson, L. D. Fay, B. L. Schoepke, and
W. T. Magee.
1969. Digestible energy requirements for winter maintenance
of Michigan white-tailed does. J. Wildl. Manage. 33(3):482-490.
Ullrey, D. E., W. G. Youatt, H. E. Johnson, A. B. Cowan, R. L. Covert, and
W. T. Magee.
1972. Digestibility and estimated metabolizability of
aspen browse for white-tailed deer. J. Wildl. Manage. 36(3):885-891.
1975. Nutrient content of mule
Urness, P. J., D. J. Neff, and J. R. Vahle.
deer diets from ponderosa pine range. J. Wildl. Manage. 39:670-673.
Wallmo, O. C. 1964. Arizona's
Wildl. Views 11(6):4-9.

educated deer.

Ariz. Game and Fish Dep.

Wallmo, O. C., and D. J. Neff. 1970. Direct observation of tamed deer to
measure their consumption of natural forage. Rocky Mountain For. Range
Exp. Stat. Misc. Pub. 1147. 105-110 p.
Wallmo, O. C., R. B. Gill, L. H. Carpenter, and D. W. Reichert.
1973.
Accuracy of field estimates of deer food habits.
J. Wildl. Manage.
37(4):556-562.
Wallmo, O. C., L. H. Carpenter, W. L. Regelin, R. B. Gill, and D. L. Baker.
1977. Evaluation of deer habitat on a nutritional basis.
J. Range
Manage. 30(2):122-127.
Ward, A. L. 1970. Stomach content and fecal analysis.
Methods of forage
identification.
Rocky Mountain For. and Range Exp. Sta. Misc. Pub.
1147. p. 146-158.
Watts, C. R. 1964. Forage preference of captive deer while free ranging in
a mixed oak forest. M. S. Thesis.
Pennsylvania State Univ., University
Park. 65 p.
Westoby, M., G. R. Rost, and J. A. Weis.
diets by microscopically identifying
J. Mammal. 57(1):167-172.

1976. Problems with estimating
plant fragments from stomachs.

�July,

1978

-217-

JOB PROGRESS

St at;e of
Project
Work

COLORADO
No.

W-38-R-33

Job Title

Deer-Elk

18

Plan No.

Period

REPORT

Job No.

Deer and Elk Management

Covered:

April

1,

Investigations
1

---------------------------------

Study

1977 - June 30, 1978

Personnel:
Thomas M. Pojar.
I gratefully acknowledge the technical
assistance of John Bartholow and Peter Strong of the Data Support Group,
Western Energy and Land Use Team, Office of Biological Services, USF&amp;WS.

ABSTRACT
The normal updating of all models with current harvest statistics was
accomplished with copies of appropriate simulations provided to regional
biologists.
The major effort for field workshops was concentrated in the.
southeast and northwest regions.
The northwest region acquired a relnote
computer terminal during this segment and all field workshops for the spring
of 1978 were conducted by regional personnel.
Several new simulations were
created for pronghorn populations in the southwest region and one new bighorn
sheep simulation was created in the southeast region.
Generalized results
of the population modeling program are contained in the publications listed
in the result~ section.

��-219-

DEER AND ELK MANAGEMENT

STUDY

Thomas M. Pojar

P. N. OBJECTIVE
Devise and test a statewide

deer and elk management

system.

SEGMENT OBJECTIVES
1.

Divide the state into a system of data analysis units (DAU' S) that
are deemed practical for gathering and analyzing population data.

2.

Based on currently available information, estimate population
by DAU, through the simulation modeling approach.

3.

Conduct training workshops in"volving management personnel to familiarize
them with the interpretation of the output from the model and to
incorporate empirical data from fieldmen directly associated with the
DAU into the simulation to produce a candidate management simulation.

4.

Identify data that are most useful for improving
parameters.

estimates

parameters,

of population

RESULTS
The following manuscripts

were prepared:

Pojar, T. M. 1977. Use of a population model in big game management.
Proceedings of the Western Association of State Game and Fish Commissioners,
57th Annual Conference, Tucson, Arizona, July 10-13.
(In Press).
Pojar, T. M. 1978. A management perspective of population modeling.
In:
C. W. Fowler and T. Smith (eds.). Dynamics in Large Mammal Populations.
John Wiley and Sons, Inc., New York.
(Manuscript submitted).

\

prepare:b:b P2 t'~)
Thomas M. fojar
Wildlife Researcher

��July, 1978
-221-

JOB PROGRESS

State of

REPORT

COLORADO
--------~~~~=------------

Proj ect No.

W-38-R-33

Deer-Elk

Work Plan No.

20

Job No.

Job Title

Brucellosis-Leptospirosis

Period Covered:

Investigations
I

---------------------------------

Survey of Big Game Animals

in Colorado

April 1, 1977 - March 31, 1978

Personnel: Marilyn Stevens, Joan Ritchie, Carol Ann Weinland,
Ford, Gene Grenerd, Robert McLean, and William Adrian

Richard

ABSTRACT
During the 1977 antelope season we checked 666 antelope for brucellosis
and 589 for five serotypes of Leptospira (~. canicola, grippotyphosa,
hardjo, icterohemorrhagiae, and pomona). All samples screened for brucellosis
were negative, however, we did find a significant increase (13% of those
tested) for serological titers for ~. hardj o. All other serotypes of
Leptospir~ were considered negative.

��-223-

BRUCELLOSIS-LEPTOSPIROSIS
SURVEY
OF BIG GAME ANIMALS IN COLORADO

William

J. Adrian

P. N. OBJECTIVE
To survey big game populations within the State of Colorado
presence or absence of brucellosis and leptospirosis.

for the

SEGMENT OBJECTIVE
1.

Survey Colorado's antelope populations
during the annual hunting season.

METHODS

for brucellosis

and leptospirosis

AND MATERIALS

Blood sample kits were mailed to all antelope hunters with the request
draw a blood sample from their kill and return the kit to the Colorado
Cooperative Brucellosis Laboratory in Denver, Colorado for processing.

they

The kit consists of a cardboard mailer, blood tube and stopper, plastic
bag, rubber band, paper towel and instructions for obtaining the sample
and mailing the kit.

RESULTS AND DISCUSSION
The Wildlife Research Laboratory in cooperation with the Cooperative Brucellosis Laboratory in Denver, has in 1977 completed its ninth annual survey
for brucellosis and leptospirosis in Colorado's big game populations.
Brucellosis is a specific contagious disease which primarily affects cattle,
swine, and goats.
It can, however, affect many other animals including
elk, deer, antelope and man (undulant fever, Malta fever and Bang's Disease).
It is caused by bacteria of the Brucella group and is characterized by
abortion in the female, and to a lesser extent, inflammation of the testes
and infection of the accessory sex glands in the male and infertility in
both sexes.
Leptospirosis is a febrile (feverish) disease caused by certain species of
leptospirae.
The disease occurs with various manifestations
in dogs, cattle,
pigs, sheep, goats, horses, and wildlife and is transmissable to man.
During the 1977 season we checked antelope for brucellosis and five serotypes
of leptospirosis (leptospira canicola, grippotyphosa, hardjo, icterohemorrhagiae,
and pomona).

�-224-

All licensed antelope hunters (5669) were sent blood kits and we received
2221 (39%) blood samples of which 1555 (70%) were badly hemolyzed and were
not usable.
Again in 1977, all samples tested for brucellosis were negative,
however, in sharp contrast to previous years we found 13% of the samples
screened for Leptospira had serological titers for ~. hardjo at the 1-10
level or higher.
All other serotypes for Leptospira were considered
negative.
In 1974, the last previous serological survey for these disease
in antelope, we found 4 - 1/10 and 2 - 1/40 serological titers for ~. hardjo.
We considered those titers negative due to our large sample size and the
low titers.
The 1977 data shows 55 - 1/10, 15 - 1/40, 2 - 1/160 and
1 - 1/640 titers.
During the 1978 hunting season both antelope and deer will be surveyed.
The antelope survey is being repeated to follow the trend of ~. hardjo in
antelope.
Those areas with the highest ~. hardjo incidences will be
special coded so we can contact the hunter later to find exactly where he
took his animal.
Once an infected herd can be isolated we will attempt
to isolate ~. hardjo and confirm or discredit the serological data.
The discrepancy between blood samples received and those tested for either
brucellosis or leptospirosis is due to the condition of the samples when
received at the laboratory; this is a highly variable factor from year to
year depending on weather and the lag time from kill to laboratory.
Generally, however, approximately one-third of all samples received are
very good to excellent quality and approximately one-fourth are completely
hemolyzed and unusable.

')

Prepared

by

/6~&lt;~;.t.:&gt;::;;:"
William Adrian
Wildlife Researcher

�July 1978
-225-

JOB PROGP~SS

State of

COLORADO
------~~~~~--------

Project

No.

Work

Deer-Elk
22

Plan No.

Job Title
Period

Job No.

Experimental

Covered:

Personnel:

REPORT

April

Deer Inventory

1, 1977-March

Investigations
1

-----------------------------

- Northeast

Region

31, 1978

Allen Anderson, Brian Anderson, David Bowden, Randy Fischer,
Howard Geduldig, Jennie Goldberg, William Lance, Sally McKenney,
Judy Melzer, Vicki Jameson, Don Samuelsen, Gene Schoonveld,
Lynn Sexton, and Stan Vallejos.

ABSTRACT
Two plot stake types, two plot stake colors and 4 plot spacings were tested
for differences in mean location times on 6 representative habitats of G.M.D.
20 (Big Thompson) deer winter range.
Nonparametric analytical procedures
were inconclusive regarding plot stake type or color.
The mean location
times of the 66 ft (20.1 m) spacing, however, were several times faster than
the 132 ft (40.2 m), 264 ft (80.5 m) or 528 ft (160.9 m) spacings.
On a
randomly selected square mile within each of 19 stratum, deer pellet groups
were counted on 320, 10 m2 circular sample plots located at 66 ft (20.1 m)
intervals along two pairs of randomly-selected
transects one mile (1.6 km)
in length.
Eight subsampling plans were developed from analyses of these
data and a subsampling plan involving one pair of randomly selected transects
one-half mile (0.8 km) apart was selected.
In this plan, each transect has
20 plots per group consisting of 10 consecutive plots spaced 66 ft (20.1 m)
separated by 30 chains (1,980 ft) from another set of 10 plots also spaced
66 ft (20.1 m).
It is pointed out that sample size requirements may be
adquately estimated only from counts of deer pellet groups during spring on
permanent sample plots from which all deer pellet groups have been removed
during the fall.
Buck:doe and fawn:doe ratios were estimated from 1,067 deer
classified on 20 routes, November 28, 1977 to December 29, 1977.
Ratios did
not differ significantly
(P = 0.10) between observers or methods of recording
but the 1977 buck:doe (0.170) and fawn:doe (0.665) ratios were significantly
(P = 0.10) greater than in 1976.
Buck:doe ratios decreased significantly
(P = 0.10) during the latter part of the 1977 breeding period.
For the fourth
consecutive year sample size requirements for the buck:doe ratio were impractically large.
Fawn:doe ratios were adequately sampled to be within 10
percent deviation from the true value at the 90 percent confidence level.

��-227EXPERIMENTAL

DEER INVENTORY

- NORTHEAST

REGION

Allen

E. Anderson
and
David C. Bowden

P. N. OBJECTIVE

To design appropriate sampling
reliably estimate deer numbers
on a preliminary sampling of a
four administrative
regions of

and analytical procedures necessary to
and buck:doe:fawn
ratios on winter range based
problem management unit within each of the
the state.

SEGMENT

OBJECTIVES

1.

To design and execute a preliminary sampling of deer pellet groups, or
where domestic sheep may occur, helicopter counts of deer on the entire
winter range of one Game Management Unit within each of the four
administrative
regions.

2.

To design a sampling scheme based on the means, variances, and deer
density strate estimated from Objective 1 capable of estimating mean deer
numbers +15% at a = 0.05.

3.

To sample buck:doe:fawn
ratios on 12 or more randonlly selected, all-day
walking routes on the winter range of one Game Management Unit within
each of the four administrative
regions.

This report is concerned only with the preliminary sampling of deer and elk
pellet groups and buck:doe and fawn:doe ratios on the winter range of Game
Management Unit (G.M.U.) 20 within the Big Thompson and North St. Vrain
river drainages.
METHODS

AND MATERIALS

Deer and Elk Census
On the often heavily forested and precipitous terrain characteristic
of G.M.U. 20 winter range, the distance between permanent plots is an important factor in the time spent relocating the stakes and marking the center of
each permanent plot; hence must be considered in the design of efficient
schemes for sampling numbers of deer and elk pellet groups.
Stake shape and
color may also affect relocation times.
Because of the frequently high human
use of some winter range areas, plot stakes ideally should not be obvious
to the casual observer but readily visible to the biologist.
An experiment was performed to determine the time involved in the location
deer fecal group plot stakes and to test for differences in mean location
times among the four combinations of two stake types and two stake colors.
Stake height above ground varied from about 4 to 9 inches (10.2-22.9 em).

of

�-228-

Six locations in habitats representative of the study area were selected.
Within each location 3 rep l Lc at-Lons were run. Each replication consisted of
4 plots with 66 ft (20.1 m) spacing, 4 plots with 132 ft (40.2 m) spacing, 4
plots with 264 ft (80.5 m) spacing and 4 plots with 528 ft (160.9 m) spacing.
In order to average out the effects of habitat changes on the unequal spacing
design, the 66 ft (20.1 m) spacing test was run by having flagged wood survey
stakes about 3 ft (0.9 m) in height located 528 ft (160.9 m) apart and then
a plot 66 ft (20.1 m) from the flag. A similar arrangement was performed
for the l32ft
(40.2 m) and 264 ft (80.5 m) spacings (Appendix I). Plot
relocation simulated actual field procedures on G.M.U. 20 (Anderson 1977).
That is a two-person crew walked at a normal pace and both searching for the
plot stake.
Observers and the order of the 4 stake type-color combinations were randomly
assigned.
Time to locate a stake was recorded with stop watches to the
nearest second except those stakes which could not be located in 60 minutes.
A truncated score of 3,600 seconds was recorded for these stakes.
The experimental plots were established by two persons who did not participate in the tests. Six persons, who had no previous experience in finding
plot stakes were involved in the tests, May 18, 1977 to May 30, 1977. The
stake types were reinforcing-bar and 0.375 inch (0.95 cm) angle-iron painted
glossy blue or glossy green over silver rust-preventative.
The training, field procedures and criteria for assessing deer and elk pellet
groups have been described (Anderson 1977). Notable changes included an
increase in habitat types (Appendix II) and the use of IBM PORT-A-PUNCH boards
to record data directly onto data processing cards (Patton and Casner 1970).
In 1976, data were collected on 71 square miles (183.9 km2) and an optimum
allocation of the number of square miles to be sampled per stratum (± 10
percent at P&lt;0.05) was calculated (Anderson 1977).
Since the total sample
size (59 square miles or 152.8 square kilometers) was impractically large,
efforts during 1977 emphasized the determination of the most efficient way to
subsample a randomly selected square mile (2.59 km2) and thus reduce the
sample size. Note that these estimates were based on counts of deer pellet
groups subjectively judged as "new" or voided during the previous autumn,
winter, spring, and early to late summer, hence are gross approximations.
In
addition, the catastrophic heavy rainfall of 31 July 1976 may have washed
sufficient numbers of "new" pellet groups from those plots on steep slopes
to significantly affect results since field work continued until October 15,
1976.
In 1977, counts of deer and elk pellet groups on temporary circular, 10-m2
(107.64 ft2) sample plots were made employing the following sampling scheme.
On a randomly-selected
square mile (2.59 km2) within each of 19 stratum, a
set of 10 transects each 0.1 mile (0.16 km) apart, were plotted on U. S.
Geological Survey Topographic Quadrangles.
Each transect consisted of 80
plots, spaced 66 ft (20.1 m) apart, and hence, was a mile (1.6 km) in length
and traversed the square mile (2.59 km2) in that cardinal magnetic bearing
which would place most of the transect approximately perpendicular to the
contour lines.
The transects were numbered from one to 10 in sequence.
The
transects one and five, two and six, etc., which were one-half mile (0.8 km)
apart were considered pairs.
From the five pairs of transects, two pairs

�-229-

were randomly selected and new (subjectively estimated as &lt;1 year old) and
old (subjectively estimated as &gt;1 year old) deer and elk pellet groups were
counted.
Thus, counts were made on 320 plots per square mile (2.59 km2) on
each of 19 square miles (49.2 km2) during 1977.
A variety of subsampling schemes were tested which could be applied to the
existing sampling plan.
Consider the estimation of variance in that plan
characterized as stratified and multistage since the primary units (square
miles) are stratified and then selected by simple random sampling within the
strata.
Each square mile (2.59 km2) is then systematically
sub-sampled by
random placement of the first transect within the first 0.1 mile (0.16 km)
and every other transect is systematically
located from that point.
An overestimate of the variance of tbe mean number of fecal groups per plot can be
obtained by simply obtaining the mean number of fecal groups per plot for each
sample square mile (2.59 km2) and finding the variance within stratum and
proceeding like ordinary stratified sampling; however, the finite population
correction factor is not used.
In the present study the primary interest was
in subsampling design and the sample of size 1 in each stratum does not allow
unbiased variance estimation.
In the 1976 sample, however, the average within
stratum variance based on square mile totals' was 40 percent of the variance
among all square miles ignoring strata.
The 40 percent figure is used as an
adjustment factor in the analyses presented herein.
Mule Deer Herd Structure
The statistical and sampling rationale and field procedures in estimating buck:
doe and fawn:doe ratios within C.M.U. 20 have been described (Anderson 1977).
During 1977, 6 all-day walking routes were added to the 14 routes established
in 1976.
Some modifications
in the 14 routes were made during 1977 to
increase sample size.
The 20 routes were run from November 28, 1977 to December 29, 1977.
Two methods were used in deer classification
and entered
separately on prepared forms.
Method 1 included both unclassified
single deer
and groups in which one or more individuals could not be classified as an
antlered buck, a doe, or a fawn of the current year and unspecified sex.
Method 2 included only classified single deer and those groups in which all
deer could be so classified.
RESULTS
Deer and Elk Census
The mean search times by stake type, color and spacings are listed in Table 1.
Although analysis of variance tests were performed, the tests for significant
differences among stake type combinations were made on a basis of nonparametric test for equality of median locaticn times.
Within each locationreplication and spacing set (one each for 66 ft (20.1 cm) ...
, 528 ft
(160.9 m»
the 4 location times of the stake type combinations were compared
to their median.
The number of times a particular stake-combination
was
greater than and the number less than or equal to the median of the 4 comparable locations times are listed in Table 2. A chi square test was performed
for each of the 5 sub tables to test for significant differences in median
location times.
These chi square values are reported below each subpart.
Averaged over all spacings no significant (P&lt;.05) difference in the median
location times is indicated.
However, both 132 ft (40.2 m) spacing (P&lt;.05)

�Table 1.

Mean search times (seconds) by plot stake type, color and spacings.

Angle Iron
Green

Mean

Overall
Mean per Number
of 66 ft (20.1 rn)
Segments

18.278

16.000

31.792

31.792

60.833

39.056

226.889

135.333

67.667

323.111

430.611

222.222

201.222

294.292

73.573

227.944

722.500

575.000

491.611

504.389

63.049

Plot
Spacing
ft

m

66

20.1

59.722

33.167

132

40.2

214.556

264

80.5

528

160.9

Reinforcing Bar
Blue

Stake Type and Color
Reinforcing Bar
Angle Iron
Green
Blue

Stake

Type Only

Reinforcing Bar

-

259.056

Angle Iron

-

223.847

Stake Color~
Blue

- 210.049

Green

- 272.854

I
N
l.U
0
I

�-231-

Table 2. Comparisons among 4 plot stake spacings showing the number of times
a particular stake type-color combination was greater than, less than or
equal to the median of the 4 comparable location times in seconds.

Plot
Spacing
ft

m

66

20.1

132

264

528

40.2

80.5

160.9

Overall

Plot Stake
Type and Color

Equal To
Or Less
Than Hedian
Time

Greater
Than
Hedian
Time

Reinforcing

bar, blue

8

10

Reinforcing

bar, green

7

11

Angle-iron,

blue

14

4

Angle-iron,

green

12

6

Reinforcing

bar, blue

4

14

Reinforcing

bar, green

10

8

Angle-iron,

blue

10

8

Angle-iron,

green

12

6

Reinforcing

bar, blue

7

11

Reinforcing

bar, green

9

9

Angle-iron,

blue

11

7

Angle-iron,

green

10

8

Reinforcing

bar, blue

13

5

Reinforcing

bar, green

8

10

Angle-iron,

blue

6

12

Angle-iron,

green

9

9

Reinforcing

bar, blue

32

40

Reinforcing

bar, green

34

38

Angle-iron,

blue

41

31

Angle-iron,

green

43

29

1/ Each comparison has 3 degrees of freedom; Tabular X2

Calculated
Chi Square
Value 1/

7.420

7.815 at P

8.000

1.946

5.778

4.730

0.05.

�-232-

~

and 66 ft (20.1 m) spacing (P~.07) indicate the possibility of smaller median
location time for the reinforcing bar stake.
The overall test for differences
in the two give:
&lt; Median
&gt; Median
Reinforcing
Angle-iron

bar

66
84

78
60

x2 (1) = 4.508 (P~.03).
It should be noted that an interaction is indicated between spacings and
median location time.
The 264 ft (80.5 m) spacing indicating no difference
among the stake combinations in median location times and the 528 ft (160.9 m)
spacings indicating preference for the angular iron stake, if any.
The nonparametric
procedures were used due to the existence of the extreme
values from the stakes which were not found or were difficult to locate.
The
data were very non-normal with hetergeneous variances.
Four stakes were not
found on the 528 ft (160.9 m) spacing (2 of reinforcing bar and 2 of angleiron) and one stake at the 132 ft (40.2 m) spacing was not found.
Thus, results of this experiment were inconclusive regarding plot stake color
and type as important influents on median location time.
However, the greater
ability of the sharpened point of the angle-iron stake to penetrate rock
fractures or tree roots recommend its use.
It is clear, however, that the
66 ft (20.1 m) stake spacing greatly reduced relocation time. Hence, more
time can be spent on plot search.
Analyses of counts of deer pellet groups on 6,080 sample plots showed that a
number of options are available for subsampling.
1. Only one of the two
pairs of transects need to be used.
2. Only one of the two transects within
a pair need be used.
3. Each transect of 80 plots can be considered in a
variety of subgroups.
The cases considered for each trap-sect are: (a) 8
groups of 10 plots spaced 528 ft (160.9 m), (b) 4 groups of 20 plots spaced
264 ft (80.5 m), (c) 2 groups of 40 plots spaced 132 ft (40.2 m), (d) 8
groups of 10 consecutive plots spaced 66 ft (20.1 m), (e) 4 groups of 20
consecutive plots spaced 66 ft (20.1 m), (f) 2 groups of 40 consecutive plots
spaced 66 ft (20.1 m), (g) 4 groups of 20 plots (10 consecutive plots spaced
66 ft (20.1 m) separated by 1,980 ft (603.5 m) from another set of 10 consecutive plots also spaced 66 ft (20.1 m»,
and (h) 2 groups of 40 plots (20
consecutive 66 ft (20.1 m»
spaced plots which are 1,320 ft (402.3 m) from
20 consecutive 66 ft (20.1 m) spaced plots).
A nested analyses of variance
was calculated for each data configuration using the group total as a basic
data point.
The components used in the fu~OVA are given in Table 3. The
variance which would have been obtained on a single square mile basis can then
be predicted by

+ (1 -~)

s;

+ (1 - ~)

mk
In the analysis of variance n0 adjustment was made for stratification.
An
adjusted variance was also calculated based on the 40 percent reduction for
the average within strata variance estimated, as noted above, from the 1976
data.
This factor applies only to the sf term.
The number of square miles
needed to obtain the same precision of one square mile sampled with the 320
plots are given for each subsampling plan in Table 4. For example, under

�-233-

plan g with m = 1, k = 2, h = 1 has used m = 1 of the two pairs of transects,
k = 2 of the 2 transects within a pair and h = 1 of the 4 groups of plots.
The plot group is 20 plots with 10 consecutive plots 1,980 ft (603.5 m)
spacing and 10 consecutive plots at 66 ft (20.1 m) spacings.
It would take
1.67 square miles (4.3 km2) with this sampling pattern to give the same
precision as one square mile with the full 320 plots.
Note the subsampling
pattern consists of only 40 plots.
However, adjusting for stratifification,
which should be done, it will take 2.67 square miles (6.91 km2) of this
pattern of 40 plots to have equal efficiency of the 320 plots per square mile
(2.59 km2) sample.

Table 3. Components used in the nested analysis of variance among
subsampling plans for counts of deer pellet groups on sample plots
on mile-long transects within a square mile.

No. of
Units

Source

Transects

n

miles mean

M

pairs within miles
mean sq. /KH

m

a pair

K

transect within
mean sq. /H

pairs

k

a transect

H

groups within
mean sq.

transect

h

a square mile

within

Groups within

Sample
Sizes

Component

N

Square mile
Pairs within

Variance

8
located

square/MKH

Because of the approximate nature of the values in Table 4, no further estimates of relative efficiencies sample size requirements of each subsampling
plan will be presented.
Valid but still tentative estimates of sample size
requirements can be made from counts of deer pellet groups on permanent sample
plots installed during the summer of 1978 and read during autumn 1978 and
again in spring 1979.
Several years of consecutive readings will be necessary
before a firm estimate of sample size requirements can be made.
In view of
the material presented in Table 4, which of the 8 subsampling plans should be
used on the winter range of G.M.U. 20? The 66 ft (20.1 m) spacing between
plot stakes is clearly the fastest and often the only practically feasible
spacing in the often dense timber and precipitous terrain characteristic
of
this winter range.
Since the 66 ft (20.1 m) plot spacing is employed in subsampling plans d, e, f and g, these are the plans of choice even though the
528 ft (160.9 m) spacing of plan a is most efficient.
Even though plan g
appears relatively less efficient than plans f or h, the smaller number of
sample plots per group recommend it. An adequate assessment of these candidate
subsampling plans, however, should ideally include time studies of travel
between plots within groups, between groups of plots, and search time on
plots which have been permanently marked and cleared at least once.

�-234-

Table 4. The number of square miles to be sampled for each of 8 subsamp1ing
plans if efficiency equal to that of 320 plots is desired. 1/

Plan

a

b

c

d

e

f

g

Unadjusted

m = 1
k = 2
h = 1

1.59

m
k
h

1
2
1

m
k

2/

m
k
h

1
2
1

2.47

1.40

m
k
h

1
2
1

1. 99

20

1. 32

m
k
h

1
2
1

1.80

40

h

1
2
1

m
k
h

1
2
1

2.32

m
k
h

1
2
1

4.30

m
k
h

1
2
1

1.89

m
k
h

1
2
1

3.22

m
k
h

1
2
1

1.52

m
k
h

1
2
1

2.29

m
k

1
2
1

1.67

m
k
h

1
2
1

2.67

20

1
2
1

1. 36

m
k

1
2
1

1. 90

40

h

h

m = 2
k = 2
h = 1

Number of Square Miles
Adjusted

Number
Plots
per
Group

m
k
h

m
k
h

m
k
h

2
2
1

1
1
1

1.31

2.04

2.06

h

m
k
h

m
k
h

2
2
1

2
2
1

1.77

3.60

10

10

20

m
k
h

1
1
1

3.66

40

1/

- Note number of plots sampled per square mile is number of plots per group
times mkh.

1/ Adjusted

variance based on the 40 percent reduction
strata variance estimated for the 1976 datA.

for the average within

�-235-

Elk pellet groups have received only cursory attention herein.
But the
estimated sample size (+ 10 percent at P = 0.05) of 83 square miles (215 km2)
as compared to 58 (150.2 km2) for deer in 1976 (Anderson 1977) suggests that
much larger sample sizes may ultimately be necessary.
Improved stratification,
specifically for elk, should reduce the variance since ITlanysquare miles of
lower elevation deer winter range in G.M.D. 20 support few, if any, elk.
Mule Deer Herd Structure
Total counts by Methods 1 and 2 for observer 1 and observer 2 on 20 routes
are listed in Table 5. Mean group size differed significantly
(P&lt; 0.05)
among strata (Table 6). Buck:doe and fanw:doe ratios are compared between
methods for each observer and between observers for each method and between
observers for both methods combined (Table 7). No significant (P = 0.10)
differences were detected in these comparisions.
But a comparison of the
1976 and 1977 buck:doe and fawn:doe ratios showed a significant (P = 0.10)
increase in both ratios during 1977. When the same comparisons were limited
to the original 14 routes and observer 1 (A.E.A.), however, buck:doe ratios
did not increase but fawn:doe ratios increased significantly
(P = 0.10) in
1977. A comparison of buck:doe and fawh:doe ratios obtcined during peak
(prior to December 12) and late (after December 12) breeding periods (Anderson
and Medin [1967]) shows no change in fawn:doe ratios but a significant
(P&lt; 0.10) decrease in the buck:doe ratios.
A regression of the 1976 fa~l:doe ratios (Y) on the 1976 mean pellet groups
per plot (X) within 13 stratum yielded a correlation coefficient of -0.159.
But the number of deer recorded (Y) during the 1976 herd structure sampling
regressed on the 1976 mean pellet groups per plot (X) within 14 stratum
yielded a significant (P = 0.02) correlation coefficient of 0.592.
The foregoing analyses suggest speculations that mean group size may be
changing over the month long sampling period and that habitat differences
between stratum probably influence both visibility and sociality of deer.
The 1977 increase in the fawn:doe ratios appears genuine but the increase
in buck:doe ratio may be more a function of observer 2 encountering a few
large aggregations on one of the six new routes.
The general lack of significant differences in buck:doe and fawn:doe ratios between observers or
between methods has held constant for 4 years.
Significantly
fewer bucks
seen during the late breeding period is expected and likely associated with
their lesser activity.
The lack of significant correlation between fawn:doe
ratios and mean pellet groups per plot seems to negate the hypothesis that
fawn:doe ratios were higher at lower winter range densities (Bowden and
Anderson 1976 ).
In Table 8 it is once again clear that the buck:doe ratio is an intractable
sampling problem on winter range.
It is probable that the present 20 routes
will generally sample the fawn:doe ratio within 15 percent of the
true ratio at P = 0.05 and often within 10 percent of the true ratio at

P = 0.10.

�-236Table 5. Number of deer classified by two observers using two methods on
20, all-day walking routes on Big Thompson (G.M.U. 20) winter range, November
28, 1977 to December 29, 1977 .

Method 11./
Fawn
UncI

Observer

Strata

Buck

Doe

1

2

0

0

0

3

0

0

4

0

5

Method il/
Doe
Fawn Total

Total

Buck

0

0

2

2

1

5

0

0

0

1

4

2

7

0

0

0

0

0

0

0

0

0

0

0

0

0

0

10

10

20

5

0

0

0

0

0

0

1

1

2

6

1

1

0

1

3

1

3

1

5

7

1

2

2

1

6

0

7

4

11

7

0

0

0

1

1

0

5

5

10

8

0

3

0

3

6

0

4

4

8

8

0

1

0

2

3

1

4

3

8

9

0

3

1

2

6

0

15

11

26

9

0

2

0

19

21

5

45

32

82

10

1

4

4

10

19

3

40

26

69

10

0

0

0

0

0

2

12

3

17

11

0

0

0

0

0

4

5

4

l3

11

1

3

2

3

9

5

21

18

44

13

1

1

0

2

4

4

22

19

45

l3

1

4

2

2

9

3

60

38

101

14

0

0

0

5

5

6

31

18

55

15

0

0

0

0

0

3

11

7

21

6

24

11

51

92

40

302

207

549

Total
F:D

.4583

F:D

.6854

SE

.l337

SE

.0335

B:D

.2500

B:D

.l325

SE

.0725

SE

.0274

----------------------------------------------------------_._-------------------

�-237-

Table 5. Number of deer classified by two observers using two methods on 20,
all-day walking routes on Big Thompson (G.M.U. 20) winter range, November 28,
1977 to December 29, 1977 (cont.).

Method 11/
Fawn
UncI

Observer

Strata

Buck

Doe

2

2

0

0

0

3

0

1

4

0

5

Total

Method il/
Doe
Fawn

Total

Buck

0

0

1

0

0

1

0

1

2

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

6

5

3

14

5

0

0

0

0

0

0

0

0

0

6

0

0

0

0

0

1

2

2

5

7

0

0

0

3

3

1

0

0

1

7

0

0

0

0

0

1

0

0

1

8

0

0

0

0

0

0

0

0

0

8

1

6

5

5

17

3

10

8

21

9

0

0

0

0

0

1

32

20

53

9

0

0

0

0

0

3

20

20

43

10

0

4

1

5

10

4

27

24

55

10

0

0

0

0

0

1

5

2

8

11

2

2

0

3

7

6

20

8

34

11

0

0

0

2

2

10

23

21

54

13

0

1

3

2

6

4

41

35

80

13

0

7

3

9

19

3

20

12

35

14

1

2

1

7

11

2

10

7

19

15

0

2

1

5

8

1

5

4

10

4

25

14

42

85

48

220

166

434

F:D

.5600

F:D

.7545

SE

.1458

SE

.0569

B:D

.1600

B:D

.2182

SE

.0927

SE

.0518

]) See text for explanation.

Total

�-238Table 6. Group size differences of mule deer sighted among strata of the
Big Thompson (G.M.U. 20) winter range, November 28, 1977 to December 29, 1977.
Statistics

95% Con. Interva1-

of Group Size
SE

n

Stratum

1/

Lower

Upper

2

6

12

4

1. 50

.5

3

9

33

3

3.00

1.00

4

o

o

o

5

36

164

11

3.27

.65

6

13

47

4

3.25

.63

7

33

127

12

2.75

.52

8

63

399

21

3.00

.71

1.52

4.48

9

231

1,925

43

5.37

.62

4.12

6.62

10

178

1,446

29

6.14

.66

4.79

7.49

11

163

921

41

3.98

.41

3.15

4.81

l3

299

2,031

69

4.33

.40

3.53

5.l3

14

90

448

23

3.91

.44

3.00

4.82

15

39

181

12

3.25

.64

1,160

7,734

272

To test whether the mean group size is the same in each stratum,
following ANOVA:
d.

Source

f.

SS

MS

Among

strata

11

284.88

25.90

Within

strata

260

2502.06

9.62

Total

(corrected)

271

2786.94

-1/ Computed
deer.

for only those stratum with 20 or more sightings

1/ Significant

at P

0.05.

we have the

F

2.692:./

of one or more

�Table 7. Comparisons of buck:doe and fawn:doe ratios sampled on Big Thompson (G.M.U. 20) winter range November 28,
1977 to December 29, 1977 between the 1976 and 1977 samples, and between the first and second halves of the 1977
sampling period.
90% c..r ,
Conf. Int. Values
(~ Width).!.! +

Ratio and
No. Deer

Observers,
Methods

B:D
46 326

Observer 1

Method 1
.2500

vs. Method 2
.l325

.1175

.1218

.2393

-.0043

N.

S.

B:D
52 245

Observer 2

Method 1
.1600

vs. Method 2
.2182

-.0582

.2571

.1989

-.3153

N.

S.

F:D
218 326

Observer 1

Method 1
.4583

vs , Method 2
.6854

-.2271

.2502

.0231

-.4773

N.

S.

Ratio Comparison

Ratio
Diff.

Conc1usion-~/

I
N

F:D
180 245

Observer 2

Method 1
.5600

B:D
88 522

Method 2

F:D
373 522

vs. Method 2
.7545

LV
\D

-.1945

.2319

.0374

-.4264

N.

S.

Observer 1 vs. Observer 2
.1325
.2182

-.0857

.0866

.0009

- .1723

N.

S.

Method 2

Observer 1 vs. Observer 2
.6854
.7545

-.069l

.0960

.0269

-.1651

N.

S.

B:D
98 571

Methods
1 and 2

Observer 1 vs. Observer 2
.1411
.2122

-.0711

.0762

.0051

-.1473

N.

S.

F:D
398 571

Methods
1 and 2

Observer 1 vs. Observer 2
.6687
.7347

-.0660

.0932

.0272

-.1592

N.

S.

B:D
150 995

Methods and
Observers
Combined

1976
.1226

vs.

1977
.1703

-.0477

.0459

-.0017

-.0936

Sign.

F:D
556 995

Methods and
Observers
Combined

1976
.3726

vs.

1977
.6649

-.2903

.1096

-.1807

-.4000

Sign.

I

�Table 7. Comparisons of buck:doe and fawn:doe ratios sampled on Big Thompson (G.M.U. 20) winter range November 28,
1977 to December 29, 1977 between the 1976 and 1977 samples, and between the first and second halves of the 1977
sampling period.

Ratio and
No. Deer

Observers,
Methods

B:D
70 521

Observer 1 (14 origina1 routes)

1976
.1321

vs.

F:D
268 521

Observer 1 (14 origina1 routes)

1976
.3893

vs,

B:D21

Routes between 11-27
and 12-11 Observers
and Methods combined

1976
.1818

vs.

"

1976
.3636

vs.

B:rJ!-1
98 571

1977 Observer and
Methods Combined

F:rJ!-1
398 571

1977 Observer and
Methods Combined

15

66

Ratio Comparison

Ratio
Diff.

90% C. 1.
Conf. Int. Values
(~ Width).!.! +
-

Conc1usionll

1977
.1369

-.0048

.0383

.0335

-.0431

N. S.

1977
.6598

-.2705

.1057

-.1648

-.3762

Sign.

1977
.3182

~ .1364

.4070

.2706

-.5434

N. S.
I

F:D21
34

66

N

+:-

1977
.8182

-.4546

.5484

.0938

-1.003

N. S.

Pre 12-11
.3358

vs, Post 12-12
.1198

.2160

.0920

.3080

.1240

Sign.

Pre 12-11
.7226

vs.

Post 12-12
.6889

.0337

.3470

.3806

-.3133

N. s.

II 90% confidence interval about the difference between the sample ratios.

11 Not significant (N. S.) or significant (Sign.) at P = 0.10.
31 Comparisons based on only 4 routes.
41 Based on V = min (n1 - 1, n2 - 1) = 6 degrees of freedom.

0
I

�-241-

Table 8. Number of routes required for sampling buck:doe and doe:fawn
ratios based on 20, all-day walking routes on G.M.U. 20 winter range,
November 28, 1977 to December 29, 1977.

No. Deer
Components

Buck

Doe

Ratio

SE

Buck:Doe

98

571

.1716

.0379

1/
Percent-

Conf. Level

(I-a)

.95

.90

.80

05

1,710

1,167

688

10

428

292

173

15

190

130

77

20

107

73

44

Buck

Doe

Ratio

SE

398

571

No. Deer

Fawn: Doe

1/
Percent-

1/

Deviation

.6970
Conf. Level

.0379

(1-0.)

.95

.90

.80

05

104

71

42

10

26

18

11

15

12

8

5

20

7

5

3

from true ratio.

�APPENDIX I
PLOT LAYOUT
FOR TEST OF PLOT DISTANCE,

Two randomly-selected

STAKE TYPE AND COLOR

observers for each

compartment
~

Wood survey stakes

•

Steel stakes at 66' (20.1 m), 132' (40.2 m)
and 264' (80.5 m) from wood survey stakes
For 528' (160.9 m) plot interval the only
wood survey stake is at the starting
point

Designated Section Corner (SW)
I

t&lt;1

.j:--

~~/264'

ft

N

(80.5 m) (or some other arbitrary distance)

I

't7
I~'

Plot Interval (ft)

I
66

1

H
U

~

(20.1 m)

528 (160.9 m)
2~----~----~----~----~~--~----~----~--------r-~----~'-----+----~

en

~

H

3

4

ti}&amp;

(:).

A

Go

(;)#,1

G~

Q.

B

COMPARTHENT

0,.

G~I

e·

G"
C

Go&gt;

S'

264

(gO.5

m)

132

(40.2

m)

�-243APPENDIX

II

HABITATS ON DEER WINTER RANGE OF G.M.U. 20
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.

14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.

Pipo-O
Pipo-D
Pipo-Psme
Psme
Psme-Pipo
Pico-O
Pico-D
Pico-Psme
Pico-Potr
Potr
D. Mead., + Arfr
D. Mead., - Arfr
Wet Mead.
Ir. Mead.
SAL Mead.
Cemo-Putr
Putr-Cemo
JUNI-Shrub
Riparian
Logged-recent
Rock-outcrop
Rock talus
Dry Gull Bott.
Water
Spr.-fir
Cult. Fields
Surfaced road
Unsur. road

Ponderosa pine, open stand
Ponderosa pine, dense stand
Ponderosa pine-Douglas fir
Douglas fir
Douglas fir-ponderosa pine
Lodgepole pine-open stand
Lodgepole pine-dense stand
Lodgepole pine-Douglas fir
Lodgepole pine-aspen
Aspen
Dry meadow, with fringed sage
Dry meadow, without fringed sage
Wet meadow
Irrigated meadow
Willow meadow
Mountain mahogany-bitterbrush
Bitterbrush-mountain mahogany
Juniper spp-other shrub species

Dry gulley bottom
Spruce-fir
Cultivated

fields

�-244-

LITERATURE

CITED

in migratory
Anderson, A. E., and D. E. Medin.
[1967]. The breeding season
mule deer. Game Info. Leaflet No. 60. Colorado Game, Fish and Parks
Dept. 4 p.
Anderson, A. E. 1977. Experimental deer inventory, northeast region.
Job
Progress Report.
Pp. 227-250.
In Colo. Div. Wildl. Game Res. Rept.,
July, Part 2. 125-305 pp. (Processed).
1976. Evaluation of herd structure
Bowden, D. C., and A. E. Anderson.
Work Plan 19, Job 2, Job Progress Report.
PP. 505-554.
methodology.
Div.
Wildl.
Game
Res.
Rept.,
July,
Part
2.
253-561 pp.
In Colorado
(Processed).
Patton, D. R., and W. B. Casner.
1970. Port-A-Punch recording and computer
summarization of pellet count data. U.S.D.A. Forest Service Research
Note RM-170.
8 p.

Prepared

by --.--:L='=tt!::::Yb=-J2..L1u----b::C::.:..- -",U,...::...=.,-~-=-=-' _,~-,--Allen E. Anderson
Wildlife Researcher

_

�July

1978

- 245--

JOB PROGRESS

REPORT

State of

COLORADO
------------~---------------

Project

No.

W-38-R-33

Plan No.

22

Job No.

Experimental

Deer Inventory-Piceance

Work

Job Title
Period

Covered:

Personnel:

April

_

Deer Elk Investigations
1 (NW)

----------~~------------------Basin-Northwest

Region

1, 1977 - June 30, 1978

D. Freddy, D. Bowden, D. Bartmann, L. Carpenter, B. Gill, K.
Karrow, C. Moore, D. Osborne, R. Kolecki, L. Strong, T. Tietz.

ABSTRACT

A stratified sampling system capable of estimating density of mule deer
pellet groups within ± 15% of the mean 90 percent of the time was established
for the Piceance Basin.
This system employed a square-mile as the sample
unit which was then subsampled by 2 transects, ~ mile apart, each containing
Sampled
10 temporary plots, 107.64 ft2 in size, spaced at 0.1 mile intervals.
density of new groups for the winter period of 1977-78 was 15.94 + 2.06 (P &lt; .10)
per square-mile.
Projected population estimates were higher than-expected ~nd
probably resulted from errors in aging "new" pellet groups.
A permanent versus
temporary plot test was established on 10 square-miles in 1977 to quantify the
differences in estimates of new groups per square-mile by the two types of
plots.
Data from 1978 indicated temporary plots significantly inflated number
of new grounds per square-mile (P &lt; .10). Aerial versus ground methods of
classifying deer as to sex and age-were compared.
Discrepancies
of up to 20
fawns: 100 does did not result in significant differences between methods.

��-247-

EXPERIMENTAL

DEER INVENTORY-PICEANCE

BASIN-NORTHWEST

REGION

David J. Freddy

PROGRAM NARRATIVE

OBJECTIVE

Design appropriate sampling and analytical procedures
deer numbers and buck:doe:fawn ratios in the Piceance
Unit 22.

to reliably estimate
Basin, Game Management

SEGMENT OBJECTIVES
1.

Sample pellet group density to aid in designing a permanent sampling
system capable of estimating deer numbers within + 10% at P &lt; .10.

2.

Investigate sampling methods
and age ratios.

that might be used to estimate

deer sex

METHODS AND MATERIALS
Pellet Group Density
A long-term system for estimating deer pellet group densities in the
Piceance Basin was established based on a preliminary sample in 1976 (Freddy
1977). Winter range boundaries were slightly refined from 1976 due to the
presence of domestic sheep. The 606 square-mile winter range area to be
sampled for pellet group density was divided into 3 major strata (Freddy 1977)
(Fig. 1). Each strata was further divided into 3 blocks of nearly equal size.
Blocking was done to insure representative sampling of the entire strata to
account for potential changes in deer distribution within strata.
Sampling
effort was allocated between strata (by block) by optimal allocation (Cochran
1977) based on data acquired in 1976. Square-mile sample units were drawn
independently wirhout replacement for each block and thus by definition
(Cochran 1977) there were 9 sampling strata.
A total of 54 square-miles were
selected at random; 12 in strata 1, If in strata 2, and 26 in strata 3.
Square-mile sample units were sampled for pellet group density using 2
transects with 10, 107.64 ft2 (.001 ha) circular plots spaced at 0.1 mile
(0.16 km) intervals (Freddy 1977). Transects were selected from 10 possible
transects spaced at 0.1 mile (0.16 km) intervals (Freddy 1977). One transect was
selected at random from transects 1-5 with the second transect systematically
located ~ mile from the first transect.
Transects and plots were located by
pacing with the aid of a hand-held compass and topographic maps (Freddy 1977).
Plots were delineated by a 5.85 ft (1.784 m) chain revolving about a metal rod.
Transects intercepted maximum elevation change within the square-mile and were
oriented on either a true north or east bearing.
No stakes or markers were
installed to relocate transects or plots.

�-248-

Four persons working in two, 2-man teams, sampled 2 square-miles per day.
Rules for examining pellet plots and definitions of pellet groups are
given in Freddy (1977).

Permanent

Plot Test

While designing a long-term sampli.ng system, the question arose as to the
necessity of using pern~nently marked pellet plots and transects.
When most
of ,the wintering deer population migrates to higher elevations outside winter
range boundaries in late spring, only a small percentage of that population
resides on that winter range in summer.
Empirical methods such as aerial
mapping of winter distribution of deer and subjectively aging pellet groups
according to season of deposition may be sufficient to identify winter pellet
groups less than 1 year old, thus eliminating two major reasons for using
permanent plots which are more time consuming to install, maintain, relocate,
and read.
To ascertain quantitative advantages or disadvantages of permanent and temporary
plots, a permanent plot test was designed.
Ten square-miles
(25.9 km2)
representative
of the Piceance Basin and having a high deer pellet group density
were selected.
A grid of 10 transects at 0.1 mile (0.16 km) intervals each
with 10 plots at 0.1 mile (0.16 km) intervals was delineated for each squaremile (Freddy 1977).
Five pairs of transects with one permanently marked transect
and one temporary transect in each pair one were established on each square-mile.
Permanent and temporary transects alternated with the first transect in each
square-mile having been randomly selected as a temporary transect.
With this
design the mean and variance of new groups per square-mile can be compared for
types of sampling systems (permanent vs. temporary plots) using a paired t-test
(Sokal and Rohlf 1969).
Permanent and temporary plots were again located by pacing with the aid of a
hand-held compass and topographic maps.
Numbers of new and old pellet groups
were determined for both temporary and permanent plots.
For the initial
establishment,
new groups were subjectively determined on both types of plots.
Permanent plots were cleared of all pellet groups, marked with a 2 ft steel
stake painted orange and tagged with an aluminum tag. High visibility yellow
flagging was tied to trees, etc. to aid in relocating permanent plots.
All
transects intercepted maximum elevation change within each square-mile and
were oriented on a true east bearing.
This test will be conducted for 3 years.
As an aid to subjectively "aging" pellet groups found on temporary plots,
aging plots were established on 5 microsights.
Known new pellet groups were
collected after recent snowfalls during the winter of 1977-78 and placed on
aging plots at monthly intervals from December through March.
These groups
were examined prior to conducting the permanent plot test to establish aging
criteria for new groups.
Sex and Age Ratios
Aerial vs. ground methods to classify deer as to bucks, does, and fawns, have produced significantly different results (Freddy 1977). To further test the effect of

�-249-

methodology, deer buck:doe and fawn:doe ratios were sampled on two, 100
square-mile areas (259 km2) by both aerial and ground methods.
Four
observers having 9, 5, 5, and 3 years experience in classifying deer,
classified deer as to bucks, does, and fawns during early December.
Two
observers were assigned to each 100 square-mile (259 km2) area (Areas A &amp;
B).
Both areas A and B were considered to be high deer density areas.
Within each area, 14 one square-mile sample units were selected at random,
without replacement, for each observer.
Square miles selected were restricted
so they could not have adjoining boundaries but could have cornmon corners.
Each observer then walked a route of his choice through each sample unit and
classified as many deer as he could without duplicating counts of the same
deer.
These walking routes were situated to enhance observations of deer and
minimize terrain variance.
Two square-miles were sampled by each observer
each day.
To standardize criteria for identifying does, fawns, and bucks, all
observers classified deer together for one day prior to conducting the test.
Each observer then classified deer from a Bell r,4713-1helicopter approximately
1 week after walking the sample units.
Each observer classified deer from the
helicopter on the same sample units he had walked and in the same sequence
he had walked them.
This design resulted in 14 paired observations of buck:
doe and fawn:doe ratios for each observer.
Additionally, buck:doe and fawn:doe
ratios of observers classifying deer within the same 100 square-mile
(259 km2)
sample area could be compared
to test for observer differences.
Ratio
estimator tests were used for all statistical comparisons (Cochran 1977; 181).

RESULTS
Pellet

AND DISCUSSIONS
Group Counts

group
counts were conducted from 18 May through 27 June.
A mean of
15.94 + 2.06 (P &lt; .10) new groups were found per square mile (6.15 + 0.80 km2)

Pellet

(Table-I, 2). A-two-way analysis of variance (unequal numbers within cells)
revealed significant differences in new and old pellet groups between the 3
major strata and no significant differences in new or old groups between blocks
within strata (P ~ .05).
Regressions of new groups (Y) per square-mile on old
groups (X) per square-mile were positive and significant for all strata.
Correlation coefficients
(r) for individual strata were 0.686, 0.438, and 0.617
for strata 1-3 respectively and the pooled correlation coefficient was 0.740.
These positive relationships are similar to those previously generated (Freddy
1977) and support the validity of delineating strata on the basis of total
groups.
Starting and ending times were recorded for each transect.
Mean time per
transect was 93.9 + 2.8 (SE) minutes which was not significantly different
than the 92.6 + 1.5 (SE) minutes per transects for REP 1 in 1976 (Freddy 1977)

(P ~ .05).
Deer numbers were estimated using 13 pellet groups/day/deer
and projecting
the 15.94 new groups per 20 plots to a square mile and then projecting this
figure to 606 square miles of sampled winter range.
Specific deer-days use

�-250-

Table 1. Location of square mile sample units and sampled pellet group
densities, Piceance Basin, Colorado, 1977.

Strata

Block

Sample
Unit If

Location

Transects
Sampled

I

I

1-1-1

Sec 31 T2N R98W

3, 8

37

9

46

1

1

1-1-5

See 5

2,

7

34

6

40

1

1

1-1-12

See 13 TIN R99W

1, 6

34

4

38

1

1

1-1-23

See 25 TIN R99W

2,

7

19

8

27

1

2

1-2-4

See 13 TIS R99W

2,

7

34

o

34

1

2

1-2-9

See 23 TIS R99W

4, 9

40

3

43

1

2

1-2-15

See 27 TIS R99W

4, 9

29

5

34

1

2

1-2-16

See 32 TIS R99W

1, 6

26

7

33

1

3

1-3-15

See 4

T3S R98W

2,

7

42

6

48

1

3

1-3-27

Sec 7

T3S R97W

1, 6

32

10

!~2

1

3

1-3-40

See 21 T3S R97W

1, 6

62

24

91

1

3

1-3-42

See 29 T3S R97W

1, 6

34

7

41

2

1

2-1-3

See 3

T2N R98W

2,

7

46

10

56

2

1

2-1-43

See 6

TIN R97W

5, 10

27

3

30

2

1

2-1-45

See 4

TIN R97W

4,

9

44

6

50

2

1

2-1-65

See 17 TIN R97W

2,

7

49

7

56

2

1

2-1-86

See 33 TIN R97W

4,

9

64

17

81

2

2

2-2-10

See 3

TIS R98W

5, 10

53

9

62

2

2

2-2-20

See 10 TIS R98W

3, 8

31

9

40

2

2

2-2-32

See 18 TIS R97W

2,

7

48

14

62

2

2

2-2-51

See 25 TIS R98W

1, 6

81

18

99

2

2

2-2-84

See 25 T2S R98W

5, 10

30

9

39

2

3

2-3-12

See 31 TIS R95W

3, 8

48

4

52

2

3

2-3-24

See 3

T2S R95W

3, 8

33

28

61

2

3

2-3-27

See 9

T2S R96W

3, 8

63

23

86

2

3

2-3-45

See 13 T3S R97W

1, 6

57

21

78

2

3

2-3-65

Sec 28 T3S R96W

2,

7

45

3

48

2

3

2-3-83

See 5

5, 10

61

33

94

TIN R98W

T4S R96W

Pellet-groups/20 plots
Old
New
Total

�-251-

Table 1. Location of square mile sample units and sampled pellet group
densities, Piceance Basin, Colorado, 1977. (Cont'd).

Location

Transects
Sampled

Pellet-group/20 plots
Old
New
Total

Strata

Block

Sample
Unit It

3

1

3-1-2

See 35 T3N R99W

3, 8

75

27

102

3

1

3-1-4

See 31 T3N R98W

2,

7

85

23

108

3

1

3-1-5

See 5

T2N R99W

4,

9

39

9

48

3

1

3-1-6

See 4

T2N R99W

3, 8

70

23

93

3

1

3-1-12

See 9

T2N R99W

5, 10

109

31

140

3

1

3-1-19

See 14 T2N R99W

5, 10

91

22

113

3

1

3-1-50

See 21 TIN R96W

5, 10

70

42

112

3

1

3-1-53

See 25 TIN R97W

5, 10

64

13

77

3

1

3-1-65

See 35 TIN R96W

5, 10

100

48

148

3

2

3-2-7

See 5

2,

7

80

23

103

3

2

3-2-12

See 11 TIS R97W

3, 8

45

23

68

3

2

3-2-14

See 7

TIS R96W

4, 9

86

25

III

3

2

3-2-33

See 18 TIS R94W

4, 9

38

14

52

3

2

3-2-49

See 27 TIS R97W

5, 10

58

8

66

3

2

3-2-54

See 28 TIS R96W

2,

7

60

27

87

3

2

3-2-63

See 36 TIS R97W

2, 7

68

22

90

3

2

3-2-64

See 6

5, 10

81

15

96

3

2

3-2-17

See 11 TIS R96W

4,

9

83

22

105

3

3

3-3-6

See 16 T2S R97W

4, 9

65

24

89

3

3

3-3-10

See 18 T2S R96W

5. 10

99

54

153

3

3

3-3-20

See 21 T2S R96W

4, 9

64

22

86

3

3

3-3-23

See 28 T2S R97W

2,

7

58

19

77

3

3

3-3-27

See 30 T2S R96W

3, 8

56

13

69

3

3

3-3-32

See 25 T2S R96W

5, 10

85

58

143

3

3

3-3-51

See 5

2,

7

45

17

62

3

3

3-3-69

See 17 T3S R96W

5, 10

60

21

81

TIS R95W

T2S R97W

T3S R96W

�Table 2. Numbers of new pellet groups counted on square-mile sample units within 9 sampling strata and
perti~ent statistics, Piceance Basin, Colorado, 1977.

Block 1

Strata 1
Block 2

Block 3

Block 1

Strata 2
Block 2

Block 3

Block 1

Strata 3
Block 2

Block 3

9

0

24

10

9

4

27

23

24

6

3

7

3

9

28

23

23

54

4

5

6

6

14

23

9

25

22

8

7

10

7

18

21

23

14

19

7

9

3

31

8

13

33

22

27

58
I

42

22

17

N
\..rI
N

I

nh

4

4

4

2:Yh

27

15

47

Yh

6.75

3.75

Nh

44

S2
h

4.92

13

15

48

22

21

5·

6

9

9

8

33

59

112

238

179

228

11.75

8.60

11.80

18.67

26.44

19.89

28.50

45

44

87

88

85

70

72

71

8.92

69.58

28.3

16.7

155.47

157.03

38. 1J.

300.29

-

5

---------------------------------------------------------------------------------------------------------

2:Nh= 606

yst = 15.94 V(yst) = 1.46

S(yst) = 1.21

df = 29

c.r.= 15.94 + 2.06 (P ~ .10)

�-253-

were unknown so population estimates were made for several possibilities
(Table 3). All population estimates were considerably higher than the
estimate of 29,820 for the same winter period made by Bartmann (1977)
from aerial counts of deer. The large difference in estimates of population
size are probably due to errors in aging pellet groups and are not artifacts
of the sampling design (Personal Communication Dave Bowden).

Table 3. Estimates of deer population size for varying deer-days use,
Piceance Basin, Colorado, 1977. Estimates based on 13 pellet groups/day/deer.
Deer
Days

Estimated
Mean Population

210

45,887

+5825

180

53,535

+6796

150

64,242

+8155

120

80,303

±10,193

Confidence

Interval

P &lt; .10

The design of 2 transects per square mile was logistically feasible.
The
precision about the mean (± 12-13%) was also acceptable.
Thus, the candidate
sampling design appears promising as an estimator of pellet group density.
The primary problem appears to involve aging of pellet groups which could
be solved by using permanent pellet plots.
Sample size requirements estimated from data collected in 1976 indicated
that 43 square miles should be sampled to obtain a precision of ± 10% of x at
2
P ~ .10. This precision was not achieved by sampling 54 square miles (140 km )
in 1977.
Permanent

Plot Test

1977
Permanent plots were established and cleared from 6 July through 17 August,
1977. Numbers of new, old, and total groups per square mile on temporary
and permanent plots were:
37.0 + 3.1 (SE) new, 164.7 + 7.8 (SE) old, 201.7
± 8.3 (SE) total and 40.2 ± 3.5 (SE) new, 167.4 ± 12.2-(SE) old, 207.6 ± 15.0
(SE) total, for temporary and permanent plots, respectively (Table 4). There
were no significant differences in numbers of new, old and total pellet groups
per square mile when estimated by either temporary or permanent plots (P ~ .01).
These "pre-treatment" data suggest that differences between transect types occurring in subsequent years will be due to problems of aging pellet groups on
temporary plots and not due to differences in transect and/or plot locations.

�-254-

Table 4. Location of square-mile sample units sampled by temporary and
permanent transects and summary of sampled pellet group densities, Piceance
Basin, Colorado, July-August, 1977 .
Sample
Unit

Location

Transecta

4-18

Sec 18 TIS R96W

1

64

5

69

2

60

9

69

3

40

15

55

4

33

17

50

5

24

8

32

6

32

14

46

7

42

5

47

8

58

12

70

9

34

7

41

10

30

8

38

E

204

40

244

E

213

60

273

1

27

5

32

2

41

9

50

3

38

7

45

4

43

14

57

5

26

5

31

6

27

5

32

7

49

6

55

8

52

2

54

9

59

6

65

10

38

8

46

E

199

29

228

E

201

38

239

1

31

3

34

2

39

11

50

3

16

4

20

4

24

4

28

5

32

4

36

6

45

8

53

7

36

18

54

8

31

10

41

9

20

1

21

10

31

3

34

E

135

30

165

L-

170

36

206

1

43

16

59

2

28

11

39

3

29

16

45

4

23

13

36

5

19

7

26

6

24

4

28

7

19

3

22

8

15

5

20

9

22

6

28

10

18

7

25

E

132

48

180

E

108

40

148

4-19

4-30

4-31

Sec 19 TIS R96W

Sec 30 TIS R96W

Sec 31 TIS R96W

Pellet GrouEs
Old New Total

b
Transect

Pellet GrouEs
Old New Total

--------------------------------------------------------------------------------

�-255-

Table 4. Location of square-mile sample units sampled by temporary and
permanent transects and summary of sampled pellet group densities, Piceance
Basin, Colorado, July-August, 1977 . (Cont'd).
Sample
Unit

Location

Transect

4-12

Sec 12 TIS R97W

1

29

12

41

2

23

5

28

3

32

7

39

4

52

7

59

5

39

3

42

6

38

8

46

7

37

11

48

8

30

7

37

9

37

8

45

10

47

22

69

E

174

41

215

E

190

49

239

1

36

8

44

2

29

5

34

3

50

3

53

4

10

1

11

5

17

3

20

6

19

4

23

7

29

4

33

8

20

4

24

9

27

4

31

10

30

5

35

2:

159

22

181

E

108

19

127

1

33

7

40

2

46

13

59

3

26

15

41

4

39

15

54

5

57

6

63

6

57

6

63

7

34

9

43

8

37

11

48

9

34

8

42

10

33

4

37

E

184

45

229

E

212

49

261

1

21

6

27

2

17

3

20

3

26

5

31

4

45

9

54

5

37

5

42

6

25

1

26

7

30

7

37

8

35

14

49

9

40

12

52

10

48

12

60

E

154

35

189

E

170

39

209

1

39

9

48

2

33

8

41

3

25

5

30

4

26

4

30

4-13

4-24

4-25

4-14

Sec 13 TIS R97W

Sec 24 TIS R97W

Sec 25 TIS R97W

Sec 14 TIS R97W

a

Pellet GrouEs
Old New Total

Transect

b

Pellet GrouEs
Old New Total

----~-------------------------------------------------------------------------

�-256-

Table 4. Location of square-mile sample units sampled by temporary and
permanent transects and summary of sampled pellet group densities, Piceance
Basin, Colorado, July-August, 1977 • (Cont'd).
Sample
Unit
4-14

4-23

Location

a
Transect

Sec 14 TIS R97W

Sec 23 TIS R97W

Grand Sum

Pellet GrouEs
Old New Total

Transect

b

Pellet GrouEs
Old New Total

5

19

3

22

6

24

5

29

7

30

4

34

8

32

8

40

9

41

7

48

10

29

9

38

z

154

28

182

l:

144

34

178

1

9

6

15

2

31

9

40

3

34

9

43

4

32

3

35

5

28

8

36

6

27

6

33

7

34

17

51

8

30

14

44

9

47

12

59

10

38

6

44

l:

152

52

204

l:

158

38

196

H

1647 370

2017

H

1674 402

2076

plots

aOdd-numbered

transects

contained

temporary

bEven-numbered

transects

contained

permanently

staked plots

In 1977 time per transect for temporary transects was 67.7
and for permanent transects 140.4 + 4.2 (SE) minutes.

+ 1.8 (SE) minutes

1978
In 1978, the permanent plot test was conducted from 26 April through 31 May.
Two persons working as a team evaluted 2 permanent and 2 temporary transects
daily.
On 25 April, 1978 aging plots were examined to establish criteria for judging
new groups.
Criteria were:
1) individual pellets associated in recognizeable
groups,
2) pellets dark, uniform, and intensely colored,
3) sheen present
on entire surface of most pellets, and 4) some cracking of the pellet surface.
On 23 May, aging plots were again examined.
Criteria for new groups were:
1) individual pellets still closely associated in recognizeable groups,
2)
pellets dark in color but fading on some pellets producing a mild two-toned
color effect,
3) sheen present, although not uniform over entire pellet, and
4) cracking and peeling of the pellet surface more evident.

�-257-

There was a significant difference in numbers of new groups per square-mile
estimated by temporary and permanent plots (P &lt; .10 but not P .2. .05).
Temporary plots averaged 9 groups per square-mile (3.5/km2) more than
permanent plots, (Table 5), about a 14 percent increase.
These data indicate
temporary plots inflate the estimate of new groups per square-mile.
On
permanent plots, some subjectivity was needed in determining new groups,
as obvious "old" pellet groups had washed onto the plots.
These groups
were not tallied in new group totals.

Table 5. Sampled pellet group densities on temporary and permanent transects
on 10, square-mile sample units, Piceance Basin, Colorado, April-May 1978.
Sample
Unit

Transect

4-18

4-19

4-30

4-31

New Pellet
Groups

Transect

1

18

2

33

3

22

4

10

5

24

6

11

7

18

8

24

9

24

10

23

L:

106

L:

101

1

18

2

17

3

25

4

18

5

13

6

18

7

16

8

23

9

12

10

18

L:

84

L:

94

1

17

2

10

3

13

4

16

5

24

6

14

7

20

8

12

9

14

10

11

L:

88

L:

63

1

27

2

11

3

9

4

9

5

22

6

8

a

b

New Pellet
Groups

�-258-

Table 5. Sampled pellet group densities on temporary and permanent transects
on 10, square-mile sample units, Piceance Basin, Colorado, April-May 1978.(Cont'd).
Sample
Unit

Transect

4-31

7

2

8

7

9

24

10

14

a

New Pellet
Groups

Transect

84
4-12

1

17

2

13

3

9

4

13

5

7

6

7

7

16

8

4

9

7

10

16

53

1

15

2

14

3

19

4

6

5

12

6

6

7

14

8

1

9

6

10

20

66
4-24

47

1

10

2

13

3

11

4

14

5

19

6

10

7

9

8

17

9

16

10

12

65
4-25

New Pellet
Groups

49

56
4-l3

b

66

1

9

2

15

3

16

4

15

5

10

6

7

7

4

8

9

�-259-

Table 5. Sampled pellet group densities on temporary and permanent transects
on 10, square-mile sample units, Piceance Basin, Colorado, April-May 1978.
(Cont'd).
Sample
Unit

Transect

4-25

New Pellet
Groups

Transect

9

14

10

8

I

53

I

54

1

19

2

9

3

23

4

8

5

19

6

7

7

8

8

28

9

17

10

23

I

86

I

75

1

10

2

6

3

12

4

8

5

7

6

7

7

5

8

8

9

13

10

13

I

47

I

42

Grand Sum

II

735

II

644

aOdd-numbered

transects

contained

temporary

bEven-numbered

transects

contained

permanently

4-14

4-23

a

b

New Pellet
Groups

plots
staked plots

In 1978, time per transect was 68.8 + 1.43 (SE) minutes for temporary transects
and 85.1 ± 3.04 (SE) minutes for permanent transects.
These values were
significantly different (P &lt; .05). Differences between mean times for temporary
transects for 1977 and 1978-were not significant (P &lt; .05).
Of the 500 permanently staked plots, 4 were missing.
In two instances yellow
flagging remained to denote general location of the missing plot. Missing
plots were restaked in an estimated location and recleared.
The orange stakes
and yellow flagging greatly aided in finding permanent plots.
There was virtually
no problem in relocating permanent plots spaced at 0.1 mile (0.16 km) intervals.

�-260-

Sex and Age Ratios
Ground routes were conducted from 29 November through 5 December and aerial
counts were conducted from 9 December through 11 December, 1977.
Significant differences in fawn:doe ratios between aerial and ground methods
were detected for only 1 of 4 observers (observer F, Tables 6, 7). There
were no significant differences between air and ground methods in buck:doe
ratios for any observer (P ~ .10). Differences of up to 20 fawns:lOO does
were not statistically significant.
This indicated a great deal of variability
within both the aerial and ground samples.
In determining variances of fawn:
doe ratios, all calculated covariance terms were negative.
This suggested
that different fractions of the deer population occuring on a sample unit were
possibly more visible by one method or that observers inconsistently classified
deer between methods.
There were several neck-banded deer observed on the
study area. Neck-banded deer seen during ground routes were often seen on the
same sample unit during aerial sampling, suggesting no large distributional
shift of deer during the experiment.
Within area A there were no differences
in fawn:doe or buck:doe ratios between observers within methods (P ~ .10).
However, in area B there were significant differences in buck:doe ratios
between observers within both methods and there was also a significant difference in fawn:doe ratios between observers within the aerial sample (P &lt; .10).

ACKNOWLEDGMENTS
Dr. David C. Bowden, Colorado State University,
statistical design and calculations.

LITERATURE
Cochran, W. G. 1972.
York.
428pp.

Sampling

provided

techniques.

1969.

in

CITED
John Wiley and Sons, New

Freddy, D. J. 1977. Experimental deer inventory-Piceance
Div. Wildl. Game Res. Rep. July, Part 2:251-273.
Sokal, R. R., and F. J. Rohlf.
San Francisco.
776pp.

assistance

Biometry.

Basin.

Colo.

W. H. Freeman and Co.,

Bartmann, R. M. 1978. Piceance deer study-estimating parameters of
population dynamics.
Colo. Div. Wildl. Game Res. Rep. July Part 1:
In Press.

�Table 6. Number of deer classified as to sex and age by 4 observers on paired aerial and ground square-mile
sample units, Piceance Basin, Colorado, 29 November through 11 December, 1977.

AREA A
(GROUND
SAMPLE)

AREA A
(AERIAL
SAMPLE)

OBSERVER

SAMPLE
UNIT

BUCKS

F
F
F
F
F
F
F
F
F
F
F
F
F
F

1-1
1-2
1-3
1-4
1-5
1-6
1-7
1-8
1-9
1-10
1-11
1-12
1-13
1-14

1
0
8
2
2
2
1
2
2
0
1
5
2
2

TOTALS
RATIOS

14

122
30
200
352
15 BUCKS: 100 DOES:61 FAWNS

F
F
F
F
F
F
F
F
F
F
F
F
F
F

1-1
1-2
1-3
1-4
1-5
1-6
1-7
1-8
1-9
1-10
1-11
1-12
1-13
1-14

4
7
2
2
1
5
3
2
1
11
3
5
1

TOTALS
RATIOS

14

47
251
195
493
19 BUCKS: 100 DOES:78 FAWNS

0

DEER CLASSIFIED
DOES
FAWNS
1
6
26
24
8
17
29
12
7

6
32
16
9
7

12
6
18
25
15
23
26
14
18
29
23
23
16
3

2
4
10
12
5
7
19
8
6
8
24
8
5
4

11
. 8
14
16
5
17
18
7
16
18
20
28
15
2

TOTAL

OBSERVER

SAMPLE
UNIT

BUCKS

4
10
44
38
15
26
49
22
15
14
57
29
16
13

C
C
C
C
C
C
C
C
C
C
C
C
C
C

2-1
2-2
2-3
2-4
2-5
2-6
2-7
2-8
2-9
2-10
2-11
2-12
2-13
2-14

7
5
4
1
0
0
3
1
1
7
0
1
1
0

TOTALS
RATIOS

14

31
166
109
306
19 BUCKS: 100 DOES:66 FAWNS

C
C
C
C
C
C
C
C
C
C
C
C
C
C

2-1
2-2
2-3
2-4
2-5
2-6
2-7
2-8
2-9
2-10
2-11
2-12
2-13
2-14

4
2
1
0
1
0
3
1
0
3
2
1
6
2

TOTALS
RATIOS

14

26
163
140
329
16 BUCKS:100 DOES:86 FAWNS

27
21
34
43
21
45
47
23
35
58
46
56
32
5

DEER CLASSIFIED
DOES FAWNS
TOTAL
15
34
5
6
2
8
15
22
7
42
2
5
3
0

5
26
11
0
7
8
20
29
1
18
10
2
20
6

10
19
2
4
3
3
12
20
4
20
3
7
2
0

4
29
11
0
3
8
12
16
3
26
9
2
12
5

32
58
11
11
5
11
30
43
12
69
5
13
6
0

13
57
23
0
11
16
35
46
4
47
21
5
38
13

I
N

•...

0\

I

�Table 6. Number of deer classified as to sex and age by 4 observers on paired aerial and ground square-mile
sample units, Piceance Basin, Colorado 29 November through 11 December, 1977. (Cant'd) .

AREA B
(GROUND
SAMPLE)

OBSERVER

SAMPLE
UNIT

BUCKS

40
3
29
7
4
5
0
5
23
14
46
41
10
46

G
G
G
G
G
G
G
G
G
G
G
G
G
G

4-1
4-2
4-3
4-4
4-5
4-6
4-7
4-8
4-9
4-10
4-11
4-12
4-13
4-14

7
1
0
0
2
0
2
5
6
3
2
0
0
3

TOTALS
RATIOS

14

120
323
31
172
18 BUCKS: 100 DOES:70 FAWNS

G
G
G
G
G
G
G
G
G
G
G
G
G
G

4-1
4-2
4-3
4-4
4-5
4-6
4-7
4-8
4-9
4-10
4-11
4-12
4-13
4-14

4
0
1
1
0
1
6
5
1
3
3
3
2
0

TOTALS
RATIOS

14

148
234
30
412
13 BUCKS: 100 DOES:63 FAWNS

3-1
3-2
3-3
3-4
3-5
3-6
3-7
3-8
3-9
3-10
3-11
3-12
3-13
3-14

5
0
10
3
2
2
0
0
4
2
7
7
0
4

TOTALS
RATIOS

14

273
131
96
46
35 BUCKS: 100 DOES: 73 FAWNS

B
B
B
B
B
B
B
B
B
B
B
B
B
B

3-1
3-2
3-3
3-4
3-5
3-6
3-7
3-8
3-9
3-10
3-11
3-12
3-13
3-14

11
2
5
4
0
4
0
3
6
10
1
0
0
6

TOTALS
RATIOS

14

387
216
119
52
24 BUCKS: 100 DOES:55 FAWNS

B
B
B
B
B
B
B
B
B

18
1
13
4
2
2
0
2
13
7
17
19
7
26

38
12
17
14
6
9
2
22
19
17
12
18
0
30

17
2
6
0
0
1
0
3
6
5
22
15
3
16

27
3
15
11
3
6
2
6
11
10
7
8
0
10

DEER CLASSIFIED
FA\\TNS TOTAL
DOES

TOTAL

BUCKS

B
B
B
B
B

AREA B
(AERIAL
SAMPLE)

DEER CLASSIFIED
DOES
FAWNS

SAMPLE
UNIT

OBSERVER

76
17
37
29
9

19
4
31
36
37
20
26
0
46.

23
10
8
3
6
4
9
10
48
9
22
4
1
15

35
7
11
8
8
1

34
26
36
13
22
18
11
4

17
9
9
2
3
2
6
7
36
4
12
0
0
13

22
2
5
5
5
0
21
16
25
9
12
12
10
4

47
20
17
5
11
6
17
22
90
16
36
4
1
31

61
9
17
14
13
2
61
47
62
25
34
33
23
8

I

tv
(j\

tv
I

�-263Table 7. Fawn:doe and buck:doe ratios resulting from ground and aerial
classifications of deer, Piceance Basin, Colorado, 1977.

Observer

Fawn:doe ratio (+SE)
Ground
Aerial

Buck:doe ratio (+SE)
Ground
Aerial

F

.6100 + .0529

.7769 + .0624

.1500 + .0387

.1873 + .0412

C

.6566 + .0714

.8589 + .1114

.1867 + .0400

.1595 + .0400

B

.7328 + .1005

.5509 + .0640

.3511 + .0671

.2407 + .0424

G

.6977 + .0447

.6325 + .0245

.1802 + .0361

.1282 + .0224

��-265July, 1978

JOB PROGRESS

State of

COLORADO
------~~~~~--------

Proj ect No.

W-38-R-33

Deer-Elk

Investigations

I
---------------------------Experimental Deer Inventory - Southeast Region
~~-----------------------

Work Plan No.
Job Title

REPORT

Period Covered:

22

Job No.

c

April 1, 1977 - June 30, 1978

Personnel:
Katherine Ortega, Bryan Gross, John Ernst, Michael A. Knapp,
Thomas. Valenta, Bert A. Jellison, Stan Ogilvie, Dwayne Finch, Cathy Craig,
Thomas A. Lytle, Willie Travnicek, Harold Hood, Jeff Kogutt, Dr. David
Bowden (CSU Statistics Department), Thomas M. Pojar.
To the many landowners
who expressed their interest and cooperation in this study by allowing us
trespass permission to their property I offer my sincere appreciation.
ABSTRACT
Permanent plots were located, marked and cleared of pellet groups on 24
square miles of the 1026 square mile study area near Canon City. Twentyseven herd structure routes were used to estimate buck:doe:fawn ratios.
Three hundred seventy nine deer were observed of which 336 were classified.
The ratio of bucks to does was .1803(SE = .0290) and the fawn: doe ratio was
.6557 (SE = .0454).

��-267-

EXPERIMENTAL DEER INVENTORY
SOUTHEAST REGION
Thomas M. Pojar

P. N. OBJECTIVE
Design appropriate sampling and analytical procedures necessary to reliably
estimate deer numbers and buck:doe:fawn ratios of selected management units
in the southeast region of the state.

SEG}ffiNTOBJECTIVES
1.

Design and execute a preliminary sampling system to estimate
pellet group density on Game Management Units 58 and 581.

2.

Design a sampling scheme based on the means variances, and deer
density strata estimated from objective 1 capable of estimating mean
deer numbers ± 20 percent at a = 0.10.

3.

Sample buck:doe:fawn
walking routes.

ratios on 12 or more randomly

METHODS

selected,

deer

all-day

AND MATERIALS

Pellet Group Density
Based on information on deer distribution and relative abundance collected
from temporary plots during the previous segment (Pojar 1977) the study area
boundaries were adjusted.
The overall area was reduced from 1,510 to 1026
square miles (Figure 1). The criterion for eliminating portions of the area
was the total lack of sign (either new or old pellet groups) and an arbitrary
judgement as to whether or not the area was potential deer habitat.
Areas
were retained even if few pellet groups were found, or if groups were found
in similar habitat.
The conjecture being, that if the area appeared to be
suitable habitat, based upon evidence that deer currently inhabit areas of
similar vegetative type, then the reason that little deer sign was present
may be that the population is in a trough of its temporal undulations and
an increase in population size would most likely result in habitation.
Because of the apparent difficulty of distinguishing "new" pellet groups
consistently, it was determined that cleared permanent plots would be necessary.
The square mile was retained as the basic sample unit but a new sampling
scheme within the square mile was needed because the logistics of locating
permanent plots had to be considered.
From a pilot study done in the northeast
region (Unit 19) (Allen Anderson, personal communication) it was determined
that location of permanent plot markers spaced one chain apart required about

�-268-

half the time to find as markers spaced two, four or eight chains apart.
The time to locate a marker was adjusted for travel time between markers.
Therefore the sampling scheme within a square mile was altered from
having a grid of plots located at 0.1 mile intervals covering the entire
area to a system of two one-mile transects with permanent plots located
at one chain intervals.
The transects were randomly selected in the
following manner:
Each square mile had 10 potential north-south transects
spaced 0.1 mile apart and numbered one through ten from west to east.
Using a random numbers table the first transect selected corresponded to
the first random number encountered between one and five, and the second
transect was arbitrarily located 0.5 miles east. Therefore if transect
number 3 was randomly selected as the first, then transect number 8 would
automatically be the accompanying transect for that square mile.
The
plots on each transect were numbered I through' 80 from north to south (Figure 2).
The permanent plot marker consisted of three-eigths inch steel rebar 18
inches long. The rebar stakes were driven into the ground 8 to 10 inches
and the top six inches of the stake was painted with royal blue enamal.
The plots at each end of the transect (number I and 80) were marked with
two stakes driven side-by-side and blue plastic surveyors tape was tied
to prominent objects nearby to facilitate location of the transect endpoints.
In addition, plot number one and every deca-plot thereafter was
marked with a metal tag indicating the section, transect, and plot number.
The transects and plots were located using a hand-held Silva Type #lSTD
compass, a steel 2-chain surveyors tape, and an abney level.
The circular 0.001 hectare plots were searched twice by two observers,
first in a clockwise direction then in a counter-clockwise
direction.
All deer pellet groups were classified as "new" or "old" by predetermined
criteria (Pojar 1977) and cleared from the plot. For each plot, the date;
identification by block, section, transect, and plot number; vegetative
type of immediate area (See appendix 1) coded observer numbers; and number
of new and old pellet groups were recorded on Porta-punch computer cards.
A detailed description and map of each square mile sampled was recorded
standard field forms (Appendix II).
Herd Structure

on

Estimates

The method described by Bowden and Anderson (1976) was used to estimate the
buck:doe:fawn ratios.
The only modification was that we used one observer
per route rather than two.
DESCRIPTION
See Pojar

(1977).

OF AREA

�-269-

RESULTS' AND DISCUSSION
Revision

of Study Area Boundaries

Revision of the exterior study area boundary necessitated also the revision of some of the block boundaries.
The original purpose of
establishing blocks was to ensure that there were sample square miles
from all portions of the study area yet maintain randomness of the samples.
The new block boundaries were located so all blocks would contain similar
total areas. This made it acceptable to select an equal number of random
square miles per block.
The new alignment resulted in 14 blocks with an
average size of 73.3 (range 57 to 81) square miles.
Two square miles
were randomly selected from each block.
Pellet Group Density
Permanent plots were located, marked, and cleared on 24 of the 28 square
miles during this segment.
The mean number of new groups found on sample
plots per square mile was 10.83. Adjusting these data to mean groups per
100 plots for comparison with the data from the previous segment, the mean
is 6.77 (SE = 2.83) compared with 6.47 (SE = 0.95) for the previous segment.
Any further use of this data would be unwise because of the substantial
evidence that classification of groups into the "new" or "old" category is
inaccurate.
Data to be collected during the next segment can be used to
estimate deer density and to evaluate the efficiency of various sampling
schemes within the square mile.
Herd Structure

Estimates

Twenty-seven all-day walking routes, to estimate the herd structure in terms
of bucks, does, and fawns, were completed between 13 and 21 December, 1977.
Four observers that trained together in the classification of deer made all
the observations.
A total of 379 deer were seen and of these, 336 were
classified (33 bucks, 183 does, and 124 fawns). The ratio of fawns to does
was .6557 (SE = .0454) and the ratio of bucks to does was .1803 (SE = .0290).
The ratio of f awns observed during this segment (1977) was significantly
(P &lt; .05) greater than the ratio of fawns (.4275, SE = .0598) observed during
last segment (1976). However, there was no significant difference (P &gt; .20)
in the rat f o of bucks in 1976 (.3043, SE = .0967) and 1977 (.1803, SE = .0290).
LITERATURE

CITED

Bowden, David C., and Allen E. Anderson. 1976. Evaluation of herd structure
methodology.
pp. 507-554.
In Colorado Div. of Wildl. Game Research
Report, July, Part Two. 253-561pp.
Pojar, Thomas M.
1977. Experimental deer inventory.
Southeast Region.
pp. 275-285.
In Colorado Div. of Wildl. Game Research Report, July
1977 Part Two. pp. 125-305.

�-270-

Table l. Legal descriptions of randomly selected square miles searched for
pellet groups and the number of groups found.

Block

Total
No. of
Miles2

Randomly
Selected
Mile2

I

78

5

12S

73W

10

63

13S

72W

21

52

15S

73W

2

3

4

5

6

7

8

9

74

72

77

80

78

78

70

80

Legal DescriEtion
Township Range Section

Transect
Number

No. of Pellet GrouEs
Old
New
Total

5
10
2
7

18
57
29
29

1
6
0
2

19
63
29
31

3

I

48
70
9
36

7

6
5
10

0
0

55
71
9
36

1
6
1
6

0
50
28
35

0
5
3
1

0
55
31
36

5
10
3
8

0
0
29
20

0
0
1
0

0
0
30
20

5
10
2
7

*
*

&gt;~

*

*"ie

51
47

4
1

55
48

5
10
5
10

197
209
142
183

4
18
2
7

201
227
144
190

4
9
5
10

244
165
95
109

42
31
11
9

286
196
106
118

5
10
4
9

126
48
281
243

7
3
20
10

133
51
301
253

1
6
3
8

17
6
34
16

2
0
2
0

19
6
36
16

67

15S

74W

14

36

14S

72W

27

64

15S

72W

7

7

13S

71W

14

54

15S

71W

3

26

16S

70W

26

67

18S

7moJ'

12

6

15S

70W

20

33

16S

71W

I

21

15S

72W

35

51

16S

73W

13

18

15S

73W

26

35

16S

73W

2

48

SON

12E

20

78

49N

lOE

2

1

---------------------------------------------------------------------------------

�-271-

Table 1. Legal descriptions of randomly selected square miles searched for
pellet groups and the number of groups found (cont'd).

Block

Total
No. of
Miles2

Randomly
Selected
Mile2

10

64

18

SON

12E

22

43

17S

73W

33

11

81

19
47

12

66

Legal DescriEtion
Township Range Section

17S
17S

73W
7lW

13
31

45

17S

7lW

36

46

17S

71W

35

Transect
Number

5
10
5
10
4
9
1
6
5
10
I

6
13

57

10

18S

73W

16

36

18S

73W

31

5
10
I

6
14

71

49

48N

lIE

5

59

48N

HE

14

5
10
I

6
Total

*

No. of Pellet GrouEs
Old
New
Total
85
70

4
4

89
74

*
*

*
*

*

*

*
*

*
*

*
*

*
*

*
*
*

120
55
119
l38

7
7
1
6

127
62
120
144

128
45
70
60

8
0
5

l36
45
71
65

103
49
114
151

2
6
10
8

105
55
124
159

3,978

269

4~247

*

I

Mean

165.7500

11.2083 176.9583

Std. Error

27.9801

3.0892

Data not collected during this segment.

30.3237

�-272-

Table 2. Buck:doe
27 all-day walking
December, 1977.

and fawn:doe ratios based on 336 deer classified on
routes in G.M.U.'s 58 and 581, between 13 and 21

Buck:Doe

Ratio

= .1803,

SE

.0290

Np. Routes Required
Confidence

Level

(I-a)

Percent*

95

90

80

5

1180

813

483

10

295

203

121

20

74

51

30

Ratio

= .6557,

Fawn:Doe

SE

.0454

No. Routes Required
Confidence

Level

(I-a)

Percent*

95

90

80

5

211

145

86

10

52

36

22

20

13

9

5

*Deviation within specified percentage
are based on the percentage

of the true ratio but calculation
of the sample ratio.

�-273-

Figure 1. Status of Canon City study area, 1978, after reV1Slon of total
area boundary and block boundaries.
Randomly selected square miles within
each block are outlined and numbered.

�-274-

•• ONE SQUARE MILE ••
1
I

2

3

J\'

4

5

,

6

7,

8,

9,

10

T

Plot III

I

N

I
I

I

I

,/Plot

1180

•

•

•

I
I

I
.I

i

t

,

Figure 2. Sampling design for estimating the pellet group density within
square mile.
Each transect consists of eighty consecutive plots at one
chain intervals numbered sequentially from north to south.
If transect
number three were drawn at random from transects 1-5, then number 8 is
selected as the second transect.

a

�-275-

Appendix
Vegetative

I

type classification

Code

Description

01

Pinon-Juniper

with grass or bareground

02

Pinon-Juniper

with shrubby

03

Ponderosa

pine - Open stand with grass and shrubby

04

Ponderosa

pine - Dense

05

Ponderosa

pine - Grass understory

06

Ponderosa

pine - Open - shrubby understory

07

Douglas

08

Aspen

09

Oakbrush

10

Oakb:r;ush- Open, grass understory

11

Mountain

mahogany

12

Grassland

- Dry

13

Grassland

- Wet

14

Bare ground

15

Misc.

understory

understory
understory

stand - bare ground or duff understory

fir

- Dense

�-276-

Appendix II

BLOCK

_

SECTION

_
&amp;

TRANSECTS
SECTION

TOWNSHIP __

CORNER

NW

RANGE

SECTION __

_

NE

SW

SE

OWNERS

A

_

B

_

C

_

SPECIAL

INSTRUCTIONS

FROM LAND OWNERS

�-277-

BLOCK

_

DESCRIPTION

SECTION

OF SECTION

CORNER

:,".

Prepared by

/ IL-t'&gt;K..(,,'./
..;~:."" .-'l1 "'?, i-IN)
.,~.
v

h"~.

'/

'

Thomas M. P6jar
/
Wildlife Researcher

&gt;

��-279-

JOB PROGRESS

July,

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

W-38-R-33

Work Plan No.

Deer-Elk

22

1978

Investigations

Job No.

1

--------------------------------Job Title
Experimental Deer Inventory - Southwest Region
--~--------------------~------------~-------------------Period

Covered:

Personnel:

April 1, 1977 through June 30, 1978

Roland C. Kufeld, Jim Olterman, David C. Bowden, Timms Fowler,
Harold L. Haeffner, Sam Creacy, Howard Geduldig

ABSTRACT
Deer classification counts were made between November 16, and December 4,
1977 on 25 "walking type" routes located at random throughout the winter
range portion of Game Management Unit 62. Resulting ratios and 90 percent
confidence limits were 28.1 ± 6.8 bucks per 100 does, and 46.2 + 4.9 fawns
per 100 does. Aerial deer classification counts were made on December 9 and
10, 1977 on the same 25 routes on which "walking type" counts were made.
Routes were covered by 2 observers seated side by side in a Hiller Soloy
helicopter.
Observers classified deer independently and without collaboration.
Resulting ratios and 90 percent confidence limits were:
Observer x, 22.5
+ 4.2 bucks/lOO does and 76.1 + 7.8 fawns/lOO does; Observer y, 23.5 + 5.1
bucks/100 does and 72.2 + 8.2 fawns/lOO does. The difference in doe:fawn
ratios between aerial and ground counts was significant at the 10 percent
level. Helicopter counts were made during February 11-18, 1978 on 193 quarter
section quadrats located randomly throughout the same area. The deer population
in Unit 62 with 90 percent confidence limits is estimated at 17,884 ± 4042
animals.
Thirty-one percent of the deer were found in strata 8, which is an
89 square-mile (230.5 km2) area encompassing the lower part of Log Hill Mesa.
Sixty-eight percent of the deer occur in the south half of the unit while 32
percent occur in the north half. This coincides with results of the Fall 1976
and 1977 ground, and 1977 helicopter deer classification counts, which show
much higher fawn:doe ratios on the south half of the unit.

��-281-

EXPERIMENTAL

DEER INVENTORY

SOUTHWEST

REGION

Roland C. Kufeld

P.N.O. OBJECTIVE
To design appropriate sampling and analytical procedures necessary to
reliably estimate deer numbers and buck:doe:fawn ratios on winter range
based on a preliminary sampling of a problem management unit within each
of the four administrative regions of the state.
SEGMENT OBJECTIVES

1.

To design and execute a helicopter count of deer on the entire
winter range of Game Management Unit 62.

2.

To design a sampling scheme based on the means, variances, and deer
density strata estimated from Objective 1 capable of estimating mean
deer numbers +15% at a ~ 0.05.

3.

To sample buck:doe:fawn ratios on 12 or more randomly selected,
all-day, walking routes on the winter range of Game Management Unit 62.
DESCRIPTION

OF THE STUDY AREA

This report deals with the portion of the statewide
that is being conducted in the southwest region.

deer inventory

study

The area selected for study is the winter range portion of Game Management
Unit 62, which encompasses 651.75 square miles (1688.0 km2) on the east
side of the Uncompahgre Plateau.
This area extends from Whitewater to
Ridgway.
It is bounded on the east by the Gunnison River between Whitewater
and Delta, and the western edge of the farmland that lies along the west side
of Highways 50 and 550 from Delta to Ridgway.
The western boundary is the
8000 foot (2438 m) contour level along the east side of the Uncompahgre Plateau.
The area is characterized by flat mesas separated by deep canyons which run
from southwest to northeast.
Vegetation is predominately pinyon-juniper
forest and sagebrush parks.
The higher elevations are covered by thick stands
of Gambel oak and the lower elevations approach a desert type.
METHODS AND MATERIALS
One hundred ninety-three 160-acre sample quadrats were selected at random
within 8 strata into which the area was divided.
The number of quadrats
in each strat.um was weighted according to the area within the strata and
available knowledge of relative deer density.

�-282-

The location of each quadrat was plotted on U.S.G.S. topographic maps and
aerial photos.
During the summer of 1976 each quadrat corner was located
and marked with a l' x 2' (.3 x .6 m) rectangular piece of florescent orange
painted masonite nailed in the tops of trees, or placed on a steel fencepost
if no tree was available at the corner.
These markers subsequently faded to
a pale yellow color within one year, especially those on the northeast and
northwest corners which faced south. More than half of these were replaced
during the summer of 1977, with a 12" x 24" x .100" (30.5 x 61 x .25 cm)
Accurene (ABS) Ter polymer marker, Federal orange in color. These were
purchased from P.A.R. Marketing Association, 635 Wetmore Drive, Wichita,
Kansas, 67209. They are extremely tough, do not crack or break. even when
shot and no fading has been noticed after nearly a year of use. Remaining
masonite markers will be replaced with Accurene markers during the summer of
1978.
When random selections of quadrat locations were made, some contiguous quadrats
were chosen.
It was found that deer were spooked from adjacent quadrats during
the count.
Contiguous quadrats were deleted during the summer of 1977, and
replaced with an equal number of non-contiguous ones. This was done through
random selection.
New quadrats were marked with accurene markers.
Deer classification counts were made between November 16, and December 4, 1977,
on 25 "walking type", circular routes located at random throughout the area.
Routes averaged 5 miles (8 km) in length.
Each route was situated so that it
ran through as many quadrats as possible in the 5 miles (8 km). The count was
made by 2 observers with I observer per route. One observer ran 12 routes and
the other ran 13 routes.
The number of routes ran in each stratum by each
observer was about equal.
Since each observer ran half of the routes their
data were combined to derive totals for all of Unit 62.
Aerial deer classification counts were made on December 9 and 10, 1977 on the
same 25 routes on which "walking type" counts had been made between November
16 and December 4. Each route encompassed about 3 square miles (7.8 km2).
All 25 routes were covered by 2 observers seated side by side in a Hiller Soloy
helicopter.
Each group of deer encountered was classified by both observers
working independently and without collaboration.
The aerial observers were
not the same as those who made the ground counts.
The count required 12.2
hours of flying time. Both days were warm and clear with no wind. There was
no snow cover.
During the period February 11 through 18, inclusive, deer were counted in the
193 quadrats.
The count required 32.5 hours of flying time in a Hiller Soloy
helicopter.
Last year the same count required 46 hours (Kufel~ 1977). After
the entire area had been covered on the 1978 flight, stratum 8 was flo'NO a
second time to determine the repeatability of the census system.
This stratum
contains 38 quadrats and the second flight took another 5.5 hours of flying
time. Five days elapsed between the first and second flights of stratum 8.

�RESULTS AND DISCUSSION
Deer Classification
Ground Classification

Counts

Counts

A total of 521 deer was classified

comprised

of 84 bucks,

from the ground in 1977 which was
299 does, 138 fawns.

The 1977 buck:doe ratio of 28.1 and fawn:doe ratio of 46.2 were not
significantly different, at the 10 percent significance level, from the
previous years ratios of 20.4 bucks and 51.3 fawns per 100 does (Table 1).

Table 1. Comparison of 1976 and 1977 buck:doe
from ground classification data.

and fawn:doe

ratios calculated

Ratio

Mean

1976
90% Conf. limits

Mean

1977
90% Conf. limits

Buck:doe

20.4

12.7 to 28.2

28.1

21. 3 to 34.9

Fawn: doe

51.3

43.4 to 59.2

46.2

38.2 to 54.1

Lower fawn:doe ratios were recorded in strata 1 thru 4 (the north half of
Unit 62) than in strata 5 thru 8 (the south half).
Fawn:doe ratios in strata
1-4 averaged 39 fawns/IOO does compared with 50 in strata 5-8. These differences were not significant at the 10 percent significance level, however (Table 2).
During the previous year (1976) 38 and 64 fawns/100 does were observed in the
south and north halves, respectively, and that difference was significant at
the 10 percent level (Kufeld 1977).

Table 2. Classification of deer observed on 25, 5-mile walking
Game Management Unit 62 during November and December, 1977.
Bucks/100 does
Mean 90% Conf. limits

routes in

Fawns/IOO does
Mean 90% Conf. limits

Stratal'!

Bucks

Does

Fawns

Total

1-4

35

103

40

178

33.9

21.1 to 46.9

38.8

29.7 to 47.9

5-8

49

196

98

340

25.0

17.5 to 32.5

50.0

38.3 to 61.7

Total of
Unit 62

84

299

138

521

28.1

21.3 to 34.9

46.2

38.2 to 51.1

1/
Strata 1 thru 4 represent
the south half.

the north half of the unit.

Strata 5 thru 8 represent

�There were no significant differences between observers at the 10 percent
significance level in fawn:doe or buck:doe ratios observed.
The mean
differences between observers and 90 percent confidence limits were 10.9
± 16.6 for fawn:doe ratios, and 9.0 ± 10.8 for buck:doe ratios.
There was a significant difference at the 10 percent significance level
in fawn:doe ratios, but not in buck:doe ratios between those observations
where all deer in a group were classified as compared to observations
where some deer in the group were not classified (Table 3).

Table 3. Comparison of buck:doe and fawn:doe ratios between completely
and partially classified groups of deer based upon data derived from ground
classifications.

Partly classified
Completely

groups

classified

groups

Bucks /100 Does
Mean 90% Conf. limits

Mean

Fawns/IOO Does
90% Conf. limits

28.6

21.0 to 36.1

35.1

23.7 to 46.6

27.9

19.0 to 36.8

51.0

42.7 to 59.3

According to 1976 and 1977 data the current sampling level of 25 routes is
sufficient to permit estimation of doe:fawn ratios within 20 percent of the
mean with 95 percent confidence (Table 4). Variation in buck:doe ratios
was much greater than fawn:doe ratios, and the number of routes required for
accurate estimation of buck:doe ratios is quite large (Table 3). Esti.mation
of buck:doe ratios within 20 percent of the mean with 95 percent confidence
would require 54 routes based on 1977 data, and 130 routes based on data
from 1976 (Kufeld, 1977).

Table 4. Number of deer classification ground routes needed to estimate
buck:doe and fawn:doe ratios within a given percent of the ratio at various
confidence levels.
Confidence

Level

Buck:Doe ratio
95%
90%

80%

95%

5%

862

592

352

434

298

177

10%

216

148

88

109

75

45

15%

96

66

39

49

34

20

20%

54

37

22

28

19

11

X% of ratio

Fawn:Doe ratio
90%
80~~

�-285-

Helicopter

Classification

Counts

Two aerial observers classified a total of 723 and 591 deer, respectively
(Table 5). There were no significant differences between observing at the
10 percent significance level in buck:doe or fawn:doe ratios observed.
The
numerical differences between ratios derived by the 2 observers were very
small. The differences between the means and 90 percent confidence limits
between observers were 1.0 + .02 for buck:doe ratios and 3.9 + 0.12 for fawn:
doe ratios.
Observed buck:doe ratios for all of Unit 62 were 22.5 for
Observer X and 23.5 for Observer Y. Fawn:doe ratios were 76.1 for Observer
X and 72.2 for Observer Y (Table 5). Since ratios determined by both
observers are very close, it is suggested that use of these data in population
formulas, etc. employ Observer XIS ratios as his were based on the larger
number of deer.

Table 5. Classification of deer by helicopter on 25, 3-square mile areas
in Game Management Unit 62 during December 9 and 10, 1977.~/

Area'!:} Bucks

Observer

Does

Fawns

3/
Bucks/lOa does90% Conf
limits
Total Mean

3/
Fawns/100 does90% Conf
limits
Mean

X

Strata

1-4

12

71

33

116

16.9

7.4 to 26.4

46.5

33.5 to 59.5

Y

Strata 5-8

70

293

244

607

15.6

6.5 to 24.8

48.4

36.4 to 60.5

X

Strata 5-8

10

64

31

105

23.9 18.7 to 29.1

83.3

75.7 to 90.8

Y

Strata 5-8

61

238

187

486

25.6 19.0 to 32.3

78.6

69.4 to 87.7

X

Total Unit
62

82

364

277

723

22.5 18.3 to 26.7

76.1

68.3 to 83.9

Total Unit
62

71

302

218

591

23.5 18.4 to 28.6

72.2

64.0 to 80.4

Y

~/ Counts were made on the same 25 routes on which ground counts were made
between November 16 and December 4.

1/ Strata

1 thru 4 represent the north half of the unit.
represents the south half.

3/

Strata 5 thru 8

Since there were no significant differences between observers, and ratios
determined by both observers were very close, it is suggested that use
of these data in population formulas, etc. employ observer XIS ratios as
his were based on the larger number of deer.

�-286-

Lower fawn:doe ratios were recorded in strata 1 thru 4 (the north half
of Unit 62) than in strata 5 thru 8 (the south half).
Observer X recorded
47 fawns/100 does in strata 1-4 and 83 in strata 5-8. This difference was
significant at the 10 percent significance level.
The number of 3-square mile (7.8 km2) helicopter deer classification areas
needed to estimate doe:fawn ratios within a given percent of the ratio at
various confidence levels is shown in Table 6. The number of areas needed
by each observer was relatively close although slightly more areas were
needed by Observer Y. For Observer X the current sampling level of 25
areas is sufficient to permit estimation of doe:fawn ratios within 10 percent
of the mean with 90 percent confidence.
Due to the greater variability in
estimating buck:doe ratios the current level of sampling is sufficient to
permit estimation of buck:doe ratios within 20 percent of the mean with 90
percent confidence.
2
Table 6. Number of 3-square mile (7.8 km ) helicopter deer classification
areas needed to estimate buck:doe and fawn:doe ratios within a given percent
of the ratio at various confidence levels.
Confidence
Number of Areas
for Buck:Doe Ratios
80%
95%
90%

Level
Number of Areas
for Fawn: Doe Ratios
80%
90%
95%

Observer

X% of Ratio

X

5%

511

351

209

153

107

63

10%

128

88

53

39

27

16

15%

57

39

24

17

12

7

20%

32

22

15

10

7

4

5%

683

469

279

188

129

77

10%

171

118

70

47

33

20

15%

76

53

31

21

15

9

20%

43

30

18

12

9

5

Y

Comparison

of Helicopter

and Ground Classification

Counts

There was a significant difference at the 10 percent significance level
between fawn:doe ratios derived by the helicopter and ground classification
techniques when data for all of GMU 62 were combined (Table 7). Fawn: doe

�-287-

ratios obtained by the aerial technique were much higher (Helicopter = 76
fa¥llS/100 does, Ground = 46 fawns/100 does).
There was no significant
difference at the 90 percent level between the 2 techniques in buck:doe
ratios.

Table 7. Comparison buck:doe and fawn:doe
and ground classification data.

Type of Count
Ground

Helicopter

ratios calculated

from helicopter

Observer

Mean

Bucks/100 Does
90% Conf. limits

Mean

Fawns/100 Does
90% Conf. limits

A

32.7

20.6 to 44.7

51.7

39.1 to 64.3

B

23.7

18.4 to 29.0

40.8

30.0 to 51.6

A &amp; B

28.1

21.3 to 34.9

46.2

38.2 to 54.1

X

22.5

18.3 to 26.7

76.1

68.3 to 83.9

y

23.5

18.4 to 28.6

72.2

64.0 to 80.4

Helicopter

Deer Census

The count went very well during 1978. There was good snow cover on all
but the lowest quadrats.
The snow was about 2 feet (.61 m) deep at the
8,000 foot (2438 m) level, so the observers believed very few, if any, deer
were above the census area. No deer were seen on the uppermost quadrats.
The several changes made in the system to alleviate problems encountered
during the previous year's flight (Kufeld, 1977) helped a great deal.
These were:
(1) replacement of painted masonite markers with accurene
markers;
(2) deletion of contiguous quadrats and replacement with noncontiguous quadrats;
(3) this year when a group of deer was encountered
the counters left their location on the grid, flew to the deer and counted
them before the group became scattered.
The location of that group within
the quadrat was then noted, and the counters returned to their grid location
and continued to grid the quadrat.
This helped to keep groups of deer
separate.
The total deer population in Unit 62 with 90 percent confidence limits is
estimated at 17,884 ± 4,042 animals (Table 8). Thus, the true population
(assuming the estimate is accurate) would fall somewhere between 13,842 and
21,926.
Analysis of the 1978 data show that the percent sample of 193 quadrats is
sufficient to estimate the population within 25 percent of the mean number
of deer per square mile (2.59 km2) with 90 percent confidence (Table 9).
The same analysis based on 1977 data showed 193 quadrats were enough to
estimate within 15 percent of the mean with 95 percent confidence (Kufeld,

1977).

�Table 8. Results of helicopter deer quadrat census of the winter range portion of Game Management Unit 62 conducted
between February 11 and 18, 1978.
Deer Eer Quadrat
Standard
Mean
Deviation

Deer per
Sq. Hile
(2.59 km2)

Deer POEu1ation in Strata
Total
90%
Deer
Confidence Limits

Stratum Area
Km2
Mi2

No. of Quadrats
N
n

1

51.50

133.39

206

12

16

1.333

4.618

5.332

275

2

110.00

284.90

440

35

104

2.971

6.555

11.884

1,307

3

101.25

262.24

405

23

20

0.870

2.380

3.480

352

4

122.25

316.63

489

38

300

7.895

9.395

31.580

3,861

5

58.00

150.22

232

13

50

3.846

7.381

15.384

892

6

79.75

206.55

319

25

374

14.960

28.167

59.840

4,772

Strata

Deer
Counted

I
N

CIJ
OJ

I

7

39.75

102.95

159

9

51

5.667

11.597

22.668

901

8

89.25

231.16

357

38

588

15.474

23.214

61.895

5,524

-651.75 1688.03

2607

193

1503

6.86011

0.939611

27.44JlI

11 These are stratified random values.
21 90 percent confidence limits indicate a population of between 13,842 and 21,926 deer.

17,884

± 4,0421:.1

�-289-

The higher number of quadrats needed on the basis of 1978 data is due
to greater variation in the 1978 data. Deer appeared more concentrated
during 1978, and, in a number of cases, higher deer counts were made
on individual quadrats than during 1977.
Table 9. Number of quadrats needed to estimate the popUlation within a
given percent of the mean deer per square mile at various confidence levels.
No of quadrats needed per
Confidence Level
95%
90%

X% of Mean Deer/ 2
Square Mi (2.59 km )
10%

489

424

15%

347

298

20%

269

230

25%

220

188

Total population estimates for stratum 8 for the 2 flights made to determine
repeatability of the census technique were 5,524 and 4,528 deer, respectively.
A paired t test showed no significant difference at the 10 percent significance
level between these 2 popUlation estimates.
Relative distribution of deer throughout the unit was similar to that found
last year (Kufeld, 1977). Based on 1978 data, 31 percent of the deer
popUlation estimated for all of Unit 62 occurs in stratum 8 (Table 8), which
is an 89 square mile (230.5 km2) area encompassing the lower part of the Log
Hill Mesa. Last year, data showed 48 percent of the deer population occurred
in stratum 8.
Also (based on 1978 data) 68 percent of the deer population in Unit 62
occur in Strata 5, 6, 7 and 8 (Table 8), which represents the south half of
the Unit, or roughly the area south of the 25 Mesa road. Only 32 percent
of the deer wintering in Unit 62 occur in Strata 1, 2, 3 and 4 which is, in
general, the area north of the 25 Mesa road. The differences in population
sizes between the south and north halves of the unit is significant at the
.01 significance level. Data from last year (Kufeld, 1977) showed 79 and 21
percent of the popUlation in the south and north halves of the unit,
respectively.
This coincides with the results of the Fall 1976 and 1977
ground and 1977 helicopter deer classification counts which show much higher
fawn:doe ratios on the south half of the area (Tables 2 and 5, and Kufeld, 1977).
LITERATURE

CITED

Kufeld, Roland C. 1977. Experimental deer inventory-southwest
region.
Colo. Div. Wildl. Game Res. Rept. July, 1977, Part 2:287-297.
Prepared by:

~CI

~~

Roland C. Kufeld
Wildlife Researcher

C

��July,

1978

-291JOB PROGF~SS

State of

COLORADO
----------------------------

Project

No.

W-38-R-33

Job Title
Period

Deer-Elk

23

Work Plan No.

REPORT

Job No.

Investigations
I

---------------------------------

An i_m_a_l
__a_n_d
__P_e_nS,_u~p~p_o_r_t
__F_a_c
__l·_I_i_t_i_e_s
__f_o_r
__D_e
__e_r_-_E_l_k
__R_e_s_e_a_r
__c_h

Covered:

_

April 1, 1977 - June 30, 1978

Personnel:
Paul H. Neil, Bruce Trindle, Kate Trindle, Sally Harpster, Linda
Foerster, Lynn Stevens, Dan Parkinson, Ted Rodriguez, John Torres, Barry VanSant,
Chris Solop, Chuck Perrin, and R. B. Gill.

ABSTRACT
A total of 40 mule deer fawns were obtained during the Segment.
Thirty-six
of the 40 were successfully hand-reared and trained for research projects in
Kremmling, Fort Collins, Piceance Basin and Poudre Canyon.
Rearing success
was attributed to formula, good weather and improved hand-rearing techniques.
In conjunction with fawn hand-rearing activities an experiment was conducted
to test the humoral and cell-mediated immunocompetence
of mule deer fawns obtained from wild does vs those obtained from tame does.
It was concluded that
wild does appeared to be better mothers than tame does if kept under a minimum
of stress which was revealed by the higher plasma gamma globulin levels in
fawns from wild does which appeared to be a result of the ingestion of more
colostrum.
Pen expansion and construction continued wit.h the completion of the
runway system and the 2 acre elk pen which is now occupied by 8' research elk.
Electricity was brought in to the area and a water system was designed.
Construction is expected to continue into the next segment.

��-293-

ANIMAL AND PEN SUPPORT
FACILITIES FOR DEER-ELK RESEARCH
Paul H. Neil

PROGRAM NARRATIVE
To provide
facilities

OUTLINE

and maintain populations of captive big game animals and pen
to support big game research programs.
SEGMENT OBJECTIVES

1.

To develop a big game research

2.

To coordinate rearing, training, maintenance, and research
captive, tame animals under one research support facility.

3.

To integrate animal and physical plant support facility manpower
monetary requirements under a single budget.
METHODS

and holding

facility.
activities

with

and

AND MATERIALS

Ten wild does were trapped in the Piceance winter range and transported to the
Fort Collins facility for the purpose of obtaining fawns for hand-rearing and
training.
A total of 40 fawns were obtained during the segment from captive,
tame does, wild does and orphans that were brought in by field men.
In conjunction with hand-rearing and training an experiment was designed in
an effort to evaluate the effects of stress on the immune system of hand-reared
fawns. As a result of previous work conducted during the last segment, it was
believed that the presence of a stress induced immunosuppression
or deficiency
was a major factor responsible for the high fawn mortality experienced in
previous years. Twenty fawns were selected for an evaluation of humoral and
cell-mediated i~~unity.
The formula used to hand-rear the 40 fawns consisted of 2 parts whole milk to
I part canned evaporated milk with an addition of 1/10 the total volume per
day of buttermilk.
This formula was obtained from Mr. Floyd Blunt of the Wyoming
Game and Fish Commission at Sybille, Wyoming.
In addition to hand-rearing fawns, facility expansion and construction continued.
Materials were ordered, fence posts were set and construction of the interconnecting runway system, isolation pens and shelters was continued.
RESULTS AND D1SCUSSION
Thirty~six of the 40 fawns were successfully hand-reared and trained for digestion
cage research and for food habits trials.
Rearing success was attributed to

�-294-

the milk formula, good weather and improved rearing management techniques.
Sixteen of the fawns were transported to the Kremmling facility in September
for incorporation into research programs being conducted at that location.
Details of the results of the humoral and cell-mediated immunity experiment
can be obtained from several papers:
Trindle, B. D., L. D. Lewis, and L. H.
Lauerman, 1978. Evaluation of stress and its effects on the immune system
of hand-reared mule deer fawns, J. Wildl. Dis. (In Press)., and Trindle, B.
D., L. D. Lewis, and L. H. Lauerman, 1978, Techniques for evaluating humoral
and cell-mediated immunity in mule deer fawns J. Wildl. Dis. (In Press).
During the immune system experiment a technique for anesthetizing the fawns
for the purpose of obtaining skin biopsys had to be developed.
A standard
Heibrink
in-circuit wick vaporizor gas anesthesia machine, equipped with a
one-liter bag, canine mask, and 2 oxygen cylinders was used with Methoxyflurane
gas. Induction and recovery were very smooth, the depth of anesthesia easily
controlled and muscle relaxation very good. Details and results of the 58
anesthetics that were performed can be obtained from; Trindle, Bruce D., L.
D. Lewis.
1978. Anesthesia of mule deer (Odocoileus hemionus) fawns. J.
Appl. Ethology (In Press).
Pen and facility expansion and construction continued slowly but consistently.
Unfavorable soil conditions hindered post-hole digging and placement, however
the runway system was completed as well as the elk pen. Eight elk used in
W-38-R Work Plan 17, Job 2 now occupy one 2 acre pen. Construction of five
more holding pens for deer, twelve isolation pens, 3 digestion cages and a
weigh room facility continued.
Electricity was brought to the area by Public
Service Company and a water system is planned for next segment.
A septic
system for the lab and office area was also installed.

(~/
Prepared

;~j V 1 .I~rl'7 ~()

by _"\,,-"'-...t:--.:Q~.--=C_( __V__-=f!-:-·j_1 ---,-:'! =-!..( .: __
Paul H. Neil
Wildlife Tech II

-,--_

�July,

-295-

JOB PROGRESS

State of

1978

REPORT

COLORADO
--------~~~~~----------

Project No.

Deer and Elk Investigations

W-38-R-33

1
---------------------------Job Title
Elk Population and Ecological Studies
~~~====~~~====~~~~------------------Work Plan No.

Period

Covered:

Personnel:

24

Job No.

April 1, 1977 - June 30, 1978

George D. Bear and Paul H. Neil

ABSTRACT
Field personnel listed population dynamics and range requirements as the
most prevalent problem
areas in the Colorado elk Management program.
Pertinent literature was reviewed and a research study plan was prepared.
A preliminary evaluation was made of proposed elk capturing and marking
techniques.
Two elk calves were captured using a helicopter, and one
was marked with a radio-telemetry collar. Four other elk were captured
with Clover traps, modified for capturing elk, and three were marked
with radio-telemetry collars.
These techniques will need some modification,
but will be useful in determining elk mortality and movement patterns.

��-297-

ELK POPULATION

AND ECOLOGICAL

STUDIES

George D. Bear

P. N. OBJECTIVE
The main objective this first year will be to determine the status of
knowledge in elk ecology and to list, by priority, the deficiencies of
information which are necessary for attaining and maintaining the elk
management goals specified in THE STRATEGY OF TODAY FOR WILDLIFE TOMORROW.
A long range eLle- research plan will be prepared following this review
process.
SEGMENT OBJECTIVES
1.

Outline

elk management

problems.

2.

Review literature.

3.

Prepare a Program Narrative.
METHODS AND RESULTS

Initial work consisted of determining what problems are prevalent in the
Colorado elk management program.
Questionnaires were sent to management
personnel concerned with elk management; and they listed the following
as being the most important aspects for future research:
source and degree
of miscellaneous mortality (other than hunting); a system for obtaining
more accurate population estimates; determine carrying capacity of elk
ranges and competition among wild ungulates for range resources; effect
of various human activities en elk distribution; and identification "range"
for individual herd units.
Pertinent literature was reviewed and a study plan prepared.
This study
plan was submitted as a Program Narrative for Federal Aid Project W-126-R-2,
Work Plan 3, Job 2 (Elk Population and Ecological Studies).
Since a major portion of the proposed study depends on capturing and marking
elk, some procedures and equipment were briefly evaluated to determine their
effectiveness.
The technique of using a helicopter to capture elk calves
and mark them with radio collars, such as done in Idaho, was tested in Rocky
Mountain National Park during June 1978. This was done by hovering over
young elk calves, thus frightening them. They would then try to hide by
lying down and remaining motionless while the helicopter landed nearby and
the observer caught the calf, and collared it. Of course this means the elk
must be in open terrain to permit use of the helicopter.
This technique
appears to be very efficient and permits capture of calves that would not be
located or caught by ground crews. More work must be done to determine the

�-298-

peak of calving season.
This preliminary test was too early and only
three calves were seen; two were easily captured, while the third was in
a heavily timbered area and the helicopter could not land. Only one
radio collar was available for this test. Movements were traced for
this calf as the herd migrated toward their summer range. The radio
transmitter was attached to the calf with an expandable collar which
would expand as the animal grew in size. However, the collar expanded
prematurely and the calf lost the collar four weeks after capture.
Therefore these collars must be modified so they will be more dependable.
It does appear that this technique of capturing and marking calves has
promise and can be useful in studies of calf elk mortality.
During February, 1978 an attempt was made to capture and mark elk using
the methods being considered for population research.
Bait stations
were established on the Pulliam Ranch, near Fort Collins, Colorado.
Livestock salt and alfalfa hay were used for bait.
Elk frequented the
stations the first day, and after one week there were 15-20 elk using one
station.
Two elk sized Clover traps were placed at the station; four elk
(3 cows and I calf) were captured during the first three nights of trapping,
then the elk abandoned the bait station.
The first three elk (2 cows and
1 calf) were successfully tranquilized, out-fitted with radio collars, and
released.
The fourth elk (adult cow) was severly injured with the tranquilizing equipment and died at the trap site. Based on this limited
experimentation using Pneu-dart and cap-chur equipment and consultations
other scientists it was concluded the pneu-dart syringe (with pre-measured
dosages of succinyl) and rifle would best suit our needs for tranquilizing
elk in the Clover traps. Also, it was found the traps should be placed
out-of-sight of each other, so as to minimize the stress on a trapped elk
while workers are processing an animal in another trap.
The radio transmitters and collars have worked very well for tracking
movements on the elk as they migrated to their summer range. We will need
a more sensitive receiver and antennae system if we are to obtain reliable
information on elk activity patterns, such as feeding, walking, bedding,
etc. It appears such systems are available for purchase.

d

Prepared

by

e::P .

(;l.,r&lt;'~

•.
~.
__ " _yz.
George D. Bear
Wildlife Researcher

_

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                  <text>JOB PROGRESS

State of

COLORADO

Project No.

W-101-R-20

Work Plan No.
Job Title

Game Range Investigations
Job Nc.

4

la
------------~--------------

Inventory of Range Manipulation Projects in Colorado
------~~~~-=~~~~~--~~~~~~--~----------------

Period Covered:
Personnel:

REPORT

April 1, 1977 - June 30, 1978

Roland C. Kufeld, and Regional

Game Biologists

ABSTRACT
An inventory has been made of all range vegetation modification projects
completed through 1976, in Colorado, west of Interstate Highway 25, on lands
administered by the u.S. Forest Service and Bureau of Land Management.
The
inventory also included projects completed on the Southern Ute and Ute
Mountain Indian Reservations through 1969. Acreages treated were: Forest
Service - 253,877, Bureau of Land Management -343,364, Indian Reservations68,169.

��-301-

INVENTORY OF RANGE MANIPULATION

PROJECTS

IN COLORADO

Roland C. Kufeld

P. N. OBJECTIVE
To collect information which describes proposed and completed range vegetation
modification projects that are located in Colorado, and their effects on the
range, wildlife and livestock; and to provide desired IBM listings of these
data to cooperating agencies upon their request.
INTRODUCTION
Game managers need to know the location, extent, kinds and effects of typeconversion projects which have already modified game ranges in their areas, so
they can anticipate the probable impact of similar projects in the future.
Also, they need to know in advance where, when and how the range is going to
be modified so they can determine the probable effects of proposed projects
on wildlife, and make recommendations for maintaining or improving wildlife
habitat.
The purpose of this study is:
(1) To determine the characteristics of each
of the range type - conversion projects that have been conducted in Colorado;
(2) How these projects have affected wildlife;
(3) To establish an interagency information exchange system designed to keep all land management agencies
mutually advised on proposed type-conversion projects and results of past
projects.

METHODS
An inventory has been made of all range vegetation modification projects
completed through 1976, in Colorado, west of Interstate Highway 25, on lands
administered by the U.S. Forest Service and Bureau
of Land Management.
The
inventory also included projects completed on the Southern Ute and Ute Mountains
Indian Reservations through 1969.
Descriptive information concerning projects completed through 1969, was assembled
from files in field offices of the U.S. Forest Service, Bureau of Land Management
and Bureau of Indian Affairs.
After 1969, data were collected through an interagency information exchange system under which the U.S. Forest Service and
Bureau of Land Management notified the respective Colorado Division of Wildlife
Regional Office one year in advance of their plans for range vegetation modification.
A joint field inspection was conducted which provided Colorado Division of
Wildlife the opportunity to make recommendations concerning the project and, in
some cases, to contribute financially to the improvement of wildlife habitat.
A
standard inventory form was also completed at that time (Fig. 1). This form is
the same as that used to inventory projects completed prior to 1969. Upon
completion of treatment, information was obtained from the land management agency

�-302-

concerning the dates the work has performed, the acres involved, and any
details concerning the project if different from that planned.
Provisions
of the information exchange system also called for post-treatment evaluations
to be made on each project 2, 5, and 10 years after treatment using a
standard form (Fig. 2).
Inventory data were placed on ADP cards and original forms placed on file
by project.
This was handled by the Colorado Division of Wildlife Big Game
Research Section.
Data listings concerning vegetation modification work were made
available to all cooperating agencies upon request.
RESULTS AND DISCUSSIONS
Acreages of rangeland treated through 1976, in Colorado, by the U.S. Forest
Service and Bureau of Land Management, and through 1969, by the Bureau of
Indian Affairs are shown in Tables 1, 2 and 3. Table 4 shows combined data
for the U.S. Forest Service and Bureau of Land Management.
Data are listed
by vegetation type and kind of treatment, and show the number of acres seeded
as part of treatment.
The Forest Service and Bureau of Land Management have
treated 253,877 and 343,364 acres respectively through 1976, for a total of
597,241 acres.
Through 1969, 68,169 acres were treated on the 2 Ute Indian
Reservations.
Most of the work has been done in sagebrush and Pinyon-juniper
types with browse type ranking third. Most browse type work involved control
of Gambel oakbrush.
The Forest Service and BLM treated 325,394 acres of
sagebrush, 131,422 acres of pinyon-juniper, and 29,072 acres of browse through
1976. Most work in sagebrush involved spraying, and most work in pinyon-juniper
involved chaining.
In browse type most work has involved spraying, chaining
and burning.
The bulk of spraying was done before 1966, in the browse type.
Since then, chaining became popular and more recently many controlled burns
have been completed.
Acreages treated by individual National Forest, BLM District and Indian
Reservation are shown according to vegetation type and kind of treatment
Tables 5 through 22~ Total acreages are summarized in Table 23.

in

A discussion of project characteristics, such as equipment used, chemical
herbicides, plant species used for reseeding and environmental characteristics
has been presented by Kufeld (1968a and 1970).
Effects of range vegetation modification projects on range, livestock and
wildlife as determined from field inspections of projects completed through
1965 have been discussed by Kufeld (1968a and 1968b). Some 2 and 5 year posttreatment evaluations were made on projects completed after 1969. Data from
these evaluations were accumulated on file for several years until enough 2
and 5 year post-treatment evaluations were made to provide a basis for a
meaningful data analysis.
Due to the workload and limited available manpower
within the Division of Wildlife Regions some scheduled evaluations were omitted
or delayed.
This caused a discontinuity of data which made it virtually
impossible to derive any useful information from the relatively limited posttreatment evaluation data that has already been collected.
For that reason

�-303v,'hcncompleting the blanks on this form follow the instructions in the
Interagency Council on Wildlife Ecology booklet entitled "A System for
Inventory and Evaluation of Vegetation Hodification Projects".

IMPORTAl~T :

GFP #4A COMPLETED
Colorado

VEGETATION

PROJECT DESIGNATION
1. Number _______________________________

II*

LAND STATUS
3. Natl. Forest
4. BLi.1District
5. State Agency
6. SCS Area
7. Indian Agency

III*

LOCATION
9. G. F. &amp;P. Region~

IV

PROJECT

Game, Fish and Parks Division

1*

12.

MODIFICATION

2.

Name~

District

Name

Allotment

or Unit

10. W.C.O. District

_

County
Tmvnship

_

_

11. Game Ngmt.
Unit No. ----

13. Drainage.
Range

_
Sections.

P.M.

_

Dominant
SITE DESCRIPTION
14. Ve~,=ta tive Type__________________________
15. Spe cies
16.:" ::::::.
Co-Ii
a t 1.011_______________________
1. I •
1. opo graPny
_
18. Percent Slope
19. Exposure
_ 20. Soil Type
----21. Erosion Condition
_
2lA. Hethod Used to Determine Erosion Condition'--_
22. Range Condition for Livestock
For Deer
For Elk
22A. Method Used to Determine Range Condo Livestock
Deer
E1k
_
23. Forage Production for Livestock
For Deer
For E1k~
_
23A. Hethod Used Deter. Forage Prod. Livestock
Deer
Elk
24. Vegetative Cover
25. Shrubs
26. Herbs ------------26A. Grass
27. Litter
28. Ground.
_
28A. Method Used to Determine Vegetative Cover &amp; Composition
'---------------29. Agencies
30. Date.
_
Remarks

*

--------------------------------------------------------------------------

V

VI

CLIMATIC DATA
31. Avg. Annual Precipitation
33. Avg. Annual Temperature
33B. Avg. July Temperature
35. Avg. Snow Depth on Area

Seasonal
32. Distribution
33A. Avg. Jan. Temperature
34. Length GroHing Season -----36.*Station Reporting

RANGE USE
(At Time of Treatment)
37. Kind of Livestock
39. Carrying Capacity
41. Stocking Rate

38. ,~ Season of Use
40. Grazing System
Remarks

j

*

-------------------------------_._----Fig. 1.

Form used for recording

descriptive

inventory

data.

�-30443.*
_

42.~ Kind of
Wildlifc,

------

44A. How Use Determined
Remarks

44. Amount
of Use ------

Sca s on
of Use~----------

----------- 45. Agency --------------

46. Date

---------

---------------------------------------------------------

VII

TREATMENT
47.-1&lt;No. Acres
Treated

48.* Purpose of
Treatment
50. Equipment Used
52.·k Equipment, Chemicals
54.* How Seeded
56.* Species Se-e-d~e-d--------------------------

-----------------------------~------~~----------------~------------------------------------

----------------

49.* Kind of Treatment
51.* Date Treated
53.* Seeded: Yes
No
55.* Date Seeded----

,--------

~-----------------------

57.* Rate Seeded
58.* Protection Given
Location from Other Projects
Remarks

--------------------------------------------------------------------------------------------------------------------MAP OF PROJECT LOCATION
R

R

1

2

3

6

1

2

3

4

5

1

6

5

4

7

8

9

10

11

12

7

8

9

10

11

12

~

18

17

16

15

14

13

18

17

16

15

14

13

~
~
~

22

23

24

T

.

',1

:

19

I

I

20

21

22

I

I

I

23

rj

24

19

20

21

I

29

28

27

26

25

30

29

28

27

26

25

31

32

33

34

35

36

31

32

33

34

35

36

6

5

4

3

2

1

6

5

4

3

2

1

7

8

9

10

11

12

7

8

9

10

11

12

18

17

16

15

14

13

18

17

16'

15

14

13

19

20

21

22

23

24

19

20

21

22

23

24

30

29

28

27

26

25

30

29

28

27

26

25

31

32

33

3!~

35

36

31

32

33

34

35

36

e Lf o r t

should

be ma d e

T

..};

;

30

"flnggcd"
are of p r i ma r y i mpo r t a nc e,
to o b t a in Lnf o rrna tion.

ItC'l1lS

Form Completed By
(:;hO\v t i t Lc and agency

and special

Title
Agency
Dnt c
of aLl persons pn r t i.c i.p.it Lng in r-omp lc tLon

_
of Lorru)

�-305IMPORTAl."lT:When completing the blanks on this form follow the instructions in
the Interagency Council on Hildlife Ecology booklet entitled "/\.
System for Inventory and Evaluation of Vegetation Modification Projects".

GFP 1f4B
RESULTS AND EFFECTS OF VEGETATION
Colorado

I.

*

II. *

III.

*

MODIFICATION

PROJECTS

ON HABITAT

Game, Fish and Parks Division

PROJECT DESIGNATION
1. Number

2.

Name

.

_

Allotment
District or Unit
Name
LAND STATUS
Natl.
Forest
3.
4. BU1 District
5. State Agency
6. SCS Area
7. Indian Agency
8. Private (Name and Address) ---------------------------------------11. Game Mgmt.
LOCATION
Unit No.
10.
W.C.O.
District
9. G.F.&amp;P Region ~---------____________________
13. Drainage-----12. County
Range
Sections ------- P •M.
Township

'"l'7

r.=PT7i7"-.·"

..Ly.

J...J.1. ••..,L...I,-",..Lr..J

•..
-;

.•...
.:

:'0,,17

••••••
..1....

••....•.
\..~.I.l.

,;~.1~7
••.._ .••
~.L

18.

19.
23.

24.
25.
27.

29.

_

.•
:r:"'f'1
•••..

After

14.
15.
16.
17.

_
_

Increase Decrease

Plant Suecies

Production for Livestock
Production for Deer
Production for Elk
Deer
Elk
Used to Determine Forage Prod. Livestock --Plant
S])ecies
Decrease
Increase
After
Vegetative Cover
_
_
22. Litter
21. Grass
Shrubs
20. Herbs ---------Bare Ground .--------~--------------------------------------------Method Used to Determine Vegetative Cover &amp; Composition.
_
Kill Percent
26. How Kill Determined ----------------Success Percent
28. How Success Determined
_
Agencies Making Determination.
30. Date __---------

Forage
Forage
Forage
Method

Remarks ---------------------------------------------32.

Season of Use

33.

Type Grazing

Syatem

31.

Kind of Livestock

34.
36.
39.

Present Carrying Capacity
35. Date
_
Present Stocking Rate ------37. Increase of
38. Decrease of --No Change
40. How Carrying Capacity and Stocking Rate Determined ----------------_._---------------------------------Continued
Fig. 2.

Form used for recording

on Reverse

Side

post-treatment

evaluation

data.

�-306·-

41.
43.

______________
42. Date
Agencies Making Determination
'---Kind of
44. Season
45. Amount
Wildlife
of Use
of Us e.
-----~~------Hare Use By
47. Less Use By

46.

TREATMENT
Animal
Species

_
_
_
_

EFFECTS

(48)

(ff9)

(50)

(51)

(52)

(53)

(54)

Benefieial

Detrimental

1'To
Effect

Unknown

How
Determined

Agencies

Date

.----_._----

---------------

-.&amp;.'-_,.~••..• .
v

,

·

1.,

.Io..

" •••• ..,;,

.

Form Completed

by

Title
(Date)

Agencies

_

(Show name, title and Agency of all persons participating

in completion

of form)

�-307-

Table 1. Acreages of range land treated through
U. S. Forest Service.

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow
Spray &amp; Harrow
Spray &amp; Plow
Pitting
Other

38
17
52
37
8
1
3
6
1

3714
2057
l3855
33084
1448
200
2374
530
1020

3714
2057
13855
1986
1448
200
2374
530
1020

Subtotal

163

58282

27184

Seed only
Harrow
Plow
Spray

2
2
1
2

76
71
15
287
-----

76
71
15
47
-------- -.----

Subtotal

7

449

209

Harrow
Plow
Spray
Plow &amp; Harrow

2
2
3
1

180
325
375
40

130
325
20
40
------

Subtotal

8

920

565

Seed only
Harrow
Plow
Spray
Chain
Rotobeat
Plow &amp; Harrow
Spray &amp; Harrow
Spray &amp; Plow
Pitting

24
11
40
187
1
1
6
1
2
3

2659
2059
11211
120161
300
128
1309
160
l35
960

2659
2059
11211
995
300
128
1309
160
l35
960

Subtotal

276

l39082

19916

Harrow
Spray

1
6

65
2910

65
0

Subtotal

7

2975

65

Seed only
Plow
Spray
Chain

4
1
19
23

210
523
l2l70
4839

195
523
985
3269

Kind of
Treatment

Grass

Perennial

Forbs

Sagebrush

Rabbitbrush

Browse

by the

No. of
Projects

Vegetative
Type

Meadow

1976, in Colorado,

------------------------------------------------------

.

-------------~---------

�-308-

Table l. Acreages of range land treated through 1976, in Colorado,
u. S. Forest Service. (Cont I d) .

..

Vegetative
Type
Browse

(Contld)

Conifer

Pinyon-Juniper

Aspen

Annual Weeds

Abandoned

Accidental

Lands

Burn

All Vegetative
Types

by the

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Burn
Spray &amp; Plow
Hand Cutting
Brush Cutter
Burn

18
1
3

8897
40
150

4560
40
0

1

712

0

Subtotal

70

27541

9572

Seed only
Plow
Burn

13
2
1

985
299
205

985
299
205

Subtotal

16

1489

1489

Seed only
Harrow
Chain
Burn
Brush Roller

1
1
12
1
1

150
460
12472
200
160

150
460
ll484
200
160

Subtotal

16

13442

12454

Seed only
Harrow

6
5

2892
836

2892
836

Subtotal

II

3728

3728

Seed only
Harrow
Plow
Spray
Spray &amp; Plow

9
5
1
2
1

ll96
276
40
1210
ll5

ll66
276
40
0
ll5

Subtotal

18

2837

1597

Seed only
Harrow
Plow
Plow &amp; Harrow

5
1
4
1

206
5
56
5

206
5
56
5

Subtotal

II

272

272

Seed only

5

2860

2860

Subtotal

5

2860

2860

608

253877

799ll

Grand Total

&amp;

�-309Table 2. Acreages of rangeland
Bureau of Land Management.

treated through 1976, in Colorado,

by the

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grass

Seed only
Harrow
Plow

5
1
6

458
50
2868

458
50
2868

Subtotal

12

3376

3376

Seed only
Plow
Spray

1
1
1

200
400
152

200
400
0

Subtotal

3

752

600

Seed only
Harrow
Plow
Spray
Chain
Burn
Rotobeat
Brush Cutter
Plow &amp; Harrow
Spray &amp; Plow
Plow &amp; Burn
Chain &amp; Burn
Other

40
6
116
78
27
5
5
3
5
5
6
1
1

14250
3102
44636
95409
17834
670
575
1728
2013
4365
1503
200
27

14250
3102
44466
1807
7980
670
100
0
2013
4365
1503
200
27

Subtotal

298

186312

80483

Seed only
Plow
Spray
Rotobeat

1
4
11
1

373
2317
11170
169

373
2317
2920
0

Subtotal

17

14029

5610

Seed only
Harrow
Plow
Spray
Chain
Burn

1
2
1
1
1
1

65
477
84
500
350
55

65
477
0
500
350
55

Subtotal

7

1531

1447

Seed only
Plow

2
1

2200
200

2200
200

Perennial

Forbs

Sagebrush

Rabbitbrush

Browse

Pinyon-Juniper

------------------------------------------------------------------------------

�-310Table 2. Acreages of rangeland treated through 1976, in Colorado,
(Cont' d) .
Bureau of Jtand Management.
No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Spray
Chain

1
122

1000
114580

0
82762

Subtotal

126

117980

85162

Seed only

1

40

40

Subtotal

1

40

40

Seed only
Plow
Other

1
1
3

200
200
525

200
200
525

Subtotal

5

925

925

Plow
Spray
Spray

2
1
1

167
934
380

167
0
380

Subtotal

4

1481

547

Seed only
Plow

5
1

2137
1275

2137
1275

Subtotal

6

3412

3412

Spray

5

1553

1150

Subtotal

5

1553

1150

Seed only
Plow
Plow &amp; Harrow

1
1
1

200
320
60

200
320
60

Subtotal

3

580

580

Seed only

4

832

832

Subtotal

4

832

832

Seed only
Harrow

25
1

10536
25

10536
25

Subtotal

26

10561

10561

Grand Total

517

343364

194725

Vegetative
Type

Kind of
Treatment

Pinyon-Juniper
(Cont'd)

Aspen

Saltbush

Greasewood

Desert

Shrub

Half Shrub

Annual Weeds

Abandoned

Accidental

Lands

Burn

All Vegetative
Types

by the

&amp; Chain

�-311-

Table 3. Acreages of rangeland
Bureau of Indian Affairs.

Acres
Treated

Acres Seeded as
Part of Treatment

Harrow
Plow
Spray
Rotobeat
Plow &amp; Burn

1
10
3
1
1

750
6866
6400
700
110

0
6866
1900
0
110

Subtotal

16

14826

8876

Chain

1

220

0

Subtotal

1

220

0

Seed only
Plow
Chain
Burn

3
2
5
2

11684
574
35235
5630

11684
574
21060
4600

Subtotal

12

53123

37918

Grand Total

29

68169

46794

Kind of
Treatment

Sagebrush

Pinyon-Juniper

All Vegetative
Types

Table 4. Acreages of rangeland treated through 1976, in Colorado,
u.S. Forest Service and Bureau of Land Management.

by the

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow
Spray &amp; Harrow
Spray &amp; Plow
Pitting
Other

43
18
58
37
8
1
3
6
1

4172
2107
16723
33084
1448
200
2374
530
1020

4172
2107
16723
1986
1448
200
2374
530
1020

Subtotal

175

61658

30560

2
2

76
71

76
71

Vegetative
Type

Kind of
Treatment

Grass

Meadow

by the

No. of
Projects

Vegetative
Type

Browse

treated through 1976, in Colorado,

Seed only
Harrow

------------------------------------------------------------------------------

�-312-

Table 4.

Acreages of rangeland treated through 1976, in Colorado,
Service and Bureau of Land Management.
(Cont'd).

by the

u.s. Forest

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Vegetative
Type

Kind of
Treatment

Meadow

Plow
Spray

1
2

15
287

15
47

Subtotal

7

449

209

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow

1
2

3
4
1

200
180
725
527
40

200
180
725
20
40

Subtotal

11

1672

1165

Seed only
Harrow
Plow
Spray
Chain
Burn
Rotobeat
Brush Cutter
Plow &amp; Harrow
Spray &amp; Harrow
Spray &amp; Plow
Plow &amp; Burn
Chain &amp; Burn
Pitting
Other

64
17
156
265
28

16909
5161
55677
2802
8280
670
228

1

16909
5161
55847
215570
18134
670
703
1728
3322
160
4500
1503
200
960
27

Subtotal

574

325394

100399

1
1

373
65
2317
2920

(Cont'd)

Perennial

Forbs

Sagebrush

Rabbitbrush

Browse

5

6
3
11
1

7
6
1

3

o
3322
160
4500
1503
200
960
27

Seed only
Harrow
Plow
Spray
Rotobeat

4
17
1

373
65
2317
l4080
169

Subtotal

24

17004

5675

Seed only
Harrow
Plow
Spray
Chain
Burn

5
2
2

275
477
607
12670
5189
8952

260
477
523
1485
3619
4615

20
24
19

o

------------------------------------------------------------------------------

�-3l3-

Table 4. Acreages of rangeland treated through 1976, in Colorado,
U.S. Forest Service and Bureau of Land Management.
(Cont'd).

by the

Vegetative
Type

Kind of
Treatment

Browse

Spray &amp; Plow
Hand Cutting
Brush Cutter &amp;
Burn

1
3

40
150

40

1

712

o

Subtotal

77

29072

11019

Seed only
Plow
Burn

13
1

985
299
205

985
299
205

Subtotal

16

1489

1489

Seed only
Harrow
Plow
Spray
Chain
Burn
Brushro11er

3
1
1
1
134
1
1

2350
460
200
1000
127052
200
160

2350
460
200
94246
200
160

Subtotal

142

131422

97616

Seed only
Harrow

7
5

2932
836

2932
836

Subtotal

12

3768

3768

Seed only
Plow
Other

1
1
3

200
200
525

200
200
525

Subtotal

5

925

925

Plow
Spray
Spray

2
1

167
934
380

167

1

Subtotal

4

1481

547

Seed only
Plow

5
1

2137
1275

2137
1275

Subtotal

6

3412

3412

Spray

5

1553

1150

Subtotal

5

1553

1150

(Cont'd)

Conifer

Pinyon-Juniper

Aspen

Saltbush

Greasewood

Desert Shrub

Half Shrub

&amp; Chain

No. of
Projects

2

Acres
Treated

Acres Seeded as
Part of Treatment

o

o

o
380

�-314-

Table 4. Acreages of rangeland treated through 1976, in Colorado,
U.S. Forest Service and Bureau of Land Management.
(Cont'd).
No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow
Spray &amp; Plow

10
5
2
2
1
1

1396
276
360
1210
60
115

1366
276
360
0
60
115

Subtotal

21

3417

2177

Seed only
Harrow
Plow
Plow &amp; Harrow

9
1
4
1

1038
5
56
5

1038
5
56
5

Subtotal

15

1104

1104

Seed only
Harrow

30
1

13396
25

13396
25

Subtotal

31

13421

13421

1125

597241

274636

Vegetative
Type

Kind of
Treatment

Annual Weeds

Abandoned

Accidental

Lands

Burn

All Vegetative
Types

by the

Grand Total

�-315-Table 5.
Forest.

Acreages

of rangeland

treated through 1976 on the Arapahoe

Acres
Treated

National

Acres Seeded as
Part of Treatment

Vegetation
Type

Kind of
Treatment

No. of
Projects

Grass

Seed only

9

362

362

Subtotal

9

362

362

Harrow

1

11

11

Subtotal

1

11

11

Seed only
Plow
Spray

2
3
6

123
121
3278

123
121
0

Subtotal

11

3522

244

Spray

1

100

0

Subtotal

1

100

0

Seed only

2

20

20

Subtotal

2

20

20

Seed only

2

295

295

Subtotal

2

295

295

Grand Total

26

4310

932

Meadow

Sagebrush

Browse

Abandoned

Accidental

Lands

Burn

All Vegetative
Types

Table 6. Acreages
National Forest.

of rangeland

treated through 1976, on the Grand Mesa

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grass

Seed only
Harrow
Plow
Spray

4
2
1
14

307
70
285
14474

307
70
285
1054

Subtotal

21

15136

1716

-------------------------------------------------------------------------------

�-316-

Table 6. Acreages of rangeland
(Cont'd).
National Forest.

treated through 1976, on the Grand Mesa

Acres
Treated

Acres Seeded as
Part of Treatment

Vegetative
Type

Kind of
Treatment

No. of
Projects

Meadow

Seed only
Harrow
Spray

1
1
1

12
60
47

12
60
47

Subtotal

3

119

119

Spray

2

355

0

Subtotal

2

355

0

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow

4
1
1
10
1

563
505
600
8475
681

563
505
600
200
681

Subtotal

17

10824

2549

Spray
Chain
Burn

3
1
1

190
60
60

60
60
60

Subtotal

5

310

180

Seed only

1

54

54

Subtotal

1

54

54

Chain

2

626

626

Subtotal

2

626

626

Seed only

2

226

226

Subtotal

2

226

226

Grand Total

53

27650

5470

Perennial

Forbs

Sagebrush

Browse

Conifer

Pinyon-Juniper

Aspen

All Vegetative
Types

�-317-

Table 7. Acreages
National Forest.

of rangeland

Vegetative
Type

Kind of
Treatment

Grass

Meadow

Perennial

Forbs

Sagebrush

Browse

Aspen

Annual Weeds

All Vegetative
Types

treated through 1976, on the Gunnison

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow

1
2
6
2
2

100
450
3590
2922
350

100
450
3590
0
350

Subtotal

13

7412

4490

Plow

1

15

15

Subtotal

1

15

15

Harrow
Plow
Spray
Plow &amp; Harrow

2
2
1
1

180
325
20
40

180
325
20
40

Subtotal

6

565

565

Plow
Spray
Plow &amp; Harrow
Spray &amp; Plow

4
38
2
1

1257
33392
207
5

1257
0
207
5

Subtotal

45

34861

1469

Spray
Spray &amp; Plow

3
1

2208
40

925
40

Subtotal

4

2248

965

Harrow

1

150

150

Subtotal

1

150

150

Seed only
Harrow

1
1

900
100

900
100

Subtotal

2

1000

1000

Grand Total

72

46251

8654

�-318-

Table 8.
Forest.

Acreages

of rangeland

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Plow
Spray
Plow &amp; Harrow

17
5
1

5019
3362
3

5019
0
3

Subtotal

23

8384

5022

Seed only
Plow

1
4

46
56

46
56

Subtotal

5

102

102

Seed only

1

2400

2400

Subtotal

1

2400

2400

Grand Total

29

10886

7524

Vegetative
Type

Kind of
Treatment

Grass

Abandoned

Accidental

Lands

Burn

All Vegetative
Types

Table 9. Acreages
National Forest.

of rangeland

Vegetative
Type

Kind of
Treatment

Grass

Meadow

Sagebrush

treated through 1976, on the Pike National

treated

through 1976, on the Rio Grande

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow
Spray &amp; Plow
Other

5
3
7
3
1
1
1

142
346
681
8229
15
1913
1020

142
346
681
75
15
1913
1020

Subtotal

21

12346

4192

Spray

1

240

0

Subtotal

1

240

0

Seed only
Harrow
Plow
Spray

1
3
1
3

23
410
73
522

23
410
73
40

-------------------------------------------------------------------------------

�-319-

Table 9. Acreages of rangeland
(Cont'd).
National Forest.

treated through 1976, on the Rio Grande

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Vegetative
Type

Kind of
Treatment

Sagebrush
(Cont'd)

Plow &amp; Harrow
Spray &amp; Plow

1
1

200
130

200
130

Subtotal

10

1358

876

Harrow
Spray

1
3

65
860

65
0

Subtotal

4

925

65

Seed only
Plow
Burn

1
1
1

20
523
48

20
523
0

Subtotal

3

591

543

Seed only

2

30

30

Subtotal

2

30

30

Grand Total

41

15490

5706

Rabbitbrush

Browse

Conifer

All Vegetative
Types

Table 10. Acreages
National Forest.

of rangeland

Vegetative
Type

Kind of
Treatment

No. of
Projects

Grass

Seed only
Plow

1
3

10
220

10
220

Subtotal

4

230

230

Plow
Spray

4
8

1490
4193

1490
0

Subtotal

12

5683

1490

Seed only

1

4

4

Subtotal

1

4

4

Sagebrush

Annual Weeds

treated through 1976, on the Roosevelt

Acres
Treated

Acres Seeded as
Part of Treatment

-------------------------------------------------------------------------------

�-320-

Table 10. Acreages of rangeland
National Forest.
(Cont'd).
Vegetative
Type
Abandoned

Kind of
Treatment
Lands

All Vegetative
Types

Table 11.
Forest.

Acreages

treated through 1976, on the Roosevelt

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Plow &amp; Harrow

1

5

5

Subtotal

1

5

5

Grand Total

18

5922

1729

of rangeland

treated through 1976, on the Routt National

Vegetative
Type

Kind of
Treatment

Grass

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow

4
3
2
2
1

518
105
165
201
60

518
105
165
196
60

Subtotal

12

1049

1044

Seed only
Harrow
Plow
Spray
Spray &amp; Harrow

8
2
3
44
1

124
88
225
16559
160

124
88
225
0
160

Subtotal

58

17156

597

Spray
Hand Cutting

1
2

67
120

0
0

Subtotal

3

187

0

Seed only

1

205

205

Subtotal

1

205

205

Seed only

1

50

50

Subtotal

1

50

50

Sagebrush

Browse

Conifer

Aspen

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

�-321-

Table 11. Acreages
Forest.
(Cont'd).

of rangeland

Vegetative
Type

Kind of
Treatment

No. of
Projects

Annual Weeds

Seed only
Harrow
Plow

5
2
1

184
95
40

154
95
40

Subtotal

8

319

289

Seed only
Harrow

1
1

40
5

40
5

Subtotal

2

45

45

Grand Total

85

19011

2230

Abandoned

Lands

All Vegetative
Types

Table 12. Acreages
National Forest.

of rangeland

Vegetative
Type

Kind of
Treatment

Grass

Sagebrush

Rabbitbrush

Browse

Pinyon-Juniper

treated through 1976, on the Routt National

Acres
Treated

Acres Seeded as
Part of Treatment

treated through 1976, on the San Isabel

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Harrow
Plow
Plow &amp; Harrow

3
1
3
1

1219
590
2138
750

1219
590
2138
750

Subtotal

8

4697

4697

Harrow
Spray

1
2

220
565

220
0

Subtotal

3

785

220

Spray

1

1000

0

Subtotal

1

1000

0

Seed only

1

150

150

Subtotal

1

150

150

Harrow

1

460

460

Subtotal

1

460

460

-------------------------------------------------------------------------------

�-322-

Table 12. Acreages of rangeland
National Forest.
(Cont'd).

treated through 1976, on the San Isabel

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Vegetative
Type

Kind of
Treatment

Annual Weeds

Harrow

1

6

6

Subtotal

1

6

6

Grand Total

15

7098

5533

All Vegetative
Types

Table 13. Acreages
National Forest.

of rangeland

Vegetative
Type

Kind of
Treatment

Grass

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow
Pitting

6
4
4
5
1
6

509
376
154
1801
50 .
530

509
376
154
500
50
530

Subtotal

26

3420

2119

Seed only
Harrow
Plow
Spray
Chain
Plow &amp; Harrow
Pitting

1
2
17
6
1
1
3

300
216
5028
6678
300
100
960

300
216
5028
0
300
100
960

Subtotal

31

13582

6904

Seed only
Chain
Burn
Hand Cutting
Brush Cutter
Burn

1
14
15
1

25
3430
8289
30

25
2110
4000
0

1

712

0

32

12486

6135

Sagebrush

Browse

Subtotal

treated through 197Q, on the San Juan

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

&amp;

-------------------------------------------------------------------------------

�-323-

Table 13. Acreages of rangeland
(Cont'd).
National Forest.

treated through 1976, on the San Juan

Acres
Treated

Acres Seeded as
Part of Treatment

Vegetative
Type

Kind of
Treatment

No. of
Projects

Conifer

Seed only
Plow

10
2

901
299

901
299

Subtotal

12

1200

1200

Seed only
Chain
Burn

1
1
1

150
1499
200

150
1499
200

Subtotal

3

1849

1849

Seed only
Harrow

1
2

316
326

316
326

Subtotal

3

642

642

Seed only
Spray
Spray &amp; Plow

2
1
1

108
910
115

108
0
ll5

Subtotal

4

ll33

223

Seed only

1

100

100

Subtotal

1

100

100

ll2

34412

19172

Pinyon-Juniper

Aspen

Annual Weeds

Abandoned

Land

All Vegetative
Types

Grand Total

Table 14. Acreages
National Forest.

of rangeland

Vegetative
Type

Kind of
Treatment

No. of
Projects

Grass

Seed only
Plow
Spray

1
4
1

150
658
419

150
658
0

Subtotal

6

1227

808

Seed only

1

64

64

Subtotal

1

64

64

Meadow

treated through 1976, on the Uncompahgre

Acres
Treated

Acres Seeded as
Part of Treatment

-------------------------------------------------------------------------------

�-324-

Table 14. Acreages of rangeland
(Cont'd).
National Forest.

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow

6
2
7
39
1

l386
620
2417
30569
121

l386
620
2417
455
121

Subtotal

55

35113

4999

Spray

2

1050

0

Subtotal

2

'1050

0

Seed only
Spray
Burn

1
9
1

15
8265
500

0
0
500

Subtotal

11

8780

500

Chain
Brush Roller

9
1

10347
160

9359
160

Subtotal

10

10507

9519

Seed only
Harrow

1
1

2100
60

2100
60

Subtotal

2

2160

2160

Spray

1

300

Subtotal

1

300

0

Grand Total

88

59201

18050

Kind of
Treatment

Sagebrush

Browse

Pinyon-Juniper

Aspen

Annual Weeds

All Vegetative
Types

through 1976, on the Uncompahgre

No. of
Projects

Vegetative
Type

Rabbitbrush

treated

0
,

�-325-

Table 15. Acreages
National Forest.

of rangeland

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Harrow
Plow
Spray
Plow &amp; Harrow
Spray &amp; Harrow
Spray &amp; Plow

4
2
5
5
1
1
2

397
120
945
1676
220
200
461

397
120
945
161
220
200
461

Subtotal

20

4019

2504

Seed only
Spray
Rotobeat

2
31
1

140
15930
128

140
300
128

Subtotal

34

16198

568

Spray
Chain

2
8

1340
1349

0
1099

Subtotal

10

2689

1099

Seed only
Harrow

1
1

200
300

200
300
----

Subtotal

2

500

500

Harrow

1

75

--- 75

Subtotal

1

75

75

Harrow

2

165

165

Subtotal

2

165

165

Grand Total

69

23646

4911

Vegetative
Type

Kind of
Treatment

Grass

Sagebrush

Browse

Aspen

Annual Weeds

Accidental

Burn

All Vegetative
Types

treated through 1976, on the White River

�-326-

Table 16.
District.

Acreages of rangeland treated through 1976, on the Craig BLM

Vetetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Sagebrush

Seed only
Harrow
Plow
Spray
Chain
Burn
Rotobeat

21
2
19
21
9
5
3

7932
217
15421
19954
4620
670
420

7932
217
15421
0
1992
670
100

Subtotal

80

49234

26332

Harrow
Burn

2
1

477
55

477
55

Subtotal

3

532

532

Seed only
Chain

1
19

1200
25507

1200
11519

Subtotal

20

26707

12719

Seed only

1

200

200

Subtotal

1

200

200

Spray

1

15

0

Subtotal

1

15

0

Seed only

3

792

792

Subtotal

3

792

792

Seed only
Harrow

9
1

3995
25

3995
25

Subtotal

10

4020

4020

Grand Total

118

81500

44595

Browse

Pinyon-Juniper

Saltbush

Half Shrub

Abandoned Lands

Accidental Burn

All Vegetative
Types

�-327Table 17. Acreages of rangeland treated through 1976, on the Glenwood
Springs B.L.M. District.
No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Seed only
Plow
Spray
Chain
Rotobeat
Plow &amp; Harrow
Spray &amp; Plow
Plow &amp; Burn
Chain &amp; Burn
Other

4
32
20
5
1
4
4
5
1
1

397
5317
23379
1708
75
663
2765
1330
200
27

397
5147
407
168
0
663
2765
1330
200
27

Subtotal

77

35861

11104

Spray

1

500

500

Subtotal

1

500

500

Chain

1

910

910

Subtotal

1

910

910

Seed only

1

40

40

Subtotal

1

40

40

Plow
Spray

2
1

167
934

167
0

Subtotal

3

1101

167

Spray

1

138

0

Subtotal

1

138

0

Seed only

1

40

40

Subtotal

1

40

40

Seed only

3

2472

2472

Subtotal

3

2472

2472

Grand Total

88

41062

15233

Vegetative
Type

Kind of
Treatment

Sagebrush

Browse

Pinyon-Juniper

Aspen

Greasewood

Half Shrub

Abandoned Lands

Accidental Burn

All Vegetative
Types

�-328-

Table 18. Acreages of rangeland treated through 1976,
B.L.M. District.

on the Montrose

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grass

Seed only
Plow

1
3

100
988

100
988

Subtotal

4

1088

1088

Seed only
Plow
Spray

1
1
1

200
400
152

200
400
0

Subtotal

3

752

600

Seed only
Harrow
Plow
Spray
Chain
Plow &amp; Harrow
Spray &amp; Plow
Plow &amp; Burn

4
1
55
27
4
1
1
1

864
1665
20096
45841
4891
1350
1600
173

864
1665
20096
400
1000
1350
1600
173

Subtotal

94

76480

27148

Seed only
Plow
Chain

1
1
1

65
84
350

65
0
350

Subtotal

3

499

415

Plow
Chain

1
59

200
55414

200
44552

Subtotal

60

55614

44752

Plow

1

200

200

Subtotal

1

200

200

Spray &amp; Chain

1

380

380

Subtotal

1

380

380

Plow

1

1275

1275

Subtotal

1

1275

1275

Spray

1

1150

1150

Subtotal

1

1150

1150

Perennial Forbs

Sagebrush

Browse

Pinyon-Juniper

Saltbush

Greasewood

Desert Shrub

Half Shrub

-------------------------------------------------------------------------------

�-329-

Table 18. Acreages of rangeland
B.L.M. District.
(Cont'd).

treated through 1976, on the Montrose

Vegetative
Type

Kind of
Treatment

Annual Weeds

Seed only
Plow
Plow &amp; Harrow

1
1
1

200
320
60

200
320
60

Subtotal

3

580

580

Plow

3

1226

1226

Subtotal

3

1226

1226

174

l39244

78814

Accidental

Burn

All Vegetative
Types

Grand Total

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Table 19. Acreages
B.L.M. District.

of rangeland

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Grass

Seed only
Harrow
Plow

3
1
3

219
50
1880

219
50
1880

Subtotal

7

2149

2149

Plow
Spray
Chain

2
4
1

2053
4000
l300

2053
1000
l300

Subtotal

7

7353

4353

Seed only
Plow
Spray
Rotobeat

1
4
11
1

373
2317
11170
169

373
2317
2920
0

Subtotal

17

14029

5610

Chain

23

11098

9851

Subtotal

23

11098

9851

Sagebrush

Rabbitbrush

Pinyon-Juniper

treated through 1976, on the Canyon City

�-330-

Table 19. Acreages of rangeland
B.L.M. District.
(Cont'd).

treated through 1976, on the Canyon City

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Vegetative
Type

Kind of
Treatment

Half Shrub

Spray

2

250

0

Subtotal

2

250

0

Seed only

2

150

150

Subtotal

2

150

150

Grand Total

58

35029

22113

Accidental

Burn

All Vegetative
Types

Table 20. Acreages
B.L.M. District.

of rangeland

Vegetative
Type

Kind of
Treatment

No. of
Projects

Grass

Seed only

1

139

139

Subtotal

1

139

139

Seed only
Harrow
Plow
Spray
Chain
Rotobeat
Brush Cutter

11
3
8
6
8
1
3

5057
1220
1749
2235
5315
80
1728

5057
1220
1749
0
3520
0
0

Subtotal

40

17384

11546

Seed only
Spray
Chain

1
1
20

1000
1000
21651

1000
0
15930

Subtotal

22

23651

16930

Other

3

525

525

Subtotal

3

525

525

Sagebrush

Pinyon-Juniper

Saltbush

treated through 1976, on the Grand Junction

Acres
Treated

Acres Seeded as
Part of Treatment

-------------------------------------------------------------------------------

�-331-

Table 20. Acreages of rangeland
B.L.M. District.
(Cont'd).

treated through 1976, on the Grand Junction

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Desert Shrub

Seed only

5

2137

2137

Subtotal

5

2137

2137

Seed only

8

2693

2693

Subtotal

8

2693

2693

Grand Total

79

46529

33970

Accidental

Burn

All Vegetative
Types

Table 21. Acreages
Indian Reservation.

of rangeland

Vegetative
Type

Kind of
Treatment

Sagebrush

Pinyon-Juniper

All Vegetative
Types

treated through 1969, on the Southern Ute

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Harrow
Plow
Spray
Plow &amp; Burn

1
9
2
1

750
4366
4500
110

0
4366
0
110

Subtotal

13

9726

4476

Seed only
Plow
Chain
Burn

3
2
4
2

11684
574
15675
5630

11684
574
1500
4600

Subtotal

II

33563

18358

Grand Total

24

43289

22834

�-332-

treated through 1969, on the Ute Mountain

Table 22. Acreages
Indian Reservation.

of rangeland

Vegetative
Type

Kind of
Treatment

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Sagebrush

Plow
Spray
Rotobeat

1
1
1

2500
1900
700

2500
1900
0

Subtotal

3

5100

4400

Chain

1

220

0

Subtotal

1

220

0

Chain

2

42546

42546

Subtotal

2

42546

42546

All Vegetative
Types

Grand Total

6

47866

46946

Table 23. Acreages
and B.L.M. District

of rangeland
in Colorado.

Browse

Pinyon-Juniper

National

Forests

B.L.M. Districts

treated through 1969, on each National

Forest

Land Mgt.
Agency

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Uncompahgre
Gunnison
San Juan
Grand Mesa
White River
Routt
Rio Grande
Pike
San Isabel
Roosevelt
Arapahoe

88
72
112
53
69
85
41
29
15
18
26

59201
46251
34412
27650
23646
19011
15490
10886
7098
5922
4310

18050
8654
19172
5470
4911
2230
5706
7524
5533
1729
932

Subtotal

608

253877

79911

Montrose
Craig
Grand Jct.

174
118
79

l39244
81500
46529

78814
44595
33970

-------------------------------------------------------------------------------

�-333-

Table 23. Acreages
and B.L.M. District

of rangeland
in Colorado.

treated through 1969, on each National
(Cont'd).

Land Mgt.
Agency
B.L.M. Districts
(Cont'd)

Forest

No. of
Projects

Acres
Treated

Acres Seeded as
Part of Treatment

Glenwood Sprgs.
Canyon City

88
58

41062
35029

15233
22113

Subtotal

517

343364

194725

Grand Total

1125

597241

274636

�-334-

the post-treatment evaluation phase was discontinued.
The descriptive
inventory of projects as they are proposed and completed is also being
discontinued due to limited availability of manpowe r , This entire job
was initiated as a "bookeeping" service to management by research.
All of
the data summarized in this report were gathered and/or submitted by management personnel.
Research is discontinuing its participation because
this appears not to be a priority management need.

LITERATURE

CITED

Kufeld, Roland C. 1968a. Inventory of range manipulation projects in
Colorado.
Colo. Div. Game, Fish and Parks. Game Res. Rept, July,
Pt. 1. 121p.
Kufeld, Roland C. 1968b. Range type-conversion
their impact on deer, elk and sage grouse.
Game and Fish Comm. Proc. 48:173-187.

programs in Colorado and
Western Assn. State

Kufeld, Roland C. 1970. Inventory of range manipulation projects in
Colorado.
Colo. Div. Game, Fish and Parks.
Game Res. Rept., July,
Pt. 1. 59-94.

�-335July,

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

W-I0I-R-20

Plan No.

4

Work

Covered:

Personnel:

Game Range

J~ ~.

Investigations

3

-------------------------------

Experimental Improvement of Oakbrush
on Deer. Elk and Cattle - Hightower Mountain

Job Title
.Period

1978

April

Roland

1, 1977 - June 30, 1978

C. Kufeld

ABSTRACT
Effects of controlled burning, spraying with 2,4,5-TP and anchor chaining
of Gambel oak vegetation on 92 kinds of forbs, 19 grasses and 16 shrubs,
and on elk, deer and cattle use were measured and documented 2 and 5 years
after treatment on a site 20 miles east of Collbran, Colorado.
With adjustments for normal vegetation production changes measured on control
units data indicate that, after 2 years, summer vegetation production increased
50 percent on sprayed areas and 20 percent on chainings, and decreased 17
percent on burns.
Most of the increase 2 years after spraying occurred among
grasses which increased 295 percent.
Most of the 2nd year increase on chainings and decrease onburns occurred among shrubs.
Production of forbs, grasses
and shrubs declined on all treated areas between 2 and 5 years after treatment.
Except for a minor increase among forbs on burns, and no change due to treatment
among forbs on chainings, vegetation production for all plant categories on
all 3 treatments was below pre-treatment levels after 5 years (Spray = -14%,
chain = -19%, burn = -26%).
This decrease was attributed to drought conditions
which prevailed between 2 and 5 years after treatment plus a severe frost
which occurred during late spring of the 5th year.
Most of the decrease after
5 years occurred among grasses on sprayed areas, and shrubs and grasses on
chainings and burns.
Data were grouped according to elk, deer and cattle forage.
An explanation of
these terms is presented.
Percentage changes in forage production after 2 and
5 years due to burning, spraying and chaining are discussed.
Burned and chained areas when compared with control and spray areas showed
increased levels of total cell contents,
soluble carbohydrate, copper and
zinc in most plant categories (forbs, grasses and shrubs) after 2 years.

�-336-

Elk showed distinct preference for burned areas after 2 and 5 years with
sprayed areas ranking second and chainings third.
Elk use on burns increased
45 and 54 percent, respectively, after 2 and 5 years.
Deer did not appear
to exhibit preference for any particular treatment after 2, 3 or 5 years.
Cattle showed 6 times more preference for burns.
After 2 years than for
sprayed or chained areas.
No preference was shown by cattle for sprayed or
chained areas after 2 years, or for any of the 3 treatments after 6 years.
Preference of elk and cattle for burns occurred despite decrease in forage
production on burns as compared with higher levels produced on sprayed areas
and chainings.
Attractiveness
of burns to elk and cattle may have been related
to an observed increase in several nutrients and minerals or to changes in
cover.
The overall effect of burning was creation of openings interspersed
with patches of standing oak vegetation suitable for escape cover and shade.
All preference data are based upon fecal group densities and may not accurately
reflect actual time spent on any treatment.
The study was, however, designed
to attempt to minimize possible biases encountered in use of fecal counts to
measure habitat preference.

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EXPERIMENTAL IMPROVEMENT OF OAKBRUSH
ON DEER, ELK AND CATTLE - HIGHTOWER MOUNTAIN
Roland C. Kufeld

P. N. OBJECTIVE
To determine the extent to which deer, elk and cattle forage production and
game use can be increased and maintained by chaining, spraying and controlled
burning on over-age Gambel oak winter game ranges.
SEGMENT OBJECTIVE
To determine the extent of vegetation composition, production and nutrient
content changes, and the extent of deer and elk use changes which have resulted
from burning, spraying and chaining.
METHODS AND MATERIALS
Study area design and sampling scheme have been described by Kufeld (1970 and
1971). Implementation of burning, spraying and chaining treatments on the study
areas as well as procedures and species used in reseeding have been discussed
by Kufeld (1972).
Vegetation Measurements 2 and 5 Years
After Burning, Spraying and Chaining
Two and 5 year post-treatment vegetation measurements were made in 600 permanent meter square plots on the Hightower Mountain oak study area during the
summers of 1973 and 1976, using the same procedures employed during the 1970
pre-treatment evaluations as described by Kufeld (1971). The pre-treatment
measurement period was July 10, through August 13, 1970. The 2 year posttreatment measurement period was July 9, through August 16, 1973. The 5 year
post-treatment measurement period was July 6, through August 6, 1976. Prior
to beginning vegetation measurements observers were trained in use of the
vegetation measurement techniques, and in identification of all plants on the
study area.
Pre-and post-treatment vegetation production data were computerized and subjected
to covariance analysis by the Colorado State University Statistics Department.
Statistical significance of vegetation production changes was tested at the .05,
.10, .15, .20 and .25 significance levels.
It was necessary to test for
significance at lower levels because the study involved only 2 replications for
each treatment.
Inclusion of more replications would have increased the study
area size, workload and manpower requirements to the point the study would not
have been feasible.
Determination

of Nutrient

Quality Change~

Samples of plant species designated by Kufeld (1971) as "moisture indicator
plants" and used to reflect changes in vegetation moisture content during

�-338measurement periods were collected in one unit representing each type of
treatment
(Units 5, 6, 7, 8) between July 24, and 27, 1973, inclusive on
November 12, 1974, and between July 19 and 22, 1976 inclusive.
Composite
samples of each species weighing approximately
100 grams green weight were
collected throughout each of the 4 habitat improvement units.
In treated
units samples were collected from spots where it was obvious that burning,
spraying and chaining had been severe.
Samples were analyzed by the Division Research Laboratory to determine
changes in content of dry matter, protein, energy and trace mineral values
2 and 5 summers and 3 winters after treatments were applied.
Soil samples were also collected from each of the 4 areas from which plant
nutrient samples were obtained.
These were analyzed by the Colorado State
University Soils Laboratory to determine any differences among areas in soil
properties.
Elk, Deer and Cattle Use Measurements 2 and 5
Years After Burning, Spraying and Chaining
Elk use was measured one year before and 2 and 5 years after treatment.
The
same schedule was followed for deer except that deer use was also measured
after 3 years, because deer use during the 2nd year appeared relatively low,
and it was deemed desirable at that time to collect additional data for deer.
Accumulated elk and deer pellet groups were removed from all pellet plots
about September 1st. Groups deposited during the fall, winter and spring
were counted during early May of the next year.
Cattle use was evaluated one year before, and 2 and 6 years after treatment.
A 5-year evaluation could not be made because the area is under a rest-rotation
grazing system, and was not grazed during the 5th year.
Cattle use was
measured by counting cow-chips deposited in pellet plots during the current
season after the grazing season ended and cattle had been removed.
RESULTS

AND DISCUSSION

Vegetation Changes 2 and 5 Years After
Burning, Spraying and Chaining
Vegetation
vegetation

production estimates discussed throughout
which occurs below a level of 5 feet.

this report refer to

All production changes have been adjusted for normal year to year fluctuations
in plant growth as measured on control units.
Thus, vegetation changes on
burned, sprayed and chained areas discussed in this report reflect the response
which plants actually exhibited to the treatment.
Adjustments were made by
computing the level of vegetation production one would expect to find on a
treated area following treatment had treatment not occurred.
This expected
value, hereafter referred to as
is derived by multiplying the ratio of preto post-treatment
forage production on the control areas to pre-treatment

E,

�-339-

production on the treated areas. The difference between the expected
post-treatment production (E) on the treated areas and the actual production
recorded there is the change produced as a result of treatment.
Percentage
production changes mentioned herein always relate to pre-treatment levels.
Effects of Burning on Forbs, Grasses and Shrubs
Summer vegetation production decreased 27 percent 5 years after burning.
This represents a further decline from the 17 percent decrease recorded
after 2 years (Table 1, Fig. 1). Most of the reduction occurred among shrubs
which were down 38 percent during both the 2 year and 5 year evaluations.
The shrub declines after 2 and 5 years were significant at the a.05 and a.l0
levels respectively (Table 1).
The shrub species mainly responsible for the overall decrease in shrub
yields as well as in overall vegetation production after 2 and 5 years were
AfLtem-LoJ..a. bU.de..n:ta.;ta and SymphoJzA..ecu1.po,6 Cltb1L6 (Tables 2, 3, and 4). These
showed decrease of 92 and 51 percent, respectively, after 2 years, and 94
and 51 percent after 5 years.
AfLtem-L6J..a. bude..n;ta.;ta
did not resprout after
burning.
Those plants exposed to fire were completely killed.
SymphoJzA..e~po,6
Cltb1L6 sprouted profusely, especially the first summer after burning, however,
overall production was still substantially lower 2 and 5 years after treatment.
Other shrubs among which much sprouting and regrowth was noted after burning
were Ame1.anc..h.i..eA CltM-60.u..a., AfLtem-L6J..a. c..a.n.a., PJULn1L6 vbtgil'u..a.na. and QUeAc..LL6
gambettii.
Like SymphoJzA..c..~po,6
CltbLL6 most of the sprouting of these 4 species
occurred during the first summer and slowed considerably the second summer
(Fig. 2). Percentage production changes for the 4 species after 2 years were
-21%, +149%, -7% and +11% respectively (Table 3).
Five years after burning production of Ameil1nc..h.i..eA CltM-60.u..a. and Ptwn1L6
vbtginA..a.na. had changed very little from the level measured after 2 years.
Production of AJttem-L6ia. c..a.n.a.and QUeAeLL6 gambeLeil,
however, showed increases
of +280% and +38% respectively after 5 years (Table 4). Fig. 2, shows that
the vegetation stand was still substantially more open after 5 years than it
was before burning.
Grasses on burned areas showed a decrease of 5 percent after 2 years and 30
percent after 5 years (Table 1, Fig. 1).
Changes in production of individual
grass species on the burned areas, in terms of pounds per acre, were relatively
minor after 2 years and, with 2 exceptions, after 5 years, even though some
species showed fairly large percentage
changes (Tables 3 and 4). The 30 percent
decline in grasses after 5 years was largely attributable to declines in Poa.C~e..x ge..ye..Jui.. and Fe...6tuc..a. thunbe..Jui.. which decreased 257 and 157 lbs/ac.,
respectively.
Poa. sp. and C~e..x ge..ye..Jui.. were combined in this study because it
was often difficult to separate them when clipping vegetation in meter square
sampling plots.
Changes in production of individual forb species measured in pounds per acre
were relatively small after 2 and 5 years even though percentage changes shown
by some species were relatively large (Tables 3 and 4).

�-340-

Effects

of Spraying

on Forbs, Grasses and Shrubs

Spraying resulted in the largest total vegetation increase of the 3 treatments tested after 2 years.
Vegetation production increased 50 percent 2
years after spraying.
Most of the increase occurred among grasses which
were up 295 percent and produced an additional 658 lbs/ac (air dry wt).
This was significant at the a.Ol level (Table 1, Fig. 1). Five years after
spraying this large increase was completely eliminated, and total vegetation
production showed a net decrease of 14 percent.
Most of this decrease
occurred among grasses which were measured at 16 percent below pre-treatment
levels (Table 1, Fig. 1). Stimulated grass production after 2 years and the
ensuing decline between 2 and 5 years is evident in Figure 3, which shows
photos taken at a permanent photo point 1 year before and 1, 2 and 5 years
after spraying.
Several individual grasses were primarily responsible for the 2-year increase
as well as the ensuing 5th year decline.
Poa sp. and C~ex gey~
together
produced 212 percent or 489 lbs/ac more air dry herbage after 2 years; however
5 years following spraying they showed a net decrease of 55 percent or 200
lbs/ac from pre-treatment levels.
Other grasses which showed proportionately
large increases in air dry herbage after 2 years and subsequent declines in
production between 2 and 5 years we re AgJtopYJton.sp , (2 yrs = +42 lbs/ ac, 5 yrs
+33 lbs/ac), BJtomU6 an.emaiU6 (2 yrs = +28 lbs/ac, 5 yrs = +13 lbs/ac), VactylU6
glomenata (2 yrs = +8 lbs/ac, 5 yrs = 0 lbs/ac), Elymu¢ glau~u¢ (2 yrs = +27
lbs/ac, 5 yrs = +7 lbs/ac), and Stipa sp. (2 yrs = 24 lbs/ac, 5 yrs = +2 lbs/ac)
(Tables 5 and 6).
Forbs also increased after 2 years although not as dramatically as did grasses.
Forbs were up 15 percent or 79 lbs/ac, air dry weight.
They also suffered a
subsequent decline, and after 5 years forb production was 5 percent (16 lbs/ac)
below pre-treatment levels (Table 1, Fig. 1). Individual forbs which reflected
the largest increases after 2 years but subsequent declines between 2 and 5
years were:
A~hLttea lan.ulo~a (2 yrs = +49 lbs/ac, 5 yrs = +5 lbs/ac), Galium
sp. (2 yrs = +39 lbs/ac, 5 yrs = +5 lbs/ac), and TaJtaxa~um o66lein.ale (2 yrs =
+30 lbs/ac, 5 yrs = 0 lbs/ac).
Some forbs which decreased by relatively large
amounts after 2 years and which recovered somewhat between 2 and 5 years after
spraying were:
LathYJtU¢-vleia (2 yrs = -53 lbs/ac, 5 yrs = -27 lbs/ac) and
Lupln.u¢ ~genteu¢ (2 yrs = -37 lbs/ac, 5 yrs = -15 lbs/ac) (Tables 5 and 6).
The kill on QueJt~U¢ gambettLi
first summer after spraying.

was estimated at 80 to 90 percent during the
Extensive initial kills were also observed for
Amefun.~MeJt aiM6o~,
AJdem.v.,la .:t:JUdentata,Pnunus vlJtgln.lan.aand Sympho~~~po.6
albU6. Two years after treatment total shrub production on spray areas
exceeded by 4 percent or 35 lbs/ac that which would have been expected without
spraying (Table 5). This relatively minor change reflects a situation where
many shrubs, and particularly those 5 just mentioned, have sprouted back within
2 years to produce approximately as much herbage as before treatment (Fig. 3).
Like forbs and grasses, shrubs also declined between 2 and 5 years after
spraying.
After 5 years shrub production, less than 5 feet in height, was
14 percent below pre-treatment levels.
The same 5 species were responsible
for the bulk of this decline in shrub abundance (Table 6). Above 5 feet in

�-341-

height, however, a great deal of top sprouting was evident, after 5 years
on, Qu~e~
gamb~
plants which had initially appeared killed by spraying
(Fig. 3).
Effects of Chaining

on Forbs, Grasses and Shrubs

Production of vegetation on chained sites increased 20 percent 2 years after
chaining, but exhibited a net decrease of 19 percent after 5 years (Table 1,
Fig. 1).
Shrubs, which produced 19 percent or 167 more Ibs/ac of air dry vegetation,
accounted for most of the 2-year increase.
The increase in shrubs was
significant at the a.15 level (Table 1, Fig. 1). Most of the higher shrub
production was due to increase of 33 percent or 107 lbs/ac in Qu~e~
gamb~,
65 percent or 90 lbs/ac in Pnun~ vinginiana and 33 percent or 35 Ibs/ac
in Ameianehi~
atni6olia. These 3 species sprouted profusely after chaining
(Fig. 4). SymphonieMpo/.) atblL6 produced 14 percent or 31 Ibs/ac less forage
2 years after chaining, and resprouting was not as noticeable on this species
as it was in burned and sprayed areas (Table 7).
Shrub production dropped to 12 percent below pre-treatment levels after 5
years.
The species responsible for the net decline after 5 years were the
same 4 which caused the increase after 2 years, plus Antemi.6ia tnidentata.
Qu~e~
dropped to 23 percent or 45 lbs/ac above pre-treatment levels.
Pnun~,
Amei.anehi~, SymphonieMp0.6 and Antemi.6ia .tft.A..dentata
declined to 20, 12, 39
and 45 percent below pre-treatment levels,respectively
(Table 8).
Forbs increased 25 percent or 108 Ibs/ac on chaining after 2 years, but
reverted to pre-treatment levels after 5 years (Table 1, Fig. 1). Four species
were largely responsible for the 2 year increase and subsequent 5th year decline.
These were:
Aduliea ffinui.0.6a(2 yrs = +29 Ibs/ac, 5 yrs = +l3 Ibs/ac), A/.);t~~og~on
(2 yrs = +16 Ibs/ac, 5 yrs = -11 Ibs/ac), Galium (2 yrs = +25 lbs/ac,
5 yrs = +20 Ibs/ac), and La;thyn~-vieia (2 yrs = +20 lbs/ac; 5 yrs = -15 Ibs/ac)
(Tables 7 and 8).
Chainings also supported higher grass production after 2 years with an increase
of 17 percent or 65 Ibs/ac (Table 1, Fig. 1). Two year post-treatment changes
in production of individual grass species on the chainings in terms of pounds
per acre were relatively minor (Table 7). Grasses, however, were primarily
responsible for the net decrease in total vegetation production on chainings
after 5 years.
The 5th year, grasses had declined 48 percent or 227 lbs/ac
below pre-treatment levels.
This was due largely to a 71 percent or 473
Ib/ac decrease in production of Poa. and Can:ex geyeni (Table 8).
Contribution of Seeded Species to Vegetation
and Chained Areas.

Production

in Burned,

Sprayed

Plummer et al. (1970) were able to retard regrowth of Gambel oak thickets
for 14 years by seeding heavily after treating by burning or mechanical means.
Therefore, all areas treated as part of the Hightower Mountain oak study were
seeded at a rate of 18.9 Ibs per acre with the following species:
Metllo;tU,6

�342-

o66~ein~,
Medieago ~ativa (Ladak, Rambler and Nomad strains), AghOPYhon
e~tatum, AghOPYhon ~ntekmedium, BhOm~ ~n~,
Vactyl~ glom~,
F~tuea
ov~na, CeheOeahp~6 montan~, and Punohia ~dentata.
Seeding rates for
each species were described

by Kufeld

(1972).

Seeding success after 2 years was negligible.
Part of the reason is believed
to be extremely dry weather conditions during the winter, spring and summer
of 1972, following seeding in November, 1971. U.S. Weather Bureau records
from Collbran show precipitation during January through August, 1972 was only
31 percent of normal and only 53 percent of normal at Glenwood Springs.
During the first summer new plants of seeded species could only be found
in the fire lanes constructed around the burns, and in small patches within
one burn.
These consisted of only sparse stands of AghopYhon ~ntekmedium
and Bhom~ ~n~.
Precipitation improved somewhat the second year.
Between January through
,August, 1973 precipitation was 84 percent of normal in Collbran and III
percent of normal in Glenwood Springs.
Seeded species began to appear the
second summer and Aghophyon ~ntekme~m
and Bhom~ ~neh~
began to form
fair stands in the fire lanes and part of one burn (Fig. 5). AghopYhon
~ekmedium only produced 12 lbs/ac of air dry forage in the burns including
fire lanes, and Bhom~ ~n~
produced 3 lbs/ac.
The only other seeded
species which produced more than 1.00 lb/ac was Vacty~
glomehata which
produced 1.12 lbs/ac in the chainings and 8.09 lbs/ac in spray areas.
Since
this grass was already present in one spray area prior to treatment it is
felt that higher production was due to existing plants being released by
spraying and not to the seeding effort (Tables 3, 5 and 7).
During the 5th summer production of seeded species was still negligible and
relatively unchanged from that recorded after 2 years.
AghopYhon ~ntekmedium,
Bhom~ ~neh~ and Vactyl~ glomehata produced 9, 7 and 3 lbs/ac respectively
in the burns.
Vactyl~ glomehata also produced 5 lbs/ac in the chainings.
The only other species which produced more than 1.00 lb/ac was AghopYhon
e~tatum which produced 1.48 lbs/ac in the chainings.
Considering that seeded species only established themselves in heavily
distributed spots in one burn, such as fire lOanes and parts of the chainings
where bulldozer tracks had dug deeply it appears that better seedbed preparation is needed than was afforded by the treatments used in this study.
It
may be more effective to burn, aerially seed as soon as the ashes are cool,
and, since burning in oakbrush is usually patchy, pipe harrow the spots where
burning is not severe.
This would plant seed as well as remove burned snags
from the created openings.
Among Treatment

Comparisons

of Vegetation

Responses

Forbs increased from 14 to 25 percent after 2 years on all 3 treatments.
After
5 years forbs showed an increase of 7 percent on burns, a decrease of 5 percent
on sprayed areas, and no change due to treatment on chainings.
The differences
in forb production among treatments were not significant at even the 25 percent
significance level after either 2 or 5 years (Table 1, Fig. 1).

�-343-

Grass production 2 years after spraying was significantly higher than
burning (P &lt; .10) or chaining (P &lt; .15). Air dry production of grasses
on the sprayed areas increased 295 percent or 658 lbs/ac compared with a
5 percent or 16 lb/ac decrease on burns, and 17 percent or 65 lb/ac
increase on chainings.
Grasses decreased from 30 to 47 percent after 5
years on all 3 treatments.
There were no significant differences even
at the 25 percent level among treatments in grass production after 5
years (Table 1).
Shrub production in burns 2 and 5 years after treatment was significantly
lower (P &lt; .05 after 2 yrs; P &lt; .15 after 5 years) than sprayed or chained
areas.
Shrubs on burned areas showed a 38 percent decrease after 2 and
5 years.
On sprayed and chained areas shrubs increased 4 and 19 percent,
respectively, after 2 years but decreased 14 and 12 percent, respectively,
after 5 years.
The difference in shrub production between sprayed and
chained units was significant after 2 years, but only at the a.20 level.
It was not significant even at the a.25 level after 5 years (Table 1).
Production of forbs, grasses and shrubs declined on all treated areas
between 2 and 5 years after treatment (Fig. 1). Except for a minor increase
among forbs on burns, and a no change due to treatment situation among forbs
on chainings, vegetation production for all plant categories on all 3
treatments was below pre-treatment levels after 5 years.
It is probable
that these declines were a result of abnormally dry weather conditions which
prevailed between 2 and 5 years after treatment plus a severe frost which
occurred during late spring of the 5th year. U.S. Weather Bureau records
for Collbran, Colorado for the period January thru August show that precipitation was only 45 percent of normal the 3rd year after treatment, 109
percent of normal the 4th year, and 61 percent of normal the 5th year.
In
addition to dry weather the 5th year (the year measurements were made) a
killing frost occurred for 3 nights on June 13, 14 and 15, when many plants
were in initial stages of spring growth.
This had a very visible adverse
impact on vegetation in the area. The effects of the frost and dry conditions
on vegetation were obvious for the remainder of the 5th summer, and appeared
more severe on treated than control areas.
Since production changes have been adjusted for normal year to year fluctuations
in plant growth as measured on control units, the 5th year vegetation declines
on burned, sprayed and chained areas reflect the response of plants to the
treatments.
Thus, a decline on a treated area represents a negative treatmentinduced effect on vegetation which did not occur to vegetation growing on
control areas.
Since vegetation on controls was subject to the same weather
conditions, the occurrence of declines among all plant categories on all
treated areas suggests that removal of the overstory ata time of drought plus
effects of the 5th year late spring frost exposed the understory plants to
conditions which they could not tolerate.
Therefore they exhibited a greater
production decline during the 5th year than would have occurred had overs tory
vegetation not been removed.

�-344-

Changes in Elk, Deer and Cattle Forage 2 and 5 Years After Burning,
and Chaining

Spraying

Definition of "Elk, Deer and Cattle Forage"--Classification
of study area
plants as elk or deer forage for purposes of determining production changes
was based on the average, relative occurrence of those plant species in 48
elk and 99 mule deer food habits studies reported in the literature (Kufeld
1973; Kufeld, Wallmo and Feddema 1973). Plants considered cattle forage are
those classified as "desirable" and "intermediate" by the U.S. Forest Service
in their Region 2, Range Analysis Handbook for the foothill shrub, aspen
week, and mountain meadow vegetation types.
Since animal preference for individual plant species varies from location
to location the validity of applying "average" occurrence data to this study
area may be open to question.
However, these averages do represent a great
deal of food habits research, and constitutes the best information available
to date on the food these animals have been known to consume and the relative
amounts consumed.
For elk the following classifications are used: highly valuable, valuable
and least valuable (Kufeld 1973).
For deer plants are classified as heavily
eaten, moderately eaten, or lightly eaten (Kufeld, Wallmoand Feddema
1973).
Although the terminology used in the elk and deer references is different
the basis for classifying plants according to relative consumption is exactly
the same.
"Highly valuable" plants for elk and "heavily eaten" plants for
deer, by definition, generally
comprised a major part of a food sample
(usually at least 20 percent) in food habits studies where reported.
In a
few cases, plants which comprised less than a major portion of the food
sample were classified as "highly valuable" or "heavily eaten" if their reported
contribution to the diet was far in excess of their reported vegetative
composition.
"Valuable" plants for elk "moderately eaten" plants for deer
usually comprised between 5 and 20 percent of the food sample.
"Least valuable"
plants for elk and "lightly eaten" plants for deer comprised less than 5 but
more than I percent.
Not all plants on the Hightower Mountain oak study area were classified
according to their relative preference as elk and deer forage.
Some had never
been reported in the literature as eaten by elk or deer during some seasons
or at all during the year. The absence of such plants from the literature
possibly suggests that they are of minor importance as elk or deer forage.
Therefore, for purposes of the Hightower Mountain oak study they will not
be considered elk or deer forage.
Definition of "Available Forage"--Data and discussions relating to available forage are based on the assumption that due to factors such as degree
of plant maturity and snow cover, plant availability to elk and deer on the
study area will consist of only shrubs during winter; grasses and shrubs during
spring and fall, and forbs, grasses and shrubs during summer.
Discussions
of cattle forage relate only to summer at which time all forage is available.

�-345-

Definitions of seasons used here are: Winter = December, January, February;
Spring = March, April, May; Summer = June. July, August; Fall = September,
October, November.
All data are based on measurements made during summer.
Procedure for Expressing Changes in Available Forage--A pre-to posttreatment ratio using 1.00 as a basis to represent the level of forage
production before treatment with adjustment for normal year to year changes
measured on control areas was computed for elk, deer and cattle (Table 9).
A single value for elk and deer forage was derived by weighting lbs/ ac of
highly valuable elk forage and heavily eaten deer forage by a factor of "3",
valuable and moderately eaten by "2", and least valuable and lightly eaten
forage by "1". A total weighted value was then computed.
For cattle forage
desirables were weighted by "2" and intermediates by "1".
Effects of Burning on Available Elk Forage--After 2 and 5 years burned
areas supported less available elk forage during all 4 seasons of the year
than existed before treatment.
1~0 years after burning the level of available
forage production compared to 1.00 was 0.64 for winter, 0.84 for spring, 0.80
for summer and 0.67 for fall.
The 5th year spring, summer and fall elk-forage showed additional declines.
Winter forage exhibited a slight increase.
Pre-to post-treatment forage
production ratios after 5 years were: winter - 1:0.74, spring - 1:0.60,
summer - 1:0.62 and fall 1:0.60 (Table 9).
Effects of Spraying on Available Elk Forage--Spraying
produced considerably more available elk forage during spring, summer and fall after 2 years
than existed before treatment.
This was mainly due to large increases in
grasses which occurred 2 years after spraying.
No change attributable to
treatment was recorded in production of available winter elk forage.
Preto 2 year post-treatment ratios of elk forage production were: 1:1.01 for
winter, 1:2.09 for spring, 1:1.72 for summer and 1:1.94 for fall.
Production of available forage for all seasons declined substantially the
5th year from 2-year levels.
This was largely the result of the decreases
in grasses which we re recorded after 5 years.
Pre-to 5 year post-treatment
ratios of elk forage were: winter - 1:0.85. spring - 1:0.72, summer - 1:0.76,
fall - 1:0.77 (Table 9).
Effects of Chaining on Available Elk Forage--Chained
areas supported
increased amounts of available winter, spring, summer and fall elk forage 2
years after treatment.
Respective pre-to post-treatment elk forage production
ratios were:
1:1.24, 1:1.21, 1:1.13, and 1:1.12.
Production of available elk forage declined for all seasons the 5th year
from 2-year levels.
Pre-to 5 year post-treatment ratios of elk forage were:
winter - 1:0.75, spring - 1:0.66, summer - 1:0.60, fall - 1:0.65 (Table 9).
Effects of Burning on Available Deer Forage--Two and 5 years after
treatment burned areas supported less available deer forage during all 4 seasons

�-346-

of the year than existed before burning.
Pre-to post-treatment forage
production ratios after 2 years were: winter - 1:0.54, spring - 1:0.59,
summer - 1:0.84, fall - 1:0.66.
The 5th year relatively small additional declines were recorded for spring
and summer forage while slight increases compared to 2 year levels were
observed in winter and fall forage.
Pre-to 5 year post-treatment forage
production ratios for winter, spring, summer and fall, respectively were
1:0.65, 1:0.52, 1:0.72, and 1:0.78 (Table 9).
Effects of Spraying on Available Deer Forage--Spraying produced more
available deer forage during spring, summer and fall after 2 years than
existed before treatment.
Production of available winter deer forage did
not change from pre-treatment levels.
Pre-to 2 year post-treatment ratios
of deer forage production were: winter - 1:1.02, spring - 1:1.40, summer _
1:1.20, fall - 1:1.13.
Production of available spring, summer and fall deer forage declined substantially the 5th year from 2-year levels.
Production of available winter
deer forage still remained relatively unchanged from pre-treatment levels.
Pre-to post-treatment forage production ratios the 5th year were: winter1:0.98, spring - 1:0.82, summer - 1:0.79, fall - 1:0.91 (Table 9).
Effects of Chaining on Available Deer Forage--Available
forage production
increased for all seasons 2 years after chaining.
Respective winter, spring,
summer and fall pre-to post-treatment forage production ratios were: 1:1.23,
1:1.17, 1:1.18 and 1:1.26.
A decline from 2-year levels was recorded for all seasons in available deer
forage during the 5th year.
Pre-to 5 year post-treatment ratios of deer
forage were: winter - 1:0.98, spring - 1:0.76, summer - 1:0.82, fall - 1:0.96
(Table 9).
Effects of Burning on Cattle Forage--Summer cattle forage declined
slightly from pre-treatment levels after 2 years, and a further decline was
recorded after 5 years.
Two- and 5-year pre- to post-treatment forage production
ratios were 1:0.95 and 1:0.76 (Table 9).
Effects of Spraying on Cattle Forage--Substantial
increases in summer
cattle forage were observed on sprayed areas after 2 years.
The pre- to
2-year post-treatment forage production ratio was 1:2.17. A steep decline
occurred between 2 and 5 years after spraying.
The 5th year cattle forage
production was below pre-treatment levels with a ratio of 1:0.84.
The 2
year increase and 5th year decline was due to the sharp initial increase and
subsequent decline in grass abundance (Table 9).
Effects of Chaining on Cattle Forage--Chaining
resulted in an increase
of 1:1.24 in cattle forage after 2 years.
Five years after chaining cattle

�-347-

forage had declined to a level below pre-treatment levels at which
time a ratio of 1:0.70 was recorded.
The 5th year decline was due to
a 48 percent decrease in grass production (Table 9).
Nutrient and Mineral Content of Forage Collected
Sprayed and Chained Areas Following Treatment.

on Control, Burned,

Results of soil tests and nutrient and mineral analysis of plants collected
during the second summer and third winter after treatment were discussed
in detail by Kufeld (1975). These data indicate burned and chained areas,
when compared with control and sprayed areas, showed increased levels of
total cell contents, soluble
carbohydrate, copper and zinc in most plant
categories (forbs, grasses and shrubs).
Laboratory
completed.

analysis of plants collected during the 5th summer has not been
Results of those analyses will be presented in a future report.

Elk Use Changes

2 and 5 Years After Burning,

Spraying

and Chaining

Elk, deer and cattle use were measured through use of fecal counts.
The
possibility exists that these ungulates may not defecate in patterns related
to feeding activity.
If they defecate more frequently at bedding sites or
in transmit to and from preferred feeding areas then inferences to habitat
type preferences for feeding would be biased.
The study was originally
designed to attempt to minimize effects of the possible occurrence of such
a bias. Each of the treated units was established with a 30 acre treated
portion with a surrounding 30 acre untreated area suitable for aninlal resting
or bedding.
Fecal plots were distributed throughout both sections.
Elk pellets were significantly more abundant in burned units 2 years after
treatment despite declines in available, seasonal elk forage on burns compared
with sizeable increases on sprayed and chained areas.
Sprayed areas ranked
second and chainings third in pellet abundance.
Calculated densities of elk
per square mile increased 45 percent on burns during the fall, winter and
spring period, and decreased 48 percent on sprayed areas and 70 percent on
chainings (Table 10). Compared with control areas, however,-changes
in elk
use due to burning, spraying and chaining were not statistically significant
a~ the a.10 level even though percent changes were relatively large.
Five years after treatment elk continued to exhibit preference (as indicated
by relative pellet abundance) for the burned areas with sprayed areas still
ranking second and chainings third. Compared with the pre-treatment level
elk per square mile during the Fall-Winter-Spring period increased 54 percent
on burns, 27 percent on sprayed areas and 12 percent on chainings (Table 10).
Although the increase in elk use on burns after 5 years was higher than after
2 years (54% compared with 45%) the increase was not significant (P &gt; .10) as
was the 2 year increase.
The 5 year increases on sprayed and chained areas,
likewise, were not significant.
The primary reason for lack of significance among treatments in the covariance
analysis during the 5th year was due to the relatively large error term caused

�-348-

by high variation in elk use between the 2 sprayed areas.
One sprayed
area received use of 50 elk per square mile, while the other received only
15. There is no apparent explanation for this variation.
Elk use on the
other units which received similar treatments and on the controls was
relatively consistent during all years when measurements were made.
Reasons for apparent preference of elk for burned areas over those sprayed
and chained cannot be pinpointed.
Two factors, however, which may have
influenced the attractiveness of burned areas to elk are changes in cover
and plant nutrient content.
Cover Changes
Burning did not denude the area, but varied from almost total incineration
of all vegetation with just a few dead snags left standing (Fig. 6) to only
partial removal of the understory with little or no damage to the larger oaks
(Fig. 7). On much of the area most of the understory was removed with partial
destruction of the overstory.
The overall result of burning was creation of
openings interspersed with patches of standing oak vegetation suitable for
escape cover and shade (Fig. 8). This situation has lasted for 5 years to date.
In sprayed areas most of the shrub vegetation was either totally or partially
killed, but dead material remained standing for the first few years leaving
a thicket only slightly more open and accessible to large animals than before
treatment.
Five years after spraying dead trees and branches had begun to
sag and fall over against other trees making access difficult (Fig. 3).
After 2 and 5 years chained areas were very open with.little escape cover
and no trees large enough to provide shade (Fig. 4). Forest openings have
been found to receive increased use by elk and deer (Patton, 1974; Pearson,
1968, Reynolds, 1962a, 1962b, 1966a, 1966b; Wallmo 1969). However, during
the fall of 1973, when 2 year post-treatment ungulate use measurements were
being made this study area received unusually heavy hunting pressure particuarly for deer as efforts were being made to reduce the deer herd. Hunting
pressure coupled with the lack of cover in chainings may have caused elk to
avoid them during that period, and resulted in the substantial drop in elk use
recorded on chainings 2 years after treatment (Table 10). Exceptionally heavy
hunting pressure did not occur the 5th year and elk use on chainings was more
in line with that on other areas.
Plant Nutrient

Content

Changes

Elk preference for burned areas may have been influenced to some extent by
increased levels of certain nutrients and minerals such as total cell contents,
soluble
carbohydrate, copper and zinc. As described by Kufeld (1975) increased
amounts of these were found after 2 years in most plant categories (forbs,
grasses and shrubs) on the burned areas as compared with control and spray areas.
Since relatively little difference was observed between the burn and chaining
in plant content of these nutrients and minerals, however, any influence
exerted by increased plant nutrient and mineral content must have been coupled

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by other factors such as the previously mentioned cover changes
changes in elk movements within and outside the study area.

Deer Use Changes

2, 3 and 5 Years After Burning,

Spraying

or inexplicable

and Chaining

Deer did not appear to exhibit preference (based on fecal abundance) for
any particular treatment after 2, 3 or 5 years.
Within each treatment deer
use exhibited a sizeable increase on one replication, but this was offset
by either a decrease or only a small increase on the other replication.
After 2 years, data show average decreases in deer use associated with the
treatment on each pair of treated areas. Deer use on sprayed areas during
fall-winter~spring
periods decreased 6 percent and decreases of 25 and 53
percent were recorded on the burned and chained areas, respectively (Table
11). The large drop in deer use on chainings after 2 years was significant
at the a.05 level. As discussed in the section on elk use this was speculated
to be, at least in part due to unusually heavy hunting pressure in the study
area vicinity during the fall of the second year, coupled with the lack of
overstory in chained areas.
An average increase of 75 percent in deer use occurred on the sprayed areas
after 3 years but this was due to a large increase on one replication.
Deer
use on the other spray area changed very little.
Average decreases of 11
and 27 percent respectively were recorded on burned and chained areas.
None of the changes due to treatment were significant at the a.05 significance
level.
Five years after treatment average percentage increases of 79 percent, 67
percent and 17 percent occurred on the sprayed, chained and burned areas,
respectively.
In each case, however, the size of the increase was primarily
due to a large increase on one replication.
None of the changes due to
treatment were significant at the a.05 significance level (Table 11).
Cattle Use Changes

2 and 5 Years After Burning,

Spraying

and Chaining

Control Areas
Data show cattle use (based on fecal counts) increases for both control areas
after 2 years with an average increase of 4.1 cow days use per acre (Table 12).
Much of the increase in control unit 6 is believed due to movement of cattle
around unit 7 (a burned area) and between units 5 (a sprayed area) and 7 to
which cattle appeared to be attracted.
Unit 6 lies directly between units 5
and 7. Thus, there was very little, if any, actual change after 2 years in
cattle use on the control areas.
Data also show very little difference in cattle use on control areas between
levels recorded before and 6 years after treatment.
Use in Unit 4 increased

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..2 cow days per acre and use in Unit 6 decreased 1.3. The average
~ontrol units was an increase of 1.0 cow days per acre.

for both

Burned Areas
The most preferred treatment by cattle 2 years after treatment was definitely
burning, even though cattle forage production decreased on burns compared
to sizeable increases on sprayed and chained areas.
Both burns showed the
largest increases in cattle use after treatment, and these were significant
at the a.10 level (Table 12). Cattle use in burned units increased 15.2
and 22.1 cow days per acre respectively, and averaged 18.7. Cattle preference for burned areas was also obvious from field observations.
Preference
for burns after 2 years was approximately 6 times greater than for the
sprayed or chained areas.
The reasons for cattle preference for burns over sprayed and chained areas
during the 2nd year have not been specifically identified.
Possible reasons
are creation of openings interspersed with patches of cover and shade in
burned areas and increased amounts of certain nutrients and minerals in
plants found on burns.
These changes may have also been responsible for
increased elk preference for burned areas over those sprayed and chained,
and are discussed in detail in the previous section of "elk use changes 2
and 5 years after burning, spraying and chaining".
After 6 years cattle use in units I and 7 increased 5.2 and 0.4 cows days
per acre, respectively, and averaged 2.8. These minor increases were not
significant at the 10 percent significance level, and do not suggest
a preference for burned areas 6 years after burning (Table 12).

Sprayed Areas
One spray area (Unit 5) showed a substantial increase in cattle use after 2
years as a result of spraying, while the other (Unit 2) showed a slight decrease (Table 12). According to the data, which show an average increase of
3.3 cow days use per acre in sprayed areas, spraying would appear to be the
second most preferred treatment by cattle during the 2nd year.
Since the
cattle use change on the combined spray areas is not significant at the a.10
significance level, and in light of circumstances described in the following
discussion on chaining, it is not valid to conclude that cattle preferred
the sprayed areas more than the chained areas at the end of the 2nd year
following treatment.
Cattle use after 6 years decreased 7.4 cow days per acre in Unit 2 and increased 2.2 in Unit 5. The average for both units was a decrease of 2.6
cow days per acre which was not significant at the 10 percent significance
level.

�-351-

Data do not reflect a tendency of cattle to prefer or avoid sprayed areas
after either 2 or 6 years.

Chained Areas
No cattle use was recorded in chained unit 3 during the 2nd or 6th year
after treatment (Table 12). This unit lies in 2 cattle allotments, and
due to surrounding thick vegetation is difficult for cattle from both
allotments to find. It is also on the opposite side of a creek from one
allotment, and high water during the 2nd year barred access to cattle
from that allotment.
This was not a factor during the 6th year, however,
since the creek was dry. The almost impenetrable oak thicket in unit 3
was opened substantially by chaining, and it is felt cattle use would have
occurred after treatment if cattle had been able to get to the area.
Cattle use in unit 8 dropped from 18.3 cow days per acre before chaining
to 8.7 two years after chaining, and 6.5 after six years (Table 12). During
the 2nd year it appeared that cattle were simply drawn off unit 8 into the
more preferred burned area of adjacent
unit 7. This occurred even though
forage conditions in unit 8 were improved for cattle by chaining.
During
the 6th year, however, cattle did not exhibit a preference for burns, yet
use in unit 8 continued to decline slightly.
The 2nd and 6th year declines
attributable to chaining were not significant at the a.10 level.
In view
of the lack of statistical significance of these declines and the inability
of cattle to find and get into unit 3, it is not valid to conclude that
chained areas were any less preferred by cattle than sprayed areas. Data
do not reflect a tendency of cattle to prefer or avoid chainings after either
2 or 6 years.

SUMMARY AND TENTATIVE

CONCLUSIONS

1.

The effects of controlled burning, spraying with 2,4,5-TP herbicide
and anchor chaining of Gambel oak type vegetation were studied at a
site on the Grand Mesa National Forest, 20 miles east of Collbran,
Colorado, between 1969 and 1976. Vegetation production and deer,
elk and cattle use were measured one year before and 2 and 5 years
after treatment.
The study area consisted of eight 60 acre areas
which included 2 controls, 2 burns, 2 sprayed areas and 2 chainings
arranged in a randomized block design.
The center 30 acres of each
area were treated. The surrounding 30 acres were left intact to
provide cover for deer, elk and cattle.

2.

Responses of 92 kinds of forbs, 19 grasses and 16 shrubs to burning,
spraying and chaining have been documented at intervals of 2 and 5
years after treatment.
Spraying resulted in the largest increase
in vegetation production (+50%) after 2 years, with chaining second

�-352-

(+20%), and burning third (-17%). Burning actually caused a net
decrease in vegetation production after 2 years. Treatment methods
are arranged, as follows, according to those which produced the most
to the least amounts of vegetation (percentage wise) within each plant
category after 2 years:
Forbs, I-chaining, 2-spraying, 3-burning;
Grasses, I-spraying, 2-chaining, 3-burning; Shrubs, I-chaining, 2spraying, 3-burning.
Production of forbs, grasses and shrubs declined on all treated areas
between 2 and 5 years after treatment.
Except for a minor increase
among forbs on burns, and a no change due to treatment situation
among forbs on chainings, vegetation production for all plant categories
on all 3 treatments was below pre-treatment levels after 5 years (Spray
-14%, chain = -19%, burn = -26%).
Still, some treatments resulted in
less of a net decrease than others.
Treatment methods are arranged
as follows according to those which produced the most to the least amounts
of vegetation (percentage wise) within each major plant category after
5 years:
Forbs, I-burning, 2-chaining, 3-spraying; Grasses, I-spraying,
2-burning, 3-chaining; Shrubs, I-spraying, 2-chaining, 3-burning.
3.

Net vegetation declines due to the 3 treatments after 5 years may have
been a normal response of vegetation to treatments.
It seems probable,
however, that these declines were at least partially, if not entirely,
the result of abnormally dry weather conditions which prevailed between
2 and 5 years after treatment, plus a severe frost which occurred during
late spring of the 5th year, just prior to the 5 year vegetation measurement period.
Since vegetation on controls was subject to the same
weather conditions, the occurrence of net declines among all plant
categories on all treated areas suggests that removal of the overs tory
during a period of drought plus effects of the 5th year late spring frost
exposed the understory plants to conditions they couldn't tolerate.
Therefore they exhibited a greater production decline during the 5th year than
would have occurred had overstory vegetation not been removed.

4.

Seeding success after 2 and 5 years was negligible. Part of the reason
is believed to be extremely dry weather conditions during the first winter,
spring and summer after seeding when precipitation at the nearest weather
station was only 31 percent of normal.
Only 2 seeded species, Ag~opy~on
intenm~dium and B~omU6 in~
showed any success.
These established
themselves only in part of one burn, and in heavily disturbed spots such
as fire lanes and parts of chainings where bulldozer tracks had dug deeply.
Thus, it appears that better seedbed preparation is needed than was
afforded by the treatments used in this study.
It may be more effective
to burn, aerially seed as soon as the ashes are cool, and, since burning
in oakbrush is usually patchy, pipe harrow those spots where burning is
most severe.
This would plant seed as well as remove burned snags from
the created openings.

�-353-

5.

Spraying produced the most available spring, summer and fall elk
forage of the 3 methods tested after a period of 2 years.
Two years
after spraying the level of available forage production compared to
a pre-treatment value of 1.00 was 1:2.09 for Spring, 1:1.72 for
Summer and 1:1.94 for Fall. Chaining, however, produced the most
winter forage with a ratio of 1:1.24.
Spraying was second in production of available spring, summer, and fall forage.
Burning was lowest
in available elk forage during all seasons.
Five years after treatment available elk forage produced during all
seasons showed net declines on all treated areas compared with pretreatment levels.
Still, some treatments supported a little more
than others.
Production of available elk forage on sprayed areas
was slightly higher than that produced on chainings and burns during
all seasons.
Production on chainings and burns was about the same
during all seasons of the year. Seasonal 5th year pre-to post-treatment forage production ratios ranged from 1:0.72 to 1:0.85 on sprayed
areas, 1:0.60, to 1:0.75 on chainings, and from 1:0.60 to 1:0.74 on
burns.

6.

Spraying produced the most available spring and summer deer forage of
the 3 methods tested after 2 years (ratios of 1:1.40 and 1:1.20
respectively).
The most winter and fall forage was produced by chaining
(ratios of 1:1.23 and 1:1.26, respectively).
Spraying was second in
production of available spring and summer forage after 2 years.
Burning
was lowest in available deer forage during all seasons.
Five years after treatment available deer forage produced during all
seasons was below pre-treatment levels on all treated areas.
Still, some
treatments supported a little more than others.
Spraying and chaining
were comparable in available deer forage production.
Both methods
produced more during all seasons than burning.
Seasonal 5th year pre-to
post-treatment forage production ratios ranged from 1:0.79 to 1:0.98 on
sprayed areas, 1:0.76 to 1:0.98 on chainings, and 1:0.52 to 1:0.78 on
burns.

7.

After 2 years spraying produced by far the most summer cattle forage of
the 3 methods tested with chaining second and burning third.
Burning
resulted in a slight net decrease compared with pre-treatment levels
after 2 years.
Five years after treatment available cattle forage was below pre-treatment
levels on all treated areas, and pre-to post-treatment ratios for all
treatments were relatively close (spray-1:0.84, burn-l:0.76, chain 1:0.70).

8.

Burned
showed
copper
during

and chained areas when compared with control and sprayed areas
increased levels of total cell contents,
soluble carbohydrates,
and zinc in most plant categories (forbs, grasses and shrubs)
the 2nd summer and 3rd winter.

�-354-

9.

Elk showed distinct preference for burned areas after 2 and 5 years,
with sprayed areas ranking second and chainings third. Elk use on
burns increased 45 and 54 percent, respectively, after 2 and 5 years.
Deer did not appear to exhibit preference for any particular treatment
after 2, 3 or 5 years. Cattle showed 6 times more preference for burned
areas after 2 years than for sprayed or chained areas. No preference
was shown by cattle for sprayed or chained areas after 2 years, or for
any of the 3 treatments after 6 years. Preference of elk and cattle
for burns occurred despite decreases in forage production on burns as
compared with higher levels produced on sprayed areas and chainings.
Attractiveness of burns to elk and cattle may have been related to an
increase in those nutrients and minerals previously discussed or to
changes in cover. The overall effect of burning was creation of openings
interspersed with patches of standing oak vegetation which may have been
more suitable for cover and shade.
LITERATURE

Ju1ander, Odell.
1955. Determining
Range Manage. 8(4):182.

CITED
grazing use by cow-chip counts.

J.

Kufe1d, Roland C. 1970. Experimental improvement of oakbrush on deer,
elk and cattle ranges - Hightower Mountain.
Colo. Div. Game, Fish
and Parks.
Game Research Rept. Proj. W-I0I-R-12, WP 4, J3. July,
1970, Part 1, pp. 113-126.
Kufe1d, Roland C. 1971. Experimental improvement of oakbrush on deer, elk
and cattle ranges - Hightower Mountain.
Colo. Div. Game, Fish and
Parks.
Game Res. Rept. Proj. W-I0I-R-13, WP4, J3. July, 1971, Part
1. pp. 23-86.
Kufe1d, Roland C. 1972. Experimental improvement of oakbrush on deer, elk
and cattle ranges - Hightower Mountain.
Colo. Div. Wildlife.
Game
Research Rept. Proj. W-I0I-R-14, WP4, J3. July, 1972, Part 2. pp. 81-97.
Kufe1d, Roland C. 1973. Foods eaten by the Rocky Mountain
Manage. 26(2):106-113.

elk.

J. Range

Kufe1d, Roland C. 1975. Experimental improvement of oakbrush on deer, elk
and cattle ranges - Hightower Mountain.
Colo. Div. Wildlife.
Game
Res. Rept. Proj. W-I0I-R-17, WP4, J3. July, 1975, Part 1. pp. 25-92.
Kufe1d, Roland C., O. C. Wa11mo, and Charles Feddema.
1973. Foods of the
Rocky Mountain mule deer. USDA Forest Service, Rocky Mtn. For. and
Range Exp. Sta. Res. Pap. RM-lii.
31p.
Plummer, A. Perry, Donald R. Christensen, Richard Stevens and Kent R. Jorgensen.
1970. Intermediate wheatgrass, smooth brome and other herbs open and
control Gambe1 oak thickets.
In Highlights, results and accomplishments
of game range restoration studies. Utah Div. of Fish and Game. Pub. No.
70-3. pp. 26-30.

�-355-

LITERATURE

CITED (Cont'd)

Patton, David R. 1974. Patch cutting increases deer and elk use of a
pine forest in Arizona.
J. For. 72(12):764-766.
Pearson, Henry. A. 1968. Thinning, clearcutting and reseeding affect
deer and elk use of ponderosa pine forests in Arizona.
U.S. Forest
Ser., Rocky Mtn. For. and Range Exp. Sta. Res. Note RM-119, 4p.
Reynolds, Hudson, G. 1962a. Use of natural openings inaponderosa
pine
forest in Arizona by deer, elk and cattle. U.S. Forest Service,
Rocky Mtn. For. and Range Exp. Sta. Res. Note No. 78, 4p.
Reynolds, Hudson, G. 1962b. Effect of logging on understory vegetation
and deer use in a ponderosa pine forest in Arizona by deer, elk and
cattle.
U.S. For. Ser., Rocky Mtn. For. and Range Exp. Sta. Res.
Note No. 80. 7p.
Reynolds, Hudson, G, 1966a. Slash cleanup in a ponderosa pine forest
affects use by deer and cattle. U.S. For. Serv., Rocky Mtn. For.
and Range Exp. Sta. Res. Note RM-64, 3p.
Reynolds, Hudson, G. 1966b. Use of openings in spruce-fir forests of
Arizona by elk, deer and cattle. U.S. For. Ser., Rocky Mtn. For.
and Range Exp. Sta. Res. Note RM-66, 4p.
Wallmo, O. C. 1969. Response of deer to alternate strip clearcutting
of lodgepole pine and spruce-fir timber in Colorado.
U.S. For. Ser.,
Rocky Mtn. For. and Range Exp. Sta. Res. Note RM-141, 4p.

Prepared by

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_
Roland C. Kufeld
Wildlife Researcher

�-356-

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AFTER I AFTER
AFTER
AFTER
AFTER: AFTER
2 YEARS 15 YEARS 2 YEARS 15 YEARS 2 YEARSI 5 YEARS
BURNING

SPRAYING

CHAINING

Fig. 1. Percentage changes in air dry forage production recorded within
forb, grass and shrub categories 2 and 5 years after burning, spraying
and chaining.

�-357-

One year before burning

(July, 1970).

One year after burning

(July, 1972).

Fig. 2. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1, 2, and 5 years after burning.
(Fig. continued on next page).

�-358-

Two years after burning (July, 1973).

Five years after burning (July, 1976).
Fig. 2. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1, 2, and 5 years after burning.

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One year before spraying (July, 1970).

One year after spraying (July, 1972).
Fig. 3. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1, 2 and 5 years after spraying.
(Fig. continued on next page).

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Two years after spraying (July, 1973).

Five years after spraying (July, 1976).
Fig. 3. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1, 2 and 5 years after spraying.

�-361-

One year before chaining (July, 1970).

One year after chaining (July, 1972).
Fig. 4. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1, 2 and 5 years after chaining.
(Fig. continued on next page).

�-362-

Two years after chaining (July 1973).

Five years after chaining (July 1976)

Fig. 4. Vegetation conditions at a permanent photo point on the Hightower
Mountain oak study area 1 year before and 1, 2 and 5 years after chaining.

�-363-

Fig. 5. Section of one burn where Agropyron intermedium and Bromus inermis
began to become established two years after burning (July 1973).

Fig. 6. Most effective burn obtained in burned units. Vegetation was almost
completely burned with just a few dead snags left standing.

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Fig. 7. Least effective burn obtained in burned units. Only part of the
understory was removed with little or no damage to larger oaks.

Fig. 8. The overall result of burning was creation of openings interspersed
with patches of standing oak vegetation suitable for escape cover and shade.

�Table 1.

Changes in air dry forage production recorded within forb, grass and shrub categories 2 and 5 years after burning, spraying and chaining.

Time After
Treatment

Plant
Type

2 years

Forbs

5 years

Change Duell
To Burninglbs/ac Percent

Change Due
To SEraying
lbsl ac Percent

Change Due
To Chaining
lbsl ac Percent

+ 63.47

+ 14.49

+ 78.51

+ 14.96

+107.53 + 23.34

Grasses - 16.48

- 5.24

+658.05

+295.33

+ 64.60 + 16.70

Shrubs

-305.36

- 38.18

+ 34.83

+

4.28

+166.83 + 18.54

TOTALS

-257.37

- 16.59

+780.48

+ 50.27

+340.08 + 19.89

Forbs

+

+

2.27

- 16.22

-

-

- 29.99

- 43.36

- 15.78

6.81

Grasses -116.31

4.51

0.67 -

Confidence Level Where Treatment Difference is significan~1
Control
and Burn

Control
and Spray

Control
and Chain

.10
.05

Burn
and Spray

Burn
and Chain

.10
.15

.05

Spray
and Chain

.15
.05

.20

0.23

-226.62 - 47.50

I

V1
I

w

cr-

Shrubs

-268.97

- 37.84

-100.58

- 13.90

- 98.48 - 12.31

TOTALS

-368.44

- 26.53

-192.16

- 13.82

-288.97 - 18.87

.10

.15

.15

1/

Derived through comparison of treated and control areas with adjustments for normal year to year changes in forage production recorded on
the control areas.

l/Indicates the highest confidence level at which treatment differences are significant.
at the n.05, .10, .15, .20 or .25 level are shown.

Only treatment differences that are significant

�Table 2. Changes in production of air dry summer vegetation recorded on the control areas 2 and 5 years following burning, spraying
and chaining.

Plant Species

Achillea lanulosa
Agastache urticifolia
Agoseris glauca
Allium acuminatum
Androsace septentrionalis
Aquilegia coerulea
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia btennis
Artemisia dracunculus
Artemisia ludoviciana
Aster-Erigeron
Astragalus haydenianus
Balsamorhiza sagittata
Berberis repens
Calochortus gunnisonii
Capsella bursa-pastoris
Castilleja linariaefolia
Chenopodium berlandieri
Cirsium centaureae
Collomia linearis
Commandra umbellata
Cymopterus purpureus
Delphinium nelsoni
Delphinium occidentale
Descurainia sophia
Disporum trachycarpum

Percent ComEosition
1976
1973
1970

1.41
0.26
0.00
0.02
0.01
0.00
0.01
0.01
0.09
0.00
0.00
0.03
3.66
0.00
0.00
0.01
0.07
0.00
0.07
0.04
0.43
0.00
0.10
0.01
0.00
0.40
0.00
0.00

3.26
0.26
0.04
0.13
0.00
0.00
0.00
0.00
0.15
0.00
0.00
0.02
2.82
0.00
0.00
0.02
0.09
0.00
0.06
0.04
0.59
0.01
0.17
0.02
0.00
0.65
0.01
0.00

2.88
0.19
0.00
0.04
0.00
0.00
0.02
0.00
0.25
0.00
0.00
0.03
2.68
0.00
0.00
0.04
0.00
0.00
0.01
0.04
0.36
0.10
0.27
0.00
0.00
0.27
0.13
0.00

Lbs of Vegetation
Produced Eer Acre
1976
1973
1970
Forbs
24.15
4.52
0.00
0.30
0.18
0.00
0.18
0.18
1.49
0.00
0.00
0.42
63.64
0.00
0.00
0.18
1.19
0.00
1.31
0.71
7.26
0.00
1.84
0.18
0.00
6.90
0.00
0.00

52.28
4.16
0.59
1.90
0.00
0.00
0.00
0.06
2.20
0.00
0.00
0.36
45.50
0.00
0.00
0.24
1.43
0.00
1.01
0.59
9.58
0.12
2.80
0.24
0.00
10.65
0.12
0.00

42.59
2.74
0.00
0.54
0.00
0.00
0.30
0.00
3.39
0.00
0.00
0.36
38.96
0.00
0.00
0.59
0.00
0.00
0.18
0.48
4.94
1.37
3.98
0.00
0.00
3.98
2.08
0.00

Change
/
1970-19731
percent~/
1bs/ac

+28.13
-0.36
+0.59
+1.61
-0.18
0.00
-0.18
-0.12
+0.71
0.00
.0.00
-0.06
-18.14
0.00
0.00
+0.06
+0.24
0.00
-0.30
-0.12
+2.32
+0.12
+0.95
+0.06
0.00
+3.75
+0.12
0.00

+116.50
-7.89

+48.00

-28.50

+20.00
-22.73
+31. 97
+51.61
+54.31

Change
1970-1976
percent
1bs/ac

+18.44
-1. 78
0.00
+0.24
-0.18
0.00
+0.12
-0.18
+1.90
0.00
0.00
-0.06
-24.68
0.00
0.00
+0.42
-1.19
0.00
-1.13
-0.24
-2.32
+1.37
+2.14
-0.18
0.00
-2.91
+2.08
0.00

+76.35
-39.47

+128.00

-------------------------------------------------------------------------------------------------------------------------------------------

I

w
c-.

'"

I

-38.79

-31.07
+116.13
-42.24

�Table 2. Changes in production
and chaining.
(Cont'd).

Plant Species

Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium
Gentiana heterosepala
Geranium
Gilia aggregata
Grinde.Iia aphanactis
Hackelia leptophylla
Helemium hoopesii
Hydrophyllum capitatum
Hydrophyllum fendleri
Iris missouriensis
Lactuca scariola
Lappula redowskii
Lathyrus-Vicia
Lepidium densiflorum
Ligusticum
Linum lewisii
Lomatium leptocarpum
Lupinus argenteus
Madia glomerata
Medicago sativa
Melilotus offic.in"a.lis
Mertensia franciscana
Mertensia lanceolata
Moldavica parviflora

of air dry summer vegetation

recorded

Percent Composition
1970
1973
1976

Lbs of Vegetation
Produced per Acre
1970
1973
1976

0.02
0.00
0.00
1.08
0.01
0.37
0.00
0.21
0.05
0.30
0.00
0.12
0.00
0.00
0.00
7.32
0.00
1.57
0.20
0.00
7.58
0.23
0.00
0.00
0.49
0.00
0.00

0.07
0.04
0.00
1.01
0.02
0.18
0.01
0.28
0.00
0.12
0.01
0.10
0.00
0.00
0.00
6.78
0.01
1.29
0.19
0.00
4.63
0.01
0.00
0.00
0.48
0.00
0.00

Forbs
0.19
0.10
0.00
1.01
0.01
0.15
0.01
0.50
0.00
0.09
0.01
0.03
0.00
'0.00
0.00
4.63
0.00
1.04
0.12
0.00
3.08
0.28
0.00
0.00
0.05
0.00
0.02

(CoIlt'd).
0.42
0.00
0.00
18.62
0.12
6.30
0.00
3.45
0.83
4.94
0.00
2.02
0.00
0.00
0.00
124.43
0.00
27.12
3.63
0.00
131.50
3.75
0.00
0.00
7.97
0.00
0.00

on the control

1.13
0.65
0.00
16.12
0.30
2.85
0.12
4.22
0.00
1. 78
0.12
1.49
0.00
0.00
0.00
107.83
0.12
20.82
3.33
0.00
75.83
0.18
0.00
0.00
7.14
0.00
0.00

2.97
1.31
0.00
14.39
0.12
2.20
0.18
6.36
0.00
1.13
0.18
0.42
0.00
0.00
0.00
65.84
0.00
14.63
1.84
0.00
45.26
3.57
0.00
0.00
0.59
0.00
0.36

areas 2 and 5 years following

Change
1/
1970-1973 2/
1bs/ac
percen~

+0.71
+0.65
0.00
-2.50
+0.18
-3.45
+0.12
+0.77
-0.83
-3.15
+0.12
-0.54
0.00
0.00
0.00
-16.59
-0.12
-6.30
-0.30
0.00
-55.67
-3.57
0.00
0.00
-0.83
0.00
0.00

-13.42
-54.72
+22.41
-63.86
-26.47

-13.34
-23.25
-8.20
-42.33
-95.24

-10.45

burning,

spraying

Change
1970-1976
1bs/ac
percent

+2.56
+1.31
0.00
-4.22
0.00
-4.10
+0.18
+2.91
-0.83
-3.81
+0.18
-1. 61
0.00
0.00
0.00
-58.59
0.00
-12.49
-1. 78
0.00
-86.24
-0.18
0.00
0.00
-7.38
0.00
+0.36

-22.68
-65.09
+84.48
I

-77.11

W

Cl'

"I

-47.08
-46.05
-49.18
-65.58
-4.76

------------------------------------------------------------------------------------------------------------------------------------------

�Table 2. Changes in production of air dry summer vegetation recorded on the control areas 2 and 5 years following burning, spraying
and chaining. (Cont'd).

Plant Species

Oenothera caespitosa
Opuntia
Orthocarpus luteus
Osmorhiza
Pachistima myrsinites
pedicularis grayi
Penstemon crandallii
Penstemon strictus
Phacelia heterophylla
phlox longifolia
Polemonium foliosissimum
polygonum aviculare
polygonum douglasii
potentilla pulcherrima
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Senecio serra
Silene menziesii
Smilacina stellata
Solidago sparsiflora
Stellaria
Swertia radiata
Taraxacum officinale
Thalictrum fendleri

Percent COIDEosition
1976
1973
1970

0.00
0.00
0.00
0.95
0.28
0.00
0.19
0.03
0.00
0.00
0.00
0.00
0.00
0.21
0.00
0.00
0.12
0.00
0.00
0.00
0.00
0.06
0.11
0.80
0.00
0.17
1.95

0.00
0.00
0.00
0.87
0.09
0.02
0.04
0.02
0.00
0.00
0.04
0.00
0.00
0.27
0.00
0.00
0.05
0.00
0.01
0.06
0.00
0.05
0.05
1.73
0.00
0.44
1.59

Forbs
0.00
0.01
0.00
0.45
0.29
0.06
0.11
0.08
0.00
0.04
0.01
0.00
0.07
0.25
0.00
0.00
0.01
0.00
0.01
0.00
0.01
0.03
0.00
0.68
0.01
0.28
0.74

Lbs of Vegetation
Produced Eer Acre
1976
1973
1970

Change 1/
1970-197~
percentY
1bs/ac

Change
1970-1976
percent
1bs/ac

(Cont'd).
0.00
0.00
0.06
0.00
0.00
0.00
13.74
16.06
1.31
4.88
0.30
0.00
0.65
3.09
0.30
0.54
0.00
0.00
0.00
0.00
0.54
0.06
0.00
0.00
0.06
0.00
4.10
3.45
0.00
0.00
0.00
0.00
0.71
2.02
0.00
0.00
0.12
0.00
0.89
0.00
0.00
0.00
1.01
0.83
1.90
0.83
27.30
13.44
0.06
0.00
6.66
2.85
24.92
32.71

0.00
+0.06
0.00
-2.32
-3.57
+0.30
-2.44
-0.24
0.00
0.00
+0.48
0.00
+0.06
+0.65
0.00
0.00
-1.31
0.00
+0.12
+0.89
0.00
-0.18
-1.07
+13.86
+0.06
+3.81
-7.79

0.00
+0.18
0.00
-9.75
-0.77
+0.71
-1.61
+0.59
0.00
+0.65
+0.06
0.00
+0.89
0.00
0.00
0.00
-1.84
0.00
+0.12
0.00
+0.12
-0.48
-1.90
-3.87
+0.12
+0.89
-22.66

0.00
0.18
0.00
6.30
4.10
0.71
1.49
1.13
0.00
0.65
0.12
0.00
0.89
3.45
0.00
0.00
0.18
0.00
0.12
0.00
0.12
0.54
0.00
9.58
0.12
3.75
10.05

-14.44
-73.17
-78.85

+18.97
-64.71

+103.10
+133.83
+23.82

-60.74
-15.85
-51.92
I

w

c00
I

.
-28.76
+31.25
-69.27

------------------------------------------------------------------------------------------------------------------------------------------

�Table 2. Changes in production
and chaining.
(Cont'd).

Plant

Species

of air dry summer vegetation

recorded

Percent
1970

Lbs of Vegetation
Produced per Acre
1970
1973
1976

Composition
1973
1976

Forbs
Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
veronica americana
veronica biloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii
Wyethia amplexicaulis

on the control

areas 2 and 5 years following

burning,

spraying

Change
1/
1970-19732/
1bs/ac
percen~

Change
1970-1976
1bs/ac
percent

+7.69
+82.85
+106.12
-28.95

(Cont'd).

0.00
0.00
0.08
0.14
0.00
0.00
0.22
0.06
0.18
0.14

0.02
0.00
0.08
0.16
0.00
0.00
0.22
0.14
0.37
0.14

0.00
0.00
0.01
0.22
0.00
0.00
0.28
0.14
0.45
0.12

0.00
0.00
1.25
2.32
0.00
0.00
3.87
1.01
2.91
2.26

0.30
0.00
1.25
2.44
0.06
0.00
3.39
2.02
5.65
2.14

0.00
0.00
0.18
2.91
0.00
0.00
4.16
1.84
6.01
1.61

+0.30
0.00
0.00
+0.12
+0.06
0.00
-0.48
+1.01
+2.74
-0.12

-12.31
+100.00
+93.88
-5.26

0.00
0.00
-1.07
+0.59
0.00
0.00
+0.30
+0.83
+3.09
-0.65

Total .Forbs

31.86

30.03

22.97

545.35

478.50

328.02

-66.85

-12.26

-217.33

-39.85

Agropyron
Agropyron cristatum
Agropyron intermedium
Bromus anomalus
Bromus inermis
Bromus tectorum
Carex
Dactylus glomerata
Elymus glaucus
Festuca ovina
F'estuca thurberi

4.85
0.00
0.00
2.15
0.01
0.04
0.00
0.00
1.00
0.00
0.00

2.17
0.00
0.00
1.31
0.23
0.00
0.00
0.00
0.28
0.00
0.02

0.25
0.00
0.00
1.28
0.00
0.00
0.00
0.00
0.38
0.00
0.09

Grasses
84.52
0.00
0.00
36.58
0.18
0.59
0.00
0.00
16.18
0.00
0.06

35.27
0.00
0.00
20.40
3.81
0.00
0.00
0.00
4.10
0.00
0.36

3.63
0.00
0.00
18.14
0.00
0.00
0.00
0.00
4.94
0.00
1.13

-49.25
0.00
0.00
-16.18
+3.63
-0.59
0.00
0.00
-12.07
0.00
+0.30

-58.27

-80.89
0.00
0.00
-18.44
-0.18
-0.59
0.00
0.00
-11. 24
0.00
+1.07

-95.71

+5.13

+25.64

I

w
0'&gt;
xo
I

-44.23

-74.63

-50.41

-69.49

------------------------------------------------------------------------------------------------------------------------------------------

�Table 2. Changes in production of air dry summer vegetation recorded on the control areas 2 and 5 years following burning. spraying
and chaining. (Cont'd).

Plant Species

Percent ComEosition
1970
1973
1976

Lbs of Vegetation
Produced Eer Acre
1970
1973
1976

Change 1/
1970-197J2/
1bs/ac
percenr-

Change
1970-1976
1bs/ac
percent

Grasses (Cont'd).
Hordeum brachyantherum
Juncus balticus
Koeleria cristata
Orzyopsis hymenoides
Phleum pratense
Poa-Carex geyeri
Sitanion hystrix
Stipa

0.25
0.01
0.01
0.00
0.00
12.74
0.00
0.97

1.00
0.00
0.05
0.00
0.00
13.52
0.00
0.91

Total Grasses

22.04

19.49

0.39
0.00
0.02
0.00
0.00
23.74
0.01
0.60

4.10
0.12
0.24
0.00
0.00
221.61
0.00
16.06

14.87
0.00
0.89
0.00
0.00
218.64
0.00
14.45

5.00
0.00
0.24
0.00
0.00
344.08
0.12
8.56

26.76

380.24

312.79

385.83

+10.77
-0.12
+0.65
0.00
0.00
-2.97
0.00
-1.61

+262.32

-1.34
-10.00

+0.89
-0.12
0.00
0.00
0.00
+122.46
+0.12
-7.49

+21. 74

+55.26
-46.67
I

w

-67.45

-17.74

+5.59

+1.47

"

0
I

Shrubs
Acer glabrum
Amelanchier alnifolia
Artemisia cana
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
populus tremuloides
Prunus v.irginiana
Purshia tridentata

0.00
7.27
0.06
0.40
0.06
0.00
0.02
0.12
0.00
0.00
5.94
0.11

0.00
7.13
0.03
0.53
0.11
0.00
0.01
0.22
0.00
0.00
5.58
0.20

0.00
9.72
0.03
0.91
0.11
0.00
0.00
0.12
0.00
0.00
8.83
0.15

0.00
123.95
1.07
7.20
0.95
0.00
0.30
2.08
0.00
0.00
103.91
1.78

0.00
115.39
0.48
9.16
1.67
0.00
0.12
3.75
0.00
0.00
92.67
2.91

0.00
139.71
0.54
14.57
1.37
0.00
0.00
1.72
0.00
0.00
132.46
1.90

0.00
-8.56
-0.59
+1.96
+0.71
0.00
-0.18
+1. 67
0.00
0.00
-11.24
+1.13

-6.91
+27.27

+80.00
-10.82
+63.33

0.00
+15.76
-0.54
+7.38
+0.42
0.00
-0.30
-0.36
0.00
0.00
+28.55
+0.12

+12.72
+102.48

-17.14
+27.48
+6.67

------------------------------------------------------------------------------------------------------------------------------------------

�Table 2. Changes in production of air dry summer vegetation recorded on the control areas 2 and 5 years following burning, spraying
and chaining. (Cont'd).

Plant Species

Percent ComEosition
1976
1973
1970
Shrubs

Quercus gambellii
Ribes
Rosa nutkana
Symphoricarpos albus

Total Shrubs
TOTAL ALL PLANTS

17.92
0.43
1.35
12.44

18.75
0.40
2.01
15.51

12.86
0.74
1.85
14.94

Lbs of Vegetation
Produced Eer Acre
1973
1976
1970

Change
/
1970-197~
2/
percenr
1bs/ac

Change
1970-1976
percent
1bs/ac

(Cont'd).
308.57
6.96
23.43
2l3.35

302.44
6.l3
32.00
248.62

184.50
9.64
26.59
211.58

-6.l3
-0.83
+8.56
+35.27

-1.99
-11.97
+36.55
+16.53

-124.07
+2.68
+3.15
-1.67

-40.21
+38.46
+13.45
-0.78

---

---

---

---

---

---

46.10

50.48

50.27

815.32

724.67
---

+21.77

+2.74

-68.87

-8.68

---

1606.61

1438.52

-112.53

-6.55

-280.61

-16.32

---

---

---

793.55
---

100.00

100.00

100.00

1719.14

I
VJ

•....
"
I

1/ Reflects normal year to year changes in forage production recorded on the control areas.
-2/ Percent changes are only shown for the more abundant plants.

�Table 3.

Changes in production of air dry summer vegetation
Percent composition
1/
E 73 -

recorded on the burned areas two years following burning.
Lbs of vegetation
produced per acre

A

Plant species

1970

1973

1970

1973

E 73

1bs/ac

111.82
0.30
0.95
8.80
0.00
0.00
0.00
0.00
4.28
0.00
0.18
3.81
105.33
0.00
0.48
0.00
1.07
0.06
1.67
8.92
8.15
0.24
0.30
2.02
0.06
0.00
0.36
0.00

92.84
0.05
21.23
18.27
0.00
0.00
0.03
0.10
4.40
0.00
1. 78
2.96
77 .18
0.00
0.24
0.00
1.28
0.59
4.32
0.20
22.84
0.12
1.80
1.19
0.01
0.00
1.42
0.00

+ 18.97
+ 0.24
- 20.27
- 9.47
0.00
0.00
- 0.03
- 0.10
- 0.12
0.00
- 1.61
+ 0.85
+ 28.15
0.00
+ 0.24
0.00
- 0.21
- 0.54
- 2.65
+ 8.72
- 14.69
+ 0.12
- 1.51
+ 0.83
+ 0.05
0.00
- 1.06
0.00

Change due to burning ~
Confidence level
percent ~
where significant 4/

Forbs
AcJUlle.a £.anU£o.6a
Aga..6taehe. ~ei6otia
Ag 0.6 e.m 9 £.auea
Alliwn aewn,[natwn
AndJr.0.6«ce .6 e.pte.nW.o na.ti.6
Aqu,[,te.g,[a eo~U£e.a
A'W.b'[;" dJr.wnmoncU
Altab'[;" rUM uta
Alte.nan,[a eonge..6ta
A~e.m,[;.,,[a b,[e.nn,[;.,
A~e.m,[;.,,[a dlr.cteuneU£U.6
~e.m,[;.,,[a ludov,[eiana.
Mt~-EJUg~on
Mbtaga.£.U6 hayde.n,[anU.6
Ba£..6a.moiliza .6ag,[ttata.
B~bV1M
lte.pe.11.6
Ca.£.oeho~U.6 9unn.i.-~0 nil
Cap.6e.lia buMa-pa..6tom
Ca..6~e.ja
tinan,[ae.6o£.,[a
Che.nopocUwn be.Jt£.ancUru
CiJt.6,[wn ee.i1ta.Ulte.ae.
CoUom,[a tine.alli
Commandlr.ct wnbe.liata.
Cymopt~U.6 pUltpUlte.U6
Ve.£.ph,[n,[wn nwon,[
Ve.£.ph,[n,[wn oeeide.nta..e.e.
Ve..6eUlta,[n,[a .6oph,[a
V,[;.,POltwn .tJtaehyea.Jtpwn

2.57
0.00
0.02
0.17
0.00
0.00
0.03
0.02
0.18
0.00
0.11
0.21
6.50
0.00
0.01
0.00
0.06
0.04
0.34
0.01
1.05
0.00
0.07
0.05
0.00
0.00
0.01
0.00

8.66
0.02
0.07
0.68
0.00
0.00
0.00
0.00
0.33
0.00
0.01
0.30
8.14
0.00
0.04
0.00
0.08
0.00
0.13
0.69
0.63
0.02
0.02
0.16
0.00
0.00
0.03
0.00

5.96

5.00

42.88
0.06
0.36
2.85
0.00
0.00
0.48
0.30
2.97
0.00
1. 78
3.45
107.95
0.00
0.24
0.00
1.07
0.59
5.59
0.24
17.31
0.00
1.19
0.89
0.00
0.00
0.12
0.00

+ 20.44
- 51.82

- 2.70

.10

I

w
-.J

- 90.00
+ 28.74
+ 36.48

- 16.67
- 61.45
- 64.33
- 83.51

tv
I

�Table 3. Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 73 l!

1970

1973

E 73

lbs/ac

2.38
2.44
0.06
15.05
0.00
1.78
0.00
2.74
0.00
0.18
0.00
0.12
7.43
0.00
0.00
36.40
0.00
1.13
2.32
0.00
78.51
0.00
0.30
0.00
0.59
0.00
0.12

16.47
11.67
0.12
9.89
0.00
0.70
0.00
10.05
0.00
0.37
0.00
0.09
18.85
0.83
0.00
31.65
0.00
1.51
8.19
0.00
77.58
0.00
0.01
0.00
0.01
0.00
0.01

- 14.09
- 9.24
- 0.06
+ 5.16
0.00
+ 1.08
0.00
- 7.31
0.00
- 0.19
0.00
- 0.03
- 11.42
- 0.83
0.00
+ 4.75
0.00
- 0.38
- 5.87
0.00
+ 0.93
0.00
+ 0.29
0.00
+ 0.59
0.00
+ 0.11

Change due to burning ~
Confidence level
percent 11 where significant 11

Forbs (contd)
EJUogonwn wnbe,Ua,twn
Ep~ob~wn pa~culatum
F~agaJUa am~cana
GaUwn
Ge.ntiana heX~o-6e.pa1.a
GMaMwn
Gilia
agg~e.ga.ta.
GJUnd~a aphana~
Hack~a
te.ptophylfa
Heie.Mwn hoopuU
Hy~ophyttwn
cap~wn
Hy~ophyttwn
6e.ndt~
I W mU,-60 uJUe.n-6,u,
Lactu.c.a -6caJUota
Lapputa ~e.dOW6
W
Lathy~U6 - V~ua
Le.p~~wn de.n-6~6to~wn
Ug u.-6licwn
Unwn lw,u,U
Lomaliwn te.ptoc~pwn
Lu.p~nu.-6~g e.nte.U6
Madia gtom~a
Me.dicago -6aliva
M~otU6
o6Munat,{J.,
M~e.H-6~a 6Mnw cana
M~e.n-6~a tance.ota.ta.
Motdav~ca p~v~6toM

0.37
0.01
0.01
0.68
0.00
0.09
0.00
0.50
0.00
0.06
0.00
0.01
1.12
0.05
0.00
2.20
0.00
0.12
0.54
0.00
8.13
0.00
0.00
0.00
0.00
0.00
0.00

0.18
0.19
0.00
1.17
0.00
0.14
0.00
0.21
0.00
0.01
0.00
0.01
0.58
0.00
0.00
2.81
0.00
0.09
0.18
0.00
6.04
0.00
0.02
0.00
0.05
0.00
0.01

2.04

4.98

6.07
0.18
0.12
11.42
0.00
1.55
0.00
8.21
0.00
1.01
0.00
0.12
18.85
0.83
0.00
36.52
0.00
1.96
8.92
0.00
134.54
0.00
0.00
0.00
0.00
0.00
0.00

- 85.55
+ 52.19

+154.81
- 72.77
I

w

"I

W

- 60.57
+ 15.01

- 24.99
- 71.68
+

1.20

.25

�Table 3. Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
Qroduced Qer acre

Percent comQosition
Plant species

1970

1973

E 73 11

1970

1973

E 73

1bs/ac

0.00
2.14
0.00
0.18
0.00
0.00
2.80
1.31
0.00
4.22
0.06
0.06
0.00
5.95
0.00
0.48
1.49
0.30
1.19
0.18
0.00
0.06
1.43
28.01
0.00
23.02
2.02

0.00
33.61
0.30
1.53
0.00
0.00
0.86
1.59
0.30
0.01
0.09
1.78
7.43
7.78
0.00
0.12
1.03
0.01
0.12
0.89
0.00
0.01
1.82
26.33
0.00
38.72
2.54

0.00
- 31.47
0.30
1.35
- 0.00
0.00
+ 1.94
- 0.28
- 0.30
+ 4.21
- 0.03
- 1.72
7.43
- 1.84
0.00
+ 0.36
+ 0.46
+ 0.29
+ 1.07
- 0.71
0.00
+ 0.05
- 0.39
+ 1.68
0.00
- 15.70
- 0.52

Change due to burning ~
Confidence level
percent 11 where significant 11

Forbs (contd )
Oeno~h~a
ca~p~o4a
Opun.tia
OJdhOC.MPU6 .tuteU6
04mollJUza
Pac.{IMtima mYM-&lt;-~e4
PecUc.u1.aJW, 9tr.a.y-&lt;Pen4~emon CJtanda.tW
Pen4~emon 4:tJUc.;tu6
Phac.e.t,[a h~~ophy.t.ta
Ph.tox .tong-&lt;-6o.t-&lt;-a
Po.temo Mum 60.t,[04-&lt;-M-&lt;-mum
Po.tygonum av-&lt;-c.~e
Po.tygonum doug.ta4~
Po~enXi.Ua
pu.£c.heNUma
Ranunc.u.£U6 -&lt;-namoenU6
Rumex CI!M pU6
Rudbec.kia mo~ana
Senec.-&lt;-o amb~4-&lt;-o-&lt;-d~
Senec.-&lt;-o -&lt;-~egeJVUmU6
S en ecio 4 eJlJl.a
SUene menz-&lt;-~~
SmUac.-&lt;-na 4~e.t.tam
So.t,[dago 4 PaM-&lt;-6,totr.a.
S~e.t.tM-&lt;-a
Sw~a
tr.a.cUam
Tatr.a.xac.um o66-&lt;-c.-&lt;-na.te
ThaLi..ctJtum 6endbuu

0.00
0.34
0.02
0.11
0.00
0.00
0.25
0.17
0.02
0.00
0.00
0.11
0.07
0.39
0.00
0.01
0.17
0.00
0.00
0.00
0.00
0.00
0.25
0.78
0.00
0.99
0.20

0.00
0.16
0.00
0.01
0.00
0.00
0.21
0.10
0.00
0.32
0.00
0.00
0.00
0.46
0.00
0.04
0.12
0.02
0.09
0.01
0.00
0.00
0.11
2.16
0.00
1.79
0.16

0.00
5.65
0.30
1.78
0.00
0.00
4.04
2.85
0.30
0.00
0.00
1.78
1.25
6.54
0.00
0.12
2.91
0.00
0.00
0.00
0.00
0.00
4.16
12.97
0.00
16.59
3.33

-

- 93.63

+226.74
- 17.50

.05

I

w
'-J

'"
I

- 23.58
+ 44.56

- 21.63
+ 6.38
- 40.55
- 20.30

�Table 3.

Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
Eroduced Eer acre

Percent comEosition
Plant species

T~agopogon dub~um
T~agopogon p~aten6~
Uilic.a MO~c.a
Va1.vUana oc.udenta.-eM
V~OMc.a ainvUc.ana
V~OMc.a bUoba
V~g~CVta. mu1ti6.to~
V.{.ofu ~ugu.t06a
V~o.e.a.!.&gt;he.UonU
Wye;t~a ainpie:Uc.a~

Total Forbs

1970

0.02
0.00
0.00
0.02
0.00
0.02
0.34
0.04
0.27
0.14
30.09

1973

E 73 .lI

I

1970

Forbs (contd)
0.30
0.06
0.00
0.36
0.00
0.30
5.59
0.71
4.46
2.32

0.06
0.00
0.00
0.03
0.01
0.06
0.15
0.39
0.30
0.25
38.76

499.31

1973

E 73

lbs/ac

0.83
0.06
0.00
0.36
0.18
0.77
1.96
5.12
3.87
3.21

8.84
0.06
0.00
0.38
0.06
0.30
4.90
1.43
8.65
2.20

- 8.01
-

+

-

+

-

+

0.00
0.00
0.02
0.12
0.48
2.94
3.69
4.78
1.01

Change due to burni ng Y
._
Confidence level
percent }j where significant 11

- 59.97
- 55.30
+ 46.15

--

--

501.57

438.10

+ 63.47

+ 14.49

4.22
0.00
0.00
0.60
0.00
0.26
0.10
0.00
0.00
0.00
0.50

2.34
0.00
0.93
0.41
0.22
0.06
0.00
0.02
0.11
0.00
0.00

1.89

70.01
0.00
0.00
9.99
0.00
4.34
1.61
0.00
0.00
0.00
8.33

30.27
0.06
12.07
5.35
2.91
0.83
0.00
0.30
1.43
0.00
0.00

29.21
0.01
0.01
5.57
0.21
0.07
1.61
0.01
0.00
0.00
49.96

1.06
0.05
+ 12.06
0.22
+ 2.70
+ 0.76
- 1.61
+ 0.29
+ 1.42
0.00
- 49.96

+ 3.63

"

V1
I

Grasses
Ag~opYMn
Ag~opy~on ~tatum
Ag~opy~on ~nt~eMum
B~omU!.&gt;anoma1.U6
&amp;0 mU!.&gt; ~n e.J1J10.,
BMmU!.&gt;tecXMum
CMex
VacXyiU!.&gt; giom~a.ta
EiymU!.&gt;gfuUC.U6
FeAtuc.a ov~na
FeAtuc.a th~bvU

I

w

+

+

-

- 3.95

�Table 3. Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
Qroduced Qer acre

Percent comQosition
Plant species

Holtdewn bJW..c.hyan.:the!twn
] UYlC.U6 bal;Uc.U6
Koe.teiUa CJl1J.,:tata
Oltzyo P.6-u' hymeno-&lt;'dv.,
Phi.ewn p!ta:te1t6 e
Poa-Caltex geyeiU
S-&lt;-:tarUon hy;.,:tJt-&lt;-x
SUpa

1970

0.18
0.11
0.28
0.00
0.02
15.65
0.19
0.89

1973

E 73 11

1973

E 73

1bs/ac

0.03
0.13
0.09
0.00
0.03
17.72
0.03
0.89

Grasses {contd}
3.03
0.36
1.90
1.67
1.13
4.64
0.00
0.00
0.36
0.42
16.60
259.98
229.23
3.15
0.36
14.87
11.54

10.99
0.16
17.40
0.00
0.36
256.49
3.15
13.38

- 10.63
+ 1.51
- 16.27
0.00
+ 0.06
- 27.26
2.80
- 1.84
-

1970

. 2/
Change due to burnlng Confidence level
percent Y where significant 11

- 96.75
- 93.50

.15

- 10.63
- 13.78
I

w

Total Grasses

23.01

23.03

382.20

297.92

314.41

- 16.48

0.00
41. 57
10.53
12.55
0.12
1.07
15.23
2.74
0.71
0.00
9.87
0.00

0.00
52.82
4.23
165.93
0.62
0.01
0.40
27.30
0.12
0.00
10.61
0.00

0.00
- 11. 25
+ 6.30
-153.38
- 0.51
+ 1.06
+ 14.82
- 24.56
+ 0.59
0.00
- 0.74
0.00

- 5.24

Shrubs
Ac.e!t g.tabltwn
Amelanc.h-&lt;.e!t a.trU6oUa
A!t:temi6 -&lt;'0.,c.ana
A!t:temi6-&lt;.a :tJt-&lt;-den:ta:ta
Ce!tC.OC.altpU6 mon:tanU6
Chlty;., othamnsu. deplte6;" tv..
ChltY-60:thamnU6 natv.. eos tv..
Chlty;., othamniu, v-u, ud16.tolttv..
C!ta:taegU6 e!ty:thltopoda
POpulU6 :t!temulo-&lt;.dv.,
PltunU6 v-i.!tg-&lt;.rUana
PUItJ.,lua fuden:tata.

0.00
3.45
0.58
7.90
0.02
0.00
0.06
0.92
0.01
0.00
0.71
0.00

0.00
3.19
0.81
0.96
0.01
0.08
1.17
0.21
0.06
0.00
0.77
0.00

3.39

0.00
56.74
9.52
130.37
0.36
0.00
1.01
15.17
0.12
0.00
11.90

0.00

- 21.29
+148.91
- 92.44

- 89.98

- 6.93

"

0I

�Table 3.

Changes in production of air dry summer vegetation recorded on the burned areas two years following burning (contd).
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 73 JI

1970

1973

E 73

lbs/ac

Change due to burning ~
Confidence level
percent ~
where significant if

Shrubs (contd}
QuVtQU!.l gambe..ttU
TUbe!.&gt;
RO-6a nuti&lt;.ana
Sympho~Qa4po-6
albU!.l

Total Shrubs
TOTAL ALL PLANTS

10.56
2.41
1.74
18.54

14.70
1.10
1.38
13.78

46.90

38.21

--

--

100.00

100.00

11.06

2.60
23.11

174.98
40.39
29.20
308.63

190.15
14.10
17.84
177.90

171. 51
35.55
39.88
359.65

+ 18.64
- 21. 46
- 22.03
-181 .75

+ 10.87
- 60.35
- 55.25
- 50.54

.10

778.38

494.38

799.73

-305.36

- 38.18

.05

1659.90

1293.87

1551.24

-257.37

- 16.59

I
W

"
"
I

JlReflects

the amount of forage one would expect to find on a treated area following treatment had treatment not occurred.

~Derived through comparison of burned and control areas with adjustments for normal year to year changes in forage production recorded on
the control areas.
~Percent

changes are only shown for the more abundant plants.

if Indicates the highest confidence level at which changes due to burning are significant.
a .05, .10, .15, .20 or .25 level are shown.

Only changes that are significant at the

�Table 4.

Changes

in production

of air dry summer vegetation

Percent
Plant Species

Com~osition

1970

1976

~ 7611

2.57
0.00
0.02
0.17
0.00
0.00
0.03
0.02
0.18
0.00
0.11
0.21
6.50
0.00
0.01
0.00
0.06
0.04
0.34
0.01
1.05
0.00
0.07
0.05
0.00
0.00
0.01
0.00

5.28
0.01
0.01
0.02
0.00
0.00
0.00
0.02
0.26
0.00
0.00
0.08
7.95
0.02
0.04
0.00
0.00
0.59
0.09
0.16
1.11
0.02
0.10
0.00
0.00
0.00
0.01
0.00

5.27

recorded

on the burned areas five years following

Lbs of Vegetation
Produced per Acre
A

1970

1976

E 76

Lbslac

54.30
0.12
0.06
0.18
0.00
0.00
0.00
0.24
2.50
0.00
0.00
0.89
81.84
0.18
0.36
0.00
0.00
6.42
0.83
1.55
11.12
0.18
0.95
0.00
0.00
0.00
0.12
0.00

75.63
0.04
0.36
5.14
0.00
0.01
0.79
0.02
6.78
0.01
1.78
2.96
66.08
0.01
0.24
0.03
0.01
0.59
0.76
0.16
11.78
1.37
2.57
0.05
0.01
0.01
24.76
0.01

- 21.32
+ 0.08
- 0.30
- 4.96
0.00
- 0.01
- 0.79
+ 0.22
- 4.28
- 0.01
- 1.78
- 2.06
+ 15.76
+ 0.17
+ 0.12
- 0.03
- 0.01
+ 5.83
+ 0.07
+ 1.39
- 0.65
- 1.19
- 1.62
- 0.05
- 0.01
- 0.01
- 24.64
- 0.01

burning.

Change Due to Burnin~~
Confidence Level
Percent11
Where Significant ~I

FORBS
Achillea lanulosa
Agastache urticifolia
Agoseris glauca
Allium acuminatum
Androsace septentrionalis
Aquilegia coerulea
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia biennis
Artemisia dracunculus
Artemisia ludoviciana
Aster-Erigeron
Astragalus haydenianus
Balsamorhiza sagittata
Berberis repens
Calochortus gunnisonii
Capsella bursa-pastoris
Castilleja linariaefolia
Chenopodium berlandieri
Cirsium centaureae
Collomia linearis
Comandra umbel lata
Cymopterus purpureus
Delphinium nelsoni
Delphinium occidentale
Descurainia sophia
Disporum trachycarpum

4.75

42.88
0.06
0.36
2.85
0.00
0.00
0.48
0.30
2.97
0.00
1.78
3.45
107.95
0.00
0.24
0.00
1.07
0.59
5.59
0.24
17.31
0.00
1.19
0.89
0.00
0.00
0.12
0.00

- 28.20

- 63.16

,
co,

...,

w

+ 23.85

-

5.54

-----------------------------------------------------------------------------------------------------------------------------------------------

�Table 4.

Changes

in production

of air dry summer vegetation

Species

Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium
Gentiana heterosepala
Geranium
Gilia aggregata
Grindelia aphanactis
Hackelia leptophylla
Helenium hoopesii
Hydrophyllum captitatum
Hydrophyllum fendleri
Iris missouriensis
Lactuca scariola
Lappula redowskii
Lathyrus-Vicia
Lepidium densiflorum
Ligusticum
Linum lewisii
Lomatium leptocarpum
Lupinus argenteus
Madia glomerata
Medicago sativa
Melilotus officinalis
Mertensia franciscana
Mertensia lanceolata
Moldavica parviflora

1970

1976

0.37
0.01
0.01
0.68
0.00
0.09
0.00
0.50
0.00
0.06
0.00
0.01
1.12
0.05
0.00
2.20
0.00
0.12
0.54
0.00
8.13
0.00
0.00
0.00
0.00
0.00
0.00

0.10
0.38
0.00
0.70
0.00
0.06
0.00
1.26
0.01
0.00
0.00
0.00
0.71
0.10
0.00
1.21
0.00
0.06
0.08
0.00
4.77
0.00
0.09
0.00
0.00
0.00
0.01

on the burned areas five years following

Lbs of Vegetation
Produced Eer Acre

Percent ComEosition
Plant

recorded

~ 7J-./

Forbs Cont' d
6. 07

1.39

3.30

0.18
0.12
11.42
0.00
1.55
0.00
8.21
0.00
1.01
0.00
0.12
18.85
0.83
0.00
36.52
0.00
1.96
8.92
0.00
l34.54
0.00
0.00
0.00
0.00
0.00
0.00

(continued).

. 2/

A

1970

burning

1976

E 76

Lbs/ac

1.01
4.16
0.00
7.38
0.00
0.65
0.00
l3.50
0.12
0.00
0.00
0.00
7.79
1.07
0.00
12.13
0.00
0.59
0.77
0.00
47.05
0.00
0.83
0.00
0.00
0.00
0.06

43.33
23.35
0.12
8.83
0.01
0.54
0.18
15.14
0.00
0.23
0.18
0.02
18.85
0.83
0.01
19.32
0.01
1.06
4.53
0.01
46.31
0.01
0.01
0.00
0.00
0.01
0.36

- 42.32
- 19.18
- 0.12
- 1.45
- 0.01
+ 0.11
- 0.18
1.64
+ 0.12
- 0.23
- 0.18
- 0.02
- 11.06
+ 0.24
- 0.01
- 7.19
- 0.01
- 0.46
- 3.76
- 0.01
+ 0.74
- 0.01
+ 0.82
0.00
0.00
- 0.01
- 0.30

-

Change Due to Burnln~_
Confidence Level
Percent }j
Where Significant

i/

- 97.67
- 16.47

- 10.84
I

w

so
"
I

- 58.68
- 37.21

+

1.60

--------------------------------------------------------------------------------~.--------------------------------------------------------------

�Table 4.

Changes

in production

of air dry summer vegetation

Percent

COIDEosition

Plant Species

1970

1976

Oenothera.caespitosa
Opuntia
Orthocarpus luteus
Osmorhiza
Pachistima myrsinites
Pedicularis grayi
Penstemon crandallii
Penstemon strictus
Phacelia heterophylla
Phlox longifolia
Polemonium foliosissimum
Polygonum aviculare
Polygonum douglasii
pulcherrima
Potentilla
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Senecio serra
Silene menziesii
Smilacina stellata
Solidago sparsiflora
Stellaria
Swertia radiata
Taraxacum officinale
Thalictrum fendleri

0.00
0.34
0.02
0.11
0.00
0.00
0.25
0.17
0.02
0.00
0.00
0.11
0.07
0.39
0.00
0.01
0.17
0.00
0.00
0.00
0.00
0.00
0.25
0.78
0.00
0.99
0.20

0.00
0.38
0.01
0.03
0.00
0.00
0.11
0.19
0.00
0.29
0.00
0.08
0.49
0.53
0.00
0.00
0.27
0.00
0.00
0.00
0.00
0.00
0.00
0.18
0.00
1.02
0.05

~ 7J:-/

recorded

on the burned

areas five years

Lbs of Vegetation
Production Per Acre

following

Change

1976

E 76

Lbs/ac

0.01
100.82
0.30
0.70
0.01
0.71
1.94
6.03
0.30
0.65
0.02
1.78
111.43
6.54
0.01
0.12
0.26
0.01
0.12
0.01
0.12
0.01
0.02
9.24
0.12
21.78
1.02

- 0.01
- 97.25
- 0.18
- 0.34
- 0.01
- 0.71
- 0.93
- 4.24
- 0.30
+ 2.14
- 0.02
- 0.95
-106.32
- 0.95
- 0.01
- 0.12
+ 2.72
- 0.01
- 0.12
- 0.01
- 0.12
- 0.01
- 0.02
- 7.51
- 0.12
- 10.96
- 0.49

(continued).

2/

Due to Burning Confidence Level
Percent1/
Where Significant

A

1970

burning

i/

Forbs Con' t

0.00
5.65
0.30
1.78
0.00
0.00
4.04
2.85
0.30
0.00
0.00
1.78
1.25
6.54
0.00
0.12
2.91
0.00
0.00
0.00
0.00
0.00
4.16
12.97
0.00
16.59
3.33

0.00
3.57
0.12
0.36
0.00
0.00
1.01
1.78
0.00
2.80
0.00
0.83
5.12
5.59
0.00
0.00
2.97
0.00
0.00
0.00
0.00
0.00
0.00
1.72
0.00
10.82
0.54

- 96.46

.10

- 48.00
- 70.39

.20
w

oo

.0
I

- 95.41
- 14.55
+1056.46

.25

- 81.33

.05

- 50.30

--------------------------------------------------------------------------------------------------------------------------------------------.
.

�Table 4.

Changes

in production

of air dry summer vegetation

Percent

ComEosition

recorded

on the burned

areas

five years

Lbs of Vegetation
Produced Eer Acre

Species

1970

1976

~ 7J-/

1970

1976

burning

(continued).

2/
Due to BurningConfidence Level
Percent1/
Where Significant ~/

Change

A

Plant

following

E 76

Lbs/ac

0.30
0.06
0.00
0.45
0.01
0.30
6.02
1.30
9.19
1.65

+

Forbs Cont'd
Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
veronica americana
veronica biloba
Vigiuera multiflora
Viola rugulosa
Viola shel tonii
Wyethia amplexicaulis
Total Forbs

1.01
0.00
0.00
0.36
0.00
0.00
4.58
0.89
1. 78
2.32

+

0.71
0.06
0.00
0.09
0.01
0.30
1.44
0.41
7.41
0.67

300.33

+

6.81

3.01
0.01
0.01
4.96
0.00
0.07
1.61
0.01
0.00
0.00
158.21
3.69

+ 39.28
+ 0.35
+ 8.85
+ 19.13
+ 6.54
+ 0.05
+ 17.90
+ 2.73
+ 0.12
0.00
-156.72
- 2.32

0.10
0.00
0.00
0.04
0.00
0.00
0.44
0.09
0.19
0.24

0.30
0.06
0.00
0.36
0.00
0.30
5.59
0.71
4.46
2.32

--

--

--

--

--

30.09

30.02

499.31

307.14

0.02
0.00
0.00
0.02
0.00
0.02
0.34
0.04
0.27
0.14

-

-

-

-

-

-

- 23.94

00

- 80.59
+ 40.74

--+

2.27

GRASSES
Agropyron
Agropyron cristatum
Agropyron intermedium
Bromus anomalus
Bromus intermis
Bromus tectorum
Carex
Dactylus glomerata
Elymus glaucus
Festuca ovina
Festuca thurberi
Hordeum brachyantherum

4.22
0.00
0.00
0.60
0.00
0.26
0.10
0.00
0.00
0.00
0.50
0.18

4.14
0.04
0.85
2.30
0.69
0.01
1.77
0.28
0.01
0.00
0.14
0.12

70.01
0.00
0.00
9.99
0.00
4.34
1. 61
0.00
0.00
0.00
8.33
3.03

42.29
0.36
8.86
24.09
6.54
0.12
19.51
2.74
0.12
0.00
1. 49
1. 37

I

w

+1307.21

+386.09

+1114.81

- 99.06
- 62.96

-----------------------------------------------------------------------------------------------------------------------------------------------

•....
I

�Table 4.

Changes in production of air dry summer vegetation recorded on the burned areas five years following burning (continued).

Percent Composition
Plant Species

1970

1976

i 761/

Lbs of Vegetation
Produced per Acre
A

1976

1970

E 76

Lbs/ac

Change Due to Burning 2/
Confidence Level
Percent1/
Where Significant i/

Grasses Cont'd
Juncus balticus
Koeleria cristata
Orzyopsis hymenoides
Phleum pratense
Poa-Carex geyeri
Sitanion hystrix
Stipa

Total Grasses

---

1.90
4.64
0.00
0.36
259.98
3.15
14.87
---

0.00
0.36
0.00
1.43
148.28
0.59
13.38
---

0.16
4.64
0.01
0.36
403.64
37.50
7.93
---

26.36

382.20

271.51

387.82

0.11
0.28
0.00
0.02
15.65
0.19
0.89

0.00
0.04
0.00
0.13
14.49
0.06
1.30

-23.01

29.15

- 0.16

- 4.28
- 0.01
+ 1.07
-255.36
- 36.90
+ 5.45

+ 68.75

-116.31

- 29.99

- 63.26

I
W
OJ
N

I

SHRUBS
Acer glabrum
Amelanchier alnifolia
Artemisia can a
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depress us
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
Populus tremuloides
Prunus virginiana
Purshia tridentata

0.00
3.45
0.58
7.90
0.02
0.00
0.06
0.92
0.01
0.00
0.71
0.00

0.00
5.54
1.84
1.67
0.01
0.12
0.16
1.00
0.02
0.00
1.22
0.01

4.62

0.00
56.74
9.52
130.37
0.36
0.00
1.01
15.17
0.12
0.00
11.90
0.00

0.00
52.16
18.08
15.76
0.06
1.13
1.61
9.81
0.24
0.00
13.26
0.06

0.01
63.96
4.76
263.98
0.51
0.01
0.03
12.57
0.12
0.01
15.16
0.01

- 0.01
- 11.79
+ 13.32
-248.22
- 0.45
- 1.12
+ 1.57
- 2.75
+ 0.12
- 0.01
- 1.90
+ 0.05

- 18.44
+280.00
- 94.03

.25

- 21.91
- 12.53

-----------------------------------------------------------------------------------------------------------------------------------------------

�Table 4.

Changes

in production

of air dry summer vegetation

Percent
Plant

Species

1970

recorded

Lbs of Vegetation
Produced Eer Acre

ComEosition
1976

on the burned areas five years following

;;;76))

1970

1976

E 76

Lbs/ac

burning

(continued).

Change Due to Burning ~j
Confidence Level
Percent1/
Where Significant

i/

Shrubs Cont'd
Quercus gambellii
Ribes
Rosa nutkana
Symphoricarpos albus

10.56
2.41
1. 74
18.54

14.23
0.63
2.64
14.51

Total Shrubs

46.90

Total All Plants

100.00

174.98
40.39
29.20
308.63

144.89
6.90
28.25
149.65

104.63
55.92
33..13
306.22

+ 40.26

+ 38.48

- 49.02
- 4.88
-156.57

- 87.66
- 14.73
- 51.13

43.61

778.38

441.86

710.82

-268.97

- 37.84

100.00

1,659.90

1,020.51

1,388.95

-368.44

- 26.53

7.59
2.39
22.26

.10
I

w
(1J

w
I

!/ Reflects

the amount of forage one would expect to find on a treated

l/ Derived

through comparison
on the control areas.

3/
- Percent

and control

areas with adjustments

changes are only shown for the more abundant

i/ Indicates
a .05, .10,

of burned

area after treatment
for normal

had treatment

not occurred.

year to year changes

in forage production

recorded

plants.

the highest confidence level at which changes
.15, .20 or .25 level are shown.

due to spraying

are significant.

Only changes

that are significant

at the

�Table

5.

Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying.
Lbs of vegetation
eroduced eer acre

Percent comeosition
Plant species

1970

1973

E 7311

1970

Change due to sera~ing ~
Confidence level
where significant ~
percent '}j

1973

"E 73

1bs/ac

104.86
0.48
3.15
1.01
0.00
0.95
0.00
0.00
3.21
0.00
0.18
1.90
47.34
0.65
1. 13
0.06
1.19
0.00
0.83
0.00
29.86
9.99
1.78
0.24
0.06
20.58
2.26
0.12

55.63
0.16
3.54
3.05
0.12
0.01
0.00
0.02
0.35
0.00
7.61
0.56
43.93
0.01
0.01
0.24
1.93
0.00
3.40
0.10
19.78
0.12
0.02
0.01
0.01
27.26
2.12
0.18

+ 49.23
+ 0.31

+ 88.50

- 0.39
- 2.03

- 10.89
- 66.80

Forbs
AQhittea lanuto~a
AgM.taQhe uJr;t{.u6oUa
AgMeJ1M glauQa
AWum aQumincttum
AneVtMaQe ~epteVl.w..on~
Aquilegia QoeJtutea
AJta.b~ eVtummoncU.
AJta.b~ IUMu;ta
~enania
Qong~ta
AtLtem~ia bien~
AtLtem~ia eVtaQunQut~
AtLtem~ia ludoviQiana
MteJt - Eug eJton
M:tJtagal~ haydenian~
B~amotr.hiza ~agi.t.ta.ta
BeJtbew f!.epe~
CaloQhotLt~ gun~oili
CapuUa bUMa-pMtoW
CMtitteja
Unaniae60lia
ChenopocU.um bVtlancU.Vti
C~ium
QentaUlteae
CoUomia UneaW
ComMeVta umbeUctta
CymopteJt~ pUltpUlte~
Velphinium nWoru.
Velphinium oQudentale
V~QUltainia Mphia
V~potr.um .tJta.QhyQaltpum

1.56
0.01
0.00
0.03
0.13
0.00
0.00
0.00
0.01
0.00
0.46
0.04
3.70
0.00
0.00
0.01
0.10
0.00
0.26
0.01
0.92
0.00
0.00
0.00
0.00
1.08
0.01
0.01

4.15
0.02
0.12
0.05
0.00
0.05
0.00
0.00
0.14
0.00
0.01
0.07
2.00
0.02
0.06
0.00
0.05
0.00
0.04
0.00
1.10
0.40
0.08
0.01
0.00
0.76
0.10
0.01

3.61

2.85

25.69
0.18
0.06
0.48
2.20
0.00
0.06
0.06
0.24
0.00
7.61
0.65
61.44
0.00
0.00
0.18
1.61
0.00
4.40
0.12
14.99
0.00
0.00
0.00
0.00
17.66
0.18
0.18

- 0.12

0.94
0.00
- 0.02
+ 2.86
0.00
- 7.43
+ 1.34
+ 3.42
+ 0.64
+ 1.12
- 0.18
- 0.74
0.00
- 2.57
- 0.10
+ 10.08
+ 9.87
+ 1.77
+ 0.22
+ 0.05
- 6.68
+ 0.14
- 0.06

+

I

.20

+

7.78

- 38.27

+ 50.95

- 24.50

w
0&gt;

.c-

�Table

5.

Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying (contd).
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 73 l!

1970

1973

E 73

0.19
0.00
0.00
59.12
0.30
3.87
0.00
1.84
0.59
0.00
1.84
3.33
0.00
0.06
1.90
34.68
0.77
8.74
1.25
0.00
67.39
0.00
0.00
0.06
4.76
0.95
0.00

0.81
0.00
0.18
20.55
0.59
2.80
3.54
1.38
0.00
0.02
0.12
0.52
0.00
0.01
0.48
87.37
0.12
13.56
2.46
0.00
104.34
0.03
0.00
0.01
6.18
0.01
0.00

lbs/ac

2/
Change due to s~raling Confidence level
percent '}j
where significant 11

Forbs (contd}
E~ogonum umbellatum
EpUob-&lt;-um paMc.u1atum
FlW.gaJU..a. ameJUc.ana
GaUum
Gent,{.ana het~o4epala
G~aMum
GU,{.a aggJtegata
aphana~
G~ndet,{.a
Hac.ketia £eptophy£la
He£eMUll1 IWOPMU
HydJLophy£lum c.ap,{.tatum
HydJLophy£lum 6endfeJU
I Jt.U, m-&lt;.440 wUeM-&lt;.4
Lac.tuc.a 4 c.~o£a
Lappufa Jtedow,~W
LathyJtU6 - V-&lt;-Ua
Lep,lMWYI deM-&lt;-6£oJtum
UgMtiC.um
L,i,num £ew~u
Lomat-&lt;-um £eptoc.aftpum
Lup,lnU6 aftg enteM
MaMa g£om~ata
MeMc.ago 4ativa
Me£UotU6 o6Mc.bt~
MeJtteM-&lt;-a 6lW.l1w c.ana
MeJtten6-&lt;-a £anc.eo£ata
Moldav-&lt;-c.a paftv-&lt;-6£olW.

0.02
0.00
0.01
1.44
0.01
0.38
0.02
0.07
0.01
0.00
0.00
0.04
0.00
0.00
0.03
6.08
0.00
1.06
0.16
0.00
10.85
0.04
0.00
0.00
0.41
0.00
0.00

0.06
0.00
0.00
2.35
0.02
0.15
0.00
0.07
0.02
0.00
0.07
0.17
0.00
0.00
0.07
1.40
0.03
0.37
0.06
0.00
3.16
0.00
0.00
0.00
0.24
0.04
0.00

1.35

5.63
0.88
6.64

0.30
0.00
0.18
23.73
0.24
6.19
0.30
1.13
0.12
0.06
0.00
0.71
0.00
0.00
0.48
100.81
0.00
17.66
2.68
0.00
180.93
0.59
0.00
0.00
6.90
0.00
0.00

+

0.38
0.00
- 0.18
+ 38.57
- 0.30
+ 1.06
- 3.54
+ 0.46
+ 0.59
- 0.02
+ 1.72
+ 2.81
0.00
+ 0.05
+ 1.43
- 52.69
+ 0.65
- 4.82
- 1.21
0.00
- 36.95
- 0.03
0.00
+ 0.05
- 1.42
0.94
0.00

+187.73

.15

+ 38.02
+ 33.28

I

w

00
\J1

I

- 60.31

.10

- 35.52
- 49.17

.15

- 35.41

- 22.99

�Table 5 .

Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying (contd).
Lbs of vegetation
eroduced eer acre

Percent comeosition
Plant species

1970

1973

E 73 .l!

A

1970

1973

E 73

lbs/ac

0.00
0.00
0.00
8.33
3.15
0.00
0.48
1.13
0.00
0.36
3.21
0.36
0.18
2.68
0.00
0.00
2.91
0.24
0.06
1.37
0.00
0.54
2.80
40.80
4.16
45.56
21.65

0.00
0.00
0.00
3.92
1.42
68.98
0.31
1.75
0.00
0.01
2.68
0.01
1.77
12.03
0.06
0.00
0.59
0.36
0.12
0.89
0.00
0.15
0.73
25.49
17.33
15.68
23.34

0.00
0.00
0.00
4.41
1.73
- 68.98
+ 0.16
- 0.62
0.00
+ 0.35
+ 0.54
+ 0.35
- 1.59
- 9.35
- 0.06
0.00
+ 2.33
- 0.12
- 0.06
+ 0.48
0.00
+ 0.39
+ 2.07
+ 15.31
- 13.17
+ 29.88
- 1.69

Change due to serating Y
onfidence level
percent 11
where significant 4/

Forbs (contd)
Oeno~h~o eae4p~o~a
Opurz.:Ua.
OfLthoea!1.pU6 .e.ute~
O~moJrlUza
PaefUJ.,:U.ma mYM.i..~e4
PecUeu.£~
gJta.y.i..
Pe~~emon eJta.ndatt£.&lt;..
Pe~~emon .6Wd~
Phaeelia
h~~ophytta
Phlox long.i..6oUa
Polemon.i..um 6olio~.i..M-£mum
Polygonum av.i..eu.£a!1.e
Polygonum doug£.M«
Po~e~a
pu.£eh~a
Ranuneu.£U6 b!amo enU6
Rumex CAMp~
Rudbee~a
mo~ana
Senee.i..o amb~o~.i..o.i..de4
Senee.i..o .i..~eg~U6
Senee.i..o ~~
Silene
menue4.i...i..
Sm.i...tae.i..na~~e.t.t~
Solidago
~paM.i..6loJta.
S~eUa!1..i..a
Sw~a
~acUMa
Ta!1.axaeum o66.i..e.i..na.te
Tha.Lt~um
6end.t~

0.00
0.00
0.00
0.28
0.31
0.14
0.09
0.19
0.00
0.00
0.02
0.00
0.02
0.62
0.00
0.00
0.10
0.02
0.00
0.00
0.00
0.01
0.10
0.76
0.18
0.41
1.85

0.00
0.00
0.00
0.32
0.16
0.00
0.02
0.04
0.00
0.02
0.13
0.01
0.01
0.10
0.00
0.00
0.12
0.01
0.00
0.07
0.00
0.03
0.14
1.65
0.16
1.70
0.86

1.51

0.00
0.00
0.00
4.58
5.29
2.32
1.49
3.15
0.00
0.00
0.30
0.00
0.30
10.11
0.06
0.00
1.67
0.36
0.00
0.00
0.00
0.18
1.67
12.55
2.91
6.72
30.63

+112.51
+121.96
- 35.47

I

w

00
0I

- 77.75
+395.83

+283.67
+ 60.08
- 75.98
+190.52
- 7.22

.15

�Table

5.

Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying (contd).
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 73 l!

1970

2/
Change due to s~ra~ing Confi dence level
percent 11
where signi fi cant !!I

1973

E 73

lbs/ac

- 27.23
0.00
+ 5.12
+ 5.58
0.00
0.00
+ 0.01
- 2.20
- 2.55
- 9.15

- 19.89
- 32.49
- 76.21

Forbs (contd)
dub~um
Tnagopogon
TfUlgopogon pJtate.IU,u,
UJt;tlc.a diocea
Va1.vU.ana oc.ude.n;t:.aLU,
Ve.no~c.a am~c.ana
VeJ1.o~c.a biloba
V~gu,Le.J1.amuUL6,toJ1.a
V~oR.a nuguR.o.6a
V~oR.a -6he.UonM.
Wye.:t1ua ampR.e.uc.auR.i6

Total Forbs

0.06
0.00
0.08
0.50
0.00
0.00
0.00
0.33
0.25
0.77

0.04
0.00
0.33
0.64
0.00
0.00
0.00
0.38
0.22
0.11

0.95
0.00
1.43
8.27
0.00
0.00
0.06
5.53
4.04
12.67

1.07
0.00
6.54
14.27
0.00
0.00
0.06
8.86
5.29
2.85

28.30
0.00
1.43
8.69
0.00
0.00
0.05
11.06
7.84
12.00

--

--

--- ---

---

36.07

24.82

598.22

603.40

524.89

+ 78.51

+ 14.96

47.52
0.00
0.48
32.53
0.00
5.71
0.00
9.69
28.97
0.00
0.00

5.98
0.00
0.01
4.31
0.00
0.00
0.00
1.61
1.74
0.00
0.00

+ 4].54
0.00
+ 0.47
+ 28.22
0.00
+ 5.71
0.00
+ 8.09
+ 27.23
0.00
0.00

+694.45

+358.33
+ 64.24
.05
I

w

.15

Grasses
AgnopyJ1.on
AgnoptjJwlt cJlM:tcttum
Agnopynon ~nte.~e.cUum
BnomU.6 anoma1.U.6
BnomU.6 ~lteJ1.InL.6
BfLOmU.6te.ctonum
CMe.X

VactyR.U6 gR.omvw.;ta
ER.ymU.6gR.auc.U.6
Futuc.a
ov~na
Futuc.a
thMb~

0.86
0.00
0.00
0.47
0.00
0.00
0.00
0.10
0.41
0.00
0.00

2.06
0.00
0.02
1.33
0.00
0.25
0.00
0.36
1.39
0.00
0.00

14.33
0.00
0.00
7.73
0.00
0.00
0.00
1.61
6.84
0.00
0.00

+654.44
.05
+503.70
+]569.36

00

"I

�Table 5.

Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying (contd).
Lbs of vegetation
Eroduced Eer acre

Percent comEosition
Plant species

1970

1973

E 73 11

1973

E 73

lbs/ac

0.00
0.00
4.94
0.00
2.91
719.50
0.00
28.61

0.00
0.00
3.12
0.00
0.36
230.73
0.06
4.71

0.00
0.00
+ 1.81
0.00
+ 2.56
+488.77
- 0.06
+ 23.90

1970

Change due to sEra~ing ~
Confidence level 4/
percent '}j
where significant -

Grasses (contd)
Ho~deum b~achyanth~um

J unCU6 baLtic.UI.l
Koetvu.a c.tr.M:ta:ta
O~zyopf.,b., hymeno,[de-6
Phieum p~axen-6e
Poa-CMex geyvu.
S,[tal'uon hY-6Wx
Stipa

0.00
0.00
0.05
0.00
0.02
14.15
0.00
0.32

0.00
0.00
0.25
0.00
0.11
29.52
0.00
1.15

15.01

0.00
0.00
0.83
0.00
0.36
233.87
0.06
5.23

+211.84

.15

+507.32

I

co

00
00
I

Tota 1 Grasses

16.37

270.86

36.43

880.86

222.81

+658.05

0.00
115.33
0.00
65.13
0.00
0.59
1.84
9.16
0.00
0.42
100.87
0.00

0.24
126.63
0.00
54.88
0.00
0.65
0.00
14.13
0.00
0.01
89.48
0.00

- 0.24

+295.33

Shrubs
Ac~ giab~um
Ametanc.hl~
aini60Ua
M;temb.,,[a cana
AtLtemb.,,[a Wdentaxa
C~C.OCMpUl.l montanU6
Ch~I:fMthamnU6 dep~e-6f.,w.,
CMI:ff.,othamnu» naw.,ecs w.,
Ch~l:ff.,
othamnU6 vb., ud-&lt;-6iMU6
C~aegw.,
~ytMopoda
PopufU6 ~emufo'[de-6
P~unU6 v~g,[niana
PuM hla Wdentaxa

0.01
8.13
0.00
2.56
0.00
0.04
0.00
0.48
0.00
0.00
6.10
0.00

0.00
5.44
0.00
3.34
0.00
0.03
0.09
0.37
0.00
0.02
3.96
0.00

7.98

5.73

0.24
136.03
0.00
43.12
0.00
0.65
0.00
7.85
0.00
0.00
100.34
0.00

- 11.30
0.00
+ 10.25
0.00
- 0.06
+ 1.84
- 4.97
0.00
+ 0.41
+ 11.39
0.00

- 8.92
+ 18.67

- 35.19
+ 12.73

.10

�Table

5. Changes in production of air dry summer vegetation recorded on the sprayed areas two years following spraying (contd).
Lbs of vegetation
produced per acre

Percent composition
Plant species

1970

1973

E 73 Jj

1970

Chanqe due to spraying ~
Confidence level
percent ~
where significant 11

1973

E 73

lbs/ac

202.16
6.42
9.28
337.53

202.46
9.84
31.67
306.49

- 0.30
- 3.42
- 22.40
+ 31.04

- 34.74
- 70.71
+ 10.13

813.91

+ 34.83

+

Shrubs (contd)
QUeJtC.CL6

gambe.UU

R.tbu
RO-6 a

nuA:kana
Symphoh.tc.~po-6 albU-6
Total Shrubs

12.33
0.66
1.40
15.85

9.71
0.33
0.43
15.02

-

--

47.56

38.75

12.90
2.08
19.75

206.57
11.18
23.20
263.01

--

--

792.18

848.74

--

--

- 0.15
.10

4.28

-TOTAL ALL PLANTS

100.00

100.00

1661.27

2333.00

1552.52

+780.48

I

+ 50.27

1/
- Reflects the amount of forage one would expect to find on a treated area after treatment had treatment not occurred.
2/

-- Derived through comparison of sprayed and control areas with adjustments for normal year to year changes in forage production recorded
on the control areas.
~Percent

changes are only shown for the more abundant plants.

~Indicates
the highest confidence level at which changes due to spraying are significant.
a .05, .10, .15, .20 or .25 level are shown.

Only changes that are significant at the

w

OJ
'!)
I

�Table

6. Changes in production

of air dry summer vegetation

Percent
Plant Species

1970

COIDEosition

1976

E7J./

recorded

on the sprayed

areas five years following

Lbs of Vegetation
Produced Per Acre
A

1970

1976

E76

lbs/ac

50.44
0.18
0.00
0.06
_0.00
0.00
0.06
0.00
3.87
0.00
0.00
2.26
59.77
0.00
0.00
0.24
0.00
0.00
1.67
0.89
11.48
4.10
0.00
0.00
0.00
6.36
1.55
0.18

45.31
0.11
0.06
0.86
0.12
0.01
0.10
0.00
0.54
0.01
7.61
0.56
37.61
0.01
0.01
0.59
0.01
0.01
0.60
0.08
10.20
1.37
0.02
0.00
0.01
10.20
37.14
0.18

+ 5.12
+ 0.07
- 0.06
- 0.80
- 0.12
- 0.01
- 0.04
- 0.00
+ 3.32
- 0.01
- 7.61
+ 1.70
+22.16
- 0.01
- 0.01
- 0.36
- 0.01
- 0.01
+ 1.07
+ 0.81
+ 1.28
+ 2.74
- 0.02
- 0.00
- 0.01
- 3.84
-35.60
0.00

spraying.

Change Due To SEra~ingl/
,,/ Confidence Level 4/
Where SignificantPercent~

Forbs
Achillea lanulosa
Agastache urticifolia
Agoseris glauca
Allium acuminatum
Androsace septentrional is
Aquilegia coerulea
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia biennis
Artemisia dracunculus
Artemisia ludoviciana
Aster-Erigeron
Astragalus haydenianus
Balsamorhiza sagittata
Berberis repens
Calochortus gunnisonii
Capsella bursa~pastoris
Castilleja linariaefolia
Chenopodium berlandieri
Cirsium centaureae
Collomia linearis
Commandra unbellata
Cymopterus purpureus
Delphinium nelsoni
Delphinium occidentale
Descurainia sophia
Disporun trachycarpum

1.56
0.01
0.00
0.03
0.13
0.00
0.00
0.00
0.01
0.00
0.46
0.04
3.70
0.00
0.00
0.01
0.10
0.00
0.26
0.01
0.92
0.00
0.00
0.00
0.00
1.08
0.01
0.01

4.24
0.02
0.00
0.00
0.00
0.00
0.00
0.00
0.32
0.00
0.00
0.19
4.95
0.00
0.00
0.02
0.00
0.00
0.14
0.08
0.97
0.35
0.00
0.00
0.00
0.54
0.13
0.01

3.19

2.71

25.69
0.18
0.06
0.48
2.20
0.00
0.06
0.06
0.24
0.00
7.61
0.65
61.44
0.00
0.00
0.18
1.61
0.00
4.40
0.12
14.99
0.00
0.00
0.00
0.00
17.66
0.18
0.18

+ 11.31

I

w

\0

0
I

+58.93

+177 .48
+12.57

-37.63

------------------------------------------------------------------------------------------------------------------------------------------

�Table 6.

Changes in production

of air dry summer vegetation

Percent COIDEosition
Species

1970

1976

Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium
Gentiana heterosepala
Geranium
Gilia aggregata
Grindelia aphanactis
Hackelia leptophylla
Helenium hoopesii
Hydrophyllum capitatum
Hydrophyllum fendleri
Iris missouriensis
Lactuca scariola
Lappula redowskii
Lathyrus-Vicia
Lepidium densiflorum
Ligusticum
Linum lewisii
Lomatium leptocarpum
Lupinus argenteus
Madi a glomerata
Medicago sativa
Melilotus officinalis
Mertensia franciscana
Mertensia lanceolata
Moldavica parviflora

0.02
0.00
0.01
1.44
0.01
0.38
0.02
0.07
0.01
0.00
0.00
0.04
0.00
0.00
0.03
6.08
0.00
1.06
0.16
0.00
10.85
0.04
0.00
0.00
0.41
0.00
0.00

0.03
0.01
0.01
1.99
0.00
0.24
0.10
0.05
0.02
0.00
0.00
0.08
0.00
0.00
0.00
2.24
0.00
1.24
0.l3
0.00
3.89
0.00
0.00
0.00
0.03
0.00
0.00

Plant

E76-V

recorded

on the sprayed

areas

five years following

Lbs of Vegetation
Produced Per Acre

Change

A

1970

1976

E76

0.42
0.06
0.12
23.73
0.06
2.91
1.19
0.65
0.18
0.00
0.00
0.95
0.00
0.00
0.00
26.71
0.00
14.81
1.61
0.00
46.93
0.00
0.00
0.00
0.42
0.00
0.00

2.12
1.31
0.18
18.35
0.24
2.16
5.31
2.08
0.00
0.01
0.18
0.15
0.01
0.01
0.48
53.35
0.01
9.53
1.36
0.01
62.27
0.57
0.00
0.00
0.51
0.01
0.36

lbs/ac

spraying.

(Continued).

2/

Due To SEralin~
Confidence Level 4/
3/
Pe r cen t+
Where Significant-

Forbs

1.36

3.85
0.70
4.40

0.30
0.00
0.18
23.73
0.24
6.19
0.30
1.l3
0.12
0.06
0.00
0.71
0.00
0.00
0.48
100.81
0.00
17.66
2.68
0.00
180.93
0.59
0.00
0.00
6.90
0.00
0.00

-

1.71
1.25
- 0.06
+ 5.38
- 0.18
+ 0.76
- 4.12
- 1.43
+ 0.18
- 0.01
- 0.18
+ 0.80
- 0.01
- 0.01
- 0.48
- 26.64
- 0.01
+ 5.28
+ 0.25
- 0.01
- 15.35
- 0.57
0.00
0.00
- 0.10
- 0.01
- 0.36

-

+ 29.34
+ 34.98

I

w

-o

•....
I

- 49.94

.20

+ 55.41
+ 18.06
- 24.64

- 19.14

----------------------------------------------------------------------------------------------------------------------------------------------

�~

Table

6.

Changes in production

of air dry summer

Percent
Plant

Species

Oenothera caespitosa
Opuntia
Orthocarpus luteus
Osmorhiza
Pachistima myrsinites
Pedicularis grayi
Penstemon crandallii
Penstemon strictus
Phacelia heterophylla
Phlox longifolia
Polemonium foliosissimum
Polygonum aviculare
Polygonum douglasii
Potentilla pulcherrima
Ranunculus inameneus
Rumex crispus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Senecio serra
Silene menziesii
Smilacina stellata
Solidago sparsiflora
Stellaria
Swertia radiata
Taraxacum officinale
Thalictrum fendleri

1970

0.00
0.00
0.00
0.28
0.31
0.14
0.09
0.19
0.00
0.00
0.02
0.00
0.02
0.62
0.00
0.00
0.10
0.02
0.00
0.00
0.00
0.01
0.10
0.76
0.18
0.41
1.85

vegetation

COID2osition
E7f}./
1976

0.00
0.00
0.26
0.29
0.25
0.00
0.00
0.06
0.00
0.05
0.04
0.06
0.00
0.25
0.00
0.00
0.14
0.05
0.02
0.00
0.00
0.00
0.00
0.49
0.31
0.76
1.00

recorded

on the sprayed

areas five years following

Lbs of Vegetation
Produced Per Acre
1970

1976

Forbs

(Cont'd)

0.00
0.00
0.00
4.58
5.29
2.32
1.49
3.15
0.00
0.00
0.30
0.00
0.30
10.11
0.06
0.00
1.67
0.36
0.00
0.00
0.00
0.18
1.67
12.55
2.91
6.72
30.63

0.00
0.00
3.09
3.45
3.03
0.00
0.00
0'.65
0.00
0.59
0.48
0.71
0.00
2.91
0.00
0.00
1.67
0.59
0.18
0.00
0.00
0.06
0.00
5.89
3.63
9.04
11.90

spraying.

(Continued).

Change

E76

lbs/ac

0.01
0.18
0.01
1.80
4.45
165.56
0.71
6.65
0.01
0.65
0.59
0.01
26.53
10.11
0.06
0.01
0.15
0.36
0.12
0.01
0.12
0.09
0.01
8.94
34.67
8.82
9.41

0.01
0.18
+ 3.08
+ 1.65
1.42
-165.56
- 0.71
- 6.00
- 0.01
- 0.06
- 0.12
+ 0.70
- 26.53
- 7.20
- 0.06
- 0.01
+ 1.52
+ 0.24
+ 0.06
- 0.01
- 0.12
- 0.03
- 0.01
- 3.05
- 31.04
+ 0.22
+ 2.48

Due To s2rayingI/
Confidence Level 4/
Where SignificantPercent~/

-

-

-

+

-

91.86
31. 90

I

w

\D
N

I

-

71.18

+1033.33

+
+

34.14
89.53
2.49
26.39

.10

--------------------------------------------------------------------------------------------------------------------------------------------

�Table 6. Changes in production of air dry summer vegetation recorded on the sprayed areas five years following spraying.

Percent ComEosition
Plant Species

1970

1976

Elf):i

Lbs of Vegetation
Produced Per Acre
1970

.

1976

E76

lbs/ac

(Continued).

2/
Change Due To SErayingConfidence Level 4/
3/
PercentWhere Significant-

Forbs (Continued)
Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
Veronica americana
Veronica biloba
Viguiera mul tiflora
Viola rugulosa
Viola shel tonii
Wyethia arnplexicaulis

Total Forbs

0.06
0.00
0.08
0.50
0.00
0.00
0.00
0.33
0.25
0.77

0.04
0.00
0.20
1.14
0.00
0.00
0.01
0.56
0.21
0.41

0.95
0.00
1.43
8.27
0.00
0.00
0.06
5.53
4.04
12.67

0.42
0.00
2.44
13.74
0.00
0.00
1.25
6.66
2.50
4.88

0.95
0.01
0.20
10.39
0.01
. 0.01
0.06
10.09
8.34
9.00

36.07

28.72

598.22

343.60

-

-

0.54
0.01
2.23
3.35
0.01
0.01
1.18
3.43
5.84
4.12

359.82

-

16.22

0.62
0.00
0.01
3.83
0.00
0.00
0.01
1.61
2.09
0.00
0.19

+

33.17
0.00
0.11
13.18
0.00
0.00
0.01
0.30
6.83
0.00
0.19

-

+
+

-

+

-

-

+

32.27

-

-

33.96
70.03
45.82

-

4.51

Grasses
Agropyron
Agropyron cristatum
Agropyron intermedium
Bromus anomalus
Bromus inermis
Brornus tectorurn
Carex
Dactylus glomerata
Elymus glaucus
Festuca ovina
Festuca thurberi

0.86
0.00
0.00
0.47
0.00
0.00
0.00
0.10
0.41
0.00
0.00

2.85
0.00
0.01
1.42
0.00
0.00
0.00
0.11
0.74
0.00
0.00

14.33
0.00
0.00
7.73
0.00
0.00
0.00
1.61
6.84
0.00
0.00

33.78
0.00
0.12
17.01
0.00
0.00
0.00
1.31
8.92
0.00
0.00

+
+

-

+

+ 343.61

- 18.52
+ 327.45

-----------------------------------------------------------------------------------------------------------------------------------------------

I

w

'-0
W
I

�Table 6.

Changes

in production

of air dry summer vegetation

1970

1976

E76!/
Grasses

Hordeum brachyantherum
Juncus balticus
Koeleria cristata
Orzyopsis hymenoides
Ph.leum pratense
Poa-Carex geyeri
Sitanion hystrix
Stipa

0.00
0.00
0.05
0.00
0.02
14.15
0.00
0.32

0.00
0.00
0.00
0.00
0.15
13.74
0.00
0.42

on the sprayed

areas five years following

Lbs of Vegetation
Produced Per Acre

Percent Composition
Plant Species

recorded

26.35

1976

E76

0.06
0.00
0.00
0.00
1.78
163.44
0.06
5.00

0.01
0.00
0.83
0.01
0.36
363.10
0.71
2.79

1970

lbs/ac

spraying.

(Continued).

. 2/
Change Due To_ S~~~y~
3/
Confidence Level 4/
PercentWhere Significant-

(Continued)

0.00
0.00
0.83
0.00
0.36
233.87
0.06
5.23

0.05
0.00
- 0.83
- 0.01
+ 1.43
-199.66
- 0.65
+ 2.20

+

- 54.99
+ 78.98
I

w
-.0
.I"-

Total Grasses

16.37

19.44

270.86

231.49

274.84

- 43.36

0.24
136.03
0.00
43.12
0.00
0.65
0.00
7.85
0.00
0.00
100.34
0.00

0.00
107.06
0.00
57.57
0.00
0.77
0.00
5.71
0.00
0.00
119.07
0.00

0.24
153.32
0.01
87.31
0.00
0.65
0.00
6.51
0.01
0.01
127.91
0.00

- 0.24
- 46.26
- 0.01
- 29.74
0.00
+ 0.12
0.00
- 0.80
- 0.01
- 0.01
- 8.83
- 0.00

- 15.78

Shrubs
Acer glabrum
Amelanchier alnifolia
Artemisia cana
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
Crataegus erythropoda
Populus tremuloides
Prunus virginiana
Purshia tridentata

0.01
8.13
0.00
2.56
0.00
0.04
0.00
0.48
0.00
0.00
6.10
0.00

0.00
8.88
0.00
4.73
0.00
0.06
0.00
0.48
0.00
0.00
10.06
0.00

10.87

9.08

- 30.17
- 34.06

- 12.23
6.91

--------------------------------------------------------------------------------------------------------------------------------------------

I

�Table 6.

Changes

in production

of air dry summer vegetation

Percent

recorded

on the sprayed

areas five years following

A

Plant Species

1970

1976

E76])

12.33
0.66
1.40
15.85

9.11
0.46
0.85
17.21

8.85

--

--

47.56

51.84

1970

1976

(Continued).

2/

Lbs of Vegetation
Produced Per Acre

ComEosition

spraying.

E76

lbs/ac

Change Due To S2rayin~
Confidence Level 4/
Percent1/
Where Significant-

Shrubs (Continued)
Quercus gambellii
Ribes
Rosa nutkana
Symphoricarpos albus

Total Shrubs

-

13.59
9.83
16.09
54.09

-

11. 01
63.50
61.13
20.73

---

123.51
15.48
26.32
260.96
---

622.83

723.42

- 100.58

-

13.90

1.92
19.03

206.57
11.18
23.20
263.01

109.91
5.65
10.23
206.86

---

--792.18

-

I

w

Total All Plants

-1/ Reflects

the amount

of forage

]j Derived

through comparison
on the control areas.

3/ Percent

-

~/ Indicates

changes

100.00

one would

of sprayed

1661.27

100.00

expect

to find on a treated

and control

the highest

confidence

level at which

1390.10

- 192.16

area after treatment

areas with adjustments

are only shown for the more abundant

a.05, .10, .15, .20 or .25 level are shown.

1197.94

for normal

-

had treatment

'"

lJ'

13.82

I

not occurred.

year to year changes in forage production

recorded

plants.

changes

due to spraying

are significant.

Only changes

that are significant

at the

�Table 7 •

Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining.
Lbs of vegetation
~roduced Eer acre

Percent com~osition
1/
E 73 -

A

A

Plant species

1970

1973

1970

1973

E 73

lbs/ac

26.82
2.50
0.18
0.65
0.30
0.00
0.36
0.24
1.13
0.00
0.00
2.14
85.94
0.00
0.00
0.00
0.48
0.00
3.87
0.18
3.75
0.00
2.62
0.00
0.00
1.07
0.00
0.00

86.78
5.77
0.06
3.45
0.00
0.00
0.00
0.06
5.06
0.00
0.18
2.91
77.86
0.00
0.00
0.06
0.71
0.18
0.65
0.54
2.26
0.12
1.07
0.00
0.00
0.12
0.59
0.00

58.08
2.30
10.61
4.19
0.02
0.00
0.02
0.08
1.67
0.00
0.01
1.84
61.45
0.00
0.00
0.01
0.57
0.01
2.99
0.15
4.94
0.12
3.97
0.00
0.00
1.65
0.12
0.00

+ 28.70
+ 3.47
- 10.55
- 0.74
- 0.02
0.00
- 0.02
- 0.02
+ 3.38
0.00
+ 0.17
+ 1.08
+ 16.41
0.00
0.00
+ 0.05
+ 0.14
+ 0.17
2.33
+ 0.39
- 2.68
0.00
- 2.90
0.00
0.00
- 1.53
+ 0.48
0.00

Change due to chaining ~
Confidence level
percent Y \ where signifi cant !y

-."-'

Forbs
AQhittea lanuio~a
Ag~taQhe untiQi60lia
AgMV1.M glauQa
Allium aQum.tnlttum
AmiJtMaQe Mp.te.n:tJUon~
Aqu.tleg-ta QoeJtuiea
AJtabM MummoncU.
AfW..bM IUM uta
A~enania Qong~.ta
AuemM-ta b.tennM
Aue.mMia dJtaQunQui~
AuemM-ta ludov.tQ.{.ana
M.teJt - EJUgeJto n
M.tJW..gal~ haydeni ..an~
B~amoM.tza
MgUtata
BeJtbV1.M ~epe~
CaloQhou~
gunnMonil
Cap~e.Ua buMa-p~.tow
C~.t.tlleja
lin~e6olia
ChenopocU.um beJtlancU.~
C~.tum Qen.ta~eae.
CoUom-ta line.aW
ComandJta umbe.Uata
Cymop.teJt~ p~p~e.~
Ve.lph.tnium nwoni
Ve.lph.tnium oQQiden.tale
V~ Q~a.tn.ta ~ 0 ph-ta
VMpo~um ~aQhy~pwn

1.50
0.14
0.01
0.04
0.02
0.00
0.02
0.01
0.06
0.00
0.00
0.12
4.78
0.00
0.00
0.00
0.03
0.00
0.21
0.01
0.20
0.00
0.15
0.00
0.00
0.06
0.00
0.00

4.38
0.30
0.00
0.16
0.00
0.00
0.00
0.00
0.25
0.00
0.01
0.15
3.93
0.00
0.00
0.00
0.03
0.01
0.03
0.03
0.11
0.01
0.05
0.00
0.00
0.01
0.03
0.00

3.47

3.68

+ 49.42
+150.75

.25

I

w

+202.28

.15

'-0

'"
I

+ 58.80
+ 26.71

.20

-

- 54.29
- 73.02

�Table

7.

Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining
Lbs of vegetation
~roduced ~er acre

Percent com~osition
Plant species

1970

1973

E 73 lJ

1970

1973

E 73

lbs/ac

1.49
2.38
0.00
44.79
0.00
7.32
0.00
0.00
0.42
0.89
0.00
0.18
6.25
0.00
0.00
104.20
0.00
23.08
0.71
0.00
40.68
0.00
0.00
0.18
3.63
0.00
0.18

12.27
0.65
0.00
19.88
0.00
2.83
0.00
0.36
0.02
1.05
0.00
0.74
7.26
0.06
0.00
83.86
0.00
17.35
0.87
0.00
54.09
0.00
0.00
0.01
0.53
0.00
0.01

- 10.78
+ 1.72
0.00
+ 24.91
0.00
+ 4.49
0.00
- 0.36
0.40
- 0.16
0.00
0.57
- 1.01
0.06
0.00
+ 20.34
0.00
+ 5.73
- 0.16
0.00
- 13.41
0.00
0.00
+ 0.17
+ 3.10
0.00
+ 0.17

(contd).

Change due to chaining ~
Confidence level
percent 11
where significant 11

Forbs (contd)
E.iUogonum umbilla;tum
Ep~ob~um pa~culatum
Fltaga.iUa amvUcMa
Gallum
GentIana heteJto~epala
GeJtaMum
Gilia aggltega,ta
GJUnde£ia aphana~
Hacketia leptophiflta
HueMum hoop~MHifdltophifltum capUa.tum
Hifdltophifltum 6endlvU
Iw m,u.,MuJUeM,u.,
Lactuc.a ~caJUola
Lappula Itedow~W
Lathiflt~ - V~ua
Lep~~um deM~6loltum
Ug~ticum
Unum lw,u., MLomatium leptoc.altpum
Lup~n~ altgei1teu,6
Madia glomeJtata
Medic.ago ~ativa
M~otu,6
o6Mu~
MeJtte~.(a 6JtanWc.Ma
MeJ1;te~~a lMc.eo~
Moldav~ca paltv~MOIta

0.25
0.00
0.00
1.26
0.00
0.34
0.00
0.02
.0.08
0.16
0.00
0.06
0.41
0.00
0.00
5.20
0.00
1.21
0.05
0.00
5.24
0.00
0.00
0.00
0.03
0.00
0.00

0.07
0.12
0.00
2.21
0.00
0.36
0.00
0.00
0.02
0.05
0.00
0.01
0.32
0.00
0.00
5.02
0.00
1.10
0.04
0.00
2.04
0.00
0.00
0.01
0.19
0.00
0.01

1.19

4.82
1.00
3.20

4.52
0.00
0.00
22.96
0.00
6.25
0.00
0.30
1.37
2.91
0.00
1.01
7.26
0.06
0.00
96.77
0.00
22.60
0.95
0.00
93.80
0.00
0.00
0.00
0.59
0.00
0.00

-

- 87.88
+125.31
+158.69

.25
I

w

\0
'-l

I

- 13.93
+ 24.25
+ 33.03
- 24.79

.15

�Table

7.

Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining (contd).
Lbs of vegetation
Qroduced Eer acre

Percent comEosition
Plant species

1970

1973

E 73 11

1970

1973

E 73

lbs/ac

0.00
0.00
0.54
4.10
4.46
0.00
0.00
0.00
0.77
0.30
6.42
0.36
0.48
13.20
0.00
0.00
2.97
0.06
0.12
0.06
0.00
0.00
1.01
27.18
0.00
4.82
17.72

0.00
0.00
0.12
5.24
3.46
0.00
0.08
0.10
0.30
0.01
21.95
1.31
0.71
6.51
0.00
0.00
0.55
0.01
0.71
0.89
0.18
0.00
0.00
15.82
0.00
9.71
13.82

0.00
0.00
+ 0.42
1.14
+ 1.00
0.00
- 0.08
- 0.10
+ 0.48
+ 0.29
- 15.52
- 0.95
- 0.23
+ 6.69
0.00
0.00
+ 2.43
+ 0.05
- 0.59
- 0.83
0.18
0.00
+ 1.01
+ 11.36
0.00
- 4.90
+ 3.90

Change due to chaining ~
Confidence level
percent '}j
where significant 11

Forbs (contd)
oenosheno: c.aupUo.6a
Opun..Ua
OlLthOC.MPM tuteU6
O.6mOJtlUza
Pac.hMtima mYM).nUu
Pe~c.~
g~ay).
PeYl.6temonMandaUM..
PeYl.6temon.6~c.tM
Phac.eUa hetVtophyUa
Phlox tong).6ot).a
PotemoMum 60U0.6M.6).mum
Potygonum av).c.uiMe
Potygonum dougta.6)')'
Potentitta puic.h~a
RanUYlc.uiU6).namoenM
Rumex MM pM
Rudbec.k).amontana
Senec.).oamb~o.6).o)'du
Senec.).o)'nteg~U6
Senec.).o.6 ~a
Silene menz).u).).
Smilac.).na.6teUata
SoUdo.go .6paM).6toM
SteUa.JUa
SweJttio.Md).ata
TMaXac.um o66).c.).nate
Thili~um
6 endleJt).

0.00
0.00
0.01
0.33
0.68
0.00
0.02
0.01
0.02
0.00
0.14
0.07
0.01
0.31
0.00
0.00
0.09
0.00
0.00
0.00
0.01
0.00
0.00
0.42
0.00
0.23
0.98

0.00
0.00
0.03
0.20
0.21
0.00
0.00
0.00
0.04
0.02
0.33
0.02
0.02
0.67
0.00
0.00
0.15
0.00
0.01
0.00
0.00
0.00
0.05
1.32
0.00
0.24
0.84

0.00
0.00
0.12
6.13
12.91
0.00
0.36
0.18
0.30
0.00
2.44
1.31
0.12
5.47
0.00
0.00
1.55
0.00
0.06
0.00
0.18
0.00
0.00
7.79
0.00
4.16
18.14

-

- 21.70
+ 28.82
I

w

'"'coI
- 70.73
+102.84

-

+ 71.77
- 50.41
+ 28.25

.15

�Table 7 . Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining (contd).
Lbs of vegetation
produced per acre

Percent composition
Plant species

1970

1973

E 73 11

A

1970

1973

E 73

1bs/ac

- 50.63
0.00
- 1.19
0.00
0.00
0.00
+ 0.24
0.00
6.12
2.96

Change due to chaining EI
Confidence level
where significant ~
percent Y

Forbs {contd}
Tnagopogon dubium
Tnagopogon pnateYiJ.&gt;.u..
Umc.a. dio.a:a:
ValeniMa. oc.uderU:al.M
Venonic.a amenic.ana
venonic.a biloba.
V,[guiena mu£ti6.tona
V,[o.ta nug u.tO-6a.
V,[o.ta -6heV:onil
WIje.tJua amp.te:Uc.a~

Total Forbs

0.10
0.00
0.24
0.00
0.00
0.00
0.08
0.02
0.22
0.91

0.12
0.00
0.16
0.00
0.00
0.00
0.07
0.03
0.08
0.62

1.78
0.00
4.22
0.00
0.00
0.00
1.43
0.30
4.04
17.13

2.44
0.00
3.03
0.00
0.00
0.00
1.49
0.59
1.72
13.26

53.06
0.00
4.22
0.00
0.00
0.00
1.25
0.59
7.84
16.23

--

--

--

--

--

26.56

26.29

483.67

531 .91

21.00
0.18
0.00
2.74
3.63
0.36
0.00
1.13
0.12
0.00
12.85

- 95.40

+ 18.79

-

- 78.00
- 18.27

424.38

+107.53

+ 25.34

1.22
0.00
0.00
0.22
0.02
0.04
0.00
0.00
0.12
0.00
0.00

1.05
0.01
0.00
0.14
0.17
0.02
0.00
0.05
0.01
0.00
0.66

0.55

21.89
0.00
0.00
3.98
0.36
0.65
0.00
0.00
2.14
0.00
0.00

9.13
0.01
0.00
2.22
7.61
0.01
0.00
0.01
0.54
0.00
0.06

+ 11.86
+ 0.17
0.00
+ 0.51
- 3.98
+ 0.35
0.00
+ 1.12
- 0.42
0.00
+ 12.79

+129.86

I

w

'"'"
I

Grasses
Agnopynon
Agnopynon c.wtatum
Agnopynon intenmedium
Bnomu.o MomalCk6
Bnomu.o inenm.u..
Bnomu.o tec.tonum
Canex
Dady.tCk6 g.tomeAa.ta
Uljmu.o g.taUc.Ck6
F utuc.a avina
F u,tuc.a thunbeni

.10

+ 23.10

�Table7.

Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining (contd).
Lbs of vegetation
produced per acre

Percent composition
Plant species

1970

1973

E 73 Jj

1973

E 73

1bs/ac

0.30
0.12
0.83
0.00
0.00
402.25
0.00
5.89

9.70
0.00
0.04
0.00
0.00
425.78
0.30
5.94

- 9.40
+ 0.12
+ 0.80
0.00
0.00
- 23.53
- 0.30
- 0.05

1970

Change due to chaining ~
Confidence level
percent 11
where significant 11

Grasses {contd}
Ho~deum b~aQhyanth~um
Jun.QU6 ba£;t{.QU6
KoelvUa
CJrM:ta:ta
O~zyOP.6-W hymen.oJ.dv.,
Ph.teum pM:teYl.6 e
Poa-C~ex
gey~
Silan.J.on. hy/.):tJUx
Supa

0.15
0.00
0.00
0.00
0.00
23.77
0.02
0.37

0.02
0.01
0.04
0.00
0.00
19.74
0.00
0.30

25.22

2.68
0.00
0.00
0.00
0.00
431.57
0.30
6.60

- 5.53

- 0.90

I
.p-

o

Total Grasses

25.92

22.21

470.17

451.37

386.77

+ 64.60

0.00
138.52
2.62
35.33
0.00
1.07
0.00
0.00
5.17
0.00
228.57
0.00

0.00
103.87
3.33
44.44
0.00
0.01
0.00
1.71
5.29
0.00
138.12
0.00

0.00
+ 34.65
- 0.71
- 9.11
0.00
+ 1.06
0.00
1.71
- 0.12
0.00
+ 90.45
0.00

+ 16.70

Shrubs
AQ~ g.tab~um
Amelan.QhJ.~ a.tn.J.6oUa
Mte.mi..I.J-i.a Qan.a
M:tem-i.6J.a :tJUden.:ta:ta
C~QOQ~PU6 mon.:tan.u-6
CMY-60:thamn.U6 depM-6-6U6
Ch~y/.)o:thamn.U6 n.au-6ecs U6
CMy/.) oxhamnu» v-W ucLi.MMU6
C~egu-6
~y:tMopoda
Po pu1.u-6 :tftemu1.oJ.dv.,
P~UYl.U6v~gJ.n.J.ana
PuM hJ.a ;0uden.:ta:ta

0.00
5.98
0.42
1.95
0.00
0.00
0.00
0.05
0.30
0.00
8.42
0.00

0.00
6.52
0.13
1.81
0.00
0.05
0.00
0.00
0.27
0.00
11 .19
0.00

5.86

7.91

0.00
111.58
7.49
34.91
0.00
0.00
0.00
0.95
5.29
0.00
154.88
0.00

-

+ 33.36
- 21.43
- 20.49

- 2.25
+ 65.48

0
I

�Table 7. Changes in production of air dry summer vegetation recorded on the chained areas two years following chaining (contd).
Lbs of vegetation
produced per acre

Percent composition
Plant speci es

1970

1973

E 73 Jj

A

1973

E 73

1 bs/ac

432.22
0.00
37.77
185.51

325.24
0.00
59.45
216.11

+106.99
0.00
- 21.68
- 30.60

1970

Change due to chaining ~
3/
Confidence level
percent where signiflcant 11

Shrubs (contd)
Quvr.c.uo gambellil
TUbe;.,
ROM. ntdk.a.na.
Sympho~c.~po~
albuo

17.81
0.00
2.40
10.18

20.54
0.00
1.89
9.11

18.64
3.57
12.68

331.82
0.00
43.54
185.45

+ 32.89
- 36.47
- 14.16

-Total Shrubs

47.51

51.50

875.93

1066.79

899.95

+166.83

.20

+ 18.54

.15
I
-l&gt;0

•....

Total All Plants

100.00

100.00

1829.77

2050.07

1709.99

+340.08

I

+ 19.89

1/
- Reflects the amount of forage one would expect to find on a treated area following treatment had treatment not occurred.
~Derived
through comparison of chained and control areas with adjustments
on the control areas.
1/Percent

for normal year to year changes in forage production recorded

changes are only shown for the more abundant plants.

1IIndicates the highest confidence level at which changes due to chaining are significant.
a .05, .10, .15, .20 or .25 level are shown.

Only changes that are significant

at the

�Table 8.

Changes in production of air dry summer vegetation recorded on the chained areas five years following chaining.
Lbs of vegetation
Eroduced Eer acre

Percent comEosition
Plant Species

1970

1976

,

E 761/

-

1970

1976

E 76

1bs/ac

26.82
2.50
0.18
0.65
0.30
0.00
0.36
0.24
1.13
0.00
0.00
2.14
85.94
0.00
0.00
0.00
0.48
0.00
3.87
0.18
3.75
0.00
2.62
0.00
0.00
1.07
0.00
0.00

60.19
2.20
0.00
0.12
0.00
0.00
0.12
0.00
0.89
0.00
0.00
4.10
46.99
0.00
0.00
0.00
0.00
0.00
2.20
0.54
3.63
0.12
0.18
0.00
0.00
0.24
0.24
0.00

47.31
1.51
0.18
1.18
0.02
0.01
0.59
0.01
2.58
0.01
0.01
1.84
52.61
0.01
0.01
0.03
0.00
0.01
0.53
0.12
2.55
1.37
5.66
0.00
0.01
0.62
2.08
0.01

+12.89
+ 0.69
- 0.18
- 1.06
- 0.02
- 0.01
- 0.48
- 0.01
- 1.68
- 0.01
- 0.01
+ 2.27
- 5.62
- 0.01
- 0.01
- 0.03
- 0.00
- 0.01
+ 1.67
+ 0.42
+ 1.08
- 1.25
- 5.48
- 0.00
- 0.01
- 0.38
- 1.84
- 0.01

Change due to Chaining l/
Confidence level 4/
.
3/
percent where significant -

Forbs
Achillea lanulosa
Agastache urticifolia
Agoseris glauca
Allium acuminatum
Androsace septentrional is
Aquilegia coerulea
Arabis drummondi
Arabis hirsuta
Arenaria congesta
Artemisia biennis
Artemisia dracunculus
Artemisia ludoviciana
Aster-Erigeron
Astragalus haydenianus
Balsamorhiza sagittata
Berberis repens
Calochortus gunnisonii
Capsella bursa-pastoris
Castilleja linariaefolia
Chenopodium berlandieri
Cirsium centaureae
Collomia linearis
Comandra umbellata
Cymopterus purpureus
Delphinium nelsoni
Delphinium occidentale
Descurainia sophia
tri.s porum trachycarpum

_____

M.

1.50
0.14
0.01
0.04
0.02
0.00
0.02
0.01
0.06
0.00
0.00
0.12
4.78
0.00
0.00
0.00
0.03
0.00
0.21
0.01
0.20
0.00
0.15
0.00
0.00
0.06
0.00
0.00

5.24
0.20
0.00
0.01
0.00
0.00
0.01
0.00
0.08
0.00
0.00
0.33
4.14
0.00
0.00
0.00
0.00
0.00
0.17
0.05
0.33
0.01
0.02
0.00
0.00
0.02
0.02
0.00

3.07

3.50

•

+27.24
+45.55

.,..I
0
N

I

+123.61
-10.69

+42.32

_

�Table 8.

Changes in production of air dry summer vegetation recorded on the chained areas five years following chaining.

Percent comEosition
Plant Species

Eriogonum umbellatum
Epilobium paniculatum
Fragaria americana
Galium
Gentiana heterosepala
Geranium
Gilia aggregata
Grindelia aphanactis
Hackelia leptophylla
Helenium hoopesii
Hydrophyllum capitatum
Hydrophyllum fendleri
Iris missouriensis
Lactuca scariola
Lappula redowskii
Lathyrus-Vicia
Lepidium densiflorum
Ligusticum
Linum lewisii
Lomatium leptocarpum
Lupinus argenteus
Madia glomerata
Medicago sativa
Melilotus officinalis
Nertensia franciscana
Nertensia lanceolata
Moldavica parviflora

1970

0.25
0.00
0.00
1.26
0.00
0.34
0.00
0.02
0.08
0.16
0.00
0.06
0.41
0.00
0.00
5.20
0.00
1.21
0.05
0.00
5.24
0.00
0.00
0.00
0.03
0.00
0.00

1976

0.02
0.13
0.00
3.10
0.00
0.49
0.00
0.02
0.00
0.02
0.00
0.00
0.35
0.01
0.00
2.78
0.00
1.35
0.11
0.00
2.09
0.01
0.00
0.00
0.01
0.00
0.00

Lbs of vegetation
Eroduced Eer acre

E 76 J:j

1970

Forbs

(Cont'd).

1.19

3.29
0.80
2.13

21

A

4.52
0.00
0.00
22.96
0.00
6.25
0.00
0.30
1.37
2.91
0.00
1.01
7.26
0.06
0.00
96.77
0.00
22.60
0.95
0.00
93.80
0.00
0.00
0.00
0.59
0.00
0.00

(Cont'd).

1976

E 76

1bslac

0.18
1.49
0.00
37.41
0.06
5.89
0.00
0.24
0.00
0.18
0.00
0.00
3.81
0.06
0.00
36.10
0.06
18.08
1.19
0.00
24.03
0.12
0.00
0.00
0.12
0.00
0.00

32.29
1.31
0.01
17.75
0.01
2.18
0.18
0.55
0.02
0.67
0.18
0.21
7.26
0.06
0.00
51.21
0.01
12.19
0.48
0.01
32.28
0.01
0.00
0.00
0.04
0.01
0.36

-32.11
+ 0.18
- 0.01
+19.66
+ 0.05
+ 3.71
- 0.18
- 0.31
- 0.02
- 0.49
- 0.18
- 0.21
- 3.45
0.00
0.00
-15.10
+ 0.05
+ 5.89
+ 0.71
- 0.01
- 8.25
+ 0.11
0.00
0.00
+ 0.07
- 0.01
- 0.36

Change due to Qhaining Confidence level 4
where significant _I
percent 11

+110.76

.10

+170.12

.05
I

-I"0

w
I

- 47.54
- 29.50
- 48.29
- 25.57

�Table 8.

Changes in production of air dry summer vegetation recorded on the chained areas five years following chaining.

Percent composition
Plant Species

Oenothera caespitosa
Opuntia
Orthocarpus luteus
Osmorhiza
Pachistima myrsinites
Pedicularis grayi
Penstemon crandallii
Penstemon strictus
Phacelia heterophylla
phlox longifolia
Polemonium foliosissimum
polygonum aviculare
polygonum douglasii
Potentilla pulcherrima
Ranunculus inamoenus
Rumex crispus
Rudbeckia montana
Senecio ambrosioides
Senecio integerrimus
Senecio serra
Silene menziesii
Smilacina stellata
Solidago sparsiflora
Stellaria
Swertia radiata
Taraxacum officinale
TlJalictrum fendleri

1970

0.00
0.00
0.01
0.33
0.68
0.00
0.02
0.01
0.02
0.00
0.14
0.07
0.01
0.31
0.00
0.00
0.09
0.00
0.00
0.00
0.01
0.00
0.00
0.42
0.00
0.23
0.98

1976

0.00
0.00
0.06
0.06
0.24
0.00
0.00
0.05
0.03
0.16
0.02
0.09
0.10
0.83
0.00
0.00
0.26
0.00
0.00
0.00
0.00
0.00
0.00
0.13
0.00
0.04
0.52

Lbs of vegetation
produced per acre

E 76 Y

1970

Forbs

(Gont'd).
0.00
0.00
0.12
6.13
12.91
0.00
0.36
0.18
0.30
0.00
2.44
1.31
0.12
5.47
0.00
0.00
1.55
0.00
0.06
0.00
0.18
0.00
0.00
7.79
0.00
4.16
18.14

1976

E 76

1bs/ac

0.00
0.00
0.65
0.83
3.33
0.00
0.00
0.65
0.48
1. 78
0.24
0.95
1.19
9.28
0.00
0.00
2.85
0.06
0.06
0.00
0.00
0.00
0.00
1. 72
0.00
0.54
6.96

0.01
0.18
0.12
2.41
10.86
0.71
0.17
0.38
0.30
0.65
4.88
1.31
10.61
5.47
0.01
0.01
0.14
0.01
0.71
0.01
2.12
0.01
0.00
5.55
0.12
5.46
5.57

- 0.01
- 0.18
+ 0.54
- 1.57
- 7.53
- 0.71
- 0.17
+ 0.28
+ 0.18
+ 1.13
- 4.64
- 0.36
- 9.42
+ 3.81
- 0.01
- 0.01
+ 2.72
+ 0.05
- 0.65
- 0.01
- 2.12
- 0.01
- 0.00
- 3.83
- 0.12
- 4.93
+ 1.38

(Cont'd).

Change due to Chatnirtg~j
3/
Confidence level 4/
percent where significant -

- 65.38
- 69.33

I
.t:-

o
-l"-

I

+ 69.57

- 68.93

.20

+ 24.84

---------------------------------------------------------------------------------------------------------------------------------------------

�Table 8.

Changes

in production

of air dry summer vegetation

Percent
Plant

Species

Tragopogon dubium
Tragopogon pratensis
Urtica dioica
Valeriana occidentalis
Veronica americana
Veronica biloba
Viguiera multiflora
Viola rugulosa
Viola sheltonii
Wyethia amplexicaulis

1970

0.10
0.00
0.24
0.00
0.00
0.00
0.08
0.02
0.22
0.91

--Total Forbs

26.56

recorded

0.05
0.00
0.06
0.00
0.00
0.00
0.07
0.08
0.05
0.31

-24.33

areas five years following

Lbs of vegetation
produced per acr~

composition
1976

on the chained

E 76 Y

1970

Forbs

(Cont'd)
1. 78
0.00
4.22
0.00
0.00
0.00
1.43
0.30
4.04
17.13

Change
E 76

1bs/ac

0.54
0.00
0.71
0.00
0.00
0.00
0.89
0.95
0.65
4.16

1.78
0.01
0.60
0.01
0.01
0.01
1.54
0.54
8.34
12.17

- 1.25
- 0.01
+ 0.11
- 0.01
0.01
- 0.01
- 0.65
+ 0.41
- 7.68
- 8.01

- 65.79

0.23

---

--

483.67

290.25

290.92

- 0.67

16.59
1.49
0.00
6.66
0.42
0.59
1.01
4.70
0.36
0.00
12.73

0.94
0.01
0.00
1.98
0.02
0.02
0.01
0.01
0.65
0.00
0.19

+15.65
+ 1.48
- 0.00
+ 4.69
+ 0.40
+ 0.58
+ 1.00
+ 4.69
- 0.30
0.00
+12.54

(Cont'd).

due to Ghaining 11
3
Confidence level 4
percent _I
where significant _I

1976

---

chaining.

I

.I&gt;-

o

Grasses
Agropyron
Agropyron cristatum
Agropyron intermedium
Bromus anomalus
Bromus inermis
Bromus tectorum
Carex
Dactylus glomerata
Elymus glaucus
Festuca ovina
Festuca thurberi

1.22
0.00
0.00
0.22
0.02
0.04
0.00
0.00
0.12
0.00
0.00

1.46
0.13
0.00
0.58
0.03
0.05
0.09
0.40
0.03
0.00
1.16

21.89
0.00
0.00
3.98
0.36
0.65
0.00
0.00
2.14
0.00
0.00

------------------------------------------------------------------------------------------------------------

+237.07

------------------------------~--

U1

I

�Table 8.

Changes in production of air dry summer vegetation recorded on the chained areas five years following chaining.
Lbs of vegetation
Eroduced Eer acre

Percent comEosition
Plant Species

Hordeum brachyantherum
Juncus balticus
Koeleria cristata
Oryzopsis hymenoides
Phleum pratense
Poa-Carex geyeri
Sitanion hystrix
Stipa

1970

0.15
0.00
0.00
0.00
0.00
23.77
0.02
0.37

1976

0.04
0.00
0.01
0.00
0.08
15.68
0.05
0.62

A

E 76 Y

1970

Grasses

(Cont'd)

44.28

2.68
0.00
0.00
0.00
0.00
431.57
0.30
6.60

1976

E 76

1bs/ac

0.48
0.00
0.06
0.00
0.83
196.81
0.54
7.20

3.26
0.00
0.01
0.01
0.01
670.06
3.54
3.52

- 2.78
0.00
+ 0.05
- 0.01
+ 0.82
-473.24
- 3.00
+ 3.68

(Cont'd).

Change due to Chaining '!:./
Confidence level 4/
3/
where significant percent -

- 70.63
+104.39

.,..I
0
0I

Total Grasses

25.92

470.17

20.40

250.46

477 .08

-226.62

0.00
110.15
3.93
39.20
0.00
0.00
0.00
0.00
6.48
0.00
157.67
0.00

0.01
125.77
3.75
70.69
0.00
0.01
0.00
0.79
5.29
0.01
197.43
0.00

- 0.01
-15.62
+ 0.18
-31.50
0.00
- 0.01
- 0.00
- 0.79
+ 1.19
- 0.01
-39.76
0.00

- 47.50

Shrubs
Acer glabrum
Amelanchier alnifolia
Artemisia cana
Artemisia tridentata
Cercocarpus montanus
Chrysothamnus depressus
Chrysothamnus nauseosus
Chrysothamnus viscidiflorus
crataegus erythropoda
Populus tremuloides
Prunus virginiana
Purshia tridentata

0.00
5.98
0.42
1.95
0.00
0.00
0.00
0.05
0.30
0.00
8.42
0.00

0.00
8.19
0.36
3.49
0.00
0.00
0.00
0.00
0.59
0.00
12.89
0.00

7.99

12.53

0.00
111.58
7.49
34.9l
0.00
0.00
0.00
0.95
5.29
0.00
154.88
0.00

- 12.42
+ 4.76
- 44.55

+ 22.47
- 20.14

----------------------------------------------------------------------------------------------------------------------------------------------

�Table 8.

Changes in production of air dry summer vegetation recorded on the chained areas five years following chaining.
Lbs of vegetation
Eroduced Eer acre

Percent comEosition
Plant Species

Quercus gambellii
Ribes
Rosa nutkana
Symphoricarpos albus

1970

17.81
0.00
2.40
10.18

1976

18.49
0.02
2.42
8.82

E 76 'y

1970

Shrubs

(Cont'd)

12.79
3.29
12.22

A

331.82
0.00
43.54
185.45

1976

E 76

243.32
0.24
28.79
111.64

198.40
0.01
49.39
184.00

(Cont'd) .

Change due to Chaining 1/
Confidence level 4/
percent 1/
where significant Ibs/ac

+44.92
+ 0.22
-20.61
-72.36

+ 22.64
- 41.72
- 39.33

--Total Shrubs

47.51

55.27

875.93

701.42

799.90

-98.48

- 12.31
I

...,

---Total All Plants

100.00

100.00

1829.77

1242.13

1531.09

-288.97

""0

I

- 18.87

-1/ Reflects the amount of forage one would expect to find on a treated area after treatment had treatment not occured.

1/ Derived through comparison of sprayed and control areas with adjustments for normal year to year changes in forage production recorded
on the control areas.
-3/ Percent changes are only shown for the more abundant plants.

i/ Indicates the highest confidence level at which changes due to spraying are significant.
.10, .15, .20 or .25 level are shown.

Only changes that are significant at the a.05,

�-408Table 9. Two and 5 year changes recorded
elk, deer and cattle forage. 1/ l/ 1/.

in available

seasonal air-dry

Animal

Season-

Pre- to Post-treatment Forage Production Rati~/
Burning 6/
Spraying
Chaining
2 yrs.
5 yrs.
2 yrs.
5 yrs.
2 yrs.
5 yrs.

Elk

Winter

1:0.64

1:0.74

1:1.01

1:0.85

1: 1. 24

1:0.75

Spring

1:0.84

1:0.60

1:2.09

1:0.72

1:1.21

1:0.61

Summer

1:0.80

1:0.62

1:1. 72

1:0.76

1:1.13

1:0.60

Fall

1:0.67

1:0.60

1: 1. 94

1:0.77

1:1.12

1:0.65

Winter

1:0.54

1:0.65

1:1. 02

1:0.98

1: 1. 23

1:0.98

Spring

1:0.59

1:0.52

1: 1.40

1:0.82

1:1.17

1:0.76

Summer

1:0.84

1:0.72

1: 1. 20

1:0.79

1: 1.18

1:0.82

Fall

1:0.66

1:0.78

1:1.l3

1:0.91

1: 1. 26

1:0.96

1:0.95

1:0.76

1:2.17

1:0.84

1:1. 24

1:0.70

Deer

4/

Cattle Summer

l/ Table

is based on the assumption that due to such factors as degree of
plant maturity and snow cover, plant availability to elk and deer will consist
of only shrubs during winter; grasses and shrubs during spring and fall; and
forbs, grasses and shrubs during summer. Data in this table are based on
measurements made during summer.

l/

Classification of study area plants as elk and deer forage was based on
the average relative consumption of those plant species eaten by elk in 48
food habits studies and by deer in 99 food habits studies reported in the
literature (Kufe1d 1973; Kufe1d, Wa11mo and Feddema 1973).

1/ Plants

considered cattle forage are those classified as "desirable" and
"intermediate" by the U. S. Forest Service, Region 2, Range Analysis Handbook
for the foothill shurb, aspen weed, and mountain meadow vegetation types.

-4/ Seasons:
May; Summer

Winter = December, January, February; Spring = March, April,
= June, July, August; Fall = September, October, November.

l/ The ratio of elk and deer forage produced

before and after treatment
adjusted for normal year to year changes measured on control areas.
A single
value for elk and deer forage was derived by weighting pounds per acre of
"highly valuable" or "heavily eaten" forage by a factor of "3", "valuable"
or "moderately eaten" forage by "2", and "least valuable" or "lightly eaten"
forage by "1". A total weighted value was then computed.

. d th roug h comparlson
.
. h ad·Justments
-6/ D erlve
0 f treate d an d contro 1 areas Wlt
for normal year to year changes in forage production recorded on the control
areas.

�Table

10.

Elk use recorded

1/
Year-

Block 1

on treated

Control
Block

2

areas one year before

Ave.

Block 1

and 2 and 5 years

after burning,

spraying

Elk Density Per Sguare Mile
Burn
Block 1
Ave.
Block 2

SEray
Block

and chaining.

2

Ave.

Block 1

Chain
Block 2

Ave.

1 yr. before

8.4

10.0

9.2

11.8

18.6

15.2

11.8

13.6

12.9

21.8

18.6

20.2

2 yrs. after

6.5

18.4

12.5

24.9

35.0

30.1

6.5

11.7

9.1

6.5

9.8

8.3

5 yrs. after

15.1

21.6

18.3

46.6

46.6

46.6

15.1

50.0

32.6

50.0

40.1

45.1

Change

Before

to 2 yrs. After

% Change

+ 3.3

+14.9

- 3.8

-11.9

+36~;

+98%

-29%

-59%

~I

0
-0

Change

Before

to 5 yrs. after

% Change

Change

due to treatment,

% Change

Change

due to treatment,

due to treatment,

% Change

due to treatment,

Before

+31.4

+19.7

+24.9

+99%

+207%

+153%

+123%

+ 9.4

- 8.4

-19.1

+45%

-48%

-70%

+16.4

+ 6.9

+ 4.9

+54%

+27%

+12%

2/
to 2 yrs. After-

Before

Before

+ 9.1

to 2 yrs. After

to 5 yrs. After

Before

to 5 yrs. After

--J)

Dates pellet plots were cleared and accumulated pellets subsequently counted
treatment - 9-3-73 and 5-8-74; Five year post-treatment
- 8-31-76 and 5-4-77.

were:

Pre-treatment

- 9-3-70

and 5-5-71;

Two year post-

Jj
Change in use due to treatment use computed by comparing the ratio of pre-treatment
to 2 and 5 year post-treatment
elk densities
mile in the control areas with the pre-treatment
to 2 and 5 year post-treatment
elk densities per square Inile in the treated areas.

per square

I

�Table

11.

Deer use recorded

1/
Year-

Block

on treated

1

areas one year before

Control
Block 2

Ave.

Block

1

and 2, 3 and 5 years after burning,
Elk Density
Burn
Ave.
Block 2

spraying

Per S~u~a~r~e~M=i~l~e~
Spray
Block 1
Block 2

and chaining.

~n~L__~----------

Ave.

Block

1

wlain
Block 2

Ave.
15.9

22.7

6.5

15.2

25.2

19.4

30.2

16.0

25.2

15.2

13.6

16.8

1 yr. before

13.5

7.4

2 yrs. after

16.7

9.8

18.4

4.9

15.0

15.0

16.6

19.1

16.6

21.5

11.6

20.2

6.0

23.2

17.1

17.1

27.9

20.0

19.3

15.5

18.6

40.3

3 yrs. after

24.1

28.4

18.5

26.5

17.6

24.2

21.6

13.5

20.0

25.9

21.6

33.3

5 yrs. after
Change

Before

to 2 yrs. After

- 1.0

- 8.5

- 6%

-53%

+11.9

- 4.3

11%

+74%

-27%

+ 1.5

+ 9.9

+ 8.2

- 1.8

+52%

+ 7%

+62%

- 9%

-

- 8.3

- 5.6

- 6%

-53 %

-25%

+12.0

- 4.2

- 2.4

+75%

-27%

-11%

+11. 4

+ 9.7

+79%

+67%

- 0.3

- 6.0

0%

-26%

- 0.1

- 2.5

0%

% Change
Change

Before

to 3 yrs. After

% Change
Change

Before

to 5 yrs. After

% Change
Change

due to treatment,· Before

% Change due to treatment,
Change

due to treatment,

Before
Before

% Change due to treatment,
Change

due to treatment,

% Change

to 2 yrs. Afte~/

due to treatment,

After

to 3 yrs. After

Before
Before

to 2 yrs~

to 3 yrs"

After

to 5 yrs. After

Before

to 5 yrs '.After

+ 3.6
+17%

.08

_1/ Data pellet plots were cleared and accumulated
pellets subsequently counted were:
Pre-treatment
- 9-3-70 and 5-5-71;
9-3-73 and 5-8-74; Three year post-treatment
- 9-6-74 and 5-30-75; Five year post-treatment
- 8-31-76 and 5-4-77.

]j Change in use due to treatment was computed
square mile in the control

areas with

Two year post-treatment

by comparing the ratio of pre-treatment
to 2, 3, and 5 year post-treatment
elk densities per
the ]Jre-treatment; and 2 and 5 year post-treatment
elk densities per square mile in the treated areas.

I
.p.

•.....
0
I

-

�Table

12.

Cattle use of control and treated areas one year before and 2 and 6 years after burning,

Cow Days Use/Acrell
2 yrs. After

11

Change in Cow Days UselAcre
1 yr. Before
1 yr. Before
to 2 yrs. After
to 6 yrs. After

Unit

Treatment

1 yr. Before

4

Control

0.0

3.0

2.2

+ 3.0

+ 2.2

6

Control

1.3

hl

&amp; 6

Control

0.7

4.8

~
1.1

+~
+ 4.1

+ 1.0

Burn

4.4

19.6

9.6

+15.2

+ 5.2

7

Burn

31.7

12.:..9.

+22.1

+ 0.4

&amp; 7

Burn

~
7.0

25.7

9.8

+18.7

+ 2.8

4

2
5

6 yrs. After

spraying

and chaining.

Significance of Change from Control
Areas at a .10 Level
1 yr. Before
1 yr. Before
to 2 yrs. After
to 6 yrs. After

-~

Yes

No
I

~,

Spray
Spray

8.7

.lJl

7.4

1.3

- 1.3

- 7.4

10.9

5.2

+~

+~

+ 3.3

- 2.6

0.0
-11.8

2 &amp; 5

Spray

5.9

9.2

3.3

3

Chain

0.0

0.0

0.0

0.0

8

Chain

18.3

8.7

6.5

- 9.6

3 &amp; 8

Chain

11 Cow defecation
11 Post-treatment

_ 11

rate is 11.5 chips per day as determined

by Julander

•....
•....
I

Yes

No

No

No

(1955).

data for all units have been adjusted for differential cattle stocking rates which existed one year before and 2 and 6 years
after treatment.
Data have also been adjusted for differences in stocking rates between 2 cattle allotments which comprise the study area.
Units 1, 5, 6, 7, 8 and a portion of 3 are in Buzzard allotment, and 2, 4 and the remainder of 3 are in Porter allotment.

11 Data for chained areas were not averaged.

High creek water during the 2nd year, and difficulty for cattle in finding Unit 3 caused the
absence of cattle in Unit 3 during the 2nd and 6th year after treatment, even though the area had been substantially improved for cattle by
chaining.

��July, 1978
-413-

JOB PROGRESS REPORT

State of

COLORADO
------------~--~------------

Project No.
Work Plan No.
Job Title

Job No.
1
8
----~~--~---------------Digestible Nutrient and Trace Mineral
Content of Deer and Elk Forage Plants During Winter

----~~~~~~--~----~~~~~~~~--~~~----------

Period Covered:
Personnel:

Game Range Investigations

W-101-R-20

April 1, 1977 - June 30, 1978

Roland C. Kufeld, Marilyn Stevens

ABSTRAcr
Nutrient and trace mineral levels were determined for 45 Gambel oak and
45 big sagebrush samples collected during January, 1976. Each species was
collected from 45 sites throughout the western half of Colorado.
Collection
sites for each species represented 9 major areas of the state and 3 vegetation
types. For Gambel oak the median crude protein level from the 9 areas is 5.2
percent, and the 95 percent confidence interval ranges for 4.6 to 5.8 percent.
For big sage the median crude protein level is 10.1 percent and the 95 percent
confidence interval ranges from 8.8 to 10.6 percent.
Medians and 95 percent
confidence intervals for other nutrients and trace minerals are presented.

��-415-

DIGESTIBLE NUTRIENT AND TRACE MINERAL
CONTENT OF DEER AND ELK FORAGE PLANTS DURING WINTER
Roland C. Kufeld

P. N. OBJECTIVE
To estimate the average digestible nutrient content values and degree of
variation in digestible nutrient content of selected range forage plants
during winter.
SEGMENT OBJECTIVE
To determine the degree of variation in digestible
selected range forage plants during winter.

nutrient

content of

INTRODUCTION
Proper mangement of big game range requires knowledge of the nutrient content
of existing forage plants.
Nutrient content of forage species which make up
the diet of an animal must be high enough to meet the animals minimum requirements.
Evaluation of plant nutrient levels on an area of rangeland involves chemical
analysis of each species of forage on the area, which is consumed by the
animal being managed, to determine content of various nutrients and their
degree of digestibility.
Such an analysis is very time consuming and expensive.
If a number of ranges were being evaluated the time and cost of a detailed
analysis of each plant species on each range would be prohibitive.
Evaluation
of a number of ranges would be feasible only if it were possible to apply an
average nutrient content value to each of the forage species encountered.
Plant nutrient content levels are influenced by many factors including season
of the year, climate, weather, altitude, light and soil constituents such
as parent material, texture, depth, nutrient content, moisture and temperature
(Laycock and Price, 1970). Cook and Harris (1950) concluded that "the nutrient
content of forage is influenced by many interdependent factors and the result
is the additive or mass effect of all factors operating simultaneously".
Therefore, use of average nutrient content data in range evaluations will be
feasible only if variation due to all factors is reasonably small. The purpose
of this study is to determine if variation in nutrient content of individual
range forage species growing under widely diversified conditions and in
different geographical locations is small enough to permit use of average
nutrient content values in range evaluations.
This phase of the study considers nutrient content variation in 2 shrub species,
Gambel oak (Quercus gambellii) and big sagebrush (Artemisia tridentata).
Nutrient content of these species was measured during winter because that is
considered the most critical time of the year for big game in Colorado.

�-416-

METHODS AND MATERIALS
During January, 1976, five 70 to 100 gram, green weight, composite samples
of two species, Gambel oak and big sagebrush were collected from 9 widely
separated areas throughout the western half of Colorado, but within the
normal winter range of deer and elk. The 5 collection sites within each
major area were widely separated geographically, but were situated within
the same vegetation type. Vegetation types and major areas of the state
where oak and big sagebrush samples were collected are shown in Table 1.
Table 1. Major areas within Colorado where winter samples of Gambel oak
and Big Sagebrush were collected.
Vegetation type where collections were made
.
.
Gambel oak Collections
Big Sagebrush Collections
Ponderosa pine
Big
Gambel
Gambel oak
Pinyon-Juniper
Gambel-oak
Sagebrush
Pinyon-juniper
oak

_

1 Steamboat
Sp.-Hayden

Meeker-Rifle

Denver Castlerock

Craig

Piceance Strawberry

Steamboat
Sp-HaydenMeeker

2 SiltCarbondaleEagle

Naturita

Durango Pagosa Sp.

Middle
Park

Naturita

SiltCarbondaleEagle

3 DoloresMancos

Colo. SpCanyon City

LaVeta Rye

Gunnison

Cortez Durango

Black CanyonPaoniaCedaredge

Oak collections consisted of current annual growth stems with leaves and acorns
excluded.
Big sagebrush collections consisted of current annual growth stems
with leaves attached but seed stalks excluded.
Samples of big sagebrush were
sealed in plastic bags, refrigerated immediately upon collection and frozen
as soon thereafter as possible to minimize vaporization of essential oils which
may result in unrealistically high digestion coefficients.

~Or[lOnor II[~ l~rn~lLW~ aIr Oll~~lD ~r~[~1r~lln~ 'n I

J

, 1a p

fi
ITil
analyzed in the Division of Wildlife Research Laboratory.
Samples were analyzed
to determine content of the following nutrients and trace minerals:
Total cell
contents, crude protein, ether extract, soluble ash, soluble carbohydrate, total
cell walls, acid insoluble ash, lignin, hemicellulose, cellulose, calcium,
copper, iron, magnesium, manganese, phosphorus, potassium, sodium, zinc (Harris
1970; Adrian 1973). The remaining portion of each samples was retained for
determination of digestible dry matter and digestible ~nergy. That analysis
has not yet been completed.
RESULTS AND DISCUSSION
Median values and 95 percent confidence intervals for nutrient and trace mineral
content in Gambel oak and big sagebrush collected during January throughout
the western half of Colorado are presented in Tables 2, 3 and 4.

�-417-

Table 2. Medians, 95 percent confidence intervals and non-parametric
tolerance levels for nutrient content in Gambel oak collected during January
throughout Colorado.ll

Nutrient
Dry matter

content

Percent Content in Plant
MeClian
95% ConfiClence Interval
64.20

Non-parametric Lower I
Bound For 70% Point 1

62.80 - 65.80

62.00

CELL CONTENTS
Protein

5.20

4.60 -

5.30

4.60

Soluble Carbohydrate

27.21

26.05 - 30.73

25.31

Ether Extract

3.90

2.87 -

4.76

2.38

Soluble Ash

4.06

3.72 -

4.21

3.71

Total Cell Contents

39.72

38.65 - 43.56

38.62

CELL WALLS
Lignin

23.98

22.11 - 26.02

20.98

Hemicellulose

10.26

7.73 - 11. 78

6.97

Cellulose

24.35

22.03 - 26.69

18.43

Holocellulose

33.95

33.01 - 36.19

30.21

Insoluble

0.61

0.49 -

0.86

0.32

60.28

56.44 - 61.35

54.64

Ash

Total Cell Walls

1/

- Data are based on collections of 5 samples from widely
within each of 9 major areas throughout Colorado.

separated

locations

l/This represents the lowest 5 sample mean value collected from the 9 major
areas.
One can be 95 percent sure that at least 70 percent of the area
mean nutrient values encountered in Colorado, during January, would exceed
the minimum value listed for each nutrient.

�-418-

Table 3. Medians, 95 percent
tolerance levels for nutrient
throughout Colorado.11

Nutrient
Dry matter

content

confidence intervals and non-parametric
content in Big Sage collected during January

Percent Content in Plant
Median
95% Confidence Interval

53.00

Non-parametric Lower
Bound For 70% Poin~1

51.60

52.40 - 55.00
CELL CONTENTS

Protein

10.10

8.80 - 10.60

8.60

Soluble Carbohydrate

43.28

40.03 - 46.32

39.92

Ether Extract

5.38

4.74 -

6.12

3.82

Soluble Ash

4.32

4.07 -

4.43

4.02

Total Cell Contents

62.71

60.61 - 64.97

/

59.30

CELL WALLS
Lignin

11.84

11.40 - 12.41

11.35

Hemicellulose

6.50

6.19 -

6.84

5.98

Cellulose

18.48

16.49 - 20.50

16.08

Holocellulose

25.62

22.99 - 27.10

22.86

Insoluble Ash

0.25

0.20 -

0.49

0.14

Total Cell Walls

37.29

35.03 - 39.39

34.46

II

- Data are based on collections of 5 samples from widely separated
within each of 9 major areas throughout Colorado.

locations

l/This represents the lowest 5 sample mean value collected from the 9 major
areas.
One can be 95 percent sure that at least 70 percent of the area
mean nutrient values encountered in Colorado, during January, would exceed
the minimum value listed for each nutrient.

�-419-

Table 4. Medians, 95 percent confidence intervals and non-parametric
tolerance levels for mineral content in Gambel oak and big sage collected
during January throughout Colorado.ll

Plant

Trace
Mineral

Content in Plant (PPM)
Median
95% Confidence

Oak

Calcium

6240

Copper

10

9 -

10

9

Iron

66

48 -

72

46
1620

Big
Sage

1/

Interval

Non-parametric Lower
Bound for 70% Pointl/

5440 - 6760

5200

Magnesium

1920

1660 - 2140

Manganese

200

118 -

390

88

Phosphorus

230

180 -

260

170

Potassium

7680

7280 - 7920

6800

Sodium

76

48 -

88

46

Zinc

30

24 -

32

24

3000 - 3680

2920

Calcium

3360

Copper

16

10 -

18

7

Iron

128

108 -

178

108

Magnesium

1160

980 - 1320

880

Manganese

38

36 -

42

34

Phosphorus

490

430 -

530

430

Potassium

10120

9280 -10680

8960

Sodium

90

84 -

102

66

Zinc

26

20 -

29

18

- Data are based on collections of 5 samples from widely
within each of 9 major areas throughout Colorado.

separated

locations

llThis represents the lowest 5 sample mean value collected from the 9 major
areas. One can be 95 percent sure that at least 70 percent of the area mean
mineral values encountered in Colorado, during January, would exceed the
minimum value listed for each mineral.

�-420-

Analysis of variance shows significant differences at the 5 percent level
among vegetation types, among areas, and among areas within vegetation
types for most nutrients in both oak and big sagebrush.
Nutrient content
in big sagebrush was generally more variable than in oak. Even though these
significant differences occur, the actual differences are relatively small
(Tables 2 and 3). For oak the median crude protein level from the 9 major
areas was 5.2 percent, and the 95 percent confidence interval ranged from
4.6 to 5.3 percent.
For big sagebrush the median crude protein level was
10.1 percent, and the 95 percent confidence interval ranged from 8.8 to
10.6 percent.
The data reflect relatively little mid-winter variation from
one locality to another in content of any of the nutrients for Gambel oak
and big sagebrush (Tables 2 and 3).
I

Plant content of some trace minerals was fairly uniform while others were
more variable (Table 4). Trace mineral content might be expected to be more
variable than nutrient content due to local soil concentrations of certain
minerals throughout the state.
One-sided non-parametric tolerance intervals can be applied to further
minimize the decision-making error incurred in projecting nutrient and trace
mineral levels based on average values (Dixon and Massey, 1969). It is
desirable that nutrient and trace mineral levels be underestimated rather
than overestimated to avoid overestimation of the ability of the range to
supply the minimum needs of grazing animals unless a particular trace mineral
is toxic above certain levels (i.e. lead selenium).
This could lead to
overstocking.
One-sided non-parametric tolerance intervals permit determination
of the portion of the overall population of nutrient and trace mineral values
that could possibly be encountered for a given species, which, with 95
percent confidence, exceeds the smallest sample value. For example, using
the sample means, which range from 4.6 to 5.4 percent crude protein from the
9 areas where oak was collected, one can be 95 percent sure that at least 70
percent of the area crude protein values encountered in Gambel oak in Colorado
during January would exceed the minimum value of 4.6 percent.
By using the
minimum value a manager could, therefore, be reasonably sure his estimate of
protein production is on the conservative side. One sided non-parametric
tolerance intervals for nutrients and trace minerals in Gambel oak and big
sagebrush are presented in Tables 2, 3, and 4.
LITERATURE

CITED

Adrian, W. J. 1973. A comparison of a wet pressure digestion method with
other commonly used wet and dry-ashing methods.
Analyst 98:213-216.
Cook, C. Wayne, and Lorin E. Harris.
1950. The nutritive value of range
Utah
forage as affected by vegetation type, site and state of maturity.
State Agric. Exp. Sta. Tech. Bull. No. 344. 45pp.
Dixon, W. J., and F. J. Massey, Jr. 1969. Introduction
analysis.
McGraw-Hill, 3rd ed. 638pp.

to statistical

�-421-

LITERATURE
Harris, L. E.
animals.

CITED

(Cont'd)

1970. Nutrition research techniques for domestic and wild
Pub. by Lorin E. Harris, 1408 Highland Dr., Logan, Utah. Vol. 1.

Laycock, William, A., and Donald A. Price.
1970. Factors influencing
quality.
In: Range and wildlife habitat evaluation - a research
symposium.
USDA For. Servo Misc. Pub.No. 1147. pp.37-47.

Prepared

by

f&lt;~
o . "J C' %"'~r
1
/M

' J!j&lt;.',

/1..-6-.--t('(,.·uJ~

I

,

.."

Roland C. Kufeld
Wildlife Researcher

.

forage

�</text>
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                  <text>October 1978

JOB PROGRESS REPORT

State of

COLOP~O
--------~~~~~-----------

Project No.
Work Plan No.
Job Title

Migratory

W-88-R-23

Bird Investigations

1
J~ ~. ----------------------------------

1

Waterfowl Production Surveys
------------------------------~--------------------------------------

Period Covered:

April 21, 1977 to June 30, 1978

Personnel:
M. Nail and Staff, Monte Vista National Wildlife Refuge;
J. Creasy and G. Deutsher, Brown's Park National Wildlife Refuge; F. N. Folks,
C. Jensen, and C. Solace, Utah State Division of Wildlife Resources;
S. Petersburg, and J. Alston, Dinosaur National Monument; M. Strong, Bureau
of Land Management; S. Bissell, R. Clark, D. Denny, H. Donoho, J. Ellenberger,
J. Frothingham, J. Lorentzson, T. Rauch, W. Russell, G. Saville, R. Velarde,
R. Weldon, B. Widhalm, S. Steinert and M. Szymczak.
ABSTRACT
Water conditions for waterfowl were far below average for duck production in
all areas except North Park and Brown's Park. Reliable duck breeding pair
estimates were obtained in only two of the six trend areas in 1977, Yampa
Valley and Brown's Park. The post-nesting season population of 1,872 in
northwest Colorado was 20 percent above the 1976 level, primarily because
of excellent production on the Green River.
Canada goose (Branta canadensis)
production in north-central Colorado was 13 percent below the 1976 level but
about average.
An aerial survey on the White, Roaring Fork, Colorado and
Gunnison rivers totaled 418 adult geese. Aerial counts of nesting Canada
geese in the San Luis Valley resulted in an estimate of 100 breeding pairs,
exclusive of the Monte Vista National Wildlife Refuge.

��-3-

WATERFOWL

PRODUCTION

SURVEYS

Michael R. Szymczak
Steven F. Steinert
P.N. OBJECTIVES
1.

To estimate the number of duck breeding pairs, by species, on selected
major waterfowl nesting areas in Colorado.

2.

To estimate the number of goose breeding pairs, and in some cases,
obtain production data on selected goose nesting areas in Colorado.

3.

Compile data and submit reports to appropriate state personnel and
federal agencies for use in establishing hunting season recommendations.
SEGMENT OBJECTIVES

1.

To estimate the number of duck breeding pairs, by species,
San Luis, Cache 1a Poudre, South Platte and Yampa Valleys,
North Park and Brown's Park.

2.

To estimate the number of goose breeding pairs in the San Luis Valley
and west-central Colorado and obtain goose breeding pair and production
data on the Yampa, Little Snake and Green Rivers in northwest Colorado
and in north central Colorado.

3.

Compile data and submit appropriate
METHODS

in the
and in

reports.

AND MATERIALS

Present duck breeding pair and production surveys consist of a breeding
pair inventory of only major production areas.
The 1977 duck breeding pair surveys were conducted during the period of
May 9 to June 30. Surveys in North Park and the Cache 1a Poudre and South
Platte valleys were conducted exclusively from the air. Ground counts were
made in the Yampa Valley and Brown's Park. In the San Luis Valley intensive
aerial counts were adjusted for visibility by air-ground comparison studies.
Pair estimates for the Monte Vista National Wildlife Refuge in the San Luis
Valley were obtained from nesting transects.
All survey methods and sample
areas for ducks remained the same as in previous years.
Surveys of Canada goose production were conducted within the period of
April 21 to June 15. Production estimates for Moffat County were made
from as complete a count as possible of hatched or active nests and brood
size. Population estimates of north central Colorado were obtained from
counts of goslings and adults conducted from the ground during the period
in which the birds were flightless.
Population estimates in west-central
Colorado were estimated from aerial surveys.

�In the San Luis Valley a survey of expected productive breeding pairs was
conducted by flying transects in defined sample blocks as described by
Lowry (1974). Four evenly-spaced transects were flown in each 1 x 3 mile
selected sample block on the Rio Grande and Conejos rivers, while a simple
total count was conducted in sample blocks containing ponds or lakes, only.
Each selected section was flown with both fixed-wing and helicopter aircraft
and the results compared.
All fixed-wing flying was accomplished with a Cessna 185 aircraft and the
helicopter flying with a Bell 47G3B. Two observers were used in sampling
by transects with the fixed-wing aircraft while one observer was used for
the helicopter flights.
RESULTS AND DISCUSSION
A winter snow pack much less than normal resulted in water conditions
considered far below average for duck production in most areas. The
San Luis Valley entered into its fourth consecutive dry year. Only North
Park and Brown's Park approached average conditions.
Extremely heavy
rains in mid-May in the South Platte Valley produced local flooding and
nest destruction.
Reduced water flows in the Yampa and Little Snake
Rivers resulted in a reduced number of islands for goose nesting and
increased predation of established nests.
In north central Colorado
many of the reservoirs were not filled and many nesting structures on
those areas were not occupied at a normal rate. In spite of mild
weather in early spring, goose nesting chronology was later than normal
throughout most of the state.
A variety of mechanical failures compounded by unrealistic mathematical
projections resulted in the decision not to compute duck breeding pairs
estimates for the four major trend areas:
San Luis Valley, North Park,
South Platte Valley, and Cache la Poudre Valley.
Unrealistic visibility
rates probably resulted from abnormally small sample sizes of some species
on air-ground comparison transects in the San Luis Valley.
The North
Park data were affected since San Luis Valley visibility rates are
applied to North Park data. Recorder failures eliminated the South
Platte and Cache la Poudre data. Usable data were collected in the
Yampa River Valley and Brown's Park (Table 1).
A post-nesting season population of 1,872 Canada geese was estimated in
trend areas in Moffat County in northwest Colorado (Table 2). Both
total estimated number of geese and gosling production were below 1976
levels along the Yampa and Little Snake Rivers (Tables 3 and 4). In
contrast, total geese and gosling production on the Green River were
above 1976 levels, particularly in Dinosaur National Monument.
Overall,
both total geese and production remain above the lO-year average.
Results of the 1977 Canada goose production survey in north central
Colorado are presented by individual ~rea in Table 5. Gosling production
was down from 1975 levels in all trend areas except Denver (Table 6).

�-5-

Table 1. Summary of Colorado's
in selected areas, 1977.

duck breeding

ground population

Percent Change
From LongFrom
term
Average
1976

Total Est. Breeding Pairs
Long-term
Average J..I
1976
1977

Area

San Luis Valley

34,759

27,855

North Parkll

16,535

17,115

South Platte
Valley

10,062

6,773

Cache la Poudre
Valley

6,927

3,662

estimates

Yampa Valley

2,189

2,145

2,755

+ 2.1

- 20.5

Brown's

1,009

593

1,147

+ 70.2

- 12.0

71,021

59,307

Park

Total

1/ San Luis Valley

and North Park averages are based on results of 1964
through 1976 and 1968 through 1976 surveys, respectively, because of
changes in survey methods utilized prior to those dates.
Figures for
other areas are 22 year averages.

II Aerial

counts corrected by species from visibility
in the San Luis Valley.

ratios obtained

�-6-

Table 2. Number of Canada geese observed
Moffat County, Colorado, 1977.

Nesting
Pairs

Area

and estimated

Non-nesting
Adults

Total
Adults

production,

Estimated
Goslings

No. Total
1/ Birds

Yampa River
Craig-Juniper

Springs

25

209

259

72

331

Juniper Springs-Cross
Mountain

15

122

152

48

200

Lily Park

14

80

108

50

158

Subtotal

54

411

519

170

689

61

166

288

210

498

Dinosaur National
Monument

50

105

205

249

454

Subtotal

111

271

493

459

952

22

l33

177

54

231

187

815

1,189

683

1,872

Green River
Brown's

Little

Park

Snake River

Grand Total

1/

- Calculated
nests.

using average brood size observed

and number of successful

�-7-

Table 3.
1977 .

Total estimated

number of Canada geese observed,

Moffat

Percent

County,

From 1976

Change
From 1967-76
Average

1977

1976

1967-76
Average

689

690

565

0.1

+ 21.9

Brown's Park

498

405

334

+ 23.0

+ 49.1

Dinosaur National
Monument 1/

454

211

326

+115.2

+ 39.3

231

255

303

9.4

- 23.8

1,872

1,561

1~528

+ 19.9

+ 22.5

Area

Yampa River
Green River

Little Snake River

Total
1/

Not surveyed

Table 4.

until 1970.

Estimated

number of Canada goose goslings,

Moffat County,

1977.

Percent Change
From 1967-76
From 1976
Average

Area

1977

1976

1967-76
Average

Yampa River

170

257

169

- 33.9

+

Brown's Park

210

133

127

+ 57.9

+ 65.4

Dinosaur National
Monument 1/

249

104

117

+139.4

+112.8

Little Snake River

54

91

97

- 40.7

- 44.3

Total

683

585

510

+ 16.8

+ 33.9

0.6

Green River

1/

Not surveyed until 1970.

�-8-

Table 5. Results of the north central Colorado goose census, June 14
and 15, 1977.

Production
Area

Wellington

Subtotal
Fort Collins

Subtotal

Water Area

No.
Broods

Terry Lake
Water Supply and
Storage #4
Launer Pond
Douglas Reservoir
Stewart Pond
North Poudre #1
1/
Dry Creek ReservoirNorth Poudre #2
North Poudre #5
Bureau of Standards
Pond 111
Bureau of Standards
Pond ft2
Divide fl8
Elder Reservoir
Annex #8 1./
Van Sant Pond
Cobb Lake
Dale Pond
Watson
Curtis Lake

2
0
5
0
0
0
5
0
1

Total
No.
Goslings

Total No.
Adults &amp;
Yearlings

Total
Birds

6

48

54

o

1
12
9

1
28
9

16

o
o
o

1

1

25

48
10

48
35

o

4

4

6

2

8

o

o

26

26

o
o

o
o

2])

1

1
24
12
27
18
80

37

4
34

2

37
5
58
38
208
30
189
49

o

o

26
181
12
109
49

21

215

615

830

Peterson Ponds
1
Herring Lake
3
Maxwell Pond
0
College Lake
5
Dean Acres
0
Claymore Lake
6
Sterling Gravel Pits
5
Lindenmeier Lake
0
Grey Lakes
10
Novak Reservoir
1
Winick Ponds
3
Flatiron Gravel Pits
1
Anderson's Pond
4
4
Parkwood Lake
Kitchel Lake
10
Timnath Reservoir
0
Romi1y Gravel Pits
6
Fossil Creek Reservoir 10
1
Horseshoe Lake
3
Wolaver Ponds

3
18

2

2
6

5
28
33
171
8
71
106
97
68
3
17
6
45
31
70
18
37
61
6
20

73

288

613

901

3
4

44

10
33
153
8
50
88
97
24

1

2

11

6

o
18

o
21
18

o

4

2

10
15

35
16
26
18
16
19

44

o
21
42
4
14

------------------------------------------------------------------------

�-9-

Table 5. Results of the north central Colorado goose census, June 14
and 15,1977 (cont.).

Production
Area

Loveland

Water Area

Flatiron Reservoir
Boedecker Reservoir
Flatiron Gravel Pits
Big Thompson River
McNeil Reservoir

1
5
1
7

Subtotal
Boulder

No.
Broods

Ish Lake
Crystal Lake
Terry Lake
Faivre Ponds
Rest Home Pond
Valmont Reservoir
Boulder Valley Farm
Eddy Pond

1

Subtotal
Denver

Ketring Lake
14
Centennial Pond
0
Columbine Country Club 5
Chatfield Country Club 5
Bowles Lake
7
Kings Lake
Tule Lake
2
Grant Reservoirs
5
Marston Reservoir
Pinehurst Country Club 2
Clairfield Reservoir
11
Kendrick Lake
0
Federal Center
3
Sloans Lake
11
Standley Lake
4
Denver City Park
2
Colo. Blvd. &amp; Quincy
0
Blackmer Reservoir
1
Mohn Estate
1
Reservoir No. 3
0

-Sub t o t a L

Grand Total

Total
No.
Goslings

Total No.
Adults &amp;
Yearlings

Total
Birds

7
34
22
3
59

1
41
10
2
128

8
75
32
5
187

125

182

307

5

2

19
27
19
22
36
8

19
167
21
98
64
10
13

7
38
194
40
120
100
16
21

142

394

536

51

59
25
14
20
89
97
47
18
35
51

110
25
41

6

o
27
29
26
38
7

49
115
135
54

o

39
14
214
12
24
48
17
7
40

35
46
61
97
39
23
284
21
30
48
20
8
40

357

924

1,281

1,127

2,728

3,855

17
11
10
43

o
9

70
9
6

o
3
1

54

1/ Geese located on pond 1 mile north of Dry Creek.
2/ Goose on nest.
3/ Geese located 1, mile north of Annex and ~ mile west of No.8.

�-10-

The number of adults observed increased only in the Wellington trend
area (Table 7). The total number of adult geese observed was down
for the second consecutive year and was considerable below the 8-year
average.
The aerial survey in west-central Colorado total 143 geese on the White
River, 235 on the Colorado River, 18 on the Roaring Fork River and
22 on the Gunnison River (Table 8). The total number of birds counted
in 1977 was much higher than in 1976, however, the count date was quite
different; March 31, 1976 and May 9, 1977. A particular count period
has not yet been selected for the west-central survey.
The 1976 survey
was flown too early and the 1977 survey too late.
The estimated number of Canada goose breeding pairs in the San Luis
Valley in 1977, exclusive of the Monte Vista National Wildlife Refuge,
increased over 1976 levels (Table 9). Estimates obtained from both
helicopter and fixed-wing counts were essentially equal.

�-11-

Table 6. Number of Canada goose goslings
production trend areas, 1977.

produced

in north central Colorado

No. of Goslings
1969-1976
1976
Average

Percent Change
From
From
1969-1976
1976

Area

1977

Wellington

215

278

261

- 22.7

- 17.6

Ft. Collins

288

401

285

- 28.2

+

Loveland

125

144

93

- 13.2

+ 34.4

Boulder

142

162

225

- 12.3

- 36.9

Denver

357

304

284

+ 17.4

+25.7

Total

1,127

1,289

1,148

- 12.6

- 1.8

in north central

Colorado

Table 7. Number of adult Canada geese observed
production trend areas, 1977 .

No. of Adults
1969-1976
Average
1976

1.1

Percent Change
From
From
1969-1976
1976

Area

1977

Wellington

615

555

749

+ 10.8

- 17.9

Ft. Collins

613

653

703

6.1

- 12.8

Loveland

182

291

229

- 37.5

- 20.5

Boulder

394

430

619

8.4

- 36.3

Denver

924

995

1,265

7.1

- 27.0

Total

2,728

2,924

3,565

6.7

- 23.5

�-12--

Table 8.

West central Colorado

Area

Canada goose breeding

Singles

pair survey 1977.

Total
1977
1976

Pairs

Groups

Goslings

3

5

19

0

32

9

Rio Blanco L. - Rangley

11

24

37

4

9J=.!

36

RangleY-Utah

1

3

8

0

15

2

15

32

64

4

1431)

47

0

3

0

0

6

Silt-DeBeque

8

20

83

0

l3l

DeBeque Palisade

5

8

0

0

21

Palisade-5th

0

2

14

0

18

2

7

43

4

15

40

140

4

23s-!-1

134

2

6

4

0

18

25

Hotchkiss-Delta

0

0

0

0

0

Delta-Grand

0

11

0

0

22

Subtotal

0

11

0

0

22

6

Grand Total

32

89

208

8

4181/

212

White River
Meeker-Rio

Blanco L.

Line

Subtotal
Colorado

River

Glenwood

Springs-Silt

St. Bridge

5th St. Bridge-Utah

Line

Subtotal

59!:.!

Roaring Fork River
El Jebel-Glenwood

Springs

Gunnison River

Junction

II Does not include goslings observed.

�-13-

Table 9. Results of the San Luis Valley Canada goose breeding
survey, 1975-77.

Nesting Pairs

Year

pair

Projected Total Number
Grouped Birds
Non-nesting Pairs

1975
59

59

110

Helicopter

92

63

64

Fixed wing

77

69

30

Helicopter

100

101

130

Fixed wing

100

91

97

Helicopter

1/

Fixed wing-

1976

1977

1/ F;xed
w;ng
.
1e te.
~
~
counts 1ncomp

Prepared

£L

·7YlMLR2~

by

Michael R. SZym~a
Wildlife Researcher

C

.J/;;..(l!J.U
?

Steven F. SteInert
Wildlife Technician

II

��October

-15-

JOB PROGRESS

State of

1978

REPORT

COLORADO
--------~~~~~-----------

Work Plan No.
Job Title

2

Job No.

Studies of Canada Goose Populations

Period Covered:
Personnel:

Migratory

W-88-R-23

Project No.

Bird Investigations
6

in Colorado Transplant

_
Areas

April 1, 1977 to March 31, 1978

M. Conner, J. Corey, W. Dolezal, D. Flenthrope, D. Hopper,
J. Lorentzson, R. Oakleaf, R. Velarde, and M. Szymczak.

ABSTRACl'
A record number of 300 Canada geese (Branta canadensis) banded outside
Colorado were reported recovered within north central Colorado dUYing the
1976-77 hunting season. Bantry No. 2 Reservoir in southern Alberta was
added to the list of production areas contributing geese to the harvest
in north central Colorado.
A significant number of wild trapped local
geese from the Edmonton-Camrose area represents further evidence that
restoration flocks in that vicinity are producing geese which winter in
north central Colorado.
A total of 582 geese were banded on production
areas in north central Colorado, while only 37 were banded in the San Luis
Valley.

��-17-

STUDIES OF CANADA GOOSE
POPULATIONS IN COLORADO TRANSPLANT
Michael

AREAS

R. Szymczak

P. N. OBJECTIVES
1.

To document the breeding
central Colorado.

range of Canada

2.

To document the distribution of harvest and estimate the survival
rate of the resident foothills Canada goose population.

3.

To document the distribution
the San Luis Valley.
SEGMENT

of harvest

geese wintering

of Canada

in north

geese nesting

in

OBJECTIVES

1a.

Plot the banding locations of Canada geese (Branta canadensis)
banded outside north central Colorado and reported recovered in
north central Colorado.

b.

Participate in the cooperative
southern Alberta.

2a.

Trap and band 500 Canada geese on production
along the foothills north of Denver.

and moulting

b.

Complete,
cards.

schedules

3a.

Trap and band at least 200 Canada
areas in the San Luis Valley.

b.

Complete,
cards.

Canada

submit and file appropriate

banding

geese on production

submit and file appropriate

METHODS

goose banding

banding

operation

in

areas

and recovery

and moulting

schedules

and recovery

AND MATERIALS

Band recoveries listed on computer printouts provided periodically by the
Migratory Bird Populations Station were sorted and plotted as to banding
and recovery location, sex, and age at time of banding, and estimated
breeding area for birds associated with north central Colorado.
Banding of flightless adults and goslings was accomplished through drive
trapping operations in southern Alberta, north central Colorado and the
San Luis Valley of Colorado.
The banding in Alberta was accomplished by
representatives
of state, provincial and federal conservation agencies
including Colorado.

�,...18-

Banding schedules and recovery reports pertaining to the San Luis Valley
and north central Colorado were completed and submitted to the Bird
Banding Laboratory.
Record keeping for the Alberta operation was the
responsibility of the Alberta Fish and Wildlife Division.
RESULTS AND DISCUSSION
Hi-Line

Population

Foreign Recoveries
During the 1976-77 hunting season a record 300 Canada geese banded outside
of Colorado were reported recovered in north central Colorado (Table 1),
of which 153 were classified as wild trapped locals when banded (Table 2).
The third year of recovery information resulting from banding in southern
Alberta indicated that Bantry No. 2 seems to produce geese that are
members of both the Pacific Flyway Rocky Mountain population and the HiLine population.
Six of 191 geese banded at Bantry in 1976 were recovered
in north central Colorado.
Geese from Bantry had been recovered predominantly in the Pacific Flyway in the past, with an occasional recovery in
the Hi-Line area. A significant number of wild trapped locals from the
Edmonton-Camrose
area are beginning to appear in the north central Colorado
harvest indicating that area is definitely becoming a Hi-Line production
area. Nesting areas east of 1110 longitude in southern Alberta, Murray,
Seven Persons, Cavan and Schuler continued to be represented in the north
central Colorado harvest.
A few birds were recovered that were supposedly
banded at Scope Reservoir, however, conflicting information indicates
those birds may not have been banded at Scope.
Birds banded in Saskatchewan originated mostly, as in the past; from areas
located in the three degree blocks (49-107, 49-108, 49-109) in southwestern
Saskatchewan.
Banding programs in that area have been terminated, therefore,
the number of recoveries of birds from that area should be decreasing.
Birds transported and released at Manito Lake and near North Battleford
in east central Saskatchewan were also noted in the harvest in north central
Colorado.
Geese from Phillips County, Montana, a traditional Hi-Line breeding area
were well represented in the harvest.
In addition, birds banded along
the Big Horn and Yellowstone Rivers in Treasure, Rosebud, Custer, Prairie,
Dawson and Big Horn counties in Montana were reported taken in north
central Colorado.
Geese from restoration areas in Weston, Crook and
Sheridan counties in northeast Wyoming were represented in the harvest.
No
other known production areas were represented in the north central Colorado
harvest to any great extent.
Geese banded at the Wheatland Reservoir moulting area in Albany County,
Wyoming were again represented in the harvest, as were birds banded as
moulters in the Liverpool Bay area in Mackenzie, Northwest Territory.
The
latter birds are small Ganada geese, persumably members of the shortgrass
prairie population, banded by Dr. Tom Barry, Canadian Wildlife Service,
incidental to the banding of white-fronted geese (Anser albifrons).

�-19-

Table 1. Banding areas outside Colorado of Canada geese recovered
north central Colorado during the 1976-77 hunting season.

Area

in

Direct

Indirect

14
0
0
6
3
0
5
10
3
0
16
1
1
1
0
1
0
1

9
1
11
0
4
2
3
3
3
1
4
1
0
0
1
0
1
0

0
0
0
0
4
0
0
0
0
0
0

3
2
2
5
0
6
5
1
1
2
1

0
20

3
5

Alberta
Edmonton-Camrose-Vegreville
Area
Kenilworth Lake
Dowling Lake
Bantry 112
Scope Reservoir
Grassy Lake
Murray Lake
Seven Persons
Cavan Lake
Many Islands Lake
Schuler Lake
Milk River Ridge Reservoir
Twelve Mile Coulee
Taber Lake
Fincastle Lake
Unnamed, E. of Scope Reservoir
Louisiana Lake
Bassano Area
Saskatchewan
Cypress Hills (Maple Creek)
Cypress Hills Area
Pontiex-Notukeu R.
Vidora Area
Battleford Area
Manito Lake Area
Val Marie-Frenchman
River Area
East of Quill Lakes
Kiyiu Lakes
Eastend Area
Regina Area
Mackenzie-Northwest
Liverpool

Territory

Bay Area

Smoke River Delta
Harrowby Bay

�-20-

Table 1. Banding areas outside Colorado of Canada geese recovered in
north central Colorado during the 1976-77 hunting season (continued).

Direct

Area

Indirect

Central Flyway
Montana
Phillips County
Treasure County
Garfield County
Valley County
Rosebud County
Prairie County
Custer County
Big Horn County
Dawson
County

20
4

30
1

1

2

o

1

1

1

1

o

7

4
1

o
3

o

o

2

o

1

o
o
o

1
6
3
4

North Dakota
Adams County
South Dakota
Stanley County
Wyoming
Platte County
Albany County
Goshen County
Crook County
Weston County
Sheridan County
Fremont County

1

3
3

o

o

1
1

o

2

o

2

4
2

3

o

5

Nebraska
Perkins

County

Kansas
Phillips

County

New Mexico
Mora County
San Miguel County
Socorro County

2

--------------------------------------------------------------------------

�-21-

Table 1. Banding areas outside Colorado of Canada geese recovered in
north central Colorado during the 1976-77 hunting season (continued).

Area

Central Flyway

Direct

Indirect

1
1

o

o

1

o

1

o

1

o

4

o

2

o
o
o

1

138

162

(cont.)

Texas
Randall County
Deaf Smith County
Mississippi

1

Flyway

Minnesota
Olmsted County
Wisconsin
Brown County
Missouri
Charito County,

Swan Lake

Pacific Flyway
Utah
Box Elder County
Oregon
Harney County
Idaho
Fremont County
Oneida County
Caribou County

Total

1
1

�-22-

Table 2. Banding locations
locals which were recovered
hunting season.

of Canada geese classified as wild trapped
in north central Colorado during the 1976-77

Year of Banding
Before
1970

Area

1970

1971

1972

1973

1974

1975

1976

7

9

3
1

5

Alberta
Edmonton-Camrose
Dowling Lake Area
Bantry 112
Scope Reservoir
Grassy Lake
Murry Lake
Seven Persons Lake
Cavan Lake
Many Islands Lake
Milk River Ridge Reservoir
Taber Lake
Fincastle Lake
Unnamed, E. of Scope
Bassano Area

2
1

2
1

1
3
1

2
1

3

4

9
3

1

2

2

16
1

1
1

1
1

Saskatchewan
Cypress Hills (Maple Cr.)
Cypress Hills Area
Pontiex-Notukeu
R. Area
Vidora Area
1
Val Marie-Frenchman
River Area
E. of Quill Lakes
Eastend Area

1
1

1

2
2

2

3

1
2

Central Flyway
Montana
Phillips Co.
Treasure Co.
Custer Co.
Big Horn Co.
Rosebud Co.
Dawson Co.

1

1

1

8

1

5

5

1
1
1
1

North Dakota
Adams Co.

14

1

2

---------------------------------------------------------------------------

�-23-

Table 2. Banding locations
locals which were recovered
hunting season (continued).

of Canada geese classified as wild trapped
in north central Colorado during the 1976-77

Year of Banding
Before
1970

Area

Central Flyway

1970

1971

1972

1973

1974

1975

1976

(continued)

Wyoming
I

Fremont Co.
Sheridan Co.
Goshen Co.

I
I

Pacific Flyway
Utah
1

Box Elder Co.

I

Oregon
2

Harney Co.
Idaho
I

Fremont Co.
Oneida Co.
Caribou Co.

Total

I
I

2

I

2

8

9

28

35

68

Banding
Banding was conducted for the fourth year on production areas in north
central Colorado.
A total of 582 geese were banded, bringing the four year
combined total to 3,075 (Table 3). Geese were captured at 13 locations
with 51.9 percent of the birds banded being goslings (Table 4).

�-24-

Table 3. Number of Canada geese banded on production
central Colorado 1974-1977 .

areas in north

Year

Adults

Goslings

Total

1974

751

315

1,067

1975

406

195

601

1976

357

468

825

1977

280

302

582

Table 4. Location and number, by age and sex, of Canada geese banded on
production areas in north central Colorado, 1977.

Location

Males

Adults
Females

Goslings
Fema l.es
Males

Total

11

11

13

10

45

11

9

5

10

35

Faivoir Ponds

5

3

16

14

3~/

Johnson's

2

0

0

0

2

Grey Lakes

7

8

21

19

55

Cobb Lake

27

26

0

0

53

Rocky Ridge Reservoir

0

0

4

3

7

Fossil Creek Reservoir

9

10

14

20

53

No. 8 Annex

26

22

0

0

48

Stone Mtn. Studios

3

2

12

4

21

Kitchel

Reservoir

5

7

17

16

45

McNeil Reservoir

24

13

41

39

117

Boedecker

17

21

10

14

62

147

132

153

149

582

Loveland
Va1mont

Total

(Ready Mix)
Reservoir

Pit

Reservoir

1/ Includes one adult of unknown sex.

�-25-

San Luis Valley Population
The attempt to band Canada geese on production areas in the San Luis Valley
in 1977 was essentially unsuccessful.
A total of 37 birds were captured
on three areas: Monte Vista National Wildlife Refuge, Sheep Pond and the
Rio Grande Management Area.
Thirty-three of the birds captured were
goslings.
We have been successful in reaching our quota of 200 birds in
only one of four years (Table 5).

Table 5. Numbers of Canada geese banded on production
in the San Luis Valley, 1974-1977.

and moulting

areas

Total

Year

Adults

Goslings

1974

4

6

6

1975

14

5

19

1976

55

149

204

1977

4

33

37

Prepared

by

71/.M'P k-'al
Michael R. SzynlC
Wildlife Researcher

C

��October

-27-

JOB PROGP~SS

State of

1978

REPORT

COLORADO
------~~~~~---------

Project No.

Migratory

W-88-B-23

9
Job No.
2
Monitor Banding of the Shortgrass Prairie
Canada Goose Population in Southeastern Colorado

~~~~-------------------------

Work Plan No.
Job Title

Period Covered:
Personnel:

Bird Investigations

January

23, 1978 through March 31, 1978

B. Dupire, M. Elkins, G. Nugent, M. Potter, E. Prenzlow, J.
Slater, J. Stevenson, M. Szymczak, J. Tanke and K. Wagner.

ABSTRACT
The trapping and banding effort directed at Canada geese (Branta canadensis)
in southeast Colorado resulted in 237 birds being banded~
The distribution
of harvest through the 1976-77 hunting season of Canada geese banded in
southeastern Colorado indicated the provinces of Alberta and Saskatchewan
and the state of Colorado continue to be the major harvest areas.

��-29-

MONITOR BANDING OF THE SHORTGRASS PRAIRIE
CANADA GOOSE POPULATION IN SOUTHEASTERN COLORADO
Michael

R. Szymczak

P. N. OBJECTIVE
To continually document, through monitor banding and analysis of recovery
data the annual and long term status of the southeastern Colorado (Arkansas
Valley) segment of the shortgrass prairie Canada goose population.
SEGMENT OBJECTIVES
1.

Band a m~n~mum of 1,000 Canada geese in southeastern
post-season period.

Colorado

2.

Prepare and submit banding
and progress reports.

and return reports,

3.

Analyze band recovery data for geese banded in southeast Colorado to
determine (1) distribution of harvest, (2) recovery rate, and (3)
survival rate.

schedules,

band recovery

during the

METHODS AND MATERIALS
Trapping

and Banding

All birds banded in southeast Colorado in 1978 were captured with baited
cannon nets, and the age and sex determined through cloacal and tail feather
examination.
All banding schedules, including recapture information, were
submitted to the U. S. Fish and Wildlife Service's Bird Banding Laboratory,
Patuxent, Maryland.
Because of work schedule conflicts, banding and recovery tapes were not
requested from the Bird Banding Laboratory.
Information on the distribution
of harvest was obtained from computer printouts that are provided periodically
by the Bird Banding Laboratory.
RESULTS AND DISCUSSION
Trapping crews again encountered the same difficulties in 1978 that were
present the previous year.
Two Buttes Reservoir, the primary trap site, was
not being used by large numbers of Canada geese, probably because of low water
levels.
Permission to trap at Turk's Pond, the major roosting site, could not
be obtained.
Trapping efforts were moved to Nee Noshe Reservoir, located
about 45 miles north of Two Buttes Reservoir.
Trapping efforts at Nee Noshe
resulted in the capture and banding of 237 geese: 101 adult males, 23
immature males, 83 adult females and 29 immature females.

�-30-

Distribution

of Harvest

The recovery distribution of shortgrass pralrle geese banded in southeastern
Colorado (Table 1) has not changed significantly since the information was
summarized through the 1972-1973 season (Szymczak 1974). The major change
of note has been the steady increase in the percent of shortgrass birds
taken in the Hi-Line population area in north central Colorado.
The major
harvest areas are still the provinces of Saskatchewan and Alberta, and
Colorado.
The Nebraska and Texas Panhandles remain as harvest areas of
note but the percent of take in Nebraska fluctuates considerably.

LITERATURE

CITED

Szymczak, M. R. 1974. Arkansas Valley Canada goose flock management
studies.
Colo. Div. of Wildl., Game Res. Rept., Fed. Aid Proj.
W-88-R.
October.
p. 55-87.

Prepared

by

21U.J1?~
Michael R. Szymczak
Wildlife Researcher

C

�Percentages

Table 1.

of total band recoveries,

1951-55

Area

Arkansas

Valley post-season

Five Year Averages
1966-70
1961-65
1956-60

bandings,

by area and year of recovery,
Recover;t:Year
1974-75
1973-74

1971-75

1971-72

1972-73

0.5
2.4
0.5
3.3

2.8
4.2
1.4
8.5

0.4
5.3
1.3
7.1

0.9
7.9

17.6
16.9

16.0
14.7
1.3

19.1
17.3

2.2

1.8

Far North Above 530
N.W. Territories
Alberta
Saskatchewan
Total
Provinces

-

-

-

9.8

7.1

10.2

9.1

1.0
5.0
0.8
6.8

28.6
18.3
0.1

35.3
10.5

28.9
11.5

19.0
21.8
0.8

15.6
18.1
0.6

11.9
21.8
0.5

0.8
0.1
0.7
0.1
5.4

0.4

0.3

0.6

0.5
0.3
5.8

0.7
0.3
0.5
0.3
6.3

1.3
0.1
1.2
0.5
5.0

1.9
1.0
1.4

-

-

Below 530

Alberta
Saskatchewan
B.C., Manitoba,

Ontario

Central F1;t:wa;t:
Montana
North Dakota
Wyoming
South Dakota
Nebraska
Colorado
Southeast and Other
North Central
Total
Kansas
Oklahoma
New Mexico
Texas
Panhandle
Waggoner Ranch
Gulf Coast
Total
Pacific

Fl;t:\vay

MississiEEi

F1ywa;t:

Mexico

Total Number
Of Recoveries

6.5

5.2

8.8

all bandings.

1975-76

1976-77

Total
Number
Recoveries

0.9
8.6
0.9
10.4

1.3
4.4
0.6
6.3

333

8.6

16.2
21.6
0.9

17.6
19.5
0.6

967
644
10

25.0
16.7
0.3

28
5
26
11
226

0.7
0.1
0.7
0.3
5.8

0.6
0.7
10.6

2.7
0.4
2.2

0.9
0.9
5.1

37.8
5.8
43.6
1.8
0.9

27.4
7.3
34.7

0.9
1.9

8.2

26.8
6.4
33.2
0.9
3.7
2.7

27.7
7.5
35.2
1.9
0.6
1.9

31.4
0.2
0.4
0.4

44.6
1.9
46.5
1.0
0.5
0.5

24.6
5.6
30.3

4.6
1.5
0.5
6.6

5.5
1.2
0.8
7.5

5.6
1.0
0.6
7.2

2.4
1.0

5.3
1.3

7.8
0.9

6.4
0.9

5.7
1.3

3.4

8.5
0.7
2.8
12.0

6.7

8.7

7.3

7.0

0.8

0.7

0.9

1.4

1.4

0.4

0.3

0.1

-

-

-

-

25.5
0.4
0.9
1.5

30.9
0.3
0.3
0.7

28.4
0.5
0.4
0.4

2.3
2.0
0.8
5.1

2.8
2.2
0.2
5.2

2.4

2.2

0.1

-

1.4
0.7

-

1.8

0.9
0.6

0.9

1,218
18
29
37

677

770

712

799

211

142

225

113

108

159

31.5
0.5
0.8
1.0

166
56
21
248 .

4.3
1.4
0.5
6.4

59

1.5

5

0.1

1

0.02

0.1

748

I

w
•....
I

34.3
5.1
39.4
0.9
0.1
0.6

-

% of Total
Recoveries

3,865

��-33-

JOB PROGRESS

October
REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Work Plan No.

Job Title
Period

Covered:

Personnel:

Migratory

W-88-R-23

1978

Bird Investigations

7
Job No.
3
Population Characteristics of
Mallards Wintering in West Central Colorado

---------------------------------

April 1, 1977 to March 31, 1978

G. Bock, J. Corey, J. Ellenberger, J. Frothingham, J. Gray,
J. Gumber, J. Leslie, D. Miller, J. Miller, S. Steinert, M.
Stone and R. Hopper.

ABSTRACT
The fifth consecutive year of the study of wintering mallards in west
central Colorado was completed during Segment 23. The aerial survey in
January yielded an estimated winter population of 13,860 birds, considerably
less than the two previous two years but equal to the long-term average.
Ground counts of nearly 5,300 birds produced a sex ratio of 114.5 males:lOO
females.
This ratio was higher than those of 99.2:100 and 107.5:100
obtained in 1976 and 1977, respectively.
Almost 2,400 mallards were banded
during the postseason period in 1978, bringing the total banded sample to
8,352 for the five years of study (1974-1978).
Only 271 recoveries have
been received from these bandings through March 1, 1978, which actually
represents less than four complete years of recovery information.
Between
57 and 68 percent of these recoveries, depending upon age and sex class,
occurred in the western portion of Colorado.
Canada, Idaho and Utah were
the locations for most of the remaining recoveries.

��-35-

POPULATION CHARACTERISTICS OF
MALLARDS WINTERING IN WEST CENTRAL COLORADO
Richard M. Hopper

This report presents results of the fifth consecutive year of study of
the mallard population wintering in west central Colorado.
Included is a
summarization of all data collected since initiation of this investigation
during the 1973-74 Segment year.

PROGRAM NARRATIVE

OBJECTIVE

1.

To estimate population parameters of mallards wintering in west central
Colorado by age and sex category; specifically (1) population size,
(2) recovery rates, (3) survival rates, (4) mean life span, (5) sex
ratio, and (6) geographic distribution of the harvest.

2.

To develop a management plan for the population
in west central Colorado.

of mallards

wintering

SEGMENT OBJECTIVES
1.

Conduct an aerial count of wintering numbers of mallards during the
first or second week of January as part of the annual mid-winter survey
conducted throughout the United States.

2.

Conduct a minimum of two ground counts (sex ratio) of 500 mallards
each in each of the following two concentration areas: (1) Grand
Junction-Highline
Lake area, and (2) Montrose-Delta area.

3.

Trap and band 1,500 mallards during the post season period, including
750 in the Grand Junction-Highline
Lake area, and 750 in the MontroseDelta area. Distribute the sample equally among the four age and sex
classes.

4.

Preparation and submission of banding
return reports, and progress reports.

schedules,

band recovery

and

METHODS AND MATERIALS
All procedures remained essentially the same as in Segment 22 (Hopper 1977)
The aerial count was made on January 9, 1978 and involved flying over all
known waterfowl concentration areas in the Uncompahgre-Gunnison-Colorado
River Complex of west central Colorado.
Personnel of the Northwest Region
again conducted the count, utilizing a Cessna 185 aircraft and a crew
consisting of a pilot and two observers.
Sex ratio counts were made during the period January 28-February 1 in the
Grand Junction-Highline
Lake area, and during the period February 8-10 in
the Montrose-Delta area.
In the northern location, sex ratio counts were

�~36-

conducted at Highline Lake, Walker Wildlife Area and the Colorado River
west of Grand Junction.
Sex ratio counts were obtained only at Sweitzer
Lake near Delta in the southern area.
Salt Plains type traps, baited with corn, were employed to capture mallards
for banding.
Banding was done during the period January 25-February 12.
All birds were aged according to the wing-aging technique (Carney 1964),
and recapture data were maintained for each banding location.
Banding
schedules and recovery reports were prepared and submitted to the Bird
Banding Laboratory.
RESULTS AND DISCUSSION
Aerial Census
Table 1 shows the results of the 1978 January inventory of wintering
mallards, as well as the counts from the previous four years.
The 1978
estimate was 13,860 birds, which was lower than the 1976 (23,125) and 1977
(16,280) figures, but higher than the first two years of the study.
The
long-term (1959-1978) average for this population is 13,493.
The area
covered in the inventory was presented in more detail in the last report
for this job (Hopper 1977).

Table l. January
1974-1978.

Location

Inventory

of mallards

wintering

in west central Colorado

1974

1975

1976

1977

1978

Highline

Lake

2,400

4,700

10,200

6,900

6,100

Colorado

River

3,110

1,830

1,545

3,390

3,046

700

1,345

9,100

2,215

3,756

0

485

270

225

208

0

0

1,900

1,800

450

2,975

3,400

110

1,750

300

9,185

11,760

23,125

16,280

13,860

.

II

Uncompahgre

River

Gunnison

River-

Sweitzer Lake
Burlingame

Total

II Includes

Pond

North Fork of Gunnison

River to Hotchkiss.

�-37-

Sex Ratio Counts
Results of sex ratio counts in 1978 are shown in Table 2 by date and
location.
Overall ratios obtained in 1976 and 1977 are also presented
for comparison.
Nearly 5,300 mallards were counted in 1978, producing a
sex ratio of 114.5 ma1es:100 females for all areas combined.
The northern
and southern portions produced nearly identical ratios.
For some unknown
reason, the proportion of males in the sample counts appeared to have
steadily increased since initiation of these counts in 1976, with ratios
of 99.2, 107.5 and 114.5 males per 100 females, respectively, for the
years 1976 through 1978.

Table 2. Sex ratio counts of mallards in west central Colorado during
winter of 1978, with overall ratios from 1976 and 1977 for comparison.

Location

Date

No.
Males

No.
Females

Total

Males:
100 Females

01-28-78

282

241

523

117.0

01-29-78

288

250

538

115.2

01-31-78

272

232

504

117.2

01-30-78

266

254

520

104.7

01-31-78

279

240

519

116.2

02-01-78

272

236

508

115.2

01-30-78

88

76

164

115.8

1,747

1,529

3,276

114.2

02-08-78

278

222

500

125.2

02-09-78

258

242

500

106.6

02-10-78

261

239

500

109.2

02-10-78

272

228

500

119.3

1,069

931

2,000

114.8

Total (1978)

2,816

2,460

5,276

114.5

1976

1,741

1,755

3,496

99.2

1977

1,375

1,279

2,654

107.5

High1ine

Lake

Walker Wildlife
Area

Colorado

River

Subtotal

(North)

Swietzer Lake

Subtotal

(South)

the

�-38-

Trapping

and Banding

Trapping was continued during the 1978 post season period (January and
February), representing the fifth consecutive year of banding effort ..
Nearly 2,400 mallards were banded in the two general areas, bringing the
total banded sample for the five years (1974-1978) to 8,352 (Table 3).
The age composition of the sample in 1978 was about even for adults and
subadults, while for all years combined it is still in favor of subadults.
The adult female segment continued to be the least represented age and sex
class in the overall sample, although some years it was as common as the
other three. The 1978 sample ran high to the male portion of the population,
which is typically the situation when using the Salt Plains type trap.

Table 3. Number and age and sex composition of mallards banded postseason
in west central Colorado, 1978, and totals for the period 1974-1978.

AM

Age and Sex
SM
AF

SF

Total

Grand Junction Area

505

413

302

260

1,480

Delta Area

247

303

128

238

916

752

716

430

498

2,396

1,304

1,049

851

1,061

4,265

960

1,327

656

1,144

4,087

2,264

2,376

1,507

2,205

8,352

Year and Location

1978

Total
1974-1978
Grand Junction Area
Delta Area

Total

Preliminary

Distribution

of Band Recoveries

As of March I, 1978, only 271 recoveries had been obtained from the first
four years of banding.
These recoveries represented only those birds that
had been shot and/or found dead during the hunting season (September 1January 31). Actually, these recoveries are for less than four complete
years, since it cannot be assured that all recoveries from the 1977-78
hunting season are processed until September or October of 1978. This
simply means that in postseason banding studies there is about one and onehalf years of lag between time of banding and the time that all recoveries
from those bandings are fully processed.

�-39-

The recovery distribution of the 271 recoveries received thus far is
presented in Table 4 by age and sex class.
The Pacific Flyway portion of
Colorado was the major recovery location for all four age and sex classes,
accounting for from 56.7 to 68.3 percent of the total recoveries.
Adult
females showed a somewhat higher affinity for this area than did the other
three classes.
Idaho and Canada were the next most common recovery
locations for adult males, while Canada and Utah accounted for most of the
remaining recoveries of subadult males and both ages of females.
Few
recoveries occurred in the eastern portion (Central Flyway) of Colorado;
less than five percent of the total recoveries of each age and sex class.

Table 4. Recovery
Colorado 1974-1977

distribution of mallards
(all years).

banded postseason

in western

Percent
AM

of Total Recoveries
Age and Sex
SM
AF

Alberta

7.1

8.1

9.8

10.0

Saska t chewan

2.9

3.0

2.4

6.7

Subtotal

10.0

11.1

12.2

16.7

Colorado

57.1

58.6

68.3

56.7

Idaho

11.4

5.1

2.4

6.7

Montana

1.4

3.0

0.0

0.0

New Mexico

2.9

0.0

0.0

0.0

Oregon

0.0

1.0

2.4

0.0

Utah

5.7

7.1

4.9

8.3

Washington

0.0

0.0

2.4

0.0

Wyoming

4.3

2.0

2.4

5.0

Subtotal

82.8

76.8

82.8

76.7

Colorado

2.9

4.1

0.0

3.3

Kansas

0.0

2.0

0.0

1.7

Montana

1.4

0.0

2.4

1.7

Nebraska

0.0

1.0

0.0

0.0

New Mexico

0.0

1.0

0.0

0.0

Recovery

Location

SF

Canada

Pacific Flyway

Central Flyway

----------------------------------------------------------------------------

�-40-

Table 4. Recovery distribution of mallards banded postseason in western
Colorado 1974-1977 (all years).

Recovery Location

AM

Percent of Total Recoveries
Age and Sex
SM
AF

SF

Central Flyway (continued)
Texas

1.4

1.0

0.0

0.0

Wyoming

0.0

2.0

0.0

0.0

Subtotal

5.7

11.1

2.4

6.7

1.0

0.0

0.0

Louisiana

0.0
1.4

2.4

0.0

Subtotal

1.4

0.0
1.0

2.4

0.0

Total

99.9

100.0

99.8

100.1

Number recoveries

70

99

41

611:/

Mississippi Flyway
Iowa

.
f one C01orad0 recovery was not d·
d
-1/ Spec~·f·
~c 1ocat~ono
eterm~ne,
percentages were calculated on the basis of only 60 recoveries.

so above

LITERATURE CITED
Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification by means of wing plumage. U. S. Fish and Wi1d1. Serv., Spec.
Sci. Rept., Wildl. No. 82. 47 pp.
Hopper, R. M. 1977. Population characteristics of mallards wintering in
west central Colorado. Colo. Div. of Wi1d1., Fed. Aid Game Res. Rept.,
Oct. pp. 33-39.

Prepared by ----6,.q-7f}--&lt;L"'""~::=....~~·~"'9"-~~+~_·+-~-Y.'I;f=-Ae-,~.
_
~hard
M. Hbpp:r~
Wildlife Researcher

�October

-41-

1978

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Work Plan No.
Job Title

Migratory

W-88-R-23

Bird Investigations

Job No.
8
Monitor Banding of Eastern
Colorado Wintering Mallard Populations

----------~~-------------------

3

----------~~~~~~~~~~~~~~~~~~--------------

Period Covered:

April 1, 1977 to March 31, 1978

Personnel:
M. Babler, G. Brown, L. Budde, J. Carsella, A. Chappell, D. Coven,
J. Corey, C. Crawford, M. DePra, B. Dupire, K. Dillinger, M. Gardner, B.
Goforth, J. Jackson, J. Jones, B. Leasure, R. Lopez, J. Lorentzson, T.
Lynch, F. Marcoux, T. Martin, K. Moser, R. Oehlkers, J. Pogorelz, F.
Rinella, C. Roberts, L. Rottman, W. Rupke, H. Spear, J. Sweeting, J. Tanke,
E. Wagner, K. Wagner, J. Young, R. Zaccagnini, and R. Hopper.

ABSTRACT
The transfer of field operations for the monitor banding program in eastern
Colorado from research to management was completed during Segment 23.
Regional personnel did all of the actual banding work, except at Bonny
Reservoir.
Research personnel contributed by holding an orientation
meeting for all banders, which included a review of the instructions manual
and training session on aging techniques.
This year's banded sample
totaled over 5,600 mallards, which was in excess of the 4,000 minimum quota
established.
The age composition of the sample was low, especially for
females. An updated analysis of the recovery data will be conducted next
segment.

��-43-

MONITOR BANDING OF EASTERN
COLORADO WINTERING MALLARD POPULATIONS
Richard M. Hopper

This report presents results of the postseason mallard banding program
in eastern Colorado during Segment 23 and discusses the transfer of this
monitoring activity from a research to a management function.
Hopper
(1977) described the purpose of the original study, general accomplishments, reasons for transferring field responsibilities
to the regional
management system, and role of the research section in the program.

P. N. OBJECTIVES
1.

To establish monitor banding of wintering mallard populations
eastern Colorado as an annual management function.

in

2.

To continually document, through monitor banding and analysis of
recovery data, the annual and long-term status of eastern Colorado
wintering mallards to provide a basis for annual hunting season
recommendations.
SEGMENT OBJECTIVES

1.

Instruct management personnel in the field conduct of a postseason
mallard banding program by (a) holding wing-aging classes, and (b)
assisting and training management personnel in the field.

2.

Band a minimum of 4,000 mallards during the postseason period,
including a minimum of 500 birds in each of the following general areas
of the South Platte Valley and Arkansas Valley: (a) Denver-Greeley,
(b) Fort Collins-Loveland-Windsor,
(c) Greeley-Fort Morgan, (d) Fort
Morgan-Sterling,
(e) Sterling-Julesburg,
(f) Bonny Reservoir, (g)
Manzanola-Lamar,
and (h) Two Buttes Reservoir area. Divide the banded
sample in each area equally among the four age and sex classes.

3.

Conduct an updated analysis of band recovery data, including the
following major determinations for important population segments of
mallards wintering in eastern Colorado: (a) distribution of the harvest,
(b) recovery rates, and (c) survival rates.

4.

Prepare and submit banding
and progress report.

schedules,

band recovery

and return reports,

METHODS AND MATERIALS
Procedures and equipment remained essentially the same as in the previous
year (Hopper 1977), with the exception that less time was spent by research

�-44-

personnel (Federal Aid Project W-88-R) in field training of management
personnel.
An orientation meeting and review of the instructions manual
prepared last year were again conducted for the management personnel
involved.
RESULTS
Instruction

AND DISCUSSION

of Management

Personnel

The same management personnel selected for the eight banding crews last
year were again assigned to the 1978 crews.
Crew assignments by region
and unit remained the same as last year, as did responsibilities
of the
crew leaders and assistant crew leaders (Hopper 1977).
The orientation
meeting and review of the instruction manual covered all phases of the
trapping and banding operation as outlined in the last report, including
a training session on aging techniques.
Management personnel in the Northeast and Southeast regions did all of the
postseason banding in eastern Colorado in 1978 without field assistance
from the Research Section, except in the case of Bonny Reservoir.
Thus,
as a result of the success experienced in training management personnel
in trapping and banding operations over the past two winters, it can be
concluded that the transfer of monitor banding responsibilities
from
research to management is now complete.
To clarify responsibilities
in regard to this banding program, the Reseach
Section (Federal Aid Project W-88-R) will continue to: (1) hold the master
permit issued to the Colorado Division of Wildlife by the U. S. Fish and
Wildlife Service for the banding of migratory game birds, (2) issue subpermits to crew leaders and assistant crew leaders to cover their banding
activities, (3) order bands and issue them to the banding crews and
maintain an updated inventory of bands used and in stock, (4) supply
necessary banding and recapture forms, (5) fill out final banding schedules
and recovery and return reports and submit them to the Bird Banding Laboratory, (6) maintain all banding and recovery records, (7) periodically
update tge analysis of band recovery data and provide results to managers,
and (8) serve as a consultant on any technical aspects of the program.
Banding
Over 5,600 mallards were banded during the January-February
period of
Segment 23 (Table 1). As with last year, this total exceeded the 4,000
minimum quota by a significant amount.
This was due in part to the large
sample obtained at Bonny Reservoir.
Only one banding location, the Two
Buttes area, failed to contribute a good sample to the overall program.
All banding schedules and band recovery and return reports were prepared
and submitted to the Bird Banding Laboratory.
The age and sex composition of the banded sample differed from that of most
years, in that a low age ratio was obtained, especially for females.
Subadult females have nearly always outnumbered adult females in samples
of past years, but this year the reverse was true. This was basically a
reflection of low productivity in this population during the summer of 1977.

�-45-

Table 1. Mallards banded postseason by age and sex in the eight eastern
Colorado banding areas, January-February
1978.

Banding Area

AM

Number of Ducks Banded
Age and Sex
SM
AF
SF

Bonny Reservoir

717

380

348

149

1,594

Sterling-Julesburg

158

163

174

105

600

Ft. Morgan-Sterling

229

147

63

60

499

Greeley-Ft.

302

79

65

50

496

Denver-Greeley

295

257

234

205

991

Ft. Co11ins-LovelandWindsor

196

216

233

136

781

Arkansas

Valley

247

170

90

80

587

Two Buttes Area

38

23

6

25

92

2,182

1,435

1,213

810

5,640

Morgan

Total

Updated Analysis

of Band Recovery

Total

Data

Banding and recovery tapes were requested from the Bird Banding Laboratory
in mid-September 1977 for mallards banded postseason in eastern Colorado
during the period 1963-64 through 1975-76 and for recoveries from these
bandings during the 1964-65 through 1976-77 hunting seasons.
This request
was made this late in the Segment to insure that all recoveries from the
1976-77 hunting season had been processed and included in the recovery tape.
The tapes were not received until mid-January 1978, and then we were informed
in early March that the banding tape contained errors and that a new tape
would be forthcoming.
This new tape was received in early April following
the end of Segment 23. Thus, no work was accomplished on the banding
analysis objective outlined for this Segment.
This work will be undertaken
in Segment 24.

LITERATURE

CITED

Hopper, R. M. 1977. Monitor banding of eastern Colorado wintering mallard
populations.
Colo. Div. of Wildl., Fed. Aid Game Res. Rept., Oct.
pp. 41-47.

Prepared

by

:~d/~

P

/

t"

;;;!~:)-'4

Richard M. Hopper;;
Wildlife Researcher

•

��October

-47-

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Work Plan No.
Job Title
Period

Bird Investigations

9
3
J~ ~.
--------~----------------------Migration and Mortality Characteristics
of
D_u_c_k
__P_o_p~u
__l_a_t_i_o_n_s
__l_·n
__t_h
__e__I_n_t_e_r_-_M_o_u_n
__t_a_i_n
__V_a_l_l_e~y~s
o_f
__C_o_l_o_r_a_d_o _

Covered:

Personnel:

Migratory

W-88-R-23

1978

July 15, 1977 to March

31, 1978

M. Nail and Staff, Monte Vista National Wildlife Refuge; R.
Darnell and Staff, Alamosa National Wildlife Refuge; M. Conner,
J. Corey, D. Flenthrope, R. Hopper, R. Oakleaf, J. Wagner, K.
Wagner, and M. Szymczak, Colorado Division of Wildlife; K. Storey
and W. Riley, Colorado State University.
ABSTRACT

Totals of 3,877, 4,193 and 3,012 ducks were banded in North Park, South Park
and the San Luis Valley, respectively in 1977.
Gadwall, wigeon, shoveler,
redhead, and L. scaup made up 25.4, 14.9, 11.2, 10.1 and 9.7 percent of the
harvest, respectively, during the early duck season in North Park.
In South
Park the mallard, green-winged teal, gadwall and pintail made up 55.9, 13.2,
8.8 and 8.8 percent of the harvest during the early season.
In the San Luis
Valley during early seasons the mallard has been the major species of harvest
since 1963, but its importance, according to percentage in the harvest, has
declined steadily from 86.9 percent in 1963 to 43.7 percent in 1977.

��-49-

MIGRATION AND MORTALITY CHARACTERISTICS
OF DUCK POPULATIONS IN THE INTER-MOUNTAIN
VALLEYS OF COLORADO
Michael R. Szymczak

P. N. OBJECTIVES
1.

To investigate migration, mortality, recovery distribution and relationships among populations of selected species of ducks present in North
Park, South Park and the San Luis Valley during the mid-July through
mid-September period.

2.

To document the species composition of ducks harvested during early
October seasons, should seasons occur in North Park, South Park and the
San Luis Valley.

3.

To examine the feasibility of establishing early special duck seasons
in September in the high mountain park areas or to examine the feasibility of special early October seasons in South and North Park similar
to what has been recommended for the San Luis Valley.

4.

To establish procedures to monitor the effect of early special duck
seasons on duck populations in the high mountain park areas if granted.
SEGMENT OBJECTIVES

1.

Trap and band ducks in North Park, South Park and the San Luis Valley
during the mid-July through mid-September period as designated in
Program Narrative Outline.
Complete, submit and file appropriate
banding schedules and recovery cards.

2.

Collect and analyze data conce~ning the species composition of the
harvest during early October seasons in North Park, South Park and the
San. Luis Valley utilizing wing collection barrels and results of the
U. S. Fish and Wildlife Service's Parts Collection Survey.

3.

Analyze band recovery data through the 1976 recovery year for mallards,
pintail and green-winged teal banded during the pre-season period in
North Park, South Park and the San Luis Valley to determine the feasibility of requesting early seasons in the high mountain area.

4.

Prepare

5.

If a special high mountain duck season is requested and granted, design
a program for population monitoring which may include banding, harvest
surveys, wing collection or other activities.

progress

report.

�-50-

METHODS AND MATERIALS
Ducks were captured from August 6-September 7 in South Park, August 8September 14 in the San Luis Valley and August I-September 15 in North Park.
Primarily "salt plains" type bait traps were used (Szymczak and Corey 1976).
In North Park night lighting from airboats was used to capture most gadwall
and wigeon.
The age and sex of all birds captured and banded were determined.
Wings from ducks bagged during the October 1, 1977 through October 14, 1977
period were collected in North and South Parks through the use of voluntary
collection barrels (Hoffman and Braun 1975).
Barrels were placed at Walden
Rese!"voir (2 barrels), Lake John Annex (1 barrel) and at Cowdrey (1 barrel)
in North Park and at Antero Reservoir (3 barrels) in South Park.
Some
additional wings were collected at MacFarlane Reservoir.
All wings collected
were classified by species, age, sex and location, and in some cases, periods
of harvest.
The species composition of the harvest in the San Luis Valley
was obtained at the Central Flyway, U. S. Fish and Wildlife Service's Parts
Collection "wing bee." A computer print out listing the species, age, sex,
and time of harvest of birds whose wings were obtained through the Parts
Collection Survey for the years 1971 through 1976 was obtained for waterfowl
reported as harvested in North Park (Jackson County), South Park (Part
County) and the San Luis Valley (Saguache, Rio Grande, Alamosa, Conejos and
Costilla counties).
Banding and recovery tapes plus recapture and return information covering all
pre-season banding and resulting recoveries for ducks banded in North Park,
South Park, San Luis Valley and the high country areas west of the San Luis
Valley through the 1975-76 recovery year were received from the Bird Banding
Laboratory.
Recovery year 1976 was not included.
In addition, computer
tapes listing ducks recovered or recaptured in the above areas but banded
elsewhere was also received.
A tape containing seven programs for use in band recovery analysis was
obtained from the Northern Prairie Wildlife Research Station.
One program,
RCOUDIST, is described by Coward in (1977), five programs, EDITOR, RCOUTAB,
LLPLOT, BANDAREA and MAP are outlined by Davenport (1977) and the final
program BANDRCOU is documented by Coward in and Davenport (1973).
In addition,
the program SURVIVAL (Johnson 1974) was incorported into BANDRCOU.
All
programs with the exception of MAP were converted for use on the CDC 6400
and CYBER systems.
All converted programs except RCOUDIST and BANDAREA have
been utilized, however, only portions of the resulting outputs have been
analyzed.
RESULTS AND DISCUSSION
Totals of 3,877, 4,193 and 3,012 ducks were banded in North Park, South Park
and the San Luis Valley, respectively, in 1977. The species composition of
the birds banded in each area are presented in Table 1, 2 and 3. Quotas
of 300 mallards for each sex of adult and immature birds were met only for
males in North Park.
In North Park, green-winged teal quotas of at least
500 total birds were not achieved, while gadwall quotas of 300 of each sex
were met.
Slightly over 11,000 birds were banded in the three areas
(Table 4).

�-51.Table 1. Number of ducks banded, by species,
pre-season period, 1977.

in North Park during

Age and Sex
LM
AF

the

IF

LF

TOTAL

223

237

36

1,162

9

449

234

6

1,643

3

29

303

2

17

721

52

5

3

36

6

6

108

94

29

3

28

21

3

178

Blue-winged and
Cinnamon Teal

8

11

1

1

6

0

27

Redhead

7

0

12

13

0

6

38

1,494

659

91

1,053

506

74

3,877

AM

1M

Mallard

316

316

34

Pintail

650

295

Gadwall

367

American

Wigeon

Green-winged

Teal

Total

Table 2. Number of ducks banded by species,
pre-season period, 1977.

in South Park during

Age and Sex
LM
AF

the

IF

LF

TOTAL

191

175

12

897

1

200

128

3

1,000

0

0

23

0

0

39

AM

1M

Mallard

244

258

17

Pintail

509

159

Redhead

16

Ring-necked

Duck

3

0

0

2

0

0

5

Green-winged

Teal

985

417

0

357

259

0

2,018

125

30

0

62

17

0

234

1,882

864

18

835

579

15

4,193

Blue-winged and
Cinnamon Teal
Total

�-52-

Table 3. Number of ducks banded by species,
the pre-season period, 1977. 1/

in the San Luis Valley during

Age and Sex
LM
AF

IF

LF

Total

7

251

219

8

931

144

0

270

l32

2

965

473

141

0

112

63

0

789

Blue-winged and
Cinnamon teal

126

30

0

53

44

1

254

Redhead

l3

6

10

25

6

4

64

Canvasback

1

0

0

0

0

0

1

Gadwall

0

2

3

0

0

2

7

1

0

0

0

0

0

1

1,241

559

20

711

464

17

3,012

Wildlife

Refuge

AM

1M

Mallard

210

236

Pintail

417

Green-winged

American

Total

teal

wigeon

1/
- Includes 194 ducks banded by the Monte Vista National
and Alamosa National Wildlife Refuge.

Species Composition

of the Harvest

A total of 268 wings were collected during the early duck season in North Park
in 1977. About 62 percent of the wings were obtained at Walden Reservoir.
Gadwall was the major species of harvest (25.4 percent) in North Park for
the second consecutive year (Table 5). The American wigeon was second in
importance with 14.9 percent.
In 1975 and 1976 the gadwall and wigeon
combined comprised about 50 percent of the harvest (Table 6). Their combined
total in 1977 dropped to about 40 percent.
The redhead made up about 10
percent of the harvest for the second consecutive year. Wigeon and shovelers
were more prominent in the harvest at Walden Reservoir than in the other
three areas, while blue-winged and/or cinnamon teal were important in the
Cowdrey area.
The mallard comprised 6.0 percent of the harvest in 1977
compared to 6.1 percent in 1976 and 6.9 percent in 1975. Wigeon, shovelers
and pintail were harvested at a greater rate on opening weekend than during
the remainder of the period (Table 7). The importance of gadwall and redheads increased after the opening weekend.

�-53-

Table 4. Number of ducks banded, by species, in North Park, South Park and
the San Luis Valley during the pre-season period 1977 . 1)

AM

1M

Age and Sex
AF
LM

IF

LF

Total

Mallard

770

810

58

665

631

56

2,990

Pintail

1,576

598

10

919

494

11

3,608

1,552

587

3

497

343

3

2,985

Gadwall

367

5

32

303

2

19

728

Blue-winged and
Cinnamon teal

259

71

1

116

67

1

American

53

5

3

36

6

6

Redhead

36

6

22

61

6

10

Canvasback

1

0

0

0

0

0

3

0

0

2

0

0

4,617

2,082

129

2,599

1,549

106

11,082

Wildlife

Refuge and

Green-winged

teal

515
109

wigeon

141
1
5
Ring-necked

Total

1/

duck

- Includes 194 ducks banded by the Monte Vista National
Alamosa National Wildlife Refuge.

In South Park at Antero Reservoir, the mallard was by far the dominant species
of harvest (55.9 percent) throughout the early season in 1977 (Table 8). In
1976 the wigeon and the three species of teal comprised over 70 percent of the
harvest while the mallard made up only about 9 percent (Table 6).
The mallard was the major species of harvest (43.7 percent) in the San Luis
Valley in the early season in 1977, according to the 71 wings collected
through the parts collection mail survey (Table 6). The remainder of the
harvest was composed of primarily pintail, gadwall and green-winged teal.
The mallard has been the major species of harvest during early seasons in
the San Luis Valley since at least 1963 (Hopper et a1. 1975, Table 6). The
mallards importance, however, has been declining steadily, from 86.9 percent
of the harvest in 1963 to 43.7 in 1977. The importance of other species
has fluctuated from year to year, but generally gadwall, green-winged teal,
pintail, and possibly blue-winged and/or cinnamon teal have been increasing
in importance.

�-54-

Table 5. Species composition of the harvest in North Park during the
October 1-14, 1977 early duck hunting season according to wings voluntarily
placed in collection barrels.

Species

Walden
Reservoir

Lake John
Annex

Area
Cowdrey
Area

Gadwall

34 (20.5)-Y

15 (34.1)

American wigeon

34(20.5)

N. shoveler

MacFar1ane1/
Reservoir -

Total

6(18.2)

l3(52.0)

68(25.4)

2(4.5)

2(6.1)

2(8.0)

40(14.9)

24(14.5)

3(6.8)

1 (3.0)

2(8.0)

30(11.2)

Redhead

16(9.6)

10(22.7)

0(0.0)

1(4.0)

27(10.1)

Lesser scaup

19(11.4)

1(2.3)

4(12.1)

2(8.0)

26(9.7)

Blue-winged or
Cinnamon teal

5(3.0)

3(6.8)

l3(39.4)

2(8.0)

23(8.6)

Mallard

6(3.6)

6(l3.6)

4(12.1)

0(0.0)

16(6.0)

Pintail

11(6.6)

2(4.5)

1(3.0)

1(4.0)

15(5.6)

Green-winged teal

9(5.4)

0(0.0)

2(6.1)

2(8.0)

l3(4.9)

Ruddy

7(4.2)

0(0.0)

0(0.0)

0(0.0)

7(2.6)

Ring-necked duck

0(0.0)

2(4.5)

0(0.0)

0(0.0)

2(0.7)

Common merganser

1(0.6)

0(0.0)

0(0.0)

0(0.0)

1(0.4)

44

33

25

Total
1/
2/

166

Wings collected only on October 1-2, 1977 .
Percent of total in parentheses.

268

�Table 6. Percent species composition of the harvest in North Park, South Park and the San Luis Valley during early
October duck hunting seasons 1971-1977 based on U. S. Fish and Wildlife's Parts Collection Mail Survey and Independent Wing Barrel Survey.

Species

NP

Mallard
Gadwall
Green-winged teal
Wigeon
Blue-winged teal/
cinnamon teal
Pintail
Shoveler
Redhead
Ruddy duck
Ring-necked duck
Lesser scaup
Common merganser
H. merganser
Bufflehead
Unknown

33.3
33.3

Total Number of
Wings Examined

25.0
0.0
0.0
0.0
8.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

12

1971
SP

SLV

NP

66.7
0.0
0.0
0.0

65.6
6.1
7.1
3.0

18.8
18.8
6.3

0.0

1972
SP

1973
SLV

NP

50.7
14.1

21.1

6.9
20.7
41.4

31.3

0.0
50.0
0.0
50.0

0.0

3.4

0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

3.0
13.1
2.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

0.0
12.5
6.3
0.0
6.3
0.0
0.0
0.0
0.0
0.0
0.0

0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

1.4
5.6
5.6
0.0

0.0
20.7
0.0
0.0
0.0
0.0

9

99

16

2

71

33.3

1.4
0.0
0.0
0.0
0.0
0.0
0.0

SP

NP

60.6

14.3

9.1

50.0
0.0
0.0

2.0
2.0

3.4
0.0
0.0
0.0
0.0

29

SLV

a

1974
SP

SLV

75.0
0.0
0.0
25.0

47.7
10.2
11.4

15.9

88

9.1
11.1
5.1
1.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

7.1
21.4
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

99

14

4

-------------------------------------------------------------------------------------------------------------------

3:4
3.4
8.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

I
VI
VI

I

�Table 6. Percent species composition of the harvest in North Park, South Park and the San Luis Valley during early
October duck hunting seasons 1971-1977 based on U. S. Fish and Wildlife's Parts Collection Mail Survey and Independent Wing Barrel Survey (continued).
1975
SP

SLV
47.1

NP

1976
SP

SLV

NP

1977
SP

SLV

0.0

66.7

44.9

22.2

63.6

43.7

6.0

55.9

6.7

22.2

0.0

25.4

8.8

Species

NP

Mallard

27.01/

66.7

6.9-

-

-

6. 1

8.7

Gadwall

13.5

33.3

20.6

53.3

0.0

Z8.1

2.2

0.0

16.7

7.2

25.0

20.6
Green-winged teal

8.1

0.0

4.4

7.7
Wigeon

16.2

0.0

2.9

28.9
B1ue-winged/
cinnamon teal
Pintail

10.8

0.0

5.9

15.4
8.1

0.0

11.8

5. 1
Shoveler

0.0

Redhead

5.4

Ring-necked duck

0.0

Lesser scaup

8.1

Common merganser

0.0

H. merganser

0.0

Bufflehead

0.0

Unknown

0.0

6.7

0.0

8.8

25.0

13.3

16.7

4.8

3.3
0.0

0.0

2.7

2.2

0.0

1.5

13.3

0.0

11.2

5.4

0.0

0.0

0.0

0.0

1.6

0.0

0.0

0.0

0.0

0.0

0.0

0.0

1.6
0.2
5. 1
0.0

0.0

0.0
0.0

1.5

0.0

0.0

0.0
0.0
0.0

28.3

4.4

4.3
Ruddy duck

0.0

21.4

0.0

3.4
2.7

13.3

0.0

0.0

1.1

0.0

0.0

0.0

7.0

0.0

0.0

0.0

0.0

1.1

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

19.1
4.5
13.5
7.9
2.2
0.0

11.1

0.0

4.9

4.9

0.0

27.3

14.9

4.4

0.0

0.0

8.6

2.9

0.0

0.0

5.6

8.8

11.1

9.1

11.2

0.0

0.0

0.0

10.1

2.9

0.0

0.0

0.0

2.6

1.5

1.1

0.0

0.0

0.0
0.0
0.0
0.0
0.0

0.0

89
6
15
68
3
37
Total No. Wings
9Z
374
494
Examined
1/ Italic numbers indicates figures obtained from wings collected in wing barrels.

0.7

0.0

22.2

0.0

9.7

0.0

0.0

0.0

0.4

1.5

0.0

0.0

0.0

0.0

11.1

0.0

0.0

0.0

0.0

0.0

0.0

0.0

9

11

268

68

12.7
12.7
7. a
1.4
I
Vl

16,9
4.2
0.0
0.0
1.4
0.0
0.0

0.0
0.0
0.0
71

cI

�-57-

Table 7. Species composition of the harvest by time period at Walden Reservoir, Lake John Annex and the Cowdrey area during the October 1-14, 1977
duck season.

Species

October 1-2
Percent
No.

Time Period
October 3-9
Percent
No.

No.

Entire 1/
Percent

Gadwall

22

14.0

33

38.8

55

22.7

American wigeon

30

19.0

8

9.4

38

15.7

N. shoveler

23

14.6

5

5.9

28

11.6

Redhead

13

8.3

14

16.5

27

11.2

Lesser scaup

16

10.2

8

9.4

24

9.9

Blue-winged or
Cinnamon teal

13

8.3

8

9.4

21

8.7

Mallard

11

7.0

5

5.9

16

6.6

Pintail

13

8.3

1

1.2

14

5.8

8

5.1

3

3.5

11

4.5

Ruddy

7

4.5

0

0.0

7

2.9

Common merganser

1

0.6

0

0.0

1

0.4

Green-winged

Total

1/

teal

157

85

242

No harvest occurred after October 9, 1977 .

The federal parts collection mail survey produces very small and thus,
unreliable samples of wings for North Park and South Park. Throughout the
three years (1975, 1976 and 1977) of the wing barrel survey, the results of
the two independent surveys do not agree.
Band Recovery Analysis
Analysis of band recoveries is now in a preliminary stage. Additional
information is being generated from the computer tapes and recovery data
for the 1976-77 and 1977-78 hunting seasons are being extracted by hand

�-58-

Table 8. Species composition of the harvest by time period at Antero Reservoir, South Park, Colorado during the October 1-14, 1977 duck hunting season.

Time Period
October 3-14
Percent
No.

Entire
Percent
No.

Species

October 1-2
Percent
No.

Mallard

31

60.8

7

41.2

38

55.9

5

9.8

4

23.5

9

13.2

Gadwall

4

7.8

2

11.8

6

8.8

Pintail

6

11.8

0

0.0

6

8.8

wigeon

3

5.9

0

0.0

3

4.4

Blue-winged or
Cinnamon teal

0

0.0

2

11.8

2

2.9

Redhead

2

3.9

0

0.0

2

2.9

Ruddy

0

0.0

1

5.9

1

1.5

Cornmon merganser

0

0.0

1

5.9

1

1.5

Total

51

Green-winged

American

teal

17

68

from the computer printouts received periodically from the Bird Banding
Laboratory.
Therefore, analysis of the information extracted from the
banding and recovery tapes would be premature
at this time. In addition,
tables compiled to date will not be presented in this report since they
may be changed and most likely will be duplicated in forthcoming reports.

�-59-

LITERATURE

CITED

Cowardin, L. M. 1977. Analysis and machine mapping of the distribution
of band recoveries.
U. S. Fish and Wildl. Servo Spec. Sci. Rep.
Wildl. 198. 8 p.
Davenport, D. A. 1977. Computerized tabulation and display of band
recovery data. U. S. Fish and Wildl. Servo Spec. Sci. Rep. Wildl.
199. 7 p.
Hoffman, R. W., and C. E. Braun.
1975. A volunteer
Colo. Div. Wildl. Game Inform. Leafl. No. 101.

wing collection
3 p.

station.

Hopper, R. M., A. D. Geis, J. R. Grieb, and L. Nelson, Jr. 1975. Experimental duck hunting seasons, San Luis Valley, Colorado, 1963-1970.
Wildl. Monogr. No. 46. 68 p.
Johnson, D. H. 1974. Estimating rates from banding of adult and juvenile
birds.
J. Wildl. Manage. 38(2):290-297.
Szymczak, M. R., and J. F. Corey.
Plains duck trap in Colorado.
l3 p ,

Prepared by

1976. Construction and use of the Salt
Colo. Div. Wildl. Div. Rept. No.6.

4

'itL:/J
7?
Michael R. Szymczak ~~
Wildlife

Researcher

4-

��-61-

October 1978

JOB PROGPillSSREPORT

State of

COLORADO
--------~~~~=------------

Project

No.

Work Plan No.
Job Title

Migratory

W-88-R-23
3

Job No.

------------------------

Morphological

Period Covered:

Differences

Bird Investigations
10

---------------------------------

Between Blue-winged

and Cinnamon

Teal

April 1, 1977 through March 31, 1978

Personnel:
B. Aufdengarten, R. Darnell, R. Ellis, P. Feiger, W. McDermitt,
M. Nail, L. Vaughn, U. S. Fish and Wildlife Service; Dr. P. Baldwin, Dr. J.
Law, Dr. R. Ryder, L. Sweanor, R. Thompson, K. Woodruff, Colorado State
University; M. Connor, J. Corey, D. Flenthrope, R. Hopper, R. Oakleaf, S.
Steinert, and R. Stark, Colorado Division of Wildlife.
ABSTRACT
Investigations of the morphological differences between blue-winged teal
(Anas discors) and cinnamon teal (Anas cyanoptera) were initiated in May
1977. Twenty females and 5 males of each species were collected in May and
June 1977 for use as a reference collection and for testing possible
speciating techniques.
One hundred and fifty blue-winged teal eggs were
collected in June 1977 from the Valentine National Wildlife Refuge in Nebraska
and 150 cinnamon teal eggs were collected in June from the Alamosa and Monte
Vista National Wildlife Refuges in the San Luis Valley of Colorado.
Eggs
were incubated, hatched, and the ducklings raised at the Fort Collins Wildlife Research Station.
Two hundred and twenty-nine ducklings hatched and 130
constituted the study sample, including 63 blue-winged teal (31 males, 32
females) and 67 cinnamon teal (30 males 37 females).
Twelve measurements
were taken weekly on the sample birds from the 5th through the 16th weeks of
age and monthly from the 20th week of age through March 1978. Eye color was
recorded weekly using the Munsell Book of Color and photographs of the head
and throat regions were taken periodically of randomly selected birds.
Preliminary analysis of data from 23 measurements taken on the adult birds
collected in 1977 indicate that the twelve measurements taken on the young
birds, especially culmen center and culmen ridge, appear to be promising
for distinguishing the two species. Measurements will be continued through
September 1978 at which time the data collected will be analyzed by computer.

��-63-

MORPHOLOGICAL
BETWEEN BLUE-WINGED

DIFFERENCES
AND CINNAMON TEAL

Robert S. Stark

Anas discors discors is one of two subspecies of blue-winged teal occurring
in North America.
It breeds from southern Ontario and Quebec south into
the northeastern United States, west through the north central United States
and Canada to California and north through central British Columbia into
Alaska.
The northern cinnamon teal, (Anas cyanoptera septentrionalium)
hereafter referred to as cinnamon teal, is one of five subspecies occurring
in the western hemisphere, but the only subpseciesto inhabit North America.
It breeds from the front range of the Rocky Mountains to the west coast and
from southern British Columbia and Alberta through Arizona and New Mexico
into north central Mexico (A.O.U. 1957, Delacour 1956, Johnsgard 1975).
Both species have major fall migrations through the central and southern
United States into Mexico and Central and South America (A.O.U. 1957,
Kortright 1953, Johnsgard 1975).
The breeding ranges of the two species overlap along the Rockies from
northern New Mexico to Alberta and British Columbia and west to eastern
Washington, Oregon, and northern California.
A problem of species identification exists in this area of overlapping ranges.
Females of the two
species appear identtical and pose the major problem, although immature males
may appear similar prior to 8 weeks of age, at which time the cinnamon teal
male acquires the red eye color of the adult (Spencer 1953). Most of the
descriptions of differences between the two species in the literature are
only relative and of limited value in accurate species identification.
In the major duck production areas of Colorado, blue-winged and cinnamon
teal are important breeding species representing an average of 11 percent
of the breeding population during the period 1954-75 (Szymczak 1977).
Combined, they are second only to the mallard (Anas platyrhynchos) in
annual production in Colorado and have contributed up to nearly 20,000 birds
to the state's breeding population in recent years.
During the pre-season banding period in Colorado, for the years 1971-74,
over 75 percent of the blue-winged and cinnamon teal banded were classified
as unidentified teal because they could not be accurately identified as
one species or the other (R. Hopper, pers. corom.). Knowledge of possible
population differences is essential to the formulation of management recommendations, but is not currently available for blue-winged and cinnamon teal
within the state. Special early teal seasons currently being conducted in
various parts of the U. S. cannot be justified in much of Colorado's high
country until the status of both blue-winged and cinnamon teal populations
within the state are determined.
A management scheme could then be formulated which would give greatest consideration to the species with the
poorest population status. This can be accomplished only after a reliable
technique is developed for distinguishing between the two species in the
field, involving one which does not result in the death of the birds.

�P. N. OBJECTIVES
1.

To compare
sex.

the morphology

of blue-winged

and cinnamon

teal by age and

2.

To develop a field technique for distinguishing between blue-winged and
cinnamon teal based on morphological diffferences, primarily for use
during the pre-season banding period (August an4 September).
SEGMENT OBJECTIVES

1.

Review literature on methods of rearing ducks in captivity
morphology of blue-winged and cinnamon teal.

2.

Collect 25 blue-winged
(May and June).

3.

Collect up to 150 eggs of each of the two species and artificially
incubate, hatch, and rear the young.

4.

Examine and describe plumage characteristics and eye color of each pen
reared bird at weekly intervals from mid-July at least through October.
Measure various body parts, including the bill, tarsus, toe and wing
weekly during this same period.
Maintain photograph records of randomly
selected birds of each species and sex. Obtain these same descriptions,
measurements and photographs for the 50 adult birds immediately following
their collection.

5.

Retain the pen reared birds over winter
following spring, summer and fall.

6.

Compile and analyze data and prepare
in appropriate technical journals.

teal and 25 cinnamon

teal during

to continue

and on

the spring

the study the

reports of findings

for publication

REVIEW OF LITERATURE
The classification of blue-winged and cinnamon teal has been described by
Kennard (1919), Delacour and Mayr (1945), Snyder and Lumsden (1951), Delacour
(1956), and the American Ornithologist's Union (1957).
The life history and distribution of blue-winged and cinnamon teal have been
discussed by Bent (1923), Phillips (1923), Bennett (1938), Kortright (1953),
Spencer (1953), the American Ornithologist's Union (1957), Wheeler (1965),
Johnsgard (1975), and Connelly (1977).
The comparative morphology of blue-winged and cinnamon teal has been studied
or described by Sclater (1912), Kortright (1953), Spencer (1953), Johnsgard
(1975), Palmer (1976), Dwyer (1976), Connelly (1977), and Wallace and
Ogilvie (1977).
General morphological characteristics of blue-winged teal have been described
by Sclater (1912), Kennard (1919), Allen (1920), Bent (1923), Bennett (1938),
Kortright (1953), Delacour (1956), Glover (1956), Dane (1966 and 1968), Greij
(1973), Johnsgard (1975), and Palmer (1976).

�-65-

General morphological characteristics of cinnamon teal have been described
by Sc1ater (1912), Stephens (1921), Bent (1923), Kortright (1953), Spencer
(1953), De1acour (1956), Johnsgard (1975), and Palmer (1976).
Bird weight and its importance in taxonomy has been discussed by Weatherbee
(1934), Stewart (1937), Baldwin, Prentiss and Kendeigh (1938), Nice (1938),
Amadon
(1943), Be11rose and Hawkins (1947), Nelson and Martin (1953), and
Smart (1965).
Sex and age determination of waterfowl have been described by Hochbaum
(1942), Hanson (1949), Southwick (1953), Go110p and Marshall (1954), Dane
(1968), and Pettingill (1970). Determining the age and stage of incubation
of bird eggs has been described by Westerskov (1950) and Weller (1956).
Plumages and molts have been described by Lillie (1932), Lesher and Kendeigh
(1941), Weller (1957), Dzubin (1959), Humphrey and Parkes (1959 and 1963),
Stresemann (1963), Smart (1965), Amadon (1966), and Oring (1968).
Hubridization involving either blue-winged or cinnamon teal has been studied
by Deane (1905), Swarth (1915), Rothschild and Kinnear (1929), Sibley (1957),
Wilson and Ven Den Akker (1948), Childs (1952), Cockrum (1952), Anderson
and Miller (1953), Johnsgard (1960), Harris and Wheeler (1965), Hall (1968),
and Lahrman (1971).
Methods for artificially incubating and hatching birds eggs have been described
by Ro1nik (1943), Hanson (1951), Dill and Lee (1970), Ellis (1974), F1ieg
(1974), Greenwell (1974), Lint (1974), and Sheraw (1975).
Techniques for captive1y brooding and raising waterfowl have been described
by Greenberg (1949), Dill and Lee (1970), Martin (1972), Block (1974),
Smart (1974), Strutz (1974), Wilson (1974), and Sheraw (1975).
Methods of taking bird measurements and the relative importance of different
masurements have been described by Reichenberger (1923), Bergto1d (1925),
Myers (1928), Baldwin, Obserho1ser and Worley (1931), Pettingill (1970),
and Ferguson (1974).
The success of releasing captive reared birds into the wild has been
studied by Lincoln (1934), Errington and Albert (1936), Williams and
Kalmbach (1943), and Brakhage (1953).
METHODS AND MATERIALS
Collection of Adult. Teal
Thirteen cinnamon teal (5 males, 8 females) and one female blue-winged teal
were collected in the San Luis Valley, Colorado on 16 and 17 May, 1977.
On 26 May 1977, 12 blue-winged teal (5 males, 7 females) were collected at
the Red Lion Property near Crook, Colorado. Only females in the presence of
.a male were collected.
An additional 24 female teal were collected concurrently with the
egg collection phase of the study described below; 12 blue-winged
teal from the Valentine National Wildlife Refuge in north central
Nebraska and 12 cinnamon teal from the Alamosa and Monte Vista

�-66-

National Wildlife Refuges in the San Luis Valley of Colorado.
Species
identification of these 24 females was deferred until young males from
their respective clutches acquired nuptial plumage in the spring and summer'
of 1978 while in captivity at the Fort Collins Wildlife Research Station.
All collections were made with a shotgun and all 50 birds (5 males and 20
females of each species) were weighed and tagged with an identification
number within 24 hours after collection.
They were also measured and then
frozen for use as a reference collection of adult wild teal. The form used
to record the measurements taken is presented in Appendix A.
Egg Collections
On 7 June 1977, 150 suspected blue-winged teal eggs (and 12 adult female
blue-winged teal associated with the nests) were collected from the Valentine
NWR and transported to the Fort Collins Wildlife Research Station.
Nests
were located by flushing incubating females with a 160-foot cable dragged
through nesting habitat between two jeeps, as described by Ladd (1969).
All nests were located and marked prior to collecting eggs and all eggs
were indivuda1ly identified by nest and female at the time of collection.
The eggs were transported in a portable incubator designed and built by
J. Corey, Colorado Division of Wildlife.
The incubator operated from a car
battery and maintained a temperature of 990F and a hygrometer reading of 84oF.
During the period 6-21 June 1977, 150 suspected cinnamon teal eggs (and 12
adult female cinnamon teal associated with the nests) were collected from
the Alamosa NWR and the Monte Vista NWR. Nests were located by crews of
4-6 people walking systematically through likely nesting habitat and
flushing incubating females.
Collection and transport procedures were
similar to those used during the Nebraska collection, except some clutches
were placed in a chest cooler with cotton wadding and hot water bottles and
flown to Fort Collins.
The ether wafer in the incubator at the Fort Collins Wildlife Research
Station ruptured on 27 June 1977, resulting in over-heating and death of
75 embryos in eggs collected in Colorado.
An additional 75 cinnamon teal
eggs were collected on 30 June and I July from the Alamosa NWR to insure
that an adequate sample of cinnamon teal was obtained.
Hatching

and Rearing of Young

The eggs were not cleaned prior to setting in the incubator, but excessively
dirty eggs were scraped with a knife.
All eggs were weighed and the length
and width measured prior to placement in the incubator.
A Humidaire
Model 50 automatic incubator and a David Bradley incubator were used for
incubating and hatching the eggs at the Fort Collins Wildlife Research
Station.
The incubator was set to maintain a temperature of 99.50F and a
hygrometer reading of 86oF. The automatic turning device on the incubator
was set to turn the eggs every 2 hours.
The hatcher was set at a temperature
of 990F and a hygrometer reading of 94oF. The eggs were not turned in the
hatcher.
During the incubation period the water pump and hygrometer bottles
were filled daily with distilled water.

�-67-

Trays in the hatcher were separated into five compartments and each was
enclosed by hardware cloth. Each clutch of eggs was placed in an individual compartment in the hatcher when the first egg of the clutch pipped.
Ducklings from each clutch were thereby separated from other clutches after
hatching for accurate identification.
The ducklings were allowed ~o dry
several hours after hatching before being removed from the hatcher.
Upon
removal, they were weighed and banded with individually numbers plastic
bands and recorded by clutch.
After banding the ducklings were placed in a battery brooder with the temperature maintained at 90oF. Newspaper was placed on the floor of each of
the five brooder compartments and an unmedicated duck starter feed prepared
by the Golden West Milling Company, Longment, Colorado was sprinkled on the
paper to induce feeding.
Two waterers were placed in each brooder compartment and changed 4-5 times daily.
Feed was placed in small feeders and
changed twice daily after the ducklings were eating.
The ducklings were fed an unmedicated duck maintenance feed prepared by the
Golden West Milling Company after 12 days of age. At this time they were
also removed from the battery brooder, rebanded with individually numbered
aluminum bands and placed in a brooder house.
The brooder house was 8' x 12'
and the floor was covered with wood shavings litter.
Four large waterers
and three feeders were placed on the litter and changed three times daily.
Four 250 watt infared heat lamps were placed above the litter and left on
24 hours a day.
At 3 weeks of age the ducklings were allowed access to a 5' x 8' section of
a concrete pond adjacent to the brooder house.
The ducklings were then
allowed to swim daily to induce functioning of the oil gland and preening.
At 5 weeks of age the ducklings were rebanded permanently
with size 5 U. S.
Fish and Wildlife Service bands and moved to an outside pen. The pen
measured 30' x 75' and was completely enclosed in chicken wire, including
the top. Several small shelters were placed in the pen to provide shade,
and a concrete pond 8' x 36' was built to provide water.
Feed was changed
twice daily in three feeders.
The pond water was changed every third day.
The teal were moved in November 1977 to an 18' x 32' pen for wintering.
This facility was divided into three compartments each 6' x 32'. The pen
was raised 2 feet above the ground and had a welded wire floor to allow
waste material to pass through.
Each compartment was divided into two
6' x 16' sections, with one entirely enclosed with weatherproofing material
and provided with eight 250 watt infared heat lamps to maintain an above
freezing temperature.
The other section was weatherproofed on all except
one side but not provided with heat. A small l' x l' door, allowed passage
between the two sections.
Feeders and waterers were placed in the heated
section and changed twice daily.
Birds surviving into November were
separated by week of hatching into the three compartments of the winter
pen. The teal remained in the winter pen through the end of the Segment,
but will be moved back to the outside pen in April 1978.

�-68-

Measurements

and Morphological

Descriptions

The teal were measured weekly from the 5th through the 16th weeks of age
and monthly from the 20th week of age through March 1978. Five standard
ornithological measurements were taken on the teal including weight,
length of tarsus, length of exposed culmen, length of bill from gape, and
width of bill at widest anterior part (Baldwin et a1. 1931).
The weight
was measured in grams and all other measurements in millimeters.
Seven
additional measurements were taken based on the data from the adult wild
teal (Table 1). See Appendix B for the form used in recording measurements and other information for each bird.
Eye color was identified by
hue, value, and chroma with the Munsell Book of Color from the Kollmorgan
Corporation, Baltimore, Maryland, and recorded weekly.
Photographs of the
eye, cheek,
side of head, throat, and breast were taken periodically of
randomly selected birds throughout the study period.

RESULTS AND DISCUSSION
Adult Teal
Preliminary results of measurements of adult teal collected in May 1977
include mean value and range for each sex and species (Table 1). The
promising measurements for distinguishing between the species based on these
data include: culmen center, culmen ridge, culmen side, length of lower bill
center, and length of lower bill ridge.
Only females collected in the
presence of a male were included in this preliminary analysis.
Hatching

Success and Measurement

of Young

Two hundred and twenty-nine eggs hatched for a total hatching success of
76.3%, including 106 of the Nebraska eggs (70.6%) and 123 Colorado eggs
(82.0%).
One hundred and seventy of the 229 birds that hatched survived
through March 1978 for a 74.2% survival rate.
Twenty-five ducklings either lost their plastic bands or hatched in the
incubator.
These ducklings were classified as unidentified and were maintained but not included in the sample of birds measured and described
during the study. An additional 15 cinnamon teal were also eliminated from
the study in order to maintain similar sample sizes for each species.
Thus,
the sample size was 130 birds, of which there were 63 blue-winged teal (31
males, 32 females) and 67 cinnamon teal (30 males, 37 females).
The total number of measurements taken through March 1978 was over 26,000.
Measurements of the teal will continue through September 1978, at which
time due to the volume, all data collected in the study will be analyzed
by computer.
Photographs will also be taken through September 1978 and will
be used to demonstrate possible morphological differences between bluewinged and cinnamon teal which are not subject to quantitative analysis.

�-69-

Table 1. Range and mean values for some measurements of adult male and
female blue-winged and cinnamon teal collected in May 1977. 1/
Cinnamon Teal
Mean 2/
Range

Blue-winged Teal
Mean
Range

Weight J:.../

394

355-415

362

355-395

Wing

201

185-220

183

180-184

Tarsus

41.1

39.7-42.2

38.6-40.6

Foot

43.9

42.6-45.5

39.3
41.8

Middle toe

37.2

36.0-37.5

35.5

33.0-37.0

Culmen center

44.2

42.1-46.5

39.2

37.0-42.4

Culmen ridge

51.4

49.8-53.5

46.1

44.1-48.9

Culmen side

48.7

46.8-51.0

43.3

41.0-46.3

Maximum width upper bill

18.2

17.9-18.6

17.9

16.6-18.6

Width at nares

18.0

17.9-18.4

16.6

16.4-17.0

Posterior width upper bill

16.3

15.6-16.8

14.8

14.4-15.0

Lower bill center

40.8

38.6-42.6

37.7

35.7-39.9

Lower bill ridge

49.9

48.2-52.3

44.8

41.0-48.9

Lower bill posterior width

13.7

13.1-14.1

12.1

10.7-13.0

Lower bill anterior width

12.2

11.8-12.4

12.0

11.7-12.1

Lower bill nail length

6.4

5.5-7.1

5.6

5.2-6.3

Lower bill nail width

5.4

4.5-5.8

5.2

4.9-5.5

Nares length

4.2

3.8-4.5

3.7

3.4-3.9

Upper bill nail length

8.5

7.8-9.0

7.7

7.3-8.1

Upper hill nail width

5.4

5.1-5.7

5.4

5.1-5.8

Height of bill base

17.0

14.8-18.9

16.2

15.5-16.9

Height of bill nares

13.0

12.0-13.9

12.2

11.8-12.7

Height of bill anterior

10.4

9.8-11.4

9.1

8.7-9.4

. h t:2/
Welg

388

335-425

382

315-455

Wing

182

173-186

171

153-181

Tarsus

38.8

36.9-40.9

38.2

37.1-40.1

Sex and Measurement
Males (5 of Each Species)

39.4-42.8

Females (8 of Each Species)

----------------------------------------------------------------------------

�-70-

Table 1. Range and mean values
female blue-winged and cinnamon

Cinnamon Teal
Mean 2/
Range

Sex and Measurement

Females

for some measurements of adult male and
teal collected in May 1977 (cont.). 1./

(8 of Each Species)

Blue-winged
Mean

Teal
Range

(continued)

Tarsus

38.8

36.9-40.9

38.2

37.1-40.1

Foot

41.6

39.0-44.7

40.1

37.3-41. 9

Middle toe

35.4

34.0-37.5

33.8

31.5-35. 0

Culmen center

43.3

41.15-45.6

38.5

36.5-39.45

Culmen ridge

49.0

46.0-51. 9

44.6

43.4-46.2

Culmen side

45.7

43.4-48.5

41. 9

41.1-43.4

Maximum width upper bill

17.4

16.5-18.6

17.2

16.7-17.8

Width at nares

16.7

16.1-17.5

16.0

15.2-16.8

Posterior

15.2

14.2-16.2

14.4

13.6-15.2

Lower bill center

39.4

34.1-43.2

35.4

31.8-36.9

Lower bill ridge

46.7

43.6-49.0

42.0-45.1

width

12.9

11. 5-13. 6

43.5
11. 9

Lower bill anterior width

11.8

11.3-12.1

11.6

11.1-12.4

Lower bill nail length

6.3

5.3-6.8

6.3

5.3-7.1

Lower bill nail width

5.8

4.9-6.7

5.2

4.6-5.6

Nares length

3.5

3.0-3.9

3.3

2.9-3.8

Upper bill nail length

8.5

7.8-9.5

7.8

6.8-8.9

Upper bill nail width

5.8

5.0-6.3

5.5

5.0-6.2

Height of bill base

16.4

15.3-18.1

16.0

15.1-16.9

Height of bill nares

12.7

11.9-13.2

11.8

11.3-12.5

Height of bill anterior

9.2

8.8-9.6

8.8

8.6-9.2

1/ All females

of a male when collected.

width upper bill

Lower bill posterior

1/ Weight

in the presence

in grams and all other measurements

in millimeters.

11.0-12.9

�-71-

LITERATURE

Allen, G. M.
Amadon, D.

1920.
1966.

CITED

Pattern development
Avian plumages

in teal.

and molts.

Auk. 37:558-564.

Condor 68:263-278.

American Ormithologist's Union.
1957. Check-list of North American
birds.
5th Ed. Lord Baltimore Press, Inc., Baltimore. 691pp.
Anderson, W., and A. W. Miller.
1953. Hybridization of
blue-winged teal in northern California.
Condor 55:152-153.
Baldwin, S. P., H. C. Oberholser, and L. G. Worley.
1931. Measurements
of birds.
Sci. Publ. of the Cleveland Mus. Nat. Hist. Cleveland. 165pp.
_____ , and S. C. Kendeigh.
Auk 55:416-467.

1938.

Variations

Bellrose, F. C., and A. S. Hawkins.
Auk 64:422-430.
Bennett, L. J.
Collegiate
Bent, A. C.
anseres

1947.

in the weight of birds.

Duck weights

in Illinois.

1938. The blue-winged teal: its ecology and management.
Press, Inc., Ames, Iowa.
144pp.

1923. Life histories of North American waterfowl:
(Part 1). U.S. Natl. Museum Bull. 126. 250pp.

Bergtold, W. H.
27:59-61.

1925.

The relative value of bird measurements.

order

Condor

Block, M. 1974. Artificial lighting.
In D. O. Hyde (ed.): Raising
wild ducks in captivity.
E. P. Dutton and Co., Inc., New York.

319pp.

Brakhage, G. K. 1953. Migration and mortality of ducks hand reared and
wild trapped at Delta, Manitoba.
J. Wildl. Manage. 17:465-477.
Childs, H. E., Jr. 1952.
Condor 54:67-68.

Hybrid between a shoveller

and a blue-winged

teal.

Cockrum, E. L. 1952. A check-list and bibliography of hybrid birds in
North American north of Mexico.
Wilson Bull. 64:140-159.
Connelly, J. W., Jr. 1977. A comparative study of blue-winged and
cinnamon teal breeding in eastern Washington.
M.S. Thesis.
Washington
State University, Pullman.40pp.
Dane, C. ·W. 1966. Some aspects of breeding
teal. Auk 83:389-402.
1968. Age determination
32:267-274.

biology

of blue-winged

teal.

of the blue-winged

J. WiIdl. Manage.

�-72-

Delacour, J.
London.

1956. Waterfowl
232pp.

and E. Mayr.

1945.

_____ , 1946. Supplementary
58: 104-110.

of the world.

Vol. 2.

The family Anatidae.

Country Life Limited,

Wilson Bull. 57:3-55.

notes on the family Anatidae.

Dill, H. H., and F. B. Lee (eds.).
and Wildl. Servo 154pp.

1970.

Wilson Bull.

Home grown honkers.

u.S. Fish

Dwyer, G. L. 1976. Competition and hostile behaviors of blue-winged
cinnamon teal in western Montana.
M.A. Thesis.
University of
Montana, Missoula.
77pp.

and

Dzubin, A.
1959. Growth and plumage development of wild-trapped juvenile
canvasback (Aythya valisneria).
J. Wildl. Manage. 23:279-290.
Ellis, J. 1974. Using an artificial
Raising wild ducks in captivity.
York.
319pp.
Errington, P. C., and W. E. Albert.
mallard ducks.
Bird Banding.

incubator.
In D. o. Hyde (ed.):
E. P. Dutton and Co., Inc., New

1936. Banding
7:69-73.

studies of semi-domesticated

Ferguson, W. 1974. The importance of weighing ducklings.
In D. o. Hyde
(ed.): Raising wild ducks in captivity.
E. P. Dutton and Co., Inc.,
New York.
319pp.
Flieg, G. M. 1974. Incubators and methods.
In D. O. Hyde (ed.): Raising
wild ducks in captivity.
E. P. Dutton and Co., Inc., New York.
319pp.
Glover, F. A. 1956. Nesting and production of the blue-winged teal (Anas
discors Limmaeus) in northwest Iowa. J. Wildl. Manage. 20:28-46.
Gollop, J. B., and W. H. Marshall.
1954. A guide for aging duck broods
in the field. Miss. Flyway Council Tech. Comm.
14pp.
Greenberg, D. D. 1949. Raising game birds in captivity.
Co., Inc., New York. 224pp.

D. Van Nostrand

Greenwell, G. A. 1974. On helping ducklings
(ed.): Raising wild ducks in captivity.
New York.
319pp.

out of the egg. In D. O. Hyde
E. P. Dutton and Co., Inc.,

Greij, E. D. 1973.
Effects of sex hormones
teal. Auk 90:533-551.

on plumages

of the blue-winged

Hall, F. A., and S. W. Harris.
1968. Hybrid of blue-winged
in north western California.
Condor 70:188.

teal shoveller

�-73-

Hanson, H. C. 1949. Methods of determining age in Canada geese and other
waterfowl.
J. Wildl. Manage. 13-177-183.
1951. Notes on the artificial
Manage. 15-68-72.
Harris, S. W. ,and R. J . Wheeler.
cinnamon teal in northwestern
HochbAum,
H. A.
examination.

propagation

of wood ducks.

1965. Hybrid of blue-winged teal x
California.
Condor. 67:539-540.

1942. Sex and age determination of waterfowl
Trans. N. Am. Wildl. Conf. 7:299-307.

Johnsgard, P. A. 1975. Waterfowl of North America.
Bloomington and London.
575pp.
Kennard, F. H. 1919.
Auk 36:455-460.

J. Wildl.

Notes on a new subspecies

by cloacal

Indiana Univ. Press,

of blue-winged

teal.

Kortright, F. H. 1953. The ducks, geese and swans of North America.
Wildl. Manage. Inst., Washington D. C. 476pp.
Ladd, W. N., Jr. 1969. Relationship of predation and land use practices
to duck nesting activities on Valentine National Wildlife Refuge,
Nebraska.
M.S. Thesis.
Colorado State University, Fort Collins.
116pp.
Lahrman, F. W. 1971. Hybrid cihnamon teal and blue-winged
Regina.
The Blue Jay 29:28.

teal at

Lesher, S. W., and S. C. Kendeigh.
1941. Effect of photoperiod
of feathers. Wilson Bull. 53:169-180.
Lillie, F. R. 1932.
44: 193-211.

The physiology

of feather pattern.

on molting

Wilson Bull.

Lincoln, F. C. 1934. Restocking of marshes with hand-reared mallards not
proved practicable.
Yearbook of Agric., U.S. Dept. Agric. 1934:310-313.
Lint, K. C. 1974. The artificial incubator.
In D. o. Hyde (ed.): Raising
wild ducks in captivity.
E. P. Dutton and Co., Inc., New York. 319pp.
Martin, R. M.
157pp.

1972.

Wild fowl in captivity.

Myers, E. C.

1928.

Taking body weights of birds.

Nelson, A. L., and A. C. Martin.
17:36-42.

1953.

Nice, M. M. 1938. The biological
Banding 9: 1-11.

significance

John Gifford Ltd., London.

Auk 45:334-338.

Game bird weights.

J. Wildl. Mange.

of bird weights.

Bird

�-74-

Oring, L. W.
380.

1968.

Growth, molts, and plumages of the gadwall.

Auk 85:355-

Palmer, R. S. 1976. Handbook of North American birds.
Univ. Press, New Haven, Conn. and London.
521pp.

Vol. 2. Yale

Pettingill, o. S., Jr. 1970. Ornithology
Publ. Co., Minneapolis, Minn. 524pp.

and field.

in laboratory

Burgess

Phillips, J. C. 1923. A natural history of the ducks Vol. 2. Houghton
Mifflin Co., Boston, Mass.
490pp.
Reichenberger, E.
40:244-247.

1923.

Remarks on methods in measuring

Rolnik, V. 1943. Instructions
Wildl. Manage. 7:155-162.

for the incubation

Rothschild, W., and N. B. Kinnear.
Brit. Ormith. Club. 49:93-99.

1929.

birds.

Auk

of eider duck eggs.

List of hybrid ducks.

Sclater, W. L. 1912. A history of the birds of Colorado.
Co., London.
567pp.
Sheraw, D. 1975. Successful duck and goose raising.
Co., Pine River, Minn. 203pp.

J.

Bull.

Witherby

and

Stromberg Publ.

Sibley, C. G. 1957. The evolutionary and taxonomic significance of
sexual dimorphism and hybridization in birds.
Condor. 59:166-191.
Smart, G. 1965. Development and maturation of primary feathers of red
head ducklings.
J. Wildl. Manage.
29:533-536.
Synder, L. L., and H. G. Lumsden.
1951. Variations in Anas cyanoptera.
Royal Ontario Mus. of Zool. Occas. Paper No. 10. pp.I-18.
Southwick, C. 1953. A system of age classification for field studies
of waterfowl broods.
J. Wildl. Manage. 17:1-18.
Spencer, H. E., Jr. 1953. The cinnamon teal (Anas cyanoptera Vieillot):
its life history, ecology and management.
M.S. Thesis. Utah State
Agricultural College, Logan.
184pp.
Stephens,

F.

1921.

Eclipse plumage of cinnamon teal.

Stewart, P. A.

1937.

A preliminary

Stresemann,

1963.

The nomenclature

E.

Condor. 23:194.

list of bird weights.

Auk 54:324-332.

of plumage and molts.

Auk 80:1-8.

�-75-

Strutz, C. 1974. Winter care of wild ducks in captivity.
In D. O.
Hyde (ed.): Raising wild ducks in captivity.
E. P. Dutton Co., Inc.,
New York.
319pp.
Swarth, H. S. 1915. An apparent hybrid between species of the genera
Spatula and Querquedula.
Condor.
17:115-118.
Szymczak, M. R. 1977. Waterfowl production surveys.
Wildl., Fed. Aid Game Res. Rept., Oct. pp.1-14.

Colo. Div. of

Wallace, D. I. M., and M. A. Ogilvie.
1977. Distinguishing
and cinnamon teals. British Birds 70:290-294.
Weatherbee, K. B. 1934. Some measurements
Bird Banding 5:55-63.

blue-winged

and weights of live birds.

Westerskov, K. 1950. Methods for determining
J. Wildl. Manage. 14:56-67.

the age of game bird eggs.

Wheeler, R. J. 1965. Pioneering of blue-winged teal in California,
Oregon, Washington and British Columbia.
Murrelet 46:40-42.
Williams, C. S., and E. R. Kalmbach.
transported juvenile waterfowl.

1943.

Migration

and fate of

J. Wildl. Manage. 7:163-169.

Wilson, C. 1974. Artificial and natural ponds.
In D. O. Hyde (ed.):
Raising wild ducks in captivity.
E. P. Dutton and Co., Inc., New
York.
319pp.
Wilson, V. T., and J. B. Van Den Akker.
1948. A hybrid cinnamon teal blue-winged teal at Bear River Migratory Bird Refuge, Utah. Auk 65:316.

_S~_s;~L=&gt; r..L..:l==--_

Prepared by ----I.f7u~(/~rz-4____L...:._·/
Robert S. Stark

�-76-

APPENDICES

�-77APPENDIX

BLUE-WINGED

TEAL CINNAMON

TEAL REFERENCE

Specimen Number

_

Species

_

Location

_

Wing Length

Date Collected

SHEET

---------------------Age

Sex

'-----------

_

Culmen:

Eye Color

------------

Maximum Width

Center

Ridge

Posterior

-------------------

Ridge

Width

---------

Nail Length
Length of Nares

_
_

At Base

End of Nail

------------------Toe

---------------

Side

---------------

-------------------

Width

Anterior

At Nares

----------------~

Width

Nail Width

Nail: Length

-----------------

At Posterior

Comments:

Minimum Width

-------------------

Posterior

Height of Bill:

_

--------------------

Width at Nares
Bill:

Length of Middle

----------------

Center

_

Length of Tarsus

-------------------

Length of Foot

Lower

DATA COLLECTION

-----------------------------------------------------------------------

Weight

Bill:

A

_

----------------

-------------------_

Width

-------

-----------------

---------------------------

�-78APPENDIX

Specimen

Number

_

Species

_

Sex.

_

Hatching

Date.

Date of Measurement
Age
Eye Color
Weight
Length of Tarsus
Culmen:

Center
Ridge
Side

Maximum

Width of Bill

Width of Bill at Nares
Lower Bill: Center
Ridge
Height

of Bill:
Anterior

Length of Head
Comments:

_

B

�-79October 1978

JOB PROGRESS REPORT

State of

COLORADO
-----------------------------

Project No.

W-88-R-23

Work Plan No.

6

Job Title

Migratory
Job No.

Migratory

Bird Investigations
1

---------------------------------

Bird Publications

----------------~----------------------------------~----------------

Period Covered:
Personnel:

April 1, 1977 through March 31, 1978

William J. Adrian, David R. Anderson, Clait E. Braun, Howard D.
Funk, Richard M. Hopper and Michael R. Szymczak.

ABSTRACT
Publications
as follows:

planned for and accomplished

under this job for Segment 23 are

Braun, C. E. 1978. Molt and age determination of band-tailed pigeons.
Accepted by J. Wildl. Manage.
To be published in July, 1978 issue.
Braun, C. E. 1978. Trichomoniasis in band-tailed pigeons.
Calif. Fish and Game. To be published in 1978.

Accepted by

Hopper, R. M., H. D. Funk, and D. R. Anderson.
1978. Age specificity
mallards banded postseason in eastern Colorado.
J. Wildl. Manage.
42(2):263-270.
Szymczak, M. R., and W. J. Adrian.
1978. Lead poisoning
southeast Colorado.
J. Wildl. Manage. 42(2):299-306.

in

in Canada geese in

Szymczak, M. R. 1978. Steel shot use on a goose hunting area in Colorado.
Tentatively accepted for publication in The Wildlife Society Bulletin.

Prepared by __/~::A-=-" _t_·-.:~=--:-.t.-.:.12==-,'_-~~.(:~'-··:__:_--~~)L..~-=u'--'--".C-;.=~..:....=·-=--_
Howard D. Funk
Section Chief
Small Game Research

��October

-81-

JOB PROGRESS

State of

REPORT

COLORADO
--------~~~~~-----------

Proj ect No.

Job Title

Job No.

7

Bird Investigations
~l

_

Investigation of American Coots in Colorado
--------~------------------~----~~~~------~----~-----

Covered:

Personnel:

Migratory

W-88-R-23

Work Plan No.

Period

1978

April 1, 1977 to March 31, 1978

W. P. Gorenzel and R. A. Ryder, Colorado State University;
Braun and H. D. Funk, Colorado Division of Wildlife.

C. E.

ABSTRACT
Investigations concerning distribution and status, breeding densities, reproductive success, habitat utilization, timing of migration, and hunter harvest
of coots (Fulica americana) were conducted in Colorado between August 1976
and March 1978. Four study areas were selected to represent breeding sites
in the major topographic areas of the state, one in the eastern plains, two
in high mountain valleys, and one west of the Continental Divide.
Coots
were distributed throughout most of Colorado, breeding in most areas and
resident year round in low numbers, primarily along the western boundaries of
the plains.
The nesting season extended from mid-April to mid-August.
Peak
nest initiation dates ranged from 25 April to 6 June, while peak of hatch
varied from 30 May to 4 July.
Two hundred and eleven nests were located
with nesting and hatching success of 82 and 90 percent, respectively.
Average clutch size wasS.2.
Clutch size decreased as the season progressed.
Predation accounted for 76 percent of the unsuccessful nests.
Coots were
found to breed primarily in cattail (Typha spp.) and bulrush (Scirpus spp.)
dominated marshes.
Bulrush was the preferred nest cover type. Breeding
densities ranged from 14 to 35 successful nests/ha of cattail or bulrush.
Spring migration began in March, peak numbers were recorded in April.
Peak
concentrations of fall migrating coots were recorded in September.
Review
of harvest data and results from hunter questionnaires indicated coots were
of minor importance in Colorado.

�-82-

RECOMMENDATIONS

1.

Migration counts should continue
late summer and fall to document
of hunting seasons.

on all study areas, especially in
coot abundance in relation to timing

2.

Additional data gathering efforts concerning breeding densities
habitat relationships and productivity should be continued.

3.

Information gathering efforts concerning
survival should be continued.

brood counts and immature

�-83-

INVESTIGATION

OF AMERICAN
Warrner

COOTS IN COLORADO

P. Gorenzel

The American coot (Fulica americana Gmelin) is the largest and most aquatic
member of the order Gruiformes, family Rallidae, in North America.
The coot,
hereafter used in reference to the North American subspecies F. a. Americana,
breeds primarily on fresh water wetlands.
Highest nest densities occur
on glacial marshes of the north central United States and south central
Canada.
In winter coots utilize both brackish and fresh water habitats.
Important wintering areas are the Central Valley of California, the
coastal marshes of Texas and Louisiana, and wetlands of Florida
(Fredrickson et al. 1977).
State and federal regulations list the coot as a migratory game bird.
As such it is classed as "waterfowl" and is hunted during the regular
duck season. Although coots can constitute a large percentage of the
waterfowl harvest in certain areas of the United States, overall hunting
interest in coots is low nationwide (Fredrickson et al. 1977).
Limited data exist concerning the status of the coot in Colorado.
General
information on nest observations, early and late occurrences, and distribution has been summarized by Bailey and Niedrach (1965) and Sclater
(1912). Little is known about habitat preferences, timing of habitat
use,productivity,
and levels of hunting mortality within Colorado.
This
study was initiated to provide data for the management of coots in Colorado.
P. N. OBJECTIVES
The major objective of this study is to obtain data necessary for the
development of a management plan for coots breeding in and migrating
through Colorado.
Data considered necessary are those relating to distribution, breeding densities and production in the eastern plains, high
mouritain valleys,and west of the Continental Divide, descriptions of
habitats utilized according to vegetation, water characteristics,
documentation of timing of spring and fall migration, and present levels of
reported harvest.
Hypotheses that have been developed are:
1.

Distribtuion of coots in Colorado is determined by the occurrence
cattail (Typha spp.) and/or bulrush (Scirpus spp.) and dominated
marshes.

of

2.

Densities of breeding coots, nesting success and productivity differ
significantly between occupied habitats in the eastern plains, high
mountain valleys, and west of the Continental Divide.

3.

Timing of spring and fall migrations differs significantly between
areas east of the mountains, high mountain valleys, and west of the
Continental Divide.

�-84-

4.

Hunting in Colorado is the most important
locally produced coots.

source of mortality

for

SEGMENT OBJECTIVES
1.

Investigate selected habitats for coot occurrence and circulate
questionnaires to determine distribution and status in Colorado.

2.

Select and describe study areas representative of general wetlands
on the eastern plains, intermountain valleys, and west of the Continental Divide.

3.

Conduct weekly counts to document
migra tion s •

4.

Conduct nest searches

5.

Collect information concerning
attitudes in Colorado.

6.

Compile data, analyze results and prepare progress report.

the timing of spring and fall

to obtain breeding

data.

harvest levels of coots and hunter

METHODS AND MATERIALS
Three methods were used to investigate
in the selection of study areas:

coot occurrence

in Colorado

1.

Extensive

2.

Field trips in conjunction

3.

Questionnaires sent to field personnel of the Colorado Division
Wildlife and amateur ornithologists (Appendix).

and aid

field trips as part of planned research.
with graduate

courses.
of

Responses from the questionnaire, personal observations, and results from
the Colorado Bird Distribution Latilong Study (Kingery and Graul 1978)
were combined to plot coot distribution and status according to the latilong system.
In this system, the state was divided into distinct areas
based on latitude and longitude (Fig. 1). Coot status was then determined
within each area.
Study areas were selected based on the following

criteria:

1.

Sufficient area and cover to provide adequate numbers
breeding coots.

2.

Stable water levels.

3.

Similarity

of the area to general wetlands

(25 pairs) of

in the region.

�I·
I•·
I

I
CXl
\Jl

I

,,

.

.

I

----"'.,.. ..--Fig. 1.

COLORADO

trating

Scale of "-liles
I~

11.J.l

• __ ..

~~=-:..:-'~~u

�-86-

4.

Accessibility.

5.

Freedom

from excessive

external

interference.

Counts were conducted on a weekly basis on all study areas to determine
migration periods and the size of breeding coot populations.
A 20 X
Bausch and Lomb spotting scope and 8 x 40 Bushnell binoculars were used.
Counts were standarized by counting from the same location at each area,
at approximately the same time of day for a given area. Weather conditions
and other factors were noted on a field form (Appendix).
Each study area was mapped using a staff compass and tape. Islands of
emergent vegetation were plotted by pacing and with the aid of a hand
held compass and an Edscorp field range finder.
A Dietz planimeter was
used to measure total wetland area and area of emergent components.
Aquatic vegetation was sampled along transects using a method modified
from the Atlantic Waterfowl Council (1972). Transects were established
100 and 200 m apart based on the size of the area. Sampling intervals
were 35 to 100 m apart on each transect.
A rake was used to collect submerged vegetation.
Vegetation density ratings were based on amounts
recovered and were ranked as follows: dense, moderate, sparse.
Scientific
names follow Harrington (1964). At each station, water depth and bottom
consistency were noted.
Marked poles were placed on all study areas to
record water level fluctuations.
Water samples were taken during the 27
June to 1 July period using a Hach Kit model AL-36B.
Soil survey information of adjacent uplands, obtained from the Soil Conservation Service,
U. S. Department of Agriculture, and samples along aquatic transects were
used to identify marsh substrate types.
Nest searches were made on all study areas by systematically wading through
all emergent vegetation.
If the water was too deep to wade, the search
was conducted by boat. A nest was considered to b~ any platform-like
structure holding one or more eggs. In those instances where a nest was
discovered after hatching, the platform was identified as a nest if
numerous small shell fragments were present.
Each nest, when found, was
numbered and marked with a small cardboard tag. In addition, a piece of
yellow surveyor's tape was attached to vegetation approximately 0.6 to
12 meters away from each nest in such a manner as to be conspicuous to
the observer.
Each piece of tape was marked with a grease pencil to
indicate the nest number, direction, and distance to the nest.
Once found,
nests were revisited once a week until hatching.
A number of nest site measurements were recorded, including type of cover
vegetation; its condition, i.e., dead or alive; height of a representative
sample of the vegetation; nest concealment, i.e., was the nest concealed
on one, two, three, or four sides, and/or from above; depth of water; and
distance to open water, open water being the nearest channel or body of
water.
On each visit to a nest, the height of the eggs above the water,
the number of eggs and their condition; i.e., cold, warm, or pipped, were
recorded.

�-87-

Initiation dates of nests were classed into weekly intervals.
The
initiation dates of nests found during laying were calculated by allowing
one egg laid per day. For nests found after laying, the initiation date
was based on the date of hatch, assuming a 23-day incubation period and
one egg laid per day. Dates of hatch were also grouped into weekly
intervals, and hatch was defined as the time at which all eggs had hatched.
A successful nest was considered to be a nest in which one or more eggs
hatched.
Average clutch size and hatching success calculations were based
on data from successful nests only. Brood counts were made during regular
weekly counts.
To determine hunting pressure and harvest levels of coots, wing barrels
with appropriate signs were placed at waterfowl hunting areas in Colorado.
In addition, a questionnaire (Appendix) was designed to investigate hunter
attitudes.
Students from the Colorado State University Wildlife Techniques
class were assigned to check stations to question hunters leaving the
Wellington State Wildlife Area on the opening weekend of the waterfowl
season.
DESCRIPTION

OF STUDY AREAS

Four study areas were selected, one representing the eastern plains region,
two representing intermountain valleys, and one west of the Continental
Divide.
The Beebe Draw study area is in Weld County, T4N, R65W, Section 34, at an
elevation of 1,446 m. The dominant emergent vegetation consisted of
cattails.
Estimates of total wetland area and the emergent components for Beebe Draw
and the other study areas are presented in Table 1. For all areas, Scirpus
spp. refers primarily to hardstem bulrush (Scirpus acutus) and softstem
bulrush (~. validus).
For Lake John and Hog Lake, total areas given in
Table 1 represent that area searched for nesting coots. Total area in
these two study areas, including broad expanses of open water and emergents
not searched, was approximately 100 ha and 52 ha, respectively.
Aquatic sampling at Beebe Draw indicated the dominant aquatic was
Ceratophyllum demersum.
Potamogeton spp. were found at only 45 percent of
the sampling stations with a density rating of sparse at each. Maximum
water depth was 2.3 m and fluctuated approximately 120 cm between April
1977 to March 1978. The substrate consisted of sand and loamy sand.
The Lake John Annex study area is in Jackson County, north central Colorado,
T9N, R81W, Section 2, at an elevation of 2,500 m. Scirpus spp. represented
the dominant emergent.
The dominant aquatics were Char a spp. and
Potamogeton pectinatus.
Maximum water depth and fluctuation were 1.2 m
and 50 cm, respectively.
The substrate was composed primarily of sandy
loam and silty clay loam.

�-88-

of four study areas, Colorado,

1977.1/

Beebe Draw

Lake John

Ice Pond

Hog Lake

Typha spp.

3.45/47•9

0.01/0.4

0.4/44•4

0.04/1.2

Scirpus spp.

1.03/

2.76/99•6

0.34/37•8

2.06/61.5

2.26/31.4

0/0

0/0

0/

0.46/6.4

0/0

0.16/17•8

Total Area of
Emergents

7.20

2.77

0.90

3.35

Total Area of
Open Water and
Emergents

16.7

18.1

14.4

24.4

Table 1.

Vegetational

Sciq~us
americanus
Other

1/

- Upper number
emergents.

components

•
14 3

area in hectares,

lower number

percentage

0

1.25/

.
37 3

of total

Ice Pond, near Buena Vista, is in Chaffee County in the central mountain
region of Colorado, T13S, R78W, Sections 5, 6, 7, 8. Elevation is 2,425 m.
Emergents consisted of nearly equal proportions of Typha spp. and Scirpus
spp., and lesser amounts of spikerush (Eleocharis macrostachya), rushes
(Juncus balticus), and sedges (Carex spp.). Aquatic growth at Ice Pond
was the most abundant of all study areas; species present were Myriophyllum
exa1bescens, Polygonum amphibium, Potamogeton filaformis, and Ranunculus
aquati1is.
All received moderate to dense ratings.
Maximum water depth
and fluctuation were 1.5 m and 7.5 cm, respectively.
Information
concerning a soil survey for the Ice Pond area has not yet been obtained,
transects indicated a sand and gravel bottom.
The Hog Lake study area is in Moffat County, in extreme northwestern Colorado, TlON, Rl03W, Section 9. It is part of Brown's Park National Wildlife
Refuge.
Elevation is 1,634 m. Scirpus spp. represented the dominant
emergent.
The common reed (Phragmites communis) and salt grass (Distichlis
stricta) were also present, growing as emergents.
Myriophyllum exalbescens,
Potamogeton pectinatus and ~. vaginatus were common aquatics.
Maximum
water depth and fluctuations were 1.6 m and 25 cm, respectively.
No soil
survey exists for the Hog Lake area; aquatic transects indicated a silt
and clay substrate.
Water samples taken on all study areas during the 27 June to 1 July period
indicated basic water (pH&gt;7.0) in every instance (Table 2).

�Table 2.

Results of water analysis, coot study areas, June 1977.1/

Study Area

Free Acidity

Total
Acidity

Phenolphthalein
Alkalinity

Total
Alkalinity

Hardness

pH

Beebe Draw

0

17

0

308

410

9.3

Lake John Annex

0

0

0

120

513

10

Ice Pond

0

11

0

103

103

8.9

Hog Lake

0

0

34

239

599

9.8
I

cc

\0
I

1/ Except for pH, all results are expressed as mg of CaC03/liter of H2O.

�-90-

In order to standardize comparisons and facilitate evaluations between
areas, each study area was classified according to the new wetlands and
deep water habitat classification system for the United States (Cowardin
et ale 1977). Under this system, it was necessary to divide Lake John
Annex, Ice Pond, and Hog Lake into two systems, one representing the
lake-like portions, the other the marsh portions (Table 3).
RESULTS AND DISCUSSION
Distribution

and Status

Data obtained from 107 returned questionnaires, field investigations, and
published records were compiled to determine coot distribution and status
according to the latilong system (Table 4).
Coots were found to breed throughout most of Colorado.
Major breeding
areas were Brown's Park on the western slope, North Park, and the San
Luis Valley.
On the plains, coots were locally abundant where suitable
habitat occurred.
Nesting coots were primarily associated with cattail
or bulrush marshes.
Small numbers may breed in high altitude lakes
dominated by sedges or willows (Salix spp.), such as Lake San Cristobal
near Lake City or Beaver Lake near Marble.
Coots were resident year
round in low numbers «1,000) mainly along the western boundary of the
plains, from near Fort Collins south to the Pueblo area.
The presence
of wintering coots was dependent on the severity of winter weather.
Coots, present in the Fort Collins area durLng the winter of 1976-77,
were absent during the 1977-78 winter when water areas froze over.
Spring Migration
Weekly counts were initiated in 1977 to determine timing of migration at
only two areas, Beebe Draw and Lake John. Restrictions due to course
requirements limited visits to Ice Pond, while Hog Lake was not selected
as a study area until well after spring migration had ended.
The first signs of migrational movement in 1977 were noted at Beebe Draw
during the 14-20 March period (Table 5). Peak numbers were recorded
during the week of 11-17 April.
Coots first arrived in the North Park
area during the week of 4-10 April.
Migration pe2ked during the 18-24
April period, one week later than at Beebe Draw. At both locations, coot
numbers decreased and then stabilized by the third week after the peak
had occurred.
The range of migrational movement extended over seven
weeks at Beebe Draw, but only five weeks at Lake John.
Ice Pond was first visited on 25 April, weeks after ice thaw and the
arrival of coots.
It is assumed the peak of migration occurred prior to
25 April.
Hog Lake was not visited until 23 May, when 200 coots were observed.
However, counts by Refuge personnel over the 14 March to 22 May period give
some indication of coot movements.
It would appear coots first arrived
at Hog Lake during the first two weeks of March, and peaked during the
first two or three weeks of April.

�Table 3.

Wetlands and deep water habitat classifications of study areas.

Area

System

Subsystem

Class

Subclass

Dominance
Type

Water Regime
Semiperm. flooded

Beebe Draw

Palustrine

None

Emergent
wetland

Persistent

Typha spp.
Scirpus spp.

Lake John Annex

Pauls trine

None

Emergent
wetland

Persistent

Scirp_u_~a_c_1..!~'!~Semiperm. flooded

Lacustrine

Littoral

Aquatic bed

Submerged
algal

Cha~a spp.

Perm. flooded

Palustrine

None

Emergent
wetland

Persistent

Typha spp.
Scirpus spp.

Perm. flooded

Lacustrine

Littoral

Aquatic bed

Submerged
vascular

MxrioEhxllum
exalbescens
Polygonum
amEhibium

Perm. flooded

Palustrine

None

Emergent
wetland

Persistent

Scirpus acutus

Semiperm. flooded

Lacustrine

Littoral

Aquatic bed

Submerged
vascular

M~rioEhxllum
exalbescens
Potamogeton spp.

Perm. flooded

Ice Pond

Hog Lake

I
1.0
I-'
I

�-92-

and status in Colorado.-1/

Table 4.

Coot distribution

B

B

B

R

R

B

B

B

B

B

R

R

R

R

B

B

B

b

R

R

B

B

R

B

B

B

b

B

1/ Classification

by latilong blocks follows the system of Kingery and Graul
R = resident year round, B = breeding, b = very likely breeding,
bird found in suitable habitat.

1978.

Counts conducted in 1978 indicated migrational movements beginning by midMarch on two areas, and somewhat sooner at Hog Lake (Table 6). Lake John
was still frozen on 31 March. Water levels at Beebe Draw were extremely
low, supposedly in preparation for dragline operations.
The extent to which
this might affect migration is unknown, therefore marshes peripheral to
Beebe Draw will be monitored.
Nesting
By the end of August, 211 nests had been located; 77 at Beebe Draw, 67 at
Lake John Annex, 30 at Ice Pond, and 37 at Hog Lake. Tables 7 and 8 present
nesting chronology of coots on all study areas. Hog Lake was not searched
until after the peak of hatch occurred, therefore data from that area are
not comparable.
At Beebe Draw, the peak of nest initiation occurred
during the week of 25 April, at Lake John the peak occurred during the week
of 6 June, six weeks later than at Beebe Draw. The peak was less distinct
at Ice Pond, with six nests initiated during the week of 9 May. Due to
the small number of nests for which initiation dates could be determined,
the peak was unknown for Hog Lake. Hatching was completed by the end of
July on all areas except Hog Lake. One nest was still active there on 2
August.
This nest hatched successfully.
The peak of hatch occurred during
the week of 30 May for Beebe Draw, 4 July for Lake John, and 30 May through
12 June at Ice Pond. The first complete search of Hog Lake was made on 7
June, at which time 19 of 32 nests located had already hatched.
This would
indicate a nesting season well ahead of Lake John and possibly similar to
Beebe Draw.
Table 9 presents a summary of coot breeding data. Table 10 illustrates the
frequency distribution of clutch size. These data show variability in
modal clutch size from a high of nine for Beebe Draw to a low of six for
Lake John. Over 90 percent of the nests contained 5 to 11 eggs each.

�Table 5.

Coot numbers observed during spring migration on study areas, 1977.
Week Beginning
AEril
11
18
25

2

9

16

23

115

81

54

50

95

394

172

172

199

75

85

280

200

Location

14

March
21

Beebe Draw

15

58

105

132

178

168

Lake John

-

-

-

- }j

146

851

706

Ice Pond

-

-

-

-

-

-

305

2/
Hog Lake -

100

-

500

-

680

28

4

I

May

I

'"

W

I

~/ Coots first arrived in North Park (C. Braun, pers. comm.).
2/
- Counts from 14 March to 16 May conducted by Refuge personnel.

345

�-94-

Table 6.

Location

Beebe Draw

Coot numbers

observed during weekly counts, Colorado,

27 February

0

Week Beginning
6 March
13 March

0

1978.

20 March

5

11

Lake John

0

0

Ice Pond

31

79

179

307

Hog LakJJ

5

35

1/
- Counts for weeks beginning 27 February and 6 March conducted by Brown's
Park National Wildlife Refuge personnel.

The fate of coot nests and eggs in 1977 was examined to delineate losses
(Tables 11, 12). Overall nesting success was 82 percent.
For all study
areas, approximately 76 percent of the unsuccessful nests were destroyed
by predators.
This was especially apparent at Lake John where about 88
percent of the unsuccessful nests were destroyed.
High nesting success
of coots is often attributed to defense of the nest by both adults and the
nest location over water.
However, predation was the major cause of nest
failure in this study in 1977.
Most egg loss was attributed to predation and desertion.
Desertion may
have been human or predator induced, or the result of infertile or addled
eggs. In addition, eggs found in the water may have been infertile or
addled.
Crawford (1975) found that frequent nest checks did not cause
excessive egg abandonment.
At Lake John almost 39 percent of all eggs
laid were lost, compared to an average of less than 12 percent for all
other areas.
The average number of coot eggs lost per successful nest of
known size was 0.80. When unsuccessful nests were included, the average
number of eggs lost per nest increased to 1.47.
Statistical analyses of selected breeding data were conducted to determine
if differences existed between study areas. Average clutch size varied
from a high of 8.7 for Beebe Draw to a low of 6.5 for Hog Lake. Tests for
analysis of variance followed by a Student-Newman-Keuls
multiple range test
(Sokal and Rohlf 1969) indicated a significant difference existed (P&lt;0.05)
in clutch size only between Beebe Draw and Hog Lake. Nest data from Hog
Lake were derived from late season nests which probably accounts for the
difference.
Other investigators (Crawford 1975, Fredrickson 1970, Gullion
1954) reported that clutch size of coots varied seasonally.
An examination of clutch size in relation to date of nest Lnf.t La t Lon suggested a decrease in clutch size as the season progressed.
For example,
weekly average clutch size at Beebe Draw decreased from a high of 9.7
early in the season to a low of 5.0 late in the season (Table 13).

�Table 7.

Number of coot nests initiated per week on four study areas, Colorado, 1977.
Week Beginning
30 May 6 June 13 Jun

20 Jun

27 June 4 Ju1 11 Ju1 18 Ju1

18 Apr

25 Apr

2 May

9 May

16 May

23 May

Beebe Draw

1

17

14

14

10

2

3

5

0

0

0

0

0

0

Lake John

0

0

3

0

3

12

13

19

10

3

0

0

1

1

Ice Pond

0

5

3

6

5

4

0

2

2

2

0

1

0

0

Hog Lake

?

?

?

3

?

2

1

?

2

0

0

0

0

0

Area

I
1.0
VI
I

Table 8.

Number of coot nests hatching per week on four study areas, Colorado, 1977.
Week Beginning
20 June
13 June

-------

---

4 July

11 July

18 July

5

2

5

0

0

2

8

8

13

9

1

6

5

4

2

1

2

1

5

3

1

2

0

2

0

30 May

Beebe Draw

10

23

15

7

Lake John

0

0

0

Ice Pond

0

6

Hog Lake

?

?

Area

~~--.

27 June

23 May

6 June

�Table 9.

Coot breeding

data, Colorado,

1977.

Beebe Draw

Lake John

Ice Pond

Hog Lake

Overall

Number of nests

77

67

30

37

211

Successful

69

41

27

36

173

89.6

61.2

90.0

97.3

82.0

546

326

212

52

1,136

8.7 + 1.9

8.0 + 2.2

7.8 + 2.8

6.5 + 1.0

8.2 + 2.2

3-14

4-16

2-14

5-8

2-16

26

59

21

5

111

95.2

81.9

90.1

90.4

90.2

Nesting

nests

success

(%)

1/

Number of eggsAverage

clutch siz~/

Range of clutch sizes
Number of unhatched
Hatching

success

3/

eggs-

(%)

1/ Number

of eggs based on successful
Pond, and 8 - Hog Lake.

1/ Mean + one standard
3/

-

Number

of unhatched

deviation

nests of known size: 63 - Beebe Draw; 41 - Lake John, 27 - Ice

of mean.

eggs for successful

nests only.

I
\0
0\
I

�-97-

Table 10.
1977 .

Frequency of occurrence of clutch size for the coot in Colorado,

Clutch Size

Beebe Draw

Ice Pond

Lake John

Hog Lake

Overall

Percent

2

0

1

0

0

1

0.7

3

1

1

0

0

2

1.4

4

3

0

1

0

4

2.9

5

1

3

2

2

8

5.8

6

3

3

10

1

17

12.2

7

7

2

7

4

20

14.4

8

11

9

7

1

28

20.1

9

14

1

6

0

21

15.1

10

12

2

3

0

17

12.2

11

9

3

3

0

15

10.8

12

1

1

0

0

2

1.4

13

0

0

0

0

0

14

1

1

1

0

3

15

0

0

0

0

0

16

0

0

1

0

1

2.2

0.7

�-98-

Table 11.

Fate of coot nests, four study areas, Colorado,

1977.

Number of Nests
Lake John
Ice Pond
Hog Lake

Nest Fate

Beebe Draw

Successful

69

41

27

36

173

Unsuccessful

8

26

3

1

38

Lost to Predators

5

23

0

1

29

Avian

0

2

0

0

2

Manunalian

1

0

0

0

1

Unknown

4

21

0

1

26

1

3

1

0

5

2

0

2

0

4

77

67

30

37

211

Deserted
Unknown

failure

Total nests

Total

�-99-

Table 12.

Fate of coot eggs, four study areas, Colorado,

Number of Eggs
Lake John
Ice Pond

1/
1977.-

Hog Lake

Total

191

47

1,025

98

o

o

126

o

5

o

o

5

Mammalian

9

o

o

o

9

Unknown

19

93

o

o

112

Deserted

10

38

13

3

64

Dead embryo

2

o

o

o

2

Buried in nest

o

2

o

o

2

In water

8

17

4

1

30

Broken

1

2

2

o

5

16

11

10

1

38

Total lost

65

168

29

5

267

Total laid

585

435

220

52

1,292

Egg Fate

Beebe Draw

Hatched

520

267

Lost to predators

28

Avian

Unknown

loss

-1/Eggs from successful and unsuccessful
nests of known size only.

nests, except Hog Lake - successful

�-100-

Table 13.
1977 .
Clutch
Size

Clutch size in relation to date of nest initiation, Beebe Draw,

Week Beginning
IS Apr

25 Apr

2 May

9 May

14

16 May

23 May

30 May

6 June

1

1

2

1

13
12

1

11

3

5

1

10

5

2

5

6

2

3

2

2

2

3

4

9

1

S
7

1

2

6

2

5

1
1

4

2

3

1
1

N=l

N=17

N=13

N=14

N=9

N=l

N=3

N=5

x=9.0

x=9.7

x=9.5

x=S.S

x=S.4

x=7.0

x=5.0

x=5.4

�-101-

T-tests for differences in average clutch size before and after the median
date of nest initiation at Beebe Draw and Ice Pond indicated a highly
significant decrease (P&lt;O.Ol) in clutch size on both areas. Data from Hog
Lake could not be tested because they were incomplete.
Data from Lake John
were not tested due to predation and probable renesting, which may have
affected clutch size.
On each visit to a nest, the height of the eggs above the water was
measured to examine effects of water fluctuations on nest survival.
Field
observations indicated that coots increased the height of the nest and eggs
above water as the laying-incubation period progressed.
At-test
indicated a highly significant increase (P&lt;O.Ol) in the average height of
the eggs above water from the time of nest initiation to late in incubation.
A basic assumption was that nests were free to move up or down with the
supporting vegetation as water levels fluctuated.
Field observations
supported this assumption, no nest failure or egg loss could be attributed
to water level changes.
Habitat Use
Table 14 presents number and percentage of nests by cover type. Overall,
Scirpus was the preferred nest cover type. At Beebe Draw, 61 percent of
nests found were in cattail, which represented about 48 percent of the
emergent cover.
Scirpus (about 14 percent of emergent cover), was utilized
to an extent greater than its occurrence, having about 31 percent of the
nests.
A similar situation existed at Ice Pond. Bulrush was the only
nesting cover utilized at Lake John. At Hog Lake, the majority of nests
were found in bulrush, which represented the dominant robust emergent.
It
should be noted however, that nearly every stand of cattail at Hog Lake
contained a coot nest.

Table 14. Number and percentage
areas, Colorado, 1977. !/

of coot nests by cover type of four study

Beebe Draw

Lake John

Ice Pond

Hog Lake

Overall

Scirpus spp.

24/31.2

67/100

18/60.0

34/91•9

143/67.8

Typha spp.

47/61.0

0/0

10/33•3

3/8•1

60/28•4

6/7.8

0/0

1/3.3

0/0

7/3•3

0/0

0/0

1/3•3

0/

1/0.5

Scirpus and
Typha Mixture
Other

1/

Upper number

number of nests, lower number

percentage

0

of nests.

�-102-

Nest cover type in relation to date of initiation was examined for Ice
Pond and Beebe Draw (Table 15). Aproximately 89 percent of the nests
initiated before the peak at Beebe Draw were in cattail.
No such preference
was noted at Ice Pond. After the peak of nest initiation, cover preference
became less distinct.

Table 15.

Nest cover type in relation

to date of nest initiation.!/

Before Peak

After Peak

Scirpus spp.

1/7

18/

Typha spp.

16/7

26/

1/0

4/1

0/0

0/1

Scirpus and Typha
Carex spp.

1/

Upper number

Mixture

nests at Beebe Draw, lower number

11
3

nests at Ice Pond.

Characteristics of cover utilized by coots at nest sites are illustrated in
Tables 16-18. Measurements recorded and analyzed were taken at the time
nests were first located to illustrate conditions at the time of nest
initiation.
Nests found late in the hatch or after hatching were not
included.
The numbers 0-4 (Table 16) indicate concealment afforded by the
vegetation at the nest site on none, one, two, three, or four sides. The
classifications lA, 2A, 3A, 4A, indicate concealment on one side and from
above, two sides and above, etc. The average height of vegetation at the
nest site was: Beebe Draw - 60.2 cm, Lake John - 38.7 cm, Ice Pond - 76.8 cm,
and Hog Lake - 62.5 cm. Table 17 indicates water depth by class at the
nest site. The average depth at the nest was: Beebe Draw - 71.0 cm, Lake
John - 50.5 cm, Ice Pond - 36.2 cm, and Hog Lake - 36.2 cm. The distance
from the nest to open water by class is indicated in Table 18. The average
distance of the nest from open water was: Beebe Draw - 8.2 m, Lake John 4.4 m, Ice Pond - 3.1 m, and Hog Lake - 3.7.
Nest site measurements at Lake John indicated many nests were located in
poor cover and in shallow water compared to other areas. These factors may
have allowed better nest visibility and accessibility to predators and
resulted in the high nest failure at Lake John. Chi-square tests indicated
nest success or failure was independent of both cover class and water depth
class (P&gt;0.05). However, the strength of the independence of the cover
class-nest success or failure relationship was not great enough to warrant
complete acceptance (0.05&lt;P&lt;0.10).

�-103-

Table 16.

Concealment

at coot nest sites, four study areas, Colorado,

1977.

Sides

Beebe Draw

Lake John

Ice Pond

Hog Lake

0

1

29

0

2

1

7

5

0

1

2

11

10

0

1

3

12

9

6

0

4

17

2

6

1

1A

0

0

0

0

2A

1

2

0

0

3A

6

2

3

0

4A

20

8

15

2

Table 17.
1977 .

Depth

Number of coot nests by water depth, four study areas, Colorado,

Beebe Draw

Lake John

Ice Pond

Hog Lake

0-50 cm

1

38

27

14

50-100 cm

71

25

2

o

&gt;100 cm

4

4

1

o

Table 18.' Number of coot nests by distance
areas, Colorado, 1977.

Distance

from open water,

four study

Beebe Draw

Lake John

Ice Pond

Hog Lake

0-2 m

10

34

17

16

2-4 m

16

7

5

14

&gt;4 m

51

26

8

7

�-104-

Breeding

Densities

Breeding densities (Table 19) were based on the number of successful nests
per hectare of emergent habitat.
In addition, the breeding density within
a given emergent component was examined.
Density based on total emergent
area included species which were little used or not used for nesting, i.e.,
Carex spp., Distichlis stricta, Eleocharis spp., Phragmites
communis, and
Scirpus americanus.
A better comparison of breeding densities between areas
is one based on the common robust emergents, i.e" Scirpus and Typha spp.
On that basis, the breeding densities for three study areas were similar
with that for Ice Pond being at least twice as great.
The reasons for this
are unclear.
Ryder (1958), Weller and Spatcher (1965), and Weller and
Fredrickson (1973) noted that coots were most abundant at a cover-water
ratio of 50:50.
Sparsely vegetated marshes with numerous pools of open
water appeared most attractive to nesting coots. None of the marshes in
this study had a cover-water ratio of 50:50.

Table 19. Breeding
emergent habitat.

density as number of successful

nests per hectare of

Cover Type

Beebe Draw

Lake John

Ice Pond

Hog Lake

Scirpus spp.

16.5

14.8

47.0

16.0

Typha sPP.

13 .6

0

22.5

75.0

Combined Scirpus
and Typha

15.4

14.8

35.1

17.1

Carex spp.

0

0

16.7

0

Total Emergents

9.6

14.8

30.0

10.7

In March and April 1977, segments of the emergent zone at Beebe Draw were
burned.
Burning failed to kill any of the emergents, but did clear most
litter from the previous year's growth.
The burned area of 3.13 ha consisted
mostly of Scirpus americanus.
No nests were found in the three-square,
although brood platforms and foraging broods were observed.
Six nests
(representing all the nests found in the burned area) were found in 0.73 ha
of burned cattail.
The resultant breeding density of 8.2 for the burned
cattail was substantially less than that of the remaining unburned cattail
and bulrush with 16.9 successful nests per hectare.
Nests in burned areas
were initiated somewhat later than most other nests, possibly due to the
lack of cover before new vegetative growth had sufficiently developed.

�-105~

The owners of Beebe Draw have undertaken an emergent removal program.
In
September a portion of the emergents along the west shore were removed
using a dragline.
In December the water was drawn down. Future plans
include use of the dragline along the south shore.
If completed,it
will
allow comparison of coot breeding densities before and after treatment.
However, by the last week of March 1978 operations had not commenced.
Brood Counts
Brood counts of coots were initiated on all areas during June and continued
through August.
Results were difficult to interpret.
Gullion (1956)
suggested caution in the evaluation of brood counts.
He cited the tendency
of parent coots to split up the brood and to feed them in different parts
of their territory, often hidden from view. Another problem was the
difficulty in distinguishing beteen adults and older immatures over long
distances and often poor light conditions involved.
The largest number of
young recorded on one occasion was 76 for Beebe Draw, 17 for Lake John, 76
for Ice Pond, and 97 for Hog Lake. The low number of immatures recorded at
Lake John could possibly indicate poor brood survival.
Fall Migration
Weekly counts were conducted to determine summer movements and initiation of
migration (Table 20). Counts at Beebe Draw indicated decreasing numbers of
coots through July and continuing low numbers through most of August.
However, three marshes adjacent to Beebe Draw showed substantial increases
beginning in August, continuing into September, with each holding up to 150
coots. This may represent a late summer influx from other areas and/or
movement from Beebe Draw in response to an apparent poor supply of ~quatic
vegetation.
Coot numbers increased gradually at Ice Pond beginning in late
August, and fluctuated at Hog Lake after an initial influx in June. A
considerable buildup was observed at Lake John starting in early August.
Fall migration peaked during the week beginning 12 September at Beebe Draw
and Ice Pond, and the week of 19 September at Lake John. The peak occurred
earlier at Hog Lake with high numbers recorded during the weeks of 22 August
and 5 September.
Counts were discontinued on all study areas by the end of
October.
An attempt to visit all study areas on 19-20 November was cut short due to
a winter storm.
Ice Pond was visited on 19 November, it was still ice free
and eight coots were present.
Correspondence with Brown's Park personnel
indicated only 45-50 coots were still present on Hog Lake by 11 November.
Harvest
In Colorado, waterfowl harvest records are derived from mail surveys sent to
approximately three percent of the small game license buyers.
Colorado
Small Game Hunter Harvest surveys from 1954 through 1972 did not give specific
harvest figures for coots, but instead grouped them in the category of "Others

��-107-

and Unknown."
This group included coots, mergansers, goldeneyes,buffleheads and unknowns.
Data from more recent small game harvest surveys are
presented in Table 21 (H. Riffel, pers. corum.). An estimated 4,645 coots
were harvested in Colorado in 1976 (Sorensen et al. 1977). Despite liberal
daily bag limits of 15 and 25 in the Central and Pacific Flyways,respectively, coot harvest is low in Colorado.

Table 21.

Year

Coot harvest

in Colorado

Estimated

1973-76.

Harvest

Percent

of Duck Harvest

1973

4,579

1.50

1974

5,054

1.41

1975

2,924

1.16

1976

4,645

2.29

To further determine hunting pressure and harvest levels of coots, wing
barrels with appropriate signs were placed at Lake John (1), Walden Reservoir
(2), Cowdrey (1), MacFarlane Reservoir (1), Antero Reservoir (1), Hog Lake
(1), and Wellington (2). Wings deposited in barrels over the 1-14 October
period indicated a harvest of 443 ducks and eight coots, or a coot harvest
representing 1. 8 percent of the duck harvest.
To investigate hunter attitudes, wildlife students from Colorado State University questioned hunters at the Wellington State Wildlife Area on 1-2
October 1977. Of 86 hunters polled, only 15 indicated they normally hunt
coots. Of the remainder, 36 cited bad taste as the reason for not hunting
coots.
The general concensus was that coots were not worth shooting and
that ducks were preferred.
The results were imilar to those obtained at
Wellington in 1976 when only 11 of 54 hunters indicated they normally
hunted coots.

LITERATURE

CITED

Atlantic Waterfowl Council.
1972. Techniques handbook of waterfowl
development and management.
Habitat Development and Management
Atlantic Waterfowl Council.
217 pp.
Bailey, A. M., and R. J. Niedrach.
Nat. Hist., Denver.
Vol. I.

1965. Birds of Colorado.
454 pp.

habitat
Corum.

Denver Mus.

1977. ClassifiCowardin, L. M., V. Carter, F. C. Golet, and E. T. LaRoe.
cation of wetlands and deep-water habitats of the United States.
(An
operational draft).
U. S. Fish. Wi1dl. Servo 100 pp.

�-108-

Crawford, R. D. 1975. Breeding biology of American coots in relation to
age. Ph. D. Thesis. Iowa State Univ., Ames. 42 pp.
Fredrickson, L. H. 1970. Breeding biology of American coots in Iowa..
Wilson Bull. 82(4):445-457.
_____ , J. M. Anderson, F. M. Kozlik, and R. A. Ryder.

1977. American coot
(Fulica americana). Pages 123-147 in G. C. Sanderson, ed. Management
of migratory shore and upland game birds in North America. Int. Assoc.
Fish and Wildl. Agencies. Washington, DC.

Gullion, G. w. 1954. The reproductive cycle of American coots in California. Auk 71(4):366-412.
1956. An observation concerning the validity of coot brood counts.
J. Wildl. Manage. 20(4):465-466.
Harrington, H. D. 1964.
Swallow Press, Inc.

Manual of the plants of Colorado.
Chicago. 666 pp.

2nd Ed.

The

Kingery, H. E., and W. D. Graul, eds. 1978. Colorado bird distribution
latilong study. Colo. Field Ornithologists and Colo. Div. Wildl.
Denver. 58 pp.
Ryder, R. A.
Thesis.

1958. Coot-waterfowl relationships in northern Utah.
Utah State Univ., Logan. 219 pp.

Sclater, W. L. 1912. A history of the birds of Colorado.
Co. London. 576 pp.
Sokal, R. R., and F. J. Rohlf.
San Francisco. 776 pp.

1969.

Biometry.

Ph. D.

Witherby and

W. H. Freeman and Company.

Sorensen, M. F., S. M. Carney, and E. M. Martin. 1977. Waterfowl harvest
and hunter activity in the United States during the 1976 hunting
season. U. S. Fish Wildl. Servo Admin. Rep. 26 pp.
Weller, M. W., and L. H. Fredrickson. 1973. Avian ecology of a managed
glacial marsh. The Living Bird 12:269-291.
Weller, M. W., and C. S. Spatcher. 1965. Role of habitat in the distribution and abundance of marsh birds. Iowa State Univ. Agr. and Home
Econ. Exp. Sta., Spec. Rep. 43:269-291.

Prepared

byJt/~dJ~
~
Warner Paul Gorenzel
Graduate Research Asst.

Approved bY_--=~_~---=----_"
--=2.==----_" ..LL~~~::::::::..:::=::..
Clait E. Braun
Wildlife Researcher

_

�-109-

APPENDIX

�-rioSURVEY OF COOTS IN COLORADO

1.

District

2~

To your knowledge, do coots occur in your present district? Yes___
If no, please answer number 9 only and return survey form.

3.

If coots are known to occur in your district, please list major loeations
where they have been observed~
_

40

Please list wetlands
pairs of coots

50

Which wetland vegetation types occur where you see coots during June and
July (Leo cattails, bulrush, willow" etc.?)
_

6.

WhiCh month(s)

~----------------------------------------------------------No

where you have observed at least 10-15 or more breeding
~
_

of the year are coots most commonly present

in your district?

10 Have you encountered coots in your district during 1 December through
31 March?

8.

9..

Yes

No

If yess where?

_

Have you encountered any hunters who retain coots in the bag in your
district?
Yes_
No
• If yes, where?

_

0

List any major tvetlands in your district where you de not normally observe
coots
_

�-111-

Survey of Coots in Colorado

1.

COl.mty(or counties)

_

2.

To your knowledge, do coots occur in the area you listed

above?

._--

Yes

No

If no, please answer mnnber 8 only and return survey form.

3.

Do coots breed :in your area?
If yes, please list

Yes

No

_

wetlands where you have observed at least

10-15 or nore breeding pairs of coots °

Please be specific.

4.

Please llst

5.

Whichwetland vegetation types occur where you see coots during
June and July (i.e. cattails, bulrush, willow, etc,o)

other major areas where coots have been observed.

_

6.

Whichmonth(s) of the year are coots mist carmonly present in your
area?_·
--------

1.

Are coots present :in your area during most or all of the 1 December
through 31 Marchperiod?

8.

Yes

No

If yes, where?

List any major wetlands in your area where you do not normally observe
coots.

--------------------------------------

�-112-

Observer

-------------------------------~ Date

Location

-------~-------------Time

Estimated Cloud Cover -------~%
Wind:

None

Precip:

Medium

Strong

Rain

Mist

Sleet

Gusty
Hail

Hrs.

FO or CO

Temp

Light

None

to_____

Direction
Snow

Fog

Steady

Intermit
Number Observed:
Coots

Adults

--------

----------- Imma----------------

Ducks:
Mal

Pin

GHT

BWT

Cin Teal

Gad

Wid

sMv ----

Geese

Rud

Buffl

Red

Scaup

Ringneck

_

Other

_

_

G-Eye.

_

Can

------------------------------

Other Speci es :

Other Conditions, Changes, Comments:

_

-----------------------------

May 1977

�Check Station #

Location
Date

T~e

St~rt

. Name of Observer

Time Stop

------------------

ASK ALL HUNTERS LEAVING WITH COOTS:
Do you
normally
hunt coots?
-

If no,
why?
Comments

-------------------------

/I
Bagged

(/Hit
Not
Retrieved

ASK ALL HUNTERS LEAVING WITHOUT COOTSz
Hra.
Spent
Hunting

Do you
ever hunt
coots?

If no ,
why?
Comments

Hrs.
Spent
Hunting

--

I

lI\,...;

I

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                  <text>January,

-1-

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Work Plan No.
Job Title
Period

Bighorn

W-41-R-28

1979

Sheep and Mt. Goat Investigations

Job No.
16
1
Evaluation of Trace Mineral Availability
~t~o~B~l~·cgh~o~r~n~S~h~e~e~p~o~n~~W~i~n~t~e~r~R~a~n~ga=e~s~i~n~C~o~l~o~r~a~d
-_

----------~---------------------

Covered: June 1, 1977 - June 30, 1978

Personnel:

Robert

E. Keiss

ABSTRACT
...I'r ace mineral formulations were d ave Loped and tested for animal preference
and physiological effect upon the animal.
It became increasingly difficult
to evaluate the effects of trace mineral supplementation
on a wild ruminant
population so two biological parameters were tested to measure the amount
of trace mineral and do preliminary evaluations as to the feasibility of
using these for biological measurements.
Deer and elk hair samples collected
from throughout Colorado were evaluated for trace mineral content as well as
bighorn'sheep,fecal
samples.
-:

,,: ...

,:".

~:.;,.

��-3-

Evaluation of Trace Mineral Availability
to Bighorn Sheep on Winter Ranges in Colorado
Robert E. Keiss
P. N. OBJECTIVE
The objective of this study is primarily directed towards evaluating certain
nutritional supplements on bighorn sheep herds in an effort to improve lamb
survival and viability of bighorn sheep to bring about an overall increase
in the total numbers of animals within the State of Colorado.
These general
objectives will be attempted by implementing and evaluating the following:
1.

Develop trace mineral formulations which will be used and
accepted by bighorn sheep on their native ranges.

2.

Measure the amount of each mineral mix formulation used by
bighorn sheep herds to ascertain seasonal preferences and
consumption.

3.

Plan research facilities which can be used for trace mineral
metabolism studies in bighorn sheep. These facilities will
be utilized to measure the nutritional requirements of bighorn
sheep.

4.

Evaluate the possibility of using trace mineral mixes and the
establishment of "artificial mineral licks" as a possible
carrier for chemotherapeutic drugs for treatment of bighorn
sheep against Protostrongylus infestation.
SEGMENT OBJECTIVES

1.

Formulate a trace mineral mix which can be placed on bighorn sheep winter
ranges to supplement the nutritional trace mineral requirements of bighorns.

2.

Measure the amounts of trace mineral mix consumed by bighorn
determine seasonal preferences.

sheep and

3.

Measure trace mineral content of bighorn
fecal content to potential trace mineral

and relate

4.

Determine the effects of trace mineral mix consumption
of bighorn sheep.

sheep fecal material
deficiencies.

on the physiology

Procedures
In their publication, "The Biochemistry of Blue, Snow, and Ross' Geese",
Hanson and Jones (1976) state "There are, in effect, three general kinds of
approaches to gaining insights into the mineral metabolism of animals:
1)

�-4-

in vitro studies, exemplified by the use of excised sections of intestine
or the incubation of isolated cells, portions of cell membranes or of a
specific organelle: 2) laboratory studies using living, but confined,
orthodox experimental animals; and 3) studies of free-ranging wild animals,
an approach particularly exemplified by investigations of the mineral content of the keratin structures.
Although admitting that the objectives
and precision of the latter two approaches differ
from each other as does
also the variability of their environments, the third approach affords the
opportunity to profit from the results of an almost infinite number and
variety of natural experiments that a lifetime of laboratory work with caged
animals could not duplicate."
During the last segment of this project (W-41-R-27) procedures were described
on the development and utilization of experimental trace mineral supplements
for bighorn sheep on winter ranges. Without the benefit of large numbers of
bighorn sheep to use as experimental animals and elaborate pen facilities to
conduct strictly controlled trace mineral experiments, it has become increasingly difficult to analyze or measure any physiological changes in those
animals consuming trace mineral supplements.
Several different parameters on the wild-free ranging animal can be measured
which will give a index of the animal's mineral status. Among these parameters
are hair and feces. During this segment these two parameters were measured
in an effort to test the feasibility of using them for detecting trace mineral
deficiencies in wild ungulate populations.
Hair is composed of keratin and the ability of hair to act as a recording
filament of post nutritional state of the animal has been reported by Flyn
(1974).
It is evident that the mineral content of hair reflects the minerals
found in the animals environment and that the use of hair can be used to
evaluate the adequacy of the mineral diet.
Warner and Anke (1960) found that levels of iron, manganese, copper, molybdenum,
and cobalt in cattle hair were representative of amounts of these elements in
the forages from soils of diverse parent materials.
Anke (1967) established
the following levels as representative of dietary sufficiency for dairy cattle:
magnesium, 750-800 ppm; sodium, 200 ppm; zinc, 115 ppm; manganese, 4.0 ppm;
copper, 7 ppm; molybdenum 0.35 ppm; and cobalt, 0.05 ppm.
During the 1975 big game hunting season a state-wide collection of hair samples
from both deer and elk was made. These samples were collected at big game
check stations and an effort was made to collect at least ten samples from
male animals from each of the big game management areas where the animals were
harvested.
Approximately 5 grams of hair was plucked from the hind quarter
and placed in paper sacks. These sacks were identified as to game management
unit and returned to the Fort Collins laboratory.
Hair samples were removed and washed in a mild detergent using an ultrasonic
cleaner.
The samples were subjected to the cleaner for approximately one
minute, followed by a liberal rinsing in tap water.
The tap water rinse was
followed by 4 changes of distilled water.
A final rinsing was made using
diethyl ether.
These treatments were employed to eliminate exogenous trace
minerals on hair.

�-5-

Approximately 0.5 gm hair was placed in a glass, screw cap test tube and
digested using 3 ml of a digestion mixture (2 parts HN03 and 1 part HCL04).
Pre-digestion was allowed to proceed overnight with the screw caps loosened
on the tubes. The caps were then tightened securely and the tubes placed
in a waterbath at 700C and allowed to digest, under pressure, for approximately
4 hours.
Appropriate dilutions were made with distilled water and all determinations
were made using a Perkin-Elmer 303 atomic absorption spectrophotometer.
The following elements were determined from each of the deer and elk hair
samples:
Copper, Iron, Magnesium, Zinc, Calcium, Potassium, and Sodium.
(Manganese levels were below detectable limits for the sample dilutions
used).
Sample~ were collected on the basis of the Big Game Management Units and
all chemical analyses were carried out on these samples.
During the data
anlaysis phase of this study, it became evident that comparisons of data
between animals from these areas was not practical because these game management areas are not necessarily based upon any biological herd basis.
All
data has been grouped into the Data Analysis Unit areas and comparisons made
(Table 1 and 2). Because of this re-grouping of data it resulted in unequal
subgroups, but the advantages outweighed the disadvantages.

Table 1.

Deer Data Analysis

Units

(DAU).

DAU

DAU

Name

and

D-l
D-2
D-3
D-4
D-6
D-7

Little Snake
Bear's Ears
North Park
Red Feather
Rangely
White River

D-8
D-9
D-lO
D-11
D-12
D-l3
D-14
D-15
D-16
D-17

State Bridge
Middle Park
Boulder
Book Cliffs
Grand Mesa
Maroon Bells
Red Tables
Cottonwood Creek
Cripple Creek
Bailey

Game Management

Units

Included

(1, 2, 201)
(4, 5, 14, 441)
(6, 16, 17, 161)
(7, 8, 9, 19)
(10, 21)
(11, 12, 13, 22, 23, 24, 25, 26,
31, 32, 33, 34, l31)
(15, 35, 36)
(18, 28, 27, 37)
(20, 29, 38, 381 )
(30)
(41, 42, 52, 411, 521)
(43, 47, 471)
(44, 45)
(48, 56, 481)
(49, 57, 58, 581, 59)
(39, 46, 50, 51, 391, 461, 501,
500, 511)

�-6-

Table 1.

Deer data anlaysis units.

(Continued).

DAU

DAU

D-18
D-19
D-20
D-21
D-22
D-23
D-24
D-25
D-26
D-27
D-29
D-30
D-31
D-32
D-33

Glade Park
Uncompahgre
Crawford
West Elk
Taylor Park
LaSal
Ground Hog
Powderhorn Creek
Saguache
Westcliffe
Mesa Verde
San Juan
Rio Grande
Trinidad
Great Divide

Table 2.

Name and

DAU

E-1
E-2
E-3
E-4
E-5
E-6

Cold Springs
Bear's Ears
North Park
Poudre River
Strawberry
White River

E-7
E-8
E-9

E-14
E-15
E-16
E-17
E-18

Gore Pass
Troublesome
St. Vrain
Yellow Creek
Roan Bluffs
Piney
Williams Fork
Grand Mesa
Avalanche Creek
Frying Pan
Collegiate Range
Kenosha Pass

E-19

Glade Park

E-12
E-l3

(40)
(61, 62)
(53, 63, 64)
(54)
(55, 551)
(60)
(70, 71)
(65, 66, 67)
(68)
(69, 82, 84, 86)
(72, 73, 74)
(75, 77, 78, 751)
(76, 79, 80, 81)
(83, 85, 112, 851)
(3)

Elk data analysis units.

DAU

E-lO
E-ll

Game Management Units Included

Name and

Game Management Units Included
(201)
(4, 5, 14, 441)
(6, 7, 16, 161)
(8, 19)
(11)
(12, l3, 23, 24, 25, 26, 33,
34, l31)
(15, 27)
(17, 18)
(20,29,38,381)
(22)
(31, 32)
(35, 36)
(28, 37)
(41, 42, 52, 411, 521)
(43, 471)
(44, 45, 47)
(48, 56, 481)
(39, 46, 50, 51, 391, 461, 500,
501, 511)
(40)

�-7-

Table 2.

Elk data anlaysis

units.

(Continued).

and

DAD

DAU Name

E-20
E-21
E-22
E-23
E-24

Uncompahgre
Gunnison River
Buffalo Peaks
Eleven Mile
Disappointment
Creek
Lake Fork
Saguache
Sangre de Cristo
Grape Creek
Mesa Verde
Hermosa
San Juan
Rio Grande
Trinchera

E-25
E-26
E-27
E-28
E-29
E-30
E-31
E-32
E-33

Game Management

Unit Included

(61, 62)
(63, 64, 53, 54, 55)
(49, 57, 58)
(58, 59)
(70, 71)
(65, 66, 67)
(68)
(82, 86)
(69, 84)
(72 , 73)
(74)
(75, 77, 78, 751)
(76, 79, 80, 81)
(83, 85, 851)

Previous studies have established that domestic sheep ingest substantial amounts
of soil during their normal grazing activity.
(Healy and Ludwig 1965; Healy
1967). These early studies were designed to assess the effects of soil on
the wear of the incisor teeth, but since this time other implications of ingestion of soil in large amounts have been studied.
These ingested soils serve
as an important source of trace mineral in the diets of grazing animals.
Analysis of plant materials for trace element content is usually undertaken
to measure plant performance or to measure suitability to meet the animal
requirements for proper nutrition.
Where soil-plant-animal studies are
involved the collection of herbage samples free of soil contamination is
difficult.
This contamination takes the form of surface contamination, soil
adhering to roots, earthwQrm casts, etc., but all of these must be considered
important in the nutrition of the animal.
Ash content of feces from grazing animals gives a measure of soil content
and corrections can be made for the ash contribution of undigested herbage.
Treatment of ash with dilute acid gives an acid insoluble residue which may
serve as a measure of soil content.
Collection of feces in the field and
determination of acid insoluble residues are discussed by Healy and Ludwig
(1965). If feces output and soil content of the feces are known then the
quantity of soil ingested per day can be determined.
Feces output data are
usually available only from experimental animals under controlled conditions
and can vary with diet, age of animal, sex of animal, seasonal endocrine
rhythms, etc.
There are different factors which will influence soil ingestion by grazing
animals.
Soil type can be very important.
Well drained soils which have a

�-8-

strong structure are associated with low pasture contamination.
Soils
with a weak structure and poor drainage are associated with high contamination
(Healy 1968b). From the pattern of soil ingestion reported in Australia
(Arnold 1966) it is likely that physical properties of soils influence the
ingestion of soil by grazing animals.
The higher the stocking rate the higher
the amount of soil ingested
(Healy 1968; Arnold 1966).
Earthworm populations seem to correlate well with soil ingestion by grazing
animals.
Earthworms tend to deposit casts upon the soil surface and these
can be easily taken by animals grazing close to the grounds.
Seasonal
variations of soil ingestion where animals are grazing pastures the year
around show marked differences.
Ingestion is generally higher in winter months
when grass growth is poorest.
With new plant growth in spring and summer soil
ingestion tends to be at the lowest levels.
Seasons vary and a wetter than
normal spring or fall, or drier summer, may reflect differences in soil intake.
In cases where a specific trace mineral could be deficient as a result of
inadequate amounts of the element in forage, the deficiency could be made up
by soil ingestion.
As discussed previously the amount of soil ingested can
depend upon soil type, season, management practices, and even differences in
individual animals under similar conditions.
These factors may explain in
part the observations that at times only a percentage of animals in a specific
herd will show deficiency symptoms, and that only a percentage of farms on
a given soil type are affected.
Trace elements in soil are absorbed by the roots of plants, and trace element
concentration found in the herbage is a reflection of the soil from which it
comes.
In addition soil may have a direct effect on animal nutrition by
furnishing trace minerals directly.
As soil passes through the alimentary
tract it is subjected to the full range of influences involved in the digestive
process.
Conditions in the alimentary tract may increase the availability of
elements in the soil so that the element concentration in the digestive liquors
may be substantially increased.
Decreases in the concentration of an element
in the liquors may occur if the soil is capable of absorbing such an element
or removes it from solution by incorporation into an insoluble compound.
If changes in trace element concentration occurs in the alimentary tract, then
ingested soil would be expected to influence blood and tissue composition
and the general well-being of the grazing animals.
Various factors influencing soil ingestion have been briefly considered and
implications for animal health discussed.
It is usually considered that soils
influence animal nutrition by the quantity and quality of the herbage they
produce, so that the classical sequence is soil to plant to animal.
However
a direct soil to animal effect needs to be taken into consideration because
of the ingestion of soil.
During this segment a pilot study was designed and implemented in an effort to
develop laboratory procedures and test for differences in soil ingestion by
bighorn sheep from several different areas in the State. Bighorn fecal
samples were collected from the Lodore Canyon - Dinosaur National Monument;

�-9-

Jump Steady area west of Buena Vista; Love Ranch study area on Chalk
Creek; Rampart Range west of Colorado Springs.
To test for differences
between adult and lamb fecals the two groups were selected from Rocky
Mountain National Park. Collections were made in early summer when lamb
fecal pellets were easily distinguished from adult fecal pellets by size.
Pellet groups were collected from these study areas during June 1977 to
minimize seasonal variation.
A sample of ten pellet groups were selected
from each collection and all analytical procedures were made on these
fecal samples.
All fecal samples were air dried, ground in a Wiley Mill, and thoroughly
mixed prior to processing.
A two gram sample of fecal material was placed in a tared Gooch crucible
with a frittered glass disc bottom and ashed at 5000C over night.
The
crucibles were cooled, weighed and the percent ash was calculated as the
weight of the residue remaining in the crucible.
The acid soluble ash
portion of the residue was determined by adding O.IN HCI to the residue
and allowing the solution to drain from the crucible.
Several changes of
acid were added followed by distilled water rinse. The crucibles were then
dried to constant weight and the percent acid in soluble ash was computed.
Percent organic matter and percent acid soluble ash were computed by
difference.
A second 2 gm fecal sample was weighed into a 250 ml Erlenmeyer flask and
50 ml of 0.1 NHCI added. An additional 50 ml of O.IN HCl was placed
in a section of dialysis tubing and each end tied off. This tube was placed
in each flask so that the final concentration of the solids to fluid was
1:50. The flasks containing fecal material and O.lNHClwere
placed on a
rotary shaker and agitated for 24 hours. After this length of time the
dialysis tubing was removed and the contents transferred to a screw cap test
tube. All trace element determinations were made on this supernatant fluid.
A Perkin-Elmer 303 Atomic Absorption Spectrophotometer was used for the
calcium, magnesium, sodium, potassium, manganese, iron, copper and zinc.
Vanadate-molybdate
method was used for phosphorus determinations.
No statistical descriptions on comparisons of the data have yet been made.
Tabular summaries of the raw data are presented in Tables 3-28.

The

�-10-

Table 3. Calcium anlaysis
units in Colorado.

DAU

(ppm) of deer hair samples

from data analysis

N

Nean

Standard
Deviation

Range

D-2

Bear's Ears

20

1142

294

714-2036

D-4

Red Feather

39

1434

270

774-2088

D-6

Rangely

8

1257

161

1100-1530

D-7

White River

84

1729

471

916-4404

D-8

State Bridge

38

1588

296

1082-2200

D-9

Niddle

18

1591

432

967-2187

D-I0

Boulder

6

1331

274

991-1726

D-12

Grand Nesa

16

1815

519

896-2651

D-13

Maroon

3

1175

250

941-1439

D-14

Red Tables

9

1414

261

1136-1823

D-15

Cottonwood

4

1081

137

931-1250

D-16

Cripple

26

1553

725

771-3109

D-19

Uncompahgre

28

1043

155

748-l385

D-20

Crawford

23

1528

677

796-2717

D-21

West Elk

4

1248

420

876-1852

D-24

Ground Hog

9

2255

220

1951-2743

D-25

Powder Horn Cr

19

2349

386

1956-3100

D-26

Saguache

3

2285

196

2103-2493

D-27

Westcliffe

8

2307

305

2071-2864

D-30

San Juan

3

2292

499

1926-2861

D-33

Great Divide

12

1056

221

761-1559

Park

Bells

Creek

Creek

�-11-

Table 4. Magnesium analysis (ppm) of deer hair samples from data analysis
units in Colorado.

DAU

N

Mean

Standard
Deviation

Range

D-2

Bear's Ears

20

331

86

254-655

D-4

Red Feather

39

383

77

228-525

D-6

Rangely

8

391

50

339-490

D-7

White River

84

465

136

243-1132

D-8

State Bridge

38

420

66

302-557

D-9

Middle Park

18

412

101

244-579

D-I0

Boulder

6

323

37

258-366

D-12

Grand Mesa

16

378

76

219-550

D-13

Maroon Bells

3

322

75

238-381

D-14

Red Tables

9

325

55

267-443

D-15

Cottonwood Creek

4

293

45

263-360

D-16

Cripple Creek

26

477

227

180-808

D-19

Uncompahgre

28

318

68

182-474

D-20

Crawford

23

473

240

199-889

D-21

West Elk

4

246

37

204-278

D-24

Ground Hog

9

736

65

585-797

D-25

Powder Horn Cr

19

696

51

612-799

D-26

Saguache

3

711

29

678-729

D-27

Westcliffe

8

717

54

642-783

D-30

San Juan

3

714

49

682-771

D-33

Great Divide

12

282

73

144-419

�-12-

Table 5. Sodium analysis (ppm) of deer hair samples from data analysis
units in Colorado.

DAU

N

Mean

Standard
Deviation

Range

D-2

Bear's Ears

20

1187

473

221-2079

D-4

Red Feather

39

728

691

207-2423

D-6

Rangely

8

1916

604

592-2635

D-7

White River

84

457

331

102-1715

D-8

State Bridge

38

801

342

225-1465

D-9

Middle Park

18

653

316

243-1242

D-10

Boulder

6

2320

522

1612-2972

D-12

Grand Mesa

16

942

372

245-1940

D-13

Maroon Bells

3

1389

521

996-1191

D-14

Red Tables

9

1559

657

592-2595

D-15

Cottonwood Creek

4

1930

293

1575-2289

D-16

Cripple Creek

26

778

645

158-2426

D-19

Uncompahgre

28

572

350

181-1877

D-20

Crawford

23

965

836

126-2382

D-21

West Elk

4

1634

124

1466-1752

D-24

Ground Hog

9

195

58

99-285

D-25

Powder Horn er

19

269

108

l38-590

D-26

Saguache

3

213

44

162-241

D-27

Westcliffe

8

238

62

145-326

D-30

San Juan

3

346

III

218-418

D-33

Great Divide

12

971

311

512-1436

�-l3-

Table 6. Potassium analysis (ppm) of deer hair samples from data analysis
units in Colorado.

DAU

N

Mean

Standard
Deviation

Range

0-2

Bear's Ears

20

407

183

122-644

D-4

Red Feather

39

219

138

90-565

D-6

Rangely

8

5lO

144

197-650

0-7

White River

84

118

118

4-643

D-8

State Bridge

38

235

108

75-503

D-9

Middle Park

18

176

143

29-479

D-1O

Boulder

6

603

82

549-768

0-12

Grand Mesa

16

278

106

86-514

0-13

Maroon Bells

3

277

50

224-323

D-14

Red Tables

9

449

l31

254-625

0-15

Cottonwood Creek

4

550

lO6

413-648

D-16

Cripple Creek

26

216

157

31-598

D-19

Uncompahgre

28

225

174

60-981

0-20

Crawford

23

268

235

23-621

D-21

West Elk

4

390

87

309-484

0-24

Ground Hog

9

50.

18

28-76

0-25

Powder Horn Cr

19

60

43

24-204

D-26

Saguache

3

44

l3

29-53

D-27

Westcliffe

8

45

20

20-84

0-30

San Juan

3

69

31

35-96

D-33

Great Divide

12

333

98

192-507

�-16-

Table 9. Zinc analYSis
(ppm) of deer hair samples from data analysis
units in Colorado.

DAU

N

Mean

Standard
Deviation

Range

D-2

Bear's Ears

20

133

12

113-164

D-4

Red Feather

39

131

15

D-6

111-187

Rangely

8

142

8

D-7

132-154

White River

84

141

37

92-357

D-8

State Bridge

38

139

18

112-178

D-9

Middle Park

18

145

11

124-166

D-lO

Boulder

6

149

7

141-158

D-12

Grand Mesa

16

137

12

117-162

D-13

Maroon Bells

3

136

6

129-139

D-14

Red Tables

9

149

23

128-202

D-lS

Cottonwood Creek

4

128

10

117-142

D-16

Cripple Creek

26

120

14

90-156

D-19

Uncompahgre

28

121

13

94-150

D-20

Crawford

23

130

10

112-157

D-21

West Elk

4

132

18

118-158

D-24

Ground Hog

9

118

6

104-124

D-25

Powder Horn Cr

19

126

IS

104-155

D-26

Saguache

3

114

4

110-118

D-27

Westcliffe

8

127

17

110-161

D-30

San Juan

3

129

12

125-143

D~33

Great Divide

12

127

IS

103-161

�-17-

Table
units

10. Calcium
in Colorado.

analysis

DAD

(ppm) of elk hair samples

from data analysis

N

Mean

Standard
Deviation

Range

E-2

Bear's Ear

lO

1187

228

693-1566

E-6

White River

10

1105

160

821-1359

E-8

Troublesome

lO

1400

307

933-1603

lO

1048

216

681-1304

E...,13Williams

Fork

E-14

Grand Mesa

10

1296

417

882-2219

E-16

Frying Pan

10

1458

588

929-2290

E-21

Gunnison

10

1462

271

989-1859

E-25

Lake Fork

lO

1786

406

1302-2364

E-32

Rio Grande

lO

1593

281

1058-2066

River

Table 11. Magnesium
units in Colorado.

analysis

(ppm) of elk hair samples

from data analysis

DAD

N

Mean

Standard
Deviation

Range

E-2

Bear's Ear

lO

220

50

118-260

E-6

White River

10

241

28

200-292

E-8

Troublesome

lO

258

41

191-321

E-13

Williams

10

225

30

184-278

E-14

Grand Mesa

lO

246

43

185-308

E--16 Frying Pan

10

260

62

201-363

E-21

Gunnison

10

263

44

201-236

E-25

Lake Fork

lO

314

52

247-427

E-32

Rio Grande

10

275

34

215-323

Fork

River

�-18Ta0le 12. Sodium analysis (ppm) of elk hair samples from data analysis
li3its in Colorado.

DAU

N

Mean

Standard
Deviation

Range

E-2

Bear's Ear

10

114

35

54-168

E-6

White River

10

196

63

122-321

E-S

Troublesome

10

352

156

149-485

E-13

Williams Fork

10

495

258

153-823

E-14

Grand Mesa

10

393

227

103-667

E-16

Frying Pan

10

386

200

l34-659

E-21

Gunnison River

10

264

147

80-474

E-25

Lake Fork

10

252

164

83-484

E-32

Rio Grande

10

271

195

89-664

Table l3. Potassium analysis (ppm)
units in Colorado.

of elk hair samples from data analysis

DAU

N

Mean

Standard
Deviation

Range

E-2

Bear's Ear

10

195

51

108-279

E-6

White River

10

328

92

209-503

E-S

Troublesome

10

538

294

128-920

E-13

Williams Fork

10

765

368

241-1257

E-14

Grand Mesa

10

837

354

206-1266

E-16

Frying Pan

10

543

268

175-762

E-21

Gunnison River

10

394

275

76-809

E-25

Lake Fork

10

327

237

125-753

E-32

Rio Grande

10

331

219

93-685

�-19-

Table 14. Iron analysis (ppm) of elk hair samples from data analysis units
in Colorado.

DAU

N

Mean

Standard
Deviation

Range

E-2

Bear's Ear

10

33

18

11-81

E-6

White River

10

53

21

20-97

E-8

Troublesome

10

33

12

13-50

E-13

Williams Fork

10

42

19

17-65

E-14

Grand Mesa

10

52

20

31-99

E-16

Frying Pan

10

51

16

35-73

E-21

Gunnison River

10

49

22

20-87

E-25

Lake Fork

10

20

9

11-37

E-32

Rio Grande

10

25

12

15-46

Table 15. Copper analysis (ppm) of elk hair samples from data analysis
units in Colorado.

DAU

N

Mean

Standard
Deviation

Range

E-2

Bear's Ear

10

15

l3

5-47

E-6

White River

10

7

2

5-10

E-8

Troublesome

10

14

4

8-21

E-13

Williams Fork

10

l3

4

7-20

E-14

Grand Mesa

10

15

6

9-26

E-16

Frying Pan

10

16

7

8-26

E-21

Gunnison River

10

21

4

16-28

E-25

Lake Fork

10

21

5

10-29

E-32

Rio Grande

10

22

5

16-26

�-20-

Table 16. Zinc analysis
in Colorado.

(ppm)

of elk hair samples

from data analysis

DAD

N

Mean

Standard
Deviation

Range

E-2

Bear's Ear

10

75

27

56-146

E-6

White River

10

63

10

44-79

E-8

Troublesome

10

66

6

60-80

E-l3

Williams

10

63

3

56-67

E-14

Grand Mesa

10

63

7

52-77

E-16

Frying Pan

10

53

25

54-73

E-21

Gunnison

10

62

4

57-68

E-25

Lake Fork

10

60

5

53-68

E-32

Rio Grande

10

61

4

56-68

Table

17.

Fork

River

of bighorn

sheep fecal samples.

N

Mean

Standard
Deviation

Range

10

20.74

2.73

15.74-24.72

RMNP-(Adult)

10

44.54

16.86

18.54-63.73

RMNP-(Lamb)

10

43.61

21.38

12.56-66.96

Jump Steady

10

26.72

7.87

20.60-46.80

Love Ranch

10

18.58

1.37

16.81-19.71

Rampart

10

16.75

2.03

l3.52-19.76

Lodore

Table

Percent

Canyon

18.

ash content

of bighorn

N

Mean

Standard
Deviation

Range

10

11.96

2.22

7.26-15.75

RMNP-(Adult)

10

39.40

19.08

8.14-64.21

RMNP-(Lamb)

10

38.40

22.64

6.24-63.44

Jump Steady

10

20.90

9.03

12.01-44.08

Love Ranch

10

11.67

1.22

10.02-l3.48

Rampart

10

9.42

3.06

5.51-14.00

Lodore

Percent

total ash content

Canyon

acid insoluable

units

sheep fecal samples.

�-21-

Table 19.

Percent

acid soluble

ash content

of bighorn

sheep samples.

N

Mean

Standard
Deviation

Lodore Canyon

10

8.77

2.03

4.27-11.01

RMNP

(Adult)

10

5.15

2.60

2.50-10.40

RMNP

(Lamb)

10

5.21

2.02

3.22-9.44

Jump Steady

10

5.72

1.63

2.72-8.53

Love Ranch

10

6.92

0.79

5.34-7.56

Rampart

10

7.33

1.71

5.14-10.37

Table 20.

Calcium

(%) content of bighorn

Range

sheep fecal samples.

N

Mean

Standard
Deviation

Range

Lodore Canyon

10

2.19

0.52

1.25-2.75

RMNP

(Adult)

10

1.59

0.33

1. 38-1. 69

RMNP

(Lamb)

10

1.65

0.52

1. 23-2.69

Jump Steady

10

2.89

0.47

2.06-3.63

Love Ranch

10

1.68

0.12

1. 63-2.00

Rampart

10

2.78

0.52

1.81-3.63

Range

Table 21.

Magnesium

(%) content of bighorn

sheep fecal samples.

N

Mean

Standard
Deviation

Range

Canyon

10

0.41

0.12

0.14-0.53

RMNP

(Adult)

10

0.30

0.15

0.l3-0.65

RMNP

(Lamb)

10

0.24

0.08

0.14-0.39

Jump Steady

10

0.28

0.06

0.21-0.39

Love Ranch

10

0.21

0.02

0.18-0.24

Rampart

10

0.82

0.26

0.49-1. 20

Lodore

�-22-

Table 22.

Sodium

(%) content of bighorn sheep fecal samples.

N

Mean

Standard
Deviation

Range

Lodore Canyon

10

0.14

0.11

0.05-0.43

RMNP

(Adult)

10

0.26

0.l3

0.08-0.55

RMNP

(Lamb)

10

0.23

0.29

0.05-1.03

Jump Steady

10

0.08

0.02

0.05-0.13

Love Ranch

10

0.12

0.04

0.08-0.23

Rampart

10

0.18

0.05

0.10-0.25

Table 23.

Potassium

(%) content of bighorn sheep fecal samples.

N

Mean

Standard
Deviation

Range

Lodore Canyon

10

0.47

0.16

0.25-0.63

RMNP

(Adult)

10

0.76

0.25

0.38-1.13

RMNP

(Lamb)

10

0.51

0.36

0.25-1. 30

Jump Steady

10

0.l3

0.09

0.07-0.38

Love Ranch

10

0.61

0.15

0.50-0.93

Rampart

10

0.09

0.02

0.05-0.12

Table 24.

Manganese

(ppm) content of bighorn

sheep fecal samples.

N

Mean

Standard
Deviation

Lodore Canyon

10

176

62

80-263

RMNP

(Adult)

10

314

103

140-500

RMNP

(Lamb)

10

313

96

175-425

Jump Steady

10

575

167

375-875

Love Ranch

10

159

12

145-175

Rampart

10

358

90

145-500

Range

�-23-

Table 25.

Iron (ppm) content of bighorn sheep fecal samples.

N

Mean

Standard
Deviation

Range

Lodore Canyon

10

61

8

50-75

RMNP (Adult)

10

633

277

250-1025

RMNP (Lamb)

10

1124

823

250-2025

Jump Steady

10

575

167

375-875

Love Ranch

10

159

12

145-175

Rampart

10

358

90

145-500

Table 26.

Copper (ppm) content of bighorn sheep fecal samples.

N

Mean

Standard
Deviation

Lodore Canyon

10

9.1

4.0

7.5-17.5

RMNP (Adult)

10

11.8

4.3

5.0-15.0

RMNP (Lamb)

10

10.8

4.7

5.0-17.5

Jump Steady

10

52.0

26.8

12.5-92.5

Love Ranch

10

15.8

12.5-20.0

Rampart

10

7.8

3.5
1.8

Table 27.

Range

5.0-12.5

Zinc (ppm) content of bighorn sheep fecal samples.

N

Mean

Standard
Deviation

Range

Lodore Canyon

10

40.5

10.6

25.0-65.0

RMNP (Adult)

10

60.8

22.8

37.5-112.5

RMNP (Lamb)

10

55.0

28.1

32.5-110.0

Jump Steady

10

61.5

46.6

15.0-175.0

Love Ranch

10

53.8

10.6

45.0-72.5

Rampart

10

77 .0

17.2

60.0-115.0

�-24-

Table 28.

Phosph~rus

(%) content of bighorn sheep fecal samples.

N

Mean

Standard
Deviation

Range

Lodore Canyon

10

0.56

0.15

0.36-0.90

RMNP (Adult)

10

0.99

0.43

0.53-1.80

RMNP

(Lamb)

10

1.45

0.83

0.75-2.65

Jump Steady

10

0.64

1.25-3.70

Love Ranch

10

2.50
1.25

0.50

0.80-2.15

Rampart

10

0.88

0.15

0.58-1. 08

LITERATURE

CITED

Anke, M. 1965. Mineral and trace element content of cattle hair as
indicators of Ca, Mg, P, K, Na, Fe, Zn, Mn, Cu, Mo, and Co nutrition.
II. Relationships to cutting depth, hair type, hair color, hair age,
animal age, lactation-stage, and pregnancy.
Arch. Tierernahr.
15(6) :469-485. (BioI. Abs , 47:92059).
Anke, M. 1967. Mineral and trace element content of cattle hair as an
indicator of Ca, Mg, P, K, Na, Fe, Zn, Mn, Cu, Mo, and Co nutrition.
V. The mineral supply of dairy cows on soils of various geographical
origin measured by the mineral content black cattle cover hair and of
red clover.
Arch. Tierernahr. 17(1-2):1-26.
BioI. Abs. 48(1-2179).
Arnold, G. W., W. R. McManus, I. G. Bush. 1966. Studies in the wool
production of sheep.
5. Observations on the teeth wear and carryover
effects.
Aust. J. Exp. Agric. and Anim. Husb. 6(20):101-107.
Flynn, A., A. W. Franzman, P. D. Arnson, O. A. Hill.
(1974). Determination
of Past Trace Element Uptake in a Wild Animal by Longitudinal Analysis
of the Hair Shaft. Naturwissenschaften
61(8):362.
Hansen, H. C., R. L. Jones.
Ross' Geese.
Southern

(1976). The Biochemistry of Blue, Snow, and
Illinois University Press.

Healy, W. B., T. G. Ludwig.
(1965). Wear of sheep's teeth.
of ingested soil. N. Z. J. Agric. Res. 8(4):737-752.
Healy, W. B. (1968). Ingestion
Res. 11(2):487-499.

of soil by dairy cows.

I.

The role

N. Z. J. Agric.

Werner, A., M. Anke.
1960. Trace Element Content of Beef Cattle Hair as
an aid in recognizing deficiency phenomena.
Arch. Tierernahr.
(10):142-153. (Chem. Abs. 54:16574).

Prepared

by

--,ZlJ..~~~~&amp;~'!.~~r-~~a.~~.~;
••.•.•........
Robert E. Keiss
Wildlife

Researcher

�January,

1:J19

-25JOB FINP~ REPORT

State of

COLORADO
----~~~~~-------W-41-R-28

Project No.
Work Plan No.
Job Title

Bighorn Sheep and Mountain

Goat Investigations

23
Job No.
Bighorn Sheep Lungworm Control
By Biological Control of Snail Intermediate
1

Hosts
--~~----~----------------------------------------

Period Covered: June 1, 1977 - May 31, 1978
Personnel:

F. E. Latson, and T. N. Woodard

ABSTRACT
Land snail literature pertinent to the bighorn sheep (Ovis canadensis) lungworm (Protostrongylus sp.) complex was compiled.
A literature review
on land snails was begun and the completed portions are presented in this
report. Literature relating to snail systematics, methods of collecting,
examining, and preserving snails, and research needs will be synthesized
and incorporated into a final report during the next segment. The report
will be published in either Division of Wildlife Special Report format
or the Division Report format with Project W-126-R, Work Plan 1, Job 7 funds.

��-27-

BIGHORN SHEEP LUNGWORM CONTROL
BY BIOLOGICAL CONTROL OF SNAIL INTERMEDIATE

HOSTS

Ed Latson and Thomas N. Woodard

P. N. OBJECTIVE
To test the effects of burning treatments as a tool to reduce populations
of the snail intermediate hosts for bighorn sheep lungworm parasites.
SEGMENT OBJECTIVES

1.

Complete

compilation

of land snail literature.

2.

Prepare

a literature

review on land snail ecology.

3.

Publish results

in the Division

of Wildlife

Special Report

format.

RESULTS AND DISCUSSION
All segment objectives were not completed during this segment.
Completed
portions are incorporated in this report.
The final literature review will
be published during the next segment under the auspices of W-126-R, Work
Plan 1, Job 7.
The original Program Narrative Objective (above) was never met because
preliminary sampling indicated it would be exhorbitantly costly to obtain
samples of sufficient size to test burning effects on snail populations.
But we decided to publish the literature review portion of this job because
of its contribution to knowledge of the bighorn sheep lungworm life cycle.

Intermediate

A Literature Review of Land Snails;
Hosts in the Bighorn Sheep Lungworm Life Cycle
INTRODUCTION

The bighorn sheep, Ovis canadensis
is commonly infected with Protostrongylus
species lungworms.
These lungworms have been shown to be a factor in the
etiology of disease in bighorn sheep, particularly in lambs.
Transplacental
transmission of this parasite is important in the production of disease in
the lambs, but the life cycle of the parasite requires an intermediate host
which can be any of several small terrestrial molluscs.
Understanding this
disease as a prelude to control requires an understanding of the intermediate
hosts of this parasite.
This report is a compilation of some of what is
known about these snails.

�-28-

LUNGWORM

AND SNAIL BIOLOGY

Protostrongylus

Life Cycle
A discussion of the life cycle of the parasite will facilitate
standing the discussion of the intermediate host snails.

under-

Specific and detailed information on the complete life cycles of Protostrongylus
spp. found in bighorn sheep is not known; however, the basic features for
each cycle are apparently similar. Pillmore (1958) noticed that adults are
dioecious and occur in the lungs. The first stage larva (Ll) emerges from an
egg produced by the female in the lungs and moves up respiratory passages to
the pharynx where it is swallowed and passes through the digestive tract.
Ll larvae are 240-330 microns long.
For further deveLopment the larva must penetrate the foot of a suitable snail.
In the snail the larva grows to 550-700 microns, and the gut becomes distended
with food granules.
The second stage (L2) is reached when the larva casts the
Ll cuticle and shrinks within it. The L2 cuticle soon develops heavy transverse
striations especially along the dorsal surface.
After a time the second molt
occurs resulting in the larvae being ensheathed in two shed cuticles.
The
outer cuticle is thin and hyaline, the inner is thick and striated.
The infective stage is reached after the second molt when the inner ensheathing
cuticles are lost and granulation in the larval gut decreases.
Infection of the sheep results when infective larvae are ingested by the host
and move via an unknown route from the gut to the lungs, where development is
completed.
Larval Production
Numbers of larvae in the feces of bighorn sheep vary with the level of infection
of the host, the season of the year, and other poorly understood factors.
Pillmore (1959) found that larval output varies from sample to sample and pellet
to pellet in individual sheep. He found a range of 30 to 815 larvae per fecal
pellet in 11 samples collected on a single day from two penned ewes. He
(Pillmore 1955) and others report a peak larval discharge in late winter and
spring months.
Larval Ecology
The first stage larvae can withstand dessication, thermal extremes, and are
capable of survival over prolonged periods of time. Pillmore (1959) kept
larvae frozen in feces for over a year without significant loss of viability.
Honess (1942) kept feces containing larvae for nearly eight months in an
unheated outbuilding with little loss of viable larvae.
Forrester and Senger
(1963) noted that larvae in feces can survive high temperature dessication
while free larvae survive better in a moist situation.

�-29-

Pillmore (1959) stated that Protostrongylus
larvae can pass through the
intestine of a sheep and be excreted in the feces a second time, and may
also pass through the intestinal tract of carnivores without harm.
Infection

of the Snail

No information is available on the route of penetration into the snail for
P. stilesi, but the entry of Muellerius capillaris has been described.
Kassai (1958) described entrance as a direct penetration of the epithelium
of the snail. Hobmaier and Hobmaier (1934) felt entrance to be through
mucous glands.
All three investigators agreed that the larvae becomes highly
activated by the presence of the snail and actively penetrates by whatever
route.
Size, age, and species of snail have been found to affect the ability of
larvae to penetrate the snail. Kassai (1958) mentioned Joyeux and Gaud's
belief that Muellerius larvae only infect small species of snails with "loose
mesoderm", or larger species of snails only when they are young.
However,
Kassai did not think size or age was important and cited his own infection
of Helix pomatia, the French edible snail, with 1,000 to 1,500 larvae.
He
did not specify genus and species of larvae used. He mentioned that all
snails have a one-layer clyindrical epithelium.
Latson (1977) found one and
two-whorl Pupilla muscorum to contain a significant number of larvae in
collections made in April.
An adult P. muscorum has over 4 whorls, a newborn P. muscorum has slightly less than one whorl.
A single one-whorl and
a single two-whorl snail were observed to contain a single mature larva
each.
Gerichter (1948) found that several lungworm species could infect nearly
all the snail species he studied; however Protostrongylus kochi (now called
P. rufescens) developed in only a few species.
Pillmore (19~8) listed the following species of snails as susceptible to
invasion by protostrongylid lungworms in Colorado:
Pupilla muscorum, P.
blandi, Vallonia cyclophorella,
V. pulchella, Gastrocopta armifera, G.
pentodon, Pupoides albilaris and Vertigo concinula.
Forrester

(1962) attempted

Deroceras,

Euconulus,

unsuccessfully

Oreohelix,

Retinella,

to infect species of Allogona,
Triodopsis, Zacoleus and zonitoides.

Latson

(1977) experimentally infected one Gastrocopta sp. and three Discus
He also found 2 naturally infected Euconulus fulvus.
The genera
now known to support infection include:
Pupilla, Vallonia, Vertigo, Gastrocopta
Discus, Pupoides, and Euconulus.
cronphitei.

Rate of development and intensity of infection are definitely related to
species of snails for most, if not all, lungworms (Gerichter 1948, 1951;
Kassai 1958; Rose 1973). Rate of development of individual larvae in the same
snail also can be variable (Kassai 1958; Gerichter 1948). Gerichter noted
that in concurrent infections with more than one species of lungworm, each

�-30-

developed at its normal rate. A heavier infection causes, in some cases,
slower development (Gerichter 1948; Pillmore 1955). Pillmore (1956) observed
that development rates for P. stilesi, in Pupilla, Vallonia and Vertigo
varied greatly, the second occurring 11 to 60 days post-infection.
Monson
(1971) observed that the second molt for P. stilesi occurred 14 days postinfection in V. pulchella.
These experiments were not carried out under
controlled conditions.
Humidity, temperature, health of the snails or
other factors could have influenced the development rate.
Latson (1977) found infective larvae in snails from April to October.
Collections were not made between October and April due to cold weather.
The highest percentage of immature larvae were seen in late April and late
July, presumably indicating recent infection.
These times correspond to
time of increased moisture.
In late June a small percentage were immature
indicating most of the immature larvae seen in April had matured by June.
No immature larvae were seen in September.
June and September were both
dry periods.
The precise time at which the larva will be infective if ingested by the
sheep is not known. Pillmore (1955) felt that some development or maturation
past the second molt would be required before the larva would become infective.
Gerichter (1948, 1951) described a pre-infective stage following the second
molt for Muellerius capillaris, but found no morphological evidence for a
similar situation with P. rufescens.
Rate of development in the snail is directly dependent on temperature.
Development ceases at temperatures slightly above freezing.
Larvae of P.
refescens underwent no development below 80C in one experiment.
Gerichter
(1951) infected snails of the species Helicella barbesiana, H. vestalis
jopponensis and Monacha syriaca with P. rufescens larvae and then cooled the
snails from room temperature to 40 -80C. After 50 days without development
the snails were returned to room temperature and the larvae developed normally.
Gerichter (1948) observed that activity of the snail and its nutritional state
did not affect rate of larval development for larvae of several species of
lungworm in five species of snails. Kassai (1958) found the same with one
exception:
snails starved for six months before infection would not support
development of larvae of a Cystocaulus sp.
The snail does not appear to react markedly to infection.
Hobmaier and Hobmaier (1930) mentioned a tubercle-like proliferation in response to P. refescens,
but Kassai (1958) noted only pigment formation and saw no proliferative reaction
with larvae of a Cystocaulus sp. in several species of snail. He also found
that previously infected snails were readily reinfected.
Latson (1977) observed
several P. muscorum with pigment concentrations of the same shape and size as
a mature ensheathed larva.
The presence of larvae in the snail seems to have little effect on the viability
of the snail. Kassai (1958) experimentally infected Helix pomatia, a large
snail, with 1,000 to 1,500 larvae without any observed ill effects on the snail.

�-31-

Gerichter (1948) saw infections of 200 larvae per snail without any observed
ill effects on the snail. Lange (Personal Communication) observed over 60
Protostrongylus
larvae in a single Pupilla.
Latson (1977) has seen 48
larvae in a single V. pulchella; the snail appeared to be normal.
Kadenatsh (1969) found that Protostrongylus
tauricus of European hares
leaves the snail after developing to the infective stage.
The larvae that
left the snails were found coiled and covered by dried mucus on plants and
container walls.
Hares were experimentally exposed to these larvae and
successfully infected.
Pillmore (1957, 1961) Lange (Personal Communication)
and Latson (1977) have seen snails containing only the larval sheaths of
Protostrongylus
stilesi, suggesting that a similar situation might occur
with the species in bighorn sheep.
Infection

of the Sheep

Ingestion of the snail (or a free larva, if such do exist naturally) by a
sheep initiates the final phase of the cycle. The infective larva penetrates
the gut and by an unknown route reaches the lungs where it completes development to the adult stage.
Ingestion of the snail probably occurs accidentally
while the sheep is consuming plants among which snails are found (Pillmore
1955). Ingested larvae can also go to a fetus transplacental1y, either directly
or possibly after a period of dormancy within maternal tissues (Hibler et al.
1972, 1974).
Snails
The snails thus far implicated as intermediate
snails with similar habitats and life cycles.
factors in their ecology are discussed below.

hosts are all small terrestrial
Some of the more important

Life Span
Boycott (1934) said that in the terrestrial snails of Britain. including all
the genera and some of the same species found in the study area, all but the
larger snails have a normal life span of 9 to 15 months.
He believed the
greatest mortality occurred among eggs and infant young.
Walton (1970) believed
the life spans of snails of the genera Ashmunella, Sonorella and Monodenia
might be quite long, up to 16 years in Ashmunella.
These snails are large
snails, up to 2.5 cm in diameter.
Life Cycle
Very little is known about life cycles of snails suitable as intermediate
hosts.
All are hermaphroditic, and many are capable of self-fertilization.
Most of them are oviparous but Pupilla is ovoviviparous.
Whitney (1938)
studying V. pulchella in Indiana, found that egg laying occurred at 24-hour
intervals under favorable conditions.
For ten individual snails the average

�-32-

number of eggs per snail in three months was 53. Hatching occurred at
about 12 days and average time from hatching to maturity was 59 days.
She raised four successive generations from isolated snails to demonstrate
self-fertilization.
produces an immature snail enclosed in a membranous sac. The adult
carries several embryonic snails in the uterus, releasing them as they mature.
Latson (1977) found Pupilla muscorum to release embryos in response to
moisture following periods of drying. He found the average number of embryos
per adult P. muscorums collected from a wild population to vary from 1.7 in
the dry months of June to .37 in July following a period of daily rain showers.
In laboratory cultures he found a cycle of 3 or 4 days of drying followed
by watering of the culture dishes was required to obtain reproduction.
Most
of the young snails were produced during the first day after moistening.
Young snails were enclosed in a membrane when released from the adult, but
penetrated it within a few hours. The young snails were highly susceptible
to drying or exclusive moisture.
Pupilla

Behavior
The primary determinants of behavior seem to be temperature and humidity.
It is commonly stated that terrestrial mulluscs are nocturnal due to cooler
temperatures and higher humidity.
Increased humidity is thought by many
authors to be most important, but Dainton (1954) found decreasing temperature
to be the most important stimulus for the grey field slug (Agriolimax reticulatus).
Responses to changes of O.loe per hour were observed with decreasing
temperature stimulating activity.
The response was reversed at temperatures
between 200 and 300e when rising temperature stimulated activity.
Such reactions
would serve to prevent activity under desiccating conditions and could direct
movement toward locations with optimum conditions.
Pillmore (1959) observed active snails twice, once in September following a
light snow which was melting, and once in May on a northwest slope near melting
snow. Latson (1977) found active snails only once. This was of the 15th of
July at 9:00 a.m. The elevation was 3,780 m. The area was wet from recent
rains and the air was humid. The area had a south-southwest exposure.
Moisture

and Snails

Moisture is one of the most important factors in a snail's environment. The
eipthelium of a snail gives no protection against desiccation and movement
necessitates secretion of a slime trail containing considerable moisture.
When
moisture is available in droplet form it can be absorbed by either oral or
cutaneous absorption (Likhachev 1962). Hyman (1967) stated Helix pomatia can
survive water loss of 50% of body weight for 10 to 11 months.
Wells (1944)
observed large fluctuations in weight of H. pomatia maintained under constant
conditions and found estivation was associated with decreased body weight even
under conditions of high humidity.
Boycott (1934) stated that most land snails
and slugs are easily drowned.
Pillmore (1957) found no snails in areas occupied
by snow banks that lingered into summer. Latson (1977) did not find snails

�-33-

in areas where moisture tended to remain such as depressions
in high elevation collection areas.

or flat areas

Snails very widely in their resistance to extremes of moisture.
Vitrina
with its thin shell and large shell opening, can survive only a
short time when removed from its humid environment among fallen leaves if
it is active when removed.
Conversely, Sphincterochila boissieri, in the
Negev Desert of the eastern Mediterranean, is found scattered over the rocky
surface in direct sun (Machin 1967).

alaskana,

Likhachev (1962) stated snails found in areas of low humidity tend to be
small and have thick shells. Machin (1967) found a correlation between dryness of habitat, thickness of shell, smallness of shell opening and thickness
of epiphragm.
Of the three species of Helicidae he examined, S. boissieri
lives in the driest environment and is smaller, has a thicker shell, a thicker
epiphragm
and a smaller aperture.
Estivation

and Hibernation

Snails survive periods of extreme conditions by estivation and hibernation.
Hibernation usually occurs in winter, the process beginning by sealing the
shell with an epiphragm of hardened mucus and calcareous material.
Estivation
is a similar process beginning with secretion of a thinner film of dried mucus.
It usually occurs in response to desiccation.
Both conditions are associated
with greatly slowed metabolic processes (Hunter 1964). Water loss is reduced
in these states thus enabling survival for long periods, even as long as six
years (Comfort 1964).
Minerals

and EH

These two parameters are closely related because solubility of various compounds
is dependent on pH. Karlin (1961) indicated snails able to utilize insoluble
calcium could survive in areas of acid soil. Burch (1955) studied ecological
factors of soil affecting distribution of snails in eastern Virginia and concluded
the primary conditions of soil which limit distribution of land snails are
calcium, magnesium and organic matter.
Potassium, phosphorus and pH were assumed
not to be limiting factors.
Snails living on calcareous substrates have heavier shells indicating their
shells are thicker (Hyman 1967). Boycott (1934) lists 45 of 108 snail species
as indifferent to a calcareous substrate, and one species as not found in
calcareous areas. Hyman (1967) mentioned that Carrick (1942) found Agriolimax
agrestis tolerant of a wide range of pH.
Nutrition
Hyman (1967) stated terrestrial pulmonates eat what is available and are not
very selective.
Snails examined for various digestive enzymes have been found
to have a wide complement of carbohydrases, including cellulases which some

�-34-

feel to be of bacterial

origin (Owen 1966). Likhachev (1962) mentioned
and Euconulus feed on fungi and decomposing plant
remains, and molluscs do not use conifers as food, possibly because of the
hardness and pitch content of the needles.
Walton (1963) observed Oreohelix
refused lettuce and died after a few weeks.

Pupillidae,

Vallonidae

Temperature
The effect of temperature extremes on snails found in the Pike's Peak area
is not clear.
Pillmore (1959) attributed a decrease in a population of snails
to unseasonably cold weather in November of 1957. Latson (1977) found dead
snails on 20-21 October with the shells containing dried tissue possibly
indicating reyent death.
These snails must be able to withstand freezing to
survive winter.
Possibly a period of acclimitization is required before they
can withstand freezing.
Whitney (1938) found recovery of v. pulchella after refrigeration to be nearly
independent of duration of refrigeration.
Likhachev (1962) described Vertigo
and Euconulus as thermophobic.
Monson (1971) cultured infected V. pulchella
in an incubator at 250C.
Vegetation
Vegetation provides food and shelter for snails.
By providing a suitable
microhabitat vegetation can directly influence presence or absence of snails.
Karlin (1961) discussed the value of reporting vegetation when collecting
mollusks.
He found important correlations between vegetation and mollusks
including an almost complete absence of mollusks in coniferous forests.
Distribution

and Habitat

Mircohabitat is the most important factor in distribution of snails.
Often
in the most inhospitable area a small bush or rock may provide suitable habitat.
Likhachev (1962) noted mountains have a more varied malacofauna due to more
mircohabitats.
Getz (1974) found a positive correlation between species
diversity of snails both with increased moisture and with diversity of tree
species.
Boycott (1934) said as habitat improve, species from poorer habitats
are still present and new ones are added. Kerney (1968) and Evans (1968)
described Pupilla muscorumand Vallonia spp. as open ground species in Britain.
Pillmore
(1955) found snails in alpine areas on Pike's Peak frequently were
associated with alpine avens as well as alpine sedges and grasses.
Aspen groves,
moist meadow edges, and under sites of logs also were productive.
On Buffalo
Peaks he found a similar situation, but often a higher concentration of snails
in alpine were associated with alpine clover. A single clump of alpine clover
yielded 183 shells.
Blood (1963), in a study of a California bighorn range,
found Vallonia cyclophorellaandPupilla
sp. to be abundant on wet grass and
under pieces of wood and debris around forested margins of bunch grass slopes
at about 1,550 m (5,100 ft) elevation.

�-35-

Latson (1977) found snails to occupy limited habitat types, depending upon
species.
P. muscorum was more widely distributed than other species.
This
species could survive high elevations and dryness, but much of the study
areas was unsuitable because of excessive elevation, lack of protection from
cold and drying westerly winds, excessive moisture, lack of suitable vegetation, or combinations of these and other factors.
P. muscorum was most
numerous on a south to southeast slope, below 3,800 m, among Alpine Avens
(geum rossii) Kobresia sp. or beside rocks.
Usually there was some protection
from west wind, but snails were not found in areas with lingering snow drifts.
The soil was well drained.

Pillmore's (1955) descriptions of snails collected on Buffalo Peaks and Pike's
Peak sheep ranges show P. muscorum to be the most common snail in alpine areas.
At lower elevations P. blandi was found most commonly.
Bequaert and Miller (1973) in their discussion of P. blandi and P. muscorum
indicate P. muscorum is found to 620 N latitude and is found only at high
elevations (2,000 to 3,700 m) in New Mexico.
P. blandi is found only 520 N
latitude and is found at lower elevations (1,500 to 2,800 m) in New Mexico
and Arizona.
blandi was the most commonly infected snail found by Pillmore (1955)
on Pike's Peak. Forty-five of 92 were infected in his Area 6, the only place
he found infected snails on Pike's Peak. He also found 3 of 20 Vallonia
cyclophorella
and I of 7 Vertigo concinnulla infected.
Hunter and Pillmore
(1954) reported finding infected snails on a west facing slope below a rocky
outcropping on which sheep sign was seen. Snails were collected about the
bases of Deschampsia caespitosa (tufted hair grass).
This location is at a
lower elevation than most of Latson's (1977) coilections.

Pupilla

Latson (1977) found Vertigo spp. to be most common on moderate north slopes
beside rocks and under alpine avens or in meadow edges and aspen groves.
These
habitats are mose moist than optimum for P. muscorum.
Vertigo was the most
common snail found on north slopes and was the only snail found infected on a
north slope.
It is probably the species most likely to occur and be infected
on northern exposures.
Its usual location adjacent to rocks beneath the soil
surface would limit chances of its being ingested.
Latson (1977) reported Euconulus fulvus, the other species he found infected,
to occur in meadow edges and aspen groves and on southern exposures with more
protection and moisture than those in which Pupilla muscorum were not numerous.
These represent relatively moist and protected environments.
Dispersal

of Snails

Efficient passive dispersal is one advantage possessed by a small, hermaphroditic
organism able to remain inactive for long periods while attached to pieces of
substrate such as leaves or bits of grass. Wind and water can easily carry these
for long distances.
Karlin (1969) raised several questions
concerning how
isolated stands of aspen he examined were invariably populated by a snail

�-36fauna; how similar species were very widely distributed in a similar habitat;
and how burned areas were repopulated.
He suggested the possibility of small
populations in evergreen forests too sparse to be readily found, but which
could repopulate aspen forests growing up after a fire. Perhaps a better
explanation is that this is an example of efficient dispersal.

LITERATURE

CITED

Allen. R. W. 1962. Parasitism in bighorn sheep on the desert game range
in Nevada.
Trans. Desert Bighorn Sheep Council 6:69.
Allen, R. W. 1964. Additional notes on parasites of bighorn sheep on the
desert game range, Nevada.
Trans. Desert Bighorn Sheep Council 8:5.
Baker, H. B.

1955.

Land snail dispersal.

Nautilus

71:141-148.

Bensink, A. H. A. Observations on the biology and population
land snails in a Quebec apple orchard.
Thesis.
490pp.
Library of Canada, Canadian theses on microfilm #4575.

dynamics
National

of

Bequaert, J. C. and W. B. Miller.
1973. The Mollusks
The University of Arizona Press, Tucson, Arizona.

of the Arid Southwest.

Blood, D. A. 1963. Parasites from California bighorn
British Columbia.
Can. J. Zool. 41:913-918.

sheep in southern

Boycott, A. E.
of Ecology

1934. The habitats
22:1-38.

of land mollusca

in Britain.

Jour.

Buechner, H. K. 1960. The bighorn sheep in the United States, its past,
present and future. Wildlife Monograph 4.
Burch, J. B. 1955. Some ecological factors of the soil affecting the
the distribution and abundance of land snails of eastern Virginia.
Nautilus 69:62-69.
Burch, J. B. 1956. Distribution of land snails in plant associations
eastern Virginia.
Nautilus 70:60-64, 102-105.
Burch, J. B.
Dubuque,

1962.
Iowa.

How to Know the Eastern Land Snails.

in

Wm. C. Brown Co.,

I

Chamberlain. R. V. and D. T. Jones.
1929. A descriptive catalog of the
mollusca of Utah.
Bull. of the Univ. of Utah 19(4):Bio. Series 1(1).
Chitwood, B. G. and M. B. Chitwood.
1937.
of nematomorphs.
Nautilus 50:130-135.
Comfort, A. 1957. The duration
of London 32(6):219-241.

Snails as hosts and carriers

of life in molluscs.

Croll, N. A. 1972. Behavior of larval nematodes.
Wright (eds.) Behavioral Aspects of Parasite

Proc. Malac. Soc.

In E. U. Canning and C. A.
Transmission.
Academic Press.

�-37-

Dainton, Barbara H. 1954. The activity of slugs.
I. Induction of
activity by changing temperature.
Journ. of Experimental Biology
31:165-187.
Dikmans, G. 1931. Two new lungworms from North American ruminants and
a note on the lungworms of sheep in the United States.
Proc. U.S.
Nat'l. Mus. 79:article 18.
Dikmans, G. 1937. Protostrongylus rushi, a new lungworm from the mountain
sheep, Ovis canadensis.
Sb. Rabat po Gel'mintol.
Skrjabin., PI 126.
Dikmans, G. 1943. The lungworm, Protostrongylus rushi Dikmans, 1937, of
the mountain sheep Ovis canadensis.
Proc. Helminthol. Soc. Wash.
D. C. 10:8.
Dikmans,

G.

1957.

frosti Honess,

A note on the specific identity of Protostrongylus
1942. Proc. Helminthol. Soc. Wash. D.C. 24;116.

Dougherty, E. C. and F. C. Goble.
1946. The genus Protostrongylus
Kamenskii, 1905 (Nematoda: Metastrongylidae)
and its relatlves.
Preliminary Note. Journ. of Parasit. 32:7-16.
Evans, J. G. 1968. Changes in composition of land molluscan populations
in North Wiltshire during the last 5000 years.
Symp. Zool. Soc. of
London. 22:293-317.
Forrester,

D. J.

1962.

Land mollusca as possible intermediate hosts of
a lungworm of bighorn sheep in western
Proc. Mont. Acad. of Sci. 22:82-89.

Protostrongylus

Montana.

stilesi,

Forrester, D. J. 1971. Bighorn sheep lungworm pneumonia complex.
pp.158173 in J. W. Davis and R. C. Anderson (eds.) Parasitic Diseases of
Wild Mammals.
Iowa State University Press, Ames, Iowa.
Forrester, D. J. and C. M. Senger.
1963. Effects of temperature and
humidity on survival of first-stage Protostrongylus stilesi larvae.
Expt. Parasit. 13:83-89.
Forrester, D. J. and R. C. Littell.
Lungworm Infections in bighorn

1976.
sheep.

Influence of Rainfall on
J. Wildl. Diseases 12:48-51.

Gerichter, Ch. B. 1948. Observations on the life history of lung
nematodes using snails as intermediate hosts.
Am. J. Vet. Res.
9(30):109-112.
Gerichter, Ch. B. 1951. Studies on the lung nematodes
in the Levant.
Parasitology 41:166-183.
Getz, L. L. 1974. Species diversity of terrestrial
Smokey Mountains.
Nautilus 88:6-9.

of sheep and goats

snails in the Great

�-38-

Gleich, J. G., F. F. Gilbert, and N. P. Kutscha.
Terrestrial Gastropods from Central Maine.
Henderson, J.
Wyoming.

1924. Mollusca of Colorado,
Univ. of Colo. Stud. 13(2).

1977. Nematodes in
J. Wildl. Diseases 13:43-46.

Utah, Montana,

Idaho and

Hibler, C. P., R. E. Lange and C. J. Metzger.
1972. Transplacental
transmission of Protostrongylus
stilesi in bighorn sheep.
J. Wildl.
Dis. Assoc. 8:389.
Hibler, C. P., C. J. Metzger, T. R. Spraker and R. E. Lange.
1974. Further
observations on transplacental transmission of Protostrongylus stilesi
in bighorn sheep. J. Wildl. Dis. Assoc. 10:39-41.
Hitchcock, A. S. 1971.
(2nd ed.). Manual of the Grasses of the United
States.
Dover Publications, New York.
Hobmaier, A. and M. Hobmaier.
1930. Life history of Protostrongylus
(Synthetecaulus) rufescens.
Proc. Soc. Exptl. BioI. and Med.
28:156-158.
Hobmaier, A. and M. Hobmaier.
localization of lungworms
Honess, R. F.
Wyoming.

1934. Route of infection and the site of
in molluscs.
Science 80:229.

1942. Lungworms of domestic sheep and bighorn
Univ. of Wyo. Agr. Expt. Sta. Bull. 255:1-24.

sheep in

Howe, D. L. 1964. Life cycle of lungworms.
No. 11, Job No. lW.

Wyo. FW-3-R-ll.

Work Plan

Howe, D. L. 1965. Life cycle of lungworms.
No. 11, Job No. 11, lW.

Wyo. FW-3-R-12.

Work Plan

Hunter, G. N. and Pillmore, R. E. 1954. Hunting as a technique in
studying lungworm infestations in bighorn sheep. Trans. North Am.
Wildlife Conf. 19:117.
Hyman, L. H.
Inc.

1967.

The Invertebrates.

Volume 6, Mollusca

I.

McGraw-Hill,

Kadenatsh,

A. M. 1969. Biological investigation of Protostrongylus
pulmonary parasite of hares.
In S. N. Boev (ed.)
Contributions to Helminthology.
Translated from Russian by the
Israel Program for Scientific Translations.
Jerusalem.
350pp.
tauricus,

Karlin, E. J. 1961. Ecological relationship between vegetation and the
distribution of land snails in ~ontana, Colorado, and New Mexico.
Am. Midland Natr. 65:60-66.
Kassai, T. 1958.
8:223-236.

Larvae of Protostrongylins

in snails.

Acta. Vet.

�-39-

Kerney, M. D. 1968. Britain's fauna of land mollusca and its relation
to the Post Glacial Thermal Optimum.
Symp. Zool. Soc. of London
22:273-291.
Latson, F. E. 1977. The distribution and ecology of intermediate host
snails of Protostrongylus
spp. lungworms of bighorn sheep on Pikes
Peak, Colorado.
M.S. Thesis, Colorado State University, Fort Collins.
64pp.
Likhachev, I. M. and E. S. Rammel'meier.
1952. Terrestrial Mollusks of
the Fauna of the U.S.S.R.
Translated from Russian and published
for the National Science Foundation, Washington, D.C. by the Israel
Program for Scientific translations.
Jerusalem.
Machin, J. L. 1967. Structural adaptations for reducing water loss in
3 species of terrestrial snails.
Jour. Zool. 152:55-65.
MacMillan, G. K.
59:121-124.

1946.

Notes on some southwestern

Pupillidae.

Nautilus

Marr, J. W. 1961. Ecosystems of the East Slope of the Front Range of
Colorado.
Boulder, Univ. of Colo. Press.
Mason, C. F. Snail population, beech litter production and the role of
snails in litter decomposition.
Oecologica 5:215-239.
Monson, R. A. 1971. Experimental transmission of Protostrongylus
to bighorn and mouflon sheep hybrids.
M.S. Thesis. Colorado
University, Ft. Collins.
49pp.
Monson,

stilesi

State

R. A. and G. Post. 1972. Experimental transmission of Protostrongylus
to bighorn-mouflon hybrids.
J. Parasitol. 58(1):29-33.

stilesi

Mozley, A. 1954. An Introduction to Molluscan
Population Studies of Freshwater Molluscs.
London.
Neck, R. W. 1976. Micro-distribution
talus slope.
Sterkiana 62:19.

Ecology.
Distribution
H. K. Lewis and Co.,

and

of land snails in an artificial

Owen, G. 1966. 2. Digestion.
In K. M. Wilbur and C. M. Yonge
Physiology of Mollusca, Vol. II. Academic Press.

(eds.)

Pillmore, R. E. 1955. Investigations of the life history and ecology
of the lungworm, Protostrongylus
stilesi.
Fed. Aid Div. Quart. Rept.
Colo. Dept. of Game and Fish:6l.
Pillmore, R. E. 1957. Lungworm and its relationship to bighorn sheep
management.
Proc. 37th Ann. Conf. of Western Assoc. State Game and
Fish Comm. 198.

�-40-

Pillmore, R. E. 1958a. Life Cycles of the lungworm genus Protostrongylus
in Colorado.
J. Colo. Wyo. Acad. Sci. 4:44.
Pillmore, R. E. 1958b.
Study of the lung nematodes of game animals.
Aid Div. Quart. Rept. Colo. Dept. of Game and Fish:l.
Pillmore, R. E. 1959. Study of lung nematodes of bighorn sheep.
Aid Div. Quart. Rept. Colo. Dept. of Game and Fish:73.

Fed.

Pillmore, R. E. 1961. Study of lung nematodes of bighorn sheep.
Aid Div. Quart. Rept. Colo. Dept. of Game and Fish:69.

Fed.

Pilsbry, H. A. 1948. Land mollusca of North America.
of Philadelphia Monographs No.3.

Acad. of Nat. Sci.

Post, G. and K. B. Winter.
1956-1957.
Life cycle of lungworms.
FVl-3-R-4,
Work Plan No. 11, Job No.2.
Randolph, P. A. 1973. Influence of environmental variability
snail population properties.
Ecology 54:933-955.
Retzer, J. L. 1956.
7(1):22-32.

Fed.

Alpine soils of the Rocky Mountains.

Wyo.

on land

J. Soil. Sci.

Riggle, R. S. 1976. Quantitative Examination of gastropod and soil
relationships in an oak-hickory forest in the lower Illinois Valley
region~
Sterkiana 62:1-17.
Rose, J. H. 1973. The lungworms
in Parasitology 11:559-570.

of domestic pigs and sheep.

Rufi, V. G. 1961. Life cycle of lungworms.
Wyo. Dept. of Game and Fish:60.
Russo, J. P. 1956. The desert bighorn
Ariz. Game and Fish Dept.
Walton, M. L. 1963.
76:127-131.

Fed. Aid Div. Quart. Rept.

sheep in Arizona.

Length of life in West American

Walton, M. L. 1970. Longevity
Nautilus 83:109-112.

Advances

in Ashmunella,Monadenia

State of

land snails. Nautilus

and Sonorella.

Wells, C. P. 1944. The water relationships of snails and slugs III.
Factors determining activity in Helix pomatia.
Jour. Exptl. BioI.
20:79-87.
Whitney, M. E. 1938. Some observations on the reproductive cycle of a
common land snail, Vallonia pulchella.
Influence of Environmental
factors.
Proc. Indiana Acad. Sci. 47:299-307.

�-41-

Winter, K. B. 1956. Life cyc1~ of 1ungworms.
Wyo. Dept. of Game and Fish:86.

Fed. Aid Div. Quart. Rept.

Yom-Tov, Y. and M. Ga1un. 1971. Note on feeding habits of the desert
snails Sphincterochila boissieri.
Ve1iger 14:86-88.

Prepared by
F. E. Latson

��January,

1979

-43JOB

State

C_O_L_O_RA_D_O

of

Project

r

Work Plan No.

Period

REPORT

_

W-41-R-28

No.

Job Title

FINAL

Bighorn

Sheep and Mountain

Job No.

24

Goat Investigations

E_x
__t_e_n_d_i_n_g
__B_i_g_h_o_r_n
S_h_e_e_p
__R_a_n_g_e_s

Covered:

Personnel:

June 1, 1977 - May 31, 1978

George D. Bear

ABSTRACT

Approval was not obtained for the bighorn transplants into Gunnison area
and in Cache la Poudre River Canyon, therefore no work was initiated in
those localities.
A final evaluation was made of 1975 Cache la Poudre
Canyon bighorn transplant in May, 1977 and the Cebolla Creek transplant
in May, 1978. A manuscript will be published as a special report by the
Colorado Division of Wildlife.

Prepared

by
George D. Bear
Wildlife Researcher

_

��January,

-45-

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

W-41-R-28

Bighorn
Job No.

Sheep and Mountain

1

Job Title

Burning to Improve and Enlarge Bighorn

Period Covered:

June 1, 1977-

Goat InvestigatioTIfl

~2~5

Work Plan No.
Prescribed

1979

_
Sheep Ranges

June 30, 1978

Personnel:
Dr. J. Barrows, L. Balic, and R. Yancik, Department of Forest
and Wood Sciences, Colorado State University; D. Canalte, T. Dailey, D.
Milchunas, and T. Woodard.

ABSTRACT
Treatment plots were delineated and pretreatment forage sampling was completed
during this segment.
Analyses of clip-plot data have not been completed.
Preliminary results indicate that precision of biomass estimates will be
poor and sampling intensity and/or size of quadrats will have to be increased
during post treatment sampling years.
Three bighorn sheep were successfully reared and trained for use in this
study. Food habits data were collected during the field training periods.
Forbs, grasses, and shrubs comprised 53.0, 40.2, and 6.8 percent, respectively,
of the composite diet of all sheep for three sets of grazing trials.

��-47-

PRESCRIBED BURNING TO IMPROVE AND
ENLARGE BIGHORN SHEEP RANGES
Thomas N. Woodard

P. N. OBJECTIVES
1.

To test the hypothesis that prescribed burning of understory
vegetation improves the production and quality of bighorn sheep
forage in a ponderosa pine-Douglas fir timber zone.

2.

To test the hypothesis that bighorn sheep use of burned
increases compared to adjacent unburned areas.

areas

SEGMENT OBJECTIVES
1.

Raise and train up to 4 bighorn

2.

Construct

bighorn

sheep pen facilities.

3.

Delineate

burned

and unburned

4.

Conduct pre-treatment
sampling plots.

5.

Coordinate pre-burn, burn, and post-burn activities with U.S. Forest
Service and Colorado State University Fire Science personnel.

treatment

vegetation

METHODS

lambs for use in foraging

trials.

plots on study areas.

sampling

and establish

permanent

AND MATERIALS

Three contiguous sites were delineated with flagging tape and metal stakes
placed along the boundaries.
Site 1 is 3.9 ha in scattered ponderosa pine-Douglas
fir type with numerous rock outcroppings on an east facing slope of 15-30 degrees.
Site 2 is 5.4 ha in mountain shrub type with east and south aspects on a
slope of 0-10 degrees.
Site 3 is 1.9 ha in perennial grassland on level
terrain.
The treatments will consist of burning and non-burning three each
0.3 - to 0.9 -ha plots per site; totaling nine plots for each treatment over
the three sites. Treatment plots were paired (replications) by apparent
homogeneity within each site with burning randomly assigned to one plot in
each pair (Fig. 1). Plots within replications are approximately equal in
size (Table 1).
The prescribed burning is scheduled during the August-October,
1978 time
period.
Fire Science personnel at Colorado State University have measured
pre- fire fuel loadings, determined the fire prescription,
(Appendix I) and will
monitor physical factors during the burning and measure post-fire fuel loadings.
U.S. Forest Service and Larimer County Sheriffs Department personnel will
conduct the burning with assistance from the Fire Science Department and
Colorado Division of Wildlife.

�-48-

Table 1. Treatment plot sizes for the response
prescribed burning study.

of bighorn

sheep to

Plot

srlte-1/

Replication

Treatment-

1

1

U

0.62

1

1

B

0.58

1

2

B

0.67

1

2

U

0.64

1

3

B

0.67

1

3

U

0.67

2

1

B

0.84

2

1

U

0.85

2

2

U

0.93

2

2

B

0.94

2

3

U

0.91

2

3

B

0.92

3

1

U

0.35

3

1

B

0.34

3

2

B

0.31

3

2

U

0.32

3

3

B

0.31

3

3

U

0.29

1/1
2

3

1:./

U
B

Ponderosa pine - Douglas
Shrub land type
Grassland type
Unburn
Burn

fir type

2/

Hectares

�-49-

Vegetation

Measurements

The following sampling schemes and techniques will be used to measure
vegetation response to prescribed burning.
Plant Cover
The rated microplot technique (Morris 1973) will be used to estimate aerial
plant cover in each treatment plot. Two hundred fifty, one- x two-inch
microplots will be located systematically in each treatment for representative coverage.
Aerial cover for selected species, forage classes, and total
understory, will be measured by rating the area occupied in each microplot
to the nearest one-tenth (rating score of one for each 1/10 area).
The
ratings for the 250 microplots will then be summed to obtain a measurement
value for each treatment plot. Measurements will be made during August,
each year for the duration of the study. Pretreatment sampling was conducted
during this segment.
Overall and within treatment sampling intensity may
be changed based on variances obtained in analysis of pretreatment sampling
data.
Herbage Yield
A clip and weight technique, as described below, will be used to estimate
above-ground biomass of flora during August of every year for the duration
of the study. A restricted random sampling design will be used to select
locations of 30 sampling units (0.25m2 rectangular quadrats) in each
treatment plot. Transect lines will be established to provide systematic
coverage of the entire plot. At regular intervals along each line a compass
heading and the number of steps on the heading will be randomly selected.
The sample location will be established by pacing the prescribed number of
steps on the compass heading and placing the quadrat frame next to the right
foot. All vegetation within the quadrat frame will be ground-level clipped,
separated, and placed in paper sacks labelled by species, date, treatment
plot, and quadrat number.
After the vegetation is clipped and sacked at
each sample location the investigator will return to the original transect
line, pace another interval and repeat the procedure.
Clipped vegetation
will be dried at 1000C until weight stabilizes (~ 24 hrs) and weighed to the
nearest O.lg.
During pretreatment sampling, completed this segment, all understory plant
species were separated and sacked separately.
The woody stem portions of
shrubs were sacked separately from leaves and current annual growth.
Individual plant species selected for measurements during post treatment
years will be chosen based on availability on the study area and importance
as a forage species.
All other plants will be composited according to forage
class (shrubs, grasses, and forbs).
Overall and within treatment sampling
intensity during post treatment years may be changed based on variances
obtained in analysis of pretreatment data.

�-50-

Forage Quality
Vegetation samples will be collected in late summer-early fall each posttreatment year in each treatment plot for forage quality analyses.
Plant
parts simulating those chosen by tame sheep during grazing trials, described
below, will be hand-plucked from plants grazed by the animals or from the
immediate area.
Separate 50-g samples w.i Ll be collected by treatment plot
and individual animal (3) for each plant species comprising 2 percent or
more of the diet of tame bighorn sheep.
Samples will be sealed in plastic
bags and frozen for later analysis.
Samples will be analyzed for crude
protein, calcium, phosphorus, acid detergent fiber, lignin, cell wall
constituents, gross energy, and in vitro digestibility.
Bighorn

Sheep Food Habits Measurements

Grazing trials will be conducted with tame, trained bighorn sheep to
measure response to treatments.
Advantages and problems associated with
direct observation of forage consumption by tame animals have been reviewed
by Wallmo and Neff (1970). Three male bighorn sheep lambs were successfully
reared and trained during this segment using techniques modified from those
described by Baker and Hobbs (1977). The training regime for these lambs
included 3 sets of grazing trials conducted during winter 1977-78.
The
procedures, study design, and study areas described by Baker and Hobbs (1977)
were utilized and the raw food habits data are presented in this report.
The following sampling scheme will be utilized during post treatment years
to measure bighorn sheep response to prescribed burning.
Grazing trials will
be conducted during three 10-day sampling periods conducted in November,
January, and March each winter.
Three tame bighorn sheep will be allowed
to graze one at a time for one hour each during AM and PM hours each of the
last 9 days of each sampling period.
The resulting sampling intensity will
consist
of six, I-hour grazing trials per day (2 trials each for 3 animals).
During the first day the animals will be allowed to graze freely on the study
area to become familiar with the vegetation and data will not be collected.
Starting points for individual trials will be the approximate center of
treatment plots and will be selected randomly without replacement for each
animal.
The result will be a start of a grazing trial for each animal in
each of the 18 treatment plots. When an animal is released at a starting
point its grazing route will not be influenced unless it leaves the study
area.
If, after five minutes the animal does not reenter the study area of
its own free will, it will be led to the nearest treatment plot center and
again allowed to graze. A portable tape recorder will be used to record
data during trials.
Data collected will include animal number; starting
time; starting treatment plot; number of bites of forage by species, plant
part, and treatment plot occupied; amount of total and grazing time spent
in each plot; and incidental activities and behavioral changes.
Data Analysis
Measured vegetation and animal variables will be analyzed with analysis
of variance.
Analysis of covariance will be done to adjust post treatment

�-51-

(dependent) cover values and herbage yields by respective
(independent) values.

pretreatment

STUDY AREA LOCATION
The study area is located in the Wintersteen Park area of the Cache la
Poudre River Drainage, 2 kilometers northeast of Rustic Larimer County,
Colorado.
RESULTS AND DISCUSSION
Vegetation

Measurements

Pretreatment plant cover and herbage yield sampling was completed during
August, 1977. A total of 110 species were encountered during microplot
sampling.
The number of plant species clipped, sacked, and weighed separately
totaled 121 including 26 grasses or grasslike species, 77 forbs, and 18 shrubs.
Analyses of these data have not been completed and no results are presented
in this report.
Preliminary results indicate that precision of estimates
will be poor and sampling intensity and/or size of quadrats will have to
be increased during post treatment sampling years.
All species comprising
2 percent or more of the above-ground biomass, based on this pretreatment
sampling, will be clipped and weighed separately during post treatment years.
Bighorn

Sheep Food Habits Measurements

Twenty-one plant
species comprised 90.3 percent of 77,058 bites observed
during grazing trials conducted during winter 1977-78 (Table 2). Percentages
of forbs, grasses, and shrubs in the diet increased, decreased, and remained
consistently low, respectively, as winter progressed (Fig. 1).

Table 2. Percentage composition of sheep diet across 3 animal and all
vegetation types. Species are ranked in order of importance across all
months.

Species

Antennaria

sp.

Calamagrostis

canadensis

Poa sp.
Phleum

pratense

Eriogonum

umbellatum

November

January

March

All Months

36.7

40.2

45.1

39.9

4.9

20.2

7.7

10.1

10.5

1.1

14.6

8.8

10.9

2.4

1.3

5.9

2.4

6.3

8.1

5.0

�-52-

Table 2. Percentage composition of sheep diet across 3 animal and all
vegetation types.
Species are ranked in order of importance across all
months.
(Cont'd).

Species

November

January

March

2.2

5.2

1.5

2.9

2.5

3.2

1.5

2.5

0.8

2.1

inermis

2.7

1.3

1.4

2.0

sp.

3.3

1.1

0.3

1.9

1.7

0.6

0.8

1.2 .

tridentata

0.4

2.7

0.4

1.1

sp. (leaves)

1.2

0.4

1.1

0.9

Muhlenbergia
Carex

sp ,

Potentilla
Bromus

hippiana

Sporobolus
Stipa

coma ta

Purshia
Salix

montana

All Months

2.2

Unidentified

forb

1.7

0.4

Deschampsia

caespitosa

1.4

0.4

0.2

0.8

(green needles)0.5

1.0

1.1

0.8

Pinus

ponderosa

0.9

Boutelous

gracilis

0.5

1.4

0.4

0.7

Fragaria

sp.

1.0

0.9

0.1

0.7

balticus

0.4

0.3

1.8

0.7

Juncus

Agropyron

smithii

1.4

0.1

0.2

0.7

Solidago

sp.

0.8

1.1

0.1

0.7

% Grasses

45.3

38.6

33.2

40.2

% Forbs

49.9

51.7

59.9

53.0

% Shrubs

4.8

9.7

6.9

6.8

34,405

22,648

20,005

77 ,058

Total Bites

�-53N

1

~---

SITE

BOUNDARIES

---REP.

BOUNDARIES

-

BOUNDARIES

--·-·-PLOT

1-1 =SITE - REPLICATION

SCALE:
&lt;&gt;

&lt;&gt;

&lt;&gt;&lt;&gt;&lt;&gt;
&lt;&gt; -e- -e- -e-

Fig.

1 in.. = 200 ft.

1.

BURN PLOTS

Design of the prescribed
burning to
improve and enlarge bighorn sheep ranges

study.

�-54-

60

FORBS

50

••••

LLJ

c

40

z
••••
Z
LLJ

o
a:

30

LLJ
a,

20

10

NOV

Figure 2.

JAN

MAR

Composition of bighorn sheep diet totaled across all vegetation
types and animals during winter 1977-78.

�-55-

The plant species listed in Table 2 and present on the prescribed burn
study area will be clipped and weighed separately regardless of availability
during post treatment years.

LITERATURE

CITED

Baker, D ..L., and N. T. Hobbs.
1977 . Systems modeling big game
populations simulations of the carrying capacity of the Rocky
Mountain National Park elk winter range. Pages 125-195 in Colorado
Division of Wildlife Game Research Report. July, Part Two. 125-305pp.
Morris, M. J. 1973. Estimating understory plant cover with rated
microplots.
D.S.D.A. Forest Service, Rocky Mountain Forest and
Range Experiment Station, Research Paper RM-104. 12pp.
Wallmo, O. C., and D. J. Neff.
1977. Direct observation of tamed deer
to measure their consumption of natural forages.
Pages 105-110
in Range and wildlife habitat evaluation - A research symposium.
D.S.D.A. Forest Service. Misc. Publ. 1147.

Prepared

by ~Q;;

Ai

w~J

Thomas N. Woodard
Wildlife Researcher

�-56-

APPENDIX 1
Report to Colorado Division of Wildlife

PRESCRIBED FIRE PLANNING
AND ANALYSIS FOR WILDLIFE HABITAT

Prepared By:

Jack S. Barrows
Principal Investigator

and

Richard F. Yancik
Graduate Research Asst.

March 1978

Colorado State University
Department

of Forest and Wood Sciences

Fort Collins, Colorado 80523

�-57-

WINTERSTEEN
PRESCRIBED

ARAPAHO-ROOSEVELT

PARK

BURNING PLAN

NATIONAL FOREST, R-2

RED FEATHER LAKES RANGER DISTRICT

�-58-

I.

INTRODUCTION

The proposed Wintersteen Park prescribed fires are contained within the
ponderosa pine (Pinu~ ponderosa) zone along the Colorado Front Range in the
Arapaho-Roosevelt National Forest.

The purpose of these prescribed burns is

to establish a research site for the Colorado Division of Wildlife Bighorn
Sheep Habitat Study Project headquartered in Fort Collins, Colorado.

Agencies

involved with the purposed prescribed burns are the Colorado Division of Wildlife, U.S. Forest Service (Arapaho-Roosevelt National Forest), and Colorado
State University (Fire Science Program).

II.

OBJECTIVES
The primary objective of the prescibed burns is to establish a research

site to evaluate the impacts (favorable and/or unfavorable) of fire on Rocky
Mountain bighorn sheep (Ovis canadensis) and their habitat.
Specific objectives of the prescribed fire studies are as follows:
1.

To test the hypothesis that prescribed burning of understory
vegetation improves the production and quality of bighorn sheep
forage in a ponderosa pine-Doug las-fir timber zone.

2.

To test the hypothesis that bighorn sheep use of burned areas
increases compared to adjacent unburned areas.

III.

DESCRIPTION OF THE AREA
The Wintersteen Park burns are located in the Cache La Poudre River Drain-

age, two kilometers northeast of Rustic, Larimer County, Colorado (refer to map
and photo).

The area is adjacent to private land ownership.

Access to the

area is somewhat limited as a result of nearby private land ownership.

Primary

access is via the Delatour Boy Scout Ranch or the Leroy-Robinson Road, which
is suitable for 2-wheel drive pick-up trucks (refer to map).

�-59SUMM..~Y OF PERTIEt.'T

INFORK"-TION

Legal Description:

Section 27

Elevation:

8200

Aspect:

South-Southeast

Slope Range:

3-50 percent

Topographic

Features:

Township·9N

Range 73W

Large rock outcrops exists near ridgeline.
Swale-lL~e

feature and aspect create erratic

wind situations.
Fuel Types:

sent the distribution
components
(1) Decandent

ponderosa pine overstory.

grass species and big sagebrush
eonstitutes

Tables 1 and 2 pre-

Three fuel types exist.

of fuel loading by fuel

for each of the three fuel types.
Heterogeneous

(Artemisia tridentata)~

the surface fuel component.

understory

of bunch-

and fine woody material

Ground fuel component

is primary pine

needle litter.
(2) Homogeneous
bunchgrass

species.

(3) Homogeneous

surface fuel component of big sagebrush intermixed with
Ground fuel component is herbaceous

surface fuel compon~nt dominated by bunchgrass

with isolated clumps of big sagebrush.
of herbaceous

Ground fuel component

litter.

Planned Burn Area:
Unit

Total:

and shrub leaf litter.

Acreage

I-IS

1.44

1-2N

1.64

1-3N

1.66

2-lW

2.08

2':2S

2.32

2-3E

2.26

3-lS

0.83

3-2N

0.76

3-3N

0.76

9

13.75

species

consist mainly

�-60-

Water Resources:
area.

There are no live streams within or adjacent to the

An intermittent stream exists in the area; however, it will not pro-

vice a sufficient water supply.

Reliable water resources are the Poudre

River and Red Feather Lakes.

IV.

PRESCRIBED BURNING PRESCRIPTIONS

Ponderosa Pine Fuel Type
(1) Time of year:

Between September 1 and November 15, 1978
when grasses are 70% cured.

(2) Time of Day for Ignition:
(3)

Between 1000 and 1400 hours.

Fuel Moisture Ranges at 1000 hrs.:
Ground fuel (litter):

4 - 10%

Herbaceous vegetation:

4 - 8 %

One-hour timelog fuels:

4 - 5 %

Ten-hour timelog fuels:

4 - 12%

0

0

(4)

Air Temperature:

45 F to 80 F

(5)

Relative Humidit~:

5% to 18%

(6)

Wind Velocit~:

Between 3 mph and 10 mph.

(7) Wind Direction:

Preferably south-southeast.

(8) State of Weather:

Clear to partly cloudy with no major fronts
approaching.

Less than .2 inch of precipi-

tation 2 days prior to burning date.
(9) Atmospheric Stability:

Smoke dispersal is not critical.

However a

temperature inversion should be avoided.
Big Sagebrush Fuel Type
(1) Time of year:

Between September 1 and November 15, 1978
when grasses are 70% cured.

(2) Time of Day for Ignition:

Between 1000 and 1400 hrs.

(3) Fuel Moisture Ranges at 1000 hrs.:
Ground fuel (litter):

4 - 10%

Herbaceous vegetation:

4 - 7 %

One-hour timelog fuels:

4 - 10%

Ten-hour timelog fuels:

4 - 12%

�-61-

0

(4) Air Temperature:

0
45 F to 80 F

(5) Relative

5% to 15%

(6)

Humidity:

Wind Velocity:

3 mph and 10 mph with gusts

Between

to

15 mph.
(7) Wind Direction:

Preferably

(8) State of Veather:

Clear to partly

south-southeast.

approaching.
tation
(9) Atmospheric
Bunchgrass

Fuel

Stability:

cloudy with no major

fronts

Less than .2 inch of precipi-

2 days prior

Smoke dispersal

to burning

date.

not critical.

Type

(1) Time of Year:

Between

September

when grasses
(2)

Time of Day for Ignition:

(3)

Fuel Moisture

Ranges

Between

are 70% cured.

1000 and 1400 hrs.

fuel

Herbaceous

(litter):

4 - 10%

vegetation:

4 - 7 %

One-hour

timelog

fuels:

4 - 8 %

Ten-hour

time10g

fuels:

4 - 10%

0

0

(4) Air Temperature:

45 F to 80 F

(5) Relative

5% to 15%

(6) Wind Velocity:

15, 1978

at 1000 hrs.:
Ground

Humidity:

1 and November

Between

3 mph and 12 mph with gusts

to

15 mph.
(7) Wind Direction:

Preferably

(8) State of Weather:

Clear to partly

south-southeast.

approaching.
tation
(9) Atmospheric

Stability:

cloudy with no major

Less than

2 days prior

Smoke dispersal

fronts

.2 inch of precipi-

to burning

not critical.

date.

�-62-

V. POTENTIAL FIRE BEHAVIOR
The "predicted" potential fire behavior data to be presented in the latter
part of this section were derived using site-specific fuels and weather data
as inputs to predictive models for estimating potential fire behavior.

These

fire behavior estimations are "rough" and are to be viewed with caution. These
fire behavior estimations, when used in conjunction with experience and knowledge, were useful in the burning prescription formulation designed to achieve
the objectives set forth earlier.
These fire behavior predictions were useful for estimating the magnitude
of changes in fire behavior as burning conditions varied for each fuel type.
Ponderosa Pine Fuel Type
. From
1.9

To
13.6

(2) Flame Length (feet)

1.S

4.3

(3) Fireline Intensity (BTU/ft. second)

16.4

126.6

(4) Reaction Intensity (BTU/ft:/minute)

2247.4

2S08.5

3.S

4.7

(1) Rate of Spread (feet/min.)

2.4

20.3

(2) Flame Length (feet)

1.6

4.9

(3) Fireline Intensity (BTU/ft. second)

19.2

182.6

(4) Reaction Intensity (BTU/ft~/minute)

2028.4

2278.3

3.8

7.2

(1) Rete of Spread (feet/min.)

4.2

S4.9

(2) Flame Length (feet)

1.3

5.3

(3) Fireline Intensity (BTU/ft. second)

14.6

217.4

(4) Reaction Intensity (BTU/ft~/minute)

1270.S

1447.1

2.7

8.8

(1) Rate of Spread (feet/min.)

(S) Crown Scorch Height (feet)
Big Sagebrush Fuel Type

(S) Crown Scorch Height (feet)
Bunchgrass Fuel Type

(S) Crown Scor~h Height (feet)

�-63-

Summary.

The foregoing analysis showed minimum and maximum values for the

range of prescription factors analyzed.

Following a fire weather study of

probable wind and fuel moisture values a decision prescription will be prepared that will tighten the lower and upper limits for rate of spread, flame
length and fireline intensity.
VI. PREBURN PREPARATIONS
Weather Station
A portable weather station will be located on the proposed burn area.

The

weather station should be operational three days prior to the burning date.
On-site data also will be obtained using fuel moisture sticks and a belt weather
kit.
A special spot-weather forecast will be obtained prior to burning and
every hour (or sooner if necessary) during the burning operation.

The special

spot-weather forecast and on-site information will be the basis for decisionmaking on burning operations.
Control Line
A five-foot wide black line will be established around the portion of
each unit to be burned.

A six-foot wide "black" line will be used between

adjacent units to be burned (refer to map).
be established.

Water resource availability should

VII. FIRING PLAN
Each fuel type will have a Firing Plan so as to account for changes in
burning conditions.

Burning priority will be, if conditions exist, as follows:

(1) Ponderosa pine type, (2) Big sagebrush type, and (3) Bunchgrass type.
Firing will be conducted by the designated Ignition Crew only under supervision of Firing Boss.
Any modifications to the Firing Plans, incluiing fireing techniques and
sequences, must be approved by the Fire Boss and formally announced to all
personnel involved with the prescribed burn.
Ponderosa Pine Fuel Type (Units 1-1, 1-2, and 1-3
The Firing Plan (portrayed on the map) is based on a south or southeast
windflow.

Strip firing down slope at intervals of 50 to 75 feet will be used

�to allow short head fires to spread with the wind and slope.

Within some

units, there are small areas with fuel conditions that will reguire a backing fire down slope to achieve the proper intensity and/or objectives.

These

areas will be identified by the Firing Boss and Fire Behavior Research Boss.
Wind directions other than those previously mentioned will not necessarily
preclude burning operations.

The firing sequence and techniques used will be

modified, in agreement with Fire Boss, by the Firing Boss to include changing
burning conditions.

Care should be exercised when burning near ridgeline.

With a south or southeast wind, the firing sequence will be Unit 1-1,
Unit 1-2, and Unit 2-1 or Unit 1-3.

Refer to the map for strip firing and

black lining information.
Big Sagebrush Fuel Type (Units 2-1, 2-2, and 2-3)
The Firing Plan (see map) for these units is similar to the Firing Plan
for the Ponderosa pine fuel type, with the exception of stripping intervals and
type of fire.

Critical burning conditions (lower end of burning presciption)

coupled with brush fuels may necessitate reducing stripping intervals to 40 feet
or using a backing fire down slope for control purposes.

A backing fire down

slope may deem necessary to achieve proper intensity for favorable vegetation
responses.
The firing sequence, with a south or southeast wind, will be Units 2-1,
2-2, and then 2-3.

This firing sequence is subject to modification depend-

ing upon burning conditions.
Bunchgrass Fuel Type (Units 3-1, .3-2, and 3-3)
The Firing Plan (refer to map) for this fuel type requires a backing
fire down slope to achieve the desirable intensity for vegetation responses.
Burning conditions resembling the lower end of prescription provide the
basis for the Firing Plan associated with this fuel type.
With a south or southeast wind, the firing sequence will be Units 3-1,
3-2, and then 3-3.

�-65(To be prepared by Control Organization)

VIII. CONTAINMENT

AND MOP-UP PLAN

IX. BRIEFING AND ORIENTATION
X. SAFETY PRECAUTIONS
XI. MANPOWER

NEEDS

XII. EQUIPMENT NEEDS
XIII. SMOKE AND VISIBILITY

HAZARDS

XIV. PUBLIC RELATIONS
XV . ENVIRONMENTAL

ANALYSIS

�Table 1.

Pre-fire fuel loading contributed by ground and surface fuel components of the Wintersteen Park prescribed burning site.

Loading by Ground and Surface Fuel Components (Tons/Acre)

---------.Forest Floor

Downed Woody Size Classes
Fuel Type

Avg. Planar
Slope (percent)

Total})

Herbaceous Vegetation

-0.0-0.24"

0.25-0.99"

1.0-2.99"

3.0" + sound

3.0" + rotten

Litter

live

dead

Fuel Loading

_------- -~--

..

.4638

.1013

.2362

6.2386

.7787

.4877

.1017

.2372

8.9589

.0548

.2905

.1297

.3026

3.2628

Ponderosa pine

35.0

.1593

.1696

.2362

Shrub

18.3

.2575

.6666

Bunchgrass

9.2

.0869

.1506

}j

.2023

_ .._---.-

----------Footnotes:

.3041

Includes loading contributed by shrub components.

---

I
0"
0"
I

�Table 2.

Pre-fire fuel loading contributed hy shrub comporrents of the Wintersteen Park prescribed burning site.

Fuel Loading Contributed by Shrub Components (Tons/Acre)

��-69-

JOB PROGRESS

State of

COLORADO
-------------------------------

Project

No.

Work

Job Title

3
Trapping

Covered:

Personnel:

REPORT

Bighorn Sheep
and Mountain Goat Investigations

W-4l-R-28

Plan No.

Period

.Ianua r y 1979

Job No.
and Translocating

June 1, 1977

through

1
------------=--------------------

Bighorn

Sheep

June 30, 1978

R. Schmidt, M. Connors, V. Feuerstein, M. Middleton, F. Weber,
·P. Yates,W.
Rutherford, plus WCO's and Area Supervisors in
specific localities.

ABSTRAcr

Five separate bighorn sheep trans1ocations were done during this segment.
Twelve sheep were released at Riverside, Chaffee County; 17 an Langhoff Gulch,
Chaffee County; 20 at the South Fork of Conejos River, Conejos County; 22 at
Alamosa River, Conejos County; and 20 at Rampart Range (Monument Creek), El
Paso County, for a grand total of 91. All translocated sheep were either
neck-collared or ear-tagged, and were treated with the antihe1minthic
drug
Fenbendazole as a means for controlling lungworm.
Monitoring of the effects
of drug treatment was done on sheep populations translocated during the
previous year, as well as on many populations treated in place.
Monitor data
generally show that drug treatment is effective in reducing or eliminating
lungworm infestation for a period of several months, but must be repeated
annually for long-term results.

��-71-

TRAPPING AND TRANSLOCATING

BIGHORN SHEEP

Robert L. Schmidt and William H. Rutherford

P. N. OBJECTIVE
The principal objective of this study will be to increase the total number
of bighorn sheep within the State of Colorado by either introducing bighorn
into areas which were historically bighorn sheep ranges and where no animals
are in evidence today, or by adding additional animals to small remnant herds.
By expanding bighorn sheep from crowded areas into other areas it is hoped
that they will be better able to utilize the range and increase their numbers.
Also the introduction of new animals, especially rams, into remnant populations
should improve the reproductive capacity and livability of the lambs by
improving the gene pool and lowering the percentage of inbreeding.
SEGMENT OBJECTIVES
1.

Trap and translocate

between

100 and 150 bighorn

sheep.

2.

Treat all captured bighorn
control of Protostrongylus

3.

Monitor all translocated bighorn sheep for the degree of parasitism by
collecting fecal pellet groups and checking them by standard laboratory
procedures.

sheep with chemotherapeutic
spp. lungworm parasitism.

drugs for the

}ffiTHODSAND MATERIALS
Baiting, trapping, drug treatment, and monitoring procedures are described in
detail in the Job Progress Report for 1976-77 (Schmidt 1978) and will not be
repeated here.
RESULTS
Five separate bighorn sheep translocation operations were conducted during
this segment.
Information on these is presented in Tables 1, 2, 3, 4, and 5.
All bighorn sheep listed in Tables 1 through 5 were treated with Fenbendazole,
an antihelminthic drug that is efficacious against adult lungworms.
This
treatment was done to insure that the animals were "clean" when released at
the new sites.
Follow-up monitoring of lungworm larvae output was done on sheep that were
translocated during 1976-77, as well as being done on numerous herds that
received drug treatment without being translocated.
Results of this
monitoring are presented in Table 6.
With few exceptions, Table 6 shows that larval output from treated sheep is
generally low. Data from Pikes Peak show that re-infection with new populations of lungworms, with a corresponding increase in larval output, occurs

�-72-

within a year following treatment.
This particular bighorn sheep range
carries a high level of infectious third-stage larvae in snails, so that
re-infection of treated sheep is a near certainty.
This suggests that
treatment in such situations, in order to be considered successful over
a period of several years, must be done annually.
Data from Chalk Creek are exactly opposite from what would be expected,
giving rise to suspicions that two different groups of sheep were involved.
This was confirmed by WCO Travnicek, who reported that the sheep that had
previously been trapped, treated and marked had moved away from the area
and that a separate group of unmarked sheep had taken their place.
These
data, therefore, are meaningless and are included only for the purpose
of pointing out some of the pitfalls involved in monitoring treated sheep.

Table 1. Bighorn sheep trapped at Boulder Park, Pikes Peak, released at
Riverside, Chaffee County, TI3S, R79W, Sec. 1.

Date

Sex

Age

Ear Tag Color
and Number

3-23-78

M

Yr1g.

Red/White

- 2

None

"

M

Yrlg.

Red/White

- 3

None

"

F

9

Red/White

- 4

Red/Black

- 63

"

F

10

Red/White

- 5

Red/Black

- 67

"

M

Lamb

Red/White

- 7

None

"

F

10

Red/White

- 8

Red/Black

- 68

"

F

9

Red/White

- 9

Red/Black

- 65

"

F

10

Red/White

- 10

Red/Black

- 66

3-28-78

M

Yr1g.

Black - 90

None

"

F

10

Black - 93

Red/Black

- 74

"

F

9

Black - 95

Red/Black

- 73

"

F

2

None

Red/Black

- 71

Neck Collar Color
and Number

�-73-

Table 2. Bighorn sheep trapped at Sugarloaf Mountain, Tarrya1l Creek;
released at Langhoff Gulch, Chaffee County, T12S, R79\-1, Sec. 16.

Date

Sex

Age

Ear Tag Color
and Number

Neck Collar Color
and Number

1-26-78

M

Lamb

Yellow - 1

None

"

M

Lamb

Yellow - 2

None

"

M

Lamb

Yellow - 3

None

"

F

Lamb

Yellow - 4

None

"

M

Lamb

Yellow - 5

None

"

M

Yr1g.

Yellow - 6

Yellow - 50

"

F

3

Yellow - 7

Yellow - 45

"

F

3

Yellow - 8

Yellow

"

M

3

Yellow - 9

White - 7

"

F

8

Yellow - 10

White - 29

"

F

4

Yellow - 11

Yellow - 26

"

M

2

Yellow - 12

Yellow

- 47

"

M

Yr1g.

Yellow - 13

Yellow

- 48

"

F

4

Yellow - 14

Yellow - 46

"

M

Yrlg.

Yellow - 15

Yellow

- 49

"

F

4

Yellow - 16

Yellow

- 44

"

F

6

Yellow - 17

White - 22

Remarks

- 18

Recapture
1976

from

Recapture
1976

from

�-74-

Table 3. Bighorn sheep trapped at Arrowhead, Poudre Canyon, Larimer County,
released at South Fork of Conejos River, Conejos County, T35N, R4E, Sec. 36.

Date

Sex

Age

Ear Tag Color
and Number

Neck Collar Color
and Number

2-9-78

M

Lamb

Black - 1

None

"

M

Lamb

Black - 2

None

"

M

Lamb

Black - 3

None

"

M

Lamb

Black - 4

None

"

M

Lamb

Black - 5

None

"

M

Lamb

Black - 6

None

"

F

6

None

"

M

Lamb

Black - 7

"

F

7

None

"

F

Lamb

Black - 8

None

"

M

Lamb

Black - 9

None

"

F

5

Black - 10

Unnumbered

"

F

4

None

White/Black

- 42

"

F

5

None

White/Black

- 43

"

F

4

None

White/Black

- 44

"

F

7

None

White/Black

- 45

"

F

4

None

White/Black

- 46

"

F

Yrlg.

None

White/Black

- 47

"

F

Yrlg.

None

White/Black

- 48

"

F

5

None

White/Black

- 50

White/Black

Remarks

- 40

None
White/Black

- 41

Yellow collar
with radio
transmitter

�-75-

Table 4. Bighorn sheep trapped at Almont, Taylor River Canyon, released
at Alamosa River near Platoro, Conejos County, T36N, R5E, Sec. 4.

Date

Sex

Age

Ear Tag Color
and Number

Neck Collar Color
and Number

2-22-78

F

7

White - 93

Red/Black

- 28

"

F

3

White - 95

Red/Black

- 30

"

F

6

White - 96

Red/Black

- 31

"

F

3

White - 85

Red/Black

- 32

"

F

4

White - 89

Red/Black

- 33

"

F

Yrlg.

Black - 67

Red/Black

- 35

"

F

5

White - 81

Red/Black

- 36

"

F

Yrlg.

White - 92

Red/Black

- 37

"

F

4

White - 91

Red/Black

- 38

"

F

Yrlg.

White - 98

Red/Black

- 39

"

F

3

White - 83

Red/Black

- 40

"

F

2

Black - 66

Red/Black

- 41

"

M

Lamb

Black - 63

None

"

M

Yrlg.

Black - 64

None

"

M

Lamb

Black - 65

None

"

M

Lamb

White - 82

None

"

F

Lamb

White - 87

None

"

F

Lamb

White - 90

None

"

F

Lamb

White - 94

.None

"

F

Lamb

White - 97

None

"

M

Yrlg.

White - 99

None

"

M

2

White - 86

None

Remarks

Collar on
upside down

�-76-

Table 5. Bighorn sheep trapped at Trickle Mountain,
on Rampart Range, El Paso County, Tl1N, R67W.

Saguache

Creek; released

Date

Sex

Age

Ear Tag Color
and Number

Neck Collar Color
and Number

3-8-78

F

Lamb

Black - 75

None

"

F

Lamb

Black - 76

None

"

M

Lamb

Black - 77

None

"

M

Lamb

Black - 78

None

"

F

Lamb

Black - 79

None

"

F

Lamb

Black - 80

None

"

M

Lamb

Black - 81

None

"

F

Lamb

Black - 82

None

"

F

Lamb

Black - 83

None

"

M

Lamb

Black - 84

None

"

M

Yrlg.

Black - 86

None

"

M

Yrlg.

Black - 87

None

"

M

Yrlg.

Black - 88

None

"

F

Yrlg.

None

Red/Black

- 50

"

F

3

None

Red/Black

- 51

"

F

2

None

Red/Black

- 52

"

F

6

None

Red/Black

- 53

"

F

4

None

Red/Black

- 54

"

F

4

None

Red/Black

- 56

"

F

Yrlg.

None

Unnumbered yellow collar with
This sheep
radio transmitter.
found dead 5-16-78, collar
recovered.

Remarks

�-77-

Table 6. Lungworm larvae output as determined by laboratory analysis of
bighorn sheep pellet groups collected before and after drug treatment.

Location and
Treatment Status

Collection
Date

No. of
Sample
Groups

Frequency
1/
By Output Category 2
4
0
1
5
3

2/
Average-

Dinosaur National Monument
Pre-treat
6-5-77
Post-treat
4-30-78

14
46

14
46

0
0

0
0

0
0

0
0

0
0

0.00
0.00

C ross M ountaln. 3/
Post-treat
Post-treat
Post-treat
Post-treat

6-8-77
8-3-77
10-6-77
4-30-78

22
15
21
20

15
15
15
18

7
0
6
2

0
0
0
0

0
0
0
0

0
0
0
0

0
0
0
0

0.32
0.00
0.29
0.10

Cow Creek, RMNpl/
Post-treat

6-9-77

6

6

0

0

0

0

0

0.00

Apishapa Canyo~/
Post-treat
Post-treat
Post-treat

6-24-77
8-11-77
9-29-77

24
7
25

16
7
5

8
0
19

0
0
1

0
0
0

0
0
0

0
0
0

0.33
0.00
0.84

Lone Pine Creek1/
Post-treat
Post-treat

6-27-77
7-27-77

5
2

4
0

1
2

0
0

0
0

0
0

0
0

0.20
1.00

Rampart Range
Pre-treat
Post-treat

8-8-77
3-18-78

11
26

3
14

6
7

2
4

0
1

0
0

0
0

0.91
0.69

11-15-77
12-28-77
1-15-78
2-26-78

4
21
8
20

0
1
2
0

2
7
4
10

1
9
2
6

1
2
0
3

0
2
0
0

0
0
0
1

1. 75
1. 86
1.00
1.80

Chalk Creek
Pre-treat
Post-treat

1-10-78
1-28-78

11
12

5
0

4
4

2
5

0
3

0
0

0
0

0.73
1. 92

5/
Tarrya11 RangePre-treat

1-l3-78

27

1

21

5

0

0

0

1.15

5/
Mount Evans-Pre-treat

1-11-78

21

0

2

6

6

2

5

3.10

Pikes Peak~/

-------------------------------------------------------------------------------

�-78Table 6. Lungworm larvae output as determined by laboratory analysis
bighorn sheep pellet groups collected before and after drug treatment

Collection
Date

No. of
Sample
Groups

Almont-Taylor Canyon
Pre-treat
Pre-treat
Post-treatment

1-10-78
2-14-78
2-24-78

14
21
21

Dillon Mesa
Pre-treat
Post-treat

2-14-78
3-17-78

21
20

GrantPre-treat
Pre-treat
Pre-treat

1-18-78
1-21-78
1-31-78

19

Trickle Mountain
Pre-treat
Pre-treat
Post-treat

Location and
Treatment Status

Frequency
1/
By Output Categoryo
1
2
3
4
5

14

o

o

7

7

6

15
0

o
o

o
o
o

of
(cont.).

Average-

o
o
o

1.50
1. 71
0.00

2.57
0.35

21

o

o

3
7

8

6

3

1

13

o

o

o

o

o
o

8

2

o

21

1

13

6

1

o
o

o

4

6
4

3

9

1-10-78
3-4-78
4-14-78

25
20
44

7
13
41

17
6
3

1
1
0

o
o
o

Bison Reservoir
Post-treat

4-14-78

20

10

7

2

Poudre Canyon
Pre-treat
Post-treat

1-15-78
2-1-78

51
29

3
29

22

26
0

2/

5/

o

o

1. 95
1.89
1.33

o
o
o

o
o
o

0.76
0.40
0.07

o

o

1

0.80

o
o

o
o

o
o

1.45
0.00

1

1/

- Larval output category values are assigned as follows: Larvae absent - 0;
1-50 larvae/gram of feces - 1; 51-250 larvae/gram - 2; 251-500 larvae/gram - 3;
501-1000 larvae/gram - 4; 1000+ larvae/gram - 5.

1/ This

is a weighted value established by multiplying category values by
frequencies, summing, and then dividing the summation by the total number of
sample groups.
Generally, values lower than 1.00 indicate output levels low
enough to be no cause for concern.

3/

- These are transplanted sheep that were treated at the time of capture.
treatment larval output data were not collected.
~/ These data cannot be considered post-treatment
8-10 months prior to pellet group collection.

1/ Post-treatment

data are not available

because

treatment

Pre-

was done

at the time of this writing.

�-79-

LITERATURE

CITED

Schmidt, R. L. 1978. Trapping and translocating bighorn sheep.
Federal Aid P-R Progress Report, Project W-41-R-27, January:

Prepared

by
Wildlife

Researcher

~

Colorado
57-75.

�TABLE OF CONTENTS
Bighorn

Sheep and Mountain

Goat Investigations

(W-41-R-28)
Page

Bighorn Sheep Lungworm Control by Biological Control
of Snail Intermediate Hosts •.......•.........•.............

25

Evaluation of Trace Mineral Availability to Bighorn Sheep
on Winter Range in Colorado •....•.........••.........•.....

I

Extending

43

Bighorn

Sheep Ranges

Prescribed Burning to Improve and Enlarge Bighorn
Sheep Ranges •.•....•..•.•...•••••.........•••.•............

45

Trapping

....•................•

69

••......•......•.•.......

81

Raptor

and Translocating

Investigations

Bighorn

Sheep

(W-124-R-6)

Bald and Golden Eagle Nes ting Studies
Bald and Golden Eagle Winter Population
Ferruginous
Nesting

Hawk Nesting

Performance

Osprey Nesting

Surveys

.•........•....

87
111

Studies

of Peregrine

Falcons

in Colorado

121
95

Studies

Peregrine Flacon Habitat Protection Activities
In Colorado ...•...•.••....•.••...•....•...•.........•......

171

Physical and Biological Analysis of Colorado
Peregrine Nesting Habitat ..•.••........••••..•...•....•...•

137

Prairie

Falcon Breeding

Population

Prairie

Falcon Nesting

Studies

Characteristics

Studies

.... 105

.......•.•.•.•..................

99

Reintroduction
and Augmentation of Peregrine
Falcon Production ••••.........•..••..•.....•........•......

163

�January

1979

-81-

JOB PROGRESS

REPORT

State of ----~~~~~------------COLORADO
Project No.

W-124-R

Work Plan No.
Job Title
Period

Job No.

Bald and Golden Eagle Nesting

Covered:

Personnel:

I

Raptor Investigations

'C.

1

Studies

April 15, 1977 through February

28, 1978

Gerald R. Craig, Alden Forbes, Rickie Londe, Joe Frothingham,
James McKinley, Wayne Russell and Gordon Saville, Colorado
Division of Wildlife
ABSTRACT

A total of 464 golden eagle nests on record necessitated orientation of
nesting investigations toward aerial flights to make most efficient use of
field time. Flight routes were delineated and 330 nests were checked during
the season •. An estimated 144 young were produced for an average of 1.48
young produced and 1.47 young fledged per successful nests.

��-83-

BALD AND GOLDEN EAGLE NESTING

STUDIES

Gerald R. Craig

Eagles are the largest aerial predators and occupy a unique niche which
makes them most likely to be affected by man caused changes in the ecosystem.
Golden eagles occupy regions in which massive habitat changes are likely to
occur as a result of land, water and energy developments and thus they may
serve as indicator or key species which could be monitored to assess the
results of the changes.
The nest sites of golden eagles are often used for
many decades which should provide some rigidity in a census system to monitor nest site occupancy and reproduction.
Approximately 250 golden eagle
nests have been documented and cataloged.
A sample of these nests has been
identified to evaluate the effectiveness of aerial census in determining
occupancy and reproduction.
pnly two active bald eagle nests have been documented in Colorado although
the state is generally considered to be peripheral as nesting habitat.
Recent reclassification has designated bald eagles as endangered species in
Colorado thus each site should receive protection from disturbance and habitat degradation.
Prior to such protection, it will be necessary to delineate
important hunting areas and buffer zones adjacent to the nest sites.
P.N.

OBJECTIVES

The objectives of this study are: (1) to estimate the breeding numbers and
obtain production data of bald and golden eagles nesting in Colorado; (2)
to identify important nesting areas and associated hunting habitats of bald
and golden eagles in Colorado; and (3) to compile data and submit reports
to associated state personnel and federal agencies for use in delineating
and protecting eagle nest sites.
SEGMENT OBJECTIVES
la.

Continue to locate and map nesting sites of golden and bald eagles
throughout Colorado.
Nest searches will be conducted with fixedwing aircraft and in some instances with helicopter.
Additional
field work will be done from the ground by vehicles.
Photographs
of nest sites will be taken and pertinent information recorded about
physical features of the habitat.

lb.

Nesting areas will be stratified by such features as climate, elevation and habitat type. Sample areas then will be delineated which
are representative of important nesting areas.
The sample areas
throughout the state will be flown with a fixed-wing aircraft in late
April and early May to ascertain the number of active sites.
The same
sites again will be checked from the air in June to determine the
nesting success and productivity of the sample sites.
The percent
of active sites, percent of successful pairs, and total production
will be extrapolated for each region of the state.

�-84-

2a.

When nesting areas are visited in la, details will be recorded as
to characteristics
of the nesting habitat.
Nesting eagles also will
be observed from a distance to locate and map key hunting areas.

2b.

Radio transmitters may be attached to several young bald eagles and
a sample of young golden eagles to follow their movements after they
fledge.
Color markers mayor
may not be used to mark the eagles as
well.

2c.

A sample of sites will be selected and visited to determine the prey
species present at the nest sites.
This information will be valuable
in assessing the prey composition and availability to eagles.

3.

Analyze

all data obtained

METHODS

and prepare

a report of the findings.

AND MATERIALS

During 1977, emphasis was placed on assessing activity and production of
golden eagles through use of fixed-wing aircraft.
An effort was made to
check all nests recorded during previous years, but solitary or extremely
isolated sites were excluded in interest in saving time. Primary areas of
inventory were in northwestern and southwestern portions of the state.
The
first flights were made in late April and May to locate active nest sites.
Sites were considered active if one or both adults were present at the site
and if there was evidence that new material had been added to the nest.
Nesting activity of some adults were more advanced than others and incubating
adults were often encountered.
In addition to recording presence and activity
of adults, and stage of nest repair, observers also took polaroid photographs
of each site from the aircraft to aid relocating the nests during return
visits.
At times, prior scheduling or weather conditions did not permit flights and
nests had to be checked from the ground.
Since incubating adults are often
difficult to distinguish from their nests when viewed from the ground, a
predator call was used to cause them to move on the nest.
Those nests which were determined to be active were revisited in late May
and June to record nesting success.
The number of young and approximate ages
were recorded during these flights.
Young were considered to be of fledging
age if they were completely feathered and no white down was visible on their
heads.
For tabulation purposes, young present in nests at least two weeks
prior to fledging were considered as fledged since mortality is insignificant
which might occur from that age to fledging.
The two known bald eagle nests present in the state were observed at a distance several times during the breeding season to determine activity and
production.
The nests were not visited at any time and special precautions
were made to avoid any activity which might disturb the adults.

�-85-

RESULTS AND DISCUSSION
Golden Eagle Nesting

Results

Table 1 su~marizes results of the 1977 field season.
The total number of
young fledged (99) is deceptive because many of the nests could not be
revisited before the young had fledged.
Analysis of those sites for which
the number of young produced and fledged are known indicates that only one
young did not fledge of 102 which were produced.
(Table 2). Therefore,
it is probable that nearly all the young produced also fledged, so a figure
of 1.11 young fledged per active nest may be appropriate.
Twenty-seven
sites were recorded which produced 47 young, but the observers had been unable to return prior to fledging so no fledging success could be obtained.
Assuming that these sites also experienced low mortality between the time
the young were observed and should have fledged, they can be added to 70
sites with known outcomes to obtain 97 sites which produced and fledged
143 young, or 1.47 young per successful site.

Table 1.

Statewide

Total of Golden Eagle Nesting Activity

Nests on record
Nests not found in 1977
Nests not checked
Nests checked
New nests found
Inactive nests
Active nests
Young produced
Young fledged
Young produced per active nest
Young fledged per active nest
Active nest per nest checked

Table 2.

Productivity

of Successful

- 1977

Totals
464
10
124
330
36
198
132
147
99
1.11
0.75
0.40

Golden Eagle Nests - 1977

Known productivity and fledging success was av?ilab1e for 70 sites, while
only productivity information was available for 27 additional sites.
Due
to extremely low mortality between production and fledging, no young were
assumed to have died in the adjusted figures below.

Successful sites
Total Young Produced
Total Young Fledged
Young Produced/Successful
Nest
Young Fledged/Successful
Nest

Known Values

Adjusted

Values

70
102
101
1. 46
1. 44

70 + 27
102 + 42
101 + 42
1.48
1. 47

97
144
143

�-86-

Bald Eagle Nesting

Results

As in 1976, the two bald eagle nests produced 2 young at each site. No
attempt was made to band or colormark the young at the time. We are proceeding cautiously in monitoring these eagles in order to avoid disturbance which might cause abandonment of the sites.

Prepared

by

G,~ R. ~~

Gerald R. Craig
Sr. Wildlife Biologist

�January

-87-

1979

JOB PROGRESS REPORT

State of ----~~~~~------------COLORADO
Project NQ. -----------------------W-124-R
Work Plan No.
Job Title

Job No.

~2~

Bald and Golden Eagle Winter Population

Period Covered:
Personnel:

I

Raptor Investigations

April 15, 1977 through February

_
Surveys

28, 1978

Erwin Boeker and Alan Jenkins, U. S. Fish and Wildlife Service;
Gerald Craig, Joe Frothingham, Ann Potter and Gordon Saville,
Colorado Division of Wildlife; C. Eugene Knoder, National Audubon Society; and Wayne Russell, Colorado Division of Wildlife

ABSTRACT
Midwinter golden and bald eagle flights were continued on census areas in
the San Luis Valley, northeastern and northwestern Colorado.
Inclement
weather conditions interferred with the northwest golden eagle flightc?:D-SL
forced cancellation of the Yampa River bald eagle flight.
An experimental
census was conducted in southwest Colorado to test the feasibility of expanding
the work into that region.
The total number of golden eagles continued to decline in the San Luis Valley
and northwest and increased slightly in the northeast.
All areas were typified
by low numbers of juveniles which represents poor reproduction.
The total
number of bald eagles in all areas continued to decline.

��-89-

BALD AND GOLDEN EAGLE WINTER POPULATION

SURVEYS

Gerald R. Craig

Golden eagle winter population trend information is obtained by annually
censusing sample areas and recording the number of eagles observed per 100
square miles.
The censuses yield area estimates which are of use in extrapolating probable eagle populations throughout the region.
The aerial census flights are designed to be compatible with similar study areas in other
western states.
The U. S. Fish and Wildlife Service then compares data
from states throughout the region to obtain population information for golden eagles throughout the western United States.
Bald eagle population information is obtained by censusing concentration
areas, primarily river courses throughout the state. As with golden eagle
trend counts, aerial flights are made in midwinter when the population reaches
its peak.
Rather than an area estimate, all bald eagles sighted are recorded
since the census tends to be linear in fashion.
The censuses also yield valuable information about those areas which are preferred by wintering bald
eagles.

P.N.

OBJECTIVES

The objective of this study is to obtain winter population trend information
for bald and golden eagles on selected wintering areas in Colorado.
The
information will be used to estimate wintering populations of bald and golden eagles throughout the state.

SEGMENT OBJECTIVES
1.

Aerial

Counts of Wintering

Golden Eagles:

Once annually an aerial flight will be made on census areas in the San
Luis Valley, northeastern and northwestern portions of the state.
These census areas were established in 1972 and the procedures will be
essentially the same. Random transects will be flown throughout each
study area with a Cessna 185 aircraft and all eagles observed within
~ mile of each side of the transects will be counted and classified as
adult or juvenile.
From the transects, an area estimate will be obtained
as to eagles per 100 square miles.
Identical census areas are also
flown in Wyoming, Idaho, Montana, North Dakota, New Mexico and Nevada
by the U. S. Fish and Wildlife Service and cooperating agencies.
Information forthcoming from these states are then collected and analyzed by
the Fish and Wildlife Service to obtain population estimates for the
West. Age ratio information which is obtained provides an indication
of the previous breeding season's reproduction.

�-90-

2.

Aerial Counts of Wintering

Bald Eagles:

Since bald eagles tend to congregate primarily along river courses
and impoundments, aerial flights will be made along major river
courses throughout the state and a direct count will be rnade of all
bald eagles observed.
The eagles will be classified as to age in
order to obtain information about reproduction.
The flights wf.Ll, be
made in January when the highest concentration of eagles are usually
present.
Flights will be made along the South Platte River, Yampa River,
White River and Rio Grande River.
It is proposed to expand the censuses
to the Colorado River and possibly the Gunnison, Dolores and San Juan
Rivers.
3.

Compile data and prepare annual progress and final reports and submit
them to appropriate personnel and agencies.

METHODS

AND MATERIALS

Midwinter golden eagle flights were conducted in the census areas in the San
Luis Valley, northeastern and northwestern Colorado according to procedures
described in Segment Objective 1. The northeastern census area constitutes
approximately 3,000 square miles in Weld and Logan Counties.
North-south transects totaling approximately 600 miles were established randomly throughout the
area. Observers count all eagles observed within ~ mile of either side of the
aircraft, so this accounts for actual coverage of 300 square miles or 10% of
the total census area. The transects are flown at speeds averaging 100-120 mph
at altitudes of 100 to 300 feet, depending upon the topography.
Census areas
in the San Luis Valley and northwest portion of the state are flown in a similar manner.
Transects account for a 10% sample of the San Luis Valley.
The
area encompasses 2,500 square miles which is the majority of the valley lands
of the San Luis Valley.
The northwestern census area includes Moffat and Rio
Blanco Counties and is the largest with 4,100 square miles.
Due to extended
flight time, the number of transects were reduced so that 7% of the area is
covered.
Flights were conducted on the following dates: northeastern Colorado,
January 4, 1978; San Luis Valley, January 14, 1978; and northwestern Colorado,
January 19, 1978.
Winter bald eagle counts are conducted in the San Luis Valley in conjunction
with golden eagle flights mentioned above.
Since bald eagles are generally
distributed throughout the San Luis Valley, it is possible to obtain an
effective area estimate of bald eagles during the same flight for golden
eagles.
Bald eagles are also censuses in conjunction with golden eagle flights
in northwestern Colorado, but the effectiveness of this method has declined
over the past six years and will probably be discontinued.
To compensate,
direct counts are being made along the Yampa River from Craig to Dinosaur
National Monument.
Unfortunately, inclement weather in 1978 forced termination
of the flight before the census was complete.
A direct count is also made of
the South Platte River upon completion of the golden eagle flight of northeastern Colorado.
The course of the river is flown at altitudes of 100 to 200
feet and all eagles are counted along the river from Fort Morgan to Greeley.
In an effort to expand the censuses to southwestern Colorado, an experimental
flight was made of major drainages in February, 1978. The following river
drainages were flown:
San Juan, Piedra, Los Pinos, Animas, Florida, La Plata
and Dolores.
I

�-91-

RESULTS AND DISCUSSION
Tables 1 and 2 summarize
eagles.

the results of aerial

flights

for bald and golden

In northeastern Colorado, the total number of golden eagles increased slightly
from 1977, but a general decline is still evident.
The extremely low percentage of immature eagles observed indicates that the population is still
experiencing low reproduction.
The decline of golden eagles in northwest
Colorado has continued since 1972 and 1973 when rabbit populations were high.
Poor reproduction is also indicated by the low number of juveniles in the
population.
The general population decline was also indicated in the southern
portion of the state on the San Luis Valley census area. Tentatively, it
appears that reproduction may be up slightly in this population.
Extremely low numbers of bald eagles have been observed in recent years on the
northwest Colorado flights.
Despite the slight increase from 1 to 4 bald
eagles from 1977 to 1978, insufficient numbers are present to warrant use of
the estimates.
The high rabbit population in the early 1970's may have served
to attract large numbers of bald eagles away from the riparian areas they
normally frequent.
A slight increase was evident in the San Luis Valley bald
eagle population, but the figures are well below the high observed in 1976 when
the flights were initiated.
Along the South Platte, bald eagle numbers have
also continued to decline.
A significant increase in percent of juveniles in
the population (68%) indicates a decline in the adult segment and not an increase in immatures.
A three-hour flight of major drainages and impoundments in southwestern Colorado yielded a total of 41 bald eagle observations.
It is recommended that
an aerial flight be established in that region to assess future population
trends.
Table 1.

Golden Eagle Aerial Census - 1972-1978

Northeastern Colorado - 1972-1978
(10% sample of 3,000 sq. mi.)

Date
1/24/1973
1/16/1974
1/22/1975
2/19/1976
1/13/1977
1/4/1978

Adults
16
19
22
17
18
18

Juveniles

Unknown

Eagles per
100 sq.mi.

Est. of Total
Eagles

8
3
5
3
1
3

0
0
0
0
0
0

8.0
7.3
9.0
6.7
6.3
7.0

240
220
270
200
190
210

�-92-

Table 1 (continued)
Northwestern Colorado - 1972-1978
(7% sample of 4,100 sq. mi.)

Date
1/25 &amp; 2/26,1972
1/23/1973
1/22 &amp; 1/29,1974
1/29/1975
1/26/1976
1/19/1977
1/19/1978

Adults

Juveniles

Unknown

Eagles per
100 sq.mi.

Est. of Total
Eagles

86
35
6
11
29
29
23

80
14
8
8
3
5
5

97
70
47
17
10
2
0

91.6
41.5
21.2
12.5
14.6
12.5
9.8

3,757
1,700
871
514
600
514
400

San Luis Valley, Colorado - 1976-1978
(10% sample of 2,500 sq. mi.)

Date
1/29/1976
2/16/1977
1/14/1978

Table 2.

Adults

Juveniles

17
14
10

9
3
5

Eagles per
Unknown Total 100 sq.mi.
2
1
0

28
18
15

11.2
7.2
6.0

Est. of Total
Eagles
280
180
150

Bald Eagle Aerial Census - 1972-1978

South Platte River, Colorado - 1972-1978
(exact count)
Date

Adults

Juveniles

Unknown

Total

% Juveniles

1/24/1973
18
l3
0
31
42%
1/16/1974
16
15
0
31
48%
1/22/1975
28
14
0
42
33%
2/19/1976
19
10
0
29
35%
-------------The South Platte River was not flown this year-------------1/4/1978
8
17
0
25
68%

�-93-

Northwestern Colorado - 1972-1978
(7% sample of 4,100 sq. mi.)

Date

Adults

Juveniles

Total

*
*
*0

35
4
5
1
11
1
4

1/25 &amp; 2/26/72 ,~
1/23/1973
*
1/22 &amp; 29/1974 *
1/29/1975
1
1/26/1976
7
1/19/1977
1
1/19/1978
4

4
0
0

Eagles per Est. of Total
100 sq. mi.
Eagles
12.2
1.4
1.7
0.4
3.8
0.4
1.4

500
57
71
14
157
14
57

*No distinction was made between adults and juveniles on these flights

San Luis Valley, Colorado - 1976-1978
(10% sample of 2,500 sq. mi.)

Date

Adults

Juveniles

Total

1/29/1976
2/16/1977
1/14/1978

28
12
18

12
6
8

40
18
26

Eagles per Est. of Total
100 sq. mi.
Eagles
16.0
7.2
10.4

400
180
260

Yampa River, Colorado - 1977-1978
(exact count)

Date
1/19/1977
1/19/1978

Adults

Juveniles

Total

% Juveniles

14
17
17.6%
3
-----------cancelled due to inclement weather--------------

Southwestern Colorado - 1978
(exact count of San Juan, Piedra, Los Pinos, Animas, Florida, La Plata and
Dolores Rivers and associated reservoirs)

Date

Adults

Juveniles

Total

% Juveniles

2/ /1978

32

9

41

22%

R.

e~

Prepared by ~
Gerald R. Craig,

. Wildlife Biologist

��January

1979

-95-

JOB PROGRESS REPORT

State of

COLORADO
----~-------------------

Project No.

Raptor Investigations

W-124-R

Work Plan No. ~~--------------II
Job Title

Osprey Nesting

Job No.

Studies

Period Covered: April 15, 1977 through February
Personnel:

1

28, 1978

Terry Becker, Gerald Craig, Steve Porter, Mike Stiehl,
John Wagner, Colorado Division of Wildlife

ABSTRACT

Extremely poor reproduction continued to plague the intermountain osprey
population in Colorado.
A single young was fledged from only one of 13
occupied nests in 1977.

��-97-

OSPREY NESTING

INVESTIGATIONS

Gerald R. Craig

Although migrant osprey are regularly sighted in Colorado, only two unsatisfactory nesting records are reported in the literature.
In the late 1960's
R. A. Ryder reported a pair of osprey nesting on Grand Lake in Grand County
and another pair were observed breeding successfully at Electra Lake in
La Plata County.
Since initiation of a breeding survey in 1972, the number
of known osprey nests have expanded to 18, most of which are located in Grand,
Jackson and Larimer Counties.
The present investigation is designed to determine nest site requirements and monitor breeding success.

P.N.

OBJECTIVES

The objectives of this study are: (1) to monitor productivity of nesting
osprey, (2) to identify nesting habitat requirements and implement measures
to encourage occupancy by additional pairs, (3) to compile data and submit
reports to appropriate state and federal agencies to assist them in delineating
and protecting key nest sites.
.

SEGMENT OBJECTIVES
la.

Continue

to locate and map nesting

sites of osprey throughout

Colorado.

lb.

All known nest sites will be visited annually to establish reproductive
success.
All unhatched eggs will be collected and analyzed to ascertain the cause of hatching failure.

2a.

Habitat features such as key hunting areas, topography, vegetative
type, climate, and geology will be recorded for each nest site in
an attempt to establish habitat requirements.

2b.

Artificial nests will be constructed in localities which offer potential for occupancy by osprey.
The sites will be monitored annually
to determine the success of the endeavor.

3.

Analyze all data obtained

and prepare a report of the findings.

METHODS ~~D ~~TERIALS
Due to scheduling difficulties, most nests were not investigated until late
July and August.
All recorded nests were visited to determine production,
band young and collect egg fragments.
In addition, the presence and activity
of adults were recorded.
A fixed-wing flight was scheduled to check known
nest sites and locate additional nesting areas.

�-98-

RESULTS AND DISCUSSION
Only one nest succeeded in fledging young in 1977 out of a total of 13 sites
which were occupied by breeding osprey.
A second nest (site LAl) may have
fledged young prior to nest checks of August 10 and 11, but the young were
not visible in the vicinity.
Table 1 summarizes the results.

Table 1.

Osprey nesting

Site

Activity

GRI
GR2
GR3

success in Colorado,

1977.

Productivity

Date Checked

Adults present
Adults present
Adults present

0
0
0

7/7/77, 8/2/77
8/1/77
8/1/77

GR4~'&lt;

Adults present

0

8/2/77

GRS

Adults present

Unknown

7/31/77

GR6
GR7*
GR8*
GR9

Inactive
Adults present
Inactive
Adults present

GRlO*
JAI
JA2
JA3
JA4

Adults present
Adults present
Inactive
--------------Nest
Adults present

JAS
JA6
LAI
LA2

Adults present
Inactive
Adults present
Adults present

o
o

o

7/31/77
7/?/77, 8/1/77
8/2/77
8/1/77

Comments

Egg fragments below
nest
Adults building onto
nest
Could not climb nest,
adults not defensive.
Probably no young.
Nest destroyed by wind
Nest destroyed by wind
Adult in incubating
posture

8/1/77
7/15/77, 8/10/77
8/10/77
Nest tree blown down
and tree could not be located from ground or air---o
8/17/77, 8/29/77 Adult in incubating
posture
o
.8/18/77
8/20/77
Unknown
8/10/77, 8/11/77
o
8/10/77, 8/15/77 Nest in poor repair
2

This population continues to exhibit extremely low reproduction which is not
sufficient to maintain itself.
Drought conditions in the summer of 1977
caused extreme drawdown of the reservoirs in the Shadow Mountain-Grand Lake
complex.
Low water levels increased accessibility to some nests and mayor
may not have reduced fishing opportunity which caused incubating adults to
abandon their eggs to forage for themselves.

Prepared

by

C,11M~

Q ~

G~ardR: cr~ig
Sr. Wildlife

Biologist

�January

~99-

JOB PROGRESS

State

REPORT

COLORADO

of

Project

----------------------No.

Raptor

W-124-R

--------------------

Job No.

1
--------------------------------

No.

Job Title

--------------------------------------------------------------------

-----------------

Prairie

Covered:

Personnel:

III

Investigations

Work Plan

Period

1979

Falcon Nesting

April

Studies

15, 1977 through

February

28, 1978

Terry Becher, Gerald Craig, James McKinley
Colorado Division of Wildlife.

and Michael

Stiehl,

ABSTRACT
Statewide prairie falcon nesting studies indicate a reproduction of 3.24
young per successful pair which is down slightly from the previous year.
The effectiveness of this portion of the study was reduced because of a
delay in initiating field investigations.
The intensive study area in
northeastern Colorado yielded good occupancy of nest sites and slightly
higher reproduction.

��-101-

PRAIRIE

FALCON

NESTING

Gerald

R. Craig

STUDIES

In Colorado, the pralrle falcon ranks high in popularity for falconry and
demand will undoubtedly increase since this is the only large falcon which
can legally be removed from the wild.
As harvest pressures increase, it
will be necessary to monitor the populations to assure that harvest is not
detrimental to the well being of the population.
Preliminary investigations
indicate that the number of occupied nests have remained fairly stable over
the past decade.
However, detailed investigations
are required to establish
distribution and productivity on a statewide basis.

P. N. OBJECTIVE
The objectives

of this study are:

1.

To document the breeding range and estimate
of prairie falcons in Colorado.

2.

To obtain production data at selected nesting areas
and estimate total production of prairie falcons.

throughout

3.

To delineate

nesting

habitat

falcons.

4.

To document
Colorado.

movements

and mortality

requirements

SEGMENT

the number

of prairie

of prairie

falcons

of breeding

pairs

the state

throughout

OBJECTIVES

1.

Locate and map all known nesting sites of prairie falcons throughout
Colorado.
From data obtained through approach 3, extrapolate the number
of breeding pairs potentially occupying similar habitat types throughout
the state.

2.

Establish study areas in select habitats which represent important nesting
areas.
Study areas will be selected that represent shortgrass prairie,
foothills and mountain nesting populations and productivity will be
determined and compared between areas.

3.

Physical and biological parameters of each nest site which is visited
will be recorded on appropriate field forms and analyzed to establish
those features which favor occupancy by prairie falcons.
The field
information will be gathered in conjunction with approach 1.

4.

When nests are visited to determine productivity,
the young will be
banded with Fish and Wildlife Service lock-on bands and their movements
will subsequently be traced through reports filed with the Office of
Migratory Bird Management.
Should the occasion permit, transmitters
will be placed on several breeding adults to determine extent of hunting
ranges.
Radio transmitters will also be placed upon young at fledging
and their movements and activities will be monitored.

5.

Compile

data and prepare

annual

and final reports.

�-102-

METHODS

AND MATERIALS

Known nest sites were visited during courtship and early incubation to determine the presence of breeding adults.
Invariably all sites could not be
visited early in the season and some only received visitation once after the
young were produced.
Of those sites which were checked, a sample of active
nests was selected and an attempt was made to return to each of the sample
sites and determine reproductive success.
Young were banded with U.S. Fish
and Wildlife Service bands when nests were visited to determine brood size,
locate prey remains, and record other information about the nest.
In addition, the following information was recorded on field forms:
elevation,
topography, geology of the nest cliff, major vegetative communities, potential hunting areas, distance and direction to disturbance factors, location
of nest site, height from nest to top and bottom of the cliff, dimension of
the nest ledge, and presence of other raptors in the vicinity.

RESULTS

AND DISCUSSION

Statewide nesting studies were hampered by inability to employ field assistants that were experienced in locating and observing nesting prairie falcons.
As a result, much of the early nesting information was not gathered in 1977.
In particular, the number of active nest sites may be exaggerated since some
of the sites which were classed as inactive may have been occupied earlier in
the season.
A sample of sites were visited in late May and June and reproduction was obtained for those nests.
Although the percent of active sites
was reduced (94% in 1976 to 73.2% in 1977), the number of young produced
per successful pair was reduced only slightly (from 3.37 to 3.24).
Following
is a synopsis of the statewide reproduction:
Total number of nests checked
Number of active nests
Number of successful nests
Number of young produced
Percent of sites which were active
Young produced per successful pair
Young produced per active nest

82

60
50
162
73.2%
3.24
2.7

The intensive study area in northeastern Colorado continued to exhibit good
reproduction.
Thirty-five sites were occupied by breeding pairs for an
85.4% occupancy rate.
An average of 4.44 eggs were laid, 4.48 young were
produced per successful nest and 3.64 young fledged per successful nest
(see Table 1).

�-103-

Table 1.

Prairie

falcon reproductive

success

in northeast

Colorado,

1977.

Site

Nest
occupancy

Number
of eggs

Number
of young

Number
fledged

WE 1

Active

5

5

4

WE 2

"

5

5

5

\.JE3

"

5

5

5

WE 4

"

4

4

o

WE 5

Inactive

WE 6

Active

5

3

3

WE 7

4

4

o

Female killed

5

4

2

2 young died of tick infestation

WE 9

"
"
"

5

3

o

All young died of tick infestatior

WElOA

Inactive

WEllA

Active

3

3

3

WE12

Inactive

Great Horned Owl using site

WE13

Active

Single adult male present

WE14A

"

4

4

4

WE14B

"

4

1

1

WE16A

"

6

6

6

WE17

Inactive

WE18

Active

5

1

1

WE19

5

5

o

WE20

"
"

WE2l

"

4

4

4

WE22

4

4

4

WE23

"
"

4

3

3

WE24

"

5

5

5

l.JE25

.,

WE30

Inactive

WE32

Active

1

o

o

WE34

"

4

4

4

WE35

5

5

4

WE37

"
"

4

3

3

WE38

"

o

o

o

WE40

"

?

2

2

WE 8

Comments
5th young removed

for falconry

Adult shot

Great Horned Owl using site

Yound died of tick infestation
Single adult male present

Single adult female present

5th young removed

for falconry

�-104-

Table l.

Prairie

falcon reproductive

success in northeast

Colorado,

1977 . (Cont.)

Site

Nest
occupancy

Number
of eggs

Number
of young

Number
fledged

WE4l

Active

4

3

3

\-lE42

"

5

5

5

WE43

?

4

4

WE44

"
"

5

4

4

WE45

"

?

4

4

WE47

"

5

5

5

WE48

"

0

0

0

Pair present

WE49

"

5

4

3

4th young removed for falconry

WE50

Inactive

Great Horned Owl using site

Totals

120

112

91

Percent

of sites which were active

85.4%

Average

clutch size

= 4.44

per successful

pair

= 4.48

Young fledged per successful

pair

= 3.641/

Young fledged per active site

= 2.60

Young produced

= 3.20

Young produced

Comments

per active site

1/ Fledging success is raised to 3.72 if the 3 young are included which were
taken for falconry.

Prepared

j'J
fJ Ill).;?
.
by__~~~~~~~~~~
__ ~~~~~~Yr
Gerald R. Craig
\j
Sr. Wildlife Biologist

_

�January

1979

-105-

JOB PROGRESS REPORT

State of

Colorado

----------.-------------

Project No. W-124-R
Work Plan No.
Job Title

III

Raptor Investigations
Job No.

II

Prairie Falcon Breeding Population

Period Covered:

Characteristics

Studies

April 15, 1977 through February 28, 1978

Personnel:
Gerald Craig, and James McKinley,
Steve Platte, Brigham Young University.

Colorado Division

of Wildlife;

ABSTRACT
The total number of marked adult prairie falcons has been expanded to 41 for
the study area. Between the 1976 and 1977 breeding season, 6 marked adults
failed to return which gives an estimated mortality of 30%. Age structure
of the population is developing as ages are established from previously
banded adults. Ages of 11 falcons indicates an average age of 3.91 years
which provides a corresponding annual mortality of 26 percent.

��-107-

PRAIRIE FALCON BREEDING
POPULATION CHARACTERISTICS STUDIES
Gerald R. Craig and Steve Platte

Before any management activities or protective measures can be implemented,
it is necessary to understand population dynamics of the target species.
Prairie falcon life tables have been developed from Fish and Wildlife Service
band returns, but the calculated adult mortality is generally believed to be
unrealistic due to inconsistencies caused by illegal shooting.
Also, a large
number of falcons must be banded to yield sufficient returns to permit analysis.
An alternative method is being investigated which involves the monitoring of a
population of marked prairie falcons.
A disjunct prairie falcon population in
northeast Colorado will be trapped, banded and color marked at breeding sites,
then will be monitored annually to determine the loss of breeding adults from
the population.
Annual adult mortality can then be determined by direct
observation of the sample population.
P. N. OBJECTIVE
The objective of this study is to document nest site fidelity
of members of breeding pairs of prairie falcons.

and replacement

SEGMENT OBJECTIVES
1.

A study area in Northeastern Colorado will be delineated which encompasses
approximately 20 pairs of breeding prairie falcons.
The area is physically
isolated from the nearest breeding pairs by 10 to 15 miles, thus it will
be easier to monitor recruitment and replacement of adults within the
population.

2.

Attempts will be made to trap all adults in the study area, place Fish
and Wildlife Service bands and colored leg streamers on them to identify
individuals.
If some falcons are already banded, their age and banding
location will be obtained from banding records.

3.

Vocalization of individuals will be recorded and subjected to spectrographic analysis with the intent of distinguishing adults without
capturing and handling them. The technique will first be tested upon
identified individuals to prove its effectiveness.

4.

The study area will be revisited annually and loss of mates or shifts
in breeding pairs will be determined by observing colormarked individuals,
recording and comparing vocalizations and trapping falcons and noting
banding numbers.
Any unmarked adults will be trapped, banded and colormarked.

5.

Analyze

the data and prepare

annual progress

reports

and a final report.

�-108-

METHODS AND MATERIALS
Prairie falcon nests were visited in May and June and attempts were made
to capture the adults by using various techniques.
Care was taken to avoid
prolonged disturbance of the adults while they were incubating eggs. Those
adults that were captured were color marked with short nylon streamers on
either tarsus.
Streamers were placed on alternate legs of pairs which
occupied adjacent nests in order to distinguish them should a switch of mates
occur. u.S. Fish and Wildlife Service bands were also placed on the other
tarsus of captured falcons to provide certain identification.
While the birds
were in hand various measurements were taken in an effort to develop techniques
to distinguish individual falcons.
On subsequent visits to the nests,
vocalization of each adult was recorded with a portable tape recorder and
parabolic microphone.
The tape recordings were later analyzed spectrographically
in an attempt to determine the feasibility of identifying the vocalizations
of individuals.
On succeeding years, each site was revisited to determine replacement of adults
and possible exchange of mates with other sites. The leg streamers permitted
partial identification, but most of the adults had to be recaptured to check
band numbers.
Previously unmarked adults were marked with streamers and bands
as described above. Additional tape recordings were taken on subsequent years
to compare vocalizations of the same individual which were recorded on previous
years.
Description

of Study Area

The study area includes the Chalk Bluffs and Pine Bluffs in Wild County, Colorado.
The habitat is typical short grass prairie interspersed with sandstone outcroppings.
A relict population of timber pine is present in scattered locations
on to Pine Bluffs and a few cottonwoods are present along intermittent stream
courses.
RESULTS AND DISCUSSION
The first action was to determine how many of the 20 prairie falcons marked
last year survived one year and returned to the study area. The color leg
markers were easy to detect and 14 falcons that were marked were found. The
recruitment standard is the number of young per pair that need to be produced
to maintain a stable population.
The recruitment standard associated with 70
percent survivorship (1976-77 estimate) falls in the 2.86 to 4.29 young per
pair range. The 1977-78 sample (Table I) includes 41 marked adult prairie
falcons (14 from 1976 and 27 from 1977).
Two eyries were inactive where single adults had been marked the previous year.
Single marked falcons were replaced at two locations.
One of the replacement falcons
was one year old (molting out of immature plumage) and the other was two years
old (already banded when trapped).
Two marked falcons (a pair) were both replaced at one eyrie and one of them, an adult female, was trapped and marked.

�-109-

Table I. Possible recruitment
priarie falcon survivorship.

No. Return

standards

Ad. Survivorship

considering

variation

in immature

Recruitment Standard
Immature Surv. Rates
.30
.25
.20

40

.98

0.17

0.20

0.25

39

.95

0.34

0.41

0.51

38

.93

0.53

0.63

0.79

37

.90

0.72

0.86

1.08

36

.88

0.93

1.11

1.39

35

.85

1.14

1.37

1.71

34

.83

1.37

1.65

2.06

33

.80

1.62

1.94

2.42

32

.78

1.88

2.25

2.81

31

.76

2.15

2.58

3.25

30

.73

2.44

2.93

3.67

29

.71

2.76

3.31

4.14

28

.68

3.10

3.71

4.64

27

.66

3.46

4.15

5.19

26

.63

3.85

4.62

5.77

25

.61

4.27

5.12

6.40

The expanded study included 35 eyries as compared to 28 in 1976. Eighteen
sites were active and 10 were inactive.
Fifteen of the 18 active sites were
active in 1977 (two changed single adults and one changed both adults) and
three were inactive (all had single adults present).
Eight of the 10 sites
which were inactive in 1977 were active in 1977 (three had immature females
and one had a two year old female) and two were inactive (one had a single
adult and the other had nothing).
In 1976 35 percent of the sites were
inactive and in 1977 only 17 percent were inactive.
Thirty-one active pairs
of falcons were observed this year.
They hatched 112 young (3.61/occupied
site) and fledged 92 (2.97/occupied site).
Two observers banded approximately
110 nestling and 27 adult prairie falcons.
Seven of the adult falcons were
already banded when trapped.
The six adult falcons already banded when trapped
last year all survived.
The oldest bird in the two groups is six years old,
however, the age of several are still on request from the Bird Banding Lab,
U.S. Fish and Wildlife Service.
The age of 11 of 13 already banded falcons
is known.
The average age of these birds is 3.91 years (range 2 to 6). If
we consider these birds a "sample" then the corresponding average annual

�-1l0-

mortality rate is 26 percent (1.0/3.91, range in recruitment standards of
2.29-3.44).
This year's sample indicates 30 percent per year and Enderson
(1969) estimates 23 percent per year. Next years sample (41 marked birds)
should be sufficiently large enough to serve as a good estimator of adult
mortality because we appear to be in close agreement with other estimates
available.
Six of the females trapped were molting out of immature plumage, indicating
they were one year old. All six falcons successfully bred and fledged
young.
Tape recordings of 20 individuals
graphic analysis.

(9 banded)

LITERATURE

are available

for spectro-

CITED

Enderson, J. H. 1969. Peregrine and Prairie Falcon life tables based
on band-recovery data, pp. 505-509.
In Peregrine falcon populations,
J. J. Hickey, Editor, Univ. Wisconsin Press, Madison, 596p.

Prepared

r&gt;.

Ii /)

n.

/I

.

by __~lQ.~~~~~~V(~~~~~~:~~
Gerald R. Craig
\J
Sr. Wildlife Biologist

_

�January

-111-

JOB PROGRESS

State

Work

No.

W-124-R

-------------------

Pl~n No.

Job Title
Period

REPORT

COLORADO

of

Project

1979

Job No.

Investigations
1

---------------------------------------------------

Covered:

Personnel:

IV

Raptor

April

15, 1977 through

February

28, 1978

William Andersen, Otero Junior College; Gerald
McKinley, Colorado Division of Wildlife.

Craig

and James

ABST~~CT
Ferruginous hawk productivity was obtained from two study areas in southeast
Colorado and another in northeast Colorado.
Nests on one of the areas in
southeast Colorado were manipulated where necessary to improve stability
or suitability as nest sites.
Approximately
2.03 young were produced per
nesting attempt on all study areas.
The managed area, however, produced
3.1 young per nesting attempt while the unmanaged areas produced 1.8 and
1.6 young per nesting attempt.
The primary cause of nesting failure are
high winds which capsize the bulky nest structures.

��-113-

FERRUGINOUS
Gerald

R. Craig

HAWK NESTING

STUDIES

and William

C. Andersen

Ferruginous hawks are strongly associated with the shortgrass pralrle community of eastern Colorado, although they do occur throughout the state in
limited numbers.
Their populations has undoubtedly declined within the past
century as large portions of their habitat have been converted to agricultural
lands.
Presently, approximately
100 nests have been located, the majority of
which are on or adjacent to the Pawnee and Comanche National Grasslands.
Ferruginous hawks are extremely sensitive to human encroachment and habitat
degradation and additional investigations are underway to establish population trends, tolerance limits and develop techniques to enhance reproduction.

P.N. OBJECTIVE
The objectives

of this study are:

1.

To document the breeding
eastern Colorado.

numbers

2.

To delineate habitat parameters and disturbances
ferruginous hawks in eastern Colorado.

3.

To evaluate potential techniques to enhance nesting
expansion of ferruginous hawk breeding populations.

SEGMENT

and productivity

of ferruginous

impacting

pairs

hawks

in

breeding

or encourage

OBJECTIVES

1a.

Locate, map and photograph all known nest sites of ferruginous
the eastern Colorado plains.

hawks

on

lb.

Known nest sites will be observed from a distance to establish the presence of courting or incubating adults.
Occupied sites will be revisited
prior to fledging and the young will be counted, banded, and an attempt
will be made to identify all prey items present.

2.

Habitat features such as vegetative type, type of nesting structure, topography, soil type, climate, vicinity of human habitation, roads, and other
disturbances will be recorded for those nests identified in la and lb.
The physical and biological features will be evaluated to establish those
parameters which favor successful nesting of ferruginous hawks.

3a.

A sample of a predetermined number of nests will be stabilized with wire
baskets in an attempt to enhance nesting success.
Production of the
manipulated nests later will be compared with unaltered sites to determine the effectiveness of the efforts.

3b.

Between one and two dozen artificial nest structures will be placed in
suitable habitats which are not occupied by breeding pairs.
All breeding
pairs adjacent to each treatment area will be located prior to placement

�-114-

of the nest structures and will be monitored after placement of the
structures to be certain that the structures actually encourage
pioneering by new pairs and do not cause relocation of adjacent pairs.
4.

Analyze

the data and prepare annual progress reports and a final report.
METHODS AND MATERIALS

Four areas of primary investigation were established on the eastern plains.
Areas 1 and 2 encompassed the southern and northern portions of the Comanche
National Grasslands, respectively, in southeastern Colorado.
Area 3 included
the eastern and western portions of the Pawnee National Grasslands, respectively, in northeastern Colorado.
Ferruginous hawks have proven to be prone to abandonment if their nests are
visited during the courtship and incubation periods, but they will tolerate
visitation after the young have hatched.
This behavior mandated that the
study be carried out in three phases:
Phase 1. Nests within all study areas were checked in April and May to
determine the presence of courting or incubating adults.
Observations were
made from distances of at least one half mile (.83 km) from a vehicle and
observers remained stationary for a duration of no more than ten seconds.
Activity and nest locations were recorded after departure from the area.
No other data was taken at the time.
Phase 2. Nests were revisited between June 1 and June 25 when the young were
old enough to band but not developed enough to attempt to fledge prematurely.
At that time, young were banded with U.S. Fish and Wildlife Service lock-on
type bands.
Sizes 7B and 7D were placed upon the tarsus of each young.
Data
collected during this visit included:
age and general health of young, color
phases of young and adults, lining material in nests, and prey items present
in and below nests.
Pellets present in and below the nests were collected
and catalogued for analysis.
No habitat data was collected during this phase
to reduce the length of stay at each nest to an absolute minimum.
Phase 2
began in Area 1 and proceeded sequentially through Areas 2 and 3 since previous studies have shown that nesting activities in the north tend to lag
behind southern areas by five to seven days. At the conclusion of Phase 2,
all study areas were rechecked for previously unknown nests of renesting pairs.
Phase 3. Final visits were made to the areas from early through mid-August
after the young had fledged and were no longer in vicinity of the nests. At
that time, all the remaining data was collected such as nest site measurements,
habitat information, disturbance factors, and maps were drawn.
In addition,
modifications were made at selected sites to improve their stability or make
them more suitable for future occupancy.

�-1l5-

Modification

or management

Designation

Techni

O.

Nest was not altered

1.

techniques

used on nests

are as follows:

ue
in any way.

Top the nes t. This required removal of several previous years
nest material to reduce the bulk of the nest and thereby improve
its stability in strong winds.

2.

Remove restricting branches. Tree growth and/or annual addition
of nesting material cause obstructions which make it difficult
for adults to approach or depart the nest.
The restricting
branches may also reduce the surface area of some nests.

3.

Shift nest center.
As additional "pads" of nesting material are
added annually, the nest may begin to lean in one direction until
it eventually topples over.
The nest is realigned over a vertical
axis to improve support.

4.

Widen nest.
As nesting material accumulates, some nests become
cone shaped and the nest platform is reduced.
In other instances,
the supporting tree limbs and trunk did not permit the hawks to
construct a wide nest platform.

5.

Reinforce base.
Wire netting and other materials are used to
"shore up" nest bases that have degenerated or lack adequate
support for the weight or bulk of the nests.

6.

Provide artificial base.
In some cases it is best to remove
the old nest entirely, construct an artificial base of welded
wire and replace a portion of the nest on the new base.

7.

Move the nest to new site.
Where human disturbance repeatedly
causes nesting failure, or the nest tree is an inadequate
support structure, extreme action must be taken to move the
entire nest to a superior location.

8.

Provide predator protection.
In extremely rare circumstances,
mammalian predators (such as packrats) may endanger the eggs or
young.
Depending upon the type of predation, different solutions
are used.
A fairly effective measure is to nail tin sheathing
around the tree trunk to make it unclimbable.

9.

Provide alternate nest.
This approach is less extreme than 7
and consists of constructing an artificial nest in the vicinity
of the existing nest.
The adults can then select the superior
site, or renest at the new site should their original nest be
destroyed.
This is generally a good insurancp policy for each
site where possible.

�-116-

RESULTS

AND DISCUSSION

Ferruginous hawks frequently construct large, bulky nests in solitary deciduous trees on the prairie.
Generally, the trees are not occupying optimal
growing conditions and are stunted.
The combination of poor supporting
structure and large area appear to make many nests susceptable to high winds.
Previous investigations
tended to bear out this hypothesis, so in 1976 those
sites in Area 1 were modified using the above mentioned techniques where
necessary to improve nest stability.
None of the nests in Area 2 or 3
received this treatment prior to the 1977 field season, so it was possible
to compare nesting successes of the three study areas.
As can be seen in Table 1, Area 1 experience 100 percent nesting success
while Areas 2 and 3 encountered nesting successes of 64 and 52 percent,
respectively.
The nesting successes in Areas 2 and 3 appear to be representative of reproductions which was normally encountered on previous years.
As would be expected, the number of young produced per nesting attempt was
also much greater for Area 1 than the two unmanaged areas.
The young per
successful nests did not fluctuate much among the three areas and really
does not enter into consideration
since itfu more a function of biological
and physiological
factors such as prey availability and fecundity.
During Phase 3 in 1977, eight of 29 nests in Area 2 received treatment to
improve their qualities and 18 of 29 nests were improved in Area 3. The
three study areas will be monitored again in 1978 to determine the final
results of this proposed management action.

�-117-

Table 1.

Ferruginous

hawk productivity

and nest site management.

Study Area 1
Site
no.

Young
produced

Management /
1
tech. usecF-

Manage. tech.
still required

FH-13(CS)

3

1

0

FH-14(CS)

3

1, 4

0

FH-1S(CS)

2

1, 2, 4

9

FH-16(CS)

4

0

0

FH-17(CS)

3

1, 3, 4

0

FH-18(CS)

2

1, 2, 4

0

FH-19(CS)

3

0

0

FH-20(CS)

3

1, 2, 3

0

FH-21(CS)

4

1

0

FH-22(CS)

4

1, 4

0

FH-23(CS)

3

0

0

FH-24(CS)

4

7

9

FH-2S(CS)

4

0

0

FH-26(CS)

2

1

0

FH-27(CS)

3

Total

47 young

Active nests:
Successful

IS

nests: IS

Percent

success:

- - - -

-------

1, 2, 3, 4

0

Young per successful
Young per nesting

nest:

3.1

attempt:

3.1

100%

- - - - - - - - - - - - - - - - - - - - - - - - - -

�-118-

Table l.

Ferruginous

hawk productivity

and nest site management.

(Continued)

Study Area 2
Site
no.

Young
produced

FH-l(CN)

0

0

7

FH-2(CN)

0

1, 3

0

FH-3(CN)

2

6

0

FH-4(CN)

3

1, 3

0

FH-5(CN)

4

0

6

FH-6(CN)

0

1, 3

0

FH-7(CN)

2

1, 4

8

FH-8(CN)

3

0

0

FH-9(CN)

3

0

0

FH-1O(CN)

0

1, 4

0

FH-ll (CN)

4

0

0

FH-12(CN)

2

0

6

FH-28 (CN)

0

1, 3

0

FH-29 (CN)

2

1, 4, 5, 9

0

Total

25 young

Management2/
tech. used=-

Manage. tech.
still required

Active nests:

14

Young per successful

Successful

nests:

9

Young per nesting

success:

64%

Percent

nest:

2.8

attempt:

1.8

�-119-

Table 1.

Ferruginous

hawk productivity

and nest site management.

(Continued)

Study Area 3
Site
no.

Young
produced

Managemen~~/
tech. use

Manage. tech.
still required

FH-1(P)

3

6

0

FH-2(P)

3

3, 4

9

FH-3(P)

4

1

0

FH-4(P)

3

0

0

FH-5(P)

5

1

9

FH-6(P)

2

1

0

FH-7(P)

3

3, 5

9

FH-8(P)

3

0

7

FH-9(P)

0

6

0

FH-10(P)

0

6

0

FH-ll (P)

0

0

0

FH-12(P)

0

0

0

FH-13(P)

2

0

0

FH-14(P)

3

0

0

FH-15(P)

3

2, 3

0

FH-16(P)

2

2, 3, 9

0

FH-17(P)

4

1

9

FH-18(P)

0

1, 2

0

FH-19(P)

4

0

5

FH-20(P)

0

0

9

FH-21{P)

0

9

0

FH-22(P)

0

0

0

FH-23(P)

2

1, 2

0

FH-24(P)

0

7

0

FH-25(P)

0

0

0

- - - - - -------

- - - - - - - - -

- - - - - - - - -

- - - -

�-120-

Table 1.

Ferruginous

hawk productivity

and enst site management.

(Continued)

Study Area 3
Site
no.

Young
produced

FH-26(P)

0

0

9

FH-27(P)

0

7

9

FH-28(P)

0

2

0

FH-29(P)

0

Total

46 young

Management2/
tech. used-

Manage. tech.
still required

1, 2, 3, 4, 5

0

Active nests:

29

Young per successful nest:

3.1

Successful

nests:

15

Young per nesting attempt:

1.6

Percent success:

52%

1/
2/

Management

techniques were applied in 1976.

Management

techniques

Prepared

by

applied at the end of 1977 field season.

(J,wJ;J R. ~
Gerald R. crai~
Sr. Wildlife Biologist

�-121-

JOB PROGRESS

State of

Work

W-124-R

Plan No.

v

Job Title
Period

1979

REPORT

COLORADO
No.

Project

January

Nesting

Covered:

Personnel:

Raptor
Job No.

Performance

April

of Peregrine

15, 1977 through

Daniel Berger and Gerald
James Enderson, Colorado

Investigations

1

----------------------------

Falcons

February

in Colorado

28, 1978

Craig, Colorado
College.

Division

of Wildlife;

ABSTRACT
In 1977, eleven occupied peregrine nest sites were checked for reproduction.
Broods at six of the eleven sites were augmented with captive produced
young and fledged 1.16 young per pair, five other sites fledged 0.80
young
per pair.
Four new nesting pairs were located in 1977.
Egg shell condition remained poor with approximately
20%. thinning.
Pesticide analysis of
9 eggs are presented.
Samples of avian prey were collected near two nest
sites and pesticide analysis indicates high levels in violet-green
swallows,
robins and white-throated
swifts.

��-123-

NESTING PERFORMANCE
FALCONS

OF PEREGRINE

IN COLORADO

Gerald R. Craig and James H. Enderson
The need to document population trends and reproductive success is essential to the recovery of the American peregrine falcon (Falco peregrinus
anatum).
Indeed, regular monitoring of occupied nesting sites (eyries) may
be the only effective means to establish if the population is expanding or
declining.
Since 1965, Enderson (1965) and more recently Enderson and
Craig (1974) have conducted annual surveys to follow reproductive success
of peregrine nesting in Colorado and the rest of the Rocky Mountain region.
These surveys have established that the number of productive pairs of
peregrines in the Rocky Mountain Region have steadily declined over the
past decade.
In addition to observing the eyrie sites, the investigators
collected and analyzed contents of addled eggs and measured shell thickness.
These efforts documented significant egg shell thinning and alarming residues of DDE. This valuable baseline data must continue to be collected to
monitor the decline in pesticide levels which hopefully will occur.
In
addition to the continued eyrie site survey, it is recommended that samples
of principal avian prey species should be collected from representative
hunting areas. The prey samples would then be analyzed for pesticide
residues in an effort to identify those species which may carry excessive
levels of DDE.
P.N. OBJECTIVES
The objectives

of this study are:

1.

To document the number of breeding
productivity in Colorado.

pairs of nesting

peregrines

and their

2.

To survey potential habitats to locate previously
grines and document their reproductive success.

3.

To document egg shell thinning and pesticide
peregrines in Colorado.

4.

To collect samples of principal avian prey species utilized by nesting
peregrines and analyze the samples for pesticide residues.

5.

To compile data and submit reports to appropriate state and fedetal
personnel and the Rocky Mountain/Southwest
Peregrine Falcon Recovery
Team for use in evaluating recovery efforts.

unknown nesting

residues

pere-

in eggs of nesting

SEGMENT OBJECTIVES
la.

Visit all nest sites throughout Colorado which have been occupied within
the past three years and observe them from a distance with spotting
scopes and binoculars to establish the presence of breeding adults.
All occupied sites will be revisited periodically throughout the nesting season to document reproductive success.

�-124-

lb.

Prior to, or immediately after hatching of the eggs, nest sites will be
visited and egg shell fragments and addled eggs will be collected for
pesticide analysis.
Egg shells will be measured for a thickness index
according to standardized methods.
Egg contents will be shipped to
the Fish and Wildlife Research Laboratory at Patuxent for analysis.

lc.

Successful nests will be visited prior to fledging of the young and
they will be banded and color marked.
These sites will be kept under
surveillance to determine actual fledging success.

2.

Favorable tlabitats will be surveyed for potential nesting sites. The
surveys will be conducted from the ground, but where necessary a helicopter may be used in remote regions. When new pairs are located, they
will be surveyed as outlined in approaches la, lb and lc.

3.

Ten to twenty individuals of each of 5 principal avian prey species will
be collected from representative hunting areas utilized by breeding
peregrines.
This will be done at two sites annually.
The individuals
of each species will be combined into a single sample for pesticide
residue analysis.
The analysis will be accomplished by the Fish and
Wildlife Service Laboratory at Patuxent.

4.

Compile data and submit reports to appropriate state and federal personnel and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team.
METHODS AND MATERIALS

Nesting Investigations.
Methods
cribed in the Segment Objectives

for nesting investigations
enumerated above.

have been des-

Pesticide Analysis of Prey. Because peregrines in the region are heavily
contaminated with DDT and PCB residues, we collected adult individuals of
nine potential prey species within 8.3 km (5 mi) of two active eyries
(Table 1). The species were selected on the basis of high prevalence, and
both migratory, non-migratory, herbivorous and insectivorous forms are
included.
Specimens were taken by shotgun and stored frozen. After thawing, the birds
were weighed, plucked, and the feet, beak and large and small intestine
removed.
The remainder was finely chopped, weighed and a sample taken for
pooling.
Species pooled samples were frozen in acetone-washed foil and
submitted for analysis.
Analytical procedures followed by the WARF Institute,
Wisconsin were as follows:

Inc. at Madison,

Each sample is allowed to thaw in the refrigerator overnight.
A 10
gram aliquot is weighed into a 250 ml beaker and mixed with a spatula
with 150 grams of sodium sulfate. At the same time an aliquot is
weighed into a preweighed 100 ml beaker for moisture determination and
placed in a 40°C oven for 2 weeks. After 2 weeks, weigh the beaker
and dry sample back and calculate percent of moisture.
The 10 gram
portion is allowed to dry overnight and is transferred to a 43 x 123

�-125-

Whatman extraction thimble and plugged with glass wool.
Place in a
large soxhlet extraction apparatus and extracted with 50:50 Ethyl
Ether:Petroleum Ether for 8 hours.
Sample is removed and taken down
just to dryness on a steam bath.
Sample is brought to 25 ml with
25% Toluene in Ethyl Acetate.
A 5 ml aliquot is transferred to Gel
Permeation apparatus.
The settings are 28 mins discard, 14 mins
collect and 2 mins wash. The sample is taken down on a flash evaporator just to dryness and made to 10 ml for injection.
Injection:
Hewlett Packard 57l0A with Ni63 Election capture detector,
with allta injector and hooked to a Hewlett Packard integration computor
model 3352C.
Column:
1.5% OV-17 + 1.95% QF-l on 80/100 G.C.Q.
Column Temperature:
200°C; Detector Temperature:
300°C; DDT Retention time of approximately
9.8 mins; Injection Temperature:
250°C; Carrier:
95% argon; 5% Methane
Flow 31 ml/min.
Lipid Determination:
from the 25 ml volumetric containing the sample a
5 ml aliquot is pipetted into a preweighed 2 dram vial. These are
placed into a 40°C oven for 2 days, desiccated,weighed
and percent lipid
calculated.

Table 1.

Data on prey species collected

for residue analysis

Aver. wt. of
individual
No.
homogenate
collected
(g)

Species

Individual wt.
pooled
(g)

Pool wt.
submitted
(g)

Eyrie CC
7

60

10

70

7

51

10

70

Red-winged
blackbird

5

45

10

50

Yellow-rumped
warbler

7

10

4

28

Flicker

7

97

15

105

Robin
Brewer's

blackbird

Eyrie CR
Violet-green

swallow

7

10

4

28

White-throated

swift

7

23

5

35

Clark's nutcracker

7

95

15

105

Mourning

7

81

15

105

7

96

15

105

Flicker

dove

�-126-

RESULTS AND DISCUSSION
Productivity

of Nesting

Peregrines

In 1977, 11 nest sites were occupied by pairs of falcons and a lone adult
male was present at another site early in the breeding season.
Young were
produced at nine of the sites, of which six broods were augmented with a
total of 9 captive produced young.
However, three of the sites failed
prior to fledging and only 11 young fledged for an overall nesting success
of 1.0 young per adult pair. The six sites which were augmented with captive produced young fledged 1.16 young per pair while the three unmanipulated
sites fledged 0.80 young per pair assuming that one large downy young survived despite its absence from site 30 on 6 August.
Observations of the 11
active sites are summarized in Table 2.
The percentage of occupied sites increased from 30% in 1976 to 39% in 1977
(Table 3) due to location of four previously undiscovered pairs.
Only
three nest sites continued to remain occupied out of 27 historic nests
which were known in 1970 (Table 4). Thus, increased occupancy of nests
was an artifact of expanded field investigations and does not represent
stabilization or recovery of the population.
Location

of New Nesting

Pairs

Two observers devoted the majority of the field season toward investigating
potential habitats in an effort to locate new nesting pairs.
In all, 52
localities were visited, of which 33 cliffs appeared suitable, but not
always ideal, for occupancy by peregrines.
One pair of falcons were located
where vague reports indicated their presence a decade ago. This site was
designated site 29. None of the other three new sites (sites 28, 30 and 31)
were located as a result of this effort.
Site 28 was reported by a professional biologist who happened on the pair while on vacation; site 30 was
located as a result of a raptor inventory for a federal agency; and site 31
was discovered by amateur biologists conducting an impact assessment.
Egg, Shell Thinning

and Pesticide

Residues

Table 5 summarizes egg shell thickness of all eggs collected in Colorado from
1973 through 1977. The shell condition continues to remain poor at approximately 20% thinning.
Nine intact wild eggs were available for pesticide analysis.
Three eggs were
analyzed from site 6, three from site 26 and one each from sites 27, 15 and
9. Table 6 summarizes the results of analysis obtained from the Fish and
Wildlife Research Laboratory at Patuxent.

�-127-

Table 2.

Results

of peregrine

nesting attempts,

Colorado,

1977.

Eyrie

Visits

Eggs

Young

Results

6*

5

4,4

0,0

5 April, copulation, food transfer; 17 April, removed first clutch; 8 May, incubating 2nd clutch;
11 May, removed 2nd clutch, placed dummy eggs; 23
June, 2 young put in eyrie; 29 June, female disappeared, young removed.

7*

5

3

3

13 April, copulation; 28 April, not incubating;
12 May, removed clutch, replaced with dummies; 13
June, 3 young in; 17 July 3 young
fledging.

15*

6

3

2

15 April, copulation; 22 April, laying; 6 May,
removed clutch of 3, dummies in; 26 May, 3 young
put in; 7 June, 3 young present; 21 June, 2 young
fledging.

9*

4

3

0

27 April, pair present; 12 May, copulation; 24
May, clutch removed, dummies in; mid-June, 2 young
in, early August, site abandoned.

29

9

7

0

9 April, male; 11 April, male; 12 April, male;
13 April, male; 2 May, pair, food transfer; 15
May, adult; 17 May, incubation doubtful; 5 June,
possible nest exchange; 25 June, no birds.

30

1

?

1

27 June, female and one 4-week-old young;
peregrines heard but not seen, 6 August.

26*

4

4,2

0

15 April, incubation, food transfer; 21 April,
removed first clutch; 19 May, removed second
clutch, dummies in; 20 June, 2 young in; early
July, young eaten, adults gone.

27*

4

3

2

25 March, copulation; 8 April, pair present; 29
April, incubation; 7 May, removed clutch, dummies
in; 20 June, 2 young in; fledged in mid-July.

31

1

?

1

11 July, adults and 1 flying young.

28

2

?

2

Mid-June, 2 young on ledge; late July, 2 flying
young (Craig).

25

2

?

0

11 May, pair flying about cliff; 28 June, no
birds (helicopter).

*Young added to increase productivity.

�-128-

Table 3.

Occupancy

and productivity

of Colorado

peregrine

eyries, 1972-1977.

Year

1972

1973

1974

1975

1976

1977

of eyries visited

15

23

24

26

27

31

if of occupied sites

11

12

9

8

8

12

of adult pairs

8

11

7

6

5

11

if of lone adults

3

1

2

1

iff

iff

if of young fledged

2

pair 1/
.
% of sites occupl.ed -4/

% of sites w/adult pairs

1
])

])

1
112/

0.2

11
1.6

0.8

6
1.2

73%

52%

38%

31%

30%

39%

53%

49%

29%

27%

26%

35%

if of young/adult

5

1.0

1/

One site contained 3 young, but adult female and young could not be found 2 to
3 weeks after they should have fledged.
Thus, probable fledging success is
reduced to 5.

1/

Captive produced young were placed at two sites to artificially
tion by 4 young.

augment produc-

1/ An average.of at least 1.25 young fledged per pair is considered necessary to
sustain

the wild population.

~/

Between 10% and 20% of the eyrie sites would normally
unoccupied in any particular year.

5/

Five captive

produced young augmented

reproduction.

be expected

to be

�Table 4.

Site
1
2
3
4
5
6
7
8
9

&gt;':10
&gt;':11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31

Pre1964

+
+
-1-

Occupancy of Colorado peregrine falcon eyries, 1964-1977.

1964

1965

P
P

P
P
A
A
M

P

+

1966

1967

1968

1969

1970

1971

1972

1973

1974

1975

1976

1977

P
P

P

P
P
P
P
P

M
P
P

P
P

P
P

P
P
P
A
P
P
P
P
P

P
P
P
P
P
P
P
P
P
P

V

V
V
M
V
P
P1/
PV
P
V2/
PV
V
V
P
V
V
V
V
V
V
V
V
V
P

V
V
V
V4/

V
V
A

M
P
M
V
P
P
P
V
P
V
P
V
V
V
P
V
V
V
V
V
V
V
V
V

V
V
V
V
M
P
P
V
P
V
V
V
V
V
P
V
V
V

P
P
A
P

V

P

+
+
+
+
+
+
+
+
+

P
A
F
P
V

P
M
P

V
V
V
V

V
V
V
V

].1

V
V
V
V

V
V

~

A

+

A

V

V

V
A

A
V
V
V
V
V
V
V
V
V
V

p-

P
P
V
A
V
V
V
V
V4/
PV
V
V
V
V
V
V
V
V
P
P
P

I

,......
N

I.D

I

V

V
V
V
V
V
P
P
P
P
P
P
P

P = Pair, M = Male, F = Female, A = Lone Adult, V = Site was Vacant, Blank Spaces = No Data Available.
=The se are neighboring sites (approximately I mile apart) and this could represent a shift of one pair from
site 10 to site 11.

1/ Lone adult male observed throughout season, attracted yearling female in June

1/ An adult female was found

?:../ Adult male replaced by a yearling male m Ldway through incubation.

dead in the vicinity of the
eyrie, could not confirm
presence of a pair
4/ Pair consisted of an adult male
and vear1ing female

�-130Table 5.
Eyrie

Year

1
2

1973
1973
1974
1974
1974
1974
1975
1971

"

"
"
5

"

6

"
"
"
"
"

"
"

"
"

"

"

"
"

"
"
"

"
1975

"
"

1976

"

"
"
"
"

"

"

1977

"

"
"

"

"

"

"
"
"

"
"
7

"
"
"
"
"
"
8

1974
1976

"

"

1977

"
"

9

1973
1977

"

"

10
11

1974
1975

"
"

"
"

"

15

"
"
"
"

"

1974
1975
1977

"
"

26

1977

"
"

"
"

Eggshell condition of Colorado peregrines.
Thickness (rrun).!./
W/Membrane
w/o Membrane
.325
.319
.305
.299
.287
.282
.259
.259
.259
.231
.284
.278
.294
.288
.300
.244
.246
.203
.292
.236
.295
.216
.259
.259
.185
.2032/
-- 2/
.3022/
.2262/
.3052/
.2292/
.272.1962/
.216.267
.191
.287
.203
.279
.201
.2692/
.3002/
-- 2/
.2742/
.2032/
.193.2692/
.279
.272
.249
.188
.300
.302
.244
.269
.274
.216
.244
.19l
.2542/
.2482/
.3072/
.2392/
.3232/
.2512/
.292.229=.297
.305
.297
.244
.284
.234
.305
.203
.252
.292
.218
.310
.226
.284
.201
.269
.196
.267
.211
.302

Ratcliff's
Index
1.62
1.52
1.43
1.39

Remarks

Hatched
1.49
Hatched

1.32
1.32
1.43
1.35
1.48
1.40
1.42
1.43
1.45

1.42
1.41
1.36

Broken
Hatched
Hatched
Broken
Small Embryo
27 Day Embryo
Hatched
Hatched
Died at Hatch
Large Embryo
Large Embryo
Hatched
Hatched
27 Day Embryo
No Development
Hatched
Hatched
27 Day Embryo
Addled
Addled
Hatched
Hatched
Hatched
Hatched

1.33

1.65
1.72
1.62
1.64
1.61
1.61

1.55
1.50
1.49
1.41
1.34
1.58

Ha tched
Ha tched
No development
Hatched
Spoiled
Spoiled
Spoiled

Hatched
26 Day Embryo
26 day embryo
No Development

�-131-

Table 5.
Eyrie

Eggshell condition of Colorado peregrines.

Year

26

1977

"
"

"

27

1977

"

Thickness (rnm)'Y
W/Membrane
W/O Membrane
.2672/
.3072/
.267.244
.246
.259

.196

(Continued)

Ratcliff's
Index
1.42
1.57
1.35
1.43
1.50
1.46

Remarks
No Development
Cracked
Cracked
24 Day Embryo
Hatched
Cracked

.185
"
"
"
"
------------------------------------------------------------------------------Means
1977
.280 (n=26)
1976
.283 (n=9)
1975
.273 (n=B)
1974
.280 (n=7)
19733/ .295 (n=3)
Pre 1940- .395

.209 (n=L'Z)
.211 (n=9)
.235 (n=7)
.289 (n=3)

u Measurements taken around waist of egg
'!:../ From second clutch of eggs

1/ Eggshells from Al'ta., Sask ., Montana; Anderson and Hickey, 1972.
Proceedings of XVth Int. Orith. Congr.

In

�Table 6.

Chemical

residues

in Colorado

peregrine

eggs, 1977.

(Values given are in parts per million.)

Site

21
26:1,1 - 26:1,3

Compound

26:1,4

6:1,3

6:1,4

6:2,4

27:1,1

15:1,3

9:1,3

43

19

8.3

10.9

11. 8

36.6

12.2

0.14

0.11

--

--

0.33

0.10

--

--

0.26

DDE 1/

39.5

34

DDD

0.21

--

DDT

0.37

0.43

0.55

Dieldrin

0.15

0.11

0.22

0.80

0.33

0.35

0.10

0.39

0.18

0.14

0.21

1.10

0.38

0.37

0.24

1.00

0.22

0.41

--

0.14

--

0.22

0.15

--

--

--

Heptachlor

epoxide

Oxych1ordane

--

--

--

cis - Chlordane

--

--

--

--

--

--

1
--

I-'

W
N

--

--

--

--

--

0.11

0.18

0.11

0.10

3.30

9.70

4.40

0.06

--

0.13

0.11

0.08

3.00

--

--

Es t. Toxaphene

--

--

HCB

0.25

11

3.40

II Corrected

for dehydration;

Trans - Nonach1or

--

1

0.10

cis - Nonach1or
Endrin
0.16

Mirex
Est. PCB -

21 Example:

85% H 0 taken as content of fresh egg.
2
means 1st egg of 1st clutch at site 26.

means none detected.

1.10

1.06

0.70

�-133-

Pesticide

Residues

in Prey

Results of the WARF Institute analysis of the prey samples taken in the
vicinity of eyries 7 and 15 are given in Appendix 2 and summarized in Table 7.
Six species of birds analyzed for residues approached or exceeded 1 PPM DDE,
but three species were on the order of 2 PPM or higher (Table 7). The mourning dove and flicker appear to have low residue levels, and despite their
abundance and appearance as an important prey species, are probably not a
major source of DDE to peregrines.
Clark's nutcrackers are taken by peregrines, but contain low residue.
The robin is seemingly a major potential
source of DDE; it is at least moderately contaminated, is apparently often
taken, and is uniformly common. Two other species are potentially a problem.
The white-throated swift is moderately contaminated and is often caught.
Violet-green swallows bear very high DDE and were the most abundant species
seen. If they are taken by peregrines frequently they are a clear threat.
The two species of blackbirds are moderately contaiminated, are apparently
favored prey, and are common at several eyries.
The accuracy of these assessments depend on several variables and should be
considered tentative.
Birds were collected at only two sites; residue values
found at one site may not be obtained elswehere.
Data on prey remains may not
reflect the actual frequency with which a species is taken. Featners trom
small birds could be easily overlooked, and conspicuous feathers, such as
those from flickers, may bias upwards the estimate for that species.
Finally,
almost no prey remains were collected at three eyries (11, 5, 3) from 19731977.

�-134-

Table 7.

Chlorinated

hydrocarbon

residues

in prey species.

ppm (wet weight)

% Fat

DDE

DDT1J

Dieldrin

Heptachlor
expoxide

PCB

6.2

2.07

0.07

0.07

0.02

0.12

4.3

0.84

&lt;0.01

&lt;0.01

0.03

&lt;0.10

2.5

1.28

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.10

Yellow-rumped
warbler

4.0

0.82

0.01

0.01

&lt;0.10

Flicker

2.6

.

&lt;0.01

0.06

&lt;0.01

0.01

0.02

&lt;0.10

Violet-green
swallow

5.5

5.81

0.05

&lt;0.01

0.12

0.31

White-throated
swift

5.3

1. 75

-&lt;0.01

&lt;0.01

&lt;0.01

0.12

Clark's nutcracker

4.5

0.03

&lt;0.01

0.01

0.01

&lt;0.10

Mourning

3.5

0.08

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.10

4.2

0.04

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.10

Species
Eyrie 7
Robin
Brewer's

blackbird

Red-winged

blackbird

Eyrie 15

dove

Flicker

1/

Maximum

value,

there may be interference

from PCB's.

�Appendix

1

Colorado
Colorado

College
Springs,

WARF INSTITUTE, INC.
Madison, Wisconsin

\-J'ARF
No. 7090099-010S

Colorado
RESULTS
VG-1

WS-2

RB-3

BB-4

MD-5

YW-6

RR-7

FF-S

FF-9

CN-10

DDE

5.S1

1. 75

1. 2S

0.S4

O.OS

0.S2

2.07

0.04

0.06

0.03

DDD

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

DDT

O. 05*)~ &lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

0.07'/''/'&lt;0.01

&lt;0.01

&lt;0.01

PCB

0.31

0.12

&lt;0.10

&lt;0.10

&lt;0.10

&lt;0.10

0.12

&lt;0.10

&lt;0.10

&lt;0.10

Dieldrin

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

0.01

0.07

&lt;0.01

0.01

0.01

BHC

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

Lindane

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

HCB

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

Endrin

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

H.E.

0.12

&lt;0.01

&lt;0.01

0.03

&lt;0.01

0.01

0.02

&lt;0.01

0.02

0.01

0.08

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

&lt;0.01

% Moisture

67.1

64.8

68.8

68.4

70.4

67.9

69.6

69.3

70.6

67.8

% Fat

5.50

5.32

2.48

4.25

3.48

3.98

6.21

4.21

2.57

4.54

(gm)

10

10

10

10

10

10

10

10

10

10

(gm)

4.23

4.77

5.89

8.13

8.69

6.11

7.93

9.58

7.75

8.98

Chlorinated

Screen*

Oxych1ordan

(ppm)

)""h~ResidueWet Height
****Moi~ture

(ppm)

Wet Weight

--*No other chlorinated insecticides
)bb~Weight of sample for analysis.
Limit
Limit

of detection
of detection

Prepared

by

=
=

0.10 ppm - PCB
0.01 ppm - Chlorinated
wet weight.

--= .,. \...

.••..

detected.

**Represent macium value, may be interference
****Weight of sample for Moisture Determination.
screen,

un.;t

Biologist

oxych1ordan,

moisture,

fat, residue

wet weight,

from PCB's.

moisture

I
•.....
w

lr1
I

��January,

-137-

1979

JOB PROGRESS REPORT

State of ----~~~~~------------Colorado

Work Plan No.
Job Title:

Raptor Investigations

W-124-R

Project No.

Job No.

v

~2~

Physical and Biological Analysis of Colorado Peregrine

Period Covered:

April 15, 1977 through February 28,

_
Nesting Habitat

1978

Personnel:
James Enderson and Rod Ogilvie, Colorado College;
Daniel Berger,
Beth Braker, William Burley, Marcy Cottrell, Gerald Craig, Terrence Leyden,
and Ann Potter, Colorado Division of Wildlife.

ABSTRACT
Various physical and biological factors were analyzed for six peregrine falcon
nest sites in southern Colorado.
Physical features including cliff height,
geology, elevation and distance to permanent sources of water were recorded
and compared.
Vegetative communities were delineated and transects were
established in dominant communities to compare prey abundance.
At one site
an adult female peregrine was telemetered and her movements were monitored
for 42 days during the young rearing period.

��-139-

PHYSICAL AND BIOLOGICAL ANALYSIS OF
COLORADO PEREGRINE NESTING HABITAT
James H. Enderson

and Gerald R. Craig

A variety of eyrie (nest) sites and habitats have been historically utilized
by peregrine falcons and thus it is difficult to evaluate and protect potential
nesting areas. All known historic and currently active eyrie sites should be
evaluated to record all physical and biological features of the nesting cliff
and associated hunting areas. Land use practices and disturbance factors
should also be noted to establish tolerances of breeding peregrines to man's
activities.
The resulting key characteristics which are common to all sites
will assist investigators in locating previously unknown active eyries, and
managers will be able to limit or eliminate adverse activities which may disturb breeding peregrines.
Virtually nothing is khown of the extent of hunting ranges of breeding
peregrines and so knowledge of their habitat requirements is incomplete.
The
activities of breeding adults will be monitored with telemetry during the
young rearing period.
This will allow important hunting areas to be delineated
and protected.
P. N. OBJECTIVES
The objectives

of this study are:

1.

To establish physical parameters which are essential
eyrie sites by peregrine falcons.

2.

To establish biological parameters which are common to all eyrie
sites currently or historically occupied by peregrine falcons.

3.

To delineate those human activities
by nesting peregrine falcons.

4.

Adults at eyrie sites will be trapped after their young have hatched and
radio packages will be affixed to them to monitor their movements and
hunting ranges.

5.

To assemble the data and prepare a report of the results for use in
designating new and potential eyrie sites by wildlife agencies and land
managers.

and disturbances

to occupancy

of

which are tolerated

SEGMENT OBJECTIVES
1.

Eight known historic and currently occupied peregrine eyrie sites in
Colorado will be visited and the following physical aspects will be
recorded; topography, geology, elevation, snow depth and precipitation,
mean temperature, soil, presence and distance to water, and cliff
characteristics.
In addition, a series of photographs will be taken of
each site. Each year, eight new sites will be surveyed until all known
sites have been studied.

�-140-

2a. The vegetative types of the habitat within a distance of 15 miles of
eight nesting cliffs will be cataloged whenever possible, appropriate
Forest Service vegetative maps will be utilized and meadows and other
potential hunting areas will be located on maps.
2b. Through use of standardized census techniques, avian prey diversity and
abundance will be calculated for the months of May, June and July. This
will be done at least two sites annually.
Censuses will be run once
monthly in two localities for each habitat type represented.
3.

Human activities, land use practices and audio and visual disturbances
will be noted at each site and within an area of 15 miles of each site.
If breeding pairs are present, they should be observed to note their
reactions to potential disturbances.

4.

At two sites, adult peregrines will be trapped and equipped with radio
packages.
This will be accomplished after the young have hatched and
will extend for a period of approximately 3 weeks.
The adults will be
monitored as they embark upon hunting forays and locations will be triangulated primarily from the ground.
Periodically, hunting adults will
be tracked using a fixed-wing aircraft to follow the radio signals.
All
locations of radio marked falcons will be recorded on appropriate topographic maps of the region.

5.

Compile data forms and photographs taken during 1, 2 and 3 into notebooks.
Prepare and submit reports of the results to appropriate state personnel,
federal agencies and the Rocky Mountain/Southwest
Peregrine Falcon Recovery
Team.
METHODS AND MATERIALS

Physical Analysis.
Physical analysis of nesting cliffs and the immediate vicinity
was accomplished by visiting the site and making a visual inspection of the area.
Photographs were taken of the nest cliff as well as panoramas were compiled of
the surrounding area from vantage point at the top of the cliffs.
Vegetative
information was also recorded during the visits and was further augmented from
timber maps of the region.
Prey Abundance.
Line transect
counts of bird populations were taken in June
and July at the six eyries under study. Two or three 805 m (0.5 mile) long line
transects were established within 8 km of each eyrie to quantify bird populations
within the hunting range of resident peregrines.
One transect near each eyrie
was established in the major riparian habitat, and the other transects were set
in the most prevalent plant communities in each region (Table 1). All transects
except one were continuous; the site 5 transect in forest habitat was broken
into two segments.
Bird counts were completed prior to 1000 hours by an observer walking the 805
m line in 45-60 minutes, and returning after a 15 minute wait.
Species and
numbers of individuals were recorded for all seen or heard, and unidentified
birds were classed in three size groups.
Observers alternated on different
count days.

�-141-

Table 1.

Transect

Line-transect

information

on prey counts.

It of Counts

Habitat

Count Period

Site 15,
I

Cottonwood-willow

II

Imm. ponderosa,

III

Mature ponderosa,

12

27 May-23

July

red cedar

12

27 May-IS

July

oak

12

2 June-23

July

10

13 June-IS

July

10

13 June-21

July

10

4 June-20

July

11

5 June-14

July

9

4 June-21

July

Site 11,
I

Riparian

II

Ponderosa,

juniper,

oak

Site 26,
I

Meadow,

II

Ponderosa,

Douglas

III

Spruce-fir,

aspen-Douglas

willow-birch
fir, oak
fir

Site 5,
I

Cottonwood-willow

10

24 June-22

July

II

Douglas

10

24 June-20

July

12

4 June-24

July

aspen

12

6 June-17

July

meadows

3

5 July-7 July

3

5 July-7

fir, spruce

Site 7,
I

Cottonwood-willow

II

Douglas

fir, spruce,

Site 3,
I

II

Cottonwood-willow,
Pinyon-juniper,

oak

July

�-142-

The transects used sampled over 65% all available habitat in every case but
one. An estimate of apparent rank of the eyries in relation to prey is
the source index in terms of the % of the area samples.
This ranking is
only approximate because unsampled habitat with high bird biomass could
raise an eyrie's rank, or low biomass in unsampled habitat could lower the
rank, had transects been run in all habitats.
The ranking is most reliable
where high percentages of the habitat available was sampled.
Stream-side cottonwood-willow
or willow-birch habitat was sampled in every
case. This habitat, situated in drainages, varied between about 1950 ill and
2700 m in elevation between eyries (Table 1). Transects in this habitat
at lower elevation tended to yield higher average total biomass values than
those at higher elevations (Table 5). This habitat transect at the three
lowest elevations yielded average biomass exceeding 3.0 kg while those three
at higher elevations ranged from about 2.5 kg (Site 26) to about 1.8 kg
(Sites 5, 7). Biases perhaps entering into this analysis include the nature
of adjacent habitat and the areal extent of vegetation type. The cottonwoodwillow transect at Site 3, with extensive adjacent meadow, approached 6 kg
of bird biomass, and must represent a good source of vulnerable prey despite
the limited sampling (three transect counts).
But even though bird biomass in cottonwood-willow
habitat was always substantially higher than in other habitats at each eyrie, especially for the
two most vulnerable classes of birds, the low frequency of the habitat reduced
its potential as a prey source to low significance at all but one site (Site
3) (Table 6). However, this habitat may be important to peregrines as a
source of upland prey passing over the bottom land or going there for water,
two factors our census techniques might not have revealed.
At three eyries (15, 11, 26) ponderosa forest in various stages of mlxlng
with brush or Douglas fir yielded very high source indexes of vulnerable
species compared to other habitats (Table 7). This is due both to high bird
biomass and to extensive areal coverage.
Mainly because of this, these three
regions rank well above 5 and 7, sites where spruce-fir mixed with Douglas
fir cover large areas yet are poor in bird biomass.
These findings assume
that, at all sites, unsampled habitats are not big enough or rich enough to
provide peregrines with important sources of food.
Although transects at eyries 7 and 5 sampled vegetational types comprlslng
only 65 and 48% of the total habitat, respectively, the source indexes of
vulnerable prey were very low. It is apparent these sites are situated in
relatively poor locations from the standpoint of prey abundance, although
this does not necessarily mean that inadequate numbers of vulnerable prey are
present or exposed to predation.
Radio Telemetry.
We attempted to trap the adult males at three active eyries
for instrumentation with radio beacons.
None was caught, but a female was
tagged at eyrie 15 to illustrate preferred hunting habitat, range size, and
time and duration of hunting sorties.

�-143-

The transmitter was attached to a single tail feather with nylon thread on
7 June and remained operative until the study was terminated 19 July. Two
receiver stations were established, one 0.3 km south of the eyrie, the other
5.8 km north on a knoll of Devil Mountain, and both provided a maximum unobstructed overlook of the region.
Portable transceivers provided communications
between the stations.
Beacon signals were sought simultaneously
with both receivers at five minute
intervals when the falcon was perched, and at 30 second intervals when flying.
Receiver stations were operated at total of 90.2 hours on 19 different days.
Time, bearing, and signal strength were recorded at each station.
USGS 7.5
minute series topographic maps of the region were used to plot signal positions
by triangulation.
Points were plotted only when both receiver stations obtained
signals at the same time and points of doubtful accuracy were not plotted.
Range of signal reception is highly variable.
Line-of-sight range can exceed
32 km, but this is reduced to a few km if obstructions lie between transmitter
and receiver.
If the transmitter antenna is in line with the receiver antenna,
such as during a straight-away departure or approach, no signal is received.
Due to the broken terrain of the region, relatively few signals farther than
9.6 km were received by both stations.
Weather affected reception; calm,
overcast days correlated with strong reception, stormy days with lightning
caused poorer reception.
Error in bearing determination is thought to be plus
or minus ten degrees or arc, variation that could result in a position error
of a few kilometers for single determinations at the edge of reception range.
Where many readings are taken, this error would tend to cancel out.
RESULTS AND DISCUSSION
Physical Attributes

of 6 Nest Sites

Data were obtained on the physical attributes of four active eyries (one
with a lone male) and two inactive eyries in southwestern Colorado in an 80
km-wide strip adjacent to the New Mexico line (Table 2). The eyries averaged
2664 m (8763 ft) in elevation and none were within 8 km of surface elevations
below 1976 m (6500 ft).
The vertical cliffs containing the nest-ledges ranked
from 22 m to 198 m in height, averaging 94 m (174 ft). Height above the
nearest major drainage, usually containing a permanent stream, averaged 398 m
(1309 ft.). No eyrie was farther than 4 km from the nearest river or lake and
the mean distance was 2.4 km.
Nest ledges showed no pattern in regard to direction of exposure, fully onehalf of the ledges faced some heading with a northward component.
None were
exposed to the southern quadrant (1350-2250).
Invariably the nest ledges
were on very irregular cliff faces including Cretaceous sandstones (3 sites),
Cenozoic lavas and breccias (2 sites), and Pennsylvanian limestone and shales
(1 site). All of the sites command a dominant view of surrounding mountains
and lowlands and all overlook permanent streams (5 sites) or lakes (1 site).
Factors relating to the quality of peregrine habitat must be functions of
the physical and biological attributes of the region.
Easiest to describe

are

�Table 2.

Physical

characteristics

of peregrine

eyries.

Eyrie
Cha racteri stic

15

11

26

5

7

3

Means

Elevation of eyrie {m)

2402

2538

L873

2584

3040

2546

2664

Cliff height (m)

24

22

137

198

129

53

94

Exposure (degrees)

300

100

90

285

20

110

Height above
major drainage (m)

445

365

274

314

407

5135

398

Elevation of drainage (m)

1957

2173

2599

2270

2633

1961

-

Distance to nearest
permanent water (km)

I
I-'

r--

~
I

2.9

3.5

2.2

0.6

Geologic formation

Picture
Cl iff
sandstone
on Lewis
Shale

Mesa Verde
sandstone,
Menefee
Formation

Hermosa
Formation
marine
shale,
1imestone
sandstone

Lavas and
brecci as

Lavas and
breccias

Mesa Verde
sandstone
on Mancos
Shale

Geologic age

Upper ,
Cretaceous
90 million

Upper
Cretaceous
90 million

Pennsylvanian -",
250 mi 11ion

Oligocene
33 mi 11ion

Oligocene
33 mi 11ion

Upper
Cretaceous
90 million

1.3

4.0

2.4

�-145-

physical attributes.
Although our selection of eyries for study was not
random among sites in the Rocky Mountains, none selected is physically atypical.
An obvious factor among the eyries studied is great relief above surrounding
lowlands, the differential always exceeding 300 m. Height of the actual cliff
appears relatively unimportant.
That four in six of the nest cliffs were
about 100 m high, or higher, may relate more to selection of sites high above
the surrounding lowlands than selection of high cliffs ~
se. Another factor,
as yet unmeasured, relates to the "dominance" of the cliff, however high its
face, over the surrounding landscape.
This condition depends on the slope
conditions above and below the cliff, and on the contour of the land adjacent
to the face. In any case, an analysis of the physical attributes of eyrie
locations may not yield the same conclusions now when the population is depressed that it would were the population large.
Biological

Conditions

of 6 Nest Sites

Regional flora. We estimated the percent coverage of nine major plant communities within an 8.3 km radius of each of the six eyries (Table 3). These
estimates were made by visually sub-dividing the area extending 8.3 km from
the eyrie into many small sectors.
Each sector was then viewed with binoculars
and percentages of each vegetational type in the sector estimated.
Finally,
sector percentages were summed for each vegetational type. Error is reduced
at most sites by the over-look nature of the nesting cliff.
We found high variation between sites in regard to frequency of plant community types. Three eyries (26, 5, 7) are predominantly surrounded by mesic
transition or spruce-fir forests, usually included substantial tracts of seral
aspen forest. One site (3) overlooks mainly pinyon-juniper woodland and
cleared pasture land. One site (11) occupies mainly cleared ponderosa forest,
and oak brushland.
The remaining site (15) is surrounded by a mix of meadow,
brushland, transition forest, and most importantly, mid-aged to mature ponderosa forests. All eyries are near cottonwood-willow riverbottom habitat, but
such habitat never exceeds 5% of the total.
Flora at nest-cliff.
Vegetation immediately adjacent to the nest-cliff varied
greatly from site to site. The following is a vegetational description, by
site, of the major plant species within 0.83 km of the eyrie.
Site 15 - The steep slopes around the cliff are covered with moderately dense
Gambel's oak and mountain mahagony on the north and west, and with mixed
pinyon pine, juniper, and mountain mahogany on the east and south facing slopes.
Ponderosa pine and a few Douglas firs grow in more moist situations on the
lower slopes. About 15% of the slopes near the cliff are barren due to the
steep pitch and low moisture content of such extremely fine shale-derived soils.
Other shrub species (in approximate order of frequency) are: serviceberry,
cliff rose, sagebrush, chamisa, currant, skunkbush, and snowberry.
Sparse
grasses, opuntia cactus, yucca, are also prevalent.
Site 11 - A nearly uniform stand of brush species surrounds this site. The
dominant species are Gambel's oak, serviceberry, and mountain mahogany.
Northfacing slopes adjacent to the cliff and those in the ravines below have a

�-146-

Table 3. Frequency
falcon eyries.

of vegetational

types within

a 8.3 km radius of peregrine

Estimated frequency by eyrie (%)
Vegetational Type

15

11

26

5

7

3

Meadow, pasture

12

5

5

13

24

32

Cultivated land

&lt;1

1

&lt;1

0

0

2

Cottonwood-wt 11ow

5

1

&lt;1

2

4

&lt;1

Mixed brush1and

18

37

5

&lt;1

3

15

Ponderosa forest

46

27

17

5

8

0

Aspen forest

&lt;1

7

13

33

21

0

Pinyon-juniper woodland

5

10

0

0

0

49

Transition forest*

13

10

37

25

14

2

Spruce-fir forest

0

2"

20

21

26

0

*Doug1as fir, ponderosa pine, white fir , limber pine, in order of
significance.

�-147-

sparse growth of young Douglas firs, and a few ponderosa pine in open stands.
Above the cliff some oak has been cleared exposing a mosaic of grass patches.
Site 26 - Immediately below the cliff is a mature transition forest containing 60% Douglas fir, 25% limber pine, 10% ponderosa pine, and 5% whf. te
fir. Understory species are snowberry, chokecherry, currant, and do~vood.
Meadows a few hundred meters below the cliffs are bordered often with large
pure stands of mature aspen. Slopes facing eastward below the cliff may have
dense stands of Gambel's oak.
Site 5 - The mesic slopes below this site have thick stands of Douglas fir,
with some blue spruce and a few aspens. Above the cliff Douglas fir and
limber pine predominate.
The valley floor is open meadow with narrow-leaf
cottonwoods and blue spruce near the river. Mature ponderosa pine exist
on the gentle slopes adjacent to the pastureland.
Site 7 - This cliff outcrops above a mountainside thickly forested with
Douglas fir, and near the top mature stands of Englemann spruce. Above the
cliff the ground is 40% meadowland with large stands of iwnature aspen, many
of which show heavy wind damage.
The river is about 0.5 km below the cliff
down a steep mountain side: its margins are lined with cottomvoods, willows,
blue spruce, Douglas firs, and, on the drier slopes, ponderosa pine.
Site 3 - Above the cliff a mature pinyon-juniper woodland extends over the
mesa.
Immediately below the cliff is a steep area of mud and rock slides on
degraded Mancos shale. Stands of Gambe1's oak, mountain mahogany, with
serviceberry and snowberry occur on the more stable slopes.
Be Low the long
talus slopes 'mature stands of pinyon pine and juniper border cleared pastureland. Although all six eyries under study were at elevations within a 600
m range, surrounding plant community frequencies (Table 3) varied greatly
between sites. Sites 5 and 26 are the most mesic; subalpine and transition
forests and seral aspen are prevalent.
Site 7, although similar, is drier
(Eastern Slope) and meadow is more frequent.
The other two are in brushland
with open montane forest (11) or in pinyon-juniper woodland with extensive
meadow and pasture1and (3).
Prey Abundance.
Several biases were probably present in the collection of
the transect count data. Early in the period, the four observers involved
in counts were not equally competent, especially in regard to bird calls and
songs. This bias is probably less important for the most common and conspicuous
species.
As the suwner progressed birds often recorded by song may have been
increasingly overlooked as the breeding season passed.
Individuals of each species for the round trip on the transect were divided
by two to yield the mean number of individuals seen on the 805 m transect on
a given day. These means were in turn averaged to give the mean number of
individuals of each species for each transect in the study period (Appendix
1) .

Each species was classified in one of three arbitrary categories of vulnerability to peregrine predation.
We based these determinations of the relative
amount of time each species was seen to fly in the open at least 15 m from

�-148-

the ground or other cover.
"A" category included forms often seen away
from cover, ego Clark's nutcrackers;
"B" category included forms often
seen in cover but given to occasional long flights in the open, ego robins
and flickers; "e" category included apparently non-vulnerable species
such as rofous-sided towhees.
Large species such as buteos, ravens, and
large waterfowl were given a "not applicable" (N.A.) rating.
Table 4
includes all species seen, their vulnerability ratings and approximate body
weight.
Next, the mean number of individuals for each species recorded on the transect for all counts was multiplied by the species approximate body weight.
This yields an index of prey biomass for each transect.
Table 5 expresses
these indexes of biomass for each transect by vulnerability rating.
Unidentified birds were classified as small (sparrow-sized), medium (robinsized), or large (rock dove-sized) and were assigned corresponding weights.
Never were bird (prey) populations sampled in more than three habitats near
any eyrie, but often those habitats sampled were prevalent in the area and
sometimes contained two or three vegetational types. An index of the prey
source value of each habitat sample within 8.3 km (5 mi.) of the eyrie can
be derived by multiplying the freqeuncy of that habitat (% of all habitats)
by the bird biomass for each vulnerability group (Table 6). The result is
an estimate of prey biomass tempered by the amount of the corresponding
habiatat.
Sometimes transects sampled more than one vegetational type (Table
3); in those cases the frequencies of the appropriate types were summed.
In
Table 6, unidentified birds usually accounted for a minor part of the total
biomass, and since most unidentified birds were small forms, they probably
can be discounted as significant prey for peregrines.
Table 7 sunnnarizes the value of habitats adjacent to the six eyries.
Source
indexes for vulnerability classes A and B only were used, assuming class e
birds, which generally had low biomass, are not very significant as prey
sources.
Radio Tracking

of a Nesting Peregine

Although most data points are concentrated within five kilometers of the
eyrie, 10 of 15 sorties included excursions more distant than 9 km. Hunting
sorties exceeding about 9 km from the eyrie occurred in all directions except
north, northwest and south.
The greatest concentration of position points
lie centered 2 km east of the eyrie, we suspect the bird was loafing and not
hunting during many of these flights.
Much of these latter data were obtained
in the period 30 June - 3 July shortly after the young fledged.
In this
period the falcon often spent over 5 hours at a time in that area without
returning to the eyrie.
At least half of the falcon's time away from the eyrie appears to have been
spent in canyons or valleys characterized by open meadows and ecotonal
connnunities. Areas at higher elevations were rarely visited; few data points
are located above 2400 m (8000 ft.). Very little time was spent in the continuous ponderosa forest north of the eyrie, an area within reception range of
both receiver stations.

�-149-

Table 4. Species, body weights,
on prey transects.

.Species

Black-Crowned Night Heron
Mall ard
Blue-winged Teal
Common Merganser
Turkey Vulture
Goshawk
Coot er 's Hawk

Sheo-shinned Hawk
Red-tai1ed Hawk
Kestrel
Blue Grouse
Sora
American Coot
Ki11deer
Common Snipe
Spotted Sandpiper
Mourning Dove
Pygmy Owl
.Poor-wi11
Common Nighthawk
White-throated Swift
Rufous Hummingbird
Broad-tailed Hummingbird
Belted Kingfisher
Common Flicker
Lewis' Woodpecker
Yellow-bellied Sapsucker
Williamson's Sapsucker
Hairy Woodpecker
Downy Woodpecker
Cassin's Kingbird
Ash-throated Flycatcher
Say s Phoebe
Empidonax Flycatcher
Western Wood Pewee
Violet Green Swallow
Tree Swa 11ow
Rough-winged Swallow
Sarn Swa 11ow
C1 iff Swallow
Gray Jay
Steller's Jay
I

and vulnerability

ratings

Body weight
(g)

Sourcea

NA
NA
400
NA
NA
NA
NA
140
NA
140
NA
75
435
85
112

48
130
100
50
75
38
3
3.5
155

140
120
70
70
80
25
40
30
20
12
15

3

1

of birds seen

Vulnerability

NA
NA
A
NA
NA
NA
NA
B

1

NA
A
NA
C
A
A

1

A
A

1

3
1

.1
1
3
3
3

2
3
3

1
2
3
3

B
C
C

A
A
B
B
A
B
A

3

B
B

3

B

1

B

3
3

A

1

A

B

3

C

3

C

19

3

20

1
1
1

A
A
A
A

3

A

16

17
17
90
100

3

B

3

B

�-150-

Table 4. Species, body weights,
prey transects.
(Continued).

Species
Scrub Jay
Magpie
Clark's Nutcracker
Crow
Raven
Black-capped Chickadee
Mountain Chickadee
White-breasted Nuthatch
Red-breasted Nuthatch
Pygmy Nuthatch
Brown Creeper
House Wren
Benwi ck 's Wren
Nockingbird
Catbi rd
Robin
Hermit Thrush
Swainson's Thrush
Verry
Western Bluebird
Mountain Bluebird
Townsend's Solitaire
Blue-Gray Gnatcatcher
Ruby-crowned Kinglet
Cedar l&gt;laxwing
·Starling
So 1itary Vi reo
Warbling Vireo
Orange-crowned Warbler
Virginia's Warbler
Yellow Warbler
Yellow-rumped Warbler
Black-throated Gray Warbler
Grace's Warbler
MacGillivray's Warbler
Yellow-breasted Chat
Wilson's Warbler
Western Meadowlark
Red-winged Blackbird
Scott's Oriole
Northern (Bullock's) Oriole
Brewer's Blackbird
Brown-headed Cowbird
western Tanager
Rose-breasted Grosbeak
Black-headed Grusbeak
Lazuli Bunting
EVening Grosbeak
Cassints Finch

and vulnerability

ratings of birds seen on

Body weight
(g)

Source

Vu1nerabi 1ity

90
150
145
NA
NA
12.5
12.5
17
10
7
8

3
2
2

B
A
A

1
3
3
3
3
3
3

11

11
/0
39
80
30
30
33
33
30
27
7
6.7
25
82
17
13
12
12
12
14
13
12
12
28
12
145
60
70
70

70
50
29
111'\

or .

45
lj

52

25

NA
NA
C
C
C
C
C
C
C
C

3
3

A
B
B

1

1
3
3
1
1
2
3.
1
1

C
C
C
B
A
B
C
C
B

1

A

3

3
3

3
3
3

3
3
3
3
1
3

1
2
3
3
3

1
2
1

C
C
C
C
C
B
C
C
C
B

C
A
A
A

B
A
A
,.,
0
0

3

ts"

3
3
3

B

C

B

�-151Table 4. Species, body weights,
transects.
(Continued).

Species
Pine Grosbeak
Pine Siskin
American Goldfinch
Lesser Goldfinch
Green-tailed Towhee
Kufous-sided Towhee
Vesper Sparrow
Lark Sparrow
Sage Sparrow
Gray-headed Junco
Tree Sparrow
Chipping Sparrow
White-crowned Sparrow
Song Sparrow
Total Species:

and vulnerability

ratings of birds seen on prey

Body weight
(g)

Source

60
12

3

C

3

A

14

3

14
38
42
27
23
18
22
18
13.5
25

3
3

A
A
C
C
B

25

1
3

Vulnerability

3"

B

3

C
C
C
C
C
C

1
3

1
3
3

105 (A-31~ B-28, C-37, NA-9)

al) Poole, E. L. 1938. Weights and Wing Areas in N. A. Birds.
2} Weights from fresh specimens.
3} Weights estimated from those known in same size range.

Auk 55:511-517.

�-152-

Table 5. Average prey hiomass indexes by vulnerability class for linetransects.
Average bird biomass per transect
by vulnerability class (g)
UI;~
B
C

It Counts

A

I

12

1362

1097

395

200

3054

II

12

1001

448

335

90

1874

III

12

466

1065

493

l34

2158

I

10

642

1838

896

185

3561

II

10

1053**

660

376

l35

2224

I

10

1129

896

328

63

2436

II

11

430

658

320

120

1528

III

9

290

785

403

163

1641

I

10

825

684

159

151

1819

II

10

136

300

159

168

763

I

12

790

659

301

141

1891

II

12

19

266

191

91

567

I

3

4197

994

416

153

5760

II

3

342

277

252

48

919

Transect

Total

Site 15

Site 11

Site 26

Site 5

Site 7

Site 3

* Unidentified birds, not given vulnerability rating, but classified small,
medium, or large. Category "small" greatly predominates.
1::*

Value is 481 excluding biomass of mourning doves passing over habitat enroute
to meadow areas.

�Table 6.

Relative

values of habitats

adjacent

to peregrine

A

eyries as prey sources .

Average Biomass by
Vulnerability (9)
A
B
C
UI

Habitat

~nsect

.

A X Bb
Ba
Habitat Freg. (%)

A

B

C

UI

Source Index

CR-!

Cottonwood-willow

1362

1097

395

200

5

68

54

20

10

CR- II

Ponderosa (imm.)

1001

448

335

90

46

460

206

154

41

CR- I II

Ponderosa (mat.) - oak

466

1065

493

134

31

144

330

152

41

PP-I

Cottonwood-willow

642

1838

896

185

1

6

18

9

2

PP-II

Ponderosa-brushland

1053

660

376

135

64

674

422

241

86

HH-I

Meadow, willow-birch

1129

896

328

83

6

68

54

20

5

HH-II

Ponderosa, D. fir, oak

430

658

320

120

59

254

388

189

71

HH- III

Spr~ce-fir, aspen

290

785

403

163

33

96

259

133

54

WC-I

Cottonwood-willow

825

684

159

151

2

17

14

3

3

tvc- II

Douglas fir, spruce

136

300

159

168

46

63

138

73

77

CC-1

Cottonwood-willow

790

659

301

141

4

32

26

12

6

CC-II

Douglas fir, spruce, aspen

19

266

191

91

61

12

162

117

56

I'IV- Ie

Cottonwood-willow,

4197

994

416

153

33

1385

328

137

50

rW-IIc

Pinyon-juniper,

,342

277

252

48

64

219

177

161

31

oak

meadow

al~here transects occurred in more than one vegetation type (Table II), frequencies of each type were summed.
blast two digits dropped.
CMesa

Verde transects too few (3) for comparability.

I
•....•
V1

w
I

�-154-

Table

7.

Transect

Summary

of habitats

as prey sources.

Source Inc.Jexa

Habitat

% of Total

Habitat·

CR-I

Cottonwood-willow

122

5

CR-II

Ponderosa (imm.)

666

.46

CR-I II

Ponderosa lmat.), oak

474

31

Totals

1262

PP-I

Cottonwood-willow

PP-II

Ponderosa, brushland

24

1

1096

64

Totals

1120

HH-I .

Neadow, willow-birch

122

6

HH-II

Ponderosa, D. fir, oak

642

59

HH-I II

Spruce-fir, aspen

355

33

Totals

1119

WC-I

Cot tonwcod-wt 11ow

31

2

WC-IJ

Douglas-fir, spruce

211

46

Totals

242

CC-I

Cottonwood-willow

58

4

CC-II

Douglas fir, spruce, aspen

174

61

Totals

232

MV-I

Cottonwood-willow,

MV-II

Pinyon-juniper,

meadow

1713

33

396

64

oak

Totals
aSum of vulnerability

2109

classes A and B.

bSum of percentages for sampled vegetational
c(tota1 source index/% of habitat sampled).

types.

ApparentC
Rank

82

3(15)

65

2{l7)

98

4(11 )

48

5(5)

65

6(4)

97

1 (21)

�-155-

The time and duration of hunting sorties follow no clear pattern.
Sorties
were scattered throughout the daylight hours.
A single sortie lasted from
30 minutes to over seven hours. Frequently, trips extended beyond the
observation period, so that either exact time of departure or time of return
could not be ascertained.
Often sorties extended into more than one region.
For example, on 3 July the falcon spent time 1) traveling to the south along
a river, 2) 3 to 9 km west of the eyrie, and 3) about 16 km west of the eyrie.

Prepared by

R. ~.~

~
Gerald R. Craig
Sr. Wildlife Biologist

�APPENDIX
Peregrine Prey Counts:

I

Average Number Per 805 m (0.5 mi) TRANSECT

.

Speci es

Transect
CRl

CR2

CR3

PPl

PP2

HHl

HH2

HH3

WCl

WC2

CCl

CC2

Heron, Blk-cr Night
Ma 11ard

.92

Teal, Bl-winged

.17

Merganser, Common

.17

.10

.63

Duck, UD

.33

.15

.50

Vulture, Turkey

.04

1.00

.40

Goshawk
Hawk, Sharp-shinned

MV2

.17

.33

.13

.27

.06

.05

.10

.83

.04

.05
I

I

.04

.08

.10

Hawk, Cooper's

.04

.04

.15

Hawk, Red-tailed

.08

.13

Falcon, Peregrine

.08

Kes tre 1

MVl

I-'

.11

V1

.05

cI

.33

.33

.35

.05
.20

.06

.18

.15

.05

.05

.08
.17

.04

Grouse, Blue

.83
.05

.06

.25

.08

Sora

.17

Coot, American

.55

Ki 11deer

2.63

Snipe, Common

.08

Sandpiper, Spotted

.83

Dove, Mourning

2.38

.25

.17

1.35
3.33

2.13

l.95

4.40

.10

2.60
.09

.10

.50

2.46

.33
3.67

.50

�APPENDIX
Speci es
Owl, Pygmy

CRl

CR2

CR3

PPl

PP2

.

.05

Nighthawk, Common

.15

Swift, Wh-throated

.08

Hummingbird, Br-tailedl .29

.17

Hummingbird, Rufous

'.08

Hummingbird, Ud.

.29

Kingfisher, Belted

.67

Flicker, Red-shafted

1.75

.25

.17

.08

HH2

HH3

WCl

WC2

CCl

CC2

.05

.78

.35

1.45

.08

.25

Sapsucker, Williamson

.25

.60

.73

1.50

.04

2.L9

.92

.05

.05

.35

.15

.10

.09

.39

.30

1.00

.33

.45

.05
.17

1.88

.08

I

.30
.10

Sapsucker, Yel-belliedil.54

4.75

MCl

MV2

.33

.50

1.40

.13

.04
1.36

1.1

1.15

.55

.75

.66

3.17
I
•....

.70

VI

-...J

.05

.17
.08

.04

.40

Woodpecker, Downy

.21

.04

.60

Woodpecker, Ud.

I .13

.08

.20

Kingbird, Western

I .02

Flycatcher, Ash-thr

.08

Phoebe, Say's

.08

Pewee, Western Wood

2.50

.08

2.30

.17

.10

Woodpecker, Hairy

Flycatcher, Olives i ded

HHl

.05

i

Poor-will

Woodpecker, Lewis'

(Cont. )

I

.15

.05

I

.32

.22

.45

.36

.28

.25

.23

.06

.05

.05

.10

.14

.29

.34

.95

.15

.79

.83

.04
.08
.05

.10

.10

.05

1.00
."17

.17

1.50

.14
.10

1.58

.50
.04

5.25

1 ~15

2.00

1.05

.61

.45

.55

.46

.38

.33

.33

�Speci es
Flycatcher,
(Empidonax)

CRl
I
P·50

Flycatcher, Ud.

: .04

Swa 11ow, Vi01et-green

15.58

Swa 11ow, Tree

0.46

Swallow, Rgh-winged

1.13

APPENDIX I

(cont.)

CR2

CR3

PPl

PP2

HH1

HH2

HH3

HCl

WC2

CCl

CC2

.58

.42

(.25

.20

.90

1.09

1.11

.75

.40

.21

.08

.04

.20

.10

.09

.11

.05

2.13

9.30

17.95

14.00

6.4

3.50

.30

.96

1.10

.05

1.05

.27

.66

.

1.35

.05

.21

Swallow, Ud.

1.08

4.6

1.29

.17

.33

1.0

.79
.50

.04

.17

.17

.33

.17

.10
.08

.50

.25

.25

1.05

.64

.30

1.96

Jay, Gray

.17

Jay, Stell err s

.75

Jay, Scrub

4.13

1.00

.29

3.60
.80

2.20

4.00

1.00

.25

Raven, Common

.25

1.17

.13

CrO\&lt;/,Common

.67

.17

Nutcracker, Clark's

.04

.63

.63

.20

.80

Chickadee, Blk-capped

1.13

0.42

.50

1.60

1.15

.35

.55

.86

.33

1.20

.95

.50

.35

.45

.09

.35

.27

.10

.05

.25

.80
.06

.42

.85

.25

1.05

2.27

6.39

Nuthatch, Wh-br

.63

.50

1.67

2.10

.35

.30

.96

1.50

.05

.18

1.56

1.56

1.10

.50

.65

.05

.08

.08

.67

.33

3.67
.17
2.33

1.00

4.33

1.17

.17
.15

._.

Nuthatch, Red-br

3.21

.11

0.33

1.88

V1

.05

.05

.08

.54

I
•....

.38
1.83

Chickadee, Mountain

.04

.20

.20

Magpie, Blk-bi11ed

Nuthatch, Pygmy

MV2

.04

Swallow, Barn
Swallow, Cliff

MV1

.10

.58
3.00

.10

.25

2.83

.17

.08

.04

1.67

1.00
.33

.66

.50

00
I

�APPENDIX I
Spe~i~2

CRl

Creeper, Brown

I

Dipper (Water Ouzel)

I

Hren, House

!6.88

Hren, Bewi ck IS

I .08

Mockingbi rd

I

Catbird

I

Robin

CR2

CR3

PPl

HHL

HH2

.0'5

HH3

1.54

.06

.79

8.25

.04

1.25

.04

.15

.95

.05

.25

.17

2.71

.17

MVl

MV2

.67

.50

.65

Bluebird, Western

I .96

Bluebird, Mountain

I

Solitaire, Townsend's

I .04

Blue-gray

3.17

.17

2.38

7.25

1.60

4.65

2.00

5.44

4.80

1.80

5.13

.75

.63

.85

.85

.15

2.41

2.06

.15

1.20

.17

1.96

.06

.15

.10

.04
.05

1.21

.21

1.45

.80

I
•....

.58

.17
1.4

4.00

V1

'"
I

0.27

.06
.83

.10
.66

.21

1.08

.25

.40
,05

Kingley, Ruby-cr

.05

.25

.09

1.17
3.85

.41

.10

.09

6.83

Waxwing, Cedar

I .25

Starling

I .08

.08

Vireo, Solitary

I .54

.54

1.25

.70

.30

.30

.96

.60

Vireo, Warbling

i2.58

.04

1.50

10.65

1.10

.75

2.32

4.06

.04

.10

.05

.05

0.73

3.33

.60

1.35

.05

i

I

1.17

.29

.10
.05

I

CC2

.05

Veery

Warbler, Virginia's

CCl

.05

Thrush, Swainson's

I
I

WC2

.66
1.04

7.16

\~arbler, Org-cr

WCl

.15

Thrush, Hermit

Gnatcatcher,

PP2

(Cont.)

0.9

1.85

3.67

.10
.06

1.41

.71

2.17

.04

1.25

.65

.46

.46

.50

.17

�APPENDIX I
Speci~

CRl

Warbler, Ye lIow

10.38

Warbler, Yellowrumped
Chat, V-breasted
Warbler, Townsend's

CR2

CR3

PPl

PP2

.

.10
0.75

2.13

2.10

(Cont. )
HHl

HH2

.50
.45

.45

2.09

HH3

WCl

.06

2.35

5.50

3.3

VlC2

CCl

CC2

7.04
.80

1.08

.63

MVl
13.83

1.42
1.00

.05

Warbler, MacGillivray

.08

Warbler, Wilson's

.04

\~arbler, Ud.

.04

.

Warbler, Grace's
Spa rrow, House

I .13

Meadowlark, Western

I

.08

.21

1.13

.92

1.05

.25

.25

.25

.35

.11

.40

.22

.05

.45

.05

.17
1:00

.05

.04

.32

.29

Blackbird, Red-winged

3.04

Oriole, Bullock's

.25

5.0
1. 70

.09

.06

3.35

25.67

Oriole, Scotts

I
•....•

.17

l.67

Blackbird, Brewer's

2.54

Grackle, Common

.42

Cowbird, Br-headed

.08

Blackbird, Ud.

.17

.83

.79

.40

1.90

.13

.55

.27

.20

.11

1.04

3.15

.11

5.50

2.83

6.50

.66

.06
•...

Tanager, Western

.46

l.58

1.42

Grosbeak, Rose-br

.95

.85

.10

2.14

.05

Grosbeak, B1k-headed
Bunting, Lazuli

MV2

.96

I

.08

l.33

.75

.06

.10

.30

.08

.05
.15

.20

.18

.10
.25

.67

0\

0
I

�•

APPENDIX I
Speci es
Grosbeak, Evening

CR1

CR2

CR3

PP1

I

(Cont. )

PP2

.

HHl

.05

.15

HH2

HH3

.17

Finch, House

CCl

CC2

MVl

.05
.17

.05

Siskin, Pine

.08

Goldfinch, Lesser

.25

Towhee, Gr-tai1ed

.04

.83

.96

4.1

3.30

Towhee, Rufous-sided

.58

5.67

5.42

.95

3.05

.63

.29

.25

.10

9.10

.68

5.33

.45

.35

.13
.17

.10
.08
.17

I

1.83
.66

.04

I

MV2

.08

Grosbeak, Pine

Sparrow, Lark

WC2

.25

Finch, Cassin's

Sparrow, Vesper

~'JC1

.50

I
&gt;-'
(J\

.13

Junco, Gray-headed

.71

Sparrow, Tree

2.82

.05(?)

Sparrow, Chipping

.25

4.58

l.50

Sparrow, Wh-crowned

6.55

3.45

3.65

l.09

1.8

.15

.75

.06(?) .05

.10

1.85

.30

.06

l.46

I

.15

.05

5.10

.78

.60

.56

l.5

&gt;-'
I

l.58

.08

1.17

7.83

.42

.05

.05

Sparrow, Song
Sparrow, Ud.

.95

.55

4.50

4.71

.66

Ud. Small

4.5

2.29

3.67

6.0

2.70

2.50

5.23

7.50

4.45

3.45

5.13

4.08

3.17

1.67

Ud. Medium

1.88

0.79

1.13

1.35

.85

.65

.59

.72

.70

1.05

.92

.42

1.50

.33

.06

.05

.05

Ud. Large

.05

.13

-

��-163January 1979

JOB PROGRESS

COLORADO

State of
Project No.

W-124-R

Work Plan No.

v

Job Title

Raptor Investigations
Job No.

3

------------------------

Reintroduction

Period Covered:
Personnel:

REPORT

and Augmentation

of Peregrine Falcon Production

April 15, 1977 through February 28, 1978

Daniel Berger and Gerald Craig, Colorado Division of Wildlife;
James Enderson and Rod Ogilvie, Colorado College.

ABSTRACT
Management activities were implemented to increase peregrine falcon reproduction in Colorado.
Two breeding pairs were "recycled" to double the
number of eggs they produced. Although subsequent natural events caused
the nests to fail, the technique proved successful.
Four additional sites
were manipulated by replacement of wild eggs with dummy eggs and subsequent
replacement with broods of captive produced young. The two techniques
succeeded in doubling the natural production.

��-165-

REINTRODUCTION
PEREGRINE

AND AUGMENTATION
FALCON

Gerald

OF

PRODUCTION

R. Craig

Current breeding populations and reproductive success of peregrine falcons in
Colorado are so low that activities must be undertaken to increase the number
of young produced or released into the wild.
Poor natural production can be
boosted by three techniques, the first of which is termed "recycling or double
clutching".
This procedure has been developed and proven with captive peregrines and should be applied to wild peregrines.
The technique takes advantage of the inherent ability of nesting peregrines to recycle and lay a
second clutch of eggs should the initial clutch be destroyed within a certain time period.
The second technique is to place captively produced young
with wild adults to supplement poor production.
The third approach is to
release captive produced peregrines into the wild through a technique known
as "hacking" whereby the young are reared and released at potential nesting
sites in hopes they will return at maturity and breed.
The three techniques
afford the manager a variety of ways to react to specific problems at particular nest sites and salvage the population from extinction due to poor or
no natural recruitment.

P.N. OBJECTIVES

The objectives

of the study are:

1.

To induce production of a second clutch of eggs from wild breeding
peregrines through "double clutching or recycling".

2.

To augment
in nests.

3.

To release additional young to the wild by "hacking"
young from abandoned or potential eyrie sites.

4.

To monitor the results of the efforts, compile data and submit reports
to appropriate state and federal agencies and the Rocky Mountain/Southwest Peregrine Falcon Recovery Team.

poor natural

production

SEGMENT

1.

by placing

captive

produced

young

captive-produced

OBJECTIVES

Breeding pairs of peregrines will be observed to determine dates of
initiation of egg laying.
Within a week to 10 days after completion
of the full clutch of eggs, the eyrie will be visited and all the
eggs removed and artificially incubated.
Approximately
two weeks after
removal of the eggs, the pair will recycle and lay a second clutch.
The
second clutch mayor
may not be replaced with dummy eggs which the adults
will be permitted to incubate.
Dummy eggs should be substituted in situations where there may be concern about the adults' ability to incubate
the eggs without breaking them.
After a suitable period (if dummy eggs
were substituted), the site will be revisited and the dummy eggs will be

�-166-

replaced with chicks from the eggs which were incubated and hatched in
captivity.
If the adults were permitted to hatch their own eggs, they
will be permitted to continue to rear and fledge them.
Since there will
be a 28 day difference in age between the young produced from the first
clutch and those produced from the second clutch, the young from the first
clutch will be placed in other wild eyries containing similarly aged broods.
Several representatives
may also be retained for captive propagation
purposes if similarly aged broods cannot be located.
2.

As in 1, breeding pairs will be kept under surveillance to determine
initiation of egg laying.
Shortly after completion of theclutch, the
eyrie site will be visited and all the eggs removed and replaced with
dummy eggs which the adults will be permitted to incubate.
Since the
wild eggs usually are thin-shelled they will be artificially incubated
to avoid their being accidentally crushed by the adults.
After several
weeks, the site will be revisited and young peregrines will be exchanged
for the artificial eggs.
Up to four young may be placed at each site
to assure the maximum number of young are fledged.

3.

Captive produced young may be released at unoccupied or potential sites
without the benefit of protection or care from adults through the technique of "hacking".
Young falcons of three to four weeks of age will
be placed on a suitable ledge at a potential reintroduction
cliff site.
They will then be cared for and fed by human attendants until they are
flying and capable of feeding themselves.
In this manner, the young
falcons will return to the site at which they were reared and hopefully
breed.
This approach requires constant attendance and observation in
order to protect the vulnerable young and insure they have sufficient
food while they are in the eyrie.
Because of this, the first two techniques will receive priority attention.
If there are no additional
adult breeding pairs as required by approaches 1 and 2, then young will
be placed into the wild using this technique.

4.

Compile and submit reports to appropriate state personnel, federal agencies and the Rocky Mountain/Southwest
Peregrine Falcon Recovery Team.

METHODS

AND MATERIALS

Segment objectives 1, 2 and 3 describe the methods of augmentation and reintroduction.
A sufficient number of active peregrine nests were available
for manipulation
to absorb all the young produced in captivity in 1977.
Reintroduction
activities were delayed, therefore, until the future when
additional young are available for release through the "hacking" technique.
Eggs were incubated in portable incubators while in transit from wild nests
to more permanent facilities at the Peregrine Fund's project at Fort Collins,
Colorado.
All eggs were incubated at the facility and young produced from
the eggs were subsequently reared there as well.

�-167-

RESULTS

AND DISCUSSION

Recycling

Efforts

Sufficient field data was obtained at two peregrine nests to permit the
recycling process to be implemented.
Following is a description of the
activities which took place at sites 6 and 26.
Site 6
The first egg was laid on April 9 and an egg was laid every other day until
the fourth and final egg was deposited April 15. The nest was visited on
April 17 and the entire clutch was removed.
The pair at this site have a
history of nesting failure early in incubation (probably a result of the
shell-thinning process), so it was decided to remove their eggs almost
immediately after the clutch was completed.
The pair resumed courtship and
copulation and selected another ledge on the cliff face in the immediate
vicinity of the first nest attempt.
Egg laying was initiated April 29 and
the final four egg clutch was removed May 10 and replaced with dummy eggs.
The pair was permitted to incubate the dummy eggs until June 6, when their
natural eggs began to hatch at the Fort Collins facility.
On June 6, the
four dummy eggs were removed and replaced with two 14 day old prairie falcon
chicks.
This was done to further delay the adults until young peregrine
falcons of the proper age were available to be placed in the wild.
The
adults immediately began feeding and brooding the prairie falcons.
On
June 23, two 12 and 14 day old peregrine falcon chicks were exchanged for
the older prairie falcon chicks.
The adults accepted the change without
apparent concern.
The adults cared for the two young until June 27 when the
male
suddenly disappeared.
It appeared that the adult male was not caring
for the young adequately and the decision was made to remove the young on
June 28. Both young were placed in a wild nest in New Mexico which contained
only two young.
All four fledged successfully.
At site 6, two of the four eggs in the first clutch were cracked and did
not hatch despite special handling and high humidity conditions in an
attempt to reduce water loss.
The other two eggs hatched, but one of
the young died shortly afterward.
Three young were hatched from the second
clutch of four eggs and all survived to be placed into the wild.
Site 26
The first clutch of four eggs was removed on April 21. The stage of egg
development placed completion of the clutch and subsequent initiation of
incubation at April 21. The first egg of the clutch was probably laid
April 14. The pair moved to another cliff approximately
1/4 mile away,
recycled and laid its second clutch May 15. The site was visited May 19
when only two eggs were discovered.
Both eggs were replaced with dummy
eggs which the adults accepted and incubated.
The site was again visited
June 9 and replaced the dummy eggs with two 16 day old prairie falcon chicks.
The adults accepted them and cared for them until June 20 when they were
replaced with two 16 day old peregrine chicks.
The adult was present and
defended the nest, but the female was not seen.
The site was observed the
first week of July, but no falcons were seen.
On July 21, the nest was
visited and the remains of partially feathered young were discovered.
They
were approximately
25 days old when they died.
Neither adult was present.

�-168-

The four eggs in the first clutch all died early in incubation.
Both eggs
in the second clutch were infertile and one egg had a dent in the side and
the contents were leaking out.
Although neither site 6 or site 26 succeeded in fledging young the recycling
attempts were successful.
The adults readily adjusted their behavior from
incubating eggs to caring for young when the dummy eggs were replaced with
young falcons.
Failures of both sites can probably be attributed to the
death of at least one of the mates (possibly both at site 26) and remaining
adult's inability to care for the young.
Augmentation

Efforts

Four other sites (sites 7, 9, 15 and 27) were manipulated by replacing wild
clutches with dummy eggs, then subsequent replacement with captive produced
and wild young.
Unlike the recycling project, the wild pairs were not
encouraged to renest and lay a second clutch.
Simple replacement of existing clutches was implemented when pairs were encountered that had been incubating eggs in excess of ten days. After an interval of 10 days, the likelyhood of renesting is greatly reduced and generally not worthy of the risk.
Artificial incubation and augmentation were unqualified success at each of
the four sites. Young prairie falcons were first placed under the adults
to be certain the adults would not react negatively and also to maintain
adults at the nests until young peregrines could be placed in the nest.
Due to remoteness of one site, young peregrines were placed in the nest
without the benefit of prior experimentation with prairie falcons.
As
expected, the adults readily accepted them. When the nests were visited
and the dummy eggs were replaced with young falcons, several dead quail
were also left on the nest ledge.
In every case, the adults returned
after the climber departed and began feeding the young.
Table 1 summarizes the assumed productivity of the six sites.
It is possible
to know exactly what the productivity would have been had the sites not been
manipulated, but certain results can be assumed.
As an example, site 6 would
not have produced the second clutch, but may have been successful in hatching
two young.
This may be questionable since two of the eggs were cracked and
it is possible the female may have abandoned the entire clutch if egg contents had matted her breast feathers.
But, on the optimistic side, we can
assume that she would have hatched two young, but these would have died when
she late~ disappeared.
An optimistic estimate of natural reproduction for
the 6 sites would be 3.3 eggs per pair, 1.83 young produced per pair and .67
young fledged per pair.
Table 2 summarizes the actual productivity of the managed sites. As can be
seen, productivity was doubled, 4.33 eggs were laid per pair; 2.33 young
were produced per pair; and 1.33 young fledged per pair. The number of
young returned to the wild increases to 2.0 per pair when one considers the
two young that were moved from site 6 to the New Mexico nest. Despite loss
of reproduction from sites 6 and 9, an adequate number of young were produced
to push reproduction above the 1.25 young per pair which is normally considered necessary to sustain a stable population.

�-169-

Table 1.

Assumed natural productivity

of managed

sites.ll

Site

No. eggs which
would have been
produced

No. young which
would have
hatched

No. young which
would have
fledged

6

4

2

0

7

3

3

3

9

3

2

0

15

3

2

1

26

4

2

0

27

3

0

0

Totals

20

11

4

3.3 eggs per pair; 1.83 young per pair; .67 young fledged per pair.

II

Natural production is estimated
not been manipulated.

Table 2.

Actual productivity

for managed

of managed

sites as though they had

sites.

No. young
retur~edll
to palr -

No. young
fledged
by pair

No. young
returned
to wild

2

0

4])

3

3 (1 c. p ,)

3

3

3

2

2 (2 c. p .)

0

0

15

3

3

3 (3 c.p.)

3

3

26

6

0

2 (2 c. p .)

0

0

27

3

0

2 (1 c.p .)

2

2

Totals

26

14

14

8

12

Site

No. eggs
produced

No. young
hatched

6

8

6

7

3

9

4.33 eggs per pair; 2.33 young produced per pair; 1.33 young fledged per
pair; 2.0 young returned to wild per pair.

II

Where indicated by (c.p.), captive produced young augmented

production.

21 The adult female disappeared shortly after the young were placed in the
nest, the young were subsequently

Prepared

by

~

v&lt; ~

Gerald R. C~aii
Sr. Wildlife Biologlst

removed and in another nest in New Mexico.

��January

-171-

1979

JOB PROGRESS REPORT

State of

COLORADO

Project No.

W-124-R
-------------------v
Work Plan No.
Job Title

Peregrine

PeriQd Covered:
Personnel:

Raptor Investigations
Job No.

4

Falcon Habitat Protection

Activities

April 15, 1977 through February

Marcy Cottrell,
of Wildlife.

in Colorado

28, 1978

Gerald Craig, and Ann Potter, Colorado Division

ABSTP~CT
An occupied peregrine falcon nest was kept under surveillance through joint
efforts of the U.S. Forest Service and Colorado Division of Wildlife.
The
two agencies entered into a Memorandum of Understanding to manage the site
and provide protection to the falcons. As well as preventing trespass, the
observers gathered behavioral information about the falcons.

��-173-

PEREGRINE

FALCON HABITAT PROTECTION

ACTIVITIES

IN COLORADO

Gerald R. Craig
During the breeding season, peregrines are vulnerable to disturbance from
human interference at the nesting cliff.
If the falcons are agitated during
critical reproductive periods, eggs or young may chill or the site may be
abandoned.
While many sites are remote, several locations in Colorado
receive excessive human visitation and it is necessary to assign personnel
to observe the sites, note disturbances which occur and limit harassment or
unnecessary activities in the area.
P.N. OBJECTIVES
The objectives

of this study are:

1.

To reduce or eliminate human disturbances
sites.

occurring

at specific

eyrie

2.

To protect important feeding areas or eyrie sites located on private
land through cooperative agreement, lease, purchase or exchange of
estate.

3.

To prepare reports of the results of the endeavors.
SEGMENT OBJECTIVES

1.

Two observers will be stationed at eyrie sites and will keep them under
constant surveillance from initiation of incubation until after the
young have fledged.
The observer will be situated in such a position
that his presence will not disturb the falcons and yet he will be
capable of viewing the vicinity and note any intruders.
The observer
should be present at the site during the daylight hours.
If possible,
the observer should have access to a packset radio to communicate with
Division of Wildlife field personnel should assistance be required.
This work will also be coordinated with the Special Agent-in-Charge at
the Denver Federal Center.
In addition to maintaining surveillance,
the observers will be expected to keep field notes of various activities
of the breeding peregrines.

2.

When important feeding areas of peregrines are located on private lands,
an effort will be made to contact the landowner and negotiate to assure
that the area remains suitable as a hunting area for peregrines.
In
general, the landowner will be encouraged to continue to provide the
habitat types preferred by key prey species, continue to plant crops
which support the prey or undertake other activities which continue to
benefit the peregrine's prey. No funding request for this activity is
included at this point since it is hoped that negotiations may be accomplished without disbursement of funds to the landowner.

�-174-

3.

Prepare reports of the activities accomplished under this job and provide them to the appropriate state and federal personnel and the
Rocky Mountain/Southwestern
Peregrine Falcon Recovery Team.
MATERIALS

AND METHODS

Two observers were employed jointly by the U.S. Forest Service and the
Colorado Division of Wildlife to keep an active peregrine eyrie under surveillance from April 18 through August 31, 1977. The site was kept under
surveillance during daylight hours from several vantage points which provided views of the surrounding area and nesting cliffs. The observation
points were located at a sufficient distance from the nest site to avoid
disturbance of the falcons.
Binoculars and spotting scopes were frequently
used to scan the vicinity for intruders, although it was soon discovered
that the falcons were the best watchdogs and would vocalize and dive at
intruders in the vicinity of the nesting cliff. The observers were also
equipped with radios to contact local Wildlife Conservation Officers or
Forest Service field personnel in case assistance was required.
In addition to protecting the falcons from direct human harassment, the
observers recorded the falcons' behavior throughout the breeding season.
Particular effort was made to record any response the falcons made to
man-related disturbances.
RESULTS AND DISCUSSION
The peregrine nest site near Chimney Rock in the San Juan National Forest
poses several unique problems.
The first is that a pair of peregrine
falcons have occupied the site since prior to 1943. Second, during a period
of 9 years, the falcons were not present and the Forest Service began development of Indian ruins adjacent to the site. Plans were underway to continue
development and expand access to permit tours of the ruins when the falcons
returned to their historic nest in 1973. As a result of their presence, the
Forest Service curtailed further development of the tours because of potential
disturbance to the falcons.
This caused much concern and focused public
attention on the falcons.
In 1976, the Forest Service and Division of Wildlife decided it would be prudent to station two observers to discourage trespass and offer some protection to the falcons while management plans were
formulated and appropriate land closures implemented.
In March of 1977, the Forest Service and Division of Wildlife consumated a
Memorandum of Understanding (Appendix A) concerning management of the
Chimney Rock site. The agencies agreed to meet annually to discuss and
approve activities proposed in the vicinity which might impact upon the
falcons or their hunting areas. Action plans will be developed which will
afford protection of the falcons and hopefully will permit development of
the ruins.
Supervised experimental tours of the ruins were conducted periodically in
July and August.
The tours provided the opportunity to monitor responses
of the falcons, yet terminate the activity if the falcons responded

�-175-

negatively.
While another season will be required, preliminary results
indicate that the falcons will tolerate some tours toward the latter portion of their breeding cycle. The frequency of the tours appears to be
more important than the number of people present, although the amount of
noise generated may also be a limiting factor. Additional experimentations
will be required to develop and refine guidelines for the tours.
In 1977, the observers recorded less than half a dozen encounters with
trespassers and all were sightseers who departed when requested.
It is
likely the number of encounters will increase as the presence of the
falcons becomes better known.
Probably the greatest disturbance to the
pair occurs when biologists visit the nest to remove eggs and augment
production.
While the visits are held to a minimum of less than 30
minutes, the falcon's normal routine may be upset for the entire day.
The disturbance underscores the need to avoid unnecessary excessive
visitations to the nests.

Prepared

by

GAhnM
P ~
Ger;l([ R: cr:~ig
Sr. Wildlife

Biologist

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                  <text>April

JOB PROGRESS

State of

COLO~~O
---------=~~~=------------

Project

No.

1979

REPORT

Game Bird Survey

W-37-R-32

22
Job No.
------------------------------~Evaluation of Nesting Cover Preferences
of
Job Titl e _~_P~h.Ee~al£s~aunJ-!t:..is2.......l!..·
nU-R!)£e"'"lo.!la~t~i:,.J,oL!n!_t~o:L.._Wh!!!.!se£!a:..!:t:..._fF~a;Jr!o
n!,llig--11M~e:..!:t:..!..h~o~d~s~
_

Work

1

Plan No.

Period

Covered:

Personnel:

April

1, 1978 through

Jack Corey,

LeRoy Haeffner,

March
Kirk

31, 1979
Snyder

and Warren

Snyder.

ABSTRACT

The program narrative was revised and submitted for approval following extensive literature review and contacts.
A study area was selected and a cooperative agreement with the lessee and management personnel was arranged which
included lessee application of treatments on the Division property and on
additional private acreage.
First and second year treatment plots were
selected, measured and herbicides were applied to first year treatment plots
in mid-August 1978 by the lessee.
Pheasant trapping resulted in instrumenting and releasing 29 hens in late February and early March 1979.
Limited
monitoring was initiated and additional equipment was constructed and put
into use. Height-density
sampling of wheat stubble and other residual cover
was initiated.

��-3-

EVALUATION OF NESTING COVER PREFERENCES OF
PHEASANTS IN RELATION TO WHEAT FARMING METHODS
Warren D. Snyder
P. N. OBJECTIVE
1.

To document the relative importance of wheat stubble, green wheat, and
other vegetative cover for (1) pheasant nest site selection, and (2)
successful production of young in the wheatlands of northeast Colorado.
Other variables pertinent to understanding the basic ecology of the
nesting pheasant in the Tablelands include determination of primary
limiting factors to reproduction and brood survival.

2.

Upon documentation that pheasants use wheat stubble extensively for
initial spring nesting and that adequate sample sizes can be obtained in
the primary study area, a second primary objective will be to determine
if mini-till summer fallow, using combinations of post harvest applied
herbicides and sweep tillage, increases pheasant nesting success when
compared to conventional summer fallow methods.
The economic and
practical aspects of mini-till farming will also be determined as they
apply to the farmer and to Division of Wildlife properties.
SEGMENT OBJECTIVES

1.

Literature

review,

interviews

and coordination

2.

Selection

of Study Area.

3.

Design plot layout and apply treatments.

4.

AnaLysis

meetings.

and evaluation.
METHODS AND MATERIALS

1.

Literature

Review,

Interviews

and Coordination

Meetings

Literature pertinent to this study was reviewed including information on
farming methods and telemetry procedures and equipment.
Contacts with
agricultural research personnel from Colorado State University and its
Extension Service, the U. S. Soil Conservation Service, and the U. S. Agricultural Research Service were made along with discussion and field tours
with farmers and herbicide applicators.
Meetings and contacts to select the
location, to develop study design, to coordinate application of treatments
and to select monitoring equipment were conducted.
2.

Selection
Primary

of Study Area

criteria used in selecting

a study location

a.

Its proximity

to the research headquarters.

b.

Its yearlong

containment

of a resident

included:

flock of pheasants.

�-6-

Tree and Shrub Rows
1 - F - M - 8.25 ac ,
---------1 - F - .C - 8.25 ac •
.!,.;.--,-

2 - F - M - 7.2 ac.
- F - C - 7.2 ac.

Tree and

Shortgrass
Pasture
Potential
4 - S - C &amp; !Ii

(Undefined)

r - _4_-_ ..: _

'- :'h:::t- 1.0 ac.

l

I

Tall - 11 .5 ac.

I

4 - F- M -

:

4 - F -C - Tall - ~ .5 ac ,

1- -

-

1_-

- '- -

_
-

I

- - -l

4 - F -C - Short -\ .0 ac ,

,

I
I

,

I
I-

.J

Fig. 2
The Sand Draw Intensive study Area (West ~ of Sec. 19).
First year (F) and Second year (S) Conventionally tilled (C) and
Mini-tilled eM) plots and their respective acreages are illustrated.

�-7-

necessitate use of the contact herbicide, glyphosate (Roundup) applied by
boom type sprayer at the rate of ~ gallon per acre. The herbicide 2,4-D
Amine will be used in substitution at a rate of 1 pound per acre when broadleavedforbs
alone are present.
These treatment options will be used on
Division property and the lessee will select a contact herbicide of his
choice for use on private land.
b.

Conventional

Summer Fallow Tillage

Treatments were not enacted on the first year stubble plots during this
segment.
The lessee will initiate stubble ground tillage using an offset
disc in late April or May, timed to coincide with conventional summer fallow
peak activity in the surrounding region.
Repetitious tillage will be
continued as needed through the summer up to wheat sowing time.
4.

Analysis
a.

b.

and Evaluation

Trapping

and Monitoring

of Nesting

Hens

1.

Hen pheasants were trapped using night lighting techniques
described by Hoffman (1975). Two initial birds were trapped in
southwest Phillips County and released on the site while testing
equipment and training personnel.
The remainder were obtained
within the extensive study area. Trapping was initiated in late
February 1979 and completed during the first weekend in March
during intervals when the ground froze and permitted access.

2.

Captured hens were equipped for monitoring with solar-powered
backpack type transmitters, constructed by Wildlife Materials Inc.
and weighing approximately 15 grams.
Flat ~ inch wide nylon
elastic material was used in harnesses constructed by procedures
similar to those described by Brander (1968).

3.

Monitoring procedures followed those described by Kuck (1968) and
others.
Two 30-ft permanent towers were erected on high points
of the Sand Draw property and were located approximately 3/4 mi
apart.
Each contained a pair of 8 ft yagi high-gain antennas.
In
addition, a single 8 ft antenna was mounted in the back of each of
two pickups.
These were mounted to telescope upward to a height
of approximately 16 to 20 ft when fully extended.
Two hand-held
three-element antennas were also used in foot and vehicular searches
and monitoring.
Two 24 channel receivers were used in conjunction
with these antennas.

Environmental
1.

Monitoring

Height-density measurements were initiated in March 1979 to obtain
indices of cover quality on all residual cover within and proximal
to the intensive study area.
Samples were obtained on perennial
herbaceous cover, roadsides, and wheat stubbl~
The Robel et ale
(1970) method as modified by Kirsch (1977) was used on all residual
vegetation and will be repeated in subsequent years.
Samples were
obtained diagonal to the direction of drill rows in wheat stubble
fields.
Recording forms were prepared to permit obtaining 100

�-8-

samples at 25 points in each field.
stubble height samples were obtained
was also obtained in each field.

In addition, 25 vertical
and the drill row spacing

2.

Precipitation

gauges were set up on the study site in March 1979.

3.

Mapping of vegetative cover types and edges was initiated but
not completed in March 1979 using eight inch to the mile aerial
photos and a grid system for calculating acreages and distance
(Snyder 1966).
Study Area

The Division of Wildlife's Sand Draw Wildlife Area was selected as the primary
work area for this nesting evaluation study. The Sand Draw property contains
210 acres, comprised of the NW~ and the SW part of the NE~, S19, TlON, Range
43W in southeast Sedgwick
County, Colorado.
The elevation approximate 3,750
ft and annual precipitation averages 16 to 17 inches.
The average growing
season is 143 days.
Soils are of the Rago-Richland-Kuma
associations being
deep, nearly level and of loam texture (Brubacher and Moore 1969). An earth
constructed dam backs water over several acres of land on the site (Fig. 1)
when runoff water occasionally fills the reservoir, however, the site is dry
a majority of the time. This pond is located on the Sand Draw drainage which
drains the locality.
Dryland wheat farming predominates in the nine section extensive study area
(Fig. 1). Several deep irrigation wells have recently been installed by
neighboring
farmers to use for irrigating corn under center-pivot systems.
Only three occupied residences are present in the nine sections attesting to
the low human population and large farm size in the locality.
Pheasants are the predominant wildlife in the region, however, the Sand Draw
Property was planted extensively to shrubs and trees in 1949 which now
permits part time occupancy by mule deer and bobwhite.
Mourning doves and
cottontails use the site extensively and waterfowl are occasional visitors
when water is present.
Coyotes, red fox, great-horned owls, striped skunks,
badgers, raccoons, crows and magpies are the primary predators in the vicinity.
Feral cats and dogs also use the site and many non-game species reside at,
or visit the property.
RESULTS AND DISCUSSION
An agreement was made with the lessee, Gary Schuler, with approval of management personnel, wherein the owner's share of wheat harvest from the Sand
Draw would be paid to the lessee in compensation for his (1) treatment of
an additional acreage on his private land using prescribed mini-till procedures, (2) extra efforts expended in using two different farming methods on
the small fields within the Sand Draw, and (3) use of his private lands for
monitoring by Division personnel.
Environmental

Conditions

Precipitation averaged below normal throughout 1978 in the study region so
that the late, deep tillage of wheat stubble on the Sand Draw, required to

�-9-

bury a growth of downy brome, permitted excessive drying and little soil
moisture storage prior to wheat planting in September.
Fortunately, approximately one inch of rain was received in late August approximately one week
after the August 18 application of herbicide to the treatment plots.
This at
least partially incorporated the atrazine into the top soil where it could
act as a pre-emergent to suppress weeds and volunteer wheat.
However, little
additional precipitation was received until late October 1978 when an approximate half-inch germinated the seeded wheat on the second year plots.
Tree and shrub rows on the Sand Draw property protected these narrow fields
so that light snows that followed in November and December remained in
place protecting the young wheat plants to a degree from the record breaking
prolonged winter cold temperatures that persisted into February 1979. In
contrast, the light snows blew off the majority of the larger wheat fields
on adjacent private lands and increased winter loss was evident.
The impact
of this on selection of a second year treatment and control sites is uncertain
at this time.
Fields treated with atrazine in August 1978 are illustrated in Fig. 2.
Regrettably, of the approximate 22.5 acres treated on private land to the
south of the Sand Draw, approximately 11 acres contained short stubble
atypical of stubble in other fields. This acreage would be very marginal
for nesting.
Therefore, the treated stubble and the cultivated controls were
divided into two segments each, to distinguish between short and tall stubble
in analysis and evaluation.
Hen Pheasant Monitoring
Thirty hens were trapped, instrumented with transmitters and released onto
the study area. One of these was killed by a prairie falcon at the time of
release, however, the transmitter was recovered and placed on a second hen.
One transmitter malfunctioned prior to placement in the field so 29 hens
were released for monitoring.
All were located with tracking equipment at
least once during the remainder of March and most were located several times
in spite of malfunctioning rental receivers and extended periods of rain,
snow and high winds.
All birds remained in the vicinity of Sand Draw even
though
the first two released had been imported.
Observations indicated
the transmitter equipped hens probably comprised less than half the hens
resident in the study area.

LITERATURE

CITED

Brander, R. B. 1968. A radio-package
Manage. 32(3):630-632.

harness

for game birds.

J. Wildl.

Brubacher, J. K., and T. R. Moore.
1969. Soil Survey of Sedgwick County,
Colorado.
U.S.D.A. Soil Conservation Service.
U. S. Gov. Printing Off.
Washington, D. C. 61 p. + append.
Hoffman, D. M. 1975. Pheasant mortality investigations.
Game Research Report, April:5-35.
Wildlife.

Colo. Div. of

�-10-

Kirsch, L. 1977 (Rev.). Instructions for use of height density pole for
measuring residual vegetation on grassland wildlife habitats.
2 p.
Mimeo Rept.
Kuck, T. 1968. Movements and behavior of pheasants during the breeding
cycle as determined by radio-tracking.
M. S. Thesis, South Dakota State
Univ.
73 p.
Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert.
1970. Relationships between visual obstruction measurements and weight of grassland
vegetation.
J. Range Manage. 23(4):295-297.
Snyder, W. D. 1966. A technique for mapping wildlife habitat in farmland
areas.
Colo. Dept. of Game, Fish and Parks.
Game Infor. Leaflet #43.
1 p.

Prepared

__
by _.L;fi~, U::::.-.;:·~-=:t,-"'}::...:'iiI~-=:v~)--"~=--:=l~~..::..{.::::".~,,,,,'''-''--7.---====-)
Warren D. Snyder
,/
Wildlife Researcher C

�April 1979
-11-

JOB PROGRESS

REPORT

State of

COLORADO

Project No.

W-37-R-32

Work Plan No.

3
Job No.
9a
Evaluation
of the Effects--o'f~C~h~a-n--g-e-s--l~'n------------------Hunting Regulations on Sage Grouse Populations

Job Title
Period Covered:

Game Bird Survey

April 1, 1978 through March 31, 1979

Personnel:
J. Dingman, Univ. of Denver; R. A. Ryder, Colo. State Univ.;
D. Benson, C. Braun, A. Chappell, D. Covic, C. Crawford, S. Emmons, K. Giesen,
D. Gore, J. Hobbs, J. Jackson, S. Palm, B. Petersen, S. Porter, B. Quinlan,
H. Spear, J. Wagner, Colorado Division of Wildlife.

ABSTRACT
Investigations concerning the relationships of hunting to breeding population
levels, and survival estimates of sage grouse (Centrocercus urophaisanus) in
North Park, Colorado, initiated in 1975, continued in 1978. Numbers of known
leks increased as 7 new leks were located.
Average number of cocks per lek
increased 27% over 1977. Total estimated spring population size was between
5,469 and 8,754 birds.
In 1978, 258 sage grouse (204 males, 54 females)
were banded.
A 16 day hunting season was allowed in North Park in 1978 with
bag and possession limits of 3 and 6. Data from wings (N = 724) collected
at check stations, wing barrels and field checks revealed that the harvest
was comprised of 53% young of the year, 20% yearlings and 27% adults.
Estimated annual survival from wing analysis was 65% for females and 50% for
males.
Estimated nesting success from examination of wings of adult and
yearling females was 51% with more adults (60%) than yearlings (38%) being
successful.
Ovarian analysis (N = 82) indicated that 92 (yearlings) to 98%
(adults) of all females ovulated.
Total estimated harvest based on check
station data was 1,200-1,400 birds.
Sixty-three bands were reported by
hunters.
Direct recovery rates varied from 10.1 (adults) to 13.2% (yearlings)
for males.
Banded samples of females were too small for calculation of age
specific recovery rates. The overall direct recovery rate for females was
10.7%. The constant annual survival estimate for all females was 63.4 + 6.1%,
for adult males it was 43.5 + 6.3%, while for yearling males it was 85.3 +
15.7%. The 95% confidence i~terval was narrow (24% range) for adult males
and all females and broad (61% range) for yearling males.
Estimated fall
population size was 12,620 - 16,460 birds.

�-12-

RECOMMENDATIONS

1.

The project should be extended one year to allow for completion of data
collection concerning lek attendance patterns of male and female sage
grouse.

2.

Banding should be de-emphasized for estimating survival.
It should
continue to be used as a tool to estimate changes in harvest rate between
areas in North Park.

3.

Counts of male sage grouse present on leks should continue with emphasis
from mid April to mid May.

4.

Three check stations should be operated the opening weekend of the sage
grouse hunting season in 1979 along with volunteer wing collection barrels
when check stations are not open. The new check station would be near
Gould.

5.

The sage grouse season in North Park in 1979 should be a minimum
days with bag and possession limits of 3 and 6.

of 23

�-13-

EVALUATION
OF THE EFFECTS OF CHANGES IN
HUNTING REGULATIONS ON SAGE GROUSE POPULATIONS
Clait E. Braun

Knowledge of the impacts of hunting on sage grouse is limited, as no studies
on this subject have been published.
Impacts of hunting can be measured
through examination of changes in numbers of birds present on leks in spring
if lek counts have a relationship to population size, and through analysis
of ban dings (harvest rate). Data presented in this report concern these
problems and represent the mid ~arct 1978 through mid March 1979 interval.
P. N. OBJECTIVE
The objectives of this study are to increase hunter opportunity and recreational use of sage grouse in North Park, Colorado.
Hypotheses which are
being tested are:
a.

Hunter harvest does not affect spring counts of strutting
spring counts of females under any regulations.

b.

Counts of females during the peak of attendance on strutting grounds
a reliable index to overall numbers of sage grouse in spring.

c.

Hunter success and total harvest

d.

Nesting success and brood size to 15 August, while important in determining age composition of the harvest, have no relationship to hunter
success and total harvest.

e.

Differences
and nesting

are a function

exist between yearling
success.

of August

males or peak

are

precipitation.

and adult hens in percent

ovulating,

SEGMENT OBJECTIVES
la. Twenty males will be captured and radio transmittered in March and early
April in northwestern North Park. Transmittered males will be stratified
by age (1- and 2+) and will be followed daily until early June.
lb. Twenty females will be captured and radio transmittered in March and
early April in southeastern North Park. Transmittered females will be
stratified by age (1- and 2+) and will be followed daily until early June.
2.

Counts of all birds present will be made at least 4 times from 1 April
to 15 May on all known strutting grounds in North Park.

3.

Aerial and ground searches will be utilized
grounds within North Park.

4.

Population size will be estimated in Spring through use of ratios of
males to females on strutting grounds and known sex composition of the
population
available from previous work and harvest surveys.

to locate additional

strutting

�-14-

5.

Wing collection barrels will be placed at strategic locations throughout
North Park and will be available throughout the hunting season.
A goal
of 500 wings is desired.

6.

Number and location of marked birds harvested will be obtained
field checks of hunters and voluntary mail reporting.

7.

Age composition
wings collected
barrels.

8.

Compile

through

of the harvest will be determined through examination of
during field checks and through use of wing collection

data, analyze

results

METHODS

and prepare final reports.

AND MATERIALS

Counts of male and female sage grouse present on leks were periodically made
by project personnel and District Wildlife Managers following prescribed
procedures (Braun and Beck 1976). Counts were made from 19 March through
30 May at times within one-half hour of sunrise.
Searches for new or relocated
leks were made through ground and aerial reconnaissance.
Male and female
sage grouse were trapped at night along roads and on leks where they roosted.
Some birds, primarily hens, were captured at dawn on leks through use of
cannon nets. Most birds were captured through use of a spotlight with a back
pack or hand held power source and long handled nets (Braun and Beck 1976).
Sage grouse banded and released were marked with serially numbered aluminum
bands (size 16 for males, 14 for females) and colored plastic and aluminum
bands coded to year.
Some birds were additionally marked with radio transmitters.
Age and sex classification
of birds captured followed Eng (1955).
Check stations were operated at 2 locations (State Line and Willow Creek Pass)
on the opening weekend (9 and 10 September) and the second Sunday (17 September).
Each station was operated from about 1000 to 1800 MDT, depending upon
traffic load. Data obtained per party were: County of origin, number of
hunters, hours hunted (total of all hunters), birds observed, birds bagged,
birds lost, number of banded birds and location of where each was obtained,
and area hunted.
One wing was obtained from each bird checked, with ovaries
being taken or examined from a sample of hens.
These data were recorded on
tags with wings being individually marked with corresponding information
concerning actual sex of that bird.
Wings were frozen and stored for later
analysis.
In addition to the 2 check stations, wing barrels and signs (Hoffman and
Braun 1975) were placed along Colorado 14 near Muddy Pass, along Colorado 14
near Gould, north of the junction of Colorado 125 and 127 along Colorado 127
and at Willow Creek Pass along Colorado 125. Volunteer stations were in
operation the entire season near Muddy Pass and Gould and on all days that
manned check stations were not operating on Colorado 127 north of its junction
with Colorado 125 and along Colorado 125 at Willow Creek Pass.

,

Collected wings were thawed and classified to age (chicks, yearlings,
adults) and sex following procedures outlined by Eng (1955), and Beck et ale
(1975). Hatching dates were calculated for chicks of the year using data
from Pyrah (1963).

�-15-

_Ovaries collected were stored in AFA (alcohol, formalin and acetic acid)
and later examined for presence of ovulated follicles as described by Kabat
-.et al. (1948).
Description

of Area

North Park lies completely in Jackson County, Colorado.
This relatively
high altitude (7,800 to 8,500 ft) are~ is essentially a closed basin, as
it is encircled by mountain ranges.
Drainage is to the north, with lowest
elevations occurring north of Cowdrey.
Vegetation is principally a sagebrush (Artemisia spp.)-grassland
type with stream courses being dominated
by native grasses, sedges, and deciduous shrubs.
Detailed geographic,
-geologic, vegetational and climatic features of the area have been described
by Gill (1965), Carr (1967), Beck (1975), Braun and Beck (1976) and will be
treated in the final report.
RESULTS

AND DISCUSSION

Counts of Sage Grouse on Display

Areas

Counts of male and female sage grouse present on leks were initiated on 19
March (Denmark and Raven) and continued until 30 May (Alkali Lake, Bighorn,
Boettcher Lake Junction, Spring Creek #1 and Spring Creek #2). During this
interval 609 counts were made of 34 active leks. Average number of cocks
per lek (high count only) was 39.5 (Table 1). This is substantially higher
than the 31.1 males/lek counted in 1977 and slightly higher
than the 39.1
males/lek counted in 1968-72 (Table 2). Number of leks known to occur in
North Park increased (+7) in 1978. Two new leks were found by following
radio marked females, 3 were found from the ground, while 2 were found from
the air. One additional new lek was observed from the air west of Pole
Mountain Reservoir but was not relocated from the ground.
From 1973 to 1977 average number of cocks counted per lek remained reasonably
constant (27.7 to 33.1). During this same period, number of known leks
increased from 17 to 26, suggesting either better coverage of North Park or
an expanding sage grouse population.
In 1978, number of cocks counted per
lek increased 27.0% over the 1977 level and number of known leks increased
from 26 to 34 (30.8%).
It is quite evident that despite better coverage,
the sage grouse population in North Park has been expanding.
The expansion
in 1978 is further documented if only leks with 10 or more males are
considered (Table 3).
Estimation

of Spring Population

Size

Reliable estimates of total numbers of sage grouse in North Park during the
breeding period are difficult to derive.
Major problems involved are those
relating to how many males and females of the total population are present
on leks when peak counts are attained and how many of the actual number of
active leks are known and counted.
There is some evidence to suggest that
only 50% of the male segment of the population of lekking species is present
on a ground when peak counts of males are attained (Robel 1969, Rippin and
Boag 1974).
If it is assumed that peak counts of male sage grouse represent

�-16-

Table

1.

Peak lek counts

Males

Lek

Alkali Lake
A.rapahoe
Aspen
Bighorn
Boettcher Lake Jet.
Canuck
Case
Cheyenne 1./
Coalmont
Cowdrey 115
Deer Creek
Delaney Butte
Denmark
Eagle 1./
Fish Hatchery
Hound
Los t Creek 111
Monahan
Owl Creek
Peregrine 1./
Prague 1/
Pronghorn
Ptar 1./
Railroad
Raven
Ridge Road
Riley
Roth
Spring Creek III
Spring Creek 112
Spring Creek #4
Thrasher 1/
Ute 1./
Walden
Wattenberg
II

56
61
24
47

86
21

4
127
21
1

41
4

80
11
97
28
43

o
15
17
35
10
50
35
94

65
5
1
76

63
3
65
17
18
21

1,342

Totals
Average/Active

Lek

N =

1/ Newly

of sage grouse,

located

39.5
34

in 1978.

North

Park,

Date(s)

10 May

9 May
8 April
10 April
1 May
3 May
4 April
2 May
4, 12 &amp; 24 April
30 March, 1 May
20 April
22 April
24 April
9 &amp; 15 April
4 s 11 April
26 April
15 April
All dates
13 April, 4 May
2, 6 &amp; 8 May
27 April
7 &amp; 17 April
12 April
6 April
11 &amp; 24 April, 3 May
28 April
18 April
19 April
8 May
16 May
8 April
2 May
21 April
10 April
26 May

1978.

Females

Date(s)

20
42
10
19

3 April
8 April
8 April
4 April
4 April
10 April
12 April
22 April
12 April
All dates
5 April
5 &amp; 25 April
"7 April
8 April
11 April
10 April
6 April
All dates
8 April
20 April
9 May
8 April
12 April
6 April
7 April
12 April
10 April
All dates
4 April
4 &amp; 11 April
All dates
19 May
9 May
4 April
7 April

84

10
1
2

28

o
40
3
40
8
43
13
40

o
9

3
3
6
39
34
52
47
2

o
50
29

o
3
2

19
7

708
20.8

�Table 2.

Peak 1ek counts of male sage grouse, North Park, Colorado, 1973-1978.

Lek

Alkali Lake
Arapahoe (1977) ~/
Aspen (1977)
Bighorn (1976)
Boettcher Lake Junction
Boettcher Flats
Canuck (1974)
Coalmont
Cowdrey tI1
Cowdrey 113
Cowdrey #5 (1973)
Deer Creek
Delaney Butte
Denmark (1977)
Fish Hatchery
Hound (1974)
Lake John #2
Lost Creek 111
Lost Creek If2
Monahan Draw
Owl Creek (1976)
Pronghorn (1977)
Railroad (1975)
Raven (1977)
Ridge Road
Riley (1973)
Roth (1974)

(1968-1970)1/

(1968-1971)
(1968-1971)

Average
1968-1972

1973

1974

1975

1976

1977

1978-V

77.8

81

72

68

39

-

-

-

-

-

-

86.6
7.0

59
0

-

-

39.4
9.2
5.0

47
0
0
30
37
4

46
0
29
27
0
0
19
11
13

50
0
22
29
0
0
9
27
0

36
46
21
52
76
0
27
21
0
0
5
31
0
58
64
21
0
47
0
1
16
15
41
73
32
9
8

56
61
24
47
86
0
21
21
0
0
1
41
4
80
97
28
0
43
0
0
15
10
35
94
65
5
1

(1970-1972)

56.6
13.3

46
62
0
30
28
0
0
5
36
0

-

-

-

-

-

(1970-1972)

58.7

67

-

-

13.5
54.4
27.4
56.2

0
13
10
11

69
33
0
18
0
6

81
27

(1968-1971)

62
69
0
0
1
3

-

-

36
12

33
15
10

- .
76.4

-

-

86

o-

33
0
1
14

87

-

-

27
12
14

32
18
9

-----------------------------------------------------------------------------------------------------------------

I
I-'

....•
I

�Table 2.

Peak 1ek counts of male sage grouse, North Park, Colorado, 1973-1978 (continued).

Average
1968-1972

Lek

Spring Creek 111
Spring Creek 112
Spring Creek 114
Walden Reservoir (1973)
Wattenburg III
Wattenburg 112

(1968, 1970-1972)

1973

1974

1975

1.5
27.2

46
39
9
38
0
24

33
15
10
37
0
22

49
14
3
34
0
18

19

17

19

19

37.6
75.8
18.7

-

(1968-1969)

Number of Leks

, 1976

1977

'197sl/

49
8
37
0
16

45
23
3
26
0
12

76
63
3
18
0
21

21

26

34

11

I
f-'

co

Average/Active Ground

39.1

33.1

27.7

30.9

31.9

1/ Seven leks found in 1978 are not listed but are included in number of leks and averages.

31.1

I

39.5

These 7 leks are

listed in Table 1.

l/ Year of initial location if after 1972.
1/ Years of data included in average for 1968-1972 if inactive 1 or more years during that time interval.

�-19-

50% of the mal~ population in an area, then at least 2,684 (1,342 x 2) cocks
occurred in North Park in the spring of 1978. It is quite obvious that all
lek locations are not known in a given year. If it is assumed that 90% of
the active leks have been located, then a minimum of 3 (.10 x 34) unknown
leks should occur somewhere within North Park. Assuming that these 3 leks
have an average of 39 cocks during the peak period of male attendance, they
represent a minimum of 234 additional cocks (39 x 3 = 117 x 2 = 234). Thus
the total estimated number of cocks in North Park in Spring 1978 was 2,918.
This is substantially higher than the 1,928 estimated for North Park in 1977
(Table 4).
Table 3. Number of male sage grouse/stable lek, North Park, Colorado,
1973-1978. 1/
Year

Average No. Males/Stable Lek

No. of Stable Leks

1973

36.7

15

1974

30.8

17

1975

35.6

16

1976

38.0

17

1977

37.3

21

1978

47.3

28

!/ A stable lek is one that has 10 or more males.

Table 4.
1975-78.

Estimated spring population size, sage grouse, North Park, Colorado,

Year

No. of
Cocks
Counted

No. of
Leks
Counted

Estimated
No. of
Leks

Total
Male
Population

Total
Female
Population

Total
Population

1975

588

19

24

1,486

2,972

4,458

1976

712

22

26

1,682

3,364

5,046

1977

809

26

31

1,928

3,856

5,784

1978

1,342

34

37

2,918

5,836

8,754.

�-20-

Beck (1977) found a late winter sex ratio of 69 hens to 31 cocks, a value
remarkably similar to the sex ratio of adult and yearling birds in the fall
harvest (Braun and Beck 1976).
If it is assumed that the spring sex ratio
of the North Park sage grouse population is similar to that observed in
late winter and early fall, then 2 hens occur for every cock in the population.
Thus, the total spring female population
can be estimated based
on the estimated number of cocks in the spring population.
Therefore,
there should have been a minimum of 5,836 hens in the 1978 spring population
(2,918 x 2 = 5,836).
Thus the estimated total spring population of sage
grouse within North Park in 1978 was 8,754 birds (Table 4). Data are
becoming available (Emmons 1979) to indicate that more than 50% of the male
population is present on a lek if counts are made when most males are
displaying.
If peak lek counts represent 75% of the males in a population
then only 1,823 males (1,342 x .25 = 335 + 1,342 = 1,677, 3 x 39 = 117 x .25
29 + 117 = 146, 146 + 1,677 = 1,823) were present in the spring population.
This would indicate that there were 3,646 females (1,823 x 2 = 3,646) for
a total spring population size of 5,469 birds (3,646 + 1,823).
This is
37.5% less than the estimate of 8,754 birds based on only 50% of the males
being present during peak counts.
It is quite obvious that the basic
assumption concerning percent of the male population present during periods
of peak lek counts is critical.
Nevertheless, it would appear that the
method for estimating spring population size has merit.
In all probability,
the actual spring population was somewhere between 5,400 and 8,800 birds.
Capture

and Banding

Intensive efforts to capture and band sage grouse in North Park were not
made in 1978.
Sage grouse were captured and banded in conjunction with
radio telemetry studies, personnel training and preliminary design of a
new study.
In all, 258 birds were newly banded between 25 February and
10 May (Table 5).

Table 5. Sex and age composition
North Park, Colorado 1978.

Month

1-

Males
2+

February

1

March

of newly banded

sage grouse by month,

Total

1-

Females
2+

Total

Totals

0

1

0

0

0

1

2

5

7

0

1

1

8

April

78

65

143

22

21

43

186

May

25

28

53

10

0

10

63

Totals

106

98

204

32

22

54

258

�-21-

During the course of trapping activities in 1978, 28 different birds were
recaptured (27 males, 1 female).
Of this sample, 1 bird (male) was from
1975 (5+ years old upon recapture), 8 were from 1976, 16 were from 1977,
while 3 were banded in 1978. Of interest was a male banded as a chick in
1977 that was recaptured on a lek about 4 miles from initial banding site.
It did not recruit to the closest lek (about 1 mile from where banded).
No direct trap casualties occurred in 1978 although 1 bird released with a
radio transmitter was found dead the next day, apparently the result of a
crash landing upon release.
Hunting

Season Data Collection

The sage grouse season in North Park (Unit 12) in 1978 opened at sunrise
on 9 September and closed at sunset on 24 September.
Season length was
16 days, the daily bag limit was 3, with a possession limit of 6. Season
length and bag limits were identical to 1977.
Sage grouse hunters were not
required to have a permit in 1978 as the special sage grouse hunting permit
was discontinued after the 1977 season.
Check stations and field checks
were continued in 1978 similar to the 1974-1977 period.
Check Stations
Two check stations were operated in 1978. These stations were located at
the State Line on Colorado 127 and at Willow Creek Pass on Colorado 125.
As in previous years of operation (1974-1977), each station was open from
about 1000 to 1800 MDT, depending upon traffic load. Both check stations
were operated on the 9th, 10th and 17th.
Staffing patterns (at least 2
research personnel and 3 District Wildlife Managers) were the same at both
check stations during all 3 days of operation.
Data obtained per party
were: County of origin, number of hunters, hours hunted (total of all
hunters), 'birds observed, birds bagged, birds lost, number of banded birds
and location where each was obtained, and area hunted within North Park.
Most importantly, one wing was obtained from each bird that was checked.
Few birds (53) were completely wingless.
Ovaries (N = 82) were collected
from adult and yearling females for laboratory analysis.
Sex from gonadal
inspection was ascertained for 163 young of the year and whole body weights
were obtained for 29 grouse.
In addition to the 2 check stations, wing barrels and signs were placed
along Colorado 14 near Muddy Pass, on Colorado 14 near Gould, north of Three
Way on Colorado 127 and at Willow Creek Pass on Colorado 125. Volunteer
stations were in operation the entire season near Muddy Pass and Gould, and
for all days that manned check stations were not operating north of Three
Way and at Willow Creek Pass.
Research and regional management personnel
field checked hunters whenever possible.
During the 3 days of check station operation, 350 hunters with 480 sage
grouse (1.4 birds per hunter) were checked.
These hunters reported observing
4,922 sage grouse.
Undoubtedly some duplications are present in the 4,922
grouse reported observed.
Hunter efficiency was low, 9.8% (480 birds harvested ~ 4,922 birds observed), and 29 birds (5.7% of those retrieved +
those reported lost) were reported crippled and lost. Comparative data from
1974 through 1978 are presented (Table 6).

�-22-

Table 6.

Sage grouse harvest

statistics,

North Park, Colorado,

1974-1978.11

Year

No.
Hunters
Checked

No.
Birds
Observed

No.
Birds
Harvested

Hunter
Efficiency
(%)

Crippling
Loss

Birds
per
Hunter

1974

730

6,062

785

7.7

5.1

1.1

1975

738

5,735

551

9.6

7.1

0.7

1976

595

3,393

459

13.5

5. 7

0.8

1977

353

3,303

385

11.7

10.6

1.1

1978

350

4,922

480

9.8

5.7

1.4

11 Only those statistics

collected

at check 'stations.

As in other years in the 1974-1977 period, distribution of harvest and hunting
pressure was not uniform within North Park.
Peterson Ridge-MacFarlane
Reservoir (36.7% of the harvest and 21.1% of the hunters) was by far the leading
harvest area in 1978, probably more the result of opening the Arapaho
National Wildlife Refuge to sage grouse hunting than more birds being present
in the area.
The Lake John area had 29.4% of the hunters but only 18.1% of
the harvest, Walden Reservoir had 18.3% of the hunters and only 10.6% of
.
the harvest, while the Ridge Road area had 14.6% of the hunters and 13.7% of
the harvest.
All other areas within North Park each had less than 10% of
the hunters and harvest.
Comparative data are presented (Table 7). Hunter
interest and harvest continued to decline at Independence Mountain, Pole
Mountain, and Spring Creek-Owl Ridge in 1978. Numbers of sage grouse counted
on leks in spring have decreased in the Independence Mountain area in recent
years.
However, counts of male grouse on leks and number of leks have
increased in all other areas within North Park.
It does not appear that
hunter pressure parallels changes in sage grouse distribution or numbers.
Most hunters contacted at check stations were asked whether or not they
normally hunted sage grouse in North Park.
Of the 343 hunters responding,
251 (73.2%) reported they normally hunted sage grouse in North Park, 66 (19.2%)
were first time sage grouse hunters, while 26 (7.6%) reported they normally
hunted sage grouse elsewhere.
Comparative data for the 1974-1978 period are
presented (Table 8). Obviously, increasing season length and bag limits had
little effect on where hunters hunt sage grouse as 1974 was the most restrict've season (3 days, 2 and 4 bag and possession limits) in the 5 year period.
The most liberal seasons were in 1977 and 1978 (16 days, 3 and 6 bag and
possession limits).
Other areas in Colorado open for sage grouse hunting
either had shorter seasons or smaller bag and possession limits in the
1975-1978 interval.

�Table 7.

1 1
Sage grouse harvest and hunting pressure within North Park, Colorado 1974-1978. _ ~I
Spring
Peterson
Creek-Owl
Ridge- MacRidge
Far1ane Res.
%
%
Hunt.
Harv.
Hunt. Harv.

Michigan
River
SE

Eagle
Hill
%
Hunt. Harv.

Year

Walden
Pole
Independence
Mountain
Reservoir Ridge Road
Lake John
Mountain
%
%
%
%
%
Hunt. Harv. Hunt. Harv. Hunt. Harv. Hunt. Harv. Hunt. Harv.

1974

4.7

6.0

18.1

16.4

12.4

15.4

9.6

11.3

4.7

3.8

20.3

19.6

17.9

13.4

2.2

1.3

10.0

12.8

1975

3.1

3.8

24.3

24.7

14.2

16.7

11.1

15.6

1.8

2.0

17.2

14.3

13.6

6.2

2.6

3.8

10.4

9.8

1976

2.3

3.0

28.2

21.3

13.0

12.0

13.0

21.1

2.9

4.5

18.5

21.3

10.9

6.3

3.1

5.9

8.0

5.0

1977

1.4

0.5

28.9

23.9

15.8

12.2

16.9

20.8

3.2

6.5

16.3

12.7

8.0

8.1

2.0

2.1

7.4

13.2

1978

0.9

0.0

29.4

18.1

18.3

10.6

14.6

13.7

2.0

2.5

21.1

36.7

6.0

3.1

1.7

5.6

6.3

9.6

%

Hunt. Harv.

I

N

w

11 Data from check stations only.
~I Totals may not approximate 100% as each year some hunters and birds harvested could not be allocated to a particular
zone.

,

I

�-24-

Table 8. Previous sage grouse hunting
North Park, Colorado 1974-1978.

experience

of sage grouse hunters,

Year

Normally Hunt
in North Park
%
No.

Normally Hunt
Elsewhere
No.
%

First Time Sage
Grouse Hunter
No.
%

1974

183

68.3

20

7.4

65

24.3

1975

432

64.9

47

7.0

187

28.1

1976

366

64.9

40

7.1

159

28.1

1977

211

64.9

32

9.8

82

25.2

1978

251

73.2

26

7.6

66

19.2

Origin of 163 hunter parties was ascertained at check stations.
Eighty-five
(52.1%) originated in the Denver metropolitan area (Jefferson - 39, Adams - 18,
Denver - 15, Arapahoe - 13), 34 (20.9%) were from Larimer County, and 20
(12.3%) were from Boulder County.
All other counties each comprised less than
5% of the hunter parties.
These data are similar to those collected in
previous years.
Appreciable numbers of Jackson County residents (North Park)
are not checked at check stations although they normally comprise about 10%
of the sage grouse hunters in North Park (questionnaire data 1974-1977).
Analysis

of Wings

A total of 724 sage grouse wings was collected in North Park during the 1978
hunting season.
Of this total, 427 were obtained at the 2 check stations in
North Park, 12 were collected at the Green Mountain Reservoir check station
in Middle Park, 173 were obtained from wing barrels (Gould - 87, Willow Creek
Pass - 61, Three Way - 19, Muddy Pass - 6), 7 were received in mail envelopes
while the remainder (105) were obtained through field contacts.
Distribution
of the wings collected by time of collection was similar in 1977 and 1978
(Table 9).
These data strongly suggest that the objectives of increasing hunter opportunity and spreading the hunters and harvest over time are being achieved
with longer seasons.
Approximately
the same number of sage grouse hunters
hunted in North Park in 1977 and 1978 as in the 1974-1976 period, but hunter
pressure has greatly declined on opening weekend (see Table 6). These data
graphically illustrate that hunters are utilizing the longer season as more
hunters are hunting later in the season.

�-25-

Table 9. Time distribution
Colorado 1977-1978.

of sage grouse wings received,

North Park,

1977

1978

No. of
Wings

% of
Total

No. of
Wings

% of
Total

1st Weekend
(9-10 &amp; 11-77)
(9-9 &amp; 10-78)

425

67.3

448

61. 9

1st Week
(9-12 thr 16-77)
(9-11 thr 15-78)

39

6.2

103

14.2

2nd Weekend
(9-17 &amp; 18-77)
(9-16 &amp; 17-78)

83

13.1

47

6.5

2nd Week
(9-19 thr 23-77)
(9-18 thr 22-78)

33

5.2

57

7.9

3rd Weekend
(9-24 &amp; 25-77)
(9-23 &amp; 24-78)

52

8.2

69

9.5

Totals

632

100.0

724

100.0

Age and sex structure of the 1978 harvest as ascertained from examination of
wings is presented (Table 10) along with comparative data from 1974-1977.
As
can be readily ascertained (Table 10), sex composition of the harvest changed
only slightly in 1978 from previous years; there was a higher percentage of
yearling and adult males in the harvest in 1978 than in most previous years.
In 1978, young of the year comprised 53% of the harvest, the highest in the 5
years of intensive wing surveys.
There were fewer adults in the 1978 harvest
than in other years of the 1974-1977 period, indicating that the population
is increasing.
Estimated turnover (from wing analysis) of adult male and female sage grouse
in North Park was 48.1 and 38.6%, respectively, in 1978 (Tables 11 and 12).
The 5 year average for the North Park population was 49.2% for adult males
(50.8% annual survival) and 34.5% for adult females (65.5% annual survival):
These estimates are based on the assumption that in a stable population,
percent yearlings must equal annual loss of adults.
In an increasing population, percent yearlings must more than replace adult loss and/or adult
survival must be better than that estimated.
Percent yearlings (Table 10)
has been good each year since 1974, indicating that the population of sage
grouse in North Park has been expanding since at least that date.

�Table 10.

Age and sex

composition of the sage grouse harvest, North Park, Colorado 1974-1978.

Yearlings
Females
%
No.

Adults
Females
%
No.

Year

Males
%
No.

Immatures
Females
%
No.

1974

171

48.9

179

51.1

350

50.1

49

35.5

89

64.5

138

19.8

45

21.4

165

78.6

210

30.1

1975

101

47.6

111

52.4

212

42.0

52

46.8

59

53.2

111

22.0

55

30.2

127

69.8

182

36.0

42.3

46

39.3

71

60.7

117

23.5

49

28.8

121

71.2

170

34.2

Total
%
No.

Males
%
No.

Total
%
No.

Males
%
No.

Total
%
No.

I
N
(J'\

I

1976

104

49.5

106

50.5

210

1977

136

46.9

154

53.1

290

45.9

47

38.2

76

61.8

123

19.5

48

21.9

171

78.1

219

34.6

1978

184

47.8

201

52.2

385

53.2

62

43.4

81

56.6

143

19.8

67

34.2

129

65.8

196

27.1

5 Year
Average

48.1

51.9

47.3

40.5

59.5

20.7

27.0

73.0

32.0

�-27-

Table 11. Estimated turnover
Colorado 1974-1978. 1/

of adult male sage grouse, North Park.

Adults

Yearlings
Number

Year

Number

%

1974

45

47.9

49

52.1

94

1975

55

51.4

52

48.6

107

1976

49

51.6

46

48.4

95

1977

48

50.5

47

49.5

95

1978

67

51.9

62

48.1

129

50.8

Average

1/ Data

from wing collections

Adults
Number

Totals

49.2

only.

Table 12. Estimated turnover
Colorado 1974-1978. 1/

Year

%

of adult female sage grouse, North Park,

Yearlings
Number

%

%

Totals

1974

165

65.0

89

35.0

254

1975

127

68.3

59

31.7

186

1976

121

63.0

71

37.0

192

1977

171

69.2

76

30.8

247

1978

129

61.4

81

38.6

210

65.5

Average

1/ Data

from wing collections

34.5

only.

The data (Tables 10, 11, and 12) suggest
North Park sage grouse population.
approximates

the following

conclusions

about the

1:1.

1.

Sex ratio at hatching

2.

Differential survival favoring females starts in the first year of life
and is most pronounced in the older age classes.

�-28-

3.

Production of young in the 1974-1978 interval
to good but has not been excellent in anyone

has ranged from average
year.

4.

Overwinter survival of young grouse to the yearling class has been good
and has been more than adequate to maintain the population.

5.

The annual turnover rate for females is approximately
males it is approximately 49%.

6.

Females comprise
the population.

34%, while for

about 68% (67.7) of the adult and yearling

segment of

Wings from 210 females (adult - 129, yearling - 81) were classified as to
primary feather molt.
An estimate.d 59.7% (77 of 129) of the adult hens
and 38.3% (31 of 81) of the yearling females were successful nesters in
1978. This gave an overall success rate of 51.4%, similar to that calculated
for 1977. The young to adult hen (including yearling hens) ratio in the
harvest was 1.8:1, while the young per successful hen ratio was 3.6:1
(Table 13).
Table 13. Sage grouse nesting
Colorado 1974-1978.

success and production

rates, North Park,

Young per
Successful Hen

Year

Estimated Nesting Success
Yearlings
Total
Adults

Young per Hen

1974

65.4

46.1

58.7

1.4: 1

2.3:1

1975

53.2

39.0

48.6

1.1: 1

2.3:1

1976

52.9

26.8

43.2

1.1: 1

2.5:1

1977

59.3

32.7

50.3

1. 2: 1

2.0:1

1978

59.7

38.3

51. 4

1. 8: 1

3.6:1

Hatching dates were calculated for 383 chick sage grouse for which usable
wings were available (Table 14).
As in earlier years, it is obvious that the chick aging technique is not
precise.
In the real world, females probably do not hatch later than males.
Data presented in Table 14 suggest that hatching was spread in 1978 with
an obvious peak prior to 15 June. Renesting by hens that lost their initial
clutch was apparently common in 1978 as well over 10% of all chicks hatched
after 28 June.

�-29-

Table 14.

Sage grouse hatching

dates, North Park, Colorado

Males
No. Hatching
% of Total

Time Interval

1978.

Females
No. Hatching
% of Total

May 18-24

5

2.7

2

1.0

May 25-31

45

24.6

9

4.5

June 1-7

64

35.0

33

16.5

June 8-14

20

10.9

56

28.0

June 15-21

15

8.2

30

15.0

June 22-28

14

7.7

15

7.5

June 29-Jul~ 5

9

4.9

22

11.0

July 6-12

11

6.0

17

8.5

July 13-19

0

0.0

14

7.0

July 20-26

0

0.0

1

0.5

0

0.0

1

0.5

183

100.0

200

100.0

July 27-August

2

Totals

Ovarian Analysis
Ovaries were collected from 82 hunter harvested sage grouse in 1978 and were
analyzed following procedures described by Meyer et al. (1947), Kabat et al.
(1948) and Buss et al. (1951). Of the total, 79 females (96.3%) had ovulated
with a higher percentage (98.2%) of adults ovulating than yearlings (92.0%)
(Table 15). Data presented indicate that a higher percentage of adults than
yearlings ovulate each year.
Estimate

of Total Harvest

Since no permit was required of sage grouse hunters in North Park in 1978,
questionnaires could not be used to estimate total harvest outside of the
statewide harvest survey.
The inadequacies of the statewide small game
harvest survey have been documented in previous years.
For Unit 12, North
Park, it normally overestimated hunter numbers by 29% and total harvest by
85% (Table 16).

�-30-

analysis,

North Park, Colorado

1975-1978.

Table 15.

Sage grouse ovarian

Age Class

1975

Percent
1976

Ovulating
1977
1978

Yearling

84.6

75.0

96.6

92.0

33/39

33/

44

28/29

25/27

Adult

92.8

87.8

98.3

98.2

64/69

65/74

59/60

54/55

All
Females

89.8

83.1

97.8

96.3

97/108

98/118

87/

79/82

Table 16. Comparative
North Park 1974-1977.

data, number

Number Ovulating/Examined
1977
1978
1975
1976

of sage grouse hunters

89

and total harvest,

Number of Sage Grouse Harvested
Statewide Survey
This Study

Year

Number
This Study

of Hunters
Statewide Survey

1974

960

1,759

1,174

2,509

1975

1,187

1,600

1,053

1,973

1976

975

877

829

1,402

1977

842

880

1,069

1,749

Statewide Survey
Harvest = +85.0

4 Year Average

% Error:

Hunter Numbers

+29.1,

Total

Without a questionnaire survey of sage grouse hunters who hunted in North
Park and without reliance upon the statewide small game survey, it is possible
to estimate total harvest from check station data.
In 1977, 353 of the 842
hunters (permittees) (41.9%) were contacted at check stations with 385 of
the 1,069 grouse (36.0%) projected harvested (questionnaire survey of North
Park hunters) being checked.
In 1978, 350 hunters were contacted with 480
grouse.
Since season length, bag limits and check station operation were
the same in 1977 and 1978, it is reasonably safe to assume that the same
percentage of the hunters and kill were checked in both 1977 and 1978. Thus
if 350 = 42% of the total hunters and 480 = 36% of the total kill, then 100%
of the hunters and harvest in 1978 should be 833 and 1,333, respectively.
If birds per hunter (from check station) (1.4) are used, then the total
harvest estimate is 1,166 grouse (833 x 1.4).
It is reasonable to conclude

�-31-

that 1,200 - 1,400 sage grouse were harvested in North Park ia 1978. This
total is just slightly higher than in the previous 4 years (Table 6). This
is to be expected, as the population of sage grouse in North Park is
increasing and production in 1978 was good. Production in 1978 and 1974
were similar (53.2 and 50.1% young in the harvest, respectively, Table 10).
Harvest in 1974 was 1,174 birds, a figure comparable to that estimated for
1978 (1,200 - 1,400) with cruder techniques.
Band Recoveries
Sixty-three bands were reported from birds harvested during the 1978 hunting
season in North Park. Five additional bands were reported in 1978 from
predator kills (4) and trap casua1ties(1).
Of the 68 total recoveries, 21
were from females (1975 = 3, 1976 = 3, 1977 = 9, 1978 = 6), while 47 were
from males (1973 = 1, 1975 = 2, 1976 = 8, 1977 = 12, 1978 = 24). Of the
banded birds harvested, numbered bands were not present (either missing or
shot off) for 2 (1 male, 1 female).
One of the birds (female) was banded
in 1977. For recovery purposes, the female was assumed to have been banded
as an adult, while the male was assumed to have been banded in 1977 as a
yearling.
Recovery matrices are presented (Tables 17 and 18).

Table 17.

Year

Male sage grouse banding

Number Banded

1973

and recovery

1974

data, North Park 1973-1978.

Number Recovered
1976
1975

1977

1978

Yearlings
1973

80

1974

54

1975

139

1976

120

1977

183

1978

106

6

4

6

1

0

1

6

5

3

1

0

18

6

7

2

16

5

6

20

10
14

Adults
1973

99

1974

88

1975

152

1976

114

1977

123

1978

98

7

4

1

0

1

0

8

5

1

0

0

10

4

2

0

16

3

2

11

2
10

�-32-

Table 18.

Year

Female sage grouse banding

Number Banded

1973

and recovery

1974

data, North Park 1973-1978.

Number Recovered
1976
1975

1977

1978

Yearlings
1973

41

1974

22

1975

62

1976

71

1977

101

1978

32

2

2

4

1

2

0

2

1

1

0

0

6

2

1

2

2

4

2

6

5
5

Adults
1973

68

1974

27

1975

68

1976

74

1977

133

1978

24

5

1

0

1

1

0

2

0

0

1

0

6

2

4

1

2

2

1

14

3
1

It is apparent from data in the recovery matrices that few males banded as
yearlings or adults live longer than 4 years (only 2 recoveries past 4
years of age). Both males living past 4 years were harvested when 6 years
old. Females live longer than males, with 5 and 6 year old hens being
recorded.
Obviously each male cohort turns over in about 4 years. while it
takes about 7 years for each female cohort.
Direct recovery rates for males banded in 1978 varied from 10.1 (adults)
to 13.2% (yearlings).
For females, direct recovery rates were 4.5 (adults)
to 15.6% (yearlings).
While adequate samples of females were not banded,
the direct recovery rates for males and females combined were within the
range observed in 1973-1977.
Hunters in 1978 did not exploit the sage
grouse population at a higher rate than in earlier years.
Contingency chi-square tests for differences due to age were conducted
between adult and yearling males and adult and yearling females.
Significant (P&lt;0.05) differences were found in survival and recovery rate estimates
between yearling and adult males, but not between yearling and adult females.
Consequently, data for both age classes of females were pooled and survival
and recovery rate estimates were calculated through Program Estimate.
Data
for age classes of males were analyzed separately through Program Brownie
(Brownie et ale 1978).

�-33-

Data for females best fit Model 3 (analysis assuming constant survival
rates).
The constant annual survival rate was 63.38 + 6.14% (SE) (95%
confidence interval = 51.35 - 75.42) and the recoverY-rate was 6.77 +
.83% (SE) (95% confidence interval = 5.14 - 8.40) The estimate of 63%
annual survival is not different from the 65% estimated from wing data
(Table 12).
Data for males fit either Model HI (annual survival and recovery rates are
year specific, and yearlings have different survival and recovery rates
from those of adults) or Model H2 (same assumptions as HI, plus reporting
rates differ for newly banded birds).
A likelihood ratio test of Model HI
versus Model H2 suggested that HI is more appropriate for these data.
The constant annual survival rate for adult males was 43.48 ± 6.34% (SE)
(95% confidence interval = 31.05 - 55.91) with an annual recovery rate of
8.48 + 1.00% (SE) (95% confidence interval = 6.52 - 10.43). The constant
annual survival estimate for yearling males was 85.26 + 15.68% (SE) (95%
confidence interval = 54.53 - 115.98) with an annual recovery rate of 11.51 +
1.30% (SE) (95% confidence interval = 8.96 - 14.05). Confidence intervals
for adult males and all females were relatively narrow (24% range), while
that for yearling males was broad (61% range).
It is believed these differences are related to differences in vulnerability between years of
yearling cocks to hunting.
Reasons for the apparent differences are not
known.
It is doubtful that yearling males survive at an annual rate of 85%.
More likely, yearlings are surviving at a rate of 60 to 70% per year.
If
survival data for yearling and adult males are pooled (they cannot be
statistically), an estimated survival rate for all males would be about 50%
(relative recovery rate method).
This rate is -similar to that estimated
from wing data alone (Table 11).
Estimates

of Fall Population

Size

Estimates of fall population size are difficult to derive, primarily because
of unknown vulnerability and survival of chicks and inexact survival estimates.
Provided that all birds of each age and sex class are equally
available and vulnerable to hunting and using average survival estimates
(63% for all females, 43% for adult males and 70% for yearling males), the
1978 fall population size was estimated based on number of banded birds
alive at the start of the hunting season.
Of the 482 banded males potentially alive in 1978, bands were received from
47, while bands were received from 21 of the 318 banded females potentially
alive.
If 47% (wing survey data) of the estimated 1,300 birds harvested
(this report based on check station data) were adults and yearlings, then
611 of the birds harvested were older than one year. Of this number, 63
(68 - 5 miscellaneous recoveries) (10.3%) were banded. Thus, there should
have been about 5,932 adult and yearling sage grouse alive in North Park in.
the fall of 1978 (if 10.3% = 611 birds, then 100% = 5,932 birds).
In 1978,
immatures comprised 53% (wing survey data) of the harvest, thus there should
have been about 6,689 chicks in the 1978 fall population.
Thus, the total
population in the fall of 1978 should have been about 12,621 birds.
This
estimate is probably conservative.

�-34-

North Park,
Table 19. Estimated number of banded sage grouse alive,
Colorado, fall 1978.

Year

No. of Birds Banded
Males
All Females
Adults
Yearlings

Alive in 1978
Females
Males

1973

80

99

109

3

17

1974

54

88

49

6

8

1975

139

152

130

30

33

1976

120

114

145

57

57

1977

183

123

234

181

147

1978

106

99

56

205

56

482

318

Total Potential Alive, Fall, 1978

Using band recoveries alone, 63 of the estimated 1,300 birds harvested (7.9%)
were banded. If 800 banded birds were alive (Table 19) in the fall population
then it was comprised of a total of 16,460 birds (if 7.9% = 1,300, then
1% = 164.6 birds and 100% = 16,460).
LITERATURE CITED
Beck, T.D.I. 1975. Attributes of a wintering population of sage grouse,
North Park, Colorado. M. S. Thesis. Colorado State Univ., Ft. Collins.
49 pp.
1977. Sage grouse flock characteristics and habitat selection in
winter. J. Wildl. Manage. 41:18-26.
_____ , R. B. Gill, and C. E. Braun. 1975. Sex and age determination of
sage grouse from wing characteristics. Game Inf. Leaflet No. 49
(Revised). Colo. Div. Wild1. 4 pp.
Braun, C. E., and T.D.I. Beck. 1976. Effects of sagebrush control on
distribution and abundance of sage grouse. Colorado Div. Wildlife,
Final Rept., Fed. Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84.
Brownie, C., D. R. Anderson, K. P. Burnham, and D. S. Robson. 1978.
Statistical inference from band recovery data - a handbook. U.S.D.I.
Fish and Wi1dl. Servo Resour. Publ. No. 131. 212 pp.

�-35-

Buss, I. 0., R. K. Meyer, and C. Kabat. 1951. Wisconsin pheasant reproduction studies based on ovulated follicle technique. J. Wildl. Manage.
15:32-46.
Carr, H. D. 1967. Sage grouse and sagebrush control.
rado State Univ., Fort Collins. 106 pp.

M. S. Thesis, Colo-

Emmons, S. R. 1979. Evaluation of censuses of male sage grouse. Colorado
Div. Wildlife, Prog. Rept., Fed. Aid Proj. W -37-R, Work Plan 3,
Job 9b.
Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios
from wings. J. Wildl. Manage. 19:267-272.
Gill, R. B. 1965. Distribution and abundance of a population of sage grouse
in North Park, Colorado. M. S. Thesis. Colorado State Univ., Fort
Collins. 187 pp.
Hoffman, R. W., and C. E. Braun. 1975. A volunteer wing collection station.
Game Inf. Leaflet No. 101. Colo. Div. Wildlife. 3 pp.
Kabat, C., I. O. Buss, and R. K. Meyer. 1948. The use of ovulated follicles
in determining eggs laid by the ring-necked pheasant. J. Wildl.
Manage. 12:399-416.
Meyer, R. K., C. Kabat, and I. O. Buss. 1947. Early involutionary changes
in the post-ovulatory follicles in the ring-necked pheasant. J.
Wildl. Manage. 11:43-49.
Pyrah, D. 1963. Sage grouse investigations. Idaho Fish and Game Dept.,
Wildlife Restoration Div., Job Comp L, Rept., Fed. Aid Proj. W-125-R.
71 pp.
Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male
sharp-tailed grouse. J. Wildl. Manage. 38:616-621.
Robel, R. J. 1969. Movements and flock stratification within a population
of blackcocks in Scotland. J. Anim. Ecol. 38:755-763.

Prepared by __

-=a_~_'_--=2~._~r-.=-==~__

Clait E. Braun
Wildlife Researcher

��-37-

April 1979

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~----------Game Bird Survey

Project No.

W-37-R-32

Work Plan No.

Job No.
~~~~9~b~~--~~-----3
Evaluation of the Effects of Changes in Hunting Regulations
on Sage Grouse Populations:Evaluation
of Censuses of Males

Job Title
Period Covered:
Personnel:

1 January through 30 September

1978.

Clait Braun, Jim Dingman, Steve Emmons, Ken Giesen, Brett
Petersen, Colorado Division of Wildlife.

ABSTRI\.CT
Investigations concerning daily lek attendance and spring movements of
individual male sage grouse (Centrocercus urophasianus) were initiated
in 1978 in North Park, Colorado.
Seventeen cocks were equipped with
radio transmitters and studied on 4 leks from late March through
mid-June.
Peak cock attendance occurred from 29 April to 10 May on the 3
largest leks, 3.5 to 5 weeks after peak hen attendance.
Lek attendance of
transmittered juveniles increased to 1-15 May (86.7%) and decreased thereafter
(45.2%). Adult cock attendance increased to 93.2% (1-15 May) and decreased
to 64.1% (16-30 May). Twenty-nine percent of all off-lek locations were
within 0.5 km of a lek and 32.3% were attributed to late observer arrival
and disturbance.
Juvenile cocks visited from 2 to 4 leks, remaining on each
for an average of 3.4 days. Adults visited 1 to 2 leks, remaining on the
alternate lek for only 1 day. Juveniles and adults moved 32.7 and 19.4 km,
respectively, during the breeding season. Four juvenile cocks moved from
the study area including a move of over 25 km in 7 days. An adult cock
captured off a lek moved 12 km to a lek not in the study area. Males typically
moved over 1 km from leks to feeding and loafing sites. Subsequent mid-day
moves over 150-200 m resulted from disturbance.
Lek roosting for juvenile
and adult cocks was 44.4 and 65.5%, respectively.
Sagebrush (Artemisia spp.)
canopy coverage and height at feeding-loafing sites averaged 29.3% and 45.5
cm, respectively.
Eighty percent of the 'roosting locations occurred in
sagebrush with a canopy coverage of less than 20% and height less than 22.5
cm. Two adults were killed by predators during the spring and 3 juveniles
were harvested during the 1978 sage grouse hunting season.

�-38RECOMMENDATIONS

1.

A minimum 6f 20 cocks (10 juveniles,
during the 1979 season.

10 adults) should be monitored

2.

Trapping effort should be concentrated in the vicinity of Alkali Lake
and Boettcher Lake Junction leks, and along roads between them. Trapping
and intensive counts should be discontinued on the Bighorn (access
difficulties) and Wattenberg #2 leks. Movements to these Z leks'by
transmittered cocks should be monitored.

3.

Trapping and banding of cocks should be initiated by 15 March or earlier
depending on weather conditions and lek attendance.
Snow removal should
be initiated in early March to facilitate movement in the study area.

4.

A minimum of 3 juvenile and 3 adult cocks should be monitored
given time beginning on 20 March continuing through 30 June.

5.

Two different selected individuals
monitored at 1 to 2 hour intervals
the field season.

6.

All transmLttered cocks should be monitored for roosting
2 or 3 times per week throughout the field season.

7.

All radio transmitters should be adequately tested for signal constancy
under temperature extremes at least 3 days prior to expected day of use.

8.

Temporary markers (surveyor's lath) should be utilized to enable accurate
identification of feeding-loafing and roosting sites for later vegetation
analysis.

9.

Vegetation
throughout

10.

Four or 5 line transects
the current 3 lines.

at any

(1 of each age class) should be
throughout 2 or 3 days per week during

sites (2100 MST)

analysis should be performed concurrently with other operations
the field season.
It should not be concentrated late in June.
should be used to analyze vegetation

instead of

�-39-

EVALUATION OF THE EFFECTS OF CHANGES IN HUNTING REGULATIONS
ON SAGE GROUSE POPULATIONS:EVALUATION
OF CENSUSES OF MALES
Steven R. Emmons

Sage grouse are widely distributed throughout sagebrush dominated rangelands
in western North America.
Throughout most of their range, state wildlife
agencies routinely collect population data and regulate hunter opportunity
and harvest.
Sage grouse management has primarily been based on counts of
cocks on leks in April and estimates of nesting success and brood size
obtained in July and August.
Observations concerning the daily movements and activities of male sage grouse
during the breeding season have been reported (Dalke et al. 1960, 1963, Gill
1965, Carr 1967, May 1970, Wallestad and Schladweiler 1974, Braun and Beck
1976). Although peak counts of male and female sage grouse on leks have
been used to estimate trends in breeding season population size, little is
known concerning the daily attendance patterns of individual males to a
particular lek. Recent studies of other lekking grouse, black grouse (Lyrurus
tetrix) (Robel 1969) and sharp-tailed grouse (Pedioecetes phasianellus)
(Rippin and Boag 1974), indicate that less than 50% of the male population
is present on a lek at any given time. Data presented in this report concern
breeding season activities and habitat selection of male sage grouse and
represent mid-March through July 1978.
P. N. OBJECTIVE
The objective of this study is to increase knowledge concerning daily activity
patterns of male sage grouse during the breeding season {late winter, spring
and earlv summer).
Samples stratified bv age (adult and yearling) and
experience (trapped on and off leks) will be used to document daily lek
attendance of males.
Hypotheses which are being tested are:
a.

Adult males attend leks daily during the breeding

season.

b.

Juvenile males attend leks less than 3 days per 7 day period.

c.

Adult males display on only one lek.

d.

Juvenile

e.

Males (both adults and juveniles) trapped off leks rarely go to leks
during the breeding season of initial capture.

males display on more than one lek.

SEGMENT OBJECTIVES
1.

Males will be trapped at night while roosting, marked with color-coded
bands and equipped with radio transmitters.
A sample of 20 is desired
including adults and juveniles trapped on and off leks. Trapping will be
conducted from mid-March through April by spotlighting and netting or
cannon-netting.

�-40-

2.

All transmittered males will be located each morning
MST from late March through May.

3.

Daily activity patterns from 0430 to 2100 MST will be monitored for
selected males of each age class (adult and juvenile) at intervals
throughout the life of individual transmitters.

4.

Sites utilized for feeding,
classified as to vegetative
graphic features.

5.

Counts will be made of males and females on 4 selected
area throughout the breeding season.

6.

Compile

data, analyze

loafing and roosting will be analyzed and
composition, cover and height, and physio-

results,
METHODS

from 0430 to 0730

and prepare progress

leks in the study

repo~t.

AND MATERIALS

Sage grouse were captured at night along roads and on leks while roosting.
Most birds were located using a spotlight with a backpack
or hand-held power
source and captured with long-handled nets (Pyrah 1959, Braun and Beck 1976).
A vehicle-mounted
cannon net (Lacher and Lacher 1974) was used on one occasion.
Sage grouse banded and released were marked with serially numbered aluminum
leg bands (size 16) and individually coded colored plastic bandettes.
Selected
males were equipped with 14 gm, very high frequency (VHF) radio transmitters
using tail clips (Bray and Corner 1972). Weights and primary molt were
determined for all captured birds.
Age and sex classification of birds
captured followed Eng (1955).
All transmittered males were located daily each morning using portable
receivers with hand-held 3-element yagi antennas.
Selected males were
located periodically (every 1 to 2 hours) throughout selected days to ascertain
daily activity patterns.
All locations were recorded on standardized forms
(Appendix A).
Vegetative cover measurements for radio locations were made along 150 feet
(45.72 m) of line (three 50-foot [15.24 m] lines originating from a central
point) using Canfield's (1941) line intercept technique.
Vegetative height
of the dominant species, big sagebrush (Artemisia tridentata vaseyana),
greasewood(Sarcobatus
vermiculatis) and rabbitbrush (Chrysothamnus spp.), was
measured to the nearest 0.25 cm.
Counts of male and female sage grouse present on leks were made daily by the
principal investigator, supplemented with periodic counts by project personnel,
following prescribed procedures (Braun and Beck 1976). Counts were made from
23 March through 30 May between 0430 and 0730 MST. Radio locations were taken
concurrently with lek counts.
A transmittered cock was determined to be on a
lek if signals indicated the bird's presence on or near the lek. Signals
indicating the bird was more than 0.1 km from the periphery of a lek were
recorded as an off lek location.

�-41-

DESCRIPTION

OF AREA

The study area is near Lake John in the northwest quarter of North Park,
Jackson County, Colorado (Fig. 1). It is bounded on toe north by Independence
Mountain, the west by Boettcher Ridge and Sheep Mountain, the south by the
North Fork of the North Platte River, and the east by the North Platte River
(Fig. 2). Elevations are from 2400 m along the North Platte River to 2964 m
on Independence Mountain.
Vegetation is dominated by a sagebrush-grassland
type with scattered hay meadows and greasewood flats. Detailed geographic,
geologic, vegetational and climatic features have been described by Gill (1965),
Carr (1967), Braun and Beck (1976) and Beck (1977)
and will be treated in
the final report.
RESULTS
Male sage grouse were captured
juvenile cock was instrumented
(Tab;Le 1). Late capture dates
late winter storms and delayed

AND DISCUSSION

and marked from 18 March through 1 May.
One
on 25 February as an "experimental" bird
were related to poor access resulting from
melting.

Capture efforts were concentrated along Jackson County Roads 7 and 7A (J.C. 7
and 7A) and on Alkali Lake, Bighorn, and Boettcher Lake Junction leks. Access
difficulties
on the Bighorn Ranch (location of Bighorn lek) resulted in
shifting capture efforts to include Wattenberg
#2 lek and along J. C. 9A
(Fig. 2).
Seventeen sage grouse cocks were equipped with 16 radio transmitters.
Age
class and capture site stratification was: 8 adults and 5 juveniles caught on
leks, 1 adult and 3 juveniles caught off leks (Table 1). Difficulty in
locating roosting adults off leks during the breeding season accounted for
the low representation of that age and capture site class in the sample.
Transmitters remained functional an average of 58.4 days, ranging from 4
to over 129 days (Table 1). Five transmitters were operational beyond
termination of the intensive research period (9 June) and were periodically
located from mid-June through mid-August.
Breeding

Activities

Lek Attendance
Counts of male and female sage grouse present on leks in the Lake John study
area were initiated on 28 March and continued through 30 May. During this
interval 204 counts were made on 4 active leks (Table 2).
Peak hen attendance occurred from 2 to 16 April and declined rapidly thereafter
(Fig. 3). A second lower peak occurred between 7 and 15 May, a situation
previously reported (Dalke et al. 1963, Eng 1963, Gill 1965).
The rapid
increase in cock attendance was similar to the late spring situation described
by Jenni and Hartzler (1978). The prominent decrease from 21 to 25 April
resulted from disturbance (Alkali Lake lek 22 and 23 April, Bighorn lek 21-29
April, Boettcher Lake Junction lek 22 and 23 April) and late observer arrival
(Wattenberg #2 lek on 21, 23 and 25 April).

�-42-

:.\.~.,.
.. !
. 1
r -

J

I~ •.

Fig -, 1.

North Park, Colorado and Lake John study area.

�-43-

SCALE:

Fig. 2.

0.6 in. = I ml.

Location of active leks in Lake John study area, North Park, Colorado.

�Table 1.

Summary,

telemetry

data,

17 male

sage grouse,

North Park,

Bird
Number

Age

Capture
Location

Date of
Capture

Date of
Last Location

7035

Juv

J .C. 7

25 Feb

31 Mar

7036

Juv

J .C.

22 Mar

6 Apr

Ad

7037

Ad

6759

Ad

7039

J .C.

7

28 Mar

7

29 Mar

Boettcher

29 Mar

Boettcher

Colorado

Duration
(Days)

Tot~l
Locatlons

1/
-

Transmitter
Life
(Days)

30

76

"Experimental"

15

23

27

Moved

from study area

Moved

from study area, killed

30 Mar

2

3

81

3 May

3511

20

67

Intermittant

25 Apr

27!!./

33

Killed

Ad

Alkali

31 Mar

22 May

7040

Ad

Alkali

31 Mar

27 Apr

27

7042

Ad

Bighorn

31 Mar

9 June

70

7043

Juv

Alkali

3 Apr

13 Apr

7044

Juv

Alkali

3 Apr

2543

Ad

Wattenberg

4 Apr

18
54'i..!

67]../

99

Poor off-lek locations
difficulties

11~1

12

19

Moved

from study area

21 Apr

19~/

10

24

Moved

from study area

8 Apr

4

4

4

Transmitter

53

Shot 9 September

1978

20

Shot 9 September

1978

Shot 9 September

1978

Juv

Boettcher

5 Apr

27 May

52

7101

Juv

J.C.9A

11 Apr

24 Apr

131Q/

11

6907

Ad

Boettcher

14 Apr

9 June

56

54

66
89

7147

Juv

Boettcher

22 Apr

9 June

47

501/

7183

Juv

Alkali

29 Apr

9 June

41

37

129~/

7190

Ad

Boettcher

May

9 June

39

42

96

11 Not

including

1 day status

81
- Includes

2 days no signal

obtained.

21 Includes

11 days no signal

obtained.

undetermined.

101 Includes

3 days no signal

obtained.

15 days no locations

11 Includes

27 days no signal

obtained.

~/ Includes

12 days no signal

21 Includes

2 days status

lost by 7040 and reused

21 Includes

]j Includes

taken.

transmitter

~I

~I Transmitter

by eagle

(by coyote?)

32

7051

site.

bird

52

6221

capture

Remarks

34!:./

52

6943

1978.

undetermined.
obtained.

on 7183.

Bird obs. stlutting
14 May 1978

on Alkali

failure

on

due to access
I

"'"
"'I"

�-45-

.-.-.,

160

I
150
r-..
Q)

OJ)
Cd
H

I
140

/

~ 130

\

.-I

~ 120

/

Q)

H 100

P

0
(Jl

90

13
Q)

.fJ\,'

f:I.&lt;

-e
p
Cd

I

.

70

.-I
Cd

0

50

I

/

.w

.

I:'

\

I:

I

\1

\

\
\
\
\

'0

\

60
50

\

\

I
N
N

N

r--.

March

tr)

I
.-I

0
.-I
I
\0

tr)

.-I
I
.-I
.-I

0

l()

N

I

I

0
M
I

N

\0
N

\0

.-I

April

Sage grouse attendance

..--j

tr)

•

40

I

I
.-I

\0

tr)

.-I
I
.-I
.-I

'M
13
Cd
H
IO-!

.w

20

p
Q)

()

H
Q)

0

tr)

N

I

I

.-I

H

p

30

N
\0

Q)

(Jl

". 0···· ()... ,

0
.-I
I

;:.::

Q)

\

10

N

.-I
Cd

.w
.w

0.•••0,

.-I
M
I

0

"d

&amp;

t(

p

Q)

\I

• Females
\0.
/

Q)

CIl

T r ansnu.t t ere d \ I
.Iuven L'1e--.. \ I
Males
\I

'0 .. '. 0 •..•
0'···

Fig. 3.

•\

70

\

\

~

H

\

\ 0

Llo'

\0
N

,/®

\

\

•

(Jl

80

\

~

Transmittered
~dult
0
Males

,I I.:Id

10

0

\

I .:....

20

\

\

(.~""~.'
\
,..
I.'

30

90

\0

I

•

40

\

l--.//~~

'-.....0\

,

Z

0
IO-!

/

._a_e

/

\1

I

~

.-I
Cd

0

I

H
Q)

I·

'\{ 1°'"

100

I

60

;:.::

4-l

••
\
\

-:-0

.'

(Jl
Q)

.-I

i;

80

\

.

/

/
•

Q)

.-I
Cd

\

. I
\

(Jl

~

\

I

\

1

110

I

,

\

/
•

Q)
Q)

~

\

\

&lt;r:
;:.,

'-'

I

/

•

..--j
N

0
M
I
\0
N

May

on 4 leks in North Park, Colorado,

1978.

p.,

�-46-

Table 2.

Peak counts of sage grouse on 4 leks, North Park, Colorado

1978.

Females

Date(s)

10 May

20

4 April

44

29 April

19

4 April

Boettcher Lake
Junction

84

1 &amp; 3 May

84

4 April

112

21

26 May

7

7 April

Lek

Males

Date(s)

Alkali Lake

56

Bighorn

Wattenberg

Data from 10 birds (5 adults and 5 juveniles) were used for evaluating daily
attendance patterns (Table 3). Poor data were available from 7 cocks.
Data
from these birds were eliminated from analysis for the following reasons:
intermittent locations and movement from the study area (7035), injury and
subsequent depredation (7039), intermittent transmitter (6759), transmitter
failure after 4 days (2543), and movement from the study area after 1 to 4
days (7037, 7043, 7044) (Table 1).
Late capture dates resulted in only 1 juvenile with attendance data for March:
7036 was present on a lek for 1 (11.1%) of 9 early morning locations.
No comparable data are available for adult cocks during March (Table 3).
Lek attendance for transmittered juvenile cocks was 70.8% of 48 locations in
April.
Juvenile attendance increased during April from 52.6(1-15
April) to
82.8% (16-30 April) (Table 3). .This significant increase (~&lt;O .10) supports
other observations of increased numbers of juvenile cocks attending leks
following peak hen attendance in early April (Patterson 1952, Dalke et al.
1963, Eng 1963, Gill 1965, Jenni and Hartzler 1978). Human disturbance and
late arrival by research personnel accounted for 35.7% of 14 days juveniles
were not present on leks (Table 4). Individuals were within 0.5 km of a lek
on 28.6% of the off-lek locations.
Lek attendance of instrumented adults in April was 67.0% of 100 morning locations
(Table 3). The decrease from 71.7 to 63.0% (1-15 to 16-30 April) was not
significant (P&gt;0.05) indicating a generally stable level of adult cock attendance through~ut the month.
Disturbance by eagles and late observer arrival
accounted for 15.2 and 6.1%, respectively, of April off-lek locations (Table 4).
On 6 occasions individuals were within 0.5 km of a lek. One adult provided
data possibly indicating the existence of a non-breeding segment of the male
population, a situation described for other lekking grouse (Robel 1969, Rippin
and Boag 1974).
No. 6943 was absent from a lek between 6 and 18 April but
displayed on Alkali Lake lek the remainder of the period from 1 April to 22
May.
This period of non-attendance
followed peak hen attendance (3 April)
and was a period when average hen attendance remained relatively high
(averaging 4.4 hens per day).
Only one other male (age undetermined) accompanied 6943 for one day during this l3-day period in contrast to the
flocks of non-breeding black grouse reported by Robel (1969).

�Table 3.

Stratification of lek attendance by male sage grouse in North Park, Colorado 1978.

Bird
Number

Harch
16-31
On Lek Off Lek

Aril
1-15
On Lek Off Lek

16-30
On Lek Off Lek

May
1-15
On Lek Off Lek

16-3Q
On Lek Off Lek

Juveniles
7036

1

8

2

4

7051

-

-

6

4

13

1

2

1

5

2

-

-

6

1
11.1

8
88.9

10
52.6

9
47.4

1
11.1

8
88.9

-

-

3

12

8

6

15

0

3

1

14

1

10

4

13

2

6

5

15

0

6

6

0

1

10

4

14

1

8

4

-

-

-

-

-

33
71.7

13
28.3

34
63.0

20
37.0

13
55
93.2

1
4
6.8

8
25
64.1

4
14
35.9

7101
7147
7183
Subtotals
%

Monthly
Subtotals
%

14

1

3

5

1

14

1

5

6

0

1

11

4

6

6

24
82.8

5
17.2

39
86.7

6
13.3

14
45.2

17
54.8

Off = 23
30.3

On = 53
69.7

Off = 14
29.2

On = 34
70.8

Adults
6943
7042
7040
6907
7190
Subtotals
%

Monthly
Subtotals
%

-

-

On = 67
67.0

Off = 33
33.0

On = 80
81.6

Off = 18
18.4

I

+:-...J
I

�Table 4.
1978.

Month
March
06-31)

April
0-15)

April
06-30)

Monthly

stratification

of attributes

of off-1ek

Age
Class

Total
Off-Lek
Locations

Distance Less
Than 0.5 km

Juvenile

8

1 02.5)

Adult

0

Juvenile

locations

for male sage grouse, North Park, Colorado

Attributes of Off Lek Locations
Late Arrival
By Observer
Eagle

(%)
Other

0

0

0

0

0

0

0

9

2 (22.2)1:/

0

0

2 (22.2)

Adult

13

0

1 (7.7)

0

0

Juvenile

5

2 (40.0)1:/

1 (20.0)

0

2 (40.0)

Adult

20

6 (30.0)1/

1 (5.0)

5 (25.0)

0

I

+:C/)

May
0-15)

May
06-31)

I

Juvenile

6

3 (50.0)1/

3 (50.0)

0

1 06.7)

Adult

4

1 (25.0)1/

3 (75.0)

1 (25.0)

0

Juvenile

17

6 (35.3)1/

4 (23.5)

0

0

Adult

14

7 (50.0)]/

7 (50.0)

0

0

11 Includes 1 day accounted for by late arrival or disturbance
11 Includes 3 days accounted for by late arrival or disturbance
11 Includes 4 days accounted for by late arrival or disturbance
------

�-49-

Juvenile cock attendance increased (P&gt;O.OS) for 1-15 May (86.7% attendance)
(Table 3). The decrease in lek atte~dance to 45.2% for 16-31 May is significant (P&lt;O.OS) and corresponds with the trend observed by Jenni and Hartzler
(1978) a;d others (Eng 1963, Gill 1965). However, during the period of 15-31
May the primary researcher was not present on the study area for 4 days and
a late snowstorm created access difficulties to one lek for 2 additional days.
These factors may have influenced the size of this decrease.
Average attendance for the entire month was 69.7% of 76 locations.
Approximately
35% (34.8)
of the May off-lek locations were attributed to disturbance and late arrival
by the observer.
Individuals were within 0.5 km of a lek on 39.1% of the offlek locations (Table 4).
Lek attendance by transmittered adult males increased (P&lt;O.OS) to 93.2% for
1-15 May and decreased (~&lt;O.OS) to 64.1% during 16-31 May (Table 3). All 4 offlek locations recorded for 1-15 May were attributed to eagle disturbance and
late observer arrival.
This period was when peak counts of males occurred on
the 2 largest leks (Alkali Lake and Boettcher Lake Junction) (Table 2) with
high counts occurring in the entire study area (Fig. 3). This increase was in
contrast to conditions reported elsewhere where marked declines occurred 4 or
5 weeks following peak hen attendance (Dalke et ale 1960, Eng 1963).
Braun
and Beck (1976) noted that numbers of adult males "steadily decline following
peak hen attendance .••..
" This was not observed in this study.
Transmittered adults attended leks 81.6% of the locations for May. Adults were
within 0.5 km of a lek on 44.4% of the off-lek locations (Table 4).
Attendance of juvenile cocks increased through 10 May although a marked
decrease for 6-10 April occurred (Fig. 3). This decrease may have been the
result of extremely small sample size (only 1 transmittered juvenile) from
23 March to 12 April.
Decreased juvenile attendance after 15 May reflected
total cock attendance.
This was alsc influenced by observer absences and a
snowstorm.
Lek attendance probably was closer to 50 or 60% for transmittered
juveniles during this period.
Fl uctuations in lek attendance of adults (Fig. 3) can be attributed to 6943
not attending a lek from 6 to 18 April, frequent disturbance of leks (21-25
April), and observer absence and access difficulties (16-20 May) (Fig. 3).
Frequent disturbances from 21 to 25 April did not decrease lek attendance of
transmittered juveniles because none was present on Bighorn (21-23 April)
and Alkali Lake leks (22 April) on days of disturbance.
Five adult locations
can be attributed to these disturbances.
Periods of high lek attendance by
transmittered adults and juveniles corresponded with similar periods for
total males counted.
Movements
Interlek
movements by juvenile cocks were found to be more common than
previously reported (Dalke et ale 1963, Gill 1965, Braun and Beck 1976).
Juveniles averaged 4.2 moves (range 1 to 13) during the breeding season (Table
5). These results do not include moves to leks outside the study area (7036)
and possible moves unrecorded on 5 days when research personnel were absent
from the study area.

�-50-

Table 5. Movements
Colorado 1978.

No. Interlek
Bird
No.

April

7036

1!1

7051

0

7101

2

7147

4

May

71/

1/ Includes

Moves
Breeding
Season

sage grouse between

4 leks, North Park,

No. Different Leks
Breeding
Season
May
April

11/

21/

1

1

2

2

8

131/

2

4

4

13

21.!I,1/

1

7183

Totals

by juvenile

41/

Average
Time/Visit (Days)
Breeding
Season
April
May

21/

1.51/

2

19.0

2

2.3

4

4

1.5

3

3

4

4

2

3.51/

1.51/
8.5

18.0
2.3

2.1

1.9

3.4

3.4

3.3

3.41/

a move in March.

J) Includes a lek visited

in March.

1/ Includes

April and May.

a move between

The increase in interlek moves from April (1.75/bird) to May (4.5/bird) was
not significant (R:0.05) indicating similar lek fidelity throughout the
breeding season.
Number of leks visited increased (P&gt;0.05) from April (1 to 2)
to May (2 to 4). Each juvenile visited from 2 to 4 leks (average 2.6) during
the breeding season.
Distances between leks averaged 5.2 km, ranging from 2.4 to 8.1 km (Table 6).
Average distance per juvenile interlek move was 4.7 km. Moves commonly
occurred between Alkali Lake and Boettcher Lake Junction leks (47.6%), and
Alkali Lake and Wattenberg #2 leks (23.8%).
The number of interlek moves
between Alkali Lake and Boettcher Lake Junction leks was unexpected because
shorter distance and topography should have increased movements between Alkali
Lake-Wattenberg
#2 rather than Alkali Lake-Boettcher Lake Junction.
The reason
for this discrepancy is unknown but may be related to lek size (Table 2).
Juveniles averaged at least 17.4 km (range 2.4 to 54.2 km) in interlek moves
during the spring.
Juveniles remained on a lek for an average of 3.4 days, ranging from 1 to 34
non-consecutive
days where visits to that lek were intermixed with off-lek days.
The average number of consecutive days per visit was 2.4, with a maximum
of 15 days (7051).

�-51-

Table 6. Interlek distances (km) and sage grouse movements,
area, North Park, Colorado 1978.

Alkali Lake
No.
Movements
Distance
2+
Between
1-

Bighorn
Distance
Between

No.
Movements
2+
1-

Bighorn

5.1

2

0

Boettcher
Lake Junction

4.5

10

0

2.4

2

0

Wattenberg

3.2

5

0

8.1

1

0

fI2

Lake John study

Boettcher Lake
Junction
No.
Movements
Distance
2+
Between
1-

7.6

1

2

Interlek movements for individual juveniles varied.
Bird 7051 was sedentary,
remaining on or near the lek of initial capture from 5 April through midMay, finally moving to another lek on 25 May. No. 7147 moved 13 times
between 4 leks and remained on each for 3 consecutive days or less (may have
been on 1 lek for up to 8 days, with 4 days unrecorded due to late arrival
and observer absence).
Other individuals represented intermediates between
these 2 extremes.
Interlek moves by adults were infrequent.
Bird 7190 was the only adult
observed to attend 2 leks. It appeared on Wattenberg #2 lek on 9 May, while
displaying on Boettcher Lake Junction lek the remainder of the period from
2 to 30 May. The bird was also in the vicinity of Wattenberg #2 from 6 to
9 June, indicating reduced fidelity to the normal lek late in the season.
Although other individuals left leks for considerable lengths of time (e.g.
6943 between 6 and 18 April) all displayed on only 1 lek.
Juvenile
moves averaged 32.7 km throughout the breeding season (Table 7).
Mean distance per move was 2.0 km. This was shorter (K&lt;0.05)
than the
average interlek move observed for this group (4.7 km) indicating many short
moves of 1.0 km or less that were not between leks.
Adult cocks moved an average of 19.4 km (Table 7). Average distance per
move (1.1 km) was considerably shorter (P&lt;0.05) than that of juveniles.
This
was attributed to the large number of relatively long (&gt;1.5 km) interlek
movements of juveniles.
Thirty-six percent of all off lek locations were beyond 1.5 km (Table 8). OfT-lek
distances beyond 2.0 km comprised 10.3 and 14.5% of all off-lek locations for
juveniles and adults, respectively. These distances were greater than maximum
distances reported by others (Carr 1967, Wallestad and Schladweiler 1974).
Long distance movements from the study area were recorded for 4 juvenile and
1 adult transmittered sage grouse (Table 1). Between 7 and 13 April, 7036
moved a minimum of 25.7 km to near a lek (Spring Creek 114) southeast of
Walden.
Subsequent contact was lost after 14 April.

�-52-

Table 7. Distances (km) traveled by male sage grouse during the breeding
North Park, Colorado 1978.

Bird
Number

Consecutive 1/
Locations (A)-

season,

Number of
Moves '.!:./
(B)

Total
Distance

Mean
Distance
Between A

Mean
Distance
Between B

Juveniles

7036

20

14

26.3

1.32

1.88

7051

23/20/7/101/

19

29.6

.49

1.56

7101

4/7

7

12.6 .

1.15

1.80

7147

6/19/13

24

66.0

1.74

2.75

7183

19/7

18

28.9

1.11

1.61

Subtotals

155

82

163.4

1.05

1.99

6943

5/17/19

16

21.2

.52

1.33

7040

31

12

9.2

.30

.77

7042

31/18/6

22

15.2

.28

.69

6907

14/20/9/7

19

18.2

.36

.96

7190

18/4/13

16

33.1

.94

2.07

Subtotals

212

85

96.9

.46

1.14

Total

367

167

260.3

.71

1.56

Adults

1/ Includes

2/

Includes

1/ Groups

locations

on a lek.

interlek moves.

of consecutive
were taken.

locations

are separated

by a slash (/) when no locations

�-53-

Table 8. Frequency distribution of distances for off-lek
male sage grouse in North Park, Colorado 1978.

J uvenl'1es-1/

2/
AdultsNumber
Percent

locations

of 10

Total
Percent

Distance
Class (km)

Number

Percent

0.01 - 0.25

2

2.6

1

1.4

3

2.0

0.26 - 0.50

17

21.8

18

26.1

35

23.8

0.51 - 0.75

5

6.4

5

7.2

10

6.8

0.76 - 1.00

9

11.5

14

20.3

23

15.6

1. 01 - 1.25

3

3.8

8

11.6

11

7.5

1.26 - 1. 50

6

7.7

6

8.7

12

8.2

1.51 - 1. 75

15

19.2

3

4.3

18

12.2

1. 76 - 2.00

13

16.7

4

5.8

17

11. 6

2.01+

8

10.3

10

14.5

18

12.2

Totals

78

100.0

69

99.9

147

99.9

1/ Distances

measured

from nearest

])

measured

from cocks normal

Distances

Number

lek.
lek.

Two juveniles moved from the study area on 7 April, 5 days after the night
of instrumentation.
Bird 7043 was located 2.5 km north of Walden Reservoir
(northwest of Walden) on 12 April and thereafter contact was lost. No. 7044
was located approximately 6 km south of Wattenberg #2 lek on 18 April.
Subsequently, the bird moved to Alkali Lake lek and near Bighorn lek before signals
were lost on 22 April.
Long distance moves were periodically recorded for no. 7035 ("experimental"
bird) from 25 February to 31 March (Fig. 4). Total recorded distance was a
minimum of 80 km.
The only adult cock captured off a lek was 7037 (28 March).
It was
located 6.7 km southwest of the study area on 30 March.
From 31 March to
17 April, signals indicated the bird was near Aspen lek west of Delaney
Butte (12 km southwest of the study area), although poor access (snow)
prohibited accurate locations.
Accurate locations were periodically obtained
from 24 April to 9 May: three within 0.2 km and one within 1.9 km of Aspen lek.
This individual probably had not returned to its normal lek (Aspen) prior to

�-54-

N

Capture
Site
E

W

s
Each concentric circle represents 5.0 km with the center located at
Walden, Colorado.
Number of locations at each site is indicated (no
number means only 1 location).

Fig. 4. Movements by juvenile
25 February to 31 March 1978.

sage grouse 7035 in North Park, Colorado,

�-55-

capture in the Lake John study area. All data indicate 7037 returned
to his normal lek and remained in the vicinity throughout the spring.
Although not monitored during the early morning strutting period, the bird
probably regularly displayed on the Aspen lek. This individual did not
represent a member of a non-breeding segment of the male population despite
being captured while roosting off a lek.
Daily Activity

Patterns

Selected transmittered individuals of both age classes were monitored at 1
to 2 hour intervals on 21 and 24 May. Two birds were located 6 times between
0720 and 1645 MDT on 21 May. Seven locations were obtained between 0630
and 2120 MDT on 24 May for 2 juveniles and 1 adult. More than one morning
location were also obtained for some birds throughout the field season.
Juvenile 7051 remained within 150 m of the original location (1.8 km westnorthwest of Alkali Lake lek) throughout 21 May. A flock size of 2 birds
was determined by intentionally flushing the birds at 1645 MDT. The birds
flew north and appeared on Boettcher Lake Junction lek on 22 May (Fig. 5).
Two juveniles (7051 and 7147) in a flock of 3 birds, were accidentally
flushed at 0730 MDT on 24 May. The birds flew 1.2 km south and remained at
this location (1.7 km northwest of Alkali Lake lek) from 0915 to 2100 MDT
(Fig. 5). Periodic locations obtained on 7 and 9 June also indicate daytime
moves of less than 100 m except when disturbed (bird flew 1.1 km) (Fig. 5).
On two occasions, juveniles were present on a lek in early morning and
relocated later in the day (Fig. 5). All birds present on Boettcher Lake
Junction lek flushed for no apparent reason on 10 May. Bird 7051 was later
located 3.2 km south of the lek. On 9 June, bird 7183 flew 2.1 km northwest
of Wattenberg #2 lek between 0545 and 0930 MDT and remained at this location
until 1130 MDT when locations were discontinued.
Although based on small samples, juveniles fly considerable distances (2-3 km)
when leaving a lek. Undisturbed birds subsequently move only 150-200 m.
Long moves over 1.0 km during the day result from disturbance.
Earlier in
the breeding season juveniles may be more mobile during the day. This was
observed with 7036 averaging 2.1 km (range 0.8 to 3.7 km) between morning and
afternoon locations from 23 March to 6 April.
Juveniles remained in a
particular flock until disturbance scattered the flock or intermixing
occurred at a lek and subsequent break up resulted in changed flock composition.
Adult 6907 was located from 0720 to 1615 MDT on 21 May. Total distance
traveled was 1.5 km with the longest move (1.0 km) occurring between 0720
and 1000 and involved crossing J. C. 7. Subsequent moves were less than 0.3
km (Fig. 6). Bird 7190 was present on Boettcher Lake Junction lek at 0630
MDT on 24 May and subsequently moved 1.8 km south (4 of 7 locations).
It
returned to the lek by 2120 MDT to roost (Fig. 6). On 9 June, this individual
was accidentally flushed 1.1 km where it remained until observations were
discontinued at 1100 MDT.
Three additional movements by adults from a lek were monitored (Fig. 6).
Distances from the leks ranged from 1.0 to 2.9 km. Five afternoon locations
of adults present on leks in the morning averaged 1.3 km (range 0.0 to 2.6
km) from the lek.

�-56-

N

Bighor1J
Lek

6/7

•

(3)

s
Each concentric circle represents 1.5 km with the center located at Alkali
Lake Lek. Dates are indicated for first location of the day (e.g. 6/7) and
number of locations obtained at each site is indicated in parentheses.
Arrows indicate direction of travel.

Fig. 5. Daily activity
Colorado, 1978.

patterns

of juvenile

male sage grouse, North Park,

�-57-

N

Bighorr(")
Lek U

w

sw

S

Each concentric circle represents 1.5 km with the center located at Alkali Lake
Lek.
Dates are indicated for first location of the day (e.g. 5/21) and number
of locations obtained at each site is indicated in parentheses.
Arrows indicate
direction of travel.

Fig. 6. Daily activity patterns·
Colorado. 1978.

of adult male sage grouse, North Park,

�-58Adults had activity patterns similar to that of juveniles, i.e., relatively
long distance dispersal from the lek (although shorter distances of 0.4 km
occurred) with subsequent short moves.
Disturbance usually resulted in
moves of over 1.0 km.
Roosting
Roosting data were obtained from capture sites, evening locations (after
1700 MST) and one night when transmittered birds were located (24 May)
(Table 9). Five of 8 (62.5%) juveniles captured were roosting on leks.
Two juveniles captured off leks were trapped on 22 March or earlier (Table 1).
Capture site data are biased towards lek-roosting birds because trapping
effort was most successful on leks throughout the field season.
Early in the breeding season (22 March to 2 April) 5 of 8 (62.5%) possible
roosting sites were off leks.
On 3 consecutive evenings (31 March to 2
April) 7036 roosted on Boettcher Lake Junction 1ek where it was present the
mornings of 1 and 2 April.
No birds were present on the 1ek on 3 April due
to human disturbance.
Subsequently, 7036 moved from the study area and
contact was lost.
Three transmittered juveniles wert located at 2100 MDT on 24 May.
Birds
7051 and 7147 roosted 1.7 km northwest of Alkali Lake 1ek, the same site
used throughout the day. Both birds were present on Wattenberg #2 1ek
(4.9 km southeast) at 0645 MDT on 25 May (Fig. 5). No. 7183 roosted south
of Wattenberg #2 1ek (precise location not determined) despite being on the
1ek the mornings of 24 and 25 May.
Eight of 18 (44.4%) juvenile roosting locations were on leks (Table 9).
Highest on lek roosting occurred in April (87.5%). Although differences
Cf&lt;0.05) exist between months, more data are necessary to verify observed
trends.

Table 9. Transmittered
Colorado 1978.

male sage grouse roosting

sites, North Park,

May

Totals

Age Class

Location

February

March

April

Juvenile

On

0

1

7

0

8

Off

1

5

1

3

10

1

6

8

3

18

On

7

8

4

19

Off

1

5

4

10

8

13

8

29

Subtotals
Adult

Subtotals

�-59-

Only one adult cock was captured off a lek. This individual subsequently
moved from the study area, remaining near Aspen lek throughout the ~pring.
Early evening locations (1700 to 2000 MST) were obtained for 5 adults in
late March and early April.
Two adult cocks (6759 and 7039) were displaying
with 28 other cocks on Boettcher Lake Junction lek at 1645 MST on 31 March.
This was the only documented evening display in the study area during 1978.
No. 6759 was also present in the evening on 1, 2 and 4 April.
Between 1 and
5 April, 3 birds were located a total of 8 times in the evening: 37.5% were
on leks. Off lek locations averaged 1.8 km (range 0.8 to 2.6 km) from leks.
Three adults were roosting on leks at 2100 MDT on 24 May. One (7190) had been
1.8 km south of Boettcher Lake 12k between 1200 and 1745 MDT (Fig. 6). Four
additional off-lek locations were recorded in the evening in early May.
Adult cocks roosted on leks 65.5% of the total locations obtained (Table 9).
No difference (f&gt;0.05) existed between the 2 age classes except during March
(f&lt;0.05). Adults had strong fidelity
for lek roosting.
However, 34.5% of
the roost locations were off leks. Although not different (P&gt;0.05) than the
11.1% of the juveniles trapped off leks, extreme difficulty in locating adults
off leks was encountered.
This difficulty was partly because of poor visibility of roosting sage grouse in heavy sage off leks. Reasons why adult
cocks are more difficult to detect off leks remain unclear.
Vegetation

Analysis

Sagebrush height and canopy coverage measurements were obtained at 101 daytime feeding and loafing sites, and 40 roosting sites. Measurements of forb
and grass coverage were obtained for all locations, including the 4 leks.
Approximately 55% (55.5) of all feeding and loafing sites occurred in sagebrush with a canopy coverage of 20-50% (Table 10). This contrasts with the
80% use in these coverage classes reported by Wal1estad and Schladweiler
(1974). Sagebrush use in North Park in the 0.1-20.0% class was 2.6 times
greater than reported for Montana (Wa11estad and Schladwei1er 1974). Reasons
for this difference are unknown, but may be related to differences in
vegetation types.
Sagebrush canopy coverage averaged 29.3% (range 2.9
to 68.8%), similar to the 28 and 32% for winter and summer cover, respectively,
in Montana (Eng and Schladweiler 1972, Wallestad and Schladweiler 1974).
Approximately 80% (80.2) of the daytime feeding and loafing locations were
in sagebrush with a height of 20 to 70 cm (Table 11). The average maximum
height for the 101 locations was 45.5 cm (range 15 to 88 cm). This may
reflect actual height distribution of sagebrush in the area instead of
selection by the birds (Wallestad and Sch1adwei1er 1978).
Eighty percent of the roosting locations (including 26 on leks) occurred in sagebrush with a canopy coverage of less than 20% (Table 10). The average canopy
coverage· for the 40 locations was 11.9%, ranging from 0.7 (on Alkali Lake lek,
Table 12) to 49.3%. Six of 14 (42.9%) off-lek roosting sites were in sagebrush
with &lt;20% canopy coverage. The average canopy coverage was 24.8%. The large number
of on-lek roosting sites resulted in differences (f&lt;0.05) between feeding-loafing
and roosting sites. This contrasts with the similar vegetative parameters of

�-60-

winter cover in North Park reported by Beck (1977). Apparently, lek position
is stimulus enough to overcome normal selection to roost in heavier cover.
No difference (~&lt;0.05) existed between canopy coverage of feeding-loafing
and off-lek roosting sites.
Table 10. Frequency distribution of male sage grouse feeding-loafing
and
roosting sites by sagebrush canopy coverage class, North Park, Colorado 1978.

Canopy
Coverage.
Class 1./

Feeding-Loafing
Number of
Locations

Sites
Percent

Roosting
Number of
Locations

Sitesll
Percent

0.1-10.0

10

9.9

28

70.0

10.1-20.0

24

23.8

4

10.0

20.1-30.0

22

21.8

0

0.0

30.1-40.0

24

23.8

6

15.0

40.1-50.0

10

9.9

2

5.0

50.1+

11

10.9

0

0.0

Totals

101

100.1

40

100.0

II Expressed as percent of fully closed canopy.

II Includes

26 on-lek roosting

sites.

Twenty-nine (72.5%) roosting locations occurred in sagebrush less than 20 cm
in height (Table ll)~ Average sagebrush height was 22.5 cm (range 9 to 68 cm)
compared with 40.7 cm for the 14 off-lek roosts (~&lt;0.05).
This reflected
the apparent desire to roost on leks.
Sagebrush canopy coverage averaged 6.7% (range 0.7 to 13.4%) on leks. Grasses
(Agropyron smithii, Koeleria cristata, Bouteloua gracilis, Calamagrostis
montanensis, Poa secunda, and Stipa lettermani), fringed sagebrush (Artemisia
frigida) winterfat (Eurotia lanata), phlox (Phlox bryoides) and snakeweed
(Gutierrezia sarothrae) each contributed an average of 1.0% or more coverage.
Total vegetative coverage averaged 23.1% (range 18.0 to 32.6%) for the 4
leks (Table 12). Sagebrush height averaged 15.9 cm (range 9 to 27 cm).
Band Recoveries
Five transmittered cocks were recovered between 25 April and 9 September
(Table 1). Two adults were found dead using the telemetry equipment.
Bird
7039 appeared emaciated and was unable to fly more than 2 m on 11 and 12
April.
Subsequent transmitter failure prevented capture attempts to

�-61-

determine thebird's condition and possible adverse effects caused by the
transmitter.
Signals were received on 25 April and the bird's remains were
located in heavy sage 4.2 km north of Boettcher Lake Junction lek (1.0 km
from the 12 April location).
Appearance of the remains and the presence
of coyote scat indicated a coyote (Canis latrans) as the possible predator.
Remains of adult 7037 were located 0.2 km north of Aspen lek on 12 June.
Evidence indicated a golden eagle (Aquila chrysaetos) as the predator.
Three transmittered juveniles (7051, 7101, and 7147) were harvested in the
Lake John area on 9 September during the 1978 sage grouse hunting season.
Flock sizes were 1, 12 and 3 birds, respectively.
All had molted the radio
transmitters.
Table 11. Frequency distribution of male sage grouse feeding-loafing
roosting sites by sagebrush height class, North Park, Colorado 1978.

Class

(cm) 1/

Feeding-Loafing
Number of
Locations

Sites
Percent

.
Roostlng
Number of
Locations

and

S.ltes-2/
Percent

0.1-10.0

0

0.0

6

15.0

10.1-20.0

9

8.9

23

57.5

20.1-30.0

15

14.9

2

5.0

30.1-40.0

20

19.8

1

2.5

40.1-50.0

11

10.9

0

0.0

50.1-60.0

15

14.9

6

15.0

60.1-70.0

20

19.8

2

5.0

70.1-80.0

8

7.9

0

0.0

80.1-90.0

3

3.0

0

0.0

101

100.1

40

100.0

Totals

1/ Measurement
]j

Includes

of height of tallest living plant.

26 on-lek roosting

sites.

�-62-

Table 12.

Vegetative

cover

(%) of 4 leks, North Park, Colorado

1978.

LEK
Wattenberg

Bighorn

Boettcher
Lake 1/
Junction-

0.65

7.31

5.45

l3.36

6.69

Fringed sagebrush
(!:... frigida)

7.15

1.43

1.82

4.27

3.67

Snakeweed
(Gutierrezia
sarothrae)

0.00

2.72

0.56

1.24

1.l3

Winterfat
(Eurotia lanata)

8.14

0.10

0.66

0.58

2.37

Moss phlox
bryoides)

0.66

2.58

1.22

4.51

2.24

0.00

0.29

0.02

0.08

0.10

0.00

0.11

0.09

0.50

0.18

Grasses

4.47

6.01

6.28

7.74

6.l3

Total1/

21.15

20.57

18.00

32.62

23.09

Plant Species

Alkali
Lake

Big sagebrush
(Artemisia
tridentata)

tI2

Average

(Phlox

Bluebell
bakeri)

(Mertensia

Low daisy
pumilus)

(Erigeron

1/ Two separate parts of lek combined to form composite
1/ Includes additional species not found on all leks.

values.

�-63-

LITERATURE

CITED

Beck, T.D.I.
1977. Sage grouse flock characteristics
in winter.
J. Wildl. Manage. 41:18-26.

and habitat

selection

Braun, C. E., and T.D.I. Beck. 1976. Effects of sagebrush control on
distribution and abundance of sage grouse.
Colorado Div. Wild.
Final Rep., Fed. Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84.
Bray, O. E, and G. W. Corner.
1972. A tail clip for attaching
to birds. J. Wildl. Manage. 36:640-642.
Canfield, R. H. 1941. Application of the line interception
sampling range vegetation.
J. For. 39:388-394.

transmitters

method

in

Carr, H. D. 1967. Effects of sagebrush spraying on abundance, distribution,
and movements of sage grouse. M. S. Thesis.
Colorado State Univ., Fort
Collins.
106 pp.
Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. Schlatterer.
1960. Seasonal movements and breeding behavior of sage grouse in Idaho.
Trans. N. Am. Wildl. and Nat. Res. Conf. 25:396-407.

management

_____ , and
1963. Ecology, productivity, and
of sage grouse in Idaho. J. Wildl. Manage. 27:810-841.

Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios
from wings.
J. Wildl. Manage. 19:267-272.
1963. Observations on the breeding biology of male sage grouse.
J. Wildl. Manage. 27:841-846.
_____ , and P. Schladweiler.
1972. Sage grouse winter movements and
J. Wildl. Manage. 36:141-146.
habitat use in central Montana.
Gill, R. B. 1965. Distribution and abundance of a population of sage grouse
in North Park, Colorado.
M. S. Thesis. Colorado State Univ., Fort
Collins.
185 pp.
Jenni, D. A., and J. E. Hartzler.
1978. Attendance at a sage grouse lek:
implications for spring census. J. Wildl. Manage. 42:46-52.
Lacher, J. R, and D. D. Lacher.
Manage. 28:595-597.

1964.

A mobile cannon net trap.

J. Wildl.

May, T. A. 1970. Effects of sagebrush control on distribution and abundance
of sage grouse.
Colorado Game, Fish and Parks Div., Job Compl. Rep.,
Fed. Aid Proj. W-37-R-23, Work Plan 3, Job 8a. 23 pp.
Patterson,
Comm.

R. L. 1952. The sage grouse in Wyoming.
Sage Books, Inc., Denver.
341 pp.

Wyoming Game and Fish

�-64-

Pyrah, D. B. 1959.
Sage grouse population trend and trapping study.
Wyoming Game and Fish Comm., Job Compl. Rep.,
Fed. Aid Proj. W-50-R-8.
27 pp.
Rippin, A. B., and D. A. Boag.
1974. Recruitment to populations
sharp-tailed grouse.
J. Wildl. Manage. 38:616-621.
Robel, R. J. 1969. Movements and flock stratification within
of blackcocks in Scotland.
J. Anim. Ecol. 38:755-763.
Snedecor, G. W., and W. G. Cochran.
Iowa State Univ. Press, Ames.

1976.
Statistical
593 pp.

of male

a population

Methods.

6th Edition.

Wallestad, R., and P. Schladweiler.
1974. Breeding season movements and
habitat selection of male sage grouse.
J. Wildl. Manage. 38:634-637.

Prepared

by_~~-=-'-=~:=-_p-,---=~:....::.!.-=...:.:=-:.:.=::=--/_·_/
Steven R. Emmons
Graduate Research Assistant

Approved

by_----'t&amp;~--=V=__ _=2=:.:..... ----'~'_L..L::..=::::..::..::=__
Clait E. Braun
Wildlife Researcher

_

_

�-65-

APPENDIX

A

Sage Grouse Location

Data Form

Trans. No.

Date

Chan. No.
Freq.

Time
Band No.

Pulse Rate
Temp.

Color Code
Ppt.

Wind

%C.

C.

LOCATION
S. G.
Other

ACTIVITY

_

FLOCK COMPOSITION
Total

----- M'-----

F

----

Unk.

-----

VEGETATION

% Veg. C. ____________

Veg. Height

_

Veg. Comp.

_

TOPOGRAPHY
Slope

% Snow C.

Aspect

-----

Other

-----------------------------

_

��April 1979
-67-

JOB PROGRESS

State of

COLORADO
-----------------------------

Project

No.

REPORT

Game Bird Survey

W-37-R-32

Work Plan No.
Job No.
9c
3
--~~~--~--~--~~~--------Evaluation of the Effects of Changes in Hunting Regulations
Job Title
on Sage Grouse Populations:Evaluation
of Censuses of Females
Period

Covered:

Personnel:

1 January

through 30 September

1978.

Clait Braun, Steven Emmons, Kenneth Giesen,
Colorado Division of Wildlife.

Brett Petersen,

ABSTRACT
This investigation concerned breeding behav'Lo r of female sage grouse
(Centrocercus urophasianus) and was initiated in 1978 in North Park, Colorado.
Twenty hens (8 adults and 12 yearlings) trapped on or near 2 leks were fitted
with radio transmitters, and were located daily during April and May. Adult
females attended 1 lek for only 1 day per nesting attempt, while yearlings
attended 1 or 2 leks for 1 to 5 days (not consecutive) during 1st nesting
attempts and 1 lek for 1 or 2 days during 2nd nesting a t temp t s . Nests were
located under sagebrush (Artemisia spp.) or complexes of sagebrush and rabbitbrush (Chrysothamnus spp.). Nest sites'were 400 m to 5.0 km from leks visited,
and were located on gentle slopes of 0-30%. Height of sagebrush in areas
utilized for nesting averaged 13.5 to 59.8 cm and cover averaged 14.8 to 42.6%.
Fourteen nests were located; completed 1st clutches of adult hens averaged
7.5 eggs (6-9), while yearlings averaged 6.5 eggs (6-8). Renests of yearling
females (2) resulted in 7 eggs, and 1 adult laid 6 eggs. Length of incubation
was 26-27 days for both age classes.
Eggs were laid at the rate of 1 per 30
hours, with a 1 day interval every 3rd day. Forty-six percent of all completed
clutches hatched.
Hatchability of eggs was 72.7%.

�-68-

RECOHt1ENDATIONS

1.

A minimum
monitored

of 20 female sage grouse (12 adults, 8 yearlings)
during late Harch through late June 1979.

should be

2.

Trapping should be concentrated in the vicinity of Spring Creek lek
1, 2, and 4, and along roads connecting the 3 areas.

3.

Trapping and banding should be initiated no later than 20 Harch, depending
upon lek attendance and accessibility.
Roads should be plowed between
15-25 Harch.

4.

All transmittered birds should be located at sunrise and sunset to
ascertain lek attendance.

5.

Hens with known nests should be monitored for: movements from nests
during laying, distance traveled to feeding sites wh i Le incubating and
amount of time away from nests each day during incubation.

6.

Roosting sites of females should be located.

7.

All locations of transmittered birds should be marked with surveyor's
tape to allow for detailed vegetation analysis.

8.

Vegetation analysis should be conducted of known use areas concurrently
with other operations
to allow for accurate evaluation of vegetation
use.

9.

Hens with broods should be recaptured and have new radio units attached,
to allow greater time for recapture of chicks and evaluation of growth
rates.

10.

Known age chicks should be captured at 1 week intervals for measurement
of carpal length, primary feather growth and weight gain.

U's

�-69-

EVALUATION OF THE EFFECTS OF CHANGES
ON SAGE GROUSE POPULATIONS:EVALUATION

IN HUNTING REGULATIONS
OF CENSUSES OF FEMALES

Brett E. Petersen

Management of sage grouse has centered around 3 basic approaches: counts of
cocks on leks and estimates of nesting success and brood size. The general
life history of this .species has been described (Patterson 1952), with
particular emphasis on male lek displays (Scott 1942, Wiley 1973).
The secretive behavior of female sage grouse has been a hinderance to
collecting biological information; consequently, some aspects of their biology
is not known. Many studies have investigated nesting and brood requirements,
habitat usage, and physiographic aspects of areas utilized (Rasmussen and
Griner 1938, Patterson 1952, Klebenow 1969, May and Poley 1969, Poley 1969,
Martin 1970, May 1970, Peterson 1970, Wallestad and Pyrah 1974).
These
studies have dealt primarily with vegetation and habitat types, not with
behavioral acitivities of females.
No study has investigated daily and
seasonal attendance patterns of female sage grouse on leks. Fidelity of
sage grouse hens to leks has been mentioned by Scott (1942), Dalke et al.
(1960, 1963), Gill (1965), and Wallestad and Schladweiler (1974). Daily lek
attendance by females was mentioned by Lumsden (1968). Johnsgard (1973)
compiled data on sage grouse nesting phenology from many studies.
Data
presented in this report are from the initial year of field work, March
through August 1978, and include information on breeding season activities,
nesting phenology, and habitat selection by female sage grouse in North Park,
Jackson County, Colorado.
P. N. OBJECTIVES
The objective of this study is to supplement existing biological knowledge of
daily activity patterns of female sage grouse during early spring to summer.
Three aspects of these patterns will be examined and stratified by age (adult
and yearling): 1) daily activity patterns during the breeding period, 2)
nesting phenology and activities during egg laying and incubation, and 3)
activities during brooding and growth of chicks.
Hypotheses being tested are:
a.

Adult females visit 1 lek during the breeding

season.

b.

Adult females attend the same lek for more than 1 day during
breeding season.

c.

Adult females that have lost nests return to the lek initially

d.

Adult females that have lest nests attend the lek for 1 day during
their 2nd breeding period.

e.

Yearling

females visit more than 1 lek during the breeding

seasen.

f.

Yearling females attend more than 1 day on each lek visited
the breeding season.

during

g.

Yearling

females that have lost nests do not attempt

the

to renest.

visited.

�-70-

SEGMENT OBJECTIVES
1.

Females will be located where they roost and trapped from March through
April using spotlights and long handled nets; some hens will be captured
through use of cannon nets placed on leks. A total of 20 birds is
desired, with equal numbers of adults and yearlings.
All females will
be marked with serially numbered aluminum and color-coded plastic leg
bands, with each bird being equipped with a tail-mounted radio transmitter.

2.

Each transmittered female will be located daily between
from date of attachment until radio failure.

0430 and 0730 MST,

3.

Selected hens of both age classes will be periodically
the day to record activity patterns.

monitored

4.

Sites, located from following transmittered females, will be described as
to vegetation composition, percent cover, height of vegetation, and
physiographic features.
Sites will be grouped as to: feeding, loafing,
nesting or roosting.

5.

Counts of females and males present on each lek will be made daily in
the study area. Counts will begin in March and continue through May.

6.

Nests will be located during the egg laying period by following transmitter
equipped females.
Information will be gathered regarding nest location,
date located, and progress of nesting activity until completed or lost.
Eggs will be weighed and length and width measured.

7.

Known-age chicks will be captured with each chick being individually
marked using color-coded bands, weighed, with primary lengths being
measured.
Broods will be located and captured weekly.

8.

Data will be compiled,

analyzed

and

a progress

during

report prepared.

METHODS AND MATERIALS
Female sage grouse were located at night while roosting on leks and along
roads. Most birds were located using spotlights and captured with a long
handled net (Fyrah 1959). Additional hens were captured on leks with cannon
nets (Dill and Thornsberry 1950). All birds were banded with serial numbered
aluminum leg bands (size 14) and individual plastic color-coded bandettes.
Females were equipped with VHF radio transmitters (164 MHz frequency range)
weighing 14 gm, using tail-clip mounting (Bray and Corner 1972).
Sex and age
of birds captured were ascertained (Eng 1955), with weight and wing molt
being obtained prior to release.
Radio-equipped females were located each morning during April and May using
a portable receiver and hand-held 3-element yagi antenna.
Additional locations
were taken throughout the day on different radio-tagged hens to identify daily
activity patterns.
Slope, aspect, percent vegetation, percent sagebrush and
average height of sagebrush were recorded on a standardized form at time of
location.

�-71-

Vegetative cover measurements were made using the line intercept technique
(Canfield 1941). A total of 45.72 m (150 ft) was examined at all locations,
except nest sites and breeding areas at leks. An additional 30.48 m (100 ft)
of transects were utilized at these sites. At each location a central point
was randomly chosen with compass directions being used to locate each 15.24 m
(50 ft) line to be examined.
Vegetative height was recorded for big sagebrush
(A. tridentata), silver sagebrush (A. cana), black greasewood (Sarcobatus
v;rmiculatus) and rabbitbrush to th; nearest cm. All vegetation (by species)
was measured to the nearest 0.5 cm along transects.
Daily counts of female and male sage grouse on the 3 principal leks in the
study area were made, from 29 March through 1 June, following prescribed
techniques (Braun and Beck 1976). Two additional leks, located as the study
progressed, were counted as often as time permitted.
Radio-equipped hens were
located concurrently with lek counts.
Their locations with respect to a lek
were classified by use of triangulation of radio signals.
If transmittered
hens were determined to be on a lek, visual location was attempted.
Transmittered birds not on leks were located after completion of counts.
Locations
were plotted on U. S. Geological Survey topographic maps with movements being
measured to the nearest 0.1 km.
Description

of Area

The center of the study area was 14 km southeast of Walden, Colorado in southeast North Park, Jackson County, Colorado (Fig. 1). It is bounded on the
north by the Michigan River, on the east by Owl Creek, on the south by Owl
Ridge, and on the west by Colorado Highway 125 (Fig. 2). Elevations were
from 2463 m along the Michigan River to 2684 m on top of Owl Ridge.
Spring
Creek drains the area toward the northwest into the Illinois River.
Two
county roads divide
the area, Jackson County Road 2l(J.C. 21) (north-south),
and Jackson County Road 32 (J.C. 32) (east-west).
Jack County Road 25
(J.C. 25) (north-south) is located a maximum of 1.4 km west of Owl Creek
towards the north end of the area and is located south of Owl Creek in the
south end. Colorado Highway 14 (east-west) is located a maximum of 1.7 km
south of the Michigan River.
Five leks are known to occur in the area. The 2 main leks (Spring Creek #'s
1 and 2) are located along J.C. 21 and are 2.4 km apart.
One minor lek (Spring
Creek #4) is located south of J.C. 32, west of J.C. 21, and 2.4 km west of
Spring Creek #1 lek. A second minor lek (Eagle) is located 1.9 km NNE of
Spring Creek #4 lek. The 3rd minor lek (Peregrine) is located 0.9 km east of
Colorado Highway 125, 6.0 km northwest of Eagle lek (Fig. 2).
Small native meadows and scattered greasewood flats occur along Spring Creek,
with sagebrush-bunchgrass
dominating the remaining vegetation.
Areas along
the Illinois River, Owl Creek, and Michigan River are managed as hay meadows
by private landowners.
Detailed geographic, geologic, vegetational and
climatic features of North Park have been described by Beck (1975) and Braun
and Beck (1976). A more extensive description of the area will be completed
for the final report.

�-72-

Fig. 1.

Female

sage grouse study area, North Park, Colorado

1978.

�-73-

i

N
PEREGRINE
LEK
I
tl)
N

~

I

I

r

I
I
I

,,
I

o
o

&lt;!

0::

o
-.J

o
U

SCALE

Fig. 2.

16 mm ;; I km

Spring Creek study area, North Park, Colorado, 1978.

�-74-

RESULTS AND DISCUSSION
Trapping

and Transmitter

Life

Sage grouse hens were captured (18 with long handled nets, 2 with cannon
nets) and equipped with radio transmitters from 26 March through 10 May.
Twenty females were instrumented with 18 tail-mounted VHF t ransm.i tt.ers . Three
hens were equipped with the same radio, due to its loss by the 1st bird and
death of the 2nd hen. The transmitter remained on the 3rd bird until transmitter failure.
Inaccessibility and difficulty in capturing females resulted
in late transmitter attachment.
Seventy percent of all radios were placed 0Ii
birds after the peak of female lek attendance.
Transmitters operated an average of 62 days (range: 0-151) (Table 1). Maximum
transmitter life was determined from date of activation until the date signals
were last received.
If transmitters were known to be intermittent, their
operating life was not calculated.
Seven transmitters placed on 9 hens worked
less than 20 days or were intermittent, probably due to temperature shut off.
Two transmitters known to operate at intervals were retrieved after being
lost by their hens. These radios were tested and intermittance was attributed
to cold temperatures.
Five of the short life transmitters were retrieved
during or after the study. All radios were lost either by retrix molt or
slippage from the tail feathers.
The latter occurred because grooves in the
tail-clip were too large for adequate clamping of the feathers.
Lek Attendance
Data on lek attendance for 1st nesting attempts were collected on 8 hens (2
adults and 6 yearlings), while attendance for 2nd nesting attempts was known
for 4 birds (1 adult and 3 yearlings).
None of the other hens attended a
lek after transmitter attachment and prior to nesting
(Table 2). Nine radiotagged birds, including the 8 transmitters atta~hed in March and the 1st week
of April, were eliminated from analysis due to short working duration and
intermittent signal transmission.
Consequently, all birds equipped prior to
peak hen attendance (Table' 3) were eliminated.
Two of 6 yearling hens attended 2 leks during the 1st nesting attempt.
One
moved over 6.3 km from 1 of the main leks (Spring Creek 111) to Peregrine lek,
while the other hen moved 2.4 km between the 2 main leks. The remaining 4
yearlings and all adults attended 1 lek.
Two yearlings were observed on different dates with displaying males away
from known leks. These sites were examined later to determine if they were
possible leks. No males were observed in the area at those times.
It was
speculated that mating occurred away from leks but both yearlings were
located on leks at later dates.
Both females attended the lek closest to
these observations, with 1 hen also attending the next closest lek.
Number of days each hen visited a lek varied from 1 to 5 (Table 2). Adults
attended a lek for 1 day only, while yearlings visited from 2 to 5 days
= 2.5). Most yearling females (83.3%) visited leks on 2 days. Two
attended on consecutive days, while the other 3 had at least 1 day between
visits.
One hen visit~d a single lek on 5 different days during an 11 day
period.

ex

�Table 1.

Transmitter life and number of locations, radio-marked female sage grouse, North Park, Colorado 1978.

Date of
Capture

Band
Number

Age

Date Last
Located

Tot~l 1/
Locatlons-

26 March
1 April
1 April
2 April
2 April

4681
4682
4683
4684
4517

Adult
Yrlg.
Yrlg.
Adult
Adult

30 March
8 April
1 April
3 April
14 May

4
1

3 April
5 April
5 April
7 April

4686
4687
4688
2021

Yrlg.
Yrlg.
Adult
Yrlg.

4 April
16 April
27 August
22 May

1

8 April
10 April
10 April
10 April
11 April
11 April
14 April
24 April
24 April
2 May
10 May

4690
469l
4692
4693
4694
4697
2022
4715
4716
7192
4726

Yrlg.
Adult
Adult
Yrlg.
Yrlg.
Adult
Yrlg.
Yrlg.
Adult
Yrlg.
Yrlg.

26 June
4 August
17 July
10 May
17 July
20 April
22 May
27 August
27 August
7 September
17 July

. 1u d'lng capture slte.
.
-1/ N ot lnc
~/ Transmitter attached to 3 birds.

1/ Did not tranmit each day.

!/ No signal Eeceived for 5 days.
~/ Last location was 144 days after activation.

Transmitter Life
(Days)

8~/
7

o

o
1

8

8

10
41
63
52
41
26
53

Remarks

Transmitter fell off
Transmitter found on ground
Transmitter malfunction

1

4:))
l~/
17~/ 4/

~/
50
85
87
99
47
98

Transmitter caught in
sagebrush
Bird found dead
Transmitter found on ground
Intermittent transmitter
Transmitter caught in
sagebrush

I
-...J
\J1

I

1

9

Transmitter failure
Transmitter found on ground
Transmitter found on.ground

34
30
43
29
21

43
140
140
151
107

Transmitting distance poor

�-76-

Table 2.

Lek attendance

by female sage grouse, North Park, Colorado

1978.

Number of Days
on Leks

Number of
Leks Visited

1

1

2

0

1

1

2

1

0

3

0

2

2

2

0

2

0

0

5

1

0

1

0

0

2

6

1

3

First Nesting AttemEt
Adult
Yearling

Totals

Table 3.

Second Nesting AttemEt
Adult
Yearling

Peak counts of sage grouse on 5 leks, North Park, Colorado

1978.

Number of
Counts

High Count
of Females

Date(s)

High Count
of Males

Spring Creek 111

62

55

4 April

76

8 May

Spring Creek If2

58

29

4 &amp; 11 Ap r Ll.: 63

16 May

Spring Creek /14

34

0

3

8 April

25

8

8 April

11

9 &amp; 15
April

20

3

20 April

17

2, 6 &amp; 8
May

Lek

Eagle

(found 7 April)

Peregrine

(found 20 April)

Date(s)

Four hens (1 adult and 3 yearlings) renested.
These hens were located on only
1 lek prior to renesting.
The adult female apparently attended the same lek
where she had originally visited.
No leks were visited between capture and 1st
nesting attempt.
However, the original capture site was less than 300 m from
the lek later attended for renesting.
An additional adult female was captured
at the same time and subsequently visited the lek closest to capture prior to
her 1st nesting attempt.
Two of 3 yearling female sage grouse renesting used
the same lek prior to both nesting attempts.
The other yearling hen was
captured in May and it was not known which lek had been visited prior to the
1st nesting attempt.
Number of days between loss of nest and attendance on a lek varied from 1 to 3.
The nest of the adult hen was predated during the afternoon; 3 mornings later
she visited a lek. Yearling females visited a lek either the next morning or
2 days after their nests were lost.

�-77-

Of the 4 hens renesting, both nests were located for 2 individuals (1 adult
and 1 yearling).
Only 1 of the 2 nests was found for the other 2 yearling
hens.
Of these 2 hens, the 1st nest of one was predated and the hen returned
to the original lek visited prior to her 1st nesting attempt the following
day. Due to loss of the transmitter in dense sagebrush, no 2nd nest was
located for this female.
The 2nd female was captured on 10 May while roosting
on a lek. A brood patch was present at the time of capture, indicating an
unsuccessful 1st nesting attempt.
The 2nd nest was located on 28 May.
Counts of birds present on the 2 main and 3 minor leks were made as often as
feasible.
Counts were made from 1/2 hour before to 1 hour after sunrise from
29 March through 1 June. Two leks were newly located in 1978, one on 7 April
and the other on 20 April, 1.9 and 6.3 km from the main study area, respectively.
Both were located by radio-tracking females.
Peak hen attendance on leks occurred the week of 4 April, with a 2nd smaller
peak during the week of 8 May (Fig. 3). Numbers of hens present on leks
increased, then subsequently declined over a 2 week period.
The 2nd peak
occurred 5 days after a major snow storm (20 em total depth), which began on
the afternoon of 3 May and lasted until 6 May. A 2nd storm occurred on 17
and 18 May, followed by a smaller 3rd peak 5 days later.
Occurrence of 2
peaks has been noted by Dalke et al. (1963) and Eng (1963).
Movements
Movements of hens were segregated by age class (adults and yearlings) and
analyzed for: distance to nesting areas from lek visited, distance from nests
to loafing and feeding areas while laying and incubating, and distance from
nest to brood areas for successful females, or distance moved after nest
failure for those unsuccessful.
Lek visitation was observed for hens which laid 11 of 14 clutches.
Adults
(N = 4) moved an average of 3.3 km (range: 0.4-5.0 km) from lek visited to
the nest site, while yearlings (N = 7) moved an average 1.7 km (range:
0.6 - 3.9 km) (Fig. 4). Nests were within 2.0 km of leks attended 85.7 and
25.0% of the time for yearlings and adults, respectively.
The remaining nests
were greater than 3.5 km from leks visited.
Sixty-four percent of all nests
were located within 2.5 km of a lek compared with 68% in Montana (Wallestad
and Pyrah 1974).
Inter-lek movements, by yearlings only, occurred twice during 1978. The 2
hens each visited 2 leks. One female nested 0.6 km from the 2nd lek attended,
while the remaining yearling nested closest to the original lek. Inter-lek
movements were observed by Dalke et al. (1960) and Lumsden (1968); Wallestad
and Pyrah (1974) also observed this type of movement and measured the distance
(1.7 km) from the nest to the closest lek visited.
They observed 1 hen v Ls Lt;
4 leks, finally nesting closest to the original lek visited.
Nine of 11 nests
in this study were located closest to the lek visited.
The remaining 2 nests.
both by the same adult female, were 1.2 km closer to the nearest lek than the
1 actually visited.

�-78-

60

----)t-

-

-

Spring Creek III
-

-i(

Spring Creek 112

50

40

30

f

(J)

CJ)

r-I

cU

I \

CJ)

I

s
"'"

0
CJ)

20

\

\ /
\f

I
I

H

~;::l

\

r

4-1

(

:z;

I
I
I
I
I
10

I

l

-~

.)1.-

./

0
\0

r-I

N

C"")

If')

I

I
r-I

r-N

March

0
r-I
I
\0

If')

r-I
I
r-I
r-I

April

0
N
I
\0

r-I

If')

0

N
I

C"")

If')

.--!

\0

I

I
r-I

N

N

_
~.

0
r-I
I
\0

If')

r-I
I
r-I
r-I

0
N
I

\0

r-I

If')

N
I

r-I
C"")

I

.--!

\0

N

N

May

Fig. 3. Female sage grouse attendance on 2 leks, North Park, Colorado.
Data presented represents the high count for each 5 day interval.

�-79-

Yearling

c===J First Nest
~

Second Nest

1
Ul

.j.J

Ul
Q)

Z
4-1

0

0
1-1
Q)

'@ 4

Adult

;::l

z

~First

Nest

3
~

Second Nest

2

0.0-

0.5

0.61.0

r.a-

1.6-

LS

2.0

2.12.5

2.63.0

Distance

3.13.5

3.64.0

4.14.5

Ll.6-

5.0

(km)

Fig. 4. Distance (km) from 1ek to nest site, female sage grouse, North Park,
Colorado, 1978 (N= 11). !!

!! For females of known 1ek attendance.

�-80-

Nests of 2 adults and I yearling with unknown lek attendance were located.
One nest (adult) was 0.4 km from a lek with all locations obtained being
nearest this lek. The other nests were located 0.4 and 0.6 km closer to
Spring Creek #2 than #1. Both females remained in the vicinity of their nests.
One additional yearling hen whose nest was not located prior to transmitter
failure utilized the same area f o'r 16 days following Lek attendance.
The
center of activity was 1.6 km from the lek. The nest may have been located
in this area, however, the clutch was not completed prior to transmitter
failure.
This distance was only 0.1 km less than the average for yearlings
with known nest locations.
Renests of 1 adult and 1 yearling female were located.
The 1st nesting
attempt, of the adult, 4.2 km from the lek visited, was destroyed by a
predator during a snow storm in early May. The 2nd attempt was 3.6 km from
the same lek and 0.9 km from the original nest site. This nest was predated
in early June. No additional data were gathered on this hen due to transmitter failure prior to nest destruction.
The 1st attempted nest of the
yearling hen was 1.6 km from the lek visited.
This nest was abandoned in
May, possibly because of predator activity as 1 of 6 eggs was missing.
The
2nd nesting attempt, which was successful, was 1.8 km from the same lek and
0.3 km from the original nest.
No data on movements during laying were obtained for 2 of 14 hens due to
transmitter malfunction.
Both radios transmitted intermittently and only
nest locations were obtained.
One transmitter WaS found entangled (antenna)
in the dense sagebrush above the nest. This nest was abandoned prior to
clutch completion.
Movements of females during egg laying were normally restricted to within 0.6
km of the nest site. The longest movements of yearlings (N = 8) averaged 0.7
km (range: 0.4 - 1.4 km), while for adults (N = 4), longest movements averaged
1.1 km (range: 0.4 - 2.0 km) (Fig. 5). Long movements during this period were
uncommon for both age classes.
Observations during laying indicated an affinity to specific areas.
In the
case of 2 long feeding moves by an adult, the areas utilized were less than
0.1 km apart, but 4 days apart in use. Movements were not random and centered
around 2 or 3 specific areas. Many locations were within 0.1 km of each other.
Length of movements during egg laying by renesting hens decreased.
Movements
of adults decreased from 0.6 to less than 0.4 km, on 1st and 2nd nesting
attempts, respectively, while movements of yearlings decreased from 0.5 to
0.4 km.
Onset of incubation limited movements of hens to feeding excursions.
Feeding
areas were no further than 0.2 km from a nest, with the majority usually
within 0.1 km. Feeding occurred once a day for each hen, either prior to
sunrise or after sunset.
These observations contradict prior data indicating
feeding at both times (Gill 1965). Time of feeding did not change during
incubation.
One yearling hen was observed to feed during the day on 1
occasion, however, this movement was only a few meters from the nest.

�-81-

Yearling

c===J First Nest

5

~

Second Nest

4

3

2

1
CIl

P
(l)

::c:
4-1
0

0

l-l
(l)

..a
S

;:l

Z

~First

Adult

~

4

Nest

Second Nest

3

2

0.00.5

0.61.0

1.11.5

1. 62.0

2.12.5

Distance (km)
Fig. 5. Distance traveled away from nest site during egg laying, North Park,
Colorado 1978 (N = 12).

�-82-

Data on movements of hens after completion of nesting, either successfully
with broods or by themselves after nest loss, were limited to 7 birds (3
adults and 4 yearlings).
Locations of these hens were obtained during early
to late summer at 1 to 2 week intervals.
Successful hens (2 adults and 2 yearlings) with broods traveled toward ridge
tops and plateaus within 1 or 2 days of hatching.
This movement may be
attributed to the moist conditions prevailing in 1978. Klebenow (1969) noted
that precipitation and its variability in Idaho accounted for differential
movements between years.
In drier years in North Park hens with broods
usually moved toward meadows and pastures (Gill 1965, Poley 1969, May and
Poley 1969, May 1970). Distances traveled varied from 0.6 to 2.3 km, with no
differences noted between yearlings and adults.
Length of time spent on
ridges depended upon hatching date relative to onset of vegetation desiccation.
The earliest brood hatched remained on ridges for 9 weeks, compared with 1 of
the latest broods hatched, which remained for 4 weeks.
During this time,
movements along ridges varied from less than 0.1 km in 4 weeks to 1.0 km in
2 days. All successful hens with broods eventually moved to meadows on the
east side of the study area.
Movements by unsuccessful hens (1 adult and 2 yearlings) varied greatly,
possibly due to the moist conditions.
One unsuccessful yearling hen, which
abandoned its nest early, moved the greatest distance of this group. Daily
movements of 2.6 km for short periods of 2 to 3 days were common.
This hen
finally settled along the Illinois River, 4.7 km from the nest site. The
other 2 unsuccessful hens moved daily, no more than 1.5 km, prior to settling
along Owl Creek. Movements from nesting sites to meadows were 2.8 and 2.1 km.
These 2 hens moved into meadow areas in less than 2 weeks, compared with 5
weeks for the 1st yearling hen.
Nesting
Fourteen nests were located during 1978, with data collected on: date nest
was located, number of eggs at time of location, condition of nest (construction), date each additional egg was laid, number of eggs at completion of
clutch, date incubation started, and hatching date. Information was gathered
on nests by checking at frequent intervals and noting changes, if any, while
the hen was foraging some distance from the nest.
Eight of 14 nests were located while hens were still laying.
Five of the
remaining 6 were being incubated when found (Table 4). The remaining nest
was located after abandonment, due to the transmitter becoming entangled in
the sagebrush covering the nest. This problem was also reported in Montana
(Walles tad and Pyrah 1974). One additional nest was abandoned because of
observer interference.
This nest was located on the apparent 1st day of
incubation.
The hen flushed from the nest and was not located again on the
nest or within 1 km of it. Eight other hens were flushed from nests, but
all returned after the observer left the area.
Ten of the 14 nests had known dates for last lek visitation by their hens.
Number of days from last lek visitation to onset of egg laying was highly
variable.
The minimum was 3 days (1 adult), while the maximum was 13 days
(Table 5). The average number of days between apparent mating and laying was
8.5 days, with 1st nesting attempts averaging 8.1 and 2nd nesting attempts

�Table 4.

Timing of reproductive activities of female sage grouse, North Park, Colorado 1978.

Age

Band
Number

Lek(s) Attended

Adult

4517

Adult

Date Attended
Lek Last

Date Nest
Located

Clutch
Size

Eggs
Hatched

Unknown

Unknown

14 May

4

-

1st

4688

Spring Creek 112

6 April

27 April

9

8

1st

Adult

4691

Spring Creek III

6 May

15 May

6

4

1st

Adult

4692

Spring Creek III

10 April

30 April

7

-

1st

Adult

4692

Spring Creek 111

8 May

22 May

6

-

2nd

8

6

1st

Nesting Attempt

Adult

4716

Unknown

Unknown

3 May

Yr1g.

2021

Spring Creek 112

16 April

5 May

6

-

1st

Yr1g.

2022

Spring Creek III
Peregrine

20 April

7 May

8

-

1st

Yrlg.

4690

Spring Creek III

20 April

10 May

6

-

1st

Yr1g.

4690

Spring Creek III

15 May

26 May

7

7

2nd

Yr1g.

4694

Spring Creek tf2

17 April

2 May

7

3

1st

Yr1g.

4715

Spring Creek 111
Spring Creek tf2

10 May

22 May

6

-

1st

Yr1g.

4762

Spring Creek III

12 May

28 May

7

4

2nd

Yr1g.

7192

Unknown

Unknown

3 May

6

-

1st

-

Remarks

Clutch not completed,
man caused abandonment

I
00
W
I

�Table 5.

Sage grouse nesting phenology, North Park, Colorado 1978.

No. of Days
Between
Breeding and
Egg Laying

Date
Layingl/
Began-

No. of
Days to
Complete
Clutch

Date
Clutch
Completed

No. of Days
Between
Completion
of Clutch and
Incubation

Date
Incubation
Began

Length of
Incubation
(Days)

Date
Hatched

Nest
Attempt

27 April

26-27

23 May

1st

2

20 May

27

15 June

1st

29 April

1

30 April

Destroyed

6 May

1st

9

29 May

1

30 May

Destroyed

12 June

2nd

1 May

11

12 May

1

13 May

27

8 June

1st

10

26 April

8

3 May

2

5 May

Destroyed

13 May

1st

20 April

6

26 April

11

6 May

1

7 May

Destroyed

23,May

1st

4690

20 April

13

1 May

7

8 May

2

10 May

Abandoned

14 May

1st

Yrlg.

4690

15 May

6

21 Hay

8

29 May

1

30 May

27

25 May

2nd

Yrlg.

4694

17 April

12

29 April

10

9 May

1

10 May

27

5 June

1st

Yrlg.

4715

10 May

4

14 May

8

21 May

1

22 May

Abandoned

22 May

1st

Yrlg.

4762

12 May

10

22 May

9

31 Hay

0

31 May

27

26 June

2nd

Yrlg.

7192

Unknown

-

1 May

8

9 May

2

11 May

Abandoned

19 May

1st

Age

Band
Number

Date Attended
Lek Last

Adult

4517

Unknown

Adult

4688

6 April

Adult

4691

6 May

3

9 May

9

18 May

Adult

4692

10 April

9

19 April

11

Adult

4692

8 May

12

20 May

Adult

4716

Unknown

Yrlg.

2021

16 April

Yrlg.

2022

Yrlg.

1/ Dates calculated by backdating using 3 days for each 2 eggs laid.

I
0&gt;

"'I"

�-85-

averaging 9.3 days. Length of time from possible mating to laying decreased
for yearling 4690. Her 1st nest was initiated 13 days after last visiting
a lek, while her 2nd nest was started within 6 days of loss of the 1st nest.
Timing of egg laying was constant.
If an egg was laid in the early morning
on 1 day, the next egg was laid the following afternoon.
Thus, eggs were
laid 30 hours apart, followed by 1 day off, with the cycle being repeated.
However, egg laying may not always commence in the morning, but possibly in
the afternoon.
In 4 cases (1 adult and 3 yearlings) more than 1 day occurred
between cycles.
In these situations a 2 day rest was observed.
This could
account for the appearance of some back-dated 1st eggs being laid in the
afternoon.
Data obtained appear to indicate that 2 eggs were laid every 3
days.
The interval from completion of a clutch to initiation of incubation was
observed for 8 females.
This interval averaged 1.2 days (range: 0-2 days)
(Table 5). The median interval from clutch completion to incubation
initiation was 1 day (n = 7).
Data from 13 nests were used to calculate clutch size for 1st or 2nd nesting
attempts.
Ten nests were classified as 1st nesting attempts, while the
remalnlng 3 were known to be 2nd attempts (Table 4). The nest of adult 4691
was designated a 1st nesting attempt, as only 1 nest was found.
Another
adult (4692) originally captured at the same time was attempting a 2nd nest
concurrently with 469l's apparent 1st attempt.
Average clutch size for all
completed nests during 1978 was 6.8 eggs (Fig. 6).
First nesting attempts of 4 adult and 6 yearling females had average clutch
sizes of 7.5 and 6.5 eggs, respectively.
Of the 10 nests, 4 were successful (3 adults and 1 yearling), 3 were abandoned (yearlings), and 3 were
destroyed by predators (1 adult and 2 yearlings).
Second nesting attempts of 1 adult and 2 yearling females had average clutch
sizes of 6.0 and 7.0 eggs, respectively.
Of the 3 nests, 2 were successful
(yearlings) and 1 was destroyed by predation (adult). Data for both 1st and
2nd nesting attempts were available for 2 hens (1 adult and 1 yearling).
Clutch size of the adult decreased from 7 to 6 eggs, while clutch size of
the yearling increased from 6 to 7 eggs. The remaining yearling was captured
after loss of her 1st clutch (based on the presence of a brood patch) and
prior to attending a lek for a 2nd nesting attempt.
Length of incubation was similar for all successful nests.
Five hens were
checked daily from initiation of incubation until hatching.
The nest of a
6th hen was believed to have been found on the 1st day of incubation.
All
hens hatching their clutches left the nest with chicks on either the 26th
or 27th day of incubation.
In 2 situations, hatching was known to begin on
the 26th day, with the broods leaving the nest either· the night of the 26th·
or morning of the 27th day. Movements away from the nest appeared to occur
at dawn or dusk.
Hatchability of eggs varied greatly during 1978, with 72.7% of all eggs
hatching (n = 80). Eggs of adults had a higher hatching rate (78.3%) (N = 33)
than those of yearlings (66.7% (N = 47). Hatchability of eggs in individual
clutches ranged from 67 to 89% for adults and from 42 to 100% for yearlings.
Low hatchability appeared to be related to snow storms that occurred during

�-86-

Yearling

c===J First Nest
~

Second Nest

4

3

2

CJJ

1

.jJ

CJJ
Q)

Z
4-l
0
!-I

0

Q)

~;::l

z

4
Adult

3

~First

Nest

~second

Nest

2

5

Fig. 6.

1978.

6

7
Number of Eggs

8

9

Number of sage grouse nests by clutch size, North Park, Colorado

�-87-

laying.
Hens with clutches completed prior to snow storms had higher
hatchability (89%) than hens still laying when snow storms occurred (42%).
Hatchability of clutches, initiated after snow storms, was higher than those
only partially complete when snow storms occurred.
Egg Size
During laying and incubation, eggs were weighed and measured.
Seventy-one
eggs were weighed prior to incubation, yielding an average weight of 46.0 gms
(range: 40-55 gms). Eggs of adults (N = 24) averaged 49.0 gms (range:
44-55 gms), while eggs of yearlings (N = 47) averaged 44.6 gms (range: 40-50
gms). Eggs of an adult hen not examined until the 19th day of incubation
weighed an average of 38.4 gms (range: 38-39 gms).
If weights of these eggs
were average and had typical weight loss, then 10.6 gms of water (21.6% of
total weight) were lost during the 1st 19 days of incubation.
Egg length and width varied greatly.
Egg dimensions (n = 80) averaged 55.3 x
38.6 mm. Average length and width for adults and yearlings were 55.8 x 39.2
and 55.0 x 38.1 mm, respectively.
The range in length and width for eggs of
adults was 53.0-58.9
and 38.0-40.7 mm, respectively (Fig. 7). Ranges of
length and width for eggs of yearlings were 53.5-56.8 and 37.6-38.9 mm,
respectively.
Growth of Chicks
Chicks from 5 (2 adults and 3 yearlings) of 6 broods were captured the day
they left the nest; 22 chicks were weighed.
No other measurements were
obtained at that time. Average weight for all chicks was 31.8 gms. Chicks
of yearling hens averaged 31.2 gms (range: 24-33 gms), while chicks of adult
hens averaged 32.5 gms (range: 29-36 gms). Average chick weight per brood
ranged from 30.0 (1 adult and 1 yearling) to 34.2 gms (adult).
Chicks with radio-marked females were followed throughout the summer.
Locations were made at 1 and 2 week intervals and continued until loss of the
transmitter or brood break up. Eleven chicks were recaptured, of which 4 were
recaptured twice.
Chicks were banded with color-coded, numbered bandettes
when large enough to retain them (8 days of age). Two of the 6 broods were
not located after the 1st day, due to transmitter failure.
Broods of adult
hens (N = 2) supplied data on growth for 8 of the 11 birds.
Two other broods
(1 adult and 1 yearling) supplied 2 and 1 recapture each. Measurements were
taken of weight, carpal and primary lengths at each capture.
Accurate weights (+2 gms) were recorded up to 22 days of age. Weight gain
was estimated using mean weight of chicks leaving nests compared with weights
at time of capture.
Weight gain per day ranged from 1.5 gms/day from 0-8
days of age to 7.8 gms/day from 0-18 days of age (Table 6). Average weight
gain for 8 chicks from 18 to 22 days of age was 6.6 gms/day.
This increased
to 10+ gms/day at an interval of 18 to 31 days of age.
Wing measurements
(±l mm) were recorded for 11 chicks (Table 6). Emergence of
juvenile primary IX pegan at 11 days of age, while juvenile primary X emerged
at about 22 days of age. Replacement of juvenile primary I with adult primary
I began at approximately 23 days, after 22 and before 31 days of age. Juvenile
primary II was replaced "at approximately 31 days, while juvenile primary III
was molted between 31 and 45 days, closest to 38 days of age. Juvenile
primary IV was replaced at 45 days (Table 6).

�-88-

A

Adult

Y

Yearling

o

Second Clutch

Y = Clutches for
l
1 Individual
60.0

59.0

58.0

57.0

Y

,......

A©

A

~

'-"
..c 56.0
.w
co

t=:
&lt;lJ

A

VI

H

55.0

V

54.0

y
53.0

A

36.0

37.0

38.0

39.0

40.0

41.0

42.0

43.0

Width (nun)

Fig. 7. Average egg length and width of sage grouse clutches, North Park,
Colorado, 1978 (N = 12).

�Table 6.

Juvenile sage grouse growth data, North Park, Colorado 1978.

Age in
Days

Band
Color-Number

Weight
(gms)

Average Weight
Gain (gros/day)

Carpal
Length (mm) X

IX

VIII

Primary LenBth (mm)
VI
IV
VII
V

III

II

I

8

Blue 10

44

1.5

-

E

E

8

40

48

52

51

49

48

45

11

White 14

75

3.9

88

E

Q

19

53

59

65

67

65

63

61

11

White 94

76

4.0

94

E

E

20

54

62

67

69

68

67

64

18

Yellow 64

166

7.5

132

E

24

50

85

95

97

96

92

86

79

18

Yellow 63

149

6.5

130

E

25

49

85

96

97

95

90

85

78

31

Yellow 63

300+

-

181

24

54

106

121

129

127

118

102

100

56~/

18

Yellow 57

172

7.8

132

E

30

50

82

92

95

96

90

85

78

31

Yellow 57

300+

-

177

22

53

107

129

133

127

124

109

E

53~/

21

Red 13

159

6.1

-

E

11

68

94

99

104

102

95

83

76

22

Red 13

162

5.9

141

E

10

72

97

103

105

103

94

83

75

22

Red 18

156

5.6

143

E

13

78

99

104

106

103

93

81

72

22

Red 19

184

6.9

151

E

9

70

98

107

109

110

105

95

78

22

Red 38

166

6.1

142

2

17

75

98

104

106

103

93

79

71

31

Yellow 65

300+

-

181

24

54

106

121

129

127

118

102

100

5~/

45

Red 18

300+

-

232

82

120

162

169

161

151

E

58~/ 107~/141~/

~/ Adult primary.

I
00
1.0
I

�-90Mortality
One radio-equipped yearling female was found dead on 4 April, the morning
after release, less than 0.2 km from the release site. Death appeared
to be caused by internal injuries sustained from landing in dense
sagebrush.
Tracks in the snow indicated the bird had walked 10 m prior to
collapsing.
The hen was frozen, indicating that death occurred during the
night.
No additional dead radio-tagged hens were located, although 1 yearling was
believed to have been predated during incubation.
This hen had incubated 16
days prior to nest predation.
All 7 eggs had been eaten, but no blood or
large amounts of feathers were found in the vicinity of the nest.
No signals
were received after the nest was destroyed on 23 May.
It is believed the
transmitter was destroyed when the hen was predated.
No radio-marked hens were recovered during the 9-24 September hunting season.
One hen banded, but not radio-equipped,
in the study area in 1978 was harvested
along the Illinois River.
Vegetation

Analysis

Sagebrush height and cover measurements were taken at 127 locations.
Measurements were also obtained on coverage of forbs (by species) and grasses at all
but nest sites.
P~ior to examination areas were classified as: breeding (leks),
nesting, roosting, feeding, and loafing and feeding sites.
Breeding sites on
3 leks were described, as were 14 nest sites. Roosting sites (N = 16) consisted of locations of hens during the night.
The remaining 94 sites were
segregated into feeding (N
25) areas utilized by hens during incubation,
and loafing and feeding (N = 69) areas used by females prior to incubation.
Transmittered hens attended only 3 of the 5 leks in the study area in 1978;
therefore, vegetation of mating areas on these 3 leks was classified.
Sagebrush averaged 5.3 cm (range: 3.2-7.2 cm) in height and covered 7.3% (range:
3.9-10.3%) of the area (Table 7). All mating areas had less than 26% (range:
21.2-25.8%) ground cover and had sagebrush 20 cm in height.
This taller
sagebrush may supply some protection for hens.
Nesting sites were described from measurements of sagebrush height and cover
only, due to the difficulty of ascertaining understory species coverage.
Average sagebrush height surrounding nests was 34.9 cm (range: 11.1-59.8 cm).
Approximately
57% (57.1) of the nests were located in areas of sagebrush
taller than 30 cm (Table 8), while all nests were located under sagebrush
bushes or complexes of sagebrush and rabbitbrush with heights ~30 cm.
Roosting areas were typified by sagebrush less than 15 cm in height with 85.7%
being in sagebrush area 210 cm in height (Table 9). Average height was 5.9 cm
(range: 3.7-13.7 cm) with only original trap sites examined.
Feeding areas
had sagebrush height of less than 30 cm with 72% being in areas 215 cm in
height (Table 9). Average height was 12.9 cm (range: 4.6-26.4).
Sixty-nine loafing and feeding locations were identified and examined from
over 200 locations classified.
These areas showed the greatest variability.
No areas were utilized where sagebrush had average heights of less than 5
cm. The greatest usage in sagebrush areas was in the range of 15.1 to 30.0
cm (53.6%) (Table 10). The most commonly utilized class of sagebrush height
was 15.1-20.0 cm (24.6%). Areas of taller sagebrush were commonly used during
inclement weather.

�-91-

Table 7. Vegetative
Colorado 1978.

cover

(%) of sage grouse mating sites, North Park,

Vegetative
Species

Sagebrush
Fringed

(Artemisia

sagebrush

Peregrine

3.9 (3.2)1./

10.3(7.2)1./

.9

1.2

3.4

1.8

1.7

3.0

4.1

1.2

0.1

0.3

11.9

10.2

8.8

0.1

0.1

26.0

21.1

(~.frigida)

(Gutierrezia

Moss phlox

(Phlox bryoides)

Low daisy

(Erigeron pumilus)

sarothrae)

Grasses
(Sedum stenoEetalum)

2/

Totals-

1./ Average height

1/ Including

7.6(5.5)1/

spp.)

Sankeweed

Stonecrop

Leks
Spring Creek #2

Spring Creek III

23.4

(cm).

species not listed that occur on only 1 lek.

Table 8. Average height (cm) of sagebrush
North Park, Colorado 1978.

Average Height (cm)
Surrounding Nests

Number

surrounding

of Nests

sage grouse nests,

Percent

of Nests

10.0-19.9

2

14.3

20.0-29.9

4

28.6

30.0-39.9

3

21.4

40.0-49.9

3

21.4

50.0-59.9

2

14.3

14

100.0

Totals

�-92-

Table 9. Average height (cm) of sagebrush at roosting
female sage grouse, North Park, Colorado 1978.

and feeding areas of

Number of
Roosting Sites

Percent

Number of
Feeding Sites

Percent

0.0-5.0

4

25.0

2

8.0

5.1-10.0

10

62.5

6

24.0

10.1-15.0

2

12.5

10

40.0

15.1-20.0

3

12.0

20.1-25.0

3

12.0

25.1-30.0

1

4.0

25

100.0

Average
Height (cm)

16

Totals

100.0

Table 10. Frequency of sagebrush height classes used by female sage grouse
for loafing and feeding areas, North Park, Colorado 1978.

Classes

(cm)

Number

of Locations

Percent

0.0-5.0
5.1-10.0

2

3.0

10.1-15.0

9

13.0

15.1-20.0

17

24.6

20.1-25.0

10

14.5

25.1-30.0

10

14.5

30.1-35.0

6

8.7

35.1-40.0

6

8.7

&gt;40.1

9

13.0

Totals

69

100.0

�-93-

•
LITERATURE

CITED

Beck, T.D.I.
1975. Attributes of a wintering population of sage grouse,
North Park, Colorado.
M. S. Thesis. Colorado State Univ., Ft. Collins.
49 pp.
Braun, C. E., and T.D.I. Beck. 1976. Effects of sage brush control on
distribution and abundance of sage grouse. Colorado Div. Wildl. Final
Rep., Fed. Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84.
Bray, O. E., and G. W. Corner.
1972. A tail clip for attaching
to birds.
J. Wildl. Manage. 36:640-642.
Canfield, R. H. 1941. Application of the line interception
sampling range vegetation.
J. For. 39:388-394.

transmitters

method

in

Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. Schlatterer.
1960. Seasonal movements and breeding behavior of sage grouse in Idaho.
Trans. N. Am. Wildl. and Nat. Res. Conf. 25:396-407.
1963. Ecology, productivity, and manage----- , and
ment of sage grouse in Idaho. J. Wildl. Manage. 27:810-841.
Dill, H. R., and W. R. Thornsberry.
1950. A cannon-projected
capturing waterfowl.
J. Wildl. Manage. 14:132-137.

net trap for

Eng, R. L. 1955. A method for obtaining sage grouse age and sex ratios
from wings.
J. Wildl. Manage. 19:267-272.
1963. Observations on the breeding biology of male sage grouse.
Wildl. Manage. 27:841-846.

J.

Gill, R. B. 1965. Distribution and abundance of a population of sage grouse
in North Park, Colorado.
M. S. Thesis.
Colorado State Univ., Ft.
Collins.
185 pp.
Johnsgard, P. A. 1973. Grouse and quail of North America.
Nebraska Press. Lincoln.
553 pp.

Univ. of

Klebenow, D. A. 1969. Sage grouse nesting and brood habitat
Wildl. Manage. 33:649-662.

in Idaho.

Lumsden, H. G. 1968. The display of the sage grouse.
and Forests.
Res. Rep. 83. 94 pp.

J.

Ontario Dept. Land

Martin, N. W. 1970. Sagebrush control related to habitat and sage grouse
occurrence.
J. Wildl. Manage. 34:313-320.

�-94-

May, T. 1970. Effects of sagebrush control on distribution and abundance
of sage grouse.
Colorado Div. Wi1d1. Job Comp1. Rep. Proj. W-37-R-23,
Work Plan 3, Job 8a. pp. 115-138.
, and B. Poley.
1969. Spring and summer movements of female sage
grouse in North Park, Colorado.
Biennial Western States Sage Grouse
Workshop.
6:173-178.
Patterson, R. L. 1952.
Denver.
341 pp.

The sage grouse of Wyoming.

Sage Books, Inc.

Peterson, J. G. 1970. The food habits and summer distribution of juvenile
sage grouse in central Montana.
J. Wi1dl. Manage. 34:147-155.
Poley, B. 1969. Effects of sagebrush control on distribution and abundance
of sage grouse.
Colorado Div. Wild1. Job Compl. Rep. Proj. W-37-R-22,
Work Plan 3, Job 8a. pp. 61-86.
Pyrah, D. B. 1959. Sage grouse p0pulation trend and trapping study.
Wyoming Game and Fish Corom. , Job Compl. Rep. Fed. Aid Proj. W-50-R-8.
pp. 38-64.
Rassmussen, D. I., and L. A. Griner.
1938. Life history and management
studies of the sage grouse in Utah, with special reference to nesting
and feeding habits.
Trans. N. Amer. Wi1dl. Conf. 3:852-864.
Scott, J. W.

1942.

Mating behavior

of the sage grouse.

Wa11estad, R., and D. Pyrah.
1974. Movement and nesting
in Central Montana.
J. Wildl. Manage. 38:630-633.
___

Auk 59:472-498.
of sage grouse hens

, and P. Schladweiler.
1974. Breeding season movements and habitat
J. Wild1. Manage. 38:634-637.
selection of male sage grouse.

Wiley, R. H. 1973. Territoriality and non-random mating in sage grouse,
Centrocercus urophasianus.
Animal Behavior Monographs 6, Part Two.
83 pp.

Prepared

by

.:D~ e. ..AJ~
Brett E. Petersen
Graduate Research

Approved

Asst.

by_---Lt&amp;~V=-=-· ~2.=___. 4-'~~~~C1ait E. Braun
Wildlife Researcher

�-95-

April 1979

JOB FINAL REPORT

COLORADO

State of
Project

No.

Work Plan No.
Job Title
Period Covered:

Game Bird Survey

W-37-R-32,
3

Job No.

10

Investigations of the Distribution and Status
of Sagebrush and Sage Grouse in the Moffat County Area
January 1, 1976 to March 31, 1978

Personnel:
Carolyn Atchley, Kathleen Bieniasz, C1ait Braun, Dan Bricco (BLM),
Charles Brown, John Corey, Larry Crooks, John Ellenberger, Dale F1enthrope,
Howard Funk, Charles Hector, Richard Hoffman, William Howard, Roland Kufe1d,
Ann Leckler, Robert Mangus, Deirdre Plummer, William Roland, George Steele,
Louis Vidakovich, Claude ItJhite,Charles Woodward, and Donald Hoffman.
ABSTRACT
Objectives of this investigation were to ascertain the distribution and status
of sagebrush (Artemisia spp.), delineate the distribution of sage grouse
(Centrocercus urophasianus), and identify past ';'ndpresent trends of sage
grouse populations within the Moffat County Area. Field work was started in
January 1976 and concluded in March 1978. Background information gathered
from management surveys indicated peak populations of sage grouse may have
occurred in the Moffat County area in 1969 and lows in 1965 and 1975, indicating a possible ten-year population cycle. However, count data gathered
over the years were not consistently obtained and new grounds were not searched
for on a systematic basis in the vicinity of old grounds.
Thus, the information on number of birds present between years as we LL as on possible peaks
or declines in the total population are suspect. Approximately three percent
of the federal land in Moffat County has been altered intentionally through
various habitat manipulation projects in past years. Within best (Class 1)
ranges, approximately 25 percent of lands owned or controlled by private landowners has been altered to bring land under cultivation or to improve pastures
for grazing purposes, based upon a ten percent sample of the land area.
Within areas considered as former range for sage grouse (where agriculture
has largely eliminated sagebrush habitat), the percentage of land altered
was approximately 74 percent, based upon a 13 percent sample of the land area.
Big sagebrush (/'::.
tridentata) ranges within Moffat and western Routt counties
were separated into three classes of range based upon quality and potential
of habitats.
Class 1 range (highest quality habitat and highest potential for
sage grouse) consisted of 1,887 mi2 (488,748 ha) in Hoffat County and III mi2
(28,750 ha) in western Routt County. Class 2 range (fair quality habitat and
fair potential for sage grouse) consisted of 520 mi2 (134,684 ha) in Moffat·
County and none in western Routt County.
Class 3 range (poorest quality
habitat and poor potential for sage grouse) consisted of 434 mi2 (112,409 ha)
in Moffat County and none in western Routt County.
Former ranges, considered
as Class 4 non-ranges, consisted of areas where dry land cultivation has reduced
former sage grouse habitats to mere remnants.
These consisted of 168 mi2
(43,513 ha) in eastern Moffat County and 169 mi2 (43,772 ha) in western Routt

�-96-

ABSTRACT

(Continued)

County.
In addition, 1,902 mi2 (492,633 ha ) of Moffat County we re nonsagebrush habitat and were also considered as non-ranges for sage grouse.
More strutting grounds (59) were located and counted in Moffat and western
Routt counties during 1977 than had been known to exist in any previous year.
During 1977, 1,977 cocks and 2,441 total birds were counted on these 59
active strutting grounds for an average of 33.51 cocks per ground and an
average of 41.37 total birds per ground.
One additional strutting ground
was located by a Wildlife Conservation Officer during the study, bringing
the total number of active strutting grounds in 1977 to 60. Brood counts
along established routes were revived in 1976 and 1977 within the Moffat
County area.
In 1976, L 37 birds per mile (0.85/km) were counted in 412.6
miles (663.9 km) driven along ten brood count routes.
In 1977, 1.27 birds
per mile (0.79/km) were counted in 905.1 miles (1,456.3 km) driven along 11
brood count routes.
Eleven important wintering areas were identified through
winter period searches of potential habitats.
These contained approximately
182 mi2 (47,139 ha) of winter range. Two important brood rearing ra~ges were
identified through brood counts and miscellaneous brood observations.
These
contained approximately 146 mi2 (37,815 ha) of summer range. A total of 748
sage grouse wings was collected in 1976, and 717 in 1977, using 16 wing barrel
collection stations, miscellaneous check station collections, hunter contacts,
and wing survey envelopes.
Fifty percent of all sage grouse wings collected
in 1976 and 1977 came from the large area of Class 1 range northwest of
Craig (west of Colorado Highway 13, north of U. S. Highway 40, and east of
the Little Snake River).

�-97-

INVESTIGATIONS OF THE DISTRIBUTION AND STATUS
OF SAGEBRUSH AND SAGE GROUSE IN THE MOFFAT COUNTY AREA
Donald M. Hoffman

Distribution and status of sage grouse are intimately associated with the
distribution and status of sagebrush lands since sagebrush furnishes most of
its needs.
With extensive sagebrush-grassland
rangelands, Moffat County
provides year-long habitat requirements for many sage grouse, the dominant
upland gamebird.
The Moffat County area provides many thousands of man-days of recreation for
hunters pursuing sage grouse as well as other game species as elk, mule deer,
antelope, blue grouse, sharp-tailed grouse, geese, ducks, doves, and cottontails.
The Moffat County area is also important for agricultural and energy production. Agricultural activities date back to the homestead period following
World War I. Vast quantitites of coal, oil, and natural gas occur, which are
economically profitable to exploit.
In recent years, mining and drilling
activity has greatly increased, resulting in reduced wildlife habitats.
This
increase in the exploitation of natural resources was accompanied by more
restrictive seasons for game animals, especially sage grouse.
Very little
was known about relationships between apparent declines in populations of
sage grouse and increased habitat disturbance.
In view of the vastly expanded
search for and the exploitation of energy resources and the lack of information upon which to base management recommendations for sage grouse hunting,
it became necessary to study sage grouse populations and alterations to their
habitat.
Historical records of sage grouse distribution, populations, and hunting
seasons, along with sage grouse studies from 1939 through 1961 in Colorado
were summarized by Rogers (1964). Specific procedures, map locations, and
time periods for securing strutting ground counts, brood counts, and check
station data were outlined and turned over to management personnel in the
early 1960's for implementation.
Numbers of strutting ground counts conducted
from 1961 to 1977 by Wildlife Conservation Officers (W.C.O.'s) have varied
considerably.
Brood counts by W.C.O.'s have been too sporadic within recent
years to be of value, with only a very few exceptions.
The Cedar Mountain
Grouse Check Station has not been operated since 1973 and the Blue Mountain
Grouse Check Station has not been operated since 1958. With the low numbers
of sage grouse counted on strutting grounds in Moffat County in 1975 (663
total birds and 539 cocks) plus only 17 active strutting grounds censused,
administrators within the Division of Wildlife became concerned about populations of sage grouse within the Moffat County area and this investigation
was funded.
P. N. OBJECTIVES
To (1) ascertain the distribution of sagebrush in the Moffat County area,
(2) determine the status (i.e., principally alteration) of sagebrush in
this area, (3) delineate the distribution of sage grouse in the Moffat
County'area, and (4) identify the past and present trend of sage grouse
popUlation status in this area.

�-98-

DESCRIPTION

OF AREA

The Study Area
The study area (Fig. 1) consisted primarily of Moffat County in its entirety
but also included some inventory work in western Routt County from the
vicinity of Hayden to the Moffat County Line. Western Routt County was
included because some occupied sage grouse ranges and formerly occupied
ranges in western Routt County are contiguous with those in eastern Moffat
County.
Moffat County is located in the extreme northwestern corner of Colorado,
bordered by Wyoming on the north, Utah on the west, Rio Blanco County on the
south, and Routt County on the east. Moffat County was created in 1911 from
the western two-thirds of Routt County.
Main drainages include the Green River which flows from north to south through
western Moffat County, the Yampa River which flows from east to west in the
southern part of the County, and flows into the Green River approximately 3
miles (4.8 km) from the Utah border, and the Little Snake River which flows
from northeast to southwest through the middle of the County, and flows
into the Yampa River in Lily Park.
Climate
Moffat County climate is classified as semi-arid with 8 to 20 inches (20.350.5 cm) of precipitation annually (Table 1). Precipitation is mainly
concentrated in the late summer to early winter.
The mean annual precipitation normal for 6 stations was 12.99 inches (33.0 cm) and the annual mean
temperature for five stations was 43.4°F (6.30C) during years of record
(Table 1). The elevation of the County varies from approximately 4,600
to
11,045 feet (1,402-3,367 m) and the town of Craig is 6,240 ft (1,902 m) above
sea level.
Miscellaneous

Information

The County seat of Moffat County is Craig, located at the junction of Colorado
Highway 13 and U. S. Highway 40 in the eastern part of the County.
Agriculture,
for the most part, is dryland farming with a small percentage dependent upon
irrigation.
Agriculture has been considered the "staple" economy, with mining
and other energy-related activties becoming more and more significant in
recent years.
The first of 2 units of a coal fired electrical generating plant (Yampa Power
Plant) is presently being constructed south of Craig in Moffat County.
The
first unit is scheduled to begin operating in the fall of 1979, the second
unit in the spring of 1980, and a third unit is to be built later at the
same location.
Coal production from 3 operating mines, including 2 surface
mines (Colowyo and Trappers) and 1 underground mine (Wise Hill Number 5)
in Moffat County during 1977 was 1,080,677 tons. One additional surface
mine had no production and 1 additional underground mine was being prepared
for production during 1977. Coal production for Moffat County for 1976 was
507,010 tons according to records secured from the Colorado Division of ~ines.

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�-100-

Table 1. Annual precipitation normals and annual mean temperatures
stations within Moffat and western Routt counties.

Station

Elevation
(ft)

Annual
Precipitation
Normal
(in)

Brown's Park

a
5,600

8.07

Craig

6,240

13.78

Dinosaur

6,000a

11.77

Hamilton

6,235

Hayden
Maybell

Annual
Mean
Temperature

1975 (10 mo)

d

1941-1970

b

1975 (11 mo )

c

c

l7.22

1951-1960

6,337

16.11 d

1941-1970

6,000a

10.99

c

b

1974-1975
(15 mo)

Total

77.94

216.7

Average

12.99

43.3

a
b
c
d

Period of
Record

(OF)

b

c

for

Data from U. S. Geol. Survey, Topographic

Map (1954).

Data from Dept. of Atmospheric

Colo. State Univ.

Services,

Data from U. S. Dept. of Commerce

(1964) Weather

Summary

(1951-1960) .

Data from U. S. Dept. of Commerce

(1973) Weather

Summary

(1941-1970).

Historical

Human Population

Statistics

Table 2 presents the human population growth of Craig and Moffat County since
1910 (Moffat County Planning Department 1977). The increase in population
during the 1970's can be attributed mainly to industrial-energy
developments
brought about by a national energy shortage and large scale deposits of coal,
oil, natural gas, and oil shale within the Moffat County area.
BACKGROUND

INFORMATION

RANGE
Winter Range
Sage grouse winter ranges within a Montana study area were characterized by
large expanses of dense sagebrush on land having little if any slope (Eng and

�-101-

Schladweiler 1972).
Sage grouse tended to concentrate on winter ranges in
Central Montana, with from 200 to 300 birds observed.
Marked females showed
an affinity for winter areas approaching that shown for strutting grounds.
Removal of sagebrush from winter areas by control projects greatly reduces
their capacity to support wintering sage grouse (Eng and Sch1adweiler 1972).
In Idaho, winter concentrations of sage grouse were usually found where
snow was less than 6 inches (15.2 cm) deep (Dalke et a1. 1963).

Table 2.

Historical

population

growth of Craig and Moffat

1910-1977.

Moffat

City of Craig

Year

County

County

a

1910

392

1920

1,294

5,129

1930

1,418

4,861

1940

2,123

5,086

1950

3,080

5,946

1960

3,984

7,061

1970

4,205

6,525

1977b

6,657

10,221

a

Moffat
County.

County was created

b Preliminary

Nesting

in 1911 from the western

report from a Special U. S. Census

and Brood Rearing

two-thirds

taken in April

of Routt

1977.

Range

Walles tad and Pyrah (1974) equipped 31 sage grouse hens with radio transmitters and located 22 nests from these hens and 19 additional nests.
Sixtyeight percent of the 22 nests of radio-equipped hens occurred within 1.5 mi
(2.5 km) of the strutting ground where the hens were captured.
All nests
occurred in sagebrush stands with a canopy coverage that exceeded 15 percent.
A 3 km (1.9 mi) radius of an occupied 1ek (strutting ground) is considered'
to be the probable breeding-nesting
complex by the Western States Sage
Grouse Committee (Braun et ale 1977).
Gill (1965) found 87 percent of 23
nests within 2 mi (3.2 km) of a strutting ground in North Park, Colorado
studies.

�-102-

Klebenow (1969) considered shrub cover for sage grouse nesting as essential
in Idaho and stated that shrub control projects in nesting habitats must
be carefully considered since removal of shrubs can result only in the
elimination of sage grouse from the area. Wallestad and Pyrah (1974) stated
that the welfare of this unique bird should be considered in programs
designed to modify or eliminate sagebrush.
In a Montana study area, broods utilized sagebrush-grassland
benches early
in the summer (June and July) and moved to greasewood bottoms and/or alfalfa
fields as the forbs on the higher elevations became desiccated.
Broods
remained in these bottom types until late August and early September and
then moved back into sagebrush (Walles tad 1971).
Effects

of Range Alterations

(General)

The fact that farming and ranching activities in the arid west have both
damaged and improved sage grouse habitats have long been recognized
(Edminster 1954). Although sage grouse habitat may be improved with some
of its range devoted to farming and with people living and traveling nearby,
it is also true that there is a rather close limit of tolerance by sage
grouse to these conditions, basically because it is a wilderness species.
Intensive farming practices have taken over some areas of sage grouse range
and have excluded the species from all but the fringes (Edminster 1954).
Effects

of Spraying

Sagebrush

The effects of sagebrush control on the distribution and abundance of sage
grouse in North Park, Colorado were studied from 1963-1975 (Braun and Beck
1976).
Spraying of sagebrush within the study area resulted in decreases in
cover and relative frequency of sagebrush and forbs. Number of active
strutting grounds within the original study area declined from 7 to 6 from
1963 to 1974. Maximum numbers of male sage grouse counted on strutting
grounds within the original study area decreased significantly (P&lt;0.05)
during the period from 1959 to 1974. Block spraying of sagebrush eliminated
nesting by sage grouse and greatly reduced sage grouse use of treated areas.
While centers of sprayed strips were largely avoided by sage grouse, nesting
and other use of edges of sprayed strips did occur, apparently the result of
low percentage sagebrush kill.
INVENTORY
Strutting

Ground Counts, Moffat

County 1958-1977

Continuous counts of sage grouse on strutting grounds have been made since
1958. Table 3 lists numbers of active strutting grounds counted, numbers
of total birds and high male counts on strutting grounds in Moffat County
by Rogers (1964) and Wildlife Conservation Officers during the years of
record (1958-1977).
Apparent trends in total birds and highest numbers of males counted by Rogers
(1964) and Wildlife Conservation Officers in Moffat County from 1958 to 1977
are shown in Fig. 2. These data suggest possible sage grouse population

�-103-

Table 3. Summary of sage grouse strutting ground counts, Moffat County,
1958-1977 . a

Number
Counts

High
Count
Males

High
Count
Total
Birds

Average No.
Males/Ground

Average
Total No.
Birds/
Ground

30

58

588

679

19.6

22.6

1959

24

56

708

962

29.5

40.1

1960

16

35

569

569

35.6

35.6

1961

18

39

766

825

42.6

45.8

1962

66

1,010

1,190

1963

106

760

837

1964

89

564

672

1965

87

503

548

1966

71

571

581

1967

70

1,086

1,147

1968

71

1,449

1,666

Year

Number
Active
Grounds
Counted

1958

1969

29

69

2,164

2,283

74.6

78.7

1970

28

61

1,522

1,803

54.4

64.4

1971

25

57

1,299

1,374

52.0

55.0

1972

27

49

1,037

1,268

38.4

47.0

1973

22

1,008

1,123

45.8

51.0

1974

22

910

1,098

41.4

49.9

1975

17

539

663

31.7

39.0

1976

24

48

1,032

1,254

43.0

52.3.

1977

20

36

1,052

1,157

52.6

57.9

a Data from Rogers (1964) and Game Management Section.

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e

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2,000

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1,600

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Total Birds

• High Male Count
1966
YEA

1968

1970

1972

1974

1976

R

Fig. 2. Trends in numb~r of sage grouse counted on strutting grounds by Wildlife Cons~rvation Officers in Moffat
County, 1958-1977.

,

0
,

• I-'

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I

�-105-

fluctuations in a ten year cycle from 1965 to 1975 with a major peak in
populations occurring in 1969 and lows occurring in 1965 and 1975. Populations have apparently been on the upswing since the last low in 1975.
As expected, higher average numbers of both total birds and males were
tallied during years of peak populations (1961 and 1969) than during years
of lows (1958 and 1975) (Table 3). However, records indicate that numbers
of active strutting grounds counted annually by Rogers (1964) and W.C.O.'s
have varied considerably during the years of record (1958-1961 and 1969-1977,
Table 3). Because of this wide variation in number of active grounds counted,
the average number of total birds and males counted on strutting grounds are
probably better indicators of populations than total number of birds and
males.
Unfortunately, no record of the number of active grounds counted is
available for the years 1963 to 1968. This greatly reduced the value of
comparisons of trends using averages (Fig. 3).
Brood Counts,

1953 and 1958-1961

Testor and Boeker counted 9 roadside sage grouse trends in Moffat County in
August of 1953 (Rogers 1964). They drove 357 mi (574 km) and saw 461 birds
for an average of 1.26 birds per mi (0.78/km).
Sage grouse counted on brood
routes in the Northwest Region adapted from Rogers (1964) are summarized in
Table 4. Brood counts on sage grouse in Moffat County have been too sporadic
within recent years to be of value with only a very few exceptions.
Harvest

Statistics

1962-1977

Prior to 1968, sage grouse harvest statistics were obtained through mail
surveys only on a regional or statewide basis.
No breakdown was made by
county and Small Game Management Units were not established until 1968.
For these
reasons
comparisons of kill information from mail surveys can
be made for the Moffat County area only since 1968.
Small game Management Units established within Moffat County for reporting
harvest information from 1968-1973 are shown in Fig. 4(A) and from 1974-1977
in Fig. 4(B). Table 5 lists information on hunting seasons and bag limits
from 1962-1977, and Table 6 lists harvest information compiled from Small
Game Harvest Surveys from 1968-1977.
Fig. 5 shows apparent trends in number
of sage grouse harvested and number of hunters during the period 1968-1977.
An apparent peak in numbers of sage grouse harvested is found in 1969,
coinciding with peak numbers of sage grouse counted on strutting grounds
(Fig. 2). Another apparent peak in numbers of sage grouse harvested (1972)
may be partially explained by increased hunting pressure during a year with a
more liberal hunting season than usual during a year when populations were
still fairly high (Table 3, Fig. 2). Correlations between apparent population
levels and numbers of sage grouse reportedly harvested were, however, quite
low during this period of record (1968-1977).
Cedar Mountain

Check Station Data 1962-1973

Rogers (1964) lists sage grouse harvest data for the Cedar Mountain Check
Station, northwest of Craig, for the years 1953-1961 and for Blue Mountain
Check Station, near the town of Dinosaur, for the years 1955-1958.
Table 7
lists sage grouse harvest data gathered by management personnel of the Northwest Region at the Cedar Mountain Check Station for the period 1962-1973.

�100

•\

80

.\' .

~
tal

en

60

X

(No Record of Number of Grounds)

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tal

&gt;
20

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tal

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/

•
Average Number Total 3lrds Per Ground

--e

•

Average Number Males Per Ground

YEAR
Fig. 3. Trends in average number of sage grouse counted on strutting grounds by Wildlife Conservation Officers in
Moffat CQunty, 1958-1977.

I
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I

�-107-

Table 4.

Summary of sage grouse brood counts, Hoffat County,

1958-1961.

a

Year
Item

Areas counted

1958

1959

1960

1961

Hoffat and
W. Routt Co.

Hoffat and
W. Routt Co.

Hoffat Co.

Hoffat and
W. Routt Co.

Number

routes

6

5

3

5

Number

counts

14

26

11

19

273.9

749.9

325.2

558.8

Total miles
Total birds
Birds/mile
a

Data from Rogers

1,453

1,442
5.26

1.94

1,397

683
1. 22

4.30

(1964).

According to these data, an average of 165 (range 23 to 411) sage grouse
hunters were checked with an average of 211 (range 30 to 524) sage grouse
harvested.
Average number of sage grouse harvested per hunter has ranged
from a low of 0.82 in 1963 to a high of 2.19 in 1971 and hours required to
kill a sage grouse have ranged from a low of 0.90 in 1971 to a high of 6.57
in 1973.
METHODS

AND MATERIALS

Range
A search of literature was made of studies which described
or near the Moffat County area.

vegetation

within

Major areas of big sagebrush-grassland
range were mapped on 0.5-in per mi
County
highway maps utilizing range resource maps of the Soil Conservation
Service, traverse and sketch of range vegetation types, and sketches of
vegetation types from available aerial photographs and mosaics.
Records of locations, amounts, and types of range alterations by Federal
Agencies were secured from the Fort Collins Office of the Colorado Division
Wildlife.

of

A sample of 20 landowners within prime sage grouse ranges of Moffat County
was
interviewed to estimate percentages of native big sagebrush habitat
altered to bring land under cultivation or to improve pastures for grazing
purposes.
A sample of 10 landowners within the prime dryland cultivated areas
of western Routt and eastern Moffat counties was interviewed to estimate percentages of native big sagebrush habitat altered to bring land under cultivation,
primarily for the raising of wheat.
Records of all miscellaneous sage grouse observations (both during the summer
and winter periods) and locations and causes of mortalities were recorded to
outline possible ranges by period of use.

�-108-

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MOFFAT COUNTY

". ~

COLORADO

A.l'J66-197:l

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MOFFAT COUNTY
COLORADO

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....

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Fig. 4.

Designation of Small Game Management Units~ Moffat County.

�-109-

Table 5.

Sage grouse hunting

seasons, Moffat

.
a
County area 1962-1977.

Area or Small Game
Management Unitsb

Bag
Limit

Possession
Limit

Aug. 17-19

2

4

All

Aug. 17-19
Aug. 17-19;
Aug. 31-Sept.

2

4

All, except

3

6

Moffat County W. of county roads
9 and 17, and N. of U. S. Hwy.
40 and Colo. Hwy. 318

Sept. 12-14
Sept. 15-20

2
2

4
4

All
Mainly

Sept. 11
Sept. 12-19

2
2

2

4

All
W. Moffat County (mainly Blue
and Cold Spring Mountain areas)

Sept. 10-11
Sept. 12-18

2

4
6

All
Mainly

Sept. 9-10
Sept. 11-17

2

3

4
6

All
Moffat County W. of Colo. Hwy.
13, W. of County Roads 17 &amp; 57

1968

Sept. 14-16

3

6

5, 6 and 7

1969

Sept. 13-16

3

6

5, 6 and 7

1970

Sept. 12-21

3

6

5, 6 and 7

1971

Sept. 11-13
Sept. 11-19

2
2

4
4

5

Sept. 11-19

1

2

6 and 7 (E. of Colo. Hwy 318, S.
of Green River &amp; Vermillion Creek)
7 (remainder)

1972

Sept. 9-17

2

4

5, 6 and 7

1973

Sept. 8-10

2

4

5, 6 and 7

1974

Sept. 14-16
Sept. 14-20

2
2

2
2

14, 16 and 18
20

1975

Sept. 13-15

2

2

14, 16, 18 and 20

1976

Sept. 11-13

2

4

14, 16, 18 and 20

1977

Sept. 10-16

3

6

14, 16, 18 and 20

Year

Hunting Season
Dates

1962
1963

1964

1965

1966

1967

a Information
Regulations.

2

from Division

3

of Wildlife,

b Open areas by County and specific areas
Units (1968-1977, Revised in 1974).

for below

Blue Mountain

Blue Mountain

Small Game Surveys

(1962-1967);

area

area

and Small Game

Small Game Management

�8,000

o
o

7,000

6,000

5,000

Il::

r:z::I4,OOO

I
I-'
I-'

o

o
I

III

~
:&gt;

%_o/O~

~

o

3,000

j\
/

/
2,000

/0--

0 &lt;,

-

1,000

o

&lt;,

/

\

\

0

'0

-

Hunters

1969

1970

.". ,0

--

0-

--- Birds Harvested
1968

\

/

/

",

\

j

&lt;,

o

\

_

.-

-'0

'- '0"'-

Fig. 5. Trends in number of sage grouse harvested and number of hunters
for Small Game Management Units 6 and 7 (1968-1973) or 16, 18, and 20
(1974-1977) (from Small Game Harvest Surveys).

1972

1971
YEA

R

1973

1974

1975

1976

1977

�-111-

Table 6. Sage grouse harvest statistics
Highway 13, 1968-1977.a

for Moffat County west of Colorado

Small Game
Management Units

Number
Hunters

Total
Harvest

6
7

612
517
1,129

1,967
2 085
4,052

6
7

1,351
575
1,926

5,656
1,644
7,300

6
7

1,635
459
2,094

3,896
1 043
4,939

6
7

1,131
401
1,532

3,857
1,193
5,050

6
7

2,470
456
2,926

6,528
1,294
7,822

6
7

988
219
1,207

2,153
328
2,481

16
18
20

955
291
125
1,371

1,807
770
341
2,918

16
18
20

776
51
184
1,011

2,046
256
767
3,069

16
18
20

981
208
179
1,368

2,752
428
406
3,586

16
18
20

678
85
212
975

1,486
233
738
2,457

Totals

15,539

43,674

Average

1,554

4,367

Year

1968
Subtotal
1969
Subtotal
1970
Subtotal
1971
Subtotal
1972
Subtotal
1973
Subtotal
b

1974

Subtotal

1975

Subtotal
1976

Subtotal
1977

Subtotal

a Data from Div. of Wildl., Wildl. Mgmt. Section
information) •
b
Small Game Management Units revised in 1974.

(published and unpublished

�Table 7.

Sage grouse harvest data from Cedar Mountain Check Station, 1962-1973.a

Year
Item

1962

1963

1964

Total number hunters

411

377

23

Total hours hunted

1,331

1,364

59

0.82

30
1.30

4.40

1. 97

Total birds bagged
Average birds/hunter
Hours hunted/bird

524
1.27
2.54

310

1965

1966

1967

28

87

166

109

136

192

95

152

209

85

213

291

449

291

490

187

560

1,241

47

83

289

210

236

141

189

0.95

1. 74

1. 93

1. 74

270
1.41

208

1. 68

2.19

0.93

0.90

1.81

2.57

1.01

2.14

1. 23

1.81

0.90

3.97

6.57

1968

1969

1970

1971

1972

1973

Total cocks

92

44

9

1

35

69

69

74

71

45

15

27

Total hens

189

109

21

46

48

110

71

87

95

89

21

36

Total old birds

144

82

15

3

49

92

62

87

87

95

23

49

Total old cocks

35

16

o

1

19

23

20

45

28

27

13

22

Total old hens

109

66

15

2

30

69

42

42

59

68

10

27

Total young birds

137

71

15

44

34

87

78

74

79

39

13

14

Total young cocks

57

28

9

o

16

46

49

29

43

18

2

5

Total young hens

80

43

6

44

18

41

29

45

36

21

11

9

Percent cocks

32.7

28.8

30.0

2.1

42.2

38.5

49.3

46.0

42.8

33.6

41.7

42.9

Percent old cocks

12.5

10.5

0.0

2.1

22.9

12.8

14.3

28.0

16.9

20.1

36.1

34.9

Percent young cocks

20.3

0.0

19.3

18.0

25.9

13.4

5.6

7.9

70.0

97.9

57.8

25.7
61.5

35.0

67.3

18.3
71.2

30.0

Percent hens

50.7

54.0

57.2

66.4

58.3

57.1

Percent old hens

38.8

43.1

50.0

4.3

30.0

26.1

27.8

42.9

28.1

20.0

93.6

22.9

20.7

28.0

35.5
21.7

50.7

28.5

36.1
21.7

38.5

Percent young hens

15.7

30.6

14.3

Percent young birds

48.8
51.2

46.4

50.0

93.6

41.0

55.7

46.0

47.6

29.1

36.1

22.2

53.6

50.0

6.4

59.0

48.6
51. 4

44.3

54.0

52.4

70.9

63.9

77.8

Percent old birds
a

I
f-'
f-'
N

I

Data from Division of Hi1d1ife Northwest Regional Office.

�-113-

INVENTORY
Strutting

Ground Counts

Sage grouse were counted on all known active strutting grounds in Moffat
County during the Spring of 1976. Attempts were made to locate and count
as many old and new grounds as possible rather than attempting replicate
counts on fewer grounds. A crew of 3 men using 3 four-wheel drive vehicles
searched various areas in Moffat and western Routt counties from mid-April
through May. Procedures for locating strutting grounds through ground
searches and methods for securing counts were similar to those outlined by
Rogers (1964).
Previous to actual strutting ground counts in 1977, considerable effort was
made to locate wintering flocks of sage grouse through routine ground
searches.
The mild open winter of 1976-77 proved valuable in this activity.
Seven new strutting grounds were located in vicinities where wintering flocks
of sage grouse (especially males) had been observed periodically, starting
as early as January, 1977. Three early morning helicopter searches, using
nine and one-half hours of flying time were made from March 22-24, 1977,
resulting in locating 7 new grounds, relocating 2 grounds, and counting
birds on 3 active grounds.
Also in 1977, a crew of 3 men using 3 four-wheel drive vehicles counted all
known active strutting grounds and searched for additional new grounds
during the period April I-May 31.
Brood Counts
Sage grouse were counted along 10 routes in Moffat and western Routt counties
in August 1976 using procedures similar to those outlined by Rogers (1964).
All routes were run in the early morning commencing as soon as it became
light enough to see well.
Based upon encouraging results obtained in counting
sage grouse along these ten brood routes, the number of routes counted in
1977 was expanded to 11 and both early morning and late evening counts were
made. Morning routes were again usually started as soon as it became light
enough to see well and attempts were made to end the late evening routes
about dark. Routes were run going away from the sun whenever possible for
improved visibility.
Distribution

Checks

Observations of sage grouse, excluding birds observed during strutting
ground counts and brood route counts, were recorded by date, time, location,
number of birds, and major vegetation cover type from January 1, 1976 through
March 31, 1978. Locations and numbers of birds were plotted on Moffat County
maps on a scale of one-fourth inch per mile.
Separate maps were used to
plot summer period (April I-November 30) and winter period (December I-March
31) observations.
Different symbols were used to separate observations on
the maps by year.
Harvest Data Collection.--Sage grouse wings were collected during the
1976 hunting season through the use of 16 wing barrel collection stations and

�-114-

miscellaneous hunter contacts.
Locations of wing barrel collection stations
were similar to that of 1977 (Figs. 15 and 16) except in 1976 wing barrels
were placed near U. S. Highway 40 along the Wolf Creek Road, along the Sand
Wash Ridge Road, South of Powder Wash Camp, and near the Wyoming State Line
along Moffat County Road 2. With a 3 day season in 1976, collections were
made at the end of the second day and following the close of the season.
The wings were then frozen and stored until being processed.
Sage grouse wings were again obtained from Moffat and western Routt counties
during 1977 through use of 16 wing barrel collection stations placed within
these areas, a grouse research check station operated in Middle Park on
opening weekend, and wing survey envelopes distributed by the Wildlife Management section.
Locations of 4 wing barrel collection stations were changed for 1977 because
of the low number of sage grouse wings collected at them during 1976. The
other 12 were placed in sites similar to 1976. With a longer season in 1977
(seven days) collections were made at the end of the second day, end of the
third day, midweek, and following the close of the season.
These were again
stored in a freezer until being processed.
RESULTS AND DISCUSSION
Range
General Vegetation

Types

Sage grouse are commonly found in 2 main types of ranges: sagebrush-grassland
pra~r~e type and salt-desert shrub type. Generally these types are separate
but may also be intermixed (Edminster 1954).
Harrington (1954) described the semi-desert vegetation zone in northwestern
Colorado, of which the sagebrush plant community is an integral part, as
supporting dense stands of big sagebrush with varying mixtures of herbaceous
plants in the understory.
He states the sagebrush type is best developed
below 7,000 ft (2,134 m) in middle western Colorado, but may extend to more
than 10,000 ft (3,048 m) in a few locations.
At its lower border the type
merges with saltbrush (Atriplex spp.) and greasewood (Sarcobatus vermiculatus)
plant communities.
At its upper border, the boundary between sagebrush and
mountain shrub types is usually clear cut, but its contact with pinon pine
(Pinus edulis)-juniper
(Juniperus spp.) is frequently diffusive.
Vegetation within the big sagebrush-grassland
plant community in Moffat and
western Routt counties is similar to that described for sage grouse ranges
within the Piceance Basin (Krager 1977). Vegetation is predominantly big
sagebrush with scattered serviceberry (Amelanchier spp.) interspersed with
open areas. Moffat County also contains large areas of saltbrush-greasewood plant communities within its sage grouse ranges, particularly in its
western parts.
Within the Moffat County area (Moffat and western Routt counties), big sagebrush-grassland ranges are either bordered by or interspersed with saltbrushgreasewoodpinon
pine-juniper, rough and rocky, mountain shrub, aspen (Populus
tremuloides), or spruce (Picea spp.)-Douglas fir (Pseudotsuga taxifolia) types.

�-115-

The mountain shrub type normally
chokecherry (Prunus virginiana),
Specific

Vegetation

contains serviceberry, oak (Quercus spp.),
and mountain mahogany (Cercocarpus montanus).

Descriptions

A literature search to locate information concerning specific vegetation
descriptions for the Moffat County area was made unsuccessfully
in libraries
of Colorado State University and the Colorado Division of Wildlife.
Land Ownership

Status, Moffat

County 1976

Table 8 lists amounts of land in Moffat County by ownership as determined
from information secured from various Federal and State agencies.
More than
one-half of the total land area of 3,035,530 acres (1,228,479 ha) in Moffat
County is administered by Federal agencies (51.3%), compared with 42.1
percent in private ownership and 6.6 percent in State ownership.

Table 8.

Land ownership

in Moffat

County 1976.

Ownership

Acres

% of Total

U. S. Government
Bureau Land Management

(BLM)

1,354,552

44.62

National

(NPS)

149,444

4.92

41,763

1.38

12,380

0.41

1,558,139

51. 33

191,525

6.31

9,120

0.30

Subtotal

200,645

6.61

Private Land

1,276,746

42.06

Totals

3,035,530

100.0

Forest

Park Service

Service

(FS)

Bureau Sport Fisheries
(BSF &amp; W)

and Wildlife

Subtotal
State of Colorado
State Land Board
Division

(State School Lands)

of Wildlife

(DOW)

�-116-

Range Alterations

on Federal Lands in Moffat County

Locations of all known range conversions of Federal Lands in Moffat County
are shown in Fig. 6. Table 9 summarizes range conversions on Federal lands
by type of habitat manipulation and acreages.
The largest amount of range
conversions on Federal lands in Moffat County has been through spraying
(39.2%), followed by plowing and seeding (25.0%), chaining (19.5%),
reseeding (14.6%), burning and reseeding (1.2%), and rotobeating (0.5%).

Table 9. Summary
County. a

of habitat manipulation

projects

Type of
Manipulation

Map
Symbol

on Federal

lands, Moffat

Number
Acres

Percent
of Total

(A)

Seed (not fire reseedings)

4,597

9.5

(A)

Fire reseedings

2,455

5.1

Subtotals

7,052

14.6

(B)

Spray

18,919

39.2

(C)

Chain

2,607

5.4

(C-M)

Chain, plow and seed

1,500

3.1

(L)

Chain or rail and seed

5,302

11.0

Subtotal

9,409

19.5

(H)

Rotobeat

240

0.5

(M)

Plow and seed

12,056

25.0

(N)

Burn and seed

570

1.2

48,246

100.0

Totals

a Includes all projects
(FS) lands.

on Bureau Land Management

(BLM) and Forest

Service

It was determined that approximately 3.1 percent of the Federal lands in
Moffat County has been altered intentionally through various habitat
manipulations.
Except for one small area of 60 acres (24.3 ha) on the Routt
National Forest, all intentional habitat manipulations on Federal lands in
Moffat County have been on BLM lands.

�:..
-

',,",

-

1,.-1'°-

1"'1[

:,..-

I~

I=~

....

.•...

.Jill

(1\:

II)i.
410

•

~

..".

!~

,"

1---"

(i"

L.-:-:.,&gt;1!l.s::J.,&lt;
L~_E-

~"
t-

~,

f-"

1'-

t- ~
It(;

VT

IJlS

I"&gt;

T-

&gt;:;.~

~~:

L:

,~
t..

t-J ~

; ; .... .;1

-C- j\

•
~

-117--

I\(

:

�-118-

Range Alterations

on Privately

Owned or Controlled

Lands

Within Prime Sage Grouse Ranges, Moffat County
Twenty private landowners within prime sage grouse ranges in Moffat County
were interviewed to determine the past and future extent of habitat alterations on native sage grouse ranges on private lands. Landowners interviewed
owned and/or controlled the grazing rights on lands averaging 26,671 acres
(10,794 ha) (range 320 to 139,520 acres or 130 to 56,464 ha) amounting to
10.3 percent of the total deeded land within Moffat County.
Table 10
summarizes information gathered in this survey.
It was determined that approximately 25.2 percent of the deeded land has been
altered in the past in order to bring land under cultivation or to improve
pastures for grazing purposes.
These individuals plan to alter an additional
1.7 percent of their land within the next ten year period, if economic
conditions are favorable to do so.
Table 11 lists amounts and percentages of croplands on 16,535 acres (6,692
ha) owned or leased by the 20 landowners interviewed within Moffat County
during 1976. By use, 79.4 percent of the croplands were used to grow dryland small grains (wheat or barley), 13.8 percent to grow dryland hay, and
6.8 percent to grow irrigated hay.
Amounts of Big Sagebrush Sprayed Area
in Two Important Brood Rearing Ranges
Better cattle ranges such as found on Cold Spring Mountain and Blue Mountain
contain extensive big sagebrush habitat where range managers have improved
large areas of pastures for livestock grazing purposes through spray projects.
These better cattle ranges are also preferred by sage grouse for summer and
brood rearing ranges, based upon brood count information gathered in this
investigation.
Cold Spring Mountain.--Within
56~5 mi2 (14,634 ha) of high quality big
sagebrush habitat on Cold Spring Mountain, extensive field surveys indicated
that 7.1 mi2 (1,839 ha) (12.6%) have been sprayed.
Of this sprayed area, 1.4
mi2 (363 ha) were sprayed by the BLM in 1968 and the remainder have been
spray projects on private or state school lands.
Within 29.9 mi2 (7,744 ha) of fair quality big sagebrush range on Cold Spring
Mountain surveys indicated that 10.3 mi2 (2,668 ha) (34.4%) have been sprayed.
Of this area, 3.1 mi2 (803 ha) were sprayed by the BLM in 1966 and the
remainder have been spray projects on private or state school lands.
In combining both high and fair quality big sagebrush range on Cold Spring
Mountain, it was determined that approximately 20.1 percent of the big
sagebrush range has been sprayed.
Blue Mountain.--Within
a 4 township area (144 mi2 or 37,297 ha) on Blue
Mountain, it was determined there were 81.5 mi2 (21,109 ha) of hi~h quality
big sagebrush habitat.
Extensive field surveys indicated 21.6 mi
(5,595 ha)
(26.5%) of this big sagebrush range have been sprayed (Fig. 5). Of this
sprayed area, 3.2 mi2 (829 ha) were sprayed by the BLM in two separate
projects during 1965 and 1966 and the remainder have been spray projects
primarily on private lands.

�Tcbl.

Ic .

sUffillIary af habitat

manipul.ations

within

prime

sage

grouse

Number Acres Controlled
Pvt.
Stat.
-'l~.e~.'_'s'_"E'___ Sch. Landa_

range,

Moffat

County.a

Landowner'
Code

of
Oporntion

1-76

Sheep &amp;
Cattla
Ranching

1,969

920

2-76

Sheep &amp;
Cattle
Ranching

16,000

3,000

3~76

Cattle
Ranching

11, 000

4-76

Cattle
Ranching

980

5-76

Sheep &amp;
Cattle
Ranching

340

340

6-76

Landowner
Only

520

520

7_76

Cattle
&amp;
She.p
Ranching

37,000

B-76

Cattle
Ranching

r.soo

9-76

Cattle
Ranching

17,,500

__________________

Typ&amp;

Doedod

J ppr ox ,

BLM

NY~-

150

Method

hcres
(Pvt.)

No.

A.l.terad

%

Appr-ox ;

Acres

Method

No.

"

Plennod

rr.z

640

3,529

1,365
(Deeded &amp;
Pvt. Lease)

Burn I plow,
seed

3,000

24,000

46,000

15
( Deeded)

Plow,
seed

1,920

29,080

11 2,000

unknown

Plow &amp; seed
spray

320
~Deeded &amp;
Pvt. Lease)
300
(Deeded)

DisK or brushcutter
&amp; seed

9.0

100

Brushcutter

2.8

Brushcu t ter
seed

8B.2

20

Plow

5.9

3,540

Z,560

2,000

Total

47.2

&amp;

harrow,

&amp;

!tail,
seed

plow,

&amp;

0.1

0.0

or

&amp;

0.0

seed

&amp;

Unknown

0.0

2,500

12,000

53,500

8,000
(Deeded &amp;
Pvt. Lease)

Brushcutter,
cultivate
&amp;.. seed

20.5

160

1,280

2,940

90
(Deeded)

Disk or
(horses)

plow
&amp; seed

6.0

2,500

59,840

90,150

4,000
(Deeded)
3,400

Aerial

sprayed

22.9

seed

19.4

10,160

325
( Deeded)

Plow

&amp;

500

Brushcutter,
cultivate
&amp; seed

1.3

0.0
Unknown

Aerial

160

Disc,
seed

spray

(ueeded)

10-76

800

Cattle
Ranching
&amp; Dryland"
Parainq

4 r200

800

350
( Ueeded)

&amp;

4,520
(Deeded &amp;
Pvt. ueaee )

Spec. tiller,
aeed or aerial
spray or burn

320

240
(Deeded)

Hand

teams-

75.0

0.0

3,76C

3,560
(Deeded &amp;
Pvt. Lease)

Brushcutter,
plow, &amp; seed

94.7

0.0

1,980

900
( Deeded)

Hand

45.5

138,240

139,520

260
(Deeded)

Brushcutter,
burn,. plow,
seed

1,920

5,231

t,880
(O.eded "Pvt. Lease)

Brushcutter
or
disk &amp; burn,.
plow &amp; seed

56.B

Plow or brushcutter

25.0

240

Sheep, Cattle &amp; Dry
land farCll1ng

12-76

Cattla
Ranching
&amp; Dryland
Farlling

320

rl_76

Dryland
farming

720

14-76

Cattle
Ranching
&amp; Dryland
Farming

lr980

15-76

She.p &amp;
Cattle
Ranching

1,280

16-76

Cattle
Ranching
&amp; Dryland
farming

1,531

1,780

17-76

Cattle
Ranching
&amp; ory1ond
Farming

320

320

18-76

Sheep
Ranching

12,000

19-76

Cattle
Ranching

5,000

2,000

20-76

Cattle
Ranching

160

50

Total.

US .121

16.380

5,480

960

320

11-76

l,840

3,040"

640

160
(Deeded)

I-nforaat1on

from landowner

200
(Deeded)

Te8Jf'l &amp; burn

1,280

9,000

17,280

600
(Oaeded)

Brushcutter
seed

9,410

50
(Deeded)

Team,
plow

4,080

5,120

24 •.160

362.4BO

15,280

533,420

30,210
out of 119,980
(Unk.-out)

12.6%

26.111'

10.2"

17.6"

25.2%

••cur •••in

1976-1977 •.

p1m1

burn

33.7

plow

15.4

&amp;

0.0

100.0

200

10.1

Brushcutter
,.
plow, &amp; seed

0.0

20.3
&amp;

&amp;

&amp;

i
i-'
i-'

teams

105,440

10.1'%

intarvlcwa

&amp;

BO,640

at Total. Sampled
o

&amp;

12,800

Ill.500
Pctrc:entllg..

Disc,
burn,..
seed

BO

2.4

Brushcutter
r
plow &amp; seed

0.0

1.7

o

8.6

600

23.8

70

2,055
out of 119.980
(Unk..-out)
1.7"l.

0."
Brushcutte-r
seed
Disk

&amp;

seed

&amp;

8.6

33.3

\0
I

�-120-

Table 11. Summary of cropland use within prime sage grouse range, Moffat
County 1976. a

Landowner
Code

Dry1and
Hay

Acres of CroE1ands by TYEe
Dry1and
Wheat or Barley

Irrigated
Hay

1-76

0

300

65

2-76

0

0

0

3-76

400

0

100

4-76

80

60

5-76

0

0

100

6-76

0

0

0

7-76

1,000

5,000

0

8-76

0

0

90

9-76

0

0

200

10-76

80

270

0

11-76

0

1,560

0

12-76

40

200

0

13-76

15

3,545

0

14-76

400

500

0

15-76

0

0

260

16-76

200

1,600

80

17-76

70

90

0

18-76

0

0

200

19-76

0

0

0

20-76

0

0

30

Totals

2,285

13,125

1,125

Percentages
a

13.8

79.4

0

6.8

Basis--A sample of 20 landowners (same individuals listed in Table 10).

�-121-

Range Alterations Within Prime Wheat
Growing Areas, Moffat and Western Routt Counties
During the early phases of mapping big sagebrush ranges within the Moffat
County area, it became apparent that, up to a point, man-made range alterations could benefit sage grouse through creating openings and edge effect.
Sage grouse strutting grounds are quite often located in cultivated fields,
at salting locations, and in irrigated meadows.
It also became apparent
that extensive man-made range alterations, such as have occured through the
years within prime wheat growing areas (vicinities of Hayden in Routt County
and Craig in Moffat County) have eliminated former high quality sage grouse
ranges.
Ten wheat farmers within prime wheat growing areas in Moffat County, vicinity
of Craig, and Routt County, vicinity of Hayden (Figs. 7 and 8) were interviewed to determine the extent of past habitat alteration on formerly
occupied sage grouse ranges.
Table 12 summarizes information gathered in
this survey.
Landowners interviewed owned or controlled 27,395 acres (11,087 ha) (range
1,140 to 6,700 acres or 461 to 2,711 ha) amounting to 12.7 percent of the
total land outlined in Figs. 7 and 8.
It was determined that 70.5 percent of the private lands and 54.9 percent of
the state school lands within the sample had been converted to dryland
cultivated fields, plus an additional 3.2 percent of the private lands had
been converted to improved pastures or irrigated fields.
Thus, approximately
74 percent of the land within the sample had been altered for agriculture
purposes.
An average size farm was determined to consist of 1,333 deeded acres (539 ha)
and 2,740 total acres (1,109 ha). Of the ten landowners interviewed, 4 were
strictly wheat farmers, 5 combined wheat farming with cattle ranching, and 1
combined wheat farming with sheep ranching.
Range Reduction Within Three
Selected Prima Agricultural Areas
Twentymile
Area, Routt County.--Big sagebrush ranges within the Twentymile area southeast of Hayden were mapped from aerial photographs for the
years 1954 and 1969 and from field surveys (1977) on an area of 3~ x 3~ mi
(11.4 mi2 or 2,953 ha). Big sagebrush range reduction and land ownership
within this area are shown in Fig. 9 (A-D). Big sagebrush range in 1977
comprised less than 30 percent of the historical range and a strutting
ground comprised of 3 sites listed by Rogers (1964) has been abandoned.
The majority of the range reduction has occurred since 1954 and has been due
mainly to dryland cultivation activity, plus some coal mining activity.
Twentymile area contained the highest concentration of sage grouse in Routt
County during the early 1960's (Rogers 1964), but only remnant populations
of sage grouse now exist. Wildlife Conservation Officer J. Hicks found a
group of 12 males using transient strutting grounds one and one-half mi
(2.4 km) and one-half mile (0.8 km) east of the original sites during 1977
and 1978, respectively.
This range is classified as former range for all
practical purposes.

�-122-

Fig. 7.

Area of former sage grouse range, vicinity of Craig, Moffat County.

�-123-

~~

~'!Buck M~~.

l

. ..

"

Fig. 8.
County.

Area of former sage grouse range, vicinity

r

of Hayden, Western Routt

�Table 12.

Summary of habitat manipulations within prime wheat growing areas, Moffat and western Routt counties.
Number Acres Controlled
Private School
Total
BLM
Lands
Leased

a

Number Acres and Percentages Altered
School
Lands Percent
Purpose
Percent

County

Landowner
Code

Type of Operation

Routt
Routt

1-77
2-77

VJbeat farming
VJbeat farming, cattle ranching

320

-

829

-

1,149

305

95.3

Dryland cult.

1,100

1,000

-

-

2,100

1,200
60
1,260

57.1
2.9
60.0

Dryland cult.
Irrigated cult.

3,800

-

5,100

900

69.2

Dryland cult.

-

-

2,058

1,838

Dryland cult.

-

1,400

1,000

89.3
71.4

-

500

6,700

4,500

72.6

80

3,410

2,830
180
200
3,210

Deeded

Subtotals
Routt

3-77

VJbeat farming, sheep ranching

1,300

-

Routt

4-77

VJbeat farming

2,058

-

Routt

5-77

VJbeat farming

1,400

-

Moffat

6-77

VJbeat farming, cattle ranching

3,500

2,700

Moffat

7-77

VJbeat farming

1,330

2,000

Subtotals
Moffat

8-77

VJbeat farming, cattle ranching

980

-

160

-

1,140

Subtotals
Moffat
Moffat

9-77
10-77

VJbeat farming, cattle ranching
VJbeat farming, cattle ranching

680
658

2,000
1,000

-

-

2,680
1,658

Subtotals
Subtotals

13,326

Totals
Average number acres deeded land per farm = 1,333
Average number acres controlled per farm

= 2,740

--a Information from landowner interviews, 1977

8,700

4,789

580 27,395

Private
Lands

Purpose

629

75.9

Dryland cult.

2,000

52.6

Dryland cult.

Dryland cult.

-

85.0
5.4
6.0
96.4

Dryland cult.
Irrigated cult.
Improved pasture

-

-

-

750
195
945

76.5
20.0
96.5

Dryland cult.
Improved pasture

1,200

44.8

Dryland cult.

1,000
75
1,075

60.3
4.5
64.8

Dryland cult.
Improved pasture

15,523
240
470

70.5
1.1
2.1

16,233

73.7

Dryland cult.

0

0.0

-

-

Dryland cult.
Irrigated cult.
Improved pasture

2,629

54.9

-

2,629

54.9

Dryland cult.

I
•...
.,..
N

I

�-125-

N

A. Historical

B. 1954

D.

Land ownership, 1977

Key: (A, B, &amp; C)
_

~
(D)

Big sagebrush-grassland
Non-'z:ange'
and altered land
~
strutting
ground
0 Private land
lID Energy Fuels Corp.
'JiIlP Nature Conservancy Inc.

C. 1977
Fig. 9. Reduction of big sagebrush range by periods and land ownership
within Twentymile Area, Routt County.

�-126-

Twin ~esa~utt
CouniY.--Big sagebrush ranges within the Twin Mesa
area, southwest of Hayden, were mapped from aerial photographs for the years
1953 and 1969 and from field surveys (1977) on an area of 3~ x 3~ mi (11.4 mi2
or 2,953 ha). Big sagebrush range reduction and land ownership within this
area are shown in Fig. 10 (A-D). Big sagebrush range in 1977 comprised less
than 25 percent of the historical range and a strutting ground listed by
Rogers (1964) has been abandoned.
The majority of the big sagebrush range
reduction occurred prior to 1953, from agricultural activities (dryland
farming) dating back to the homestead period following World War I.
Twin Mesa contained only a light population of sage grouse during the early
1960's (Rogers 1964), probably because range reduction for agricultural
purposes (wheat farming) had largely occurred prior to 1953. Only remnant
sage grouse habitat and sage grouse populations now exist in this area.
This range is classified as former range for all practical purposes.
Axial Basin, Moffat County. ---Big sagebrush ranges within the Axial
Basin area, southwest of Criag, were mapped from aerial photographs for the
years 1953 and 1973 and from field surveys (1977) on an area of 3~ x 3~ mi
(11.4 mi2 or 2,953 ha).
Big sagebrush range reduction and land ownership within this area is shown in
Fig. 11 (A-D). Loss in big sagebrush range by 1977 had reached approximately
50 percent, but the potential for future extensive range losses are high.
Past range loss has been mainly due to agricultural development (dryland
and irrigated cultivation and improving pastures for grazing).
A strutting
ground recorded by Rogers (1964) was still active during 1977 with a high
cock count of 58. In 1959, W.C.O. William Roland counted highest cock
numbers of 31 (Rogers 1964), indicating the 1977 population may be even
higher than in 1959. The location of this ground has apparently moved to
the southwest approximately one-fourth to one-half mile sometime during the
period 1962 to 1976. Rogers (1964) considered the population of sage grouse
in the Axial Basin area to be light.
The amount of land now owned by ColoWyo Coal Company (Fig. 11 D) plus
increased coal mining activity with the opening of the large ColoWyo Strip
Mine nearby indicates probable future range reductions through coal mining
activities.
Extensive blocks of BLM land adjoining this range both to the
north and west may, however, allow for this population's continuing existence.
Classes

of Sage Grouse Ranges, Moffat and Western Routt Counties

Classes of sage grouse ranges within Moffat County are shown in Fig. 12 and
within western Routt County in Fig. 13, based upon surveys 1976-1978.
Descriptions

of Sage Grouse Range Classes

Class 1
Habitat

- Good to excellent quality habitat for sage grouse consists mainly
of dense high quality big sagebrush-grassland
(Fig. 14) with
overall altered habitats for agriculture and grazing consisting
of less than 26 percent by 1978. Areas of sparse, low sagebrush
and intermediate densities and heights may be present, but are
usually much more limited than are found in Class 2 and Class 3
ranges.

�-127-

N

A. Historical

•

B. 1953

D.

Land ownership,

Key: (A,

1977

B, &amp; C)

Big sagebrush-grassland

(D)

~.

type

~

strutting ground

II//I/A

BLM

Non-range and alte

c::J Private land

C. 1969
Fig.10• Reduction of-big sagebrush range by periods and land ownership
within Twin Mesa Area, Routt County.

red land

�-128-

•.
(-------3%
A.

Historical

rot

=~==-=~--)
.•
B.

1953

.D~
•

;

I

j

;

r

-

I

L

D.

Land ownership,

Key:
'"j

1977

(A, B, &amp; C}
Big sagebrush-grassland
Non-range and altered land

strutting
Private
Col~

ground
land
Coal Co.

BLM

c. 1977
Fie:011.
Reduction of big sagebrush range by periods and land ownership within
Axial Basin Area, Moffat County.

�""""

Z

-129~
z

;:)
0

0
....

~
IL.

0

::i!

~J

,;-

N

0

Jj j

~J •~

•. ~I~~
.~

~

~

i

::

~

E

"l
i!:

::

1m!. ~

•

fl ]

i

:

N

i J1

~
~

t

i10
t

'.l

:

~
i

0

�-130-

""""z

�I

r-'
W
r-'
I

Fig. 14. Class 1 big sagebrush range (foreground) bordered by pinon-juniper type and rough-rocky areas,
Moffat County.

�-132-

Populations

- Numbers of observations (Table 13) largely reflect amount of
time spent within Class 1 ranges, but are included to indicate
rough comparisons of populations.
Of 3,455 sage grouse observed
in miscellaneous checks during the winter period, 3,212 were
observed within Class 1 ranges.
Of 253 birds and seven broods
observed in miscellaneous checks during the summer period (excludes
strutting ground and brood counts), all of the 253 birds and six
broods were observed within Class 1 ranges.
Most (53 of 60 or
88.4%) of the known active strutting grounds were located within
Class 1 ranges.

Importance

- Survival and welfare of sage grouse populations is thought to
depend directly upon amounts of Class 1 ranges.
These contain
habitats used by sage grouse for summer, winter, and/or yearlong
ranges.

Extent of Area - Moffat County, 1,887 mi2 (488,748 ha)
Western Routt County, 111 mi2 (28,750 ha)
Class 2
Habitat

- Fair quality habitat for sage grouse on a yearlong basis, but these
habitats may become very important for survival during winters of
heavy snowfall within primary (Class 1) ranges. Habitat consists
mainly of intermediate height and density big sagebrush, but also may
include some open areas and intergrade with saltbrush greasewood vegetative types. Areas are normally drier than Class 1 ranges, affording
good and excellent habitat for sage grouse during winter periods.

Populations

- The relatively small number of sage grouse observed in Class 2
ranges (243 birds in the winter periods and one brood in the summer
periods) largely reflects upon the disproportionate
amount of time
spent within Class 2 ranges in comparison with Class 1 ranges (Table
13). Limited winter period observations and amounts of field signs
observed, however, indicate that Class 2 ranges become increasingly
important to sage grouse as the winters become more and more severe.
Five active strutting grounds out of 60 (8.3%) were located in
Class 2 ranges.

Importance

- These areas contain habitats used by sage grouse primarily
during winter periods and occasionally during brood rearing periods.
Overwinter survival of the species may depend upon Class 2 ranges
during severe winters.

Extent of Area - Moffat County, 520 mi2 (134,684 ha)
Western Routt Count, none
Class 3
Habitat

- Poor quality habitat for sage grouse on yearlong basis consists
mainly of lower density and height sagebrush than found in Class 2
ranges.
Open areas and saltbrush-greasewood
type become more
prominent than in better ranges.
Class 3 ranges are normally even
drier than Class 2 ranges, so may afford wintering habitats under
the most severe winter weather conditions.

�Table 13.

a
Miscellaneous sage grouse observations by range classes, Moffat County 1976-1978.
Classes of Range
1

b

3

2

Period

Birds

Broods

Mortalities

Birds

Broods

Mortalities

Birds

Broods

Mortalities

Birds

Summer

253

6

15

0

1

0

0

0

0

0

Winter

3,212

8

243

o

1

o

12

o

1

o

12

Totals

3,465

6

23

243

1

o

4c
Broods Mortalities

0

0

o
o

o

a Includes all miscellaneous observations from Figs. 19 and 21. Does not include strutting ground or brood counts.
b Refer to Fig. 12.
c

Non-range and former range.

I
t-'

W
W
I

�-134-

Populations

- Only one hen was observed within Class 3 ranges during the
study period (Table 13), but comparatively little field time was
spent within these ranges.
One small strutting ground (East Gee
Flats) was located and counted by W.C.O. Roger Lowry during the
early 1970's.

Importance

- These areas provide travel lanes between better ranges and may
provide wintering habitats under very severe winter weather conditions in Class 1 and 2 ranges.

Extent of Area - Moffat County, 434 mi2 (112,409 ha)
Western Routt County, none
Class 4 (Non-range
Habitat

and Former Range)

Habitat within areas classified as non-ranges consist of pinonjuniper, rough
and rocky, mountain shrub, aspen, and spruce-fir,
even though sage grouse may utilize fringes of most of these types.
Habitat within areas classified as former ranges (Figs. 6 and 7)
are primarily dryland cultivated fields interspersed with remnant
stands of high quality big sagebrush.
Overall altered native range
for agriculture and grazing was approximately 74 percent by 1978.

Populations

- Only 12 birds were observed within Class 4 range (Table 13),
but remnant populations are known to exist in a few locations.
Two active strutting grounds (29 Road 1 and 2) out of a total of
60 (3.3%) were located within Class 4 ranges.
These are located
in an area where approximately 45 percent of the native range
(largely big sagebrush-grassland)
exists, compared with an overall
average of only 2 percent for most of the area. Three abandoned
strutting grounds (Twentymile, Twin Mesa, and Fortirication Creek)
were located within Class 4 former ranges.
County, Non-range 1,902 mi2 (492,633 ha)
Former range 168 mi2 (43,513 ha)
Western Routt County, Non-range, not determined
Former range 169 mi2 (43,772 ha)

Extent of Area - Moffat

INVENTORY
Importance

of Strutting

Grounds

Breeding activity of the sage grouse centers on the strutting ground (or lek).
Strutting grounds in Moffat County were found to be key activity areas within
wintering-nesting
complexes as earlier designated by Wall~stad and Schladweiler
(1974) in Montana.
The fact that strutting grounds hold a strong attraction
for male sage grouse from the earliest heavy snowfall of the fall period to
the end of the breeding period in late spring often made possible the identification of potential sites through observations of male birds or field sign.
Actual strutting grounds were identified through repeated observations of
strutting males on specific sites during the breeding period.

�-135-

High counts of male sage grouse on strutting grounds provides a reasonably
accurate method of determining breeding population trends (Dalke et ale
1963). Jenni and Hartzler (1978) reported that peak numbers of cocks on
strutting grounds can be estimated to within 90 percent by the highest of
3 counts between ~ hr before and l~ hr after sunrise during the first 3 or
the second and third weeks after the peak of breeding activity.
Strutting

Ground Counts

During 1976, 3 new grounds were located and counted and 6 old grounds, which
were no longer counted by Wildlife Conservation Officers, were relocated.
Considerable time and effort was used to become familiar with the various
sage grouse ranges and to locate access roads and trails.
Counts under this
study were made independent of counts secured by Wildlife Conservation Officers.
Table 14 lists a summary of all sage grouse strutting ground counts made
during 1976. A minimum of 961 sage grouse of both sexes were counted on 27
active strutting grounds in Moffat County.
Seven-hundred sixty-six males
were observed on 24 of these grounds for an average of 31.92 cocks per ground.
Numbers of males could not be separated from the total number of birds on
3 strutting grounds counted, so counts made on only 24 or the 27 total grounds
were used to determine this average number of cocks per ground.
In 1977 the number of known active strutting grounds within Moffat and western
Routt counties was more than doubled (Table 15). This was accomplished
through an intensive effort of a ground search crew following helicopter
flight searches and ground searches in the winter period under ideal weather
conditions.
The fact that many flocks of sage grouse males over-wintered in
the vicinity of their strutting grounds also aided this effort.
Totals of
1,977 cocks and 2,441 total birds were counted on 59 act~ve strutting grounds
during the spring period of 1977 for an average of 33.51 cocks and 41.37 total
birds per ground (Table 15).
Table 16 compares counts of sage grouse on 24 strutting grounds counted in
1977 which were also counted during the spring period of 1976. A total of
900 cocks was counted in 1977 compared with 766 in 1976 (percent change
+17.5).
This increase in numbers of cocks counted is most probably a result
of the good hatch which occurred during 1976. More counts were, however,
made in 1977. Due to differences in times when the 1976 and 1977 counts were
started, numbers of total birds counted are not comparable (the 1977 counts
caught the peak of hen attendance, while the 1976 counts did not).
Table 17 lists all strutting grounds according to status: active, relocated,
or abandoned.
Of these 73 grounds, 21 are regularly censused each spring by
W.C.O.'s (active grounds), 11 have not been counted by W.C.O.'s in recent
years, but were relocated by Project personnel during 1976 or 1977, 27 are
new grounds located by Project personnel during 1976 or 1977 (plus 1 additional new ground (80 Road) located by w.c.o. Ch.arles Hector in 1976), and
13 have apparently been abandoned.
A total of 60 active strutting ground
locations was known in 1977, including 1 (Bear Creek) counted only by w.c.o.
L. Vidakovich in 1977. Locations of all known strutting grounds in 1977 are
shown in Figs. 15 and 16.

�Table 14.

Summary of highest cock and total bird counts on sage grouse strutting grounds, Moffat County, Spring 1976.

W.C.O.
District
Rangely (L. Vidakovich)

Name of Ground

Karren Ranch
Escalante
State Line
Haslim Cow Camp

Number
Counts

Date

Time
(A.M.)

1
1
1
1

5-14-76
5-13-76
5-13-76
5-13-76

6:15
6:50
5:40
6:30

Highest Count (Cocks)
Number
Cocks
Counter(s)
D. Hoffman,
D. Hoffman,
D. Hoffman,
D. Hoffman,

J. Corey
J. Corey
J. Corey
J. Corey

Subtotals
1

4-23-76

7:00

D. Hoffman

Brown's Park (R. Lowry)
Subtotals

Powder Wash Hill

Craig N. W. (C. Brown)

North Fork Big Gulch 2
Big Gulch 1
Grassie Reservoir
Lay Creek
Bard Gulch
Greasewood Gulch
Upper 19 Road
Spring Creek 1
Cross Mountain 1
Big Hole Gulch
Cox Ranch
Timberlake Creek 1
Timberlake Creek 2
Big Hole Butte
Dressler Gulch

2
1
1
1
2
1
2
2
1
2
1
2
2
1
1

4-20-76
4-21-76
5-9-76
4-13-76
4-13-76
4-22-76
4-14-76
4-15-76
4-16-76
4-24-76
4-15-76
5-5-76
5-7-76
5-6-76
4-22-76

5:25
5:26
6:53
5:30
6:50
6:10
5:45
6:05
5:30
5:27
6:50
7:05
6:40
7:45
6:28

D. Hoffman,
T. Howard
T. Howard
D. Hoffman,
D. Hoffman,
D. Hoffman,
D. Hoffman
D. Hoffman
D. Hoffman
T. Howard
J. Corey
D. Hoffman
D. Hoffman
D. Hoffman
T. Howard

Fan Rock

1

5-4-76

6:10

D. Hoffman, J. Corey

T. Howard
J. Corey
J. Corey
J. Corey

Subtotals
Slater (C. Woodward)
Subtotals

Total
Birds

26

30
45
45
65
185

26
26

26
26

26

-

31
3
21
32
32
30
32
25
18
19
20
29
60
17
36
405

32
4
21
32
32
30
42
28
19
19
21
31
62
17
38
428

113
113

117

------------------------------------------------------------------------------------------------------------------------

ll7

I
I-'
v.:&gt;
(j\

I

�Table 14. Summary of highest cock and total bird counts on sage grouse strutting grounds, Moffat County, Spring 1976
(continued).
Highest Count (Cocks)
W.C.O.
District

Name of Ground

Craig S. W. (W. Roland)

Juniper
Axial Basin
Dry Lake 2
Deception Creek
Round Bottom
Yellowjacket Road

Number
Counts

Date

Time
(A.M.)

1
2
2
1
1
1

4-16-76
4-20-76
4-21-76
4-21-76
4-25-76
4-20-76

6:40
6:00
6:00
7:00
6:05
5:10

Subtotals

Counter(s)

J. Corey
J. Corey
J. Corey
J. Corey
J. Corey
J. Corey

Number
Cocks

Total
Birds

24
28
57
14
55
18
196

24
31
58
14
57
21
205

I
•.....
W

-..J
I

Totals

766

36
Summary:
Number of active grounds counted
Average number counts per ground
Average number cocks per ground
Average number total birds per ground

=
=
=
=

27
1.33
31.92
35.59

961

�Table 15. Summary Qf highest cocks and total bird counts on sage grouse strutting grounds, Moffat and western Routt
counties, Spring 1977.
W.C.O.
District
Rangely (L. Vidakovich)

Highest Count
Date

Karren Ranch
Escalante
State Line
Haslim Cow Camp

1
1
1
1

5-3-77
5-3-77
5-4-77
5-4-77

5:55
6:30
5:50
6:30

D. Hoffman
D. Hoffman
D. Hoffman
D. Hoffman

65
62
92
40
259

75
70
98
40
283

Powder Wash Hill
Gee Flats
Snake River West
Beaver Basin
Coffee Pot Spring
Summit Spring
Goodman Draw
Sand Wash Road

2
2
2
1
2
2
1
2

4-21-77
4-25-77
4-18-77
4-25-77
4-21-77
5-20-77
5-20-77
3-22-77

5:05
6:00
6:45 •
6:30
6:15
7:00
6:10
7:40

D. Hoffman, L. Crooks 55
D. Hoffman
64
L. Crooks
12
D. Hoffman
101
D. Hoffman, L. Crooks 26
D. Hoffman
19
13
D. Hoffman
D. Hoffman
16
306

57
67
12
107
38
19
14
16
330

North Fork Big Gulch 2
Grassie Reservoir
Lay Creek
Bord Gulch
Greasewood Gulch
Upper 19 Road
Spring Creek 1
Spring Creek 2
Spring Creek 3
Cross Mountain 1
Cross Mountain 2
Big Hole Gulch
Cox Ranch
Timberlake Creek 1
Timberlake Creek 2
Timberlake Creek 3
Big Hole Butte
Dressler Gulch
Sevenmi1e Reservoir
Pole Gulch
Cottonwood Gulch
West Timberlake Creek

1
3
1
1
3
2
1
3
2
3
2
4
2
3
3
2
2
2
2
2
2
2

3-31-77
4-11-77
4-5-77
3-31-77
4-26-77
4-20-77
4-7-77
4-20-77
5-3-77
4-21-77
4-21-77
4-29-77
4-27-77
4-29-77
5-7-77
4-7-77
4-26-77
4-18-77
4-1-77
4-7-77
3-23-77
4-16-77

5:55
7:30
6:05
6:35
6:31
5:15
6:00
6:45
6:27
5:10
5:30
6:55
7:05
5:45
5:45
6:50
7:16
6:57
6:00
6:00
8:20
8:25

D. Hoffman
L. Crooks
D. Hoffman
D. Hoffman
T. Howard
D. Hoffman
D. Hoffman
D. Hoffman
T. Howard
L. Crooks
L. Crooks
T. Howard
T. Howard
D. Hoffman
D. Hoffman
L. Crooks
T. Howard
T. Howard
D. Hoffman
L. Crooks
D. Hoffman
L. Crooks

22
6
34
23
52
33
24
31
16
25

24
8
78
23
54
53
44
31
18
25

13

13

34
16
55
43
21
34
27
14
37
8
5

38
16
55
43
69
34
28
15
76
32
12

Name of Ground

Subtotals
Craig N. W. (C. Brown)

Total
Birds

Time
(A.M.)

Counter(s)

Subtotals
Brown's Park (Vacant)

(Cocks)
Number
Cocks

Number
Counts

------------------------------------------------------------------------------------------------------------------------

I
i-'
W
00
I

�Table 15. Summary of highest cocks and total bird counts on sage grouse strutting grounds, Moffat and western Routt
counties, Spring 1977 (continued).

W.C.O.
District
Craig N. W. (C. Brown)
(continued)

Highest Count (Cocks)
Number
Cocks
Counter(s)

Name of Ground

Date

Time
(A.M.)

Thornburg Well
Lone Tree
Scandinavian Gulch
Conway Spring

2
2
3
2

4-20-77
4-28-77
4-29-77
4-22-77

6:05
6:30
6:10
6:15

L. Crooks
D. Hoffman
T. Howard
T. Howard

36
22
45
3
679

38
24
47
3
901

Fan Rock
Fourmile Creek 1
Fourmile Creek 2
Fourmile Creek 3
Fly Creek
Slater Park

2
5
2
2
1
2

5-3-77
4-16-77
4-5-77
5-7-77
5-11-77
5-21-77

6:05
6:25
7:00
6:30
6:00
5:35

L. Crooks
L. Crooks
L. Crooks
D. Hoffman
D. Hoffman
L. Crooks

116
34
27
40
8
36
261

118
36
39
48
8
38
287

Subtotals
Slater (C. Woodward)

Subtotals

I
I-'

Craig S. W. (W. Roland)

Juniper
Axial Basin
Dry Lake 2
Deception Creek
Round Bottom
Yellowjacket Road
Horse Gulch
Duffy Mountain
Morgan Gulch 1
Morgan Gulch 2
Boxelder Gulch

4
3
1
1
1
2
2
3
2
4
1

4-22-77
4-16-77
4-7-77
4-7-77
4-6-77
4-25-77
4-2-77
5-4-77
4-11-77
4-26-77
4-30-77

5:50
5:30
5:50
6:20
5:55
7:00
6:05
6:50
6:50
6:15
6:30

D. Hoffman
D. Hoffman
J. Corey
J. Corey
D. Hoffman
L. Crooks
D. Hoffman
L. Crooks
D. Hoffman
L. Crooks
D. Hoffman, L. Crooks

31
58
35
22
36
54
4
27
8
39
60
374

33
78
92
30
80
56
8
37
10
40
62
526

29 Road 1
29 Road 2
80 Road
Elkhead Creek

1
2
2
1

3-25-77
5-5-77
3-25-77
5-9-77

6:30
5:45
7:20
6:15

D. Hoffman
D. Hoffman
D. Hoffman
D. Hoffman

4
62
20
12
98

4
68
30
12
114

1,977

2,441

Subtotals
Hayden (C. Hector)

Subtotals
Totals
SUMMARY:.

Total
Birds

Number
Counts

119
Number of active ground counted
Average number counts per ground
Average number cocks per ground
Average number total birds per ground

=
=
=
=

59
2.02
33.51
41.37

w

\0
I

�-140-

Table 16. Comparison of high cock and total bird counts on 24 sage grouse
strutting grounds, 1976 and 1977.

Year
Number
Counts

1976
Number
Cocks

'fotal
Birds

Number
Counts

1977
Number
Cocks

Total
Birds

Karren Ranch

1

26

30

1

65

75

Powder Wash Hill

1

26

26

2

55

57

North Fork Big Gulch 2

2

31

32

1

22

24

Big Gulch 1

1

3

4

2

0

0

Grassie Reservoir

1

21

21

3

6

8

Lay Creek

1

32

32

1

34

78

Bord Gulch

2

32

32

1

23

23

1

30

30

3

52

54

Upper 19 Road

2

32

42

2

33

53

Spring Creek 1

2

25

28

1

24

44

1

18

19

3

25

25

Big Hole Gulch

2

19

19

4

34

38

Cox Ranch

1

20

21

2

16

16

Name of
Ground

Gulch

Greasewood

Cross Mountain

1

Timberlake

Creek 1

2

29

31

3

55

55

Timberlake

Creek 2

2

60

62

3

43

43

Big Hole Butte

1

17

17

2

34

34

Dressler

1

36

38

2

27

28

Fan Rock

1

113

117

2

116

118

Juniper

1

24

24

4

31

33

Axial Basin

2

28

31

3

58

78

Dry Lake 2

2

57

58

1

35

92

1

14

14

1

22

30

1

55

57

1

36

80

1

18

21

2

54

56

33

766

806

50

900

1,142

Deception

Gulch

Creek

Round Bottom
Yellowjacket

Road

Totals
Summary:

number

counts per ground

Average number

cocks per ground

Average

Average

number

total birds per gound

=

1. 38

2.08

31. 92

37.50

33.58

47.58

�Table 17.

Status

Status of sage grouse strutting grounds, Moffat and western Routt counties, 1976 and 1977.
~~
~
Craig N. W.
(C. Brown)

Active 1976 and 1977
North Fork Big Gulch 2

Craig S. W.
(W. Roland)

Juniper

W~i~l~dlife
Conservation Officer District
Hayden
Slater
Brown's Park
(C.
Hector)
(C. Woodward)
(Vacant)

Fan Rock

Powder Wash Hill

29 Road 1

Rangely
(L. Vidakovich)

Karren Ranch
Escalante

Gee Flats

Grassie Reservoir

Axial Basin

Lay Creek

Dry Lake 2

State Line

Bord Gulch

Deception Creek

Greasewood Gulch

Round Bottom

Haslim Cow Camp
Bear Creeka

Upper 19 Road
I
I-'
.l::I-'
I

Timberlake Creek 2
Relocated 1976 and Active 1977
Spring Creek 1
Cross Mountain 1
Big Hole Gulch
Cox Ranch
Timberlake Creek 1
Relocated 1977
Spring Creek 2

Snake River West
Beaver Basin

Cross Mountain 2

Sand Wash Road
Goodman Draw
New 1976 and Active 1977
Big Hole Butte

80 Roadb

Yellowjacket Road

Dressler Gulch
New 1977
Sevenmile Res.

Horse Gulch

Fourmile Creek 1

Coffee Pot Spring

29 Road 2

Timberlake Creek 3

Duffy Mountain

Fourmile Creek 2

Summit Spring

Elkhead Creek

Pole Gulch

Morgan Gulch 1

Fourmile Creek 3

�Table 17.

Status of sage grouse strutting grounds, Moffat and western Routt counties, 1976 and 1977 (continued).
Wildlife Conservation Officer District
Slater
Brown's Park
Hayden
(C. Woodward)
(Vacant)
(C. Hector)

Craig N. W.
(C. Brown)

Craig S. W.
(W. Roland)

New 1977 (continued)
Cottonwood Gulch

Morgan Gulch2

Fly Creek

Boxelder Gulch

Slater Park

Status

West Timberlake Creek

Rangely
(L. Vidakovich)

Thornburg Well
Lone Tree
Scandinavian Gulch
Conway Spring
Spring Creek 3
Abandoned 1977 but Active 1976
Big Gulch 1
Abandoned Prior to 1976
North Fork Big Gulch 1

Dry Lake 1

Clarida Ranch
Fortification Creek
Big Gulch 2
Big Gulch 3
Big Gulch 4

a Counted by L. Vidakovich (1976 and 1977)
b

Located by C. Hector (1976).

c Eliminated by Yampa Power Plant

.-

High Mesac

I

.j:--

Sandwash Ridge Road

Twin Mesa

Hayshed

Twentymile

N
I

�\!;;;:;;====_._

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::son 0

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..

;g
0:1&gt;

"_.

.0.__

,~J~ :z~

d:
go
~.,..~
•... .., ~
c:

5-1
i~!,,0
'"

s,g

~
_
l~:'~
_ o:Ti~

~(!,[t

4::)"'1&amp;

i

~ ~. ~
as
~.

[ ~

h~~
l

.l:r

~~l~:..

~i~;';
~
~
8.

'"

I

.~

-(l]T-

~

~,
....,..==,~,..==~=-=~=---==~~~====

-;..~

, ,

~

~
+1,1

.'O;'1I!:i

'

....

�""'" z

: is"
P;l

=i~.
n§:

f

~
~
~

J

i i
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J

• I ••

~
~
~
~
~

i

-144-

~]1~
~ ~g s

ne~

.'l.J]

:!:~::

~~~~
••
i!~~

�-145-

Of the 13 grounds listed as abandoned, 2 small grounds were active during
1976 (Big Gulch 1 and High Mesa) but apparently faded out of the picture
during 1977, and 11 were apparently abandoned prior to 1976. Of 12 strutting
grounds listed as relocated, 7 were found on or very close to the same sites
where earlier recorded, 1 had moved approximately one-fourth mile (0.4 km),
2 had moved approximately 1 mile (1.6 km), and 2 had moved approximately
one and one-half miles (2.4 km). From these data, it was concluded that
the 1977 breeding population of sage grouse in the Moffat County area was
higher than in 1976. These data indicate an increasing population but further
research is needed to prove or disprove the hypothesis that sage grouse
populations in the Moffat County area are cyclic.
Land ownership status and approximate elevations of 60 sites used by sage
grouse for strutting grounds in the Moffat County area in 1977 are presented
in Table 18. Types of areas utilized by sage grouse for strutting grounds
in the Moffat County area in 1977 are listed in Table 19. Natural clearings
along or near tops of ridges (21 sites) and natural meadows or clearings in
bottoms (14 sites) were the 2 most common types of areas utilized for
strutting grounds.
Brood Counts
With very few exceptions during recent years, brood counts of sage grouse
for management purposes within the Moffat County area have been conducted
sporadically by W.C.O.'s and Regional Biologists because the reliability of
the technique has been questioned.
A method suitable to measure reproductive success and pre-hunting season populations of sage grouse was desired
for improved management of the species, so brood route counts were revived.
1976
A total of 412.6 mi (663.9 km) was driven on 10 brood count routes in 1976
and 564 sage grouse (22 hens, 16 cocks, 90 young, and 436 unclassified) were
counted, for an average of 1.37 birds per mi (0.8S/km) (Table 20). Seventeen
separate broods were counted along these routes before the juveniles became
too large to distinguish from the adults, averaging 5.29 young per brood.
A young to adult ratio of 2.37:1 indicated that sage grouse in Moffat and
western Routt counties experienced good reproductive success in 1976.
Seven
additional sage grouse broods not counted on routes averaged 3.86 young per
brood, bringing the total number of broods
tallied in 1976 to 24.
1977
Table 21 summarizes data from 33 sage grouse brood counts completed along 8
routes in Moffat County (Fig. 15), and 2 routes in western Routt county (Fig.
16), plus one extra route (Sunbeam) in Moffat County.
Counts were made
during the period July 12 through August 11, 1977. Three early morning and
2 late evening counts were secured along 4 routes and 1 or more counts were
secured along the other 7 routes.
A total of 905.1 mi (1,456.3 km) was driven
on these routes and 1,150 sage grouse (151 hens, 179 cocks, 532 young, and
288 unclassified) were counted, for an average of 1.27 birds per mi (0.79/km).
This was slightly lower than the 1.37 birds per mi (0.8S/km) counted along
412.6 mi (663.9 km) on 10 routes in 1976.

�-146-

Table 18. Land ownership
strutting grounds, Moffat

A.

status and approximate elevations
and western Routt counties 1977.

of 60 sage grouse

Number
Grounds

Percent

Private

33

55.0

BLM

22

36.6

State Land Board

3

5.0

DOW

1

1.7

FS

1

1.7

Total

60

100.0

14

23.3

30

50.0

11

18.3

1

1.7

4

6.7

60

100.0

Land Ownership

Status

Owner

B.

Approximate

Elevations

Elevational

Range

6,000-6,500

ft

(1,829-1,981

m)

6,500-7,000

ft

(1,981-2,134

m)

7,000-7,500

ft

(2,134-2,286

m)

7,500-8,000

ft

(2,286-2,438

m)

8,000-8,500

ft

(2,438-2,591

m)

a

Total

a From U. S. Geol. Survey Topographic

Maps.

�-147-

Table 19. Types of areas utilized by sage grouse for strutting
Moffat and western Routt counties 1977.

Number

Type of Area

Natural

clearing

Natural meadow
Natural

or clearing

clearing

Old cultivated

along or near top of ridge
i.nbottom

at intermediate

field

height

along ridge

(go-back land)

Sites

grounds,

a

Percent

21

32.4

14

21.5

5

7.7

5

7.7

Cultivated

field

(wheat)

5

7.7

Sagebrush

cleared

and/or reseeded

4

6.1

Stock pond or reservoir

3

4.6

Irrigated

2

3.1

Old burn

2

3.1

Salt lick

2

3.1

Well site

1

1.5

Road

1

1.5

Totals

65

100.0

a

hay meadow

Five of the 60 strutting
65 sites).

grounds

contained

two sites

(sample based upon

One hundred and eleven separate broods were counted along these routes in 1977
averaging 4.79 young per brood. This average number of young per brood was also
lower than the 5.29 young per brood counted in 1976. A young to adult ratio
of 1.61:1 indicated that sage grouse in Moffat and western Routt counties
experienced only fair overall reproductive success in 1977, and less reproductive success than during 1976 when a young to adult of 2.37:1 was observed.
Seven additional sage grouse broods not counted on routes averaged 4.86 young
per brood, bringing the total number of broods observed in 1977 to 118.
During 1977, 111 of 151 hens (74%) were observed with broods.
Earliest broods were observed during the last week of May 1977, indicating
an early hatch.
Sizes of the young observed along routes confirmed this
early hatch.
Most of the young were difficult to distinguish from adult
hens by August 10, 1977.

�Table 20.

Summary of sage grouse brood counts, Moffat and western Routt counties, 1976.
Sage Grouse Counted
Number
Adults
Minutes Hens Cocks Young Unc1ass. Total Broods

Young/
Brood

Number
Brood1ess
Hens

Birds/
Mile

Date

Time

Miles

Great Divide

8-19-77
8-26-76

A.M.
A.M.

23.6
22.7

110
130

0
0

0
0

0
0

8
28

8
28

0
0

-

0
0

0.34
1. 23

Timberlake

8-5-76
8-24-76

A.M.
A.M.

24.7
31.1

100
120

2
0

0
0

4
0

0
12

6
12

1
0

4.00

-

1
0

0.24
0.39

Big Hole

8-4-76

A.M.

28.4

110

2

3

8

0

13

1

8.00

1

0.46

Spring Creek

8-6-76
8-25-76

A.M.
A.M.

26.8
26.9

120
110

2
1

0
5

9
9

17
1

28
16

2
1

4.50
9.00

0
0

1.04
0.59

Powder Wash

8-11-76
8-31-76

A.M.
A.M.

24.0
30.0

110
130

1
0

0
0

0
0

19
41

20
41

0
0

-

1
0

0.83 .....I
1. 37 +:-co

105

3

0

19

30

52

3

6.33

0

2.43

Name of Route

County

-Moffat

I

Blue Mountain

8-10-76

A.M.

21.4

Cold Spring

8-14-76

A.M.

15.2

160

3

0

11

236

250

3

3.66

0

16.45

Juniper-Axial

8-3-76
8-27-76

A.M.
A.M.

28.1
28.3

105
100

5
0

0
8

16
0

10
6

31
14

3
0

5.33

2
0

1.10
0.49

E1khead

8-18-76
8-30-76

A.M.
A.M.

30.1
28.8

115
110

1
0

0
0

4
0

6
10

11
10

1
0

4.00

-

0
0

0.37
0.35

Slater Park

8-20-76

A.M.

22.5

110

2

0

10

12

24

2

5.00

0

1.07

Totals

412.6

1,845

22

16

90

436

564

17

Averages

25.8

-

-

-

-

-

-

Routt

115.3

-

5
5.29

-

1.37

�Table 21.

Summary of sage grouse brood counts, Moffat and western Routt counties 1977.

Young/
Brood

Number
Brood1ess
Hens

Birds/
Mile

Sage Grouse Counted
County

Number
Adults
Minutes Hens Cocks Young Unc1ass. Total Broods

Date

Time

Miles

Great Divide

7-15-77
7-26-77
7-28-77
7-21-77
8-7-77

A.M.
A.M.
A.M.
P.M.
P.M.

24.9
26.5
25.9
26.9
26.0

150
155
135
148
130

6
6
6
11
1

3
0
0
4
0

17
25
28
53
5

19
0
0
0
23

45
31
34
68
29

5
6
6
10
1

3.40
4.17
4.67
5.30
5.00

1
0
0
1
0

1.81
1.17
1.31
2.53
1.12

Timberlake

7-19-77
7-20-77
7-29-77
8-8-77
8-11-77

A.N.
A.M.
A.M.
P.M.
P.M.

33.9
34.0
36.3
35.4
34.2

115
205
140
135
150

5
5
8
4
2

0
0
0
4
23

10
10
40
0
0

0
60
32
18
43

15
75
80
26
68

2
2
8
0
0

5.00
5.00
5.00

3
3
0
4
2

0.44
2.21
1
2.20
O.73 ~
1.99 1

7-27-77
8-8-77
8-9-77
8-2-77
8-3-77

A.M.
A.M.
A.M.
P.M.
P.M.

27.0
25.8
27.0
27.0
26.9

145
120
120
150
120

1
0
3
2
2

1
10
0
0
0

4
0
12
10
6

0
0
0
0
0

6
10
15
12
8

1
0
3
2
2

4.00

0
0
0
0
0

0.22
0.39
0.56
0.44
0.30

Spring Creek

7-13-77
7-22-77
8-8-77
7-28-77
8-1-77

A.M.
A.M.
A.M.
P.M.
P.M.

26.9
28.2
28.2
27.8
27.1

125
150
125
135
110

7
6
4
2
1

1
0
0
2
0

0
32
16
9
6

0
0
0
0
0

8
38
20
13
7

0
6
4
2
1

5.33
4.00
4.50
6.00

7
0
0
0
0

0.30
1.35
0.71
0.47
0.26

Powder Wash

7-12-77
7-26-77

A.M.
A.M.

27.0
29.3

105
135

0
0

0
7

0
0

0
0

0
7

0
0

-

0
0

0.00
0.24

Name of Route

Moffat

Big Hole

-

4.00
5.00
3.00

----------------------------------------------------------------------------------------------------------------------

I-'

�Table 21.

Summary of sage grouse brood counts, Moffat and western Routt counties 1977 (continued).

Young/
Brood

Number
Brood1ess
Hens

Birds/
Mile

Sage Grouse Counted
Number
Adults
Minutes Hens Cocks Young Unc1ass. Total Broods

Date

Time

Miles

Blue Mountain

7-14-77

A.M.

22.9

l30

6

35

29

24

94

6

4.83

0

4.10

Cold Spring

7-18-77

A.M.

20.7

180

30

48

130

21

229

22

5.91

8

11.06

Juniper Axial

7-28-77
7-25-77
8-4-77

A.M.'
P.M.
P.M.

27.0
27.1
29.7

105
105
l30

0
0
0

0
0
0

0
0
0

0
0
0

0
0
0

0
0
0

-

0
0
0

0.00
0.00
0.00

Name of Route

County

Moffat (cont . )

I
I-'

Sunbeam

8-9-77

A.M.

23.4

115

1

30

2

0

33

1

2.00

0

1. 41

E1khead

7-21-77
.8-3-77
8-4-77

A.M.
A.M.
A.M.

24.8
26.0
26.1

115
160
155

11
7
14

10
0
1

5
27
56

0
0
48

26
34
119

1
7
l3

5.00
3.86
4.31

10
0
1

1.05
1. 31
4.56

Slater Park

7-22-77
7-27-77

A.H.
A.H.

23.4
21.8

110
120

0
0

0
0

0
0

0
0

0
0

0
0

-

0
0

0.00
0.00

Totals

905.1

4,428

151

179

532

288

1,150

111

Averages

27.4

-

-

-

-

-

-

Routt
--

l34.2

40
4.79

1. 27

V1
0
I

�-151-

The majority of sage grouse observations secured during the brood rearing
period were made near existing water sources and nearby meadows (Fig. 17).
This concentrating of birds near waterholes was more pronounced than
found during 1976 and probably was accented by an extended drought throughout
western Colorado (Fig. 18).
Distribution

Checks

Number of birds observed and number of specific locations were greatly influenced by the fact that more time was spent searching areas nearest to headquarters (Craig) during the study. No attempt was made to plan field work
so similar amounts of time and travel could be spent in all ranges.
For this
reason, these data have value for determining distribution by winter and
summer periods for outlining specific areas such as brood rearing and wintering areas, but limited value for comparing populations.
Winter Periods 1976-l978.--Fig. 19 shows winter period distribution of
sage grouse in Moffat County by numbers of birds and locations.
Mortalities,
flocks, and field signs (tracks, roosts, etc.) were noted throughout the
study period and used to determine winter distribution.
Eleven important wintering areas were identified (Fig. 20). These contained
182 mi2 (47,139 ha) of winter range.
In addition,
wintering flocks of sage
grouse (particularly males) were found within 1.5 mi (2.4 km) of over 90
percent of 41 active strutting grounds checked for wintering activity (19761978).
Cocks from nearby wintering flocks were observed to visit most (22 of
36) strutting grounds during the winter periods.
Probable causes of 13 winter
period mortalities are listed in Table 22.
Summer Periods 1976-l977.--Fig. 21 shows summer period distribution of
sage grouse in Moffat County by numbers of birds and locations.
Mortalities,
broods, flocks, and field signs (tracks, roosts, etc.) were noted throughout
the study period and used to determine summer distribution.
Strutting ground
counts and brood counts are recorded under inventory and are not included in
these data.
Two important brood rearing areas were identified (Fig. 20). These consist
of the Class 1 ranges on Cold Spring Mountain and Blue.Mountain.
Broods
observed along brood count routes and miscellaneous brood counts were used
to determine that these two areas are important brood rearing areas.
These
two areas contain approximately 146 mi2 (37,815 ha) of summer range. In
addition, broods were commonly observed throughout most of Class 1 ranges.
Probable causes of 34 non-winter period mortalities are listed in Table 23.
Road kills (13 birds), drownings in stock tanks (8 birds), and predator kills
(7 birds) were the three most probable causes of mortalities during this
period.
HARVEST DATA COLLECTION
Wing Collections
Wing barrel collection stations were first developed by the Arizona Game and
Fish Department and modified by Hoffman and Braun (1975) for use in Colorado.
These have proven valuable for gathering harvest information for management
purposes for a variety of small game species.
Sex and age determination of
the sage grouse wing samples was accomplished by Clait Braun using methods
described by Beck et al. (1975).

�I

""'

VI
N

I

Fig. 17.

Sage grouse broods in brood rearing habitat in meadow area, Cold Spring Mountain, Moffat County.

�I

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V1

w
J

Fig. 18.

Big sagebrush range with scattered junipers along a brood count route, Moffat

County.

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MOFFAT COUNTY
COLORADO

®
•

Winbrlng
KilG02

Syabol

.....®

are

IU.le.2

Hu. ant.
_taring
pcj)Ulotloa

Cold

Spring

56.5

1

Po~Mt!.u.h

25.1

200-390

Blue

Mountain

89.9

2

Blg

20.2

50-100

Toto].

146.4

3

'llIornl&gt;ur9h

'.4

sc-ree

MOuntain

Holo

10
11

c&gt;.lch

rouraile ~

14.5

50-100

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GrNMwod

19.0

200-300

16.2

100-200

CrHIC.

12.2

50-100

North rork Blg Gulch

15.1

50-100

n.c.ption

13.1
14.2

50-100
200-300

14.0

50-100

Spr1n;

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�-157-

Table 22.

Probable

causes of winter

period mortalities

a
1976-l978.

Number Birds

Age and Sex of Birds

3

Mature males (2)
Juvenile female (1)

3

Unknown

age males

3

Juvenile

males

2

Mature male (1)
Juvenile female (1)

Unknown

2

Juvenile male (1)
Unclassified
(1)

Total

13

Cause of Death

Predator

killed on strutting

Poached

from strutting

ground

ground

Hit power line
Road killed on gravelled

a Four month periods

road

(December

1-March

(3)

(3)

31).

1976
Sage grouse wings collected during 1976 through use of 16 wing barrels and
miscellaneous hunter contacts and number of wings collected were: Great
Divide = 129, Cedar Mountain = 127, Dinosaur = 105, Timberlake = 90, Cold
Spring Mountain = 52, Deception Creek = 35, Moffat County Road 4 = 31,
Maybell = 24, Axial Basin = 19, California Park Road = 16, Juniper = 13,
Wolf Creek = 7, Sand Wash = 7, Slater Creek = 6, Powder Wash = 0, Sand Wash
Ridge = 0, North Park Check Station = 44, and miscellaneous hunter contacts
43. In all, 748 wings of sage grouse were collected during September 11-13,
1976.
In considering the different sage grouse ranges from which wing barrels
probably sampled the kill, 401 wings (53.6%) came from the area northwest
of Craig (west of Colorado Highway 13, north of U. S. Highway 40, and east of
the Little Snake River), 112 wings (15.0%) from Blue Mountain, 67 wings (9.0%)
from the area southwest of Craig (west of Colorado Highway 13, south of U. S.
Highway 40, and east of Deception Creek), 52 wings (7.0%) from Cold Spring
Mountain, 22 wings (2.9%) from the area east of Colorado Highway 13 to north
of Hayden, and seven wings (0.9%) from Sand Wash.
An additional 87 wings
(11.6%) were secured from check stations in North Park and through miscella~eous hunter contacts.
Age and sex composition of the wing sample in 1976 is listed in Table 24.
Females comprised 84.1 percent of the adult segment of the harvest, 64.3
percent of the subadu1t or yearling segment of the harvest, and 57.4 percent
of the immature or juvenile segment of the harvest.
These data suggest
differential mortality occurred in the Moffat County area in 1976. These
data further suggest that segregation of sexes had started in the Moffat
County area prior to the hunting season.

�-158-

Table 23.

Probable

causes of non-winter

Cause of Death

mortalities

Number Birds

Road killed on pavement

Road killed on gravelled

road

1976-1977.a

Age and Sex

of

Birds

8

Juvenile male (1)
Yearling female (1)
Mature females (2)
Juvenile unknown
sexes (4)

5

Juvenile male (1)
Yearling male (1)
Juvenile female (1)
Mature female (1)
Juvenile unknown
sex (1)

8

Juvenile unknown
sexes (8)

Drowned

in stock tank

Predator

killed on strutting

ground

5

Mature male (1)
Unknown age males (3)
Mature female (1)

Predator

killed off strutting

ground

2

Unknown age male (1)
Juvenile female (1)

Poached

in deer season

2

Mature males

Hit power line

1

Juvenile

Hit fence

1

Mature male

Unknown

2

Juvenile male (1)
Unclassified (1)

Total

34

a Eight month periods

(April I-November

(2)

female

(1)

(1)

30).

Analysis of wing molt patterns of adult and subadult females revealed that 97
of 132 (73.5%) adults and 49 of 74 (66.2%) subadult or yearling hens were
probably successful nesters in 1976. These data and the high percentage of
immatures in the harvest sample (63.6%) indicate that nesting success and
production were good in the Moffat County area in 1976. The percentage of
yearlings in the harvest (15.4%) indicates reproduction during 1975 was at
least fair.
In 1976, 2.3 chicks per hen (3.3 per successful hen) were present
in the harvest.
No comparable data are available for the Moffat County area
for the previous years.
A sample of 108 sage grouse wings collected in 1976
in Middle Park indicated 2.5 chicks per hen and 3.8 chicks per successful hen.

�Age and sex composition of the sage grouse harvest, Moffat and western Routt counties 1976-1977.

Table 24.

Males
Year

No.

%

Immatures
Females
%
No.

Total
No.

%

Males
%
No.

Yearlings
Females
%
No.

Total
%
No.

Males
%
No.

Adults
Females
%
No.

Total
%
No.
I
I-'

1976

203

42.6

273

57.4

476

63.6

41

35.7

74

64.3

115

15.4

25

15.9

132

84.1

157

21.0 ~

1977

121

39.3

187

60.7

308

43.0

77

45.8

91

54.2

168

23.4

89

36.9

152

63.1

241

33.6

�-160-

Approximate dates of hatching determined by backdating, utilizing stages of
primary molt were estimated for all but one chick in the harvest sample
(N = 439). Estimated hatching dates ranged from May 20 to July 15, with the
overall average peak of the hatch occurring during the period of June 10-14.
This peak was estimated to be approximately 10 days later than the overall
average for Cold Spring Mountain at elevations from 8,000 to 8,500 ft (2,4382,591 m) and approximately 7 to 9 days earlier than the overall average for
the Axial Basin-Juniper-Deception
Creek area at elevations varying from 6,200
to 6,500 ft (1,890-1,981 m). Peak numbers of hens on strutting grounds
located at intermediate elevations such as in the Great Divide area probably
occurred during the week of April 11-17, 1976. Here, as with peak hatching
dates, adjustments must be made for peak period of hen attendance of strutting
grounds according to elevations.
The average age of 188 male chicks was 91.5 days compared with an average
of 251 female chicks of 93.0 days. This variation was small, considering
difficulty in assigning accurate ages from growth charts.

age
the

1977
Wing collection barrels were placed at 15 sites in Moffat County (Fig. 15)
and at 1 site in western Routt County (Fig. 16). Number of wings collected
were: Cedar Mountain = 137, Dinosaur = 104, Cold Spring Mountain = 102,
Great Divide = 85, Timberlake = 56, Fourmile Creek = 49, California Park
Road = 36, Moffat County Road 4 = 27, Deception Creek = 21, Lay Creek = 21,
Moffat County Road 2 = 5, Maybell = 5, Juniper = 5, Axial Basin = 5, Elk
Springs = 2, Ralph White Reservoir = 1, Middle Park Check Station = 40, and
wing survey envelopes = 16. In all, 717 wings of sage grouse were collected
during September 10-16, 1977. Thirty-one wings from sharp-tailed grouse, and
24 wings from blue grouse were collected in wing barrels, incidental to sage
grouse wing collections.
During 1976, 748 wings of sage grouse, 12 wings of
sharp-tailed grouse, and 34 wings of blue grouse were collected.
In considering the different sage grouse ranges from which wing barrels
probably sampled the kill, 331 wings (46.2%) came from the area northwest of
Craig (west of Colorado Highway 13, north of U. S. Highway 40 and east of the
Little Snake River), 106 wings (14.8%) from Blue Mountain, 102 wings (14.2%)
from Cold Spring Mountain, 91 wings (12.7%) from the area east of Colorado
Highway 13 to north of Hayden, 31 wings (4.3%) from the area southwest of
Craig (west of Colorado Highway 13, south of U. S. Highway 40 and east of
Deception Creek), 40 wings (5.6%) from the check station in Middle Park, and
16 wings (2.2%) from the wing survey envelopes.
Age and sex composition of the sage grouse harvest sample for 1977 are listed
in Table 24, along with comparable data for 1976. During 1977, females
comprised 63.1 percent of the adult segment of the harvest, 54.2 percent of
the yearling segment, and 60.7 percent of the immature segment.
During 1977
243 adult and yearling hens were harvested out of 409 adult and yearling
sage grouse (59.4%), indicating there was no selection shown towards smaller
birds (hens and/or broods).

�-161-

Analysis of wing molt patterns of adult and yearling females in the sample
revealed that 82 of 152 (53.9%) adult hens and 51 of 91 (56.0%) yearling
hens were probably successful nesters in 1977, compared with 73.5 percent
adult hens and 66.2 percent yearling hens probably being successful in 1976.
Immatures made up 43.0 percent of the total harvest sample in 1977, compared
with 63.6 percent in 1976. In 1977, 1.3 chicks per hen (2.3 per successful
hen) were present in the harvest sample, compared with 2.3 chicks per hen
(3.3 per successful hen) for 1976. From these data it was concluded that
overall reproductive success within the Moffat County area was only fair during
1977, even though reproductive success in the higher elevational ranges was
good, with 52.2 percent immatures in the harvest sample.
Reproductive success
within the lower and intermediate elevational ranges in 1977 was poor, with
only 34.8 percent immatures in the harvest sample.
The percentage of yearlings
in the harvest sample in 1977 (23.4%) confirmed the previously known fact that
overall reproductive success in 1976 was good.
Approximate dates of hatching determined by backdating, utilizing stages of
primary molt were estimated for all chicks in the harvest sample (N = 308).
An estimated hatching period from May 23 to July 8 was determined for 1977,
compared with a hatching period from May 20 to July 15 determined for 1976.
Estimated peak of the hatch for 1977 occurred during the period June 8 to
June 14, similar to that found in 1976 (June 10-14).
The peak of the hatch
on Cold Spring Mountain (elevations 8,000-8,500 ft or 2,438-2,591 m) was
delayed approximately five days in comparison with other ranges.
An average
age determined for 121 male chicks of 90.0 days was similar to the average
age determined for 187 female chicks of 90.2 days.
LITERATURE

CITED

Beck, Thomas D. I., R. Bruce Gill, and C1ait E. Braun.
1975.
Sex and age
determination of sage grouse from wing characteristics.
Game Infor.
Leaflet No. 49 (Revised).
Colo. Div. of Game Fish and Parks.
4 p.
Braun, Clait E., and Thomas D. I. Beck.
1976. Effects of sagebrush control
on distribution and abundance of sage grouse.
Job Final Report W-37-R-29,
Work Plan 3, Job 8a. Colo. Div. Wildl. pp. 21-84.

--- , Tim Britt, and Richard O. Wallestad.
of sage grouse habitats.

Wild1.

1977. Guidelines
Soc. Bull. 5(3):99-106.

for maintenance

Dalke, Paul D., Duane B. Pyrah, Don C. Stanton, John E. Crawford,
F. Schlatterer.
1963. Ecology, productivity, and management
grouse in Idaho. J. Wild1. Manage. 27(4):811-841.

and Edward
of sage

Edminster, Frank C. 1954. American game birds of field and forest.
Scribner's Son, New York.
pp. 114-134.
Eng, Robert L., and Philip Schladweiler.
and habitat use in Central Montana.
Gill, R. Bruce.
1965. Distribution
grouse in North Park, Colorado.
Collins.
187 pp.

Charles

1972. Sage grouse winter movements
J. Wildl. Manage. 36(1):141-146.

and abundance
M. S. Thesis.

of a population of sage
Colo. State Univ., Fort

�-162-

Harrington, H. D. 1954.
Denver.
666 p.

Manual of the plants of Colorado.

Sage Books,

Hoffman, Richard W., and C1ait E. Braun. 1975. A volunteer wing collection
station.
Game Infor. Leaflet No. 101. Colo. Div. Wildlife.
3 p.
Jenni, Donald A., and Jonathan E. Hartzler.
1978. Attendance at a sage
grouse 1ek:imp1ications for spring censuses.
J. Wi1d1. Manage. 42(1):46-52.
K1ebenow, Donald A. 1969. Sage grouse nesting and brood habitat
J. Wi1d1. Manage. 33(3):649-662.

in Idaho.

Krager, Ron. 1977. Survey of sage grouse strutting ground complexes and
seasonal use areas within the Piceance Basin Wildlife Habitat area.
Colo. Div. Wildlife.
46 p , Mimeo.
Moffat County Planning Department.
area of Moffat County.
67 p.

1977.

Comprehensive

plan for the Craig

Rogers, Glenn E. 1964. Sage grouse investigations in Colorado.
No. 16. Colo. Game, Fish and Parks Dept. 132 p.

Tech. Pub.

Wa11estad, Richard o. 1971. Summer movements and habitat use by sage grouse
broods in central Montana.
J. Wi1d1. Manage. 35(1):129-136.
_____ , and Duane Pyrah.
1974. Movement and nesting of sage grouse hens in
central Montana.
J. Wi1d1. Manage. 38(4):630-633.

----- , and Philip Sch1adwei1er.

1974. Breeding season movements and habitat
selection of male sage grouse. J. Wi1d1. Manage. 38(4):634-637.

Prepared by

~~o?!an~~
Wildlife Researcher

C

�April 1979

-163-

JOB PROGRESS REPORT

State of

COLORADO
------------------------------

Project No.
Work Plan No.
Job Title
Period Covered:
Personnel:

Game Bird Survey

W-37-R-32

Job No.
11
Vulnerability and Population
Characteristics of Sage Grouse in Hoffat County
3

--------~-----------------------

April 1, 1978 through March 31, 1979

Tom Beck, Clait Braun, John Corey, Larry Crooks, Jim Dingman,
Howard Funk, Ken Giesen, Dave Hoart, Don Hoffman, Richard Hoffman,
Tom Lines, Debra Martin, Russ Mason, Sue McElderry, Lou Vidakovich,
Nancy Watkins, Colorado Division of Wildlife.

ABSTRACT
Population characteristics, harvest statistics and vulnerability to hunting
were investigated for sage grouse (Centrocercus urophasianus) in Moffat and
Western Routt counties, Colorado.
Counts of males on 54 leks averaged 37.2,
up from 33.5 cocks/lek counted on 59 leks in 1977. Nine new leks were located
in 1978. Counts of sage grouse along 7 brood routes in 1978 averaged 2.85
birds/mile.
Five hundred and six sage grouse were banded in 1978 (421 young
of the year, 38 yearlings and 47 adults) primarily through use of cannon nets
and drive traps. Free permits were required of all sage grouse hunters in
Moffat County in 1978. A total of 2,006 was issued, of which 1,502 permit
holders hunted and harvested 5,185 sage grouse. Hunter success was good
(75.1%) and each successful hunter bagged an average of 4.2 grouse. Approximately 66% of the total harvest occurred during the initial weekend of the 9
day season. Check stations and volunteer wing collection barrels were used to
obtain harvest statistics and wings. A total of 770 hunters was checked at
3 check stations.
These hunters harvested 1.9 grouse each. Hunter success
was best at Cold Spring Mountain and Blue Mountain.
Analysis of 2,418 wings
from sage grouse bagged in Moffat and western Routt counties indicated that the
harvest was comprised of 68.7% young of the year, 9.9% yearlings and 21.4%
adults.
There were 3.1 chicks per female in the harvest sample. The sex
ratio at hatching approximated 1:1 with differential survival favoring females
occurring in the older age classes.
There were 2 females to each male in the
adult and yearling segment of the fall population.
Estimated turnover of adult
males was 43.3%, while for adult females it was 33.2%. Young males were
slightly more vulnerable than young females to hunting (9.2 vs 7.8%). The
overall harvest rate was 9.5%, not markedly different than that documented
from 1973-1978 in North Park.

�-164-

RECOMMENDATIONS

1.

Counts of males on selected leks should be continued as a management
function.

2 ...Trapping and banding should be expanded with a uu.n unum goal of 500 birds
to be banded, 100 each in 5 different units of Moffat County.
3.

Counts of birds along established

brood routes should be discontinued.

4.

The free permit requirement
should be continued.

5.

Check stations should be operated at 3 locations on the initial 2 weekends
of the sage grouse season in Moffat County in 1979 (Cedar Mountain, Maybell,
Dinosaur) .

6.

If data collected continue to be similar to those collected in North Park,
the intensive sage grouse study in Moffat County should be discontinued
at the end of the 1979-1980 segment.

7.

Collections of sage grouse wings from Moffat County should be continued
as a management function.

for all sage grouse hunters in Moffat County

�-165-

VULNERABILITY AND POPULATION
CHARACTERISTICS OF SAGE GROUSE IN MOFFAT COUNTY
C1ait E. Braun and Donald M. Hoffman

Until recent years, sage grouse have been hunted annually in Colorado, with
season lengths typically of 3 days and bag and possession limits of 2 and 4.
Intensive bandings of adult and yearling sage grouse along with experimental
hunting seas~ns in North Park, Jackson County, Colorado have indicated that
only about 10% of the fall population of these segments is annually harvested.
Data from this area indicate that yearling males are most vulnerable, with
adult males and hens of both age classes having lower rates of exploitation.
Knowledge concerning population characteristics and havest rates are important
if maximum allowable recreational opportunity through hunting is to be achieved.
If turnover rates are moderately high (40-60%) and recovery rates (and by
relationship, vulnerability rates) are low, conservative seasons and bag limits
have little merit.
This report covers the initial year of a study to investigate population characteristics and vulnerability to hunting of sage grouse in
Moffat County and adjacent areas in Routt County, Colorado.
Period covered
is from mid-March 1978 through February 1979.
P. N. OBJECTIVES
1.

Estimate young of the year vulnerability
adults and yearlings of each sex.

2.

Provide reliable

estimates

3.

Provide reliable

data on characteristics

4.

Estimate

of harvest

to hunting

in relationship

to

rates (recovery rates).
of the harvest.

turnover and survival rates.
SEGMENT OBJECTIVES

1.

Counts of males and females will be conducted on a mlnlmum of 28 leks
within Small Game Management Units 16 and 18 during April and May. A
minimum of 4 early morning counts will be made per 1ek. Additional new
leks will be located through ground searches.

2.

Counts of chicks and adults will be made along 7 established routes within
Small Game Management Units 16 and 18 during the period July 5 to August
20. A minimum of 2 early morning and two late evening counts will be made
per route.

3a. Chicks will be captured and banded within brood rearing areas during July,
August and early September through use of cannon nets, drive traps and/or
night lighting.
A minimum of 300 (150 male and 150 female) chicks is
desired.
Aluminum and year color coded leg bands will be used to identify
birds by location.

�-166-

3b. Adult and yearling sage grouse will be banded in conjunction with the
banding of chicks whenever possible.
Aluminum and year color coded leg
bands will be used to identify birds and location.
4a. Sage grouse hunting will be by free permits, unlimited in number, in
Moffat County.
All hunters will be mailed a questionnaire within 1 week
of the end of the hunting season.
One follow up letter will be sent 3
weeks later.
4b. Check stations will be operated at least during both days of the opening
weekend near Cedar Mountain, Dinosaur and Cold Springs Mountain to collect
harvest data, wings, and compliance with the permit requirement.
4c. Wing barrel collection stations will be located at 16 sites to sample
the harvest within Moffat and Western Routt counties.
A sample of 600
wings is desired.
4d. Numbers and locations of banded birds shot will be recorded
hunter checks, check stations and hunter questionnaires.

from field

4e. Hunter success will be determined
stations.

and check

from hunter questionnaires

4f. Age and sex composition of the harvest sample of wings will be determined
in Fort Collins following the open season.
4g. Total harvest will be calculated
5.

Vulnerability

estimates

6.

Data will be compiled,

from the hunter questionnaire

survey.

will be made based upon band recoveries.
results analyzed,

and a progress

report prepared.

METHODS AND MATERIALS
Three men using 3, 4-wheel drive vehicles counted all known leks within the
primary study area (Fig. 1) during late March to mid-May 1978. Counts were
also made on leks on Cross Mountain and west of the Snake River.
Attempts were made to make a minimum of 4 early morning counts per lek where
possible using procedures described by Rogers (1964). L~ks counted by District
Wildlife Managers within the primary study area were also counted by project
personnel in 1978.
Intensive efforts were again made to locate as many new leks as possible.
Searches of wintering areas during December 1977-March 1978 were made to locate'
sites showing a definite attraction to cocks.
Eight and one-half hours wRre
flown
in 3 morning helicopter flights during April 22-24, to search for
new leks. Routine early morning searches along roads and trails were made
using procedures outlined by Rogers (1964).
Locations of all known active leks within Moffat
were mapped for future reference.

and Western Routt counties

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Fig. 1.

Moffat County, Colorado sage grouse study area.

Pr1aary
Brood

•

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0\ecIc

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�-168-

Two early morning and 2 late evening counts were completed along 7 established
brood routes using procedures described by Rogers (1964). Early morning routes
were usually started as soon as it became light enough to see well and most
evening routes were started at 6:00 P.M. MDT. Routes requiring extra time
were started earlier.
Routes were run going away from the sun whenever possible
for improved visibility.
Sage grouse were trapped in July and August at night where they roosted along
trails and throughout the early morning and evening where they concentrated
in meadows.
Methods used involved spotlighting and capture with long handled
nets, wire drift fences with a cannon net at the end of the fence, bumper
mounted cannon nets, and modified lily pad traps (Braun and Beck 1976, Lacher
and Lacher 1964, Gullion 1965).
Birds captured were weighed, measured,
classified as to age and sex (Eng 1955, Beck et al. 1975) and banded with
serially numbered aluminum leg bands (size 16 for males, 14 for females)
and plastic bandettes color coded for year and location (5 different areas in
Moffat County).
Sage grouse hunters in Moffat County in 1978 were required to have in their
possession while hunting a free numbered permit. -Permits, unlimited in number,
were available from Division of Wildlife offices in Fort Collins, Grand Junction,
Meeker, Craig, and Denver, all license agents in the Moffat County area, and
all project and management personnel working in Moffat County.
Questionnaires
were sent to all permittees immediately following the end of the sage grouse
hunting season in Moffat County.
One follow up letter and questionnaire were
sent in mid-October to all non-respondents
to the initial mailing.
Check stations were operated at 3 locations (Cedar Mountain, Maybell and
Dinosaur) on both weekends (except Maybell which was not open during the 2nd
Saturday) of the hunting season.
Each station was operated from about 0800
to 1900 MDT, depending upon traffic load. Data obtained per party were:
county of origin, number of hunters, hours hunted (total of all hunters), birds
bagged, birds lost, number of banded birds and location of where each was
obtained, area hunted, and information on previous sage grouse hunting
experience.
One wing was obtained for most· birds checked with ovaries being
taken or examined from a sample of yearling and adult hens.
Efforts were made
to ascertain sex by gonadal inspection for a sample of birds.
These data were
recorded on tags with wings being individually marked with corresponding
information concerning actual sex of that bird. Wings were frozen and stored
for later analysis.
In addition to the 3 check stations, wing barrels and signs (Hoffman and Braun
1975) were placed at 16 locations in Moffat and Western Routt counties,
including the 3 check station sites. Volunteer wing collection stations were
in operation the entire season at all sites. Wings were collected following
the 1st weekend, midweek, and following the 2nd weekend.
Collected wings were
frozen for later analysis.
Collected wings were thawed and classified to age (chicks, yearlings, adults)
and sex following procedures outlined by Eng (1955) and Beck et al. (1975).
Hatching dates were calculated for chicks of the year using data from Pyrah
(1963).
Ovaries collected were stored in AFA (alcohol, formalin and acetic acid) and
later examined for presence of ovulated follicles as described by Kabat et al.
(1948).

�-169-

DESCRIPTION

OF AREA

The primary study area (Fig. 1) consisted of that portion of Moffat County
north of Colorado Highway 318 from the Utah state Line to Sunbeam, east of
Cedar Springs Draw (Moffat County Road 23), north of T4N from near Escarpment
Peak to the junction of Colorado Highway 13 and Moffat County Road 45, west
of Colorado Highway 13 and west of Moffat County Road 101. In addition,
harvest data were collected from Blue Mountain and western Routt County with
inventory data (brood route counts) also being collected from western Routt
County.
Moffat County is located in extreme northwestern Colorado, bordered by Wyoming
on the north, Utah on the west, Rio Blanco County on the south and Routt
County on the east. The major drainages include the Green River which flows
north to south through western Moffat County, the Yampa River which flows east
to west in the southern part of the county to its intersection with the Green
River, approximately 3 miles (4.8 km) from the Utah border, and the Little
Snake River which flows northeast to southwest through the middle of the county
where it joins the Yampa River in Lily Park.
The climate in Moffat County is semi-arid with 8 to 20 inches (20.3-50.8 cm) of
precipitation annually.
Precipitation occurs mainly in late summer to early
winter.
Mean annual precipitation for 6 weather stations was 12.99 inches
(33.0 cm). The annual mean temperature for 5 stations was 43.30F (6.30C)
during years of record.
Elevation in Moffat County varies from approximately 4,600 to 11,045 feet
(1,402-3,367 m). The county seat, Craig, located in eastern Moffat County,
is 6,240 feet (1,902 m) above sea level.
2
Sagebrush ranges comprise approximately 60% (2,841 mi ) of the total land
area of Moffat County.
Non-sagebrush ranges consisting of pinon-juniper,
rough and rocky mountain shrub, aspen, or spruce-fir types comprise most of
the remaining 40% (1,902 miles2) of the total land area.
RESULTS AND DISCUSSION
Counts of Sage Grouse on Display Areas
In 1978, 2,008 cocks and 2,163 total birds were counted on 54 grounds.
This
is an average of 37.2 cocks/lek (Table 1), slightly higher than the average
of 33.5 cocks/lek counted on 59 leks in all of Moffat and Western Routt
counties in 1977. District Wildlife Managers (DWM's) counted 19 active leks
within the primary study area in Moffat County in 1978 with 757 cocks and 780
total birds (average = 39.8 cocks/lek, 41.1 total birds/lek).
In addition
DWM's secured counts on 6 large leks outside the primary study area within
Moffat County which were not counted by project personnel.
These 6 leks had
a total of 634 cocks and 637 total birds (105.7 cocks and 106.2 total birds/
lek).

�Table l.

Sage grouse 1ek counts, Moffat County 1978.

Lek

No. of
Counts

--No. of
Sites

Brown's Park (Vacant)
Gee Flats
Powder Wash Hill
Snake River West
Beaver Basin
Coffee Pot Spring
Summit Spring
Goodman Draw
Sand Wash Road (Ab~ndoned)
East Sand Wash Road (New)
Total

3
4
4
3
4
2
3
0
4
27

1
1
1
2
1
1
2
0
1

D.W.M.
District

First
High
a
Count

1
2

3

-

3

Highest Count (Cocks)
Date

Time
A.M.b

Counter(s)

4-18-78
4-11-78
4-8-78
5-10-78
4-26-78
5-2-78
5-10-78

5:25
5:35
7:15
6:15
6:15
8:20
7:05

D. Hoffman, L. Crooks
L. Crooks, J. Corey
D. Hoffman
D. Hoffman, L. Crooks
1. Crooks
D. Hoffman, L. Crooks
D. Hoffman, L. Crooks

4-26-78

4:45

L. Crooks

No. of
Cocks

Total
Birds

55
69
8
153
29
18
42
0
28
402

60
70
8
179
29
18
43
0
28
435

I
f-'

-..J

0

Craig, Northwest (C. Brown)
North Fork Big Gulch 2
Grassie Reservoir
Lay Creek
Bord Gulch
Greasewood Gulch
Upper 19 Road
Spring Creek 1
Spring Creek 2
Spring Creek 3
Cross Mountain 1
Cross Mountain 2
Big Hole Gulch
Cox Ranch
Timberlake Creek 1
Timberlake Creek 2
Timberlake Creek 3
Big Hole Butte
Dressler Gulch
Sevenmi1e Reservoir
Pole Gulch

I

4
4
2
4
3
4
4
4
4
5
3
4
4
4
4
4
4
,4
4
4

1
1
2
1
1
1
1
1
1
1
1
1
1
1
2
1
2
1
1
1

1
2

1

1

1
1
4
5

1
1
4
2
1
2
1
1
2

4-8-78
4-8-78
4-27-78
4-8-78
4-21-78
4-3-78
4-3-78
4-3-78
4-20-78
4-28-78
4-12-78
4-3-78
4-3-78
4-26-78
4-7-78
3-25-78
4-14-78
4-6-78
4-4-78
4-13-78

5:30
5:45
5:30
6:16
6:00
7:00
5:45
6:20
5:55
5:40
6:55
7:05
7:15
5:00
6:10
7:20
6:10
6:05
5:50
5:25

J. Corey
J. Corey
J. Corey
J. Corey
J. Corey
L. Crooks
1. Crooks
L. Crooks
J. Corey
D. Hoffman
D. Hoffman
D. Hoffman
D. Hoffman
D. Hoffman
L. Crooks
D. Hoffman
D. Hoffman, J. Tanke
D. Hoffman
D. Hoffman
D. Hoffman, B. Dupire

37
22
57
37
20
30
24
42
12
31
5
37
20
53
66
20
37
25
7
87

-----------~------------------------------------------------------------------------------------------------------------

41
28
59
43
20
31
27
44
12
31
5
51
20
53
71
53
38
37
7
87

�Table 1.

D.W.M.
District

Sage grouse 1ek counts, Moffat County 1978 (continued).

Lek

No. of
Counts

Craig, Northwest (C. Brown) (continued)
3
Cottonwood Gulch
4
West Timberlake Creek 1
4
Thornburg Well
4
Lone Tree
4
Scandinavian Gulch
4
Thornburgh Gulch (New)
4
Mud Spring Draw (New)
West Timberlake Creek 2
3
(New)
107
Subtotal

No. of
Sites

First
High
Counta

Highest Count (Cocks)
Date

Time
A.M.b

Counter(s)

No. of
Cocks

Total
Birds

1
1
1
1
1
1
1

-

2
4
2
1
1
2

4-27-78
4-10-78
4-17-78
4-7-78
4-3-78
4-10-78
4-25-78

5:00
7:35
5:35
6:00
6:35
5:35
5:05

L. Crooks
D. Hoffman
L. Crooks
D. Hoffman
D. Hoffman
L. Crooks
L. Crooks

38
3
32
24
42
17
76

38
3
35
24
45
17
79

1

-

5-11-78

6:00

L. Crooks

47
948

51
1,050

4
5
5
4
18

1
1
3
1

3
1
4
1

4-20-78
3-25-78
4-20-78
5-1-78

5:55
6:35
5:10
7:05

L. Crooks
D. Hoffman
L. Crooks
L. Crooks

36
18
136
11
201

38
19
l38
12
207

Craig, Southwest (W. Roland)
Juniper 1
Axial Basin
Dry Lake 2
Deception Creek
Round Bottom
Horse Gulch
Duffy Mountain
Morgan Gulch 1
Morgan Gulch 2
Boxe1der Gulch 1
Juniper 2 (New)

4
4
4
3
3
2
4
4
4
4
4

1
1
2
1
1
1
2
1
2
1
1

2

4-4-78
3-22-78
5-9-78
5-4-78
4-27-78
4-27-78
4-21-78
3-22-78
4-6-78
4-12-78
4-4-78

6:10
6:10
6:10
6:15
6:00
6:30
6:15
7:55
6:35
6:30
6:25

L. Crooks
D. Hoffman
D. Hoffman
D. Hoffman, L. Crooks
L. Crooks
J. Corey
D. Hoffman
D. Hoffman
L. Crooks
L. Crooks
L. Crooks

45
34
56
17
50
35
14
4
43
44
15

48
34
59
17
51
35
14
4
44
44
16

1

4
1
2
3
2

------------------------------------------------------------------------------------------------------------------------

"-I

I-'
I

Slater (C. Woodward)
Fourmile Creek 1
Fourmile Creek 2
Fourmile Creek 3
Cowboy Reservoir (New)
Subtotal

4

I
I-'

�Table L

D.W.M.
District

Sage grouse 1ek counts, Moffat County 1978 (continued).

No. of
Counts

Lek

Craig, Southwest (W. Roland) (continued)
Morgan Gulch 3 (New)
4
4
Boxe1der Gulch 2 (New)
Temple Gulch (New)
4
52
Subtotal

Highest Count (Cocks)

No. of
Sites

First
High
Counta

Date

Time
b
A.M.

1
1
1

4
3
3

4-12-78
4-12-78
4-12-78

6:15
6:55
5:25

204

Total

54
=

b Times are MST to 4-29-78 and MDT after 4-30-78.

3.8
37.2

Average number cocks/1ek

a Leks with a minimum of 4 counts.

Total
Birds

24
44
32
457

25
45
35
471

2,008

2,163
-....

Number of active leks counted

Average number total birds/1ek

J. Corey
J. Corey
J. Corey

No. of
Cocks

I
f--'

SUMMARY:

Average number counts/1ek

Counter(s)

"&lt;..

40.1

N
I

�-173-

Comparisons were made of numbers present on 45 leks counted by project
personnel in 1977 and 1978 (Table 2). A total of 1,714 cocks was counted in
1978 compared with 1,387 in 1977 (+23.6%).
Total birds counted were 1,855
in 1978 and 1,805 in 1977 (+2.8%). These data, along with an estimated 17.5%
more cocks in 1977 compared with 1976, indicates the breeding population of
sage grouse in Moffat County is increasing.
Nine out of 54 leks counted in 1978 (16.7%) had more than 1 location where
birds regularly displayed compared with 5 out of 60 counted in 1977 (8.3%).
Leks with more than 1 display site were considered as 1 lek in both years.
A minimum of 4 counts was secured on 42 of the 54 leks counted in 1978. The
1st count on the 42 grounds resulted in the highest cock count on 17 leks
(40.5%), compared with the 2nd count being highest on 11 leks (26.2%), the
3rd count being highest on 6 leks (14.3%), with the 4th or 5th count being
highest on 8 leks (19.0%)
Nine new leks were located and counted in 1978, and one lek aGtive in 1977
was abandoned.
Intensive searches of wintering areas which had a definite
attraction for cocks resulted in the location of 4 new leks. Routine early
morning searches along roads and trails provided an additional 4 new leks,
with 1 additional new lek being located through helicopter flights.
Fifteen leks within Moffat County were counted by both project personnel and
DWM's in 1978 (Table 3). An average of 44.3 cocks/lek was obtained (54 counts)
by project personnel compared to an average of 44.1 cocks/lek (31 counts) on
different mornings by 3 DWM's.
Even though project personnel made more counts
per lek (3.6) than DWM's (2.1) and there were variations in maximum numbers
of cocks counted on all but 1 lek, overall averages were similar.
Total birds
averaged 48.4 per lek in 54 counts by project personnel, compared with 45.5
in 31 counts by 3 Dl~'s.
The higher average number of total birds recorded
by project personnel probably was because DWM's began their counts later
(early April 1978) than project personnel (March 20, 1978).
Known sage grouse leks in Moffat County were classified as active, new or
abandoned in 1978 (Table 4). Some leks were not classified in 1978 due to
reduction in area inventoried or isolated location.
These were:
D.W.M. District

No. of Leks

Names
State

Rangely

(L. Vidakovich)

4

Karren Ranch, Escalante,
Line, Haslim Cow Camp

Slater

(C. Woodward)

3

Fan Rock, Fly Creek,

Hayden

(C. Hector)

4

29 Road 1, 29 Road 2, 80 Road,
Elkhead Creek

Craig SW (W. Roland)

1

Yellowjacket

Craig NW (C. Brown)

1

Conway Spring

Total

13

Slater Park

Road

Locations of 50 active leks within the primary study area (Fig. 1) and 17
active leks outside the primary study area were mapped for future reference.

�Table 2.

Peak counts of sage grouse, 45 leks, Moffat County 1977 and 1978.
Year

Lek
Gee Flats
Powder Wash Hill
Snake River West
Beaver Basin
Coffee Pot Spring
Summit Spring
Goodman Draw
North Fork Big Gulch 2
Grassie Reservoir
Lay Creek
Bord Gulch
Greasewood Gulch
Upper 19 Road
Spring Creek 1
Spring Creek 2
Spring Creek 3
Cross Mountain 1
Cross Mountain 2
Big Hole Gulch
Cox Ranch
Timberlake Creek 1
Timberlake Creek 2
Timberlake Creek 3
Big Hole Butte
Dressler Gulch
Sevenmi1e Reservoir
Pole Gulch
Cottonwood Gulch

No. of
Counts

1977
No. of
Cocks

Total
Birds

No. of
Counts

1978
No. of
Cocks

Total
Birds

2
2
2
1
2
2
1
1
3
1
1
3
2
1
3
2
3
2
4
2
3
3
2
2
2
2
2
2

64
55
12
101
26
19
13
22
6
34
23
52
33
24
31
16
25
13
34
16
55
43
21
34
27
14
37
8

67
57
12
107
38
19
14
24
8
78
23
54
53
44
31
18
25
13
38
16
55
43
69
34
28
15
76
32

3
4
4
3
4
2
3
4
4
2
4
3
4
4
4
4
5
3
4
4
4
4
4
4
4
4
4
3

55
69
8
153
29
18
42
37
22
57
37
20
30
24
42
12
31
5
37
20
53
66
20
37
25
7
87
38

60
70
8
179
29
18
43
41
28
59
43
20
31
27
44
12
31
5
51
20
53
71
53
38
37
7
87
38

-------------------------------------------------------------------------------------------------------------

I
I-'
-..j

~
I

�Table 2.

Peak counts of sage grouse, 45 leks, Moffat County 1977 and 1978 (continued).

.
Year
No. of
Counts

1978
No. of
Cocks

Total
Birds

12
38
24
47
36
39
48
33
78
92
30
80
8
37
10
40
62

4
4
4
4
4
5
5
4
4
4
3
3
2
4
4
4
4

3
32
24
42
36
18
l36
45
34
56
17
50
35
14
4
43
44

3
35
24
45
38
19
138
48
34
59
17
51
35
14
4
44
44

1,805

169

1,714

1,855

No. of
Counts

1977
No. of
Cocks

Total
Birds

West Timberlake Creek 1
Thornburg Well
Lone Tree
Scandinavian Gulch
Fourmile Creek 1
Fourmile Creek 2
Fourmile Creek 3
Juniper
Axial Basin
Dry Lake 2
Deception Creek
Round Bottom
Horse Gulch
Duffy Mountain
Morgan Gulch 1
Morgan Gulch 2
Boxelder Gulch

2
2
2
3
5
2
2
4
3
1
1
1
2
3
2
4
1

5
36
22
45
34
27
40
31
58
35
22
36
4
27
8
39
60

Total

98

1,387 .

Lek

Sununary:
Ave. no. counts/lek
Ave. no. cocks/lek
Ave. no. total birds/lek

3.8

2.2

38.1

30.8
40.1

41.2

I
I-'

-...J

V1
I

�-176Table 3. Comparison of sage grouse numbers, 15 leks 'counted both by D.W.M. 's
and project personnel, Moffat County, 1978.

D.W.M. District

Lek

High Counts
D.W.M.'s
Project W-37-R
Total
Total
Cocks
Birds
Cocks
Birds

Brown's Park

Gee Flats

52

52

55

60

(C. Brown)

Beaver Basin

133

150

153

179

Subtotals

185

202

208

239

Craig, NW

Cross Mountain 1

31

31

31

31

(C. Brown)

Cross Mountain 2

3

3

5

5

Grassie Reservoir

13

13

22

28

Lay Creek

62

62

57

59

Bord Gulch

22

22

37

43

Greasewood Gulch

30

30

20

20

Upper 19 Road

18

19

30

31

North Fork Big Gulch 2

40

40

37

41

Subtotals

219

220

239

258

Slater

Timberlake Creek 2

83

86

66

71

(C. Woodward)

Subtotals

83

86

66

71

Craig, SW

Juniper 1

33

33

45

48

(W. Roland)

Axial Basin

73

73

34

34

Dry Lake

55

55

56

59

Deception Creek

14

14

17

17

Subtotals

175

175

152

158

662

683

665

726

Totals
Summary:
Ave. no. cocks/lek
Ave. no. total birds/lek

44.1

44.3
45.5

48.4

�Table 4.

Status

Sage grouse 1ek status, Moffat County 1978.

Craig NW (C. Brown)

Active (1978)
North Fork Big Gulch 2
Grassie Reservoir
Lay Creek
Bord Gulch
Greasewood Gulch
Upper 19 Road
Spring Creek 1
Spring Creek 2
Spring Creek 3
Cross Mountain 1
Cross Mountain 2
Big Hole Gulch
Cox Ranch
Timberlake Creek 1
Timberlake Creek 2
Timberlake Creek 3
Big Hole Butte
Dressler Gulch
Sevenmi1e Reservoir
Pole Gulch
Cottonwood Gulch
West Timberlake Creek 1
(Skyline Road)
Thornburg Well
Lone Tree
Scandinavian Gulch
New (1978)
Thornburgh Gulch
Mud Spring Draw
West Timberlake Creek 2
Abandoned (1978) but Active (1977)

District Wildlife Manager
Craig SW (W. Roland)
Slater (C. Woodward)

Juniper 1
Axial Basin
Dry Lake 2
Deception Creek
Round Bottom
Horse Gulch
Duffy Mountain
Morgan Gulch 1
Morgan Gulch 2
Boxe1der Gulch 1

Fourmile Creek 1
Fourmile Creek 2
Fourmile Creek 3

Brown's Park (Vacant)

Gee Flats
Powder Wash Hill
Snake River West
Beaver Basin
Coffee Pot Spring
Summit Spring
Goodman Draw

I
I-'
'-J
'-J

I

Juniper 2
Morgan Gulch 3
Boxe1der Gulch 2
Temple Gulch

Cowboy Reservoir

East Sand Wash Road

Sand Wash Road

�-178Brood Counts
Seven sage grouse brood routes in Moffat and Western Routt counties were each
counted 4 times (2 in early A.M., 2 in late P.M.) in July 1978 (Table 5). A
total of 2,174 sage grouse (252 hens, 325 cocks, 665 young, and 932 unclassiComparative data for
fied) was counted in 762.0 miles (2.85 birds/mile).
1976, 1977 and 1978 are presented (Table 6).
The total of 1,214 sage grouse counted in the 14 morning counts is compared
with 960 in the 14 evening counts for approximately 21% fewer birds in the
evening counts.
One hundred and thirty-nine separate broods, averaging 4.8
young, were counted along these routes.
The young to adult ratio of 1.2:1
was lower than recorded both in 1976 (2.4:1) and 1977 (1.6:1). Numbers of
broods counted in 1978 (139) were higher than in 1976 (17) and 1977 (111).
Numbers of young per brood in 1978 (4.8) were
the same as in
1977 (4.8). Twenty-six additional sage grouse broods not counted along
routes during July 1978 averaged 3.9 young, bringing the total number of
broods observed in 1978 to 165. During 1978, 113 of 252 hens (45%) were
observed without broods along the routes.
The earliest brood observed in 1978 was on May 25, indicating some hens
brought off broods early. A late spring snowstorm in early May 1978, however,
caused many hens to recycle.
This resulted in various sizes of young ranging
from approximately 2 to almost 10 weeks old being observed during July 1978.
Capture

and Banding

Intensive efforts to capture and band sage grouse in the 1978 field period
were initiated on 13 July and continued through 29 August.
Age and sex data
of the banded sample are given in Table 7.
Trapping success was not uniform in Moffat County in 1~78 as birds were.captured
in only 2 different areas, Cold Spring Mountain (461 blrds) and Mud Sprlng Draw
(45 birds).
Of the trapping methods used, stationary cannon nets at t~e end
of wire drift fences were most successful (61 captures in 5 attempts, x = 12.2,
range 6-20). Lily-pad traps were next, with 202 captures in ~8 tra~ days,

ex

=7.2, maximum in 1 trap = 33), with bumper mounted cannon nets (197
captures in 37 attempts, x = 5.3, range 0-23), and spotlighting (105 captures
in 96.4 hours, x = 1.1 birds/hr) being somewhat less productive.
Spotlighting
was non-selective for age or sex of grouse captured, whereas, cannon nets
and lily pad traps were selective for juveniles.
Obviously, composition of
the sage grouse population was not reflected in the banded sample (83.2% chicks,
7.5% yearlings, 9.3% adults, 51.5% chick males, 65.8% yearling males, 51.1%
adult males).
It would appear that sex classes of chicks were captured in about
the same ratio they occurred in the population (see harvest data).
Prior to 1978, no sage grouse had been banded in Moffat County.
Consequently,
no recaptures of birds banded in prior years were recorded.
In addition to
the 506 sage grouse newly banded, 54 recaptures were recorded (up to 3 different
captures of some birds) with 5 additional birds being trap casualties (4 due
to broken necks, 1 due to suffocation).
Mortality associated with trapping
was low (0.9%, 5 7 565).

�Table 5.

Observations of sage grouse along brood routes, Moffat and Western Routt counties, 1978.

Name of
Route

Sage Grouse Counted
Adults
Hens Cocks Young Unc1ass.

Total

No. of
Broods

Young/
Brood

No. of
Brood1ess
Hens

Birds/
Mile

Date

Time

Miles

Minutes

Great Divide

7/8/78
7/11/78
7/16/78
7/20/78

AM
AM
PM
PM

24.6
23.6
24.0
23.9

127
155
142
140

11
4
11
7

4
8
5
5

30
20
55
28

24
78
38
0

69
110
109
40

6
4
11
7

5.00
5.00
5.00
4.00

5
0
0
0

2.80
4.66
4.54
1.67

Timberlake

7/6/78
7/l3/78
7/19/78
7/21/78

AM
AM
PM
PM

34.2
32.6
32.5
32.5

l30
230
158
145

2
14
l3
15

3
4

0
48
57
59

0
1
27
23

5
67
105
101

0
11
11
11

4.36
5.18
5.36

2
3
2
4

0.15
2.06
3.23
3.11

3
2
2
2

4.67
5.00
4.50
4.00

0
0
0
0

1.69
1.05
1.02
1.72

8

4

-...J

Big Hole

7/14/78
7/17/78
7/19/78
7/24/78

AM
AM
PM
PM

24.8
24.8
25.5
25.6

125
140
120
120

3
2
2
2

0
0
0
0

14
10
9
8

25
14
15
34

42
26
26
44

Spring Creek

7/7/78
7/l3/78
7/14/78
7/17/78

AM
AM
PM
PM

23.8
23.5
34.6
34.6

100
100
160
140

4
4
8
4

0
0
0
1

18
23
44
11

0
0
2
0

22
27
54
16

4
4
8
3

4.50
5.75
5.50
3.67

0
0
0
1

0.92
1.15
1.56
0.46

Powder Wash

7/10/78
7/14/78
7/22/78
7/23/78

AM

AM
PM
PM

27.9
30.0
30.5
33.4

l35
140
155
180

4
1
4
3

0
0
0
4

6
1
22
l3

0
27
0
0

10
29
26
20

1
1
4
3

6.00
1.00
5.50
4.33

3
0
0
0

0.36
0.97
0.85
0.60

7/12/78
7/18/78
7/17/78
7/18/78

AM
AM
PM
PM

21.7
21.7
22.3
22.1

195
185
160
195

56
54
17
5

46
91
58
71

47
63
53
19

102
328
100
94

251
536
228
189

9
14
12
5

5.22
4.50
4.42
3.80

47
40
5
0

11.57
24.70
10.22
8.55

Cold Spring

I
f-'

------------------------------------------------------------------------------------------------------------------------

1.0
I

�Table 5.

Observations of sage grouse along brood routes, Moffat and western Routt counties, 1978 (continued).

Name of
Route
E1khead

Sage Grouse Counted
Adults
Hens Cocks Young Unc1ass.

Total

No. of
Broods

Young/
Brood
7.00

Date

Time

Miles

Minutes

7/11/78
7/21/78
7/13/78
7/25/78

AM
AM
PM
PM

27.5
27.5
24.9
27.4

115
110
135
120

1
1
0
0

0
11
2
0

7
0
0
0

O'
0
0
0

8
12
2
0

1
0
0
0

762.0

4,057

252

325

665

932

2,174

139

Totals

Birds/
Mile

0
1
0
0

0.29
0.44
0.08
0.00

-

-

113
2.85

4.8

Averages

Table 6.

No. of
Brood1ess
Hens

I
f-'
00
0
I

Observations of sage grouse along brood routes, Moffat County area, 1976-1978.

Year

Period
of
Counts

1976

Sage Grouse Counted
Young

Unc1ass.

Total

No. of
Broods

Young/
Brood

Brood1ess
Hens

Birds/
Mile

16

90

436

564

17

5.3

5

1.37

151

179

532

288

1,150

111

4.8

40

1.27

252

325

665

932

2,174

139

4. 8

113

2.85

Time

No. of
Counts

No. of
Routes

No. of
Miles

Adults
Hens Cocks

8/3-8/30

AM

16

10

412.6

22

1977

7/12-8/11

AM
PM

23
10

11

905.1

1978

7/6-7/25

AM
PM

14
14

7

762.0

�-181Table 7. Sex and age composition
Moffat County, Colorado 1978.

of newly banded

sage grouse by month,

Month

Chick

1+

Males
2+

Total

Chick

1+

Females
2+

Total

July

67

6

8

81

80

8

3

91

August

150

19

16

185

124

5

20

149

Totals

217

25

24

266

204

13

23

240

Hunting

Season Data Collection

The sage grouse season in Moffat County in 1978 opened at sunrise on 9
September and closed at sunset on 17 September, 2 days longer than in 1977.
Bag and possession limits were 3 daily and 6 in possession (in the aggregate
with sharp-tailed grouse), the same as in 1977. Check stations were operated
from about 0800 to 1900 hrs MDT on September 9, 10, 16 and 17 at Cedar Mountain
and Dinosaur and about the same hours on September 9, 10 and 17 at Maybell
on Colorado Highway 318. In addition, volunteer wing collection barrels were
placed at Axial, Cedar Mountain (when the check station was closed), Deception
Creek, Dinosaur (when the check station was closed), Elkhead Reservoir,
Juniper, Lay Creek, Lower Big Gulch, Maybell at 318 (when the check station
was closed), Maybell City Park, Moffat County Road #2, Moffat County Road #3,
Moffat County Road #4, Moffat County Road #101, Wolf Creek Road, Hayden and
the Slater Grouse Camps.
Origin and Distribution

of Wings

Number

from the wing barrels

of wings received

Axial
Cedar Mountain
Deception Creek
Dinosaur
Elkhead Reservoir
Juniper
Lay Creek
Lower Big Gulch
Maybell at 318
Maybell City Park
Moffat County Road #2
Moffat County Road #3
Moffat County Road #4
Moffat County Road #101
Wolf Creek Road
Slater Grouse Camps
Hayden

were:

Sage Grouse

Blue Grouse

17
160
20
93
9
20
84

2

Sharp-tailed

2

3
1

7

58
54
44
134
83
99
21
69
34

9

1

4
7

31

23

Grouse

�-182-

A total of 1;312 sage grouse wings was obtained at the 3 check stations
(Cedar Mountain - 558 sage grouse, Dinosaur - 417 sage grouse, Maybell - 337
sage grouse, 17 blue grouse), while 98 sage grouse wings were received from
miscellaneous sources (Middle Park check station - 55, North Park check
stations - 12, brought in to Craig office - 14, brought in to Fort Collins
office - 19). In all, 2,418 sage grouse wings from Moffat (2,373) and western
Routt counties (45) were received.
Time of harvest was identifiable

for wings from 2,326 sage grouse

(Table 8).

Table 8. Time distribution of sage grouse wings received, Moffat and western
Routt counties, Colorado 1978.

Time Period

Number

Percent

September

9-10

1,745

75.0

September

11-15

224

9.6

September

16-17

357

15.4

2,326

100.0

Total

Check Stations
During the 4 days of check station operation (3 days at Maybell), 770 hunters
with 1,490 sage grouse (1.9 birds per hunter) were checked.
These hunters
reported observing 15,295 sage grouse.
Some duplications are undoubtedly
present in the 15,295 birds observed.
Hunter efficiency was low at 9.7%
(1,490 birds harvested + 15,295 birds observed) and 122 birds (7.6% of those
retrieved + those lost) were reported crippled and lost (Table 9).
It is readily apparent that differences occurred in hunter pressure, birds
seen, hunter efficiency and birds per hunter in the 3 areas sampled by check
stations.
Hunter efficiency was lowest and birds per hunter were highest in
the area where hunters observed the highest number of birds (primarily Cold
Spring Mountain).
It is also apparent that where hunter pressure was relatively
high birds per hunter was lowest, suggesting that sage grouse in these areas
were more difficult to approach and harvest.
Small game units in Moffat County were subdivided into harvest zones to
examine hunter pressure and harvest by area (Fig. 2). Data available from
check stations illustrate that hunter pressure and harvest were not uniformly
distributed in Moffat County in 1978 (Table 10).

�-183-

Table 9.

Sage grouse harvest

statistics,

1/
1978.-

Hunter
Efficiency
%

Crippling
Loss

Birds
per
Hunter

651

12.7

8.1

1.4

3,474

435

12.5

6.9

2.2

112

6,707

404

6.0

7.6

3.6

770

15,295

1,490

9.7

7.6

1.9

Only those statistics

collected

Check Stations

No.
Hunters
Checked

No.
Birds
Observed

Cedar Mountain

459

5,114

Dinosaur

199

Maybe11~/

All Stations

1:./

Moffat County, Colorado

l/ No check on 16 September.
9, 10, 16 and 17 September.

Table 10. Hunter pressure,
Moffat County 1978. 1:./

Statistic

No.
Birds
Harvested

at check stations.

Data from Cedar Mountain and Dinosaur represent
Data from Maybell represent 9, 10 and 17 September.

harvest

and hunter

success by harvest

zone,

16A

l6B

Harvest
l6C

Zone
l8A

l8B

20A

Total hunters

(%)

22.7

37.5

2.5

9.0

2.3

26.0

Total harvest

(%)

16.4

28.5

3.0

22.0

0.9

29.2

1.4

1.5

2.3

4.8

0.8

2.2

Birds/hunter

1:./

Check station data only.

Check station data indicate that while about 37% of the hunters hunted in
zone l6B and harvested about 28% of the birds bagged, smaller percentages of
hunters in zones 20A (26%) and l8A (9%) harvested significant portions of th~
birds bagged (29.2 and 22.0%, respectively).
Obviously, the Cold Spring
Mountain and Blue Mountain areas were the most productive in terms of hunter
success in 1978.
Most hunters contacted at check stations were asked whether or not they
normally hunted sage grouse in Moffat County.
Of the 727 hunters responding,
502 (69.1%) reported they normally hunted sage grouse in Moffat County, 81

�.

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Harvest zones, Moffat County, Colorado.

==:='C" ..J\"~"

ZONES

I

�-185-

(11.1%) normally hunted sage grouse elsewhere, while 144 (19.8%) were first
time sage grouse hunters (Table 11). These data are not markedly different
from those gathered in North Park (Braun 1978).

Table 11. Previous sage grouse hunting
Moffat County, Colorado 1978.

experience

of sage grouse hunters,

First Time Sage
Grouse Hunter

Normally Hunt
In Moffat County
%
No.

Normally Hunt
Elsewhere
No.
%

~.

%

502

81

11.1

144

19.8

69.1

Origin of 347 hunter parties was ascertained at check stations in 1978. Of
this total, 129 (37.2%) were from Moffat County, with 46 (13.3%) from Mesa
and 41 (11.8%) from Rio Blanco counties.
Counties in the Denver metropolitan
area (Jefferson - 9.2%, Adams - 5.2%, Arapahoe - 3.2%, Denver - 3.2%, and
Douglas - 0.6%) contributed 74 parties or 21.3% of the total. Other counties
contributing over 2% were
Boulder (3.5%), and Garfield (3.2%). All other
counties (11 different) and states (Wyoming and Utah) each contributed less
than 2% to the total and collectively totaled 34 parties (9.8%).
Wing Analysis
Age and Sex Composition.--All wings received were usable.
Moffat County
was subdivided into 5 areas for wing analysis.
These areas and corresponding
harvest zones (Fig. 2) were: 1) Eastern Moffat and Western Routt counties
(east of Colorado 789 to the Hayden-Slater Road, north of U. S. 40) including
wing collection barrels at Elkhead Reservoir, Moffat County Roads #'s 2 and
101, Slater Grouse Camps and Hayden, and wings from this area collected at
the Middle Park check station (harvest zone l4A). All but 45 wings were from
within Moffat County.
2) North Central Moffat County excluding data from the
Maybell Check Station and wing barrel at Colorado 318 and U. S. 40. This area
was west of Colorado 789, to the Little Snake River north of U. S. 40 (harvest
zones l6A and B). Included was the Cedar Mountain check station and wing
barrel, wing barrels at Lay Creek, Lower Big Gulch, Maybell City Park, Moffat
County Roads #'s 3 and 4 and miscellaneous collections.
3) The area of Cold
Spring Mountain which was sampled primarily by the Maybell check station and
wing barrel (harvest zone l8A, B and C). Some overlap occurred with this subunit
and North Central Moffat County. 4) Dinosaur.
This subunit contained all of
Blue Mountain north of U. S. 40 and included the Dinosaur check station and
wing barrel and the wing barrel on the Wolf Creek Road (harvest zone 20A).
5) Moffat County South. All area in Moffat County south of U. S. 40 was
included in this subunit including wing barrels at Axial, Deception Creek, and
Juniper (harvest zones l6C, 20B and 26A) (Table 12).

�Table 12.

Age composition of the sage grouse harvest, Moffat County, 11 Colorado, 1978.

No.

Area

11

Young
Percent

Yearlin~
Percent
No.

No.

Adults
Percent

Total

Percent of
Total Sample

Moffat County East-

207

77.8

23

8.7

36

13.5

266

11.0

North Central

839

71.8

58

5.0

272

23.2

1,169

48.3

Cold Spring

246

62.3

66

16.7

83

21.0

395

16.3

Dinosaur

330

62.1

90

17.0

111

20.9

531

22.0

Moffat County South

40

70.2

3

5.3

14

24.5

57

2.4
I
I-'

Totals
Average

68.7

-11 Including Western Routt County (45 wings).

2,418

516

240

1,662

9.9

00
0'1
I

21.4

100.0

�-187-

It is obvious from data in Table 12 that harvest was not uniform throughout
Moffat County.
Almost no harvest (2.4% of the total) occurred south of U. S.
40, even though apparently good populations exist in this area.
It is also
apparent that production at Blue Mountain and Cold Spring Mountain, while
good in 1978, was less than in other areas. However, percent yearlings in
both of these areas was average (16.7, 17.0%) in 1978, suggesting good
production in 1977 and survival of those young to 1978. All other areas
had less than 10% yearlings in the harvest sample suggesting poor production
in 1977 and/or survival of the 1977 cohort to 1978. Comparable data for
1976-1978 are presented in Table 13.
From the data presented (Table 13) it would appear that production of young
is consistently good each year at Blue Mountain (Dinosaur) and Cold Spring
Mountain.
It is possible that production in 1975 at Cold Spring Mountain was
either poor or survival of the yearling class to 1976 was poor.
Production
at the other areas has varied with 1977 being the poorest year.
It is probable
that sample sizes of wings have been inadequate in both 1976 and 1977 in the
Moffat County East (54, 91) and South (67, 31) areas.
Age and sex composition of the harvest for Moffat County during
varied (Table 14). These data suggest the following:

1976-1978

1.

The sex ratio at hatching approximates 1:1 with possibly some differential
survival favoring females occurring before chicks are 3-4 months of age.

2.

Differential survival
being most pronounced

3.

There are 2 hens to every male in the spring breeding population (3 year
average = 68.7% females, 31.3% males of the total adults and yearlings).

4.

Production
production

5.

Production in 1975 must have been average
yearlings in the 1976 harvest.

favoring females occurs in the older age classes;
in adults.

of young in 1976 and 1978 was good to excellent
in 1977 being poor to average.

with

to good based on percentage

of

Turnover.--Estimated
turnover (from wing analysis) of adult male and
female sage grouse in Moffat County was 43.3 and 33.2%, respectively, in
the 1976-1978 period (Tables 15 and 16). Estimated survival of adult males
was 56.7%, while for females it was 66.8%. These estimates are based on the
assumption that in a stable population, percent yearlings must equal annual
loss of adults.
In an increasing population, percent yearlings must more
than replace adult loss and/or adult survival must be better than that
estimated.
Percent yearlings (Table 14) has not been good (except in 1977).
Consequently, there should have been no increase in the 1978 breeding population unless survival of the 1976 cohort of chicks to 1978 was excellent.
Survival of males (56.7 vs 50.8%) is higher in Moffat County than in North
Park.
Survival of females in the 2 areas is essentially identical (33.2 vs
34.5%) (Braun 1978).

�Table 13.

Age composition of the sage grouse harvest, Moffat county1~ Colorado 1976-1978.

l1

1976

Yearlings
1977

1978

1976

Adult
1977

1978

77 .8

33.3

19.8

8.7

20.4

40.7

13.5

34.0

71.8

14.5

29.0

5.0

21.4

37.0

23.2

73.1

59.8

62.3

7.7

17.6

16.7

19.2

22.5

21.0

Dinosaur

61.6

60.4

62.1

15.2

16.0

17.0

23.2

23.6

20.9

Moffat County South

71.6

25.8

70.1

13.4

29.0

5.3

14.9

45.2

24.5

1976

Young
1977

1978

11
Moffat County East-

46.3

39.6

North Cen tra1

64.1

Cold Spring

Area

-

Average

63.8

-11 Including western Routt County.

II In percent.

42.6

68.7

15.4

23.9

9.9

20.8

33.5

21.4

I
f-'

co
co
I

�Table 14.

Age and sex composition of the sage grouse harvest, Moffat and Western Routt counties, Colorado 1976-1978.

Year

Males
No.
%

Immatures
Females
No.
%

Total
No.
%

Males
No.
%

Yearlings
Females
No.
%

Total
No.
%

Males
No.
%

Adults
Females
No.
%

Total
No.
%

1976

203

42.6

273

57.4

476

63.6

41

35.7

74

64.3

115

15.4

25

15.9

l32

84.1

157

21.0

748

1977

121

39.3

187

60.7

308

43.0

77

45.8

91

54.2

168

23.4

89

36.9

152

63.1

241

33.6

717

1978

825

49.6

837

50.4

1,662

68.7

77

32.1

163

67.9

240

9.9

141

27.3

375

72.7

516

21.4

2,418

3 Year
Average

Total
Sample

I
I-"
00

'"
I

47.0

53.0

63.0

37.3

62.7

l3.5

27.9

72.1

23.5

�-190-

Table 15. Estimated turnover
Colorado 1976-1978. 1/

of adult male sage grouse, Moffat

No.

Yearlings
Percent

County,

Year

No.

Adults
Percent

1976

25

37.9

41

62.1

66

1977

89

53.6

77

46.4

166

1978

141

64.7

77

35.3

218

1/ Data

43.3

56.7

3 Year Average

from wing collections

Table 16.

Estimated

Colorado

1976-1978. 1/

turnover

Totals

only.

of adult female sage grouse, Moffat

Adults
Percent

County,

No.

Yearlings
Percent

Totals

Year

No.

1976

132

64.1

74

35.9

206

1977

152

62.6

91

37.4

243

1978

375

69.7

163

30.3

538

3 Year Average

1/ Data

66.8

from wing collections

33.2

only.

Nesting Success and Production.--Molt
of adult and yearling sage grouse
was examined to estimate nesting success (Table 17). Differences in estimated
nesting success closely parallel differences in percentages of young in the
harvest (Table 12). Considering all areas within Moffat County, about onehalf (50.3%) of the yearling
and two-thirds (65.3%) of the adult females were
successful nesters in 1978. About 61% (60.8) of all hens were successful.
These data are higher than the estimated overall nesting success of 30.0% in
1977, and 48.3% estimated in 1976. Production of young in 1978 was excellent
(Table 17) with only Blue Mountain and Cold Spring Mountain having less than
3 chicks/hen in the harvest.
The high chick/hen ratio (5.0) for eastern
Moffat County is probably an artifact of either hunter and/or sage grouse
behavior.
Chick/hen ratios in 1978 were higher than in either 1976 (2.3) or
1977 (1.2) (Table 18).

�Table 17.

Estimated sage grouse nesting success and production, Moffat County, Colorado 1978.

Area

Yearlings
No. Successful

Moffat County East

13/

North Central

25/41

17

%

Adults
No. Successful

%

All Hens
No. Successful

%

Chicks/Hen

Chicks/
Successful Hen

76.5

17/24

70.6

30/41

73.2

5.0

6.9

61.0

151/204

74.0

176/245

71.8

3.4

4.8

Cold Spring

18/

45.0

32/59

54.2

50/

50.5

2.5

4.9

Dinosaur

25/63

39.7

38/79

48.1

63/142

44.4

2.3

5.2

Moffat County South

1/2

50.0

7/9

77.8

8/11

72.7

3.6

5.0

Totals

82/162

60.8

3.1

5.1

Average

40

245/

327/

375

50.3

99

538

65.3

I
f-'
I.D
f-'
I

�Table 18.

Estimated sage grouse nesting success and production, Moffat County, Colorado 1976-1978.

Yearlings
1978
1977

1976

Adults
1977

1978

1976

All Hens
1978
1977

1976

Chicks/Hen
1978
1977

Area

1976
28.6

28.6

76.5

50.0

58.3

70.6

40.0

47.4

73.2

1.7

0.9

5.0

Moffat County East

36.2

6.4

61.0

51.9

25.6

74.0

46.1

18.4

71.8

2.3

0.9

3.4

North Central

25.0

45.0

80.0

80.0

54.2

66.7

59.4

50.5

1.9

0.0

3.2

2.5

Cold Spring

20.0

28.6

39.7

50.0

35.7

48.1

40.6

33.3

44.4

2.2

3.0

Dinosaur

62.5

14.3

50.0

80.0

0.0

77.8

72.2

10.0

72.7

2.7

0.8

3.6

Moffat County South

14.9

50.3

55.7

39.6

65.3

48.3

30.0

2.3

1.2

35.1

60.8

3.1

Average

I

2.3

r-'
1.0
N

I

�-193-

Hatching Dates
Ages and hatching dates were calculated for 1,662 chick sage grouse
harvested in Moffat County in 1978 (Table 19). Hatching dates varied from
4-10 May to 20-26 July, with an obvious peak between 1-4 June. Hatching was
spread over the longest period in the eastern, north central and Cold Spring
Mountain areas and was shortest on Blue Mountain and south of U.S. 40.
Renesting by some hens that lost their initial clutch probably occurred in
all areas with the Cold Spring Mountain area having an obvious second peak
of hatching during the 29 June-12 July period.
Data presented in Table 19
do not argue strongly for analyzing chick hatch dates by area. Despite area,
elevation, differing weather conditions, etc., most chicks (~70%) hatched
within a 30 day period from late May to late June. Only slight 7-10 day
shifts in timing of peak hatch and in importance of renesting can be
expected from year to year. Once range and normal peak of hatching are
documented, annual calculation of hatching dates would appear to have little
value for management.
Ovarian Analysis
Ovaries were collected from 64 hunter harvested female sage grouse in 1978
and were analyzed following procedures described by Meyer et al. (1947),
Kabat et al. (1948), and Buss et al. (1951). An additional 77-ovaries were
classified in situ at check stations.
Data presented (Table 20) indicate
that a high percentage of both yearlings and adults ovulated in 1978.
Comparable data for years prior to 1978 are not available.
Hunter Questionnaire
A total of 2 006 permits
was issued for hunting sage grouse in Moffat County
in 1978. Ba;ed on 770 hunters contacted at check stations, only 65 (8.4%) of
those hunting did not have the required permit. .If this percentage reflects
the hunter universe in 1978. then it is possible that ther~ were 2,174 sage
grouse hunters (2,006 + 168) in Moffat County in 1978. However, when it is
realized that of the 2,006 permittees, only 1,502 (Table 21) hunted, then it
is reasonable to conclude that there were no more than 1,628 sage grouse
hunters in Moffat County in 1978 (1,502 + 126).
Origin of permittees was primarily from Moffat (44.0%), Rio Blanco (9.5%),
and Mesa (8.9%) counties.
Counties in the Denver metropolitan area (Adams,
Arapahoe, Denver, Douglas, Jefferson) each contributed less than 8% of the
total permittees but collectively comprised 19.7% of the total. Other
counties providing more than 1% of the total were Routt (4.0%), Boulder
(2.9%), Garfield (2.2%), and Larimer (1.3%). All other counties and states
each comprised 1.0% or less of the total permittees.
These data are not
markedly different from those collected at check stations.
Questionnaires were sent to all permittees immediately following the sage
grouse season in Moffat County (22 September).
Responses were received from
1,213 permittees.
On 16 October follow up letters and questionnaires were sent
~~ all non-respondents and 353 additional responses were received.
In all,
1,566 permittees (78.1%) responded (Table 21). Fifty-eight questionnaires
(2.9%) were undeliverable.
Mean values calculated for permittees responding

�1

Table 19.

Estimated sage grouse hatching dates, Moffat County, Colorado 1978.

Females

Males
East

N.C.

Cold
Spring

/

Dinosaur

N.C.

Total

East

0.4

0.9

6.2

0.8

2.8

2.5

South

Cold
Spring

May 4-10
May 11-17

0.7

0.7
1.8

12.5

5.1

11.1

9.2

3.8

7.4

2.1

27.1

12.5

10.7

23.2

24.3

6.7

17.4

4.4
29.8

28.3

31.2

32.0

18.5

24.8

23.8

May 18-24
May 25-31
June 1-7
June 8-14
June 15-21
June 22-28
June 29-Ju1y 5
July 6-12
July 13-19

7.1
7.1

34.5

33.3

11.1
0.0
1.0

Total Sample

99

7.3

6.7

16.7

18.3

22.0

33.3

23.5

21. 3

23.9

26.7

25.6

8.3

12.0

5.3

3.8

18.9

0.0

9.9

11.5

8.5

6.3

4.6

4.8

7.6

12.2

6.4

8.2

6.6

6.3

4.2

9.6

3.4

8.5

5.4

16.3

1.9

12.4

4.5

8.2

4.2

3.0

2.8

1.3

12.4

4.2

3.9

2.0

3.0

6.1

8.5

0.6

0.9

0.5

2.8

0.5

1.4

0.7

0.1

141

166

16

825

108

I
•.....
\0

23.8

14.5

403

-1/ In percent of total.

20.8

25.0

0.2

July 20-26

1.9

15.7

7.3

5.1

8.3

18.4

8.9

18.1

Total

19.4

16.6

16.2

South

0.1

2.1
1.0

Dinosaur

436

105

164

24

837

~
I

�-195-

Table 20.

Age Class

Sage grouse ovarian

analysis,

Moffat

Number Ovulatory/Examined

County,

Colorado

1978.

Percent

Ovulatory

Yearling

98.2

Adult

98.8

Total
Percent

139/

141
98.6

to the follow up letter were used to project for the 440 (21;9%) nonrespondents.
This undoubtedly somewhat inflated total estimates as the
available evidence indicates that non-respondents
are even less successful
than respondents to the follow up letter (5 of 8 questionnaires received
after the cut off date indicated that the permittees did not hunt).
Data presented indicate that about 75% of the permittees "hunted and harvested
(including crippling loss) an estimated 5,185 sage grouse in Moffat County.
About 75% of all hunters were successful and harvested 4.2 grouse each.
These values are considerably higher than those reported for North Park
(Braun 1978).
Time period of hunting was received for 1,586 hunts which resulted in 3,891
grouse being bagged.
Based on this sample, 61.3% of the hunts resulted in
66.1% of the total harvest during the opening weekend.
The 5 week days of
the season attracted 20.6% of the hunters and 19.0% of the harvest, while
18.0% of the hunts resulted in 14.8% of the grouse being bagged the 2nd
weekend.
A total of 474 of the 1,214 hunters responding (39.0%) reported achieving
the bag limit at least 1 day during the season.
Of this sample, 220 hunters
achieved the bag limit on 1 day, 219 did on 2 days, 21 did on 3 days, 10 did
on 4 days, while 2 each achieved the bag limit on 5 and 6 days.
Obviously,
the longer seasons, while resulting in increased hunter opportunity, would
not necessarily result in increased levels of harvest.
The questionnaire was designed to examine harvest and hunter
activity by
Small Game Units and harvest zones (Fig. 2). Data presented (Table 22)
indicate that 53% of the hunters were in Unit 16 and harvested about 48% of
the total birds taken in Moffat County.
The next most important units were
18, 20, and 14. Few hunters hunted in Unit 26 in Moffat County.

�-196-

Table 21.

Moffat County sage grouse hunter questionnaire data, 1978.

N2
No. in sample

1,566

353

1,213

78.1

Projected
For

Projected
For

440

2,006

21.9

100.0

Percent of total permittees

60.5

17.6

No. of hunters

983

231

Percent of total hunters

81.0

65.4

77 .5

65.4

74.9

No. of non-hunters

230

122

352

152

504

Percent of non-hunters

19.0

34.6

22.5

34.6

25.1

No. of successful hunters

775

157

932

196

Percent successful hunters

78.8

68.0

76.8

89.0

75.1

Percent success of permittees

63.9

44.5

59.5

44.5

56.2

No. of hunter days
Days/hunter
No. of grouse bagged

2,030
2.1
3,239

497
2.2
652

Grouse/hunter

3.3

2.8

Grouse/successful hunter

4.2

4.2

Grouse/permittee

2.7

1.8

No. birds lost
Birds lost/hunter
Total harvest
Percent crippling loss

293
0.3
3,532
8.3

109
0.5
761
14.3

1,502

288

1,214

2,527
2.1
3,891
3.2

2.5
402
0.3
4,293
9.4

634
2.2
806

1,128

3,161
2.1
4,697

2.8

3.1

4.2

4.2

1.8

2.3

86
0.3
892
9.6

488
0.3
5,185
9.4

�-197-

Table 22. Hunter activity and harvest
harvest zones, Moffat County, Colorado

by Small Game Management
1978.

Units and

Unit
(Harvest Zone)

No. of
Hunters

Percent
of Total

No. of Birds
Harvested

Percent of
Total

l4(A)

l38

10.8

380

9.8

16

679

53.0

1,865

47.9

(A)

227

17.7

604

15.5

(B)

356

27.8

1,051

27.0

(C)

96

7.5

210

5.4

220

17.2

802

20.6

(A)

127

9.9

517

l3.3

(B)

80

6.2

231

5.9

(C)

13

1.0

54

1.4

169

13.2

655

16.8

(A)

153

11. 9

603

15.5

(B)

16

1.2

52

1.3

26(A)

33

2.6

80

2.1

Unknown

43

3.4

109

2.8

Totals

1 ,2821:./

100.0

3,891

100.0

18

20

1/
- Total does not equal 1,214 as some hunters

hunted

in more than

1 zone.

Vulnerability
Forty-eight band recoveries were reported during and after the 1978 hunting
season, all from bandings in 1978. Of this total, 40 were reported on the
questionnaire survey.
Other sources were check stations and mail reporting.
Of the 8 not reported on the questionnaire survey, 6 were obtained at check
stations, 1 personally, and 1 from a wing barrel.
Many bands were reported
2 or 3 times because of the various methods of collection (check stations,
questionnaires and reports by mail).

�-198-

Nineteen bands (1- = 16, 1+ = 1, 2+ = 2) were from females, while 29
(1- = 20, 1+ = 4, 2+ = 5) were from males (Table 23). Data presented
indicate that males, principally adults and yearlings, are slightly more
vulnerable to hunting than females.
Essentially no differences existed in
vulnerability between female age classes and only slight differences existed
between chick females and chick males.
These data do not suggest that hunters
are selective towards sma lLe r birds.
On the contrary, selection would seem
to be towards larger birds.
However, differences in vulnerability between
males and females are probably related to differences in flushing distances,
size of target and flight speeds.
Also, it should be noted that only small
samples of adults and yearlings were banded in 1978. Present data do not
suggest that chicks are more vulnerable to hunting than older age classes.

Table 23.

Sage grouse banding

and recovery

data, Moffat County,

Colorado

No.
Banded

No.
Recovered

Harvest

Chicks

204

16

7.8

Yearlings

13

1

7.7

Adults

23

2

8.7

Subtotals

240

19

7.9

Chicks

217

20

9.2

Yearlings

25

4

16.0

Adults

24

5

20.8

Subtotals

266

29

10.9

506

48

9.5

Age and
Sex Class

Rate

(%)

Females

Males

Totals

1978.

�-199LITERATURE

CITED

Beck, T.D.I., R. B. Gill, and C. E. Braun.
1975.
Sex and age determination
of sage grouse from wing characteristics.
Game Inf. Leaflet No. 49
(Revised).
Colorado. Div. Wildl.
4 pp.
Braun, C. E. 1978.
Evaluation of the effects of changes in hunting regulations on sage grouse populations.
Colorado Div. Wildlife, Prog.
Rept., Fed. Aid Proj. W-37-R, Work Plan 3, Job 9. pp. 1-30.
---

, and T.D.I. Beck.
1976.
Effects of sagebrush control on distribution
and abundance of sage grouse.
Colorado Div. Wildlife, Final Rept.,
Fed. Aid Proj. W-37-R, Work Plan 3, Job 8a. pp. 21-84.

Buss,

I. 0., R. K. Meyer, and C. Kabat.
studies based on ovulated follicle

Eng, R. L.
wings.

1951.
Wisconsin pheasant reproduction
technique.
J. Wildl. Manage. 15:32-46.

1955. A method for obtaining sage grouse
J. Wildl. Manage. 19:267-272.

Gullion, G. W. 1965.
Improvements in methods
grouse.
J. Wildl. Manage. 29:109-116.
Hoffman, R. W., and C. E. Braun.
station.
Game Inf. Leaflet

age and sex ratios

for trapping

and marking

1975.
1975. A volunteer wing
No. 101. Colo. Div. Wildlife.

from

ruffed

collection
3 pp.

Kabat, C., I. O. Buss, and R. K. Meyer.
1948.
The use of ovulated follicles
in determining eggs laid by the ring-necked pheasant.
J. Wildl. Manage.
12:399-416.
Lacher, J. R., and D. D. Lacher.
Manage. 28:595-597.

1964.

A mobile

cannon net trap.

J. Wildl.

Meyer, R. K., C. Kabat, and I. O. Buss.
1947.
Early involutionary
changes
in the post-ovulatory
follicles in the ring-necked pheasant.
J. Wildl.
Manage. 11:43-49.
Pyrah, D. G.
Wildlife
71 pp.

1963.
Sage grouse investigations.
Restoration Div., Job Compl. Rept.,

Rogers, G. E. 1964.
Sage grouse investigations
Game, Fish and Parks, Tech. Publ. No. 16.

Prepared

by

~£~
Clait E. Braun
Wildlife Researcher

Donald M. Hoffman
Wildlife Researcher

4W"

C

C

Idaho Fish and Game Dept.,
Fed. Aid Proj. W-125-R.

in Colorado.
132 pp.

Colo. Dept.

��-201-

JOB PROGRESS

State of

October

1979

REPORT

COLORADO
--------~~~~~-----------

Proj ect No.
Work Plan 1,0.

9

Job Title

Dynamics

Population

Period Covered:
Personnel:

Game Bird Survey

H-37-R-32

Job No.
and Habitat Relationships

~5

_

of Blup Grouse

April 1, 1978 to March 31, 1979

L. Alexander, D. Benson, J. Brache, C. Braun, J. Claassen, R.
Clippinger, J. Corey, K. Duncan, H. Funk, J. Gerrans, K. Giesen,
W. Heicher, D. Hoart, D. Hoffman, R. Hoffman, R. Oakleaf, L.
Rottman, M. Sellitto, B. Sigler, M. Smith, S. Steinert, and W.
Woodward.
ABSTRACT

Investigations concerning the effects of hunting on blue grouse (Dendragapus
obscurus) populations, stability of breeding population levels, and habitat
relationships of blue grouse were initiated in 1975 and continued in 1978
on two areas in northwestern Colorado.
The breeding population at Eiby Creek
has remained essentially stable at about 10 hectares/territorial
male since
1976, whereas, the Green Mountain population has increased slightly from 9.2
hectares/territorial
male (1975) to 7.7 hectares/territorial
male (1978).
The sex ratio in the breeding population approximated 1:1. Evaluation of 8
nests located since 1975 indicated an average clutch size of 6.0 eggs.
Egg
loss reduced the hatch by some 32 percent with 75 percent of the nests
hatched successfully.
Peak of hatch varied by almost 3 weeks over the 4 years
of study, but mainly occurred in mid- to late June. Estimated nesting success
from examination of wings was 72 percent for Middle Park and 81 percent for
the Eagle area. Adults exhibited a more constant level of nesting success
from year to year and an overall higher success than yearlings.
Sixty-two
broods were observed in 1978 with a composite average of 4.9 (Green Mountain)
and 5.1 chicks per brood (Eiby Creek).
Average annual summer mortality of
chicks was 45 percent for Green Mountain and only 28 percent for Eiby Creek.
Pooled data on breeding birds banded in 1977 and resighted in 1978 gave an
estimated mortality rate for the breeding population of 39 percent.
Production
and mortality estimates suggest there exists a surplus of birds in the fall
population and in most years this surplus exceeds 40 percent.
Only a small
portion (5%) of the fall population was removed by hunting, compared to what
could be safely harvested (23%) without adversely affecting the subsequent
spring population.
Another survey was conducted in 1978 in order to evaluate
the combined blue grouse-big game season. As in 1977, few big game hunters
(5.5%) actually took advantage of the longer season, but their general
response to the more liberal season was highly favorable (59.7%).
The
additional harvest resulting from the longer season amounted to about a 26
percent increase in the total kill. Vegetation maps were prepared for the
Green Mountain study area and characteristics of breeding, nesting and brood
areas were described.

��-203-

HABITAT

POPULATION DYNAMICS AND
RELATIONSHIPS OF BLUE GROUSE
Richard W. Hoffman

Blue grouse are the most widespread member of the upland game birds in Colo2
rado, occurring in varying densities over more than 51,800 km2 (20,000 mi )
of diverse habitats and terrain throughout the state (Rogers 1968).
Furthermore, blue grouse rank first in terms of population numbers and annual harvest
among resident grouse species.
Yet, Colorado has had a long history of
conservative seasons on blue grouse, in spite of its abundance and wide
distribution.
This conservatism has mainly resulted from: (1) lack of data
upon which to base management recommendations, and (2) the misconception that
hunting can be a major mortality factor on blue grouse.
Data presented in
this report concern these problems and represent results of population studies
initiated in 1975.
P. N. OBJECTIVE
Major objectives of this study are to (1) increase the harvest of blue grouse
in Colorado (double present harvest estimates) without harm to breeding populations in subsequent years, (2) to identify differences in breeding densities
due to differing habitats, and (3) to document the stability of breeding
densities over time.
SEGMENT OBJECTIVES
1.

Review literature concerning (a) techniques for marking, capturing, and
censusing grouse, (b) methods of aging and sexing blue grouse, and (c)
population dynamics, behavior, and habitat requirements of blue grouse,
and (d) effects of hunting on grouse populations.

2.

Trap and individually mark 50 birds (including adults,
juveniles) on each of the two selected study areas.

subadults,

3.

Estimate breeding densities on the study areas through
acoustical census and systematic search.

the use of

4.

Estimate nesting

5.

Vegetatively describe areas studied and correlate
with features of the environment.

6.

Manipulate

7.

Compile data, analyze results,

success and production

hunting

Reference
materials
discussed

on the study area.

season and obtain harvest

METHODS

and

blue grouse densities

data.

and prepare progress

reports.

AND MATERIALS

is made to Hoffman (1976) for a detailed discussion of methods
used in this study. Any deviations or additions to these are
in the Results and Discussion section.

and

�-204-

DESCRIPTION

OF STUDY AREAS

Research is presently being conducted on two selected study areas of differing
habitat types in northwestern Colorado.
These areas are described in detail
in previous reports (Hoffman 1976 and 1977) and are depicted in Figs. 1 and 2.
RESULTS AND DISCUSSION
Breeding

Survey

Timing
In 1978, grouse were observed on the breeding range at both study areas in
early April.
First arrivals were mostly males.
Few hens were present during
the first 2-3 weeks of April, thus display was much curtailed.
Displays were
limited to the predawn hours and probably associated with territorial establishment rather than sexual responsiveness.
Birds still spent most of their
time in the trees.
The frequency and intensity of displays increase about the third week of
April.
This increased intensity of display corresponded with an increase in
the number of hens observed on the study areas. Activities associated with
breeding and density of breeding grouse peaked during the first week of May.
Timing of breeding events in 1978 was similar to 1975 and 1976, and approximately one week later than in 1977.
Display by males was concentrated in the early morning period between 0430
and 0600 MST. Occasional display occurred after dusk, but such display was
sporadic and involved fewer birds. Males spent more time on the ground after
the hens arrived before flying to trees to roost and feed. Once the snow
cover disappeared (late April-early May), both males and females spent almost
all of their time on the ground, roosting only at night.
Males were effectively finished displaying by late June-early July, although
flutter flights were heard into late July-early August.
The actual time of
departure of males from the breeding areas was unknown.
Males were observed
on Green Mountain year round, but it was uncertain whether they were the same
males that utilized the area for breeding.
At Eiby Creek, few males were
found after the first week of August, except at higher elevations near the
coniferous forest.
Displays
Display of the male blue grouse consisted mainly of flutter flights, strutting
and hooting.
These displays have been described by several authors (Rogers
1968, Hjorth 1970, Blackford 1963, Harju 1974, and Stirling and Bendell 1970).
Flutter flight as used here is analogous to the flutter flight described by
Harju (1974) and included either (1) jumping into the air with wings beating
and turning a half circle before landing in full display, or (2) flight from
ground to tree, or tree to tree with loud crackling wing beats at the termination of the flight.
The full display or strutting position consisted of
the tail being raised to the vertical and fanned to a complete half circle;

�GREE

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Fig.

1.

Green Mountain

study

area.

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EBY CREEK AREA
COLORADO

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�-207-

wings extended slightly out and down so the wing tips dragged on the ground;
eye combs and gular sacs expanded and engorged with blood so they became
deep red; and the white rosette of cervical feathers around the gular sacs
extended until forming a complete circle.
Several variations of this display
were observed, especially when the bird was hooting.
During hooting the wings were often flat against the body.
The white cervical
feathers were seldom fully extended and the gular sacs were only partially
visible.
The tail was generally in the normal position and unspread, but
sometimes it was raised and not fanned, raised and fanned, or fanned partially
and not raised.
Hooting was the five note type characteristic of the dusky
race. Pulsations could be seen in the neck region when the bird was hooting.
Males selected open sites within their territories for performing their
displays and could almost always be located in the early morning at their
favorite display spot. However, their display activities were not confined
to these selected spots. When a hen was heard or seen, the male would
frequently strut or fly in her direction.
Males would sometimes display in
trees, but usually only if they were disturbed on the ground or aroused while
feeding or roosting in a tree.
Flutter-flights were the most commonly heard and observed type of display.
Flutter-flights, as noted by Caswell (1954) and Harju (1974), had a chain
reaction response that triggered flutter-flights from surrounding territorial
males.
It is believed for this reason, that one function of the flutter-flight
was to proclaim territoriality to other males.
Flutter-flights also served
as a long distance display to attract hens to the territory and advertise
the presence of a territorial male.
There were some mornings when flutterflights could be heard up to 500 meters away.
Hooting was considered a close up display designed to attract and stimulate
the hen. Whenever a male was observed hooting there was usually a hen nearby.
Males could be stimulated to hoot by the play-back of a tape recorded hen
call at distances less than 50 meters.
At distances greater than 50 meters
the males initially responded with a wing flutter.
Often times, the male
would move towards the source of the call in the strutting position stopping
occasionally to perform wing flutters.
As he approached closer, he stopped
wing fluttering and started hooting.
If disturbed, he would fly to a tree
and continue hooting.
Hooting was variable in terms of audibility, but
seldom could be heard over 30 meters away. As in this study, Harju (1974)
found no evidence to suggest that hooting served to proclaim territory and
intimidate intruders or neighboring males.
Communal display, described by Blackford (1958, 1963) and noted by Harju
(1974), was not observed on Green Mountain or at Eiby Creek.
There were
several instances when 5 or 6 males became concentrated in a small area
where several hens were present.
However, none of the males crossed over
territories of adjacent males in order to display collectively as a group.
Little active display was noted by hens other than cackle calls and wing
sounds.
There were undoubtedly more displays performed by hens but due to
their more secretive nature and greater occupied range, fewer hens than males
were encountered in the field. When disturbed they either sat motionless on

�-208-

the ground or in a tree, or else they flushed wildly out of sight.
Cackling
was the main call produced by hens which stimulated males to flutter or hoot
depending on how close they were to each other.
This call was referred to
as the "quaver cry" by Stirling and Bendell (1970) and was the chief call
performed by hens in Wyoming (Harju 1974). A call resembling the precopulatory "whinny" described by Stirling and Bendell (1970) was heard only
once and may be a less important part of dusky grouse display than the cackle
call.
Hens also produce a type of wing flutter when landing, which often initiated
a response from nearby males.
The wing flutter was not the same as that
performed by males.
Hens landed with a constant wing beat, while males
terminated their flight with a more distinct, louder flapping.
Territoriality

and Density

Blue grouse males on the study areas were territorial and returned to the
same territory year after year, though the boundary of the defended area
changed occasionally.
New territories were located in some years, but for
the most part, the same territories were occupied each year. Thus, the newly
recruited males were attracted to and filled vacant territories.
Some yearling
males were territorial, and only one adult male captured on Green Mountain
was not territorial.
All other adult males captured on the study areas were
territorial.
Even through territorial males were frequently observed in close proximity
to one another, it is believed territorial boundaries seldom overlapped.
There were known instances when a male was disturbed that he would fly into
another male's territory without being pursued by the resident male. However,
such behavior was usually observed during the early stages of territorial
establishment in mid-April.
Attempts to force a male off his territory in
late April were unsuccessful.
Territory size was determined for 9 banded males seen at least 5 times each
in 1977 and 1978 on Green Mountain.
Territory size ranged from 1.2 to 1.9
hectares, with a mean size of 1.6 hectares.
Harju (1974) reported a mean size
of 0.7 hectares (range 0.4-1.1 hectares) per territory in Wyoming.
Bendell
and Elliott (1970) found that sooty grouse occupied territories of 0.4 to
0.8 hectares in a dense population and 1.6 to 2.8 hectares in a sparse population. Martinka (1972) reported a mean territory size of 0.8 hectares
(range = 0.4 to 1.5 hectares) for blue grouse in Montana, while Boag (1966)
found that blue grouse in Alberta occupied territories of 0.22 to 0.92
hectares.
Total number of territorial males recorded during the peak of breeding activities was used as an index to population size. It was impossible to traverse
the entire area sufficiently intensively to determine the total population.
The problem was further compounded by the fact that non-territorial males and
females undergo rather extensive daily movements and were more difficult to
find to accurately census.
Since the study areas under investigation differed
in terms of habitat type, ecological density rather than total territorial
males counted was used for comparison of densities between study areas.
In

�-209-

the estimation of ecological density, any area on the breeding range where
grouse were observed was considered habitable.
For Eiby Creek and Green
Mountain the estimates were 302.5 (60.5% of total area) and 146.7 hectares
(80.9% of total area), respectively.
Table 1 presents the change in numbers of territorial males and ecological
densities over a 4 year period on Green Mountain and a 3 year period at Eiby
Creek.
Breeding surveys were not conducted at Eiby Creek until 1976.
Since
then, the population density has shown little fluctuation.
Similarly, Green
Mountain supported a stable population in 1975 and 1976, with slight increases
in 1977 and 1978. Ecological density of males appeared to be slightly
higher on Green Mountain than at Eiby Creek, however, the difference was nonsignificant (P&gt;.05)
.
Table 1. Numbers and ecological densities
Green Mountain and Eiby Creek, 1975-1978.

of territorial

male blue grouse,

Ecological Density(ha/Bird)
1975
1976
1977
1978

Study Area

Size
(ha)

No. Territorial Males
1975
1976
1977
1978

Green Mtn.

181.3

16

16

17

19

9.2

9.2

8.6

7.7

Eiby Creek

500

ND

29

29

30

ND

10.4

10.4

10.1

Number of hens on the breeding range could not be accurately ascertained
from the breeding survey.
It was possible to estimate the number of hens
on the study area just after most of the broods had hatched.
At this time
hens were very responsive to a tape recorded chick call and were easily
located.
These data are presented in Table 2, along with the total population
estimate.
The data are given from 1976 to 1978 for Green Mountain only.
Due to the smaller size and isolated nature of Green Mountain from surrounding
breeding populations, the area was easier to search and the hens observed
were probably resident birds.
Data from 1975 are excluded because less
time was spent searching for broods than in other years and both the field
crew and dog lacked experience in finding broods.
Table 2.

Population

estimate

of blue grouse on Green Mountain,

1976-1978.

Terr.

Males
Non-Terr.

Tota]

Sex Ratio
Males: Females

Total
Population

Year

Succ.

Females
Unsucc.

1976

13

4

17

16

3

19

1:1.1

33

1977

17

4

21

17

3

20

1:1.1

41

1978

15

5

20

19

4

23

1.2:1

43

Total

�-210-

Some hens and non-territorial males undoubtedly escaped detection in all
years, therefore the total population estimate was minimal.
The sex ratio
indicated by these estimates did not deviate significantly from 1:1 (P&gt;.05).
Data collected in other studies similarly suggest a 1:1 sex ratio in the
spring population (Zwickel 1972, Bendell et al. 1972).
Nesting

Parameters,

Hatching

Dates and Production

Natality
Few nests were located during the course of study because of the difficulty
in finding nesting hens. All nests were located on Green Mountain while
performing other duties, as no concerted effort was made to conduct nest
searches on either study area. Clutch size and hatching success for the nests
examined are presented in Table 3. Sample size was too small to judge
whether there was a significant change in natality rate between years.
Similarly, there were insufficient data for comparison between adult and
yearling hens or between clutches laid early and late in the season.

Table 3.

No.
Nests

eJ:-I

Natality

rate among blue grouse on Green Mountain.

Clutch Size
Mean Range

Total
Eggs
Laid Unhatched

6.0

44

4-8

6

II

Depredated

Deserted

Total
Eggs
Hatched

6

2

30

Ave. No.
Young per
Female

3.7

II Includes all nests located from 1975-1978.

21

One nest abandoned

before

laying was completed.

Egg loss, including one nest that was deserted before laying was completed,
reduced the hatch by some 32 percent.
Boag (1966) reported an approximate 18
percent egg loss of Alberta blue grouse, whereas, Zwickel (1975) reported
what he considered a conservative estimate of 54 percent hatching success or
a maximum estimate of 46 percent egg loss. Overall, 50 percent (4) of the
nests hatched successfully, 25 percent (2) were partially successful, 12.5
percent (1) were destroyed by predators, and 12.5 percent (1) were deserted.
Nesting success for 6 of 8 nests completely or partially hatched was 75
percent.
Floristic and physiognomic characteristics of each nest site were recorded
and are discussed under "Habitat Investigations".
Hatching

Dates

Fig. 3 presents the hatching curves for blue grouse in the Eagle and Middle
Park areas.
Time of hatch was &amp;imilarbetween
areas, so the data were grouped.

�-211-

70
1975.
•
1976 0
0
1977 .-----.
1978 0-----0

60

(!)

50

~

:J:

U
~

I40

•

t-

1\

z

I

w
u
a:::
W
c, 30

\

I

\

,/

\

1

II

yl

/ \

/

20

!
/

10

/

1

/

\

\

\

\

I

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1

\

I

/1
•
I

II

\

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/

//

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I

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///

8-14

1-7

15-21 22-28

29-5

JUNE
WEEK OF HATCHING
,

Fig. 3.

Blue grouse hatching curves, 1975-1978.

6-12

JULY

13-19 20-26

�-212-

All chicks were aged according to procedures outline by Zwickel and Lance
(1966). The estimated age was adjusted for bias as described by Refield and
Zwickel (1976). Peak of hatch varied by as much as 3 weeks over the 4 years
of study.
Examination of the hatching curves show that no second peak,
indicative of renesting, was evident in any year.
This does not imply
renesting did not occur, but the contribution of renesting to production of
young was insignificant.
Nesting

Success

Since hens with broods are easier to locate than those without, this factor
tends to produce an inflated estimate of nesting success when calculated from
field observations.
Consequently, analyses of the wing molt pattern of
hunter harvested birds was used as the best estimate of nesting success
(Braun 1971).
The technique provides a minimum estimate as some successful
hens will have completed their molt by mid- to late September and cannot be
distinguished from unsuccessful hens.
This problem is compounded during
years when hens initiate nesting earlier such as in 1977. Few yearlings
were identifiable in 1977 and many adult hens were in the advance stages of
molt.
Therefore, nesting success was probably bett~r in 1977 than what the
data indicate.
Combined nesting success of adult and yearling hens was relatively constant
(range = 68.6 - 76.4%) from year to year in Middle Park (Table 4). Overall,
72.0 percent of the adult and yearling female wings collected in Middle Park
were from successful nesters.
This estimate is in agreement with the 75.0
percent nesting success determined from 8 nests found on Green Mountain.
Based on field observations of successful (52) and unsuccessful (15) hens,
estimated nesting success on Green Mountain from 1975 to 1978 was 77.6
percent.
Again the range of values (75.0 - 80.9%) was relatively constant
from year to year.
Nesting success varied more within and between age classes than noted for
combined success of both age classes.
Adult hens exhibited less variation
(68.5 - 80.9%) and an overall higher (74.2%) success than yearlings (x = 64.0%,
range = 33.3 - 82.1%).
Reasons for this difference are uncertain, but may
be an artifact of sample size (especially for yearlings).
Zwickel (1975)
detected no significant difference in nesting success of yearling and adult
hens.
Too few wings were collected from the Eagle area in 1975 and 1976 to estimate
nesting success in these years.
In 1977, only 4 yearling hens were identifiable from the wing sample.
Consequently, the 1977 estimate of nesting
success was for adults only. Combined success was greater in 1978 (80.9%)
than 1977 (57.1) and compared favorably with field estimates (1977 = 53.6%,
1978 = 72.7%).
Production

and Juvenile

Mortality

From mid-June until mid-August 1978, complete counts were made on 62 broods
including 32 on Green Mountain and 30 at Eiby Creek.
Some broods were
counted more than once, but not within the same month.
Results of brood
surveys from 1975 to 1978 are given in Table 5.

�-213-

Table 4.

Estimated

nesting

success, Middle

Park and Eagle,

1975-1978.

Eagle
Estimated Nesting
Adults
Yearlings~1

Year

Middle Park
Estimated Nesting Success
Adults(N)l/ Yearlingsll
Total

1975

80.3(71)

33.3(21)

69.6(92)

ND ]j

ND

ND

1976

68.3(60)

82.1(28)

72.7(88)

ND

ND

ND

1977

68.5(143)

68.7(16)

68.6(159)

57.1(63)

1978

80.9(115)

65.2(46)

76.4(161)

81.0 (58)

Total

74.2(389)

64.0(111)

72.0(500)

68.6(121)

IS!!j(4)

80.8(26)

Success
Total

57.1(63)
80.9(84)
70.7(147)

11 Total sample of wings examined.

II Due to sma 11 samp 1··
e Slzes
not be representative

ln some years, es t·
lma te d nes t·
lng
of this age class.

success may

]j No data.
il Inadequate

Yearly

sample.

average brood sizes recorded
Green Mountain

3.9
1975
4.4
1976
1977 = 3.8
1978 - 4.9

were as follows:
Eiby Creek

1975
1976
1977
1978

No Data

3.4
3.9
5.1

Average brood size was similar between areas in 1977 and 1978. The apparently
smaller brood size at Eiby Creek in 1976 was attributed to a larger sample
of broods counted in late July-early August, whereas, primarily early to midJuly counts were made on Green Mountain.
Rogers (1968) found average brood
sizes of 4.1, 2.6 and 3.9 in 3 years of study in western Colorado.
Since some doubt exists as to whether blue grouse broods can be accurately
counted until after they are 4 weeks old (Boag 1966), late June and early to
mid-July counts may be underestimated.
Brood counts in August may also be
misleading due to shuffling of chicks between broods, formation of gang
broods, and brood breakup and dispersal.
Therefore, observed differences
in brood size from June to August may not accurately reflect loss of chicks
during this period (Table 6). The gradual attrition in size of broods over
the summer months is the least reliable method of evaluating chick mortality
(Zwickel and Bendell 1967), but was the only data available for determining
losses in this study. Average annual summer mortality of chicks was 43.6
percent for Green Mountain ar.d only 28.0 percent for Eiby Creek.

�-214-

Table 5. Average brood size, range, and number
intervals, 1975 to 1978. 1/

of broods

observed

by monthly

1975

Green Mountain
1976
1977

1978

1976

Mean

ND'!:/

5.3

5.1

6.2

5.0

4.6

6.5

Range

ND

3-7

4-7

5-9

4-6

3-6

6-7

ND

3

11

9

2

7

2

Mean

5.2

4.1

4.0

5.4

3.7

4.1

4.9

Range

2-7

2-6

1-7

3-7

1-7

2-9

3-8

Sample Size

4

7

10

9

9

10

l3

Mean

3.0

3.0

2.5

3.7

2.2

3.6

5.0

Range

1-5

1-4

2-6

1-4

1-10

3-8

13

14

5

20

17

Month

Eiby Creek
1977

1978

June

Sample

Size

July

August

Sample

Size

1/ Only distinct

6

1

broods with full counts are included.

1/ No data,

late hatching

Production

and Breeding

Estimated
presented
following
studies:

total production in relation to breeding population levels is
in Table 7 for both study areas.
Calculations are based on the
assumptions which are supported by data collected in this and other

year, no full counts obtained.

Populations

(1)

A 1:1 sex ratio exists

(2)

Estimated

(3)

Mean annual brood size and monthly estimated
the production of chicks on the study areas

(4)

Immigration

percent

equals

in the breeding

nesting

success

emigration

population

is correct
brood

sizes represent

�-215-

Table 6.

Mortality

Area

of juvenile

blue grouse from late June to mid-August.

Year

Mean Brood Size
June
August

1975

5.21/

3.0

42.3

1976

5.3

3.0

43.4

1977

5.1

2.5

51.0

1978

6.2

3.7

40.3

1975-78

5.5

3.1

l.3.6

1976

5.0

2.2

56.0

1977

4.6

3.6

21.7

1978

6.5

5.0

23.1

1976-78

5.0

3.6

28.0

Mortality

(%)

Green Mountain

Eiby Creek

1/ Mean brood

size for July as no full counts were obtained

in June due to

late hatch.

Zwickel (1965) and Bendell and Elliott (1967) reported a mean annual death
rate of approximately 30 percent for breeding populations of blue grouse on
Vancouver Island, British Columbia.
Of 6 males and 4 females banded on Green
Mountain between April and mid-June 1977, 6 (4 males, 2 females) were
recensused in spring 1978. During the same period 4 males and 4 females were
banded at Eiby Creek, and 5 (2 males, 3 females) were reobserved in 1978.
The pooled data on birds banded in 1977 and resighted in 1978 gave an
estimated mortality rate of 39 percent.
Assuming this estimate is constant
from year to year, one can determine from the estimate of total production
the minimum replacement requirements necessary to maintain a stable breeding
population (Table 8).
Production by mid-August rather than total production was used in the computations because mid-August figures more closely approximate production and
survival of juveniles until fall. The replacement requirement then becomes
an estimate of the fall to spring mortality of juveniles that should occur
in a stab~e population.
Both populations under investigation were essentially
stable.

�Table 7.

Area

Estimation of the annual production of young on Green Mountain and Eiby Creek.

Total
Production

Mortality
June-August

Total
Production
by mid-August

Total
Population
by mid-August

Percent
Gain

3.9

43

42.3

25

57

44

12

4.4

53

43.4

30

62

48

68.6

12

3.8

46

51.0

22

56

39

76.4

14

4.9

69

40.3

41

79

52

Total~J
Breeding
Population

Nesting
Success
(%)

No. Hens
With
Broods

1975

32

69.6

11

1976

32

72.7

1977

34

1978

38

Year

Average
Annual
Brood Size

Green Mountain

I
N

•....•
C]\

Eiby Creek

I

1977

58

57.1

17

3.9

66

56.0

29

87

33

1978

60

80.9

24

5.1

122

23.1

94

154

61

!/ Excludes non-territorial males.

�-217-

Table 8. Estimates of the fall to spring loss of juveniles
maintain a stable breeding population.

to

Total
Breeding
Population

Annual Mortality
Breeding
Populationl/

Total
Production
by mid-August

Fall to Spring
Mortality of
Juveniles~j (%)

1975

32

12

25

52

1976

32

12

30

60

1977

34

13

22

41

1978

38

15

41

63

1977

58

23

29

21

1978

60

23

94

75

Year

Area

necessary

Green Mountain

Eiby Creek

!/ Number

of birds expected to be lost from the population
annual mortality rate of 39 percent.

-2/ Morta 1·lty necessary

.
.
to malntaln

assuming

a constant

the popu 1·atlon.

It is apparent from the data presented in Table 8 that production was more
than sufficient to replace natural losses in the breeding population.
Excluding the data from 1977 at Eiby Creek, at least 41 percent (Green
Mountain 1977) and up to 75 percent (Eiby Creek 1978) of the juveniles alive
at the end of summer would either have to succumb to mortality or disperse
to other areas between fall and spring in order for the breeding population
to remain stable.
For Green Mountain, the average annual fall to spring loss
of juveniles required to maintain the population from 1975 to 1978 was 56
percent.
These data suggest there are always surplus birds in the fall
population and in most years this surplus exceeds 40 percent.
Harvest
Hunting

Season

The 1978 blue grouse season opened on 9 September and closed on 8 October.
Blue grouse hunting was also permitted between 14 October and 14 November in

�-218-

conjunction with the deer, elk and combined deer elk seasons.
Season length
was 57 days with daily bag and possession limits of 3 and 6 birds, respectively.
Season structure, bag limits, and length from 1975 to 1978 are presented in
Table 9.
Table 9.

Blue grouse hunting

Year

Hunting
Season
Dates

1975

1976

1977

1978

Middle

seasons,

Colorado,

1975-1978.

Season
Length
(days)

Bag
Limit

Possession
Limit

9/13-10/5

23

3

6

Unit 80 and all units
west of Interstate 25
except portions of Unit
52

9/11-10/10
9/11-10/10
and
10/16-10/26

30
40

3
3

6

Same as 1975
Unit 28 (Middle Park) only

9/10-10/9

30

3

6

10/15-11/15

27

3

6

West of Interstate 25 and
Unit 80
West of Interstate 25
and Unit 80 when open to
deer or elk hunting

9/9-10/8
10/14-11/14

30
27

3
3

6
6

Same as 1977
Same as 1977

6

Open Areas

Park

As in 1977, a check station was again operated in 1978 at the Prairie Point
Campground along Highway 9 at the south end of Green Mountain Reservoir.
The
check station was operated opening weekend from about 1000 to 1800 MST, depending upon traffic load. All vehciles were stopped on the highway, but only
hunters were directed to pull off at the check station.
Data obtained per
party included:
County of origin, number of hunters, hours hunted, birds
observed, birds bagged per hunter, area hunted, and location where each bird
was harvested.
One wing was removed from each bird provided no wings had been
deposited in barrels.
Whenever a bird was missing one or both wings, the
hunter was questioned as to what he did with the wings.
Thus, the status of
all wings was recorded as follows: (1) hunter disposed of wings, (2) hunter
deposited wing in barrel, and (3) wing collected at check station.
Sex by
gonadal inspection was ascertained for all young of the year whenever possible.
Ovaries were collected from adult and yearling hens and crops were collected
from all birds when present.
Whole body weights were obtained for all grouse
that were not eviscerated.

�-219-

In addition to the check station, 16 volunteer wing collection stations were
available to hunters throughout the entire season at various locations in
Middle Park. Hunters were also contacted opportunistically
in the field
whenever possible.
Themail
wingsurveywasdiscontinuedin1978.thus.no
wings were obtained from this source.
A total of 1,006 blue grouse wings were collected from Middle Park in 1978.
Wing collection stations accounted for 87.0 percent (875 wings) of the wings
collected.
Number of wings collected from each barrel were as follows:
Ute
Pass 19, Williams Peak Road 50, Spring Creek 169, Lawson Ridge 3, Trough
Road 85, Kremmling 29, Troublesome 16, Corral Creek 80, Cottonwood Pass 16,
Willow Creek 57, Gore Pass 154, Pinto Creek 23, and Chimney Rock 136. Cottonwood Pass, Parshall Divide, and Willow Creek represent new barrel locations.
No barrels were placed at City Reservoir and Rock Creek in 1978. The
remaining wings were collected at check stations (105 = 10.4%) in North Park
(11) and Middle Park (94) and from miscellaneous collections (26 = 2.6%).
Comparative data from 1975 to 1978 are presented in Table 10.

Table 10. Number and source of blue grouse wings collected
from 1975 to 1978.

Wing Barrels
No.
%

Year

Check Stations
No.
%

Mail Survey
No.
%

in Middle Park

Miscellaneous
No.
%

Total

1975

120(10)1/

65.6

26

14.2

o

0.0

37

20.2

183

1976

292(13)

84.9

49

14.2

3

0.9

o

0.0

344

1977

501(15)

82.7

87

14.4

18

2.9

o

0.0

606

1978

875(16)

87.0

105

10.4

o

0.0

26

2.6

1,006

83.6

267

12.5

21

1.0

63

2.9

2,139

1,788

1/ Number in parentheses

represents

number of wing barrels

used.

It is apparent from data presented in Table 10 that number of wings collected
each year increased by almost 2 fold over the number collected the previous
year. Liberalization of the blue grouse season partially contributed to the
increased number of wings collected.
However, relocation of wing barrels to
more productive sites plus use of additional wing barrels were largely
responsible for yearly differences.
During the 2 days of check station operations, 222 hunters with 294 blue grouse
(1.3 birds per hunter) were checked.
Total birds reported observed by these
hunters was 758. Some duplications are undoubtedly present in the observations.
Of the 222 hunters checked, 45.4 percent were unsuccessful, 19.8 percent

�-220-

harvested 1 bird, 21.3 percent harvested 2 birds, and 13.5 percent harvested
the daily bag limit of 3 birds.
Only 9 birds were reported wounded and not
retrieved for an estimated crippling loss of 3.1 percent.
Hunter efficiency
was calculated at 38.8 percent (birds bagged;
birds observed ~ 100). Comparative data from 1975 through 1978 are presented in Table 11.

Table 11.

Blue grouse harvest

statistics,

Middle Park, Colorado,

1975-1978.11

Year

No.
Hunters
Checked

No.
Birds
Observed

No.
Birds
Bagged

Hunter
Efficiency
%

%
Successful
Hunters

Crippling
Loss
%

Birds
per
Hunter

1975

112

158

45

28.4

30.4

4.4

0.4

1976

138

141

61

43.2

33.4

8.2

0.4

1977

228

480

226

47.1

39.5

4.0

1.0

1978

222

758

294

38.8

54.6

3.1

1.3

11 Check stations operated on the Williams Peak road in 1975 and 1976 and at
the Prairie

Point campground

in 1977 and 1978.

In order to evaluate the effectiveness of wing barrels for sampling, Middle
Park hunters were questioned as to what they did with the wings from the birds
they harvested.
This information is summarized below along with similar data
collected in 1977:
.
1978
1977
Percent wings deposited in barrels
60.7
50.6
Percent wings collected that should have been deposited
20.3
18.9
Percent wings disposed
17.4
28.3
Percent wings of unknown status
3.0
0.8
Distribution of harvest and hunting pressure was not uniform within Middle
Park.
Spring Creek was the leading harvest area in 1978 accounting for 18.8
percent of the harvest and 30.7 percent of the hunters.
Gore Pass was next
with 19.3 percent of the hunters and 17.2 percent of the harvest.
Chimney
Rock accounted for 14.0 percent of the harvest but only 6.4 percent of the
hunters, whereas, Piney had 9.7 percent of the harvest and 15.2 percent of
the hunters.
Blue Ridge ranked third in terms of hunters pressure (16.5%)
but only made up 7.4 percent of the harvest.
Corral Creek and Willow Creek
had 8.0 and 5.7 percent of the harvest, respectively.
All other areas within
Middle Park each had less than 4 percent of the harvest and hunter pressure.
The only other year for which comparative data are available is 1977. The
Leading harvest areas in 1977 were Chimney Rock (16.0% of harvest), Piney
(15.5%), Gore Pass (14.1%), Blue Ridge (12.7%), Spring Creek (12.2%), Corral
Creek (9.1%), and Ute Pass (6.9%). Major hunting areas in 1977 were Spring
Creek (33.0% of hunters), Blue Ridge (23.4%), Gore Pass (14.8%), Piney (11.5%),
and Chimney Rock (8.1%).

�-221-

Most hunters contacted in 1978 originated from the Denver metropolitan area.
The counties of Jefferson (49), Denver (31), Arapahoe (28), and Adams (22)
accounted for 130 of 206 (63.1%) hunters for which origin was ascertained.
Local hunters from Summit (23) and Grand (7) counties made up 14.6 percent
of the hunters contacted. This figure is probably under estimated as
appreciable numbers of local residents (especially from Grand County) are
not contacted at the check station. All other counties each comprised less
than 5 percent of the hunter contacts. Similar data have been collected in
previous years.
Distribution of the wings collected in 1978 according to time of collection is
given in Table 12, along with comparative data from 1975 through 1977. These
data are presented to illustrate how more liberal seasons have spread the
harvest and hunting pressure over a longer period of time. Whereas, hunting
pressure has probably remained the same, pressure has declined opening weekend
as more hunters are hunting later in the season.
Table 12. Time distribution of blue grouse wings collected, Middle Park,
Colorado, 1975-1978. 1/
1976
%
No.

1977
No.

1978

Time Period

1975
%
No.

1st weekend

53

44.2

121

41.4

188

37.5

288

32.9

1st week

9

7.4

36

12.3

20

4.0

99

11.3

2nd weekend

20

16.7

36

12.3

63

12.6

109

12.5

2nd week

0

0.0

11

3.8

28

5.5

13

1.5

3rd weekend

8

6.7

23

7.9

44

8.8

55

6.3

3rd week

11

9.2

10

3.4

7

1.4

13

1.5

4th weekend

19

15.8

21

7.2

18

3.6

23

2.6

4th week

8

2.7

11

2.2

10

1.1

5th weekend

15

5.2

12

2.4

119

13.6

Experimental
Season (1976)
10/16 to 10/26

11

3.8

Deer season

37

7.4

67

7.7

Elk fleason

63

12.6

44

5.0

Combined season

10

2.0

35

4.0

501

100.0

875

100.0

Totals

120

100.0

292

1/ lUng collections from barrels only.

100.0

%

No.

%

�-222-

Age and sex composition of the 1978 harvest sample as ascertained from
examination of wings is presented in Table 13. Data from 1975 to 1977 are
included for comparison.
There were significant changes in age and sex
structure of the harvest in 1978 from previous years.
In 1978, immatures
comprised 67.2 percent of the harvest, the best representation of immatures
during the 4 years of intensive wing collections in Middle Park. Fewer adults
were present in the 1978 harvest sample than in previous years.
Since some yearlings will have completed their primary molt by mid- to late
September, they cannot be distinguished from adults.
Therefore, it is
believed yearlings are under-represented
in the harvest in all years.
The
problem is most evident in "early" years (1977) because the birds initiate
their primary molt earlier.
Also, the problem of separating adults and
yearlings is more pronounced for males than females because males start their
primary molt before most females.
Thus in 3 (1975, 1976 and 1977) of 4 years
of data, there were significantly fewer males than females comprising the
yearling segment of the harvest.
Sex ratio of adults and immatures approximated 1:1 in all years except 1978 when there were significantly more immature
males than females.
Reasons for the deviation in sex ratio of chicks in 1978
is uncertain, but may be related to a greater harvest of male chicks late in
the season.
Eagle
A total of 465 blue grouse wings was collected in the Eagle area during the
1978 season.
Ten wing collection stations accounted for 97.6 percent (454
wings) of the wings obtained.
Number of wings collected from each barrel
were as follows:
Eiby Creek North 78, Eiby Creek South 26, Cabin Creek 5,
Coffee Pot Springs 67, Gypsum 8, Brush Creek 28, Milk Creek 73, Red and White
Mountain 68, Wolcott 38, and Red Sandstone Road 63. The remaining wings
were collected from the mail wing sllrvey (3) and at the Idaho Springs big
game check station (8). Due to low hunter pressure and scattered distribution
of hunters, no check station was operated in the Eagle area in 1978. Approximately 26 percent more wings were collected in 1978 (465) than in 1977 (342).
This increase was probably due to the relocation of wing barrels to more
productive sites rather than an increa~e in harvest.
Data presented in Table 14 were in agreement with the same information
collected in Middle Park (Table 12). In general, liberalization of the
season spread the hunter pressure and harvest over a longer period of time,
thus reducing the opening weekend syndrome.
Age and sex composition of the 1977 and 1978 harvest samples from the Eagle
area is shown in Table 15. Inadequate samples of wings were collected in
1975 (21) and 1976 (67) to permit reliable analysis.
The blue grouse harvest
samples from the Eagle area are similar to data collected in Middle Park for
the same years.
From 1977 to 1978 there was a substantial decline in the
adult segment of the harvest, with a corresponding increase in harvest of
juveniles.
Data from wing analyses suggest excellent production of young
in 1978 and possibly better survival; an analogous conclusion was drawn from
data presented in Tables 5 and 6.

�Table 13.

Age and sex composition of the blue grouse harvest, Middle Park, Colorado, 1975-1978.

Adults
Females
%
No.

Total
%
No.

Males
%
No.

Yearlings
Females
%
No.

Tota1
%
No.

Males
%
No.

Immatures
Females
%
No.

Total
%
No.

Year

Males
%
No.

24.1

71

41. 8

5

2.9

15

8.8

20

11. 7

36

43

79

46.5

41

25.3

30

17.7

21.2

1975

18.8

132

38.7

15

4.4

28

8.2

43

12.6

74

92

166

48. 7

64

27.0

68

19.9

21.7

1976

23.9

277

45.9

3

0.5

16

2.6

19

3.2

164

27.2

50.9

144

307

22.1

23.7

133

143

1977

11.9

237

24.4

36

3.7

46

4.7

82

8.4

364

37.4

67.2

116

654

12.5

29.8

121

290

1978

I
N
N
W

I

�-224-

Table 14. Time distribution
Colorado, 1977-1978.

of blue grouse wings collected,

1977

Eagle Area,

1978
Total

No. of
Wings

% of
Total

127

37.1

126

27.1

1st week

48

14.0

62

13.3

2nd weekend

21

6.2

54

11. 6

2nd week

27

7.9

9

1.9

3rd weekend

10

2.9

17

3.7

3rd week

10

2.9

18

3.9

4th weekend

10

2.9

29

6.2

4th week

5

1.5

14

3.0

5th weekend

11

3.2

44

9.5

Deer season

39

11. 4

62

13.3

Elk season

15

4.4

27

5.8

Combined

19

5.6

3

.7

342

100.0

Time Period

No. of
Wings

1st weekend

season

Totals

% of

465

100.0

As previously noted, not all yearlings could be identified in the harvest
samples, therefore creating what appeared to be deficiency.
The Eagle data
showed a greater deficiency of yearling females in 1977 and yearling males
in 1978.
In Middle Park, the deficiency of yearlings was most pronounced
for males in all years.
The observed sex ratio of immatures in 1978 was
exactly 1:1 which was incongruent with the unbalanced sex ratio favoring
males that was noted for the Middle Park sample in 1978. Otherwise, the sex
ratio of adults and immatures did not deviate significantly from 1:1 and
was in agreement with the Middle Park data.
Utilization
Direct evidence from banding data and indirect evidence based on the attributes
of the populations under investigation indicate that present levels of
harvest have minor effects on grouse populations.
Available data suggest
there is a high and variable loss of grouse during their first year.
Further-

�Table 15.

Age and sex composition of the blue grouse harvest, Eagle Area, Colorado, 1977-1978.

Adults
Females

Males

Males

Total

Yearlings
Females

Imroatures
Females

Males

Total

%

No.

%

No.

%

N-o-.--

No.

%

No.

%

26.7

171

50.7

Year

No.

%

No.

%

No.

%

87

25.8

65

19.3

152

45.1

10

3.0

4

1.2

14

4.2

81

24.0

90

1977

62

13.4

59

12.8

121

26.2

9

2.0

26

5.6

35

7.6

153

33.1

153

1978

,

Total

%

N-o-.--

I
N

33.1

306

66.2 ~
I

�-226-

more, production of young greatly exceeds the number necessary to replace
natural losses in the breeding population.
The excess birds are expendable
and may be harvested without adversely affecting the subsequent spring
breeding population.
While few birds were banded, the percentage of banded birds shot (4.8%)
indicate hunters removed a negligible portion of the fall population (Table
16). The juvenile segment of the population sustained the bulk of the kill
(13.5%) followed by hens (1.9%). No banded males were reported shot. Bendell
and Elliott (1967) reported approximately 5 percent of the hens and chicks
banded on their study areas on Vancouver Island were shot each year, while
very few (0.7%) adult males were taken, Mussehl (1960) found that hunters
removed 7 (1957) and 12 (1958) percent of the grouse banded in the Bridger
Mountains, Montana, most of which were juveniles.

Table 16. Number of grouse available
and Eagle, Colorado, 1976-1978.

Year

Males
No.
%
Banded
Shot

to and shot by hunters,

Females
No.
%
Banded
Shot

Chicks
No.
%
Banded
Shot

Middle

Park

Total
No.
%
Banded
Shot

1976

6

0.0

13

7.7

5

0.0

24

4.2

1977

20

0.0

15

0.0

14

21.4

49

6.1

1978

9

0.0

25

0.0

18

11.1

52

3.8

Totals

35

0.0

53

1.9

37

13.5

125

4.8

Hickey (1955) says that gallinaceous birds can safely withstand a hunting kill
equivalent to about one-half their annual mortality rate. Based on a 4 year
·average for Green Mountain (Table 7), production contributes to approximately
a 46 percent increase in the population.
In a stable population there must
be an annual loss from fall to fall of this amount.
Therefore, according to
Hickey (1955), the population can absorb a harvest of 23 percent.
This
represents a minimum estimate because the annual mortality of adults is not
taken into account.
Even so, the calculated yield (23%) surpasses the estimated harvest of 4.8 percent and further substantiates the conclusion that
hunting has no detrimental effects on blue grouse populations.
Combined

Blue Grouse - Big Game Season

Evaluation of the 1978 combined blue grouse - big game season followed the
exact procedures used in 1977. A total of 1.500 big game hunters were
surveyed at the Idaho Springs check station during the 1978 deer (500), elk

�-227-

(500), and combined deer-elk (500) seasons.
Hunters were contacted opportunistically at the check station.
Only one hunter from each party was
surveyed and only residents were included in the sample.
To insure a
diversity of hunters were questioned, sampling periods included days at the
beginning, middle, and end of each season.
From data presented in Table 17 it is apparent that few big game hunters
took advantage of the longer grouse seasons in 1977 (6.3%) and 1978 (5.5%).
Based only on those hunters aware the grouse season was open, participation
was 13.1 percent in 1977 and 9.8 percent in 1978. Even though participation
was low, general response of hunters to the combined blue grouse - big game
season was highly favorable in both years.
The primary complaint of those
hunters opposed to the season was their concern about the potential increase
in number of hunters (big game + grouse hunters).
However, less than one
percent of all hunters contacted during the 1977 and 1978 big game seasons
were strictly grouse hunters.
More hunters were aware of the longer grouse
season in 1978 (55.1%) than in 1977 (48.1%), but participation remained
relatively constant.
Therefore, only a small percentage of hunters were
interested in hunting grouse during the big game season.
Due to good production this year, number of grouse observed increased
substantially in 1978 (4,892) compared to 1977 (3,415).
Consequently, hunters
that took advantage of the longer season were more successful in 1978 (48.5%
success, 2.3 grouse/successful
hunter) than in 1977 (27.4% success, 2.0
grouse/successful
hunter).
As a result, fewer hunters killed more grouse in
1978 than in 1977. Based on wing collections from barrels in Middle Park,
22.0 and 16.7 percent of the total grouse harvest occurred during the big
game seasons in 1977 and 1978, respectively.
In 1977 the projected estimate of the total harvest of blue grouse in Colorado was about 26,200 birds as determined from the statewide small game survey.
Whereas, this is probably an inflated estimate, the additional harvest
resulting from the longer season in 1977 (22% of total harvest) was somewhere around 5,800 grouse, or about a 26 percent increase over the 11 year
(1966-1976) average annual harvest of 21,100 blue grouse.
Results of the
1978 small game survey were not available, but preliminary indications are
that the harvest was up from 1977.
Habitat
Vegetation

Investigations

Characteristics

Green Mountain lies in the southwest corner of Middle Park, one of three
major intermountain parks in Colorado.
Unlike the other intermountain parks
characterized by broad, rolling grass (South Park) or sagebrush (Artemisia
spp.) (North Park) covered f1oor.s, Middle Park is mountainous and locally
heavily forested; however, sagebrush types still predominate.
The study area
includes portions of both the sagebrush and Douglas-fir
(Pseudotsuga
menziesii) vegetation zones as described in Harrington (1964).
Scattered to
dense stands of Douglas-fir predominate above 2700 meters and extend downward
to 2600 meters.
Open areas below this elevation are vegetated primarily with
big sagebrush (!. tridentata).
Intermediate areas are best described as a
mixed conifer-aspen (Populus tremuloides)-shrub
association.
This zonal
pattern of vegetation is not distinct as extensions of various sagebrush
types occur throughout the Douglas-fir zone creating a patchwork of vegetation types.

�Table 17.

Combined blue grouse-big game season hunter questionnaire data, Idaho Springs, Colorado, 1977 and 1978.

No. in sample

1978
Combined

Deer

Elk

1977
Combined

Total

Deer

Elk

500

500

502

1,502

500

500

500

1,500

Total

No. big game hunters aware of season

253

260

210

723

259

285

283

827

% big game hunters aware of season

50.6

52.0

41.8

48.1

51.8

57.0

56.6

55.1

No. big game hunters hunting grouse

40

25

30

95

14

37

31

82

% big game hunters hunting grouse

8.0

5.0

6.0

6.3

2.8

7.4

6.2

5.5
I

tv
tv

% big game hunters aware of season

that hunted grouse

15.8

9.6
2,102

14.3
620

l3.1
3,415

5.4
1,384

13.0
2,053

10.9
1,455

9.8
4,892

Total grouse observed

693

% big game hunters observing grouse

24.8

41.8

18.5

28.4

32.4

35.6

37.8

35.0

Grouse observed/big game hunter

1.4

4.2

1.2

2.3

2.7

4.1

2.9

3.2

Total grouse harvested

l3

28

12

53

17

28

32

77

Grouse/big game hunter hunting grouse

0.3

1.1

0.4

0.6

1.2

.7

1.0

0.9

52.0

52.2

52.0

52.1

60.0

57.0

62.0

59.7

% negative

l3.4

9.2

10.4

11.0

14.0

11.0

9.6

11.5

% indifferent

34.6

38.6

37.6

36.9

26.0

32.0

28.4

28.8

% hunters favorable of combined blue

grouse-big game season

'F

�-229-

Canopy coverage for the entire study area is about 40 percent.
Seventy percent of the crown cover is Douglas-fir, 27 percent aspen, and 3 percent Rocky
Mountain juniper (Juniperus virginiana) and mountain maple (Acer glabrum).
Understory cover consists of a mosaic of shrub and grass-forb types.
Snowberry (Symphoricarpos spp.), common juniper (l. communis), rose (Rosa
acicularis), chokecherry
(Prunus virginiana), and serviceberry (Amelanchier
spp.) are the major understory shrubs; yarrow (Achillea lanulosa), pussy toes
(Antennaria spp.), northern bedstraw (Galium boreale), and peavine (Lathyrus
spp.) are the major forbs encountered in the understory, and sedge (Carex spp.)
and bluegrass (Poa spp.) are the major grass and grasslike species.
Unforested areas constitute about 60 percent of the study area of which in
excess of 90 percent is dominated by sagebrush.
Other shrubs, forbs, and
grasses (or grasslike) commonly found in association with the sagebrush include:
snowberry, rabbitbrush (Chrysothamnus spp.), serviceberry, rose, bitterbrush
(Purshia tridentata), pussy toes, arrowleaf balsamroot (Balsamorhiza sagittata),
paintbrush (Castelleja spp.), sulphur flower (Eriogonum umbellatum), common
lupine (Lupinus argenteus), bedstraw, sedge, wheat grass (Agropyron spp.),
Junegrass (Koleria cristata), needle-grass (Stipa spp.), and bluegrass.
Plant
nomenclature is according to Weber (1972) and Nelson (1969).
Several factors have markedly influenced the composition and structure of
vegetation on Green Mountain.
The earliest record of disturbance was between
1880 and 1890 when the northern portion of the study area was set on fire by
a local homesteader to create a supply of firewood.
Much of the area burned
has reverted to sagebrush with charred logs, stumps, and patches of Douglasfir dotting the landscape.
Around 1930 the Douglas-fir was selectively
logged as a local source of ties for contructing the railroad.
Few trees
were removed, but enough to open up the habitat and allow the development of
a dense understory and in some areas, invasion of aspen.
The most extensive disturbance occurred in 1962 when sagebrush dominated
areas below the Douglas-fir zone were sprayed with 2,4D to enhance grass
production for livestock.
The degree of mortality in shrub species was not
uniform throughout the sprayed area, thus vegetative composition and structure
were altered moreso at some locations than others.
Adequate time has since
elapsed for the sagebrush to recover except in a few small areas where spraying
was most effective.
Dead or partially damaged sagebrush plants were still an
important component of ground cover in such areas.
Past grazing history indicates moderate to heavy pressure on portions of the
area by domestic livestock.
Present grazing includes light summer use by
cattle mostly at the lower elevations near available water.
Mule deer
(Odocoileus hemionus) use the area from late spring until early winter.
Besides grazing, no other use is presently made of the area.
Vegetation

Type Mapping

Vegetative classification and description of the Green Mountain study area
was done in accordance with procedures outlined by Kuchler (1955). This
method of vegetation mapping is essentially a derivation of Braun-Blanquet's
(1951) system of floristic and physiognomic description of plant communities.
Vegetation maps have been prepared elsewhere in Colorado using this method
(Braun 1969, Medin 1962).

�-230-

Aerial photographs (approximately 7.0 cm:l km) of the study area were used
as base maps.
Before going into the field, the photos were carefully
examined and every area of vegetation appearing dissimilar from adjacent
areas was delineated by a line drawn directly on the photo.
The minimum
area of vegetation bounded by a line on the photo was approximately .5
hectares.
No types were omitted within the study area boundary.
Upon delineation of all areas apparent on the map, each area was then inspected
in the field.
The first task was to walk about the area being examined from
end to end and across, observing the vegetation critically and field checking
the boundaries shown on the photo.
Salient features of the area where
recorded on prepared data sheets.
Consecutive numbers were assigned to each
area inspected and were entered both on the data sheet and the photograph.
Each area received its own number regardless of any similarities to previously
inspected areas.
To better facilitate preparation of the final map, a
photograph was taken of each vegetative unit. Number of the individual
exposure was recorded along with the corresponding number of the unti where
the photograph was taken.
The next step was to record the vegetative physiognomy and other physical
features of the area including: slope gradient, exposure and character.
The
physiognomic classification
(Table 18) reveals the appearance and structure
of the plant community (vegetation unit), i.e., height and density of every
item listed, and in addition, such special features as may be present.
A
detailed description of the classification scheme and its application is
presented by Kuchler (1949). Slope gradient and exposure were measured with
an Abney level and compass.
After the physiognomic formula for an inspected area had been established,
attention was focused on the floristic character of the vegetation.
All
species occurring on the area were recorded by name and each species was
assigned numerical values describing its coverage and sociability (Table 19).
Coverage is understood to mean the percentage of ground that would be covered
if the full spread of the species were projected vertically to the ground.
Sociability refers
to the distribution of a species within the area of
vegetation under consideration.
Undoubtedly, some species which only very
few specimens were present in a given type escaped detection.
This was also
true for "early or late flowering" species that peaked prior to or after the
period of field work, which was designed to coincide with the peak of plant
development (mid-June to mid-July).
Additional features such as the presence and amount of bare ground, stumps,
logs and rocks were recorded for each vegetation unit. Comments on physiography, soil, water condition, whether the area was logged or burned, and any
other pertinent information were noted on the data form.
The result of the laboratory and field activity was a set of lists and notes
and a series of a.erial photos on which the vegetation units were outlined.
Each unit received its own number corresponding to the number in the lists
where the particular type was analyzed.
The next step was the preparation
of the base map which shows the exact outline of each individual vegetation
unit and its number.

�-231-

T abl e 18 .

"
Ph"YSlognomlc"1 c aSSl"f" lcatlon

0f

."
1/
vegetatlon.-

CAPITAL LETTERS:
Herbaceous Vegetation:
G: Graminoids
H: forbs
L: lichens and mosses

Woody Vegetation:
B: evergreen broadleaf
D: deciduous broadleaf
E: evergreen needleleaf
N: deciduous needle leaf
0: without leaves
SMALL LETTERS:
Group I:

Height:
t: tall;

m: medium

Minimum height of trees:
Minimum height of herbaceous
tall;

1: low;

Height of trees:
Height of herbaceous

plants:

25 m
2 m
10-25 m
~-2 m

plants:

10 m
~ m

Maximum
Maximum

height of trees:
height of herbaceous

s: shrubs;

Minimum

height:

1 m

z: dwarf shrubs;

Maximum

height:

1 m

plants:

Group II: Density:
c: continuous

growth

i: interrupted;

plants usually

p: plants scattered

do not touch

singly, or in groves or patches

r: rare, yet conspicuous
b: barren; vegetation

largely or entirely

absent

Group III: Special Features:
e: epiphytes

u: palms

j:

v: bamboos

lianas

k: succulents

w: adquatic vegetation

q: cushion plants

y: tree ferns and tuft plants

1/ From Kuchler

(1955).

�-232-

Table 19.

Floristic

classification

of vegetation.1/

Coverage:

Sociability:

+ Very sparsely present; cover

1

Growing

singly
or tufted

very small
2

Grouped

Plentiful but less than 1/20
of the area

3

In small patches

2

Covering

1/20 - 1/4 of the area

4

In extensive

3

Covering

1/4 - 1/2 of the area

5

In great crowds

4

Covering

1/2 to 3/4 of the area

5

Covering

greater

- From Kuchler

(1955).

1

1/

patches

than 3/4 of the area

Preparation of the base map involved sorting through the data forms, notes
and photographs and rearranging the vegetation units into combinations
encompassing similar floristic and physiognomic classifications.
Woody
plants including trees and shrubs were given primary emphasis during the
first phase in the sorting and combining process.
Units within these major
vegetation types were then manipulated to produce categories based on similar
floristic and structural characteristics.
After assigning each unit to a
major vegetation type and combining similar vegetation units, the final map
was traced off the base map, lumping together all similar units which were
adjacent to each other.
All similar units were then reassigned the same
number to simplify reading the map.
Total area mapped as planimetered from a U. S. Geologic Survey topographic
map (scale 1:62500) was 181.3 hectares (1.8 km2). Area encompassed by each
vegetation type and unit was calculated by the weight method (Welch 1948,
p. 85). An exact model of the vegetation map was traced onto high quality
bond paper.
The model was cut out and weighed on an analytical balance to
the nearest milligram.
Each vegetation type was then cut out and weighed.
Finally, the weight of each individual unit was recorded.
Percentage of the
total area encompassed by each vegetation type and unit was computed by
dividing the weight of each type and unit by the total weight.
The percentages were then multiplied times the total area (181.3 hectares) to estimate
the area of each vegetation type and unit.
Five major vegetation types and 26 vegetation units were recognized on the
Green Mountain study area (Figs. 4 and 5). Vegetative summary tables
(Appendix) were compiled from the data sheets and are presented as supplementary information describing the physiognomy, floristics, and physiography
of each vegetation unit. Examination of the vegetation map might indicate
that the vegetation types and units were clearly distinct.
While the major

�-233-

GREEN MOU

,

~

)('

J[&amp;~~

~

"

••..
•...

..

~
~&gt;~

)

~;&gt;

•...•..•..•...

~

')l'

l

,I(

~
;"'''

'"

II

l

~
~

r"

,..f"'" ~'"

x~
x {

'5c

~

'(l

~~

- - "~)

~

~

:

8 '1I:S..JI~~""'.....r

'x~

~
.=.... ...

(i)

Artemisia - mixed shrub (77.1).V ~
Pseudotsuga - mixed shrub (43.7)DID
Pseudotsuga - Populus - mixed
shrub (44.4)
Populus - mixed shrub (13.2)
Carex-A gropyron - scattered

~~

~~~~~~~~--~

0

~

§

t&lt;::l\

-;

E§§

10·.
••

~

Surface aNa

~~

.•••

L~

=::::0

~~UJ

••

~

in hectares

A~

:.

~

:

:

.~.L
••:.~
G

e

o

~'X:'X
x

t~ lV ~ ) r

~

e

-

,~I~

~

~-

1/
-

'~

,~~.

rn

shrub (2.9)

~~

j.

~,

,I

l~
~~

H

e:

••

"

)(

~

SCALE
I

0

~~

1

~2

I(

I km ~ ~

'l...

1.., •••1.., ••••

~
LJ

Fig. 4.

Major vegetation types, Green Mountain study area.

V

�-234-

GM

1

Artemisia-mixed

GM

2

Artemisia-Symphoricarpos-mixed

shrub-scattered

GM

3

Artemisia-Chrysothamnus-mixed

shrub

GM

4

Artemisia-Amelanchier-mixed

GM

5

Artemisia-mixed

GM

6

Artemisia-Agropyron-Chrysothamnus

GM

7

Dense Pseudotsuga-Symphoricarpos-mixed

shrub

GM

8

Dense Pseudotsuga-bareground-scattered

shrub

GM

9

Dense Pseudotsuga-Juniperus-mixed

shrub-scattered

Pseudotsuga

shrub-scattered

shrub-rock-scattered

Pseudotsuga

Pseudotsuga

Pseudotsuga

shrub

GM 10

Dense Psuedotsuga-Acer-mixed

GM 11

Semi-open

Pseudotsuga-Symphoricarpos-mixed

GM 12

Semi-open

Pseudotsuga-mixed

GM 13

Open Pseudotsuga-Artemisia-Symphoricarpos~mixed

GM 14

Open Pseudotsuga-Prunus-mixed

GM 15

Open Pseudotsuga-Artemisia-mixed

shrub

GM 16

Open Pseudotsuga-Juniperus-mixed

shrub-rock

GM 17

Semi-open

GM 18

Open Pseudotsuga-Populus-mixed

shrub

GM 19

Dense Pseudotsuga-Populus-mixed

shrub-rock

GM 20

Scattered

GM 21

Low Populus-mixed

GM 22

Medium

Populus-mixed

shrub

GM 23

Scrub Populus-mixed

shrub

GM 24

Medium

Populus-Pseudotsuga-Prunus-mixed

GM 25

Medium

Populus-Carex-Lathyrus-mixes

GM 26

Carex-Agropyron-mixed

shrub

shrub-rock

shrub
outcrop
shrub

shrub

Pseudotsuga-Populus-Symphoricarpos-mixed

Pseudotsuga-Populus-Artemisia-mixed

shrub

shrub-rock

shrub

shrub

grass-scattered

Fig. 5. Vegetation units, Green Mountain
information presented in Appendix).

shrub

shrub

study area

(supplementary

�-235-

GREEN MOUNTAI

Fig. 5.

N

SCALE

o

I km

�-236-

GREEN

Fig. 6. Major vegetation types
and blue grouse use areas, Green
Mountain study area.
~Approximate
location of
'( breeding territories
Approximate boundary of

[[IJ] breeding territories

~: Brood concentration areas

I~

N

SCALE

o

MOUNTAIN

�-237-

types and some units were distinct, most units were not, with the majority
of units gradually integrating into adjoining units.
In some cases, the
transition was so gradual as to warrant establishment of a separate unit.
When the transition zone was more abrupt, the boundary line was placed midway
through the area of overlap.
Thus, admixtures of adjacent units were
frequently present along their boundaries.
"Islands" of dissimilar vegetation within a larger, individual unit were
usually too small «.5 ha) for recognition. Unless the "islands" were of
known importance in terms of grouse utilization, they were ignored.
Such
"islands" of vegetation that were delineated on the map include portions or
all of the following unit~: Artemisia-Symphoricarpos-mixed
shrub-scattered
Pseudotsuga, scrub Populus-mixed shrub, and low Populus-mixed shrub.
Characteristics

of Breeding Areas

Blue grouse males on the study area were territorial and defended chosen
territories throughout the breeding season (mid-April to early July).
All
territories (22) located from 1975-78 were marked on aerial photos.
Territory
locations were subsequently plotted on the vegetation map to facilitate their
description in relation to vegetative features of the area (Fig. 6). Delineation of territorial boundaries was limited to 9 territories where the resident
male was banded, and observed 5 or more times within the same breeding season.
Otherwise, only an approximate location of the territory was determined.
No territory was located completely in anyone vegetation unit as portions of
at least two and up to four units were encompassed within the boundaries of
all territories on Green Mountain.
Breeding territories were primarily located
where the semi-open and dense Pseudotsuga units (Fig. 5,_ units 7 through 12)
integrated into typically more open units such as the Artemisia units (Fig. 5,
units 1, 2 and 4) and the open Pseudotsuga units (Fig. 5, units 13 and 20).
Ten of 22 territories were associated with the semi-open Pseudotsuga-Popu1usSymphoricarpos-mixed
shrub (unit 17) and Artemisia-Symphoricarpos-mixed
shrubscattered Pseudotsuga (unit 2) vegetation units.
The latter unit (2) was
made up of many small openings not all of which were depicted on the vegetation
maps.
Features found in common among all territories examined include: (1) some form
of tree cover, (2) shrub thickets, (3) edges, and (4) some degree of openness
in both the canopy and understory cover.
Stands of Douglas-fir were the major
vegetational component on 21 of 22 territories, though aspen was frequently
present but in small amounts.
The only territory void of Douglas-fir was
associated with an aspen-sagebrush type. This does not imply that grouse
selected Douglas-fir over other species of trees for breeding habitat, but
reflects the high incidence (70% of canopy coverage) of Douglas-fir in the
overstory.
Aspen appeared to be an important part of breeding habitat.
It was much less
abundant in the overall tree cover than at sites where territorial males were
observed.
This was attributed to the fact that (1) territorial males were
most frequently found while performing their displays, and (2) aspens occurred
along the edges of coniferous stands and openings which were the same areas
preferred by males for displaying.

�-238-

Territorial males were seldom found more than 25 meters from an opening and
avoided areas with dense canopy and understory cover except along the edges.
During early to mid-April when most ground cover was beneath the snow, the
birds spent the majority of time in trees utilizing small, dense clumps of
conifers in an otherwise open habitat.
As phenological changes progressed
and the birds reverted to ground dwelling habits, shrub thickets became
important during resting and feeding activities while trees were primarily
used as escape cover and for roosting.
Some males performed their displays
considerable distances (&gt;50 m) from coniferous cover and therefore relied
on nearby shrub thickets for escape cover. Areas characterized by an
interrupted mixture of low growing vegetation (grasses and forbs), bare
ground, stumps, logs, and patches of shrub cover were preferred over areas
with continuous or homogeneous ground cover.
Whether shrub cover was essential in site selection of territories is uncertain. At the time of territorial establishment, most shrubs are totally or
partially snow covered.
Therefore, possibly only tree cover and relative
amount of open space nearby are important in site selection.
Furthermore,
since blue grouse have been documented to breed in a variety of habitat
types, this suggests that structural characteristics of the vegetation and
not species composition is an important factor in breeding habitat selection.
Characteristics

of Nest Sites

Eight nests were located from 1976 to 1978. Each nest site was described
in
detail as to aspect, slope, altitude and vegetative cover. All nests had
some type of cover immediately above and surrounding the nest. The cover
was mainly in form of shrub clumps.
One nest was partially protected by a
log, while another was adjacent to the trunk of a Douglas-fir.
Two nests
were found under sagebrush, two beneath saplings of Douglas-fir, one at the
base of a Douglas-fir (approx. 6 m high), two in the center of a sagebrushsnowberry clump, and one in the center of a serviceberry-sagebrush
clump.
Maximum distance from the nest to the nearest tree was 25 meters.
Height of
vegetation immediately above the nest averaged 81 cm and ranged from 56 to
104 cm. General appearance of the habitat around the nest was open to semiopen.
Of the 8 nests located, 4 were found in transition zones (edges) between the
following units: (1) dense Pseudotsuga~Symphoricarpos-mixed
shrub and ArtemisiaAmelanchier-mixed
shrub-scattered Pseudotsuga, (2) dense PseudotsugaSymphoricarpos-mixed
shrub and semi-open Pseudotsuga-Populus-Syrnphoricarposmixed shrub, (3) semi-open Pseudotsuga-Populus-Symphoricarpos-mixed
shrub and
Artemisia-mixed shrub-scattered Pseudotsuga, and (4) semi-open PseudotsugaPopulus-Symphoricarpos-mixed
shrub and Artemisia-Symphoricarpos-mixed
shrubscattered Pseudotsuga.
Two of the other 4 nests were in the ArtemisiaAmelanchier-mixed
shrub-scattered Pseudotsuga vegetation unit. One nest was
found in the Artemisia-Syrnphoricarpos-mixed
shrub-scattered Pseudotsuga unit
and the other one was located in the scattered Pseudotsuga-Populus-Artemisiamixed shrub-rock unit. Nest sites were in the same vegetation units selected
for breeding. However, only 3 nests were situated within the boundaries of
a territory.
All nests were between 2500 and 2740 meters elevation.
Sample size was
probably inadequate to evaluate the importance of slope and aspect; however,

�-239-

neither feature was believed to be significant factor in nest site selection.
Exposure of nests included all major compass directions, except south.
The
majority of nests faced north and east which are also the primary exposures
of the study area. Slope measurements at the nest site ranged from 6 to 25
percent.
Characteristics

of Brood Areas

Over 90 observations of blue grouse broods were made on Green Mountain from
1975 to 1978. At one time or another, broods were observed within most all
vegetation units shown in Figure 5. However, there were definite concentration areas as nearly 90 percent of all brood sightings were made in the open
and semi-open habitats on the east face of Green Mountain.
These areas are
depicted in Figure 6 and were the focal areas for evaluation of brood habitat.
Broods used areas where vegetation had interspersions of plants of various
life forms.
Such areas were generally along the edge of brush or free cover
and provided a high degree of concealment.
Almost invariably, any brood
encountered in the field was originally observed on the ground, but they
seldom ventured more than 30 meters from brush or tree cover and utilized
this type of cover for resting and for escape when disturbed.
There was no
evidence of the necessity for open water near brood cover. Only two sources
of open water occurred on Green Mountain and no broods were ever found at
these sites. Evidently the birds obtained their moisture requirements from
succulent foods and condensation of moisture on plants.
Summer brood range partially overlapped with breeding areas, but there
tended to be a greater diveristy of plant species, more ground cover (especially herbaceous),
and a mosaic pattern of more open vegetation units in
brood habitat than in breeding habitat.
The distribution of broods closely
approximated the distribution of aspen on the study area. Examination of
Figure 6 indicates that a major portion of the brood range encompassed the
semi-open Pseudotsuga-Populus-Symphoricarpos-mixed
shrub vegetation unit.
However, even in this predominantly Douglas-fir unit, broods were most
frequently found near or in the many small clearings (Artemisia-Symphoricarposmixed shrub-scattered Pseudotsuga) scattered throughout this unit. Characteristically, these openings were often bordered by aspen. At lower elevations
broods were found along the edge of the coniferous forest where aspen was
commonly present.
Vegetation units favored by broods in these areas included:
low Populus-mixed shrub, scrub Populus-mixed shrub, medium Populus-mixed shrub,
medium Populus-Carex-Lathyrus-mixed
shrub, ,Artemisia-Chrysothanrrlus-mixed shrub,
Artemisia-Amelanchier-mixed
shrub-scattered Pseudotsuga, open PseudotsugaPopulus-mixed shrub, and open Pseudotsuga-Artemisia-Symphoricarpos-mixed
shrub.
LITERATURE

CITED

Bendell, J. F., and P. W. Elliott.
1967. Behavior and the regulation
numbers in blue grouse.
Can. Wildl. Servo Rept. Ser. 4. 76 pp.
_____ , D. G. King, and D. H. Mossop.
1972.
blue grouse in a declining population.
1165.

of

Removal and repopulation of
J. Wildl. Manage. 36(4):1153-

�-240-

Blackford, J. L. 1958. Territoriality and breeding behavior
of blue grouse in Montana.
Condor 60(3):145-158.

of a population

1963. Further observations on the breeding behavior
population in Montana.
Condor 65(6):485-513.

of a blue grouse

Boag, D. A. 1966. Population attributes
Alberta.
Can. J. Zool. 44:799-814.

of blue grouse in southwestern

Braun, C. E. 1969. Population dynamics, habitat, and movements of whitetailed ptarmigan in Colorado.
Ph. D. Dissertation.
Colorado State Univ.
189 pp.
1971. Determination of blue grouse sex and age from wing characteristics.
Colorado Div. Game, Fish and Parks. Game Info. Leaflet No.
86. 4 pp.
Braun-Blanquet, J. 1951.
Germany.
pp. 58-66.

Pflanzensoziologie.

Springer-Verlag,

Wien,

Caswell, E. B. 1954. A preliminary study of the life history and ecology
of the blue grouse in west-central Idaho. M. S. Thesis. Univ. Idaho.
105 pp.
Harju, H. J.
grouse.

1974. An analysis of some aspects of the ecology of dusky
Ph. D. Dissertation.
Univ. Wyoming.
142 pp.

Harrington, H. D. 1964.
Denver.
666 pp.

Manual of the plants of Colorado.

Sage Books,

Hickey, J. J. 1955. Some American population research on gallinaceous
birds.
Pages 326-396 in A. Wolfson, ed. Recent studies in avian
biology.
Univ. of Illinois Press, Urbana.
479 pp.
Hjorth, I. 1970. Reproductive behavior in Tetraonidae,
reference to males.
Viltrevy 7(4):183-596.

with special

Hoffman, R. W. 1976. Population dynamics and habitat relationships of blue
grouse.
Colorado Div. Wildl. Job Prog. Rept. Fed. Aid Proj. W-37-R.
April 1976. p. 135-152.
1977. Population dynamics and habitat relationships of blue grouse.
Colorado Div. Wildl. Job Prog. Rept. Fed. Aid Proj. W-37-R.
April,
1977. p. 83-104.
Kuchler, A. W. 1949. A physiognomic classification
Assoc. Amer. Geographers 39(3):201-210.

of vegetation.

1955. A comprehensive method of mapping vegetation.
Amer. Geographers 45(4):404-415.

Ann.

Ann. Assoc.

Martinka, R. R. 1972. Structural characteristics of blue grouse territories
in southwestern Montana.
J. Wildl. Manage. 36(2):498-510.

�-241-

Medin, D. E. 1962.
herd.
Colorado
Proj. W-l05-R.

An ecological investigation of the Cache la Poudre deer
Dept. Game and Fish.
Job Completion Rept. Fed. Aid
July 1962. p. 187-204.

Mussehl, T. W. 1960.
Blue grouse production, movements, and populations
the Bridger Mountains, Montana.
J. Wildl. Manage. 24(1):60-68.
Nelson, R. A. 1969. Handbook
Tucson.
331 pp.

of Rocky Mountain

plants.

in

Dale Stuart King,

Redfield, J. A., and F. C. Zwickel.
1976. Determining the age of young
blue grouse:a correction for bias.
J. Wildl. Manage. 40(2):349-351.
Rogers, G. E. 1968. The blue grouse in Colorado.
Parks.
Tech. Publ. No. 21. 63 pp.
Stirling, I., and J. F. Bendell.
grouse.
Syesis 3:161-171.
Weber, W. A.
Boulder.

1970.

1972. Rocky Mountain
437 pp.

Welch, D. S. 1948. Limnological
New York.
381 pp.

Colo. Div. Game. Fish and

The reproductive

flora.

methods.

Colorado

behavior

Assoc. Univ.

McGraw-Hill

of blue

Press,

Book Co., Inc.,

Zwickel, F. C. 1965. Early mortality and numbers of blue grouse.
Dissertation.
Univ. British Columbia, Vancouver.
153 pp.
1972. Removal and repopulation of blue grouse
population.
J. Wildl. Manage. 36(4):1141-1152.

Ph. D.

in an increasing

1975. Nesting parameters of blue grouse and their relevance
populations.
Condor 77(4):423-430.

to

_____ , and J. F. Bendell.
1967. Early mortality and the regluation
numbers of blue grouse.
Can. J. Zool. 45(5):817-851.

of

_____ , and A. N. Lance.
1966. Determining
J. Wildl. Manage. 30(4):712-717.

the age of young blue grouse.

�· APPENDIX

�-243-

DESCRIPTIVE
Legal
Elevatlonal
Range:

SUMMARY OF VEGETATION TYPE !iA.P. GREEN HOUNTAIN - BLUE GROUSE INVESTIGATIONS
Description:
T25. RSOW. Sections
2. 3, la, 11
County:
SucmiL...
2500-2865
Cl
Surface
Area:
~
Aerial
Photo
So.
U.S.F.S.
Ft6CN 10-3-72
AND SUPPLEMENTARY
FEATURES

VEGETATIVE CLASSIFICATION

VEGETATION
Classification

and

1/

Physiognomic

GM 1

GM-2

GM3

Qi-4

GM-5

BziOztEIp

Bz1DdElp

BziOzi

BziDszpElp

EziOzpElp

Features

BzpCl1Dzp

UNIT NUMBER
GM-7

GM-S

EmcliDzp

Erne

GH-IO

GM-II

GH-12

cx-r )

[rnliD •.p Ez p Emi D'Lz p

EmiDzp

Em i De p

ElpBsi.I)zi

GM-9

Exposure

(Degrees Azimuth)

38-85

36-46

45-180

350-90

160-180

180

340-35

125

310

28

20-32

315

20-')0

Gradient

(Percent)

18-30

20-25

5-17

5-25

25-27

10-12

14-28

10

10-20

17

18-25

25-35

17-2~

14.7

4.5

10.0

3.S

2.2

18.3

.7

7.2

.7

4.0

2.9

Area

(Hectares)

Plant Species
List 11
Woody
Trees:
Acer glabrum
Juniperus
vlrginiana
Populus
tremuloldes
Pseudotsuga
menzlesii
vood Shrubs:
Acer glabrum
Amelanchier
a1n1£011a
Arctostaphylos
uva-urs!
Artemisia
cana
Artemisia
tridentata
Ceano thus velutinus
Cercocarpus montanus
Chrysothamnus
spp.
Ho Lod Ls cus dUOlOSUS
Juniperus
communis
Juniperus
vlrginiana
NahonLa repens
Pachystima
myrsinices
Populus tremuloides
Prunus virginiana
Pscudo t s uga mcnetes i 1
Purshia tridentata
Rhus trilobata
Ribes spp ,
Rosa acicular
Is
Rubus deliciosus
Sambucus racemosa
Shepherdia
canadensis
Symphoricarpos
oreophi1us
Tetradymia
canescens
Vaccinium
spp.
Forbs:
Achillea
lanulosa
Anemone spp.
Antennaria
epp ,
Aqullegia
caeru1en
Arabis
spp.
Arnica
spp.
Artemisia
frigida
Artemisia
ludoviciana
Aster spp.
Astragalus
spp.
Balsamorhiza
sagittata
Calochortus
gunnisonii
Campanu1a rotundifolia
Castelleja
spp •
Chaenactis
doug1asii
Chenopodium spp •
Cirsium
spp.
Clematis
columbiana
Clematis
hirsutissima
Comandra umbellata
Crepi.s spp.
Cryptantha
spp.
Cynogo1ossum of ficinale
Delphinium
spp ,
Descur at.nfa spp.
Dodecatheon
pulchellum
E.bilobium spp.
Erigeron
spp.
Eriogonur.; spp.
Erysimum spp.
Pr-aga r-La cpp .
Frasera
speciosa
Galium borea1e
Geranium fremontii
HapLopappus spp ,
Heuche r a spp.
Helenium hoopesii
He1ianthel1a
qudnquene rv f s
Ipomopsis
agg regat;a
Lathyrus
spp.
Linum lewisii
Lithospermum
spp •
Lupinus
argentius
Mertensia
spp.
Oxytropis
tippo
Penstemon
spp.
Phacelia
spp.
Phlox spp ,
Potentilla
spp.
Primu1a spp •
Pseudocymopterus
montanus
Pulsatilla
patens
Saxifraga
bronchia lis
Senecio
spp.
Smilacina
spp.
So lidago
spp.
Taraxacum officinale
Tragopogon
dub ius
ntal1c:trum
tippo
vto La spp.
Graminoids:
Agropyron
spp.
Bromus $PP.
Carex spp.
Deschampsia
caespitosa
Fes cuca spp.
Iris
missouriensis
Koleria
gracilis
Oryzopsis
hyeeno Ldea
Phleum pratense
Poa spp.
Sitanion
longifolium
Stipa
spp.

co;rrsoc'!:.l COy

see

COY

41.9

roc

COy

See

COy

FLORISTIC CHARACTERISTICS
Soc
COy
See
COy
Soc COy

+
,

+-1
4

+

1
1

+
1-2

+
2-3

1-3

3-4

1-4

3-4

+
1-2

1-2

1-2

1
+-1

+-1
+-1

+-1
+-1
+-1

1-2
1
+-1

+
1

+-1

+-1

1-2
1
1-2

1
1-4

1-2

+-1

2-3

+

3
1-3

1-2

1-3
1-3
1
1

+
+-1

1-2
1
1

+-1

1

+-1
+-1
+-1

+-1

+

2-3
1

1
1
1-2
2

1-2

1-3 +-1

1
1-2
1

1
2-3
1
1

1-2

1-3

+-1
,

1-4

+-1
4

Soe

1-2

3

COy

Soc

COy

+-1
3-4

3 +-1
1-2 3-4

1-2
1

+ 1
1-2
1

1-3

according

to

Cov

+

1

+-1

1-2

+
+-1

2
1-2
1

1-2

2-3
1-2 +-1
+-1

+-1
+-1

1-3
1

+

1-2

+-1

2-3
1

+-1
1

2-3
1
2

1-2

1-2
1

1-2

1-2

+-1

1 3

+

+-1

1-3

1-3
1-2
1
1

+
+

+-1

1-3
1
1-2
2-3

2

+-1
1

1-3

See

3 +-1
1-4
2

+
1-2
3

1

1-2

1-2

2-3

1-3

1-2

+-1

1-2

-+-1

1-3
1
2

+
+
+-1

+-1

+
1

1
+-1

1-3
]

+-1
+-1

+

+-1

+-1

+-1

+-1
+-1
1-2

+-1
1-2
1-2
+-1
+-1

1

1-2

1

1-2

1
1

+-1

+-1
1-2
+-1
+-1
+-1

+-1

+-1

1-2

1 +-1
1-2
+-1
+-1

1-3
1
3

1-2

1-2

+-1

+

+-1
+-1

1-2
1

+ 1

+-1

1

2-'3

1 2 + 1

+-1

1-2

1-2
+-1

1

+-1

1-2
1-2

1-2
1-2

2

1 2 + II+-1
2-3
1

+-1
+-1

2-3
1-3
1-2
1
1-2

+-1

+-1

1-2

+-1

1-2

+-1
+-1

3

+-1
+-1

1

+-1

+-1

1

+-1

1

+-1

+-1

+-1
1-2
1
1

1-2
1

+-1

+-1

1-2

+-1

1

+··1

+-1

+-1

+

+-1

1-2
3
1

1-3
3
1

1-3

1-3
1

+-1
1

1-2
3

+-1
1

1-2
1-3
1

+
1-3

+-1

1
1-3

+-1

1-2

+-1

+-1

+-1
+-1
+-1

+-1

2
1-3
1

+-1

1-2

1
+-1

+-1
1-2 +-1
3

2

+·-1

2

1-2

1
+-1

1-2
1-2 "'-1
1

+-1
+-1

+-1

+-1

+'
+

1-2

+-1

+
+
+-1
+-1

+
1-2

+-1

+-1

+-1

+-1
+-1

+-1

+
+
1
1
1-3
1-2
1
1
1

+

1-3

+-1
+-1

+
1-2
+-1

+
+-1

1-3
2
1
1

+-1

+

+-1

+-1

+-1

+

1-2
1
1-2

+-1

2

+

+

1-2

1
1-2
2
1
1-2
2

1-2
+-1
+-1

+-1
1-3
2
1

+
+-1

1-2

1-2

+-1

1-2

+-1
+-1

+-1
+-1

+

1

+-1
+-1

+-1
+-1
+-1

+-1

&gt;-1

+-1

1
1-2

+
+

+-1

1-2

+-1

+

+-1

1

+-1

+-1
+-1

+-1

+

+

+

+-1

+

1-2
+-1
2-3

1-3
1
1-3

+-1

1-2
+-1
1

+-1
2-3

1-2
1
1-3

+-1

1-2
1-2
1-2

+-1

(1955).

the

1-2

+

1
1-2
1-2

1-2

1-2
1

1-2
+-1
1-2

1-3

1-2

1
1-3

~2

+-1

1-2

+
+-1

1-2
2-3

+-1

1-2
1-2
1-2

+

1-2
1
1-2

+
+e-I

+-1

+-1

1-2

1-3
1
1-3

1-2

+-1

+-1

2 3

1-3

+-1

+-1

+-1
+-1
1-2

1 2 +
1-2
1-3 +-1

1-2
1-2

1-3

+-1

1

+-1

+-1

1

+-1

+-1
+-1
2-)

1
1 +-1
1-3 2-3

+

1
1-2
+-1

1

+-1

+-1

1 2 +-1
1 +-1
1-2 1-2

Kuchler

phys Lognced c formulae

reads

left

to

right

with

the

1
1
1-3

+-1
+
1-2

2
1
1-3

1

+-1
+-1

1-2
1
1

+-1

1-3

+

1-2

+

most

ccnsp t cucus

+-1

1

+-1
+-1

+-1

+-1

-------------1/ Assigned

Soc

1-2

+-1
+-1

1-2

+-1

1-2
1
1

COy

1 2

1-4

1-2

+-1
+-1

Soc

3
1-3

2-3

+
1
1
+-1
+-1

COy

1-2

+
+

+

Soc

type

p.Laced at

the

beginning.

�-244DESCRIPTIVE
SUMMARYOF VEGETATION TYPE HAP. GREEN t«)UNTAIN - BLUE GROUSE INVESTIGATIONS
LeBa1 Description:
T25. aBOW. Sections
2. 3. 10, 11
Count.y: ~
Range:
2500-2865
m
Surf3ce
Area:
181. 3 be
Aerial
Photo
No.
U.S.F .5.
F16CN
10-3-72

Elevational

VEGETATIVE CLASSIFICATION

Classification
Physiognomic

and

VEGETATION
GM-19

GM-14

GH-IS

GM 16

GK-17

GK-lS

EmpDszp

ElpRzpDzp

EmlpDszp

EmizpDlzp

EmOlnpBzp

£miDIp

35

90-190

65-120

18-46

35-90

2-20

UNIT NUMBER
GM-20

GM-22

GH-23

GM-24

GH-25

CM-26

DcczpHGlc

DlczlHGlc

DmpEmpElspDszp

DmiGlpHl{'

GlpHlpDsp

25-70

20

30-35

30-40

322-72

18-20

11-22

6-22

2.5

GH-21

DzpBzpEmpDlp 61cszpHGlc

1..1

Exposure

(Degrees

Gradient

(Percent)

Area

Features

AND SUPPLEMENTARY FEATURES

Azimuth)

Plant Species
List3/
Woody
Trees:
Acer glabrum
Juniperus
virginiana
Populus
tremuloides
Pseudotsuga
menziesii
Woody~Shr~u~bs~,
Acer glabrum
AmeLanch i.e r alnifoUa
Arctostaphylos
uva-ursi

1.5

5.3

.9

(Hcc::ares)

35

12-25

17

Cov2/Soc2/

COY Soc

COY Soc

17-30
23.0

COY Soc

34

15-25

3

1-2

1-2

15.9

4.3

.4

1-3

3
1-3

2-3

4
1-3
3-4
1-4
3
1-3
~~~--~--~~~~--~--~----~~~~~~~~~--------------------------------------------------1-2
1-3
+-1
+-1
2-3
1-2
1-2

5-16

2.9

FLORISTIC CHARACTERISTICS
Cov
Soc
CoY Sac
COY

3

2

9-15

Soc

1-2

.•..,

10-20

1.

2.6

2.9
COy

20-45

305-10

+-1
1-2
+-1

.•..,
.•..,
.•..,

.•..,

1-3
3

1-3 .•..1
Artemisia
cana
1-3
1-2
1-3
1-3
1-2
1-2
1-3
Artemisia
tridentata
2
Ceanothus
velutinus
2
1-2
Cercocarpus
montanus
1
Chrysothamnus
spp.
+-1
+-1
.•..,
1
Holodiscus
dumosus
1
1
.•..
,
1-2
Juniperus
communis
1
+
+-1
1
+-1
Juniperus
virginiana
1
+-1
1
+-1
2-3
Mahonia repens
+-1
+-1
+-1 . 2-3
+-1
+
+-1
2-3
+-1
.•..
,
+
Pachystima
myrsinites
3
1
Populus
tremuloides
2-3
1
2-3
2-3
Prunus virginiana
+
1
1-2
1-2
1
1
1
1-2
1
Pseudotsuga
menz1esi1
.•..
,
1
1
Purshia
tridentata
1-3
1-3
+
Rhus tdlobata
1-2
2-3
+
+-1
1
+-1
1
1
Ribes spp ,
+-1
+-1
.•..,
1
Rosa acieularis
+-1
2
+-1
Rubus de Hc tosus
1-2
Sambucus racemosa
+
2
Shepherdia
canadensis
+-1
1-2
1-2
1-2
1-2
1-2
+-!.
1
S)'!lIphoricarpos
oreophi1us
1-2
1-2
Tetradymia
cane scens
Vaecinium
s pp ,
+-1
Forbs:
1
+-1
+-1
1-2
+-1
1-2
+-1
Achillea
ranutcsa
1
+
Anemone spp .
2-3
1
+-1
2-3
2-3
+-1
2-3
Antennaria
spp ,
+
+
+
1
Aquilegia
caerulen
+
1-2
+
Arabis
spp ,
+-1
Arnica
spp ,
+-1
Artemisia
frigida
Artemisia
1udoviciana
+-1
+
+-1
+
+-1
Aster
spp.
2-3
1
+-1
2
+-1
2
+-1
Astragalus
spp .
+-1
1-3 +-1
1
1-2
+-1
1-2
1-2
+-1
+-1
1
Balsamorhiza
sagittata
+
1
+
1
1
Calochortus
gunnisonii
+
+
+
1-2
+
1
Campanu1a rotundifolia
+-1
1
+-1
1
+-1
+-1
Castelleja
spp .
Chaenactis
douglasii
+
+-1
Chenopodium
spp.
+-1
1-2
1
+-1
+-1
1-2
Cirsium
s pp .
+-1
1-2
Clematis
columbiana
Clematis
hirsutissima
+-1
Comandra umbel1ata
+-1
+-1
Crepis
spp.
+
+
Cryptantha
s pp ,
Cynogolossum
offie1nsle
+-1
Delphinium
spp.
Descurainia
spp.
+
Dodecatheon
pulche11um
+-1
+
Ebilobium
spp.
1
1-2
+-1
1-2
Erigeron
spp ,
+-1
1
+-1
3
1
3
Eriogonum
spp.
+
1
1
Erysimum spp.
Fragaria
spp.
Frasera
speclosa
+
+-1
+-1
2-3
Galium borea1e
+-1
+-1
+-1
Geranium
fremontii
+-1
Haplopappus
spp ,
+
Heuchera
spp.
Heleniuro hoopesii
Helianthe11a
quinquenervls
Ipomopsis
aggregata
1-2
1-2
Lat.hyrus
s pp ,
1-2
1-2
+-1
1-2
+-1
1-2
Linum lewisii
+
1
+-1
1
Lithospermum
spp ,
1
1-2
1
1-2
Lup:inus argent ius
+
1-2
Mertensia
spp .
+-1
+
2
Oxyrropis
spp.
1
+
1
+-1
Penstemon
spp ,
+-1
1.
Phacel1a
spp.
+
Phlox spp.
+-1
+
Potent.illa
spp ,
+-1
-+-1
Primula
spp .
ssecdccvecptervs
montanus
+-1
+-1
Pulsatilla
patens
2-3
+-1
Saxifraga
bronchlalis
+-1
+
+
Senecio
spp .
Smilaeina
spp.
+
Solidago
spp.
+-1
1-2
+-1
1-2
Taraxacum
officinale
+-1
1
Tragopogon
dub ius
Thalictrum
spp ,
Viola
spp.
Graminoids:

....,

.•..,

.•..,

.•..,

.•..,

.•..,

Soc

3-4

1-3

.•..,

1-2

+

.•..,

.•..,

Cov

Soc

3-4
2-3

1-3
1

2-3

1-2

+
1

2
1-2

+-1

1-3

+-1

1-2

.•..,

+-1
+-1

+-1

1

+-1
1

1-2
+-1

1-2
1

.•..,

+-1

+-1

1-2

+-1

1-2

+-1

+-1

1-2

+-1

2-3
+-1
+-1

.•..,

1-2

+-1

+-1
+-1
1-2

1

.•..,

.•..,

+-1

2-3

....,

.•..,

+
+-1
+-1

2-3
2-3

1-2

1-2

.•..,

1

1

1-2
1

1

.•..,
.•..,

+-1
+-1
1-2
2

1

+-1
+-1

4-5

+-1

.•..,

.•..,
.•..,

.•..,
.•..,

.•..,

Cov

COY Soc

See

+
+

••

+-1

1-2

2
1-2

+-1
+-1

1-2
1-2

+
+

+
+-1

�-245April

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Game Bird Survey

W-37-R-32

Job TitLe

Job No.~
21
Minimum Tillage Techniques for
Establishing Shrubs in Clump Plantings

Period

April 1, 1978 through March 31, 1979

Work Plan No.

Covered:

Personnel:

David C. Bowden

1979

(Statistician,

l

_

CSU) and Warren D. Snyder.

ABSTRACT
Spring measurements of weed competition showed a reduced effectiveness among
all treatments in spring 1978 for several reasons.
The impacts of the
previous spring 1977 hail on the loam soil sites was noted in dramatically
increased mortality.
Spring survival and winter die-back and mortality
information was obtained on all sites. Below normal precipitation conditions
persisted through the spring and summer of 1978 in the locality, which was
accompanied by a devastating outbreak of grasshoppers in the sandhills.
A
complete defoliation in June 1978, along with some consumption of the cambium
layer resulted in all sandhill sites. No fall growth or survival data were
obtained.
Analysis of weed competition, shrub growth, and shrub survival
data was completed by Dr. Bowden.
Preparations were begun on a final report.

��-247-

MINIMUM TILLAGE TECHNIQUES FOR
ESTABLISHING SHRUBS IN CLUMP PLANTINGS
Warren D. Snyder

P. N. OBJECTIVE
To evaluate ground cover techniques, pre-emergent herbicide techniques, a
combination of the two, and cultivation techniques for establishing shrubs
in clump p1antings in eastern Colorado.
SEGMENT OBJECTIVES
1.

To apply pre-emergent herbicide treatments to the Eptam~Tref1an
Casoron plots and to apply cultivation to the control.

2.

To measure and evaluate
tition among treatments

and

shrub survival, shrub growth and weed compeand soils in relation to weather variables.

METHODS

AND MATERIALS

Methods and materials used in this segment were summarized in previous
segments starting in 1975 and therefore will not be repeated here.

work

RESULTS AND DISCUSSION
Environmental

Measurements

Precipitation
Below average precipitation was recorded for 1978 in the study region.
The
only significant precipitation was received in May when approximately 2.75
inches was recorded.
This rainfall was not consistent over the entire area
and rangelands to the southeast of the Tamarack Headquarters remained dry
with little spring green up. The annual total for 1978, approximating 9.30
inches, was considerably below the 15 to 16 inch annual average for the
study region.
Grasshoppers
High densities of ~rasshoppers in combination with little green vegetation
in rangelands prompted high concentrations and rapid defoliation of the
shrubs on the Red Lion, Sharptai1 and 1-76 sites in the sandhil1s during
June and July 1978. Little green foliage remained by early July and by
August the bark and green cambium layer on new growth had been consumed.
Efforts were not made to control these hoppers because of (1) the lateness
in initial detection of the problem, and (2) the vast quantities of replacement hoppers in surrounding rangelands.
It is assumed that near total
mortality of all shrubs occurred as a result of the devastation.

��-249April

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Game Bird Survey

W-37-R-32
Job No.

22

Work Plan No.
Job Title

Upland

Period

April 1, 1978 through March

Covered:

1
---------------------------------

Game Publications
31, 1979

T.D.I. Beck, C. E. Braun, K. M. Giesen,

Personnel:

1979

R. W. Hoffman,

D. H. Nish.

ABSTRACT
Publications

planned

for and accomplished

under this job for Segment

32 are as

follows:
Beck, T.D.I., and C. E. Braun.
Condor 80:241-243.

1978.

Weights

of Colorado

sage grouse.

Braun, C. E. 1978. Effects of changes in hunting regulations on sage grouse
harvest.
Proc. Ann. Conf. Central Mtns. and Plains Section, The Wildl.
Soc. 23:Abstract.
Braun, C. E., D. H. Nish, and K. M. Giesen.
1978. Release and establishment
Nat. 23:661-668.
of white-tailed ptarmigan in Utah.
Southwest.
Giesen, K. M. 1978.
Auk 95:761-762.

Egg retrieval

by incubating

Giesen, K. M., and C. E. Braun.
1979.
juvenile white-tailed ptarmigan.

white-tailed

ptarmigan.

A technique for age determination
J. Wildl. Manage. 43:508-511.

of

Hoffman, R. W., and.C. E. Braun.
1978. Characteristics
and status of ruffed
grouse and blue grouse in Colorado.
Western Birds 9:121-126.
Hoffman,

R. w.

Prepared

by_C-=J_·~_~...:::;-:=:-:-=·td=~~""",-,--:-·
cV-_,,",,-&gt;£=a:::...tz:.=~--!..£=-__
Howard D. Funk
Section Chief
Small Game Research

1978.

Pikes Peak ptarmigan.

Colo. Outdoors

27(6):12-13.

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                  <text>July,

-1-

JOB PROGRESS

State of

REPORT

COLORADO
--------~~~~~---------Deer-Vehicle

W-125-R-5

Project No.
Work Plan No.
Job Title

1979

1

Job No.

Deer Vehicle Accidents

Period Covered:

Accident

Investigations

1, 2, 3, 4, 5, and 6

Statewide and Methods

and Devices

July 1, 1978 - June 30, 1979

to Reduce
Them

Personnel: James D. Fleming, Kenneth R. Kincaid, Dale F. Reed, Area
Supervisors and Wildlife Conservation officers,

ABSTRACT
Methods, devices, or structures related to reducing the number of deer-vehicle
accidents were evaluated or experimentally tested.
These were highway
lighting, underpasses, overpasses, 2.44-m fences and one-way deer gates, and
deer guards. Forty deer were killed in the highway lighting study area,
twenty-two and eighteen with lights off and on, respectively.
Background and
target luminance measurements were taken at 27 of the 39 accident sites.
The
overall calculated mean indices were 1.845 ± 1.538 (SD) and 1.754 ± 0.849 (SD)
for transition and full lighting respectively.
No major changes occurred with
underpass use. The experimental procedure for testing a net arch at the Dowd
deer overpass was completed for two seasons.
A combined total of 43 and 18
crossings occurring during control (without net arch) and variable (with net
arch), respectively, were reviewed during video tape replay. Although selected
behavioral differences were noted during the seasons, the combined data have
not shown significant differences at the 95 percent confidence level.
The
average reduction of deer-vehicle accidents adjacent to six 2.44-m fences was
77.5 percent.
No data were obtained on deer guard prototypes.

��-3-

DEER-VEHICLE ACCIDENTS STATEWIDE
AND METHODS AND DEVICES TO REDUCE THEM
Dale F. Reed

P. N. OBJECTIVE
Locate and examine potentially critical deer-vehicle accident areas in
Colorado and recommend methods or structures which may reduce deer-vehicle
accidents in these areas.
Subsequently, measure the effects of methods
recommended and investigate deer responses to various experimental structures.

SEGMENT OBJECTIVES
1.

Monitor

potentially

2.

Test the effects of highway
vehicles.

3.

Evaluate

responses

4.

Evaluate

deer responses

5.

Evaluate deer responses
highways.

6.

Evaluate

responses

critical

deer-vehicle
lighting

Monitoring

areas statewide.

on number of deer killed by

of deer to highway underpasses.
to highway overpasses.
to 2.44-m

(8-ft) fencing adjacent

to

of deer to guards.

DESCRIPTION
1.

accident

potentially

OF AREAS

critical deer-vehicle

accident

areas statewide.

Generally,.Myers
(1969), Pojar (1972a) , and Woodard (1973a, 1973b,
1974, 1975, 1976a) described the areas most intensively monitored.
The completion of deer-vehicle accident reports (includes location)
has been required statewide since 1974. Hence, the potential existed
for the detection of new problem areas statewide.
2.

Evaluating the effects of highway lighting
by vehicles.

Highway

82 - Jammeron

Pojar et al. (1975) described
3.

Evaluate

responses

on number of deer killed

Curve

this area.

of deer to highway

underpasses.

The Vail deer underpass (Reed et al. 1975), a small structure under
1-70 east of Avon, two concrete box underpasses under 1-70 east of

�-4-

Eagle, twin bridges structure under 1-70 west of Eagle, and twin
bridge structures east of Rifle were monitored for deer use or
behavioral responses.
4.

Evaluation

of deer responses

to highway

Dowd Junction
Reed
5.

(1976a) described

Evaluate

deer responses

- 1-70

the area of the Dowd deer overpass.
to 2.44-m

Highway
The Highway

overpasses.

(8-ft) fencing adjacent

82 - Diamond

S

82 study area has been described

Highway

by Pojar

(1972b).

82 - Carbondale

This study area was described

Interstate

to highways.

by Reed (1975).

70 - Avon, Edwards,

Vail and Eagle

The Interstate 70 Avon and Edwards study areas have been described
by Woodard (1973b), and the Vail study areas by Reed (1971), and
the Eagle study areas by Reed (1974).
6.

Evaluating

responses

of deer to guards.

Trail Gulch-Dotsero
Reed et al. (1974) described

METHODS
1.

Monitoring

potentially

this area.

AND MATERIALS

critical

deer-vehicle

accident

areas statewide.

Methods have been described by Pojar (1972a) and Woodard (1976a).
Because no new critical areas were delineated, methodology was limited
to tabulation of statewide Wildlife Conservation Officer kill reports.
2.

Evaluating the effects
by motorists.

of highway

lighting

on number of deer killed

Methods were described by Woodard (1976b).
Luminance measurements
were taken at 27 kill sites.
Luminance values (foot-Iamberts) were
recorded with a spotmeter, Spectra Model UBA, and were taken on a
target at the kill site and on the background as viewed by an
approaching motorist.
The target was a taxidermy mount of the left

�-5-

half (transverse section) of a female mule deer and the actual
spotmeter recording area was a 26-cm diameter circle midway between
the shoulder and hip. Spotmeter readings were taken from a height
of 1.3 m and a dLs t ance of 15 m , For background luminance readings
the spotmeter was placed 60 m from the kill site, resulting in a
target area diameter of 102 cm. The taxidermy mount was removed
during this background measurement.
Measurements were taken between
0400 and 0600 MST to minimize traffic interference.
Background
luminance (Lb) and target luminance (Lt) measurements were transformed
into visibility indices (VI) by the following equation (Gallagher and
Meguire 1974):

VI
(DGF)

and where DGF

where C

L~Lb
Lb

Relative contrast
sensitivity for
the recorded
background luminance

= Disability glare factor = 1.0

RCS values were obtained from standard tables (Gallagher and Meguire
1974). VI values can then be compared to standards established for
urban lighting requirements.
These values also provide a better
description of the visibility of deer in the lighted portion of the
study area; which, in light of the poor horizontal uniformity ratio
in the lighted section (&gt;15.0:1), should allow a more refined analysis
of the effectiveness of highway lighting.
3.

Evaluate responses

of deer to highway underpasses.

Seven underpasses were monitored for deer use as described by Reed
(1971). Observations at the Vail deer underpass were continued
as described by Reed (1976b).
4.

Evaluation

of deer responses

to highway overpasses.

Methods were previously described by Reed (1976a) and Reed and Beck
(19.77). A net arch, similar to those used on pedestrian overpasses
was changed to control (without arch) or to variable (with arch)
every three days during the fall migration of 1978 and the spring
.migration of 1979. During both seasons the video housing was
positioned to cover the distant approach areas (approximately 20 m
beyond the bridge entrance and exit).
5.

Evaluate deer responses

to 2.44-m (8-ft) fencing adjacent

to highways.

Methods and materials have been described by Pojar (1972b) and Woodard
(1976c). Benefit-cost analysis followed procedures described by
Howe (1971). Costs for which values were available include initial
cost of 2.44-m fence and associated passage structures.
Estimates of
maintenance costs were not available.
Total cost was reduced by the
cost of the appropriate length of 1.07-m (42-inch standard right-of-way

�-6-

fence which was not built because of the 2.44-m fence).
Benefits
for which a value was available was the potential savings accrued
by a reduction in deer-vehicle accidents and related vehicle repairs.
Savings by reduction in personal injury and death were omitted because
of difficulties in obtaining reliable values.
No value was assigned
to deer. Therefore, benefit-cost analysis is essentially from a
highway department accounting stance. All future benefits were
discounted at an annual rate of 5.5 percent.
No monetary values
have been assigned for aesthetic considerations, either positive
or negative.
6.

Evaluating

responses

of deer to guards.

Methods were described

by Reed et al. (1974).

RESULTS AND DISCUSSION
Statewide Deer-Vehicle

Accidents

Two Division of Wildlife Regions, Southwest and Northeast, have taken
over the responsibilities. of maintaining their deer-vehicle accident
files. Area analyses of deer-vehicle kills will not be generated unless
requested because of manpower requirements.
Generally no new critical deer-vehicle accident areas were detected.
A
total of 3,365 deer-vehicle accident kills were reported in Colorado in
1978; an increase of 54.8 percent over 1977. Regional totals were
Southwest 1,148, Northwest 1,486, Northeast 490, and Southeast 241. By
March of 1979 some potentially critical areas had a higher than average
number of kills, likely because of the winter severity (greater snow
accumulation and lower temperatures).

Highway Lighting Evaluation
Deer occupied the highway lighting study area for eight weeks beginning
4 February.
Eight weeks of lights off and eight weeks of lights on data
(lights off and on for alternate weekly periods) were collected when deer
were in the study area. Although deer were in the general study area for
an extended period, most had not concentrated or begun to cross the highway
in high numbers until mid-February when the south-western exposures began
to "open up".
Snow accumulation adjacent to the study area at higher
elevations was considerably greater than in any previous years of study
(Table 1).
Forty deer were killed during an estimated total of 1,481 crossings (Table
2). The d8er crossings to deer kill ratio with lights off was less than
the ratio with the lights on 33.6 and 41.2, respectively.
However, by
chi-square analysis, there was no significant difference (P &gt; 0.50) between
these ratios.
In addition, the ratios are less than the combined ratio data

�-7-

Table 1.

Adjacent average snow depth, mean d~ily temperature, mean deer

counted and number of accidents in the lighting study, 1974 through 1979.

Adjacent
Average Snow
Depth
Year

(cm)I)

1974

51.0

1975

Mean
Daily
Temperature
(CO)

Mean Count

No. of Accidents

-3.5

27.1 en::71)

13

46.4

-2.8

40.3 (n=98)

22

1976

27.5

-1.7

0.3 (n=6S)

0

1977

8.0

-2.3

0

(n=5)

0

1978

34.9

-2.1

33.1 (n=53)

9

1979

61.8

-5.2

48.5 en=31)

40

Total

84

II

- Snow depth measurements taken at three sites northeast and above the
study area.

Table 2. Estimated deer crossings and total accidents in the Highway 82
lighting study area during 1979.

Off

On

Total

Estimated' crossings

740

741

1,481

Total accidents

22 .

18

40

Crossing/accident

33.6

41.2 ..
...........

�-8-

obtained in other years of study. The differences, however, are not
significant (P &gt; 0.05 and P &gt; 0.25 off and on, respectively) according
to chi-square analyses.
It is reasonable to postulate that crossings
per kill ratios may change over extended periods of time because of
the annual increases in traffic volume.
Background and target luminance measurements were taken at 27 additional
sites. The overall means of the visibility indices were 1.845 ± SD 1.538
and 1.754 ± SD 0.849
for transition and full lighting. respectively
(Tables 3 and 4). These visibility index means are near the level where
70 percent of the motorists can see a target at satisfactory separation
distance (Gallagher and Meguire 1974). To attain a level where 85-95
percent of the motorists can see a target at satisfactory separation
distance, a visibility index value of 2.6 - 3.6 is required.
Only 6 of
the 39 indices were within or above this range. Probably inherent in
this problem was the drab pelage of deer which, by eye, readily blended
with the lighted highway surface when in certain locations.
For example, accident 24 (Table 4) involved a target and background with
very low contrast despite its occurrence under full lighting.
At these
locations, just beyond the lamps and when the background was relatively
well lighted ( ~ 1.0 cd), it is estimated that an increase in horizontal
illumination would not substantially increase the contrast or likelihood
of motorist visual discrimination.

Highway Underpasses
Seven underpasses were checked for deer use during this segment.
Moderate
(30-90) to high (&gt; 90) numbers of deer passages occurred through two of
the underpasses, one located west of Vail and one east of Avon.
Extremes
of openness or tunnel effect were represented by these two structures.
The primary stimulus of a given underpass structure to approaching deer
may be termed the "openness effect".
Calculated as follows:
height x width

(or open-end
length

surface area)

the openness
effects of the Vail and Avon underpasses were 0.3 and 0.03
(metric measurements),
respectively.
There are several factors to consider
before relating openness effect to deer behavioral response.
Additionally,
any reasonable attempt to relate openness effect to deer .ise must consider
deer density and motivation at the structure.
This raises several questions,
e.g. does deer behavioral response and/or motivation change with changing
deer density (numbers of individuals and groups, and size of groups) and/or
with changing sex and age structure?
Only a few daily observations (n = 9) were made at the Vail deer underpass
in June and July of 1979. With 1974 through 1979 combined, 77.4 percent
of 319 groups of deer did not exhibit flight reaction when westbound
vehicles traveled by the area (Table 5). Likewise, of 1,021 instances

�-9-

Table

3.

Visibility

index measurements

from accident

sites in transition

lighting.

Relative
Contrast
Sensitivity
(Percent)

Background
luminance

Target
luminance

No.

(fL)

(fL)

1

0.0193

0.0253

0.3109

2.211'];/

0.1198

2

0.0104

0.115

10.0577

1.367 2/

2.3953

3

0.0655

0.0055

- 0.9160

5.525

- 0.8817

5

0.103

0.0105

- 0.898

7.3

- 1.142

6

1.480

0.0079

- 0.9947

27.7 'l:_'

- 4.800

7

0.124

0.0125

- 0.8992

8.23

- 1.289

9

0.043

0.0325

- 0.244

4.1

- 0.174

10

0.226

0.0030

- 0.9867

11.8

- 2.0285

11

0.330

0.00325

- 0.990

14.4

- 2.484

12

0.061

0.0094

- 0.846

5.25

- 0.774

13

0.235

0.0255

- 0.891

12.05

- 1.870

14

0.060

0.0048

- 0.920

5.2

- 0.833

15

0.285

0.004

- 0.986

·13.325

- 2.289

16

0.080

0.0165

- 0.794

6.2

- 0.858

18

0.086

0.130

0.512

6.5

0.580

19

0.180

O.Oll

- 0.939

10.4

- 1.701

21

0.180

0.026

- 0.856

10.4

- 1.551

22

(0.690)

0.284

- 0.600

20.31

- 2.134

23

0.690

0.012

- 0.9826

20.31

- 3.477

26

0.253

0.247

0.0236

12.53

- 0.0517

Contrast

Visibilityl'
Index

3/

-----------------------------------------------------------------------------

�-10-

Table

3. Visibility

lighting.

index measurements

from accident

sites in transition

(Cont.Lnued ),

Relative
Contrast
Sensitivity
(Percent)

No.

Background
luminance
(fL)

Target
luminance
(fL)

27

0.066

0.260

·2.9394

5.55

·2.842

28

0.279

0.136

- 0.5125

13.2

-1. 179

31

0.041

0.0056

- 0.8390

4.0

-0.5847

32

0.280

0.0091

- 0.9675

13.22

-2.228

33

0.400

0.0197

- 0.9508

15.85

-2.625

36

0.045

0.475

9.5556

4.25

7.075

Positive and negative
respectively.

Contrast

values indicate

frontlighting

V·1Sl
·b·l·
1/
1 ;t.tyIndex

and backlighting,

21
-

Derived

from data presented

lIThe background

by Technical

Committee

Report of the CIE 1972.

at this site involved a snow covered emergency lane and
right-of-way.
The snow was lost before measurement.
A value for similar
conditions (No. 23) was used.

�-11-

Table

4.

Visibility

index measurements

from accident

sites in full lighting.

Contrast

Relative
Contrast
Sensitivity
(Percent)

Visibility
Index

0.390

- 0.381

19.62

-1.303

1.100

0.209

- 0.810

24.44

- 3.449

17

0.240

0.470

0.958

12.2

2.036

20

0.197

0.069

- 0.650

10.95

-1. 240

24

0.185

0.178

- 0.0378

10.57

-0.070

25

0.237

0.455

0.9198

12.10

1.939

29

0.075

0.120

·0.600

5.95

0.622

30

0.500

0.110

- 0.780

17.67

- 2.401

34

0.420

0.140

- 0.6667

16.24

-1. 886

35

0.580

0.330

.- 0.4310

18.98

-1.425

37

0.620

0.270

- 0.5645

19.51

-1. 919

38

0.440

0.108

- 0.7545

16.63

-2.186

'39

0.225

0.480

1.133

11.77

2.323

Positive and negative
respectively.

values

frontlighting

and backlighting,

Background
luminance

Target
luminance

No.

(fL)

(fL)

4

0.630

8

1/
-

indicate

1/

�-12-

of vehicles traveling west when deer were near the underpass entrance
and the west bound lanes, 918 did not elicit flight reaction from 319
different individuals or groups (1-14 deer).
Twenty-three vehicles
elicited flight reaction from 1-50 percent of the individuals from
seventeen different groups ranging in size from one to ten deer.
Eighty-one vehicles elicited flight reaction from 51-100 percent of
the individuals from fifty-five different groups ranging in size from
one to eight. Previous to these observations it was hypothesized that
deer at distances of 15 to 30 m from moving traffic were frequently
caused to leave the area. Observations indicate that when vehicles stop
at or when bicyclists travel by the underpass, most deer exhibit immediate
and intense flight reaction.
Five groups of deer exhibited moderate to
intense flight reaction to unknown stimuli (where observer was unable to
determine potential cause).
Table 5. The number and percent of deer groups not exhibiting flight
reaction (Hediger 1950, Scott 1958), and the number and percent of
westbound vehicles not eliciting flight reaction at the Vail deer underpass
west of Vail.
Groups of Deer

Year

Numbers of Individuals
or Groups II Near
Westbound Traffic

Percent Not
Exhibiting
Flight
Reaction

Westbound
Number Traveling
West When Deer
Present

Vehicles
Percent Not
Eliciting
Flight
Reaction

1974

37

86.5

198

97.5

1975

67

71.6

251

89.2

1976

78

78.2

243

89.7

1977

59

86.4

148

92.7

1978

62

74.2

150

83.3

1979

16

56.2

31

64.5

Totals/Avg.

319

77.4

1,021I1

89.9

II
Number of deer in groups ranged from 2 to 14.

Twenty of 38 tracker-trailers

elicited flight reaction.

�-13-

The behavioral modes of 298 deer exits (leaving the underpass to the
south) were 49.7, 32.2, and 18.1 percent, trotting, walking, and bounding,
respectively.
Combined data 1974 through 1979 yielded a walk:trot:bound
ratio of 1.8:2.7:1.0 (Table 6). Wariness had been noted irrespective of
the mode of exit in many instances.
The predominant exit behavior of
trotting supports the continued reluctance of deer to use a structure
of this size and character (Reed et al. 1975).

Table 6.
underpass

The number of exits observed at the south end of the Vail deer
and the walk:trot:bound
ratio of exit behavior.

Year

No. of Exits
Observed

1974

35

4.0:12.5:1.0

1975

73

3.3:2.4:1.0

1976

55

1.8:6.3:1.0

1977

46

4.0:2.7:1.0

1978

70

0.7:1.4:1.0

1979

19

0.3:1.8:1.0

Totals/Avg.

298

1.8:2.7:1.0

Highway

Walk: Trot: Bound
Ratio During Exits

OVerpasses

The video time-lapse surveillance system was operated at the Dowd
experimental-net-arch
overpass during two seasons, fall of 1978 and spring
of 1979, for 36 and 27 consecutive nights, respectively.
A total of 61
crossings were examined during video replay (Table 7).

Number of Crossings
More crossings occurred under control than variable during each of the
seasons (Table 7). A test for independence shows that the difference
between the number of control and variable crossings during the alternate
three-day periods is not significant (X2 = 1.357, df = 3, P &gt; 0.50).
However, differences, as determined from a similar analysis (X2 = 19.599,
df = 4, P &lt; 0.005), in pre-experimental
data (three-day periods during
four seasons:
fall 74-75 and spring 75-76) are sufficiently great that
this statistical method is inapplicable.

�Table 7. The number of deer crossings and approaches examined on video replay at the experimentalnet~arch overpass and the crossings per approach (C:a) ratios for two migration seasons (Fall 1978 ~prtngs 197~)~

VARIABLE· .

CONTROL
Crossings
1978
Fall

25

Distant
.:
. 2/'
.. ..
..'
'1!
Approach- Approach- C:a

2

2

6.2;1

Crossings

Distant
Approach

Approach

4

5

6

.

.

TOTAL

C :a

crossings

0.4:1

29
I
•.....

"'I"

1979
Spring

18

8

12

0.9:1

14

Totals!Avg.

43

10

14

1.8:1

18

..........

."

4

2

2.3:1

32

9

8

1.1:1

61

......,.

l!Denotes a deer' that entered the overpass area not encompassed' by an entrance and exit trackbed
and that was oriented toward the structure.
~!Denotes a deer that entered either' the entrance or exit trackbed area without crossing the
overpass.

�-15-

Crossings:approach

Ratios

More crossings and approaches (Table 7) occurred during control (43 and
24 respectively) than during variable (18 and 17 respectively).
However,
variable crossings per approach ratios were not significantly (P &gt; 0.50)
smaller than that for control.
Generally, it would be expected that the
variable C:a ratio would be smaller if there were a greater reluctance
of deer to cross under a net arch (i.e. more reluctant animals either
approached and failed to cross, or made more than one approach before
crossing).
Any ratio difference, however, is diminished if, as was
discussed previously, the difference in the number of crossings is
considered inapplicable.

Duration

of Hesitations

and Crossings

The control and the variable (net arch) means of the combined seasons
(Table 8) of both the duration of hesitations and the duration of
crossings are not significantly different (P &gt; 0.20 and P &gt; 0.40,
respectively).
Generally, if the net arch was expected to result in
more reluctance to cross, then longer hesitations would be expected.
The means for the duration of hesitations for the fall of 1978 were
not significantly different (P &gt; 0.20). However, the variable mean was
significantly (P &lt; 0.05) less in the spring of 1979. Also, animals being
more reluctant in crossing under the net arch (variable) would be expected
to do so more hurriedly (more and/or faster trotting and bounding).
The
durations of crossings for the fall of 1978 were significantly (P &lt; 0.05)
greater, whereas the duration of crossings for the spring of 1979 were
not significantly (P &gt; 0.50) different.
Differences between the spring and fall migrations are equally perplexing.
The variable duration of hesitations and the variable duration of crossings
were both significantly (P &lt; 0.05 and P &lt; 0.05) greater in the fall of 1978
than in the spring of 1979, whereas the control duration of hesitations
and control duration of crossings were not significantly (P &gt; 0.05 and P &gt;
0.50 respectively) different.
Possibly the data of the two seasons should
not be combined.
The differences may be related to the observed and
postulated dissimilarities between the seasons, some of which are as follows:
Fall

Spring

First net arch experience

Potentially

Good physical condition
High wariness (hunting related)
Most mature females with
fawns at side
Maternal-fawn bond strong
No dawn activity
Very high motivation to
migrate (more direct movements
and less time consumption likely
causally related to increasingly
inclement weather)

Poor physical condition
Moderate wariness
Most mature females
parturient

second net arch experience

Maternal-yearling
bond weak
Dawn activity
High motivation to migrate (more indirect
movements and more time consumption
likely causally related to increasingly
inclement weather)

�Table 8. The mean duration
at the experimenta1-net-arch

(seconds) ·0£ hesitations
overpass.·

i~~

and crossings

during

, ··s.r~~~~·"

'"

control

and variable

p1/

TOTAL,

Hesitations

....

2/

Control

10. 2±12. 4- Cn=42)

16. 3±-19.4 (n=49)

13. 5±16. 7 (n=91)

Variaole

14. 3±14. 3 (n;'20)

7. 2±4. 6 (n=20)

10. 2±-lO.4 (n=47)

p

&gt;

0.20

Crossings
•....I
Control

7.2±12.4(n=25)

8.7;tlO.2(n=18)

s.oeu. 4 (n=43)

0\

I

p &gt; 0.40

24.2±19.8(n;'4)

Variable

1/
- Independent

2/

- Standard

.

t statistic.

deviation.

6 .4±B.2 (n=14)

10. 4±13. 3 (n=l8)

�-17-

Behavioral

Responses:Crossing

Ratios

There were more animals exhibiting behavioral responses (BR) and more
instances of behavioral responses per crossing (C) during variable than
control for most of the responses studied (Table 9). Although the ratios
for instances of behavioral response per crossing are larger for the
variable except for muzzle-to-ground,
they were not significant (P &gt; 0.50).
Therefore, differences in these behavioral responses may be due to chance
and not to the reluctance of animals in crossing under a net arch.

Conclusions
Based upon the above analyses, it does not appear that an important
of reluctance of deer to cross under a bridge net arch was reached.
statistical analyses used in this job will be reviewed and possibly
in the final report.

Responses

to Highway

level
The
expanded

Fencing

Six 2.44-m (8-ft) fences were evaluated as to the reduction of highway
deer kills after installation.
The range of reduction was 61.1 to 89.5
percent with a cumulative average of 77.5 percent (Table 10). Since
this evaluation is based upon year-to-year comparisonsa the cumulate
average changes each segment.
The reduction in kills adjacent to the
Avon, Edwards, Eagle, and Diamond S fences increased, whereas, it
decreased for the Vail and Carbondale fences.
The January-March winter
period of 1977 was exceptionally mild and may have resulted in fewer kills
at some of the fences.
Data from severe winters would not be discarded
from the sample, similarly, data from mild winters probably should not
be discarded, either.
No important changes occurred in the use of 28
one-way deer gates located in the Vail, Avon, Edwards, and Eagle fences.
Only minor changes would be shown in benefit:cost ratios previously
described (Reed and Beck 1977), if they were recalcula~ed for this
segment.
They will be presented in the final report.
A manuscript
draft titled "Methods of red~cing deer-vehicle a~~idents:benefit-tost
analysis" has been in preparation.

a

An acknowledged common fallacy in biological investigations is to
compare groups separated by time. For evidence so obtained, cause-andeffect should be attributed cautiosly.
Changes in deer populations,
weather conditions, deer response to fence, traffic conditions, and
land use can affect the number of highway deer kills.

�-18Table 9. The numhe.r of ae.Le.ct
ed .fiefu&amp;t..ora1
responses e.:xliiIiited
by deer
approaching or crossing an experimental-net:"'arch overpass during two seasons
(fall 1978 and spring .1979).and ca1culated'behavioralresponse
ratios.
No. of Animals
No~ of
No. 6f
I
Instances Crossings BR :C!
Exhibiting
a
of BR, (BRi) , ..(C) .
BR (BR )

Behavioral
Responses (BR)

BR. :BR
1.

a

a,

Muzz1e-to-Ground

. 2/

-

Control

15

33

43

0.35:1

O. 77 :1

2.20 :1

6

9

18

0.33:1

0.50:1

1.50:1

Control

16

25

43

0.37:1

0.58:11.56:1

Variable

8

15

18

0.44:1

0.83:1

1.88:1

Control

39

102

43

0.91:1

2.37:1

2.62:1

Variable

18

55

18

1.00 :1

3.06:1

3.06:1

Control

48

91

43

1.12 :1

2.12:1

1.90:1

Variable

22

47

18

1.22 :1

2.61:1

2.14:1

Control

7

9

43

0.16:1

0.21:11.29:1

Variable·

2

7·

18 '

0.11:1

0.39:1

Variable

3/

Muzz1e-to-Structure

!!_/

Low-Head 51

•
.
HeS1.tat1.on
-61

Alert: Stance

71

3.50:1

llc denotes the number of crossings during the two seasons.
l/Muzz1e~to~ground

denotes a deer lowering its muzzle to the ground.

21Control was a 2.48 m wide bridge and the variable was the same bridge with
a net arch similar to those used on'pedestrian overpasses.

4/M uzz 1e-to--structure
'
d"enotes a deer"1 ower1.ng
~
-,
its muzzle to the bridge deck or
raising its muzzle to the bridge railing.
2_/Low-head denotes a lowering of the head where the axis of the neck declines
(posterior to anterior) below the horizontal.
' .
"f
-6/C essat1.on
0f f orwar·d movement

or 1•0 second or more. '

L/Alert stance denotes a position where the head and neck are above horizontal
and the 'ears are erect with the vertical axis of the ear either'perpendicular
to horizontal or inclined forward.

�-19-

Table 10. The mean annual number or pre- and post-installation deer
highway kills and percent reduction for six 2.44-m fences adjacent to
Interstate 70 and Highway 82.

Fence (Hwy)

Lengthbf
hwy fenced
km (mles)

Mean annual
preinstallation
"mortality,

Mean annual
post •...
installation
mortality

Vail (1-70)

2.4 (1.5)

36' (3)1/

12.2 (9)1/

66.1

Avon (1-70)

3.6 (2.3)

28 (1)

4.2 (6)

85.1

Edwards (1-70)

3.6 (2.3)

27 (1)

7.0 (6)

74.1 '

Eagle (1-70)

7.7 (4.8)

167 (1)

17 .6 (5)

89.5

Diamond S (82)

1.8 (1.1)

10 (3)'

1.6 '(7)

84.3 .

Carbondale (82) 1.8 (1.1)

14 (5)

4.8 (4)

66.1 .
,77.5 .

Cumulative Avg.

];/

Percent
reduction
mortalitv·

(n) denotes the number' of years of pre-' and pos ti-dns t afLat.Lon data.

Deer movements lateral or parallel to 2.44-m fences have been furth~r
documented at the Vail, Avon, and Eagle fences (Table 11). Based on
lateral movements of radio-collared, neckbanded, or observed deer, females
and males have moved lateral to the fence for mean distances of 0.578 and
0.709 km, respectively.

Response of Deer to Guards
No deer guard prototypes were tested during this segment.
LITERATURE CITED
Gallagher, V. P., and P. G. Meguire. 1974.' Contrast requirements of urban
drivers. Fed. Highway Admin. Rep. Number FHWA-RD-74-76. 72pp.
Hediger, H.

1950.

Wild animals in captivity.

Butterworth, London.

19pp.

�-20-

Table 11. Female and male deer mean lateral movements (km) along or adjacent
to the field side or highway side of 2.44-m fences at Vail, Avon, and Eagle
as de termfned unde r several conditions ...

Vail

Avon.

Eagle·

Female. . ...
Male •.

Side of Fence

Male

Field Side
Apparent 1/ 0.900(n=3)~10.·500(n=2) 0.578(n';"2)
ObserVed

1-' 0.272(n=8)

0.588 (n=107) 0.762 (n=34)

0.350(n=1) 0.280(n=1)

Highway Side
Apparent
Observed

0.617 (n=3)
0.350(n=1)

0.500 (n=5f10 .506 (n=4).--

0.480 (n=4)
1.207 (n=2)

Observed and 5/
haraased

0.975 (n=4)

0.646 (n=5)

l/ Movements

estimated by noting the locations of radio-collared or neckbanded
deer· at selected· pOints along the fences· at widely separated· time periods.

~l (n) denotes number of lateral df.s tiancesc
3t Movements

-

observed· during their duration.

!!./ Included one distance

that was the "net" lateral movement occurring during
a period of 45 minutes.· Twenty.:...two
changes in direction or mini-lateral
movemenbs occur red during this -"net"·movementn : The mean mini-·lateral
movement was l40.9±154.8(SD) m,

~/ Animals were usually chased with a vehicle at night until they either
escaped· from the fenced side of the highway or "broke" back against the
vehicle •.

�-21-

Howe, C. W. 1971. Benefit-cost analysis for water system planning.
Amer. Geophysical Union. Washington, D. C. 144pp.
Myers, G. T. 1969. An investigation of deer-auto accidents.
Pages
147-178 in Game Res. Rept., July, Part 2. Colo. Game, Fish and
Parks Div. Fed. Aid.
Pojar, T. M. 1972a. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 267-275 in Game Res. Rep., July, Part 3.
Colo. Div. Wildl. Fed. Aid.
1972b. An evaluation of deer-proof fence length required to
prevent deer movements on or across high speed highways.
Pages
305-310 in Game Res. Rep., July, Part 3. Colo. Div. Wildl. Fed. Aid.

----- , R. A. Prosence, D. F. Reed, and T. N. Woodard.
of a lighted animated deer crossing
87-91.

sign.

1975. Effectiveness
J. Wildl. Manage. 39(1):

Reed, D. F. 1971. Deer underpass evaluation.
Pages 341-351 in Game Res.
Rep., July, Part 3. Colo. Div. Wildl. Fed. Aid.
1974. An evaluation of 8-foot fence length required to prevent
deer movements on or across high speed highways.
Pages 313-320
in Game Res. Rept., July, Part 2. Colo. Div. Wildl. Fed. Aid.
1975. An evaluation of 8-foot (2.44-m) fence length required to
prevent deer movements on or across high speed highways.
Pages
315-320 in Game Res. Rept" July, Part 2. Colo. Div. Wildl. Fed. Aid.
1976a. Evaluation of deer overpasses.
Pages 247-251 in Game
Res. Rept., July, Part 2. Colo. Div. Wildl. Fed. Aid.
1976b. Evaluation of deer underpasses.
Pages 223-229 'in Game
Res. Rept., July, Part 2. Colo. Div. Wildl. Fed. Aid.

----- , T. M. Pojar, and T. N. Woodard.
deer guards.

J. Range Manage.

1974. Mule deer responses
27(2):111-113.

_____ , T. N. Woodard, and T. M. Pojar.
mule deer to a highway underpass.

1975. Behavioral
J. Wildl. Manage.

to

response of
39(2):361-367.

, and T. D. I. Beck.
1977. Deer-vehicle accidents statewide and
~---methods
and devices to reduce them. Pages 23-41 in Game Res. Rept.,
July, Part 1.

Colo. Div. Wildl. Fed. Aid.

Scott, J. P. 1958. Animal behavior.
Press, Chicago.
349pp.

2nd ed.

University

of Chicago

Woodard, T. N. 1973a. Monitor potentially critical deer-vehicle accident
areas statewide.
Pages 163-170 in Game Res. Rept., July, Part 2.
Colo. Div. Wildl. Fed. Aid.

�-22-

1973b. An evaluation of deer-proof fence length required to
prevent-deer movements on or across high speed highways.
Pages
197-202 in Game Res. Rept., July, Part 2. Colo. Div. Wildl. Fed. Aid.
1974. MOnitor potentially critical deer-vehicle accident areas
statewide.
Pages 293-298 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. Fed. Aid.
1975. Monitor potentially critical deer-vehicle accident areas
statewide.
Pages 295-303 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. Fed. Aid.
1976a. Monitor potentially critical deer-vehicle accident areas
statewide.
Pages 215-221 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. Fed. Aid.
1976b. Effects of highway lighting on number of deer killed by
vehicles.
Pages 231-236 in Game Res. Rep., July, Part 2. Colorado
Div. Wildl. Fed. Aid.
1976c. An evaluation of deer-proof fence length required to prevent
deer movements on or across high speed highways.
Pages 239-244 in
Game Res. Rep., July, Part 2. Colorado Div. Wildl. Fed. Aid.

Prepared

by:----::-"-:If:-/_'u-=··,_·'-:{

_·",;-_1-:'

_tu_, _._j__'~_.

Dale F. Reed
Wildlife Researcher

_

�July, 1979

-23-

JOB PROGRESS

State of

REPORT

COLORADO
--------~~~~~----------

1

Work Plan No.
Job Title

Guidelines

Period Covered:
Personnel:

Big Game Investigations

W-126-R-2

Proj ect No.

Job No.
for Wildlife

1

---------------------------------

Habitat Manipulation

July 1, 1978 - June 30, 1979

W. H. Rutherford,

W. D. Snyder

ABSTRACT
Literature review and interviews have provided a background of information
on techniques and procedures for manipulation of wildlife habitat.
This
information has been compiled, organized and written into a manuscript for
a guideline manual that is about 70% complete.
Completed portions have
been turned over to an editor.

��-25-

GUIDELINES
William

FOR WILDLIFE

H. Rutherford

HABITAT MANIPULATION
and Warren

D. Snyder

P. N. OBJECTIVE
To publish a comprehensive manual of directional
guidelines for sustaining and enhancing wildlife
consumptive and non-consumptive uses on Division

habitat management
populations for
of Wildlife properties.

SEGMENT OBJECTIVE
1.

Assemble information relevant
a looseleaf manual format.

2.

Review draft version
selected referees.

to wildlife

of the wildlife

METHODS

habitat manipulation

habitat manipulation

into

manual with

AND MATERIALS

With literature review and personal interviews having been essentially
completed during the previous segment, activity during this segment was
limited to organizing, compiling and writing the manuscript.
Methods
and materials are self-explanatory.

RESULTS
Extensive data compilation and writing were accomplished during the segment.
The format for presentation was established and subsequently revised and
refined.
Effort was concentrated on bringing the voluminous information
together in a condensed orderly arrangement.
At the end of this segment (June 30) the sections on basic habitat design
considerations, basic habitat management concepts (by groups of species),
description of manipulation procedures and techniques, and application of
procedures and techniques to various wildlife species and habitats, are
approximately 80% complete.
A great deal of the rough draft of these
sections has been turned over to the Division of Wildlife staff technical
editor.
The description of general procedures not species-related,
and the
final section on procedures for evaluating the effects of habitat manipulation,
remain to be drafted.
Activity during the next segment will be directed
.
toward completion of the drafting, review and revision, and setting up
printing and publishing.

Prepared

by:

��-27-

JOB PROGRESS

State

0

f

REPORT

W-126-R-2

Work Plan No.

Period

1979

C.=.O.:_:L=-O:..:RAD=:..:O'--_

Proj ect No.

Job Title

July,

Big "Game Investigations
Job No.

1

Deer and Elk Management

Covered:

~2~

_

Study

July 1, 1978 to June 30, 1979

Personnel:
Colorado Division of Wildlife:
Regional Wildlife Biologist staff
members; John Ellenberger, Hal Burdick, Jim Olterman, Marc Elkins, Jack
Vayhinger; Area Supervisors and District Wildlife Managers; Thomas M. Pojar.

ABSTRACT
All simulations and data analysis unit boundaries for mule deer, elk, and
pronghorn antelope in the NW, SW, and SE regions were reviewed by three
levels of management; Regional Wildlife Biologists, Area Wildlife Supervisors,
and District Wildlife Managers.
These reviews resulted in updating and
various degrees of revisions of the simulations.
The boundaries of five
DAU's were changed and new simulations created for the new alignments.
Currently there are simulations for 38 mule deer populations, 35 elk populations,
15 pronghorn antelope populations, 2 bighorn sheep populations and one mountain
goat population for a total of 91 big game population simulations.
The game
management units included in the DAU's are listed in Appendix I. Some special
population simulations were created for an area the Division was considering
leasing and for special research areas.
Various aspects of the simulating process,
in the papers cited in the Results Section

utility and status
of this report.

are discussed

��-29-

DEER AND ELK MANAGEMENT

STUDY

Thomas M. Pojar

P. N. OBJECTIVE
Devise and test a statewide deer and elk management

system.

SEGMENT OBJECTIVES
1.

Divide the state into a system of data analysis units (DAU's) that
are deemed practical for gathering and analyzing population data.

2.

Based on currently available information, estimate population
by DAU, through the simulation modeling approach.

3.

Conduct training workshops involving management personnel to familiarize
them with the interpretation of the output from the model and to
incorporate empirical data from fieldmen directly associated with the
DAU into the simulation to produce a candidate management simulation.

4.

Identify data that are most useful for improving
parameters.

estimates

parameters,

of population

RESULTS
The following manuscripts

were prepared:

Pojar, T. M. 1979. Population modeling in Colorado.
In T. M. Pojar and D. Strickland (Eds.). 1979. A workshop on the
Status and Application of Big Game Population Modeling.
Ft. Collins,
Colorado, Jan. 16-17. 1979. 53 pp.
Pojar, T. M., and D. Strickland.
1979. (Eds.). A workshop on the status
and application of big game population modeling.
Ft. Collins, Colorado,
Jan. 16-17, 1979. 53pp.
Salwasser, H., and T. M. Pojar.
1979. Simulation modeling of pronghorn
populations.
Submitted to: J. Yoakum (Ed.) Pronghorn Management.
Wildlife Management Institute.

C
Prepared

by:

~6~
Wildlife

Researcher

�-30-

APPENDIX I

�-31APPENDIX I
The current (1978) data analysis unit alignment
ELK
DAU

Identifier

Name

Units

E 1

CS

Cold Springs

201

E 2

BE

Bear's Ears

4, 5, 441, 14

E 3

NP

North Park

6, 7, 16, 161

E 4

PR

Poudre River

8, 19

E 5

ST

Strawberry

11

E 6

WR

White River

12, 13, 131, 23, 24, 25,
26, 33, 34

E 7

GP

Gore Pass

15, 27

E 8

TR

Troublesome

17, 18

E 9

SV

St. Vrain

20, 29, 38, 381

E10

YC

Yellow Creek

22

Ell

RB

Roan Bluffs

31, 32

E12

PI

Piney

35, 36

E13

WF

Williams Fork

28, 37

E14

GM

Grand Mesa

41, 411, 42, 52, 521

E15

AC

Avalanche Creek

43, 471

E16

FP

Frying Pan

44, 45, 47

E17

CR

Collegiate Range

48, 56, 481

E18

KP

Kenosha Pass

50, 500, 501, 51, 511, 46,
461, 39, 391

E19

GL

Glade Park

40

E20

UN

Uncompahgre

61, 62

E21

GR

Gunnison River

63, 53, 54, 55,551

E22

BP

Buffalo Peaks

49, 57, 58

E23

EM

11-Mile

58, 59

E24

DC

Disappointment Creek

70, 71

E25

LF

Lake Fork

66, 67

E26

SA

Saguache

68

E27

SD

Sangre de Cristo

82, 86

E28

GC

Grape Creek

69, 84

E29

MV

Mesa Verde

72, 73

E30

HE

Hermosa

74

E31

SJ

San Juan

75, 751, 77, 78

E32

LG

Lower Rio Grande

80, 81

E33

TI

Trinchera

83, 85, 851

E34

UG

Upper Rio Grande

76, 79

E35

CI

Cimarron

64, 65

�-32-

MULE DEER

DAU

Identifier

Name

Units

D 1

LS

Little Snake

1, 2, 201

D 2

BE

Bear's Ears

4, 441,

5,

14

D 3

NP

North Park

6,

161,

17

D 4

RF

Red Feather

7,8,9,19

D 5

SP

South Platte

87,
92,
99,

D 6

RA

Rangely

10

D 7

WR

White River

11, 12, 13, 131, 22,
23, 24, 25, 26, 31, 32,
33, 34

D 8

SB

State Bridge

15, 35,

36

D 9

MP

Middle Park

18, 28,

27,

37

D10

BO

Boulder

20,

29,

38,

381

Dll

BC

Book Cliffs

30,

21

D12

GM

Grand Mesa

41,

411,

42,

52,

521

D13

MB

Maroon Bells

43,

47,

471

D14

RT

Red Tables

44,

45

DIS

CO

Cottonwood Creek

48,

56,

481

D16

CC

Cripple Creek

49,

57,

58,

581,

59

D17

BA

Bailey

39,
51,

391,
500,

46, 461,
511

D18

GP

Glade Park

40

D19

UP

Uncompahgre

61,

D20

CR

Crawford

53

D21

WE

West Elk

54

D22

TP

Taylor Park

55,

D23

LA

LaSal

60

D24

GH

Groundhog

70,

71

D25

PC

Powderhorn Creek

66,

67

D26

SA

Saguache

68

D27

WC

Westcliffe

69,

D28

AV

Arkansas Valley

97,98,102,103,106,107,
108, 109, 1l0, Ill,

16,

88, 89, 90, 91,
93, 94, 95, 96,
100, 104, 105

50,

501,

62

551

84,

86

D29

MV

Mesa Verde

72,

73,

74

D30

SJ

San Juan

75,

751,

77,

D32

TR

Trinidad

83,

85,

ll2

D33

GD

Great Divide

3

78

ll3,

114

�-33-

MULE DEER
(Cont'd)
DAU

Identifier

Name

D35

LG

Lower Rio Grande

80, 81

D36

UG

Upper Rio Grande

76, 79

D37

SD

Sand Dunes

82

D39

FM

Fruitland Mesa

63

D40

CI

Cimmeron

64, 65

Units

PRONGHORN ANTELOPE
PI

ESC

Escarpment

10, l1, 12, 13,14, 17

P2

HAR

Hardpan

22, 23, 24, 28, 29, 30, 31

P3

NP

North Park

6, 7, 8, 9

P4

SH

Sandhills

15, 16, 18, 19, 20, 21, 25,
26, 27

P5

HW

Haswell

45, 46, 51, 52

P6

HU

Hugo

36, 37, 38

P7

TH

Thatcher

49, 50, 55, 56, 59, 60, 61,
66

P8

YO

Yoder

34, 35, 41, 42, 43, 44

P9

GD

Great Divide

3

P10

MA

Maybell

5

Pll

VB

Vermillian Bluffs

2

P12

CH

Cheyenne

39, 40, 47, 48, 53, 54

P13

TO

Tobe

57, 58, 62, 63, 64, 67, 68

P14

VG

Villa Grove

74

PIS

SW

Saguache West

73

P16

MV

Monte Vista

77

BIGHORN SHEEP
B1

PP

Pikes Peak

59

B2

™

Trickle Mountain

68, 681

MOUNTAIN GOAT
Gl

SA

Sawatch

48, 481

��-35July,

JOB PROGRESS

State of

COLORADO
------------------------------

Project

No.

W-126-R-2

Plan No.

1

Work

Job Title

REPORT

Deer-Elk

J~ ~.

Investigations

---------------------------------

Brucellosis-Leptospirosis
Survey of Big Game Animals in Colorado
--~~~~~~-=~~~~~~~~~~~~~~~~---------------------June

30, 1979

Personnel:
Joan Ritchie, Carol Ann Weinland,
Marilyn Stevens, William Adrian.

Richard

Period

1979

Covered:

July

1, 1978 through

Ford, Gene Grenard,

ABSTRACT
During the 1978 hunting season blood collection kits were mailed to all
antelope hunters (4,625) and 13,181 deer hunters.
The serum received from
successful hunters was tested for serological titers for brucellosis and
five serotypes of leptospira (~. canic61a, grippotyphosa,
hardjo,
icterohemorrhagiae,
and pomana).
All deer and antelope serum tested for
brucellosis were negative.
All deer and antelope serum was considered
negative for all serotypes of leptospira except ~. hardjo.
Deer had less
than 1% as reactors while antelope continued to have about 10% incidence of
reactors to L. hardio.

��-37-

BRUCELLOSIS-LEPTOSPIROSIS
SURVEY
OF BIG GAME ANIMALS IN COLORADO
William

J. Adrian

P. N. OBJECTIVE
To survey big game populations within the State of Colorado
or absence of brucellosis and leptospirosis.

for the presence

SEGMENT OBJECTIVE
1.

Survey Colorado's deer populations for brucellosis and leptospirosis
during the annual hunting season and continue
to monitor antelope
for serological titers for L. hardjo.

METHODS AND MATERIALS
Blood sample kits were mailed to deer hunters with the request they draw
a blood sample from their kill and return the kit to the Colorado Cooperative
Brucellosis Laboratory in Denver, Colorado for processing.
A maximum of 300
sampling kits were sent to hunters in each deer unit.
If more than 300 permits
were issued for one unit the names were randomly selected.
Blood sample kits
were mailed to all antelope hunters.
The kit consists of a cardboard mailer, blood tube and stopper, plastic
rubber band, paper towel and instructions for obtaining the sample and
mailing the kits.

bag,

RESULTS AND DISCUSSION
The Wildlife Research Laboratory in cooperation with the Cooperative Brucellosis
Laboratory in Denver has, in 1978, completed its tenth annual survey for
brucellosis and leptospirosis in Colorado's big game populations.
Brucellosis is a specific contagious disease which primarily affects cattle,
swine, and goats.
It can, however, affect many other animals including elk,
deer, antelope and man (undulant fever, Malta fever and Bang's Disease).
It
is caused by bacteria of the Brucella group and is characterized by abortion
in the female, and to a lesser extent, inflammation of the testes and infection
of the accessory sex glands in the male and infertility in both sexes.
Leptospirosis is a febrile (feverish) disease caused by certain species of
leptospirae.
The disease occurs with various manifestations in dogs, cattle,
pigs, sheep, goats, horses, and wildlife and is transmissible to man.

�-38-

Leptospira hardjo was first reported in the continental u.s. in 1951 and
first isolated from cattle in 1960 (Roth, 1960). ~. hardjo infections were
considered to be limited to the bovine until it was recently isolated from
humans (Stallman, 1972), sheep (Andreani, 1975), opossum (Brockie 1975), and
horses (Myers 1976).
Spread of ~. hardjo through the U.S. has been rapid. Recent reports from
Georgia and Alabama indicate leptospira reactor in cattle of approximately
22% for ~. hardjo and 10% for ~. pomona.
Nationally, the react of rate for
L. hardjo in cattle is about 7% (Rubin, 1976). Since 1967, the U.S. Animal
Health Association, Leptospirosis Committee has considered L. hardjo
infections in cattle a major economic threat (Glosser et al~ 1974).
Colorado's first serologic evidence for L. hardjo infection came in 1974.
Of 646 antelope serum samples, 4 had titers of 1:10 and 2 had titers of
1:40. In 1977, serologic evidence of L. hardjo infection became more
substantial.
Of 666 antelope samples,-55 had titers of 1:10, 15 had titers
of 1:40 and 3 had titers of greater than 1:160 by the macroscopic
agglutination technique (Adrian and Keiss 1977).
The 1978 deer and antelope
summarized in Table 1.

brucellosis-leptospirosis

survey data are

Species

Kits Mailed

Sample
Received

Sample
Hemolyzed

Antelope

4,625

1,477

991

486

o

427

52

Deer

13,181

1,663

1,173

976

o

485

5

The data in Table 1 is available by game management
incidence of ~. hardjo upon request.

Brucellosis
Neg
Pos

Leptospirosis
Neg
Pos

unit with its corresponding

Attempts to collect antelope in high reactor. areas and isolate~.
hardjo
has been unsuccessful.
Antelope transplants from areas with L. hardjo reactors
will be screened in the field and those animals with serological titers for
~. hardjo will be necropsied and cultured followed by the appropriate
histopathological
studies.

LITERATURE

CITED

Adrian, W. J., and R. E. Keiss.
1977.
for serologic titers to brucellosis
Diseases:
13:429-431.

Survey of Colorado's wild ruminants
and leptospirosis.
J. of Wildlife

Andreani, E. E., Santarelli, and R. Diligenti.
1975.
sheep. Natural infection by the Hardjo serotype.
Di Medicina Veterinaria Di Pisa.
27:33-40.

Leptospirosis in
Annali Della Facolta

�-39-

Brockie, R. E. 1977. Leptospirae infections
New Zealand.
Vet. J. 25:4, 89-95.

of rodents

in North Island

Glosser, J. W., C. R. Sulzer, G. C. Reynolds, and D. Whitsett.
1974.
Isolation of Leptospira serotype Szanajizak from dairy cattle in
Oregon.
In proceedings.
78th Ann. Meeting U.S. Animal Health Assoc.
Roanoke, VA. 74:119-125.
Myers, D. M. 1976. Serological studies from isolation of serotype
Hardjo and Leptospira bifloxa strains from horses of Argentina.
of Clin. Micro. 3:548-555.

J.

Roth, E. E., and M. M. Galton.
1960. Isolation and identification of
Leptospira Hardjo from cattle in Louisiana.
Am. J. Vet. Res. 21:422-427.
Rubin, H. L.

1976.

Bovine leptospirosis.

Florida Vet. J. 6:16-19.

Stallman, N. D. 1972. The isolation of a strain of Leptospira, serotype
Hardjo from a patient in Southern Queensland, Australia.
Vet. J. 48:497.

Prepared

".

Z:

bY:M_y~

~

//

,

,/'

fI2

(.L ~~~

1 liam J. Adria~
Wildlife Researiher

/

"-

��July 1979

-41-

JOB PROGRESS

State of

REPORT

COLORADO

--------~----~--------------

Proj ect No.

W-126-R-2

Work Plan No.

I

Job Title

Job No.

Animal and Pen Support Facilities

Period Covered:
Personnel:

Deer-Elk

Invest~gations
4__ ~

for Deer-Elk

_

Research

July 1, 1978 - June 30, 1979

Paul H. Neil, John Anderson, Dan Parkinson, Valerie Thomas,
Barry VanSant, Don Bay, Steve Torbit, Arnold O'l.s en , Lynn Sexton,
and R. Bruce Gill.

ABSTRACT
During the 1978-1979 segment, emphasis was directed towards completion of
the deer/elk research facility and the raising and training of seven mule
deer fawns for use in metabolism cages.
Three additional fawns were added
to this group during mid-winter 1979.
These all metal metabolism cages were completed and installed for an in vivo
study of mule deer body composition by use of tritiated water.
A sixty day
feeding trial was performed on six mature mule deer. Techniques of injection,
sample collection and response to feeding levels were-investigated.
More
detailed information regarding sample preparation, data collection and
analysis is described under W~126-R-2, Work Plan 2, Job 1.
Pregnancy scans were conducted on four mature does within the facility
using an ultra-high sound wave fluoroscope.
Three of the four does were
shown to be pregnant with twins. Detailed information on the scanner system
and techniques are described under W-126-R-2, Work Plan 5, Job 1.
Early winter grazing trials using three tame mule deer were conducted during
November 1978. The study area is the proposed experimental controlled burn
site described under W-126-R-2, Work Plan 4, Job 1. The seven day trial
had to be terminated early due to heavy snowfall which altered the environmental
conditions being sampled.
Additional heavy snowfall during the remaining
winter months prohibited further trials. Although the November trial was
considered a training trial, plant species comprising 2% or more of the diet
as well as other plant species taken are listed in this report.
Pen and facility expansion continued during the segment with the installation
of a complete watering system and electrical system for each pen. A Toledo
digital read-out scale was incorporated into the runway system.
Lab and
office facilities were also completed.

��-43-

ANIMAL AND PEN SUPPORT
FACILITIES FOR DEER-ELK RESEARCH

Paul H. Neil

P. N. OBJECTIVES
To provide and maintain populations of captive big game animals
facilities to support big game research programs.

and pen

SEGMENT OBJECTIVES
1.

To continue
facility.

to expand the big game research

2.

To coordinate rearing, training, maintenance, and research activities
with captive wild and tame animals under one research support facility
manager.

3.

To integrate animal and physical plant support
monetary requirements under a single budget.

METHODS

and animal holding

facility manpower

and

AND MATERIALS

Materials and equipment were ordered and various aspects of construction
were put on bid in order to complete the final portions of the expansion
program at the deer-elk facility.
A Toledo digital read-out scale was
ordered for installation and three metabolic cages were constructed.
A water system and an electrical system were designed and installed to
meet the needs of the entire facility.
The young-adult conservation corps of the Bureau of Reclamation was enlisted
to assist in completion of the lab and office facility and other projects.
Seven orphan mule deer fawns were obtained from Division of Wildlife field
personnel and hand reared and trained for use in the metabolism cages.
Three additional fawns were recruited during mid-winter of 1979 as a result
of severe winter conditions.
The formula used for hand-rearing was the
same used during the last segment and consisted of 2 parts whole milk to
1 part canned evaporated milk with an addition of 1/10 the total volume
per day of buttermilk.

RESULTS AND DISCUSSION
All ten of the orphaned
cage programs.
Rearing

fawns were successfully reared for metabolism
success was again attributed to the formula, good

�-44-

weather, and improved rearing management techniques.
Because of the
young age of the 10 orphans, six mature mule deer were used for a sixty
day feeding trial in the newly constructed metabolism cages.
During the first phase of this program techniques of injection of tritiated
water as a tracer, sample collection and response to feeding levels were
investigated.
Detailed methods and data showing predicted weight loss
and observed weight loss for the animals is described under W-126-R-2,
Work Plan 2, Job 1.
Construction of the three metabolism cages being used for the body
composition study of mule deer by use of tritiated water allows for complete
urine and feces collection.
Certification for use of radionuclides was
obtained and project approval obtained from the Colorado State University
Radiation Safety Committee.
During fall of 1978 the only remalnlng breeding buck died of apparent
"chronic wasting disease".
He was replaced by capturing a wild yearling
buck. A Pneu-dart rifle and a 10 mg dose of succinycholine chloride were
used for capture.
Drug reaction time was 5 minutes, immobilization time
was 15 minutes, and recovery time was 10 minutes.
Following capture,
the antlers were sawed off and the buck was released into the Animal
Research facility with the does.
Pregnancy scans were conducted in the spring of 1979 on four mature does
within the facility using an ultra-high soundwave fluo.roscope. The scanner
revealed that three of the four does were pregnant with twins. Detailed
information on the scanner system and techniques are described under
W-126-R-2, Work Plan 5, Job 1.
Early winter mule deer grazing trials using three tame deer were conducted
during November 1978 in the Poudre Canyon area west of Fort Collins, Colorado.
The study area is proposed as an experimental controlled burn site described
under W-126-R-2, Work Plan 4, Job 1. The seven day trial began November
28, 1978 but had to be terminated on December 2, 1978 due to heavy snowfall
which altered the environmental conditions being sampled.
This trial was
considered a training trial since it could not be completed.
Plant species
comprising 2% or more of the total diet as well as other plant species taken
are listed in Table 1.
Pen and facility expansion continued during the segment and is presently
in the final stage. Automatic watering devices were installed in each
large holding pen and isolation pen. Electrical outlets and lights were
also installed in strategic locations.
A 16' x 16' weigh room is presently
being constructed to accommodate the digital read-out scale that was
incorporated into the runway system.
The scale is capable of weighing
both deer and elk accurately and at any ambient temperature.
Four castrated male elk and one cow elk now occupy a 2 acre pen. All
of the deer will be moved to the expanded facility area early in the next
segment.
The Division of Wildlife Research Animal herd presently consists
of the following:
Deer - 10 mature does, 3 immature does, one breeding

�-45-

buck, and 8 male castrates (mature and immature);
Elk - 4 mature male
castrates and one mature cow; Bighorn sheep - 3 mature castrate rams,
1 yearling ram, and 2 yearling ewes; Rocky Mountain Goats - 2 yearling
nannies, one yearling billy and 4 nanny kids.

Prepared

by:

~-=-'-"'~(:--:A:_~_"
__
" -:fo:-::-i._'

._-,,-iJ_;:)~.....-c
__·k__

Paul H. Neil
Wildlife Technician

II

�Table 1.

Mule deer grazing trials Poudre Canyon Area, November,
Shrubs

Grasses
Muh1enbergia montana
Poa species 5.07%
Carex species 3.90%
Hesperoch1oa kingii
Agropyron spicatum
Agroypyron smithii
Agropyron trachycau1um
Agropyron sub secundum
Bromus techtorum
Bromus species
Boute1oua gracilis
Danthonia parryi
Stipa comata
Ko1eria cristata
Sitanion hystrix

1978.

12.12%

Populus tremu10ides (dry leaves) 14.32%
Pinus ponderosa (dry needles) 6.17%
Pinus ponderosa (green needles) 5.24%
Acer glabrum 4.08%
~hia
tridentata 3.97%
Prunus virginiana 2.45%
Ribes cereum 2.14%
Arctostaphylos ~~-ursi
Jamesia americana
Salix sp.
Moss sp.
Ame1anchier a1nifo1ia
Pseudotsuga menziesii (green-dry needles)
Artemisia tridentata
Rubus de1iciousus
--Rosa acicu1aris
Juniperus scopu1orum

Forbs
Potenti11a fissa 13.92%
Artemisia frigida 4.43%
Erigonium umbe11atum 2.71%
Aster porteri 2.5%
Artemisia 1udoviciana
Oxytropis species
Campanu1a species
Chrysops is vi110sa
Erigeron caespitosus
Tragopogon dubius
Geranium sp.
Geranium parryi
Thermopsis divaricarpa
Antennaria sp.
Potenti11a sp.
Potenti11a pucherrima
Potenti11a hippiana
Ba1samorhiza sagittata
He1ianthus pumi1us
Gallium borea1e
Lupinus greenei
Aster 1avens
Grinde11ia suba1pina
Aster glaucodes
Erigeron speciosus
Achi11ia 1anu1osa
Solidago nana
Unknown forbs
Agoseris sp.
Mahonea rep ens
Arenaria fend1eri
Cryptantha sp.

I
.jO-

0\
I

�-47-

July, 1979

JOB PROGRESS

State of

REPORT

COLORADO
------------------------------

Project No.

W-126-R-2

Work Plan No.

1

Job Title

Nutrient

Digestible

Period Covered:
Personnel:

Deer-Elk
Job No.

Investigations

6
------------~-------------------

Content of Deer and Elk Winter Forage Plants

July 1, 1978 through June 30, 1979

Roland C. Kufeld, Marilyn

Stevens

ABSTRACT
In-vitro digestible dry matter levels were determined for 45 Gambel oak
(Quercus gaffibellii)and 45 big sagebrush (Artemisia tridentata) samples
collected during January, 1976. Each species was collected from 45 sites
throughout 9 major areas of the state and 3 vegetation types. For Gambel
oak the median in-vitro digestible dry matter level from 9 areas is 27.82
percent, and the 95 percent confidence interval ranges from 26.98 to 29.33
percent.
For big sage the median in-vitro digestible dry matter level is
49.76 percent, and the 95 percent confidence interval r~nges from 47.15 to
53.09 percent.
During January, 1979, five 70 to 100 gram, green weight
composite samples of two species, serviceberry (Amelanchier alnifolia)
and mountain mahogany (Cercocarpus montanus) were collected from 8 widely
separated geographic areas throughout the western half of Colorado.
These
will be analyzed for digestible dry matter and trace mineral content.

��-49-

DIGESTIBLE

NUTRIENT CONTENT OF DEER AND ELK
WINTER FORAGE PLANTS
Roland C. Kufeld

P. N. OBJECTIVE
To estimate the average digestible nutrient content values and degree
of variation in digestible nutrient content of selected range forage plants
during winter.

SEGMENT OBJECTIVE
To determine the degree of variation in digestible
selected range forage plants during winter.

nutrient

content of

METHODS AND MATERIALS
In-vitro digestibility was determined for 45 samples of Gambel oak
(Quercus gambellii) and 45 samples of big sagebrush (Artemisia tridentata),
which were collected and analyzed for nutrient content in the manner described
by Kufeld (1978). The technique for determining in-vitro digestibility was
described by Tilley and Terry (1963) and modified by Pearson (1970).
Inoculum
used was from a holstein cow.
During January, 1979, five 70 to 100 gram, green weight, composite samples
of two species, serviceberry (Amelanchier alnifolia) and mountain mahogany
(Cercocarpus montanus) were collected from 8 widely separated areas throughout
the western half of Colorado, but within the normal winter range of deer and
elk. The 5 collectio~ sites within each major area were widely separated
geographically, but were situated within the same vegetation type (Table 1).
Collections of both species consisted of current annual growth stems with
leaves excluded.
Each sample was air dried and ground in a Wiley Mill.
They will be analyzed to determine content of the following nutrients and
trace minerals:
Total cell contents, crude protein, ether extract, soluble
ash, soluble carbohydrate, total cell walls, acid insoluble ash, lignin,
hemicellulose, cellulose, calcium, copper, iron, magnesium, manganese,
phosphorus, potassium, sodium, zinc (Harris 1970; Adrian 1973).
In vitro
dry matter digestibility of each sample will also be determined (Tilley
and Terry, 1963; Pearson, 1970).

RESULTS AND DISCUSSION
Analysis of variance showed significant differences at the 5 percent level
among vegetation types, among areas, and among areas within vegetation types
for in vitro digestible dry matter sagebrush, but no significant differences
were~ound
in Gambel oak. Despite the significant differences in

�-50-

sagebrush actual differences in both sagebrush and Gambel oak were
relatively small (Table 2). For big sagebrush the median in-vitro
digestible dry matter from the 9 major areas was 49·,76 percent, and the 95
percent confidence interval ranged from 47.15 to 53.09 percent.
For Gambel
oak the median in-vitro digestible dry matter level was 27.82, and the 95
percent confidence interval ranged from 26.98 to 29.33 percent.
The data
reflect relatively little mid-winter variation from one locality to another
in content of any of the nutrients for big sagebrush and Gambel oak.
Table 1. Major areas within Colorado where winter samples of serviceherry
and mountain mahogany were collected.
Vegetation Type Where Collections Were Made
Serviceberry Collections
Mtn. Mahogany
Gambel oak-Serviceberry
Pinyon-juniper
Mtn. Mahogany
1.

Steamboat
Meeker

SpringsGrand Junction

Fort Collins

Collections
Pinyon-juniper
Glenwood
Eagle

Sp

Springs-

2.

Rifle-Eagle

Naturita

Denver-Idaho

3.

Black Canyon

East side of
Uncompahgre
Plateau

Rifle

Grand Junction

4.

Durango

Cortez

Montrose

Cortez-Durango

Colo. Sp.
Canon City

Table 2. Medians, 95 percent confidence intervals and non-parametric tolerance
levels for in-vitro digestible dry matter content in Gambel oak and big
sagebrush collected during January throughout Colorado!/.

Plant

Percent content in plant
Median
95% Confidence Interval

Gambel oak

27.82

26.98 - 29.33

26.60

Big Sagebrush

49.76

47.15 - 53.09

45.48

II

Non-parametric lower
bound for 70% poin~1

- Data are based on collections of 5 samples from widely separated
within each of 8 major areas throughout Colorado.

locations

IIThis represents the lowest 5 sample mean value collected from the 9 major
areas.
One can be 95 percent sure that at least 70 percent of the area
mean nutrient values encountered in Colorado, during January, would exceed
the minimum value listed for each nutrient.

�-51-

Medians rather than means were used to describe "average" in vitro digestible
dry matter levels in plants among the 9 geographic areas to avoid the influence
of abnormally high or low values which might tend to raise or lower a mean.
In this situation it was believed that a median would provide a more
representative estimate of the "average" digestible dry matter which one
could expect to encounter in the field. Medians are also suited for use
with one-sided non-parametric tolerance intervals discussed below.
One-sided non-parametric tolerance intervals can be applied to further
minimize the decision-making error incurred in projecting digestibility levels
based on average values (Dixon and Massey, 1969). It is desirable that
digestibility levels be underestimated rather than overestimated to avoid
overestimation of the ability of the range to supply the minimum needs of
grazing animals.
This could lead to overstocking.
One sided non-parametric
tolerance intervals permit estimation of the portion of the overall population
of digestibility values that would likely be encountered for a given species,
which, with 95 percent confidence, exceeds the smallest sample value.
For
example, using the sample means, which range from 26.60 to 29.50 percent
in-vitro digestible dry matter from the 9 areas where oak was collected, one
can be 95 percent sure that at'least 70 percent of the area digestible dry
matter values encountered in Gambel oak in Colorado during January would
exceed the minimum value of 26.60 percent.
By using the minimum value a
manager could, therefore, be reasonably sure his estimate of digestible dry
matter production is on the conservative side. One sided non-parametric
tolerance intervals for in-vitro digestible dry matter in big sagebrush and
Gambel oak are presented~n
Table 2.

LITERATURE

CITED

Adrian, W. J. 1973. A comparision of a wet pressure digestion method
with other commonly used wet and dry-ashing methods.
Analyst 98:213-216.
Dixon, W. J., and F. J. Massey, Jr. 1969. Introduction
analysis.
McGraw-Hill, 3rd ed. 638p.

to statistical

Harris, L. E. 1970. Nutrition research techniques for domestic and wild
animals.
Pub. by Lorin E. Harris, 1408 Highland Dr., Logan, Utah.
1(2):1401-3519.
Kufeld, R. C. 1978. Digestible nutrient and trace mineral content of deer
and elk forage plants during winter.
Colorado Division of Wildlife.
Game Res. Research Report July (3):413-421.
Pearson, H. A. 1970. Digestibility trials:
In-vitro techniques.
P. 85-92.
In:
Range and wildlife habitat evaluation - a research symposium.
USDA For. Servo Misc. Publ. No. 1147. 220p.
Tilley, J. M. A., and R. A. Terry.
1963. A two stage technique for the
in-vitro digestion of forage crops. J. Br. Grassl. Soc. 18:104-111.
Prepared

by

£taA-(,O/ C ;Kf_J1.if
i

Roland C. Kufeld
Wildlife Researcher

��-53-

July, 1979

JOB PROGRESS REPORT

State of

COLORADO
----------------------------

Work Plan No.
Job Title

Big Game Investigations

W-126-R-2

Project No.

1

Multispecies

Job No.
Investigations:

7

----------------------------------

Big Game Research Publications

Period Covered: July 1, 1978 - June 30, 1979
Personnel:
Dr. L. H. Carpenter, R. B. Gill, Dr. O. C. Wallmo, D. J. Freddy,
D. L. Baker, N. T. Hobbs, R. M. Bartmann, A. E. Anderson, Dr. D. C. Bowden,
R. C. Kufeld, D. F. Reed, T. N. Woodard, P. H. Neil, T. M. Pojar, W. H.
Rutherford.

ABSTRACT
Ten publications were completed during the 1978-79 segment.
Three papers
were prepared for presentation at professional society meetings or workshops.
Eight publications are nearing the completed draft stage or are in a state
of manuscript review and revision.

��-55-

BIG GAME RESEARCH PUBLICATIONS
R. BRUCE GILL

P. N. OBJECTIVE
To publish the results of research conducted under the auspices of Federal
Aid Project W-126-R in a variety of professional journals and other indexed
publishing media to insure widespread dissemination and availability of
this information to natural resource managers and ecological scientists.

SEGMENT OBJECTIVES
Following is a list of publication working titles that will be prepared
and/or published under this job in Segment 2. The publication outlet is
tentative depending upon acceptance by the particular periodical.
1. Journal of Wildlife Management
patterns of mule deer at pasture".

- "Twenty-four

hour activity

2. Journal of Ecology - "Mule deer forage diversity
salubrity".
3. Journal of Range Management - "Effects
2,4-D herbicide on mule deer winter range",

and dietary

of time of application

4. Colorado Division of Wildlife Special Report - "Permits
success for special deer seasons in Middle Park".

of

and hunter

5. Journal of Wildlife Management - "A sampling system for estimating
deer pellet group density in Pinyon-juniper woodland".
6. Journal of Wildlife Management
mule deer at various activities".

- "Heart rate measurements

of

7. Journal of Wildlife Management - "Comparison of aerial and ground
measurements of sex and age ratios of mule deer populations".
8. Journal of Wildlife Management - "Rearing and training
Mountain elk for nutritional investigations".

Rocky

9. Journal of Wildlife Management - "Elk summer diet selection
nutritional composition in Rocky Mountain National Park".
10. Journal of Range Management
by grazing animals from microscopic

and

,....
"Estimation of the diets selected
analysis of the feces - warning 2".

11. Journal of Animal Science - "Effects of ambient
in vivo digestibility in mule deer".

temperature

on

�-56-

12. Journal of Wildlife Management - ·"Xylazine as an immobilizing
agent for captive elk (Cervus elaphus nelsonii)"
13. Journal of Wildlife Management - "Statistical models and sampling
design in estimating buck:doe and fawn:doe ratios in Colorado mule deer".
14. Journal of Wildlife Management
fawns in relation to habitat".

- "Mortality

of radio-collared

15. Journal of Mammalogy - "Composition and behavior
groups in free-ranging Rocky Mountain mule deer".
16. Colorado Division
trace minerals in bighorn

of Wildlife Division
sheep nutrition".

of maternal

Report - "Sources

of

17. Journal of Range Management - "Variation in nutrient content
levels of gambel oak (Quercus gambellii) and big sagebrush (Artemisia
tridentata) collected from diverse sites during winter".
18. Wildlife Society Bulletin - "Benefit-cost
to reduce deer-vehicle accidents".

analysis

19. Journal of Wildlife Management - "Effectiveness
lighting in reducing deer-vehicle accidents".

PUBLICATION

of methods

of highway

PROGRESS

Fourteen articles relating to the Colorado Division of Wildlife's Big
Game Research Program were published or prepared for presentation at
professional society meetings and workshops during the 1978-79 segment.
These are listed below.
Publications:
Baker, D. L., D. E. Johnson, L. H. Carpenter, O. C. Wallmo, and R. B.
Gill.
1979. Energy requirements of mule deer fawns in winter.
J. Wildl. Manage. 43(1):162-169.
Beck, T. D. I., K. R. Kincaid, and D. F. Reed.
1978. Accidental
parturition in a mule deer.
Southwest. Natural. 23:7-5-706.
Carpenter, L. H., O. C. Wallmo, and R. B. Gill.
1979. Forage diversity
and dietary selection by wintering mule deer. J. Range Manage. 32(3):
226-229.
Hobbs, N. T., and D. L. Baker.
1979. Rearing and training elk calves
for use in food habits studies.
J. Wildl. Manage. 43(2):568-570.
Reed, D. F., K. R. Kincaid,
deer fall from cliffs.

and T. D. I. Beck.
1979. Migratory
J. Wildl. Manage. 43(1):272.

mule

�-57-

Schmidt, R. L., C. P. Hibler, T. R. Spraker, and W. H. Rutherford.
An evaluation of drug treatment for lungworm in bighorn sheep.
Wildl. Manage. 43(2):461-467.

1979.

J.

Strong, L. H., and D. J. Freddy.
1979. Number of pellets per mule
deer defecation.
J. Wildl. Manage. 43(2):563-564.
Trindle, B. D., and L. D. Lewis.
1978. Methoxyflurane anesthesia in
mule deer (Odocoileus hemionus) fawns. J. Wildl. Dis. 14:519-522.
Trindle, B. D., L. D. Lewis, and L. H. Lauerman.
1978. Evaluation
of stress and its effects on the immune system of hand-reared
mule deer fawns (Odocoileus hemionus).
J. Wildl. Dis. 14:523-537.
Trindle, B. D., L. D. Lewis, and L. H. Lauerman.
1979. Techniques for
evaluating humoral and cell-mediated immunity in mule deer fawns
(Odocoileus hemionus).
J. Wildl. Dis. 15:25-31.

Papers Presented

at Society Meetings

and/or Workshops

Carpenter, L. H. 1979. Twenty four hour activity patterns of mule deer
at pasture Soc. Range Manage. Annual Meeting. Casper, WY
Freddy, D. J. 1979. Measuring heart rates of mule deer using a repeatertype telemetry system.
Internatl. Conf. Wildl. Biotelemetry.
Laramie,
WY.
Pojar, T. M.
Workshop.

1979. Population
Ft. Collins, CO.

modeling

in Colorado.

Pop. Model.

In addition to the above listed publications and papers, several manuscripts
were in the process of drafting, pre-publication review or publication.
Segment Objectives 4, 6, 10, 13, 14, 17 were all in various stages of review
and/or drafting.
Minimal or no progress was made toward publication
objectives 3, 5, 7, 11, 15, 16, 18, and 19.

Prepared

by:
R. Bruce Gill
Big Game Section Chief

��July, 1979

-59-

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

W-126-R-2

Work Plan No.
Job Title
Period

Job No.

1

Multispecies

Covered:

Big Game Investigations

Investigations:

8
----------~---------------------

Big Game Publication
Library Services
July 1, 1978 - June 30, 1979

Editing and

Personnel:
M. W. Herschopf, Dr. O. C. Cope, R. B. Gill, A. E. Anderson, D.
L. Baker, R. M. Bartmann, T. G. Baumann, G. D. Bear, T. D. I. Beck, Dr. L. H.
Carpenter, D. J. Freddy, R. E. Keiss, R. C. Kufeld, P. H. Neil, T. M. Pojar,
D. F. Reed, W. H. Rutherford, and T. N. Woodard.

ABSTRAcr
During the segment 6 Division of Wildlife publications were in various stages
of editing and publication.
Big Game Research Federal Aid Job Progress Reports
for the 1977-78 segment were written edited and printed.
Twenty-four publications
were purchased and placed on file in the Research Center Library.
Twelve
theses were ordered, duplicated, and placed on file at the Research Center
Library.
Approximately 40 copies per Big Game Research scientist were duplicated
from various Journals and periodicals and distributed to the requestor.
Twentythree literature searches were conducted for various Big Game Researchers upon
request.

��-61-

BIG GAME PUBLICATION EDITING AND
LIBRARY SERVICES

R. B. Gill

P. N. OBJECTIVE
To provide a centralized support program for Big Game Research technical
editing and library services so that Big Game Research scientists can
allocate additional time to the conduct of actual research.

SEGMENT OBJECTIVE
To provide coordinated, efficient, and economic editing and library services
to all Colorado Big Game Research programs (Federal Aid Project W-126-R).

SUMMARY OF SERVICES
Publications

Submitted

for Editing and DOW Publication

1.

Bear, G. D. 1978. Evaluation of fertilizer and herbicide applications
on two Colorado bighorn sheep winter ranges.
Colo. Div. Wildl.
Div. Rep. No. 10. 75p.

2.

Milchunas, D. G., M. J. Dyer, O. C. Wallmo, and D. E. Johnson.
1978.
In vivo/in vitro relationships of Colorado mule deer forages.
Colo. Div. Wildl. Spec. Rep. No. 43. 44p.

3.

Bear, G. D. 1979. Evaluation of bighorn transplants in two Colorado
localities.
Colo. Div. Wildl. Spec. Rep. No. 45. In Press.

4.

Carpenter, L. H., R. B. Gill, D. J. Freddy, and L. E. Sanders.
1979.
Distribution and movements of mule deer in Middle Park, Colorado.
Colo. Div. Wildl. Spec. Rep. No. 46
In Press.

5.

Kufeld, R. C. 1979. History and current status of the mule deer
population on the east side of the Uncompaghre Plateau.
Colo.
Div. Wildl. Div. Rept. No. 11. In Press.

6.

Neil, P. H., T. N. Woodard, and D. L. Baker.
1979. Procedures for
rearing wild ruminants in captivity.
Colo. Div. Wildl. Game
Info. Leaflet No. 105. In Press.

i

�-62-

Publications

Purchased

with W-126-R Funds for Research Center Library

Gold, H. J. 1977. Mathematical modeling of biological systems. An
introductory guidebook.
John Wiley and Sons, Inc. New York.
357p.
Reid, J. T. (Chmn) 1968. Body composition
Acad. Sci. Publ. 1598. 521p.

in animals and man.

Natl.

Gill, J. L. 1978. Design and analysis of experiments in the animal
and medical sciences.
Vol. 1. Iowa State Univ. Press, Ames, IA
50010. 409p.
Gill, J. L. 1978. Design and analysis of experiments in the animal
and medical sciences.
Vol. 2. Iowa State Univ. Press, Ames,
IA 50010.
301p.
Gill, J. L. 1978. Design and analysis of experiments in the animal
and medical sciences.
Vol. 3. Iowa State Univ. Press, Ames, IA
50010.
173p.
Wang, L. C. H., and J. W. Hudson.
(eds.). 1978. Strategies in cold.
Academic Press, New York.
Natural torpidity and thermogenesis.
715p.
Arnold, G. W., and M. L. Dudzinski.
1978. Ethology of free-ranging
domestic animals.
Elsevier Scientific Publishing Co., Amsterdam.
198p.
International Mountain Goat Symposium.
1977. Proc. 1st Inter. Mountain
Goat Symposium, Kalispell, Montana, Feb. 19. W. Samuel and W. G.
Macgregor (eds.) British Columbia, Fish and Wildl. Branch (Victoria).
243p.
Harborne, J. B. 1977. Introduction
Press, New York.
243p.

to ecological

biochemistry.

Academic

Keeler, R. F. et al., eds. 1978. Effects of poisonous plants on livestock.
Joint United States-Australian Symposium on Poisonous Plants, Utah
State University, 1977. Academic Press, New York.
600p.
Kleiber, M. 1975. The fire of life; an introduction to animal energetics.
Rev. ed., Huntingdon, N.Y., Robert E. Krieger Publ. Co. Inc. 453p.
Bekoff, M., ed. 1978. Coyotes;
biology,
Academic Press, N. Y. 384p.
Gladfelter, H. L. 1978.
by radio telemetry.
23. 27p.

behavior,

and management.

Movement and home range of deer as determined
Iowa Consr. Comm., Iowa Wildl. Res. Bull. No.

Masters, R., and Northeast Deer Study Group.
1978. Deer trapping, marking
and telemetry techniques.
State Univ., of N.Y., College of Environmental
Science and Forestry.
Newcomb, New York.
72p.

�-63-

Sosebee, R., and H. A. Wright (eds.). 1977. Research highlights
brush and weed control; range and wildlife management 1977.
Texas Tech. Univ., Lubbock, Texas.
54p.

noxious
V. 8.

Clarke, R. T. J., ed. 1977. Microbial ecology of the gut, edited by
R. T. J. Clarke and T. Bauchop.
Academic Press, New York.
410p.
Fletcher, John L. ed. 1978. Effects of noise on wildlife, edited by
John L. Fletcher and R. G. Busnel.
Academic Press, New York.
305p.
Follis, T. B. 1972. Reproduction and hematology of the Cache elk herd.
Ph.D. Dissertation, Utah State University.
Utah Div. Wildlife
Resources.
Publ. No. 72-8. 133p.
Hutchinson, G. E. 1978. An introduction
Univ. Press, New Haven.
260p.
Young, E., ed. 1975.
Books, Hollywood,

to population

ecology.

The capture and care of wild animals.
FL. 224p.

Yale

Ralph Curtis

Gruell, G. E., and L. L. Loope.
1974. Relationships among aspen, fire,
and ungulate browsing in Jackson Hole, Wyoming.
Forest Service,
U.S.D.A. and National Park Service, U.S. Dept. Interior.
33p.
Hanson, H. C. 1962. Dictionary of ecology.
(Crown Publishers), New York.
382p.

Philosophical

Library

Martin, R. E., J. E. Dealy, and D. L. Caraher.
(eds.) 1977.
Proceedings of the western juniper ecology and management workshop,
Bend Oregon, January 1977. U.S. Forest Service, Pacific Northwest
Forest and Range Experiment Station, Portland.
U.S. Forest Service
Gen. Tech. Report PNW-74.
177p.
Schmidt, J. L. (ed.) 1978. Big game of North America; ecology and
management, compiled and edited by John L. Schmidt and Douglas L.
Gilbert.
A Wildlife Management Institute Book, qtackpole Books,
Harrisburg, Pa. 494p.

Theses Ordered,

Duplicated,

and Placed on File at Research

Maloiy, G. M. o. 1968. The physiology
the red deer (Cervus elaphus L.).
Aberdeen, Scotland.
192p.

Center Library

of digestion and metabolism in
Ph.D. Thesis, Univ. of Aberdeen,

Morton, M. A. 1976. Nutritional values of important mule deer winter
forage plants in the Bridger Mountains, Montana.
M.S. Thesis,
Montana State Univ., Bozeman, Montana.
104p.
McBride, R. T. The status and ecology of the mountain
concolor stanleyana) of the Texas-Mexico Border.
SuI Ross State University.
160p.
(1976?)

lion (Felis
M. S. Thesis,

�-64-

Simpson, A. M. 1976. A study of the energy metabolism
cycles of captive red deer. Ph.D. Thesis.
194p.

and seasonal
Univ. of Aberdeen.

McCollum, M. T. 1973. Habitat utilization and movements of black bears
in southwest Oregon.
M.S. Thesis, California State University,
Humboldt, Arcata.
67p.
Moss, H. H. A study of black bear in the San Gabriel Mountains.
Thesis, California Polytechnic University, Pomona.
63p.

H. S.

Olmsted, C. E. III. 1977. The effect of large herbivores on aspen
in Rocky Mountain National Park. Ph.D. Thesis, University of
Colorado, Boulder.
141p.
Gipson, P. S. 1972. The taxonomy, reproductive biology, food habits
and range of wild Canis (Canidae) in Arkansas, Ph.D. Dissertation,
University of Arkansas.
186p.
(Courtesy A. E. Anderson)
Humphreys, I. D., III. 1977. Hunter selectivity of mule deer harvested
during the San Andreas special season hunts 1973-1976.
M.S. Thesis,
New Mexico State University, Las Cruces.
22p.
Lucich, G. C. 1977. Plant taxa eaten by doe and fawn mule deer in
northcentral Colorado.
M.S. Thesis, Colorado State University,
Fort Collins.
35p.
Sheehy, D. P. 1975. Relative palatability of seven Artemisia taxa to
mule deer and sheep. M.S. Thesis, Oregon State University, Corvallis.
147p.
Sheriff, S, L. 1978. Computer model for mountain lion populations.
Thesis, Colorado State University, Fort Collins.
287p.

List of Literature
Center Library

Searches

Performed

for Big Game Researchers

Bears
Black Bears - principally North American references
Cervus canadensis
Condition factors of deer
Parturition of deer, elk and moose
Retrospective search on Odocoileus hemionus
Deer enclosure studies
Alaskan black-tailed deer
Deer physical characteristics and condition (6691F)
Deer physical characteristics and condition (6691)
Coyote
Mountain goats
Mountain lion, cougar, puma, Felix concolor

M.S.

by Research

�-65-

Big game forage requirement in the western U.S
Immobilization and tranquilizers for mammals
Radio telemetry of wildlife
Pinyon-juniper management and mule deer
Pinyon-juniper deer habitat
Effects of highway, road, and pipeline construction on wildlife
Improvement of wildlife habitat by soil fertilization
Habitat improvement techniques (015413)
Effects of power lines on big game
Heart rate and energy expenditure

Reprint Duplication
In addition to the above listed services, the Research Center Library also
duplicated approximately 40 articles per Big Game Research Scientist and
technician or approximately 700 total articles for the segment for the
Big Game Research Section.

Prepared

by:
R. Bruce Gill
Big Game Section Chief

��July, 1979

-67-

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~----------

Project No.

2

Work Plan No.
Job Title Nutritional
Period Covered:

Big Game Investigations

W-126-R-2

Job No.

Basis for Quantifying

July 1,

1

---------------------------------

Capacity of Winter Ranges to Support Deer

1978 to June 30, 1979

Personnel:
Len H. Carpenter, D. J. Freddy, Larry L. Strong, Steve C. Torbit,
Linda E. Sanders, Linda J. Robinson, Timm J. Kaminski.

ABSTRACT
Fifteen adult mule deer were divided into 3 groups of 5 deer and subjected
to 3 levels of food intake. Weight losses corresponded to decreasing food
levels. Deer subjected to the lowest food offering (291 g/day) exhibited
severe nutritional stress after 42 days. Urine output was greatest for
deer in the low food intake treatment.
Determination of daily maintenance
energy requirements will be made when laboratory analyses of feed, urine,
and fecal samples are completed.

��-69-

NUTRITIONAL BASIS FOR QUANTIFYING
CAPACITY OF WINTER RANGES TO SUPPORT DEER
Len H. Carpenter

P. N. OBJECTIVE
Develop procedures
to support deer.

for quantifying

the capacity

of Middle Park winter

ranges

SEGMENT OBJECTIVES
1.

Estimate

energy requirements

2.

Estimate

energy costs associated

3.

Develop sampling methodology for estimating
availability of deer forage in winter.

annual production

4.

Test the efficacy
carrying capacity

of a computer simulation
of winter ranges.

model for estimating

5.

Estimate

requirements

protein

METHODS

of adult mule deer during winter.
with selected

activities

of mule deer.
and

of mule deer fawns in winter.

AND MATERIALS

Segment objectives 3 and 5 are being investigated by Larry L. Strong and
Steve C. Torbit respectively.
Both are graduate students at Colorado State
University.
Detailed descriptions of methodology and results of their
studies will be presented next segment as Graduate of Science Theses.
Work on objective two involved publication of previous results.
A manuscript
entitled "Heart rate as a predictor of energy expenditure of mule deer" has
been submitted to the Journal of Wildlife Management.
Most of the effort
expended by the principal investigator was devoted to segment objective 1.
Description of this work follows.
Details of simulation modeling will be
presented in later segments.

Estimating

Energy Requirements

of Adult Mule Deer in Winter

Methodology of this experiment closely followed that of Baker et al. 1979.
Fifteen, 18-month-old mule deer were divided into 3 groups of,S deer each.
Deer were ranked from lightest to heaviest and randomly allotted into 5
successive trios. This was done to disperse various sizes of deer equally
over all treatments.
The 3 treatments were feed levels and were as follows:

�-70-

1) Ad libitum (as determined from 14 days of intake for all 15 deer),
2)
one-half of ad libitum,
and 3) one-fourth of ad libitum.
The food offered
was a concentrate alfalfa pellet (Pawnee Special).
Composition analysis
of this ration is presented by Baker et al. 1979.
Deer were maintained on these feed levels for 10 weeks beginning January 1,
and continuing until March 12, 1979. Each deer was randomly assigned to
an individual isolation pen for duration of the experiment.
Every other
week, beginning with the second week of the experiment, one deer (randomly
chosen) from each treatment was assigned to a digestion cage for complete
nitrogen and energy balance experiments.
Two days were for acclimization
only and no data (other than intake) were recorded.
During the next 5
days all feces and urine were collected daily and intake was continually
monitored on a daily basis.
Every Monday during the experiment all deer
were weighed.
All deer were offered snow ad libitum.
Quantities of feces and urine were measured and recorded daily, however,
weekly outputs were of most interest.
Feces weights (wet) were recorded
to the nearest gram. A 10 percent aliquot of each day's sample was combined
and frozen for future analyses of energy, and protein and dry matter.
Daily
urine samples for each deer were filtered through cheese cloth, quantity
in milliliters determined, and a 10 percent aliquot taken composited and
Lab analyses
frozen.
Urine was acidified with H2S04 during collection.
for energy and protein for each urine and fecal sample will be made.
In
addition, measurements of creatinine will be made on each urine sample.
Daily intakes were monitored for each deer by weighing orts. Each deer in
a treatment received the same amount of feed daily for the 70-day period.
From the 14-day pre-trial period it was determined that mean intake
for the 15 deer was 1165 + 64 (SE) g. Consequently deer receiving 50
percent feed level (Treat;ent B) received 583 g daily and deer receiving
25 percent of ad libitum received 291 g each day. The five deer assigned
to the ad libitum level were offered 2500 g daily.
This was done to
compensate for increased intakes that might occur during the 10-week period.
Digestible and metabolizeable energy and protein and dry matter digestibility
coefficients will be determined for each deer following methods outlined by
Baker (1979). Comparisons will be made between body weight losses and
creatinine levels in the urine of each deer. Creatinine levels will also
be compared to creatinine levels measured in mule deer where body composition
using tritiated water has been determined.

RESULTS AND .DISCUSSION
Much of the laboratory analyses have not been completed
of the results will be presented in later segments.

consequently

most

There were no significant differences (P &lt; .05) in weekly intakes among the
5 deer in treatment A during the experiment.
Average daily intake for the
5 ad libitum deer were 925.9 + 28.7 (SE) g. This was a 240 g per day decrease
as~ompared
to pre-trial meas~rements.
All deer in Treatments Band C
essentially consumed all food offered each day.

�-71-

All deer in each treatment lost weight during the experiment (Fig. 1). As
expected deer in Treatment C lost the most weight.
The relatively large
weight loss of deer in Treatment A is not fully understood.
These deer
did decrease their intakes during the course of the experiment.
Deer
in Treatment C lost weight very quickly and steadily (Fig. 1). Two deer
in this group starved to death during the experiment and one was removed
from the study and refed and one died one week after the experiment
concluded.
The first deer starved to death after 7 weeks; the second
deer starved after 9 weeks and the third deer was removed from the study
after 7 weeks.
Removal of these deer from the experiment results in the
apparent weight gain in Treatment C deer in Figure 1. Overall the average
percent body weight loss for Treatment C deer was 28.1 ± 9.1 (SD) %.
Deer in Treatment B lost weight steadily during the experiment but only
one deer in this group exceeded a loss of 20 percent of initial body weight.
Percent of initial body weight losses for Treatment B deer varied from
3.1 to 21.5 with a mean of 17.1 ± 3.1 (SD). Greater variability was
exhibited by deer in Treatment A with regard to body weight changes.
All
deer lost weight with one deer in this group losing 25.3 percent of initial
body weight.
Minimum weight loss exhibited by a deer in this group was
8.6 with a mean percent of 14.7 ± 6.5 (SD) occurring for the entire treatment.
Weight losses for Treatment A corresponded to weekly intakes.
Decrease
in intake by the ad libitum deer is not understood but may be due to
absence of a roughage to enhance palatability of the concentrate over an
extended period of time.
Certain differences in daily urine volumes by treatment were recorded.
Some
problems were encountered by blowing and drifting snow which accumulated on
urine trays and inflated daily volumes.
This occurred during two days for
one deer in Treatment A and one deer in Treatment C
The degree of dilution
was impossible to determine so no correction of this error was made.
Average daily urine volume for Treatment A deer was 976.8 ± 145.7 (SE) mI.
Daily urine volumes for Treatments Band C were 725.3 ± 56.5 (SE) and
1607.5 ± 197.1 (SE) ml, respectively.
It appears that nutritionally
stressed deer increase daily urine output.
This could be expected if body
catabolism is occurring and body catabolic products must be excreted by the
kidneys.
Subjectively, Treatment C deer ate more snow each day but it did
not appear the difference in snow intake was as great between treatments
as was urine output.
Perhaps laboratory analyses of protein and creatinine
in urine samples will aid interpretation of daily urine volumes.

Simulation

of Mule Deer Carrying

Capacity

Much of the simulation modeling work accomplished in this segment was
devoted to "tuning" the model specifically to the unique physiological inputs
of mule deer. The original ruminant nutritional model constructed by
Natural Resources Ecology Laboratory scientists at Colorado State University
was constructed largely from literature available regarding domestic ruminants.
Several runs were made substituting published values for mule deer forage

�-72-

intake, forage chemical constitution, forage digestibilities, daily energy
budgets, and nitrogen excretion rates to evaluate the model responses.
Sensitivity analyses were run to identify the most critical input values.
Then adjustments were made in the model so that it more closely simulated
deer nutritional physiology.
Next segment activities will involve simulating
deer nutritional responses to variations in forage intake quantities and
qualities.
Then empirical experiments will be designed to test the model
predictions.

LITERATURE

CITED

Baker, D. L., D. E. Johnson, L. H. Carpenter, O. C. Wallmo, and R. B. Gill.
1979. Energy requirements of mule deer fawns in winter.
J. Wildl.
Manage. 43(1):162-169.

Prepared

by:
Len H. Carpenter
Wildlife Researcher

�-73-

--15

• _.0.-.,.

...............
n · · 0-._.-0Treatment
~
'-'-0...

-

•z••

\
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UI

U
II:IJII:
UI

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\

-5

D.

o, ....

\
.... ....

...

\

,

"'0

'\
'\

-

10

\

,,

0......

" , Treatment

,,

0,
'\

15

'\

q

-

C)
Z
C

,,

,

z
UI

\

'0

&gt;Q

o

A

'\
'\

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20

'0----0 ....

:J:

C
D

"".... , /

/'\

\

//

o

u

\

\\
\

25

\
\
\
\

b
2

pretrial

1

2

3

4

5

6

7

8

trial

WEEK

Figure 1.

Mean weekly weight change (percent) compared to initial weight
of adult mule deer on 3 different levels of food intake.

9

10

��July,

-75-

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~--~----------

Project

No.

W-126-R-2

Plan No.

2

Work

1979

Deer-Elk
Job No.

Investigations
2~ __ ~

_

Job Title __~S~n~o~wm~~o~b~i~l~e~H~a~r~a~s~sm~e~n~t~o~f~M~u~l~e~D~e~e~r~o~n~C~o~l~d~._W~i~n~t~e~r~
_
Period

Covered:

Personnel:
D. Bowden,

July 1, 1978 - June 30, 1979

D. Freddy,
L. Sanders.

T. Kaminski,

W. Bronaugh,

L. Robinson,

L. Carpenter,

ABSTRACT

Five wild female deer were trapped and equiped with heart rate telemetry
systems.
These deer were systematically harassed by people and snowmobiles.
Observed reactions of deer will be presented in next segment's report.
Heart rate measurements on tame deer at pasture during winter 1978 are being
incorporated into a manuscript for publication.
Developmental
progress and
performance of the heart rate system employed in this study have been
summarized in a publication to be published in July, 1979.

��-77-

SNOWMOBILE HARASSMENT OF MULE DEER
ON COLD WINTER RANGES

David J. Freddy

P. N. OBJECTIVE

Evaluate whether snowmobile activity on winter ranges inhabited by deer
decreases the ability of deer to survive winter by modifying activities
of deer so as to significantly increase energy expended.

SEGMENT OBJECTIVES
1.

Monitor heart rates of deer during various daily activities
energy costs of these activities during winter.

2.

Ascertain

3.

Evaluate observed reactions of deer to harassment in terms of
appropriate heart rate measurements and corresponding energy costs.

4.

Evaluate the energy cost of harassment to deer in relation
estimated energy budget of deer (concurring study).

5.

Examine alternatives
harassing deer.

activities

of deer responding

to minimize

to prescribed

potential

detrimental

to quantify

harassment.

to a total

effects of

METHODS AND MATERIALS
Five wild female deer, yearling and older, were trapped near the Junction
Butte Research Center and equipped with heart rate telemetry systems (see
Freddy 1978 for trapping and surgical procedures).
Deer were released
and heart-rates were monitored opportunistically.
These 5 deer along with 4 neck-banded deer were harassed by 1 and 2 persons
walking and 1 or 2 snowmobiles moving on prescribed routes.
Reactions of
these deer were visually and telemetrically monitored from 4 observation
huts placed at strategic points on Junction Butte.
At least 2 observers
monitored deer during harassment bouts.

RESULTS AND DISCUSSION
1979
Data from harassment bouts are in the process of computer
will be discussed in next segment's report.

analysis

and

�-78-

1978
Heart rate data collected from 3 tame deer at pasture and 2 wild deer
is being assimulated into a manuscript for publication.

1977-1979
Developmental progress and performance of the heart rate telemetry
system employed in this study has been summarized in the publication:
Freddy, D. J. 1979. Measuring heart rates of mule deer using a repeatertype telemetry system.
In Proc. Second Intern'l Wildl. Biotelemetry
Sym., Laramie, Wyo., July, 1979. In Press.
Copies of the manuscript
available upon request.

LITERATURE

CITED

Freddy, D. J. 1978. Snowmobile harassment of mule deer on cold winter
ranges.
Colo. Div. Wildl. Game Res. Rep. July, Part 2: 137-144.

Prepared

by:

/'

")

-L

. "

~(iM--t--ci . ~~ '~T':.(_ti~£'~vDavid J. Freddy
Wildlife Researcher

0

�-79-

July, 1979

JOB PROGRESS REPORT

State of

COLORADO
--------~~~~~-----------

Project No.
Work Plan No.
Job Title

Big Game Investigations

W-126-R-2

Job No.

2

3

Deer Investigations--EstimatingParameters
of Piceance Basin
--------------~~-------nD~e~e~r~P~o~p~u~lra~t~i-o=n~D~y~n~a~m~1'·c-S~------------------

Period Covered:

July 1, 1978 - June 30, 1979

Personnel:
R. M. Bartmann, T. G. Baumann, Dr. D. C. Bowden, Dr. L. H.
Carpenter, W. Erickson, K. Clowser, D. Flenthrope, M. Hamady, E. Killinger,
R. Penn, and C. Perron.

ABSTRAcr
The 1978-79 winter deer census estimate in Unit 22 was 28,994 + 4,788
deer (90 percent confidence level) compared to the predicted p~pulation
of 36,993. The buck:doe and fawn:doe ratios were estimated by two
methods; one was the regular non-random procedure and the other a stratified
random quadrat technique.
Resultant ratios for both methods were similar,
but the quadrat method allowed placement of confidence limits about the
estimated ratios which, at the .10 level, were for the buck:doe, 13.3:100
2.6, and for the fawn:doe, 58.2:100
5.2. Adjustment of the fawn:doe
ratio for antlerless harvest yields a pre-season ratio of 55:100. The
1978-79 winter was comparable in severity to the 1972-73 winter. As a
res1llt, estimated deer winter mortality was also similar, 44 percent
compared to 41 percent for 1972-73. The estimated deer loss was 16,233
± 3,990 (90 percent confidence level) and was comprised of 11,302 fawns,
4,459 adult does and 571 adult bucks. The 1978 deer harvest for all
seasons was 9,277. Of 5,500 either-sex permits available for Unit 22,
only 3,388 were issued.

±

_

±

��-81-

ESTIMATING

PICEANCE

DEER STUDY

PARAMETERS

OF POPULATION

DYNAMICS

R. M. Bartmann

P. N. OBJECTIVE
To develop and test a method for estimating
deer populations in the Piceance Basin

density of over-winter

mule

SEGMENT OBJECTIVES
1.

To analyze

and publish results of deer distribution

studies.

2.

To estimate the size of the wintering
Management Unit 22.

deer population

in Game

3.

To estimate the sex and age structure
in Game Management Unit 22.

of the wintering

deer population

4.

To estimate

the annual productivity

5.

To estimate

the annual winter mortality

6.

To estimate annual hunter harvest

of the Piceance

deer population.

of the Piceance

of the Piceance

deer population.

deer population.

METHODS AND MATERIALS
Deer Density
See Bartmann
Population

(1974a).

Structure

The 1978 post-season deer sex and age structure data were collected by two
procedures.
One was the previously used non-random method whereby number
of deer classified was maximized by spending most of the flight time over
flying higher deer density areas (Bartmann 1974a). The second method
entailed classifying deer on mile2 (2.6 km2) quadrats located in a
stratified random manner in Game Management Unit 22. The expected deer
distribution for early December was delineated on a map of the Piceance
Basin.
The total area was then divided into high and low density areas
based on expected deer densities.
The high density area, containing 2
532-miles2 (1378 km2) was arbitrarily divided into 5, 106 or 107-mile
(275 or 277 km2) strata.
The low density area, containing 283-miles2
(733 km2) was divided into 3, 94 or 95-mile2 (244 or 246 km2) strata.

�-82-

2
2
Six mile
(2.6 km ) quadrats were randomly selected from each of the 8
strata for a total of 48 quadrats.
The only restriction was that no 2
quadrats could be contiguous along their sides although corners of
diagonally adjacent quadrats could touch. Quadrat boundaries were drawn
on aerial photos which were used to locate and define quadrats during
the classifications.

Productivity
See Bartmann

(1975).

Winter Mortality
See Bartmann

(1974a and 1975).

Hunter Harvest
See Bartmann

(1974b).

RESULTS AND DISCUSSION
Publication

of Deer Distribution

Work

Attempts to summarize the deer distribution and banding data have been
unproductive due to the limited amount of time that could be devoted to
this job. Plans are to publish these data in a Division Special Report.

Deer Density
The 1978-79 winter deer census was made January 3-5, Deep snow accumulated
during December causing rapid changes in deer distribution and, consequently,
an earlier than usual census. A Hiller 12EJ3 helicopter with a Soloy
turbine conversion was used with Dr. Len H. Carpenter again serving as
second observer.
2
2
The 1,304 deer counted converts to 43.6 deer/mile
(2.6 km ) or
28,994 ± 4,788 total deer (90 percent confidence level) on the Piceance
winter range (Table 1). This figure is considerably lower than the 36,993
deer population that was predicted (Table 2). As usual, several possible
reasons can be offered for the discrepancies. some of which are inherent
in the sampling design.
In addition, there seemed a reluctance by deer
to run from the helicopter as they did in past years.
This was also
noticed during the post-season classifications in mid-December.
Whether
or not this condition actually existed and what effects it may have had on
Ithe count is only speculation.
Another consideration is that there may be
an upper limit of deer density at which counting efficiency begins to
decrease.
Unless the 1977-78 population estimate was too high, there

�-83-

Table 1.
Number of deer counted on 120 \-mile2 (0.6 km2) quadrats on the
Piceance winter range in Game Management Unit 22, January 3-5, 1979.
Quad.

Deer

Quad.

Deer

Quad.

Deer

Quad.

Deer

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30

19
24
25
13
13
14
19
7
0
1
0
19
0
8
19
19
32
28
29
33

31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60

30
29
8
20
13
27
1
4
5
6
0
0
20
0
29
8
6
1
0
0
0
3
0
0
7
12
3
0
0
0

61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90

5
0
31
33
9
54
19
21
0
10
15
0
3
3
0
7
6
12
0
38
5
0
0
3
0
0
0
14
15
19

91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
llO
111
112
ll3
ll4
115
116
ll7
ll8
119
120

13
2
3
15
8
4
4
0
0
0
16
13
10
6
3
4
0
0
1
1
36
3
2
22
7
4
47
1
19
17

II

9
30
14
40
24
33
0
0
6

n = 120

90% Confidence

Interval

IX = 1,304
x

10.9

10.9 ± (1.658) (1.09) = 10.9 + 1.8

s
sx

11.98

Total POEulation

1.09

C.Il.= llO%

28,994

± 4,788

Estimate

�-84-

Table 2. Calculations to predict
Game Management Unit 22.

the 1978-79 winter deer population

in

Bucks

Does

Fawns

Total

3,206

18,956

12,684

34,846

96

476

1,237

1,809

1978 summer population

3,110

18,480

11 ,447

33,037

Fawns apportioned

5,723

5,724

8,833

24,204

Est. 1977-78 winter pop.
Est. 1977-78 winter mortality

Adjusted

50:50

1978 summer pop.

1978 fawn prod. est.
(55 fawns:100 does)

Predicted

13,233

pop.

8,833

24,204

13,233

46,270

estimate

7,478

1,648

151

9,277

1,355

22,556

13,082

36,993

2,811

21,641

12,541

36,993

1978 pre-season
1978 harvest

33,037

1978-79 winter pop.

Predicted population
corrected for 1978
post-season buck:doe:fawn
ratio (13:100:58)

�-85-

seem no logical reasons for the population not to have increased over the
past year. Thus the predicted population figure seems the most appropriate
to use for the present.

Population

Structure

The two classification methods yielded similiar fawn:doe and buck:doe
ratios (Table 3). Nearly 50 percent more deer were classified/hour with
the non-random method, but this was partly due to spending a greater
proportion of time in higher deer density areas plus being able to
classify deer continually without the "dead" time needed to fly between
quadrats.
If only the time spent classifying deer on quadrats is
considered, the number of deer classified/hour is more equitable between
the two methods.
The main advantages of the quadrat method is the ability to place
confidence limits about the estimated ratios, and the applicability of
the data to the entire deer population, including unsampled individuals.
Precision of the fawn:doe and buck:doe ratios at the 90 percent confidence
level was + 9 and + 20 percent, respectively.
This is close to that
achieved by Freddy-(1977) (+ 10 and + 21 percent, respectively) when he
sampled a 791-mile2 (2049 k;2) area in the Piceance Basin with 9 strata
and 3 mile2 (2.6 km2) quadrats per stratum.
That he achieved similar
precision with 21 fewer quanrats may reflect the difference in deer
distribution the two years.
:Because deer distribution during the 1978
classifications was adversely affected by early deep snow, the comparison
will be repeated in 1979.

Productivity
The 1978 deer season in Unit 22 included issuance of 3,388 of 5,500
available either-sex permits.
The Game Management Section's random
harvest survey results for all seasons show 1,648 does and 151 fawns
harvested.
These figures include an arbitrary inflation of 25 percent
to acknowledge
wounding loss and illegal kill. The post-season fawn:doe
ratio of 58:100 was then adjusted to provide a pre-season ratio of 55:100
for a pre-season production of 13,233 fawns.

Winter Mortality
The severity of the 1978-79 winter was similar to that of 1972-73.
Snow
began accumulating in early December, and low temperatures prevented
much melting and settling of the snowpa~k.
The mean snow depth at the
Little Hills Station for December through February of 15.6 (39.6 cm)
inches was 2.8 (7.1 cm) inches greater than for the same period in
1972-73, but the mean temperature, 23.6°F, was 2.30F higher.
The

�-86-

Table 3. Comparison of non-random and stratified random quadrat methods
for estimating the sex and age structure of the mule deer population in
Game Management Unit 22, December, 1978.
Method
Non-random

Parameter

Total fyling time
Total deer classified
Deer classified

per hour

9.3 hours

Stratified

Random Quadrats

15.0 hours

1,193

1,292

128

86
12 minutes

Mean time per quadrat
Total bucks

87

100

Total does

697

750

Total fawns

409

442

Buck:doe

ratio

12.5:100

13.3:100

+

Fawn:doe

ratio

58.7:100

58.2:100

+ 5.2

2.6J

(P &lt; .10)

�-87-

severity of the winter is reflected in the mortality data with 172 deer
found on the 56 ~ x ~-mile (0.6 x 1.3-km) sample units.
This computes
to a mean 3.07 deer/sample unit or 16,233 ± 3,990 total dead deer (90
percent confidence level) on the winter range. Precision of the estimate
is + 24 percent which is the best achieved to date.
Age and sex composition of the carcasses found with unknowns apportioned
among the categories is 69 percent fawns, 27 percent adult does and 4
percent adult bucks.
This compares closely with the 1973 percentages
of 70, 25 and 5 for the same classes, respectively.
Numerical breakdown
of the 1978-79 mortality is 11,302 fawns, 4, 459 adult does and 571 adult
bucks.
This is a 44 percent reduction of the 36,993 predicted winter
population.

Hunter Harvest
The 1978 regular deer season structure was the same as in 1977. A 5-day
deer-only season ran October 14-18 and an II-day combined deer-elk season
ran November 4-14. A post-season was held December 2-6 for holders of
unfilled either-sex permits for Unit 22. Of 5,500 permits available,
2,200 were for the separate season and 3,300 for the combined season.
Only, 2,038 separate season and 1,350 combined season permits were
actually issued.
The archery season, September 2-24 and the muzzleloader season, September 9-17 were both either-sex.
An estimated 9,277
total deer were harvested during all seasons (Table 4).
As usual, the report of the 1978 check station results is inconsistent
with previous years.
Information from all check stations operated in
1978 is lumped together and age class data by sex are not separated for
the Idaho Springs station.
Predicted yearling percentages of bucks and
does in the adult pre-season population were 65 and 24, respectively.
Percentages from pooled check station data are similar, 59 and 22,
respectively (Table 5).

LITERATURE

CITED

Bartmann, R. M. 1974a. Piceance deer study-population density and
structure.
P. 363-370.
In Game Research Report. Colo. Div. of
Wildl., Denver.
3(Part 2):185-398.
(Proc.).
Bartmann, R. M. 1974b. Piceance deer study-productivity
and mortality.
P. 371-380.
In Game Research Report.
Colo. Div. of Wildl., Denver.
3(Part 2):185-398.
(Proc.)
Bartmann, R. M. 1975. Piceance deer study-productivity
and mortality.
P. 371-380.
In Game Research Report.
Colo. Div. of Wildl., Denver.
3(Part 2):195-504.
(Proc.).
Freddy, D. J. 1978. Experimental deer inventory-Piceance
Basin-Northwest
Region.
P. 245-263.
In Game Research Report, Colo. Div. of Wildl.,
Denver.
3(Part 2):137-298.
(Proc.).
,,/l-

Prepared

.

/J

by:/C-.---1.
J •.•.
l..-&lt;4:' 1&gt;-1~)/.:~{,rYu';.v .....
'i
Richard M. Bartmann
Wildlife Researcher

�-88-

Table 4.

Summary of the 1978 deer harvest

II

in Unit 22 for all seasons.-

Bucks

Does

Season

Adult

Fawns

Archery

15

0

Muzzle-loader

26

Antlered-only
E·lt h er-sex-21

Adult

Fawns

Total

0

0

15

0

23

0

49

3,011

0

0

0

3,011

669

26

996

54

1,745

Antlered-only

3,111

0

0

0

3,111

E·lt h er-sex-2/

646

22

629

49

1,346

7,478

48

1,648

103

9,277

Separate

Combined

All

1/All figures
wounding

include an arbitrary
loss and illegal kill.

inflation

liThe separate and combined season either-sex
kill data from the December post-season.

Table 5. Deer check station results
seasons in Game Management Unit 22.

of 25 percent

harvest

to acknowledge

estimates

for the 1978 separate

also contain

and combined

Sex

Mature

Yearling

Fawn

Unknown

Total

Male

252

369

61

181

863

Female

292

83

39

77

491

�GAME

MANAGEMENT UNIT

22

~-....•

-PICEANCE-

~

,i1=
I·

,

~

\

.
• \.e..c.

{

~\\

,-

7

Fig. 1. Locations of two general study areas on the Piceance juniper-pinyon-shrubland
winter range for conducting mule deer winter forage selection trials.

I
00
\0
I

��-91-

July, 1979

JOB PROGRESS REPORT

State of

COLORADO
------------------------------

Project No.

W-126-R-2

Big Game Investigations

4
2
Work Plan No.
Job No.
Deer Investigations--Food Habits Technique Development
Job Title __~a~n~d~~F~o~r~a~gce~P~r~e~f~e~r~e~n~c~e~s~o~f~M~u~1~e~D==e~e~r~i~n~t~h~e~P~i~c~e~a~n
_

---------------------------------

Period Covered:

July 1, 1978 - June 30, 1979

Personnel: R. M. Bartmann, T. G. Baumann, Dr. D. C. Bowden, G. Bryant,
J. Cribari, W. Erickson, and L. Logren.

ABSTRACT
Forage selections of 2 groups of tame mule deer were made on 2 winter
ranges:
sagebrush steppe and juniper-pinyon-shrubland.
The results are
being prepared for publication in the Journal of Range Management.
Five
series of deer grazing trials were conducted from October, 1978 through
April, 1979 on the Piceance juniper-pinyon-shrubland
winter range. Forage
samples were collected for. all species comprising 2 percent or more of the
total bites during each period on each study area for bite/weight conversions
and nutritional analyses.
The grazing data are still being prepared for
computer summarization.

��-93-

•

DEER INVESTIGATIONSFOOD HABITS TECHNIQUE DEVELOPMENT AND
FORAGE PREFERENCES OF MULE DEER IN THE PICEANCE

BASIN

R. M. Bartmann

P. N. OBJECTIVE
To evaluate possible bias in the use of tame deer in forage preference
studies and to estimate forage preferences of mule deer on pinyon-juniper
winter range in the Piceance Basin.

SEGMENT OBJECTIVES
1.

Analyze data from previous food habits techniques
and publish pertinent results.

development

2.

Prepare a detailed
forage preferences

3.

Maintain

4.

Select specific

5.

Prepare

6.

Conduct deer foraging

7.

Rate forage availability

8.

Test necessity

9.

Conduct deer grazing comparison trials to evaluate
nutritional background on forage selections.

10.

Tabulate

work

study plan for estimating fall-winter-spring
of deer in the Piceance Basin.

and train tame deer.
sites for food habits sampling.

an herbarium

of plants from the Piceance

Basin.

trials.
during deer foraging

of converting

data and prepare

trials.

bite data to a weight basis.

annual progress

influence

of

report.

METHODS AND MATERIALS
Tame Deer Grazing Evaluations
Two groups of tame mule deer were hand-raised and maintained at Division
facilities near Kremmling and at the Little Hills Game Experiment Station
near Meeker.
The former area is a sagebrush steppe winter range and the
latter a juniper-pinyon-shrubland
winter range.
Neither deer group had
any previous foraging experience on the other's area.
Comparisons of

•

�-94-

the forage selections of both groups on both areas were made in October,
1978. A paper covering the results of this work is being prepared for
publication in the Journal of Range Management.

Plant Collections
Only token plant collecting was done in 1978 but this work is being
continued by a temporary employee in 1979. Emphasis is being placed on
collecting and identifying unknown species eaten by tame deer during
winter grazing trials and getting better specimens of some species
previously collected as well as collecting any new species encountered.
While over 150 plants have already been collected and tentatively
identified, identifications have not been verified nor have herbarium
mounts been prepared.
Scientific names are being standardized with
those available through the computerized Plant Information Network at
Colorado State University.

Tame Mule Deer Maintenance

and Training

Based on previous tame deer grazing evaluations at Little Hills, it was
decided that maximum exposure of deer to be used for grazing studies to
native forage was preferred to artificial feeding in a pen. Consequently,
all the deer were moved to an approximate 100 acre (41 ha) pasture in
April, 1978. They were kept there, except during grazing trials, until
the 1978 hunting seasons when they were moved back to the pens at the
Little Hills headquarters for safety reasons.
In mid-December they were
returned to another 140 acre (57 ha) pasture where they were kept the
rest of the winter when not being used in grazing trials.
After the
last trials in April, 1979, they were moved back to the original pasture
for the rest of the summer.
While in the pastures, the deer were supplied
small amounts of alfalfa hay and concentrate feed to enable checking their
condition each day and to keep thenl used to loading in a truck.

Mule Deer Forage Selections

on Winter

Range

Forage selections of tame mule deer were recorded during 5 periods on
2 study areas during the 1978-79 winter.
Grazing periods coincided with
deer occurrence on various portions of the winter range from early October
through April.
A sixth scheduled grazing period in early January was
cancelled for logistical reasons.
The winter range portion sampled each
month is shown below:

Month

Oct.

Nov.

Winter range
portion sampled

High

Mid

Dec.

Jan.

Feb.

Mar.

Apr.

Low

Low

Mid

High

�-95-

This plan was not strictly adhered to this past winter due to the late
spring and persistent snow cover which affected deer distribution.
Thus,
March trials were held on low elevation winter range and April trials on
mid-elevation winter range.
Two major habitat sub-divisions of the Piceance juniper-pinyon-shrubland
winter range were recognized:
juniper-pinyon-mixed
browse and juniper2
pinyon-sagebrush.
A general study area of 55 to 65 miles2 (142 to 168 km )
for each was delineated and grazing trials were conducted both places during
each grazing period (Fig.l).
Two observers and 8 tame mule deer were used to conduct grazing trials.
A grazing trial is defined as I observer following 1 deer for I to 1Yz hours
and recording the number of bites by species and plant parts taken. All 8
deer were observed each day for 3 days for a total of 24 trials per study
area. A different grazing site was used each day to expose the deer to a
variety of vegetation and terrain features.
Sites were spaced Yz to I mile
(0.8 to 1.6 km) apart. During the trials, the deer were kept overnight,
and during the day when not being grazed, in temporary pens 24 x 24 feet
(7.3 x 7.3m) constructed on each site of 8 x 8 foot (2.4 x 2.4m) game
damage panels. Water was always available and a small quantity of alfalfa
hay and concentrate feed was provided each night.
The deer were allowed 1 day to familiarize themselves with an area before
the start of grazing trials. For grazing observations, the deer were
released in groups of 4 but only 2 were observed.
Four trials were held
in the morning and 4 in the afternoon.
The first group of deer was started
in a random direction from the pen and allowed to roam and forage at will.
Their direction of travel was influenced only if they started following
roads or trails or tried to return to the pen prematurely.
Each succeding
deer group was started in a direction 900 clockwise from the preceding
one. After all deer had been observed, they were walked to the next
site and penned overnight.
Procedures were repeated the following 2 days
after which the deer and pens were moved to the second study area and the
entire process repeated.
Individual deer were alternated between observers and time of day observed.
The sequence of study areas was also alternated between grazing periods.
An extra deer that was raised and trained with the others was included
in the grazing trials. This deer was substituted on several occasions
for other deer with injuries.
After completion of trials on each study area, the major forage species,
on a bite basis, were determined and those comprising 2 percent of more
of the total bites were collected for bite-weight conversions.
A minimum
of 25 simulated "deer bites" per species was handpicked, oven-dried at
1050C for 24 hours, and weighed to the nearest 0.0001 gram. Conversion
factors for species not collected were based on those for similar forages.
Twenty-five to 50-gram (dry weight) samples of forages comprising 2
percent or more of the total bites were collected on each study area for
estimation of crude protein content in vitro dry matter digestibility .

•

�-96-

Samples were comprised of plant portions eaten by the deer.
Samples were
stored in moisture-proof bags in a freezer until they could be oven-dried
at 600C for 24 hours and ground and stored in glass jars.
Forage availability is largely a function of plant density, stage of
development, growing conditions and, most importantly in winter, snow
cover.
Since even a small change in snow depth can affect the relative
availability of species, it was deemed most practical to use a subjective
rating system at the time grazing trials were conducted.
The availability
ratings of abundant, locally abundant, common, and locally common follow
those of Neff (1974) except "uncommon" was substituted for his term "rare".

RESULTS
Mule Deer Forage Selections

on Winter

Range

Results of the 1978-79 winter deer grazing trials are currently being
summarized.
The computer program used to summarize the elk grazing data
from Work Plan 3, Job I is being modified to handle the Piceance data.
A trial run of the October grazing data is being made to check the
revised program but is not yet completed.
Bite/weight conversion factors have been calculated but cannot be applied
until the computer summaries are available.

LITERATURE

CITED

Neff, D. J. 1974. Forage preferences of trained mule deer on the Beaver
Creek watersheds.
Arizona Game and Fish Dept. Spec. Rept. No.4.
61p.

/'J

Prepared

by:

-e

•

/&amp;-t {..(~

.•.•...••

q{

.J) '(

.f;;:Z~~. _.

Richard M. Bartmann
Wildlife Researcher

�GAME

M~.NAGEMENT UNIT

22

-PICEANCE-

"...

9---.l..-J_'

"._~...•••

1IrDi'::'.::1ime.~

••••

\

"-'--""

IF.I&lt;

r

i~

;JJ .

_,I( i

r

.~
.~

~

I
I.

.~~l
I'

.

J

',/(

,-I

u'

J~.

".'\ t;\ I~'"\'
_ I r /'/~__,_, (/) 1"
r ~'" \ " ) .1 \
---

\!

.i-&gt;: -(/

,~

\. \

I

1('\.

V/~

Fi~. 1.
Locations of two general study areas on the Piceance
winter ran~e for conducting mule deer winter forage selection

"---'.;;;0-)\'

juniper-pinyon-shruhland
trials.

I
\0
--.J

I

��July, 1979

-99-

JOB PROGRESS

State of

COLORADO
------------------------------

Project

No.

Work Plan No.
Job Title
Period Covered:

REPORT

W-126-R-2

Deer Investigations

2

Experimental

Job No.
Deer Inventory

5 (1st year) Northeast

- Northeast

Region

Region

June 30, 1978 - July 1. 1979

Personnel:
Allen Anderson, A. William Alldredge, Jay Bucholz, Vera Chen,
Alexis Duxbury, James Enote, Cheryl Fimbel, Randy Fischer, Vicki Jameson,
Gene Schoonveld.

ABSTRACf
Based on counts of pellet groups on 40, permanent, randomly selected,
107.64 ft2 (10 m2) circular plots within each of 25 stratified, pro~ortionallY
allocated and randomly selected square miles (2.59 km2) from 310 mi
(802.9 km2)
of G.M.D. 20 winter range; mean densities with 95 ~ercent confidence limits
were 17.37 &lt; 31.26 &lt; 45.15 deer per mile2 (2.59 km) and .30 &lt; 4.12 &lt; 7.94
elk per mile2 (2.59-km2) from September 11, 1978 to May 30, 1979. Minimum
mean densities with 95 percent confidence limits derived from direct counts
of mule deer and elk on 93, stratified, proportionally allocated, and randomly
selected ~-mile2 (.648 km2) quadrats were 8.03 &lt; 13.52 &lt; 18.93 deer per mile2
(2.59 km2) and .12 &lt; 1.53 &lt; 5,46 elk per mile2 (2.59 kmL), respectively, from
February 21, 1979 t~ April-7, 1979. Buck:doe ratios (+ SE) and fawn:doe
ratios (± SE) sampled on 20, all-day walking routes by-2 observers per route,
November 21, 1978 to December 22, 1978; were .2241 + .0371 (N=874) and
.4860 + .0282 (N=1061), respectively.
From 1976 to-1978 there was a significant
(P &lt; 0~10) increase in both the buck:doe and fawn:doe ratios but no significant
(P &lt; 0.10) change in either ratio from 1977 to 1978. Sample sizes for these
statistics were estimated to be with X percent and at the cited probability
level as follows:
counts of pellet groups-deer, + 20 percent at P &lt; 0.20 and
elk, grossly inadequate; minimum density quadrat ~ounts-deer, approached +
20 percent at P &lt; 0.20, and elk, + 100 percent at P &lt; 0.10; buck:doe rati~s,
approached 20 percent at P &lt; 0.20-and fawn:doe ratios + 10 percent at P &lt; 0.10.

��-101-

EXPERIMENTAL DEER INVENTORY
Northeast Region
Allen E. Anderson
and
David C. Bowden
P. N. OBJECTIVE
To design appropriate sampling and analytical procedures necessary to reliably
estimate deer numbers and buck:doe:fawn ratios on winter range based on a
preliminary sampling of a problem management unit within each of the four
administrative regions of the state.

SEGMENT OBJECTIVES
1.

To estimate densities of deer pellet groups, or where domestic sheep
may occur, helicopter counts of deer on the entire winter range of
one Game Management Unit within each of the four administrative
regions.

2.

To establish permanent pellet group plot locations according to a
sampling scheme capable of efficiently estimating mean deer numbers
+ 15% at a = 0.05.

3.

To sample buck:doe:fawn ratios on 12 or more randomly
walking routes on the winter range of Game Management
selected administrative regions.

selected, all-day,
Units within

This report is concerned only with the preliminary sampling of deer and elk
pellet groups, the minimum bound estimate of deer numbers, and buck:doe and
fawn ratios on the winter range of Game Management Unit (G.M.U.) 20 within
the Big Thompson and North St. Vrain river drainages.
METHODS

AND MATERIALS

Counts of Pellet Groups
Field work during 1976 and 1977, analyses of resultant data, and some
empirical decisions formed the bases for the sampling plan adopted for
installation of permanent pellet group plots (Anderson 1977, Anderson
and Bowden 1978). The sampling plan can be characterized as stratified
multistage as follows.

and

2
The primary sampling units (square mile, 2.59 krn ) were proportionally
allocated among 11 blocks or strata and selected by simple random sampling
within the strata (Fig. 1). The subsampling plan was plotted to scale on
both topographic maps and on aerial photos.
First, the initial reference

�-102-

point was selected, this was generally the most accessible surveyed and
marked section corner.
Each square mile (2.59 km2) was systematically
subsampled by random placement of the first transect (1 mile, 1.6094 km
in/length) within the first 0.1 mile (0.16 km) from the survey marker.
Nine additional transects were systematically located from the first
transect at 0.1 mile (0.16 km) intervals.
Transects traversed the square
mile (7..59 km2) sample unit in that cardinal direction which would place
most transects approximately perpendicular to the contour lines.
The
transects were numbered from 1 to 10 in sequence.
The transects 1 and 5,
2 and 6, etc., which were one-half mile (0.8 km) apart were considered
pairs.
From the 5 pairs of transects 1 pair was randomly selected.
Along
each transect of this pair, there was a random placement of 10 consective
plots spaced 66 ft (20.1 m) and separated by 1,980 ft (603.5 m) from
2
another set of 10 plots also spaced at 66 ft (20.1 m). In effect, each ~mile
(.648 km2) of each square mile (2.59 km2) was sampled by 10 plots. Plots were
107.64 ft2 (10 m2) in area, circular, with their centers marked by numbered,
angle-iron stakes painted glossy blue over silver rust preventative.
A representative sample of the subsampling scheme is shown in Fig. 2. We
used xerox copies of the subsampling plan on the topographic maps and aerial
photos as aids in the installation of 40 plots on each of 26 randomly
selected square miles (2.59 km2). The 1,040 sample plots were established
from May 24 to September 10, 1978 by 2, 2-person crews using 2 chain topographic
steel tapes with trailer, abney levels, surveyor's pins and Silva Type 15+
hand-held compasses.
In general, each sample unit required about 2 days or
8 person days to complete.
Surveyor's staff compasses were used initially but proved to be too slow
and sensitive to deflection by fences, ore deposits, and power lines.
Transects
10 chain-segments, individual plots, and routes to transects and segments were
marked for relocation by nailing 4 inch x 4 inch (10.2 cm x 10.2 cm) galvanized
metal markers and tying surveyor's tape to trees, logs or bushes.
Where
permissible, steel fence posts were used as transect or segment markers.
Deer and elk pellet groups were counted and removed from the sample plots,
September 11 to November 8, 1978 by two, 2-person crews following specific
definitions and procedures (Appendix A). We attempted a total pellet group
individual pellet removal by sifting through the litter layer to mineral soil.
On a few plots, the depth and composition of the litter made this procedure
impractical.
Each sample plot was assigned to one of 28 habitats (Appendix B).
The general criteria for vegetation was the dominant and codominant species
within 30 ft (9.14 m) of the plot stake, All data were punched directly on
data processing cards using IBM PORT-A-PUNCH boards (Patton and Casner 1970).
Deer and elk pellet groups were counted and removed from the permanent plots
April 24 to May 30, 1979. Data were recorded on a prepared form (Appendix
C). After one day of training with emphasis on differentiating
between deer
and elk pellet groups, the counts were conducted over two time periods;
about one-half of the sample was completed on each stratum by May 10
(replication 1) and the remaining by May 30 (replication 2). Sample units

�-103-

and individuals for each 2 person crew were randomly selected.
However,
series of snows mainly at the higher elevations made a shambles of the
stipulated random order during both replications.

a

Minimum Bound Estimate
This estimate is based on the actual number of deer and elk counted by two
observers on ~ mile2 (.648 km2) quadrats and was first used on the Cache
la Poudre river drainage winter range (Bowden and Anderson 1975, Bowden and
Anderson 1976). Since standard errors thereon approximated 9.4 and 20.8
percent of their mean values and the means appeared empirically reasonable,
this procedure was employed on the same sampling universe as the pellet
group counts to provide an additional independent estimate of deer and elk
numbers.
Field work by 2 observers extended from February 21, to April 7,

1979.
It was initially estimated that the 2ime available would permit a total
sample of about 90, ~ mile2 (.648 km ) quadrats from the 1,236 quadrats
present.
Within each of 11 strata, there was a simple random selection of
quadrats following classification and a separate and sequential numbering
of quadrats believed to support relatively "low" or "high" numbers of
wintering deer. The number of quadrats selected was a proportional allocation
within and among strata of "low" and "high" quadrats but with not less than
2 quadrats of each deer density classification per stratum. A total of 93
quadrats were selected for sampling; 66 of which were judged as supporting
"low" and 27 "high" numbers of deer. Quadrat distribution is shown in Fig.
3. These were segregated into 24 groups, with 2-5 proximate quadrats per
group. The groups were numbered and sampled in a random order at the approximate rate of one group per day. Generally, sampling began at daylight
and ended at dusk.
The actual search procedure for deer by 2 observers was influenced by the
topography, prevailing wind, snow conditions, accessibility, and the amount
and spatial distribution of negative cover characteristic of each quadrat.
Prior to search and within the vicinity of the quadrat we used cutants from
a topographic map of the quadrat to decide on the search strategy.
In
general, we traversed each quadrat by walking or snowshoeing as completely
as possible using binoculars from ridgetops, outcrops or other vantage
points.
Search time ranged from about 45 minutes to 3 hours. We recorded
the time of each observation but made no attempt to record age or sex of
either deer or elk on the herd structure from (Appendix D).

Mule Deer Herd Structure
Field and analytical procedures used to estimate buck:doe and fawn:doe
ratios on 20 all-day walking routes with 2 observers have been described
(Anderson 1977, Anderson and Bowden 1978) as well as the procedures to

1

The 1 square mile difference in the 2 procedures
inadvertent omission of an isolated square mile.

was due to the

�-104-

randomly select the locale of each route
(Fig. 4) (Anderson 1977:233).
A representative route is shown in Fig. 5 to reiterate one important
procedure; two methods were used in deer classification and entered
separately on prepared forms (Appendix D). Method I included both
unclassified single deer and groups in which one or more individuals could
not be classified as an antlered buck, doe, or a fawn of the current year
and unspecified sex. Method 2 included only classified single deer and
those groups in which all deer could be so classified.
We used a modification
(Appendix E) of the criteria proposed by Dasmann and Taber (1956) to help
identify sex and age classes.
Just prior to actual sampling, observers spent
one deer making and comparing simultaneous independent classifications of
the same groups of mule deer. Approximately 50 groups of mule deer were
classified in these training excursions.

RESULTS
Counts of Deer and Elk Pellet Groups
Deer and elk pellet groups counted and removed during plot establishment
in 1978 on each of 26 sample units are described statistically for all
habitats combined (Appendix F), by individual habitats (Appendix G) and for
those 10 habitats sampled by 30 or more plots (Table 1). These data provide
a gross index to relative numbers of deer and elk among sample units in the
recent post. The utility of the cumulative mean pellet groups-habitat
relationship is tenuous beyond the observation that up to eightfold differences
between habitats are associated wtih the large variances characteristic of
most sample units.
Up to 53 deer pellet groups were counted and removed from
a single plot and 177 groups from a 10 plot segment.
The basic statistics and pellet groups derived estimate of deer and elk
density and sample size requirements are given in Table 2 and 3, respectively.
Note tbat about 132 mile2 (341.9 km2) of G.M.U. 20 winter range was excluded
from this sample, hence these estimates are not valid for the entire winter
range.
The mean density with 95% confidence limits of 17.37 &lt; 31.26 &lt; 45.15
deer per square mile (2.59 km2) applies then, to about 70 per~ent of the
winter range.
Due to a couple of 10-plot segments with very high counts of
deer pellet groups, variances were very large within strata 5, 6, and 9
(Table 2); hence the wide confidence interval and large estimated sample
sizes.
Thus, the current sample size suggests that the above mean estimate
is only within about 20 percent of the true mean at P = 20. The mean
density with 95% confidence limits of 0.30 &lt; ~.12 &lt; 7.94 elk per square
mile (~.59 km2) (Table 3) is even more vari~ble.
Estimated sample size
requirements for estimating elk densities are essentially unattainable at
any level of preC1Slon.
It may now be possible with a better knowledge of
relative elk distribution and density to devise a sampling scheme which
would yield useful levels of variance.

�-105-

Table 1. Summary of pellet group counts on G.M.U. 20 winter range occurring
within 28 habitats on 26 randomly-selected
square miles.
Only those habitats
sampled by 30 or more plots are included.

Winter Range Habitat
Open ponderosa
Ponderosa

(G.M.U. 20)

pine

pine-Douglas

Douglas

fir

Douglas

fir-ponderosa

fir

pine

1/

Elk-V
Mean
SD

No.
Plots

Deer-Mean
SD

246

4.2

6.00

.28

.73

167

2.5

3.49

.32

.68

38

2.1

4.50

.10

.31

140

2.2

3.40

.18

.47

Open lodgepole

pine

39

1.3

2.60

.70

1. 17

Dense lodgepole

pine

42

0.5

0.97

.61

1.03

fir

47

2.3

4.30

.66

1.24

sage

115

0.9

2.00

.11

.60

mahogany-bitterbrush

89

3.6

6.50

0

0

54

0.2

0.70

0

0

Lodgepole

pine-Douglas

Dry meadow with fringed
Mountain

Rock outcrops

1/

.

.

- Includes both "old" and "new" pellet groups (Appendlx A) counted
establishment, September 11 to November 8, 1978.

on plot

�-106-

2
Table 2. Estimated densities of deer on 310 square miles (802.9 km ) of
G.M.U. 20 winter range based on counts of deer pellet groups on 40 permanent
randomly located 107.64 ft2 (10 m2) circular plots within each of 26 stratified,
proportionally
allocated and randomly selected square miles (2.59 km2). !/

Stratum

No. SamEle Units
Total
Selected

Mean Pellet Groups Per
10 Plots Per Day

Variance
00-5)

Coefficient
Variation (%)

1

26

2

.0077305

6.090106

101

2

25

2

.0119177

4.923084

59

3

34

3

.0108380

3.317595

53

4

28

2

.0029583

0.280987

57

5

29

2

.0191740

73.528455

141

6

30

3

.0228510

27.260672

72

7

30

3

.0127344

15.511025

98

8

28

2

.0226472

11.572833

48

9

32

3

.0266496

26.812108

61

10

22

2

.0195380

13.883112

60

11

26

2

.0143766

0.447872

15

310

26

.015691

Deer Per Square Mile

(2.59

Mean

31. 26

SE

5.00

CV (%)

16.0

95% Conf. Limit

km2)11

17.37 - 45.15

------------------------------------------------------------------------------

�-107-

2
Table 2. Estimated densities of deer on 310 square miles (802.9 km ) of
G.M.U. 20 winter range bas d on counts of deer pellet groups on 40 permanent
randoNly located 107.64 ft (10 m2) circular plots within each of 26 stratified,
2
proportionally allocated and randomly selected square miles (2.59 km ). II
(Continued).

1

Estimates

2
2
of Sample Size (No. Mile - 2.59 km ) R'equlrements- 31
d

1
Percent

.95

.90

.80

5

259

237

200

10

163

132

93

15

101

76

50

20

66

48

30

25

46

32

20

liThe pellet

group deposition period ranged from September 11, 1978 to May
30, 1979, with intervals between counts of 104, 10-plot segments of 168 to
282 days. Hence, the necessity for the mean pellet groups per 10 plots
per day statistic.

llBased on a mean defecation rate of 13.0 groups per deer per day as generalized
from information in Neff (1968).
l/With

15 degrees of freedom and present allocation

proportions

6

�-108-

2

Table 3. Estimated densities of elk on 310 square miles (802.9 km ) of
G.M.U. 20 winter range based on counts of elk pellet groups on 40 permanent,
randomly located 107.64 ft2 (10 m2) circular plots within each of stratified,
proportionally allocated and randomly selected square miles (2.59 km2).

Stratum

Mean Pellet Groups Per
10 Plots Per Day

No. Sampled
Units
Total
Selected

Variance

(10

1

26

2

5.0955

5.192824

2

25

2

2.0905

0.15824

3

34

3

2.7105

0.00530

4

28

2

0.6155

0.07576

5

29

2

1.3733

0.56581

6

30

3

1.8583

0.29329

7

30

3

5.5685

4.05090

8

28

2

0

0

9

32

3

.6250

.07812

10

22

2

0

0

11

26

2

.0020681

310

26
Elk Per Square Mile

Estimates
Percent

2
(2.59 km )

Mean

4.12

SE

1.20

CV (%)

29.1

95% Conf. Limit

-5)

.30 - 7.94
2

of Sample Size (No. Mile
1
a
95
90

2
- 2.59 km ) Reguirements
80

5

310

310

301

10

299

271

223

15

259

216

155

20

218

168

109

25

181

131

79

l/See Table 2 for explanation
2/Based on a mean defecation rate of 12.0 groups
per elk per day as generalized from information in (Neff 1968).

�-109-

Minimum

Bound Estimate

Estimated minimum numbers of mule deer and elk are detailed in Tables 4 and
5 for mule deer and elk, respectively.
In the case of mule deer,
stratification of quadrats to "low" and "high" densities was successful;
mean densities differed significantly (P &lt; 0.02) and there was a 37.8%
reduction in variance due to stratification.
The overall minimum mean
(+ SE) density was 13.48 + 2.24 mule deer per mile2 (2.59 km2) within
the sampling universe of 309 mile2 (247,295 km2) comprising about 70
percent of G.M.U. 20 deer winter range.
For elk, the stratification
designed for mule deer did not reduce the variance even though the high
density mean exceeded the low density mean by a factor of about 7. The
2
overall minimum mean (± SE) density was 1.53 + 0.93 elk per mile2 (2.59 km ).
Estimated sample sizes to be within 20 percent of the true mean at P = .20
were 116 quadrats for mule deer and 676 quadrats for elk. Thus, the 93
quadrats sampled approached adequacy only for mule deer (Table 6).

Herd Structure
The data are summarized by route within each of 15 strata for 3 observers
and 2 methods of classification in Tables 7 and 8. Buck:doe and fawn:doe
ratios are compared by classification method, observer, and years in Table
9. Excluding years, most sample ratios did not differ significantly (P &lt;
0.10). Exceptions include: fawn:doe of observer 1, method 1 vs. method 2;
buck:doe of method 1, observer 1 vs. observer 3; buck:doe or methods 1 and
2 combined, observer 1 vs. observer 3; and fawn:doe of methods 1 and 2
combined, observer 2 vs. observer 3, The significant difference between
the fawn:doe ratio of methods 1 vs. method 2 is inexplicable and has not
previously occurred in several years of herd structure sampling.
Observer
3 participated in only 6 routes and this might be a factor in the
significant differences in the ratios obtained by this observer and observers
1 and 2.
The following ratios obtained in 1978 were significantly (P &lt; 0.10) larger
than in 1976: buck:doe of observer 1, buck:doe, observers and methods of
classification combined, and fawn:doe, observers and methods of classification
combined.
From 1977 to 1978, the fawn:doe ratio of observer 1, methods of
classification combined, decreased significantly (P &lt; 0.10).
Since mean group size of wintering mule deer may conceivably affect
classification, the 1977 and 1978 mean group sizes are listed by stratum
in Table 10 and tested for significant (P &lt; 0.05) differences (Table 11).
Unlike the 1977 sample, mean group size in 1978 did not differ significantly
(P &lt; 0.05) between strata.
There was, however, a significant (P &lt; 0.01)
decrease from 1977 to 1978 in mean group size observed within strata 10 and
11 (Tables 10 and 11). The overall mean (+ SE) group size observed in
1978 was 3.67 + .25 mule deer.
Estimates of sample size requirements (number of routes) indicate that in
1978, the 20 routes yielded fawn:doe ratios that were + 15 percent of the
true ratio at (P &lt; 0.05) but the buck:doe ratio would ~equire III routes

�-110-

2
Table 4. Estimated minimum densities of mule deer on 309 square miles (802.3 km )
of G.M.U. 20 winter range based on ground counts of deer by 2 observers of 93 2
subjectively stratified proportionally allocated, and randomly selected, ~-mile
(.648 km2) quadrats, February 21, to April 7, 1979.
Relative
Density
Low

High

Deer per2~-Mi1e2
(.648 km ) Quadrat

No. Quadrats
Total
Selected

Mean

Variance

1

94

6

.5000

1.5000

Mean

SE

2

92

7

.28571

.5714

1.92218

.51701

3

115

8

2.6250

49.4107

4

101

7

2.2857

12.9048

5

93

6

3.1667

18.9667

6

100

7

.7143

2.2381

7

81

6

3.1667

52.9667

8

76

5

5.2000

55.7000

9

68

5

.4000

.8000

10

50

3

3.3333

9.3333

11

92

6

.3333

.6667

962

66

1

10

2

2.0000

2

4

2

3

21

4

Stratum

D.F.
24

90% Conf. Int.
1.92218

1.0376 - 2.8068

95% Can£. Int.
1.92218

.8851 - 2.9893

8.0000

Mean

SE

1.0000

2.0000

8.50488

1.75084

2

.5000

.5000

11

2

2.5000

4.5000

5

23

2

1.0000

2.0000

6

20

2

5.0000

18.0000

7

39

3

29.6667

274.3333

8

36

3

5.3333

25.3333

9

60

4

5.2500

64.9167

10

38

3

9.6667

72.3333

11

12

2

9.5000

12.5000

274

27

D. F.
5

90% Can£. Int.
8.50488

4.977 - 12.033

95% Conf. Int.
8.50488

4.003 - 13.006

Low and High Deer Density Quadrats Combined
Deer Per Quadrat
Mean
SE
3.38067 .55908

D.F.
6

Total Deer
Mean
4178.51

SE
691.03

Deer Per Mile

2

2
(2.59 km )

Mean
13.52

SE
2.24

�-111-

2

Table 4. Estimated mlnlmum densities of mule deer on 309 square miles (802.3 km )
of G.M.D. 20 winter range based on ground counts of deer by 2 observers of 93 2
subjectively stratified proportionally allocated, and randomly selected, ~-mile
(.648 km2) quadrats, February 21, to April 7, 1979. (Continued).

Low and High Deer Density Quadrats

Combined

2
Total Deer

Deer Per Quadrat
Mean

SE

D.F.

Mean

2.2945 - 4.4670

4178.51

95% Conf. Int.
3.38067

2.0126 - 4.7487

Mean

90% Conf. Int.

90% Conf. Int
3.38067

SE

Deer Per Mile

2836 - 5521

2488 - 5869

SE

90% Conf. Int.
l3.52

95% Conf. Int.
4178.51

2
(2.59 km )

9.14 - 17.81
95% Conf. Int.

l3.52

8.03 - 18.93

�-112-

2
Table 5. Estimated m1n1mum densities of elk on 309 square miles (802.3 km )
of G.M.U. 20 winter range based on ground counts of deer by 2 observers on 93
subjectively stratified, proportionally allocated, and randomly selected,
~-mile2 (.648 km2) quadrats, February 21, to April 7, 1979.
No. Quadrats
Total
Selected

Nean

Variance

6

0

0

92

7

.28571

.23810

3

115

8

0

0

4

101

7

.42857

1.28571

5

93

6

.50000

1.50000

6

100

7

.14286

.14286

7

81

6

0

0

8

76

5

0

0

9

68

5

0

0

10

50

3

.33333

.33333

II

92

6

0

0

962

66

1

10

2

0

2

4

2

0

3

21

2

1.0000

2.0000

4

II

2

.5000

.50000

5

23

2

0

0

6

20

2

0

0

7

39

3

8

36

3

0

0

9

60

4

0

0

10

38

3

11

12

2

274

27

Density

Stratum

Low

1

94

2

High

Elk Per Quadrat

Mean

SE

D.F .

.15283

.06960

14

90% Conf. lnt .
.15283

.0303 - .2754
90% Conf. Int.

.15283

.0104 - .30212

0

Mean

SE

0

1.18673 1.00663

D.F.
2

90% Conf. Int.
1.18673

161. 3333

7.3333

.0949 - 4.1261

95% Conf. Int.
1.18673

.0949 - 5.5183

.3333

.3333
0

Low and High Elk Densitl

Elk Per ~-mile2
(.648 km2) Quadrat

Quadrats

0

Combined

Total Elk

2
Elk Per Mile 2 (2.59 km )

A

Hean

SE

D.F.

.38203 .7.2963
2
90% Conf. Int.
.38203 .0291 - 1.0525
95% Conf. Int.
•38203 .0291 - 1.3701

Mean
472.19

SE

283.83
90% Conf. Int.
472.19
36 - 1301
95% Conf. Int.
472.19
36 - 1694

Mean
1.53
1.53
1.53

SE
.92
90% Conf. Int .
.12 - 4.20
95% Conf. Int .
.12 - 5.46

�-113-

Table 6. Number of randomly selected ~-mile2(.648 km2) quadrats to be within X
percent of the mean at 3 confidence levels for mule deer and elk on G.M.U. 20
deer winter range.
Estimates based on ground counts by 2 observers from February
21 to April 7, 1979.
Percent

Species

II

Mule Deer-

Elk~J

of Mean

Confidence

Level

5

95
944

90
864

80
727

10

640

512

354

20

280

195

116

10

1,136

1,099

1,024

20

914

825

676

50

386

301

200

100

126

92

57

!/Based on optimum allocation (Snedecor and Cochran 1967:525):
New strata
were created by collapsing high and low density strata in strata 1, 2, 3, 4,
5, 6, 11 (Fig. 3) and using the 15 new strata in the allocation.

�Table 7. Summary of mule deer recorded on 20 all-day, walking routes by observer and method of classification,
November 21, 1978 to December 22, 1978, on G.M.U. 20 winter range.

Observer

Route

Stratum

1

1
2
3
4
5
61./
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20

2
3
4
5
5
6
6
7
7
8
8
9
9
10
10
11
11
13
13
14
15
L:

0
0
0
0
0
0
0
0
0
0
0
0
0

F:D
SE
B:D
SE
2

2
3
4
6

3
4
5
6

Method I 1../
Buck
Doe
Fawn
0
0
0
1
0
3
0
0
2
0
0
0
3
7
6
0
8
16
2
0

0
1
0
5
0
0
0
1
0
7
48
= .3333
= .0827

0
0
0
1
0
1
0
0
0
0
0
0
1
2
2
0
0
8
1
0
0
16

UncI.

Total

Buck

0
0
0
2
3
8
0
0
2
0
1
0
2
3
1
0
14
3
3
2
1
45

0
0
0
4
3
12
0
0
4
0
1
0
6
12
10
0
27
27
6
2
2
116

0
1
0
3
0
1
0
5
12
4
4
0
8
4
1
12
6
5
2
7
5
80
F:D
SE

2
0
1
9
0
3
0
2
21
4
10
17
71
21
26
32
20
57
15
27
7
345
= .5246
= .0402

B:D
SE

= .2319
= .0518

0
2
1
5

0
2
3
10

= .1458
=

.0981
0
0
0
0

0
0
0

0

0
0
0
0

2/
Method 2 Fawn
Doe

2
0
0
0

2
0
0
0

1
0
2
3
0
0
0
2
8
2
5
9
46
8
11
22
7

25
9
17
4
181

0
0
1
6

---------------------------------------------------------------------------------------------------------------

Total
3
1
3
15
0
4
0
9
41
10
19
26
125
33
38
66
33
87
26
51
16
606

0

4
5

21

I

•....•
•....•
.p..
I

�Table 7. Summary of mule deer recorded on 20 all-day, walking routes by observer
November 21,1978 to December 22, 1978, on G.M_U. 20 winter range. (Continued).

and method of classification,

21

11

Observer

Route

Stratum

Buck

2

7
8

7
7
8
8

0
0
0
1
0
0
0
0
1
0
0
2

9

10
11
13
14
15
16
18
19

9

10
10
11
11
13
14
I:
F:D
SE
B:D
SE

3

1
5
6
12
17
20

2
5
6
9
l3
15
I:
F:D
SE
B:D
SE

llIncludes

groups wherein

Method 1 Fawn
Doe
0
0
2
0
3
0
1
0
3
1
0
9

Total

Buck

0
0
6
3
6
5
3
2
8
1
1
37

0
0
8
4
9
5
5
2
12
4
1
52

0
3
4
7
1
9
3
0
4
2
8
49

1
22
8
22
19
30
16
0
20
21
20
194

= .4444
= .2953
= .2222
= .1501
0
0
0
1
0
0
1

0
0
0
3
0
0
3

0
0
0
0
0
0
0

0
0
0
3
0
0
3

0
0
0
7
0
0
7

0
0
0
3
l3
6
22

= 0

= 0
= .3333
= 0

9

13
22
4
0
13
9
8
103

F:D
SE

= .5309
= .0522

B:D
SE

= .2526
= .0411

0
0
2
17
76
18
113
F:D =
SE =
B:D
SE

one or more deer or solitary

on route 6 because

1
11
6

Total
2
36
18
38
33
61
23
0
37
32
36
346

,_.I
,_.
\Jl

I

]j Includes groups wherein all deer were classified

llTwo days required

0
0
1
0
0
0
1
0
0
2
0
4

UncI.

Method 2 Doe
Fawn

deer could not be classified.

and all solitary

of snowstorm.

deer were classified.

0
0
0
4
26
l3
43
.3805
.0706

= .1947

= .0302

0
0
2
24
115
37
178

�-116-

Table 8. Summary of mule deer recorded on 20 routes by 3 observers on G.M.U.
20 winter range, November 21, to December 22, 1978. Deer recorded by Methods
land 2 are combined.
Observer
1
F:D = .4987
SE = .0401
B:D = .2177
SE = .0521

Route

Stratum

Buck

Doe

Fawn

Total

1
2
3
4
5
6
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20

2
3
4
5
5
6
6
7
7
8
8
9
9
10
10
11
11
13
13
14
15

0
0
3
0
1
0
5
12
4
4
0
8
4
1
12
11
5
2
7
6
86

2
0
1
10
0
6
0
2
23
4
10
17
76
28
32
32
28
73
17
27
7
395

1
0
2
4
0
1
0
2
8
2
5
9
47
10
13
22
7
33
10
17
4
197

3
1
3
17
0
8
0
9
43
10
19
26
131
42
46
66
46
111
29
51
17
678

0
2
1
5
0
3
4
8
1
9
3
0
5
2
8
51

0
2
3
10
1
22
9
22
22
30
17
0
23
22
20
203

0
0
1
6
1
11
7
9
13
22
5
0
13
11
8
107

0
4
5
21
2
36
20
39
36
61
25
0
41
35
36
361

0
0
0
4
13
6
23

0
0
2
20
76
18
116

0
0
0
4
26
13
43

0
0
2
28
115
37
182

L:

2
F:D = .5271
SE = .0450
B:D
SE

.2512
.0427

2
3
4
6
7
8
9
10
11
13
14
15
16
18
19

3
4
5
6
7
7
8
8
9
10
10
11
11
13
14
L:

3
F:D = .3707
SE = .0713
B:D = .1983
SE = .0304

1
5
6
12
17
20

2
5
6
9
13
15
L:

1

�Table 9. Comparisons of buck:doe and fawn:doe ratios sampled on Big Thompson-North
drainage (G.M.U. 20) winter range; November 21, 1978 to December 22, 1978, November
29, 1977 and November 22, 1976 to December 26, 1976.

Ratio

90% C. I.

Method,
Observer

. Comparlson. 1/
R atlo

3

Method

(4)
.3333

B:D

(3)

o

F:D
2

Method

(11)
.2222
(13)
.4444

B:D

F:D
1

F:D
Method

2

F:D
Method

2

F:D
Method

2

F:D
Methods

F:D

1 &amp; 2

+

.1386

.0302

5

.1995

.0778

-.3805

.0706

5

-.2383

-.5228

-.0304

.1383

14

.2132

-.2739

NS

-.0865

.3234

14

.4830

-.6559

NS

-.0919

.0922

20

.0671

-.2509

NS

-.2046

.0976

20

-.0363

-.3730

S

-.0590

.0156

5

-.0275

-.0904

S

.1623

.1025

5

.3688

-.0442

NS

.0450

.0674

14

.1636

-.0737

NS

-.0187

.0773

14

.1174

-.1548

NS

-.0579

.0510

5

.0449

-.1607

NS

- .1504

.0878

5

.0265

-.3273

NS

-.0763

.0168

5

-.0434

-.1103

S

.1537

.0999

5

.3550

-.0476

NS

1 vs 2

(135)
.1947
(156)
.3805

NS

1 vs 2

(243)
.2526
(297)
.5309
1 vs 2

(425)
.2319
(526)
.5246
(135)
.1947
(156)
.3805
(243)
.2526
(297)
.5309

Obs. 2 vs Obs. 3

(243)
.1947
(297)
.3805

B:D

D. F.

Obs. 1 vs Obs. 2

(425)
.2976
(526)
.5122

B:D

. 2/
Conc 1US10rr-

SE Diff.

Ratio Diff.

Obs. 1 vs Obs. 3

(425)
.1357
(526)
.5429

B:D

B:D

Method

(57)
.14
(66)
.32

B:D

St. Vrain river

28, 1977 to December

(135)
.2526
(156)
.5309

Ob s , 1 vs Ob s . 3

(481)
..•1220
(592)
.5243

(139)
.1983
(159)
.3707

,_.I
,_.
-...J
I

�Table 9. Comparisons of buck:doe and fawn:doe ratios sampled on Big Thompson-North
drainage (G.M.U. 20) winter range; November 21, 1978 to December 22, 1978, November
December 29, 1977 and November 22, 1976 to December 26, 1976. (Continued).

Ratio

Method,
Observer

. Comparlson. 1/
Ratlo

Methods

Obs. 1 vs Obs. 2

1 &amp; 2

(481)
.2857
(592)
.4805

B:D

F:D
Methods

(254)
.2512
(310)
.5271

St. Vrain river
28, 1977 to

90 % C.1.
SE Dif.

D.F.

+

.0345

.0674

15

.1526

-.0836

NS

-.0466

.0775

15

.0892

-.1824

NS

-.0529

.0524
.0843

14
14

.0393
-.0079

-.1452
-.3049

NS

-.1564

-.0943

.0506

13

-.0046

-.1840

S

-.1403

.0811

l3

.0033

-.2839

NS

-.1134

.0335

l3

-.0540

-.1727

S

-.1194

.0621

13

-.0094

-.2293

S

-.0766

.0525

19

.0142

-.1674

NS

.1700

.0558

19

.2664

.0736

S

-.0485

.0327

19

.0080

-.1051

NS

.2126

.0578

19

.3126

.1127

NS

Ratio Diff.

Conclusion

Y

Obs. 3 vs Obs. 2

1 &amp; 2

B:D

F:D

(139)
(254)
.1983
.2512
(159) .3707 (310) .5271
Observer
1976 vs 1978
1
Methods

B:D

1 &amp; 2

F:D
Observers
Combined
Methods

(317)
.1321
(389)
.3893

(481)
.2265
(592)
.5296

S

1976 vs 1978

1 &amp; 2

B:D

F:D

(476)
(874)
.1226
.2360
(582)
(1061)
.3726
.4920
Observer 1 1977 vs 1978
Methods

1 &amp; 2
B:D

F:D
Observers
Combined
Methods

(372)
.1411
(544)
.6687

(481)
.2177
(592)
.4987

1977 vs 1978
(669)
(874)

1 &amp; 2
B:D

F:D

.1716
(969)
.6970

.2202
(1061)
.4844

1/ Sample sizes are in parenthesis.

2/ Significantly

(S) or not significantly

(NS) different

at P &lt; 0.10.

I
&gt;-'
&gt;-'
00
I

�Table 10. Statistics of group size of mule deer by strata obtained on 20, all-day walking routes, G.M.U. 20
winter range, November 21, 1978 to December 22, 1978 and November 28, 1977 to December 29, 1977.
1978 95% Conf. Interval
2

Stratum

LX

LX

n

2
3
4
5
6
7
8
9
10
11
13
14
15

3
3
7
27
39
92
98
233
187
177
297
90
55

9
5
17
121
265
536
448
1493
1319
691
2187
632
255

1308

7978

-

X

SE(X)

1
2
3
8
12
23
31
56
45
67
69
22
17

3
1.5
2.33
3.375
3.250
4.0
3.161
4.161
4.156
2.642
4.304
4.091
3.235

.500
.333
.730
1.023
0.576
.385
.412
.523
.225
.440
.756
.532

356

3.67

.251

1977 95% Conf. Interval

Lower

Upper

2.805
2.375
3.337
3.102
2.192
3.424
2.519

5.195
3.947
4.985
5.210
3.092
5.184
5.663

Total
Among Strata
Within Strata

SS
3172.2
158.4
3013.8

!/ Not significant at P = 0.05.

D.F.
355
12
343

-

X

SE(X)

Lower

Upper

3.00
5.37
6.14
3.98
4.33
3.91
3.25

.71
.62
.66
.41
.40
.44
.64

1.52
4.12
4.79
3.15
3.53
3.00

4.48
6.62
7.49
4.81
5.13
4.82

I

•....•
•....•
\0
I

ANOVA to test whether the 1978 mean group size is similar in each stratum.
Source

-

MS

F

13.20
8.79

1.50.!/

�-120-

Table 11.

Mean group size comparisions

within strata, 1977 vs 1978.

1/

Stratum

t-stat-

D. F))

P

3

1.342

2.8

.5 &lt; P &lt;.8

5

- .105

15.6

P &lt;.2

6

0

13.7

P

7

-1. 605

31.0

.8 &lt; P &lt;.9

8

- .200

31.8

P &lt;.2

9

1.635

76.4

.8 &lt; p

10

2.354

59.3

.98 &lt; p &lt;.99

11

2.864

64.4

.99 &lt; p &lt;.999

13

.049

134.5

P &lt;.2

14

- .204

33.7

p &lt;.2

15

.018

23.7

p &lt;.2

1/

- t-test for unequal variances

(Li 1964:143).
D. F.

t

1
2
8

+ (1-8)

n -1
1

Where 8
S 2
2

/n2

n-l
2

2

= 0

�-121-

at the corresponding percent deviation from the mean and probability
levels (Table 12). Adequate samples sizes for fawn:doe ratios but
impractically large sample size for buck:doe ratios have been characteristic
of G.M.U. 20 winter range herd structure since sampling began in 1976.

DISCUSSION
Based on preliminary calculations using 1976 and 1977 deer pellet group
data from 13,180 temporary plots on GMU 20 winter range, the variance
associated with the estimate of the mean numbers of deer ~as unexpectedly
large. Most of this variance is associated with the extremely high counts
of deer pellet groups on one or more 10-plot segments within 3 of the 11
strata (Table 2). The localized distribution of large numbers of deer
pellet groups has also been noted on Cache la Poudre river drainage winter
range (Bowden et ale 1969). While it might be possible to reduce the
associated variance by adding 2 additional sample units to each of the 3
strata, it now appears improbable that any feasible sampling intensity
will result in an estimate of mean deer numbers within + 15 percent at
a 0.05 as stated in objective 2. This does not negate the counts of pellet
groups as a useful technique but emphasizes the need to increase sample size
by about 6 sample units as indicated and to continue the biannual counts
of pellet groups on an experimental basis for several years.
This will
permit a realistic assessment of annual variations; hence, an improved
estimate of sample size adequacy.
There was a slight overlap in the wide 95 percent confidence limits about
the mean deer densities estimated from the pellet group counts and the
minimum bound estimate.
Both mean values appear feasible since 27.6 to 61.0
deer per square mile (2.59 km2) have been extrapolated from pellet group
counts on the Cache la Poudre river drainage winter range, 1962-65 (Medin
1976:82) and 16.4 + 3.42 deer per square mile (2.59 km2) by the minimum
bound estimate on that same winter range during 1975.
The sampling procedure for herd structure appears adequate for the fawn:doe
ratio but inadequate for the buck:doe with any feasible sampling intensity.
The 20 routes have been delineated on topographic maps and aerial photos in
final form; the sampling procedure is now operational thus fulfilling
objective 3.

LITERATURE

CITED

Anderson, A. E. 1977. Experimental deer inventory, northeast region.
Job
Prog. Rep. Pp. 227-250.
In Colo. Div. Wildl. Game Res. Rept., July
Part 2. 125-305p. (Processed).
Anderson, A. E., and D. C. Bowden.
1978. Experimental deer inventory northeast region. Job Progress Report,
Pp. 225-244.
In Colo. Div.
Wildl., Game Res. Rept., July, Part 2. 137-298p.
(Processed).
Bowden, D. C., A. E. Anderson, and D. E. Medin.
1969. Frequency distributions
of mule deer fecal group counts.
J. Wildl. Manage. 33(4):895-905.

�-122-

Table 12. Number of routes required for sampling buck:doe and fawn:doe
ratios based on 20 all-day routes on G.M.U. 20 winter range, November 21,
1978 to December 22, 1978. Methods 1 and 2 and observers are combined.

No.
Buck Doe

Ratio

SE

160

.2241

.0371

714

Conf . level

Percent

1/

-

(l-a)

•95

.90

.80

5

1002

685

404

10

250

171

101

15

III

76

45

20

63

43

25

No.
Fawn Doe

Ratio

SE

347

.4860

.0282

714

Conf • level

1/

(l-a)

.95

.90

.80

5

123

84

50

10

31

21

13

15

14

9

6

20

8

5

3

Percent-

. .
-l/D eVlatl0n

f rom true ratio.

�-123-

Bowden, D. C., and A. E. Anderson.
1975. Evaluation of herd structure
methodology.
Job Progress Report. Pp. 505-554.
In Colo. Div. Wildl.,
Game Res. Rept.,July, Part 2. 253-561p.
(Processed).
Bowden, D. C., and A. E. Anderson.
1976. Evaluation
methodology.
Job Progress Report. Pp. 227-250.
July, Part 2. 125-305p.
(Processed).

of herd structure
In Colo. Div. Wildl.,

Dasmann, R. F., and R. D. Taber.
1956. Determining structure in Columbian
black-tailed deer populations.
J. Wildl. Manage. 29(1)~78-80.
Li, J. C. R. 1964. Statistical
Ann Arbor, Michigan.
658p.

inference.

I.

Edwards Brothers,

Inc.,

Medin, D. E. 1976. Modeling the dynamics of a Colorado mule deer population.
Ph.D. Diss., Colorado State Univ., Fort Collins.
167p.
Neff, D. J. 1968. The pellet group count technique for big game trend,
census, and distribution:
a review.
J. Wildl. Manage. 32(3):597-614.
Patton, D. R., and W. B. Casner.
1970.
summarization of pellet count data.
Note RM-170.
Fort Collins, CO 8p.

Port-a-Punch recording and computer
U.S.D.A. Forest Service Research

Snedecor, G. W., and W. G. Cochran.
1967. Statistical
Iowa State University Press, Ames, Iowa. 593p.

Prepared by:

(LC/(i,Y,

c;: (~t----dt/~6-()-'r&lt;-/

Allen E. Anderson
Wildlife Researcher

methods.

6th ed.,

�-124-

APPENDIX

A

�-125-

APPENDIX

COUNTING

A

DEER PELLET GROUPS

Definitions

and Procedures

Deer Pellet Group - Five or more deer pellets of the same general size,
shape, hardness and color judged to have been continuously voided at
the place where observed.
Pellets redistributed by water of other
agents to within the plot are not recorded.
Pellets strewn across the
plot are counted as a group if about one-half of the total linear distance
or its midpoint falls within the plot as measured with a steel pocket tape.
Groups occurring on the plot periphery are counted if about one-half of
their total area falls within the plot.
"New" Deer Pellet Group - Pellets are typically
deposited during the previous year.

shiny, often soft and

"Old" Deer Pellet Group - Pellets are typically not shiny, generally
hard and sometimes wholly or partially concealed by litter.
Search Procedures - The area of search is defined by the light metal chain,
1.784 meters in length revolving about the metal rod driven firmly into the
ground.
The plot size is 0.001 hectare or 10 square meters or 107.64 square
feet. On steep slopes the chain is held horizontal.
~.Jhennecessary, the
plot boundary is defined by dropping a pebble from the 1.784 meter mark
on the chain.
Each plot is searched twice (clockwise and counterclockwise)
with the two observers changing positions with the change in search direction.

�-126-

APPENDIX

B

�-127-

APPENDIX
HABITATS

1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.

Pipo-O
Pipo-D
Pipo-Psme
Psme
Psme-Pipo
Pico-O
Pico-D
Pico-Psme
Pico-Potr
Potr
D. Mead., + Arfr
D. Mead., - Arfr
Wet Mead.
Ir. Mead.
SAL Mead.
Cemo-Putr
Putr-Cemo
JUNI-Shrub
Riparian
Logged-recent
Rock-outcrop
Rock talus
Dry Gull Bott.
Water
Spr.-fir.
Cult. Fields
Surfaced roads
Unsure road

ON DEER WINTER

B
RANGE

OF G.M.U.

20

Ponderosa pine, open stand
Ponderosa pine, dense stand
Ponderosa pine-Douglas fir
Douglas fir
Douglas fir-ponderosa pine
Lodgepole pine-open stand
Lodgepole pine-dense stand
Lodgepole pine-Douglas fir
Lodgepole pine-aspen
Aspen
Dry meadow, with fringed sage
Dry meadow, without fringed sage
Wet meadow
Irrigated meadow
Willow meadow
Mountain mahogany-bitterbrush
Bitterbrush-mountain
mahogany
Juniper spp-other shrub species

Dry gulley

bottom

Spruce-fir
Cultivated

fields

�-128-

APPENDIX C

�-129-

APPENDIX C
FORM FOR RECORDING PELLET GROUP COUNTS

G.M.U. 20
PELLET GROUP COUNTS
W-126-R
WP2 -

SAMPLE UNIT ___

STRATUM
J.:"If.

OBSERVERS

------

IDlE BEGAN

Plot I

TRANSECT(S)

ELK
New Old

TOTAL

I 'fRAN.
11

2

12

3

13

4

14

5

15

!

I

16

I

S

I

9

I

I

10

..

Total
COmI:lents, Weather,

HR

-

-

DEER
I
Plot -- New Old

1

6

_

-------------------

TIME FINISHED

__QEER
New Old

TR.-\...'-;" •

J5

17

I
19 I

MIN

-

ELK
New Old

I

I,
I

18

i

20

I

Total
etc.

------------------------------

�-130-

APPENDIX D

�-131-

APPENDIX D

Form for recording deer herd structure and counts of
deer and elk on quadrats

----- Time:----to---- Observer ---------N, R
w.
Area:
------ Legal Descr.: ___ 1;..../4_, S __ , T
Date:

B

D

F

Strata

---

Unc.

L

Random No.
TIME

B

D

F

Unc.

~

TIME

�-132-

APPENDIX E

�-133-

APPENDIX E
SOME CRITERIA FOR DISTINGUISHING

ATTRIBUTE

ANTLERED MALES

SEX AND AGE OF MULE DEER

DOES

BODY SIZE

HEAD SHAPE

DEEP MUZZLE
OFTEN APPEARS
VERY THICK

NECK

THICK

PELAGE

BEHAVIOR

DARK FOREHEAD
CONTRASTING
WITH LIGHT
MUZZLE
OFTEN SOLITARY
OR SOMEWHAT
APART FROM
GROUP

LESS DEEP MUZZLE.
DORSAL SURFACE
OF MUZZLE MAY
APPEAR CURVED
WITH VISIBLE
VEINS
OFTEN LONG,
THIN
OFTEN LESS COLOR
CONTRAST THAN
MALE ON HEADMUZZLE AREA
USUALLY TAKES
LEAD IN GROUP
TRAVEL, SOMETIMES AGGRESSIVE
TO FAWN

FAWNS
SMALLER THAN HOST
ADULTS ESPECIALLY
IN LENGTH OF HEAD
AND SIZE OF hTHITE
RUMP PATCH
SHORT MUZZLED,
HIGH CROWNED. LONG
EARS IN PROPORTION
TO HEAD SIZE
RELATIVELY SHORT
AND THICK
EARS AND NECK
MAY APPEAR
"FUZZY" OR
"WOOLY"
USUALLY CLOSE TO
DOE. EARS OFTEN
CLOSE TOGETHER
AND NECK ARCHED
WHEN ALAID1ED AND
RUNNING OR
TROTTING

�-134-

APPENDIX

STATISTICAL
WITHIN

DESCRIPTION

EACH SAMPLE UNIT,

F

OF TOTAL COUNTS OF DEER AND ELK PEL.LET GROUPS
MAY 24 TO SEPTEMBER 10, 1978.

�-135-

N

Mean

S

40
40
40
40

3.9750
.1250
.4500
0.0000

4.7528
.4043
.6385
0.0000

19
19
19
19

5.4737
.2632
.0526
0.0000

5.2108
.4524
.2294
0.0000

8
8
8
8

1.6250
0.0000
.2500
0.0000

2.0659
0.0000
.7071
0.0000

5
5
5
5

4.0000
0.0000
.4000
0.0000

5.8310
0.0000
.8944
0.0000

2

13.0000
3.5000
0.0000·
0.0000

9.8995
3.5355
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

Stratum 10, Sq Mi 13
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Stratum 7, Sq Mi 29
Habitat 1, Open Ponderosa pine
Old deer
New deer
Old elk
New elk
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New lek
Habitat 11, Dry Meadow with Fringed Sage
Old deer
New deer
Old elk
New elk
Habitat 16, Mountain Mahogany-Bitterbrush
Old deer
New deer
Old elk
New elk

2
2
2

Habitat 18, Juniper-Shrub
Old deer
New deer
Old elk
New elk

2
2

2
2

�-136-

N

Mean

S

1
1
1
1

1.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

3
3

3
3

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

22
22
22
22

2.2273
0.0000
0.0000
0.0000

2.9266
0.0000
0.0000
0.0000

14
14
14
14

.2857
0.0000
0.0000
0.0000

.6112
0.0000
0.0000
0.0000

2
2
2
2

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

Stratum 7, Sq Mi 29 (cont.)
Habitat 19, Riparian
Old deer
New Deer
Old elk
New elk
Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk
Stratum 7, Sq Hi 19
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 5, Douglas Fir-Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk

------------------------------------------------------------------------------

�-137-

Stratum

7, Sq Mi 8

Habitat

1, Open Ponderosa

Old
New
Old
New
Habitat
Old
New
Old
New
Habitat
Old
New
Old
New
Habitat
Old
New
Old
New
Habitat
Old
New
Old
New
Habitat
Old
New
Old
New
Habitat
Old
New
Old
New

Pine-Douglas

Fir-Ponderosa

deer
deer
elk
elk

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

4
4
4
4

1. 2500
.2500
0.0000
0.0000

2.5000
.5000
0.0000
0.0000

4
4
4
4

1. 7500
0.0000
0.0000
0.0000

3.5000
0.0000
0.0000
0.0000

7
7
7
7

.5714
0.0000
0.0000
0.0000

1.1339
0.0000
0.0000
0.0000

2
2
2
2

1.0000
.5000
0.0000
0.0000

0.0000
.7071
0.0000
0.0000

3

8.6667

3

.6667

3
3

0.0000
0.0000

3.7859
1.1547
0.0000
0.0000

Pine

deer
deer
elk
elk
8, Lodgepole

1
1
1
1

Pine

deer
deer
elk
elk
7, Dense Ponderosa

1. 8387
.3519
0.0000
0.0000

Pine

deer
deer
elk
elk
6, Open Lodgepole

1. 3333
.1333
0.0000
0.0000

Fir

deer
deer
elk
elk
5, Douglas

15
15
15
15

S

Pine

deer
deer
elk
elk
3, Ponderosa

Mean

Pine

deer
deer
elk
elk
2, Dense Ponderosa

N

Pine-Douglas

Fir

-----------------------------------------------------------------------------~-

�-138-

N

Mean

S

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

3
3
3
3

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

1.0000
0.0000
.5000
0.0000

1.4142
0.0000
.7071
0.0000

2
2
2
2

1.5000
0.0000
0.0000
0.0000

.7071
0.0000
0.0000
0.0000

3
3
3
3

1.6667
0.0000
0.0000
0.0000

2.8868
0.0000
0.0000
0.0000

13
13
13
13

3.0000
.6923
.2308
0.0000

3.5824
1.7022
.4385
0.0000

1
1
1
1

0.0000
0.0000
2.0000
0.0000

0.0000
0.0000
0.0000
0.0000

Stratum 7, S9 Mi 8 (cont.)
Habitat 19, Riparian
Old deer
New deer
Old elk
New elk
Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk
Stratum 6, S9 Mi 28
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 4, Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 5, Douglas Fir-Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 6, Open Lodgepole Pine
Old deer
New deer
Old elk
New elk

--~------------------------------------------------------------------------------

�-139-

-------

Stratum

6, Sq Mi 28 (cont. )

Habitat

7, Dense Lodgepole

Old
New
Old
New
Habitat
Old
New
Old
New
Habitat
Old
New
Old
New
Habitat
Old
New
Old
New
Habitat
Old
New
Old
New

Pine-Douglas

deer
deer
elk
elk

deer
deer
elk
elk

0.0000
0.0000
0.0000
0.0000

11
11
11
11

1.5455
0.0000
.1818
0.0000

2.8413
0.0000
.4045
0.0000

5
5
5
5

3.8000
.2000
0.0000
0.0000

4.3818
.4472
0.0000
0.0000

1
1
1

1.0000
2.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

3
3
3
3

14.0000
3.3333
0.0000
0.0000

11. 7898
3.21Lf6
0.0000
0.0000

9

3.2222
.4444
0.0000
0.0000

2.8186
.8819
0.0000
0.0000

1

II, Dry Meadow with Fringed Sage
deer
deer
elk
elk

Habitat

1, Open Ponderosa

Old
New
Old
New

0.0000
0.0000
0.0000
0.0000

10, Aspen

6, Sq Mi 4

Habitat

1
1
1
1

Pine-Aspen

Stratum

Old
New
Old
New

S

Fir

deer
deer
elk
elk
9, Lodgepole

Mean

Pine

deer
deer
elk
elk
8, Lodgepole

N

Pine

deer
deer
elk
elk
3, Ponderosa
deer
deer
elk
elk

Pine-Douglas

Fir

9
9
9

---------------------------------------------------------------------------------

�-140-

N

Mean

6
6
6
6

.1667
0.0000
0.0000
0.0000

.4082
0.0000
0.0000
0.0000

7
7
7
7

1. 2857
.7143
0.0000
0.0000

1.1127
.9512
0.0000
0.0000

7
7
7
7

4.8571
1. 2857
0.0000
0.0000

3.8048
1. 8898
0.0000
0.0000

8
8
8
8

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

31
31
31
31

4.4516
.1613
0.0000
0.0000

4.6607
.4544
0.0000
0.0000

2
2
2
2

9.0000
.5000
0.0000
0.0000

0.0000
.7071
0.0000
0.0000

S

Statum 6, Sq Mi 4 (cont. )
Habitat
Old
New
Old
New

Fir-Ponderosa

17, Bitterbrush-Mountain

deer
deer
elk
elk
6, Sq Mi 12

Habitat

1, Open Ponderosa

Pine

deer
deer
elk
elk

Habitat
Old
New
Old
New

Mahogany

21, Rock Outcrop

Stratum

Old
New
Old
New

Pine

deer
deer
elk
elk

Habitat
Old
New
Old
New

5, Douglas

deer
deer
elk
elk

Habitat
Old
New
Old
New

Fir

deer
deer
elk
elk

Habitat
Old
New
Old
New

4, Douglas

3, Ponderosa

deer
deer
elk
elk

Pine-Douglas

Fir

------------------------------------------------------------------------------

�-141-

Stratum 6, Sq Mi 12

(cont. )

Habitat

Mahogany-Bitterbrush

Old
New
Old
New

16, Mountain

deer
deer
elk
elk

deer
deer
elk
elk

Habitat
Old
New
Old
New

Mean

S

5
5
5
5

5.6000
.6000
0.0000
0.0000

3.7148
.5477
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

1
1
1
1

1.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

8
8
8
8

2.0000
.1250
.3750
0.0000

2.0702
.3536
.7440
0.0000

11
11
11
11

2.3636
.1818
0.0000
0.0000

3.5573
.4045
0.0000
0.0000

4
4
4
4

.7500
0.0000
0.0000
0.0000

1.5000
0.0000
0.0000
0.0000

18, Juniper-Shrub

Habitat
Old
New
Old
New

N

21, Rock Outcrop

deer
deer
elk
elk

Stratum 5, Sq Mi 21
Habitat
Old
New
Old
New

3, Ponderosa

Pine-Douglas

deer
deer
elk
elk

Habitat
Old
New
Old
New

Pine

deer
deer
elk
elk

Habitat
Old
New
Old
New

1, Open Ponderosa

4, Douglas

deer
deer
elk
elk

Fir

Fir

-----------------------------------------------------------------------------

�-142-

N

Mean

S

5
5
5
5

0.0000
.2000
0.0000
0.0000

0.0000
.4472
0.0000
0.0000

10.0000
.6667
0.0000
0.0000

2.6458
.5774
0.0000
0.0000

5
5
5
5

25.4000
1.6000
0.0000
0.0000

13.8311
1.6733 '
0.0000
0.0000

4
4
4
4

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

8
8

.3750
0.0000
.6250
.1250

.7440
0.0000
.7440
.3536

19
19
19
19

.5263
0.0000
.8421
.0526

1.2188
0.0000
1.2140
.2294

Stratum 5, Sq Mi 21 (cont.)
Habitat 5, Douglas Fir-Ponderosa Pine
Old deer
New deer
Old elk
New elk

Habitat 16, Mountain Mahogany-Bitterbrush
Old deer
New deer
Old elk
New elk

3
3
3
3

Habitat 17, Bitterbrush-Mountain Mahogany
Old deer
New deer
Old elk
New elk
Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk

I

Stratum 4, Sq Mi 28
Habitat 6, Open Lodgepole Pine
Old deer
New deer
Old elk
New elk

8
8

Habitat 7, Dense Lodgepole Pine
Old deer
New deer
Old elk
New elk

�-143-

Stratum

4, Sq Mi 28 (cont. )

Habitat

9,

Old
New
Old
New

Old
New
Old
New

12, Dry Meadow Without

deer
deer
elk
elk

Stratum

5, Sq Mi 13

Habitat

2, Dense Ponderosa

Old
New
Old
New

Old
New
Old
New

Old
New
Old
New

6, Open Lodgepole

2.5000
.3333
.6667
.3333

1
1
1
1

2.0000
0.0000
3.0000
0.0000

0.0000
0.0000
0.0000
0.0000

7, Dense Lodgepole

deer
deer
elk
elk

Fringed

3
3
3
3

0.0000
0.0000
1.3333
0.0000

0.0000
0.0000
1. 5275
0.0000

10
10
10
10

.8000
0.0000
1.4000
.2000

1.3166
0.0000
1. 0750
.4216

1
1
1
1

1.0000
0.0000
1.0000
0.0000

0.0000
0.0000
0.0000
O. 0000,

10
10
10
10

.2000
.1000
.1000
0.0000

.6325
.3162
.3162
0.0000

Sage

Pine

deer
deer
elk
elk

Habitat

1. 6667
.1111
.7778
.1111

Pine

deer
deer
elk
elk

Habitat

9
9
9
9

S

la-Aspen

deer
deer
elk
elk

Habitat

Mean

Pine-Aspen

deer
deer
elk
elk

Habitat
Old
New
Old
New

Lodgepole

N

Pine

-----------------------------------------------------------------------------

�-144-

Stratum

5, Sq Mi 13 (cont.)

Habitat

9, Lodgepoie

Old
New
Old
New

11, Dry Meadow with Fringed

0.0000
0.0000
1. 7078
0.0000

4

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

16
16
16
16

2.5625
.0625
.6250
0.0000

3.1405
.2500
1.1475
0.0000

3
3
3
3

2.0000
0.0000
1.0000
0.0000

2.6458
0.0000
1. 7321
0.0000

5
5

1.4000
.2000
0.0000
0.0000

.8944
.4472
0.0000
0.0000

Sage

4
4
4
4

12, Dry Headow without

Fringed

28, Unsurfaced

4
4

Road

deer
deer
elk
elk

1, Open Ponderosa

Pine

deer
deer
elk
elk

Habitat

3, Ponderosa

Pine-Douglas

deer
deer
elk
elk

Habitat

6, Open Lodgepole

deer
deer
elk
elk

Sage
4

Habitat

Old
New
Old
New

0.0000
0.0000
2.7500
0.0000

9

3, Sq Mi 13

Old
New
Old
New

3.9370
0.0000
.4410
0.0000

9

Stratum

Old
New
Old
New

1. 6667
0.0000
.2222
0.0000

9

deer
deer
elk
elk

Habitat
Old
New
Old
New

9

S

Pirte-Aspen

deer
deer
elk
elk

Habitat
Old
New
Old
New

Mean

deer
deer
elk
elk

Habitat
Old
New
Old
New

N

Fir

Pine

5
5

-----------------------------------------------------------------------------

�-145-

N

Mean

S

4
4
4
4

1.0000
0.0000
1.0000
.2500

.8165
0.0000
1.4142
.5000

6
6
6
6

0.0000
.1667
.6667
.1667

0.0000
.4082
.8165
.4082

2
2
2
2

3.5000
0.0000
0.0000
0.0000

2.1213
0.0000
0.0000
0.0000

2
2
2
2

3.5000
.5000
2.0000
0.0000

.7071
.7071
1.4142
0.0000

2
2

2.5000
0.0000
0.0000
0.0000

3.5355
0.0000
0.0000
0.0000

8.5000
0.0000
.5000
0.0000

.7071
0.0000
.7071
0.0000

Stratum 3, Sq Mi 13 (cont.)
Habitat 7, Dense Lodgepole Pine
Old deer
New deer
Old elk
New elk
Habitat 8, Lodgepole-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 9, Lodgepole Pine-Aspen
Old deer
New deer
Old elk
New elk
Habitat 10, Aspen
Old deer
New deer
Old elk
New elk
Habitat 20, Recent Logged Area
Old deer
New deer
Old elk
New elk

2

2

Stratum 4, Sq Mi 5
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk

2
2
2
2

-----------------------------------------------------------------------------

�-146-

Stratum

4, Sq Mi 5 (cont. )

Habitat

3, Ponderosa

Old
New
Old
New

4, Douglas

Old
New
Old
New

Old
New
Old
New

6, Open Lodgepole

7, Dense Lodgepole

16, Mountain

deer
deer
elk
elk

1. 7143
0.0000
0.0000
0.0000

3.3022
0.0000
0.0000
0.0000

6
6
6
6

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

l3
l3
13
13

.9231
0.0000
0.0000
0.0000

1. 2558
0.0000
0.0000
0.0000

4
4
4
4

2.7500
0.0000
.2500
.5000

5.5000
0.0000
.5000
1.0000

4
4
4
4

.2500
0.0000
.5000
0.0000

.5000
0.0000
.5774
0.0000

1
1
1
1

1.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

Pine

Pine

deer
deer
elk
elk

Habitat

7
7
7
7

Pine

deer
deer
elk
elk

Habitat
Old
New
Old
New

Fir-Ponderosa

deer
deer
elk
elk

Habitat
Old
New
Old
New

5, Douglas

S

Fir

deer
deer
elk
elk

Habitat

Mean

Fir

deer
deer
elk
elk

Habitat
Old
New
Old
New

Pine-Douglas

N

Mahogany-Bitterbrush

-----------------------------------------------------------------------------

�-147-

N

Mean

S

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

8
8
8
8

6.8750
.2500
.1250
0.0000

7.0191
.7071
.3536
0.0000

13
13
13
13

5.0000
.1538
.2308
0.0000

6.6207
.3755
.4385
0.0000

16
16
16
16

4.0000
.0625
.3125
0.0000

5.8878
.2500
.6021
0.0000

3
3
3
3

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

Stratum 4, Sq Mi 5 (cont. )
Habitat
Old
New
Old
New

deer
deer
elk
elk

Habitat
Old
New
Old
New

19, Riparian

21, Rock Outcrop

deer
deer
elk
elk

Stratum 3, Sq Mi 8
Habitat
Old
New
Old
New

4, Douglas Fir

5, Douglas

Fir-Ponderosa

deer
deer
elk
elk

Habitat
Old
New
Old
New

Fir

deer
deer
elk
elk

Habitat
Old
New
Old
New

Pine-Douglas

deer
deer
elk
elk

Habitat
Old
New
Old
New

3, Ponderosa

Pine

21, Dry Gully

deer
deer
elk
elk

-----------------------------------------------------------------------------

�-148-

Stratum

3, Sq Mi 4

Habitat

1, Open Ponderosa

Old
New
Old
New

Old
New
Old
New

6, Open Lodgepole

7, Dense Lodgepole

8, Lodgepole

deer
deer
elk
elk

1.0000
0.0000
1.0000
0.0000

0.0000
0.0000
0.0000
0.0000

7
7
7
7

2.4286
.1429
.4286
0.0000

1. 9881
.3780
1.1339
0.0000

2
2
2
2

.5000
0.0000
.5000
0.0000

.7071
0.0000
.7071
0.0000

3
3
3
3

2.0000
0.0000
1.0000
0.0000

2.6458
0.0000
0.0000
0.0000

4
4
4
4

.5000
0.0000
0.0000
0.0000

.5774
0.0000
0.0000
0.0000

2
2
2
2

0.0000
0.0000
.5000
0.0000

0.0000
0.0000
.7071
0.0000

14
14
14
14

2.2857
.0714
1.1429
0.0000

5.7703
.2673
2.0327
0.0000

Pine

Pine

Pine

deer
deer
elk
elk

Habitat
Old
New
Old
New

Fir-Ponderosa

deer
deer
elk
elk

Habitat
Old
New
Old
New

5, Douglas

1
1
1
1

Fir

deer
deer
elk
elk

Habitat
Old
New
Old
New

4, Douglas

S

Fir

deer
deer
elk
elk

Habitat
Old
New
Old
New

Pine-Douglas

deer
deer
elk
elk

Habitat
Old
New
Old
New

3, Ponderosa

Mean

Pine

deer
deer
elk
elk

Habitat

N

Pine-Douglas

Fir

-----------------------------------------------------------------------------

�-149-

S

N

Mean

3
3
3
3

.3333
0.0000
1.3333
0.0000

.5774
0.0000
.5774
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

6
6
6
6

3.3333
0.0000
.1667
0.0000

4.1312
0.0000
.4082
0.0000

12
12
12
12

1.5000
.0833
.2500
.0833

2.1106
.2887
.4523
.2887

Stratum 3, Sq Mi 4 (cont. )
Habitat
Old
New
Old
New

19, Riparian

21, Rock Outcrop

deer
deer
elk
elk

Habitat
Old
New
Old
New

Sage

deer
deer
elk
elk

Habitat
Old
New
Old
New

11, Dry Meadow with Fringed

deer
deer
elk
elk

Habitat
Old
New
Old
New

Pine-Aspen

Deer
deer
elk
elk

Habitat
Old
New
Old
New

9, Lodgepole

28, Unsurfaced

Road

deer
deer
elk
elk

Stratum 2, Sq Mi 11
Habitat
Old
New
Old
New

Pine

deer
deer
elk
elk

Habitat
Old
New
Old
New

1, Open Ponderosa

3, Ponderosa

deer
deer
elk
elk

Pine-Douglas

Fir

-------------------------------------------------------------------------------

�-150-

N

Mean

l3
13
13
13

1.2308
.0769
0.0000
.0769

1.4806
.2774
0.0000
.2774

7
7
7
7

2.4286
.1429
.4286
0.0000

3.7353
.3780
.5345
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

5.0000
1.0000
.5000
0.0000

7.0711
1.4142
.7071
0.0000

18
18
18
18

1.4444
.0556
.3889
0.0000

2.6172
.2357
.6077
0.0000

11
11
11
11

.8182
0.0000
.2727
0.0000

1.1677
0.0000
.4671
0.0000

S

Stratum 2, Sq Mi 11 (cont.)
Habitat 5, Douglas Fir-Ponderosa
Old deer
New deer
Old elk
New elk
Habitat 8, Lodgepole Pine-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk
Habitat 23, Dry Gully
Old deer
New deer
Old elk
New elk
Stratum 2, Sq Mi 4
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 5, Douglas Fir-Ponderosa Pine
Old deer
New deer
Old elk
New elk

-------------------------------------------------------------------------------

�-151-

N

Mean

S

9
9
9
9

.1111
0.0000
0.0000
0.0000

.3333
0.0000
0.0000
0.0000

11
11
11
11

2.4545
.1818
1.4545
0.0000

2.6968
.6030
1.2933
0.0000

12
12
12
12

1.9167
.2500
.8333
.2500

1.8320
.8660
.8348
.8660

3
3
3
3

1.3333
0.0000
.6667
0.0000

.5774
0.0000
.5774
0.0000

7
7
7
7

1.0000
0.0000
2.1429
0.0000

1.8257
0.0000
1.7728
0.0000

3
3
3

1.0000
0.0000
.3333
0.0000

1.0000
0.0000
.5774
0.0000

Stratum 2, Sq Mi 4 (cant.)
Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk
Stratum 1, Sq Mi 22
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 5, Douglas Fir-Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 6, Open Lodgepole Pine
Old deer
New deer
Old elk
New elk
Habitat 8, Lodgepole Pine-Douglas Fir
Old deer
New deer
Old elk
New elk

3

-----------------------------------------------------------------------------

�-152-

Stratum

1, Sq Mi 22 (cont.)

Habitat

11, Dry Headow with Fringed

Old
New
Old
New

Old
New
Old
New

Old
New
Old
New

S

1
1
1
1

1.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

2.0000
0.0000
.5000
0.0000

0.0000
0.0000
.7071
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

13
13
13
13

1.6154
.1538
.7692
.0769

1.6602
.3755
.8321
.2774

4
4
4
4

.5000
0.0000
0.0000
0.0000

.5774
0.0000
0.0000
0.0000

15
15
15
15

3.0000
.2667
.3333
0.0000

3.3594
.7988
.6172
0.0000

18, Juniper-Shrub

deer
deer
elk
elk

Habitat

Mean

Sage

deer
deer
elk
elk

Habitat

N

19, Riparian

deer
deer
elk
elk

Stratum 1, Sq Mi 18
Habitat
Old
New
Old
New

Pine-Douglas

Fir

deer
deer
elk
elk

Habitat
Old
New
Old
New

3, Ponderosa

4, Douglas

Fir

deer
deer
elk
elk

Habitat

5, Douglas

Old deer
New deer
Old elk
Ne~&gt;Jelk

Fir-Ponderosa

Pine

-----------------------------------------------------------------------------

�-153-

N

Mean

3
3
3
3

2.0000
0.0000

1

S

Stratum 1, Sq Mi. 18 (cont.)
Habitat
Old
New
Old
New

Fir

1. 7321
0.0000

1.3333

.5774

0.0000

0.0000

1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

4
4
4
4

.2500
0.0000
0.0000
0.0000

.5000
0.0000
0.0000
0.0000

37
37
37
37

.7297
.1081
0.0000
0.0000

.9902
.3148
0.0000
0.0000

2
2
2
2

.5000
.5000
0.0000
0.0000

.7071
.7071
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

19, Riparian

deer
deer
elk
elk

Habitat
Old
New
Old
New

Pine-Douglas

deer
deer
elk
elk

Habitat
Old
New
Old
New

8, Lodgepole

1
1

21, Rock Outcrop

deer
deer
elk
elk

Stratum 9, Sq Mi 20
Habitat
Old
New
Old
New

deer
deer
elk
elk

Habitat
Old
New
Old
New

16, Mountain

deer
deer
elk
elk

Habitat
Old
New
Old
New

11, Dry Meadow with Fringed Sage

Mahogany-Bitterbrush

19, Riparian

deer
deer
elk
elk

�-154-

N

Mean

6
6
6
6

6.1667
.1667
0.0000
0.0000

8.0850
.4082
0.0000
0.0000

19
19
19
19

2.6316
.1053
.4737
0.0000

3.7448
.3153
.6067
0.0000

10
10
10
10

3.5000
.1000
0.0000
0.0000

4.6007
.3162
0.0000
0.0000

3
3
3
3

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

6.4118
.4118
0.0000
0.0000

10.5834
.7123
0.0000
0.0000

S

Stratum 8, Sq Mi 28
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 5, Douglas Fir-Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk
Habitat 28, Unsurfaced Road
Old deer
New deer
Old elk
New elk
Stratum 9, Sq Mi 22
Habitat 1, Open ponderosa Pine
Old deer
New deer
Old elk
New elk

17
17
17
17

------------------------------------------------------------------------------

�-155-

N

Mean

S

5
5
5

4.4000
0.0000
0.0000
0.0000

4.8270
0.0000
0.0000
0.0000

8
8
8
8

3.3750
.2500
0.0000
0.0000

2.2638
.7071
0.0000
0.0000

1
1
1
1

10.0000
2.0000
0.0000
0.0000

0.0000
0.0000
0 ..0000
0.0000

6
6
6
6

10.6667
1.1667
0.0000
0.0000

10.8382
1.3292
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

3.5000
0.0000
0.0000
0.0000

.7071
0.0000
0.0000
0.0000

13
13
13
13

1.8462
.1538
0.0000
0.0000

1.8640
.5547
0.0000
0.0000

Stratum 9, S9 Mi 22 (cont.)
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New elk

5

Habitat 5, Douglas Fir-Ponderosa Pine
Old deer
New Deer
Old elk
New elk
Habitat 6, Open Lodgepole Pine
Old deer
New deer
Old elk
New elk
Habitat 16, Mountain Mahogany-Bitterbrush
Old deer
New deer
Old elk
New elk
Habitat 18, Juniper-Shrub
Old deer
New deer
Old elk
New elk
Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk
Stratum 11, Sq Mi 8
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk

---------------------------------------------------------------------------~---

�-156-

N

Mean

S

.1667
0.0000
0.0000
0.0000

.3892
0.0000
0.0000
0.0000

15
15
15
15

.2000
.1333
0.0000
0.0000

.4140
.5164
0.0000
0.0000

26
26
26
26

3.4231
.0385
0.0000
0.0000

4.8922
.1961
0.0000
0.0000

1
1
1
1

1.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

2
2
2
2

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

8.5000
.3000
0.0000
0.0000

6.5870
.4830
0.0000
0.0000

Stratum 11, Sq Mi 8 (cont.)
Habitat 11, Dry Meadow with Fringed Sage
Old deer
New deer
Old elk
New elk

12
12
12
12

Habitat 16, Mountain Mahogany-Bitterbrush
Old deer
New deer
Old elk
New elk
Stratum 10, Sq Mi 22
Habitat 1, Open ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 13, Wet Meadow
Old deer
New deer
Old elk
New elk

Habitat 16, Mountain Mahogany-Bitterbrush
Old deer
New deer
Old elk
New elk

10
10
10
10

�-157-

N

Mean

1
1
1
1

2.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

19
19
19
19

9.4211
.8947
1.1579
0.0000

6.8094
1.3289
1.4245
0.0000

4
4
4
4

1.7500
0.0000
.2500
0.0000

2.3629
0.0000
.5000
0.0000

5
5
5
5

1.6000
0.0000
.6000
0.0000

1.8166
0.0000
1.3416
0.0000

5.0000
.5000
3.0000
0.0000

1.4142
.7071
1.4142
0.0000

10.0000
.6667
.6667
0.0000

3.0000
1.1547
1.1547
0.0000

0.0000

0.0000

0.0000
0.0000
0.0000

0.0000
0.0000
0.0000

S

Stratum 10, Sq Mi 22 (cont.)
Habitat 18, Juniper Shrub
Old deer
New deer
Old elk
New elk
Stratum 8, Sq Mi 11
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 3, Ponderosa Pine-Douglas Fir
Old deer
New deer
Old elk
New elk
Habitat 5, Douglas· Fir-Ponderosa Pine
Old deer
New deer
Old elk
New elk

Habitat 12, Dry Meadow without Fringed Sage
Old deer
New deer
Old-elk
New elk

2
2
2
2

Habitat 17, Bitterbrush-Mountain Mahogany
Old deer
New deer
Old elk
New elk

3
3
3
3

Habitat 21, Rock Outcrop
Old deer
New deer
Old elk
New elk

7
7
7
7

�-158-

N

Mean

S

9
9
9
9

4.1111
.7778
0.0000
0.0000

5.32000
1.2018
0.0000
0.0000

15
15
15
15

3.2000
.2667
0.0000
.0667

2.4842
.5936
0.0000
.2582

6
6
6
6

.3333
0.0000
0.0000
0.0000

.8165
0.0000
0.0000
0.0000

8
8
8
8

.2500
0.0000
0.0000
0.0000

.4629
0.0000
0.0000
0.0000

1
1
1
1

0.0000
1.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

1
1
1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

Stratum 11, Sq Mi 14
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 2, Dense Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 11, Dry Meadow with Fringed Sage
Old deer
New deer
Old elk
New elk
Habitat 16, Mountain Mahogany-Bitterbrush
Old deer
New deer
Old elk
New elk
Habitat 19, Riparian
Old deer
New deer
Old elk
New elk
Habitat 28, Unsurfaced Road
Old deer
New deer
Old elk
New elk

------------------------------------------------------------------------------

�-159-

N

Mean

S

2

2
2
2

1.0000
0.0000
0.0000
0.0000

1.4142
0.0000
0.0000
0.0000

19
19
19
19

1.6842
1.0526
0.0000
0.0000

2.6885
.9113
0.0000
0.0000

1
1

10.0000
7.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

15
15
15
15

3.0000
.2667
0.0000
0.0000

6.5465
.5936
0.0000
0.0000

5
5
5

1.4000
0.0000
0.0000
0.0000

2.6077
0.0000
0.0000
0.0000

3
3
3
3

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

1
1

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

Stratum 9, Sq Mi 7
Habitat 1, Open Ponderosa Pine
Old deer
New deer
Old elk
New elk
Habitat 11, Dry Meadow with Fringed Sage
Old deer
New deer
Old elk
New elk

Habitat 12, Dry Meadow without Fringed Sage
Old deer
New deer
Old elk
New elk

1
1

Habitat 16, Mountain Mahogany-Bitterbrush
Old deer
New deer
Old elk
New elk
Habitat 19, Riparian
Old deer
New deer
Old elk
New elk

5

Habitat 22, Rock Talus
Old deer
New deer
Old elk
New elk
Habitat 24, Water
01d·deer
New deer
Old elk
New elk

1
1

�-160-

N
Stratum 9, Sq Mi 7

Mean

S

0.0000
0.0000
0.0000
0.0000

0.0000
0.0000
0.0000
0.0000

(cont.)

Habitat 21, Rock Outcrop
Old deer
.New deer
Old elk
New elk

1
1
1
1

�-161-

APPENDIX G
STATISTICAL DESCRIPTION OF TOTAL COUNTS OF DEER AND ELK PELLET GROUPS
BY HABITAT WITHIN EACH SAMPLE UNIT, MAY 24 TO SEPTEMBER 10, 1978.

�-162-

1

SO '11

18

DEER
N···
40
:

··-:······MEAN
(2: C250

'-___

.:

••••••
:..:

S
2 • 8 4 1 9 .::.

.~~ .~

i

...

-' __.: __

_~_:_:_~._i:..__

.;

&gt;j

VALUE

0

1

2

3

4

5

7

Q

14;

FREO.

14

11

2

5

4

1

,1

1

1,

8

III
1;

!
--:--:::::
-"":.':-:.-'.-';"
--,:, ..-r.. -,
....::,_:\.:
..
.,_)?::""'~\/=:.- .;-....,..

I.•..
Ei K • &gt;

);

40
.••..&lt;

.•. /

r:~~

...•...•...

s

M EA"J
.5000

N

••..••.~.~.~.~.~

...",:.

.7511

2~;i/~.-·;.&gt;? ~ .•....•••.•..

S T RA TIT: 5~~__
2_:, S::-O"';-:~-:-I:--'-:l'--. 8"';--"---'---'-OLD

DEER

1
'--.

.. --,---.,.-~-----.

NEW Df~R
.----~---.
N
MEA~
:

.

-.

·VALUE.
F REO.

____;_;___;__--S
.5335

.1500

..

&gt;

3

...
.

------------_._-····1
&gt;3···

·0

36

----

·----~l

---_ ..-- _.,--.

'",

40

OLD

-. --.

__ ._._-_.

. 1 ..

ELK

_--------._-_.- ------ -___,---

..

N
40

S
.7157

MEAN
.4750

----'rAT·UE--·. ..6- '-1' -2----- --.-..
---.----.----.-----1j
FREO.

26

9

5

NEI,! ELK
N

MEAN

40

.0250

VALUE

0

1

FREo.

39

1

------::0--

S

.1581

--------------

�-163- .
--.------------.-.-.~--~--=-=----------5TRATUM
1
50 ~t
22

DEER
--.,--.

--- .. N

._-

40

---.-- ------,------::-MEAN

"S

1.8500

2.326~

!----::-_.__._---_.

---.------~~~--.-~-

VALUE
FREO~

0
11

-----

.

1.·.·••

&gt;EtK

(:;'"

1
12

__

2
10

3
1

4
2

5
1

,.__,.___.,- _-...,.--,-,-

.-

....,.....

.

10
1

--. --.-_ ---.-~

.

.i:'l

..,...........•....••.•.•.••...
:....•:.•••.•.•
·.•••·.•·..•••••.•
Ji:.:......
•••.••••

r-'-' ---'-N---' ._.MEAN~-" -·--::--S----···-· '-"-"-'
'_.
-,~
..
~
!i

40

1.200e

1.2649

OLD DEER
N.i
.... 40

'.•... MEA N
1.725

:---- -~-'~:"".'."'.-,-"'.'-,-,--,-"
....

VALUE

0

1

2

FREO.

11

12

10

4

3
1

.:.=_..::_..:_.~~_._'-'--' -_.-,':-:_'

MEa\N
.1250

N

40

..~~:.;_.-'

8;

7

Ii

1

_' -.~~

.5633
-------

.. --

.. _-----::-:0---:--

..-.-

....

.3

FREO.
38'
.1
r----:'__ .-.~- ..-...-.-':-:.-.--.-.:..-:...:-----.--:....-.
OLD

6
1

S

--:VAlUE···.····· •.··.O··

_.

5
1

2

..;_.-.-------:--;.-:,-.-.-...:..::..

ELI&lt;

__ ._-----MEAN__ -----_ _----- _-_- -----..-.

N
40

•...

....

..

S

1.2442

.1.1250
0

1

3

4

FREO.
15 13
8
1
---------_._. __ ---

2

VALUE

2

5
1

.

NE W ELK
N----,;if-A-N - ..-------

40
VALUE

FREO.

.0750

----0

3

39

1

---=-s---.4743

�-164-.STRATUM

2

4

DEE P
·--··--·-N-·---p.1-EAN-·-----

40

1.2250

--"$---:----. -r-r-

-

2.6a41

-'--\TAiu{-'-~ ·o-i-----i--~-3-··--f-,--1-1--'"12
FREO.

25

I)

4

2

1

1

"_7----

1 ,.

...-, -,-_:
:.-.~

_E_L ~._ ~~ __ : __.. .-'C.......:
.._.;_;.
•.• ~
N

MEAN

40

.2750

=v: l UE~-~~-FPfO •.
'~-.

---' ..- - -._.

·-i-·-·------~~~::',~':I

30

\,9

t

1..

.._'-_

S TR AlUM

OLD

'-.-,

.5057

2

4

DEER

f

"ME AN C

-'.~~40

'-. -. -~;:-:

1~1500

-S:--, &lt; .. :]

.3918

\:
•

1

0
25

VALUE

FREO.

6

2
4

_'-- -"-.'-. -.-~--""-'--.
..

3

6

10

2

1

~

r-,

ME AN
.0750

N

40

-VALUE ---- a
FREO.

OLD

S
.3499

1

2

38 .".. 1

1

ELK

_'_--'-'N
40

._-----_._------_Mf.AN
.2750

--V]:CUF. ---- cf' ---. f ----'2.---.-- -----.-30

FI?EO.

NEW

q

1

ELK

..
-N---MFA·~--------

40

0.0000
_ ..

VAL UE

0

FREO.

40

!
:

--, ,

--:

DEER ..

NEW

i

$

o.coo~

_--._--_..-:..

�-165-

STRATUM

11

~Q.'H

2

DEER

.-.-----,-.
-.S...--_ ..

----- -N-..
-·-.
-.'-_M f A ~
..

..

€500
.:······1
.•...

40

.

.--:,'

"

,-

.. ···i

!

-'T- -~~2-'--"-3~-:- ~---~~ &lt;\O~ l'~'~

.---:"VA l~"':':'::~--\
FREO.

2.6942

:.. '.'

17

q

4

3

3

2

1

1:

~.,
-'-.

:.

! .' ELK'
N

"'EA~

40

.2250

S
.4229

--------.---.

VALUE
FREO.

0
31

2

STRATUM'

OLD

DEER

VALU!:

0

1

FREO.

18

8

-------:---.

-..

....
,'.::

DEE R

NEW

N

3
3

---:"'-'.---:-:--~~--:---:
.

&lt;

"'EA~

S

.2667

.0750

40

4
1

=»:L uT--6-' -·-1----FREO~

31

3

.

._---------'- ..- ._--------_.

OLD

__
,'_

.

._.--_ ...• _._.---'----"---

ELK

I

·----·-N -_------...,--_._----------------MEAN .
S~
• 17r:;0

40

.3848

..'ffCUE---·-o---'-i-------·-----·--FREO.

NEW

33

7

ELK

._-_MEA~._ ._._----..-

N

40
VAt UE
FREO.

.0500
0
38

1
2

&lt;:
.2207

---

�-166-

STRATUM

SO '1 I

3

4

DEER
N

40

.

.------S---- ---'--.-----:;-~

MEAN

······1.5500

3.6581

\ ~
I

VALUE
FREO.

0

1
7

22

2

3

4

2

4
2

5

.6

22

1

1

1

_'--_.ELK

r
;

M EA~

S

.7250

1.3585

3

4

;~L-OE

FREO.
STRATUM
OLD

DEER
::::-,::;,MEA N~_?7------S

----N--

4 O· ....10 5000

3 • 6 2.3 3·

_,_ '_. __ R_._·_·_:·····i__~-:·-·.r-:::w~~·~.~_
..__..~.__.
VALUE

0

1

FREO.

22

7

.

. ....

2
5

.. ,.,....
:...
;:
....:..
:.:;......
,..
.; &lt;.
:.: J
:.!

. _~_." ., _:~) ..... .._'"'_", __;_;;';
3
1

5
2

4

2

22
1

NEW DEEP
-----,.-.--------~_ .. ---_. __._._ ~-.--'~--N

!"lEAN

40

00500

VALUE

---

··FREOo

38

OLD

O·

s

.... -.-.-~-

1

..

---:-----.

-_._ - .....
...

0

MEA.N

«5

7250

1 ••3585

_---:::-----::--.

VALUE
FREe.

.. -

2

N

.

..~

.2207

ELK
-,..,--_ .. _ .._.
40

..

0

1

25

q

__ --_._._-_ _---_.
..

..

2

3

7

2

3

1

__

.-

NEW·ELK
---j.J----;{EA·N·-------·---S40

000000

VALUE

0

FREO.

40

000000

--_.

-----

�-167-

DEER

·MEAN

N
40

....

S

4.7250
-----

VALUE

0

FREQ.

12

1
4

1,.3770
.. _-------,. _:,.::

_ _._-_._-_
..

2

3

4

4

2

3
4

4
3

'.5

6'

2

1

"

7

A
3

11
1

?

,

18
1

19
1

20
1

25
1

.22'50
l

j

._.

..

._.

_

VALUE
F REO~

OLD

DEER

VAL UE'
FREO.

1
5

0
12

3

4

3

NEW. DEER
N

MEAN

40

.1250

-------.--..

-----.------

VALUE······O
.1
F RE o.
36 .
3

_._.:_--- -----.--.--

OLD

.•

:.:._.;'

2
.. 1

.:__-"-_ -,_.....__........_._.__./ .... i·. i .'.•.•.•.....
/

.. -~..:.......------- ..-,

- ......_~.'"-.:_.~

ELK

s

.MEAN

N
40

.22150

.4797

",

.

')

1

;

\lALUE
FREO.

NEW
..

0
32

1
7

2

1

ELK

,-_ .._-- ._._-----_.

N
~O

__ ._-------:::---

MEAN
0.0000

VALUE

0

FREO.

40

S
0.0000

----

�-168-

DEER
.-------$-----~.---.

-.---ij-----;.,-E4N
40

200250.

204336
.... ,

_.---.-- ----,-

--T ~-- 2-----3 ---5--- 6--=; .............,1:--:1-----'-

VALUE

0

fREQo

14

6

10

2

5

1

1

1
..............

ELK

/

:. ":1

------------'-'---'-:-:------N

I'tE&amp;N

40

0671)0

--_._-----_. __ ._._-VALUE .: 0
1
2
26
fREOo
6
4
'STRATUM

3
--

SO MI

3

4
···.1

.~

13

OLD DEEP
N .•···.

40

VALUE

0

FREQo

15

1

.,--------.

NEW

--.-.'-.

--

DEER
N

MEAN

40

01000

.------_._--_.

VALUE

FREQo

s
03038

.
__
._---------------_._-_._---_
0
1
..

36

__

4

OLD ELK
---------rl(----·"tiiEa~.
40

----$-------------

06250

100786

.

_ ._-----_._----_:.._----_._-----...
---------..
---_._._-----j'
VALUE

0

1

2

3

4

FREQo

27

6

3

3

1

----------------.---NEW

ELK
N

MEIi~

S

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._~ '.' _""""_;.;;_---'----'-"'"'-.-'-"'-""-'=--'-~-----'_--j

11
4

q

1

12
1

19
1

22
1

27
1

44
1

ELK
______

" __

. _.

-'-

. --'-'-_--"

N

MEh~

S

40

0.0000

0.0000

VALUE·

0

--'--. -. -. ---'-.--"'-'.

....•••.••••
';....•.•••••••.••••••
,•••••.
r ••.·:•••::: ••••
'..•":::.): ..•••.•••.•••••••••.•.•
'•'••i.··. f··.•··.;' ..••..•...•.•.•...•.....•........•
'..'}.•...•••..
,••••.•••.•••.•••...••.•••••••••••.•.•.••••••
'.:~_
••..•
,i

FRE 0.' .40
..

__

,~-~===

.-:::----

'

.'_..,.;;
..•.
"'...

:"'::' :.~:::::""

..•.
,,,-

..

. _" .:::. -.'.::':'.::::.;?'":;:.;:.' - :-_ . "_-

9

STRATUM

OLD

"_ ...

DEER
N
40

._--_. _. ---~------.--'VAL
F RE Q.

NEW

DEER

_.

._ .._.

MEAN

40

.4500

_. '

._. .-"-~...:

0

1

2

3

30

3

···6

1

ELK

------_ .. _._ .._---_.:------_.

--"

MEAN

40

0.0000

NEW

__ __ ._---_
_-----------..,,------,......,...-.....,S
.

..

0.0000

-----_. --::-------------_. __ . --------

a

VALUE
FREO.

40

ELK

-----N----riEAN---------s-·
40

'0.0000

0.0000

-..··'\FATUE·----·O------------ ..--------·--·---·
F PE O.

40

__ __;

.8458

FREO.

N

.c_"-_;:__--'--._ ..;._"-~..;."'_..;.,
....-...:.......o •..;.:;....,..;...o;;c--'---'-._..;.,'--"'-_~._--".;..'-_;__---'-_---'--'-

S

VALUE

OLD
.-

._._.

N

----

�---_._-----_
STRATUM

-184-

_------------------SO ~I
13

10

.._ ..

D~ER

13
14
111

...............
'_.

-'_-~.'

~_;..;.;_

~_~;,__-::-::--__;__;;__;;_,,

_

_;__--,-,__;_ _

__;_~_--..;,"

15

18
1

'...-.;.:.... : ,',&lt;;,:::,,:;;:&lt;;::::/ :,:..;:,:.,-

, "".••..
"':.:,...;.,
...•....
&gt;:•...
,' ,.::':::':
..•••...
I/i {:;
__ ~: ii

STRATUM
OLD

DEEP
.- ~-~i~0T'T·-~·?S;;~:--::--V2~::z;~?TI'03017'0TIIY1537T:?i7T1

,.....---:-N---·-- -11 E..~~-.-.

40

---.- -.-. ---~"...,.. __ ._---::--...._,..----------.--.
. VALUE
0··
1

FREO.

3~

:1

__ _---_. -.-._._- --'
.

.--_ -_._-_._-----::

OLD

--.~

.

,-

..•..

'-' :_._---

Ell&lt;
~-

MEA N

s

40

.4500

• 6 3~ 5

--VALL'E--'-- o-----T---'?-·----------------- ..-..-----FREO.

NEW

25

l~

3

ELK

----s

;-.--·---·N------MT.i t-j- ---'_'
40

0.0000

.------ .... _ •.
VALUE

_- --_
0

FREO.

40

.....

G.oooe

- ----....

_-_ _----- _-- ---_._------::---:::---:--.
...

..

�-185STRATUM

SO '11

10

2~

DEER
-'---'-N ------M{AN----------

40

4.5250

--S----~--------·

'---.

5.8000

.~.:'

".;ELK

.

M-E4N------·-..:.----S-

---=-N:-' --

40

"

','

-----------

...

~~----

:::I;( .' ': .

.

:.:.:- .,_ .. ;. :',:'

':";:'::;;'_:'"
,"

••••.;..c_...•.•.._.. "'-'-'--'------'--

, 0.0000

0.0000

OLD DEER
·N-·--·-~-M-EA~40

4.4250

--VA LtJf-'~-b~ ······l~·-~~~-~--...
- --~~~~"'-~...
-..-'-.~-:"---'-'-=--::c-'--'"-~'__::"__'__'_":'--'-~=-=-=---__::__'_:::::'__~=__~
FREO.

11

9

4

3

3

1

1

1

1

1

1

1

.~---.-----.--.-..--.,....--

NEW

DEER

··------N-..-----,.;4-TAN-----40

.1000

S

.3038

--- V A CUE- -6-:--- 1

FREO.

36

4

.-'-_._---- --.---:.- ._._----_.----._--_._-----_. ---- ._----_._---_... __ -~
.

OLD

ELK

----N------M E A N----·-----·-S·-··------- .-- ...
-----40

O.OCOO

0.0000

···---\,.Al-UE-·----- ·-O------·-··~·- -.- _..__
r_.

···_···

• -

.. -.-.-

... ----

FREO •. 40
NEW

ELK

-------N------ME
40

A-fC'

0.0000

VALUE

0

,F REO.

40

----------S-·----·-··----

0.0000

_.---.

---------_

�.

STRATUM

11

..=.lg..u.:~

SO '11

_

14

DEER

·---N---·----MEA-:tiI---· 2~4250

40.

---_._-_-- VALUE
FREO.

--'--'--$-'---

.

3.7682
...

0
13

•._. ----.~.

1

2

9

5

~'---., ---

3
4

5
1

4
4

--------.-,

1

-"""7""----.. ----.-

..-. :-:-:'-~~

ELK

MEA"!
.0250

N

40

STRATUM

------~-;-:-;_-·7

S

.1581.

11

--..
-_.-.
-

N

M E A~

4C

.• 3000

.-VA CUE T--::'o-. '-~l~---c--z- -_ -. 3
F R E Q.

OLD

33

j.

_J

_":': .: .- ~

1

ELK

--·----N----MFAN-------f----40

0.0000

0.0000
.

----_._--_ .._._-_._._-----_ .._-_._._---_._-------_
... .

VALUE
F!(EO.

NEW

0

40

ELK

MTA-N--------S-------·--·-··-·---

·----N

.0250

40
VAL UE
FREO.

a
39

1
1

.15111

-:

-. - -----~ ,

.

�-187-

STRATUM

11

DEER
------ N
i

'.

------s----- ---.

ME A N-

40·

.8250

:';:&gt;

··1.5002

.

:

&lt;0 ~---2------4--· 5-~~~:

VAL~
FR~O.

26

6

2

4

1

1

ELK

----.

N
40

s

MEAN

o.oeco

0.0000

VAL U{--· 0-------,---------

FREQ.

40
SO

111

I

8

OLD DEER
---.--- N----- --.-~.---.
~.-------.-.-....S
\&gt; :
...

..

•..

1.3395'
.: ..

--'[AI uE-c~o·---f~2--4
FREO.

27

:

..;-:._'

-'_:':::

;

6

3

---5":--;':'

3

1

NEw.:··OFER
40

~EAN
.1000

'-------,.--.,VALUE
0
FREO.
38

2

N

S

.4414

2

-------"--------

OLD ELK
----------_ .. __ -_.
N
~EAN
.

40
,---_._

..

0.0000

..

0.0000

.-- --_ .. __ ._-, ... _._--------_
.

VALUE

FREe.

NEW

__ __ ._--_ _._----S

ELK

----N--..
- ME-A-N40

0.0000

-VALUE' --0-----FREQ.

40

---

.. - ..

0
40

--------s
0.0000

�-188-

BERTHOUD

,__.,
I MILE

+
N

Fig. 1.

Twenty-six proportionally and randomly selected square mile
2
(2.59 km2) sample units among 11 strata sampling about 310 mile
(803 km2) of G.M.U. 20 winter range whose upper limit is defined
by the 8,500 ft (2,591 m) contour line. Deer and elk pellet groups
are counted and removed from 40, 10-m2, permanent sample plots
randomly located in 4, 10-plot segments within each sample unit.

�-189-

Fig. 2.

A representative sample of the subsampling plan of the square mile
(2.59 km2) sample unit.
The 4 horizontal lines are the randomly
located segments, each consisting of 10, 10 m2 circular, permanent
plots spaced at 66ft. (20.1 m) intervals on transects I and 6.

�-190-

BERTHOUD

,___.
I MILE
N

Fig. 3.

Distribution of a proportionally allocated, stratified (by 2
relative density) and randomly selected sample of 93 ~-mile
(.648 km2) quadrats on which mule deer and elk were counted
to provide the "minimum bound estimate" of their densities,
February 21, to April 7, 1979.

�-191-

!

£1

II

._._.-i-._._._.
I
7
-.-.-.~j_.-.
I

j

i
j'l9

!

BERTHOUD

-- +

I MILE

N

Fig. 4.

The randomly selected locales of 20 herd structure routes
are indicated by their assigned numbers (smaller) among 15
strata.
The boundaries of strata 6-15, however, are in error
as drafted.

�-192-

Fig. 5.

2
A representative herd structure route.
The square mile (2.59 km )
is the randomly selected locale within stratum 2. The route was
laid to see the maximum number of deer, the only stipulation being
that the route had to include some portion of the randomly selected
square mile (2.59 km2).

�July,

-193.-

JOB PROGRESS

State of
Project

1979

REPORT

C.=..O.:..,;L::,.O:...:RAD=:....:O:__
_
No.

Work Plan No.

Big Game Investigations

W-126-R-2

Job No.

2

~5~(~S~E~)

_

Job Title __~E~x~p~e~r~im~e~n~t~a~l~D~e~e~r~I~nuv~e~n~t~o~r~y~-~S~o~u~t~h~e~a~s~t~R~e~g~i~o~n~
__~
_
Period

Covered:

July 1, 1978 - June 30, 1979

Personnel:
Brian Gross, Rick Binford, Dave Roche, Bill Kendall, Howard
Geduldig, Eric Shropshire, Charlie Wallmo, Dr, David C, Bowden~. and
Thomas M. Pojar.

ABSTRACT
Permanently marked pellet group plots that had been cleared pI;'eviously were
revisited during this segment and the newly deposited groups were counted
and cleared.
This process was accomplished on 24 randomly selected square
miles of the total 1026 square miles in the study area near Canon City,
Data from this effort were used to estimate the relative efficiency of
several sampling designs under consideration.
Herd structure was estimated
by walking 27 routes throughout the study area during t.he period December
11 to December 17. This season~ 272 deer were observed; 260 of which were
classified; 45 bucks, 156 does, and 59 fawns for a ratio of 29B;100D;38F.
Compared to the 1977 ratios of 18B;100D:66F~ the B:D ratio is not significantly
different but the F:D ratio is significantly
(:P &lt; ,005) lower,
Comparison
of three years' data (1976-1978) indicates there was no different (P &gt; .10)
in the B:D ratios (1976 data, 30B: 100D:43F) and the F:D ratio in 1977
(66F:I00D) was significantly higher (P &lt; .10) than in 1976 (43F:100D) or
1978 (38F:I00D).
The 1976 and 1978 F:D ratios did not 'differ significantly
(P &gt; .10).

��-195-

EXPERIMENTAL DEER INVENTORY
SOUTHEAST REGION

Thomas M. Pojar

P. N. OBJECTIVE
Design appropriate sampling and analytical procedures necessary to reliably
estimate deer numbers and buck:doe:fawn ratios of selected management units
in the southeast region of the state.

SEGMENT OBJECTIVES
1.

Design and execute a preliminary sampling system to estimate
pellet group density on Game Management Units 58 and 581.

2.

Design a sampling scheme based on the means variances. and deer
density strata estimated from objective 1 capable of estimating mean
deer numbers + 20 percent at a = 0.10.

3.

Sample buck:doe:fawn
walking routes.

ratios on 12 or more randomly

METHODS

selected.

deer

all-day

AND MATERIALS

Pellet Group Density
The sampling design within a square mile was described in Pojar (1978).
Eight additional square miles were established (plots marked and cleared)
during this segment bringing the total sample square miles to 32. All of
the 14 blocks (strata) contain two sample square miles with the four blocks
(6. 7. 8, and 10) that exhibited an easily definable
higher density (Table
1) of pellet groups assigned one additional square mile each.
The data
from the 24 square miles that had cleared 'permanent plots were used to
calculate the expected variance. assuming for the time being that the
addition of eight more sample square miles will not change the variance
drastically.
Timing trials were conducted to help determine the most time-efficient
sampling scheme.
These time trials included:
time to search plot. time
to locate plot at a one chain interval and at a two chain interval.
An electronic digital stop watch was used to measure the time it took to
search a sample of the plots and to measure the time it took to locate stakes
at one and two chain intervals.

�-196-

Table 1.

Block

Density of pellet groups by block, 1978 data.
Number
Square Mile

2
No. Mi
Sampled

Mean NO.2
Groups/Mi

S2

1

78

2

15.5

84.5

2

74

2

49.5

264.5

3

72

2

47.5

924.5

4

77

2

23.0

968.0

5

80

1

32.00

3241/

6

78

2

100.0

450.0

7

78

2

169.5

26,220.5

8

70

2

106.5

13,612.5

9

80

2

35.5

12.5

10

64

1

94.0

2,796.Q}:j

11

81

0

12

66

2

44.0

0

13

57

2

28.5

544.5

14

71

2

50.0

8.0

l/Estimated from an average coefficient of variation over blocks of
56.257%.

Since some of the sampling schemes considered would use plot spacing of
two chains rather than one chain it was important to know if the plot
location time differed between one and two chain intervals. Therefore,
in each of 13 blocks the following procedure was conducted. Perpendicular
to the starting point of the first transect encountered, the two-person
crew installed a short transect (using a hand-held compass and a steel
surveyors chain) consisting of eight stakes. The first stake was set at
a one chain distance, the next at a two chain distance and so on until
eight stakes were installed. Fourteen such transects were established.

�-197-

At the end of the field season a two-person crew that had not installed
the stakes for a particular transect recorded their time to locate each
stake using only a hand-held compass and pacing for distance.

Herd Structure Estimates
As in the past, the study used the method described by Bowden and Anderson
(1976) to estimate the herd structure.
Again, as in the past, I modified
their technique by using only one observer per route rather than two.

DESCRIPTION
See Pojar

OF AREA

(1977).

RESULTS AND DISCUSSION
Pellet Group Density
New pellet groups were counted and cleared from permanent plots that had
been previously cleared on 24 sample square mile. Permanent plots were
marked and cleared during this segment on eight additional square miles,
giving a total of 32 square miles with permanent plots (Table 2). The
first opportunity for data analysis with a full compliment of sample
square miles will be next segment.
However, the variance based on the
data from 24-square miles was used in examining various within-square
mile sampling schemes.
This same analysis will be done next segment when
data from all 32 square miles are available to verify the initial findings.
Search time was recorded on each of the first ten plots encountered on
eleven blocks.
The times obtained were from a good representation of the
various vegetative, terrain, and pellet group density conditions encountered
across the study area. The mean search time per plot was 173.63 seconds
with a standard error of 11.21 (n = 106).
The mean time to locate stakes at one chain intervals, was 29.44 seconds
(n = 49) and to locate stakes at two chain intervals took 63.36 seconds
(n = 49) (Table 3). Since the travel time between stakes at two chain
intervals would be twice that of one chain interval the recorded time for
the two chain test was divided by two to make it comparable to the one chain
test. When the adjustment for travel time is made, there is little practical
difference in search time between one and two chain intervals (29.44 vs. 31.68).
This is contrary to the results of a similar test by Anderson and Bowden (1978)
and is probably due to the denser ground cover and rougher terrain found on
their test area.
A linear regression was run on the number of new groups found per square mile
on the cleared plots this segment and number found when first clearing the
plots (i.e. uncleared plots) last segment (Fig. 1). The correlation was
significant (P &lt; .01) with a slope of 3.72 which was highly significantly

�-198-

Table 2. Number of new pellet groups found in 1978 by transect, section and
block.
For the legal description of the sample sections, see Pojar (1978).

Block

Section

Transect
Number

Number of
Groups

1

5

5

2

10

20

2

3

7

6

1

32

6

29

5

24

10

14

1

9

6
1

60
41:./

6

15

5

1

10

0

3

33

8

12

2

14

7

18

5

28

10

17

5

35

10

80

5

45

10

40

2

103

7

24

63

2

52
67

3

36
64

4

7
54

5

67
26l:!

6

6

33
70~_/

Mean Groups
Per Mi2

Variance

15.5

84.5

49.5

264.5

47.5

924.5

23.0

968.0

38.5

84.5

109.0

468.0

----------------------------------------------------------------_------------

�-199-

Table 2. Number of new pellet groups found in 1978 by transect, section and
block. For the legal description of the sample sections, see Pojar (1978) .
(Continued) .

Block

Section

Transect
Number

Number of
Groups

7

21

4

213

9

71

5

24

10

31

1

12

6

36

5

24

10

0

4

84

9

105

5

34

10

38

1

19

6

14

3

16

8

25

5

41

10

53

5

49

10

51

5

32

10

57

4

35

9

22

1

0

6

2

5

30

10

14

1

37

6

7

51

75l:-1
8

18
35
6S'!)

9

48

78

10

18
43'!:/

3c}:/
11

1c}:/
472:/

12

45
46

Mean Groups
Per Mi2

Variance

129.0

18,031.0

95.0

7,203.0

37.0

32.0

94.3

30.3

29.5

1,512.5

44.0

0.0

------------------------------------------------------------------------------

�-200-

Table 2. Number of new pellet groups found in 1978 by transect, section and
block.
For the legal description of the sample sections, see Pojar (1978).
(Continued).

Block

Section

Transect
Number

Number of
Groups

13

10

5

23

10

24

1

7

6

5

5

24

10

28

1

22

6

26

36

14

49

59

Mean Groups
Per Mi2

Variance

29.5

612.5

50.0

8.0

II
Based on only 28 plots,

52 stakes had been removed.

llThese sections first cleared in 1978 so the observer was forced to make
the judgement concerning whether or not the groups were "new" or "old".

Mean number

groups per square mile

62.5937,

SD

56.40, S

2

= 3,180.96.

�•

200 1

(73,284)
180
00

I

0"1

I

•.

1601

I-l

140

r--.
...-l

.~
::E:

N

Cl)

/

/

Y = 17.40 + 3.72x
r = .8998

/

p...
f/J

P.
;:l
0

120

I-l
t.!&gt;
.!J
Cl)

~
~

I

100

N

o
,_.

Cl)

I

p...

~

Cl)

80

z
60
40

20

I

, •

0
0

20

Figure 1.

80
2
New Pellet Groups per Mi 1977

40

60

--.-

100

Relation of the number of new pellet groups counted on uncleared
(1977 data) and cleared (1978 data) plots.

�-202-

different (P &lt; .001) from zero. This is added evidence (Pojar 1977) that
permanent plots are necessary to get an accurate estimate of the number of
new groups.

Table 3. Results of time trials
and two chain intervals.

(in seconds)

to locate stakes at one

Range
Mean

Standard

Deviation

Max

Min

One Chain

29.44

28.50

190.00

11.00

Two Chain

63.36

69.48

418.70

24.40

Relative efficiency of several within-square mile sampling systems were
evaluated in terms of time efficiency and in terms of reducing the overall
variance.
The time to read plots and travel from plot to plot was taken
from the field measurements, while the time to travel from headquarters
in Canon City and to locate transects was estimated based on field experience.

Table 4. Comparison of the efficiency of six sampling schemes in relation
to current sampling scheme.
The constraint that 100 to 120 persons days
per year are available to accomplish the work has been applied in all cases.

Number of
Transects

Plots/
Trans.

Plot
Spacing
(Chains)

PlotS/Mi2

2

80

1

160

1

10

2

1

10

1

Total
Plots

Variance
Reduction (%)

28

4480

Standard

10

150

1500

40%

1

10

153

1530

38%

20

1

20

110

2200

35%

2

10

1

20

93

1860

35%

2

40

2

80

52

4160

34%

2

40

1

80

52

4160

30%

2
Total Mi

�-203-

By sub-sampling from the 160 plots per square mile and calculating the
variance, it was apparent that to more efficiently estimate the pellet group
density of the total area, it would be necessary to put less effort in each
square mile and sample more square miles.
The relative efficiency of six
potential sampling schemes are outlined in Table 5. The greatest reduction
in variance is accomplished by maximizing the number of square miles sampled
even though the total number of plots searched is only about one-third the
number searched under current sampling.

Herd Structure

Estimates

The third season of collecting herd structure data via the all-day walking
route technique was completed (Table 5). Three experienced observers and
myself walked 27 routes during the period December 11 and 17. We observed
272 deer, 260 of which were classified, 45 bucks, 156 does, and 59 fawns.
In comparing the three years data (Figures 2 and 3), the fawn:doe ratio
was significantly (P &lt; .10) higher in 1977 than in 1976 or 1978. There was
no difference (P &gt; .10) between the ratios in 1976 and 1978. There was no
difference (P &gt; .10) in any of the buck:doe ratios.
For comparison of annual
variation the number of routes required to estimate the herd structure ratios
within a specified confidence level and within a specified percentage of the
true ratio are presented in Table 6.
LITERATURE
Anderson,

CITED

A. E., and D. C. Bowden.
1978. Experimental deer inventory,
northeast region.
Job Prog. Rept., pp. 225-244. In Colo.
Div. Wildl. Game Res. Rept., July, Part 2. 137-29~

Bowden, D. C., and A. E. Anderson.
1976. Evaluation of herd structure
methodology.
Job. Prog. Rept., pp. 507-554. In Colo. Div.
Wildl. Game Res. Rept., July, Part 2. 253-561-.Pojar, T. M. 1977. Experimental deer inventory, southeast region.
Job
Prog. Rept., pp. 275-286. In Colo. Div. of Wildl. Game Res.
Rept., July, Part 2. 125-305.
Pojar, T. M. 1978. Experimental deer inventory, southeast region.
Job
Prog. Rept., pp. 265-277. In Colo. Div. Wildl. Game Res. Rept.
July, Part 2. 137-297.

�-204-

Table 5.

The 1978 herd structure data from the Canon City study area.

Block

Section

Date

B

D

F

Unc

Total

Observer

1

5

12-12

63

12-12

o
o

o
o

o
o

o
o

HG

1

o
o

2

52

12-12

4

11

1

1

17

TP

2

67

12-12

3

3

o

1

7

CW

3

36

12-13

2

11

9

o

22

TP

3

64

12-13

o

o

o

o

o

CW

4

17

12-13

1

o

HG

54

12-13

o

o

o
o

1

4

o
o

o

ES

5

6

12-15

2

9

2

2

15

HG

5

67

12-15

4

o

o

1

5

ES

6

6

12-14

2

o

o

o

2

HG

6

33

12-14

o

o

o

o

ES

7

7

12-17

6

23

8

o
o

37

TP

7

21

12-11

6

39

11

2

58

TP &amp; CW

8

18

12-14

o

o

o

o

o

CW

8

35

12-14

2

14

6

1

23

TP

9

48

12-16

o

o

o

o

HG

9

78

12-17

1

5

1

7

ES

10

18

12-16

o

o

o

ES

10

43

12-15

2

2

TP

11

19

12-15

o
o

o
o

o
o
o
o

4

2

o

6

CW

11

47

12-17

2

12

7

2

23

HG

12

45 &amp; 46

12-17

1

1

o

o

2

CW

13

10

12-16

6

8

8

o

22

TP

13

36

12-16

3

14

4

2

23

CW

14

49

12-11

o

o

o

o

o

HG

14

59

12-11

o

o

o

o

o

ES

E

45

156

59

12

272

x = 1.661
SE = 2.019
C.V. =

.8254

ES

5.778 2.185

.444

x

10.074

9.044 3.464

.751

SE

14.188

.6388 .6307 .592

C.V.

.710

�-205-

Table 6. Comparison of annual variation in the number of routes to estimate
herd structure at three given confidence levels and within three specified
percentage levels of the true ratio (x).

1976

1977

1978

Buck:doe
Confidence level

Buck:doe
Confidence level

Buck:doe
Confidence level

X

95 .

90

80

95

90

80

95 -,

90

80

5

5066

3497

2082

1180

813

483

1853

1276

758

10

1267

875

521

295

203

121

463

319

189

20

317

219

130

74

51

30

115

80

47

Fawn: doe
Confidence Level

Fawn:doe
Confidence level

Fawn: doe
Confidence level

95

90

80

95

90

80

95

90

80

5

983

678

404

211

145

86

1168

804

478

10

246

170

101

52

36

22

292

201

119

20

62

44

26

13

9

5

73

50

30

X

�FAWN:DOE

RATIO

80

70

60
Ul
&lt;ll
0

~

0
0
•.....•

50

40

l-l
&lt;ll

p..,
Ul

~
t1j

I
N

30

o
0I

J:&lt;.o

20

10

0
1976

1977
YEAR

Figure 2.

Fawn:doe

ratio with 90 percent

confidence

interval.

1978

�50

rJ)
aJ
0
Q
0
0
.-4

J.&lt;
aJ

p..

40

30

1

-.;

-------1

20

rJ)

§
co

r:r..

10
0
1976

1977
YEAR

Figure 3.

Buck:doe ratio with 90 percent confidence interval.

1978

I
N

0

-...J
I

��July, 1979

-209-

JOB PROGRESS

State of

COLORADO
Deer-Elk

Project No.
Work Plan No.
Job Title

Investigations

Job No.

2

Experimental

Period Covered:
Personnel:

REPORT

Deer Inventory - Piceance

~_5_C
Basin ~ Northwest

_
Region

July 1, 1978 - June 30, 1979

D. Freddy, T. Kaminski,
D. Bowden.

L. Robinson, W. Erickson, R. Bartmann,

ABSTRACT
Density of new pellet groups was estimated on each of 10 square-mile sample
units using equal numbers of permanently staked pellet plots and temporary
plots.
There was no significant difference in numbers of new pellet groups
peT square-mile between types of sample plots (P &lt; .05).
There were
55.70 ± 4.50 (SE) and 57.50 ± 4.00 (SE) new groups. per square-mile on
temporary and permanent plots, respectively.
Time to complete a transect
was 65.8 ± 2.3 (SE) minutes for temporary transects and 79.7 + 2.1 (SE)
minutes for permanent transects.

��-211-

EXPERIMENTAL

DEER INVENTORY-PICEANCE

BASIN-NORTHWEST

REGION

David J. Freddy

P. N. OBJECTIVE
Design appropriate sampling and analytical procedures to reliably
deer numbers in the Piceance Basin, Game Management Unit 22.
SEGMENT

OBJECTIVES

Determine the efficiency of permanent and temporary
pellet group densities in pinyon-juniper woodland.
METHODS
Methods

have been presented

estimate

plots in estimating

AND MATERIALS

in detail

in Freddy

(1978).

Criteria in 1979 for subjectively "aging" new pellet groups on temporary
plots were:
1) individual pellets associated in recognizeable groups;
2) pellets usually dark, intensely, and uniformly colored;
3) sheen present
on nearly the entire surface of most pellets;
4) pellet surface usually
smooth in texture; and 5) cracking of pellet surface uncommon.
Criteria for
old groups were:
1) individual pellets not always in recognizeable groups;
2) color of pellets faded and often "bleached" gray on at least one surface;
3) sheen seldom present but occasionally occurred on bottom surfaces of
pellets;
4) pellet surface usually rough in texture;
5) cracking of pellet
surface common; and 6) pellets often settled into duff.
Criteria for new pellet groups were developed from examining known new
pellet groups collected in October, December, February, and April 1978-79
and placed during these months on "aging" plots within 5 microsights.
Groups
were examined in late April to determine appropriate cr~teria.
Criteria for
old groups were developed from examining pellet groups placed on "aging" plots
within the same 5 microsights during winter of 1977-78.
RESULTS AND DISCUSSION
The permanent plot test was conducted from 26 April through 30 May.
Two
persons working as a team evaluated 2 permanent and 2 temporary transects
daily.
There was no significant difference in numbers of new pellet groups per
square-mile found on temporary and permanent plots (P &lt; .05). There were
55.70 + 4.50 (SE) and 57.50 + 4.00 (SE) new groups per-square-mile on
temporary and permanent plots, respectively.
These data differ from 1978
when temporary plots significantly inflated the number of new pellet groups
per square7mile (Freddy 1978).

~

�-212-

In 1979, time to complete a transect was 65.8 + 2.3 (SE) minutes for
temporary transects and 79.7 ± 2.1 (SE)minute;-for permanent transects.
Only 2 of 500 permanently
restaked in approximately

Table 1.
transects
April-May

staked plots were missing.
Missing
the same location and recleared.

plots were

Sampled pellet group densities on temporary and permanent
on 10, square-mile sample units, Piceance Basin, Colorado,

1979.

Square-mile
Sample
Unit

Transecta

New Pellet
Groups

Transect

4-18

1

15

2

10

3

16

4

9

5

9

6

6

7

9

8

17

9

12

10

11

61

4-19

New Pellet
Groups

53

1

10

2

17

3

3

4

9

5

8

6

14

7

9

8

10

9

16

10

12

46
4-30

b

62

1

17

2

11

3

12

4

17

5

26

6

16

7

15

8

17

9

9

10

19

L:

79

L:

80

----------------------------------------------------------------------------

�-213-

Table 1. Sampled pellet group densities on temporary and permanent transects
on 10, square-mile sample units, Piceance Basin, Colorado, April-May 1979. (Cont'd).
Square-mile
Sample
Unit

Transecta

New Pellet
Groups

Transect

4-31

1

11

2

2

3

21

4

8

5

8

6

12

7

5

8

8

9

11

10

13

56

4-12

1

9

2

6

3

5

4

10

5

5

6

17

7

10

8

16

9

11

10

25
74

1

9

2

11

3

11

4

11

5

11

6

6

7

8

8

5

9

9

10

5

48
4-24

New Pellet
Groups

43

40

4-13

b

38

1

13

2

14

3

15

4

12

-------------------------------------------------------------------------------

�-214-

Table 1. Sampled pellet group densities on temporary and permanent transects
on 10, square-mile sample units, Piceance Basin, Colorado, April-May 1979. (Cont'd).
Square-mile
Sample
Unit

New Pellet
Groups

Transect

5

21

6

10

7

11

8

12

9

19

10

12

Transect

a

b

New Pellet
Groups

4-24
(Cont'd)

60

79

4-25

1

8

2

13

3

13

4

11

5

7

6

9

7

13

8

6

9

17

10

17

58

4-14

56

1

13

2

10

3

15

4

9

5

5

6

7

7

9

8

18

9

9

10

10

51
4-23

54

1

5

2

13

3

8

4

8

5

7

6

8

7

4

8

12

�-215-

Table 1. Sampled pellet group densities on temporary and permanent transects
on la, square-mile sample units, Piceance Basin, Colorado, April-May 1979. (Cont'd)
Square-mile
Sample
Unit

a
Transect

New Pellet
Groups

Transect

4-23
(Cont'd)

9

15

10

Grand Sum

a
b

b

55

557

575

transects

contained

temporary

Even-numbered

transects

contained

permanently

LITERATURE

plots.
staked plots.

CITED

Freddy, D. J. 1978. Experimental deer inventory - Piceance
Div. Wildl. Game Res. Rept. July, Part 2:245-263.

by:

~;;i,

'&gt;....i-4(/tt.(_

David J. Fri(cfdy
Wildlife Researcher

14

39

Odd-numbered

Prepared

New Pellet
Groups

y

Basin.

Colo.

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                  <text>JOB PROGRESS

COLORADO

State of
Project

REPORT

No.

W-126-R-2

Deer-Elk

Investigations

3
1
Work Plan No.
Job No.
------~--~~----~~~~~--~
Systems Modeling Big Game Populations;
Simulations of the Carrying
Capacity of the Rocky Mountain National Park Elk Winter Range
Job Title
Period

Covered:

July

1, 1978 - June 30, 1979

Personnel:
D. L. Baker,
and J. E. Ellis.

N. T. Hobbs,

D. G. Milchunas,

L. Stevens,

D. M. Swift,

ABSTRACT
Botanical and chemical composition of diets of tame elk were examined on winter
and summer ranges in Rocky Mountain National Park, Colorado from November 1976
to September 1978. Heasured parameters of diet quality were then used to predict
overwinter animal condition and to estimate relative carrying capacities of major
winter habitat types •
.For 1976-1977, graminoids dominated winter diets (x = 61%) but declined in
importance from 66% of observed bites in November to 44% in February, as browse
consumption increased.
Browse and grass contributed equally to diets through
the winter of 1977-78, accounting for more than 90% of bites each month.
Diet quality was influenced to some extent by changes in chemical composition
of major forage species.
The first year dietary crude protein declined from
5.8 to 4.9% while IVDDM decreased from 49 to 40%.
The following year dietary
crude protein declined from 5.4 to 4.6%; IVDDM, from 42 to 37%. During 19761977 mean overwinter dietary protein was highest in dry grassland (6.3%) and
lowest in mesic meadow communities (4.6%) while dietary IVDDM ranged from 47%
in aspen to 35% in grasslands.
In 1977-78, protein ranged from a high of 5.6%
in willow to a low of 4.6% in mesic meadow; IVDDM was highest (39%) in aspen and
lowest (35%) in willow.
Despite year to year variation in forage quality, elk
were able to maintain a relatively stable diet quality throughout winter by
shifting the forage class mix of diets.
Simulation modeling of elk energy and nitrogen status predicted more favorable
overwinter animal condition during 1976-77 than during 1977-78.
Simulated weight
losses were relatively small the first winter (10% of body weight) but were
much more substantial the second year (20-30% of body weight).
Habitat types
could not be consistently ranked on the basis of their ability to contribute to

-,

�-232-

maintenance of animal condition.
Carrying
quantity of forage present was not related
feeding in those communities.

capacity of habitats based on the
to nutritional status of elk

Summer elk_diets were dominated by grass and grasslike plants during 1977
and 1978 ex = 63%.and 64%, respectively).
Graminoids maintained this
dominance throughout the sampling period (July~September).
For both 1977 and 1978, forage classes exhibited a declining content of
nutrients from July to September.
Crude protein content of grasses (across
types and years) declined from 14% in July to 10.5% in September while IVDDM
decreased from 60 to 56%.
Shrub species decreased in protein from 14.1 to 8.8%
and IVDDM from 32 to 26%.
Forbs decreased in protein and IVDDM from 18.3 to
10.8% and from 61 to 48%, respectively.
Diet quality like forage quality declined as summer progressed.
For 1977,
crude protein in diets (across animals and communities) decreased from 17 to
10% while IVDDM dropped from 54 to 43%.
The following year, diet protein
declined from 15 to 10% but IVDDM did not decline over the same time interval.

�-233-

SYSTEMS MODELING BIG GAME POPULATIONS:
OF THE ROCKY MOUNTAIN NATIONAL

SIMULATIONS OF THE CARRYING
PARK ELK WINTER RANGE

CAPACITY

N. T. HOBBS and D. L. BAKER

P. N. OBJECTIVE
To develop and test a model simulating carrying capacity of elk winter ranges
using Rocky Mountain National Park (RMNP) as a study population.

SEGMENT OBJECTIVES
1.

To analyze botanical
RMNP.

and chemical

composition

2.

To begin model simulations describing the relationships between
Rocky Mountain National Park elk herd and its food resources.

3.

To collect and analyze botanical
diets in RMNP.

and chemical

of elk winter

composition

diets in

the

of summer elk

METHODS AND MATERIALS
Methods and materials employed in the winter and summer studies have been
previously described (Hobbs and Baker 1977, Baker and Hobbs 1978).

RESULTS AND DISCUSSION
Botanical and Chemical Composition
of Elk Winter Diets
A detailed description relating the bqtanical composition· and nutritional
quality of tame elk diets to forage quality, advancing season and plant
communities of upper montane winter range is presented in a Ph.D. Dissertation
written by N. T. Hobbs (Appendix I).

Model Simulations

of Elk Nutritional

Status

Simulated changes in elk nutritional status and predicted carrying capacities
of upper montane habitat types for winter are presented and discussed in
detail in the above Ph.D. Dissertation (Appendix I).

A
V

�-234-

Botanical

Diet Composition

and Chemical Composition
Summer Elk Diets

of

1977 and 1978

Elk summer diets were similar with respect to forage class for both years
of study (Table 1). For both summers graminoids were chosen most frequently
in all vegetation types sampled.
Grasses mantained this dominance throughout
the summer sampling period (July-September).
The principal species consumed
were sedges (Carex spp.), tufted hairgrass (Deschampsia caespitosa), and rushes
(Juncus spp.).
Shrub species were encountered and consumed primarily in willow
park and krummholz-ecotone
communities.
Important shrubs included leaves, stems
and inflorescences of willow (Salix brachycarpa, Salix planifolia) and blueberry
(Vaccinium spp.).
The alpine tundra community provided the majority of forbs
found in the diet.
White marsh-marigold
(Caltha leptosepala), alpine avens
(Geum rossii) and Parry clover (Trifolium parryi) were of major consequence and
equally chosen.

Forage

Quality

1977 and 1978

Chemical composition of the principal summer forage species was similar for
both Year 1 and Year 2 (Table 2). Forage quality as measured by crude protein
content and in vitro digestible dry matter (IVDDM) showed similar patterns of
change with advancing season.
In general, measures of forage quality declined
in all vegetation classes from July to September.

Diet Quality

1977 and 1978

Diet quality like forage quality declined as summer progressed (Table 3). There
appeared to be no differences in dietary crude protein or IVDDM among vegetation
types.
During Year 1, IVDDM in diets declined from a mean (across animals and
communities) of 54% during July to 43% in September.
The second year, diet IVDDM
did not decline over the same time interval.
Crude protein in diets declined
over summer during Year 1 from 17% to 10% and from 15% to 10% the following year.

Prepared

by:

APtlM at J5?~

--~D~a-n~L~.~B~a~k~e-r~~~~--~-Wildlife

Researcher

�Table 1. ~l"an p erceu t age with .90(;confidence Lntervals of observcd bites of plant species in diets of tame elk on subalpine tundra summe r range in Colorado during July-September 1977 and 1978, sununarized by h"bitnt type. !};_/

July
Tax)-/

Year

x

C.1.

IHllow-Pnrk
Augu-st
X
C.1.

September
X

C.1.

x

July
C.1.

Habitat Type
Krununholz-Ecotone
August
X

C.1.

September
X
C.1..

July
X

C.1.

Alpine-Tundra
August
X
C.l.

September
X

C.1.

GRMf1NOllJS

----.-

Ca1amagt:0stis
c anadens is
~arex s pp ,
Deschampsia
caespitosa

Juncus
balticus
Kobresia
mvosuroides c/
All grasses--

1977
1978
1977
1978
1977
1978
1977
1978
1977
1978
1977
1978

.5

1

4

6

tr

2

3

3

o

51
42

16

2
36
42

2
1

2

1

36
34

46
43

20
23

49
53

2

5
3

4
3

12
15

11

5
7
1

11

o

5

9
5

7

10

3

1
11

'l

tr

15
15
tr

6
12
14

o

o

o

60
60

2
4

13
15

63
64

69
64

2

40

4

4

9

10

2

33

7

8

6
3
8
4

72
68

18
10

62
62

13

o
14
29

27

tr
tr
38
43

18

9

o
o

47
51

15
24

13

6
5
4

3

13

10

2

2
2
7
11
3

30
13
12
28

11
4

tr
67
73

6

o
23
33
4
9

30

11

34

o
o

o

o
17

40
31

4

8
11
7
7

2

2

16

5
2

3
2

3

54
46

10
6

9

12
2

15
27
10
3
2
7
1

o
59
59

49
40

23

22

5

5
7
11

6
7

28

17

o
1

9

10

69
78

3
3

15
I

SHRUBS

N
I..U

Salix
brachycarpa
Salix
plal1ifolia
Vaccin.lum
----spp.
,,'
All shrubs--'

1977
1978
1977
1978
1977
1978
1977
1978

5

6
15

3
7
7

8

12

11
12
31
26

6

12
12
8

11
4
11
9
8

16

13
30
26

2
4

1
3

3
8
11

17
11
6

4
6
5

17
15
6

14

12

0.5

7

14

15
12

7

9

6

o

o

10
18

13
16
28

13
10
35

tr
17
24

17

3
7

3
8

tr

o

o

22

7

tr
tr
24
27

3

o

3

3

9

20

1
3
7

19

12

5

7
8

7
4

13

12

23

1

2
4

o
o

15

12
24

7
2

5
11

4
12
17

4

1

15
12

3

9

6

7
7
7
5

4
9

2
2

14

28
21

33

4
7

3
6

7

9

16

7

2

tr

4
3
6

o

o
o

tr
5

7

6

19

10

5

5
6
2

11

FORBS
Caltha
1eptosepala
Ceum rossii
----Polygonum
histortoides
Trifolium
nanum

TrifOlium
pauyi
A11 SIH:ubs·~)
Total Bites-'!-/

!};_/

1977
1978
1977
1978
1977
1978
1977
1978
1977
1978

2
10
0
0
1
0
0
0
0
3

1977
1978

8
15

1977 7245
1978 3813

2

25

2
7

4

12

tr

o
1

tr
1

o

3

o

tr
tr

tr
tr
tr

o

o

tr

tr

o

4

7
11

4544
5630

7
6
1

o

o
o
o

5

15

10

9

6262
6706

9
8

tr

tr
4
5
2

tr

o
5

22

o

2
3
1

1
1

1

1

4

tr

10

10

8

7

14
12

6965
7548

Q/ Species includes those which contributed 2% or more of total bites during any month.

tr

1

2

7

12

o
o

10

17

tr

4

2

4

o

tr

1

14

11

5

5

13
tr

36
31

10
11

25
13

2

3
1

tr
2

o

tr

1

3

1

5
2
2

4

6

16

7

4
7

34
31

11
8

1

6180
7139

5595
3886

Means are calculated across individual animal diet percentages for each month.

2

9
9
6
2
1

16

7941
6719

6340
4500

Sum of total bites across animals.

1

tr
1

1

tr
tr

6774
6274

~/ Includes all species selected, not just those in table.

i/

4

9

6

11

V1
I

�Table 2.
Percent crude
summer range in Colorado

protein and IVDDM of principle forage
during July-September
1977 and 1978.

Tax~/

Year

July

species

consumed

Percent

of Dry Hatter

Crude Protein
August

by tame elk on subalpine

IVDIJH b7
August

September

.July

10.8/

c_r:j

C 5::./

cS:./
69
59
58
53
61
65
52
61
60(±8.3)
60(±5.9)

c 5::./
63
61
52
63
57
64
54
C 5::./
57(+5.6)
62 (±:2.0)

42
48
32
35
20
20
31(+18.5)
34 (i:Z3. 7)

39
37
30
30
17
18
29 (+18. 6)
28(E16.2)

tundra

September

GRAHlNOIDS
Calamagrostis
Carex

canadensis

spp.

Deschampsia
Juncus

caespitosa

balticus

Kobresia

myosuroides

X Graminoids

(±90% C.I.)

C 5::./

1977
1978
1977
1978
1977
1978
1977
1978
1977
1978
1977
1978

17.2
15.0
12.8
11.5
16.1
14.6
13.8
14.3
15.0(±2.3)
13.9(±1.9)

cS:./
cS:./
14.5
15.3
8.7
15.3
11. 8
15.2
12.0
cS:./
11.8(±2.8)
15.3(± .1)

1977
1978
1977
1978
1977
1978
1977
1978

16.4
18.0
16.8
14.0
10.4
9.5
14.5(+6.0)
13.8(i).1)

11.9
13.4
13.2
14.0
9.4
10.3
11. 5(+3. 3)
12.6
4)

1977
1978
1977
1978
1977
1978
.1977
1978
1977
1978
1977
1978

C 5::./
17.6
17.3
15.5
C 5::./
C 5::./
22.7

12.7
14.7
11. 4
13.7

C 5::./

cs

10.7
9.8
8.9
14.6
7.0
11.3
C 5::./
. C 5::./
9.4(±2.1)
12.0 (±3.3)

57

c 5::./
47
60
53
62
45
61

C s_l

C 5::./
51(+6.5)
61(~). 36)

SHRUBS
I

Salix

brachycarpa

Salix

planifoli~

Vaccinium

spp.

X Shrubs

(±90% C. I.)

(:!).

8.6
8.9
8.8

10.7
7.9
7.9
8.4(+0.8)
9.2 (i:2. 4)

N
W

26
32
27
34
17
21

(J\

I

23 (+9.3)

29(±11.8)

FORBS
Caltha

~tosepala

Geum rossii
----Polygonum

bistortoides

Trifolium

~

Trifolium

parryi

X Forbs

~/ Species

contributing

2% or·more

£/

digestible

dry matter.

In vitro

s./ Sp ec Le s did not occur
&lt;7

in diet

C 5::./

C 5::./
C !:./
20.0
16.6

of total bites

that month.

8.3
9.1
C
C

C 5::./

16.2/
C5::.
C 5::./
21.8
c 5::./
15.3
14.9

observed

c/

c/
C c/
C c/
C c/
C

£/

16.6
c 5::./
12.5
9.1

during

any month.

76.0
73.5
43.7
46.4
C 5::./
C 5::./

67.6
C 5::./
C s_l

C 5::./

62.4
60.0

64.8
58.7
38.4
38.5
C 5::./
60.9
c 5::./
C

s/

68.9
C 5::./
57.4
53.7

40.5
48.0
C c/

c/
c/
C c/
C c/
C £/
C
C

56.2
C &amp;1.
4f!.4
48.0

of'

�Table 3. Mean percentage with 90 percent confidence intervals
diets of tame elk in 3 sub-alpine-tundra habitats in Colorado,
Percent

for crude proteiri and IVDDM in summer
during July to September 1977 and 1978.
of Dry Matter

Crude Protein
August

September

July

IVDDM~_l
August

September

Habitat

Year

July

Willow-park

1977

16.4(±0.R)

12.9(±1.0)

9.8(±0.6)

53(±5.5)

50(±0.8)

41(±loO)

1978

14.4(±O.9)

14.0(±0.6)

10.1(±0.1)

49(±4.0)

47(±8.1)

SO(±0.6)

Krummholzecotone

1977
1978

16.4(±0.6)
14.7(±0.6)

13.0(±0.6)
15.0(±0.5)

10.0(±0.4)
1lo4(±0.0)

S6(±5.6)
47 (±4. 7)

52 (±lo 8)
55(±12.1)

42(±2.3)
49 (±O. 4)

Alpine tundra

1977
1978

17.1(±0.81)
1S.2(±0.3)

14.5(±lo5)
14.2(±0.6)

10.8(±0.4)
10.1(±O.4)

53(±7.1)
47(±2.9)

59(±2.7)
47(±9.2)

47(±1.4)
51(±2.92)

I
N

w

~
I

a/ In vitro digestible

dry matter.

�-238-

APPENDIX I

�-239-

DISSERTATION

WINTER DIET QUALITY AND NUTRITIONAL STATUS
OF ELK IN THE UPPER MONTANE ZONE, COLORADO

Submitted by

N. Thompson Hobbs
Fishery and Wildlife Biology

In partial fulfillment of the requirements
for the Degree of Doctor of Philosophy
Colorado State University
Fort Collins, Colorado 80523
Fall 1979

�-240-

ABSTRACT OF DISSERTATION
WINTER DIET QUALITY AND NUTRITIONAL STATUS
OF ELK IN THE UPPER MONTANE ZONE, COLORADO
Studies were conducted

to relate forage resources on upper montane

winter range in Colorado to nutritional

quality and botanical

of elk winter diets and to simulated elk nutritional

composition

status during 1976

to 1978.
Standing crop biomass of winter range herbage was highly variable
between years and among plant cowmunities,

ranging from 216.6 g • m-2 in

willow during 1976 to 67.5 g • m2 in grassland

the following year.

Browse species contained more crude protein,
lignin and less cellulose
than grasses.

and in vitro digestible

Browse nutritional

cell solubles;' and
dry matter

(IVDDM)

quality remained constant with advan-

cing season, while grasses decreased

in IVDDM and crude protein but

increased

in fiber content.

Grasses and browse contained less crude

protein,

and browse less IVDDM, during 1977-1978 than during the

previous year.
Graminoids
in importance,

dominated

diets

(X = 61%) during

1976-1977, but declined

from 66% of observed bites in November

as browse consumption

increased.

to 44% in February,

Browse and grass contributed

to diets through the winter of 1977-1978,

accounting

equally

for more than 90%

of bites each month.
Diet quality was influenced
quality; from November

to some extent by changes in forage

to March 1976 to 1977, dietary crude protein

�-241-

declined from 5.8 to 4.9%, and IVDDM declined from 49 to 40%.

Over the

same period the following year, dietary crude protein declined from 5.4
to 4.6%; IVDDM, from 42 to 37%.

During 1976-1977, mean overwinter

dietary protein ranged from a high of 6.3% in dry grassland to a low of

4.6% in mesic meadow communities, while dietary IVDDM ranged from 47% in
aspen to 35% in grasslands.

During 1977-1978, protein was highest

(5.6%) in willow and lowest (4.6%) in mesic meadow, while IVDDM was
highest

(39%) in aspen and lowest (35%) in willow.

variation

Despite year-to-year

in forage quality, elk were able to maintain a relatively

stable diet quality over time and space by shifting the forage class mix
of diets.
Simulation modeling of elk energy and nitrogen
more favorable overwinter

1977-1978.

status predicted

animal condition during 1976-1977 than during

Simulated weight losses were relatively

small the first

winter (10% of body weight) but were much more substantial
year (20-30% of body weight).

Habitat types could not be consistently

ranked on the basis of their ability to contribute
animal condition.

the second

to maintenance

of

Carrying capacity of habitats based on the quantity

of forage present was not related to nutritional

status of elk feeding

in those communities.
Temporal changes 1n the quality and quantity of forage resources
appeared to be related to density-independent
we should expect some stochastic variation
individual elk and in performance

in nutritional

of elk populations.

could be reduced by providing wintering
choices and with browse-grass

effects of weather.

Thus,

status of

This variation

elk with a diversity of habitat

mixtures within habitats.

�-242-

Effects of variation

in forage resources on elk nutritional

are examined in the context of current theory on competition,
and diet selection,

and population

status

habitat

regulation.

N. Thompson Hobbs
Fishery and Wildlife Biology Department
Colorado State University
Fort Collins, Colorado 80523
Fall, 1979

�-243-

TABLE OF CONTENTS
Page

Chapter
I.

INTRODUCTION

1

II.

STUDY OBJECTIVES

3

III.

LITERATURE REVIEW

6

The Problem:

Herbivory and Nutrient Limitation

6

Constraints on Diet Selection

9

Models of Diet Choice

11

i) Theoretical Models

11

ii) Empirical Models

14

Methods of Diet Analysis

16

i) Observation of Diet Choices of
Tame Animals

16

ii) Assessment of Dietary Preference

19

iii) Measurement of Forage Quality

21

Elk Winter Food Habits

23

IV.

STUDY AREA

25

V.

METHODS .

29

V.

Diet Observations

29

Forage Quality

31

Diet Data Analysis . . . .

32

Estimation of Herbage Biomass

33

Simulation Modelling of Nutritional Status

34

Carrying Capacity Estimation

38

RESULTS ...
Herbage Biomass

40
40

�-244-

TABLE OF CONTENTS
Chapter

VI.

VII.

Page
Diet Botanical Composition

42

Forage Quality

52

Diet Quality .

63

Nutritional Status

73

Carrying Capacity

79

DISCUSSION

83

Herbage Biomass

83

Forage Quality

85

Diet Botanical Composition

88

Diet Quality: Interactions of Forage Quality
and Diet Botanical Composition . . . . . .

90

Elk Nutritional Status and Carrying Capacity

96

CONCLUSIONS

. . . . . . . .

100

Theoretical Implications

100

i) Optimal Foraging and Competition Theory

100

ii) Theory of Population Regulation

102

iii) Habitat Selection

103

Management Implications

105

i) Population Management

105

ii) Habitat Management

106

LITERATURE CITED
APPENDICES
1.

Rearing and Training Elk Calves for
Use in Food Habits Studies

2.

Plant Species Biomass Data

108

�LIST OF FIGURES
Figure
1.

2.

3.
4.

Page
Control model of hypothesized interactions
among variables examined in this study. Arrows
indicate influence of one variable on another .

4

Geographic location of Rocky Mountain National
Park in Colorado
.

26

Structure of generalized model of ruminant
energy and nitrogen metabolism

36

Total abovegroung ,biomass production at t4e
end of the growing season during 1976 and 1977
in upper montane habitats, Colorado. Values
are indicated by Xj non-overlapping verticle
lines indicate significant difference at P
.05.
Solid lines are for 1976, dashed for 1977. Abbreviations are: WILW
willow, PPSH
ponderosa
pine shrub, WSHM
wet shrub meadow, ASPN
aspen,
SAGE
sagebrush, WGSM
wet meadow, GRSL
grassland, MGSM
mesic meadow
.

41

Biomass of grass at the end of the growing season
during 1976 and 1977 in upper montane habitats,
Colorado. Mean values are indicated by X; nonoverlapping verticle lines indicated significant
difference at P = .05. Solid lines are for 1976,
dashed for 1977. Abbreviations are: WILW = willow,
PPSH
ponderosa pine shrub, WSHM
wet shrub meadow,
ASPN
aspen, SAGE
sagebrush, WGSN
wet meadow,
GRSL = grassland, MGSN
mesic meadow
.

43

Biomass of current annual growth of shrubs during
November of 1976 and 1977 in upper montane habitats,
Colorado. Mean values are indicated by Xj nonoverlapping verticle lines indicate signHicant
differences at P = .05. Solid lines are for 1976,
dashed for 1977. Abbreviations are: WILW
willow,
PPjSH
ponderosa pine-shrub, WSHM
wet shrub
meadow, ASPN
aspen, SAGE
sagebrush, WGSH
wet
meadow, GRSL
grassland, MGSM = mesic meadow

44

=

=

5.

=

7.

=

=

=

=
=

6.

=

=

=

=

=

=

=
=

=

=

=

=

=

=

Biomass of forbs at the end of the growing
season during 1976 and 1977 in upper montane
habitats, Colorado. Mean values are indicated
by X; non-overlapping vertical lines indicate
significant difference at P
.05. Solid lines
are for 1976, dashed for 1977. Abbreviations are:
wnw willow, PPjSH = ponderosa pine-shrub, WSHH =
wet shrub meadow, ASPN
aspen, SAGE
sagebrush,

=

=

=

=

�-246-

LIST OF FIGURES
Page

Figure

8.

9.

10.

11.

12.

l3.

14.

15..

WGSM = wet meadow, GRSL = grassland, MGSi"l= mesic
meadow

45

Botanical composition of elk winter diets estimated across habitat types on upper montane winter
range, Colorado, during 1976-77 and 1977-78 ...

50

Relationship between proportion of forage classes
in prewinter herbage biomass and their proportion
in overwinter diets of elk on upper montane winter
range, Colorado, during 1976 and 1977. Each point
represents the mean forage class percentage across
5 elk in one habitat type during one winter. Symbols are: S = shrubs, G = grass, F = Forbs ...

51

Relationship between proportion of 4 most common
species in Aspen, Grassland, Ponderosa-Pine Shrub,
and Sagebrush habitats and their mean percentage
.in winter diets across 5 elk on upper montane
winter range, Colorado, during 1976-77 and 1977-78

54

Forage class composition of elk winter diets in
upper montane habitats during 1976-1977 and
1977-78 . . . . . . . . . . . . . . . . . . .

55

Cell wall content of grasses consumed by elk
on upper montane winter range, Colorado.
Numeral 2 indicates a value for 1977-78;
numeral 1 is a value for 1976-77. For
regression statistics see Table 4 . . . . .

59

Protein content of grasses consumed by elk
on upper montane winter range, Colorado.
Numeral 2 indicates a value for 1977-78;
numeral 1 is a value for 1976-77. For
regression statistics see Table 4 . . . .

60

In vitro dry digestible dry matter (IVDDM)
for grasses consumed by elk on upper montane
winter range, Colorado. Numeral 2 indicates
a value for 1977-78; numeral 1 is a value for
1976-77. For regression statistics see Table 4 .....

61

In vitro digestible dry matter (IVDDM) of
winter diets of 5 elk on upper montane winter
range, Colorado, during 1976-77 and 1977-78 ..
Numeral 1 indicates a value for 1976-77.
Numeral 2 is a value for 1977-78. For
regression statistics see Table 6 . . . . . . . . . . ..

64

�-247-

LIST OF FIGURES
Figure
16.

17.

Page
Protein content of winter diets of 5 elk on
upper montane winter range, Colorado, during
1976-77 and 1977-78. Numeral 1 indicates a
value for 1976-77; numeral 2 is a value for
1977-78. For regression statistics see
Table 6
. . . . .
. . . .
Protein content of winter diets of 5 elk
in upper montane habitats, Colorado. X
indicates a mean across animals; vertical
bars are Tukey's Q simultaneous confidence
intervals for eight comparisons; nonoverlapping bars indicate significant differences at P
.05. Solid lines are for
1976-77, dotted for 1977-78. Habitat abbreviations are: WILW
willow, PP/SH
ponderosa pine-shrub, WSHM
wet shrub
meadow, ASPN = aspen, SAGE = sagebrush,
WGSM = wet meadow, GRSL = grassland,
MGSM
mesic meadow

65

=

=

=

=

=

18.

.

69

In vitro digestible dry matter (IVDDH)
content of winter diets of 5 elk in upper
montane habitats, Colorado. X indicates
mean value across animals; verticle bars
are Tukey's Q simultaneous confidence
intervals for 8 comparisons. Non-overlapping
bars indicate significant difference at p
0.05. Solid lines are for 1976-77, dotted for
1977-78. Habitat abbreviations are: WILW
willow, PP/SH
ponderosa pine-shrub, WSHM
wet shrub meadow, ASPN
Aspen, SAGE
sagebrush, WGSM
wet meadow, GRSL
grassland,
MGSM
mesic meadow . . . .
. . . . .

70

Cell wall content of winter diets of 5 elk
in upper montane habitats, Colorado. X
indicates mean value across animals; vertical
bars are Tukey's Q simultaneous confidence
intervals for 8 comparisons. Non-overlapping
bars indicate a significant difference at
p
0.05. Solid lines are for 1976-77,
dotted for 1977-78. Habitat abbreviations
are: WILW
willow, PP/SH = ponderosa pineshrub, WSffif wet shrub meadow, ASPN = aspen,
SAGE
sagebrush, WGSM
wet meadow, GRSL
grassland, MGSM = mesic meadow
....

71

=

=

=

19.

=

=

=

=
=

=

=

=

20.

=
=

=

Simulated changes in total weight, lean
body and fat or an adult elk on upper

=

�-248-

LIST OF FIGURES
Figure

21.

22.

23.

24.

25.

Page
montane winter range, Colorado, during NovemberMarch 1976-77 ..
.

74

Simulated changes in total weight, lean body and
fat of a calf elk on upper montane winter range,
Colorado, during November-march 1976-77 ...

75

Simulated changes in total weight, lean body, and
fat of an adult elk on upper montane winter range,
Colorado, during November-March 1977-78 ...

76

Simulated changes in total weight, lean body, and
fat of a calf elk on upper montane winter range,
Colorado, during November-March 1977-78 ....

77

Relationships between browse content of winter
diets of 5 elk and mean crude protein content of
grasses on upper montain winter range, Colorado,
during 1976-77 and 1977-78

93

Relationship between in vitro digestible dry matter
(IVDDM) and browse content of diets of 5 elk on
upper montane winter range, Colorado, during
1976-77 and 1977-78 . . . . . . . . . . . . . . . . . ..

95

�-249-

LIST OF TABLES
Page

Table
1.
2.

3.

4.

5.

6.

7.

8.

Nutritional quality of wild ungulate winter
forages in the Rocky Mountain region

8

Summary of estimates of forage class composition of elk winter diets in the Rocky Mountain
region

24

Site characteristics and principle plant species
of upper montane vegetation types in Rocky
Mountain National Park, Colorado
.

27

Parameter specifications for the ruminant
simulation model runs representing elk on
upper montane winter range, Colorado

37

Mean percentage of observed bites of plant
species in diets of 5 elk on upper montane
winter range in Rocky Mountain National Park,
Colorado, during November-March 1976-77 and
1977-78, summarized by habitat type

46

Percentage of 4 most common species in biomass
of upper montane habitats, Colorado and in
winter diets of 5 elk feeding in those habitats
during Novembe r-Ha rch 1976-77 and 1977-78 . . .

53

Chemical constituents and in vitro digestibility of
principle forage species consumed by 5 elk on upper
montane range, Colorado, during November-March 1976-77
and 1977-1978 . . . ..
.

57

Regression coefficients and statistics for in vitro
digestible dry matter (IVDDM), crude protei~ ~
cell wall of grasses' from upper montane winter range,
Colorado, as a function of advancing season.
Independent variable is days where 15 October
day 1. See Fig. 2-4 for plots
....

62

Regression coefficients and statistics for winter
diet crude protein and in vitro digestible dry
matter (IVDDM) of 5 elk on upper montane winter
range, Colorado, as a function of advancing season.
Independent variable is days where 15 October
day 1.
See Fig. 5 and 6 for plots . . . ..
.....

66

=

9.

=

10.

Mean percentage chemical constituents and
in vitro dry matter digestibility of winter
diets of 5 elk in upper montane habitats in

�-250-

LIST OF TABLES
Table

Page
Colorado during November-March
1977-1978
. . . ..

11.

12.

1976-77 and
. ....

68

Simulated estimates of energy and nitrogen
status of elk in upper montane habitats
during November-March of 1976-77 and 197778 in Colorado
. . . . . . ..
....

80

Estimates of nutritional status and earring
capacity for elk in upper montane habitat
types during winter 1976-77 and 1977-78 in
Colorado
.

81

�-251-

Chapter 1
INTRODUCTION
Why study animal diets?

Diet selection controls the transfer of

energy and materials between trophic levels.

As such, it affects the

function of ecosystems and is, in turn, influenced by ecosystem change.
The diets animals choose determine the partitioning

of food resources

between species, and this division of resources affects the structure of
communities.

Important consequences

direct way, the determination
effects on performance

of animal condition, and more indirectly,

of populations

of animal diets, therefore,

and condition of habitats.

furthers our understanding

interactions within ecosystems,
animal populations

of diet selection include, in a

and contributes

Study

of fundamental

to wise management

of

and the habitats they occupy.

Winter in the Rocky Mountain region is generally a period of nutritional deprivation

for wild ruminants.

Dormant winter plants are less

digestible and contain less protein than summer forage (Dietz 1967,
Milchunas et al. 1978), deep sQow may render large portions of forage
supplies unavailable

(Loveless 1967, Regelin 1976, Leege and Hickey

1977), and conversion of traditional winter range to other uses relentlessly shrinks suitable winter habitat.

Given these constraints

on

available winter food, selection of adequately nutritious diets may
become difficult.
Although winter food habits of elk have been extensively

studied

(for review see Kufeld 1972), our knowledge of elk diets remains incomplete.

Predicting

range carrying capacity ultimately

depends on

�-252-

estimates of diet and forage quality (Wallmo et al. 1977), yet estimates
of chemical composition

and digestibility

of elk winter diets are not

available, and information on the nutritional quality of elk forage is
scant (Ward 1971, McCullough

1969:105-113).

Few studies have described

the relation between winter diets and plant communities chosen for
feeding.

Consequently,

although some vegetation types are thought to be

better winter habitat for elk than other communities,

these differences

have never been demonstrated

Overwinter changes

on a nutritional basis.

in forage quality and availability can markedly alter elk nutritional
status (Flook 1970); however, dietary responses of elk to within-season
variation in forage quality have not been examined.
whether elk are selective or non-selective
generalists,

grazers or browsers.

feeders, specialists or

Description

of diet composition has

not yet extended to prediction of elk nutritional
fundamental to understanding

status, an extension

their trophic ecology.

What are the potential contributions
diets?

It remains unclear

of a study of elk winter

A long term outcome is a more thorough understanding

of how

variation in food resources affects the resource utilization patterns of
ungulates.

A more immediate result offers application to management

problems.

The afore mentioned depletion of winter habitat for elk

populations

requires that elk ranges must be improved in quality to

mitigate reductions in their size.
identification

Habitat improvement depends on

of plant communities and individual forages which can

best provide for maintenance
winter diets contributes

of animal condition.

Thus, study of elk

to development of theory of trophic relations

of ungulates to the ecosystems they occupy and provides information
useful for management

of elk populations

and their winter habitat.

�-253-

Chapter II
STUDY OBJECTIVES
The global objective

of this study was to describe the processes

affecting the nutritional

quality of elk diets and to predict the poten-

tial effects of variation

in diet quality on animal condition.

A schematic of the variables
interrelationships

is presented

examined in this study and their

in Fig. 1.

diets is determined by the interaction
with nutritional

Nutritional

of diet botanical

quality of forages eaten.

quality of elk
composition

Diet choice, in turn, is

influenced by the quality and quantity of available
influenced by the location and time of feeding.

foods, which are

The quality of diets

affects the amount of food an elk can consume and influences
logical condition.

To examine this system of interacting variables,

subobjectives

of this study were to

1.

the prewinter

Estimate

its physio-

standing crops of potential

the

elk food in

habitats used for feeding.
2.

Measure the chemical composition
forages throughout

3.

Determine

of those

the winter.

the botanical

lation to advancing

and digestibility

composition

of elk winter diets in re-

season and plant communities

chosen for

feeding.
4.

Evaluate the effects of pre-winter

forage availability

over winter forage quality on diet botanical
5.

Estimate the nutritional

composition.

quality of elk diets in relation to

habitat types chosen for feeding, and overwinter
forage resources .
-. ;.....

and

changes in

�-254-

Temporal

locatiof

rcation

Forage

&lt;.
Diet

/orage
botanical

_. .._. __ /iet_:ali\_
SIMULATED
VARIABLES

Figure 1.

quality

comrSition

MEASURED
VARIABLES

Voluntary

of feeding

intake

-Animal

condition

1

.

Food requirements

Control model of hypothesized interactions among variables
examined in this study.
Arrows indicate influence of one
variable on another.

�-255-

6.

Using simulation modelling, predict the nutritional status of
elk which results from diets of differing quality.

7.

Compare estimates of habitat carrying capacity based on forage
quantity with predictions of elk nutritional status.

8.

Examine annual variation in the above variables.

�-256-

Chapter III
LITERATURE
The Problem:

Herbivory

REVIEW

and Nutrient Limitation

Animals ... spend the greater part of their lives eating.
Feeding is such a universal and commonplace business that we
are inclined to forget its importance.
The primary driving
force of all animals is the necessity of finding the right
type of food and enough of it. Food is the burning question
in animal society, and the whole structure and activities of
the community are dependent upon questions of food supply.
(Charles Elton 1927:56)
Despite Charles Elton's early emphasis on food as the principal
limitation

of animal populations,

many ecologists

the seeming paradox of food limitation
(Hariston et al. 1960, Van Valen 1973).
paradox

is that the world is sometimes

have been perplexed

of herbivores

in a "green world"

A parsimonious
"brown"

during the dry season, temperate grasslands

by

solution to this

(observe tropical savanna

in winter).

A more resolute

answer is that all that is green is indeed not food.
The fundamental
adequate nutrients

problem confronted
from a ubiquitous

Plants are ultimately
White (1978:77),

is extraction

because,

in the words of

any species which was adequately

to a herbivore

of

but poor quality source of food.

poor foods for herbivores

"Presumably,

and freely available

by herbivores

throughout

nutritious

its life cycle would soon

be exterminated."

Thus, natural selection has resulted in active defense

against herbivory

through chemical and physical

Freeland

adaptations

(Levin 1976,

and Janzen 1974, Erlich and Raven 1974, Whittaker

1970, Berger

1977, Leopold

1976, Labov 1977) and through dilution of nutrients

quired by herbivores,
drates (White 1978).

particularly

nitrogen,

with structural

re-

carbohy-

�-257-

Dilute concentrations

of nutrients are characteristic

forages in the Rocky Mountain
during plant maturation

region (Table 1).

of winter

This dilution occurs

and growth as fibrous tissue is elaborated more

rapidly than cell solubles.

Consequently,

winter diets of wild ungulates

are commonly low in protein and digestibility.
Fulgham

(1978) found mule deer (Odocoileus hemionus)

diets in early

winter in northern Utah contained less than 8.5% protein and less than
50% in vitro digestible

dry matter (IVDDM).

Mule deer winter diets in New Mexico contained
were only 23% digestible

12% protein, but

(Boeker et al. 1972).

Wallmo et al. (1977) estimated that protein content of mule deer
diets during late winter in central Colorado contained

less than 40%

digestible drymatter and less than 7% protein.
Pronghorn antelope

(Antilocapra americana) winter diets on the

shortgrass prairie in Colorado contained 7% protein and were less than
40% digestible

(Schwartz 1977).

Although evolution of symbiosis between cellulolytic

bacteria and

their ruminant hosts would seem to'offer the ruminant a virtually unlimited supply of energy contained in plant fiber, there are important
constraints' on the availability

of that energy.

First,. tint rumen envi-

ronment must provide sufficient nitrogen for microbial

growth since

activity of fiber digesting bacteria is retarded by deficiencies
nitrogen

(Van Glswyk 1970, Schwartz and Gilchrist

1975).

in

Although some

nitrogen enters the rumen through recycling of urea (Wales et al. 1975,
Robbins, et al. 1974, Allen and Miller 1976 Nolan and Leng 1973, Westra
and Hudson 1979), a significant
vided by forage substrates.

exogenous portion must always be pro-

For ruminants consuming fibrous forages,

rates of cellulose digestion and microbial protein synthesis are directly

�Table 1. Nutritional quality of wild ungulate winter forages in the Rocky Mountian region.
Dry Matter
Digestibility (%)

Crude Protein (%)
Authority

Grass

Forbs

Shrubs

Grass

Forbs

Shrubs

Central Colorado

Milchunas et al. 1978

3.7-5.0

3.0-5.0

4.5-7.6

33-52

8-29

29-64

NW Colorado

Gibbs 1978

NE Co lorado

Schwartz ·1977

N. Central Colorado

Young and Bailey 1975

S. Wyoming

Ward 1971

N. Central Colorado

Dietz 1967

N. Central Colorado

Short et al. 1966

S.W. Colorado

Dietz 1958

8.2-13.5

Utah

Smith 1954

6.2-11.0

Area

7.6-80

7.8

Grass

Forbs

Shrubs

I

1.6-2.5

5.2-10.2
6.2-10.0

Energy (kcal/g)

37-46

14-35

52

38-50

40-53

22-59

N
\Jl

CO
I

8.0

7.6-11.1
1.97-2.24

�-259-

related to nitrogen content of the forage (Campling et al. 1962, Hume et
al. 1970, Van Glswyk 1970)
An additional

constraint on energy assimilation

volves bulk limitation of voluntary

intake.

by ruminants in-

Fore-stomach

digescion

limits the amount of fibrous forage which can be consumed per unit time
to that amount which can pass the rumen (Ammann et al. 1973, Campling
1970).

Rate of passage is a function of forage particle size reduction

which, in turn, is controlled by physical and chemical characteristics
of the forage and the quality of the rumen environment
Smith 1968, Mertins 1973, Milchunas
this limitation on voluntary

et al. 1978).

(reviewed by

Thus, it follows from

intake that ruminants on poor quality diets

can not compensate

for the dilution of nutrients

forage consumption

rates (Janis 1976).

in forage by increasing

The importance of these two interacting phenomena,

nutrient limita-

tion of rumen bacteria and bulk limitation of voluntary

intake, is that

deficiencies

in nutrients, particulary

nitrogen,

acquisition

of energy in the face of seemingly abundant forage supplies.

Constraints

on Diet Selection

A solution to this dilemma is selective

can impede a ruminants

feeding, consumption

of

only those plants and plant parts which contain relatively high concentrations of nitrogen and other nutrients.

There is abundant evidence

(reviewed in later sections) that grazing ruminants can exert such diet
selection.
The ability of a rumiant to feed selectively
aspects of its physiology,

anatomy, and ecology.

is limited by several
Physiological

require-

ments for energy and nitrogen limit the extent to which animals can
consume rare foods.

A ruminant's requirement

for energy and nitrogen is

�-260-

functionally
1975).

related to its body weight raised to the 3/4 power (Kleiber

As a consequence

of this relation,

greater total nutrient intake requirements

large bodied animals have
but smaller requirements

unit of body mass than animals of smaller body size.
these allometric

relationships

per

The importance

ot

for diet selection is that large animals

must eat large quantities of forage, but can consume relatively

low

quality forage, while small animals can eat relatively

less, but must

consume foods with higher concentrations

(Janis 1976, Bell

1971, Jarman 1974, Geist 1974).

of nutrients

Thus, large herbivores

by their total intake requirements

to eat relatively common foods while

small animals, with their lower total food requisites,
foods which are more scarce.
feed more selectively

are constrained

can afford to eat

It follows, then, that small herbivores

than large herbivores.

A corollary to these physiological

constraints

on diet selection by feeding morphology.

is the limit placed

Herbivores with a narrow jaw

can select among plants and plant parts which can not be separated by
animals with larger mouth parts (Hoffman 1973, Schwartz 1977).
for a wide variety of North American

and African ungulates,

Further,

feeding

style has been correlated with anatomy of the digestive tract.

Hoffman

(1973) classified African wild ruminants into roughage feeders (eating
mainly grass), selective feeders (eating browse tips, forbs, and new
growth of grasses) and transitional
Selective

forms (eating grasses and browse).

feeders tended to have small rumens with few "barriers"

rumen and omasum to impede passage of ingestion.
contains a high proportion

in the

Since their diet

of cell solubles, selective feeders can rely

in large part on enzymatic digestion of these soluble sugars and proteins
and consequently

do not require protracted

\.

I

retention of forage fiber for

�-261-

microbial

fermentation.

In contrast,

roughage eaters obtain a large

portion of their energy from digestion
of efficient

fiber digestion

between digestive

of fiber.

correlate

is a large rumen and physical barriers

organs to slow down rate of passage and allow more

complete fermentation

of ingested forages.

rumen anatomy and feeding style observed
pronghorn

A necessary

A similar correlation

between

for bison, cattle, sheep, and

was observed by Schwartz and Ellis (submitted).

In addition to these physiological
feeding style, diet selection
animal's ecology.

Weins (1977) and Cody (1974a) have suggested
are food specialists,

tend to be food generalists.

costs of searching

correlates

for rare food items by the specialist

rare foods.

Jarman's

relates of social organization
hypothesis,

particularly

that

while habitat

They reasoned that increased
cause it to use

its habitat in a more fine grained manner, thus increasing
available

of

is further limited by aspects of the

animals which are habitat generalists
specialists

and anatomical

the amount of

(1974) elegant work on ecological

cor-

in African antelope substantiates

this

with the considered

unit of diet selection

includes plant parts as well as plant species.
Models of Diet Choice
i) Theoretical

Models

Given the above constraints
theoretical

on diet selection,

basis, the outcomes of diet choice?

to exhaustively

review theory of diet selection.

can we predict,

on a

No attempt will be made
(For such review, see

Rapport and Turner 1977, Pyke et al. 1977, Schoner 1971, Cody 1974b,
Ivlev 1967).

Instead, the essential

lems associated

with its application

be discussed.

features of that theory, and probto diets of large herbivores

will

�-262-

Optimal foraging theory relies on three assumptions

1977, Pulliam 1974).
foraging behavior

It is assumed that behavior

in particular,

is a heritable

(Pyke et al.

in general, and

trait, that environ-

mental resources constrain the range of forage choices, and that natural
selection acts to favor inheritance
maximal fitness.

In general, such behaviors

energy intake, minimize
nutrient balance

of those behaviors which convey

time spent feeding (Schoener 1971) or optimize

(Westoby 1974, Pulliam 1975, Katz 1974).

ever, that at any time, in any population,
optimal feeding behavior.
distribution

are thought to maximize

we would expect to see sub-

This is the case because the frequency

of feeding traits must by definition

include some behaviors

less optimum than others and, further, because consumption
present environments
most appropriate

Note, how-

reflect past selective pressures

for survival and reproduction

choices in

and may not now be

(Weins 1977, Rapport and

Turner 1977).
With one exception,
extremely

(Ellis et al. 1976) optimal foraging models are

similar; in the words of Pyke et a1. (1977), "the same essen-

tial result being reached independently
This result is obtained,
Upon encountering

by no less than 9 authors".

in the form of McArthur

(1972), as follows.

a food item, a consumer must choose to either eat the

item, or search for another.

Since either choice eventually

initiates a

new search, the better choice is the one which promises to yield more
energy (or other optimized

unit) per unit time.

Deciding to eat the

food requires its "pursuit"; pursuit time being the temporal interval
from discovery of food to its consumption.

An animal should pursue an

item if and only if during the time the pursuit would require, a "better"
forage item could not be located.

Stated mathematically,

if Pj is

�-263-

pursuit time per calorie ingested of food item j, and ~ and § are the
then item j should be

mean pursuit and search time per calorie obtained,
eaten if and only if
P. ~ ~ + S
J

Thus, this formulation
its associated

accounts for the amount of energy acquired and

temporal costs.

There are limitations
by large herbivores

to application

(Westoby 1974).

of this model to diets selected

First, in the sense of a predator

pursuing prey, .pursuit time for a herbivore
Rather than the time necessary
herbivore

can more realistically

to assimilate

nutrients

sumed by carnivores

be thought of as the interval required

can justifiably

trivial in consideration

Assimilation

Thus, differences

in assimilation

animal tissue, forages are extremely heterogenous

intake by ruminants

rates for food con-

be assumed equal since their food is

of predator diet choices.

their energy and nutrients

to zero.

to capture food once found, pursuit for a

once captured.

of uniformly high quality.

reduces essentially

can be extracted.

rate are

In contrast to
in the rate at which

Further,

since voluntary

is bulk limited on most natural. diets, inclusion of

slowly. digested material decreases
ingest during subsquent

the. a~ountof

time intervals.

forage an animal can

Consequently,

if "pursuit time"

is expanded to include the time from which an item is eaten to the time
at which it has passed from the animal, then the above theory can be
realistically

applied to a ruminant herbivore.

Note, however, that this

realism is achieved only at the expense of an enormous increase in mathematical complexity,
nonlinear
1971).

since rate functions

(Milchunas et al. 1978, Mertins

for passage of forages are
1973, Mautz and Petrides

�-264-

Further complexity

results from the heterogeneity

bility and nitrogen content.

This variation

perties to the process of diet choice.
consider maximization
probably

of one variable

of forage digesti-

conveys multivariate

That is, it is not sufficient

to

(i.e., energy), since plants are

chosen on the basis of several nutritional

criteria.

attempts have been made to include nutrient constraints
models

pro-

Some

in optimization

(Pulliam 1975, Katz 1974, Westoby 1975); but, in general, these

simple formulations

are not up to the task of integrating

complex inter-

actions between forage chemical constituents.
The most mechanistic

approach to modelling

diet selection

is that

of Ellis et al. (1976) who developed a conceptual model of the flow of
foods from the habitat to the animal.
array of ecosystem variables
requirements,

Flows were controlled by a large

including factors affecting consumer food

consumer preference

and selectivity,

and food availability.

ii) Empirical Models
It is important to differentiate

between models which predict total

3mount of food eaten and those which predict the kinds of foods selected
and their relative contribution
been predicted

to the diet.

While voluntary

intake has

fairly well on the basis of physical and chemical charac-

teristics of forage (reviewed by Mertins 1973, Campling 1970) the prediction of composition

of diets based on forage constituents

with notable lack of success.
"Numerous associations
palatability

has met

Thus, Marten (1978: 1472) concluded that

~eported between plant

characteristics

and plant

to ruminant animals have proven to be highly situation

specific, making them worthless
Marten and Anderson

as general selection criteria."

(1975) compared nutritive value of 12 common

annual weed species with their relative palatability

for sheep and found

�-265-

no association between forage preference

and its protein and mineral

content or its dry matter digestibility.
Ivins (1952) concluded that selective grazing was not determined by
the chemical composition of plants.
cattle preference

Though he did find differences

for different species of forage, these differences

could not be consistently predicted on a nutritional
Radwin and Crouch (197~) studied preference
(Odocoileus hemionus columbianus)
fiber characteristics,
non-reducing

in

basis.

of black-tailed

deer

in relation to forage protein content,

total available carbohydrates,

reducing and

sugars, dry matter, mineral content, crude fat, essential

oil content, moisture, and dry matter and cellulose digestibility.

They

found no consistent chemical basis for plant species preference:
Tucker et a1. (1976) found no correlation between chemical composition of Douglas fir (Pseudotsuga menziesii)

foliage and mule deer preference.

Scotter (1965) examined caribou (Rangifer tarandus) preferences
species of lichens and found no relationship

for

between their protein

content and frequency of consumption.
Alexander
preferences

(1979) found no consistent relationship between forage

of mule deer and forage nutritional

It is paradoxical

values.

that despite these findings, wild and domest~c

ruminants have been shown repeatedly to select diets of higher nutritional quality than the mean quality of herbage on offer (for review see
Klein 1970, Longhurst et al. 1968, Arnold 1964, White 1978). This
apparent paradox can be explained in terms of the unit of diet selection.
If animals do not discriminate

between plant species in choosing

their diets but instead between plant parts or groups of species, then
analysis of diet selection on a species basis will fail to accurately

�-266-

predict preference.

For example, protein content of a group of species

may be related to preference
preference

for that group, while within the group, no

is exerted for individual

protein content.

Likewise,

species on the basis of their

there may be good correlation

between pref~r-

ence for plant parts and their chemical constituents,

a correlation

which remains undiscovered

studied is plant

species.

when the unit of selection

This is the case because there is often greater variation

in

nutritive value between plant parts of the same species than between
whole plants of different

species (Arnold 1960, Van Dyne and Heady 1965,

Cowan et al. 1970, Short et al. 1972, Bailey 1967, Aldous 1945).
though there is widespread

Al-

evidence that ruminants are selective for

plant parts (Hirst 1975, Jarman 1974, Sinclair

1974c, Arnold 1960, Neyer

et al. 1957, Bluser et al. 1960), there has been only limited study of
mechanisms

of preference

for groups of species (Kossak 1976).

Methods of Diet Analysis
The quality of a herbivore's
tween the botanical
and digestibility

composition

tions and measurements

phenomena

be-

of its diet and the chemical composition

of the plant species and plant parts consumed.

of these interacting

i) Observation

diet results from the interaction

Study.

requires methods for food habits observa-

of forage quality.

of Diet Choices of Tame Animals

Methods for study of diets of large herbivores
sively reviewed elsewhere
Ward 1970, Van Dyne 1968).

have been exten-

(Schwartz 1977, Medin 1970, Free et al. 1971,
Rather than duplicate

cussion here will focus on advantages

those efforts, dis-

and limitations

of the technique

employed in this study, bite counts with tame animals, in relation to
other methods.

�-267-

Observation
cant advantages
its applicability

of diet choices of tame animals offers several signifiover other food habits methods.
to controlled

Most advantageous

is

sampling designs (Wallmo and Neff 1970).

That is, diets can be observed at a variety of locations and at various
times since when and where the animals feeds is controlled by the investigator.

For example, tame animals are the only means to quantify diet

selection within plant communities when a variety of habitats are used
by the species being investigated.
cal composition

It is impossible

to estimate botani-

of diets within a vegetation

type based on fecal samples

collected there since locations of herbivores

when consuming and defeca-

ting a given meal may be different
inherent in interpretation
examination

(Collins 1979).

Similar problems are

of rumen samples (Medin 1970).

of feeding sites and distant observation

Although

of feeding activity

of wild animals provides knowledge of where a plant was eaten, neither
method is particularly

useful when animals are nocturnal

or seclusive

(Wallmo et al. 1973).
Tame animals facilitate accurate description
tion of diets.

As previously

discussed,

rumiants can be selective for

plant parts as well as plant species; variation
result in substantial

differences

of chemical composi-

in parts eaten can

in chemical composition

(Arnold 1960, Bailey 1967).

Therefore,

position alone is inadequate

for sampling diet quality.

tame animals allows description

knowledge

of diets

of botanical

com-

Observation

of

of plant species and plant parts eaten,

and thereby permits collection of forage samples which more accurately
reflect chemical composition
Certain assumptions
criticized,

however.

of forages consumed.

required by the tame animal technique have been

Obviously,

it must be assumed that artificial

�-268-

environments

do not bias natural diets selected by tame animals compared

to their wild counterparts.

There is conflicting

evidence on the valid-

ity of this assumption.
There are data which suggest that early experience may affect die~
choice of ruminants.

Young animals may learn preferences

(Edwards 1976) or early nutritional
selection

experience may influence latter diet

(Arnold and Miller 1977, Langlands

1968). However, Bergerud

from adults

1969, Longhurst et al.

(1972) observed that tame bottle-reared

caribou

removed from the dam at I day of age had similar diet preferences
wild animals as adults.

to

Schwartz and Nagy (1973) used microhistological

analysis of feces to compare diets of tame and wild pronghorn

antelope

and found that with the exception of one forage species, variation
diet choices was greater between wild pronghorn

in

than between wild and

tame animals.
It has also been suggested that nutritional
during grazing experiments
1968).

This hypothesis

status of the animal

may influence diet choice (Longhurst et al.

has been investigated

by a variety of workers

(Schwartz and Nagy 1973, Reglin et-al. 1976, Bartman et al. 1978) who
found no effect of artificial

supplementation

on the composition

of tame

animal diets.
Although the possibility

remains that diets of tame ungulates may

not exactly reflect composition

of diets of wild ungulates,

sufficient evidence to suggest that this inaccuracy
however, is a problem not unique to observation
histological

there is

is small.

Bias,

of tame animals.

Micro-

analysis of feces has been shown to be discrepant with

estimates of diet composition

measured by other methods

(Alexander et

al. 1978, Anthony and Smith 1974, Smith and Shandruk 1979, Johnson et

�-269-

al. 1978, Hoover 1971, Ward 1970, Adams et al. 1962, Vara et al. 1978,
Slater and Jones 1971); rumen samples overestimate

forage species which

have slow passage rates (Bergerud and Russel 1964, Medin 1970), and
feeding site observations
shrubs and overestimate
1973).

of wild animals will probably underestimate
grasses in wild herbivore

Given these drawbacks of other techniques,

tame animals previously

diets (Wallmo et al.
and the advantages

of

discussed, potential bias in the tame animal

technique seems a trivial problem.
ii) Assessment

of Dietary Preference

A preferred
proportionately
environment

food species has been defined as one which occurs
more frequently

(Petrides 1975).

in the diet of an animals than in the

This definition

has given rise to a pleth-

ora of indexes of dietary preference which attempt to weight the proportions of items in the diet by some measure of their availability.
(Chamrad and Box 1968, Jacobs 1974, Heady 1964, Cock 1978, Petrides
1975, Chesson 1978).

In their rudimentary

form, these indexes can

usually be reduced to a simple fraction where the percentage

of an item

in the diet is divided by the biomass, density, or frequency of the item
in the habitat.
contribute

For several reasons, .such arithmetic manipulations

little to our understanding

First, it is reasonable
some foods in an exploratory

of diet choice.

to assume that any herbivore will select
way (Pyke et al. 1978).

constantly test the food potentials

of its environment

basis of past tests (Foreland and Janzen 1974).
the animal consumes an exceedingly

That is, it will
and act on the

If during that testing

rare species, then that species would

be calculated as being "highly preferredll

simply because it was rare.

�-270-

Second, if the unit of diet selection is not plant species; that
is, if a herbivore does not discriminate
basis, then preference

between plants on a species

indexes will shed little light on diet choice.

Third, common species in the environment
able to the grazing animal despite differences
biomass

(Westoby 1974).

tend to be equally avail~
in their density or

This is the case because those species are so

abundant that searching time is zero for all of them.
their superabundance,

However,

more rare species would be calculated

despite

as

"preferred".
Finally, there are acute problems with comparison
indexes measurement
differences

in one area to areas where there are even slight

in mosaics of forage.

out that preference
species.

Westoby

(1974) and Heady (1964) point

for one species is affected by availability

Thus, a plant that is highly preferred

unpalatable

of preference

of other

when surrounded by

neighbors might be rejected when its associated plants were

eaten more often.

It follows that preference

is only a relative charac-

teristic of a plant; this fact strongly limits the utility of preference
indexes.
Although the preference
attempts to measure,

index has shortcomings,

"selectively",

the diet selection process.

a real attribute

of

are being selective,

(Ellis et al.

feeding occurs when foods are consumed in proportion

with their availability.
diet selectivity,

it

Animals which consume foods more frequently

than they occur in the environment
1976); nonselective

is doubtlessly

the phenomena

Although

this paper will address problems of

diet data will be presented

rather than as preference
indexes discussed above.

as simple perc~ntages

indexes in light of the limitation

of those

�-271-

iii) Measurement

of Forage Quality

Evaluation

of the nutritional

volves measurement

quality of forages for ruminants

of forage fiber characteristics,

in-

protein content, and

digestibility.
The Weende system of proximate

analysis

(Henneberg and Stohmann

1960) fractions plant dryrnatter into crude fiber and nitrogen free extract (NFE); crude fiber being less digestible
ies have shown this fiber breakdown

than NFE.

Several stud-

to be poorly correlated with digest-

ibility (reviewed by Van Soest 1966, 1968); often the crude fiber portion
was more digestible

than NFE (Ely and Moore 1959).

An alternative

system of fiber analysis was developed which used acid and neutral detergent to break down plant dryrnatter into components of uniform composition and digestibility
Soest 1970, Fonnesbeck

(Van Soest 1966, 1967, 1968, Goering and Van

and Harris 1970, Robbins et al. 1975).

This

scheme partitions

plant material into cell solubles and cell wall and

then fractionates

the cell wall into its principle

cellulose, hemicellulose,

and lignin.

fiber components,

Cell solubles are completely

digested enzymatically;

microbial digestion

extent of lignification

of cellulose and hemicellulose

Although

fiber characteristics

bility and rate of digestion

of cell wall depends on the
(Van Soest 1973).

have been correlated with forage digesti(Denium and Van Soest 1969, Smith et al.

1971, McLeod and Minson 1972, Minson 1971, Wilson et al. 1971), rate of
passage

(Smith et al. 1960), and voluntary

et al.(1975:68)

cautioned that

ships between feed constituents
as biologically
vestigated

significant

tI

•••

intake (Mertins 1973), Robbins

statistically

and digestibility

or meaningful

are adequately understood

significant

relation-

should not be viewed

unless the relationships

and causatively

related."

in-

Thus,

�-272-

evaluation

of forage quality on the basis of fiber characteristics

be strongly coupled with knowledge of the physiological
volved in ruminant digestion and assimilation

processes

(see Milchunas

must
in-

et al. 1978

for an example of such merger).
Protein is often in short supply in diets' of grazing ruminants
(White 1978, Klein 1962, 1965, Holter et al. 1977, Elliot and Topps
1964).

Microbial protein synthesized

essential
1976).

in the rumen contains all the

amino acids and has a biological value of about 80% (Hendricks

Although amino acid composition

microbial

protein synthesis

of nitrogen

of dietary protein can affect

(Hume 1970), on low protein diets the amount

in forage is of much greater significance

to the ruminant

than is the forage amino acid mix (Hume et al. 1970).
measurement
provides

of nitrogen content by Kjeldahl methods

Consequently,

(A.O.A.C. 1965)

a good measure of the ability of the forage to contribute to

the animals's protein status.

Conversion

crude protein values by mUltiplying
proportion

of forage nitrogen content to

Kelldahl N times the stociometric

of nitrogen in protein molecules

(6.25) achieves no improve-

ment on the original measure, but is a ubiquitous

practice and has

become the standard index of forage quality with respect to protein.
Robbins et al. (1974f

and Sullivan

(1962) found crude protein con-

tent of forage was correlated with digestible protein.

Based on their

work, it can be assumed that true protein digestibility

of a variety of

forages is greater than 90%.
Measurement
previously
completeness
predicting

of forage dry matter digestibility

discussed

forage chemical characteristics

of forage digestion.
forage digestibility

integrates all the
in a measure of

In vitro laboratory methods for

allow much more rapid analysis than was

�-273-

previously

possible with conventional

for review of development

in vivo trials (see Johnson

of the in vitro technique).

In vitro determi-

nations have been shown to be correlated with in vivo predictions
matter digestibility

(Tilley and Terry 1963, Milchunas

Robbins et al. 1976, Ruggiero and Whelan 1976).
is that dry matter digestion
in approximately

the percentage
animal.

What is more important

is related to digestible

et al. 1971).

of dry

et al. 1978,

energy in forages

a 1 to 1.ratio (Robbins 1975, Milchunas

Moir 1961, Rittenhouse

1966

et al. 1978,

Thus, IVDDM can be used to predict

of energy in the forage which can be digested by the

However,

in vitro values can be affected by innoculum

sources

(Robbins et al. 1975 Palmer et al. 1976, Ward 1971), inoculum handling
time (Schwartz and Nagy 1972) and specific
by Scales 1972).

Consequently,

(reviewed

despite the accuracy of in vitro values

compared with in vivo determinations,
technique

in vitro procedures

the precision

requires some caution in application

of the in vitro

to range situations

(Hi.Lchunas and Baker, submitted).
Elk Winter Food Habits
Diets of elk during winter have been extensively
Rocky Mountain Region
studies:

(Table 2).

There is one compelling

elk diets during winter were dominated

forbs were universally

studied in the

minor diet contributors.

trend in these

by grass and browse;
In general, browse

dominated diets in areas where snow covered the grass sinusia; grasses
were more frequently

consumed in relatively

however, that a large number of previous

snow free areas.

food habits studies have relied

on analysis of rumen samples a measure of diet composition.
fied browses has shorter rumina 1 retention
gested by Milchunas

Note,

If ligni-

times than grasses as sug-

et al. (1978), then rumen samples would tend to

�-274-

overestimate
prevalent

the dietary contribution

of grasses.

food habit technique, examination

Further,

of feeding sites, has also

been shown to inflate estimates of grass consumption
1973).

Consequently,

(Wallmo et al.

the widely held belief that elk are "grazers" may

result, in part, from bias in previous

Table 2.

the other

food habits methods.

Summary of estimates of forage class composition
diets in the Rocky Hountain region.

Area
(Authority)

Diet Composition C%)
Grass Forbs Shrubs

of elk winter

Food Habits
Technique

Hontana
Singer 1979
Cons tan 1972
Knight 1970
Nackie 1970
Schallenberger 1965
Knight 1970
Kirsch 1962
Rouse 1957
Norris and Schwartz 1957

12
60

88

84
100
37
14

o

o

21

42

1

88

7

90
100

o

85
5
3

7422
79

6
10
7

57
14

fecal samples
rumen samples
feeding sites

65

8

27

rumen samples

44

24

93

32
7

rumen samples
rumen samples

97

3

direct observation

21

rumen samples

7

30
10

feeding sites
feeding sites
feeding sites
rumen samples
feeding sites
rumen samples
rumen samples
feeding sites
rumen samples

10
6

o

Colorado
Hansen and Reid 1975
Boyd 1970
Stevens 1969

20

Idaho
Denio
Wyoming
Anderson 1953
Murie 1951
Canada
Cowan 1947
New Hexi co
Lang 1958

77

2

�-275-

Chapter IV
STUDY AREA
Investigations

were conducted in Rocky Mountain National Park on

the eastern slope of the Continental Divide between 2500 and 2800 m in
elevation

8 km west of the town of Estes Park, Colorado

(R73W, T4W)

(Fig. 1).

The total area of elk winter range within the park boundary

is about 6000 ha.
Glacial moraines extending eastward from the front range mountains
separate the wintering

area into 4 parallel valleys.

in these valleys and form the headwaters

Streams flow east

of the Big Thompson River.

Soils are derived from decomposed granite and mica-shists.
Vegetation
(1967).

is typical of the upper montane climax region (Marr

Lodgepole pine (Pinus contorta) and douglas fir (Pseudotsuga

menziesii)

dominate the northfacing

ponderosa)

and big-sagebrush

prevalent

on southern aspects.

Ponderosa pine (Pinus

(Artemisia tridentata)
Vegetation

strongly influenced by differential
cultivation.

slopes.

communities

on the valley floors is

moisture availability

and past

Eight habitat types (described in Table 3) are used by elk

for feeding during the winter and all were investigated
Approximately
pers. com.).

are

2,000 elk occupy this area during the winter (Stevens

Other ungulates,

including mule deer (Odicoileus hemionus)

and bighorn sheep (avis canadensis),
Winter weather is usually mild.
January mean minimum temperature
maximum temperature

in this study.

are also present.
During both years of the study,

at Estes Park was 9 C, while mean

was 8 C during 1976 and 1977 and 2 C the following

�-276-

year (National Oceanic and Atmospheric Administration

1977, 1978).

Because of strong westerly winds (up to 75 kph) and frequent abovefreezing temperatures, protracted snow accumulation is rare .

Rocky~
Mountain
National
Park

• Fort Collins

U
o Denver

• Grand Junction

e Pueblo

COLORADO

Figure 2.

Geographic location of Rocky Mountain National Park in
Colorado.

�-277-

Table 3.

Site characteristics and principle plant species of upper
montane vegetation types in Rocky Mountain National Park,
Colorado.

Vegetation Type

General Site
Characteristics

Principal Plant Species

Grassland

Patchily distributed
grass communities on
open xeric sites

Muhlenbergia montana
Stipa comata
BOUteloua gracilis
Danthonia parryi

Ponderosa pine-shrub

Large open canopy
parklands on south
exposed slopes with
shallow, rocky soils

Danthonia parryi
Muhlenbergia montana
Bouteloua gracilis
Carex heliophila
Ribes spp.
PUrShia tridentata

Sagebrush

Open canopies of big
sagebrush (Artemisia
tridentata) on south
slopes with deep,
well-drained soils

Muhlenbergia montana
Carex heliophia
Agropyron spicatum
Stipa comata
Artemisia tridentata
Purshia tridentata

Willow

Found in wet areas
along stream courses;
closed canopy of
willow (Salix spp.)

Calamogrostis
Carex spp.
Salix spp.

Aspen

Small, patchily
distributed communities.ort mesic to wet
sites

Calamogrostis canadensis
Phleum pratense
Poa pratensis
populus tremuloides

Mesic meadow

Large areas formerly
cultivated; moderately drained soils

Poa pratensis
Phleum pratense
Juncus balticus
Muhlenbergia richardsoni

Wet shrub meadow

Open-canopied shrub
meadow on hygric sites

Calamagrostis canadensis
Carex spp.
PfiIeUm pratense
Glyceria grandis
Salix spp.
BetUla glandulosa
Alnus tenuifolia

canadensis

�-280-

ensuring

independent observations.

As each animal grazed, one observer

recorded

(on a tape recorder) the plant community in which the animal

was feeding, the number of bites of each plant species eaten, and a
description

of plant parts selected.

Distance

from observers

varied from 3 to 0.1 M, depending on ease of identifying

to anima~s

plants con-

sumed, either from the plant being grazed or, when it was entirely
eaten, from adjacent plants of the same species.
Forage samples were collected for all species contributing
more of total bites observed during each grazing trial.
plant material,

2% or

Fifty grams of

taken from no fewer than five locations, was composited

into plastic bags and frozen.
ing an additional

Bite weight was estimated by hand-pluck-

25 sample "bites" of these species, drying the samples

at 100 C for 24 hr, weighing the entire sample to the nearest ±0.001 g,
and dividing the total weight by 25.
Between grazing trials, the elk were kept in Fort Collins, Colorado,
and were provided with alfalfa hay ad libitum.
animals were maintained
transported

During the trials,

in a O.l-ha enclosure on the study area and were

each day to and from release points in a modified double-

axle, 2-horse trailer.

Food intake was limited to the forage ingested

in the field, 20 g of grain used as a reinforcer for loading"into
trailer

(Hobbs and Baker 1979), and approximately

Artificial

supplementation

the

500 g alfalfa hay.

of natural diets and restricted

grazing time

have been shown to have no effect on diet choices of tame pronghorn
antelope

(Antilocapra

americana)

22, Regelin et al. 1976).

and mule deer (Schwartz and Nagy 1973:

�-281-

Forage Quality
Crude protein

(Kjeldahl N x 6.25), ash, and dry matter of forage

samples were determined by procedures
Cell wall constituents

described by A.O.A.C.

(CWC) , acid detergent

(1965).

fiber (ADF) , and acid de-.

tergent lignin were determined by the procedures

summarized by Colburn

~nd Evans (1967) and Bailey and Ulyatt (1970).

Analyses were performed

sequentially

by Robbins et al. (1975).

on single samples, as recommended

In vitro digestible

dry matter (IVDDM) was determined by techniques

described by Tilley and Terry (1963) and Pearson
trial was conducted for each year's samples.

(1970).

One in vitro

Rumen innocula were ob-

tained from wild elk collected from the study area.

Rumen fluid was

collected from 5 animals (3 adults, 2 calves) in March 1977 and from 2
animals (1 adult, 1 calf) in March 1978.· Rumen contents were transported in prewarmed
tory.

insulated chests from the study area to the labora-

Elapsed time from animal death until innoculum was buffered and

in tubes was approximately

1.5 hr.

Before innoculation,

equal portions

of rumen fluid from each animal were mixed to reduce variation

in fluid

quality.
Because the rumen fluid obtained in 1978 was extremely viscous and
could not be vacuum filtered, undigested
the fluid by centrifugation.

These unconventional

biased estimates of digestibility.

Digestion

forages were lower in 1978 (Year 2) (P
this bias was compensated

substrate was separated from
procedures

eoefficients

apparently

of standard

= 0.001) than in 1977 (Year 1).

for by regressing

standard coefficients

Year 2 against those of Year 1.

The resulting

plied to Year 2 grasses was IVDDM

= 0.29 + 0.68 IVDDM(R2 = 0.84, P =

0.004, n

correction

of

equation ap-

= 8); the equation for browse and perennial forbs was IVDDM =

0.04 + 0.95 IVDDM(R

2

= 0.99, P = 0.003, n = 5).

�-282-

Diet Data Analyses
Botanical

composition

total observed bites.

of elk diets was calculated as percentages

Chemical composition

of diets was calculated

the sum of forage chemical values times weighted diet percentages.
percentages

were weighted by multiplying

bite weight and normalizing

of
as
Diet

bite frequency times estimated

to sum to 100 (Schwartz et al. 1977:162).

Because data expressed

as percentages

are frequently not distrib-

uted normally and therefore are not amenable to routine statistical
analyses,

transformations

normal distribtion.

(such as arc sin) may be used to obtain a

Diet botanical

compositions

centages, but arc sin transformation

were expressed as per-

did not alter estimates of the

variance beyond the third decimal place and biased estimates of means.
Butchner and Kemp (1974) found no difference
of variance.using

transformed

unequal number of observations
diet percentages

in power between analysis

and untransformed

data, and no effect of

among percentages.

For these reasons,

were treated as normal variates

and were not trans-

formed.
Dietary differences

among animals, years, and habitat types were

analyzed by a factorial

analysis of variance

for a repeated measures

design, repeating" over 2 years with 5 animals as replicates and 8 habitat types as treatments.

Tukey's Q simultaneous

comparisons were used

to detect differences

between mean responses.

chemical constituents

between years were analyzed with paired t-tests.

Seasonal variations

in chemical content of diets and forages were cal-

culated with simple linear and polynomial
indicated,

differences

in mean responses.

and associated

For discussion

and Draper and Smith (1966).

Differences

regressions.

probabilities

in forage

Unless otherwise

refer to differences

of the above tests, see Hicks (1973)

�-283-

Estimation

of Herbage Biomass

Standing crops of herbacious biomass and current production

of

shrubs were estimated at the end of the growing season during 1976 and
1977.

Sample dates were August 20-September

for shrubs.

20 for herbs and Nov. 1-10

Sampling dates were delayed for shrubs to allow for leaf-

fall.
Sample units were stands representing
ducing vegetation

types (Table 1).

each of the major forage pro-

Stands were chosen as follows.

itat maps, prepared by Dave Stevens, Park Biologist,
eate the vegetation

types.

x 100 m) were subjectively

For each vegetation

Hab-

were used to delin-

type, 8 macro-plots

(100

identified and 4 of these areas were randomly

selected as stands to be sampled.

Stand boundaries

were paced off

following a compass bearing and steel stakes were placed in the corners
of each of these stands to permanently
Each stand was subsampled

mark their location.

for herbacious

lar plots (1 x \ m); shrub production

species with 30 rectangu-

with each stand was measured

in 10

2m2 circular plots.

Subsample plots were allocated according

following procedure.

Ten stakes were placed at random intervals along

one border of each stand.

Three \m2 herbacious

plots and one 2m2 shrub

plot were randomly located along 100 m transects originating
stake and running perpendicular

to the

to the stand border.

at each

All random inter-

vals were specified using a random number table and were delimitated

in

to field by pacing the specified distances.
Summarizing

the sampling design, 8 community types were sampled in

4 stands per type, each stand being subsampled with 30 herbacious
and 10 shrub plots.

Clip and weigh methods

plots

(Swift and French 1972) were

�-284-

used to measure plant biomass in each subsample plot.

Herbacious

material was ground level clipped and separated by species for the
dominant grasses and lumped into forage classes for minor grasses and
forbs.

Current growth of shrubs was clipped between ground level and

2.5 m high, the vertical dimension being defined by a steel rod placed
perpendicular

to the plot frame.

Dominant shrub species were separated;
o

minor species were composited.

Clipped samples were dried at 100 C for

48 hr and weighed to the nearest 0.1 g.
Biomass of each species or forage category within a stand was calculated as the me~n dryweights across all subsample plots within the
stand.

Biomass of a forage category in a vegetation type was then cal-

culated as the mean of 4 stand subsample means.

Standard errors were

calculated based on a pooled estimate of the variance across 4 stands.
Differences

in means were analyzed by a nested factorial analysis of

variance for a repeated measures design, repeating over 2 years with 8
habitat types as treatments and 4 stands as replicates.
Simulation Modeling of Nutritional

Status

The ruminant simulation model used in analysis of elk nutritional
status was developed by D.-M. Swift at the Natural Resource Ecology
Laboratory,

Colorado State University.

For a detailed description of

the structure of the model, calculation of flows, and an example implementation, see chapters 5 and 6 of Ellis and Parton (1978).

A brief

discussion of the essential features of the model will be presented
here.
The ruminant model is a generalized mathematical

representation

of

the flows of nitrogen and energy through an individual runinant animal

�-285-

(Fig. 3).

It is structured so that minor changes in parameter specifi-

cations allow the model to represent the nitrogen and energy metabolism
for any ruminant species and for any sex and age class.
Model objectives include prediction of 1) rates of forage consumption, 2) rates of digestion and metabolism
partitioning

of energy and nitrogen, 3)

of energy and nitrogen within the body, and 4) losses of

energy as nitrogen from the body.

Objectives

3 and 4 allow estimation

of changes in the animal's total weight, and fat and lean body mass.
These weight fluxes are used in this study as indicators of elk nutritional status.
The model is a difference equation model operating at one day time
steps.

That is, values of flows and state variables are calculated on a

per day basis.

Parameters specified on the model to represent an elk

during winter are outlined in Table 4.

Minimum and maximum daily tem-

peratures were taken from the National Oceanic and Atmospheric

Adminis-

tration records (1976, 1977, 1978) for Estes Park, Colorado.
Principal input data used to drive the model were the nitrogen content and IVDDM of diets.
point estimates.

These values were entered into the model as

For modelling exercises it was assumed that differ-

ences in diet quality, regardless of their statistical

significance,

were real differences.
This assumption was made for two reasons.
integration of two variates
dicted responses

First, the model allowed

(diet IVDDM and nitrogen) in a set of pre-

(weight change patterns). This synthesis was not pos-

sible in the statistical

comparisons which treated diet quality measures

as independent observations.

�-286-

Rumen
Microbial
Protein X21

Figure 3.

Structure of generalized
gen metabolism.

model of ruminant energy and nitro-

�-287-

Parameter spe ci.fi ca t i ons for the ruminant simulation model
runs representing elk on upper montane winter range, Colorado.

Table 4.

Parameter

Value

Source

Length of winter

Nov. 10-March 25

Specified to coincide
with diet sampling
periods

Lower critical temperature

-10 C

Gates and Hudson
(1979)

Basil metabolic

79 x kgO.75 (kcal)

rate (BMR)

Activity costs

BMR x 0.55

Methane production

0.52 x gross energy
intake

Forage dry matter to
gross energy conversion

4.3

Maloiy et al. (1970)
Craighead et al.
(1973) and Gates and
Hudson (1978)

Rumen volume

estimates based on
Nagy and Reglin
(1975 )

Adult body weight

217 kg

Dean et a1. (1976)

Calf body weight

87 kg·

Dean et a1. (1976)

Adult fat weight

33 kg

Flook (1970)

Calf fat weight

10 kg

Flook (1970)

Growth rate

w

= 372_262~-·00086t

t

= years,

w

Second, this assumption
variation

= weight in kg

allowed examination

in diet quality on animal condition.

tigation of the biological

McEwan (1975)

significance

ever, because of the measurement

of effects of small
Thus, it allows inves-

of observed differences.

How-

error implicit in diet quality input

�-288-

data, differences
caution.

in model output among habitats must be considered with

That is, if there were no statistical

quality measures among habitats,
condition,

differences

then differences

in diet

in predicted

animal

no matter how large, would not be repeatable given several

replications

of the diet sampling.

Two sets of model runs were made.
was parameterized

In the first series, the model

to represent an adult and a calf elk during both years

of the study and was driven with across habitat estimates of diet quality by month.

These model runs provided my best estimate of elk nutri-

tional status in relation to the entire winter range.
In the second series of model runs, differences
status among different

habitats were investigated.

in elk nutritional
The model was param-

eterized for a calf during year 1 and an adult during year 2 and driven
with across month estimates

of diet quality by habitat.

Thus, for each

year and in each habitat the model was used to predict overwinter

changes

in animal condition.
Carrying Capacity Estimation
Following
estimated

the example of Wallmo et al. 1977, carrying capacity was

on the basis of forage quantity by calculating

the number of

elk required per unit time to remove ~ the total standing crop biomass
of winter f orage :

carrying capacity

= total herbage biomas . ha-1 • ~ = elk • day • ha -1
elk forage intake • day

-1

Forage intake estimates were based on model predictions
take when the model parameters
(Table 4).

were specified

of voluntary

in-

to represent an adult elk

This estimate of carrying capacity does not include any

�-289-

prediction

of the animals nutritional

status.

Therefore,

also following

Wal1mo et a1. 1977, all carrying capacity estimates based on forage
quantity are presented

in conjunction

with estimates of animal condition

based on measured diet quality and predicted

food requirements.

however, that these condition estimates would be absolutely
given the predicted

Note,

accurate

elk densities only if a 50% removal of total biomass

did not reduce elk diet quality below the measured values.

�-290-

Chapter VI
RESULTS
Herbage Biomass
Biomass of total standing crops, forage classes and dominant species was estimated within 20% of the mean with 80% confidence.

Individ-

ual species, other than the dominants, were estimated with considerably
less precision.

Species data are presented in Baker and Hobbs (1979)

and Hobbs and Baker (1978); results presented here will be limited to
forage classes and total biomass.
There was extensive variation in total standing crop biomass between years and among habitats (Fig. 4).

During year 1, total herbage

biomass ranged from a high of 216.6 g • m-2 in willow to a low of 72.4 g
• m-2 in grassland; the following year the high and low standing crops
were 189.8 g • m-2 in willow and 67.5 g • m-2 in grassland.
years, total biomass was significantly
grassland, ponderosa pine-shrub,

greater in willow than in aspen,

or mesic meadow (P &lt; 0.05, Fig. 4).

Standing crops declined significantly
(year effect P

in all habitats between years

= 0.006, year x habitat interaction P = 0.6).

this decline, species composition was not substantially
vegetation

During both

Despite

changed in most

types (Appendix 2).

Differences

in total biomass resulted largely from variation in

grass standing crops, since grasses contributed
biomass in all communities.
and mesic vegetation

the major portion of

Biomass of grasses was greater in the wet

types (wet meadow, mesic meadow, willow, and wet

shrub meadow) than in aspen and the more xeric communities

(sagebrush,

�-291-

WILW
SAGE

225

WGSM

WSHM

....--..

(\j
I

E 175

MGSM

N

0'1

(f)
(f)

125

&lt;!

m

75

25

Figure 4.

GRSL

PP/SH

2

0

ASPN

N

Total aboveground biomass production at the end of the growing season during 1976 and 1977 in upper montane habitats,
Colorado. Mean values are indicated by X; non-overlapping
vertical lines indicate significant difference at P=.OS.
Solid lines are for 1976; dashed for 1977. Abreviations
are: WILW = willow, PPSH = ponderosa pine shrub, WSffrl= wet
shrub meadow, ASPN = aspen, SAGE = sagebrush, WGSM = wet
meadow, GRSL = grassland, MGS~l = mesic meadow.

�-292-

grassland, and ponderosa pine-shrub)

(Fig. 5)

Biomass of grasses de-

clined in all habitats the second year (year effect P

= 0.003, year x

habitat interaction P = 0.67).
Current growth of shrubs was greatest in willow and sagebrush,
intermediate

in ponderosa pine-shrub, wet shrub meadow, and aspen, and

least in wet meadow, grassland, and mesic meadow (Fig. 6). There was no
between year difference in shrub production in any habitat (year effect
P

= 0.18).
Forb biomass was highly variable between years, increasing in some

habitats (willow, aspen, wet meadow and mesic meadow), decreasing in
others (ponderosa pine-shrup,

grassland, sagebrush) and remaining con-

stant in a single vegetation type (wet shrub meadow) (Fig. 7).

Thus,

annual variation was sufficient to rearrange the habitat differences

in

forb biomass established by the first years data (year x habitat interaction P

= 0.06).

Diet Botanical Composition
Browse and grasses dominated winter diets selected by experimental
animals (Table 5).
mesic vegetation

The most frequently consumed grasses in wet and

types were bluegrass

(Poa pratensis),

pratense), Canada reedgrass (Calamagrostic

timothy (Phleum

canadensis), and sedges

(Carex spp.). Important browses included aspen leaves (Populus
tremuloides),

leaves and stems of willow (Salix spp.), and stems of

shrubby cinquefoil

(Potentilla fruiticosa).

Principal species eaten in

xeric communities were mountain muhly (Muhlenbergia montana), blue grama
(Bouteloua gracilis), needle and thread (Stipa comata), and bitterbrush
(Purshia tridentata).

Although elk feeding on fallen leaves has not

been widely reported, I observed that frequent consumption of fallen

�-293-

WILW
WGSM
MGSM

WSHM

••......•.
N
I

E
ASPN

(f)
(f)

«

100

~

SAGE

o
m

GRSL

PP/SH
50

Figure 5.

Biomass of grass at the end of the growing season during 1976
and 1977 in upper montane habitats, Colorado.
Mean values are
indicated by X; non-overlapping vertical lines indicate significant difference at P=.05. Solid lines are for 1976,
dashed for 1977. Abreviations are: WILW = willow, PPSH =
ponderosa pine shrub, WSHM = wet shrub meadow, ASPN = Aspen,
SAGE = sagebrush, WGSM = wet meadow, GRSL = grassland, MGSN =
mesic meadow.

�-294-

25

WILW

SAGE

I
I

I

t
I

t
1
20
(\J
I

E
0\

WSHM
ASPN

15

I

(j)
(j)

«
~

0
CD

PP/SH
10

WGSM
GRSL

I:

L

Figure 6.

Biomass of current annual growth of shrubs during November of
1976 and 1977 in upper montane habitats, Colorado. Mean
values are indicated by X; non-overlapping vertical lines
indicate significant difference at P=.OS. Solid lines are
for 1976; dashed for 1977. abreviations are: WILW = willow,
PP/SH = ponderosa pine-shrub, WSHJl = west shrub meadow, ASPN
= aspen, SAGE = sagebrush, WGSM
wet meadow, GRSL
grassland, MGSM
mesic meadow.

=

=

=

�-295-

ASPN

GRSL.

I

SAGE

....--..
N

WGSM
1
1

I

I

E
01 20

PP/SH

WSHM

(J)
(J)

-r

~
0

15

CD

WILW
1

I

MGSM
I

I
I

I

Figure 7.

Biomass of forbs at the end of the growing season during 1976
and 1977 in upper montane habitats, in Rocky Mountain National Park, Colorado.
Mean values are indicated by X; non-overlapping vertical lines indicate significant difference at
P=.OS. Solid lines are for 1976; dashed for 1977. Abreviations are: WILW = willow, PPjSH = ponderosa pine-shrub,
WSill1= wet shrub meadow, ASPN = aspen, SAGE = sagebrush, WGSH
= wet meadow, GRSL = grassland, MGSN = mesic meadow.

�Table 5.

Mean percentage

of observed

Novembe r-Ha rch 1976-1977

bites of plant species

and 1977-1978

summarized

in diets of 5 elk on upper montane winter
by habitat

Habitat

Willow

Het
meadow

Nesic
meadow

Grassland

Taxa~/

Year~j

X%

inermis

1977
1978

5
tr

1977
1978

0
0

1977
1978

16
10

4
19

1
8

8
7

2
1

0

1

1977
1978

6
8

3
5

21
16

4
5

5
17

2
1

1
tr

SE

X%

SE

X%

11
3

5
1

12
4

SE

range, in Colorado,

during

type.~/

X%

SE

Type

Sagebrush

Ponderosa
pine-shrub

Aspen

Wet
shrub

Across
all
types

---

---

X%

SE

X%

SE

X%

SE

X%

SE

X%

SE

1
0

0.4
0

2
2

1
1

2
tr

0.3

2
1

0.4
0.4

4
1

0.4
0.2

5

1
1

1
6

0.3
2

0
tr

1
2

0.2
0.4

----

Grasses
Rromus

~outeloua

gracilis

Calamagrostis

~rex

canadensis

spp.

3

0
0

3
1

tr
0

tr

5
4

1
8

0.4

3

0
0

0
0

0
0

2
14

1
1

11
10

1
1

4
7

tr
tr

t.r
tr

5
3

2
1

13
17

1

6

8

1

I
N
\D

JlIncus balticus

1977
1978

tr
tr

1977
1978

0
0

1977
1978

tr

Phleum pratense

1977
1978

8
3

2
1

7
7

Poa E_!:"atensis

1977
1978

31

4
4

26
12

1977
1978

0
0

1977
1978

55
3S

~Iuhlenber~

Muhlenbergia

montana

.richardsoni

Slipa comata
All graminoids~/

11

1
4
9

0.4
1

0.4
6

1
1

0.5
0.4

tr
0

18
35

2
2

34
52

0.4

tr

0
0

tr
0

3
4

1
1

5

19
2

6

1

44
16

5
6

1
1

4
2

1
1

0
0

4
4

61
61

4
4

59
41

1

2
3

15
18

10
10

tr

9
3

38

12
6

4
tr

1

14
S

2
1

7
4

48
50

3
4

69
61

83
69

3
S

2

0
1

1
5

8
8

0.4

2

tr
0

1
3

71
64

2

tr
tr

tr
tr

tr
tr

1

0
0

tr
tr

11

tr
2

tr
0

0
0

0
0

0
0
6
1

1
7

7
4

0

24
41

4
4

I

9
14

2
1

0.4

tr
1

0.4

8
4

3
1

5
4

22
6

3
1

25

0
0
S
3

o-

tr
1

S9
45

7

2
1

3

2
4
4

61
50

3
3

�Table 5.

Continued
Habitat Type

Willow

Wet
meadow

Nesic
meadow

Grassland

SE

X%

0.4

Sagebrush

Ponderosa
pine-shrub

X%

X%

Aspen

Wet
shrub

Across
all
types

X%

X%

X%

Taxa!?/

Year~j

X%

SE

X% .SE

X%

Alnus tenuifolia
(leaves)

1977
1978

2

0.4
0.4

6
tr

tr

o
o

o
o

o
o

o
o

2
tr

tr

Populus treOluloides

Deaves)

1977
1978

tr

o
o

o
o

tr
tr

o
o

tr
tr

32 4
36 3

o
o

7

1

5

0.4

Prunus virglnlanu
Heaves)

1977
1978

o

o

o

o
o

15 3
15 3

o
o

o

o
o

o
o

Salix spp.
(leaves)

1977
1978

26 5
26 3.

7 4
5 1

3

1

tr

7

1

o

o
o

o
o

3
9

Salix spp,
(stems)

1977
1978

2

33 3

15 6

2
5

1
0.4

o
o

o
o

o
o

6

Purshia tridentata

1977
1978

o

0

tr

13 4
21 7

21 5

o

tr

tr
tr

SE

SE

SE

SE

SE

SE

Br ows e

·(StemS)

1

o

o

7

2

o

3

1

0

o

10 5
10 4

6
12

2

1

6

0.4

1

13

0.2

1
3

1 0.6
3

1

0.4
0.2

10 3
15 5
6

2

3

. 14

1

10 0.5

o
o

o

8

10
1

3

1

6

2

I
N

1.0

.....•
(stt!~

1977
1978

Rosa woodsii
(stem~

1977
1978

o

tr

tr
3

All browse~/

1977
1978

42 7
64 4

26 8
32 8

13
28

1977

o

tr

1978

o

o

1977
1978

tr

tr

1977
1978

2

1

2

1

0,4

4

3
2

4,756
6,300

Polenlilla

fruticosa

4

1

1 0.3
1 0.3

1

0.4

1
2

tr
tr

3
tr

1

o

2
tr

3

21 3
32 5

20 6
29 6

28 3
29 5

tr

20

4

o

10 2

5

8
5

1
2

3

1

1

0.3

1

0.4

2

0.4
0.4

0.3

30

3

11 3

11

2

1

1

18 4

10 4

6 1

3

2,800
14,580

15,885
9,105

12,485

21,575
25,685

28,850
17,790

tr
1

2
5

0.3

1

16

2

5

22

2

7

I

0.2
0.3

0.3
0.3

2
tr

5

38 4
53 4

32 3

56 4

6

o

tr

3

8

o

o

3

o
o

tr

6 4
40

44 3

Forbs
Eriogonurn umbellatum

Potentilla

spp,

All forbs~.I

Total bites~/

!/Neans are calculated

1977
1978

o

21,000
21,600

across 5 animals'

!?/Species include those which contributed

~/1977 = 1976-77, 1978 = 1978-79.

1

0.5
0.4

diet percentages

where percentages

2

9,930

0.4
1

o

tr

3

6

2

6

27,3:)0

155,250
131,910

26,910

were pooled for each animal over all months.

2% or more of total biles during any lIJonth.

0.4

�-298-

willow and aspen leaves.

This feeding behavior might have been over-

looked in previous studies because of the methods used.

Obviously,

no

evidence of leaf removal would be present when feeding sites were
examined.

Furthermore,

leaves were often eaten at the base of shrubs

where they were obscured from view by the surrounding
quently, observation

Conse-

from a distance would probably overlook such feed-

ing (Wallmo et al. 1973).

Because biomass of fallen leaves can be quite

large (Baker and Hobbs 1978) and their nutritional
high (see latter sections), elk consumption
tially increases the food potentials
Variation

herbage.

in composition

quality is relatively

of fallen leaves substan-

of habitats where leaves occur.

of diets among the five animals was rela-

tively small (Table 5).

There was no difference among animals (p &lt;

0.07) in the proportion

of grass or browse in their diets in any habitat

during either year.

Most standard errors of estimates of grass and

browse composition

of diets within individual habitat types were less

than 15% of the means.

Across-habitat

than those of within-habitat

estimates.

standard errors were even lower
The lack of variation

in diet

estimates

suggests that all animals responded similarly to the extant

variation

in forage res~urces.

When available,
material.

Bluegrass

consistently

green herbage was preferred to senescent plant
remained green for most of the first winter and was

sought out and consumed.

During Year 2, a colder winter,

senesced by November and was consumed less frequently

0.0001).

Animals also selected green tillers of timothy and sedges in

preference

to standing-dead

Initially,

(P

=

bluegrass

material of the same species.

grasses dominated Year-l winter diets (75%, November),

but the proportion

of browse increased and the proportion

of grass

�-299-

declined by 6 percentage

points per month until grass and browse were

eaten with almost equal frequency
March (Fig. 8).

On the other hand, grass and browse comprised approxi-

mately equal proportions
accounting

(45% of bites) during February and

of diets throughout

the second winter, togeth~r

for more than 90% of total bites each month.

a consistently

small proportion

of total observed bites (less than 10%)

during both years except in the xeric communities,
(Eriogonum umbellatum)

Forbs comprised

where sulfurflower

was an important diet item (Table 5).

More browse and less grass was consumed during Year 2 than during
Year 1 (P

= 0.001, Table 5).

This increase in browse intake was ob-

served across all habitat types (year x habitat interaction,

P

= .76).

Increases in browse consrunption in willow, wet meadow, aspen, wet shrub
meadow, and mesic meadow resulted primarily
of willow stems.

In xeric communities,

more frequent consumption

of bitterbrush

from increased

consumption

the increase in browse reflected
stems.

In both years, frequency of browse consumption

was greater in

aspen, willow, and wet shrub meadow than in other vegetation
0.05, Table 5).

types (P =

The aspen, willow~ and wet shrub meadow communities

contained the largest biomass of shrubs (Fig. 6), suggesting
was consumed according to its availability.
browse consumption

that browse

However, differences

in

among habitats,

although statistically

significant,

were small compared to differences

in browse availability

among habi-

tats.

Browses were consumed in greater amounts than would be predicted·

on the basis of their prewinter biomass

(Fig. 9).

Diets contained no

less than 13% browse in grassland, mesic meadow, and wet meadow communities where shrubs contributed

less than 3% of total plant biomass.

Thus, we can conclude that elk were highly selective
ularly in habitats with a small biomass of shrubs.

for browse, partic-

�-300-

76-77

(f)

w
~

co

Shrubs

....J
~

o
~

~
o

Ix

40
Shrubs

Dec I

Fig. 8.

Ja n l

Feb I

Mar ,.

Botanical composition of elk winter diets estimated across
habitat types on upper montane winter range, Colorado, during
1976-77 and 1977-78.

�-301-

100

Consumed more often
than expected
&lt;,

G

I

80

I

.- - s-

J-W

o

60

z

S

S

-,

/.

s- '\ I G

G G

I

G

(\

\\

s

I
I

G G

/

.

I
G

G

G

s
&lt;,

&lt;,

/
/

G /

......_-

_ _..- ;7

Consumed less often
than expected

60

80

100

0/0 IN BIOMASS

Figure 9.

Relationship between proportion of forage classes in prewinter herbage biomass and their proportion in overwinter
diets of elk on upper montane winter range, Colorado, during
1976-77 and 1977-78. Each point represents the mean forage
class percentage across 5 elk in one habitat type during one
winter. Symbols are: S
shrubs, G
grass, F
forbs.

=

=

=

�-302-

In contrast to this selective

feeding on relatively

rare browse,

elk also included large amounts of common forages in their diets in most
habitats.

Between 40% and 75% of bites were chosen from the 4 most

abundant

(based on prewinter biomass) plant species in all habitat ex-

cept mesic and wet meadow (Table 6).
ponderosa-pine

grassland

and

shrub, these species were consumed totally non-selective-

ly; that is, the percentage
not different
biomass

In aspen, sagebrush,

of common species in the overwinter

from the proportion

diet was

of those species in the prewinter

(Fig. 10).

This contrasting

evidence for selective and non-selective

feeding

styles can be reconciled by the following observation.

In all habitats,

diets contained

(Fig. 11).

a similar mixture of grasses and browse

gardless of the relative availability
contained

substantial

choices.

implications

this

The independence

mixture of diets from the grass-browse

tat biomass had important
discussed

In some habitats this

feeding style; in other communities

forage mix resulted from non-selective
grass-browse

of grass and browse, elk diets

amounts of both forages.

required a highly selective

Re-

composition

for maintenance

of the
of habi-

of diet quality,

later.

Forage Quality
Chemical

composition

grasses showed differences

and digestibility

typical of these forage classes (Van Soest

1973, Short et al. 1974) (Table 3).
than browse, a larger proportion
lulose (CW-ADF) and cellulose
protein

analyses of browse and

Grasses contained more cell wall

of the cell-wall material

(ADF-lignin),

as hemicel-

and less as lignin.

Crude-

content was greater in browse tissue, while IVDDM was greater

for grasses.

�-303-

Table 6.

Percentage of 4 most common species in biomass of upper
montane habitats, Colorado, and in winter diets of 5 elk
feeding in those habitats during November-March 1976-77 and
1977-78.

Habitat

Year

% of Total
Bites

% of
Biomass

Grassland

1977
1978

43
44

47
43

Ponderosa pine-shrub

1977
1978

42
48

48
45

Sagebrush

1977
1978

53
55

58
59

Willow

1977
1978

52
79

90
92

Aspen

1977
1978

78
72

75~/
66

Mesic meadow

1977
1978

7
27

77
72

Wet shrub meadow

1977
1978

60
58

79
72

Wet grass meadow

1977
1978

26
34

81~/
78

~/Includes leaf biomass estimate from 1978.

�-304-

100

Consumed
than

more often
expected

80

f-

60

w

o
z

40
~
o

Consumed less often
than expected

40

20
0/0

Figure 10.

IN BIOMASS

Relationship between proportion of 4 most common species in
Aspen, Grassland, Ponderosa-pine shrub, and Sagebrush habitats and their mean percentage in winter diets across 5 elk
on upper montane winter range, Colorado, during 1976-77 and
1977-78.

�-305-

80

o Grass

,._eo 60,._,._
40f.-

/

(f)

20

rn

t- .:

v1/ .:
V --

-r:

_J

80

~
0
I-

V--

nvQ
v- --

/

/

V

/

v-

l/r-

v .:
1/ .: I1/

vv::--

/

--

"-

I

Ix ~

~

40~

20 -

V

v-:/: -.

/
/

--

./:
' --

V -,/

n n
.•. ::

!J')

,._

0
Q)

Figure 11.

/

:.

v- ---

/

..c

.2

C'I

n, n,

CI)

c: c:

,/~

"0

Q)

0

r-

J::.
!J')
::J
,._
Q)

0

~:
h

V

I-,

V

,._

0
(!)

/

V
/
.-

/

1/

/

V

,/

~

v

V --

V

'r.

v:v:/: :

3:

3:

0

.~
!J')
Q)

0
Q)

::i; ~

Q)

-

/: .

h

"0

-

/:

/

L/nl-, V

c:
!J')
!J')

.,...

V
V

V

V
l/

v:_
v-r.:

0

"0

v1/

/

./

l,V
l/

~

V

/

,/

v:

--

,/

:1-,

1/
1/

r-

V

/

/

,/ -,/:-

V

----

~

V

::

,..., .:

1/
1/-

17"

V
V
l/

1/
1/

-V
1/ .:

--

v:1/~:: v-v::h
h
/

/

--

--

,

1./- --

1/'"":
1/ ---/

17" - -

,.... --

t-

V

~

-s ,._ 60~

0

/
/

vr.

I-

/

V
V
V
l/
Vr.
-V .:

V

w

r

V

I

m

DForbs

OShrubs

0
"0

0
Q)

:;: ~

[7

~

1/

V

;":

1/......,

/" --

l/ .:
1/:-

' --

~::

/-

" ..

/

V --

,/

1,./ .:

,/ --

--

: :1-,

..c
,._::J
c:
Q)
Q.

If)

&lt;[

J::.

-

(/)

3:
0
"0

V:-

3:
0

0
Q)

~ ~

~

Forage class composition of elk winter diets in upper
montane habitats during 1976-1977 and 1977-1978.

�-306-

Browse leaves were intermediate
ibility and crude-protein
cell-wall material,

to grass and browse stems in digest-

content, but contained the least amount of

and thus the greatest amount of cell solubles.

The

ADF fraction of cell wall was lower in leaves than in browse stems or
grass; hemicellulose

was thus most concentrated

in leaves.

Cell solubles in browse leaves and stems were not completely
gested, as evidenced by the discrepancy
percentages

between their cell-soluble

(l-CW) and IVDDM coefficients

(Table 7).

Short et al.

(1974) found that cell solubles in browse tissue were essentially
digested, but Milchunas
mine.

Presumably,

et al. (1978:9-10)

these conflicting

the extent of rupture of cell walls.
to variation

di-

100%

reported results similar to

findings result from differences
Whether these differences'are

in
due

in the browse material analyzed or to sample-preparation

methods is not clear.
Forbs were generally minor diet items.
tescant perennial,

was the single exception

stems in chemical composition,
The chemical composition
patterns

Sulfurflower,

a suffru-

and was similar to browse

IVDDM, and fiber characteristics.
of grasses and browse showed different

of change with advancing

season.

Protein content, cell wall,

and IVDDM of browse species did not change appreciably

during winter

(Slope f 0: P &lt; 0.37); whereas cell wall of grasses increased and their
crude protein and IVDDM declined from November until late February,
the trends reversed
I observed
grass quality.

when

(Fig. 12-14; Table 8).

some year-to-year

differences

in the temporal changes in

IVDDM declined more rapidly during Year 2 than Year 1 (P

= 0.04), but there was no difference in the .rate of change of crudeprotein or cell wall between years (P &gt; 10, Table 8).

Despite this

�Table 7.

Chemical
Colorado

constituents and in vitro digestibility
of principal forage species
upper montane range dlHing Novembe r-Ha rch 1976-1977 and 1977-1978.
Percent

Taxa~!

yearE.!

Number of
samples

inermis

1977
1978g!

by 5 elk on

of dry matter~!

ADF'=_/

cwci!

conswned

Lignin

Crude
Protein

1VDDMY

X

SE

X

SE

X

SE '.

X

SE

X

SE

4
3

62

2.7

33

1.5

4.8

0.4

4.2

0.8

47

5.0

1977
1978

3
2

72
78

1.5
1.2

36
39

0.5
0.0

5.3
5.5

0.2
0.7

4.6
4.3

0.4
0.3

41
47

5.8
3.4

1977
1978

5

64
72

1.9
1.3

36
40

1.4
1.4

4.1
5.3

0.4
0.6

4.7
3.9

0.5
0.4

43
43

2.2
3.3

Carex spp.

1977
1978

5
5

68
73

2.8
1.8

36
36

1.5
1.9

3.6
6.3

0.6
2.7

4.3
4.6

0.5
0.3

45
42

2.0
2.3

Juncus balticus

1977
1978

g!

1

79

1977'
1978

5
5

74
77

1.6
0.7

36
44

1.0
4.1

4.6
4.0

0.4
0.2

4.0
3.5

0.2
0.2

41
43

2.2
3.4

pratense

1977
1978

5
4

60
70

1.9
2.3

31
38

2.6
1.8

4.3
5.8

1.1
0.2

4.2
3.9

0.8
0.9

50
50

2.8
4.2

Poa pratensesis

1977
1978

4

61
73

2.8
1.4

34
40

1.8

3.3
4.8

0.3
0.4

5.0
4.0

0.7
0.5

50
44

2.4
1.4

Stipa comata

1977
1978

5
1

68
75

1.8

36
39

1.7

4.9
6.4

1.0

4.9
3.4

0.6

46
44

1.0

X eraminoids

1977

3

66

0.8

35

0.7

4.4

0.2

4.5

0.1

45

1.3

1978

7

75

1.1

[~O

0.8

5.1

0.4

3.9

0.2

44

1.4

1977
1978

1

'49

g/

1977
1978

5
5

32
41

2.0
1.5

23
29

1.3
1.1

8.0
14

2.0
0.7

5.6
3.9

0.3
0.2

44
39

1977
1978

5
5

34
49

1.0
1.4

26
38

1.0
2.5

13
23

2.1
1.6

6.0
p.2

0.2
0.4

34

1.1

29

2.6

Graminoids
Bromus
--,

Bouteloua

gracilis

Calamagrostis

Muhlenbergia

Phleum

Browse

canadensis

montana

41

2.9

1.0

0

......,

38

3.1

r

leaves

Alnus tenuifolia

~opulus

tremuloides

Salix spp ,

r

w

38

25

42

7.0

0.7
2.3

�Table 7.

Continued
Percent

cwc&lt;l/

ADFr:./

of dry matter~.!

Lignin

Crude
Protein

lVOmlY

----

Number of
samples

X

SE

X

SE

X

SE

X

SE

X

SE

2
1

32
42

0.2

23
43

0.2

11
5

0.6 "

5.1
4.8

0.04

42
40

0.9
0.7

3

33
44

0.7
2.5

24
37

1.5
4.1

11
14

1.0
5.2

5.9
5.0

0.4
0.5

40
36

3.0
3.5

1977
1978

5
5

42
57

3.5
1.9

33
45

3.0
1.6

15
21

1.5
1.0

8.1
6.0

0.1
0.4

40
35

1.0
1.9

1977
1978

5
4

55
59

0.6
0.6

41
44

0.6
0.8

16
17

1.2
0.2

5.4
4.6

0.1
0.1

26
23

2.1

1977
1978

4
4

48
56

1.1
0.9

36
45

1.5
3.4

20
25

0.1
0.7

8.1
6.8

0.2
0.5

35
29

0.8
2.9

Rosa woodsi

1977
1978

2
1

49
52

0.5

36
43

0.9

12
16

1.0

4.9
4.6

0.4

38
35

2.0

X browse

1977
1978

4
4

49
56

2.4
1.4

36
44

1.7
0.4

16
20

1.7
2.0

6.6
5.5

0.9
0.5

35
30

3.1
2.9

Eriogonuro umbellatum

1977
1978

4
3

43
55

2.0
0.3

34
43

2.0
1.1

17
18

1.6
1.1

7.8
6.3

0.3
0.2

30
25

1.0
3.0

Potentilla

1977
1978

1

52

'&amp;_/

1977

2

49

4.5

34

0.5

14

3.0

6.1

0.3

28

2.0

1978

1

55

0.3

43

1.1

18

1.1

6.3

0.2

25

3.0

across monthly

samples;

Year':./

Taxa!?.!

1977
1978

Prunus virginiana

X browse
Browse

leaves

.

stems

Salix spp.
Potentilla

Purshia

fruiticosa

tridentata

I

-----

stems

Forbs

spp.

X forbs

!/X is the mean of percentages
~/Species

contributing

~/Cell-wall
~/Acid

constituents.

detergent

SE is standard

2% or more of total bites observed

£/1977 = winter of 1976-77, 1978 = winter of 1977-78.
fiber.

!/In vitro

digestible

g/Species

did not occur in diet that year.

dry matter.

11

33

error of mean.

during anY,month.

31

4.3

w

0
00
I

�-309-

GRASS

CELL

WALL
2

~~2

75
2

70
_J
_J

~

2

I

2

/I/~

I

~

~
2

/~

I

Nov I

Dec I

2

65

_J
_J

w

&lt;..)

Figure 12.

.Jcn I

Feb I

Mar I

Apr I

Cell wall content of grasses consumed by elk on upper
montane winter range, Colorado. Numeral 2 indicates a value
for 1977-78; numeral 1 is a value for 1976-77. For regression statistics see Table 4.

�-310-

GRASS PROTEIN
6.5

6.0

5.5
z
w

I0---

0::

o,
w

4.5

0

::J

0::
L)

~

0

3.5

2
I

2

2
2
2

Novl

Figure 13.

Dec I

Janl

Feb I

Marl

Apr I

Protein content of grasses consum~d by elk on upper
montane winter range, Colorado. Numeral 2 indicates a
value for 1977-78; numeral 1 is a value for 1976-77. For
regression statistics see Table 4.

�-311-

GRASS DIGESTIBILITY

60
55
50

o
~

o

&gt;

~
o

2
2

35

Nov I

Figure 14.

Dec I

Janl

Feb I

Mar I

Apr I

In vitro dry digestible dry matter (IVDDM) for grasses consumed by elk on upper montane winter range, Colorado. Numeral 2 indicates a value for 1977-78; numeral 1 is a value
for 1976-77. For regression statistics see Table 4.

�-312-

Table 8.

Regression coefficients and statistics for in vitro digestible
dry matter (IVDDM), crude protein content, and cell wall of
grasses from upper montane winter range, Colorado, as a
function of advancing season. Independent variable is days
where 15 October
day 1. See Fig. 2-4 for plots.

=

Dependent
variable

Year'p_/

IVDDM

Equation

R2

P

SE£./

0.31

0.007

5.0

0.50

0.003

2.6

0.38

0.001

0.9

1978

= 55.8 - 0.24X + 0.001X2
Y = 51.8 - 0.18X + 0.0008X2
Y = 6.S - 0.04X + 0.0002X2
Y = 5.8 - O. Ol}X + O. 0002X2

0.46

0.005

0.7

1977

Y

= 56.7 + 0.21X - 0.0010X2

0.23

0.02

5.2

1978

Y

= 62.6 + 0.20 - 0.0009X2

0.45

0.001

2.7

1977
1978

Crude protein

Cell wall

~/Includes

1977

Y

only those species for which a sample was collected each month.

=

'p_/1977 winter of 1976-77, 1978
£/Standard

error of estimate.

= winter of 1977-78.

�-313-

annual variation,

these data suggest that the nutritional

quality of

grasses generally declines until March, then begins to increase.
Measures of forage quality were markedly lower during Year 2 than
Year 1 (P &lt; 0.02, Table 7):

Mean crude protein in grass was 13% less;

protein in browse sterns, 15% less; and mean crude protein of browse
leaves, 17% less.

IVDDM of grasses did not change between years, where-

as browse sterns were 14%, and browse leaves 10%, less digestible

the

All forage classes increased in ewe between years; browse

second year.

sterns and grasses increased 12%; leaves, 25%.
Diet Quality
The previous section dealt with the chemical composition
forages consumed by experimental
composition

and digestibility

on the proportions

animals.

However, the exact chemical

of the diets (diet quality) also depended

of the various forages selected.

Diet q~ality was

estimated quite precisely because chemical composition
classes varied little, and forage-class
very similar.

of the

As a consequence

composition

of animal diets were
statistical

differ-

ences between means resulted in large part from small variances

rather

than from large differences
variances

of this precision,

within forage

in mean values.

can contribute to statistical

is of greatest biological
As discussed,

Note that, although small

significance,

the value of means

importance.

the nutritional

quality of forage grasses decreased

steadily through the winter (Fig. 12-14), while nutritional
browse remained constant.

quality of

The quality of elk diets (calculated across

habitats by month) also decreased over the winter (Fig. 15 and 16, Table
9), but the rates of decline and magnitudes
from those of grasses (Tables 8 and 9).

of change sometimes differed

Dietary crude protein decreased

�-314-

DIET

IVDMD

45
0
~

0

&gt;

~

22

35

22

2

0

2

30
Nov I

Figure 15.

Decl

Jan I

Feb I

Mar I

Apr I

In vitro digestible dry matter (IVDDM) of winter diets of 5
elk on upper montane winter range, Colorado, during 1976-77
. and 1977-78. Numeral 1 indicates a value for 1976-77. Numeral 2 is a value for 1977-78. For regression statistics
see Table 6.

�-315-

DIET

7.0

PROTEIN

z
W

I0

~

n:::

a,
w
0

:J

n:::
l)

1

2

21

~
0

2
2

2

Novl

Figure 16.

Decl

Janl

Feb I

Marl

Aprl

Protein content of winter diets of 5 elk on upper montane
winter range, Colorado, during 1976-77 and 1977-78. Numeral 1 indicates a value for 1976-77; numeral 2 is a value for
1977-78. For regression statistics see Table 6.

�-316-

Table 9.

Regression coefficients and statistics for winter diet crude
protein and IVDDM of 5 elk on upper montane winter range,
Colorado, as a function of advancing season. Independent
variable is days where 15 October
day 1. See Fig. 5 and 6
for plots.

=

Dependent
variable
IVDDM

Year~/

Equation

R2

P

SE~/

1977

= 52 - 0.19X + 0.0007X2
Y = 46 - 0.18X + 0.0007X2
Y = 6.2 - 0.009X
Y = 5.8 - 0.02X + 0.00008X2

0.82

0.00001

2.1

0.64

0.0005

1.8

0.56

0.00002

0.4

0.60

0.00003

0.3

1978
Crude protein

1977
1978

~/1977

Y

= winter of 1976-77; 1978 = winter of 1977-78.

~/Standard error of estimate.

�-317-

about 20% from November

through March during both years, while grass

crude protein decreased

about 30%.

During Year 1 the rates of decline

of both dietary IVDDM and crude protein were lower than those of grass
IVDDM and protein.

In the second year the rates of decline for dietary

IVDDM and protein did not differ from those of grass IVDDM and protein.
In Year 2, although the rates of decline in dietary crude protein
did not differ significantly
levels were consistently
protein levels.

from those of grass, dietary crude-protein

higher (:1 percentage

On the other hand, Year 2 IVDDM levels were consistent-

ly higher in grasses than in the diet.
elk maintained
available

point) than grass crude-

Thus, in Year 2, as in Year 1,

a diet higher in crude protein than levels generally

grasses but, unlike in Year 1, were unable to also maintain

higher level of digestibility

than was generally

available

During both years, there was no significant

a

in grass.

change in dietary cell

wall content during winter.

Constancy

of diet cell wall content con-

trasted with the overwinter

variation

in grass cell wall content

(Fig.

Diet quality, like forage quality, declined between years.

Diets

15).

contained

less crude protein during Year 2 than during the previous year

(P = 0.0001).
ties:

However,

these differences

grassland, ponderosa

dietary-protein

pine-shrub,

were limited to three communiand aspen (Table 10, Fig. 17);

content in all other communities

did not change signif-

icantly (P &lt; 0.05), despite the marked reductions

in forage protein

content.

Dietary IVDDM was lower the second year in all habitats

grassland,

sagebrush,

18).

and ponderosa

pine-shrub

(P = 0.05, Table 10, Fig.

Dietary crude protein declined most sharply in grassland

ponderosa

pine-shrub

(Fig. 17).

Thus, constancy

except

and

of IVDDM in diets was

�-318-

Table 10. Mean percentage chemical constituents and in vitro digestibility of winter diets of 5
elk in upper montane habitats in Colorado, during November-~larch 1976-1977 and 1977-1978.
Percent
CWC"r}_/

of dry matter

ADFr:_/

Lignin

Crude
protein

IVDm1&lt;!i

Year~/

X

SE

X

SE

X

SE

X

SE

----X
SE

Grassland

1977
1978

53
61

0.1
0.3

33
48

0.6
0.5

9.3
13.6

0.2
0.7

6.3
5.1

0.1
0.1

35
39

0.7
1.2

Mesic meadow

1977
1978

61
62

0.5
1.6

34
38

0.2
1.3

4.7
11.0

0.4
1.0

4.6
4.5

0.1
0.5

46
36

0.5
1.0

Ponderosa

1977
1978

60
61

0.2
2.8

35
50

0.2
1.0

8.2
8.3

0.9
0.9

5.5
4.8

0.1
0.2

40
37

1.0
0.7

Sagebrush

1977
1978

64
63

1.6
3.8

35
52

0.5
1.1

7.7
8.7

0.8
1.0

5.2
4.8

0.2
0.1

38
38

0.3
1.2

Willow

1977
1978

52
58

0.6
0.3

32
41

0.2
0.3

8.1
17.8

0.5
0.2

5.6
5.6

0.1
0.2

42
35

0.5
0.2

Aspen

1977
1978

53
59

1.7
1.2

31
37

0.7
0.2

5.2
12.0

0.4
1.0

5.2
4.6

0.1
0.2

47
39

0.5
1.7

Wet shrub-meadow

1977
1978

59
60

0.8
1.0

35
40

0.2
0.4

6.8
15.0

0.5
0.6

5.2
5.2

0.1
0.2

43
34

0.5
0.4

Wet meadow

1977
1978

61
63

0.8
2.3

35
41

0.5
0.5

6.2
12.7

1.1
2.2

5.1
5.0

0.1
0.3

45
37

1.6
1.0

Habitat

Tukey's

pine-shrub

cF/

Among habitats

6.0

2.8

3.7

0.60

4.1

Between years

4.5

2.1

2.8

0.45

3.1

~/1977 = winter of 1976-77, 1978 = winter of 1977-78.
~/Cell-wall

constituents.

r:_/Aciddetergent

fiber.

~/In vitro digestible

dry matter.

e/Tukey's Q simultaneous confidence intervals for 8 (among habitats) and 2 (between years) comparisons. Means must differ by this value to be significantly different at P
0.05.

=

�-319-

DIET

7

PROTEIN
GRSL

z
w

I0

0:: 6
0....

w

0

:::&gt;
0::

WILW

H

l.)

PP/SH

\H \h
WSHM

ASPN

SAGE

WGSM

I
I

~

~

0

MGSM

·4
I

4

Figure 17.

Protein content of winter diets of 5 elk in upper montpne
habitats, Colorado. X indicates a mean across animals;
vertical bars are Tukey's Q simultaneous confidence intervals for eight comparisons; non-overlapping bars indicate
significant differences at p = 0.05. Solid lines are for
1976-77, dotted for 1977-78. Habitat abbreviations are:
WILW
willow, PPjSH
ponderosa pine-shrub, WSHM
wet
shrub meadow, ASPN
aspen, SAGE
sagebrush, WGSM
wet
meadow, GRSL = grassland, MGSM = mesic meadow.

=

=

=

=

=
=

�-320-

DIET

50

DIGESTIBILITY
ASPN

MGSM
WGSM

45

WSHM
WILW
PPISH

0

~

0

&gt;

40

~

0

35

Figure 18.

Jf:

\

GRSL

SAGE

I
I

N

'r

I

In vitro digestible dry matter (IVDDM) content of winter
diets of 5 elk in upper montane habitats, Colorado. X indicates mean value across animals; verticle bars are Tukey's
Q simultaneous confidence intervals for 8 comparisons. Nonoverlapping bars indicate significant difference at p
0.05.
Soild lines are for 1976-77, dotted for 1977-78. Habitat
abbreviations are: WILW = willow, PP/SH = ponderosa pineshrub, WSHM
wet shrub meadow, ASPN
aspen, SAGE
sagebrush, WGSM
wet meadow, GRSL
grassland, MGSM
mesic
meadow.

=

=
=

=

=

=

=

�-321-

70

DIET

CE LL WALL
WGSM

65

PP/SH WSHM

_J
_J

I

I

&lt;!
_J
_J
W

60

(..)

~
0

I

WILW

:5:

55

ASPN
I

I
I

Hr

F

GRSL MGSM
I
I

I

I

r:

50

Figure 19.

Cell wall content of winter diets of 5 elk in upper montane
habitats, Colorado. X indicates mean value across animals;
vertical bars are Tukey's Q simultaneous confidence intervals
for 8 comparisons.
Non-overlapping bars indicate a significant difference at p
0.05. Solid lines are for 1976-77,
dotted for 1977-78. Habitat abbreviations are: WILW
willow, PP/SH
ponderosa pine-shrub, WSHM
wet shrub meadow, ASPN - aspen, SAGE
sagebrush ,-WGSM
wet meadow, GRSL
grassland, MGSM
mesic meadow.

=

=

=

=

=

=

=

=

�-322-

associated

with sharply reduced crude protein, while stable dietary

protein occurred most often in habitats where dietary IVDDM declined.
Cell wall content of diets increased between years in aspen and grassland; in all other habitats proportions

of dietary cell wall remained

= 0.05, Table 10, Fig. 19).

These annual trends in diet

constant

(p

nutritional
mixtures

quality were influenced by changes in the browse-grass

of elk diets, an influence discussed

in more detail later.

During both years, dietary IVDDM values were highest in aspen and
protein content of diets was consistently
in mesic meadow (Table 10).
types according

high in willow but usually low

Despite these generalities,

to nutritional

ranking habitat

quality of diets selected was not possi-

ble for two reasons.
First, there were significant
actions for diet protein,

(P &lt; 0.05) year-X-habitat

IVDDM, CW, ADF, and lignin.

annual changes in diet quality were not proportional

inter-

That is, because
across habitats,

Year 1 ranking of habitats by diet quality differed significantly

from

that of Year 2 (Fig. 17-19).
Second, the differences
(Table 10, Fig. 17-19).
contained

habitat

During the first winter, diets in grassland

a large proportion

consequently

in diet quality among habitats were small

of sulfurflower,

more crude protein

(P &lt; 0.05).

(X

Diets in other communities

the most protein,

selected in aspen, sagebrush,

forb, and

= 6.3%) than diets in any other

and 5.6% protein, most having close to 5%.
willow contained

a high-protein

contained between 4.5

The following year, diets in

significantly

or mesic meadow

more than the diets

(P &lt; 0.05).

Otherwise,

dietary crude protein did not differ between habitats during Year 2, and
protein

in diets again ranged from 4.5 to 5.6%.

Dietary IVDDM did not

�-323-

differ in aspen, willow, mesic meadow, and wet shrub meadow during Year
1; diets in these habitats were more digestible
pine-shrub,

sagebrush,

and grassland

than those in ponderosa

(P &lt; 0.05).

The subsequent year,

dietary IVDDM values were higher in aspen and grassland
shrub meadow (P &lt; 0.05); otherwise,
habitats.

Cell wall percentage

than in wet

dietary IVDDM did not differ between

of diets was smaller in aspen, grass-

land, and willow than in any other habitat during year 1 (P &lt; 0.05).
There were no differences

in proportions

of dietary cell wall among the

other habitats that year or among any habitats during year 2 (p &gt; 0.05).
Nutritional

Status

As a result of the observed variation
differences

in predicted

in diet quality there were

animal condition between years, within winters,

and among some habitats.
Simulated weight fluxes for a calf during Year 1 were a 5% increase
in lean body weight, a 6% decrement

in total weight, and a complete ex-

haustion of fat stores (Fig. 20).

Adults during the first winter were

predicted

to gain 3% in lean body, and loose 60% of fat reserves and 10%

of total body weight (Fig. 21).
lost 6% of their prewinter

The following year, simulated

adults

lean body, 20% of their total weight and all

of their fat stores (Fig. 22).

When the model was parameterized

for

calves consuming Year 2 diets, estimated weight loss were 20% of lean
body, 100% of fat and 30% of total weight (Fig. 23).
weight, adults were predicted

In terms of total

to loose twice as much weight, and calves

5 times more weight during Year 2 relative to their losses the first
year.
During Year 1, simulated

changes in lean body weight of calves and

adults indicated diets chosen that year contained

above maintenance

�-324-

220

212

204

196

-0'1

x

»&gt;

- body
- - --

Lean

---

~

I-

:r:
(.!)

w

:s:

32

- - -Fat- ....
..•..

&gt;-

Cl

0

co

....

.....

....

.•....

24

&lt;,

.•....
.•....
&lt;,
&lt;,
&lt;,

16

••...
"- .•.•.

.•.•.

.... .•.•.

8

Dec I

Figure 20.

Jan I

Feb I

Mar I

Simulated changes in total weight, lean body, and fat of an
adult elk on upper montane winter range, Colorado, during
November-Narch 1976-77.

�-325-

90

Cl

82

~
••......

I-

:c

(!)

~

/

78

---

--- _-- - -_
Lean

body

/

w

3:

&gt;- 74

0

0

--

en

Fat

... ...

8

.•.. ...•.

4

Dec I

Figure 21.

Jan I

•.. ...

•.. •.•.

... •.•.

Feb I

•.•..

•.•.

•.•..

•.•...

•.•.. ...

Mar I

Simulated changes in total weight, lean body, and fat of a
calf elk on upper montane winter range, Colorado,
November-March 1976-77.

during

�-326-

220

210

200

-.._.. 190

-_ --- --

Q')

Lean

~

tI
(!)

body

w
3:

-- --

....
Fat
.•..

&gt;-

•...

Cl

0
CD

..•.

..•.

..•.

20

•...

....•

....

•...

•...
"-

....•

....•
....•
•....

10

•...

•...

....•

•...
....•.

Dec 1

Figure 22.

Jan 1

Feb

....•

•...

•...

•...

Mar 1

Simulated changes in total weight, lean body, and fat of an
adult elk on upper montane winter range, Colorado, during
November-March 1977-78.

�-327-

-

78

01

-_ --_

Lean body
-_

~

t-

:I: 72
(!)

w

5
&gt;a 66
0

m

_ _Fat
6

0

Figure 23.

Dec I

.•...

.•. ...

Jan

•... .•.

.•...

.•. .•...

.•....

•...

Feb I

Mar I

Simulated changes in total weight, lean body, and fat of a
calf elk on upper montane winter range, Colorado, during
November-March 1977-78.

�-328-

levels of protein until mid-winter,

at which time the diet protein

levels equalled simulated protein requirements

(Fig. 20, 21).

The

following year, lean body declined slowly until fat reserves were
depleted,

at which point sharp declines in lean body were predicted

(Fig. 22, 23). This result implies that protein balance per se during
Year 2 was only slightly negative and that a large portion of simulated
lean body deficits incurred that year resulted from catabolism
teinaceous

of pro-

tissue to meet energy requirements.

Fat reserves were predicted
the beginning

to be maintained

for a short period at

of winter during Year 1 and then decline steadily through-

out the remainder of the season (Figs. 20, 21).
of that year provided

Thus, diets in November

energy in excess of simulated

requirements,

while

during the following months, dietary energy was at submaintenance

levels.

During Year 2, fat was drawn down rapidly in both adults and calves from
the beginning

of winter

(Fig. 22, 23).

in calves by mid-February,
weight accelerated

Fat stores were entirely depleted

at which time losses of lean body and total

rapidly.

Results of model runs for diets chosen in the various habitats were
summarized

in Table 11.

Variation

in energy status of simulated

elk was

greater between years across habitats than within years among habitats.
This was the case because diet digestibility
over space but showed large annual changes

was relatively
(Fig. 18).

In most habitats during both years predicted
exceeded energy gains.
aspen community

The single exception

energy requirements

to this pattern was in the

during Year 1 where diets provided above predicted

tenance levels of energy.
balances

constant

That same year, simulated

overwinter

main-

energy

in willow, mesic meadow, wet meadow and wet shrub meadow were

�-329-

only slightly negative.
grassland.

Energy deficits were greatest during Year 1 in

The following year, energy deficits were greatest in willow

and wet shrub meadow and were smallest in aspen and grassland.
Simulated nitrogen balance prior to depletion

of fat stores was

less variable than was elk energy status (Table 11).
predicted

nitrogen flux ranged from a loss of 1.52 g • d-1 in mesic

meadow to a gain of 2.89 g • d-1 in willow.
provided nitrogen

in excess of simulated

During Year 2, predicted
habitats.

During Year 1,

Diets.in most habitats

requirements

the first year.

nitrogen fluxes were slightly negative

Again, maximum simulated nitrogen

in most

losses were encountered

mesic meadow (-3.38 g • d-1), and maximum gains in willow

in

(+2.39 g

• d-1).

Carrying Capacity
Consistent

with trends in total standing crop biomass,

carrying capacities

for elk based on forage quantity were greater in wet

and mesic habitats than in drier communities
years, wet meadow and willow could provide
quirements

estimated

of more elk than other vegetation

could support the smallest number.

(Table 12).

During both

for total forage intake retypes, while grassland

However, note that predicted

nutri-

tional status of elk in the various habitats was often at odds with
habitat carrying capacity estimates.
largest estimated

with the

support capacity based on forage quantity, predicted

weight losses of individual
smaller calculated

That is, in communities

elk were often larger than in habitats with

carrying capacities.

Thus, estimation

capacity based on total biomass gave poor estimates

of carrying

of the capability

those areas to provide for elk nutrition.
Annual variation
pronounced

in habitat carrying capacity estimates

than yearly variation

in herbage standing crops.

was less
This was

of

�Table 11.

Simulated estimates of energy and nitrogen
1977-78 in Colorado.

status of elk in upper montane

Energy Balance

habitats

(kcal)

during Novembe r-Ha rch of 1976-77 and

-1

Overwinter

per day

Grassland

1977
1978

-133308
-249749

- 966
-1810

+1.28
-1.09

-38
-15

75
129

Mesic Neadow

1977
1978

- 8219
-318324

--230759

-1.52
-3.38

-04
-26

1210
102

1977
1978

- 85747
-292816

- 621
-2121

+1.12
-2.30

-14
-21

116
110

1977
1978

- 96154
-269005

- 696
-1949

+0.20
-2.25

-20
-18

103
121

Habitat

Ponderosa

Pine Shrub

Sagebrush

Weight

Total
change

Time until
Fat Depletion

Year

Nitrogen Flux (g • d )
prior to Fat Depletion

(%)

Willow

1977
1978

9958
--350989

-

72
-2543

+2.89
+2.39

-04
-33

998
92

Aspen

1977
1978

+ 7045
-248990

+ 51
-1804

+0.51
-3.36

+02
-18

131

Wet Shrub Meadow

1977
1978

- 48186
-375223

- 349
-2719

+0.43
-0.61

-06
-38

206
86

Wet Headow

1977
1978

- 21376
-297938

- 154
-2158

-0.24
-1.22

-02
-22

465
108

~/Time until total kcal deficit

= 90% of kcals in fat stores.

No value given when animal is in positive

energy balance.

I

w
w
0
I

�-331-

Table 12.

Estimates of nutritional status and carrying capacity for elk in upper montane
habitat types during winter 1976-77 and 1977-78 in Colorado.
Nutritional

Status

Forage~/

Body Weight~/

Year~/

Intake (kg • d-1)

change (%)

-1
(elk days • ha )

Grassland

1977
1978

5.22
5.06

-38
-15

69
66

Hesic Headow

1977
1978

5.26
4.89

-04
-26

132
117

Ponderosa Pine Shrub

1977
1978

5.21
4.90

-14
-21

79
49

Sagebrush

1977
1978

5.09
4.92

-20
-18

79
49

Willow

1977
1978

5.45
5.09

-04
-33

174
218

Aspen

1977
1978

5.45
4.98

+02
-18

127
101

Wet Shrub Headow

1977
1978

5.26
4.93

-06
-38

154
139

Wet Headow

1977
1978

5.33
4.95

-02
-22'

189
117

Habitat Type

Carrying Capaci t'f~/

!/1977 = winter of 1976-77, 1978 = winter of 1977-78.
~/Based on predictions

of simulation model.

~/Predicted total weight change from Nov. 10 to Harch 25 based on simulation model
estimates of adult animals.
~/Carrying capacity calculated as herbage biomass
elk-I. day-I).

(kg/ha) . total intake (kg·

�-332-

the case because
diminished

reduced diet quality during the second year resulted in

rates of predicted

forage intake, and consequently,

tively higher carrying capacities.

rela-

�-333-

Chapter VII
DISCUSSION
The standing crop biomass of winter range herbage varied between
years and among habitats.

Simultaneously,

forage changed annually and within winters.
sources influenced
and estimates
tritional

quality of elk

Variation

in forage re-

elk diet quality, predictions

of carrying capacity.

status (herbage quantity,

will be discussed

nutritional

individually.

of nutritional

Each of these influences

status,
on elk nu-

forage quality, and diet quality)

The significance

of variation

in animal

condition and range carrying capacity will then be examined.

Following

this discussion,

of temporal

conclusions

and spatial variation

will be drawn on the importance

in food resources

in several theoretical

and

applied contexts.
Herbage Biomass
Variability

in herbage biomas~ between years resulted from across

habitat declines in grass standing crops and the differential
of forbs.

Shrubs production

Variation

responses

was constant.

in grass biomass appeared to be closely related to mois-

ture availability.

Differences

in grass production

among habitats

re-

flected moisture gradients between sites; wet and m~sic habitats had
greater standing crops of grasses than did dry vegetation
S). Annual differences

types (Fig.

in standing crop biomass of grasses appeared to

be related to decreased precipiation

during the winter prior to the Year

2 growing season (Oct.-Mar. 1976-77).

Although both Year 1 and Year 2

�-334growing seasons were preceeded by abnormally
tation measures

in Estes Park, Colorado,

dry winters, total precipi-

from October-March

1976-77 was

67% less than for the same time period the previous year (National
Oceanic and Atmospheric

Administration

1975, "1976, 1977).

With the

exception

of blue gramma, all of the dominant grasses were cool season

perennials;

growth of these grasses in the spring depends on water

stored in the soil profile, as well as early spring precipitation
(Veihmeyer and Hendrickson

1950, McMinn 1952, Gates 1968, Eddleman and

Minos 1972).

overwinter

deficits

Presumably,

in soil water potential

drought during Year 1 resulted in

at the beginning

of Year 2 which were not compensated
tation.

of the growing season

for by early growing season precipi-

Sneva (1977) found yield of mature crested wheatgrass

cristatum),

a cool season perenial,

0~70) with precipitation

was significantly

correlated

(r

=

during plant dormancy.

Shrubs, with their deep root systems and high root/shoot
less sensitive

(Agropyron

to variation

in soil moisture

ratios are

than are more shallowly

rooted grasses with their smaller root surface relative to shoots
(Anderson et aI. 1972:177).

We might expect, therefore,

that shrub pro-

duction would not decline as much as that of grasses as a consequence
reduced soil moisture.

However, note that shrub production,

its relatively patchy distribution
measure

because of

in space, is much more difficult

than is biomass of the more evenly distributed

of

grasses.

to

Conse-

quently, the standard errors of means of shrub biomass were much larger
than those associated

with estimates

of grasses biomass

Hence, failure to detect year differences
resulted from sampling error.

(Appendix 3).

in shrub production

might have

�-335-

The differential

response of forbs in the various habitats is dif-

ficult to explain and doubtlessly

resulted from a variety of influences.

However, a plausible

for the observed year x habitat inter-

explanation

action in forb biomass involves the differences

in herbage structure

in

the wet and dry communities.
Hufstader
grasslands

(1976:435) suggested that dominant species in California

tended to be limited by availability

tion of sub-dominants
Extending

of water, while produc-

was more closely coupled to light availability.

this prediction

to the current data, forbs, being sub-

dominants, would be light limited.

Further,

since most of the forbs in

the study area were warm season annuals, they would have been less
affected by early growing season soil water than would the cool' season
grasses.

In the wet and mesic types, where forb biomass increased be-

tween years, grasses formed a nearly continuous
sihusia.

The Year 2 increase in forbs in these vegetation

have resulted from increased light penetration
to the forb layer as a consequence

above the forb
types could

through the grass canopy

of decreased grass production.

the dry types, where forb biomass declined,
continuous

overstory

In

grasses do not grow in a

sward above forbs; hence, we would expect no increase in

light availability

to the forbs during Year 2.

Further,

a large portion

of dry type forbs were perennials

which would be expected to be more

sensitive to soil water potential

deficits than the more ephemeral

annuals in the wet types.
Forage Quality
Annual variation

in the crude protein content and digestibility

range herbage has been previously
1979, Gibbs 1978).

Usually,

documented

the nutritional

of

(Dietz 1967, Pease et al.
quality of herbage declines

�-336-

as herbage production

increases.

This reduction in herbage quality

occurs because the concentration
well as the digestibility
growth.

of crude protein in plant tissue as

of that tissue is diminished during plant

As the plant matures, plant structural carbohydrates

orated more rapidly than cell solubles.
total production,

are elab-

It follows that increases in

while elevating the total amount of crude protein and

cell solubles per unit area of range, will result in decreased concentrations of crude protein and cell solubles and increased cell wall per
unit of plant biomass.

It is surprising,

therefore, that elk forages

during Year two, a year with markedly reduced herbage production,

con-

tained more dilute concentrations

of crude protein, and digestible dry

matter and higher concentrations

of cell wall.

This surprising

result can be explained by the difference between

changes in the overall quality of herbage on offer and changes in forage
material actually selected.

Note that plant samples selected for nutri-

tional analysis were chosen to mimic the material selected by grazing
elk.

Based on numerous studies of diet selection by wild and domestic

reminants

(reviewed by Klein 1971)", we can assume that the forage in-

cluded in diets was generally of higher nutritional
the total available plant biomass.
of potential

quality than that of

That is, there existed a finite set

diet items which exceeded the mean quality of herbage, and

this set was included to greater or lesser extent in elk diets depending
on their dietary selectivity.
very selectively.

I showed that elk could, indeed, feed

Thus, a reduction in total production,

while ele-

vating the mean quality of herbage, would also reduce the absolute size
of the set of higher quality forages.

Consequently,

we might expect the

quality of forage included in an elk's diet would more closely resemble

�-337-

the quality of herbage on offer.

This convergence

could plausibly

re-

sult in a reduction in the quality of diet items selected despite an
increase in herbage quality.
The decrement

in forage protein, digestibility

and cell solubles

between years could a~so have been due to earlier and more complete
senesence of forage plants the second year.

During Year 1, there

appeared to be much more green material available
As previously

discussed

throughout

the winter.

this forage WaS often consumed by my experimen-

tal animals and was frequently

included in forage samples.

The fol-

lowing year, there appeared to be much less green herbage present on the
winter range, and it was rarely collected.

A shift in the composition

of forage samples to include more dead material during Year 2 could
easily explain the observed annual differences
Overwinter
curvilinear

changes in nutritional

patterns

quality of grasses similar to the

I observed for grasses

and decrease in protein,

digestibility,

in forage quality.

(an increase in cell wall

from November

through February,

followed by a reversal of the trends) have been reported previously
(Willard and Schuster
no explanation
rated.

1973, Sims et al. 1971, Burzlaff

for this temporal variation

I believe the overwinter

1971

). However,

has been previously

elabo-

changes in forage quality can be ex-

plained by two processes.
The decline in quality can be attributed
from periodic

snowmelt percolating

leaching cell solubles.

to the effects of water

through senescent plant tissue and

Leaching can rapidly remove substantial

of soluble plant dryrnatter, including

sugars and amino acids (for review

of effects of leaching on plants, see Tukey 1971).
browses

amounts

I analyzed with their highly lignified

Presumably,

the

cell walls and waxy

�-338-

cuticles were more resistant to leaching and, as a result, did not show
these overwinter

declines in nutritional

quality.

The increase in grass quality in the spring probably resulted from
upward translocation

from roots to shoots.

Green-up of perennial

grasses

was observed during March of both years, and this green material was
included in animal diets and in forage samples.
material

An increase in the live

in March samples would account for the observed inceases in

forage quality.
Botanical

Composition

Elk were observed to feed selectively

on browses while frequently

consuming very abundant forages (primarily grasses) and rarely eating
forbs.

How can these feeding patterns be integrated with what we know

about ungulate diet selection?
Diet selection by large herbivores
ological and anatomical
of these limitations
relatively
requirements

constraints

like elk is influenced by physi-

on their food choices.

The result

is that animals of large body size must consume

common foods.

This is the case because energy and nutrient

of mammals are functionally

related to their body weight

raised to the 0.75 power (Kleiber 1975).

Consequently,

animals have smaller nutrient requirements

per unit of body mass, but

larger total nutrient requirements,

large bodied

than do animals of smaller size

(Jarman 1974, Geist 1974, Jannis 1976).

The rate at which a ruminant

can consume a high fiber diet is limited by rumen fill (Ammann et al.
1973, Campling et al. 1970).

Rumen fill, in turn, is also related to

body size; large bodied ruminants tend to have larger rumens per unit of
body mass than do small ruminants
intake is determined

(Hoffman 1973).

Since total nutrient

by the amount of food consumed and the concentrations

�-339-

of nutrients

in that food, the above allometeric

relationships

constrain

large ruminants to eat large amounts of forage, but permit them to
consume foods of relatively
consume relatively

low quality.

common food items.

the largest North American herbivores,

In order to do this, they must

Therefore,

since elk are one of

we expect that the food dimension

of their niche would include a large portion of relatively

common foods.

In most cases, these common foods were grasses.
How can we account for the intense selectivity
ulate that browse consumption

for browse?

was related to its higher protein content

relative to grass and elaborate more fully on this advantage
tion on diet quality.
non-selective

In short, however, selectivity

in grass.

styles are not incompatible.

sufficient

requirements,

non-selective

feeding

feeding fine-tuning

the

of diets.

The apparent rejection of forbs was probably
tial distribution

feeding

Indeed, the two feeding modes interact to

total intake, and selective

nutrient concentrations

the protein

Thus, selective and non-selective

provide for different nutritional
providing

in the sec-

for browse allows

feeding on common grasses by supplementing

which is deficient

I spec-

and physical size.

tended to be obscured by an overstory

related to their spa-

In the verticle plane, forbs
of grasses in communities

where

grasses formed dense swards (aspen, wet shrub meadow, wet meadow, willow
and mesic meadow).
least visually,

As a result, forbs might be less easily detected,

than their prewinter

the grass canopy physically

biomass might suggest.

interfered with consumption

at

Further,

of forbs.

Forb

intake was greatest in communities

where the canopy of grasses was rela-

tively open (sagebrush,

and ponderosa

grassland,

pine-shrub).

�-340-

Horizontally

distributed,

forbs were more dilute in the environment

than browse or grass, and more evenly distributed
communities,

grass was ubiquitous:

than browse.

In most communities,

In all

browse, al-

though not necessarily

abundant, was highly visible and concentrated

patches in the habitat.

As a result of this patchy distribution,

once discovered,

could be consumed in large quanities.

in

browse,

In contrast,

forbs were neither overwhelmingly

abundant nor concentrated

Consequently,

of forbs required extensive searching

frequent consumption

in patches.

time.
Plant size may also have interfered with consumption
elk.

Individual

of forbs by

forb plants tended to be smaller than either grasses or

browses;

The size of plants and plant parts which an animal can in-

dividually

select is bounded by the animals mouthput size; large mouths

are limited to consumption
1973).

of relatively

large forage items (Hoffman

Many forbs were probably smaller than elk could individually

select from the surrounding

sward of other plants.

Diet Quality:

of Forage Quality

Interactions

and Diet Botanical Composition
Although elk diet quality was influenced by changes in forage quality, elk partially

compensated

for decrements

in forage quality by

shifting the botanic mix of their diets.
This compensation
nutritional

was possible because of the large differences

quality between grasses and shrubs.

leaves contained higher concentrations

in

Browse stems and fallen

of protein than senescent grasses.

Browse provided a better source of digestible protein than grasses
because true protein digestibility

of browses is greater than 90%

(Robbins et al. 1975:44, Short and Reagor 1970:965).

However, grasses

�-341-

contained more substrate for microbial
their relatively

than browse because of

small amounts of lignin and large percentages

lose and hemicellulose.
characteristics,

digestion

As a result of these differences

in fiber

grasses contained more IVDDM than browses.

IVDDM and digestible

Because

energy can be assumed to be approximately

(Moir et al. 1961, Rittenhouse

et al. 1971, Milchunas

equal

et al. 1978:19),

grasses were a better source of energy per unit of dry-matter
Apparently,

of cellu-

intake.

then, these two forage classes were complementary

foods, grasses provided a more completely digestable
and browse contained more protein.

These protein and energy character-

istics allow elk to vary the chemical composition
shifting the proportions

source of energy

of their diets by

of grass and browse which they consume.

I

observed that such shifts permitted my elk to maintain a relatively
stable diet quality, despite large variation

in forage quality among

habitats and between years.
Spatially,

diet quality was stabilized by selection of diets con-

taining similar proportions

of browses and grass in the different habi-

tat types.

Although the proportion

significantly

between habitats,

great as the differences

of browse in the diets differed

these differences

were not nearly so

in browse availability.

Because the animals

actively sought out browse in browse-poor

habitats,

the browse~grass

ratios of the diets were much more similar than the browse-grass

mix-

tures of the habitats themselves.

in

browse (grassland),

consumption

In the habitat most deficient
of a browse-like

greater than in any other vegetation
larities in botanical

composition

in diet chemical composition

type.

forb (sulfurflower)

As a result of these simi-

of diets among habitats,

were few.

was

differences

�-342-

Temporally,

the protein and cell wall content of diets changed

little between years in most habitats, despite large annual decrements
in forage protein and increments in forage cell wall.

Again this rela-

tive constancy can be attributed to changes in the grass-browse
of elk diets.

Frequency

mixture

of browse consumption was inversely correlated

with the crude-protein

content of grass (Fig. 24), suggesting that elk

responded to decrements

in grass protein by consuming more high-protein,

low cell wall browse.

Thus, during the first year, browse consumption

increased as winter progressed,

an increase that prevented diet protein

from declining as rapidly as the protein content of grasses.

Paradox-

ically, browse consumption was constant the subsequent year at about 40%
of observed bites throughout

the winter.

Consequently,

dietary protein

declined just as rapidly as the protein in grass.
Differences

in elk diet botanical

result from differences

composition between years might

in animal age; it is possible that yearling elk

select diets which differ from those chosen by calves.
have been observed for caribou (Bergerud 1970).
annual differences

Such differences

However, I suggest that

in diet composition were related to variation

protein content of forages during the two years.

in

Because protein con-

tents of browse and grass was very similar during November of the first
year, predominantly

grass diets were no less rich in protein than browse

diets and had more IVDDM.
and consistently

As winter progressed,

more browse was consumed.

grass protein declined

In November of Year 2, mean

protein content of grass was 20% less than that in browse and 25% lower
than that in grass of the previous year.

Thus, compared to Year 1, high

browse diets during November of Year 2 contained substantially
protein than predominantly

grass diets.

more

As a result of increased browse

�-343-

R2 =0.38

x

60

x

x

x
x

x

xx

50

x
x

P =0.00001
S.E. of Est. = 11.5

x

I-

w
0 40

x

z

~

w
(f)
30

5

~x

x

0:::

x

x

x
x
x

20

x

~x
x

0

rn

x

~

x
x

~

x

0

10

x

x
x

3.0

4.0

5.0

6.0

x CRUDE PROTEIN IN GRASS

Figure 24.

Relationships between browse' content of winter diets of 5
elk and mean crude protein content of grasses on upper
montane winter range, Colorado, during 1976-77 and 1977-78.

�-344-

consumption

throughout

the second year, diet protein content did not

decrease appreciably

in most habitats despite markedly

trations of protein

in forage.

I speculate

reduced concen-

that browse intake did not increase with advancing

season the second year, as it did the first, because digestion

of high-

browse diets is inefficient.

Diet IVDDM declined as the percentage

browse in the diet increased

(Fig. 25).

browse much above November

Presumably,

increases

of

in dietary

levels of the second year were too costly of

dietary digestibility.
Diet selection by elk apparently
stable dietary protein was maintained
variable diet digestibility.
" ... it is reasonable

involved a compromise:

between years at the expense of

Ammann et al. (1973:200) concluded,

to assume that a ruminant could not maintain

itself on a diet much below 50% digestible
universally

Relatively

less than 50% digestible

energy."

Elk diets were

during winter; however, had the

animals eaten no browse, diet IVDDM would have been close to, or exceeded, 50%.
What is the significance
and protein

resulting

of the tradeoff between diet digestibility

from increased browse consumption?

First, browse

protein could help maintain

a rumen environment

favorable to digestion

of unlignified

Rates of microbial

protein

cellulose

grass fiber.

digestion

dietary nitrogen
and Gilchrist

levels (Hume et al. 1970, Van Gylswyk 1970, Schwartz

1975).

grass.

and

in the rumen have been shown to be affected by

Hence, the relatively high protein content of

browses could contribute,
deficient

synthesis

in vivo, to increased digestion

Such associative

of protein-

effects have been widely reported

(Elliot and Topps 1963, Dietz 1967:136, Egan and Moir 1965, Milchunas
al. 1978:11).

et

�-345-

•

•

50

xx

x
x

~
~

R2 =0.39

0
0

x xxx

P &lt; 0.00001

&gt;

S.E. of Est. =3.3

~

0

20
10

20
0/0

Figure 25.

30

BROWSE

40

50

60

70

IN DIET

Relationship between in vitro digestible dry matter (IVDDM)
and browse content of diets of 5 elk on upper montane winter
range, Colorado, during 1976-77 and 1977-78.

�-346-

Second, protein is particularly
condition during winter.

important for maintaining

animal

At the beginning of winter, elk have abundant

fat stores (Flook 1970:41-46) but no comparable "reserves" of amino
acids.

Consequently,

catabolism of fat results only in weight loss,

while negative protein balance must incur more immediate deleterious
physiological
viability,

effects, including loss of muscle strength, reduced fetal

and decreased resistance to disease (reviewed by Robinson

1977:15, Harper et al. 1977: 570-571).
deficiencies
animal.

Consequently,

dietary protein

may be more costly than energy deficits to the wintering

Thus, the energy penalty imposed by browse consumption

is

probably more than offset by its protein contribution.
Elk Nutritional

Status and Carrying Capacity

Could an elk have survived a winter period consuming diets of the
nutritional

quality I observed?

Predicted overwinter weight losses of

adult elk based on across habitat diets did not exceed 20% of prewinter
weight.

Decalesta

starvation

et al. (1975, 1977) found that mule deer died from

after losing 30% of their total body weight.

result, and the observation

that model predictions

Given this

of weight changes of

animals consuming poor quality diets tend to overestimate
(Torbit, in press), it is plausible

weight loss

that diets I observed would have

allowed adults to survive both winters of the study.

However, estimated

condition losses were much greater during the second year.

As a result

of the decrease in diet digestibility,

elk energy status was much less

favorable during Year 2 and predicted

losses of fat were more rapid than

for the previous year.
Across habitat estimates of weight losses of simulated calves were
similar to the annual and overwinter

trends in condition of adults

�-347-

except that weight losses were far more severe the second year.

These

predicted losses (30% of total body weight) had they been incurred by a
calf would probably have resulted in its death.

Remember, however, that

these weight loss estimates may be inflated, and that Year 2 diets useq
to drive the model were estimated from forage choices of yearling animals.

Caribou calves have been shown to select diets which differ from

those selected by their adult counterparts
animals have smaller mouthparts

(Bergerud 1972).

Since young

and lower total forage intake require-

ments than adults, they can be more selective in their food choices than
older, larger animals.

Consequently,

we might expect that a calf during

Year 2 would have selected a higher quality diet than I observed for
yearling animals, and as a result, might not have experienced

the extreme

losses of condition predicted by the model.
During both years, measured diets of elk were not nutritionally
adequate to meet predicted energy ~equirements.

However, simulated ni-

trogen balance was positive, or only slightly negative, except when lean
body was catabolized
these modelling

to meet energy demands.

Thus, I conclude from

results that elk protein requirements

did not greatly

exceed protein content of diets.
This conclusion is supported by several observations

on nitrogen

balance of ruminants .. From studies of deer and domestic ruminants,
McCullough

(1969:108) concluded that 7% protein diets are "adequate"

elk during winter.

for

Red deer (Cervus elaphus) maintained body weight on

diets containing 5% crude protein (Maloiy et al. 1970:844).
et al. (1975:74) concluded that apparent protein digestibility

Robbins
of browse

diets eaten by deer (Odocoileus sp.) would be negative when forages
contained less than 4.8% crude protein.

Mertins

(1973:41) estimated

�-348-

that for diets containing

40% digestible

carbohydrate,

rumen microbes

require approximately

6 g crude protein per 100 g of ingested dry matter

to ensure "adaquate"

fermentation

plausible

rates.

Based on these findings,

it is

that elk diets containing much less than 5% crude protein

would fail to meet the animals' metabolic
would also result in decreased

requirements

rates of carbohydrate

for protein and
digestion.

During

both years and in a variety of habitats, protein content of elk diets
was close to 5%.
Condition

estimates based on diets selected in the various habitats

were consistent with the relative uniformity
diets selected in those communities.

of nutritional

quality of

That is, habitats could not be

ranked on the basis of diet quality; similarly, when estimates of dietary
nitrogen and digestibility
was no predictable
status.

were integrated

ordering of habitats based on simulated nutritional

During both years, predicted

weight losses were smallest in the

aspen community.

However,

animal condition

showed strong interactions

among the other communities,

Despite these interactions,
habitat comparisons:
vorable,

in the ruminant model, there

of

between years.

one clear pattern emerged from the

in habitats where elk energy status was most fa-

elk nitrogen balance was most negative.

where diets contributed
duced insufficient

estimates

Conversely,

habitats

most to nitrogen status were those which pro-

energy.

In most instances,

across habitat estimates

of diets resulted in less rapid loss of condition

(lean body and fat)

than diets selected in any single habitat.
Similar to the findings of Wallmo et al. (1977), differences

in

predicted

animal condition

among habitats were often at variance with

predicted

carrying capacities based on total intake of forage and total

�-349-

biomass; That is, weight losses were often greatest where herbage biomass was largest.

The discrepancy

between estimates

based on forage quantity and estimates
forage quality illustrates
supportable

population

considered

simultaneously

of carrying capacity

of animal condition based on

that both individual

animal condition and

density are important attributes which must be
in evaluation

of elk habitat.

�-350-

Chapter IX
CONCLUSIONS
Theoretical

Implications

i) Optimal Foraging and Competition

Theory

A large body of literature suggests that the feeding patterns of
animals are predictable

(Rapport and Turner 1977, Pyke et al. 1977,

Schoner 1971, Ivlev 1961).

We can predict these patterns because a

species is thought to be uniquely adapted to capture, handle, and assimilate a specific range of food items.

This range defines the food dimen-

sion of the animal's niche (Hutchinson

1957).

The food niche has traditionally
petitive

interactions

1978).

Competition

been viewed as the outcome of com-

for limiting food resources

(Cody 1974a, Diamond

theory predicts that in a species packed environment

natural selection results in divergence of morphological
zation traits such that an individual's

diet choices are restricted to

those food items not consumed by other species.
cal constraints
phenoplastic

But, apart from anatomi-

on food choice, diet selection is probably influenced by

"decisions".

If the diet selection process is governed in

part by volition and responds to density independent
variation

resource utili-

temporal and spatial

in food resources rather than density dependent competition

for those resources, then shouldn't we expect diets to be relatively
labile and unpredictable?

Shouldn't diet selection be more exploratory

than habitual?
When resources are relatively abundant, selective pressure for
"optimum"

resource utilization

tion in those traits increases.

patterns relaxes, and phenotypic
Wiens (1977) concludes that:

varia-

�-351-

... many of the differences between coexisting species, which
may indeed have been adaptive under conditions of resource
competition, may become diluted during periods of resource
abundance.
At these times, departure from optimum predicted
by theory may be commonplace and competition may be relaxed or
absent. Close study, of course, will always reveal different
species to be, in fact different.
But it is wrong to attribute these differences indiscriminately to competitive pressures ....
I observed a large degree of variation
of elk winter forages and concomitant
tanical composition

of elk diets.

in the quality and quantity

variation

in the chemical and bo-

Thus, resource ultilization

patterns

were more variable than fixed and appeared to respond to changes in food
resources rather than past competitive

pressure.

Can we generalize

on

the nature and direction of shifts in elk diets?
Theory of optimal diets (Rapport and Turner 1977, Pyke et al. 1977,
Cody 1974b, Schoener 1971) predicts that as resources become abundant
relative to demands, individuals

should become increasingly

their diet choices, restricting

their consumption

which are most profitable
spent feeding.

selective

in

of food items to those

in terms of energy or nutrients

gained or time

Based on this theory, we expect that elk should have

been more selective during the first year than the second and that
during both years, selectivity

should have declined as winter progressed.

We expect these patterns because 1) forage quality and quantity was
greatest during Year 1, 2) we can assume forage biomass declined over'
the course of both winters due to forage consumption
vores and losses to litter, and 3) nutritional

by native herbi-

quality of the most

abundant forages (grasses) was observed to decline with advancing
Assuming that increased

frequency of browse consumption

equated with increased selectivity

season.

can be

(Fig. 9), then the observed patterns

of diet selection by elk were diametrically

opposed to the predictions

�-352-

of optimal foraging theory.
by relatively

ubiquitous

grasses, while the following year, the amount

of browse consumed increased
browse consumption

Diets during the first year were dominated

substantially

during Year 1 increased concurrently

the quality and (presumably)

the quantity of available

Shifts in the relative selectivity
Wiens'

in all habitats.

of diet quality and nutritional

tively common foods.
declined,

forage.

during periods of

When food resources were "abundant"

(relative to Year 2), non-selective

animal requirements

with declines in

of elk are consistent with

(1977) ideas on relaxed selection pressures

resource abundance.

Further,

during Year 1

feeding incurred no penalty in terms

status; at the beginning

of Year 1,

could be met by diets which were drawn from relaAs winter progressed

the quality of common grasses

this decline in resource quality required increased

tivity for browse to compensate

for reductions

selec-

in grass protein.

Elk became more selective when common foods became nutritionally
less adequate compared with foods which were more rare.
therefore,

It appears,

that elk diet choice can be strongly influenced by animal

requirements

relative to food resources.

When foods are abundant,

however, there may be little selective pressure to select the "best" of
those foods.
ii) Theory of Population

Regulation

How does the observed variation
quality and nutritional
animal numbers?
Predicted
that relatively

in elk forage resources,

diet

status fit current ideas on the regulation of

Several observations

are relevant here.

changes in elk nutritional

status demonstrated

clearly

small changes in the quality of elk diets could cause

large changes in animal condition.

Such variation

in individual

�-353-

nutritional
Predicted

status could substantially

perturb population

adult weight losses during year 2, had they been incurred by a

pregnant cow, would probably have prevented production
(Thorne et al. 1976).

the predictions

of a viable calf

Simulated weight losses of calves that year would

have resulted in their death.

not.

performance.

The important

of the model are absolutely

What is significant

is that variation

idea here is not whether
accurate;

they probably are

in food resources and its

effect on individual

animal condition can exert large influence on elk

population

That deficits in high quality food can affect

numbers.

ungulate population

dynamics has been demonstrated

in a variety of eco-

systems (Sinclair 1973, 1974a,b,c; Klein 1965, 1970; Caughley 1970;
Ellis 1970).
From a theoretical

standpoint,

these observations

concur with the

ideas of T. C. R. White (1978) who proposed that a ubiquitous
limitation

on the growth of animal population

food in the environment.

and potent

is a relative shortage of

For elk the relative shortage appeared to be a

lack of winter food items which provided adequate amounts of nitrogen
and digestible

dry matter.

quality interacted;
in digestiblity

As discussed,

these shortcomings

in diet

diets which were high in nitrogen tended to be low

and vice-versa.

The food shortage confronted by elk

during winter is relative rather than absolute because at the end of
winter a large proportion

of pre-winter

herbage biomass

remained, a

fraction of the food supply which could have been eaten and was not.
iii) Habitat Selection
This study did not investigate
based on comparison

habitat choices of elk.

of diet botanical

composition

However,

and nutritional

qual-

ity among habitats, we can infer that elk are capable of effectively

�-354-

utilizing a wide variety of habitat patches.
generalists

That elk can be habitat

is supported by two lines of evidence.

First, the increase in the proportion

of browse in diets which oc-

curred between years was observed in all habitat types.

That is, re-

gardless of the animals feeding location, the amount of browse it consumed increased and these increments were proportionately
the different habitat types.
independently
quirements

equal among

Thus, elk appeared to be using the habitats

and diet composition

appeared to respond to animal re-

rather than to differences

in the forage composition

of habi-

tats.
Second, elk were able to select diets of similar nutritional

quality

from habitats which were markedly different in plant species composition
and total biomass.

The quality of the diets selected in all habitats

was so similar that the habitats could not be individually
basis of diet quality.

ranked on the

Thus, elk were able to make efficient use of all

the habitats they encountered.
The capability of elk to be generalists

in their choice of habitats

resulted directly from their ability to be selective in their food
choices.

Theory predicts that habitat generalists

cialists

(Wiens .1976, Covich 1976).

should be food spe-

That is, if habitat patches are

used in a fine grained manner, food utilization

within habitats should

be relatively more coarse grained.

quality of elk diets was

relatively

Nutritional

stable over space because elk could be intensely selective

for higher protein forages (browses and sulfurflower).
selectivity,

This intense

where food items are consumed far more frequently than

their frequency in the environment,

is a good example of coarse grained

food selection which permits fine grained use of habitats.

�-355-

Management

Implications

i) Population

Management

The observed temporal variation
duction has important implications

in forage quality and herbage profor management

of elk populations.

Plant biomass in all habitat types appeared to be strongly influenced by
annual variation

in precipitation.

but in terms of management,
vegetation

its significance

types total production

with the first.

This is hardly a surprising
is substantial.

result,

In some

was 50% less the second year compared

These changes reduced estimates

of habitat carrying

capacity based on total production.
Nutritional

quality of elk winter forages was quite labile within

seasons amd between years.
fluctuations

This variation

is consistent with annual

in browse protein reported by Dietz (1967:45), Pease et al.

(1979:58), and Gibbs (1978:137) and fits the pattern of overwinter
changes in grass quality observed by Willard and Schuster
et al. (1971), and Burzlaff
variation

in availability

fluctuations

(1971).

(1973), Sims

While it is widely recognized

that

of winter forages can result from year-to-year

in primary production' and snow cover (Severinghaus

1947,

Moen and Evans 1971, Gilbert et al. 1970, Wallmo and Gill 1972), my data
indicate that less-obvious

annual and within-season

quality of elk winter forage can substantially

changes in the

influence their food

resources.
It is plausible
potentially

that variation

affect elk. population

goals which include maintenance

in forage quality and quantity can

densities.

Consequently,

of elk populations

at stable densities

may not be realistic given these density independent
resources.

management

influences

on food

�-356-

ii) Habitat Management
From a nutritional

standpoint, elk were able to effectively

a wide variety of upper montane habitat types.
differences

utilize

Although there were

in predicted animal condition among habitats, these dif-

ferences were usually small.

When the differences

in elk nutritional

status among habitats were large (e.g. between willow and aspen during
Year 2) they tended to be compensatory.

That is, habitats where elk

energy status was most deficent, nitrogen balance
tion) was most positive, and vice versa.

(prior to fat deple-

Consequently,

predicted nutri-

tional status based on across habitat estimates of diet quality was
generally more favorable than predicted
habitat estimates of diet quality.
tory relationship,

I conclude, based on this compensa-

that winter habitat diversity is an important manage-

ment objective for maintenance
permits generalism

condition based on individual

of elk populations.

Habitat diversity

in habitat choice; habitat generalism facilitates

choice of diets which provide for different nutritional

requirements.

Within habitats, diversity of forage classes contributed
tenance of animal condition.

The ability of elk to vary the quality of

their diets depended on a gradient of forage digestibilities
contents.

to main-

and protein

This gradient occurred whenever the habitat contained a mix-

ture of grass and browse.

Thus, forage diversity,

diversity appeared to be significant
tives to wintering

as well as habitat

in providing nutritional

alterna-

elk.

It was clear from the modeling exercises that a very important
energy resource for elk during winter was fat reserves.

Depletion

of

fat stores before the end of winter always resulted in model predictions
of unaceptable

condition loss.

These fat stores are particularly

�-357-

important when winter food resources are temporally variable as I observed them to be.

Without a large "buffer" of stored energy, unpre-

dictable variation in food supply would have much greater effect on
nutritional

status than was observed.

Indeed, ever during nutritionally

"good" years like the first year of this study, some fat was required to
compensate for energy deficits during winter.

Consequently,

in managing

elk habitats, we can not ignore the fundamental importance of interactions between summer food supplies (which provide fat stores) and winter
foods (which provide a portion of overwinter maintenance
in providing for the year round nutritional

/

requirements)

status of elk.

�-358-

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R. B. GILL, L. H. CARPENTER, AND D. W. REICHART.
1973. Accuracy
of field estimates of deer food habits. J. Wildl. Manage. 37:
566-562.

WARD, A. L. 1970. Stomach content and fecal analysis: Methods of
forage identification.
Pages 146-158 in Range and wildlife habitat
evaluation:
A research symposium. U.~ Dep. Agr. Misc. Publ. No.
1147. 220 pp.
1971.
Wyoming.

In vitro digestibility

of elk winter forage in southern

J. Wildl. Manage. 35(4):681-688.

WEINS, J. A. 1976. Population responses to patchy environments.
Rev. Ecol. Syst. 7:81-120.
1977.
. 590-597.

On competition

in variable environments.

Annu.

Am. Sci. 65:

WESTOBY, M. 1974. An analysis of diet selection by large generalist
herbivores.
108:290-304.
WESTRA, R., AND R. HUDSON. 1979. Urea recycling in ~~apiti. In n.
Boyce, ed., North American elk: Ecology, behavior, and management.
Univ. Wyoming, Laramie.
(In press).
WHITE, T. C. R. 1978. The importance of a relative shortage of food in
animal ecology. Oecologia 33:71-86.
WILLARD, E., AND J. L. SCHUSTER.
1973. Chemical composition of six
great plains grasses as related to season and precipitation.
26:37-38.

---

WILLSON, A. D., W. C. WEIR, AND D. T. TORELL. 1971. Comparison of
methods of estimating the digestibility of range forage and browse.
J. Anim. Sci. 32:1046-1050.
WHITTAKER, R. H. 1970. The biochemical ecology of,higher plants.
Pages 43-70 in E. Sondheimer and J. B. Simeone, eds., Chemical
ecology. Academic Press, New York. 332 pp.
YOUNG, D. L., AND J. A. BAILEY. 1975. Effects of fire and mechanical
treatment on Cercocorpus montanus and Ribes cereum. J. Range
Manage. 28:495-497.

�July, 1979

-373-

JOB PROGRESS

REPORT

State of

C.:..O:.._:L.:..O:....:RAD=-'O
_

Proj ect No.

W-I26-R-2

Work Plan No.

3

Big Game Investigations
Job No.

~2~

Job Title __~E~I~k~P~o~p~u~l~a~t~i~o~n~a~n~d~E~c~o~l~o~gLy~S~t~u~d~i~e~s~
Period Covered:

_
_

July 1, 1978 - June 30, 1979

Personnel:
George D. Bear, *Ronald A. Green, Paul H. Neil, and Douglas
Cannalte, Rocky Mountain National Park personnel:
George Wagner, Jack
Gartner, Dan Davis, Charles Logan, Larry VanSlyke, Dave Stevens, Lorin
Casebeer, Jim Protto, Bert McLaren.

ABSTRACT
Eighty elk were trapped in three localities during a one-month period from
January 23 to February 22, 1979. Elk-sized Clover traps (Clover 195_) baited
with salt and alfalfa hay were used to capture the elk. Succinylcholine
chloride was used to tranquilize the larger animals for ease in handling.
Twenty-eight elk were marked with telemetry collars and twenty-two with
eartags at the Rocky Mountain National Park trap site; four elk were marked
with telemetry collars at each Masonville and Longmont Reservoir trap sites.
Elk at the latter two trap sites migrated westward to higher elevations within
Rocky Mountain National Park in the spring of the year. A portion of the
collared elk within Rocky Mountain National Park migrated west over the
Continental Divide to the Colorado River drainage; the remainder of the elk
moved to alpine ranges along the Continental
Divide.
Twenty-five elk calves were marked with telemetry collars containing mortality
sensors.
Two of these calves died when 2-3 weeks old. One was killed by a
bobcat and the other died from an unknown factor.
Calf:cow ratios appeared to
be very low on the study area; however, more data are needed to determine the
factors effecting this ratio.
Other mortality data were very minimal.
Investigations into the elk habitat selection and activity portion of this
program was primarily limited to testing equipment.
It was found that an
observer could accurately detect the activity of elk wearing an activity collar
94% of the time. The location error using a 7-element beam antennae was 11.2
(± 6.8) ha; which was considerably less than that for a 3-element antennae.

*Ronald A. Green is a graduate student enrolled at Colorado State University
working under contract to DOW to study elk activity patterns and relate them
to habitat selection.

�-374-

Only the activity data for May have been analyzed.
Information was collected
for two radio collared ~lk for a 24-hour period, and a third elk for only a
sunset sampling period.
A 24-hour activity budget indicated that the animals
were spending 55.8% of their time feeding, 36.8% resting, and 7.4% moving.
They were least active during daylight hours and most active at night.
The
coniferous type was used most (34.2%) over a 24-hour period with the aspen
type used 16.6% of the time.
Willow, wet meadow, and grassland types were
the primary feeding sites at night, while the coniferous type was most
important feeding site in daylight hours.
Feeding activity was more evenly
distributed throughout the habitat types at sunset and sunrise.
Willow and
grassland types were the two types used the most for resting at night, while
the coniferous type was used most for resting during the daylight period.

�-375-

ELK POPULATION

AND ECOLOGICAL

STUDIES

~eorge D. Bear

P. N. OBJECTIVES

1.

Develop techniques to more accurately
population levels.

and precisely

2.

Define natality

of selected elk populations.

and mortality

problems

estimate

elk

SEGMENT OBJECTIVES
1.

Construct

elk traps.

2.

Trap and mark elk on the winter range for distribution,
and habitat preference studies.

3.

Capture and mark elk calves for mortality

mortality,

studies.

METHODS AND MATERIALS
Trap Construction
Elk were to be captured with a portable elk-sized Clover trap (Clover 195_),
which is a larger version of the Clover traps commonly used to capture deer.
These traps consist of a frame made of 1.25 inch (OD) black pipe; with the
overall dimension of each trap being 4 feet wide, 8 feet long, and 6 feet
high. The pipe framework is covered with heavy 4 inch mesh nylon netting.
Twenty traps were constructed for use in this study.

Trapping

and Marking

Three trapping localities were selected for this study:
(a) Green Ridge,
approximately two miles southeast of Masonville;
(b) Longmont Reservoir,
approximately 5 miles east of Lyons; and (c) Rocky Mountain National Park
(Morraine Park and Horseshoe Parkin the eastern portion).
Major emphasis
was placed at the Rocky Mountain National Park site where 10 traps were
used, while two traps were placed at each of the other two sites.
The trap sites were prebaited with a 50-lb block of livestock salt and
alfalfa hay. After elk started visiting the areas traps were hauled in
and erected.
Since trap sites within Rocky Mountain National Park were
situated away from roads, a helicopter was used to ferry traps in and out.
It only required 2-4 days for elk to become accustomed to traps. Trapping
was conducted only for a week at the Longmont Reservoir and Masonville sites,
and for a month (January 23-February 22) in Rocky Mountain National Park.
Traps were baited with good quality leafy-alfalfa.

�-376-

Generally a crew of 4-6 persons were used to handle elk in traps.
In the
early stages of trapping all elk were tranquilized with succinycholine
chloride; however it was found animals could be cleared from traps faster
if calves and yearlings were manually handled and larger elk were
tranquilized before handl~ng.
It was also found a blindfold placed over
the subdued elk's eyes further reduced struggling.
A Pneu-dart cartridge-powered
rifle and darts were used to administer the
tranquilizing drug.
The darts are plastic projectiles carrying pre-measured
amounts of powdered succinychlorine chloride.
Since the work was being
done in sub-zero temperatures (January-February)
these darts with pre-measured
powdered drug were easier to work with than liquid-filled darts.
Captured elk were marked with a telemetry collar or eartags.
The collars
were adjustable telemetry collars prepared for big game animals by Telonics,
Mesa, Arizona.
They are an hermetically sealed unit, with magnetic switching,
and a lithium battery with 3-4 year life expectancy; weighing approximately
550 grams.
These collars were green in color.
Thirty-two of the collars were
equipped with mortality sensors.
Normal pulse rate was 550 ms, and 850 ms
in the mortality mode.
The mortality switch is activated if the elk does not
move its head (or collar) within a five-hour period.
Four of the collars
were equipped with the activity switch option.
Pulse rate of these collars
is 550 ms when the elk's head is up and 850 ms when the elk has its head down
feeding.
The receiver is a 200-channel unit (TR-2) with a scanner operating
on 148-149 MHZ frequency.
Each animal or collar is monitored on an individual
channel.
Elk not fitted with telemetry collars were marked with large (2-inch) livestock
eartags.
The tags were yellow with black numerals on them for individual
identification.
Duplicate tags were placed on each animal; one in each ear.
In past years visitors to the National Park objected to large collars placed
on elk, therefore, one objective of this segment was to investigate the
iei.,Sibility of using a less conspicuous marking device, such as ear tags , yet
one that will be readily detected by the research observer.

Locating

Marked Elk

Radio-collared elk were located 1-3 times a week, depending on accessibility.
Majority of the monitoring work was done from a vehicle, with occasional
foot travel or aircraft surveys when they migrated off winter range.
Several
types of antennas were used for tracking or locating collared elk. A monopole
whip antennae with a magnetic base was used when searching an area with a
vehicle or aircraft.
Once a signal was received one of the more directional
antennaes (loop, H-beam, 3-element beam, or 7-element beam) was used to
triangulate or locate the animal.
The latter antennaes were more accurate,
but also larger; thus, the type of antennae used depended on the situation
or condition of use.
Information gathered on eartagged
opportunities permitted.

elk was incidental

and obtained

whenever

�-377-

Marking

Elk Calves

Elk calves were located and captured either by searching calving grounds
on foot or with a helicopter.
Calves during the first few days after birth
generally avoided danger by lying down and remaining motionless.
Then as
they got older they tended to jump up and run if they suspected they had
been detected.
Therefore, only calves 1-3 days old could be caught by a
person on foot. By using a helicopter a researcher could capture calves
until they were 20-25 days old. When a calf was first observed the pilot
separated it from the cow or other adults.
Then the helicopter hovered
20-30 feet directly over the calf, this would generally stimulate it to
try to hide.
Once the calf laid down, the helicopter would land nearby
permitting a person(s) to get out. With older calves it was necessary for
the helicopter to return to a position above the calf to divert its attention
away from a capturer until he could get close enough to catch the calf.
The calf was blindfolded and placed in a large cotton bag. It was sexed,
weighed (to the nearest pound), and the following measurements were recorded
(to the nearest centimeter):
total body length, shoulder height, and girth.
Age of the calf was estimated based on hoof coloration, total body coordination,
body size, and other characteristics of general physical appearance.
A
telemetry collar was then placed on its neck.
Calf telemetry collars consisted of the telemetry unit made by Telonics
attached to a two-inch collar.
The collar was made from a two-inch cotton
firehose with the telemetry unit attached to it with pop-rivets, and the
ribbon antennae inserted inside the hose for protection.
Overall diameter
of the collar was reduced by folding the collar and fastening each fold with
elastic, thus forming an expanding collar that would expand as the calf grew.
Each of these collars contained a mortality sensor as in the telemetry units
placed on larger animals trapped last winter.
Each calf was located 3-4 times a week to determine if it was still alive and
moving.
The same equipment described earlier was used to locate calves.

Activity

Radio~Collars

Of the 28 radio-collars, four collars were equipped with activity switches.
When these collars are displaced forward from a vertical position as in a
feeding activity, a tip switch causes the transmitter to emit a higher pulse
rate. By monitoring the change in pulse rate and signal strength, feeding,
bedding and moving activities can be discriminated.
To test accuracy and reliability of these collars to accurately detect
activity patterns, an activity collar was placed on a tame elk at the
Colorado Division of Wildlife's wild animal research facilities at the Colorado
State University Foothills Campus.
During one-minute intervals, one person
would record activity pattern of elk from visually observations while a second
person would record activity patterns indicated by telemetry signals.

�-378-

Table 1.
January,

Elk trapped and ear tagged in Rocky Mountain
1979.

National

Park,

Sex

Est. Age
(years)

Est. Wt.
(lbs)

Body
Condition

Drug
Dosage (mg)

Time For
Restraint
(min)

Restraint
Time
(min)

1

F

Calf

250

Good

10

5

41

2

M

Calf

250

Good

10

5

38

3

F

1

400

Good

17(5)

36

27

4

F

Calf

250

Good

10

10

19

5

M

1

350

Good

17

Partial

6

F

Calf

200

Good

10

5

19

7

F

Calf

200

Good

None

8

F

Calf

200

Good

None

9

F

Calf

Good

None

10

M

Calf

200

Good

None

11

F

Calf

200

Good

None

12

F

Calf

200

Good

None

13

F

Calf

200

Good

None

14

M

Calf

200

Good

None

15

F

Calf

200

Good

None

16

M

Calf

200

Good

None

17

F

Calf

200

Good

None

18

F

Calf

225

Good

None

19

F

Calf

200

Good

None

20

M

1

350

Poor

15

4

44

21

M

Calf

200

Good

None

22

F

Calf

250

Good

None

Eartag
Number

�-379-

Table 2.

Elk trapped and collared in January, 1979.

Est. Age Est. wt.
(years)
(lbs)

Body
Condition

Time For
Drug
Restraint
Dosage (mg)
(min)

Restraint
Time
(min)

Collar
Number

Sex

10

M

1

400

Good

17

8

20

20

F

2

400

Good

17

6

24

30

F

2

400

Good

None

40

F

5

600

Good

17

9

58

50

F

1

300

Good

None

60

M

1+

450

Fair

17

5

33

70

F

1

400

Good

15

4

6

80

F

2+

400

Poor

None

90

M

7+

700

Good

26

7

64

100

F

2

400

Fair

None

110

F

4

450

Fair

17

4

55

120

F

3

450

Good

*
20(5)

22

21

130

F

1

300

Fair

None

140

F

1

350

Good

None

150

F

2

375

Fair

None

170

F

2+

450

Fair

17

2

34

180

F

2+

450

Good

20

Partial

200

F

1

300

Fair

None

210

F

2+

450

Fair

17(5)*

Partial

220

F

1

350

Good

17

Partial

260

F

3+

500

Fair

17

15

270

F

+
3

450

Good

None

------------------------------------------------------------------------------

2

�-380-

Table 2.

Collar
Number

Elk trapped and collared

in January,

1979.

(Continued).
Time For
Restraint
(min)

Restraint
Time
(min)

Sex

Est. Age
(years)

Est. wt.
(lbs)

Body
Condition

Drug
Dosage (mg)

280

F

1

350

Good

None

290

F

1

300

Good

None

300

M

2

400

Good

17

9

320

F

2+

450

Good

17

Partial

330

F

2

450

Good

15

Partial

340

M

1

450

Good

20

2

350

F

3

Good

None

360

F

3

450

Fair

17 (5)(5)

370

F

2

450

Good

None

380

F

2

350

Good

10(5)

26

5

None

M

+
5

650

Fair

20

15

16

600

F

Calf

250

Good

10

3

10

750

F

3+

500

Good

20

11

35

850

F

3+

500

Good

None

900

F

2

400

Good

20

5

42

* Second dosage required.

11

58

Partial

�-381-

Triangulation

Accuracy

To measure elk location measurement by accurately using triangulation estimating
methods, one person placed radio-collars throughout the study area and
marked these locations on black and white aerial photographs (Scale 1:15840)
while a second person recorded three bearings from different preselected
positions using a directional antennae.
Triangulated locations were
determined by plotting the three bearings on aerial photographs and determining
the center of the resultant triangle
formed.
The error of location was
calculated by first measuring the distance between the triangulated location
and true location and using this distance was used as the radius to scribe
a circle with the triangulated location as the center.
The area of this circle
was used as the measure of triangulation error.

Habitat

Selection

and Activity

Measurements

Pre-selected points throughout the study area were used to determine elk
locations by triangulation.
For each confirmed location; time of day, radiocollar frequency, bearings and habitat type were recorded.
This procedure
was conducted on a 24-hour basis with the 24-hour period divided into four
sampling intervals.
The intervals will be:
(1) 2 hours before and after
sunset and sunrise, respectively (2) daylight hours and (3) nighttime, and
(4) 2 hours before and after sunrise.
To collect activity data, one sampling period of 3 consecutive days will be
conducted per month by monitoring of the activity collars around the clock
with the aid of a second person.

RESULTS AND DISCUSSION
Trapping

and Marking

Eighty elk were trapped during a one-month period, January 23-February
22, for an overall trapping success of 61 percent.
Twenty-two (14 calves,
7 yearling males, and 1 adult bull) were released unmarKed once the
desired quota of that sex and age class was obtained.
Twenty-two elk
(20 calves and 2 yearlings) were marked with eartags in Rocky Mountain
National Park (Table 1). Twenty-eight elk (18 adult cows, 6 yearling cows,
2 adult bulls, and 2 yearling bulls) in RMNP were fitted with telemetry
.collars.
Four of the collars had activity switches and 24 had mortality
sensors or switches.
The four activity collars were placed on adult cows.
Four telemetry collars were placed on elk (1 calf, and 3 adult cows) at
Masonville; four collars on elk (2 adult cows, 1 yearling cow, and 1
yearling bull) at the Longmont Reservoir trap site (Table 2). Two cows
which were captured at Masonville and collared in February 1978 still had
active collars.
Trapping and handling of elk was very successful.
Of the 80 elk trapped
only 1 cow (a yearling) died.
This cow had received 5 mg dosage of succinyl,
however, other elk of this size and age were successfully tranquilized with
a 17 mg dosage.
A field necropsy did not reveal any injuries or abnormal
conditions, therefore, death was attributed to excitability of this particular
animal and stress of being handled.

�-382-

The mean dosage of succinyl for tranquilized elk was .040 + .005 mg/lb
body weight.
This was greater than the dosage (.034 mg/lb) recommended
by Boyd (1968)*. However, Boyd was working with penned elk; also,
weights we recorded for elk were estimates not measured weights (Table 2).
A rule-of-thumb we found to be useful during the trapping period was: 10
mg dosage for calves, 17 mg for yearlings, and 20 mg for adult cows. Of
course those dosages were varied if the animal varied from the normal
(i.e. very poor body condition, very excitable, an exceptionally large
yearling bull, etc.). Mean reaction time or time for drug to take effect
was 9 ± 8 minutes, and mean recovery time 30 + 18 minutes.
Obviously there
was considerable variation in individual
reaction to the drug. Because
some elk were so slow to recover from the drug, it was necessary to leave
one person to "baby-sit" the elk until it recovered, while the rest of the
crew continued working with other animals.
Therefore, it was often faster
to manually handle smaller animals than to tranquilize them.

Population

Distribution

The five collared elk at Masonville remained on Green Ridge throughout the
winter.
Then in June they migrated to the Storm Mountain area; remained
there for 10-14 days, then four of them migrated to the alpine range in
northeastern Rocky Mountain National Park (Fig. 1). The fifth cow remained
at Storm Mountain.
Two cows were collared on Green Ridge in February, 1978;
they had followed a similar time schedule and migration pattern in the spring
of 1978. The time of migration seems to be related to snow melt and vegetation
"green-up" at higher elevations.
Migrations were very sudden, requiring only
2-3 days for each segment of the migration.
The four elk collared at Longmont Reservoir followed a similar migration
pattern.
Only this group migrated northwesterly to higher elevations in
J.a t e April.
They remained around a large meadow or park (Big Elk Meadows)
in w1 -ch new vegetation growth ("green-up") had occurred in early April.
Then as the snow melted at higher elevations in early June they migrated
westward into Rocky Mountain National Park (Fig. 1).
The 28 elk telemetry collared in Rocky Mountain National Park remained on
their winter range in the easterly portion of the Park until spring.
There
was considerable movement back and forth among winter concentration sites in
Morraine Park, Beaver Meadows, and Horseshoe Park.
In late Mayas
snow
melted and corridors opened along ridgetops twelve elk migrated westerly
over the Continental Divide to the Colorado River headwaters area (Fig. 2).
There were still 3-6 feet of snow remaining in timbered areas and severe
spring snowstorms still swept higher elevations.
This group of elk settled
in green meadows along the Colorado River for the calving season.
It was
mid-to-1ate June before the other collared elk migrated up Forest Canyon to

*

Boyd, R. J. 1968. Experimental
Aid Rept., W-38-R.
pp. 223-236.

immobilization

techniques.

Colo. Fed.

�I

("")

co
("")
I

Corter
lake

.

..
~

N

5eola:

Figure 1.

Migration

routes of radio-collared

elk trapped at Masonville

and Lyons.

t

1 in. =1 mi.

�I
&lt;r

00
C"l

I

Carter
lake

.

.

.

~

N

j
!&gt;cale:

Figure 2.

Migration

routes of radio-collared

elk trapped in Rocky Mountain

National

Park.

I ir. -! ml

�-385-

to alpine areas along the Continental Divide, and up Fall River to alpine
and willow flats at the headwaters of Big South Fork of Cache la Poudre
River. By late June only four collared elk were still on winter range. At
that time very few elk were seen at these lower elevations.

Mortality
Little mortality occurred among radio collared elk during this segment.
Starvation appeared minimal; six adult cows, 1 yearling (#13 eartag),
and 2 calves (#2 eartag) were incidently found throughout winter months.
These deaths were assumed to be the result of starvation.
It would be
very difficult to numerically document such mortality in Rocky Mountain
National Park because predator appear to be numerous and known deer and
elk carcasses were quickly (2-7 days) dismantled and eaten.
The calf collared at Masonville in February 1978 died 1 month after it was
marked.
However, coyotes had dismembered the carcass before it was recovered.
The femur bone marrow was solid and white in color, the rumen was nearly
full; so it was concluded death was not caused by starvation.
The cow wearing telemetry collar #210 died during a severe spring snowstorm
in mid-June.
She was on winter range in Horseshoe Park on June 11, then on
June 14 she was found (via the mortality sensor) in a willow flat on the
headwaters of Cache la Poudre River.
This is a distance of 8 miles (direct
or airline miles) over alpine range traversed in a maximum of 4 days.
She
was lying on a grassy knoll between large snow fields.
She was extremely
thin, her pelage very rough looking, and the bone marrow was red and
gelatinous.
She had a large amount (approximately one quart) of puss-like
mucus in her uterus.
Her mammaries were filled with milk.
It appeared she
had developed an infection in the reproductive tract after giving birth;
and because she was in a starved condition was unable to recover from
additional stresses.
On June 20 another adult cow (5-6 years old) was found dead in the Horseshoe
Park vicinity.
She was lying in a grassy aspen glen a few feet from a
small stream.
She had molted winter pellage and appeared to be in excellent
body condition.
The only abnormal condition discovered in a field necropsy
was about a pint of green colored mucous in the uterus.
She had milk in
her mammaries.
It was surmised a reproductive disorder may have contributed
to her death, also.
Twenty-five calves were captured and collared during the calving season
(Table 3). Only one calf did not appear to be in excellent health.
It
appeared to have a mild case of diarrhea, but must have recovered, since
it is still alive and thriving.
One of the collared calves was killed by
a bobcat during a spring storm in which it snowed 12-15 inches.
It ate
the hindquarters and viscera, then buried the front portion of the carcass
in some debris.
The calf was approximately 2-3 weeks old. Another collared
calf died or was killed, however, coyotes devoured most of the carcass before
it could be found and examined.
This calf also was approximately two weeks
old.

�-386-

Table 3. Elk calves collared in Rocky Mountain National Park during
May-June, 1979.
Collar
Number

Date
Tagged

Sex

Weight
(lbs)

Shoulder
Ht (cm)

Estimated
Girth
Total
(cm) Length (cm) Age (days)

610

6-18

M

30

65

56

102

1

620

6-22

M

35

63

57

96

2

630

6-28

M

69

81

81

114

21+

650

5-31

F

45

660

6-28

M

38

63

66

86

5

670

6-26

F

51

71

65

105

10

680

6-28

F

68

85

86

120

21+

690

6-28

M

58

75

71

112

14

700

6-26

F

55

63

76

118

14

710

6-28

F

40

63

62

99

7

720

6-28

M

68

78

81

122

21

730

6-26

M

38

71

62

111

5

740

6-19

M

67

72

87

118

21

750

6-22

F

33

63

58

101

2

760

6-26

F

72

71

77

140

21

770

6-6

M

50

71

67

109

10

780

5-31

F

790

6-28

F

49

71

70

121

10

800

6-26

F

54

66

76

119

14

810

6-26

M

56

63

72

118

14

820

6-28

M

67

80

81

130

21

830

6-28

F

45

76

71

117

14

840

5-31

M

10

1

2

-------------------------------------------------------------------------

�-387-

Table 3.
May-June,

Elk calves collared
1979.
(Continued).

in Rocky Mountain

Collar
Number

Date
Tagged

Sex

Weight
(lbs)

*680

6-4

M

28

**680

6-15

F

39

Shoulder
Ht (em)

National

Girth
(em)

Park during

Total
Length (em)

Estimated
Age (days)
2

62

62

4

103

* This calf died on June 8; collar reused June 15.
** This calf died on June 25; collar reused June 28.

One day during the calving season a cow was seen running about a clump of
willows in Beaver Meadows.
Upon closer inspection a large coyote came
running out of the willows carrying the front half of an elk calf.
The
calf was very small, probably less than a day old. It was speculated that
this calf also was a victim of coyote predation.
These were the only calf mortalities that could be documented.
All other
calves observed appeared to be healthy.
Calf:cow ratios obtained while
traveling about the area were low even late in calving season when the
calves should be traveling with cows and therefore more visible (Table 4).
Table 4. Elk observations

at weekly

intervals

during the calving

season.

Number Observed
Marked
Elk

Date

Cows

May 28-31

124

3

(2)

June 4-9

386

10

(3)

122

4

June 11-16

165

18

(11)

29

3

June 18-23

337

38

(11)

89

11

June 25-30

197

33

(17)

34

6

(Calves/100

cows)

Bulls

2

�-388-

General Comments
This research project is only in the initial stages, much more data are
needed to elucidate factors contributing to low calf:cow ratios in" this
herd.
These data will be gathered in future research efforts of this
project.
Now that trapping and marking techniques have been tested more
emphasis will be placed on estimates of population size during the next
segment.
The eartags were readily seen by observers on the ground and
in aircraft, so they will be used in the marking program for the population
study. Also more data will be obtained concerning distribution and movements
and habitat selection of this elk population.

Triangulation

Accuracy

Considerably more time was spent in testing the accuracy of the triangulation
system than first anticipated.
The accuracy of the system was originally
tested using a 3-element antenna utilizing three azimuths to determine a
location.
The location error was calculated as a mean circular area
surrounding the true location of the telemetry collar with 90% confidence
limits.
The radius of the circles surrounding each location was the linear
distance between the true location and determined location.
The 3-element antenna system proved dissatisfactory resulting in a mean
location error of 62.5 (± 28.7) hectares.
Due to the small areas of some
habitat types and considerable interspersion in certain regions, it was
felt that an error this large was unacceptable.
To remedy this problem,
the 3-element antenna was replaced with a 7-element antenna in hopes of
obtaining greater directional accuracy within the system.
Again three
azimuths were used to determine a location.
After 6 locations it was
discovered that reading the compass with the radioreceiver headphones on
was causing errors in azimuth readings by as much as 10 to 15 degrees.
All
previous locations with the 3-element antenna had also been determined by
reading the compass with headphones on. The remaining 10 locations were
then determined by always removing the headphones first.
This significantly
(a = .10, P = .009) reduced the mean location error from 24.3 (+ 5.8) ha
to 11.2 (± 6.8)ha.
A mean location error of 11.2 (± 6.8) ha was felt to be an acceptable
error if the habitat types were defined such that all types had areas
greater than the lower 90% confidence limit of the mean location error
(4.4 ha). Types smaller than this were included in the surrounding habitat
type.
It soon became apparent after a day of determining elk locations within
habitats, that using three azimuths to locate an animal was much too
slow. It was felt that an insufficient number of locations (one per hour)
were being obtained to calculate a reasonable estimate of habitat selection.
It was decided to use only two azimuths vs. three to locate an animal.
This increased the locations recorded per hour by 2.5-4 times depending on
distance between triangulation points and weather conditions.
A reassessment
of the location error was done to determine if the accuracy using two
azimuths was significantly different from the accuracy with three azimuths.

�-389-

Using the 10 locations previously determined with the 7-element antenna
and three azimuths, 3 combinations of two azimuths each were formed from
each location.
Heezen and Tester (1967) reported that triangulations with
angels of 20 degrees and less or 160 degrees and greater generally produced
grossly inaccurate results.
My data on a limited number of sample points
(n = 10) suggested that this was true of angles less than 30 degrees.
Since
this angle can be somewhat controlled under field conditions, locations
determined by a triangulation angle of less than 30 degrees were separated
from the others and analyzed separately.
The two azimuth triangulations
produced a mean location error of 11.9 (± 5.4) ha compared to 11.2 (± 6.8)
ha for the three azimuth triangulations.
A t-test (a = .10) suggested that
these means were not significantly different (P =.50).
The two azimuth
triangulation with a 7-element antenna is the system currently being used
to collect habitat selection data.
A t-test was run between the two azimuth triangulations with an angle less
than 30 degrees and those with an angle greater than 300 to test whether
triangulation angle had an effect on the accuracy of location.
A significant
difference (a = .10, P = .05) was found between the two. Care must be
taken in the field when selecting monitoring points so an adequate triangulation
angle is maintained.
It was not only important to locate the animal within certain area limits of
the true location but also to correctly place that animal's location into
the correct habitat type. For each location, the actual habitat type and
the habitat type for the triangulated location was recorded.
A Chi-square
goodness-of-fit test was run to determine whether any difference existed
between observed and expected habitat types. There were no significant
differences (a = .10, P = .25 and P = .51, respectively) between the two
azimuth and three azimuth triangulations.
The two azimuth method located
elk in correct habitat types 70% (± 5.7) of the time vs. 80% (± 10.1%) of
the time for the three azimuth method.

Activity

Collars

Time did not permit collection of any additional data.
It was previously
reported that elk activities were correctly detected by the telemetry
equipment and operator 94% of the time. A Chi-square goodness-of-fit
test was run to see if differences occurred between observed and expected
activities.
No significant difference ~ = .10, P = .64) was apparent.
The 90% C.I. was 94 + 2.1%.

Habitat

Selection

and Activity

Data

Habitat selection and activity data were obtained during May and June.
Analysis of the May data are nearly completed, and the June data are
currently being analyzed.

�-390-

Habitat Types
After
types
taken
Baker

analysis of the triangulation accuracy data, the following habitat
were defined on the study area. Major species of these types were
from the biomass estimates and species composition reported by
and Hobbs (1978).

1.

Wet meadow (WTMD) - major species are Carex spp., Ca1amagrostis
canadensis, G1yceria grandis, Ph1eum pratense, Juncus ba1ticus,
Deschampsia caespitosa, associated forbs and some Salix spp.

2.

Willow (WILU)- dominated primarily by Salix spp. in relatively
dense stands with an understory of Carex spp., Ca1amagrostis
canadensis, associated forbs, some Ph1eum pratense, and Poa spp.

3.

Coniferous (CONF) - dominated primarily by Pinus contorta and
Pseudotsuga menziesii.
Ranges from dense stands with virtually
no understory to broken areas with small patches of either GRSL or
WTMD.
Other areas are broken by small aspen patches or are old
aspen stands in late stages of succession.

4.

Aspen (ASPN) - dominated by Populus tremu10ides in stands generally
defined greater than 10-12 hectares.
Understory of primarily of
forbs, Poa spp., Phleum pratense, Ca1amagrostis canadensis, Bromus
inermis and Rosa spp.

5.

Grassland (GRSL) - drier sites dominated by Muh1enbergia montana,
Danthonia parryi, Bouteloua gracilis, Stipa comata, Potentilla spp.,
Koeleria cristata, Carex heliophila, Erogonum umbellatum, Agropyron
spp., and associated forbs.

6.

Ponderosa pine-grassland
(PPGS) - primarily a grassland
associated parklike overstory of Pinus ponderosa.

7.

Ponderosa pine-shrub (PPSH) - this type ranges from areas dominated
by Pinus ponderosa with a shrub understory of Purshia tridentata,
Artemisia tridentata and Chrysopsis villosa to open areas dominated
primarily by Artemisia tridentat~.
Other species associated with
this type are Agropyron spp., Muhlenbergia montana, Stipa comata,
Bout~loua gracilis, Carex heliophila, Koeleria cristata, and forbs.

8.

Willow-park (WLPK) - (elevation 3050 - 3355 m) This type is a mixture
of mesic sites, wet marshes and xeric upland sites. Dominant species
are Salix spp., Betula glandulosa, Carex spp., Calamagrostis canadensis,
Deschampsia caespitosa and Caltha leptosepala.

9.

Krumholz (KRHZ) - (elevation 3355 - 3660 m) This type is an ecotone
between the subalpine forest and alpine tundra.
Salix spp. dominate
and are interspersed by Abies lasiocarpa and Picea engelmannii.
Deschampsia caespitosa is characteristic of the understory.

type with an

�-391-

10.

Alpine Tundra (ALPN) - (elevation 3660+ m) This region varies from
rolling terrain to sharp peaks and ridges.
Vegetation is dominated
by Kobresia meadow.
Other species are: Deschampsia caespitosa,
Silene acaulis, Trifolium spp., and Carex spp.

11.

Subalpine forest (SPFR) - (elevation similar to WLPK) - Dominated
primarily by Abies lasiocarpa and Picea engelmannii.
Associated with
the WLPK type. In its lower elevational range it grades into the CONF
type and into the ALPN and KRHZ types in its upper elevational range.

Activity

Patterns

Activity patterns to be recorded were divided into three categories:
feeding, resting and moving.
A feeding pattern was one in which feeding
was the dominate activity.
It may include such other activities such as
standing and walking, therefore, a minimum of 5 minutes was usually spent
monitoring the telemetry signal so the dominating activity could be
detected.
Resting was defined when the animal was immobile whether in a
standing or bedding posture.
Moving was defined as walking or running.

Habitat

Selection

and Activity

Data for May

Activity data were collected from two radio collared elk during one 24-hour
period.
Animal 360 provided data for 75% of a second daylight sampling
period in addition to the one complete 24-hour period.
A third elk (380)
contributed activity data for one sunset sampling period.
The data were
analyzed by individual elk and across sampling intervals and for 24-hour
periods across the month.
A 24-hour activity budget across elk indicated that animals were spending
55.8% of their time feeding, 36.8% resting and 7.4% moving (Fig. 3). The
daylight interval indicated that animals were least active during this time
with 52.3% of the time spent resting.
Feeding occupied 38.7% of the daylight
period and 9% moving (Fig. 4). It appeared from the interspersion of feeding
and resting activities that animals rest for only short periods at a time.
The sunset period was by far the most active feeding period with 90.3% of the
time occupied by this activity.
Equal amounts of time (4.2%) were spent
resting and moving.
Animals were quite active through out the night with
only 26.7% of the time spent resting.
They feed for 68.3% of the time through
the night.
Feeding activitiy was reduced to approximately 50% during the
sunrise interval with most of the feeding done before and about an hour after
sunrise.
Resting consumed 41.7% of the time with most occurring after
sunrise.
Some individual variation did occur between animals in activity patterns
(Fig. 5-8). Across the 24-hour period, elk 360 and 370 feed in nearly
equal amounts.
Elk 360 moved about much more than 370 and rested 12%
less. Both elk exhibited the pattern of resting the most during daylight
hours and feeding most intensively during sunset hours and slowly decreasing
feeding intensity as night hours passed and sunrise approached.

�-392-

24-HOUR ACTIVITY BUDGET, MAY 1979

{

36.8%
RESTING

Figure

55.8;6
FEEDING

3. Mean activity percentages across elk across the 24-hour
period during May, 1979, Rocky Mountain National Park.

�DA~~IGHT*
10.33 hours

NIGHT-5.33 hours

33.7%

52.3%

26.7%

58.3%

Feedin~

Resting

Resting

Feeding

I

C")
0\
C")

I

50%
Feeding

90.3%
Feed.ing

41. '7%
Resting

Fi::ure 4. Mean activity percentages across elk across the 24-hour sampling intervals
during !·~ay1979, Rocky !·~ountain Na:.ional Park.

�-394-

24-HOUR ACTIVITY BUDGET, lAAY 1979

BLK360

29. 5)~
RESTING

Figure

56.3~
FEEDING

5. fiiean activity
percentages
for elk 360 a cro s s the
period May 1979, Rocky Mountain National
Park.

24-hour

�-395-

24-HOUR ACTIVITY BUDG~T, MAY 1979
ELK 3'to

41.5fo
RESTINI}

56. 91~
FBEDING

Figure

6. Mean activity percentages for elk 370 across the 24-hour
periqd May 1979, Rocky Mountain National Park.

�JAY;_, L(_j·;e.'-

10.33

hours

NIGliT- 5.33 hours

/...----r·------ ·,
~5
/

. 9 .~

-.~.-T---.~-_ .-....~.

"",-

(

~Iovin€~

3°~~
Feeding
)

,-'
14
,:')
4:J.

.J

(

Pestine;

\

\

26.7);

63.3%

\ Pesting

Feeding

\

-.

,+,.;,2:'0.'

I

-o

ResNnr,.~
~

0'1
CV)

/'

~_.._

__ -.-

.-:"
,,/,

---....___ ..•..
".

I

16. 7!'o

//

Hoving

!

I

\\"

Peedine

rHS !~.L
",m

Q
••J&lt;)!'".

F i;,:ure

7.

I

(1?
')-',._;'
U_; • ..) ..'')

( ~~
l hours

)

33.3)~
Pc' s t '1 ncr
-..

•

--

(j

SmH7I'3I:::

~1T.eanac t:t vi ty perc ('n ta[':e
s for e 1;.: 360 aero S3 the 2L~-hour samplinr;
intervals
during
I'~ay 1979, Rocky rfoountain
Na t Lona L Park.

50;~
Feeding

(4

hours)

�.!)AYL ~;:i.Tw. 10.33

hours

HIG.flT-

I

I

I

!

r

38.3/

r
,I
I

F'e e d l ng

.50..3) hours

.

I

---.~

/'
,/

/
(26.7,(;
, Resting

59 (..,_,f
.•

0/,)

I
I

:P.e3tine

73.yi
Peeding

~

\
I

r-,

a-

--'__

~-

M

'

I

91.7i;
Feedine

S UlJ S l1T (4 hour s )
Fieur'e

8.

50:;&amp;
Resting

50:'s
Feeding

sun ;~IS};: (l.t he ur s )

l':ean activity
percentages
for elk 370 across
the 24-hour
in torvals
dur ing ~.~fJ.y1979, Rocky l'1.oun tain Natl onn I Parka

sampling

�-398-

Habitat selection data (Table 5) indicated that animals used the coniferous
type most over a 24-hour period (34.2%) with greatest use during daylight
hours and least during night.
Aspen, the second most frequently used type
(16.6%) was utilized in nearly equal amounts during daylight and sunset
hours.
Wet meadow, willow, grassland and ponderosa pine-grassland types
were utilized in nearly equal amounts (11%, 12.4%, 10.3% and 11.2%,
respectively) across the 24-hour period.
All four types were used most
during sunset hours through early morning hours and used least during
daylight hours.
Of the seven types utilized across the 24-hour period,
ponderosa-pine
shrub was least used, being used only during night and
early morning.

Table 5. Mean habitat selection percentages across animals by 24-hour
segment and across the 24-hour period, May 1979, Rocky Mountain National
Habitat

Type

Daylight

Night

Sunrise

Sunset

24-Hour Period

WTMD

0%

23.3%

5.5%

29.2%

11 %

WILW

0

26.7

22.2

16.7

12.4

CONF

64

0

22.1

12.5

34.2

ASPN

21.7

6.7

11. 1

20.8

16.6

GRSL

6.3

33.7

0

0

10.3

PPGS

6.8

0

27.8

20.9

11. 2

PPSH

0

9.9

11.1

0

4.2

Park.

During the daylight segment, activity was greatest in the coniferous type
with feeding (27.7%) and resting (31.7%) activities constituting the
majority of the total activity.
Aspen was used approximately a quarter
of the daylight time (21.7%) with resting (12.3%) being the predominate
activity and feeding (8.1%) second (Tables 6-10).
Feeding was clearly the dominate activity during the sunset interval with
wet meadow (25%) and ponderosa pine-grassland
(20.9%) types being utilized
most. Willow, coniferous and aspen types were also used considerably for
feeding during this time. Feeding during night was primarily associated
with wet meadow, willow and grassland types, 20%, 15%, and 18.3%, respectively.
Willow and grassland types were the two types used most for resting at night.
As previously mentioned, resting periods appeared to be quite interspersed
among other activities.
During early morning feeding was associated primarily
with willow and ponderosa-pine grassland types. Coniferous, aspen, ponderosa

�-399-

pine-grassland and ponderosa
during early morning hours.

pine-shrub

were all utilized

for resting

No obvious reactions to weather conditions were observable.
Temperature
appeared to have little effect on activity or selection of habitat type.
Elk used open meadow types through the night whether the temperature was
close to freezing or considerably warmer.
There were not enough windy
days to establish any type of behavioral response to wind. Elk were observed
feeding during a heavy rainstorm in open meadow types. There was no wind
or lightning associated with the storm and animals appeared to pay no
attention to the rain.

LITERATURE

CITED

Baker, D. L., and N. T. Hobbs.
1978. Simulations of the carrying capacity
of the Rocky Mountain National Park elk winter range.
Colo. Div. Wildl.
Fed. Aid. W-38-R-32, WP 17, Job 2. Prog. Rep., Game Res. Rep. July 1978.
Heezen, K. L., and J. R. Tester.
1967.
triangulation with special reference
Manage. 31(1):124-141.

Evaluation of radio-tracking by
to deer movements.
J. Wildl.

Boyd, R. J. 1968. Experimental immobilization techniques.
Colo. Dept.
Game, Fish and Parks.
Game Res. Rep. July Part 3 :223 - 236.
Clover, M. R. 1956.
Game. 42:199-201.

Prepared

;..lJ .'/ .A

by:

Single gate trap for deer.

/.,

/"/

6" fi.··· ....r

G~;;::if.:·::a~.~
Wildlife

Researcher

.~.&lt;te£N~
Ronald Gr en
Graduate Student

~&amp;,~

(

Calif. Fish and

�-400-

Table 6. Activity percentages by habitat type across animals across the
daylight sampling interval, May 1979, Rocky Mountain National Park.
Habitat

Type

Feeding

Resting

Moving

0%

0%

0%

CONF

27.7

31

5.3

ASPN

8.1

12.3

1.3

GRSL

3.1

1.9

2.3

PPGS

o

6.8

o

WTMD
WILW

PPSH

Table 7. Activity percentages by habitat type across animals across the
night sampling interval, May 1979, Rocky Mountain National Park.
Habitat

Type

Feeding

Resting

Moving

WTMD

20~~

3.3%

0%

WILW

15%

11. 7%

0

CONF

0

0

0

ASPN

6.7

0

0

GRSL

18.3

10

5

PPGS

0

0

0

PPSH

8.3

1.7

0

�-401-

Table 8. Activity percentages by habitat type across animals across
the sunset sampling interval, May 1979, Rocky Mountain National Park.
Habitat

Type

Feeding

Resting

Moving

WTMD

25%

0%

4.2%

WILW

14.6

2.1

0

CONF

12.5

0

0

ASPN

14.6

2.1

4.1

GRSL

0

0

0

PPGS

20.9

0

0

PPSH

Table 9. Activity percentages by habitat type across animals across
Sunrise sampling interval, May 1979, Rocky Mountain National Park.
Habitat

Type

Feeding

Resting

Moving

WTMD

2.7%

0%

2.7%

WILW

19.5

2.7

0

CONF

8.3

11.1

2.7

ASPN

0

8.3

2.7

GRSL

0

0

0

PPGS

19.5

8.3

0

PPSH

0

11.1

0

the

�-402-

Table 10. Activity
the 24-hour period,

Feeding

Resting

WTMD

9.1%

.7%

1.1%

WILW

9

3.4

0

CONF

15.8

15.7

2.8

ASPN

7.5

7.2

1.7

GRSL

5.5

3

2.1

PPGS

6.7

4.4

0

PPSH

1.8

2.2

0

Habitat

Type

percentages by habitat type across animals
May 1979, Rocky Mountain National Park.
Moving

across

�July,

-403-

JOB PROGRESS

State of

COLORADO
----------------------------

Project

No.

Work Plan No.
Job Title
Period

W-126-R-2

REPORT

Bighorn

4

1979

Job No.

Sheep and Mountain

Goat Investigations

1

---------------------------------

Prescribed Burning to Improve and Enlarge Bighorn Sheep Ranges
--------------------~------~--------------~--~~--------~----~-----

Covered:

July 1, 1978 - June 30, 1979

Personnel:
Dr. P. N. Omi, Colorado
Service; P. Neil, and T. Woodard.

State University;

J. Bustos,

U.S. Forest

ABSTRACT
Burning of treatment plots was postponed for 1 year t6 allow for further
preparations.
Burning is planned for August-September,
1979.
Three additional bighorn sheep were successfully reared and trained for
use in this study.
Food habits data were collected during a field training
period in November.
Forbs, grasses, and shrubs comprised 63.6, 31.8, and
4.4 percent, respectively, of 21,951 bites observed in 3 habitat types.
Seventeen plant species comprised 2 percent or more of the diet in at least
one of the habitat types.
Plant production estimates
the previous segment.

were calculated

from clip-plot

data collected

��-405-

PRESCRIBED
ENLARGE

BURNING
BIGHORN

TO IMPROVE AND
SHEEP RANGES

Thomas N. Woodard

P. N. OBJECTIVES
1.

To test the hypothesis that prescribed burning of understory
vegetation improves the production and quality of bighorn sheep
forage in a ponderosa pine-Douglas fir timber zone.

2.

To test the hypothesis that bighorn sheep use of burned areas
increases compared to adjacent unburned areas.

SEGMENT

OBJECTIVES

1.

Continue maintenance and training
use in grazing trials.

2.

Raise 3-5 additional

bighorn

sheep.

3.

Complete pretreatment
sheep forage samples.

quality

analysis

4.

Coordinate pre-burn,
Service and Colorado

burn, and post-burn activities with U.S. Forest
State University Fire Science personnel.

METHODS

of hand-reared

bighorn

of previously

sheep for

collected

bighorn

AND MATERIALS

Methods and materials were described
during this segment are as follows.

by Woodard

(1979).

Changes

made

The prescribed burning was not conducted during 1978 as scheduled and is
planned for the August-September,
1979 time period.
Colorado State University
Fire Science, U.S. Forest Service, and Larimer County Sheriffs Department
personnel will be involved in the burning as described by Woodard (1979).

Vegetation
Forage

Measurements

Quality

Vegetation samples will be collected in November, January, and March each
post-treatment
year for forage quality analyses.
Plant parts simulating
those chosen by tame bighorn sheep during grazing trials, described below,
will be hand-plucked.
Separate 50-g samples will be collected by treatment
(burned or unburned) for each plant species comprising 2 percent or more of

�-406-

the diet of tame bighorn sheep within each of the 3 habitat types.
Samples
will be analyzed for crude protein, calcium, phosphorus, acid detergent
fiber, lignin, cell wall constituents, gross energy and in vitro digestibility.

Herbage Yield
Pre-treatment vegetation clip samples collected during the previous
were analyzed and production estimates are presented.

Bighorn

segment

Sheep Food Habits Measurements

Grazing trials will be conducted with tame, trained bighorn sheep to measure
response to treatments.
Three additional bighorn sheep lambs were successfully
reared and trained during this segment.
The tame captive bighorn sheep herd
now consists of 3 castrate males, I intact male, and 2 ewes.
Grazing trials will be conducted during 10-day sampling periods in November,
January, and March each winter after the first postburn growing season.
During the first day the animals will be allowed to graze freely on the study
area to become familiar with the vegetation and data will not be collected.
During each of the last 9 days of a sampling period 4 tame bighorn sheep will
be allowed to graze one at a time for I hour each day. Starting points
for individual trails will be the approximate center of the boundary line
between treatments in a replication and will be selected randomly without
replacement for each animal.
The result will be a start of a grazing trial
for each animal in each of the 9 replications.
Individual plant species selected for biomass, cover, and quality measurements
during post-treatment years will be chosen based on availability and/or
importance as a forage species (Woodard 1979). One grazing trial set with
3 tame bighorn sheep was conducted during November of this segment to test
procedures and estimate importance of forage species for subsequent individual
measurements for the first post-treatment year.

STUDY AREA LOCATION
The study area location was described

by Woodard

(1979).

RESULTS AND DISCUSSION
Preparation

of Study Area

The prescribed burn was not conducted during this segment as planned because
of administrative problems and incomplete environmental assessment preparations
by the USFS. Fire Science personnel at Colorado State University updated and
revised the fire prescription (Appendix I). The burn is scheduled for the
coming segment, and the result will be a year's delay in completion of project
objectives.

�-407-

Selection

of Plant Species for Individual

Measurements

Plant species comprising 2 percent or more of the diet of tame bighorn
sheep within each habitat type will be selected for individual sampling
for production, cover and nutritional composition.
Seventeen species
(Table 1) comprised 2 percent or more of the diet in at least one of the
habitat types during the November grazing trials and will be sampled
individually during the first post-treatment year.
Table 1. Percentage composition of tame bighorn sheep diets during November,
1978. Entries are forage species comprising 2 percent or more of the diet
by habitat type.
Percentage ComEosition by Habitat
Ponderosa Pine
Shrubland

Species

3.0

Agropyron

smithii

Agropyron

spicatum

3.6

Agropyron

trachycaulum

5.7

Antennaria

sp.

2.5

Artemisia

frigida

Artemisia

ludoviciana

umbellatum

11.7

33.5

20.2

3.3

2.8

4.5

kingii

7.7

3.4

Muhlenbergia

montana

16.1

7.9

5.2

9.9

Poa spp.

(leaves)

Total Bites

6.9

5.4
3.1

Stipa comata
Thermopsis

4.1

2.3

fissa

Prunus virginiana

77 .5

2.7

Hesperochloa

Potentilla

2.0

2.9

Carex spp.
Eriogonum

Type
Grassland

divaricarpa

2.3

3,784

6,727

11,440

�-408-

Production estimates
based on clip sampling during 1977 on the study area
are presented in Tables 2, 3, and 4. The majority of these species will
be clipped and weighed separately during post-treatment years regardless
of importance in bighorn sheep diets.
Table 2. Herbage production in ponderosa-pine-Douglas
fir type in the
prescribed burn study area during 1977. Entries are g/m2 + 90 percent
confidence intervals.
Species

2
g/m (± 90% CI)

Litter

730.3 + 350.1

Juniperus scopulorum
(needles and current annual growth)

40.6 +

50.3

I. scopulorum

25.4 +

57.7

15.2 +

19.2

9.5 +

12.6

9.3 +

23.2

A. tridentata
(leaves and current annual growth)

5.7 +

4.2

Muhlenbergia

4.4 +

3.5

3.8 +

6.2

3.5 +

4.0

3.1 +

0.7

2.6 +

7.6

2.4 +

0.8

Artemisia
Purshia

(stems)

tridentata

tridentata

Ribes cereum

(stems)
(stems)

(stems)

montana

~. tridentata

(current annual growth)

Rub1lS deliciosus

(stems)

Carex spp.
Juniperus

communis

Hesperochloa

(needles and current annual growth)

kingii

Agropyron

smithii

2.4 +

1.9

Potentilla

fissa

2.3 +

2.6

(stems)

2.1 +

5.4

(current annual growth)

1.9 +

5.5

(stems)

1.7 +

4.9

(current annual growth)

1.5+

0.7

Prunus virginiana
~. cereum
Physocarpus

monogynus

~. deliciosus

�-409-

Table 2. Herbage production in ponderosa-pine-Douglas
fir type in the
prescribed burn study area during 1977. Entries are g/m2 ± 90 percent
confidence intervals.
(Continued).
Species

g/m

Artemisia

ludoviciana

Poa spp.
-P. virginiana
Agropyron

(current annual growth)

spicatum

J. communis

(stems)

Aster porteri
Populus

tremuloides

(stems)

2

(± 90% CI)

1.3 +

1.6

1.3 +

1.8

1.3 +

3.3

1.2 +

0.3

1.2 +

3.5

1.1 +

2.9

1.1 +

3.1

Artemisia

frigida

1.0 +

1.4

Chrysopsis

villosa

1.0 +

1.0

1.0 +

2.9

Arctostaphylos

uva-ursi
-----

Table 3. Herbage production in the shrubland type in the prescribed burn
study area during 1977. Entries are g/m2 ± 90 percent confidence intervals.
Species

g/m

2

(± 90% CI)

Litter

659.4

+ 13.8

Artemisia
Purshia

tridentata

tridentata

(stems)
(stems)

71.6 + 53.8
54.6

+ 5.9

Ribes cereum

(stems)

30.9 + 41.1

A. tridentata

(leaves and current annual growth)

20.6 + 14.1

P. tridentata

(current annual growth)

12.8 +

4.4

12.6 +

7.6

11.6 +

0.5

10.5 +

7.8

Muhlenbergia
Artemisia
Poa spp.

montana

frigida

�-410-

Table 3. Herbage production in the shrubland ~ype in the prescribed burn
study area during 1977. Entries are g/m2 ± 90 percent confidence intervals.
(Continued).
Species

g/m2

(± 90% CI)

Chrysopsis villosa

3.6 +

0.5

R. cereum (current annual growth)

3.6 +

2.7

Juniperus scopulorum (needles and current annual growth)

3.3 +

0.9

Bouteloua gracilis

3.2 +

1.9

Antennaria spp.

2.9 +

0.6

Carex spp.

2.8 +

0.8

Agropyron smithii

2.1 +

2.0

Potentilla fissa

2.0 +

1.1

Stipa comata

2.0 +

1.5

Jamesia americana (stems)

1.9 +

5.6

Lupinus spp.

1.9 +

3.5

Rubus deliciosus (stems)

1.7 +

4.3

Thermopsis divaricarpa

1.7 +

1.6

Physocarpus monogynus (stems)

1.5 +

4.2

Eriogonum umbellatum

1.4 +

0.9

Astragalus flexuosus

1.3 +

1.8

Chrysothamnus viscidiflorus

1.3 +

3.9

Danthonia parryi

1.3 +

3.4

Oxytropis spp.

1.3 +

2.2

Hesperochloa kingii

1.1 +

1.1

Artemisia ludoviciano

1.1 +

1.1

Achillea lanulosa

1.1 +

0.5

R. deliciosus (current annual growth)

1.0 +

2.4

�-411-

Table 4. Herbage production in the grassland type in the prescribed burn
study area during 1977. Entries are g/m2 ± 90 percent confidence intervals.
2

(± 90% Cr)

Species

g/m

Litter

235.7

+ 98.5
+ 12.5

Artemisia

frigida

31.4

Agropyron

smithii

30.6 +

+ 3.0

Poa spp.

23.9

Stipa comata

20.2 +

Artemisia

tridentata

Oxytropis

spp.

Purshia

Bouteloua

montana

gracilis

!. tridentata
Achillea

(stems)

(leaves and current annual growth)

lanulosa

Carex spp.
Ribes cereum

(stems)

4.6

14.3 + 22.3
9.7 +

tridentata

Muhlenbergia

(stems)

9.6

0.6

7.6 + 22.2
5.7 +

4.6

4.9 +

1.8

4.2 +

2.9

3.8 +

1.8

3.5 +

0.6

3.1 +

6.6

Antennaria

spp.

3.1 +

4.1

Agropyron

spicatum

3.0 +

1.4

2.9 +

3.7

2.0 +

2.6

Koleria cristata

1.9 +

1.4

Aster laevis

1.5 +

1.9

Danthonia

1.1 +

1.2

1.1 +

3.2

Lupinus

spp.

Potentilla

fruticosa

parryi

P. tridentata

(current annual growth)

�-412-

Intensified sampling and/or larger plot sizes will be utilized during post
treatment years to improve production estimates for important species with
poor estimates under the described sampling scheme.

LITERATURE

CITED

Woodard, T. N. 1979. Prescribed burning to improve and enlarge bighorn
sheep ranges.
Colo. Div. Wildl. Game Res. Rep. January:4S-67.

Prepared

by:
Thomas N. Woodard
Wildlife Researcher

�-413-

APPENDIX I

�-414-

Report to the Colorado Division
of Wildlife

Wintersteen
Updated Prescribed

Park
Burning Plan

Philip N. Omi, Principal

Investigator

February 15, 1979

Colorado Stace University
De~artment

of Forest and Wood Sciences

Fort Collins, CO

80523

�-415-

This report updates the original burning plan by Barrows and Yancik.
(1978) for the Wintersteen prescribed fires. The plan was written for burns
scheduled in the period September 1
necessitated

November IS, 1978.

This update is

due to postponement of the burns until 1979, the replacement 'of CSU's

investigators on the project, and the availability of new information pertaining to the proposed burning operations.
The objectives of the proposed burnings, the description of the study area,
and a summary of pertinent geographic information for the burning sites are
.co~tained in the original burning plan.

Changes in the prescribed burning pre-

scriptions, in assessed potential fire behavior, and preburn preparations are
addressed in this report.

A. CONDITIONS

WA..l{RA.NTINGREVISION

OF PREVIOUSLY

SPECIFIED

BURNING

PRESCRIPTIONS

Revisions in the original burning prescriptions are necessitated by:
1) Additional fuel inventory info~ation
2) Demonstrated

inconsistencies

collected in summer, 1978;

in the parameters specified in the

original prescription; and
3) Synoptic weather information not previously considered.
1. Updated information on fuel inventory
The herbaceous, shrub, and litter fuel components of proposed burn plots
were resampled in summer, 1978, and compared ,vith the previous year's inventory.
The I-hour and lO-hour timelag downed woody fuels (0 - 1/4" and 1/4 - 1" diameter fuels, respectively) were not resampled, the assumption being that fuel
loadings for downed woody materials would remain relatively constant between
years.
The second year's inventor] indicated generally higher levels of herbaceous
live and dead fuels in the bunchgrass fuel type, with little or no difference
in herbaceous fuels of the shrub and ponderosa pine fuel types.

Contributions

to fuel loading attributable to shrub components within each fuel type appeared
negligible, with the exception of an apparent increas8 in the bunchgrass fuel
type.

These inferences seem to indicate that the relatively ,·let precipitation

year preceding the second inventory primarily affected the bunchgrass fuel type.
wnile plausible,

statements must be considered in light of the random

errors associated with any saI:l.pling
technique.

�2) Revisions

-416to previously specified weather parameters

Inconsistencies
moisture) windspeed,

in originally prescribed' weather specifications

(fuel

and relative humidity were discovered after failure to

match these specifications

with the existing historical weather data base at the

Red Feather weather station.

The conflicting weather parameters were re-specified,

based primarily on adjustments

to relative humidity.

3) Synoptic weather

considerations

Personal communications
Laboratory

with Mr. Jack Cohen of, the Northern Forest Fire

revealed the necessity for consideratibn of trends in the jet stream

track aloft in planning the proposed burning dates.

Of special importance is

the northerly migraticn of the jet stream which can be expected in mid-September.
The impact of this migration on fire potential is of sufficient magnitude to
warrant consideration
specified.

of an earlier range of burning dates than originally

Crucial to this consideration will be the extent to which herba-

ceous fuels have cured prior to burning.

B. REVISED PRESCRIPTIONS
The following prescription
decision-making,

for the stndy area represents a guide for

based on the above considerations.

.
1) Time of Year: Between August 15 - September 15, 1979 provided
grasses
,
are 70% cured.
2)

Time of Day for Lan LtLon ;

3) Fuel Moisture

Be tween

1000 and 1400 hours.

Ran£es at 1000 hours:
Ground fuel (litter):

6 - 10%

Herbaceous vegetation:

4-

7%

One-hour timelag fuels:

4 -

7%

Ten-hour timelag fuels:

7 - 12%

4) Air TemDerature:

45°F to 80°F

5) Relative Humiditv:

12% to 35%

6) Hind Velocity:

Between 3 mph and 10 mph

7) Wind Direction:

Preferably south-southeast.

8) State of Weather:

Clear to partly cloudy with no major fronts
approaching.

Less than .2 inch of precipi-

tation 2 days prior to burning date.

�-417Smoke dispersal is not critical.

9) Atmospheric Stability:

Boweve r

a temperature inversion should be avoided.
C. POTENTIAL FIRE BEHAVIOR
The updated fuel inventory, collected as it was following a relatively
wet precipitation year, did not produce substantially different fire behavior
potential when analyzed using computer models.

As such, the predicted ranges

in Barrows and Yancik (1978) are retained insofar as they represent only rough
approximations

to expected ranges of rate of spread, flame length, fireline

intensity,reaction

intensity and crown scorch height.

The seasonal progression in herbaceous curing will be of crucial importance
to fire potential estimation.

Trends in herbaceous curing will be highly

sensitive to precipitation during the pre-burn period.
ing precipitation

Procedures for monitor-

levels are described in the following section.

D. PP~BURN PREPARATIONS
Beginning July 1, 1979, visits will be made to the study area to monitor
curing trends in the three fuel types.

A simplified sampling scheme will be

designed to measure percent curing in herbaceous fuels.

The resampling interval

will be no greater than two weeks, and will aid in establishing any curing trends.
~fuen fuels have reached at least a 50% cured stage, it is highly recommended
that cooperators on this project convene in the study area to finalize plans.
At this meeting, final·strategies

for control lines and amendments to the

firing plan of Barrows and Yancik (1978) will be considered.

Ten days prior

to this meeting, updates to this burning plan will be completed as necessary and
distributed to cooperators.
In addition. the lOOO-hr timelag fuel moisture content and energy release
component readings at the Red Feather weather station will be monitored starting June 1.

These readings will roughly indicate impacts of prolonged drought

periods on fire potential throughout the S~~2r

season.

A portable weather station TNill be established on the proposed burn site
at least 2 weeks in advance of the proposed burn date.

The measures from

this station will augment spot-weather forecasts obtained from the National
T..Jeather
Service.

�-418-

References Cited

Barrows, J. S. and R. F. Yancik.

1978.

and analysis for wildlife habitat.
Division of Wildlife.

Prescribed fire planning
Report to the Colorado

�July, 1979

-419-

JOB PROGRESS REPORT

State of

COLORADO
------------------------------

Project No.

W-126-R-2

Bighorn

Sheep and Mountain

Goat Investigations

Work Plan No.

4

Job No.

----------------------------------

2

Job Title __~S~e~a~s~o~n~a~l~D~i~e~t~a~r~y~P~r~e~f~e~r~e~n~c~e~s~o~f~B~ibg~h~o~r~n~S~h~e~e~p~
_
Period Covered: July 1, 1978

- June 30, 1979

Personnel: Institute of Arctic and Alpine Research Mountain Research Station
personnel, D. Baker, D. Canalte, T. Dailey, B. Gill, T. Hobbs, L. Stevens,
and T. Woodard.

ABSTRACT
Grazing trials with 3 tame bighorn sheep were conducted in alpine type
during winter.
Grasses, forbs, and shrubs comprised 65.9, 30.8, and 3.3
percent, respectively, of the over winter diet.
Estimated sample sizes required for estimating herbage production by
selected plant species in different habitat types with a clip and weigh
technique are presented.

��-421-

SEASONAL DIETARY PREFERENCES
BIGHORN SHEEP

OF

Thomas N. Woodard

P. N. OBJECTIVES
1.

To describe and quantify
bighorn sheep.

seasonal

diet selection

2.

To evaluate
forages.

potential

and digestibility

3.

To test the hypothesis that forage yields in seasonal bighorn sheep
habitats can be quantified within acceptable tolerance limits with
available fiscal resources.

4.

To quantify

nutritional

bighorn

sheep forage yields within

of Rocky Mountain

of bighorn

sheep

seasonal habitats.

SEGMENT OBJECTIVES
1.

Develop

2.

Conduct bighorn sheep grazing

3.

Initiate measurements to estimate bighorn
quality in seasonal habitat types.

a detailed

study plan.

METHODS

trials in selected

seasonal

habitat

sheep forage quantity

types.

and

AND MATERIALS

Food Habits
Diet selection of individual bighorn sheep ultimately determine what
vegetation is utilized to meet energy and nutrient requirements.
Evaluation
of the quantity and quality of food resources depends on the estimation of
the array of plants selected by bighorn sheep and the relative importance of
each plant species in the animal's diet (Hobbs and Baker 1977). Forage
selection is a complex process which can vary over time and space (Ellis
et al. 1976). Under Objective 1 of this study, dietary selection of 3 to
6 bighorn sheep previously reared and trained (Woodard 1979) for food habits
experiments were quantified during grazing trials in selected seasonal
habitats.
A grazing trial consisted of two observers counting from close
distance the number of bites by species consumed by one sheep feeding in
selected habitats.
Trials were terminated when animals lost interest in
feeding.
A portable tape recorder was used to record data during trials.
Data collected included animal name, starting time, starting point, weather

�-422-

conditions, number of bites of forage by species, and relevant bighorn
sheep behavior.
Animals were transported daily to and from the Division of Wildlife
Research Center or maintained in holding pens near study area locations
during each sampling period.
Each animal was allowed to graze at will
the first day of a sampling period in each habitat type for exposure
to new surroundings and available forage. The following 3-4 days, a
grazing trial was conducted with each animal in a selected vegetation
type. Starting points for grazing trials were selected randomly
within a selected vegetation type. This procedure was followed for
vegetation type 1, listed below. Food habits data in types 2, 3, and
4, listed below, will be collected in conjunction with an existing
prescribed burning study described by Woodard (1978). Bite count data
from 3 non-burn plots in each of the 3 habitat types will be analyzed
to estimate bighorn sheep diets in these types. The sampling scheme
will be as described by Woodard (1979).
Descriptions of habitat types selected for grazing trials follow those
of Hobbs and Baker (1977) and Marr (1961). They are:
1)

Alpine Tundra Climax Region
Elevation is approximately 3480 m and higher.
Topography is
gently to steeply rolling terrain interspersed with precipitous
cliffs, talus slides, and boulderfields.
Dominant vegetation
are sedges and forbs. Common species include Deschampsia
caespitosa, Kobresia sp., Si1ene acau1is, Trifolium sp., Carex
sp., Geum rossii, and Po1ygonum bistortoides.

2)

Upper Montane Forest Climax Region - Xeric Grassland Type
Elevation is approximately 2440 m. Terrain is level and common
forage species include Muh1enbergia montana, Stipa comata, Poa
sp., Agropyron sp., Boute1oua gracilis, Artemisia frigida, Potenti11a
sp., and Oxytropis sp.

3)

Upper Montane Forest Climax Region - Sagebrush Type
Elevation is approximately 2450 - 2460 m. Topography is a gentle
slope of 0-10 degrees on east and south aspects.
Common forage
species include Artemisia tridentata, Purshia tridentata, plus
those present in the xeric grassland community described above.

4)

Upper Montane Forest Climax Region - Ponderosa

Pine - Douglas Fir Type

Elevation is above 2460 m. Topography is a 15-30 degree slope with
numerous rock outcroppings on south and east aspects.
Common plant
species include Pinus ponderosa, Pseudotsuga taxifo1ia, Juniperus
sp., Ribes sp., Rosa sp., Potenti11a sp., Achillea 1anulosa, Artemisia
frigida, Thermop~divaricarpa,
Poa sp .. Agropyron sp., Bromus sp.,
and Festuca sp.

�-423-

These vegetation types were chosen based on bighorn sheep seasonal habitat
selection propensities as reported by Bauman (1978), Bear and Jones (1973),
Geist and Petocz (1977), Pallister (1974), Shannon et al. (1975), Stelfox
(1976), and Stewart (1975).
Grazing trials with 3 tame bighorn sheep were conducted in the Alpine
Tundra Climax Region during November, January, and March of this segment.

Forage Quality Evaluation
Nutritional values of individual plants eaten by sheep will be assessed
by chemical determinations of percent dry matter, nitrogen, acid detergent
fiber, lignin, cell wall constituents and in vitro digestibility of species
contributing at least 2 percent of total intake.
One-hundred gram samples
were handplucked from plants grazed by tame animals during food habits
studies; plant parts taken simulated as closely as possible those selected
by tame sheep. At least 25 individual plants were composited to form each
sample; these were frozen for subsequent analysis.
Triplicate la-gram samples from each composite will be analyzed for chemical
constituents.
Dry matter digestibility will be measured in vitro using
procedures described by Tilley and Terry (1963) and modified by Pearson
(1970). Triplicate 0.5 g subsamples of each sample will be digested with
rumen inoculum collected from a domestic cow.

Forage Yields
Sample size requirements utilizing a clip and weigh technique, described
below, for estimating above-ground forage yields were determined during
preliminary sampling in seasonal habitat types. The accepted precision
level was within 20 percent of the mean 80 percent of the time for forage
species comprising 2 percent or more of the diet of bighorn sheep as
determined by grazing trials with tame animals.
Twenty, 30.5 x 61.0 cm
quadrats were randomly located within each of 3 of the 5 habitat types
selected for food habit studies.
All vegetation within the quadrat frame
was ground-level clipped, separated and placed in paper sacks labelled
by species, date, habitat type and quadrat number.
Forage species comprising
2 percent or more of the diets were sacked separately.
All other plants were
composited according to forage class (shrubs, grasses, and forbs).
Clipped
vegetation was dried at 1000 C for 24 hours and weighed to the nearest 0.1 g.
Desired precision levels of estimates of individual forage species aboveground biomass were not possible with available fiscal resources and
time allocations utilizing the clip and weigh technique alone (D. Baker, L.
Carpenter, and T. Hobbs; personal communication).
Because of this, two
approaches to estimating biomass will be investigated.
First, a double
sampling system utilizing the clip and weigh technique and an electronic

�-424-

capacitance meter (Neal and Neal 1973) will be initiated.
This work will
be combined with an existing study in which a clip and weigh technique
alone is being used to estimate herbage yield (Woodard 1979).
Vegetation
in 180 quadrats, placed with a restricted random sampling design, will
be ground-level clipped each August and separated by selected species
and forage category in each of 3 habitat types of the Upper Montane Climax
Region (Woodard 1979).
The size of the quadrats will be 30.5 x 61.0 cm
to correspond with the herbage meter size and facilitate the double
sampling scheme.
Before this vegetation is clipped a herbage meter
reading will be taken.
An additional 900 meter readings will be taken at
randomly placed locations in each of the 3 habitat types.
The resulting
double sampling ratio will be I meter read and clipped quadrat to 5 meter
read only quadrats.
The predictive equation resulting from regression of
total herbage weight on meter readings from the small double samples will
be used to estimate weights from the large meter read only samples.
Selected forage species and forage class weights will be estimated by
calculating the percentage contribution of each to the total clip plot
weights and multiplying
the results by the total herbage weights estimated
from the large meter read only samples.
A second approach to estimating above ground biomass of selected forage
species and forage categories will be to investigate the relationship
between cover and biomass.
Six, 0.3 - 0.9 ha tracts have been delineated
in each of 3 habitat types in an existing study (Woodard 1979).
Two
hundred fifty microplots
(Morris 1973) and 30 clip-plots as explained by
Woodard (1979) are established in each of the 18 tracts each August.
The
resulting cover measurement values of selected forage species and forage
categories will be regressed on the corresponding weights.
If there is
a correlation it will be tested for repeatability
in succeeding years.
If
correlation is high and repeatable, resulting predictive equations can be
used to estimate biomass from microplots.
Forage sampling was not conducted during this segment but vegetation
clip data collected previously in habitats 2, 3, and 4 were analyzed for
estimates of sample size requirements and results are presented in this
report.

Statistical

Analysis

Food Habits
Statistical treatment of diet data (percentages) has been reviewed by
Hobbs and Baker (1977).
Similar analysis using t intervals will be used
to make inferences on the precision of diet estimates.
Multivariate
analysis of variances
permit hypothesis testing for:

(Harris

1975, Peden

1972) will

�-425-

1.

Differences in forage choices across animals
periods and between vegetation types.

2.

Differences
types.

in forage

quality

between

between

sampling

sampling

periods

and vegetation

Stepwise regression analysis will be used in an attempt to relate bighorn
sheep forage preference to several measured attributes of vegetation.

Forage Yields
Sample sizes required to estimate, by clipping only, biomass of selected
forage species within 20 percent of the mean with 80% confidence were
estimated after preliminary samplinr by the following standard formula:

N

(0.20

-

x)

2

The important quantity when the double sampling approach (clip plots
and herbage meter) is used will be the product derived by multiplying
the percentage contribution of each selected forage species and category
to the total clip plot weights by the total herbage weights estimated
from the large meter read only samples.
The precision of this product
will be calculated with specific formulas (D. Bowden, personal communication).
The precision will depend on the strength of the correlations of selected
species weights to total clip plot weights and total clip plots weights to
meter readings (D. Bowden, personal communication).

STUDY AREA LOCATION
The alpine study area is Niwot Ridge in the India Peaks area of the Colorado
Front Range approximately
6.4 km west of Ward.
It is bounded on the south
by North Boulder Creek and on the south by South St. Vrain Creek and lies
in T1N, R73W.
The Upper Montane Forest Climax Region study area will be located in the
Wintersteen Park area of the Cache la Poudre River drainage, 2 km northeast'
of Rustic.
It lies in T9N, R73W, Sec. 27.

RESULTS
Bighorn

Sheep Food Habits

Twelve
during

forage species
grazing trials

AND DISCUSSION

comprising 91.9 percent
conducted during winter

of 28,624 bites observed
1978-79 in an alpine tundra

�-426-

habitat type (Table 1). Percentages
diet are depicted in Figure 1.

of grasses,

forbs, and shrubs

in the

Table 1. Percentage composition of bighorn sheep diet across 3 animals in
alpine tundra type during winter 1978-79. Entries are mean percentages
with 90 percent confidence intervals for plant species contributing I percent
or more to the total diet.
Species are ranked in order of importance across
all months.

Species

Percent in Diet
-N-o-v-e-m-b-e-r--------J-a-n-u-a-r-y--------M-a-r-c-h--------A-I-l-M--o-n-t-h

Carex rupestris

3.9 +

4.4

31.2 + 23.6

24.9 +

3.6

22.0 +

9.1

Trisetum

25.8 +

6.8

24.8 + 19.0

16.6 + 12.0

21.8 +

6.4

Campanula
rotundifolia

21.4+

9.9

10.1 +

8.6

5.6 +

6.8

11.3 +

8.8

Kobresia

myosuroides

13.8 + 27.3

9.7 + 13.5

10.0 +

4.8

11.2 + 13.3

Agropyron

scribneri

3.4 +

8.7

5.8 +

3.5

8.3 +

7.8

6.4 +

5.9

Arenaria

fendleri

8.2 +

1.4

4.8 +

7.1

3.2 +

2.7

5.0 +

3.9

1.2 +

0.4

10.9 + 15.0

4.6 +

5.9

spicatum

Trifolium
dasyphyllum

3.4 +

3.6

3.0 +

4.9

3.0 +

5.3

3.1 +

4.4

Salix planifolia
(stems)

2.5 +

4.9

2.5 +

2.5

1.3 +

2.6

2.0 +

1.2

Festuca

0.9 +

0.8

0.4 +

0.1

3.3 +

2.2

1.7+

1.1

Poa spp.

3.4 +

6.9

1.4+

1.0

0.4 +

0.7

1.6 +

2.0

Salix planifolia
(leaves)

3.5 +

6.6

0.1 +

0.1

0.9 +

1.3

1.2+

1.1

Grasses

52.1 + 22.9

73.8 + 16.7 66.5 + 14.8

65.9 + 12.2

Forbs

41.8 + 16.1

23.4 + 19.5 31.3 + 15.7

30.8 + 11.9

Shrubs

6.0 + 11.6

2.8 +

Total Bites

7,500

10,154

Polygonum
bistortoides

brachyphylla

3.2

2.3 + 4.0

3.3 +

10,970

28,624

2.3

�-427-

Forage Quality Evaluation
Chemical analysis of the major species in the sheep diet is in progress
but not yet completed.

Forage Yields
Based on preliminary sampling desired prec~s~on levels of estimates of
individual forage species above-ground biomass cannot usually be attained
with time and fiscal restraints utilizing the clip and weigh technique
Table 2 lists calculated sample size
(plot size = 0.25 m2) alone.
estimates for selected plant species in 3 habitat types with precision
set to within 20 percent of the mean, 80 percent of the time.

Table 2. Calculated clip sample size estimates for precision within 20
percent of the mean with 80 percent probability for herbage production
of selected forage species in 3 habitat types. Estimates are based on
30 randomly placed 0.25 m2 preliminary clip plots.
Habitat
P. Pine - D. Fir

Species
Agropyron

smithii

U

1284

gracilis

Carex spp.

Grass

144

21 -

237 -

630

312 - 1007

98 -

225

40 -

213

309

34 -

158

178 - 1272'

A. spicatum
Bouteloua

Type
Shrub

122 -

374

57 -

156

Hesporochloa

kingii

262 -

528

242 - 1040

Muhlenbergia

montana

182 -

469

61 -

166

138 - 1286

319 -

622

47 -

210

16 -

66

134 -

358

30 -

70

Poa spp.
Stipa comata
Antennaria

species

1289

285 -

796

116 -

753

Artemisia

frigida

423

65 -

279

26 -

79

Eriogonum

umbellatum

245

286 -

787

250 -

320

437 -

625

187 -

330

52 -

337

Lupinus

sp.

Oxytropis

spp.

212 -

1294

�-428-

Table 2. Calculated clip sample size estimates for precision within 20
percent of the mean with 80 percent probability for herbage production
of selected forage specie in 3 habitat types. Estimates are based on
1
30 randomly placed 0.25 m preliminary clip plots.
(Continued).
Habitat Type
P. Pine - D. Fir
Shrub

Species

Grass

Potentilla

fissa

62 -

359

124 -

921

Thermopsis

divaricarpa

410 -

510

100 -

432

893

Artemisia tridentata
(leaves and ca~7)

166 -

724

55 -

372

197 - 1028

A. tridentata

408 -

764

136 -

340

452 - 1152

50 -

178

14 -

42

26 -

604 -

632

106 -

607

634

516 - 1293

142 -

645

1058

(stems)

Litter
Purshia tridentata
P. tridentata

(cag)

(stems)

148

.
.
b ase d on up
-l/R anges represent mlnlmum to maXlmum
samp l'e Slze estlmates
to 6 different preliminary sample sets.

2/

- cag

= current

annual growth.

LITERATURE

CITED

Bear, G. D., and G. W. Jones.
1973. History and distribution of bighorn
sheep in Colorado.
Colo. Div. Wildl. Fed. Aid.Job Final Rep. Proj.
W-41-R, vJork Plan 1, Job 1. 232p.
Ellis, J. E., J. A. Weins, C. F. Rodell, and J. C. Anway.
1976.
conceptual model of diet selection as an ecosystem process.
Theoret. Biol. 60:93-108.

A
J.

Geist, V., and R. G. Petocz.
1977. Bighorn sheep in winter:
do rams
maximize reproductive fitness by spatial and habitat segregation
from ewes? Can. J. Zool. 55:1802-1810.
Harris, R. J. 1975. A primer of multivariate
Press, New York.
332p.

statistics.

Academic

�-429-

Hobbs, N. T., and D. L. Baker.
1977. Systems modeling big game populations
simulations of the carrying capacity of the Rocky Mountain National
Park elk winter range. Colo. Div. Wildl. Game Res. Rep., July, Part
III: 12S-30S.
Marr, J. W. 1961. Ecosystems of the east slope of the Front Range in
Colorado.
Univ. of Colo. Studies Series in BioI., No.8.
Univ.
of Colo. Press, Boulder.
134p.
Morris, M. J. 1973. Estimating understory plant cover with rated microplots. USDA For. Servo Rocky Mtn. For. and Range Exp. Sta. Res.
Paper RM-104.
12p.
Neal, D. L., and J. L. Neal.
1973. Uses and capabilities of electronic
capacitance meters for estimating standing herbage.
J. Br. Grassl.
Soc. 28(2):81-89.
Pallister, G. L. 1974. The seasonal distribution and range use of
bighorn sheep in the Beartooth Mountains, with special reference
to the West Rosebud and Stillwater herds. M.S. Thesis, Montana
State Univ., Bozeman.
67p.
Pearson, H. A. 1970. Digestibility trials:
In vitro techniques. Pp.
8S-92 In H. A. Paulsen, and E. L. Reid (Cochairman) Range and
wildlife habitat evaluation - A research symposium.
USDA For.
Servo Misc., Publ. 1147. 220p.
Peden, D. G.
plains.

1972. The tropic relations of Bison bison to the shortgrass
Ph.D. Thesis, Colo. State Univ., Fort Collins.
134p.

Shannon, N. H., R. J. Hudson, V. C. Brink, and W. D. Kitts.
1975.
Determinants of spatial distribution of Rocky Mountain bighorn
sheep. J. Wildl. Manage. 39(2) :387-401.
Stelfox, J. G. 1976. Range ecology of Rocky Mountain
Can. Wildl. Servo Rep. Series Number 39. SOp.

bighorn sheep.

Stewart, S. T. 1975. Ecology of the West Rosebud and Stillwater
bighorn sheep herds, Beartooth Mountains, Montana.
M.S. Thesis.
Montana State Univ., Bozeman.
129p.
Tilley, J. M., and R. A. Terry.
1963. A two-stage technique for
in vitro digestion of forage crops. J. Br. Grassl. Soc. 18(2): 104-111.
/)

1/
Prepared by:

--/
).., .:...t. ) . \.', .. _ .• '

.~ ;,' //[t'

)A'

:,

.

/!

C&lt; /J-,,:(;(jl ,{r;//

homas N. Woodard
Wildlife Researcher

r

/ /

�-430-

70

60

50

.._,
&lt;!)
or&lt;

q

40

C

or&lt;

u

~
Qj

U
H

a.J
P-&lt;

30

20

10

November

Figure

1.

January

Harch

Diet by forage category of 3 tame bighorn sheep
during winter 1978-79 in alpine tundra type on
Niwot Ridge, Coloradoo

�-431-

JOB PROGRESS

State of

COLORADO.
------------------~----------

Project

No.

Work

Period

Covered:

Personnel:

REPORT

4

Bighorn
Johne's
July

Robert

1979

Big Game Investigations

W-126-R-2

Plan No.

Job Title

July,

3
Job No.
Sheep Investigations--Investigations
of
Disease in Selected Bighorn Sheep Populations

--~~~~----~-----------------

1, 1978 - June 30, 1979

E. Keiss, Timothy G. Baumann, Gene G. Schoonveld.

ABSTRACT
Methods for the isolation and identification of Mycobacterium
paratuberculosis
were developed and a total of 281 separate bighorn sheep fecal pellet groups
were cultured.
Two cultures from the Grant area were positive presumptive
for the presence of ~. paratuberculosis.
Distribution, habitat preferences and age-sex composition of bighorn sheep
(Ovis canadensis) and Mountain goat (Oreamnos americanus) in the Mt. Evans,
Colorado vicinity were studied during grounds and aerial surveys conducted
during the summer of 1978. Minimum population estimates of 205 bighorn
sheep and 55 mountain goats were obtained on the basis of non-duplicate
counts.

��-433-

INVESTIGATIONS OF JOHNE'S DISEASE
IN SELECTED BIGHORN SHEEP POPULATIONS
ROBERT E. KEISS AND TIM BAUMANN

P. N. OBJECTIVE
The primary objective of this study will be to enhance the knowledge of
the status of Johne's Disease in wild ungulate populations with emphasis
on bighorn sheep and mountain goats.

SEGMENT OBJECTIVES
1.

To develop a Johne's Disease diagnostic capability
Collins Wildlife Research Center Laboratory.

2.

To provide the Colorado Division of Wildlife with a diagnostic
service for culturing Mycobacterium paratuberculosis in tissues
and/or feces of clinically ill or dead animals suspected of having
Johne's Disease.

3.

To ascertain the viability of~. paratuberculosis in fecal pellets
of bighorn sheep and mountain goats over time and under field
conditions.

4.

To test for the presence
populations.

5.

To relate population dynamics of bighorn sheep populations and
mountain goat populations of Mt. Evans to Johne's Disease infections.

of M. paratuberculosis

at the Fort

in bighorn

sheep

PROCEDURES
For detailed procedures used to culture Mycobacterium paratuberculosis
in
this segment refer to the PROGRAM NARRATIVE for Project W-126-R-2, Work
Plan 4, Job 3.
Laboratory

Culture

Before any culture work could be carried out for this project, it was
first necessary to set up the laboratory capability to do the work.
Glassware and supplies were obtained and the procedures developed which
were ultimately successful in culturing and propagating~.
paratuberculosis
on culture media.
The following collections of bighorn
cultured in the laboratory.

sheep fecal pellet groups were

�-434-

1.

On January 18, 1978 a total of 17 pellet groups were collected from
bighorn sheep trapped from the Chalk Creek herd.
These fecal samples
were cultured on August 29, 1978 and all groups were negative.

2.

On January 26, 1978 a total of 41 pellet groups were collected from
bighorn sheep trapped from the Tarryall Mountain herd.
These fecal
samples were cultured on August 14, 1978 and all groups were negative.

3.

On February 9, 1978 a total of 34 pellet groups were collected from
bighorn sheep trapped from the Poudre River herd.
These fecal samples
were cultured on August 29, 1978 and all groups were negative.

4.

On February 22, 1978 a total of 20 pellet groups were collected from
bighorn sheep trapped from the Almont Triangle herd.
These fecal
samples were cultured on August 24, 1978 and all groups were negative.

5.

On February 23, 1978 a total of 9 pellet groups were collected from
bighorn sheep trapped from the Pikes Peak herd. These fecal samples
were cultured on August 24, 1978 and all groups were negative.

6.

On March 1, 1978 a total of 39 pellet groups were collected from
bighorn sheep trapped from the Mt. Evans herd near Grant.
These
fecal samples were cultured on August 9, 1978 and all groups were
negative.

7.

On January 12, 1979 a total of 51 bighorn sheep pellet groups were
collected from the Grant herd by picking up from the ground where they
were deposited.
These pellet groups were cultured on January 18,
1979 and two pellet groups were positive presumptive for ~. paratuberculosis.

8.

On January 26, 1979 a total of 15 pellet groups were collected from
bighorn sheep trapped from the South Platte River Canyon herd.
These
fecal samples were cultured on January 29, 1979 and all groups were
negative.

9.

On February 12, 1979 a total of 26 pellet groups were collected from
the Grant herd by picking up from the ground where they were deposited.
These pellet groups were cultured on February 14, 1979 and all groups
were negative.

10.

On March 28, 1979 a total of 29 pellet groups were collected from bighorn
sheep trapped from the Grant herd.
These fecal samples were cultured
on March 29, 1979 and none of them were positive for M. paratuberculosis.

Bighorn

Sheep and Mountain

Goat Population

Data on Mt. Evans

All population data were collected by Mr. Timothy G. Baumann during the
summer of 1978 and the following information was taken from his report
which is on file in the Northeast Regional Office - Ft. Collins, Colorado.

�-435-

During May-September
1978, 2152 bighorn sheep and 1133 mountain goats were
located in the Mt. Evans region, including duplications.
The geographical
distribution of collared and uncollared bighorn sheep observed during this
study suggests that three relatively distinct ewe-subadult segments exist
within the Mt. Evans population.
Summer ranges of the "Mt. Evans" segment
were delimited during 1978 based upon relocations of yellow-collared
and
white-ear-tagged
sheep; these individuals were trapped and marked on
Goliath Peak during 1977.
The "Mt. Evans" sheep occupied cirques and
mountaintops near the heads of the Scott Gomer and Chicago Creek drainages.
Sheep marked with orange or red collars and/or yellow, black or orange
ear tags frequented summer ranges above and below treeline in the Threemile
and Deer Creek drainages.
These sheep (marked at two Grant trap locations
during 1977 and 1978) also utilized south-facing slopes below treeline in
the Geneva Creek and Platte River Valleys during movements associated with
use of the Grant salt station.
The existence of a third ewe-subadult
segment, the "Frozen Lake" sheep, was suggested by 256 sightings (including
duplications)
of unmarked sheep in the Arrowhead Mountain - Geneva MountainMount Bierstadt - Epaulet Mountain region during May-September.
Since
sightings of marked sheep were not recorded in this region during 1977-1978,
these sheep are treated as a separate ewe-subadult segment of the Mt. Evans
population in this report.
Interchange between sheep marked at Grant and
those marked on Goliath Peak was not detected during this study.
Hence,
exchange of individuals among the three ewe-subadult segments during summer
1978 was assumed to be negligible.
Two areas of ram concentration were also identified during spring 1978.
Bands of mature rams were freqently located in the Sugarloaf Peak and
Rosedale Peak regions during May and June.
Sightings of orange-ear-tagged
rams on Rosedale Peak and white-ear-tagged
rams in the Clear Creek watershed,
recorded during this study, suggest that rams which frequent the Platte River
Valley during fall and winter concentrate on Rosedale Peak while those which
range through the Clear Creek watershed concentrate in the Sugarloaf Peak
region.
These two ram segments probably drift throughout the entire study
area and intermingle during the rut, as the sighting of a white-ear-tagged
ram near Grant during 1977 rut (R. Mason, personal commuication)
suggests.
Mountain goats which inhabit the Mt. Evans study area may be reproductively
isolated from those which inhabit the Continental Divide west of Guanella
Pass.
These goats are not marked, therefore this theory cannot be proven
or discounted at present.
Since goat crossings of the heavily traveled
Guanella Pass Road have not been observed by or reported by D.O.W.
personnel in the area, interchange between these two "sister herds" is
assumed to be minimal.
Results regarding mountain goat distribution,
abundance and population structure reported below refer to goats east of
Guanella Pass which are considered as a distinct population in this report.
Distribution.
Patterns of sheep and goat concentration
changed in the
Mt. Evans region as the summer progressed, apparently in response to
shifts in the distribution and availability of green forages.
Shannon
et ale (1975) postulated a similar influence of forage quality and
availability on the seasonal distribution of bighorn sheep in British

�-436-

Columbia, as did McFetridge (1977) for mountain goats in Alberta.
The
frequency of ram band observations decreased during summer and so shifts
in ram distribution were less apparent than those exhibited by ewe-subadult
segments.
Figure 1 illustrates sheep and goat distribution in the study area during
May-June, when lambing and kidding activities reached peaks.
The first
newborn lamb was located on 23 May; the first newborn kids were seen on
30 May. One lamb was born in the Beartrack Cirque during the 21-27 June
interval; by this time most lambs and kids had been born. Peaks in
kidding activities were also recorded during late May and early June by
Brandborg (1955) in Idaho, Debock (1970) in British Columbia and Richardson
(1971) in the Black Hills of South Dakota. Lambing peaks for bighorn
sheep in the Northern Rocky Mountains have been recorded during late May
and early June by Smith (1954), Woodard (1971), and numerous other authors.
Apparently, peaks of mountain goat and bighorn sheep parturition occur in
close temporal sequence throughout the Northern Rocky Mountains.
Lambing and kidding areas were identified in the Mt. Evans study area
based on observations of solitary (presumably parturient) females and
females with newborn in steep and cliffy terrain during late May and early
June. Lambing areas were identified on Gray Wolf Mountain, Geneva Mountain,
Kataka Mountain, Mt. Logan and in the Chicago Lakes, Frozen Lake and Camp
Creek cirques.
These 7 lambing areas are characterized by southerly aspects
(with the sole exception of the northwest-facing Gray Wolf areas),
interspersion of 600-vertical rock cliffs with 50-600 boulder-turf terrain
and the occurrence of persistent snowfields during spring.
Interspersion
of green forage, water and escape cover, common to each of the 7 lambing
areas, was probably the key to concentrated use of these sites by parturient
ewes and ewes with newborn lambs.
The Chicago Lakes Cirque and the cirque between it and Gray Wolf Mountain
were primary kidding areas during 1978 as indicated by the abundance of
female goats with newborn kids which were observed there during June.
These two kidding areas are similarly characterized by 500-vertical,
south-facing terrain where free water and green forage were abundant during
May and June. Kidding may occur on south-facing cliff and bench sites
above Vance, Peede and Bear Creek drainages where goat use was also recorded
.during May and June. Use of subalpine cliffs as kidding areas has been
reported by Hibbs (1965) in the Collegiate Range of Colorado and by Debock
(1970) in the Canadian Rocky Mountains of British Columbia. Maximum daily,
non-duplicate counts of kid and adult goats increased on Mt. Evans tundra
during May-August 1978. This trend also suggests that some kids are born
in subalpine areas and follow their dams to tundra summer ranges.
By mid-July green-upon the tundra was widespread and the availability of
new green herbaceous growth increased as snowmelt progressed, especially
in cirque headwalls.
Sheep and goats made frequent use of snowbed sites
during July and August.
By this time availability of free water had
noticeably decreased on the Geneva Mountain, Kataka Mountain and Mt. Logan
lambing areas.
These shifts in the distribution and availability of
forage and water resources may have initiated changes in sheep and goat

�-437-

.
1

/
.'7

(

----7--

r=

~

Ram

~

Mount Evans Sheep

Conc.entrations

t;_:;:;::;j Mounta;n Goal.
~

Gaal.

~

Frozen Lake Sheep

and Mount Eyans Sheep

~

Goals Clnd Frozen Lake Sheep

illIIIl Grant Sheep
Fig. 1. Distribution of bighorn sheep. and mountain goats
in the lwunt Evans, Colorado vicinity ~~y - June

1978.

�-438-

distribution observed during June and July (Fig. 2). Sightings of ram
bands on Sugarloaf Peak, and sign of recent use on Rosedale and Bandit
Peaks during late August indicated that minor shifts in ram distribution
had also occurred.
By September many tundra plants were cured out, especially turf sites,
and the distribution and availability of green forage was basically
restricted to mesic swales and snow accumulation sites. The Grant
ewe-subadult segment expanded its summer range to include the Rosalie
Peak-Deer Creek and Roosevelt Lakes cirque regions during September while
the Frozen Lake and Mt. Evans segments reverted to distribution patterns
similar to those observed during late May (Fig. 3). Small ram bands
(2-9 animals) were seen in and out of ewe~subadult company during late
August and September in areas where they were not seen during May-August.
Geist (1971) noted similar trends in ram distribution for bighorn sheep
herds in the Canadian Rocky Mountains.
Goat distribution during September
was not noticeably different from that of July-August; although, as stated
above, the abundance of goats on the tundra was maximum during late summer.
Distributional trends indicating similar concentrations of goats on tundra
during summer were reported by Hjeljord (1973) in Alaska, McFetridge (1977)
in Alberta and Schultz (1973) and Johnson (1975) in Colorado.
The effects of "salting" on the movements and distribution of the Mt.
Evans ewe-subadult segment and mountain goats in the vicinity is somewhat
obscure since salt was placed near the center of their summer ranges (Fig. 4).
Large quantities of soil were ingested by sheep at bait stations on Rogers
Peak and Goliath Peak during late August and September.
A relationship
between late summer use ..
of these areas by the Mt. Evans sheep and the
availability
of salt cannot be ascertained as "pre-salting" distribution
data are not available.
Hebert and Cowan (1971) reported that the attraction
of big game to natural salt licks was related to availability of sodium
during periodic sodium stress, which occurs when an increase of succulent
growth in their diets decreases sodium retention.
Although one might
expect that sodium stress should be most intense during July and August
when succulent herbaceous growth was most abundant (rather than during
September when availability of such feed was apparently reduced) a delayed
response to this stress may have occurred.
Goats ingested significant
quantities of soil at bait stations above Lincoln Lake and Tumbling Creek
throughout the summer. Movements of marked Grant sheep between the Camp
Creek cirque and the Grant salt station over straight line distances
of ~ 5.1 km, suggest that availability of salt has a significant effect
on the distribution of these sheep. Heimer (1973) noted a similar influence
of salt availability on the distribution of Dall sheep in Alaska as
did Brandborg (1955) and Hebert and Cowan (1971) for mountain goats.
Relocations of marked "Mt. Evans" sheep during July and August suggest that
"handouts" offered by tourists along the Mt. Evans road affect an artificial
concentration of sheep in this vicinity.
Handout items taken during summer
1978 included granola bars, sandwiches, dougnuts, saltine and "animal"
crackers, "doritos", wine, and chlorox, which was inadvertently made
available to the sheep.
Sheep actively sought handouts by approaching

�-439-

I

/

~

Ram Concentrations

~

Mounl E~ons. Sheep

t:;:;:;:j Mounlain Gool.
~

Goals and Mount Evan.

~

Frozen Lake Sheep

Sheep

[[ill] Gran! Sheep

Fig. 2.

Distribution of bighorn sheep and mountain goats
in the Hount Evans, Colorado vicinity July
August 1978.

�-440-

\
.\

~

Mounl E.ans Sheep

[::-:-:-4

Mounlain Goals

~

Goals and Mounl Evans Sheep

~

Frozen Lake Sheep

~

Goals and Frozen Lake Sheep

UIllI1J

Gran I Sheep

~

Goats and Grant Sheep

Fig. 3~ Distribution of bighorn sheep and mountain goats
in the Mount Evans, Colorado vicinity during
September 1978.

�-441-

Fig. 4.

.

Locations at ~hich salt ~as placed during ·summer
1978; Hount Evans, Colorado vicinity.

�-442-

tourists and accepting hand-held foods.
Although the nutritional effects
of this "supplemental
feeding" are unknown, the concentration of sheep on
Mt. Evans is apparently a conditioned response to it. Daily movements
of yellow-collared
ewes between the Gray Wolf Mountain lambing and nursery
area and the Mt. Evans Crest House (~ 1.4 km straight-line distance) were
observed on several occasions during July.
Successive relocations of
yellow-collared
and white-ear-tagged
sheep and daily observations of
unmarked sheep in the area indicate that the Mt. Evans Crest House was
near the center of activity for many of these sheep.
Abundance.
Summation of maximum non-duplicate
counts obtained for each
age-class (Tables I and 2) indicate that a minimum of 205 bighorn sheep
and 55 mountain goats inhabited the study area during summer 1978. Although
these maximum class counts are not accurate or precise estimators of
population sizes, they do provide the best available population structure
information and absolute minimum population figures.
If minimum population
estimates derived by summation of maximum non-duplicate
class counts recorded
during summer 1978 are considered to arbitrarily represent 70% of the total
Mt. Evans mountain goat and bighorn sheep populations, then extrapolated
populations of 79 goats and 293 sheep may be derived.
Based on the extent
and intensity of sampling of sheep and goat band composition during this
study,
I estimate that 65-75 mountain goats and 225-250 bighorn sheep
inhabit the Mt. Evans region.
Martin and Stewart (1977) reported maximum
"non-duplicate"
counts of 208 sheep and 68 goats for the Mt. Evans study
area, however, these "minimum population" figures are not substantiated
by daily observation summaries presented in their report.
Their 1977
estimates must have been based on sums of animals (including unclassified
individuals) recorded for specific geographic regions over a period of days
and the assumption that no movements between these regions occurred.

Sheep Population Structure.
Lamb/ewe ratios obtained for the Mt. Evans
and Grant ewe-subadult segments of the sheep population (81/100 and
72/100 respectively)
compare favorably to those summarized by Buechner
(1960).
Since the Frozen Lake segment is composed of unmarked sheep,
the likelihood exists that these animals did not receive antihelminthic
drug treatment during 1977-1978 and were therefore not marked.
The
relatively low lamb/ewe ratio (44/100) observed for the Frozen Lake
segment during summer 1978 may indicate that reproductive success has
been reduced in this herd segment as a result of lungworm infection.
However, larval loads recorded for a small number of fecal samples collected
during June 1978 do not suggest high levels of lungworm infection in
sheep which utilized the Frozen Lake region.
Since survival rates beyond the yearling age class are generally believed
to be high in mountain sheep populations
(Buechner 1960, Geist 1971), a
yearling/adult
ratio of 24/100 (calculated from pooled maximum class
counts collected during this study) suggests that population stability will
be approximated as the annual mortality rate of the adult and yearling age
classes combined approaches 24%, providing that population composition
remains relatively constant.
Year to year variations of birth rate and
lamb survival, which are apparently not uncommon in bighorn populations,

�Table 1. Mount Evans. Colorado bighorn sheep population size and age-sell;structure
based on maximum non-duplicate class count s,Hay - September 1978.

Herd

3/4
7

Mount Evans

5/8 1/2
5

6

1/4

yJ'

Af.

y~

L

Hinimum
herd
size

3

4

26

5

21

77

Frozen Lake

--

--

--

1

4

16

7

7

35

Grant

6

2

5

2

6

39

5

28

93

estimates

Lamb
Ewe

-

Yearlinfj
Ewe

81

35

ToO

100

44
100

69
100

72

28
100

Wo

Ram
Ewe

a

Male
Female

I

-i"-i"-

W
I

All

Herds

13

7

11

6

14

81

17

56

(6%) (3%) (5%) (3%) (7%) (40%) (8%) (28%)

a

Rams excluding

yearling males.

205

69

38

1.6

63

Wo

'iOo

TOo"

100

�Table 2. Hount Evans, Colorado mountain goat population
cstimateso Hay •..September 19180

size and agu-uex structure

Source of
estimate

ABe-sex composition

Maximum class
counts

Arf#

A~

Y

K

Hinimum
population

4

18

17

168

55

Pooled data
for all banda completely
classified. Hay September

aincludinB

3 pairs of twins.

Kids
Nannieo

Yearlin8s
Nannies

Kid
Older animals

89

94

41

100

IOO

100

96
100

83
100

61
100

I
.p.p.pI

�-445-

will necessitate frequent re-evaluation of these ratios in the future.
A ram/ewe ratio of 56/100 and favorable ratio of young (yearling,
1/4 and 1/2 curl) rams to older (5/8 and 3/4 curl) rams (~ 1.5/1),
indicated by maximum class counts (Table 1), do not suggest that the
current intensity of ram harvest is adversely affecting the population
structure.

Mountain Goat Population Structure.--Interpretation
of mountain goat
age ratios is difficult due to classification bias which results from
the limited degree of sexual dimorphism in goats and rapid growth of
yearling and two-year-olds.
Physical measurements of mountain goats
published by Brandborg (1955) Lentfer (1955) and Richardson
(1971)
indicate that reliable discrimination of adult males from adult females
and differentiation
of long yearlings and two-year-olds from adults on
the basis of horn and body characteristics
is difficult, at best, over
distances at which they are commonly observed in the field.
In order
to minimize classification bias, many authors have employed kid/adult,
yearling/adult
and kid/older animal ratios in their analyses of mountain
goat population composition.
A minimum of 1978 Mt. Evans mountain goat population estimate, age ratios
based on maximum class counts, and age ratios computed from pooled data
for bands in which every goat was aged and sexed are presented in Table 2.
Brandborg (1955) suggested that estimates obtained by pooling data collected
during an entire field season were most accurate indices of goat
population structure than those obtained from maximum class counts.
In
an attempt to refine estimates of population composition based on maximum
class counts recorded during this study, monthly age ratios were computed
from observations of bands in which all goats were aged (Fig. 5) and those
in which all goats were aged and all adults sexed (Fig. 6).
Curves
constructed for kid/adult, kid/older animal (Fig. 5) and kid/adult female
(Fig. 6) ratios illustrate an increase in the proportion of kids in the
population during July-September
1978. Curves constructed for yearling/
adult and yearling/adult
female ratios show a coincident decrease in the
proportion of yearlings in the population.
These trends were probably
affected by concentration of goats on the tundra during late summer,
resulting in increased opportunities
to observe adult females with kids,
and a decrease in the ability to reliably distinguish yealings from
adults as they approached adult size.
Although kid/adult female and
yearling/adult
female ratios computed for August and September were not
significantly different from those for May-July (P &lt; .10), the decreasing
widths of 90% confidence intervals for these ratios (Fig. 6) support the
theory that these trends occurred.
Since kid/adult female and yearling/adult
female ratios were computed
from close-range observations
(all goats were aged and all adults were
sexed), these ratios are the most accurate estimators of kid production
and survival available.
The similarity of kid/adult female ratios
presented in Table 2 with those computed from close-range observations
during August and September (Fig. 6) suggests that the true ratio of
kids/adult females was somewhere in the range of 82-94 kids per 100 adult

�-446-

......
70

YEARLINGS
100 ADULTS

.71

KIDS
100 OLDERANIMALS
KIDS
100 ADULTS

60

SO
NUMBER
OF

.'

47

40

YOUNGER
100 OLDER

..

30
GOATS
20

e_

,
,,
I

..

22

.

20

...•

...

'17 •

14~

10

Y K K

Y K K

Y K K

Y K K

Y K K

AOAA

AOAA

AOAA

AOAA

AOAA

MAY

JUNE

JULY

AUG

SEPT

NUMBER
OF BANOS

10

41

28

2S

61

NUMBER
OF GOATS

2S

120

200

111

244

Fig. 5. ~fonthly kid/older
animal, kid/adult,
and yearling/adult
ratios
observed in the ~fount Evans, Colorado mountain goat population,
Hay September 1978. Data are from bands in which all goats were aged
(g~ = 700), including duplications.

�-447-

YEARLINGS

••••••

100 ADULT FEMALES
KIDS
100 ADULT FEMALES

100

138

105

117 101

102

96

90

OF

•
94·

2----

87·~1

80
NUMBER

104

96

.

83

70
60
0

57.

71

••
.
•

YOUNGER 50
100 OLDER

53 .

•
••

••

43·

40

••
••
•

41

41

GOATS

.

30

27.00 ••••• 28.

20

10

16

o
-24

-1

-32

y

K

y

A~

Al

A~

MAY

K
Y
At

K
A~

Al

JUNE

Y

K

Y

K

A~

A~

At

Ai

J UL Y

·AUG

SEPT

NUMBER
OF BANDS

5

17

19

34

5

NUMBER
OF GOATS

14

97

n&amp;

64

%04

Fig. 6. ~!onthly kid/adult
female and year Lf.ng Zadul t; female ratios
(and
90% confidence intervals)
observed in the :lount Evans, Colorado moun-«
tain goat population.
aay - September 1978. Data are from bands in
which all goats we're aged and all adults sexed (g""" = 393 goats), including duplications.

�-448-

females during summer 1978. Likewise, comparison of yearling/adult female
ratios presented in Table 2 and Figure 6 suggests that the ratio of
yearlings/adult females was between 73 and 94 per hundred.
Martin and Stewart (1977) reported age ratios of 26 kids/100 adults,
24 yearlings/100 adults, and 19 kids/100 older animals and suggested
that the Mt. Evans mountain goat population was approaching stability.
Age ratios computed from data collected during May-September 1978 indicate,
to the contrary, reproductive vigor and suggest the potential for population growth.
Three pairs of twins were observed without duplication during
June and again during September, indicating that approximately 17% of
adult females in the population successfully weaned kids during 1978. These
observations of twinning are also indicative of reproductive vigor.
Brandborg
(1955) suggested that ratios on the order of 50-70 kids per 100 females were
typical of mountain goat productivity during "average years on most ranges".
Bailey and Johnson (1977) found that kid/older animal ratios for introduced
mountain goat populations averaged 59/100 during the first 15 years of their
existence while a ratio of 28 kids/100 older animals was "average" for
native and introduced populations which had been established for at least
16 years.
Many authors have concluded that kid survival is highly variable
and related to severity of winter weather as indicated by disparity between
spring yearling/adult female ratios and kid/adult female ratios obtained
during the previous summer. Hall and Bibaud (1978) reported an average
annual kid mortality rate of 50% for kids in Willmore Provincial Park,
Alberta during 1974-1978.
Clearly, comparison of age ratios obtained during
summer 1978 with those published for other populations (see Schultz, 1973)
indicates that reproduction is good and yearling survival is excellent in
the Mt. Evans mountain goat population.
These trends indicate the potential
for increased mountain goat harvest in the Mt. Evans region.
~[nterspecific Intolerance.--Bighorn
sheep were observed in visual contact
with mountain goats during 92 minutes between 6 June and 23 August, 1978.
All such observations occurred when both sheep and goats were feeding.
No overt aggressive behavior was exhibited by either species during these
observations.
Sheep sometimes ceased feeding as mountain goats approached;
this behavior was most prevalent among lambs.
Sheep and goats commonly
reacted as conspecifics while feeding together as Cowan (1944) and
Brandbord (1955) reported.
Klein (1953) reported several instances of an
avoidance response by Dall sheep elicited by approaching mountain goats in
Alaska.
Petocz (1973) noted an increase in the frequency of overt aggressive
behaviors exhibited by bighorn rams and mountain goats in response to deep
snow conditions.
The potential for interspecific intolerance between
mountain sheep and goats is apparently greatest during winter when availability
of forage is reduced due to snowfall.
Assessment of this intolerance during
periods of winter stress is a topic for future study.

LITERATURE

CITED

Bailey, J. A., and B. K. Johnson.
1977. Status of introduced mountain goats
in the Sawatch Range in Colorado.
Pro. First Int. Rocky Mountain Goat
Symp., February 19, 1977. Kalispell, Montana.

�-449-

Brandborg, S. M. 1955. Life history and management
in Idaho. Idaho Fish and Game Dept., Bull. 2.

of the mountain
142p.

g@at

Buechner, H. K. 1960. The bighorn sheep in the United States, its
past, present and future. Wildl. Monogr. 4. 174p.
Cowan, I. Mc T. 1944. Report of wildlife studies in Jasper, Banff and
Yoho National Parks in 1944 and parasites, diseases and injuries
of game animals in the Rocky Mountain National Parks, 1942-1944.
Wildlife Service, Dept. of Mines and Resources, Ottawa, Canada.
83p.
(mimeo).
DE Bock, E. A. 1970. On the behavior of mountain goat (Oreamnos
americanus) in Kootenay National Park. M.Sc. Thesis, The Univ.
of Alberta, Edmonton.
Geist, V. 1971. Mountain sheep; a study in behavior and evolution.
Univ. of Chicago Press, Chicago and London.
383p.
Hall, W. K., and J. A. Bibaud. Goats and their management in Alberta.
Pro. N. Wildl. Sheep Council, April 12, 1978. Penticton, British
Columbia.
Hebert, D., and I. Mc T. Cowan.
the ecology of the mountain

1970.
goat.

Natural salt licks as a part of
Can. J. Zool. 49:605-610.

Heimer, W. E. 1973. Dall sheep movements and mineral lick use. Final
Report P. R. Proj. W-17-2, W-17-3, W-17-4, W-17-5.
Alaska Dept.
Fish and Game. 35p.
Hibbs, L. D. 1965. The mountain goat of Colorado.
Colo. State Univ., Ft. Collins.
152p.

Unpubl. M.S. Thesis.

Hjeljord, O. 1973. Mountain goat forage and habitat preference
J. Wildl. Manage. 37:353-362.

in Alaska.

Johnson, B. K. 1975. Summer and autumn, 1975 study of mountain goats in
the Collegiate Range of Colorado.
Unpubl. report on file Colorado
Div. Wildl. Res. Center Library, Fort Collins.
28p.
Klein, D. R.
Alaska.

1953. A reconnaissance study of the mountain goat in
Unpubl. M.S. Thesis. Univ. of Alaska, Fairbanks. 121p.

Lentfer, J. W. 1955. A two year study of the rocky mountain
the Crazy Mountains, Montana.
J. Wildl. Manage.

goat in

Martin, L. M., and D. W. Stewart.
1977. 1977 summer census of the Mt.
Evans bighorn sheep and rocky mountain goat populations.
Unpubl.
report on file Colorado Div. Wildl. Res. Library. Fort Collins.
29p.

�-450-

McFetridge, R. J. 1977. Strategy of resource use by mountain goat nursery
groups. Pro. First Int. Mountain Goat Symp., February 19, 1977.
Kalispell, Montana.
Petocz, R. G. 1973. The effect of snow cover on the social behavior of
bighorn ra~s and mountain goats. Can.J. Zool. 51:987-993.
Richardson, A. H. 1971. The rocky mountain goat in the Black Hills.
South Dakota Dept. Game, Fish and Parks Bull. No.2.
24p.
Schultz, R. D. 1973. The Mount Shavano rocky mountain goat herd~Summer
1973. Unpubl. report, on file Colorado Div. Wildl. Res. Library,
Fort Collins.
17p.
Shannon, N. H., R. J. Hudson, V. C. Brink, and W. D. Kitts.
1975.
Determinants of spatial distribution of rocky mountain bighorn
sheep. J. Wildl. Manage. 39:387-401.
Smith, D. R. 1954.
and management.

The bighorn sheep in Idaho; its status life history
Idaho Game and Fish Dept. Wildl. Bull. No.1.
154p.

Woodard, T. N. 1971. Bighorn sheep lamb production, survival and
mortality in the Sangre De Cristo Mountains, Colorado.
Unpubl.
M.S. Thesis.
Colorado State Univ., Fort Collins.
84p.

,,--. -,
Prepared by: I

's

/~t~r_l/' G.:-

Wildlife Researcher

�-451-

JOB PROGRESS

State of
Project

1979

REPORT

COLORADO
W-126-R-2

No.

Job Title

_

P~r~o~n~g~h~o~r~n~P~o~p~u~l~a~t_i~o~n~S~t~u~d~y~

_

Covered:

Personnel:

Big Game Investigations
l

Work Plan No.

Period

July,

5

January

Job No.

1, 1979 to June 30, 1979

Thomas M. Pojar

ABSTRACT
A literature search was conducted and a Program Narrative for the pronghorn
antelope (Antilocapra americana) study was completed.
In addition, an
Environmental Assessment Report was written for this project and is also
included as part of this progress report.

Prepared

by:~~~~
~~~~
__ +_~Thomas M. P jar
Wildlife Researcher

��-453-

PROGR ...
~1 NARRATIVE
(Research)

State:
Project
1.

Colorado
Title:

Study

A.

Big Game Investigations

Title:
Pronghorn Investigations:
Pronghorn Population Study

Project No. W-126-R
Work Plan 2, Job I

NEED
Historically
the pronghorn antelope (Antilocapra americana) in
Colorado has experienced a precipitous decline from an estimated
two million animals in 1860 to a close brush with extinction in 1918
when their numbers we thought to be less than one thousand (Hoover
et al. 1959).
The near extermination
of pronghorn from Colorado
was presumably the result of year-round market hunting.
Legal
protection was afforded the pronghorn in 1898 but actual protection
was not realized until about 1920. With effective protection the
populations
increased to the point that in 1945 the first hunt in
47 years was held, mostly because of crop da~age complaints
(Hay
et al. 1961).
Today the pronghorn is a prized big game animal and
the demand for the privilege to hunt this species exceeds the
available pennits by a margin of four to one (Colorado Division of
Wildlife.
1977).
Nutrition

and Reproduction

The reproductive
potential of the pronghorn is among the highest of
any wild ungulate in Colorado.
O'Gara (1970) reported that 3 to 7
corpora lutea are commonly found in this species, indicating that a
similar number of ova were ovulated, fertilized and developed into
spherical blastocysts.
Usually no more than two of these blascocysts
survive, but Ellis's (1970) review of r~ported fetal rates from five
western states still indicates 1.92 fetuses per doe.
It follows
that a species with a potential for a high fetal rate also has
potential for a high recruitment rate.
The recruitment
rate, as
measured by aerial fa,va to doe ratio estimates, varies a good de21
be twe en populations.
Hithin Colorado, the f awn r do e ratios vary
from the low 40's in the Northeast region (Figure 1) to double that
rate in the Northwest region (Figure 2). The recruitment rate in
these two regions is at least stable or increa3io8, however the
recruitment
rate in the southeast region exhibits a significant
decline (Figure 3). The Thatcher populations shows the steepest
decline of any po?ulation in the southeast, going from 86 fawns per
100 does in 1967 to 43 f a:v ms per 100 does in 1974 (Figure 4).

�-454-

According to Caughley (1976) the fecundity pattern (number of
young born) of an ungulate population can react to environmental
stress by:
(a) increasing the mean age at puberty;
{b) reducing
the pregnancy rate; and (c) lowering ~he mean litter size; Any
of these factors would result in lower late summer fawn:doe ratios
and may provide one of the key manifestations of declining
demographic vigor.
The nutritional plane of the reproductive female has been associated
with ovulation and fertilization success; fetus viability and early
post natal survival; and lactation which affects subsequent surn.mer
fawn survival (Verme 1969). Most of the fetal growth takes place
during the last third of the gestation period (}~ynard and Loosli
1969) which suggests this is one of the nutritionally critical time
periods for the pronghorn (Ellis 1970). Ewes on a low plane of
nutrition during the last half of pregnancy produced small, weak
lambs, most of which died at birth or soon after (Thomson and
Thomson 1948).
Pronghorn inhabit a great diversity of range types (Humphrey 1962).
From a very thorough review of pronghorn food habits literature
Schwartz (1977) stated that most investigators agree that browse
and forbs are preferred by pronghorn but grasses are readily used
especially during the spring green-up period when grasses are highest
in nutrltlon.
In short, pronghorns are herbiverous opportunists,
taking forage that is available, with some selectivity for plants
and plant parts that are most nutritious.
The nutritional difference in habitat types can be detected by
measuring the physiological status of the local animal populations
(Seal et al. 1978). These authors and Seal and Hoskinson (1978)
were ahle to correlate habitat conditions with selected blood
parameters of popul.etLons of white-tailed deer (Odocoileus v i rz i n i anu s)
and pronghorn.
Blood urea nitrogen (BL~) appeared to be the most
reflective of nutritional status of the animal and it was the most
stable in terms of not being affected by handling of the animal.
However, excessive muscular activity and excitation will elevate BUN
after about 6 hours (Trout 1976) Barrett and Chal~ers (1977) verified this by
holding wild trapped pronghorn in captivity and taking blood samples
on successive days. The BUN immediately after capture was 33.20
mg/dl (n = 48) and the following day it was 51.62 mg/dl Cn = 23).
Using blood parameters to assess physical condition of the animal
has the distinct advantage that a sample can be obtained without
sacrificing the subject.
There has been some research on the
relation of nutritional status and hair shaft diameter or hair root
diameters in horses (Godbee et al. 1979), humans (~utrition research
1968, Bradfield 1971, Bradfield 1968), and bovine (Martin et al. 1969,
Haaland 1977). These studies showed good correlation with nutritional intake
but, t o my knowledge, this procedure has not been tested on wi Ld ungulat-es.
Other than simply weighing or taking certain measurements on the
.animal, most other me t hod s of assessing body fat required sac ri ficing
the animal.
Anderson et al. (1969) tested the relationship of six
such methods with the percent petroleum ether extractable carcass

�-455-

fat.
They concluded that the kidney fat index was the best
single index of winter carcass fat, however for a reliable mean
vdlue large sample sizes would be required.
There is ample evidence that nutrition is related to the recruitment rate of a population.
One of the best means of assessing
whether or not the nutritional plane is affecting the reproduction
of a population is to compare it to a population of the same species
that is in healthy condition (Caughley 1977).
There is definitely
a need to attempt to determine what is causing the alarming decline
in the late SUDLT11er
fawn:doe ratios in the Thatcher and other southeast
region populations. Assessment of the range'in meeting thenutritional
requirements of the pronghorn is an important consideration.
Comparison of the
Thatcher population with a highly productive population in the
northwest in terms of physical condition, size, ovulation rate and
fetal rate would be of great assistance in determining
the factors
associated with the low fawn:doe ratios.

Genetics
The genetic
aspects of wild animal populations have largely been
ignored in favor of environmental
factors and overt population
characteristics
(Ryman et al. 1977 and Smith et al. 1976).
Electrophoresis is a relatively new technique that is used for analyzinE
the amount of genetic variation in populations
(Norgan et al.. 197L,).
Loss of genetic variability reduces a popUlation's
ability to adapt
to environmental
change (Nei et aI , 1975, Ayala 1968).
Lm17 ge~l£'tic
variability
is frequently associated with restricted gene flow and
small breeding populations
in the recent past (Smith et al. 1977).
Bonnell and Selander (1974) cite the elephant seal (~1irounga
an~stirostris)
as a typical example of a species that exhibit.s very
low genetic variability.
Today's elephant seal population is derived
from as few as 20 animals as this species survived near extinction
in the 1890's.
These authors speculate that genetic variability
is
not essential for the survival of a species, but that the elephant
seal, lacking a pool of variability with which to adapt to changing
conditions is especially vulnerable
to environmental
change.
.'-Ioose
(Alces al.ces ) in Sweden also exhibit very low genetic variation and
Ryman et al. (1977) speculate this may be due to near eradication of
the species during the 19th century.
Even with the low variability,
the eleccrophoretic
technique is capable of detecting genetic
diversity between populations, which the authors cautiously suggest
may be correlated with antler size.
Genetic variation has been used successfully
to identify genetically
distinct subpopu1ations
within a species (Ryman et al. 1977, Manlove
et a1. 1975, Manlove et a1. 1978, Smith et a1. 1976).
There are
efforts now to associate population attributes with certain genotypes.
Smith (personal communication
1979) has some preliminary results
vh Lch indicate that a particular genetic pattern in wh i.te+ t a i.Le d
deer can be associat2d ~ith time of breeding (and therefore also ti~e
of f awn i.ng) . This results in a double f awn i n g peak in wha t app ea rs

�-456-

to be a distinct population.
Other ?opulation attributes that
can be linked to some discernible genetic characteristic include:
different fertility rates in pigeons (Frelinger 1972),/different
survival rates of juvenile blue grouse (Dendragapus obscurus)
depending on genotype (Redfield 1974), pelage variation in natural
populations of mice (Peromvscus polionotus) (BoHen and Dawson 1977),
artificially induced increased heterogygosity resulted in increased
life span and reduced juvenile mortality in mice (Chai 1959), and
Simon (1978) presents evidence that lack of genetic variability in
hatchery trout, due to using small numbers of parents to produce
brood stock, has resulted in fish that are more vulnerable to
stressful situations as evidenced by increased blood glucose levels
and that some nutrition and disease problems were in fact largely
inbreeding problems.
Selander (1970) has examined aspects of the
genetic structure of wild populations of the house mouse (Mus
musculus) as they are affected by the behavioral mechanism of
territoriality.
His analysis indicates that wild populations of
the house mouse are characterized by fine-scale genetic subdivisions
which is achieved through territorial behavior even in the absence
of physical or ecological barriers.
It seems reasonable, with the amount of evidence accumulating as to
the usefulness
of genetic evaluation in population studies, that
there is a need to include genetic assessment in the pronghorn
population study. Attempts to associate demographic and genetic
attributes in ~~hite-tailed deer are just now beginning to emerge
in the literature (Ramsey et ale 1979). The potential of detecting
genetic differences in the two pronghorn populations selected for
study seems very good. The eastern plains population is small.
about 600 animals, and undoubtedly was derived from a very limited
gene pool with little opportunity for increasing heterogenicity
through imigration.
The northeast region, by contrast, is large
(5000 animals) and may never have been as drastically depleted as
the eastern populations.
In addition, periodic migration into that
population from Wyoming, triggered by severe winter storms may
provide a continued recharging of the gene pool.
Disease
Pronghorn are susceptible to many diseases carried by domestic
livestock.
Pronghorn on the eastern plains may have rr~re direct
contact w i t h domestic livestock than those in the northwest area
of the State because of co~~on use of water sources.
Several of
these diseases are abortive or cause an increase in neonatal
mortality.
Trueblood and Post (1959) isolated vibriosis (Vibrio
i_etus) in a po puLa t Lon of wyoming pronghorn.
Howe (1970) found
evidence that pronghorn may be more susceptible to this disease
than domestic livestcck.
Johnes disease causes reduced lactation
in affected animals which could lead to und2rnourished young,
decreasing their chances for survival.
This disease is cOmmon
in domestic livestock and is probably transmissable to pronghorn
(Beth IHlliams, personal comraunLc at..i on ) . Brucellosis (Bruc clla

�-457-

abortus) and leptospirosis
(LeptosDira 2omona and~.
zanari
zanoni) are abortive bacterial diseases.
In 1485 pronghorn
blood samples tested for leptospriosis
from 1969 to 1974 in
Colorado, 80 of them were positive and of 5,272 blood" samples
screened for brucellosis only one was weakly positive (Adrian

1976) .
Leptospirosis
is defintely found in Colorado pronghorn populations,
however the extent of infection rate should be much more intensively
monitored in the two populations selected for the study.
Hore
intensive screening for vigriosis and Johne's Disease should be
initiated because of the high likelihood the Colorado pronghorn
populations are infected with these diseases.
Any of these
diseases could have a dramatic affect on the reproductive
performance of pronghorn populations.
Hunting

Disturbance

of the Rut

There is some evidence that double f awn Lng peaks occur in some
southeast region pronghorn populations.
Elkins (unpublished data)
made weekly helicopter surveys of selected management units and
his data show a fawn:doe ratio peak in mid-June, followed by a
sharp decline, then another peak in mid-July.
There is speculation
that this double fawning peak phenomenon is the result of conducting
the animal hunt in September which coincides
with the rut (f"renzlow
et al. 1968). Hunting pronghorn during the mating season could
disrupt breeding behavior and harem formation resulting in a
significant number of does passing the first estrus without getting
herd.
This theory is weakened somewhat by data (Elkins, unpusblished)
from a population that is hunted in November and exhibits a similar
double peak in summer fawn:doe ratios.
From extensive observations on a small herd of pronghorn in an
enclosure
(Ellis and Deblinger 1975) found that the yearling female
breeding for the first time, came into estrus and were bred one
month later than the mature females in the herd.
If this were a
common pattern in large wild populations
then it is possible the
late fawning of the younger females is responsible for the second
peak in the fawn:doe ratios.
If the factors associated with the apparent double far...'l1ing
peak are
to be elucidated, then the first step would be to verify that two
peaks in fawning do, in fact, occur.
If they do, then it is critical
to deteroine whether or not hunting during the rut is a factor.
Hanagement action that results in f awn s being born later in the
summer could very easily reduce the survival of the f awn s because
it is generally kno~m that smaller fawns have lower witer survival.
Problem

Definition

Low and declining f awn c do e ratios on the eastern plains may be a symp t orn
of a more serious environ~ental
problem.
Low annual production invariably
results in more conservative ha rve s t s wh i ch may only compound the

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�-459-

problem (Salwasser
causitive factors.

and Pojar

1979) without

an understanding

of the

Several major hypotheses can be generated with relation to the
problem of low natality and/or high mortality of young in pronghorn
population.

B.

1.

Measurement of annual
counts are inaccurate

2.

Pronghorn
pregnancy
resulting

3.

Lack of genetic variability results in population symptoms that
are similar to low nutritional plane, i.e. sensitivity to
environmental stress resulting in reduced demographic vigor.

4.

Diseases such as leptospirosis
rates or natality rates.

5.

Hunting during the pronghorn rut either interrupts
behavior sufficiently to depress conception rates,
of hunting impedes the physiology of reproduction.

6.

Predation of newborn
survival rates.

pronghorn production i.e. classification
and tend to underestimate
real productivity.

nutrition during the critical last trimester of
and/or the period from birth to weaning is inadequate,
in low in utero or post natal fawn survival.

fawns

or brucellosis

significantly

reduces

fawn survival

rutting
or the stress

pronghorn

OBJECTIVE
To determine causes for low fawn:doe
in southeastern Colorado.

C.

reduces

EXPECTED

RESULTS

ratios

of pronghorn

populations

OR BENEFITS

The successful determination of factor(s) limiting pronghorn fawn
production and/or survival will allow DOW wildlife managers to develop
more realistic strategies for affecting pronghorn population increases
consistent with strategic plan goals.
D.

APPROACH
Existing hypotheses of factors limiting pronghorn production will be
examined and ranked in priority of probable importance.
Study plans
will be prepared to test the priority hypotheses.
As these study plans
are developed they will be appended to and become a part of this document.
The following are brief procedural outlines of methods for examining
the major hypotheses.

1.

Production

measurement

error.

A sampling scheme using strips flown by helicopter will be designed
and tested with the object of estimating fawn:doe estimates within
plus or minus 10 percent of the true population value at the 95%
confidence level.

�-460-

Repetitions and cross strips may be flown to determine
of the method.
2.

Low nutritional

reliability

plane.

Blood samples, weights, and measurements will be taken from
live-trapped pronghorn during the spring and compared with
like parameters obtained from northwest region pronghorn.
Eviscerated carcass weight of hunter harvested animals will
be compared by sex and age-class for the two areas in the
fall. If there is any indication that a difference in
these measurements exists between the two areas then forage
diversity and yield potentials for these habitats will be
initiated using a method similar to that of Tiedeman (1978).
In addition, the fetal rate of live-trapped female pronghorn
from both populations will be determined by ultra-sound scanning.
3.

Lack of genetic variability.
Blood, liver, kidney, and muscle tissue samples will be
collected from hunter harvested pronghorn from the two areas.
These samples will be subjected to electrophoretic genetic
screening.

4.

Disease
In conjunction with the brucellosis-leptospirosis
survey
pronghorn populations statewide will be monitored for incidence
of positive reactors and this will be related to productivity
of the various populations.

5.

Hunting

disturbance

Two or more pronghorn populations will be selected with one
hunted during the rut and the other hunted post-rut.
Pregnancy
and fetal rates for each population will be monitored using
ultra-sound scanning.
After three years of this regime the
treatments will be reversed and comparisons will be made for
treatment effects due to hunting.
6.

Predation

on newborn

fawTIs.

..

This hypothesis will not be persued unless substantially
evidence that it is a serious factor surfaces.

more

Schedule
Fiscal Year

Period

Activitv

1978-79

Problem analysis
plan preparation

1979-83

Test measurement error hypothesis
Initial tests for low nutritional plane
Collect initial sample of blood and
tissue for genetic screening

and seudy

Jan.-June

�-461-

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�-462-

Schedule
(Cont'd)
Activity

Fiscal Year
1979-83

(Cont'd)

1983-88

Period

Test reliability of ultrasound scanning in detecting
fetal rate.
Relate incidence of disease
with productivity statewide

July-June

Further testing of nutritional
hypothesis if necessary
Test hunting disturbance hypothesis

July-June

Personnel
Thomas M. Pojar

Principal

Estimated

Investigator

Annual Cost
Cost

(01) Personal Services
(21) Operating Supplies and Services
(28) Travel Expenses
(31) Capital Expenditures
Total
E.

$12,000.00
$ 7,000.00
1,500.00
500.00
$21,000.00

LOCATION
This study initially will locate in the Southeast Region with a
study area for comparison purposes located in the Northwest Region.
Specific areas will be selected as individual study plans are prepared and maps of study area locations "Jill be included in study
plans.

F.

RELATED

FEDERAL PROJECTS

W-126-R,

Work Plan 1, Job 3.

�-463-

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�-465-

LITE~~TURE

CITED

Anderson, A. E., D. c. Medin, and D. P. Gchs.
1969.
Relationships
of
carcass fat indices in 18 wintering mule deer.
Proc. Western Assoc.
Game and Fish Comm , 49: 329-3tfO.
Ayala, F.
1968.
Genotype,
162(3861):1453-1459.

environment,

and population

numbers.

Barrett, M. W., a~d G. A. Chalmers.
adult free-ranging pronghorns.

1977.
Clinicochemical
values
Can. J. Zool. 55:1252-1260.

Bonnell, M. L., and R. K. Selander.
variation and near extinction.

1974.
Elephant seals:
Science 184:908-909.

Bowen, W. W., and W. D. Dawson.
variation in old field mice
J. ~uillffial.
58(4):521-530.

Science

for

Genetic

1977. Genetic analysis of coat pattern
(Peromyscus polionotus) of western Florida.

Bradfield, R. B.
1968.
Changes in hair associated with protein-calorie
malnutrition.
In McLance, R. and E. ~1. Widdowson (eds.).
Calorie
deficiencies
and protein deficiencies.
London, Churchill (from
Godbee 1979).
Caughley, G.
1976.
Wildlife management and the dynamics of ungulate
populations.
In Coaker, T. H. (ed.).
Applied Biology, Vol. I.
Academic Press, New York.
358pp.
Caughley, G.
1977.
Ana Ly sLs of vertebrate
and Sons, New York.
234pp.
Chai,

C. K.
1959.
50(5):203-208.

Life span in inbred

populations.

and hybrid

Colorado Division of Wildlife.
1977.
Today's
Wildlife.
Colorado Division of Wildlife.
96pp.

mice.

John IHley

J. Heredity

strategy ...Tomorrow's
Planning Section, Denver.

Ellis, J. E.
1970.
A computer analysis of f awn survival in the pronghorn
antelope.
Ph.D. Thesis.
University of California, Davis.
70pp.
Frelinger, J. A.
1972.
The maintenance of transferrin
pigeons.
Proc. Nat. Acad. Sci. 69(2):326-329.

polymorphism

in

Godbee, R. G., L. M. Slade, and L. M. Lawrence.
1979.
Use of protein
blocks containing urea for minimally managed broodmares.
J. Animal
Sci. 48(3):459-463.
Haaland, G. L., J. K. Matsushima,
C. F. Nockels, and D. E. Johnson.
1977.
Bovine hair as an i~dicator of calorie-protein
status.
J. Anim.
Sci. 46:826.
(irom Godbee 1979).

�-466-

Hay, K. G., G. N. Hunter,
Colorado 1949-1958.
No.8.
112pp.

and L. Robbins.
1961.
Big ga~e management in
Colorado Dept. of Game and Fish, Tech. Bull.

Hoover, R. L., G. E. Till, and S. Ogilvie.
1959.
Colo. D~pt. Fish and Game. Tech. Bull. No.4.

The antelope
1l0pp.

Humphrey, R. R.
234pp.

Press

1962.

Range

ecology.

The Ronald

of Colorado.

Co., New York.

Hanlove, M. N., J. C. Avise, H. O. Hillestad. P. R. Ramsey, M. H. Sruith, and
D. O. Straney.
1975.
Starch gel electrophoresis
for the study of
population genetics in white-tailed
deer.
Proc. 29th Ann. Conf. SE
Game and Fish Commrs.
29:392-403.
Manlove, M. N., M. H. Smith, H. O. Hillestad, S. E. Fuller, P. E. Johns, and
D. O. Straney.
1978.
Genetic subdivision in a herd of white-tailed
deer as de~onstrated by spatial shifts in gene frequencies.
Proc.
Ann. Conf. SE Assoc. Fish and Wildl. Agencies.
30:487-492.
Martin, Y. G., ~.].J. Miller, and D. H. B'lacknom , 1967.
Wound healing,
hair growth, and biochemical measures as affected by subnormal protein
and energy intake in young cattle.
Amer. J. Vet. Res. 30:355.
(from
Godbee 1979).
Maynard, L. A., and J. K. Loosli.
1969.
Ani~al
McGra~-Hill
Book Co., New York.
613pp.

nutrition,

6th edition.

Morgan, R. P. II, J. A. Chapman, L. A. Noe, and C. J. Henny.
1974.
Electrophoresis as a management
tool.
Trans. NE Section of The Wildlife
Society, 31st NE Fish and Wildlife Conf. p. 63-71.
Nei, ~'" T. ~laruyama, and R. Chakraborty.
1975.
The bottleneck effect
and genetic variability
in populations.
Evolution 29(1):1-10.
Nutrition Research.
1968.
Objective
kwa sh Lc rko r . Nutr. Res. 26:330.

measurement of hair changes
(from Godbee 1979).

O'Cara, B. W.
1970.
Intra-uterine
mortality in the pronghorn.
of the 4th Antelope States l.Jorkshop. 82-92.

in

Proceedings

Ramsey, P. R., J. C. Avise, M. H. Smith, and D. F. UrJston.
1979.
Biochemical variation and genetic heterogeneity
in South Carolina
populations.
J. Wildl. Hanage. 43(1): 136-142.
Redfield, J. A.
1974.
Genetics and selection at the Ng locus
(Dencragaous obscurus).
Heredity 33(1):69-78.
Ryrnau ,

deer

in blue grouse

N., C. Be ckman , G. Brunn-Petersen,
and C. Reuterwall.
1977. Variability
of red cell enzymes and gEnetic implications of management policies in
Scandinavian
moose CAlces alces) Hereditas 85:157-162.

�-467-

Salwasser, H., and T; M. Pojar.
populations.
Second Draft.
Schwartz, C. C. 1977.
prairie, Colorado.
Collins.
113pp.

1979. Simulation modeling
Xerox copy.
73pp.

of pronghorn

Pronghorn grazing strategies on the short grass
Ph.D. Thesis, Colorado State University, Fort

Seal, U. S., and R. L. Hoskinson.
1978. Metabolic indicators of habitat
condition and capture stress in pronghorn.
J. Wildl. Manage. 42(4):
755-763.
Seal, U. S., ~. E. Nelson, L. D. Mech, and R. L. Hoskinson.
1978. Metabolic
indicators of habitat differences in four Minnesota deer populations.
J. Wildl. Manage. 42(4):746-754.
Selander, R. K. 1970.
American Zoologist

Behavior and genetic variation
10(1):53-66.

in natural

populations.

Simon, R. C. 1978. Fish genetics laboratory (report).
In: Thomas G.
Scott, Helen C. Schultz, and Paul H. Eschmeyer (eds.).
Sport Fishery
and Wildlife Research.
1975-76.
U.S. Fish and Wildlife Service, U.S.
Government Printing Office, Denver, Colorado.
Smith, H. H., H. O. Hillestad, H. N. Hanlove, R. L. Harchinton.
1976. Use
of population genetics data for the management of fish and wildlife
populations.
41st N. Amer. "\..'ildl.
and Nat. Resources Conf. p. 119-133.
Smith, M. H., H. O. Hillestad, M. N. Manlove, D. O. Straney, and J. M. Dean.
1977. Xanage~ent implications of genetic variability in loggerhead
and green sea turtles.
XIII International Congress of Game Biologists.
p. 302-312.
Thomson, A. M., and W. Thomson.
1943. Lambing in relation
the pregnant ewe. Brit. J. Nutrition, 2:290-305.

to the diet of

Tiedeman, J. A. 1978. Middle Park inventory analysis; habitat type
developments.
USFS Rocky Mountain Forest and Range Exp. Sta. Res.
Paper.
(In Press).
Trout, L. E. 1976. Blood analysis of Id2ho pronghorn.
Proceedings
the 7th Biennial Pronghorn Antelope Workshop.
pI22-126.
Verme, L. J. 1969. Reproductive patterns of white-tailed
nutritional plane.
J. ~.J'ildl.
Manage. 33(4) :381-887.

of

deer related

to

�-468-

Howe,

D. L.
1970.
Hiscellaneous
bacterial diseases.
In Davis, J. W.,
L. H. Karstad, D. O. Trainer (eds.).
Infectious diseases of Wild
HalT!ITlals.Iowa State University Press, Ames.
421pp.

Adrain, W. J.
1976.
Brucellosis-Leptospirosis
survey of big game
animals in Colorado.
In Game Research Report, Part 2, Colorado
Division of Wildlife pp. 555-559.
Trueblood, M. S., and G. Post.
1959.
Vibrosis as a factor in the
reproduction
of antelope (Antilocapra americana).
J. An.irn,Vet.
Med. Assoc. 13~:562.
Ellis, J. E., and R. D. Deblinger.
1975.
Delayed breeding
in young pronghorns.
Manuscript.
4 xerox pages.
Prenzlow, E. J., D. L. Gilbert,
patterns of the pronghorn.
Division of Game Research.

and parturition

and F. A. Glover.
1968.
Some behavior
Colo. Dept. of Game, Fish and Parks,
Special Report No. 17.
16pp.

�-469-

COLORADO

FEDE~~L AID PROJECT W-130-R

ENVIRONMENTAL

ASSESSMENT

Pronghorn

Ecology

Studies:

Pronghorn

Population

Study

Program Leader
R. Bruce Gill, Wildlife Research

Supervisor
Harold M. Swope, Hildlife

Leader

Manager/Research

for

Colorado

Department of Natural
Division of Wildlife

Active Dates:

July 1, 1979-June

Resources

30, 1988

Prepared

May 15, 1979
Fort Collins, Colorado

�-470-

Table

of Contents
Page

I.

Introduction

1

II.

Proposed

1

III.

Description

IV.

Environmental

V.

Mitigating

Measures

VI.

Unavoidable

Adverse

VII.

Irreversible

Commitment

VIII.

Alternatives

to the Current

IX.

Consultation

and Coordination

Action
of the Environment

2
2

Impacts

4
5

Impacts
of Resources
Program

5

5
5

�-471-

SUMMARY

This report describes a research program designed to determine the
cause of reduced reproduction and/or fawn survival of a pronghorn population
in Southeastern Colorado.
The number of viable fa,vns produced by this
population has declined at an alarming rate over the past ten years.
It
is generally accepted that a decline in a populations ability to produce
young is a sure symptom that the population is experiencing environmental
stress.
Many times this stress is the result of man's activities, therefore
if the factor(s) can be identified, remedial action can also be identified
and initiated.
Some of the factors that could cause reduced reproduction
in pronghorn antelope are:

1.

Low nutritional plane during the critical last trimester of
pregnancy and/or the period from birth to weaning is inadequate,
resulting in low in utero or post natal fawn survival.

2.

Lack of genetic variability results in population symptoms that
are similar to low nutritional plane, i.e. sensitivity to environmental stress resulting in reduced demographic vigor.

3.

Diseases related to abortion or fawn mortality,
brucellosis, vibriosis, and Johne's Disease.

4.

Hunting during the pronghorn rut either interrupts
behavior sufficiently to depress conception rates,
of hunting impedes the physiology of reproduction.

5.

Predation
rates.

of newborn

fawns significantly

reduces

such as leptospirosis,

rutting
or the stress

pronghorn

survival

Minor negative impacts are expected to the human environments within
the study areas however, none of these are judged to be irreversible.
Substantial
positive aesthetic and financial impacts will be realized in future years with
the successful completion of this program.

�-472-

I.

Introduction
The number of young animals added to a population annually is determined
by the intrinsic reproductive rate of the species and the neonatal survival
rate.
The pronghorn antelope (Antilocarpa americana) have an intrinsic
reproductive capacity that is among the highest of any ungulate in Colorado.
The pronghorn possess the ability to produce many young, consequently the
population can expand rapidly under ideal conditions.
Likewise, the young
are most susceptible to mortality when conditions are less than ideal
which keeps the population from expanding beyond the ability of the
habitat to support it.
Therefore, an important manifestation
that the
population is functioning under environmental
stress is a reduction in
the number of young born and/or a reduction in young survival.
The
Colorado Division of Wildlife estimates the population production by
making late summer herd structure counts.
These counts are made from the
air (usually helicopter) and the animals are classified as to bucks (males
one year and older), does (females one year and older), and fawns (young
of the year).
An index of the reproductive success of the population for
that fawning season is gained by expressing the ratio of the total number
of fawns observed in relation to the total number of does observed in
terms of fawns per 100 does.
The fawn:doe ratio in a demographically
vigorous population in Western United States is in the neighborhood of
90 fa,vns per 100 does (Ellis, 1970).
The fawn:doe ratio in Southeastern
Colorado exhibits a statistically significant decline since 1967.
In
recent years fawn:doe ratios in the 30's and 40's are not uncommon.
The
factor(s) responsible for low fawn:doe ratios are likely associated with
nutrition, disease, genetic composition of the population, or abnormally
high predation.

II.

Proposed

Action

When an apparent anomaly in a wild population is detected it is a challenge
of appreciable magnitude to isolate the factor(s) that are responsible.
One of the more fruitful methods is to compare population parameters and
environmental
conditions of the affected population with a population
that is in a healthy state.
In the Northwest region of Colorado the
observed fawn:doe ratios are consistently in the 70's and 80's which
are the highest reported anywhere in the state.
Therefore, a pronghorn
population from the Northwest will be compared with one that exhibits
one of the sharpest declines in fawn:doe ratio from the Southeast region,
The nutritional plane of the reproductive female has been associated
Hith ovulation and fertilization
success; fetus viability and early
post natal survival, and lactation which affects subsequent summer fawn
survival.
The nutritional plane of the Southeast and Northwest populations will be compared by weights, measurements
and blood parameters
that are indicators of nutritional intake.
Weights and measurements will
be collected from hunter harvested animals and from live-trapped animals.
Blood samples will be taken from the live-trapped animals.
Diseases that are known or suspected to affect the reproductive success
of pronghorn populations are:
leptospirosis,
brucellosis, vibriosis,

1

�-473-

and Johne's Disease.
Serological tests are available to reliably
test for the presence of these disease organisms, with the exception
of Johne's Disease.
Blood samples from live-trapped pronghorn
will be collected to test for disease.
This project will work
cooperatively with the Pathology Department of Colorado State University to develop methods of detecting Johne's Disease in pronghorn.
Blood and tissue samples will be collected from hunter harvested
animals to estimate genetic diversity of the two populations being
Objectives

III.

studied.

of this study are:

1.

Test current

method

2.

Evaluate nutritional
it to reproduction.

3.

Determine the incidence of diseases
reproduction or survival of fawns.

4.

Screen

for genetic

Description

of estimating
plane

fawn:doe

of pronghorn

diversity

ratios.

populations

that are known

of two populations

and relate

to affect

selected

for study.

of the Environment

The pronghorn range in the Southeast region is characterized
by rolling
short-grass prairie with little topographic relief.
The greatest proportion of the area is arid grassland used for grazing domestic livestock.
Along the Arkansas River there is some irrigated farmland.
The human
population for the area as a whole is very sparse with highest densities
along the Arkansas River.
The Northwest region is also very sparsely inhabited by humans.
The
area is used for grazing domestic livestock but at very low stocking
rates because of its meager production of forage species utilized by
domestic livestock.
The terrain has much more topographic relief than
the Southeast, but it would still not be considered steep or precipitous.
The predominant vegetative type is the sagebrush steppe type.
IV.

Environmental

Impacts
Aerial

Surveys

Surveys to estimate pronghorn population parameters will be done by
helicopter.
The impacts associated with helicopter surveys are
expected to be:
a) noise from a low flying helicopter will be a
disturbance to human inhabitants and will cause an escape reaction in
wild and domestic livestock,
b) observance of a low flying helicopter
may cause anxiety on the part of humans that an emergency of some
sort exists,
c) low flying aircraft within sight of roads may distract
motorists and result in a motor vehicle accident, d) landing of the

�-474-

helicopter for refueling will cause soil impaction and vegetation
trampling in the immediate area of the refueling operation, e) fuel
spillage will defoiliate and possibly kill the vegetation directly
affected.
Trapping

Operation

and Blood

Collection

Live-trapping
pronghorn will be done in late winter and will be
accomplished by driving them into a corral trap with a helicopter.
Considerable
trampling of the vegetation and possibly some wind and/or
water erosion is expected as the result of a trapping operation.
Trapping will intensively affect one of two acres with a less intense
affect on about five acres surrounding the trap.
Pronghorn will be driven by helicopter up to five miles to the trap
site.
Although pronghorn have evolved as the high-speed, long-range
runners of the prairie, being driven by helicopter will stress them
to some degree, if not physically at least psychologically.
Once in
the trap, handling will cause some degree of physical and psychological
stress and could result in two to five percent mortality.
A sample of about 30 females from the live-trapped animals of each
population will be immobilized with the drug M-99 and the tranquilizer
rompun.
These females will be scanned by ultra-sound equipment to
detect the number of fetuses they are carrying.
The ultra-sound scan
is harmless so the only impact involved is the affect of the drug.
Ungulates have a wide tolerance for M-99, but dosages for pronghorn
have not been well documented so it is possible some mortality could
result.
A blood sample (10 ml) will be collected via the jugular vein of each
live-trapped animal.
No impact is expected from these collections
because approved methods in veterinary science will be used to collect
the samples.
Most of the diseases that can be detected by seriologic
tests are transmissable
to domestic livestock.
If pronghorn are
carriers then detection and remedial action would be a tremendous
positive financial impact on the domestic livestock industry.
Collection

of Data from Hunter

Harvest

Pronghorn

Hunters will be inconvenienced
by the necessity to stop at hunter check
stations to allow data to be collected from the animal they harvested.
Other hunters will be contacted either in the field or in their camps.
Collection of the necessary data should take between 15 and 30 minutes.
A positive impact will be a better informed public because of personal
contact with individuals to explain the need for pronghorn research
and management to maintain vigorous and healthy natural populations.

�-475-

V.

Mitigating

Measures
Aerial

Surveys

If field refueling is necessary it will be done on areas that are
already void of vegetation so the effect of trampling or spillage will
be minimized.
Such areas might be roads that are used only intermittently
or suitable arroyos.
Any areas where domestic livestock are confined will be avoided by at
least a half mile lateral distance and if a group of domestic livestock
are in the line of travel we will attempt to gain sufficient altitude
to minimize the escape reaction of the animals.
In no instance will a
flight be made directly over an inhabited farmstead; at least a half
mile lateral distance will be allowed.
An attempt to contact all
inhabitants on the study areas will be made to explain the project and
the need for helicopter flights.
This hopefully will alleviate some
of the anxiety of hearing a low flying helicopter.
The flights will be on pre-determined
flight lines so there will be
no actual pursuit of pronghorn groups.
Unless a group of pronghorn
would run directly along the flight line, no group should be harassed
for more than five minutes.
Trapping

and Blood

Collections

All precautions that are kno\vu to reduce pronghorn trap mortality will
be employed.
One of the major precautions being the lining of trap
fences with burlap which eliminates challenging of the fences by
the animals.
If pronghorn are to be held for any length of time they
will be put in darkened, confined enclosures which has a calming affect
on them.
The trapping will mostly be done before
dormancy which will minimize the affect

plants emerge
of trampling.

from winter

Much of the information necessary for this project can be obtained
from animals trapped by regional management operations.
These activities
are not sponsored by Federal Aid funds.
Most workers agree that immobilization of pronghorn with drugs is more
risky then with other ungulates because of their highly variable
physiological
state which affects their reaction to drugs.
M-99 was
chosen as the drug to use because of the wide tolerance margin ungulates
exhibit for it. The drug will be administered with a hand held syringe
rather any form of projectile which will eliminate some of the hazards
and ensure that the prescribed dose is administered.
Careful records
will be kept of dosages and animal response to better delineate the
acceptable dosage range.

�-476-

For the ultra-sound scan, the immobilized animal will be positioned
on its side with an attendant assigned strictly to maintain its head
position to avoid the possibility of aspirating rumen fluid.
The
animal will be blindfolded with a soft fabric to reduce the chance
of eye injury and further alleviate any stress associated with handling.
There is no pain caused by the ultra-sound scan and no harmful side
effects.
It is used regularly on unaesthesized
humans and has been
proven harmless to human fetuses.
VI.

VII.

Unavoidable

Adverse

Impacts

1.

Hithin the corral traps the vegetation ~vill be damaged by the sharp
hoofs of the animals even if the plants are dormant.
Outside the
trap the plants will be trampled by human foot traffic but damage
is expected to be slight.

2.

Two to five percent trap mortality is expected.
Of the animals
that are immobilized, it is expected that five to ten percent will
suffer mortality.

3.

LO'tvlevel helicopter flights will cause a few instances of disturbance
to humans.
This will be quite limited, however, since so few humans
inhabit either study area.

4.

Helicopter flights will cause temporary disruption
and possibly domestic livestock activities.

5.

Collection of data from hunter harvested animals will detain
divert the hunters activities for 15 to 30 minutes.

Irreversible

Commitment

Alternatives

or

of Resources

There will be no irreversible
of this project.
VIII.

of pronghorn,

to the Current

commitment

of resources

as the result

Program

The objective of this project is to identify the factors related to an
alarming decline in the productivity of Southeast region pronghorn
populations.
Some of the alternatives to this proposed project are:

IX.

1.

Do not fund the project

2.

Do not conduct

3.

Conduct only the segments
no negative impacts.

Consultation

with Federal

Aid money.

this study.
of this study

that are judged

to have

and Coordination

The general concept of this program has been reviewed
and management personnel within the Colorado Division

with administration
of Wildlife.
In

�-477-

addition, the following have been consulted for information
of the more technical aspects of the program.
Dr. David C. Bowden, Statistical
Colorado State University.

Analysis,

Department

Beth Williams, Physiology of Drug Immobilization,
Department, Colorado State University.
Dr. James E. Ellis, Pronghorn Behavior,
Laboratory, Colorado State University.
Dr. Donald J. Nash, Genetics,
Colorado State University.

Department

Natural

on some

of Statistics,

Veterinary

Resources

of Zoology

Science

Ecology

and Entomology,

��-479-

JOB PROGRESS

State of

COLORADO
------------------------------

Project

No.

W-126-R-2

.Work Plan No.
Job Title
Period

6

Seasonal

Covered:

Dietary

Selection

July,

1979

REPORT

Bighorn

Sheep-Mountain

Job NO.

~l

_

of Rocky Mountain

Goats

and Forage

Quality

Goat Investigations

July 1, 1978 - June 30, 1979

Personnel:
T. V. Dailey, T. N. Woodard, R. B. Gill, J. E. Ellis, M. L. Stevens,
N. T. Hobbs, D. L. Baker, D. Cannalte, K. Pals, University of Colorado Mountain
Research Station Personnel, G. D. Bear, S. J. Dailey.

ABSTRACT
Work during the 1978-1979 segment was aimed primarily at determining the
dietary selection of hand-reared Rocky Mountain goats in the alpine tundra
vegetation zone.
This involved capturing, rearing, and training three
mountain goat kids for use in grazing trials during the winter of 1978-1979.
Across all months forbs were the most frequently chosen forage class.
Graminoids were somewhat less important in the diet relative to forbs.
Salix planifolia accounted for virtually all of the browse consumed, but
was essentially the only browse species available on the study area.

��-481-

SEASONAL DIETARY SELECTION AND FORAGE
QUALITY OF ROCKY MOUNTAIN GOATS
T. V. Dailey and T. N. Woodard

P. N. OBJECTIVES
1.

To describe
goats.

and quantify

the seasonal diet selection of Rocky Mountain

2.

To evaluate the chemical constitution
selected by Rocky Mountain goats.

and digestibility

of forage

SEGMENT OBJECTIVES
1.

Obtain, rear, and train mountain

goat kids for food habits study.

2.

Select study areas for vegetation

and food habits sampling.

3.

Conduct grazing trials during winter,

4.

Begin summer grazing trials.

spring and summer 1978-1979.

METHODS AND MATERIALS
Capture, Rearing and Training of Mountain

Goat Kids

Three mountain goat kids were captured from free-ranging nannies in the
Collegiate Range in Colorado during June-July 1978. Two kids were
captured by hand in June and the third was taken in a clover trap in July.
Capture efforts were terminated in July due to the difficulty of handling
and taming wild kids at this late date. Kids were bottle fed a formula
consisting of two parts whole milk to one part condensed milk with baby
vitamins and buttermilk being added at alternate feedings each day. Grain
and alfalfa hay were fed ad libitum.
Training for grazing trials began as soon as the kids had readily accepted
the handler.
Initially the kids were fed their milk formula in the back
of a specially enclosed pickup truck. The kids were conditioned to load
at the sound of a whistle.
Gradually trips were made with the mountain
goats to various mountain sites to accustom them to transportation and
familiarize them with native vegetation.
At this time the kids would be
unloaded and allowed to graze freely.
The whistle would then be used to
reload the kids into the pickup where they would then be given their
milk formula as a reward.
Raisins were substituted for the milk reward
once weaning was completed.
The original goal in obtaining mountain goat kids called for the capture
of at least six kids which would decrease total sample variation resulting

�-482-

Forage Evaluation
Nutritional evaluation of individual plants eaten by goats will be assessed
by chemical determinations of percent dry matter, nitrogen, acid detergent
fiber, lignin, cell wall constituents and in vitro digestibility of species
contributing at least two percent of total bites.
Fifty gram samples
were handplucked from plants in the immediate vicinity of forage grazed
by mountain goats during a trial. Plant parts were collected to represent
those eaten by the hand-reared mountain goats. At least 25 individual
plants of each species were composited to form each species sample and
were frozen for analysis at a later date.
RESULTS AND DISCUSSION
Grazing trials were conducted during November, January and March.
A trial
was attempted in May but was cancelled due to poor road conditions and
failure of the over-snow machine to navigate across icy snowpatches typical
of this season.
In November graminoids were the most frequently chosen forage class, with
Kobresia myosuroides, Trisetum spicatum, and Carex rupestris showing the
greatest use (Table 1). Forbs were chosen with slightly less frequency.
Campanula rotundifolia constituted the bulk of the forb diet. Browse
consumption was relatively small compared to graminoids and forbs.
Salix
planifolia accounted for99.97 percent of the browse consumed, and ranked
third when compared to individual plant species (Table 1). Trisetum
spicatum, Kobresia myosuroides, and Campanula rotundifolia were collected
for future chemical analysis.
Table 1. Composition of Rocky Mountain goat diet in alpine tundra type in
November 1978. Entries are percentages of total individual animal bites
for each plant species and mean percentages with 90% confidence intervals
calculated across 3 animals.

Mean + Confidence

Plant Species

Lanea

Shava

Oreo

Kobresia myosuroides

8.2

20.3

19.3

15.9 + 11.3

Trisetum

15.7

16.4

10.7

14.3 +

16.6

21.2

4.7

14.2 + 14.4

(leaves)

9.3

13.2

3.2

8.6 +

8.5

(stems)

6.9

7.8

0.8

5.2 +

6.4

(catkins)

0.4

0.2

0.7

0.4

spicatum

Salix planifolia

5.2

Interval

�-483-

from individual animal variation in diet selection.
Since, this goal was
not reached in the spring of 1978, attempts are presently (June 1979) being
made to capture four mountain goat kids to augment numbers of experimental
animals.
To facilitate this, a helicopter was used to place six clover
traps on top of Sheep Mountain in the Collegiate Range.
(*Editorial - If
you look critically at between animal and within animal variances you'll
probably discover that 6 animals are not much of an improvement over 3).

Study Area

Selection

The area selected for alpine ecosystem grazing trials was the University
of Colorado Mountain Research Station on Niwot Ridge near Nederland,
Colorado.
Niwot Ridge has areas of acceptable and accessible habitat which
are not utilized by domestic livestock, eliminating the influence of that
variable on mountain goat diet selection.
Hibbs (1965) reported that Rocky Mountain goats inhabitat alpine tundra
habitats at all seasons of the year.
Consequently grazing trials will be
conducted year round in the alpine tundra zone.
The alpine zone begins at
approximately
3505 m (11,500 ft) and continues upward in elevation.
This
zone is characterized by low growing grasses, sedges, forbs, and finger-like
patches of spruce and pine (Krummholz).
The winter study area is about
3505 m in elevation with moderately steep slopes and snow-free ridge tops.

Grazing

Trials

During the winter of 1978-1979 forage selections of the three mountain
goat kids was quantified at the Niwot Ridge winter study area.
A grazing
trial consisted of one observer counting from close distance (less than
2 m) the number of bites of each plant species consumed by one goat feeding
for approximately
one hour.
A tape recorder was used to record data during
the grazing trials.
During each trial period the observer handplucked 25 forage samples of each
species comprising two percent or more of the diet.
These "simulated bites"
approximated the size of individual real bites ingested by the foraging
animal.
An average weight per bite will be derived from the 25 simulated
bites.
Estimates of diet composition based on these weights will be compared
to diet composition derived from bite counts.
On each starting date the animals were transported from Fort Collins to
the Mountain Research Station.
During the January and March trials an
over-snow machine transported the goats to holding pens located just below
timberline.
One to two days of pre-trial grazing were conducted to allow
mountain goats to sample and adjust to available forage, as well as permit
dry run data collection by the observer.
After this pre-trial period
three grazing trials were conducted each day until approximately
3,000
bites were documented for each animal.
This normally required about three
days of trials depending on weather conditions.

�-484-

Table 1. Composition of Rocky Mountain goat diet in alpine tundra type
in November 1978. Entries are percentages of total individual animal bites
for each plant species and mean percentages with 90% confidence intervals
calculated across 3 animals.
(Continued).

Lanea

Shava

Oreo

Carex rupestris

17.0

3.7

16.8

12.5 + 12.9

Campanu1a

rotundifo1ia

9.7

13.3

13.0

12.0 +

6.7

Arenaria

fend1eri

2.9

7.8

5.0

5.2 +

4.1

Oreoxis

alpina

10.4

0.0

4.5

5.0

+ 8.8

6.6

1.8

6.2

4.9

+ 4.5

Geum rossii

Mean

+ Confidence

Plant Species

Pedicu1aris

pa1ustris

4.8

1.3

1.5

2.5 +

3.3

Trifolium

dasyphy11um

2.1

5.1

0.9

2.4 +

3.7

% Graminoids

43.9

41.9

54.0

46.6

+ 10.9

% Forbs

39.4

35.7

38.9

38.0

+ 3.4

% Browse

16.6

21.5

5.7

14.6 + 13.6

% Unknown

o

0.9

1.5

Total Bites

2,336

2,565

2,208

Interval

7,109

During January, graminoids and forbs were chosen with similar frequency.
Of
the graminoids, Carex rupestris, Trisetum spicatum, and Kobresia myosuroides
showed the greatest use (Table 2). Campanu1a rotundifo1ia constituted a
large portion of the forb intake.
Browse consumption was relatively small
compared to graminoids and forbs.
Salix p1anifolia accounted for all of the
browse consumed, and ranked fifth when compared to individual plant species
(Table 2). All plant species listed in Table 2 were collected for future
chemical analysis.
During March forbs were the most frequently chosen forage, with Trifolium
dasyphy11um being especially important (Table 3). Graminoids and browse
were chosen with considerably less frequency.
Carex rupestris was the
most frequently chosen individual graminoid.
Salix p1anifo1ia constituted
virtually all of the browse diet, and ranked fourth when compared to individual
plant species (Table 3). All plant species listed in Table 3 were collected
for future chemical analysis.

�-485-

Table 2. Composition of Rocky Mountain goat diet in alpine tundra type in
January 1979. Entries are percentages of total individual animal bites for
each plant species and mean percentages with 90% confidence intervals
calculated across 3 animals.
Species listed are those which contribute 2
percent or more of total bites.

Mean + Confidence

Plant Species

Lanea

Shava

Oreo

Carex rupestris

46.5

7.8

7.6

22.5 + 37.8

Campanula

rotundifolia

13.6

15.1

23.6

16.4 +

Trisetum

spicatum

3.5

20.6

12.9

12.4 + 14.4

Kobresia

myosuroides

9.2

11.1

2.8

8.5

+

7.3

7.3

4.9

11.3

7.8 +

5.4

(leaves)

3.5

2.9

7.6

4.7 +

4.3

(stems)

3.8

2.0

2.7

2.8 +

1.5

(catkins)

0.0

0.0

1.0

0.0

0.0

Geum rossii

7.9

5.6

10.0

7.5 +

3.7

Oreoxis alpina

4.0

2.6

7.0

4.5 +

3.8

Arenaria

obtusiloba

0.05

7.6

1.3

3.4 +

6.8

Arenaria

fendleri

1.5

3.4

2.8

2.6 +

1.6

Trifolium

dasyphyllum

0.7

3.7

1.1

2.0 +

2.7

Polygonum

bistortoides

1.1

3.0

1.5

2.0+

1.7

% Graminoids

61. 2

47.2

25.6

44.7 + 30.2

% Forbs

31.4

47.8

62.7

47.3 + 26.4

% Browse

7.3

4.9

11.3

7.8 +

Total Bites

3,790

3,966

2,208

9,964

Salix planifolia

Interval

9.3

5.4

�-486-

the browse consumption.
From November to March, the percent of forbs and
graminoids in the mountain goat diets showed a definite trend as forb
consumption increased while grass consumption decreased (Figure 1). The
percent of browse in the diets showed some small changes during the winter
period.
Table 4. Percent composition of mountain goat diets across 3 animals in
alpine tundra during winter 1978-79. Entries are mean percentages with
90 percent confidence intervals for plant species contributing 2% or more
to the total winter diet.
Species are ranked in order of importance across
all months.

Carex

rupestris

November

January

March

All Months

12.2 + 12.9

22.5 + 37.8

9.5 + 12.0

14.6 + 20.6

Trifolium

dasyphyllum

2.8 +

3.7

2.0 +

2.7

24.2 + 24.5

10.6 + 10.4

Campanula

rotundifolia

12.0 +

6.7

16.4 +

9.3

3.9 +

5.9

10.4 +

1.3

Trisetum

spicatum

14.4 +

5.2

12.4 + 14.4

3.4 +

3.3

9.3 +

6.2

14.6 + 14.4

7.2 +

5.4

8.1 + 10.9

9.2 +

6.0

(leaves)

9.3 +

8.5

4.4 +

4.3

7.0 +

9.0

6.6 +

1.8

(stems)

5.3 +

6.4

2.8 +

1.5

1.1 +

2.0

2.7 +

2.6

16.0 + 11.3

8.5 +

7.3

2.5 +

2.5

7.9 +

2.8

Oreo.1{
is alpina

4.8 +

8.8

4.1 +

3.8

9.4 + 10.7

6.7 +

3.0

Geum rossii

4.8 +

4.5

7.5 +

3.7

6.2 +

4.4

6.5 +

3.3

Arenaria

obtusiloba

0.0

3.4 +

6.8

6.3 +

9.9

3.8 +

4.7

Arenaria

fendleri

5.4 +

2.6 +

1.6

3.1 +

0.7

3.5 +

1.5

Senecio

canis

0.0

1.0+

1.0

4.8 + 10.6

2.7 +

6.0

Salix planifolia

Kobre~ia

myosuroides

4.1

% Forbs

37.9 +

3.4

44.9 + 26.4

72.8 + 21.2

55.0 + 19.1

% Graminoids

46.3 + 10.9

47.8 + 30.2

19.1 + 10.4

35.4 + 13,.7

% Browse

14.6 + 13.6

7.2 +

8.1 + 10.9

7.4 + 11.0

11,485

28,558

Total Bites

7,109

5.4

9,964

�-487-

Table 3. Composition of Rocky Mountain goat diet in alpine tundra type in
March 1979. Entries are percentages of total individual animal bites for
each plant species and mean percentages with 90% confidence intervals
calculated across 3 animals.
Species listed are those which contribute 2
percent or more of total bites.

Lanea

Shava

Or eo

Trifolium

28.0

36.1

7.9

24.2 + 24.5

Carex rupestris

17.4

8.1

3.4

9.5 + 12.0

Oreoxis alpina

4.4

7.1

16.5

9.4 + 10.7

Salix planifolia

15.1

7.5

2.2

8.1 + 10.9

12.6

6.9

1.9

7.0 +

9.0

2.2

3.7

13.0

6.3 +

9.9

Geum rossii

4.0

5.5

9.1

6.2 +

4.4

Senecio canis

0.1

2.4

11.9

4.8 + 10.6

Campanula

rotundifolia

8.1

1.8

2.3

3.9 +

5.9

Heuchera

spp.

0.0

3.4

7.1

3.5 +

6.0

Trisetum

spicatum

1.1

4.3

4.7

3.4 +

3.3

Arenaria

fendleri

3.1

2.7

3.5

3.1 +

0.7

Kobresia myosuroides

4.2

1.9

1.5

2.5 +

2.5

Polygonum

2.6

2.2

1.6

2.1 +

0.8

% Forbs

59.6

73.3

84.7

72.7+21.2

% Graminoids

25.4

19.2

13.0

19.1 + 10.4

% Browse

15.1

7.5

2.2

8.1 + 10.9

Total Bites

3,624

4,068

3,793

dasyphyllum

(leaves)
Arenaria

obtusiloba

bistortoides

Mean

+ Confidence Interval

Plant Species

Across all months forbs were the most frequently chosen forage class.
The
grass Carex rupestris was ranked the highest when compared to individual
plant species (Table 4). Graminoids were somewhat less important in the
diet relative to forbs.
Salix planifolia accounted for virtually all of

�-488-

LITERATURE

CITED

Hibbs, L. D. 1965. The mountain goat of Colorado.
State Univ., Fort Collins.
152pp.

Prepared

by:

Approved

by:

~;~~~:'r~Y
?r- f(?L'f~

~~:~~.t/ev~es::rc:
;J:~e2:_,ij
Thomas N. Woodard
Wildlife Researcher

I

M.S. Thesis.

Colorado

�-489-

70

60

50

.w
&lt;lJ
.--!

~

j::;
'r-!

40

.w

C
&lt;lJ

..,

U
Q)

c,

30

20

10

Shrubs

November

Figure

1.

January

March

Diet by forage category of 3 tame Rocky Mountain goats
during w i nr e r 1978-79 in alpine
tundra
on Niwot Ridge,
Colorado.

��-491-

JOB PROGRESS

State of

--------~~~~~----------

Project

No.

1979

REPORT

COLORADO

Job Title

Big Game Investigations

W-126-R-2

Work Plan No.

Period

July,

7

J~

~.

1

----------~---------------------

B~l~a~c~k~B~e~a~r~I~n~v~e~s~t~i~g~a~t~i~o~n~s~

Covered:

July 1, 1978 through

_

June 30, 1979

Personnel: Thomas D. I. Beck, Rick Danvir,
Green, Dan Miller, Bill Boydstun; Colorado

James Haskins, Dave Kenvin,
Division of Wildlife

Larry

ABSTRACT
The Black Mesa-Crystal Creek area in west-central Colorado was selected as
the principal study area.
A closed season on bears was obtained from the
Colorado Wildlife Commission for the study area and a buffer zone.
Capture
efforts began on 29 May 1979 and by end of segment four bears were captured
a total of six times.
Capture success was lower than expected.
The use of
a mixture of Ketamine and Rompun has proved very satisfactory.

��-493-

BLACK BEAR INVESTIGATIONS
Thomas D. I. Beck

P. N. OBJECTIVES
1.

Develop techniques
populations.

to accurately

2.

Determine

3.

Describe black bear population dynamics sufficiently to allow
analysis of various harvest and habitat manipulations.

habitat preferences

and precisely

estimate black bear

of selected black bear populations.

SEGMENT OBJECTIVES
Same as P. N. Objectives.

METHODS AND MATERIALS
Study Area Selection
Field personnel with the Colorado Division of Wildlife (DOW) were contacted
for suggestions on study areas. Harvest data for the previous 30 years were
reviewed as well. General surveys were conducted in 5 possible study areas.
Surveys primarily consisted of looking for bear sign, vehicle access, and
the proportion of desired vegetation types (spruce-fir, aspen, Gambel's oak).
Discussions with personnel from DOW and U.S. Forest Service were conducted
to get a feeling for the receptivity of a bear study by local management
people.
This was of great importance when it became evident that all bear
hunting would have to be stopped in the study area during the duration of
the study. The need for a bear hunting closure severely restricted the choice
of study areas as it would have been difficult to obtain a closure on areas
of moderate to high historic harvest and areas frequented by bear outfitters.
Finally, an area without domestic sheep grazing was needed as sheep herders
are allowed to kill unlimited numbers of bears and have historically done so.

Capture and Marking
Capture efforts commenced on 29 May 1979. Bears were captured using Aldrich
spring-activated snares, set in baited cubbies, blind trail sets, and baited
trail sets. Snaring techniques were basically those described by Flowers
(1977). Primary bait items were carp (Cyprinus carpio), beaver (Castor
canadensis) and marmots (Marmota flaviventris).
Snares were set throughout
the study area but out of sight from roads and maintained trails.
All snare
sets were marked with signs on nearby trees warning people to stay clear of
the snare and a snared bear.
Snares were attached to green drag logs to

�-494-

reduce
needed

incidence of foot injury to captured bears and to reduce the time
to rebuild and reset the snare.
Snares were checked daily.

Snared bears were immobilized with a combination of ketamine hydrochloride
(Ketamine) and xylazine hydrochloride
(Rompun).
The ketamine hydrochloride
(100 mg/cc) was freeze-dried and concentrated to 200 mg/cc, then mixed
with xylazine hydrochloride
(100 mg/cc) in a 2:1 ratio.
A six-foot prod
pole was used to administer the drug.
Data on dosage used, time needed to
immobilize, and duration of immobilization were collected.
Immobilized
bears were tattooed, ear-tagged with Richey rubber ear tags (calf size), and
some we re instrumented with a Telonics radio transmitter collar.
Numerous
physical measurements were taken of each bear and a premolar was removed for
subsequent aging.
A completed bear capture data form is attached as Appendix
A. All of the measurements are recognizeable
from the form notation with
the possible exception of CAL and MzW.
These represent the canine alveolar
length and greatest width of second molar on the mandible as described by
Gordon and Morejohn (1975).
Lab estimates of age will be determined by
cementum annuli counts (Willey 1974).

Habitat

Selection

Detailed techniques for habitat type mapping and radio telemetry tracking
were reviewed.
However, insufficient effort on these phases of field work
was made to warrant description of methods.
The habitat type mapping techniques
will be improved in the next segment and will be described in detail in the
next segment report.

RESULTS
Study

Area

AND DISCUSSION

Selection

The B]ack Mesa - Crystal Creek area southeast of Crawford was selected as
the principal study area.
The study area is roughly bounded by Smith Fork
and Sink Creek on the north, Highway 92 on the west, the Gunnison River on
the south, and Curecanti Creek on the east.
The study area will be modified
to include all the seasonal ranges of radio-collared
females.
Such
modifications
will likely add areas west of Highway 92 on the north rim of
the Black Canyon and remove areas in the Virginia Creek and Smith Fork
drainages.
The final study area will likely be 80-90 square miles.
The
nearest town is Crawford, population 250, and fewer than 20 people live
within the principal study area.
Major drainages are Iron, Dyer, Crystal,
Mesa, and Curecanti Creeks.
Elevations range from 6,000 to 11,000 ft above
sea level.
Annual precipitation
averages 10.8 inches in Crawford (elevation
6,800 ft) and is considerably
higher in the upper reaches of the study area.
Average growing season in Crawford is 137 days and this would be similar to
the oakbrush dominated areas of the study area.
The study area has significant areas of six vegetation communities:
irrigated
grass meadows , sagebrush steppe, oakbrush chaparral
aspen, spruce-fir,
and dry mountain meadows.
These types are listed in ascending order of

�-495-

elevation but form a mosaic of communities with changes in aspect and
elevation. Primary human uses of the area are timber harvest, cattle
grazing, and big game hunting. Evidence of a long history of overgrazing
and current overgrazing abound. The livestock use in the area is all
cattle except for a sheep operation on private land in lower Curecanti
Creek. Most of the study area is administered by the Gunnison National
Forest. Cooperation of landowners has been generally excellent. However,
stlouldseasonal ranges extend west of Highway 92 it is probable that access
across private lands will be difficult to obtain.
The need for a hunting closure to protect adult female bears and minimize
adv~rse
contacts with hunters was.stressed by both John Beecham, biologist
for Idaho Fish and Game, and Ralph Flowers, Washington Forest Protection
AIHltlciation
Animal Damage Control Supervisor (pers. communication). This
requtrement for a study area soon became the most critical and was a
deciding factor in the final selection. The Black Mesa area (Game
Managemant Unit 63) haa historically supported a low licensed harvest
(~ 5 per year), had few if any nuisance bear problems, low numbers of sheep,
and nc outfitters dependent on the area for bear hunting. Also, unlicensed
hlit:rv(;l~twaa thought to be very low. However, unlicensed harvest has been
hiRh~r than earlier thought but still probably has not been high enough to
I3dvli'lndydhct
bear density. The Colorado Wildlife Commission decided on
January 1979 to support the closure of the study area to bear hunting. Their
intent 1~ to ~upport the closure for the study duration but regulations can
emly b~ cHltablt~hed for one year at a time. The area closed to bear hunting
in 1979 wa~ all of Game Management Unit 63, that part of GMU 54 west of
~G~p er~~k, and that part of GMU 53 south of Smith Fork and Sink Creek.
S1~n~ wHh th@ e1CH'1ure description and maps were posted in the area. The
11unt1nBclo~ur~ do~s not apply to unlicensed killing by livestock owners.

GapturM ~ucc~~~ WB~ paor in June 1979 with only six captures of four different
An ,!ldultboar, 310 lb was first caught on 29 May and later caught at
g cubby 0.4 m1l~~ aw~y an 29 June, then weighing 290 lb. This boar slipped the
Unt radio collAI'aHI!;!'f
five days and wore the second until his death on
12 ,July! H@ Wf.l~ ~hDt seueh of the Gunnison River so was out of the closed
~rIR, Th~ Initial capture of this bear convinced me not to use aspen drag
;!_f.JgEl.
Th~ b~aI'had Ch~WCH!down a standing 6" DBH aspen and both ends of
thg 7" diam~t~I' dr~B lag, reducing the drag from 9 ft to 2 ft in length.
6ubN@qu.nt captuI'~~with conifer drag logs have been satisfactory. AlSO lb
h~af WR§ cauBht twlc~, an 13 and 27 June. The captures were 2 miles apart.
Aft tlftuH hmal~ (l5~ Lb) was caught on 13 June and slipped her radio collar
th§ §gm~ dRY. A 28 Ih cub was caught 17 June. The sow was observed with the
ruh from a dl§tlnc~ of 200 yds but never appeared during the handling. Five
of th~ §ix c.ptUI'~~wlr~ at the same location. Snaring was restricted to
t'fl@ m;¥·bf!!§h And low{!lr~lil1~vl!ltion
aspen stands because of access. An
UfPj§tH~llyw~H;M&amp;y"'Jun~ parIcd resulted in lush understory throughout the
bf,\lIH'!3,

fl~~hfjH~h

IHliUHl.

�-496-

Initially a dosage of 2.5 mg/lb of Ketamine/Rompun
was used.
This rate
was not adequate, always requiring subsequent injections.
A dosage of
3.0 mg/lb appears to be suitable.
One problem with a light dose or
accidental underdosing
(by underestimating
weight) is that we have not
seen any progressive symptoms of drugging.
Whenever the dose was adequate
the bear slumped with no warning and when the dose was inadequate the bear
stayed alert and active.
Therefore, a dosage of 3.0 mg/lb will be used in
the future and try to overdose rather than underdose.
Overall the drug
works well.
Breathing rates stay high enough to keep bear well ventilated
(avg. 14.6/min for bears older than cubs).
Pulse rate averaged 72.8
beats/
min for the 5 bears older than cubs.
The cub had a pulse of 132 beats/min
and breathing rate of 30/min.
A potential problem with drug would be in
handling treed bears.
As there are no progressive symptoms it would be
difficult to rope the bear to the tree prior to complete immobilization and
having a bear fall from high in an aspen would likely contribute to high
trapping mortality rates.
Bears dosed at 2.5-3.0 mg/lb remained immobilized
at least 60 minutes and usually longer.
I believe it is easier on the bear
to recover without human present so I do not have data on total time of
immobilization.
Further analysis of this drug combination will be prepared
at the end of the 1979 field season.
Most of the 1979 field season has been, and will be, devoted to learning the
study area and capturing as many bears as possible.
Radio telemetry data
collection will increase as more bears are collared.

LITERATURE

CITED

Flowers, R.
1977.
The art and techn~que
Forest Protection Assoc., 37p.
Gordon, K. R., and V. Morejohn.
lower canine and lower molar
Willey,

C. H.

1974.

Aging

1.1

by:

black bears

(

,~:
. .JI~:vp"..!;

/1

bears.

Washington

1975.
Sexing black bear skulls using
measurements.
J. Wildl. Manage. 39(1):40,44.
from first pre-molar

J. Wildl. Manage. 38(1):97-100.

Prepared

of snaring

['

.

-O€. c A....

Thomas D. I. Beck
Wildlife Researcher

tooth sections.

�-497-

APPENDIX

A

�-498-

APPENDIX A

BEAR CAPTURE DATA FORM

--,-E_;:~=7--------~~~--~-----

BEAR CAPTURE DATA FORM

TAG NO.
TATTOO
TRANS. FREQ. 147. 'Itt 18
SEX
AGE'---

-¥

_

Date i?O J"U.L'i 1'n1 Capture
site216.Z X 'ig~2.3 Workers "BECK MJLI.£R.
Est.
Wt. If5S lb. Act. Wt.
IDS 1/'.
Drug Ke-tAho\;l1f! lR.o:"pv-n
Dosage:
3.0
mg/lb;
~cc.
Injection
site
Rl hip - TM
Method
:raJ, 'Po\e
Time of injection
~13~~=O Time of immob. __ ~I~~~~~~_ Time
left
bear
ILtSO
Time of recovery
Ul\know!')
Progressiv-e--s~y~m~p~t~o-m-s~=n:o-~~-~-~~b~~a~r
__ ~s~J~~~~p~.~d-Lh~O~M~~S~~~I~9-~rO~s~,~1~Dn~_
I

and
Convulsions:
Recurrence

INa;,

OlAf.
Time of

onset

~~~

Duration
----------Supp. Drugs
.lVONE
Air Temp. -"'4,~8~o.:...F-;__ -:-_
Rectal
Temp 1tX:&gt; () F
Pulse
~Ll /min
Respiration
/1
/min
Foot snared
LF Swollen
Chance for injury
6000
Est.
Age
S+
Lab est.
Sex
Color
Blond
Photo taken Yes
Ear
tag
F (0
Tattoo
G,
Trans.
Freq.
Itf7. Q'l1 g
Tooth removed -..::.....!-------P.
Collar
color
OrA~'3e
Pelage
c ond i.t Lon __ _::P....::o::..::o:..:.Y"
r+
Neck girth
£/1.5 CM
Chest girth.
6,8.0 0"
Wt. lOS
Total
length
IOO£m
Anus to anterior
of sternum
length
'87. 0 cm
CAL: 15.~ hi""
/5.8 •.••
,.., M2~.J: /0.3 ",,,,
10.0 •.••""
P4-Ci length
3/.3 m •••, 31.&amp;.j h"lm p4-Ci
length
d(D.O )of"'"
Z•O.", ••• :1in. width between C1 30.1mlYl Hin. width between
p4 3fo.'-It'II"t HI width
,
Hi width
,
Nursing
NO
Nipple
diam. L= TOP 4.1,.",.."
-1-.-"'-""-"'--' '7.iJ,,,,,,., R:1VP /).8,.,." ~.O,.,,,., , 8'.0 In""
Fore pad
width R lo.ScW!,
".2. c •.•.• Fore pad length
R SSe ••.•,
Hind pad width
tl.?c,.."
9.:l.c,."
Hind pad length
I~.::(t:,"
,
/3.Di:......
Vulva condo
non~~.J.rDtAS
Physiol.
I
Samples
Taken ~~~o~~~c~
1
Unusual
Phys.
Char. ~AJ~O~~~~~
_
Other bears
in area ~M~~~~~b
__~S~€~E~~~
_
Outside
influences
__ ~~~D~M~~~
~-r __ ~~
__ -.~
_
Comments Thumb hooL IH·bl:.e" PI?
nq~d .£9!-' ~~IJ/e
ckep

------------------------

yes

£

Ii.

a

'.0,,.,,

cu.I

r

»

I

�July,

-499-

JOB PROGRESS

Job Title

Mountain

8

Job No.

Lion Population

Dynamics

1

---------------------------------

--~------~----~--------~--------------------------------

Covered:

Personnel:

Big Game Investigations

W-126-R-2

No.

Work Plan No.

Period

REPORT

COLORADO

State of
Project

1979

July

Allen

1, 1978 - June 30, 1979

E. Anderson

ABSTRACT
A review of the published and unpublished literature on the mountain lion
was essentially completed with about 524 references read, catalogued and
filed under 26 subject headings.
A detailed study plan was not prepared
because I underestimated
the time required to fulfill objectives of Work
Plan 2, Job 5. A revised time table for the preparation of a detailed
study plan is recommended.

��-501-

MOUNTAIN

LION POPULATION

DYNAMICS

Allen E. Anderson

t

P. N. OBJECTIVE
Develop improved mountain lion inventory procedures and expand on knowledge
of mountain lion population dynamics and predator-prey relationships.
Specific
objectives are:
1.

Estimate density and population
populations.

size of selected mountain

2.

Develop an improved method
which is also economically

3.

Assess mortality rates of young mountain
maternal-filial
bond separation.

4.

Assess dispersal

5.

Estimate the impact of hunting and removal on mountain
dynamics.

6.

Measure the inter-relationships
large ungulate populations.

to reliably estimate mountain
feasible to apply.

and subsequent

lion

lion densities

lions after the period of

fate of sub-adult

between mountain

mountain

lions.

lion population

lion populations

and

SEGMENT OBJECTIVE
1.

Preparation of a detailed study plan describing the specific research
strategies which will be employed to meet the previously stated specific
objectives.
This includes the selection of specific study site(s).

METHODS AND MATERIALS
A comprehensive review of the published and unpublished
mountain lion was undertaken as the first step.

literature

on the

RESULTS AND DISCUSSION
The literature review was essentially completed.
About 524 references
were read, catalogued and cross referenced under 26 subject headings.
Because
of an underestimate of the time required to fulfill the objectives of Work
Plan 2, Job 5 (Experimental Deer Inventory), the literature review was the
only activity accomplished.

�-502-

Since the ultimate success of the study will depend largely on both the
suitability of the study areas selected and the field methodology adopted,
I now believe that the time table for the preparation of a study plan is
unrealistically short. The revised research time table should include
synthesis and publication of the literature; field work trips with researchers
currently involved in field investigations of mountain lions to review
potential methodologies; during selection of study area(s) and preparation
of a draft study plan during fiscal year 1979-80. Fiscal year 1980-81 will
involve testing methodologies and revising the study plan into final form.
Full-scale field work would commence during fiscal year 1981-82.

Prepared

by:

~5j~
Allen E. Anderson
Wildlife Researcher

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                  <text>-1October

JOB PROGRESS

State of
Project
Work

1979

REPORT

COWRADO

No.

Plan No.

Job No.

1

Job Title

Waterfowl

Period

April

Covered:

Migratorv

H-88-R-24

Production

Bird Investigations

1
------------------------------------

Surveys

23, 1978 to June 30, 1978

Personnel:
M. Nail and Staff, Monte Vista National Wildlife Refuge;
J. Creasy, and Staff, Brown's Park National Wildlife Refuge; F. N. Folks,
Utah State Division of Wildlife Resources; R. Clark, J. Corey, J. Ellenberger,
J. Frothingham, R. Keiss, J. Lorentzson, T. Martin, D. Hasden, J. Olterman,
G. Saville, R. Velarde, S. Steinert and Hike Szymczak, Division of Wildlife.
ABSTRACT
Water conditions for duck production were variable with poor conditions in
the San Luis Valley, average conditions in North Park, the Yampa Valley and
Brown's Park and excellent conditions in the Cache La Poudre and South Platte
valleys.
Conditions for goose production were considered excellent in the northwest and west central portion of the state but below average in north central
Colorado.
The total number of duck breeding pairs was estimated to be 72,526,
22 percent above the long term average.
The mallard (anas platyrhynchos)
was the major breeding species.
The post-nesting season population of Canada
geese (Bx-anta canadensis) in northwest Colorado was estimated to be 1710, 9
percent below 1977 level but 7 percent above the long term average.
In
north central Colorado, gosling production wa s down about 20 percent from
both 1977 and 1969-77 average levels.
Aerial counts on the Colorado, Gunnison
and White Rivers indicated an increase in the nesting population in all areas
except the Gunnison River.

��-3-

WATERFOHL

PRODUCTION

SURVEYS

Michael R. Szymczak
Steven F. Steinert

P.N. OBJECTIVES
1.

To estimate the number of duck breeding pairs,
~ajor waterfowl nesting areas in Colorado.

by species,

on selected

2.

To estimate the number of goose breeding pairs, and in some cases,
obtain production data on selected goose nesting areas in Colorado.

3.

Compile
federal

data and submit reports to appropriate state personnel and
agencies for use in establishing hunting season recommendations.

SEGMENT

OBJECTIVES

1.

To estimate the number of duck breeding pairs, by species, in the San
Luis, Cache la Poudre, South Platte and Yampa Valleys, and in North
Park and Brown's Park.

2.

To estimate the number of goose breeding pairs in the San Luis Valley
and west-central
Colorado and obtain goose breeding pair and production
data on the Yampa, Little Snake and Green Rivers in northwest Colorado
and in north central Colorado.

3.

Compile

data and submit

appropriate

METHODS

reports.

AND MATERIALS

Present duck breeding pairs and production surveys
pair inventory of only major production areas.

consist

of a breeding

The 1978 duck breeding pair surveys were conducted during the period of
May 8 to June 22.
Surveys in North Park and the Cache la Poudre and South
Platte valleys were conducted exclusively from the air.
Ground counts were
made in the Yampa Valley and Brown's Park.
In the San Luis Valley intensive
aerial counts were adjusted for visibility by air-ground comparison studies.
Pair estimates for the Monte Vista National Wildlife Refuge in the San Luis
Valley were obtained from nesting transects.
All survey methods and sample
areas for ducks remained the same as in previous years.
Surveys of Canada goose production were conducted within the period of April
25 to June 23. Production estimates for Moffat County were made from as
complete a count as possible of hatched or active nests and brood size.
Population estimates for north central Colorado were obtained from counts
of goslings and adults conducted from the ground during the period in which

�-4-

the birds were flightless.
Estimates of the Colorado, Gunnison and White
Rivers population were obtained from direct counts from a fixed-wing
aircraft.· A scheduled survey in the San Luis Valley of axpec t ed productive
Canada goose breeding pairs was not conducted because of inclement weather.
All flying was accomplished with a Cessna 185 aircraft.
Two observers
were used when sampling by transect while one observer was used when
sampling sections on flying river courses.
RESULTS AND DISCUSSION
In general, water conditions for duck production were much better in 1978
than in 1977. Conditions in the San Luis Valley continue to decline
because of a combination of dry weather and an increase in circular
sprinkling from wells as opposed to flood irrigation.
Water conditions
in North Park were below average, as a cool, wet spring retarded the
initiation of the irrigation season. However, conditions in the South
Platte and Cache la Poudre valleys were excellent with all ponds and
ditches full of water.
In the Yampa Valley a heavy snowpack resulted in
some temporary ponds retaining water into the nesting season.
But the
permanent ponds dependent upon irrigation water were low, because of delay
in irrigation.
Conditions for Canada goose production were excellent along the rivers in
northwest Colorado.
Normal water flows resulted in the average number of
usable islands throughout the nesting season. High water levels were
not experienced until after the peak of hatch.
In north central Colorado
reservoir water levels were higher than in 1977. However, a 26 inch
snowfall near the peak of hatch definitely reduced production from expected
estimates.
The estimated number of duck breeding pairs were very similar to 1976
levels, and 22 percent above thE! long-term average (Table 1). Comparisons
with 1977 numbers cannot be made as no estimates are available for most
areas in 1977.
The number of pairs in the San Luis Valley remained stable in spite of a
valley-wide decline in habitat conditions.
A decrease in the North Park
duck population was more than compensated for by large increases in the
South Platte and Cache la Poudre Valleys.
The mallard made up 29 percent
of breeding pair population; an increase over the 1976 level, but still
far below the long te~~ average (Table 2). The mallard was followed in
descending order of abundance by gadwall, green-winged teal and blue-winged
or cinnamon teal, each composing more than 10 percent of the population.
The post-nesting season population of Canada geese on trend areas in
northwestern Colorado was projected to be 1,710 birds in 1978 (Table 3),
down 9 percent from 1977 but 7 percent above the 1967-77 average (Table 4).
Estimated gosling production
(773) was 13 percent above the 1977 level and
40 percent above the 1967-77 average (Table 5).

�-5-

Table 1. Summary of Colorado's
in selected areas, 1978.

duck breeding

Total Estimated

pair population

Breeding

Pairs'

2/

1978

Area

1977])

Long-terrrrAverage

estimates

Percent
From

1977

Change

From Long
Term Average

+

San Luis Valley

28,524

27,855

North Park

9,633

17,U5

43.7

South Platte Valley

19,182

6,773

+ 183.2

Cache la Poudre Valley

12,674

3,662

+ 246.1

Yampa Valley

1,708

2,189

2,730

- 22.0

37.4

Brown's Park

805

1,009

1,141

- 20.2

29.4

Total

1/
2/

Estimates

72,526

not obtained

2.4

+ 22.4

59,276

for most areas in 1977.
are based on results of 1964 through
because of changes in survey
Figures for other"areas are 22 year

San Luis Valley and North Park averages

1977 and 1968 through 1977 surveys, respectively,
methods utilized
averages.

prior to those dates.

~/ Aerial counts corrected
San Luis Valley.

by species from visibility

ratios obtained

in the

Results of the 1978 production survey in north central Colorado are presented
by individual area in Table 6. Gosling production in north central Colorado
is down 20 percent from 1977 levels and 21 percent from the 1969-77 average
(Table 7). Production declined in all trend areas except Boulder and
Loveland.
The number of adults observed was up 20 percent from 1977 levels
but still down 6 percent from the 1969-77 average (Table 8).
The aerial survey of breeding Canada geese in west central Colorado was
flown on April 20, 1978. The results are presented in Table 9 and compared
with 1977 figures in Table 10. The 1977 survey was conducted on May 9.
In spite of the variation in survey dates between years, the presence of
more single geese during the earlier 1978 survey most likely indicates an
increase in the number of active breeding pairs.

�-6-

Table 2.

Species

composition

of Colorado's

1978 duck breeding

pair population.

,

Number of Breeding
Species

1978

1976-U

Pairs
1954-76
Average

Mallard

21,3l3

17,324

27,865

29.4

24.4

51.9

Blue-winged and
Cinnamon teal

9,267

7,432

5,761

12.8

10.5

10.7

Gadwall

14,311

4,205

5,283

19.7

5.9

9.8

Pintail

3,557

2,980

3,607

4.9

4.2

6.7

Shoveler

5,866

21,145

3,439

8.1

29.8

6.4

Green-winged
teal

12,739

9,136

2,743

17.6

12.9

5.1

543

3,623

2,314

0.7

5.1

4.3

978

2,750

1, l38

1.3

3.9

2.1

Other Divers

3,952

2,426

1,560

5.3

3.4

2.9

Total

72,526

71,021

53,71~1

Redhead
American

wigeon

llComparative

Percent
1978

SJ2ecies Com2osition
1954-76
Average
1976

data for 1977 not available.

. .
-2/S' peCles composltlon
regardless of changes

. d
compute d f rom data from a 11 areas for th e 22 year perlO
in survey method.

�-7-

Table
areas

3. Number of Canada geese observed
in Moffat County, Colorado, 1978.

and estimated

production

on trend

Nesting
Pairs

NonNesting
Adults

Total
Adults

Estimated
No.
1/
Goslings-

Total
Geese

22

105

149

65

214

23

93

139

106

245

Lily Park-~/

98

29

127

Subtotal

386

200

586

Area

Yampa River
Craig-Juniper

Springs

Juniper Springs Cross Mountain

Green River
Brown's

Park

73

96

242

285

527

Dinosaur National
Monument
(Colorado, Utah)

39

l30

208

198

406

Subtotal

112

226

450

483

933

20

61

101

90

191

937

773

1,710

Little

Snake River

Grand Total

1/ Calculated

using average

1/ Not surveyed.

Calculated

brood

size and number

totals are averages

of successful
of 1973-77

nests.

totals.

�-8Table 4. To al e timated
trend areas, 1978.

number

of Canada

geese obs rved, }foffat County

Percent

1977

Change
From
1967-77
Average

14.9

+

1.7

5.S

+

1.0

383

10.6

+

6.0

231

296

17.3

- 35.4

1,710

1,872

1,604

8.7

+

of Canada

goose goslings,

County

trend

1978

1977

1967-77
Average

586

689

576

Park

5'2.7

498

349

Dinosaur National
Monument
1/
(Colorado-Utah)-

406

454

191

Area

Yam a River

------

From

Green River
Brown's

Little

Snake River

Total

1/
- Not surveyed

until

Table 5. Estimated
areas, 1978.

+

6.6

1970.

numbers

Moffat

Percent

Change
From
1967-77
Average
1977

1978

1977

1967-77
Average

200

170

169

+ 17.6

+ 18.3

Park

285

210

135

+ 35.7

+111.1

Dinosaur National
Monument (Colorado,
Itah) _V

198

249

154

- 20.5

+ 28.6

90

54

93

+ 66.7

3.2

773

683

551

+ 13.2

+ 40.3

Area

Yampa

River

From

Green River
Brown's

Little

Snake River

Total

1/

- Not surveyed

until

1970.

�-9-

Table 6.

Production
Area

Results

of the north central

No.
Broods

Water Area

Wellington
Terry Lake
Water Supply and
Storage No. 4
Launer Pond
Douglas Reservoir
North Poudre No. 1
Dry Creek Reservoir
North Poudre No. 3
North Poudre No. S
Bureau of Standards
Divide No. 8 ]j
Elder Reservoir
Annex No. 8
Van Sant Pond
Cobb Lake
Watson Lake
Curtis Lake

l/
No. 1

0
0
4
1
0
1
12
6
0
1
3
1

Subtotal
Fort Collins
Peterson Ponds
Dixon (Herring) Lake
Miller (Maxwell) Pond
College Lake
Dean Acres
Claymore Lake
Sterling Gravel Pits
Lindenmeier Lake
Grey Lakes
Novak Reservoir
Anderson Pond's
Parkwood
Timnath Reservoir
Romily Gravel Pits
Fossil Creek Reservoir
Schuelke Reservoir
Wolaver Ponds

Subtotal

0
5
2
3
1
0
9
2
4

2
2

Colorado

goose census,

June 19, 1978.

Total No.
Goslings

Total No. Adults
and Yearlings

Total
Birds

35

44

79

18
8
0
0
16
1
0
3
35
18
0
5
23
9
4

26
5
2
14
10
2
2
25
134
12
20
17
117
144
69

44
13
2
14
26
3
2
28
169
30
20
22
140
153
73

175

643

818

0
16
9
8
14
2
43
0
41
8
8
15
61
21
11
9
9

9
19
21
199
18
64
202
81
45
4
31
39
77
10
21
6
4

9
35
30
207
32
66
245
81
86
12
39
54
138
32
32
15
13

276

850

1,126

�-10-

Table 6. Results
(Continued).

Production
Area

of the north central Colorado

Water Area

Loveland
Flatiron Reservoir
Boedecker Reservoir
Flatiron Gravel Pits
Kauffman Gravel Pits
Big Thompson River
McNeil Reservoir
Reservoir No. 12

goose census, June 19, 1978.

No.
Broods

Total No.
Goslings

Total No. Adults
and Yearlings

Total
Birds

1
5
6
2

2
24
24
6
12
51
14

2
85
12
4
8
43
12

4
109
36
10
20
94
26

133

166

299

4
11
8
22
0
57
15
8
6

28
19
46
24
32
29
184
8
9
4

32
30
81
32
54
29
241
23
17
10

166

383

549

27
0
25
5
8
8
0
7
0
11
0
35
0

60
31
175

87
31
200
75
52
160
36
31
15
52
l3
385
35

Subtotal

Boulder
Ish Lake
Crystal Lake
Terry Lake
Faivre Ponds
Rest Home Pond
Sawhill Ponds
Valmont Reservoir
Boulder Valley Farm
Eddy Pond
Angus Ranch Pond

351/
0

Subtotal

Denver
Ketring Lake
Centennial Pond
Bowles Lake
Kings Pond
Tule Lakes
Grant Reservoirs
Marston Reservoir
Pinehurst Country Club
Clairfield Reservoir
Kendirck Lake
Federal Center
Sloans Lake
Standley Lq_ke

0
6
2
2
3
2
0
3
0
0

70
44
152
36
24
15
41
13
350
35

-------------------------------------------------------------------------

�-11-

Table 6. Results of the north central Colorado
(Continued).

Production
Area

Hater Area

Denver (Continued)
Denver City Park
Colo. Blvd. and Quincy

No.
Broods

goose census,

Total No.
Goslings

June 19, 1978.

Total No. Adults
and Yearlings

Total
Birds

25
5

180
2

205
7

Subtotal

156

1,228

1,384

GRAND TOTAL

906

3,270

4,176

1/ Includes

2/

Includes

4
1

2 adults and 5 goslings

on pond 1 mile north of Dry Creek.

geese located on pond located !2 mile west of No.8.

1/ Goslings and adults not distinguished,
percent

therefore separation based on
on total birds which were goslings during the previous 5 years.

�-12-

Table 7. Number of Canada goose goslings
production trend areas, 1978.

Area

1978

produced

No. of Goslings
1969-1977
1977
Average

in north central

Percent
From
1977

Change
From
1969-1977

Wellington

175

215

256

-18.6

-31. 6

Fort Collins

276

288

285

- 4.2

- 3.2

Loveland

133

125

97

+ 6.4

+37.1

Boulder

166

142

216

+16.9

-23.1

Denver

156

357

292

-56.3

-46.6

Total

906

1,127

1,146

-19.6

-20.9

Table 8. Number of adult Cana.da geese observed
production trend areas, 1978.

Area

1978

No. of Goslings
1969-1977
1977
Average

in north

central

Percent
From
1977

Colorado

Colorado

Change
From
1969-1977

Hellington

643

615

734

+ 4.6

-12.4

Fort Collins

850

613

693

+38.7

+22.7

Loveland

166

182

224

- 8.8

-25.9

Boulder

383

394

594

- 2.8

-35.5

Denver

1,228

924

1,227

+32.9

0.0

Total

3,270

2,728

3,472

+19.9

- 5.8

�-l3-

Table 9.

West central

Colorado

Area

White

Canada goose breeding

Singles

Pairs

pair survey,

1978.

Croups

Goslings

River
6

1

o

o

Rio Blanco L. - Rangely

21

14

17

o

Rangely-Utah

4

11

o

o

31

26

17

o

o

2

9

o

26

31

15

4

DeBeque-Palisade

1

1

o

o

Palisade-5th

3

1

o

o

4

4

19

o

34

39

43

4

2

o

12

o

Hotchkiss-Delta

o

o

o

o

Delta-Grand

4

5

o

o

Subtotal

4

5

o

o

GRAND TOTAL

71

70

72

4

Blanco L.

Meeker-Rio

Line

Subtotal
Colorado

River

Glenwood

Springs-Silt

Silt-DeBeque

St. Bridge

5th St. Bridge-Utah

Line

Subtotal
Roaring

Fork River

El Jebel-Glenwood
Gunnison

Springs

River

Junction

�-14-

Table 10.
Comparison of the number of Canada goose singles and pairs observed
on aerial surveys in west central Colorado, Spring 1977 and 1978.
Number Observed
Singles

Pairs

Area

1978

1977

1978

1977

White River

31

15

26

32

Roaring

Fork River

2

2

o

6

Colorado

River

Glenwood Springs-5th
Street Bridge

30

13

35

33

Bridge-Utah

4

2

4

7

Hotchkiss-Delta

o

o

o

o

Delta-Grand

4

o

5

11

71

32

70

89

Gunnison

Line

River

Junction

Total

Prepared

by__-+~~~~~~
Michael
Wildlife

__~~~~~~~~~~_
Szymczak
Researcher C

Steven F. Steinert
Wildlife Technician

II

�-15October

JOB PROGRESS

State of

1979

REPORT

COLORADO
----------------~-----------

Work Plan No.
Job Title

2

Job No.

Studies of Canada Goose Populations

Period Covered:
Personnel:

Migratory

W-88-R-24

Project No.

Bird Investigations
6

--------------------------------in Colorado Transplant

Areas

April 1, 1978 to March 31, 1979

J. Corey, D. Flenthrope, R. Hopper, J. Lorentzson,
R. Velarde, and M. Szymczak

R. Oakleaf,

ABSTRACT
A total of 258 Canada geese (Branta canadensis) banded outside Colorado were
reported recovered within north central Colorado during the 1977-78 hunting
season. Analysis of those bandings did not conclusively indicate any new
production areas for Hi-Line Canada geese. Trapping operations in southern
Alberta, north central Colorado and the San Luis Valley of Colorado resulted
in 2,788, 603, and 139 geese being banded in the respective areas.

��-17STUDIES OF CANADA GOOSE
POPULATIONS IN COLORADO TRANSPLANT
Michael

AREAS

R. Szymczak

P. N. OBJECTIVES
1.

To document the breeding
central Colorado.

range of Canada geese wintering

2.

To document the distribution of harvest and estimate
of the resident foothills Canada goose populations.

3.

To document the distribution
San Luis Valley.

of harvest

in north

the survival

of Canada geese nesting

rate

in the

SEGMENT OBJECTIVES
lao

Plot the banding locations of Canada geese (Branta canadensis) banded
outside north central Colorado and reported recovered in north central
Colorado.

lb.

Participate in the cooperative
southern Alberta.

2a.

Trap and band 500 Canada geese on production
the foothills north of Denver.

and moulting

2b.

Complete,
cards.

schedules

3a.

Trap and band at least 200 Canada geese on production
areas in the San Luis Valley.

3b.

Complete,
cards.

Canada goose banding

submit and file appropriate

submit and file appropriate

banding

banding

operati.on in

areas along

and recovery

and moulting

schedules

and recovery

METHODS AND MATERIALS
Band recoveries listed on computer printouts provided periodically by the
Migratory Bird Populations Station were sorted and plotted as to banding
and recovery locations, sex, and age at time of banding, and estimated
breeding area for birds associated with north central Colorado.
Banding of flightless adults and goslings was accomplished through drive
trapping operations in southern Alberta, north central Colorado and the San
Luis Valley of Colorado.
The banding in Alberta was accomplished by
representatives of state, provincial and federal conservation agencies,
including Colorado.
Banding schedules and recovery reports pertaining to the San Luis Valley
and north central Colorado were completed and submitted to the Bird Banding
Laboratory.
Record keeping for the Alberta operation was the responsibility
of the Alberta Fish and Wildlife Division.

�-18-

RESULTS AND DISCUSSION
Hi-Line Population
Foreign Recoveries
During the 1977-78 hunting season, 258 Canada geese banded outside Colorado
were reported recovered in north central Colorado (Table 1), of which 128
were classified as wild trapped locals when banded (Table 2). The fourth
year of recovery information from banding in southern Alberta did very
little to alter the accepted distribution of the Hi-Line breeding range.
Locations banded for the first time in the sununer of 1977, Mary (93 banded),
Fincastle (236 banded), Cowoki (41 banded), Coleman (15 banded), Byemoor
(17 banded) and Erskine (12 banded) lakes resulted in only 5 recoveries in
north central Colorado, 3 from Fincastle and 1 each from Coleman and Erskine.
Mary Lake is well within the Rocky Mountain Population breeding range;
Fincastle is thought to be Rocky Mountain and or.ly 3 recoveries from the HiLine area will not change the designation; and Cowoki, Coleman, Byemoor and
Erskine are located along the hypothesized north-south boundary of the Rocky
Mountain Hi-Line populations from near Brooks to Stettler.
One recovery
each from Coleman and Erksine is not sufficient for population designation
and additional bandings are needed.
An abnormally high number of first
year recoveries from Grassy Lake (3% of the birds banded) occurred in north
central Colorado during 1977-78.
But geese from Grassy have been predominantly Rocky Mountain oriented in the past. Geese nesting at Schuler, Cavan,
Seven Persons and Murray lakes continue to be contributors to the population
wintering in north central Colorado, as do birds breeding in the EdmontonCamrose area.
Bandings in the Hi-Line area in southwest Saskatchewan were terminated in
1975, therefore the number of recoveries of banded birds from that area
has been going down steadily.
The only area in Saskatchewan contributing a
significant number of banded birds to the north central Colorado harvest in
1977-78 was the North Battleford area in east central Saskatchwan.
The
North Battleford geese were transported to the area and released as part of
a restoration effort.
Phillips County, a traditicnal Hi-Line breeding area in north central
Montana, was well represented in the harvest, as were Custer, Rosebud and
Treasure counties located along the Yellowstone River in the eastern portion
of the state.
Geese from restoration areas in northwestern Wyoming also
were well represented in the harvest.
Geese banded at the Wheatland Reservoir moulting area were again represented
in the harvest.
Small Canada geese captured in large moulting flocks in
the Anderson River Delta area were also well represented in the harvest for
the third consecutive year.
Banding
Banding was conducted for the fifth and final consecutive year on production
areas in southern Alberta and north central Colorado. A resume of the number
and location of geese banded in southern Alberta during the five year period
is presented in Table 3. In north central Colorado, a total of 603 geese

�-19-

was banded, bringing the five year total to 3,678 (Table 4). Geese were
captured at 13 locations with 50.1 percent of the birds banded being
goslings (Table 5).
San Luis Valley Population
A total of 139 Canada geese was trapped and banded on production areas in the
San Luis Valley (Table 6). The number of adults and goslings trapped is
presented in Table 7. A total of 405 geese has been trapped and banded over
the five year period.
An attempt to capture birds will be made again in 1979.

Table 1. Banding areas outside Colorado for Canada geese recovered
central Colorado during the 1977-78 hunting season.

Area

Alberta
Edmonton-Camrose-Vegreville
Kenilworth Lake
Hardisty Area
Erskine Lake
Dowling Lake
Coleman Lake
San Francisco Lakes
Louisiana Lakes
Bantry /12
Hays Reservoir
Grassy Lake
Fincastle Lake
Schuler Lake
Cavan Lake
Seven Persons Lake
Murray Lake

Direct

Area

4

Indirect

7
1

1
1

3
1

1
3
3
5
7
3
6

3

5
4

3
5

Saskatchewan
Manito Lake Area
North Battleford Area
Regina Area
Eyebrow Lake
Pelican Lake
Cypress Hills (Maple Creek)
Cypress Hills Area

2

7

1
1

1
1
1
2

British Columbia
Okanagan Valley
Mackenzie-Northwest
Territory
Beverly Lake-Thelon River
Anderson River Delta-Harrowby

in north

2

1
Bay

4

12

�-20-

Table 1. Banding areas outside Colorado for Canada geese recovered
central Colorado during the 1977-78 hunting season (continued).

Area

Central

in north

Direct

Indirect

9
1

33

1
4
6

1

Flyway

Montana
Phillips Co.
Sheridan Co.
Prairie Co.
Custer Co.
Rosebud Co.
Treasure Co.
Big Horn Co.

2

North Dakota
Stutsman Co.

4

5
5
1

1

South Dakota
Harding Co.
Perkins Co.
Hughes Co.
Wyoming
Big Horn Co.
Sheridan Co.
Campbell Co.
Crook Co.
Weston Co.
Goshen Co.
Albany Co.
Laramie Co.

2

4

o

2

11

4
1
3
8

1
3
5
1

1

Nebraska
Keith Co.
Kansas
Phillips

11:/

Co.

New Mexico
Mora Co.
San Miguel Co.
Socorro Co.
Chaves Co.
Texas
Randall Co.
Deaf Smith Co.

21/
y!1

21:/
21/
gll

11/
,11
.1.11

1-

�-21-

Pacific Flyway
Washington
Spokane Co.

1

Idaho
Gem Co.
Canyon Co.
Lake Co.
Bannock Co.
Utah
Rich Co.

2

California
Siskiyou Co.
Lassen Co.
San Diego Co.

1
1

Total

160

98

1./ Banded pos tseason.
1/ Banded

postseason,

taken 2 days later.

Table 2. Banding locations of Canada geese classified as wild trapped locals
which were recovered in north central Colorado during the 1977-78 hunting
season.

Year of Banding
Area

Alberta
Edmonton-CamroseVegreville Area
Erskine Lake
Dowling Lake
Coleman Lake
Louisiana Lakes
Bantry 112
Hays Reservoir
Grassy Lake
Fincastle Lake
Cavan Lake
Seven Persons Lake
Murray Lake
Schuler Lake

Before
1971

1971

1972

1973

1974

1975

1976

2

3

1

1977

1
1
1
2
2
2

1

1
1
2

1
1
4
2

1
2

2
1
1

7
3
6
3

�-22-

2.
Banding locations of Canada geese classified as wild trapped locals
Table
which we r e recovered in north central Colorado during the 1977-78 hunting
season.

Year of Banding
Before
1971

Area

Saskatchewan
Cypress Hills (Maple
Hills)
Cypress Hills Area
Val Marie-Frenchman
R. Area

1971

1972

1973

1974

1975

2

1
1

2

Flyway

Montana
Phillips Co.
Sheridan Co.
Prairie Co.
Custer Co.
Rosebud Co.
Treasure Co.
Big Horn Co.

3

1

3

2

3

4

11

1

4
2
2

1

1
4
6
1

1

1

\-Jyoming
Big Horn Co.

2

Flyway

Washington
Spokane

Co.

1

Idaho
Gem Co.
Lake Co.

1
1

Utah
Rich Co.

1

Total

8
I

1

North Dakota
Stutsman Co.

Pacific

1977

1

British Columbia
Okanagan Valley
Central

1976

5

2

8

3

14

20

32

44

�Table 3.

Location and number of Canada geese banded in southern Alberta 1974-78.

Numbers of Geese Banded
1977
1976
1975

Common Name

Location
Lat.-Long.

C.P.R. Res.

4950-11236

7 N Coaldale

90

84

139

1

101

Little Bow Res.

5012-11240

6 SW Travers

221

66

130

76

57

Milk R. Ridge Res.

4922-11227

6 S Raymond

327

118

227

5

77

Knight Ranch L.

4914-11227

20 S Stirling

6

111

73

106

334

Many Islands L.

5009-11004

28 NE Medicine Hat

5

69

Schuler L.

5020-11006

1 S Schuler

136

91

62

Cavan L.

4956-11024

14 SE Medicine Hat

79

74

91

40

83

Grassy L.

4948-11139

4 E Grassy Lake

107

224

273

206

70

Unnamed

5002-11141

18 E Vauxhall

91

32

35

45

12-Mi1e Coulee

5014-11139

6 E Rolling Hills

69

51

8

60

22

Bantry tI2

5022-11135

7 SE Tilley

325

82

191

67

77

Murray L.

4948-11057

5 SW Seven Persons

45

56

61

61

Seven Persons L.

4952-11054

1 S Seven Persons

60

84

70

Scope Res.

5005-11156

7 E Vauxhall

152

164

Unnamed

5047-11225

3 E Bassano

39

41

Barkenhouse L.

5054-11221

9 E Bassano

56

12

Nearest Town

1978

41
230

268

45

1974

--------------------------------------------------------------------------------------------------------------

I
N
W
I

�Table 3.

Location and number of Canada geese banded in southern Alberta 1974-78 (continued).

Common Name

Location
Lat.-Long.

Nearest Town

1978

Numbers of Geese Banded
1977
1976
1975

Keho L.

4957-11301

12 NW Picture Butte

202

78

110

101

Taber L.

4948-11206

2 E Taber

7

77

121

96

Lake Newell

5027-11158

9 S Brooks

St. Mary's Res.

4921-11311

12 NE Cards ton

115

Ross L.

4909-11255

16 E

Cards ton

184

174

Mary L.

4901-11313

13 SE Cardston

120

93

Fincastle L.

4950-11158

4 W Purple Springs

Coleman L.

5126-11151

13 S Hanna

llO

15

Cowoki L.

5035-11142

7 E Brooks

189

41

W. J. Reid L.

5052-11203

6 E Gem

154

Shooting L.

5211-11220

8 S Gadsby

98

Oldman River

4955-11152

7 N Purple Springs

16

Bow River

4958-11137

11 NW Burdett

3

1974

46
55

ll5

98
I

tv

.eI

Total

236

2,788

1,878

1,800

1,442

1,346

�-25Table 4. Number of Canada geese banded on production
central Colorado 1974-1978.

areas in north

Year

Adults

Goslings

1974

751

315

1,0671/

1975

406

195

601

1976

357

468

825

1977

280

302

582

1978

301

302

603

2,095

1,582

3,678

1/

Includes

one goose of unknown

Total

age.

Table 5. Location and number, by age and sex, of Canada geese banded
production areas in north central Colorado 1978.

Location

Adults
Males
Females

Males

Goslings
Females

on

Total

Rocky Ridge Reservoir

8

5

7

11

31

Grey Lakes

5

6

15

15

i.1

Romily Gravel Pits

6

8

15

14

43

Poudre Pre-Mix

9

7

8

2

26

Reservoir

No. 8

13

18

17

6

54

Timnath

Reservoir

2

3

17

24

46

Sterling

Gravel Pits

39

36

9

12

96

Boedecker

Reservoir

7

6

8

7

28

Watson Lake

0

0

3

2

5

McNeil Reservoir

14

18

11

19

62

Loveland

'+

5

9

12

30

Dixon Reservoir

0

0

9

4

l3

Valmont

31

51

23

23

128

l38

163

151

151

603

Total

Flat Iron

Reservoir

,

�-26-

Table 6. Numbers of Canada geese banded on production
in the San Luis Valley, 1974-1978.

and moulting

areas

Year

Adults

Goslings

Total

1974

4

6

6

1975

14

5

19

1976

55

149

204

1977

4

33

37

1978

18

121

139

Table 7. Location and number, by age and sex, of Canada geese banded on
production areas in north central Colorado.

Males

Location

Adults
Females

Males

Goslings
Females

Total

Monte Vista National
Hildlife Refuge

0

1

4

4

9

Russell Lakes

2

6

34

31

73

2

1

13

14

30

Rio Grande Management
Area

3

3

9

12

27

Total

7

11

60

61

139

Blanca Wildlife
ment Area

Prepared

by

Manage-

7Ji¥.til~
Wildlife

Researcher

�-27October

JOB PROGRESS

State of

1979

REPORT

COLORP~O
------------~--~----------W-88-R-24

Proj ect No.

Bird Investigations

2
9
Job No.
---------------------------------Monitor Banding of the Shortgrass
Prairie
~C~a~n~a~d~a~G~o~o~s~e~P~o~p~u~l~a~t~l~·o~n~~i~n~S~o~u~t~h~e~a~~~~~t~e~r~n~C~
_

Work PIau No.
Job Title
Period Covered:
Personnel:

Migratory

January 22, 1979 through March 31, 1979.

J. Corey, K. Wagner and M. Szymczak.

ABSTRACT

Post-season trapping in southeastern Colorado resulted in 353 banded Canada
geese (Branta canadensis), 340 of which were classified as members of the
short grass prairie populations.
The percentage of recoveries of southeast
in
Colorado banded geese reported taken during 1977-78 north of 530 latitude
Alberta reached 12.5 percent, the highest level in the last 15 years.
The
percentage of recoveries taken in southeast Colorado dropped below 20 percent
for the first time since banding began in 1951. Analysis of banding data
from 1958-65 and 1967-77 provided no evidence that survival rates of juveniles
and adults banded post-season in southeast Colorado were different.
Survival
of geese was estimated to be 74.44 + 1.61 from 1958-65 and 80.00 + 1.18 for
1967 through 1977.

��-29-

MONITOR BANDING OF THE SHORTGRASS PRAIRIE
CANADA GOOSE POPULATIONS IN SOUTHEASTERN COLORADO
Michael

R. Szymczak

P. N. OBJECTIVE
To continually document, through monitor banding and analysis of recovery
data the annual and long term status of the southeastern Colorado (Arkansas
Valley) segment of the short grass prairie Canada goose population.
SEGMENT OBJECTIVES
1.

Band a mlnlmum of 1,000 Canada geese in southeastern
post-season period.

Colorado

2.

Prepare and submit banding
and progress reports.

and return reports,

3.

Analyze band recovery data for geese banded in southeast Colorado to
determine (1) distribution of harvest, (2) recovery rate, and (3)
survival rate.

schedules,

band recovery

during

the

METHODS AND MATERIALS
All birds banded in southeast Colorado in 1978 were captured with baited
cannon nets, and the age and sex determined through cloacal and tail feather
examination.
All banding schedules, including recapture information, were
submitted to the U. S. Fish and Wildlife Service's Bird Banding Laboratory,
Patuxent, Maryland.
Information on the distribution of harvest and the scheme of recoveries by
year of harvest was obtained from computor printouts that are provided periodically by the Bird Banding Laboratory.
The banding and recovery matrices
were subjected to programs BROWNIE and ESTIMATE in order to obtain estimates
of recovery and survival rates (Brownie et al. 1978).
RESULTS AND DISCUSSION
Trapping

and Banding

Trapping efforts were hampered again in 1979 by the lack of geese roosting
at Two Buttes Reservoir and denial of access to Turk's Pond, their major
roosting area, for the majority of the trapping period.
Most of the birds
banded were captured at Turk's late in the banding period.
The number of
birds banded by location is presented in Table 1.
Distribution

of Harvest

The distribution of recoveries of southeast Colorado banded geese during the
1977-78 hunting season was quite different than the distribution recorded
for 1976-77 (Table 2). The percentage of recoveries north of 530 latitude

�-30-

in Alberta reached the highest level in the last 15 years, while the percentage of recoveries in Saskatchewan below 530 latitude also reached a record
high level. The take in southeast Colorado dropped below 20 percent for the
first time since banding began in 1951. A record high 9 percent of the
recoveries were reported taken in north central Colorado, probably indicating
increased use of that area by shortgrass prairie geese. The total number of
recoveries (88) was the lowest since the 1951-52 season, probably reflecting
reduced banding success in the past few years.

Table 1. The number and location
Colorado 1979.

Adults

Location

of Canada geese banded in southeastern

Males
Juveniles

Adults

Females
Juveniles

Unknown

Turk's Pond
Small geese

79

51

97

79

1

Large geese

3

3

3

4

o

8

6

9

10

o

90

60

109

93

1

John Martin
Small geese

Total

Recovery

and Survival

Rates

Banding and recovery summaries, presented in Tables 3 and 4, were analyzed
to obtain recovery and survival rate estimates.
Unfortunately, the data had
to be divided into two sets because banding was not done in 1966. Since two
age groups, adults (ASY) and juveniles (SY) , were identified during postseason banding, the data were first tested to see whether juveniles and adults
have similar recovery and survival rates. The tests involved model Ho' which
assumes recovery and survival rates to be independent of age, against Hl, which
permits those parameters to differ between juveniles and adults (Brownie et
ale 1978, p. 56-113).
Both models allow recovery and survival rates to vary
from year to year.
The results of the contingency chi-square tests of data sets in Tables 3 and
4 provided no evidence that recovery and survival rates for juveniles and
adults were different (Table 5). Additional analysis, using a Z-test statistic
also indicated there is no evidence that survival rates of the two age groups
are different.
The same test comparing
recovery rates provided no evidence
that recovery rates of the two age groups were different for birds banded in
1958-65, but some indication of a difference in juvenile and adult recovery
rates for geese banded in 1967-77.

�Table 2.

Percentages

of total band recoveries,

Area

1951-55

Far North Above

Five Year Averages
1956-60 1961-65
1966-70

post-season

bandings,

by area and year of recovery,
Recover}':Year
1973-74
1974-75

1971-75

1971-72

1972-73

0.5
2.4
0.5
3.3

2.8
4.2
1.4
8.5

0.4
5.3
1.3
7.1

0.9
7.9

17.6
16.9

16.0
14.7
1.3

19.1
17.3

2.2

1.8

all bandings.
Total No.
1977-78 Recoveries

1975-76

1976-77

0.9
8.6
0.9
10.4

1.2
4.3
0.6
6.1

1.1
12.5
1.1
14.7

16.2
21.6
0.9

17.8
19.6
0.6

12.5
28.4

% of Total
Recoveries

-

-

-

-

-

9.8

7.1

10.2

9.1

1.0
5.0
0.8
6.8

28.6
18.3
0.1

35.3
10.5

28.9
11.5

19.0
21.8
0.8

15.6
18.1
0.6

11.9
21. 8
0.5

0.8
0.1
0.7
0.1
5.4

0.4

0.3

1.3
0.1
1.2
0.5
5.0

1.9
1.0
1.4

6.5

0.5
0.3
5.8

0.7
0.3
0.5
0.3
6.3

5.2

-

-

-

-

25.5
0.4
0.9
1.5

30.9
0.3
0.3
0.7

28.4
0.5
0.4
0.4

31.4
0.2
0.4
0.4

34.3
5.1
39.4
0.9
0.1
0.6

44.6
1.9
46.5
1.0
0.5
0.5

24.6
5.6
30.3

2.3
2.0
0.8
5.1

2.8
2.2
0.2
5.2

4.6
1.5
0.5
6.6

5.5
1.2
0.8
7.5

5.6
1.0
0.6
7.2

2.4
1.0
3.4

8.5
0.7
2.8
12.0

2.4

2.2

0.8

0.7

0.9

1.4

1.4

0.4

0.3

0.1

-

-

8.8

346

8.7

979
670
10

24.7
16.9
0.3

28
5
26
13
234

0.7
0.1
0.7
0.3
5.9

1,243
18
31
39

31. 4
0.5
0.8
1.0

168
56
21
250

cl.4
0.5
6.3

59

1.5

5

0.1

1

0.03

Below 530

Alberta
Saskatchewan
B.C., Manitoba,

Ontario

Central Flyway
Montana
North Dakota
Wyoming
South Dakota
Nebraska
Colorado
Southeast and Other
North Central
Total
Kansas
Oklahoma
New Mexico
Texas
Panhandle
Waggoner Ranch
Gulf Coast
Total
Pacific

Valley

530

N. W. Territories
Alberta
Saskatchewan
Total
Provinces

Arkansas

Flyway

MississiEEi
Mexico
Total Number
of Recoveries

Fll!vay

0.6

-

0.1

0.6
0.7
10.6

1.4
0.7

2.7
0.4
2.2

0.9
0.9
5.1

37.8
5.8
43.6
1.8
0.9

27.4
7.3
34.7

5.3
1.3

7.8
0.9

6.7

8.7

1.8

2.3
8.0

0.9
1.9

8.6

26.8
6.4
33.2
0.9
3.7
2.7

27.6
7.4
34.4
1.8
0.6
1.8

18.2
9.1
27.3

6.4
0.9

5.5
1.2

2.3

7.3

6.7

2.3

2.3
2.3

0.9
0.9

0.6

0.1

748

677

770

712

799

211

142

225

li3

108

163

88

3,957

4.-'_

I

w

•....
I

�Table 3.

Number of Canada geese banded in 1958-65 and resulting recoveries 1958-77.

Year
Banded

Number
Banded

1958

1959

1960

1961

1962

1958

347

33

27

17

8

1959

217

o

21

9

7

1960

250

o

16

6

12

306

o

29

13

18

1962

334

o

o

26

18

1963

369

o
o
o

o
o
o

27

1961

335

780

o

o
o

o
o

1965

o
o

o
o
o

o

1964

o
o

1958

176

21

16

6

o
o
o
o

25

o
o
o
o
o
o

Number and Year Recovered
1966
1967
1968
1969

1963

1964

1965

5

8

2

2

2

3

1

4

4

2

1

1

o

1

o
o

8

4

6

3

2

1

12

4

7

3

2

3

12

5

13

3

2

35

21

14

4

11

24

12

15

o

o
o

o

55

4

4

1

4

16

16

8

7

3

16

10

3

14

o

15

5

o
o
o
o

o

10

o
o
o

o
o
o

1970

1971

1972

1973

1974

1975

1976

1977

1

2

o
o

o

1

1

o

1

1

o
o
o

o

o

o
o

o

o
2

1

3

2

1

1

o
o

o
o
o
o

2

4

1

1

2

o

1

1

1

5

2

5

2

o

2

1

1

o

o

10

4

4

11

3

1

8

1

2

2

o

40

20

22

12

13

9

6

7

3

2

5

o

o

3

1

o

o

o

o

o
o

1

1

2

1

1

o

1

o

o

1

o
o
o

o
o

o

1

o

4

7

8

7

3

2

1

2

2

1

o
o
o
o

1

8

o
o
o

10

o

1

1

1

1

1

o

o

1_

1

o
o

o

12

o

o
o
o
o
o

40

24

9

14

6

2

2

5

2

1

3

1

o

1

1

o
o

16

11

11

13

5

2

8

5

o

1

o

1

o

15

7

8

1

o

3

1

1

3

1

o
o

o
o

Adults

Juveniles

1959

298

1960

167

1961

248

1962

123

1963

399

1964

251

o
o

1965

198

o

o
o

1

o

I
W
N
I

�-33-

Table 4. Number of Canada geese banded in southeast Colorado and resulting
recoveries 1967-77.

'Year
Banded

Number
Banded

1967

1968

1969

Number and Year Recovered
1970 1971 1972 1973 1974

1967

389

29

26

10

9

8

6

6

3

3

1

2

1968

646

35

17

29

25

10

10

2

6

3

3

1969

905

39

42

15

28

12

8

8

6

358

21

6

6

4

2

7

3

532

o

23

16

21

15

7

10

1

1972

694

o

34

18

10

17

10

499

14

16

10

6

205

o

6

8

8

3

1975

357

6

7

237

o
o

17

1977

o
o
o

6

405

o
o
o

19

1976

o
o
o
o
o

30

1974

o
o
o
o
o

o
o
o
o
o
o

25

1973

o
o
o
o
o
o
o
o

21

1971

o
o
o
o
o
o
o
o
o

22

1970

o
o
o
o
o
o
o
o
o
o

o

6

1967

285

18

10

5

12

9

8

5

3

2

o

1

1968

270

o

16

9

8

4

6

3

2

2

3

o

1969

184

o

9

11

11

3

5

3

2

o

1

1970

241

o

18

9

4

l3

o

o

2

2

1971

301

o
o

10

13

5

6

4

1

337

o

17

17

5

8

8

4

1973

65

o
o

5

2

3

1

2

132

o
o

o

1974

8

2

5

2

1975

71

4

1

514

11

114

o

o
o

21

1977

o
o
o

3

1976

o
o

o
o
o
o
o
o
o

19

1972

o
o
o

o
o
o
o
o
o
o
o
o

o

6

1975

,1976 1977

Adults

Juveniles

o
o

o

o
o

o
o

o

o

o
o
o

o

o

o

o

�-34-

Table 5. Comparison of juvenile and adult recovery and survival
for Canada geese banded post-season in eastern Colorado.

Parameter

or Test

Banded 1967-77
Adult
Difference

Subadult

rate estimates

Z Value

P

Mean recovery

rate

5.75 + 0.57

4.69 + 0.24

-1.06

-1. 71

0.09

Mean survival

rate

84.68 + 8.73

80.08 + 3.22

-4.60

-0.49

0.64

Z Value

P

Test of Ho vs HI

X2

12.20, df

21, P

0.93

Goodness

of fit test of Ho

X2

85.23, df

95, P

0.75

Parameter

or Test

Banded 1958-65
Adult
Difference

Subadult

Mean recovery

rate

8.51 + 0.69

8.43 + 0.40

- .08

-0.10

0.92

Mean survival

rate

73.50 + 4.63

74.08 + 2.07

.38

0.12

0.90

Test of Ho vs HI

X2 =20.72,

Goodness

X2 =154.44, df

of fit test of Ho

df

16, P = 0.19
l30, P = 0.07.

Adult and juvenile bandings and recoveries were then pooled (Tables 6 and 7)
and tested for model fitness using models developed for bandings of adult
birds only (Brownie et al. 1978, p. 13-55). Analysis and testing of 1958-65
bandings indicates Modell,
which assumes time-specific survival and recovery
rates is the appropriate model for the data set. A test of Modell
vs.
Model 0 (assumes time-specific survival and recovery rates where first year
recovery rates are different from recovery rates of previously-banded
cohorts)
indicated rejection
of Modell
(Table 8). However, considering the geese
are banded post-season, there is no reason to expect the variation in re~overy
rates as assumed under Model O. In addition,
the rejection of Modell
is
not conclusive considering the large X2 and z values are mainly attri.buted
to 1965 (Table 8). The goodness of fit test indicates Modell
is acceptable
(X2 = 81.24, df = 81, P = 0.47). Recovery and survival rates under Modell
for 1958-65 are presented in Table 9.

�Table 6.

Number of Canada geese banded in southeast Colorado, 1958-65 and resulting recoveries 1958-77.

Year
Banded

Number
Banded

1958

1959

1960

1961

1962

1963

1964

1965

1958

523

54

43

23

12

9

9

6

2

5

4

1

1959

515

0

46

25

23

12

11

5

1

2

1

1960

417

0

0

43

26

9

26

16

4

6

1961

554

0

0

0

44

18

22

19

12

14

Number and Year Recoverd
1966
1967
1968
1969

1970

1971

1972

1973

1974

1975

1976

1977

0

1

2

0

0

0

1

0

0

3

1

1

0

0

0

0

0

1

0

4

2

1

2

0

1

1

0

2

0

0

6

4

4

4

3

4

2

2

1

0

0

I

w
Ul

1962

457

0

0

0

0

36

30

22

5

14

4

3

3

5

1

1

3

1

1

1

1

1963

768

0

0

0

0

0

75

45

23

18

17

7

4

10

4

1

5

2

1

1

1

1964

586

0

0

0

0

0

0

40

23

26

23

9

6

19

8

1

9

1

2

3

0

1965

978

0

0

0

0

0

0

0

70

47

28

23

12

16

10

7

10

4

2

5

0

I

�-36Table 7. Number of Canada geese banded in southeast Colorado
recoveries 1967-77.

Year

Number
Banded

1967

1968

1969

Number and Year Recovered
1970 1971 1972 1973 1~74

1967

674

47

36

15

21

17

14

11

1968

922

0

51

26

37

29

16

1969

1,089

0

0

31

50

53

1970

599

0

0

0

39

1971

836

0

0

0

1972

1,032

°

0

1973

564

0

1974

337

1975

and resulting

1975

1976

1977

6

5

1

3

13

4

8

6

3

18

33

15

10

8

7

30

10

19

4

2

9

5

0

42

26

34

20

13

14

2

0

0

0

42

51

24

18

25

lLf

0

0

0

0

0

35

16

19

11

8

0

0

0

0

0

0

0

14

10

13

5

428

0

0

0

0

0

0

0

0

22

10

7

1976

920

0

0

0

0

0

0

0

0

38

18

1977

353

0

0

0

0

0

0

0

0

°
°

0

12

Table 8. Chi-square and Z values obtained from a test of Model 1 vs Model 0,
that first year recovery rates and/or survival rates are the same as those
from cohorts banded in previous years.
Values
Cohort

X2

Z

1959

.07

- .26

1960

.00

.05

1961

.04

- .20

1962

4.77

2.18

1963

.85

.92

1964

3.84

-1. 96

1965

9.96

3.16

Total

19.52

3.88

2
X

19.52, df

7, P

0.007

Z

3.88, df

7, P

0.07

�-37-

Table 9. Recovery and survival rates of Canada geese banded
Colorado 1958-65 and 1967-77.

Recovery

(± SE)

Rate

in southeast

Survival Rate

(± SE)

95% c. I.

95% c. I.

Estimate

10.33 + 1.33

7.72 - 12.93

87.5Lf+ 9.67

68.59 - 106.48

1959

9.13 + 1.04

7.10 - 11.15

47.91 + 5.34

37.44 -

1960

10.27 + 1.14

8.03 - 12.50

83.34 + 8.53

66.62 - 100.07

1961

8.10 + 0.86

6.41 -

9.79

71.47 + 7.49

56.79 -

86.16

1962

6.05 + 0.74

4.61 -

7.49

80.89 + 7.88

65.45 -

96.33

1963

9.13 + 0.78

7.61 - 10.65

64.31 + 5.92

52.71 -

75.91

1964

8.45 + 0.79

6.90 - 10.01

85.63 + 7.76

70.43 - 100.84

1965

5.53 + 0.52

4.51 -

Year

Estimate

1958

Arithmetic

Mean

95% C. I.

6.54
74.44 +

8.06 - 9.50

71.29 - 77.60

7.10 + 0.98

5.17 -

9~O3

1968

5.94 + 0.62

4.72 -

7.16\

1969

3.19 + 0.37

2.42 -

3.93

1970

6.09 + 0.58

5.09 -

7.08

1971

6.16 + 0.49

5.21 -

7.11

1972

3.89 + 0.36

3.19 -

4.59

1973

'6.22 + 0.48

5.27 -

7.16

1974

3.61 + 0.38

2.86 -

4.36

1975

3.95 + 0.42

3.12 -

4.77

1976

4.36 + 0.41

3.55 -

5.17/

1977

2.97 + 0.35

2.28 -

3.66

95% C. I.

Mean

1.61

8.78 + 0.37

1967

Arithmetic

58.39

80.00 + 1.18

77.69 -

82.32

4.86 + 0.19
4.49 - 5.24

------

�-38-

Analysis of the 1967-77 data set indicates Model 2, which assumes recovery
rates to be year specific, but survival to be constant, is the appropriate
model.
Examination of the "Goodness of Fit" tests indicates both Hodel 1
(X2 = 42.41, df = 45, P = 0.58) and Model 2 (X2 = 46.76, df ,= 54, P = 0.75)
fit the data. A test of Model 2 vs Model 1 indicate Model 2 cannot be
rejected in favor of Modell
(X2 = 5.89, df = 9, P = 0.75).
Since Model 2
has fewer parameters (12) than Modell
(21), substantially better precision
of the estimators is achieved under Hodel 2. Recovery and survival rate
information for 1967-77 is given in Table 9.
Examination of both recovery and survival rates indicate the recovery rates
were higher and survival rates lower during 1958-65 compared to 1967-77.
Unfortunately, since the two sets of data best fit different models, testing
of differences of the most precise estimates are not possible.
However,
since Modell
was also considered a suitable fit for the 1967-77 data, tests
of differences were conducted using estimates derived
under Model 1 for both
data sets (1958-65, S = 74.44
1.61, T = 8.78
0.37; 1967-77, S = 78.80
2.09, f = 5.00 ± 0.20). A 2-test of the difference in survival as estimated
under Modell
indicated there was no evidence,of different survival rates
between the two groups (2 = 0.78, P = 0.44). There was, however, a significant difference in recovery rates (2 = 9.75, P&lt; .0001).

±

±

±

Generally there are indications of declining recovery rates and increasing
survival rates over the 19 year period.
Population inventory figures indicate
an increasing population from 1958 to 1972 (Table 10). Since 1972, the
population size according to counts has been very erratic.
Accurate estimates
of the total population have been difficult because of the change in distribution with more shortgrass birds wintering in north central Colorado.
Age
ratio information as an index of production indicate a high percentage of
immatures in the population in only 2 out of the last 6 years.
An increase
or decrease in the percent of immatures in the population has predictably
not been paral1e11ed by population numbers, primarily because of the nonproductive sub-adult component of the population.
Under the current rate of
survival, the population should continue to increase given average production.

LITERATURE

CITED

Brownie, C., D. R. Anderson, K. P. Burnham, and D. S. Robson.
1978.
Statistical inference from band recovery data - A hand book. U. S. Fish
and Wild1. Serv., Resour. Pub1. 131. 212 p.

Prepared

by

77'1DIA:L)

e.:

;J

.~~

Michael R. Szym~~
Wildlife Researcher

C

�-39-

Table 10. Results
geese 1958-79.

of the January

inventory

of short grass prairie

Canada

Year

No. of Geese

Year

No. of Geese

1958

60,300

1969

112,399

1959

76,010

1970

147,414

1960

77,709

1971

145,349

1961

103,355

1972

147,300

1962

80,133

1973

256,710

1963

93,940

1974

145,780

1964

81,221

1975

103,674

1965

103,435

1976

216,753

1966

110,485

1977

192,942

1967

106,652

1978

106,062

1968

127,903

1979

140,191

1/ Unreliable

estimate.

1/

��-41October

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Work Plan No.
Job Title
Period

Bird Investigations

3

7
Job No.
------------~-------------------Population Characteristics
of
Mallards Wintering in West Central Colorado

------------------------------~~-------------------------------------

Covered:

Personnel:

Migratory

W-88-R-24

1979

April 1, 1978 to March 31, 1979

G. Bock, J. Corey, D. Coven, J. Ellenberger, J. Frothingham, J.
Gray, J. Gumber, J. Leslie, J. Hiller, D. Owerrs , S. Steinert,
M. Stone, and R. Hopper.

ABSTRACT
The sixth consecutive year of the study of wintering mallards in west central
Colorado was completed in Segment 24. A wintering population of 12,691 birds
was estimated for,1979, based on the aerial survey conducted in January.
This
figure was less than the January 1978 estimate (13,860) and the long-term
average (13,493 for 1959-1978).
Ground counts of 3,360 birds produced a sex
ratio of 110 males:l00 females, which was lower than the 1978 ratio (114.5:
100). Nearly 1,800 mallards were banded during the postseason period in 1979,
bringing the total banded sample to 10,141 for the six years of study (19741979).
About 350 recoveries have been obtained from these bandings through
August 1, 1978, which represents only five years of recovery information.
The
Pacific Flyway portion of Colorado accounted for the majority of the recoveries
ranging from 58 to 71 percent of the total, depending upon age and sex class.
Most of the remaining recoveries occurred in Canada and Idaho.

��-43-

POPULATION CHARACTERISTICS OF
MALLARDS WINTERING IN WEST CENTRAL COLORADO
Richard M. Hopper

This report presents results of the sixth consecutive year of study of the
mallard population wintering in west central Colorado.
Included is a sumnlary
of all data collected since initiation of this investigation during the 197374 segment year.
P. N. OBJECTIVES
1.

To estimate population parameters of mallards wintering in west central
Colorado by age and sex category; specifically (1) population size, (2)
recovery rates, (3) survival rates, (4) mean life span, (5) sex ratio,
and (6) geographic distribution of the harvest.

2.

To develop a management plan for the population
in west central Colorado.

of mallards

winterin8

SEGMENT OBJECTIVES
1.

Conduct an aerial count of wintering numbers of mallards during the first
or second week of January as part of the annual mid-winter survey conducted throughout the United States.

2.

Conduct a minimum of two ground counts (sex ratio) of 500 mallards each
in each of the following two concentration areas: (1) Grand JunctionHighline Lake area, and (2) Montrose-Delta area.

3.

Trap and band 1,500 mallards during the postseason period, including 750
in the Grand Junction-Highline Lake area, and 750 in the Montrose-Delta
area. Distribute the sample equally among the four age and sex classes.

4.

Preparation and submission of banding
reports, and progress reports.

schedules,

band recovery

and return

METHODS AND MATERIALS
All methods remained the same as in Segment 22 (Hopper 1977). The aerial
inventory was conducted on January 3, 1979 and involved flying over all kno~vn
waterfowl concentration areas in the Uncompahgre-Gunnison-Colorado
River
Complex of west central Colorado.
Personnel of the Northwest Region again
made the survey, utilizing a Cessna 185 aircraft and a crew consisting of a
pilot and two observers.
Sex ratio counts were made during the postseason banding period on the dates
of January 26, 29, 31 and February 4 and 6 at Highline Lake and on January 25
and February 5 at Burlingame Pond near Delta.
Salt Plains type traps, baited with corn, were again used to capture mallards
for banding after the close of the hunting season.
Specifically, banding was

done during the period January 25 and February

7, 1979.

All birds were aged

�-44-

according to the wing-aging
maintained for each banding
were prepared and submitted

technique (Carney 1964), and recapture data were
location.
Banding schedules and recovery reports
to the Bird Banding Laboratory.

RESULTS AND DISCUSSION
Aerial

Census

Results of the 1979 January inventory of wintering mallards, in addition to
counts from the previous five years, are presented in Table 1. The 1979
estimate of 12,691 was about 1,200 birds short of the number observed in 1978
(13,860) and about 2,200 less than the average for the period 1974-1978
(14,842).
The long-term (1959-1978) average for this population is 13,493.
The area covered in this count was presented in more detail in a previous
report for this job (Hopper 1977).
Table l. January
1974-1979.

Location

inventory

of mallards

wintering

in west central Colorado

1974

1975

1976

1977

1978

1979

High1ine

Lake

2,400

4,700

10,200

6,900

6,100

4,200

Colorado

River

3,110

1,830

1,545

3,390

3,046

2,856

Gunnison

River

700

1,345

9,100

2,215

3,756

2,850

0

485

270

225

208

1,785

0

0

1,900

1,800

450

0

2,975

3,400

110

1,750

300

1,000

9,185

11,760

. 23,125

16,280

13,860

12,691

Uncompahgre
Sweitzer

River

Lake

Burlingame

Total
5-Year Mean

Pond

(1974-1978)

= 14,842

Sex Ratio Counts
Table 2 lists the sex ratio counts taken in 1979 by date and location.
Overall
ratios obtained during each of the previous three years are also shown for
comparison.
Over 3,350 mallards were counted in 1979, resulting in a sex
ratio of 110.1 ma1es:lOO females for all areas combined.
Northern and southern
portions of the study area were not too different, yielding 110.8 and 107.8
males per 100 females, respectively.
Prior to 1979, the proportion of males
in the counts steadily increased each year since 1976, but this may have been
due to sampling error rather than a reflection of a real change in the sex ratio.

�-45-

Table 2. Sex ratio counts of mallards in west central Colorado during
January and February 1979, with overall ratios from 1976 and 1977 for
comparison.

No.
Males

No.
Females

Total

01-26-79

278

264

542

105.3

01-29-79

278

254

532

109.4

01-31-79

284

260

544

109.2

02-04-79

272

228

500

ll9.3

02-06-79

264

236

500

lll.9

1,376

1,242

2,618

llO.8

01-25-79

276

246

522

ll2.2

02-05-79

109

III

220

98.2

385

357

742

107.8

Total

1,761

1,599

3,360

llO.l

1976

1,741

1,755

3,496

99.2

1977

1,375

1,279

2,654

107.5

1978

2,816

2,460

5,276

ll4.5

Date

Location

Highline

Lake

Subtotal
Burlingame

Subtotal

(North)
Pond

(South)

Trapping

Males:lOO
Females

and Banding

Trapping and banding was continued during the 1979 postseason period (late
January and February), representing the sixth consecutive year of banding
effort.
Nearly 1,800 mallards were banded in the two general areas in 1979,
resulting in a total banded sample of 10,141 for the six years (1974-1979)
of study (Table 3). The age composition of the trapped sample favored the
subadult class in 1979, much the same as it has in past years.
The adult
female segment continued to be the least represented of the four age and
sex classes, although in the earlier years it was as common as the other
three.
Preliminary
The number

of recoveries

Distribution
obtained

of Band Recoveries

from the first five years of banding

as

�-46-

Table 4.
Colorado,

Recovery

Recovery distribution of mallards
1974-1978 (all years).

Location

AM

banded postseason

in west central

Percent of Total Recoveries
Age and Sex
SM
AF

SF

Canada
Alberta
British Columbia
Saskatchewan

5.4
0.0
3.2

6.0
0.0
3.0

8.2
0.0
2.0

9.9
1.4
5.6

Subtotal

8.6

9.0

10.2

16.9

Arizona
Colorado
Idaho
Montana
New Mexico
Oregon
Utah
Washington
Wyoming

0.0
58.1
8.6
2.1
2.1
0.0
5.4
0.0
4.3

1.5
60.4
3.7
2.3
0.0
0.7
6.7
0.0
2.3

0.0
71. 5
2.0
0.0
0.0
2.0
6.2
2.0
2.0

0.0
57.8
5.6
0.0
0.0
0.0
9.9
0.0
4.2

Subtotal

80.6

77.6

85.7

77.5

Colorado
Kansas
Montana
Nebraska
New Mexico
Texas
Wyoming

7.5
0.0
1.1
0.0
0.0
1.1
0.0

5.3
1.5
0.0
3.0
0.7
O. 7
1.5

0.0
0.0
2.0
0.0
0.0
0.0
0.0

2.8
1.4
1.4
0.0
0.0
0.0
0.0

Subtotal

9.7

12.7

2.0

5.6

Iowa
Louisiana

0.0
1.1

0.7
0.0

0.0
2.0

0.0
0.0

Subtotal

1.1

0.7

2.0

0.0

Total

100.0

100.0

99.9

100.0

Number of Recoveries

93

134

49

71

Pacific

Flyway

Central E'Lyway

Mississippi

Flyway

�-47-

of August 1, 1978 totaled 347. These recoveries represent only those birds
that had been shot and/or found dead during the hunting season (September 1January 31).
Table 3. Number and age and sex composition of mallards banded postseason
in west central Colorado, 1979, and totals for the period 1974-1979.

AM

Age and Sex
SM
AF

SF

Total

209

328

86

329

952

296

242

134

165

837

505

570

220

494

1,789

1,513

1,377

937

1,390

5,217

1,256

1,569

790

1,309

4,924

2,769

2,946

1,727

2,699

10,141

Year and Location

1979
Grand Junction

Area

Delta Area

Total
1974-1979
Grand Junction

Area

Delta Area

Total

The geographic distribution of these 347 recoveries is shown in Table 4 by
age and sex class.
The Pacific Flyway portion of Colorado was the major
recovery location for all four age and sex classes, accounting for from
58.1 to 71.5 percent of the total recoveries.
Adult females showed a somewhat higher affinity for this area than did the other three classes.
Canada
and Idaho were the next most common recovery locations.
Literature

Cited

Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification
by means of wing plumage.
U. S. Fish and Wildl. Serv., Spec. Sci.
Rept., Wildl. No. 82. 47 pp.
Hopper, R. M. 1977. Population characteristics
of mallards wintering in
west central Colorado.
Colo. Div. of Wildl. Fed. Aid Game Res. Repto,
Oct. pp. 33-39.

Prepared

by

~
kL. L"}/J'"'Hopp;r;;;~
.·~t&gt;t:.~ ,

/

, (
~/

.

'/~ c ..{

Rf.char d M.

Wildlife

Researcher

'=

C

��-49October

JOB PROGRESS

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

Work Plan No.
Job Title

W-88-R-24
3

Monitor Banding

Period Covered:

1979

Migratory
Job No.

Bird Investigations

8
---------------------------------

of Eastern Colorado Wintering

Mallard Populations

April 1, 1978 to March 31, 1979

Personnel:
M. Babler, M. Bauman, G. Brown, L. Budde, J. Carsella, J. Corey,
C. Crawford, M. DePra, K. Dillinger, M. Gardner, B. Goforth, K. Green, D.
Homan, T. Howard" J. Jackson, B. Leasure. R. Lopez, J. Lorentzson, S. Lucero,
T. Lynch, F. Marcoux, K. Moser, R. Oehlkers, C. Pabst, J. Pogorelz, F. Rinella,
C. Roberts, W. 'Rupke, H. Sp ear , S. Steinert, M. Szymczak, K. Wagner, R.
Zaccagnini, and R. Hopper.
ABSTRACT
The banded sample for Segment 24 totaled 4,606 mallards, or about 600 birds
over the minimum quota established for eastern Colorado.
An updated analysis
of banding data was initiated during the segment, based on 76,197 bandings
between 1963-1964 and 1975-1976, and 6,475 recoveries from the 1964-1965
through 1976-1977 hunting seasons.
Several analyses involving recovery
distributions and recovery and survival rate estimates are being conducted
by management unit of banding and age and sex class.

��-51MONITOR BANDING OF EASTERN
COLORADO WINTERING MALLARD POPULATIONS
Richard M. Hopper

This report summarizes Segment 24 results of the program involving monitor
banding of mallard populations in eastern Colorado during the post season
period.
Hopper (1977) described the purpose of the original study, general
accomplishments, reasons for transferring field responsibilities
to the
regional management system, and the role of the Research Section in the
program.
Hopper (1978) presented results of the first phase of this investigation, which related to the first P. N. Objective listed below.
These
results concluded that the transfer of monitor banding responsibilities
from
research to management was now complete, and that the first objective of
this study was thereby met. The 1978 report also outlined the responsibilities of the Research Section with regard to this banding program.
Thus,
the Segment 24 report presented here addresses the second P. N. Objective.
P. N. OBJECTIVES
1.

To establish monitor banding of wintering mallard
Colorado as an annual management function.

populations

in eastern

2.

To continually document, through monitor banding and analysis of recovery
data, the annual and long term status of eastern Colorado wintering
mallards to provide a basis for annual hunting season recommendations.
SEGMENT OBJECTIVES

1.

Band a mlnlmum of 4,000 mallards during the post season period, including
a minimum of 500 birds in each of the following general areas of the
South Platte Valley and Arkansas Valley: (a) Denver-Greeley,
(b) Fort
Collins-Loveland-Windsor,
(c) Greeley-Fort Morgan, Cd) Fort MorganSterling, (e) Sterling-Julesburg,
(f) Bonny Reservoir, (g) ManzanolaLamar, and (h) Two Buttes Reservoir area. Divide the banded
sample in
each area equally among the four age and sex classes.

2.

Conduct an updated analysis of band recovery data, including the following
major determinations for important population segments of mallards
wintering in eastern Colorado: (a) distribution of the harvest, (b)
recovery rates, and (c) survival rates.

3.

Prepare and submit banding
and progress report.

schedules,

band recovery

and return reports,

METHODS AND MATERIALS
Procedures and equipment remained essentially the same as in the previous year
(Hopper 1977), with the exception that participation in the actual banding
effort by research personnel (Federal Aid Project W-88-R) was limited to
Bonny Reservoir.
The Research Section has retained responsibility for banding
at Bonny Reservoir because of clo~ely related studies anticipated for the
future.

�WATERFOWL MANAGEMENT UNITS

K

N E

A

WYOMING

I

"&gt;:J

OKLAHOMA

k'
NEW

GFP-R-M-l

Fig. L

Waterfowl Management

MEXICO

Units in colorado.

I."

'~.

�-53-

RESULTS AND DISCUSSION
Banding
Over 4,600 mallards were banded during January and February of 1979, or
Segment 24 (Table 1). This number again exceeded the 4,000 quota established
for eastern Colorado.
Only one location, the Arkansas Valley, failed to
contribute a representative sample to the banding program.
All banding
schedules and band recovery reports were prepared and submitted to the Bird
Banding Laboratory.

Table 1. Mallards banded post season by age and sex in the eight eastern
Colorado banding areas, January-February
1979.

Banding Area

AM

Number of Ducks Banded
Age and Sex
SM
AF
SF

Bonny Reservoir

400

400

248

231

1,279

Sterling-Julesburg

114

103

112

119

448

Fort Morgan-Sterling

125

119

116

136

496

Greeley-Fort

141

126

124

117

508

176

182

135-

206

699

Fort Collins-Loveland-Windsor

226

129

124

126

605

Arkansas

Valley

11

4

4

3

22

Two Buttes Area

251

175

30

93

549

1,444

1,238

893

1,031

Morgan

Denver-Greeley

Total

I

Total

Updated Analysis

of Band Recovery

4,606

Data

Banding and recovery tapes were received from the Bird Banding Laboratory in
Segment 24 for mallards banded post season in eastern Colorado during the
period 1963-1964 through 1975-1976 and for recoveries from these bandings
during the 1964-1965 through 1976-1977 hunting seasons.
These tapes were
placed on file at the Colorado State University Computer Center, and the
Computer Center User Services was hired to run the desired programs.
The banding tape was subjected to a program that tallied numbers of mallards
banded by: (1) management unit (Fig. 1), (2) latitude and longitude (10minute block), (3) year, and (4) age and sex class.
Table 2 presents a

�-54-

summary of the resulting printout by management unit and age and sex class,
while Table 3 lists numbers banded by year and age and sex class.
Over 76,000
birds were banded during the l3-year
period of study (44,453 males and 31,744
females).
Total banded samples were consistent among management units at
about 10,000-12,000 each, except for units 10-13 (Table 2).' These latter
four units will be combined for purposes of the analysis because of the
poorer individual samples obtained.
Yearly samples varied from 3,382 to
7,153 (Table 3).
Table 2. Numbers of mallards banded post season in eastern Colorado
ment unit and age and sex class (1963-64 through 1975-76).

by manage-

Management
Unit

Adult

Male
Subadult

Total

Adult

Female
Subadult

Total

Grand
Total

1

3,161

2,923

6,084

1,688

2,246

3,934

10,018

2

3,571

2,912

6,483

2,177

2,135

4,312

10,795

3

3,052

3,013

6,065

2,008

2,528

4,536

10,601

4

3,224

2,667

5,89l

2,247

2,282

4,529

10,420

6

3,910

3,130

7,040

2,253

2,509

4,762

11,802

9

3,248

2,733

5,981

2,114

2,458

4,572

10,553

10

451

490

941

205

408

613

1,554

11

765

477

1,242

832

786

1,618

2,860

12

2,167

1,338

3,505

1,110

1,022

2,132

5,637

13

817

404

1,221

387

349

736

1,957

Totals

24,366

20,087

44,453

15,021

16,723

31,744

76,197

The recovery tape was also subjected to several programs involving recovery
distributions and recovery and survival rate estimates for 6,475 recoveries.
Printouts of recovery distributions included numbers and percent of total
birds recovered for each management unit of banding and age and sex class.
Recovery locations were tabulated by latitude and longitude (degree block),
flyway, state, and Colorado management unit.
Separate printouts were also
obtained for first-year and all-year recoveries.
Although these programs
involving recovery distribution were run during the segment, the resulting
printouts have not been reviewed and interpreted.
This work will be accomplished during Segment 25, along with generating some additional printouts
of recovery distribution by 10-day intervals of the hunting season and by
year class of birds.

�-55-

Table 3. Numbers of mallards
and age and sex class.

banded post season in eastern Colorado

Banding
Year

Adult

Male
Subadu1t

Total

Adult

1963-64

2,177

1,244

3,421

1964-65

3,135

1,237

1965-66

2,293

1966-67

by year

Female
Subadu1t

Total

Grand
Total

484

684

1,168

if,589

4,372

726

634

1,360

5,732

2,164

4,457

1,066

1,548

2,614

7,071

2,090

2,069

4,159

1,136

1,722

2,858

7,017

1967-68

1,914

1,844

3,758

1,405

1,990

3,395

7,153

1968-69

1,210

1,219

2,429

1,145

872

2,017

4,446

1969-70

910

791

1,701

645

1,036

1,681

3,382

1970-71

1,466

1,441

2,907

1,221

1,207

2,428

5,335

1971-72

1,823

1,566

3,389

1,448

1,709

3,157

6,546

1972-73

1,862

1,523

3,385

1,677

989

2,666

6,051

1973-74

1,568

1,371

2,939

1,393

1,278

2,671

5,610

1974-75

1,868

1,812

3,680

1,350

1,562

2,912

6,592

1975-76

2,050

1,806

3,856

1,325

1,492

2,817

6,673

Totals

24,366

20,087

44,453

15,021

16,723

31,744

76,197

Re cove ry and survival rate analyses were initiated based on the new methods
described by Brownie et al. (1978). Specifically, a portion of the recovery
data was analyzed through programs BROWNIE and ESTIMATE during the segment,
and the remaining data will be processed by these same two programs in Segment
25. These analyses include recovery and survival rate estimates by management unit of banding and by age and sex class.
The printouts thus far
generated from these programs were still in the process of review at the end
of Segment 24, thus no results can be presented here. Hopefully, the complete
analyses will be finished during next segment and a report prepared shortly
thereafter.

�-56-

LITERATURE

CITED

Brownie, C., D. R. Anderson, K. P. Burnham, and D. S. Robso~.
1978.
Statistical inference from band recovery data - a handbook.
U. S. Dept.
of the Int., Fish and Wildl. Servo Resour. Publ. No. 131. 212 pp.
Hopper, R. M. 1977. Monitor banding of eastern Colorado wintering mallard
populations.
Colo. Div. of Wildl. Fed. Aid Game Res. Rept., Oct.
pp. 41-47.
1978.
lations.

Prepared

by

Monitor banding of eastern Colorado wintering mallard popuColo. Div. of Wildl. Fed. Aid Game Res. Rept., Oct. pp. 41-45.

;t:J- -iaP/:; - )/D#2/
Richard M. Hopper
Wildlife Researcher

i'

C

C

�-57-

JOB PROGRESS

October

REPORT

State of

COLORADO
--------~~~~~-----------

Project

No.

1~-88-R-24

Job Title
Period

Job No.

9

_

Migration and Hortality Characteristics
of
Duck Populations in the Inter-Mountain Valleys of Colorado

Covered:

Personnel:

Higrat_ory Bird Investigations

3

Work Plan No.

1979

April 1, 1978 to Harch 31, 1979

H. Nail and Staff, Honte Vista National Wildlife Refuge; R.
Darnell and Staff, Alamosa National Wildlife Refuge; J. Corey,
Flenthrope, Dean Flenthrope, R. Oakleaf~ J. Wagner, K. Wagner,
S. Steinert, L. Stevens, and M. Szymczak, Colorado Divsion of
Wildlife.

Dale

ABSTRAGr
Totals of 2,875, 3,316 and 4,912 ducks were banded in North Park, South Park
and the San Luis Valley, respectively, in 1978. The major species of harvest
during the early season in North Park in 1978 were the gadwall (20%), northern
shoveler (18%), mallard (11%), blue-winged or cinnamon teal (10%), and lesser
scaup (10%). In South Park the mallard made up 44 percent of the harvest
during the early season followed by the pintail (15%), American wigeon (12%),
and green-winged teal (12%). In the San Luis Valley the mallard was the major
species of harvest (40%) during the early season, followed by the gadwall
(24%), American wigeon (11%), and northern shoveler (11%).

��-59HIGRATION AND MORTALITY CHARACTERISTICS
OF
DUCK POPULATIONS IN THE INTER-MOUNTAIN VALLEYS OF COLORADO
Michael

R. Szymczak

P. N. OBJECTIVES
1.

To investigate migration, mortality, recovery distribution and relationships among populations of selected species of ducks present in North Park,
South Park and the San Luis Valley during the mid-July through midSeptember period.

2.

To document the species composition of ducks harvested during early October
seasons, should seasons occur in North Park, South Park and the San Luis
Valley.

3.

To examine the feasibility of establishing early special duck seasons in
September in the high mountain park areas or to examine the feasibility
of special early October seasons in South and North Park similar to what
has been recommended for the San Luis Valley.

4.

To establish procedures to monitor the effect of early special duck
seasons or, duck populations in the high mountain park areas if granted.
SEGMENT OBJECTIVES

1.

Trap and band ducks in North Park, South Park, and the San Luis Valley
during the mid-July through mid-September period as designated in Program
Narrative Outline.
Complete, submit and file appropriate banding schedules
and recovery cards.

2.

Collect and analyze data concerning the species composition of the harvest
during early October seasons in North Park, South Park and the San Luis
Valley utilizing wing collection barrels and results of the U. S. Fish
and Wildlife Service's Parts Collection Survey.

3.

Analyze band recovery data through the 1976 recovery year for mallards,
pintail and green-winged teal banded during the pre-season period in North
Park, South Park and the San Luis Valley to determine the feasibility of
requesting early seasons in the high mountain area.

4.

Prepare

5.

If a special high mountain duck season is requested and granted, design
a program for population monitoring which may include banding, harvest
surveys, wing collection or other activities.

progress

report.

METHODS

AND MATERIALS

Ducks were captured from August 9-September 13 in South Park, August 12September 15 in the San Luis Valley and July 26-September 22 in North Park.
Primarily "salt plains" type bait traps were used (Szymczak and Corey 1976).
In North Park, night lighting from airboats was used to capture most gadwall
and wigeon.
The age and sex of all birds captured and banded were determined.

�-60-

Wings from ducks bagged during the September 30, 1978 through October 13, 1978
period were collected in North and South Parks through the use of voluntary
collection barrels (Hoffman and Braun 1975). Barrels were placed at Walden
Reservoir (2 barrels), Lake John Annex (1 barrel) and at Cowdrey (1 barrel)
in North Park, and at Antero Reservoir (3 barrels) in South'Park.
Some
additional wings were collected at Delaney Butte Lake and on Case Flats.
All wings collected were classified by species, age, sex and location, and
in some cases, periods
of harvest.
The species composition of the harvest
in the San Luis Valley was obtained at the Central Flyway, U. S. Fish and
Wildlife Service's Parts Collection "wing bee."
Computerized analysis of banding and recovery tapes of birds marked through
1975 in North Park, South Park, the San Luis Valley and the high country
areas west of the San Luis Valley continued using programs outlined by
Szymczak (1978). In addition, most results for mallard, pintail and greenwinged teal were updated through the 1977-78 recovery year utilizing periodic
printouts of recovery information received from the Bird Banding Laboratory.
Information concerning all aspects of the duck populations in the banding
areas including breeding pairs, production, pre-season and winter population
size, hunting season and harvest was compiled
and written into draft form
to be included in the final report.
RESULTS AND DISCUSSION
Banding
Totals of 2,875, 3,316 and 4,912 ducks were banded in North Park, South Park
and the San Luis Valley, respectively, in 1978. The species composition of
the birds banded in each area are presented in Tables 1, 2 and 3. Quotas of
300 mallards of each sex of adult and immature birds were essentially met for
both ages of males in all areas, but for females only in the San Luis Valley.
In the three areas, fewer total pintails were banded in 1978 than in 1977,
however, the age ratios were more equally balanced.
The gadwall quota in
North Park of 300 adults of each sex was not met because of mechanical
problems with the airboat.
The total number of birds banded was 11,103
(Table 4).
Species Composition

of the Harvest

A total of 597 wings was collected during the 1978 early duck hunting season
in North Park.
Seventy-two percent of the wings were collected at Walden
Reservoir where the major species of harvest was the gadwall, followed by the
shoveler (Table 5). The mallard made up a substantial portion of the harvest
at both Lake John Annex and the Cowdrey area. The mallard comprised over 11
percent of the total harvest compared to 6 to 7 percent during the 1975-77
period in North Park (Table 6). The percent of the harvest composed of wigeon
has been declining since 1975, while the shoveler harvest has generally been
increasing.
Sixty-six percent of the wings in the sample were from birds
taken the first two days of the season (Table 7).
The mallard made up 43.5 percent of the ducks harvested during the early season
at Antero Reservoir (Table 6). The pintail was second in importance in the
harvest followed by the green-winged teal and wigeon.
Ninety percent of the
wings were from birds taken the first 2 days of the season (Table 8).

�-61-

Table l. Number of ducks banded, by species,
pre-season period, 1978.

in North Park during

Age and Sex
LM
AF

AM

1M

Mallard

315

347

4

Pintail

317

268

Gadwall

303

the

IF

LF

Total

175

242

2

1,085

12

224

214

12

1,047

26

5

100

12

5

451

28

7

4

15

12

2

68

102

39

0

25

21

°

187

Blue-winged and
cinnamon teal

2

3

2

4

2

2

15

Redhead

4

1

6

6

1

4

22

1,071

691

33

549

504

27

2,875

in South Park during

the pre-

American

wigeon

Green-winged

teal

Total

Table 2. Number of ducks banded by species,
season period, 1978.

Age and Sex
LM
AF

AM

n1

Mallard

301

296

6

Pintail

328

356

Redhead

5

American

wigeon

Gadwall
Green-winged

teal

Blue-winged and
cinnamon teal

Total

IF

LF

Total

132

259

5

999

2

l30

321

4

1,141

1

0

16

0

22

4

1

0

1

°

0

0

6

0

1

0

1

i

0

3

330

263

0

97

154

0

844

39

145

0

17

100

0

301

1,007

1,063

8

394

835

9

3,316

�-62Table 3. Number of ducks banded by species in the San Luis Valley during the
pre-season period, 1978. 1/

AM

1M

Age and Sex
LM
AF

IF

LF

Total

28

506

589

19

2,246

Mallard

540

Pintail

178

363

o

130

291

1

963

Green-winged teal

277

131

3

115

87

o

613

Blue-winged and
Cinnamon teal

110

391

5

83

288

2

879

Redhead

5

15

5

5

15

10

55

Shoveler

o

o

1

o

o

o

1

Gadwall

o

25

34

13

27

32

131

American wigeon

2

11

2

5

1

3

24

1,112

1,500

78

857

1,298

67

4,912

Total

1/ Includes 1,261 and 763 ducks banded by the Alamosa and Monte Vista National
Wildlife Refuges, respectively.

Table 4. Number of ducks banded, by species, in North Park, South Park and the
San Luis Valley during the pre-season period 1978. 1/
Age and Sex
LM
AF

IF

LF

Total

38

813

1,090

26

4,330

987

14

484

826

17

3,151

709

433

3

237

262

o

1,644

Gadwall

303

52

39

114

40

37

585

Blue-winged and
Cinnamon teal

151

539

7

104

390

4

1,195

American wigeon

34

19

6

21

13

5

98

Redhead

14

17

11

27

16

14

99

Shoveler

o

o

1

o

o

o

1

3,190

3,254

119

1,800

2,637

103

11,103

AM

1M

Mallard

1,156

1,207

Pintail

823

Green-winged teal

Total

1/
and 763 ducks banded by the Alamosa and Monte Vista National
- Includes 1,261
Wildlife Refuges, respectively.

�-63-

Table 5. Species composition of the harvest in North Park during
September 30-0ctober 13, 1978 early duck hunting season according
voluntarily placed in collection barrels.

the
to wings

Area

Species

\.Jalden
Reservoir

Lake John
Annex

Cowdrey
Area

Delaney
ButtesCase Flats

Total

Gadwall

96(22.3)

11(13.9)

8(11.0)

4(28.6)

119(20.0)

87(20.2)

9(11.4)

11(15.1)

3(21.4)

110(18.4)

Mallard

30(8.6)

17(21.5)

18(24.7)

3(21.4)

68(11.4)

Blue-winged or
Cinnamon teal

45(10.4)

9(11.4)

8(11.0)

0(0.0)

62(10.4)

Lesser

52(12.1)

2(2.5)

8(11.0)

0(0.0)

62(10.4)

37(8.6)

11(13.9)

5(6.8)

1(7.1)

54(9.0)

28(6.5)

11(13.9)

2(2.7)

0(0.0)

41(6.9)

22(5.1)

3(3.8)

4(5.5)

2(14.3)

31(5.2)

17(3.9)

4 (5.1)

8 (11.0)

1(7.1)

30(5.0)

10(2.3)

1(1.3)

0(0.0)

0(0.0)

11(1.8)

Ruddy

6(1.4)

1(1.3)

1 (1.4)

0(0.0)

8(1.3)

Bufflehead

1(0.2)

0(0.0)

0(0.0)

0(0.0)

1(0.2)

79

73

14

Northern

Shoveler

scaup

American

wigeon

Redhead
Green-winged

teal

Pintail
Ring-necked

Total

duck

431

597

Forty percent of the 45 wings collected during the early season in the San
Luis Valley were from mallards, while gadwall comprised 24 percent (Table 6).
The percent of the harvest composed of mallards during early season has been
generally declining since at least 1963 (Table 9).
Band Recovery

Analysis

Analysis of the banding data through the 1977-78 recovery year 1S continuing
at this time. Results of the analysis will be presented in the final
report.

�Table 6. Percent species composition of the harvest in North Park and South Park during early OctolJer duck
hunting seasons 1975-1978 according to independent wing barrel surveys and in the San Luis Valley based on U. S.
Fish and Wildlife Service's Parts Collection Survey.

__

.

Species

NP

1975
SP 1/

SLV

NP

1976
SP

S1V

NP

1977
SP

S1V

NP

1978
SP

SLV

Mallard

6.9

-

47.1

6.1

8.7

44.9

6.0

55.9

43.7

11. 4

43.5

40.0

Gadwall

20.6

-

20.6

28.1

2.2

6.7

25.4

8.8

12. 7

20.0

6.5

24.4

7.7

-

4.4

7.2

25.0

19.1

4.9

4.9

12.7

c;

')

,_

12.0

4.1f

2.9

21.1j

28.3

4.5

14.9

4.4

7. a

9.0

12.0

11. J

5.9

8.8

25.0

13.5

8.6

2.9

1.4

10.4

7.6

8.9

H.8

4.8

3.3

7.9

5.6

8.8

16.9

5.0

15.2

0.0

4.4

2.7

2.2

2.2

11. 2

0.0

4.2

18.4

1.1

11.1

1.5

11. 2

5.4

0.0

10.1

2.9

0.0

6.9

0.0

0.0

Green-winged
American

teal

wigeon

28.9

Blue-winged/
Cinnamon teal

15.4

Pintail

5.1

-

J.

Redhead

4.3

-

Ruddy duck

1.6

-

0.0

1.6

0.0

0.0

2.6

1.5

0.0

1.3

0.0

0.0

0.2

-

0.0

1.1

0.0

1.1

0.7

0.0

1.4

1.8

0.0

0.0

0.0

7.0

0.0

0.0

9.7

0.0

0.0

10.4

0.0

0.0

0.0

0.0

1.1

0.0

0.4

1.5

0.0

0.0

2.2

0.0

1.5

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

92

89

68

71

92

45

Northern

shoveler

Ring-necked

3.4

duck

Scaup

5.1

Merganser

0.0

Hooded merganser

0.0

Bufflehead

0.0

-

494

1/ Wings not collected

I
0'&gt;

Total No.
Wings Examined

.

68

374

in South Park in 1975.

268

597

.1-....
I

�-65-

Table 7.
Species composition of the harvest by time period at Walden
Reservoir, Lake John Annex, and the Cowdrey area during the September
October 13, 1978 duck season.

30-

Species

Sept. 30-0ct. 1
No.
Percent

Oct. 2-8
No.
Percent

Oct. 9-13
No.
Percent

.
~
2/
E'ntlre
~easonNo.
Percent

Gadwall

72

18.3

27

19.7

3

11.5

119

20.0

87

22.1

11

8.0

5

19.2

110

18.4

Mallard

35

8.9

21

15.3

9

34.6

68

11. 4

Lesser scaup

37

9.4

24

17.5

1.

3.8

62

10.4

Blue-winged or
Cinnamon teal

51

12.9

9

6.6

o

0.0

62

10.4

American wigeon

31

7.9

17

12.4

2

7.7

54

9.0

Redhead

32

8.1

8

5.8

o

0.0

41

6.9

24

6.1

4

2.9

2

7.7

31

5.2

14

3.6

8

5.8

4

15.4

30

5.0

4

1.0

6

4.4

o

0.0

11

1.8

Ruddy

6

1.5

2

1.5

o

0.0

8

1.3

Bufflehead

1

0.3

o

0.0

o

0.0

1

0.2

Northern

shoveler

Green-winged

teal

Pintail
Ring-necked

Total

duck

137

1/ Includes

26

597

14 birds taken either at Delaney
the old Fish Hatchery.

Buttes Lakes or on Case Flats near

1/ Includes

time period.

40 birds taken during an unknown

�-66Table 8. Species composition of the harvest by time period at Antero
Re servo Lr , South Park, Colorado during the September 30-0ctober 13, 1978 duck
hunting season.

Time Period
Oct. 2-13
No.
Percent

Entire Season
No.
Percent

Species

Se2t. 30-0ct. 1
No.
Percent

'1allard

40

48.2

0

0.0

40

43.5

Pintail

14

16.9

0

0.0

14

15.2

9

10.8

2

22.2

11

12.0

Cr eerr-w i.nge d teal

6

7.2

5

55.6

11

12.0

Blue-winged

5

6.0

2

22.2

7

7.6

Gadwall

6

7.2

0

0.0

6

6.5

Common merganser

2

2.4

0

0.0

2

2.2

Shoveler

1

1.2

0

0.0

1

1.1

Total

83

American

wigeon

teal

Table 9. Percent of mallards
San Luis Valley 1963-78. 1/

92

9

in the early October

duck season harvest

in the

Year

Percent

Year

Percent

1963

86.9

1971

65.6

1964

73.8

1972

50.7

1965

77 .8

1973

60.6

1966

73.7

1974

47.7

1967

71.6

1975

47.1

1968

75.8

1976

44.9

1969

67.5

1977

43. 7

1970

54.9

1978

40.0

1/ 1963 through

1970 from Hopper

et al. 1975.

�-67-

LITERATURE

CITED

Hoffman, R. W., and C. E. Braun.
1975. A volunteer
Colo. Div. Wildl. Game Inform. Leafl. No. 101.

wing collection
3 p.

station.

Hopper, R. M., A. D. Geis, J. R. Grieb, and L. Nelson, Jr. 1975. Experimental duck seasons, San Luis Valley, Colorado 1963-1970.
Wildl.
Monogr. No. 46. 68 p.
Szymczak, M. R. 1978. Migration and mortality characteristics
of duck
populations in the Inter-mountain valleys of Colorado.
Colo. Div. of
Wildl., Game Res. Rept., Fed. Aid Proj. W-88-R.
October.
p. 47-59.
Szymczak, M. R., and J. F. Corey.
Plains duck trap in Colorado.
13 p.

Michael R. Szymczak
Wildlife Researcher

1976. Construction and use of the Salt
Colo. Div. Wildl. Div. Rept. No.6.

��-69-

October

1979

JOB FUl.AJ.. REPORT

COLORADO

State of
Project

No.

Work Plan No.
Job Title
Period

Vl-88-R-24
3

Morphological

Covered:

Higratory
Job
Differences

No.
Between

Bird Investigations
10

Blue-winged

and Cinnamon

Teal

April 1, 1977 to March 31, 1979

Personnel:
B. Aufdengarten, R. Darnell, R. Ellis, P. Feiger, W. McDermitt,
M. Nail, L. Vaughn, U. S. Fish and Wildlife Service; Dr. P. Baldwin, Dr. J.
Law, L. Martin, Dr. R. Ryder, L. Sweanor, R. Thompson, K. Woodruff, Colorado
.State University; M. Conner, J. Corey, D. F#~·n.thrope; S. Harpster, R. Hopper,
R. Oakleaf, S. Steinert, and R. Stark, Colorado Division of Wildlife.
ABSTRACT
Investigations of differences in selected morphological characteristics between
blue-winged teal (Anas discors) and cinnamon teal (Anas cyanoptera) were
initiated in May 1977. Twenty females and 5 males of each species were
collected in May and June 1977 for use as a reference collection and for testing
potential techniques for distinguishing the species.
One hundred and fifty
eggs of each species were incubated, hatched, and ducklings raised at the
Fort Collins Wildlife Research Station.
Twenty-three measurements, mostly
involving the bill, were taken on the wild teal, while only 10 were taken
on the captive teal due to non-significant differences between the species or
to the difficulty of obtaining an accurate measurement on live birds.
The
captive birds were measured weekly from the 5th through the 16th week of age
and monthly from the 20th through the 56th week of age. Data were analyzed
for the 5th through the 8th, 12th, 16th, 20th, and 56th weeks.
Descriptive
statistics indicated that cinnamon teal bills were larger than bluewing bills
and significant differences (P~.05) existed between wild females of the 2
species for 16 of the 23 variables measured and for all 10 variables measured
on the captive females after the 8th week of age. Although the sample size
of wild males was small, 14 of the 23 variables were significantly different,
while all 10 of the variables were significantly different for the captive
males after the 5th week of age. Discriminant analyses for the l6-week old
captive teal indicated that 96.2% of the females and 100.0% of the males were
classified accurately when all 10 variables were included in the analysis,
but it was felt that measuring 10 variables would not be practical for field
use, and therefore, 4 were selected for use in the field based on discriminating power and ease of taking accurate measurements.
The 4 variables
selected were the lengths of the (1) center of the culmen, (2) maximum exposed
culmen, (3) culmen from the gape, and (4) center of the cu~men x the width
'
of the culmen at the gape. Discriminant analyses using logarithms of the 4
variables and based on discriminant coefficients calculated from each week's
data indicated that 100% of the captive males and &gt;92% .of the captive females

�-70-

ABSTRACT

(Continued)

were classified correctly after the 8th week of age.
Both sexes of the wild
teal were classified correctly (100.0%) by discriminant analysis using the
4 sel~cted variables and based on a discriminant coefficient calculated from
the measurements taken on the wild birds.
An average discriminant coefficent
was calculated from the weekly data for the captive teal from the 8th, 12th,
16th, 20th, and 56th weeks.
The average coefficient was used in a discriminant analysis of captive birds and essentially the same results were obtained
as when the analyses were based on weekly coefficients.
The average
coefficient was also used in a discriminant analysis of the wild teal and
all (100.%) were correctly classified, demonstrating the possibility of
developing discriminant coefficients from correctly identified sample birds
for use in classifying unidentified teal in the field.

�-71-

MORPHOLOGICAL
BETWEEN BLUE-WINGED
Robert

DIFFERENCES
AND CINNAMON

TEAL

S. Stark

Anas discors discors is one of 2 subspecies of blue-winged teal occurring in
North America.
It breeds from southern Ontario and Quebec south into the
northeastern United States, west through the north central United States
and Canada to Califonria and north through central British Columbia into
Alaska.
The northern cinnamon teal, (Anas cyanoptera septentrionalium)
hereafter referred to as cinnamon teal, is one of 5 subspecies occurring
in the western hemisphere, but the only subspecies to inhabit North America.
It breeds from the front range of the Rocky Mountains to the west coast and
from southern British Columbia and Alberta through Arizona and New Mexico
into north central Mexico (A.O.U. 1957, Delacour 1956, Johnsgard 1975).
Both
species have major fall migrations through the central and southern United
States into Mexico and Central and South America (A.O.U. 1957, Kortright
1942, Johnsgard 1975).
The breeding ranges of the 2 species overlap along the Rockies from northern
New Mexico to Alberta and British Columbia and west to eastern Washington,
Oregon, and northern California.
A problem of species identification,
especially among females, exists in this area of overlapping ranges.
Descriptions of differences between blue-winged and cinnamon teal are only
relative in most of the literature.
Sclater (1912) indicated that the female
cinnamon teal is darker and more spotted on the breast and neck than the
blue-winged teal, with only the upper throat and chin being unspotted.
Johnsgard (1975) suggested that cinnamon females show tiny, dark spotting
on the cheek, chin, and throat, while this characteristic is lacking in the
bluewing females.
These areas appear larger and light buffy to whitish in
bluewings and yellowish in cinnamons.
Wallace and Ogilvie (1977) noted this
coloration of the loral spot and also suggested that bluewing females have
a more distinct contrast between the dark crown and eyestripe than do cinnamon
females, which have a more uniform head pattern.
Bennett (1938), Kortright (1942), Johnsgard (1975), Palmer (1976), Dwyer
(1977), and Wallace and Ogilivie (1977) described the cinnamon teal female
as more brownish or reddish, especially on the breast and abdomen feathers,
compared with the more grayish cast of the bluewing female.
Kortright (1942)
and Wallace and Ogilvie (1977) warned against rust staining and Delacour (1956)
and Palmer (1976) described the cinnamon female as varying in color, showing
a tendency to a pale, as well as a dark phase.
Plumage color is probably of
questionable value as a means of distinguishing females of the 2 species
(Kortright 1942).
Spencer (1953) and Bellrose (1976) suggested that the shape of the forehead
of cinnamon teal rises more gradually to the top of the slightly flattened
head, whereas, in bluewings this rise is more abrupt.
Spencer (1953) also
mentioned the shape of the feather tract at the junction of the culmen and
forehead as a possible difference between the species.
The junction is

�-72supposedly "v" shaped in bluewings and "U" shaped in cinnamons, but he found
enough variation in one brood to reject it as useful characteristic.
The blue wing coverts of both species have also been examined as a possible
means of distinguishing
the species.
According to Spencer '(1953) and Bellrose
(1976), the blue coverts should have a slightly waxy tone in bluewings and a
more chalky tone in cinnamons.
Spencer (1953), however, found no appreciable
difference in color, hue, intensity, or pattern in the specimens he examined
and attributed most differences to individual variation.
The size and shape of the bill has been the most promising means of distinguishing females of the 2 species.
Kortright (1942) described the bill as
larger in the cinnamon teal and more constricted toward the base, while in
bluewings it is the same width throughout.
Spencer (1953) measured the length
of the exposed culmen, length of the upper mandible, length of the lower
mandible and maximum breadth on the chord of 7 blue-winged and 7 cinnamon teal
and suggested that the only measurement which showed satisfactory freedom
from overlap between the 2 species was the length of the exposed culmen.
The
results of other investigations have also found no overlap in the ranges of
culmen length for the 2 species (Phillips 1923, Duvall cited in Spencer 1953),
but Johnsgard (1975) noted that the species have overlapping measurements for
both bill length and width.
Johnsgard (1975) suggested that cinnamon teal have slightly longer soft
lateral margins of the upper mandible distinctly drooping over the lower
mandible near the tip, creating a semi-spatulate appearance when viewed from
the side. Wallace and Ogilivie (1977) also noted this profile, but warned
that bluewings can show the same characteristic at times.
Male blue-winged and cinnamon teal present a similar problem of identification
prior to about 8 weeks of age, or until the cinnamon male acquires the red eye
color of the adult (Spencer 1953). They are easily distinguishable
in any
plumage after that time. The tracheal bulla can also be used to distinguish
the males, being 2-3 times larger in the cinnamons and rather differently
shaped (Phillips 1923, Palmer 1976) .. It was not suggested whether this trait
is useful for distinguishing females or males less than 8 weeks of age.
The size and shape of the lachrymal bone appears to be the best characteristic
for distinguishing blue-winged and cinnamon teal (Spencer 1953) but it is not
applicable to banding studies.
It may, however, aid in accurately identifying museum specimens in which the bone has been retained.
In the major duck production areas of Colorado, blue-winged and cinnamon teal
are important breeding species and represented, jointly, an average of 11%
of the breeding population during the period 1954-75 (Szymczak 1977).
Combined, they are second only to the mallard (Anas platyrhynchos) in annual
production in Colorado and have contributed up to nearly 20,000 birds to
the state's breeding population in recent years.
During the pre season banding period in Colorado, for the years 1971-74, over
75% of the blue-winged and cinnamon teal banded were classified as unidentified teal because they could not be accurately identified as one species or
the other (R. Hopper, pers. comm.).
Knowledge of possible population
differences is essential to the formulation of management recommendations,
but is not currently available for blue-winged and cinnamon teal within the

�-73-

state.
Special early teal seasons currently being conducted in various parts
of the U. S. cannot be justified in much of Colorado's high country until
the status of both blue-winged and cinnamon teal populations within the state
are determined.
A management scheme could then be formulated which would
give greatest consideration to the species with the poorest population status.
This can be accomplished only after a reliable technique is developed for
distinguishing between the 2 species in the field, involving one which does
not result in the death of the birds.
P. N. OBJECTIVES
1.

To compare
sex.

the morphology

of blue-winged

and cinnamon

teal by age and

2.

To develop a field technique for distinguishing between blue-winged and
cinnamon teal based on morphological differences, primarily for use
during the pre-season banding period (August and September).
METHODS AND MATERIALS
Collection

of Adult Teal

Thirteen cinnamon teal (5 males, 8 females) and 1 female blue-winged teal
were collected in the San Luis Valley, Colorado on 16 and 17 May 1977. On
26 May 1977, 12 blue-winged teal (5 males, 7 females) were collected at the
Red Lion State Wildlife Area near Crook, Colorado.
Only females in the
presence of a male were collected.
An additional 24 female teal were collected concurrently with the egg collection phase of the study described below, 12 blue-winged teal from the
Valentine National Wildlife-Refuge
in north central Nebraska and 12 cinnamon
teal from the Alamosa and Monte Vista National Wildlife Refuges in the San
Luis Valley of Colorado.
Species identification of these 24 females was
deferred until young males from their respective clutches acquired nuptial
plumage in the spring and summer of 1978 while in captivity at the Fort Collins
Wildlife Research Station.
All collections were made with a shotgun and all 50 birds (5 males and 20
females of each species) were weighed and tagged with an identification
number within 24 hours after collection.
They were also measured and then
frozen for use as a reference collection of adult wild teal. The form used
to record the measurements taken in presented in Appendix A.
Egg Collections
On 7 June 1977, 150 suspected blue-winged teal eggs (and 12 adult female bluewings associated with the nests) were collected from the Valentine NWR and
transported to the Fort Collins Wildlife Research Station.
Nests were located
by flushing incubating females with a 48.8 m cable dragged through nesting
habitat between two vehicles, as described by Ladd (1969). All nests were
located and marked prior to collecting eggs and all eggs were individually
identified by nest and female at the time of collection.
The eggs were

�-74-

transported in a portable incubator designed to operate from a car battery
and maintained a temperature of 37.2oC (990F) and a hygrometer reading of
28.90C (840F) (Flieg 1974, Stromberg 1975).
During the period 6 to 21 June 1977, 150 suspected cinnamon teal eggs (and
12 adult female cinnamons associated with the nests) were collected from
the Alamosa and Monte Vista National Wildlife Refuges.
Nests were located
by crews of 4-6 people walking systematically through likely nesting habitat
and flushing incubating females.
Collectipn and transport procedures were
similar to those used during the Nebraska collection, except some clutches
were placed in a chest cooler with cotton wadding and hot water bottles and
flown to Fort Collins.
The ether wafer in the incubator at the Fort Collins Wildlife Research Station
ruptured on 27 June 1977, resulting in over-heating and death of 75 embryos
in eggs collected in Colorado.
An additional 75 suspected cinnamon teal eggs
collected on 30 June and 1 July from the Alamosa National Wildlife Refuge
to insure than an adequate sample of cinnamon teal was obtained.
Hatching

and Rearing

of Young

The eggs were not washed prior to setting in the incubator, but excessively
dirty eggs were scraped with a knife (Sheraw 1975). All eggs were weighed
and the length and width measured prior to placement in the incubator.
A
Humidaire Model 50 automatic incubator and a David Bradley incubator (hatcher)
were used for incubating and hatching the eggs at the Fort Collins Wildlife
Research Station.
The incubator was set to maintain a temperature of 37.50C
0
(99.5 F), and a hygrometer reading of 30.0oC (86.00F) (Ellis 1974, Flieg
1974, Stromberg 1975).
The automatic turning device on the incubator was
set to turn the eggs every 2 hours (Stromberg 1975). The hatcher was set
at a temperature of 37.2oC (99.00F) and a hygrometer reading of 34.4°C
(94.0oF) (Ellis 1974, Flieg 1974, Stromberg 1975).
The eggs were not turned
in the hatcher and during the incubation period the water pans and hygrometer
bottles were filled daily with distilled water (Stromberg 1975).
Trays in the hatcher were separated into 5 compartments and each was enclosed
by hardware cloth.
Each clutch of eggs was placed in an individual compartment in the hatcher when the first egg of the clutch pipped.
Ducklings from
each clutch were thereby separated from other clutches after hatching for
accurate identification
(Dill and Lee 1970). The ducklings were allowed to
dry several hours after hatching before being removed from the hatcher
(Grow 1972). Upon removal, they were weighed and banded with individually
numbered plastic leg bands and recorded by clutch.
After banding, the ducklings were placed in a battery brooder with the temperature maintained at 32.2oC (90.00F) (Dill and Lee 1970, Lee 1974). Newspaper was placed on the floor of each of the 5 brooder compartments and an
unmedicated starter feed (28.5% crude protein) (F. Lee, pers. comm.) prepared
by the Golden West Milling Company, Longmont, Colorado was sprinkled on the
paper to induce feeding (Smart 1974). Two waterers were placed in each
brooder compartment and the water was changed 4-5 times daily.
Feed was
placed in small feeders and changed twice daily after the ducklings were
eating.

�-75-

The ducklings were fed an unmedicated duck maintenance feed (18.3% crude
protein) (F. Lee, pers. comm.) after 12 days of age (Dill and Lee 1970). At
this time they were also removed from the battery brooder, as recommended
by Sheraw (1975), rebanded with individually numbered aLuminum leg bands and
placed in a brooder house.
The brooder house was 2.4 x 2.7 m and the floor
was covered with wood shavings litter (J. Law, pers. comm.).
Four large
waterers and 3 feeders were placed on the litter and the water and feed
were changed 3 times daily.
Four 250 watt infrared heat lamps were placed
above the litter and left on 24 hours a day (Lee 1974).
At 3 weeks of age the ducklings were allowed access to a 1.5 x 2.4 m section
of a concrete pond adjacent to the brooder house.
The ducklings were allowed
to swim daily to induce functioning of the oil gland and preening (Bauer 1974).
At 5 weeks of age the ducklings were rebanded permanently with size 5 U. S.
Fish and Wildlife Service leg bands and moved to an outside pen. The pen
measured 9.1 x 22.9 m and was completely enclosed in chicken wire, including
the top. Several small shelters were placed in the pen to provide shade,
and a concrete pond 2.4 x 11.0 m was built to provide water.
Feed was
changed twice daily in the 3 feeders.
The pond water was changed every
third day.
The teal were moved in November 1977 to a 5.5 x 9.7 m pen for wintering.
This facility was divided into 3 compartments, each 1.8 x 9.7 m. The pen was
raised 0.6 m above the ground and had a welded wire floor to allow waste
material to pass through.
Each compartment was divided into 2, 1.8 x 4.9 m
sections, with one entirely enclosed with weatherproofing
material and
provided with 8, 250 watt infrared heat lamps to maintain an above freezing
temperature (Strutz 1974). The other section was weatherproofed
all all
except one side, but not provided with heat. A small 0.3 x 0.3 m door
allowed passage between the 2 sections.
Feeders and waterers were placed in
the heated section and chaged twice daily.
Birds surviving into November
were separated by week of hatching into the 3 compartments of the winter pen.
The teal remained in the winter pen through March 1978 and were moved back
to the outside pen in early April 1978.
Measurements

of Wild and Captive Teal

In this study, culmen referred to the upper mandible, mandible was used to
describe the lower mandible, and bill referred to both the upper and lower
mandibles.
Wild Teal
Twenty-three measurements, based on Baldwin et ale (1931), were taken on the
wild teal including the tarsus (TAR), length of the center of the culmen
(CCEN) (Fig. 1), length of the maximum exposed culmen (MEC) (Fig. 1), length
of the culmen from the gape (CGAPE) (Fig. 2), width of the culmen at the
widest anterior point (WIANT) (Fig. 3), width of the culmen at the gape
(WIGAPE) (Fig. 3), length of the center of mandible (LBCEN) (Fig. 4), length
of the exposed mandible (LEM) (Fig. 4), height of the bill at the base (BHGT).
(Fig. 5), length of the foot (LFOOT), length of the middle toe (LMTOE),
width of the culmen at the nares (WINARES), width of the mandible at the
widest anterior point (MWIANT), width of the mandible at the gape (MWIGAPE),

�-76-

---------_
...... --~---

-

Fig. 1. Length of the center of the culmen
maximum exposed culmen (MEC).

(CCEN) and length of the

•
Fig. 2.

Length of the culmen to the gape (CGAPE).

�--77-

Fig. 3. Width of the culmen at the widest anterior
of the culmen at the gape (WIGAPE).

Fig. 4. Length of the center of the mandible
exposed mandible (LEM).

point

(WIANT) and width

(LBCEN) and length of the

�-78-

LHC

Fig. 5. Length of the head and total culmen (LHC) and height of the bill
at the base (BHGT).

�-79-

length of the culmen nail (LCNAIL), width of the culmen nail (WCNAIL),
length of the mandible nail (LMNAIL), width of the mandible nail (ill1NAIL),
length of the nares (LN) , height of the bill at the nares (BHGTN), height
of the bill at the nail (BHGTNAIL), wing length (WINGL), and weight (WGT).
Captive Teal
Ten measurements were taken on the captive teal in the sample at weekly
intervals from the 5th through the 16th week of age and monthly from the 20th
through the 56th week.
These measurements included the first 9 listed above,
plus the length of the head and total culmen (LHC) (Fig. 5). The wing
length was not taken on the captive birds due to the tips of the primaries
breaking off in the pen. Weight was recorded for the captive birds but was
not included in the data analysis due to the artificial conditions under
which the birds were raised.
Variations

in Sample Size .

The sample size varied almost weekly throughout the study for a variety of
reasons.
The most common cause was birds escaping while being measured or
escaping from the catch box they were held in prior to being measured.
Deaths
were also a cause of variation, but measurements were initiated on a replacement bird after the death of a bird in the sample.
Some of the birds were
extremely active during the measuring process and if it was felt that an
accurate measurement was not being obtained, the value for the variable was
not recorded.
Computer

Programs

The graphs and descriptive statistics were programmed at Colorado State
University for a Cyber 172 computer, and the discriminarit analyses and ~tests were programmed in the statistical package for the Social Sciences
(SPSS) (Nie et ale 1975). A detailed discussion of discriminant analysis
can be found in Nie et ale (1975) and Kleinbaum and Kupper (1978).
RESULTS AND DISCUSSION
Descriptive

Statistics

and Hean Compari.sons

Wild Teal
Data for the wild teal were initially analyzed with descriptive statistics,
including the mean, standard error of the mean, standard deviation, and
range (minimum and maximum values) (Table 1 and 2). Also included in this
analysis was ~-test information for interspecific mean comparisons of the
variables for both sexes.
Significant differences (P~.05) existed between the 2 species of wild females
for all but 7 of the 23 variables (Table 1). The cinnamon females had larger
mean values for all variables except the weight, length of the mandible nail,
and width of the culmen nail.
The length of the center of the culmen (CCEN)
was the only variable with no overlap in ranges between the 2 species.

�Table 1.

Descriptive statistics and t-tests for wild female blue-winged and cinnamon teal.

Blue Winged Teal
Mean + SE
SD

Range

Cinnamon Teal
Mean + SE
SD

t-test
Range

DF 1/

Variable

N

WGT

20

370.75 + 8.478

37.91

315.00-455.00 20

359.25 + 9.454

42.28

280.00-440.00 37.56

.371

WINGL

20

174.25 + 1.63

7.31

153.00-184.00 20

180.68 + 1.018

4.43

173.00-190.00 31.57

.002

TAR

20

38.70 +

.255

1.14

37.10- 41.00

20

39.22 +

.224

1.00

36.90- 40.90

37.37

.135

LFOOT

20

40.37 +

.313

1.40

37.30- 42.50

20

41.38 +

.374

1.67

38.00- 44.70

36.87

.045

LMTOE

20

34.72 +

.302

1.35

31.50- 37.00

20

35.92 +

.295

1.32

34.00- 38.50

37.98

.007

CCEN

20

38~38 +

.226

1.01

36.50- 40.10

20

43.09 +

.347

1.55

40.75- 45.85

32.70

.001

MEC

20

44.49 +

.294

1.32

42.25- 47.10

20

48.68 +

.355

1.59

45.65- 51.90

36.74

.001

CGAPE

20

42.24 +

.265

1.83

40.15- 45.20

20

45.95 +

.331

1.48

43.40- 48.50

36.26

.001 ~

WIANT

20

17.05 +

.131

.59

16.10- 18.25

20

17.61 +

.144

.64

16.50- 19.20

37.67

.006

WINARES

20

15.92 +

.136

.61

14.50- 16.80

20

16.95 +

.120

.54

16.15- 18.55

37.44

.001

WIGAPE

20

13.98 +

.139

.60

12.85- 15.25

20

14.65 +

.181

.81

13.00- 16.15

35.08

.005

LBCEN

20

35.78 +

.360

1.61

31.80- 38.30

20

39.49 +

.338

1.51

36.65- 43.20

37.85

.001

LEM

20

43.62 +

.318

1.42

41.05- 47.20

20

47.24 +

.382

1.71

43.60- 49.95

36.79

.001

MWIGAPE

20

11.76 +

.139

.62

10.45- 13.00

20

12.36 +

.164

.73

11.20- 13.60

37.06

.009

MWIANT

20

11.32 +

.096

.43

10.75- 12.40

20

11.86 +

.072

.32

11.30- 12.40

35.07

.001

LMNAIL

20

6.26 +

.126

.55

5.50- 7.10

20

6.11 +

.115

.51

4.85- 6.80

36.50

.375

WMNAIL

20

5.30 +

.078

.34

4.60- 6.25

20

5.84 +

.115

.51

4.45- 6.70

33.25

.192

NL

20

3.38 +

.056

.25

2.90- 3.80

20

3.70 +

.068

.30

3.15- 4.15

36.73

.001

LCNAIL

20

7.98 +

.114

.51

6.80- 8.90

20

8.17 +

.112

.50

7.05- 8.80

38.00

.248

WCNAlL

20

5.65 +

.077

.34

5.00- 6.25

5.51 +

.088

.39

5.00- 6.30

37.33

.249

BHGT

20

15.62 +

.166

.74

14.60- 16.90

16.09 +

.212

.95

14.80- 18.10

35.90

.086

N

P

.

o
I

�Table 1.

Descriptive

statistics

and ~-tests

for wild female blue-winged

Variable

N

Blue '.,Tinged
T~_a_l
Mean + SE
SD

BHGTN

20

11.73 +

.114

.51

10.90- 12.60

20

BHGTNAIL

20

8.61 +

.099

.44

7.95-

20

1/ Weight

in grams and all other measurements

2/

-

Calculated

_

Range

9.75

and cinnamon

teal (continued).
t-test

Teal

N

Cinnamon
Hean + SE
12.44

+ .134
9.03 + .143

.60

11.00- 13.25

37.01

.00.1

.64

7.25-

33.88

.022

SD

--------

DF

Range

9.90

p

in millimeters.

d.f.:
u

=

(~i 2) 2
+

nl

s2

n2

(;~}G:Y
ml-l

m2-l

I

co
I-'
I

�Table

2.

Descriptive

statistics

Blue

and

Winged

+ SE

t-tests

for

wild

male

b l.ue-w i.nged

and

Teal

SD

Range

N

362.00 + 9.695

21.68

335.00-395.00

5

5

182.60 +

.748

1.67

180.00-184.00

TAR

5

39.30 +

.460

.92

LFOOT

5

41.82 +

.631

LMTOE

5

35.50 +

CCEN

5

MEC

cinnamon

t e a L,

Cinnamon

Teal

+ SE

t-test

SD

Range

DF

P

394.00 + 10.654

23.82

355.00-415.00

7.93

5

201.20 +

6.492

14.52

185.00-220.00

4.11

.057
.047

38.60- 40.60

5

41.13 ±

.437

.98

39.70- 42.25

6.74

· 02 II

1. 41

39.35- 42.80

5

43.90 ±

.470

1.05

42.60- 45.50

7.39

.033

.775

1. 73

33.00- 37.00

5

37.20 +

.300

.67

36.00- 37.50

5.17

.096

39.21 +

.980

2.19

37.05- 42.40

5

44.24 +

.791

1.77

42.10- 46.50

7.66

.004

5

46.07 +

.885

1.98

44.10- 48.95

5

51.39 +

.666

1. 49

49.85- 53.50

7.43

.002

CGAPE

5

43.32 +

.897

2.00

41.00- 46.35

5

48.71 +

.700

1. 56

46.80- 51.00

7.55

.001

~

WIANT

5

17.93 +

.337

.75

16.65- 18.65

5

18.17 +

.191

.43

17.55- 18.50

6.32

·558

I

WINARES

5

16.20 +

.357

.80

15.00- 17.00

5

18.02 +

.211

.47

17.25

18.40

6.50

.005

WIGAPE

5

15.16 +

.465

1.04

14.45- 17.00

5

16.29 +

.206

.46

15.60- 16.80

5.52

.068

LBCEN

5

37.69 +

.829

1. 85

35.65- 39.90

5

40.85 +

.924

2.07

38.60- 43.40

7.91

·034

LEM

5

44.80 + 1.308

2.93

41.05- 48.95

5

49.92 +

.852

1. 90

48.25- 52.35

6.88 .

.013

MWIGAPE

5

12.12 +

.440

.98

10.70- 13.10

5

13.71 +

.185

.41

13.10- 14.15

5.38

.021

MWIANT

5

11.97 +

.064

.14

11.75- 12.15

5

12.18 +

.106

.24

11.80- 12.35

6.62

.133

LMNAIL

5

5.59 +

.208

.46

5.20-

6.35

5

6.45 +

.266

.60

5.50-

7.10

7.55

·034

WMNAIL

5

5.20 +

.097

.22

4.95-

5.45·

5

5.42 +

.247

.55

4.50-

5.85

-5.21

.446

NL

5

3.68 +

.097

.22

3.45-

3.95

5

4.18 +

.119

.27

3.80-

4.50

7.69

.012

LCNAIL

5

7.70 +

.139

.31

7.35-

8.10

5

8.47 +

.215

.48

7.85-

9.00

6.83

.020

WCNAIL

5

5.45 +

.130

.29

5.10-

5.80

5

5.38 +

.094

.21

5.15-

5.70

7.29

·677

BHGT

5

16.23 +

.241

.54

15.55- 16.85

5

16.96 +

.663

1. 48

14.80- 18.70

5.04

·348

BHGTN

5

12.37 +

.227

.51

11.75- 13.00

5

13.03 +

.334

.75

12.00- 13.95

7.04

· 146

BHGTNAIL

5

9.11 +

.119

.27

8.70-

5

10.38 ±

.286

.64

9.85- 11.40

5.34

.009

Variable

N

WGT

5

WINGL

Mean

9.35

Mean

~"

�0
0

&lt;D
:::I'

b
0

E
E

z
w
o

o

0
:::I'

0
0
N

en

0
0
:::I'
(\1

I
(X)

W
I
0
0

&lt;D
:::I'

E
E

0
0
0
:::I'

Z
W

o
o

0
0

BWT

N
(D

0
0
:::I'

~.OO

I

5.60

11.20

16.80

22.1I0

28.00

33.60

39.20

1I11.80

50. LID

~ 1

51:

AGE IWEEKSI

Fig. 6. Comparative profile
of the length
blue-winged and cinnamon teal.

of the c·enter of the culmen (CCEN)measurements for female
.

�Table 3.

Descriptive statistics and t-tests for female blue-winged and cinnamon teal at 5 weeks of age.

Variable

N

Blue Winged Teal
Mean + SE
SD

TAR

33

37.94 +

.186

CCEN

33.

36.77 +

MEC

33

CGAPE

Range

N

Cinnamon Teal
Mean + SE
SD

1.07

35.35 - 40,15

33

39.04 + .205

.233

1.34

34.70 - 39.25

33

41.67 +

-

.242

1.39

39.05 - 44.80

33

39.71 +

.217

1.25

WIANT

33

15.76 +

.082

WIGAPE

33

14.79 +

LBCEN

33

LEM

t-test
Range

DF

P

1.18

36.85 - 41. 30

63.41

.001

38.13 + .276

1.59

34.90 - 41.20

62.22

.001

33

43.53 + .300

1.72

40.35 - 47,00

61.28

.001

37.10 - 42.30

33

41.03 + .297

1.70

36.95 - 44.85

58.68

.001

.47

14.95 - 17.00

33

16.20 + .096

.55

14.80 - 17.30

62.63

.001

.070

.40

14.05 - 15.75

33

15.52+.110
-

.63

14.45 - 16.50

54.29

.001

32.65 +

.269

1.54

29.70 - 35.10

33

38.38 + .273

1.57

30.25 - 36.60

63.98

.062

33

40.56 -+

.250

1.44

36.90 - 43.30

33

42.09 + .285

1.64

38.05 - 45.90

62.96

.001 I

BHGT

33

7.80 +

.072

.42

7.00 -

8.65

33

8.55 + .066

.38

7.90 -

63.50

.001 t

LHC

33

75.88 +

.318

1.83

71.70 - 80.35

33

77.37 + .428

2.46

72.35-82.15

59.09

.007

-

00

9.30

�Table

4.

Descriptive

statistics

and

Blue

Teal
SD

Range

N

Mean

1.02

36.75 - 40.25

33

1.35

36.35 - 40.90

Winged

+ SE

.!_-tests

for

female

blue-winged

and

cinnamon

teal

Cjnnamon
+ SE

at

6 we ek s of

age.

t-test

Teal
SD

Range

DF

P

39.21 + .187

1.04

37.00 - 41.05

61. 94

.001

33

39.98 + .285

1.59

37.40 - 43.20

60.15

.001

Variable

N

Mean

TAR

32

CCEN

32

MEC

32

1.41

40.00 - 45.85

33

45.69 + .3l7

1.77

42.70 - 49.10

58.74

.001

CGAPE

32

+ .181
38.46 + .239
43.36 + .250
41. 69 + .. 206

1. 16

39.40 - 44.15

33

43.49 + .299

1. 67

40.05 - 46.70

54.98

.001

WIAJ.1"T

32

16.18 + .077

.43

15.35 - 17.25

33

16.63 + .090

.50

15.35 - 17.65

60.43

.001

WIGAPE

32

14.98 + .061

.35

14.35 - 15.70

33

15.84 + .094

.55

14.70 - 16.90

53.37

.001

LBCEN

32

34.45 + .249

1.41

31. 60 - 36.70

33

35.37 + .293

1. 65

32.30 - 38.85

60.47

.027

LEM

32

1.12

39.90 - 44.95

33

44.58 + .302

1. 68

41.30 - 48.85

53.56

.001

BHGT

32

42.57 + .199
8.15 -+ .052

.29

7.65 -

8.70

33

8.81 + .053

.30

8.00 -

9.35

61.94

.001 ~

33

80.87 + .394

2.20

77.05 - 85.00

58.25

.001

LHC

32

38.34

78.99 + .304

1.72

74.85 - 82.60

I
I

�Table 5.

Descriptive statistics and t-tests for female blue-winged and cinnamon teal at 7 weeks of age.

Variable

N

Blue Winged Teal
Mean + SE
SD

Range

N

Cinnamon Teal
Mean + SE
SD

TAR

33

38.85 + .172

.98

37.00 - 40.30

35

39.50 + .196

CCEN

33

39.28 + .212

1.22

37.00 - 41.40

35

MEC

33

44.30 + ~236

1.33

41.40 - 46.95

CGAPE

33

42.60 + .204

1.16

WIANT

33

16.48 + .079

WIGAPE

33

LBCEN

t-test
Range

DF

P

1.16

37.10 - 42.10

64.55

.019

41.38 -+ .271

1.60

37.70 - 44.40

62.70

.001

35

47.18 + .313

1.85

43.65 - 51.05

61.67

.001

40.75 - 45.00

35

44.94 + .269

1.59

42.05 - 48.05

61.92

.001

.44

15.55 - 17.60

35

16.92 + .076

.45

15.80 - 17.75

64.57

.001

15.13 + .073

.40

14.35 - 16.40

35

16.00 + .087

.51

15.15 - 17.05

63.94

.001 I

33

35.20 + .259

1.44

32.40 - 37.70

35

36.69 + .270

1.60

33.70 - 39.85

65.00

.001 I

LEM

33

43.19 + .210

1.18

40.85 - 45.70

35

45.93 + .248

1.46

43.25 - 48.25

64.14

.001

BHGT

33

8.42 + .073
-

.41

7.85 -

9.35

35

9.11 + .058

.34

8.30 -

9.95

60.43

.001

LHC

33

80.16 + .291·

1.65

76.60 - 83.85

35

82.78 + .414

2.42

78.05 - 87.75

58.49

.001

co

&lt;1'

�Table 6.

Descriptive statistics and t-tests for female blue-winged and cinnamon teal at 8 we eks of age.

Blue Winged Teal
Mean + SE
SD

Range

N

Cinnamon Teal
Mean + SE
SD

.94

37.50 - 40.40

34

39.75 + .198

39.41 + .207

1.19

36.80 - 41. 65

34

33

44.65 + .237

1.36

41.85 - 46.80

CGAPE

33

42.85 + .178

1.02

WIANT

33

16.66 + .074

WIGAPE

33

15.33 + .072

Variable

N

TAR

33

39.05 + .164

CCEN

33

MEC

t-test
Range

DF

P

1.15

37.20 - 42.00

63.15

.008

42:.07 + .271

1.58

38.95 - 44.90

61. 19

.001

34

47.94 + .293

1.71

44.75 - 51.40

62.63

.001

41.00 - 44.80

34

45.66 + .257

1.50

42.20 - 48.50

58.30

.001

.43

15.60 - 17.85

34

17.10 + .086

.50

15.70 - 18.05

64.00

.001

.41

14.60 - 16.20

34

16.16 -+ .088

.51

15.15 - 17.45

62.87

.001 I
co
--J

LBCEN

33

35.27 + .249

1.43

32.90 - 37.85

34

37.32 + .264

1.54

34.15 - 40.00

64.87

.001 I

LEM

33

43.16 + .207

1.19

40.85 - 45.65

34

46.36 + .291

1.70

Id.85

-

49.00

59.15

.001

BHGT

33

8.60 + .058

.33

8.10 -

9.30

34

9.23 + .053

.31

8.55 -

9.80

64.22

.001

LHC

33

80.12+.277

1.59

76.70 - 83.75

34

83.76 + .351

2.05

79.55 - 88.40

62.07

.001

�Table 7.

Descriptive statistics and _!-tests for female blue-winged and cinnamon teal at 12 we eks of age.

Blue Winged Teal
SD
Mean + SE

Range

N

Cinnamon Teal
Mean + SE
SD

.99

37.45 - 40.70

22

40.12 + .269

39.38 + .220

1.23

36.70 - 41.55

22

32

44.71 + ;250

1.37

42.20 - 46.75

CGAPE

32

42.67 + .202

1.04

WIANT

32

16.87 + .084

WIGAPE

32

LBCEN

Variable

N

TAR

32

39.30 + .192

CCEN

32

MEC

t-test
Range

DF

P

1.26

37.45 - 41.85

39.77

.015

41.87 + .316

1.48

38.95 - 44.40

39.10

.005

22

47.97 + .321

1.51

44.60 - 49.85

42.14

.001

40.95 - 44.50

22

45.40 + .299

1.40

41.90 - 48.10

38.38

.001

.47

15.85 - 17.85

22

17.23 + .085

.40

16.40 - 18.00

47.04

.005

15.55 + .072

.36

14.90 - 16.40

22

16.26 -+ .082

.38

15.75 - 17.00.

44.93

.001 I

32

35.12 + .250

1.38

32.70 - 37.60

22

37.11 + .357

1.67

33.90 - 39.50

39.35

.001 I

LEM

32

43.14+.240

1.23

40.40 - 45.50

22

46.15 + .316

1.48

42.30 - 48.35

41.49

.001

BHGT

32

8.79 + .061

.34

8.15 -

9.50

22

9.53 + .065

.31

9.05 - 10.05

46.22

.00]

LHC

32

80.27 + .320

1.63

77.80 - 83.85

22

83.41 + .405

1.90

79.10 - 86.75

42.13

.001

00
00

�Table 8.

Descriptive

statistics

and ~-tests

for female blue-winged

and cinnamon

teal at 16 weeks

Teal
SD

Range

N

Cinnamon
Mean + SE

of age.

t-tcst

Variable

N

Blue Winged
Mean + SE

Teal
SD

Range

DF

P

TAR

32

39.49

+ .193

.92

37.90 - 40.85

27

40.34

+ .231

1. 18

37.90 - 42.10

41. 93

.006

CCEN

32

39.23

+ .273

1.23

36.55 - 4l.40

27

41.90

+ .353

1.56

38.50 - 44.45

40.41

.001

MEC

32

44.68

+ .296

1.33

42.40 - 46.90

27

48.31

+ .340

1.55

45.20 - 51.30

42.59

.001

CGAPE

32

42.65 + .237

1.08

40.90 - 44.70

27

45.65 + .313

1.39

42.00 - 48.15

40.02

.001

WIANT

32

16.88 + .102

.52

15.70 - 17.95

27

17.37 + .103

.47

16.35 - 18.25

43.88

.005

WIGAPE

32

15.61 + .078

.40

14.70 - 16.55

27

16.51 + .102

.46

15.90 - 17.55

40.09

.001 do
\0

I

LBCEN

32

35.29 + .305

1.36

32.90 - 37.85

27

37.38

+ .357

1.63

34.60 - 40.05

42.27

.001

LEM

32

43.14 + .269

1.20

40.75 - 45.80

27

46.27 + .345
-

1.55

42.20 - 48.85

40.62

.001

BHGT

32

8.80 + .070

.35

8.10 -

9.55

27

9.60 + .075

.35

9.10 - 10.40

43.46

.001

LHC

32

80.11

+ .387

1.73

76.70 - 84.20

27

83.71 + .465

2.25

79.05 - 88.50

41.84

.001

�Table 9.

Descriptive statistics ilnd t-tests for female blue-winged and cinnamon teal at 20 weeks of age.

Variable

N

TAR

30

39.34 + .177

CCEN

30

MEC

Blue Winged Teal
Mean + SE
SD

Range

N

Cinnamon Teal
Mean + SE
SD

.96

37.90 - 40.90

25

40.58 + .219

39.07 + .249

1.32

36.35 - 41.65

25

30

44.37 + .252

1.34

41.95 - 47.25

CGAPE

30

42.48 + .198

1.05

WIANT

30

16.76 + .097

WIGAPE

30

15.60 + .068

t-test

- P

Range

DF

1.08

38.65 - 42.50

46.56

.001

41.84 + .307

1.51

38.85 - 44.60

46.61

.001

25

48.21 + .352

1.71

44.70 - 51.35

43.31

.001

40.50 - 44.60

25

45.72 + .319

1.55

41. 90 - 48.30

39.28

.001

.51

15.60 - 17.70

25

17.36 + .103

.49

16.25 - 18.25

49.69

.001

.37

14.90 - 16.55

25

16.58 + .099

.48

15.75 - 17.50

42.24.

.001 I
IJ)

a

LBCEN

30

35.29 + .238

1.33

33.00 - 38.00

25

37.35 + .335

1.62

34.05 - 39.80

43.05

.001 I

LEM

30

43.05 + .229

1.22

40.55 - 45.70

25

46.38 + .334

1.62

42.00 - 48.85

42.14

.001

BHGT

30

8.86 + .071

.38

8.10 -

9.50

25

9.70+

.078

.38

9.00 - 10.40

49.07

.001

LHC

30

80.05 + .319

1.71

77.10 - 83.65

25

84.10 + .475

2.29

78.40 - 88.60

41.56

.001

�Table

10.

Descriptive

statistics

Blue Winged
Mean + SE

and J:.-tests for female blue-winged

and cinnamon

teal at 56 we ek.s of age.

Teal
SD

Range

N

Cinnamon
Mean + SE

Teal
SD

Range

DF

P

t-test

Variable

N

TAR

29

39.47 + .196

1.03

37.15 - 41.15

26

40.41 + .2t16

1.25

37.85 - 42.35

47.66

.002

CCEN

29

39.31 + .272

1.34

36.40 - 41. 75

28

42.31 + .295

1.53

39.05 - 44.95

50.81

.001

MEC

29

44.72 + .247

1.30

42.45 - 47.15

28

48.53 + .335

1. 74

45.40 - 51. 70

47.38

.001

CGAPE

29

42.70 + 2.15

1.11

40.80 - 45.00

28

45.78 + .285

1.47

42.05 - 48.20

47.86

.001

WIANT

29

16.87 + .106

.54

15.50 - 17.95

28

17.36 + .097

.49

16.20 - 18.20

50.36

.001

WIGAPE

29

15.61 + .080

.42

14.65 - 16.50

28

16.63 + .096

.49

15.85 - 17.65

49.68

.001

I
\D
I-'

LBCEN

29

35.05 + .248

1.30

33.05 - 37.75

28

37.29 + .328

1.69

33.95 - 39.60

47.98

.001

LEM

29

43.03 + .239

1.24

40.45 - 45.35

28

46.40 + .300

1.55

41.90 - 48.95

48.93

.001

BHGT

29

8.90 + .085

-

.43

8.05 -

9.55

28

9.77+.071

.37

9.05 - 10.55

49.22

.001

LHC

29

80.03 + .289

1.59

76.70 - 83.40

28

83.88 + .428

2.19

79.40 - 88.65

45.32

.001

I

�Table 11.

Descriptive statistics and ~-tests for male blue-winged and cinnamon teal at 5 weeks of age.

Blue Winged Teal
Mean + SE
SD

Range

N

Cinnamon Teal
Mean + SE
SD

1.21

36.80 - 41.30

29

39.80 + .238

37.81 + .177

.98

36.00 - 39.35

29

31

43.52 + ~224

1.25

40.85 - 45.60

CGAPE

31

41.50 + .190

1.06

WIANT

31

16.08 + .069

WIGAPE

31

LBCEN

Variable

N

TAR

31

38.63 + .217

CCEN

31

MEC

t-test
Range

DF

P

1.28

36.85 - 41.85

57.11

.001

39.46 + .264

1.42

.36.40 - 42.35

49.41

.001

29

45.57 -+ .228

1.23

43.05 - 47.50

57.84

.001

39.10 - 43.40

29

42.74 + .230

1.24

39.90 - 45.20

55.29

.001

.38

15.25 - 16.80

29

16.60 + .099

.53

15.55 - 17.50

50.69

.001

15.15 + .064

.36

14.35 - 15.85

29

15.94 + .116

.63

14.95 - 17.30

43.76

.001

31

34.18 + .233

1.30

31.90 - 37.15

29

34.74 + .294

1.58

31.00 - 38.10

54.25

.140 I

LEM

31

42.82 + .241

1.34

LfO.70

29

44.05 -+ .321

1.73

39.05 - 47.00

52.83

.003

BHGT

31

8.23 + .067

.37

7.45 -

8.85

29

8.84 + .088

.47

7.95 -

9.70

53.38

.001

LHC

31

78.82 + .273

1.52

76.20 - 83.00

29

80.30 + .440

2.37

74.50 - 85.10

47.18

.006

- 45.50

I
1.0
N

�Descriptive

statistics

and _.S_-testsfor male blue-winged

Variable

N

Blue Winged
+ SE

Teal
SD

Range

N

TAR

34

39.20 + .197

1. 12

37.40 - 41.50

30

CCEN

34

39.61

+ .178

1.05

36.30 - 41. 30

MEC

34

45.58 + .211

1.23

CGAPE

34

43.69 + .202

_-

WIANT

34

WIGAPE

34

LBCEN

Table

12.

and cinnamon

teal at 6 we eks of age.

Cinnamon
Mean + SE

t-test

Teal
SD

Range

40.42 -+ .188

1.03

38.40 - 41.95

61 .DO

.001

30

41.89 -+ .252

1. 38

39.45 - 44.40

53.23

.001

42.05 - 47.35

30

48.29 + .189

1.04

46.15 - 50.05

60.80

.001

1.16

41.05 - 45.60

30

45.58 + .192

1.05

43.10 - 47.85

6 J .00

.001

16.49 -+ .064

.37

15.65 - 17.10

30

17.12 + .093

.51

16.30 - 18.05

52.43

.001

15.35

+ .065

.37

14.40 - 16.30

30

16.33 + .100

.55

15.25 - 17.65

50.19

.OOl

34

35.96

+ .234

1.34

32.75 - 39.00

30

37.17 + .237

1. 30

35.00 - 39.70

60.76

.001

LEM

34

44.88 + .241

1.39

42.10-47.15

30

47.05 + .251

1.37

44.60 - 49.55

60.51

.001

BHGT

34

8.52 + .061

.34

7.80 -

9.15

30

9.19 + .069

.38

8.40 - 10.25

59.22

.001

LHC

34

82.12 + .275

1.59

77.30 - 85.40

30

84.40 + .350

1.92

79.90 - 88.35

56.34

.001

Mean

DF

P

I

&lt;o
w
I

�Table

13.

Descriptive

statistics

and t-tests

for male blue-winged

Teal
SD

Range

N

and cinnamon

teal at 7 weeks

Cinnamon
Mean + SE

of age.

t-test

Variable

N

Blue Winged
Mean + SE

Teal
SD

Range

TAR

33

39.66+

.171

.98

38.00 - 41. 70

30

40.70 + .200

1. 09

38.30 - 42.50

58.60

.001

CCEN

33

40.67 + .161

.93

37.60 - 42.15

30

43.70 + .233

1.28

41.85 - 46.80

52.54

.001

MEC

33

46.74 + .212
-

1. 22

43.30 - 48.60

30

50.36 + .177

.97

47.75 - 52.45

60.02

.001

CGAPE

33

44.80 + .205

1. 18

42.10 - 46.35

30

If 7.69

+ . 171

.93

45.50 - 50.10

59.94

.001

WIANT

33

16.86 + .060

.34

16.10 - 17.45

30

17.46 + .082

.45

16.65 - 18.30

54.09

.001

WIGAPE

33

15.55 + .051

.29

15.00 - 16.45

30

16.53 + .091

.50

15.40 - 17.50

46.22

.001

LBCEN

33

36.94

+ .238

1.37

33.60 - 39.45

30

38.85 + .256

1.40

36.05 - 41.95

60.10

.001

LEM

33

45.82 + .237

1. 36

42.75 - 48.00

30

49.03 + .221

1.21

46.60 - 51.25

60.97

.00l

BHGT

33

8.75 + .062

.36

8.15 -

9.65

30

9.50 + .080

.44

8.75 - 10.85

55.84

.00]

LHC

33

83.51 + .257

1.48

79.70 - 86.50

29

87.20 + .293

1. 58

83.60 - 89.75

57.76

.001

DF

P

I
'.D
.j&gt;

I

�Table

14.

Descriptive

statistics

and t-tests

for male blue-winged

and cinnamon

teal at 8 weeks

of ag~.

-Variable

N

Blue Winged
Mean + SE

Teal
SD

Range

N

TAR

33

39.85

+ .184

1.05

38.15 - 41. 90

30

40.98

+ .201

1. 10

CCEN

33

40.94

+ .167

.96

37.95 - 42.95

30

44.56 + .240

MEC

33

47.28

+ .202

1. 16

43.80 - 48.85

30

51.33

CGAPE

33

45.16

+ .205

1.18

42.40 - 46.95

WIANT

33

17.07

+ .061

.35

WIGAPE

33

15.72

+ .065

.37

l-tcst

Cinnamon
Mean + SE

Teal
SD

Range

DF

P

38.30 - 42.55

59.85

.001

1.31

42.90 - 47.70

52. (,7

• (lUI

+ .170

.93

49.55 - 53.90

60. ]3

· 001

30

48.51 + .177

.97

46.35 - 51.00

1)0.4 J

.O()l

16.20 - 17.65

30

17.74 + .07Lf

.41

17.00 - 18.50

57.67

.001

14.85 - 16.65

30

16.73 + .075

.41

15.95 - 17.90

58.91

.001

I

.c
VI

I

LBCEN

33

37.20

+ .220

1.26

34.00 - 39.40

30

39.72+.257

LEM

33

46.14 + .220

1.26

42.75 - 47.85

30

49.85

BHGT

33

8.93

+ .053

.31

8.30 -

9.55

LHC

33

83.83

+ .252

1.45

80.40 - 86.45

1.41

36.40 - 42.85

58.54

.00]

+ .192

1.05

47.70 - 51.90

60.52

.001

30

9.66 + .078

.42

8.90 - 10.70

52.27

.001

29

88.36

+ .315

1. 70

84.00 - 90.90

55.50

· 00 I

�Table

15.

Descriptive

statistics

and _!:_-testsfor male

blue-winged

and cinnamon

teal at 1: we eks of age.
-.

Variable

N

Blue Winged
Mean + SE

Teal
SD

Range

N

Ctnnamon
Hean + SE

Teal
SD

t-test

r

DF

Range

32

40.17 + .169

.96

38.75 - 42.45

21

41.29 ._
+ .246

1.07

39. J 5 -

CCEN

33

40 .96 + .1SLf

.88

38.60 - 42.50

22

44.85 --+ .259

1. 23

!~2.70 - ~7. 1(l

34.02

.001

MEC

33

47.55 -+ .201

1. 16

44.40 - 49.40

22

51.62+.193

.87

SO.OO -

53.60

50.13

.001

CGAPE

33

45.15 + .207

1. 19

42.00 - 46.85

22

48.74 + .229

1.05

47.00 - 'i1.0S

!~b.49

· no 1

~nANT

33

17.31 + .058

.33

16.50 - 17.85

22

17.95 + .095

.43

17.15 - 18.85

34 ..5.5

• O() I

WIGAPE

33

16.02

+ .054

.31

15.35 - 16.80 .

22

16.88 + .091

.42

16.16 - 17.55

:n.Hl

· ()()I

I. '\.

L5

:'!,

TAR

J6.

· ll() 1

,
1.0

0'
I

LBCEN

33

37.06 + .209

1.20

34.30 - 39.30

22

39.62 + .338

1.53

36.60 - 42.90

35.00

• ()() 1

LEM

31

45.85 + .235

1.31

42.80 - 47.85

22

49.86 + .276

1. 23

47.90 - 52.50

.',4.0"3

.00 I

BHGT

33

9.19 + .050

.29

8.60 -

9.80

22

10.04 + .069

.31

9 .3(~ - 1(l.()(J

\9.'&gt;8

• ()O I

LHC

33

83.99 + .282

1. 62

79.90 - 87.00

22

83.88 + .345

1.56

85.1)0 -

43.52

·()O1

lJ

1.JO

._------

�Table 16.

Descriptive statistics and t-tests for male blue-winged and cinnamon teal at 16 weeks of age.

Variable

N

TAR

32

40.32 +

.177

CCEN

32

41.00 +

MEC

32

47.58 +

Blue Winged Teal
Mean + SE
SD

Cinnamon Teal
Mean + SE
SD

t-test
DF
P

Range

N

1.01

38.80-43.15

28

41.53 +

.208

.99

39.45-43.20

48.07

.001

.168

.87

38.80-42.30

29

44.63 +

.238

1.15

43.00-47.15

45.33

.001

.221

1.09

44.40-49.40

29

51.80 +

.196

.95

49.80-54.10

50.50

Range

.001

,
CGAPE

32

45.23 +

.210

1.04

42.45-46.75

29

48.70 +

.191

.97

46.85-51.15

50.73

.001

WIANT

32

17.39 +

.068

.37

16.60-18.05

29

18.07 +

.090

.48

17.10-19.10

47.09

.004

WIGAPE

32

15.99 +

.048

.34

15.10-16.85

29

17.00 +

.081

.48

16.15-18.15

40.46

.001

LBCEN

32

37.20 +

.241

.34

34.65-39.75

29

39.70 +

.290

1.48

36.70-42.75

49.01

.001

LEM

32

46.08 +

.233

1.25

42.95-47.80

29

49.57 +

.284

1.44

49.95-52.50

48.75

.001

BHGT

32

9.22 +

.061

1.16

8.65- 9.70

29

10.12 +

.069

.40

9.45-11. 30

50.15

.001

LHC

32

84.10 +

.303

1.48

80.45-87.00

29

88.47 +

.305

1.54

84.70-91. 40

50.96

.001

I

'"

-...J
I

�Table 17.

Descriptive statistics and t-tests for male b Lue=w Lnged and cinnamon teal at 20 weeks of age.

Variable

N

Blue Winged Teal
Mean + SE
SD

Range

N

Cinnamon Teal
Mean + SE
SD

t-test
Range

DF

P

TAR

29

40,21 + .167

.90

38.70-42.45

23

41.65 + .178

.81

40.00-42.90

42.42

.001

CCEN

29

41.09 + .152

.82

38.75-42.40

24

44.54 + .254

1.16

42.75-47.05

32.18

.001

MEC

29

47.43 + .194

1.04

44.25-49.30

24

51. 70 + .249

1.10

49.55-54.30

39.23

.001

CGAPE

29

45.23 + .189

1.02

42.45-46.85

24

48.74 + .202

.90

47.45-51. 30

43.94

.001

WIANT

29

17.26 + .071

.38

16.50-17.80

24

17.96 + .096

.49

16.85-18.60

37.73

.001

WIGAPE

29

16.05 + .066

.36

15.10-16.65

24

17.02 + .093

.41

16.30-17.65

36.74

.001

LBCEN

29

37.29 + .204

1.10

35.35-39.45

24

39.64 + .299

1.35

37.05-42.35

35.57

.001

LEM

29

46.17 + .218

1.18

43.10-47.75

24

49.82 + .221

.98

48.40-52.25

45.03

.001

BHGT

29

9.27 + .057

.31

8.50- 9.85

24

10.13 + .072

.34

9.35-10.75

39.59

.001

LHC

29

84.21 + .256

1.38

80.85-86.85

24

89.16 + .287

1.40

86.45-91. 40

42.87

.001

I
1.0
00
I

�Table 18.

Variable

Descriptive

N

statistics

Blue Winged
Mean + SE

and t-tests

for male blue-winged

Teal
SD

Range

N

and cinnamon

teal at 56 weeks

Cinnamon
Mean + SE

of age.

t-test

Teal
SD

Range

DF

P

TAR

29

40.21 +

.194

1. 03

37.75-42.85

24

41.61 + .210

1.03

39.50-43.25

48.76

.001

CCEN

29

41.19 +

.175

.92

39.05-43.00

25

44.73 + .245

1. 22

43.15-47.55

44.44

.001

MEC

29

47.93 +

.204

1.14

44.95-49.80

25

52.27 + .220

1.10

49.65-54.75

50.08

.001

CGAPE

29

45.39 +

.209

1.15

42.30-46.05

25

48.77+

.211

1. 05

46.95-51. 45

50.77

.001

WIANT

29

17.31 +

.086

.45

16.50-18.15

25

18.01 + .095

.48

17.10-19.05

49.77

.001

25

16.97 + .109

.54

15.95-18.30

44.32

.001

I

WIGAPE

29

16.00 +

.077

.41

15.20-16.95

LBCEN

29

36.98 +

.193

1.12

34.40-38.95

25

39.56 + .325

1. 63

36.65-43.30

39.55

.001

LEM

29

45.99 +

.222

1.19

42.95-47.55

25

49.72

+ .240

1. 20

47.30-52.45

50.08

.001

BUGT

29

9.27

+

.058

.30

8.65- 9.90

25

10.20

+ .074

.37

9.50-10.90

46.98

.001

LHC

29

84.03

+

.292

1. 59

80.55-87.45

25

88.83

+ .307

1. 54

85.70-91.40

50.39

.001

\0
\0

I

�-100-

Although the sample size of males was small (5 of each species) there were
significant differences (P&lt;.05) between the males for 14 of the 23 variables
(Table 2). The cinnamon males had larger mean values for all variables
except the width of the culmen nail. There was overlap in the range of
all variables except the length of the center of the culmen' (CCEN), maximum
exposed culmen (MEC), width of the culmen at the nares, and the height of
the bill at the nail.
Captive

Teal

The captive teal grew rapidly from the 5th through the 8th week of age, at
which time they essentially obtained full growth (Fig. 6). No apparent
increase in size occurred after the 8th week and variations after that time
were attributed to experimental error.
To avoid excessive repetition, the
presentation of data was restricted to representative weeks after full
growth was obtained.
Included, in addition to the 5th through the 8th weeks
were measurements for the 12th, 16th, 20th and 56th weeks.
The descriptive statistics for the penned birds indicated that significant
differences in mean values existed between bluewings and cinnamons, of both
sexes, for each of the 10 variables in all age classes except the 5th week.
Even in the 5th week all but 1 variable (LBCEN) were significantly different
for both males and females (Tables 3-18). Females of the 2 species showed
greater similarity than males, while bluewings of both sexes had smaller
mean values for all 10 variables in every time period.
Overlap

in Range of Variables

The 2 species of captive females showed overlap of ranges for each variable
in every time period (Tables 3-10). Males had overlap of ranges for all
variables through the 7th week (Tables 11-13), but in the 8th week (Table 14)
there was no range overlap in the maximum.exposed
culmen (MEC), and in the
12th through 20th weeks (Tables 15-17) there was no overlap in the 3 culmen
measurements
(CCEN, MEC, CGAPE) or in the length of exposed mandible (LEM).
There was no overlap in the length of the center of the culmen (CCEN) or in
the length of exposed mandible (LEM) in the 56th week (Table 18).
The variables with no overlap in range between the species were largely
consistent between the wild and captive teal. Exceptions were (1) lengths of
the exposed mandible (LEM) and culmen to the gape (CGAPE) in males, and (2)
length of the center of the culmen (CCEN) in females.
Results

of Previous

Investigations

Data from this study generally support claims of previous investigators that
cinnamon teal have larger bills than bluewings.
There were no cases in which
the penned bluewings had a mean value for any bill variable which was larger
than the value for the cinnamons, and only the weight, length of the mandible
nail, and the width of the culmen nail were larger in wild bluewings than
in wild cinnamons.

�-101-

Spencer (1953) included measurements for the length of the lower mandible
(=LEM) and the w Ld t h of the upper mandible (=WIANT) for both males and females
but felt there was no distinctive difference between the species and that
they could not be reliably identified based on these characteristics.
Data
from this study indicated significant differences (P.2_.003)in the 2 species
for both sexes of captive birds in all time periods for each of these 2
characteristics (Tables 3-1S). The wild females showed significant differences
for both characteristics, P.2_.001 and P.2_.006,respectively (Table 1); and
the males, with a sample size of only 5 for each species, showed significance in the LEM (P.2_.013),but no significant difference in the WIANT
(P.2_.55S)(Table 2).
Dwyer (1977) recommended using the ratio of the length of the culmen to the
length of the head as a means of distinguishing the species (Fig. 2). He
suggested that if the length of the culmen is greater than the length of the
head (ratio &gt;1.0) the bird is a cinnamon and if it is less (ratio &lt;1.0) the
bird is a bluewing.
A t-test was used to examine this characteristic on the
penned females (16th we~k data) and a significant difference existed (P.2_.002).
Twenty-eight percent of the bluewings, however, had a culmen-head ratio of
~l.O and 26% of the cinnamons had a ratio of &lt;1.0. These results suggest that
only about 75% of the birds could be classified correctly based on this
characteristic.
The shape of the culmen has also been recommended as a characteristic for
distinguishing the species (Kortright 1942, Bellrose 1976, Johnsgard 1976).
They described the culmen of cinnamon teal as more constricted toward the
base, while in bluewings it is the same width throughout.
The difference
between the width of the culmen at the widest anterior point and at the gape
(Figs. 5, 6) was compared interspecifically using a t-test (16th week data)
and no significant difference was found (P.2_.4S6males-:-Pi.613 females).
This
relationship was also tested on the wild birds, and again, no significant
difference was evident (P':::_.256
females, P.2_.112males).
The length of the culmen has been the most popular characteristic used for
distinguishing the species, and several authors have documented culmen
lengths (Table 19). The measurements are similar with the exception of those
given in Spencer (1953) and the North American Bird Banding Manual (USFWS
and CWS 1977), who have cited the culmen chord length, which is longer than
the other given values.
The culmen chord length compares favorably with the
maximum exposed culmen (MEC) (Fig. 3), but the exact measurement taken is
open to interpretation.
The various other culmen lengths cited are similar
to the length of the center of the culmen (CCEN) (Fig. 2) and are assumed
to be the same variable.
These values suggest that cinnamon teal have
longer culmens (both CCEN and MEC) than bluewings and this difference was
significant for the wild teal (P&lt;.004) and the penned birds (P&lt;.005) for all
age classes in this study (Tables l-lS).
Discriminant

Analyses

Previous investigations and the descriptive statistics from this one indicated
that cinnamon teal have significantly larger bills than bluewings in nearly
all dimensions after 5 weeks of age (Tables l-lS).
It is not safe, however,

�Table 19.

Reported culmen lengths from descriptions of blue-winged and cinnamon teal.

Source

BWT Males
Culmen
Length
Range

Sclater (1912)

38.1

Spencer (1953)

46.5

45.0-47.0

45.5

45.0-46.0

50.5

50.0-51. 0

50.0

49.0-51. 0

Duvall (cited in Spencer 1953)

-

-

38.7

36.5-41. 0

-

-

43.0

41.0-46. 0

Phillips (1923)

-

39.0-42.0

-

38.0-40.0

-

42.5-48.5

-

41.4-47.0

Delacour (1956)

38.1

-

-

-

43.2

-

-

-

-

-

-

-

43.9

39.0-47.0

42.3

39.0-44.5

42.0

-

40.0

-

-

44.0-47.0

-

42.0-44.0

-

-

&lt;48.0

-

-

-

&gt;48.0

Wild

39.2

37.0-42.4

38.4

36.5-40.1

44.2

42.1-46.5

43.1

40.7-45.8

Penned

41.0

38.8-42.3

39.2

36.5-41. 4

44.6

43.0-47.1

41.9

38.5-44.4

Snyder and Lumsden (1951)
Palmer (1976)
Bird Banding Manual (1977)

BWT Females
Culmen
Length
Range

CT Males
Culmen
Range
Length

CT Females
Culmen
Length
Range

43.2

I
I-'
0
N

I

�-103-

to classify unknown birds based solely on these differences.
A discriminant
analysis programmed in SPSS (Nie et al. 1975), was used to classify females,
by species, based on the 16th week data for each variable (Table 20).
None of the variables individually classified more than 89.1% of the known
birds correctly, even though all variables showed significant differences
between the species.
The best discriminator (89.1% correct) was the height
of the bill at the base (BHGT), which has not been commonly used to distinguish these species.
The most common variable for identifying the species
is the length of the center of the culmen (CCEN), which was significantly
different (P2.OOI) and yet classified only 71.7% of the females correctly
when used as the single criterion.
The other variables, indpendently, classified only 57.7%
to 82.6% correctly (Table 20). These results indicated that
there was enough overlap in the range of the variables between the 2 species
to prevent correct classification of all birds, but that 2 or more variables
might be used to obtain an acceptable degree of accuracy.
The number of females correctly classified from a stepwise discriminant
analysis, including all 10 variables for the 16th week of age, was 96.2%
(Table 21). The F values to enter or remove variables from the analysis
(Table 22) indicated that the maximum exposed culmen (MEC) was the best
discriminator, followed by the width of the culmen at the gape (WIGAPE) and
the width of the culmen at the widest anterior point (WIANT). The length of
the center of the culmen (CCEN) was a poor discriminator with only the length
of the center of the mandible (LBCEN) and the length of the exposed mandible
(LEM) having less discriminating power.
The 3 culmen lengths (CCEN, MEC, CGAPE) were the easiest measurements to take
accurately on live birds, so 6 additive and multiplicative combinations of
these 3 variables were included in a second analysis in.addition to the 10
original variables, the ANTAREA (CCEN X WIANT), and the GAPEAREA (CCEN X
WIGAPE).
The addition of the computed variables had no effect on the number
of birds correctly classified (Table 21). The GAPEAREA became the best
discriminator at every step in the analysis (Table 22).
Using all the variables would be useful for classifying museum specimens
where time and conditions permit a thorough e~amination of the bird.
This
number of measurements would not be practical for field use when handling
large numbers of live birds, and a maximum of 3 or 4 measurements was
considered practical for field purposes (R. Hopper, pers. comm).
The 3 culmen lengths (CCEN, MEC, CGAPE) showed little overlap of ranges
between the 2 species, and as relatively easy measurements to take accurately
on live birds, it is recommended that these 3 measurements be taken on live
birds in the field. The GAPEAREA (CCEN X WIGAPE) was the best discriminantor
(Table 22) and it is recommended as a 4th variable for classifying wild teal
even though the width of the culmen at the gape (WIGAPE) was reasonably
difficult measurement to take accurately.
When the GAPEAREA was included in the discriminant analysis with the 3 culmen
lengths, the percentage of birds (16th week females) correctly classified was
93.5, only 2.7% less than when all 10 variables were included in the analysis.
It was felt that these 4 variables (CCEN, MEC, CGAPE, CAPE AREA) represented
the optimal combination if discriminating power and reliability of obtaining
accurate measurements under field conditions.

�-104-

Table 20. Classification of 16th week female blue-winged and cinnamon teal
based on single variable discriminant analysis.

Variable

Actual
Group

Classification Based on Single Variable
Predicted Group
BWT
CT
% Correctly
N
#
%
#
%
Classified

BWT

24

12

50.0

12

50.0

CT

22

7

31.8

15

68.2

BWT

24

15

62.5

9

37.5

CT

22

4

18.2

18

81.8

BWT

24

15

62.5

9

37.5

CT

22

1

4.5

21

95.5

BWT

24

15

62.5

9

37.5

CT

22

2

9.1

20

90.9

BWT

24

13

54.2

11

45.8

CT

22

5

22.7

17

77.3

BWT

24

16

66.7

8

33.3

CT

22

0

0.0

22

100.0

BWT

24

13

54.2

11

45.8

CT

22

5

22.7

17

77.3

BWT

24

16

66.7

8

33.3

CT

22

2

9.1

20

90.9

BWT

24

19

79.2

5

20.8

58.7

TAR

71.7

CCEN

MEC

78.3

CGAPE

76.1

WIANT

65.2

WIGAPE

82.6

LBCEN

65.2

78.3

LEM

89.1

BRGT
CT

22

0

0.0

22

100.0

BWT

24

17

70.8

7

29.2

CT

22

3

13.6

19

86.4

LRC

78.3

�Table 21.
Classification of 16 week old blue-winged
measured variables and 8 computed variables.

Actual
Group

Classification Based on 10 Variables
Predicted Group
% Correctly
BWT
CT
-%
Classified
N
%
1t
II

BWT

29

29

100.0

o

0.0

and cinnamon

teal based on 1) 10 measured

variables

and 2) 10

Actual
Group

Classification Based on 18 Variables
Predicted Group
BWT
___Q
% Correctly
Classified
%
%
N
II
if

BWT

29

29

100.0

o

0.0

I
I-'

o

96.2

96.2
CT

24

2

8.3

22

91.7

CT

24

2

8.3

22

91. 7

\..Jl

I

�-106-

Table 22. Values for F to enter or remcve from the analysis
from the 16th week.

Variable

F to Enter or Remove

Variable

for female data

F to'Enter

or Remove

TAR

.797l3

TAR

.77106

CCEN

.71409

CCEN

.04717

HEC

80.35929

MEC

.24992

CGAPE

2.32524

CGAPE

"k

WIANT

8.08248

WIANT

.09346

WIGAPE

14.20781

WIGAPE

LBCEN

.07732

LBCEN

.08406

LEM

.58078

LEM

1. 06936

BHGT

5.71997

BHGT

8.24020

LHC

1.l3105

LHC

.99765

J_

,~

CCEN

+ MEC

CCEN

+ CGAPE

MEC

+ CGAPE

10.85685

CCEN x MEC

.31835

"i'\

CCEN x CGAPE

,;'c

MEC x CGAPE

.27329

CCEN x WIANT

9.03993

x WIGAPE

86.17016

CCEN

";~

Contributed

Classification

so little

to the analysis

With Logarithm

they were not entered.

Transformations

of Measurements

All additive and multiplicative
combinations of the variables could be included
by logarithm transformation of the data without increasing the number of
variables.
All subsequent discriminant analyses were run using log transformations of the 4 variables.

�-107-

Table 23 presents the classification results of the penned birds based on
Although
discriminant analysis for data from the 5th,·6th and 7th weeks.
significant differences existed between the species for the variables at
those ages, the females could not be classified with an acceptable degree
of accuracy.
The males were classified reasonably well' after the 5th week
of age.
Tables 24 and 25 present the results for the weekly measurements
(CCEN, MEC,
CGAPE, GAPE AREA) from the 8th through the 56th weeks.
Ninety-two percent or
more of the females were classified correctly after the 8th week (89.6%
correct at the 8th week) and 100% of the males were classified accurately
in each time period.
The weekly classifications were based on discriminant
coefficients computed from each week's respective data (Table 26).
Aging birds to the week in the field is not practical (R. Hopper, pers. comm.) ,
so average coefficients were developed from the weekly measurements to be
used for classifying the birds in each time period (Table 26). These
coefficients were developed from each bird's average measurements of the 4
variables for the 8th through the 56th weeks data. Tables 24 and 25 present
the results of classifying the birds using the average coefficients.
Although there was a slight drop in the number of birds correctly identified,
the results were essentially the same as the weekly coefficient classifications,
demonstrating the possibility of developing coefficients to be used in classifying all wild birds older than 7 weeks of age.
All of the wild teal were classified accurately (100% correct) when the
average coefficients, developed from the penned birds, were used in the
discriminant analysis, as well as when the classification was based on
coefficients developed from the measurements of the wild birds.
CONCLUSIONS
The descriptive statistics from this study indicated that the average cinnamon teal bill is significantly larger than the average bluewing bill in most
dimensions.
These differences, however, cannot be used reliably to classify
the birds under all conditions.
Fledging or pre-fledged young are growing
rapidly through the 7th week of age and are very similar in appearance.
A
discriminant analysis using 4 variables (CCEN, MEC, CGAPE, GAPEAREA) classified 77.3%, 84.4% and 89.6% of the females correctly in the 5th, 6th and 7th
week age groups, respectively.
Aging young birds to the week is impractical
and due to the rapid growth prior to fledging (at approximately the 7th week
of age), it is recommended that only fully fledged young (8 weeks of age)
and older birds be classified in areas where both species occur.
Male
cinnamon teal acquire a red eye color at approximately 8 weeks of age (Spencer
1953) and should not present a problem of identification after that time.
The classification of the penned teal based on weekly discriminant coeffici'ents
accurately identified 92% or more of the birds in all age groups after the
8th week (89.6% at 8th week).
Essentially the same results were obtained
when the teal were classified with the calculated average discriminant
coefficients.
The male and female wild teal were classified correctly (100.0%)
b~sed on the coefficient derived from the measurements taken on them as well
as from the average coefficients from the penned teal.

�Table 23. Classification
of male and female blue-winged
ments from the 5th, 6th and 7th we eka of age.

Female
Predicted Grou12
BWT
CT

and cinnamon

teal from discriminant

-

Age
Weeks

Actual
Group

N

It

%

It

%

5

BWT

33

26

78.8

7

21.2

% Correctly
Classified

analysis

based on measure-

Male
Predicted Grou12
BWT
CT
%
%
It

% Correctly
Classified

Actual
Group

N

It

BWT

31

29

93.5

2

6.5

CT

29

6

20.7

23

79.3

m.JT

34

31

91.2

3

8.8

CT

30

1

3.3

29

96.7

77.3

6

CT

33

8

24.2

25

75.8

BWT

32

28

87.5

4

12.5

86.7

84.4
CT
BWT

7

32
32

6
28

18.8
87.5

26
4

81. 3
12.5

93.8

BWT

33

33

100.0

0

0.0

CT

30

1

3.3

29

96.7

89.6
CT

,

35

3

8.6

32

91.4

I
f-'
0
00
I

98.4

�T~b1e 24. Classification of females from discriminate analysis based on measurements
from the average measurements for the 8th, 12th, 16th, 20th, and 56th weeks of age.

Age
Weeks

Actual
Group

N

Based on Weekly Coefficients
Predicted Group
BWT
CT
% Correctly
%
%
Classified
fI
It

8

BWT

33

30

3

9.1

weeks

Actual
Group

Classification Based on Average
Predicted Group
BWT
CT
-1'-- %
%
N
fI

BWT

33

31

93.9

2

6.1

CT

34

6

17.6

28

82.4

BWT

28

26

92.9

2

7.1

CT

22

2

9.1

20

90.9

Classification

90.9

for selected

12

34

4

11.8

30

88.2

BWT

28

25

89.3

3

10.7

22

1

4.5

21

95.5

% Correctly
Classified

92.0

92.0
CT

Coefficient

88.1

89.6
CT

of age and

I
I-'

0
\.D

I

16

BWT

24

23

95.8

1

4.2

BWT

24

22

91. 7

2

8.3

CT

22

2

9.1

20

90.9

BWT

29

28

96.6

1

3.4

91. 3

95.7

20

CT

22

1

4.5

21

95.5

BWT

30

29

96.7

1

3.3

92.5

94.5

56

CT

25

2

8.0

23

92.0

CT

24

3

12.5

21

87.5

BWT

26

25

96.2

1

3.8

BWT

26

24

92.3

2

7.7

2

7.4

25

CT

27

4

14.8

23

85.2

88.7

94.3
CT

27

92.6

�Table 25. Classification of males from discriminant analysis based on measurements
from the average measurements for the 8th, 12th, 16th, 20th and 56th weeks of age.

Classification
Age
Weeks

Actual
Group

N

Based on Weekly Coefficient
Predicted Group
BWT
CT
% Correctly
-----%
%
Classified
#
If

8

BWT

33

33

100.0

0

0.0

for selected weeks of age and

Classification
Actual
Group

N

#

Based on Average
Predicted Group
BWT
CT
%
#
%

BWT

33

33

100.0

0

0.0

CT

30

0

0.0

30

100.0

BWT

33

33

100.0

0

0.0

100.0

12

CT

30

0

0.0

30

100.0

BWT

33

33

100.0

0

0.0

16

21

0

0.0

21

100.0

BWT

27

27

100.0

0

0.0

100.0
CT

21

0

0.0

21

100.0

BWT

27

27

100.0

0

0.0

CT

26

0

0.0

26

100.0

BWT

29

29

100.0

0

0.0

CT

20

0

0.0

20

100.0

B1-n

28

27

96.4

0

0.0

CT

25

1

3.6

25

100.0

100.0

20

CT

26

0

0.0

26

100.0

BWT

29

29

100.0

0

0.0

56

20

0

0.0

20

100.0

BWT

29

29

100.0

0

0.0

CT

25

0

0.0

25

100.0

I

100.0

100.0

I
f-'
f-'

0

100.0

100.0
CT

% Correctly
Classified

100.0

100.0
CT

Coefficient

98.1

�Table 26. Discriminant
selected variables.

coefficients

for male and female blue-winged

5th

6th

7th

Age
8th

CCEN

-1.12330

-1. 52485

1. 37793

MEC

1. 26050

1.10058

CGAPE

-0.50180

CCEN x ~HGAPE

1.48303

Sex and Variable

and cinnamon

teal based

on logarithms

of 4

(weeks)
12th

16th

20th

56th

Average

0.85718

1.58840

1. 62867

1. 83478

1. 39327

0.39853

-1. 04561

-0.53744

-1.46134

-1. 50137

-0.82238

-0.99182

-1. 01860

-0.23086

-0.07288

-0.64182

-0.28234

-0.12084

1.00642

-0.38338

-0.74354

1. 94036

-1. 75782

-1. 31429

01.46650

-1. 62366

-1. 82512

-1. 84465

-0.40231

Female
---

I

~
~
~

Male

I

CCEN

-0.42716

-0.46837

-0.25696

-0.45993

0.19629

-0.26700

-0.16847

-0.40863

0.32521

MEC

1. 48719

1.18044

1. 51032

1.41713

1.04596

1.42890

-0.70383

-1. 09556

1. 56043

CGAPE

-0.95610

-0.40907

-0.64524

-0.10855

-0.16809

-0.61241

-0.41397

-0.25223

-0.40837

CCEN x WIGAPE

1. 23357

1. 48052

1. 76384

1. 76458

1. 60258

2.45476

-1. 48265

-1. 02654

1.45743

�-112-

This demonstrates the possibility of developing a set of coefficients
derived from measurements from wild teal to use as an aid in identifying
fledged or older birds in the field.
Initial identification of the wild
specimens
would present a problem in areas wr.ere both species occur unless
collections were made in the spring when pair bonds would provide an assumed
accurate identification of the species.
Regional collections and the development of coefficients for those regions would minimize any potential problems
of geographical variation in the species which could exist throughout their
ranges in the western U. S.
Minor errors in measurements, of even 2-3 mm, could result in incorrectly
classifying an unknown bird, especially if the bird's measurements fell into
the range of overlap of the 2 species.
It is recoIT@ended that measurements
be taken carefully to maximize the probability of classifying the bird as
the proper species.
Some birds are too active in the hand to be measured
accurately and it is recommended these be released unbanded, or if banded,
classified as unidentified teal.
LITERATURE

CITED

American Ornithologists' Union.
1957. Check-list of North American
5th Ed. Lord Baltimore Press, Inc., Baltimore.
691 pp.

birds.

Baldwin, S. P., H. C. Oberholser, and L. G. Worley.
1931. Measurements of
birds.
Sci. Publ. of the Cleveland Mus. Nat. Hist., Cleveland.
165 pp.
Baur, R. V. 1974. Bantam chickens for hatching
Hyde, ed. Raising wild ducks in captivity.
Inc., New York.

ducks.
Pages 88-90 in D.
E. P. Dutton and Co.,

o.

Bellrose, F. C. 1976. The ducks, geese and swans of North America.
Stackpole Books, Harrisburg, Pa. 544 pp.
Bennett, L. J.
Collegiate
Delacour, J.
London.

1938. The blue-winged teal: its ecology and management.
Press, Inc., Ames, Iowa. 144 pp.

1956. Waterfowl
232 pp.

of the world. Vol. 2.

Dill, H. H., and F. B. Lee (eds.).
and Wildl. Servo 154 pp.

1970.

Country Life Limited,

Home grown honkers.

U. S. Fish

Dwyer, G. L. 1977. Competition and hostile behaviors of blue-winged and
M. A. Thesis, University of Montana,
cinnamon teal in western Montana.
Missoula.
77 pp.
Pages 96-98 in D. o. Hyde,
Ellis, J. 1974. Using an artificial incubator.
E. P. Dutton and Co., Inc. New York.
ed. Raising wild ducks captivity.
Flieg, G. M. 1974. Incubators and methods.
Pages 103-104 in D. O. Hyde,
ed. Raising wild ducks in captivity.
E. P. Dutton and Co., Inc., New
York.

�-113-

Grown, o. 1972. Modern waterfowl management
Bantam Assoc., Chicago.
359 pp.

and breeding

Johnsgard, P. A. 1975. Waterfowl of North
Bloomington and London.
575 pp.

America.

guide.

American

Indiana Univ. Press,

Kleinbaum, D. G., and L. L. Kupper.
1978. Applied regression analysis and
other multivariable methods,
Duxbury Press, Belmont, Calif.
556 pp.
Kortright, F. H. 1942. The ducks, geese and swans of North America.
Manage. Inst., Washington, D. C. 476 pp.

Wildl.

Ladd, W. N., Jr. 1969. Relationship of predation and land use practices
to duck nesting activities on Valentine National Wildlife Refuge,
Nebraska.
M. S. Thesis.
Colorado State University, Fort Collins.
116 pp.
Lee, F. B. 1974. Artificial brooding of ducklings.
Pages 129-136 in D.
Hyde, ed. Raising wild ducks in captivity.
E. P. Dutton and Co.,
Inc., New York.

o.

Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and D. H. Bent.
1975. Statistical package for the social sciences.
McGraw-Hill,
New York. 675 pp.
Palmer, R. S. 1976. Handbook of North American birds.
Press, New Haven, Conn. and London.
521 pp.

Vol. 2.

Yale Univ.

Phillips, J. C. 1923. A natural history of the ducks.
Mifflin Co., Boston.
490 pp.

Vol. 2.

Houghton

Sclater, W. L. 1912. A history of the birds of Colorado.
Co., London.
567 pp.
Sheraw, D. 1975. Successful duck and goose raising.
Pine River, Minn.
203 pp.

Witherby

Stromberg

Smart, G. 1974. Inducing difficult species to begin eating.
182 in D. O. Hyde, ed. Raising wild ducks in captivity.
and Co., Inc. New York.
Snyder, L. L., and H. G. Lumsden.
1951. Variations
Royal Ontario Mus. Zool. Occas. Paper 10:1-18.

Publ. Co.,

Pages 177D. P. Dutton

in Anas cyanoptera.

Spencer, H. E., Jr. 1953. The cinnamon teal (Anas cyanoptera
its life history, ecology and management.
M. S. Thesis.
Agricultural College, Logan.
184 pp.
Stromberg, J. 1975. A guide to better hatching.
Fort Dodge, Iowa. 100 pp.

and

Vieillot):
Utah State

Stromberg Pub1. Co.,

Strutz, C. 1974. Winter care of wild ducks in captivity.
in D. O. Hyde, ed. Raising wild ducks in captivity.
Co., Inc. New York.

Pages 186-193
E. P. Dutton and

�-114-

Strutz, C. 1974. Winter care of wild ducks in captivity.
in D. O. Hyde, ed. Raising wild ducks in captivity.
and Co., Inc. New York.
Szymczak, M. R. 1977. Waterfowl production surveys.
Fed. Aid Game Res. Rept., October.
pp. 1-14.

Pages 186-193
E. P. Dutton

Colo. Div. Wildl.,

U. S. Fish and Wildlife Service and Canadian Wildlife Service.
1977.
American bird banding manual.
Vol. 2. Bird banding techniques.
Un-numbered pages.

North

Wallace, K.I.M., and M. A. Ogilivie.
1977. Distinguishing
cinnamon teals. British Birds 70:290-294.

and

&amp;-'-L--.l__

Prepared by ---,/'--'-'&lt;./l)-=-~.::.._[_'-f--,-J_s_-_ --,-S
Robert S. Stark
Graduate Research Assistant

/
/
--)'-'j
-llzo:'/
Approved by_~,~~.~,.~~~~~~l~~j,{~·_//
__/_~~';~·
{Chard
M.
Hopper
~'j.r
~
Wildlife Researcher C
_'~~-~'~~~'~'A~

_

blue-winged

�-115-

APPENDIX

BLUE-WINGED

Specimen Number

TEAL CINNAMON TEAL REFERENCE

Weight

DATA COLLECTION

Date Collected

-----------------------

Species

_

Location

A

SHEET

------------------Age

Sex'-----------

_

----------------------------------------------------------------------

Eye Color

---------------------------

_

Wing Length

_

Length of Tarsus

Length of Foot

_

Length of Middle Toe ---------------

Culmen:

Ridge

Bill:

Center ----------Maximum

Lower Bill:

_

Width

Minimum Width ----------------

Width at Nares -----------

Posterior

Center

Width

Anterior

-------

Nail Length'---

Height of Bill:

----------------

_
Width '---------------

Nail Width

Length --------

At Base --------------------At Posterior

Comments:

_
Nail:

Width

Ridge

-----------------------

Posterior

Length of Nares

Side --------

End of Nail

-----------------Width

At Nares -----------------

��-117-

JOB PROGRESS

State of

October

1978

REPORT

COLORADO
Migratory

H-88-R-24

Proj ect No.
Work Plan No.

4

Job No.

Bird Investigations
7

--------------------------------Job Title
Effects of Land Use Changes on Mourning Doves
--~~~~~~~~~~~~~~~~~~~~------------------Period Covered:
Personnel:

January

1, 1978 to December

31, 1978

T. E. Olson, R. A. Ryder, L. Sweanor, Colorado State University;
C. E. Braun, H. D. Funk, Colorado Division of Wildlife.

ABSTRACT
Investigations concerning distribution, nesting densities, success rates, and
production of mourning doves (Zenaida macroura) in habitat types involved in
current changes in farming practices were conducted between January and
December 1978. Four study areas in northeastern Colorado ",'ereselected to
represent short-grass prairie, mid-grass prairie, winter wheat (Triticum
aestivum) and corn (Zea mays).
Shelterbelts 2nd study plots were searched
,veekly for nests from May to October 1978. Censuses were conducted for each
area every other week. A total of 194 nests was observed, including 176 in
she1terbelts, 8 in short-grass prairie, 7 in mid-grass prairie, 1 in wheat,
and 1 in corn. Overall nesting success was 45.1%.
Short-grass (50.0%) and
mid-grass (57.1%) prairie nests were more successful than were those in
shelterbelts (45.4%). Nesting density was higher for shelterbelts (40.93
nests per ha) than for any other habitat type. Chinese elm (Ulmus pumila)
was the most common tree found in 11 shelterbelts and was the species most
often used for nesting.
The nesting season was from 15 May to 7 September.
There we re no nest initiations or failures in September.
Average clutch size
was 1.94 + 0.12. Weather was the most important cause of nest failures.
Peak numbers of doves observed occurred during mid-- to late August.
Number
of doves observed has increased from 1968 to 1978 on 3 of 4 study areas,
despite increases in crop production.

�-118-

RECOMMENDATIONS

1.

Data collection concerning nesting densities,
productivity iu native prairie, winter wheat,
shelterbelts should be continued.

success rates, and
irrigated corn, and

2.

Censuses conducted every other week
be continued on all study areas.

throughout

the field season

3.

Crop statistics for the Great Plains
ascertain trends in land use.

in general

should

should

be examined

to

�-119-

EFFECTS OF LAND USE CHANGES ON MOURNING

DOVES

Thomas E. Olson

INTRODUCTION
The mourning dove is the most important and popular game bird in North
America and is widely known as an appealing songbird (Nelson 1976).
Keeler
(1977) estimated an annual harvest in North America of about 50 million
birds. About 500,000 doves are annually harvested in Colorado (Braun 1976).
A member of the Order Columbiformes, Family Columbidae, there are 2 recognized subspecies of mourning doves: Z. m. carolinensis (eastern) and Z. m.
marginella (western).
The dividing lin; between the 2 subspecies is thebreak between the eastern deciduous forests and the midwestern grasslands
(Aldrich and Duvall 1958).
It has been established that mourning doves nest not only above ground (in
trees, shrubs, on buildings, etc.), but also on the ground.
Studies have
shown that ground nesting doves make a substantial contribution to overall
production, even though nesting densities are considerably lower than above
ground (Giezentanner 1970, Strong 1971, Ryder 1972).
In recent years farming practices have undergone tremendous changes.
This
is especially true for northeastern Colorado and most of the Great Plains.
Shelterbelts have been removed, native prairie has been converted to dryland
farming and land previously dry farmed is now irrigated for row crops.
Since mourning doves nest on land that is undergoing changes in farming use,
knowledge of how alterations in farming practices affect mourning dove nesting
densities and production is needed.
No previous investigations of the effects
of land use changes on mourning doves have been undertaken in the Great
Plains region.
This study was initiated to examine the effects of such
changes on nesting densities and production.
This report presents data
collected in the initial year of study.
P. N. OBJECTIVES
The major objective
is to ascertain the effects of recent land use changes
on nesting densities and production of mourning doves to assess impacts of
such changes in the Great Plains.
Data to be collected include distribution,
nesting densities, and production of doves in various habitat types involved
in the current changes in farming practices and land use. Hypotheses which
have been developed are:
1.

Significant

changes in land use patterns

2.

Occurrence of native rangeland
doves in the Great Plains.

3.

Mourning dove nesting densities, hatching success, and fledging success
differ signficantly among shelterbelts, native rangeland, dryland
farming, and irrigated cropland habitats.

4.

Season and amount of use differ significantly

determines

have occurred

in the Great Plains.

the distribution

by vegetative

of mourning

types.

�-120-

SEGMENT OBJECTIVES
lao

Review literature concerning mourning dove nesting densities, nesting
success, and production in various habitats with particular emphasis
on the Great Plains region and northeastern Colorado. .

lb.

Examine crop statistics of ASCS, SCS, ARS, etc., and annual reports of
Colorado Agricultural Statistics to ascertain trends in land use.

2.

Select study areas representative of shelterbelts, native prairie,
dryland farming, and irrigated farming in eastern Colorado.
Particular
attention will be given to areas where established coo count or breeding
bird census routes occur.

3.

Estimate breeding densities
coo count censuses.

4.

Establish and make nest searches
habitat type.

5.

Ascertain nesting success of at least 25 nests per habitat
surveillance of known nests.

6.

Estimate productivity per nesting
surveillance of known nests.

7.

Prepare data, analyze results, and prepare progress
data using standard statistical tests.

through repeated

use of direct counts and

of 3 study plots in each major

pair per habitat

type through

type through

report.

Analyze

REVIEW OF LITERATURE
The life history of the mourning dove has been described by Moore and Pearson
(1942), Hanson and Kossack (1963), Keeler (1977), and Madson (1978).
Additional information concerning physical characteristics and distribution
has been reported by Lano (1927), Bastin (1952), and Swank (1955).
Numerous mourning dove nesting studies have been conducted.
These studies
have taken place primarily in 3 areas of the United States, the midwest
(McClure 1943, 1946, 1950; Boldt and Hendrickson 1952; Randall 1955; Carter
1957; LaPointe 1958; Hanson and Kossack 1963,K1ataske 1966; Crawford 1970;
Schroeder 1970; Faanes 1977), the south (Pearson and Moore 1939, Moore and
Pearson 1942, Lowe 1956), and the southwest (Nice 1922, 1923; Swank 1955;
Davis and Sintz 1973; Davis et a1. 1974). Fichter (1959) investigated the
breeding biology of the mourning dove in Idaho. Hon (1956), Downing (1959),
and Ryder (1972) have reported the significance of ground nesting in mourning
doves.
Early and late season nesting dates have been discussed by Gander
(1927), Angus (1934), and Hunt (1978). Bailey (1944), Weller (1959),
Nelson (1976), and Rees (1978) have noted dove nest sites in diverse and
unusual habitats.
Calculating
by Mayfield

nesting success by number of days of exposure
(1961, 1975) and Miller and Johnson (1978).

has been examined

�-121-

Mourning dove food habits have been studied by Martin et al. (1951),
Hanson and Kossack (1963), Griffing et a1. (1977), and Madson (1978).
Migration and movements have been discussed by Hoore and Pearson (1942),
Hanson and Kossack (1963), Funk (1965), Braun (1976), ana Ruble and Urban
(1977) •
Harvest data have been presented
and Keeler (1977).

by Hanson and Kossack

(1963), Braun

(1976),

Changes in land use and farming practices have been documented by Van Deusen
(1976, 1978), Carter (1977), Smith (1977), and Boldt (1978). The Colorado
Agricultural Statistics (1952, 1957, 1962, 1967, 1972, 1977) were used to
obtain information concerning changes in total acreage for dryland crops
and irrigated row crops within Colorado.
Validity of the mourning dove coo count census and factors affecting it have
been analyzed by Hanson and Kossack (1963), LaPerriere and Haugen (1972),
Keeler (1977), Baskett et al. (1978), and Sayre et al. (1978).
Sampling methods used in habitat evaluation have been examined by Grue and
Silvy (1977) and Adams and Gentle (1978). Methods used in population studies
have been reported by Lowe (1956), Jumber et al. (1957), and Eberhardt (1978).
METHODS AND MATERIALS
Study areas were selected based on the following

criteria:

1.

Similarity to habitats throughout the Great Plains (native pralrle,
dryland agriculture, irrigated cropland, and shelterbelts).

2.

Proximity to established
census routes.

3.

Accessibility.

mourning

dove coo count and breeding

Plots were randomly chosen within the selected study areas.
marked by stakes at the corners.
All plots were immediately
mourning dove coo count or breeding bird census routes.

bird

Boundaries were
adjacent to

Plots and shelterbelts were searched weekly for nests.
Active nests (eggs
or young in nest or adult present on nest) discovered were marked nearby with
red plastic tape. A number was assigned to each nesting attempt and was
written on the tape along with date of discovery.
A different number was
assigned to each nesting, rather than to each nest because of possible nest
reuse. Mourning doves were not marked and could not be individually distinguished.
Each active nest was observed weekly until it either failed or
fledged young. Although it has been stated that fledging can occur as early
as 11 days after hatching (Swank 1955, Hanson and Kossack 1963, Davis et al.
1974), nests were considered successful only if young disappeared from the
nest after their 14th day. Age of nestlings was estimated by comparing
them to photographs and descriptions of known age nestings (Hanson and
Kossack 1963).

�-122-

One change in the weekly schedule of nest searches was made for the corn
plots.
Searching entire plots became difficult after mid-July due to
height and extreme density of the plants.
Consequently, each plot was
divided into 4 subplots in late July, just as tassles of the plants emerged.
One subplot from each plot was searched completely once per' month.
The Nunn, Briggsdale, and Yuma routes were censused every other week.
The
coo count route at Yuma consisted of 20, 3 minute stops spaced 1 mile apart,
while the 2 breeding bird census routes contained 50 such stops at one-half
mile intervals.
Censuses were repeated the following week if bad weather
conditions caused postponement.
Number of doves seen, number of doves
heard, and total coos heard were recorded.
Each route was censused until
zero doves were heard on 2 consecutive censuses.
Direct counts were conducted during nest searches each week. Number of
doves observed, the number of the plot or shelterbelt, type of activity
(nesting, feeding, flying, etc.) and time spent searching were recorded
during all nest searches.
DESCRIPTION

OF STUDY AREAS

Four study areas were selected.
Within those areas, the following habitats
typical of the Great Plains were represented: native prairie, dryland crops,
irrigated row crops, and shelterbelts.
Location and dominant vegetative
type for each study area are presented (Table 1). Eighteen plots and 11
shelterbelts and tree claims (hereafter referred to as shelterbelts) in the
4 study areas were searched for nests throughout the field season (14 May
through 13 October) (Table 2).

Table 1.

Study area location and vegetative

types, northeastern

Colorado.

Location

County

Dominant

Nunn

Weld

Short-grass

Weld-Morgan

Winter wheat

(NE)

Briggsdale

(SE)

Vegetative

Type

prairie

Y~a

(So.)

Yuma

Mid-grass

prairie,

corn

Abarr

(E)

Y~a

Mid-grass

prairie,

corn

The Nunn study area, including 8 short-grass pralrle plots and 3 sheiterbelts,
is located within the Pawnee National Grassland in Weld County.
Five plots
(totaling 40 ha) were composed of short vegetation
m in height) with shrubs
virtually absent.
AIlS plots were in TlON, R66W, Section 23. These plots
had been previously established and were added because of the availability
of past nesting data (Ryder 1972).
In contrast, the 3 remaining short-grass
prairie plots (10 ha each) contained vegetation up to 0.75 m in height and
included shrubs, predominantly saltbush (Atriplex spp.). The plots were
located at TION, R66W, Section 28; T10N, R66W, Section 13; and TlON, R65W,
Section 5.

«~

�-123-

Table 2.

Vegetative

Vegetative

Type

Short-grass

prairie

type, location

and total area of selected

study plots.

Number of Plots

Total Area
in Plots (lla)

Nunn

8

70.0

Winter wheat

Briggsdale

2

40.0

Fallow

Briggsdale

2

40.0

Location

Irrigated

corn

Yuma

3

60.0

Mid-grass

prairie

Yuma

3

30.0

11

4.3

Shelterbelts

11

All Areas-

11 Including Abarr.
Dominant grasses in the short-grass pralrle plots were blue grama (Bouteloua
gracilis) and buffalo grass (Buchloe dactyloides).
Taller species (midgrasses)
such as red threeawn (Aristida longiseta), needleand thread (Stipa comata),
and sand dropseed (Sporobolus cryptandrus) were also present, but were not
abundant.
Forbs found included lambsquarter (Chenopodium album), prairie
pepperweed(Lepidium
densiflorum), and sunflower (Helianthus annus) (Costello
1944).
Sunflowers were abundant in roadside borrow pits. Other frequently
occurring species were saltbush (Atriplex spp.) andpricklypear
(Opuntia spp.).
The 3 shelterbelts selected for observation in the Nunn study area (TlON, R65\J,
Section 18; TlON, R66W, Section 12; TllN, R63W, Section 31) were 0.3, 0.2,
and 0.4 ha in size, respectively.
Dominant species were Chinese elm, pine
(Pinus spp.), and juniper (Juniperus spp.).
The Briggsdale study area is located in southeast Weld County and northwest
Morgan County.
Nest searches were conducted in 2 w Lnt.er wheat plots, 2
fallow plots, and 3 shelterbelts.
Wheat and fallow plots (T7N, R6lW, Section
26 and T6N, R60W, Section 26) were 20 ha each in size and were divided into
2 groups.
Each group contained 1 wheat and 1 fallow plot adjacent to each
other. Wheat plots were seeded in fall 1977 and harvested in mid July 1978.
Due to agronomic practices, wheat and fallow plots will be reversed in 1979.
Large areas of short-grass prairie were also 'present within th~s study area.
Species composition was similar to that described for the Nunn area. No
plots were established in short-grass prairie at Briggsdale.
The most common tree species observed in the 3 Briggsdale shelterbelts were
Chinese elm and locust (Robinia spp.). Locations were T7N, R6lW, Section 35;
T7N, R6lW, Section 36; and T7N, R60W, Section 5. Sizes were 0.2, 0.9, and
0.4 ha, respectively.

�-124-

The Yuma study area is in western Yuma County in the sandhill region of
northeastern Colorado.
Three irrigated corn plots, 3 mid-grass prairie plots,
and 3 shelterbelts were searched for nests.
Twenty ha corn plots were
irrigated by center pivot
systems.
Corn was seede4 in spring
before the start of the field season in mid May and was harvested in early
October for grain, as opposed to silage.
Corn plots were located at TIS,
R48W, Section 4 and T2S, R47W, Section 25 (2 plots established within the
latter section).
Mid-grass prairie plots were located at TIN, R48W, Section 31; T15, R47W,
Section 20; and T1S, R47W, Section 21. Like the short-grass plots, the midgrass plots were 10 ha in size. Vegetation height was greater (up to 1 m)
than that observed in short-grass prairie.
Species composition included a
higher percentage of shrubs than did the short-grass prairie plots.
Mid-grass, or mixed prairie contained a combination of short grasses, such
~s blue grama, and taller grasses, including prairie sandreed (Calamovilfa
longifolia), needleandthread, western wheatgrass (Agropyron smithii), and
sand dropseed.
Dominant forbs were lambsquarter and spiderwort (_l'radescantia
occidentalis).
Shrubs consisted mainly of sand sage (Artemisia filifolia)
and yucca (Yucca spp.) (Costello 1944, Ruangpanit 1977). As in the Nunn
study area, sunflowers were abundant in roadside b orrow pits.
Yuma study area shelterbelts (TIS, R47W, Section 20; TIS, R47W, Section 23;
TIS, R46W, Section 31) were 0.1, 0.4, and 0.1 ha in size, respectively.
Chinese elm was the dominant species.
The Abarr study area is in Yuma County, 19 mile3 south of Yuma. Two shelterbelts (both located at T2S, R47W, Section 10) were selected for observation
throughout the field season.
The dominant species was Chinese elm. Shelterbelts were 0.8 and 0.5 ha in size. No plots were established in mid-grass
prairie or irrigated corn, although both habitat types were present.
Species
composition of mid-grass pralrle at Ab~rr was similar to the description
given for the Yuma study area.
Data collected for all trees contained in the 11 shelterbelts were examined.
Chinese elm was the dominant species, comprising 57.4% of all trees searched
(Table 3). Other important species included locust, juniper, Russian olive
(Elaeagnus angustifolia), and pine (Pinus spp.). Mean height and diameter
(dbh) of the trees were 6.5 m, and 19.8 cm, respectively.
Of the 1,015 trees
searched, 873 (86.0%) were living.
RESULTS AND DISCUSSION
Habitat

Selection

During the 1978 field season, 194 nests were observed (Table 4). Of that
total, 176 (90.7%) were located in shelterbelts (175 in trees, 1 in shrubs).
Nesting attempts and productivity of ground nesting mourning doves were low.
Only 7.8% of all nesting attempts and 9.7% of the total number of young
produced occurred in short-grass and mid-grass prairie.
Nesting in corn and
wheat was insignificant, as evidenced by only 1 attempt (0.5%) in each. Downing
(1959) estimated that mourning doves in a relatively treeless area of north-

�-125-

western Oklahoma nested on the ground 71% of the time. Ground nests in
that study were responsible for 59% of all young produced and were found
primarily in areas of sand sage, grassland, and wheat stubble.

Table 3. Species composition
Colorado 1978.

of 11 shelterbelts

searched,

northeastern

Number

Percent of
Total

Chinese elm

583

57.4

Locust

118

11.6

Juniper

110

10.8

Russian olive

83

8.2

Pine

77

7.6

25

2.5

Green ash

16

1.6

Other

3

0.3

Total

1,015

100.0

Species

Cottonwood

(Populus deltoides)

The shelterbelt habitat was also predominant in nests located per hour
searched (Table 5). More than 2 tree nests (2.01) were located per hour
searched.
Nests were found at considerably lower rates in all other habitats
searched.
This suggests a preference by mourning doves to nest in trees
wherever possible.
Nesting cover in trees in northeastern Colorado is limited
and widely spaced.
Due to the scarcity of trees nests in shelterbelts are
concentrated and easily discoverd.
Nesting

Success

Overall nesting success was 45.1% (Table 4). Short-grass (50.0% successful)
and mid-grass (57.1% successful) prairie nests were more successful than were
nests in shelterbelts.
One reason could be that ground nests weren't as
susceptible to wind damage.
Differences in rates of nesting success among
short-grass prairie, mid-grass prairie and shelterbelt habitats were nonsignificant (T-test, R:0.50). It must be noted that the figures for prairie
nests were based on small sample sizes. Neither of the nests located in
wheat and corn was successful.
Downing (1959) found ground nests (29% successful) to be less successful than tree nests (49%) in northwestern Oklahoma.

�-126-

Table 4. Number and success of nests located in study plots and shelterbelts in northeastern Colorado 1978.

Habitat Type

Number
of Nests

Number
Successful

Number
Failed

Percent
Successful

Shelterbelts

176

80

96

45.4

8

4

4

50.0

7

4

3

57.1

Hinter wheat

1

0

1

0.0

Irrigated

corn

1

0

1

0.0

Abandoned

building

1

0

1

0.0

194

88

]06

Short-grass
Mid-grass

prairie

prairie

Totals

45.4

X

Table 5. Nests located per hour searched, by habitat
Colorado 1978.

Nests/Hour

Habitat

Short-grass

Searched

2.01

Shelterbe1ts

Mid-grass

type, northeastern

prairie

prairie

0.10
0.13

Winter wheat

0.03

Irrigated

0.02

cocn

Overall success rate (tree and ground nests combined) was 45.4%. That figure
is somewhat lower than nesting success reported in other studies.
Randall
(1955) reported the highest success rate of any investigation (71.1% in
North Dakota).
Nesting success reported in Iowa was 61.1% (Jumber et al.
1956), in Kansas it was 57.4% (Schroeder 1970), while in Nebraska it was 47.0%
(McClure 1946, 1950).
Only LaPointe (1958) reported a lower success rate
(26% in Nebraska).

�-127-

Nesting success also was calculated using a method introduced by Mayfield
(1961, 1975). The method is based on days of nest exposure.
For purposes
of calculation 14 days were used as the number of days in both incubation
and nestling stages. Due to small sample sizes, short-grass and mid-grass
prairie nests were combined.
In addition, success was determined for nests
in she1terbe1ts and all nests combined.
Success rates as calculated by this method were low (Table 6). Successes
for she1terbe1t nests, native prairie nests, and all nests combined were 28.2,
30.7, and 27.8%, respectively.
No other mourning dove nesting studies have
reported success rates derived by Mayfield's method.
Other investigators
have reported (traditional method) success rates from 47.0% in Nebraska
(McClure 1946) to 71.1% in North Dakota (Randall 1955).
In this study,
nests in native prairie were more successful (30.7%) than were those in
trees (28.2%).
Table 6. Comparison of nesting
traditional percentages.

success using Mayfield's

Habitat Type

Mayfield's

Method

method

and

~raditiona1

Percentage

---------------------------------------------She1terbe1ts

28.2

45.4

Native prairie

30.7

53.3

All nests combined~/

27.8

45.4

a/

- Includes 1 unsuccessful
abandoned building.

nest in each of the following:

corn, wheat,

Nesting Density
Nesting density was higher for she1terbe1ts (40.93 nests per ha) than for any
other habitat type (Table 7). Mid-grass prairie had a higher density than
did short-grass prairie.
Based on small sample sizes (1 nest located in each
type), both wheat and corn had extremely low densities (0.03 nests per ha and
0.02 nests per ha, respectively).
No nests were located in fallow.
The high density of tree nests suggests that she1teberbe1ts play a dominant
role in mourning dove production in northeastern Colorado. It would appear
that production in she1terbe1ts is far more important than production from
ground nests.
This is critical because many she1terbe1ts in the Great Plains
are being removed (Van Deusen 1978).

�-128-

Table 7.

Nesting

densities

by habitat type northeastern

Colorado

1978.

Habitat Type

Number
of Plots

Total Area
in Plots (ha)

Number
of N~sts

Density
(nests/hel)

Shelterbelts

11

4.3

176

40.93

8

70.0

8

0.11

3

30.0

7

0.23

vJinter wheat

2

40.0

I

0.03

Irrigated

3

60.0

I

0.02

2

40.0

0

0.0

Short-grass
Mid-grass

Fallow

prairie

prairie

corn

Nesting density for all ground nests (0.071 nests per hal in this study was
similar to that reported for ground nests in Oklahoma (0.062 nests per ha)
(Downing 1959). Nesting densities in shelterbelts reported in previous
studies have varied greatly.
Downing (1959) estimated a density of about
4.5 nests per ha in Oklahoma.
In contrast, Fichter (1959) found 50 nests
per ha of orchard in Idaho. Figge et al. (1953) reported an average of 194
young fledged per ha in a Colorado pear (RY.rus spp.) orchard.
Nest Sites
Trees predominated as nest sites. Of 194 nests observed, 175 (90.7%) were
located in trees. This suggests a strong preference for trees as nest sites.
As others have reported (Carter 1957, McClure 1943), mourning doves appear
to select trees as nest sites according to availability.
Chinese elm was
the species most often used for nesting.
One hundred forty-three nests
(81.8% of all tree nests) were located in that species (Table 8). Chinese
elm was also the most common tree found in 11 shelterbelts.
Moore and Pearson
(1942), Klataske (1966), and Swank (1955) have suggested that doves prefer to
nest in evergreens, especially early in the season.
Data from the 1978 field
season do not support conclusions of these authors.
Although 18.4% of all
trees searched were conifers, only 12.6% (juniper and pine with 6.3% each)
of the nests were located in them (Tables 3 and 8).
Nests were found from 1 to 11 m above the ground.
Few nests higher than 11 m
have been reported (Moore and Pearson 1942). Davis and Sintz (1973) observed
nests less than 1 m above the ground, but stated that over 90% were located
at a height of at least 1 m. Hean nest height for 175 nests observed in
northeastern Colorado was 3.4 m. This figure is intermediate between
extremes in nest height reported by Boldt and Henrickson (1952) in North
Dakota (1.8 m) and by Pearson and Moore (1939) in Alabama (7.8 m).

�-129-

Table 8. Tree species used as nest sites in 11 shelterbe1ts,
Colorado 1978.

northeastern

Species

Number of Nests

Chinese elm

143

81.8

Pine

11

6.3

Juniper

11

6.3

Russian olive

4

2.3

Locust

2

1.1

Cottonwood

2

1.1

Green ash

2

1.1

175

100.0

Total

Percent

of Total

Mourning doves apparently prefer tall, sturdy trees (McClure 1943).
Results
from 1978 support this idea. Mean height (8.2 m) and dbh (28.8 cm) of trees
used as nest sites exceeded mean height (6.5 m) and dbh (19.8 cm) of all
trees observed.
Davis and Sintz (1973) had similar findings in New Mexico.
Horizontal limbs of living trees were predominantly chosen by doves as nest
sites. Only 16 nests (9.1%) were located in forks of trees, as opposed to
159 (90.9%) on horizontal limbs. Davis and Sintz (1973) stated that 50% of
the nests they observed were in forks.
Similarly, Nice (1922) found 33% of
the dove nests she located in Oklahoma to be in forks of trees.
Seven nests
(4.0%) were found in dead trees, a low value as 14% of all trees observed
were dead. Increased cover and protection could be the reasons for the
apparent preference for living trees. Old nests of other species, primarily
mockingbird (Mimus polyglottus) were used for 7 mourning dove nesting attempts.
Mourning doves appear to require a minimum distance between nests.
Only
twice were 2 active nests observed at the same time in the same tree. Moore
and Pearson (1942) also found only 2 such occurrences in a sample size of 680
nests.
Although doves appear gregarious in some nesting situations (density
in shelterbelts was 40.93 nests per ha), this could be the result of limited
amounts of preferred nesting habitat.
Cover of ground nests varied.
Atriplex spp. and Opuntia spp. supplied little
cover for the nests observed in short-grass prairie.
Those nests were
completely exposed.
Nesting sites in mid-grass prairie contained greater
cover, supplied primarily by Artemisia filifolia and Yucca spp. The unsuccesSful nests in wheat and corn had the greatest cover.
Both were well hidden
and unexposed to the sun for much of the day.

�-130-

Nest platforms in trees were reused 27 times. Twenty-five (14.3% of all
tree nests) were reused once, while 2 others (1.1%) were reused twice.
Schroeder (1970) found that 22.1% of the nests he observed were used more
than once, including 1 nest that was reused 3 times. Ground nests were
not reused in this study.
Nesting

Season

The nesting season during 1978 was from 15 May (1st recorded nest initiation)
to 7 September (date of fledging of last nestlings).
Because field research
did not begin until mid May the start of the nesting season was not known.
Nests were initiated from 15 May to 27 August.
It is quite certain that
nests were initiated prior to 15 May due to the presence of recently used
nests at the beginning of field season.
In northern states, such as North Dakota, early May has been established
as the beginning of nesting activity (Randall 1955).
1aPointe (1958)
and Carter (1957) reported that nesting activity for both Nebraska and
Iowa started in mid April.
Most previous nesting studies have indicated
that the nesting season ends later than early September.
Only one investigation (Lowe 1956) reported an earlier close (late August for Georgia).
Moore and Pearson (1942) reported the latest end-of-season date, late
October for Alabama.
June, July and August were the most important months for dove production
(Table 9) in shelterbelts with peak activity in mid July (Table 11). These
3 months were the most important for dove production in native prairie
(Table 10), however, the peak of nesting activity for short-grass prairie
was much earlier (early June) (Table 11). No definite pattern occurred in
mid-grass prairie.
Several authors have reported peak nesting activity
occurring before mid July. McClure (1943) stated that June was the month
of greatest activity in Iowa. Similarly,
LaPointe (1958) concluded that
activity in Nebraska peaked in early June.
Results from this study are
more closely aligned to those reported in New Mexico (Davis and Sintz 1973),
Texas (Swank 1955), Idaho (Fichter 1959), and Oklahoma (Downing 1959). All
of the above authors placed the peak of nesting activity in July.

Table 9. Nesting
month, 1978.

activity

in 11 northeastern

Colorado

shelterbelts

by

Month

Nests
Initiated

Percent
of Total

Broods
Fledged

Percent
of Total

fuy

23

13.1

0

0.0

June

55

31.3

21

26.2

July

74

42.0

23

28.8

August

24

13.6

33

41.2

September

0

0.0

3

3.8

176

100.0

80

100.0

Totals
--- ---

�-131-

Table 10. Nesting activity in ll.native pralrle (short-grass and mid-grass
combined) study plots in northeastern Colorado, by month, 1978.
,

Percent
of Total

Nests
Initiated

Percent
of Total

Broods
Fledged

~y

4

26.7

0

0.0

June

4

26.7

4

50.0

July

5

33.3

2

25.0

August

2

l3.j

2

25.0

September

0

0.0

0

0.0

Totals

15

100.0

8

100.0

Month

Corn and wheat are not included in Table 11 because only 1 nest was located
in each type. The nest in wheat was active from 29 May until 4 June, while
the nest in corn was active from 28 June until 3 July.
No nests were initiated in any habitat type in September (Tables 9 and 10).
More importantly, there were no nest failures (including abandonment) in
September.
These 2 points suggest that the effect of September hunting on
nesting and dove production in Colorado is minimal.
Results from other
studies have shown hunting to have negative effects on late season nesting.
Schroeder (1970) found 37 of 48 active nests (77%) abandoned in Kansas
during the period of 1 to 11 September, 1962. Davis and Sintz (1973)
suggested that the nesting population in southern New Mexico could be
better protected by delaying the opening of hunting season until mid
September.
It would appear that no such delay is necessary in northeastern
Colorado.
Productivity

per Nesting

Pair

Average clutch size was 1.94 + 0.12. Two-egg clutches were most prevalent
(Table 12). Eighteen I-egg clutches (9.3% of all clutches) and 6, 3-egg
clutches (3.1%) were found. Lowest percentages for eggs hatched and successful nests were found in the I-egg clutches.
However, some of the I-egg
clutches might have been 2-egg clutches had they not failed after only 1
egg was laid.
This could be a contributing factor toward the apparent low success for 1egg clutches.
Hatching and successful nest percentages for 2- and 3-egg
clutches were not significantly different.
Although 50% of the 3-egg clutches
were successful, only 1 fledged all 3 nestlings.
Average clutch size appears
to be quite consistent throughout the United States.
Figures indicated for
Minnesota (1.8) by Faanes (1977) and for Kansas (1.9) by Schroeder (1970)
are comparable to the 1.94 reported for this study.

�-l32-

Table 11. Number of active nests per habitat type by week, 14 May to 9
September, northeastern Colorado, 1978.

She1terbe1ts

Short-grass
Prairie

Mid-grass
Prairie

l4-20 May

10

1

0

21-27 May

19

4

0

28 May-3 June

27

5

0

4-10 June

45

5

1

11-17 June

47

3

1

18-24 June

41

3

1

25 June-1 July

39

2

2

2-8 July

43

2

3

9-15 July

51

1

3

16-22 July

58

1

2

23-29 July

51

1

2

30 Ju1y-5 August

50

0

3

6-12 August

37

0

2

13-19 August

23

0

2

20-26 August

16

0

3

27 August-2 September

9

0

1

3-9 September

3

0

0

Week

Number of young produced per successful nesting attempt also seems to be
consistent throughout the country. During the 1978 field season, 165 young
were produced. This represents .85 young per nesting attempt and 1.88
young per successful nest. Eight previous studies reviewed by Fichter
(1959) reported a range of young produced per successful nest of 1.59 to
1.94.

�-133-

Table 12. Number and success of 1-, 2-, and 3-egg clutches
doves, northeastern Colorado, 1978.

of mourning

Clutch Size

Number

Eggs
Hatched

Percent
Hatched

Successful
Nests

Percent
Successful

Young
Produced

1

18

3

16.7

2

11.1

2

2

170

186

54.7

83

48.8

156

3

6

10

55.5

3

50.0

7

Totals

194

199

x

88
52.9

165
45.4

Productivity per nesting pair per habitat type was estimated using the
method discussed by Fichter (1959). The highest number of active nests in
any 1 week was considered the minimum number of nesting pairs.
The minimum
numbers of nesting pairs for shelterbelts, short-grass prairie, and midgrass prairie were 58, 5, and 3, respectively.
The following estimates for
productivity per nesting pair were calculated: 2.57 for shelterbelts, 1.60
for short-grass prairie, and 2.67 for mid-grass prairie.
Productivity per
nesting pair for all habitat types combined was 2.70. These results suggest
relatively low productivity per nesting pair, especially in short-grass
prairie, when compared to previous investigations.
Productivity per nesting pair has varied among studies conducted in different
states.
Number of young produced per nesting pair in previous investigations
has ranged from 2.0 in Georgia (Lowe 1956) to 6.7 in Texas (Swank 1955).
There appears to be no correlcation between number of young fledged per
nesting pair and latitude (and thus, with length of breeding season), as
evidenced by the fact that southern states represented both the highest and
lowest recorded values.
Causes of Nest Failures
Weather caused nearly one-half (45.3%) of the 106 recorded nest failures
(Table 13). Any nest that was observed at some time to contain eggs or
young, but did not fledge at least 1 nestling was considered a failure.
Of
the weather caused failures, 32 were lost due to wind, while 10 were attributed to hail storms.
Three dead adult doves were found on or near nests .
after a hail storm. All 32 nests lost due to wind were in trees. Nice
(1922) and Faanes (1977) also discussed the importance of weather.
Conversely, Fichter (1959) and Schroeder (1970) both stated that weather played
a minor role in nest destruction.
Of the remaining unsuccessful nests, 29 (27.4%) failed due to unknown causes,
26 (24.5%) were abandoned, and only 3 (2.8%) could be definitely attributed

�-134-

to predation.
Two tree nests were predated in the egg stage.
Round holes
were visible in the eggshells, suggesting an avian predator, such as a
black-billed magpie (Pica pica).
The ground nest that was predated contained
eggshells with large oblong holes in them, possibly the result of thirteenlined ground squirrel
(Spermophilus tridecemlineatus)
predation.
Faanes
(1977) in Minnesota, Klataske (1966) in Nebraska, and Boldt and Hendrickson
(1952) in North Dakota all reported substantial numbers of unsuccessful
nests due to abandonment.

Table 13. Causes of failures
northeastern Colorado, 1978.

of tree and ground nests of mourning

doves,

Cause

Number of Nest Failures

Weather

48

45.3

Unknown

29

27.4

Abandonment

26

24.5

Predation

3

2.8

106

100.0

Total

Percent

of Total

Coo Counts
Coo count censuses were used to establish patterns in cooing activity and
number of doves present throughout the 1978 season (Table 14). Because no
study plots were established adjacent to the breeding bird census route at
Abarr, it was censused only twice. Only data from the 1st 20 stops were
recorded.
Data from the other 3 routes varied greatly.
Cooing activity
peaked for both Yuma and Briggsdale in late May and early June, while cooing
at Nunn peaked in early July. The timing of peak numbers of doves seen
for the Nunn and Yuma routes (26 and 14 August, respectively) agrees with
the findings of a 1962-1963 study in Colorado (Funk 1965). His data revealed
that peak numbers of doves throughout most of the state occurred during the
1st 3 weeks of August.
The peak in total number of doves observed on the
Briggsdale route (112 seen on 29 May) represents a deviation from Funk's
results.
Results of coo counts and breeding bird censuses at Nunn, Briggsdale, Yuma,
and Abarr were obtained from the files of R. A. Ryder, L. C. McEwen, and
the Colorado Division of Wildlife for 1968-1978.
An upward trend was apparent
in 3 of the 4 routes (Table 15). There has been an apparent increase in the
number of doves at Nunn, Briggsdale, and Yuma.
There was no discernible
trend in doves observed on the breeding bird census route at Abarr.

�-135-

Table 14. Results of mourning dove call count censuses, northeastern Colorado,
starting 23 May 1978.

Date of Census

Doves
Heard

'total
Calls

Doves
Seen

Nunn

23 May
6 June
20 June
4 July
18 July
1 August
11 August
26 August
5 September
19 September
5 October

60
38
23
81
17
11
6
3
2
0
0

508
130
67
370
52
35
12
8
6
0
0

44
32
32
29
27
22
9
67
20
10
0

Briggsdale

29 May
14 June
27 June
11 July
24 July
8 August
19 August
31 August
12 September

79
77
44
68
50
5
6
0
0

423
366
182
347
187
10
11
0
0

112
81
59
65
58
49
28
25
2

Yuma

25 May
8 June
22 June
6 July
13 July
20 July
1 August
10 August
14 August
27 August
9 September
23 September

32
175
49
194
234
43
Discontinued - extreme wind
36
15
113
26
Discontinued - rain
125
36
141
36
48
17
0
0
0
0

26 May
15 June

38
27

Location
of Route

Abar~/

~/ First 20 stops only.

183
127

46
53
60
80
35
93
125
72
33
50
41
17

�Table 15.

Results of coo counts and breeding bird censuses, northeastern Colorado, 1968-1978.

Year

Nunn (BBC)
Date of
Total Doves
Seen and Heard
Count

Briggsdale (BBC)
Date of
Total Doves
Count
Seen and Heard

Yuma (CC)
Date of
Total Doves
Seen and Heard
Count

Abarr (BBC)5.:/
Date of
Total Doves
Count
Seen and Heard

1968

2 July

9

14 June

20

6 June

20

28 June

28

1969

26 June

22

22 June

62

31 May

32

28 June

22

1970

16 June

31

23 June

58

29 May

46

25 June

20

1971

15 June

42

15 June

77

9 June

61

22 June

26

1972

14 June

51

15 June

61

24 May

42

22 June

43
I
•....

1973

14 June

12

17 June

49

31 May

34

20 June

75

1974

18 June

34

13 June

30

M ayb/

58

25 June

63

1975

7 June

38

5 June

44

22 May

67

24 June

39

1976

12 June

50

10 June

63

20 May

84

22 June

62

1977

18 June

51

15 June

87

27 May

121

23 June

57

1978

11 June

69

18 June

112

20 May

116

29 June

33

1978

6 June~i

70

a/
14 June-

158

25 Ma~/

78

a/
15 June-

27

~I Conducted by this investigator.

hI Only month and year given.
~/ Data for Abarr from first 20 stops only.

w

0'\
I

�-137-

Changes

in Land Use

Hectares planted to both grain corn and winter wheat have increased in
eastern Colorado since 1950. From examination of Colora~o Agricultural
Statistics (1952 through 1977), it is apparent that the upward trend in
grain corn is most important.
Area planted to winter wheat in Weld County
has increased only slightly, while area planted to grain corn in Yuma
County has increased substantially (Table 16). Dryland wheat study plots
for this investigation are located in Weld County, while irrigated corn
plots are located in Yuma County.
Two additional counties, Baca and Phillips, were randomly selected to better
ascertain trends in agricultural land use in eastern Colorado.
Both counties
had increased numbers of hectares planted to grain corn, but only Baca
County showed an increase in winter wheat production (Table 17). Phillips
County had no apparent trend in winter wheat production.
Data from these 2
counties indicate the magnitude of the upward trend in grain corn production.
Increased use of center-pivot irrigation systems is chiefly responsible for
the change in corn production.
The rise in popularity of center pivot irrigation systems has paralleled
the increase in percent of all corn harvested that is irrigated in Yuma
county (Table 18). The percentage has increased from 0.2 in 1950 to 95.2
in 1975 (Colorado Agricultural Statistics 1952 through 1977). Without this
relatively new system of irrigation, such a tremendous shift from nonirrigated corn to irrigated probably could not have taken place.
There was an increase in area used for crop production in all 4 counties.
Additional land used for crops probably was converted from native prairie.
The conversion of land from native prairie to cropland i~ important to
mourning dove production.
During 1978 nest density was higher in native
prairie (0.15 nests/ha) than in either dryland wheat (0.03) or irrigated
corn (0.02).
Another major land use change that could have an important impact on mourning
dove production is the removal of shelterbelts.
Depression era shelterbelts
have been removed throughout the Great Plains during the 1970' s. .Reasons
that have been given for removal include decadence of the trees within the
shelterbelts, excessive water usage, and interference with center pivot
irrigation
systems (Van Deusen 1976). Northeastern Colorado has been no
exception to this practice.
Shelterbelt removal has been noted on the study
area at Yuma. Nesting density in shelterbelts (40.93 nests/ha) was much
higher than densities in prairie, corn, and wheat (Table 7).
The land conversion presently occurring in the Great Plains may be partly
beneficial to mourning doves.
Although potential nest sites have undoubtedly
been reduced by changes in farming practices, mourning doves have apparently
been able to adapt.
This is reflected by the increase in doves observed
from 1968 to 1978 on 3 of 4 study areas in northeastern Colorado.
The increase
in grain production in the Great Plains could result in more high quality
food being available to doves in the form of waste grain.
The advantage of
more food may outweigh the disadvantage of decreased nesting sites.

�-138-

Table 16. Area planted in winter wheat and grain corn, Weld and Yuma
counties, Colorado, 1950-1975.

Year

Crop

County

Number of
Hectares Planted

1950

Winter wheat

Weld

60 ,81#/

1955

Winter wheat

Weld

20 ,52~/

1960

Winter wheat

Weld

76,432

1965

Winter wheat

Weld

82,195

1970

Winter wheat

Weld

84,178

1975

Winter wheat

Weld

80,940

1950

Corn

Yuma

44 ,71gP./

1955

Corn

Yuma

23,3lgP../

1960

Grain corn

Yuma

21,344

1965

Grain corn

Yuma

10,927

1970

Grain corn

Yuma

21,044

1975

Grain corn

Yuma

54,230

~/ Statistics available only for number of hectares harvested.
~/ Statistics available only for all purpose corn.

�-139Table 17. Area planted in winter wheat and grain corn, Baca and Phillips
counties, Colorado, 1950-1975.

Year

County

Hectares Planted
in Grain Corn

1950

Baca

1 ,4l0~j

30,45(~j

1955

Baca

2 ,OO~/

48 ,ll#/

1960

Baca

1,300

103,560

1965

Baca

800

116,000

1970

Baca

15,600

139,600

1975

Baca

19,840

80,000

1950

Phillips

13,07~/

49,26#/

1955

Phillips

7,60~/

44,15#/

1960

Phillips

6,080

47,420

1965

Phillips

3,080

57,600

1970

Phillips

8,000

54,000

1975

Phillips

17,000

48,000

Hectares Planted
in Winter Wheat

~/ Statistics available only for number of hectares harvested of corn for all
purposes.

l/ Statistics available only for number of hectares harvested.
Table 18. Number of hectares of irrigated and non-irrigated corn harvested
1950 through 1975, Yuma County, Colorado.

Year

Hectares of Irrigated
Corn Harvested

Hectares of Non-Irrigated
Corn Harvested

Irrigated Corn
As % of Total

1950

104

44,096

0.2

1955

316

22,732

1.4

1960

1,628

16,812

8.8

1965

4,560

4,040

S3.0

1970

15,000

3,000

83.3

1975

47,600

2,400

95.2

�-140-

LITERATURE

CITED

Adams, G. D., and G. C. Gentle.
1978.
Spatial changes in waterfowl habitat,
1964-74 on two land types in the Manitoba Newdale Plain.
Can. Wildl.
Servo Rep. Ser. 38. 29 pp.
Aldrich, J. W., and A. J. Duvall.
of the mourning dove.
Condor

1958. Distribution
60:108-128.

Angus, H. L.

1934.

Late nesting

notes

for doves.

Bailey, A. M. 1944.
56:171-172.

The history

of a mourning

and migration

of races

Bird Lore 36:172.

dove nest.

Wilson

Bull.

Baskett, T. S., M. J. Armbruster, and M. W. Sayre.
1978.
perspectives for the mourning dove call-count system.
Am. Hildl. and Nat. Res. Conf , 43:163-180.

Biological
Trans. N.

Bastin,

Wilson

E. W.

1952.

Flight-speed

of the mourning

dove.

Bull. 64:47.

Boldt, C. E. 1978.
Stand manaeement in endangered prairie woodlands.
111. In Proc. 1977 West. For. Conf., West. For. Conserv. Assoc.,
Portland, Ore.
Boldt, W. J., and G. O. Hendrickson.
1952. Mourning dove production
North Dakota shelterbelts, 1950. J. Wild1. Manage. 16:187-191.

Pg.

in

Braun, C. E. 1976.
Trapping and banding doves.
Final Report, Fed. Aid to
Wi1d1. Rest. Proj. W-88-R, Work Plan 4, Job 3. Game Res. Rept.
Oct.
1976.
pp. 93-112.
Carter, D. L. 1957. Production of the mourning dove at Lewis,
M. S. Thesis, Iowa State Univ., Ames.
68 pp.

Iowa, 1956.

Carter, L. J. 1977 . Soil erosion: the problem
spent on it. Science 196: 409-411.

the billions

Colorado Agricultural
Statistics.
112 pp.
Agr. Bull. 1-52.

1952.

persists

despite

1950 Final report.

Colo. Dept.

1957.

1955 Final report.

Colo. Dept. Agr. Bull. 1-57.

94 pp.

1962.

1960 Final report.

Colo. Dept. Agr. Bull. 1-62.

105 pp.

1967.

1965 Final report.

Colo. Dept. Agr. Bull. 1-67.

89 pp.

1972.

1970 Final report.

Colo. Dept. Agr. Bull. 1-72.

88 pp.

1977 •

1975 Final report.

Colo. Dept. Agr. Bull. 1-77.

94 pp.

Costello, D. F. 1944.
Important species
Colorado and Wyoming.
Ecol. Monogr.

of the major
14:107-134.

forage types in

�-141-

Crawford, R. D. 1970. Mourning dove and blackbird
pine planting.
Iowa Bird Life 40:65-67.

production

in a Missouri

Davis, C. A., and G. K. Sintz.
1973. Nesting of mourning doves at San
Simon Cienega, southwestern New Mexico.
New Mexico State Univ. Agric.
Exp. Sta. Res. Rep. 265. 8 pp.
_____ , M. C. Conway, and A. H. Starkey.
1974. Nesting season of the
mourning dove in the Mesilla Valley of south-central New Mexico.
Mexico State Univ. Agric. Exp. Sta. Res. Rep. 271. 8 pp.

New

Downing, R. L. 1959. Significance of ground nesting by mourning
northwestern Oklahoma.
J. Wildl. Manage. 23:117-118.

doves in

Eberhardt, L. L. 1978.
Manage. 42:1-31.

J. Wildl.

Transect methods

for population

Faanes, C. A. 1977. Reproductive success of mourning
generating plant. Loon 49:201-204.

studies.

doves near a nuclear

Fichter, E. 1959. Mourning dove production in four Idaho orchards
some possible implications.
J. Wildl. Manage. 23:438-447.

and

Figge, H. J., G. I. Crawford, H. M. Swope, W. W. Sandfort, D. E. Owens,
and H. M. Boeker.
1953. Game bird surveys:mourning dove study.
Colorado Federal Aid Proj. W-37-R-6.
Pittman-Robertson
Quarterly,
13(3):242.
Funk, H. D. 1965. Mourning dove migration in Colorado.
Fish and Parks, Game Info. Leaflet No. 26. 2 pp.
Gander, F. F.

1927.

Nesting

of the western

mourning

Colo. Div. Game,

dove.

Auk 44:418.

Giezentanner, J. B. 1970. Avian distribution and population fluctuations
on the shortgrass prairie of north central Colorado.
M. S. Thesis,
Colorado State Univ., Fort Collins.
113 pp.
Griffing, J. P., P. E. Sawyer, and C. A. Davis.
1977. Mourning dove body
weights in southeastern New Mexico during and after a dry spring and
early summer.
Southwest. Nat. 21:545-547.
Grue, C. E., and N. J. Silvy.
1977. Mourning dove breeding habitat in
Texas-final report.
U. S. Fish and Wildl. Service, Accel. Res. Proj.
14-16-0008-954.
100 pp.
Hanson, H. C., and C. W. Kossack.
1963. The mourning
Illinois Dept. Conserve Tech. Bull. 2. 133 pp.

dove in Illinois.

Hon, W. H. 1956. The status of a ground-nesting population of eastern
mourning doves in coastal North Carolina.
M. S. Thesis.
North Carolina
State Univ., Raleigh.
65 pp.
Hunt,

L. B. 1978. Extreme nesting dates for the mourning
Illinois.
Wilson Bull. 90:458-460.

dove in central

�-142-

Jumber, J. F., H. O. Hartley, E. L. Kozicky, and A. M. Johnson. 1957. A
technique for sampling mourning dove production. J. Wildl. Manage.
21:226-229.
Keeler, J. E., Chairman. 1977. Mourning dove (Zenaida macroura). Pg. 275298. In G. C. Sanderson, ed. Management of migratory shore and upland
game birds in North America. IntI. Assoc. of Fish and Wildl. Agencies,
Washington, D.C. 358 pp.
Klataske, R. D. 1966. Mourning dove nesting success and nest site selection
in a sandhill region of Nebraska. Nebraska Bird Review 34:71-75.
Lano, A. 1927. Western mourning dove (Zenaidura macroura marginella) in
Arkansas. Auk 44:419.
LaPerriere, A. J., and A. O. Haugen. 1972. Trends of mourning dove
populations in Boone County, Iowa farm groves. Iowa State J. Sci.
46:417-424.
LaPointe, D. F. 1958. Mourning dove production in a central Nebraska
shelterbelt. J. Wildl. Manage. 22:439-440.
Lowe, J. I. 1956. Breeding density and productivity of mourning doves on
a county-wide basis in Georgia. J. Wildl. Manage. 20:428-433.
Madson, J. 1978. The mourning dove. Conserv. Dept., Winchester Group.,
Olin Corp., East Alton, Ill. 114 pp.
Martin, A. C., H. S. Zim, and A. L. Nelson. 1951. American wildlife and
plants: a guide to wildlife food habits. Dover Publications, New York.
500 pp.
Mayfield, H. F. 1961.
Bull. 73:255-261.
1975.
456-466.

Nesting success calculated from exposure.

Suggestions for calculating nest success.

Wilson

Wilson Bull. 87:

McClure, H. E. 1943. Ecology and management of the mourning dove Zenaidura
macroura (Linn.), in Cass County, Iowa. Iowa State Univ. Agric. Exp.
Sta. Res. Bull. 310:356-415.
1946.

Mourning doves in Nebraska and the west.

Auk 63:24-42.

1950. An eleven year summary of mourning dove observations in the
west. Trans. No. Amer. Wildl. Conf. 15:335-346.
Miller, H. W., and D. H. Johnson. 1978. Interpreting the results of nesting
studies. J. Wi1d1. Manage. 42:471-476.
Moore, G. C., and A. M. Pearson. 1942. The mourning dove in Alabama.
Alabama Dept. Cons., Misc. Pub1. 36 pp.

�-143-

Nelson, G. 1976. Mourning dove nesting success in artificial and natural
nests in Central Minnesota.
Proc. Iowa Acad. Sci. 83:112-115.
Nice, M. M. 1922. A study of the nesting of mourning
Auk 39:457-474.
1923.
37-58.

A study of the nesting of mourning

doves, Part 1.

doves, Part 2.

Auk 40:

Pearson, A. M., and G. C. Moore.
1939. Nesting habits of the mourning
dove in Alabama.
Trans. N. Am. Wildl. Conf. 4:468-473.
Randall, R. N. 1955. Mourning dove production
J. Wildl. Manage. 19:157-159.

in central North Dakota.

Rees, J. R. 1978. Mourning
Ornith. J. 34:15-16.

in sagebrush.

dove ground nesting

Ruangpanit, N. 1977. Primary productivity of sandhills
Dissertation.
Colorado State Univ., Fort Collins.
Ruble, P., and D. Urban.
1977. Seasonal movements
dove. Ohio Fish and Wildl. Rept. 6. 14 pp.

Colo. Field

grassland.
165 pp.

Ph. D.

of the Ohio mourning

Ryder, R. A. 1972. Avian population studies on the Pawnee Site, 19681971. Grassland Biome, U. S. IntI. BioI. Program, Tech. Rept. 171.
62 pp.
Sayre, M. W., R. D. Atkinson, T. S. Baskett, and G. H. Haas.
1978.
Reappraising factors affecting mourning dove perch cooing.
J. Wi1d1.
Manage. 42:884-889.
Schroeder, M. H. 1970. Mourn~ng dove production
planting.
J. Wi1d1. Manage. 34:344-348.
Smith, L. 1977. No she1terbe1ts,
Conserve News. 42(19):10-13.

in a Kansas osage orange

no shelter, for the Great Plains.

Strong, M. A. 1971. Avian productivity on shortgrass pra1r1e of north
central Colorado.
M. S. Thesis, Colorado State Univ., Fort Collins.
70 pp.
Swank, W. G. 1955. Nesting and production
Ecology 36:495-505.

of the mourning

dove in Texas.

Van Deusen, J. L. 1976. She1terbe1t renovation in the Great Plains. Pages
181-186.
In She1terbe1ts on the Great Plains.
Proc. Symp. Great P1ai~s
Agric. Counc. Pub1. 78. 218 pp.

�-144-

Van Deusen, J. L. 1978. Shelterbelts on the Great Plains:what's
happening? J. Forestry 76:160-161.
Weller, M. W.
24-25.

Prepared by

1959.

Mourning dove nest in cattail.

~~~~_'nv.~~=A~_t_-_.
~CJ
__~~_7__ .

_

Thomas E. Olson
Graduate Research Assistant

Approved bY_--,,~~V=-.:-'---"S:~
. ..!..~M-...=':'_
Clait E. Braun
Wildlife Researcher

__

Iowa Bird Life 29:

�October

-145JOB PROGRESS

State

0f

Project

REPORT

.:::CO:::_LO::::::_:RAD:.::::..:::::..:O::__
_
No.

Job No.

6

Job Title

Bird Investigations
~l~

_

~M~l~·g~r~a~t~o~r~y~B==i~r~d~P~u~b~l~i~c~a~t~i~o~n~s~

Covered:

Personnel:

Migratory

W-88-R-24

Work Plan No.

Period

1979

April 1, 1978 through March

_

31, 1979

C. E. Braun, R. L. Boyd, K. W. Harmon, R. M. Hopper, J. A. Jackson,
C. D. Littlefield, T. D. Ray, V. H. Reid, R. M. Stabler, F. J.
Ward, J. A. White, M. R. Szymczak.

ABSTRACT

Publications
as follows:

planned

for and accomplished

under this job for Segment

24 are

.Braun, C. E., K. H. Harmon, J. A. Jackson, and C. D. Littlefield.
1978.
Management of National Wildlife Refuges in the United States:its impacts
on birds.
Wilson Bull. 90:309-321.
Braun, C. E., and R. L. Boyd.
Nat. 24:198-200.

1979.

Albinism

in mourning

doves.

Southwest

Braun,

C. E., V. H. Reid, T. D. Ray, and R. L. Boyd.
1979.
of white-winged doves in Colorado.
Condor 81:96.

Additional

Braun,

C. E., F. J. Ward, J. A. White, and R. L. Boyd.
1979.
Plumage
aberrancies in band-tailed pigeons.
Southwest Nat. 24:In Press.

Hopper, R. M. 1978. Evaluation of pothole blasting for waterfowl
Colo. Div. of Wildl., Spec. Rept. No. 44. 21 pp.

records

in Colorado.

Stabler, R. M., and C. E. Braun.
1979. Effects of a California-derived
strain
of Trichomonas gallinae on Colorado band-tailed pigeons.
Calif. Fish
and Game 65:56-58.
Szymczak, M. R. 1978.
Steel shot use on a goose hunting
Wildl. Soc. Bull. 6(4):217-225.
White,

area in Colorado.

J. A., and C. E. Braun.
1978. Age and sex determination
band-tailed pigeons.
J. Wildl. Manage. 42:564-569.

~

Prepared

..

,-

('

?

---

by ___.!,~~"""","",:::.&lt;."'.&lt;S:e.::.)~~::I.GL""-. __;:cS!=--=:::2;h
__ ~_~ __ ~-h..::.·
Howard D. Funk
Section Chief
Small Game Research

_

of juvenile

��-147-

O-ctober 1979

INTERIM
JOB FIK~

State of

P~ORT

Colorado
----~~~~~----

Project No.

W-88-R-24
7

'l-lork
Plan No.
Job Title

Investigation

Bird Investigations

Job No.
of American

1

Coots in Colorado

---------=------------------------------------------------

Period Covered:
Personnel:

Migratory

August 1, 1976 through March 31, 1979

K. M. Giesen, H. J. Griese, W. P. Gorenzel, G. C. Miller and R.
A. Ryder, Colorado State University; C. E. Braun a.nd H. D. Funk,
Colorado Division of Wildlife

ABSTRA.CT
Distribution, status, breeding densities, reproductive success, habitat
utilization, timing of migration, and hunter harvest of coots (Fulica
americana) were investigated at 4 study areas in Colorado between August
1976 and November 1978. Coots were distributed in suitable habitats throughout Colorado, breeding in most areas and resident in low numbers along the
eastern foothills.
Coots were associated primarily with Typha and Scirpus
marshes.
Spring migration began in early March and peaked the second to
third week of April.
Nesting extended from mid April to early August.
Peak
nest initiation dates ranged from 25 April to 12 June, while peak of hatch
varied from 30 May to 4 July. A total of 354 nests was located, with nesting
and hatching success of 80.2 and 90.4%, respectively.
Average clutch size
was 8.4 and decreased as the season progressed.
Predation accounted for
64.2% of the unsuccessful nests.
Typha and Scirpus were most frequently
used for nest cover. Nest site measurements were related to local conditions
at each study area. Breeding densities ranged from 1.0 to 35.1 successful
nests per hectare of Typha and Scirpus.
Counts of juvenile coots ~ 30 days
of age were a useful index to productivity.
Movements of coots to study
areas increased in late summer, beginning in mid to late July for adults and
mid to late August for juveniles.
Timing of peak concentrations of fall
migrating coots was variable, ranging from late August to mid September in
1977, and mid September to mid October in 1978. Harvest data and hunter
responses to questionnaires indicated coots were of minor importance in
Colorado.

�-148-

RECOMMENDATIONS
Marshes managed for coots should contain primarily persistant emergent
vegetation (i.e., Scirpus acutus and/or ~
spp.) interspersed with
open water.
The cover:waterratiowithin
the emergent zone should approach
50:50.
Interspersion may be maintained by mechanical means, burning, and/or
water manipulations;
however, timing of such activities should be restricted
to the November-March
period.
Water levels of 30 cm or greater and sufficient
to flood emergents should be maintained during the spring migration and
breeding period.
Decreasing water levels which expose emergents during the
nesting period are undesirable.
The growth of aquatic food species (i.e.,
Potamogeton spp. and Chara spp.) should be encouraged, possibly through
periodic drawdown and bottom exposure.
Late summer counts of coots 30 days
of age or older should be undertaken as an index to productivity.
Research concerriing the timing of fall migration and hunting seasons should
be conducted, with the objective of selecting a hunting season coincident
with peak migration periods.
Attempts to increase hunter and manager awareness and interests in coots should be undertaken.

�-149-

TABLE OF CONTENTS
Page
LIST OF TABLES •

150

LIST OF FIGURES.

152

INTRODUCTION . .

153

ACKNO~iLEDGMENTS.•

•

153

I'

P. N. OBJECTIVE

154

METHODS •..•.

154

Distribution and Status .
Study Area Selection •.
Physical and Botanical Descriptions
Census Techniques •
Nesting Data.
Harvest . . • • •

154
155
155
156
156
157

DESCRIPTION OF STUDY AREAS
Location. • . . •
Description • • •
Water Chemistry •

157
157
157
159

RESULTS AND DISCUSSION
Distribution and Status
Spring Migration.
Nesting • • • • • • • . .
Chronology • • .
Nesting Statistics
Nest Site Characteristics .•
Breeding Densities ••
Nonbreeders . • • •
Brood Counts ••.•••
Summer Movements.
Fall Migration. •
Harvest

161
161
165
168
168
174
181
187
190
190
192
193
196

.. . .

SUMMARY •••

198

LITERATURE CITED •

200

�-150-

LIST OF TABLES

Table
1

Vegetation components in hectares of
4 study areas, Colorado, 1977-78 .

158

Results of water analyses, 4 study areas,
Colorado, June 1977 and 1978
....

160

Relationship of coot breeding distribution
and vegetation type in Colorado ..•

164

Relationship of coot distribution in fall
and winter with vegetation type in Colorado.

165

Dates of first territorial displays, peak
of spring migration, and initiation of first
nests on 4 study areas, Colorado, 1977 and 1978.

169

6

Coot production

175

7

Fate of coot nests on 4 study areas,
Colorado, 1977-78.
• .•.....

176

Fate of coot eggs on 4 study areas,
Colorado, 1977-78. . •
• ...

177

Frequency of occurrence of clutch size
for the coot in Colorado, 1977-78.

179

Number of nests and clutch size in
relation to week of nesting season,
Colorado, 1977-78 •.•..•...•

180

Number and percentage of coot nests by
cover type on 4 study areas, Colorado, 1977-78

182

Number of nests and cover type in relation
to date of nest initiation at Beebe Draw in
1977 and Ice Pond in 1977 and 1978 • . . . •

183

Number of coot nests and nest site vegetation
condition in relation to week of nest initiation,
4 study areas, Colorado, 1977-78 .
. .•.

184

Nesting success (%) by cover type, 4 study
areas, Colorado, 1977-78 • .
• ...

185

2

3

4

5

8

9

10

11

12

13

14

data, Colorado,

1977-78 .•

�-151-

LIST OF TABLES

(CONTINUED)

Table
15

16

17

18

19

20

21

22

Concealment afforded by vegetation
coot nest sites on 4 study areas,
Colorado, 1977-78 •..••••..

at
185

Number of coot nests by water depth
on 4 study areas, Colorado, 1977-78.

186

Number of coot nests by distance to
open water on 4 study areas, Colorado,
1977-78 .••.••...•••....

186

Relationship of coot nest success and
nest site cover, Lake John, Colorado, 1977

187

Breeding densities of coots as number of
successful nests per hectare of emergent
habitat on 4 study areas, Colorado, 1977-78.

188

Coot numbers observed on 4 study areas,
Colorado, mid May through June, 1977-78.

191

Dates of hatching vs. dates when young coots
were first observed on 4 study areas, Colorado

.

192

Nesting success and highest number of
young coots 30 days of age or older seen
on 4 study areas, Colorado, 1977 and 1978.

193

23

Coot harvest

196

24

Coot and duck harvest as obtained from
wing barrels and check stations on
Colorado hunting areas, 1-14 October
1977, and 30 September-13 October 1978

in Colorado,

1973-76 •••

197

�-152-

LIST OF FIGURES
Figure
1

2

3

4

5

6

7

8

9

10

Representation of areas examined for
coot presence, August 1976-November
1978. More than 1 site is included in
most specific locations ...

162

Coot distribution and status in Colorado.
Classification by latilong blocks follows
the system of Kingery and Graul (1978).
R = resident year round, B = breeding,
b = very likely breeding, bird found in
suitable habitat . • • . . . . • . . . .

163

Coot numbers observed during spring migration
on 4 study areas, Colorado, 1977. Coots first
arrived in North Park week of 4 April (C. Braun,
pers. commn.).
Counts from 14 March to 16 May
by Brown's Park NWR personnel ........••

166

Coot numbers observed during spring migration
on 4 study areas, Colorado, 1978. Numbers for
Beebe Draw include counts from 5 peripheral
~arshes.
Counts from 27 February-12 March at
Hog Lake by Brown's Park NWR personnel
.•.••

167

Number of coot nests initiated per week on
3 study areas, Colorado, 1977 •

170

··.········

Number of coot nests initiated per week on
4 study areas, Colorado, 1978.
Number of coot nests hatching per week on
4 study areas, Colorado, 1977 .

..··

Number of coot nests hatching per week on
4 study areas, Colorado, 1978.

..··

Coot numbers observed during late summer
and fall, 4 study areas, Colorado, 1977 •
Coot numbers observed during late summer
and fall, 4 study areas, Colorado, 1978.

··· · · · ·

171

·······

172

· · · · ·· ·

173

········

194

········

195

�-153-

INVESTIGATION OF AMERICAN COOTS IN COLORADO:
AND TEMPORAL RELATIONSHIPS OF THE AMERICAN

PRODUCTION, SPATIAL
COOT IN COLORADO

Warner P. Gorenzel

INTRODUCTION
The American coot (Fulica americana Gmelin) is the largest and most
aquatic member of the order Gruiformes, family Rallidae, in North America.
Two subspecies occur in the United States with the Hawaiian coot (F. a.
alae Peale) being restricted to the Hawaiian Islands.
F. a. americana
(Gmelin) is widespread in North America, breeding prima~ily on fresh water
wetlands.
Highest nest densities occur on glacial marshes in north central
United States and south central Canada.
In winter coots (hereafter used
in reference to the North American subspecies) utilize both brackish and
fresh water habitats.
Important wintering areas are the Central Valley of
California, coastal marshes of Texas and Louisiana, and wetlands of Florida
(Fredrickson 1977).
State and federal regulations list the coot as a migratory game bird.
As such, it is grouped with ducks and geese as "waterfowl" and is hunted
during waterfowl seasons.
Although coots can constitute a large percentage
of the waterfowl harvest in localized areas, hunter interest in coots nationwide is low (Fredrickson 1977).
Major studies of aspects of the biology, behavior, and ecology of coots
have centered in California, Utah, Iowa, Wisconsin, Manitoba, and Alberta.
Gullion (1952, 1953, 1954) examined territorial behavior and the reproductive
cycle in California.
In Utah, Ryder (1958) examined coot-waterfowl relationships.
Crawford (1975) studied breeding biology in relation to age; Ryan
(1978) examined behavior of breeding coots in relation to sex and age, while
Fredrickson (1967) and Sooter (1941) discussed aspects of reproduction,
ecology and management of coots in Iowa. Coot ecology and management in
Wisconsin were reported by Jahn and Hunt (1964) and Bartelt (1977). Kiel
(1955) in Manitoba and Smith (1961) in Alberta examined coot nesting and
habitat; Smith (1961) ~~so discussed censusing of coot populations.
Limited data exist concerning the status of the coot in Colorado.
General
information on nest observations, early and late occurrences, and distribution
has been summarized by Sclater (1912) and Bailey and Niedrach (1965). Little
is known about habitat preferences, timing of habitat use, productivity,
and levels of hunting mortality within Colorado.
This study was initiated
in 1976 to improve the knowledge and management of coots in Colorado.
ACKNOWLEDGMENTS
Financial assistance was provided by The Accelerated Research Program for
Migratory Shore and Upland Game Birds and the Colorado Division of Wildlife,
Federal Aid to Wildlife Restoration Project W-88-R.
The support of these
agencies is gratefully acknowledged.

�-154-

Special thanks are expressed to my major professor, Dr. R. A. Ryder,
Department of Fishery and Wildlife Biology, Colorado State University,
and my research advisor, Dr. C. E. Braun, Colorado Division of Wildlife,
for their guidance, advice, and encouragement.
Their high standards
of professionalism
were inspiring.
It was my privilege to have worked
with them.
I also thank committee member, R. G. Walter, Department
of Botany and Plant Pathology, Colorado State University, for advice
concerning aquatic aspects of the study and for critical review of the
thesis, and H. D. Funk, Colorado Division of Wildlife, for his review
and comments concerning the thesis.
I am deeply grateful to James Creasey, refuge manager, Brown's Park
National Wildlife Refuge, and his staff, Gerald Deutscher and James
Sellers, for their assistance and hospitality throughout the field
investigations.
I am grateful to the Beebe Draw Gun Club for access
to its property and the Mt. Princeton Fishing Club, especially R. M.
Stabler for access to Ice Pond.
The assistance of Colorado Division of
Wildlife personnel in obtaining harvest data pertaining to coots and for
use of equipment is acknowledged.
Sincere appreciation is due fellow
graduate students K. M. Giesen, H. J. Griese. and G. C. Miller, who
accompanied me in the field, and helped make this study so enjoyable.
Gratitude is expressed to all who returned questionnaires concerning coot
observations in Colorado.
Most of all, I am indebted
standing and support.

to my wife, Ellie, for her patience,

under-

P. N. OBJECTIVE
The major objective
was to obtain data necessary for development of a
management plan for coots breeding in and migrating through Colorado.
Data considered necessary were those relating to distribution, breeding
densities and production in the eastern plains, high mountain valleys
and west of the Continental Divide, descriptions of habitats used, timing
of spring and fall migrations, and present levels of reported harvest.
Hypotheses examined were:
(1) Distribution of coots in Colorado is
determined by the occurrence of cattail (Typha spp.) and/or bulrush
(Scirpus spp.) dominated marshes, (2) Densities of breeding coots, nesting
success and productivity differ (~ &lt; 0.05) between occupied habitats in
the eastern plains, high mountain valleys, and west of the Continental
Divide, (3) Timing of spring and fall migrations differs between areas
east of the mountains, high mountain valleys, and west of the Continental
Divide, (4) Hunting in Colorado is the most important source of mortality
for locally produced coots.
METHODS
Distribution
Two methods
Colorado:

and Status
were used to investigate

coot distribution

and status in

�-155-

1.

Observation of wetlands encountered
study areas throughout the state.

while traveling

between

2.

Questionn~ires sent to field personnel of the Colorado Division
of Wildlife and amateur ornithologists.

Date or period of observation, dominant emergent vegetation, geographic
location, presence or absence of coots, and coot status were recorded
for each wetland visited.
Vegetation was classified as either cattail
and/or bulrush, or other.
Omitted were roadside ditches and wetlands
completely filled with cattail and/or bulrush, with no open water.
"Other" included areas with sedges (Carex spp.), willows (Salix spp.),
or grasses (Family Gramineae) as the dominant'emergents,
or areas
without any emergents, such as many lakes, reservoirs, and stockponds.
Geographic location classifications were eastern plains, high mountain
valleys, and west of the Continental Divide.
High mountain valleys
refer specifically to North, Middle, and South Parks, and the San Luis
Valley.
Coots were considered to be breeding if territorial behavior
(as defined by Gullion 1952) and nests and/or young coots were observed
during 15 April through 15 August, and nonbreeding if gregarious and
nonterritorial.
Coots observed from 16 August through 14 April were
classed as migrating or wintering.
Responses from questionnaires, observations of wetlands, and results
from the Colorado Bird Distribution Latilong Study (Kingery and Graul
1978) were combined to plot coot distribution.
Coot status (i.e.,
breeding, resident, migrant only, etc.) was determined for each area.

Study Area Selection
Study areas were selected to represent the major geographic and coot
breeding areas in Colorado.
Criteria for selection included:
(1)
Sufficient area and cover to provide adequate numbers (25 pairs) of
breeding coots, (2) Similarity to other wetlands in the region, (3)
Stable water levels, (4) Accessibility,
(5) Freedom from excessive
external interference.

Physical

and Botanical

Descriotions

Each study area was mapped using a staff compass and tape. Emergent
vegetation was plotted by pacing and with the aid of a hand held compass
and field range finder. A planimeter was used to measure size of total
wetland and emergent areas. Aquatic vegetation was sampled along
transects using a method modified from the Atlantic Waterfowl Council
(1972). Transects were established 100 and 200 m apart depending upon
size of the area with sampling intervals 35 to 100 m apart.
A rake
was used to collect submerged vegetation.
Vegetation density ratings
(dense, moderate, sparse) were based on amounts recovered.
Scientific

�-156-

plant names follow Harrington (1964). At each sampling station, water
depth and bottom consistency were recorded.
Marked poles were placed
on all study areas to record water level fluctuations.
Water samples
were analyzed for free and total acidity, phenolphthalein and total
alkalinity, hardness, and pH using a Hach Chemical Company Water
Ecology Kit, Model AL-36B.
Soil survey information of adjacent uplands,
obtained from the Soil Conservation Service, and samples from along
aquatic transects were used to identify marsh substrate types.

Census Techniques
Counts of coots on all study areas were conducted weekly or bi-weekly
with a 20X spotting scope and 8 X 40 binoculars.
Counts were standardized by counting from the same location on each area at approximately
the same time of day for a given area. Weather conditions and other
factors were recorded on each visit.

Nesting

Data

Nest searches were systematically made at approximately weekly intervals
on all study areas by wading through emergent vegetation or from a boat.
A nest was considered to be any platform with 1 or more eggs. In those
instances when a nest was discovered after hatching, the platform was
considered a nest if shell fragments were present.
Each nest was
numbered and marked with a small cardboard tag. In addition, yellow
surveyor's tape was attached to vegetation approximately 0.6 to 12 m
away in -such a manner as to be conspicuous to the observer.
Each tape
was marked to indicate nest number and direction, and distance to the
nest.
Once found, nests were revisited every 7-11 days until hatching.
Type of cover vegetation, its condition, i.e., dead or alive, height
of a representative sample of the vegetation, nest concealment, i.e.,
was the nest concealed on 1, 2, 3, or 4 sides, and/or from above, depth
of water, and distance to open water were recorded for each nest site.
Height of the eggs above water, number of eggs and their condition, i.e.,
cold, warm, or pipped, were recorded on each nest visit.
Initiation dates of nests found during laying were calculated by backdating, assuming 1 egg laid per day. For nests found after laying,
initiation dates were based on hatch dates, assuming a 23 day incubation
period and 1 egg laid per day. Hatch was defined as the time at which
all eggs had hatched.
A successful nest was one in which 1 or more eggs
hatched.
Average clutch size and hatching success calculations were
based on data from successful nests only. Statistical tests follow Sokal
and Rohlf (1969).

�-157-

Harvest
To measure hunting pressure and coot harvest levels, volunteer wing
collection barrels with appropriate signs were placed at waterfowl
hunting areas in Colorado (Hoffman and Braun 1975). Published and
unpublished harvest data were examined.
A questionnaire was designed
to investigate hunter attitudes.
Students from Colorado State
University, Wildlife Techniques class were assigned to check stations
to question hunters leaving the Wellington State Wildlife Area on the
opening weekend of the waterfowl season in 1976 and 1977.

Description

of Study Areas

Location
Study areas were located at Beebe Draw (T4N, R65W, S34) near LaSalle,
Weld County, on the eastern plains; Lake John (T9N, R81W, S2), near
Walden, Jackson County, in North Park; Ice Pond (T13S, R78W, S5, 6, 7,
8), near Buena Vista, Chaffee County, in the central mountains; and
Hog Lake (T10N, R103W, S9), part of Brown's Park National Wildlife
Refuge (NWR), Moffat County, in extreme northwestern Colorado.
Description
Beebe Draw was a deep emergent marsh at an elevation of 1,446 m. Cattail
was the dominant emergent (Table 1). Maximum water depth was 2.3 m
and fluctuated approximately
122 cm between April 1977 to November 1978.
The substrate consisted of sand and loamy sand.
In March and April 1977, and April 1978, segments of the emergent zone
(3.1 and 0.8 ha respectively) were burned.
Burning did not kill any
emergents, but did clear away litter from previous years growth.
In
January 1978, water levels were drawn down in preparation for dragline
operations.
This resulted in exposed emergents and mudflats but did not
completely drain the area. A large number of carp (Cyprinus carpio)
either died or were exposed to predation.
Dragline operations commenced
during the first week and continued through the last week of April. .Low
water levels were maintained until the second week of May. Thereafter,
water levels were allowed to increase, flooding all previously exposed
mudflats and some emergents by the end of June. A bulldozer was used in
late June to flatten high spoil banks left by the dragline.
Approximately 4.2 ha, or 58.3% of the emergents were removed by the
dragline, covered by spoil banks, or cut off from any water source.
The
total area of open water and emergents was reduced by 21.0% from 16.7
to 13.2 ha. In 1977, only 2 aquatic species were sampled at 20 stations.
Coontail (Ceratophyllum demersum) was found at 16 stations, and was rated
moderate to dense at 10. Sago pondweed (Potamogeton pectinatus) was found
at 9 stations and was rated sparse at each.
In 1978, 4 aquatic species
were sampled at 22 stations.
Coontail decreased in abundance, being
found at only 6 stations and was rated moderate to dense at 3. Sago

�Table

1.

Vegetation

components
Beebe

in hectares

spp.

.
S clrpus
spp.

2

Scirpus
alnericanus

areas,

Lake

Draw

Colorado,

1977-78.

John

Ice Pond

Hog Lake

1978

1977

1978

1977

1978

1977

1978

_
1
3.4::&gt;(47.9)

1.52(50.5)

0.01(0.4)

0.01(0.4)

0.40(44.4)

0.40(44.4)

0.04(1,2)

0.04(1.3)

1. 03( 14.3)

0.49(16.3)

2.76(99.6)

2.76(99.6)

0.34(37.8)

0.34(37.8)

2.06(61.5)

1.81(58.4}

2.26(31.4)

0.78(25.9}

O(O}

0(0)

0(0)

0(0)

0(0)

0(0)

1977
Typha

of 4 study

I
I-'
l/1

3

0:&gt;

0.22(7.3)

Total area of
en1ergents

7.20

3.01

2.77

Total area
open H20 &amp;
emergents

16.7

13.2

18. 1

lparentheses
2Hardstem

indicate
bulrush

313eebe Draw:
Hog Lake:
Distichlis

percentage

(Scirpus

0(0)

0.16(17.8)

0.16(17.8)

1.25(37.3)

1, 25( 40.3)

2.77

0.90

0.90

3.35

3.10

18. 1

14.4

14.4

0(0)

0.46(6.4)

Other

of total

24.4

emergents.

a cutu s ) and s oIt s t ern bulrush

mixed Scirpus
and Typha
stricta,
Phragmit~1munis.

24.4

sppr : Ice Pond:

(~

va l ldu s },

Eleocharis

macrostachya,

Jllnclls

balticus,

Carex'
---

spp.:

I

�-159-

pondweed increased, being found at 10 stations, and rated moderate to
dense at 6. Horned pondweed (Zannichellia palustris), fo.lnd at 3 stations,
represented a minor component of the aquatic vegetation.
Chara spp.,
fresh water algae and important coot foods (Jones 1940), were not
recorded in 1977. In 1978, Chara was found at 11 or 50% of the sampling
stations and was rated moderate to dense at 9. Chi-square analysis
indicated a relationship between abundance rating of Chara and site
condition (P &lt; 0.05).
Of the 9 stations with a high rating, 7 represented
sites disturbed by dragline operations and/or exposed during drawdown.
The Lake John study area, locally known as Lake John Annex, included a
shallow bulrush dominated marsh of 18.1 ha, and a broad expanse of open
water, totaling approximately 100 ha at an elevation of 2,500 m. The
dominant aquatic was Chara spp., found at 24 of 45 stations and rated
moderate to dense at each. Other important aquatics included sago pondweed and water milfoil (Myriophyllum exalbescens).
Maximum water depth
and fluctuation were 1.4 m and 56 cm, respectively.
The substrate was
composed of sandy loam and silty clay loam.
Ice Pond at 2,425 m elevation, was 14.4 ha in area. Emergents consisted
of nearly equal proportions of cattail and bulrush, with lesser amounts
of spikerush (Eleocharis macrostachya), rushes (Juncus balticus), and
sedges (Carex spp.).
Interspersion between emergents and pools of open
water was well developed.
Aquatic growth was the most abundant of all
study areas; species present were water milfoil, smartweed (Polygonum
amphibium), Potamogeton filaformis, and Ranunculus aquatilis.
Each
received moderate to dense ratings.
Maximum water depth and fluctuation
were 1.5 m and 11 cm, respectively.
The substrate consisted of stony
sandy loam.
Hog Lake at 1,634 m elevation, was a shallow bulrush dominated marsh,
characterized by a well developed interspersion of emergents and open
water.
Common reed (Phragmites communis) and salt grass (Distichlis
stricta) were also present, growing as emergents.
Water milfoil,
sago pondweed, and Potamogeton vaginatus were common aquatics.
Maximum
water depth and fluctuation were 1.6 m and 33 cm, respectively.
No
soil survey exists for the Hog Lake area; aquatic transects indicated a
silt and clay substrate.
During the winter of 1977-78, muskrat (Ondatra
zibethica) activity resulted in a decrease of 0.25 ha of hardstem bulrush.

Water Chemistry
Water at all study areas was basic (pH&gt; 7.0( (Table 2). Factors
influencing chemical characteristics of water include source, organic
matter content, and amount of photosynthetic activity.
In Colorado, most
water sources are basic.
Sources of water for study areas, besides rainfall, were:
1) Beebe Draw - irrigation water and seepage, 2) Lake John water diverted from the North Fork of the North Platte River, 3) Ice Pond natural springs, and 4) Hog Lake - seepage and water pumped from the Green
River.

�Table

2. Results

Area

of water

analyses,

4 study areas,

Colorado,

June

1977 and 1978.

1

Free
acidity

Total
acidity

Phenolphthalien
alkalinity

Total
alkalinity

Hardness

pH

Beebe

Draw

0

11-17

0

137-308

325.,.410

9.3-9.6

Lake

John

0

0

0-51

120-137

513-564

10.0

Ice Pond

0

0-11

0-34

103-120

103

8.9 -9.7

Hog Lake

0

0

34

239 -274

599-667

9. 8 -9.9

1

Except

for pH, all results

expressed

as mg of CaC0 /liter
3

of H 0.
2

I
f-'
C1'

o
I

�-161-

RESULTS AND DISCUSSION
Distribution

and Status

Two hundred and thirty wetlands were investigated for coot presence
from August 1976 through November 1978 (Figure 1). Notably underrepresented were the far eastern plains, especially along the South
Platte and Arkansas rivers.
Data obtained from 108 returned
questionnaires, field investigations, and published records were compiled
to determine coot distribution and status according to the latilong
system (Figure 2).
Major breeding areas were Brown's
Coots bred throughout most of Colorado.
Park on the western slope, and North Park and the San Luis Valley in the
high mountain valleys.
On the plains, coots were locally abundant where
Sclater (1912) and Bailey and Niedrach (1965)
suitable habitat occurred.
listed the coot as common in summer, breeding mainly on the northeastern
plains and occasionally in mountain parks up to 3,000 m. Sclater (1912)
indicated coots were not abundant on the western slope. Cooke (1897)
reported coots breeding in Middle and South Parks, and Keyser (1902)
observed coots with young near Buena Vista, probably at Ice Pond. Breeding
coots were not observed in Middle or South Parks during this study.
Nesting coots were primarily associated with cattail and bulrush marshes
(Table 3). A Chi-square test of independence between the presence or
absence of breeding coots and the occurrence of cattail and/or bulrush
dominated marshes indicated a strong relationship (~ &lt; 0.001) in each
region and statewide.
A similar analysis for the migration-wintering
period (Table 4) also indicated a strong relationship (~ &lt; 0.001) statewide.
The hypothesis that distribution was related to the occurrence
of cattail and/or bulrush
marshes was accepted.
Not all cattail or
bulrush marshes were used by breeding, migrating, or wintering coots, but
they were used in strong preference to other vegetation types. Other
vegetation types used for nesting included tamarix (Tamarix gallica),
spikerush, willows, and sedges.
Sedges growing as emergents were characteristic of many high altitude wetlands utilized by coots, especially
wetlands in North Park.
Coots were resident in low numbers « 1,000) mainly along the western
boundary of the plains, from near Fort Collins south to Pueblo.
The
presence of wintering coots was dependent on the severity of winter
weather.
Coots, present near Fort Collins during the winter of 1976-77,
were absent during the 1977-78 winter when water areas froze.
In the
Fort Collins area, coots associated with wintering waterfowl and were
observed feeding on feces and waterfowl carcasses.
Bailey and Niedrach
(1965) listed the coot as uncommon in winter, and along with Sclater
(1912) mentioned a few wintering at Barr Lake northeast of Denver.
Christmas bird counts sponsored by the National Audubon Society (19601977) and listed in American Birds have consistently noted coots along
the Front Range from Fort Collins to Pueblo.

�COLORADO
SEDGWiCK

lOGAN

WEL.D

MOFFAT

•

PHILLIPS

o
RIO BLANCO

•

•

MORGAN

WASHINGTOI'4 YUMA

ADAMS

•

ARAPAHOE
GARf"IElD

L.INCOL.N I KIT CARSON

ELBERT

MESA

I
I-'

•

CHEYENNE

I

\CROWLEY I

J

,

KIOWA

() • ,...".,..".,..,.,=-:
BENT
SAN MIGUEL

PROy/ERS

•

DOL.ORES
LAS

MONTEZUMA

ANIMAS

BACA

o

o
LA PLATA

Figure 1.
November

Representation
of areas exami.ned for coot presence,
1978.
More than onc site is included in most spccific

August 1976 locations.

'"

N
I

�r'-'-'-'l-'-'
I
gB

°

I

B

,

~

I

~

'-i .-..

_._._._.-.

.~
-, B

._. '--'_'"1

.-~

R

!

R

I

i

~

I

.,.. -.

°

. _.,
...

I~

i,

B

.~~~~~~~~

I
,

I

!-s-J
I
!.."' ,_

V

NiP

-R L-~-~-I

.".:;

rr

I

11..
r-Il::-~ ~ yt'- L._._. . .' . _.1. _.l~ ._.jl._._.i_L._._. _l._. _._.lL._. _.J
I

...,

b

8

F'i gu r c 2.
Coot d is t r ibut i on and status
in Colorado.
Classification
by latilong
blocks
follows
the system
of Ki n ge r y and Graul (1978).
R -= resident
year
round,
13:-: breeding,
b:: very likely bn'ccllng,
bird found in suitable
habitat.

�Table 3.

Relationship

of coot breeding

distribution

and vegetation

type in Colorado.

Nurnb er of Areas

Area

Dominated

by Typha or Scirpus

Dominated

With
breeding
coots

None
or
nonbreeders

Total

With
breeding
coots

None
or
nonbreeders

Total

45

31

76

15

136

151

63.13

&lt;0.001

17

10

27

1

58

59

42.02

&lt;0.001

by other vegetation

2
X (df=l)

Probability

Statewide
Eastern

plains

I
f-'
0-

High mountain
valleys

8

1

9

13

35

48

12.44

&lt;0.001

West of Continental Divide

20

20

40

1

43

44

25.45

&lt;0.001

.~

I

�-165-

Table 4. Relationship of coot distribution
vegetation type in Colorado.

Number of Areas
._._--_._-----_._.--_._----_._----,-_
.._----_--_- ..
Dominated by
Dominated hy
Typha or Scirpus
other vegetation

Status
Coots present
Coots not present
To t a Ls

29.40, df

in fall and winter with

40
24
64

26
92
ll8

1, P &lt; 0.001

Spring Migration
Weekly counts were initiated in 1977 to document migration at only 2
areas, Beebe Draw and Lake John (Figure 3). Ice Pond and Hog Lake were
not visited until after the peak of migration.
However, counts by Brown's
Park National Wildlife Refuge personnel over the 14 March to 22 May 1977
period were used to indicate coot migration at Hog Lake.
In 1978, regular
visits to all areas began by mid March (Figure 4).
At Beebe Draw and Hog Lake coots first arrived in late February or early
March. Peak numbers were recorded in 1977 and 1978 during the second
week of April.
In 1978, low water conditions at Beebe Draw influenced
coot activities.
Numbers recorded ranged from 7-14 during the peak
migration month of April with no discernible trend. To identify the
peak of migration, counts included coots observed on 5 peripheral marshes.
Observations indicated no substantial increase in coot use on adjacent
marshes, suggesting that coots by-passed the area.
At Lake John coots first arrived in 1977 and 1978 during the first week
of April. Arrival date was dependent on ice thaw. In both years,
numbers increased rapidly for 2 weeks, then decreased to resident levels
within 3-4 weeks.
Although coots arrived at Ice Pond earlier than Lake
John, peak numbers were also recorded the third week of April.
The
decrease of coot numbers from peak to resident levels in 1977 and 1978
was consistent with other ireas.
Timing of migration and relative numbers observed on each area were
consistent between years.
The general pattern statewide was arrival in
late February or early March (first week of April at Lake John),
increasing to peak numbers during the second to third week of April,
then decreasing to resident levels by the second to fourth week of May.
Timing of peak migration was similar for the eastern plains and west of
the Continental Divide.
The peak occurred I week later in the high
mountain valleys.
The literature suggests a similar pattern of coot
migration in Colorado.
Cooke (1897) reported coots reached northern
Colorado the last part of March; Sclater (1912) reported that coots
arrived the second half of March; while Niedrach and Rockwell (1939)
observed first arrivals in early March, with most coots returning the
first week of April.

�-166-

900

,1\

,

I ,

aoo
fi)

,

0 BEEBE DRAW

i \

LAKE JOHN·
0----",
0-- ....•ICE POND

700

\

i ~
,.
••
:

e··· -_.• HOG LAKE

,.,

\,

,
,

\

'..1

600

r··

5
0
0

I

••

500

lL.

0
CD

I
I

I

\

".

.••...~

•

:

z

\

.

\

I•

W

::&gt;

'

.~
, ,

0:

:E

I
I

I

,

300

•,

••

\

""\

\
\

\

200

.~

.

\

\ \

_.I

\0--"'"
\

\
100

\

,
'0-":

~
FE8.

\..7__

1_4__

2_1__

MARCH

2_8J

,-4__

1_1__

'_8__

APRIL

2_5-,

2

9

16

23

MAY

WEEK BEGINNING
Figure 3. Coot numbers observed during spring migration on 4 study a r e as ,
Colorado,
1977. Coots fi r s t arrived in North Park week of 4 April (C. Braun,
pers.
c orrrrn; }, Counts from 14 March -16 May at Hog Lake by Brown's NWR
personnel.

�-167-

900

BEEBE DRAW

800

G-----i)

LAKE JOHN

•••••••

HOG LAKE

ICE POND

700

1·'-,
, .,

•

I.•,

600

.''.\.

I

'.\.,

I

\

,•

~

0
0

••

0 500
IL
0

.,.,

I

Ct:
lLI
CD 400

2
:&gt;

z

300

200

100

si
FEB.

15

8

MARCH

APRIL

22

MAY

WEEK BEGINNING

Figure 4. Coot numbers observed during spring migration on 4 study areas,
Colorado,
1978. Numbers for Beebe Draw include counts from 5 peripheral
marshes.
Counts from 27 February
- 12 March at Hog Lake by Brown's
Park NWR pe rsonnel.

�-168-

Nesting
Chronology
Breeding coots defend a territory which serves as a mating, nesting,
feeding, and brood rearing area (Gullion 1953, Ryder 1958). Behaviors
associated with territories include patrolling, charging, splattering,
fighting, and paired display (Gullion 1952). Gullion (1953) reported
that coots in California displayed year round if resident on the
breeding area. On the Colorado study areas, no coots were resident
year round, therefore such displays were used to indicate the onset of
breeding.
The dates of first territorial displays were recorded (Table 5).
Territorial displays were observed at Beebe Draw, Ice Pond, and Hog Lake
1-3 weeks before the peak of migration in 1977 and 1978. At Lake John
displays were not observed until the peak or 1 week after the peak of
migration in 1977 and 1978. The range of time between first territorial
displays and the appearance of first nests suggested a shortened territory
establishment-to-nest
initiation cycle in the high mountain valleys.
At
Beebe Draw and Hog Lake the range was 4-5 weeks, but was only 1.5-3 weeks
at Lake John and 2.5 weeks at Ice Pond.
Nest initiation ranged from mid April to mid July (Figs,S
and 6).
The peak, defined as the largest number of nests initiated during a week,
varied between areas and years. Peak nest initiation in 1977 for Beebe
Draw, Lake John, and Ice Pond were the weeks beginning 25 April, 6 June,
and 9 May, respectively.
Hog Lake was selected as a study area too late
in 1977 to make a complete determination of initiation dates.
In 1978,
peak nest initiation occurred the weeks of 29 May and 12 June for Beebe
Draw, 29 May for Lake John, 15 and 22 May for Ice Pond, and 8 May for Hog
Lake.
Habitat disturbances affected nest initiation at Beebe Draw.
In March
and April 1977, 3.1 ha of emergents were burned.
Six nests in burned areas
were initated somewhat later than most other nests, possibly due to a lack
of cover or support 'before new vegetative growth had sufficiently developed.
Fredrickson (1970) and Weller
and Spatcher (1965) noted dElayed nesting
until new vegetation
was sufficiently high to provide a substrate for
attaching nests.
In 1978, low water conditions at Beebe Draw delayed nest
initiation by 6 weeks compared to 1977. Smith (1961) indicated coots will
not nest until water levels flood the emergents.
Dates of hatch ranged from mid May through early August (Figs. 7 and 8).
Peak hatching dates in 1977 were the weeks beginning 30 May at Beebe Draw,
4 July at Lake John, and 30 May and 6 June at Ice Pond. Peak of hatch
could not be determined for Hog Lake due to the late date when nest searches
were initiated.
In 1978, peak dates for Beebe Draw, Lake John, Ice Pond,
and Hog Lake, were the weeks of 26 June, 3 July, 19 June, and 12 June,
respectively.

�Table 5. Dates of first territorial
displays,
4 study areas, Colorado, 1977 and 1978.
First Territorial

Display

Area

1977

1978

Beebe
Draw

23 March

4 April

Lake
John

23 April

24 April

Ice
Pond

2

Hog
Lake

2

and initiation

Week of Peak Migration

of first nests

Week of First

on

Nests

1977

1978

1977

1978

11 April

. 1
10 Apr Il

18 April

8 May

18 April

17 April

2 May

15 May

2

1 April

1Based on observations
2Area visited

1

peak of spring migration,

19 March

11 April

from peripheral

marshes.

too late to make determination.

17 April

10 April

25 April
2

1

17 April

17 April

I
i-'
0\
\0
I

�20

R

18

I'

16

Z

a:

10

W
CO
~

:J
Z

ICE POND

,

i

,,

I
I

4

"I

I
II

18
APRIL

"

25

2

~~\

/

9

16

I

\,,

'

'.

a

\

" X" "~

2

I
I-'
._...

\,

" r-:

6

LAKE JOHN

\

,r

8

BEEBE DRAW

\,,
\,,

,r'''
,

0

....

\

"

f3 14
12

\

/

~

U.

._

/\

~
23

MAY

30

6

13

20

27

4

JUNE

II

18

25

JULY

WEEK BEGINNING

Figure

5.

Number of coot nests initiated

per week on 3 study areas,

Colorado,

1977.

�18

C/)
~

CI)

14

'.

,/

\

'-;'1'

,

~

,
,

6

f

4

1,1

/.

2

:'

I
I-'

-....J

,

I-'

I

'

\

\

\

-.

'

,

:" I

_l'~

II

\

I

~'~

'W"

I

f

.

,

\

' e.. ~·o,

I

. ..

.

HOG LAKE

'"1P

I

I : .
I '., \
I
.'

8

LAKE JOHN

..

~

'.

a::
Z

e····· ..0

o

010

BEEBE DRAW

e---o ICE POND
'.

u,

:::::&gt;

•

,

W
Z 12

W
CD

•e-----e

..~ ...
. .
9,
..
.

16

/

_L~

17

24

I

8

APRIL

15

22

MAY

29

5

12

19

26

:3

JUNE

10

17

JULY

WEEK BEGINNING
Figure

6.

Number

of coot nests

initiated

per week on 4 study areas,

Colorado,

1978.

�24
22

•

0 BEEBE

e- -.

18

..,. -

(/)

~

(/)

16

LAKE

DRAW
JOHN

~

ICE POND

G· . ....•

HOG LAKE

ltJ

Z 14
LL.

o 12
a::
~

/

10

r.

l

\

"

,_ ... "

~

::&gt; B

~--~,

6

I

I
II

4
2

e,

23

\,,

I

I

"

WJ..

I

\

~

/ • ".'..-.'

-t: ••••••
--\.

~,
30

MAY

6

. 13

N
I

\,

,f..~

"G

I
16

'0....

..._,

\

'./

....

I
f-'

,

,,

2

'.

I

I

20

27

JUNE

·····~·.'-I
'.J
~
4

II

18

25

JULY

WEEK BEGINNING
Figure

70

Nurnb e r of coot nests

hatching per week on 4 study areas,

Colorado,

�18

g
16

.~.
..
.. ..
... ..
. ···
.
:'.
,

,

14

l(/)

W
Z

12

."

u, 10
0

0::

,

8

::&gt;

:

.

.
.
.

6

Z

,

4

f

//
8

=," :

f'

22

29

5

MAY

JOHN

A

\

1':

\,t

I"~

\~" "

\

I
I-'

-.....J

W
I

,

::_(:\~~:\ ",
'"

~:

..e

rot

f
I,~/

,=

15

-

f

.

2

\

JI, "X
'

,

LAKE

\

1

,

~ ..· -0

DRAW

,~

:/

'0

,

W
CO
~

,
,
,
,
,
,
,

BEEBE

$-----e ICE POND
G· ..... e HOG LAKE

,

CJ)

•

12

~

",

,

"
,
"',
"
A'-'","
"
/
-, '~

19

JUNE

26

:3

10

17

24

31

JULY

7

14

AUG

WEEK 8EG~NN!NG

Figure

80

Number

of coot nests

hatching

per week on 4 study areas,

Colorado,

19780

�-174-

The nesting season extended from mid April to mid August.
Beebe Draw and
Hog Lake had the earliest dates of nest initiation, Ice Pond was intermediate, and Lake John the latest.
Maximum nesting season length, from
first nest to last hatching was 12 weeks at Beebe Draw and Lake John,
15 weeks at Ice Pond, and 17 weeks at Hog Lake. The shortest nesting
season was 5 weeks at Beebe Draw. Rockwell (1912) stated the nesting
season extended from late April through July at Barr Lake, the earliest
nest being discovered on 27 April, and the latest nestswi~heggs
observed
on 21 July.
In Utah, Ryder (1961) found the first nests in early April,
with the peak occurring the third week of April, or 2-3 weeks earlier
than the closest Colorado study area, Hog Lake.
The typical sequence in the pattern of nest initiation was a low number
of early nests, followed within 1-3 weeks by an increase to peak numbers.
Thereafter, number of new nests declined, followed by 1 or more lower
peaks.
Similar patterns of nest initiation with second lower peaks were
reported by Ryder (1961) and Fredrickson (1970). Kiel (1955) also observed
second hatching peaks.
Fredrickson (1970) suggested the second peak may
represent renesting or late nests of young birds.
Crawford (1975) reported
that nest initiation date was related to age, with younger birds nesting
later than older birds.
Renesting was observed by Gullion (1954) after
desertion or predation, and by Ryder (1961) and Vaa (1972) after intentional nest destruction.
Renesting was assumed to have occurred on all study
areas.
An additional factor accounting for late nests may have been second
nests after a first successful nest.
Second nests have been reported from
California (Gullion 1954) and Utah (Ryder 1961). Ryder (1961) reported
that 13% of the coot pairs on his study area were double brooded.
Second
nests were not documented in this study.
It was suspected that 1 nest at
Ice Pond and 5 at Hog Lake were second nests.

Nesting

Statistics

A total of 354 nests was located (Table 6). There were differences in
the number of nests per area for 1977 and 1978, respectively:
Beebe Draw,
77, 6; Lake John, 67, 42; Ice Pond, 30, 29; and Hog Lake, 37, 66. The
dramatic decrease at Beebe Draw was related to low water levels.
Reduction
in the number of nests and even abandonment of nesting areas due to decreasing or low water levels have been observed (Weller et ale 1958, Krapu et
ale 1970, Vaa 1972, Weller and Fredrickson 1973). The increase in nests
at Hog Lake related to the late initiation (7 June) of nest searches in
1977. The late date of search increased the likelihood earlier nests were
not identified due to decay or vegetative growth.
Gullion (1954) noted
the rapidity with which coot nests decayed and sank after hatching; he
noted 1 nest disintegrated within 48 hours of the end of incubation.
Overall, 284 nests hatched, for a nesting success of 80.2%. Changes in
nesting success between 1977 and 1978, respectively, were observed:
Beebe
Draw, 89.6%, 33.3%; Lake John, 61.2%, 76.2%; Ice Pond, 90.0%, 79.3%; Hog
Lake, 97.3%, 81.8%. Nesting success calculations for Beebe Draw in 1978

I

�Table 6.

Coot production

data,

Production
statistic
Number
nests

Number
eggs1

1977-78.

Beebe
Draw

Lake
John

Ice
Pond

Hog
Lake

Overall

83

109

59

103

354

71

73

50

90 .

284

85.5

67.0

84.7

87.4

80.2

of

Successful
nests
Nesting
success

Colorado,

(0/0)

I
I-'

of

-....J

562

603

373

549

2,087

Average
2
clutch size

8.6+2. 3

8.3+1.2

7.5+2.2

8.8+2.3

804+2.0

Range of
clutch sizes

3-14

4-16

2-14

5-13

2-16

Number of
3
unhatched eggs

32

89

26

53

200

Hatching
success

94.3

85.2

93.0

9003

9004

(%)

INumber of eggs based on successful
Pond, 50; Hog Lake, 62.

nests

of known size:

Beebe Dr aw, 65; Lake John,

ZX + 1 So Do
3Number

of unhatched

eggs f r crn successful

nests

of known size.

75; Ice

U1
I

�-176-

were based on only 6 nests and were not considered representative.
Nesting
success at Hog Lake in 1977 appeared abnormally high.
The late search dates
at Hog Lake resulted in a greater proportion of nests being found in the
later stages of incubation or after hatching.
Nesting success was biased
upward by inability to identify and ascertain the fate of earlier nests.
Statistical tests concerning selected production data exclude Beebe Draw
in 1978 and Hog Lake in 1977 as they were not representative.
There was
no difference
&gt; 0.05) in nesting success between areas.
For all areas
combined, nests initiated before and during the peak weeks of nest
initiation showed similar nesting success (78.3%) compared to nests initiated
after the peak (82.9%).
Nesting success of coots is typically high, being
commonly reported in the 85-97% range (Miller and Collins 1954, Hunt and
Naylor 1955, Kiel 1955, Ryder
1961, Smith 1961, Fredrickson
1967, Krapu
et al. 1970, Vaa 1972).
The high reported success may be due to search
methods and/or search intervals which introduce bias (i.e.) Hunt and Naylor
1955).

(K

Examination of the causes of nest failure indicated 64.2% (76.3% in 1977,
50.0% in 1978) of the unsuccessful nests were destroyed by avian and
mammalian predators (Table 7). This was especially apparent at Lake John
in 1977 where 88.5% of the unsuccessful nests were destroyed by predators.
In 1978, only 30.0% or 3 unsuccessful nests at Lake John were attributed
to predation.
The decrease in predation may have accounted for the increased
nesting success and lower number of nests at Lake John in 1978. High nesting
success of coots is often attributed to defense of the nest by both adults
and the nest location over water.
However, predation was the major cause of
nest failure in this study.
Other causes of failure included desertion (12.9%),
and unknown or other causes (22.9%).
Two nests in the latter category at
Beebe Draw were destroyed by hail.
Table

7.

Fate of coot nests on 4 study areas,

Nest fate

Beebe
Draw

Lake
John

Successful
Unsuccessful
Lost to predators
Avian
Mammalian
Unknown
Deserted
Unknown or other failure
Total nests

71
12
6
0
2
4
2
4
83

73
36
26
4
1
21
5
5
109

Colorado,

Number of Nests
Ice
Hog
Pond
Lake
50
9
4
4
0
0

1
4
59

90
l3
9
2
1
6
1
3
103

1977-78.

Total
284
70
45
10
4
31
9
16
354

All studies have indicated desertion to be a minor cause of nest failure.
High rates of predation similar to Lake John have been reported only from
California
(Anderson 1957), where 54.3% of all nests were destroyed by
predators.
Anderson (1957) related the predation to nests located on dikes.

�-177-

Weather factors can also be a major cause of nest loss. Failure due to
wind and/or wave action has been reported (Sooter 1941, Provost 1947,
Harris and Marshall 1957). Fluctuating or decreasing water levels have
been important causes of nest failure on reservoir habitats (Wolf 1955,
Johnsgard 1956). Wind and/or wave destruction of nests were not observed
on any study area; 1 nest remained intact after being moved 1.5 m by waves
at Lake John. Nest failure due to wind and/or wave action and drawdown
was observed at Walden Reservoir in North Park.
Overall hatching success was 90.4%. Hatching success, based on successful
nests only, was consistently high on all areas, ranging from 81.9% at Lake
John in 1977 to 96.8% at Ice Pond in 1978. There was no difference
&gt; 0.05)
in hatching success between areas.
Other studies have usually reported 90%+
hatching success.

(R

Sixty-eight percent (68.5%) of egg loss was attributed to predation and
desertion (Table 8). Desertion may have been human or predator induced,
or the result of infertile or addled eggs.
Crawford (1975) found that
frequent nest checks did not cause excessive nest or egg abandonment.
Gullion
(1954) and Ryder (1961) noted eggs in the water as a factor in egg loss.
Gullion (1954) suggested that after a certain number of chicks had hatched,
the desire to incubate was overcome by the stimulus to brood, resulting in
eggs being ejected from the nest as it was converted for brooding.
In this
study eggs in the water accounted for 9.9% of egg loss. The average number
of coot eggs lost per successful nest of known size was 0.8. When unsuccessful
nests were included, the average number of eggs lost increased to 1.6.
Particularly high losses were noted at Lake John where 31.1% of all eggs
laid were lost, compared to less than 16.6% for all other areas.
Table 8.

Fate of coot eggs on 4 study areas, Colorado,

Egg fate
Hatched
Lost to predators
Avian
Mammalian
Unknown
Deserted
Dead embryo
Buried in nest
In water
Broken
Unknown or other loss
Total eggs lost
Total eggs laid

1

Beebe
Draw

Number of Eggs
Lake
Ice
John
Pond

530
35
0
16
19
13
2

514
116
20
3
93
60

1

5
22
4
24
232
746

11
1
19
82
612

1

347
20
20
0
0
15
0
0
11
2
15
63
410

1977-78.

Hog
Lake

Total

496
61
9
13
39
26
0
6
6
0
29
128
624

1,887
232
49
32
151
114
3
12
50
7
87
505
2,392

1

Eggs from successful and unsuccessful nests for all years except Hog Lake
nests of known size only.

1977 ; successful

�-178-

On each visit to a nest, the height of the eggs above the water was measured
to examine the effects of water fluctuations on nest and egg survival.
Field
observations indicated coots increased the height of the nest and eggs above
water as the laying-incubation
period progressed.
A t-test indicated an
increase
&lt; 0.001) in the average height of the eggs above water from the
time of nest initiation to late in incubation. It was assuned that nests were
free to move up or down with the supporting vegetation cs water levels
changed.
However, during periods of markedly decreasing (&gt; 20 em) water
levels, nests would eventually rest on the supporting vegetation.
Therefore,
only data from Ice Pond, (where water fluctuations were minimal) and Lake
John (floating nests) were used in statistical tests.
Gullion (1954)
indicated egg nests were frequently converted to brood nests.
Egg nests
averaged 10-15 em high at the rim; brood nests were larger, about 20 em
high due to the addition of fresh materials.
Data from Colorado suggest
coots were in the process of converting egg nests to brood nests prior to
and during hatching.
Ryan (1978) found brood platforms we re built exclusively
by males with an age related trend in the time spent building platforms.
One-year-old males only added new materials to the egg nest; young were
brooded there.
Older males built separate brood platforms, and did not use
the egg nest for brooding.
In Colorado, 34.2% of the nests showed no increase
in height, possibly the result of brood nests being built elsewhere.

(R

Average clutch size for all areas over 2 years was 8.4 eggs. Average clutch
size for each area in 1977 and 1978, respectively, was:
Beebe Draw, 8.7,
8.0; Lake John, 8.0, 8.7; Ice Pond, 7.8, 7.0; and Hog Lake, 6.5, 9.2.
There was a difference
&lt; 0.001) in clutch size between years only at
Hog Lake.
ANOVA followed by Student-Newman-Keuls
multiple range tests
indicated a difference
&lt; 0.05) in clutch size between Beebe Draw
and Hog Lake in 1977; Hog Lake and Ice Pond, Lake John and Ice Pond in 1978;
and for both years combined, between Ice Pond and all other areas.
It would
appear that clutch size was smaller in high mountain areas.

(R

(R

The model clutch size was 9 eggs (Table 9) and 90% of the nests contained
5-11 eggs each.
Fredrickson (1970) obtained similar results in Iowa where
model clutch size was 10 eggs, and 90.3% of the nests held 6-11 eggs.
Clutch size data from all areas, except Hog Lake 1977, were grouped starting
with the first week of nesting for each area (Table 10). Weekly average
clutch size decreased from a high of 9.8 eggs early in the nesting season to
a low of 5.0 late in the season.
Student-Newman-Keuls
mu l.t Lp Le range tests
indicated means 9.8-8.7 were different
&lt; 0.05) from all means 7.3 or less.
For statistical tests, data from weeks 9-13 were grouped.
Fredrickson (1967)
found a similar decrease in average clutch size from 11.1 during the first
week of nesting to 5.3 for the seventh week.
Gullion (1954), Ryder (1958),
Vaa (1972), and Crawford (1975) also noted a decrease in clutch size as the
season progressed.
The seasonal variation in clutch size may explain the
low average clutch size of 6.5 for Hog Lake in 1977. Nest data from Hog
Lake were derived from late season nests.
Crawford (1975) related variation
in ~lutch size to the age of the nesting birds.
Young coots had smaller
clutches than older coots.
He suggested that differences in average clutch
size between studies were related to the age structure of the breeding
population.

(R

�-179-

Table 9.

of clutch size for the coot in Colorado,

Frequency

of occurrence

Beebe
Draw

Lake
John

Ice
Pond

Hog
Lake

Overall

Percent

2

0

0

1

0

1

0.4

3

1

0

2

0

3

1.2

4

3

1

1

0

5

2.0

5

1

3

7

4

15

6.0

6

4

10

3

4

21

8.4

7

7

14

8

11

40

16.0

8

11

14

16

4

45

18.0

9

14

17

5

16

52

20.8

10

13

4

2

10

29

11.6

11

9

6

3

5

23

9.2

12

1

1

1

7

10

4.0

l3

0

1

0

1

2

0.8

14

1

1

1

0

3

1.2

15

0

0

0

0

0

0.0

16

0

1

0

0

1

0.4

1977-78.
Clutch
Size

�-180-

Table 10. Number of nests and clutch
nesting
season,'. Colorado"
1977-78.;

size

Week of Nesting

Clutch
size

1

16

1

2

3

4

5

6

7

in relation

to week of

Season

8

9

10

11

12

13

15
14

1

1

1

13

2

12

2

2

3

2

1

11

2

6

6

5

3

1

10

2

6

6

8

5

2

9

4

7

12

13

12

2

8

3

4

7

12

9

5

1

2

7

1

1

5

3

6

8

8

3

6

1

2

2

2

3

6

3

1

2

1

3

2

1

1

2

1

5
4

1

1
1
1
1
1

1

1

1
3

3
1

2
N

15

29

42

48

42

25

21

13

2

3

1

1

X

9.7

9.8

9.0

8.9

8.7

7.3

6.3

5.9

7.0

7.0

5.0

5.0

�-181-

Nest Site Characteristics
Cattail and bulrush were commonly used for nest sites (Table 11), with
bulrush most frequently used. At Beebe Draw in 1977, 61.0% of the
nests were in cattail, which represented 47.9% of the emergent cover.
Bulrush (14.3% of emergent cover) also was utilized to an extent greater
than its occurrence, having 31.2% of the nests.
In 1978, all nests at
Beebe Draw were in cattail.
With drawdown, most emergents were exposed.
With rising water levels, cattails were the first emergents to be flooded.
At Ice Pond, bulrush (37.8% of emergent cover) provided cover for 62.7%
of all nests. One nest at Ice Pond was in Carex. Bulrush was the only
nesting cover used at Lake John. At Hog Lake, most nest (95.1%) were
found in bulrush, however it should be noted that nearly every stand of
cattail contained a coot nest. All nests were located ove~ water.
Nest cover type in relation to date of nest initiation was examined for
Beebe Draw and Ice Pond (Table 12). In 1977, 88.9% of the nests at
Beebe Draw initiated before or during the peak week were in cattail.
No
such preference was noted at Ice Pond, with 40.7% of the nests in cattail
before the peak. After the peak of nest initiation, cover preference
became less distinct at Beebe Draw, with 54.2% of the nests in cattail.
The opposite occurred at Ice Pond, with 65.6% of the nests being found in
bulrush after the peak, and only 28.1% in cattail.
The differences in
the timing of vegetation use between the 2 areas indicates that species
of vegetation (cattail vs. bulrush) was not the most important factor
regarding nest site location.
Other factors such as the actual presence
of vegetation to serve as a nest foundation, water depth, and the presence
of other territorial coots were undoubtedly important.
Other studies have shown coots to use many vegetation types beside the
common robust emergents as a nesting substrate:
tree or shrub species such
as aspen (Populus spp.) or willows (Smith 1961), non-persistent emergents
such as Equisetum spp., Sparganium spp., Iris spp., Acorus spp., Sagittaria
spp., or Scolochloa spp. (Friley et al. 1938, Kiel 1955, Krapu et al. 1970),
and even flooded wheat fields (Kennedy 1974). All habitats used shared one
characteristic; they were standing or growing in water.
Nests on dry
land or in trees have been infrequently recorded.
Gullion (1954)
suggested that such nests were built during high water conditions, and
were later exposed when water levels decreased.
Provost (1947) believed
the variety of species used indicated that no single cover type was preferred, but that density of cover was important.
Weller and Spatcher
(1965) stated the species was irrelevant; instead vegetation to serve
as a substrate and standing in water were required.
Kiel (1955) noted
cover used varied with water fluctuations.
Examination of nest site vegetation condition, i.e., predominately dead
or alive, indicated the importance of dead or carry-over vegetation
(Table 13). Carry-over vegetation was used heavily by early nesters for
the first 3-6 weeks of the nesting season.
Thereafter a shift to live
cover occurred.
Crawford (1975) noted the shift may relate to territorial
behavior.
Early breeding coots establish a territory, nest in the avail-

�Table

11.

Number

and percentage

of coot nests

by cover

type on 4 study areas,

Colorado,

1977-78.

Lake
John

Ice
Pond

Hog
Lake

Overall

Cover

type

Beebe
Draw

Scirpus

spp.

24( 28.9) 1

109(100)

37(62.7)

98(95.1)

268(75.7)

Typha sppo

53(63.9)

0(0)

20(33.9)

5(4.9)

78(22.0)

Scirpus and
Typha mixture

6(7.2)

0(0)

1(1.7)

0(0)

7( 2.0)

Other

0(0)

0(0)

1(1.7}

0(0)

1(0.3)

Iparentheses

indicate

percentage

of nests.

I
•....
CXl
N
I

�-183-

able (dead) vegetation, and exclude later arrivals from nesting.
Later
in the season, new vegetation emerges, and is used by late nesters.
Weller
and Spatcher (1965) stated nest initiation preceded maturation of green
vegetation; nests were placed in stems of plants of the previous year.
Late nests were in green vegetation, especially in areas where there was
no vegetation early in the season.
Fredrickson (1970) also observed delayed
nesting until new vegetation developed.
Gullion (1954) and Harris and
Marshall (1957) remarked on the importance of dead materials for nesting.

Table 12.
initiation

Number of nests and cover type in relation to date of nest
at Beebe Draw in 1977 and Ice Pond in 1977 and 1978.

Cover type

Before Peak
Beebe Draw
Ice Pond

After Peak
Beebe Draw
Ice Pond

Scirpus spp.

1

16

18

21

Typha spp.

16

11

26

9

Sciq~us and Ty:eha mixture

1

0

4

1

Other

0

0

0

1

Nesting success was consistently higher for nests in cattails than in
bulrush (Table 14). When only Beebe Draw and Ice Pond were examined
(both areas with more equal proportions of cattail and bulrush compared
to Lake John and Hog Lake), nesting success was higher in cattail than
bulrush (91.8% vs. 77.0%).
Reasons for the differences are unclear.
There
are no similar comparisions in the literature.
A number of measurements were recorded at the time nests were first located
to illustrate conditions at the time of nest initiation.
Data from nests
found late in incubation or after hatching were not included in the analyses.
The numbers 0-4 (Table 15) indicate concealment afforded by vegetation at
the nest site on none, 1, 2, 3, or 4 sides. The classifications lA, 2A,
3A, 4A, indicate concealment on 1 side and from above, 2 sides and above,
etc. The majority of nests (77.9%) on all areas except Lake John were
located in what was considered good cover (classifications 3, 4, 2A-4A).
Conversely, 65.1% of all nests at Lake John were in poor cover. As
incubation progressed, 2 factors improved concealment:
1) new emergent
growth, and 2) the tendency of coots to pullover
green shoots of emergent
vegetation to form a canopy over the nest. Forty percent (39.7%) of all
nests had at least a few shoots pulled over the nest. Miller and Collins
(1954), using a comparable cover concealment rating, found 55.0% of coots
nests in good cover. Provost (1947) judged 55% of coot nests to be in
poor cover. Weller and Spatcher (1965) stated coots were tolerant of
open conditions.
Gullion (1954) reported that green leaves from surrounding
plants were pulled down and worked into the nest, but that overhead structures
were not built.
Ryder (1958) observed coots pulling and bending vegetation
adjacent to the nest, resulting in the formation of a canopy.

�Table 13. Number of coot nests and nest site vegetation
4 study areas, ..Colorado .• 1977-78. '

condition in relation

to week of nest initiation,

Week Beginning on nEEMonday
April

July

June

May

Area

3

4

1

2

3

4

5

1

2

3

4

1

2

3

Beebe
Draw

1
1/0

13/4

10/4

5/9

1/9

0/2

0/6

0/5

0/3

0/0

0/0

0/0

0/0

0/0

Lake
John

0/0

0/0

3/0

0/0

4/1

13/7

14/11

8/21

2/9

1/4

'0/2

0/1

0/1

0/1

Ice
Pond

1/0

7/0

3/0

7/0

12/2

10/3

1/2

1/3

1/1

0/3

0/1

0/1

0/0

0/0

3/0

2/0

13/1

11/5

6/5

1/4

1/4

0/3

0/2

0/4

0/0

0/0

0/1

0/0

Hog
Lake
-I

Nests

in predominantly

dead cover /nests

in predominantly

live cover.

I
f-'
IX&gt;
..j:I

�-185-

Table 14.
1977-78.

Nesting

success

(%) by cover type, 4 study areas, Colorado,

Cover type

Beebe
Draw

Lake
John

Ice
Pond

Hog
Lake

Overall

Scirpus spp.

70.8

67.0

81.1

86.7

76.5

Typha spp.

92.4

90.0

100.0

92.3

Scirpus and
Typha mixture

83.3

100.0

85.7

100.0

100.0

Other

Table 15. Concealment
study areas, Colorado,
Sides of
concealment

afforded by vegetation
1977-78.

at coot nest sites on 4

Beebe
Draw

Lake
John

Ice
Pond

Hog
Lake

0

1

48

1

3

1

8

8

0

5

2

13

15

1

15

3

14

15

8

10

4

17

3

11

6

lA

0

0

0

0

2A

1

2

0

4

3A

6

6

3

8

4A

21

12

35

22

�-186-

The average height of vegetation at the nest site was: Beebe Draw, 63 cm;
Lake John, 36 cm; Ice Pond, 75 cm; and Hog Lake, 80 cm. The average depth
of water and range of depths at the nest were:
Beebe Draw, 69 cm, 24-142
cm; Lake John, 50 cm, 0.5-119 cm; Ice Pond, 37 cm, 2-101 em; and Hog Lake,
41 cm, 21-64 cm (Table 16). The average distance of the nest from open
water and range of distances were:
Beebe Draw, 7.7 m, 0.3-35.1 m; Lake
John, 4.1 m, 0-22.9 m; Ice Pond, 2.8 m, 0.3-18.3 m, and Hog Lake, 3.7 m,
0-22.9 m (Table 17). Miller and Collins (1954) indicated 59.2% of coot
nests were in vegetation 90 cm or taller, while Krapu et a1. (1970) gave
an average height of 84 cm. Gullion (1954) reported most nests were
within 60-90 cm of the water, with 122 cm the maximum.
Provost (1947)
and Sooter (1941) gave average distances of 9.9 and 16.2 m, respectively,
from open water.

Table 16.

Number of coot nests by water depth on 4 study areas, Colorado,

1977-78.
Beebe
Draw

Lake
John

Ice
Pond

Hog
Lake

0-49 cm

4

65

51

67

50-99 cm

74

38

7

13

100~149 cm

4

6

1

c

Depth

Table 17. Number of coot nests by distance
Colorado, 1977-78.

to open water on 4 study areas,

Distance

Beebe
Draw

Lake
John

Ice
Pond

Hog
Lake

0-1. 9 m

15

58

37

46

2-3.9 m

16

13

9

27

&gt; 4 m

52

38

13

30

The wide range in nest site measurements illustrates not only the widespread
use of the emergent zone by coots, but also basic differences in the nature
of the study areas.
The poor nest concealment at Lake John related to a
lack of carry-over vegetation on portions of the study area. A shortened
growing season due to the high altitude of Lake John also influenced nest
concealment and the height of vegetation at the nest. Differences in water

�-187-

depth at the nest site related to different bottom profiles and water
regimes.
Differences in the average distance to open water were a result
of different patterns of vegetation growth.
Beebe Draw, with the highest
average distance to bpen water, had a wide emergent zone with little interspersion of open water.
Ice Pond, with the lowest average distance, had a
narrow emergent zone with many emergent islands and oper. water channels.
Nest site measurements at Lake John indicated many nests were located in
shallow water and poor cover compared to other areas.
These factors may
have allowed better nest visibility and accessibility to predators and
resulted in the high nest failure at Lake John. Only data from 1977 were
included in the analyses, since predation was the major cause of nest failure
that year. T-tests indicated there was no difference (f &gt; 0.05) in the
average water depth at successful vs. unsuccessful nests.
Chi-square
analysis indicated no relationship
&gt; 0.05) between wa~er depth class
and nest success or failure.
Chi-square analysis indicated a relationship
&lt;
0.05) between cover and nest success or failure.
However, results
were the opposite of those expected, with nest success being related to
poor cover (Table 18). Results were explained by examination of the Lake
John nesting habitat.
Twenty-five of the successful nests in poor cover
were found in 2 emergent islands located 30 m or more from the shore and
surrounded by broad expanses of open water.
Vegetation in these islands
was exposed to wind, wave, and ice action, resulting in little carry-over
and concealment for the beginning of the breeding season.
Nesting success
was higher on these islands (71.4%) than on other portions of the marsh
(50.0%), probably due to poor access to mammalian predators.

(I

(I

Table 18. Relationship
John, Colorado, 1977.

of coot nest success and nest site cover, Lake

Number of Nests
Category

Successful

Good cover

10

13

Poor cover

31

13

Total

41

26

i

4.63,

df

Unsuccessful

1, P &lt; 0.05

Breeding Densities
Breeding densities were estimated from the number of successful nests
per hectare of emergent habitat (Table 19). In addition, the breeding
density within a given emergent component was examined.
Density based
on total emergent areas included species which were little used or not

�Table 19. Breeding densities of coots as number
habitat on 4 study areas,
Colorado" 1977 -78. )
Beebe
Draw

Cover type
Total
emergents
Carex

9.6/0.7/7.0

spp.

Scirpus

spp.

Typha spp.

of succes sful nests per hectare

Lake
John
1

14.8/11.

0/0/0
16.5/0/11.2

6/13. 2

0/0/0
14.8/11.

13.6/1.3/9.9

6/13.2

0/0/0

of emergent

Ice
Pond

Hog
Lake

30.0/25.6/27.8

10.7/17.4/14.0

16.7/0/8.3

0/0/0

47.0/41.2/44.1
22.5/22.5/22.5

16.0/28.7/22.0
I
I-'

75.0/50.0/62.5

co
co
I

Combined
Scirpus and
Typha spp.

15.4/1.

11977 breeding

0/10. 2

density/1978

14.8/11.6/13.2
breeding

35.1/31.

density/ 1977 -78 average

1/33. 1
breeding

17. 11,29. 2/22. 8
density.

�-189-

used for nesting, i.e., sedges, spikerush, salt grass, common reed, and
common three-square, (Scirpus americanus).
A better basis for breeding
density is that based on the common robust emergents, Typha and Scirpus
spp. Breeding densities ranged from a low of 1.0 successful nests per
hectare of cattail and bulrush at Beebe Draw in 1978 to a high of 35.1
at Ice Pond in 1977. The breeding density for Beebe Draw in 1978 was
not representative of normal conditions, but documents the importance
of water and habitat conditions during the breeding season.
The 1977
Hog Lake breeding density was also not considered representative due
to the late date of nest search.
Densities for Ice Pond and Hog Lake
were 2-3 times greater than for the other areas, although ANOVA (ex~
cluding Beebe Draw in 1978 and Hog Lake in 1977) indicated a difference
(K &lt; 0.05) only between Lake John and Ice Pond.
In 1977 and 1978, segments of the emergent zone at Beebe Draw were burned.
Burning failed to kill any emergents and increase open water area.
In
1977, the burned area of 3.1 ha consisted mostly of common three-square.
No nests were found in the three-square, although brood platforms and
foraging broods were observed.
Six successful nests (representing all
nests in the burned area) were found in 0.7 ha of burned cat t a t l.. The
resultant breeding density of 8.2 for the burned cattail was substantially
less than that of the remaining unburned cattail with 15.1 successful
nests/ha.
In 1978, no nests were found in the burned areas,
Measures of breeding or nesting density in the literature are not comparable
to this study. The areas measured often are not clear, i.e., emergent
vegetation only or emergent and water areas. Also, apparently all nests,
successful or not, were counted.
A given measure of breeding density useful for one area may not be appropriate for another.
For example, in the
pothole country of Manitoba density based on total emergent and open water
area may be useful, since the total area may be apportioned by coots. However, in Colorado, the study areas had broad expanses of open water, making
it inappropriate for use as a breeding density measure.
The closest comparison in the literature is that of Ryder (1961), who calculated the
highest coot nest densities as over 400 nests/100 acres of available cover
(&gt;9.9 nests/ha of cover).
As calculated, breeding density was in part determined by nesting success.
However, the actual number of nesting pairs appears related to interspersion
of open water and emergent vegetation.
Weller apd Fredrickson (1973) noted
that highest coot densities occurred at a 50:50 cover:water ratio.
Ryder
(1961) showed nest densities increased as the proportion of open water to
available nesting cover increased.
Smith (1961) observed that in the year
of highest coot population, the majority of Type 4 and 5 wetlands were in
an "open" (0-33% covered with vegetation) or "half" (34-66% covered with
vegetation) cover class. On the Colorado study areas, the ~road expanses
of open water made a comparison of cover to water inappropriate.
Measurements of interspersion within the emergent zone were not undertaken, however,
ocular estimates indicated Ice Pond and Hog Lake had the greatest interspersion.

�-190-

Nonbreeders

Counts of coots conducted on each visit to a study area were lower in
1978 than in 1977 (Table 20). This was due to a decrease or absence
of nonbreeders in 1978. Nonbreeding coots were identified by their
gregarious behavior and lack of territorial activities.
They normally
moved about as a group and avoided territorial or breeding coots.
Usuually seen on open water, they were more conspicuous than breeders.
On the study areas, coots occurring primarily in open water were the
only ones which could be seen and counted.
Based on a knowledge of
nest locations, it was estimated that no more than 50% of the breeders
were observed on a given occasion.
Marsh vegetation hid most breeders
or they remained concealed.
Assuming that a relatively constant
proportion of the breeding coots was observed on each count, and that
there were no substantial changes in the number of nesting pairs, then
count differences were due to a decrease in the number of nonbreeders.
At Hog Lake and Ice Pond, it is believed there were only small changes in
the number of nesters between 1977 and 1978. At Lake John Annex, there
were fewer nests in 1978 and 1977, but not enough for count differences.
At Beebe Draw low water levels greatly reduced the number of nesters in
1978. The effects of low water levels on nonbreeders is unknown.
However,
no more than 12 nonbreeders in 1 group were observed at BEebe Draw in
1978. Thirty to 50 nonbreeders were commonly seen at Beebe Draw in 1977.
Observations on peripheral marshes also indicated a decrease in nonbreeders
from 1977. Ryder (1963) and Crawford (1975) observed nonLreeders, but
did not mention any change in numbers between years.
Crawford indicated
the majority were first year birds.
Smith (1961), in Alberta, observed
no unpaired or nonbreeding coots.
The lack of sightings may relate to the
pothole habitat and the territorial behavior of coots.

Brood Counts
Counts of coot broods were initiated on all areas with the first appearance of young and continued through August.
Gullion (1956) suggested
caution in the evaluation of counts of coot broods.
He cited the tendency
of parent coots to divide the brood and to feed chicks in different parts
of their territory, often hidden from view.
In additio~, young coots may
be left unattended in dense cover while part of the brood forages with the
The range in time from hatching of first nests to the first
adults.
observation of chicks varied from 1 to 30 days, a result
of chicks remaining in the emergent vegetation and staying out of view (Table 21). Gullion
(1956) stated that to obtain accurate results, the observer must be familiar
with coot territorial behavior, know the territory size, and make several
counts daily for a week.
These requirements could not be met in this study.
Gullion (1956) implied that counts of young coots over 25 days of age
may be more productive than counts of younger coots.
By 30 days of age,
young coots move into more open water and are more visible (Gullion 1954).

�Table 20.

Coot numbers

observed

on 4 study areas,

Colorado,

Week Starting

mid May through June,
Eli

on n

1977 and 1978.

Monday

May

June

Area

3

4

5

1

2

3

4

Beebe Draw

50/111

95/25

69/--

53/14

76/4

66/--

73/9

Lake John

172/--

199/124

101/74

104/--

193/20

186/12

219/--

Ice Pond

75/--

85/59

80/43

80/--

75/54

92/37

118/--

Hog Lake

280/--

200/105

--/75

155/--

234/58

244/102

436/--

--

11977/1978.

I
I-'
\0
I-'
I

�-192Table 21. Dates of hatching
on 4 study areas, Colorado.

vs. dates when young coots were first observed

1977

1978

First Nests
Hatched

First Young
Observed

First Nests
Hatched

First Young
Observed

Beebe Draw

Week of
23 May

10 June

Week of
26 June

23 July

Lake John

Week of
13 June

30 June

Week of
12 June

25 June

Ice Pond

Week of
30 May

6 June

Week of
15 May

1 June

Week of
15 May

13 June

Area

Hog Lake

Brood members become gregarious during this period, and while more independent
of adults, they remain on territory.
Young coots at this stage were easily
identified by a whitish breast, throat, and head contrasting with the darker
back.
Smith (1961) stated that estimates of coot production could best be
made in late summer when most young coots had reached the Class III (fully
feathered but flightless) stage because of their habit of gathering in open
water.
To ascertain if counts of young coots 30 days of age or older were a useful
index of trends in coot productivity, counts were compared to nesting success
(Table 22). Analysis required a knowledge of local conditions, i.e., the
location and water levels of nearby marshes, and their use by coots.
Adult
and juvenile coots will leave a marsh as water levels recede and it becomes
dry. This situation was particularly applicable to Beebe Draw and Hog Lake,
where nearby marshes with breeding coots occurred.
A late summer buildup
of adult and juvenile coots suggested an influx from other areas.
Counts
used in the analysis represent the highest number of young 30 days of age
or older seen on anyone
occasion with most or all of the young being produced on the given area.
Comparison of high counts and nesting success
as determined from nest searches indicated a positive relationship.
At
Lake John the high count in 1977 was 17 young and nesting success was 61.2%.
In 1978, 44 young were seen and nesting success increased to 76.2%.
Simple
linear correlation indicated a strong positive association (r = 0.91, K &lt;
0.005) when data from all areas were combined.
It would appear that trend
in number of young seen may be a useful index to productivity.
Smith (1961)
compared the total number of young observed to counts of adult coot pairs
made earlier in the season.
His data also indicated a strong positive
correlation (r = 0.99, ~ &lt; 0.001).
Summer Movements
N~mbers of adult coots increased gradually on the study areas starting in
mld to late July.
Increases in numbers of juvenile coots 30 days of age
or older were observed beginning mid to late August.
There were 2 exceptions
to the general pattern.
At Beebe Draw in 1977, numbers of coots decreased

�-193-

through July and continued low through most of August.
However, 3 marshes
adjacent to Beebe Draw had substantial increases beginning in August,
continuing into September, with each containing up to 150 coots.
The
buildup on peripheral marshes represented a late summer influx from other
areas and/or movement from Beebe Draw in response to an apparent poor
supply of aquatic vegetation.
In 1978, the reverse occurred.
With the
drying of a peripheral marsh and an improved aquatic food supply on the
study area, numbers of coots increased steadily at Beebe Draw beginning
in mid August.
Numbers of coots on adjacent marshes remained low compared
to 1977. An influx of adult and juvenile coots on Hog LaKe in 1977 began
about 2 weeks earlier than the general pattern found on other areas. Movements on to Hog Lake were in part related to pumping schedules and water
levels on nearby marshes.
Water levels were allowed to recede on some
marshes in Brown's Park NWR as the summer progressed.
Juvenile coots were
observed on 1 occasion crossing the Green River in the direction of Hog
Lake. Ryder (1958) observed a progressive increase in coot numbers in
the latter half of August on larger bodies of water.
Broods were seen
moving overland from drying natal marshes.
Ward (1953) saw similar movements
in Manitoba.
Postbreeding adults and nonbreeders began to concentrate by
mid July to undergo molt, and were joined by 5 week or older juveniles in
early August.
Bergman (1973) observed concentrations of up to 50,000 coots
during late August and September in Manitoba.
Table 22. Nesting success and highest number of young coots 30 days of age
or older seen on 4 study areas, Colorado, 1977 and 1978.

Location

Number Counted
1977
1978

Beebe Draw

34

Lake John
Ice Pond
Hog Lake

(%)

Nestin_g_Success
1977

1978

2

89.6

33.3

17

44

61.2

76.2

46

42

90.0

79.3

35

81.8

Fall Migration
Counts to document fall migration of coots indicated differences in timing
of peak numbers between 1977 and 1978. Peak numbers were recorded in 1977
on all areas during late August to mid September, before the waterfowl
hunting season of 1-14 October (Fig. 9). In 1978, peak numbers were observed 5 weeks later at Beebe Draw, 4 weeks later at Lake John 1 week
later at Ice Pond, and 6 weeks later at Hog Lake (Fig. 10). P~ak numbers
occurred before the 30 September-13 October 1978 waterfowl season at Ice
Pond and during or after the hunting season for the other areas.
In
November 1977 and 1978, coot numbers decreased to minimal levels.
No
coots were known to winter on the study areas.
Compared to spring migration, fall migration occurred over a more widespread
~eri~d~ with l~ss dramatic changes in numbers, and with less consistency
1n t1m1ng.
Ba1ley and Niedrach (1965) stated the m~jority of coots.in

�-194-

3300
•
._

•
-- __

BEEBE DRAW
LAKE JOHN

0-- __ ICE F-OND

e·· .···e HOG LAJ&lt;E
,

•

~

o

o
u,

o
a::
IJ.l

m
~

:::&gt;

z

•... ... ...••••..
.

.

.

...

..

'.
·0

~

". J!i.....
/.
~~

.

'.
:

•

_/

~.

•• 0

0

•••••

. -.fiT

" ••
~"

~ &amp;
//

..
~ ..~v.'.

"

~

~

"

~

''''0

26

3
1
8
15
22
29
5
~---------------~
~------------~
12

AUGUST

19

SE~TEMt:lER

10

17

24

OCTOBEI-&lt;

WEEK BEGINNING

Figure 9. Coot numbers
4 study areas,
Colorado,

observed
1977.

during

late

summer

and fall,

�-195-

•

•

• - BEEBE DRAW

----0

LAKE JOHN

--.

ICE

.

,

FOND

e····· ··e HOG LAKE

/

~ ..,

i\ \

I

I,

I

" ..i

I

'"

,r
,
I

\
\

!

\.-'~

I
,,

j

~

o
o

~

o

,I

LL

o

1/

m

~

::&gt;

z

I

I'
I

".

/

..

I

,,-

I

0::
ltJ

,

...

.....••

",

..

-----i'
. ...._ .. ...•
.'

-

'

°3~1~~7~~14~~2~1---2~8---L4~~Il--_JJ8--_J25--_J2--~9L---JL6---2L3---30~~

~

JULY

AUGUST

'----

~

_J

SEPTEMBER

OCTOBER

WEEK BEGINNING
I

Figure 10. Coot numbers obse rved during
4 study areas,
Colorado,
1978.

late summer

and fall,

�-196-

Colorado migrated south by the last of October; Ryder (1958) reported coot
numbers in Utah declined greatly each fall after 15 October.
Ward (1953)
observed migration in several waves from Delta, Manitoba.
The first wave
peaked in late August or early September, and departed before mid September.
A second buildup occurred through the rest of September into early October.
Coot numbers dropped suddenly the first week of October.
Successive migratory
waves may explain the fluctuating numbers observed on some Colorado areas.
Harvest
In Colorado, waterfowl harvest records are derived from mail surveys
sent to approximately 3% of the small game license buyers.
Colorado
Small Game Hunter Harvest surveys from 1954 through 1972 did not give
specific harvest figures for coots, but instead grouped them in the
category of "others and unknown."
This group included coots, mergansers,
goldeneyes, buffleheads, and unknowns.
Data from more recent harvest
surveys (Table 23) indicate an average harvest of 4,300 coots for the
1973-76 period CH. Riffel, pers. comm., M. F. Sorensen, S. M. Carney,
and E. M. Martin, Admin. Rep. OMBM U.S. Fish &amp; Wildl. Servo unpubl.
data).

Table 23.

Year

Coot ha.rvest in Colorado,

Estimated

1973-76.

Harvest

Percent

of Duck Harvest

1973

4,579

1.6

1974

5,054

1.4

1975

2,924

1.2

1976

4,645

2.3

To further determine hunting pressure and harvest levels of coots, volunteer
wing collection barrels were placed at Lake John (1), Walden Reservoir (2),
Cowdrey (1), MacFarlane Reservoir (1), Antero Reservoir (1), Hog Lake (1),
and Wellington State Wildlife Area (2). In addition, wildlife students
collected wings at the Wellington check stations.
Wings obtained during
1-14 October 1977 indicated a harvest of 443 ducks and 8 coots, or a coot
harvest representing 1.8% of the duck harvest (Table 24). Wings collected
during 30 September-13 October 1978 indicated a coot harvest representing
2.0% of the duck harvest.
These harvest percentages are similar to those
estimated statewide from 1973-76.

�-197-

Table 24. Coot and duck harvest as obtained from wing barrels and check
stations on Colorado hunting areas, 1-14 October 1977, and 30 September13 October 1978.

1978

1977
Duck
wings

Coot
wings

Duck
wings

268

2

557

1

Antero Reservoir

68

0

92

0

Hog Lake

42

0

4

0

Wellington

65

6

106

14

Total

443

8

759

15

Area
North Park

1

1
Includes wing barrels
reservoirs.

at Lake John, Cowdrey, Walden,

Coot
wings

and MacFarlane

To investigate hunter attitudes, students surveyed hunters at Wellington
Wildlife Management Area on the opening weekends of the 1976 and 1977
waterfowl seasons.
Of 86 hunters polled in 1977, only 15 (17.4%) indicated
they normally hunted coots.
Of the remainder, 36 cited bad taste as the
season for not hunting coots.
Similar results were obtained in 1976
when only 11 of 54 hunters (20.4%) indicated they normally hunted coots.
The general concensus was that coots were not worth shooting and that
ducks were preferred.
Despite liberal daily bag limits of 15 and 25 in the Central and Pacific
flyways, respectively, coot harvest is low in Colorado.
Coot harvest in
the Central Flyway is the lowest of all flyways (Fredrickson 1977). Hunter
interest in coots is low. Out of 67 questionnaires returned by Division
of Wildlife personnel, only 7 indicated having encountered hunters who
retained coots in the bag. In Wisconsin, Bartelt (1977) reported only 18%
of hunters normally hunted coots; Fredrickson (1977) reported similar
figures from California.
Bartelt (1977) also indicated the unretrieved
coot harvest was 45 and 25% of the total coot harvest for 1975 and 1976
waterfowl seasons, respectively.
Unretrieved coots were found in this
study; however, thorough searches of hunting areas were not conducted.
The hypothesis concerning hunting mortality and locally ?roduced coots
was not tested.
In view of the timing of coot fall migration, low hunter
interest, and low coot harvest, it is unlikely hunting is an important
source of mortality for locally produced coots.

�-198-

SUMMARY
Investigations concerning distribution, status, reproductive success,
breeding densities, habitat utilization, timing of migration, and hunter
harvest of coots were conducted in Colorado between August 1976 and
November 1978. Four study areas were selected to represent breeding
sites in the major topographic areas of the state, I in the eastern
plains, 2 in high mountain valleys, and 1 west of the Continental Divide.
Coots bred throughout Colorado with major breeding areas in Brown's Park
on the western slope and North Park and the San Luis Valley.
On the
plains, coots were locally abundant where suitable habitat occurred.
Breeding and migrating coots were associated primarily with cattail and
bulrush dominated marshes (~&lt; 0.001). Other emergent vegetation used
for nesting included tamarix, spikerush, willows, and sedges.
Large
lakes and reservoirs were used by non-breeders in summer and by migrants
in spring and fall. Coots were resident year round in low numbers « 1,000)
mainly along the western boundary of the plains.
Coots first arrived in late February or early March (first week of April
for Lake John), building to peak numbers during the second to third week
of April.
A delay of 1 week in peak migration occurred in the high mountain
valleys.
Coot numbers decreased to resident levels by the second to fourth
week of May.
Onset of breeding was 1-3 weeks before to I week after peak migration periods,
with a shortened territory establishment-to-nest
initiation cycle in the
high mountain valleys.
Nest initiation dates ranged from mid April to mid
July; peak initiation dates ranged from 25 April to 12 June. Water levels
and cover condition affected timing of nest initiation.
Dates of hatch
ranged from mid May through early August; peak hatching dates ranged from
30 May to 4 July.
A total of 354 nests was located, with nesting and hatching success, respectively, of 80.2 (range of 33.3-97.3%), and 90.4% (range from 81.9-96.8%).
There were no differences (~ &gt; 0.05) in nesting or hatching success between
areas.
Average clutch size was 8.4 and ranged from 6.5 to 9.2 depending
upon area. Differences
&lt; 0.05) in average clutch size existed between
areas.
Clutch size decreased as the season progressed.
Predation was the
major cause of nest failure, accounting for 64.2% of all unsuccessful nests.
Desertion and other factors accounted for the remainder.

(K

Cattail and bulrush were the cover types most commonly used for nesting.
All nests were located over water.
Early nesters depended
heavily on
dead or carry-over vegetation.
Water levels and cover condition affected
nest site selection.
Concealment afforded by the vegetation and other nest site measurements at
time of nest initiation were variable and depended on local conditions.
Cover concealment at Lake John was less important to nest success than nest
location in relation to distance to shore and surrounding open water.

�-199-

Breeding densities (the number of successful nests per hectare of cattail
and bulrush) ranged from 1.0 to 35.1. Factors influencing number of
nesting coots were water levels and interspersion of open water and
emergent cover.
Nonbreeding coots were observed on all study areas; however, fewer were
observed in 1978 than 1977. Reasons for the change are unclear.
Counts
of young coots less than 30 days of age were judged unsatisfactory as an
index to productivity.
However, counts of young coots 30 days of age or
older showed a strong positive association when compared to nesting
success (r = 0.91, ~ &lt; 0.005).
Late summer movements of adult and juvenile coots onto the study areas
were observed.
Decreasing water levels on peripheral ma~shes and abundant
aquatics on the study areas were contributing factors.
IDcreases in adult
numbers were observed starting in mid to late July, and in juveniles
beginning mid to late August.
.
Peak concentrations of fall migrating coots were recorded in September in
1977. Compared to 1977, peak numbers in 1978 occurred 1-6 weeks later
depending upon the area. Fluctuating numbers observed on the study areas
suggested coot migration in several waves.
Coot numbers dropped to minimal
levels during November during both years.
Over 2 hunting seasons, coots represented 1.9% the total waterfowl harvest.
Hunter questionnaires indicated that fewer than 20% of waterfowl hunters
normally hunt coots.
It is unlikely hunting is an important source of
mortality for coots in Colorado.

�-200-

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habitat development and management. Habitat Develcpment and
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Bailey, A. M., and R. J. Niedrach. 1965.
Mus. Nat. Hist. Vol. I. 454pp.

Birds of Colorado.

Denver

Bartelt, G. A. 1977. Aspects of the population ecology of the American
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Bergman, R. D. 1973. Use of southern boreal lakes by post-breeding
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143pp.

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Crawford, R. D. 1975. Breeding biology of American coots in relation
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Wilson

_____ , Chairman. 1977. American coot (Fulica americana). Pages 123147. In G. C. Sanderson (ed.). Management of migratory shore and
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The American

Gullion, G. W. 1952. The displays and calls of the American coot.
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Condor

1954. The reproductive cycle of American coots in California.
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�-201-

Harrington, H. D. 1964. Manual of the plants of Colorado.
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666pp.

2nd Ed.

Harris, S. W., and W. H. Marshall.
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Ecology
37:689-701.
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1974. Unusual nesting attempts
J. Wi1d1. Manage. 38:937.

Keyser. L. S.
355pp.

1902.

Birds of the Rockies.

by waterfowl

McClory

in southern

&amp; Co., Chicago.

Kie1, W. H. 1955. Nesting studies of the coot in southwestern
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Rockwell, R. B. 1912. Notes on the wading birds of the Barr Lake region,
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�-202-

Ryder, R. A.
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1958. Coot-waterfowl relationships
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219pp.

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Sooter, C. A.
americana
120pp.

1969.

Biometry.

W. H. Freeman

and Co.

1941. Ecology and management of the American coot Fulica
americana Gmelin.
Ph.D. Thesis.
Iowa State Univ., Ames.

Vaa, S. J. 1972. Use of waterfowl production areas by ducks and coots
in eastern South Dakota.
M. S. Thesis.
S. Dakota State Univ.,
Brookings.
31pp.
Ward, P. 1953. The American
Conf. 18:322-329.

coot as a game bird.

Trans. N. Am. Wildl.

Weller, M. W., and L. H. Fredrickson.
1973. Avian ecology
glacial marsh.
The Living Bird 12:269-291.

of a managed

_____ , and C. S. Spatcher.
1965. Role of habitat in the distribution
and abundance of marsh birds.
Iowa State Univ. Agric. Exp. Stn.
Spec. Rep. 43. 31pp.
and J. B. Low.
1958. Effects of habitat deteron bird populations of a small Utah marsh.
Condor 60:220-226.

---- , B. H. Wingfield,
ioration

Wolf, K. 1955. Some effects of fluctuating and falling water levels on
waterfowl production.
J. Wildl. Manage. 19:13-23.

Prepared

by __ Wt__~_rltJ2.&lt;.
__
2_. _t1._(!)_r_~-,=,~
__ ~'IQK~
Warner P. Gorenzel
Graduate Research Assistant

Approved

by __ ~~~~~~.~~=-.~~~~
Clait E. Braun
Wildlife Researcher

_

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